Aquatic Snakes

Aquatic Snakes,
Diversity and Natural History

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Aquatic Snakes

This book is dedicated to the present and future herpetologists who

will study aquatic snakes and promote their conservation.

ISBN 9798218194901
Copyright © 2024 John C. Murphy
Publisher JCM Natural History
with Herpetological Conservation International

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 Cover Photos

Cover Photos

The front cover photo was taken on Orchid Island (Lanyu) off the southeastern part of
Taiwan on June 30, 2020. It happened to be at low tide that night, so there were many sea
kraits active between the crevices of the coral reef and the tide pool (the species appeared that
night including Laticauda semifasciata, Laticauda colubrina and Laticauda laticaudata). Since
Laticauda semifasciata rarely crawls out of the sea surface, I crouched in a shallow tidal pool
and waited for it to swim to my predetermined position. I used DSLR (Nikon D850) with a
manual wide-angle lens (LAOWA 15mm f4) and off-camera flashlight shooting this photo.
Yu Sing Lin

On the back cover a Northern Water Snake, Nerodia sipedon, lies at the edge of a man-
made canal shortly after emerging from hibernation. Will County, Illinois. JCM

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Aquatic Snakes

Contents
3 Cover Photos
15 Co-Authors to Aquatic Snakes
17 Acknowledgments
19 Preface
22 1. Introduction
38 2. Aquatic Snake Biology
50 3. Aquatic Snake Geography
72 4. Ancient but Extant Lineages
72 Trachyboa boulengeri Peracca, 1910
74 Trachyboa gularis Peters, 1860
75 Anilius scytale (Linnaeus, 1758)
75 Anilius phelpsorum Roze, 1958
76 Xenopeltis unicolor Reinwardt, 1827
77 Liasis fuscus, Peters 1873
79 Liasis olivaceus Gray, 1842
79 Malayopython reticulatus Schneider, 1801
81 Python bivittatus Kuhl, 1820
82 Python molurus (Linnaeus, 1758)
83 Python natalensis Smith, 1840
85 Python sebae Gmelin in Linnaeus, 1789
87 Eunectes beniensis Dirksen, 2002
87 Eunectes deschauenseei Dunn and Conant, 1936
88 Eunectes murinus (Linnaeus, 1758)
89 Eunectes notaeus Cope, 1862

95 5. File Snakes
98 Acrochordus arafurae McDowell, 1979
101 Acrochordus granulatus Schneider, 1799
102 Acrochordus javanicus Hornstedt, 1787

107 6. Odd Scaled Snakes

113 7. Aquatic Vipers
113 Agkistrodon conanti (Gloyd, 1969)
114 Agkistrodon piscivorus (Lacépède, 1789)
116 Craspedocephalus malabaricus (Jerdon, 1854)

120 8. The Homalopsidae

129 Brachyorrhos albus (Linnaeus, 1758)
130 Brachyorrhos gastrotaenius (Bleeker, 1860)
131 Brachyorrhos pygmaeus Murphy & Voris, 2020
131 Brachyorrhos raffrayi (Sauvage, 1879)
131 Brachyorrhos wallacei Murphy, et al. 2012

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132 Calamophis jobiensis Meyer, 1874

133 Calamophis katesandersae Murphy, 2012
133 Calamophis ruuddelangi Murphy, 2012
134 Calamophis sharonbrooksae Murphy, 2012
134 Karnsophis siantaris Murphy & Voris, 2013
135 Bitia hydroides Gray, 1842
136 Cantoria violacea Girard, 1858
138 Cerberus australis (Gray, 1842)
139 Cerberus dunsoni Murphy, Voris, and Karns, 2012
140 Cerberus microlepis Boulenger, 1896
141 Cerberus rynchops (Schneider, 1799)
143 Cerberus schneiderii (Schlegel, 1837)
145 Dieurostus dussumieri (Duméril, Bibron & Duméril, 1854)
147 Djokoiskandarus annulata (De Jong, 1926)
147 Enhydris chanardi Murphy & Voris, 2005
148 Enhydris enhydris (Schneider, 1799)
150 Enhydris jagorii (Peters, 1863)
152 Enhydris innominata (Morice, 1875a)
153 Enhydris longicauda (Bourret, 1934)
154 Enhydris subtaeniata Bourret, 1934
155 Erpeton tentaculatum Lacépède, 1800
157 Ferania sieboldii (Schlegel, 1837)
158 Fordonia leucobalia (Schlegel, 1837)
160 Gerarda prevostiana (Eydoux & Gervais, 1837)
163 Gyiophis maculosa (Blanford, 1881)
163 Gyiophis salweenensis Quah, et al. 2017
164 Gyiophis vorisi (Murphy, 2007b)
165 Heurnia ventromaculata Jong, 1926
165 Homalophis doriae Peters, 1871
166 Homalophis gyii (Murphy, Voris & Auliya, 2005)
167 Homalopsis buccata (Linnaeus, 1758)
169 Homalopsis hardwickii Gray, 1842
169 Homalopsis mereljcoxi Murphy et al. 2012
172 Homalopsis nigroventralis Deuve, 1970
173 Homalopsis semizonata Blyth, 1855
174 Hypsiscopus indonesiensis Hamidy et al. 2023
174 Hypsiscopus matannensis (Boulenger, 1897)
175 Hypsiscopus murphyi Bernstein et al. 2022
176 Hypsiscopus plumbeus (Boie, 1827)
177 Kualatahan pahangensis (Tweedie, 1946)
178 Mintonophis pakistanicus (Mertens, 1959)
179 Miralia alternans (Reuss, 1834)
180 Myanophis thanlyinensis Köhler, et al. 2021
181 Myron karnsi Murphy, 2011
182 Myron resetari Murphy, 2011
182 Myron richardsonii Gray, 1849
184 Myrrophis bennettii (Gray, 1842)
185 Myrrophis chinensis (Gray, 1842)

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186 Myrrophis dakkrongensis Nguyen et al. 2024

186 Phytolopsis punctata Gray, 1849
188 Pseudoferania polylepis (Fischer, 1886)
190 Raclitia indica Gray, 1842
191 Subsessor bocourti (Jan, 1865)
193 Sumatranus albomaculata (Duméril, et al., 1854)

195 9. Aquatic Coral Snakes

195 Micrurus nattereri Schmidt, 1952
196 Micrurus surinamensis (Cuvier, 1816)

199 10. Aquatic Cobras (Najinae)

199 Naja annulata Peters, 1876
201 Naja christyi Boulenger, 1904
201 Naja nana Collet and Trape, 2020
203 Naja melanoleuca Hallowell, 1857
203 Naja subfulva (Laurent, 1955)
204 Naja peroescobari Ceríaco, Marques, Schmitz & Bauer 2017
204 Naja (Boulengerina) guineensis Broadley, et al., 2018
204 Naja (Boulengerina) savannula Broadley, et al., 2018
204 Naja kaouthia Lesson, 1831

207 11. Semi-Aquatic Hydrophiinae

207 Micropechis ikaheka Lessons, 1830
208 Austrelaps labialis (Jan, 1859)
208 Austrelaps ramsayi (Krefft, 1864)
209 Austrelaps superbus (Günther, 1858)
209 Hemiaspsis damelii (Günther, 1876)
210 Hemiaspis signata (Jan,1859)
211 Pseudechis porphyriacus (Shaw, 1794)
212 Notechis scutatus (Peters, 1861)

215 12. Sea Kraits

216 Laticauda colubrina (Schneider, 1799)
219 Laticauda crockeri (Slevin, 1934)
220 Laticauda schistorhyncha (Günther, 1874)
221 Laticauda saintgironsi (Cogger and Heatwole, 2005)
223 Laticauda guineai (Heatwole et al 2005)
223 Laticauda frontalis (de Vis, 1905)
224 Laticauda laticaudata (Linnaeus, 1758)
225 Laticauda semifasciata (Reinwardt, 1837)

227 13. Seasnakes

228 Aipysurus apraefrontalis (Smith, 1926)
229 Aipysurus duboisii (Bavay, 1869)
229 Aipysurus eydouxii (Gray, 1849)
232 Aipysurus foliosquama (Smith, 1926)
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232 Aipysurus fuscus (Tschudi, 1837)

233 Aipysurus laevis (Lacèpéde, 1804)
235 Aipysurus mosaicus (Sanders et al, 2012)
236 Aipysurus pooleorum (Smith, 1974)
237 Aipysurus tenuis (Lönnberg and Anderson, 1913)
239 Emydocephalus annulatus Krefft, 1869
239 Emydocephalus ijimae Stejneger, 1898
241 Emydocephalus orarius Nankivell, et al. 2020
242 Parahydrophis mertoni (Roux, 1910)
243 Ephalophis greyae (Smith, 1931)
243 Hydrelaps darwiniensis (Boulenger, 1896)
244 Hydrophis annandalei (Laidlaw, 1901)
244 Hydrophis anomalus (Schmidt, 1852)
245 Hydrophis atriceps (Günther, 1864)
246 Hydrophis belcheri (Gray, 1849)
246 Hydrophis bituberculatus (Peters, 1872)
247 Hydrophis brookii (Günther, 1872)
248 Hydrophis caerulescens (Shaw, 1802)
249 Hydrophis cantoris (Günther, 1864)
249 Hydrophis coggeri (Kharin, 1984)
250 Hydrophis curtus (Shaw, 1802)
252 Hydrophis cyanocinctus (Daudin, 1803)
254 Hydrophis czeblukovi (Kharin, 1984)
254 Hydrophis donaldi (Ukuwela, Sanders, and Fry 2012)
255 Hydrophis elegans (Gray, 1842)
256 Hydrophis fasciatus (Schneider, 1799)
257 Hydrophis gracilis (Shaw, 1802)
258 Hydrophis hardwickii Gray 1834
259 Hydrophis hendersoni Boulenger 1903
259 Hydrophis inornatus (Gray, 1849)
260 Hydrophis jerdonii (Gray, 1849)
261 Hydrophis kingii (Boulenger, 1896)
261 Hydrophis klossi (Boulenger, 1912)
262 Hydrophis laboutei (Rassumussen & Ineich, 2000)
262 Hydrophis lamberti (Smith, 1917)
263 Hydrophis lapemoides (Gray, 1849)
264 Hydrophis macdowelli (Kharin, 1983)
265 Hydrophis major (Shaw, 1802)
266 Hydrophis mamillaris (Daudin, 1803)
266 Hydrophis melanocephalus (Gray, 1849)
267 Hydrophis melanosoma (Günther, 1864)
268 Hydrophis nigrocinctus (Daudin, 1803)
269 Hydrophis obscurus (Daudin, 1803)
270 Hydrophis ornatus (Gray, 1842)
271 Hydrophis ocellatus Gray 1849
271 Hydrophis pachyceros Fisher 1855
271 Hydrophis parviceps Smith 1935
272 Hydrophis pacificus (Boulenger, 1896)

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273 Hydrophis peronii (Duméril, 1853)

274 Hydrophis platurus (Linnaeus, 1766)
277 Hydrophis schistosus (Daudin, 1803)
279 Hydrophis semperi Garman, 1881
280 Hydrophis sibauensis (Rasmussen et al., 2001)
281 Hydrophis spiralis (Shaw, 1802)
282 Hydrophis stokesii (Gray, 1846)
283 Hydrophis stricticollis Günther, 1864
284 Hydrophis torquatus Günther, 1864
285 Hydrophis viperinus (Schmidt, 1852).
286 Hydrophis vorisi (Kharin, 1984)
286 Hydrophis zweifeli (Kharin, 1985)

288 14. Radiant Snakes and Gemsnakes

288 Lycodonomorphus bicolor (Günther, 1893)
289 Lycodonomorphus laevissimus (Günther, 1862)
290 Lycodonomorphus leleupi (Laurent, 1950)
291 Lycodonomorphus mlanjensis Loveridge, 1953
291 Lycodonomorphus obscuriventris Fitzsimons, 1963
291 Lycodonomorphus rufulus (Lichtenstein, 1823)
292 Lycodonomorphus subtaeniatus Laurent, 1954
292 Lycodonomorphus whytii (Boulenger, 1897)
293 Boaedon upembae (Laurent, 1954)
295 Liopholidophis sexlineatus (Günther, 1882)
296 Thamnosophis lateralis Duméril, Bibron and Duméril, 1854
296 Thamnosophis stumpffi (Boettger, 1881)

299 The Dipsadidae Section

300 15. Bamboo Snakes, Pseudoxenodontidae
300 Pseudoxenodon bambusicola Vogt, 1922
301 Pseudoxenodon baramensis (Smith, 1921)

303 16. The Basal Dipsadids

303 Thermophis baileyi (Wall, 1907)
303 Thermophis shangrila Peng et al, 2014
304 Thermophis zhaoermii Guo, Liu, Feng & He, 2008

307 17. New World Mud Snakes

307 Farancia abacura (Holbrook, 1836)
308 Farancia erytrogramma Palissot De Beauvois, 1801

311 18. Wetland Coniophanes

311 Coniophanes bipunctatus Günther, 1858
311 Coniophanes quinquevittatus Duméril, et al. 1854

313 19. Neotropical Swamp Snakes

313 Hydromorphus concolor Peters, 1859
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315 Hydromorphus dunni Slevin, 1942

315 Tretanorhinus mocquardi Bocourt, 1891
315 Tretanorhinus nigroluteus Cope, 1861
316 Tretanorhinus taeniatus Cope, 1861
316 Tretanorhinus variabilis Duméril, et al., 1854

320 20. Neotropical River Snakes

324 Hydrodynastes bicinctus Herrmann, 1804
324 Hydrodynastes gigas (Duméril et al, 1854)

327 21. Neotropical Fishing Snakes

327 Helicops angulatus (Linnaeus, 1758)
328 Helicops apiaka Kawashita-Ribeiro et al, 2013
329 Helicops boitata Moraes-da-Silva et al, 2019
329 Helicops carinicaudus (Wied-Neuwied, 1825)
329 Helicops cyclops Cope, 1869
330 Helicops danieli Amaral, 1938
330 Helicops gomesi Amaral, 1921
331 Helicops hagmanni Roux, 1910
331 Helicops infrataeniatus Jan, 1865
332 Helicops leopardinus Schlegel, 1837
333 Helicops modestus Günther, 1861
334 Helicops nentur Costa, et al., 2016
335 Helicops pastazae Shreve, 1934
335 Helicops petersi Rossman, 1976
336 Helicops phantasma Moraes-da-Silva et al, 2021
337 Helicops polylepis Günther, 1861
337 Helicops scalaris Jan, 1865
338 Helicops tapajonicus Da Frota, 2005
338 Helicops trivittatus (Gray, 1849)
338 Helicops yacu Rossman, 1975
340 Hydrops caesurus Scrocchi et al. 2005
340 Hydrops triangularis Wagler, 1824
341 Hydrops martii (Wagler, 1824)
342 Pseudoeryx plicatilis (Linnaeus, 1758)
343 Pseudoeryx relictualis Schargel et al, 2007

345 22. Neotropical Bush Snakes

345 Gomesophis brasiliensis Gomes, 1918
346 Ptychophis flavovirgatus Gomes, 1915
347 Thamnodynastes chaquensis Bergna and Álvarez, 1993
348 Thamnodynastes hypoconia (Cope, 1860)
349 Thamnodynastes lanei Bailey, Thomas, and da Silva, 2005
349 Thamnodynastes pallidus (Linnaeus, 1758)
351 Thamnodynastes ramonriveroi Manzanilla and Sánchez, 2005
352 Thamnodynastes strigatus (Günther, 1858)

354 23. The Semi-Aquatic Painted Snakes

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Aquatic Snakes

354 Lygophis anomalus (Günther, 1858)

356 Erythrolamprus miliaris (Linnaeus, 1758)
358 Erythrolamprus poecilogyrus, (Wied-Neuwied 1825)
359 Erythrolamprus semiaureus (Cope, 1862)
360 Erythrolamprus breviceps (Cope, 1860)
360 Erythrolamprus cobellus (Linnaeus, 1758)
361 Erythrolamprus frenatus (Werner, 1909)
362 Erythrolamprus jaegeri Günther, 1858
363 Erythrolamprus taeniogaster (Jan, 1863)
363 Erythrolamprus reginae (Linnaeus, 1758)
364 Erythrolamprus zweifeli (Roze, 1959)
365 Erythrolamprus macrosomus (Amaral, 1936)
366 Erythrolamprus epinephalus (Cope, 1862)
366 Erythrolamprus albiventris (Jan, 1863)
366 Erythrolamprus fraseri (Boulenger, 1894)
367 Erythrolamprus lamonae (Dunn, 1944)
367 Erythrolamprus melanotus (Shaw, 1802)
368 Sordellina punctata (Peters, 1880)

369 The Cosmopolitan Watersnake Section

372 24. Mountain Keelbacks
372 Hebius annamensis (Bourret, 1934)
373 Hebius chapaensis (Bourret, 1934)
374 Hebius craspedogaster (Boulenger, 1899)
375 Hebius deschauenseei (Taylor, 1934)
375 Hebius modestus (Günther, 1875)
376 Hebius nigriventer (Wall, 1925)
377 Hebius octolineatus (Boulenger, 1904)
378 Hebius popei (Schmidt, 1925)
379 Hebius parallelum (Boulenger, 1890)
379 Hebius sauteri (Boulenger, 1909)
380 Hebius taronensis Smith, 1940
381 Hebius venningi (Wall, 1910)
382 Herpetoreas burbrinki Guo, et al., 2014
382 Herpetoreas pealii (Sclater, 1891)
383 Herpetoreas platyceps (Blyth, 1854)
383 Herpetoreas sieboldii Günther, 1860
384 Herpetoreas tpser Ren, Jiang, Huang, David and Li, 2022
384 Herpetoreas xenura (Wall, 1907)
384 Rhabdops aquaticus Giri, et al., 2017
385 Rhabdops olivaceus (Beddome, 1863)
386 Pseudagkistrodon rudis Boulenger, 1906

389 25. Keeled Water Snakes

391 Amphiesma stolatum (Linnaeus, 1758)
393 Tropidonophis doriae Boulenger, 1897
393 Tropidonophis elongatus (Jan, 1865)
394 Tropidonophis mairii Gray, 1841
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395 Tropidonophis multiscutellatus (Brongersma, 1948)

396 Tropidonophis negrosensis (Taylor, 1917)
397 Tropidonophis punctiventris (Boettger, 1895)
398 Tropidonophis truncatus (Peters, 1863)
398 Rhabdophis auriculatus (Günther, 1858)
399 Rhabdophis lineatus (Peters, 1861)
400 Rhabdophis rhodomelas (Boie, 1827)
401 Rhabdophis tigrinus (Boie, 1826)
402 Rhabdophis formosanus (Maki, 1931)
403 Rhabdophis lateralis Berthold, 1859
403 Atretium schistosum (Daudin, 1803)
404 Fowlea schnurrenbergeri (Kramer, 1977)
405 Fowlea yunnanensis Anderson, 1879
406 Fowlea asperrimus (Boulenger, 1891)
407 Fowlea flavipunctata (Hallowell, 1860)
408 Fowlea melanzostus (Gravenhorst, 1807)
409 Fowlea piscator (Schneider, 1799)
410 Fowlea punctulatus (Günther, 1858)
411 Fowlea tytleri (Blyth, 1863)
412 Xenochrophis bellulus (Stoliczka, 1871)
413 Xenochrophis cerasogaster (Cantor, 1839)
413 Xenochrophis maculatus (Edeling, 1864)
415 Xenochrophis trianguligerus (Boie, 1827)
417 Ceratophallus vittatus (Linnaeus, 1758)

420 26. Asian Stream Snakes & Relatives

420 Smithophis arunachalensis Das et al., 2020
421 Smithophis atemporalis Giri et al., 2019
421 Smithophis bicolor (Blyth, 1854)
422 Smithophis linearis Vogel, et al. 2020
422 Opisthotropis alcalai Brown and Leviton, 1961
423 Opisthotropis andersonii (Boulenger, 1888)
424 Opisthotropis atra Günther, 1872
424 Opisthotropis cheni Zhao, 1999
424 Opisthotropis cucae David et al., 2011
424 Opisthotropis daovantieni Orlov, et al., 1998
425 Opisthotropis durandi Teynié, et al., 2014
426 Opisthotropis guangxiensis Zhao, Jiang and Huang, 1978
427 Opisthotropis hungtai Wang, et al., 2020
427 Opisthotropis haihaensis Ziegler et al., 2019
428 Opisthotropis jacobi Angel & Bourret, 1933
428 Opisthotropis kikuzatoi (Okada & Takara, 1958)
429 Opisthotropis kuatunensis Pope, 1928
430 Opisthotropis lateralis Boulenger, 1903
430 Opisthotropis latouchii (Boulenger, 1899)
431 Opisthotropis laui Yang, Sung & Chan, 2013
431 Opisthotropis maculosa Stuart & Chuaynkern, 2007
433 Opisthotropis maxwelli Boulenger, 1914

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Aquatic Snakes

433 Opisthotropis rugosa (Lidth De Jeude, 1890)

434 Opisthotropis shenzhenensis Wang et al., 2017
434 Opisthotropis spenceri Smith, 1918
435 Opisthotropis tamdaoensis Ziegler, David & Vu, 2008
436 Opisthotropis typicus (Mocquard, 1890)
436 Opisthotropis voquyi Ziegler et al., 2018
437 Opisthotropis zhaoermii Ren et al., 2017
438 Trimerodytes aequifasciatus (Barbour, 1908)
438 Trimerodytes annularis (Hallowell, 1856)
439 Trimerodytes balteatus (Cope, 1895)
440 Trimerodytes percarinatus (Boulenger, 1899)
441 Trimerodytes yapingi Guo, Zhu, and Liu, 2019
441 Trimerodytes yunnanensis Rao and Yang, 1998
441 Trimerodytes (=Paratapinophis) praemaxillaris Angel, 1929

444 27. African Water Snakes

444 Afronatrix anoscopus (Cope, 1861)
445 Natriciteres bipostocularis Broadley, 1962
446 Natriciteres fuliginoides (Günther, 1858)
447 Natriciteres olivacea (Peters, 1854)
447 Natriciteres pembana (Loveridge, 1935)
448 Natriciteres sylvatica Broadley, 1966
449 Natriciteres variegata (W. Peters, 1861)
449 Helophis schoutedeni (De Witte, 1942)
450 Hydraethiops laevis Boulenger, 1904
450 Hydraethiops melanogaster Günther, 1872
452 Limnophis bangweolicus (Mertens, 1936)
453 Limnophis bicolor Günther, 1865
453 Limnophis branchi Conradie, Deepak, Keates and Gower, 2020

455 28. The Palearctic Watersnakes

456 Natrix maura (Linnaeus, 1758)
464 Natrix natrix (Linnaeus, 1758)
464 Natrix helvetica (Lacépède, 1789)
464 Natrix astreptophora (López-Seoane, 1884)
479 Natrix tessellata (Laurenti, 1768)

490 29. The Thamnophiine Clade

491 Liodytes alleni (Garman, 1874)
492 Liodytes pygaea (Cope, 1871)
494 Liodytes rigida (Say, 1825)
495 Clonophis kirtlandii (Kennicott, 1856)
496 Regina grahamii Baird and Girard, 1853
498 Regina septemvittata (Say, 1825)
500 Nerodia cyclopion (Duméril, Bibron & Duméril, 1854)
500 Nerodia floridana (Goff, 1936)
501 Nerodia clarkii (Baird and Girard, 1853)

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504 Nerodia erythrogaster (Forster, 1771)

506 Nerodia fasciata (Linnaeus, 1785)
508 Nerodia harteri (Trapido, 1941)
509 Nerodia paucimaculata (Tinkle and Conant, 1961)
510 Nerodia rhombifer (Hallowell, 1852)
512 Nerodia sipedon (Linnaeus, 1758)
515 Nerodia taxispilota (Holbrook, 1838)
517 Thamnophis atratus Kennicott, 1860
518 Thamnophis bogerti Rossman and Burbrink, 2005
519 Thamnophis conanti Rossman and Burbrink, 2005
520 Thamnophis lineri Rossman and Burbrink, 2005
521 Thamnophis brachystoma (Cope, 1892)
522 Thamnophis butleri (Cope, 1889)
523 Thamnophis chrysocephalus (Cope, 1885)
523 Thamnophis couchii (Kennicott, 1859)
524 Thamnophis cyrtopsis (Kennicott, 1860)
526 Thamnophis elegans (Baird and Girard, 1853)
528 Thamnophis eques (Reuss, 1834)
530 Thamnophis errans Smith, 1942
531 Thamnophis exsul Rossman, 1969
532 Thamnophis fulvus (Bocourt, 1893)
533 Thamnophis gigas Fitch, 1940
534 Thamnophis godmani (Gunther, 1894)
535 Thamnophis hammondii (Kennicott, 1860)
536 Thamnophis marcianus (Baird and Girard, 1853)
537 Thamnophis melanogaster (Peters, 1864)
538 Thamnophis copei (Dugès in Cope, 1879)
539 Thamnophis foxi (Rossman and Blaney, 1968)
539 Thamnophis mendax Walker, 1955
540 Thamnophis nigronuchalis Thompson, 1957
541 Thamnophis ordinoides (Baird and Girard, 1852)
542 Thamnophis postremus Smith, 1942
542 Thamnophis proximus (Say, 1823)
544 Thamnophis pulchrilatus (Cope, 1885)
545 Thamnophis radix (Baird and Girard, 1853)
546 Thamnophis rossmani Conant, 2000
547 Thamnophis rufipunctatus (Cope, 1875)
549 Thamnophis saurita (Linnaeus, 1766)
550 Thamnophis scalaris Cope, 1861
551 Thamnophis scaliger (Jan, 1863)
553 Thamnophis sirtalis (Linnaeus, 1758)
556 Thamnophis sumichrasti (Cope, 1866)
557 Thamnophis unilabialis (Tanner, 1985)
559 Thamnophis validus (Kennicott, 1860)

563 30. Natricids of Uncertain Status

563 Anoplohydrus aemulans Werner, 1909
563 Isanophis boonsongi (Taylor and Elbel, 1958)

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Aquatic Snakes

564 Hydrablabes periops (Günther,1872)

564 Hydrablabes praefrontalis (Mocquard, 1890)

566 31. African Rain Forest Water Snakes

566 Grayia caesar Günther, 1863
567 Grayia ornata (Bocage, 1866)
569 Grayia obscura Chaney, Greenbaum and Pauwels 2023
570 Grayia smithii (Leach, 1818)
571 Grayia tholloni Mocquard, 1897

574 32. Colubridae

574 Crotaphopeltis barotseensis Broadley, 1968
574 Crotaphopeltis degeni (Boulenger, 1906)
576 Platyceps insulanus (Mertens, 1965)
577 Platyceps rhodorachis (Jan, 1863)
577 Philothamnus angolensis Bocage, 1882
578 Philothamnus hoplogaster (Günther, 1863)
579 Oocatochus rufodorsatus (Cantor, 1842)
579 Xenelaphis ellipsifer Boulenger, 1900
580 Xenelaphis hexagonotus (Cantor, 1847)

582 33. Conservation

609 Literature Cited

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Aquatic Snakes

Co-Authors to Aquatic Snakes

Justin M. Bernstein
Center for Genomics
University of Kansas

Myke Clarkson
Herpetological Conservation International
57556 29 Palms Hwy #467
Yucca Valley, CA 92284

Jason A. Fantuzzi
EcolSciences, Inc. JFantuzzi@[Link]
75 Fleetwood Dr # 250, Rockaway, NJ 07866

Daniel Jablonski
Department of Zoology,
Comenius University in Bratislava,
Ilkovičova 6, Mlynská dolina, 84215,
Bratislava, Slovakia

Michael J Jowers
Departamento de Zoología, Facultad de Ciencias
Avenida de la Fuente Nueva s/n, 18071
University of Granada, Spain

Gunther Köhler,
Senckenberg Research Institute
Frankfurt am Main, Germany

Zachary Loughman
208 University Drive
West Liberty University
West Liberty WV, 26074

Jean-Jay Mao
Department of Forestry and Natural
Resources, National Ilan University
No. 1, Sec. 1, Shennong Rd., Yilan City,
260007 Yilan County, Taiwan

Konrad Mebert
Global Biology and Viper Specialist Group IUCN
15, 5242 Birr, Switzerland
IDECC; Institute of Development
Ecology, Conservation and Cooperation
Via G. Tomasi di Lampedusa 33, 00144 Rome, Italy

15
Aquatic Snakes

John C Murphy
Science & Education
Field Musuem
Chicago, IL USA

Juan M. Pleguezuelos
Department of Zoology,
Faculty of Sciences,
Granada University
E-18071 Granada, Spain

Nathan Rusli,
Indonesia Herpetofauna Foundation
Bogor, Indonesia

Xavier Santos,
5. Research Centre for Biodiversity and Genetic Resources (CIBIO/InBIO)
University of Porto,
R. Padre Armando Quintas,
4485-661 Vairão, Portugal

Marion Segall
Natural History Museum
Cromwell Rd, South Kensington, London SW7 5BD, UK

16
 Acknowledgments

Acknowledgments
The idea for this book was initially conceived while I was working on Homalopsid
Snakes, Evolution in the Mud (2007). Discussions with Harold Voris (Field Muse-
um), Daryl Karns (Hanover College), and Myke Clarkson provided additional ideas
and motivation. Rune Midtgaard (RepFocus) generously supplied the distribution
maps from his website. The maps used in the chapter on Natrix are from Schöneberg
et al. (2023). Justin Berstein, Konrad Mebert, Myke Clarkson, Michael Jowers, and JJ
Mao provide extra assistance with editing. The photographs were made possible by
about 120 photographers from more than 30 countries. The co-authors provided the
text to their areas of specialization, species groups and species accounts. To all my
sincerest thanks. JCM

17
Aquatic Snakes

A Python bivittatus from Sakaerat Environmental Research Station, Thailand. Photography JCM

18
 Preface

Preface
I shall not pretend to assert that they are as numerous as land snakes; but it is
very likely that one hundred species at least of this tribe exist in the waters of the
ocean, lakes, and rivers.
C. S. Rafinesque 1817. Dissertation on water snakes, sea snakes and sea ser-
pents. Philosophical Transaction

When Rafinesque wrote this, 14 species of snakes were considered water snakes. The
idea for a global overview of aquatic snakes has attracted my attention since I finished the
Homalopsid Snakes, Evolution in the Mud in 2007. Books on sea snakes are available: Mal-
colm Smith’s classic The Sea-Snakes, William Dunson’s The Biology of the Sea Snakes, Harold
Heatwole’s Sea Snakes, and Ivan Ineich and Pierre Laboutes’ Sea Snakes of New Caledonia, to
name a few. However, most information on aquatic snakes remains scattered in hundreds of
journals and books.
Snakes living in fresh and brackish waters had been mostly overlooked. Snakes inhab-
iting marine environments were restricted to a relatively few lineages, while snakes using
freshwater were found in many more lineages. Furthermore, snakes use aquatic habitats in
many ways that may not cause them to be easily recognized as aquatic. For example, some
North American snakes spend the winter in water-filled crayfish burrows and are terrestrial
or semi-aquatic the rest of the year.
This volume is perhaps best considered an illustrated, annotated checklist of aquatic and
semi-aquatic snakes in marine and freshwater environments. The biggest challenge was de-
ciding what species to include and which to exclude. While the intent was to have accounts
for all aquatic snakes, where do you draw the line on the continuum? How much evidence
do you need to label a species semi-aquatic? I am confident this volume covers the range of
snakes that are traditionally considered semi-aquatic. However, knowledge of some of these
animals is poor or absent, while others are some of the best-studied reptiles.
Snakes are derived from lizards, but surprisingly few saurians are semi-aquatic, and no
extant lizards are known to be aquatic. Bauer and Jackman (2007) found 73 species in 11 fam-
ilies using freshwater habitats. They found no aquatic lizards inhabiting the Nearctic or Pale-
arctic regions. Still, the Neotropics, the Oriental, and Australian regions support the greatest
diversities of freshwater forms, particularly in the families Gymnophthalmidae, Scincidae,
and Varanidae. Pauwels et al. (2008) listed 153 snakes (about 5% of all snakes recognized at
the time) that inhabit freshwater. This number covers 44 genera in six families (Acrochordi-
dae, Boidae, Colubridae, Elapidae, Homalopsidae, and Viperidae), with the greatest numbers
in the Oriental (64 species) and Neotropical Regions (39 species).
Here, the definition of aquatic is expanded to include marine species and species that are
terrestrial, arboreal, or fossorial but enter the water and use aquatic situations and resources.
I have included species considered terrestrial or fossorial because there is some evidence, they
are also using aquatic habitats and resources. The marine, brackish, and freshwater snakes
discussed here provide an overall picture of the extant snakes using fluid habitats. This work
lists more than 420 species (about 10.3% of snakes) in 106 genera, 31 subfamilies and fami-
ly-level clades. But not all these species are aquatic or semiaquatic, and they have been includ-
ed because their close relatives use watery habitats. Note that I (JCM) compiled the chapters
with no authors listed. The species accounts are uneven because knowledge is limited on
many species, different authors have written accounts, and I have intentionally tried to keep
the size of this volume down.

19
Aquatic Snakes

A note on species and subspecies

Definitions of species and subspecies are hotly debated. While species are frequently
regarded as natural units, subspecies are considered arbitrary human constructs. This point
seems to have been first argued by Wilson and Brown (1953). Others note that the “arbi-
trary” label could be applied against any higher taxonomic rank below the phylum (Avise and
Mitchell 2007).
Zoologists have used the Linnaean system for almost three centuries, and doing away
with it and replacing it with some other system seems improbable. De Queiroz (2021) has
updated the subspecies concept to base them on the same criteria as species – separately
evolving metapopulation lineages. They differ from species in that they have not entirely
separated from each other. I have included subspecies (trinomials) in the species accounts be-
cause they are used in much of the literature – undoubtedly, some of them are invalid, others
are likely valid, and yet others are likely to be elevated to species status. Some species might
be synonymized once more detailed information will become available.

Figure P.1. A Natrix natrix from Marchegg, Austria.

Photography by Kai Kolodziej.

On the next page. The homalopsid snake Enhydris jagorii from Bung Ka Loh.
Photography by Daryl Karns.

20
 Preface

21
Aquatic Snakes

1. Introduction
Lake Songkhla looks like many other wetlands from the shoreline (Figure 1-1), but it is
the largest wetland complex in Thailand and comprises three shallow basins. One hundred
and fifty years ago, Songkhla was a bay open to the ocean. Since then, a series of barrier is-
lands were expanded with human assistance and gradually enclosed the basins. The islands
now form the Sathing-Phra Peninsula and almost wholly isolate Lake Songkhla from the Gulf
of Thailand (Figure 1-2). Only one opening to the sea is located at the south end of the lake
near Songkhla. Annandale (1916) described the lake as no more than 4.9 m deep. Since then,
the lake has continued to accumulate silt, and today, the three basins have a mean depth of
1.2 m and a maximum depth of 2.4 m (Leelawathanagoon et al. 1997). The water levels and
chemistry changes accompanying the lake’s formation left an unusual mixture of freshwater
and marine fauna.
Lake Songkhla has perhaps the most diverse assemblage of water-dwelling snakes glob-
ally. One sea snake (Hydrophis brookii family Hydrophilidae), one file snake (Acrochordus
granulatus, family Acrochordidae), the pipe snake (Cylindrophis sp. family Cylindrophiidae),
at least one species of keelback (Fowlea piscator - family Natricidae), and at least four species
in four genera of Asian-Australasian Mud Snakes (family Homalopsidae) live in the lake or
along its wet edges.
Aquatic is an easily applied label if the snake never comes out of the water, like some sea
snakes or the Tentacled Snake (Erpeton tentaculatus). They are easily labeled semi-aquatic if
they are like the North American water snakes of the genus Nerodia, the European Natrix, or
the sea kraits of the genus Laticauda. All of these enter the water to hunt or search for mates
but bask in bushes and on rocks close to the water, and some mate and lay eggs on land. Then,
there are terrestrial or arboreal snakes that escape predators by diving into the water, such as
Philodryas olfersii (Abegg and Mario-da-Rosa 2018).
Snakes have evolved aquatic lifestyles many times and to varying degrees in dozens of
different lineages. Snakes living in still, turbid water and feeding on fish can be expected to
have different adaptations than snakes feeding on crustaceans living in bedrock streams with
boulders and fast-moving water. Making this more complex and confusing are snakes that
evolved aquatic lifestyles and then returned to a terrestrial existence.
Pauwels et al. (2008) proposed that freshwater snakes forage in the water and cannot sur-
vive without aquatic prey and frequent submersion. They suggested that subtle external ana-
tomical characters betray aquatic habits in freshwater snakes. Morphological specializations
include dorsally oriented nostrils, often close together, some of which have valved nares that
exclude water from the upper respiratory system and allow the snake to inhale at the surface
without being seen by predators. The eyes, too, tend to be dorsally oriented so that a snake
lying in the water can view the sky (or the water) above for potential prey or predators. The
eyes of aquatic snakes are often reduced in size, and sometimes, the iris matches the color of
the surrounding scales. Other species have a brightly colored iris that makes the eyes stand
out compared to the surrounding scales. Aquatic snakes frequently have a dorsal pattern of
transverse bands and keeled dorsal scales.
None of these characteristics are shared by all aquatic snakes, and some traits, like trans-
verse bands or keeled scales, are shared with many terrestrial, arboreal, or exclusively marine
species. The freshwater Tentacled Snake, Erpeton tentaculatum, is highly aquatic, possibly
the most specialized freshwater snake, and may never leave the water or do so only on rare
occasions (Figure 1-3). Its eyes are relatively large and lateral, not dorsolateral or dorsal, and
it has longitudinal stripes, not transverse bands.

22
 1. Introduction

Figure 1.1. Lake Songkhla,

the largest lake in Thailand
looks very much like many
other wetlands but it supports
a diverse aquatic snake fauna.
Photography by JCM

Figure 1.2. A view of Lake

Songkhla from space is mostly
isolated from the Gulf of Thai-
land but remains connected by
an opening at the south end of
the lake. Image taken 13 Febru-
ary 2008. Photo credit: NASA,
WorldWind [Link].

23
Aquatic Snakes

Thus, the ancestors of modern aquatic snakes were adapted for life underground, on the
ground, or in the trees. Some of their descendants moved into freshwater, brackish water, or
marine environments and evolved traits for those environments. Some returned to life on
land, and others evolved into more specialized aquatic lifestyles.
Extant snakes are organized into about 45 family and subfamily-level clades extend-
ing backward in time. Estimated dates for snake origins are complicated by the evolution of
squamate snake-like morphs that appear in the fossil record. Snakes and lizards shared an
ancestor in the lower Jurassic, about 190 MYA (Harrington and Reader 2017). During this
time, snake lineages branched. Some existed for a while before becoming extinct. Others
survive to the present day.
The evolution and adaptations of marine snakes have received considerable attention
(Dunson 1975; Heatwole 1999; Sanders et al. 2008). Sea snakes are in the subfamily Hy-
drophiinae – a clade nested in the larger elapid clade. All share fixed front fangs and venom
glands, including coral snakes, cobras, and mambas. Marine elapids have three ancestors
and have adapted to life in the oceans differently. They form two or three lineages: the sea
snakes (Hydrophis + Apisuryus = hydrophiids) with 62 species and the sea kraits with eight
species in the genus Laticauda. Both clades have paddle-like tails to improve their swimming
ability, but the hydrophiids give birth to young in the water and have lost the broad ventral
scales. Sea kraits are oviparous, laying eggs in terrestrial situations, and retain their broad
ventral scales. The only thing sea kraits do in the water is hunt. Despite living in the oceans,
sea snakes and sea kraits still rely on freshwater for drinking (Lillywhite et al. 2008; Bonnet,
Brischoux 2008; Lillywhite et al. 2014). This requires the sea snakes to stay close to river del-
tas or areas of heavy rainfall. Freshwater draining off the land into the ocean remains on the
surface before it mixes with seawater, and rainwater floats on the ocean’s surface and forms a
lens of freshwater before mixing with the heavier seawater.
Ambush predators can use water for concealment while retaining much of the terrestrial
morphology for life on land. Other snakes hunt shorelines and search small pools without
entering the water, and a few snakes hunt from branches and strike aquatic prey from a perch
over the water. Arboreal snakes can use aquatic resources without the apparent morphologi-
cal modifications to aquatic habitats, a reminder of the remarkable plasticity we see in snakes.
Asad et al. (2020) reported that the mock viper, Psammodynastes pictus, is restricted to Sun-
daland and hunts fish of the genus Rasbora from branches overhanging the water. Trobisch
and Gläßer-Trobisch (2011) observed hunting behavior in captive juvenile Natrix tessellata
and reported hunting from branches over the water. The Burmese Vine Snake, Ahaetulla
fronticincta, is highly arboreal but feeds exclusively on fish (Figure 1-4). It inhabits bushes
and other low vegetation in mangrove forests in coastal Myanmar (Wogan and Vogel, 2020).
Snakes that hunt from branches or shorelines without entering the water are not includ-
ed here because, technically, they do not spend much time in the water. However, snakes that
hunt from the water are included – even though many of them are habitat generalists. And
maybe taking prey from the shoreline
Not surprisingly, the largest snakes often use water. The Green Anaconda (Eunectes mu-
rinus) and its relatives are highly aquatic. The largest pythons (Python sebae, P. natalensis, P.
bivittatus, P. molurus, and Malayopython reticulatus) are less dependent on water than the
anacondas (aquatic boas). Yet the pythons still spend substantial time in the water, hunt from
the water, and use the water for thermoregulation. Not only does water provide buoyancy
for massive bodies, but it allows the snakes to conceal themselves from prey and predators.
One sea snake, the Yellow-bellied Sea Snake (Hydrophis platurus), has become pelag-
ic, drifting with ocean currents (Figure 1-5). The only time it is seen on land is when it

24
 1. Introduction

Figure 1.3. The freshwater Tentacled Snake, Erpeton tentaculum.

Photography by Alex Mason.

Figure 1.4. Gunther’s Whipsnake Snake, Ahaetulla fronticincta, is highly arboreal but feeds
exclusively on fish.
Photography by Paul Freed.

25
Aquatic Snakes

washes up on beaches from Africa’s east

coast to the coastlines of western North
and Central America. The Yellow-bellied
Sea Snake aggregates along slicks or drift
lines. Floating debris accumulates in the
slicks, and the snake may remain for days
or weeks before a change in wind speed
or current direction breaks them up. Ag-
gregations of Hydrophis platurus drift, the
aggregate, may contain just a few animals
or several thousand, including juvenile and
adult snakes. Other animals inhabiting the
drifts are jellyfish medusa, fish, porpoises,
sea turtles, and sea birds, often following
floating debris lines. Snakes aggregate
here because the slicks are a place to lo-
cate food and mates. The degree to which
these snakes have adapted to the marine
environment is significant, given that they
represent a recent evolutionary radiation.

Figure 1-5. The Yellow-bellied Sea

Snake, Hydrophis platurus, has be- Was the snake ancestor aquatic?
come pelagic, drifting with ocean
currents. Photography by May
Zabe.
Evidence from morphology and molecules leaves lit-
tle doubt that snakes evolved from lizards. However, the
transition between lizards and snakes is among the most
controversial topics in evolution, partly because of the
lack of well-preserved fossils. Three competing hypoth-
eses for the lifestyle of the ancestral snake -- burrowing,
aquatic, and terrestrial origins -- have been debated for
more than a century.
Cope (1869) implied the snakes’ ancestors could have
been aquatic when he placed snakes and mosasaurs (Fig.
1-6) in the Pythonomorpha. Caldwell and Lee (1997), Ca-
prette et al. (2004), and others who have worked on Cre-
taceous marine lizards also suggested an aquatic snake
Figure 1.6. Reconstruction of liv- ancestor. Vidal and Hedges (2004) tried to rule out an
ing Mosasaurus missouriensis. The aquatic origin for snakes based on the molecular results
fossil remains were discovered in
suggesting snakes were not related to varanoid lizards –
2012 in a layer of shale by a min-
ing company in southern Alberta the mosasauroids’ closest living relatives.
in Canada. The lizard was about Camp (1923) proposed snakes evolved from a grass-
6.5 meters long with a skull that land anguimorph lizard (a terrestrial ancestor). Today,
was slightly less than a meter long. anguimorph lizards remain possible candidates for the
Artist credit: Nobu Tamura. snake ancestor, and there are living grass-swimming an-
guimorphs. Anguimorph lizards and snakes share paired
male reproductive organs with ornamentation and venom
glands.
26
 1. Introduction

Mahendra (1938) proposed that the snake ancestor was a burrowing lizard. This hypoth-
esis gained broad support and was the most well-accepted idea regarding the snake’s ancestral
lifestyle. Mahendra noted that a ring of scales surrounded the eye in most advanced snakes.
Burrowing forms like the threadsnakes and blindsnakes (Leptotyphlopidae and Typhlopidae),
the shield-tailed snakes (Uropeltidae), and the Amazonian pipe snakes (Anilius) have the eye
covered by a single large scale. Mahendra viewed all these snakes as primitive or basal to
more advanced snakes. He placed them at the evolutionary tree’s base and concluded that
the ancestral snake must have been fossorial. Today, these snakes are recognized as some of
the oldest lineages, the dawn snakes, threadsnakes, and blindsnakes — and have been called
scolecophidians – but they are not monophyletic (Heise et al. 1995).
An early terrestrial relative of snakes was the 95-million-year-old Najash rionegrina,
which had two rear legs and a sacrum. The remains of Najash included partial skulls. Howev-
er, other fossil snakes possessed hindlimbs, but their pelvic bones were not directly attached
to the vertebrae. These are older than Najash; all were marine (Haasiophis, Pachyrhachis, and
Eupodophis).
The widespread acceptance of the burrowing ancestor hypothesis was partly due to Walls’
(1940) work on snake eye anatomy. In a list of differences between snake and lizard eyes, Walls
clarifies that snake eye anatomy is quite distinct from lizard eye anatomy. The most dramatic
difference between the lizard and snake eyes is the snakes’ inability to change the shape of
their lenses to focus an image onto the retina. Muscles attached to the lens for focusing in liz-
ards are absent in snakes. Snakes must move the lens towards or away from the retina to focus
their eyes using the enlarged peripheral iris muscles. Also, most snakes lack a fovea, although
some Asian tree snakes have re-evolved the structure. Walls also explained the cone-like struc-
ture of rods in snake retinas (and those of other tetrapods) resulting from the loss of rods in
response to diurnal activity and then secondarily redeveloping rods from cones. He thought
cone cells were transformed into rod cells, which became known as the transmutation hypoth-
esis. The modifications Walls (1940) found in snake eyes suggested to him they resulted from
a burrowing lifestyle.
Shine and Wall (2008) found two ecomorphs for lizards that had lost their legs: legless
burrowing lizards have elongated trunks, small heads, short tails, and constant body widths;
and surface-active legless species have short trunks, broader heads, long tails, and more vari-
able body widths. They suggested that a long tail in surface dwellers helps escape predators if
the tail can be automized. Long tails are probably beneficial for locomotion when the animal
uses lateral undulations (side-to-side movement). On the other hand, burrowing lizards need
to push through the substrate, which requires a different arrangement of muscles and bones
in which the rib cage and trunk muscles form a rigid but flexible cylinder. This pipe-like mor-
phology transfers force to the digging head, and a long tapering tail is not well suited for this
lifestyle.
Regarding the aquatic origins of snakes, Shine and Wall observe that living semi-aquatic
lizards have well-developed limbs and long tails. The tails are essential in swimming and,
therefore, are unlikely to be lost. Many living lizard species are burrowers, and by implication,
Shine and Wall infer that snakes were more likely to have evolved from a burrowing ancestor
than an aquatic one.
Molecular clock dates produced by Vidal et al. (2007) suggest lizards last shared an ances-
tor with snakes 166.4 MYA. Other authors using similar methods estimated that this ances-
tor lived 194–145 MYA. Molecular clock dates suggest the scolecophidians (leptotyphlopids,
typhlopids, and anomalepidids) shared an ancestor with all other snakes about 155.6–151.9
MYA. Hugall et al. (2007) placed the snake - Anguimorph lizard divergence at about 155 MYA

27
Aquatic Snakes

Figure1.7. Threadsnakes and Blind Snakes have been long considered to be models for the ancestral
snake Photography by JCM.
and the separation of the scolecophidians from the rest of the snakes at about 105 MYA.
Sanders et al. (2008) place the anguimorph-snake divergence date at about 140 MYA and the
divergence of the scolecophidians from the rest of the snakes at about 110 MYA.
Yi and Norell (2015) argued in favor of snakes’ burrowing origin based on inner ear
morphology. They noted that the fossil snake Dinilysia is a stem snake close to the most
recent common ancestor of living snakes; according to most phylogenetic analyses, it has
inner ear morphology like some burrowing squamates. Their model predicted that both
Dinilysia and the reconstructed ‘hypothetical ancestor of crown snakes’ are classified as bur-
rowing forms based on a comparison with modern species of known ecology. Burrowing is
a predominant lifestyle in most basal lineages of crown snakes; therefore, snakes must have

Figure 1.8. The glass lizards may be readily mistaken for a snake. They are anguimorph lizards.
Photography by JCM.

28
 1. Introduction

had a burrowing origin.

Again, looking at inner ear structure, Palci et al. (2017) used a larger dataset with more
taxa and subdivided the species into five traditional ecological categories. They recognized
that the closest outgroup to snakes might be iguanians, anguimorphs, or mosasauroids. The
results placed Dinilysia close to the existing snake Xenopeltis and found that homalopsids
were close to Dinilysia. The placement contradicts the claim that a sizeable spherical sacculus
(their ‘vestibule’) is diagnostic of burrowing forms. Such morphology can also be found in
semi-aquatic snakes, and it is absent in some typical obligate burrowers like scolecophidians
or semifossorial snakes such as Calabaria and Brachyurophis.
Scolecophidians are often considered the most basal lineage(s) of crown snakes, histor-
ically and in recent phylogenetic analyses (Figure 1.7). Supposing this is an accurate phylo-
genetic position for these snakes, it might be expected that Dinilysia would display an inner
ear morphology like scolecophidian snakes if burrowing were indeed primitive for snakes.
However, the sharp differences between the inner ears of Dinilysia and scolecophidians are
not easily understandable if all share a burrowing ancestry.
The ancestral snake’s lifestyle remains uncertain; it could have been a burrower, a swim-
mer, a terrestrial species, or a species using a combination of these ecological lifestyles. Fur-
ther discussion of this topic awaits additional fossil discoveries and studies of extant species.
Ancient Aquatic Snakes
The Eocene began at 55.8 MYA and ended at 33.9 MYA. During this time, the continents
were drifting toward their present positions. It was a time when the Earth was a "greenhouse,"
with global warming due to increased atmospheric carbon dioxide (Bohaty and Zachos 2003).
The distribution of Eocene species was remarkably different than what we see today. There
were palm trees in Alaska and the northern Rockies, crocodiles on Elsmere Island above the
Arctic Circle, and forests covering much of Antarctica. Primates spread from Asia to Europe
and North America during this time, and the rivers, estuaries, and oceans contained snakes,
including giant aquatic snakes
The snake family Palaeopheidae is known exclusively for their vertebrae and ribs. They
inhabited both hemispheres, from the Upper Cretaceous to the Eocene (McDowell 1987).
Their fossils are always associated with rock formed in watery environments. The vertebrae
tend to be tall and narrow, and the ribs are only slightly curved, characteristics found in the
most aquatic snakes living today, the sea snakes. Those who study Palaeopheidae fossils con-
sider the group relatives of the boids, but others have suggested that they are close to the file
snakes (the acrochordids).
Size estimates for the paleophids range from half a meter to nine meters or more, and
while some lived in near-shore environments such as estuaries and mangroves, others were
using open ocean habitats far from shore. Holman (2000) and Rage et al. (2003) have re-
viewed the family, known chiefly from vertebrae and ribs, and have described paleophids as
booid-like.
Some species were highly modified for life in the water. They had laterally compressed
vertebrae and bodies for swimming, high neural spines on all vertebrae (to aid the lateral
compression of the body), synapophyses for rib articulation low on the vertebrae, and ribs
that were only slightly curved (assisting lateral body compression). The vertebrae of at least
one species contained marrow cavities, which may have been involved in regulating buoy-
ancy, increased erythrocytes, or both. Moreover, like some modern sea snakes, the laterally
compressed species likely could not move on land. Other family members showed only slight
modifications for aquatic life and were probably somewhat terrestrial.

29
Aquatic Snakes

The extinct colubroid families Anomalophiidae and Russellophiidae are also suspect-
ed of being aquatic (Rage et al. 2003). Palaeophis colossaeus was described by Rage (1983)
based on vertebrae collected in Mali. The vertebrae were 34 mm long. We do not know how
many vertebrae the snake had, but given that a typical boa or python has about 270, it is
likely this snake could have been 9.180 m or more than 30 feet. Another huge paleophiid is
Pterosphenus schucherti, described from coastal North America. It ranged from Texas to New
Jersey. Remains from Florida indicate that an individual snake died at least 300 km from the
nearest mainland during the late Eocene, where it was buried with cartilaginous fish, bony
fish, and an ancient whale.
Ancient aquatic snakes disappeared near the end of the Eocene, and their extinction
is likely linked to a changing climate. Global cooling at the start of the Oligocene occurred
as oceanic circulation was altered with Antarctica's disconnection from Australia and South
America. The formation of the South Pole's continental ice sheet followed the extinction of
many warm-water species
Four new fossil aquatic snake species ranging from 167–143 MYA in age were described
by Caldwell et al. (2015). These four species push the earliest known snakes backward by
nearly 70 million years - into the mid-Mesozoic. Thus, their origin coincided with the known
radiation of most other major groups of squamates in the mid-Jurassic: the time of the final
stages in the break-up of Pangaea into Laurasia and Gondwana. These new records for early
snakes fill a significant chronological gap predicted by molecular phylogenetics.
In stratigraphic order, the fossil snakes Caldwell et al. recognized are Parviraptor es-
tesi, from rocks dated at ~167 MYA, from the Middle Jurassic, from Southern England,
followed by the North American Diablophis gilmorei dated at ~155 MYA from the Upper
Jurassic of Colorado, USA; which appears to be a contemporary of Portugalophis lignites
from the Upper Jurassic of Guimarota, Portugal; the youngest species was Eophis under-
woodi from rocks dated at ~150–140 MYA positioned in time at the Jurassic-Cretaceous
border from an outcropping near Swanage, Dorset, Southern England. Portugalophis lig-
nites remains were deposited in a coal swamp. Eophis and Parviraptor remains came from
mixed coastal lakes, pond systems, and riparian environments, and Diablophis was recov-
ered from an epicontinental deposit several hundred kilometers from the nearest shoreline.
Caldwell et al. wrote,
"It is also possible that snakes forming these island assemblages arrived as
secondarily aquatic invaders. Secondary invasions of marine environments
characterize the subsequent evolutionary histories of numerous clades of fossil
and modern snakes." Thus, some of the earliest known snakes – were likely
semi-aquatic or aquatic.
Cretaceous (91–95 MYA) were also aquatic. Pachyophis woodwardi and Pachyrhachis prob-
lematicus, had laterally compressed bodies, small heads, and pachyostoic ribs and vertebrae
(thick, dense bone lacking marrow). These fossils came from carbonate rocks deposited in an
inter-reef basin, suggesting they lived in shallow water on a carbonate platform (Caldwell and
Albino, 2001). A group of at least three species (P. problematicus, Haasiophis terrasanctus, and
Eupodophis descouensi) had hind limbs. Their remains come from marine deposits in the Med-
iterranean area of the ancient Tethys Ocean or its immediate vicinity (Rage and Escuilié, 2002).
Not all Cretaceous snakes were aquatic. Apesteguia and Zaher (2006) describe Najash rione-
grina, from the Cenomanian (Upper Cretaceous) of Rio Negro Province, Argentina. Najash
is unique among snakes (fossil and living) in that it has two sacral vertebrae that separate
the trunk vertebrae with ribs from the caudal vertebrae without ribs. Thus, this animal has a

30
 1. Introduction

Fig. 1.9. The Green Anaconda, Eunectes murinus, (Family Boidae) is the largest living snake and
it is highly aquatic. Photography by William W. Lamar.

pelvis, and articulating with the sacrum were robust functional legs. Additionally, this spe-
cies was from a terrestrial deposit, with the skull showing characteristics associated with an
underground lifestyle. The authors wrote that it was perhaps "…a surface-dwelling species
that would occasionally use tunnels produced by burrowers."
The fossil remains of the boid, Titanoboa cerrejonensis, from La Guajira in northeastern
Colombia, imply the snake reached 12.8 m and a weight of 1,135 kg. It is the largest snake
ever discovered, and it was aquatic. The remains were dated 58 to 60 MYA (Head et al. 2009).

Extant Aquatic Snakes

Extant aquatic snakes of all lineages reach their highest diversity in tropical South-
east Asia and Australasia (Heatwole 1999). However, among the advanced terrestrial
snakes, some boodontines, colubrines, dipsadids, and natricids use coastal environments
and enter brackish water or full seawater. Virtually all other marine species are restrict-
ed to Asia and Australia. The only exception is the pelagic hydrophiid Hydrophis (Pela-
mis) platurus, which has dispersed into the eastern Pacific and the western Indian Ocean).
Without a doubt, Asia and Australasia have the most remarkable diversity of aquatic species.
At first, this seems to be due to the abundance of shallow-water habitats. However, the
answer may have more to do with the geographical origins of advanced snake lineages. The
best-known and most committed aquatic snakes are the fixed front-fanged, paddle-tailed
sea snakes (Hydrophiinae: Elapidae), which spend all or most of their life in water and have
two or more terrestrial elapid ancestors. The unusual, semi-aquatic sea kraits (Laticauda:

31
Aquatic Snakes

Figure 1.10. Phylogeny of the extant snakes from Pyron and Burbrink (2012) Clades with a blue
check mark contain at least one aquatic or semi-aquatic species. This demostrates that aquatic
life styles evolved in at least 16 subfamily and family level clades over the past 150 million years.

32
 1. Introduction

Elapidae) have paddle tails, broad ventral scales,

and terrestrial tick parasites, and they only enter
the water to feed. Instead, sea kraits bask, mate,
and deposit their eggs in terrestrial locations.
The file snakes (Acrochordidae) seem to
spend their entire lives in the water and show
numerous adaptations to an aquatic lifestyle, sug-
gesting they, and their ancestors, have been liv-
ing in the water for an exceptionally long time.
Some Oriental-Australian rear-fanged water
snakes (Homalopsidae) give the impression that
they spend slightly less time in the water than sea
snakes and file snakes.
And, as previously mentioned, many of the na-
tricids (Natricidae or Natricinae) are semi-aquatic.
However, most species' time in the water is poorly
known. This is also the case for many species in
other lineages, snakes considered semi-aquatic.
Life in the water changes the rules snakes face on
land. Figure 1.11. A preserved specimen of the
Water supports an animal's body weight and homalopsid Cerberus schneiderii. Note the
is slow to lose or gain heat. Salt in the ocean ren- dorsally oriented eyes and nostrils. The nos-
ders the water undrinkable for snakes and many tril valve on the left side is open, the nostril
other animals. Salt also increases the density of valve on the right side is closed. Photogra-
phy by JCM.
water, making animals more buoyant. Warm wa-
ter reduces the need for basking, but some aquatic
snakes live in relatively cold water that originates
on mountaintops. Water may lower the risk of predation from terrestrial and aerial predators.
Of course, aquatic environments expose their inhabitants to a new set of predators, parasites,
and physical environmental challenges.
Aquatic habitats provide abundant food resources in the form of invertebrates, fish, and
amphibians and present a variety of new hiding places, such as the intertidal burrow system, the
cavities under a riverbank, or the layer of detritus covering the bottom of the pond or stream.
All snakes are likely capable of swimming, but some species are much better at it than others.
Many snake lineages have at least a few species adapted to aquatic habitats. Still, five lineag-
es (file snakes, the homalopsids, elapids, dipsadids, and the natricids) contain the most species
adapted for life in water. Snake adaptations to water can include modifications to many or all organ
systems. However, all adaptations for an aquatic existence do not occur in every species of aquat-
ic snake. Some species have extreme adaptations to life in water, others appear to have very few.
Dorsally positioned nostrils and eyes allow the snake to breathe and observe without ex-
posing the head or body to the air. When diving, valves open and close the nostrils, and the
trachea opens opposite the internal nares to exclude water from the respiratory tract.
Lateral compression of the body increases the surface for swimming; the ribs may be
less bowed and elevated to exaggerate the surface area on the snake's sides. The tail may be
flattened, compressed slightly at the base, or turned into a paddle with exaggerated fin-like
flaps above and below the vertebrae. Ventral scales are broad in land-dwelling snakes so the
snakes can grip surfaces; in some highly aquatic snakes, the ventral scales narrow (i.e., the file
snakes), and the belly scales are similar in size to the dorsal scales. This reduction in ventral

33
Aquatic Snakes

scales aids the snake in flattening its body for an increased surface area when swimming. The
snakes most adapted for aquatic life swallow food while submerged and give birth in the water.
Neill (1958) summarized the literature and included personal observations on snakes using
brackish and seawater. In some cases, the snakes are directly observed in the water. In other
instances, he assumes their saltwater use based upon their proximity to it. Many subsequent
papers also discuss this phenomenon. Thus at least some semi-aquatic or mostly freshwater
snakes can tolerate high concentrations of sodium ions in their blood (hypernatremia), or
avoid absorbing them. This allows them to venture into brackish and marine environments to
forage, find mates, or disperse into new habitats for at least a short time. Even some terrestrial
species are occasionally seen swimming in the ocean. The list is extensive. The following is
organized by family and based on Neill (1958) and other more recent literature.
Tropidophiidae. Neill also found the Cuban trope, Tropidophis m. maculatus, beneath
debris at the edge of weed-grown tidal flats near Havana, Cuba.
Booids. Candoia carinata, Island distribution; Neil found one on Morotai Island, Mo-
luccas, under masses of rooted coconuts at the bases of coconut trees growing above the tidal
zone and notes one specimen was found swimming in the sea at Natterer Bay, New Guinea.
An African Sand Boa (Eryx jaculus) was found beneath logs close to the beach in Bougie,
Algeria.
Pythonidae. Hart et al. (2012) found that hatchling Python bivittatus could survive for
five months in brackish water and for more than a month in full seawater.
Colubridae. The Florida kingsnake (Lampropeltis getula floridana) Carr (1940) listed it
as “occasional” in a salt marsh. A close ally, the speckled kingsnake (L. g. holbrooki) was found
to be as common in the salt marsh as in the woods of southeastern Texas.
Psammophiidae. The European Malpolon monspessulanus has been reported multiple
times from brackish and saltwater habitats (Deso et al 2021).
Natricidae. Populations of these snakes adjacent to brackish or marine environments
can be expected to occur in saline waters on occasion. The green water snake (Nerodia cy-

Figure 1.12, Natrix tessellata.

Photography by Kai Kolodziej.

34
 1. Introduction

Figure 1.13. Acrochordus arafurae, Gregory River.

Photography by Scott Eipper.
clopion), the Brown Water Snake, (Nerodia taxispilota), the Glossy Water Snake (Lithodytes
rigida), and Graham’s Water Snake (Regina. grahami) in brackish marshes. The European
water snake (Natrix natrix) has been reported in the sea along the Mediterranean coast. The
Viperine Water Snake (Natrix maura), the Eastern Garter Snake (Thamnophis s. sirtalis) the
Ribbon Snake (T. sauritus) are frequent visitors to saline water. (Neill 1958)
Dipsadidae. The North American Mud Snake (Farancia abacura) and the Rainbow
Snake (Farancia erytrogramma) use tidal marshes. While the aquatic neotropical genus Tre-
tanorhinus uses freshwater habitats, some inhabit mangroves.
Elapidae. The Indian cobra (Naja naja) is an adaptable but essentially terrestrial species,
occurring in southern Asia’s forests, fields, gardens, and human-modified situations. It enters
the water readily and has been observed swimming in the sea (Gharpurey, 1954, p. 54). The
Australian black snake (Pseudechis porphyriacus) has been found swimming in the middle of
Botany Bay. The Australian taipan (Oxyuranus scutellatus) is abundant around the shores of
the Gulf of Carpentaria. And other Australian elapids (species not named) may swim across
estuaries (Kinghorn, 1956). And, of course, marine hydrophiids can survive in full seawater
as long as they have access to freshwater for drinking.
Viperinae. The Common Viper (Vipera berus) is terrestrial but swims readily and has
been found many miles at sea off the coast of Ireland (M. A. Smith, 1951).
Crotalinae. Some rattlesnakes (Sistrurus and Crotalus) use saline habitats. These include
the Dusky Pygmy Rattlesnake (Sistrurus miliarius barbouri) and the Eastern Diamondback
Rattlesnake (Crotalus adamanteus).
Thus, freshwater snakes may take a first step into transitioning into brackish or salt wa-
ter by exploring those habitats. In the reverse situation, brackish water or marine snakes may
transition back to freshwater. Cerberus microlepis, Laticauda crockeri, and Hydrophis semperi
are in genera that are mostly marine but those species live in freshwater

35
Aquatic Snakes

Terrestrial and Freshwater Snakes Visit the Oceans

Coastal populations of snakes that usually inhabit freshwater or terrestrial habitats may
venture into brackish or full seawater. Nerodia spp. and in particular Natrix tessellata have
evolved brackish- and low saline tolerant seawater populations (e.g., Mebert 2004, 2005;
Tuniyev et al. 2011; Brischoux et al. 2017; Pauwels et al. 2020; Rautsaw et al. 2020). Snakes
may get washed downstream into the ocean or brackish water habitats during flooding. Re-
ports of Reticulated Pythons, Eastern Diamondback Rattlesnakes, and many other species
swimming between islands are documented. They suggest that these species may intention-
ally enter the oceans (a hypertonic environment) because they will find food, mates, or a
new territory.
Figure 1.14 shows a Thamnophis ordinoides on the Oregon coast just north of Port Or-
ford in August 2012. The snake traveled a considerable distance to reach the ocean. The mo-
tivation was unknown. Was the snake lost or intentionally entering the marine environment?
This individual snake was not washed down a stream during a flooding event (the evidence
is the long track in the sand). Nor was it transported to the edge of the Pacific Ocean by
humans.

Figure 1.14. (A) A photo from the parking lot taken the same day I (SB) saw the snake. (B) The
second photo shows the view from the beach towards the ocean. (C) The third photo shows the
snake tracts go right into the ocean. (D-E) The snake tracks in more detail. (F) The snake, a
Thamnophis ordinoides. Photography by Sabine Baer.
On the next page. The major lineages containing numerous aquatic species are Homalopsidae, the
elapid subfamily Hydrophiinae, some dipsadids, and the natricids. All clades have independently
evolved species with a viviparous reproductive mode, and the evolution of viviparity could have
occurred multiple times within a clade. The image on the following page illustrates eggs with devel-
oping embryos from an Enhydris enhydris. Photography by Daryl Karns

36
Aquatic Snakes

37
Aquatic Snakes

2. Aquatic Snake Biology

Marion Segall
Acquiring an aquatic lifestyle is associated with physical and chemical constraints relat-
ed to the properties of the water bodies used by vertebrates. The physicochemical properties
of water affect many aspects of life, from their locomotion to sensory signal propagation
and homeostasis (Houssaye & Fish, 2016; Vogel, 1994). These extrinsic constraints are driv-
ing the evolution of aquatic snakes and have led to convergences (e.g., (Gazzola et al., 2014;
Stayton, 2011; Vincent et al., 2009) and some specific adaptations. As mentioned in this
book, acquiring an aquatic lifestyle has evolved multiple times independently in snakes all
over the globe, and species use the aquatic medium to different extents. For all these rea-
sons, aquatic snakes are a great model to study evolutionary convergences and adaptations.
This chapter aims to overview the convergences and specificities of aquatic snakes, from
their anatomy to their behavior.

Feeding and Drinking

Feeding underwater, and especially prey capture behavior, is mainly constrained by the
density and viscosity of water; these hydrodynamic forces work to oppose a strike. The head
of snakes, and especially its external morphology, is directly affected by those constraints.
The head of aquatic-foraging snakes presents convergent features such as dorsally positioned
eyes and nostrils, a short, stout, and a dorso-ventrally flattened head (Esquerré & Keogh,
2016; Segall et al., 2016). The dorsally positioned nostril allows aquatic snakes to breathe
at the water’s surface without exposing their head. Aquatic snakes have evolved cutaneous
respiration to some degree. In Hydrophis, up to 23% of its oxygen intake is cutaneous. This
led to increased vascularization of their head and brain (Palci et al., 2019). However, snakes’
primary oxygen intake process remains pulmonary respiration (Heatwole & Seymour, 1975;
Pratt & Franklin, 2010; Udyawer et al., 2016). The dorsal position of their eyes allows pred-
ator and prey detection from above (Heatwole, 1999; Trapp & Mebert, 2011); it should be
most beneficial for bottom-dwelling ambush species. In addition, the dorsal position of the
eye may lead to an increase in binocular vision, which produces a better 3D image and more
accurately targets prey (see the section on the sensory systems). The short, stout head, with
a barely visible head-neck transition, provides aquatic snakes with a more streamlined body
and allows them to reduce the forces opposing an open-mouth strike underwater (Segall et
al., 2019). However, aquatic adaptation in snakes seems related to the extent to which they
use water suggesting potential evolutionary trade-offs between aquatic and terrestrial life-
styles. For instance, the head shape of aquatic snakes is more hydrodynamic and efficient at
capturing prey than one of semi-aquatic species (Segall et al., 2020).
The relationship between head shape hydrodynamics and prey capture success has not
been established yet as we lack data on most behavioral aquatic snake species, especially in a
natural context. Aquatic snakes are elusive, and field observations of their foraging behavior
are rare or insufficient to provide quantitative data or comparative analysis. However, the
diet of aquatic snakes is well-known for many species. It is diverse, from generalist/oppor-
tunistic snakes fishing for any prey (i.e., Cerberus) to highly specialized species eating only
specific species of eels (i.e., Laticauda sp.), fish eggs, or crustaceans.
Dietary specialization in aquatic snakes is associated with morphological adaptations.
In homalopsids, crustaceans specialists have broad heads and short jaws, while piscivorous
species have longer, slender heads and jaws (Fabre et al., 2016), a pattern that seems to apply
38
 2. Aquatic Snake Biology

Figure 2.1: Morphological variability in the skull of aquatic snakes. The phylogeny has been adapted
from Pyron et al. (2014), color indicates snake families. The silhouettes of the main items in the diet
are indicated on the right: fish, anurans, crustaceans, freshly molted crustaceans (black and white),
fish-eggs. Skulls from CT scanned specimens (top to bottom) AMNH-R11970, AMNH-R115651,
FMNH229747, CAS12391, FMNH229816, AMNH-R152305, AMNH-R161755, AMNH-R14169,
AMNH-R161752,AMNH-R19316, AMNH-R159304, AMNH-R160212, AMNH-R101360,
AMNH-R162317, AMNH-R103188.

39
Aquatic Snakes

to most aquatic snakes (see below). In hydrophiines, specialization toward burrowing prey
has led to the independent evolution of microcephalic species (Sherratt et al., 2018, 2019).
Microcephalic sea snakes specializing in eels show convergence and high rates of evolution.
However, microcephalic goby-specialists do not. While both are fish, eels and gobies present
distinct challenges for snakes: eels are elongated, fierce prey, probably associated with a high
manipulation cost, and require an efficient swallowing process.
In contrast, gobies are bulky prey that requires a large gape size through the maintenance
of larger head width. This functional hypothesis has been validated on a phylogenetically
large sample of aquatic snakes. Yet, the difference in head shape between the two dietary
categories (i.e., bulky vs. elongated) is not striking, especially in light of the great head shape
variability between all aquatic snakes (Segall et al., 2020). More data on the cost associated
with feeding on different prey are needed, such as manipulation time, maximum gape size, or
the number of pterygoid walks required to swallow the prey. Such data will establish the con-
tribution of functional constraints related to diet on the head shape of snakes. Unfortunately,
for most aquatic species, such behavioral data are lacking.
Morphological adaptations have been found in the skull bones used in aquatic snakes.
Most snakes have a hyperkinetic skull composed of over 20 independent bones, and 16 are
used in feeding. A study on a phylogenetically large sample of aquatically feeding snakes
showed these bones could be divided into functional modules associated with prey capture,
restraint, and swallowing (Rhoda et al., 2020). The variation in the shape of each module is
related to dietary specialization, either toward durophagy or piscivory. Durophagy in aquatic
snakes consists of a dietary specialization toward crustaceans and evolved independently in
homalopsids and North American natricids. Anatomically, this specialization is associated
with a shorter lower jaw providing a better bite force and a reduction of the number of teeth
and smaller and stouter teeth, while piscivorous species have long slender jaws with numer-
ous, long, and sharp teeth that allow them to generate fast prey capture and get a good grip
on slippery prey (Figure 2.1)(Rhoda et al., 2020; Savitzky, 1983).
There are two types of crustacean predators in both snake families, the hard-shelled
(Liodytes, Fordonia, Cantoria) and the soft-shelled (Regina, Gerarda) specialists. While the
adaptations described above apply to all these species, their “sub-specialization” has led to
additional morphological convergences. The hard-shelled specialists have blunter teeth,
thickened stomachs, larger gapes, and consume smaller prey (Jayne et al., 2018). Unlike their
non-durophagous relatives, crustaceans specialists, except R. septemvittata, use their bodies
to restrain for capturing and handling their prey, allowing them to reduce their consumption
time (Gripshover & Jayne, 2021; Tumlison & Roberts, 2018).
Some crustacean feeders have evolved a unique feeding behavior known as crab-ripping
that consists of dismembering the prey’s legs and chelipeds. In the case of Gerarda, the whole
body of large prey, is torn apart, thus circumventing the gape-size limitation (Gripshover &
Jayne, 2021; Jayne et al., 2018). Snakes usually swallow their prey whole, limiting the size of
the prey they can ingest (Gans, 1974). Besides the crustacean specialists, the only report of
food processing in snakes is the evisceration of large toads by the colubrid, Oligodon fascio-
latus (Bringsøe et al., 2020). Other dietary specializations in aquatic feeding snakes have led
to spectacular anatomical adaptations, such as the Hydrophiine snakes that feed on fish eggs
such as Emydocephalus sp., Aipysurus eydouxii, or Aipysurus mosaicus. These species have
reduced teeth (Fig. 2.1) and derived labial scales for scraping fish eggs from the substrate
(Guinea, 1996; Shine, Bonnet, et al., 2004), reduced fangs, atrophied venom glands, and low
venom quantity and toxicity (Gopalakrishnakone & Kochva, 1990; Healy et al., 2019; Li et
al., 2005).

40
 2. Aquatic Snake Biology

Among aquatic snake lineages, several species independently invaded saline environ-
ments (Murphy, 2012) and hence have to maintain their water balance. Snakes living in
marine environments need a source of fresh water. This dependency is reflected in their
distribution along coasts or estuaries (Brischoux et al., 2012; Lillywhite & Tu, 2011; Mur-
phy, 2012) and their rainfall-dependent behavior and distributions (Bonnet & Brischoux,
2008; Lillywhite et al., 2008). The few species that live in the open ocean, such as Hydrophis
platurus, do not have access to freshwater from land. Still, they drink from rainwater lens-
es that form at the ocean’s surface during the rainy season and suffer severe dehydration
during seasonal drought (Lillywhite et al., 2019). Piscivorus snakes suffer further dehydra-
tion from a diet of fish that generates an excess of nitrogen that must be excreted through
urine, thus increasing the need for freshwater (Lillywhite & Ellis, 1994). When snakes are
too dehydrated, they stop eating, probably to avoid additional water loss (Edwards et al.,
2021). The dehydration tolerance in marine snakes seems to depend on their access to
freshwater and a gradient of habitat specialization.
Estuarine species like Nerodia (Edwards et al., 2021), Acrochordus: (Lillywhite et al.,
2014), Laticauda: (Lillywhite et al., 2008), and hydrophiines (Lillywhite et al., 2015) have a
lower tolerance than sea snakes. In addition, marine snakes have less permeable skin allow-
ing them to decrease water loss through skin evaporation (Edwards et al., 2021; Lillywhite
et al., 2009). To cope with salt and dehydration, sea snakes, acrochordids and at least one
homalopsid, Cerberus rynchops, have evolved supralabial or sublingual salt glands (Dunson
& Dunson, 1973, 1979). Combined with cloacal excretions, they can reduce the excess salt
they absorb. However, unlike other marine vertebrates that maintain normal natremia, ma-
rine snakes can sustain high salt concentrations (Brischoux et al., 2013) and seem only to
use their glands when natremia reaches critical levels. Therefore, osmoreceptors probably
alter the snake the snake’s condition (Dunson et al., 1971).

Sensory System
Snakes possess an arsenal of senses, from the light and UV spectra, olfaction, and vom-
erolfaction to mechanoreception and even audio-reception. However, air and water have
different physical properties that affect signal propagation, such as refractive index, vis-
cosity, and density. Thus, the sensory system and the preferred sensory modalities used by
species depend on the media they use. The sensory system of aquatic snakes presents con-
siderable differences from their terrestrial relatives and some novelties, such as phototactic
tails. Among hydrophiines, most of the species of Aipysurus show a behavioral response to
light stimulation on their tail, especially in the green and blue spectra, a feature found in
other elongated aquatic vertebrates (Crowe-Riddell, Simões, et al., 2019). Other elapid sea
snakes show no sensitivity to light along their body, neither does Aipysurus mosaicus. The
ancestral origin of this different sense remains to be determined. The adaptive value of a
photosensitivity body is not well understood, but the tail of sea snakes resembles a paddle
and increases their swimming efficiency (see the section on locomotion). Phototaxis could
be an adaptive feature if it decreases the risk of tail injuries or predation. However, there is
no evidence of how this compares to other sea snakes. The tail of sea snakes also presents
scale organs of unknown functions. These may involve mechanoreception (Crowe-Riddell,
Williams, et al., 2019). However, compared to the mechanosensitive scale organs found on
the head of sea snakes, the ones on the tail have a simpler structure that suggests less sen-
sitivity. Other dermal structures in the tail could be involved in sensory modalities such as
phototaxis, but further investigations are needed.

41
Aquatic Snakes

Highly aquatic snakes present various degrees of mechanoreceptor coverage, from the
entire body and shark-like skin of acrochordids (Figure 2.2)(Povel & Van Der Kooij, 1997)
to the tentacles of
The tentacles of Erpeton (Catania et al., 2010) and the scale sensilla on the head of hy-
drophiines (Crowe-Riddell et al., 2016; Crowe-Riddell, Williams, et al., 2019) detect water
movement. These kinds of sensory organs have recently been characterized in Helicops sp.
(García-Cobos et al., 2018). Similar mechanosensitive scale organs are present in terrestrial
snakes. Still, these scale sensilla are more protruding and more abundant in sea snakes, indi-
cating a better sensitivity than their terrestrial relatives (Crowe-Riddell et al., 2016). However,
experiments on Hydrophis (Lapemis) curtus revealed that they can detect a low amplitude
pressure variation they might use to detect a nearby swimming fish. However, this might have
lower sensitivity than other aquatic animals (Westhoff et al., 2005). Similar results were found
with the hearing abilities of sea snakes (Chapuis et al., 2019). Yet, some highly aquatic species
rely on mechanosensitive cues to detect and catch their prey. Acrochordids, for instance,
catch prey with their whole body, with no visual cues (pers. obs.; Dowling, 1960). Erpeton
tentaculatum uses its mechanosensitive tentacles to detect prey in the absence of light, yet,
this species seems to rely on both visual and mechanical cues for prey detection (Catania et
al., 2010). Mechanoreception in other aquatic snakes has not been functionally investigated.
Still, Subsessor bocourti can strike fish without visual information if the prey touches any part
of its body. This suggests that their skin is sensitive to touch and can elicit a prey-capture be-
havior (MS pers. obs.). Many aquatic snake species use tactile cues, natricids, acrochordids,
homalopsids, and even hydrophiines (Drummond, 1979; Heatwole, 1999; Jayne et al., 2018;
Rossman et al., 1996; Shine, Brown, et al., 2004; Voris et al., 1978). More controlled experi-
ments to evaluate the sensory modalities aquatic snakes use are needed as mechanoreceptors
could be sensitive to water pressure or tactile stimuli (Crowe-Riddell et al., 2016).
Aquatic snakes most likely use several senses to detect and catch their prey, as in Erpeton.
Previous works demonstrated that the preferred sensory modalities used by aquatic snakes
depend on many factors, such as their diet, their prey capture strategy, and the availability
of sensory cues (Kutsuma et al., 2018; Vincent et al., 2005; Waters & Burghardt, 2013). Yet,
because of the properties of water, the senses of aquatic snakes have functionally diverged
from their terrestrial relatives. The most striking –albeit obvious— example of these adap-
tations is the loss of vomerolfaction and olfaction. Vomerolfaction and olfaction are crucial
to trail airborne compounds in terrestrial snakes (Halpern et al., 1997) and to elicit tongue
flicks if no other sensory cues are available (Zuri & Halpern, 2003). Because they cannot use
their primary olfactory system underwater, aquatic sea snakes have lost most of their olfac-
tory receptor genes (Kishida et al., 2019), while the amphibious Laticaudinae retain most of
these genes (Kishida & Hikida, 2010). Despite losing this chemosensory system, most aquatic
snake taxa rely on vomerolfaction and olfaction. Its efficiency underwater largely depends
on the ecology of species. Although tongue-flicks in some aquatic species, such as acrochor-
dids or hydrophiines, are discrete thanks to an elongation of their long and thin tines (pers.
obs., Gove, 1979; Heatwole, 1999), the flicking behavior might warn prey of the predator’s
presence. This may be a trigger some species have exploited to their advantage by using a
“tongue-fishing” technique. Such lingual luring has been reported in semi-aquatic piscivores
(Figure 2.3). In North America, natricids, such as Thamnophis atratus (Welsh & Lind, 2000),
Nerodia rhombifer and Nerodia sipedon (Czaplicki & Porter, 1974), Nerodia clarkii compres-
sicauda (Hansknecht, 2008; Hansknecht & Burghardt, 2010) use lingual luring to attract
fish. The behavior was recently reported in Thamnodynastes strigatus, a semi-arboreal South
American dipsadid that catches fish while hanging on branches (Mario-Da-Rosa et al., 2020).

42
 2. Aquatic Snake Biology

This peculiar strategy is used in

an ambush context in the pres-
ence of visual and chemical cues
(Hansknecht & Burghardt, 2010).
The availability and reliabil-
ity of both chemical and visual
cues depend on the environment’s
properties and the species’ ecolo-
gy. For instance, aquatic ambush
predators that need a fast and
traceable signal to strike quickly
and accurately cannot only rely
on chemical cues to detect prey.
Because the diffusion of mole-
cules underwater is too slow in
Fig.2.2. The strongly keeled scales of Acrochordus. Pho-
lentic environments (e.g., ponds,
tography by JCM
lakes, swamps) or too fast and un-
traceable in lotic environments (e.g.,
streams, rivers). Snakes using ambush strategies might therefore rely more upon visual cues.
But active foragers can use chemical signals’ excellent persistence and traceability underwa-
ter to trail prey over long distances. Such behavior-dependent sensory modalities have been
demonstrated in Acrochordus arafurae; active-foraging males use chemical cues, while am-
bush females use vision (Vincent et al., 2005). Generally, piscivores species rely mostly on vi-
sual and chemical cues depending on the properties of the environment (Catania et al., 2010;
Czaplicki & Porter, 1974; Kutsuma et al., 2018; Shine, Brown, et al., 2004; Udyawer et al.,
2020). Crustacean and fish-egg specialists rely on chemical cues (Mark Waters & Burghardt,
2005; Shine, Bonnet, et al., 2004).
Interestingly, some aquatic snakes, such as acrochordids and hydrophiines have a pecu-
liar tongue morphology and flicking behavior that consists of protruding only their long and
thin tines. While terrestrial snakes usually extend the fused base of their tongue (MS pers.
obs., Gove 1979; Heatwole 1987). Counterintuitively, snakes do not use their forked tongue
to stereo-sample chemicals from their environment, and they can trail prey with a single tine
(Parker et al., 2008). However, flicking their forked tongue generates a recirculation of the
flow to get a better sample of volatile molecules by “mixing” the air (Ryerson & Schwenk,
2014). How the peculiar tongue morphology and flicking behavior of aquatic snakes affect
the chemical sampling in these species has yet to be investigated. Unfortunately, we lack com-
parative data for the tongue morphology and flicking behavior of most aquatic snakes despite
the relative ease of access to these data and the recent efforts to maintain aquatic snakes in
captivity.
The importance of visual cues in aquatic snakes might have been underestimated. Most
snakes respond to visual cues, epecially moving objects (Catania et al., 2010; Drummond,
1985; Shine, 2005), but their acuity is relatively poor (Drummond, 1985; Hart et al., 2012),
albeit better than some small mammals (Baker et al., 2007). Using visual cues probably de-
pends on the properties of water, such as turbidity or depth, on the activity period during
which species forage, the foraging strategy, and the size of the eyes and their respective po-
sition. The size and position of the eyes determine the binocular vision field (i.e., the area
where the visual field of each eye overlaps), which allows for depth perception. The dorsally
positioned eyes of aquatic snakes might grant them a larger binocular field of vision, thus

43
Aquatic Snakes

enhancing their visual-spatial resolution, which would benefit species living in complex
habitats. Yet, those have not been measured in most aquatic snakes. Among hydrophiines,
species foraging among rocks (i.e., Aipysurus laevis) seem to have a visual acuity slightly
better than their “bottom cruising” relatives (i.e., H. curtus, H. major) (Hart et al., 2012).
Despite their limited acuity, aquatic snakes present adaptations of their visual system, such
as the accommodation mechanisms in semi-aquatic Thamnophis, Nerodia, and Natrix
(Schaeffel & de Queiroz, 1990; Schaeffel & Mathis, 1991) or the adaptive convergence of the
diversification of visual pigments in hydrophiines that provide them with a better vision in
the parts of the visible spectrum in their environment (Simões et al., 2020). Similarities have
been found in the visual genotypes of Hydrophis and Helicops, namely a polymorphism in
the pigment opsin. At the same site is a unique feature among reptiles that, if the two alleles
were expressed, it could provide aquatic snakes with better color vision (Hauzman et al.,
2017; Simões et al., 2015; Simões et al., 2020). Still, additional work is needed to validate
this functional hypothesis.
The importance of the contribution of vomerolfaction, olfaction, and vision in aquatic
snakes is reflected in the endocranial space allocated to both senses. The olfactory and
optic tract morphology are related to aquatic species’ activity period. Diurnal snakes have
an optic area larger than nocturnal species, while the olfactory tract is bulkier in nocturnal
snakes. Many species, however, can be active day and night and show an intermediate mor-
phology (Segall et al., 2021). Besides its reflection in the endocranium, the activity period
is related to eye size, with diurnal snakes having larger eyes (Da Silva et al., 2017; Liu et al.,
2012). However, aquatic snakes also have a relatively thick spectacle (Da Silva et al., 2017)
which does not seem to prevent light transmission (Simões et al., 2016) and might protect
the eyes from fluid pressure. It would be interesting to test whether the spectacle thickness
depends on the maximum pressure experienced by snakes. Or if it depends on the maxi-
mum depth encountered during diving. Or, does it depend on the physical properties of the
media, such as the viscosity and density of the water used by “mud” snakes. Or the speed of
the stream flow for snakes living in rivers.
Finally, while the sense of balance has rarely been functionally investigated in snakes,
it is likely to play a crucial role in these limbless tetrapods, especially those moving in a 3D
environment that offers no (or negligible) body support, such as water. Balance control is
essential in locomotion and is therefore developed in the following section.
Locomotion
All aquatic snakes use undulations to move underwater; they generate a wave that
propagates from their head and extends to their tail thus inducing forward motion at a
constant frequency. However, the amplitude of the wave increases along the body (Fran-
cois Brischoux et al., 2010; Jayne, 1985). Some sea snakes can swim backward by generat-
ing reverse undulations (tail-to-head), probably similar to the backward swimming of eels
(D’Août & Aerts, 1999). The pelagic sea snake, Hydrophis platurus, often uses this behavior
to chase fish swimming nearby. The specific eel-like morphology and the paddle-like tail of
sea snakes probably facilitate this backward motion. Yet, the homalopsid Subsessor bocourti,
which does not have a specialized tail seems able to swim backward (MS pers. obs.)
Snakes show various degrees of aquatic adaptation depending on how much time they
spend in the water. More aquatic species, such as acrochordids or hydrophiines (Lillywhite,
1996; Shine & Houston, 1993), present a lateral compression of their body that increases
their swimming efficiency (Brischoux & Shine, 2011). In addition, most sea snakes can ex-
pand their skin ventrally. At the same time, the semi-aquatic natricine, Nerodia sipedon, can

44
 2. Aquatic Snake Biology

actively compress its body lat-

erally through rib movements
to increase the body surface
that is used to generate pro-
pulsion during swimming
(Pattishall & Cundall, 2008).
The skin of aquatic snakes
also demonstrates several de-
grees of adaptations related to
locomotion and habitat use
(Martínez et al., 2021). Most
highly aquatic snakes’ bodies
are uniformly covered in small,
rough scales, even their belly.
While the hydrodynamic ad-
vantage of such skin roughness
has not been characterized in
snakes, the ridges in shark skin
are associated with drag reduc-
tion and thrust enhancement
(Oeffner & Lauder, 2012). An-
other –yet to be functionally
tested—morphological char-
acteristic providing aquatic
snakes with a hydrodynamic
advantage is their almost ab-
sent head-neck transition. Un-
like most other snakes, except
fossorial species (see Discus-
sion), the tips of the mandibles
are barely noticeable with-
out palpation, giving aquatic
species a streamlined profile,
which might prevent the for-
mation of vortices at the rear
of the jaw, thus reducing drag.
While snakes’ elongated,
slender body is hydrodynami-
cally advantageous as it reduc-
es pressure drag, the absence
of limbs requires specific ad-
aptations to prevent their body
from rolling - absent a hard Figure 2.3. Nerodia tongue luring.
substrate to stabilize them. To Photography by Kerry A. Hansknecht
control their buoyancy and
balance, aquatic snakes have
evolved anatomical specifici-
ties, such as an elongated lung,

45
Aquatic Snakes

Figure 2.4. Water moccasin, Agkistrodon piscivorus, (A) displaying and (B) coiling at the water’s
surface. Photography by Seth LaGrange.

that can extend up to 80% of the total body length of the animal (Cook & Brischoux, 2014;
Jayne, 1985; Lillywhite et al., 2012). Some semi-aquatic species, like the heavy-bodied wa-
ter moccasin Agkistrodon piscivorus, demonstrate remarkable buoyancy and balance control
while swimming and resting at the surface (Herault et al., 2021) (Fig. 2.x). However, more
aquatic species, such as Nerodia, sink when they stop swimming (Jayne, 1985). Such impres-
sive stability without a solid substrate requires fine sensorimotor control of balance.
Several sensory organs achieve balance in snakes—the inner ear, sense linear and rota-
tional accelerations of the head. The somatosensory system comprises mechanoreceptors on
the joints, muscles, and tendons. And, along the spinal cord, which is sensitive to stretching,
generates a proportional reflex response (Crowe, 1992). In addition, the proprioceptive sys-
tem and the mechanoreceptors in the skin may provide information on pressure. Some stud-

46
 2. Aquatic Snake Biology

ies have demonstrated a relationship between the different structures involved in balance:
the inner ear (Palci et al., 2017; Yi & Norell, 2015), skin mechanoreceptors (Crowe-Riddell
et al., 2016), the cerebellum (Macrì et al., 2019; Segall et al., 2021) and visual system (Simões
et al., 2020) vary with the ecology and behavior of aquatic species. However, we know little
about the functional relevance of those anatomical variations, nor do we see the contribu-
tion of the different systems in maintaining balance and their available integration.
Discussion
Aquatic snakes are a fantastic study system for many reasons. First, independently ac-
quiring an aquatic lifestyle during the evolutionary history of snakes and their wide geo-
graphic distribution make them a fertile ground for studying morphological and function-
al convergences and, more generally, in our understanding of the evolutionary success of
snakes through exceptional adaptability. Second, their ecological and behavioral richness,
from semi-aquatic and semi-arboreal species such as Psammodynastes or Grayia, that catch
fish from branches to the aquatic sea snakes that scrape fish-egg, Aipysurus eydouxii or Em-
ydocephalus sp., or the semi-fossorial homalopsids Brachyorhos, Calamophis, and Hypsicop-
sis or the natricids Regina and Liodytes that exploit burrow systems, provide an opportunity
for the study of niche invasion or habitat adaptation. Finally, despite a growing scientific
interest, there is much to discover about aquatic snakes’ ecology, behavior and evolution.
Among these various topics, attention should be drawn to the natural history of these
snakes as these data are the base for most evolutionary studies and could help us unravel the
ecological origins of snakes. This debate is fueled by the many convergences –or at least re-
semblances—between fossorial and aquatic species. For example, fossorial snakes also bear
sensilla on the head. However, flatter than those of aquatic species (Povel & Van Der Kooij,
1997), they have thick spectacles (Da Silva et al., 2017) and small eyes (Liu et al., 2012), no
noticeable head-neck transition, dorsally positioned eyes and a short, stout and flat head
(Esquerré & Keogh, 2016). Besides these resemblances, some aquatic species use burrows
from crustaceans or coral reefs to forage or hide, which might have led to selective pres-
sure toward more streamlined (or smaller) heads (Fabre et al., 2016; Sherratt et al., 2018).
Therefore, classifying these species as aquatic might result in biased conclusions regarding
the ecological origin of snakes.
Similarly, fossorial habits are attributed to species with distinct ecologies, from leaf-lit-
ter snakes, burrow-users, and active burrowers. In contrast, each ecology is probably asso-
ciated with different mechanical constraints, resulting in various degrees of selective pres-
sure. Palci and colleagues emphasized this issue in their study on the inner ear of snakes
(Palci et al., 2017).
We need a better understanding of the ecology of extant snakes and how they inter-
act with their environment to understand what drives their evolution. For instance, one
might consider that aquatic snakes roughly share the same ecology as they use water as a
medium. Yet, each aquatic environment has its constraints, from quiet ponds to streams,
brackish water to the deep ocean. Each habitat impacts the biology of snakes and should
be differentiated while investigating the evolution of aquatic organisms. Drawing a more
accurate description of the body of water they inhabit, along with their habitat use (i.e., to
what extent the animals use water as a resource) would allow us to understand the evolution
of aquatic snakes better. Knowledge of habitat use and the quantifiable aspects of ecology
would increase our understanding of snakes’ evolution and ecological origins.
To conclude this chapter with a thought, or more precisely, a lead that might de-
serve attention to shed light on the origin of snakes. The key might lie in semi-fossorial/

47
Aquatic Snakes

semi-aquatic species such as Cylindrophis ruffus, homalopsids such as Fordonia, Gerarda,

Cantoria, Myron, and the natricid Liodytes, as well as other aquatic species that use burrows
or actively burrow in the mud. These species’ inner ear and endocranium demonstrate simi-
larities (Allemand et al., 2017; Palci et al., 2017) and an intriguing convergence with the fossil
species Dinilysia patagonica (Palci et al., 2017).

On the next page. The homalopsid Ferania sieboldi inhabits

the large rivers that drain the Himalayas. Photography by
Avrajjal Ghosh.

48
Aquatic Snakes

49
Aquatic Snakes

3. Aquatic Snake Geography

Lakes, rivers, swamps, marshes, peatlands, mangroves, estuaries, and oceans provide
habitats for much of Earth’s biodiversity. Marine, brackish, and saline ground water compose
97% of all the water on Earth. Freshwater is only 2.5% of the planet’s water. Most freshwater is
in the polar ice caps, groundwater, and soil moisture. Liquid freshwater at the surface is 0.01%
of Earth’s total water in lakes, swamps, and rivers. Lakes contain about 87% of fresh surface
water, including 29% in the African Great Lakes, 22% in Lake Baikal in Russia, 21% in the
North American Great Lakes, and 14% in other lakes. Swamps have most of the remaining
freshwater, followed by rivers, most notably the Amazon River. The atmosphere contains a
mere 0.04% water (USGS, WWW).
The areas of the globe and the aquatic snakes they support are summarized here by con-
tinents, island groups, and oceans. Maps in this chapter were made using ESRI and the World
Surface Water Layer.

North America and the Caribbean

Aquatic North American snakes represent three family/subfamily level clades, Dipsadi-
dae, Natricidae, and Viperidae.
Dipsadidae snakes are diverse in Central and South America. But relatively few species
occur in North America, and only Farancia is aquatic. Grazziotin et al. (2012) found the phy-
logenetic placement of North American dipsadids unstable and controversial.
There are three semi-aquatic vipers in North America. First, the Massasauga Rattlesnake,
Sistrurus cantenatus, uses an unconventional aquatic habitat – flooded crayfish burrows for
over-wintering. But they also use mammal burrows and logs for over-wintering. Secondly,
the two semi-aquatic viper species are the Northern Cottonmouth, Agkistrodon piscivorus,
and the Southern Cottonmouth, A. conanti. These snakes spend much of their time in shal-
low water swamps, catching fish, frogs, and other prey. Wüster et al. (2008) found the New
World pitvipers diverged from their Asiatic sister group at approximately 24 MYA, and the
first radiation of the New World clade occurred at about 22 MYA.
Most North American aquatic snakes are in the family Natricidae, and aquatic and
semi-aquatic taxa occur in five genera forming the clade Thamnophiini. Clonophis, Nerodia,
Liodytes, Regina, and Thamnophis. These species shared an ancestor with the Eurasian genus
Natrix about 29 MYA, and radiated about 12 MYA in North America (Depaak et al. 2022).
The Thamnophiini inhabit the most extensive wetlands in North America, the western
US, and Mexico. All of these species are viviparous
The Gulf Coastal Plain extends from the southeast coast of the US to southern Texas and
into Mexico and expands inland at some locations for hundred miles. The region covers over
250 million square kilometers. The area has high biodiversity and was long overlooked as a
global biodiversity hotspot. Noss et al. (2015) showed the Gulf Coastal Plain is older, more
climatically stable than usually assumed, and rich in species, some of which are endemics.
They consider the region a biodiversity hotspot. The Lower Mississippi Valley (including the
delta) was North America’s largest forested wetland ecosystem. The floodplain extends from
the confluence of the Mississippi and Ohio rivers at Cairo, Illinois, to the Gulf of Mexico. The
drainage basin encompasses 41% of the continental United States. The floodplain is a mosaic
of ridges, swales, meander belts, and swamps that support a diverse and ecologically rich
forested wetland ecosystem.

50
 3. Aquatic Snake Geography

Figure 3.1. Wetlands and rivers in the Canada and the USA.

The area also includes the Everglades of southern Florida – a shallow, slow-moving river
from Lake Okeechobee to Florida Bay dominated by grass. Water levels drop during the dry
season except for the deeper locations. As a result, the region supports rich biodiversity and
many invasive species that arrived via the pet trade, including the highly publicized invasive
Burmese Python.
The Cuatro Ciénegas Basin is a Chihuahuan Desert wetland in Coahuila, Mexico. It
includes several natural springs that supply over 200 small ponds and wetlands, covering
1500 square kilometers. The wetlands are part of the UNESCO-designated Cuatro Ciénegas
biosphere reserve. The basin has long been known for its exceptional biodiversity, including
North America’s most significant number of endemic species (Stein et al., 2000). The region is
a biological island on the Sierra Madre Oriental’s northern extension and ranges in elevation
from about 740 m on the grassy basin floor to above 3,000 m in the surrounding mountains.
Abundant freshwater originates in springs that emerge from hundreds of interconnected,
spring-fed pools and spring-fed lakes with endemic fish, aquatic snails, and crustaceans.
Semi-aquatic snakes found here are in the genera Thamnophis and Nerodia.
To the south, the trans-volcanic belt in southern Mexico supports seven endemic
semi-aquatic Thamnophis eques populations that have been recognized as subspecies.
The Gulf of California has about 300 estuaries and other wetlands and is the northern-

51
Aquatic Snakes

Figure 3.2. The wetlands and rivers of Middle America.

most limit for the distribution of mangroves in the Eastern Pacific. On the west coast of the
Gulf of California, mangroves are distributed from the Cape region to the center of the Baja
California peninsula. These are primarily in small bays, estuaries, and isolated pockets. On
the Gulf ’s eastern side, mangrove forests are distributed southward from Tiburon Island in
Sonora to Sinaloa and Nayarit in large coastal lagoons with extensive mangroves. On the
Pacific side, the largest mangrove forests occur inside the coastal lagoons of Magdalena Bay.
(Aburto-Oropeza et al. 2008).
The region contains the endemic semi-aquatic garter snake, Thamnophis validus, a poly-
typic species that use mangroves and freshwater habitats. The area also supports the Yel-
low-bellied Sea Snake, Hydrophis platurus, an elapid. It is unclear if Hydrophis platurus is a
resident population that migrates in and out of the area or is composed of waifs.
Relatively few semi-aquatic and aquatic snakes occur in Central America. The snake
genera containing aquatic species are the tropidophiid genus Trachyboa is represented in
Panama; the dipsadid genera: Hydromorphus, Tretanorhinus, and Erythrolamprus, and the
natricid genera Nerodia, and Thamnophis. Central America covers about half a million km2
with steep topography concentrating wetlands in lowland and coastal areas. Lake Nicaragua is
the largest body of fresh water in Central America. The limestone plain of the Peten in north-
ern Guatemala has a mosaic of lakes, rivers, and wetlands, including the Laguna del Tigre, a
vast expanse of palm forests, flooded forests, grasslands, and mudflats, which together form
the largest Central American wetland. Coastal Belize has a chain of saltwater and freshwater
lagoons surrounded by flooded forests, mangroves, and swamp forests, along much of its
coastline. The Caribbean coasts of Honduras, Nicaragua, and Costa Rica have tidal lagoons
with mangroves, swamp forests, and palm swamps that stretch inland and cover much of the
landscape. Mangroves occur on the Atlantic and Pacific coasts, and the largest concentrations
occur in the Gulf of Fonseca, Nicoya, and the Gulf of Panama, Hydrophis platura is a likely
resident. Unconfirmed records of the sea krait Laticauda colubrina from the Pacific coasts of
several Central American countries: Nicaragua (Villa, 1962), Mexico (Alvarez de Toro, 1982),
and El Salvador are also noted by Cogger and Heatwole (2006).
There are no sea snakes in the Caribbean (but see Shuntov 1965; Hernández- Cama-
cho et al. 2006) or Atlantic, and very few aquatic snakes on Caribbean islands. However, the
Mangrove Water Snake, Nerodia clarki (a natricid), is in the Bahamas, and the tropical water
snake, Tretanorhinus variabilis (a dipsadid), is present in the Cayman Islands, Cuba, and on

52
 3. Aquatic Snake Geography

some islands off the coast of Central America. Trinidad’s continental fauna has six aquatic
species, the booid Eunectes murinus, the dipsadids Hydrops triangularis, Helicops angula-
tus, two species of Erythrolamprus (one of these is found on Tobago), and one species of
Thamnodynastes.

South America
South America has a rich aquatic snake fauna, undoubtedly due to the large river ba-
sins of the Orinoco and Amazon and many smaller rivers, and the ancient Pebas Wetlands.
The Orinoco River Delta of Venezuela and Colombia covers 36,000 square kilome-
ters and is dominated by a brackish shoreline of mangrove forests. The Llanos are on the
western side of the Orinoco River in western Venezuela and northern Colombia. It covers
450,000 square kilometers and is one of South America’s most extensive inland wetlands.
The llanos have a winter dry season with a summer wet season that produces annual flood-
ing. Savannas dominate the region.
Aquatic snakes in the region are numerous and include members of the genus Anilius
and aquatic burrowing species estimated to have a minimum age of 72.1 Ma (Head 2015).
booid genus Eunectes, and the dipsadid genera Erythrolamprus, Hydrops, Helicops, Hydro-
dynastes, Pseudoeryx, Tretanorhinus and the elapid genus Micrurus.
Reynolds et al. (2013, 2014) found the anacondas of the genus Eunectes to be the sister
of Epicrates. Eunectes appear to have evolved by the middle Miocene (Hoffstetter and Rage,
1977; Hsiou and Albino, 2009) based on the fossil remains of Eunectes stirtoni. Epicrates and
Eunectes diverged about 25.3 Ma (19.2–32.5 Ma). Thus, Eunectes appears to have separated
from South American Epicrates in the Oligocene (19.2–32.5 Ma), followed by subsequent
diversification into the four extant species.
There are two species of aquatic coral snakes (Elapidae), Micrurus surinamensis in the
Amazon basin and M. natteri in the Orinoco basin.
The aquatic dipsadid genera include Erythrolamprus, Hydrops, Helicops, Pseudoeryx,
and Hydrodynastes. The genus Erythrolamprus contains 55 species, of which 48 occur in
South America. While most are terrestrial, at least a few are semi-aquatic. They tend to feed
on fish, frogs, and lizards. They are oviparous and tend to be less than a meter in length. The
genus is in the clade Xenodontini.
The clade Hydropsini includes three aquatic genera Hydrops, Helicops, and Pseudoeryx.
Hydrops contains three species restricted to South America east of the Andes and the con-
tinental island of Trinidad.
Hydrops martii and H. triangularis have overlapping distributions and occur in syn-
topy, especially along the central Amazon basin and the Iquitos region. Hydrops caesurus
has a distribution outside of the Amazon basin. Hydrops martii is restricted to tropical and
subtropical moist broadleaf forests. The distribution of H. triangularis is associated with
tropical and subtropical moist broadleaf forests, and it is present in tropical and subtrop-
ical grasslands, savannas, and shrublands in Venezuela and Bolivia. The distribution of
H. caesurus is associated with flooded grasslands and savannas, in Brazil, Paraguay, and
Argentina. The distribution does not overlap with the other two species. Ecological barriers
of river basins likely drive the distributions of these species. One biome, tropical and sub-
tropical dry broadleaf forest, is between the northernmost localities of H. caesurus and the
southernmost localities of H. triangularis. Neither species has been recorded in this biome
(von May et al. 2019).
Helicops contains 20 species with distributions in South America, mostly east of the
Andes. The genus is poorly studied but interesting because some populations have been
53
Aquatic Snakes

Figure 3.3. The wetlands and rivers of South America.

54
 3. Aquatic Snake Geography

suggested to have facultative viviparity and cryptic diversity (Murphy et al. 2020; Schöneberg
and Köhler 2022).
The genus Pseudoeryx occurs in Amazon lowland rainforests and the Chaco of northern
Argentina and Paraguay (Giraudo and Scrocchi 2002) it is poorly known. However, the dis-
covery of a second species of Pseudoeryx, in the Lake Maracaibo Basin supports previously
hypothesized paleogeographic reconstructions of the changing course of the Orinoco River
(Schargel et al. 2007).
The two species of Hydrodynastes are large (approach or exceed three meters) and wide-
spread. Zaher et al. (2019) found Hydrodynastes to be the sister to the terrestrial and arbo-
real Oxyrhopus and Siphlophis. In addition, the highly aquatic Tretanorhinus was found to
be paraphyletic by Zaher et al. (2019). The genus also occurs in Cuba, and Central America.
Gomesophis the Brazilian endemic is the sister to Thamnodynastes (Zaher et al. 2019).
The 21 species in the genus Thamnodynastes are not monophyletic (Zaher et al. 2019) and it
has and has many semi-aquatic-arboreal species specialized to feed on frogs.
The Amazon River Basin covers 300,000 square kilometers, is 3,000 km long, and floods
5-15 meters annually. It is the world’s largest river, with a flow that results in about one-sixth
to one-fifth of all the world’s fresh water. It is so large that it creates its own weather patterns.

The Ancient Pebas Wetland

In what is now the Western Amazonia, an ancient wetland was likely the incubator for
aquatic snake evolution on the South American continent. Hoorn et al. (2010) reviewed the
characteristics of this mega-wetland and its phases of development. The region consisted
of fluvial systems that originated on the Amazonian Craton and were directed towards the
sub-Andean zone and the Caribbean. In the Early Miocene, the fluvial systems were primar-
ily replaced by lakes, swamps, tidal channels, and marginal marine embayments, forming a
mega-wetland.
At its maximum extent, the wetland covered over 1.5 million square kilometers compris-
ing much of the present western Amazonian lowlands. Descriptions of the snake fossils from
the Pebas system have not yet been published. Salas-Gismondi et al. (2015) report on the
crocodilians from two species-rich bonebeds from late Middle Miocene proto-Amazonian
deposits of northeastern Peru and some highly aquatic snakes throughout the Neogene.
In Amazonia, marine incursions are recorded as thin beds in the Middle to Upper Mio-
cene fluvial strata and contain evidence of marine and coastal taxa (foraminifera, mangrove
pollen).

The Pantanal
The Pantanal is the largest tropical wetland and the world’s most extensive flooded grass-
lands. The region is a vast alluvial floodplain in southwestern Brazil’s upper Paraguay River
basin. The subsidence that created the basin probably occurred about 2.5 Ma and was subject
to severe climatic changes during the Quaternary. It was a sandy desert during the last glacial
period 13,000 years ago. About 80% of the Pantanal floodplains are submerged during the
rainy seasons, supporting a diverse collection of aquatic plants and animals.
The gradual slope of the basin receives runoff from the upland Planalto highlands and
slowly releases the water through the Paraguay River and its tributaries. The concave for-
mation resulted from a pre-Andean depression of the earth’s crust, related to the Tertiary

55
Aquatic Snakes

Figure 3.4. The development and reduction of the Pebas Wetland. From Hoorn
and Wesselingh, 2010.

56
 3. Aquatic Snake Geography

Andean orogeny. It constitutes a huge inland delta, in which several rivers flowing from the
surrounding plateau merge, depositing their sediments and erosion residues.
The Pantanal is bounded by mostly arid habitats - the Chiquitano dry forests to the west
and northwest, the arid Chaco dry forests to the southwest, and the humid Chaco to the
south. The Cerrado savannas lie to the north, east, and southeast. Throughout the year, tem-
perature varies about 6.0 °C, with the warmest month being November (with an average tem-
perature of 26 °C) and the coldest month being June (with an average temperature of 20 °C).
Europe
Europe has few aquatic snake species, all in the natricid genus Natrix. However, Zaher
et al. (2019) found Natrix to be the sister to all New World natricids. Mediterranean Sea Del-
tas. The Rhone River Delta forms France’s most important wetland, the Camargue, covering
about 9,000 square kilometers. The dealt has a hot, dry summer and cool, wet winters. About
a third of the Camargue comprises lakes or marshland. Its brine ponds provide one of the few
European habitats for the greater flamingo.
Po River Delta – Venice. The Regional Park of the Po River Delta is in the provinces of
Ravenna and Ferrara, in the Emilia-Romagna region, was established in 1988 to safeguard

Figure 3.5. The wetlands and rivers of Europe.

the native flora and fauna of this area. It is a unique environment and was declared a human
heritage site by UNESCO in 1999.
Coastal Salt Marshes in northern Europe are extensive, and mudflats occur along the
Atlantic Ocean and Europe’s North Sea coastlines.
Northern boreal and subarctic peatlands compose more than half of the world’s wet-
lands. Unfortunately, much of this is at latitudes well above where most snakes can survive.
However, there are records of Natrix close to the Arctic Circle.
57
Aquatic Snakes

A collision of continental plates about 12 MYA raised new mountains in central Europe
and created the most extensive lake ever known, the Paratethys Sea. This inland sea’s fauna
included the smallest known whales. Palcu et al. (2021) estimate the lake once contained over
1.77 million cubic kilometers of water (more than ten times the volume found in all of today’s
fresh- and saltwater lakes combined). However, a changing climate caused the lake to shrink
at least four times dramatically in its five-million-year life. Water levels and salinity fluctua-
tion resulted in aquatic species passing into extinction. Species that could survive changing
salinities repopulated the lake when it expanded. The Paratethys ceased to exist 6.7 -6.9 Ma
when erosion formed an outlet.

Africa & Madagascar

Wetlands are abundant in sub-Saharan Africa, including the Congo River Swamps,
the Inner Niger Delta, the Sudd of the Upper Nile, and the Okavango Delta. Six snake
families contain aquatic species. The Pythonidae has two species of giant python that are
semi-aquatic (P. natalensis and P. sebae). A lineage within the elapid genus Naja has at least
four semi-aquatic species. The family Lamprophidae has aquatic snakes in two lineages Ly-
codonomorphus (about eight species) and one species of Boaedon. Africa has a radiation of
natricids composed of six genera, five of which are endemic Afronatrix, Limnophis, Helophis,
Hydraethiops, Natriciteres, and Natrix. The family Colubridae has one semi-aquatic genus
with several species in the endemic Crotaphopeltis. For many years snakes in the endemic
African genus Grayia were considered natricids. The four species are now placed in their own
family Grayidae.
Madagascar has two genera of semi-aquatic snakes that are poorly known, Liopholidophis
and Thamnosophis. Both are in the sub-family level clade Pseudoxyrhophiinae.
The Nile Delta is (or was) one of the largest river deltas, but much of the area is now
farmland and no longer floods due to the Aswan Dam. The Nile Crocodile and hippopot-
amus are two aquatic vertebrates once widespread in the delta during antiquity. They have
been replaced by 39 million people living on the delta.
The Congo River Basin is in west equatorial Africa and is sometimes known as the Con-
go. The drainage basin originates in the highlands of the East African Rift system with input
from many rivers. The young age (1.5 to 2 MYA) and active uplift of the East African Rift at
the headlands results in the river’s huge annual sediment load (Beadle 1981). The basin is 3.7
million square kilometers and has extensive, undisturbed tropical rainforests and significant
wetlands. The drainage basin empties into the Gulf of Guinea on the Atlantic Ocean. The cli-
mate is equatorial tropical, with two rainy seasons and high temperatures all year. The Congo
River is the second longest river in Africa, shorter only than the Nile, and it is the second
largest river in the world in discharge volume, following the Amazon. It is also the world’s
deepest river, with depths of over 220 m (Oberg et al. 2009).
The genus Crotaphopeltis reaches its highest diversity with many sympatric species in the
Congo River Basin. These snakes are considered primarily mesic savanna dwellers. However,
some, such as C. hotamboeia have been found in coastal fynbos (areas with sclerophyllous
vegetation) less than 200 m from the Indian Ocean in South Africa. Aquatic snake genera of
the Congo Basin include Afronatrix, Crotaphopeltis, Grayia, Helophis, Hydraethiops, Limno-
phis, Lycodonomorphus, Natriciteres, and Naja.
The Blue and White Nile confluence forms the Sudd in southern Sudan. The Sudd creat-
ed a mosaic of wetlands, and at one time, it was an impenetrable barrier to navigation on the
Nile. The shallow, flat inland delta covers an area of about 30,000 square kilometers but can

58
 3. Aquatic Snake Geography

Figure 3.6. The wetlands and rivers of Africa.

be as large as 130,000 km2 depending on the inflowing waters. The Sudd consists of mean-
dering channels, lagoons, reeds, and papyrus marshes. Aquatic snakes inhabiting the Sudd
include Natriciteres and Grayia.
The Okavango River empties into a basin in the Kalahari Desert and forms an inland
delta. The Okavango Delta is famous for its wildlife. However, despite the high diversity, the
delta supports few aquatic or semi-aquatic snake species. Six species of snakes representing
the genera Crotaphopeltis, Limnophis, and Natriciteres approach the northern edges of the
delta. The Okavango River system remains under studied in Zambia, central Angola, and
Botswana.
The Nile Delta covers about 25000 km2, but the outer edges have been eroded, and
59
Aquatic Snakes

some coastal lagoons have increasing salinity levels because of their connection to the Medi-
terranean Sea. Since the Delta no longer receives an annual supply of nutrients and sediments
from upstream due to the construction of the Aswan Dam, the soils of the floodplains have
lost the yearly input of nutrients. The Suez Canal is east of the Delta and enters the coastal
Lake Manzala northeast of the Delta. To the northwest are three other coastal lakes or la-
goons: Lake Burullus, Lake Idku, and Lake Mariout. The Delta has a hot desert climate, as
does the rest of Egypt. Still, its northern portion is the wettest region in the country and has
relatively moderate temperatures, with highs rarely surpassing 31°C in the summer. There
appear to be only one aquatic snake species in the Nile Delta, Natrix tessellata (El Din 2006).

Australasia
Semi-aquatic and aquatic snakes in Australasia include the genera Liasis (Pythonidae),
Acrochordus (Acrochoridae), Cerberus Djokoiskandarus, Pseudoferrania, Myron, Fordonia,
Heurnia (Homalopsidae), Tropidonophis (Natricidae), Hemiaspis, Laticauda, Aipysurus, Em-
ydocephalus,

n wetlands are distinctive due to seasonal dry periods with high evaporation rates and
low rainfall. Thus, saline wetlands and lakes are common due to the high evaporation rates.
Kakadu National Park is about 20,000 km2 in the Northern Territory of Australia. One
of Kakadu’s best-known landmarks is the Yellow Water billabong. A wetland near the small
settlement of Cooinda. The billabong, which floods and connects other waterways during
the wet season, attracts millions of migratory birds yearly. In addition, aquatic snakes in the
genera Liasis, Acrochordus, Cerberus, Fordonia, Myron, Pseudoferrania, and Tropidonophis
inhabit the area.
The Coorong, Alexandrina Lake, and Albert Lake in South Australia support a diverse
waterbird assemblage that visits the 1405 square kilometers of wetland. The area comprises
23 individual wetlands. Semi-aquatic snakes present includes Notechis and Pseudechis.
Barmah Forest, Victoria, is a 285-square-kilometer forest wetland that provides natural
flood control for the Murray River. The wetlands support 13 vegetation communities– in-
cluding the greatest extent of river red gum in Victoria – which provides vital habitats for
native bats, parrots, possums, and the Red-bellied Black Snake Pseudechis porphyriacus.
Macquarie Marshes, in northwestern NSW, have swamps, lagoons, and channels on
the floodplains of the Macquarie River. Semi-aquatic snakes present include Austrelaps and
Hemiaspis.
Mareeba Tropical Savanna and Wetland Reserve, Queensland, also known as the Ma-
reeba Wetlands, comprises 12 interconnecting lagoons, creeks, and channels across 20.25 ha
reserve.
Aquatic snake genera include Liasis, Pseudoferrania, Tropidonophis, and possibly Hemias-
pis. Many of Australia’s wetlands are coastal and essential to this work because they support a
significant diversity (about 38 species) of sea snakes and several homalopsids. Coral reefs in
Australian waters are also crucial to marine snake communities.
Ashmore Reef National Nature Reserve and Cartier Island Marine Reserve (the Re-
serves) are two discontinuous exposed open ocean platform reefs on the northwestern edge
of the Sahul Shelf in northwest Australia, in the Timor Sea. Ashmore Reef comprises an area
of approximately 583 km2 and includes three small, vegetated islands, several sand cays, two
lagoons, and an extensive reef. Cartier Island covers about 167 square kilometers and is about
25 nautical miles southeast of Ashmore Reef. Cartier Island is an unvegetated sand cay sur-

60
 3. Aquatic Snake Geography

Figure 3.7. Wetlands of Australiasia.

rounded by a coral reef with an extensive reef flat and no lagoon. The Reserve includes
the seabed and substrata to 1000 meters depth. The Reserves are protected for their high
biodiversity, including sea snakes. The Reserves are in an isolated oceanic location and thus
undisturbed; they have extraordinary cultural value. Declines in biodiversity are especially
troublesome when they occur in isolated locations relatively protected from most major
anthropogenic disturbances.
Lukoschek et al. (2013) examined declines in sea snakes at Ashmore Reef based on sur-
veys conducted between 1973 and 010 recorded the highest abundances (average of 42–46
snakes per day) and species richness (nine species) in 1973 and 1994. In 2002 abundance
had declined by over 50% (21 snakes per day representing five). Since 2005 abundances
have been consistently low (1–7 snakes per day), with just two species, Aipysurus laevis,
and Emydocephalus annulatus, recorded in significant numbers. Despite extensive searches
since 2005 (especially in 2010), five species of sea snake historically abundant at Ashmore
Reef have not been sighted and are presumed locally extinct. These species include three
Timor Sea endemics Aipysurus apraefrontalis, Aipysurus foliosquama, Aipysurus fuscus, and
one Australasian endemic Aipysurus duboisii. Declines in the abundance and diversity of
sea snakes at Ashmore Reef cannot be attributed to differences in survey methods over the

61
Aquatic Snakes

years.
In New Guinea, the Wasur National Park is a massive wetland region in the Indonesian
province of Papua. It was dubbed the Serengeti of Papua owing to its incredible biodiversity.
The area supports many waterfowl species and migrant birds. However, the invasion of alien
species such as water hyacinth and the Giant Sensitive Tree threatens the park’s naturally
flooded grassland systems.

Asia
Asia, the largest continent, has diverse climates and landscapes ranging from the arctic to
tropical, tundra to rainforest. Southeast Asia is wet, while western Asia is dry. The Himalayas
influence the monsoon by forcing a thermal low to form and draw moisture into the atmo-
sphere during the summer. The dynamic range of the climates and landscapes has created
numerous and often extensive wetlands where biodiversity can thrive and evolve.

China
The Pearl and Yangtze River Deltas have approximately 650,000 km2 of wetlands. Of
that, 250,000 km2 have been reserved, and 400,000 km2 are used for rice paddies and fish-
ponds. The great plains of the Yellow and Yangtze River basins in eastern China contain
some of the most extensive wetlands in the country and the greatest concentration of large
freshwater lakes. The total area of lakes is over 0.22 million square kilometers. The Yangtze
basin is noteworthy for its lakes, including the Dongting lakes in Hunan Province, the Wuhan
lakes in Hubei Province, Poyang Lake in Jiangxi Province, a chain of large lakes including
Shengjin in southwestern Anhui Province, Cao Hu in central Anhui Province, and Tai Hu,
Hongze Hu and Gaoyou Hu in Jiangsu Province. Many lakes are fringed with marshes. Ex-
tensive marshes occur in ancient lakes’ dried-out beds, old river channels (along the Yellow
River), and seasonally flooded areas (An et al. 2007). Aquatic and semi-aquatic snakes in-
clude: Acrochordus, Thermophis, Hypsiscopus, Myrrophis, ?Subsessor, Enhydris, Amphiesma,
Amphiesmoides, Atretium, Fowlea, Hebius, Herpetoreas, Natrix, Opisthotropis, Rhabdophis,
Rhabdops, Trimerodytes, and Xenochrophis.

South Asia
The South Asian (Fig.3.17) region includes Bangladesh, India, Nepal, Myanmar, Paki-
stan, and Sri Lanka. The region is rich in aquatic snake diversity Acrochordus, Python, Cer-
berus, Enhydris, Dieurostus, Ferania, Fordonia, Gerarda, Gyiophis, Homalopsis, Mintonophis,
Myanophis. Hydrophis, Amphiesma, Hebius, Fowlea, Rhabdophis, Xenochrophis are all repre-
sented.

Bangladesh
Much of Bangladesh is low-elevation terrain with many riparian systems. It has a 580
km coastline on the Bay of Bengal. The delta plain of the Ganges (Padma), Brahmaputra (Ja-
muna), and Meghna Rivers and their tributaries occupy 79% of the country. The Hill Ranges
make up only nine percent of the country. Thus, the country is characterized by a broad delta-
ic plain subject to frequent flooding and a small hilly region crossed by swiftly flowing rivers.
Roughly 80% of the landmass is the alluvial lowland. The Bangladesh Plain, with alti-

62
 3. Aquatic Snake Geography

tudes up to 105 meters above sea level, occurs in the plain’s northern part, but most are less
than 10 m above sea level. The south coastal region is at sea level. With such low elevations
and numerous rivers, water—and concomitant flooding—is a predominant physical feature.
About 10,000 square kilometers of Bangladesh are covered with water and are routinely
flooded during the monsoon season. The alluvial soils in the Bangladesh Plain are nutri-
ent-rich due to heavy silt deposits carried downstream during the rainy season. The rivers of
Bangladesh mark both the nation’s physiography and the people’s life. About 700 in number,
these rivers generally flow south. The rivers also drain excess monsoon rainfall into the Bay
of Bengal. Thus, the great river system is simultaneously
the country’s principal resource and its most significant hazard. The network of four
river systems flowing through the Bangladesh Plain drains 1.5 million square kilometers.
The numerous channels of the Padma-Meghna, its tributaries, and smaller parallel rivers
that flow into the Bay of Bengal are called the Mouths of the Ganges. During the annu-
al monsoon, the rivers of Bangladesh flow at about 140,000 cubic meters per second, but
during the dry season, the water flow is reduced to 7,000 cubic meters per second (Islam
2014).

India
India has about 14,500 km of inland navigable waterways with twelve significant rivers
and a total catchment area exceeding 2,528,000 square kilometers. All major rivers of India
originate from one of the three main watersheds: The Himalaya and the Karakoram ranges;
Vindhya and Satpura ranges in central India; and the Sahyadri or Western Ghats in west-

Figure 3.8. South Asian wetlands.

63
Aquatic Snakes

ern India. The Himalayan river networks are snow-fed and have a perennial water supply
throughout the year. The other two river systems depend on the monsoons and shrink into
creeks during the dry season. The Himalayan rivers that flow westward into Punjab are the
Indus, Jhelum, Chenab, Ravi, Beas, and Sutlej.
The Ganges-Brahmaputra-Meghana system has the largest catchment area of about
1,600,000 square kilometers. The Ganges Basin has a catchment of approximately 1,100,000
square kilometers. The Ganges originates from the Gangotri Glacier in Uttarakhand. It flows
southeast, draining into the Bay of Bengal. The Brahmaputra originates in Tibet, China,
and enters India in Arunachal Pradesh, then flows west through Assam. The Brahmaputra
merges with the Ganges in Bangladesh, where it is known as the Jamuna River. The Cham-
bal, another tributary of the Ganges, via the Yamuna, originates from the Vindhya-Satpura
watershed. The river flows eastward. Westward-flowing rivers from this watershed are the
Narmada and Tapi, which drain into the Arabian Sea in Gujarat.
The Western Ghats are the source of all Deccan rivers, including the Godavari River,
Krishna River and Kaveri River, draining into the Bay of Bengal. These rivers constitute 20%
of India’s total outflow. However, the heavy southwest monsoon rains cause the Brahmapu-
tra and other rivers to distend their banks, often flooding surrounding areas.
The Pichavaram Mangroves. Tamil Nadu India’s wetland ecosystem is widely distribut-
ed from the cold and arid in the Ladakh region of Jammu and Kashmir and those with the
wet and humid climate of peninsular India. Most wetlands are directly or indirectly linked to
river networks. Mangrove forests border the Indian coastline in sheltered estuaries, creeks,
backwaters, salt marshes, and mudflats. Mangroves cover 4,461 square kilometers or about
7% of the world’s mangrove forests. Prominent mangroves are in the Andaman and Nicobar
Islands, the Sundarbans delta, the Gulf of Kutch, and the deltas of the Mahanadi, Godavari,
and Krishna rivers. Parts of Maharashtra, Karnataka, and Kerala also have extensive man-
grove forests. The Sundarbans delta is home to the largest mangrove forest. It lies at the
mouth of the Ganges and spreads across areas of Bangladesh and West Bengal. The Sundar-

Figure 3.9. The mangrove dwelling homalopsid Gerarada prevostiana.

Photography by Avrajjal Ghosh.

64
 3. Aquatic Snake Geography

bans is a UNESCO World Heritage Site but is identified separately as the Sundarbans (Ban-
gladesh) and the Sundarbans National Park (India). A complex network of tidal waterways,
mudflats, and small islands of salt-tolerant mangrove forests intersects the Sundarbans. The
Rann of Kutch is a marshy region in northwestern Gujarat and the bordering Sindh province
of Pakistan. It occupies a total area of 27,900 square kilometers.
The Kerala Backwaters are a chain of lagoons and lakes lying parallel to the Arabian Sea
coast in the state of Kerala in southern India. The network includes five large lakes linked by
canals; some constructed, others natural, and the area is fed by 38 rivers (Bassi et al. 2014).
Myanmar
The Irrawaddy River is the country’s largest river originating from the N’mai and Mali
rivers; it flows relatively straight from north to south before emptying into the Andaman Sea
at the Irrawaddy delta. Its drainage basin covers about 404,200 square kilometers. The delta
is the lowest expanse of land in Myanmar; it spreads out from the limit of tidal influence at
Myan Aung to the Bay of Bengal and the Andaman Sea, about 290 km from the south end of
the Ayeyarwady River. The region consists of mangroves and freshwater swamp forests. The
Irrawaddy Delta holds Myanmar’s largest remaining area of mangrove forest. However, over
the past thirty years, an estimated 75-83 percent of mangroves have been lost.

Nepal
Nepal is about 800 kilometers along its Himalayan axis and 150 to 250 kilometers across.
Nepal can be divided into three belts on the south-to-north axis: Terai, Pahad, and Himal. It
is divided into three major river systems from east to west: the Koshi, the Gandaki/Narayani,
and the Karnali - all are the Ganges’ tributaries. The Ganges-Yarlung Zangbo/Brahmaputra
watershed coincides with the Nepal-Tibet border; however, several Ganges tributaries form
in Tibet. These tributaries have cut through the highest mountains in deep gorges. They flow
south through the Middle Hills, then join in a parallel configuration before crossing the Ma-
habharat Range and emerging onto the plains where they deposited alluvial fans exceeding
10,000 square kilometers (Jha 2006).

Sri Lanka
The Kirila Kelle wetlands are found a short distance inland from the southern Sri Lank-
an city of Matara. The wetlands are a part of the Nilwala River basin and comprise some 3.0
km2 of wetlands. Portuguese colonialists built canals to mitigate flooding of the Nilwala river
into Matara township and aid in the cropping of a further surrounding 24 square kilometers
of rice and other farming. The Kirila Kelle Wetlands are the focus of a sustainable Environ-
mental Conservation Community Development program. The program is being developed
through Ecotourism and community development through a provincial integrated develop-
ment strategy managed and implemented by the community for the community.
The Muthurajawela is a marsh complex in the southern region of the Negombo lagoon,
30 km north of Colombo. The Muthurajawela is 30.68 square kilometers and is the country’s
largest saline coastal peat bog. The marsh is notable for its unique and highly diverse ecosys-
tem and is listed as one of the 12 priority wetlands in Sri Lanka. “Muthurajawela” translates to
“Swamp of Royal Treasure” (Kotagama and Bambaradeniya, 2006).

65
Aquatic Snakes

Figure 3.10. The Crab-eating Snake Fordonia leucobalia is the only species that has a distribution
extending from peninsular India to Australia and New Guinea. Today it is considered one spe-
cies. That may change in the future. Photography by Harry Ward-Smith.

Pakistan
The Indus River is one of Asia’s longest rivers. The headwaters, fed by glacier meltwater
and rivers in the Himalayan, Karakoram, and Hindu Kush ranges, originate in the mountains
of the Tibetan Plateau near Lake Manasarovar. The river supports ecosystems with temperate
forests, plains, and arid landscapes.
The Indus runs through the Ladakh region of Jammu and Kashmir, India, towards the
Gilgit-Baltistan region of Pakistan and the Hindukush ranges. It then flows in a southerly
direction along the entire length of Pakistan to merge into the Arabian Sea near the port city
of Karachi in Sindh. The drainage area exceeds 1,165,000 square kilometers and is one of the
largest rivers in the world in terms of annual flow. The northern part of the Indus Valley, with
its tributaries, forms the Punjab region, while the lower course of the Indus is known as Sindh
and ends in a large delta (Kahn and Arshad 2014).

Southeast Asia
Southeast Asia has the most extraordinary diversity of aquatic snakes. Seven families
contain 44 genera. Aquatic and semi-aquatic snakes found in this region include the acro-
chordid genus Acrochordus; pythonid genera Liasis, Malayopython, Python; the xenopeltid
genus Xenopeltis; the cylindrophid genus Cylindrophis; the homalopsid genera: Bitia, Canto-
ria, Cerberus, Enhydris, Erpeton, Fordonia, Gerarda, Homalophis, Homalopsis, Hypsiscopus,
Karnsophis, Kualatahan, Miralia, Myrrophis, Phytolopsis, Raclitia, Subsessor, Sumatranus; the
elapid genera: Aipysurus, Hydrophis, Laticauda; the natricid genera Amphiesma, Amphies-
moides, Anoplohydrus, Atretium, Fowlea, Hebius, Herpetoreas, Hydrablabes, Isanophis, Opisth-

66
 3. Aquatic Snake Geography

Figure 311. Southeast Asia has many rivers and wetlands as well as a diverse aquatic snake fauna.
Historically the wetlands have expaded and contracted . It also has the most family level clades,
genera, and species of aquatic snakes than another region of the planet.

otropis, Pseudagkistrodon, Rhabdophis, Rhabdops, Smithophis, Trimerodytes, Tropidonophis,

Xenochrophis.
The Mekong begins at the Tibet Plateau and runs through China, Laos, Cambodia, and
Vietnam, draining 625,000 square kilometers. The Mekong Delta is in southern Vietnam.
Sundaland (Fig.3.22) is a biogeographical region of Southeastern Asia corresponding
to a landmass exposed during the last 2.6 Ma when sea levels were lower than today. It
includes the Malay Peninsula on the Asian mainland, the large islands of Borneo, Java,
Sumatra, and their small islands.
The area covered by Sundaland is about 1.8 million square kilometers. The land area
of Sundaland has expanded, and the contracted considerably during the past two million
years, and the exposed land area is approximately half of the maximum historic area.
Sundaland’s western and southern borders are marked by deep water and the Indian
Ocean. The eastern boundary of Sundaland is Wallace’s Line, the east boundary of Asia’s
terrestrial mammal fauna, and the western edge of the Indomalaya and Australasia biogeo-

67
Aquatic Snakes

Figure 3.12. The plate on the opposite page illustrates the expansion and contractions of the Sunda
Shelf wetlands. Map A shows the Sunda Shelf in the present, figure G shows the Sunda Shelf 21,000
YBP. The other maps show views of the region in between the present and 21,000 YBP. From Voris
(2000).

68
 3. Aquatic Snake Geography

Figure 3.13. The distribution of terrestrial, fossorial arboreal, and freshwater snakes is shown in
brown , the distribution of marine snakes is shown in blue.
graphic provinces. The islands east of Wallace’s Line are Wallacea, a separate biogeograph-
ical region considered part of Australasia. Wallace’s Line is a deep-water channel never
crossed by any land bridges (Bird et al. 2005).
Large portions of Sundaland were most recently exposed from approximately 110,000
to 12,000 YBP during the last glacial period. When the sea level was decreased by 30–40
meters or more, land bridges connected the islands of Borneo, Java, and Sumatra to the
Malay Peninsula and mainland Asia. Because the sea level has been 30 meters or lower
throughout much of the last 800,000 years, the isolation of Borneo, Java, and Sumatra as
islands have been uncommon throughout the Pleistocene. In the late Pliocene, the exposed
area of Sundaland was reduced compared to the present. At the Last Glacial Maximum, the

Figure 3.14. The homalopsid Cantoria violacea is an example of a snake that occurs in freshwater,
brackish water, and marine habitats, Photography by Harry Ward Smith.

sea level fell by approximately 120 meters, and the entire Sunda Shelf was exposed (Bird et
al. 2005).
The equator runs through central Sumatra and Borneo. Thus, Sundaland is in the
tropics. Consequently, rainfall is the primary determinant of regional variation. Most Sun-
daland receives over 2,000 millimeters of rain annually, and rainfall exceeds evapotrans-
piration throughout the year. Thus, there are no predictable dry seasons like elsewhere in

69
Aquatic Snakes

Southeast Asia.
In summary, aquatic snakes are found throughout the distribution of snakes (Figure
13.23), but they have evolved multiple times in multiple lineages. Some have recent origins,
while others are more ancient. In addition, they have adapted to numerous aquatic micro-
habitats that differ in water chemistry, temperature, substrate, flow rate, and prey availability.
However, Southeast Asia’s Sunda Shelf, the Amazon Basin, the Pantanal, the Congo Basin,
North America’s Gulf Coastal Plain, and the coastal waters of Australasia stand out in sup-
porting the rich diversity of aquatic species.

Figure 3.15. The Banded Water Cobra Naja annulata.

Photography by Vaclav Gvozdik

On the next page. A Green Anaconda surfaces in a Venezuela River. Eu-

nectes murinus is the largest snake and inhabits the rivers of the Amazon
and Orinoc basins of South America. Photography by William W. La-
mar.

70
Aquatic Snakes

71
Aquatic Snakes

4. Ancient but Extant Lineages

The Henophidia was established by Hoffstetter (1939) for a group of snakes that in-
cluded boas, pythons, and “anilioids.” Various studies added or subtracted groups of snakes
to this superfamily in the 21st century. However, the Henophidia is no longer considered
monophyletic (Reynolds et al., 2014). These families may be best thought of as a collection
of ancient lineages that have survived to the present day.

Eyelash Boas – Tropidophiidae

The family contains two genera of ancient Neotropical snakes, Trachyboa and Tropi-
dophis. Both have been considered booids but form an independent clade that diverged rel-
atively early in snake radiation (Wilcox et al. 2002). The left lung is absent, and the tracheal
lung is well-developed. Males have cloacal spurs, and pelvic elements are present. Tropi-
dophis are terrestrial-arboreal snakes that are sometimes called wood snakes or tropes. How-
ever, there is evidence that the two species of Trachyboa are semi-aquatic.
Trachyboa contains two poorly known species: Trachyboa boulengeri inhabits the Choco
forests of Panama, Colombia, and Ecuador, and Trachyboa gularis occurs in the dryer coastal
areas of Ecuador. Both species are terrestrial, sedentary snakes with the peculiar defensive
behavior of coiling into a flattened disk or forming a rigid stick-like posture when disturbed
(Arnett et al. 1992). Much of what we know about these snakes come from captive animals.
Successful captive maintenance requires cool ambient temperatures. Both species prey on
frogs and toads but thrive on live fish in captivity. Litters
average 5–6 young.
They are unusual in appearance because of their very
rugose scales and the projecting scales from above the eye.
Both species will eat anurans but prefer fish; they seem to
do well at temperatures between 18−26C. At temperatures
Figure 4.2. Trachyboa boulengeri. above 21C, they stop feeding (Arnett et al. 1992). Trachy-
From Peracca 1910. boa have a hypapophysis (small projections) coming from
the ventral side of each vertebra; these are frequently found
in aquatic snakes.

Northern Eyelash Boa

Trachyboa boulengeri Peracca, 1910
Distribution and Habitat. Streams with gravel substrates and swamps in Panama, Ecua-
dor, and Colombia (Valle del Cauca) (Figure 4.1) (Ortega-Andrade et al. 2010). Type locality:
none given.
Natural History. The literature on this species’ food
habits is mainly based on captivity specimens. The diet is
composed of frogs and tadpoles (Pristimantis, Rhinella, and
Gastrotheca), as well as fish Dwyer et al. (2018) observed
a sub-adult in a temporary pond along a small stream in
Figure 4.1. The distribution of
Esmeraldas Province, Ecuador. The snake had its head and
Trachyboa boulengeri
about four centimeters of its body underwater. A catfish

72
 4. Ancient but Extant Lineages

Figure 4.2. A Trachyboa boulengeri from Esmeralda, Ecuador.

Photography by Ross J. Maynard.

(family Astroblebidae) in the same pool was coaxed to swim close to the snake. The snake
immediately seized and constricted the fish. After 30 minutes, the snake released the fish
without having killed it. Thus, coiling is for control rather than killing it. At the same locality,
at 2130 h, a subadult T. boulengeri in damp leaf litter next to a permanent pond in the prima-
ry forest was feeding on a juvenile leptodactylid frog, Leptodactylus melanonotus. The authors
report observing 23 individuals of T. boulengeri, of which eight were displaying “fishing” be-
havior. Shawn Harrison (personal communication) found these snakes in a similar situation
- submerged in a small stream with gravel and rock substrate (Figure 4.3).

Figure 4.3. The microhabitat of Trachyboa boulengeri. The snake in the insert was collected in the
stream shown in the larger photograph. Photography by Sean Harrison.

73
Aquatic Snakes

Southern Eyelash Boa

Trachyboa gularis Peters, 1860
Distribution and Habitat. Dry, coastal Ecuador. Type lo-
cality: Guayaquil, Ecuador (Figure 4.4).
Natural History. A poorly known species. Its habits are
assumed to be similar to Trachyboa boulengeri. The species is
probably extinct due to the the descruction of its habitat. The
savanna habitat for this species is "almost entirely gone." It has
become cattle pasture and banana plantations. All known lo-
Figure 4.4 The distribution of Tra-
chyboa gularis calities are in the vicinity of human settlements (Arteaga-Na-
varro and Cisneros-Heredia 2021).

Neotropical Pipe Snakes

The family Aniliidae is often considered monotypic with a single species, Anilius scytale.
However, a second species, Anilius phelpsorum Roze, is likely valid. The Neotropical Pipe
Snakes have a small gape and form the sister to the large gape wood snakes of the family
Tropidophiidae. As unlikely as this may seem, the relationship has been demonstrated in
multiple molecular investigations, given the morphological and ecological differences. They
form the Amerophida clade composed, and members share constriction to subdue prey and
viviparity. The molecular work suggests the two families shared an ancestor in the Cretaceous
about 89.1 Ma (Burbrink et al. 2020). Like many other ancient snake clades, the species in this
family have a remnant pelvic girdle and hind limbs.
The South American Pipe snakes are found in the tropics of northern South America
from southern and eastern Venezuela, Guyana, Suriname, and French Guiana south through
the Amazon Basin of Colombia, Ecuador, Peru, Bolivia, and Brazil. Despite the fact it has
been called the “Trinidad Pipe Snake,” its presence in Trinidad is problematic.
These snakes may be active at any time of day but are often found at night. It uses forests
and open areas and is usually associated with aquatic situations. Prey handling, described by
Marques and Sazima (2008), included seizing and coiling around small, elongated fish. Want-
ing to see how the snake would deal with larger prey, they fed the snake an amphisbaenian.
The lizard was bitten on the anterior body; the snake held its grip for 25 seconds. Horizontal
coils contacted the lizard’s body as the prey twisted around its long axis in a defensive move.
The snake loosened its hold and constricted the prey again. The pipe snake continued to bite
and constrict the lizard for about 30 minutes until the lizard died. The snake released its bite
hold and coils, swallowing the lizard headfirst.
Martins and Oliveira (1998) reported the fish Synbranchus marmoratus, the lizard Am-
phisbaena vanzolinii, and the snake Atractus torquatus in the diet of this species. The diet of
Anilius in the Brazilian Amazonia basin is based on the analysis of 162 specimens, including
Amphisbaenians (81.25%), Tantilla melanocephala (12.5%), and caecilians (6.25%). Anilius
scytale forages at night on the ground and in aquatic environments. Swallowing prey head-
first may minimize the risk of injury to the predator
(Maschio et al. 2010). It seems likely that this species is fossorial and may be semiaquat-
ic. The pipe snakes reach a length of about one meter. The body is cylindrical, hence the name
pipe snake. The tail is short, flatted, and raised in a defense display to draw attention away
from the head. Ventral (208-254) and subcaudal (10-15) scales are relatively small - about
twice the size of dorsal scales. The body is bright red with black transverse bars on the dorsal

74
 4. Ancient but Extant Lineages

and ventral surfaces.

Neotropical Pipe Snake

Anilius scytale (Linnaeus, 1758)
Distribution and Habitat. Forest leaf litter, swamps, and
streams in Amazonian Brazil, Venezuela? Trinidad?, Colombia,
Bolivia, Ecuador, French Guiana, Guyana, and Peru. In Venezu-
ela is known to reach 700 m in elevation. Type locality: “Indiis.”
Figure 4.5. The distribution Natural History. The diet consists of elongated amphibians
of Anilius scytale. and reptiles. Ferreira et al. (2020) report litters of 4-15 young 150-
180 mm in total length.

Orinoco Pipe Snake

Anilius phelpsorum Roze, 1958
Distribution and Habitat. Uncertain, likely restricted to the Orinoco Basin, including the
Delta Amacuro and maybe Trinidad. Type locality: “La base del Auyantepui, Estado Bolívar,
Venezuela. This species has been neglected and little is known about it. Many authors have
synonomized it with Anilius scytale

Figure 4.6. An Anilius scytale from the Napo region of Ecuador.

Photography by JCM

75
Aquatic Snakes

Sunbeam Snakes
Two species compose the Asian Sunbeam Snake family Xenopeltidae. Xenopeltis unicolor
is widespread, ranging from India to the Philippines. The other X. hainanensis is restricted to
the Chinese island of Hainan. Most authors (e.g., Orlov et al. 2000) regard these snakes as fos-
sorial. However, they are also semi-aquatic and are sometimes collected in gill nets by fishers
(Karns et al. 2020); Stuart (2004) found Xenopeltis unicolor to be the most traded snake at U
Minh Thuong National Park in southern Viet Nam. Asian Sunbeam snakes slightly exceed a
meter in length and feed on other snakes and many other vertebrates. Gilbert and Goodyear
(2019) report an adult X. unicolor preying on an adult Asian Bullfrog, Kaloula pulchra, in
Bangkok, Thailand. The extent to which these snakes are aquatic needs further investigation.
The Mexican Burrowing Boa family Loxocemidae is monotypic. Loxocemis bicolor is
considered a member of the Constrictores clade resurrected by Georgalis et al. (2020) and oc-
curs in Mexico and Central America. Mo-
lecular studies suggest Loxocemus is related
to the Asian sunbeam snake. However, it is
not known to be aquatic.

Sunbeam Snake
Xenopeltis unicolor Reinwardt, 1827
Distribution and Habitat. South and
Southeast Asia from India to the Philippines
and throughout the Indo-Chinese Peninsu-
la and Indonesia. Type locality: Java. While
this snake is terrestrial and inhabits lowland
and lower montane forest, scrub, and dis-
turbed habitats, it prefers wet ground and
burrows and enters bodies of water (Figure
4.7).
Natural
History. A
n o c tu r n a l ,
fossorial, and
semi-aquat-
ic snake
Figure 4.7. The distribution of typically en-
Xenopeltis unicolor. c o u n t e r ed
on land near Figure 4.8. The Sunbeam Snake, Xenopeltis uni-
water. It feeds on various prey, including color. (A-B) photographs were taken in Borneo,
frogs, snakes, lizards, and small mammals. and Thailand. Photography for A and B by Harry
Its detention is specialized for grasping Ward-Smith (C) a juvenile feeding on an Enhydris
and swallowing hard-bodied prey, such as enhydris in Thailand. Photography by JCM
skinks. Larger prey is constricted smaller
prey is eaten alive. Male combat has been re-
ported on a roadside verge consisting mainly of tall grass. The snakes displayed body-bridg-
ing behavior, twisting continuously while moving forward. After approximately 10 minutes,
they coiled around each other and bit each other for about one minute. Then, they continued

76
 4. Ancient but Extant Lineages

body bridging behavior and, once aligned, simultaneously pushed their anterior downwards
onto each other. Clutches of up to 17 eggs have been reported. Of 26 females, five ranging
from 653 to 1020 mm in body length contained eggs from 8 to 20 mm. The smallest gravid
female was 653 mm, was caught in January, and had eggs of 12 mm. The individual with the
largest ovarian eggs was seen at the end of July.

The Pythons
There are no highly specialized aquatic
pythons that converge with the anacondas –
the highly aquatic boas. However, some py-
thon species are best considered semi-aquat-
ic. The largest pythons (Malayopython
reticulatus, Python sebae, Python natalensis,
Python bivittatus) use aquatic situations to
hunt and thermoregulate (Murphy and Hen-
derson 1997, Walters et al. 2016, Murphy and
Crutchfield 2019). The largest pythons are
habitat generalists, but they are also frequent-
ly associated with water. The large body size
may be possible because of their buoyance
in water makingmoving more efficient. The
four species of short-tailed pythons (Python
curtus Group) are also frequently associated
with water, and the question of whether they
are semi-aquatic is controversial even though
they are often associated with streams, stream Figure 4.9. Brongersma's Short-tailed Python,
Python brongersmai, from Kaeng Krachan Na-
banks,and poorly drained soils, some have
tional Park, Thailand. (A) an adult, (B) an adult
specifically stated that they are not aquatic, in water. Photography by Rushen Bilgin
but see Figure 4.9b.

Australian Water Python,

Liasis fuscus, Peters 1873
Distribution and Habitat. Liasis fuscus is distributed acrosstropical north Australia, from
Broome in Western Australia to thecoast of central Queensland in the east. It also occurs in
southernNew Guinea, from Merauke, Irian Jaya eastward through the lowlands of Western Province,
Papua. Type locality: Port Bowen (=Port Clinton), Queensland, Australia (Figure 4.10).
Natural History. It is associated with seasonal bodies of water (floodplains, billabongs)
as well as permanent water (lakes and rivers).Northern Australia is highly season with a very
pronouncedwet season that expands water python’s habitat, and the aquatic habitats are very
productive resulting in high population densities, like Fogg Dam Northern Territory At this
locality fuscus reaches one of the highest concentrations of any vertebrate predators.
The Australian Water Python, Liasis fuscus, hunts in and out of the water and is probably
the most aquatic species in the clade. Its relative, the Olive Python, Liasis olivaceus, also uses
streams and associated flood plains. However, it does wander far from water and frequents
rock outcrops (Auliya 2006, Madsen et al. 2006, Walters et al. 2016, Ujvari et al. 2918).

77
Aquatic Snakes

Figure 4.11. Australian Water Python, Liasis fuscus. From the Tully area.
Photography by Scott Eipper.
The Australian Water Python can maintain a high body
temperature throughout the year without showing visible
basking behavior. However, cool nights during the dry season
depress snake activity (Shine and Madsen 1995)
A five-year field study by Shine and Madsen (1997) fo-
cused on ways the Dusky Rat’s (Rattus colletti) abundance
can influence the reproductive rates of its predator, the Wa-
Figure 4.10. The distribuition of ter Pythons (Liasis fuscus), on the Adelaide River floodplain
Liasis fuscus. in tropical Australia. The python preys almost exclusively on
the Dusky Rat. Rat numbers varied dramatically over the five
years of the study. Feeding rates of pythons were highest when rats were abundant and the
snakes were in good condition; that is, they had a good mass relative to body length. The
proportion of adult female pythons reproduced each year was closely tied to rat abundance.
However, reproductive output per litter, that is, offspring size and fecundity relative to mater-
nal body size, was unaffected by prey availability. Female pythons reproducing in years with
abundant rats were in better condition after oviposition.
Water Pythons use facultative maternal brooding with shivering thermogenesis to warm
their eggs. Shine et al. (1997) found that both nest-site selection and maternal care influence
incubation temperatures. They experimentally simulated three thermal regimes typical of dif-
ferent natural nests in their study population. These were (1) hot, stable temperatures typical
of nests laid in the burrows of varanid lizards -a constant 32°C; (2) lower and more variable
temperatures typical of nests laid inside tree root boles, either with maternal attendance, diel
range 27.1°–32.9°C; or (3) in root boles but without the maternal presence (24.3°–32.9°C).
The authors incubated 187 eggs from 15 clutches obtained from field-caught gravid pythons.
The temperature treatment during incubation strongly affected incubation periods, body siz-
es, body shapes (mass and tail length relative to snout-vent length), initial growth rates, escape
behavior, and willingness to feed. In addition, the authors detected strong maternal effects

78
 4. Ancient but Extant Lineages

on all these traits and interactions between maternal outcomes and incubation regimes.
Thus, a female python’s decisions about where she lays her eggs and whether she remains
and broods them through development have sig-
nificant consequences for the phenotypes of her
offspring.

Olive Python
Liasis olivaceus Gray, 1842
Distribution and Habitat. Endemic to north-
ern Australia, it is polytypic. Two subspecies are
recognized: Liasis o. olivaceus in Queensland and Figure 4.12. The distribution of the Ol-
ive Python, Liasis olivaceus.
the Northern Territory and Liasis o. barroni Smith,
1981 in Western Australia (Type locality Pin-
drunna, 32 km north-west of Tambrey, WA. The
Western Australian subspecies is the second-lon-
gest snake in Australia and is likely a valid species.
Habitat includes rocky terrain, including gorges
and rocky areas near water. It shelters in caves and
rock crevices but uses hollow logs and burrows un-
der rocks. Type locality: Port Essington, Northern
Territory (L. o. olivacea), Australia (Figure 14.12).
Natural History. Prey includes most verte-
brates. It hunts from ambush and actively forages
in the water in search of prey. In waterholes, it will
submerge and capture prey while concealed by the
water. Mating occurs from May to mid-July. Eggs
require a gestation period of 81–85 days, and the
female lays 12–40 eggs in late spring. The average
clutch size is about 19 eggs. Incubation is about
50 days. Hatchlings are approximately 35 cm in
length. (Smith 1981).

Reticulated Python Figure 4.13. The Olive Python, Liasis oli-

vaceus from Sybella Creek. Photography
Malayopython reticulatus Schneider, 1801 by Scott Eipper.
Distribution and Habitat. The species is native
to Southeast Asia and occurs primarily in the Indonesian Archipelago. It has been reported
from northeast India, but its presence is poorly documented. Its distribution includes many
islands reinforcing its aquatic behavior. It uses both primary forests, secondary forests, and
human-modified environments. Forest swamp habitats are used, as well as riparian en-
vironments that transect more open agricultural landscapes (Auliya 2003). Type locality:
None given by Schneider (Fig. 4.14).
Three subspecies are recognized. Malayopython reticulatus saputrai: Selayar Island, In-
donesia. Malayopython reticulatus jampeanus, and Malayopython r. reticulatus. The first
two are from very restricted ranges in the eastern Indonesian Archipelago.
Natural History. This is likely the longest extant species of snake. Malayopython is high-
ly adaptable, occupying a wide range of habitats from lower montane forests (up to eleva-

79
Aquatic Snakes

tions of at least 1500 meters), agricultural areas, scrubland, man-

groves, and cities. Prey is generally small to medium mammals
(small deer, wild pigs, rats, cats, dogs, goats, and other domestic
species). Prey is killed by constricting. And there are well-docu-
mented cases of large specimens killing and consuming humans. Figure 4.14. The distri-
It is an excellent swimmer and has been reported at sea far bution of the Reticulated
from land. It has also colonized many small islands. For example, Python, Malayopython re-
it was the first snake species to reach the island of Krakatoa after ticulatus.
its eruption.

Figure 4.15. The Reticulated Python, Malayopython reticulatus.

Photography by Kenneth Chin.

Reproduction. Much of what we know about Malayopython reproduction in nature

comes from the Sumatra population (Shine et al. 1999). Testes are enlarged in October com-
pared to April. Males engage in combat over females (Murphy and Crutchfield, 2018). Fe-
males lay their eggs from March through May. According to Ross and Marzec (1990) and
Malkmus et al. (2002), egg-laying occurs from February to March in Sabah, East Malaysia.
Auylia’s (2006) reproduction schedule for West Kalimantan reticulatus seems to be correlated
with wet and dry seasons. Mating occurs from October to December. Egg laying starts at the
end of the wet season and lasts from January to March-April, so the eggs are incubated during
the dry season. Hatchlings are present at the beginning of the wet season in August-October.
Various authors estimate the incubation time at 55–105 days, suggesting that females in some
populations may retain eggs during development.
Females in the Sumatra population may not reproduce every year and may hold for other
localities. Reproduction depends on the available food supply and the hunting success of in-
dividual females. Clutch sizes reported in the literature range from 8–124, with many clutches
in the 20–40 range. Maturation may occur in two-year-old males and three-year-old females
that are captives. Age to maturity in the wild is undoubtedly longer. Cox (1991) suggests five
years.

80
 4. Ancient but Extant Lineages

Burmese Python
Python bivittatus Kuhl, 1820
Distribution and Habitat. Eastern Nepal, India (Arunachal
Pradesh, S Bhutan, Bangladesh, Myanmar, Thailand, Laos,
Cambodia, Vietnam, S China including Hainan and Hong
Kong; Sichuan, Guangxi, Guangdong; Indonesia (Java, Bali,
Sulawesi). Type Locality Java (Mertens 1930). Introduced into
southern Florida (Figure 4.16).
Figure 4.16. The distribution Natural History. Prey taken by Python bivittatus is diverse.
of Python bivittatus. Stomach contents within its natural distribution are almost
always terrestrial or semi-aquatic vertebrates. They include
frogs, monitor lizards, birds,
bats, rodents, pangolins,
porcupines, langurs, jack-
als, mousedeer, hog deer,
chital, barking deer, pigs,
chinkara, civets, and leop-
ards.
Reproduction occurs
in the early spring, with fe-
males producing 12–36 eggs
in March or April. Females
attend to the eggs, usually
until they hatch, wrapping
around them and rapidly
contracting their muscles
to raise the temperature of
the eggs by several degrees.
However, their muscle
twitch and metabolic rate
were negatively correlated
with nest temperature af-
ter hatchling maternal care
ceased.
Hatchlings are 480 to
790 mm and weigh 75 to
165 g. Sexual maturity is
probably reached at about
Figure 4.17. The Burmese Python, Python bivittatus.
2 m in males and 2.6 m in
Photography by Artur Tomaszek.
females, although maturity
at smaller sizes has been reported. It seems likely that reaching maturity in less than two or
three years is an artifact of captivity. Willson et al. (2014) said a female, 2.1 m in total length
with a 1.85 m SVL with 11 viable eggs in the Florida population.
Molecular evidence for facultative parthenogenesis in this species was provided by Groot
et al. (2003). In addition, a female Python bivittatus from the Artis Royal Zoo in Amsterdam
produced eggs for five consecutive years that contained embryos while she was isolated from

81
Aquatic Snakes

males.
The aquatic nature of bivittatus may be best demonstrated by its successful colonization
of the Florida Everglades, a riparian environment dominated by grass and canals. Smith et al.
(2021) found this python uses aquatic habitats within its native range. Movements were pos-
itively associated with aquatic habitat features such as water bodies (i.e., ponds, irrigation ca-
nals) and agriculture (i.e., rice paddy), suggesting that aquatic elements are essential in habi-
tat selection. Smith et al. (2021) suspect that the Burmese Python’s use of connected systems
of aquatic agriculture in their native range may help explain their success in the Everglade
wetlands of Florida. Rice paddies are often wetland-like habitats and support diverse wading
bird communities. In addition, the vegetation that typically grows at the edges of ponds and
irrigation canals may serve as refugia in areas highly modified by humans.
The Burmese Python has been reported swimming in the ocean. Li-Wei Chung et al.
(2016) observed this snake swimming between two islands near the China coastline. It was
about 2 km from its origin of, Lesser Kinmen Island, and about 1 km from its destination
(Major Kinmen Island).

Indian Python
Python molurus (Linnaeus, 1758)
Distribution and Habitat. The Indian Python tends to
follow rivers into dry and upper-elevation habitats that it
would otherwise not be able to colonize. They are present
from Pakistan’s Indus Valley eastward into India, Nepal,
Bhutan, and western Bangladesh, then southward into pen-
insular India (O’Shea 1998; Barker and Barker 2008). Schle-
Figure 4.18. The distribution ich and Kastle (2002) suggest that P. molurus and P. bivittatus
of the Indian Python, Python may be sympatric in the Terai of Nepal. Shaw et al. (1942),
molurus. writing about the snakes of Sikkim and Bengal, recognized
a light and dark phase of this snake but suggested the dark
phase was from northeast India and the lighter phase from
southwest India. They considered it common up to about 1,219 m and noted that Wall re-
ported it from 1,828 m. Pope (1935, 1961) says it from 1,500 m and 1,650 m, and it has been
reported from 2,000 to 2,500 m (Whitaker 1978; Ernst and Zug 1996; Kabish in Schleich and
Kastle 2002; Whitaker and Captain 2004). Type locality: Java (Mertens 1930) (Figure 4.18).
The Java holotype may be based on a feral specimen or population.
Its northern and eastern borders are poorly understood. In part, this is due to the
long-standing confusion with bivittatus. It is possible, even probable, that the current distri-
bution of this species represents a combination of its natural distribution and human intro-
ductions. People have been moving this snake around for centuries. See Stothers (2004) for
evidence that humans transported it in ancient times. Vyas (2011, 2013) describes moving
rescued specimens of molurus from urban areas to agricultural lands and forest reserves.
Natural History. A habitat generalist uses open rocky habitats, deserts, cultivated lands,
grasslands, marshes, scrub forests, mangrove forests, deciduous forests, wet tropical forests,
and urbanized areas. The variety of habitats and its ability to survive in agricultural and ur-
banized lands suggests this is a highly adaptable snake. Despite its plasticity, P. molurus is the
only large constricting snake considered threatened. This is probably justified since much
of its distribution coincides with a human population that exceeds one billion humans. The
snake could be easily extirpated through overhunting, habitat destruction, and pollution. In

82
 4. Ancient but Extant Lineages

Figure 4.19. Python molurus. Photography by JCM.

India, it occurs throughout the country, avoiding the extreme elevations and xeric desert
areas. Perhaps the most prominent and best-studied population of P. molurus can be found
in Keoladeo National Park, near Agra in Rajasthan. Whitaker (1993) suggests that the popu-
lation consisted of 100 adults. Keoladeo is a temporary monsoonal wetland and remains dry
from April–July. Colder months (December–February) corresponded to the mating season
(Bhupathy and Ramesh 2010) when snakes were found near burrows and basking. Pythons
were active during the day (0600 to 1800 h) in winter and summer. Winter activity was diur-
nal and was throughout the day in the spring, becoming crepuscular in summer. The burrow
temperatures correlated with the ambient temperature during the day, but nighttime bur-
row temperatures were warmer than the ambient temperature. Prey activity peaked in the
morning and evening hours throughout the study. The decline in sightings towards the sum-
mer may be due to the emigration of pythons or a decrease in conspicuous basking activity
(Krishnan et al. 2009).

South African Python

Python natalensis Smith, 1840
Distribution and Habitat. It ranges from the equator in
Kenya and the Central Congo DRC southward through Angola,
Zambia, Tanzania, Malawi, Mozambique, Zimbabwe, Botswana,
Namibia, and South Africa. Type locality: Port Natal = Durban,
Natal, South Africa. Broadley (1984) suggests natalensis favors
moist, rocky, well-watered wooded valleys, plantations, and
Figure 4.20. The distribution of
Python natalensis.
bush country and observed that they are rarely far from water.
In Kruger National Park, Piennar (1966) describes its habitat as
moist, rocky, well-wooded valleys, reed beds, and bush country
83
Aquatic Snakes

Figure 4.21. Southern African Rock Python, Python natalensis, at the edge of the Cuando River in Bo-
tswana, Kwando Lagoon Camp, north of Moremi Wildlife Reserve and to the west of Chobe National
Park. Photography by Martha de Jong-Lantink.

but notes that it seldom ventures far from permanent water (Figure 4.20).
Natural History. Hatchling snakes feed on mice, rats, birds, lizards, and frogs. Adult
South African Pythons sometimes hunt from ambush, including from a submerged posi-
tion. Fitzsimons (1930) describes a python striking and overpowering a duiker buck when it
stepped into a shallow pool. He also reports this species lying along a tree branch waiting for
prey. Thomas (1985) says a natalensis captured a dove from an arboreal ambush as the bird
flew into a water hole to get a drink.
Males may follow receptive females for as long as two months—Python natalensis mates
in the autumn. The number of eggs laid is correlated to the female body size and clutches of
23–69 eggs (Alexander 2018). The eggs are 142–156 g, and no visible embryo is present when
the eggs are laid. Females lay their eggs in anteater or aardvark burrows, cavities in rocks,
cavities among tree roots, and accumulations of vegetation such as leaf litter, grass clumps,
and sugar cane. Eggs are deposited in December and January, but egg laying in October and
November did occur. The female coils around the eggs and produces body heat (thermogen-
esis) to raise the temperature of the developing embryos to about 38–40°C. Captive snakes
were less likely to attend to their eggs. Additionally, the captive females paid little attention to
the young after hatching.
Monitor lizards, mongooses, rats, and jackals are egg predators, and their presence may
encourage the female to stay with their eggs. Incubation is about 60 days. Hatchlings are 450–
600 mm long. Female Python natalensis do not eat while attending eggs and may lose 40% of

84
 4. Ancient but Extant Lineages

their body mass during the attending period. Alexander (2018) saw that females continue
to care for neonates after the eggs hatch. Female pythons would transmit heat to the eggs
but also to the hatchlings. The female’s skin turns dark, almost black, when basking outside
the burrow. Females may only reproduce once every two or three years.

North African Python

Python sebae Gmelin in Linnaeus, 1789
Distribution and Habitat. It inhabits the dense semi-ev-
ergreen and semi-deciduous forest and more open savanna
environments, marshland, and gallery forest. Often it occurs
close to water. It has been found in swamps, trees, and shrub
savanna in the Gambia (Starin and Burghardt 1992). It oc-
curs in the lower Tana River basin, at low elevations in the
Figure 4.22. The distribution
Lake Victoria Basin, and in the coastal lowlands of Kenya
of Python sebae.
(Broadley 1990). In southeastern Nigeria, Luiselli and col-
leagues (2001, 2007) report that it uses a vari-
ety of habitats, noting that it is most abundant
in coastal mangroves and spends considerable
time in the water. In Somalia, it has been found
in xerophilic open woodland, lowland ever-
green thicket and scrub, lowland dry evergreen
forest, lowland moist evergreen forest, and ri-
parian formations. Pakenham (1983) discussed
a possible waif in the Pemba Islands off the
coast of Tanzania, confirming its tolerance to
salt water (Figure 4.22).
Ranges south of the Sahara from Senegal
eastward to Ethiopia, extending southward
into northern Angola as far south as Ambriz
on the coast, the Shaba Province of Zaire, Ken-
ya, and Tanzania. Type locality: Guaira dans le
Bresil (in error). Guaira may refer to Guayrá in
Figure 4.23. The North African Python, Py- present-day Paraguay and under the control of
thon sebae.. Photography by JCM Spanish conquistadors in the 1550s. Or it may
also refer to a locality in the former Dutch Bra-
zil. The specimen was most likely collected in West Africa on a return trip from the New
World. The type locality is thus in West Africa (Åhlander in Uetz 2020). Rarely does it go
above 2,250 m in elevation?
Natural History. The diet in natural and urbanized habitats in southern Nigeria was
studied by Luiselli et al. (2001). They found it fed mostly between 4:00 PM and 12:00 AM
in the suburban areas, whereas, in the natural habitat, it feeds mainly during the day (7:00
AM to 7:00 PM). The urbanized population preys primarily on human commensals such as
poultry, rats, goats, and dogs. It fed on rodents, fruit bats, and crocodiles and monitor liz-
ards in more natural habitats. Predation in suburban areas often occurs near rivers because
the snakes hide in riparian vegetation and riverbanks and uses the waterways as dispersion
routes through heavily populated areas. Comparing the body size of snakes in natural hab-

85
Aquatic Snakes

itats versus urbanized areas, they found smaller body sizes, averaging 2.9 m (1–4.5 m), than
those from forested regions, which averaged 3.4 m (1–5.5 m). Python predation on large prey
(dogs) was unusual in urban areas, possibly because of the relatively smaller body size. The
hatchlings and young pythons fed on the giant rats of the genus Cricetomys, among the largest
West African rodents. Adults take much larger prey, including antelope that may weigh more
than the snake.
Hatchlings captured in the wild and raised at the London Zoo included a female that
grew to 8 or 9 feet [2.4–2.7 m] in five years and mated when she was five years and ten months
old (Flower, 1925). Lederer (1956) had two females, one 11 feet (3.35 m) and another 11 feet
9 inches (3.58 m), that laid eggs; both were five years of age.
Size. Hatchlings are 570-710 mm. Hatchlings captured in the wild and raised at the Lon-
don Zoo included a female that grew to 8 or 9 feet [2.4–2.7 m] in five years and mated when
she was five years and ten months old (Flower, 1925). Lederer (1956) had two females, one 11
feet (3.35 m) and another 11 feet 9 inches (3.58 m), that laid eggs at five years of age.

Aquatic Boas
Booid taxonomy and systematics have changed radically over the past several decades,
while knowledge of the natural history of many booid snakes remains poorly known. How-
ever, the largest boas, the highly aquatic anacondas (Eunectes), are relatively well known. The
oldest and largest member of the clade (Titanoboa) was aquatic, and today, the most aquatic
members of the family are the anacondas of the genus Eunectes.

Anacondas, Eunectes
The Anacondas are Neotropical snakes found east of the Andes. Four extant species and
one fossil species compose the genus. The clade includes the largest snake, the Green Ana-
conda, Eunectes murinus, which reaches at least 8 m and has the greatest mass (in females)
of any extant snake (250 kg). Eunectes is derived from a Greek word that means “good swim-
mer.” The aquatic habitats of these snakes are well documented. Pizzatto et al. (2007) found
an aquatic habitat to be a synapomorphy for this genus. The large body size, a relatively long
tail, and the dorsal positioning of the eyes and nares are all traits associated with life in water.
Anacondas are not the longest snakes but the most massive (a high mass/length ratio).
The giant snakes have the most specialized habitat requirements—they are almost entirely
aquatic. The Green Anaconda is the species that receive the most attention, but three other
species are smaller, and they remain poorly studied. Hard evidence for a Green Anaconda
exceeding nine meters is elusive. However, the Green Anaconda is an apex predator in the
Amazonia’s aquatic and terrestrial ecosystems, thus of considerable ecological importance.
The timing of reproduction varies with the species and location but tends to be associ-
ated with the dry season. Females attract males with a pheromone, resulting in polyandrous
breeding aggregations with multiple males coiling around a female, forming breeding balls.
This mating system has been observed in three of the four species in the genus.
Sexual cannibalism has been reported in Eunectes and its sister clade, Epicrates. Females
benefit from the post-copulatory meal through the additional nutrients and energy that can
be given to their offspring. Facultative parthenogenesis in captive females that were not in
contact with males for more than eight years was documented by Shibata et al. (2017).

86
 4. Ancient but Extant Lineages

Beni Anaconda
Eunectes beniensis Dirksen, 2002
Distribution and Habitat. Known only from the vicinity
of the Type locality: Type locality: Bolivia (Beni: Trinidad)
(Figure 4.24).
Natural History. Poorly known. In one of the few studies
on this snake, De la Quintana et al. (2017) used radiotelem-
etry to study the home range and habitat use of nine adult
Beni Anacondas. The authors located the snakes 242 times
Figure 4.24. The distribution in the wet and 255 in the dry seasons. The mean wet season
of the Beni Anaconda, Eunect- home range was 25.81 ha (6.7 to 39.4 ha), while the mean
es beniensis.
dry season home range was 0.29 ha (0.13 – 0.42 ha). Beni

Figure 4.25. The Beni Anaconda, Eunectes beniensis.

Photography by Myke Clarkson.

Anacondas seem to prefer swamps while avoiding forest and rice fields. However, individ-
ual snakes’ habitat use appears to vary based on the habitats available within their home
range. Notably, rice fields were avoided by most individuals.

DeSchauensee’s Anaconda
Eunectes deschauenseei Dunn and Conant, 1936
Distribution and Habitat. Wetlands, swamps, and sea-
sonally flooded areas of the Amazonian savanna in the
Brazilian states of Pará and Amapá and in French Guiana
(Dirksen and Henderson 2002). Including Marajó Island and
the adjacent regions at the mouth of the Amazon River and
Figure 4.26. The distribution several drainages French Guiana. It may also be found in ad-
of the DeSchauensee’s Ana- jacent Suriname, and the range may be disjunct. DeSchauen-
conda, Eunectes deschauenseei see’s Anaconda is a lowland species occurring between sea
level and 300 m above sea level. Type locality: Marajo I; Pará,

87
Aquatic Snakes

Northeast Brazil. Habitat loss and degradation may be a problem for this species (Figure
4.25).

Green Anaconda
Eunectes murinus (Linnaeus, 1758)
Distribution and Habitat. The Green Anaconda uses
aquatic habitats in tropical wet forests, tropical dry forests, and
savanna Distribution and Habitat. The Green Anaconda uses
aquatic habitats in tropical wet forests, tropical dry forests, and
savanna at elevations below 240 m. In Venezuela, it is wide-
Figure 4.27. The distribution spread in the Orinoco drainage system and is most likely found
of the Green Anaconda, Eu- throughout the country; there is the possibility of an isolated
nectes murinus. population in the Maracaibo Basin (Barrio-Amorós and Man-
rique 2007). It is endemic to the Amazon Basin and peripheral
drainage systems and is
restricted to rivers and
wetlands. Populations
are known in Colombia,
Venezuela, Guyana, Su-
riname, French Guiana,
Ecuador, Peru, Brazil,
Bolivia, and probably
Paraguay. A population
also exists on the con-
tinental island of Trini-
dad, near the mouth of
the Orinoco River and
the northern edge of the
range (Figure 4.27). Type
locality: “America.”
Natural History.
Monitored neonates were
always in shallow water
(less than 20 cm deep).
Their cryptic coloration
made them difficult to
see in water with hya-
cinths and grasses (Ri-
vas et al. 2016). The only
well-studied population
of Green Anacondas is in
the Venezuelan Llanos.
The Llanos is a mosaic
of savannas, rivers, flood
Figure 4.28. The Green Anaconda, Eunectes murinus. plains, lagoons, palm,
Photography by William W. Lamar.
and gallery forests. Ra-
dio-tracked snakes were

88
 4. Ancient but Extant Lineages

found in the water 86% of the time and at the water’s edge the other 14% of the time they
were located (Rivas, 1999). Rivas (2000) found that anacondas have relatively distinct home
ranges. During the dry season, they have a mean home range of about 25 ha. Wet season
home ranges are more extensive and average 37.4 ha. They are active during the day, with
activity peaking in the evening. Late-night activity was expected.
The Green Anaconda is an ambush predator (Rivas, 2000) but may also actively forage
for food (Allen, 1963). Müllner and Linsemair (2007) observed an individual that had fed on
a nest of fledgling Hotzans, a bird known for over-water nesting. Populations living in wet-
lands with a dry season, like the llanos, may experience increased prey densities as wetlands
reduce their size in the dry season, and fish and other aquatic animals are concentrated in
small areas. Rivas (2000) reports that some individuals move to rivers in the dry season, while
others remain in drying pools and estivate as the water evaporates.
Gravid females are relatively sedentary and probably do not forage. Parturition occurs at
the end of the wet season between October and late December, with a gestation of 202.6 days.
Most births occur in the evening after the peak heat has passed, and birthing lasts between
20 to 40 minutes, but some females take up to 145 minutes. Litter size was correlated with the
mass of the female and averaged 29.4. The neonates average about one percent of the mass of
the female. The sex ratio at birth, calculated from all the newborns obtained in the study, was
even. Neonate anacondas did not show any sexual size dimorphism. Neonates in captivity
have been reported to grow at 2.13 mm/day (Lamonica, 2007), while wild individuals grow
much slower (0.068–0.074 mm/day) (Rivas et al. 2016).
Neill and Allen (1962) observed a captive female E. murinus give birth to four young
and 19 undeveloped eggs. After parturition, the female ingested the undeveloped eggs. Rivas
(2000) found relatively large numbers of stillborn young and attributed it to the stress of han-
dling during the study. Still, this seems true for captive animals that should be acclimated to
captivity and handling.
Neonates have a diet composed of birds, the same prey as young adult snakes. The ab-
sence of small caiman, hatchling turtles, fish, and frogs in the diet was unexpected, although
adult anacondas rarely eat fish and amphibians. The small relative prey size, their low feeding
frequency, and their lack of mobility suggest a low metabolism. The young prefer stagnant,
shallow water covered by aquatic vegetation, essentially the same habitat used by the adults
(Rivas et al. 2016). Adults capture prey as large as adult capybaras (40–55 kg), adult white-
tailed deer (55–70 kg), and full-grown spectacled caimans (35–55 kg). Anacondas are dietary
generalists who take any prey they can subdue and swallow. Anacondas hunt almost exclu-
sively from the water.

Yellow Anaconda
Eunectes notaeus Cope, 1862
Distribution and Habitat. Distributed from about 15°S in
Bolivia and Brazil to 32°S in Argentina. It inhabits wetlands, in-
cluding swamps, seasonal floodplains, palm savanna, marshes,
gallery forests, and oxbow lakes. In Argentina, it is restricted to
the northeast portion of the country in the states of Corrientes,
Figure 4.29. The distribution Chaco, Entre Rios, Formosa, Misiones, and Santa Fe. In Bolivia,
of the Yellow Anaconda, Eu- it occurs in the northeast border region with Brazil. Here it is
nectes notateus. sympatric with the Green Anaconda. In Brazil, the Yellow Ana-
conda inhabits the western Pantanal in Mato Grosso, and Mato

89
Aquatic Snakes

Figure 4.30. The Yellow Anaconda, Euneces notaeus. Photography by JCM.

Grosso do Sul. In addition, there is an outlying, isolated population in São Paulo Province.
In Paraguay, the species is widespread, inhabiting the Paraguay, Parana, and Pilcomayo River
Basins. In Uruguay, it is known from only one specimen from Achaval Province (Figure 4.29).
The type locality is the Paraguay River and its tributaries.
Natural History. Prey items were collected from 49 males and 37 females by Waller et al.
(2007). They found all vertebrate classes represented. Rodents were eaten most often (67%)
by both sexes. The largest female examined was 3.1 m and contained hairs from a Capyba-
ra. The most common prey was the Water Rat (Holochilus chacarius). Birds found in nine
snakes included cormorants, limpkins, and wading species. Three individuals had eaten the

90
 4. Ancient but Extant Lineages

semi-aquatic dipsadid snake Hydrodynastes gigas.

Reproduction. Waller et al. (2007) found sexual maturity is reached in males at a body
length of 1.28–1.43 m. The smallest reproductive female was 1.47 m in body length. Mating
occurs in September and October. Females showed synchronous reproductive cycles. Ovidu-
cal eggs were found in October and November. Neonates are born in April. The average litter
size for wild-caught snakes is 11, and the size of the litter is positively correlated with the size
of the female.

Malagasy Booids, Family Sanziniidae

Habitat use and foraging habits of the Malagasy Ground Boa genus Acrantophis are poor-
ly known. However, Gardner et al. (2017) observed Acrantophis dumerili in a pond (in a cave)
with most of its body submerged. They also reported A. madagascariensis submerged in a
pond at a different location. In both instances, they suggest the snakes were foraging from
ambush. Both observations occurred in semi-arid areas lacking abundant surface water. In
such contexts, diverse species may visit water to drink and thus occur at higher densities than
in the surrounding landscape, increasing the probability of encountering prey. These snakes
may not be using aquatic habitats regularly.

Figure 4.31. The Malagasy Ground Boa, Acrantophis dumerili. Photography by JCM.

Asian Pipe Snakes, Family Cylindrophiidae

The thirteen species of Cylindrophis are considered fossorial in much of the literature.
However, Cylindrophis ruffus composes part of the Tonlé Sap snake harvest, which is collect-
ed using gill nets. A telemetered individual in southern Thailand stayed at the exact under-
water location for several weeks, not moving more than a few meters (Murphy et al. 1999,
Karns et al. 2005, Brooks et al. 2009). Cylindrophis is superficially similar in its ecology to the

91
Aquatic Snakes

Western Hemisphere Anilius scytale. The two families share similar, superficial appearances,
ecology, and defense behaviors. These snakes are likely adapted to inhabit flooded burrows.
Cylindrophis is widespread in southeastern Asia, ranging from Myanmar, Laos, Viet-
nam, Cambodia, Thailand, and the Malay Archipelago, including Singapore, both penin-
sular Malaysia and eastern Malaysia (Borneo); in the Indonesian Archipelago, they occur
in the Greater Sunda Islands (Borneo, Sumatra, and Java, as well as some of their offshore
islands), Sulawesi, the Lesser Sunda Islands (Lombok, Komodo, Flores, Sumbawa, Timor
including Timor-Leste, and east to the Maluku Islands (Halmahera, Wetar, Damar, Babar,
and into the Tanimbar Archipelago). The eastern distributional limit, sometimes given as the
Aru Islands off the southwestern coast of New Guinea, is questionable. They are known from
Sri Lanka but are not known from India. They are present in southeastern China (Fujian,
Hong Kong, and Hainan Island).
Pipe snakes of the genus Cylindrophis have a relatively blunt rounded head with large
plates, not distinct from the anterior body, the eyes are small, and a mental groove is present.
The body is uniformly cylindrical; the ventral scales are narrow and barely wider than the
dorsal scales; they have a pair of pelvic spurs in both sexes; the tail is exceptionally short tail,
often with aposematic ventral coloration. The body is cylindrical, with a nearly uniform
diameter, thus the name pipe snakes. The eyes have round or vertically subelliptic pupils.
The head has large plate-like shields, with the nostrils in a single nasal scale; a loreal scale
is present; the postocular scale is small. The dorsal scales are smooth, in 17 to 23 rows de-
pending on the species. They range in size from small (120 mm) to medium sized snakes
(1000 mm). The life history and habits of these snakes need more documentation (Figure
4.34). Some Cylindrophis have a coloration and pattern that makes them appear very similar
to North American Farancia.

Figure 4.32. The Oriental Pipe Snake, Cylindrophis ruffus.

Photography by Kenneth Chin.

92
 4. Ancient but Extant Lineages

On the next page. A Little File Snake, Acrochordus granulatus, is in a

puddle. Photography by Kenneth Chin.

93
Aquatic Snakes

94
Aquatic Snakes

5. File Snakes
Myke Clarkson and John C Murphy

The family Acrochordidae is a fully aquatic snake lineage comprised of three extant
species (Acrochordus javanicus, Acrochordus granulatus, and Acrochordus arafurae) and one
extinct species (Acrochordus dehlmi). The extant species are found throughout the Indo-Aus-
tralian region, ranging as far west as three file snake species range from the North West coast
of India and as far East as the Philippines Sea and the Solomon Islands, with Acrochordus
granulatus occupying the most extensive range of the three (McDowell 1979).
Fossil records from the Miocene of the extinct Acrochordus dehmi indicate this species
had a more inland distribution earlier in its history, with fossil evidence found in Siwalik, Pa-
kistan, Nepal, northern India, and Thailand. Only a single fossil in Thailand has been found
within the group's extant range, suggesting an Asian origin with later dispersal to Indonesia
and Australia. Australian fossil records only date to the Pliocene, indicating Australia as a
more recent range for the group. In addition to a more inland range, Acrochordus dehmi was
the largest of the known Acrochordus, with an approximate size of ~3 meters (Head et al. 2007
and Sanders et al. 2010).
The highly aquatic and unique file snakes have loose, baggy skin and minor, rough
scales, each with sensory or-
gans. Their short tails are
round, prehensile, flat, and
sword-like; the eyes and val-
vular nostrils are dorsally or
dorsolateral. Their lack of
muscle tone is spectacular and
can only be fully appreciated
by handling one; a physiolog-
ical feature restricting them
to an entirely aquatic lifestyle.
The rough, file-like surface
of the skin and its loose folds
are used in catching and hold-
ing slippery, mucus-covered
fish. Their maxillary teeth
are similar in size, long, and
fluted on the sides and pos-
terior surfaces. The fluted
condition is found in many
fish-eating snakes. File snakes
usually have more than 100
teeth. Their thick bodies,
slow movements, sluggish
metabolism, and complete
reluctance to leave the water Figure 5.1. Acrochordus javanicus 15 kg, 1.95 cm from Bangkok,
render them helpless out of Thailand. Mr. Eakasak Kanjaruke is the person in the photo. Pho-
the water (McDowell 1979). tography by Kittipong Jarutani.

95
 5. File Snakes

The extant file snakes range in size from the 1.1 m Acro-
chordus granulatus to the 2.9-meter A. javanicus, though
fossil evidence of Acrochordus dehmi suggests a much
larger size earlier in their evolution, with specimens.
Although the A. javanicus size record has been ques-
tioned because it is old and may be an error, most large
specimens reach about 2 m. File snakes tolerate various
salinity levels and inhabit fresh, brackish, and marine en-
vironments.
The Little File Snake (Acrochordus granulatus) is
the most marine of the three species and has been found
15–20 km offshore. It has the highest blood volume of
any snake (13% of its body weight, most snakes are in
the 5–6% range), and red blood cells compose 50% of its
blood volume (most snakes are in the 25–30% range).
Thus, it has the largest oxygen-carrying capacity of any
snake. Underwater the file snake’s heartbeat is slow, but
in anticipation of taking a breath as the snake surfac-
Figure 5.2. A dorsal view of the head es, the heartbeat increases so that more oxygen can be
of A. javanicus that is covered with al-
taken up by the blood flowing through the lungs. Like
gae. Photography JCM
other aquatic snakes, file snakes exchange blood gas-
es through their skin, increasing their dive time. File
Snakes living in freshwater do not have the high blood
volume or the oxygen-carrying capacity found in ma-
rine species.
Ecological studies on the Arafura File Snake sug-
gest that it may not have a specific home range; instead,
they wander through the murky water. Some individual
file snakes stay at one location for a day or more, while
others may move almost a kilometer. The freshwater
Arafura file snake lives in a seasonally wet climate, and
their wetland habitat expands and contracts with the
rains as they follow the water. During the dry season,
populations may be highly concentrated in small areas,
which offers an opportunity for mating. Multiple males
compete for females in these situations, and it seems
likely that sperm competition is present in this species,
as it is in most snakes. With the rain, the area covered
by water expands, and the snakes disperse to distant lo-
cations, lowering the density of the snakes. Like all an-
Figure 5.3. Note that acrochordids
imals, file snakes are significantly impacted by climatic
lack plate-like scales on the top of the
head. Photography JCM
events. Madsen and Shine found high rainfall late in the
wet season resulted in abundant fish populations and fat
file snakes the following year.
Like many aquatic snakes, acrochordids laterally compress their body when swimming.
The compression produces a keel on the belly and increases the body’s surface area, making
the snake a more efficient swimmer.
Questions about file snake relationships were largely unresolved until DNA sequencing
and genetic comparisons became possible. While some authors hypothesized that acrochor-
96
Aquatic Snakes

dids were related to boas and pythons, others considered

them colubrids. DNA sequences suggest that file snakes
are the sister species to all the advanced snakes, the Caeno-
phidia, and last shared an ancestor with them at about 90.7
Ma (Vidal et al. 2009). Sanders et al. (2010) examined re-
lationships within the genus using mitochondrial and two
nuclear gene sequences. Acrochordus javanicus is the sister
to A. arafurae + A. granulatus, counter to expectations from
superficial ecology, external phenotype, and former taxon-
omy. They reviewed and revised fossil calibrations for dat-
ing snake divergences. A Bayesian relaxed-clock analysis of
the two nuclear loci found interspecific variations among
extant species during the Miocene about 16 and 20 MYA,
making this group one of the oldest extant marine snake
lineages predating actual sea snakes (Hydrophiinae), which
diverged ~7 MYA, and sea kraits (Laticaudinae) which first
appeared ~13 MYA (Sanders et al. 2010).
New morphological data for A. arafurae, and their
molecular timescale supported the placement of the fossil
taxon A. dehmi within the Acrochordus crown group as the
sister to A. javanicus among nominate species. The study
highlights the three Acrochordus species as old and highly
Figure 5.4 Scales on Acrochor-
distinct lineages. dus javanicus are differentiat-
File Snakes share unspecialized head scales with the boas ed on various regions of the
and pythons, but they are united with colubroids in that they body. Note the lateral scales
lack vestigial limbs; males have spines on their hemipenes, in the top photo show three
and they have a well-developed vomeronasal system. Traits cusps. Scales on the dorsum
unique to acrochordids include skin sense organs, a passive have strong keels but no cusps,
joint between the frontal and parietal skull bones, and the and the midventral scales are
shape of the head on the ribs. Acrochordids also have an spine-like. Photography by
JCM.
unusual lung morphology, with a double row of small lung
pouches on the trachea and a more folded intestinal tract
than seen in other snakes. Acrochordids can remain sub-
merged for as long as 2.3 hours. When they surface, they take
about five breaths per minute. The first several breaths oxy-
genate the blood, and the last fills the multi-chambered lung.
All three species show sexual dimorphism in body and
tail sizes. Reproduction is seasonal in all three species. Ovu-
lation occurs about July, and parturition follows five or six
months later.
Male file snakes mature at about six years of age; females
mature at about nine years. Only a small proportion of fe-
males are reproductive in any given year, and only the most
prominent females reproduce frequently. Acrochordus arafu-
rae average about 16 young (with a range of 9–25). One cap-
Figure 5.5. Scales on the snout
tive Acrochordus arafurae gave birth to a single young after have a sense organ. Photogra-
seven years of isolation, suggesting file snakes can use fac- phy by JCM.
ultative parthenogenesis to reproduce (Dubach et al. 1997).

97
 5. File Snakes

The three extant species of these aquatic snakes compose this ancient Asian – Austral-
asian family. One species is mostly freshwater (Acrochordus arafurae), one is primarily ma-
rine (Acrochordus granulatus), and one (Acrochordus javanicus) is a habitat specialist. It uses
streams running through river deltas and it is likely exposed to brackish water. Some authors
suggest file snakes are constrictors; however, how they restrain prey with folds of skin covered
with scales with keels and cusps. This is not the same as constriction in other snake lineages.
Sensory organs occur on tubercles on the interstitial skin between their scales and each scale.
These likely provide information about the presence and position of prey. Like many snakes,
acrochordids use both ambush and active foraging hunting strategies. Most hunting behavior
occurs at night.
During the day, file snakes take refuge by staying in the shadows created by vegetation
overhanging the water. They may even move with the clouds to confuse predatory birds
(Shine 1985).

Arafura File Snake

Acrochordus arafurae McDowell, 1979
Distribution and Habitat. This species is restricted to the
freshwater drainages of New Guinea, and Australia connected
to the Arafura Sea (including the Gulf of Carpentaria). Shine
(1986) found this snake in Northern Territory billabongs during
the dry season, expanding their range into inundated grassland
with the onset of wet-season flooding. In New Guinea, known
from the Mimika and Lorentz Rivers (Irian Jaya), the Fly-Strick-
Figure 5.6. The distribution
land River system, including Lake Daviumbo and Lake Mur-
of the Arafurae File Snake,
Acrochordus arafurae.
ray, the smaller rivers (Bensbach, Binaturi, Oriomo, Pahoturi)
of southwestern Western Province, Papua New Guinea, and
the Aramia River drainage. In Australia, they are present in the
Daly, Alligator, Koolatong, and McArthur River systems, North-
ern Territory, and the Leichardt, Gilbert, Mitchell, Lukin, Edward, and Archer River systems
of Queensland. The Australian distribution is along the coast from the Kimberly region of
Western Australia to the eastern (Coral Sea) coast of the Queensland Peninsula. Although
the Fly and Aramia Rivers of New Guinea open into the western end of the Coral Sea, they
are connected by swampy lowlands to the rivers draining into the Arafura Sea. Type locality:
Lake Daviumbo, western Province, New Guinea (Figure 5.6).
Natural History. Telemetered file snakes were sedentary during daylight hours but moved
extensively at night (Shine and Lambeck 1985). Sex differences in habitat use during the wet
season were evident. Females were often collected in deep-water (more than one meter) nets,
and males were collected in shallow-water nets (less than one meter deep). Most (88%) of the
females from deep water were adults, while many females from shallow water were juveniles.
Most males (87%) and immature females (69%) were in shallow water. Adult females occur
in both habitat types (54% in deep water). During the dry season, a female sex bias related to
water depth or maturity was found. The proportions of juvenile to adult males did not differ
between deep and shallow water in either comparison.
Shine (1986) found sexual dimorphism in A. arafurae is consistent with the hypothesis
that sexual selection is responsible for body length sexual dimorphism. Female reproductive
success depended on body size—larger females produce larger and more frequent litters. In
contrast, the reproductive success of males probably does not rely so much on body size—

98
Aquatic Snakes

multiple males in shallow water court females. The observa-

tions of several males around one female suggest that male-
male aggression may not occur.
However, sexual dimorphism in A. arafurae involves
more than body length. The longer tails of males presumably
reflect the need to fit the hemipenes into the base of the tail:
this tail-length dimorphism is widespread in reptiles. Sexual
differences were also apparent in the prey taken by each sex,
and adult females ate much larger prey than did adult males
(mean prey masses 291 g in females as opposed to 24 g in
males). Prey-size differences were evident even in males and
females of the same body size.
A four-year mark-recapture study of the Arafura File-
snake in the “wet-dry” Australian tropics of Kakadu National
Park by Houston and Shine (1994a) showed less movement
on moonlit nights. Trap sites under overhanging vegetation
consistently caught more snakes than trap sites in open wa-
ter, and juvenile snakes were mainly caught in areas covered
in floating vegetation. The authors suggest that the prefer-
ence for nights, traveling primarily through heavily vegetat-
Figure 5.7. The skull of an Acro- ed areas, is adaptive in reducing their vulnerability to pred-
chordus javanicus. Photography ators (especially visually hunting birds). The movements of
by JCM marked filesnakes suggest they are nomadic instead of hav-
ing fixed home ranges. Larger snakes moved greater distanc-
es; males were more likely to remain at their initial capture site than females, but dispersal
patterns of males and females were similar in most respects. Filesnakes sometimes move
between billabongs during the wet season when large expanses of shallow water connect the
billabongs.
Some individual file snakes stay at one location for a day or more, while others may move
almost a kilometer. The freshwater Arafura File Snake lives in a seasonally wet climate, and
its wetland habitat expands and contracts with the rains as they follow the water. Population
dynamics in these snakes are driven by rainfall in northern Australia. During the dry season,
populations may be highly concentrated in small areas that offer mating opportunities. Mul-
tiple males compete for females in these situations, and it seems likely that sperm competition
is present in this species, as it is in most snakes. With the rains, the area covered by water
expands, and the snakes disperse to distant locations, lowering the density of the snakes. Like
all animals, file snakes are impacted by climatic events, and Madsen and Shine (2000) found
high rainfall late in the wet season resulted in abundant fish populations and fat file snakes
the following year.
The following description of A. arafurae dive physiology and behavior is from Pratt
(2009). File snakes respire bimodally, using both atmospheric and aquatic (cutaneous) gas
exchange. Pratt (2009) investigated its ability to up-regulate cutaneous oxygen uptake to
compensate for the temperature-induced increases in metabolic rate. She found cutaneous
oxygen uptake was independent of temperature, and the snake met all elevated metabolic
demands by increasing atmospheric oxygen uptake. Consequently, the maximum dive dura-
tion was reduced by 70%, from 77 minutes at 20°C to 28 minutes at 32°C. Cutaneous oxygen
uptake significantly contributes to dive duration. When the ability to respire aquatically was
removed (severe hypoxia), dive duration was reduced by up to 30%.

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 5. File Snakes

Figure 5.8. The Arafurae File Snake, Acrochordus arafurae.

Photography by Scott Eipper

Acrochordus arafurae infrequently feeds on large meals; thus, the post-eating metabol-
ic response requires more energy. After fasting for one month, peak oxygen consumption
was up to 12 times the standard metabolic rate. The substantial reductions in dive duration
following feeding may reduce the snake’s vulnerability to predators. Acrochordus arafurae is
prone to predation bybirds of prey and aquatic predators such as crocodiles and large fish.
Simulated avian predation did not change dive or surface duration or proportion of time at
the surface or spent active. However, a larger number of longer dives were observed with
fewer long surface intervals suggesting an increase in cutaneous oxygen uptake.
At night, A. arafurae became more active; its reduced dive duration and increased sur-
face-active suggest it is foraging. The mean dive time was 6.6 minutes, with 85% of dives less
than 10 minutes. The maximum dive duration was 153 min. Snakes were located at a mean
depth of 0.62 m; however, they occasionally dove to 6 m, but very infrequently.
Pratt et al. (2010) remotely monitored diving in free-ranging A. arafurae. The average
dive depth was 0.62 m, and 95% of dives had an average depth of one meter or less. The
average dive duration was 6.6 min, and 84% of dives were terminated within 10 min, but
all snakes performed dives 450 min during the 14-day observation period. The authors hy-
pothesized that dive behavior was strongly influenced by predation pressure. The short dives
within the aerobic dive limits reduced the time they needed to spend at the surface on each
breathing bout, reducing the risk of bird predation. Predation is a strong selective force that
might alter the time allocation during dive cycles.
Growth increments of snakes (71 males, 77 females) recaptured after periods of more
than six months were used to estimate growth curves. The low metabolic rates, and slow
growth rates suggest these snakes mature at ages older than most other snakes. Male file-
snakes mature at about five years, and females mature at about seven. Male grew more slowly
than females as juveniles and matured at smaller body sizes (87 cm in males vs. 117 cm SVL
in females) (Houston and Shine 1994b).

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Aquatic Snakes

Little File Snake

Acrochordus granulatus Schneider, 1799
Distribution and Habitat. It occurs on both coasts of the
Indian Peninsula, Sri Lanka, and is widespread in the Indo-Chi-
nese and Malayan regions; it is also present in the Philippine Is-
lands; the Greater Sunda Islands, Sulawesi, and Flores and Timor
in Lesser Sundas; the coasts of New Guinea and is present in the
Lorentz, Sepik, and Fly-Strickland river systems. In addition,
Figure 5.10. The distribu- it occurs on the islands of Biak and Japen Islands in Geelvink
tion of the Little File Snake, Bay; Kiriwina Island, Trobriands, Mussau Island and the Wil-
Acrochordus granulatus. laumez Peninsula of New Britain in the Bismarck Archipelago,
Bougainville Island, and at least Morgusaia (Shortland Group),
Santa Isabel, Malaita, and Guadalcanal
Islands in the Solomon Islands. It in-
habits coastal marine waters and some
freshwater lakes and rivers. Type lo-
cality: None given (Figure 5.10).
Natural History. The habitats of
A. granulatus reported in the litera-
ture (Cantor, 1847; Flower, 1899; and
Boulenger, 1912) include marine and
freshwater environments in coastal
rivers of south and southeastern Asia.
Dunson and Dunson (1973) found
this species in full seawater and fresh
water. They also reported on snakes
collected over a shallow reef flat ex-
posed at low tide near Townsville,
Queensland.
The diet includes gobies and
goby-like fish that are small and de-
mersal, often in caves, under rocks,
and crevices. These fish reproduce
and care for their young in burrows,
suggesting that A. granulatus feeds by
exploring crevices and burrows along
the bottom. Four of the fish species
eaten by A. granulatus are also preyed
upon by the venomous sea snake, Hy-
drophis (Lapemis) hardwickii (Voris
and Glodek 1980)
Figure 5.12. (A) The Little File Snake,
Acrochordus granulatus. (A-B) Photogra-
phy by Harry Ward-Smith. (C). Photog-
raphy by Law Ing Sid.

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 5. File Snakes

The Little File Snake produces small litters of 6–12 (mean = 4.8 offspring) and they have
a relative clutch mass (RCM) that is lower than in its larger relatives.
Like other marine-dwelling species, it drinks freshwater and has a dehydration threshold
for the first drinking response. A deficit of 7.4% of the original body mass will trigger it to
drink from the freshwater lens on the ocean's surface. The thirst mechanism in this species is
more sensitive than that of sea snakes. The volume of water ingested increases with increas-
ing dehydration. In marine environments, freshwater is obtained at the water’s surface, and
A. granulatus tests the water with tongue flicks between swallows of water. Large volumes of
freshwater may be consumed, and approach 50% of the snake’s body mass (Lillywhite et al.
2014).
The Tiger Shark is a known predator of this snake in Northern Australia (Lyle and
Timms, 1987).

Karung
Acrochordus javanicus Hornstedt, 1787
Distribution and Habitat. Southeast Asia - a Sunda-
land-Indochinese species. Indonesia (Borneo, Java, Kaliman-
tan, Sumatra; the range in eastern Indonesia is unknown);
Cambodia; Malaysia (Malayan Peninsula and East Malaysia);
Singapore; Thailand; and Vietnam.

The Type Locality of Acrochordus javanicus Hornstead

There is some confusion in the literature concerning the type locality of A. javanicus.
Shaw (1802:573) suggests the type specimen came from a “large pepper-ground near San-
gasan” Java. Smith (1943: 132) reports the type locality as Java. Steubing (1991:329) reports
the type locality as Bantam (also spelled Bantem), W. Java. And McDiarmid et al. 1999:229
reports both of these literature accounts and make no judgment as to the actual type locality.
My translation of Hornstead follows, parentheses with italics are Hornstead’s words, and
parentheses with bold are my interpretation. He wrote,
During my stay on the island of Java, in the years 1783 and 1784, I had the
pleasure, during a journey from Bantam, of discovering one of the largest
snakes there are in the Indies, and one that until this time has evaded discovery
and thus the scrutiny of natural scientists. The snake was found inside a big
pepper forest at Tangaran. A Chinese in our travel party brought him (the
snake) alive to Batavia, using a forked bamboo stick to fix/hold the head. As
the snake was too big to be stored in alcohol, I had it skinned, and the Chinese
chopped up the meat, which they fried and boiled and used for delicious
courses/dishes. I stored the skin in arrak and it is now part of His Majesty
the King´s priceless collection. When the snake was opened we found, beside
an abundance of semi-digested fruits, five young of (halfannat kvarters) (20-
25 cm) length, which probably were the reason for the bulky/bulging belly/
stomach of the female snake.
Although this snake had all the characters other snakes possess, it struck me
as peculiar that it lacked, not only the scales under the belly and tail (= ventral
and subcaudal scales), which are the only the holy Creator have bestowed upon

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Aquatic Snakes

this naked genus so that they may move with speed from one place to the next,
and which are also a common feature for all snake types known to date; (but
also lacked) the rings and wrinkles (annuli et rugae) characterizing the two
latter genera of snakes in von Linnaeus’ system. In contrast to other snakes,
which possess smooth and shiny skin, this snake was instead covered by warts
all over, warts which were rough and covered the upper as well as the lower
side (of the snake). Thus, in this respect it could not be placed in any genus of
hitherto known snakes, why I call it Acrochordus, and whose description I here
have the honor of submitting to the Royal Academy of Science.
Thus, Hornstead (1787) described Acrochordus javanicus from a specimen (Figure 5.15)
collected in a pepper forest at “Tangaran,” Java. I have not found any localities on Java with
this spelling; however, about 68 km northwest of Bantem in West Java is Tangerang, now a
suburb of Jakarta located on the Cisadane River. The 18th-century Cisadane drainage un-
doubtedly supported swamp forests with large stands of the tree Elaeocarpus macrocerus
(family Elaeocarpaceae), a tree with a fruit that looks remarkably like a pepper (Whitten et al.
1996; Wiart 2006). Thus, I am restricting the type locality of A. javanicus to Tangerang, Java,
Indonesia (~ 6º10’50”S 106º37’55”E).
Natural History. Only three of 100 hundred dissected A. javanicus examined by Shine et
al. (1995) contained identifiable prey remains. Two contained fish, and one contained a large
aquatic snail that may have been a secondary prey item. Most of the snakes also included
fishhooks embedded in their stomach walls.
Males mature at about 1.0 m, and females mature at about 1.14 meters. About 64% of
adult females were reproductive. Litter sizes ranged from 13–52 offspring (mean = 29.3)
and were significantly correlated with maternal snout-vent length. The relative clutch mass
(RCM) ranged from 0.15 to 0.47 (mean = 0.29). However, this estimate of RCM will be much
lower than the RCM at parturition, because of the increase in embryonic mass through ges-
tation (Shine, 1986).

Figure 5.15 The plate from Hornstead (1787) is the type of of A. javanicus.

103
 5. File Snakes

Figure 5.16. A captive specimen of Acrochordus javanicus.

Acrochordus is capable of sustaining breath holding due to its effective elimination of

CO2 through its skin. The snake has a regulated mode of breathing that maximizes its oxygen
usage and allows it to stay submerged for up to 90% of the time. The control system govern-
ing its breathing is sensitive to hypoxia and ends breath holds when oxygen stores are almost
depleted. Despite having a relatively small lung volume, Acrochordus has a well-vascularized
lung that enables it to breathe without conflicting with the buoyancy requirements of a bot-
tom-living lifestyle (Glass and Johansen 1976).
Shedding skin can be problematic for file snakes since they live in a low-friction envi-
ronment. According to Lillywhite (1989), the keratin layer comes loose from the body, and
the snake crawls out of it. Occasionally, however, the skin being shed can tighten around the
snake’s body and form a knot. In this case, the snake moves the knot towards its tail to push
the skin off.
Based on the biology of A. javanicus, it is unlikely that the current harvest will signifi-
cantly reduce wild populations (Shine et al. 1995). Factors such as the precipitation regime,
extensive and inaccessible habitat, lack of efficient capture techniques, high reproductive out-
put, and low economic value of the skins contribute to a sustainable harvest.

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Aquatic Snakes

On the next page. The odd-scaled snake, Xenodermus javanicus shows off its
unusual head and tail scalation . Photography credit Herpetoeditor CC-
BY-SA-4.0 and Wikapedia. The original image was cropped and the resolu-
tion increased to 300 dpi.

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 5. File Snakes

106
Aquatic Snakes

6. Odd Scaled Snakes

Jason Fantuzzi
While the Xenodermidae live only a marginally semi-aquatic lifestyle, they are here
included given their close phylogenic relations with the fully aquatic Acrochordoidea, and
to their extremely unusual ecomorphologies which may lend to at least a semi-aquatic ex-
istence. One of the most basal clades of the Caenophidia, the Xenodermidae are distributed
from the Greater Sunda islands of Indonesia northward through southeastern and eastern
Asia. The family reaches its highest diversity in Vietnam, which is home to seven Achalinus,
four of which are endemic (Zeigler, [Link]. 2020), and all species of the Fimbrios/ Parafimbrios
group. Several recent phylogenies have recovered an origination timeline for this unusual
group of snakes dating back to at least 45 mya and as early as 80 mya, at the end of the Cre-
taceous period (Burbrink & Pyron, 2011; Pyron, [Link]. 2014). While the timeline of origin
for these taxa is well realized, the relationships of the family within the Caenophidia is still a
matter of uncertainty, and even the most recent publications position them as incertae sedis,
a most unresolved placement. The relationships between the Xenodermidae and Acrochor-
didae were recognized as early as 1893 when the great taxonomist George Albert Boulenger
placed them within the same clade, the Acrochordinae based solely on morphology and anat-
omy. Phylogenies more than 100 years afterward have solidified that relationship, however
the exact placement of the two regarding the rest of Colubroidea has flipped back and forth
more than five times since 1998 (Durso, online. 2016). Notwithstanding, most studies recover
an earlier occurrence and more distant relation of the acrochordids to the colubroids. Current
investigators (Kraus & Brown, 1998; Kelly, [Link]. 2003; Oguiura, [Link]. 2010; Pyron, [Link]. 2013,
2014; Figueroa, [Link]. 2016) have recovered the xenodermids (sometimes coined Xenoderma-
toidea) as sister to the Colubroidea, together with Acrochordoidea, whereas both previous
and later phylogenies placed them as basal within the superfamily, with a caveat of varying
degrees of support. Recent phylogenies (Pyron, [Link]. 2013; Figueroa, [Link]. 2016; Zaher, [Link].
2019) found high support for a sister relationship between Stolizckia and Xenodermus, with
the former occurring earlier. These and other current studies have also found high support
for two distinct groups of Achalinus, though no further recognitions have been suggested at
the present. Once included in this group were the “wood snakes” (Xylophis spp.), now nested
within the Pareidae (V. Deepak, [Link]. 2019). The Xenodermidae are represented by 6 genera
comprising 28 species, all of which possess odd scalation, hence the family name, Xenos
(Greek for “strange” or “odd” and dermus, “skin”). Typically characterized by undivided sub-
caudals, juxtaposed (and often keeled) dorsolateral scales, and perhaps most intriguingly, by
pronounced discoid nasal scales. The arrangement, size, and structure of this bizarre scala-
tion is perhaps most pronounced in the type genus and monotypic, Xenodermus javanicus.
As its species epithet “javanicus” implies, populations of this species are unequivocally more
abundant on the Greater Sunda island of Java, the species’ type locality, than anywhere else
within its highly disjunct range. Despite an apparently widespread geographic distribution
according to historic records from the Isthmus of Kra border areas of Burma (Robinson &
Kloss, 1920), Thailand, and Peninsular Malaysia, although the latter two countries continue
to produce much more recent observations than the former. The bizarre scalation of this
species begins at the glossy and contrastingly pearly and black ventral scales and undivided
subcaudals, then upward to the matte grey dorso-lateral scales which are somewhat obliquely
arranged and largely juxtaposed to one another, finally followed by three ridges of osteo-
derm-like vertebral scales.

107
 6. Odd Scaled Snakes

The cephalic scalation is no less unusual than the

body, appearing mostly granulated throughout the head
with undifferentiated labial and gular scales. Within the
Xenodermidae, this species is the only taxon which lacks
any large, colubroid cephalic scales save for the enlarged
nasals and internasals. Dorsal scales number 40-50 at mid-
body (DeLang, 2017). Ventrals of males range from 171-
177, with 147-165 subcaudals. Ventrals and subcaudals of
females, 176-186, and 133-150, respectively. Labial scales
while slightly enlarged, remain small enough to number up
to 20 and 25 upper and lower, respectively: truly a non-col-
ubroid trait. The peculiarity of X. javanicus does not end
with its strange scalation. With a tail to body length per-
centage of up to 42% (50.3% according to De Rooij, 1917),
these snakes might possess the longest tails of any ser-
pents, alongside Scaphiodontophis and Urotheca spp. of
Central America. Males possess longer tails (38-44%) than
females (34-38%). Given a relatively high degree of pre-
hensility, the tails of X. javanicus may be used to traverse
low streamside vegetation while seeking semi-arboreal an-
uran prey, and it has been suggested to re-consider a sole-
ly burrowing or fossorial natural history of these snakes.
While we do not doubt the observations of authors involv-
ing the observed burrowing habits of Xenodermus, those
specimens found commonly occupying rice paddy fields
may have simply adapted to these anthropogenic alter-
ations as secondary hab-
itat due to an abundance
of anuran prey sources.
Most primary habitat
harboring these snakes
occurs as low to mid-el-
evation rainforest in hilly
country offering plentiful
Figure 6.1. The unusual scale ar-
streams, swamps, and
rangements of the Odd Scaled
Snake, Xenodermus javanicus.
other forested wetlands.
From M. Smith, 1943. While Xenodermus can
hardly be considered
the strongest swimmers, Figure 6.2.A. Distributioon of the fami-
observations have been made of javanicus swimming or ly Xenoddermatidae. B. Distribution of
rather “floating” (T. Charlton, personal communication) the genus Achalinus. C. The distribu-
in pursuit of anuran larvae and small fish prey. tion of Fimbros. D The distribution of
Most morphologically like Xenodermus are three Parafimbrios. E. The distribution of
species of equally rarely known and possibly anuran and Paraxenodermus. F. The distribution of
Stoliczkia. G. The distribution of Xeno-
fish-eating snakes from the northeastern Indian states
dermus.
of Meghalaya, Assam, Mizoram, and the Indo-Malayan
Island of Borneo. Until very recently, these three spe-
cies (including the newly discovered S. vanhnuailianai)

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Aquatic Snakes

Figure 6.3. The Odd Scaled Snake, Xenodermus javanicus. PT Karyanusa Eka Daya (Oil Palm Plan-
tation) in Muara Wahau Sub-district, Kutai Timur District, Kalimantan Timur Province - Indone-
sia Photography by Mediyansyah.

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 6. Odd Scaled Snakes

comprised the genus Stoliczkia. The genus name was erected for these unusual snakes by
Dr. Thomas Caverhill Jerdon honoring Dr. Ferdinand Stoliczka, an early Moravian (now
Czech Republic) zoologist who performed extensive surveys in India including the Hima-
layas, where he met an untimely fate, dying of spinal meningitis commonly known as Acute
Mountain Sickness, or AMS (Boelens [Link]. 2011). A recent updated phylogeny of the group
found borneensis as sister taxon to X. javanicus (Deepak, [Link]. 2021). Applying a multilo-
cus dataset including new mitochondrial and nuclear gene sequence data as well as sever-
al morphological character differences of the cephalic scutellation resulted in the authors’
creation of the genus name Paraxenodermus (gen. nov.) to represent “Stoliczkia” borneensis.
As earlier stated, Stoliczkia share a slim, laterally compressed body form with Xenodermus
javanicus including similarly long, flagellate tails comprising about a third or more of the
total body length. However, unlike the latter, the former possesses distinctly keeled lateral
scales and contrastingly enlarged cephalic scutation particularly from the parietals to the
internasals. Dorsal scales of S. borneensis number 30 at mid-body, with 210 ventrals and 124
subcaudals (De Rooij, 1917). Steubing, [Link]. 2014 report scale counts of 31-35 at mid-body,
with ventral scales of females ranging from 205-210 with 117-124 undivided subcaudals.
Upper labials number 10 or 11 with the sixth and seventh touching the eye, though these
may be separated from the eye by a row of subocular scales (Steubing, [Link]. 2014).
Of Stoliczkia and Paraxenodermus, perhaps the latter is the best known ecologically,
with a handful of observations and several photographs circulating online or published in
regional guides. One of these images portrays the species traversing rainforest dipterocarp
fronds within proximity of a forested stream. Indeed, it has been suggested that the elongat-
ed tails of this group (including Xenodermus) atypical to this family of serpents facilitate life
ecologies amongst a three-dimensional environment at least to some extent above the forest
floor, possibly in search of arboreal anuran prey. Besides these random, extremely sporadic
observations, this genus is represented by six specimens, of which four are accompanied by
legitimate institutional and cataloguing records. Even less is known about their behavioral
ecology.
The genus Achalinus is unquestionably the most speciose of the group with 26 spe-
cies currently recognized (Uetz, [Link]. 2023). Achalinus are small (60cm-75cm) burrowing
forms indigenous to the eastern aspect of Asia, from southern Japan’s Ryukyu Archipelago
and thenceforth southward through China including Taiwan, and into the hills and val-
leys of northern Vietnam. The stomach of a specimen (USNM 81513) contains earthworm
remains (Pope, 1935). Yamasaki and Mori, 2017 reported bimodal seasonal activity peak-
ing in early summer and then again in autumn, often coinciding with the appearance of
earthworms on the ground surface. They reported annual activity patterns that agreed with
most snakes of temperate regions, however unlike most of those examples, the hatchlings
of A. spinalis remained underground in cavities until the following spring/early summer.
A female from Chungan Hsien contains seven well-developed eggs, one of which mea-
sures 19x7mm (Pope, 1929). Pope opined on the fragile nature of collected Achalinus: “It is
extremely delicate and dies more readily than almost any other snake. Soon after death it
dries and hardens in a most unusual way. This drying up might lead one to conclude that
it is a secretive burrower but the greatly reduced rostral is anything but a characteristic of
burrowing species. It shows not the slightest signs of defensive behavior when handled.”
The “bearded” burrowing odd-scaled snakes: Fimbrios and Parafimbrios represented
by two species apiece. The generic names are from fimbria, Latin for “a fringe”, an allusion
to the strangely enlarged labial scales of these taxa, unique to the genera. It is highly possi-
ble that these fringes contain sensory epithelia (akin to the rostral appendages of Erpeton,

110
Aquatic Snakes

a homalopsid also from Indochina) which enable these burrowing and small-eyed snakes to
seek out annelid worms in complete darkness and largely incapable of tongue-flicking while
tunneling through porous substrate for prey. While recent phylogenies (Ziegler, [Link]. 2018,
Zaher, [Link]. 2019) place these two genera at a later node within the family, morphologically,
this group appears to fall somewhat intermediately between Achalinus and the Xenodermus/
Stoliczkia group, featuring the stouter bodies of the former with the oddly juxtaposed dor-
solateral scales and fragmented cephalic scales of the latter. Ecologically, they are typical of
all xenodermids in sharing a secretive lifestyle not far from wet habitats such as streams and
swamps, often sheltering beneath stones and decomposing logs.

On the next page.

A Northern Cottonmouth, Agkistrodon piscivorus threatens the

photographer with envenomation by opening its mouth and ex-
posing the white lining. Photography by William W. Lamar

111
 6. Odd Scaled Snakes

112
Aquatic Snakes

7. Aquatic Vipers
Very few vipers are considered aquatic or semi-aquatic. The reason for this is open to
speculation.
Copperheads and Cottonmouths of the genus Agkistrodon are distributed in North
America from the northeastern and central United States southward through peninsular
Florida and southwestern Texas, and in Central America on the Atlantic versant from Tam-
aulipas and Nuevo León southward to the Yucatan Peninsula, Belize, and Guatemala. In
addition, they occur along the Pacific coastal plain and lower foothills from Sonora south
through Guatemala, El Salvador, Honduras, and Nicaragua to northwestern Costa Rica.
There are eight extant species of Agkistrodon. They are semiaquatic to terrestrial and are
often found near water. Two species, A. contortrix, and A. bilineatus are also found in dry
habitats, often far from permanent streams or ponds.
Underwater hibernation by the Eastern Massasauga, Sistrurus catentatus, has been doc-
umented but poorly studied. While this is a mostly terrestrial species, it may spend the
winter submerged in a crayfish burrow (Yagi et al. 2020). Sistrurus are known to swim across
relatively larges bodies of water. Hullinger et al. (2018) reported a Western Massasauga, Sis-
trurus tergeminus, removed from the gut of the percid fish,
Sander vitreus, at Wilson Reservoir Russel Co., Kansas.
Submerged hibernation has also been reported in Vi-
pera berus (Viitanen 1967). While the risk of drowning re-
mains a problem for these snakes, spending the winter un-
derwater may increase the survival rate in some situations.

Figure 7.1. The distribution Florida Cottonmouth

of the Florida Cottonmouth, Agkistrodon conanti (Gloyd, 1969)
Agkistrodon conanti.
Distribution and Habitat. Marshes, lakes, ponds,
streams, roadside ponds, drainage ditches, coastal islands, mangrove swamps, and rice
fields from southern Florida to Savannah, Georgia, and west to southeastern Alabama.
A medium- to large-bodied semi-aquatic pit viper with an average adult size of 76–122
cm. Maximum size 189.2 cm (Gloyd and Conant, 1990; Conant and Collins, 1991), with a
tail/total length ratio of 0.15–0.19 in males and 0.13–0.18 in females.
Geography and color patterns distinguish the Florida Cottonmouth from related spe-
cies. The head is typically brown with vertical stripes along the snout on the rostrals, prena-
sals, and first upper labials (Gloyd and Conant, 1990). Dark stripes appear on the lower jaw
extending from the mental to the first four or five infralabials (Gloyd and Conant, 1990). A
dark cheek stripe is bordered above and below by pale stripes and is often present in adults,
although it may be indistinguishable from the ground color in adult A. piscivorus (Gloyd
and Conant, 1990; Conant and Collins, 1991).
Natural History. Agkistrodon conanti is a generalist predator with a diet composed of
invertebrates, fish, amphibians, reptiles, birds, and mammals. They will also feed on carrion
(Gloyd and Conant 1990).
A heron rookery on Seahorse Key, (a small island adjacent to the northern Florida pen-
insula in the Gulf of Mexico) contains a population of cottonmouths that feed on the fish
dropped by the birds. Young et al. (2008) report that a fish-based olfactory stimulus, without
associated ground-borne vibration or motion, triggered a foraging response in the Seahorse
Key cottonmouths. Snakes from the two study sites reacted differently to the olfactory stim-

113
 7. Aquatic Vipers

Figure 7.2. The Florida Cottonmouth, Agkistrodon conanti. Photography by Kaylyn Cullen.

ulus; those with less ground cover, more rats, and fewer snakes exhibited a weaker foraging
response. Sandfoss et al. (2018) reported that cottonmouths unexpectedly abandoned the
Seahorse Key rookery in April 2015 and did not return.
Males are larger than females and compete for females using combat behavior. The litter
size ranges from 1-15; the neonates are 21-27 cm in total length. Parturition occurs in August
and September.

Northern Cottonmouth
Agkistrodon piscivorus (Lacépède, 1789)
Distribution and Habitat. Found throughout the southeastern USA from southeastern
Virginia to central Georgia, east of the Appalachian Mountains, north to southern Illinois
and eastern Kansas, and southward into central Texas in the west (Fig. 7.3).
Hybridization with the Florida Cottonmouth occurs in the mid-Atlantic coastal plains
in southern North Carolina to the coastal plains in southeastern Louisiana. Diagnosis may
be difficult for some individuals in this area without additional morphological and molecular
data.
Natural History. Males consume significantly more fish, amphibians, and arthropods
than females, whereas females consume significantly more reptiles and mollusks than males.
Therefore, when expressing dietary preferences using the relative mass of the prey, arthro-
pods, mollusks, and mammals are no longer essential prey sources for either males or females
relative to the overall mass of prey consumed (Vincent et al. 2004).
Delisle et al. (2019) conducted a year-long telemetric habitat selection study on two
Northern Cottonmouth populations in northeast Texas. The home range selection of both
males and females exhibited a preference for edges and avoidance of pasture. In addition,

114
Aquatic Snakes

both sexes selected microhabitats with thick vegetation in

the spring and summer. However, in the fall, male microhab-
itat selection was influenced by lower temperatures and fe-
male microhabitat use appeared to be driven by parturition.
Life history data for the Cottonmouth was described by
Hill and Beaupre (2008) for a population inhabiting a “rif-
fle–pool” creek system (Rocky Grove) located in the Ozark
Figure 7.3. The distribution of
Mountains in northwest Arkansas. One hundred and for-
Agkistrodon piscivorus.
ty-two individual snakes were captured 283 times from Au-
gust 1996 through September 2003. Mean snout–vent length
(SVL) of adult Rocky Grove A. piscivorus (males 60.8, females 54.9 cm) was among the small-
est reported for any population of A. piscivorus. Rocky Grove A. piscivorus also exhibited a
low degree of sexual size dimorphism (SSD) compared to other localities, and mean female
SVL was 90.3% that of males. Growth rates were the slowest reported for any temperate zone
pitviper (males 0.151, females 0.178 cm/month). Limited data also indicate low reproductive

Figure 7.4. Northern Cottonmouth, Agkistrodon piscivorus, (A) profile of an adult. (B) a juvenile from
Union Co., Illinois, (C) an adult from North Carolina (D) an adult from Union Co., Illinois, (E) a
juvenile in a defense posture. Photography by (A-B, D-E) JCM; (C) Jason Fantuzzi.

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 7. Aquatic Vipers

output for this population in terms of frequency of reproduction and litter size. Female
reproduction averaged less than biennial, as evidenced by the consistently low proportion
of pregnant to non-pregnant females (18.4%) and the reproductive histories of individual
female snakes. Litter size averaged 4.1. Mating at Rocky Grove may occur in late summer
and spring. Agkistrodon piscivorus at this study site may be limited in energy acquisition
rates relative to the conspecifics in other parts of the range.
Eskew et al. (2009) characterized the foraging strategy and microhabitat use of cotton-
mouths at Ellenton Bay, an isolated Carolina Bay freshwater wetland on the Savannah River
site in South Carolina. They measured the habitat characteristics of 55 ambush sites used by
51 individual cottonmouths located visually at night and compared them to 225 randomly
selected locations. They found Northern Cottonmouths exhibited an ontogenetic shift in
foraging strategy, with juveniles using predominately ambush foraging around the edge of
the wetland. At the same time, adults were most often encountered actively moving with-
in the wetland. Juveniles selected foraging microhabitats that differed from random and
consisted of mud substrate with sparse vegetation. Adults occupied a greater variety of mi-
crohabitats that did not differ from random sites. In addition, free-ranging cottonmouths
exhibited ontogenetic shifts in diet: juveniles consumed mostly salamanders, while adults
ate a greater variety of prey, including other snakes and birds.
Survivorship was estimated by Rose et al. (2010) for the Northern Cottonmouths in
central Texas. The annual probability of survival at Honey Creek, Comal County, Texas, a
spring-fed stream flowing 3.2 km to its confluence with the Guadalupe River, was 0.81 and
was consistent with estimates for similar species of snakes.
Siegel et al. (2009) found that males and females store sperm, allowing for the disso-
ciation of reproductive event timing between the sexes. Thus, the only reproductive event
coordinated between the sexes is copulation, which is likely to occur in the fall and the
spring. In females, the atrophy and activity of the reproductive organs vary concurrently
with vitellogenesis and the mating seasons. In males, spermatogenesis peaks in the summer,
independent of the mating season.

Asian Bamboo Vipers

Craspedocephalus occurs in Asia from the Indian Subcontinent Sri Lanka, Thailand,
and the Sunda Shelf. About 14 species are recognized. Most species in the genus are relative-
ly small and arboreal, with prehensile tails. Often they are green in color, but some species
also have yellow, black, orange, red, or gold markings. One
species shows unexpected aquatic foraging behavior and is
highly polymorphic in coloration.

Malabar Bamboo Pit Viper

Craspedocephalus malabaricus (Jerdon, 1854)
Distribution and Habitat. Endemic to the Western
Figure 7.5. The distribution of
Craspedocephalus malabaricus. Ghats along southern and western India at 600–2,000 m
ASL in elevation. The type locality is the Western Ghats of
southwestern India. It inhabits riparian forests and inhabits
hill streams and torrents. The forests are dense wet rainforests, evergreen forests and decid-
uous forests, where it may be found on the ground, on rocks in stream beds, on low vege-

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Aquatic Snakes

Figure 7.6 Craspedocephalus malabaricus.

Photography by (A) Vaclav Gvozdik; (B,D) © B R. Ansil. (C) Dilshad Kanniyan.

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 7. Aquatic Vipers

tation, or in shrubs. Recently it was found in the Korba District of Chhattisgarh. The species
prefer semi-arboreal and arboreal habitats facilitated by their prehensile tail but sometimes
exploit the habitats on the ground (Fig. 7.5). Note that this snake was long placed in the genus
Trimeresurus.
Natural History. Riparian habitats are frequently used (Whitaker and Captain 2002; Ga-
nesh et al. 2010). This is a nocturnal species. During the day, they are sometimes observed
basking. They forage for food in the evening, possibly to avoid predators. Craspedocephalus
malabaricus are semi-arboreal and are often detected in the lower canopy, shrub layer, and
ground level. Prey includes small mammals, lizards, frogs, toads, birds, and their eggs. Adults
prefer rodents, whereas juveniles feed on small frogs and lizards. The tail’s white tip and prey
choice suggest the young and juveniles use caudal luring. Reproductive data for this species
is lacking. Wall (1919) reported that the breeding occurs from February to October. Sagar
(2016) said the presence of six juveniles in August and September, indicate that breeding at
Agumbe is well within the season.
Diving behavior was reported by Bhasare and Pelling (2015). They observed the tropical
moist evergreen forest of Agumbe, which is in the central Western Ghats of Karnataka State,
India. The region receives heavy annual rainfall (7,500 to 10,000 mm) during the monsoon
from June to September.
Swimming at a slow pace, the snake was observed to halt in mid-stream (approximate
depth 35–45 cm) and was followed by a short backward jolt of the leading half of the body,
causing the snake to completely submerge 8–10 cm below the surface. After the backward
jolt, the snake remained underwater for 5-6 seconds, of which, for approximately 3-4 sec-
onds, it was completely motionless. Resurfacing involved a slight forward progression during
which only the head broke the surface for about 1-2 seconds before repeating the backward
jolt action. The snake repeated this action three more times, remaining submerged for 5-6
seconds each time. After surfacing for a fourth time, the snake swam a further three meters
downstream to a submerged horizontal branch. It then wrapped its tail around the stem and
coiled its body in a typical ambush position, fully submerged approximately 5 cm under the
surface, with its head facing upwards. Underwater tongue flicks with the occasional slow
sideward movement of the head were observed. The snake remained in this position without
any attempt of surfacing to breathe for approximately 20 minutes and then emerged in a slow
and controlled manner with tongue flicking, leaving the body submerged and coiled. The
snake was not visibly gasping or heavily breathing on surfacing.
Another adult observed by the same authors had a total length of approximately 35 – 40
cm. It was in an open concrete tank (80 cm wide, 250 cm long, and 45 cm deep) at Agumbe
Rainforest Research Station. During the monsoon season, the tank is a breeding site for the
Malabar Gliding Frogs, Rhacophorus malabaricus, prey for C malabaricus. Initially, the
snake was observed floating, partially submerged, at the surface of the 30 cm deep water. The
snake remained motionless with its head under the surface, pointing downwards and floating
in an unusual sideward position due to an inflated body. After one minute, the snake surfaced
and swam a little distance. Once stopped, the snake retracted its head and anterior body and
submerged again. The snake repeated the action of partial submergence four more times for
approximately one minute during each submergence event within a 15-minute observation
period before it ceased due to heavy rain.
The drag created by the large head of a viper was proposed by Young (1990) as the reason
why there are no aquatic vipers. However, at least two of the three aquatic vipers discussed
here do fine catching prey in the water.

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Aquatic Snakes

8. The Homalopsidae
Justin M. Bernstein & John C. Murphy

The snakes of the family Homalopsidae (mud snakes) are indeed a diverse (and often
odd-looking) group of primarily aquatic snakes. Perhaps one of the most poorly known,
aquatic, caenophidian groups regarding their evolution and natural history, many of the
species in this family remain to be investigated, and speculations of their diets, behavior,
reproduction, physiology, and biogeography are inferred from what is known about conge-
ners or other homalopsids. With advances in DNA acquisition methods/technologies (i.e.,
high-throughput sequencing) and software aimed at biological inference, many answers to
the questions we have about homalopsid evolution and diversity will likely come to light
soon.
Homalopsid snakes have a family-wide distribution throughout the Old World (Figure
8.1). The Indus River Delta in Pakistan represents the westernmost range limit for Homalop-
sidae. Eastward, they are distributed throughout much of South Asia, throughout mainland
and maritime Southeast Asia, and reaching New Guinea and northern Australia at the eastern
margin of their range. Their widespread distribution and the fact that they comprise a large
part of the biomass in aquatic ecosystems of Southeast Asia (Murphy, 2007a) are evidence of
their evolutionary success. There are currently 56 described species of homalopsids, distrib-
uted across 29 genera, with new species continuously and recently being described from un-
derexplored regions (e.g., Quah et al., 2017; Kohler et al., 2021, Murphy and Voris, 2020). The
greatest species richness of homalopsids are found in mainland Southeast Asia (excluding the
totality of Indonesian islands throughout the Indonesian Archipelago). While there has yet to
be any synapomorphies discovered for Homalopsidae, a suite of morphological characters is
found amongst most representatives of the family, many of which are characteristic of aquatic
lifestyles: hypapophyses present along the length of the vertebral column; hemipenes forked
with distal end finely calyculate, with shallow cups (spines present and variable); tracheal
lung; crescent-shaped valvular nostrils; shallow rostral notch; elliptical pupils; subcaudals
and cloacal plate divided; and viviparous with a placenta-like connection to the female’s cir-
culatory system (Murphy and Voris, 2014). Although these characters are not confirmed for
all species (as many are known from one or few specimens), the family has been consistently
recovered as monophyletic in studies utilizing molecular datasets, with fanged and fangless
homalopsids as sister groups (Alfaro et al., 2008; Murphy et al., 2011; Burbrink et al., 2020;
Bernstein et al., 2021).

Natural History and Diversity

The diversity of homalopsid snakes is emphasized by their disparate ecologies, diets,
morphologies, behaviors, and habitat preferences. The current species richness of Homalop-
sidae is 56 species, amongst 29 genera. The morphological disparity of homalopsids is seen in
their color patterns, body sizes, and head morphology. Species range from stout, heavy-bod-

Previous page. The Puff-faced Watersnake, Homalopsis buccata (Homalopsidae) traverses a shallow
water habitat. The snake was photographed at Langkawi, Malaysia, Photography by Kai Kolodziej.

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Aquatic Snakes

ied snakes (e.g., Subsessor bocourti) to slender-bodied Enhydris enyhydris. The differences in
body forms are accompanied by different head shapes that contribute to dietary specializa-
tion (Jayne et al., 2018). The most drastic example of morphological disparity in this family
is Erpeton tentaculatum, the tentacled snake, named after its rostral appendages, that are
unique to this taxon. Intraspecific variation is also prevalent in many species, particularly
notable in Fordonia leucobalia, which contains forms that are only black and white, as well as
morphs that contain any combination of rust, red, and orange, or any combination of these.
The greatest morphological dichotomy of homalopsids, though, is their dentition. Mud
snakes are split into two groups: fangless homalopsids (aglyphous; three genera: Brachyor-
rhos, Calamophis, Karnsophis), and the rear-fanged group (opisthoglyphous; all other gen-
era [Taub, 1967; Gyi, 1970]). It is likely that all the rear-fanged members use venom that
help subdue their prey (Mori, 1998). While bites by rear-fanged homalopsids may produce
localized effects (e.g., swelling, burning), reports are seldomly medically significant (Mur-
phy, 2007a). It is worth nothing that bites from most species have never been documented;
some reports with locals of the Andaman Islands state that bites inflicted by Cantoria viola-
cea have resulted in death or unconsciousness (though, these reports may have confused C.
violacea with the venomous elapid Bungarus andamanensis;
(Ghodke and Andrews, 2000). Interestingly, the venom of
Cerberus contains a family of venom proteins that have yet
to be discovered in other venomous snakes (OmPraba et al.,
2010), and rear-fanged homalopsids may possess a venom
protein family (veficolins) that is unique to Homalopsidae.
More studies are needed to identify the extent of variation
Figure 8.1. The distribution of
the Homalopsidae in this group’s venom, and the respective physiological ef-
fects from bites from each species. As with all snakes, han-
dling species without knowing their exact identity and the
potency of their venom is advised against.
Mud snakes have successfully adapted to an aquatic lifestyle and are well known for
dominating the aquatic habitats they are a part of. Unlike most of the species in this fam-
ily, the fangless homalopsids have adapted to a terrestrial/semi-fossorial lifestyle (reports
are mostly limited to Brachyorrhos; Murphy et al., 2012a). The rear-fanged clade inhabits a
variety of aquatic habitats. Some species are found primarily in freshwater systems such as
rivers, estuaries, lakes (Cerberus, Homalopsis, Enhydris) and freshwater swamps (Gyiophis,
Heurnia, Raclitia). Peat swamps are inhabited by Homalopsis buccata, Phytolopsis punctata,
Homalophis doriae, and Enhydris innominata, suggesting that these species may be able to
tolerate the high acidity of the substrate. Several genera and species are well-adapted to
the brackish waters of coastal mangrove systems and tidal flats (e.g., Cerberus schneiderii,
Bitia hydroides, Cantoria violacea, Gerarda prevostiana). Many closely related species have
partitioned into different habitats, such as the freshwater Myrrophis chinensis and saltwater
tolerant Myrrophis bennettii, the latter of which also rests on shrubs and bushes, which is
atypical amongst homalopsids (Murphy, 2007a). Many species that inhabit brackish waters
have also been found swimming in open waters near the coast (Cerberus australis [King-
horn, 1929], Cerberus rynchops [Acharji and Mukherjee, 1996], Cerberus dunsoni [Crombie
and Pregill, 1999], Fordonia leucobalia [O’Shea, 1986]). It is likely that many species, es-
pecially those possessing salt glands (Dunson and Dunson, 1979) are able to tolerate both
freshwater and saltwater environments for at least a limited amount of time, as many species
are not solely found in one particular type of habitat. The differences in habitat between the
fangless and rear-fanged homalopsids are also reflected in their diet. While Calamophis and

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Karnsophis remain largely understudied due to low numbers of encountered and collected
individuals, Brachyorrhos are vermivorous, primarily feeding on earthworms (Murphy et
al., 2012a; Murphy unpublished; Sanders unpublished).
Karnsophis has been confirmed to eat frogs, but it is un-
certain if they have a frog-specialized diet or are gener-
alists (Murphy and Voris, 2013). The prey consumed by
the rear-fanged homalopsids is much more diverse, be-
ing comprised of a variety of fish (Anabantidae, Channi-
dae, Clariidae, Cyprinidae, Nandidae, Osphronemidae,
Synbranchidae), frogs (Ranidae, Rhacophoridae), and
aquatic arthropods (Thalasinnidae [mud lobsters], Pa-
laemonidae and Alpheidae [shrimp], Varunidae [crabs])
(Shine, 1991; Khan, 2002; Murphy, 2007a; Brooks et
al., 2009; Murphy and Voris, 2013; Pongcharoen et al.,
2016; Ghodke et al., 2018). A clear shift in diet during
the evolution of homalopsids can be seen in Fordonia
leucobalia, Gerarda prevostiana, and Cantoria violacea,
which form a clade. These snakes primarily feed on Figure 8.2. The tentacles of Erpeton
make it one of the most distinctive
crustaceans, particularly hard-shelled crabs (Fordonia)
snakes. Note the free end of the
and freshly molted crabs and alpheids (Gerarda and scales point anteriorly. Photogra-
Cantoria) (Murphy, 2007a; Jayne et al., 2002). Homalop- phy by Ken Catania.
sids have a variety of predators; some of these include:
predatory fish and sharks (Carcharhinus cautus) birds of
prey (Haliaeetus leucogaster, Haliastur indus, Milvus migrans), crocodiles (Crocodylus poro-
sus), lizards (Varanus indicus), other snakes (Bungarus spp. Cylindrophis spp., Naja naja),
and mammals (Bandicota indica, Homo sapiens) (Mao, 1970; Lyle and Timms, 1987; Shine,
1991; Voris and Jeffries, 1995; Murphy 2007a).
Homalopsids display a variety of behaviors that are
well known amongst snakes. These include swimming
in open waters, hiding amongst root entanglements, uti-
lizing burrows of other animals, and sidewinding when
escaping predators (Murphy, 2007a; personal observation
by JMB [Cerberus microlepis]). However, a few species
have been widely recognized for behaviors that have yet
to be identified in any of the 3,800+ species of snakes. Figure 8.3. Cerberus has well de-
The ‘crustacean-eating’ clade’s (Fordonia, Gerarda, Can- veloped, gooved rear-fangs. Here
the functional fang is adajcent to
toria) shift in diet to lobsters, crabs, and shrimp also
a replacement fang. Photography
shows a shift in how they feed. Fordonia leucobalia uses a by JCM
specialized closed-mouth strike to attack and its chin to
pin down its crustacean prey (Jayne et al., 2018). Both F.
leucobalia and G. prevostiana rip their prey’s legs off, engulfing them separately, allowing
them to eat prey larger than would normally be possible. Gerarda prevostiana also tears the
carapace off of the crab and uses body coiling (Jayne et al., 2018). Erpeton tentaculatum,
a highly aquatic homalopsid snake, distantly related to the aforementioned clade, uses its
unique rostral appendages for hunting. These appendages are mechanoreceptors which are
innervated by the trigeminal nerve and increase the snake’s sensitivity to water movement
(Catania et al., 2010). This allows the snake to wait in a rigid J-shaped posture in the water,
flick its tail to startle unsuspecting fish prey into a C-start escape response towards the

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snake’s mouth, and meet it to ultimately eat its prey (Smith et al., 2002; Catania, 2009; Catania
et al., 2010). And finally, an observation suggests the poorly known Bornean species Hom-
alophis gyii, changes color from dark reddish-brown to a complete white color within a few
minutes. The role of metachrosis in snakes is under studied. It may be a defense mechanism
or stress response once the specimen was placed into a bucket (Auliya, 2003).

Taxonomic History
The nomenclatural history of Homalopsidae has been
far from stable for centuries. The first described species of
the family was Homalopsis buccata, and was actually de-
scribed on three separate accounts before the second species
of homalopsid snake was described. Homalopsis buccata was
described by Carl Linnaeus (1707–1778), twice in the same
work (Linnaeus, 1758): first as Coluber buccata from India
(specimen lost), and again as Coluber monilis from Ameri-
Figure 8.4 Cerberus in a man- ca. Then, in 1789, Johann Friedrich Gmelin (1748–1804), a
grove swamp in Singapore. German naturalist, used an illustration from Dutch zool-
Photography by Daryl R. Karns.
ogist Albertus Seba’s Locupletissimi Rerum Naturalium
Thesauri as the iconotype for Coluber subalbidus (Gmelin,
1789); these three Coluber taxa are all now known to be H.
buccata (Murphy, 2007a). It was not until 10 years later that the second species, Hydrus en-
hydris (=Enhydris enhydris) was described by Schneider (1799).
Further contributions to homalopsids took place in the early 19th century by Heinrich
Boie (1794–1827), a German naturalist who worked on the herpetofauna of Java, which ul-
timately remained unpublished (Adler, 1989). Boie’s older brother, Friedrich Boie, published
some of Heinrich Boie’s earlier papers posthumously (Adler, 1989), which included many
new genera and species; one of these still-valid taxa is what is now recognized as Hypsisco-
pus plumbeus (Boie, 1827). Before his death, Heinrich Boie also grouped all aquatic snakes
into the family Hydrophidae, which included homalopsids except for Erpeton tentaculatum
(Murphy, 2007a). Although aquatic snakes are now, without
question, recognized as distinct evolutionary lineages in
part due to broadly-sampled systematic studies (e.g., Pyron
et al., 2013; Burbrink et al., 2020; Bernstein et al., 2021), the
Hydrophidae was the earliest grouping of aquatic snakes as
a distinct taxonomic group (Murphy, 2007a). Later, John E.
Gray (1800–1875), British zoologist and founder of the zo-
Figure 8.5. Cantoria violacea is an
ological collections of the Natural History Museum (=Brit-
exceptionally slender and elon-
gated homalopsid. Photography ish Museum, London), placed all genera of aquatic snakes
Daryl R. Karns. in the family Hydridae, which included 14 homalopsids
described by himself, eight of which are currently valid as
the following: Bitia hydroides, Cerberus australis, Homalopsis
hardwickii, Myron richardsonii, Myrrophis bennettii, Myrrophis chinensis, Phytolopsis puncta-
ta, and Raclitia indica (Gray, 1849). Continuing with the taxonomic flux of aquatic snakes,
homalopsids were placed in the group ‘Platyrhiniens’ by French zoologists Constant Duméril
(1774–1860), Gabriel Bibron (1805–1848), and Auguste Duméril (1812–1870) in their sem-
inal work Erpétologie Générale (Duméril et al., 1854); this work includes the description of
new homalopsid snakes, two of which are still valid and recognized: Dieurostus dussumieri

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 8. The Homalopsidae

and Sumatranus albomaculatus.

The recognition of Homalopsidae as a higher taxonomic group of snakes distinct from
other aquatic lineages did not happen until Charles L. Bonaparte (1803–1857), French zo-
ologist, classified them under the “Homalopsina,” a subfamily
of Dipsadidae (Bonaparte, 1845). Other than the higher-lev-
el name (likely derived from the genus Homalopsis by Kuhl
and Hasselt [1822]), no list of characters or descriptions that
grouped homalopsids in the Homalopsina was given. Because
of this inconspicuous mentioning of homalopsids as a distinct
group, credit towards naming homalopsids as a family of snakes
is often given to Giorgio Jan (1791–1866), the infamous Aus- Figure 8.6 A Gerarda near
trian botanist and zoologist, most known for the iconic snake the entrance to a crab bur-
illustrations in his Iconographie Générale des Ophidiens (Jan, row in a Singapore man-
1864). Jan (1863) used the name ‘Homalopsinae’ as a subfamily grove. Photography by Dar-
yl R. Karns
of Potamophilidae (Jan, 1863: p. 67), a family that contained
snake representatives of what we now recognize as homalop-
sids, natricids, and dipsadids. Because of this, Jan’s 1863 paper has been repeatedly cited
as the authority on the family (e.g., Boulenger, 1896; Taylor, 1965; Gyi, 1970). Although
many credit Jan (1863) as resurrecting the name, English physician and amateur herpe-
tologist Malcom A. Smith (1875–1958) recognized the ichthyologist and herpetologist Al-
bert Günther’s 1864 work Reptiles of British India as the authority on Homalopsidae; this
being because Günther (1864) elevated the Homalopsinae to family-level status based on
the arrangement of head plates compared to other colubroid snakes. Still-valid homalopsid
genera in Gunther’s (1864) work include: Bitia, Cantoria, Cerberus, Enhydris, Ferania, For-
donia, Erpeton, Homalopsis, Hypsiscopus, and Myrrophis. While
this is seemingly the first publication to specifically refer to
‘Homalopsidae’ in detail, Jan had a publication, that was appar-
ently forgotten about, titled uber die familie Homalopsidae
(Jan, 1961; Murphy, 2007a). Despite these family or subfam-
ily mentions of the group, Bonaparte (1845) has been cited as
the authority in a few widely recognized herpetological works,
such as Underwood (1967) and David and Ineich (1845).
The taxonomic contributions to Homalopsidae continued
by some of these and other naturalists, especially in regard to
close examinations of morphology and describing new species.
Edward D. Cope (1840–1897), one of America’s most prolific
herpetologists, tried to group homalopsids using vertebral and
Figure 8.7. Homalopsis was
dentition characters, leading to a diverse ‘Homalopsinae’ of 31
considered to be monotypic
genera containing currently-recognized homalopsids, but also but it is composed of sever-
dipsadids, natricids, and pseudaspidids (Cope, 1886). Belgian al species. Photography by
taxonomic herpetologist George A. Boulenger (1858–1937) Daryl R. Karns
followed Günther’s (1864) grouping of homalopsids, and added
Gerarda and Myron to the group, described Hypsirhina (=Hyp-
siscopus) matannensis, synonymized several names, and ultimately reduced Homalopsidae
to subfamily status (Homalopsinae; Boulenger 1890, 1893). The recognition of homalop-
sids as a subfamily of Colubridae was continued by Smith (1943) while revising Boulenger’s
Fauna of British India. Smith (1943) also provided traits that are shared by homalopsids:
enlarged grooved fangs on the posterior of the maxillary; valvular nares; small, dorsolater-

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Aquatic Snakes

al eyes; the tendency to have fragmented head shields; rostral

scale with shallow notch; similar hemipene structure amongst
species; hypapophyses throughout the vertebral column; a
highly aquatic lifestyle; viviparous reproduction; and a distri-
bution restricted to South Asia, Southeast Asia, and northern
Australasia. Although these groupings are shared by a major-
ity of homalopsids, none of these represent synapomorphies
of Homalopsidae (Zaher, 1999), and some of these traits are
now known to be lacking in some representatives (such as the
fangless, semi-fossorial homalopsids [e.g., Brachyorrhos]).
Perhaps one of the most notable taxonomic works on
Homalopsidae was that done by Ko Ko Gyi, a Burmese her-
petologist at Rangoon University (now the University of Yan-
gon; Myanmar). Gyi worked on homalopsid snakes, in the
Division of Herpetology of the Natural History Museum at
the University of Kansas (Kansas, USA), during a three-year
fellowship from the United Nations Educational, Scientific
and Cultural Organization. Gyi’s work included a revision of
Figure 8.8. The illustration of the entire (back then subfamily) Homalopsinae (Gyi, 1970).
Cerberus rynchops from Rus-
In this revision, Gyi (1970) recorded scalation, color pattern
sells 1796 volume, An Account
of Indian Serpents, Collected characters, and body measure-
from the Coast of Coroman- ments on 983 specimens (1038
del.. Russell used the common total in the study), ultimately
Indian name for this snake - recognizing 34 species amongst
Bockdam. 10 genera (22 species of which
were placed in Enhydris). Gyi’s
(1970) work also provided dis-
tribution maps, specimen photographs, a key to genera and
species, natural history notes, and biogeographic remarks and
hypotheses. While taxonomic rearrangements have been pro-
posed or implemented, especially regarding Enhydris (e.g., Saint
Girons, 1971), a family-wide review and revision of Homalopsi-
dae did not occur until 37 years later.
One of us (JCM) and Harold K. Voris, researchers at the
Field Museum of Natural History (Chicago, Illinois, USA) have
provided the most recent revisions to homalopsid taxonomy at a
familial level. Murphy (2007a) published a book on mud snakes,
which by this point is recognized as a family due to several stud-
ies supporting their evolutionary distinctiveness using molecu-
lar data (e.g., Voris et al., 2002). Using detailed morphological
examinations of homalopsids, Murphy (2007a) recognized 39
species of homalopsid snakes in 12 genera (though, two genera
Figure 8.9. The holotype of
and three species were considered problematic, to be revised lat- Homalophis gyii. Names in
er [Murphy and Voris, 2014]). Of these, 23 species were placed honor of the Burmese her-
in Enhydris. This book provided updated geographic distribu- petologist who revised the
tions, taxonomy, genera and species keys and illustrations, nat- homalopsid snakes in the
ural history information from the literature and personal obser- 1960s. Art work by Sara
vations, and a synopsis on homalopsid taxonomic history. With Drasner.

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 8. The Homalopsidae

the increasing ease of acquiring molecular data, it was established that several homalopsids,
primarily Enhydris, were not monophyletic (Alfaro et al., 2008); this led to the most recent
revision of homalopsid snakes by Murphy and Voris (2014). Murphy and Voris (2014) rec-
ognizes 53 species (not including an undescribed lineage from Sulawesi; Murphy and Voris,
2014) amongst 28 genera, now with only six species of Enhydris, using morphological cri-
teria. While Murphy and Voris (2014) was the most complete checklist of Homalopsidae,
more species have been described since its publication (e.g., Gyiophis salweenensis [Quah et
al., 2017], Myanophis thanlyinensis [Günther et al., 2021], Brachyorrhos pygmaeus [Murphy
and Voris, 2020]), Hypsiscopus murphyi (Bernstein et al., 2022) raising the current species
richness of the family to 56. “Genomic data from Bernstein et al. (2023) suggests that hom-
alopsids actually diversified in the Oligocene ~26 million years ago, either in Indochina
or Indochina+South Asia, and that many divergence events in the rear-fanged group were
likely due to shifts in paleorivers ~17 million years ago rather than movement into different
aquatic environments. This suggests Pleistocene sea-level fluctuations were not the main
driver of homalopsids diversification, but sea-level changes undoubtedly influences popu-
lation structure at least in the last tens of thousands of years.

Phylogenetic Placement
Homalopsids have long been recognized as caenophidian snakes and hypothesized to
be affiliated with several families. Mud snakes have been considered as closely related, or
part of, colubrids for much of their history (Boulenger 1890, 1893; Gyi, 1970). Before this,
though, homalopsids were considered as a family, transitional between his colubrid group
‘Natricina’ (=Natricidae) and hydrophiines (=Hydrophiidae), particularly noting the ‘tran-
sitional’ appearance of the fang dentition (Nicholson, 1874); it is important to note that
there is no evidence to support this as an evolutionary transition. Underwood (1874) also
suggested that acrochordids were derived from the homalopsids. While still considered as
part of colubrids, Dowling (1974) and Dowling and Duellman (1978) considered the mud
snakes to be a tribe of the natricids (as Natricinae, family Colubridae). While homalopsids
certainly do not produce the complex venom compounds as viperids, Knight and Mindell
(1994) and Underwood (1999) hypothesized a close association between the two groups of
snakes. The increased possibilities of DNA sequencing in the latter parts of the 20th cen-
tury led to many studies confirming the monophyly and distinctiveness of homalopsids as
a family distinct from the Colubridae, Natricide, Viperidae, and Hydrophiinae (Elapidae)
(Voris et al., 2002; Pyron et al., 2013; Figueroa et al., 2016; Burbink et al., 2020). Homalop-
sids have been recovered in these molecular phylogenies as either sister to a Lamprophi-
idae+Elapidae clade (Pyron et al., 2013; Figueroa et al., 2016), or outside the sister rela-
tionship between Elapidae and a group containing Colubridae+Natricidae (Harrington and
Reeder, 2017). This placement of homalopsids amongst the evolutionary tree of Serpentes
seems to be supported using genomic data as well. Having the most homalopsid species
sampled for a phylogenomics dataset, Burbrink et al. (2020) recovered Homalopsidae as
the most closely related family to the Elapoidea and Colubroidea, with all families sampled.

Biogeography
While the taxonomy and phylogenetic placement of Homalopsidae has stabilized in re-
cent years (e.g., Murphy, 2007a; Murphy and Voris, 2014; Burbrink et al., 2020), the biogeo-

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Aquatic Snakes

graphic origins of the group and the evolutionary processes that have led to the subsequent
diversification of lineages is far less understood. Mud snakes are distributed from Pakistan,
eastward into South Asia, Southeast Asia, and reach their eastern limit in northern Australia
and New Guinea. The geological history that spans these regions is extremely complex, re-
sulting from millions of years of plate collisions, tectonic uplift, river catchment events, and
fluctuating sea levels (Inam et al., 2007; Polhemus, 2007; Hall, 2009). Some of these events
happened simultaneously in short time spans, while other events were separated by millions
of years; some geological phenomena have been one-time events, and others repetitively over
the course of several epochs.
The geographic origins of homalopsid snakes are suspected to be in mainland Southeast
Asia (Murphy, 2007a), as 48% of homalopsid species are distributed throughout Indochina
(and adjacent China), with the rest known from isolated regions or islands east or west of
Indochina. Few family-wide systematic studies exist for this
group, but molecular data suggests that the fangless hom-
alopsids (Brachyorrhos [Karnsophis and Calamophis not in-
cluded in studies]) diverged from the rear-fanged clade in the
mid-Eocene (43.7–47.9 Ma; Bernstein et al., 2021). Other esti-
mates that do not include the fangless clade range from 22 Ma
Figure 8.10. Until 2014, the ge-
nus Enhydris contained most (Alfaro et al., 2008 [multilocus data]) to ~50 Ma (Burbrink
species of homalopsids despite et al., 2020 [genomic data]). It
evidence that it wss not mono- is likely that the fluctuating sea
phyletic. Above is an adult fe- levels that facilitated land con-
male Subsessor bocourti (left) nections during sea level mini-
and an adult female Enhydris ma in Sundaland lead to many
enhydris (right). both consid- of the homalopsid (and other
ered members of the genus En- faunal and floral) distributions
hydris until 2014. Photography
seen today (Woodruff, 2010;
by JCM
Bernstein et al., 2021). Multilo-
cus data suggests the possibility
of extinction of lineages throughout homalopsid evolution and
increased diversification of rear-fanged lineages ~10 Ma (Bern-
stein et al., 2021). However, studies utilizing genome-scale data
on both rear-fanged and fangless taxa with rigorous biogeo-
graphic analyses are necessary to confirm many of these find-
ings and hypotheses. ‘Genomic data (Bernstein et al., 2023a)
supports that homalopsids likely diversified in the Oligocene Figure 8.11. An Enhydris
~26 million years ago in Indochina or Indochina+South Asia, enhydris eating a Siamese
and diversified in the last east and west from there in the last Fighting Fish of the genus
~20 million years.’ Betta. Photography by JCM
The evolution of genera and species has been primarily
focused on Cerberus and Enhydris (sensu Murphy and Voris,
2014). The six species of Enhydris comprise a large proportion
of the vertebrate biomass in mainland Southeast Asia’s low-elevation freshwater systems
(Murphy, 2007a). While in-depth studies with more locus and specimen sampling are need-
ed, their reliance on aquatic systems makes it likely that river length, topography (drainage
basins, plateaus; Lukoschek et al., 2011), and sea level fluctuations (Woodruff, 2010) all influ-
enced the diversification of this genus. Molecular data suggests that populations of Enhydris
subtaeniata have been significantly structured by the lifting of the Khorat Plateau during

127
 8. The Homalopsidae

the Quaternary, and the associated river catchment event of the Mekong River (Hutchi-
son, 1989; Rainboth, 1996; Lukoschek et al., 2011). It is likely that this also had a similar
impact on the population structure of Hypsiscopus plumbeus (Bernstein et al., 2021). Of
particular interest would be why the distribution of Enhydris enhydris is widespread from
India, mainland Southeast Asia, and the islands of Sundaland compared to its conspecifics,
which have narrow distributions in Thailand, Cambodia, Vietnam, and Laos.
The brackish water Cerberus have one of the widest distributions of all snakes, which
is nearly as widespread as Homalopsidae itself. Specifically, Cerberus schneiderii ranges
from the Andaman sea coast of Thailand, eastward through the Indonesian archipelago.
Phylogenetic methods using molecular data have shown that C. microlepis and C. dunsoni
may actually be isolated, founder populations of C. schneiderii, as both former species are
recovered within the latter (Alfaro et al., 2004; Bernstein et al., 2021). Similar to many
other taxa, the distributions of Cerberus schneiderii and its congeners have likely been the
result of Pleistocene sea level fluctuations in the past few hundreds of thousands of years.
Similarly, land bridges during sea level minima have likely led to migration of Pseudofer-
ania polylepis and Myron richardsonii populations between New Guinea and Australia.
Genomic data (Bernstein et al., 2023a) supports that homalopsids likely diversified in the
Oligocene ~37 mya in Indochina and/or Sundaland.

Conservation
Unfortunately, despite their ecological importance, the conservation status of most
homalopsids snakes have either not been assessed for extinction risk by the International
Union for Conservation of Nature (IUCN) or population trends are unknown due to data
deficiency (IUCN, 2020). Out of the 56 described species, Homalopsidae has the following
IUCN statistics:
A large proportion of homalopsid species have not been assessed in over 10 years for
IUCN extinction status or are being assessed based on old taxonomy. For example, Cerbe-
rus populations in Wallacea and the Greater Sunda Islands are erroneously still recognized
as C. rynchops, rather than C. schneiderii. Ironically, C. rynchops is listed as a CITES appen-
dix III species (India), despite being one of the most abundant homalopsids, and snakes,
in Asia (Jayne et al., 1998; Murphy, 2007a). While only Cerberus microlepis (EN), Gyiophis
vorisi (EN), and Enhydris longicauda (VU) have extinction risks as endangered or vul-
nerable, almost all other species have either yet to be assessed, are data deficient, or have
not been reevaluated. Thus, any indication as to whether populations are increasing, de-
creasing, or are stable is
only available for ~9%
of species. Species that
are poorly known/en-
countered, are endemic,
or have extremely limit-
ed distributions, such
as Enhydris chanardi,
Mintonophis pakistan-
icus, Calamophis spp.,
Brachyorrhos spp., as
well as island popu-
lations of widespread

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Aquatic Snakes

species (e.g., Cerberus schneiderii), should be focused on in

future assessments.
The primary threats to homalopsid snakes are those
that are considered detrimental for most other species: hab-
itat destruction/fragmentation, pollution, and overexploita-
tion/collecting for sale in the pet trade or in local markets
Figure 8.12. Enhydsris subtae- for their meat and skin (Murphy, 2007a). Mud snakes are
niata was long confused with also commonly killed as bycatch in fishing nets. Their abun-
Enhydris enhydris until it was dance in aquatic systems likely makes them important as
removed from the synonomy aquatic indicators of ecosystem health, but fishing and hunt-
of E. enhydris. Photography by ing operations threaten many populations. In particular, the
JCM. populations of homalopsids at Tonlé Sap Lake in Cambodia
are threatened, with almost 4 million snakes collected per
year (Brooks et al., 2007). This estimate includes Erpeton
tentaculatum, Subsessor bocourti, Hypsiscopus plumbeus, Homalopsis buccata, Enhydris lon-
gicauda, and Enhydris enhydris, with the latter species comprising ~70% of all specimens
(Brooks et al., 2007). The catching of ‘low-value’ water snakes, such as homalopsids, benefit
the economy when there are declines in fish catchment (Brooks et al., 2008). Finding balanc-
es between snake hunting and fishing, along with a better understanding of the diversity of
lineages in this family and their associated taxonomy, is much needed to ascertain extinction
risks of homalopsids and to create conservation management plans.

Fangless Species

Seram Short-tailed Snake

Brachyorrhos albus (Linnaeus, 1758)
Distribution and Habitat. Currently known from Indo-
nesia, Maluku Province, specifically Seram and its satellite
islands (Ambon, Haruku, Nusa Laut, Saparua). It has also
been reported from the Banda Islands (Boettger, 1985). A
population may exist on the Indonesian Island of Pulau Bisa
(O’Shea, 2018), but molecular confirmation of this specimen
is required to designate this as B. albus (Murphy and Vo-
Figure 8.13. Distribution of ris, 2014). Populations may inhabit other islands (Banda,
Brachyorrhos albus. Batjan, Aru Islands) that have been underexplored or lack
surveys. Type locality: “Indiis” (Indonesia, Ambon). (Figure
8.14)
Identification. Distinct preocular scale usually present (also present in B. raffrayi and B.
wallacei); seven upper labials, fourth (rarely 3+4) in orbit, sixth is tallest; dorsal scales more
lanceolate (elongated, taper posteriorly) than in congeners; usually single temporal scale con-
tacting postoculars in Ambon population; two primary temporals in some Seram individu-
als. Size. Males: SVL: 337–472 mm; Tail: 32–61 mm. Females: SVL: 343–443 mm; Tail: 33–41
mm.
Natural History. Extraordinarily little is known about the natural history, evolution, and
habits of Brachyorrhos spp, and most of what is known about the genus comes from observa-

129
 8. The Homalopsidae

Figure 8.14. to the left an adult Brachyorrhos albus. To the right

are near full term from a B. albus from Ceram. Photography by
JCM.
tions of B. albus. Brachyorrhos are an early diverging lineage of fangless homalopsids that are
primarily terrestrial/semi-fossorial. There have been published reports of Brachyorrhos occa-
sionally inhabiting arboreal habitats as well (Edgar and Lilley, 1993). Brachyorrhos albus are
vermivorous, primarily feeding on earthworms (Murphy et al., 2012a; Murphy unpublished;
Sanders unpublished). They are reported to be secretive, staying hidden beneath stones and
logs, and are active at dusk to hunt prey (Kopstein, 1926). Populations on Seram were found
in lowland secondary forests, but also in plantations, gardens, and have been found in hu-
man-populated areas at all altitudes (Murphy et al., 2012a). Although its reproduction is
hardly understood, B. albus is viviparous based on one Seram female that contained four
near-term embryos (Murphy et al., 2012a). Molecular data supports Brachyorrhos as the sis-
ter group to the rear-fanged homalopsids (Murphy et al., 2011). Skull and scale morphology
suggest that Brachyorrhos and the other fangless homalopsids, Calamophis and Karnophis,
share a more recent ancestry than either of them does with the rear-fanged species (Murphy
and Voris, 2013). Molecular data also supports a very close relationship between B. albus and
B. raffrayi (Bernstein et al., 2021); further investigation is needed to see if these two species
are populations are conspecific. The natural history of other Brachyorrhos species are largely
unknown.

Buru Short-tailed Snake

Brachyorrhos gastrotaenius (Bleeker, 1860)
Distribution and Habitat. Type locality: “Amboina” (= Am-
Figure 8.15. The distribu- bon), in error. Buru – Fakal. Only known from Indonesia, Malu-
tion of Brachyorrhos gastro- ku Province, Buru Island (Figure 8.15.)
taenius. Identification. Six upper labials, fourth (rarely 3+4) in orbit,
fifth is tallest; mid-ventral stripe often present. This combination

Figure 8.16. Brachyorrhos

gastrotaenius. Photography
by JCM.

130
Aquatic Snakes

of characters separates this species from all other members of the genus. Size. Males: SVL:
342–415 mm; Tail: 42–73 mm. Females: SVL: 338–595 mm; Tail: 30–48 mm (Figure 8.16).
Natural History. See Brachyorrhos albus.

Pygmy Short-tailed Snake

Brachyorrhos pygmaeus Murphy & Voris, 2020
Distribution and Habitat. Type locality: Indonesia, Se-
ram Island. Only known from the Type locality:
Figure 8.17. Distribution of Brachy-
orrhos pygmaeus Identification. A small Brachyorrhos with a rounded
snout, a loreal scale; one preocular scale; upper labials six or
seven, the fourth upper labial bordering the orbit; the second
and third upper labials contact loreal; six lower labials; 143
ventral scales; 22 subcaudal scales. Size. Males: Data Defi-
cient. Females: SVL: 115 mm; Tail: 13 mm.
Natural History. See Brachyorrhos albus.
Figure 8.18. Brachyorrhos pyg-
maeus. Artist credit Nathalie
Aall.

Ternate Short-tailed Snake

Brachyorrhos raffrayi (Sauvage, 1879)
Distribution and Habitat. Only known from Indone-
sia, Maluku Province, Ternate. Type locality: Indonesia,
Ternate.
Identification. Six upper labials, the third, rarely 3+4,
enter the orbit, tallest upper labial is third; very pointed
snout. A preocular is present in about 35% of the speci-
mens examined, making it possible to confuse this species
Figuire 8.19. The distribution of with B. albus, which often have the preocular scale. Size.
Brachyorrhos raffrayi. Males: SVL: 453–660 mm; Tail: 56–82 mm. Females: SVL:
394–462 mm; Tail: 29–51 mm.
Natural History. See Brachyorrhos albus.

Halmahera Short-tailed Snake

Brachyorrhos wallacei Murphy, et al. 2012
Distribution and Habitat. Only known from Indone-
sia, Maluku Province, Halmahera Type locality: Indonesia,
Halmahera, Jailolo including the island or Morotai (Murphy
and Voris 2020).
Identification. Usually lacks preocular, usually seven
Figure 8.21. The distribution of (rarely six) upper labials, tallest is sixth (rarely fifth). Brachy-
Brachyorrhos walacei. orrhos wallacei is similar to B. raffrayi, which has upper labial
three in the orbit and the third is the tallest. Specimens of B.
wallacei usually have the fourth upper labial (rarely 3+4) in

131
 8. The Homalopsidae

Figure 8.20. Brachyorrhos raffrayi.

Photography by Kate Sanders.
the orbit, and the fifth is the tallest. Size. Males: SVL: 317–395 mm; Tail: 17–54 mm. Females:
SVL: 345–575 mm; Tail: 35–65 mm.
Natural History. See Brachyorrhos albus. Molecular data has shown that B. wallacei is
deeply divergent from B. raffrayi (Murphy et al. 2012a). This was also seen in Bernstein et al.
(2021) using mitochondrial and nuclear loci, with B. wallacei as sister to B. raffrayi+B. albus.
However, these datasets do not include B. gastrotaenius in a phylogenetic framework and are
based on limited molecular datasets. Thus, a thorough investigation with increased molecular
and taxonomic sampling will be needed to elucidate the species boundaries and evolutionary
relationships of this genus.

Figure 8.22. Brachyorrhos

wallacei. Photography by
JCM.

Yapen Island Stout-tailed Snake

Calamophis jobiensis Meyer, 1874
Distribution and Habitat. Only known from New Guinea, apen Island (= Jobi/Japen).
Type locality: Indonesia, Yapen Waropen Regency, Yapen Is-
land, Ansus (about 1°43’2” S, 136°7’57” E). (Figure 8.23)
Identification. Diagnosis based on a translation of the
original description164 ventrals and 10 subcaudals. Upper la-
bials six; third and fourth enter orbit, eight lower labials, one
Figure 8.23. The distribution of preocular, one postocular; pentagonal internasal contacts ros-
Calamophis jobiensis. tral with one side and penetrates between prefrontals; subcau-
dals divided. Size: Measurements not available.

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Aquatic Snakes

Natural History. Like most of the other fangless homalopsids, the known natural history
of all Calamophis is minimal and based off few specimens. Calamophis jobiensis has the least
amount of data attributed to any member of the genus. The holotype of C. jobiensis MTD (=
MTKD; Staatliches Museum für Tierkunde) 1026 was destroyed in the Dresden bombings
of World War II; no measurements were recorded from the holotype’s original description
(Meyer, 1872). It is possible that C. jobiensis has a terrestrial/fossorial lifestyle.

Andian Stout-tailed Snake

Calamophis katesandersae Murphy, 2012
Distribution and Habitat. Only known from Indonesia,
West Papua, Andai. Type locality: Indonesia, West Papua Prov-
ince, Manokwari Regency, Andai (~0°54’58” S, 134°00’25” E).
(Figure 8.11)
Identification. Distinctive laterally compressed body
Figure 8.24 The distribuition (mid-body width 75–79% of body’s height), exceptionally
of Calamophis katesandersae. short tail (3.7–3.8% of snout–vent length [SVL]); frontal pen-
tagonal but almost triangular; five or six upper labials; sev-
en lower labials; low subcaudal count (eight or nine). Size.
Males. Data Deficient, Females. SVL: 157–215 mm; Tail: 6–8
mm.
Natural History. Calamophis katesandersae may have
a more aquatic lifestyle compared to its congeners. This is
based off its morphology: laterally compressed body, nar-
Figure 8.25. Calamophis kate- row ventrals (compared to its congeners), and eyes that are
sandersae. Photography by JCM slightly more dorsolateral (compared to more dorsal in C.
ruuddelangi and C. sharonbrooksae) (Murphy, 2012). More
specimens and through investigation of morphology and natural history observations will be
needed to identify the true habitat preferences and lifestyles of Calamophis spp.

Kebar Stout-tailed Snake

Calamophis ruuddelangi Murphy, 2012
Distribution and Habitat. Known only from Indonesia,
West Papua Province localities of Ambuaki and Kebar Val-
ley. Type locality: Indonesia, West Papua Province, Manok-
wari Regency, Ambuaki in the Tamrau Mountains (~ 0°46’ S,
132°57’ E) West Papua. (Figure 8.26).
Figure 8.26, The distribution
Identification. Gracile cylindrical body, relatively short
of Calamophis ruuddelangi.
tail (12% of SVL); six upper labials, fifth tallest, three and four
shorter, ventrals 143–145, subcaudals 21–23. Size. Males. SVL:
219–232 mm; Tail: 27–29 mm. Females. Data Deficient. (Fig-
ure 8.27).
Natural History. This species may have a more terres-
trial/fossorial lifestyle compared to C. katesandersae. Molec-
ular data (Bernstein et al., 2023) shows that C. ruuddelangi
Figure 8.27. Calamophis (only one specimen in the sampling) is the sister to the genus
ruuddelangi.
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 8. The Homalopsidae

Brachyorrhos. It is likely that Calamophis and Brachyorrhos are sisters but the phylogenetic
placement of Karnsophis will be needed to confirm how the three known fangless genera are
related. However, more specimens and natural history observations are needed to confirm
this.

Arfak Stout-tailed Snake

Calamophis sharonbrooksae Murphy, 2012
Distribution and Habitat. Only known from Indonesia, West Papua Province. The two
type specimens are collected from Manokwari Regency, Mount
Arfak. A female specimen was collected from Atinyu (1°26’04” S,
132°22’58” E), Sorong Selatan Regency, near Lake Danau (O’Shea
and Kaiser, 2016). Atinyu is approximately 180 km west southwest
of the Type locality: Type locality: Indonesia, West Papua, Mount
Arfak (~ 1°05’00” S, 133°58’00” E). (Figure 8.28)
Identification. A robust Calamophis with a cylindrical body,
Figure 8.28. The distribu- tail 9.0–9.6% of the SVL; upper labials three, four and five about
tion of Calamophis sharon- equal in height; the rostral to frontal distance is greater than the
brooksae.
parietal seam. Dorsal scales are uniform brown—no center spot
of dark pigment. All other species have dorsal scales with a dark
central spot and a light outer edge. In C. katesandersae the fifth
upper labial is the tallest; the body is laterally compressed; and
the tail is about 4% of the SVL. Calamophis ruuddelangi has a
gracile, cylindrical body and a tail that is about 12% of the SVL.
Size. Males: SVL: 288–290 mm; Tail: 26–28 mm, Females. SVL:
390 mm; Tail: 18 mm (Figure 8.29).
Natural History. This species may have a more terrestrial/
Figure 8.29. Calamophis sha- fossorial lifestyle compared to C. katesandersae. O’shea and Kai-
ronbrooksae. Photography by ser (2018) mention that Calamophis may not be able to be in
JCM
sympatry with the species-rich, vermivorous elapid genus Tox-
icocalamus, based on the mutual exclusion of both genera in their respective distributions
(O’Shea and Kaiser, 2018). These hypotheses require more thorough investigation with higher
sampling and observations.

Sumatran short-tailed snake

Karnsophis siantaris Murphy & Voris, 2013
Distribution and Habitat. Only known from Indonesia,
Sumatra, Sumatera Utara (North Sumatra Province), Siantar.
However, Bleeker (1860) reported Brachyorrhos albus from
Agam (~0°15’ S, 100°05’ E) and Padang (0°57’ S, 100°21’ E),
both localities from West Sumatra Province. As Brachyorrhos
is currently considered only found in eastern Indonesia, these
accounts from Bleeker (1860) likely K. siantaris. Type locality:
Figure 8.30. The distribution Indonesia, Sumatra, Sumatera Utara, Siantar (~1°58’ N, 99°47’
of Karnsophis siantaris. E). (Figure 8.30).
Identification. Fangless, 19 scale rows not reduced posteri-

134
Aquatic Snakes

Figure 8.31. Karnsophis siantaris.

Photography by JCM.

orly, premaxillary bone absent, nasal bones greatly reduced in size, neural spine present on the
atlas (absent in Brachyorrhos). Distinguished from Brachyorrhos by its depressed skull, rostral
not visible from above; eye diameter greater than eye–mouth distance, countersunk lower
jaw, laterally compressed tail. Distinguished from Calamophis by 19 scale rows just anterior to
the vent (Calamophis has 17); divided internasal (single in Calamophis); divided nasal scale
(single in Calamophis); two postoculars (one in Calamophis). Size. Males: SVL: 415 mm; Tail:
12 mm. Females: no data. (Figure 8.31).
Natural History. This species has been confirmed to feed on frogs (unknown species;
Murphy and Voris, 2013). It is unknown if K. siantaris is a frog specialist, or if frogs are just a
part of this species’ diet.

Rear-Fanged Species

Gray’s Mangrove Snake

Bitia hydroides Gray, 1842
Distribution and Habitat. Coastal waters of southern Myanmar, Thailand, peninsular
Malaysia (Johor, Penang, Selangor, Penang Island; Charlton, 2020), Singapore, and Borneo.
There may be populations in other areas of the Greater Sun-
das (Murphy, 2007a). Type locality: Unknown. (Figure 8.32)
Identification. Extremely similar to true sea snake mor-
phologically but lacking a paddle-like tail and has eyes that
are more dorsal-facing. Scale rows at midbody 37–43 at mid-
body; ventral scales keeled; parietal scales fragmented; sin-
Figure 8.32. The distribution of gle internasal; dorsal scales small, triangular, exposed skin
Bitia hydroides. between them; head slightly distinguishable from neck; pari-
etals fragmented; head, neck, and anterior body slender with
body increasing in diameter posteriorly; single internasal
separates nasal scales; loreal present; loreal in contact with first 3 or 4 upper labials; ocu-
lar ring scales large, plate-like; upper labials 3–4 or 4–5 below the orbit; tail short 8–11% of
SVL; dorsal scales small, smooth, and triangular; dorsal scales have exposed skin between the
scales; ventrals very narrow with two rows of keels; maxillary teeth number 11–13, followed
by a gap and two enlarged grooved teeth. It has enlarged palatine teeth. Size. Males: Max SVL:
641 mm; Tail: 55 mm; | Average SVL: 338 mm; Females: Max SVL: 718 mm; Tail: 54 mm; |
Average SVL: 500 mm Juveniles: SVL: 134–172 mm | Average SVL: 153.9 mm. (Figure 8.33)
Natural History. Bitia hydroides is an intertidal species, inhabiting mud flats and estuar-
ies, and mangroves. It burrows into the mud. It is possible that the exposed, interstitial skin

135
 8. The Homalopsidae

Figure 8.33. Bitia hydroides. Muar, Malaysia.

Photography by Bruce Jayne.
of Bitia may aid in cutaneous respiration in the already low-oxygen environments it inhabits.
Like Cerberus, it may use a sidewinding-like motion on the surface of the mud. It is nocturnal
and may be seasonally active. The species is remarkably similar in appearance to sea snakes of
the genus Hydrophis, and it is often collected with Hydrophis in the sluice nets (or creeks) in
the Bassein River of Myanmar (Theobald, 1868). The diet is composed of fish (Cantor, 1847;
Boulenger, 1890) most of which are gobies, including Oxuderces dentatus (Jayne et al., 1995)
and the deep-water goby Trypauchen raha. Females mature at an early age (as young as 1
year) and have small litters, ranging from 1–10, with small neonates (Jayne et al., 1995). Ovi-
ducal eggs have been confirmed in this species, though this species has been considered as
having both viviparous or ovoviviparous modes of preproduction. Females captured between
20 November and 18 December were often gravid, and females have been found to grow to
significantly longer SVLs than males (Jayne et al. 1995).

Cantor’s Mangrove Snake

Cantoria violacea Girard, 1858
Distribution and Habitat. The Andaman Islands, Andaman coasts of Myanmar, Thai-
land, peninsular Malaysia, Sumatra, and Borneo. Type locality: Singapore (Figure 8.34)
Identification. Distinguished from all other homalopsids by the combination of 19 dor-
sal scales rows at midbody; nasals separated by internasal
scale; prefrontals make broad contact; five upper labials;
loreal contacts first 2–3 upper labials; subocular scale sepa-
rates upper labials 3–4 from orbit; 7–8 lower labials; ventral
count exceeds 243 (more ventrals than any other homalop-
sid). Size. Males: SVL: 358–902 mm; Tail: 71–193 mm (tail
18–21% of SVL | Average: 20%); Females: SVL: 620–957
Figure 8.34. The distribution of
Cantoria violacea.
mm; Tail: 127 mm (tail 12–17% of SVL | Average: 13.7%).
Males have a long, thin, stringy, gracile appearance while
females of the same size are more robust. (Figure 8.35-37)

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Aquatic Snakes

Natural History. Cantoria violacea are found in mangrove forests with mud lobster
mounds, mudflats, tidal creeks and river mouths, but will also occasionally enter nearby
freshwater environments. It utilizes mud-lobster burrows and crab holes during the day, pri-
marily being active at night. It will also use these holes or partially burrow into wet med to
escape predators (Murphy, 2007a; Voris and Murphy, 2002; Ghodke et al., 2018). It uses the
intertidal burrow system. Sworder (1923) considered this “a very rare snake” in Singapore.
This is undoubtedly due to its microhabitat, the intertidal burrow system, which is seldom
explored by herpetologists or other naturalists. Karns et al. (2002) found this species only
three times (1.1 % of the 270 snakes collected) at Pasar Ris Mangrove Park in Singapore, while
Ghodke and Andrews (2002) considered it common in the North Andaman Islands. Cantoria
violacea specializes in eating small fish and crustaceans, including alpheids (e.g., Alpheus mi-
crorhynchus, giant snapping prawn; Voris and Murphy, 2002) (Figure 8.38), and hard-shelled
crabs (Metaplax elegans; Ghodke et al., 2018). These snakes often start their attack using
their bodies to restrain and pin their prey (though, without coiling), as opposed to biting and
striking (Jayne et al., 2018). Depending on its prey, it might either swallow their prey whole
(shrimps; Murphy, 2007a) or rip their prey in pieces prior to consuming individual parts
(crabs; Ghodke et al., 2018).
The venom chemistry of this species is entirely unknown. However, Ghodke and An-
drews (2002) reported stories from Karens living in the Andaman Islands who stated that
they knew people who had been bitten by this snake, one had died, and another had been
unconscious for several days. Bite symptoms included abdominal pain, vomiting blood, and
swelling at the site of the bite. A folk remedy made from a “cactus-like plant that grows in
mangroves” was applied to the bite lesions and taken internally. They acknowledged that
Cantoria is usually considered harmless and has a similar appearance to the Andaman krait
(Bungarus andamanensis). This species forms a clade with the other crustacean-eating hom-
alopsids, Fordonia leucobalia and Gerarda prevostiana (Alfaro et al., 2008; Figueroa et al.,
2016; Burbrink et al., 2020; Bernstein et al., 2021). Cantoria is the sister genus of Gerarda
(Bernstein et al., 2023a).

Figure 8.35. Cantor’s

Mangrove Snake, Can-
toria violacea entering
a burrow in Thailand.
Photograph by Harry
Ward-Smith.

137
 8. The Homalopsidae

Figure 8.36. Cantoria vio-

lacea. Photograph by Ken-
neth Chin.

Figure 8.37. An Alpheus

pistol shrimp preyed upon
by Cantoria violacea.

Australian Bockadam
Cerberus australis (Gray,
1842)
Distribution and Habitat.
Found on the northern coasts
of Australia and the south
Figure 8.38. Cantoria violacea. coast of New Guinea. Its east-
Photography by Kenneth Chin. ern-most distribution based on
museum specimens is Edward
River (Queensland, Australia), and it may reach the tip of the Cape York Peninsula (Wilson
and Swan, 2003) though it the true limits of this species distribution is uncertain; records are
confirmed around the Gulf of Carpentaria. A single specimen is known from Waingapu,
Sumba Island, Indonesia. It is unclear if this is a waif or represents an established population
(WAM 101620; Murphy et al., 2012b). Type locality: Merauke, Dutch New Guinea. (Figure
8.39)
Identification. Dorsal scales in 23 rows at midbody, upper labials 2–4 contact loreal,
scales on the crown imbricate; those anterior to the angle of jaw lack keels; first labial does not
contact loreal; last upper labial horizontally divided; venter is mottled. This species is known
to exhibit several color morphs, such as grey, red, or even pale olive green (shown above).
Size. Males: SVL: 380–665 mm; Tail: 117 mm (tail 21–26.7% of SVL); Females: SVL: 445–756
mm; Tail: 106 mm (tail 19–22.6% of SVL). (Figure 8.40)
Natural History. All members of this genus have similar or identical habitat preferences
and behaviors. Cerberus australis are nocturnally active on mudflats, mangrove forests, and
streams that pass through these habitats. They may be found in the sea or at river mouths
(Kinghorn, 1929), or in estuaries and fresh-water streams, and can be found basking on mud-
flats (Heatwole, 1977). They are often present in groups, and, at low tide, burrow into soft
mud, leaving just the top of their head visible (Gow, 1989). This species is considered pisciv-
orous. While it has been reported to feed primarily on mudskippers, they might also feed on
other small fishes and crustaceans (Gow, 1989) in estuarine mangrove roots and the shelv-

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Aquatic Snakes

ing banks of tidal streams (Worrel, 1963); though, verification

of crustacean prey is still needed (Murphy, 2007a). Confirmed
predators include nervous sharks (Carcharhinus cautus; Lyle
and Timms, 1987). Like other members of this genus, C. austra-
lis may sidewind across mudflats to increase locomotion speed
(Kinghorn, 1929). The venom of this member of the genus is
Figure 8.39. The distribu- of little concern; stinging sensations have been reported (Gow,
tion of Cerberus australis. 1989). Data shows venom yields of 10 μl are typical, but up to 200
μl are possible; these data were from manual venom acquisition
(venom milking) and venom is shown to increase free platelet

Figure 8.40. Aus-

tralian Bockadam,
Cerberus australis.
Photography from
MAGNT.

clotting times from 12–40% and contains at least four major acidic proteins (Guinea et al.,
1992). Cerberus australis has been found to be the sister lineage to all other Cerberus in phy-
logenetic studies (Alfaro et al., 2004; Bernstein et al., 2021; Bernstein et al. 2023a).

Palau Bockadam
Cerberus dunsoni Murphy, Voris, and Karns, 2012
Distribution and Habitat. Palau Islands, Micronesia. Specifically known from Oreor
Island (=Koror Island), Arakabesang (formerly Ngerekebesang) Island, and Babeldaob Is-
land. Type locality: Micronesia: Palau Islands: Arakabesang (formerly Ngerekebesang) Island
(~7°20’ N, 134°27’ E). (Figure 8.41)
Identification. Dorsal scales in 23 rows at midbody, upper labials 1–4 contact loreal;
plate-like scales on crown rounded, juxtaposed, and thickened compared to scales of other
Cerberus species, ninth upper labial horizontally divided, ventral surface uniform black. The
9th upper labial is horizontally divided. These characters, combined with large parietal scale
fragments and a uniform black venter, make this a very distinctive species. The large, plate-
like fragments of the parietals may fuse with the temporal scales. The parietal scales in other
Cerberus are usually fragmented into small scales like scales on the crown. The scales on the
crown anterior to rictus usually lack keels; all other Cerberus except C. australis have scales
with keels anterior to the level of the angle of the jaw. Crown scales are distinctly rounded
and juxtaposed; in other Cerberus, these scales tend to be sharp-edged and slightly imbricate.
The light pigment forms an irregular ventrolateral stripe involving scale rows 1–2; this stripe

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Aquatic Snakes

involves rows 1–3 or 1–4 in other Cerberus except C. microlepis,

which has 27–29 scale rows at midbody. The ventral pattern is
uniform black or dark brown in preserved C. dunsoni, while all
other species have a mottled ventral surface or one with an irreg-
ular central stripe of dark pigment. This species tends to have a
broader, more robust head than the other species in the genus and
Figure 8.41. The distribu-
it lacks the dorsolateral pattern of bars or incomplete cross bands.
tion of Cerberus dunsoni.
Size. Males: No Data; Females: SVL: 228–687 mm; Tail: No Data.

Figure 8.42. Pulau Bock-

adam, Cerberus dunsoni.
Photography by JCM.

Neonate. SVL: 232 mm; Tail: 62 mm.(Figure 8.42)

Natural History. Cerberus dunsoni is poorly known in comparison to other Cerberus.
Populations have not recently been assessed, but Crombie and Pregill (1999) states that
Cerberus is common in Palau, but is not often collected, as it is a nocturnal snake in diffi-
cult-to-survey mangrove habitats. They also report that C. dunsoni is occasionally found in
the open ocean or along beaches, particularly near river or streams, as has bene reported for
C. australis (Kinghorn, 1929). They may cross stand vegetation between beach and man-
grove swamp habitats. While their reproductive mode has not studied, an umbilical scare
was present in a single neonate found by Pregill and Crombie (1999), so this species is likely
viviparous like viviparous like Cerberus schneiderii. They have facultative premaxillary salt
glands (Dunson and Dunson, 1979); it is uncertain if these are present in all Cerberus or just
the isolated C. dunsoni. This species has been recovered as a clade within Philippine C. schnei-
derii, and may represent a founder population of this C. schneiderii (Bernstein et al., 2021).

Lake Buhi Bockadam

Cerberus microlepis Boulenger, 1896
Distribution and Habitat. Known only from Lake Buhi and the adjacent rivers (personal
observation by JMB). Lake Buhi is a small freshwater lake in the Camarines Sur Province on
Luzon Island, Philippines. The lake is about 18 square kilometers and is positioned between
two volcanoes: Mt. Iriga (~1200 feet above sea level) on the western side of the lake and Mt.
Malinao (~3900 feet above sea level. Lake Buhi is 15 km from the coast. Type locality: Phil-
ippines. (Figure 8.43)
Identification. Dorsal scales 27 to 31 rows at midbody; all other species have 21–25 scale
rows at mid-body; the imbricate plate-like scales on the crown have a slightly thickened ap-
pearance; last upper labial is horizontally divided, and the venter is mottled. Size. Males:
Maximum TL: 768 mm; Tail: 158 mm (tail 23–27% of SVL). Females. Maximum TL: 1046
mm; Tail: 161 mm (tail 18–27% of SVL) (Figure 8.44).

140
 8. The Homalopsidae

Natural History. Collectors’ notes accompanying specimens report this species in

the shallow water fishponds at 1940–0100 hours. Personal
observations by JMB include abundant C. microlepis in the
river of San Pascual Municipality, which drains into Lake
Buhi’s southern coast. Snakes were found late at night, lay-
ing submerged on the river’s edges or swimming. Cerberus
schneiderii were sometimes within 1–2 meters of each other,
Figure 8.43, The distribution or in one case two specimens on top of one another. Locals
of Cerberus microlepis. from barangays around Lake Buhi refer to the bockadams
as ‘Aninikig’ and report them in high numbers after floods
and storms. This species is very closely related to Cerberus schneiderii, the Southeast Asian
Bockadam, which is often associated with brackish waters. Cerberus microlepis is often
recovered as a clade within C. schneiderii in phylogenetic trees (e.g., Alfaro et al., 2004,
Bernstein et al., 2021; Bernstein et al. 2023a) and may represent a freshwater-adapted pop-
ulation of C. schneiderii.

Figure 8.44. Lake Buhi Bockadam, Cerberus microlepis

Photograph by JMB

South Asian Bockadam

Cerberus rynchops (Schneider, 1799)
Distribution and Habitat. On the eastern edge of its range, it extends to the Mouth of
the Irrawaddy, Myanmar, and perhaps into the Mergui Archipelago. It is replaced by Cer-
berus schneiderii along the Andaman Sea coast of Thailand and coastal Southeast Asia. The
western edge of the distribution is likely to be the Gulf of Khambhat in the rivers and estu-
aries of Gujarat, India (Vyas et al., 2013). Type locality: “Ganjam” (Orissa State, SE India),
(~19°22’ N 85°03’ E). There are at least two other localities on India’s east coast that contain
the name “Ganjam:” Chinna Ganjam and Pedda Ganjam. Both are south of Ganjam, all
three locations are coastal and within the range of this species. Murphy et al. (2012) has
restricted the type locality to ‘Ganjam, India’ (Figure 8.44).
Identification. Dorsal scale rows 25 (rarely 23) at midbody, imbricate plate-like scales

141
Aquatic Snakes

on the crown have flat, thin appearance, keeled scales anterior to

the angle of jaw; the last two upper labials are horizontally divid-
ed; venter mottled. Cerberus rynchops can be distinguished from
all other members of the genus by its 25 (rarely 23) scale rows
at midbody, the imbricate plate-like scales on the crown have
Figure 8.45. . The distribu- a flat, thin appearance and are keeled anterior to angle of jaw;
tion of Cerberus rynchops the last two upper labials are horizontally divided; the venter is
mottled. Cerberus australis has 23 scale rows at mid-body, and
the first upper labial does not contact the loreal (it usually does
so in all other Cerberus species). Cerberus dunsoni has 23 scale rows at midbody, rounded
juxtaposed scales on the crown, and a uniform black venter. Cerberus schneiderii usually has
23 scale rows at mid-body (rarely 21 or 25). The last upper labial is horizontally divided (in-
stead of the last two in Cerberus rynchops) and the venter is mottled. Size. The largest male
was 777 mm in TL, and the largest female was 886 mm (Figure 8.46 - 8.47).
Natural History. This species, along with Cerberus schneiderii, are perhaps the most
well-known homalopsid snakes regarding natural history, in part due to their high abun-
dance and wide distributional ranges. Cerberus rynchops are coastal specialists that inhabit
mangrove forests and associated mud flats, tidal zones and creeks, and estuaries and rivers.
It has also been found at open sea (Acharji and Mukherjee, 1966), and is sometimes caught
in fishing traps together with true sea snakes (Hydrophiidae) (Anderson, 1889). At Chilka
Lake, Orissa, India, it uses seasonally flooded areas with numerous aquatic plants (Dutta,
1989). In Sri Lanka, it inhabits rivers and inland swamps, lagoons, and coastal waters (Der-
aniyagala, 1955). It is also common in human populated areas, such as near or in rice paddy
fields or near fishing communities. It is most often found in salt and brackish waters but is
also found in freshwater streams and lakes. It has even been found in fresh water more than
100 miles from the salt waters of coastal areas (Smith, 1943; Whitaker, 1969).
The diet is mostly composed of fish, primarily gobiids and carangids (Annandale, 1907;
Murphy, 2007a). Wall (1918) reports fish herding behavior. It flicks its tail, first on one side
and then the other in such a way as to make an unwary fish recede from the movement
towards its head; when coming within the snake’s field of vision, the fish is captured. Major
predators of this species include birds, such as white-bellied sea eagle (Haliaeetys [=Hali-
aeetus] leucogaster), house crows (Corvus splendens), Brahminy kites (Haliastur indus), and
pariah kites (Milyus migrans) in different parts of India (Saha 1984; Murthy and Rao, 1986).
Though not as common, other predators include mangrove crabs (Scylla serrata; Voris and
Jeffries, 1995).
Algal fouling has been reported in this species, though the effects it has on Cerberus
requires more research (Rahman and Reza, 2013). This species is extremely abundant in
India, with several published accounts noting that they might be the most or second most
common aquatic snake in swamps, shorelines, estuaries, and other systems (Keswall, 1886;
Dutta, 1989; Murthy, 1987; Whitaker, 1969).
The venom of Cerberus rynchops (and likely the entire genus) contains three known
venom protein families: metalloprotease, CRISP, and C-type lectin. It also contains unique
proteins (ryncolin1 and ryncolin2) that belong to a venom protein family unique amongst
snakes, called veficolins (OmPraba et al., 2010); it is presumed that veficolins may induce
platelet aggregation and/or initiate complement activation (OmPraba et al., 2010). Few ven-
om studies have bene performed, but injections in laboratory mice showed sluggishness,
partial paralysis, and death in 10–36 minutes, with post-mortem subcutaneous extrava-
sation of blood over the back (Alcock and Rogers, 1902). The bite and venom, however,

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Aquatic Snakes

Figure 8.46. South Asian Bockadam, Cerberus rynchops. Photography by Vivek-Sharma.

are considered medically insignificant for humans. Cerberus rynchops has been consistently
recovered as the sister species to Cerberus schneiderii (Alfaro et
al., 2004; Bernstein et al., 2021; Bernstein et al. 2023a). It is likely
that the mosaic geological history (e.g., plate tectonics, sea level
fluctuations; Hall, 2009) in the Quaternary has shaped the history
of Cerberus rynchops and Cerberus schneiderii.

Southeast Asian Bockadam

Figure 8.47. Divided up-
per labials of of Cerberus Cerberus schneiderii (Schlegel, 1837)
rynchops. Photography by
JCM. Distribution and Habitat. Throughout Southeast Asia as a
coastal specialist (though also can be found in inland streams and
aquatic habitats). Specifically, it is found along the Andaman Sea
coast of Thailand eastward to the Philippines and southward throughout peninsular Ma-
laysia. Its distribution widely continues throughout Indonesia, as far east as the Moluccan
islands of Halmahera and Seram. Type locality: Timor (Figure 8.48).
Identification. Dorsal scale rows in 23 (rarely 25) scale rows at midbody; the imbricate
plate-like scales on crown have flat, thin appearance, lack keels anterior to rictus; last up-
per labial horizontally divided; venter mottled. The background color of the venter can be
a cream, light yellow, or yellow. There seems to be significant variation in the cloacal plate,
anterior chin shield, lower labials at the first chin shield, type of scale, prefrontal plate, up-
per labials at loreal, and ocular ring within Philippine Cerberus schneiderii (Barrera Jr. et al.,
2017). Size. Males: SVL: 212–683 mm; Tail: 55–168 mm (tail 18–34% of SVL); Females: SVL:
190–901 mm; Tail: 45–184 mm (tail 18–28% of SVL). Neonates : SVL: 116–160 mm (Figure

143
 8. The Homalopsidae

8.49).
Natural History. Most of what is thought to be known
about Cerberus rynchops is attributed to Cerberus schneide-
rii, as all of the mainland Southeast Asian populations were
considered C. rynchops until Murphy et al. (2012b). They are
nocturnal and often found in the same muddy, mangrove,
Figure 8.48. The distribution of and intertidal habitats that the rest of the genus are found in
Cerberus schneiderii. (Murphy, 2007a). They have been reported as the dominant
snakes in Rhizophorus mucronata or Sonneratia sp. man-
groves of Komodo, as well as in shallow sea water (Hooger-
werf, 1954; Auffenberg, 1980). In the Philippines they are abundant in most low-elevation
aquatic systems, such as mangroves, intertidal zones, rivers, and lakes (see C. microlepis)
(Griffin, 1909; Gorman et al., 1981; Barrera Jr. et al., 2017; personal observations by JMB).
On Java they are commonly found in muddy, brackish tidal waters of the coastal zone, and
live-in holes in the mud until they are active at night (Hoesel, 1959). Based on diet data (two
Valencieannea longipinnus , a coral reef goby) of a single Malaysian C. schneiderii, there is
a possibility that C. schneiderii will enter coral reefs (L.T. Ming, personal communication
[Murphy, 2007a]). These accounts of habitat and microhabitat preference, coastal special-
ization, and nocturnal behavior have been reported for several localities, such as Thailand
(Taylor, 1965), Bali (Auffenberg, 1980), Singapore (Chim and Diong, 2013; de Lang, 2013),
peninsular Malaysia (Charlton, 2020), the Moluccas (Edgard and Lilley, 1993; Setiadi and
Hamidi, 2006;) and Sumatra (David and Vogel, 1996). They are known as ‘Aninikig’ in the
Philippines and locals find them in high numbers, especially after heavy rain events (personal
communication by JMB). Prey includes eels, crustaceans, and a variety of actinopterygian

Figure 8.49. Southeast Asian Bockadam, Cerberus schneideri. Photography by Kenneth Chin.

144
Aquatic Snakes

fish, most of which are gobies (Frith et al., 1977; Auffenberg,

1980; Gormon et al., 1981; Jayne et al., 1988; Murphy, 2007a).
Snakes will hold their mouth open while swimming to cap-
ture schooling fish. Prey tends to be small—less than 10% of
the predator’s mass. Snakes will sometimes anchor itself to a
convenient bamboo stake, anchor rope, or submerged branch
by its tail, and from this purchase swings about in the current
on the lookout for fish passing by. Cerberus schneiderii in the
Philippines will readily sidewind to escape and stay still when
approached; the Philippines C. schneiderii in the field were
found swimming in the intertidal zone at ~2300 h in March
in the Libmanan River mouth by San Miguel Bay, Camarines
Figure 8.50. Habitat for Cerbe- Sur Province, Luzon Islands (personal observation by JMB).
rus schneiderii. Photography by In the immediate area where these specimens were caught,
JCM Acrochordus granulatus is also present, suggesting sympatry
of these two aquatic snakes. This was also found to be the case
in other parts of the Philippines, were there is a significant
dietary overlap between Cerberus and Acrochordus (Gormon
et al., 1981). This species is viviparous and has 2–12, with a
direct correlation between female snake size and litter size
(Chim and Diong, 2009); though, this is likely higher as C.
rynchops can give litter sizes up to 38. Cerberus schneiderii
is recovered as sister to Cerberus rynchops; it is likely that C.
microlepis and C. dunsoni represent founder populations of
C. schneideri (Bernstein et al., 2021).

Kerala Mud Snake

Dieurostus dussumieri (Duméril, Bibron &
Duméril, 1854)
Distribution and Habitat. Coastal plain of South India
(Kerala and Tamil Nadu), from Vembanad-kole wetlands of
Kerala and Vellayani Lake to Neyyar River southern Kerala.
Figure 8.51. A Cerberus schnei- Type locality: west coast (Malabar coast) of North Kerala, In-
derii escapes into the water. dia (Figure 8.52.)
Photography by JCM Identification. Dorsal scales smooth in rows of 25 or 27,
nasal scales in contact, posterior labials horizontally divid-
ed by divided internasal, upper labials 1–3 contacting loreal;
five lower labials contact the chin shields; a mostly uniform
dorsum with lower ventrolateral stripe. The anterior edge of
the first pair of chin shields is narrower than the posterior
edge—ventral scales 146–148 in the two females. Size. Males.
no data. Unsexed specimens. Maximum SVL: 555–795 mm;
Maximum total length: 670–920 mm; Females. SVL: 491–670
mm; Tail: 74–77 mm (tail 12% of SVL) (Figure 8.53).
Natural History. Dieurostus dussumieri has a high salin-
ity tolerance but has been found in both fresh and brackish

145
 8. The Homalopsidae

Figure 8.53. Kerala Mud Snake, Dieurostus dussumieri.

Photography by Vivek-Sharma,

environments. They use marshy habitats, stagnant ponds, and

nearby areas with emergent and submerged aquatic vegetation
such as wetland rushes and water hyacinths as habitat. They
can also be found in paddy-fields (Kumar and Captain, 2011).
Dieurostus dussumieri is piscivorous, and identifiable prey
includes Anabas testudineus (Kumar and Captain, 2011). It
Figure 8.52. The distribution breeds early in June with the southwest monsoon; gestation
of Dieurostus dussumieri lasts more than five months. This species is viviparous and can
have litter sizes of 25 neonates (can have up to 32 eggs, some of
which may atrophy [Parameswaran 1954, 1962]). During the
early stages of pregnancy, a chorioallantoic placenta forms that supplies embryos with nutri-
ents after the yolk has been used (Parameswaran, 1962). Kumar and Captain (2011) mention
that theses snakes can be found buried in the mud, and also observed escape behavior in
which D. dussumieri escapes into the grasses and Ipomoea aquatica along the margins of
lakes and their flood plains. Personal communication with fisherman in Kumar and Captain
(2011) state that the bite of D. dussumieri, while only mildly venomous, is painful (though, it
is uncertain if this is due to lacerations from the teeth or the venom itself). Several hypotheses
using molecular data exist for its phylogenetic placement: it has been recovered as sister to
Myrrophis chinensis (Kumar et al., 2012); sister to a clade consisting of Hypsiscopus, Myr-
rophis, Gyiophis, and Enhydris (Quah et al., 2018); sister to Raclitia indica or all rear-fanged
homalopsis with the exclusion of Hypsiscopus (Bernstein et al., 2021); and other placements
within the rear-fanged group (Quah et al., 2017). Gyi (1970) hypothesized that D. dussumi-
eri is in the Ferania sieboldii group, and Murphy (2007a), based on morphological data, hy-
pothesized that D. dussumieri, F. sieboldii, and Mintonophis pakistanicus all share a common
ancestor. However, all of the molecular studies have varying levels of support. Bernstein et

146
Aquatic Snakes

al. (in review), used nuclear and mitochondrial DNA to recover a clade that consists of D.
dussumieri, F. sieboldii, and M. pakistanicus, which are closely related to the clade containing
Hypsiscopus, Enhydris, Myrrophis, and Gyiophis.

Trans-fly Mangrove Snake

Djokoiskandarus annulata (De Jong, 1926)
Distribution and Habitat. Found on the southern coast of New
Figure 8.54. The distri- Guinea, Trans-Fly region. Type locality: Prins Frederik Hendrik Is-
bution of Djokoiskan- land, West New Guinea (about 7°56’ S and 138°21’ E). (Figure 8.54).
darus annulatus. Identification. Mostly smooth scales but some of the posterior
dorsal scales keeled; dorsal scales in 21 rows at midbody reduced
to 19 rows posteriorly; internasal separates
prefrontal and nasals scales, frontal contacts
internasal. Size. Males: SVL: 470 mm; Tail: 108
mm (tail 18.7% of SVL) (n=1 specimen) (Fig-
ure 8.38)
Females: Total length: 227–548 mm; Tail:
Data Deficient (tail 16–17.8% of SVL); Juve-
nile: SVL: 254 mm; Tail: 44 (tail: 17.3% of SVL)
(Figure 8.55)
Natural History. Little is known about
the natural history of D. annulata. It occurs
at Abam in mangrove and Nipa palm habitats
(Parker, 1982). Parker (1982) also reported a
gravid female that was 568 mm (total length;
five eggs). The Trans-Fly region is a mosaic of
freshwater, marine, and swamps and monsoon
Figure 8.55. Trans-Fly Mangrove Snake, Djoko- and mangrove forests, and it is likely that D.
iskandarus annulatus MCZ R136063. Artist annulata inhabits these areas.
credit Lisa Kanellos.

Chanard’s Mud Snake

Enhydris chanardi Murphy & Voris, 2005
Distribution and Habitat. Restricted to vicinity of Bangkok, possibly as far west as Chant-
aburi, Thailand. No known extant population. Type locality:
Bangkok, Thailand (13° 45’ N, 100° 31’ E) (Figure 8.56). A
photograph of a live specimen Cox et al. (1998, p. 41) labeled
as Enhydris jagorii exists. However, a photograph (,http://
[Link]/photos/exotarium-oberhof/8243681837/.) of
a specimen reportedly from Mueang Nakhon Nayok, Nak-
Figure 8.56. The distribution of En- hon Nayok in Khao Yai National Park was recently seen, and
hydris chanardi.
this species may be more widely distributed than previously
thought. A photograph was also posted on social media on
November 11th, 2020 (locality: Thamai District, Chanthaburi Province, Thailand), though
this may be E. subtaeniata. All specimen of this snake come from metropolitan Bangkok area.
Enhydris chanardi appears to be endemic (or a near-endemic) to the Chao Phraya freshwater
swamp forest.

147
 8. The Homalopsidae

Figure 8.57. Enhydris chanardi.

Photography by Rica Achim Kempter.
Identification. Enhydris chanardi can be distinguished from all other Enhydris with 21
scale rows at midbody by the combination of the following characteristics: 116–125 ventral
scales, 38–60 subcadual scales (53–60 in males and 38–49 in females); two pairs of chin
shields with the second pair being the longest or equal to the first pair; lateral spots rows
4–5 or 4–6; stripe on scale row three. Dorsal scales lack striations (present in E. enhydris
and E. subtaeniata). Size. Males: SVL: 127–437 mm; Tail: 39–87 mm (tail 19.9–26% of SVL;
Females: SVL: 192–460 mm; Tail: 42–101 mm (tail 18.3–26% of SVL) (Figure 8.57).
Natural History. Despite the genus Enhydris being one of the most well-studied groups
of homalopsids, along with Cerberus, very little is known about E. chanardi. It is likely pi-
scivorous. One specimen at the Smithsonian National Museum of Natural History (USNM
83431) contains 13 well-developed embryos. Bernstein et al. (2021) obtained strong support
in a concatenated nuclear+mitochondrial tree for E. chanardi as the sister taxon to all other
Enhydris. In this study, Enhydris was consistently recovered as sister to a group containing
the reciprocally monophyletic Gyiophis and Myrrophis (strong and weak support in the con-
catenated and species tree analysis, respectively). New data (Bernstein et al. 2023a) using
genomic-level DNA supports this, and recovers Enhydris as sister to a clade containing Myr-
rophis, Myanophis, and Gyiophis, and also recovers E. chanardi as sister to all other Enhydris
species.

Rainbow Mud Snake

Enhydris enhydris (Schneider, 1799)
Distribution and Habitat. Widespread from Nepal and
eastern India to Indochina (and possibly southern China),
southward to the Greater Sunda Islands as far east as Bor-
neo. Its presence on Sulawesi is doubtful. Type locality: ‘‘In-
Figure 8.58. . The distribution diae orientalis. … lake of Ankapilly, coast of Coromandel,
of Enhydris enhydris. India.’’ (Figure 8.58)
Identification. Enhydris enhydris is a distinctly striped

148
Aquatic Snakes

Figure 8.60 Rainbow Mud Snake, Enhydris enhydris.

Photography by A. Konrad Mebert; B. JCM.
species with a small narrow head. The combination of 153–174 ventral scales, a pattern of
longitudinal stripes with no lateral or dorsal spots or bars, second pair of chin shields longer
than the first will distinguish this snake from other Enhydris. It may be most easily confused
with is its sympatric (in the Mekong drainage) sister species, E. subtaeniata, which has fewer
than 153 ventral scales; a pattern of spots on scale rows 4–6, and two or three pairs of chin
shield. When there are two pairs, the first pair is longer than the second (Figure 8.59).
Size. Males. Maximum SVL: 575 mm; Tail: 125 mm (tail 25–38% of SVL); Females. Max-

149
 8. The Homalopsidae

imum SVL: 734 mm; Tail: 148 mm (tail 20–35% of SVL).

Neonates: SVL: 143–206 mm (average=178 mm; Karns et
al. [2005]). There may be considerable size differences be-
tween populations.
Natural History. This is a highly aquatic snake that
uses a variety of stagnant, slow-moving, and shallow
aquatic habitats ranging from rice paddies, canals, ditches,
and drainpipes to lakes and rivers. It can be found in and
around urban areas; it thrives in disturbed habitats (Figure
8.60). It is considered a freshwater species. Although it has
been observed in brackish and salt water, it is not known if
these observations were due to flooding events of nearby
freshwater localities (Subaraj and Rajathurai, 2018). They
can be found in high numbers and studies using PIT tags
and radio telemetry in Thailand show that these snakes
can dominate the aquatic snake assemblages of the systems
they live in (Murphy et al., 1999; Karns et al., 2000). En-
hydris enhydris is likely to be one of the most common and
Figure 8.59. Habitat for Enhydris abundant species in its range. It will use drain tiles to move
enhydris at Ban Tha Hin, Lake around in metropolitan areas. Rarely is it far from water.
Songhkla Thailand. Stakes on the
Fish are the primary food, and specimens examined con-
left shoreline mark funnel traps
for snakes. Photography JCM
tained only fish. The most common species were cyprinids
(Rasbora sumatrana, Trichopsis vitta [=Trichopsis vittate])
and osphronemids (Trichogaster trichopterus (=Trichopo-
dus trichopterus) (Murphy, 2007a), but also include nan-
dids and anabantids (Brooks et al., 2009). Karns et al. (2005) dissected 141 specimens caught
as bycatch from gill nets in Thailand, and stomach contents only contained fish, supporting
that his is a piscivorous species. Cylindrophis jodiae (formeraly C. ruffus) has been a con-
firmed predator of E. enhydris (Brooks et al., 2009), as has Xenopeltis unicolor (Murphy,
2007a). Bandicoot rats (Bandicota indica) are also reported to feed on E. enhydris (Nandi,
1984). Females continue to hunt when gravid. The smallest females at maturity are 410 mm
(Brooks et al., 2009). Litter sizes range from 2–39 neonates. Parturition occurs likely occurs
year-round across the range, with most births timed with the onset of the monsoon season.
Reports of envenomation from this snake are in the literature; symptoms are localized (lo-
calized pain, swelling, throbbing [Acharji and Mukherjee, 1966; D’Abreu, 1912]). Molecular
systematics have consistently recovered a monophyletic Enhydris, with either E. enhydris or
E. subtaeniata as the sister taxon to a clade containing E. innominata, E. longicauda, and E.
jagorii (Alfaro et al., 2008; Karns et al., 2010a; Bernstein et
al., 2021). Genomic data (Bernstein et al. 2023a) recovers
E. enhydris

Jagor’s Mud Snake

Enhydris jagorii (Peters, 1863)
Figure 8.61 The distribution of
Enhydris jagorii. Distribution and Habitat. Probably restricted to Thai-
land’s Central Plain. Historical evidence suggests it occurs
in the vicinity of Bangkok. Only one extant population is from the northern edge of Thai-
land’s Central Plain at the Bung Ka Lo wetland, Uttaradit province, Thailand. Type locality:

150
Aquatic Snakes

Figure 8.62. Jagor’s Mud Snake, Enhydris jagorii. From northern Thailand.
Photography by JCM
“Siam” [Thailand]. (Figure 8.61).
Identification. Dorsal scales in 23–25 rows on the forebody, large, scalloped blotches or
spots on sides and dorsum, ventral scales 117–127; females have more than 50 subcaudals,
males have about 68, pupil mottled and colored like scales surrounding it. Eye diameter less
than the diameter of the nasal scale.
Note that E. jagorii, E. longicauda, and E. innominata as sister to E. longicauda and E.
innominata, but this sampling does not include E. jagorii (though, E. jagorii is likely in a clade
with the latter two taxa) are closely related. At some point, they are likely to be synonymized,
and E. jagorii is the oldest of the three names. Size. Male: Maximum SVL: 353 mm; Tail: 118
mm (tail 29–33.4% of SVL); Female: Maximum SVL: 195–445; Tail: 43– 93 mm (tail 22–26%
of SVL) (Figure 8.64). as sister to E. longicauda and E. innominata, but this sampling does not
include E. jagorii (though, E. jagorii is likely in a clade with the latter two taxa).
Natural History. Little is known about E. jagorii. Of 108 specimens, 22 contained fishes
of six families: Cyprinidae, Anabantidae, Channidae, Bagridae, Nandidae and Osphronemi-
dae (Pongcharoen et al., 2016). Gravid females were found in every month of the year except
March, December, and January. The absence of gravid females was likely the result of sam-
pling bias or sample size (Pongcharoen et al. 2016). Molecular systematics using mitochondri-
al and nuclear loci have shown that E. jagorii, E. innominata, and E. longicauda are extremely
similar or identical regarding sequence divergence (Karns et al., 2010a; Bernstein et al., 2021),
and Murphy (2007a) suggested that E. innominata and E. longicauda may be best regarded as
subspecies (‘races’) of E. jagorii, the oldest name of the three species.

151
 8. The Homalopsidae

Tay Ninh Mud Snake

Enhydris innominata (Morice, 1875a)
Distribution and Habitat. Lower Mekong drainage of
Vietnam and possibly into Cambodia (Murphy, 2007a). Type
locality: Tay-ninh, Cochinchina (= Tay Ninh Province, South
Vietnam) (Figure 8.63).
Identification. Extremely low ventral count (106 –117),
42–56 subcaudals, a highly distinctive pattern of black later-
Figure 8.63. The distribution al blotches (scalloped-shaped) arise from the ventral surface.
of Enhydris innominata. These distinguish this snake from most other Enhydris. En-
hydris jagorii has a higher ventral count (117–127), and small-
er temporal scales. Enhydris longicauda has more ventral
scales (122–136) and more subcaudal scales (52–76) than E. innominata. Enhydris chanardi
has a higher ventral scale count (110–125), small dorsolateral spots that involve two or three
scales: and larger, plate-like temporal scales. Enhydris enhydris and E. subtaeniata have stripes
in their patterns and smooth, striated dorsal scales, and elongated anterior chin shields. This
species has smooth scales without striations. The largest female measured had TL of 655 mm,
with a 50 mm tail. Size. Males. SVL: 233–390 mm; Tail: 47 mm (tail 25–33% of SVL); Females:
SVL: 176–605 mm; Tail: 39–50 mm (tail 18–24% of SVL). (Figure 8.62).
Natural History. Enhydris innominata uses artificial canals, flooded grasslands, and
Melaleuca swamp forest (Bryan Stuart, personal communication in Murphy [2007a]). This

Figure 8.64. Tay Ninh Mud Snake, Enhydris innominata.

habitat is within the Tonlé Sap-Mekong Peat Swamp Forest ecoregion of Wikramanayake et et
al. (2002). Three gravid females were 465–555 mm SVL and contained 13–32 eggs (X = 25.6).
Embryos were not detected in any of the eggs, and they were collected in South Vietnam in
late-October and November. See E. jagorii for systematics and evolution.

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Aquatic Snakes

Long-tailed Mud Snake

Enhydris longicauda (Bourret, 1934)
Distribution and Habitat. Restricted to waters and adja-
cent shorelines of the Tonlé Sap area of Cambodia. Type local-
ity: Tonlé Sap, Cambodia. (Figure 8.65).
Identification. Ventral scales 124–133, dorsolateral pat-
tern mostly black-purple with large, lateral, dark scalloped
Figure 8.65. The distribution blotches. In color pattern, it is most like E. jagorii and E. in-
of Enhydris longicauda. nominata. However, both E. jagorii and E. innominata demon-
strate sexual dimorphism of the subcaudal scales and show
less lateral compression of the tail. These two species also
have different ventral scale count ranges, E. jagorii 117–127,
and E. innominata 106 –117. Enhydris chanardi has fewer scale rows (20–22) around the neck
compared to E. longicauda (23–25), fewer ventral scales (110–122), and a distinctive pattern
of lateral spots involving 2–4 scales on scale rows 4–6. Enhydris enhydris and E. subtaeniata
have a dorsal pattern of
longitudinal stripes, but
E. subtaeniata’s pattern
includes lateral spots
on scale rows 4–6, ven-
tral scale count range
of 136–153. Enhydris
enhydris has more than
153 ventrals. Enhydris
enhydris and E. subtaeni-
ata have smooth, striated
scales, while E. longicau-
da, E. chanardi, E. jago-
rii, and E. innominata do
not. Size. Male: SVL: 503
mm; Tail: 173 mm (tail
23–35% of SVL); Fe-
male: SVL: 640 mm; Tail:
163 mm (tail 25–35% of
SVL) This taxon shows
no sexual dimorphism
in tail length, an unusual
condition for a homalop-
sid (Fig. 8.67).
Natural History.
A highly aquatic snake
that may leave the water
on rare occasions, but
as Tonlé Sap undergoes
Figure 8.66. Long-tailed Mud Snake, Enhydris longicauda. From Tonle
seasonal changes in water
Sap, Cambodia. Photography by Konrad Mebert.
level, it appears to estivate
in the mud. It may be in-

153
 8. The Homalopsidae

active during low water (Saint Girons and Pfeffer, 1972).

Fish likely compose a major portion of its diet; one speci-
men contained a Trichogaster tricopterus (Family Anaban-
tidae), and other prey items include bagrids, channids, cy-
prinids, nandids, and osphronemids (Brooks et al., 2009).
Little is known about its biology, but a female collected in
Figure 8.67 The distribution
June had 18 oviductal eggs. See E. jagorii for systematics and
of Enhydris subtaeniata.
evolution.

Kompong Mud Snake

Enhydris subtaeniata Bourret, 1934
Distribution and Habitat. Found in the wetlands of the Indochinese Peninsula, mostly
present in Mekong Drainage from Laos, Vietnam, Thailand, Cambodia; outside the Me-
kong drainage at Bung Boraphet, Thailand. Type locality: Kompong Speu, Cambodia (Fig-
ure 8.68).
Identification. This species has 136–153 ventral scales, and the color pattern contains
stripes and lateral spots. The presence of lateral spots and ventral counts separate this spe-
cies from E. enhydris, which has 153–174 ventral scales. Enhydris subtaeniata has two or
three pairs of elongated chin shields, rather than the flared condition found in E. jagorii and
its relatives (E. chanardi, E. innominata, E. longicauda). Both E. enhydris and E. subtaeniata
have dorsal scales with striations, which do not occur in other members of the genus. This
species has been long confused with Enhydris jagorii, and most of the older literature refer-
ences, particularly those outside of Thailand, to Enhydris jagorii refer to this species. Size.
SVL: 133–539 mm; Tail: 120 mm; Average SVL: 397 mm; Female: SVL: 128–743 mm; Tail:
127 mm; Average SVL: 402 mm (Figure 8.69).

Figure 8.68. Kompong Mud Snake, Enhydris subtaeniata from Thailand.

Photography by JCM.

154
Aquatic Snakes

Natural History. This species is found in shallow stagnant water and streams. It is syn-
topic with E. enhydris at many locations (Saint Girons and Pfeffer, 1972; Stuart et al., 2000;
Murphy, 2007a), but E. enhydris is numerically dominate (Murphy, 2007a). There are several
localities in central Thailand where E. enhydris and E. subtaeniata co-exists in similar num-
bers and both species have been collected in the same gill net. Prey includes fish and frogs.
This aquatic snake uses streams impoundments, padi, ditches. klongs, ponds, and probably
many other aquatic microhabitats. In Cambodia, Saint Girons (1972) and Saint Girons and
Pfeffer (1972) considered this snake to be strictly aquatic and diurnal, and they suggest it
migrates with the water as the water levels in the lake undergo season fluctuations. However,
Murphy (2007a) has seen them buried in the mud. Murphy (2007a) found three females con-
tained 7, 12, and 20 oviductal eggs, respectively. The smallest gravid female had a SVL of 572
mm. Deuve (1970) reported young born in April and May with the first rains; this assumes
his E. enhydris account pertains to this species and not E. enhydris. The phylogenetic affinity
of this species is uncertain but may be sister to all other Enhydris excluding E. chanardi or
sister to the E. jagorii+E. innominata+E. longicauda group (Karns et al., 2010a; Bernstein et
al., 2021). Genomic data supports this topology (Bernstein et al., 2023a).

Tentacled Snake
Erpeton tentaculatum Lacépède, 1800
Distribution and Habitat. Southern Thailand, Cambodia, southern Vietnam. Type local-
ity: Unknown (fide Manthey and Grossmann, 1997). (Figure 8.69).
Identification. Prominent paired rostral appendages (appendage scales arranged in a
pointed fashion, in successive annuli); dorsal scales in 34–39 at mid body, heavily keeled;
ventrals very narrow with keel on each side (left and right); the ventrals and rostral append-
ages readily separate this snake from all other homalopsids. Size. Males: Maximum SVL: 478
mm; Tail: 246 mm (tail 43–59% of SVL). Females: Maximum SVL: 536 mm; Tail: 231 mm
(tail 33–44% of SVL). Neonates. TL: 197–244 mm (Figure 8.70).
Natural History. Erpeton tentaculatum is a highly aquatic snake, living in shallow,
slow-moving, turbid water, with some emergent vegetation (Taylor, 1965; Cox, 1991; Murphy,
2007a). It likely never leaves the water, and while it may be cryptic in nature, Smith (1943) re-
ported that it is not uncommon in the country around Bangkok in ponds and sluggish waters,
and that Bourret stated it is not rare in Cambodia and Cochin China (=Vietnam). This spe-
cies is piscivorous and feeds on anabantid, bagrid, osphronemid fishes (Brooks et al., 2009).
Using a J-shaped hunting posture, it hunts from ambush, taking advantage of the C-start
response of fish to predators (Smith et al., 2002). Erpeton tentaculatum feints their body so
fish will initiate a C-start response towards the snake’s jaws
as it strikes (Smith et al., 2002; Catania, 2009). While the
rostral appendages of Erpeton have been the subject of much
research, Catania et al. (2010) found that the appendages are
sensitive mechanoreceptors that are innervated by branches
of the trigeminal nerve and respond to water movement. Ac-
Figure [Link] distribution of counts of feeding on amphibians (frogs, tadpoles) are cited
Erpeton tentaculatum. by Morice (1875b,c), along with plant matter. The amount
of plant matter and algae in the digestive tract of E. tentac-
ulatum has been found in multiple specimens (see Neill and
Allen [1956] and Saint Girons [1972]). However, given the ballistic nature of E. tentaculatum’s
strikes and the dense vegetation of its environment, this is likely secondary ingestion, as also

155
Aquatic Snakes

Figure 8.70. Tentacled

Snake, Erpeton tentac-
ulatum from southern
Thailand. Photography
by JCM.

156
 8. The Homalopsidae

suggested by Morice and Neil and Allen (1956). Cylindrophis

jodiae (formerly C. ruffus) has been a confirmed predator of
E. tentaculatum (Brooks et al., 2009). When this snake is han-
dled, it does not attempt to bite or escape; instead, it stiffens
its body (Smith, 1943). A captive snake placed in an aquarium
with one floating water hyacinth proved to be remarkably cryp-
tic. Its body posture and coloration combine to make it appear
Figure 8.71. Hunting posture part of the vegetation. Its cryptic nature is also enhanced by
of Erpeton in an aquarium the growth of algae on the skin. When placed in clean water
with a Fighting Fish, Betta the symbiotic algae are readily visible, and the fish (Betta) in
sp. Photography by JCM. the aquarium would graze along the snake’s body. Thus, besides
serving as camouflage, the algae may also help to attract prey.
Reproduction studies of Erpeton has found that litters of 5–13
young are born in July–October (Morice, 1875c; Smith, 1943;
Campden-Main, 1970; Martinez and Behler, 1988). The mean litter size is about eight (Mar-
tinez and Behler, 1988). Courtship was observed only once, on 24 February. Erpeton tentac-
ulatum has been recovered as the sister taxon of Subsessor bocourti (Alfaro et al., 2008; Quah
et al., 2017; Bernstein et al., 2021). However, Quah et al. (2018) recovered Raclitia indica as
the sister to E. tentaculatum. All these studies have low support for these relationships and
varying molecular datasets and taxa. Bernstein et al. (in review) used genomic data and found
a strongly supported relationships of Erpeton as sister to the Australasian genera Pseudofera-
nia and Myron. Thus, further investigation with more comprehensive sampling is needed to
obtain more accurate phylogenetic relationships.

Siebold’s Mud Snake

Ferania sieboldii (Schlegel, 1837)
Distribution and Habitat. Large rivers and associated flood plains from northwest pen-
insular India, Bangladesh, Nepal, and extreme western Myanmar. Possibly Bhutan (Lenz,
2012). Type locality: Jumma River, near Agra (Figure 8.32).
Identification. Dorsal scale rows 27–29; ventrals 143–156; subcaudals 28–56; prefrontals
contact loreal; upper labials in orbit 4 or 4–5; divided internasal; two postoculars; no suboc-
ulars; 7–9 upper labials; last one or two upper labials divided; 11–13 lower labials; hemipenis
extends to 14th subcaudal. Size. Males: no data. Females: Maximum SVL: 575; Tail: 105 (part-
ly damaged tail); Neonates: SVL: 172 – 191; Tail: 35–41 mm | Average SVL: 180 mm; Average
Tail: 39 mm (Figure 8.73)
Natural History. Ferania sieboldii is a poorly known homalopsid snake. Habitat includes
large rivers and associated flood plains; it has been found embedded in the mud (Gharpurey,
1944), in dead rushes in the water/mud substrate (Wall, 1897), and even underneath wood
stacks near water sources (Wall, 1908). It has been caught in fishing nets by fisherman (water
depth = 0.5 m) in the Kaliya Sot dam (Bhopoal, Madhya Pradesh State) of India (Thakur and
Watve, 2009). Prey may be exclusively fish. Reproduction: Litters of 5–7 neonates are likely.
The diet of this snake likely includes fish and frogs; Wall (1897) reported a captive F. sieboldii
ate a frog, and another specimen died trying to eat a fish. Mell (1928) described a passive de-
fensive posture and behavior of this species when disturbed: it alternated between mirrored
postures and will assume a loose coiled position. Thakur and Watve (2009) observed aggres-
sive behavior that was also supported by accounts from local fisherman of Bhopal: the snake
actively lunged and tried to bite, which often causes it to be killed once caught in fishing nets.

157
Aquatic Snakes

Gyi (1970) hypothesized it is closely related to Dieurostus dussumieri, and Murphy (2007a)
suggests that F. sieboldii, D. dussumieri, and Mintonophis pakistanica derive from a common
ancestor. Molecular data (Bernstein et al. 2023a) supports this hypothesis by Murphy (2007a).

Crab-eating Snake
Fordonia leucobalia (Schlegel, 1837)
Distribution and Habitat. Found on mainland coastal
habitats from Bangladesh, Myanmar, Thailand, and southern
Vietnam. Also, Nicobar Islands and Greater Sunda Islands
eastward to New Guinea and northern Queensland, Austra-
Figure 8.72. The distribution of lia. Type locality: Timor (Indonesia) (Fig. 8.74)
Ferania sieboldii. Identification. Dorsal scales in 25–29 rows at midbody;
five or six upper labials, third enters orbit; loreal usually ab-

Figure 8.73 . Siebold’s Mud Snake, Ferania sieboldii. a juvenile of Ferania from Lucknow, Uttar
Pradesh of India.
Photography by Vivek R Sharma.

sent; scales in ocular ring plate-like; frontal shorter than parietal; chin shields small, usually
two pair (can number 1–3), second pair contacting each other (a trait that will separate this
snake from all other homalopsids). Rear fangs grooved, exceptionally robust. Size. Males:
SVL: 394–528 mm; Tail: 57–82 mm (tail 13.5–15.9% of SVL); Females: SVL: 427–589 mm;
Tail: 46–58 mm (tail 9.8–12.6% of SVL). Neonates: SVL: 190–222 mm; Tail: 24–27 mm (Fig-
ure 8.76).
Natural History.
Found in shallow, stagnant water and streams. On occa-
sion, it may be found in monsoon forests or the open ocean
(O’Shea, 1986; Campden-Main, 1970), though it is unlikely
that they thrive in this habitat (Murphy, 2007a). This species
uses the intertidal burrow system. Karns et al. (2002) used
Figure 8.74. The distribution of
Fordonia leucobalia. radiotelemetry on three specimens in Singapore, and found
F. leucobalia to be quite sedentary (average 4.4 m over 7–10
days), primarily nocturnal, and were often found in mud
lobster mounds or foraging on tidal mud flats. The use of

158
 8. The Homalopsidae

mangrove systems,
mud flats, and crusta-
cean burrows by pop-
ulations throughout its
range is supported in
the literature (Malayan
Peninsula and Thailand
[Macnae, 1968]; Java
[Kopstein 1931, Hoesel,
1959]; Australia [Cog-
ger, 1981]; New Guinea
[Parker, 1982; O’Shea
1986]). It has been ob-
served using sidewind-
ing behavior on the
mudflats (Cogger and
Lindner, 1974). Their
diet is almost entirely
crustaceans, including:
Sarmiatium germaini,
Macrophthalmus sp.,
graspid crabs, Dotillop-
sis brevitarsis, Uca sp.
(ocyopodids, fiddler
crabs), and Thalassina
anomala (mud lob-
ster - Fig. 8.70)) (Gow,
1989; Voris and Mur-
phy, 2002). Like other
snakes, they are likely
predated on by birds
and mammals; they
have been reported to
be eaten nervous sharks
(Carcharhinus cautus;
Lyle and Timms, 1987)
and other squamates,
such as Bungarus candi-
dus and Varanus indicus
Figure 8.75. Crab-eating
Snake, Fordonia leuco-
balia. Top two are from
Penang. Photograph from
Evan Quah. The next two
are from the Northern
Territory of Australia
Photography by JCM and
Mark Sanders.

159
Aquatic Snakes

(Murphy, 2007a). They have been observed feeding in the day-

time by Nobbs and Blamires (2004), who also made two separate
observations of F. leucobalia separating the tail from the cephalo-
thorax prior to eating (behavior unique to this taxon and Gerar-
da prevostiana amongst all alethinophidians, except the natricid
Opisthotropis cf. spenceri; Noonloy et al., 2018). This has been fur-
ther observed in the field and in experimental setups: F. leucobalia
subdues its prey with its venom, then pins the prey to the mud
substrate, subsequently ripping the prey apart, usually by the legs
and sometimes by chewing (Shine and Schwaner, 1985; O’Shea,
1996; Voris and Murphy, 2002; Jayne et al., 2018). Fordonia leu-
cobalia is considered a specialist on hardshelled crabs (Jayne et
al., 2002). Litter size range from 2–17 neonates (Murphy, 2007a).
Bergman (1960) suggests that adults are sexually mature at 330
mm SVL. The mainland Asia populations tend to be a uniform
black, while Australasian populations show dramatic color poly-
morphisms with red, white, and tan color morphs. Genomic data
(Bernstein et al. 2023a) recovered a relationship in which For-
donia is the sister to the sister-pair Gerarda+ Cantoria. Fordonia Figure 8.77 Mud lobsters of
leucobalia, Gerarda prevostiana, and Cantoria violacea, the three the genus Thalassina. build
crustacean specialists of the Homalopsidae, are often strongly mounds of dirt around its
supported as a clade in molecular systematics studies (Alfaro et burrow. The mounds are
al., 2008; Burbrink et al., 2020; Bernstein et al., 2021). used by several homalop-
sid snakes for refugia and
the crustaceans are eaten
Gerard’s Crab-eating Snake by both Fordonia and Ge-
Gerarda prevostiana (Eydoux & Gervais, 1837) rarda. Photography by
Bruce Jayne.
Distribution and Habitat. A widespread coastal species from
the vicinity of the Gulf of Kutch, India eastward to Sri Lanka,
Myanmar, Thailand, Cambodia, peninsular Malaysia,
Borneo, and the Philippines. The snake is likely rare in Sri Lanka (Somaweera et al.,
2006), and the Indian and Sri Lankan populations are restricted to both countries’ western
coasts (Karunarathna et al., 2018). It is probably widespread in the Indonesian Archipelago,
but specimens are absent due to the secretive habits of this snake, other than a report from
Sarawak, East Malaysia (Borneo; Das et al., 2013). Type locality: “Manille” (= Manila, 14°37’
N, 120°58’ E, Luzon, Philippines) (Figure 8.78).
Description. A relatively small and slender species; dorsal scales in 17 rows at mid-body;
loreal present, in contact with upper labials 1–3; of 7–8 upper labials, the fourth enters
orbit; postoculars are usually two with bottom scale extending partially under the eye; first
pair of chin shields large, second pair much smaller and separated by a pair of smaller scales.
Size. Males: SVL: 282–449 mm; Tail: 42–65 mm (tail 13.5–18.4% of SVL); Females: SVL:
254–434; Tail: 40–60 mm (tail 13–16.9% of SVL) (Figure 8.77.)
Natural History. A nocturnally active snake in mangrove forests with or without mud
lobster mounds. Multiple studies have observed it as a coastal, intertidal snakes (Wall, 1921;
Lim, 1963; Taylor, 1965; Tweedie, 1983; Gharpurey, 1944; Murphy, 2007a). It has been
excavated from the mounds of mud lobsters, specifically Thalassinia anomala and T. gracilis
(Karns et al., 2002; Voris and Murphy, 2002), which occur toward the landward edge of the
mangal. It appears to be more active on two spring tide survey nights (full and or new

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Aquatic Snakes

moon; 16 snakes) compared to eight survey nights on the neap tide

(half-moon: 12 snakes). Sri Lankan populations have also been found
to only be active at night (Karunarathna et al., 2018). A variety of mi-
crohabitats are used from the landward edge of the mangal to the mud
pools in the lower tidal zone at Pasir Ris Park in Singapore. However,
Karns et al. (2002) found that it was not restricted to areas around mud
Figure 8.76. . The distribution of lobster mounds. The diet of G. prevostiana has been presumed to in-
Gerarda prevostiana clude fish, shrimp, and other aquatic animals (Cox, 1991), but recent
literature has not supported this; Voris and Murphy (2002) never found
fish in the stomach contents of examined specimens. This snake tears
its prey into pieces before engulfing (Jayne et al., 2002). This is similar
to F. leucobalia (hard-shelled crab specialist), but G. prevostiana special-
izes on freshly molted crabs and does not press their prey into the mud; rather, they rip apart the crab
carapace assisted by body coiling (Jayne et al., 2018). The only other non-homalopsid snake that rips

Figure 8.77, Gerard’s Crab-eating Snake, Gerarda prevostiana

Photography by Vivek-Sharma

their prey apart (other than scolecophidians, which may decapitate termites) scale rows square; 129
(female) ventral scales; 30/29 (female) paired subcaudals; a divided cloacal plate; eight or nine upper
labials; ten lower labials; a maximum total length of 416 mm; relative tail length ratio of 0.13; a ventral
patterning lacking a central spot on each ventral scale; the presence of a faint stripe on the lower, dorsal
scale rows; and four rows of dark spots on the dorsum. Gyiophis salweenensis can be distinguished from
G. maculosa by the shape of the dorsal scales of first three rows (square in G. salweenensis vs. ovate in
G. maculosa), the ventral scale pattern (absence of a central spot on each ventral scale vs. its presence),
and a stripe running through the scales of the lower dorsal scale row (faint stripe vs. absent). It is dis-

161
 8. The Homalopsidae

Figure 8.78. A mud lobster mound on the west coast of peninsular Thailand. The burrow is visible.
Harold Voris (left) Tanya Chanard and a graduate student (right). Photography by JCM.
tinguished from G. vorisi by its lower number of ventrals (129 in G. salweenensis vs. 142–152
in G. vorisi), lower number of subcaudals (30/29 [left/right] vs. 41–58), shape of the rostral
scale (narrow vs. broad), and the number of rows of spots on the dorsum (four vs. three). Size.
Male: no data. Female: SVL: 364 mm; Tail: 52 mm (Fig.8.77)
Natural History. Any aspect of biology or natural history of this species is solely known
from the holotype (Quah et al., 2017). The single specimen of G. salweenensis was found at
~1930 hours crossing a narrow dirt road between flooded fields. Given the abundance of
other homalopsid species that utilize flooded fields or intertidal habitats, it is likely that these
flooded fields are natural habitat for G. salweenensis. This species has the dorsally-oriented,
semilunar valvular nostrils that indicate an aquatic/semiaquatic lifestyle. Although more data
is needed, the collection time of this specimen may indicate it has nocturnal habits. Quah et
al. (2017) mention that the holotype had a puncture wound on its back, which may have come
from a predation attempt from a heron. This species is recovered as sister to Myanophis, both
of which are sister to Myrrophis (Bernstein et al. 2023a).’

Spotted Mud Snake

Gyiophis maculosa (Blanford, 1881)
Distribution and Habitat. Known only from Bajo in Ayeyar-
wady River Delta, Myanmar; and known from only two spec-
Figure 8.79. The distribu-
imens, one of which (the holotype) appears to be lost. The pop-
tion of Gyiophis maculosa.
ulation may extend to Dagon Township, Yangon (= Rangoon),
Ayeyarwady State, Myanmar (JMB, Kate Woolf; personal commu-

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Aquatic Snakes

nication). Type locality: ‘Pegu (probably in the neighborhood of Bassein)’ (= Bago, in south-
ern Myanmar) (Fig. 8.81). Identification. This species has three
pairs of chin shields, 123 ventrals, 25 scale rows on the neck, and
a dorsal pattern of blotches without a stripe. These traits will
separate it from G. vorisi which has more than 142 ventrals, a
stripe on scale rows 2–4, 26-28 (usually 27) scale rows on the
neck, a depressed head, and a bold zigzag stripe on the edge of
the ventral scales that extends onto scale rows one and two, and
a cream-colored stripe on the upper part of row two and row
three; this stripe is bordered dorsally by a row of dark spots. It is
distinguished from G. salweenensis by having ovate scales for the
first three dorsal scale rows (square in G. salweenensis), presence
of a central spot on each ventral scale (absent in G. salweenensis),
absence of a stripe running through the scales of the lower dorsal
scale row (faint in G. salweenensis). In G. maculosa, three up-
per labials contact the loreal, smallest upper labial is the eighth,
ten lower labials; 25 scale rows on the neck, and 25 scale rows at
midbody; ventrals 122–125 ventral scales; and 32–33 subcaudal
scales. Size. no data (Fig. 8.80).
Natural History. Almost nothing is known about this spe-
cies, and very little is known about the genus as a whole. It ap-
pears to be endemic to the Ayeyarwady freshwater swamp forest Figure 8.80. Spotted Mud
ecoregion of the Indochina Bioregion (Wikramanayake et al., Snake, Gyiophis maculosa,
BMNH 1913.6.12.1 labeled
2002). A specimen of G. maculosa (based on the presence of a
“blanfordi.” Photography by
central spot on the ventral scales) may have been found in Dagon, JCM.
Yangon City, but the molecular analyses and a thorough examina-
tion morphology is still needed to determine its species-level identity.).

Salween Mud Snake

Gyiophis salweenensis Quah, et al. 2017
Distribution and Habitat. Only known Mawlamyine, Mon
State, Myanmar (type locality). Type locality: close to San-
pel Cave, Mawlamyine, Mon State, Myanmar (16°22.427 N,
97°46.388 E; 44 m in elevation) (Fig. 8.81).
Figure 8.81. The distribution of Identification. This species is separated from the other
Gyiophis salweenensis.. two Gyiophis by having a unique combination of the following
characters: a narrow rostral scale; the first three dorsal scale
rows square; 129 (female) ventral scales; 30/29 (female) paired subcaudals; a divided cloacal
plate; eight or nine upper labials; ten lower labials; a maximum total length of 416 mm; rela-
tive tail length ratio of 0.13; a ventral patterning lacking a central spot on each ventral scale;
the presence of a faint stripe on the lower, dorsal scale rows; and four rows of dark spots on
the dorsum. Gyiophis salweenensis can be distinguished from G. maculosa by the shape of
the dorsal scales of first three rows (square in G. salweenensis vs. ovate in G. maculosa), the
ventral scale pattern (absence of a central spot on each ventral scale vs. its presence), and a
stripe running through the scales of the lower dorsal scale row (faint stripe vs. absent). It
is distinguished from G. vorisi by its lower number of ventrals (129 in G. salweenensis vs.
142–152 in G. vorisi), lower number of subcaudals (30/29 [left/right] vs. 41–58), shape of

163
 8. The Homalopsidae

Figure 8.82. Salween Mud Snake, Gyiophis salweenensis

Photography by Evan Quah.
the rostral scale (narrow vs. broad), and the number of rows of spots on the dorsum (four vs.
three). Size. Male: no data. Female: SVL: 364 mm; Tail: 52 mm (Fig.8.82) Natural History.
Any aspect of biology or natural history of this species is solely known from the holotype
(Quah et al., 2017). The single specimen of G. salweenensis was found at ~1930 hours crossing
a narrow dirt road between flooded fields. Given the abundance of other homalopsid species
that utilize flooded fields or intertidal habitats, it is likely that these flooded fields are natural
habitat for G. salweenensis. This species has the dorsally-oriented, semilunar valvular nostrils
that indicate an aquatic/semiaquatic

Maubin Mud Snake.

Gyiophis vorisi (Murphy, 2007b)
Distribution and Habitat. Known only from the vicinity of Maubin in the Ayeyarwady
River Delta, Myanmar. Type locality: Maubin, Ayeyarwady Division, Myanmar (about 16° 73’
N, 95° 65’ E) (Fig.8.83).
Identification. Two pairs of chin shields, 142–152 ventrals, dorsal pattern of stripes that
are lateral and dorsal, and may break up into smaller spots. All these traits separate it from
G. maculosa, a species with which it has been long confused. Gyiophis vorisi is distinguished
from G. salweenensis by having a higher number of ventrals (142–152 in G. vorisi vs. 129 in
G. salweenensis), higher number of subcaudals (41–59 vs. 30/29 [left/right]), shape of the
rostral scale (broad vs. narrow), and the number of rows of spots on the dorsum (three vs.
four) (Fig.8.84)
Size. Males: SVL: 378–450 mm; Tail: 90–102 mm (tail 24–25% of SVL) | Average SVL:
390.2 mm; Average Tail: 95.8 mm; Females: SVL: 371–505 mm; Tail: 73–86 mm (tail 16–24%
of SVL) | Average SVL: 429.6 mm; Average Tail: 81.1 mm.
Natural History. Known from the Maubin area, which is
in the Ayeyarwady Freshwater Swamp Forest ecoregion. None
of the specimens examined by Murphy (2007b) contained prey.
A 450 mm SVL female (KU 92397) had five oviductal eggs
Figure 8.83. Distribution of Gy- (Murphy, 2007b). The species inhabits a severely degraded
iophis vorisi. habitat. Wikramanayake et al. (2002) report the sedimenta-
tion rate of the Ayeyarwady River is the fifth highest in the

164
Aquatic Snakes

world because of the deforestation that has occurred upstream in recent

decades.

Mamberamo River Water Snake

Heurnia ventromaculata Jong, 1926
Distribution and Habitat. Only known from the type locality in
northwestern West Papua, Indonesia. Pionierbivak, Mamberano River,
northern New Guinea. Type locality: Pionierbivak, Mamberamo River,
NW New Guinea (Fig. 8.85.)
Identification. Dorsal scales in 27 rows; the internasal is large; pre-
frontals are fused with the loreals and separate the internasal scale from
the frontal; lower postocular extends under the orbit; upper labials eight
or nine, with the fifth entering the orbit. Size. Males: SVL: 554 mm; Tail:
148 mm. Known only from the holotype
Natural History. Known only from the type specimen, no informa-
tion exists on the natural history of H. ventromaculata. The area H. ven-
tromaculata is known from is filled with estuaries of the Mamberamo
River, freshwater swamps, and extensive mangroves areas (Muchtar and
Figure 8.84. Maubin Mud Ilahude, 2001; Richards and Suryadi, 2002). It is possible that these areas
Snake, Gyiophis vorisi. Photog- are home to H. ventromaculata, as many homalopsids have adapted to
raphy by JCM these types of ecosystems.

Borneo Mud Snake

Homalophis doriae Peters, 1871
Distribution and Habitat. Known only from the island of Borneo,
and is widespread on the island (Figure 8.86). Type locality: Sarawak,
Figure 8.85. The distribu- Borneo.
tion of Heurnia ventromac-
Identification. Dorsal scales in 29–33 rows at mid-body; suboculars
ulata
present; 11–16 upper labials with last 5–9 horizontally divided. It may
be most easily confused with H. gyii, which has 25–27 scale rows at mid-
body and more than 153 ventrals. Size. Males: SVL: 175–580 mm; Tail:
125 mm (tail 20–26% of SVL); Females: SVL: 193–696 mm; Tail: 101
mm (tail 14–18% of SVL) (Figure 8.87).
Natural History. Homalophis doriae is found exclusively in swampy
habitats and muddy rivers below 500 m in coastal Sarawak and Sabah; it
Figure 8.86. The distribu- has been found in the same locality as Hypsiscopus plumbeus (Stuebing
tion of Homalophis doriae. and Inger, 1999). It can be abundant in stagnant or slow-moving water
of rice fields and ponds and sometimes burrows in muddy river bottoms
during the day, becoming more active at night (Iskandar, 2004). Its diet
is composed of fish, has been caught in fishing nets, and is considered
a pest by aquaculturists in eastern Kalimantan (Iskandar, 2004). Con-
firmed prey items include Systomus cf. orphoides (barbs; Voris and Mur-
phy, 2002). Three gravid females from Murphy (2007a) contained 6–16 eggs or embryos. Bernstein
et al. (in review), using mitochondrial DNA, found that Homalophis doriae is sister to Phytolopsis
punctata.

165
Aquatic Snakes

Figure 8.87. Borneo Mud Snake, Homalophis doriae

Photography by Anton Sorokin.
Gyii’s Mud Snake
Homalophis gyii (Murphy, Voris & Auliya, 2005)
Distribution and Habitat. Known only from the Kapuas River drainage in Kalimantan,
Indonesia. Type locality: Kalbar, Sungei Kapuas, near Putussibau, Indonesia (Figure 8.88).
Identification. Dorsal scales in 25–27 rows at mid-body; subocular scales present; 15–16
upper labials, ones below and behind eye horizontally divided. It may be most easily confused
with H. doriae, which has 29–33 scale rows at mid body, and less than 153 ventrals. Size.
Males: no data. Females: Maximum SVL: 666 mm; Tail: 96 mm (tail 14–18% of SVL) (Figure
8.89).
Natural History. This snake is likely the sister to H. doriae. Little is known about the hab-
itat, feeding habits, and reproduction of this species. Two specimens were obtained during
periods of heavy rainfall in flooded riparian habitats. The wet season in Putussibau may be
characterized by months that exceed 400 mm of precipitation (January, February, August,
October, and November), and a total annual rainfall of 4480 mm. The Reserve is west of the
type locality and is a large complex of swamp forest, peat swamps, and lakes. This area is
subject to two annual monsoons, a northeast monsoon in October and the second monsoon
in March. Water levels in the area recede by August, at which time the floodplain lakes are
completely dry for a 3–4-month period; the area then becomes a dry savanna with high daily
temperatures. This climatic regime has produced an environment unique from the surround-
ing lowland forest and several endemic fish and bird species are
known from the region (Sebastian, 1994). It is uncertain if H. gyii
and H. doriae are sympatric. This snake has also been referred
to as the ‘chameleon snake,’ due to its potential ability to change
from a reddish-brown color to completely white within minutes
(Auliya, 2003). This phenomenon in H. gyii may be a defense/
Figure 8.88. The distribu- warning mechanism, but it is poorly understood in this species
tion of Homalophis gyii. and has yet to be published on since.

166
Aquatic Snakes

Figure 8.89. Gyii’s Mud Snake, Homalophis gyii from Kalamantan, Indonesia.
Photography by Mark Auliya.
Puffed-faced Watersnake
Homalopsis buccata (Linnaeus, 1758)
Distribution and Habitat. Found from extreme southern Thailand, peninsular Malaysia,
Indonesian Archipelago from Sumatra to Borneo. A population may exist on Sulawesi (Rooij,
1917), but this has yet to be confirmed (de Lang and Vogel, 2005; Murphy et al., 2012c). Type
locality: “Indiis” (in error; given erroneously as “India” by Manthey and Grossman1997).
“Indiis” was likely meant to mean ‘East Indies,’ now the Indonesian Archipelago. The type
locality is now Singapore (Murphy et al., 2012c) (Figure 8.93).
Identification. Upper labials 1–4 single contact loreal; two prefrontals; 33–40 dorsal scale
rows at midbody, usually reduced to less than 30 posteriorly; one postocular plus a post-
subocular; 12 (11–14) upper labials; ventral count less than 166. Homalopsis buccata can be
distinguished from H. hardwickii by the latter species having a divided loreal; Homalopsis ni-
groventralis has upper labials 1–3 contacting the loreal; Homalopsis semizonata has a divided
or fragmented loreal contacting upper labials 1–4 or 1–5 and three prefrontals; H. mereljcoxi,
has a single loreal contacting upper labials 1–4, scale rows at midbody 40–47, reduced to 30
or more posteriorly. Size. Males: SVL: 167 – 695 mm; Tail: 56–236 mm (tail 28–34% of SVL);
Females: SVL: 192–794 mm; Tail: 62–202 mm (tail 23–26% of SVL); Neonates: 227–450 mm
(Figure 8.91-92).
Natural History. Homalopsis buccata is an aquatic species that feeds on fish. Much of the
literature that reference H. buccata are referring to other species within the genus. The habitat
used by H. buccata may include a variety of aquatic habitat, from freshwater streams, ponds,
agricultural drains and irrigated fields (Cantor, 1847; Charlton, 2020), to brackish waters (Ha-
gen, 1890) and peat swamp forests (specimen notes of California Academy of Sciences speci-
men CAS 174127 [Narathiwat, Thailand]). In Malaysia, they have been found both inland and
coastally, as well as in vacated crab holes in swamps, river banks, streams, and ponds (Lim,
1964). While considered a nocturnal, Lim (1964) found it foraging during the day along the
edges of water bodies. This species may even inhabit polluted habitats, as it is still found in Kua-

167
 8. The Homalopsidae

la Lumpur’s
Klang River
(Malaysia),
which is of-
ten flood-
ed, muddy,
Figure 8.90. The distribu-
tion of Homalopsis buccata.
and polluted
(Berry and
Lim, 1967). In
Sumatra, it has been described as the most
common water snake in Medan (specifi-
cally, Deli River, in root systems). This is
another species that has been found to
sidewind (Bergman, 1951). Specific prey
items for H. buccata include fish (Tila-
pia, Lebistes, Mystus, Claris sp. Chana sp.
Puntius binotatus) and eels (Fluta alba)
(Bergman, 1951; Berry and Lim, 1967;
Lim, 1964). Deuve (1970) mentions fresh-
water crustaceans as a part of their diet,
but this refers to the Laos populations,
which likely represent what is now con-
sidered H. nigroventralis (Murphy et al., Figure 8.91. A juvenile Homalopsis buccata. Kala-
2012c). It is likely Cylindrophis sp. (C. ruffus mantan. Photography by Rob Stuebing,
or C. jodiae) are predators of Homalopsis
(Greene, 1893). Cobras and kraits may also
feed on them, as Homalopsis buccata are
susceptible to the elapids’ venom (captive
observation; Bergman, 1951). No defini-
tive breeding season has been confirmed,
and juveniles have been found year-round
in a population in Kuala Lumpur, Malaysia
(Berry and Lim, 1967). Litter sizes range
form 4–37 (embryos and eggs; Murphy,
2007a). Size of reproductive maturity varies
across studies that have looked at different
species (based on locality); Bergman (1951)
reported a 569 mm SVL females with 4 mm
eggs. Homalopsis has been recovered as the
sister to Cerberus when analyzed with ge-
netic data (Alfaro et al., 2008; Bernstein et
al., 2021; Bernstein et al. 2023a). Before the
use of molecular data, it was hypothesized
that Cerberus, Homalopsis, and Enhydris
Figure 8.92. Puffed-faced Watersnake, Hom-
alopsis buccata. Krabi, Thailand. Photography by
Harry Ward-Smith.

168
Aquatic Snakes

were closely related (Gyi, 1970). Studies using more sampling are needed to identify strongly
supported, intergeneric relationships, which are currently lacking for the family. Genomic
data (Bernstein et al. 2023a) supports that Homalopsis and Cerberus are sister to each other,
but are distantly related to Enhydris.’

Indian Puff-faced Watersnake

Homalopsis hardwickii Gray, 1842
Distribution and Habitat. Known only from type specimen, this species may be restrict-
ed to northeast India or Nepal. Its distribution is poorly known. Type locality: India (datum
questionable; Murphy et al., 2012c) (Figure 8.93).
Identification. Homalopsis hardwickii has a divided loreal contacting upper labials 1–4;
two prefrontals; scale rows at midbody 39, reduced to 28 posteriorly; one postocular and
no presubocular; 159 ventrals. Homalopsis buccata has a single loreal contacting upper la-
bials 1–4. H. nigroventralis has upper labials 1–3 contacting loreal. Homalopsis semizonata
has three prefrontals. Homalopsis mereljcoxi has a single loreal and two postoculars plus a
postsubocular and 40 –47 scale rows at midbody, reduced to 30 or more posteriorly. Known
only from the holotype. Size. Males: SVL: 511 mm; Tail: 172 mm (tail 33.7% of SVL). Data
Deficient. (Figure 8.94).
Natural History. Nothing is known about the natural history of Homalopsis hardwickii. It
is likely similar to the other members of the genus, but much
remains to be discovered.

Jack’s Watersnake
Homalopsis mereljcoxi Murphy et al. 2012
Figure 8.93. The distribution of
Homalopsis hardwickii. Distribution and Habitat. Known from lowland local-
ities in Thailand, Cambodia, and Vietnam. It is known as
far north as Bung cho, Uttaradit Province, Phichai District,
Thailand (~12° N, 104° E) and as far south as Lake Songkhla
(~7° N, 100° E), Thailand. In Vietnam, it is known as far east
as Can Tho Province (~10° N, 105° E), and as far south as
Vinh Thuan District Town in Kien Giang Province (~9° N,
105° E). In Cambodia, it is present in Tonle Sap, and likely
occurs in the lower elevations of the Chao Phraya and Me-
kong river drainage systems. It is unclear if this species inhab-
its Myanmar, which H. semizonata is native to. Type locality:
Thailand, Nakhon Ratchasima Province, Wang Nam Khieo,
Udon Sap (subdistrict), Ban Badan Reservoir (14°31’04” N,
101°58’25” E) (Fig. 8.95).
Identification. Homalopsis mereljcoxi has a single loreal
contacting upper labials 1–4; scale rows at mid-body 40–49,
reduced to 30 or more posteriorly; two postoculars plus a
Figure 8.94. One of the syn- postsubocular; and ventral counts are usually greater than
types of Homalopsis hardwickii 165. Homalopsis buccata has 33–40 dorsal scale rows at mid-
BMNH 111.18.1b Photography body, reduced to less than 30 posteriorly; one postocular
by Christinia Cumberbatch. plus a postsubocular; ventral count less than 166. Homalopsis

169
 8. The Homalopsidae

hardwickii has a divided loreal; Homalopsis nigroventra-

lis has upper labials 1–3 contacting loreal; 35–39 dorsal
scale rows at mid-body, reduced to less than 30 posteri-
orly; reverse color pattern on the venter (dark olive-gray
with white spots). Homalopsis semizonata has a divided
or fragmented loreal contacting upper labials 1–4 or 1–5;
three prefrontals; one postocular and one postsubocular.
Size. Males: SVL: 404–914 mm; Average Tail: 223.72 mm
Figure 8.95. The distribution of (tail 30–44% of SVL); Females: SVL: 400–973 mm; Aver-
Homalopsis merlejcoxi. age Tail: 181.24 mm (tail 25–34% of SVL) (Figure 8.96).
Natural History. Habitat in Thailand includes small
reservoirs, ditches, ponds, streams, and shallow wetlands
(Karns et al., 2005, 2010b). Snakes are taken as bycatch in gill nets; one specimen from Mur-
phy et al. (2012c) was hand-collected from a pond at night while it floated near the surface
in an ambush posture with its head and forebody submerged. They have also been found in
similar habitats in Cambodia: streams, rivers, irrigation canals, marshes, reservoirs, lakes,
riverbanks Saint Girons (1972). It rests in burrows or crevices in the bank during the day
and has been reported to move very little while on land (Saint Girons, 1972). Individuals re-
leased into shallow water escaped without hesitation but placed in a river with steep banks,
the snakes immediately sought refuge along the shore. In a population depletion experi-
ment at Tonlé Sap (Brooks et al., 2007), a density of one specimen per 1718 m2 was calculat-
ed. Only fish have been found in the gut of H. mereljcoxi. About 61.5% of males and 37.5%
of females contained fish as prey in Karns et al (2010). Five families of fish are confirmed as
prey items in a study of 700+ specimens: Anabantidae 57%, Channidae 7.5%, Cyprinidae
2.5%, Mastacembilidae 3%, Osphronemidae 31% (Brooks et al., 2009); fish consumed were

Figure 8.96. Jack’s Watersnake, Homalopsis merlejcoxi. Eastern Thailand

Photography byJCM.

170
Aquatic Snakes

usually less than 10% of the predator’s mass. Predators likely include Cylindrophis jodiae (for-
merly C. ruffus) (Smith, 1914a; Brooks et al., 2007). Females start vitellogenesis in November
and mating occurs in December or early January; ovulation occurs in February (Murphy et
al., 2012c). Six females contain 13–33 large ovarian follicles or recently ovulated eggs in Feb-
ruary. Gestation lasts until May (Karns et al., 2005).

Mekong Watersnake
Homalopsis nigroventralis Deuve, 1970
Distribution and Habitat. This species is known from the
Mekong River Valley from Laos and Thailand to at least Cambo-
dia. It has been reported from the Mekong River in Kratié Prov-
ince, Cambodia and Koh Kapeung and the Koh Khlap Islands.
Type locality: Ngum River Valley, Laos (Figure 8.100)
Figure 8.97. The distribu-
Identification. Upper labials 1–3 contacting the loreal; two
tion of Homalopsis nigroven-
tralis. prefrontals; 35–39 dorsal scale rows at midbody, reduced to 30
or less; one or two postocular plus one postsubocular; 10–12 up-
per labials; 159–167 ventrals; reverse color pattern on the venter
(dark olive-gray with white spots). All other Homalopsis have upper labials 1–4 contacting

Figure 8.98. Mekong Watersnake, Homalopsis nigroventralis. Thailand.

Photography by JCM.
171
 8. The Homalopsidae

loreal and a ventral pattern that is light with dark spots. Size. Males: Maximum SVL: 828 mm;
Maximum Tail: 204 mm (tail 24.6% of SVL); Females: Maximum SVL: 820 mm; Maximum
Tail: 171 mm (damaged tail; tail 20.8% of SVL) (Figure 8.101).
Natural History. Homalopsis nigroventralis utilizes aquatic habitats, and, unlike the other
members of this genus, may use streams with a moderate current and sand and rock substrate,
compared to the slow moving or stagnant water with mud substrates that other Homalopsis
utilize (Stuart et al., 2006; Murphy et al., 2012c). The streams and localities where this species
have been found have closed or open canopies, and some had grass-lined banks (Stuart et al.,
2006). All these localities are peripheral to the Mekong. Despite this, they have been collected
on mud substrates too (Bezuijen et al., 2009). It also occurs in bamboo mixed with evergreen
forest between 100 and 170 m ASL in eastern Cambodia (Stuart et al., 2006). Specimens have
been collected during the day and are sometimes caught in gillnets as bycatch (Stuart et al.,
2006). Deuve (1970) noted that H. buccata and H, nigroventralis never occurred together.
Bernstein et al. (2021; in review), using phylogenetics methods with molecular data, showed
that H. nigroventralis is strongly supported as the sister to all other Homalopsis.

Myanmar Watersnake
Homalopsis semizonata Blyth, 1855
Distribution and Habitat. This species is known from
Kawkareik, Twante near Rangoon, Mottama, and the Ir-
rawaddy Delta, Myanmar. It also occurs at Kawkareik on
east side of Gulf of Martaban. It extends southward along the
coast to Phuket Province in southwestern Thailand (Frith,
1977). It may extend along the Andaman Sea’s coastal plain
in southern Myanmar and along the Andaman coast of pen-
Figure 8.99. The distribution of
insular Thailand. Type locality: Mottama (formerly Marta-
Homalopsis semizonata.
ban), Myanmar (Figure 8.99).

Figure 8.100. Myanmar Watersnake, Homalopsis semizonata.

Photograph from (A) Jens Vindum, (B) B. Murthy (C-D) JCM.

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Aquatic Snakes

Identification. This species is distinguished from all other members of the genus by a
prefrontal composed of three scales, loreal scale divided into 2 –6 smaller scales, upper labi-
als 1–5 contacting loreal scales. All other Homalopsis have two prefrontals, and upper labials
1–4 at the loreal. The three prefrontal scales, however, may be variable in different popula-
tions; a series from Ranong, Thailand was shown to have two or three prefrontals, and some-
times even four (Murphy et al., 2012c [University of Kansas Biodiversity Institute & Natural
History Museum specimen voucher KU 92453; Pauwels and Sumontha, 2016). Size. Males:
SVL: 562 mm; Tail: 166 mm (tail 29.4% of SVL); Females: SVL: 480 – 629 mm; Tail: 136–177
(tail ~28% of SVL); Neonates: SVL: 54 mm; Tail: 170 mm (tail 31.8% of SVL) (Figure 8.100).
Natural History. Little is known about Homalopsis semizonata. They have been collected
at night during rain near a dam at the Suwansiri waterfall in the Kraburi District of Thai-
land (Pauwels and Sumontha, 2016). In Pauwels and Sumontha (2016), a specimen had nine
babies in mid-June. They also two young individuals (one male, sex of second specimen
not mentioned) on Phuket Island (Phuket Province, southeastern Thailand) actively forag-
ing in a small stream near a waterfall in mid-September and mid-October. The young male
was found syntopic with the diploglossid Limnonectes blythii. The unsexed individual was
catching a channid fish (Channa gachua) and found syntopic with the geomydid turtle Sie-
benrockiella crassicollis. It is possible that H. semizonata may be a population of H. buccata, as
phylogenies using genomic data recovers the former embedded within the latter (Bernstein
et al. 2023a), but more sampling is needed to confirm this. Because observations are limited,
it is uncertain if H. semizonata is piscivorous and nocturnal, or if day encounters and obser-
vations of other prey items have just yet to be confirmed.

Lake Towuti Paddle-tailed Water Snake

Hypsiscopus indonesiensis Hamidy et al. 2023

Distribution and Habitat. The species is known only from Towuti Lake, South Su-
lawesi. Although Matano Lake, Towuti Lake, and Mahalona lakes are nearby and
connected, Hypsiscopus indonesiensis is only recorded from Lake Towuti Lake. The
Type Locality is Lengkona Bay, Towuti Lake, South Sulawesi. (no map).
Identification. small body size (SVL <700 mm), internasal(s) do not contact lo-

Figure 8.101. Hypsiscopus indonesiensis. A. Dorsal view of head. B. Narrow ventrals C. Laterally compress-
wed tail Photography by JCM.

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 8. The Homalopsidae

real, upper labials 2–3 or 2–4 usually contact with loreal, lower labials 1–5 usually
contact the anterior chin shields, relatively short tails (10–17 % of SVL) (Murphy &
Voris, 2014) and molecularly nested in the genus Hypsiscopus (Bernstein et al., 2021;
Fukuyama et al., 2022). Hypsiscopus indonesiensis is diagnosable from all congeners
by having a laterally compressed tail, a high number of scale rows (24–25 in the neck,
25–27 at mid-body, 20–22 near the vent), a high number of ventral scales (152–159),
the blotched pattern on the dorsum and mottled pattern on the ventral surface. Nat-
ural History. Highy aquatic. Poorly known.

Lake Matanna Mud Snake

Hypsiscopus matannensis (Boulenger, 1897)
Distribution and Habitat. Type locality: Lake Matanna, South-eastern Celebes (=Sulawe-
si). Endemic to Sulawesi. Known from Lake Matana (~02°27’ S, 121°12’ E) and near Raha
(~04°51’ S, 122° 43’ E), on Muna Island in southeast Sulawesi
(Figure 8.102).
Identification. Scale at mid-body in 21 rows; single posto-
cular; and divided internasal separate it from its congener, H.
plumbeus. Size. Males: SVL: 369–392 mm; Tail: 64–71 mm (tail
17.1–18.1% of SVL). Females: SVL: 387–566; 65–78 mm (tail
Figure 8.102. The distribu- 16–16.7% of SVL).
tion of Hypsiscopus matan- Natural History. Hypsiscopus matannensis is another spe-
nensis. cies of homalopsid that is poorly known and published on.
The second specimen ever discovered was collected from a
fishpond about 200 m from the nearest river (Iskandar, 1979).
All other aspects of its natural history have yet to be discovered.

Murphy’s Mud Snake

Hypsiscopus murphyi Bernstein et al. 2022
Distribution and Habitat. It inhabits sluggish streams,
ponds, rice fields, and marshlands at elevations from sea level
up to at least 1200 m (Mell, 1922; Deuve, 1970; Murphy, 2007).
It is present in Taiwan and southern China (Zhejiang Prov-
ince) at its northernmost limit extending southward into Chi-
na (including Hainan Island), into Vietnam, Laos, Cambodia,
and Thailand. Sampling by Bernstein et al. (2022) suggests the
southern limit of this species is in Phetchaburi, Thailand. Spec-
imens from adjacent and nearby islands will need to be identi-
fied pending future molecular and morphological comparisons
(e.g., Andaman Islands and other parts of India and Myanmar
Type locality: Laos, Khammouan Province, Gnommalath
Figure 8.103. Hypsicopus District, Ban Phak Phoung, 17.581918N, 105.219978E, 167m
matannensis. Photograsphy elev., collected in water in a rice paddy near karst forest at 2110
by JCM. h, Bryan L. Stuart, Niane Sivongxay, Sengvilay Seateun, and
Monekham Davanhkham, 11 July 2014.
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Aquatic Snakes

Identification. Hypsiscopus
murphyi can be distinguished from
its congeners by having a high-
er range of ventral scales which is
nearly non-overlapping with that
of H. plumbeus (113–123), a low-
er minimum number of subcaudal
scales than H. plumbeus(30–44), a
lower range of subcaudals than H.
matannensis(43–48), and a lower
number of scales at mid-body than
[Link] (21 DSR at mid-
body)
Natural History. It inhabits
sluggish streams, ponds, rice fields,
and marshlands at elevations from
sea level up to at least 1200 m (Mell,
1922; Deuve, 1970; Murphy, 2007).
However, it may wander some dis-
tance from the water. It has been
found beneath logs, especially
during the dry season (Saint Gi-
rons, 1972).One of the paratypes
was found during the day (1420
Figure 8.104. Hypsiscopus murphyi from Hong Kong. h) two meters above the ground
Photography by Artur Tomaszeck. inside a rotted, hollow, termite-in-
fested vertical tree in a gallery ev-
ergreen forest near a stream. The
species is nocturnal or cathemeral
(Murphy, 2007). Hypsiscopus murphyi is likely syntopic with its congener, H. plumbeus, in
regions south of the Khorat Plateau in central and eastern Thailand and western Cambodia,
up to latitudes of approximately 12.48N. Like other members of the genus, this species feeds
on fishes, frogs, and sometimes crustaceans (Schmidt, 1927; Gressitt, 1941). Like all hom-
alopsids it is ovoviviparous, containing 2–18 young (Murphy, 2007); averages of nine young
per litter have been reported (Cox,1991). Two examined specimens of H. murphyi (ROM
30818 and ROM 30933) in this study contained seven and nine ova, respectively. The venom
toxicity of this species is not considered dangerous to humans, usually only causing localized
effects (if any) such as burning sensations and local swelling (Karsen, 1986)
Hypsiscopus has three species. Hypsiscopus plumbeus was considered to inhabit nearly all
of Southeast Asia, including Sulawesi. Previous research recovered H. plumbeus as paraphy-
letic with respect to the Sulawesi-endemic [Link] (Bernstein et al., 2021), a finding
confirmed here with additional sampling. Bernstein et al. (2022) molecular and morpho-
logical analyses show evidence that the populations around and north of central Thailand’s
Khorat Plateau represent a distinct lineage that warrants species-level recognition. Wile they
did not use genetic distances to diagnose this species, their genetic divergences (mitochon-
drial genes: 4.75–13.53%; nuclear genes: 0–2.48%) exceed that of species-level distances seen
in studies on homalopsids (Kohler et al., 2021) and other snakes (Ruane et al., 2018).

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Aquatic Snakes

Plumbeous Mud Snake

Hypsiscopus plumbeus (Boie, 1827)
Distribution and Habitat. Previously considered a
widespread species. Bernstein et al. split it into two spe-
cies. H. plumbeus now occupies the southern portion
of the previously widespread species. range in south-
Figure 8.105. The combined distri- ern Indochina, peninsular Thailand and Malaysia It is
bution of Hypsiscopus murphyi and
present on many, if not all, the Indonesian islands and
H. plumbeus. The blue line roughly
extends its distribution eastward to Sulawesi. One spec-
marks the boundary between the
two species. H. murphyi is to the imen (American Museum of Natural History specimen
north line and H. plumbeus is to voucher AMNH 3840) is from the Moluccan Islands,
the south. The two species are likely which would extend the eastern edge of the range if the
sympatric in the vicinity of the line locality data is correct. Type locality: Java (Figure 8.105).
Identification. Scale rows at mid-body 19, ventrals
119–142; upper labials 2–3 or
2–4 contact loreal; four or five
lower labials contact anterior
chin shields. Dorsum a uni-
form grey or black. Its con-
gener, H. matannensis has 21
scale rows at mid-body. Size.
Males: SVL: 152–420 mm;
Tail: 24–59 mm (tail 11.9–
19.1% of SVL); Females: SVL:
190–650 mm; Tail: 29–70 mm
(tail 10–17.6% of SVL). Neo-
nates: 122–160 mm (Figure
8.106).
Natural History. This is
a semi-aquatic snake and the
most terrestrial of the fanged
homalopsids. It is associat-
ed with wallows, marshes,
ditches, stream, pond banks
in the mud-root tangle 10–
20 cm below surface debris,
both day and night, and oc-
curs from sea level to at least
1200 m (Deuve, 1970; Mell,
Figure 8.106. Hypsiscopus plumbeus. A. is from Sabah, Malaysia 1922; Smith, 1943). The spe-
(island of Borneo). Photography by DRK. B is a female from cies is particularly common in
Lake Songhkla, southern Thailand. Photography by JCM. the rice paddies and drainage
ditches (Bien Hoa, Vietnam;
Main, 1970), and it will move over land. Accounts from Annandale and Robinson (Bou-
lenger, [1903]) report it is aquatic, but is found under dead trees at some distance from wa-
ter; this has also been stated by Saint Girons (1972), mentioning that prefers rice fields and
marshland, but is found away from water in the dry season. Because of common encounters

176
 8. The Homalopsidae

in ride paddies and agricultural areas, this snake is also called the ‘rice paddy snake.’ This
species is considered nocturnal (Karsen et al., 1986; Murphy, 2007a). They have also been
collected in stagnant ponds in Thailand (Murphy, 2007a). Snakes that were using a nearby
stream were more mobile than those using the wallow microhabitats. It also feeds on dead
fish. Their diet includes fish and amphibians (adults and larvae). Confirmed prey items in-
clude: the four-lined tree frog Polypedates leucomystax (Rhacophoridae), alpine cricket frog
Rana [=Fejervarya] limnocharis (Ranidae), and walking catfish Claris teysmanni (=Clarias
teijsmanni; Clariidae) (Pope, 1935; Voris and Karns, 1996; Murphy, 2007a). Confirmed pred-
ators include Cylindrophis ruffus (though, this record may represent C. jodiae), Bungarus
multicinctus (Mao, 1970; Slowinski, 1994), and Naja naja (Mao, 1970). While not confirmed,
the root tangle that H. plumbeus resides in is likely very safe from predatory birds and some
fish, but others, such as snakeheads (Ophicephalidae) likely feed on H. plumbeus. Gravid fe-
males ranged in size from 282–480 mm total length and litter sizes range from 2–18 neonates
(Murphy, 2007a). In Cambodia, neonates are recorded to be born in the spring (Saint Girons,
1972; Saint Girons and Pfeffer, 1972). In Laos, neonates have bene found throughout the
rainy season, with young being born from April through August (Deuve, 1970). Parturition
is finished by August. These snakes are known to use sidewinding and erratic ‘jumping’ to
escape (Pope, 1935; Kuntz, 1963), and have been described as having either hiding behaviors
by burying its head (Batchelor, 1963) or a tendency to flee when encountered in water, or
relentless striking and biting if provoked out of water (Kuntz, 1963; Campden-Main, 1970).
High levels of molecular divergence have been found in this group, and research currently
underway (Bernstein et al. 2023b) will likely increase the known diversity of this genus. The
venom of H. plumbeus is mild, not posing a threat to humans (Karsen et al., 1986). Bites from
this species have been followed by immediate burning sensations and localized swelling (i.e.,
no systemic effects) and been compared to that of a bee sting (Daryl Karns, personal com-
munication [Murphy, 2007]).

Pahang Mud Snake

Kualatahan pahangensis (Tweedie, 1946)
Distribution and Habitat. Kualatahan pahangensis is known from the vicinity of the type
locality Kuala Tahan, River Tembeling, Pahang, peninsular Malaysia. Its range was expanded
to include Hulu Terengganu and the Nenasi Forest Reserve in Pahang (Chan and Ahmad,
2009). Type locality: Kuala Tahan, River Tembeling, Pahang, between 500- and 1000-feet ASL
(152–308 m) (Fig.8.107).
Identification. Has 25 scale rows at midbody, eight up-
per labials, upper labials 2–3 contact loreal, dorsal scales lan-
ceolate; two pairs of chin shields five lower labials contacting
first pair. Dark lateral stripe on the outer edge of ventrals row
one, light stripe on rows 2–4. It may be confused with Gyio-
Figure 8.107. The distribution phis, which has 25 scale rows at midbody, but upper labials
of Kualatahan pahangensis. 1–3 contact the loreal; dorsal scales, particularly those in the
first row, are ovate. Both Homalophis and Phytolopsis have
25 scale rows at midbody but have more than ten upper labials, and some are horizontally
divided. Size. Males: SVL: 220–300 mm; Tail: 40–69 mm (tail 18.2–23% of SVL); Females:
SVL: 392 mm; Tail: 68.5 mm (tail 17.5% of SVL) (Fig.8.108).
Natural History. Kualatahan pahangensis is known only from very few localities in Ma-
laysia, and all natural history is based on a few specimens. Chan and Ahmad (2009) report

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Aquatic Snakes

Figure 8.108. Kualatahan pahangensis

Photography by Chan Kin Onn.

that this species inhabits primary or undisturbed secondary forests, utilizing decaying leaves
in stagnant portions of small, slowing moving streams. Charlton (2020) reports it from both
primary and secondary forests up to 300 m asl, residing in slow-moving streams.

Indus Mud Snake

Mintonophis pakistanicus (Mertens, 1959)
Distribution and Habitat. Extreme western edge of the family’s distribution in the Indus
River delta of Pakistan. Type locality: Jati, Sind, West Pakistan (Fig. 8.109).
Identification. Smooth dorsal scales, in 29 rows at mid-body; 153–162 ventrals excep-
tionally narrow on the anterior body, widening posteriorly; 8–9 upper labials, 4 or 4–5 in
orbit; internasal divided contact loreal; loreal contacts upper labials 1–3; lower labials 11–12;
chin shields absent (or exceptionally small), replaced by numerous small scales; tail excep-
tionally long. The narrow ventrals, absent or tiny chin shields, and long-tail suggest this snake
is highly derived and convergent with Erpeton tentaculatum. Males: SVL: 513 mm; Tail: 207
mm (tail 32–41% of SVL). Females: SVL: 779 mm; Tail: 208 mm (tail 21–33% of SVL) (Fig.
8.108).
Natural History. This species, while poorly known outside a few specimens, is known
to occupy large shallow pons near channels of the Indus River,
but not in the stream itself (Minton, 1966). Some of the sites in
this area are brackish water habitats, and one used to be a sea-
port (Murphy, 2007a). They are reported to be diurnal, almost
entirely aquatic, and shy in nature. Personal communication with
collectors also mentions that they have been collected from late
Figure 8.108. The distri-
April through November, and will bury themselves in the mud
bution of Mintonophis
pakistanicus. the edges of their habitats (e.g., ponds) before the onset of cool
weather (Murphy, 2007a). These habitat notes have also been cited
by Khan (2002), stating that M. pakistanicus is almost complete-

178
 8. The Homalopsidae

ly aquatic (moving poorly on

dry land), will bury in the mud
to hibernate, and inhabits back-
waters and ponds with emersed
coastal vegetation. This study
contains the only diet records of
this species, finding that they eat
fish, frogs, tadpoles, and aquatic
arthropods, and also confirms
previous reports on their shy and
non-aggressive nature. ‘Bernstein
et al. (2023a) uses mitochondri-
al DNA to recover Mintonophis
as sister to Dieurostus, both of
Figure 8.109. Indus Mud Snake, Mintonophis pakistanicus. which are sister to Ferania. The
Photography by Petrova and [Link]. morphology suggests a highly
aquatic snake with a prehensile
tail (Murphy, 2007a). Gyi (1970) and Murphy (2007a) noted that two female museum spec-
imens, AMNH 93154 (708 mm SVL) and CAS 99964 (500 mm SVL) contained 16 and 8
embryos, respectively.

Reuss’ Mud Snake

Miralia alternans (Reuss, 1834)
Distribution and Habitat. Greater Sunda Islands (Sumatra, Java, Borneo) and smaller
islands of Bangka and Beliting. Type locality: Java (Fig. 8.110).
Identification. Smooth scales, nasals in contact; dorsal scales in 19 rows at mid-body;
a tiny preocular scale; lower postocular does not extend below eye; fourth and fifth upper
labials enter orbit; the ventrals number 120–143; eight upper labials; nasal cleft contacts
second upper labial. Superficially like Raclitia indicia (but see Natural History, below). Size.
Males: SVL: 380–474 mm; Tail: 50 mm (tail 13.1% of SVL); Females: SVL: 320–490 mm;
Tail: 45–50 mm (tail 13.2–14.1% of SVL); Neonates: SVL: 123–126 mm; Tail: 14–19 mm |
Average SVL: 124.4 mm; Average Tail: 16.2 mm (Fig. 8.111)
Natural History. Miralia alternans is a rare species that is not encountered often, thus
little is known about its Natural History. It has been found in rivers, specifically the Kali
Cakung in Jakarta, Indonesia (Iskandar, 1987). Confirmed species that predate on M. al-
ternans include Cylindrophis ruffus. The small head and short tail of this species suggest it
have burrowing behavior. Clutch sizes in this species range from 6–16 (average = 9), and
litter sizes may increase with female body size; the smallest
gravid female recorded was 403 mm SVL (Bergman, 1960).
Miralia alternans is superficially like Raclitia indica. However,
the extremely gracile body of R. indica, more robust body of
M. alternans, the difference of more than 30 ventral between
the two species, and the exceptionally microcephalic condi-
Figure 8.110. The distribution of tion of R. indica suggest the two snakes have been separated
Miralia alternans.
from each other for some time and have distinct evolution-
ary trajectories. Mitochondrial DNA (Bernstein et al. 2023a)
suggests that Miralia may be sister to Myrrophis, but high

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Aquatic Snakes

Figure 8.111 .Reuss’ Mud Snake, Miralia alternans.

Photography by Rendra-Wahyudi.
levels of sampling are needed to confirm this.

Thanlyin Mud Snake

Myanophis thanlyinensis Köhler, et al. 2021
Distribution and Habitat. Only known from the Type locality: Specimen found near
East Yangon University. Type locality: Thanlyin, Yangon, Myanmar (Figure 8.112).
Identification. Differs from all homalopsids by the following combination of char-
acters: (1) dorsal scales smooth, row formula 21–21–19 or
21–21–17; (2) tail short, ratio tail length/SVL 0.185–0.204
in males, 0.160–0.167 in females; (3) nasal scales separated;
(4) 125–126 ventral scales in males, 120–122 in females; (5)
38–39 subcaudal scales in males, 32–34 in females; and (6)
hemipenis bilobed. It differs from Myrrophis by having 125–
Figure 8.112. Distribution of 126 ventral scales in males, 120–122 in females (vs. 137–162,
Myanophis thanlyinensis. 137–164, respectively), and 38–39 subcaudal scales in males,
32–34 in females (vs. 39–55, 37–52, respectively). Myanophis
thanlyinensis differs from Gyiophis by lacking dark blotches along flank (vs. blotches pres-
ent), and by having 21 dorsal scales rows at midbody (vs. 25 scale rows). Additionally, the
bilobed hemipenes present in M. thanlyinensis distinguish it from both Myrrophis and Gyio-
phis. Size. Males: SV:260–335 mm; Tail: 53–62 mm (tail 18.5–20.38% of SVL); Females: SVL:
300–362 mm; Tail: 50–58 mm (tail 16–16.7% of SVL) (Figure 8.113).
Natural History. The natural history of this monotypic genus is only known from the
holotype and three paratypes published in Köhler et al. (2021). This species only known
from open grassland that is flooded during the rainy season. Its discovery adjacent to East
Yangon University may indicate this taxon can thrive in disturbed habitats close to human
settlement. Myanophis thanlyinensis is likely nocturnal and has been found in shallow wa-
ters (<50 cm depth). A phylogenomic study (Bernstein et al. 2023a) places Myanophis as sis-
ter to Gyiophis. Other snakes syntopic with M. thanlyinensis are Enhydris enhydris, Gyiophis

180
 8. The Homalopsidae

Figure 8.113. Thanlyin Mud Snake, Myanophis thanlyinensis.

Photography by Günther Kohler.
vorisi, Amphiesma stolatum, Xenochrophis bellulus, and X. piscator.

Karns’s Mangrove Snake

Myron karnsi Murphy, 2011
Distribution and Habitat. Only known from the type locality of Indonesia Aru. Type lo-
cality: Selrutti, Kobroor, Indonesia Aru (~5°46’ S, 134°31’ E) (Fig. 8.114).
Identification. Myron karnsi is a melanistic species with narrow yellow cross bands, with
21 scale rows at mid-body; seven upper labials; single preocu-
lar, semi-divided nasal, and a black dorsum with narrow yel-
low cross bands separate this species from other two species
of Myron. Size. Males: no data. Females: SVL: 333 mm; Tail:
64 mm (tail 19% of SVL).(Figure 8.114)
Natural History. The natural history of M. karnsi is un-
known but is likely similar to its congeners.
Figure 8.114. The distribution
of Myron karnsi.
Resetar’s Mangrove Snake
Myron resetari Murphy, 2011
Distribution. Myron resetari is known only from the type
locality of Broome, Western Australia. However, it may be
more widespread. It is likely that this snake also inhabits the
Roebuck Bay area of the Dampier Peninsula (Murphy, 2011).
Type locality: Broome, Western Australia (~17°58’ S, 122°14’
E) (Fig. 8.116).
Figure 8.115. Myron karnsi. Identification. This species has 19 scale rows on the neck
Phototography by JCM. and at mid-body; one preocular; large, plate-like occipital
scales that are posterior and lateral to parietals. Myron resetari
can be distinguished from M. richardsonii by a lower scale row count on the neck and at mid
body (19 vs. 21); one preocular (two in M. richardsonii); largest upper labial is the fifth com-
pared with the largest upper labial of six or seven in M. richardsonii. Myron resetari can be
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Aquatic Snakes

distinguished from M. karnsi by a lower number of scale rows at

mid-body (19 vs. 21) a blotched pattern on a gray ground color
(as opposed to yellow bands on black ground color in M. karn-
si). Size. Males: SVL: 330 mm; Tail: 65 mm (tail 19.7% of SVL).
Females: SVL: 246 mm; Tail: 44 mm (tail 17.9% of SVL) (Fig.
8.117).
Natural History. Similar to M. karnsi, little is known about
Figure 8.116. The distribution this species. It inhabits mangroves and mudflats (Murphy, 2011).
of Myron resetari.

Richardson’s Mangrove Snake

Myron richardsonii Gray, 1849
Distribution and Habitat. Found on coastal northern Australia, from the Gulf of Car-
pentaria to Western Australia and northward to the south coast of New Guinea. It is also

Figure 8.117. Resetar’s Mangrove Snake, Myron resetari.

Photography by Gary Rethus.

present on the Aru Islands of Indonesia (Roux, 1910). Type locality: North Western Australia.
Restricted to Buffalo Creek, Northern Territory, Australia (~12°40’ S, 131°29’ E) by Murphy
(2011) (Fig. 8.118).
Identification. Has 21 scale rows on the neck and at mid-body, reduced to 17 or 19 rows
near vent; posterior dorsal scale rows above row six prominently
keeled in males; two preocular; upper labials usually number 8
or 9; lower labials 8–10; a dorsal pattern of 35–48 blotches. The
presence of 21 scale rows, two preoculars distinguish it from M.
resetari. The 8 to 10 upper labials and blotched pattern separate
it from M. karnsi, which is melanistic with narrow yellow cross
Figure 8.118 The distribu- bands and has fewer upper labials. Size. Males: SVL: 191–372;
tion of Myron richardsonii. Tail: 32–64 mm (tail 16.8–19% of SVL); Females: SVL: 255–370;
Maximum Tail: 59 mm (tail 16–17% of SVL) (Fig. 8.119).
Natural History. The natural history of M. richard-

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 8. The Homalopsidae

Figure 8.119. Richardson’s Mangrove Snake, Myron richardsonii.

Photography by Jacob Loyacano.
sonii is perhaps the most well-known out of any member from this genus. This species (as
well as the other Myron) are considered dwarf homalopsids (Murphy, 2011), not exceed-
ing 436 mm in total length; most other homalopsids exceed one meter in length (Murphy,
2007a). This species uses coastal mangroves, nearby mud flats, tidal rivers, and other aquatic
habitats. Similar to Cerberus, Cantoria, Fordonia, it inhabits the intertidal burrow system and
will bury itself into muddy substrate (Glauert, 1950; Gow, 1989, Worrell, 1963). Confirmed
prey items include gobies (Murphy, 2007a), but crustaceans have also been mentioned as prey
items (Worrell, 1963; Gow, 1989). One observation reports finding a M. richardsonii with a
nudibranch in its mouth (Nobbs and Blamires, 2004). Shine and Schwaner (1985) observed it
using constriction to subdue small fish, and Gyi (1970) suggested that the long, anterior teeth
of this species may assist in holding onto gobies and gastropods covered in slippery mucous.
Confirmed predators are Crocodylus porosus (Taylor, 1979). Litter sizes of this species are
reported to be 6–8 (Gow, 1989; Shine, 1991). The interspecific, evolutionary relationships
within Myron are currently unknown, but Gyi (1970) hypothesized that Myron and Heurnia
ventromaculata formed a group. Bernstein et al. (2021) and Alfaro et al. (2008), using molec-
ular data, both found strong support for the genus Myron to be sister to Pseudoferania polyl-
epis and genomic data also supports this sister relationship (Bernstein et al. 2023a).

Bennet’s Mangrove Snake

Myrrophis bennettii (Gray, 1842)
Distribution and Habitat. Myrrophis bennettii is endemic to China’s south coast from
Hong Kong, south and west to Hainan. Gyi (1970) examined a specimen from Java, from the
Zoölogisch Museum (University of Amsterdam; ZMA, no voucher number), but this is likely
in error according to Murphy (2007a). Type locality: China (Figure 8.120).

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Aquatic Snakes

Identification. Myrrophis bennettii has dorsal scales in 21 rows at mid-body, reduced to

17 or 15 in front of the vent; an internasal scale that does not contact the loreal; and an upper
postocular larger than lower postocular. It is most often confused with M. chinensis, which
has 23 rows of scales at midbody, 19 rows near the vent. Size.
Males: Maximum SVL: 560 mm; Tail: 120 mm (tail 21–22% of
SVL); Females: Maximum SVL: 755 mm; Tail: 96 mm (dam-
aged tail) (tail 21–24% of SVL) (Figure 8.121).
Natural History. Myrrophis bennettii inhabits saltwater
marshes and estuaries, and occasionally climbs onto shrubs,
Figure 8.120. The distribution behavior not typical for homalopsid snakes (Murphy, 2007a).
of Myrrophis bennettii. It is unclear if this behavior is an attempt at basking, or if the
snakes use branches at high tide and were left on a branch as
the tide receded. They have been reported as common in tidal
marshes in and around mangroves (Deep Bay) and partially active in the day, but still may

Figure 8.121. Bennet’s Mangrove Snake, Myrrophis bennettii

Photography by Adam Francis.

be nocturnal (Karsten et al., 1986). Other observations also mention their ability to climb is
particularly good (Karsten et al., 1986). They are not considered a rare snake and have bene
found in the sea around Hainan’s coast, which may indicate they get washed out to sea from
streams (Pope, 1935; Murphy, 2007a); this is possible as they have been found in streams at
points that drain into the sea (Romer, 1961). The Hong Kong population uses tidal marshes
(near Pak Hok Chau) along the shore at Deep Bay. Specimens have also been taken at the
mouth of the Yuen Long Creek (unofficial name) and by the shore on the Mong Tseng pen-
insula at Deep Bay (Murphy, 2007a). Using molecular data, Bernstein et al. (2021) found
strong support that Myrrophis is most closely related to Gyiophis salweenensis (the only spe-
cies of Gyiophis in the study), which together form a clade that is sister to a monophyletic En-
hydris. However, genomic data (Bernstein et al. 2023a) now supports that Myrrophis is sister
to Myanophis and Gyiophis; these three genera are all reciprocally monophyletic to Enhydris.

Chinese Mud Snake

Myrrophis chinensis (Gray, 1842)
Distribution and Habitat. Known from southern China, North Vietnam, Taiwan, Hong

184
 8. The Homalopsidae

Kong, Hainan. It may be present in northern Laos. Type locality: China (Fig. 8.122).
Identification. Myrrophis chinensis has 23 scale rows at midbody, reduced to 19 rows in
front of vent; internasal not usually in contact with loreal, upper postocular smaller than lower
postocular. Size. Males: SVL: 580 mm; Tail: 125 mm (tail 15–19% of SVL); Females: SVL: 710
mm; Tail: 100 mm (tail 12–15% of SVL); Neonates: 100–170 mm (Fig. 8.123).
Natural History. Myrrophis chinensis is a highly aquatic snake that inhabits lowlands
and is common between sea level and 200 m. Though, it as-
cends to considerable altitudes in northeastern Kiangsi and on
the Chungan plateau, the latter of which reaches an elevation of
1500 m (however, this is not to say M. chinensis is found at that
elevation). It has been recorded at 525 m elevation (Gangeku,
southwestern Fukien Province) and at 825 m (Hong San, south-
eastern Kiangsi Province) (Gressitt, 1941). Data accompanying
Figure 8.122. The distribu-
tion of Myrrophis chinensis. specimen from North Vietnam suggest it reaches 1100 m ASL
(Murphy, 2007a). This implies that this population may experi-
ence cold winter temperatures. It uses fishponds, sluggish streams, canals, and rice paddies;
many literature accounts have found this species associated with rice paddy ecosystems, thus,
agricultural practices may in part be responsible for the relatively high abundance widespread
altitudinal distribution of this species (Murphy, 2007a). The optimal time of activity for M.
chinensis is uncertain; Karsen et al. (1986) states that it may leave the water at night during
rain spells, but also reports it being active in water during the day. Kuntz (1963) also made
similar observations and records its presence in suburban Taipei, Taiwan during flooding.
They feed primarily on fish (anabantids [Anabas scandens (=Perca scandens): Chang and
Fang, 1931], cyprinids [Carassius sp., sp.: Pope, 1935], xenocypridids [Erythroculter aokii(?),
Culter alburnus: Pope and Granger, 1929; Pope, 1935], osphronemids [Macropodus opercularis
(=baviensis), Macropodus viridiauratus: Pope and Granger, 1929; Pope, 1935) and according
to the literature may take amphibians on occasion (Gressitt 1940, 1941; Kunts, 1963). Mori
(1998) found this species will swallow food underwater, which is a common behavior for most
homalopsids.
Seven females from Taiwan contained 3–32 embryo/eggs (average=16.3; Murphy, 2007a).
The smallest SVL of a gravid female in Murphy (2007a) was 396 mm, the largest was 701
mm. However, another gravid female from Canton, China is 385 mm SVL. Pope (1929) com-
mented that females from higher latitudes give birth to larger litters. Parturition likely occurs
during the summer months, though gravid females in December have been found (Smith,
1914b). Smith (1914b) mentions that parturition may take place in April, though as to which
population this pertains to is uncertain. Kuntz (1963) remarked on the behavior of M. chin-
ensis, saying that at first, it is prone to striking upon the slightest irritation, but soon becomes
more docile. It may make false strikes with lateral and/or vertical jerks of the head, along with
quick vertical jumps. It is difficult to capture in the wild when disturbed since it tends to dis-
appear or submerge into the soft mud of the rice paddies. When handled, this snake is likely
to initiate a profuse flow of semi-fluid feces with a pungent odor (Murphy, 2007a). The venom
of this species may be more potent than other homalopsids, inducing headaches, nausea, and
pain (Karsen et al., 1986).

Dak Krong Mud Snake

Myrrophis dakkrongensis Nguyen et al. 2024
Distribution and Habitat. Known only from its type locality in Dak Glong District, Dak

185
Aquatic Snakes

Figure 8.123. Chinese Mud Snake, Myrrophis chinensis.

Photography by Artur Tomaszek.
Nong Province, Vietnam southern China and northern Vietnam. The holotype is an adult
male, collected from Quang Son, Dak Glong District, Dak Nong Province, Vietnam at an
elevation of 890 m ASL. The first specimens were collected in forest wetlands, including a
wetland in a rubber plantation. [No map or photo.]

Blackwater Mud Snake

Phytolopsis punctata Gray, 1849
Distribution and Habitat. Found in peat swamps in peninsular Malaysia, Singapore, Su-
matra, and Borneo (Kalimantan, Brunei). Type locality: India (in error) (Figure 8.123).
Identification. A large bodied snake, dorsal scales in 25–27 rows; upper labials number
11–13, those behind eye horizontally divided; lower labials 12–15; ventrals 148–160; subcau-
dals 36–48; upper labial 5 or 5–6 under orbit; no subocular scales; two pairs of chin shields,
first pair very wide. Readily distinguished from Homalophis by lower number of scales at
mid-body (29–33 in Homalophis), single supraocular (2 or 3 in Homalophis), and the ab-
sence of subocular scales. Size. Murphy (2007a) reports on sizes from the literature, but as
to which sex these size ranges pertain to is uncertain; it is likely inclusive of both males and
females. Murphy (2007a) reported that the largest specimen in that study was 665 mm SVL
and 95 mm tail. This exceeds previous measurements by Erns and Zug (1996) and David and
Vogel (1996), in which the maximum total length was 450 mm; the latter study found adults
averaged 340–400 mm. Murphy (2007a) comments that most of the museum specimens ex-
amined previously were juveniles. Males have tails that are 19–21% of SVL, and females have
tails that are 8.5–17.8% of SVL (Murphy, 2007a) (Figure 8.124).
Natural History. Phytolopsis punctata has been found in a variety of habitats. Museum
notes for peninsular Malaysian specimens from the Field Museum of Natural History (FMNH
250111–250112) mention that they were collected from ‘blackwater streams.’ Malaysian spec-
imens have been collected from peat swamp forests (collected by Shahrul Anuaroast) of Pon-

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 8. The Homalopsidae

species typcally burrows in the ground, possibly in aquat-

ic habitats, and may be vermivorous (Murphy, 2007a;
Murphy et al., 2011). They have been collected in small
forest streams at night (Klang, Selangor, Malaysia: Lim
and Karmudin, 1975; Lim et al., 1995). The population in
Singapore has been considered questionable by Murphy
Figure 8.124. The distribution of
(2007a), but recent records of a specimen found in a con-
Phytolopsis punctata. crete drain at the edge of a freshwater swamp forest from
Upper Seletar, Singapore confirm a population here (Law

Figure 8.124. Blackwater Mud Snake, Phytolopsis punctata.

Photography by Ing Sind.

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Aquatic Snakes

dak Tanjung Forest Reserve in Selama Perak. And, recently, they have been reported from
within and around the Nee Soon freshwater swamp forest of Singapore (Thomas et al., 2014;
Serin et al., 2019). The fragmented nature of these habitats and the high level of fish ende-
mism in the peat swamps of Southeast Asia add an interesting dimension to this species
complex, and to homalopsid diversity. The details of its lifestyle remain virtually unknown.
A female specimen examined by Murphy (2007a) contained 15 yolked follicles (Murphy,
2007a). Its diet includes fish, with walking catfish (Clariidae: Clarias leiacanthus) as con-
firmed prey (Tan et al., 2014).
Phytolopsis punctata has been recovered in molecular phylogenies with varying levels
of support: Alfaro et al. (2008) recovered P. punctata as sister to a clade with Cerberus, Hom
alopsis, Erpeton, Subsessor, Myron, Pseudoferania, Fordonia, Gerarda, Cantoria, and Bitia, but
Bernstein et al. (2021) recovered P. punctata as sister to Bitia hydroides. Using thousands of
nuclear markers, Bernstein et al. (in review) finds strong support that P. punctata is sister to
Raclitia indica.

Macleay’s Mud Snake

Pseudoferania polylepis (Fischer, 1886)
Distribution and Habitat. Found in Australia (coastal plain of Queensland, Northern
Territory (Groote Island)) and Papua New Guinea. Groote Island. Type locality: Fly River,
Western Province, Papua New Guinea (Figure 8.129).
Identification. Pseudoferania polylepis has parietals scales that are longer than the fron-
tal; loreal single. Dorsal scales in 21–27 rows (usually 23) at mid body; 137–162 ventral
scales; upper labials 7 – 9 (usually 8), with 4–5 or 5–6 under the orbit, upper labials 2 –3 or
2–4 contact loreal.
Internasal single or double, it may or may not contact the loreal. Size. Males: SVL: 380–
689 mm; Tail: 114 mm (tail 16.5–19.6% of SVL); Females: SVL: 44–870 mm; Tail: 107 mm
(tail 16.5–21.5% of SVL) (Figure 8.130).
Natural History. This taxon is the only freshwater homalopsid in the Austral-Papuan re-
gion except for Heurnia ventromaculata from Western Papua which does not have the nasal
scales in contact. They use slow moving or stagnant bodies of freshwater, such as lagoons,
billabongs, swamps, and creeks and take refuge in vegetation
near water, such may prefer dense pandanus root systems near
overhanging banks (Gow, 1989; Murphy, 2007a). Dunson and
Dunson (1973) found this species sensitive to salt water. Shine
et al. (2004) reports them from shallowly inundated areas of
Fogg Dam in Northern Territory, Australia; these areas are
Figure 8.129. The distribution usually dry for a majority of the year and were flooded for two
of Pseudoferania polylepis. weeks before their study took place. During seasonal flooding,
their movement expands. They report that while P. polylepis is
infrequent, the two weeks of these floods results in high num-
bers of this species at night, foraging for sleeping fish. The fish allows the snake to approach
closely and then flips upwards and out of the water when touched; experiments showed that
these snakes locate their prey primarily by visual cues, especially movement, rather than
by scent or waterborne vibrations (Shine et al., 2004). Strikes are elicited by tactile cues,
especially the splash of water as a fish leaps upwards. Confirmed prey includes decapods
(shrimp fragments; Macrobrachium sp.), frogs (captive specimens), and fishes (Eleodtridae,
Megalopidae (Megalops cyprinoides) (Shine, 1991). They are known to be eaten by various

188
 8. The Homalopsidae

Figure 8.130. Macleay’s Mud Snake, Pseudoferania polylepis.

Photography by David Nixon.
birds, such as white-belled sea eagles (Accipitridae: Haliaeetus leucogaster) and black-belled
storks (Ciconiidae: Ephippiorhynchus asiaticus) (Sergo and Shine, 2015), as well as long-
necked turtles (Chelidae: Chelonia rugosa; Shine, 1991). Junqueira-de-Azevedo et al. (2016)
found the venom of this species contained low amounts of snake venom metalloproteinase
(SVMP), three finger toxin (3FTx), cysteine rich secretory protein (CRISP), and C-type
lectin (CLT); they also contain veficolins, the venom protein family discovered in Cerberus
rynchops (Fry et al., 2012). Reports on the bite of P. polylepis on humans are asymptomat-
ic (Thompson, 1935), but frogs have been found to succumb quickly, with accompanying
discoloration and oedema at the site of the bite, and death in fish have been observed <2
minutes from the time of attack (Shine, 1991). Pseudoferania polylepis has consistently been
recovered, with strong support, as the sister group to Myron when using mitochondrial and
nuclear genes (Alfaro et al., 2008; Bernstein et al., 2021; Bernstein et al. 2023a).

Selangor Mud Snake

Raclitia indica Gray, 1842
Distribution and Habitat. Known from peninsular Malaysia (Perak, Pahang, Selangor)
and Singapore. Type locality: India (probably referring to the East Indies or, broadly speak-
ing, Southeast Asia. (Figure 8.131)
Description. Supraocular exceptionally large, plate-like, preocular exceptionally small;
supraocular and upper postocular fused; lower postocular extends under eye; nasal cleft
contacts first labial; 173–175 very wide ventrals; eye diameter less than width of supraocu-
lar; seven upper labials. Size. Males: SVL: 303–372 mm; Tail: 42–53 mm (tail 13.9–14.4% of
SVL); Females: Total Length: 490 mm (Figure 8.132)
Natural History. Raclitia indica has a small, narrow head, gracile body form, short
tail, and a relatively low dorsal scale count; based on these, it has been suggested that this

189
Aquatic Snakes

et al., 2020). Whether a population get here accidentally due to human transport is uncer-
tain. Raclitia indica is recovered in phylogenies using molecular data as either the sister to
Erpeton tentaculatum (Quah et al., 2018), to a clade containing
Bitia, Phytolopsis, Homalopsis, and Cerberus (concatenated ap-
proach; Bernstein et al., 2021), or sister to Dieurostus dussum-
ieri (coalescent approach; Bernstein et al., 2021). All of these
phylogenetic relationships have low support, and further, more
comprehensive studies must be conducted.
Figure 8.131. The distribu-
tion of Raclitia indica. Bocourt’s Mud Snake
Subsessor bocourti (Jan, 1865)
Distribution and Habitat. This species is found in southern Thailand, peninsular Ma-
laysia, Cambodia, Vietnam, and Laos. It may have been introduced into southern China
where it is captive raised. Type locality: Bangkok, Thailand (Figure 133).
Identification. Subsessor bocourti is considered a massive, stout-bodied snake, exceed-

Figure 8.132. Selengor Mud Snake, Raclitia indica.

Photography by Evan Quah.

ing a meter in length; dorsal scale rows in 27–29 rows; ventrals less than 136; chin shields
petal shaped, some upper labials behind the eye are horizontally divided. Probably the larg-
est, extant homalopsid when considering weight and length; 27 scale rows at midbody; di-
vided upper labials behind the eye: 135 ventral scales, or fewer; an internasal that is single,
and may make nrrow contact with the loreal scales. Size: Males. SVL: 646 mm; Tail: 124
mm (tail 19.2% of SVL); Females: SVL: 950–1080 mm; Tail: 150 mm (tail 4.6% of SVL);
Neonates: 220 mm. Size of males and females from Murphy (2007a). Previous studies that
report on specimens (unsexed) include individuals with total lengths of 1220 mm (Smith,
1914b) and one with a 1075 mm SVL and 125 mm tail Werner (1923). Saint Girons (1972)

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Aquatic Snakes

recorded a female of 950 mm and a male of 812 mm total lengths. Neonate size data from
Flower (1899).
Natural History. Subsessor bocourti is a highly aquatic snake that uses swamps, shal-
low lakes, pools, and other stagnant water habitats, including those that are anthropogenic,
such as reservoirs, rice paddies, and fishponds (Campden-Main, 1970; Murphy, 2007a). It
is reportedly found with Acrochordus javanicus in swamps and pools, and it may move into
deep water during the hottest part of the year (Laidlaw, 1901;
Boulenger, 1903). They are considered nocturnal (Deuve,
1970), but have been found in lakes in Laos during the day-
time predating on fish (Nguyen et al., 2020). This species is
known to eat fish across multiple studies and observational
accounts (Deuve, 1970; Saint Girons, 1972; Murphy, 2007a).
A specimen from Tonlé Sap, Cambodia (555 mm) contained
a 50 mm catfish, (Bagridae: Mystus mysticetus). A neonate ob-
Figure 8.133. The distribution
tained near Kabin Buri, Thailand contained a 145 mm fresh-
of Subsessor bocourti. water eel (Monopterus albus). Several other specimens from
the same locality contained unidentified fish remains. Cap-
tive newborns will feed on small frog (e.g., Rana and Microhyla; Flower, 1899). In captivity,
females have produced 17 live young (Flower, 1899). A range of 6–17 eggs have been found
inside females in the Cambodian population (Saint Girons, 1972), and 26 embryos were
found in a female of a Thailand population (Murphy, 2007a). Using DNA, Subsessor bocourti
and Erpeton tentaculatum have both been recovered as sister taxa, though with low support
but genomic data (Bernstein et al. 2023a) strongly supports Subsessor as sister to both Hom-
alopsis and Cerberus.

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 8. The Homalopsidae

Figure 8.134. Bocourt’s Mud Snake, Subsessor bocourti.

Photography by (top) Tom Charleton and (bottom) JCM.

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Aquatic Snakes

Sumatran Mud Snake

Sumatranus albomaculata (Duméril, et al., 1854)
Distribution and Habitat. Found on the Sumatra sat-
ellite islands of Nias, Pulo, Simeuluë, Sibigo, and Sinabang;
and probably Sumatra proper. Type locality: “les environs
de Padani, île de Sumatra.” [=Padang, west Sumatera].(Fig-
ure 8.135)
Figure 8.135. The distribution of Identification. This species has smooth scales in 27
Sumatranus albomaculata. (26–28; possibly 25 based on literature accounts) rows; the
nasals are in contact; the last two or three upper labials are
horizontally divided; large internasal makes narrow con-
tact with elongated loreal; five of six lower labials contact anterior chin shields; dorsum with
scattered spots, venter mottled. Size. Males: SVL: 520 mm; Tail: 90 mm (tail 17.1–26.9% of
SVL); Females: SVL: 625 mm; Tail: 75 mm (tail 12.5–14.4% of SVL). Size data from largest
individuals from Murphy (2007a). Murphy (2007a) study also reported that the smallest in-
dividual (sex not given) was 260 mm SVL with a 70 mm tail. Gyi (1970) mentioned that male
tails are significantly longer than female tails.
Natural History. The natural history of this homalopsid is nearly unknown. A single
specimen (USNM 30767) contained three eggs and has an SVL of 520 mm (Murphy, 2007a).

On the next page. Amazonian Aquatic Coral Snake, Mi-

crurus surinamensis waiting in ambush Photography by
Konrad Mebert

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Aquatic Snakes

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Aquatic Snakes

9. Aquatic Coral Snakes

Most Neotropical coral snakes of the Micrurus genus are fossorial or cryptozoic. How-
ever, two species are semi-aquatic, M. nattereri and M. surinamensis (Passos and Fernandes
2005). Additionally, other species will readily enter the water and search for food – particu-
larly Micrurus diutius and other members of the Micrurus lemniscatus group.

Orinoco Aquatic Coral Snake

Micrurus nattereri Schmidt, 1952
Distribution and Habitat. Streams and wetlands
in the equatorial forest are associated with the upper
Orinoco and Negro Rivers, from southern Venezuela
and Colombia to northern Brazil. Type locality: Be-
tween Guaramaco and San Fernando de Atabapo in
the upper Orinoco River, Venezuela (Fig. 9.1)
Identification. Distinguished from other Micru-
Figure 9.1 The distribution of Micrurus rus (except M. surinamensis) by a frontal scale nar-
nattereri.
rower than supraoculars; only the fourth upper labial
borders the orbit;

Figure 9.2 Orinoco Aquatic Coral Snake, Micrurus nattereri.

Photography by Santiago Ayerbe.

195
 9. Aquatic Coral Snakes

head flattened with nostrils and eyes directed upwards; distinguished from M. surinamensis
by having 180–195 (instead of 156–174) ventrals in males and 193–215 (instead of 169–187)
ventrals in females. This species is smaller than surinamensis. The largest male M. natteri
was 650 mm TL, with an 80 mm tail; the largest female was 674 mm TL, with a 69 mm tail;
ventrals 180–195 in males; 193–206 in females; subcaudals 33–41 in males; 30–38 in females;
upper labials 7–8; lower labials 7–8; temporals 1+2 or 1+3; the number of triads 6–8; width
of the central triad at the level of the midbody 5–10 scales; width of interspaces 6–12 scales;
hemipenis extend 13–15 subcaudal scales (based on Passos and Fernandes 2005).

Amazonian Aquatic Coral Snake

Micrurus surinamensis (Cuvier, 1816)
Distribution and Habitat. Rivers in equatorial forest in South
America in the Guiana region, Colombia, eastern Ecuador, Peru,
Bolivia, and central Brazil. In Brazil, it occurs in Acre, Amazonas,
Roraima, Rondônia, Pará, Maranhão, Tocantins, and Mato Gros-
Figure 9.3. The distri- so. Type locality: Suriname (Fig. 9.3).
bution of Micrurus suri- Identification. Ditinguished from other species of the genus
namensis. by a frontal scale narrower than supraoculars, only the fourth up-
per labials border the orbit of the eye; head flattened with nostrils
and eyes directed upwards and distinguished from M. nattereri
by having 156–174 (nattereri 180–195) ventrals in males and 169–187 in females (instead of
193–215 in surinamensis ventrals). Rostral scale broader than high, with a triangular shape
in frontal view and visible from above; internasals less than half of the length of prefrontals;
frontal longer than broad; supra-
oculars approximately as broad
as the prefrontal; parietals about
twice longer than wide; nasal di-
vided; one preocular, longer than
high; two postoculars; fourth up-
per labial enters the orbit; first pair
of lower labials contacts behind
symphysial; first four pairs of low-
er labials touch first pair of chin
shields; two pairs of chin shields;
usually four gulars; 15 dorsal scales
rows, smooth, without reduction;
cloacal plate divided (Figure 9.4).
Natural History. Silva et al.
(2018) found the head shape of su-
rinamensis is convergent with oth-
er aquatic snakes. However, Mi-
crurus surinamensis differs from
the two semi-aquatic species of
Figure 9.4. Micrurus surinamensis.
Micrurus lemniscatus and M. spixii
Photography by Konrad Mebert.
in having a wider head, a smaller dis-
tance between nostrils, and a longer

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Aquatic Snakes

tail. Despite having an extremely conserved skull and mandible shape, M. surinamensis has
longer supratemporal and quadrate bones than in terrestrial coral snakes, suggesting a larger
gape for feeding on fish are larger and broader prey in contrast with the elongated prey of
other Micrurus. The South American coral snakes (M. lemniscatus species complex) are, to a
lesser degree, semi-aquatic and are known to feed on synbranchid eels and shallow water fish
but do not demonstrate the morphological adaptations seen in surinamensis. Tavares-Pin-
heiro et al. (2021) observed this snake feeding on a knifefish, swallowing it headfirst on the
edge of a stream. The coral snake captured the fish at night in water ca. 1.5 m deep, but the
fish was subdued and consumed on the stream bank.
Micrurus surinamensis inhabits the Amazon drainage, while M. nattereri occupies the
Orinoco drainage basin. The separation of these two basins likely resulted in the evolution
of the two taxa in the late Miocene when the Vaupes Arch semi-isolated the river systems.
However, separating the two basins is still incomplete, with the Casiquiare River in southern
Venezuela connecting to the Orinoco and Negro Rivers. The connection explains the pres-
ence of M. nattereri in the upper Negro River.
.

On the next page. The Banded Water

Cobra, Naja annulata from Kinshasa,
DRC. Photography by Vaclav Gvozdik.

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 9. Aquatic Coral Snakes

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Aquatic Snakes

10. Aquatic Cobras (Najinae)

Lake Tanganyika, the African Great Lake, is well-known for the two British explorers,
Richard Burton and John Speke, who discovered the lake in 1858. A species swarm of more
than 250 cichlid fish has made this lake famous among biologists and hobbyists because the
fish are a well-studied example of rapid evolution and are popular in the pet trade. But a
fish predator is living here that is also of interest, Boulenger’s Water Cobra (Naja annulata).
Arthur Loveridge, a Harvard University herpetologist, was one of the first to provide observa-
tions on this snake. At Kasanga, rocky promontories extend into the Lake, and the Germans
built one for their Bismarckburg military base. A natural breakwater of jumbled rocks pro-
tects these peninsulas, and it is here Loveridge observed the aquatic cobras. He wrote,

According to native reports, which my own experience confirmed in some

points and contradicted in none, when the sun rises and strikes the rocks the
cobras emerge from their retreats beneath them and bask for a short time on
the tops of the rocks. Shortly afterward...and I found none on the rocks an hour
and a half after sunup...they take to the water in search of fish. I was told on a
calm day one might see as many as ten in a morning’s fishing. We saw four in
a little over three hours. The snakes come out of the water in the evening, and
they are said to bask under the rocks. Loveridge suggested that this is probably
correct because the evenings were cool, and the snakes may absorb heat from
the rocks and avoid the cool wind blowing in mid-May.
Boulengerina Dollo (1886) was established for a cobra with aquatic habits, Boulengerina
annulata (Buchholz & W. Peters, 1876) and Boulengerina christyi Boulenger, 1904 was later
assigned to the genus. Wallach et al. (2009) returned the two species of Boulengerina to Naja.
They recognized Boulengerina as a subgenus containing four non-spitting cobras species,
including semi-aquatic and semi-fossorial species. Wüster et al. (2018) used mitochondrial
and nuclear gene sequences and morphological data to diagnose species limits within the Af-
rican Forest Cobra, Naja (Boulengerina) melanoleuca. They found deep genetic divergences
within N. melanoleuca and recognized four additional species (N. subfulva, N. peroescobari,
N. guineensis, N. savannula), two of which were new. The lifestyles of these snakes, previously
considered to be Naja melanoleuca are poorly known. However, Wüster et al. (2018) state
that Boulengerina is composed of a mixture of small and primarily terrestrial, burrowing,
and aquatic forms occupying a diversity of biomes and with distinct foraging ecologies. More
recently, another member of the genus was described, the Dwarf Water Cobra, Naja nana.

Banded Water Cobra

Naja annulata Peters, 1876
Distribution and Habitat. Distributed in central Africa, from the Cameroons to Tang-
anyika, where it is often associated with flooded forests and rarely occurs far from water. It
occurs in Burundi, Cameroon, Central African Republic, Democratic Republic of Congo, Re-
public of Congo, Equatorial Guinea, Gabon, Rwanda, and the province of Cabinda in Angola,
as well as along the Burundian, Tanzanian and Zambian shores of Lake Tanganyika. Type
locality: “Dorfe Mbusa (Eliva Sonange am Ogowe),” Gabon.
The subspecies Naja annulata stormi was considered endemic to Lake Tanganyika, but
there is a single record from Pweto on Lake Mweru, Katanga Province in the Democratic

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 10. Aquatic Cobras (Najinae)

Republic of Congo (Broadley and Cotterill 2004). Type locality:

Lake Tanganyika, Zaire. Pauwels et al. (2017) report it from the
buffer zone of the southeastern part of Minkébé National Park,
where one was killed in a river and eaten by local people. Hab-
itat includes flooded forests and savannas and is rarely far from
Figure 10.1. The distribu- water.
tion of Naja annulata. Natural History. The Banded Water Cobra is in the water
most of the time. It is an excellent swimmer and can remain
underwater for up to 10 minutes and dive to depths of 25 m (82 ft). On land, It moves slowly
and hides among rocks, in burrows made by other animals, or in overhanging tree roots at the
shoreline; It also uses artificial structures, such as bridges and jetties, to hide. It is generally
not aggressive, and if approached in water, it will swim away swiftly and, on land, it will at-
tempt to escape into the water. If threatened on land, it will rear up and spread its narrow yet
prominent hood, and it may hiss loudly, but it tends not to make any forward movements. It
will only bite when provoked. It feeds on fish and amphibians. When not looking for food, it
uses rocks, bank holes, tree holes, and root clusters for refuge. Human encounters with water
cobras are probably few, and the snake poses little danger to anyone except when it becomes
entangled in fishers’ nets or in the nets of herpetologists trying to trap it.
Venom. The LD50 of long-chain neurotoxin two from B. a. annulata venoms was 0.086
mg/kg. The venom of this little-known elapid has the lowest LD50 of any African protero-
glyph studied thus far and has a high concentration of potent postsynaptic neurotoxins.

Figure 10.2. Banded Water Co-

bra, Naja annulata. (A) N. a.
stormsi Lake Tanganyika, Tan-
zania (B) N. a. annulata from
the DRC) Photography by (A)
Wolfgang Wuster, (B) Vaclav
Gvozdik.

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Aquatic Snakes

Congo Water Cobra

Naja christyi Boulenger, 1904
Distribution and Habitat. The lower reaches of the Zaire River. The Congo Water Cobra
has a small range in the western Democratic Republic of Congo, the southern half of the
Republic of Congo, a small portion of southeastern Gabon, and the province of Cabinda in
northern Angola. It is highly aquatic and always near water, in lowland bush or wooded areas
along banks of lakes, rivers, and streams. It is restricted to
the lower Congo River and nearby wet forested areas. Type
locality: Leopoldville, Congo.
Natural History. A poorly studied species. The diet is
mostly fish. But, the literature suggests that amphibians, liz-
ards and rodents may be consumed.
It is active day and night and spends much of its time
Figure 10.3. The distribution of in the water. Not an aggressive snake; when approached in
Naja christyi. water, it will swim away; If threatened on land, it will rear up,
spread its narrow hood, and hiss. It may strike if provoked.
Like other cobras, it is oviparous.
Venom. Not well studied, but believed that the venom
is dangerously neurotoxic, like most elapids. The LD50 of
long-chain neurotoxins-2 from N. christyi venoms was 0.090
mg/kg. The venom of this elapid has the lowest LD50 of any
African proteroglyph studied thus far and has high concen-
trations of potent postsynaptic neurotoxins (Weinstein et al.
1991).
A DNA has not been included in the nolecular studies.
It is unclear where this species falls phylogenetically. Schmidt
(1923) erected a new genus, Limnonaja, for it because he con-
sidered it morphologically distant from N. annulata.

Dwarf Water Cobra

Naja nana Collet and Trape, 2020
Distribution and Habitat. Democratic Republic of Con-
go (Katanga [Zaire]). Known only from Lake Mai-Ndombe,
The type locality is the environs of the village of Bokeben.
Natural History. Locally, the common name is Musso.
The Dwarf Water Cobra feeds exclusively on fish. It is fre-
Figure 10.4. Congo Water Co- quently caught in the nets of the fishermen at Lake Mai-
bra, Naja christyi. From Kin- Ndombe. According to the villagers, it stays in the boulders
haa,DRC. Photography by Va- bordering the Lake when it is out of the water. At high water,
clav Gvozdik. it climbs into the branches of emergent vegetation border-
ing the Lake. When threatened out of the water, Naja nana
adopts the classic cobra defense behavior by raising its head and anterior body and spreading
its hood. Captives only accept fish and refuse to eat amphibians (Collet and Trape 2020).
Captive specimens have produced clutches of four eggs. The eggs are relatively large
and elongated (~70 x 24mm). Incubation at 29° C was exactly 70 days (laid June 17, first
hatching August 26 (Raw and Deacon 2021).
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 10. Aquatic Cobras (Najinae)

Fishers who handle it frequently teach their children not to

fear it. A snake hunter who took more than 300 specimens from
nets was bitten about fifteen times, and symptoms of envenom-
ation followed five times. The bites were moderate and lacked
respiratory symptoms. There was mild local pain and slight ede-
Figure 10.5. The distirbu- ma, which resolved in 24-48 hours.
ition of the Dwarf Water
Cobra, Naja nana.

Figure 10.6. The Dwarf Water Cobra, Naja nana.

Photography by Jean-Francois Trape.

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Aquatic Snakes

Forest Cobra clade

Naja melanoleuca Hallowell, 1857
Distribution and Habitat. Swamp forests and mangrove forests from Nigeria, Camer-
oon, Central African Republic, Equatorial Guinea, Gabon, Republic of Congo (Brazzaville),
Democratic Republic of the Congo (Kinshasa) (Zaire), N Angola, and Rwanda. Type locali-
ty: Gabon. Cryptic diversity in this species was described by
Wüster et al (2018), and the species was divided into five
taxa. However, the exact distributions are unclear.
Natural History. It is considered a semi-aquatic snake
because it escapes to the water and feeds on amphibians,
other snakes, and fish (Haagner and Carpenter 1988).
Clutches of 15-17 eggs are known and require about 78
Figure 10.7. The distribution of days of incubation. The venom is neurotoxic.
Naja melanoleuca. The African Forest Cobra, Naja (Boulengerina) melan-
oleuca was recently shown to be a composite species. Using
mitochondrial DNA sequences and morphology, Wuster et al. (2018) revealed deep diver-
gences and recognized five species within N. melanoleuca.

Naja subfulva (Laurent, 1955)

Distribution and Habitat. from South Kivu Province of the Democratic Republic of

Figure 10.8. Forest Cobra, Naja melanoleuca.

Photography by Myke Clarkson.

the Congo (DRC), Forest/savanna mosaic, encircling the Congo Basin, from the grasslands
of western Cameroon northward, to Bol, Chad, on the northern shore of Lake Chad, east
through the CAR and South Sudan to west Ethiopia, south through Uganda, western Kenya,
eastern DRC, Rwanda, Burundi and western Tanzania to northern Malawi, west through
Zambia and Katanga to Angola and the Lower Congo region (Fig. 6). The distribution ex-

203
 10. Aquatic Cobras (Najinae)

tends disjunctly east and south to the East African coast from Kenya to northern KwaZu-
lu-Natal (South Africa), including inland locations in western Zimbabwe.

Naja peroescobari Ceríaco, Marques, Schmitz & Bauer 2017

Distribution and Habitat. Known only from the island of São Tomé, adult male col-
lected in the vicinity of Praia Inhame, São Tomé Island, Republic of São Tomé e Príncipe. It
does not appear to be a semi-aquatic species.

Naja (Boulengerina) guineensis Broadley, et al., 2018

Distribution and Habitat from the Upper Guinea forest of West Africa, The distribu-
tion of Naja guineensis appears to be restricted to the Upper Guinea Forests of western Af-
rica, from west Togo to Liberia and Guinea. It does not appear to be a semi-aquatic species.

Naja (Boulengerina) savannula Broadley, et al., 2018

Distribution and Habitat. A banded cobra from the savanna-forest mosaic of the
Guinea and Sudanian savannas of West Africa. From Senegal and Gambia east to north-
ern Cameroon in gallery forest areas in the Guinean Forest/Savanna Mosaic. This is not a
semi-aquatic species.

Monocled Cobra
Naja kaouthia Lesson, 1831
Distribution and Habitat. They inhabit natural and anthropogenic habitats and prefer
water-associated habitats, such as swamps, and mangroves, but can also be found in grass-
lands, shrublands, and forests. Agricultural land (paddy fields), human settlements, and
cities are also used. They occur at elevations of up to 1000 m above sea level. The range
includes Bangladesh, Myanmar, Cambodia, northeast India, Bhutan, Laos, northern Ma-
laysia, Nepal, S China, Thailand, and southern Vietnam.
Type locality: Bengal, India.
Natural History. Diet includes snakes, lizards, and
fish. In Myanmar, it has been observed to enter fish traps
to prey on the fish in the trap (Kyi and Zug 2003). They
are primarily crepuscular. In rice-growing areas, they
hide in rodent burrows in the dykes between fields and
have become semi-aquatic in this type of Habitat. Ju-
veniles feed primarily on amphibians, while adults prey
Figure 10.9 The distribution of
on small mammals, snakes, and fish. When disturbed,
Naja kaouthia.
they prefer to take flight. However, when threatened,
they will raise the anterior body, spread a hood, often
hiss loudly, and strike to bite and defend themselves. Sometimes they inhabit tree holes and
areas where rodents are plentiful.
Venom. Some populations of the monocled cobra can spit venom. Adult and juvenile
N. kaouthia venom composition was investigated by Modahl et al. (2016). Alpha-cobra-
toxin (a-elapitoxin) was the only significantly toxic 3FTx (three-fingered toxin) found in
its venom, and it was equally harmful in both lizard and mouse models. The abundance
and diversity of 3FTxs and most enzyme activities did not vary between adult and juvenile

204
Aquatic Snakes

cobra venoms; however, total venom PLA2 activity and specific PLA2 isoforms did vary, with
juveniles lacking several of the least acidic PLA2s, and these differences could have both bio-
logical (related to predation) and clinical (antivenom efficacy) implications. Nevertheless, the
ubiquitous presence of a-cobratoxin in both adult and juvenile cobra venoms, with high tox-
icity toward both reptiles and mammals, represents a venom compositional strategy wherein
a single potent toxin effectively immobilizes a variety of prey types encountered across life
history stages.

Figure 10.10 Monocled Cobra, Naja kaouthia.

Photography by Avrajjal Ghosh.

Next Page. A Lowland Copperhead hunting in a pond. Photog-

raphy by Ben Fisher.

205
 10. Aquatic Cobras (Najinae)

206
Aquatic Snakes

11. Semi-Aquatic Hydrophiinae

The elapid subfamily Hydrophiinae contains 38 genera and about 200 species. It in-
cludes snakes entirely terrestrial or fossorial, and some are aquatic. This chapter examines the
semi-aquatic species minus the genus Laticauda (Chapter 12). The genus Hydrophis (Chapter
13) are the most highly aquatic member of this clade, with a paddle-like tail, but other species
also show trends toward aquatic behavior.

New Guinea Small-eyed Snake

Micropechis ikaheka Lessons, 1830
Distribution and Habitat. It inhabits rainforests and coco-
nut plantations in Papua New Guinea, West Papua, and the In-
donesian islands of Batanta, Misool, Jobi, Mios, Num, Mefoor,
Mansinam, Valise, and the Aru Islands. Type locality: Ddore,
Irian Jaya (=West Papua) (Figure 11.1).
Natural History. Diet includes squamates, conspecifics,
Figure 11.1. The distribution of terrestrial boid Candoia aspera, Sphenomorphus jobiensis,
Micropechis ikaheka. colubrid Stegonotus, and large skink Tiliqua gigas, and large
earthworms. Small mammals and frogs are also eaten (Krey
et al. 2015). Females produce clutches of 2-7 eggs in October,
which are relatively large, 40 to 65 mm long (Shine and Keogh
1996). The snakes are easiest to find in the drier weather when

Figure 11.2. New Guinea

Small-eyed Snake, Micro-
pechis ikaheka Photography
by David Williams (top) Paul
Freed bottom..

207
 11. Semi-Aquatic Hydrophiinae

they are hidden inside the coconut husk piles. It has been reported to catch fish (eels) in the
water (Krey et al. 2015).
Small-eyed snakes bite plantation workers, grass-cutters, and villagers when they en-
counter these snakes abroad at night or in the early morning, especially during the wet sea-
son (Figure 11.2).

Pygmy Copperhead
Austrelaps labialis (Jan, 1859)
Distribution and Habitat. South Australia. Kangaroo Island,
Mt. Lofty Ra, and the Fleurieu Peninsula. Island populations use
coastal dunes, woodlands, samphire flats, and agricultural areas.
Mainland populations are found in high-altitude stringybark for-
ests with a dense understory (Wilson and Swan 2003). Type local-
ity: Australia. NeoType locality: Islet 477, Pelican Lagoon, Kanga-
Figure11.3. The distribu-
tion of Austrelaps labialis. roo Island, South Australia (Rawlinson 1991).
Natural History. It is a semi-aquatic snake that feeds on
frogs, tadpoles, and lizards. Females are gravid from late spring (October) to early autumn
(March). Parturition can occur as early as February. Litters usually contain seven young but
have a maximum size of 20. Full-term embryos are 131-142 mm.

Figure 11.4. Pygmy Copperhead, Austrelaps labialis, Adelaide Hills, SA.

Photography by Shawn Scott.

Highlands Copperhead
Austrelaps ramsayi (Krefft, 1864)
Distribution and Habitat. Mesic upland habitats in New South Wales and Victoria, Aus-
tralia. Type locality: near Braidwood, NSW. NeoType locality: Moss Vale. Montane heaths,
woodlands, creek edges, and marshes with dense grasses.

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Aquatic Snakes

Natural History. A semi-aquatic snake. The diet comprises

small frogs, tadpoles, lizards and snakes, and squamate eggs. Fe-
males give birth to neonates that are about 146 mm long.

Figure 11.5 The distribu-

tion of Austrelaps ramsayi.

Figure 11.6 Highland’s

Copperhead, Austrelaps
ramsayi. The Blue Moun-
tains. Photography by Scott
Eipper.

Lowland Copperhead
Austrelaps superbus
(Günther, 1858)
Distribution and Habitat. South Australia, Tasmania, and Victoria, Australia. Type lo-
cality: Australia and Tasmania, restricted to Tasmania by Günther (1863). It prefers areas
of low vegetation near water. It has also been found on sandstone ridgetop woodland in the
Blue Mountains, west of Sydney, which is becoming rare due to
increasing fires and the spread ofurban settlements.
Natural History. The diet includes primarily small frogs,
lizards, and snakes. Reproduction probably does not occur ev-
ery year. Litter sizes average about 15 neonates.

Grey Snake
Figure 11.7 The distribu-
tion of Austrelaps superbus. Hemiaspsis damelii (Günther, 1876)
Distribution and Habitat. Dry sclerophyll forests and wood-
lands on clay soils where water bodies or gullies are present in

Figure 11.8. Lowland Cop-

perhead, Austrelaps super-
bus, Warrandyter. Photog-
raphy by Scott Eipper.

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 11. Semi-Aquatic Hydrophiinae

southeast Queensland and New South Wales, and there are isolated populations in north-east-
ern South Australia and southwestern New South Wales Australia. Type locality: Peak Downs,
Qld.
Natural History. At Macquarie Marshes in central New
South Wales, Shine (1987) found H. damelii is exclusively cre-
puscular and only active for an hour or two afterward. The diet
is primarily frogs: Litoria (Hylidae), Adelotus, Limmnodynastes,
Rana.
Figure 11.9. The distribution
of Hemiaspis damelii.

Figure 11.10. Grey Snake,

Hemiaspis damelii. Photogra-
phy by Melissa Bruton.

Black-bellied Swamp Snake

Hemiaspis signata (Jan,1859)
Distribution and Habitat. Rainforests and wet sclerophyll forest, margins of creeks, dams,
wetlands, and low-lying seasonally flooded areas along the east coast of New South Wales and
Queensland, Australia. This species may be partially estuarian.
It occurs in marshes along the east coast of Australia. Type lo-
cality: Sydney, Australia.
Natural History. Hemiaspis signata may be active all day
and well into the night (Shine 1987). The diet of H. signata is
much broader than that of H. damelii, with the recorded prey
Figure 11.11. The distribu- items including one invertebrate, 22 anurans, 68 lizards, and 13
tion of Hemiaspis signata. lizard eggs. Only two of the 22 frogs are arboreal species. Apart
from a single gecko, all the lizards consumed by H. signata were
skinks, with small diurnal species of the genus Lampropholis
being the most critical (37 records). The scincid species eaten

Figure 11.12. Black-bellied

Swamp Snake, Hemiaspis sig-
nata. Photography by Tony
Gerard <tonyg@shawneecc.
edu>

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Aquatic Snakes

also included fossorial (e.g., Anomalopus) as well as larger diurnal surface-active taxa (e.g.,
Ctenotus) (Shine 1987). Females produce 6 to 16 embryos per clutch (Shine 1977).

Red-bellied Black Snake

Pseudechis porphyriacus (Shaw, 1794)
Distribution and Habitat. The east coast of Australia, in the urban forest, wood-
lands, plains, and bushland areas of the Blue Mountains, Canberra, Sydney, Brisbane,
Melbourne, Cairns, and Adelaide. The Macquarie Marshes mark a western border to its
distribution in New South Wales, and Gladstone in central
Queensland marks the northern limit to the main popu-
lation. To the south, it occurs across central and east Vic-
toria and extends along the Murray River into South Aus-
tralia. Disjunct populations occur in the southern Mount
Lofty Ranges in South Australia and North Queensland.
The type locality is New Holland, Australia. It is often
Figure 11.13 The distribution close to dams, streams, billabongs, and other bodies of
of Pseudechis porphyriacus. water, although they can venture up to 100 m (350 ft)
away, including into nearby backyards. In particular, the
Red-bellied Black Snake prefers shallow areas with tan-
gles of water plants, logs, or debris (Shine 1981b).
Natural History. The diet includes fish, frogs, reptiles (including other snakes), and
small mammals. Fish are hunted in water - at the surface or underwater. Individual snakes
stir up the substrate in search of prey. As they grow and mature, they eat the same prey and
add larger animals. They have been reported to scavenge road-killed frogs.

Figure 11.14. Red-bellied Black Snake, Pseudechis porphyriacus.

Photography by Shawn Scott.

211
 11. Semi-Aquatic Hydrophiinae

Males engage in ritualized combat for 2 to 30 minutes and may attack males already
mating with females. They wrestle vigorously but rarely bite and engage in head-pushing con-
tests, where each snake tries to push his opponent’s head downward with his chin. Females
can gravitate from early spring to late summer (Shine et al. 1981). After that, they become less
active and congregate in nursery areas. They are viviparous; and give birth to young after 14
weeks gestation, usually in February or March. Litter sizes 8- 40. Neonates average 122 mm.
Young triple their length and increase their weight 18-fold in their first year of life. Males are
sexually mature when they reach 780 mm, and females mature at about 880 mm. Females can
reproduce at about 31 months of age (Shine 1977, 1987b).

Tiger Snake
Notechis scutatus (Peters, 1861)
Distribution and Habitat. Coastal environments, wetlands, and creeks in Australia (New
South Wales, Queensland, South Australia, Victoria, Western Australia, Tasmania). Type lo-
cality: Java (in error); neotype locality: “The Brothers,” 10km NE of Benambra, Victoria. This
is a polytypic species, and two subspecies are recognized. N. s. scutatus and N. s. occidentalis:
The latter occurs in southwestern Western Australia and had no
type locality given.
Natural History. A semi-aquatic snake. Mainland adult
snakes feed on frogs and mice, while adult island snakes mostly
feed on silver-gull chicks. Tiger snakes give birth to 20 -30 young;
the largest litter reported was 64. Tiger Snakes usually mate in
Figure 11.15 The distribu-
tion of Notechis scutatus. spring and gives birth in the summer. A released captive snake
crawled into a pond and submerged for 18 m 36 s (Cornelis and
Lettoof 2020).
Temperature preferences are plastic in some snake species.
Michniewicz and Aubret (2010) experimentally raised young Tiger Snakes in terrestrial or
semi-aquatic environments over 11 months. Young snakes raised in a semi-aquatic environ-
ment selected slightly but significantly higher mean body temperatures (30.3°C) than their
terrestrially grown siblings (29.5°C). The semi-aquatic raised young allowed their body tem-

Figure 11.16. Tiger Snake, Notechis scutatus.

Photography by Wolfgang Wuster.

212
Aquatic Snakes

perature to remain above 32°C for twice as long as the terrestrial raised snakes (4.4 hours vs
2.1 hours. Swimming speed was strongly linked to body temperature. Entering the water with
a higher body temperature (30°C versus 19°C) delayed a sharp drop in swimming speed, thus
lengthening maximum performance time.
Translocated snakes studied by Butler et al. (2005) traveled greater distances than resi-
dent telemetered snakes, although there was no difference in the frequency of movements.
The degree of cloud cover and the maximum daily temperature significantly influenced daily
activity, with snakes more likely to be active on relatively cool, sunny days. Snakes exhib-
ited bimodal peaks in daily activity avoiding the hottest part of the day. Both resident and
translocated snakes preferred grassy woodlands associated with hills but avoided escarpment
woodlands and open plains. Floodplain riparian woodland was avoided by resident snakes
but used in proportion to its availability by translocated snakes.

On the next page. New Caledonia Sea Krait, Laticauda saintgiron-

si. Photography by Jean Roger.

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 11. Semi-Aquatic Hydrophiinae

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Aquatic Snakes

12. Sea Kraits

Myke Clarkson

Sea kraits form a monophyletic clade, the Laticaudini composed of a single genus, La-
ticauda with eight species of banded paddle-tailed snakes. Seven of the eight species of this
group inhabit a mix of marine and land environments, with Laticauda crockeri being the
sole exception, restricted to a single brackish lake in the Solomon Islands. Sea kraits are am-
phibious, rather than fully aquatic like their close relatives the true sea snakes (Hydrophi-
ini.) While both groups of elapids evolved into marine environments about the same time
13‒8 million years ago (Kim et al. 2018), the two groups have taken two notably different
evolutionary paths. Laticaudini nasal scales are separated by one or two internasal scales,
have retained widened ventral scales and oviparity from their terrestrial ancestors. Their
bodies are cylindric in shape, scales are smooth, and imbricate. Sea kraits are adapted for
life on land as well as the water, they spend much of their lives in aquatic environments
searching for prey. Their time on land is for digestion, thermoregulation, mating, ecdysis
(shedding), and egg laying. Drinking freshwater on land has also been observed in this
genus, which is reliant on freshwater for hydration. The reliance on fresh drinking water is
thought to be one of the potential factors that has led to the groups characteristically patchy
distribution (Lillywhite et al. 2008).
This dependence of freshwater for drinking also lends this group highly susceptible to
impacts of climate change. As mentioned above, all sea kraits are oviparous, except for Lat-
icauda crockeri which could potentially be viviparous this has not been confirmed. Average
clutch sizes for Laticauda are 4–19 eggs. The thin shelled eggs are laid on land and have a
high permeability to both oxygen and water. Hatchlings mature rapidly, and sexual maturity
is reached in approximately 18 months in males and eighteen to thirty months in females
(Heatwole et al. 2005). Courtship can occur on water or land and can involve many males
courting a single female at once. However, unlike other species who can engage in polygyny
(including Thamnophis, Natrix, and Eunectes referenced in this book) male sea kraits do
not show signs of engaging in combat to fend off other suitors. While in other species body
pushes or tail wrestling may be employed, sea krait males seem to not directly compete
with one another. Instead males seem to have scamble competition for mating without a
influence of male body size on mating success (Sheet and Shine, 2002c). All members of the
group are venomous and of medical significance to humans, though generally perceived to
be mild mannered in nature. While their venom is highly toxic, Rasmussen et al suggested
Laticauda might also use their tails as a defense mechanism against predators, twisting them
to look like a second head, while their head can continue to forage for food in cracks and
crevices (Rasmussen et al. 2009).
Natural predators can include tiger sharks sea eagles, and larger fish. The placement
of the sea kraits has been a topic of much discussion over the years. While once classified
as their own family Laticaudidae, they were later put into the marine Elapidae subfami-
ly Laticaudinae. With improvements in gene sequencing the close relations of these ma-
rine elapids has been more clearly revealed and they have most recently been placed in the
monophyletic group Laticaudini (Kishida 2020; Pyron et al. 2011). Less debated than the
taxonomic placement of this clade is the ecological categorization with in it. Laticaudini
have been divided into three closely related complexes, though the complexes ranges in-
clude large overlap, their habitat usage (land versus water) and relation is distinct. The first
group is the “Laticauda colubrina complex” and includes L. colubrina, L. frontalis, L. guineai,

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 12. Sea Kraits

and L. saintgironsi. Members of this group are notably

more terrestrial. The second group is “L. semifasciata
complex” which includes L. semifasciata and L. schis-
torhyncha, which are the most aquatic of the sea kraits.
The third group, the “L. laticaudata complex” includes
L. laticaudata and L. crockeri and is an intermediate
group between the colubrina and semifasciata com-
plexes, spending less time on land than colubrina, but
more than semifasciata (Brischoux, et al, 2013). The Figure 12.1. The distributiuon of Lat-
sea kraits are found in both tropical and subtropical icaudini.
coastal waters in the eastern Indian Ocean, south-east
Asia, and the western Pacific Ocean. They are found as
far north as Ryukyu Islands, as far South as Vanuatu, as far East as the Solomon Islands and
as far west as the Andaman Islands (Figure 12.1). While Laticauda colubrina and Laticauda
laticauda have very expansive ranges, other members of the group like Laticauda schisto-
rhynchus and Laticauda saintgironsi are isolated to single islands (Gherghel 2016; Kishida
et al. 2020). There are unconfirmed sightings on the western coasts of Mexico, El Salvador,
and Nicaragua, but no records have been presented to back this finding. Sea kraits face many
threats including habitat loss, and under regulated harvest. Currently five of the eight species
in the genus are considered vulnerable or near threatened by the IUCN.

Yellow-lipped Sea Krait

Laticauda colubrina (Schneider, 1799)
Distribution and Habitat. Laticauda colubrina has the largest distribution of any spe-
cies in the Laticaudini group, and while there is a degree of
morphological variance throughout the species range, it is still
regarded as a singular species (Lane and Shine, 2011). In the
Indian Ocean it is found in East India, Sri Lanka, Myanmar,
the Andaman Islands, the Nicobar Islands, Western Vietnam,
Western Malaysia, and Western Indonesia. In the Pacific it
Figure 12.2. The distributiuon is found in Eastern Vietnam, Eastern Malaysia, Micronesia,
of Laticauda colubrina.
Polynesia, Solomon Islands, Vanuatu, Thailand, Philippines,
Palau, Taiwan, China, New Guinea, Timor-Leste, Japan’s
Ryukyu Islands, Australia, New Zealand, Fiji, and New Caledonia. Type locality: not given.
Yellow-lipped sea kraits are most often encountered on shallow reef shelfs, small islands, and
rocky shores which they utilize for basking and shelter. In the author’s experience, sea kraits
prefer reef rock lined shores, and can most easily be found sheltering beneath the rocks and
in roots by day, or by walking suitable shorelines and islands by night. While this is the most
common place of encounter, they can also be found in deep waters many kilometers away
from the nearest islands while foraging for prey or floating on the surface at night. Shine et al
observed that Yellow-lipped sea kraits exhibit philopatry and return to specific home islands
even when relocated distances of more than 5 kilometers (Sheety and Shine 2002a).
Identification. Head is black, with a characteristic yellow mask that typically encompass-
es the snake’s labial, nasal, prefrontal, and ocular scales. The yellow mask typically terminates
at the frontal scale on the top of the head. On the sides of the head the yellow mask divides
into two yellow stripes separated by black at the temporal scales just behind the eyes. These
stripes are variable in length and can continue just a few scales or to the back of the head

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Aquatic Snakes

Figure 12.3. Yellow-lipped Sea Krait, Laticauda colubrina.

Photography by Yu Sing Lin.
terminating before the first light colored band. While this is the most common coverage of
the yellow mask, regional variation can occur in terms of hue and saturation of the yellow, as
well as quality of yellow coverage. The body is banded with black and blue alternating bands,
often having a darker blue tone dorsally and gradually shift-
ing to a lighter blue or white ventrally. The tail can at times
have a yellow tint to it, as can the first band after the head,
however coloration of this species does vary throughout its
range as does saturation levels of the blue bands. The tail
is laterally flattened with the end tip often being a white,
blue, or light-yellow color. Ventrals are sizable, averaging
one third or larger than half the body width. Rostral is un-
divided, with a prefrontal shield usually present. As with all
Laticauda, pupils are round, nostrils lateral and nasals are
separated by inter nasals. The mid body has 21–25 longi-
tudinal rows of scales and the average animal has 213–243
ventral scales. Laticauda colubrina have 37-47 subcaudals
in males, and 29-35 in females. Average snout to vent length
on males is 875 mm, and 1420 mm for females. The average
tail length for males is 130 mm, and 145 mm for females
Figure 12.4. This is likely a mat- (Leviton, 2003).
ing aggregation of Laticauda col- Natural History. Like all sea kraits, Yellow-lipped Sea
ubrina. With a large female and
Kraits are amphibious in nature, and are most active at
many smaller males. Taken on
dusk and at night. They are not considered fully nocturnal,
Malapascua Island, Philippines
Photography Tony Gerard. due to occasional diurnal activity and thermoregulating on
land (Heatwole, et al, 2005). This species spends much of

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 12. Sea Kraits

Figure 12.5. The terrestrial habitat of Laticauda colubrina. A small satelitte island off the coast of Sa-
bah, Malyasia. At low tide it was possible to walk to the pile of rocks. About six sea kraits were on the
top in the vegetation. Photography by JCM.

its time at sea foraging for moray eels (Muraenidae) and conger eels (Congridae). While eels
make up most of this species prey, other bony fish have also been observed in stomach content
studies (Gorman, et al, 1981). Yellow-lipped Sea Kraits are active hunters, using chemorecep-
tors to locate eels hidden in rocks and crevices. Like other Laticauda this species has a unique
saccular lung allowing for deep dives up to 60 m in the search of food. Shine observed sexual
dimorphism in this species, like that seen in the Acrochordidae. Large females will search for
a single large conger eels in deep water, whereas the males forage for multiple prey consisting
of smaller moray eels. Prey are subdued rapidly by a potent neurotoxic venom with an LD50
of 0.1 mg/g IV in mice (Tan, et al, 2017). This corresponds to the morphological differences
between the smaller males and larger females with disproportionally larger heads (Shetty and
Shine, 2002b). Conversely it was found males have stronger terrestrial locomotion than females
yet decreased swimming ability. This could correspond to the fact that males spend more time
searching for mates, than females. Increased land mobility in males comes at a cost, decreased
aquatic mobility in comparison to the females (Shine and Shetty, 2001). As the most terres-
trial of the sea krait groups, the L. colubrina complex is also the fastest moving on land when
compared to L. semifasciata and L. laticauda complexes (Wang et al 2013). Breeding cycle and
clutch size vary greatly thoughout the species range. Mating takes place on shore throughout
the entire range, with breeding behavior being polyandrous. Little is known about the behavior
of hatchlings (Wright, 2011).

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Aquatic Snakes

Rennel Island Sea Krait

Laticauda crockeri (Slevin, 1934)
Distribution and Habitat. Restricted to Lake Te-Ngga-
no, Rennell Island, Solomon Islands. It is known exclusive-
ly from a single brackish lake, (2.8-6.2% salinity) where it
is sympatric withthe distantly related Laticauda colubrina.
Lake Te-nngano is the largest body of enclosed water in
the insular Pacific, spanning 155 km2 and reaching depths
of 40 meters (UNEP, 2008). The lake has a hard limestone
Figure 12.6. The distribution of bottom with several meters of fine, anaerobic, muddy sedi-
Laticauda crockeri. ment layered on top. This fine sediment renders the depths
unsuitable for foraging, and restricts both sexes to foraging
in the shallow areas on the edges of the lakes or around the

Figure 12.7. Rennel Island Sea Krait, Laticauda crockeri.

Photography by Jonathan Q. Richmond..

many small islands within the lake, as these shores are comprised of pitted limestone, with
far less fine sediment than the lake floor (Cogger, et al, 1987). Type locality: Lake Te-Nggano
on Rennell Island.
Identification. The most melanistic of the Laticauda, with 22-35 faint, alternating
bands on the body, which vary from dark brown to a dark maroon band or a dark muted
blue band alternating with black bands. As with all Laticauda, pupils are round, nostrils
lateral and nasals are separated by internasals, rostral is undivided, with a prefrontal shield
usually present. The mid body has 19–21 longitudinal rows of scales and the average animal
has 192 – 210 ventral scales. Males have 24-39 subcaudals and females 24 – 30. Snout-vent
length averages for males are 615mm and 795 mm for females. The average tail length in

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 12. Sea Kraits

males is 91 mm and 90 mm for females (Uetz et al, 2020).

Natural History. One hypothesis on the origins of this species is that Laticauda laticau-
data once inhabited the Solomon Islands, and Laticauda crockeri descended from this now
extirpated population, diverging from the parent population after the formation of Te-Ng-
gano Lake from a former lagoon (Cogger, et al, 1987). Though closely related to Laticauda
laticaudata, L. crockeri is sympatric with Laticauda colubrina. One hypothesis as to how two
species of krait could share a relatively small area is prey preferences. Laticauda crockeri are
only known to eat one of the two native species in Lake Te-Nggano, the Dusky Sleeper Goby
(Eleotris fusca). The other native fish, Pacific Shortfinned Eel (Anguilla obscura) seems to be
eaten exclusively by the lake’s population of Laticauda colubrina. Unfortunately, neither have
been documented preying on the lake’s invasive species, Mozambique Tilapia (Oreochromis
mossambicus).
Breeding of this species is presumed to be seasonal, but nothing else is known about the
species reproduction. Cogger states that there are local reports of viviparity in the species, but
these claims have yet to be confirmed (Cogger, et al, 1987). The species is listed by the IUCN
as Vulnerable since its entire extant of occurrence is a single lake. The lake is listed as a World
Heritage Site and provides the species some protection (Lane & Guinea, 2010a).

Niue Sea Krait

Laticauda schistorhyncha (Günther, 1874)
Distribution and Habitat. Known only from the small island nation of Niue, where it in-
habits and area less than 300 km². Habitat preferences are poor-
ly studied, studies from Niue are absent however the species is
quite abundant along the reefs, and numerous sea caves on the
island’s coast. (Personal comm. Dr. Jessica Cramp). Type locali-
ty: Niue Island.
Identification. The head is black with a thin off-white mask
Figure 12.8. The distribution over the eyes. Black bands are wider than the alternating blue
of Laticauda schistorhyncha. bands and this species has an overall lower band count than oth-
er sea kraits. Laticauda schistorhyncha and Laticauda semifasci-
ata can be distinguished from the rest of the genus by the rostral
scale, which is divided horizontally in these two species. Laticauda schistorhyncha and Lati-
cauda semifasciata can most easily be distinguished from one another based on band count,
Laticauda semifasciata having 30-42 body bands, compared with Laticauda schistorhynchus
having 18-31 body bands. As with all Laticauda, pupils are round, nostrils lateral and nasals
are separated by internasals (Cogger and Heatwole, 2005).
Ventral scales 195-205 compared to Laticauda schistorhyncha having 187-195 ventral
scales (Guinea, et al, 1983). Males average 38–40 subcaudal scales and females 32–35. The
average snout-vent length in males is 670 mm and 750 mm in females. The average tail length
in males is 100mm and 110 mm in females.
Natural History. Also known as the Flat-tail Sea Snake and locally known as the Katuali,
the species is abundant in sea caves around and beneath the island (Jess Cramp per comms).
Despite its local abundance, this species is listed as vulnerable by the IUCN due to its restrict-
ed range and vulnerability to severe weather events, coral bleaching, and the effects of climate
change, all of which reduce habitat complexity and prey abundance, as well as create a loss of
ideal refuge sites (Lane and Guinea 2010b). Though an official population study has yet to
be conducted, locals have noted a decline in population densities (Personal communication.
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Aquatic Snakes

Figure 12.9. Niue Sea Krait, Laticauda schistorhyncha.

Photography by Kirby Gonzalo Morejohn.
Dr. Jessica Cramp). The venom of Laticauda schistorhyncha is comprised of erabutoxins with
an LD50 of 15 mg/g IV in mice (Guinea et al. 1983). Little is known about this species repro-
duction and natural history since much of Niue’s marine environment has not been studied
due to its remote location and difficulty accessing due to the lack of safe harbors (Friedlander
et al. 2017). The venom of Laticauda schistorhynchus and Laticauda semifasciata comprise
erabutoxins with an LD50 of 15 mg/g IV in mice (Guinea et al. 1983).

New Caledonia Sea Krait

Laticauda saintgironsi (Cogger and Heatwole, 2005)
Distribution and Habitat. Predominantly known from New
Caledonia where it is locally abundant, with single specimen
sightings in the Caroline Islands and New Zealand. Like other
members of the colubrina complex, this species spends most of
its time in shallow reefs and coastal waters. It can be found both
at sea, under rocks, within crevices, inside sea caves, and taking
Figure 12.10 The distribu-
tion of Laticauda saintgi-
shelter under various forms of cover on land. Type locality: Porc-
ronsi. Red area marks the épic Island, New Caledonia.
breeding population. Identification. Prior to its description, Laticauda saintgi-
ronsi was often confused with juvenile and subadult Laticauda
colubrina, not only because of the two species’ morphological
similarities, but also because the two species can be found in the
same together in the same locations including terrestrial aggregations and aquatic feeding

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 12. Sea Kraits

Figure 12.12. Laticauda saintgironsi.

Photography by Pauline Fey.

sites (Cogger and Heatwole, 2006).

A uniquely colored member of the genus, with 21-30 glossy black bands alternated by
burnt orange bands which fade to a pale cream color ventrally. The scales of the light bands
have a dark grey tipping to them, as do the margins of the head and lip scales. Like it’s close-
ly related Laticauda colubrina, Laticauda saintgironsi has a yellow or cream upper face mask
pattern, and an undivided rostral scale. Average ventral scale count is 211-224 in males and
215-230 in females. Maximum snout to vent length is 817mm in males and 1090 mm in
females. The average tail length in males 78.9 mm and 84.2 mm in females. Males average
37–42 subcaudal scales and females 25-34. Average mid-body scale count is 21 rows in
both sexes (Heatwole et al 2005). It can be distinguished from Laticauda semifasciata and
Laticauda schistorhyncha by its undivided rostral. It is easily discernible from Laticauda
laticaudata and Laticauda crockeri by its yellow or cream face mask pattern.
Geographic separation makes the species identification simple from Laticauda col-
ubrina simple, with no known overlap, however it may also be distinguished by a lower
mid-body scale row count, and lower ventral scale counts. Another key difference is the
dark body bands on Laticauda saintgironsi often do not meet ventrally, or at least narrow
ventrally. Coloration is also a notably different, with Laticauda saintgironsi most common-
ly taking on a brown and orange banded color scheme and Laticauda colubrina most often
coming in a blue and black banded color scheme.
Natural History. This species spends most of its time in shallow waters foraging for
eels, with brief periods on shore hiding in litter, under boulders, in crevices, and vegetation.
Heatwole and Cogger report this species may travel some hundreds of meters inland on
the islands and up to 100m in elevation. This species is often found in aggregations of 30
or more individuals in single refuges and in the hundreds in some lagoon’s small islands.
Laticauda saintgironsi Females are thought to breed on average every two years with a mean
clutch size of 3.3 eggs. (Bonnet et al, 2014)

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Guinea’s Sea Krait

Laticauda guineai (Heatwole et al 2005)
Distribution and Habitat. This species is endemic to Papua
New Guinea. Habitat preferences are like that of the closely relat-
ed Laticauda colubrina. Shallow reef shelfs, small islands, rocky
shores, and at sea, as well as among roots and leaves on shore.
Type locality: Bava Island, Papua New Guinea
Identification. Closely related to Laticauda colubrina but is
Figure 12.13. The distribu- identifiable from all other members of the Laticauda colubrina
tion of Laticauda guinea. complex by having a higher number of black bands (47 or more)
which are notably more narrow than the light colored bands and
do not reduce towards the venter as much as in the others. The
species is sexually dimorphic in size, and tail length (Heatwole, et al, 2005).
Natural History. Given the recent description of this species, not much is known of it’s
natural history, however it’s close relation to Laticauda colubrina and the similarities with
in the complex, we can extrapolate some presumptions. Other members of the complex use
tongue-flicking to forage reefs for their prey, which almost exclusively consists of eels. Land
is used to shedding, breeding and egg laying. This species is listed as near threatened by
the IUCN. It is associated with coral reefs, and the loss of habitat due to coral bleaching
and elevated sea temperatures is likely to pose a threat to the species. As corals bleach, prey
abundance and loss of refuge will likely impact this species future success. (Lane and Guinea,
2010d)

Dwarf Sea Krait

Laticauda frontalis (de Vis, 1905)
Distribution and Habitat. Endemic to the islands of Efate
and Espiritu Santo in Vanuatu. Two individuals have been found
at the Loyalty Islands in New Caledonia, but these are presumed
vagrants and these islands are not considered to be a part of the
species range. (Lane and Guinea, 2010c). This species is often
found in seaside rock crevices and beneath mangrove trees when
Figure 12.14. The distribu- on land. At sea it can be found among coral reefs, and sea caves
tion of Laticauda frontalis. like the sympatric L. colubrina. Type locality: New Guinea.
Identification. A small Laticauda with a SVL of 293–717
mm, and a tail length of 34-88 mm with no notable sexual di-
morphism in this species. The species has 28-36 dark bands on the body and 3-4 dark bands
on the tail. There is no difference in the width of the bands with the sympatric Laticauda
colubrina (Heatwole et al 2005).
It can be distinguished from Laticauda semifasciata and Laticauda schistorhyncha by its
undivided rostral. Laticauda frontalis can be distinguished from Laticauda laticaudata and
Laticauda crockeri by its yellow upper lip. A characteristic shared with Laticauda colubrina
based on a lower midbody scale count of 21, lower ventral count and smaller size. Also, on
Laticauda colubrina there is a lower lateral connection between the black head and the first
black band on the neck, which is not present in Laticauda frontalis. Furthermore, the dark
colored body bands on Laticauda frontalis do not extend to the middle of the venter anteri-
orly. (Heatwole and Cogger 2005)
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 12. Sea Kraits

Natural History. It is the smaller sister to Laticauda colubrina a larger relative that feeds
exclusively on eels. This species is listed as Near Threatened by the IUCN due to continuing
decline in the area, extent, and quality of habitat. Other major threats include anthropogen-
ic disturbances such as coastal development, which disturbs nesting. This species predom-
inantly uses the intertidal zone. Rising sea levels and coastal development will impact its
survival (Lane and Guinea 2010c).

Blue-lipped Sea krait

Laticauda laticaudata (Linnaeus, 1758)
Distribution and Habitat. This species inhabits a no-
tably large range, occurring from northeastern India (in-
cluding the Andaman Islands, and the Nicobar Islands),
Malaysia, Indonesia, New Caledonia, the Loyalty Islands,
Papua New Guinea, the southwestern Pacific islands, Chi-
na, Taiwan, and Japan. (Lane et al 2010). Type locality: In-
Figure 12.15 The distribution diis. Habitat included coral reefs, on small coral islands, in
of Laticauda laticauda. coastal mangroves and the open ocean. It is usually found
at a depths of 0-15 m (Cogger 2007). It has been recorded

Figure 12.16 Laticauda laticauda. Photography by Jens Petersen.

to dive to depths greater than 80 m (Brischoux et al. 2009).

Identification. The upper lip of this species is dark brown to black with no pale yellow
lip, helping distinguish it from members of the colubrina complex. Colors in the species
vary across the range from a rich royal blue and black to a muddled dark brown and tan,
or even a dark black and beige. Total length for the epics is 910 mm in males and 1070
mm in females with an additional 110 mm in tail length for both sexes. Ventral scales are
notably enlarged ranging from third to more than half the width of the body. Nasal scales
are separated from intranasal scales, rostral is undivided, and no azygous prefrontal scales
are present. The species has 225-243 ventral scales, and 30-47 subcaudal scales, 38-47 in
makes, and 30-35 in females (Leviton et al 2018).

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Aquatic Snakes

Natural History. Laticuada laticauda are less terrestrial than members of the L. colubrina
complex, yet more terrestrial than the L. semifasciata complex, however it is slower moving
than both on land and water (Wang et al 2013).

Black-banded Sea Krait

Laticauda semifasciata (Reinwardt, 1837)
Distribution and Habitat. This species is found in the Philippines, Taiwan, China, Japan,
and Palu Api in Indonesia and inhabits shallow tropical seas and coral reefs, as well as rocky
shores. Type locality: Moluccas.
Identification. Colors range from dark grey, or black
bands alternating with varying saturation levels of blue bands,
including some very bright blue examples in parts of the spe-
cies range. Body bands notably narrow ventrally. This species
average SVL is 805 mm for males, and 1190mm for females.
Figure 12.17. The distribution It has an average tail length of 115mm for both males and
of Laticauda semifasciata. females. Ventral scales 195-205 and 32-43 subcaudal scales.
(Uetz et al 2020).

Figure 12.18 Laticauda semifasciata.

Photography by by Yu Sing Ling.

Natural History. Laticauda semifasciata is a fish specialist preying primarily on rovers

(Emmelichthyidae), tangs (Acanthuridae), and damselfish (Pomacentridae). Prey items are
subdued by the species venom, which is contains Erabutoxins

On the next page. A Turtle-headed

Seasnake, Emydocephalus annulata.
Photography by Pauline Fey.

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Aquatic Snakes

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Aquatic Snakes

13. Seasnakes
Myke Clarkson

The true sea snakes of the sub-family Hydrophiinae compose about 60 species of poorly
studied marine snakes with paddle tails and front fangs. While more scientists have been ex-
amining specimens and diving with snakes to observe their behavior in the last few decades,
there is still much to learn. These are the only living snakes to have successfully colonized the
oceans, but, of these species, most are restricted to the waters of the continental shelf near
sources of freshwater. All the species studied to date are live-bearing and females produce
relatively small litters, usually with less than 17 young and often as few as one to three young.
Megan Kerford and colleagues studied the movements of the Bar-bellied Sea Snake (Hy-
drophis elegans) at Shark Bay in Western Australia. Bar-bellied Sea Snakes are specialist preda-
tors on snake eels (Family Ophichthidae) that live in burrows on open sand flats. During high
tides, Tiger Sharks have access to most of the areas of Shark Bay, and Keford and co-workers
discovered H. elegans move into adjacent sea grass beds at high tide. The dense grass cover
provides few opportunities to forage on eels but does provide cover for the snakes to avoid
sharks.
Photoreceptors are usually associated with the head of vertebrates, but Kenneth Zimmer-
man and Harold Heatwole found photoreceptors on the tail of the Olive Sea Snake (Aipysurus
laevis). They observed the Olive Sea Snake’s tail was more often concealed during the day than
at night, and that the tail will be pulled out of the light when it is exposed. By masking parts of
the tail with tape so they could not be stimulated by light, they determined the photoreceptors
were located mostly on the dorsal portion of the tail. The nature of the photoreceptors was not
determined but the snake’s behavior revealed their presence.
Only one sea snake has become truly pelagic, drifting with open ocean currents, the Yel-
low-bellied Sea Snake (Hydrophis platura). This is not to say that it does not occur over the
continental shelf, because it occasionally gets washed up on beaches from Africa’s east coast
to the coastlines of western North and Central America. The Yellow-bellied Sea Snake aggre-
gates along slicks or drift lines. Floating debris accumulates in the slicks, and it may remain
for days or weeks before a change in wind speed or current direction breaks them up. Aggre-
gations of snakes in these drifts numbered from five to several thousands and are composed
of juvenile and adult snakes. Other animals inhabiting the drifts are jellyfish medusa, fish,
porpoises, and sea turtles, with sea birds often following the lines of floating debris. Snakes
aggregated here because the slicks are a useful place to locate food and mates. The degree to
which these snakes have adapted to the marine environment is significant given they repre-
sent a recent evolutionary radiation.
Sanders et al. (2008) used mtDNA as well as nuclear genes to examine the relationships of
the Australasian elapids and the sea snakes and estimated the time sea snakes diverged from
the other elapids. They found the sea kraits to be the sister to all other hydrophiines, and the
Melanesian Small-eye Snake (Micropechis) the sister to the remaining species. They also re-
covered a clade containing the true sea snakes, the black swamp snakes (Hemiaspis), and the
Tiger Snakes and their relatives (Notechis group).
Which of these groups form the sister to the sea snakes was unclear? However, of interest
is that tiger snakes are known to eat frogs and forage in water or along shorelines, and the
black swamp snakes are semi-aquatic. Pre-adaptations to life in the water are widespread in
this clade. The time the ancestral sea snake diverged from the terrestrial or freshwater mem-
bers is estimated at 6.2 Ma (7.9–4.7Ma) and the authors write that this is, “...an extremely brief
interval to generate ~60 species of great ecological and morphological diversity.”
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 13. Seasnakes

Short-nosed Seasnake
Aipysurus apraefrontalis (Smith, 1926)
Distribution and Habitat. Ashmore Reef, Hibernia Reef, Ningaloo Reef, Exmouth Gulf,
Barrow Island, and Shark Bay, Australia (D’Anastasi et al 2016). This species occurs in the
shallow waters around coral reefs, and in subtropical seagrass beds with a maximum estimat-
ed depth of 10 meters. They are associated with sections of reef with sparse coral and sandy
bottoms (D’Anastasi et al 2016, Voris 1972). Type locality: Ashmore Reef
Identification. A medium size snake of 108cm, SVL is 985mm (Storr et al 1986). Body is
slender and cylindrical in shape and color is a dark olive to purplish brown in base color with
variable faint olive-brown cross bands. In some specimens these bands are rather conspic-
uous, with scattered olive tipped cream-colored scales creating a speckled like appearance.
Body scales heavily overlap, and the anteriorly facing edges are pointed and are loose from
the body. They may be smooth or possess tubercles or a short keel posteriorly. Dorsal scale
count at mid-body is 17. The head is small and pointed with large symmetrical head shield
present with some fragmentation. Parietal shields are divided and larger than the neck scales.
Prefrontal scales are absent with the nasal coming in direct
contact with the frontal shield. The eye of this species has
a small black pupil with a cream-colored iris. Throat scales
are small and off white in color anteriorly and dark brown
posteriorly. Ventral is most often uniform grey-brown with
occasionally scattered white or white blotched scales. Ventral
scales are deeply notched with a median keel and often worn
Figure 13.1. The distribution of hind margins. Ventral scale count is 140-155. Subcaudals are
Aipysurus apraefrontalis. all single with a count of 18-25 and cloacal scale is divided
(Wilson and Swan 2013) (Cogger 1975). Aipysurus aprae-
frontalis can be distinguished from Aipysurus foliosquama by
lacking prefrontal scales and having 17 midbody rows as opposed to 19-21 in A. foliosquama
(Storr et al 1986).
Natural History. This species actively searches sparse coral and sandy bottoms for fish
with an affinity for eels and gobies (Voris 1972). They are commonly encountered on reef flats
hiding in dead coral rubble at low tide or along the shallow waters of the reef ’s edge or resting
beneath coral overhangs 1-2 meters below the surface during the daylight hours (Lukoschek
et al 2010a) (Cogger 2000). Once feared extinct until the species 2015 rediscovery by Sanders
et al, this critically endangered species has a very restricted range in Western Australia and
is in decline. While the cause of the severity of the decline remains unclear, climate change
induced degradation of shallow coral reef habitats resulting in bleaching events have a sig-
nificant impact on this species and its future as do trawling fisheries (Lukoschek et al 2010a).

Figure 13.2 The Short-nosed

Seasnake, Aipysurus apraefron-
talis. From Smith 1926.

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Aquatic Snakes

Reef Shallows Seasnake

Aipysurus duboisii (Bavay, 1869)
Distribution and Habitat. In Australia the species is known from Northern Territory,
Queensland and Western Australia, Timor Sea in Indonesia, New Caledonia, and Southern
Papua New Guinea. (Uetz and Hosek 2020) This species occurs in reef adjacent shallow wa-
ters, including sandy bottoms, sea grass beds, gorgonian coral
fields, and among dead broken corals. While it is typically a
shallow water dweller, they have been recorded as deep as 80
m although this is a rare exception (Lukoschek et al 2010b).
Type locality: Life, Loyalty Islands, New Caledonia.
Identification. This is a medium sized species with an
Figure 13.3. The distribution average total length of 114 cm, SVL 1010 mm (Stor et al 1986).
of Aipysurus duboisii. Body color is variable, most often with a purplish-brown base
color with pale scale edges, giving the snake a reticulated or
net like appearance. Triangular blotches often form bands down the dorsal of the snake, be-
coming narrower towards the ventral. With exception to the rostral and nasal scales, all scales
on this species have a shallow posterior notch and a mild median keel. Scale are imbricate,
and smooth with keeled tubercules. Midbody scale row count is 19 (Wilson and Swan 2013).
Ventrals number 154-181 and are approximately half as wide as the body. Subcaudals number
25-30 and cloacal plate scale is divided (Storr et al 1986).
Natural History. This snake is commonly encountered with a great deal of biofouling
made up of algae, bryozoans, tube worms, and polychaetas (Lukoschek et al 2010b). Their
diet is comprised of small reef fish including members of the families Muraenidae, Blennidae,
Scaridae, Acanthuridae, scorpaenid, and the toxic Tetrarogidae as well as eels (McCosker
1975). This species is the third most toxic known snake known to science when the venom
is administered subcutaneously with an LD50 of 0.044 mg/kg (Fry 2002). Reproduction oc-
curs every four years with gestation lasting six to seven months, and with the birthing of 4–6
offspring occurring between March and June (Fry et al. 2001). While this species is currently
listed as Least Concern by the IUCN, populations are in decline. Its patchy distribution, and
low densities do leave it vulnerable to future threats including climate change and pressures
from prawn fisheries (Lukoschek et al 2010b).

Marbled Seasnake
Aipysurus eydouxii (Gray, 1849)
Distribution and Habitat. This species is known from the South China Sea, Borneo,
Western Malaysian, Vietnam, Gulf of Thailand, Cambodia, Singapore, Papua New Guinea,
and Thursday Island in Australia. This species frequents turbid waters from 0-50 meters in
shallow bays and estuaries and is associated with muddy substrate but may be found along
hard sea floors as well. It is speculated this species may also swim upstream into rivers (Guin-
ea et al 2010). Type locality: Indian Ocean.
Identification. This is a small snake with a length averaging around 60 cm but up to
91cm long, SVL is up to 790mm (Stor et al 1986). Body is an even diameter throughout and
cylindrical in shape with imbricate scales. The body color is brownish or olive green dorsally
fading to a paler color ventrally, with 30–42 irregular yellow or brown bands (Sanders et al
2012). Banding and band count is highly variable, but often narrows ventrally. Scales within
bands usually with dark margins. Dorsal scales number 17 on the neck and narrow down to
229
 13. Seasnakes

Figure 13.4. Reef Shallows Sea snake, Aipysurus duboissi from Anse Vata, Nouméa, New Caledonia.
Photography by Jean Roger.

230
Aquatic Snakes

15 by the vent. Six lower and upper labials are presents, with the
fourth bordering the eye. The frontal scales of this species are
longer than the tip to snout length. Prefrontals are divided and
range in count from two or occasionally four, with two sets one
preocular and one post ocular. 10-11 Maxillary teeth behind the
fangs. Males possess 129–137 ventral scales, 135-146 for females.
Figure 13.5. The distribution Subcaudal count is 30–32 in males 23–27 on females. Both male
of Aipysurus eydouxii.
and female possess a median keel, however in males these devel-
op into spinous tubercles across their body (Leviton et al 2014).
Aipysurus eudouxii be distinguished from Aipysurus mosaicus, based on a higher band count,
Aipysurus eydouxii having 29 or more and Aipysurus mosaicus having 22 or less body bands
(Sanders et al 2012). Body row count distinguishes Aipysurus eydouxi from Aipysurus duboi-
sii, Aipysurus fuscus, Aipysurus laevis, and Aipysurus pooleorum which all have more than
17 scale midbody rows. This species can be distinguished from Aipysurus apraefrontalis and
Aipysurus foliosquama due
to its lack of deep median
notches on the posterior
edge of the ventral scales. It
can be distinguished from
Aipysurus tenuis by ven-
tral count, having a ventral
count of 129–146 scales as
compared to Aipysurus ten-
nis having more than 170
ventral scales (Sanders et al
2012) (Storr et al 1986).
Natural History. While
many members of the ge-
nus Aipysurus possess some
of the most potent ven-
oms known among snakes,
Aipysurus eydouxii has
evolved away from venom
in a similar evolutionary
fashion to Emydocephalus
annulatus. This is believed
to be because Aipysurus ey-
Figure 3.6. The Marbled Seasnake, Aipysurus eydouxii. The snake
douxiii is a fish egg special- was on the sand bar of Chek Jawa Singapore in the early morning.
ist, searching its habitat for It lmay have been stranded there at low tide. Photography by Ria
benthic fish eggs, which do Tan, Wild Singapore,
not require venom to be sub-
dued. With seasnake venom believed to have evolved for predation and not defense, the di-
etary shift to fish eggs has led to highly atrophied venom glands with reduced and ineffective
fangs resulting in a species that is essentially harmless (Fry et al 2005). This species is listed as
Least Concern by the IUCN, however a recent survey in Shoalwater Bay, Australia indicates
regional population decline, and the species does face threats including climate change and
as by catch in fisheries throughout its range (Guinea et al 2010).

231
 13. Seasnakes

Leaf-scaled Seasnake
Aipysurus foliosquama (Smith, 1926)
Distribution and Habitat. Ashmore Reef, Hibernia Reef, Scott Reef, Shark Bay, and
Barrow Island Australia. This species is associated with tropical reefs, sub-tropical sea grass
meadows, sand over limestone, silt, and sponges (D’Anastasi et al 2016). Type Locality. Ash-
more Reefs, Australia (Uetz and Hosek 2020).
Identification. A small seasnake measuring 55 cm long, with an SVL of 465 mm (Storr
et al 1986). Body color is a light brown to pinkish tan with a variable pattern that can include
faint bands or scattered spots. The head shields of this species are large and symmetrical,
with some fragmentation occurring around the parietal region with prefrontal scales pres-
ent. Body scales are smooth to tuberculate and have notable overlap. The edges of the body
scales have one to two points on each and are extensible free posteriorly creating the leaf
scale like appearance. Mid-body scale row count is 19-21. Ventral scales are nearly half as
wide as the body, notably keeled, and notched posteriorly, with a count of 139-153 (Storr et al
1986). Subcaudal count is 20-29. Cloacal scale is divided. (D’Anastasi et al 2016) (Wilson and
Swan 2013). Aipysurus foliosquama appears similar to Aipysurus
apraefrontalis but can be distinguished by having prefrontals
and 19-21 midbody scale rows as opposed to 17 as in A. aprea-
frontalis (Storr et al 1986).
Natural History. This species is an active predator of small
reef fish, poking its head into crevices and holes, and then
Figure 13.7. The distribution striking at fish hiding with in them. The species is thought
of Aipysurus foliosquama. to usually inhabit depths of no more than 10 m and as such
the populations have limited gene flow between them due to
deep water barriers surrounding the localities they
are known by (Lukoschek 2010c). Once abundant at
Ashmore reef and Hibernia reefs, this species went
missing from 2001 despite several dedicated surveys
in search of them. With these small locals compris-
ing their entire known geographic range the species
was believed extinct until Sanders et al rediscovered
the species at Barrow Island in 2015 (Sanders et al
2015). Today it is listed as Critically Endangered by
Figure 13.8 Aipysurus foliosquama from the IUCN, and while the causation of the species ini-
Ashmore Reef. Photography by Scott tial decline is still not fully understood, impacts from
Eipper. climate change including increased sea temperatures,
coral bleaching, and coral reef habitat degradation are
suspected threats.

Dusky Seasnake
Aipysurus fuscus (Tschudi, 1837)
Distribution and Habitat. Ashmore, Hibernia, Cartier, Scott, and Serangipatan Reefs in
the Timor Sea Western Australia. This species prefers reef and reef edge habitats ranging
from 0-12 m, but up to 30 m deep (Lukoschek et al 2010d). Type Locality. Celebes.
Identification. A short and stout member of Aipysurus with a typical length of 60cm

232
Aquatic Snakes

(though up to 96cm has been recorded), and an SVL of 250-823mm. Aipysurus can superfi-
cially resemble its sister taxon Aipysurus foliosquama, with a shorter more punctuated snout.
Coloration is brown to dark brown or purplish-brown. Pattern is most often uniform, al-
though faint cross bands can be observed towards the lower dorsal
scales. Some individuals feature a darkening in the center of each
lateral scale, causing a longitudinal striation pattern. This species
has some variable head scales including occasional prefrontals,
and most often having frontal scales, but not all. Additionally, this
species usually has a single preocular though occasionally there
Figure 13.9 The distribu- can be two and labial scales vary between 6-9. Consistent features
tion of Aipysurus fuscus. on the head includes: the presence of supraoculars and parietals
which are divided, and small temporal scales Body scales are
smooth and imbricate, with 19-21 rows of dorsal scales at midbody. Ventral scales number
155-180, and have a slight notch, 20-40 undivided subcaudals, and the cloacal scale is divided
(Storr et al 1986) (Cogger 2000).
Natural History. This species is a shallow water specialist that searches the reefs edge
in search of prey items including reef fish like gobies, eels, wrasses, and their eggs. Being a
shallow water species, Aipysurus fuses is only known from five disjunct subpopulations, with
extraordinarily little gene flow suspected between the subpopulations. All these subpopula-
tions have suffered drastic declines in the past few decades, with some estimates putting the
population decline at 70% since 1998. This species’ decline coincides with overall seasnake
declines on the reefs of the Timor Sea, where it is endemic, and it is listed as endangered with
the IUCN. While many seasnakes are stereotyped for having a calm disposition, this species
is known to bite defensively when threatened, and should be approached with caution as it
does possess a potent venom (Dunson 1975).

Olive-Brown Seasnake
Aipysurus laevis (Lacèpéde, 1804)
Distribution and Habitat. Northern seas to Exmouth Gulf off Western Australia, North-
ern Territory Australia, Southern New Guinea, and New Caledonia (Storr et al 1986). Like
other Aipysurus, this species is commonly associated with coral reefs however, unlike many of
its sister taxa can be found in a wide range and depth of habitat including reef slope, reef edge,
sandy bottoms, shipwrecks, deep reefs, inter-tidal habitats, and lagoons. While this species
may dive as deep as 133 m (Crowe-Riddell et al 2019), it more typically occurs at depths rang-
ing from 10 to 40 m (Lukoschek et al 2010e). Type locality: Locker Island, Western Australia.
Identification. Aipysurus laevis is the longest and heaviest member of the genus, with
a total body length ranging from 1.2 m up to nearly 2 m long and up to 23 cm in girth.
Midbody scale count distinguishes this species from all other
Aipysurus except for Aipyrsurus pooleorum, which it can be dis-
tinguished from by the lack of tubercles in males, its larger size
and its lighter coloration. Aipysurus laevis can be distinguished
by other members of the genus based on its larger size, lack of
tubercles in males and paler coloration. This species is high-
Fugure 13.10. The distribu- ly variable in color and pattern, ranging from dark brown to
tion of Aipysurus laevis. purplish-brown, often fading to a lighter pale brown centrally,
with a speckling of lighter colored scales often increasing to-
wards the ventral creating a sort of mosaic gradient. In lighter
233
 13. Seasnakes

specimens the ventral can be entirely off-white. Like other Aipysurus, the center of each scale
is often darker than the edges, creating a lateral striation look. The tail can vary from white
to brown, with dark brown along the dorsal ridge. Prefrontals are occasionally present, and
frontals are often present. Supraoculars and parietals are always divided, and temporals are
small. Preocular scales range from 1–3, though most often 1–2. And upper labial scale counts
number 8–10. Dorsal scales are smooth and imbricate, numbering 21–25 at midbody. Ventral
scales number 142–152, have slight posterior notch, and about are half the width of the body.
Subcaudals number 22–30 and cloacal plate is divided (Storr et al 1986) (Cogger 2000). This
species goes through an ontogenetic change, and young individuals are notable lighter in
coloration than their adult counterparts (Lukoschek et al 2010e).
Natural History. The Olive Seasnake is an active nocturnal hunter, which searches a
home range of reef for crabs, prawns, fish eggs, and fish. Olive Sea Snakes are naturally curi-
ous and are known to approach SCUBA divers, not aggressively but inquisitively, especially
at night (Lukoschek et al 2010e). Individuals of this species maintain often overlapping linear
home ranges of about 0.18 hectares for females, and 0.15 hectare for males along the edges of
the reef. While there is an documented consistency with home ranges, their daytime shelters
do not appear to be consistent (Marsh et al 1993) (Lukoschek et al 2010e). While naturally
curious, and inclined to approach divers for closer investigation, this species is known to bite
when it feels threatened or is restrained. This large species possesses the second most toxic
known snake venom when injected intramuscularly, with an IM LD50 of only 0.09 mg/kg
and human fatalities are known from Aipysurus laevis (Fry 2002) (Dunson 1975). While this
species is believed to be in decline, its large geographic range and wide variety of habitats have
resulted in a Least Concern listing by the IUCN (Lukoschek et al 2010e).

Figure 13.11. Aipysurus laevis. Baie de magenta, Nouméa, New Caledonia.

Photography by Pauline Fey.

234
Aquatic Snakes

Mosaic Seasnake
Aipysurus mosaicus (Sanders et al, 2012)
Distribution and Habitat. This species is known from the Coburg Peninsula, the Gulf of
Carpentaria, Thursday Island, Townsville, Swain Reefs, subtropical Queensland, and Shoal
Water Bay Australia, as well as West Papua (Figure 13.12) This species is associated with off-
shore waters, estuaries and tidal rivers where it is often found among soft sand bed and mud
bottom substrates. It is most often encountered between 2-22 m, with a maximum known
depth of 50 m. Type locality: Gulf of Carpentaria, Weipa, Australia.
Identification. This species is named for its dark and light scales arrangement which
creates a mosaic-like pattern of irregular cross-bands. The body
is a yellowish cream to salmon color, with brown to olive green
incomplete bands, tampering dorsally, numbering 17-22 in
males and 15-12 females. Scale row count at mid-body is 17,
ventral scales number 140–154, subcaudals 25-38, and cloacal
plate is divided. Head dark brown above and below. Aipysurus
Figure13.12. The distribution
mosaicus can be distinguished from Aipysurus eydouxii, based
of Aipysurus mosaicus.
on a lower band count, Aipysurus mosaicus having 22 or less
body bands and Aipysurus eydouxii having 29 or more. Aipy-
surus mosaicus has a concave front parietal suture compared to Aipysurus eydouxi where
it is flat and a crescent-shaped nasal rather than triangular in Aipysurus eydouxi. Aipysurus
mosaicus also has a larger premaxilla, and a smaller maxilla than sister taxon Aipyrsurus ey-
douxi. Aipysurus mosaicus can be distinguished from Aipysurus apraefrontalis and Aipysurus
foliosquama due to its lack of deep median notches on the posterior edge of the ventral scales.

Figure 13.13. Aipysurus mosaicus from Cooloola Qld, Australia.

Photography by Josh Jenson.
It can be distinguished from Aipysurus tenuis by ventral count, having a ventral count of 140-
154 scales as compared to Aipysurus tennis having more than 170 ventral scales. Body row
count distinguishes Aipysurus mosaicus from Aipysurus duboisii, Aipysurus fuscus, Aipysurus
laevis, and Aipysurus pooleorum which all have more than 17 scale midbody rows (Sanders
et al 2012).
Natural History. Like its closest relative, Aipysurus eydouxi, Aipysurus mosaicus is a ben-

235
 13. Seasnakes

thic fish egg specialist and as such has highly reduced venom glands and delivery. With this
species being new to science, much of what is known about it based on prior studies of Aipy-
surus eydouxi. Although other seasnake species in its range are in decline, this species has not
been assessed by the IUCN.

Shark Bay Seasnake

Aipysurus pooleorum (Smith, 1974)
Distribution and Habitat. Endemic to Shark Bay Australia with strays occurring as far
south as Perth, Australia (Wilson and Swan 2013) (Figure 13.14). This species is associated
with limestone reef, seagrass meadows, stromatolite, macroalgae fields, sand bottoms, rocky
rubble, and artificial human made structures like jetties and rock walls (D’Anastasi et al
2016). Type locality: Shark Bay, WA.
Identification. A moderately sized member of the genus at 114 cm long and 19 cm in
girth. Females are a dark brown to purplish brown with
oblique pale bars, and males are brown tone which radiates
to a lighter brown to tan on the flanks (Storr et al 1986).
Prefrontals are rare, frontals occasional, 1-2 preocular pres-
ent, 2-3 post oculars present, 8-10 labial scales, typically
3 primary temporal scales, supraoculars and parietals di-
vided. Dorsal scales are imbricate and smooth in females,
Figure 13.14. The distribution of but males possess tubercles, increasing in size on the lower
Aipysurus pooleorum. rows. Midbody scale row count is 20-23, ventral scale count
is 146-159. Ventral scales have a median keel and posterior
notch and are approximately half as wide as the body. Subcaudals number 25-33 and cloacal
plate is divided (Storr et al 1986).
Natural History. Aipysurus pooleorum are active foragers, taking advantage of areas with

Figure 13.15. Aipysurus pooleorum.

Photography by Scott Eipper.

236
Aquatic Snakes

high structural complexity which provide an abundance of prey and numerous sites for shel-
tering within. Aipysurus pooleorum has not been assessed by the IUCN, however conser-
vationists do believe studies on this small ranging species should be of the highest priority
(D’Anastasi et al 2016).

Arafura Seasnake
Aipysurus tenuis (Lönnberg and Anderson, 1913)
Distribution and Habitat. The true geographic range of this species is poorly known,
but is thought to include the Northwestern coast of Australia from Broome to the Dampier
Archipelago on the Pilbarra Coast though recent data suggests it can occur as far out as
Ningaloo Reef and Exmouth Gulf (Figure 13.16). It is associated with sandy bottom, seagrass
meadows, coastal rock outcroppings, reef, and marine man-made structures (Guinea et al
2010b; D’Anastasi et al 2016). Type locality: Cape Jaubert, Broome, Australia.
Identification. A large member of the genus, up to 1.3
meters long. Body color is light brown with dark brown scale
tips that create longitude lions down the dorsum and/or faint
cross bars on the sides. This species has smooth imbricate
dorsal scales that number 19 at midbody. Males possess tu-
bercles, increasing in size on the lower dorsal scales (Cogger
Figure 13.16. The distribution 2000). Preocular scale is single, however the frontal, supraoc-
of Aipysurus tenuis. ular, and parietal scales are all divided. Two postoculars are
present, and the labial scales number 7-6. Ventrals number
185-194, with a medium notch and a keel,
which is stronger in females than males.
Subcaudals number 36-27 and the cloacal
plate is divided (Storr et al 1986).
Natural History. Very little is known
about this poorly studied species, due
to it’s rarity, and often misidentification
with its sister taxon. By 2010 it was only
known by four specimens (Guinea et al
2010b), though more recent studies by
D’Anastasi et al found 11 during sur-
veys. Aipysurus pooleorum has not been
assessed by the IUCN, however conser-
vationists do believe studies on this small
ranging species should be of the highest
priority (D’Anastasi et al 2016).

Figure 13.17. Aipysurus tenuis. Photogra-

phy by Rick Stuart-Smith, [Link]-
[Link].

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 13. Seasnakes

Turtle Headed Sea Snakes - Emydocephalus

There are three members of the genus Emydocephalus all of which are diurnal, medium
sized, heavy bodied hydrophiini elapid snakes which feed exclusively on the eggs of small de-
mersal spawning fish found through chemosensory foraging (Shine et al, 2020). Due to their
exclusive diet of fish eggs, the species in this genus lack palatal, maxilla, and dentary teeth,
though they do have small teeth on the pterygoid. Members of this genus also have unique
fangs which face inward toward the midline, rather than the traditional vertical positioning
pointing downward making them ineffective (Voris, 1966). With a specialized diet of eggs,
Emydocephalus have evolved away from having fangs and a venom apparatus, making them
the only marine elapid virtually harmless to humans due to their degenerated venom system
(Minton, 1983). Emydocephalus do have a high quantity of scale sensilla compared to terres-
trial snakes, which are thought to act as a form of tactile mechanoreceptors, as well as poten-
tially functioning as hydrodynamic receptors which sense water displacement (Crowe et al
2016). Males of this genus possess a rostral spine, which was initially suspected to be used for
the consuming of fish eggs, but later revealed to be used in mating by the males, utilizing the
spine to prod potential mates in courtship.
Instead of using the rostral spine to scrape eggs off surfaces as was previously suspected,
Emydocephalus use their consolidated upper labial scales to scrape eggs off surfaces (Guin-
ea, 1996). While Emydocephalus is closely related to Aipysurus and placed within the Aipy-
surus-Emydocephalus clade with in Hydrophiini, Emydocephalus have not been shown to
possess the light sensing phototactic tail that is present in Aipysurus laevis, Aipysurus duboi-
sii, and Aipysurus tenuis. All members of this genus are dimorphic, with females attaining a
larger total size than males, and males possessing the before mentioned rostral spine, which
is absent in females (Shine et al 2020).
While members of this genus are mostly banded or blotched, Goiran et al found that
individuals living in polluted urban adjacent waters tend to be melanistic. Gorian and his
team concluded that the darker skin allows the snakes to excrete the trace elements by bind-
ing these elements to the melanin and disposing of them when shedding their skin. In turn
animal inhabiting these polluted waters slough more frequently than those in cleaner waters,
as mechanism for expelling pollutants (Gorian et al 2017).
Members of this genus occur in the shallow marine waters of Australia, New Caledonia,
the Philippine, Vietnam, and Japan. Two of the three species are strongly associated with
coral reefs, with the recently described Emydocephalus orarius being more closely associate
with (Shine et al, 2020) soft bottomed trawl grounds. There is no range overlap between the
species in this genus, making identification between the three species simplified (Crowe et
al 2019).
Foraging in the genus is conducted in an unusual slow and continuous browsing fashion
in a small, restricted home range (Gorian et al 2020). Gorian et al studied the effects tides
had on this behavior and found the genus alters where it forages based on the tides, but tides
do not make a significant impact on activity Levels. In this study, the authors found that this
species is more likely to be associated with rocky or coral habitats during high or medium
tides and sandier habitats during low tides (Gorian et al 2020).
Reproduction is seasonal, ovulation and mating occurring September- October, with
young born in May, accounting for an 8-month gestation. Neonates grow rapidly with sexual
maturity in males occurring at two years and three years for females. Emydocephalus only
reproduces every two to three years and their litter size is two offspring (Shine et al 2020).
Currently no members of this genus are threatened with extinction, however the most

238
Aquatic Snakes

recent assessment by the IUCN has noted that populations of Emydocephalus annulatus are
decreasing (Lukoschek et al, 2010).

Banded Turtle Headed Seasnake

Emydocephalus annulatus Krefft, 1869
Distribution and Habitat. Timor sea, New Caledonia, the Loyalty Islands Vietnam, Phil-
ippines, and Australia (Lukoschek et al 2010). This species has a highly disjunct range, how-
ever only slight mitochondrial and nuclear divergences occur across the range (Nankivell et
al, 2020). Type locality: Unknown, but probably the Australian Seas (Uetz and Hošek, 2020).
This species is associated with shallow clear ocean waters and
coral reefs, usually seen in water less than 40 meters in depth
(Lukoschek et al 2010).
Identification. Members of the genus is characterized by
having three upper labials, with the second one being very long.
This feature alone can distinguish them from other members
Figure 13.18. The distribution of the Hydrophiini group. As mentioned, prior, the species
of Emydocephalus annulatus. only has rudimentary teeth. Adult males posses a rostral spine
used in courtship which is absent in females. This species col-
oration is highly variable and can range from banded patterns
of off white to yellow, alternating with dark rings or a patternless dark grey to black. The
scales of this species are smooth and overlapping with 15–17 rows at the mid-body. This
species has 125–145 wide slightly keeled ventral scales and 20–33 subcaudal scales. Cloacal
scales are undivided. Adult males have small tubercles on their ventral scales. Average adult
size is 75 cm (Department of the Environment, 2021).
Natural History. Forages slowly over coral reefs and rocky environments, searching
cracks and crevices for fish eggs using scent to locate their prey, which is exclusively fish eggs
from the fish genus Gobiidae, Pomacentridae and Blenniidae. This species uses modified
labial scales to break the fish eggs from the surfaces they are adhered to (Department of
the Environment, 2021). During low tide Emydocephalus annulatus can become dangerously
entrapped in shallow pockets, leaving them suscecptible to overheating, and as such most
individuals leave the reef flats during low tide, and travel through drainage channels to reach
the deeper waters at the edges of the reef (Department of the Environment, 2021).

Ijima’s Turtle Headed Seasnake

Emydocephalus ijimae Stejneger, 1898
Distribution and Habitat. This species is associated with shallow clear ocean waters and
coral reefs but known from waters as deep as 40 meters (Lukoschek & Sanders 2010). China,
Taiwan, Japan. Type locality: Ryuku Islands, Japan
Identification. Members of the genus are characterized by having three upper labials,
with the second one being exceptionally long. As mentioned, prior, the species only has ru-
dimentary teeth. Adult males possess a rostral spine, which is absent in females. This species
has an average of 141.7 ventral scales, 32.3 subcaudal scales, and an average of 18 dorsal scales
at the midbody. When present the species has 24–29 bands, however coloration and pattern is
highly variable, and ranges from alternating light tan to yellow bands against a black to dark
grey background, or partially banded, or solid black in color (Nankivell et al, 2020).

239
Aquatic Snakes

Figure 13.19. The Turtle-headed Sea Snake, Emydocephalus annulatus. (A) a banded morph from
Baie des Citrons, Nouméa, New Caledonia, (B-C) a black morph from Baie de Magenta, New
Caledonia. Photography by Pauline Fey.

240
Aquatic Snakes

Natural His-
tory. Both Emydo-
cephalus ijimae and
Emydocephalus
orarius share simi-
Figure 13.20. The distri- lar natural histories,
bution of Emydocephalus
preying specifically
ijimae.
on fish eggs. Their
primary diet is goby and blenny fish eggs.
These are often laid attached to sandy surfaces,
and as such many animals were found to have
sand in their colons (Voris, 1966). Masunga
et al noted seasonal weight fluctuation in this
species, which could be attributed in part to
both prey availability and sea temperatures
with males attaining higher body weight ear-
ly spring to late summer (November to March
or April), when fish spawning is at its peak.
Conversely, body weights lessen in the winter,
though more dramatically in females, which Figure 13.21. Emydocephalus ijimae. Photog-
could be attributed in part to parturition raphy by Patrick Davis.
(Masunaga & Ota, 2003).

Western Turtle Headed Seasnake

Emydocephalus orarius Nankivell, et al. 2020
Distribution and Habitat. Endemic to Western Australia, confirmed from Shark Bay,
Exmouth Gulf, Pilbara coast and Broome (Nankivell et al, 2020). Type locality: Shark Bay,
WA, Australia. Unlike other members of this genus which are mostly associated with rocky
and coral areas, Emydocephalus orarius is associated with shallow ocean water with soft sandy
bottoms (Nankivell et al, 2020).
Identification. Members of the genus is characterized by
having three upper labials, with the second one being long.
This feature alone can distinguish them from other members
of the Hydrophiini group. As mentioned, prior, the species
only has rudimentary teeth. Males possess a rostral spine, ab-
Figure 13.22. The distribution sent on females. The males of this species also possess ventral
of Emydocephalus orarius. tubercles on adult individuals.
This species has an average of 144–145 ventral scales, an
average of 32.3 subcaudal scales, and an average of 17 dorsal
scales at the middy. Emydocephalus orarius can be distinguished from Emydocephalus ijimae
and Emydocephalus annulatus based on the first upper labial scale. In Emydocephalus orarius
the first upper labial scale is elongated and contacts the preocular, where as in Emydocephalus
ijimae or Emydocephalus annulatus, it is usually the second upper labial that is elongated.
Some specimens may also have an elongated first upper labial, in which case comparing ven-
tral counts. Emydocephalus orarius has an average of 144.7 compared to an average of 136.9
in Emydocephalus annulatus. This species can be distinguished by neighboring Emydoceph-

241
 13. Seasnakes

alus ijimae in having two prefron-

tal scales compared to the three
to four prefrontal scales found
in Emydocephalus ijimae. Adult
males of this species possess tu-
bercles on their ventral scales,
absent on Emydocephalus ijimae
(Nankivell et al. 2020).
Natural History. Emydoce-
phalus orarius diurnally forage
for small demersal spawning
fish eggs which it finds through
chemosensory foraging (Shine
et al 2020) in an unusual slow
and continuous browsing fash-
ion (Gorian et al. 2020). Their
primary prey, goby and blenny
fish eggs, are often laid attached
to sandy surfaces, and as such
many animals were found to
have sand in their colons (Voris
1966).

Arafura Smooth Seasnake

Parahydrophis mertoni
(Roux, 1910)
Distribution and Habitat. Figure 13.23. The Western Headed Seasnake of Emydoceph-
Papua New Guinea in the Ara- alus orarius. Lighthouse Bay, Exmouth, WA, Australia Pho-
fura Sea, and Northern Territory, tography by Alex [Link] Glen Whisson.
Queensland, Australia. This spe-
cies occurs in the coastal mangrove, estuarial mangroves, and mudflats (Heatwole 1999).
Type locality: Sungei Waskei, Wokam, Aru Island, Indonesia .
Identification. Blue gray, to olive brown dorsal patter with 40-50 irregular darker cross
bands joined at the midline, with a lighter center in each band. Head shields have yellow
spots except for on the rostral and labials which are black (de Rooij N 1917). The average
length is 50 cm. The body scales are smooth and imbricate. Dorsal scale count is 36 to 39
rows at the mid-body, ventral scale count is 153 to 161, subcaudal scales count is 29 and
35, and the cloacal scale divided. Ventral scales are character-
istically large in this species. Adults grow to a total length of
50 cm (Cogger 2000). No sexual dimorphism in SVL or tail
length seems present (Parkin and Schembri 2011).
Natural History. Parahydrophis is a monotypic genus
closely related to Ephalophis and sister to Hydrophis (Sanders
et al 2013). Both monotypic genera represent early lineages
Figure 13.24. The distribu- in the subfamily Hydrophinae and as such retain many of
tion of Parahydrophis metoni. the more primitive traits of land snakes such as a cylindrical
head and widened ventral scales. Unlike its other close rela-

242
Aquatic Snakes

tive Hydrelaps which forages

terrestrially at the waters edge
on mudflats, Parahydrophis
forage in mangroves and tidal
creek banks for small fish. The
species has also been cited as
foraging the mudflats at night,
when submerged at high tide.
Average litter size is three,
though it is undetermined if
breeding occurs throughout
the year or is seasonal (Parkin
and Schembri 2011).
Figure 13.26. Ephalophis [Link] by Victoria de Bruyn
© Broome Community Seagrass Monitoring Project.

North-western Mangrove Seasnake

Ephalophis greyae (Smith, 1931)
Distribution and Habitat. Endemic to Australia’s Northwestern coast from the Kimberley
region to Shark Bay and is associated with mangroves, estuarine mudflats and coastal shallow
flats, found at depths from 0-10 meters. (Lukoschek et al. 2010) (Heatwole 1999) Type local-
ity: Cape Boileau, Western Australia.
Identification. The body scales have a median keel forming 19 to 21 dorsal scale rows
at mid-body, 159 to 171 ventral scales, and 27 to 33 sub caudal scales with the cloacal scale
divided (Cogger 2000).
Natural History. Ephalophis is a monotypic genus closely related to Parahydrophis and a
sister to Hydrophis (Sanders et al. 2013). Both monotypic genus represent one of the earliest
lineages in the subfamily Hydrophinae and as such retain many of the more primitive traits
of land snakes such as a cylindrical head and widened ventral scales. This species forages the
waters edge at high tide, and the out of the water on the sand
flats at low tides in search of mudskippers (Oxudercidae), its
primary prey. Voris also includes gobies (Gobiidae) as a prey.
The snakes will consume both the mudskipper and its eggs
and has been observed entering the mudskippers burrows in
its search. It has also been observed entering crab burrows in
Figure 13.25. The distribuition search of hiding gobies (Voris & Voris 1983; Lukoschek et al
of Ephalophis greyae. 2010).

Port Darwin Seasnake

Hydrelaps darwiniensis (Boulenger, 1896)
Distribution and Habitat. Endemic to Australia occurring in the coastal waters of
Queensland, Northern Territory and Western Australia. Type locality: Port Darwin, North-
ern Territory, Australia
Identification. Twenty-five to 30 dorsal scale rows at mid body 163 to 72 ventral scales,

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 13. Seasnakes

and 27 to 39 subcaudal scales which may be single but occa-

sionally divided anteriorly. Cloacal scale is divided. (Cogger
2000)
Natural History. This monotypic genus and a sister
lineage to Hydrophis (Sanders et al 2013). Guinea et al. ob-
served specimens crawling onto mud flats and into crab
Figure 13.27. The distribution of
Hydrelaps darwiniensi.s burrows in search of prey, during high tide at 15:00 h. This
indicates this species feeds diurnally, and likely feeds on
land where its movement is adept for a sea snake. Guinea
also observed specimens drinking from shallow upper tidal
freshwater pools indicating that like Laticuada, Hydrelaps
may use these rainwater pools as a water source (Guinea et
al 1993).

Bighead Seasnake
Hydrophis annandalei (Laidlaw, 1901)
Distribution and Habitat. Found in the Gulf of Thailand
in Cambodia south to Indonesia, including Peninsular Ma-
laysia, Singapore, Java and Sumatra in Indonesia and Viet-
nam. A record from Brunei however is suspected to have
potentially been a washed-up vagrant and may not be a part
of the species extant range. Throughout its range the species Figure 13.28. Hydrelaps dar-
is associated with shallow muddy coastal waters with one winiensis. From Boulenger
freshwater record in the Pattani region in Thailand (Rass- 1896.
musen et al 2010). Type locality: Patani Bay, Thailand.
Identification. As the common name indicates, this species possesses a notably large
head for a sea snake, with irregular head shield and a stout body (Lim 1991). Some note-
worthy squamation detail includes a W-shaped rostral scale which is longer than it is wide,
a triangle shaped mental scale, and a large scale beneath the eye between super labials 5
and 6. Small distinct ventrals are present which number 363 plus 65 subcaudal scales Body
scales are small and highly irregular with a high body scale
count of 62-82 scale rows at the neck, and 74-97 body scale
rows. The body has 46 dark body bands against a more pale
gray backdrop, which narrow on the sides, and can fade with
age to a darker dorsal, lighter ventral gradient pattern (Uetz
and Hosek 2020) (Das 1993).
Figure 13.29. The distribution Natural History. This species is known from shallow
of Hydrophis annandalei. muddy coastal waters. Prey includes members of the family
Clupeidae which includes herrings, shads, sardines, hilsa,
and menhadens, based on stomach content studies (Voris and Voris 1983).

Anomalous Seasnake
Hydrophis anomalus (Schmidt, 1852)
Distribution and Habitat. Found in the South China Sea including Malaysia, Vietnam,
Sumatra, Java, and Borneo (Uetz and Hosek 2020). Type locality: Listed as “Rhede von

244
Aquatic Snakes

Samarang”, Java.
Identification. This species has a light whitish base ton with 30-36 darker bands which
taper dorsally creating a triangular shape. A pair of elongated shields separate the nasals, the
rostral is divided into 4-5 scales. rostral divided into four or
five scales. Dorsal scale rows at mid body are 31–35 and ven-
trals number 210– 256 (Chan et al. 2015).
Natural History. Formerly place in the monotypic genus
Thalassophios, little is known about this species ecology and
behavior, outside of it’s association with estuaries (Chan et al.
Figure 13.31. The distribution 2015). Stomach sample studies have shown Conger eels (Con-
of Hydrophis anomalus. gridae) to be one of the prey items of this species (Voris &
Voris 1983).

Figure 13.32. Hydrophis anomalus. From Schmidt 1852.

Black-Headed Seasnake
Hydrophis atriceps (Günther, 1864)
Distribution and Habitat. This species is known Indone-
sia, Malaysia, Vietnam, Thailand, Singapore, China, Taiwan
and the Philippines, and Papua New Guinea, Northern Ter-
ritory and Western Australia. Older literature includes Myan-
mar and India in this species distribution; however these re-
Figure 13.33. The distribution cords should actually be referred to as Hydrophis fasciatus. It
of Hydrophis atriceps. is associated with turbid off shore waters over sandy bottoms
5-10 km from shore at depths of 20-30 m (Rasmussen et al
2010a; Leviton et al 2003). Type locality: Thailand.
Identification. This species head is small, with a long body that becomes slender ante-
riorly. The body is a light yellow with black banding that narrows dorsally and may or may
not connect at mid body. In juveniles the bands may be complete the length of the body, be-
coming less complete with age. Beneath incomplete bands at mid body, darker spots may be
found. The head and neck are a dark olive or black color. Dorsal scales at the thickest part of
the body are hexagonal in shape and maybe be juxtaposed or slightly imbricate. Dorsal scale
245
 13. Seasnakes

row count is 25-30 at the neck, and midbody scale row count is
39-49. Ventral scales count is 323-452. This species is slightly
size dimorphic with the total length in males coming in at 1100
mm and females 990 mm (Leviton 2003).
Natural History. Little is known about the natural history
of this species. Their prey is primarily comprised of eels that
hung along sandy sea bottoms at depth of 20-30m (Sherratt et
Figure 13.34. Hydrophis atri- al. 2018). Smith reported this species is also commonly associ-
ceps. From Cao et al. 2014. ated with the mouths of rivers (Smith 1926).

Faint-Banded Seasnake
Hydrophis belcheri (Gray, 1849)
Distribution and Habitat. Known from south east Asia (Cambodia, Indonesia, Malaysia,
Philippines; Singapore; Thailand; Vietnam), Papua New Guinea, and questionably in Aus-
tralia, where only the type specimen was found. It is presumed to occur in waters over soft
bottoms of depths between 0-15 meters (Rasmussen & Sanders 2010; Cogger, 2000). Type
locality: Listed as New Guinea, though some authorities question this locals validity.
Identification. This species pattern consists of 52-70 olive-gray bands over an off-white/
yellowish backdrop. Bands widen dorsally and fade ventrally ventrally with an off-white to
white ventral coloration. This species has a medium sized head, narrow neck and elongate
body which is compressed posteriorly, and two to four times the diameter of the neck. The
head pattern is an olive green with a yellow hotshot shaped spot
running through e prefrontal scales and around the eyes. The
fourth supralabial borders the eye, though in rare occasions
two supralabials can contact the eyes, either 3 and 4 or 4 and
5. Dorsal scale count rows are 24-
27 at the neck and 32-37 at mid
body. Ventrals number 278-313
(Kharin & Cheblukov 2007).
Natural History. Hydrophis
belcheri is an eel specialist, pri-
Figure 13.35. The distribution of marily feeding on worm eels
Hydrophis belcheri. (Moringuidae) and other eels
(Anguilliformes). Though er-
rantly rumored to be the most lethal snake on the planet, the
actual LD50 when tested intramuscularly is 0.07 mg/kg which
does make this species the fifth most toxic snake when tested in-
tramuscularly (Voris & Voris 1983; Rasmussen & Sanders 2010;
Tamiya and Puffer 1974; Steinhoff, 2018).

Peters’ Seasnake
Hydrophis bituberculatus (Peters, 1872)
Figure 13.36. Hydrophis
Distribution and Habitat. Known from Phuket harbor in belcheri. From Boulengeri
Thailand and Colombo in Sri Lanka. It is thought to live over 1896.
sandy gravel bottoms, but little is known of its actual habitat

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Aquatic Snakes

preferences. (Rassmussen & Lobom 2010a) Type locality: Co-

lombo, Sri Lanka.
Identification. The head of this species is blackish above,
with a lighter whitish grey color at the supralabials, and a light
grey color around the eyes. The body is banded with black bands
and significantly narrower light colored (grey/white) interspac-
es. Males have 37-51 bands on the body, 6-10 on the tail and fe-
Figure 13.37. The distribu- males 38-50 bands on the body, 6-8 on the tail. These interspaces
tion of Hydrophis bituber- are wider ventrally than dorsally, where they average 1-1.5 scales
culatus. where they join at the top. Interspaces are narrow anteriorly, wid-
en through the body, and narrow once more posteriorly. Banding
continues on to the tail where the interspaces become thinner, approximately one scale wide.
The first two interspaces from the head are often not joined. Embryos showed a higher color
contrast than their adult counter parts (black bands with white interspace), and the holotype
showed almost not pattern, though it is unclear whether this was an older animal showing
high ontogenetic shift, or if alcohol bleaching was to blame. Dorsal scales feature a two crest-
ed median keel across the entirety of the body, with 25-29 scale rows at the neck and 43-50
scale rows at midbody. Ventral scales are distinct, approximately twice as wide as the adjoin-
ing scales, narrowing posteriorly, and number 247-290. Average total length is 110 cm in both
males and females, with only slight size variance between the sexes, with the largest recorded
male having 1 cm longer tail than the longest recorded female (Rasmussen 1992).
Natural History. From its initial description in 1872 by Peters, until Rasmussen’s re-
discovery of the species in Thailand in 1992, the holotype was the only known specimen.
Rasmussen examined 25 specimens caught by trawl from 1987-1989 (8 females, 17 males),
though the fisherman did not disclose exact positions of their fishing grounds, so despite
the additional specimens, little is still known about this species habitat preferences. Stomach
contents of the examined specimens showed five potential species of eels (Anguilliformes),
though contents were to digested to determine which species of eels. One of the eight females
contained three full term embryos.
Four females in total were collected
after, and none of others with em-
bryos, indicating November may be
the end of the species birthing period
(Rasmussen, 1992).

Brook’s Small Headed

Seasnake
Hydrophis brookii (Günther,
1872)
Distribution and Habitat. This
species is known from Indonesia,
Malaysia, and Thailand known from
subtidal muddy regions and estuar-
ies. It is also reported from a fresh-
water lake in Thailand (Rasmussen
et al 2010g). Type locality: Sarawak,
Figure 13.38. Hydrophis bituberculatus. From Peters 1873 Malaysia (Uetz and Hosek 2020).

247
 13. Seasnakes

Identification. The head of this species is very small,

black in color, with a yellow hotshoe mark on top. The
body is long and slender anteriorly which compresses
posteriorly becoming to three times the diameter of the
neck. The body is bluish white with 60-80 dark grey to
black bands which are complete anteriorly and twice the
width of the interspace, but become incomplete ventral-
Figure 13.39. The distribution of ly down the posterior of the body. Body scales are slight-
Hydrophis brookii Below. Figure ly imbricate, and hexagonal in shape often showing a me-
13.40. dian tubercle. There are 23 to 31 scale rows at the neck,
and 37-45 at midbody. Light colored ventrals are distinct
and number 328-414. Adult size is 104cm (Leviton et al. 2014; Rasmussen et al. 2011b).
Natural History. Little is known about this species. Stomach content studies reveled
prey items to include gobies (Gobiidae), worm eels (Moringuidae), and snake eels (Ophich-
thidae). (Rasmussen et al. 2010f; Voris and Voris 1983)

Figure 13.40. Hydrophis brookii .Photography by JCM (left) Figure 13.41 A Vietnamese stamp with
an artistic representation of Hydrophis brookii (right).

Dwarf Seasnake
Hydrophis caerulescens (Shaw, 1802)
Distribution and Habitat. This species is wide spread,
common in South East Asia it is known from Myanmar, Ma-
laysia, Indonesia, Singapore, and Vietnam, but rare in south
Asia including Pakistan, India, and Bangladesh. To the north
it is known from the South China Sea as well as the coasts
of Shandong and Guandong in China. It is also known from
Figure 13.42. The distribu-
Queensland Australia, and New Caledonia. It is known from
tion of Hydrophis caerules-
cens. depths from 0-25 meters above seagrass beds and muddy
subtidal regions (Rasmussen et al 2010e). Type locality label
cites as “Indian Ocean: Vizagapatam.”
Identification. As the name indicates this is a smaller species of sea snake with the
average reported length being 0.6cm. The head of this species is black with enlarged head
shields. The third and fourth supralabials contact the eyes. The body of this species is is
blue-grey above, with 41-43 black blotches or transverse bands, with 7-9 bands at the tail.

248
Aquatic Snakes

These bands taper ventrally and can become

indistinct in older adults. Body scales are
feebly imbricate and number 37-39 at mid-
body. At the thickest part of the body, these
scales are quadrangular or hexagonal in
shape. Ventral scales are barely wider than
adjacent body scales and the ventral scale
count is 266-287 (Cogger, 2000).
Natural History. Voris and Voris found
prey to include true gobies (Gobiodidae),
worm eels (Moringuidae), and burrowing
gobies (Trypauchenidae) in stomach con-
tent. Little is known about the life cycle of
this species. Fry et al reported a single brood
containing seven young. (Voris & Voris
1983; Fry et al. 2001)

Figure [Link] caerulescens. Photog-

raphy by Prathamesh Ghadekar.

Cantor’s Narrow Headed Seasnake

Hydrophis cantoris (Günther, 1864)
Distribution and Habitat. Pakistan, India, Sri Lanka,
Myanmar, to the western Malay Peninsula and Iran (Heat-
wole 1999; Rezaie-Atagholopour et al 2016). This species
Figure 13.45. The distribution habitat includes shallow coastal waters with soft floors
of Hydrophis cantoris. (Leviton et al. 2003). Type locality: Penang, Malaysia.
Identification. A former member of the genus Micro-
cephalis, this species pointed head and slender neck are considerably smaller than the rest
of their body. The head is yellow in color, and the neck is a darker black color with 20-28
faint yellowish incomplete bands that fade towards the body to a darker olive color dorsally
graduating to a yellow to off white ventrally. The notably slender neck has 23–25 to scale
rows, but as little as 21 have been reported, with a midbody scale count of 41 and ventrals
numbering 404-468. Average total length is 1450 mm for males and 1880 mm for females.
(Leviton et al. 2003; Rezaie-Atagholipoir et al. 2016)
Natural History. Very little is known about this species, and it is considered rare (Rass-
mussen et al. 2010d).

Slender-necked Seasnake
Hydrophis coggeri (Kharin, 1984)
Distribution and Habitat. This species is know from Western Australia, Vanuatu, Fiji,
the Philippines, Indonesia, and New Caledonia. It is associated with shallow coastal waters,
shallow mangroves, coral sea beds, sea grass beds, mudflats, inner reef beds, sea grass fields,

249
 13. Seasnakes

coral reef floors, mangroves and ocean floors with sediments and loose organic materials in
the 1-40 m depth range. (Palomares and Pauly 2021; Ineich and
Laboute 2002). Type locality: Port Suva, Fiji.
Identification. Hydrophis coggeri is a banded sea snake with
a thin cylindrical body, a black tail tip, and no spots or mark-
ings between the darker colored bands. Hydrophis coggeri is dif-
ferentiated from all other Hydrophis species except Hydrophis
Figure 13.46, The distribu- donaldi, Hydrophis sibauensis and Hydrophis torquatus diade-
tion of Hydrophis coggeri. ma based on ventrals that are not divided by a longitudinal fur-
row. This species has 29-30 scale rows the neck, 33-35 scale rows at midbody and 246-288
ventral scales. It can be distinguished from Hydrophis donaldi based on having 30-42 bands
around the body rather than 47-56 in Hydrophis donaldi, and the lack of strongly spinous
body scales as is present in Hydrophis donaldi. Average adult length is 1 m, with a maximum
recorded length of 1.364 m, and females being larger and heavier than males (Cogger 2002).
Natural History. Hydrophis coggeri is a highly nocturnal species which forages the sea-
floor in search of its main prey which includes eels of the families Ophichthidae and Cong-
ridae. In Fiji the preferred prey includes moray eel species (Callechelys melanotaenia, Leiura-

Figure 13.47. Hydrophis coggeri from Baie des Citrons, Nouméa, New Caledonia.
Photography by Pauline Fey.

nus semicinctus, and Myrichthys colubrinus). A specimen feeding on eel tail catfish (Plotosus
anguillaris) has been observed and this may also be common prey. This species gives birth
to 1 to 8 live offspring. Hydrophis coggeri is a highly venomous species, and it’s venom has an
intramuscular LD50 of 0.24 mg/ kg. There have been reports of this species quickly pursuing
those that provoke it and can be considered dangerous to swimmers, divers, and those who
disturb it (Palomares and Pauly 2021; Ineich and Laboute 2002).

Spine-bellied Seasnake
Hydrophis curtus (Shaw, 1802)
Distribution and Habitat. This species is very widespread with one of the most exten-
sive distributions of all sea snakes, Hydrophis platurus being one of the only species more
widespread. Hydrophis curtis range includes the Arabian Gulf (Oman, United Arab Emirates,
Iran, Kuwait, Iraq, Qatar, and Saudi Arabia) all of south Asia (India, Sri Lanka, Pakistan, and

250
Aquatic Snakes

Bangladesh), South East Asia (Indonesia, Philippines, Malaysia, Thailand, Myanmar, Brunei,
Singapore, and Vietnam) and south east to Papua New Guinea and northern Australia. The
furthest southeast recorded specimen was found in New Caledonia, but it is uncertain if this
specimen represents a vagrant individual or a breeding population. This species in known
from waters between 4-55 m, and is adapt to a wide variety of tropical shallow water habitats
including: gulfs, bays, estuaries, and occasional entries into rivers and freshwater habitats.
It is associated with waters over soft-sediments, including sands, muds, and seagrass beads
adjacent to coral reefs (Rassmussen et al 2021). Type locality: None given.
Identification. The presence of large hook like keels on the ventral and lower dorsal
scales, but absent on the upper dorsal scales easily distinguishes this species from most oth-
ers. These large hook-like spines are more prominent in males than females and may play a
role in breeding. The spines in males are most prominent during breeding season, meaning
the rugosity of the scales maybe hormonally controlled as seen in many species of frogs and
fish. This is a short stout sea snake, with three known pattern
variations. Color and pattern are variable. The most common
pattern is a darker grey dorsally graduating to a grey-whitish
ventrally, with lighter grey bands. The two less common pat-
tern and color forms include a yellowish body with black-
ish bands or a blackish grey body with darker black dorsal
Figure 13.48. The distribution of bands. In all three-color forms, the dorsal bands are typically
Hydrophis curtus. fused laterally, creating a zigzag pattern with a black tip of
tail. This species has a large head, with a notably tridentate
rostral scale, with two notches or groves where the tongue
protrudes, rather than a singular notch. The parietal scales on
this species are divided into small shields with 7–9 supralabi-
als with the second supralabial contacting with the prefrontal
scale and the third and/or fourth touching the eye. Body scale
rows number 28– 38 at the neck and 32–43 at the mid-body.
Ventral scales are present and distinguishable, number 147–
201. Ventral scales are larger (half the body width) and hex-
Figure 13.49. The “Great Hy-
drus, Hydrophis pelamidoïdes”
agonal shape anteriorly and becoming smaller posteriorly. The
is a synonym of Hydrophus average total length for this species is 80-110cm in total length,
curtus. The illustration is with no sexual dimorphism in total length. Hydrophis curtus
from Iconographia Zoologica can superficially resemble Hydrophis ornatus, but can be easily
- Special Collections Univer- distinguished from Hydrophis ornatus in having in having a
sity of Amsterdam. zigzag pattern form, rather than the even bands in Hydrophis
ornatus. Another distinguishing distinction is Hydrophis cur-
tus has divided parietal forming small shields not present in H.
ornatus (Buzás et al 2018; Avolio et al 2006; Rezaie-Atagholipour et al 2016).
Natural History. The diet includes a wide range of prey items including amphipods, cut-
tlefish (sepia), squids, and over 33 families of fish ranging from bottom dwelling benthic fish,
near bottom dwelling demersal fish, and even open water pelagic fish. The ability to be both
a habitat and prey generalist has allowed this species to occupy a considerable range. It is
believed that deep waters act as a barrier for dispersal in this species, with an average known
depth of 40 m, and a maximum known depth of 55 m. Genetic studies indicate H. curtus ex-
pansion into Australia and South Was Asia is recent or sudden. Studies in Australia indicate
annual breeding with a 6–7-month gestation period resulting in an average of 4.3 young being
born between March and June. In Australia females select shallow, sheltered areas such as the

251
 13. Seasnakes

Figure 13.50. Hydrophis curtus. Art work from Soedirman(1916).

mouths of bays, estuaries, and sea grass beds during the summer months prior to birthing,
indicating these habitats could be critically important nursery grounds to the species (Udy-
awer et al. 2016). While there is anecdotal evidence this species is in decline in Asia, due to
the widespread spread nature of it as range, it is listed as least concern by the IUCN. Buzás et
al. found the species in the United Arab Emirates, and that it is “generally inoffensive” except
for one specimen that “jumped” out of a transport box to attempt to bite the handler’s glove.
Heatwole reported this species to be “very aggressive if provoked or handled”. The venom
contains postsynaptic nuertoxins and likely myotoxins and should be treated as a potentially
lethal snake (Voris and Voris 1983; Buzás et al 2018; Rassmussen et al 2021; Rezaie-Atagho-
lipour 2012; Heatwole 1999). Taxonomic note: A review of a large number of specimens of
Hydrophis curtus and Hydrophis harwickii did not support speciation based on morpholog-
ical analysis, and while molecular data does show a deep divergence of the Indian Ocean
(H. curtus), and South East Asia Australiasia (H. hardwickii) clades, the authors will defer
to Rassmussen et al’s 2021 determination that Hydrophis curtus be viewed as a single species
until further studies are conducted (Rasmussen et al. 2021).

Annulated Seasnake
Hydrophis cyanocinctus (Daudin, 1803)
Distribution and Habitat. This species has a wide range extending west from the Arabian
Gulf (Iran, Iraq, Kuwait, Oman, Qatar, United Arab Emirates, and Saudi Arabia), throughout
south India (India including Andaman Islands and Nicobar Islands, Sri Lanka, Pakistan, and
Bangaladesh) east to south east Asia (Philippines Indonesia, Cambodia, Thailand, Vietnam,
Palua), north east to the South China Sea (China, Taiwan, and possibly Japan), and south to
northern Australia and Papua New Guinea. It was originally reported present in Solomon
Islands, but the author of this finding later retracted this. This species is associated with shal-

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Aquatic Snakes

low seas over muddy bottom with depths ranging from 0-30
m. In Pakistan they have been seasonally found in shallow
mangrove swamps during monsoons (Minton 1966; Rasmus-
sen et al 2010f). Type locality is listed as “Coromandel” (Uetz
and Hosek 2020). Authors note: Coromandel Coast likely refers
to East India.
Figure 13.51. The distribution Identification. Hydrophis cyanocinctus is a long and very
of Hydrophis cyanocinctus.
slender banded sea snake, with a dark olive or gray body
which lightens ventrally. The darker body bands number 50–75 which are broader dorsally,
tampering to a point ventrally, with widening interspace at the body, which narrows at the head
and tail. This species has juveniles presenting a black head with a yellow horseshoe-shaped
marking on top of the head with the horse-shoe marking fading with age. The head of this
species is slightly enlarged,
and the body is elongate
but not markedly slender
anteriorly. Dorsal scales
are imbricate and at the
thickest part of the body
present round or bluntly
pointed tips, and num-
ber 39–41 scale rows at
mid-body and 25–31
scale rows at the neck.
Ventral scales are only
slightly distinguishable
and number 300–359.
Average total length is
Figure 13,52. Hydrophis cyanocinctus. 1243mm, though excep-
Photography by Anslem da Silva. tionally large specimens,
as large as 1495 mm
have been recorded (Rezaie-Atagholipour et al 2016). Juveniles of Hydrophis cyanocinctus
and Hydrophis lapemoides can be easily confused, however, the mid-body scale shapes are
distinct between these two species. Hydrophis cyanocinctus having bluntly pointed scales,
and Hydrophis lapemoides having hexagonal or quadrangular scales. (Voris and Voris 1983;
Rezaie-Atagholipour et al 2016; Egan 2007).
Natural History. This species is often seen basking on the surface during the days of
warmer months. A 2013 stomach content study in Pakistan’s Hara Protected area revealed
members of the goby family (Boleophthalmus, Parachaeturichthys, Periophthalmus, Scarte-
laos, Taenoides) comprised 77% of the stomach contents, with all but one species belonging
to the mudskippers (Ocudercinae) group. Other known prey items include conger eels (Con-
geridae), pike conger (Muraenesocidae), gobies (Gobioididae), eel catfish (Plotosus lineatus),
and unidentified eels (Anguilliformes). This species produces between 3-16 live young, with
birthing seasons varying through out their range. Like many sea snake species, it is not un-
common to find this species with heavy barnacle infestations of the sea snake specializing
barnacle Platylepas ophiophila. This species has a potent myotonic venom and has caused
human fatalities. While inoffensive in the water may bite if restrained. (Buzás et al 2018; Voris
and Voris 1983; Rezaie Atagholipour et al 2013; Egan 2007).

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 13. Seasnakes

Fine-Spined Seasnake
Hydrophis czeblukovi (Kharin, 1984)
Distribution and Habitat. This species is only known from North and North West Aus-
tralia with a single record from Bam Island on the north coast of Papua New Guinea. This
species has been found as bycatch in deep water trawls and is thought to occur in deep waters
over soft substrate up to 110 m deep (Rasmussen & Lukoschek 2010). Type locality: Arafura
Sea.
Identification. As the common name of the now synonymized Geometrical sea snake
(formerly Hydrophis geometricus) would indicate, specimens of this species can feature an
elaborate and distinctive pattern consisting of two rows of pentagonal like dark grey spots
separated by thin lighter colored interspaces creating a very distinct geometrically webbed
like appearance. The spots consist of 37 large hexagonal spots running down the back, with a
second row of 36 spots, that are smaller and less distinct than the upper row. This patterning
becomes much less pronounced and indistinct ventrally, as well as along the base of the tale.
The head is brown in color, with large sublingual, with.a moderately sized triangular mental
scale. The body scales anteriorly are imbricate, but gradually become hexagonal and without
overlap posteriorly. The scales also graduate from smooth at the neck progressing anteriorly
to a weak keel consisting of 2-4 tubercles, and by midbody the weak keels transition to stout
spines. The neck of this species has 33 scale rows, and 57 scale rows at mid-body. The 306
ventral scales and 45 subcaudal scales are clearly pronounced, being twice the size of the
surrounding scales, with 1-2 slight tubercles present anteriorly.
There are four cloacal shields adjacent to each other with the
posterior shield after the cloacal opening being the largest of
the four (Kharin & Hallerman 2010).
Natural History. This species is considered rare, with very
little known about their natural history. To date, only eight
specimens of this species have been found. (Rasmussen & Lu-
Figure 13.53. The distribu- koschek 2010; Kharin & Hallerman 2010)
tion of Hydrophis czeblukovi. Taxonomy Note: Kharin described Hydrophis czeblukovi
in 1984, based on two female specimens which had been col-
lected by Professor V. P. Shuntov in 1968. In 1986, two Hydrophis geometricus, based on three
specimens from Western Australia, however upon review, Hydrophis geometricus was found
to be synonymous with Hydrophis czeblukovi, and thus not recognized here as a valid taxon
by the authors. Due to the extreme rarity of this species, little is know about its conservation
status or population trends and is listed Data Deficient by the IUCN (Kharin & Hallerman
2010; Rasmussen & Lukoschek 2010).

Rough-scaled Seasnake
Hydrophis donaldi (Ukuwela, Sanders, and Fry 2012)
Distribution and Habitat. Endemic to the Gulf of Carpenteria, Queensland Australia,
where it is commonly associated with estuarine habitats consisting of shale, mud, and/or sea-
grass bottoms. The species has been found at the mouths of the Mission River and Hey Creek
at the connection to the Albatross Bay in Weipa, Queensland. Specimens have been found via
spotlighting from a boat traveling at low speeds in the early evening over water 10 m or less
deep (Ukuwela et al 2012). Type locality: Gulf of Carpentaria, Weipa, Queensland, Australia.
Identification. Hydrophis donaldi is a recently described, smaller sea snake with a total
254
Aquatic Snakes

length between 92-101 mm. The pattern of this sea snake consists of 47-64 dark brown bands
over a lighter yellowish brown interpsace, that are wider than the interspaces. These bands
are broad dorsally which narrow latterally, and do not connect ventrally. The bands are also
darker in coloration in the first third of the body. The head is brown in color in adults, but
one juvenile specimen was found with a black head and a more highly colored body, with
turquoise blue interspaces rather than light grey and black and bands. There are 33-35 scale
rows at midbody and 29-30 at the neck. The 246-288 ventral scales are distinct and bear two
strong spines on each. Hydrophis donaldi is distinguished from all other Hydrophis species
except Hydrophis coggerii, Hydrophis sibauensis and Hydrophis torquatus by having ventral
scales that are not divided by a longitudinal keel or furrow, having 29-30 costal scale rows
around the neck and by having 246-288 ventral scales. From Hydrophis coggeri, Hydrophis
sibauensis and Hydrophis torquatus it can be identified, by
having stronger spines on the body scales, as well as other
morphological variances. Hydrophis donaldi can be distin-
guished from Hydrophis coggeri based on Hydrophis donaldi
having more body bands (37-56 compared to 30-42), a larger
head blunter head, and a lower average ventral scale count
Figure 13.54. The distribution (246-288 compared to 280-360). Hydrophis donaldi can be
of Hydrophis donaldi. differed from Hydrophis sibauensis based on having a higher
number of scale rows at the neck and more scale rows around
the neck (29-30 in Hydrophis donaldi compared to 25-26 in Hydrophis sibauensis). Hydrophis
donaldi can be differed from Hydrophis torquatus diadema in having a lower scale row count
at the neck, (33-35 compared to 35-42) (Ukulele et al 2012).
Natural History. This species was first found by Bryan Fry, and formally described in
2012. It is named after Dave Donald, who was the skipper on the boat at the time of its initial
discovery in 2000. From 2000-2012 only nine specimens were found, making it one of the
more uncommonly seen species in its poorly surveyed range. As such little is known about its
ecology (Ukuwela et al 2012).

Elegant Seasnake
Hydrophis elegans (Gray, 1842)
Distribution and Habitat. Hydrophis elegans is endemic to
the waters around Northern Australia, Indonesian West Papua,
and Papua New Guinea. The species ranges into Australia’s tem-
perate waters as far south in Australia as Kalbarri on the western
coast and Gold Coast on the eastern coast giving it one of the
furthest south distributions of any sea snake species. In terms of
Figure 13.55. The distribu- habitat this species is a generalist, inhabiting waters above sand,
tion of Hydropis elegans. and mud, coral reefs, seagrass beds, subtidal rock, subtidal gravel,
subtidal sand, rocky reef, estuaries, and even up rivers and into
freshwater habitats where it has been found by beam trawlers 12
km upstream from the sea on the Burnett River in Queensland. This species is believed to
prefer depths between 3.7-26 m, however it has been found in water as shallow as 2 m, and as
deep as 250 m. However, Limpus noted the upriver occurrences have only been recorded be-
tween December and January and may be seasonal in nature. The discovery of Hydrophis el-
egans 250 m deep in Browse Basin, Australia, marked the first known records of a sea snakes’
presence in the dark cold deep waters known as the mesopelagic or “twilight” zone, though

255
 13. Seasnakes

the frequency and importance of this deep water behavior is yet

unknown and merits more study (Limpus 1975; Milton et al 2010;
Crowe et al 2019; Department of the Environment, 2021). Type
locality: Port Essington, N. T.
Identification. Hydrophis elegans is the longest of all sea
snakes, with a maximum recorded length of 2.6 m. The body pat-
tern consist of 39-44 blackish bands over a grey or yellowish body,
which are widest along the spine and constrict ventrally. Between
each bands within the yellowish interspace lies a row of smaller
black dots that can produce a secondary black band when joined.
This species has an elongated body compressing posteriorly. The
body scales are imbricate, with 37-49 rows at the mid-body. The
345-432 ventral scales are only slightly wider than the adjacent
scales. Females are generally larger than males, with males attain-
ing an average SVL of 120 - 160 cm and females an average SVL of
140-180 cm. Female Hydrophis elegans also have larger heads than
males, and as such females prey on larger food items than their
male counterparts. Hydrophis elegans can be deciphered from the
sympatric Hydrophis major by having more bands (39-44 in Hy-
drophis elegans compared to 31-40 in Hydrophis major), a longer Figure 13.56, Hydrophis el-
egans from Kretf 1869.
overall length, and a smaller head (Kerford 2005; Department of
the Environment, 2021; Milton et al 2010).
Natural History. Hydrophis elegans is a generalist feeder with a diet including a variety
of bottom-dwelling fish such as Catfish (Euristhmus), moray eels (Muraenidae and Nettas-
tomatidae), snake eels (Ophicthyidae), gobies (Gobiidae), Sillago (Sillaginidae), snake eels
(Ophichthidae), and squid (Teuthoidea). Known predators include tiger sharks (Galeocerdo
cuvier), of which Hydrophis elegans account for 50% of young tiger sharks prey within their
range. Hydrophis elegans reaches sexual maturity around 2 years old, with the oldest known
specimen being 10 years old. This species can reproduce every year or every 2-3 years, with
recorded clutch sizes of 12-13 offspring in the Norther Prawn Fishery and Great Barrier Reef.
Studies have shown the size of the litter, and size of the resulting offspring vary with the
length of the female, with larger females producing larger quantities of bigger offspring (Voris
and Voris 1983; Fry et al 2001; Kerford 2005; Milton et al 2010; Department of the Environ-
ment, 2021)

Striped Seasnake
Hydrophis fasciatus (Schneider, 1799)
Distribution and Habitat. This species is mainly known from southeast Asia where it
occurs in Cambodia, Malaysia, Myanmar, Thailand, and Vietnam, but absent from Borneo.
It also occurs in south Asia where it is considered rare, known from Eastern India and Ban-
gladesh, but absent in Sri Lanka. It is associated with soft bottom coastal waters ranging from
0-12 meters in depth (Rasmussen & Lobo 2010c; Uetz and Hosek 2020). Type locality: None
given.
Identification. The head of this species is a dark olive to black color with yellowish tri-
angular-oval saddles forming a banded appearance of dark inverted triangles which fade to
gray ventrally. Posteriorly the body is grayish dorsally and a whitish color ventrally. Juve-
niles spots appear rhomboidal and can form concentric circles. This is a microcephalic snake,

256
Aquatic Snakes

with a small head, slender anterior and long body. Scales on the
thickest part of the body are hexagonal in shape, and slightly
imbricate, with a mid-body scale count of 47-58 rows, and a
neck scale count of 28-33 rows, noting a dramatic scale count
increase posteriorly. Ventral scales number 414-514. Males of
this species are larger than females, males having a total length
Figure 13.57. The distribu- of 1100 mm of which 100 mm makes up the tail and females a
tion of Hydrophis fasciatus. total length of 990 mm, with 75 mm comprising the tail. (Levi-
ton et al 2003).
Natural History. This microcephalic snake is a nocturnal, shallow water, eel specialist,
feeding on snake eels (Ophichthidae), worm eels (Moringuidae), burrowing (eel like) gobies
(Trypauchenidae), moray eels (Muraenidae), and false moray eels (Xenocongriae) (Voris and
Voris, 1983; Glodek and Voris 1982; Rasmussen & Lobo 2010c; Leviton et al. 2003).

Narrow-Headed Seasnake
Hydrophis gracilis (Shaw, 1802)
Distribution and Habitat. This species is widespread known from the Arabian and Oman
Gulfs (Oman, Iran, Iraq, Kuwait, Bahrain, Qatar, and United Arab Emirates), in south Asia
(Pakistan, India, Sri Lanka, and Bangaladesh), and southeast Asia (Indonesia, Malaysia,
Myanmar, Philippines, Thailand, and Vietnam), north Asia (China, Hong Kong, and Tai-
wan), south to Australia and Papua New Guinea. It is known from waters 0-30 meters deep
and associated with mangrove swamps, intertidal muddy bottom coasts, and coastal reefs.
This species appears to be rather common around eastern India, Borneo, and Northern Java
(Indonesia), but uncommon near Pakistan (Guinea et al. 2010b). Type locality: None given.
Identification. A microcephalic snake, a long slender body anteriorly which becomes
laterally compressed posterior. This species goes through a significant ontogenetic change.
Juveniles have dark lateral spots over a lighter whitish body anteriorly, which become less

Figure 13.58. Hydrophis curtus. Photography Anslem da Silva.

257
 13. Seasnakes

distinctly defined posteriorly. These blotches form a band like pattern with the lighter colored
interspace, creating a count of 40-60 bands. However, this pattern fades posteriorly with age
and the pattern become indistinct, creating an almost uniformly grey dorsal graduating to
a whitish ventral side anteriorly, with a faintly banded body anteriorly. The body scales are
imbricate, numbering 30-36 rows at mid-body, and 12-21 rows at the neck. The scales be-
come keeled with dark brown or black tubercles ventrally. Ventral
scales are divided by a longitudinal fissure and number 220-228.
Females grow larger than males, females attaining a total length of
1025 mm, 95 mm of which is tail, and males reach 950 mm, 80mm
of which is tail (Rasmussen et al 2011; Leviton et al 2003).
Natural History. Hydrophis gracilis, like many microcephalic
Figure 13.59. The distribu-
species, is an eel specialist, foraging sand beds in search of its prey
tion of Hydrophis gracilis.
(Guinea et al. 2010b).

Figure 13.60. Hydrophis gracilis. Photography by JCM.

Hardwicke’s Spine-bellied Seasnake

Hydrophis hardwickii Gray 1834
Distribution and Habitat. Persian Gulf, Indian Ocean, South China Sea north to the
coasts of Fujian and Shandong Strait of Taiwan, Singapore; Indo-Australian Archipelago;
North coast of Australia; Philippines; Pacific Ocean from Myanmar, Thailand, Vietnam, In-
donesia, China, Japan, New Guinea (no map). Type locality: India.
Identification. Hardwicke’s Spine-bellied Seasnake has long been confused with Hy-
drophis curtus. The following has been take from Leviton (2003:436). Body short, stout, neck
region not less than half as thick at midbody; head large; scales squarish or hexagonal, jux-
taposed, outer 34 rows larger than others, scale rows: & 23-31 around neck, 27-35, around
midbody, 3 25-27, 2 33-41; ventrals small, usually distinct anteriorly, not so posteriorly, in ¢
114-186, in 2 141-230; head shields entire, parietals occasionally divided; nostrils superior,
nasals in contact with one another; prefrontal usually in contact with second upper labial; 7-8
upper labials, 3-4 bordering eye; | pre- and 1-2 postoculars; 2, rarely 3, anterior temporals;
greenish or yellow-olive above, whitish below, 35—SO olive to dark gray dorsal bars, tapering

258
Aquatic Snakes

to a point laterally, occasionally encircling body, a narrow dark ventral stripe or broad irreg-
ular band occasionally present; adults often lack any pattern and are uniform olive to dark
gray; head pale olive to black, yellow markings on snout present or not. Total length 860 mm,
tail length 85 mm.
Natural History. Diet. Glodek and Voris (1982) found this species has a diet that is more
diverse than any other sea snake that it occurs with. H. hardwickii had 48 prey species in 21
families of fish as well as squid and cuttlefish (Cephalopoda) , while the other sea snakes it
occurred with had 4-7 prey species.
Relative abundance. Ward (2000) found the catch rates of Hydrophis hardwickii were
highest in the eastern part of the Gulf of Carpentaria at depths of <20 m. Species that repre-
sented ≥20% of specimens from one or more areas were: Hydrophis hardwickii, H. ornatus,
H. major and Aipysurus eydouxii. Catch rates were highest for H. elegans. Catch rates of all
species of snakes combined did not differ significantly between 1984–86 and 1989–90 and
were moderately high around Groote Eylandt where the fishery is centered. Most by-catch
species occur in areas that are not subjected to extensive trawling, and there are no data to
suggest that this fishery seriously threatens any population of sea snake. However, sea snakes
may be more vulnerable to trawler-induced effects than most other by-catch species, and
formal assessment of the status of populations is needed.
Reproduction. The reproductive cycle of H. hardwickii reported by Ward (2001) found
copulation occurs between early May and the end of July; ovulation during August–Sep-
tember; and gestation during September–December. Females produce approximately 8.5
offspring per clutch. Females breed annually. Average litter swize is is 8.4 offspring per year.

Rangoon Sea Snake

Hydrophis hendersoni Boulenger 1903
Distribution and Habitat. Myanmar. Type locality: Ran-
goon, Burma.
Identification. This species was long considered part of H.
nigrocinctus. Rasmussen et al (2011c) removed it from H. nigro-
cinctus based on it having 2-3 maxillary teeth behind the fang
(0-1 teeth in H. nigrocinctus), seven supralabials (eight in H. ni-
grocinctus), 30-33 scale rows on anterior body (27 rows in H.
Figure13.61 The distribution
of Hydrophis hendersoni.
nigrocinctus)

Plain Seasnake
Hydrophis inornatus (Gray, 1849)
Distribution and Habitat. This species is only known from it’s type specimen which was
listed as merely “Indian Ocean”. It is presumed to live as far east as the Philippines and as far
south as the Arafura Sea (Rassmussen 2010). Type locality: “Indian Ocean” (Rassmussen
2010). [no map or photo]
Identification. Body is a bluish grey dorsally and dorsally transitioning to white on the
ventral. The tail is white light black bands with a black or dark grey end of tail. Body is uni-
formly wide. Dorsal scan rows are 44 at midbody with 253 ventral scales which are distinct
and twice as wide as adjacent anteriorly, narrowing posteriorly. It is distinct from the similar
Hydrophis lamberti, and Hydrophis ornatus based externally on color pattern, and internal-

259
 13. Seasnakes

ly by several physiological traits (Rassmussen

1989).
Natural History. Prey include old world
silver sides (Atherinidae) and gobies (Gobi-
oididae), with little else known of the natural
history of this species (Voris and Voris 1983;
Rassmussen 2010).

Sadle-Backed Seasnake
Hydrophis jerdonii (Gray, 1849)
Distribution and Habitat. Occurs in the
Indian Ocean including Sri Lanka, India
and Myanmar,
Southeast Asia
including East
Malaysia, and
Vietnam, and in
the South China
Figure 13.63. The distribu-
Sea including
tion of Hydrophis jerdonii
southern Chi-
na and Taiwan.
Despite being known from India’s east coast,
there have been no records of this species oc-
curring there since 1977 and it is uncommon Figure 13.62. Hydrophis hendersoni from Bou-
in the Andaman Sea. This species is associated lenger (1903).
with gravel and sandy muddy bottoms and has
been found between 0-30 m deep (Rasmussen
& Lobo 2010; Uetz and Hosek 2020). Type
locality: Madras, India.
Identification. This species has a short
head which is gray or greenish in color on
top and white or yellow below. The body is
a lighter olive to gray with 30-38 wide black
bands which narrow laterally and encircle
the entire body. The body is nearly an equal
diameter throughout it’s length, with keeled
and imbricate scales. The scale count at mid-
dy is notably low, less than 23 scale rows. The
ventral scale count varies regionally from
225–278. Total length for this species is re-
ported at 1000 mm (Chan et al 2015.) Fig.13.64. Hydrophis jerdonii. From Smith
Natural History. Formerly in the mono- (1943).
typic genus Kerilia, this species forages the sea
floor peaking its head into crevices and holes in search of prey items which include lizard-
fish (Synodontidae) and anguilliform or eel like fishes. (Chan et al 2015; Rasmussen & Lobo
2010).

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Aquatic Snakes

Spectacled Seasnake
Hydrophis kingii (Boulenger, 1896)
Distribution and Habitat. This species is known from northern Australia, West Papua,
and Papua New Guinea. It is associated with depths ranging from 0-22 meters over soft sandy
and muddy substrates, as well as reef habitat (Milton et al 2010b). Type Locality. Listed as
North Australia .
Identification. Hydrophis kingii has a black head and throat which is followed by banding
which becomes a dorsal pattern of 45-50 darker grey to black
round to rhombic shaped saddle markings with an yellowish to
cream base color forming the innerspaces. The interspace be-
tween these rhombic like saddles can have black lining to each
scale, most prominent dorsally, but fading on the flanks. The
lateral sides have smaller dark spots. Ventrally the coloration
Figure 13.65. The distribu- becomes cream to pale brown with a black mid-ventral stripe
tion of Hydrophis kingii. and a series of dark mid lateral triangular markings which are
pale grey. The head shields are enlarged, symmetrical and un-
fragmented, with a large anterior chine shield bordering the
mental groove. The body scales are keeled and imbricate, with 37-39 dorsal scale rows at
midbody. The ventral scales are notably larger than the dorsal scales and number 324-342.
Average total length of adults is 1.5 m (Per obs; Cogger, 2018).
Natural History. Little is known of this species as it is considered rare. Most have been
found in deeper waters with varying seabeds. Prey include benthic fish including catfish (Eu-
risthmus nudiceps), and eels from the family moray eels (Muraenidae). Females give birth
annually in September after a six to seven months, producing five young on average, with a
maximum recorded litter size of eight (Cogger 2018; Milton et al 2010b; Fry et al 2001; Voris
and Voris, 1983).

Kloss’ Seasnake
Hydrophis klossi (Boulenger, 1912)
Distribution and Habitat. This species is restricted to the Gulf of Thailand, occurring in
Thailand, Cambodia, Malaysia, Singapore and Indonesia. Nothing is known of the habitat
preferences of this species (Sanders & Rasmussen 2010). Type
Locality. Thuala, Selangor, Malay Peninsula.
Identification. This species pattern consists of a greyish to
dark green dorsally, graduating to yellowish ventrally with 50-
75 darker crossbones, with a grayish to greenish head, adorned
with a yellow horseshoe marking. The head scallion includes
1 preocular scale, 1 post ocular scale, and 1 anterior tempo-
Figure 13.66. The distribution ral plate. Supralabials number 5-6, the second being the larges
of Hydrophis klossi. and the 3rd and 4th scales contact the orbit. There are 607
infra labials. Body scales are smooth or with a slight keel and
number 23-27 at the neck and 31-19 at midbody. Ventrals have two tubercles, and number
360-430. Adult length ranges from 975-1190 mm. (Kharin, 2004)
Natural History. Nothing is known of this species habitat preferences or ecology.

261
 13. Seasnakes

Laboute’s Seasnake
Hydrophis laboutei (Rassumussen & Ineich, 2000)
Distribution and Habitat. This species is endemic to the waters surrounding New Cale-
donia and associated with depths from 0-62 meters in depth. Nothing is known about the
species habitat preferences, the holotype was caught on a trawl at
a depth of 62 m (Rasmussen 2010b). Type Locality. Chesterfield
Reefs, New Caledonia (20°21.98’S, 161°04.87’E) (Rasmussen & In-
eich 2000).
Identification. This sea snake species has a black head with a
Figure 13.67. The dis- weak lighter line over the eyes, with 45-46 black bands with white
tribution of Hydrophis interspaces, and 7-8 more dark bands on the tail. One noteworthy
laboutei. pattern feature for this species is that the first two or three black
bands melt together with the black head dorsally. The white in-
terspaces are widest on the body flanks, narrowing dorsally, and
interrupted ventrally by a black ventral surface compressed of 265-
280 ventral scales. Dorsal scale rows number 26-28 at the neck and
44-46 midbody. This species can be distinguished by five allopatric
species (Hydrophis atericeps, Hydrophis elegans, Hydrophis kingii,
Hydrophis pacificus, and Hydrophis vorisi) on its notably low ven-
tral count (265-280), with the other five allopatrics having a ventral
count greater than 311. Hydrophis laboutei can be distinguished
from Hydrophis belcheri 34-37 dorsal scales at the neck, and the
first bands do not fuse together as in Hydrophis laboutei (Rasmus-
sen and Ineich 2000).
Natural History. This species was only recently described
(2000) and is only known from three specimens. Nothing is known
about this species ecology, though it is presumed to be rare (Ras-
mussen 2010b).

Lambert’s Seasnake
Hydrophis lamberti (Smith, 1917)
Figure 13.68. Hydrophis
laboutei. A specimen at Distribution and Habitat. This species occurs through out
MNHN Photographed south east Asia including Cambodia, Vietnam, Thailand, China,
by Antoine Fraysse
Taiwan, Malaysia, Indonesia, Singapore, and the Philippines. It is
- 2017, MNHN Pro-
ject:RECOLNAT (ANR- associated with coastal waters with sandy bottoms (Rasmussen &
11-INBS-0004). Gatus 2010). Type locality: Mouth of the Mekong River.
Identification. This species has a large robust head which is
grayish to brown colored and mildly patterned above and whitish below. The body’s base
color is an off-white to pale grey which lightens ventrally, covered with dark bands which are
widest dorsally and lighten and narrow ventrally in an upside-down triangular shape. Males
have 26-36 bands on the body and 4-6 on the tail, and females have 26-31 bands on the body
and 5-6 on the tail. Ventral scales are distinct, about twice as wide as the adjacent scales an-
teriorly and number 258-395 in males and 237-306 in females. Males have 37-50 subcaudals
whereas females have 34-44. Dorsal scale rows number 37-45 at the neck and 45-56 at the
midbody. Hydrophis ornata has a similar appearance but has fewer than 42 bands on the body
and tail (Rasmussen 1989; Chanard et al 2015; Leviton et al 2014).
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Aquatic Snakes

Natural History. Little is known about this species natural history; it is said to prey on
marine catfish. While this species is caught as by-catch in trawling activities, harvested in the
Philippines for export to Japan as smoked sea snake, and harvested in Thailand and Vietnam
for their skin, it is a widespread species and listed least concern by the IUCN (Das 2010;
Rasmussen & Gatus 2010).

Arabian Gulf Seasnake

Hydrophis lapemoides (Gray, 1849)
Distribution and Habitat. This species is known from the Arabian Gulf including Bah-
rain, Iran, Iraq, Kuwait, Qatar, Oman and the United Arab Emirates. In south Asia it is found
in Pakistan, India, Bangladesh, and Sri Lanka. In South East Asia
it is known from Myanmar, Thailand, Malaysia, and Singapore. It
is associated with shallow warm waters over sandy bottoms, sea-
grass beds, and reefs and is often seen basking at the surface. It has
also been occasionally found in freshwater or brackish estuaries
Figure 13.69. The distri- and as close to shore as rocky sea breaks less than 3 meters deep.
bution of Hydrophis lam- As such this species is frequently found washed up on beaches.
berti. This species is abundant in the Arabian Sea, however uncommon

Figure 13.70. Hydrophis

lamberti. [Link]
[Link]/By Fran-
co Colnago - https://
[Link]/
photos/63668789,
CC BY 4.0, htps://
[Link].
org/w/[Link]?cu-
rid=92604323

to rare throughout the remainder of its range (Rasmussen et al 2010i; Egan 2007; Garner
2013; Buzás et al 2018). Type locality: Ceylon, Madras.
Identification. Hydrophis lapemoides has a variable pattern consisting of 33-64 black or
dark grey bands over a pale light grey to yellow background. These bands are widest dorsally
and taper ventrally, narrowing down the flanks. The rings face ventrally and maybe faded or
completely absent in older specimens. The black head has a yellow or off-white horseshoe
pattern. This species is a stout Seasnake with a moderate sized head with the widest part of
the body 2-3 times wider than the neck. The body scales are keeled and hexagonal in shape,
with each scale being as broad as it is long. Towards the tail the scales become more distinctly
keeled and widen, in males these posterior keels becoming spike like tubercles. This species
could be confused with Hydrophis ornatus, however the banding on Hydrophis ornatus termi-
nate in a sharp point. This species also could be confused with Hydrophis cyanocinctus, how-
ever Hydrophis cyanocinctus is much thinner and longer than Hydrophis lapemoides (Egan
2007; Garner 2013; Buzás et al 2018)
Natural History. This species is commonly found near to shore (0-9.7km out to sea)
during the day and at dusk in the Arabian Sea, and is one of the most abundant Seasnakes in
the region. This species feeds on a wide variety of fish including flat fish (Perciformes), car-
dinalfish (Apogonidae), gobies (Gobiidae); snake eels (Ophichthidae) and other unidentified

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 13. Seasnakes

eels (Anguilliformes) wrasses (Labridae), goat fish (mullidae), Pseudochromis (Pseudochro-

midae), worm eels (Ophichthidae). Eels, flatfish, and gobies are presumed to comprise the
bulk of their diet due to ease of capture. This species actively forages for food by day and
forages on the sea floor. One was observed outside of a go-
by’s burrow. Mating takes place in May, with the birthing of
1-5 offspring occurring in November. Like many seasnake
species, it is not uncommon to find this species with heavy
barnacle infestations from the sea snake specializing barna-
cle species Platylepas ophiophila. This species is considered
generally placid, however some reports claim they can be
Figure 13.71. The distribution of aggressive when removed from the water. The venom of
Hydrophis lapemoides. this species contains postsynaptic neurotoxins and possibly
myotoxins and is considered relatively dangerous to hu-
mans. This species is wide ranging and relatively abundant and listed least concern by the
IUCN (Egan 2007; Buzás et al 2018; Voris & Voris, 1983; Rasmussen et al 2010i).

Small-headed Seasnake
Hydrophis macdowelli (Kharin, 1983)
Distribution and Habitat. Northern Australia (including Northern Territory, Queensland,
Western Australia), Loyalty Islands of New Caledonia, Papua New Guinea (Uetz and Hosek
2020) Type locality: Northern Australian shelf (Uetz and Hosek
2020)
Identification. As implied in the common name, this
snakes head is much smaller in comparison to its body. Its
body is slender anteriorly and compressed posteriorly. The
base coloration of the body is a beige to cream color becoming
Figure 13.73. The distribution more of a lighter yellow to white color ventrally, with a series of
of Hydrophis mcdowelli. 30-40 black or dark grey blotches running along the top of the
body, which extend one third of the way
down the body. The side of the body also
has three longitude series of small grey
spots with occasional black outlining.
Body scales are imbricate having a medi-
an keel, with 35-42 rows at mid-body and
252-274 ventral scales with a double keel,
one keel on each side of the median line.
This species is sexually divergent, with
both the body and ventral keels being
more obvious in male specimens. The
average length is around 80 cm. (Ineich
and Laboute 2002; Cogger 2018; Storr et
al 1986).
Natural History. Australian conti-
Figure 13.74. Hydrophis mcdowelli
Photography by Hugues Debeyser.

264
Aquatic Snakes

nental shelf in water up to 50 m deep, in river estuaries and other turbid (muddy) inshore
waters. More specifically, Small-headed Seasnakes produce around two or three young in a
litter. Gestation lasts six to seven months and young are born in September. Females appear
to reproduce every year. (Cogger 2018; Fry et al. 2001).

Olive-headed Seasnake
Hydrophis major (Shaw, 1802)
Distribution and Habitat. This species is known from Indonesia in the Arafura sea,
Northern Australia, New Caledonia, and southern Papua New Guinea. Exact habitat pref-
erences are poorly known, however it is associated with turbid waters, offshore coasts with
soft bottoms including mud and sand, and can also be found
in some tidal creeks. In New Caledonia it is thought to be
associated with coral flagstones and areas with an abundance
of seaweed species including Sargassum, Halimeda and Caul-
erpa algae. It is often found by trawlers in deeper waters with
depths ranging from 1.4-106 m (Ineich & Laboute 2002;
Figure 13.75. The distribution
Cogger 2018; Guinea et al 2010c; Courtney et al. 2010). Type
of Hydrophis majori. locality: Indian Ocean per Cogger.
Identification. The head
of this species is bulky colored
dark brown with black flecks,
the body is a yellowish to beige
color with 25-30 dark blotch-
es or saddles dorsally, that do
not form complete bands ven-
trally and decrease in size to-
wards the tail. The inner spac-
es between these saddles have
small bars about a scale wide.
Just below each dorsal saddles,
are two rows of smaller round
lateral blotches or spots. The
ventral scales are barely wider
than the adjacent scales and are
dark grey with darker tips. The
head shields are unfragmented,
large, and symmetrical and this
species usually lacks an anteri-
or chin shield or one is barely
Figure 13.76 Hydrophis major, From Anse Vata, Nouméa, New present and the rostral scale is
Caledonia Photography by Josh Jensen.
barely noticeable from above.
Body scales are imbricate with a
single low blunted keel and number 37-43 at midbody. Ventral scales have a double keel and
number 230-266 with an enlarged cloacal scale. This species is sexually divergent, with both
the body and ventral keels being more obvious in male specimens. Average total length for
this species is 1.3 meters. This species superficially resembles Hydrophis macdowelli, howev-
er its head is much notably larger and wider and there are only two rows of smaller lateral

265
 13. Seasnakes

spots in Hydrophis major compared to three in Hydrophis macdowelli (Ineich & Laboute 2002;
Cogger 2018).
Natural History. Despite it’s abundance in portions of its range, there is still a lot un-
known about this species. Stomach content studies revealed prey to include pearlfish (Car-
apidae sp.), and eeltail catfish (Plotosidae sp.). Ineich and Laboute also observed a juvenile
foraging on cardinalfish (Apogon sp.) and Gon’s cardinalfish (Archamia) in pinnacle reefs
and gulf weeds at depths of 6-15 meters. Reproduction occurs year-round, though slowing in
the colder months, with 6-12 live young being produced. This species should be approached
with caution as it is considered aggressive compared to many other sea snakes, and human
fatalities have been recorded (Ineich & Laboute 2002; Voris and Voris 1983).

Bombay Seasnake
Hydrophis mamillaris (Daudin, 1803)
Distribution and Habitat. Known from Pakistan, India and Sri Lanka waters. (Heatwole
1999) It is known to occur over soft bottom sand or mud
(Lobo and Rassmussen 2010). Type Locality. Visakhapat-
nam, Andhra Pradesh, India.
Identification. Alternatively known as the broad
banded sea snake, due to the species 44-55 black body
bands which are approximately twice as wide as the in-
terspace between then which is a yellow or grayish color.
Figure 13.77. The distribution of The bands expand slightly dorsally, and are connected at
Hydrophis mamillaris. ventrally. The head is entirely black with occasional yel-
low markings around the temporal region. Body scales
are hexagonal in shape, with a tubercules and are feebly
imbricate. Dorsal neck scale count is 25-29 scale rows, and 35-43 at midbody. Ventrals are
distinct and bicarrinate with a scale count of 302-309.
Natural History. There are only 13 reported sightings of this species since it’s description,
the most recent being in 1926 by Malcolm Smith (Smith, 1926; Lobo and Rassmussen 2010).
With the type specimen lost, there has been some debate about the validity of this taxon, con-
sidered to be a synonym of Hydrophis fascaitus however Rassmussen and Lobo believe it to
be a legitimate species. It is presumed this species travels large distances in surface and tidal
currents, like Hydrophis platurus (Lobo and Rassmussen 2010; Somaweera,2009).

Slender-necked Seasnake
Hydrophis melanocephalus (Gray, 1849)
Distribution and Habitat. While the type specimen is labeled Indian Ocean, this species
seems to be restricted to northeast Asia including China, Japan, Taiwan, Korea, Philippines,
and Vietnam. It is potentially the most abundant sea snake in Japan’s Ryukyu Archipela-
go. This species is associated with inshore waters. (Rasmussen 2010c; Goris & Maeda 2004;
Masunaga et al 2005). Type locality: Indian Ocean.
Identification. As the Latin name implies, this species has a very black head, and slender
neck, both of which are black with faint yellow spotting. The body of this species is slender,
with the forebody about 1/3 the width of the posterior body. The base color of this species is a
light yellow with 46-61 dark bands running the length of the body, and 3-8 more dark bands
on the tail, which terminates in black. Bands taper ventrally, but usually completely wrapping
266
Aquatic Snakes

the body and ventrals. The eyes are notably large, larger than the
preocular scale. The frontal scale is longer than it is wide, the pa-
rietals are as long as both the prefrontal and frontal scales, and
no loreal scale is present. There are nine supralabial scales, with
scales 3-4 or 3-5 contacting the orbit, one preocular scale, 1-2
post oculars, and typically one temporal scale, but in rare instanc-
Figure 13.79. The distribu- es 7 or 8 are present. Body scales are usually keeled, are imbri-
tion of Hydrophis melano- cate anteriorly, but only subimbricate posterity. There are 25-31
cephalus.
scales at the neck, 31-39 at midbody. Ventral scales are larger than
the adjacent scales, have two reduced keels, and are sometimes
wedged making scale counts difficult. Ventral scale count ranges
from 281-353. The average length males of this species is 114 -123 cm in females. (Mao and
Chen 1980; Maunaga et al 2005; Goris and Maeda 2004).
Natural History. Very little is known about this species. Prey include conger eels (Cong-
ridae), worm eels (Moringuidae), and snake eels (Ophichthidae). Goris and Maeda reported
this species forages by putting its head in the sand and then holds onto the prey items pulling
them from the sand. Females produce 1-8 young, with birthing season varying greatly with
locality. The venom of this species is medically significant and has an LD50 of 0.111 mg/kg
subcutaneously, with an average yield of 2.1 mg. Due to an overall lack of knowledge on the
species, it is listed as Data Deficient by the IUCN. (Rasmussen 2010c; Voris and Voris 1983;
Goris & Maeda 2004; Minton 1983).

Robust Seasnake
Hydrophis melanosoma (Günther, 1864)
Distribution and Habitat. This species occurs along the east coast of the Malay Penin-
sula including Malaysia, Indonesia and Singapore. It also occurs
along the north coast of Borneo where it is considered rare,
along Indonesian Borneo, Malaysian Borneo and Brunei and
nearby islands. There is also a disjunct population in Halma-
hera, Indonesia. It is associated with shallow coastal waters as
well as rivers (Sanders 2010b; Stuebing and Inner 1999). Type
locality: unknown.
Figure 13.80. The distribu- Identification. The body of this species is robust as the
tion of Hydrophis melanoso- common name suggests, grey dorsally that graduates to an off
ma. white to yellowish ventrally with 50-70 black cross bands that
are complete, and about twice as wide as the interspaces. The
neck and head are black with yellow or cream-colored oval
shaped pattern that runs over the nasals, prefrontals, preoculars, post ocular and temporal.
The head is distinct. The ventral is a cream or yellow color, with two tubercles on each of the
266-368 ventral scales which are slightly less than twice the size of the adjacent scales. The
cloacal scale is divided and the end of the tail is black. The dorsal scales are strongly keeled
and there are 35-45 scale rows at midbody. Average total length is 1.5 meters. This species can
be distinguished from Hydrophis caerulescens and Hydrophis fasciatus, but both of those spe-
cies have small heads. Hydrophis klossi also appears similar, however the bands are the same
width as the interspaces, where as in Hydrophis melanosoma the bands are notably wider than
the interspaces (Stuebing and Inner 1999; Chan-ard et al 2015; Das 2010).
Natural History. This species forages shallow waters for a variety of prey including

267
 13. Seasnakes

conger eels (Congridae),

worm eels (Moringuidae),
moray eels (Muraenidae)
and various other eels
(Anguilliformes). The re-
productive habits of this
species are yet unstudied.
Due to a lack of population
information, this species is
listed as Data Deficient by
the IUCN (Voris and Voris
1983; Das 2010; Sanders
2010b).

Black-banded
Seasnake
Hydrophis nigrocinctus
(Daudin, 1803)
Distribution and
Habitat. This species oc-
curs in the Indian Ocean
in Bangladesh, India, and
Myanmar. Little is known
about this species includ-
ing habitat preferences or
ecological niche. The type
specimen was found near
the tidal zone and lower
saline areas near a river
in Calcutta (Rasmussen &
Lobo 2010d; Rasmussen
et al 2011c). Type locality:
Figure 13.81. Hydrophis melanosoma. This image is from Boulenger
Sundarbans, Bengal (Levi-
1898, and is labled Hydrophis floweri. This species was later placed in ton et al 2003).
the synonomy of H. melanosoma. Identification. Hy-
drophis nigrocinctus is a
banded sea snake species
easily distinguished from all the Hydrophis in Asia by only having 1-3 maxillary teeth behind
the fang. The body is an olive color dorsally graduating to a yellowish on the flanks ventrally.
The body of this species is covered in 40-60 thin dark body bands, slightly tapering ventrally
on the flanks. The head is black and has a yellow stripe at the supraorbital with a crown pat-
tern that extends to the prefrontals. The mental scale is large and not concealed by the mental
grove. There are 27-33 scale rows at the neck and 39-45 scale rows at midbody. Ventrals are
distinct, but not twice as wide as the adjacent scales, and number 296-330. Average total

268
Aquatic Snakes

length is 1080 mm (Leviton et al 2003; Rasmussen et al 2011c).

Natural History. Very little is known about this species. It
was formerly in the genus Disteira, and members of that for-
mer genus are diurnal and prefer deep, turbid waters with sandy
[Link] prey include conger eels (Congridae). Noth-
ing is known about this species reproductive cycle. With very
little known about this species, its population size, and ecologi-
Figure 13.82. The distribu- cal preferences, this species is listed data deficient by the IUCN.
tion of Hydrophis nigrocintus.
Rassmusen et al synonymized Hydrophis walli with Hydrophis
nigrocinctus and thus Hydrophis walli is not included in this
volume. Wallach et al synonymized Hydrophis hendersoni with
Hydrophis nigrocinctus and thus Hydrophis hendersoni is also not included in this volume
(Voris and Voris 1983; Rasmussen et al 2011c; Rasmussen & Lobo 2010d; Wallach et al 2014).

Russell’s Seasnake
Hydrophis obscurus (Daudin, 1803)

Figure 13.83. Hydrophis nigrocinctus

The image is from Abhandlungen aus dem Gebiete der Naturwissenschaften 1846.

Distribution and Habitat. Hydrophis obscurus is known from east India, Myanmar, Ban-
gladesh, and southern Thailand. It is associated with river mouths, saltwater lakes, and shal-
low brackish lagoons (Rasmussen & Lobo 2010e; Leviton et al 2003; Das 2010). Type locality:
Sundarbans, India (Leviton et al 2003).
Identification. This species goes through a significant ontogenetic change, where juve-
niles are black to bluish grey with 35-55 complete bright yellow to white bands, but adults pat-
terns fade to become a completely uniform grey dorsally and yellow ventrally. Juveniles heads
are also pattered, with a black base patterned with a curved yellow marking that runs from
the snout to the sides of the parietal scales. The body scales are imbricate and have rounded
or blunted scale points. There are 19-23 scale rows at the neck and 29-37 at midbody. The

269
 13. Seasnakes

ventrals are distinct and number 300-338. Average total length

for this species is 1190 mm in males, and 1200 mm in females,
with only a slight size variance between the sexes (Leviton et al
2003; Das 2010).
Natural History. This species it is though to inhabit brack-
ish areas including lagoons and river mouths where it search-
Figure 13.84. The distribuition es for prey which are known to include tripodfish (Triacan-
of Hydrophis obscurus. thidae). Little else is known about this species. (Leviton et al
2003; Das 2010; Voris and Voris 1983)

Ornate Reef Seasnake

Hydrophis ornatus (Gray, 1842)
Distribution and Habitat. A very widespread species found west in the Persian gulf, and
the Indian Ocean east to South East Asia including Indonesia, Malaysia, Vietnam, Thailand,
Phillipines, and Myanmar, North to southern China and Japan, and south to Australia, New
Caledonia, and the Solomon Islands. This species prefers waters 30 m or deeper in depth,
although it has also been found at coral reefs and mildly turbid
inshore water as well as estuaries (Uetz and Hosek 2020; Ras-
mussen et al, 2010h). Type locality: None given.
Identification. The head color is a dark olive to black
above, and off white to yellow below, with a fragmented lighter
hotshot pattern and is large and robust. The base dorsal color-
ation is variable and can include light gray, light olive or off-
Figure 13.85. The distribution white with wide black, grey or dark brown bands that come to
of Hydrophis ornatus.
a point just above the ventral scales. In males 40-55 bands are
present while females have 40-56 bands. Scales are imbricate
or juxtaposed, and hexagonal in shape at midbody. Midbody scale count is 33-45 on males,

Figure 13.86. Hydrophis ornatus. from Ponton Méridien, Province Sud, New Caledonia.
Photography by Pauline Fey.

and 39-55 in females, with males having 209-260 ventral scales and 236-312 ventral scales in
females. Snout to vent length in males is 950 mm with a tail length of 11 mm Snout length is
860 mm in females, with a tail length of 80mm (Egan 2007; Rasmussen, 1989; Leviton et al,

270
Aquatic Snakes

2003).
Natural History. The diet of this species has been shown to vary regionally and includes
a variety of fish species that occur close to coral reefs. In Australia Hydrophis ornatus appears
to specialize in benthic and demersal fish including cardinal fish (Apogonidae), cusk eels
(Sirembo), gobies (Yongeichthys), bigeyes (Priacanthidae), goatfish (Mullidae), and threadfin
breams (Nemipteridae). In Thailand six fish families have been found in stomach content
studies including cardinal fish (Apogonidae), squirrel fish (Holocentridae), wrasse (Labri-
dae), ponyfish (Leiognathidae), parrot fish (Scaridae), and threadfin breams (Nemipteridae).
Because this species diet is known to include free swimming species, there is speculation they
may prey on sleeping fish at night. Sexual maturity is reached around 800 mm in females with
litter sizes ranging from 1-17. This species’ venom contains potent myotoxins and has caused
human fatalities (Voris and Voris 1983; Fry et al, 2001; Gardner 2013; Rasmussen, 1989).

Ocellated Sea Snake

Hydrophis ocellatus Gray 1849
Distribution and Habitat. The Northern Territory, Queensland, Western Australia, and
Tasmania.[No map.]
Identification. Long confused with H. ornatus, however Rasmussen et al. (2014) found
no evidence that H. ornatus occurs in Australia waters. (no map) but found it in New Cale-
donia. (see figure 13.87).

Figure 13.87. Hydrophis ocellatus

from Cable Beach WA, Australia
Photography by Sharmaine Don-
nelly.

Fischer’s Sea Snake

Hydrophis pachyceros Fisher 1855
Distribution and Habitat. Indian Ocean, South Chinese Sea, and Vietnam. Neotype was
obtained from a fish market at Phat Thien, Vietnam the specimen was taken from a trawler
catch by Rasmussen et al. (2007).
Identification. Diagnostic characters: Scale rows around neck 26–31; scale rows around
body 39–45; ventrals 247–297. Maxillary teeth behind fangs 7–8. Coloration: Body pale yel-
low above, white below, with pale brown transverse bands fading on the upper part of the
flanks. Head white below, black/dark above with pale supralabials and a pale ring around the
eyes.

Small-headed Sea Snake

Hydrophis parviceps Smith 1935
Distribution and Habitat. South Chinese Sea -Vietnam. Type locality: coast of Cochin

271
 13. Seasnakes

China (no map).

Identification. Head small, body slender anteriorly. Scale rows around neck 19–21;
scale rows around body 31–34; ventrals 329–348. Maxillary teeth behind fangs 6–. Color-
ation: Body olivaceous above, greyish below, with 65–0 blackish bands. Head black with no
or only a few faint paler marks.

Pacific Seasnake
Hydrophis pacificus (Boulenger, 1896)
Distribution and Habitat. This species is known from southern Papua New Guinea and
Northern Territory and Queensland in Australia. It is associated with continental shelf wa-
ters, inter-reef waters, as well as soft sediment estuaries and tidal rivers, with a known from
depths ranging from 0-50 meters. Most of the records for
this species come from shore wash-ups, trawler by-catch,
or surface captured animals basking in shallow continental
shelf waters (Milton et al 2010c; Cogger, 2013). Type local-
ity: New Britain, Papua New Guinea.
Identification. Hydrophis pacificus, which is also
known as the large-headed sea
Figure 13.88. The distribution
snake, has a large-head com-
of Hydrophis pacificus.
pared to other members of the
genus, with a relatively slender
body that is notably compressed laterally. Juveniles present 49-
72 pronounced black bands against a lighter foreground that
are complete and from the black head to the tail. The throat
in juveniles is also black with light-yellow spots behind each
eye and pale spots on the snout. Pattern becomes less dis-
tinct in adult specimens, with a darker grey dorsal side with a
mid-body lightening ventrally. Bands in adults may be faded,
the bands on the neck are usually, but not always connected
by black ventrals. Adult bands maybe divided or displaces on
the vertebral line. Thread on adults is park grey above, lighter
beneath, and has pale flecks. Head shields in this species are
distinct and regular. Body scales are imbricate and number 45-
49 rows at midbody. Ventral scales are mostly undivided, not
much broader than adjacent scales, and number 320-430. Av-
erage adult length is 1.4 m. (Wilson and Swan, 2013; Cogger,
2013)
Natural History. This fish specialist is slow to mature and
reproduce, with the approximate age of maturation occurring
at eight years, yet a total longevity estimate of only twelve years.
Litter size is reported to be 17 live young. Due to this species Figure 13.89. Hydrophis
slow reproductive rate, and the pressures from being captured pacificus. From Boulenger
as by catch from trawling, it is listed as vulnerable by the IUCN 1896.
and is thought to be in decline. While generally considered
non-aggressive this species may bite when restrained and does
posses postsynaptic neurotoxins which are considered poten-

272
Aquatic Snakes

tially lethal to humans. (Milton et al 2010c; Wilson and Swan, 2013; Tangella, 2019)

Horned Seasnake
Hydrophis peronii (Duméril, 1853)
Distribution and Habitat. This species is known from northern Australia, southern Pap-
ua New Guinea, the Solomon Islands, the Loyalty Islands, southern Indonesia, and New Ca-
ledonia in the southern hemisphere. The northern and southern hemisphere populations are
notably disjunct, with no known occurrence in the Philippines or Central Indonesia. This
species distribution picks up again in the northern hemisphere
where it is known from Cambodia, China, Malaysia, Singapore,
Taiwan, Thailand, and Vietnam. Hydrophis peronii is associated
with soft sandy bottoms adjacent to coral reef and is found in
depths from 0-60 m (Lukoschek et al 2010). Type locality: New
Holland (Modern day Australia)
Figure 13.90. The distribu-
Identification. This species is easily distinguished from all
tion of Hydrophis peroni. other Hydrophis by having pronounced spines on its head. The
head is short, and small, with very broad shallow nostrils, and
distinct eye color of off-white, with a black ring encircling a
slightly darker shade of white with black pupils. The body color is off-white to gray, lighter
below than above with 25-30 brown or black bands which taper ventrally, which can become
obscure in older specimens. On some specimens, there is a second set of smaller incomplete

Figure 13.91. Hydrophis peroni. From Jan and Sordelli 1872.

273
 13. Seasnakes

dark bands, between the primary bands. This pattern may be obscured by biofouling, with
reports of some specimens found covered in algae and seaweed. The supraocular, postoc-
ular, and adjacent scales are raised and pointed along the posterior border with projecting
spines in adults, that present as still present, yet less distinct tubercles in juveniles. Parietal
and frontal shields are indistinct, fragmented into smaller scales, and the prefrontal scales
are absent. This species’ neck is slender, and only one-third to two-thirds the width of the
diameter of the body. The body scales of this species have a strong dorsally facing spine
shape, with short keels, more distinct in males than females. This banding becomes faded
and less distinct on the tail. Ventrally narrow bands or spots may be present. Dorsal scale
count is 18-24 scale rows at the neck, 21 to 31 rows at the mid-body, ventral scale count is
140-210, subcaudal scales count is 29 and 35, and the cloacal scale divided. The average
adult total length is 1.1-1.3 meters. (Cogger 2014; O’Shae, 2005; Leviton et al 2014; Fry et
al 2001)
Natural History. Hydrophis peronii is reported to go through an ontogenetic prey shift,
with juveniles specializing in shrimp and adults hunting the ocean floor looking for bur-
rowing gobies (Gobiidae) in the genus Trypauchenidae and Oxyurichythys. This species
appears to be active both day and night, though Retfield et al. had greater success collecting
specimens on trawl by day than night. Hunting consists of prodding their smaller head and
narrow necks into the burrows of gobies, which are comprised of mud, sand, gravel, and
broken coral. The prey is quickly subdued with the species’ potent post-synaptic neurotoxic
venom, with an LD50 of 0.125 ug/g when tested intravenously in mice, and a total yield of
18.3 mg. While incredibly toxic, the snake has a small yield and no human bites or fatalities
have been recorded. This species appears to reproduce annually giving birth to up to 10
live young. In Australia gestation lasts six to seven months with birthing in March to June.
(Cogger, 2014; Voris, 1972; Voris and Voris 1983; Fry et al. 2001; Mori and Tu, 1988; Leviton
et al 2014)

Yellow Bellied Seasnake

Hydrophis platurus (Linnaeus, 1766)
Distribution and Habitat. Hydrophis platurus possess the largest natural range of any
living squamate (Rasmussen et al 2011). In the New World, in the Pacific Ocean, they have
been spotted as far south as Coastal Chile (Donoso, 1966), and as far north as Ventura
County, California (Nafis, 2020). It should be noted, that though this species can occur in
these far limits of its presumed “extended” range, due to the species low tolerance of cooler
water temperatures (Graham, 1974). These occurrences are often attributed to accidental
current drift, and likely do not represent known breeding populations. Type locality Un-
known.
H. p. platurus populations with year-round breeding are likely limited from Mexico to
Ecuador, where occurrence of the species is far greater. There have also been four sightings
of the species in the Caribbean Sea, off the coast of Colombia, though these sightings are
presumed to be due to human introduction, be it via the Panama Canal or other means
(Hernández-Camacho et al. 2006). In the Old World this species ranges throughout the
greater part of Southeast Asia and Australia, south into New Zealand and Tasmania and as
far North as Southern Japan. Hydrophis platurus also inhabits all the Indian Ocean includ-
ing the entire East African coast, the adjacent Persian Gulf and the Red Sea. Their range
in the Indian Ocean occurs all the way south to South Africa, where their occurrence is
considered rare. They have on two occasions been spotted across into the Atlantic Ocean

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in South Africa and Namibia (Branch, 1998), though this occur-

rence is considered likely an accidental current displacement and
does not represent a breeding population (Visser, 1967). Should
climate change continue to warm these waters, it is the authors
opinion that this species could see significant range expansion, as
cold waters appear to be their main barrier to further dispersal.
Figure 13.92. The distribu- As the alternate common name, the pelagic sea snake implies,
tion of Hydrophis platurus. this species is entirely pelagic. They are most often seen off trop-
ical shores, where rain is plentiful and ocean temperatures stay
between 36-18 C. While the species is capable of surviving tem-
peratures below 16 C, feeding has been documented to cease from 18-16 C, and thus areas
in this range are not common places for them to occur (Dunson and Ehlert, 1971; Graham,
1974). Though the species does inhabit the open ocean, as its movements are largely dictated
by currents, they are mostly found within 1-20 km from shore throughout their range. Due to
their “drifting” lifestyle, they can most easily be located in “debris slicks” where currents con-
verge along shorelines. These slicks can be identified by the large number of debris, and an
obvious flatness. Some have described them as looking like “small roads” on the surface of the
ocean, a description the author finds fitting for the general width and appearance. (Per obs)
Identification. Hydrophis platurus is one of the most distinct looking of all the snakes in
the subfamily Hydrophiinae. It’s laterally compressed body, side by side semi-hexagonal to
quadrangle shaped scales, flattened head, and distinctive bicolor (though variable) patterning
all contribute to the relative ease of this species identification. The common pattern is a black
colored dorsal side, yellow colored ventral side, down to the tail, which is an aberrantly spot-
ted and highly variable mix of white and black, though on some individuals the white is not
present, and instead the yellow coloration front he ventral side continues to the tail. Juveniles’
tails may be yellow instead of white, and tail coloration could be an ontogenetic change. The
bicolor arrangement is most common, though colors may vary including brown and yellow
bi-color, black with a lateral strip and yellow, aberrant, and fully melanistic. The recently de-
scribed subspecies Hydrophis platurus xanthos lacks the black and is instead solid yellow with
occasional black dots. Scales are irregular in shape varying from sub-quadrangular to hexag-
onal. These scales become very small along the body with 49–67 rows around thickest part
of body and a ventral scales count of 264–406. Their head is narrow, with an elongated snout
and large head shields. nasal shields in contact with one another; prefrontal in contact with
second upper labial; 1–2 pre- and 2–3 postoculars; 2–3 small anterior temporals; 7–8 upper
labials, 4–5 below eye but separated from border by subocular. Species snout to vent length
is dimorphic, Female reaching 880 mm and males reaching 720 mm (Leviton et al. 2014).
Natural History. Hydrophis platurus is a truly pelagic species, occurring off tropical
shores around the world. Like other Hydrophiinae, the species is reliant on rainwater for
fresh drinking water, and as such is most densely populated in areas with heavy rain fall.
Some unique characteristics of the species include their “knotting behavior” employed for
ecdysis, and their ability to swim backwards or forwards with equal ease. While the species
can dive, they are mostly encountered floating on the surface of the water. The species is pri-
marily diurnal, and most encountered in the morning hours. Upon approached carefully, the
snakes rarely dive, unless there is a significant water disturbance. While adept swimmers in
the water, this species is almost incapable of locomotion on land due to their complete lack of
ventral scales. The species feeding behavior is quite unique. The bicolor camouflage under-
neath looks like the leaf and debris they hide in while in debris slicks. As fish approach the
snake, they prey upon them using a lateral striking method, and often consume them without

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 13. Seasnakes

Figure 13.93. A Hydrophis platurus from Birubi Beach, Newcastle, New South Wales, Australia
Photography by Scott Eipper.

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Aquatic Snakes

the aid of venom. Heatwole proposed the snake may use small sensory organs on the scales
to feel the presence of prey items, similar to the predation tactics employed by acrochordids.
They can often be found among washed out leaves and sticks in these slicks. As small fish ap-
proach the debris in search for food, they become prey items for the Hydrophis platurus. Hy-
drophis platurus breed year-round, with gestation lasting about six months, giving live birth
to 1–10 offspring. There is some speculation that the offspring may stay with their mother
for the first few days of life, which would be a unique maternal behavior among sea snakes.
While human deaths are rare, the venom of Hydrophis platurus is highly potent nuerotoxin
(with other unique isotoxins) with a subcutaneous LD50 0.055 mg/kg, making it the seventh
most toxic snakes on earth when tested subcutaneously. This said the species is hesitant to
bite in the water, and the species average yield is only. (Bolaños et al 1975; Pickwell et al 1972;
Steinhoff, 2018). While a lethal bite is possible, it is unlikely due to the species hesitancy to
bite unless restrained. However, in the author’s experience this species will in fact attempt to
bite if restrained, and as such handling by an untrained professional should be avoided unless
necessary.
Taxonomy. In 2017, Brooke L. Bessesen, and Gary J. Galbreath described the Golfo Dul-
cean Sea Snake (Hydrophis platurus xanthos), as the first subspecies in the long-standing
monotypic species. They described the subspecies as having a notably smaller body, and
nearly solid yellow pattern. They also noted that this subspecies is most active at night, and
not associated with drift-lines as Hydrophis platurus platurus is. This subspecies is endemic
to the Golfo Dulce in Southern Costa Rica (Bessesen, Galbreath 2017).

Beaked Seasnake
Hydrophis schistosus (Daudin, 1803)
Distribution and Habitat. A very widespread species found as far west as the east coast of
India, east to Southeast Asia, north to southern China and south to northern Australia. The
species also appears to be a habitat generalist occurring in deep sea locations with rocky or
sandy bottoms, tidal creeks, shallow open sea, river mouths,
lagoons mangrove forests and estuaries. It has also been
found in freshwater lakes in Cambodia and India, as far up as
7l km in the Goa River in India (Rassmussen et al 2018). The
species prefers depths between 5 m and 30 m (Rassmussen
et al 2018). Animals are active night and day and night, and
most seen in marine environments with soft bottoms. This is
Figure 13.94. The distributuion
the most encountered sea snake by humans (Karithkeyan and
of Hydrophis schistosus.
Balasubramanian 2007). Type locality: Tranquebar, India.
Identification. Hydrophis schistosus is distinct from most
other sea snakes other than based on its beak shaped rostral. It shares this shape with Hy-
drophis zweifeli, which it can be distinguished from based on its 1-2 preoculars which are
lacking in Hydrophis zweifeli, and instead are fused with the supraoculars. Juveniles are dark-
er on top and lighter below, with bands that fade with age to a uniform dark dorsal gradient to
a lighter ventral side. Dorsal scales are somewhat imbricate or juxtaposed with a short central
keel. Midbody scale count in males is 43-52 and in females 48-55. Ventral count is 262-322,
with a maximum total length of 1400 mm, though average adults rarely surpass 1100 mm
total length (Rezaie-Atagholipour et al 2016; Ukuwela et al 2013).
Natural History. The beaked sea snake is active day and night, and forages for prey in-
cluding crustaceans and fish, larger specimens feeding mainly on fish, with an apparent pref-

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 13. Seasnakes

Figure 13.95. Hydrophis schistosus from Sri Lanka.

Photography by (a) Paul Freed (B-C) by Anslem da Silva.

278
Aquatic Snakes

erence for catfish. Voris and Voris found a large variety of prey items in their 1983 stomach
content study including eels (Anguilliformes), catfish (Ariidae), anchovy (Engraulidae),
lizard fish (Harpadontidae), ponyfish (Leiognathidae) eeltail catfish (Plotosidae), drums
(Sciaenidae), and even the toxic pufferfish (Tetraodontidae), and shrimp. Mating occurs
March to April. Males combating other males for mates, such as seen in some terrestrial
species, has been observed in this species. Female mature rapidly at 18 months, producing
first offspring at 24 months. Offspring are large 21-24 cm and numerous (up to 30 or more
in larger females), though only 6% of females survive to a reproductive age. Ernst and Zug’s
book, Snakes in Question: The Smithsonian Answer Book errantly declared Hydrophis
schistosus as the most toxic snake in the world, and with an LD50 of 0.1125 mg/kg SC, it is
certainly one of the more lethal snake species, but not the most toxic by any route of injec-
tion. While human fatalities are rare, they have occurred, and the species should be treated
with caution. (Voris and Voris, 1983; Karithkeyan and Balasubramanian, 2007; Rassmussen
et al 2018; Rezaie-Atagholipour et al. 2016; Fry 2012)

Lake Taal Seasnake

Hydrophis semperi Garman, 1881
Distribution and Habitat. Lake Taal, island of Luzon, Philippines is freshwater lake
and the type locality for this sea snake. The lake is a deep meso-eu¬trophic lake in a vol-
canic caldera. Taal Lake was once a navigable inlet to Balayan Bay. A series of eruptions
in the early 18th century culminated in 1754 with the larg-
est eruption that blocked the Pansipit River, blocking the
lake’s outlet to the sea. This caused the waters to rise, Since
the 1754 eruption, the surface elevation had risen from sea
level to 5 m above sea level. Lake Taal became freshwater
after centuries of precipitation. Taal Lake was declared a
Figure 13.96. The distribution of national park, Taal Volcano National Park, in 1967, it cov-
Hydrophis semperii.
ers 62,292 hectares. When Garman (1881) described this
snake, he considered it similar to Hydrophis fasciata and
H. fisheri (presumably Platurus fischeri which is now in the
synonymy of Laticauda laticauda). However, Voris (1977)
considered it similar to Hydrophis cyanocinctus that is not
present in the waters surrounding Luzon.
Other endemic species have evolved and adapted to
the desalination of the lake’s waters. The lake has a freshwa-
ter-adapted population of the fishes, Caranx ignobilis and
Sardinella tawilis, Two other endemic fish are the gobies
Figure 13.97. The Lake Taal
Gnatholepis volcanus and Rhinogobius flavoventris. The
Seasnake, Hydrophis semperi. Bull Shark, Carcharhinus leucas, was part of the lake’s eco-
Photography by Vhon Oliver S. system but were extirpated by the local people by the 1930s.
Garcia. Identification. Head slightly larger than the neck,
crown convex, snout broad, rounded ; tail about one eighth
of the total length, moderately broad. Body not compressed
anteriorly, greatest diameter about twice that of the neck. Maxillary teeth behind fangs six
to eight; scales on body imbricate and with short keel, 29–21 around neck, 37–43 around
body (increase of 10–12); ventrals 314–356, distinct throughout, bicarinate, about twice as

279
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broad as adjacent scales; precloacal scales enlarged; head black, body black with 50 to 60
narrow whitish bands of annuli, ventral scales usually black (Garman 1881; Leviton et al.
2014).
Natural History. Analysis of gut contents found gobies and eels are primary prey items.
Halfbeaks (Family Hemiramphidae) were recorded as one of the Lake Taal Sea Snake’s prey
which is considered a new prey record for sea snakes (Garcia et al. 2014).
Garcia et al. (2017) reported a distinct surfacing behavior. As the snake approaches the
surface to ventilate its lungs it arch’s its body. The motion begins with a nearly vertical,
ascent to the sur-face. The sea snake appeared to drag its body with its head as it ascended,
with the momentum causing its head and neck to break the water surface. Its head briefly
remained above the surface (1–2 s). It then formed an arch by diving back into the water
as it swam towards the bottom, as opposed to simply descending in reverse motion. This
type of biomechanics is like other sea snake species that ascend to the water surface to re-
spire. The authors recorded the behavior 2–3 times day and night. They also observed the
behavior of captive H. semperi was limited to moving from crevice to crevice and surfacing
to breathe.
Threats to the species come from aquaculture and fishing. While they may avoid aqua-
culture by staying in open water their surfacing behavior makes them susceptible to gill-
nets.

Kalimantan Seasnake
Hydrophis sibauensis (Rasmussen et al., 2001)
Distribution and Habitat. This species is known from only three specimens which were
found in the freshwater river habitats of the Sibau River in Kalimantan, Indonesia, 1000 km
upriver from the coast (Rasmussen et al 2001). Type local-
ity: Sibau Kecil, Putussibau (Kalimantan, Indonesia) (Ras-
mussen et al 2001).
Identification. Hydrophis sibuaensis appears to occur
alone in the Sibua river, with no conspecifics. This species
head is black dorsally and ventrally with light yellow dots
Figure 13.98. The distribution of or lines below creating a forward-facing arrow and dots or
Hydrophis sibauensis. blotches behind the arrow. Base color of the body is dark
grey to black and dark anteriorly than posteriorly. The ventral surface is black ventrally, but
grayish yellow from midbody back. The body is patterned with 49-58 yellow bands that do
not connect ventrally. The coloration of the bands transitions with the shade of yellow being
brighter anteriorly and transitioning to a dark yellow to light orange posteriorly. The tail is
patterned in 8-9 elongated blotches. There is one postocular scale, The dorsal scales have
a long median keel and number 15-26 scale rows at the neck and 35-37 scale rows at the
midbody. This species ventral scales are twice as wide as the adjacent scales, unfurrowed,
and number 257-264. Total length is 45.5-73.5 cm (Rasmussen et al 2001).
Natural History. This species is known only from three specimens from a single iso-
lated location. All specimens were collected during the regions two rainy seasons (July to
August and November to December). As such the river was flooding, with estimated depths
of 2 to 3 meters. The holotype was gravid with seven unborn young collected in December,
indicating this species may give birth in December, however if the breeding is seasonal or
year-round remains unknown (Rasmussen et al. 2001).

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Yellow Seasnake
Hydrophis spiralis (Shaw, 1802)
Distribution and Habitat. This species is widespread occurring in the Arabian Sea in
Bahrain, Kuwait, Iran, Iraq, Oman, Saudi Arabia, United Arab Emirates, and Qatar. In south
Asia it is known from Pakistan, India (including the Andaman and Nicobar Islands), Sri
Lanka, and Bangladesh. In southeast Asia it is known from
Indonesia, Malaysia, Myanmar, Philippines, Singapore, and
Thailand. There is also a disjunct range in New Caledonia.
This species is associated with both deep-water habitats over
sand muddy bottoms up to 50 meters deep, as well as shallow
waters over coastal reefs and sandy bottoms (Gardner 2013.)
Figure 13.99. The distribution of Type locality: Indian Ocean.
Hydrophis spiralis. Identification. This species head is moderately sized,
only slightly wider than the neck and mostly indistinct from
the body. the head color is black with a yellow horseshoe
pattern in juveniles, but in adults this head pattern fades and lends the head to be typically
mostly yellow with some black markings. The long slender body has a yellow to mustard
brown base color, which fades to a paler yellow or white on the flanks. The body is patterned
in 30-60 narrow bands (normally less than 50), which are usually
complete, and thinner than the wider interspaces. The tail often has black blotching,

Figure 13.100. Hydrophis spiralis.

Photography by Anslem da Silva.

with the end tip being black. The complete narrow black bands on a long and slender yellow
body help distinguish this species from others. There are 6-8 upper labials with the 3rd and
4th or 4th and 5th contacting the eye. One preocular scale is present, as well as 1-2 postocular
scales. The dorsal scales are imbricate and smooth with a small tubercle and nearly hexagonal
in shape, though longer than they are broad. There are 29-39 dorsal scale rows at midbody,
and 282-373 dorsal scales. This is the longest of all known sea snakes, and can reach up to
2.75 m, though 1.6-1.9 meters is more common (Egan 2007; Garner 2013; Buzás et al 2018).

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Natural History. Very little is known about the life history of this species, despite its
wide distribution. This species is encountered both during the day, and at night, with some
reports associating it with deeper water, while others shallow waters. It is occasionally seen at
the surface basking when the water is calm. Known prey items include eels including conger
eels (Congridae), and snake eels (Ophichthidae), as well as small hole dwelling fish. Clutch
sizes range from 5-15, though little else is known about their reproductive cycle. There are
vastly contrasting reports as to this species disposition, some stating it is rather aggressive,
and others stating it is placid, however this species has led to human fatalities, and its venom
contains potent myotoxins. This species is widespread, and though it is found as bycatch by
trawl fisheries, is listed as Least Concern by the IUCN (Egan 2007; Garner 2013; Buzás et al
2018; Voris and Voris 1983; Rasmussen et al 2010j).

Stoke’s Seasnake
Hydrophis stokesii (Gray, 1846)
Distribution and Habitat. A widespread species occurring in UAE, Pakistan, Oman, Pa-
kistan, India, Sri Lanka, China, Taiwan, Thailand, Malaysia, Vietnam, Indonesia, New Guin-
ea, Eastern Australia, and the Phillipines. In the Strait of Malacca it is reported too have been
observed in the thousands floating along sea slicks, similar to reports of Hydrophis platurus
by Lillywhite et al. in Costa Rica. Hydrophis stokesii is a habitat
generalist, inhabiting reefs and harbours over mud and sand
substrates found in depths ranging from 0-50 m (Uetz and Ho-
sek 2020; Lillywhite et al 2015; Sanders et al 2018). Type local-
ity: Australia.
Identification. Hydrophis stokesii is the heaviest of all the
true sea snakes often reaching a weight of 2 kg, with a record
Figure 13.101. The distribu- weight of 5.144 kg. with a notably large head, stout body, and
tion of Hydrophis stokesi.
thick neck. It is also notable for having the longest fangs of any
marine snake, up to 6.7 mm, making it capable of penetrating
neoprene wetsuits. The juvenile pattern includes alternating dark blotches along the vertebral
line and narrow incomplete dark bands against a yellow-brown, to light yellow foreground.
Two series of dark spots flank the lateral surfaces alternating from the lower lateral line to
the mid-dental line. Pattern fades with age, and alternating bands can diminish to narrow
rows of spots. Adult color ranges from off-white to dark grey, obscuring the reticulate pat-
tern. Unfragmented large head shields are present and are symmetrical. One preocular and
two postoccular scales are present, 10-12 lower labial scales are present, as are 8-10 upper
labial scales of which the second and third may meet the prefrontal scales, and upper labial
scales 4-6 contact the eyes. Body scales possess a median keel or series of tubercles and are
imbricate, numbering 46-63 at mid-body. Ventral scales at the throat consist of single rows.
Posteriorly the ventral scales become broken up and are often cryptic with the remainder of
ventral scales consisting of reduced, strongly keeled, elongated pointed scales with dentate
tips which number 226-286. The notable ventral points may be absent in juveniles. This is a
large species that can reach a maximum adult length of nearly two meters. (Wilson and Swan,
2013; Sanders et al 2018; O’Shea 2005; Cogger, 2013).
Natural History. This species forages among reefs for fish including venomous juvenile
stonefish, venomous toadfish (Batrachoididae), jawfish (Opisthognathidae), gobies (Gobi-
idae), and similar fish. This species is one of the few sea snakes stereotyped with having
aggressive behavior towards humans, reportedly pursuing them with repeated bites. Coupled

282
 13. Seasnakes

Figure 13.102. Hydrophis stokesii.

From from John Lort Stokes (1846) book Discoveries in Australia, volume 1.

with a potent post-synaptic neurotoxin and fangs long enough to penetrate 5 mm wetsuits,
it should be approached with caution, though no human fatalities have been reported (Vo-
ris and Voris, 1983; Sanders et al 2018; O’Shae 2005; Williamson, 1996).

Collared Seasnake
Hydrophis stricticollis Günther, 1864
Distribution and Habitat. Hydrophis stricticollis is endemic to the Indian Ocean and
adjacent bodies of water, including India, Sri Lanka, Bangladesh, and Myanmar. Little is
known about the habitat preferences of this species. It is associated with muddy bottoms,
turbid waters, and is known to into rivers (Rasmussen &
Lobo 2010). Type locality: India (Uetz and Hosek 2020).
Identification. Hydrophis stricticollis is a long and
slender banded sea snake with a small head, with the
posterior portion of the body being 2.5-3 times thicker
than the anterior portion. This species is commonly pat-
Figure 13,103. The distribution of terned with 45-65 dark bands which are widest dorsally
Hydrophis stricticollis and narrow ventrally, but these bands may fade entirely
with age. The body color is grey to olive dorsally and yel-
lowish ventrally. Head color ranges from olive to black
with yellow markings on the head and snout. One ante-
rior temporal scale is present, and rarely divided, as are
7-8 upper labial scales of which the second one contacts
the prefrontal scales and scales 3-4 border the eye. Scales
at the thickest part of the body are hexagonal in shape,
mildly imbricate or juxtaposed, and number 45-55 at
midbody. Ventral scales
Figure 13.104. Hydrophis number 374-452, and are
striicticolis. From Günther less than twice as large as
1864). the adjacent scales though

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Aquatic Snakes

they are distinct down the length of the body. Total length in both males and females is 1050
mm, of which 140 mm is tail in males and 90 mm is tail in females (Leviton et al 2003).
Natural History. Little is known about the natural history of this species. They are thought
to produce a viviparous brood of ten or fewer offspring measuring approximately 350 mm at
birth. Specific prey is unknown, but it is suspected to hunt near-shore environments for fish.
(Rasmussen and Lobo 2010).

West Coast Black-headed Seasnake

Hydrophis torquatus Günther, 1864
Distribution and Habitat. This species is known from the Straits of Malacca, as well as
the Gulf of Thailand including Thailand, Vietnam, and Cambodia. It is noted as particularly
abundant at the mouths of the Mekong and Chanthaburi rivers and is unique among the ge-
nus for being one of the few known species to travel up rivers. One interesting, confirmed re-
cord of this species includes Cambodia’s inland lake, Tonle Sap, well known for its abundance
of Homalopsidae. This specimen likely traveled up the Mekong
River, where the species is notably abundant, which breaks into
the Tonle Sap and Mekong Rivers in Phnom Penh, and contin-
ued up the Tonle Sap river to arrive at the lake. There is a single
record of this species in Borneo, though this is suspected to be
a vagrant. There are also several museum specimens labeled
Figure 13.105 The distribu- as China, though the locality here is suspected to be incorrect.
tion of Hydrophis torquatus. Hydrophis torquatus is associated with shallow coastal waters,
estuaries, brackish waters, and freshwater rivers mouths, and
has been documented in freshwater rivers and freshwater lakes
(Rasmussen 2010; Ineich 1996; Das 2010). Type locality: Pen-
ang, Malaysia .
Identification. Hydrophis torquatus has an elongated body
that compresses posteriorly with a moderate size head. Col-
or and pattern typically consist of 55-68 dark grey to black
bands or saddles 3-5 scales wide which are complete across the
ventral scales anteriorly. Juvenile color and pattern are more
pronounced with 46-63 well-distinguished black bands over a
white body. The body coloration consists of a greenish-grey to
greyish-tan dorsally and a yellowish cream color ventrally. The
head pattern consists of a dark grey, dark olive, or black fore-
head with a yellowish horseshoe-shaped pattern often present
Figure 13.106. Hydrophis tor- across the snout and sides of the head. Yellow spots may also
quatus from Gunther 1864, sometimes be present on the frontal and parietal shields. A
plate 25. single preocular scale, and 1-2 postocular scales are present, as
well as 7-8 supralabial scales, of which scales 2-3 contact the
eye. Scale rows number 35-42 at midbody, and the dorsal scales
are squarish in shape at the thickest portion of the body. Ventral scale count is 242-343 with
preanal scales enlarged (Chan-ard et al 2015; Murphy et al 1999; Das 2010).
Natural History. Little is known about the natural history or behavior of this species,
other than its ability to inhabit both freshwater and marine environments. It is reported to
search crevices for prey (Chan-ard et al. 2015; Das 2010).

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 13. Seasnakes

Viperine Seasnake
Hydrophis viperinus (Schmidt, 1852).
Distribution and Habitat. This species is widespread and is found in Bahrain, Bangla-
desh, Cambodia, China, India (Andaman Is., Nicobar Is.), Indonesia, Iran Iraq, Japan, Ku-
wait, Malaysia, Myanmar, Oman, Pakistan, Philippines, Qatar, Saudi Arabia, Singapore, Sri
Lanka, Taiwan, United Arab Emirates, and Vietnam. This species is notably absent from
Phillipines, and only one record off the island of Phuket exists
in Thailand. This species is associated with muddy bottoms,
warm coastal waters, lagoons, estuaries, creeks, and deeper
waters (15 - 30m). It’s enhanced ventral scales allow this spe-
cies to be slightly more mobile on land than other members
of Hydrophis, and can crawl across dry areas, though feebly
Figure 13.107. The distribution and with great effort (Rassmussen et al 2018; Uetz and Hosek
of Hydrophis viperinus. 2020; Egan, 2008). Type locality: Java, Indonesia.
Identification. Hydrophis viperinus is a relatively
stout-bodied, heavily-built snake sea snake, with wide ante-
rior ventrals, and ventrals clearly present along the length of
the body, making it fairly distinguishable from other mem-
bers of the genus. The species has a variable body pattern
often consisting of 25-34 rhomboidal spots running along
the dorsal side with a bicolored body of grey dorsally tran-
sitioning in a straight line to white on the lateral flanks and
white ventrally. Color and pattern are variable, and animals
in some localities are more commonly plain grey dorsally,
creams white ventrally with rhomboidal pattern indistinct
or diminished entirely. The head is large, wide, and distinct
from the neck, with complete head shields. Nasal shields
contact each other. Prefrontal scales are longer than they are
wide, and not in contact with the upper labials. The iris of
this species is pale with a dark round pupil. Post and pre-oc-
ular scales number 1-2, with 7-9 upper labial scales of which
3-5 border the labial scales. Body scales are juxtaposed and
hexagonal in shape with 27-34 scale rows on the neck and
37-50 scale rows at the mid-body. Adult males have spinose
protrusions on the distal edges of the body scales and the
anterior edges of the ventral scales. Ventral scales are nota-
Figure 13.108. Hydrophis vi- ble wide anteriorly, as wide as the body, which helps distin-
perinus. Art work from Ewart guish this species from the often-sympatric Hydrophis schis-
(1869). tosus. These ventral scales narrow anterior to twice the size
of adjacent dorsal scales posteriorly and number 226-274.
The average total adult length in males is 925 mm, of which 100 mm is comprised of the tail,
and 820 mm in females of which 80 mm is comprised of the tail. (Leviton, 2003; Egan 2008).
Natural History. Little is known about the natural history of this species. It is known to
forage shallow warm waters for prey, which has been documented to include flatfish (Kumo-
cocius), gobies (Gobiidae), and eels. This species gives birth to 3-5 live young in early summer
to notably large young (Rassmussen et al 2018; Egan, 2008).

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Estuarine Seasnake
Hydrophis vorisi (Kharin, 1984)
Distribution and Habitat. This species occurs in the Torres Strait, the Timor Sea, and
the Arafura Sea between Australia and Papua New Guinea, and is associated with estuarine
and shallow inshore waters (Cogger, 2018; Kharin 1984). Type locality: New Guinea (Kharin
1984).
Identification. This species head is very small, with an elongate body that is slender an-
teriorly, and compressed posteriorly. The body is a grayish white
with 70-80 light black bands which are twice as wide as the
patternless interspaces dorsally and taper ventrally. On some
specimens the bands break or are displaced along the ventral
line. The body scales imbricate, smooth anteriorly. But keeled
Figure [Link] distribu- posteriorly, and there are 29-35 scale rows at midbody. Ventrals
tion of Hydrophis voirsi. are small, not much larger than the adjacent scales posteriorly,
though slightly larger anteriorly and number 330-350 (Cogger,
2018; Kharin 1984; O’Shea 1996).
Natural History. This species is rarely encountered, and only known from two spec-
imens. There is little known about their natural history, but they are presumed to feed on
eels, and are viviparous. With almost nothing known about this species, it is listed as Data
Deficient by the IUCN (Rasmussen & Guinea 2010; Cogger 2018; O’Shae 1996).

Sepic Beaked Seasnake

Hydrophis zweifeli (Kharin, 1985)
Distribution and Habitat. Papua New Guinea, Australia (Queensland, Northern Terri-
tory). Habitat is only known from one specimen which was at the mouth of the Sepic River,
leaving he assumption it has a preference for gulfs. Type locality: “Off mouth of Sepik River,
PNG.”[No map.]
Identification. This species is very similar in appearance to Hydrophis schistosus, but
distinguishable based on the lack of preocular, which are fused with the supraocular in Hyr-
dophis zweifeli. Another distinguishing trait between these two species is the presence of a
split fourth supralabial in Hydrophis zweifeli which is absent in Hydrophis schistosus. Hy-
drophis zweifeli is distinguishable from all other sea snakes due to its beak like head. This
species has 36 body bands that taper ventrally, which are gray in color over and off-white
background. Midbody scale count is 48, ventrals number 271 (Ukuwela et al. 2013).
Natural History. This species is only known from a single specimen collected in 1966
off the mouth of the Sepik River and as such virtually nothing is known about its ecology.
Population size and population trends remain unknown (Rassmussen, 2018).

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Aquatic Snakes

14. Radiant Snakes and Gemsnakes

. The Radiant Snakes, the lamprophiids (Lamprophidae) are diverse, most are terrestrial,
others are fossorial or arboreal, and members of the genus Lycodonomorphus are semi-aquat-
ic. Family members occur in deserts, grasslands, and forests in temperate and tropical re-
gions. Radiant Snakes are part of the Elapoidea. They take vertebrate and invertebrate prey.
Some genera subdue prey with venom, others use constriction. Dental morphology is also
diverse and reflects the variety of prey they eat. Most species are oviparous. Lamprophiid
distribution is primarily in Africa and Madagascar with some representation in the Middle
East and Europe. Nine subfamilies are recognized, some of these are given family status by
some authors (Aparallactinae, Atractaspidinae, Atractaspidinae, Cyclocorinae, Lamprophi-
inae, Prosymninae, Psammophiinae, Pseudaspidinae, Pseudoxyrhophiinae)

Radiant Water Snake Clade,

Lycodonomorphus
The African Water Snake Clade is known from the countries of Angola, Belgian Con-
go, Botswana, Congo, the Democratic Republic
of the Congo, Malawi, Mozambique, Republic of
South Africa, Rwanda, Tanzania, Zambia, Zim-
babwe.
Lycodonomorphus is diagnosed by the fol-
lowing combination of characters (following
Kelly et al. 2011): body cylindrical and mod-
erately elongated; tail moderate to short; head
moderate to small, slightly broader than neck
and distinct from it; eye moderate, with vertical-
ly elliptical or sub-circular to round pupil; mid-
body scale rows 19–23 (rarely 25), vertebral row
not enlarged; dorsal scales smooth, without api-
cal pits or with two pits; ventrals 152–196, with-
out lateral keel; subcaudals paired, 37–89; cloacal
plate entire (exceptionally divided); maxilla with
18–25 teeth, subequal or slightly smaller or larg-
er posteriorly, diastema absent.
While most members of the genus are
aquatic - Lycodonomorphus inornatus seems to
be mostly, if not completely, terrestrial. A photo
is provided here for comparison purposes.

Figure 14.1. Olive Snake, Lycodonomorphus Tanganyika Water Snake

inornatus from Overberg, Western Cape, ZA Lycodonomorphus bicolor (Günther,
Province, South Africa. from Western Cape,
ZA Cape Town, Western Cape, ZA, South
1893)
Africa. Photography by Alex Rebelo. Arebe-
Distribution and Habitat. Endemic to Lake
lo23@[Link].
Tanganyika. Diurnal behavior is restricted to the
shallow water of the shoreline where they spend
their time on a rocky substrate.

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 14. Radiant Snakes and Gemsnakes

Natural History. Diet in composed entirely of cich-

lid fishes of the genera Leprologist, Chalinochromis, and
Xenotilapia. They are actively forage in the evening, but
moonlight depresses activity. Females were found gravid
throughout the year. The smallest gravid female was 50
cm in total length. Clutch sizes of 4, 5, and 8 eggs have
been reported. The only predators observed were crabs.
Figure 14.2. The distribution of Ly- Population densities of 9000 to 29,000 individuals per
codonomorphus bicolor. square kilometer were reported but estimates suggest
that as many as 38,000 snake per square kilometer are
possible (Madsen and Osterkamp 1982).
Barbault (1970) had reported a density
of 3390 snakes per square kilometer.

Dusky Bellied Water Snake

Lycodonomorphus laevissimus
(Günther, 1862)
Figure 14.3. Tanganyika Water
Snake, Lycodonomorphus bicolor
from Lake Tanganyika, County Nka-
si, RU, TZ District. Photography by
Hubert A. Szczygieł.

Distribution and Habitat. The east-

ern Cape from Transkei and Natal to
the southeast Transvaal. In Swaziland
it is restricted to the highveld region. It
in habits pools in forested streams. Type
locality: “East Indies” in error.
Natural History. More aquatic than L. rufulus, often
seen swimming, completely submersed, along the beds of
streams and rivers. Diurnal in the wild captives may be
more active at night. Highly susceptible to dehydration
and soon die without water. Diet includes frogs, tadpoles
(Rana, Phrynobatrachus, etc.), and fish (Tilapia). Large
animals are constricted but small ones are merely seized
Figure 14.4 Distribution of Lyco- and rapidly swallowed while still struggling. Prey caught
donomorphus laevissmus.
underwater are eaten without the snake surfacing for air
unless swallowing is prolonged when the snake will drag
its prey to a position above the surface before resuming (Raw 1973). Females lay 8-17 eggs
in the Southern Hemisphere’s summer.

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Aquatic Snakes

Figure 14.5. To the right. Dusky Bellied Water Snake, Lycodonomorphus laevissimus.
Photography by Kirsty Kyle.
Mulanje Water Snake
Lycodonomorphus leleupi (Laurent, 1950)
Distribution and Habitat. Southern Democratic Republic of the Congo (Zaire), eastern
Zimbabwe, western Mozambique, Malawi, and Zambia. Uses small streams, pans and vleis
(vegetated wetlands).
Natural History. It is diurnal. Feeds on frogs and their larvae. Two species have been
recognized. Females lay clutches of up to nine eggs. Broadley and
Coterill (2004) wrote, "This endemic water snake was described
from the Kundelungu plateau and also occurs on the Kibara pla-
teau in the P.N.U. between 1250 and 1810 m (Witte 1953). It was
described as a subspecies of L. whytei (Boulenger) of northern
Malawi and adjacent Tanzania, but was later thought to be con-
Figure 14.6. Distribution Ly- specific with L. rufulus mlanjensis Loveridge of southern Malawi
codonomorphus leleupi. and eastern Zimbabwe (Broadley 1967). These are now consid-
ered to be sister species, distinguished by ventral colouration:

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Aquatic Snakes

dark with yellow patches and a dark tail in L. leleupi and yellow with a few dark spots and a
dark median stripe on the subcaudals in L. mlanjensis. The nearest populations are separated
by a gap of 1000 km."

Mlanje Water Snake

Lycodonomorphus mlanjensis Loveridge, 1953
Distribution and Habitat. Eastern Zimbabwe, southern Mozambique, and Malawi. In-
habits swamps, pools, and rivers in montane grassland and savanna. Local name of the Mlan-
je Water-Snake: Chirumi (Manganja). Type locality: Ruo River, Mlanje Mountain, Malawi.
Natural History. Feeds on fish, frogs, and tadpoles.

FitzSimons Flood-plain Water Snake

Figure 14.7. Distribution Lycodonomorphus obscuriventris Fitzsimons, 1963
of Lycodonomorphus mlan-
jensis . Distribution and Habitat. The flood plains of the Lower
Shire Valley in southern Malawi in the north, through Mozam-
bique and Zimbabwe, into Kruger National Park through Eswatini (Swaziland) in the east
and southward into the Isimangaliso World Heritage Site, South Africa (Kyle et al. 2021).
Natural History. A nocturnal, but sometimes diurnal species that hunts frogs on flood-
plains. Kyle et al. (2021) reported a large gravid female laid a
clutch of eight eggs in early December 2012, confirming specula-
tions of oviparity in the species. Average egg length was 27.3 mm
(23.1–30.9 mm) and the average width was 15.9 mm (range
Figure 14.8. Distribution 15–17 mm). All eggs hatched in early February after approx-
of Lycodonomorphus ob- imal 60 days of incubation, and neonates were released at the
scuriventris capture site of the mother. Average total length of neonates was
147 mm (range 138–160 mm; N = 6) and average ventral scale
count was 173 (range 167–176; N= 5).

Common Brown Water Snake

Lycodonomorphus rufulus (Lichtenstein, 1823)
Distribution and Habitat. includes Zimbabwe, S Mozambique, Republic of South Africa
(Eastern Cape), Swaziland (it occurs throughout the country in the highveld, middleveld, low
veld Lesotho, Natal, Botswana, Malawi. Type locality: South Africa.
Natural History. Feeds on tadpoles (probably Rana angolensis). Taylor (1970) reported
one of these snakes swimming toward the bank in a stream in Rho-
desia's eastern highlands. The snake swam with the first few inches
of its body out of the water and had a tadpole in its mouth. It shift-
ed the tadpole in its mout several times while swimming (Taylor,
1970). and Lubombo regions, more common in the highveld.).
Figure 14.9. Distribution of
Lycodonomorphus rufulus.

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Aquatic Snakes

Figure 14.10. Common Brown Water Snake, Lycodonomorphus rufulus from Umgungundlovu,
KwaZulu-Natal, South Africa. Photography by Gus Benson.
Congo White-bellied Water-Snake
Lycodonomorphus subtaeniatus Laurent, 1954
Distribution and Habitat. Northern Angola to the Democratic Republic of the Congo
(Zaire) Congo (Brazzaville). Type locality: Keseki, near Kwamouth, Belgian Congo. Restrict-
ed to lake Upemba.
Natural History. A skink was found in the stomach of
one of these poorly known snakes.
Taxonomy. Branch (2018) wrote the following about this
snake. “Laurent (1954) described Lycodonomorphus subtae-
niatus based on a specimen from Keseki (DRC) and a series
of 12 paratypes, including four from Dundo, which remain
Figure 14.11. The distribu-
the only Angolan material. At the same time Laurent (1954)
tion of Lycodonomorphus sub- also described the subspecies L. s. upembae from Nyonga
taeniatus. (DRC). The type series for both taxa included material pre-
viously identified as Boaedon, i.e., B. virgatus and B. lineatus,
respectively (Laurent 1952; De Witte 1933). It was thus not unexpected that when preparing
a molecular phylogeny of house snakes of the description of their new species B. radfordii,
Greenbaum et al. (2015) found L. s. upembae embedded within Boaedon, to which it was
transferred. No new material was available to assess the generic relationships of L. subtaenia-
tus, but the current placement within Lycodonomorphus is problematic and its relationship to
B. virgatus with which has previously been confused should be investigated.”

Whyte's Water Snake

Lycodonomorphus whytii (Boulenger, 1897)
Distribution and Habitat. Inhabits small montane streams in southwest Tanzania, Mo-

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 14. Radiant Snakes and Gemsnakes

zambique, Republic of South Africa (Transvaal. Swaziland). Type locality: Fort Hill, (Chi-
pata), Malawi.
Natural History. Associated with upland streams and preys on anuran larvae and adults
(J.B. Rasmussen 2004). Loveridge (1955) wrote the follow-
ing, “Its stomach contained the hind legs of a frog (Rana
fuscigula) and the entire digestive tract was riddled with
worms. These have been identified by J. T. Lucker as Oxy-
uroidea, both sexes of a Kalicephalus, probably K. micrurus.
In the mesentery were two Dracunculus sp. and numerous
encapsuled larvae of one of the Physalopteridae.”
Figure 14.12. Distribution of Ly-
codonomorphus whytii.

Figure 14.13. Whyte’s Water Snake, Lycodonomorphus whytii..

Photography by John Valentine Lyakurwa.

White-bellied Water Snake

Boaedon upembae (Laurent, 1954)
Distribution and Habitat. Democratic Republic of the Congo (Zaire, Katanga; near
Lake Upemba). Type locality: Nyonga, Katanga, Belgian Con-
go. Laurent described this snake as a subspecies of Lycodono-
morphus subtaeniatus. Greenbaum et al. (2015) moved it to
the genus Boaedon based on molecular evidence. Laurent
(1954) commented on this species noting that Lycodonomor-
phus s. subtaeniatus and L. s. upembae bear a superficial re-
Figure 14.14. Distribution of semblance to Boaedon. This is why they were both confused
Boaedon upembae. with Boaedon lineatus, a common species often determined
by sight, without examining the number of scales. For the
typical form, this examination led to confusion with B. virgatus, whose numbers of scales
are similar, but which is nevertheless very different due to its characteristic ventral color-

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Aquatic Snakes

ation. (Laurent 1954)

Natural History. One specimen contained a catfish so it seems likely to be partly aquatic
(Greenbaum et al. 2015).

The Gemsnakes, Pseudoxyrhophiidae

The Pseudoxyrhophiinae was included in the family Lamprophiidae (Vidal et al. 2007),
but Zaher et al. (2029) raised it to family status, the Pseudoxyrhophiidae. About 88 species
in 21 genera compose the endemic Madagascar subfamily Pseudoxyrhophiinae.
Burbrink et al. (2019) found the Pseudoxyrhophiinae, the sister to a clade com-
posed of East African genera Amplorhinus and Duberria. Sister to all other pseu-
doxyrhophiines was the Socotran Island (Yemen) endemic Ditypophis. Burbrink
et al. (2023) found that the Pseudoxyrhophiidae originated in the early Miocene.
Aquatic habits have evolved in at least two genera of Gemsnakes, but they are poorly known.
The Pseudoxyrhophiinae was included in the family Lamprophiidae (Vidal et al. 2007),
however Zaher et al. (2029) upgraded it to family rank, the Pseudoxyrhophiinae. Pseudoxy-
rhophiidae is an indigenous clade to Madagascar, made up of about 88 species in 21 genera.
Pseudoxyrhophiinae is the sister group to the East African taxa Amplorhinus and Du-
berria, according to Burbrink et al. (2019). Sister to all other pseudoxyrhophiids was the
Socotran Island (Yemen) endemic Ditypophis. According to Burbrink et al. (2023), the early
Miocene is when the Pseudoxyrhophiidae first appeared. It is poorly recognized that at least
two genera of gemsnakes have adopted aquatic habits.
It appears that two other genera of Malagasy snakes, Thamnosophis (Cadle 2003; Glaw
& Vences 2007) and Madagascarophis, have semi-aquatic lifestyles (Rosa et al. 2016). For-
aging along streams has been documented for Madagascarophis meridionalis (Neaves et al.
2019). There is only one known instance of Thamnosophis mavotenda, and it was caught
in the Tsingy de Bemaraha National Park. It was swimming in a river when it was caught.
implying that it is a semi-aquatic species as well. (Glaw et al. 2009). The most aquatic species
are the three listed below, and some documented observations substantiate their frequent
usage of aquatic environments.
Closer examination will likely show more aquatic or semi-aquatic species in this family.
The genus Thamnosophis Jan 1863 is part of another radiation of Malagasy snakes. Cur-
rently, six species are included in the genus: Thamnosophis stumpffi, T. lateralis, T. in-
frasignatus, T. epistibes, T. martae, and T. mavotenda These species have 19 rows of dorsal
scales and minimal sexual dimorphism in tail length. During a recent survey in the Tsingy
de Bemaraha in western Madagascar, a single individual of a new Thamnosophis species
strongly resembles T. martae in scale arrangements but T. epistibes in live coloration. The
adjacent vegetation was disturbed by dry forest on calcareous ground. The snake proba-
bly tried to escape by swimming, having been disturbed by people walking in the river.
Jovanovic et al. (2009) report a successful predation event on the Malagasy poison frog Man-
tella aurantiaca; the observations were made in Torotorofotsy Wetlands, one of the few known
sites where M. aurantiaca occurs. The predation event was believed to be carried out by this
snake. After 30 minutes of observation on the frog Thamnosophis (probably T. lateralis, but
it could be confused with T. epistibes, a similar species that also inhabits this area.), as the
snake started to swallow the frog, it did not seem bothered by the taste. The toxins from the
frog may not bother this species. After ingestion, the snake showed no signs of intoxication.

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 14. Radiant Snakes and Gemsnakes

Six-lined Water Snake

Liopholidophis sexlineatus (Günther, 1882)
Distribution and Habitat. Madagascar (elevations up to more than 2500 m). close to
water including anthropogenic bodies of water (rice paddies) as
well as swamps, and marshes. Type locality: “Eastern Betsileo,
Madagascar.” North Fianarantsoa Province, in central Mada-
gascar. Widespread in the central highlands and south-east of
Madagascar. Additionally, there are records from Toamasina and
Mangindrano on the north end of the island.
Figure 14.18. The distribu- Liopholidophis Mocquard, 1904 is a genus endemic to Mad-
tion of Liopholidophis sex- agascar. The genus was recently revised by Cadle (1996, 1998)
lineatus. whorecognized nine species in two species groups. The Liop-
holidophis sexlineatus group contains L. sexlineatus L. varius L.
dolicocercus, L. grandidieri and L. rhadinaea. These species have 17 rows of dorsal scales and
are characterized (with the exception of L. varius) by an extreme sexual dimorphism in tail
length, the tails of the males being much longer. Accordingly, the number of subcaudals is
much higher and the total length is longer in males than in females.
Natural History. A semi-aquatic, diurnal snake that inhabits open areas near water (Glaw
and Vences 2007). Diet includes frogs in the genera Ptychadena Heterixalus, and Boophis and
the skink Trachylepis madagascariensis. It is viviparous with litters of 4-10 young reported in
December.

Figure 14.19 Liopholidophis sexlineatus.

Photography by Konrad Mebert.

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Aquatic Snakes

Lateral Water Snake

Thamnosophis lateralis Duméril, Bibron and Duméril, 1854
Distribution and Habitat. Widespread in Madagascar
[No map.] Mostly terrestrial but also found in water, in-
cluding rice paddies. Cadle (1996) examined 20 specimens
and found the largest male 729 mm total length, 212 mm
tail length (29% of total; tip of tail missing). The largest fe-
males was 820 mm total length, 234 mm tail length (29%
Figure 14.20. The distribution of of total). Natural History. Diet is mostly frogs. Clutches of
Thamnosophis lateralis. 6-13 eggs.

Figure 14.21. Thamnosophis lateralis.

Photography by Gonçalo M. Rosa.

Yellow-striped Water Snake

Thamnosophis stumpffi (Boettger, 1881)
Distribution and Habitat. Forests of northeast Madagascar. The type locality is the island
of Nossi-Bé (no map).
The stumpffi group contains T. stumpffi , T. lateralis, T. infrasignatus, and T. epistibes.
These species have 19 rows of dorsal scales and do not show an extreme sexual dimorphism
in tail length. They are distributed throughout eastern Madagascar from Montagne d’Ambre
in the north to Andohahela in the south, ranging from sea level to about 4700 ft. In addition,
T. lateralis, T. stumpffi, and T. epistibes are also known from less humid localities in western
Madagascar.
Natural History. A diurnal, mostly terrestrial snake inhabiting low elevation forests.
It uses primary closed canopy forests and forests that have been disturbed (Blumgart et al.
2017). It is threatened by habitat fragmentation, deforestation, and agricultural expansion.

296
 14. Radiant Snakes and Gemsnakes

Figure 14.22. Thamnosophis stumpffi

Photography by Paul Freed.

Next page. Pseudoxenodon bambusicola. Photography by Konrad Mebert

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Aquatic Snakes

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Aquatic Snakes

The Dipsadidae Section

The snake on the previous page is an arboreal Imantodes from Peru - its not aquatic. I use
it to mark the begining of the chapters on the dipsadids.
The Dipsadidae (or Dipsadinae) are ecologically and morphologically diverse; some are
arboreal, others are terrestrial, fossorial or aquatic. Most are oviparous, but some Helicops ap-
pear to be facultatively viviparous. Many prey on vertebrates, yet others feed on soft-bodied
invertebrates such as earthworms, snails, and slugs.
The clade contains 843 species that are mostly restricted to the neotropics, but some
occur in North America.
However, the sister clades to the dipsadids are the bamboo snakes of the family Pseu-
doxenodontidae (genera Pseudoxenodon and Plagiopholis) following Zaher et al. (2019).
They found the immediate sister to the bamboo snakes was the Tibetian hot springs Snakes
of genus Thermophis. Surprisingly. the sister to Thermophis was the North American clade
Carphophiinae that includes the genera Carphophis and Contia. This section covers the next
eight chapters each discussing one or more subclades.

Figure 1. Modified from Zaher et al. (2019) shows the position of the Bamboo Snakes as the sister to
the dipsadids and the basal position of Asian Thermophis and North American Carphophis Heterodon,
and Farancia, in the Dipsadidae.

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Aquatic Snakes

15. Bamboo Snakes, Pseudoxenodontidae

Pseudoxenodontidae contains 10 species in two genera. Several species are polytypic.
They are found in South and Southeastern Asia, from northeast India to southern China
(including Taiwan) and south into Indonesia as far east as Wallace’s Line. Little is known
about them. Bamboo Snakes spread a small hood as part of their defense behavior they are
commonly referred to as false cobras.
They are small to medium-sized oviparous snakes. Pseudoxenodon occurs along streams
in wet forests (Rahadian and Das 2012; Stuart and Heatwole 2008). Plagiopholis are more ter-
restrial (Zhong et al. 2015). The two genera (Pseudoxenodon and Plagiopholis) have similar
hemipenes morphology and molecular data confirmed their close relationship (McDowell
1987; Pyron et al. 2011). They were recovered as the sister to the Dipsadidae by Zaher et al.
(2019). The degree to which these snakes are semi-aquatic is not known. The literature sug-
gests that some of them are semi-aquatic.

Common Bamboo Snake

Pseudoxenodon bambusicola Vogt, 1922
Distribution and Habitat. Leaf litter and moist ground under large objects they are gen-
erally found in inland wetlands, or close to permanent water from southern and southeastern
China, central Laos, northern Thailand, and northern and
central Vietnam.
Natural History. Diet includes frogs of the genus Micro-
hyla. This has an impressive threat displays, including flash-
ing boldly banded ventral patterning and bright yellow col-
oration, spreading a hood, and death feigning (Pope 1935).

Figure 15.1. The distribution of

Pseudoxenodon bambusicola.

Figure 15.2. Pseudoxenodon bambusicola.

Photographed by greenlapwing@[Link].

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 15. Bamboo Snakes, Pseudoxenodontidae

Baramen Bamboo Snake

Pseudoxenodon baramensis (Smith, 1921)
Distribution and Habitat. Malaysia (Borneo, Sarawak). Type
locality: “Mount Dulit, Sarawak, North Borneo, at 1,000 metres.”
[=Banjaran Dulit, 3rd Division, Baram District, northern Sar-
awak, central East Malaysia, 1000 m, 3˚15’N, 114˚15’E]. Stue-
bing et al. (2014) notes the species is known from a few submon-
tane sites in Sabah, Sarwak, and Kalimantan at elevations above
1000 m ASL A photograph of this species (Fig. 15.4. Pseudoxeno-
Figure 15.3. The distribu- don baramensis Photography by Abhijit Das.)
tion of Pseudoxenodon bara-
mensis

Figure 15.4. Baramen Bamboo Snake Pseudoxenodon baramensis.

Photography by Fenoy Xavier.

On the next page. Is a plate from Wall's 1907 paper that described what is now Thermophis
baileyi.

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Aquatic Snakes

302
 16. The Basal Dipsadids

16. The Basal Dipsadids

It is unexpected to find the closest ancestor of New World dipsadid snakes that are most-
ly tropical exist in a landscape that looks like the one to the left. Yet, the Tibetan hot springs
snakes in the genus Thermophis represent a lineage that moved from Asia into the West-
ern Hemisphere. The literature suggests that at least some dipsadid species are semi-aquat-
ic, while others are more terrestrial. The only other Asian dipsadid is the gastropod-eating
Stichophanes ningshaanensis formerly consider a member of the genus Oligodon (Wang et al.
2014). Messenger and Wang (2015) found Stichophanes to be terrestrial and oviparous. Basal
dipsadids also contain the Carphophiinae (the North American fossorial genera Carphophis
and Contia) neither are semi-aquatic. Zaher et al. (2019) found these snakes to be the sister
to the highly aquatic mud snakes of Farancia.

Bailey’s Hot-spring Keelback

Thermophis baileyi (Wall, 1907)
Distribution and Habitat. China (Tibet = Xizang, Lhasa region), 3000-4000 m elevation
or higher. Type locality: Tibet. Dorge et al (2007) surveyed locations that offered suitable
habitat and found it closely associated with hot-springs.
Altitudinal distribution is 3600 to 4900 m asl. The species
occurred in glacial refugia associated with hot-springs
during the uplift of the Tibetan Plateau and during the
cooler periods of the ice ages.
Natural History. Diet appears to be restricted to
metamorphs of the frog Nanorana parkeri and fishes of
Figure 16.1. The distribution of the genus Schizopygopsis. The areas around hot springs
Thermophis baileyi. are used for basking on the warm ground between rocks
or along riverbanks. They are particularly active after
rain. Some individuals were observed in the warm pools of hot springs and in adjacent rivers.
The water temperatures of the hot springs were relatively constant over the season and ranged
from 29 to 33°C. River temperatures were much lower (8–11°C) and did not vary during a
given day. At ground level, temperatures rose to 35°C as early as March. Low-sulfur sites
along rivers provide food and basking sites for thermoregulation. However, the temperature
of such thermal sites plays no role, or only a secondary role, in determining the species’ pres-
ence. (Dorge et al. 2007, Hofmann et al 2014).

Yunnan Hot-spring Keelback

Thermophis shangrila Peng et al, 2014
Distribution and Habitat. Shangri-La County, Yunnan, China. The holotype was in a
grassland at the forest edge near a hot spring about 500 m in Shangri-La, Northern Yunnan,
China, at 12:30 on 23 August 2011, when it was moving towards the forest. The authors (Peng
et al. 2014) considered this snake terrestrial – but based on a limited observation.
Natural History. A diurnal and possibly terrestrial snake. The temperatures recorded at
the mouths of hot springs at the 31 known sampling sites of Thermophis exceed 40°C. There
are 229 hot springs with the temperature hotter than 40°C in the area, raising the questions
why only a few hot springs are utilized by Thermophis and why did snakes sympatric with
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Aquatic Snakes

Figure 16.2. Thermophis baileyi Photography by Sylvia Hofmann.

Thermophis in earlier geologic periods fail to survive in these regions?

Sichuan Hot-spring Keelback

Thermophis zhaoermii Guo, Liu, Feng & He, 2008
Distribution and Habitat. Found at high elevations in
the Hengduan Mountains of China. Recent investigations
show the Sichuan Hot-spring Snake is restricted to geo-
Figure 16.3, Distribution of Ther-
thermal zones above 3000 m ASL. The type locality, Litang
mophis zhaoermii. County, Suchuan, China, elevation 3700 m.
Survival of hot-spring snakes is seriously threatened. The
exploitation of geothermal resources and the construction
of hydropower stations are the major threats to its survival.

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 17. New World Mud Snakes

This is especially the case for the Sichuan hot-spring snakes. The development of hot-spring
tourism in Sichuan may encroach on T. zhaoermii habitat, resulting in habitat degradation,
water pollution, and limited access to food resources (Ren et al. 2022). Thermophis zhaoermii
only occurred in about one-quarter of the investigated hot-springs. Before the construction
of hydropower stations and exploitation of geothermal resources, Ren et al. (2022) propose
detailed population surveys should be conducted to guide policy decisions.

Figure 16.4. Thermophis zhaoermii. Photography by Sylvia Hofmann.

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Aquatic Snakes

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 17. New World Mud Snakes

17. New World Mud Snakes

The two species that compose the genus Farancia are considered some of the most aquatic
snakes in North America, but both species are known to wander overland (Steen et al. 2013).
Droughts may trigger their movement from a wetland, and excessive rainfall may also trigger
their movement. Females will also move overland to nest. Both species take aquatic prey. Sem-
litsch et al. (1988) have described juvenile emergence in both the Fall and Spring and note that
hatchlings may benefit from Fall emergence by feeding prior to their first winter. However,
Fall emergence may present the hatchlings with dried or reduced aquatic habitats while spring
emergence would have abundant habitat and food (larval salamanders).
The members of the genus are known to have maternal attendance of eggs; a highly spe-
cific diet of elongated prey; American eels (Anguilla rostrata) and the eel-like salamanders of
the genera Siren and Amphiuma. Knowledge about even basic aspects of their biology may
be subject to revision. For example, relatively recently both species but were found to make
overland movements far from water (Steen et al. 2013).

Red-bellied Mud Snake

Farancia abacura (Holbrook, 1836)
Distribution and Habitat. The Coastal Plain of the southern United States from southern
Virginia southward throughout Florida; follow the Mississippi drainage as far north as South-
ern Illinois and west to eastern Texas and the occur the Coastal Plain and range into the Pied-
mont of western Georgia. They occur in seasonal wetlands, ditches, Carolina bays, cypress
swamps, marshes, slow-moving streams, and the heavily vegetated lakes and ponds margins.
Type locality: South Carolina. This is a polytypic species, with an eastern race and a western
race and a zone of intergration between them. The Western Mud Snake, F. a. reinwardtii has
fewer than 52 red [Link] nominate race has more than 53 red bars
Natural History. These are highly aquatic and spend much of their lives in submerged
vegetation and debris. They seldom bask out of the water and are seen on rainy nights when
they may traverse roads into adjacent aquatic habitats. Prey
is usually the giant aquatic salamanders in the genera Am-
phiuma and Siren; young snakes consume other amphibians
(usually larval anurans and salamanders). However, Faran-
cia abacura feeds heavily on mole salamanders, Ambystoma,
in wetlands without fishes, despite giant salamanders being
relatively common in those habitats (Durso et al. 2013).
Eggs are laid in summer (July-August) and hatch in Septem- Figure 17.1. The distribution
ber-October, often in sandy upland habitats near water, and of Farancia abacura.
females may attend the eggs until hatching. Eggs are some-
times laid in American Alligator nests and the distribution of mud snakes is similar to, and
overlaps that of alligators. The eight-week incubation fits well with a strategy of commensal
nesting with alligators, whose eggs have about a nine-week incubation period. Several turtles
have also been found to have commensal nesting with alligators. This strategy may provide
nesting snakes and turtles, as well as their eggs, with a degree of protection from egg predators
and provide an excellent medium for egg incubation, thus decreasing maternity costs. This is
particularly true if the nest is actively defended by an adult crocodilian, because depredation
On the previous page. Farancia erytrogramma.
Photography by William W. Lamar.

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Aquatic Snakes

rates are markedly higher in undefended nests (Metzen, 1977; Crawshaw, 1991).
Large females may lay more the 100 eggs; but the average clutch size is about 25 eggs.
Hatchlings have total lengths of 158-231 mm.

Figure 17.2. Farancia abacura

Photography by Jake Scott.

Rainbow Snake
Farancia erytrogramma Palissot De Beauvois, 1801
Distribution and Habitat. There is an isolated population in southern Florida. Otherwise,
the distribution is from the Potomac drainage of southern Maryland southward through the
Coastal Plain into north-central Florida, westward into eastern Louisiana. Farancia erytro-
gramma inhabits streams that are alkaline (calcareous) to pH neutral waters. Type locality:
“l’Amerique septentrionale.” Restricted by Harper, 1940, to “the lower Cooper River, in the
vicinity of Charleston, S. C.”
Natural History. Mostly nocturnal. Adults prey almost exclusively on the common fresh-

308
 17. New World Mud Snakes

water eel, Anguilla rostrata but they will also eat salamanders.
This species was included in the study done by Durso et al. (2013)
but the sample size was too small to include in their analyses.
They observed this species feeding heavily on Ambystoma at one
fishless wetland site. Eggs are laid in early summer and hatch in
Figure 17.3. The distri-
bution of Farancia erytro-
mid-August with 75-80 days of incubation. Females may attend
gramma. the nest. They do not spend the winter in the water at the north-
ern edge of their distribution while they do so in the southern
portion of the range (Neill 1964, Steen et al. 2013). Durso and Laverick (2021) discovered a
dead specimen entangled in plastic mesh that had been placed to reduce soil erosion. The
same plastic mesh had also trapped a cottonmouth which was dead.

Figure 17.4. Farancia

erytrogramma Pho-
tography by William
W Lamar.

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Aquatic Snakes

310
 18. Wetland Coniophanes

18. Wetland Coniophanes

About 17 species of Black-striped Snakes of the genus Coniophanes range from Texas
to Peru. They are venomous with rear fangs, and have long fragile tails – presumably to aid
in escaping predators. Coniophanes snakes typically reach a maximum length of 31–46 cm
with a brown body and black stripes running down the sides and center of the back, as well
as a red or orange ventral scales. However, some species have no stripes. They spend much
of their time in burrows beneath decaying plants, in loose soils, or in leaf litter on the forest
floor. They are nocturnal and forage on the surface for small vertebrates. Coniophanes are
oviparous, depositing clutches of up to ten eggs in loose soil. Depending on the humidity and
relative temperature, the eggs hatch in about 40 days. Hatchlings are about 17 cm in length.
Most are terrestrial, leaf-litter dwelling species, at least two are semi-aquatic (Lee 1996).

Two-spotted Snake
Coniophanes bipunctatus Günther, 1858
Distribution and Habitat. Moist lowland forest, in-
cluding swampy areas, has also been taken in ciénegas
(stagnant standing water bodies) from southern Mexico to
northwest Panama. Dry forest (Köhler 2008) and disturbed
forest (McCranie 2011) are also used. Henderson and Ho-
Figure 18.1. The distribution of evers (1977) considered this species aquatic in the Orange
Coniophanes bipunctatus. Walk area of Belize. McCoy (1969) considered both species
aquatic and present in the mangrove forests and coastal la-
goons of the Yucatan. Type locality: Unknown.
Natural History. Crepuscular and nocturnal. Feeds
on fish, frogs, and lizards. Nahuat-Cervera and Barrao-Nobrga (2021) report a specimen
attempting to eat the frog Lithobates [Link] of up to six eggs were reported in
June. The image to the left is Coniophanes bipunctatus. Photo by Ruth Pecino Daniel.

Five-striped Snake
Coniophanes quinquevittatus Duméril, et al. 1854
Distribution and Habitat. Inhabits mangrove forests
and coastal lagoons in Mexico (Chiapas, Tabasco, S Vera-
cruz, Yucatan, Campeche), and northern Guatemala. The
distribution seems to be discontinuous. Type locality: Un-
Figure 18.3. The distribution of known.
Coniophanes quinquevittatus. Natural History. Poorly known. Conant (1965) found
this species and its congener C. bipunctatus in the same Ty-
pha-Pontederia-Thalia marsh southwest of Coatzacoalcos,
Veracruz. The Tiger Heron has been reported as a predator
on this species

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312
 19. Neotropical Swamp Snakes

19. Neotropical Swamp Snakes

This chapter discusses two clades of dipsadid snakes the poorly known Prawn Snakes of
the genus Hydromorphus and the Swamp Snakes of the genus Tretanorhinus. A related clade
consists of the arboreal-terrestrial Imantodes and Leptodeira (Zaher et al. 2019). While Lep-
todeira is considered semi-aquatic by some authors my (JCM) experience with them is that
they are much more arboreal and terrestrial, and are dietary specialist on frogs, tadpoles, and
frog eggs from non-aquatic foraging sites.
Perhaps the most interesting aquatic dipsadids, are the snakes in the genus Tretanorhi-
nus. Four species are known from Ecuador, Central America, Cuba, and the Bahamas. The
long head, square muzzle, and long tail suggest this snake may be convergent with the Ten-
tacled Snake (Erpeton tentaculatus) from Southeast Asia. Few herpetologists have examined
these snakes since Wilfred Neil observed them in Cuba in 1949 and reported them hunting
at night in shallow water and described them as highly aquatic. They move very slowly, use
freshwater, brackish water, and possibly full-strength sea water. Neill observed them snapping
at small fish in shallow pools stating that they were quite successful in capturing prey.

Prawn Snakes, Hydromoprhus

Hydromorphus are medium-sized snakes (males to 690, females to 797 mm TL) with
rounded heads barely distinct from the neck, and moderately short tails (13-19% of TL).
Eyes are small, with a round pupil. Nostrils are directed dorsally. Head scales are typically
except for having 1 or 3 internasals and 1 or 3 prefrontals. Nasals are usually entire but may
be partially or entirely divided below the nostrils. A loreal bordering the orbit usually present.
A single, upper preocular may be present (the lower preocular is usually fused to the loreal)
but two or more may be found Two pairs of chin shields with a mental groove variably de-
veloped or absent. Dorsal scales are smooth (except for supra-cloacal keels in one species),
in 15- 17 rows at midbody, 13-15 near the vent. Cloacal plate and subcaudals are divided.
Ventrals number 157-186, subcaudals 33-54. Dorsal color is dark gray to dark brown or olive
green somewhat paler on rows 1-2. The venter is yellowish or cream-colored with variable
dark smudging along the edges and the center of the ventrals, more abundant on the body’s
posterior portion (Villa 1990).

Peter’s Prawn Snake

Hydromorphus concolor Peters, 1859
Distribution and Habitat. Atlantic lowlands and premon-
tane slopes from Guatemala to Panama (Honduras, Nicara-
gua, Costa Rica, Panama); Colombia; reaches 900 m elevation
in Honduras. Inhabits streams and the adjacent forest floor in
wet forests. Type locality: Not given, presumably, Costa Rica.
Figure 19.1. The distribution of Natural History. Diet may include prawns as well as oth-
Hydromorphus concolor. er aquatic prey. Active day and night. When in the water, of-
ten at the bottom in debris. Active on land after heavy rains.
Clutches of four and seven eggs in June and hatch in November and December.
Previous page. A Tretanorhinus variabilis wagleri.
Photography by Tomás Michel Rodríguez Cabrera.

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Aquatic Snakes

Figure 19.2. Hydromorphus concolor.

Photography by Twan Leenders (upper two ) Fabio Hidalgo (bottom).

314
 19. Neotropical Swamp Snakes

Dunn’s Prawn Snake

Hydromorphus dunni Slevin, 1942
Known from a single specimen from Chiriqui, Panama
at 1250 m in premontane forests. Total length of 482 mm.

Swampsnakes, Tretanorhinus
The Neotropical Swampsnakes are nocturnal and high-
ly aquatic. Currently, four species are recognized, and the Figure 19.3. The distribution of Hy-
dromorphus dunni.
genus remains poorly studied. Tretanorhinus is exclusive to
the Neotropics where it inhabits fresh and brackish water
bodies such as rivers, streams, lagoons, estuaries, mangroves, and cow wells. They seem to
require a muddy or rocky substrate with aquatic vegetation where they can hide and rest. But
there are reports of individuals found out of the water, usually, after flooding, that forces the
snakes to search for another water body. The distributions imply the ability of these snakes to
disperse and survive in across saltwater barriers (Barbour and Amaral 1924; Neill 1958) since
they have colonized several islands.
Dunn (1939) described the generic traits as: “Dorsal scales keeled, striate, pitless, re-
ducing by dropping the paravertebrals, 21-17; subcaudals double; cloacal plate double; head
scales normal with at times two loreals and at times one or three prefrontals; internasals
small; pupil circular to slightly oval; maxillary teeth about 27, subequal.”
Zaher et al. (2019) found this genus to be paraphyletic with T. variabilis and T. nigrolute-
us separated by quite a few nodes, thus some generic changes may be expected.

Mocquard’s Swampsnake
Tretanorhinus mocquardi Bocourt, 1891
Distribution and Habitat. Lowland forest swamps on
the Pacific coasts from Panama to Ecuador. Type locality:
“Belize” in error. Dunn (1939) states that it is Panama City.
There is apparently one record from eastern Panama that
may be the result of dispersal along the Canal.
Natural History. Presumably, fish and amphibians are
the prey of this species.
Figure 19.4. The distribution of
Orange-bellied Swampsnake Tretanorhinus mocquardi.
Tretanorhinus nigroluteus Cope, 1861
Distribution and Habitat. Pacific and Caribbean slopes
of Mexico (Chiapas, Oaxaca, Tabasco, Quintana Roo,
Campeche), Guatemala, Honduras (Islas de la Bahia and
Utila), Nicaragua, Belize, El Salvador uncertain, Costa Rica,
Panama. Records of this species in Colombia are contro-
versial (Barquero and Arguedas (2019). Inhabits streams in
the Chocó Tropical Humid Forest and rivers and coastal la- Figure 19.5. The distribution of
goons in Quintana Roo, Mexico (Cedeno-Vazquez et al. 2021).Tretanorhinus nigroluteus.
315
Aquatic Snakes

Natural History. Diet.

Known to feed on poecilid
fish (P. sphenops and P. mexi-
cana) (Cedeno-Vazquez et al.
2021).

Striped Swampsnake
Tretanorhinus taeniatus
Cope, 1861
Distribution and Hab-
itat. The Pacific coast of
Colombia and Ecuador. In
Colombia reported from the
Valle del Cauca. In Ecuador
it is only known from Esmer-
aldas Province, at an altitude
of approximately 140 m. Type
locality: River “Sapayo”, Es-
meraldas Province, Ecuador.
Natural History. Aquat-
ic in fresh water, black water,
and salt water. When threat-
ened they dive for conceal-
ment.

Caribbean
Swampsnake
Tretanorhinus variabilis
Duméril, et al., 1854
Distribution and Habi-
tat. Swamps, streams, includ-
ing coastal situations with
brackish water. Cuba, Isla de
la Juventud, Cayman Island.
Type locality: Unknown.
Natural History. Diet.
López-Hurtado et al. (2020)
document fish, frogs, and
freshwater crabs in the diet Figure 19.6. Tretanorhinus nigroluteus Cahuita, Costa Rica.
of this species. The diversity Photography by Daniel Jablonski.
of the prey suggests this snake
uses a variety of foraging strat-
egies. Petzold (1967) reported a captive female laid eight eggs on 31 August, on wet moss on
the border of the water basin in the terrarium. Four eggs were saved, and the temperature
was 30-32 C during the day, 20-25 C at night). Egg sizes were: 33 x 18, 35 x 16, 36 x 16, and

316
 19. Neotropical Swamp Snakes

Figure 19.7. Tretanorhinus variabilis.

Photography by Tomás Michel Rodríguez Cabrera.

317
Aquatic Snakes

36 x 17 mm.; the average weight was 5.4 grams. After an

incubation period of 35 days. The total length of the new-
born ranged from the pattern of the young snakes resem-
bles that of the adults, but in the young, there is greater
contrast between the spots and the ground color. Males
possess tubercles on the shields of the lower side of the
head-on the mental, the anterior chin shields, and the first
Figure 19.8. Distribution of Tre-
tanorhinus taeniatus lower labials (here very clearly). The females lack these tu-
bercles (Petzold 1967). When disturbed the body is rolled
up like a ball with the head hidden in the middle (Petzold
1967).
This is a polytypic species that needs further investi-
gation using molecular data.
Tretanorhinus variabilis variabilis Duméril, Bibron &
Duméril 1854. Distribution: Cuba, from La Habana Prov.
Figure 19.9. The distribution of in the west, east into Guantánamo Prov. (except for region
Tretanorhinus variabilis. about Sierra Maestra, Granma Prov.). Type locality: Un-
known.
Tretanorhinus variabilis binghami Schwartz & Ogren
1956. Distribution: Cuba, Granma Prov., lowlands, and
lower foothills of Sierra Maestra. Type locality: Finca Bu-
cares, 22 km S Bueycito, in the Rio Yao, Granma Province, Cuba.
Tretanorhinus variabilis insulaepinorum Barbour 1916. Distribution: Isla de la Juventud;
Type locality: Isla de la Juventud.
Tretanorhinus variabilis lewisi Grant 1941 Type locality: North Side, Grand Cayman Is-
land, Cayman.
Tretanorhinus variabilis wagleri Jan 1863. Distribution: Cuba, Pinar del Rio Province.
Type locality: Brazil (in error)

On the next page. Hydrodynastes gigas, Photography JCM.

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Aquatic Snakes

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Aquatic Snakes

20. Neotropical River Snakes

Zachary J. Loughman and John C. Murphy

Two snakes, Herman’s Water Snake (Hydrodynastes bicinctus) and the False Water
Cobra (Hydrodynastes gigas) constitute the dipsadid tribe Hydrodynastini. Both snakes are
aquatic, with H. bicinctus rarely being found outside of forests streams, and rivers. Hydro-
dynastes gigas are among South America’s largest colubroid snakes. Specimens in excess of
2.5 m weighing more than 5 kg are frequently encountered. Like most xenodontines, both
species occur over a wide portion of northern and central South America. Hydrodynastes
bicinctus occurs primarily in Amazonia and portions of the Cerrado, while H. gigas have a
wide distribution across several South American biomes. Ecologically, both species could be
considered habitat specialists given their habitat associations with wetlands, streams, and riv-
ers. Hydrodynastes bicinctus is piscivorous and a feeding specialist. Hydrodynastes gigas have a
much more generalized diet and will readily eat any vertebrate small enough to be consumed.
Both species have enlarged opisthoglyphous teeth in the posterior portion of their upper jaw.
Several bioactive peptides have been retrieved from both species Duvernoy’s glands. Due to
this, most herpetologists consider Hydrodynastes to be rear-fanged venomous snakes.

Natural History & Diversity

Hydrodynastes bicinctus and H. gigas have both similar and different ecologies. While
both species are aquatic, H. bicinctus appears to have stronger affinities for a truly aquatic
existence. The majority of H. bicinctus found in the field are found in water. Furthermore, H.
bicinctus is frequently encountered in lotic waterways, often under rocks and in woody snags.
Wetland environs are the preferred haunts of H. gigas, particularly those choked with mac-
rophytes. Unlike H. bicinctus, H. gigas is often encountered outside of water, though aquatic
habitats are never far away. While both species have these preferences, it is possible to en-
counter H. bicinctus and H. gigas in any aquatic environment across their respective distri-
butions. Both species' populations decline in lentic and lotic systems with moderate to heavy
anthropogenic presence (Loughman 2023).
Both Hydrodynastes are diurnal and are rarely encountered in the open after dark. Hy-
drodynastes gigas often is observed basking on mats of aquatic vegetation during morning
hours; H. bicinctus rarely is encountered outside of water. Hydrodynastes gigas diel activity is
strongly associated with morning and evening hours. Refuge is sought during the heat of the
day, especially during dry seasons. During transitional months between dry and wet seasons
H. gigas is active during all hours of the day (Giraudo et al. 2014). Hydrodynastes gigas activ-
ity declines as the dry season reaches its zenith. During this time, activity is limited to early
morning hours, overcast days, and evening hours (Giraudo et al. 2014).
Little is known about H. bicinctus dietary habits other than they consume fish and am-
phibians. The feeding ecology of H. gigas has been studied extensively (Lopez and Giraudo,
2014). Hydrodynastes gigas are feeding generalists, and readily consume vertebrates they en-
counter (Lopez and Giraudo, 2014; Ballini et al. 2015). Specifically, amphibians and fish are
consumed by all demographics (Giraudo et al. 2014; Lopez and Giraudo, 2014). As H. gigas
mature and increase in size, snakes and mammals are added to their dietary repertoire. By
the time they reach adulthood, H. gigas diets include any moderate to small sized vertebrate
they encounter. Ballini et al. (2015) studied the feeding habits of a South American snake
community in eastern Argentina. Hydrodynastes gigas were part of this community and had

320
 20. Neotropical River Snakes

the evenest prey selection of any species studied. Amphibians, reptiles, fish and birds were all
consumed in this respective order – all vertebrate classes were consumed.
Hydrodynastes gigas predatory behavior is similar to other aquatic snakes. Prey when
encountered in water is captured by H. gigas waving its head, mouth agape, until contact
occurs. Prey is grasped, and depending on prey size, either rapidly consumed, or constrict-
ed, dispatched, and then consumed. Duvernoy's secretions and envenomation seem to serve
little to no role in prey acquisition. Hydrodynastes gigas is an enthusiastic feeder and will
often consume multiple prey items at once if the opportunity presents itself. Strussman and
Sazima (1970) noted a unique H. gigas feeding behavior. Large H. gigas were observed using
the posterior portion of their bodies to flush fish and tadpoles from macrophyte beds out to
deeper water. Hydordynastes gigas were observed using the first third of their bodies, which
were held in an arc, to corral flushed fish into their open mouths.
Several other unique behaviors are exhibited by these snakes outside of prey aquisition.
Hydrodynastes gigas when disturbed are well known for flattening the anterior portion of
their bodies, producing hoods similar to cobras. Interestingly, H. gigas can flatten the entire
body, not just the cervical region, and will do so if a threat is perceived. During these displays,
the head is often lifted off the ground and the tongue flicked slowly and deliberately. If the
perceived threat advances, H. gigas will use the last two-thirds of their bodies as a whip, and
bludgeon their attackers (Loughman, 2023). If this does not dissuade the attacker, thanatosis
may occur. Both adults (Marques et al. 2013) and neonates (Loughman, 2023) play dead.
Variation in thanatosis frequency occurs at the population level – some H. gigas populations
readily employ thanatosis while others rarely do (Marques et al. 2013).
Hydrodynastes are opistyghlyphous. Both species have enlarged teeth in the posterior
portion of their mouths (Loughman 2023). Bites from H. bicinctus are rare. The effect of
H. bicinctus Duvernoy’s secretions on mammals is poorly understood. Hydrodynastes gigas
Duvernoy’s Gland secretion have been studied and are well understood. Hydrodynastes gigas
Duvernoy’s secretions have high concentrations of proteins, specifically azocasinases, casein-
ases, and metalloproteinases (Hill and Mackessy, 1997). The latter enzyme is responsible for
the frequent comparison of H. gigas to Timber Rattlesnakes (Crotalus horridus). Metallopro-
teinases are primary components in several viper venoms. They are powerful enzymes that
target fibrinogen, which is used to hold cells together. While H. gigas possess metallopro-
teinases, venom delivery is very inefficient. Most people who have reactions to H. gigas bites
experience localized edema. Bigger snakes deliver more venom and often produce the most
virulent reactions.
Reproductive information for H. bicinctus is lacking, though a specimen brought into
captivity laid 17 eggs. The reproductive biology of H. gigas is well understood. Hydrodynas-
tes gigas frequently produce clutches of 30 or more eggs in human care (Loughman, 2023).
In nature, H. gigas clutches normally number from 15 to 25 eggs. Vitellogenesis occurs in
September, and oviductal eggs were present in northeastern Argentina populations from Oc-
tober through December (Giraudo et al. 2014). Hydrodynastes gigas is considered a discon-
tinuous breeder. Males are always willing to mate, and like other aquatic snakes will form
breeding balls around receptive females (Loughman, 2023). Follicles were present year-round
in road-killed animals indicating the possibility of multiple clutches being laid in a single year
(Giraudo et al. 2014). In human care, H. gigas have produced as many as 63 eggs in a single
10-month period (Z.J.L. personal observation).

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Aquatic Snakes

Taxonomic History
Both Hydrodynastes species have a tumultuous taxonomic history. Hermans Wa-
tersnake was described by Herman in 1804 as Coluber bicinctus. A type specimen was not
denoted, and the initial description was based on specimens that were lost. Wagler and Spix
(1824) described Elaps schranki from Amazonia, which ultimately was determined to be Her-
man’s C. bicinctus. Schlegel (1837) transferred to E. schranki to Xenodon. Fitizinger (1843) de-
termined X. schranki was worthy of generic recognition and described and placed Herman’s
Watersnakes into Hydrodynastes. When Dumeril, Bibron, and Dumeril (1854) described the
False Water Cobra, they initially placed the species in Xenodon as X. gigas. Jan (1863) placed
both species in Lejosophis, a genera that contained several obscure South American snakes.
Cope (1854) described Cyclagaras to house South America’s Water Cobras, and removed
them from Lejosophis. Over the next century, Hermans Watersnake would be split from False
Water Cobras two more times, and placed in Urotheca (Dunn, 1894) and then the monotypic
genera Dugandia (Dunn, 1944). Hoge (1958) determined that the then D. bicinctus should
not be recognized due to name priority, and moved D. bicinctus back into Hydrodynastes.
In 1966 Hoge somewhat unceremoniously moved L. gigas to Hydrodynastes and de-
scribed Hydrodynastes bicinctus schultzi. Hydrodynastes bicinctus subspecies were differen-
tiated by Hoge (1966) based on the extent of the postocular stripe, ventral scale patterning,
and distribution. Hydrodynastes b. bicinctus was limited to Amazonia, while H. b. schultzi oc-
curred in the Cerrado. Murta-Fonesca (2015) demonstrated morphologically that characters
overlapped extensively between both Amazonia and Cerrado populations, and relegated H.
b. schultzi to be a single nominate taxon. Carvalho et al. (2023) supported these results with
molecular phylogenetic data.
Hydrodynastes gigas was moved back to Cyclagaras by Peters and Orejas-Miranda in
1970. At the time, apical pits were thought to hold significant taxonomic validity. Hydrodyn-
astes bicinctus lacked pits, while H. gigas possessed them. Later work determined that apical
pits were not taxonomically significant for certain groups of snakes, leading Dowling and
Gibson (1970) to place C. gigas back into Hydrodynastes. By 1970, both H. bicinctus and H.
gigas were in Hydrodynastes where they have remained until today. Hydrodynastes melanogi-
gas was described from the Tocantins River basin in East-Central Brazil in the early 2000’s by
Franco et al. (2007). The only characteristic used to differentiate H. melanogigas from H. gigas
was the extreme melanism observed in the taxon. Carvalho et al. (2020) dissolved this taxon
by showing that meristically and genetically there was nothing that could be used to differen-
tiate H. bicinctus from H. gigas other than coloration. Carvalho et al. (2023) determined that
though both species have rather extensive ranges over South America, they are closely related
to each other, evolved in the mid-Pleistocene, and display little population variation.

Biogeography

Both hydrodynastinins occur in South America, and in certain portions of their

range are syntopic. Hydrodynastes bicinctus prefers more forested environments compared to
its more ecologically cosmopolitan relative. As currently understood, H. bicinctus occurs in
riverine habitats and riparian corridors in central, eastern, and northern Amazonia in Brazil,
Venezuela, French Guiana, and Guyana. Hydrodynastes bicinctus has also been recorded in
the central and southern portions of the Cerrado and along the junction of the Atlantic Coast
Forest and Cerrado in Brazil.

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 20. Neotropical River Snakes

Hydrodynastes gigas has a larger distribution than H. bicinctus. Hydrodynastes bicinctus

and H. gigas occur syntopically in central and eastern Amazonia, a small portion of the Pan-
tanal (Fischer et al. 2018) as well as central and southern portions of the Cerrado (Noguiera
et al. 2019). Hydrodynastes gigas also occur in the Chaco of northeastern Argentina, eastern
Bolivia, and western Paraguay, the Caatinga of Northeastern Brazil, and throughout the Bra-
zilian Pantanal wetlands. Across their respective distributions, both Hydrodynastes species
are strongly associated with lotic environments (Giraudo et al. 2014; Carvalho et al. 2023;
Loughman 2023). Hydrodynastes bicinctus appears to be more aquatic than H. gigas and are
rarely encountered outside of water. While encounters with H. gigas have occurred outside of
water, aquatic habitats are almost always nearby. Hydrodynastes gigas utilize a wider array of
aquatic habitats and are frequently encountered in lentic as well as lotic systems.
Carvalho et al. (2023) investigated the phylogeography of both Hydrodynastes species.
Both are relatively young species and diverged from each other within the past 1.7 million
years in the Pleistocene. Genetic evidence points towards Hydrodynastes originating in the
Tocantins River of east-central Brazil roughly 1.5 mya. By the last glacial maxima, both spe-
cies had likely reached all watersheds and habitats they currently occur in. Genetic evidence
suggests that H. gigas populations have declined slowly over the last 20 thousand years; H.
bicinctus populations have remained stable.
Often animals with distributions strongly associated with river corridors have genetic
divergence and population structure allied to specific basins and watersheds. Given their
extensive range, one would expect H. bicinctus and H. gigas to have haplotypes associated
with specific river systems. This, however, is not the case. Both Hydrodynastes species exhibit
little to no population variation across their rather extensive range. Carvalho et al. (2023) at-
tributed a lack of genetic distinctiveness to life history and ecological attributes. Both species
are large and capable of expansive movements over their lifetimes. Given their generalized
feeding habits, emigrating populations likely have little problem finding habitat, refugia, or
prey. Vicariance is not likely to occur under these conditions, explaining the lack of popula-
tion distinctiveness observed in Hydrodynastes populations.

Conservation
Hydrodynastes bicinctus and H. gigas appear to be stable across their expansive
ranges. The International Union for Conservation of Nature lists both species as Least Con-
cern. Hydrodynastes gigas is a CITES II species, which eliminates its exportation from any
country of origin without an import permit. Both species currently occur in herpetoculture.
Hydrodynastes bicinctus is rarely maintained, though a stable captive H. gigas population
exists in both North America and Europe eliminating the need for importation (Loughman
2020, 2023). Habitat destruction is the most pressing concern for H. bicinctus and H. gigas.
Pollution, degradation, and destruction of riparian and riverine habitats, as well as wetlands,
eliminate populations of both species at a local level. Given their cantankerous nature when
disturbed, H. gigas is often killed on-site when encountered by humans. Throughout its dis-
tribution H. gigas is referred to as Nacanina which translates as Evil River Snake in Spanish
and Portuguese respectively. Climate change will likely impact both species negatively. Ex-
tensive drought conditions followed by expansive fires have become much more common in
the Chaco, Pantanal, and Amazonia over the past two decades. Destruction of habitat by fires
likely displaces Hydrodynastes populations.

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Aquatic Snakes

Herrmann’s Water Snake

Hydrodynastes bicinctus Herrmann, 1804
Distribution and Habitat. Guiana, Suriname, French
Guiana, Brazil (Amazonas, Rondonia, Pará, Maranhão,
Goiás, Mato Grosso do Sul, São Paulo, Amapa, Tocantins),
Colombia, Venezuela. Type locality: None given. Neotype
locality: municipality of Novo Progresso, about 240m ASL,
state of Pará, Brazil. The species is polytypic with two sub-
Figure 20.1. The distribution of Hy- species. H. b. bicinctus: Colombia, Venezuela, Guianas, Bra-
drodynastes bicinctus. zil (Amazonas). Type locality: “none given; Rio Japurá, Bra-
zil” (fide Kornacker 1999); H. b. schultzi: Brazil (São Paulo,
Goias: HR 33: 226, Tocantins, Mato Grosso do Sul) .

Figure 20.2. Hydrodynastes bicinctus.

Photography by William W. Lamar.

False Water Cobra

Hydrodynastes gigas (Duméril et al, 1854)
Distribution and Habitat. French Guiana (along the
coast), Suriname, southern Brazil, Piauí, São Paulo, Tocan-
tins, Minas Gerais, eastern Bolivia, Paraguay, N Argentina
(Misiones, Corrientes, Entre Rios, Santa Fe, Chaco, For-
mosa), Venezuela, Colombia, Guyana, possibly Peru. Type
locality: Corrientes, Argentina. A polytypic species with
Figure 20.3. The distribution of Hy- two subspecies. H. g. gigas, and H. g. melanogigas: Tocan-
drodynastes gigas tins, Brazil. Type locality: Municipality of Palmas State of
Tocantins, Brazil. Habitat ranges from wet savannas and
grasslands to subtropical dry forests, gallery forests, shrub-
lands and a wide variety of wetlands (rivers, streams, marshes, swamps).
Natural History. The diet reported by López and Giraudo (2004) included, amphibians

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 21. Neotropical Fishing Snakes

(30%), colubrid snakes

(26%), fish (22%) and
mammals (22%). One
specimen contained
fish eggs, and repre-
sentatives of two fish
families were identified,
Synbranchidae and Cal-
lichthyidae. Two adults
regurgitated four swamp
eels (Synbranchus mar-
moratus). The shape and
behavior of swamp eels
are similar to those of
aquatic snakes. Swamp
eels breath air and may
leave the water to move
overland.
Venom. Hydrodyn-
astes gigas is rear-fanged
snake with poorly
known venom. An adult
male human was bitten
on the fourth digit of the
left hand and the snake
was attached to the
digit for approximate-
ly 30 seconds. Within
5 minutes intense local
pain developed, and at
4 hours post bite the en-
tire dorsal aspect of the
hand was significantly Figure 20.4. Hydrodynastes gigas photographed near Filadelfia, Para-
edematous. The local ef-guay. Photography by Mike Pingleton.
fects progressed and in-
volved the entire forearm, and the local pain referred to the axillary region. Mild paresthesia
and local blanching (“pallor”) were noted in the affected digit but resolved within seven days.
Thus, moderate localized symptoms may result from the bite of a juvenile H. gigas.

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326
Aquatic Snakes

21. Neotropical Fishing Snakes

The genera Helicops, Hydrops, and Pseudoeryx form a widely distributed clade in the
Amazon basin. They have small dorsal eyes and narrow ventral scales and are found in many
aquatic habitats where they feed on fish including synbranchid eels. Species that reproduce
by oviparous and viviparous modes are present in this clade as well as species suspected of
being facultatively viviparous.
Schöneberg and Köhler (2021) provide distributions and identification diagnoses for all
the species of Helicops. Helicops contains 20 species and more are likely to be described. Heli-
cops angulatus probably contains multipe cryptic species.
Neotropical water snakes in the genus Hydrops Wagler, 1830 (Serpentes, Dipsadidae,
Hydropsini) are distributed across the northern two-thirds of South America and have been
reported from Colom­bia, Venezuela, Trinidad (not present on Tobago) Guyana, Suriname,
French Guiana, Ecuador, Peru, Brazil, Bolivia, Paraguay, and Argentina. Three species are
currently recognized: Hydrops caesurus, Hydrops martii, and Hydrops triangularis.
The eel snakes of the genus Pseudoeryx are widespread in South America east of the An-
des. They seem to prefer shallow water habitats where the search for synbranchid eels. Two
species are currently recognized with one polytypic species (P. plicatilis).

Water Mapepire
Helicops angulatus (Linnaeus, 1758)
Distribution and Habitat. A widespread species: Vene-
zuela and Trinidad, Colombia, Brazil, Bolivia, Peru, Ecuador,
French Guiana. It is almost exclusively aquatic, using shal-
low water habitats (ponds, rivers or swamps, canals, flooded
Figure 21.1. The distribution of fields (Murphy, 1997; Martins and Olivera, 1998; Ford and
Helicops angulatus Ford, 2002). It is mostly nocturnal. Henderson et al. (1976),
found activity starts about 20 minutes after dark. Helicops an-
gulatus seems to show site fidelity at least in the short term.
The size of the area in which H. angulatus utilizes in a period of about three days has been
found to be approximately 30 square meters squared (Henderson et al., 1976).
Natural History. It forages for fish at night but has also been observed foraging during
the day (JCM observation, Martins and Olivera, 1998). Prey includes tadpoles, frogs, fish, and
aquatic lizards. I (JCM) observed young trailing each other in a flooded ditch at night – the
snakes in the lead may startle prey into moving and allow the individuals behind to discover
the prey more easily. Strüssmann et al. (2013) report earthworms as prey and reports H.
angulatus feeds on the giant earthworms of the genus Rhinodrillus.
Reproduction. On Trinidad and at scattered locations in Brazil it appears to be facul-

On the previous page is Helicops angulatus Photography by JCM.

327
 21. Neotropical Fishing Snakes

Figure 21.2. Helicops angulatus

Photography by Photography
by Luis Alejandro Rodriguez J.

tatively viviparous (Braz et al. 2016; Murphy et al. 2020), that is, they can give birth to live
young as well as lay eggs. Eggs and young are likely produced year-round (Martins and Oli-
vera, 1998; Ford and Ford, 2002; Murphy et al. 2020). Clutch and litter size ranges from 2-20.
Helicops angulatus does not appear to be territorial (Ford and Ford, 2002).

Apiaka Watersnake
Helicops apiaka Kawashita-Ribeiro et al, 2013
Distribution and Habitat. Known from southern Pará and
Mato Grosso State, Brazil: Alta Floresta, Guaranta do Norte,
Paranaíta, and Peixoto de Azevedo municipalities. These
localities are situated in the southern Amazon Basin along
rivers in the drainages of the Teles Pires River, all are trib-
utaries of the Tapajos River. Type locality: Teles Pires River.
municipality of Paranaíta, Mato Grosso, Brazil.
Figure 21.3. The distribution of Natural History. Sympatric with H. angulatus and H. pol-
Helicops apiaka. ylepis. It is nocturnal in small and large streams and flooded
areas. It is nocturnal and frequently observed at the type lo-
cality, in small streams and major branches of the Teles Pires River. It is active, from 1830

328
Aquatic Snakes

to 2330 h, along the margins

of the Teles Pires River, or
partially submerged in small
Figure 21.4. Helicops apiaka.
Photography by Ricardo Al-
exandre Kawashita-Ribeiro.

streams and flooded areas.

When handled, they may at-
tempt to bite (Kawashita-Ri-
berio et al. 2013).

Fire Watersnake
Helicops boitata Moraes-da-Silva et al, 2019
Distribution and Habitat. Mato Grosso, Brazil. Known
only from the type locality- the Municipality of Poconé,
Mato Grosso state, Transpantaneira Road, municipality of
Figure 21.5. The distribution Poconé, Mato Grosso state, Brazil Brazil at the northern
of Helicops boitata.
limit of the Brazilian Pantanal.
Natural History. Known only from the holotype.
It has been found in mid-afternoon and may be diur-
nal. The holotype was found in a flooded area at the
edge of an unpaved road in the Panatal. The location
corresponds to a tributary of the Bento Gomes River..

Rough Tail Watersnake

Figure 21.6. The distribution Helicops carinicaudus (Wied-Neuwied, 1825)
of Helicops carinicaudus.
Distribution and Habitat. The distribution extends
from the estuary of the Rio de La Plata along the
shoreline of Brazil to the province of Pernambuco.
The name Helicops baliogaster Cope 1885 was listed as a synonym of Helicops carini-
caudas by Boulenger 1886.
Natural History. Active during the day and at night. It is often found along the Serra do
Mar Range. Diet includes fish and amphibians. The reproductive mode is viviparous with a
vestigial shell membrane. Reproduction is seasonal with vitellogenesis occurring from Sep-
tember to December, embryos present from November to March and young present from
the end of the rainy season to the beginning of the dry season (Marques 1998, Nogueira
and Marques 1998).

329
 21. Neotropical Fishing Snakes

Figure 21.7 Helicops carinicaudus from the Guandu River, City of Seropedica,
State of Rio de Janeiro. Photography by Adam Carvalho.
Cope’s Watersnake
Helicops cyclops Cope, 1869
Known only from the type specimen. Type locality Bahia,
Brazil. It has a remarkably short snout and head compared to
H. angulatus. Murphy et al. (2020) removed it from the syn-
onymy of H. angulatus.
Figure 21.8. The distribution of
Helicops danieli .

Daniel’s Watersnake
Helicops danieli Amaral, 1938
Distribution and Habitat. Shorelines of large swamps and
lakes, and mangroves in Colombia west of the Andes. There is
a report from the lowland in the east near the Brazilian border
which may be in error. Type locality: Colombia, Santander,
Carare.
Figure 21.9. The distribution of
Natural History. Sea level to 1500 meters ASL. It is abun-
Helicops gomesi. dant in the Magdalena River basin. Activity is nocturnal and it
uses aquatic plants as a refuge. Young feed on tadpoles; switch
to fish with age. Viviparous.

Amaral’s Keelback
Helicops gomesi Amaral, 1921
Distribution and Habitat. Streams and rivers in the vicinity of Sao Paulo Province to the

330
Aquatic Snakes

Figure 21.10. Helicops gomesi.

From Amaral, 1926.

Mato Grosso do Sul and Gorias, Brazil. Type locality: Costa Pinto Station, Railway Sorocaba,
São Paulo, Brazil.
Natural History. Occurs under rocks along the edges of streams. Oviparous, a clutch of
three eggs has been reported; originally reported as vivipa-
rous, may be facultatively viviparous. Braz (2020) found a
three-barbeled catfish, Pimelodella sp. (Heptapteridae) in a
female that also contained 14 pre-ovulatory vitellogenic fol-
licles on 5 November. This suggests the snake reproduces in
the wet-warm season (November-December).

Hagmann’s Keelback
Figure 21.11. The distribution of
Helicops hagmanni Roux, 1910
Helicops hagmanni.
Distribution and Habitat. It occurs from the Amazon
estuary to the Brazilian province of Amazonas and the Venezuelan province of Amazonas.
There is a single record from northern Rondônia, Brazil. Type locality: Santarem, Brazil.
Natural History. The diet appears to be composed exclusively of fish and it forages in
the medium and upper strata of the water column in the lentic zone (de Carvalho Teixeira,
2017). de Lima Moraes and Fraga (2015) found an adult perched about two meters above the
ground, on the vegetation adjacent to a small first order stream, in a sunny clearing.
Sturaro and Gomes (2008) observed an adult Helicops hagmanni 5`16 mm SVL, weight
162 g) preying on the catfish, Rhamdia muelleri (215 mm total length and 70 g) in the field at
2300 h. The water was 20 cm deep. The snake seized the fish biting the prey’s posterior body
between the anal and post-anal fins, maybe to avoid the fish’s pectoral spines. The snake held
the prey with anterior, horizontal coils, similar to what has been observed in other colubrid
snakes. Subsequently, the snake extended constriction
until the opercula area of the catfish. Thereafter, the snake examined the prey’s body and
swallowed it, starting ingestion head first.

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 21. Neotropical Fishing Snakes

Figure 21.12. The distribution of

Helicops infrataeniatus.

Figure 21.13. Helicops hagmanni

(A) From Presidente Figueiredo,
Amazonas, Brazil. Photography
by Jairmaia (B) from Beruri,
Amazonas, Brazil. Photography
by Marco Aurelio de Sena.

Jan’s Keelback
Helicops infrataeniatus Jan, 1865
Distribution and Habitat. Southern Brazil, Uruguay, Argentina, possibly Paraguay. Type
locality was given as “Mexico” in error; other type material from Brazil.
Natural History. The diet is composed of fish (70%) and frogs (30%). Foraging behavior
likely occurs at the water’s edge and at multiple depths in the water column. The reproduc-
tive cycle is longer than other oviparous colubrids from subtropical and temperate zones of
Brazil. Vitellogenesis occurred from September to February. Oviductal embryos were found
from September to May. Births took place in September, January, February, and March, and
occurred in two peaks. Litter size ranged from 5 to 22 offspring. The differences between the
number of vitellogenic follicles and embryos, and between embryos and newborns were not
significant, thus practically all follicles were generally converted into embryos and embryos
into neonates.

Spotted Keelback
Helicops leopardinus Schlegel, 1837
Distribution and Habitat. The distribution is peripheral to the central Amazon basin
and the species is known from the northwestern Argentina to the Amazon estuary and from

332
Aquatic Snakes

Ecuador to the Brazilian state Bahia. Wetlands of Guiana, Suri-

name, French Guiana, Brazil, Bolivia, Paraguay, Ecuador, Peru,
and possibly Colombia. It is abundant in the Pantanal. Type
locality is unknown.
Natural History. Diet includes mostly fish (and the fish are
often Gymnotiformes) and about 30% anurans, most of which
are hylids. The mean litter size is 6.6 and a range of 4–11. The
reproductive cycle is seasonal, with parturition occurring late
in the wet season (simultaneous with the flooding of the Par-
Figure 21.14. The distribution
of Helicops leopardinus.

Figure 21.15. Helicops leop-

ardinus. (A) Photography by
Wolfgang Wuster (B) from
Gral Paz, Corrientes, Argen-
tina Alfredo Sabaliauskas.

aguay River). Young snakes

disperse on floating vegetation
and the floating vegetation
may be a critical habitat for H.
leopardinus, and management
plans for their maintenance
need to be developed (Ávila
et al. 2006). Ruiz-Garcia et al.
(2020) found the Crab-eating
Fox is a predator on this snake.

Olive Keelback
Helicops modestus
Günther, 1861
Distribution and Habitat. Distributed in the Brazilian province Minas Gerais, Distrito
Federal, Goias, Bahia, and Pará, Brazil
Natural History. A diurnal and nocturnal snake with a
diet of fish and small anurans. A juvenile H. modestus preyed
on a small leiuperid frog, Physalaemus cuvieri.
Live fish fed to modestus in captivity were taken while
the snake was submerged (Oliviera et al. 2016). After
Figure 21.16. The distribution tongue-flicking the snake faced and approached the fish.
of Helicops modestus. Using repeated lateral undulations, resulting in irregular si-
nusoidal flexions that pushed the body forward, typical of
pre-striking behavior. Strikes were followed by immediate bites on the fish’s head, mid-re-

333
 21. Neotropical Fishing Snakes

gions or tail. Prey was sometimes pinned against the terrarium wall. Ingestion began head-
first, with the fish either dead or alive but always immobilized. The mean swallowing time
was 16 minutes. The Duvernoy’s glands are connected to the ungrooved, blade-like maxillary
fangs by a vestibule from which the secretion is drained.
Observations of prey-handling suggests H. modestus strikes and holds fish in its mouth
while repeatedly carrying out bilateral raking motions with both maxillae. Fish are swallowed
alive but are immobilized, suggesting that the venom is associated with the immobilization of
the fish prey Oliveira et al. (2016).
One female gave birth to 20 young in March. Another 590 mm (SVL) female gave birth
on 31 January to a litter of 24 neonates. The relative clutch mass (RCM) was 0.34, a high value
but like the values reported for other Helicops. Neonates have SVLs 120-160 mm, and weigh
1.1-2.5 g. The reproductive cycle is seasonal with vitellogenesis occurring from September

Figure 21.17. Helicops modestus

Photography by Wolfgang Wuster.

to December, embryos from November to March and recruitment from the end of the rainy
season to the beginning of the dry season (Maia et al. 2015). Da Costa et al. (2021) observed
the leaf cutter ant Atta opacieps attack and kill one of these
snake crawling across a trail.

Quenya Keelback
Helicops nentur Costa, et al., 2016

Figure 21.18. The distribution of Distribution and Habitat. In southeastern Brazil it

Helicops nentur. is known from the eastern half of Minas Gerais, Brazil. It
inhabits the Bahia Interior Forest ecoregion (Tropical and
Subtropical Moist Broadleaf Forests biome), its border with the Cerrado ecoregion (Tropical
and Subtropical Grasslands, Savannas, and Shrublands biome), and the ecotone between the
Atlantic Dry Forest and the Caatinga ecoregions (Tropical and Subtropical Dry Broadleaf

334
Aquatic Snakes

Forests biome and Deserts and Xeric Shrublands biome). The type locality is a swamp in
Fazenda Papa Capim São José da Safira, in the state of Minas Gerais, Brazil.
Natural History. Inhabits lentic water bodies such as swamps. The holotype was collect-
ed while active at 0300 h, partially submerged. After collection, it gave birth to two stillborn
specimens not measured nor preserved. One specimen has an umbilical scar on ventrals
91–95. Date of collection of these two specimens suggest that reproduction in H. nentur may
occur during the rainy season (October–March) (Costa et al. 2016).

Shreve’s Keelback
Helicops pastazae Shreve, 1934
Distribution and Habitat. Ranges from eastern Vene-
zuela to eastern Ecuador – unknown from Colombia. The
type locality is on the Río Pastaza, between Canelos and the
Figure 21.19. The distribution Marañon River, Ecuador.
of Helicops pastazae. Natural History. The diet includes fish and anurans. Gar-
cia-Cobos and Gomez-Sanchez (2019) collected a clutch of 11
eggs on 20 December (early dry season). The eggs were buried under a substratum composed
of sand, small rocks and decomposing material located under a large rock separated about
two meters from the edge of the river. After 46 days of incubation, on 4 February, the eggs
started to hatch, and hatching continued until 7 February.
García-Cobos et al. (2020) studied the reproductive cycle of this species in a montane
river in the Andean foothills of Colombia. They sampled 212 snakes from 2013 to 2017 and
classified the monthly reproductive stages of the adult snakes. Females showed a marked
seasonal reproductive cycle, with secondary vitellogenic individuals mainly clustered during
the high precipitation months, ovigerous stages found throughout low precipitation months,
and egg-laying occurred at times of low precipitation at the end of the wet season to the
early dry seasons. Males produced spermatozoa throughout the year indicating year-round
reproduction at a population level. They found sexual dimorphism in tail length, with both
juvenile and adult males showing significantly longer tails than females, while adult females
were larger than males and showed greater midbody width, head length, and head width. The
reproductive activity of H. pastazae is highly seasonal in females, driven by regional rainfall
patterns, whereas males are reproductive thorough the year, being a permanent resource for
females. Defense behavior of this species includes striking with the mouth open and flatten-
ing the head and body. Predators include other snakes, birds, and carnivorous mammals.

Peter’s Water Snake

Helicops petersi Rossman, 1976
Distribution and Habitat. is known only from a very small
area in the Ecuadorian province Napo. The type locality is on the
east bank of Rio Misahualli, 1 mile northeast of Tena,
Napo, Ecuador.
Figure 21.20. The distribu-
Identification. Helicops petersi can be distinguished from
tion of Helicops petersi.
all its congeners except H. pastazae by the combination of 21–23
dorsal scale rows at midbody, reduced to 16 rows anterior to cloaca, and 135–150 ventrals.
Natural History. Arteaga (2021) states that it inhabits lentic waters in areas of evergreen

335
 21. Neotropical Fishing Snakes

Figure 21.21. Helicops pastazae. Photography by Mario Yanez Munoz.

lowland and foothill forest. It occurs in tributaries of large lakes and rivers, including small
streams and fast-flowing whitewater rivers, but it is also present in flooded habitats altered
by humans. This species occurs in open and forested areas with various degrees of human
disturbance. Peter’s Watersnake is nocturnal but can occasionally be seen moving during the
day. During the day, individuals have been found buried in the sand besides rivers. They are
sit-and-wait predators that feed on fish. Individuals have been captured in fish nets. Faced
with a potential threat, individuals make an S-coil, flatten the body and head dorsoventrally,
open the mouth, and strike. When manipulated, they rotate the body, bite, and produce cloa-
cal discharges. This species is probably oviparous. One female contained 12 small ova.

Ghost Water Snake

Helicops phantasma Moraes-da-Silva et al, 2021
Distribution and Habitat. The species is only known from
the Tocantins-Araguaia River Basin in the provinces Toncan-
Figure 21.23. Helicops phantasma from Barra do Bugres-MT,
eastern of Mato Grosso.
Figure 21.22. The distribution Photography by Ricardo Alexandre Kawashita-Ribeiro.
Helicops phantasma.

336
Aquatic Snakes

tins, Mato Grosso and Maranhão in northern Brazil. The type locality is Barra do Garças,
Mato Grosso, Brazil, at 840 m ASL.

Norman’s Keelback
Helicops polylepis Günther, 1861
Distribution and Habitat. Ranges from Brazil (Amazonas
and Pará), Colombia, Peru, and Bolivia. The type locality is the
upper Amazon.
Natural History. The diet includes Synbranchus eeland
Figure 21,24. The distribution of
Helicops polylepis.

other fishes (Flores et al. 2010;

de Carvalho Teixeira C et al.
2017)), Females have a larger
body size than males, con-
sistent with most viviparous
snakes (Carmago et al. 2021)

Hog’s Keelback
Helicops scalaris Jan,
1865
Distribution and Habitat.
Known only from a small area
in the northern border area
between Colombia and Ven-
ezuela, western and northern
of Lake Maracaibo in Venezu-
ela. The type locality is Ven-
ezuela.
Figure 21.25. Helicops polylepis. The top image is an adult, bottom
is a juvenile.
Photography by (top) Mike Pingleton; (bottom) Paul Freed.

Figure 21.26. The distribution

of Helicops scalaris.

Figure 21.27. Helicops scalaris.

Photography by Raul Carmo-
na.

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 21. Neotropical Fishing Snakes

Cobra-D’água
Helicops tapajonicus Da Frota, 2005
Distribution and Habitat. Shallow water with muddy sub-
strates at two localities on the Tapajos River close to its con-
fluence with the Amazon in the Brazilian state Pará. The type
locality is the community of Parauá, on the right bank of the
Tapajós river, in the municipality of Santarém, Pará, Brazil.
Figure 21.28. The distribution of Natural History. The specimens were found at night, near
Helicops tapajonicus. the mouth of the streams on the Tapajós River. At the time of
collection, the specimens were in the water, in a muddy area
with grasses, fallen branches and roots (Da Frota 2005)

Equatorial Keelback
Helicops trivittatus (Gray, 1849)
Distribution and Habitat. Forested streams in eastern
Pará, in northern Brazil. The type locality was given as “India
?” in error.
Natural History. Active day and night, in streams of for-
ested areas or near residences of riverside residents. Diet: fish
(Acaronia nasa, Plagioscion squamosissimus). Reproduction:
female with 26 follicles in advanced vitellogenesis in March;
Figure 21.29. The distribution of
and oviducts with marks of recent spawning in August and
Helicops trivittatus.
November; parturition in March; litters of seven to nine off-
spring.

Figure 21.30. EquatoriaL Keel-

back, Helicops trivittatus. From
Belém, Pará, Brazil Photogra-
phy by Gisele Cassunde.

Peruvian Keelback
Helicops yacu Rossman,
1975
This is the most poorly
known member of the genus. The type locality is Moropon, Departamento de Loreto, Peru.
Natural History. Unknown.

Figure 21.31. The distribution

of Helicops yacu.

338
Aquatic Snakes

Figure 21.32. This is an undescribed Helicops, from the Rio Morpon in Peru. It has been con-
fused with Helicops leopardinus. Photography by William W. Lamar.

339
 21. Neotropical Fishing Snakes

Southern Water Snake

Hydrops caesurus Scrocchi et al. 2005
Distribution and Habitat. Northeast Argentina and Para-
guay. The distribution is disjunct from other members of the
Figure 21.33. The distribution genus. It inhabits temperate and subtropical latitudes from the
of Hydrops caesurus. Paraná and Plata River basins, from Pantanal in Mato Grosso
do Sul, Brazil, through Paraguay and Paraná rivers Type local-
ity: Departamento Itapúa, Isla Paloma, Canal de los Jesuitas, Paraguay.
Natural History. Information on this species is limited. Clutches of nine and 12 eggs
have been reported (Álvarez et al., 2003, Etchepare et al. 2012). The clutch of 12 had calcar-
eous shells, they were found on 15
November Puerto Tala, Isla Apipé
Grande, Corrientes province, Ar-
gentina. The nest was on the bor-
der of the embankment of a rural
road that crosses a marsh. An em-
bryo examined was in Zehr stage
26. It measured 40.7 mm body
length (from the snout to the tail
tip).
Figure 21.34. Hydrops caesurus Pho-
tography by (A-B) Stephen Barten,
(C) Ricardo Alexandre Kawashita-Ri-
beiro.

Triangle Water Snake

Hydrops triangularis Wagler,
1824
Distribution and Habitat.
Aquatic habitats of tropical and
subtropical moist broadleaf forest,

Figure 21.35. The distribution

of Hydrops triangularis.

though it also occurs in tropical

and subtropical grasslands, savan-
nas and shrublands in the Amazo-

340
Aquatic Snakes

nas, Orinoco and Guyana drainages includes Bolivia, Vene-

zuela, Guyana, Surinam, French Guiana, Trinidad, E Peru,
Ecuador, Brazil, Colombia, and Argentina. Type locality: Ega
(= Tefé) Lago Tefé, at confluence with Rio Amazon, Brazil.
Natural History. An aquatic specialist. It is relative-
ly common in streams found in open forests and savannas
(Rivero- Blanco and Dixon 1978; Hoogmoed 1982; Murphy
1997) and they often use shady, lentic environments. Activ-
ity usually after dark when fish, especially synbranchid fish
(which have eel-body forms) are active. Females produce
clutches of 8- 34 eggs.

Amazon Water Snake

Hydrops martii (Wagler,
1824)
Distribution and Hab-
itat. Aquatic habitats in
Figure 21.37. The distribution of tropical and subtropical
Hydrops martii. moist broadleaf forest. East
Figure 21.36. Hydrops triangu- of the Andes in the Amazon
laris. Photography by (A) Nue- basin from Colombia and eastern Peru to Maranhao, Brazil.
vo, Peru William W. Lamar; (B)
Known from tributaries of Rio Amazonas, the Rio Ucayali
Trinidad, Renoir Auguste.
and Rio Marañon, Peru. Roze (1957) reported a specimen
from Rio Cairary, near Jurupary waterfall, on the boundary
between Colombia and Brazil. Type locality: Provincia Maranhao, Rio Itapicuru, Brazil.
Natural History. Pin-
heiro et al. (2020) found the
stomach contents of the pira-
nha, Serrasalmus rhombeus,
containing a partly digested
Hydrops martii. A hatchling
found in November. De Car-
valho Teixeira et al. (2017)
reported active specimens
during the day, in a stream
traversing primary forest.
They considered the snake
primarily aquatic, and suggest
it will eventually be found ac-
tive on the ground, foraging
during day and night. Dietary
information was reported
by Cunha and Nascimento
(1993) as fish (Callichthys cal-
lichthys, Pimelodella crista-
ta, Hoplias malabaricus and Figure 21.38. Hydrops martii from Momo, Peru. Photography by
Erythrinus erythrinus) and William W. Lamar.

341
 21. Neotropical Fishing Snakes

Bartlett and Bartlett (2003), reported Synbranchidae in its diet, confirming the ichthyopha-
gous habit. Reproduction is poorly known. One offspring was found in November in Ecuador
(Dixon and Soini, 1977).

Amazonian Eel Snake

Pseudoeryx plicatilis (Linnaeus, 1758)
Distribution and Habitat. It occurs east of the Andes in marshy areas and dense reedbeds
as well as in streams, ponds, and flooded areas in the northern region of the Amazonian
rainforest, including the Orinoco River basin and drainages of Guyana area. It occurs in the
Amazonas River basin and its major tributaries to the east
of the state of Pará and west of the state of Maranhão. It also
occurs along the Prata River basin (restrict to the Paraguay
River basin), in the states of Mato Grosso and Mato Grosso
do Sul in Brazil (in flooded areas of the Pantanal region),
to the east of Bolivia, north of Argentina, and southwestern
Figure 21.39 The distribution of Paraguay. Type locality: Ternataeis.
Pseudoeryx plicatilis. This is a polytypic species: Pseudoeryx plicatilis plicatilis
(Linnaeus 1758) occurs in Colombia, Venezuela, Ecuador,
Peru, Guyana, Suriname, French Guiana, Paraguay, N Argentina, Brazil. Pseudoeryx plicatilis
mimeticus Cope 1885 occurs in Bolivia.
Type locality: Bolivia, Rio Mamoré.
Natural History. The diet is com-
posed of fish with eel-like bodies Re-
productive mode is oviparous, and
available evidence suggests that females
remain with eggs throughout incuba-
[Link] viviparity maybe pres-
ent in this snake. Defensive behaviors
are shared with other water snakes and
include: dorso-ventral flattening, hid-
ing the head, and cloacal discharge. P.
plicatilis flattens the whole body mak-
ing the snake look bigger than it is, and
may intimidate visual predators, such
birds. Many snakes hide the head as
anti-predator mechanism. Pseudoeryx
may hide the head and tail at the same
time. Pseudoeryx also inverts the body
and knots the body (Cavalheri et al.
2021).

Figure 21.40. Pseudoeryx plicatilis Bottom photo shows

death feigning behavior Photography by Konrad Me-
bert.

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Aquatic Snakes

Maracaibo Eel Snake

Pseudoeryx relictualis Schargel et al, 2007
Distribution and Habitat. Irrigation canals, swamps,
flooded pastures of the Maracaibo Basin in northwest Ven-
ezuela. Known from three locations when it was described
in 2007.
Natural History. The holotype was found in an irriga-
tion channel that provides water from the Catatumbo River
to crops and pastures in the area. The channels have her-
Figure 21.41 The distribution of Pseu- baceous and aquatic vegetation, especially water hyacinths.
doeryx relictualis. The type locality is on a farm where patches of the original
vegetation, ombrofilous evergreen forest, persist. This area
undergoes periodic flooded by the Catatumbo River. The original forests in this region, have
been called “Humid Forests of the Catatumbo” and are currently considered critically endan-
gered. The specimen from Bobures, was collected in flooded pastures used for cattle farming
and where most of the original forest has been cleared. This locality is only about 100 m
from Lake Maracaibo. The specimen from Ensenada El Congo comes from a wildlife reserve
consisting of extensive swamps where the natural vegetation of flooded pastures and forests
remains almost pristine (Schargel et al. 2007).

Figure 21.42. Pseudoeryx relictualis.

Photography by Luis Alejandro Rodriguez J.

343
Aquatic Snakes

344
Aquatic Snakes

22. Neotropical Bush Snakes

Bailey (1967) first recognized the Tachymenini group. Today it contains 37 species.
However Zaher et al. (2019) found the morphologically diverse clade was only moderately
supported. It contains arboreal, terrestrial, fossorial, and aquatic species in seven genera:
Calamodontophis, Gomesophis, Pseudotomodon, Ptychophis, Tachymenis, Thamnodynastes,
Tomodon.
Calamodontophis contains two poorly known species from southern South America.
They appear to be terrestrial. Pseudotomodon – monotypic genus from western Argentina,
is not aquatic. Tachymenis has about six species distributed from Peru to Chile, that are not
aquatic. Tomodon are specialized slug and snail-eating species that are terrestrial and arboreal.
Thamnodynastes is composed of 20 small to medium size species with an enlarged pair
of fangs at the back of the maxillae. These fangs are grooved to transport venom into the bite
punctures. They are viviparous snakes, with elliptical pupils, single nasal scales (which may be
semi-divided). Some may be best regarded as semi-aquatic and terrestrial snakes, that forage
in or near the water for anurans. The semi-aquatic Gomesophis is the sister to the other mem-
bers of the clade (Vidal et al. 2010).
The two highlt aquatic Tachymenini species are both in monotypic genera. The most
recent discussion of these poorly known snakes is Gonzalez et al. (2014).

Brazilian Burrowing Snake

Gomesophis brasiliensis Gomes, 1918
Distribution and Habitat. Slow-flowing water, mud-
dy wetlands, and the margins of streams and ponds in
cold-water environments of Minas Gerais, Sao Paulo, Pa-
rana, Rio Grande do Sul, and Santa Catarina, Brazil. Type
locality: Pindamonhangaba, São Paulo, Brazil. Gomesophis
Figure 22.1. The distribution Gomeso- occurs in aquatic habitats. Two specimens were collect-
phis brasiliensis.
ed at night in an area that was originally ombrophylous
Mixed Forest (Brazilian pine forest). One collection site
was a pond with a muddy bottom and turbid water, 80 cm deep. Herbaceous aquatic macro-
phytes (emergent and floating) are often present. It often hides in burrows dug into the mud of
wetlands and banks of streams and ponds. (Gomes 1918, Franco 2000, Prudente and Brandão
1998; Lema 2002; França and Araújo 2007; Fortes et al. 2010).
Most of the distribution of G. brasiliensis is associated with the Plata Basin, the second
largest river basin in South America, The Plata Basin is threatened by hydroelectric impound-
ments. The species is restricted to elevations of 427–1235 m ASL, (mean = 808 m ASL.) with
high rainfall levels and a cool climate (Prudente 1993), although it can also be found at lower
elevation in Rio Grande do Sul with a cooler subtropical climate.
Natural History. Gomesophis are specialized earthworm-eaters, including the giant
earthworms (family Glossoscolecidae) (Marques 1996, Oliveira et al. 2003, Pereira et al. 2007,
Strussmann et al. 2013). Giant earthworms occur at many groundwater upwelling sites found
along riparian systems belonging to the Rio Verde sub-basin. It is sympatric, but not syntopic,
with Ptychophis flavovirgatus in most parts of its geographical range.

Previous Page. Thamnodynastes ramonriveroi . southwest Trinidad

Photography JCM.

345
 22. Neotropical Bush Snakes

Figure 22.2. Gomesophis brasiliensis from the Serra do Papagaio, Aiuruoca, MG, Brazil. Photog-
raphy by Frederico de Alcântara Menezes, Arthur Diesel Abegg, Bruno Rocha da Silva, Francisco
Luís Franco, Renato Neves Feio.
Fanged Water Snake
Ptychophis flavovirgatus Gomes, 1915
Distribution and Habitat. Lotic waters in the Brazilian
states of Minas Gerais, Paraná, Santa Catarina, and Rio Grande
do Sul. It uses crevices between stones in the water for refuge. It
has an elevational distribution between 316–1358 m ASL. Most
locations are above 800 m ASL, and the lower elevation locali-
ties (316– 687 m ASL) are in the State of Rio Grande do Sul, in
a seasonal-subtropical climate where vegetation can be open or
Figure 22.3. The distribution
Ptychophis flavovirgatus. Araucaria forests (Porto & Caramaschi 1988).
Natural History. Both sexes reach the sexual maturity at
a similar body size. The reproductive cycle of Ptychophis fla-
vovirgatus is long (October to June), vitellogenesis is seasonal (August and September). Six
females contained between 5 and 10 embryos in the oviducts. The recruitment probably
takes place in the winter and early spring (Scartozzoni and Marques 2004).

Thamnodynastes
Most members of this genus feed on anurans and are therefore likely associated with
wet habitats. However, many of these species are either terrestrial or are too poorly under-
stood to assess their use of aquatic habitats: Thamnodynastes are terrestrial and semi-aquat-
ic. They are viviparous, opisthoglyphous and have elliptical vertical pupils. The genus has
346
Aquatic Snakes

Figure 22.4. Ptychophis fla-

vovirgatus Bottom photo
death feigning behavior.
Photography by Pedro Ber-
nardo.

21 valid species, 11 of which

occur in Brazil.
Thamnodynastes may not
represent a monophyletic ge-
nus (Franco & Ferreira 2003;
Zaher et al. 2009; Vidal et al.
2010; Grazziotin et al. 2012;
Zaher et al. 2018), and Franco
et al. (2017)have already not-
ed the cryptic diversity asso-
ciated with its members. The
combination of taxonomic is-
sues and the absence of robust
diagnostic characters have
perpetuated the difficulty to
correctly assign individuals to
species. The Thamnodynastes
pallidus group represents an
exception to this pattern. Bai-
ley et al. (2005) defined the
group based on several scale
characters, body shape, col-
or, behavior, and hemipenial
morphology. The hemipenes
are characteristic of the group,
with no enlarged spines.

Chaco Bush Snake

Thamnodynastes
chaquensis Bergna and Álvarez, 1993
Distribution and Habitat. Wetlands including the Pantanal of Paraguay, Uruguay, Brazil,
Argentina. The type locality is Colonia Las Mercedes, San Fernando Department, Chaco,
Argentina. Bellini et al. (2014) found this semi-aquatic species inhabits dry forest (51%), wet-
lands (16%), wet savannas (14%) and marshes (10%) and (9%) suburban areas. Harrington
et al. (2018) found it is also partly arboreal. Akmentins et al (2010) reported a specimen
submerged in a small temporary pond in secondary montane forest made up mostly of alders
(Alnus acuminata).
Natural History. Diet contains hylid, microhylid and leptodactylid frogs as well as other
small vertebrates. Frogs compose about 70% of the diet Small frogs seem to be swallowed

347
 22. Neotropical Bush Snakes

with the posterior first (Bernarde et al.2000).

Argentina Bush Snake

Thamnodynastes hypoconia (Cope, 1860)
Distribution and Habitat. Stagnant and slow-mov-
Figure 22.5. The distribution ing wetlands of Brazil, Argentina, Paraguay, and Uruguay.
Thamnodynastes chaquensis. Habitats include wetlands surrounding large rivers. Ripar-

Figure 22.6. Thamnodynastes chaquensis Photography by Dan Riley.

ian situations in forests, and wetlands around farmland, urban and suburban areas. Type
locality: Buenos Aires, Argentina.
Natural History. The diet is composed mostly of am-
phibians (96%) and the occasional skink. Anuran families
making up the diet: Hylidae (34%), Leptodactylidae (16%),
Leiuperidae (11%), Bufonidae (5%), Cycloramphidae (3%),
Scincidae (3%), and unidentified amphibians (29%) (Bell-
ini et al. 2013).
Females have vitellogenic follicles from March to May,
and oviductal eggs from September to December indicat-
Figure 22.7. The distribution ing seasonal reproduction. Births occur in January, litter
Thamnodynastes hypoconia.
sizes reported range from 7-13. Neonates with umbilical
scars were found from January to March. Females may
only reproduce once every two years. Males have continuous reproductive cycle, with no

348
Aquatic Snakes

Figure 22.8. Thamnodynastes hypoco-

nia. Photography by Paul Freed.

significant difference in testicular volume between months (Bellini et al. 2013).

Snakes are found throughout the year, but encounter frequency was higher in spring
and summer (September to March) than in winter and autumn Encounters peak in March.
The daily activity maximum occurred from twilight to nightfall, with the greatest number of
encounters between 1800 and 2200 h (Bellini et al. 2013).

Mato Gross Bush Snake

Thamnodynastes lanei Bailey, Thomas, and da Silva, 2005
Distribution and Habitat. The larger streams and ri-
parian systems and their flood plains are the usual habitats
in Brazil (Mato Grosso do Sul), probably also in Argentina,
Paraguay, and northeast Bolivia. Type locality: Salobra, Mato
Groso, Brazil. Natural History. Poorly known. A semi-aquatic
snake that feeds on frogs of the genus Leptodactylus (Eversole
Figure 22.9 The distribution et al. 2016).
Thamnodynastesl lanei.

Linne’s Bush Snake

Thamnodynastes pallidus (Linnaeus, 1758)
Distribution and Habitat. Guyana, Suriname, French
Guiana, Brazil, Peru, Venezuela, Colombia, Bolivia, Ecuador.
Type locality: Indiis [in error]. They are in forests or at for-
est edges. Also known from old secondary growth. Wetlands,
riparian floodplains, swamps, floating islands of vegetation.
Natural History. Usually associated with frog choruses.
Figure 22.11. The distribution Diet is composed of mostly hylid frogs. Litters of 2-7 young.
Thamnodynastes pallidus Six near-term embryos measured 108 mm and 110 mm SVL
for two females and 115-173 mm for four males.

349
 22. Neotropical Bush Snakes

Figure 22.10. Thamnodynastes

lanei. Pedro Bernado.

Figure 22.12. Thamnodynas-

tes pallidus. Photography by
Dario Jose Alarcon Naforo.

350
Aquatic Snakes

Turimiquire Bush Snake

Thamnodynastes ramonriveroi
Manzanilla and Sánchez, 2005
Distribution and Habitat. Streams and wetlands of Ven-
ezuela (Turimiquire massiff, elevation 0-1750 m), Suriname,
Figure 22.13. The distribution Guyana, Brazil, and Trinidad. The type locality is Cerro La
Thamnodynastes ramonriveroi. Laguna, Turimiquire massif, Freites, Anzoátegui, Venezuela.
Natural History. On Trinidad often associated with
frog choruses, streams, and flooded ditches. Litters of four

Figure 22.14. Thamnodynastes ramonriverio, southern Trinidad. Photography by JCM.

351
 22. Neotropical Bush Snakes

and five embryos reported. Defense behavior includes flattening the first half of body and
when tormented they opened mouth very widely; death feigning occurred when struck with
a light stick (Manzanilla & Sánchez, 2005; personal observation
JCM).

Gunther’s Bush Snake

Thamnodynastes strigatus (Günther, 1858)
Distribution and Habitat. In lotic aquatic habitats, such as
Figure 22.15. The distribution of marshes in Argentina, Uruguay, Paraguay, and southern Bra-
Thamnodynastes strigatus.
zil. Type locality India (in error).
Natural History. Diet includes fish, frogs, and mammals.
Snakes ambush fish from the vegetation. They peering at fishes and use lingual luring to attract
them. It will also actively forage at the margins of bodies of water. Proenca et al. (2015) recov-
ered the remains of the fish Hoplias malabaricus from the stomach of this species. Reproduc-
tion. Four females (407-602 mm SVL) produced 12, 13, and 26 neonates. A fourth female had
22 oviductal eggs. Two of the litters were born in October. The neonates were 130-168 mm in
total length (average = 151 mm). The three litters had RCMs of 0.313, 0.609, and 0.309 (Braz
and Scartozzoni 2020).

Figure 22.16. Thamnodynastes cf. strigatus

Photography by Wolfgang Wüster.

On the next page. Erythrolamprus cf reginae from Peru. Photography by JCM.

352
Aquatic Snakes

353
 23. The Semi-Aquatic Painted Snakes

23. The Semi-Aquatic Painted Snakes

The Painted Snakes form a tribe level clade, the Xenodontini. Zaher et al. (2019) found
unambiguous support for a common ancestor shared by the genera Erythrolamprus, Ly-
gophis, Xenodon, and possibly Umbrivaga. The 80-plus species tend to be small (less than a
meter), feed on small vertebrates, are oviparous, and have continuous reproductive cycles
in tropical latitudes and seasonal reproduction in more temperate latitudes (Pizzatto et al.
2008). Members of the genus Erythrolamprus tend to have fewer eggs, but larger offspring
and Xenodon merremii have the largest clutches. Adult females tend to be larger than males,
but both sexes reach sexual maturity at the same size. Tail lengths and subcaudal counts in
many of these species are not sexually dimorphic.
I have not included species of Xenodon here because the evidence for semi-aquatic
habits is scarce, although Hoogmoed (1985) suggests they have valvular nares and note
they feed primarily on frogs. As these snakes become better known, they may be shown to
be semi-aquatic.
The Painted Snake name is derived from the bright colors found in many of them. Fre-
quently the bright coloration is restricted to the ventral surface. A high contrast red, orange,
or yellow venter with dark bands or stripes are found in many aquatic and semi-aquatic
dipsadids (Erythrolamprus almadensis, E. reginae, Sordellina, Helicops, Farancia), natricids
(Lithodytes, Nerodia, Smithophis), homalopsids (Enhydris, Homalophis, Homalopsis, Hypsis-
copus), and cylindrophiids (Cylindrophis).
Bright colors and high-contrast patterns, such as stripes or bands, on the ventral sur-
face of aquatic snakes, suggest that this pattern may be aposematic and useful for deterring
predators during encounters in aquatic environments (Batista et al. 2020). However, an al-
ternative hypothesis was proposed by Goldenberg et al. (2021). They found brightly colored
and patterned ventrals may have evolved for moving heat absorbed by soils to snakes when
they are in contact with the ground. Dry soil and rocks convey heat more rapidly than wet
soils. Thus, variation in soils’ thermal properties could be significant for species that live in
continuous and close contact with their substrate. In addition, melanins, a ubiquitous class
of multifunctional macromolecules, largely determine the brightness of the integument.
Melanins absorb and transform solar radiation into heat and have good electrically conduc-
tive properties. Thus, brightly colored ventral surfaces may be more than just aposematic.

Gunther’s Striped Snake

Lygophis anomalus (Günther, 1858)
Distribution and Habitat. Humid grasslands in Southern Brazil, Uruguay, Paraguay,
and northern Argentina. Type locality the Banks of the Pa-
raná.
Natural History. The diet is composed mostly of an-
urans, and occasionally reptiles. Prior to swallowing, prey
was oriented for headfirst ingestion. Sexual dimorphism in
head length, and diet composition was absent suggesting
that L. anomalus is opportunistic. Its feeding activity seems
to depend more on the season that on the day/night cycle
Figure 23.1 The distribution of (Panzera and Maneyro 2014). Females lay a clutch of about
Lygophis anomalus. 15 eggs in in December and January.

354
Aquatic Snakes

Figure 23.2 Lygophis anomalus

Photography by Alfredo Sabaliauskas.

Erythrolamprus
Fifty-five species of Erythrolamprus occur in the Antilles and on mainland South Amer-
ica from the slopes of the Andes to northern Brazil and the Chaco of Argentina. The genus
comprises one of the most morphologically diverse radiations of snakes found in the New
World. Its color patterns range from coral snake mimics to uniform colorations, banded pat-
terns, striped patterns, and complex pigmentation of scales that produce various color and
pattern effects. Within the genus are at least 9-13 subclades (Hurtado-Gomes 2026), many of
which contain one or more species with aquatic habits. Even though some of these species are
widely distributed and locally common they remain poorly studied.
Of interest, Dixon (1983) found no sexual dimorphism in ventral or subcaudal counts,
nor did he find sexual dimorphism in tail/snout-vent length (SVL) ratios in the E. reginae
group. Subcaudal and ventral counts are often sexually dimorphic in snakes with males hav-
ing more subcaudals (and longer tails) and fewer ventral scales than females (Lindell et al.
1993). These traits are closely associated with body size, locomotion, and reproduction and
are thus under continual selection. The absence of sexual dimorphism in these traits makes
the E. reginae group of ecological and evolutionary interest given that not all members of the
genus have these traits. Alencar et al. (2014) found that Erythrolamprus poecilogyrus differed
in overall body and clutch size between geographic regions. Generally, females attain larger
body size than males but do not have a higher number of vertebrae (corresponding with
ventral scale counts). They did not find intersexual differences in tail length, which may
be related to low intersexual competition among males. Diet composition was very similar
throughout the species’ range and characterizes E. poecilogyrus as an anuran specialist. Eis-
feld and Vrcibradic (2019) examined sexual dimorphism of the semi-aquatic Erythrolam-
prus miliaris in southeastern Brazil and detected sexual dimorphism in body size (snout-vent
length), with females averaging larger than males, but no sexual dimorphism in the relative
length of the tail. Eisfeld et al. (2021) analyzed the diet of E. miliaris in five different regions
along the Atlantic Forest in eastern Brazil and found the most frequent prey were fishes and
frogs. Their data suggest that younger snakes feed mainly on anurans, tending to consume

355
Aquatic Snakes

In this chapter are some, but not all of the semi-aquatic snakes in this genus. Also, I
have placed an account for Sordellina punctata, a species not closely related to Erythrolam-
prus at the end of this chapter.
Importantly, many of these snakes hold cryptic diversity that has yet to be resolved.
Their morphology is similar but confusing and molecular data will be needed to resolve
many of these relationships. The 55 known species of Erythrolamprus could easily double in
the next few decades.

Military Ground Snake

Erythrolamprus miliaris (Linnaeus, 1758)
Distribution and Habitat. Brazil, Uruguay, Paraguay,
Bolivia, Peru, Colombia, French Guiana, Guyana, Venezu-
ela, Argentina. The large distribution and polytypic nature
of this species suggest it contains cryptic diversity. Terres-
trial and aquatic situations including swamps, marshes,
streams, floodplains, and coastal habitats are used.
Natural History. The diet includes fish (including the
Figure 23.3. The distribution electric eel), amphibians, reptiles, and mammals (Farina et
of Erythrolamprus miliaris. al. 2019, Tpantiza-Tuguminago et al 2019, de Oliveira et al.
2019).The summary of the subspecies that follows is based
upon Dixon (1983).
Erythrolamprus miliaris miliaris (Linnaeus 1758). Distribution and Habitat. Guyana,
Suriname, and French Guyana. Type locality: South America.
Erythrolamprus miliaris amazonicus (Dunn 1922). Distribution from Santarem, Brazil,
south to Rio Itenez, Beni, Bolivia, east and south into Mato Grosso, Brazil. The type locality
is Santarem, Brazil.
Erythrolamprus miliaris chrysostomus (Cope 1886). Distribution and Habitat. Dense
rainforests of lowland Amazonian parts of Brazil, Colombia, Ecuador, and Peru. Type local-
ity: Upper Rio Amazonas.
Erythrolamprus miliaris merremii (Wied 1821). Distribution and Habitat. Brazil, from
Recife, Pernambuco, to Rio de Janeiro; primarily in the Atlantic rainforest and similar wet-
land sites in eastern Minas Gerais. The type locality is Sao Pedro d’Alcantara, Bahia Brazil.
Erythrolamprus miliaris orinus (Griffin 1916). Distribution and Habitat. The Southern
Coastal Atlantic Forest between the States of São Paulo and Paraná on the eastern slopes
of the Brazilian “Serra do Mar” covered by rainforest. Southeastern Brazil, from southern
Minas Gerais, south through the states of Sao Paulo, Parandi, Santa Catarina, to the north-
ern third of Rio Grande do Sul. Type locality: Sierras of Bolivia; restricted to Sao Paulo, Sao
Paulo, Brazil, by Gans (1964).
Natural History. Marques and Souza (1993) reported this snake preys upon the Frillfin
Goby, Bathygobius soporator, in tide pools. They also cited information from third parties
discussing predation on the Sleeper Goby, Guavina guavina, in mangroves. Duarte et al.
(2014) observed this snake attempting to prey on the Sleeper Goby but the snake in their
observation abandons the prey. They also observed the snake foraging in mangrove crab
(Ucides cordatus) burrows. as well as the surface of the water and the substrate (Farina et al.
2019).

356
 23. The Semi-Aquatic Painted Snakes

Figure 23.4. Erythrolamprus poecilogyrus.

Photography by Myke Clarkson.

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Aquatic Snakes

Yellow-bellied Frog Snake

Erythrolamprus poecilogyrus,
(Wied-Neuwied 1825)
Distribution and Habitat.
A polytypic species, it is present
in many forested and open hab-
itats with (or without) wetlands,

Figure 23.5. Distribution of Eryth-

rolamprus poecilogyrus.

streams, and rivers in South Amer-

ica from se Venezuela and e Guy-
ana to the Argentinian Pampas and
Chaco (Figure 23.3).
Erythrolamprus poecilogy-
rus poecilogyrus (Wied-Neuwied
1825). Distribution and Habi-
tat. Bolivia, Uruguay, Argenti-
na, Paraguay, Brazil (Amazonas,
Para, Minas Gerais, Bahia, Goiás,
SW Pernambuco, São Paulo, Rio
Grande do Sul to coastal Santa Ca-
tarina, Mato Grosso, Piauí); Type
locality: Barra de Jucú, Rio Espirito
Santo, Brazil.
Erythrolamprus poecilogyrus Figure 23.6. Erythrolamprus poecilogyrus. Photography by
caesius (Cope 1862). Distribution (A-B) Steve Barten, (C) Alfredo Sabaliauskas.
and Habitat. Argentina, Bolivia, Par-
aguay; Type locality: Santa Fe, Ar-
gentina.
Erythrolamprus poecilogyrus schotti (Schlegel 1837). Distribution and Habitat. Venezue-
la, Brazil (Sã Paulo, Bahia). Type locality: South America. Restricted to Estado do São Paulo,
Brazil by Hoge (1964)..
Erythrolamprus poecilogyrus sublineatus (Cope 1860). Distribution and Habitat. Uru-
guay, Argentina; The type locality is Buenos Aires, Argentina.
Natural History. The diet of E. p. sublineatus included anurans (54.1%), fishes (42.3%),
reptiles (2.7%) and mammals (0.9%) (Correa et al. 2016).

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Aquatic Snakes

Golden Swamp Snake

Erythrolamprus semiaureus (Cope, 1862)
Distribution and Habitat. Wetlands in savannas, grasslands, and tropical forests in Para-
guay, west and south of Iguazfi Falls, northeastern Argentina, southern and eastern Uruguay
and the southern one half of the Brazilian state of Rio Grande do Sul. Type locality: Sao Joao
de Monte Negro, Rio Grande do Sul, Brazil.
Natural History. Diet is compopsed of mostly fish.
Mature females were significantly longer, had higher body
weight, and reached sexual maturity at a greater length
than males. Males have longer tails than females. The re-
productive cycle was seasonal with greater activity in the
temperate periods of the year, although prior to the flood
Figure 23.7. The distribution of season. Erythrolamprus semiaureus invested more ener-
Erythrolamprus semiaureus. gy in reproduction than in growth, which allows them to
start reproducing at smaller
sizes compared to other pop-
ulations, without delaying
their reproduction until they
reach larger sizes. This can be
an advantageous strategy in
seasonal climates.
Erythrolamprus
cobellus Clade
The Erythrolamprus co-
bellus group, as currently rec-
ognized, includes four spe-
cies: E. breviceps, E. cobellus
(Linnaeus, 1758), E. frenata
(Werner, 1909), and E. taenio-
gaster Jan, 1863 (Fernandes et
al. 2002). E. breviceps and E.
cobellus are distributed only
in the northern portion of
Amazon basin, while E. fre-
nata occurs in central-south-
west Brazil, east Paraguay,
and north Argentina (Dixon
Figure 23.8 Erythrolamprus semiaureus. 1989; Giraudo 2001). Eryth-
Photography by Alfredo Sabaliauskas. rolamprus taeniogaster is the
most widely distributed species
of this group, with two disjunct populations, one in northeastern Brazil ranging from south
Bahia to Pernambuco and Maranhão, and another one in Amazon Basin until Lomalinda,
Colombia, south to Buena Vista, Bolivia, east to the state of Mato Grosso, Brazil (Fernandes
et al. 2002). All have dorsal scale rows 17-17-15, smooth, no apical pits.

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 23. The Semi-Aquatic Painted Snakes

Cope’s Short-headed Watersnake

Erythrolamprus breviceps (Cope, 1860)
Distribution and Habitat. Flood plains, wet agricultural
areas such as rice paddies in Suriname, northwest Brazil,
Ecuador, Colombia, Bolivia, Peru, French Guiana, Guyana.
Type locality: Suriname. Natural History. The diet includes
synbranchid eels, frogs, and earthworms. A clutch of eight
eggs has been reported.
Figure 23.9. The distribution of
Erythrolamprus breviceps.

Figure 23.10. Erythrolamprus breviceps.

Photography by Konrad Mebert

Mangrove Snake
Erythrolamprus cobellus (Linnaeus, 1758)
Distribution and Habitat. Widespread in northern South
America - Colombia, Venezuela, Trinidad, the Guianas, Bra-
zil, and Ecuador. Semi-aquatic in fresh and brackish water;
present in mangroves swamps, herbaceous swamps, rice pad-
dies, and associated streams.
Figure 23.11. The distribution Natural History. Moves overland and may be seen on
of Erythrolamprus cobellus. roads. Active late afternoon and after dark. Diet: includes a
variety of small vertebrates but fish and frogs are likely more
important in its diet than are lizards . Reported to lay eggs in June, clutch sizes less than six
eggs (Beebe 1946; Duellman 1978). This species inhabits swampy coastal areas between 0
and 20 m ASL, ditches along roads in agricultural areas, and secondary-growth vegetation
in water, vegetation near water, and occasionally elsewhere. It has been found in irrigated
watermelon plantations. It feeds mainly on fish .

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Aquatic Snakes

Figure23.12. Erythrolamprus cobellus, Trinidad. Photography by Renoir Auguste.

Werner’s Swamp Snake

Erythrolamprus frenatus (Werner, 1909)

Distribution and Habitat. Distributed mainly within

Cerrado (tropical savanna) and forested parts of the Amazon
Basin in Bolivia, Brazil, Colombia, Peru, and similar areas of
French Guiana, Guyana, Surinam, and Venezuela. Type
locality: Paraguay. Natural History. Batista et al. (2020)
Figure 23.13. The distribution of
observed five specimens in the water and two on land close
Erythrolamprus frenatus.
to water. Of 120 snakes examined only four had prey, all the
were synbranchid eels.

Figure 23.14 Erythrolamprus

frenatus Photography credit
Bruno Catossirosa.

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 23. The Semi-Aquatic Painted Snakes

Jaeger’s Ground Snake

Erythrolamprus jaegeri Günther, 1858
Distribution and Habitat. Open habitats near water
in Brazil, Uruguay, Paraguay, Argentina. A polytypic
species. E. j. jaegeri is in Uruguay, southern Brazil.
Type locality: Brazil. E. j. coralliventris is in Argentina,
Paraguay, southern Brazil. Type locality is the island
north of Concepcion, near San Salvador, northern
Paraguay. Natural History. Prey items consisted of
Figure 23.15. The distribution of anuran amphibians (82.2%), fishes (10.7%) and isopods
Erythrolamprus jaegeri.
(7.1%). Anurans in the diet consisted of Leptodactylus
latrans (35.7%), Physalaemus gracilis (28.6%) and
Pseudopaludicola falcipes (3.6%) and fish prey of
Phallocerus caudimaculatus (10.7%). Isopods could not be identified at the species level
(Correa et al. 2016).

Figure 23.16. Erythrolamprus jaegeri from Corumba, Matto Grosso, Brazil.

Photography by Vinicius Carvalho (top). and Konrad Mebert (bottom).

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Aquatic Snakes

Figure 23.17. Erythrolamprus taeniogaster.

Photography by Ricardo Alexandre Kawashita-Ribeiro.

Jan’s Swamp Snake

Erythrolamprus taeniogaster (Jan, 1863)
Distribution and Habitat. Aquatic and terrestrial habitats, including flooded forests
south of the Amazon in Brazil, to central Brazil and coastal Bahia, Para, Amapa, Paraiba,
Pernambuco, Maranhao, Mato rosso, S Ceara, and Piauí, west to Colombia, south to Bolivia
(Buena Vista), Peru, Ecuador. Type locality: Brazil. It is a polytypic species.
Natural History. Diurnal and nocturnal in forest and clearings. Diet composed of fish,
including Callichthys callichthys, Rivulus sp., Synbranchus marmoratus, and Gymnotus
carapo, as well as frogs. Litter sizes of 7-10 reported in February and October, neonates pres-
ent in May and June.

Linnaeus Royal Snake

Erythrolamprus reginae (Linnaeus, 1758)
Distribution and Habitat. Erythrolamprus reginae occurs from
the Guyana Shield, along the Venezuelan Llanos and Pantepui,
as well as in the Amazon basin of Brazil, Colombia, Ecuador,
and Peru, on the “Brejos de altitude” rainforests in northeastern
Brazil, into the coastal fragments of Atlantic Forest from the
Northeastern and Southeastern Brazil (Ascenso et al. 2019).
Figure [Link] distribution of
Erythrolamprus reginae.
The type locality was given as “Indiis” by Linnaeus but it was
restricted to Suriname by Dixon (1983). It is unclear what the
exact distribution of this snake is. Cryptic diversity has shown
numerous species were previously concealed under this name. Photo on page 357.

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 23. The Semi-Aquatic Painted Snakes

Zweifel’s Royal Snake

Erythrolamprus zweifeli (Roze, 1959)
Distribution and Habitat. Venezuela (Trujillo, Falcon, Amazonas,
etc.) and Trinidad. This species is associated with streams in
premontane and montane humid forest where it forages . The
species tolerates some level of disturbance and uses streams in
coffee plantations and other agricultue between 400-2000 m ASL.
Type locality: Rancho Grande in the state of Aragua, Venezuela,
Figure 23.19. Distribution
of Erythrolamprus zweifeli. at an elevation of 1100 meters, in a cloud forest. Identification.
Erythrolamprus zweifeli differs from E. reginae in having a salt-
and-pepper dorsal pattern (vs dorsum with dense pale and dark
paravertebral flecking in E. reginae) and in having a higher number of subcaudal scales
(69−88 vs 55−78 in E. r. regina. On Trinidad this snake hunts along forested streams for
frogs, their tadpoles and fish.

Below. Figure 23.20. Erythrolamprus zweifeli from Trinidad.

Photography JCM.

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Aquatic Snakes

Cobra D’água
Erythrolamprus macrosomus (Amaral, 1936)
Distribution and Habitat. Inhabits semideciduous forests
of the Atlantic Forest in Paraná and São Paulo states, in the
Brazilian Cerrado in Goiás, Mato Grosso, Mato Grosso do
Sul, Minas Gerais, São Paulo, and Paraná states, and in the
Chaco from Paraguay and Argentina. The distribution ex-
Figure 23.21 The distribution of tended to El Palmar National Park, province of Entre Ríos,
Erythrolamprus macrosomus. Argentina, representing the southernmost record (based
upon Ascendo et al. 2019).

Figure 23.22 Erythrolamprus

macrosomus. Photography by
Alfredo Sabaliauskas.

Erythrolamprus melanotus Clade

The status of this clade needs verification. It may belong with the E. cobellus clade, or it
may be distinct. Murphy et al. (2019) recovered Erythrogaster pseudoreginae and E. epineph-
alus as the sister to E. melanotus. Erythrolamprus pseudoreginae is endemic to Tobago and
poorly known. It has been found within several meters of streams. Erythrolamprus epinepha-
lus is a widespread mainland species complex associated with marshes and wetlands
Maps and photos of most of these are not presented here.

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 23. The Semi-Aquatic Painted Snakes

Green Marsh Snake

Erythrolamprus epinephalus (Cope, 1862)
Distribution and Habitat. Wetlands, marshes, riparian floodplains, and wet forests from
Costa Rica southward to Panama and Colombia, Peru, northwest Ecuador and westward to
Venezuela. The type locality is Truando, (New Granada) Colombia (no map)..
A polytypic species. Torres-Carvajal and Hinojosa (2020) found the species to be para-
phyletic and support recognition of the subspecies albiventris, fraseri and lamonae as distinct
species. Leaving the subspecies: E. e. bimaculatus from Colombia, Venezuela, Ecuador; with
a type locality of Colombia. “Presumably from the vicinity of Bogotá” (Dunn 1944); E.e. juve-
nalis from Costa Rica with a type locality of San Jose, Costa Rica; E. e. kogiorum from Colom-
bia (Magdalena), Venezuela (Zulia) with a type locality of Ciudad Perdida,Sierra Nevadade
Santa Marta, Departamento de Magdalena, Colombia, 1100 m elevation; E. e. opisthotaenius
from the Venezuelan states of Merida, Barinas, Táchira, Trujillo, Zulia, Falcón), and Colom-
bia; and E. e. pseudocobella from W Colombia (Andes), Ecuador; with a type locality of
Angelópolis, Colombia.

White-bellied Marsh Snake

Erythrolamprus albiventris (Jan, 1863)
Distribution and Habitat. West of the Andes in Ecuador and Colombia. The type local-
ity is western Andes of Ecuador and Guayaquil, Ecuador (No Map). Diurnal and terrestrial,
inhabits evergreen to semi-deciduous forests, cloud forests, and humid montane shrublands.
The species also occurs in areas having a matrix of pastures, plantations, and remnants of
native vegetation, as well as in gardens in urban areas. They are common in marshes, swamps,
artificial ponds, and along streams; they enter caves and abandoned mines. Most active in
the morning, crossing roads and trails, basking in open areas, or foraging on leaf-litter, soil,
or among grass or shrubs. May climb up to two meters in vegetation. They hide under logs,
agave plants, stones, and in crevices.
Natural History. They actively hunt frogs and tadpoles. Lizards are also preyed upon.
Often, they rely on crypsis to avoid predation. However, if disturbed, they flatten the body
and produce cloacal musk. Predators include snakes and domestic animals.
Breeding congregations have been observed during the rainy season in western Ecuador
(December–May). In one hole, five males and one female were coiled into a reproductive ball.
Females lay 5–10 eggs in heaps of rotten vegetation (account based on Arteaga 2020a).

Olive Marsh Snake

Erythrolamprus fraseri (Boulenger, 1894)
Distribution and Habitat. Western side of the Andes in Ecuador and Peru. The type lo-
cality is western Ecuador.
Natural History. A diurnal and terrestrial snake that inhabits humid to dry montane
shrublands, highland grasslands, high evergreen montane forests, and areas having a ma-
trix of pastures, plantations, rural gardens, and remnants of native vegetation. Individuals
occur in rural gardens, along roads, and occasionally inside houses. They also use marshes,
swamps, and streams. Most active on sunny morning, basking in open areas or foraging on
leaf-litter, mud, or in tall grass. Hide under debris on the ground. Hunt amphibians, mostly

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Aquatic Snakes

frogs and tadpoles. Flee when threatened but rely mostly on crypsis. If disturbed, they may
flatten their body and produce release cloacal musk (based on Arteaga 2020b).

Lamon’s Ground Snake

Erythrolamprus lamonae (Dunn, 1944)
Distribution and Habitat. Colombia (Andean slopes, 1,500 - 2,600 m elevation) and
southward to east central Ecuador. The type locality is Sonsón, Antioquia, Colombia at 2410
m.

Shaw’s Dark Ground Snake

Erythrolamprus melanotus (Shaw, 1802)
Distribution and Habitat. Colombia, Venezuela (Co-
jedes), Isla Margarita, Trinidad, Tobago, and Grenada.
The type locality was given as the Cape of Good Hope, Af-
rica in error. A forest and forest-edge snake that is closely
Figure 23.23. The distribution of associated with streams and ponds, it will also use cacao
Erythrolamprus melanotus. plantations and other disturbed habitats.
This is a polytypic species with three subspecies: E.
m. melanotus, E. m. neosus (Trinidad & Tobago), and E. m. lamari (Zulia, Venezuela).
Natural History. Diurnal. Diet in the Trinidad and Tobago population includes small
vertebrates, mostly frogs (Engystomops pustulosus, Pristimantis urichi) tadpoles, and lizards
(Bachia trinitatis, Gonatodes vittatus). Clutches of 2−10 eggs have been reported between
January and August.

Figure 23.24 Erythrolamprus

melanotus from Tobago Photog-
raphy by JCM.

Sordellina punctata Clade

Sordellina punctata is a poorly known species from Brazil that, until recently, had an
uncertain phylogenetic position. Molecular data place this species in the tribe Echinanterini
(Miranda et al 2012; Abbeg et al. 2022). There is a single species in the genus. It inhabits the
Atlantic Forest wetlands of Brazil.

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 23. The Semi-Aquatic Painted Snakes

Yellow-spotted Snake
Sordellina punctata (Peters, 1880)
Distribution and Habitat. Occurs in the Atlantic Forest and
southeast Brazil with confirmed specimens from the states of São
Figure 23.25. The distribu- Paulo, Paraná and Santa Catarina. Pereira et al. (2007) suggest
tion of Sordellina punctata. reports of this species from the western Mato Grosso do Sul and
all literature citations for Rio de Janeiro are questionable. Pereira
et al. also suggest this species is probably associated with water-
logged soils than aquatic habitats. Muddy and soaked soils in floodplains were reported as
habitat.

Figure 23.26. Sordellina punctata.

Photography by Wolfgang Wuster.

Natural History. A species specialized to feed on earthworm and occasionally includes

the giant earthworm in its diet (Marques 1996, Oliveira et al. 2003, Pereira et al. 2007). Pereira
et al. (2007) found the burrowing caecilian Chthonerpeton indistinctum in the stomach of S.
punctata. The caecilian is common in muddy riparian floodplains and wetlands. They did not
consider this a truly aquatic snake writing that, “It seems more associated with wetlands and
other soak soils, surrounding lakes or rivers, than water bodies itself.” However, water-soaked
soils are indeed an aquatic environment. It is not “open water,” but it is a water-filled micro-
habitat. This species is an aquatic burrower.

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Aquatic Snakes

The Cosmopolitan Watersnake Section

The Cosmopolitan Water Snakes is a family level (Natricidae) clade (Zaher et al. 2009,
Burbrink et al. 2020). Other authors consider it a subfamily (Natricinae) of the Colubridae.
Here I will refer to the clade as the family Natricidae (natricids). Most species occur north of
the equator, and the greatest diversity occurs in Asia and North American and Sub-Saharan
Africa have radiations that are monophyletic.
This manuscript was quite far along when Deepak et al. (2021) published their overview
of natricids . Ideally, the organization of the next seven chapters would have followed their
trees. The text has been adjusted but does not completely follow their results..
The oldest extant lineages found by Deepak et al. (2021) are found today in mainland
Asia and India+Sri Lanka (Amphiesmoides and Aspidura), and their estimation of the ances-
tral area suggests natricids were likely from one of these two areas. Deepak et al. (2021a, b)
also propose a single dispersal event from India plus Sri Lanka to sub-Saharan Africa and
from mainland sub-Saharan Africa to the Seychelles. Extant natricids from the Philippines
were hypothesized to have originated by dispersal from Sundaland + Wallacea. Extant na-
tricids from Australo-Melanesia were inferred to have evolved from a primarily Philippine
clade. The Sundaland + Wallacean, Philippine and Australo-Melanesian natricines were all
suspected as originating from an initial dispersal from India plus Sri Lanka. Extant natricids
from North and Central America and Europe, Central Asia, and North Africa were recipro-
cally monophyletic, and the best-fitting model suggested that the dispersal route was from
mainland Asia plus Japan to North and Central America to Europe, Central Asia, and North
Africa (Deepak et al. 2021).
Natricids are semi-aquatic or aquatic (53%) and terrestrial (28%); aquatic burrowers
(10%) or terrestrial burrowers (8%). Deepak et al. (2021) estimated the ancestral habitat of
all natricines to be terrestrial (~50%), followed by semi-aquatic/aquatic. Many natricids are
dietary generalists (37%) or aquatic generalists (25%). Some feed only on anurans (18%), and
others on worms (12%). Most vermivores are also burrowers or aquatic burrowers. A few
species (8%) are dietary specialists feeding on fish, mollusks, or crayfish. The ancestral state
estimation for all natricines shows a higher proportion of anuran or aquatic generalists than
the other six diet categories.
All North American natricids are viviparous. The only non-American viviparous natri-
cids are the Asian Pseudagkistrodon rudis and Trimerodytes annularis. These two species rep-
resent independent origins of viviparity from oviparous ancestors. When testing for associa-
tions between habit and reproductive mode, the best-fitting model was one with symmetric
transition rates among states. This suggests that there was no association between habitat and
reproductive mode (Deepak et al. 2021).
Most extant species are terrestrial or semi-aquatic and are often dietary generalists, yet
some have specialized their diet. Fully aquatic species are typically generalists, but some feed
only on crayfish or worms, small crustaceans, or fish (Deepak et al. 2021). Natricids range in
maximum size from ~115 to 1195 mm SVL, with many of the smallest species being fossorial
(Deepak et al. 2021).
Most are harmless to humans, but a few garter snakes (Thamnophis) have venom, caus-
ing localized symptoms. However, a few species of Asian Rhabdophis have venom that is
life-threatening to humans.
Natricids were present in North American in the early Miocene (Holman, 2000) and by
the late Miocene members of the genera Nerodia, Thamnophis, and Neonatrix were present.
Mead and Steadman (2017) found Nerodia fossils in the late Pleistocene site Sawmill Sink,

369
 The Cosmopolitan Watersnake Section

on Abaco Island in the Bahamas. Georgialis et al. (2019) describe natricids fossils from the
Neogene (latest Miocene or earliest Pliocene) locality of Maramena, in northern Greece. Na-
trix aff. rudabanyaensis, and another distinct species of Natrix probably inhabited nearshore
environments. The Cosmopolitan Water Snakes are discussed in the next seven chapters.
The oldest natricid lineage recovered by Deepak et al. (2021) is Amphiesmoides ornati-
ceps (White-eyed Keelback). It occurs along forested streams in southern China (including
Hainan) and northern Vietnam. Its type locality is Northern Hainan, China.

Natricid distribution. Modified from Deepak et al. 2021. The chapters covering the
various clades are given on the map.

On the next page Pseudoagkistrodon

rudis Photography by Konrad Me-
bert.

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24. Mountain Keelbacks

This chapter discusses the genera Pseudagkistrodon, Rhabdops Herpetoreas, and Hebius.
Das et al. (2020) recovered these four genera in a clade with the terrestrial-fossorial Trachis-
chium monticola and two species of Blythia. Trachischium are worm-eating, oviparous, fosso-
rial snakes from the Trans-Himalayan region of China, Tibet, and north Vietnam (Ren et al.
2018, Wang et al. 2019). The natural history of Blythia is poorly known.
Herpetoreas contains seven species distributed along the southern edge of the Hima-
layas, into northeast India and adjacent China. Five species of Herpetoreas (H. burbinki, H.
platyceps, H. sieboldii, Herpetoreas tpser, H. venningi ) are known only from subtropical and
temperate regions above 900 m (up to 3056 m) ASL. The northeast Indian Herpetoreas xenura
and H. pealii,) on the other hand, occur from 30 m up to 1170 m. Herpetoreas are semi-aquat-
ic snakes with keeled dorsal scales. The posterior maxillary teeth are the longest, in a con-
tinuous series, with the anterior ones. Body and tail slender and compressed. Two nasals,
one loreal, one anterior, and two posterior oculars. Dorsal scales are moderately elongated,
keeled, and in nineteen rows.
Giri et al. (2017) used molecular data to confirm that the aquatic Indian genus Rhabdops
also belongs to this clade. Two species of Rhabdops inhabit the Western Ghats of peninsular
India. These two species are the most aquatic members of the clade.
Snakes of the genus Hebius were formerly in the genus Amphiesma and were commonly
called keelbacks. They are terrestrial to semi-aquatic snakes that are oviparous and wide-
spread in Asia. Currently, 50 species are known – here, I discuss a sample (25%) of the species
in the genus.
David et al. (2021) recognized two informal groups among the “dark-bellied” species of
Asian natricid snakes. These two groups share: (1) 15, 17 or 19 dorsal scale rows; (2) a very
dark dorsum (dark grey, dark brown, or blackish-brown); (3) dark reticulations on the back,
often with orange or reddish-brown spots or dorsal blotches; (4) venter at least in part black-
ish-brown or black; (5) postocular stripe absent or present; and (6) strongly keeled scales at
the base of the tail in those species with 17 dorsal scale rows. These groups include species
referable to the genus Hebius, as currently conceived following Guo et al. (2014), but also the
two species formerly placed in the genera Parahelicops Bourret, 1934 and Pararhabdophis
Bourret, 1934 are now placed in Hebius.
The 48 species of Hebius are on the continuum between terrestrial to semi-aquatic. The
color pattern commonly consists of dark dorsolateral spots forming two longitudinal stripes.
Many also have a black or very darkly pigmented venter. Hebius andreae and H. leucomystax
are sisters to all the other members of the genus. Here, I have accounts for 12 species, which
may or may not be the most aquatic ones. The lack of knowledge on habitat use in these
snakes is significant. David et al. (2021) provide detailed morphological descriptions for
these snakes - I do not repeat that here
Pseudagkistrodon rudis is common in montane meadows, riparian areas, valleys, and
roadsides. It ranges from 600–2,650 m ASL. The species is polytypic, with three recognized
subspecies.

Annam Keelback
Hebius annamensis (Bourret, 1934)
Distribution and Habitat. Vietnam and southeast Laos. Type locality: Bana, Annan, Viet-
nam (now Ba Na Nature Reserve, Da Nang City, Vietnam), 1500 m elevation. Habitat in-

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 24. Mountain Keelbacks

cludes tropical wet evergreen forests and subtropical mon-

tane evergreen forests between about 300-1,500 m ASL,
although most specimens for which elevation data are
available were collected between 1,200 and 1,400 m ASL
(Stuart & Nguyen 2012a). All were collected in primary
montane evergreen forests and in close association with
Figure 24.1. The distribution of fast-running, rocky forest streams. It is mostly a noctur-
Hebius annamensis. nal and aquatic snake, although one specimen was found
on the ground. According to Stuart (2006), specimen
FMNH 258637 was collected
at night (20.30 h) in steep ter-
rain covered by wet evergreen
forest between 1,280–1,500 m
elevation. The snake was first
observed on land 50 cm from
a small, swift, rocky stream,
then it dove into the water
and swam under a rock on
the stream bottom. Specimen
ZFMK 86457 was captured at
night (20.00–21.00h) in the
leaf litter of a forest stream
at 510 m ASL, with only the
snout tip protruding from the
surface of the water (David et
al. 2015b). The diet and repro-
ductive habits of Hebius an-
namensis, remain unknown.

Vietnam Water Snake

Hebius chapaensis
(Bourret, 1934)
Distribution and Habi-
tat. Streams in Laos, northern
Vietnam (Lao Cai, Yen Bai)
and China (Yunnan). Type
locality: Chapa, Tong-King
(= Sa Pa, Lao Cai Province,
Figure 24.2. Hebius annamensis. Hoa Son Quang Binh (A), Kon North Vietnam). At eleva-
Tum (B) Vietnam. Photography by Truong Nguyen. tions of primary forest, at el-
evations between 1050 - 2046
m.
Natural History. Tropical
and subtropical evergreen montane forests between 900 and 2,046 m ASL support this spe-
cies (Ren et al. 2018). All specimens for which data are available were collected near streams
in montane, wet evergreen forests. This species is nocturnal, semiaquatic, and terrestrial;
no specimen was collected more than a few meters from water. A specimen from Laos was
373
Aquatic Snakes

swimming in submerged vegetation along the bank of a three-

meter-wide stream with moderate current; another specimen
was seen perched on a tree one meter from a stream. A spec-
imen observed by Ren et al. (2018) was actively preying upon

Figure 24.3. The distribution

of Hebius chapaensis.

tadpoles of the treefrog

Rhacophorus duboisi on the
edge of a pool. Brakels et
al. (2021) found it feeding
on the frog Amalops crem-
nobatus. The snake was un-
der a rock in a stream swal-
lowing the frog head first.

Figure 24.4. Hebius chapaensis

Photography by Kai Wang.

Kuatun Keelback
Hebius craspedogaster (Boulenger, 1899)

Figure 24.5. The distribution Distribution and Habitat. Montane forests associated with
of Hebius craspedogaster. streams in Southern China and Vietnam. Type locality: Gua-
dun, Fujian Province, China, 500-2000 m elevation or higher.
Natural History. Pope (1935) found this snake common in
the Yenping Mountains and extremely abundant about Sanchiang and Kuatim in Chimgan
Hsien. It was present in forests 1500-6000 feet ASL and is considered semi-aquatic. Pope
recovered young frogs or toads and tiny tadpoles from the digestive systems of this species.

Figure 24.6. Hebius craspedogaster. Photography by Kevin Messenger.

374
 24. Mountain Keelbacks

Chiang Mai Keelback

Hebius deschauenseei (Taylor, 1934)
Distribution and Habitat. Semi-evergreen and mixed de-
ciduous submontane and montane forest in Chiang Mai Prov-
ince, Thailand.
Natural History. Chan-ard et al. (2015), found this species
inhabits semi-evergreen and mixed deciduous submontane and
montane forest. It is nocturnal and terrestrial and occurs in the
Figure 24.7. The distribution
of Hebius deschauenseei.
vicinity of forest streams. Nothing else is known of the biology
of this rare species Nocturnal and terrestrial. The degree of its
aquatic habits are unknown.

Figure 24.8. Hebius deschauenseei, Doi Phu Kha National Park, Thailand.
Photography by Rushen Bligin.

Modest Keelback
Hebius modestus (Günther, 1875)
Distribution and Habitat. A semi-aquatic stream dweller
known from Myanmar, India, Myanmar, Yunnan, China. Type
locality, the Khasi Hills, India. It may be present in Bhutan.
There are questionable records from Vietnam. It is unknown
from Assam; Mizoram (India); Laos, Cambodia; Guizhou,
Guangdong, China; Mizoram, India.
Natural History. A poorly known species. This species
Figure 24.9. The distribution inhabits evergreen or semi-evergreen submontane forests up
of Hebius modestus.
to 1,600 m ASL, and is associated with hill or montane forest
streams. It occupies the forest litter in riparian areas. It probably

375
Aquatic Snakes

feeds on tadpoles and frogs. Based

on the photo from Jian Wang it also
feeds on fish. Wall (1926) recorded
a female, preserved between June
and August, that contained three
large eggs in its oviducts. This spe-
cies is either secretive or rare (Da-
vid et al. 2021).

Figure 24.11. Hebius

modestus, (A) is from Mt.
Mopan, Xinping Coun-
ty, Yunnan, China; (B-C)
is from Mt. Cenwang-
laoshan, Tianlin County,
Guangxi, China. Photog-
raphy by Jian Wang.

Black-bellied Keelback
Hebius nigriventer (Wall,
1925)
Distribution and Habitat. Ev-
ergreen or semi-evergreen sub-
montane forest and mixed or de-
ciduous moist montane forest from
350 up to 1,550 m ASL from Myan-
mar (Kachin State. Bhamo District;
Myitkyina District); People’s Re-
public of China (Yunnan Province.
Yingjiang County, Dehong Dai and
Jingpo Autonomous Prefecture;
Gongshan County, Nujiang Prefec-
ture; Mengla County, Jinhong Pre-
fecture.) It may also be present in
northern Laos.
Natural History. Wall (1925) noted one had swallowed two large and four small tadpoles.
Diurnal and active at dusk and early evening, at air temperatures from about 24 to 27 °C.
Mostly aquatic. It occurs in hill and montane streams, between
or on rocks in the stream bed. Another one, a female col-
lected in June, contained two large eggs, indicating that egg
laying would have taken place at the beginning of the rainy
season (David et al. 2021).

24.10. The distribution of Hebi-

us nigriventer.

376
 24. Mountain Keelbacks

Figure 24.12. Hebius nigriventer

Photography by Jian-Huan Yang.

Eight-lined Keelback
Hebius octolineatus (Boulenger, 1904)
Distribution and Habitat. SW China (Yunnan, Sichuan,
Guangxi). Type locality: Kunming Shi, Yunnan Province,
China; about 2000 m elevation.
Natural History. Mell (1922) and Pope (1935) agree that
thisi is the most common snake in cultivated fields from
Yunnanfu to Tali and Pope reports small fish and a frog in
Figure 24.13. The distribution
of Hebius octolineatus. three stomachs, suggesting that this is an aquatic snake.

377
Aquatic Snakes

Figure 24.14. Hebius octolineatus from Dali Bai, Yunnan, China.

Photography by Xavier Rufray.

Pope’s Keelback
Hebius popei (Schmidt, 1925)
Distribution and Habitat. It inhabits mountain and hill
Figure 24.15. The distribu- streams in forests at 281–900 m in China (Hainan, west to Hu-
tion of Hebius popei. nan and Yunnan) and Vietnam. Type locality: Nodoa, Hainan,
China.

Figure 24.16. Hebius popei.

Photography by Konrad Mebert.

378
 24. Mountain Keelbacks

Yunnan Keelback
Hebius parallelum (Boulenger, 1890)
Distribution and Habitat. India (Sikkim, Nagaland), Nepal
(?), Bhutan. Type locality: Sikkim. To at least 1890 m ASL.
Figure 24.17. The distribu- Natural History. Literature states its diet includes tadpoles
tion of Hebius parallelum. and fish.

Figure 24.18. Hebius parallelum from Nagaland.

Photography by Viral K. Mystery.

Kosempo Keelback
Hebius sauteri (Boulenger, 1909)
Distribution and Habitat. Taiwan, southern China (Jiangxi, Guangxi, Guangdong, Fuji-
an, Sichuan, Hubei, Hong Kong), and north Vietnam to elevations of 1200 m. Type locality:
Kosempo, Formosa (it is present in the Jiaxian District, Kaohsiung City, south Taiwan).

379
Aquatic Snakes

Natural History. A montane, semi-aquatic, forest dwell-

ing snake that feeds on tadpoles and slugs (Pope, 1935), frogs
and earthworms (Mao, 1998). Oviparous (Kuntz, 1963).

igure 24.19. The distribution of

Hebius sauteri.

Figure24.20. Hebius sauteri, China.

Photography by Gernot Vogel.

Kachin Keelback
Hebius taronensis Smith, 1940
Distribution and Habitat. David et al. (2021) reports
this species inhabits semi-evergreen submontane for-
est and mixed or deciduous moist montane forest from
about 1,000 up to at least 1,850 m ASL. The species is
thought to be diurnal. Most specimens were obtained
from small mountain streams. One was eating a frog,
Figure 24.21. The distribution of another specimen had eaten tadpoles. Others were found
Hebius taronensis.
under rocks near a stream.

380
 24. Mountain Keelbacks

Figure 24.22. Hebius taronensis. From Lake Indawgyi, Mohnyin Township,

Kachin State, Myanmar.
Photographic credit Nikolay Poyarkov.

Chin Hills Keelback

Hebius venningi (Wall, 1910)
Distribution and Habitat. Hill streams to at least 1040
m. of India (Meghalaya, Mizoram, Arunachal Pradesh),
northern Myanmar, China (Yunnan). Type locality: Haka
Chin Hills, Myanmar. It occurs berween 900-1800 m ASL.
Diagnosis. Dorsum dark grey, brown with indistinct
black checkering with a dorsolateral chain of yellow spots.
Figure 24.23. The distribution of
An incomplete yellow collar may be present. Ventral sur-
Hebius venningi.
face yellow. Dorsal scales in 17 rows, weakly keeled,
outer rows smooth. Two preoculars, nine upper la-
bials (4-6 border orbit. Ventrals 158-172, subcaudals
117-140. Total length to at least 680 mm, tail 225
mm. Natural History. The diet includes tadpoles
and frogs.

Figure 24.24. Hebius venningi

Photography by Ashok Captain.

381
Aquatic Snakes

Burbrink’s Keelback
Herpetoreas burbrinki Guo, et al., 2014
Distribution and Habitat. Xizang, China. Type locality:
Zayu County, Xizang A. R., China, at an elevation of 1889 m.
At this writing this species is known from two specimens from
the type locality.

Figure 24.25. The distribution

of Herpetoreas burbrinki .

Assam Keelback
Herpetoreas pealii (Sclater, 1891)
Distribution and Habitat. Small, slow-moving streams
with muddy bottoms that transect forests in Assam, Arunachal
Pradesh, India. Type locality: Sibsagar district of Assam (Das et
al. 2020).

Figure 24.26. The distribu-

tion of Herpetoreas pealii.

Figure 24.30. Herpetoreas pealii

Photography by Abhijit Das.

382
 24. Mountain Keelbacks

Himalayan Keelback
Herpetoreas platyceps (Blyth, 1854)
Distribution and Habitat. Kashmir, Sikkim, Punjab, As-
sam, Darjeeling, Meghalaya, Arunachal Pradesh, Himachal
Pradesh,
Jammu and Kashmir, India; Nepal (1040-3657 m ASL),
Bangla-desh, Pakistan, Bhutan, Tibet, China. Type locality:
Figure 24 27. The distribution of Restricted to “Darjiling” by Malnate (1966).
Herpetoreas platyceps. Natural History. The stomachs of specimens examined
con-tained the skink Leiolepisma himalayanum. Malnate
(1966) found three of the four snakes having fed on this

Figure 24 28. The eastern form of Herpetoreas platyceps

Photography by Abhijit Das.

lizard had swallowed their prey tail first. One snake contained eight adult and four juvenile
specimens of L. himalayanum. A female 111 mm in total length, containing two large (25 X
8 mm) eggs.

Siebold’s Keelback
Herpetoreas sieboldii Günther, 1860
Distribution and Habitat. Pakistan, India (Uttar Pradesh,
Sikkim, Punjab), Nepal, Myanmar, Bangladesh, and Bhutan.
Type locality: Sikkim, Himalaya 2500 m ASL).
Figure 24 29. The distribution Natural History. Prey includes frogs, tadpoles, frog eggs,
of Herpetoreas sieboldii. and skinks of the genus Lygosoma. A female, 435 mm. in total
length, contained five small, undeveloped eggs (Malnate 1966).
383
Aquatic Snakes

Mêdog Himalayas Keelback

Herpetoreas tpser Ren, Jiang, Huang, David and Li, 2022
Distribution and Habitat. Evergreen broad-leaf forest or the moist fields of forest mar-
gins covered with dense vegetation close to the water. Known from Beibeng and adjacent Ani
Bridge, Hanmi, and 80K, Mêdog County, southeastern Xizang Autonomous Region (Tibet),
southwestern China, at an altitude of 1087–2280 m ASL. [No Map.]
Natural History. Diurnal, at least on a cloudy or mildly rainy days. The species has a
fierce disposition and bites when handled, often holding the mouth open while on the defen-
sive. The specific epithet tpser is the acronym of “Tibetan Plateau Scientific Expedition and
Research” of China, which promoted the scientific research of the Qinghai-Tibet Plateau. All
Herpetoreas tpser were collected during The First and the Second Tibetan Plateau Scientific
Expedition and Research.

Wall’s Keelback
Herpetoreas xenura (Wall, 1907)
Distribution and Habitat. Forest floor near streams Bangladesh, Myanmar, and Assam,
India. Type locality: Cherrapunji,
Khasi Hills, Assam.
Natural History. Diet. It is
known to feed on anurans and
snakes including Hydrophylax
leptoglossa, Ingerana borealis,
Rhabdophis himalayanus (De-
cemson and Lalremsanga 2021)
Figure 24.31. Herpetoreas xe-
nura. Photography by Abhijit
Das.

Aquatic Rhabdops
Rhabdops aquaticus Giri, et
al., 2017
Distribution and Habitat. it
is restricted to the Western Ghats
region of Goa, on the north at
Karnataka and on the south to
Maharashtra, at elevations of
750–1000 m. The species is encountered mostly during the monsoon season, when it is pre-
dominantly aquatic or semi-aquatic, and can be observed along
streams in semi-evergreen forest and waterlogged habitats
At the type locality, Giri et al. found adults in streams tra-
versing the semi-evergreen forest and juveniles seen mostly on
lateritic plateaus during the monsoon; it was also encountered
Figure 24.33. The distribu- close to a village. Small individuals were seen mostly under rocks,
tion of Rhabdops aquaticus.. always in moist or waterlogged areas. Larger individuals were en-
countered primarily on large, seasonal pools, though also during

384
 24. Mountain Keelbacks

the day in wet soil under rocks.

Natural History. Activity is
primarily nocturnal; foraging oc-
curs in lentic or slowly flowing
water. Opportunistic observation
of five actively foraging individu-
als in a stream stayed submerged
for 4–12 minutes. During the day,
R. aquaticus was mostly under
rocks close to streams or on pla-
teaus. During the day, individu-
als bask on exposed rocks in the
stream. Giri et al. (2017) have no
Figure 24.34. Rhabdops aquaticus, Chalkewadi, Satara, direct observations on the diet of
Maharashtra, India. R. aquaticus. Still, remnants of ar-
Photography by Christian Langner.
thropod (seemingly crustacean)
exoskeletons were observed in the
feces of one specimen temporarily kept for observation and photography. Specimens were
observed in ephemeral pools under rocks directly alongside remnants of freshwater crabs.

Olive Ghats Keelback

Rhabdops olivaceus (Beddome, 1863)
Distribution and Habitat. Semi-evergreen hill forest Western Ghats region of penin-
sular India. A total of six individuals were encountered in three different habitats by Bho-
sale and Joshi (2014). Four were found in the Kaas Region, and two were recorded from
Chalkewadi Plateau. A specimen from Kaas Lake was found active during the day, whereas
individuals from Chalkewadi and Kaas plateau and forest stream were inactive under me-
dium-sized rocks. Four specimens from plateaus (2 from
each Kaas and Chalkewadi) were found in water-logged
conditions (>2 inches depth). Our observations suggest
that Rhabdops olivaceus chiefly inhabit the seasonal pools
on the plateaus apart from the hilly forest streams, which
can be considered among their preferred monsoon habi-
Figure 24.35. The distribution of tats. In all four localities, individuals were found under
Rhabdops olivaceus. small to medium-sized rocks in soggy conditions. This
habitat was also shared by Hemidactlyus cf. brookii (geck-
os were found on lower surfaces of the small-sized boul-
ders under which the snake was encountered), Monopterus sp., Nyctibatrachus sp., etc. The
specimen from Kaas represents the northernmost locality to date. A review of district bu-
tion records reveals its occurrence in at least four Indian states: Goa, Maharashtra, Karna-
taka, and Kerala. Their observations indicate that species are not restricted to perennial hill
streams but also occupy seasonal water bodies like the ones that develop post-monsoons on
the lateritic plateaus at Chalkewadi and Kaas.
Bhosale and Joshi (2014) observations suggest that seasonal water pools on the pla-
teaus are one of the preferred habitats of this species. Their observations in the Satara dis-
trict attest to this.

385
Aquatic Snakes

Figure 24.36. Rhabdops olivaceus, Kunnathidavaka, Kerala.

Photography by David Raju.
The patchy distribution and its small size can lead to its uncommon occurrence, thus
enlisting it as a rare species and remaining poorly known. The species seems widespread
throughout the Western Ghats data, however, is preliminary, and it is hoped that additional
information on natural history, distribution, and variation in habitat will be provided. Indeed,
significant fieldwork is required for further studies of this species.(Bhosale and Joshi 2014).
Natural History. It is a nocturnal, semi-aquatic snake found in slow-moving water.
The diet of R. olivaceus is reported as fish, slugs, earthworms, and soft-bodied invertebrates
(Whitaker and Captain 2004; Radhakrishnan 1997; Srinivasulu et al. 2014). However, Giri et
al. (2017) could not find a primary published report to confirm this.

False Habu
Pseudagkistrodon rudis Boulenger, 1906
Distribution and Habitat. Taiwan, southern China (Fujian,
Yunnan, Sichuan, Guizhou, Hunan, Jiangxi, Zhejiang, Guangxi,
Guangdong). Type locality. Tongchuan, Yunan, China. A poly-
typic species with two subspecies: Pseudagkistrodon rudis rudis
Boulenger 1906. Pseudagkistrodon rudis multiprefrontalis (Zhao
and Jiang 1981) which ranges in China (Sichuan) and has a type
Figure 24.37. The distribu-
locality: Xichang, Sichuan Province, at about 2650 m ASL.
tion of Pseudoagkistrodon
rudis.
Identification. Stout, brown body with black blotches, and
the crown maybe black, blotches separated by narrow brown-
ish-white interspaces; posteriorly the ground-color passes gradually to olive-brown, with a
median series of round and a lateral series of longitudinally elliptic blackish spots; blackish

386
 24. Mountain Keelbacks

Figure 24.38. Pseudoagkistrodon rudis

Photography by Konrad Mebert.

brown above, cream-white on the upper labials; venter grey-olive speckled with black ante-
riorly, blackish speckled with white posteriorly. Rostral one and a half times as long as deep,
scarcely visible from above; upper head-shields rugose ; internasals broader than long, nearly
as long as the prefrontals ; frontal once and one fourth as long as broad; loreal small, deep-
er than long ; three preoculars, four post oculars, and three suboculars separating the eye
from the labials ; temporals scale-like, strongly keeled ; seven upper labials ; five lower labials
in contact with the anterior chin-shields, which are shorter than the posterior. Scales very
strongly keeled, in 19-25 rows, 23 rows at midbody. Ventrals 123-156 (Zhao et al., 1998); anal
divided; subcaudals 37-65 (Maki, 1931; Zhao et al., 1998). Maximum size in total length re-
spectively male 97.5 cm and female 121.7 cm (Zhao et al., 1998).
Natural History. A diurnal-nocturnal species in mountainous forest floors, bush- and
grassland, and near streams. The diet is primarily toads, but frogs, small snakes, lizards, in-
sects, and earthworms are also prey. Viviparous, with litters of 12-27 young in late summer
and fall. Neonates 130-200 mm in total length. Defensive behavior includes flattening the
head and forebody, or the entire body, it coils up. When flattened, the oval head becomes
triangular to mimic vipers. This species has rear fangs that are kinetic and which appears to
be the specialisms for toad ingestion (Wu, 2004), and a venom gland may be able to deliver a
medically important bite to humans.

387
Aquatic Snakes

388
Aquatic Snakes

25. Keeled Water Snakes

Nathan Rusli, Gunther Köhler,
Jean-Jay Mao, and John C. Murphy

The keeled water snakes are terrestrial and semi-aquatic species. However, as a clade,
they are not well studied. All forms are thought to be oviparous, with clutch sizes small to
moderate (2 to 11). Specific data are included in the accounts of the species. Unfortunate-
ly, much of the information on Tropidonophis here is based upon Malnate and Underwood
(1988) a publication badly in need of revision.
Tropidonophis ranges from the central and southern Philippine Islands southeastward to
the Moluccas and east through New Guinea and the Bismarck Archipelago to the Trobriand
Islands and the Louisiade Archipelago. The genus also occurs in northeastern Australia. Little
is known of the life habits of these snakes. The limited collection data available suggest that
they are semiaquatic forest dwellers. One form, brongersmai, makes an extensive incursion
into the savannah region of southeastern New Guinea. Altitudinal distribution is from sea
level to 2200 meters. Stomach contents indicate that most species feed on frogs, apparently
foraged from the forest floor. The presence of tadpoles and a few small, stream-living fish
(gobiids) in stomachs is evidence of semiaquatic habits. Ground skinks (lygosomines) are
known to be eaten by two species and geckos by a third.
Rhabdophis is distributed in eastern Asia from southeastern Siberia and Japan south
through China, the eastern Himalayas, Ceylon (unknown in peninsular India), the Malay
region east to Flores, Borneo, Sulawesi, and the Philippine Islands. These are primarily terres-
trial snakes, with some being semi-aquatic. None seem highly adapted to an aquatic lifestyle,
although many eat anurans and tadpoles, and some feed on fish and earthworms. They have
rear-fangs, and a few have highly toxic venom. Some Rhabdophis specialize in feeding on
toads and sequester the toad’s poisons in nuchal glands to deter predators. They are ovipa-
rous.
About 26 species are wide-
ly distributed in Asia. Although,
most are poorly known, Rhab-
dophis subminiatus is sometimes
referred to as a water snake, and
while I (JCM) have seen about a
dozen specimens in the field none
were in water. Instead, they were
in small bushes or on the ground.
De Lang (2017) reports similar
habits.
Natricids of the genera
Fowlea and Xenochrophis are dis-
Figure 25.1 The Red-necked Keelback, Rhabdophis submin-
tributed throughout many parts
iatus is one of a subclade of these snakes that has a nuchal
gland. The gland stores molecules obtained from toads. The
of Asia. They have a cylindrical
snake usesd those molecules to deter [Link] gland body, do not exceed 2 meters in
secretions can been in this photo Photography by Konrad length, and have a round pupil.
Mebert. These snakes are oviparous, diur

389
 25. Keeled Water Snakes

Figure 25.2. The Checkered

Keelback, Fowlea flavipunctatus
is a semi-aquatic member of this
clade. ,It is dirunal and moves in
and out of the water in search of
food. Photography by JCM.

nal, semi-aquatic, and are often found near water. Members of this group are opisto-
glyphous (rear-fanged), but just a few are considered dangerous to humans.
The snakes now classified under Fowlea and Xenochrophis have undergone many tax-
onomic revisions since their respective establishments in 1868 and 1864. The genus Fowlea
was erected by Theobald (1868), to accommodate a new species, Fowlea peguensis. In his Cat-
alogue of Snakes, Boulenger (1893) considered F. peguensis a junior synonym of Tropidonotus
punctulatus, based on maxillary dentition and nasal characters. In this publication, all species
currently under Fowlea and Xenochrophis were classified under the genus Tropidonotus, ex-
cept for X. cerasogaster.
In his book Herpetology of Japan and Adjacent Territory, Stejneger (1907) demonstrat-
ed how poor the taxonomic status of Tropidonotus was. On that basis, he assigned many
to Natrix, which were used in many other herpetological works after that. After Stejneger’s
publication in 1907, Smith (1943) considered bellula, punctulata, piscator, and trianguligera
to be in the genus Natrix and listed flavipunctata, asperrimus, and melanzostus as subspecies
of Natrix piscator.
During the mid-twentieth century, X. cerasogaster was the only species of the genus Xe-
nochrophis, which remained monotypic for many years until a revision of the genus Natrix
(sensu lato) by Malnate (1960) split the snakes into five genera: Macropophis, Fowlea, Rhab-
dophis, Natrix, and Amphiesma. In 1965, Malnate and Minton found that X. cerasogaster was
congeneric with Fowlea spp., which at the time consisted of the species piscator, punctulata,
and vittata. On this basis, they moved these snakes from Fowlea to Xenochrophis (by priority).
In 1988, Malnate & Underwood published another revision of this group, reclassifying what
were then known as Macropophis maculata, Sinonatrix trianguligera, and Sinonatrix bellula
into the genus Xenochrophis.
The Xenochrophis piscator complex (comprised of X. asperrimus, X. flavipunctulatus, X.
melanzostus, X. piscator, X. sanctijohannis, X. schnurrenbergi, and X. tytleri) was revised by
Vogel & David (2006), and subsequently by the same authors in 2012. In 2019, Purkayastha
et al. reallocated the X. piscator complex, along with X. punctulatus, into the genus Fowlea,
based on molecular data.
The diagnostic characteristics that distinguish these genera from other snakes are still
undefined, and the taxonomy of this group needs further study. In 1864, Günther erected
the genus Xenochrophis for type X. cerasogaster. They share a cylindrical body with a narrow,
elongated head, a round pupil, keeled scales in 19 rows, and a divided cloacal plate. The nos-
trils are laterally positioned on the nasal scales.
Based on the original genera descriptions by Günther (1864) and Theobald (1868), the
only differences that could be found were the number of dorsal scale rows (19 in Xenochro-
phis and 17 in Fowlea) and the shape of the pupil (round in Xenochrophis and vertical in
Fowlea). However, these characters are no longer diagnostic for the two genera.
The revision of natricids by Malnate (1960) reassigned many snakes previously classified
390
Aquatic Snakes

Figure 25.3. Characters used by Malnate (1960) to distinguish the genera of natricid snakes.
(Diagrams by Nathan Rusli, adapted from Malnate, 1960).

by Boulenger (1893) as Tropidonotus. Malnate divided them into Macropophis, Fowlea, Rhab-
dophis, Natrix, and Amphiesma, based on hemipenes structure (sulcus spermaticus simple
or forked), maxillary dentition (continuous gradual, continuous interrupted, or continuous
abrupt), number of maxillary teeth (more than 33 or less than 30), internasals (narrow or
broad), and position of nostrils (dorsolateral or lateral). These characters are illustrated in
Figure 25.1, adapted from Malnate (1960).
This group’s most recent taxonomic revision (Purkayastha et al. 2019) provided detailed
morphological descriptions for X. cerasogaster, distinguishing it from all other species of Xe-
nochrophis and Fowlea. However, the reallocation of many Xenochrophis species to the genus
Fowlea was based on molecular data rather than morphology. In their study, X. trianguligerus

and X. vittatus were found to be more closely related to the genus Rhabdophis. However,
limited data provisionally kept the species in the genus Xenochrophis, along with X. bellulus
and X. maculatus which were not investigated in the study. Therefore, further research is
necessary to ascertain the phylogeny of this group, especially for X. bellulus, X. maculatus,
X. trianguligerus, and X. vittatus. In addition, further research investigating morphological
characters to determine diagnostic features of genera and species is also needed.
Twelve species (seven Fowlea, and five Xenochrophis) are recognized: Fowlea asperrim-
us, F. flavipunctata, F. melanzostus, F. piscator, F. punctulatus, F. schnurrenbergeri, F. tytleri,
Xenochrophis bellulus, X. cerasogaster, X. maculatus, X. trianguligerus, and X. vittatus. The
St. John’s Keelback, Fowlea sanctijohannis, is not treated here as a valid species, following the
taxonomy by Vogel & David (2012), who stated that there was insufficient evidence to sup-
port the validity of the species.
Buff-striped Keelback
Amphiesma stolatum (Linnaeus, 1758)
Distribution and Habitat. Widespread in South and South-
east Asia: Forests and modified human-modified habitats, in-
cluding agricultural lands in Taiwan, China (Hainan, Hong
Figure 25.4. The distribution Kong, Fujian, Jiangxi), Sri Lanka, possibly the Andaman Is-
of Amphiesma stolatum. land, India (Kerala, Tamil Nadu, Andhra Pradesh, Karnataka,
Gujarat, Madhya Pradesh, Chhattisgarh, Orissa, Uttar Pradesh,
Assam, Bihar, Maharashtra, Arunachal Pradesh, Punjab, Him-
achal Pradesh, Jammu and Kashmir, Bhutan, Myanmar, Thai-
391
 24. Mountain Keelbacks

land, Vietnam (Hoa

Binh), Laos, Cam-
bodia, Nepal, Paki-
stan. Type locality.
America (in error).
Identification.
The head is distinct
from the body. The
head and anterior
body are orange,
red-brown or blue-
grey. Upper labials
are white, becoming
gray posteriorly and
edged with black:
body olive-green or
brown with black
spots or reticulated
cross bands with
two yellow or tan
longitudinal stripes.
Ventrals are white
with some dark
spots on the outer
edge of each ven-
tral scale and counts
141-151 on males
143-149 on females
in Taiwan (Mao,
1998).
Natural His-
tory. A mostly ter-
restrial or riparian
Figure 25.5a. Amphiesma stolatum. A. Photography by Konrad Mebert. Bottom. India. zone between wet-
Photography by Daniel Jablonski. land to grassland
snake, that feeds
on a variety of prey
(e.g. fishes, frogs, toads, grasshoppers, etc.) (Norval et al.,
2005, 2007, 2008) both in and out of the water. Usually close
to water and up to 2000 m ASL. It was often associated with
human-modified habitats as well as forests. Anurans and
lizards form the largest portion of their diet (Baruah et al.
2001). In Sri Lanka, it has been reported eating the Indian
Burrowing Frog, Sphaerotheca breviceps (Dissanayake and
Wellappuliarachchi 2016). Females are known to lay 1-14
eggs in May-September and remain with the eggs. When
Figure 25.5b Ventral view of in Taiwan disturbed, it will flatten its forebody (Pope, 1935)
(Photographer: Jean-Jay Mao).

392
Aquatic Snakes

Barred Keelback
Tropidonophis doriae Boulenger, 1897
Distribution and Habitat. Forest streams in New Guin-
ea east of the Vogelkop, absent from higher elevations in the
central mountain spine. The species also is present in the
Aru Islands. Elevational records for collection sites range
from sea level (Western Province, Papua New Guinea) to
1300 meters at Moroka in the Owen Stanley Range (Central
Figure 25.6. The distribution of Province, Papua New Guinea).
Tropidonophis doriae. Identification. Dorsum brown or gray-brown with
darker crossbands that usually are more distinct and broad-
er on the anterior body and narrower and/or less well defined on the posterior body. Bands
on the body 31-47, each two to six scales long and are separated by one to three scales. bands
may be interrupted into two or four series of dark spots on the posterior third or more of the
body length. Occasional specimens have narrow bands anteriorly and a reticulate pattern of
darkened scale edges posteriorly or are uniformly brown dorsally. The chin and throat are
cream-color or yellow; some lower labials may have dark edges. Juveniles are more boldly
patterned. Size. Males to 1049 mm total length, 794 mm SVL; females to 1113 mm, 829 mm
SVL. Tail length 24.3% to 28.5% of the total length. Dorsal scales in 17-17-17 rows; ventrals
134-153, subcaudals 71-90. Pits are present on head scales (Malnate and Underwood 1988).
Natural History. Inhabits primary
and secondary forest, mostly in riparian
areas. It is mostly aquatic and mainly diur-
nal. Diet is mostly frogs but will also take
fish, frogs’ eggs and tadpoles. It is ovipa-
rous with a clutch size of between 2 – 8
(Allison et al. 1998).

Figure 25.7. Tropidonophis doriae. from Papua New

Guinea. From Malnate (1988).

Elongated Keelback
Tropidonophis elongatus (Jan, 1865)
Distribution and Habitat. Inhabits the Moluccas Island
Group, Indonesia. Including the islands of Ambon, Irian Jaya,
Ceram, Ambon, Halmahera, Salawati, Numfoor and Biak;
possibly Buru and Obi). Type locality: Molucca; restricted to
Ambon by Malnate and Underwood 1988.
Identification. A slender-bodied snake with a long tail.
Figure 25.8. The distribution of Brown above with vertebral blotches or transverse bands.
Tropidonophis elongatu. Ventral surface with brown spots mainly on the outer edges
of ventrals. On a pale background. Dorsal scales are keeled
in 15-15-15 rows. Ventrals 155-175, subcaudals 72-109 (Mal-
nate and Underwood 1988).

393
 24. Mountain Keelbacks

Natural History. Poorly known species,

de Lang (2013) found one in a fast-moving
stream at 400 m ASL.

Mair’s Keelback
Tropidonophis mairii Gray, 1841
Distribution and Habitat. Creeks,
swamps, eucalypt forests, heaths, pastures,
parks, and suburban gardens are inhabted.
In New Guinea it occurs along the east coast
of the Gulf of Papua (Central Province). In
Australia it is found from the Cape York
Peninsula, Queensland, westward through
the northern portion of the Northern Ter-
ritory to the vicinity of Darwin, and south-
ward to the Clarence River in northern
New South Wales. Malnate and Underwood
(1988) considered this species polytyp-
Figure 25.9. Tropidonophis elongatus. Top RMNH ic and names the New Guinea population
5163, female; Seri, Ambon. Lower: Lateral view west of the Gulf of Papua T. m. plumbeus
of the head and posterior body Bottom. RMNH
and the Australian population T. m. mairii.
18607b, male; Aitinjo, Vogelkop, Irian Jaya. From
Type locality: for T. m. maririi New Holland
Malnate (1988).
(= Australia). Neotype locality: Port Essing-
ton, NT; Type locality for T. m. plumbeus
Katow (now Mawatta), Western Province (Malnate and
Underwood 1988).
Identification. The dorsum is a uniform brown or
gray, that may have scales with darker edges. A dark gray
vertebral stripe or vertebral
area with vague lateral definition often is present. A
Figure 25.10. The distribution of series of small dark spots may be present on the third,
Tropidonophis mairii. fifth and vertebral scale rows. The spots may be enlarged
on the forebody. The head and nape. The top of the head
may be pale with the plates, especially the parietals, mot-
tled or marbled with dark. Paired light parietal spots may be present. The upper labials are
pale with the upper edges of all plates darkened; the last two labials may be dark. A postocular
dark stripe is present, often narrow, sometimes prominent, occasionally absent. Dorsal scales
keeled and in 15-15-15 rows. Ventrals males 133-166, in females 133-156. Size. Males to 728
mm total length, 585 mm SVL; females to 814 mm, 712 mm s-v. Tail length is 23.2% to 30.8%
of the total length (Malnate and Underwood 1988).
Natural History. Diet seems to be mostly frogs, will also feed on lizards. Active day and
night. Capable of eating the introduced and highly toxic Cane Toad. It will climb into vege-
tation.
Females lay 2-18 eggs in June. Females the selected substrates with more moisture for
egg deposition produced young with larger body size at hatching (Brown and Shine, 2004).
Brown and
Shine (2005) tested 42 gravid Tropidonophis mairii from tropical Australia for preference

394
Aquatic Snakes

Figure 25.11. Tropidonophis mairii

Photography by Shawn Scott.

of egg laying sites. Given the choice of laying their eggs in a nest containing empty eggshells of
their own species, empty eggshells of their predator, the Slatey Grey Snake, (Stegonotus cucul-
latus), or a control with no eggshells. Six of the females chose the control box (no eggshells), 16
used the box with keelback eggshells, and 20 laid their eggs in nest boxes with the slatey-grey
snake implying that the presence of eggshells may prompt the snake to lay eggs there because
other successful hatchlings took place at that site.

Many-scaled Keelback
Tropidonophis multiscutellatus (Brongersma, 1948)
Distribution and Habitat. New Guinea, the islands of
Batanta and Salawati off the western tip of the Vogelkop. It is
also present on Noemfor and Biak Islands in Geelvinck Bay,
and Liki, Manam, and Karkar Islands off the north coast of
New Guinea. On the mainland of New Guinea multiscutellatus
Figure 25.12. The distribution is distributed from the western tip of the Vogelkop to Fife Bay,
of Tropidonophis multiscutella- near the tip of the southeastern peninsula, but absent from
tus. the higher elevations of the central massif and the savannah
region in the southeast. Known from elevations from 15 m to about 1440 m. Type locality:
Alkmaar, Lorentz River, Irian Jaya (Malnate and Underwood 1988).
Identification. The dorsum uniformly brown or gray, may have dark, edged scales. A pat-
tern of dark spots is common. The spots may be irregularly spaced or linear on the third, fifth
and vertebral scale rows; they may be small or large, obscure, or prominent, those on the fifth
and vertebral rows and may fuse to form narrow crossbars. The venter is light, gray, or brown
at the sides, with short, dark transverse streaks at the outer corners of the ventrals. Dorsal

395
 24. Mountain Keelbacks

scales keeled and in 15-15-15 rows. Ventral scales 231-253 in males, and 228-254 in females.
Size. Males to 808 mm, total length, 583 mm SVL; females to 951 mm, 710 mm SVL. Tails are
25.3-34.2% of the total length.
Natural History. Diet includes fish (gobies). Females produce clutches of 2-10 eggs (Mal-
nate and Underwood 1988).

Figure 25.13. Tropidonophis

multiscutellatus Photogra-
phy by Nick Baker.

Negros Spotted Water Snake

Tropidonophis negrosensis (Taylor, 1917)
Distribution and Habitat. Stream and their vicinity on the
central Philippine islands of Mindoro, Masbate, Panay, Sico-
gen, Pan de Azucar, Negros and Cebu. At altitudes of 411-823
meters. Type locality: Canlaon Volcano, Occidental Negros,
Philippines (Malnate and Underwood 1988).
Identification. Head and nape are dark gray or black, the
Figure 25.14. The distribution
of Tropidonophis negrosensis.
snout is lighter. Paired light spots often are present on the pa-
rietals. Upper labials are pale, darkened at their upper borders

Figure 25.15. Tropidonophis negrosensis, Twin Lakes Natural Park, Negros Oriental, Philippines.
Photography by Chris Cafer.

and vertical sutures; the last scale often is heavily pigmented. A black stripe extends from the
lowest postocular to the corner of the mouth but may be obscure or absent. The dark area on
the nape is followed by three or four dark crossbands enclosing light lateral spots; occasion-
ally, the bands are broken into large dark spots. A narrow, light vertebral stripe sometimes is
396
Aquatic Snakes

present on the neck, continuing through the anterior quarter of the body. The chin and throat
are pale. Two rows of black spots along the sides of the venter. Dark speckling also may be
present at the sides. Each subcaudal bears a single bold dark spot; scattered gray flecks may
occur along the midline of the scutes. Dorsal scales keeled and in rows 19-19-17. Ventrals in
males 252-262, in females 245-261, subcaudals in males 92-100, in females 87-98. Size. The
largest male 775 mm total length, 554 mm SVL; the largest female 784 mm, 586 mm SVL. Tail
length 24.1-30.5% of the total length (Malnate and Underwood 1988).
Natural History. Diet includes frogs and their tadpoles. Clutches of 3-7 eggs (Malnate
and Underwood 1988).

Speckled Belly Keelback

Tropidonophis punctiventris (Boettger, 1895)
Distribution and Habitat. A semiaquatic snake that uses
slow-moving shallow streams that transect rainforests. Halma-
hera, Indonesia. Type locality: Konorra, Halmahera (Subasli
2020).
Identification. A brown or olive dorsum with a series of pale
Figure 25.16. The distribu- spots on scale rows 4-6. Ventral is pale with brown spots. The
tion of Tropidonophis punc- slender body with a long tail. Dorsal scales keeled in 15-15-15
tiventris. rows. Ventrals 145-160, cloacal plate divided, subcaudals 74-75.
Size. A small snake, 320 and 460 mm long total length for the
two known specimens (Malnate and Underwood 1988).
Natural History. Nothing known.

Figure 25.17. Tropidonophis

punctiventris. The male, ho-
lotype from Soa Konorra,
Halmahera. From Malnate
(1988).

397
 24. Mountain Keelbacks

Truncated Keelback
Tropidonophis truncatus (Peters, 1863)
Distribution and Habitat. Small streams on Salawati, Halma-
hera, Bacan, and Ternate Indonesia. Type locality: Dodinga, auf
der Mitte der Insel Djololo [Halmahera] (Malnate and Under-
wood 1988).
Identification. A slender cylindrical body with a moderately
long tail. Dorsum brown or dark grey with a network of black
Figure 25.18. The distribu- stripes formed by the dark edges on the scales. Maybe a uniform
tion of Tropidonophis trun-
pattern. Venter with a pale background with dark speckling. Dor-
cates.
sal scales in 15-15-15 rows, most rows keeled, except the vertebral

Figure 25.19. Tropidono-

phis truncates A female
syntype from Halmahera.
From Malnate (1988).

and para vertebral rows that are smooth. Ventrals 132-159, cloacal plate single, subcaudals
36-51. Size. Maximum length 560 mm total length. Tail about 0.2 of the total length (Malnate
and Underwood 1988).
Natural History. Nothing known.

White-lined Water Snake

Rhabdophis auriculatus (Günther, 1858)
Distribution and Habitat. Along streams in the Philip-
pines (Basilan, Bohol, Leyte, Samar, Mindanao). Type local-
ity: Philippines. A polytypic species. Rhabdophis auriculata
myersi Leviton 1970; Type locality Davao Province, Mind-
anao Island. Differs in subcaudal counts than the nominate
subspecies.
Figure 25.20. The distribution of Identification. A dark brown to olive green dorsum
Rhabdophis auriculatus. with a pale vertebral stripe on each side of this stripe is a
series of dark elongate spots, immediately below those is a
series of pale gray spots, followed by another series of dark
spots below. The first two scale rows are pale pink. The ventral scales are black with white
lateral spots.
Head short, blunt, very distinct from anterior body; rostral visible barely visible from
above; parietals longer than frontal, in contact with one postocular; two nasals, the posterior
highest; loreal higher than wide, in contact with second and third labials; preocular single,
three postoculars; temporals 1+3; eight upper labials, third, fourth, and fifth border the orbit.
Five lower labials touching anterior chin shields, which are shorter and broader than posteri-
or. Dorsal scales keeled and in 17 rows. Ventrals 150-158; subcaudals 75-91. Size. To at least
398
Aquatic Snakes

Figure 25.21. Rhabdophis auricu-

latus from Ormoc, Leyte, Philip-
pines. Photography by Stefanie
de Win.

524 mm total length, tail 145 mm (Taylor, 1922). This seems to be a small snake compared
with other members of the genus.
Natural History. Diet includes ranid frogs in all life stages (eggs, tadpoles, adults).

Zigzag-lined Water Snake

Rhabdophis lineatus (Peters, 1861)
Distribution and Habitat. Coastal freshwater swamp edg-
es in the Philippines (Basilan, Bohol, Mindanao, Samar, Leyte)
Type locality: Loquilocun, Insel Samar.
Identification. A red-brown dorsum with scattered black
spots that look like smudges of black on scales. A white stripe
on the upper labials extends to about the tenth ventral, bordered
Figure 25.22. The distribu- by black above. Dorsal scales keeled and in 19 rows, ventrals 132-
tion of Rhabdophis lineatus.
142, subcaudals 64-71. Size. To at least 625 mm total length, tail
159 mm.

Figure 25.23. Rhabdophis lineatus from Taft, Eastern Samar, Philippines.

Photography by Jeff Weinell.

399
 24. Mountain Keelbacks

Blueneck Keelback
Rhabdophis rhodomelas (Boie, 1827)
Distribution and Habitat. Brunei Darussalam, Myanmar,
Cambodia, India, Indonesia, Laos, Malaysia, Singapore, Thai-
land, Vietnam. Usually occurs in lowland areas but can be found
up to 1200m asl. Associated with aquatic habitats, such as rivers
and swamps. Type locality: Java, Indonesia.
Identification. Body robust and cylindrical, with a long
Figure 25.24. The distribu- tail. Head distinct from neck. Eye large with round pupil, di-
tion of Rhabdophis rhodome-
ameter of eye greater than distance between eye and edge of lip.
las.
Loreal present, one preocular, and three or four postoculars.
Nine upper labials (rarely 8 or 10), 4th to 6th (rarely 5th-7th)
touching eye. Scales in 19 rows on
midbody, strongly keeled. Ventrals
130-145,cloacal plate divided, sub-
caudals 67-99.
Dorsum olive grey or brown,
with distinct or indistinct dark tri-
angles. On the posterior half of the
body, there are often small, light spots
positioned laterally, in the middle of
the dark triangles. Juveniles tend to
have brighter coloration, and many
specimens have bright red markings
laterally, alternating with the trian-
gular pattern. The head is olive, with
white labials and black sutures on the
upper labials. Ventrals pale with no
markings.
Natural History. Mostly aquat-
ic. It occurs in disturbed areas such
as canals, ditches, and fishponds. It
is diurnal, actively hunting fish and
frogs. At night, specimens have been
observed resting on vegetation over-
hanging small streams and large,
fast-flowing rivers. Individuals have
also been recorded resting on water
hyacinths at night, and twigs emerg-
ing from the water in a large man-
made lake. When captured, they will
often secrete a foul-smelling musk.
Oviparous, with a clutch size of
about 5-8 eggs.
Figure 25.25. Rhabdophis
rhodomelas. Photography by
Mediyansyah.

400
Aquatic Snakes

Japanese Tiger Keelback

Rhabdophis tigrinus (Boie, 1826)
Distribution and Habitat. Coastal wetlands to montane
streams, including roadside ditches, ponds, and rice paddies
on Honshu, Kyushu, Shikoku, and the Satsunan Islands of the
Ryukyu group, Japan.
Identification. Dorsal scales are keeled in 19-17-17 rows
(sometimes to 15 rows post body). Ventral scales are 148-160
Figure 25.26. The distri-
in males and 153-162 in females; subcaudals in males 57-73, in
bution of Rhabdophis ti-
grinus.
females 52-65. Size. Males reach at least 730 mm SVL; females
reach at least 780 mm SVL.
The Japanese populations have
prominent geographical variations in
coloration. Goris (1971) found popu-
lations from Kanto (eastern Honshu),
Kinki (western Honshu), and Kyushu
have distinct dorsal color patterns.
The Kanto population expresses
many dark dorsal blotches on a red
ground color; the Kinki type has ob-
scure dorsal blotches, and the ground
color is not red; and the Kyushu type
has large, dark dorsal blotches on red
ground color.
Natural History. The diet con-
sists mainly of anurans, most often
tadpoles, toadlets (Bufonidae) and
froglets. It uses chemical and visu-
al cues to locate prey (Tanka 2002).
Home range size showed consider-
able individual variation, with home
range area ranging from 1.3 to 11.0
ha in a temperate forest in Kyoto, Ja-
pan (Kojima and Mori 2014).
Venom and Poison. Severe en-
Figure 25.27. Rhabdophis tigrinus. venomation by this species was
Photography by Yasunori Koide.
reported by Mittleman and Goris
(1974); symptoms included delayed,
spontaneous, superficial hemorrhaging and profound impairment of normal blood coagula-
tion. Severe internal hemorrhaging and hemolysis were present in two cases. Human deaths
have resulted from envenomation by this snake (Mittleman and Goris 1978). Rhabdophis
is one of three genera with nuchal glands containing cardiotonic steroid toxins (bufadien-
olides). Rhabdophis tigrinus (and R. lateralis and R. formosanus) sequesters bufadienolides
from its toad prey and stores them in the nuchal glands as a defensive mechanism (Takeuchi
et al. 2018). Gravid females with high nuchal gland toxin levels pass the compounds to their
offspring. Snake hatchlings thus also benefit from the toad-derived defenses found in their
parent.

401
 24. Mountain Keelbacks

Taiwanese Tiger Keelback

Rhabdophis formosanus (Maki, 1931)

Distribution and Habitat. Montane streams, ponds, and

nearby riparian zones, forest edges, and roadside ditches
within elevations 1600 to 3000 m in Taiwan (Mao et al., 2012)
Identification. Head is distinct from neck; body is moderate-
ly stout. The snake has a pair of nuchal glands and enlarged
rear fangs. Crown of head olive at least anteriorly; there is a
Figure 25.28. The distribution wide, curved transverse band of yellow on the nape, to which
of Rhabdophis formosanus. a black cross band adjoins in the front and rear. The upper
labials are yellow with black sutures, the black areas along
the sutures below eye
are broad. Upper body
and tail bear green-yel-
low-orange and black
spots arranged in five
alternating rows, creat-
ing a checkered appear-
ance. Ventral head is
white. Ventral scales on
body and tail are black
with (irregular) pos-
terior margins of light
green-yellow. Cloacal
scale is divided and sub-
caudals are paired. Dor-
sal scales are keeled, in
19-17-17 rows (some-
times to 15 rows post
body), Ventral scales
are 148-160 in males
and 153-162 in females.
Rhabdophis formosa-
nus has high subcaudal
Figure 25.29. Rhabdophis formosanus. Taiwan. counts (≤ 80 in males, ≤
Photography by Yu Sing Lin. 77 in females). Thus, sub-
caudal counts could dis-
tinguish R. formosanus from R. tigrinus and R. lateralis, except for the Kyushu populations of
R. tigrinus with partially overlap of the ranges (Ota & Mori, 1985; Ota et al., 1999). Males reach
at least 730 mm SVL, females reach at least 780 mm SVL. Total length up to 100 cm.
Natural History. Diet is primarily anurans but will also feed on fish and other snakes.
Females produce 8-47 eggs per clutch in summer; hatchlings measure about 160 mm in total
length. Defense behavior includes flattening the anterior body into a narrow hood.

402
Aquatic Snakes

Chinese Tiger Keelback

Rhabdophis lateralis Berthold, 1859
Distribution and Habitat. Coastal wetlands to montane
streams, including roadside ditches, ponds, and rice paddies in
China, North Korea, Russia, South Korea, and Vietnam.
Identification. Dorsum green ventral surface white ante-
riorly and dusky green posteriorly. Narrow black bars on the
Figure 25.30. The distribution of labial scales. Black transverse bands extend down the body
Rhabdophis lateralis. and fade on the tail. Spaces between bands are red anteriorly
and become green posterior-
ly. Dorsal scales are keeled, in
19-17-17 rows (sometimes to
15 rows post body). Ventral
scales are 148-160 in males
and 153-162 in females; sub-
caudals in males 57-73, in fe-
males 52-65. Size. Males reach
at least 730 mm SVL, females
reach at least 780 mm SVL.
Pope (1935) considered the
longest specimen to be 998
mm.
Natural History. Li et
al. (2008) found hibernation
occurs from the middle of
Figure 25.31. Rhabdophis lateralis from Licun Village, Fenghua October to following April.
County, Zhejiang, China. Photography by Jian Wang. Most activity occurs from May
to August. The diet is composed
of anurans. Reproduction occurs from the middle of June to the middle of July in the Beijing
area. Clutches contain six to 22 eggs, and the incubation period is 41 to 43 days.

Olive Keelbacks, Atretium, Fowlea, and Xenochrophis

The genus Atretium contained two species that were not monophyletic. They appear in
two different places in a tree with Xenochrophis and Fowlea. The Yunnan Olive Keelback is
the sister to Fowlea asperrimus and A. schistosum is the sister to a clade containing both Xe-
nochrophis and Fowlea (Zaher et al. 2019).

Olive Keelback
Atretium schistosum (Daudin, 1803)
Distribution and Habitat. Ponds, marshes, flood agricultural land, and swamps of South
Asia, Sri Lanka, India, Bangladesh, and Nepal. Possibly in Bhutan. Peninsular India south of
latitude 15 degrees north and along the east coast to Uttarakhand. Up to 1000 m ASL. Absent
from most of North India. Type locality: None given.
Identification. A uniform olive green to brown above with a yellow to orange venter. It is
a small, robust snake with a narrow head, short snout, and slit nostrils placed relatively high.

403
 24. Mountain Keelbacks

Figure 25.32. The distribution of

Atretium schistosum.
It is sometimes tinged with pink
or purplish on the flanks. The tail
length is one-third to one-fourth
of the total length. Dorsal scales
keeled in 19-19-17 rows; ventrals
129-160, cloacal plate divided,
subcaudals paired 53-85. Upper
labials number eight, third and
fourth border the orbit (or nine
upper labials with the fourth and
fifth bordering the orbit). Spec-
imens from South India have a
distinct reddish line along the
fifth and sixth or the fourth and
fifth scale rows up to the vent.
This stripe is brighter in males.
The females are between 700 and
750 mm long, and the males are
between 500- and 600-mm. Max-
imum size 870 mm total length
(Schleich & Kastle 2002; Wall
1921).
Natural History. Diet in-
cludes fish, tadpoles, frogs, and
mosquito larvae. Oviparous
clutches of 12-30 eggs laid be-
tween December and March.
Figure 25.33. Atretium schistosum. Avrajjal Gosh.

Bar-necked Keelback
Fowlea schnurrenbergeri (Kramer, 1977)
Distribution and Habitat. Slow rivers and streams, marshes, swamps, ponds, and lakes in
Nepal and northern India. Type locality: Devanandpur, Bhairawa, Nepal.
Identification. Characterized by a small checkerboard dorsal pattern; a broad, straight
crossbar on the nape; two well-defined subocular streaks, the posterior one extending from
the eye to the corner of the mouth but not meeting the nuchal crossbar; and ventral and sub-
caudal scales with broad, dark margins (Vogel & David 2006). Dorsal scales are keeled and
in 19:19:17 rows. In males, ventral scales are 132-139, and subcaudals are 71-80; in females
ventral scales are 141-152, and subcaudals are 61-79. Males average 600 mm in total length,

404
Aquatic Snakes

and females average 900 mm in average total length. Inter-

nasals are narrow anteriorly; upper labials 9-10, 4-5 border
the orbit; preocular one; the loreal is single; postoculars
2-3; sometimes a single subocular; temporal 2+2 or 2+3 or
rarely 1+2.
Zaher et al. (2019) found this species to be the sister to
Figure 25.34 The distribution of all the other members of the genus Atretium and Fowlea.
Fowlea schnurrenbergeri Natural History. A poorly studied species.

Yunnan Olive Keelback

Fowlea yunnanensis
Anderson, 1879
Distribution and Habitat.
Western Yunnan, China. Type
locality: Lianghe, Yingjiang
County; and Husa, Longch-
uan County; both in Yunnan
Province, southern China.
May also be present in Myan-
mar. To elevations of 1500 m
ASL.
Identification. Color uni-
form dark olive-brown above;
yellow beneath. Snout rather
pointed; internasals triangular
and form a short suture with
the rostral but may be exclud-
ed from the rostral by a small
azygos scale which is wedged
between them. The prefron-
tals are broader than long,
and about the same width as
the internasals. Parietals lon-
ger than the vertical, round-
ed behind, sometimes with a
portion separated from their
hinder extremities. Loreal
subquadrangular, higher than
broad. One preocular reach-
ing to the upper surface of the

Figure 25.35. Fowlea schnurren-

bergeri. Photography by Avrajjal
Ghosh.

405
 24. Mountain Keelbacks

head. Three postoculars, the lowest in contact with three labials.

Temporals 2+3. Nine upper labials, the fourth and fifth border
the orbit. Twenty-three maxillary teeth, increasing in length pos-
teriorly. Ventrals 144- 154; subcaudals 62-85. Dorsal scale rows
keeled in 19 rows. Yunnan specimens differ from the Indian spe-
cies in the narrow suture between the anterior frontals and rostral;
Figure 25.36. The distribu-
tion of Fowlea yunnanensis.
in the divided posterior frontals; in the presence of nine upper
labials, and in having slightly larger preoculars, and occipitals.
Chen et al. (2021) note that Atretium yunnanensis was orig-
inally recognized as a variant of Atretium schistosum. It was subsequently elevated to full
species status and has long been assigned to the natricine genus Atretium. They used mor-
phological and genetic evidence and found Atretium yunnanensis nested within the genus
Fowlea Theobald, 1868.
Natural History. This species inhabits paddy fields, ponds, and river valleys between 800
m and 1500 m ASL. It is diurnal, preys on frogs, fish, and rats. Defense behavior includes
biting repeatedly and violently.

Boulenger’s Keelback
Fowlea asperrimus (Boulenger, 1891)
Distribution and Habitat. Endemic to Sri Lanka. Occurs in elevations up to 1000m asl,
usually living in or near water bodies such as flooded rice fields,
lakes, marshes, ponds, and rivers. Type locality: Ceylon.
Identification. Body robust and cylindrical, tail length mod-
erate. Head distinct from neck. Eye small with round pupil, di-
ameter of eye less than or just reaching greater than distance
between eye and edge of lip. Loreal present, one preocular, and
Figure 25.37. The distribu- three (rarely two) postoculars. Nine upper labials (rarely 10 or
tion of Fowlea asperrimus. 11), 4th and 5th (rarely 5th and 6th) touching eye; 10 (rarely 9 or
11) lower labials. Scales in 19 rows on midbody, strongly keeled.
The keels form sharp lines posteriorly, except for the outer rows,
which are smooth. Ventrals 127-142, cloacal plate divided, subcaudals 75-92. Maximum total
length 890 mm for females; males much shorter (maximum length females / males 1.41),
longest male 630 mm. Relative tail length sexually dimorphic: tail 28.3–31.4 % of total length
in males, 25.7–28.9 % in females (Vogel
and David 2012).
Dorsal color pale with broad,
dark, confluent blotches anteriorly,
which become separated on the pos-
terior part of the snake. Blotches num-
ber 22-32. The anterior part can also
be described as dark with incomplete

Figure 25.38. Fowlea asperrimus from

Kalawana, Ratnapura, Sri Lanka. Pho-
tography by Susan Meyer.

406
Aquatic Snakes

crossbands, larger at the base, sometimes forming a triangular shape. The head is dark ol-
ive dorsally, with two narrow black streaks on the sides, located under and behind the eye.
Light-colored ventrals, outer edges clouded with dark grey (based on Vogel and David 2012).
Natural History. Mostly aquatic, presumably diurnal. Has been recorded from a creek
near paddy fields and observed swimming in a river during the day. Oviparous.

Yellow-Spotted Keelback
Fowlea flavipunctata (Hallowell, 1860)
Distribution and Habitat. India, Thailand, Myanmar, South China, Taiwan, West Malay-
sia, Laos, Cambodia, Vietnam, Bangladesh. Occurs in slow-moving streams, rivers, ponds,
swamps, and marshes. It is also present in modified habitats such as paddy fields and ditches.
Some working rotavator on paddy
fields make this species vulnera-
ble to opportunistic avian forag-
ers in Taiwan (Mao and Norval,
2013). Type locality: Canton River,
Kwangtung, China (now Guang-
dong).
Identification. Body robust
and cylindrical, tail length moder-
ate. Head distinct from neck. Eye
large with round pupil, the diame-
ter of eye greater than the distance
between eye and edge of lip. Loreal
present, one preocular, and three or
four (rarely two) postoculars. Ten
upper labials (rarely 8), 4th and 5th
(rarely 4th, or 5th and 6th) touch-
ing eye; 9-11 lower labials. Scales in
19 rows on midbody, rather weakly
keeled anteriorly and more strong-
ly keeled posteriorly. Ventrals 120-
143, and present sexual difference
on males 124-132 (n=20) and fe-
males 138-143 (n=28) in Taiwan
(Mao, 1998), anal divided, subcau-
dals 60-91 (Fig. 25.41b).
Dorsal color varies between
different shades of olive, brown,

Figure 25.39. (to right)_Fowlea flavi-

punctatus. Top: Sakaeret, Thailand.
Photography by Harry Ward-Smith
Middle: Laos. Photography by Daniel
Jablonski. Bottom Hong Kong. Pho-
tography by Artur Tomaszek.

407
 24. Mountain Keelbacks

orange-red or grey. Clear, dark spots arranged in 6-7 alternating

rows, producing a checkered pattern; more distinct anteriorly and
larger laterally. Dorsal scales often have pale margins. In life, some
specimens have vivid red or orange markings on the flanks.
Natural History. Mostly semi-aquatic, diurnal. Like most keel-
backs, its diet consists of fish (e.g.: snakehead fish [Channa asiati-
ca] and frogs (e.g.: Heymonsi’s Narrow-mouthed Toad [Microhyla
Figure 25.40. The distribu- heymonsi]) (Norval et al., 2010). In Taiwan, this species usually
tion of Fowlea flavipuncta- shared habitat (sympatric) with Amphiesma stolatum, Hypsiscopus
tus murphyi, Myrrophis chinensis, and Trimerodytes annularis but dif-
ferences in spatial or active time (JJ Mao, unpublish
data). Some literature and observations indicated
that the krait is an obvious predator of this snake
(Mao, 1970; Mao et al., 2006; Mao et al., 2010).
Oviparous, clutch size are 23-112 eggs, copulation
in Autumn (October to early November), par-
turition in early May, and then the egg takes two
months to hatch (JJ Mao, unpublish data).

Javanese Keelback
Fowlea melanzostus (Gravenhorst, 1807)
Distribution and Habitat. Endemic to Indone-
sia, specifically the Java, Sumatra, and Bali islands.
It occurs in lowland habitats with varying degrees
of disturbance. These habitats often have still or
slow-moving bodies of water, such as paddy fields,
ditches, or fishponds. Type locality: Batavia, Java
(now Jakarta).
Identification. Body robust and cylindrical,
tail length moderate. Head distinct from neck. Eye
average-sized with round pupil, the diameter of
eye roughly equal to the distance between eye and
edge of lip. Loreal present, one preocular, and three
(rarely two) postoculars. Nine upper labials (rarely
10), 4th and 5th (rarely 4th, or 5th and 6th) touch-
ing eye; 10 (rarely 9 or 11) lower labials. Scales in
19 rows on midbody, strongly keeled except for the
outer 2-3 rows. Ventrals 128-143, cloacal divided,
subcaudals 66-83.
Dorsal color varies between shades of yellow,
Figure 25.41. (top)_The newly hatchling snake
of Fowlea flavipunctatus in JJM lab. of Taiwan. light brown, grey, or olive. There are two forms;
(middle) The right side hemipenis of Fowlea blotched and striped. The blotched form has four
flavipunctatus of Taiwan (Photographer: rows of large black blotches on the neck and one
Jean-Jay Mao). (bottom) The ventral view of vertebral blotch behind the neck. These are fol-
large female Fowlea flavipunctatus of Taiwan. lowed by five rows of smaller blotches, which often
Photography by Jean-Jay Mao. fade posteriorly. The striped form has four dark
stripes starting on the neck and a vertebral stripe

408
Aquatic Snakes

starting behind the neck. Like the blotched form, these

markings also fade posteriorly. Some dorsal scales often have
dark margins. In life, some specimens have bright red mark-
ings laterally. The head is darker than the body, and has dark
streaks below and behind the eye. Ventrals are pale yellow or
cream, often with black outer edges.
Figure 25.42. The distribution of Natural History. Mostly aquatic, in rice paddies with
Fowlea melanzostus. shallow water, in ditches, and on the edges of fishponds. It
is primarily diurnal but occasionally observed hunting at
night. It is an active hunter with a diet consisting of fish and frogs. When captured, this spe-
cies secretes a strong musk as a defensive mechanism to deter predators. Oviparous.

Figure 25.43. Fowlea melanzostus.

Photography by Muhammad Azib.

Checkered Keelback
Fowlea piscator (Schneider, 1799)
Distribution and Habitat. Pakistan, Sri Lanka, India, Bangladesh, Bhutan, Nepal, Myan-
mar, Thailand, Laos, Vietnam, Malaysia, Singapore, and China. It occurs in aquatic habitats
such as slow-moving rivers, lakes, and marshes. Also common in modified habitats such as
canals and fishponds. Type locality “East Indies” (unspecified).
Identification. Body robust and cylindrical, tail length moderate. Head distinct from
neck. Eyes average size with a round pupil, eye diameter equal to, or greater than the dis-
tance between eye and edge of the mouth. Loreal present, one preocular, and three or four
postoculars. Nine upper labials (rarely 8 or 10), 4th and 5th (rarely only 4th or 5th) touching

409
 24. Mountain Keelbacks

eye; 10 (rarely 9 or 11) lower labials. Scales in 19 rows on midbody, weakly keeled except for
the outer 2-4 rows. Ventrals 128-154, cloacal divided, subcaudals 68-96 (based on Vogel and
David 2012).
Base color on dorsum shades of light or dark brown, grey,
yellow, or olive. The dorsal pattern usually consists of 5-7 rows
of alternating dark blotches, forming a checkered pattern. The
dark blotches may vary in size, and sometimes lighter speck-
ling is present. In addition, it has narrow dark streaks below
and behind the eye. Ventrals are usually pale yellow or cream,
Figure 25.44. The distribution often with lateral cloudy spots.
of Fowlea piscator.

Natural History. Mostly

aquatic and diurnal. It is an ac-
tive hunter with a diet consist-
ing of fish and frogs. Patel et al.
(2017) found a specimen that
had died trying to eat the fish
Anabas tertudineus. Presum-
ably, the fishes’ spines prevent-
ed the snake from swallowing
the fish and caused it to suffo-
cate. Kalki (2020) found this
snake feeding on the eggs of the
Malabar Gliding Frog (Rha-
cophorus malabaricus). When
threatened, it flattens its neck,
mimicking a cobra. Autonomy
of the tail has also been record-
ed in this species. Oviparous,
average clutch size of 30-70
eggs.

White-Spotted Keelback
Fowlea punctulatus
(Günther, 1858)
Distribution and Habitat.
Myanmar, Thailand. It occurs
in mangrove habitats and tidal
Figure 25.45. Fowlea piscator. A.
India Avrajjal Ghosh. B. Sakaeret,
Thailand Harry Ward-Smith. C.
Pha Nam Yod Waterfall, Tha Yang
District, Phethaburi Province,
Thailand. Photography by Rush-
en Bilgin.

410
Aquatic Snakes

marshes. Type locality: Unknown.

Identification. Body robust and cylindrical, tail length mod-
erate. Head distinct from neck. Eye small, with round pupil, the
diameter of eye equal to, or less than the distance between the
eye and the edge of mouth. Loreal present, one preocular, and
Figure 25.46. The distribution two or three postoculars. Nine upper labials (rarely 10), 4th and
of Fowlea punctulatus. 5th (rarely 5th and 6th) touching eye; 10 (rarely 9 or 11) lower
labials. Scales in 17 rows on midbody, smooth or weakly keeled.
Ventrals 142-154, cloacal divided, subcaudals 62-83.
Dorsum black with small pale dots, forming incomplete crossbands throughout the dor-
sum. Upper labials pale cream or white, followed by a narrow “V” pattern on the neck. Pale
ventrolateral stripes present, bordered in black. Ventrals light-colored, often with dark mar-
gins.
Natural History. Mostly aquatic, found in brackish habitats such as tidal palm forest and
mangrove swamps. Preys on fish and frogs, presumably predated by larger snakes, monitor
lizards, large fish, and wading birds. Oviparous.

Figure 25.47. Fowlea punctulatus

Photography by Montri Sumontha.

Tytler’s Keelback
Fowlea tytleri (Blyth, 1863)
Distribution and Habitat. The Andaman Islands, and pos-
sibly Nicobar Islands. Common in forested areas, presumably
in or near bodies of water. Type locality: Andaman Island.
Identification. Body cylindrical, tail length moderate.
Head distinct from neck. Eyes with a round pupil, and the
Figure 25.48. The distribution diameter of eye roughly equal to the distance between eye
of Fowlea tytleri. and edge of mouth. Loreal present, one preocular, and three

411
 24. Mountain Keelbacks

(rarely 4) postoculars. Nine upper labials, 4th and 5th touching eye; 9 (rarely 10) lower labials.
Scales in 19 rows on midbody, strongly keeled. Ventrals 131-145, cloacal divided, subcaudals
76-86.
Base color of dorsum pale brown with variable markings; striped, blotched, or both. Ju-
veniles have narrow pink-brown stripes, often broken into a series of spots posteriorly. Some
color forms have a wide “V” pattern on the neck. It is cream ventrally.
Natural History. Mostly aquatic, diurnal. Preys on frogs and fish. Oviparous.

Burmese White-Barred Keelback

Xenochrophis bellulus (Stoliczka, 1871)
Distribution: As far as currently known, X. bellulus is
endemic to southcentral Myanmar in the states of Yangon
and Bago (Lee et al. 2018).
Identification. Based on Lee et al. 2018 [two males and
two females] and an adult male specimen collected by GK,
SMF 106048): The head is olive-brown with contrasting
cream-colored vertical bars on the upper labials of which
Figure 25.49. The distribution of
Xenochrophis bellulus. one in front of the eye and one just posterior to eye reach
to upper margin of eye. Two other such vertical bars are
present in the lateral nuchal region. The body is dark grayish olive to black with cream-col-
ored checkering and anterior crossbands arranged on both sides, not reaching dorsal surface.
The eyes are large, pupil round, nostril directed dorsolaterally; prefrontal single; internasals
two; one loreal; one preocular; three postoculars; temporals 1+1 or 1+2; upper labials usually
nine, sometimes ten, with 4th–6th bordering orbit. Lower labials nine or ten, with the 1st–6th
contacting the anterior chin shields. Body scales keeled, usually in 19–19–17 rows, some-
times in 19–19–19 rows when transition from 19 to 17 rows happens posterior to midbody
(as in GK-7061 that has this transition at ventral scale 70 or 54.2% of SVL); 2–4 preventrals;
ventral scales 134–139 in males, 135–143 in females, all unkeeled; cloacal plate divided; sub-
caudals 71–79 pairs in males, 70–72 in females. SVL 323-429 mm in males, 316-321 mm in
females; tail 27.5–28.8% of total length in males, 27.0–27.2% in females.
Natural History Notes: Inhabits seasonally flooded rice fields and grassland with forest
patches. Its diet consists of frogs and fish (Lee et al. 2018).

Figure 25.50. Xenochrophis bellulus.

Photography by Gunther Kohler.

412
Aquatic Snakes

Painted Keelback
Xenochrophis cerasogaster (Cantor, 1839)
Distribution and Habitat. Pakistan, Nepal, Bangladesh, India.
Occurs in or near water bodies, such as ponds and rivers. Type
locality: Vicinity of Calcutta, India.
Identification. Body slender and cylindrical, with a long tail.
Head not distinct from neck. Eye with round pupil. Loreal pres-
ent, one preocular, and three (rarely 4) postoculars. Nine (rarely
Figure.25.51. The distribu-
tion of Xenochrophis cera- 8) upper labials, only the 4th touching eye; 10 (rarely 9 or 11) low-
sogaster. er labials. Scales in 19 rows on midbody, strongly keeled except for
the outer 2-3 rows. Ventrals 140-158, cloacal divided, subcaudals
68-80 (males).
Dorsum reddish brown, with a thick tan or light brown dorsolateral stripe on each side.
The stripe is more prominent posteriorly. In some specimens, this stripe may be rather in-
distinct. Yellow ventrolateral stripes, and yellowish on the labials. Ventrals and subcaudals
marbled red, with dark edges and pale speckling.
Natural History. Mostly aquatic, found in still water with thick emergent vegetation. Has
been observed resting on lily pads, or grasses at marsh edges. It is expected from late June to
November but absent during winter. Diurnal and active, but it is also timid and will flee to
the water in the presence of a potential threat. They feed on fish, amphibians, and shrimp.
Oviparous, known to lay eggs in early April and hatch during the rainy season (May-July).

Spotted Keelback
Xenochrophis maculatus (Edeling, 1864)
Distribution and Habitat. Belitung, Natuna Archipelago,
Riau Archipelago, Sumatra, Borneo, Brunei Darussalam, Ma-
laysia, and possibly Singapore. Occurs in lowland forest with
Figure 25.53. The distribution of
small streams and swamps. Type locality: Martapoura, pres
Xenochrophis maculatus. de Bandjermasin, Borneo (now Banjarmasin).
Identification. Body slender and cylindrical, tail length
moderate. Head distinct from neck. Eye exceptionally large
with round pupil. Loreal present, one preocular, and three postoculars. Nine upper labials,
only 4th to 6th touching eye. Scales in 19 rows on midbody, strongly keeled. Ventrals 140-156,
cloacal divided, subcaudals 95-117.
Dorsum olive brown, with four longitudinal rows of small black spots, and two rows of
yellow spots. Labials yellow or white, with black sutures. Ventrals and subcaudals yellowish
with black edges.
Natural History. Mostly aquatic, but more terrestrial than other species of this group.
Diurnal, feeds mainly on frogs. Reproductive habits unknown but presumed to be oviparous.

413
 24. Mountain Keelbacks

Figure.25.52. Xenochrophis cerasogaster.

Photography by Avrajjal Ghosh.

414
Aquatic Snakes

Figure 25.53. Xenochrophis maculatus from Marudi, Sarawak, Malaysia (Island of Borneo).
Photography by Bernard DuPont.

Triangle Keelback
Xenochrophis trianguligerus (Boie, 1827)
Distribution and Habitat. Wetlands (ponds, swamps, rice
paddies) close to evergreen forests in Southeast Asia, includ-
ing the island of Borneo (Brunei Darussalam, Sabah, Sarawak,
Kalimantan); Burma (Myanmar), Cambodia, India (Nicobar
Islands; Arunachal Pradesh); Indonesia (islands of Nias, Men-
Figure 25.54. The distribution tawai, Sumatra, Riau Archipelago, Bangka, Billiton, Java, Bali,
of Xenochrophis trianguligerus . Sulawesi, Sangihe Archipelago, Ternate); Laos, peninsular
Malaysia; Singapore; Thailand, and Vietnam. Type locality:
Java.
Identification. The head is olive on the crown, the upper labials are yellow. The body is
415
 24. Mountain Keelbacks

Figure 25.55. Xenochrophis trianguligerus.

Photography by Mediyansyah.

light brown to olive green with yellow and red highlights. Yellow and red coloration towards
the neck fading into white towards the tails with black saddle markings running the length
of the body. Laterally the body has red triangular markings with the apex near the ventral
surface. Each triangle has a pale center. Dorsal scales are keeled and in 19 rows at mid-body.
Ventrals are 132-153. Total length up to 1350 mm.
Natural History. A nocturnal, semi-aquatic snake. The diet includes fish, tadpoles, frogs,
and other small vertebrates. Females are oviparous and lay clutches of 5-15 eggs.

416
Aquatic Snakes

Striped Keelback
Ceratophallus vittatus (Linnaeus, 1758)
Distribution and Habitat. Native to Indonesia and West Malaysia. Introduced to Singa-
pore and Puerto Rico. Occurs in lowland areas up to 1000m asl, associated with water bodies
such as swamps and rice paddies. Type locality: “America” (in
error).
Identification. Body robust and cylindrical, with a long
tail. Head distinct from neck. Eye large with round pupil,
diameter of eye greater than distance between eye and edge
of lip. Loreal present, one preocular, and three postoculars.
Nine upper labials (rarely 8), 4th to 6th (rarely 4th-5th)
touching eye. Scales in 19 rows on midbody, strongly keeled.
Figure 25.56. The distribution
of Ceratophallus (Xenochrophis) Ventrals 138-155, cloacal divided, subcaudals 66-88. Base
vittatus. color of dorsum light brown, with a black vertebral stripe.
Natural History. Mostly aquatic, associated with shal-
low, still water such as rice paddy fields. It has also been found near streams and rivers, bask-
ing on stones during the day. It is a very fast-moving snake and will secrete a musk when
captured. Autonomy of the tail has also been recorded. Diurnal, actively hunting frogs and
fish. Oviparous. The invasive population in Puerto Rico is eating the critically endangered
frog Eleutherodactylus juanariveroi and the White-lipped Frog, Leptodactylus albilabris.
Note. As this book was going to press Patel et al. (2023) found Hebius beddomei (Gun-
ther, 1864) to be the sister to a clade containing Fowlea and Atretium. Freshly collected spec-
imens and existing museum material allowed us to elucidate the generic status of the species
and identify two distinct populations, one of which is described as a new species. A new ge-
nus, Sahyadriophis was erected to accommodate Sahyadriophis beddomei, and a new, second
species Sahyadriophis uttaraghati . The discovery of the new Oligocene divergent lineage,
Sahyadriophis adds the the evidence of the Western Ghats being a source of relic lineages.

Figure 25.57. Ceratophallus (Xenochrophis) vittatus.

Photography by Mediyansyah.

417
 24. Mountain Keelbacks

On the next page is Opistjhotropis cheni. From Shimentai NR, Yingde City, Guangdong, China Pho-
tography by Jian Wang.

418
Aquatic Snakes

419
 26. Asian Stream Snakes & Relatives

26. Asian Stream Snakes & Relatives

This clade contains three genera of poorly known southeast Asian snakes that may range
from almost wholly aquatic to mostly terrestrial. The genus Smithophis contains four poorly
understood fossorial-cryptozoic-semiaquatic snakes. The genus is named in honor of Mal-
colm A. Smith in recognition of his contributions to the knowledge of Asian snakes and is
sometimes called two-colored forest snakes. The degree to which these snakes are aquatic
remains to be determined. At least a few of these snakes feed on earthworms and skinks.
However, its two sister genera (Giri et al. 2019) are the highly aquatic Opisthotropis and the
semi-aquatic Trimerodytes (formerly Sinonatrix). Opisthotropis is distributed on mainland
Southeast Asia eastwards to the Philippines and the Ryukyu Archipelago of Japan. Most Opist-
hotropis are known only from a few specimens or described based on single specimens, and
most inhabit hill streams and nearby terrestrial habitats. They are known to forage at night on
juvenile crabs, earthworms, fishes, tadpoles, frogs, and shrimps. Opisthotropis is oviparous.
Trimerodytes likely originated in Hengduan Mountains (eastern Tibetan Plateau) in western
China at 23.93 Ma (17.09–31.30) (Gou et al. 2020). While most clade members appear to be
oviparous, at least one species of Trimerodytes (T. annularis) is viviparous.

Arunachal Rain Snake

Smithophis arunachalensis Das et al., 2020
Distribution and Habitat. Known from streams at six localities from eastern Arunachal
Pradesh, including Roing and Koro-
nu in Lower Dibang Valley District,
Dening in Lohit District, and three
localities in Namdapha Tiger Reserve
in Changlang District of Arunachal
Pradesh. The elevation ranges from
310 m in the foothills of Mishmi Hills
to 580 m at Namdapha Forest Camp
(Das et al. 2020).
Natural History. The holotype

Figure 26.1. The distribution of Smitho-

phis arunachalensis.

was caught while it was moving on the

side of a stream in a moist evergreen
forest. The type locality was in a de-
graded forest with extensive bamboo Figure 26.2. Smithophis arunachalensis
regeneration along both sides of the Photography by Abhijit Das.
stream. The stream habitat where the

420
Aquatic Snakes

snake was collected had large, bryophyte-covered boulders and bedrock near a pool and riffle
section. The snake was caught in dim light during heavy rain near dawn at 05:00 hours.

Mizo Rain Snake

Smithophis atemporalis Giri et al., 2019
Distribution and Habitat. Known from secondary forests
in Mizoram, India. Type locality: Mizoram University Cam-
pus, Aizawl, India, about 833 m elevation. It has an elevation-
al range of 71–1,235 m asl (Remruatpuii et al. 2020).
Natural History. It is most frequently encountered during
the monsoon season. It has been observed feeding on juve-
nile skinks and earthworms in nature and on the Bangladeshi
Figure 26.3. The distribution of Cricket Frog (Minervarya asmati) eggs in captivity (Giri et
Smithophis atemporalis. al. 2019). A female (SVL 437 mm) was observed feeding on
an earthworm (Eutyphoeus sp.; to-
tal length 263 mm) in the middle
of a road surrounded by a second-
ary forest used as a plantation. The
earthworm had crawled onto the
road and was quickly followed by
the snake. The snake bit the earth-
worm and struggled with it for about
5 min while slowly working its way
to the posterior end of its prey be-
fore swallowing it. The elapsed time
from strike to complete ingestion was
about 14 minutes (Muansanga and
Figure 26.4. Smithophis atemporalis
Lalremsanga 2020). It is oviparous,
Photography by H.T. Lalremsanga.
laying a maximum of four eggs per
clutch (Remruatpuii et al. 2020).

Brown Trapezoid Snake,

Smithophis bicolor (Blyth, 1854)
Distribution and Habitat. Assam, Meghalaya; Arunachal
Pradesh (Deban, Namdapha - Changlang district), Maharash-
tra, India; northern Myanmar; and western Yunnan, China.
Type locality: Assam, India.
Natural History. Feeds on worms and slugs (Smith 1943).

Figure 26.5. The distribution of

Smithophis bicolor.

421
 26. Asian Stream Snakes & Relatives

Jingpo Mountain Stream Snake,

Smithophis linearis Vogel, et al. 2020

Figure 26.7. The distribution of Smithophis

linearis.

Distribution and Habitat. It has been

found along streams in tropical evergreen
forests in Yunnan, China. Type locality:
Nabang Town, Yingjiang County, Dehong
Dai, and Jingpo Autonomous Prefecture,
Yunnan Province, People’s Republic of
China. Elevation ca. 467 m ASL. Known
from three localities with a 40 km radius
Figure 26.6. Smithophis bicolor (Vogel et al. 2020).
Photography by Abhijit Das. Natural History. Diet is said to com-
pose mollusks and arachnids.

Gary’s Mountain Stream Snake

Opisthotropis alcalai Brown and Leviton, 1961
Distribution and Habitat. Mindanao, Philippines.
Type locality: Cugat Creek, on the west side of Dapitan
Peak, Mt. Malindang, Zamboanga del Norte Province,
Mindanao at 2500 m ASL.
On 17 March 2019, Maglangit et al. (2023) oppor-
tunistically encountered a single individual of this spe-
cies and observed it at 19:30 hrs in heavy rainfall. The
snake was feeding on tadpoles (possibly Leptobrachium
Figure 26.8. The distribution of
lumadorum, Pelobatrachus stejnegeri, Staurois natator
Opisthotropis alcalai.
tadpoles), and macro-invertebrates in a temporary pool
at 440 meters elevation). The pool was near the fast-moderately flowing water channels of
Mambuntan Falls. The habitat was a regenerating secondary-growth forest that transitioned
to an agricultural area, approximately 10 meters distance from the locality. The vegetation
was characterized by epiphytes, ferns, mosses, shrubs, and dominant flowering plants. The
pool microhabitat was shallow (0.7 m) and covered with dead leaves (water pH = 6.7). The
location was about 95 km straight-line distance from its type locality.

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Aquatic Snakes

Figure 26.9. Opisthotropis alcalai.

Photography by Sharmaine Bonachita.

Anderson’s Mountain Stream Snake

Opisthotropis andersonii (Boulenger, 1888)
Distribution and Habitat. Hill streams and wetlands of Hong
Kong and adjacent Shenzhen City it extends north to Guang-
zhou City, China, and into northern and central Vietnam. It is a
low-elevation mountain dweller, commonly found in montane
fast or slow-flowing streams, tiniest trickles covered with bare
rocks, pools, and ditches, surrounded by moist subtropical ev-
Figure 26.10. The distribution ergreen broadleaved forests (Wang et al. 2017).
of Opisthotropis andersonii. Natural History. Inhabits small, fast water mountain streams

Figure 26.11. Opisthotropis

andersonii, Hong Kong.
Photography by Mike Pin-
gleton.

in secondary forest; probably absent from larger streams. During the day it is concealed un-
der rocks or in stream bed gravel, active at night to forage. Speculation on its diet suggests
it probably feeds on semi-aquatic earthworms. Reported to do well in captivity on a diet of

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 26. Asian Stream Snakes & Relatives

earthworms. Hatchlings are about 120 mm, most adult specimens 300-400 mm, it may reach a
maximum size of 500 mm.

Günther’s Mystery Stream Snake

Opisthotropis atra Günther, 1872
Distribution and Habitat. Unknown. Type locality: West Africa. This species has never
been rediscovered. The genus is not present in Africa. The holotype is in the Natural History
Museum in London – otherwise, this species is unknown.

Zhao’s Stream Snake

Opisthotropis cheni Zhao, 1999
Distribution and Habitat. Hill streams of Guangdong,
China. Type locality: Mt. Mang, China. Mangshan Forestry
Bureau, Yizhang County, Hunan, China, 480 m elevation fide
Zhao (2004).

Figure 26.12. The distribution of

Opisthotropis cheni.

Cuc’s Mountain Stream Snake

Opisthotropis cucae David et al., 2011
Distribution and Habitat. Hill streams of Kon Tum Pla-
teau, Kon Tum Province, Vietnam. Type locality: Chu Mom
Ray National Park, Sa Thay District, Kon Tum Province (14°
26.227’ N, 107° 43.268’ E), 740 m elevation, Vietnam.
Figure 26.13. The distribution of
Opisthotropis cucae.

Tien’s Mountain Stream Snake

Opisthotropis daovantieni Orlov, et al., 1998
Distribution and Habitat. Probably restricted to the hill
streams of the northern part of Tai Nguyen Plateau in the
Annam Mountains, S Vietnam. Type locality: Buoenloy, (14°
20’N, 108° 36’E), 750 m elevation, Ankhe District, Gia Lai
Prov., S Vietnam.

Figure 26.14. The distribution of

Opisthotropis daovantieni.

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Aquatic Snakes

Figure 26.15. Opisthotropis cheni.

Photography by: A Ho Koon Centre Hong Kong; B-D Jian Wang E. Konrad Mebert.

Durand’s Mountain Stream Snake

Opisthotropis durandi Teynié, et al., 2014
Distribution and Habitat. Louangphabang, Laos, may also
be present in adjacent Thailand. Type locality: vicinity of Muang
Ngoi (20°42’10”N, 102°41’21”E), Ngoi District, Louangphabang
Figure 26.16. The distribution Province, Laos, elevation about 370 m.
of Opisthotropis durandi.

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 26. Asian Stream Snakes & Relatives

Figure 26.17. Opisthotropis durandi.

Photography by Peter Barkles.
Guangxi Mountain Stream Snake
Opisthotropis guangxiensis Zhao, Jiang and Huang, 1978
Distribution and Habitat.
Known from the type locality of
Mount Yao (Dayao Shan), Jinxiu,
Guangxi Zhuang Automonous Re-

Figure 26.18. The distribution of

Opisthotropis guangxiensis.

gion, China (Zhao and Adler 1993).

It has also been recorded from
Guangdong (Kadoorie Farm and
Botanic Garden, 2004). Zhao (2006)
also mentioned Hunan which is not Figure 26.19. Opisthotropis guangxiensis from Mt. Yunkai,
supported with specimens. It is occursXinyi City, Guangdong, China. Photography by Jian Wang.
at elevations of 950 to 1,220 m ASL.
This species has been recorded
living in and near streams in both forests as well as open habitats. It spectulated to feed on
semi-aquatic annelids (Zhao 2006). The IUCN considers it near threatened.

426
Aquatic Snakes

Hung-Ta Chang’s Mountain Stream Snake

Opisthotropis hungtai Wang, et al., 2020
Distribution of Habitat. Hill streams of Guangdong, Chi-
na. Type locality: Heishiding Nature Reserve, Fengkai County,
Zhaoqing City, Guangdong Province, 300 m ASL. P.R. China.
Natural History. Currently known from Heishiding Na-
ture Reserve (ca 300 m ASL.) and Dawuling Forestry Station
Figure 26.20. The distribu-
tion of Opisthotropis hungtai. (ca 900 m ASL) in western Guangdong, and Mt. Wuhuang (ca
500 m ASL.) in southeastern Guangxi. The specimen from Mt.

Figure 26.21. Opisthotropis

hungtai Mt. Yunkai, Xinyi
City, Guangdong, China.
Photography by Jian Wang.

Wuhuang was collected in a

rocky stream. Besides, speci-
mens from Heishiding Nature
Reserve were found in pelitic
gutterways along the dirt path,
and specimens from Dawuling
Forestry Station were collected
in a pelitic stream. The collec-
tion sites were all surrounded
by well-preserved, dense decid-
uous forest (Wang et al. 2020).

Hai Ha Mountain Stream

Snake
Opisthotropis haihaensis
Ziegler et al., 2019
Distribution and Habits.
Known from the type locality,
the forest near Tai Chi Village

(ca 950 m ASL), Quang Ninh, northern Vietnam, and Shi-

wandashan Nature Reserve (ca 500 m ASL), southwestern
Guangxi, southern China. The distance between the two lo-
calities is approximately 150 kilometers, indicating that the
distribution area of this species is the mountain region on the
Figure 26.22. The distribution of border between China and Vietnam.
Opisthotropis haihaensis.

427
 26. Asian Stream Snakes & Relatives

Figure 26.23. Opisthotropis haihaensis.

Photography by Jian-Huan Yang.

Chapa Mountain Stream Snake

Opisthotropis jacobi Angel & Bourret, 1933

Distribution and Habitat. Japan (Ryukyu Islands:

Kumejima Island). Type locality: Kumejima Island, Okina-
wa Prefecture, Japan. This species is a highly endangered
aquatic or semiaquatic species endemic to Kumejima Island
Figure 26.24. The distribution of the Okinawa Group, Ryukyu Archipelago.
of Opisthotropis jacobi. Natural History. Field studies suggest the snake is ac-
tive almost year-round; the snake tends to be diurnal in the
warmer and nocturnal in the cooler seasons. And predation
on small freshwater crabs, Candidiopotamon kumejimense.

Kikuzato’s Brook Snake

Opisthotropis kikuzatoi (Okada & Takara, 1958)
Distribution and Habitat. Hill streams of north Vietnam
and Yunnan, China. Type locality: Chapa, Tonkin-Yunnan
border (now Sa Pa, Lao Cai Province, northern Vietnam). Re-
cently collected specimens were found at 19:00–22:00 h on
rocks along small rocky streams. The surrounding habitat
was a secondary evergreen forest of hardwood trees. The air
Figure 26.25. The distribution of temperature was 19.5–25.1 C and the relative humidity was
Opisthotropis kikuzatoi. 75–90% (Ziegler et al. 2018).

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Aquatic Snakes

Chinese Mountain Stream Snake

Opisthotropis kuatunensis Pope, 1928
Distribution and Habitat. The hill streams of Fujian, Zhe-
jiang, Jiangxi, Guangxi, and Hong Kong, China. Type locality:
Ch’ungan Hsien, Fukien Province, China.
Natural History. Diet. A female O. kuatunensis was observed
(Sung and Wag 2011) attempting to swallow an egg of Paa ex-
Figure 26.26. The distribution ilispinosa (Lesser Spiny Frog) in an artificial pond located in
of Opisthotropis kuatunensis. a secondary forest. The pond
was connected to a 2 m wide
stream. When the egg was in-
gested halfway, the snake de-
tected our presence, released
the egg, and fled. The snake
was caught for measurement
and a photographic record.
Nineteen eggs of P. exilispino-
sa at early tail-bud stage were
found in the pond. The clutch
of P. exilispinosa contains ap-
proximately 50–150 eggs, and
the entire clutch is typically
deposited together in a small
pool. The few eggs observed at
this site suggest that the snake,
or possibly some other preda-
tor, had consumed other eggs.
Eggs of P. exilispinosa are ap-
proximately 12–23 mm in di-
ameter, with embryos ranging
from 9 to 11 mm, depending
upon developmental stage.
These are the second largest
embryos of any stream-breed-
Figure 26.27. Opisthotropis kuatunensis. Top Hong Kong. Artur
ing frog in Hong Kong, aside
Tomaszek; Bottom. [Link], Shenzhen, Guangdong, China from those of the Giant Spiny
Jian Wang. Frog (P. spinosa), and their size
and oviposition as a group po-
tentially represents a substan-
tial food source for O. kuatenensis.
In a second observation at 2040 h on 11 January 2010, at Tai Mo Shan (22.54278°N,
114.28917°E, datum: WGS84; elev. 878 m), New Territories, Hong Kong, a male O. kuatun-
ensis (SVL = 32 cm; total length = 38.5 cm) was observed biting the head of a Leptolalax liui
(Leaf Litter Toad) tadpole (total length ca. 3 cm) in a pool along a natural stream in secondary
forest. The tadpole struggled and escaped after being held for 30 seconds, and we were unable
to capture it. This sighting confirms that O. kuatunensis eats tadpoles in the wild. These two
species are commonly sympatric in streams in Hong Kong, and it is likely that L. liui tadpoles

429
 26. Asian Stream Snakes & Relatives

are regular prey for O. kuatunensis. Although the natural diets of stream snakes in southern
China are largely unknown, amphibians may be important prey. Lui et al. (2021) sequenced
the mitochondrial genome of this snake.

Bicolored Stream Snake

Opisthotropis lateralis Boulenger, 1903
Distribution and Habitat. Streams and stream side
habitats in Hoa Binh, North Vietnam and Guangxi,
Guizhou, and Hong Kong, China. Type locality: Manson
Mountains, Tonkin.
Natural History. A specimen was found in a stream
next to the ranger station on Ba Mun Island (16 June) and
Figure 26.28. The distribution of
contained five yellowish eggs with maximum lengths from
Opisthotropis lateralis.
19.1 to 21.2 mm.

Figure 26.29. Opisthotropis lateralis.

Photography by Kevin Messenger.

Sichuan Mountain Stream Snake

Opisthotropis latouchii (Boulenger, 1899)
Distribution and Habitat. Inhabits the smallest forested streams in southern China -
Guangdong, Fujian, Jiangxi, Hunan, Guangxi, Guizhou, Sich-
uan. Type locality: Kuatun, Fukien Province, China.
Natural History. Diet Pope (1935) describes this species a
burrower, that feeds on earthworms. Eggs reported laid under
cascading water, partially submerged. This species lives in cold
water streams and spends much of its time under submerged
Figure 26.30. The distribution rocks.
of Opisthotropis latouchii.

430
Aquatic Snakes

Figure 26.31. Opisthotropis latouchii.

Phototgraphy by Kevin Messenger.
Lau’s Mountain Stream Snake
Opisthotropis laui Yang, Sung & Chan, 2013
Distribution and Habitat. Known from the type locality at Beifengshan Forest Park in
Mt. Gudou, Jiangmen City, Guangdong Province, China. Mt. Gudou is an isolated mountain
range situated along the coastline at the western side of the Pearl River Estuary and has been
protected as a nature reserve since 2001. The summit, known
as Shizitou, is 982 m ASL. Original vegetation of the mountain
range has been largely destroyed. Currently it supports sec-
ondary forest, as well as shrubland and grassland; the stream
valleys generally support more mature natural forest. The Ho-
lotype was found swimming in a small stream at night about
2145h, which is shallow, rocky, and about m wide, with young
Figure 26.32. The distribu- but closed-canopy natural forest. The stream drains pass Tais-
tion of Opisthotropis laui. han City, Guangdong Province, before emptying into the Pearl
River Delta.

Yellow-spotted Mountain Stream Snake

Opisthotropis maculosa Stuart & Chuaynkern, 2007
Distribution and Habitat. Thailand (Nong Khai Province). Type locality: Thailand, Nong
Khai Province, Boong Klar District, Phu Wua Wildlife Sanctuary, at marker 027 on footpath
to Lad Phleuy Waterfall, 190 m ASL.
Natural History. Known from the Phu Wua Wildlife Sanctuary, Nong Khai Province,

431
 26. Asian Stream Snakes & Relatives

Figure 26.33. Opisthotropis laui from Mt. Gudou, Taishan City, Guangdong, Chi-
na. Photography by Jian-Huan Yang.

432
Aquatic Snakes

northeastern Thailand. The Holotype was found swim-

ming at night during a heavy rain. The stream passed
through a disturbed bamboo mixed with evergreen forest
(Stuart and Chuaynkern 2007).

Figure 26.34. The distribution of

Opisthotropis maculosa.

Figure 26.35. Opisthotropis macu-

[Link] its habitat. Photography
by Bryan Stuart.

Maxwell’s Mountain Stream

Snake
Opisthotropis maxwelli Boulenger,
1914
Distribution and Habitat. South-
east of Guangdong Province, the south-
west of Fujian Province and south of
Jiangxi Province. It inhabits montane
slow-flowing streams with sand and
gravel
bottom at
elevations
between
425–1000
m ASL
Figure 26.36. The distribu-
(Wang et
tion of Opisthotropis max-
al. 2017).
welli.
Natural History. This species occurs in high-altitude slow mov-
ing streams with sand and gravel substrates and often found un-
der stones (Zhao et al. 1998). This is an oviparous species. It inhabits montane slow-flowing
streams with sand and gravel

Sumatran Stream Snake

Opisthotropis rugosa (Lidth De Jeude, 1890)
Distribution and Habitat. Sumatra, Indonesia. Type locality: Kajutanam, western Suma-
tra, Indonesia. Inhabits streams in primary lowland forests. The
type locality was previously forested and at an altitude of about
300 m ASL. The second locality is Toru River in Aek Nangali,
Batang, within a forested area at about 600 m ASL.

Figure 26.37. The distribu-

tion of Opisthotropis rugosa.

433
 26. Asian Stream Snakes & Relatives

Figure 26.38. Opisthot-

ropis maxwelli from Mt.
Muotianshi, Puning City,
Guangdong, China. Pho-
tography by Jian Wang.

Shenzhen Stream Snake

Opisthotropis shenzhenensis Wang et al., 2017
Distribution and Habitat. From the low mountain ranges
in the eastern Shenzhen and southern Dongguan, Guangdong
Province, China, including Mt. Wutong, Sanzhoutian and
Mt. Tiantou, and Mt. Yinping. All specimens found at night
in slow-flowing streams covered with bare rocks at elevations
Figure 26.39. The distribution of
between 155 and 327 m ASL Type locality: Shenzhen City,
Opisthotropis shenzhenensis. Guangdong Province, China.

Smith’s Mountain Stream Snake

Opisthotropis spenceri Smith, 1918
Distribution and Habitat. Thailand. Type locality: Ngao, Lampang Province, northern
Thailand It is also known from Nan Province. It has been collected from streams traversing
dry evergreen forest (Chuaynern et al. 2014.)
Natural History. Diet composed of earthworms. Females know to lay clutches of three
eggs. In the 100 years following its description this snake was known from only four speci-
mens and little was known about its ecology. In 2015, a location was found where the snakes
were abundant, and their feeding behavior could be observed. The snakes were seen to hunt
freshly molted crabs (soft shells). They would grab and constrict the crab until its shell burst,
then reach inside to swallow the soft inner parts. Then it made a loop in its body that it pulled
the rest of the crab through until it broke up the hardened shells, and swallowed those parts as

434
Aquatic Snakes

well. This was the first time that dismemberment as a feeding

strategy had ever been observed among a freshwater snake
species.

Figure 26.40. The distribution of

Opisthotropis spenceri.

Figure 26.41. Opisthotropis spenceri.

Photography by Montri Sumontha.

Tam Do Stream Snake

Opisthotropis tamdaoensis Ziegler, David & Vu, 2008
Distribution and Habitat. Tam Dao, Vinh Phuc Provinces, north Viet Nam. Type locali-
ty: Bac (Silver) stream, Tam Dao, Vinh Phuc Province, northern
Vietnam, ca. 750 m elevation.

Figure 26.42. The distribution

of Opisthotropis tamdaoensis.

Figure 26.43. Opisthotropis tamdaoen-

sis. Photography by Tomas Ziegler.

435
 26. Asian Stream Snakes & Relatives

Borneo Stream Snake

Opisthotropis typicus (Mocquard, 1890)
Distribution and Habitat. Found in and near streams in pri-
mary rainforest. It has been found from sea level to 900m ASL
in the Malaysian states of Sabah and Sarawak, and the sultanate
of Brunei on the island of Borneo. Type locality: Mt. Kinablau,
Sabah, Northern Borneo.
Figure 26.44. The distribu-
tion of Opisthotropis typicus.

Figure. 26.45. Opisthotropis typicus. Photography by Amaël Borzée.

Quy’s Mountain Stream Snake

Opisthotropis voquyi Ziegler et al., 2018
Distribution and Habitat. Small rocky streams in sec-
ondary evergreen forest at the Type locality: Type locality:
Tay Yen Tu Nature Reserve, Bac Giang Province, 437 m
ASL
Natural History. Nocturnal, found in and nest to
streams in secondary evergreen forest of medium and
Figure 26.46. The distribution of small hardwoods mixed with bamboo, shrubs, and vines.
Opisthotropis voquyi. The air temperature was 29.2–36.5oC and the relative hu-
midity was 55–73%. Co-occurring species: Cyclophiops
multicinctus, Opisthotropis lateralis, Sinonatrix percarina-
ta, Gekko palmatus, Sphenomorphus cryptotis, and Shinis-
aurus crocodilurus vietnamensis.

436
Aquatic Snakes

Figure 26.47. Opisthotropis voquyi

Photography by Thomas Ziegler.

Zhao’s Mountain Stream Snake

Opisthotropis zhaoermii Ren et al., 2017
Distribution and Habitat. Known only from the type lo-
cality in western China. Type locality: Zuolong Gorges, Gu-
zhang, Tujia-Miao of western Hunan, China
Natural History. Known only from the type locality
where it inhabits small, fast-flowing mountain streams in
forested areas, with water temperature and the pH between
Figure 26.48. The distribution of 19.9–21.2 °C and 7.85–7.93, respectively. Individuals were
Opisthotropis zhaoermii. seen swimming at the edge of the backwater of travertine wa-
terfalls from 2100h to 0100h at night. The holotype was col-
lected during a heavy rainstorm that caused the water to be-
come extreme turbid. When handled, individuals struggled violently and released a musky,
pungent, and enduring defensive odor. The scales of the snakes become dehydrated and
crimped quickly after leaving the water for about 10 min, but recovered rapidly when
returned to water. Similar to other members of the genus in China, this snake may prey on
earthworms, tadpoles, freshwater isopods, crabs, and small fish (Pope, 1935; Ren et al. 2017).

Figure 26.49. Opisthotropis zhaoermii

Photography by Cen Peng.

437
 26. Asian Stream Snakes & Relatives

Annulated Keelback
Trimerodytes aequifasciatus (Barbour, 1908)
Distribution and Habitat. Streams in evergreen forests to at
least 700 m in China, North Vietnam, and Laos. Type locality: Mt.
Wuzhi, Hainan Island, People’s Republic of China.
Inhabits streams and waterways in or near forests, including
ditches bordering agricultural land (Bain and Truong 2004, Orlov
Figure 26.50. The distribution of
Trimerodytes aequifasciatus.
et al. 2000, Stuart 1999). It is oviparous and is active both at day
and at night.
Natural History. The diet includes
small fish. Pope (1935) considered
this species a inhabitant of mountain
streams, always finding them in very
close proximity to streams and noted
they readily drop into the water when
disturbed.

Red-bellied Annulate Keelback

Trimerodytes annularis
(Hallowell, 1856)
Distribution and Habitat. Taiwan,
Fujian, Jiangxi, Chekiang, Anhwei, Ki-
angsu provinces China. Type locality:
Ningpo, China.
Natural History. Nocturnal, uses
rice paddies, swamps, and ponds. Diet
includes tadpoles, frogs, and fish. Fe-
males give birth to 1-14 neonates.
In 1914, Trimerodytes annularis
was reported as a freshwater aquacul-
ture pest in Taiwan (Oshima, 1914). It
was widely distributed throughout the
low-elevation rice paddies, ditches, and
freshwater wetlands of Taiwan before
the 1970s’ economic and industrial
highly development era (Maki, 1931;
Horikawa, 1941; Wang and Wang,
1956; Kuntz, 1963). From an ecological
aspect, T. annularis is a piscivorous rare
occasion on frog and high habitat fi-
delity species with a small home range,
Figure 26.51. Trimerodytes aequifasciatus. Photography
reproduction associated with the hydro-
by (A-B) Artur Tomaszek, (C) Tom Chun Wa Lee.
logical dynamic of wetlands, and pop-
ulation size can reach 1,000 snakes in a
ca.4-hectare suitable wetland-agriculture mosaic ecosystem, and good population conditions
represented on equal sex ratio or skewed female and low body injury or scar ratio (Mao, 2003).
438
Aquatic Snakes

Neonates take three years at minimum to reach maturity

size, ca. SVL 45 cm in females, lifespan 10-15 years. In
2009, a four-year general survey and habitat assessment
focused on the vulnerable Taiwanese watersnake pop-
ulation was conducted, and only two districts retained
T. annularis populations in north Taiwan (Lai and Mao,
Figure 26.52. The distribution of Tri- 2012). So far, present known populations and habitats of
merodytes annularis. this species are all on private land. One has
suffered from a semiconductor science
park expansion, habitat pollution, and
loss. Another struggle with the multi-
ple factors’ twisting of extinction vortex:
habitat destruction by landscape hazard
prevention engineering, the threat of
invasive species, small population par-
adigm, the unequal sex ratio of male
bias, predation, nematode-infected and
transmission, high body injury rate from
agriculture machine, damaged from pes-
ticide and herbicide, water source short-
age causes freshwater ecosystem collapse,
extreme climate events, etc. The present
Figure 26.53. Trimerodytes annularis.
status of T. annularis in Taiwan is critical-
Photography by Oscar Yin.
ly endangered.

Hainan Mountain Keelback

Trimerodytes balteatus (Cope, 1895)
D ist r ibu -
tion and Hab-
itat. Known
from torrents
and mountain
streams at low
Figure 26.54. The distribution to moderate
of Trimerodytes balteatus elevations from
southern Chi-
na including Hainan, Guangxi, Guangdong,
and Hong Kong), and Vietnam (Hai Phong
(Cat Ba Island), and Bac Giang (Tay Yen Tu
Nature Reserve).
Natural History. A diurnal species of-
ten hides under rocks in the water. Forages
in streams or drainage ditches near village;

Figure 26.55. Trimerodytes balteatus Photography

by Carol Kwok.

439
 26. Asian Stream Snakes & Relatives

adults were encountered competing for a single loach, where each snake bit on one end of the
fish. A strong musk is released when captured. Tail autotomy is also used to escape predators
(Pope 1935). This snake was long considered a member of the genus Opisthotropis.

Olive Annulate Keelback

Trimerodytes percarinatus (Boulenger, 1899)
Distribution and Habitat. Streams in secondary forests
in Vietnam, India, China, Taiwan, Myanmar, Laos, and Thai-
land. Type locality: Kuatun (now Guadun), Chongan, Fujian
Province, China.
Natural History. Nocturnal and arboreal on vegetation
Figure 26.56. The distribution of Tri- hanging above small puddles or pools in streams, under rocks
merodytes percarinatus
in streams, in secondary forest, in wet hill evergreen forests.
Preyed upon by the krait Bungarus fasciatus.

Figure 26.57. Trimerodytes percarinatus.

Photography by Artur Tomaszek.

440
Aquatic Snakes

Jingdong Water Snake

Trimerodytes yapingi Guo, Zhu, and Liu, 2019
Distribution and Habitat. Yunnan, China. Type locality:
Jingdong Yunnan Province, China, at an elevation of 1500
m ASL.

Figure 26.58. The distribution

of Trimerodytes yapingi.

Yunnan Annulate Keelback

Trimerodytes yunnanensis Rao and Yang, 1998
Distribution and Habitat. The distribution of Trimero-
dytes yunnanensis is disjunct; it has been reported from
southern China, Thailand (Chiang Rai Province), Vietnam
(Son La Province), and northern Myanmar (Kachin state) and
inhabits streams and ponds in montane forests ranging from
400 to 2000 m ASL.
Figure 26.59. The distribution of Natural History. Captives ate fish and tadpoles and some-
Trimerodytes yunnanensis. times held the fish above the water before swallowing them.

Figure 26.60. Trimerodytes yunnanensis

Photography by Thai National Parks.

Angel’s Mountain Keelback

Trimerodytes (=Paratapinophis) praemaxillaris Angel, 1929
Distribution and Habitat. Rocky streams in northern Laos, Thailand, and China (Yun-
nan). Type locality: Chieng-Kuang, upper Laos. Natural History. Nocturnal, in fast-moving,
rocky streams. On land, this species appears sluggish, whereas it appears much more agile in

441
 26. Asian Stream Snakes & Relatives

the water, suggesting that this species is strictly aquatic. It is

also found in a flowing rocky stream (c. 2 m broad), running
through the forest and fringed by bamboo scrub. Both Nan
River females contained digested remains of fish (scales and
a mandible) and numerous roundworms (Nematoda), while
the three males’ stomachs contained only roundworms. The
Figure 26.61. The distribution of reproductive mode of this snake is unknown, however, T.
Trimerodytes (formerly Parata-
praemaxillaris is known to have an egg tooth, suggesting it
pinophis) praemaxillaris.
is oviparous.
Ren et al. (2019) suggest this snake may belong to the genus Trimerodytes. However, they
did not assign it to the genus because Murphy et al. (2008) described it as having two pairs of

Figure 26.62. Trimerodytes (Paratapinophis) praemaxillaris.

Photography by Michael Cota

posterior chin shields in an unusual configuration (the second pair of chin shields are separat-
ed by another pair of plate-like scales). Therefore, molecular data can resolve the relationships
of these snakes.

On the next page. A Smooth-scaled Blackbelly Snake Hydraethiops melanogas-

ter from Sembe, Congo Photography by Vaclav Gvozdik.

442
Aquatic Snakes

443
Aquatic Snakes

27. African Water Snakes

Five genera of natricids occur in sub-Sahara Africa (Afronatrix, Helophis, Hydraethiops,
Limnophis, Natriciteres) and form a clade with the sister being the endemic Seychelle Islands
Lycognathophis seychellensis (Deepak et al. 2021). The Lycognathophis seychellensis lineage
likely reached the Seychelles via over water dispersal from Africa, approximately 43–25 Ma.
Deepak et al. (2021) suggested that the mainland sub-Saharan African natricids, are relatively
poor in species, and are ecologically conservative despite 30 million years of evolution, with
all species being predominantly semiaquatic or aquatic (Gibbons and Dorcas, 2004; Vitt and
Caldwell, 2009). However, Lycognathophis seychellensis appears to be the only sub-Saharan
natricid radiation that is predominantly terrestrial. It is encountered on the ground but has
also been observed in vegetation at heights of at least five meters as well as on the margins of
pools and streams. All mainland sub-Saharan African natricids are oviparous. Lycognathophis
seychellensis is probably also oviparous but that is not yet supported by direct observation.
Luiselli and Akani, (2002) showed that females of the water snakes Natriciteres fuliginoi-
des, N. variegata, Afronatrix anoscopus tended to show body temperatures inversely related
to their size (snout–vent length), and that gravid specimens tended to maintain higher body
temperatures than non-gravid specimens. However, the high and constant body temperature
exhibited at night strongly indicate that these species can maintain high and stable body tem-
peratures with little no thermoregulatory behavior.
As a clade these snakes are understudied and in need of a systematic revision. As might be
expected most are poorly known in regard to their natural history.

Brown Water Snake

Afronatrix anoscopus (Cope, 1861)
Distribution and Habitat. Water bodies in rainforests of West
Africa from Liberia, Sierra Leone, southern Senegal, Ivory Coast,
SW Burkina Faso, Guinea (Conakry), Mali, Nigeria, Cameroon,
Ghana, Togo, and Benin. Type locality: Originally reported as
Cuba (in error).
Natural History. Field research on these semi-aquatic snakes
Figure 27.1. The distribution of in southern Nigeria, have provided detailed data on their diets
Afronatrix anoscopus. (Akani and Luiselli, 2001; Luiselli et al., 1998, 2003), reproduc-
tive biology (Luiselli and Akani, 2002a), thermal ecology (Luisel-
li and Akani, 2002a), and population biology (Akani and Luiselli, 2001).
Afronatrix anoscopus is among the most abundant snakes in the aquatic habitats of rain-
forest zones in West Africa (Chippaux, 1999), including southern Nigeria (Butler and Reid,
1990). Nonetheless, until recently, little data were available on their ecology. (e.g., see Butler
and Reid, 1986, 1990). The diet is composed of tadpoles and fish, but adult amphibians are
also eaten. The tail is prehensile and anchors the snake to submerged debris while it hunts. A
total of 332 specimens (59 subadults, 131 males and 142 females) were collected in southern
Nigeria to study their diet. The adult male to female sex-ratio (0.92) was not significantly
different from equivalence, with female body length being greater than that of males. A total
of 60.2 % of the subadults, 39% of the males, and 43.2% of the females had identifiable food
items in their stomachs. The diet of all categories (subadults, males and females) consisted of
amphibians and fish, with a predominance of anuran tadpoles and the frog Silurana tropicalis.

444
 27. African Water Snakes

Subadults took significantly more

tadpoles than adult frogs.

Marsh Snakes, Natriciteres

The Marsh Snakes are found
in Sub-Saharan Africa. They are
diurnal, oviparous, and they show
tail autonomy, but seem unable
to re-grow their tail once it is
damaged. Their diet is composed
mostly of fish and anurans.

Figure 27.2. Afronatrix anoscopus from Bamenda, Cameroon.

Photography by Vaclav Gvozdik.
Southwestern Forest Marsh Snake
Natriciteres bipostocularis Broadley, 1962
Distribution and Habitat. Zambia, Democratic Repub-
lic of the Congo (Zaire: Katanga), central Angola; elevation
1320-1585 m. This is a montane species. Branch (2018)
pointed out that there is a large disjunction between the
main range of N. bipostocularis in Zambia and DRC and the
isolated records in western Angola, although the eastern re-
gions of the country remain poorly surveyed.
Figure 27.3. The distribution of
Natriciteres bipostocularis

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Aquatic Snakes

Collared Marsh Snake

Natriciteres fuliginoides (Günther, 1858)
Distribution and Habitat. A semi-aquatic snake that
will wander away from water. Guinea, Ghana, Togo, Ni-
geria, Cameroon, Central African Republic, Democratic
Republic of the Congo (Zaire; except in the east), Congo
(Brazzaville), Gabon, Sierra Leone, Liberia. Type locality:
West Africa.
Figure 27.4. The distribution of Natural History. Common in swamp forest and riv-
Natriciteres fuliginoides
ers. Haagner (1994) reported it from mangrove forest.
They are active in leaf litter during the day or found under
logs or debris in marshy areas and riverbeds. One specimen contained the remains of the
ranid frog Dimorphognathus africanus, while in captivity another ate another the small ranid
Phrynobatrachus auratus.

Figure 27.5. Natriciteres fuliginoides from Motima River, DRC.

Photography Vaclav Gvozdik.

446
 27. African Water Snakes

Olive Marsh Snake

Natriciteres olivacea (Peters, 1854)
Distribution and Habitat. Flooded forests and savannas from Sudan south to Mozam-
bique westward to Guinea and Angola. Jackson et al. (2007)
found this specxies closely assocaited with flooded forest.
Natural History. A diurnal, savanna and forest species that
does not live far from water. It swims well and feeds on frogs and
tadpoles, small fish, and some invertebrates while in the water.
Figure 27.6. The distribution of
Natriciteres olivacea.

Figure 27.7. Natriciteres olivacea (A) an adult, (B) hatchling (C) a melanistic adult (D) from Kanzi,
DRC. Photography by (A) Twan Leenders (B) Paul Freed (C-D) Vaclav Gvozdik.

Pemba Island Marsh Snake

Natriciteres pembana (Loveridge, 1935)
Distribution and Habitat. Marshes and swamps (including
coastal marshes that are brackish water) of Pemba Island, Tan-
zania; colonizes agriculture landscapes- wet rice paddies. Type
locality: Chake Chake, Pemba Island. Endemic to Pemba Island.

Figure 27.8. The distribution

of Natriciteres pembana.

447
Aquatic Snakes

Figure 27.9. Natriciteres pembana.

Photography by Frank Glaw.
Forest Marsh Snake
Natriciteres sylvatica Broadley, 1966
Distribution and Habitat. Coastal forest edges from
southeast Tanzania south to Kwa Zulu and inland to eastern
Zimbabwe border and central Mozambique. coastal forests
between elevations of 600-2000 m.
Natural History. The diet includes small fish, frogs, and
fish-eating spiders. Females lay five to six eggs in the sum-
Figure 27.10. The distribution of mer.
Natriciteres sylvatica

Figure 27.10. Natriciteres sylvatica.

Photography by Johan Marais
448
 27. African Water Snakes

Variable Marsh Snake

Natriciteres variegata (W. Peters, 1861)
Distribution and Habitat. Southern Democratic Republic
of the Congo (Zaire), Gabon, Central African Republic, Equa-
torial Guinea, Cameroon, Nigeria, Benin, Togo, Ghana, Ivory
Coast, Liberia, Sierra Leone, Guinea (Conakry), Burkina Faso.
Type locality: Ghana.
Figure 27.11. The distribution of
Natriciteres variegata.

Figure 27.12. Natriciteres var-

iegata. Photography by Vaclav
Gvozdik.

Schouteden’s Mud Snake

Helophis schoutedeni (De Witte, 1942)
Distribution and Habitat. Swamp forests in west and cen-
tral Democratic Republic of the Congo (Zaire), southeast
Brazzaville, Congo. Type locality: “Kwamouth (district du
Moyen-Congo) [et ...] Tondu, sur Ie Lac Tumba (district de
l’Equateur)” (DRC)
This is a poorly known snake Nagy et al. (2014) noted the
Figure 27.13. The distribution of
Helophis schoutedeni.
majority of distribution records come from the Eastern Congo-
lian swamp forests upriver the Congo River and its tributaries,
it raises the question about the origin of the specimens collected
in Léopoldville/Kinshasa. The strong current of the Congo River often carries floating vege-
tation islands, which occasionally introduce fauna and flora from the upriver into the Lower
Congo region. This phenomenon might also explain the findings of the Schouteden’s Sun
Snake outside the swamp forests.
449
Aquatic Snakes

Nagy et al. considered, Helophis schoutedeni a semi-aquatic snake because of its dor-
sally-oriented nostrils, narrow triangular internasals, small eyes with round pupil and stout
body.
Observations of captive specimens supported this with a specimen that preferred to stay
in water. Helophis shows an extreme similarity to the genus Hydraethiops, as de Witte (1922)
indicated in the original description.

Figure 27.14. Helophis schoutedeni.

Photography by Vaclav Gvozdik.

Smooth-scaled Blackbelly Snake

Hydraethiops laevis Boulenger, 1904
Distribution and Habitat. Small streams in primary
rainforest in Cameroon and Gabon. Type locality: Efulen,
southern Cameroon. This snake is known from three spec-
imens. The holotype was 510 mm in total length. Deepak et
al. (2021) found this species to be the sister of Heliophis.
Figure 27.15. The distribution of
Hydraethiops laevis.

Rough-scaled Blackbelly Snake

Hydraethiops melanogaster Günther, 1872
Distribution and Habitat. Rainforest streams and ponds. Gabon, Democratic Republic
of the Congo, Brazzaville, Congo, Central African Republic, and Cameroon. Type locality:
Gabon, Africa.

450
 27. African Water Snakes

Natural History. Diet includes fish and tadpoles. A female

with 13 eggs in December (Schmidt 1923). Gabon,Africa.
Natural History. A semi-aquatic species found in streams
within tropical moist forest, and in gallery forest in savannas.
It feeds on amphibians and fish including Mormyridae (ele-
phantfish) (Chippaux 2006, Mamonekene et al. 2006). Diet
includes fish and tadpoles. A female with 13 eggs was found
in December (Schmidt 1923).
Figure 27.16. The distribution of
Hydraethiops melanogaster.
African Swamp Snakes, The Limnophis Clade
The genus ranges from Zam-
bia, west to the Democratic
Republic of the Congo (Katan-
ga), E Angola, Botswana, Na-
mibia. Inhabit the freshwater
ecoregions: Cuvette Centrale,
Kasai, Lower Congo Rapids,
Ogooue - Nyanga - Kouilou –
Niari, Southern Gulf of Guin-
ea Drainages – Bioko, North-
ern Gulf of Guinea Drainages.
The head is slightly depressed,
snout rounded, eyes moderate
in size, pupil round; Smooth
scales in 19 rows; subcaudals
divided. Oviparous

Figure 27.17. Hydraethiops melanogaster. Photographs from

Twan Leenders (top), Vaclav Gvozdik (bottom two).

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Aquatic Snakes

Bangweulu Water Snake

Limnophis bangweolicus (Mertens, 1936)
Distribution and Habitat. Moist savannas and floodplains
from Zambia, west to the Democratic Republic of the Congo
(Zaire: Katanga), east Angola, Botswana, Namibia. Type local-
Figure 27.18. The distribution ity: Nsombo, Northern Rhodesia.
of Limnophis bangweolicus.

Figure 27.19. Limnophis bangweolicus.

Photography by Werner Conradie.

452
 27. African Water Snakes

Bicolored Swamp Snake

Limnophis bicolor Günther, 1865
Distribution and Habitat. southwest Angola, southern Zam-
bia (Okavango and Zambezi drainage basins), eastern andsouth-
ern Democratic Republic of the Congo (Zaire), elevation 1000 -
1800 m.
Figure 27.20. The distribu-
tion of Limnophis bicolor.

Figure 27.21. Limnophis bi-

color. Photography by Alex
Rebelo

Angolan Water Snake

Limnophis branchi Conradie, Deepak, Keates and Gower, 2020
Distribution and Habitat. northeast Angola (Lagoa Carum-
bo region). Type locality: Luele River area north of Lagoa
Carumbo, Lunda Norte Province, Angola.

Figure 27.22. The distribu-

tion of Limnophis branchi.

Figure 27.23. Limnophis branchi.

Photography by Bill Branch.

453
Aquatic Snakes

454
 28. The Palearctic Watersnakes

28. The Palearctic Watersnakes

Konrad Mebert, Daniel Jablonski
Natrix is a Palearctic genus from northern Africa and temperate Eurasia east to West-
ern China and Pakistan with five semi-aquatic snakes (natricines) currently. They attain a
total length of usually around 1 meter, with the larger Grass Snakes rarely growing up to
1.5 meters, and exceptionally longer with the record at 2.05 meters of a specimen from Krk
Island, Croatia (Schreiber 1912, cit. in Kabisch 1999). The head is well differentiated from
the body, and the tail is rather short. They have large eyes with round pupils. Depending on
the species, there are 17 to 23 dorsal scales rows, mostly keeled, less along the paraventral
rows. Teeth are aglyphic, and sharp. A Duvernoy’s gland is present. The diet consists pri-
marily of amphibians and fish. They are oviparous, with females reaching a larger body size
than males. Natricines usually swim on the surface but will submerge when disturbed or
hunting for fish and amphibians. They occasionally climb into bushes for thermoregulation,
and are usually diurnal but switch to nocturnal activity on warm nights. Predator defense is
manifold, individually, and locally variable, and can include in all species tight coiling, head
hiding under its body, excretion of musk through bilateral anal glands, triangular flattening
of the head and also the body, regurgitating undigested food, death feigning, tail vibration or
wiggling, hissing. The coloration of juvenile and adult individuals is only slightly different.
Fossil records and current distribution indicate that the genus Natrix originated in west-
ern Asia (Hecht 1930; Rage and Augé 1993; Ivanov 2001; Guicking et al. 2006). The genus
is monophyletic, a complex with five species, including the Viperine Snake Natrix maura
Linnaeus 1758, the Dice Snake N. tessellata Laurenti, 1768 and the Grass Snake group, a
complex of three species with the traditional Eastern Grass Snake N. natrix Linnaeus 1758,
the Ibero-Maghrebian Grass Snake N. astreptophora, and the Western Grass Snake N. helvet-
ica (Guicking et al. 2006a; Speybroeck et al. 2020). The ancestor of Natrix separated in the
Miocene and evolved independently into N. maura in Northern Africa, and N. tessellata in
Western Asia. The Grass Snakes (N. natrix complex) radiated north into Europe and central
Asia, also reaching a few islands by marine transgression (Guicking and Joger 2011).
Natrix tessellata and the Grass Snake group are sister to N. maura that occupies a bas-
al position, diverging from the common ancestor about 18–27 MYA. Grass Snakes and N.
tessellata split about 13–22 MYA. However, newer studies showed more concrete divergence
times from a common Natrix ancestor to be ca. 21.96 Mya , 18.39 Mya, and 9.58 Mya for
N. maura, N, helvetica/natrix, and N. tessellata clades, respectively (Supplementary files in
Jablonski et al. 2023).Intraspecific divergence for N. maura begun at approximately 5.3 MYA,
for the Grass Snakes (N. natrix complex) 6.0 MYA, and for N. tessellata 6.7 MYA (Guick-
ing et al. 2006a). In the western Mediterranean Basin, the Messinian Salinity Crisis likely
led to the split of Natrix species (maura and astreptophora) between the Iberian Peninsula
and Northern Africa, whereas latter Natrix populations subsequently split into Moroccan
and Algerian-Tunisian clades (Guicking et al. 2008; Kindler et al. 2018). Even though these
semi-aquatic snakes are good swimmers, the 14 km width of the Strait of Gibraltar (passage
from the Atlantic into the Mediterranean Sea) was a significant biogeographic barrier for a
population exchange (Guicking et al. 2002).

On the previous page. is a Natrix maura. Photography by Kai Kolodziej.

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Aquatic Snakes

Viperine Snake
Natrix maura (Linnaeus, 1758)

Xavier Santos, Juan M. Pleguezuelos and Konrad Mebert

Distribution and Habitat. Type locality: “Algiriae” = Algeria. The Viperine Snake occurs
across the African and European countries that form the Western Mediterranean. In Africa, it
is present in Morocco, Algeria, Tunisia, and coastal Libya as far east as the springs and irriga-
tion complex around Tawergha (Tauorga), District Misratah (Kramer and Schnurrenberger
1963; Bauer et al. 2017). In Europe, it is present in Spain, Portugal, France, southwestern
Switzerland, and northwestern Italy, and has sporadically been observed in Andorra, a small
country in the Pyrenean Mountains. On Gibraltar, a British Overseas Territory located at the
southern tip of the Iberian Peninsula, the species was found on the isthmus (Cortés 1982),
but no new records were reported in the last 40
years. Detailed information on distribution is
found in atlases of many of these countries such as
Portugal (Santos 2008), Spain (Santos et al. 2002),
and France (Santos and Vacher 2010). Overall,
the Viperine Snake is predominantly found along
bodies of water within open, scrubland and forest-
ed habitats. The southern distribution is limited
by the aridity of the Sahara Desert (Naulleau and
Figure 28.1. The distribution of Natrix maura. Schätti 1997). However, populations persist in
that region along wadis (periodically water-filled
valleys) such as Oued Ziz and Oued Drâa (Martínez del Marmol et al. 2019) and other small
water bodies (Bons and Geniez 1996; Schleich et al. 1996). The southernmost population
is in Assa, Morocco (ca. 28º N) near the Drâa River (de Lepiney and Nemeth 1945-46-47).
Cold climate shapes the northern limit and approximately follows the southern sections of
the French rivers Seine-et-Marne and Aube into the region of Franche-Compténear, and the
Rhone River into Lake Geneva and Canton Valais in Switzerland (Schätti 1999; Santos and
Vacher 2010). Italy’s distribution in Mediterranean habitats is limited to the northeast by the
presence of the closely related Dice Snake Natrix tessellata. Both species overlap across ca. 200
km, from the eastern Cuneo Province to the western Parma Province. Although both species
partly could compete in sympatry by using similar ecological niches (Schätti 1982), various
studies found some niche partitioning, such as smaller, shadier streams preferred by N. mau-
ra and larger, deeper water bodies in a more open landscape (Mazza et al. 2011; Metzger et al.
2011; Scali 2011); also see the Natrix tessellata account herein.
The Viperine Snake inhabits several islands in the western Mediterranean Basin, e.g., Ile
du Levant (Toulon-Hyères), Île de Yeu and Île d’Oléron, in the Atlantic close to the mainland
in France (Schätti 1999), Mallorca (Spain), Menorca (Spain), Sardinia and some adjacent
islets (Italy). Genetic and morphological studies have demonstrated that the distribution in
these islands is recent and caused by (historical) human introductions. This is the case of the
Balearic Archipelago, where the Romans could have introduced the Viperine Snake (Alcover
and Mayol 1981) from the European mainland (Guicking et al. 2006a). In addition, there are
a few reports of its presence in marine habitats, e.g., in Italy (Lanza 1983), in southern Spain
(Cabo and Olea 1978), and northeastern Spain (Fuentes and Escoriza 2015). Still, it is best

456
 28. The Palearctic Watersnakes

Figure 28.2: Natural color pattern variation of the Viperine Snake (Natrix maura) from: A) High Atlas Mountains,
central Morocco; B) northern Morocco; C) southwestern France; D) southern Spain; E) southwestern France; F)
western Switzerland; G) Sardinia Island, Italy. Photo credit: A), E), F) Konrad Mebert; B), D) Octavio Jiménez-Ro-
bles; C), H) Matthieu Berroneau; G) Laura and Bobby Bok.

457
Aquatic Snakes

known for the thriving populations on the Cies Archipelago, Pontevedra, off the Atlantic
coastal city Vigo, northwestern Spain, where they feed exclusively on marine fishes along
rocky islets and grow to large body sizes (Galán 2004, 2012).
The habitat of Natrix maura is widely influenced by the Mediterranean climate on the
Iberian Peninsula, France, and Morocco (Godinho et al. 1999; Santos et al. 1997, 2002; Santos
2008), where it occupies all natural and artificial water bodies. The Viperine Snake is rare in
the driest regions, such as Almeria, southeastern Spain. The species is frequent at elevations
between 500–600 m (Pleguezuelos and Villafranca 1997), less common up to 1200 m, with
the highest elevations records at 2050 m in the Sierra Nevada Mountains, southern Spain
(Pleguezuelos and Feriche 2003), and up to 2600 m elevation at Oukäimeden, Alto Atlas,
Morocco (Bons and Geniez 1996). It inhabits all natural water bodies, such as rivers, lakes,
streams, wetlands, ponds and lagoons, and river valleys within mountain chains with a cool
climate like the Pyrenees. The Viperine Snake also occupies artificial water bodies such as
irrigation ponds and channels, fountains, ditches, and pools, showing high flexibility to colo-
nize new water habitats if suitable prey is available. It also occupies brackish water in coastal
marshes and rocky seashores where prey is abundant (Cabot and Olea 1978; Del Canto and
Busack 2011; Galan 2012). The Viperine Snakes are easily observed swimming and foraging
in the water and can be observed basking on land near the water or under rocks. They rarely
wander off far from water. Thus, in arid African regions, the Viperine Snake is strictly linked
to sites of permanent water bodies, such as oueds and oases (Bons and Geniez 1996; Schleich
et al. 1996).
Identification. This medium-sized snake usually does not reach more than 1 m total
length, but up to 1.25 m on Cies Archipelago in the Atlantic Sea. Females are the larger sex
with males exhibiting a relatively long tail, but on average, about 20% for both sexes. Males
have 2-3 more ventral scales and 9-10 more subcaudal scales than females. The head is wider
than the neck and body. When threatened by a potential predator, it can be flattened into a
viper-like triangular shape, a typical anti-predator posture in natricine snakes. The pupil is
round and bordered by a light-colored ring. Usually, there are two preocular and two posto-
cular scales, 6-9 supralabial scales the third and fourth usually contact the eye. Dorsal scales
are arranged in 21 rows at mid-body and are heavily keeled except for the rows in contact
with the ventral scales.
The most conspicuous characteristic is the dorsal pattern, composed of bilateral alter-
nating dark, round- or rhomboidal blotches along the vertebral line from neck to tail. These
dorsal blotches often form a zigzag or slalom-shaped band at some part/s on the body. Lateral
spots alternate with the dorsal ones and may form ocelli, a ring-like blotch with a whitish
center. Two blotches join on the pileus. Infra- and supralabial scales are light with darker
colors in the contact zone between scales. The dorsal background coloration varies from
olive, brown and grey tones to reddish. The ventral coloration is grey or yellowish, with black
spots in many ventral scales. Albinotic and melanistic individuals have been found in the wild
(Schätti 1999).
The visually most intriguing morphological variation is a stripped dorsal pattern. This
pattern is composed of two dorsolateral yellow bands formed between the dorsal and lateral
blotches (bilineata morph: Schätti 1982). The striped pattern is sometimes overlooked on the
zigzag pattern in the same individual. The zigzag pattern is more common (76%, n = 917;
Santos et al. 2018) than the bilineata pattern. Often the only morph in many populations
is the zig-zag pattern. The bilineata pattern occurs across most of the species’ distribution
(Schätti 1982). Still, it is linked to open habitats with close populations showing marked dif-
ferences in the proportion of bilineata individuals (Santos et al. 2018). There is a latitudi-

458
 28. The Palearctic Watersnakes

nal gradient in total body size, with African Viperine Snakes larger than Iberian individuals
(Fahd 2001; Saint Girons 1956). However, optimal conditions can result in populations with
huge specimens like those on the Cíes Archipelago, northwestern Spain, or the population
along Lake Geneva’s rocky and fish-rich shore, near Lausanne, Switzerland (Koller and Ur-
senbacher 1996; K. Mebert unpubl. data). The number of ventral and subcaudal scales is
higher in southern populations, with a clinal variation especially marked among African in-
dividuals (Schätti 1982, 1999; Fahd 2001).
Phylogeny and phylogeography. Analyses based on cytochrome b sequences by Guic-
king et al. (2002) showed that the Viperine Snake is structured in three lineages in Tuni-
sia-Sardinia, Morocco, and Iberia-France, with genetic distances between them ranging from
3.9-4.6%. The divergence time was around 3-3.5 Ma, during the Mid or Late Pliocene. The
African clades of N. maura were older than the European ones, suggesting an African origin
of the species. Indeed, the three distinct lineages between Tunisian and Moroccan popula-
tions suggest multiple independent migrations between European and African populations
as ancient as the Messinian Crisis (Barata et al. 2007; Bauer et al. 2017). A phylogenetic study
using mitochondrial markers from the three Natrix groups (maura, tessellata, and natrix s.l.)
indicated that N. maura was basal in the phylogeny and coalescence time among the three
species was dated 13-22 Ma ago during the early Miocene or late Oligocene (Guicking et al.
2006b). The genetic structure among European populations is low, with an average genetic
distance of 1.3% (Guicking et al. 2008). Haplotype diversity is higher on the Iberian Peninsula
than on northernmost populations. The peninsula could act as Pleistocene refugia during
glaciations, with rapid recolonization towards its northern range in subsequent interglacial
periods. On the Iberian Peninsula, Guicking et al. (2008) also detected higher genetic diversi-
ty among river valleys in southern Iberia, compared to northern Iberian populations.
Fossil records of Natrix specimens in Iberia were first documented by Sanz and Sanchiz
(1980) and Sanchiz (1991). More recently, Blain et al. (2008a) described Viperine Snake fossils
at Cueva Victoria (Murcia) dated in the Upper Pleistocene (ca. 1.1 Ma). Blain et al. (2008b,
2010) found fossils at Sima del Elefante and Gran Dolina, Atapuerca (Burgos, Spain) dated
1.2 Ma ago. Other N. maura fossils have been recorded in Sète, southern France, and Cueva
Gorham, Gibraltar, by Bailón (1991) and Blain et al. (2013), respectively.
Finally, there is morphological evidence for occasional hybridization between N. astrep-
tophora and N. maura (Gonzalez de la Vega et al. 2021). Furthermore, molecular analysis
showed that past hybridization with subsequent genetic introgression happened between N.
maura and N. tessellata (Schöneberg et al. 2023), two species for which no hybrids have been
documented to date, neither from the wild nor from captivity (Mebert et al. 2011a).
Population Aspects. The Viperine Snake is one of the most abundant species within its
range in Morocco, Spain and southern France (Bons and Geniez 1996; Santos and Vacher
2010; Santos 2014) and can attain dense populations. For example, at the Jalón River, Alican-
te, eastern Iberian Peninsula, Hailey and Davies (1987a) estimated 4800 adults/ha along a 4
m wide section of the riverbank; The high density was attributed to the lack of predators and
the high availability of prey. Similar trends have been noted for the coastal wetlands such as
the Ebro Delta and many Mediterranean rivers. However, water contamination, physical hab-
itat degradation, and fish decline have caused the reduction of Viperine Snake populations
(Santos and Llorente 2009).
Hailey and Davies (1987d) and Filippakopoulou et al. (2014) have calculated growth
trajectory by ectopterygoid annual rings and observed that Viperine Snakes reach an asymp-
totic plateau as growth ceases after the acquisition of sexual maturity, when snakes start to
invest energy into reproduction (Hailey and Davies 1987a). Immature females grow faster

459
Aquatic Snakes

than males (3.4 and 2.9 cm/year, respectively, Hailey and Davies 1987d), which explains why
they attain larger body size (Filippakopoulou et al. 2014). Also, based on ectopterygoid an-
nual growth lines, the maximum lifespan was estimated to be 22 years (Filippakopoulou et al.
2014), with females living longer than males (Hailey and Davies 1987a).
Activity and Thermal Biology. Emergence from hibernation begins in March (Jaén and
Pérez-Mellado 1989; Santos and Llorente 2001a) or April in colder regions (Galán 1988).
However, it might be observed on the surface on sunny winter days in southern Iberia (Santos
and Llorente 2001a; Malkmus 1996, 2008). The seasonal activity of the Viperine Snake is ac-
centuated in spring for mating and in autumn for hatching (Galán 1988; Santos and Llorente
2001a, 2001b). During the hot summer, snakes spend copious time in the water (Hailey et
al. 1982; Duguy and Saint Girons 1966; Jaén and Pérez-Mellado 1989) or they become more
nocturnal (Hailey and Davies 1986b; Pleguezuelos and Feriche 2003; Scali 2011). Experimen-
tal studies estimated that the selected temperature of Viperine Snakes ranged from 24.7–29°
C (Spellerberg 1977; Hailey and Davies 1986a). At 10° C body temperature, snakes cannot
digest prey, whereas digestion was optimal at 35° C (Hailey and Davies 1987c). The body
temperature of active snakes covers a wide range between 14–35° C (Jaén and Pérez-Mellado
1988; Santos 2000; Hailey et al. 1982; Hailey and Davies 1987c). Reproductive females attain
higher body temperatures than other adults or immature snakes (Santos 2000), and body
temperature is higher in summer compared to spring and autumn (Hailey et al. 1982; Hailey
and Davies 1986b; Santos 2000). The thermal pattern differs for snakes in and out of the wa-
ter. Viperine Snakes were active in water at temperatures ranging from 13–34° C (Hailey et al.
1982; Jaén and Pérez-Mellado 1989; Santos 2000). They behaved as thermoconformers with
body temperature equaling water temperature. In the Ebro Delta, northeastern Spain, snakes
thermoregulated heliothermic, i.e., terrestrially on the surface, in spring, and switched to
thigmothermic basking (substrate heating, usually under rocks) during the summer (Santos
2000).
Activity patterns and home range have been estimated using radiotelemetry in the Ebro
Delta and by capture-recapture techniques in the Jalón River, Alicante, eastern Spain. Males
had larger home ranges in spring (5.8 ha, n=4) compared to summer (0.7 ha, n=7) due to
searching for mates (Santos and Llorente 1997). Similarly, snakes covered longer distances in
spring compared to summer due to reproductive activity (Santos and Llorente 1997; Duguy
and Saint Girons 1966) and the distribution of aquatic prey (Hailey and Davies 1987d). Latter
authors detected longer lineal home ranges along the river in females than in males, presum-
ably due to the larger body size of the former sex.
Reproduction. Age and size of sexual maturity varies between the sexes. Females start
reproduction at 4–5-year age at 310–350 mm of snout-vent length (SVL), whereas males ma-
ture at 2–3-year age and 220–250 mm SVL (references in Santos 2014). Mating starts when
snakes emerge from hibernation in the spring (Duguy and Saint Girons 1966; Hailey and
Davies 1987b; Santos and Llorente 2001b). A second mating period may occur in autumn
(Duguy and Saint Girons 1966). There is one clutch per season in the wild, whereas more
than one clutch per year has been observed in captivity (Petch 1991; Schleich et al. 1996). A
mating ball has been documented once with eight snakes of which probably only one was a
female due to its larger size (Torres-Porras 2008). Females will mate several times (Dumont
1979; Hailey and Davies 1987b) and maintain viable spermatozoa for several years (Quillon
and Lamouille 1978).
Spermatogenesis is postnuptial, and males start to produce new spermatozoa after mat-
ing which means that testes function were at their maximum in the summer and spermatozoa
are stored in the vas deferens until the following spring (Santos and Llorente 2001b). The

460
 28. The Palearctic Watersnakes

Viperine Snake is an oviparous species with 70 to 100% of mature females reproducing annu-
ally (Feriche and Pleguezuelos 1999; Rugiero et al. 2000; Hailey and Davies 1987b; Duguy and
Saint Girons 1966; Santos and Llorente 2001b). Vitellogenesis is prenuptial, as follicles start to
grow after hibernation (Duguy and Saint Girons 1966; Feriche and Pleguezuelos 1999; Hailey
and Davies 1987b). Females with oviductal eggs were found in June and July. Egg size ranges
from 28–37 x 14–19 mm and incubation lasts 40–45 days (Duguy and Saint Girons 1966;
Dumont 1979; Feriche and Pleguezuelos 1999; Santos and Llorente 2001b). Thus, neonates
appear in mid-August with a total length between 146 and 220 mm (Duguy and Saint Girons
1966). Clutch size ranges from 2–27, with an average of 7.3 and increases with female size
(see references in Schätti 1999; Santos 2015). Vitellogenesis is energetically costly for females,
which allocate fat bodies stored in the abdomen to grow follicles (Santos and Llorente 2004).
However, females can also mobilize other energetic sources during vitellogenesis and gesta-
tion, such as feeding and maybe resorption of unfertilized eggs or muscle (Santos et al. 2007).
Foraging and Diet. The Viperine Snake feeds on varied fish and amphibians captured
in the water. Prey types, foraging ecology and modes strongly resemble its eastern counter-
part, the Dice Snake Natrix tessellata, (see chapter herein). Fish species recorded in the diet
of N. maura vary in body size, swimming speed, position in the water column and schooling
aggregation, whereas amphibians varied according to size and water dependency from very
terrestrial to almost strictly aquatic species. Fish eaten by the Viperine Snake include the fol-
lowing: Family Anguilidae (Anguilla anguilla), Blenniidae (Salaria fluviatilis), Centrarchidae
(Lepomis gibbosus), Ciprinidae (Barbus bocagei, Barbus graellsii, Barbus haasi, Barbus sclateri,
Carassius auratus, Chondrostoma miegii, Chondrostoma polylepis, Cyprinus carpio, Phoxinus
phoxinus, Rutilus arcasii, Rutilus rutilus, Squalius cephalus), Cobitidae (Barbatula barbatula),
Gasterosteidae (Gasterosteus aculeatus), Gobiidae (Gobius gobius), Poecilidae (Gambusia af-
finis, Gambusia holbrooki) and Salmonidae (Oncorhynchus mykiss, Salmo trutta) (references
in Santos 2014), although this list is incomplete. Amphibians preyed upon include Caudata:
Salamandra salamandra, Pleurodeles waltl, Lissotriton boscai, Lissotriton helveticus, Triturus
marmoratus, Chioglossa lusitanica; anurans: Alytes muletensis, Alytes obstetricans, Alytes dick-
hilleni, Bufo spinosus, Epidalea calamita, Discoglossus galganoi, Discoglossus jeanneae, Pelo-
bates cultripes, Rana iberica, Phelophylax perezi, Rana temporaria and Hyla molleri, whereas
other vertebrates, e.g. small mammals or reptiles, are also preyed on occasionally (references
in Schätti 1999; Santos 2014). Ayres (2012) reported snakes scavenging on dead newts (Lis-
sotriton boscai) in Galicia, northwestern Spain. Additionally, immature snakes also prey on
tadpoles and egg clutches of amphibians, as well as on invertebrate prey such as earthworms,
leeches, snails, slugs and insects (references in Schätti 1999; Santos 2014). Predation on rep-
tiles such as the Morish gecko Tarentola mauritanica and small mammals has also been re-
ported (Pleguezuelos and Moreno 1989; Vericad and Escarré 1976).
Two main foraging modes were described by Hailey and Davies (1986c): 1) ambush, used
by adult snakes which consists of the visual identification of a prey, followed by ambushing
at any point in the water column through anchoring the posterior body to the substrate or
submerged plant. Upon the prey’s approach to within striking distance, the snake launches
forward with its anterior body; 2) active foraging, which consists of exploring the water and
its substrates using tongue flicking in search for chemical cues, a mode particularly common
in shallow waters.
Other studies have described alternative foraging behaviors: Naulleau (1964) described
the use of the tongue to lure fish, Martorell (1990) documented snakes ambushing fish from
outside the water by anchoring themselves on the bank of a river and partially suspended over
the water, waiting for the passage of prey, and Alarcos et al. (2012) documented a snake climb-

461
Aquatic Snakes

ing on cattail leaves to capture Hyla frogs. After the seizure of large fish, snakes tend to move
their catch to land for swallowing (Meijide and Salas 1989; Pérez de Lanuza 2011), whereas
Pleguezuelos and Feriche (2003) observed nocturnal Viperine Snakes maintaining aquatic
prey brought on land until the prey dies, before consuming them. Most foraging modes are
virtually identical as in the Dice Snake (Natrix tessellata), see chapter herein.
The proportion of fish versus amphibians depends on prey availability and local climate.
For example, in northern Iberian wetlands, amphibian represents about 50% of prey whereas
in southern and Mediterranean-type areas amphibian represents less than 25% of prey (di-
etary references in Santos 2014). Hence, the Viperine Snake is an opportunistic predator that
forages on the most commonly available prey (Hailey and Davies 1986c; Pleguezuelos and
Moreno 1989; Rugiero et al. 2000; Santos et al. 2000). However, snakes may also focus on fish
species that are easy to catch, such as blenid fish hidden under rocks in the benthic zone of
rivers (Santos et al. 2006) or submerged riparian rock formations of Atlantic islands (Galán
2004).
The Viperine Snake shows an ontogenetic shift in their diet. Immature snakes tend to
forage on tadpoles, earthworms, and aquatic arthropods (references in Santos 2014), whereas
adults can consume prey that represents 80% of the snake’s body size (Santos et al. 2006).
Hailey and Davies (1986c) found dead snakes that tried to swallow fish that were 60% of the
snake’s body size. Despite space limitation in the body cavity, gravid females continue foraging
(Feriche and Pleguezuelos 1999; Santos and Llorente 1998; Santos et al. 2006).
Predators and Defense. Viperine Snakes have many predators, including birds, such as
herons, storks, and raptors, and mammals, such as otters, wild boars, beech martens and other
meso-carnivores (references in Schätti 1999; Santos 2014). This high predation pressure varies
among localities. For example, coastal wetlands and some Mediterranean rivers with abun-
dant herons and other potential predators, the percentage of Viperine Snakes with broken
tails accounted for 18.7% on average. Given that the tail loss indicates predation failure, this
high percentage means a high predation pressure on these animals (Santos et al. 2011). Be-
cause this is an aquatic snake, parasites found in the digestive tract are abundant and include
six trematode, one cestode and six nematode species (Navarro and Lluch 1993; Navarro et al.
1987; García 1987). Parasites have also been found in the trachea (Lluch and Navarro 1986).
Viperine Snakes exhibit several morphological and behavioral defense traits that mim-
ic the sympatric European viper species, a nice example of Batesian mimicry (Aubret and
Mangin 2014, Santos et al. 2018). For example, N. maura exhibits a viper-like zigzag dorsal
pattern and displays head triangulation, a defense coiling body posture, hissing, body infla-
tion, and forward launching of its head/anterior body when threatened. These snakes also
excrete feces, a noxious liquid from its cloacal glands, and regurgitate freshly captured prey.
Experimental studies with plasticine models have demonstrated both the aposematic
and cryptic value of the viper-like zigzag dorsal pattern. The lack of such a Batesian mimicry
function in bilineata Viperine Snakes resulted in a higher attack rate on bilineata plasticine
models than zigzag ones (Santos et al. 2018). In addition, field observations showed that both
morphs differed in their defensive behaviors: zigzag snakes tended to remain immobile or flee
at shorter distances compared to bilineata snakes. As bilineata snakes are more common in
open habitats, e.g., marshes, Santos et al. (2018) concluded that snakes with the lined pattern
benefit from reduced detection or catchability by a visual predator in grass and reed-domi-
nated habitats.
Conservation. The Viperine Snake is listed in Appendix III of the Bern Convention. It
is protected by national legislation in Switzerland and Spain. The global IUCN category is
Least Concern (Mateo et al. 2009) and the same in regional IUCN Red Lists (Santos et al.

462
 28. The Palearctic Watersnakes

2002; Vacher and Geniez 2010). Despite this apparent protection, aquatic contamination,
freshwater habitat degradation, and prey decline in streams, rivers, and wetlands, threaten
some populations (Mateo et al. 2009) with observed declines in many localities (Naulleau
and Schätti 1997; Santos et al. 2002). Natural and artificial ponds have small snake popula-
tions that may face local extirpation when these habitats dry up or become contaminated.
Pesticides from agriculture and chemicals from industrial processing are taken up by aquatic
organisms and move along the trophic webs. Consequently, high amounts of organochlorine
compounds such as polychlorinated biphenyl (PCB), hexachlorobenzene (HCB), hexachlo-
rocyclohexanes (HCH) and dichlorodiphenyltrichloroethane (DDT) accumulate in Viperine
Snakes (Santos et al. 1999). Although DDT was banned from the 1970s on, its presence in
snake tissues 20 years later demonstrates that these pesticides persist in the wild for long
periods. Pesticide effects in Viperine Snakes have not been demonstrated but their negative
effects on other organisms are well known. They are presumed to be one of the leading causes
of Viperine Snake decline at the Ebro Delta, northeastern Spain (Santos and Llorente 2009).
In addition to habitat degradation, traffic mortality also affects Viperine Snakes (Llorente et
al. 2005; Meek 2009). In an extensive study, Llorente et al (2005) found that 18% of snakes
found dead on Catalan roads (northeastern Iberia) were Viperine Snakes. Despite being a
docile and harmless snake, people frequently kill it in areas with dense populations (Hailey
and Davies 1987d; Santos, pers. obs.).
Restored freshwater habitats and newly constructed artificial ponds stocked with alien
fish species are rapidly colonized by Viperine Snakes, that opportunistically feed on the rich
diet source (Rugiero et al. 2000; Pleguezuelos and Feriche 2003). The consumption of some
alien fish with hard and long dorsal spines can end fatally during ingestion, as was con-
cluded from snakes found death with recently consumed Lepomis gibbosus and Micropterus
salmoides in the stomach (Santos and García-Cardenete 2006; Alarcos et al. 2009). Large in-
troduced fish species behind dams and in big rivers can also consume Viperine Snakes as, for
example, the Pikeperch, Sander lucioperca (Pérez-Bote and Roso 2009).
Ecological competition with Natrix tessellata in areas of sympatry has also been responsi-
ble for Viperine Snake decline in Italy (Naulleau and Schätti 1997) and Switzerland (Metzger
et al. 2009, 2011). In the latter country, N. tessellata introduction at Geneva Lake appeared to
be a cause of N. maura decline (Mazza et al. 2011), whereas the existence of hybrids has long
been known only from captive specimens, historic hybridization between these species has
been indicated by molecular analysis (Mebert et al. 2011a; Schöneberg et al. 2023; chapter on
N. tessellata herein).

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Aquatic Snakes

Grass Snakes

Eastern Grass Snake

Natrix natrix (Linnaeus, 1758)

Daniel Jablonski and Konrad Mebert

Western Grass Snake

Natrix helvetica (Lacépède, 1789)

Konrad Mebert and Daniel Jablonski

Ibero-Maghrebian Grass Snake

Natrix astreptophora (López-Seoane, 1884)

Juan M. Pleguezuelos and Konrad Mebert

Grass Snakes are among the most common and best-known snakes in Europe and tem-
perate Asia. They are widespread, polytypic and experienced a complex taxonomic history
with four (Thorpe 1979) up to 13 subspecies (Kabisch 1999). Fritz and Schmidtler (2020)
present a cleaned and updated revision of Grass Snakes taxonomy. Finally, molecular-based
studies by Pokrant et al. (2016) and Kindler et al. (2017) on phylogeny and biogeography
revealed genetic barriers between three parapatric groups, partly confirming Thorpe’s mul-
tivariate analysis of quantitative traits of “then” Natrix natrix in the 1970s (Thorpe 1973,
1975a, b, 1979), and leading to the recognition of three Grass Snake species: the Eastern Grass
Snake Natrix natrix (Linnaeus, 1758), the Western Grass Snake Natrix helvetica (Lacépède,
1789), and the Ibero-Maghrebian Grass Snake Natrix astreptophora (López-Seoane, 1884);
described initially as Coluber, Natrix Linnaeus, 1758, Coluber helveticus Lacépède, 1789 and
Tropidonotus natrix var. astreptophorus López-Seoane, 1884, respectively.
However, two comprehensive works were composed of detailed information on the
Grass Snakes before their recent split into three species: Kabisch (1999) with 67 pages and
Blanke et al. (2008) with 312 pages. Therefore, chapters in the following account begin with
summary information on the Grass Snakes species as one group, where they exhibit similar
traits, but add separated information in subsequent paragraphs where the three species differ
from each other, primarily on distribution, morphology, and regional ecology, or present
examples of local studies. The order of species treatment roughly follows the chronology of
their description and body of information: natrix, helvetica, astreptophora.
Distribution and Habitat. The Grass Snake group includes three continents (Hecht
1930; Kabisch 1978, 1999). They occur across most of Europe, north to the limit of the Arctic
Circle, and east to central Asia as far as western Mongolia/China. In the south, Grass Snakes
occur in northwestern Africa and again in the Levant (eastern Mediterranean) from Lebanon,
Israel (historically), and northeast across Turkey to the southern shores of the Caspian Sea.
Grass Snakes inhabit a great variety of wetlands, from small streams to large rivers,

464
 28. The Palearctic Watersnakes

swamps, marshes, and ponds to lakes, where

they find their main prey, anurans. But it is also
found along coastal habitats and can regionally
venture away from water to feed on toads in the
humid countryside. Grass Snakes occupy cooler,
often shadier, aquatic habitats than the largely
sympatric Dice Snake N. tessellata, which prefers
more open and rockier areas and is primarily a
fish predator. The distribution of the single Grass
Snake species is as follows:
Figutre 28.3. The distribution of Natrix natrix. The Eastern Grass Snake N. natrix (type
locality: Fada mill pond, Berga-Tuna Estate,
Nyköping, Sweden) ranges from northern Eu-
rope between Norway, and Finland, with the most northern record from near Gällivare in
northern Sweden (Holm 1934 cit. in Kabisch 1999), south to Estonia, Latvia, Lithuania, Po-
land, Denmark, central (longitude) and eastern Germany, northeastern Switzerland, central
eastern Austria, northeastern Italy, Belarus, Ukraine, Czech Republic, Slovakia, Hungary, Ro-
mania, Moldova, Slovenia, Croatia, Bosnia and Herzegovina, Serbia, Montenegro, Kosovo,
Macedonia, Greece, Albania, Türkiye (Turkey), Russia, Georgia, Armenia, Azerbaijan, Syria,
northeastern Iraq, northwestern Iran, southern Kazakhstan, eastern Turkmenistan, northern
Uzbekistan, northern Kyrgyzstan, northeast to Aral and Baikal Lakes, northern Mongolia and
northwestern China. In the Republic of Georgia, the Eastern Grass Snake is expected in the
humid west, where they are observed in ratios of a couple of dozen N. natrix to one Dice
Snake N. tessellata, latter becoming more abundant in drier landscapes (Frotzler et al. 2011).
This species also inhabits some eastern Mediterranean islands and archipelagos, including
Cyclades and Cyprus but not Crete. In central Europe, there is a contact zone with some hy-
bridization between N. natrix and N. helvetica that runs approximatively along and east of the
Rhine River in Germany and northeastern Switzerland but deviates farther east and passes
through the eastern Alps shared by southeastern Germany, western Austria, and northeastern
Italy (Sindaco et al. 2013; Kindler et al. 2017; Schultze et al. 2020; Asztalos et al. 2021a; di
Nicola et al. 2021; Milko et al. 2021). Cline analyses with genetic markers and color pattern
characters revealed concordant steep clines across those hybrid zones (Fritz et al. 2023). The
presence of the species in Lebanon and Israel might be still possible, but remains currently
unconfirmed and probably very rare due to anthropogenic habitat changes during 20th Cen-
tury (see Martens 1996; Grillitsch and Werner 2009). The Eastern Grass Snake is known for
elevation as high as 2000 m in the Lesser Caucasus Mountains (Tuniyev et al. 2019) with a new
elevational record of 2350 m on the Meskheti Range (authors unpubl. data).
The range of the Western Grass Snake N. helvetica (type locality: Jorat, western Switzer-
land) continues adjacent west to that of the Eastern Grass Snake and stretches from extreme
western Germany across France, Netherlands, Belgium, Luxembourg, United Kingdom, and
reaching Scotland in the north (Cathrine 2014). In the south it ranges across most of Switzer-
land, Liechtenstein, western Austria, Italy, and the Mediterranean islands of Corsica (France),
Sardinia, and Sicily (both Italy). The highest elevation for the Western Grass Snake is 2300 m
in the Italian Alps (Camerano 1891) and 2160 m in Ticino, southern Switzerland (authors,
unpubl. data). Niche modelling of N. h. cetti from Sardinia Island found its habitat to be pos-
itively correlated with higher elevation, precipitation, and accordingly cooler temperatures
(Di Nicola et al. 2023). In the eastern Alps (Austria and Germany), the Western Grass Snake
inhabits the mountainous areas. In contrast, the Eastern Grass Snake occupies the lowlands,

465
Aquatic Snakes

with some hybridization between

them where these two different
topographic areas meet (Asz-
talos et al. 2021a). In far southern
France, N. helvetica contacts but
rarely hybridizes with the Ibe-
ro-Maghrebian Grass Snake N.
astreptophora just north of the
Pyrenean Mountains (Pokrant et
al. 2016). This hybridization is
more frequent in the northeast
than in the northwest of the Py-
Figure 28.4. The distribution of Natrix helvatica. renean Mountains (Asztalos et al.
2020).
The Ibero-Maghrebian Grass Snake N. astreptophora (type locality: La Coruña, Gali-
cia, Spain) occurs southwest of the Western Grass Snake, from the northern coast of Spain,
eastward along the southern slopes of the Pyrenean Mountains to its eastern end where they
crossed north into a few departments of southern France. It occupies the following habi-
tats (IUCN Habitats Classification Scheme v. 3.1): temperate forests, temperate grasslands,
permanent rivers/streams, bog/marshes/swamps, permanent lakes, seasonal lakes, perma-
nent pools, seasonal pools, springs, alpine wetlands, and sandy shoreline (Pleguezuelos et al.
2010). Natrix astreptophora ranges across the entire Iberian Peninsula (Portugal and Spain),
at 0–3,200 m ASL and becomes more montane in central and southern Iberia (González-
Miras et al. 2008). It is homogenously distributed and abundant in the north, inhabiting the
border of beech and mixed forests, dense scrubs, and river valleys, including those with a
high hydrologic seasonality (San-
tos et al. 2002; Escoriza 2018). To
the south, river-associated habi-
tats become more important, but
it also dwells in other artificial
biotopes, such as humid mead-
ows, ponds, and ditches (Santos
et al. 2002; Segura et al. 2007).
It is absent from most of the dry
Guadalquivir Valley and the arid
southeast, where rainfall is below
600 mm (Santos et al. 2002, 2008)
and the Balearic Islands (Kindler
Figure 28.5. The distribution of Natrix astreptophora et al. 2018b). In northwestern Af-
rica, scarce records from northern
Morocco, Algeria and Tunisia are
mainly restricted to wetlands and humid mountainous forest between 0–2000 m. Second-
arily, it occupies rivers and lagoons close to the Mediterranean coast, where amphibians are
numerous, but was occasionally observed also far from water (Schleich et al. 1996; Sindaco
et al. 2013; Kalboussi and Achour 2018; Martínez del Mármol et al. 2019). In areas of fre-
quent forest fires in on the western Iberian Peninsula, N. astreptophora was only recorded in
unburnt plots (Muñoz et al. 2021). On an aridity gradient in the western Mediterranean, the
species mainly occupies the more pristine aquatic habitats (Escoriza and Pascual 2021). Mean

466
 28. The Palearctic Watersnakes

annual temperature and precipitation values for northwestern Africa are 16.9oC and 838 mm,
respectively (Escoriza and Ben Hassine 2017). Within southwestern France, it marginally
occurs in the departments of Eastern Pyrenees, Aude and Ariege (Geniez 2015; Pokrant et al.
2016; Kindler et al. 2018a; Asztalos et al. 2020; Fritz and Schmidtler 2020).
Identification. The morphological variation of the Grass Snake group led in the past to
the description of many subspecies and regional morphs, with the most prominent ones de-
tailed in Kabisch (1978, 1999), that often do not correspond with recently presented molecu-
lar phylogenies. It resulted in many misidentifications and unclear distribution patterns. The
following accounts summarize morphological characteristics for all three species together,
but adds specifics in subsequent paragraphs where each species differs.
Commonly, the Grass Snakes are slender to robust snakes with a total length up to 150
cm, rarely more, with both Natrix natrix and N. helvetica have been reported of exceptional
maximum length > 180 cm (Kabisch 1999), whereas N. astreptophora remains smaller with
112.5 cm SVL for a female from the Iberian Peninsula (García-Antón et al. 2017). Females
attain larger size with a relatively longer tail in males, e.g., 22.7% of total length for males ver-
sus 20.2% for females in southeastern Spain (Feriche et al. 1993). Their dorsal color usually
consists of shades of grey, but brown and even reddish tones can also occur (Jablonski et al.
2022). Four rows of dorsal spots or blotches vary in size depending on geographic variation,
species, and age. Another traditional name for this group is Ringed Snake, which refers to
their white to yellow collar spanning across the neck or being separated into bilateral lunar
spots. The collar is orangish in subspecies N. natrix scutata of Transcaucasian countries and
Russia. On the body and adjacent to the light collar are large, bilateral blotches, often curved
and pointed posteriorly. The light collar may disappear in fully grown individuals, whereas
the post-collar blotches remain, most frequently in adult N. astreptophora. Melanism is wide-
spread and occurs occasionally in single individuals or represents a sizeable proportion in
some populations. The iNaturalist dataset showed that the frequency of dark body coloration
increases from south to north and west to east (Fritz and Ihlow 2022). The ventral pattern is
checkered with black blotches on white ground, with the blotches often exhibiting a concave
outer edge (Gonzalez de la Vega 2021). The black proportion of the ventral body increases
in size posteriorly to become almost entirely black. Main scale characters (Boulenger 1913;
Kreiner 2007; Scali et al. 2011) are dorsal scales keeled and usually arranged in 19 rows mid-
body, less prominent keeled to smooth in the lowest (paraventral) two rows; the keeling is
also reduced or disappears at the end of the tail. Ventral scales vary geographically but are
within 159–189 in males and 155–181 in females, and similarly for subcaudal scales vary
within 56–90 in males and 47–74 in females. All three Grass Snake species share the following
characteristics: Anal scale is usually divided. Preocular scales: 1, rarely divided. Postocular
scales: usually 3, rarely 2 or 4. Upper Labial scales: 7 (rarely 6 or 8); 3rd and 4th touch the eye.
Temporal scales: 1 (rarely 2) anterior and 2 (rarely 3) posterior. R.S. Thorpe has analyzed
more detail on geographic variation in multiple studies (see his references from 1973–1989).
In the Eastern Grass Snake Natrix natrix northern populations are usually greyish, indi-
vidually in different tones, and with four rows of small dorsal spots. In southern populations
of N. natrix, the dorsal pattern contrasts with four rows of larger dorsal blotches, and the
lateral ones extended to vertical bars. Individuals with two light dorsal lines along the body
(persa morph) occur in southeastern populations from the Balkans to Turkey, Caspian Sea,
and Lebanon (subspecies vulgaris, moreotica, scutata, and possibly syriaca). Some distinct
color pattern expression on insular populations led to descriptions of different subspecies
and morphs (e.g., N. n. fusca from Kea Island in Greece, N. n. gotlandica from Gotland, Swe-
den, N. n. cypriaca from Cyprus, or N. n. schweizeri from the southern Cyclades, e.g., Milos

467
Aquatic Snakes

Figure 28.6: Natural color pattern variation of Eastern Grass Snakes (Natrix natrix) from: A) Poland;
B) Slovakia; C) northeastern Turkey; D) southwestern Turkey; E) and F) Milos Island, Greece. Photo
credit: A), C), D) Konrad Mebert; B) Daniel Jablonski; E) Andre Schmid; F) Laura and Bobby Bok.

468
 28. The Palearctic Watersnakes

Island, Greece), that are invalid from the evolutionary point of view (Kindler and Fritz 2018).
Interestingly, N. n. cypriaca and N. n. schweizeri have similar body coloration variation: first,
the more contrastingly heavily blotched morph on light gray, without or only partly visible
light collar band, a second wholly or partly melanistic individual. And third a stippled morph
with more minor black and olive-grey flecks, the so-called picturata variation (Baier et al.
2009; Zotos et al. 2021). However, current genetic data (Kindler et al. 2013; Asztalos et al.
2021b) support the existence of only four subspecies of N. natrix without any insular endem-
ics, even though the southern subspecies, including the polymorphic Aegean populations of
N. n. moreotica, show generally a higher variability than northern ones (Jablonski et al. 2023a;
and see below). Thus, these insular phenotypes apparently represent adaptation to local en-
vironments in the Mediterranean area. Their scalation characteristics are those of the entire
Grass Snake group, however, with a restricted range in the number of ventral and subcaudal
scales, 163–188 and 53–89, respectively. The number of ventral scales of males is slightly high-
er on average than in females, whereas the number of subcaudal scales is significantly higher
in males. The Eastern Grass Snake is slimmer and smaller than western species, with patches
behind the head usually richer in yellow to orange, especially in the easternmost populations,
ssp. scutata, that is generally missing the lateral vertical bars. Natrix natrix females are more
robust and reach longer sizes than males. Specimens > 120 cm in total length are uncommon
to rare, with the largest specimen found killed on Krk Island, Croatia, and subsequently mea-
sured as 205 cm (Schreiber 1912; Kabisch 1999). The weight of the Eastern Grass Snake from
central Europe ranges in males from 100–140 g, in females 300–350 g (Günther and Völkl
1996; Kabisch 1978, 1999).
The Western Grass Snake Natrix helvetica dorsal pattern usually consists of two rows of
dorsal spots/blotches and vertically elongated lateral bars. The black nuchal blotches are often
only little indicated (Kabisch 1999). Specimens of the Italian Grass Snake N. helvetica sicula
from far southern Italy, e.g., Sicily Island and southern Calabria, are characterized for their
orange snout tip and chin. Melanistic specimens are known. The subspecies N. helvetica cetti
from the islands of Sardinia and Corsica (Vanni and Cimmaruta 2011; di Nicola et al. 2021;
Schultze et al. 2020) show a more distinctive morphology: Ventral scales: 158–178 (males
160–178, females 158–173). Subcaudal scales: 47–64 pairs (males 56–64, females 47–53).
Preocular scales 1, but often 2 at least on one side of the head. Adult individuals of the Western
Grass Snake sometimes lack the typical light collar band.
An external character of Natrix astreptophora relates to its epithet “astreptophora”
(López-Seoane 1884: p. 15) that denotes the absence of the white to the yellowish collar in
adults, which is still present in juveniles (Braña 1998; Fritz and Schmidtler 2020). Besides
the black blotches posterior to the light collar standard in most Grass Snakes, some juvenile
N. astreptophora also show an additional distinctive black parietal band anterior to the light
collar band (Pleguezuelos 2018). Most specimens show a reddish iris, the most visible external
diagnostic character to distinguish against the parapatric N. helvetica (Pokrant et al. 2016),
but also exhibit a black patch crossing diagonal over the lower anterior portion of the iris that
also occurs in some southern N. natrix and N. helvetica (Gonzalez de la Vega 2021). Newborn
body color is brown to olive or ocher after the first ecdysis, whereas adults are grey with the
occasional reddish-brown morph (Gonzalez de la Vega 2021). On the dorsum, 3–6 longitu-
dinal rows of 41–56 black spots often disappear in adults (pattern less), and rarely fuse across
the back, as in other Grass Snakes (Geniez 2015). Up to 5.4% of individuals in Cantabrian
Mountain populations are melanistic (Meijide and Pérez-Melero 1994; Albadalejo 2008). The
Ibero-Maghrebian Grass Snake has a wide head and stout muscular body, showing differences
in skull bones from the other Grass Snakes (Pokrant et al. 2016).

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Phylogeny and phylogeography. The extant diversity within Grass Snakes (Deepak et al.
2022) suggests that the group originated in Asia, afterward dispersing to the western Paleart-
ic, where they diversified into the current species of the genus (Schöneberg et al. 2023). First
comprehensive morphological studies on Grass Snakes by Thorpe (e.g., 1973, 1975a, b, 1984,
1979) were followed by molecular studies (Guicking et al. 2006; Fritz et al. 2012; Kindler et
al. 2013). They evaluated phylogeographic structure and time of divergences with subsequent
studies of hybrid zones, morphology, and osteology (Pokrant et al. 2016; Kindler and Fritz
2018; Kindler et al. 2017, 2018a,b; Schultze et al. 2020; Asztalos et al. 2020, 2021a, b), finally
resulting in significant changes in the taxonomy and phylogeny of the traditional Grass Snake
Natrix natrix, by recognizing three well-separated parapatric species. The Ibero-Maghrebian
Grass Snake N. astreptophora was the first to diverge from other Grass Snakes in Europe at
9.6–10.6 MYA, resulting from the uplift of the Pyrenean Mountains (Pokrant et al. 2016). This
was reanalyzed and adjusted to a slightly younger split by Schöneberg et al. 2023, see below.
Subsequently, the Grass Snakes in Europe split into the Eastern Grass Snake N. natrix and
the Western Grass Snake N. helvetica approximately 8.2 to 7.3 MYA (Fritz et al. 2012; Kindler
and Fritz 2018; Kindler et al. 2017). The Big-head Grass Snake N. megalocephala (Orlov and
Tuniyev, 1987) from the Caucasian region and northeastern part of the Black Sea coast was
often listed as a separate species. However, the molecular-phylogenetic analysis showed that
this taxon is only a phenotypic variation, possibly an ecomorph or ecophenotype of N. natrix,
occurs more widespread as far as Switzerland (Göçmen et al. 2011a) and was ultimately syn-
onymized with this species (Kindler et al. 2013; Asztalos et al. 2021a).
The Eastern Grass Snake N. natrix continued diversification around 6 MYA (Kindler et
al. 2017) with the following, currently recognized subspecies (Kindler and Fritz 2018; Fritz
and Schmidtler 2020; Asztalos et al. 2021a, b): N. n. natrix (Linnaeus, 1758); N. n. vulgaris
(Laurenti, 1768); N. n. moreotica (Bedriaga, 1882), N. n. scutata (Pallas, 1771), with one po-
tential subspecies, N. n. syriaca (Hecht, 1930), that requires more attention. According to
new data based on mitochondrial and nuclear DNA, N. n. natrix occurs in Sweden, Norway,
Finnland, Denmark, and most of Germany, except the western part. It forms possible broad
contact zones with the subspecies N. n. vulgaris in Poland, the Czech Republic, and Slovakia.
This subspecies is probably widespread in Poland, southern Central Europe, northeastern It-
aly at least as far south as Argenta and Ravenna, and large parts of central Balkan countries. It
is replaced in western and southern Balkan and Western Turkey by N. n. moreotica. The areas
farther east, from the Baltic republics through Ukraine, the Middle East and Iran to Mongolia
is inhabited by the subspecies N. n. scutata (Asztalos et al. 2021a, b; Jablonski et al. 2023b)
Diversification in the Western Grass Snake N. helvetica began after establishing the old-
est lineage for in the most southern (Italian) populations with an estimated mean age of
6.8 MYA. Currently, three or four subspecies are recognized: first, N. h. helvetica of western
Europe, and second N. h. sicula (Cuvier, 1829) of Italy that includes now also N. h. lanzai
(Kramer, 1970). The Tyrrhenian Grass Snakes from the Italian island of Sardinia, N. h. cetti
(Gené, 1839), and the neighboring French island of Corsica, N. h. corsa (Hecht, 1930), repre-
sent the third and fourth subspecies, or they become synonymized with each other, pending
further investigation (Fritz et al. 2012; Kindler and Fritz 2018; Fritz and Schmidtler 2020;
Schultze et al. 2020). The nominotypic Natrix h. helvetica also occurs across most of Swit-
zerland and France, and to northern Netherland including Belgium and Luxembourg (van
Riemsdijk et al. 2020; Ahnelt et al. 2021) and Great Britain, barely reaching into Scotland
(Cathrine 2014);southwards it reaches the Spanish Valle de Arán in the northern slope of the
Pyrenean Mountains. It introgresses in northwestern Italy into N. helvetica sicula, the Italian
Grass Snake, that primarily inhabits mainland Italy and Sicily Island, but also southern Swit-

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zerland and across the Alps of Western Austria into far southern Bavaria, Germany (Schultze
et al. 2020; Asztalos et al. 2021a, b).
The Ibero-Maghrebian Grass Snake N. astreptophora is the sister taxon to N. helvetica.
These two Grass Snakes species from western Europe and northweastern Africa form the sis-
ter clade to the older group of Eastern Grass Snake N. natrix (Schöneberg et al. 2023). Natrix
astreptophora split from the Western Grass Snake N. helvetica in the upper Miocene (7.3 Mya,
confidence interval 5.86–10.59 Mya; Schöneberg et al. 2023). There is also morphological and
molecular evidence for some hybridization between N. astreptophora and N. maura (Gonzalez
de la Vega 2021; Schöneberg et al. 2023). Natrix astreptophora is genetically structured into
three clades. First, the European (Portugal and Spain) and northwest African N. astreptophora
diverged 5.44 MYA from each other (Kindler et al. 2018a). Second, the North African Grass
Snakes split again into two groups on both sides of the Mouluya Valley, forming the Moroccan
and Tunisian/Algerian clades. Both previous splits likely occured as a result of the Zanclean
flooding event of the dried Mediterranean Basin at the end of the Messinian Salinity Crisis
(Kindler et al. 2018a). Latter group was subsequently re-established as N. astreptophora al-
gerica (Hecht, 1930) by Fritz and Schmidtler (2020). On the northern side of the Pyrenean
Mountains in southeastern France, N. astreptophora and N. helvetica are parapatric, with a
steep genetic transition from one taxon to the other, and rare hybridization (Pokrant et al.
2016; Asztalos et al. 2020). This contact zone had established after the mid-Holocene range
expansion from glacial refuges on the Iberian Peninsula (N. astreptophora) and southwestern
France (N. helvetica).
Population Aspects. Newborns of all three Natrix species exhibit a total length of 14–22
cm (usually 18–19 cm) and weigh around 2–4 gr (Kabisch 1999). Before their first hiberna-
tion, they reach a total length of 25–30 cm, to 50–60 cm at five years, to 70 cm at 7–8 years, and
to > 100 cm at 15 years of age; usually only females’ growth larger than 80 cm of total length,
some of them reaching 17–18 years of age (Madsen 1983; Braña 1998). Females grow faster
than males but require more time to mature and generally attain greater length and weight.
In N. natrix, males reach sexual maturity at about 50 cm total length and 50 g weight at the
age of three years, whereas males of N. astreptophora in southeastern Spain, mature at 30 cm
snout-vent length (SVL; approximately 40 cm total length). Females of all Grass Snake species
reach sexual maturity in the fourth or fifth year of life, at approximately 60 cm SVL and 80
g in weight (Jusczyk 1974; Madsen 1983; Rehák 1992; Feriche 1998). In northern Morocco a
male at 38.7 cm snout-vent length and a female at 67.8 cm snout-vent length were adults (Fahd
2001). After the age of 8–9 years, growth slows down and may become negligible (Madsen
1983). However, locally reduced food availability can result in females growing slower and to
a smaller size; for example, an insular population in Sweden that feeds exclusively on generally
smaller newts (the only available amphibians) than females from the mainland that were feed-
ing on larger anurans (Madsen and Shine 1993). The sex ratio of N. natrix for males:females
from northern populations ranges from 1:1.36 to 1:1.5, respectively (Rehák 1992, refs. in Ka-
bisch 1999), whereas those from Iran were 1:1 (Ahmadzadeh et al. 2011), for N. astreptophora
1:1.9 in northern Spain (Braña 1998) and 1:1.4 in southeastern Spain (Feriche 1998).
Grass Snakes can be very common in suitable habitats with about 1–7 individuals/ha and
reach high densities of close to one individual per square meter during aggregation for mating
and feeding when mates and prey temporarily occur at high densities (Tuner 1977; Mertens
1995; Kabisch 1999 and refs.; Sidorovich et al. 2008). The mean total home range for individ-
uals in Sweden and Germany were 21.2 ha and 16.4 ha, respectively, and the adjusted home
range that excludes large areas never used by the snakes showed a mean of 11.8 ha and 15.1 ha,
respectively (Madsen 1984; Mertens 2008), and increased to 39,7 ha in agricultural impacted

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area, where seasonally required sites were more distant to each other (Wisler et al. 2008). In-
deed, seasonally varying locations have been found for different annual activities, including
hibernation, mating, oviposition, and foraging in N. natrix in Germany and N. helvetica in
the Netherlands, with migratory distances of up to 7 km between seasonal sites (Janssen and
Völkl 2008) Maximum age by mark-recapture methods was given as 28 years from a Swedish
population (Edelstam 1989).
Activity and Thermal Biology. In Central Europe, the Eastern Grass Snake starts
emerging from hibernation (rodent burrows, rock crevices, cellars, composts, piles of leaves,
subterranean tree roots) in late March or early April and remains active until late September
to early October. In southern France, they emerge from hibernation usually on day 95 (Julian
calendar; Prodon et al. 2020). Exceptionally, this species can also be found in November or
in February, for example, during short spells of warmer temperatures, or probably is more
active throughout the year in warmer regions of southwestern Asia, e.g., Azerbaijan (Rehák
1992, D. Jablonski, unpubl. data). In spring and autumn, activity peaks in the warmest hours
of the day, usually mid-day into the afternoon, whereas the summer activity has two peaks, in
the morning, beginning between 0630 h and 0800 h, and late afternoon/evening, respectively
(Mertens 2008). During hot days, activity can occasionally be switched to nights. In contrast,
nocturnal activity can be added in the cooler spring, April and May, when high amphibian
breeding activity (e.g., Albania, Northern Macedonia, Switzerland, Germany, Mertens 1994,
2008, authors pers. obs.). Referenced accounts in Kabisch (1978, 1999) show that N. natrix
(and likewise N. helvetica) experience a wide range of environmental temperatures during
daytime activity, as low as 6° C and > 30° C, but usually somewhat between 18–25° C (Gre-
gor 1980; Kühnel 1993), with preferred body (cloaca) temperatures ranging from 29–33° C
(Mertens 1994, 2008). Severe cold weather during the winter months is survived in hiber-
nacula, sometimes with other reptiles such as the Northern Viper (Vipera berus), Smooth
Snake (Coronella austriaca), or Slow Worm (Anguis fragilis). Mertens (2008) radiotelemetry
hibernating Grass Snakes in complex rodent burrows and found their subterranean position
constantly at 5–15 cm depth, and only during a freezing period with temperatures below –10°
C snakes move into lower layers, between 25–50 cm depth.
Often, the visibly highest activity relates to the mating period in the spring months when
males search for females. For example, in a mosaic landscape of woods, pastures, agricultural
fields and ponds in Sweden, Madsen (1984) observed an average movement of 54.8 m per day
by Grass Snakes, whereas later in the summer, they remained more sedentary (average move-
ment per day 13 m). However, neither a higher movement of male Grass Snakes nor a re-
duced summer activity was found in a similar study and habitat in central Germany (Mertens
2008). Instead, larger individuals followed large toads that had left the ponds/pools into more
lightly forested sections, whereas smaller individuals remained in the ponds to forage on
water frogs. Yet only 4.8% of telemetered individuals were detected in water (Mertens 2008).
The Western Grass Snake N. helvetica yields a comparable variation of activities and
thermal biology as its eastern sister species (see accounts in Kabisch 1999). For example, in
western Switzerland, female N. helvetica used monocultures managed by a cropping system,
e.g., wheat, potato, and corn (Wisler et al. 2008). The monocultures provided suitable basking
sites, favorable foraging grounds, and low pressure from avian predators. Adjacent edge hab-
itat (dam, canal bank, forest edge) was preferred for the period of shedding and oviposition.
After oviposition, the females returned to the monocultures to hunt amphibians primarily.
The mean home range size was 34 ha (Wisler et al. 2008). While the latter study found no
nocturnal behavior, night foraging was observed in populations of the Western Grass Snake
occupying warmer regions, exemplified by the predominantly nocturnal in the Sardinian

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Figure 28.7: Natural color pattern variation of Western or Barred Grass Snakes (Natrix helvetica) from:
A) and B) Ticino, southern Switzerland; C) western Switzerland; D) Sardinia Island, Italy. Photo cred-
it: A), B), C) Konrad Mebert; D) Matteo Riccardo di Nicola.
subspecies N. helvetica cetti (Capula et al. 1994).
The activity season of the Ibero-Maghrebian Grass Snake N. astreptophora ranges between
March and October in southeastern Spain (Feriche 1998) and between the end of February
and early November in the western part of the Iberian Peninsula (Galán and Fernández-Arias
1993; Malkmus 1997), with most records from April to September (Galán 1988). In northern
Morocco, active individuals have been found between May and October (Fahd 2001), whereas
in Tunisia, N. astreptophora algerica is also active during mild winter days (Kalboussi and
Achour 2018). Communal hibernation dens with up to three individuals have been detected
in Galicia (Galán and Fernández-Arias, 1993). In southeastern Spain, the mean annual body
temperature of active individuals was 26.6° C (range 22–32° C), higher during summer with
27.8° C (24.3–32.0° C), less with 26.8° C (19.0–30.0° C) during spring and autumn (Hailey and
Davies 1986a; Hailey et al. 1982).
Reproduction. All three species of Grass Snakes appear to have a similar reproductive
base strategy, which likely has been adapted to local environmental conditions rather than
species constraints. In general, the mating period for N. natrix begins soon after emergence
from hibernation, in March/April, and can last into May/June, followed by oviposition be-
tween June and August (Feriche 1998; Kabisch 1999; authors pers. obs.). Males begin court-
ship primarily after first shedding, raise their head next to a receptive female, twitching their
bodies, frequently tongue flicking, and pressing their bodies against the female, wrapping
their tail around her tail. Then, with the opening of the female´s cloaca, one of the hemipenes,
rarely both, penetrates. Copulation takes several minutes, sometimes hours (Rehák 1992).
There are mating aggregations with up to 50 snakes (Berec et al. 2015), while some engage in
a mating ball consisting of one receptive female with up to 20 males entangling her (Kabisch
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1999). Larger males are more successful in this case, even though no direct fights for females
have been observed (Luiselli 1996).
Mating in autumn (September/October) occasionally occurs in Germany, with egg
clutches discovered in November/December (Kabisch 1999 and refs. therein). Spermatogen-
esis in males persists until the end of the mating season. After mating, sperm remains in the
female’s body, and fertilization is delayed until the onset of ovulation next spring (Berec et al.
2015). In northern Iran (N. natrix) and Spain (N. astreptophora), females start vitellogenesis
in May/early June, with the growth of ovaries continuing at least until mid-July to August
(Galán and Fernández-Arias 1993; Feriche 1998; Ahmadzadeh et al. 2011; Shiravi et al. 2012).
In cool and rainy Asturias, northern Spain, there are female N. astreptophora with oviductal
eggs during July and embryogenesis in the early stages at oviposition (Braña 1998). Males
from Iran (N. natrix) and Spain (N. astreptophora) showed signs of a postnuptial spermato-
genesis with slow and constant growth in late spring, reaching its height in August and Sep-
tember, ending in November, with sperms stored over the winter for the next spring mating
(Faghiri et al. 2011; Arragayago and Bea 1988). Sperm storage may last long, exemplified by
a solitary kept Grass Snake that produced fertilized eggs after > 2 years (Hediger 1971; cit. in
Eckstein 1993).
Oviposition ranges from June to August, with clutch size from 8–32 eggs per female (Ka-
bisch 1999). Clutch size can geographically range, for example, for N. astreptophora: 9-26 eggs
in Asturias, northern Spain (Braña 1998), 12-29 eggs in southeastern Spain (Feriche 1998),
6-50 eggs in Galicia (Galán and Fernández-Arias 1993), 10-50 eggs in all of Morocco (Bons
1967), and 11–28 eggs only in northern Morocco (Fahd 2001). Clutch size is also positively
correlated with female body size and age for all Grass Snake species (Madsen 1983; Rehák
1992; Feriche 1998). For example, petite females from a food-depauperated habitat on Got-
land Island, Sweden, laid only 2–3 eggs (Edelstam 1989), whereas huge females exceptionally
deposited around 70–100 eggs (Kabisch 1999 and refs. therein). The size of the egg ranges
from 21–40 x 11–24 mm, and the eggs increase in weight by ca. 30% during the incuba-
tion (Kabisch 1999). Clutches are deposited in manure piles, sawdust, and rotting vegetables
under rocks and rodent burrows. Several females might lay clutches in one suitable place,
sometimes with other species, e.g., Natrix maura or Aesculapian Snake Zamenis longissimus.
Communal nests have been recorded in different countries, e.g., 400 hatchling Grass Snakes
emerged from one nest in Galicia, Spain (Galán and Fernández-Arias 1993), 558 eggs in the
Czech Republic (Berec et al. 2015), and even between 3000–4000 eggs in a pile of sawdust in
eastern Germany (Kabisch 1999). Depending on their geographical location, the young hatch
from July to September after an incubation period of approximately two months.
Foraging and Diet. Grass Snakes are diurnal, terrestrial, and visually guided active hunt-
ers (Hailey et al. 1982; Hailey and Davies 1986a, b), that hold on to their slippery prey until
the ingestion process begins (Galán and Fernández-Arias 1993). However, they switch to cre-
puscular and nocturnal foraging during summer or in generally warmer regions in the south
(e.g., Hailey and Davies 1986a; Schleich et al. 1996, pers. obs.). For example, Ibero-Maghrebi-
an Grass Snakes, N. astreptophora from Galicia, Spain, avoided activity during the hot midday
hours in captivity but under natural light and temperature conditions (Galán 1988). There are
also a few reports of marine observation, some for foraging, e.g., a Grass Snake that became
entangled in a fishing net at 8–9 m depth in Norway, whereas others might be drifted away by
sea currents, one up to 47 km (Kabisch 1999 and refs.). Small prey can be swallowed under-
water, while larger prey is carried and then swallowed on land (Schleich et al. 1996). One prey,
the Natterjack Toad Epidalea calamita, avoids shelter containing the scent of N. astreptophora
(Gonzalo et al. 2008).

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Studies have revealed a broad diet for Grass Snakes, yet always with a focus on common
amphibians, primarily frogs of the genus Rana and Pelophylax, and toads of the genus Bufo,
rarely the genus Pelobates (Kabisch 1999). In contrast, other prey types are less frequently con-
sumed, e.g., poisonous amphibians such as the Fire-bellied toads Bombina spp., salamanders
and newts, lizards, invertebrates, birds, eggs, and mammals (Rehák 1992; Luiselli et al. 1997;
Scali et al. 2011; di Nicola and Bruni 2020). Grass snakes consume a lot of available food at
once and can fast for a long time, reportedly until about one year (Dobreff 1939). During the
metamorphosis of B. bufo tadpoles, N. natrix can eat many juvenile toads (Berec et al. 2015).
For more information on the diet across all three grass snake species, see also: Luiselli and
Rugiero (1991); Gled-Owen (1994); Filippi et al. (1996); Reading and Davies (1996); Luiselli et
al. (2005); Faraone et al. (2010); and a particularly extensive account given in Kabisch (1999).
Some specific diet studies on the Eastern Grass Snake N. natrix show the proportion of
fish consumed: 4.1% of the catch in Bulgaria (Beshkov and Dushkov 1981), 40.8% in Croatia
with the remainder being amphibians (Janev Hutinec and Mebert 2011), whereas a Swedish
populations fed mainly on fish from the seashores (Andrén and Nilson 1981). In an alpine
population of N. natrix in northeastern Friuli Venezia Giulia, Italy, anurans were the most
frequent diet (Bufo bufo 46%, Rana temporaria 40%), followed by urodeles (Salamandra atra
6%, Ichthyosaura alpestris 5%), but also a few lizards (Zootoca vivipara) and mice (Apodemus
sylvaticus) were consumed (Luiselli et al. 1997).
In southeastern England, the diet of small and medium-sized of the Western Grass Snake
N. helvetica is mainly based on aquatic prey with a predominance of anurans (63%, primarily
introduced Pelophylax spp.), fish (25%) and rarely a bird (1%). Grass snake usually swallowed
prey headfirst (65%), however, this depended on the prey type (Gregory and Isaac 2004). The
principal prey of N. helvetica on the Italian mainland consisted primarily of the toad species
Bufo bufo and the frog genus Rana, and rarely included rodents, fish, saurians, invertebrates
and passerine (Scali et al. 2011; Di Nicola & Bruni, 2020). The main prey of Sardinia´s N.
helvetica cetti were the frogs Discoglossus sardus and Hyla sarda, and rarely lizards such as
Podarcis tiliguerta (Scali et al. 2011). Amphibian larvae, including those of newts, are food
mainly for juveniles.
Data from Galicia, Asturias, and Levante for the Ibero-Maghrebian Grass Snake N. as-
treptophora, show the major prey are amphibians. Species include Discoglossus galganoi, Bufo
spinosus, Alytes obstetricans, and the urodele Salamandra salamandra and less often oligo-
chaetes, insects and microtines (Braña 1998; Pleguezuelos 2018). Furthermore, prey species
recorded for N. astreptophora are anurans (Epidalea calamita, Pelophylax perezi, Rana tempo-
raria, R. parvipalmata, R. iberica), urodeles (Lissotriton boscai, L. helveticus, Triturus marmor-
atus, T. alpestris) and amphibian larvae, fishes (Leuciscus sp., Phoximus sp.), other vertebrates
(Riparia riparia), and insects (Garzón 1974; Valverde 1974; Vericad and Escarré 1976; Galán
1988; Meijide 1989; Galán and Fernández-Arias 1993; Braña 1998; Díaz 2004; Galán and Fer-
reiro 2010; Ayres 2012; S. Busack per. com.). Frequency of amphibian prey is between 80–98%,
but certainly will vary by season and location. In Morocco, the anurans Discoglossus scovazzi,
Pelophylax saharicus, and the lizard Chalcides sp. have been recorded as prey (Bons 1967). In
Tunisia, N. astreptophora is a potential predator of Bufo spinosus (Ben Hassine and Escoriza
2014).
Predators and Defense. The Eastern Grass Snake N. natrix faces a broad spectrum of
vertebrate predators, from fish to reptiles, and carnivorous mammals including domestic cats
(Galán 2013), but primarily from dozens of different bird species (Kabisch 1999). Young Grass
Snakes are frequently preyed on even by invertebrates such as the Golden Ground Beetle
(Carabus auratus), but also by anurans, passerines, chickens, hedgehogs, moles, and mice (Ka-

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bisch 1999). For all Grass Snakes, the Short-toed Snake Eagle (Circaetus gallicus) has a partic-
ular reputation as a formidable snake hunter (Pleguezuelos 2018; Pleguezuelos and Ontiveros
2010). According to Zebe (1936, cit. in Kabisch 1999), an adult pair of Short-toed Snake Eagle
and their single young may consume up to 1000 snakes per season, although when preying
on larger colubrids compared to smaller vipers, the figure of 450 snakes per eagle family and
season, is more credible (unpub. data of the authors). In the Minsk region of Belarus, Short-
toed Snake Eagles almost exclusively preyed on N. natrix (Glutz von Blotzheim et al. 1971)
Likewise for the Western Grass Snake N. helvetica from Loir-et-Cher, France, where the
Short-toed Snake Eagle only fed on that species and the Asp Viper Vipera aspis (Perthuis
2008), which locally represented the most frequent and sizable snake species (Maumary et al.
2012). However, in the Montados of southern Portugal, this snake rarely appears in the diet
of the Short-toed Snake Eagle (Onofre and Sampaio 2020). Furthermore, all Grass Snakes
are preyed by herons and egrets, storks, owls, kites, falcons, Egyptian Vulture, otters, felines,
martens, and also by other snakes, in particular by the Western Montpellier Snake Malpolon
monspessulanus preying on N. helvetica and N. astreptophora, whereas the Eastern Montpel-
lier Snake M. insignitus preys on N. natrix (Díaz-Paniagua 1976; Schleich et al. 1996; Kabisch
1999; Vacher and Geniez 2010; Pleguezuelos 2018).
The Eastern Grass Snake N. natrix is infected by many parasites. From Iran, Yosse-
fi et al. (2014) reported Rhabdias fuscovenosa (Nematoda), Telorchis assula (Digenea), and
Ophiotaenia europaea (Cestoda). In Turkey N. natrix was infected by five species of Digenea:
Astiotrema monticelli, Encyclometra colubrimurorum, Macrodera longicollis, Paralepoderma
cloacicola, and Telorchis assula; two species of Cestoda: Ophiotaenia europaea and Spirometra
erinaceieuropae; and two species of Nematoda: Rhabdias fuscovenosa and larvae of Eustron-
gylides excisus (Yildirimhan et al. 2007, and refs. therein for other countries). Lewin (1992)
reported 21 types of parasites from Poland, including six adult trematodes, five meso- and
metacercariae, one cestode larva, five adult nematodes, two nematode larvae, one acantho-
cephalan and one tick. A more recent review from Poland revealed > 25 species of parasites
for N. natrix (Kuśmierek et al. 2020; Belcik et al. 2022). There is no comparable information
on the Western Grass Snake N. helvetica whereas the Digenian trematode Leptophallus ni-
grovenosus has been found in the Ibero-Maghrebian Grass Snake N. astreptophora (Navarro
et al. 1987).
Adult Grass Snakes typically flee by the approach of a potential predator, with the fleeing
distance being rather short (Schleich et al. 1996). When cornered without a reasonable escape
option, a Grass Snake may coil the body and hide the head in the center or under a body coil,
or flattens the entire trunk, and transforms the head into a triangular shape by displacing the
quadrate bones. It may hiss loudly and strike with a closed mouth in an example of Batesian
mimicry of coexisting Vipera species (de Solan et al. 2020), but they rarely bite (Kabisch 1999;
di Nicola et al. 2023). Some individuals even raise their forebody and may also spread their
necks horizontally like cobras (Schweizer 1911 cit. in Kabisch 1999; Pokrant et al. 2017; pho-
tos in [Link]). The throat can also be inflated vertically, and even open mouth strik-
ing or biting has been observed but is extremely rare (Kabisch 1999; authors unpubl. data). If
these behaviors do not deter the potential predator, a Grass Snake begins to simulate death by
adopting a state of thanatosis (faking death) by becoming limp, open its mouth and let their
tongue hang out, sometimes enhancing the deterrence effect by mouth bleeding; often also
turning on their back to display the contrasting black and white belly (Eckstein 1993; Kabisch
1999; Malkmus and Sauer 2013; unpubl. data by authors). When handling Grass Snakes, they
often regurgitate their stomach content, emit an obnoxious fluid from their anal glands, and
sometimes defecate.

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 28. The Palearctic Watersnakes

Figure 28.8: Natural color pattern variation of Ibero-Maghrebian Grass Snakes (Natrix astreptophora)
from: A), B) and G) southern France; C) northwestern Spain; D) northern Morocco; E) Portugal; F)
northern Spain. Photo credit: A), B), G) Konrad Mebert; C) and F) Octavio Jiménez-Robles; D) Ab-
dellah Bouazza; E) Benny Trapp.
The white-orange collar band may function as an anti-predator mechanism by mimicking
aposematically colored unpalatable insects (Madsen 1997). In northwestern Spain large toads
Bufo spinosus avoid juveniles of N. astreptophora that display the striking collar pattern (Galán
and Fernández-Arias 1993). Finally, defensive behaviors occur at lower body temperatures,
while aggressive behaviors are more common at higher body temperatures (Eckstein 1993).
Conservation. Grass Snakes have suffered a decline across their entire range, but most
severely in densely populated regions of central and western Europe and in landscapes heavily
used for agriculture in Europe and west Asia (Blanke et al. 2008). These human-made modifi-
cations resulted in habitat fragmentation and the decline of local amphibian populations that

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added to the threats of Grass Snakes and justified its listing as threatened in many central
European countries.
Other human activities also have very negative consequences. It is well known that
many fishermen kill these snakes due to their fear of these semi-aquatic snakes competing
for fish. High mortality caused by traffic, especially when roads cross traditional migra-
tion corridors near water, is also an important source of snake decline (e.g., Ioannidis and
Mebert 2011; Gezova and Jablonski 2018). Gruschwitz et al. (1993) included seven studies,
and Blanke et al. (2008) produced a complete summary with 24 articles on primarily con-
servation aspects of N. natrix and N. helvetica from Germany, Netherlands, Switzerland,
and Cyprus. They proposed a 4-step-concept species action plan: 1. basic investigations
on historical and present distribution/habitat; 2. protection and management of existing
populations, including maintaining and improving foraging, oviposition, and hibernation
sites; 3. re-establish former distribution (where possible) by constructing corridors between
isolated populations, and potentially support weak populations by head-starting with cap-
tive-reared snakes; 4. conduct ongoing monitoring. For the Western Grass Snake N. hel-
vetica a study by Meister et al. (2012) suggested that conservation actions in landscapes
altered by humans should focus on maintaining a habitat mosaic with anuran breeding
ponds and adequate oviposition sites. Luckily, in most regions where it is still present, the
Grass Snakes are considered a common snake species and accordingly listed with the IUCN
category Least Concern. However, the status of the Eastern Grass Snake (N. natrix) popu-
lations along its southeastern borders in some Asian countries is very poorly known (Iraq,
Lebanon, Mongolia, or Turkmenistan; Afrasiab et al. 2011; Asztalos et al. 2020). Also, the
Critically Endangered Cyprus Grass Snake is at risk of extinction, despite some recent new
findings (Blosat 2008; Zotos et al. 2021).
Alien Grass Snakes may negatively impact the conservation of native populations like
the Eastern Grass Snake N. natrix that were introduced into indigenous populations of N.
helvetica in the Netherlands, where they likely hybridize (van Riemsdijk et al. 2020). Oth-
er related examples of alien Eastern Grass Snake species introduced or accidentally trans-
located into local native Western Grass Snake populations in England, France, Italy, and
Germany can be found in Asztalos et al. (2021c). In addition, there are two locations in
Switzerland where alien N. natrix were introduced into native populations of N. helvetica
and have persisted there through decades. In the first location in western Switzerland the
alien N. natrix hybridized with the native N. helvetica (Dubey et al. 2017). In contrast, no
such introgression was found between these Grass Snake species in the second location, in
central Switzerland (Kieffer Merki et al. 2018).
Natrix astreptophora is not listed in the global IUCN red list but is categorized as Least
Concern in Spain and Portugal (Santos et al. 2002; Cabral et al. 2005), and as Near Threat-
ened in Morocco (Pleguezuelos et al. 2010). Within the Iberian Peninsula, threats differ
by region; in the northern belt, the species is rather abundant and not threatened (Galán
and Fernández-Arias 1993; Santos 2008). However, in northeastern Spain, its presence is
negatively influenced by the presence of alien fishes (Escoriza 2018). In central and south-
ern Iberia, the populations are much smaller and survive in ever fewer water bodies that
are often heavily degraded (Santos 2002). Consequently, the number of records of the Ibe-
ro-Maghrebian Grass Snake in the last 40 years decreased by half in Spain and by even
two thirds relative to the number of non-snake reptiles in Languedoc-Roussillon, southern
France, suggesting upgrading its current IUCN threat category under the A2 criterium of
the red listing process (Santos et al. 2022). In northern Morocco, most of the populations are
restricted and isolated to mountainous areas and experience high vulnerability to climate

478
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change (Martínez-Freiría et al. 2013; Escoriza and Ben Hassine 2017). In Algeria, the popu-
lations are also threatened by drastic degradation and loss of their aquatic habitats (Beddek
2017). The fungal pathogen Ophidiomyces ophidiicola, that causes snake fungal disease, has
been documented in this species (0.6% of pathogen prevalence; Blanvillain et al. 2022). Finally,
populations of N. astreptophora are genetically more diverse than those of the more northern
species Grass Snakes (Kindler et al. 2018a, b), something well known for Western Palearctic
species whose range was affected by the Quaternary glaciations; this represents an added value
for the conservation of N. astreptophora.

Dice Snake
Natrix tessellata (Laurenti, 1768)

Konrad Mebert

Two large and biologically comprehensive accounts have been composed on much of what
is known about the Dice Snake Natrix tessellata today: Gruschwitz et al. 1999 (65 pp.) and
Mebert 2011a (456 pp.). This information is summarized and complemented with subsequent
publication since then.
Distribution and Habitat. Type locality “Japidia, vulgo Cars” = Istrian Peninsula, Cro-
atia. The Dice Snake is one of the few snake species occurring naturally on three continents:
Europe, Asia, and Africa. In the latter case, it ranges at least for a few hundred kilometers into
Egypt (Baha El Din 2011), but possi-
bly being in the process of expand-
ing farther south along the Nile Riv-
er towards Sudan. Countries of well
known occurrence in approximative
west to east and north to south order
are: Germany, southern Switzerland
(native) and western to central-north-
ern Switzerland (introduced), eastern
and southern Austria, Italy, Czech Re-
public, Slovakia, Hungary, Slovenia,
Croatia (including some Adriatic is-
Figure 28.9. The distribution of Natrix tessellata. lands, e.g., Krk, Cres), Serbia, Kosovo,
Bosnia and Hercegovina, Montenegro,
North Macedonia, Albania, Moldova, Romania, Bulgaria, Greece (incl. several islands such as
Crete, Lesbos, Samos, Corfu, Kithira, Serifos, Euböa, Tinos), Russia, Ukraine, Georgia, Azer-
baijan, Armenia, Turkey, Cyprus, Syria, Jordan, Lebanon, northeastern Egypt, Israel, Iraq, Iran,
Kazakhstan, Kyrgyzstan, Uzbekistan, Turkmenistan, Tajikistan, Afghanistan, Pakistan, and in
Xinjiang Province north-western China (Mebert 2011a; Jablonski et al. 2023c, d). The single
record from southern Poland (Vlček et al. 2010) was recently confirmed with more materi-
al by Zając et al. (2019, 2022). Its limits are primarily set by unsuitable habitats, such as a)
the water-poor arid areas in north-western China, southern Iran, and Arabian countries as
far as western Egypt, and b) cold regions north of the 53° latitude and those of high eleva-
tions from the Chinese Tibetan plateau south to the Karakoram-Himalaya massif in eastern
Pakistan and India (Mebert and Masroor 2013; Jablonski et al. 2023c, d). The second most
relevant limiting factor appears to be the same or similar ecological niche occupation by a dif-

479
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ferent natricine species. For example, the Viperine Snake (Natrix maura) inhabits habitats in
south-western Europe and north-western Africa that is otherwise typically occupied by Dice
Snakes. Similarly, the Asiatic Watersnake (Fowlea piscator) appears to replace the Dice Snake
in extreme south-eastern Afghanistan (Kabul River) into southern and central Pakistan and
farther east (Mebert et al. 2013). Natrix tessellata was erroneously reported from Yemen by
a single juvenile specimen (Scortecci 1932), stored today at that Museo Civico di Storia Na-
turale, Milano, Italy, and was subsequently doubted by Leviton (1977). The author herein has
personally inspected and compared that specimen in his wider graduate research of > 500
N. tessellata (Mebert 1993, 2011b). Indeed, color pattern and scale characters did not differ
from Italian Dice Snakes, suggesting the Yemeni specimen to represent a possible mis-labeled
or mis-placed Italian N. tessellata [Link] juvenile Dice Snake was initially deposited
at a non-secured military hospital in Florence, Italy (Escuola di Sanita Militare di Firenze;
Scortecci 1932), where local Dice Snakes may easily have brought in and addeed to an enthu-
siastically composed reptile collection. On the other hand, Scortecci has been known to have
mislabeled preserved specimens before (W. Böhme, K. Mebert, unpubl. information). Anoth-
er source of erroneous reports of Dice Snakes in Yemen relates to the potential confusion with
some phenotypically very similar and piscivorous specimens of the Wadi Racer Platyceps
rhodorachis occurring in Yemen; see dark-blotched on grey and fish-devouring Wadi Racer
on p. 48 in Egan 2007). Other records from France, Belgium and other countries originate
from introduced or escaped captive specimens, without the establishment of a population,
and will not be further discussed. There is a potential that specimens from the introduced
population at the northern shore of Lake Geneva, Switzerland, could cross the lake to reach
the southern French coast (see also general accounts and references in Gruschwitz et al. 1999;
Mebert 2011a).
Inventories. Based on a renewed monitoring to assess the conservation status of Dice
Snakes in Ticino, southern Switzerland, Mebert et al. (2011b) calculated a detection prob-
ability of 61% and a site occupancy of 78% of historically known locations. On a nation-
al level, Smole–Wiener (2011) and Kammel and Mebert (2011) summarized basic data of
Natrix tessellata from southern Austria, with latter also reporting on the recolonization of
river segments by the Dice Snake after a larger river rehabilitation program and looking at
the effects of hydroelectric power plants. Other studies in Mebert (2011a) provided first or
updated national accounts on variably combined aspects including distribution, conserva-
tion status, habitat and other ecological aspects of N. tessellata, e.g., for Slovenia (Žagar et
al. 2011), Croatia (Jelić and Lelo 2011), Romania (Strugariu et al. 2011), Bulgaria, including
also morphology (Naumov et al. 2011), Ukraine (Kotenko et al. 2011), Georgia (Frotzler et al.
2011), Egypt (Baha el Din 2011), Syria (Shehab et al. 2011), Jordan (Amr et al. 2011), Iran, in-
cluding morphological variation (Rajabizadeh et al. 2011), and China (Liu et al. 2011), where-
as separate restricted surveys were conducted for Kazakhstan (Dubrovsky 1967), Kyrgyzstan
(Yakovleva 1964), Tajikistan (Said-Aliev 1979), Uzbekistan (Bogdanov 1960; Tashmuradov
1975; Vashetko 1981), Afghanistan and Pakistan (Mebert and Masroor 2013; Mebert et al.
2013; Wagner et al. 2016; Jablonski et al. 2023c). Regionally within a country, Dínçaslan et
al. (2011) showed variation of blood serum proteins and morphology in lake populations
from southwestern Türkiye, and Ahmadzadeh et al. (2011) recorded basic morphology and
reproductive development in Dice Snake populations from the south-eastern coastal region
of the Caspian Sea in Iran. Tuniyev et al. (2011) updated distribution data, accounts on color
pattern variation, activity, and conservation of Dice Snakes along the Caucasus isthmus, ex-
cept for most of Georgia.
Several new peripheral populations and surprising rediscoveries of Dice Snakes have

480
 28. The Palearctic Watersnakes

been reported relatively recently, such as the 500 km range extension across the Tarim desert
in China (Liu et al. 2011), first populations from the Baltic Sea Drainage Basin (Vlček et al.
2011), rediscoveries on the island of Cyprus (Göçmen and Mebert 2011), recolonization of
the capital of Romania, Bucharest, (Strugariu et al. 2011), and the Czech Republic, Prague
(Velenský et al. 2011), rediscovered or first confirmation on Croatian islands in the Adriatic
Sea, Krk (Vlček et al. 2015) and Cres (Burić and Baskiera 2014). The recent discoveries of
Dice Snakes in a valley of the Karakoram-Himalaya mountain range in Pakistan were as-
sessed in terms of glacial refugial expansion routes from Uzbekistan and Afghanistan and the
required transgression over a high pass at 3,700 m elevation during the warming period in the
Holocene Climatic Optimum (Mebert et al. 2013), recently supported based on genetic and
environmental data (Jablonski et al. 2023d).
Dice Snakes generally inhabit a large array of wetlands, including modest saline seas
and estuaries, freshwater lakes, ponds, rivers, streams and mountain brooks. Their aquatic
habitats are usually rich with fish, the principal prey, but they can switch to frogs where nec-
essary. Originating from wide steppes in Central Asia, they occupy relatively open and even
arid areas as long as wetlands provide food and rocky shores or dense reed mats for shelter.
These habitat characteristics are distinct from its largely sympatric Grass Snake N. natrix and
N. helvetica, which tends to occupy shadier, thus cooler and more humid, aquatic habitats and
is primarily an amphibian predator.
Identification. Summarized from Gruschwitz et al. (1999), a medium-sized snake with
a total length up to around 120 cm, claimed up to 130 cm for one insular population (Black
Sea); a narrow, pointed head; heavily keeled dorsal scales, usually 4 to 5 alternately arranged
rows of dorsal spots/blotches, some blotches of the two middle rows maybe medially fused,
usually distinct from sympatric/parapatric Grass Snakes (including N. helvetica, N. natrix)
and Viperine Snake (N. maura) by a combination of 2 to 3 preocular scales, 3 to 4 postocular
scales, 8 supralabial scales, of which the fourth or the fourth and fifth supralabial scale touch
the lower margin of the eye; further distinct from the other Natrix species by 19 scale rows
around mid-body, higher ventral scale counts, strongly keeled scales on the upper side of the
tail, and the lack of semi-lunar anterior bright/posterior black blotches on the neck as well as
the lack of lateral ocelli (ring-shaped blotches).
There is an extensive scale variation across the Dice Snake´s vast distribution, which is
underlined by a constant sexual dimorphism (Mebert 2011b, c; Werner and Shapira 2011)
with females being considerably larger, exhibiting region-specific lower number of ventral
scales (on average 5 less) and subcaudal scales (on average 10 less) than males, but females
also exhibit a more posterior reduction of dorsal scale-rows on the trunk, a less posterior
position on the tail, a greater relative tail length, body width and posterior head length, and
a narrower head, whereas head shape and size also differ between separate Greek island pop-
ulations in accordance to their principal prey type, anuran vs. fish (Brecko et al. 2011). Prin-
cipal scale characters (number of ventral and subcaudal scales) vary clinally, mostly with
increasing values from west (Italy) to east (Turkey) and from south (Egypt to Iraq) towards
north, with the highest scale number in the lowland region of Ukraine and Russia between
the northern shore of the Black Sea and the north-western coast of the Caspian Sea. In this
region, Natrix tessellata exhibits 18 ventrals more than conspecifics in Western Europe. On a
microgeographic scale, more than 50 morphological characters were sufficiently powerful to
detect significant geographic variation across relatively short distances of 40 to 100 km and
helped to identify the origin of introduced Dice Snakes without genetic data (Mebert 2011d),
albeit such was later confirmed by molecular studies (Gautschi et al. 2002).
The ground color of Natrix tessellata varies from olive to beige, grey, and brown, with 4–5

481
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rows of dorsal blotches common across its large distribution. Small spotted specimens are
common in populations from the southeastern Mediterranean between Lebanon and Egypt.
Melanistic and spotless (concolor) morphs can be locally absent to be abundant, in particular
in a region between the southern Balkan and the Caspian Sea, but occur locally over most of
its huge range, including recently discovered ones in the Czech Republic and Italy (Mebert
2011b), Slovakia (Jablonski and Kautman 2017), Montenegro (Gvozdenović and Schweiger
2014) and Slovenia (Cafuta 2011). In addition, a rare striped morph or at least one with longi-
tudinally stretched blotches is known from the Caspian Sea (Tuniyev et al. 2011), Syria, Leb-
anon, and southern Iran (Mertens 1969, E. Kramer, unpubl. data). It has occurred constantly
at Lake Iliki, Greece (Mebert 2011b).
Furthermore, Mebert (2011d) and Gautschi et al. (2002) elaborated on the high frequen-
cy of scale abnormalities in introduced Dice Snakes in Switzerland and related deformed ven-
tral scales to fused vertebrae. This and the occurrence of exceptionally short Dice Snakes with
up to 20% fewer body segments viewed in the context of inbreeding in introduced popula-
tions, following a severe bottleneck of the founding population of ca. 20 adult snakes. Brecko
et al. (2011) found a prey-dependent head shape in Dice Snakes, with those consuming frogs
exhibiting significantly narrower and more streamlined frontal heads than in populations
that consumed fish, suggesting a phenotypically plastic response to the local abundance of
prey types. Herrel et al. (2008) found that among the semi-aquatic snakes, frontal striker spe-
cies, such as N. tessellata, have generally smaller heads but also reduced frontal and projected
frontal surface areas compared to lateral strikers like Nerodia fasciata.
Histological investigations about the Duvernoy’s’ gland in the Dice Snake, a secretion
apparatus in Natricines homolog to the venom glands of viperid and elapid snakes, were con-
ducted by Gygax (1968, 1971), Ovadia (1984), Akat et al. (2011) the skin sensory organs by
Walztöhny and Ziswiler (1979), embryonic development by Korneva (1969), and karyology
in Manuilova and Pisatets (2008). Other studies on skull morphology are analysed in And-
jelković et al. (2016) and referenced in Gruschwitz et al. (1999), with the latter also containing
small chapters with references on the postcranial skeleton, hemipenis, and blood cells.
Phylogeny and phylogeography. Analyses by mitochondrial cytochrome b sequences
and nuclear ISSR-PCR genomic fingerprinting identified nine mitochondrial DNA haplotype
clades associated with animals from Europe except for southern Greece, Jordan/Egypt, Tur-
key, Caucasus, Kazakhstan, Uzbekistan, Crete, and Iran, respectively (Guicking et al. 2009;
Guicking and Joger 2011, Kyriazi et al. 2013). Genetic data indicate that N. tessellata originat-
ed in southwest Asia during the Miocene (first split off by the Iran clade at 9.6 Mya), followed
by basal radiation around the Miocene-Pliocene boundary with further differentiation of
the European lineages during the Pleistocene (Jablonski et al. 2023d). Excluding rather dis-
tinct N. tessellata from southwestern Balkan (Greece, Albania), geographic mitochondrial
differentiation among European populations is low, reflecting a severe population bottleneck
during the late Pleistocene followed by a rapid expansion (Guicking and Joger 2011, Gautschi
et al. 2002). In Asia, the Anatolian and Central Asia clades of Dice Snakes show deep diver-
gence from European populations. The two Asian clades began divergence at 3.7 MYA in the
Pliocene, with their three, respectively four, mitochondrial lineages radiating ca. 1 M years
later (Jablonski et al. 2023d). Their distributions are affiliated with specific river systems and
are further restricted by high mountains and extreme deserts.
The oldest fossil records of N. tessellata originate from the middle Pliocene and suggest
a continuous presence across the East European Plains since that period, with range contrac-
tion and expansion following the frequent climatic and topographic fluctuations (Ratnikov
and Mebert 2011). During cooler periods and after range contraction, some populations sur-

482
 28. The Palearctic Watersnakes

vived isolated in thermally warmer pockets, as for example the four remaining populations
in Germany, but also similar accounts for isolated northern populations in the Ukraine and
Russia (Kotenko et al. 2011; Litvinov et al. 2011).
Population Aspects. In ideal habitats, such as open rocky shores along fish rich water
bodies, either natural or man-made by riparian reconstruction, Dice Snakes are known to
reach high densities and large populations, even when introduced (Mebert 2011a, d; Velen-
sky et al. 2011; Metzger et al. 2011; Sterijovski et al. 2011; Tuniyev et al. 2011). Large scale
research by international teams found huge numbers of Dice Snakes at such ideal habitats.
For example, a Swiss team marked/recaptured Dice Snakes at an introduced site in Central
Switzerland and estimated a population size of > 2000 individuals along a 3 km shore-
line habitat, corresponding to about 1 snake/m (Bendel 1997). In Romania, Carlsson et al.
(2011), did a mark/recapture study along the shore of a brackish water lake next to excavat-
ed Hellenistic-Roman ruins at Histria. They individually marked > 2300 Dice Snakes along
a 200 m stretch in the first year and > 1500 snakes in the two subsequent years, estimating
a population size of > 10’000 individuals. As the ruins are the only rocky microstructure
for hibernation for the greater surrounding, likely, in the months close to hibernation, in-
dividuals from nearby water bodies within a few kilometers were also present. Sterijovski
et al. (2014) marked > 6000 Dice Snakes on Golem Grad, North Macedonia, a small island
of 18 ha, estimated to be inhabited by at least 10’000 N. tessellata, respectively > 500 snakes
per ha. Pauwels et al. (2020) reported a total of nearly 2000 Dice Snakes that were counted
basking and foraging for fish along both sides of a one-kilometer-long rocky pier penetrat-
ing the Caspian Sea, amounting to a density of 1 Dice Snake/m. Mebert, in Burton (2021),
suggested that Dice Snakes were the ideal and likely candidate for Hannibal’s “Bithynian
snake bombs”, by catapulting clay pots full of the “viper-like” Dices Snakes thrown a the
enemies’ ship during a naval battle at 184 BCE.
Natrix tessellata is also well known for entering saline water. For example, Dice Snakes
have been seen in seawater off the coasts of Crete Island (Van der Meijden 2006), the Italian
sector of the northern Adriatic Sea (between Trieste and Venice) and observed by fisher-
men a few km offshore (Mebert 2011e), and all along segments of rocky shores around the
Black Sea and offshore islands (e.g., Serpilor Island at 30 km distance from the mainland).
In the Azerbaijan sector of the Caspian Sea, Natrix tessellata has been observed on aban-
doned wooden oil drilling platforms up to 8–9 km from the mainland (Tuniyev et al. 2011).
In addition, Dice Snakes have colonized recently built artificial islands in the Kazakh sector
of the Caspian Sea (Pauwels et al. 2020). The minimal distance between the mainland and
these new islands where Dice Snakes were observed is about 50 km. Whether these snakes
were accidentally imported with barges importing limestone boulders and soil from the
mainland to construct the artificial islands or they were swimming/drifting the large dis-
tance remains unclear. Numerous photographic and anecdotal evidence exist from around
the Caspian Sea with Dice Snake seen several kilometers offshore, approaching boats, and
also climbing into boats and travel like that one Dice Snake that was riding 10 days on an
environmental survey ship from the mainland until reaching one of the artificial islands,
where it left the ship voluntarily (Pauwels et al. 2020). A study on N. tessellata from a marine
coastal (Black Sea) environment showed that these snakes could display hypernatremia
(e.g., increased plasma sodium) without any apparent ill effect on physiological and behav-
ioral traits and thus are relatively tolerant to salinity. This strongly suggests specific physio-
logical adaptations to withstand hyperosmolality (Brischoux and Kornilev 2014; Brischoux
et al. 2017). However, their salt tolerance appears to be particularly suitable to sustain large
populations in the Black Sea and Caspian Sea, which are 2–3 times less saline than the

483
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Mediterranean.
Activity and Thermal Biology. A radiotelemetric study at three different sites in Ticino,
southern Switzerland, revealed an annual linear “home range” of approx. 500 m of terrestrial
habitat parallel to the shoreline, consisting primarily of man-made rock walls along a river or
lake shore (Conelli et al. 2009, 2011). Within this home range, the radio-tracked Dice Snakes
were found only in the immediate surroundings of the open water surface, with 97% of the
locations occurring less than 20 m from the water. However, movements showed site-depen-
dent variations. At two sites, Dice Snakes foraged, mated, thermoregulated, and hibernated
in the same area. In contrast, at the third site, the snakes revealed substantial seasonal move-
ments for different summer and winter habitats). In spring, they migrated to their summer
habitat, a backwater system next to a lake, for all activities (Conelli et al. 2009, 2011). In the
fall, the snake returned to the hibernation site, a dam-like rock construction to support an
elevated road next to a floodplain. All animals studied remained within the riparian zone
during hibernation (maximum distance from water ≤ 10 m), and selected shelters of at least
1.5–4 m above the mean water level, thus, reducing the risk of drowning during late winter
high water while still in hibernation. Microclimatic conditions of hibernacula allowed ani-
mals to maintain a body temperature between approx. 4–10° C, even when external tempera-
tures decreased below 0 °C in January and February.
A radiotelemetric study was conducted at Prague Zoo, Czech Republic, where wild Dice
Snakes colonized a relatively new, rocky embankment, completed in 1989, between Vltava
River and the zoo (Velenský et al. 2011). This embankment yielded a higher slope, more
crevices, and less shade than the previous embankment. The first Dice Snakes showed up in
the mid-1990s. The population increased over the years to reach a density of approximately
180 individuals per hectare, resulting in a population size between 700–800 individuals in
2008. During the study, Dice Snakes hibernated up to 238 days to mid-April. In early spring,
the snakes migrated several hundred meters through the zoo’s territory in a few days to reach
a summer habitat along the river embankment. There, the snakes mated, foraged, thermo-
regulated, and oviposited, with some individuals regularly entering the zoo to use its gullies
and fishponds to hunt. In the fall, some Dice Snakes crawled back through the zoo to return
to their hibernacula on a slope at the other side of the zoo. In contrast, some remaining in-
dividuals hibernated directly in the river’s embankment. Oviposition happened within the
first two weeks of July, after which female snakes were left with as little as 40 days to acquire
enough food and energy for the following hibernation. Basking was frequently observed on
days with a thin cloud cover but sufficient solar radiation to warm the upper surface. On hot
summer days, Dice Snakes usually moved under rocks and practiced thigmothermy, i.e. gain-
ing sufficient body temperature by acquiring heat from the warm surface.
At the most northern population of N. tessellata near Samara along the Volga River,
Russia, Litvinov et al. (2011) investigated thermal behavior of that all-melanistic population.
They survive around southern exposed rocky riparian bluffs. Snakes were active at surface
air temperatures ≥ 14.0° C and substrate temperature ≥ 13.2° C. Females selected more open
areas than males. When internal body temperature reached 28° C, Dice Snakes moved to
cooler sites.
Reproduction. Males are generally the first gender to emerge from hibernation, sup-
posedly to activate spermatogenesis (e.g., Kärvemo et al. 2011). Test activity in males lasted
the entire season, whereas vitellogenesis in females is seasonal. Mating usually follows a few
weeks after hibernation, primarily in April–June, less frequent in March or July (Gruschwitz
et al. 1999). Mass mating and mating ball formations are common in dense populations (Me-
bert and Ott 2011), whereas late summer/fall mating rarely occurs (Gruschwitz et al. 1999;

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Ajtić et al. 2013). Females begin with their yolk deposition early in May, with large follicles
reaching a maximum level in summer, late June–early July, and oviposition during July and
early August, e.g., in Switzerland (Bendel 1997, 2001), northern Iran (Ahmadzadeh et al.
2011), and North Macedonia Ajtić et al. (2013). Latter authors also found that 54% (336
of 621 individuals) of adult females examined were gravid. The mean clutch size was nine
(range 1–20) among 699 females, where the follicles could be counted by palpation. In cen-
tral Italy, the mean clutch size was approximately 15 (Luiselli and Rugiero 2005), whereas a
maximum number of 37 eggs in captivity was given by Dimitropoulos (1989).
Foraging and Diet. Natrix tessellata primarily consumes fish in its European range
but often feeds on frogs in western Asia and occasionally adds a newt or salamander, and
even rarer a young rodent, duck, lizard, or invertebrate (Gruschwitz et al. 1999; Weiperth
et al. 2014, and references therein). Non-fish species in the diet were especially pronounced
in deserts, high mountains, and dry Asian and Mediterranean areas. Generally, any fish is
preyed that an individual Dice Snake can overcome. They forage during day, but also during
warm nights (at ca. 20°C or more), in fresh or saline water, e.g., with large populations in the
Black Sea and Caspian Sea, but have also been observed in the Adriatic and Mediterranean
Seas. Open water fish are ambushed by anchoring tail on rock or submerged water plants,
whereas ground fish are actively searched in crevices of subterranean rocks at least as deep as
10 m. In populations where N. tessellata show a white tip on an otherwise black tongue, the
snakes have been observed to slowly tongue-flick when a fish has approached near striking
distance (Mebert 2007).
Radiotelemetric studies revealed that N. tessellata, inhabiting a rocky wall and bank
along a river in Germany, descended only every 4–5 days from their terrestrial shelter to for-
age in the water up to 15 m away and 100 m along the shore. Still, they else remained on land
to rest and thermoregulate (Neumann and Mebert 2011). They were mainly active at air tem-
peratures between 20–26 °C on a partially cloudy sky. Extensive accounts were summarized
for a lake population in North Macedonia by Ajtić et al. (2013): all Dice Snakes observed
fishing were in ambush position near the shore (< 5 m offshore), sometimes in high density;
snakes were rarely observed swimming more than 100 m from the shore (n < 5); the propor-
tion of fed snakes varied across seasons; these snakes rarely foraged early in the active season
in spring (late April to early May); only 15.4% of the snakes were observed with prey in their
stomach with this proportion increased markedly (44.1%) in summer (June–August), and
the snakes ceased to fish in autumn (September–October).
A study on the stomach content of Turkish Dice Snakes showed the usual dominance
of fish (72.4% of items), followed by amphibians (14.5%), but also a few unusual stomach
items, such as insects (7.9%), gastropods (2.6%), one reptile, and one mammal (Göçmen et
al. 2011b). While the invertebrates may have been ingested indirectly through primary prey,
the reptile, a large Balkan Green Lizard (Lacerta trilineata, approx. TL 200 mm), and an adult
Forest Dormouse (Dryomys nitedula, approx. TL 320 mm), were found in large female N. tes-
sellata. The gastropod Lymnaea stagnatilis and even a juvenile Northern Viper (Vipera berus)
were found in the stomach of Dice Snakes from the Volga Basin, Russia (Bakiev et al. 2011).
Long-term studies in central Italy investigated the biological aspects of N. tessellata at
three sites (Capula et al. 2011). Among others, they found an ontogenetic shift in the lower
size limit of prey in all three study areas. The larger snakes were selected against smaller prey
sizes, as the minimum prey size increased with snake size (Luiselli et al. 2007). This is true
especially for females, likely due to the high energy demands of females for reproduction.
Likewise, a remarkable and significant inter-sexual variation in the composition of the diet is
correlated with the considerable body size differences between the two sexes (females being

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more significant than males). For subsequent digestion, gravid females selected significantly
higher substrate temperatures than non-gravid ones.
Mebert and Pölzer (2011) presented two fatal accidents of Dice Snakes by which the
hunted groundfish snapped back and locked onto the snakes’ upper jaw with subsequent
drowning of both prey and predator. Velikov (2011) illustratively presented material of Dice
Snakes feeding on introduced spiny fish, whereas Mebert (2011f) revealed the unusual stom-
ach contents of three Dice Snakes that consumed rocks or peach stones.
Predators and Defense. Various predators feed regularly on Dice Snakes: Smaller mam-
mals such as the Brown Rat (Rattus norvegicus), introduced Muskrat (Ondatra zibethicus),
Short-tailed Weasel (Mustela erminea) Least Weasel (Mustela nivalis), reptiles such as the
Large Whip Snake (Dolichophis jugularis) in Israel, and numerous bird species (especially
Ardeidae herons, Black-headed Gull (Chroicocephalus ridibundus), Kingfishers (e.g., Halcyon
smyrnensis) are listed and referenced in Gruschwitz et al. (1999).
Predators on Dice Snake on Golem Grad Island, North Macedonia, included European
Otter (Lutra lutra), with > 50 feces usually containing large amounts of snake scales and ver-
tebrae, and different birds that preyed heavily on the snakes (e.g., Bubo bubo, Ardea cinerea,
Accipiter sp., Buteo sp., Larus sp.). Nose-horned Viper (Vipera ammodytes) also commonly
preyed on Dice Snakes (Ajtić et al. 2013). The Balkan Whip Snake (Hierophis gemonensis)
and the Gull (Larus cachinnans) have been documented as predators in Croatia (Jelic and
Laus 2011a, b), and the Raccoon Dog (Nyctereutes procyonoides) in Musilová and Zavadil
(2011). Finally, ecto- and endoparasites have been frequently analysed and summarized in
Gruschwitz et al. (1999), Mihalca (2011) and Bakiev et al. (2011), but also in newer studies,
for example Kirillov and Kirillova (2019).
Active defensive behavior is manyfold and individually variable. It can include vomiting
of freshly captured food, excretion of faeces with an intense fish odor, secretion of a yellow
liquid from glandular fields next to the cloacal opening that smells of butyric acid, balling
– primarily associated with hiding the head in or under the coils of the body, violent tail
wagging, and body undulating when lifted, but also a viper-style defense, such as flattening
of the entire body, audible hissing, and fake bites. Such supposedly Batesian mimicry with
head flattening and fake striking in Dice Snakes has been found in populations in proximity
or sympatry with local vipers instead of viper-free localities (Bijelica et al. 2023). However,
in 20+ years of experience handling 1000s of Dice Snakes, the author has never observed a
true bite (K. Mebert, unpubl. Data). A death reflex, like that of the Grass Snake (Natrix natrix
group) or Hognose Snake (Heterodon platirhinos) is also part of individual defense repertoire
and a combination with reflex bleeding in the mouth area, albeit rarely.
Sympatric Natrix species. Grass Snake spp., N. natrix sensu lato, prefer small, tempo-
rary, and permanent water bodies, such as ponds in forested situations, wet meadows, and
marshes, with large breeding concentrations of their preferred prey, frogs, and toads. In con-
trast, N. tessellata is more aquatic, frequent comparatively more open water courses in rockier
and drier areas and prefers more fish. The habitat partitioning was also confirmed in sym-
patry in southern Croatia (Janev Hutinec and Mebert 2011) where the diet of Natrix natrix
consisted mainly of amphibians with the occasional open water fish, whereas N. tessellata fed
exclusively on fish, and dove deeper to hunt for benthic fish. An increased underwater vision
ability of N. tessellata compared to N. natrix has been technically investigated and confirmed
by Schaeffel and Mathis (1991). Both can be common and coexist in large reed grass habi-
tats in the lowland of Eastern Europe and semi-arid landscapes, yet N. tessellata is generally
viewed to be more aquatic (Gruschwitz et al. 1999).
Few studies compared N. tessellata with the ecologically similar Viperine Snake N. mau-

486
 28. The Palearctic Watersnakes

Figure 28.10: Natural color pattern variation of Dice Snakes (Natrix tessellata) from: A), B), C) tri-
national Lake Prespa, North Macedonia, Albania, and Greece. Such co-occurring variations are
common in some dense populations, including Lake Prespa, Lake Beysehir in Turkey, or Caspian
Sea; D) partially striped specimens, here deposited at the Museum of Natural History Geneva, Swit-
zerland, were known from Lake Yliki, Greece, and occasionally other sites from its more southeast-
ern distribution. Photo credit: A)-D) Konrad Mebert.

many has become extinct but was successfully repopulated with specimens from the nearest
populations (same river system) from the Czech Republic (Obst and Strasser 2011, Strasser
and Peters 2014, Strasser 2020). In fact, bank and shoreline reinforcements made of big stone
blocks (rip-rap) with an abundance of interstices and herbaceous vegetation or little shrub
cover appear to be particularly attractive to Dice Snakes. New rip-raps along the river be-
tween a retention basin in the forest and the watercourse farther down the village have been
colonized within less than three years after their construction (Conelli et al. 2009, 2011).
Dry stone and rip-rap walls offer many places for safe shelters and thermoregulation and are
potential sites for oviposition.

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Aquatic Snakes

ra. They are largely parapatric and occupy the same habitat, N. tessellata in southeastern
Europe and N. maura in southwestern Europe. Scali (2011) compared temperature and other
ecological variables between these two species at one of their few sites of natural sympatry
in northern Italy. Dice snakes were observed in comparatively deeper streams, being more
piscivorous and less nocturnal, possibly benefitting from such habitats along the Po River, the
principal large river system across northern Italy. These two species were also compared at a
site in Lake Geneva, Switzerland, where N. maura is native, and N. tessellata was introduced
many decades ago. Compared with the Viperine Snake, the Dice Snake occupied shore zones
that were relatively more open and inhabited more often steep slopes (Mazza et al. 2011).
Metzger et al. (2011) found a large overlap in the trophic niche between both species at the
same site, regarding seasonal preferences and prey types. A higher fecundity and a larger
body size render the alien N. tessellata probably a more competitive capacity and possibly a
key role in the population decline of the native N. maura over the last decades. Finally, much
like in other natricines, interspecific hybrids between N. natrix and N. tessellata have been
produced in captivity up to the F2 generation and are rarely observed in nature, whereas
a hybrid between N. tessellata and N. maura has been reported only once between captive
specimens (Mebert 2010; Mebert et al. 2011a), but was recently shown by molecular analysis
(Schöneberg et al. 2023).
Conservation. Major threats through habitat degradation, including water contamina-
tion, are ubiquitous for agricultural, housing, industrial, and traffic purposes. For example,
a high number of road-killed N. tessellata was recorded during the summer along a stretch
of 2.1 km on the north shore of Prokopos Lagoon near Strofylia in southern Greece (Ioan-
nidis and Mebert 2011). Extrapolated to the entire season, they estimated that> 1000 Dice
Snakes are killed on that road segment yearly. Another threat resulted in high mortality of
Dice Snakes through illegal fishing around Golem Grad Island, a protected national park in
North Macedonia. A total of 44 Dice Snakes drowned in a single small fishing net, set for
3–4 hours in the water after they became entangled themselves chasing fish embedded in
the mesh. Thousands are killed yearly in protected water around the island (Sterijovski et al.
2011, 2014).
In Central Europe, Lenz and Schmidt (2011) summarized the results of extensive mea-
surements to support the three remaining populations of Dice Snakes in western Germany.
On the river Mosel, the core habitat was improved by reconstructing the lentic water courses
with protective boulders and rock walls, providing piles of decomposing plants, branches,
and organic waste material for oviposition. They also tested different guidance and deflec-
tion systems to avoid road-kills. Open gravel areas were recreated on the river Lahn, and two
new founder-populations were started with captive-bred juveniles Dice Snakes (Trobisch and
Gläßer-Trobisch 2011). A fourth isolated population at the river Elbe in northeastern Ger

On the next [Link] Eastern Ribbon Snake, Thamnophis

saurita from central Florida. Photography JCM

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Aquatic Snakes

489
Aquatic Snakes

29. The Thamnophiine Clade

The New World natricids snakes are monophyletic, descended from an ancestor that
immigrated from the Old World. They form the clade Thamnophiine, with some of the plan-
et’s most studied serpents. Zaher et al. (2019) found the genus Storeria, the sister to all the
other New World natricids clade members. Storeria comprises five ground-dwelling cryp-
tozoic snakes that feed primarily on worms and mollusks. While they tend to live on damp
soils, often near bodies of water, they are not semi-aquatic. All species are viviparous.
The most basal Thamnophiine clade recovered by Zaher et al. (2019) included the Li-
odytes, Clonophis, Haldea, and Virginia genera. These are a mixture of fossorial, terrestrial,
and semi-aquatic species. Clonophis exploits an unusual aquatic habitat – crayfish burrows;
both Haldea and Virginia are small fossorial species. Liodytes contain some of the most
aquatic North American snakes.
The ancestor of North American natricids immigrated from Eurasia sometime before
the early Miocene (~19–18 Ma). We know this because the earliest known Western Hemi-
sphere natricid fossils date to the early Miocene, and we know they shared an ancestor be-
cause of shared anatomy and DNA.
Alfaro and Arnold compared three genes from many North American natricid snakes.
They found that they shared a distant ancestor and formed three distinct clades: a water
snake clade, a garter snake clade, and a novel semi-fossorial clade. Living around the subur-
ban quarry were members of each of the three clades. Alfaro and Arnold’s work suggested
that the most basal Western Hemisphere natricids is the fossorial Kirtland’s Snake (Clonophis
kirtlandi). While Kirtland’s Snake is unknown from the area around the quarry, there is a
population within 10 miles. Both the Kirtland’s Snake [Figure 11–6b] and its sister, Dekay’s
Snake, occur in wet prairie or grassland habitats. Kirtland’s Snake is fascinating because it
is primarily fossorial and appears to use the chimney building crayfish burrows as a hiber-
nation site (suggesting it hibernates underwater or at least in the water). Clonophis has a
fractured distribution with isolated populations widely scattered over Illinois, Indiana, Ohio,
and bordering states.
North American natricids are perhaps the best-known serpents. Garter snakes, water
snakes, and brown snakes are often abundant in urban areas with remnant habitats. Natricids
have continually adapted to aquatic habitats and reverted to a life on land, only to return to
the water. At least a few garter snakes are entirely terrestrial, but many forage along streams
or ponds, while others are aquatic.
Jean-Claude Rage proposed that natricids invaded North America from Europe via a
land bridge. Since the oldest natricid fossils are from the early Miocene, the land-bridge mi-
gration and colonization must have occurred before this date. An early Eocene land bridge
connecting Europe-Greenland-North America with a subtropical climate may explain the
expansion of these snakes into the Western Hemisphere. But how probable is it that snakes
used a high latitude land-bridge to disperse from Europe through Greenland and into conti-
nental North America in a mild or cold climate?
A snake den in an abandoned railroad yard in Portage County, Wisconsin, provided a
valuable clue. A friend had discovered the snake den near his summer house and took sev-
eral of us for a tour. The circular cement foundation was mostly covered with rusted metal;
it had been used as a giant turntable to switch railroad cars from one track to another. The
cement cistern was about 2.4 m deep and partially filled with water. Shade from the metal
covering had insulated the layer of ice and kept it from melting, but the ice cover was com-
pletely solid and transparent. Eastern Garter Snakes (Thamnophis sirtalis) and Western Fox

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 29. The Thamnophiine Clade

Snakes (Pantherophis vulpinus) were submerged beneath the ice with their posterior bodies
wrapped around debris. The snakes had been in the water since the previous fall; this was
early May. With a solid ice cover in place since January or earlier, the snakes had not filled
their lungs with air for at least five months.
Field observations and experiments by Costanzo (1986, 1989a, 1989b, 1989c) compared
submerged and non-submerged garter snakes from this Wisconsin den and found submerged
snakes in water at 5º C. reduced their oxygen consumption by 54% and their heart rate by
77% over experimental snakes hibernating in the air under the same conditions. Snakes hi-
bernating underwater lived in acidic (5.5–6.5 pH), hypoxic (dissolved oxygen was 2.9 ppm)
groundwater. And yet, the snakes were carrying out aerobic respiration, using oxygen that
diffused through their skin from the water for five months, a remarkable ability, but one that
could have been anticipated. Carpenter (1952) studied garter snakes in Michigan in the early
1950s and observed five Eastern Garter Snakes hibernating in the burrow of a crayfish. He
wrote,

“…many snakes hibernate completely submerged in water in underground

tunnels. It seems that this should be possible, provided the water is very cold.”

Costanzo’s experiments and the significance of snakes being able to spend months sub-
merged in cold water have been long overlooked. The ability of natricids to survive sub-
merged under the ice for months is valuable for surviving long Midwestern winters and could
explain how snakes dispersed from Eurasia to North America by colonizing high-latitude
land bridges. Even if the land bridges had a seasonally cold climate, these snakes could have
made the intercontinental trip.
Ulrich (1989) noted that freshwater snakes usually hibernate on land. Still, they appear
capable of aquatic hibernation, and many do not do so because of the risk of death from anox-
ia. Some terrestrial snakes are known to hibernate underwater and can do so in the laboratory
for months. This behavior is considered opportunistic in the field, as there is no evidence to
suggest that snakes can tolerate extended anoxia.
More than 100 species of natricids use aquatic environments to varying degrees; most
feed on aquatic animals. Bilcke et al. (2007) compared feeding behavior in natricids believed
to be dietary generalists and specialists. The generalists were thought to forage for food by
open-mouth searching, holding their mouth open, and sweeping it through the water until
they made contact with prey. These species strike slowly to the side, perpendicular to the
body’s long axis, and do not seem to depend upon vision. Specialists were thought to hunt
from ambush, be visually alerted by the presence of prey, and strike forward parallel to the
long axis of their body. However, Bilcke and colleagues also looked for a correlation between
prey density and strike behavior. Two European natricids (Natrix tesselata and Natrix mau-
ra) were compared to the North American Nerodia fasciata. They found that prey-capture
strategies in natricid snakes were not correlated with diet but with prey density. For example,
snakes feeding on fish in a school or concentrated in shallow water use the open mouth tech-
nique, while snakes feeding on fish at a low-density strike from ambush.

Striped Crayfish Snake

Liodytes alleni (Garman, 1874)
Distribution and Habitat. Wetlands include marshes, slow-moving streams, and
swamps in Florida and southeastern Georgia. The type locality is in the vicinity of Jackson-

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ville, Florida.
Identification. The total length is 330–500 mm. The
stripes are indistinct and located laterally on the dorsum.
The ventral surface is yellow with dark spots. Average adults
have total lengths of 500-600 mm, and the maximum re-
ported total length is 70.5 cm (Demuth 1997). Neonates are
Figure 29.1. Distribution of Liodytes 153-291 mm in TL. The head is not much wider than the
alleni.
body. The nares are directed dorsolateral, and compared to
congeners, the eye is large with a small pupil. The scales are smooth except on the dorsal area
of the tail, where they are always keeled. Females have 110-133 ventrals (males average 2-3
fewer) and 53-61 subcaudals (males have 59-69). Dorsal scale counts are 19- 19- 19 (or 19-
19-17 in some males). A single internasal, coupled with smooth dorsal scales, distinguishes L.
alleni from other members of the genus. Other scale characteristics include two prefrontals, a
single preocular, two nasals (partially divided by the nostril), three or four postoculars, loreal
present, 1 + 2 temporals, eight (sometimes seven or nine) upper labials, 11 (sometimes nine
or ten) lower labials, and a divided cloacal scute. Females are the larger sex. It can be distin-
guished from the Glossy Crayfish Snake (Liodytes rigida) by the single row of ventral spots (L.
rigida has two rows) (Dorcas et al. 2003).
Natural History. The diet is composed of hard-shelled crayfish. However, this noctur-
nal, highly aquatic species will also feed on aquatic salamanders, small frogs, tadpoles, glass
shrimp, and the larvae of dragonflies and damselflies. Predators include crayfish, large fishes,
sirens, alligators, other snakes (Coluber constrictor, Lampropeltis getula, Agkistrodon pisciv-
orus), river otters, raccoons, Red-shouldered Hawks, Great Blue Herons, Great Egrets, and
Sandhill Cranes.
Mating probably occurs in the spring, and the young are born in the late summer or early
fall. Litter size is 4-12, with larger females producing more young.

Black Swamp Snake

Liodytes pygaea (Cope, 1871)
Distribution and Habitat. Black swamp snakes are highly aquatic and extremely secre-
tive. Individuals hide under debris or vegetation at the water’s edge, particularly on sunny
spring days. Open aquatic habitats include Carolina bays, roadside ditches, sphagnum bogs,
sawgrass prairies, and the margins of heavily vegetated ponds and lakes. Large populations
occur in wetlands with dense vegetation that dry periodically and lack fish but have abundant
amphibians. They inhabit the Gulf Coastal Plain from North Carolina southward to Florida
and west to southeast Alabama. Type locality: Volusia Co., Florida.
A polytypic species with three subspecies recognized. [S. p. cyclas: Florida; Type locality:
Indian Prairie, 2-8 miles NE of Lakeport, Glades County, Florida; S. p. paludis: North Caroli-
na, South Carolina, Georgia; Type locality: Camp Davis, Near
Identification Head distinct from the body; eyes medium, about the same size as the dis-
tance between the anterior margin of the eye to the nostril. Black dorsum with faint pale lines
running through the center of three to five lateral scale rows. Ventrals metallic red with the
dorsal color extending on the anterior edges for about one-third of the width; labials grayish
olive. A small snake, maximum length, about 425 mm. Body moderately slender with a short
tail (0.15-0.25 of total length. Rounded rostral, two internasals, two prefrontals, a frontal,
two supraoculars, and two parietals. Nasal single, nostril near upper margin with a suture
extending to the labial border. Eight upper labials and nine lower labials. Two pairs of chin

492
 29. The Thamnophiine Clade

Figure 29.2. Liodytes alleni from Florida.

Photography by Jance Carter.
shields, subequal in length. Dorsal scales are smooth, rounded, and without apical pits. Lower
scale rows are widest, decreasing width dorsally; dorsal scale rows 17 at midbody reduced to
15 posteriorly. Ventrals 112-134; cloacal plate divided; subcaudals paired, 35-45 in females;
46-56 in males (Dowling 1950).
Natural History. Black Swamp Snakes are active day and night and forage in submerged
vegetation. Diet includes small fish, tadpoles, and small frogs. This species has been found

493
Aquatic Snakes

to prey on aquatic salamanders, particularly larval and pae-

domorphic mole salamanders, and leeches in South Caroli-
na. Elsewhere they feed on aquatic salamanders in the fam-
ily Plethodontidae. They can survive long (multiple years)
droughts by aestivating within dried wetlands and survive
better than other water snakes. They recover rapidly from
droughts by feeding during pregnancy and investing ingest-
ed energy directly into offspring (income breeding). Swamp
snake populations seem to thrive post-drought years when
Figure 29.3. Distribution of Li- amphibian prey is abundant. Females give birth to up to 23
odytes pygaea. live young in late summer (Winne et al. 2005).

Figure 29.4. Liodytes pygaea from North Carolina.

Photography by JCM

Glossy Crayfish Snake

Liodytes rigida (Say, 1825)
Distribution and Habitat. Eastern Texas, southeast Oklahoma, southern Arkansas, Lou-
isiana, Mississippi, southern Alabama, southern Georgia, northern Florida, South Carolina,
eastern North Carolina, and Virginia. Type locality: the southern states. A polytypic species
with three recognized subspecies. Liodytes r. sinicola: Texas, Louisiana, Florida, Alabama;
Type locality: Beaumont, Jefferson County, Texas. Liodytes r. deltae Louisiana; Type locality:
Paradis, St. Charles Parish, Louisiana.

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 29. The Thamnophiine Clade

Identification A small- to medium snake with glossy skin, maximum size 797 mm. The
head, body, and tail dorsum are uniformly chocolate brown, with two narrow, black stripes
on either side of the midline that are difficult to detect. The ventral surface is cream with two
rows of black spots, often half-moon shaped on either side of
the midline. Cream pigmentation on the venter invades the first
dorsal scale row and fades into brown. The pale centers on the
first scale row form a lateral stripe; labial scales are white. Ven-
trals 120-136; subcaudals 51-63; dorsal scales keeled, in scale
rows 19 at midbody; upper labials seven; lower labials 10; loreal
Figure 29.5. Distribution of
Liodytes rigida. scale absent; one preocular; two postoculars; one primary tem-
poral scale, secondary temporal may be one to three.
Natural History. The diet is primarily on crayfish, and its prey-handling strategy includes
attacking, then coiling to immobilize the chelipeds of the crayfish. Followed by orienting the
abdomen so the snake’s upper jaw can bite the ventral abdomen’s soft tissues, resulting in im-
mobilization or death of the prey, followed by consumption from the caudal end (Tumlison
and Roberts 2018). The crayfish Creaserinus danielae is eaten by this snake, which uses the
burrows made by this crayfish (Loughman et al. 2020). Durso et al. (2013) found this species
to be a strict dietary specialist that relies only on crayfishes. This species was notably absent
from wetlands without fishes, partly because their probability of site occupancy in this area
is strongly related to its proximity to flood plain habitats than to the abundance of their pre-
ferred prey. Females give birth to litters of 6-14 young in late summer or fall.

Kirtland’s Snake
Clonophis kirtlandii (Kennicott, 1856)
Distribution and Habitat. The species ranges from
northeastern Missouri and southeastern Wisconsin through
Illinois, southern Michigan, most of Indiana, and Ohio to
western Pennsylvania and north-central Kentucky (Conant,
1943). Tucker et al. (1977) reported localities in Illinois,
Indiana, Kentucky, and Ohio. Wet meadows and adjacent
Figure 29.6. Distribution of Clo- woodland; several populations are known from urban areas.
nophis kirtlandii.
Conant (1943) stated that Clonophis is a typical snake of the
Prairie Peninsula, where it had survived as a relict population
of the postglacial period when prairie conditions were widespread. Type locality West North-
field, Illinois.
Identification. A small (maximum total length: 622 mm), moderately stout natricid
snake. Upper labials are 4-7 (usually six) and 6-9 (usually seven). Ventrals number 121-137;
subcaudals 44-69.
Dorsal scales overlap, are strongly keeled, and slightly notched posteriorly with two faint
apical pits (absent in many individuals, according to Conant (1961). The typical scale row
formula is 19-17 with an occasional further reduction to as low as 14. Tail length constitutes
19-24% of the total length in adult females and 23-28% in adult males. The dorsal pattern
consists of four alternating longitudinal rows of 43-65 rounded black or dark brown blotches
on a brown to gray-brown (often with a red tint) ground color. Blotches tend to be indistinct
posteriorly. Some specimens may also have dark spots adjacent to the venter that alternate
with the lower row of large blotches. The venter is pink to brick red medially, becoming yel-
low anteriorly, often with dark stippling. Laterally the ventrals are gray with large black spots

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Aquatic Snakes

Figure 29.7. Liodytes rigida, northern Florida.

Photography by Jake Scott
that form two conspicuous rows. The top and sides of the head are black, dark brown, or ol-
ive, plain or inconspicuously mottled. The labials, chin, and throat are yellow or cream except
for a dark area extending down from the side of the head onto the last upper labial. Juveniles
are often so dark that the dorsal pattern is obscured; the venter is bright and like adults.
Natural History: Diet includes earthworms, leeches, and slugs. A highly secretive, noc-
turnally active species that shelters beneath logs and surface debris, and in crayfish burrows,
by day. When threatened, it flattens its body and becomes rigid. Mates in May and gives birth
to 4-15 young in August or September. Neonates are 100-170 mm in total length.
Predators include other snakes and birds. This fossorial snake uses an unusual aquatic
microhabitat, crayfish burrows. Brust et al. (2020) found a Kirtland’s Snake outside its hiber-
naculum on 26 December in Fulton Co., Illinois, with dried mud adhered to its skin at 15°C.
Conant (1943) noted that C. kirtlandii is far less aquatic than members of the genus Nero-
dia. He said that it swims readily and usually occurs in wet meadows rather than in the water.
Its habits are much more like those of Thamnophis butleri, Thamnophis s. sirtalis, and Storeria
dekayi than they are to Nerodia. The species avoids large bodies of water and frequently oc-
curs in small ponds, streams, bogs, and woodland pools that are temporary or seasonal. In
the more open habitats, its presence is usually associated with precipitation, abundant during
and after rainy periods and absent during droughts.

Graham’s Water Snake

Regina grahamii Baird and Girard, 1853
Distribution and Habitat. The margins of mud-bottom and rocky-edged marshes, oxbow
lakes, rivers, and streams in areas with abundant crayfish are frequently inhabited. Marshes
and other wetlands in Texas, western Mississippi, Louisiana, Oklahoma, southeast Kansas,
southeast Nebraska, Iowa, Missouri, Arkansas, Illinois, and western Alabama. Type locality:
Rio Salado, Texas.
Identification. A brown dorsum with a yellow venter and three light stripes running the
length of the body, a yellow, light olive, or tan lateral stripe encompasses the lower three scale

496
 29. The Thamnophiine Clade

Figure 29.8. Clonophis kirtlandii

Photography by Stephen Barten
rows. It is bordered below by an irregular, narrow black line on the outer margins of the ven-
trals and occasionally bordered above by a fainter dark line between the third and fourth scale
rows. In addition, a dark median stripe 2–3 rows in width may be present. The pattern and
a single cloacal plate will distinguish this species from other striped and semiaquatic natric-
ids. Most specimens are 460–710 mm; the maximum known
length is 1.194 m. Dorsal scale rows are keeled in 17–20 rows
at midbody; males 162–175 ventrals; females 155–178 ven-
trals; male subcaudals 51–67; female subcaudals 60–67. Head
scales are one loreal, two preoculars; two or three postocu-
lars, upper labials seven or eight; lower labials nine to eleven
(Anton 2019).
Figure 29.9. Distribution of Re- Natural History. Of 104 digestive tracts examined by
gina grahamii
Hall (1969), 32 (30%) contained food. Of the 38 food records,
33 were crayfish remains, and five could not be unidentified.
Crayfish remains were easily identified by the presence of the digestion-resistant gastroliths.
Several stomachs contained unidentified nematodes. All the crayfish in the digestive tract had
soft exoskeletons, indicating that they may have been in the process of molting at the time of
death. Crayfish had been swallowed headfirst and abdomen- first with approximately equal
frequency.
Reproduction. Females usually are capable of reproduction before their third spring. One
422-mm female carrying eggs may have been a year older than most individuals her size but
stunted in growth. Thirteen of 21 potentially reproductive females were gravid. (All nongravid
females were in the 50- to 60-cm size range.) Litter sizes range from 10 to 27, and oviductal
eggs are 9-39 (Hall 1969).

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Aquatic Snakes

Queen Snake
Regina septemvittata (Say, 1825)
Distribution and Habitat. Branson and Baker (1974) de-

Figure 29.10. Distribution of

Regina septemvittata.

scribed the optimal habitat for

Queen Snakes as small to me-
dium, shallow streams with
moderate currents and rocky
banks and bottoms. Forested
streams, ponds, and quar-
ries from Southeast Ontario,
Canada, southward into the
eastern USA, including the
states of Arkansas, southeast
Wisconsin, northeast Illinois,
Michigan, Indiana, Ohio,
western New York, Pennsyl-
vania, Maryland, Delaware,
West Virginia, Virginia,
Kentucky, Tennessee, North
Carolina, South Carolina,
northwest Georgia, Alabama,
Mississippi, southwest Mis-
souri, northwest Florida. Type
locality: Pennsylvania.
Identification. The dorsal
color is brownish to olive, with
a yellow stripe on each side,
flanking three narrow dark
dorsal stripes. Four brown
stripes mark a yellow venter.
The labials, chin, and throat
are cream to yellow. Dorsal
scales are weakly keeled, with
two apical pits, and occur in
19 rows at midbody, reduced
Figure 29.11. Regina septemvitta-
ta. Will County, Illinois.
Photography by JCM

498
 29. The Thamnophiine Clade

to 17 rows posteriorly. The venter has 118-159 ventrals, 47-89 subcaudals, and a divided clo-
aca1 plate.
Natural History. Queen snakes are dietary specialist feeding on recently molted crayfish,
and they are highly aquatic and less secretive than other crayfish snakes. They are often ob-
served along streams and rivers basking on rocks or streamside vegetation, often alongside
Northern Watersnakes (Nerodia sipedon). Queen snakes will take refuge under rocks along
stream edges. However, unlike glossy and striped crayfish snakes, queen snakes generally
avoid hard-shelled crayfish, restricting their diet to soft, newly molted crayfish. Queen snakes
are primarily active during the day and occasionally active near aquatic habitats at dusk. Mat-
ing occurs in the spring, and females give birth to 5–23 live offspring in the late summer.
Densities of 6 to 63 R. septemvittata per kilometer of the stream were estimated in central
Kentucky by Leuenberger et al. (2019).

Figure 29.12. Regina grahamii (A) DuPage Co., IL,

Photography by Nathan Kutok

Four geographically distinct mitochondrial lineages within R. septemvittata were found,

and multilocus species delimitation analyses showed support for recognizing these lineages
as different species by Flilpen (2016). Species divergence time estimates indicated all lineage
divergences for R. septemvittata occurred relatively recently during the Pleistocene. Historical
demographic analyses indicated the Northern lineage, which occupies formerly glaciated ar-
eas, had the highest recent increase in population size, likely due to post-glacial range expan-
sion. Ecological niche modeling analyses indicated lineages occupy unique niches, but niche
conservatism has been the more prominent pattern of niche evolution.
Conservation Status: Queen snakes are common in the Piedmont and mountains. They
are not protected in most of their range but are protected throughout the state of Georgia.
Observations suggest that this species may have declined in many areas of the Piedmont, pre-
sumably due to the siltation and channelization of small streams. Maintenance of clean, rocky
streams and rivers so that an abundance of crayfish remains critical for this species.

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 Aquatic Snakes

Green Water Snake

Nerodia cyclopion (Duméril, Bibron & Duméril, 1854)
Distribution and Habitat. Uses wetlands, lakes, and streams that may be open or wooded
from eastern Texas, Louisiana, Arkansas, southeast Missouri, s Illinois, extreme western Ten-
nessee, southern and western Mississippi, southern Alabama, southeastern Georgia, extreme
northwest Florida, southern South Carolina, extreme western Kentucky. Type locality: New
Orleans. Type locality: Indianola, Calhoun Co., Texas. It will also use salt marshes and has
been reported to be as common in salt marshes as in freshwater (Brown 1950; Guidry 1953).
Identification A heavy-bodied snake reaching 1295 mm. Neonates are 187-305 mm in
total length. Dorsal coloration is olive or brown, and subocular scales keep the labials from
entering the orbit. Dorsal scales are in 25 or 27 rows at midbody and are keeled to two apical
pits. Ventrals 133-148, subcaudals 57-78. Nerodia cyclopion and its sister N. floridana can be
distinguished from other North American Nerodia by the presence of one or more subocular
scales and an indistinct pattern of about 50 mid-dorsal bars alternating with a lateral series of
bars.
Natural History. A highly aquatic species that leaves the water during heavy rains and
moves as much as 100 m from the water (Kofron 1978). It basks during the day and adjusts its
behavior by season. The diet of this species seems to be opportunistic. Gibbons and Dorcas
(2004) consider this species an opportunistic aquatic feeding species that will take salaman-
ders, fish, and crayfish. However, Camp et al. (2020) proposed it is likely a fish specialist sug-
gested by its small eyes. Mushinsky and Hebrand (1977) found almost all fish in the diet of a
Louisiana population.
Reproduction. Copulation occurs in the water and on land (Krofron, 1977; Mead, 1934)
in the spring (April-May), and parturition has been reported in July and August. The smallest
reproductive female had an SVL of 637 mm. Litter sizes range from 8-37.

Florida Green Water Snake

Nerodia floridana (Goff, 1936)
Distribution and Habitat. It uses wetlands choked with vegetation and rarely occurs in
rivers or streams. N. floridana. It also uses lakes, ponds, ditches, and occasionally in brackish
waters from Southern South Carolina into northeast Georgia. The species is absent from most
of Georgia, and the gap forms two disjunct populations. The southern population is in south-
east Georgia, extends into Florida, and extends west to Alabama. The type locality is near
Leesburg, Florida, at the Alsa-Brook Prairie between Lake Griffin and Lake Yale.
Identification A heavy-bodied watersnake and the lon-
gest species of Nerodia, reaching 1880 mm. The dorsum is
olive, brown, or reddish. Dorsal scales are keeled, with two
apical pits, and organized in 25-27 midbody rows. Ventrals
129-142, subcaudals 63-84. Suboculars prevent labials from
bordering the orbit.
Natural History. The diet is composed of aquatic ver-
Figure 29.13. Distribution of Nerodia
cyclopion. tebrates (frogs and fishes). The diet has been summarized
by Gibbons and Dorcas (2004). In southern Florida, N. flor-
idana is active year-round, but on the northern edge of the
range, it hibernates/brumation during the winter. The snake can be seen basking on sunny
days. In southern Florida, it often travels overland on rainy days.
500
 29. The Thamnophiine Clade

Figure 29.14. Nerodia cy-

clopion. (A) Brazos Bend
State Park, Needville, TX.
Photography by J. Batch.
(B) Photography by Mike
Pingleton.

This species does not

respond well to droughts.
Seigel et al. (1995) stud-
ied a watersnake commu-
nity during a drought and
noted that this species
was not seen for five years
after the drought ended.
Willson et al. (2002) did
a similar study and found
this species to be extir-
pated.
Reproduction. Mat-
ing has been reported as
early as February, and
parturition occurs in
captives between June
and August. Litter sizes
as small as seven and as
large as 132 have been
reported (Gibbons and
Dorcas 2002).

Salt Marsh Snake

Nerodia clarkii (Baird
and Girard, 1853)
Distribution and
Habitat. Coastal salt
meadows, swamps, and
marshes rarely enter freshwater systems from Florida, Loui-
siana, Mississippi, Alabama, southeast Texas, and northern
Cuba. (Fossil remains known from Abaco in the Bahamas in
the late Pleistocene). The type locality is Indianola, Calhoun
Co., Texas.
Identification Dorsal coloration is highly variable, gray,
olive, black, yellow, or brick red. The pattern may be uniform,
Figure 29.15. Distribution of
Nerodia floridana. striped, or banded. Dorsal scales are keeled with two apical pits
and organized in 21-23 rows at the midbody. Subocular scales
present. Ventrals 123-138, subcaudals 57-88. Size of adults
380-760 mm; to the maximum size 933 mm; neonate size 197-266 mm;
A polytypic species with three subspecies:

501
Aquatic Snakes

Figure 29.16. Nerodia floridana. Central Florida.

Photography by Jake Scott.
Nerodia clarkii clarki (Baird and Girard, 1853). The type locality is coastal Texas and
Alabama. Adult dorsum ranges from dark gray to red-brown with four yellow longitudinal
stripes (two on each side). The venter is dark gray to red-brown with one to three rows of
pale spots.
Nerodia clarkii compressicauda Kennicott 1860. The type locality is south of Tampa Bay,
Florida. Adults can be gray, gray-olive, brown, tan, or rusty orange, with patterns of dark
longitudinal stripes or transverse bands on the dorsum. However, some individuals may be
uniform; some individuals may be almost entirely black (melanistic).
Nerodia clarkii taeniata Cope 1860. The type locality is Volusia, National Gardens, Volu-
sia County, Florida. Adults have a dorsal pattern of dark brown and pale stripes anteriorly
with dark blotches on a pale olive ground color over much of the body. Ventral surface dark
with a central row of cream or yellow spots.
Natural History. The diet includes fish and crabs, with the snakes frequently sustaining
an injury from the crabs. (Miller and Mushinsky 1990; Mullin 1994). Nerodia clarkii compres-
sicauda forages in and among the prop roots of mangroves. It has also been reported to feed
with the tidal cycle and is active day and night.
Lingual luring has been described in this species
(Hansknecht 2008). It involves the tongue curling upon itself
distally to form a conspicuous loop. The typical chemosen-
sory tongue flicking is absent, although the terminal loop
does exhibit some vertical and horizontal movement. The
duration of luring tongue flicks is significantly greater than
Figure 29.17. Distribution of Nero- chemosensory tongue flicks.
dia clarkii Reproduction. Trauth (1991) collected gravid females
and males undergoing spermatogenesis were found in July.
Therefore, parturition presumably takes place in August or early September.
Population and Biomass. Ackley and Meylan (2010) studied variation in population size
and biomass of Nerodia clarkii compressicauda population in two artificial ponds in St. Pe-
tersburg, Florida, USA. The population’s substantial biomass (25.4 kg/ha, one of the highest
observed in snakes) was maintained for at least six months until the herbicide Aquamaster™

502
 29. The Thamnophiine Clade

Figure 29.18. Nerodia clarkii Salt Marsh Snakes

Photography by JCM.
(glyphosate) was applied to the study area. Two months after application, the dead emergent
vegetation collapsed into the pond. The population declined because of emigration after the
change in habitat structure.
Conservation. Salt marsh snake populations appear to be fractured into isolated neigh-
borhoods on the order of 50–80 km, according to a study done by Jansen et al. (2007). Despite
its apparent local abundance, they suggest N. c. compressicauda needs conservation protec-
tion. The combination of extremely low dispersal, narrow habitat requirements, and, most
503
Aquatic Snakes

importantly, extensive habitat alteration resulting from coastal real estate development may
mean that N. c. compressicauda is highly susceptible to population extirpation and extinction.

Plain-Bellied Water Snake

Nerodia erythrogaster (Forster, 1771)
Distribution and Habitat. Permanent wetlands, rivers, and lakes from Iowa and Texas
eastward to Delaware and northern Florida, southward into the Mexican states of Duran-
go, Zacatecas, Coahuila, and Nuevo León. This is often considered a polytypic species, even
though little genetic evidence supports the subspecies (Makowsky et al. 2010).
Plain-bellied Water Snake, Nerodia erythrogaster alta (Conant, 1963) inhabits Mexico
(Zacatecas). The type locality is in the Rio Trujillo (also known as the Rio Florido), at the
village of Rio Florido, approximately 15 miles NW of Fresnillo, Zacatecas.
Bogert’s Water Snake, Nerodia erythrogaster bogerti (Conant, 1953). The type locality is
in the Rio Nazas, near La Goma, approximately 15 miles southwest of Lerdo, Durango.
Redbelly Water Snake, Nerodia erythrogaster erythrogaster (Forster, 1771). The neotype
locality is near Parker’s Ferry, Edisto River Swamp, Charleston County, South Carolina.
Yellowbelly Water Snake, Nerodia erythrogaster flavigaster (Conant, 1949). The type lo-
cality is Frenier Beach, St. John the Baptist Parish, Louisiana.
Copperbelly Water Snake, Nerodia erythrogaster neglecta (Conant, 1949). The type local-
ity is approximately 3 miles east of Mount Victory, Hardin County, Ohio.
Blotched Water Snake, Nerodia erythrogaster transversa (Hallowell, 1852). The type lo-
cality is Arkansas, Boundary of Creek Nation, Arkansas River.
Identification. A large watersnake, reaching 1636 mm in total length. Olive gray dorsum
with faint crossbars. The ventral surface may be red, yellow, or orange. Dorsal scales keeled
with paired apical pits; dorsals in 23-25 rows at midbody. Ventrals 132-116, subcaudals 46-90.
No suboculars.
Natural History. They will wander far from the water and spend considerable time out of
the water. They are active day and night in the warmest months of the year. They can be seen
basking on logs or shorelines in the spring and fall. They hibernate during the coldest months
of the winter. Prey includes fish and amphibians.
Nerodia erythrogaster neglecta is well studied because, in 1997, it was designated a threat-
ened species in Ohio, Michigan, and northern Indiana under the Federal Endangered Species
Act. It migrates seasonally from its summer bottomland
forests and shrub swamp habitat to more upland habi-
tats through vegetated corridors to wetland areas. How-
ever, some populations stay in their lowland habitat and
hibernate in rotten stumps and burrows made by other
animals. Sometimes these burrows are flooded and are in
saturated soils. Copperbelly Water Snakes emerge from
Figure 29.19. Distribution of Nero- their upland hibernation sites in early spring and migrate
dia erythrogaster through wooded habitats to wetlands. When the wood-
land swamps begin to dry in late spring or in early June,
the snakes again disperse and move through wooded or
vegetated corridors to their summer habitat areas. Summer activities usually center around
forests and forest edges (Conant 1951, Kingsbury and Coppola 2000).
For this reason, an upland habitat is essential for the snake’s summer foraging activities.
In late fall, Copperbelly Water Snakes seek out hibernation sites. It was believed that Copper-

504
 29. The Thamnophiine Clade

belly Water Snakes use hi-

bernation sites at elevations
higher than the flood line
in pond areas (Sellers 1991).
However, in radio-teleme-
try studies, Kingsbury and
Coppola (2000) found they
use bottomland hibernation
sites. Bottomland hiberna-
tion sites have been identi-
fied as cavities in felled tree-
root networks (Lodato 1985),
crayfish burrows (Kingsbury
and Coppola 2000), dense
brush piles, rockpiles, and
perhaps beaver and musk-
rat lodges (Sellers 1991). A
mid-winter flood and freez-
ing temperatures could be
lethal and decimate the local
population if floodplain and
riverbank areas are the only
hibernation sites available.
This species is known
to form small groups in the
spring and fall. Groups of
snakes have been observed
swimming, feeding, court-
ing, and resting together
Figure 29.20. Nerodia erythrogaster. (A) Photography by Mike Pin- (Conant 1934; Sellers 1991).
gleton (B) N. e. transversa from Kansas. Photography by JCM. Copperbelly Water
Snakes have a more extended
gestation period than other water snakes sharing their range, and their average litter size (18)
is also smaller (Schmidt and Davis 1941). Young snakes are born in the fall near, or in, the
hibernaculum and may not become active until the following spring.
The diet comprises fish, crayfish, salamanders, and frogs. It uses both active foraging and
sit-and-wait strategies to find food. Prey is seized and swallowed without constriction. Like
some other Nerodia, this species has been observed swimming underwater with its mouth
open. Platt et al. (2020) reported this species stealing food (kleptoparsitism) from a Nerodia
rhombifera. McAllister (2021) wrote a female from McCurtain Co., Oklahoma, that contained
remains of arthropods; a caterpillar, a grasshopper, beetle wings, and a harvestman. Perkins
and Eason (2017) reported them feeding on the Bowfin, Amia calva, and the Pirate Perch,
Aphredoderus sayanus.
Mating occurs April-June in the southeast USA, and parturition occurs in August-Sep-
tember. Litters of 55 have been reported. Most, however, contain 18 young. Parthenogenesis
has been documented in this species.
Predators are terrestrial and aquatic and range from other snakes and large fish to wading
birds and carnivorous mammals. Unlike the typical water snake, the plain-bellied water snake

505
Aquatic Snakes

will leave the water and try to escape over land if threatened (Gibbons and Dorcas 2004).
The Plain-bellied Water Snake is considered a conservation risk because it tends to travel
over land on roads and highways. Destruction or damage to wetlands also threatens the spe-
cies’ habitat. It is not a protected species in the southeastern states. Everitt and Philips (2019)
reported death feigning in this species.

Banded Water Snake

Nerodia fasciata (Linnaeus, 1785)
Distribution and Habitat. Lakes, marshes, ponds, and streams from Texas eastward to
Florida as far north as southern Illinois and North Carolina. In 1992, Nerodia fasciata and N.
sipedon were found at three sites in California by the US Fish and Wildlife Service. In 2009,
more than 300 banded water snakes were caught in the suburbs of Los Angeles by the Nerodia
Working Group of USFWS. Then in May 2016, the species was found in the Colorado Riv-
er basin near Yuma, Arizona, where a well-established population can be seen today.
This is a polytypic species with three subspecies.
Nerodia fasciata confluens (Blanchard, 1923) has its type locality in Butler County, Mis-
souri. They were distinguished by 11 to 17 dorsal transverse bands (Clay 1938).
Nerodia fasciata fasciata (Linnaeus, 1766) has a type locality of Carolina. It is identified
by the shape of the ventral markings that are quadrangular near the anterior margins of the
ventrals (elongated in pictiventris) and by the presence of more than 128 ventrals (fewer in
pictiventris) (Clay 1938).
Nerodia fasciata confluens has 11-17 dorsal blotches and N. f. fasciata has 19 to 30 (Clay
1938). Nerodia fasciata pictiventris (Cope, 1895) has a type locality in Gainesville, Florida. It
can be distinguished by 26-35 red-brown to black cross bands; mature individuals have lateral
interspaces that sometimes become darker than the lateral portions of the cross-bands and
thus produce an appearance of alternating dorsal and lateral spots (Clay 1938).
Identification. A large watersnake, to 1588 mm. Dorsum can be gray, tan, brown, or red-
brown with dark brown transverse bands (older individuals may be a uniform black). Ventral
surface yellow or cream marked with dark spots or transverse bars. Dorsal scales are strongly
keeled with a pair of apical pits; at midbody, the scales are in 17 rows. The ventrals are 120-
143, and the subcaudals are 50-89.
Natural History. The diet includes fish and frogs with an ontogenetic shift from fish to
frogs with maturity (Mushinsky et al. 1982). Many fish species are eaten, as well as frogs,
salamanders, small turtles, snakes, birds, crayfish, and earthworms (see Ernst and Ernst 2003
for a complete list). A Florida specimen was found to have eaten an introduced Asian clariid
catfish (Donini 2018). Lamb and Ljustina (2018) reported a specimen coiled under water
below a film of Green Frog eggs and was feeding on the spawn. Fuchs et al, (2020) found an
individual with its jaws stuck in the shell of the snail Pomacea
maculata in an apparent attempt to prey on the snail.
Females reach sexual maturity at body lengths of 450 mm
and males at 520 mm. Females provide nutrients and calories
via a placenta to their embryos. Gestation is 48-109 days, de-
pending on temperatures. Litter size varies from 9 to 50. New-
Figure 29.21. Distribution of borns are 200–310 mm long (Ernst and Ernst 2003).
Nerodia fasciata. Hybridization with Nerodia sipedon. Mebert (2005/2008)
found widespread hybridization in the contact zone between
Nerodia sipedon and Nerodia fasciata in the Carolinas with most specimens exhibiting a

506
 29. The Thamnophiine Clade

Figure 29.22. Nerodia fasciata. A. From central Florida.

Photography by JCM. B. Photography by Mike Pingleton

backcrossed hybrid status. However, the distributions of diagnostic genetic markers between
species are asymmetric, and the frequencies of snakes resembling a F1 interspecific hybrid
(50/50% fasciata/sipedon) are very low. This and the pronounced morphological characteris-
tics of N. fasciata in most of the studied hybrid zone is in accordance with the habitat relating
more with that species.
Tail autotomy has been reported in this species and some other species of thamnophiini

507
Aquatic Snakes

(N. erythrogaster, N. sipedon, Thamnophis sirtalis, T. sauritus) (Donini 2019). In addition,

Everitt and Philips (2019) reported death feigning in this species.

Brazos Water Snake

Nerodia harteri (Trapido, 1941)
Distribution and Habitat. Rock-strewn shorelines along the Brazos River system in Cen-
tral Texas and stream impoundments. Restricted to the upper Brazos River drainage, along
303 km of the stream, it is in two reservoirs; this snake has one of the smallest ranges of any
North American snake species (Scott et al. 1989). The species is not restricted to rocky rif-
fles in flowing rivers but also inhabits pools and lakes. Several sections of the Brazos do not
support this species because the rocky riffles needed by juveniles are absent or are separated
by greater distances on the uninhabited stretches of the river. Adults are not restricted to the
riffles. Scott et al. (1989) observed them swimming or on land within 3 m of the shoreline
of a river or lake, and the great majority were within one me-
ter of the water. They may follow intermittent streams a short
distance during high water. The type locality is north of Palo
Pinto, Palo Pinto County, Texas.
Identification. A small, slender water snake that reaches
900 mm (most are less than 600 mm). The dorsum is brown,
gray, or orange, with a lateral pattern of spots that may fuse on
Figure 29.23. Distribution of the midline to form saddles. Dorsal scales keeled with two api-
Nerodia harteri cal pits in 17 rows at midbody. Subocular scales absent. Ven-
trals 143-151, subcaudals 67-88.
Natural History. Active in spring and summer, they winter
underground. Fish are likely the most important prey; the species may also eat tadpoles and

Figure 29.24. Nerodia harteri.

Photography by Mark Pyle.

frogs. Males reach sexual maturity in one or two years at a body length of 42-52 cm. Females
mature in two or three years. Litters of 4-24 are born in late July – September. Litter size cor-
related with female body size.

508
 29. The Thamnophiine Clade

Concho Water Snake

Nerodia paucimaculata (Tinkle and Conant, 1961)
Distribution and Habitat. The Concho River system in Coke, Runnels, San Saba, and Tom
Green Counties. The type locality is the Colorado River south of Robert Lee, Coke County,
Texas city limits. It inhabits shallow, fast-flowing water with a rocky or gravel substrate.
Identification. Closely related to N. harteri but smaller, it reaches 652 mm. Dorsum red-
brown or olive-brown with dark brown spots. Dorsal scales are keeled with a pair of apical
pits and organized into 21 rows at mid-body. Ventrals 139-151, subcaudals 64-83. Subocular
scales absent.
Natural History. This species is a fish specialist, Green et al. (1994) found it feeds primar-
ily on cyprinid fishes but will also eat the hylid frog Acris crepitans. Feeding occurs from mid-
April to early November, and gravid females feed until early June and resume after parturition.
Specimens from lacustrine environments can be observed on the bottom or in shallow water.
Hibernation begins in late October to late November, depending upon weather and tem-
peratures. Most adults probably hibernate in burrows of other animals, particularly crayfish,
and juveniles use crevices under rocks on gravel bars (Werler and Dixon 2000). The snakes
emerge in mid-March to mid-April.
Reproduction. Males reached sexual maturity at SVLs of 380 mm and females at 460 mm.
Mating occurs in April and May; gestation is about three months, and parturition occurs from
late July to mid-August. About 85% of the females in the population are gravid each year.
Litter size ranges from four to 29 and averages 11—neonates average 175 mm in total length
(Greene et al. 1992).
Conservation. Its limited range resulted in a threatened species status. It was placed on
the state endangered list by the State of Texas in 1977. Flury and Maxwell (1981), in a detailed
report on Nerodia paucimaculata, estimated its total population at only 332-613 individuals.
In 1986 it was considered threatened. The Concho water snake was removed from the U.S.
Endangered Species list in 2011. When delisting, Janecka et al. (2021) recently monitored the
snake by conducting extensive surveys and a population genetic structure assessment. They
found no Concho Water Snakes along the Concho River, part of the previously known range,
and fewer snakes on the Colorado River than in previous studies. Several population genetic
analyses showed concordant patterns of population substructure along the Colorado River,
and genetic bottlenecks were detected. Habitat modification
and low water flow during periods of drought may have
contributed to existing patterns. The genetic-based results
demonstrate that the Concho Water Snake exists as small
subpopulations along the Colorado River.

Figure 29.25. Distribution of

Nerodia paucimaculata.

Diamondback Water Snake

Nerodia rhombifer (Hallowell, 1852)

509
Aquatic Snakes

Distribution and Habitat. Slow-moving rivers, lakes, marshes, swamps, and anthropo-
genic bodies of water from eastern Kansas, extreme southeastern Iowa, southern Illinois, and
southwestern Indiana southward through western Kentucky and Tennessee, central Alabama,
Mississippi, Missouri, Arkansas, Oklahoma, and eastern two-thirds of Texas southward into
Mexico to Tabasco and southwest-
ern Campeche.
This polytypic species has
three subspecies: Nerodia rhombi-
fera blanchardi (Clay, 1938). The
type-locality is Mexico, within a ra-
dius of 85 miles of Tampico in the
triangle formed by the Rio Tamesi
and Rio Panuco.
Nerodia rhombifera rhombifera
(Hallowell 1852) has a type-locality
in the Arkansas River and its tribu-
taries near the northern boundary
of the Creek Nation. Revised to Ar-
kansas River between Keystone and
Tulsa, Tulsa County, Oklahoma by
Conant (1969).
Nerodia rhombifera werleri
(Conant, 1953). The type locality is
the Alvarado-San Andres Tuxtla road
(Mexico No. 180), approximately 18
miles southeast of Alvarado, Veracruz,
Mexico.
Identification. This is a large water
snake reaching 1753 mm. The dorsum
is brown or gray with connected
dark brown squares or diamonds.
The venter is cream to yellow with
dark half-moon blotches. The dor-
sal scales are keeled with paired api-
cal pits; at midbody, they are in 21
rows. Ventrals 132-152, subcaudals
are 56-88. Subocular scales absent.
Natural History. The literature
on this snake is extensive and not
completely covered here. A labora-
tory investigation of the feeding be-
havior of juvenile Nerodia rhombif-
era (Savitzky and Burghardt 2000)
found neonates were negatively Figure 29.26. Nerodia paucimaculata. Photography by (A)
buoyant, foraged only in the wa- Romey Swanson (B) Scott Wahlberg (C) Paul Freed.
ter, and were successful in catching
fish. The open-mouth foraging behavior seen in adults was rudimentary in neonates. There
was an ontogenetic shift for more underwater foraging, less foraging on the surface, and in-
creased open-water use. The results were congruent with field observations.

510
 29. The Thamnophiine Clade

Diet. The prey base for this species is highly diverse. In a literature survey combined with
stomach content analysis, Kalki
et al. (2018) found 55 species of
fish, nine species of amphibians,

Figure 29.27. Distribution of

Nerodia rhombifer.
eight species of insects (some of
these may represent secondari-
ly ingested items), five crusta-
ceans, and one species each of
the following taxa: snake, turtle,
bird, and a bat. Perkins and Ea-
son (2017b) reported this snake
feeding on the Pirate Perch,
Aphredoderus sayanus.
Reproduction. Both sexes
mature in 2.5 to three years; the
smallest reproductive male was
475 mm (but most do not ma-
ture until they reach 680-800
mm, and females mature at 650
mm to 680 mm. Mating occurs
after emergence from hiberna-
tion in April and May. Partu-
rition usually occurs from late
July to early November. Howev-
er, one female from Texas had
full-term embryos in February,
suggesting that reproductive
timing in this snake is variable.
Litter size ranges from eight to
62; the mean is 26.8. Neonate
size is 200-332 mm in total
length. There is a direct cor-
relation between litter size and
female body size.
Figure 29.28. Nerodia rhombifer. Photography by (A) Mike Pin-
Northern Water Snake gleton (b) Paul Freed (c) JCM.
Nerodia sipedon (Linnaeus,
1758)
Distribution and Habitat. Natural and anthropogenic bodies of water from rivers and

511
Aquatic Snakes

lakes, stream impoundments, canals, roadside ditches, wetlands, marshes, and swamps from
Maine, southern Quebec, and southern Ontario westward to southeastern Minnesota, then
southwestward through Nebraska to northeastern Colorado. The range extends to the south-
east from Colorado throughout most of Kansas, eastern Oklahoma, and Arkansas, and east-
ward through Missouri to the Mississippi River. East of the Mississippi River, it occurs from
Canada to the Gulf Coast in Louisiana; from there, it ranges eastward to the Florida Panhan-
dle and throughout the remainder of the eastern states except below the Fall Line in Georgia
and the Carolinas. Nerodia sipedon and Nerodia fasciata are sympatric at numerous points
and hybridize in some areas (Gibbons and Dorcas 2004). The type-locality, North America,
was reported by Linnaeus (1758). Nerodia sipedon is primarily freshwater, but some coastal
populations inhabit brackish water.
This is a polytypic species.
Nerodia sipedon insularum (Conant and Clay 1937) Type-locality, Pelee Island, Lake
Erie. This pale subspecies is confined to islands in Lake Erie’s Ohio and Ontario portions.
The dorsal pattern is reduced or completely lacking, and the dorsal ground color is usually
green or brown with an occasional pinkish or orange tinge. The dorsal pattern, when visible,
is like that of the nominate subspecies. Ventrals range from 141–153 (average 146.3) in males,
and 137–152 (average 145.9) in females, with subcaudals ranging from 70–81 (average 75.6)
in males and 58–68 (average 63.3) in females (based upon Walley et al. 2012).
Nerodia sipedon pleuralis (Cope) has an unknown type-locality. Cochran (1961) gave
it as Summerville, Dorchester, South Carolina. A neotype designated by Clay (1938) is at
USNM. The dorsal pattern is like that of N. s. sipedon, but the dark dorsal markings usually
number 30 or fewer and are separated by paler interspaces that are wider than the bands
(Gibbons and Dorcas 2004). The crescentic ventral blotches often break up and disappear
with age and are not nearly as bright as those of N. s. sipedon.
Nerodia sipedon williamengelsi (Conant and Lazell). Island Creek on Ocracoke Island,
Hyde Co., North Carolina, is the type-locality. This coastal
subspecies closely resembles N. s. sipedon in scale count data
differs in having the adult dorsum essentially black with the
interspaces between the crossbands and blotches so dark that
pattern details may be obscured. The taxon is restricted to
brackish marshes along the margins of Pamlico and Croatan
sounds. This taxon occurred in salt marshes and tidal creeks
and was common in such habitats on the barrier islands. Gaul
(1996) noted clinal variation in several morphological char-
Figure 29.29. Distribution of
Nerodia sipedon. acters within its known range. These clines correspond with
clines in salinity in the coastal waters of North Carolina (Gaul
and Mitchell 2007).
Identification. A medium to large water snake that reaches 1500 mm in total length. The
dorsum is tan to gray with wide red-brown, alternating, quadrangular blotches along the
midline of the back. The venter is cream to yellow with a half-moon marking present. Older
snakes tend to become uniform black. Dorsal scales are keeled with two apical pits and are
organized into 23 rows at mid-body. Ventral scales are 123-155, subcaudal scales are 42-84.
Natural History. The Northern Water Snake is one of the best-studied aquatic snakes.
The extensive distribution from Maine and southern Canada southward to the Gulf Coast
of Alabama results in northern populations spending September to April-May in hiberna-
tion. The southern populations may not hibernate; if they do, they spend only a few days
to weeks escaping cold temperatures. Hibernation sites include rock crevices, stream banks,

512
 29. The Thamnophiine Clade

cavities formed by the roots of large trees, and animal burrows. These may be close to water
or a significant distance from water (hundreds of meters) and may be in upland situations or
lowland areas. They occasionally use beaver, muskrat lodges, or terrestrial rodent burrows for
hibernacula. They are known to hibernate alone as well as in communal groups.
Body Temperatures. Nerodia sipedon spends significant time basking along shorelines
on the substrate, vegetation, or emergent debris. Therefore, body temperatures in the field
generally range from 20.8 to 34.7 °C with a mean of 28.0 °C.
Diet. Northern Water Snakes are fish specialists. Ernst and Ernst (2004) list prey records
from free-living specimens containing 71 species of fish, 11 species of salamanders, 15 species
of anurans, one species of snake, and two species of mammals. They also report invertebrates:
crayfish, other arthropods, snails, earthworms, and leeches.
Reproduction. The population’s latitude influences the timing of maturity and the devel-
opment of gametes. Males mature at 370 to 450 mm SVL at 21-24 months. Females mature at
SVLs of 500-650 mm and 24-36 months of age (Feaver 1976, King 1986, 1994). Females grow

Figure 29.30. Nerodia sipedon williamengelsi. is a salt marsh dwelling population of this species.
Photography by JCM.

their ovarian follicles in a four-to-six-week period in the spring. They increase the yolk before
ovulation. Females carry fertilized eggs in June, and gestation is 9 to 12 weeks. Nerodia sipe-
don embryos do have a placental connection to the female. However, there is some evidence
that the placenta transfers water and minerals but not nourishment to their embryos (see
Weatherhead et al. 1999). Males produce spermatozoa after the breeding season, use some in
the fall, and store some until the following spring. Mating starts immediately after emergence
from hibernation. Males actively search for females, following pheromone trails formed by
the female. Females mate with multiple males, and litters have numerous fathers. Senter and
Senter (2020) have described courtship and mating.

513
Aquatic Snakes

Figure 29.31. Nerodia sipedon from northeastern Illinois. JCM

Defense and Predators. When confronted by a predator, Nerodia sipedon will always try
to escape, often diving into the water. If it cannot escape, it will flatten its head and body,
strike, and bite if it can. If the snake is seized, it will not hesitate to release foul-smelling clo-
acal musk. In addition to smelling bad, it may be toxic and irritating to mucus membranes.
The range of predators is diverse – predatory fishes, bullfrogs, alligators, snapping turtles,
other snakes, wading birds, raptors, gulls, carnivorous mammals, and of course, humans.

Brown Water Snake

Nerodia taxispilota (Holbrook, 1838)
Distribution and Habitat. Flowing, permanent waters such as rivers, canals, blackwater
cypress creeks, and stream impoundments on the Coastal Plain and Piedmont regions of the
southeast USA from southern Virginia southward through all of Florida and westward to
eastern Alabama. Brown Water Snakes are found throughout Georgia and South Carolina

514
 29. The Thamnophiine Clade

but are absent from the mountainous regions of extreme northwestern South Carolina and
northern Georgia. Their preferred habitat has overhanging vegetation, emergent snags, or
rocky riverbanks where snakes may bask. The type-locality was given as the South Carolina
seaboard and the Altamaha River neighborhood in Georgia.
Identification. A large water snake that reaches 1766 mm in total length. It has a thick
body with a brown dorsum and three rows of dark brown blotches. The ventral surface is
cream to tan with half-moon markings. The dorsal scales are keeled with two apical pits and
are organized into 29-31 rows at the midbody. The ventrals are 128-152; the subcaudals are
59-87.
Natural History. Brown water snakes were studied by Mills (2002) on the Savanna Riv-
er. He found they were not randomly distributed but were significantly associated with the
steep-banked outer bends of the river and the availability of potential perch sites. Sections of
the river with the highest number of captures were clustered within 200 m of backwater areas.
Seventy percent of 164 recaptured snakes were less than 250 m from their previous capture
site; however, three moved more than a kilometer. Only eight large individuals (800 mm
snout-vent length) crossed the river (approximately 100 m).
Brown Water Snakes are active in all months of the year but are most infrequently seen
from November through February in the Savanna River. However, they are likely active year-
round in Florida and much of its range. Radiotelemetry studies suggest N. taxispilota will
emerge and sometimes move short distances on warm winter days regardless of water level.
They are diurnal but may shift to nocturnal behavior at the warmest time of the year.
Diet. Brown Water Snakes are fish specialists, and the largest portion of their diet com-
prises ictalurid catfishes (Camp et al., 1980). Mills (2002) supports this with his data from
the Savannah River. Of the identifiable food items, all were fish, and 62% were catfishes in
the genera Ictalurus and Noturus. Snakes over 60.0 cm SVL consume almost exclusively cat-
fish. Although a variety of other non-fish prey has been reported in the diet, including frogs,
crayfish, a turtle, small snakes, and lizards, these observations are few. They may represent
secondarily ingested prey (Neill and Allen, 1956).
Reproduction. The seasonal reproductive cycle of Nerodia taxispilota in southeastern
Virginia was found to resemble other temperate zone colubrids by White et al. (1982). Testes
are small during April-June and largest in August, decreasing in size during September-No-
vember. Spermatozoa produced in late summer are stored in the vas deferens through winter
and used the following spring. The female cycle is annual. Vitellogenesis occurs from April
to June, ovulation is in late June, and parturition is in early to mid-September. Fecundity
increases with increasing female body size. The average total clutch size is 33.9, and the mean
size of full-term embryos is 28.0. Sequential stages of embryonic development are described.
Males average significantly smaller than females. The percentage of tail breaks is about equal
in both sexes, but adults have higher proportions of tail breakage than juveniles. Sex ratios
do not differ significantly from 1:1; however, males predominate in autumn. Males mature at
about 503 mm SVL (about two or three years of age), and females mature at about 725 mm
(about four or five years of age) (White et al. 1982, Ernst and Ernst 2003).
The Genus Thamnophis - Garter Snakes
The garter snakes, genus Thamnophis, include some of the western hemisphere’s most
abundant and well-studied snakes. A few species spend considerable time in the water; a few
are terrestrial or terrestrial-arboreal. Molecular studies in the past two decades have suggest-
ed the two species of Adelophis belong to Thamnophis, but no one has formally moved them
to the garter snake genus. There are 37 species, including the two species Adelophis.

515
Aquatic Snakes

The distribution of Thamnophis extends further north than any other squamate in the
western hemisphere, there are populations of Thamnophis ordinoides in southeast Alaska and
Canada’s Northwest Territories, and the genus is represented as far south as Panama. Garter
snakes are primarily diurnal; they use ambush and actively for-
aging to capture prey, and they use venom for subduing prey.
They have chorioallantoic and yolk-sac placentation. At least a
few species can hibernate underwater. And a few species have
been documented to have facultative parthenogenesis.
Most recently, Hallas et al. (2022) found support for two
clades of Thamnophis, a northern North American clade and a
Figure 29.32. Distribution of southern North American clade. Divergence time estimates and
Nerodia taxispilota.
biogeographic analyses suggest Thamnophis originated in the
mid-Miocene of Mexico. Their studies of morphological traits
associated with feeding ecology showed differences in particular cranial morphologies be-
tween aquatic specialists and garter snakes that are terrestrial-aquatic generalists, indepen-
dent of evolutionary history. Hallas et al. (2022) consider Thamnophis atratus, T. couchii, T.
gigas, T. hammondii, T. melanogaster, T. nigronuchalis, T. rufipunctatus, and T. validus aquatic
specialists, and grouped all the remaining species

as terrestrial or aquatic generalists. Considering all other Thamnophis as terrestrial-aquatic

generalists may be misleading. Many taxa could be partitioned into different categories based
on diet, terrestrial habits, and feeding ecology.
Aquatic-specialists occur in both clades. For example, species that hunt aquatic prey
have long, narrow muzzles to reduce hydrodynamic drag. In contrast, those with broader or

Figure 29.33.. Nerodia taxispilota from northern Florida. JCM.

mostly terrestrial diets have wider muzzles and deeper heads to accommodate an array of

516
 29. The Thamnophiine Clade

potentially large prey (Hallas et al. 2022).

Aquatic Garter Snake

Thamnophis atratus Kennicott, 1860
Distribution and Habitat. Fast-flowing streams with rocky substrates, sluggish streams
with soft bottoms, ponds, small lakes, and adjacent terrestrial habitats in woodlands, transi-
tional woodland-grass ecotones, or chaparral. It is closely tied to water, but individuals some-
times move overland. The Aquatic Garter Snake ranges from Coos and Douglas counties
in southwest Oregon southward to Santa Barbra County, California (Nussbaum et al. 1983,
Rossman et al. 1996, Ernst and Ernst 2003, Stebbins 2003).
This is a polytypic species with three subspecies. Thamnophis a. atratus is restricted to
the outer coastal area of San Francisco Bay, California. The type locality was given as Califor-
nia but limited by Fitch (1940) and Cochran (1961) to San Francisco, California.
Thamnophis a. hydrophilus Fitch, 1936 occurs north of San Francisco Bay northward to
southwest Oregon. Type Locality is Trail Creek six miles from its mouth in Jackson County,
Oregon.
Thamnophis a. zaxanthus Boundy, 1999 occurs from the Inner Coast Range from Napa
and Solano to Santa Barbara counties and the Santa Lucia Range. The type locality is two
miles south of Gilroy Hot Springs, Santa Clara County, California.
Identification. The species reaches a maximum of 1060 mm. Dorsal scales are organized
in a maximum of 19 or 21 rows. Dorsum gray, brown, dark brown, or black may have two
rows of black spots between the stripes; the vertebral strip may be distinct, indistinct, or ab-
sent. Lateral stripes are confined to rows two and three.
Natural History. Adults have been observed anchoring their tails around submerged
rocks in deep water and striking at fish swimming by (Boundy, 1999), and neonate snakes
fed on smaller prey that inhabited shallow stream margins (Lind and Welsh 1994). Lind and
Welsh (1994) found adults took a wider variety of prey, but they focused on the Pacific Giant
Salamander, Dicamptodon tenebrosis, larvae, and neotenes, in the mid-stream substrates. Ju-
venile and Consumption of large prey by adults and smaller prey by juveniles indicate a shift
in foraging strategy from frequent feeding on small prey to infrequent feeding on large prey.
Neonates have an unusual feed behavior described by Welsh and Lind (2000), while
adults are active foragers feeding on the aquatic Pacific Giant Salamanders on streambed
substrates. Although juvenile snakes use primarily ambush tactics to capture larval anurans
and juvenile salmonids along stream margins, their behavior includes lingual luring. Luring
snakes are in an ambush position and extend and hold their tongues out and rigid, with
the tongue-tips quivering on the water’s surface; apparently, this mimics insects and attracts
young fish within striking range.
Preston and Johnson (2012) investigated the diet of T. atratus in the San Francisco Bay
Area. At one-hundred and eighty-five ponds, they captured 139 T. atratus, and 60 contained
identifiable stomach contents. Native amphibians were found in 93% of the snakes contain-
ing food. Pacific Chorus Frogs (Pseudacris regilla) were the most important amphibian prey,
followed by Western Toads (Anaxyrus boreas), California Newts (Taricha torosa), and Cali-
fornia Red-legged Frogs (Rana draytonii). T. atratus at a pond was positively associated with
all native amphibian species but negatively associated with introduced American Bullfrogs
Lithobates catesbeianus. The mean species richness of native amphibians at ponds where they
detected T. atratus was also higher than that in ponds without the garters snakes (2.45 vs.
1.74), and the odds of finding T. atratus at ponds with native amphibians was 12 times greater
517
Aquatic Snakes

than at ponds without native amphibians.

Bogert’s Garter Snake

Thamnophis bogerti Rossman and Burbrink, 2005
Fiogure 29.34. The distribu- Distribution and Habitat. Oak woodland, pine-oak forest,
tion of Thamnophis atratus.. and pine-oak- madroño forest in the Mesa del Sur of Oaxaca,
exclusive of the Sierra de Juarez. The type locality is Mexico,
Oaxaca, El Tejocote, at 2377 m ASL. They are probably restrict-

Figure 29.34. Thamnophis atratus. Photograph from Mike Pingleton.

ed to elevations of 2195-2743 m (Rossman and Burbrink, 2005).

Identification. The male holotype was 298 mm. The dorsum is brown, with two rows of
black spots between the stripes. Dorsal scales are in 17 rows at the midbody. The top of the
head is unpatterned. Ventrals average 145 in males, 140 in females. Subcaudals average 70
in males and 62 in females.
The dorsum is brown, with small, irregular black spots barely visible. The light vertebral
stripe is confined to the vertebral row and is difficult to
discern on the lower edge of dorsal scale row three. The nuchal
blotches are predominantly black and fused mid-dorsally
(the vertebral stripe is separated from the interparietal
notch by 4-1 /2 scales). Prominent black barring along the
posterior margins of upper labials 2-5 extends ventrally
Figure 29.35. Distribution of
Thamnophis bogerti.
50-75% of the height of each suture; along the posterior
sutures of SL 6 and SL 7 (which includes anterior upper

518
 29. The Thamnophiine Clade

labial eight), the bars reach the Figure 29.36. Thamnophis bogerti (A) from San Vicente Nuñú,
lip. A black stripe extends along Oaxaca, MX. Photograph from José Luis Mendoza Betanzos (B)
the anterior temporal’s lower from San Agustín Loxicha, Oaxaca, Mexico. Photograph from
margin, connecting the two hfranz.
posterior most SL bars. The lower labials are unmarked except for a black bar along the lower
labials nine and ten suture. (Rossman and Burbrink, 2005). Natural History. Unknown.

Conant’s Garter Snake

Thamnophis conanti Rossman and Burbrink, 2005
Distribution and Habitat. Oak woodland is confined to the
Figure 29.37. Distribution of
Thamnophis conanti. southern interface of the Mesa Central and the Sierra Madre
Oriental near the Puebla-Veracruz state line. Recorded eleva-
tions likely to be valid range from 2134-2256 m ASL. Type locality: Mexico, Puebla-Veracruz

519
Aquatic Snakes

state line at Mexico High-

way 125 [number in error,
it is Highway 150 that ex-
tends north from Tehua-
cana, Puebla, to the state
line at Puerto del Aire
(Rossman and Burbrink,
2005).
Identification. Males
reach 520 mm. Dorsal
scales are in 17 rows at
the midbody. Top of head
unpatterned. The area be-
tween the stripes is brown,
with two rows of relatively
Figure 29.38. Thamnophis conanti small black spots between
Photograph from Mike Pingleton. the light vertebral and later-
al stripes. Ventrals average
150 in males and 144 in fe-
males. Subcaudals average 72 in males and 64 in females.

Liner’s Garter Snake

Thamnophis lineri Rossman and Burbrink, 2005
Distribution and Habitat. Confined to pine-oak forest and pine-oak- madroño forest
in the Sierra de Juarez portion of the Mesa de! Sur in Oaxaca. Elevations range from 2700-
2786 m. The type locality is Mexico, Oaxaca, Llano de las Flores, 2786 km elevation.
Identification. The adult male ho-
lotype was 307 mm. The dorsum is
brown, with two rows of spots between
the stripes. Dorsal scales are in 17 rows
at the midbody. Maxillary teeth 18-20.

Figure 29.39. Distribution of Thamno-

phis lineri.

Figure 29.40. Thamnophis lineri (A)

From Llano de las Flores, Oaxaca, MX.
Photograph from Iván Ahumada/Herp.
mx . (B) is from San Juan Bautista Atat-
lahuca, Oaxaca, MX. Photograph from
Eli Garcia-Padilla.

520
 29. The Thamnophiine Clade

The top of the head is unpatterned. Ventrals average 140 in males and 136 in females. Subcau-
dals average 62 in males and 55 in females.

Short-headed Garter Snake

Thamnophis brachystoma (Cope, 1892)
Distribution and Habitat. It occurs in old field habitats near marshes and is frequently
under rocks. They are known from southern Michigan, northwest Pennsylvania, and western
New York. The type locality is
the Allegheny River near Frank-
lin, Venango County, Pennsyl-
vania.
Identification. Maximum
total length 556 mm. Dorsum
is brown or gray-brown, the
stripes are yellow-tan, and the
area between the stripes is un-
spotted. Dorsal scales in 17
rows, occasionally 19 rows. Up-
per labials six, lower labials sev-
en. The lateral stripe is confined
to rows two and three.
Natural History. This is
a very terrestrial species of
garter snake. The diet is com-
posed of almost entirely earth-

Figure 29.41. Distribution of

Thamnophis brachystoma.

worms. Males mature at about

28.0 cm in total length; fe- Figure 29.42. Thamnophis brachystoma Photograph from (a)
males mature at 330 mm. Brandon Hunsberger and (b) Jeff Tome.
Males produce sperm as they
emerge from hibernation,
and the testes reach maximum size by late July or early August. Sperm over winter in the
vas deferens and epididymitis. Mating occurs in the spring; parturition occurs in Au-
gust-September. Neonates are 118-145 mm in total length. Lethaby and Gray (2015) esti-
mated the population size to be 93 snakes and was estimated at 26 snakes/ha. The estimat-
ed biomass was 1.42 kg. At a second site, the population was estimated to be 161 snakes.
The density of snakes was estimated at 34 snakes/ha with a total biomass of 1.72 kg.

521
Aquatic Snakes

Butler’s Garter Snake

Thamnophis butleri (Cope, 1889)
Distribution and Habitat. Mesic prairies, marsh shore-
lines, grassy shorelines of streams and canals, and old fields
Figure 29.43. Distribution of
Thamnophis butleri. in Northeastern Indiana, Ohio, Michigan, and southeast
Wisconsin, USA, and in southern Ontario, Canada. The
type locality is Richmond, Indiana.
Identification. The species reaches a maximum length of 737 mm. The dorsum is ol-
ive-brown or tan to black. Pale individuals tend to have spots in two rows. The stripes are
yellow and involve rows
2 and 3. The ventral
surface has two rows of
spots on the outer edges
of the ventrals. Ventrals
129-151, subcaudals are
49-72.
Natural History.
This species is an an-
nelid worm specialist.
Experimental evidence
suggests that they rec-
ognize the odor of
worms within days of
birth. They will eat fish
and amphibians but
have a strong preference
for worms. They search
the ground for worms
and will burrow into the
soil to find them below
ground.
Lethaby and Gray
(2016) marked indi-
viduals at two sites in
Pennsylvania and found
Figure 29.44. Thamnophis butleri from Ontario, Canada. maximum distance
Photograph from Joshua Vandermeulen. traveled by two males
at one location was 10
m over seven days. The
maximum distance traveled by a male at the second site was 70 m over 62 days. A female
traveled 37 m in 14 days, and at another location, a female traveled a maximum distance of
66 m in 21 days.
Reproduction. The smallest mature gravid female was 34.5 cm SVL, and a 32,1 cm SVL
male was observed courting a female. Thamnophis butleri of these sizes would be in their
second spring. Mature males have tubercles on their mental scale and chin shields. Mating

522
 29. The Thamnophiine Clade

occurs upon emergence from hibernation, and males scramble to mate with females. Multi-
ple males will follow a female’s pheromone trail and try to mate, forming a “ball of snakes”
(mating aggregation). Parturition July-September (Vogt 1981). Clutch size about 4-20, in-
creasing with female size. Sexually mature in 2nd or 3rd year (Fitch 1970). Litters of 4-20
have been reported, but most litters have 8-10 young that are 125-185 mm in total length
(Ernst and Ernst 2003).

Golden-headed Garter Snake

Thamnophis chrysocephalus (Cope, 1885)
Distribution and Habitat. Streams, ponds, and adjacent
terrestrial habitats in pine-oak forests of the southern Sierra
Madre Oriental in Guerrero, Oaxaca, Puebla, and Veracruz,
Mexico. The type locality Orizaba, Veracruz.
Figure 29.45. Distribution of Note: Taylor (1940) described Thamnophis eburatus from
Thamnophis chrysocephalus. the type locality of Cerro San Felipe, Oaxaca, Mexico. This
snake is a melanistic Tham-
nophis chrysocephalus.
The maximum length
is 692 mm. The dorsum is a
uniform black or brown. The
vertebral stripe is only on
the anterior body; the lateral
stripe is confined to scale row
2. Dorsal scale rows at mid-
body are in 17 rows. Ventrals
135-155, subcaudals 69-87.
Natural History. A poor-
ly known species. Reportedly
active day and night. Ob-
served swimming in streams
at night and several meters
Figure 29.46. Thamnophis chrysocephalus. Photograph from Mike
from a stream in the after-
Pingleton.
noon; observed foraging at
the bottom of a shallow pool (Rossman et al. 1996).

Sierra Garter Snake

Thamnophis couchii (Kennicott, 1859)
Distribution and Habitat. Streams, ponds, and reser-
voirs (permanent and seasonal) in oak woodlands, mon-
tane coniferous forests, grassy valleys, and chaparral in
California and western central Nevada at 91-1438 m ASL.
The type locality is the Pitt River, California.
Figure 29.47. Distribution of The maximum length is 957 mm. The dorsum is ol-
Thamnophis couchii. ive-brown to dark brown or black. Two alternating rows of
spots occur between the stripes. The lateral stripe is con-

523
Aquatic Snakes

fined to rows two and three.

The vertebral stripe is yellow
and may be visible on the an-
terior body but fades posteri-
orly. The lateral stripe may be
bright yellow or absent. Some
populations are melanistic.
Dorsal scale rows are 21 at
mid-body and resuced to 17
posteriorly, some individuals
have 19 at midbody. Ventrals
161-187, subcaudals are 68-
99.
Natural History. A diur-
nal snake with seasonal activ-
Figure 29.48. Thamnophis couchii
ity that varies with elevation.
Photograph from Mike Pingleton. Lowland populations may be
active from March to Decem-
ber, while higher-elevation populations are active for three or four months (Rossman et al.
1996). Morning basking in tall grass is followed by foraging. Diet changes with the habitat
of this highly aquatic snake. In seasonal aquatic habitats, it feeds on amphibians (anurans
and salamanders), and in permanent streams, it feeds on fish. Juveniles tend to ambush fish
and tadpoles in shallow water or from the edge of the shoreline, while adults crawl along the
bottom and tongue flick, actively foraging for prey (Drummond 1983, Lind 1990). If the prey
is large, it may be dragged into shallow water before the snake starts to swallow it. Litter size
varies from five to 38. Gravid females may form aggregation. Parturition occurs from late July
to September (Rossman et al. 1996; Ernst and Ernst 2003). Predators include a variety of birds
and carnivorous mammals; defense behavior involves escape, biting, and the release of musk.

Black-necked Garter Snake

Thamnophis cyrtopsis (Kennicott, 1860)
Distribution and Habitat. Southwestern USA from
southern Colorado, Utah, Arizona, Texas, and New Mexico;
in Mexico, it ranges from Aguascalientes, Chiapas, Tamau-
lipas, San Luis Potosí, Quéretaro, Oaxaca, Jalisco, Sinaloa,
Hidalgo, Guanajuato, Puebla, Guerrero, Nayarit, Morelos,
and southward into Guatemala. Type locality: Rinconada,
Coahuila, Mexico.
Figure 29.49. Distribution of This is a polytypic species with three subspecies.
Thamnophis cyrtopsis.
Thamnophis cryptopis collaris (Jan 1863) occurs from
southern Sonora, Durango, and Chihuahua, through west-
ern and southern Mexico to west central Guatemala.
Thamnophis crytopsis cyrtopsis (Kennicott 1860) ranges from southwestern Texas and
New Mexico; southward into Mexico (Coahuila, Nuevo León, Chihuahua).
Thamnophis crytopsis ocellatus (Cope 1880) ranges from southcentral Texas and the Ed-
wards Plateau west to the Big Bend. Type locality: Texas, near Helotes.
Identification. Dorsal scales in 19 rows at midbody. Upper labials eight. The black nu-

524
 29. The Thamnophiine Clade

Figure 29.50. Thamno-

phis cyrtopsis. Photo-
graph from (A) Mike
Pingleton, (B) Paul
Freed.

chal collar is not divided into two nuchal blotches. The dorsum of head is gray or brown.
The brown to black dorsolateral areas show some evidence of a checkerboard pattern (most
prominent anteriorly). The narrow vertebral stripe is distinct. The first three scale rows are
white, devoid of markings or having irregular small black markings; A prominent row of
black marks along the margin of the first scale row and ventral scales. Black marks are usually
lacking on lower labial sutures (except the most posterior) (Webb 1980).
Natural History. The diet comprises anurans, with some evidence that salamanders, liz-
ards, and small birds may be eaten, and some invertebrates (earthworms and Triops). Neo-
nates feed on small fish and worms. Foraging along the bank of a stream and ambushing prey
while they float or conceal themselves in stream bed rocks or algal mats have been reported.
Reproduction. The gametic cycle of both sexes of this species was examined by Goldberg
(1998). In spermiogenesis, metamorphosing spermatids and mature sperm were present.
Sperm were present in the vasa deferentia during all months (March-November) indicating
T. cyrtopsis has the potential of breeding throughout the year. Mating occurs the following
spring utilizing sperm stored overwinter in the vasa deferentia. Some mating likely occurs
during autumn. Mating coincides with hypertrophy of the kidney’s sexual segment.

525
Aquatic Snakes

The smallest reproductive female measured by Goldberg (1998) was 412 mm SVL. Mean
clutch sizes for 25 females averaged 8.4 (3-24 range). A significant positive correlation existed
between female body size and enlarged follicles (six mm, or embryos). Litter sizes from Ari-
zona and Texas ranged from 6-22 and were born in July and August (Goldberg 1998).
In prenuptial secondary vitellogenesis, the female’s reproductive cycle is like that of oth-
er Thamnophis species. Females enter hibernation with small, previtellogenic follicles. Yolk
deposition occurs in spring. Females with enlarged follicles six mm in diameter or developing
embryos were found March-June. The presence of some reproductively inactive females in
March-June suggests that not all mature females produce litters each year. About 61% of fe-
males from March-June produced a litter. It is unknown if the March and June females under-
going yolk deposition would have been produced. Data on reproductive frequency for other
species of Thamnophis suggests most females reproduce each year, although some females
may not breed yearly (Rossman et al. 1996). This appears to be the case for T. cyrtopsis fe-
males. Neonates were found from June to August. The date of birth, size of neonates at birth,
and growth rates likely show geographic and yearly variation (Goldberg 1998). Neonates are
145-281 mm in total length.

Western Terrestrial Garter Snake

Thamnophis elegans (Baird and Girard, 1853)
Distribution and Habitat. Many habitats, including coastal
marine habitats, are used but usually close to water. The species
ranges from southwest Canada into Washington state, south-
ward to Oregon, eastward to Montana, Colorado, and Nebras-
ka, and southward to Baja California Norte, Mexico. A polytyp-
ic species with at least three subspecies.
Figure 29.51. Distribution of
Other subspecies have been described, but Hallas et al.
Thamnophis elegans.
(2021) and previous authors found morphological and genetic
evidence that cast doubt on the validity of those taxa. Hallas et
al. (2021) used molecular data and found Thamnophis elegans is composed of distinct evo-
lutionary lineages, each with its geographic range and history of diversification. In addition,
Thamnophis e. elegans and T. e. terrestris show unique patterns of diversification as popula-
tions dispersed from east to west.
Thamnophis elegans elegans (Baird and Girard. 1853) ranges from central California and
southwest Oregon. The type locality is El Dorado County, California.
Thamnophis elegans terrestris (Fox 1951) ranges from extreme southwest Oregon to San-
ta Barbara County, California, west of the Central Valley. The type locality is Strawberry
Canyon, Berkeley, Alameda County, California.
Thamnophis elegans vagrans (Baird and Girard. 1853) is widespread in the western USA
(Washington, Oregon, Nevada, NW New Mexico), W Canada (British Columbia, Saskatch-
ewan, Alberta). Type locality “California” in error. Likely Santa Fe, Santa Fe County, New
Mexico [see Maslin 1958 and Maslin et al. 1959].
Identification. Maximum size 1090 mm. A highly variable species. A vertebral stripe is
usually present and can be white, red, orange, or tan but is absent in some subspecies. On
scale rows two and there is a dorsal lateral stripe that can be off-white or red. Dorsal scale
keeled (except the first row) in 17-21 mid body, and posterior rows are reduced to 15-17.
Ventral scales in males 148-185, subcaudals in males 71-101; in females ventrals are 137-177;
subcaudals are 61-91. Head scales. Upper labials eight, 4-5 border the orbit; one preocular

526
 29. The Thamnophiine Clade

Figure 29.52. Thamnophis elegans (A) T. e. elegans, Plumas County, CA (B) T. e. terrestris, San Mateo
County, CA, (c) T. e. terrestris, San Mateo County, CA, Brian Hubbs; (D) T. e. vagrans from Taos
County, NM. James N. Stuart; (E) from Coconino County, AZ Robert Dobbs (F) A juvenile T. e.
vagrans from Boulder County, CO, photo by aiknight13.
sometimes two; usually three postoculars; posterior chin shields little may be longer than
anterior pair.
Natural History. The diet is exceptionally diverse and includes gastropods (slugs), an-
nelids (worms and leeches), mollusks (snails), fishes, anurans (tadpoles, frogs), small birds,
and mammals. It also will scavenge on carrion. Foraging is usually diurnal and may occur in
water and on land. Fitch (1940) examined 69 T. e. vagrans that contained prey: 37 specimens
had aquatic prey, while 32 had terrestrial prey. Amphibians are preyed upon in the high Sierra
Nevada, California (Jennings et al. 1992), and it is also known to feed on ictalurid fish (Gimbel
and Zahratka 2021) and rodents (voles) Blais et al. 2021. Prey taken by this snake is subdued

527
Aquatic Snakes

with venom that has proteolytic and myonecrotic effects (Jansen 1987, Finley et al. 1994).
Reproduction is similar to other temperate Thamnophis. However, the timing of the ga-
metic cycles and emergence from hibernation, mating, and parturition are influenced by the
latitude and altitudes of the population (Fox 1952, 1982). Males mature at 370-420 mm total
length; females mature at 420-460 mm. Neonates are 167-276 mm and average 21.5 cm (Ernst
and Ernst 2003). Harmer (2019) observed the sea anemone Anthopleura sola consuming a
small T. e. terrestris in a tidal pool. Bowley et al. (2020) reported T. e. vagrans using thermal
hot springs in Yellowstone National Park; the water had a pH of 2.73 and a temperature of
36.8°C. Bowley et al. discuss the neutral nature of keratin and explain how it may give the
snake resistance to acid.

Mexican Garter Snake

Thamnophis eques (Reuss, 1834)
Distribution and Habitat. Permanent water with emergent vegetation in southern Arizo-
na and southwest New Mexico in the USA; in Mexico, it ranges from the vicinity of the Pico de
Orizaba northwestward to and south to Aguascalientes, Sonora, and Chihuahua, Quéretaro.
Rossman et al. (1996) showed disjunct colonies in Oaxaca and Nuevo León, Hidalgo, San Luis
Potosí, Guanajuato, Puebla, Guerrero, Nayarit, and Morelos, Mexico. It has been associated
with stock tanks, cienegas, and streams flowing from them; they also occur along large rivers,
shallow wetlands, and wet meadows. This polytypic species has ten subspecies; these have not
yet been evaluated using molecular data. The maximum size for this snake is 1240 mm in total
length (Suarez-Varon et al. 2016).
Thamnophis eques carmenensis Conant 2003 is distributed from Tlaxcala, La Lagunilla
del Carmen, Mexico. The type locality is near El Carmen, Tlaxcala, Mexico.
Thamnophis eques cuitzeoensis Conant 2003 occurs in Michoacán, Mexico. The type
locality is 100 m east of the southern end of the south–north causeway across the Lago de
Cuitzeo, Michoacán, Mexico.
Thamnophis eques diluvialis Conant 2003 is known from Jalisco, Cajititlán, and several
isolated localities. The type locality is near Villa Corona at the north end of the Laguna de
Atotonilco, Jalisco, Mexico.
Thamnophis eques eques (Reuss, 1834) ranges along the southern edge of the altiplano
and in the Sierra Madre del Sur in the states of Nayarit, Jalisco, Michoacan, Mexico Distrito
Federal, Puebla, Veracruz, and Oaxaca.
Thamnophis eques insperatus Conant 2003 is known from Michoacán, Mexico. The type
locality is on a highway 6 km southeast of Zacapu, Michoacán, Mexico.
Thamnophis eques megalops (Kennicott 1860) is known from Arizona and New Mexico in
the USA and ranges southward into Mexico along the Sierra Madre Occidental in the Chihua-
huan Desert and north of the Mexican altiplano in the Mexican states of Sonora, Chihuahua,
Durango, Guanajuato, Hidalgo, and SanLuis Potosi.
Thamnophis eques obscurus Conant 2003 is known from Jalisco, Mexico. The type locality
is in the town of Chapala, Jalisco, Mexico.
Thamnophis eques patzcuaroensis Conant 2003 is known
from Michoacán, Mexico. The type locality is the lakeshore
north of Pátzcuaro, Michoacán, Mexico.
Thamnophis eques scotti Conant 2003 occurs in Jalisco,
Figure 29.52. Distribution of Mexico. The type locality is near Magdalena, Lago de Magdale-
Thamnophis eques. na, Jalisco, Mexico.

528
 29. The Thamnophiine Clade

Figure 29.53. Thamnophis eques (A) Zacapu Garter Snake (Thamnophis eques insperatus), Photograph
from Ricardo Arredondo T. (B) Northern Mexican Garter Snake, T. e. megalops from Yavapai County.
Photograph from Scott Sprague. (D) Lake Chapala Garter Snakes, T. e. obscura Photograph from J.
Carlos Cuevas. (E) Magdalena Garter Snake, T. e. scotti from Teuchitlán, Jalisco, MX, Photograph
from Julio Alejandro Alvarez Ruiz. (F) Blue-Striped Mexican Garter Snake, Thamnophis eques virgat-
enius from Pueblo Nuevo, Durango, MX. Photograph from Heriberto Avila G. (G) Thamnophis eques
patzcuaroensis from Pátzcuaro, Michoacán, Mexico. Photograph from R. Enrique Llanos-Romero.

529
Aquatic Snakes

Thamnophis eques virgatenuis Conant 1963 occurs in the highlands of Durango and Chi-
huahua, Mexico.
Natural History. The diet comprises fish and amphibians, with juveniles taking worms.
Adults occasionally consume birds and mammals (Rossman et al. 1996). While some studies
have suggested this threatened taxon requires small, soft-rayed native fishes, Emmons et al.
(2016) found that it will prey upon larger, invasive prey such as Bullfrogs, Largemouth Bass,
and Bullheads. In Arizona, Rosen and Schwalbe (1988) found females do not show follicle en-
largement until the spring, which is unusual for Thamnophis, which often indicates follicular
growth in the fall. They reported an average litter size of 13.6, noted that neonates are 13.7-
19.4 cm in SVL, and that the introduced Bullfrogs prey on this species. Bourne et al. (2021)
reported predation by the invasive crayfish Oronectes virilis.
Conservation. Wood et al. (2018) used genome-wide nuclear markers to study the pop-
ulation structure and genetic diversity of Thamnophis eques megalops. They found that pop-
ulations along the lower Colorado River Basin are highly differentiated from sister species
and populations located further south in Mexico. Colorado River populations exhibit reduced
genetic diversity relative to populations along the Sierra Madre Occidental. Within the lower
Colorado River Basin, genetic analyses revealed highly structured genetic groups that point
to shared contemporary and historical drivers of differentiation. They found most sites in the
lower Colorado River Basin have low genetic diversity and effective population sizes below the
threshold required to retain adaptive potential.

Mexican Wandering Garter Snake

Thamnophis errans Smith, 1942
Distribution and Habitat. Pine-oak forests in Chihuahua, Du-
rango, Jalisco, Nayarit, Mexico. The type locality is Colonia Garcia,
Chihuahua, Mexico.
Dorsal scale rows 19 at midbody. Ventrals 155 to 166; subcau-
Figure 29.54. Distribution of dals 72 to 91; upper labials seven or eight; sixth labial large and
Thamnophis errans not narrowed above; vertebral and dorsolateral stripes poorly
defined, vertebral stripe covering
one scale row where visible; spots
between stripes small, those of
the outer row usually very poor-
ly de­fined or absent; top of head
dark; upper labial region pale,
except for black posterior edges
on most of the labials (modified
from Smith 1942). Addition-
ally, it has a black tongue (Fitch
1980), while other members of
the elegans-ordinoides group
have red tongues.
Natural History. Poorly
Figure 29.56. Mexican Wandering Garter Snake, Thamnophis known. Fitch (1985) reported
errans from Mezquital, Durango, Mexico. Photograph by Chris the average liter was 7.8, rang-
Grünwald. ing from 6 to 10.

530
Aquatic Snakes

Exiled Garter Snake

Thamnophis exsul Rossman, 1969

Distribution and Habitat. Dry pine forest at higher

Figure 29.55. Distribution of elevations in Tamaulipas, Nuevo León, Coahuila, Mexico.
Thamnophis exsul
The type locality is a relatively dry pine forest, 11 mi. E, 3.5
mi S San Antonio de las Alazanas, Coahuila, at about 9,100 feet elevation.
The smallest known species in the genus, having a maximum recorded length of 460
mm. Dorsum almost uniform gray-brown with three indistinct pale stripes or having three
or four alternating rows of spots or blotches with the light stripes partially or entirely sup-
pressed. Specimens from Nuevo Leon show a marked ontogenetic pattern variation. Juveniles
have three or four rows of prominent spots. The vertebral stripe is absent or faint and some-
times interrupted at intervals by dorsal spotting. Dorsal scales in 17 rows with no posterior
reduction; upper labials seven, lower labials eight; ventrals 142-150 ventrals; subcaudals 52-
56. The subadult spotted condition is not as prominent as in juveniles and is barely evident in

Figure 29.56. Thamnophis exsul from the Sierra de Pena Nevada, Nuevo Leon, Mexico.
Photograph from Alan de Queiroz.

larger adults (modified from Liner 1992). Natural History. This garter snake may be entirely
terrestrial. Unknown.

531
Aquatic Snakes

Mesoamerican Highlands Garter Snake

Thamnophis fulvus (Bocourt, 1893)
Distribution and Habitat. Endemic to the highlands of
central Chiapas, Mexico, southern Guatemala, southwest Hon-
duras, and adjacent El Salvador, at 1,400 to 3,700 m. It is known
from the montane forest: including broadleaf forests (rain and
cloud forest) and mixed and coniferous forests (pine-oak and
Figure 29.57. Distribution of pine forest), and páramo grassland with opine pine woodland
Thamnophis fulvus. within 150 m (Johnson et al. 2010; Acevedo et al. 2010; Eiser-
mann et al. 2016). The type locality is Alta Verapaz, Guatemala.
Adults reach at least 62.5 cm. Dorsum olive-brown dorsum and two alternating rows of
black spots that may increase in size
on the forebody and intrude on the
vertebral stripe. The vertebral stripe
is tan, and the lateral stripe is pale
yellow. The ventrals may have two
rows of small spots laterally, have an
indistinct mid-ventral stripe, or be
unmarked. Some specimens have
black blotches or vermiculations on
the frontal and parietal scales. The
tongue is black. Dorsal scale rows
are 19 or 21 at midbody. Ventrals
132-154; subcaudals 56-80 (based
on Rossman et al. 1996).

Natural History. Frequently

found near water. Dietary records
were two hylid frogs and tadpoles.
Reproduction is poorly known;
females are known to be gravid in
May and October. One female gave
birth to a single dead neonate 39
months after it had access to a male
(Rossman et al. 1996).

Giant Garter Snake

Figure 29.58. Thamnophis fulvus
from Francisco Morazan, Distrito
Central, Honduras. Photograph
from (a) Josue Ramos (b) Juan
Rabasa (C) San Cristóbal de las Ca-
sas, Chiapas, MX Photograph from
Eric Eberman.

532
 29. The Thamnophiine Clade

Thamnophis gigas Fitch, 1940

Distribution and Habitat. Freshwater marshes, streams, and wetlands with soft bottoms
and emergent vegetation; and their adjacent associated upland
habitats throughout the length of the Sacramento and San Joa-
quin valleys in Central California. Only about five percent of
its historical wetland/upland habitat acreage remains today. The
type locality is Gadwall, Merced County, California. Elevation-
al distribution ranges from sea level to 122 m.
Figure 29.59. Distribution of Dorsum brown to olive with a cream, yellow, or orange
Thamnophis gigas. vertebral stripe and two pale lateral stripes. Some individuals
may have a checkered pattern of black spots between the dorsal
and lateral stripes. Background coloration, prominence of the checkered pattern, and the
three yellow stripes are individually and geographically variable (Hansen 1980). Individuals
in the northern Sacramento Valley tend to be darker, with more pronounced vertebral and
lateral stripes. The ventral coloration can be cream, orange, olive brown, or pale blue with or
without ventral markings. Upper labial scales are dull brown and usually lack distinct wedge
markings.
Dorsal scales are in 23 or 21 rows; females have 23 rows more frequently than males.; the

Figure 29.60. Thamnophis gigas from Glen County, California US.

Photograph from Brian Hubbs.

scale rows are reduced posteriorly to 17. Ventrals 162-168 in males and 156-164 in females;
subcaudals range from 73-81 in males and 65-73 in females. On the head, one preocular,
three postoculars; upper labials eight, lower labials 10 or 11.
Size. The Giant Garter Snake reaches a total length of 162.6 cm. Neonates 15.2-23.2 cm
in SVL
Natural History. Diet. Anuran prey is taken more often than fish. Native treefrogs and

533
Aquatic Snakes

the introduced Bullfrog are eaten most often, and the fish captured are usually cyprinids,
including Common Carp, Fathead Minnows, Red Shiner, and Golden Shiner. Giant Gar-
tersnakes selected native frogs over any other prey type and selected all frogs, native and
introduced, more than introduced fishes. Even though invasive prey is dominant in their
environment, given a choice, Giant Gartersnakes prefer to eat the less abundant native tree
frogs. Thus, the management of native prey communities that also limits the abundance and
distribution of invasive prey will preserve historical trophic dynamics and will likely benefit
Giant Gartersnakes and other native predators of the Central Valley (Ersan et al. 2020). This
snake has been reported to eat the rodent Microtus californicus (Fulton and Munoz 2018).
Reproduction. Litter sizes range from 10 to 46, with a mean of 23.1 (Hansen and Hansen
1990). Rose et al. (2018) found average litter size from X-rays was 15.9, and captive-born
litters averaged 15.5; but captive-born litters had high stillbirth rates; of 202 neonates born,
154 survived. Fecundity is positively related to maternal snout-vent length, and there is some
evidence that larger litters contained smaller neonates. The proportion of gravid X-rayed
females was 0.50 in 2014, 0.47 in 2015, and 0.64 in 2016. Central California experienced an
exceptional drought from 2012–2015, which may have affected the reproductive output.
The Giant Garter Snake was listed as endangered on December 27, 1991. However, the
Fish & Wildlife Service reevaluated the status of the snake before adopting the final listing
rule, and it was listed as a threatened species on October 20, 1993.
The loss and subsequent habitat fragmentation are the primary threat to the Giant Gar-
tersnake throughout the Central Valley of California. Habitat loss has occurred from urban
expansion, agricultural conversion, and flood control. Habitat fragmentation has ultimately
resulted in the snake being extirpated from the southern one-third of its range in the San
Joaquin Valley. The Giant Garter Snake also faces other threats: traffic, climate change, and
predation by non-native species. Climate change has been linked to the intensity of weather
events (heat waves, droughts, and storms) that may cause mass mortality by affecting habitat
or ecosystem characteristics.

Godman’s Garter Snake

Thamnophis godmani (Gunther, 1894)
Distribution and Habitat. Pine-oak Forest and cloud
forest in the Sierra Madre del Sur of Guerrero. Record-
ed from elevations of 1768-2438 m. A record from Aca-
huizotla at 853 m is suspect because a habitat suitable for
this species would unlikely occur at such a low elevation.
The type locality is Mexico, Guerrero, Omilteme, and Am-
ula (Rossman and Burbrink, 2005).
Figure 29.61 Distribution of Tham- Dorsal scale rows 17 at midbody. Top of the head
nophis godmani. un-patterned. Two rows of small black spots between
light vertebral and lateral stripes. Ventrals average 144 in
males and 138 in females; subcaudals average 79 in males, and 71 in females (Rossman and
Burbrink, 2005).
This species was initially composed of four distinct populations; Rossman and Burbrink
(2005) described each population as a different species (T. bogerti, T. conanti, T. lineri) based
on morphology. Little is known about their natural histories. Rossman et al. (1995) report
observations by Caldwell and Rossman, who observed them under logs, tar paper, oak-pine
forests, and swimming in streams. One specimen disgorged a mouse when captured.

534
 29. The Thamnophiine Clade

Figure 29.62. Thamnophis godmani is from Chilpancingo de los Bravo, Guerrero, MX.
Photograph from Edgar Salmeron Barrera.
Two-Striped Garter Snake
Thamnophis hammondii (Kennicott, 1860)
Distribution and Habitat. Streams,
creeks, and pools in oak woodlands and
mixed oak chaparral and sparse pines,
cedars, and scrub oak from Monterey
County, California, southward into Baja
California, Mexico. It has a population
on Catalina Island and an isolated pop-
ulation in Baja California Sur. Its eleva-
tional distribution is from sea level to
2130 m.

Figure 29.63. Distribution of

Thamnophis hammondii.

Figure 29.64. Thamnophis hammondii. (A)

Photograph from Mike Pingleton. (B)
From Los Angles Co., CA. Photograph
from Kaiyuan Chen. (C) From Santa Mon-
ica Mountains National Recreation Area,
Los Angeles County, CA. Photograph from
Brad Alexander.

535
Aquatic Snakes

Natural History. Because this species was long confused with Thamnophis elegans and T.
couchii, its natural history is poorly known. The diet is composed of fish and amphibians in
all their life stages. They may also feed on annelid worms. Foraging occurs day and night, in
the water and on land. Sexual maturity of males occurs at 37.3 cm SVL, and females mature at
38.8 cm SVL (Wright and Wright, 1957). Mating occurs in the spring (April-May), and par-
turition occurs from July to October. Litter sizes of 3-36 (average 15-16) have been reported,
and neonates are 20.0-22.0 cm in total length (Cunningham 1959, Rossi and Rossi 1995).

Checkered Garter Snake

Thamnophis marcianus (Baird and Girard, 1853)
Distribution and Habitat. Deserts and dry grasslands to
premontane forests (Abarca and Solórzano 2020) in Costa
Rica. It ranges from southern Kansas to California and south-
ward into Mexico, Belize, Guatemala, possibly El Salvador,
Honduras, Nicaragua, and Costa Rica. Elevational ranges ex-
tend from near sea level to 2200 m. This is a polytypic species
with three or four subspecies. The type locality is Indian Ter-
ritory between Camp 5 and Red River (Tillman County near
Figure 29.64. Distribution of Hollister, Oklahoma, fide H. A. Dundee, cited in Cochran
Thamnophis marcianus.
1961: 182. Taylor (1949) restricted the type locality to Red
River, Arkansas. Originally the type locality in-
cluded New Braunfels, Sand Pedro, and Indiano-
la, Texas. Thamnophis marcianus is a polytypic
species.
Thamnophis marcianus bovalli Dunn 1940
occurs in Nicaragua, northern Costa Rica. The
type locality is Granada, Nicaragua. Dunn di-
agnosed it as a Thamnophis without stripes; but
with equidistant black lateral bars alternating
with black dorsal spots anteriorly. The ventrals
are black-bordered anteriorly; one preocular;
140-144 ventrals; tail about 25% of total length.
Thamnophis marcianus marcianus (Baird &
Girard, 1853) ranges southward from Texas, New
Mexico, and Oklahoma to Chihuahua, Mexico.
Type locality: Indian Territory between Camp 5
and Red River (Tillman County near Hollister,
Oklahoma). Note that Taylor 1949 restricted the
type locality to Red River, Arkansas. It originally
included New Braunfels, Sand Pedro, and Indi-
anola, Texas.
Thamnophis marcianus praeocularis ranges
Figure 29.65. Thamnophis marcianus
(Baird & Girard, 1853). Photograph from
(A) Mike Pingleton (B-C) JCM.

536
 29. The Thamnophiine Clade

southward from Mexico Yucatan, Quintana Roo, Mexico to Belize. Type locality Belize. A
broad vertebral stripe with two rows of small spots on each ventral has the fewest ventrals and
the greatest number of maxillary teeth of any marcianus population (Rossman 1971).
Thamnophis marcianus ruthveni Hartweg and Oliver 1938. From Oaxaca, Mexico. The
type locality is three miles northeast of Tehuantepec, Oaxaca, Mexico. The tan vertebral stripe
is restricted to the vertebral row. The ground color is brown.
Natural History. The species can be diurnal, crepuscular, and nocturnal activity may
change with the temperature. Seasonal activity extends from February to December in Tex-
as, northern Mexico. Central American populations are likely to be active year-round. Diet
mainly comprises amphibians, but fish, mammals, birds, and various invertebrates. Gati-
ca-Colima and Hernandez-Alvarez (2019) found a Chihuahua specimen that had eaten a
Fundalus zebrinus, an invasive species in Mexico.

Blackbelly Garter Snake

Thamnophis melanogaster (Peters, 1864)
Distribution and Habitat. Streams, lakes, and marshes in Mexico (Distrito Federal,
Durango, Guanajuato, Jalisco, Michoacán, Nayarit,
Querétaro, Zacatecas, San Luis Potosí, Aguascalientes,
Sonora, Hidalgo). Rossman et al. (1996) consider this
a polytypic species with four indefinable subspecies.
Thamnophis melanogaster canescens Smith 1942 Mexi-
co (Durango, Jalisco), the type locality is Chapala, Lake
Chapala, Jalisco. For a detailed analysis of this snake’s
Figure 29.66. Distribution of color pattern, see Gregory et al. (1983). The coloration
Thamnophis melanogaster.
and pattern are polymorphic.

Figure 29.67. Tham-

nophis melanogas-
ter. (A) Thamnophis
melanogaster ca-
nescens from Jocote-
pec, Jalisco, MX.
Photograph from
Emmanuel Guevara
Lazcano. (C) Tham-
nophis melanogaster
chihuahuaensis from
Madera, Chihuahua,
MX. Photograph
from Javier Cruz Ni-
eto (D) From Jalisco,
MX. Photograph
from Iván Ahuma-
da/[Link] .

537
Aquatic Snakes

Thamnophis melanogaster chihuahuaensis Tanner 1959. Mexico (Chihuahua, Durango).

This subspecies lacks the black ventral surface of the other subspecies.
Thamnophis melanogaster linearis Smith, Nixon and Smith 1950. The type locality is
Lerma, México.
Thamnophis melanogaster melanogaster (Peters 1864). Type locality Mexico.
Natural History. A diurnal snake, much of the year. It transitions to nocturnal behavior
in midsummer. Conant (1963) observed basking behavior in the late morning during the
winter. Diet. Prey reported as fish (57%), leeches (20%), tadpoles (16%), and lesser numbers
of earthworms, crabs, and adult frogs (Drummond 1983). Thamnophis melanogaster mainly
forages at temperatures above 20C (Manjarrez and Drummond 1996). Foraging behavior
includes swimming on the surface and diving to catch prey. Underwater searching involves
slowly swimming with the mouth closed; chemosensory cues do not seem as essential as
visual cues in this species (Drummond 1983, 1985). Reproduction is initiated by females
releasing pheromones to attract males. Females from Lake Cuitzeo, Michoacán, gave birth
in June and July, the average clutch size was 12.9, and the number of neonates was related to
female size (Ford and Ball 1977).
Adelophis are most closely related to Thamnophis melanogaster and are discussed here.
The phylogenetic placement of Adelophis within New World natricines has always been prob-
lematic. The generic name means “uncertain snake,” a reference to its unclear phylogenetic
affinities. Adelophis superficially resemble other semi-fossorial natricines, such as Storeria
and Tropidoclonion, and have historically been placed in these genera. They possess unique
scale counts and specialized osteological traits (Hallas et al. 2022). I am following Hallas et
al. (2022) and recognizing them as members of Thamnophis.

Cope’s Mountain Meadow Snake

Thamnophis copei (Dugès in Cope, 1879)

Figure 29.69. Distribution of Tham-

nophis copei.

Figure 29.70. Thamnophis copei Dugès,

1879 from Tecolotlán, Jalisco, MX.
Photograph from Iván Ahumada/
[Link] .

Distribution and Habitat. Gua-

najuato, Jalisco, and Morelos Mexi-
co. Type locality: Guanajuato, Gua-
dalajara, Mexico.
A small garter snake reaches
538
 29. The Thamnophiine Clade

39.2 cm. Dorsal scales in 15 rows; the absence of a loreal; ventrals 132-138; subcaudals; back-
ground ground color. Modified from Rossman and Wallach (1987).

Fox’s Mountain Meadow Snake

Thamnophis foxi (Rossman and Blaney, 1968)
Distribution and Habitat. Known only from the type locali-
ty in southwestern Durango, Mexico, it occurs in a damp moun-
tain meadow surrounded by pine forest.
Dorsal scales in 17 rows with no reduction; a loreal pres-
ent; 132-139 ventrals; subcaudals 48-57; dorsal pattern with one
pale stripe and four dark stripes on a gray-brown or olive-brown
Figure 29.71. Distribution of ground color; 20-21 maxillary teeth. Modified from Rossman
Thamnophis foxi.
and Wallach (1987).

Tamaulipan Montane Garter Snake

Thamnophis mendax Walker, 1955
Distribution and Habitat. Oak-pine forests in Tamaulipas, Mexico. Type locality near La
Joya de Salas, Tamaulipas, at about 6,000 ft ASL. Known speci-
mens were taken along the mountain trail (1050-2110 m ASL)
between Gomez Farias and La Joya de Salas, at various points
between Rancho del Cielo and La Joya, and Valle de la Gruta,
about 1.5 miles northwest of “Rancho del Cielo.
The largest of seven specimens was 71.0 cm in total length.
Figure 29.72. Distribution of A narrow mid-dorsal stripe bordered by a single row of black-
Thamnophis mendax. edged blotches on each side, the lower margins of blotches usu-
ally involving the lowermost body scale row or the outer edges
of the ventrals; no lateral stripes; dorsal scales in 17 at midbody,

Figure 29.73. Thamnophis mendax from Fagáceas del Noreste de México, Gómez Farías, MX. photo by
Mario Álberto Álvarez Lara courtesy of Elí García Padilla.

539
Aquatic Snakes

reduced to 15 rows posteriorly; a low number of ventrals, males have 145-149 females have
141-145; subcaudals in males 68-69 and in females 57-61 (based on Walker 1955).
Natural History. The stomachs of two specimens contained remains of salamanders of
the genus Pseudoeurycea. This is likely to be a primarily terrestrial garter snake.

Southern Durango Spotted Garter Snake

Thamnophis nigronuchalis Thompson, 1957
Distribution and Habitat. Southwest Durango and Nayarit, Mexico. The type locality
is San Luis, about 2743 m elevation Durango, Mexico. The
Nayarit record is based on a single specimen (CAS 169660)
collected from “along Arroyo Santa Rosa, west of Santa
Teresa, Mesa del Nayar, Municipio de El Nayar” by D. B.
Breedlove on 5 October 1979 at an elevation of 2,095 m.
The habitat is described as a “rocky ridge with Pinus, Quer-
cus & Arbutus dissected by a grassy spring-filled valley.”
(Woolrich-Piña et al. 2016).
Figure 29.74. Distribution of A single-stripe garter snake, medium-sized (to 57.4
Thamnophis nigronuchalis. cm), with a stocky build, closely related to T. rufipunctatus
and T. unilabials (Wood et al. 2011). Dorsal scales in 21-
19-17 scale rows; 149-165 ventrals;
63-70 subcaudals; an ovate head;
two upper labials entering the or-
bit; two preoculars; a distinct and
undivided black nuchal blotch;
a dorsal color pattern consisting
of 5-10 rows of red-brown spots
outlined with black; no dorsal or
lateral stripes; heavily pigmented
ventrals, the pigment tending to
form an irregular black longitudi-
nal stripe.
Natural History. Tanner
(1986) suggested this snake is
confined to high-elevation basins
where it uses deep pools. Rossman
observed it swimming on the sur-
face and lying on the bottom of a
mountain stream and observed
them basking near the stream on
cool days and concealed under
logs on the bank.

Figure 29.75. Thamnophis nigronuchalis (A) Durango, MX.

Photograph from Alan de Queiroz; (B) a juvenile from Du-
rango, MX. Photograph from Eric Centenero-Alcala.

540
 29. The Thamnophiine Clade

Northwestern Garter Snake

Thamnophis ordinoides (Baird and Girard, 1852)
Distribution and Habitat. Most terrestrial snakes use edge
habitats (Dixon-MacCallum et al. 2017) from southwest Brit-
ish Columbia in Canada, western Washington, western Oregon,
and northwestern California. Alaska has an isolated population
(Neuman-Lee et al. 2011). The type locality is Puget Sound. Its
Figure 29.76. Distribution elevational distribution extends to 5000 feet (1500 meters) (Stew-
of Thamnophis ordinoides. art, 1965). The northern part of the range in British Columbia
has not been well defined. The
range extends east of the Cascade
Mountains in southern Washing-
ton (Slater, 1963).
It is a polytypic species with
three subspecies:
Thamnophis o. leptocephali
(Baird and Girard 1853) Wash-
ington, USA, and a type locality
on Puget Sound.
Thamnophis o. ordinoides
(Baird and Girard 1853), the type
locality is Puget Sound.
Thamnophis o. rubristria-
ta Meek In Meek & Elliot 1899.
The type locality was restricted
to Port Angeles, Clallam Coun-
ty, Washington, by Marx (1958),
with a type locality in the Olym-
pic Mountains.
A medium-sized species
reaching a total length 96.5 cm.
Dorsal scale rows are usually 17
at midbody but occasionally 19.
There typically are seven upper
labials and eight lower labials.
Figure 29.78. Thamnophis ordinoides (A) Lake County, Ore- Ventrals 134-162, subcaudals 49-
gon. Photograph from Claire Goodfellow (B) Washington 82. Color and color patterns vary
County, Oregon. Photograph from Sabine Baer. substantially. Dorsal color is usu-
ally some shade of brown, olive,
gray, or black. A yellow, orange, red, or blue middorsal stripe usually is present, and a yellow-
ish or white lateral stripe sometimes is present. Dark spots and light fecks may occur between
the stripes. The ventral surface is usually paler than the dorsal color but maybe suffused with
dark pigment or red.
Natural History. On the Sausalito Peninsula of California, this species inhabits eroded
canyons with scrub oaks and groves of laurels, manzanita, and willows. Frogs were uncom-
mon, while salamanders were abundant, but Thompson (1951) found they feed almost exclu-
sively on slugs of the genus Ariolimax. Gregory (1978) reported the principal food of this spe-

541
Aquatic Snakes

cies on Vancouver Island to be slugs and earthworms. In Puget Sound, snakes emerge from
hibernation in late March to April, mating occurs, ovulation occurs from June 7th to June 15,
and gestation lasts about nine weeks from June 15 to August 20. Embryos number 3-15 and
average 8.8. Fall mating occurs from September to October. Females mature at about 33.0 cm
(Hebard 1951). In Saanich, British Colombia, Dixon-MacCallum and Bell (2016) captured
54 female snakes in August and found 32 (59%) were gravid females. The average litter size
was 8.3, and the range was 2-17. Dixon-MacCallum et al. (2017) found that overall, litter
depth, canopy cover, a lack of bare ground, and woody vegetation were the essential habitat
variables for determining where snakes were located. They suggest that the edge habitat is
necessary.

Tepalcatepec Valley Gartersnake

Thamnophis postremus Smith, 1942
Distribution and Habitat. It inhabits dry tropical scrub
forests that reach 1067 m in elevation and is restricted to the
Tepalcatepec Valley of Michoacan. The type locality, EI Sabi-
no, Michoacan.
Total length 101.5 cm. The vertebral stripe is indistinct—
pale brown dorsum with small, dark alternating blotches ex-
Figure 29.77 Distribution of tending the length of the body. The first three scale rows are
Thamnophis postremus. pale, mostly unicolor, but often have scattered small black
flecks. Black upper labial markings are well developed, with
a characteristic large comma or thorn-shaped mark on the next to last suture. Seven upper
labials, sometimes eight (37%). There is often more than one black mark on the infralabial
sutures. Small, paired black spots usually occur laterally on ventrals. Ventral scales (136-151)
(based upon Webb 1980); subcaudals 71 to 72; lateral light stripes not apparent; middorsal
stripe but very little lighter than ground color, scarcely or not evident; dark spots very much
reduced, al­ternating with each other in two series on either side; mid-dorsum not traversed
by spots; the outer row of dark spots on first and second scale rows greatly reduced, generally
scarcely visible; no distinct dark spots on venter (Smith 1942).
Natural History is poorly known. However, the species appears abundant near tempo-
rary pools where frogs are breeding. A 57.6 cm SVL female gave birth to 25 neonates on 20
June. The neonates were about 13.5 cm SVL and 17.6 cm in total length (Duellman 1961).

Western Ribbon Snake

Thamnophis proximus (Say, 1823)
Distribution and Habitat. They use upland habitats near
wetlands, thick growths of grass in swamps, the margins of
pools, swamps, marshes, lakes, rivers, creeks, peat bogs, desert
springs, and roadside ditches. In addition, they will use man-
groves (salty water) and savannas in the tropics. From south-
ern Wisconsin, Indiana, and the Mississippi Valley westward
Figure 29.79 Distribution of through the Great Plains to southeastern Colorado and eastern
Thamnophis proximus. New Mexico, southward through eastern Mexico to central Cos-
ta Rica. It occurs on Mexico’s Pacific coast from the vicinity of

542
 29. The Thamnophiine Clade

Acapulco, Guerrero, to the

Isthmus of Tehuantepec in
Oaxaca (Liner and Dundee
1969; Rossman 1970). The
type locality is a stone quar-
ry on the west side of Mis-
souri, three miles above the
mouth of Boyer’s River.
This polytypic species
is based mainly on Rossman
(1963):
Thamnophis proxi-
mus diabolicus Rossman
1963 from Western Texas
and New Mexico, USA. A
subspecies of Thamnophis
proximus characterized by
an olive-gray’ to olive-brown
dorsum, an orange vertebral
stripe, a narrow dark ventro-
lateral stripe, and frequent
reduction in the width of the
lateral stripe on the posteri-
or portion of the body in the
southern part of the range.
Thamnophis proximus
orarius Rossman 1963 from
southeastern Texas and Lou-
isiana; an olive-brown dor-
sum, a broad gold vertebral
Figure 29.80. Thamnophis proximus (Say, 1823). Photograph from stripe, and a lack of a dark
(A) Mike Pingleton (B) from Union Co., IL USA. Photograph ventrolateral stripe charac-
from JCM. terize it.
Thamnophis proximus
proximus Say 1823 from northeastern Texas, Oklahoma, Arkansas, and Louisiana USA;
characterized by a black dorsum, a narrow orange vertebral stripe, and a lack of a dark
ventrolateral stripe.
Thamnophis proximus rubrilineatus Rossman 1963 from Central Texas, USA; charac-
terized by an olive-brown to olive-gray dorsum, a bright red vertebral stripe, and a narrow
dark ventrolateral stripe or none.
Thamnophis proximus rutiloris Rossman 1963 from Guerrero and Oaxaca, Mexico,
southward to Costa Rica; characterized by an olive-brown dorsum, a broad grayish-tan ver-
tebral stripe, yellow-orange labials, a narrow lateral stripe, and a moderately narrow ven-
trolateral stripe.
Adults can reach 127.0 cm. A three-striped garter snake with a long tail; dorsal scale
rows 19 at midbody reduced to 17 posteriorly. Ventrals 141 to 181, subcaudals 82 to 131.
The lateral stripe on dorsal scale rows three and four on the anterior body; labials and ven-
trals without black markings; dark ventrolateral stripe absent or narrow in most popula-

543
Aquatic Snakes

tions; parietal spots fused, brightly colored, and usually large.

Natural History. A diurnal species with activity extending from March to October in
northern populations and more southern populations are likely active year-round. During
the hottest part of the year, it may become nocturnal. The diet comprises amphibians, and
will take fish and lizards occasionally. It will actively forage frogs by probing leaf litter de-
bris or grasses with its anterior body until a frog is flushed from hiding (Wendelken 1978).
Prey taken by a Brazos County, Texas population was mostly the frogs Gastrophryne olivacea,
Pseudacris clarki, Incilius valliceps, and Lithobates sp. (Clark 1974). Adams and Childress
(2018) found a dead specimen that had attempted to feed on the toad Incilius nebulifera. It is
unknown if the snake died from asphyxiation or the toxins from the toads. Ignoring age dif-
ferences, Clark (1974) found average movements for individuals in the Texas population were
2.5 m/day for males and 1.1 m per day for females. Mating occurs in the spring (April-May),
and parturition occurs in June-July. Litter size ranges from 6-18, and neonates are 13.0-17.4
cm SVL. Beutelspacher-García (2015) described arboreal aggregation behavior that may be
associated with mating.

Yellow-throated Gartersnake
Thamnophis pulchrilatus (Cope, 1885)
Distribution and Habitat. Boreal pine-oak forests from
Durango to Tamaulipas and Nuevo Leon to Oaxaca. Elevation-
al distribution ranges from 2240 to 3000 m.
A three-striped garter snake with pale stripes. The dorso-
lateral areas are either dark brown with an indistinct checker-
board pattern of black blotches or are black and confluent with
Figure 29.81. Distribution of
Thamnophis pulchrilatus. the black nuchal collar. Pale keels on most dorsolateral scales of-
ten form thin longitudinal lines. The ventrolateral stripe is black
(paler in young) or brown-black, including a regularly spaced
series of black spots or two black dots (one above the other). There are seven upper labials.
The characteristic is a prominent black, bar-like mark between the last two upper labials.
None of the infralabial sutures (except posterior most) is black (based on Webb 1980). On the
head, one preocular, three postoculars, seven upper labials, and ten lower labials. Dorsal scale
rows are in 19 rows at midbody reduced to 17 posteriorly. Ventral scales 152-173; subcaudals
68-94.
Natural History. Díaz and Aguirre-Ramírez (2014) found a female under a rock on 28
August 2013, at 2012 h. It was 43.4 cm SVL; tail length = 10.9 cm; 11 g) at Mesa del Huarache,
Municipality of Calvillo, Aguascalientes, México. The habitat was grassland with patches of
oak forest and tropical scrub forest ecotone. When they flipped the stone, the snake tried to
escape and finally formed concentric rings, hiding the head and raising its tail in sinuous
movements while secreting musk. Following capture, the snake was palpated and regurgitat-
ed a partially digested Dryophytes eximius. Similarly, on 15 August 2013, at the same locality,
another female (SVL = 510 mm; tail length = 13.2 cm; 20 g, elevation 2380 m) was discovered
basking in the grass and displaying the same defensive behavior. On 12 June 2010, at Ciéne-
ga de Quijas, Sierra el Laurel, Aguascalientes, another specimen was found beneath a rock.
It flattened the anterior half of its body, including the head, showing the bright blue in the
interspaces between the scales for at least 30 sec, then tried to escape. A 49.4 cm total length
female gave birth to a litter of seven on 21 May from Mexico’s Distrito Federal 19.1-20.7 mm
total length and weighed 2.1-2.6 grams (Garcia-Vazquez et al. 2012). Mcvay et al. 2015 re-

544
 29. The Thamnophiine Clade

covered this snake as the sister to

Thamnophis cyrtopsis.

Plains Garter Snake

Thamnophis radix (Baird
and Girard, 1853)
Distribution and Habitat.
Wet prairies, meadows, marshes,
often near streams, vacant lots,
and open woodlands from South-
ern Manitoba, Alberta, and Sas-
katchewan, Canada, southward
into the USA - Montana, North
Dakota, South Dakota, Nebraska,
Kansas, E Wyoming, E Colorado,
NE New Mexico, Oklahoma, N
Texas, Minnesota, Iowa, Wiscon-
sin, Missouri, Illinois, Indiana,
and Arkansas. Disjunct, relictual
colonies in north-central Ohio
and on the Illinois-Missouri bor-
der. In Saskatchewan, T. radix is
restricted to grasslands and aspen
Figure 29.82. Thamnophis pulchrilatus from Jalisco, MX. Photo-
parklands in the southern por-
graph from Iván Ahumada/[Link]. tion of the province (Secoy 1976).
The type locality is Racine, Wis-
consin, limited to the “vicinity of
Chicago, Cook County, Illinois” by Smith and Brown (1946).
The maximum total length of 1095 mm. A three-striped
garter snake with a black, gray, or olive-brown dorsum. The
middorsal stripe is yellow, orange, or orange-yellow, and the
lateral stripes vary from yellowish gray to blue-cream and
involve scale rows 3-4. Two rows of dark spots usually occur
Figure 29.83. Distribution of
above the lateral stripes and another row of dark spots below
Thamnophis radix.
them. The top of the head is usually dark, with two tiny pale
spots on the parietal scales. The upper labials are yellow or
cream in coloration, with prominent dark lines along the scale edges. The ventral surface is
yellow to green, often with black spots at the outer edges of the ventral scales. Dorsal scales
are in 19 or 21 at midbody and reduced to 17 posteriorly. Ventrals 135-175, subcaudals 54-88.
Upper labials 7-8, and lower labials 9-10.
Eutainia haydenii Kennicott (1860) and Eutainia radix twiningii Couse and Yarrow 1878
have been considered subspecies of T. radix. However, given the comments of Walley et al.
(2004) I do not recognize them here.
Natural History. A well-studied, mostly terrestrial garter snake, the species is active
throughout its distribution from April to October. It is diurnal; during the hottest months, it
may shift its location to crayfish burrows. Its diet is catholic, but it does consume many earth-
worms and leeches. Ballinger et al. (1979) observed these snakes feeding on larval amphibians;

545
Aquatic Snakes

they wrote, “On one occasion,

30-40 adults were actively div-
ing and capturing salamander
larvae. Once a salamander was
caught, it was carried to shore
and eaten.
Mating in this species in-
volves scramble competition
and produces small mating
balls of snakes (this is proba-
bly true for all Thamnophis).
Litter sizes range from 3-36,
with an average clutch size of
11.6 from Boulder, Colora-
do, to 17 in the Chicagoland
area. Neonates have body sizes
of 13.1 to 13.7 cm. Tuttle and
Gregory (2014) captured grav-
id females from 10 May to 3
September at Miquelon Lake
in central Alberta, close to the
northern edge of the range.
The average size of gravid fe-
males was 62.15 cm SVL. The
litter size averaged 22.7 and
Figure 29.84. Thamnophis radix. ranged from 7 to 41. There was
Photograph from (A) Tom Anton, (B) JCM. a positive correlation between
the size of females and litter
size. They presented data consistent with the hypothesis that Miquelon Lake T. radix exhibits
a relatively ‘fast’ life history. These snakes grow quickly and reach reproductive maturity early,
like more southerly populations (Tuttle and Gregory, 2012), but they produce more signifi-
cant numbers of slightly larger young than lower-latitude snakes. Additionally, the propor-
tion of adult females that are reproductive increases with SVL. Finally, despite a lower food
intake during pregnancy (Tuttle and Gregory, 2009), their results demonstrate that annual
reproduction occurs in this northern population, although the frequency remains uncertain.

Rossman’s Garter Snake

Thamnophis rossmani Conant, 2000
Distribution and Habitat. Springs, seepages, and
ditches near the San Cayetano River flow northward to
and beyond Tepic (Conant 2000). Nayarit, Mexico. The
type locality is an open roadside ditch 1.4 km northwest
of the small settlement of San Cayetano, Nayarit, Mexico.
The maximum size is 74.5 cm (Conant 2000). The
three longitudinal stripes are pale yellow-green. Central
Figure 29.85. Distribution of stripe occupying a middorsal row of scales and adjacent
Thamnophis rossmani. halves of paravertebral rows. Lateral stripes occupying

546
 29. The Thamnophiine Clade

Figure 29.86. Thamnophis rossmani

Photograph from Ivan Ahumada /[Link].
scale rows 3-4 anteriorly and rows 2 and 3 near the tail. The color between the stripes is me-
dium brown. A black blotch above the rictus. Venter is pale gray-green, darkening slightly
toward the rear of the body. Chin and throat a uniform pale yellow. Underside of the tail and
anal scute yellow, also uniform. The anterior edge of each ventral is marked with a narrow
dark gray stripe—dorsal scale rows 19 at midbody and reduced to 17 posteriorly.
Natural History. The spring rills were inhabited by small fishes, unidenti-
fied but probably cyprinids measuring roughly 25 mm to 50 mm in length. When
the rills were not in flood, there were occasional riffles along them over which the
fishes had to flip their way. On two occasions, Conant (2000) watched individu-
als of Thamnophis rossmani lying in wait and catching fish as they crossed the riffles.

Narrowheaded Garter Snake

Thamnophis rufipunctatus (Cope, 1875)
Distribution and Habitat. Perennial streams and inter-
mittent streams with pools and riffles, and a substrate of
cobbles or boulders in east central Arizona and west central
New Mexico in the USA, and southward into Chihuahua,
Coahuila, Sonora, and Durango, Mexico. The type locality
Figure 29.87. Distribution of
is S Arizona.
Thamnophis rufipunctatus. A moderate to large-sized garter snake (to 70.0 cm total
length). No stripes are present. Dorsal color is dark brown to
olive brown, usually with six rows of dark brown or rust-col-
ored middorsal spots involving 3-5 scales extending from the nape onto the tail. The ventral
pattern is irregular dark spots or a series of dark brown cross bars, often on the anterior
547
Aquatic Snakes

edge of the ventral scales.

The head is narrow, elongat-
ed, and laterally compressed,
with one loreal, two preocu-
lars, 3-4 postoculars (the low-
er may or may not contact the
4th upper labial), upper labi-
als 8-9 (usually 8), lower la-
bials 9-10 (usually l0). Dorsal
scales in 21 rows at midbody
reduced to 17 rows posterior-
ly. Ventrals range from 151-
179; subcaudals 62-88.
Natural History. At San
Francisco Hot Springs in
New Mexico Hibbits et al.
(2002) studied Thamnophis
rufipunctatus. The diet was
exclusively fish, of which a
large proportion was the in-
troduced Gambusia affinis.
They feed on native fish, in-
cluding suckers, dace, and
trout. In 1996, the San Fran-
cisco Hot Springs population
was about 7.2 adult snakes/
ha; however, they did not find
Figure 29.88. Thamnophis rufipunctatus Photograph from (A) snakes in subsequent visits.
JCM (B) Scott Sprague. Thamnophis rufipunctatus
use streams with a steep riv-
erbed slope and large rocks of uniform size. The San Francisco Hot Springs population of T.
rufipunctatus was one of the most robust in the United States, and it completely disappeared
in a decade. Christman et al. (2021) reported a Speckled Dace (Rhinichthys osculus) regurgi-
tated by a captured specimen, and the same specimen had a larval Anaxyrus microscaphus.
Hibernation for telemetered snakes in Oak Creek Canyon, Arizona, began in late Octo-
ber to late November, and snakes emerged in early to late March (Nowak 2006). Telemetered
snakes were not active outside of the hibernacula during the winter. Hibernation sites are
above the 100-year floodplain and include rock outcrops, rocky slopes, and rock piles, often
within 30-200 m from the water.
The mating occurs in the spring and early summer, and litters of 8-18 neonates are born
in July - August (Rossman et al. 1996). Males reach sexual maturity around 2.5 years, females
mature about year two, and they may reproduce every year (Rosen and Schwalbe 1988, De-
genhardt et al. 1996). Although courtship has not been described for this species, evidence
suggests mating occurs from late July into August (Degenhardt et al. 1996). Snout-vent length
for neonates ranges from 16.0 – 20.5 cm (6.3 – 8.0 in).
Narrowheaded Gartersnakes have experienced recent extirpations. At a stream in Ari-
zona, reliable snake records were known from 1992; recent surveys (before 2018) in the vi-
cinity have had mixed but mostly negative results. The colonization of the invasive Northern

548
 29. The Thamnophiine Clade

Crayfish (Orconectes virilis) may be a factor in the decline. The crustacean has been observed
in the tens of thousands in habitats that previously supported this species. The population
decrease was likely the result of crayfish predation on the neonate snakes and the native fish
prey base.

Eastern Ribbon Snake

Thamnophis saurita (Linnaeus, 1766)
Distribution and Hab-
itat. Creek-beds, streams,
lakes, wet woodlands,
swamps, and marsh areas
are found east of the Missis-
sippi from southern Ontario,
Canada, and southern Maine
southward to the Florida
Keys and the northern side of
Lake Pontchartrain in Lou-
isiana. An isolated colony
occurs in central Nova Sco-

Figure 29.89 Distribution

of Thamnophis saurita.

tia, and perhaps another in

northeastern Wisconsin. The
species is uncommon or rare
in much of the interior of the
eastern United States. Waifs
have been reported near
the Atlantis Resort on Para-
dise Island in the Bahamas
(Johnson and Sancois 2018).
Type-locality was given as

Figure 29.90. Thamnophis sau-

rita . (A) T. s. nitae from cen-
tral Florida. Photograph from
JCM. (B) Photograph from
Mike Pingleton (C) T. s. sau-
rita from se Michigan. Photo-
graph from JCM.

549
Aquatic Snakes

Carolina. The holotype is likely lost. A neotype designated by Schmidt and Conant, 1956-57
was collected 15 miles NNE Charleston, Berkeley County, South Carolina. This polytypic
species has four subspecies. The Eastern Ribbon Snake, T. s. saurita, has a brown dorsum
with three distinctive stripes, a yellow vertebral stripe, and yellow lateral stripes that alternate
with the brown dorsum. It ranges from New York to Florida, west to the Mississippi River.
Northern Ribbon Snake, T. s. septentrionalis Rossman, 1963 has a dark brown to black dor-
sum with a yellow vertebral stripe and lateral stripes. It ranges from Maine through Ontario
and Indiana. Southern Ribbon Snake, T. s. sackenii (Kennicott, 1859) has a tan or brown dor-
sum without a vertebral stripe but with yellow lateral stripes on scale rows 3-4; it ranges from
South Carolina south through Florida. The Bluestripe Ribbon Snake, T. s. nitae Rossman,
1963 has dark dorsum with no vertebral stripe but with light blue lateral stripes; on the Gulf
Coast of north-central Florida. Hallas et al. (2022) found it, and T. proximus and T. sirtalis
clade strongly supported as the sister to the remaining northern clade species, a pattern also
recovered in some other analyses.
Maximum length 101.8 cm. A two or three-striped garter snake with no black markings
on the labials or ventrals; a dark ventrolateral stripe always present and broad, dorsal scale
rows 3-4 (row 2 is occasionally involved); parietal spots frequently lacking; when present,
they are small and rarely in contact or brightly colored. A short-body (or long-tailed) Tham-
nophis is characterized by 19 scale rows at midbody, reduced to 17 rows posteriorly; upper
labials seven or eight; ventral scales 143-177, subcaudals 94-136.
Natural History. Diet. Nocturnal foraging and feeding are common, especially in the
southern part of the range and during the warm months. The diet consists of fish and adult
and larval amphibians. Older literature on this snake suggests it is insectivorous. But the evi-
dence was lacking and was likely based on secondarily ingested items.
Reproduction. Carpenter (1952) found that female ribbon snakes in southern Michigan
attain sexual maturity between two and eight years when they attain a minimum body length
of 42.1 cm; he did not present corresponding data for males. McCauley (1945) reported a
gravid female from Maryland 84.1 cm in SVL. Mating occurs in April in the northern por-
tion of the range, where emergence from hibernation occurs in late March. Mating probably
occurs earlier in the extreme southern part of the range, where no true winter hibernation
occurs. The young are usually born in August in the north, and July in the south, with con-
siderable overlap. Dates range from 2 July through 4 October. The smallest gravid female
Rossman (1963) measured had a body length of 35.0 cm.

Mexican Alpine Blotched Garter Snake

Thamnophis scalaris Cope, 1861
Distribution and Habitat. A terrestrial, montane garter snake
that uses alpine coniferous forests and grasslands in Mexico in the
Distrito Federal, Guanajuato, Jalisco, Mexico. Michoacán, Oaxa-
ca, Querétaro, Tamaulipas, San Luis Potosí, Veracruz, Morelos,
Hidalgo, and Puebla at elevations of 2103-4273 m. The type lo-
cality is Jalapa, Veracruz, Mexico.
Figure 29.91. Distribution
of Thamnophis scalaris. A small, maximum length 60.1 cm, three-striped garter
snake with the anterior stripes connected by large black-edged,
brown blotches, usually in one row on each side, but occasionally
in two alternating rows. The blotches may or may not reduce to narrow black bars posterior-
ly. The prominent nuchal blotches also are brown with black borders, and they may or may

550
 29. The Thamnophiine Clade

not be continuous with

the brown coloration of
the parietals. The ver-
tebral stripe involves at
least the adjacent mar-
gins of the paravertebral
rows. All upper labials
have the same ground
color, and there often
are short, narrow, black
wedge-shaped marks on
the upper part of some
anterior upper labial su-
Figure 29.92. Thamnophis scalaris tures. Dorsal scale rows
Photograph from Ivan Ahumada /[Link]. are in 19-19-17 rows.
The ventral scales have
irregular dusting or pep-
pering. Ventrals are 130-147; subcaudals are 50-85.
Natural History. The only prey known for this species are lizards of the genus Sceloporus
and Barisia. Two litters born in the second half of May had eight and fifteen neonates 16.8 to
18.1 cm long.

Mesa Central Blotched Garter Snake

Thamnophis scaliger (Jan, 1863)
Distribution and Habitat. Semiarid desert scrub valleys, oak and coniferous forests, and
pastures at high elevations (2500-2720 m ASL) in the Mexi-
can states of Jalisco, Aguascalientes, Michoacan, San Luis Po-
tosí, Puebla, and Guanajuato. The type locality is unknown
(Smith and Taylor 1945, Mociño-Deloya et al. 2009b).
Adult females can reach 43.9 cm; adult males reach 36.9
cm. Dorsal scales in 19 rows at midbody. A brown dorsum
and a series of black-edged dark brown blotches two scales
Figure 29.93 Distribution of long. The ventral surface is pale gray with black vermicu-
Thamnophis scaliger.
lations on the anterior edge of many ventral scales. Nu-
chal blotches are transected by the vertebral stripe. A pale
Y-shaped marking is present on the interparietal and frontal-parietal sutures. Upper labials
are cream or white with black sutures. The vertebral stripe is pale yellow. The lateral stripes
are also yellow (distinct or indistinct) and are on scale row two or two-three: seven upper
labials, nine lower labials, one preocular, and three postoculars.
Natural History. The diet consists primarily of earthworms. However, they will take larg-
er prey Mociño-Deloya et al. (2009a) reported one specimen consumed a Thamnophis scalar-
is and summarized the literature noting that they have been reported to feed on amphibians,
lizards, and rodents, and one consumed a neonate Crotalus triseratus.
Of 27 pregnant females examined by Feriche et al. (2016), 44% recently had fed. Howev-
er, females feeding during pregnancy did not produce heavier litters or larger neonates.
Feriche et al. (2016) examined the female reproductive strategy. Gravid females were
collected in the field and maintained in captivity until birth. The low percentage of gravid

551
Aquatic Snakes

Figure 29.94. Thamnophis

scaliger from Toluca, Mex-
ico, MX. Photograph from
Juan Carlos Garcia Mo-
rales.

552
Aquatic Snakes

females each year suggested they did not reproduce annually. This result probably was driven
by the seasonal constrained availability and low-energy content of earthworms, which pre-
clude females from acquiring sufficient energy to fuel more frequent reproduction. However,
females produced relatively large litter sizes, suggesting significant investment in reproduc-
tion. In addition, Feriche et al. (2016) also found that females who reproduced later benefited
most from foraging opportunities during the short rainy and high prey-availability period
and remained in better body condition.
Most females delayed reproduction until achieving a body size approximately 2.7 times
that of neonates. Also, most females matured at a body size of about 73% of the maximum
body size recorded (SVL = 439 mm), although some mature at 61% of this body size. Sug-
gesting some females delay the first reproductive event to increase body size to ensure high
reproductive output. Therefore, the range in body sizes for mature females is relatively small
(approximately 14.2 cm), which may obscure correlations between female SVL and repro-
ductive output, a typical snake pattern (Feriche et al. 2016). Parturitions occurred between
26 June and 15 August.

Common Garter Snake

Thamnophis sirtalis (Linnaeus, 1758)
Distribution and Habitat. It ranges from central Can-
ada in the Northwest Territories, Manitoba, Alberta, Sas-
katchewan, New Brunswick, Ontario, and Prince Edward
Island southward into the USA. It extends from the At-
lantic and Gulf Coasts and west to Washington, Oregon,
and California’s Pacific Coasts and southward into Mex-
ico – there is an isolated population in Chihuahua. It is
Figure 29.95 Distribution of Tham- also present in the Bahamas (Buckner and Franz 1998).
nophis sirtalis. And it maintains populations in Wood Buffalo National
Park (59°49’N, 112°W), Northwest Territories, Canada).
Wright and Wright (1957) found 154 habitats used by this species – the 20 most used were:
woods, near water, lakes, fields, meadows, swamps, borders of rivers, marshes, along streams,
gardens, roadsides, parks, clearings, margins of a ditch, margins of creeks, moist situations,
swimming in the water, in water, and low ground. To this list, I (JCM) would add some per-
sonal observations. This snake has/had populations in urban environments where they can
spend the winter in spaces around building foundations, crevices under sidewalks, and road-
ways.
Recently, Jones et al. (2023) used the Common Gartersnake, to understand the influence
of major river basins and the transition from forest to grassland within the Interior Plains on
genetic structure, they employ phylogenetic analysis, population structure analysis with spa-
tial considerations, and demographic analyses using single nucleotide polymorphism data.
Additionally, they investigate the species' historical range expansion with ecological niche
modeling and data on geographic locations and historical climate conditions.
They (Jones et al. 2023) identified four distinct phylogeographic lineages, each showing
varying degrees of connectivity. Interestingly, they observe differences in population struc-
ture patterns between genetic markers associated with sex chromosomes and those from
autosomal loci, particularly concerning the central North American lineage compared to
coastal lineages. They also found evidence supporting the existence of Pleistocene refugia
in the southeastern region, where recent secondary contact occurred during the Last Glacial

553
 29. The Thamnophiine Clade

Maximum. Furthermore, they detected signs of both northern and southern refugia in west-
ern North America.
Jones et al. found compelling evidence indicating a Pliocene origin for Thamnophis sir-
talis in the central-southeastern part of North America. This origin predates the species' rap-
id expansion across the continent, which occurred before the middle Pleistocene and was
influenced by climatic factors leading to lineage formation. They propose that major river
networks within the Mississippi watershed played a significant role in repeated westward ex-
pansion events across the Interior Plains. It's worth noting that the study supports previous
conclusions that phenotypic differences among subspecies do not reflect shared evolutionary
history.
This is a polytypic species, and the following subspecies continue to be recognized.
Thamnophis sirtalis annectens Brown 1950 ranges from east-central Texas and adjacent
Oklahoma and the Texas panhandle; SW Kansas has a disjunct population. The type locality
is a small branch of Boggy Creek, one mile east of Austin, Travis County, Texas.
Thamnophis sirtalis concinnus (Hallowell 1852) occurs in Washington and Oregon. The
type locality is in Oregon, probably in the lower Willamette Valley near Oregon City (Fitch,
1941).
Thamnophis sirtalis dorsalis (Baird and Girard 1853) ranges from the Rio Grande Valley
and north-central New Mexico southward to El Paso, west Texas, with disjunct populations
in northeast New Mexico, USA. In Mexico, there are disjunct populations in Chihuahua. The
type locality is Fort Conrad, New Mexico.
Thamnophis sirtalis fitchi Fox 1951 occurs in California, Oregon, Idaho, and Utah in the
USA. The type locality is Gray-lodge Refuge, 9 mi W of Gridley, Butte Co., California.
Thamnophis sirtalis infernalis (Blainville 1835) occurs in California. The type locality is
California.
Thamnophis sirtalis lowei Tanner 1988 occurs in Chihuahua, México; The type locality is
Yepómera, Chihuahua, Mexico.
Thamnophis sirtalis pallidulus Allen 1899 occurs from southern Quebec and the eastern
shore of James Bay to the Gulf of St. Lawrence and Nova Scotia in Canada; in the USA, it
ranges from New England to extreme northeastern Massachusetts.
Thamnophis sirtalis parietalis (Say 1823) occurs in Canada from southern Alberta and
Manitoba. It extends southward into eastern Nevada, southern Idaho, and Utah, south to
New Mexico, north Texas, and east to Iowa, Minnesota, Wisconsin, Oklahoma, and Missouri.
And it extends southward into Mexico to Chihuahua, Coahuila, and Tamaulipas. The type
locality is a stone quarry on the west side of the Missouri River, 3 miles above the mouth of
the Boyers River.
Thamnophis sirtalis pickeringii (Baird and Girard 1853) occurs in Washington, USA. The
type locality is Puget Sound, Washington.
Thamnophis sirtalis semifasciatus (Cope 1892) occurs in northeastern Illinois and ad-
jacent portions of Indiana and Wisconsin. The type locality is Aux Plaines [=Des Plaines],
Illinois.
Thamnophis sirtalis similis Rossman 1965 ranges from western Florida’s Wakulla Co. to
the Withlacoochee River of Florida.
Thamnophis sirtalis sirtalis (Linnaeus 1758) occurs in southern Canada (Ontario, Que-
bec), through the eastern USA to the Gulf of Mexico, and west to Minnesota and SE Texas;
Type locality: Canada; the neotype is from Quebec County, Quebec.
Thamnophis sirtalis tetrataenia (Cope in Yarrow 1875) California, USA. The type locality
is the Pitt [Pit] River, California.

554
Aquatic Snakes

Figure 29.96. Thamnophis sirtalis

Photograph from (A-C) Mike Pingleton; (D) Nathan Kutok; (E-F) JCM; (G) Joe Peck.

Natural History. As Rossman et al. (1996) pointed out, the extensive distribution and
plasticity of its ecological traits and life history make generalizing these aspects of its biology
nearly impossible. Its activity varies with latitude and local climates. It has an exceptional
cold tolerance. It can be active in northern Canada for five months of the year. I (JCM) once
found a young one frozen in the ice covering a pond following a few warm March days. I
chipped it out of the ice, brought it home, and released it at the capture site a month later.
At some southern locations, the species may be active year-round. While it is diurnal, it may
become active at night during warm rain. Larsen and Gregory (1989) found a snake just out
of the northern Alberta, Canada den, with a cloacal temperature of 0.5°C. Telemetered snakes

555
Aquatic Snakes

had body temperatures of 1.8 to 6.5°C (Macartney et al. 1989). Some Canadian snakes were
found to have activity ranges that were 142,000 m square while other studies report this snake’s
recapture points were within 160 m of their original capture site (Fitch 1965, Freedman and
Catling 1979).
Diet. A wide variety of aquatic and terrestrial invertebrates and vertebrates are preyed
upon. The species is positively buoyant and hunts while swimming at the surface. And it will
hunt earthworms using chemoreception to locate burrows, pushing its way into the soil to
capture them (Rossman et al. 1996). This snake will also scavenge corpses, Vanek et al. (2021)
reported a specimen from Lake Co., Illinois, feeding on a dead American Bever.
Reproduction. Mating occurs in the spring when the snakes emerge from hibernation
and in the fall when the snake aggregates at den sites. Parturition occurs from mid-summer
to early fall, but the young may be born as early as May at more southern locations. Litter siz-
es range from an average of 7.6 in Manitoba to 32.5 in Maryland. Females may not reproduce
yearly; they may extend reproduction to once every three years or more at higher latitudes.
Neonates range from 15.4 to 20.1 cm in total length.
Note: Key et al. (2019) investigated the taxonomic status of the Texas garter snake,
Thamnophis sirtalis annectens, a taxon needing conservation. They test the hypothesis that
an integrative approach to species conservation has greater utility than using only a single
data type (e.g., molecular, morphological, behavioral, or ecological). They demonstrated how
ecological niche modeling and molecular genetics could synergistically resolve taxonomic
classifications. Using a phylogenetic analysis based on molecular data, they found that some
putative T. s. annectens specimens represent a co-occurring subspecies (the Redsided Garter
Snake, Thamnophis sirtalis parietalis), which is not of conservation concern. However, both
T. s. annectens and T. s. parietalis are genetically distinctive from Thamnophis sirtalis sirtalis.
Furthermore, the ecological distinctness of T. s. annectens from T. s. parietalis suggests that
T. s. annectens is, in fact, a distinct evolutionary subunit. Only when genetic and ecological
information was included was the distinctiveness of T. s. annectens apparent.

Sumichrast’s Garter Snake

Thamnophis sumichrasti (Cope, 1866)
Distribution and Habitat. Humid montane forest with
small streams in the Mexican states of Oaxaca, Chiapas, Que-
retaro, San Luis Potosí, Tabasco, Puebla, Veracruz, and Hidal-
go. The type locality is Orizava, México.
A one-stripe or no-stripe garter snake with spotted and
Figure 29.97. The distribution blotched morphs can occur in the same litter. The spotted
of Thamnophis sumichrasti. morph has black-edged, lateral brown spots 0.5 to 2 scale rows
long on dorsal scale rows 3-5 or 3-8 with vertebral bars. Some-
times present are alternating rows or rows of spots on the first
dorsal scale rows. Spots are best developed in young animals and may disappear with age.
The blotched morph has 34-49 black-edged blotches that extend from the vertebral line to the
first dorsal scale row. The blotches are 3-3.5 scale rows long - stripes are always absent in this
morph. Comma-shaped nuchal blotches with the tails ending on the parietal scales.
Eight upper labials, ten lower labials. Dorsal scales in 19 rows and lack apical pits. Ventral
scales are 147-161, subcaudals 57-80.
Natural History. This snake has been reported to feed on the toad Incilius cristatus
(Cerón-de la Luz et al. 2020) and a Pseudoeurycea salamander (García-Bañuelos et al. 2020).

556
 29. The Thamnophiine Clade

Figure 29.198. Thamnophis sumichrasti from Agua Zarca, Tenango de Doria, Hidalgo, México.
Photography by Fernandez Ballido.

Madrean Narrowheaded Garter Snake

Thamnophis unilabialis (Tanner, 1985)
Distribution and Habitat. A highly aquatic Thamnophis
Rock-strewn stream in the Mexican states of eastern Sonora,
Chihuahua, and Durango. The type locality is 0.5 mi SW of
Bocoyna, Chihuahua, Mexico.
Adults can reach 93.5 cm. Overall this snake is brown
with little pattern, which contrasts with the coloration of
Figure 29.99. Distribution of adults from western Chihuahua that are dark brown with five
Thamnophis unilabialis.
or more rows of reddish to black spots or black-bordered red
spots on the dorsum (Lemos-Espinal and Smith 2007). Neo-
nates in Sonora show the more typical spotted pattern. Ventral
scales are 151-169, subcaudals are 65-82. In addition, it has two enlarged nape spots, six rows
of dark brown or rust-colored spots on the body, and ventral scales usually have anterior
margins heavily pigmented to form a light-and-dark, crossed-barred pattern.
Natural History. Most were found under rocks near streams; one was captured in a min-
now trap. An unvouchered specimen from Arroyo La Presita had a Gila minacae in its stom-
ach (Rorabaugh et al. 2019).

557
Aquatic Snakes

Figure 29.100. Thamnophis unilabialis. (A) From Temósachic, Chihuahua, MX. Photograph
from Jorge H. Valdez. (B) From Guadalupe y Calvo, Chihuahua, MX. Photograph from Eric
Centenero Acala (C) From Casa Blanca, Durango, MX. Photograph from Ivan Ahumada /
[Link].

558
 29. The Thamnophiine Clade

Mexican Westcoast Garter Snake

Thamnophis validus (Kennicott, 1860)
Distribution and Habitat. Marshes, meadows, canyon pools, streams, and irrigation canals
in Baja California Sur, Chihuahua, Jalisco, Sonora, Guerrero,
Sinaloa, and Nayarit. The type locality is Durango, Mexico
(in error). Conant (1969) corrected it to Río Presidio on the
coastal plain of southern Mexico. A polytypic species with
three subspecies.
Thamnophis validus celaeno (Cope, 1861) has a type lo-
cality in Cape St. Lucas, Lower California, revised to San Jose
Figure 29.101. Distribution of River, Cape San Lucas, Baja California, Mexico (Cochran,
Thamnophis validus. 1961) and distinguished by lateral pale stripes involving dor-
sal rows 1-3. Dorsal coloration is variable but with dark and
pale morphs. The dark morph is dark brown or black with little or no pattern except for the
lateral stripes. The venter is virtually uniform black or very dark brown. The dorsum is pale
gray or brown in the pale morph with 75-97 dark spots in four rows like mainland N. v. vali-
dus. The venter is white or yellow and may be immaculate or with a pale wash or stippled with
slightly darker pigment. There is a single preocular and usually three postoculars. Ventrals are
135-146 in males and 134-145 in females; subcaudals are 74-82 in males and 67-75 in females
(modified from McCranie and McAllister 1988).
Thamnophis validus isabellae (Conant, 1953) is on the west coast of Guerrero, Mexico.
The type locality is Pie de la Cuesta, Laguna Coyuca, Guerrero. They are distinguished by a
brown or chocolate brown dorsum, a pale stripe on the first two or three rows of dorsal scales
on each side, a low number of ventral scutes, and a slightly higher mean number subcaudals.
The venter gradually changes from a dull yellow on the anterior portion to a pale pinkish
posteriorly. There are usually four rows of dark spots on the dorsum, each spot occupying
only the anterior edge of anyone scale. The preocular is single, and there are three postoculars.
Ventral scales in males are 131-137, 130-138 in females, 76-81 subcaudals in males, and 68-76
in females (modified from McCranie and McAllister 1988).
Thamnophis validus thamnophisoides (Conant, 1961) Nayarit, Mexico. The type locality
is along the Rio San Cayetano also known as the Rio de Tepic on some maps and locally called
the Rio Mololoa), approximately 3.5 miles southeast of Tepic, Nayarit, Mexico. They are dis-
tinguished by a distinct pale middorsal stripe and pale scales on the first three dorsal scale
rows that form a lateral stripe on each side of the body. Dorsum coloration varies from brown
to olive brown, and the venter is a pale yellow with no dark pigment. There are a single preoc-
ular and three postoculars. Ventrals in males are 130-140, in females 127-139; subcaudals in
males 69-80, in females 65-73 (modified from McCranie and McAllister 1988).
A three-striped or two-striped garter snake that is semi-aquatic. There are 127-150
ventrals, eight upper labials (occasionally 7 or 9), and usually ten lower labials (sometimes 9
or 11) (modified from McCranie and McAllister 1988).
Size. Neonates are 16.5 cm, and adult females reach 1100 cm.
Adult males average significantly smaller than adult females. The largest measured by
Conant (1946) was a 92.5 cm female from Mazatlan, Sinaloa. The largest male was 60.5 cm.
Natural History. The species is diurnal and active year-round, but activity is reduced
from November to February (Grismer 2002). Adult Red Spotted Toads and their tadpoles,
Pacific Treefrogs, Couch’s Spadefoots, and mosquito fish are listed as prey by Conant (1969)
and Grismer (2002). Although reproduction is poorly known, Grismer observed neonates from

559
Aquatic Snakes

Figure 29.102. Thamnophis validus. Top Mexican West Coast Garter Snake (Thamnophis validus vali-
dus). Near Huatabampo, southern Sonora, Mexico. Photograph from Mike [Link] left. Cape
Garter Snake (Thamnophis validus celaeno). Baja Sur; bottom Cape Garter Snake (Thamnophis validus
celaeno) from Boca del Sierra. Photograph from Mike Pingleton.

early August to October and had a captive female that gave birth to 10 neonates in August.
At Laguna Coyuca this species inhabits mangrove forests, but the water is almost fresh.
Dunson (1980) placed adults in 100% seawater and found they lost only 0.76% of their body
weight daily. He also tested neonates and found they were not as tolerant to salinity as the
adults.

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Aquatic Snakes

On the next page. The Borneo endemic Hydrablabes periops.

Photography by (A) Paul Freed.(B) Chien C. Lee . When I
examined musuem specimens of these snakes it was clear
that there are multiple species hiding under the names Hy-
drablabes periops and H. praefrontalis. (JCM).

561
Aquatic Snakes

562
Aquatic Snakes

30. Natricids of Uncertain Status

These are species that have not been exsamined using molecular evidence to determine
how that are related to other groups.

Sumatran Keelback
Anoplohydrus aemulans Werner, 1909
Distribution and Habitat. Sumatra, Indonesia. Type lo-
cality: Pudang, Distrikt Babongan, Sumatra. This may be
Ujungpadang, Bakongan district, Aceh Provence. Known
only from the holotype.
The position of the nostrils and eyes suggests this
species is aquatic. The generic name also suggests Werner
believed this snake was aquatic in its habits (Nguyen et al.
Figure 30.1 The distribution of 2010). David and Vogel (1996) considered it a semi-aquatic
Anoplohydrus aemulans natricid inhabiting wet forests. Murphy (2007) suggested it
may be a homalopsid.

Boonsong’s Stream Snake

Isanophis boonsongi (Taylor and Elbel, 1958)
Distribution and Habitat. Northeast Thailand. Type lo-
cality: Phu Lang, Ban Khok, Na Phung, Dan Sai, Loei, Thai-
land.
Based on David et al. (2015), the posterior maxillary
teeth are much enlarged but not grooved. The nostrils are
dorsolateral and directed obliquely upwards; and the eye di-
Figure 30.2 The distribution of
Isanophis boosongi.

Figure 30.3 Isanophis boon-

songi from Lowei Province,
Thailand. Photographer cred-
it Merle J. Cox.

ameter is slightly greater than the eye-nostril distance and distinctly greater than the distance
between its lower margin and the margin of the lip. The pupil is round; dorsal scales are
strongly keeled with a thick median keel throughout the body. The body is stout; the head is
distinct from the neck, and the prefrontal scale is single.

563
 30. Natricids of Uncertain Status

Borneo Small-eyed Snakes

Boulenger (1891) described this genus as having; “Maxillary teeth small, equal, about 18;
mandibular teeth small, equal. Head small, not distinct from neck; eye small, with a round
pupil; nostril valvular, an oblique slit between two nasals; three pairs of chin-shields. Body
cylindrical, elongate; scales smooth, without apical pits, in fifteen or seventeen rows. Tail
moderate subcaudals in two rows.” This genus may be confused with Opisthotropis in the
literature. Seven specimens I (JCM) have examined had either 17 dorsal rows or 15 rows at
mid-body. A few had 21 or 19 rows just behind the head. They can have two or three pairs
of chin shields.
Bernstein and Ruane (2022) extract and sequence the DNA of homalopsid snakes from
several natural history collections and two specimens of H. periops from the Field Museum of
Natural History (FMNH). They use a high-throughput target capture approach to sequence
ultraconserved elements anchored hybrid enrichment loci and nuclear protein-coding genes
(NPCGs) commonly used in squamates.
They found the placement of H. periops supported the familial taxonomic status of Hy-
drablabes as a natricid, rejecting the hypothesis that this species is a homalopsid (Murphy
and Voris, 2014). They also obtained divergence dates of natricids slightly younger than those
of Burbrink et al. (2020). Given its distribution, the close relation of H. periops to other Asian
natricids is not unexpected. Containing H. periops (about 20.9 mya) may indicate population
dispersal into Borneo from the mainland, with subsequent extinction events outside Borneo.
Alternatively, the lineage ancestral to this clade may have dispersed into Borneo, followed by
an in situ speciation event. The divergence dates of H. periops and Trimerodytes occur at the
rise of the Hengduan Mountains of the eastern Tibetan Plateau (Western China) are consid-
ered a diversification driver of Trimerodytes about23.9 mya (Guo et al., 2020).

Olive Small-eyed Snake

Hydrablabes periops (Günther,1872)
Distribution and Habitat. Forest streams of East Malaysia,
Borneo/Sabah, Sarawak), and Indonesia (Kalimantan), and
Brunei Darussalam. The type locality is Matang, Borneo. Ele-
vational distribution from sea level to 710 m ASL. See the cover
photos for this chapter on page 601.

Figure 30.4 The distribution

of Hydrablabes periops.

Mocquard’s Small-eyed Snake

Hydrablabes praefrontalis (Mocquard, 1890)
Distribution and Habitat. Forest streams East Malaysia (Borneo: Sabah), Indonesia
(Kalimantan), and Brunei. The type locality is Mount Kinabalu, Sabah, Malaysia, Island of
Borneo.

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Aquatic Snakes

565
 31. African Rain Forest Water Snakes

31. African Rain Forest Water Snakes

The genus Grayia is widespread in the lowland forests of central Africa and present in
the countries of Angola, Benin, Burkina Faso, Burundi, Cameroon, Central African Republic,
Congo, the Democratic Republic of the Congo (Zaire), Equatorial Guinea, Ethiopia, Gabon,
Gambia, Ghana, Guinea, Guinea-Bissau, Ivory Coast, Kenya, Liberia, Niger, Nigeria, Rwanda,
S Chad, S Mali, Senegal, Sierra Leone, Sudan, Tanzania, Togo, Uganda, Zambia.
The genus was long considered part of the family Colubridae but has more recently been
placed in its own family, Grayiidae. and even more recently suggested to be the sister of the
Colubrinae (Ruane and Austin 2017; Zaher et al. 2019) and considered a subfamily of the
Colubridae (Chaney et al. 2021). The genus is endemic to tropical Africa and is the only genus
in the Grayiidae.(or Grayiinae)
Local people hunt these snakes for “bushmeat” and various medicinal uses. Unfortunate-
ly, conservative morphology makes identification difficult, and several “Grayia” samples on
GenBank included misidentified specimens in other genera. Chaney et al. (2021) used mo-
lecular and morphological data to reconcile the current taxonomy of Grayia. They describe
them as locally abundant, notoriously tricky to catch, and note that they occur in areas where
fieldwork is challenging. Habitat includes seasonal rainforest swamps, streams, and perma-
nent water bodies. The diet is composed of fish and frogs. Females deposit eggs in the leaf
litter accumulated between the buttress roots of trees during the dry season. These snakes are
unusual in depositing their clutch in small batches of three or four eggs at different times and
more than one nest site rather than laying all the eggs simultaneously. The genus contains four
species.

Long-tailed African Water Snake

Grayia caesar Günther, 1863
Distribution and Habitat. A central African forest species in
seasonal rainforest swamps, streams, and permanent water bodies
from Equatorial Guinea, C/E/S Cameroon, Democratic Republic
of the Congo (Zaire), Brazzaville, Congo, Gabon, Central African
Republic, and Angola. Type locality: Fernando Po.
A single female specimen of this species was described by
Figure 31.1 Distribution of Schmidt (1923) and was collected at Niapu. The specimen agrees
Grayia caesar.
with Boulenger’s description except for having only two posteri-
or temporals. The color is grayish-brown above, with thirty light,
narrowly black-edged crossbars, occasionally interrupted on the vertebral line. Tail faintly
crossbarred. Ventral surface is entirely uniform gray-yellow. Lower labials are immaculate,
except the last. Upper labials are pale, with dark markings mostly on the sutures. The top of
the head is black, with a faint light spot on each parietal and still fainter ones on the frontal.
A pale postocular stripe is followed by a sharply defined one from the mouth’s rictus to the
parietals’ rear corner.
Natural History. The diet seems to be composed chiefly of fish, particularly catfish. It
hunts in the water. It may also hunt from arboreal perches but see the following account.

On the previous page is Smyth’s African Water Snake, Grayia smythii. Photography by Kate Jackson.

566
Aquatic Snakes

Ornate African Water Snake

Grayia ornata (Bocage, 1866)
Distribution and Habitat. A central African forest spe-
cies present in seasonal rainforest swamps, streams, and per-
manent water bodies in Angola; the Democratic Republic of
the Congo (Zaire; except in the west); Brazzaville; Congo; Ga-
bon; Equatorial Guinea; Cameroon; Zambia; Central African
Republic. Type locality: Duque de Bragança, Bayão, Angola.
Figure 31.3 Distribution of Grayia ornata has two distinct color patterns: the typical
Grayia ornata. transverse bar color phase and the rare longitudinally striped
phase. Spawls and Branch (1995) noted the extreme similari-
ty between the cross-barred color phase of G. ornata and the
sympatric aquatic cobra Naja annulata.
The following is based on Schmidt (1923). The largest male measures 1240 mm., and the
largest female 1385 mm. The tail length varies from 0.25-0.32 in males, mean 0.28, and from
0.24- 0.26 in females, mean 0.25. The ventrals range from 148 to 155, mean 153, in males, and
from 153-160, mean 156, in females; subcaudals 69-87, mean 82, in males, 69-78, mean 74,
in females—one preocular, two postoculars, and temporals 2-3 in every specimen. The lower
anterior temporal is frequently separated from the postoculars. Five specimens have both
loreals fused with the prefrontals; three have the loreal of one side fused. The upper labials
number 8-10, and the lower 10 or 11. Three specimens from Medje and Niangara represent
the longitudinally striped form. The light ventral color extends on the sides to the middle of
the third scale row. Two interrupted black lines are distinct on this ground color, one on the
ends of the ventrals and the first scale row and a second between the first and second scale
rows. These lines consist of streaks six or seven scales in length, with interspaces of two or
three scales. The upper half of the third scale row, the fourth, and the lower half of the fifth
are occupied by a black band extending from the last upper labial. In adaption to its habitat,
the tail is markedly compressed.
Natural History. Diet is composed mainly of fish. They will bask in branches overhang-
ing the water and dive in when disturbed. Chippaux’s (1999:83) report that G. ornata can
hunt from branches overhanging streams was refuted by people interviewed by Pauwels et al.
(2002) on the Massif du Chailiu, Gabon. The species is considered an excellent climber and
likes to bask on branches overhanging the water, sometimes two to three meters high. When
alerted, they fall into the water and quickly disappear into holes in the banks. They are said
to be solitary and to hunt underwater at night, during which they actively explore every nook
and crevice for prey items. It is believed in the Lunda area of north-eastern Angola that G.
ornata live in pairs (Laurent, 1964: 102), but Pauwels et al. (2002) did not support this.
The stomach contents were usually fish (especially silurids), but frogs, shrimp, and small
rodents sometimes were in stomachs. At Boussimbi, Diangui and Iboundji Pauwels et al.
(2002) were told that the smell of G. ornata strongly attracts fish. Fisherwomen search the
places where this snake lives as they think there will be many fishes there. Perhaps in sup-
port of this belief, one of the authors observed cyprinid fishes (genus Barbus) feeding on
excrement defecated by G. ornata in the Loeme River, Kouilou Region, Republic of Congo.
Itsiba inhabitants reported that otters often destroy hoop nets to eat the entrapped fish and
dibomina (= G. ornata). Several fishermen of Diangui told Pauewels et al. (2002) sometimes
find dibomina in the stomachs of the crocodilians Crocodylus cataphractus and Osteolaemus
tetraspis.

567
 31. African Rain Forest Water Snakes

Villagers in Itsiba and Moudouma use G. ornata as the

Figure 31.4. Grayia ornata (A)
essential remedy for delivery problems during childbirth. It Kate Jackson with a G. ornata
is also used as a preventive measure a few days after a baby in a fishing net, and (B) a G.
is born; some people take the viscera of G. ornata (first dried ornata from Ganganya Brous-
in the smokehouse) and use it as a bangle around the an- se, Republic of Congo. (C-D)
kle of the baby. This bangle is believed to suck out sickness: A juvenile ornate from Uma-
“Like the dibomina can go out of the water, the sicknesses ngi, DRC, (E) Photography
can go out of the body.” The authors were also informed in by (A-B) Kate Jackson, (C-E)
Diangui that the fat of dibomina (as well as that of other Vaclav Gvozdik.
snake species, mainly Python sebae), collected while cook-
ing, can be used to treat sprains by direct application on the skin. This latter use is also current
by the Pounou at least in Mouila. In the southern Republic of Congo, the fat of this species
is applied on the skin to facilitate removing a thorn or a splinter. At Boussimbi, Diangui and
Iboundji, G. ornata is used by fishermen for two magical purposes, to obtain excellent swim-
ming ability and to be a successful fisherman (Pauwels et al. 2002).

568
Aquatic Snakes

Shadow African Water Snake

Grayia obscura Chaney, Greenbaum and Pauwels 2023
Distribution and Habitat. DRC and northeastern Angola, this cryptic species ranges
from the Middle to Upper Congo River and its tributaries. In the Middle Congo River, it is
probably sympatric with G. ornata sensu stricto. In the Middle Congo River, It is moleculary
and mporphologically G. ornata. The holotype is from Mungombe in the South Kivu Prov-
ince of the Democratic Republic of Congo.
Natural History. A single individual ontained the clariid catfish, Clariallabes platyceps.
Although the authors speculate that a Grayia ornata was discussed by Broadley in1983, was
most likely a Grayia obscura from Zambia. The prey was identified as the frog Rana ango-
lensis (now Amietia delalandii or A. chapini) . The type locality and the locations of the para-
types are located in a transitional forest between the lowland rainforest of the Congo Basin
and the foothills of the western Itombwe Plateau.

Figure 31.5. The Shadow African Water Snake, Grayia obscura.

Photography Konrad Mebert.

569
Aquatic Snakes

Smith’s African Water Snake

Grayia smithii (Leach, 1818)
Distribution and Habitat. A western and central African forest species in seasonal rain-
forest swamps, streams, and permanent water bodies in southern Sudan; Uganda; Kenya;
Tanzania; Rwanda; Burundi; Angola; Democratic Republic of the Congo (Zaire); Brazzaville,
Congo; Equatorial Guinea; Cameroon; Nigeria; Benin; Togo; Ghana; Ivory Coast; Liberia;
Guinea (Conakry); Guinea-Bissau; Senegal; Burkina Faso; Sierra Leone; Gambia; Central Af-
rican Republic; probably southern Mali; Niger. Type locality Boma, Zaire.
Leach (1818) spelled the name smythii in the original description. He named the species
after the Norwegian botanist Christen Smith (1785-1816). Some subsequent authors changed
the spelling to smithii because the spelling “smythii” was considered inappropriate. Wallach et
al. (2014) followed the original spelling, as do I.
The following is based on Schmidt (1923). This species is considerably larger than G. or-
nata. The largest male measures 1535 mm., and the largest female 1350 mm (tail incomplete).
The tail length in three males varies from 0.29-0.33 of the total; the single female with a com-
plete tail is 0.29. Ventral scales range from 149-160 in males, 157-162 in females; subcaudals
90-96 in males, 92 in one female. Dorsal scales 17-17-15, 19 on the neck on one specimen.
One preocular, two postoculars, and temporals 2-3 in all specimens. Three or four slightly
enlarged occipitals. Upper labials 7, the last very large, lower 10 or 11. There are twenty-three
narrow light crossbands, with indications on the sides of eleven more. Adults have black
crossbars, always with a row of light spots on the black scales, more or less in the middle of the
bar. In two specimens, more than thirty such bars are visible; the posterior ones are merely in-
dicated dorsally. In other specimens, the color becomes uniform on the posterior two-thirds
of the body, with ten or twelve crossbars anteriorly. The tail
scales are outlined with black, giving it a longitudinally lined
effect and a similar line edges the venter. The throat is white,
the lower labials black-edged. The parietal shields are gray,
with black borders.
Natural History. The ecology of the water snake, Grayia
smythii was examined by Akani and Luiselli (2001) using a
Figure 31.5 Distribution of
population in a seasonal rainforest swamp of the Niger Delta
Grayia smithii.
(southern Nigeria). The two sexes were similar in body sizes
(SVL) and head sizes, but males had tails significantly longer
than females. The diet consisted of frogs and fish. The major prey was the frog Xenopus trop-
icalis, followed by the fishes Tilapia sp. and Clarias sp. The adult sex-ratio was 1:1. Sloughing
of skin and oviposition occurs during the dry season in the humid enclosure of buttress roots
amongst leaf litter.
Clutch size ranged from 8 to 14 eggs per female, with a mean of 10. The smallest gravid
female was 78.2 cm SVL. In at least two to three different sites, eggs were laid in batches of
three to four eggs at a site. The eggs averaged 3.1 cm in length, 2.1 cm in width and had a fresh
weight ranging from 18,2 to 22.1 g. Maternal size significantly influenced the number of eggs
produced by females, but not their average size. A statistically significant negative correlation
existed between the number of eggs and mean egg size. Predators of this snake in the study
area were herons and fishermen.

570
 31. African Rain Forest Water Snakes

Figure 31.6. Smith’s African Water Snake, Grayia smithii. (A) is a juvenile (B) is an adult.
Photography by (A) Vaclav Gvozdik (B) Kate Jackson.

Tholloni’s African Water Snake

Grayia tholloni Mocquard, 1897
Distribution and Habitat. A savanna species in seasonal wetlands, streams, and perma-
nent water bodies in southern Sudan, Uganda, Kenya, Tanzania, N/E/S Democratic Republic
of the Congo (Zaire), Congo, Angola, Central African Republic, Ethiopia, Gambia, Senegal,
Nigeria, Guinea (Conakry), maybe Cameroon, Zambia, and southern Chad. Type locality
Brazzaville, Congo.
Schmidt (1923) examined two specimens; both specimens had incomplete tails. The

571
Aquatic Snakes

head pattern distinguishes G. tholloni from G. caesar. Also

the fifth labial, instead of broadly entering the orbit it is
separated from it by a lower postocular or enters the orbit.
Grayia caessar and G. tholloni agree in having much narrow-
er postoculars than G. smythii and G. ornata, as well as in
the larger eye. Color gray-brown above, with very indistinct
Figure 31.7 Distribution of light crossbars. The venter and underside of the tail are im-
Grayia tholloni. maculate light yellow, outlined with black at the juncture
with the dorsal color. Upper and lower labials were black-
edged, and the black between the last two upper labials con-
tinued upward as a bar across the temporal scales.

Figure 31.8. Grayia tholloni.

Photography by Konrad Me-
bert.

On the next page is a Xenelaphis hexagonotus from Singapore.

Photography by Kenneth Chin.

572
 31. African Rain Forest Water Snakes

573
Aquatic Snakes

32. Colubridae
Aquatic colubrids (or Colubrinae) are scattered in several clades, two in Africa and
one in Asia. The members of the genus Crotaphopeltis were examined for genetic structure
(Engelbrecht et al. 2017, 2019). The widespread, terrestrial Crotaphopeltis hotamboeia did
not show cryptic diversity at the species level, although there was a clear phylogeographic
structure (Engelbrecht et al. 2017, 2019). Crotaphopeltis hotamboeia diverged from its more
aquatic Central and East African sister taxa, C. degeni and C. tornieri, in the mid-Miocene, a
period characterized by a global cooling event. This caused the fragmentation of a west-to-
east African forest belt and the emergence of savanna in Africa. The habitat changes likely
induced allopatric speciation of forest and open habitat specialist species within this genus.
Diversification took place at the forest-savanna ecotone in East Africa. Intraspecific diversi-
fication for C. hotamboeia commenced during the Miocene-Pliocene transition period, ca.
6 Ma (Engelbrecht, 2017). The Early-Pliocene forest expansion, which extended into coastal
Kenya, Tanzania, and Ethiopia (Pickford, Senut, & Mourer-Chauvire, 2004; Sepulchre et al.,
2006) probably excluded C. hotamboeia from East Africa.

Barotse Water Snake

Crotaphopeltis barotseensis Broadley, 1968
Distribution and Habitat. It inhabits the papyrus swamps of
Zambia, northern Botswana (Okavango Swamp, along Chobe
River to upper Zambezi River), and Angola. It is also present in
Ethiopia. Type locality: Kalabo, Barotseland, Zambia.
Natural History. A nocturnal frog-eating snake. Anurans
were 97% of the food taken in one study. The taxonomic diver-
sity of the prey items was large, with six families represented
among the 51 anurans identified to family; however, bufonids
(39%), ranids (29%), and microhylids (25%) comprised the bulk
Figure 32.1. The distribu-
tion of Crotaphopeltis barot- of the diet. These snakes feed all year except for midwinter. The
seensis. smallest sexually mature female was 295 mm SVL. The small-
est sexually mature male was 240 mm SVL. Clutch sizes ranged
from four to 12 eggs. Females appear to yolk their ovarian folli-
cles rapidly, starting in September. Egg deposition commences
in October and continues through January. The only significant
decrease in testicular activity was in mid-winter (July). Thus,
spermatogenesis coincides with female follicular development
and ovulation (Keogh et al. 2000).

Degen’s Herald Snake

Crotaphopeltis degeni (Boulenger, 1906)

Figure 32.2. The distribu-

Distribution and Habitat. Moist savannas, swamps, and
tion of Crotaphopeltis degeni. floodplains. Southern Sudan, Uganda, Kenya, Ethiopia, Tanza-
nia (relict population), eastern Central African Republic, Cam-
eroon, and Chad. Type locality: Entebbe, Uganda.

574
 32. Colubridae

Fig.32.3. Crotaphopeltis barotseensis Photography by Piet-Hein Stutterheim.

Figure 32.4. Crotaphopeltis

degeni. Defence posture-
sPhotography by Konrad
Mebert.

575
Aquatic Snakes

The Genus Platyceps

A widespread genus of about 30 speicies tend to be habitat and dietary .generalists.
The degree to which either of the two species discussed here use aquatic envirnments is
uncertain.

Sarso Island Racer

Platyceps insulanus (Mertens, 1965)
Distribution and Habitat. Coastal environments of
Sarso Island, Farasan Archipelago in the Red Sea, Saudi
Arabia. The type locality is Insel Sarso, Farsan-Archipel,
Rotes Meer, SW-Arabien. The holotype of Platyceps in-
sulanus was collected by W. Kost on 26 November 1964,
during the Senckenberg-Museum’s Meteor Expedition in
the Indian Ocean. The type specimen was found swim-
ming in shallow waters just off-shore of the island (Mer-
Figure . 32.5 The distribution of tens, 1965). The species was characterized by a uniform
Platyceps insulanus. dull lead grey coloration of the upper parts (Mertens,
1965). The holotype is preserved at the Senkenberg Mu-
seum Frankfurt (SMF). The species is represented only
Figure 32.6. Platyceps in-
sulanus. (A) The specimen
collected by W. Kost on the
26th November 1964, in the
course of the “Meteor” - Ex-
pedition launched by the
Senckenberg-Museum in the
Indian Ocean. Its type speci-
men was found while swim-
ming in shallow waters just
off-shore of the island (from
Mertens, 1965). (B) The ho-
lotype of Platyceps insulanus
is preserved at the Senken-
berg Museum Frankfurt, SMF
60027 (Michael Franzen, Sen-
kenberg Museum Frankfurt).
Note both photos from Masseti
(2014).

576
 32. Colubridae

by this specimen, possibly subadult (Mertens, 1965) and its shed skin. The total length of the
only known specimen is 760 mm (Mertens, 1965; Gasperetti, 1988). Mertens (1965) originally
believed that Sarso was an island of the Dahlak archipelago, of the coast of Eritrea. Following
Kost’s report and the subsequent description by Mertens (1965), the snake was never found
again, either on Sarso or on other nearby islands. Masseti (2014) suggests that this species is
extinct.

Jan’s Cliff Racer

Platyceps rhodorachis (Jan, 1863)
Distribution and Habitat. A widespread species that
uses many different habitats. Known from Turkey, Algeria,
Egypt, possibly Sudan, Eritrea, Ethiopia, Somalia, Chad,
Syria, Jordan, Saudi Arabia (incl. Farasan Islands), Unit-
ed Arab Emirates, Oman, Iraq, Iran, Yemen, Afghanistan,
W Pakistan, NW India, southern Turkmenistan, southern
Uzbekistan, northwest Tajikistan, western Kyrgyzstan,

Figure 32.7 The distribution of

Platyceps rhodorachis.

southern Kazakhstan, and China

(Tibet). The type locality is Persia;
restricted to the Arabian Peninsula;
Shiraz, Iran.
Inhabits arid regions with
stone, and gravel substrates. Often
desert habitats with associated hills
and cliffs. It also penetrates desert
along the Tejen and Murgab river
valleys and is found in rocky situa-
tions bordering floodplains, includ- Figure 32.8 Platyceps rhodorachis from Natanz, Isfahan
ing dry irrigation canals (Ananjeva et Province, Iran.
al. 2006). It can also be found in Photography by Hossein Nabizadeh.
mountain steppes and juniper scrub
(Ananjeva et al. 2006). It may be found close to cultivated land and villages, and swimming in
wadis. Oviparous, lays 3-4 eggs per clutch; 4-9 per clutch have been reported in Kazakhstan
(Els et al. 2001).

The Genus Philothamnus

Angola Green Snake

Philothamnus angolensis Bocage, 1882
Distribution and Habitat. Reed beds, stream banks, Namibia, northern Botswana, Zim-
babwe, southern Mozambique, Okavango Swamp, Caprivi Strip, Angola, Democratic Repub-
lic of the Congo (Zaire; except in the north), Congo (Brazzaville), Cameroon, Central Af-
577
Aquatic Snakes

rican Republic, Sudan, South Sudan, Tanzania, Zambia. Type

locality: Capangombe, Angola. Known from sea level to 2,000
m. Natural History. The diet is composed mainly of frogs and
lizards. Females lay 5-16 eggs that hatch in December-February
(Broadley 1983).

Southeastern Green Snake

Figure 32.9. The distribu- Philothamnus hoplogaster (Günther, 1863)
tion of Philothamnus ango-
lensis Distribution and Habitat. Marshes, ponds, and rivers in
Zimbabwe, S Mozambique, Zimbabwe, Tanzania, Kenya, Ma-
lawi, E/S Democratic Republic of the Congo (Zaire), Rwanda,
Burundi, Zambia, Republic of South Africa (Transvaal, Natal,
Cape, Swaziland), Botswana, Namibia, Angola. Type locality:
Port Natal (= Durban, South Africa.)
Natural History. Young individuals will eat coleopterans,
orthopterans, and other insects. However, fish, frogs, and liz-
ards are the most frequent prey of adults. Females lay clutches
Figure 32.11. The distribu- of 3-8 eggs in early summer (Broadley 1983).
tion of Philothamnus hoplo-
gaster.

Figure 32.10. Philothamnus

angolensis. Photography by
Vaclav Gvozdik.

Figure 32.12. Philothamnus

hoplogaster. Photography by
Paul Freed.

578
 32. Colubridae

Three Asian colubrids that show aquatic habits

Frog-eating Rat Snake

Oocatochus rufodorsatus (Cantor, 1842)
Distribution and Habitat. Primorskiy Territory, Amur
Oblast in Russia, Korea, Taiwan, northeast China. The type lo-
cality is Zhejiang, Chusan island, China (Cantor, 1842).
Natural History. Diet includes frogs (Pelophylax nigro-
maculatus) including the invasive American Bullfrog (Heo et
al. 2014). Radiotracking 21 snakes over two seasons on agricul- Figure. 32.13. The distribu-
tural land during two active seasons. Lee et al. (2011) demon- tion of Oocatochus rufodorsa-
tus.
strated both sexes stayed close to aquatic habitats such as paddy
fields and agricultural ponds during breeding and non-breed-
ing periods, except when they moved to dry areas to hibernate in late fall. The use of structural
features in the habitat (ground, tree, underground, and water) depended on the temperatures,
the female’s reproductive conditions, and the time of day. Both sexes moved about 17 m daily

Figure 32.14. Oocatochus rufodorsatus.

Photography by Amaël Borzée.

and postpartum females moved farther than antepartum females. The home ranges of males
and females were 0.45 ha and 0.47 ha, respectively, and the year-round home range of this
species was approximately 1.54 ha. Therefore, conserving a population of rufodorsatus would
require aquatic and terrestrial habitats within a radius of 150 m from a pond habitat. require
aquatic and terrestrial habitats within a radius of 150 m from a pond habitat.

Ornate Brown Snake

Xenelaphis ellipsifer Boulenger, 1900
Distribution and Habitat. Primary inhabits and old secondary forests from lowlands to
1100 m in elevation. Known from six specimens from Sumatra, Indonesia, and the island of
Borneo. At least one specimen was taken from a fish trap. Other specimens were taken from
streams and stream edges. Malaysia (Malaya and East Malaysia; Sarawak). The type locality is

579
Aquatic Snakes

the Sarawak River.

Natural History. Adults probably feed on fish and rodents, while juveniles prefer fish,
frogs, and lizards (Malkmus et al. 2002). Collectors found this snake foraging in and along a
moderately flowing stream and were quick to escape when approached. The same snake was
observed along the same stream multiple times, which might suggest a small home range.

Malayan Brown Snake

Xenelaphis hexagonotus (Cantor, 1847)
Distribution and Habitat. Inhabits peat swamps, scrublands, agricultural fields and
freshwater swamps to elevations of 500 m. Indonesia (islands of Bangka, Belitung, Java,
Kalimantan, Riau Archipelago, Sumatra, Borneo); peninsular
Malaysia; Singapore; Thailand; Vietnam. Its presence in Myan-
mar is questionable.
Type locality: Great Hill of Pinang (Pulau Pinang, West
Malaysia) .
Natural History. A diurnal semi-aquatic snake that preys
Fig. 32.15. Distribution of Xe- upon fish, amphibians, and [Link] reaches a length of
nelaphis hexagonotus. 200 cm.

Figure. 32.18. Xenelaphis hexagonotus.

Photography by Teo Eng Wah (Vincent).

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Aquatic Snakes

Above is a Lake Chapala Garter Snake, Thamnophis eques obscurus, from Jocotepec, Jaloisco,
MX. Photography by Ethan Hazel. This snake and at least one other were entangled in a gill
net on the shore of Lake Chapala. This snake is considered Vulnerable under Mexican law.

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 33. Conservation

33. Conservation
Michael J. Jowers

According to the International Union for the Conservation of Nature (IUCN) globally,
9% of seasnakes are threatened, 6% are near threatened, and 34% are data deficient (Elfes et
al. 2013).
Elfes et al. (2013) reported on only two seasnakes (Aipysurus apraefrontalis and A. fo-
liosquama) threatened as Critically Endangered (CR), both reef inhabitants found at Ash-
more and Hibernia reefs, respectively, in northwestern Australia and have undergone dra-
matic population declines. Nevertheless, their assessment in IUCN assessment from 2018
reports them as Data Deficient (DD) today. One species is endangered (EN); the reef species
Aipysurus fuscus has similarly undergone population reductions (~70%). Seven species are
Vulnerable or Near Threatened primarily due to small geographic ranges and habitat degra-
dation. Two Vulnerable species are known from a single lake system each. Hydrophis semperi
is a freshwater species only found in Lake Tall in the Philippines. Still, its population status
is unknown, and Laticauda crockeri is endemic to Lake Te-Nggano on Rennell Island in the
Solomon Islands. The third Vulnerable species, Laticauda schistorhyncha, is endemic to the
island of Niue, while the Near Threatened species with restricted ranges include Laticauda
frontalis (endemic to Vanuatu) and Laticauda guineai (endemic to Papua New Guinea). Five
of the eight species in the genus Laticauda are classified as Vulnerable or Near Threatened.
Many of the seasnakes considered in this book are listed by the IUCN as Dada Defi-
cient, yet many of these species require conservation status. An example is Enhydris jagorii,
endemic to the Chao Phraya-Ta Chin basin in Thailand and threatened by habitat change
and destruction, with large areas where it is found transforming into urban and agricultural
areas and threatened by uncontrolled fishing. Despite the species-restricted distribution, it is
under the category of Data Deficient, while the evidence may suggest a Critically Endangered
status (Pongcharoen et al., 2016). Widely distributed species have been extensively studied
from a conservation perspective (Roe et al., 2003, 2006, Mebert 2011), but species restricted
to isolated locations or endemic in remote areas remain mostly Dada Deficient. Overall, for
many of the species listed in this book, despite them being listed as Least Concern, limited in-
formation, or no information at all is available on threats. Their population trends are mostly
unspecified, as well as most populations being listed as not severely fragmented, which might
result from lack of accurate data rather than actual knowledge.
Studies on the on conservation of freshwater aquatic or semi-aquatic snakes are lacking
and overall are challenging to summarize. Their worldwide distribution and taxonomic dif-
ferences as well as their natural history and freshwater biota habitat, adds implicit complexity
to assess their conservation in a review. From the data available from the IUCN there are
apparent differences in the species group assessments, which reflect their taxonomy, where
they are found as well as their economic importance. For example, the data on the risks and
threats of giant snakes (boids, pythons, anacondas) is ample, as is their conservation action.
This is in strong contrasts to those of general such Pseudoxenodon, Thermophis, Farancia,
Coniophanes, Helicops, Hydraethiops, Limnophis, Thamnophis, Hydrablabes, among others,
that remain almost blank in their IUCN assessments.
Needless to say is that such a vast group of reptiles inhabiting the world's aquatic or
semi-aquatic freshwater and marine biotas are under diverse and multiple threats. Here we
list some of the most reported or better known threats to such taxa.

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Aquatic Snakes

Habitat degradation
Sea kraits (genus Laticauda) have unique habitat requirements, being dependent on
intact shallow coral reefs, supralittoral nesting sites, and appropriate intertidal and inland
resting sites (Lillywhite et al. 2008; Bonnet et al. 2009), and minimal disturbances to these
habitats will have important consequences to seasnake populations inhabiting them. Areas
where species were common seem to have undergone dramatic population declines. For
example, ten species were common in Ashmore Reef (Minton and Heatwole 1975; Guinea
and Whiting 2005) while several surveys have only found one species in the reef (Elfes et
al., 2013). Declines have also been noticed on sea kraits with broader geographic distribu-
tions, suggesting some kind of habitat deterioration. In fact, coral reefs are threatened by
pollution, climate change, and diseases (Hughes et al. 2003; Pandolfi et al. 2003; Wilkinson
2008). Such factors might have important ecological consequences in the trophic system,
having negative impacts on seasnake prey abundance in their hunting grounds. Seasnakes
are dependent on undisturbed small islands and are more often than not encountered less
than a few meters from the water edge on beach rock which is used as necessary terrestrial
refugia (Bonnet et al. 2009). Therefore, minimal disturbances to such delicate ecosystems
have resulted in population declines (Brischoux et al. 2009)
Many aquatic or semi-aquatic snakes require both, aquatic and terrestrial habitats
and remain a dependent on aquatic habitats, and many wetland reptiles require movement
through, or extended refuge within terrestrial habitats for nesting, hibernation, migration,
dispersal, foraging, etc. Wetlands constitute vastly large and important habitats for reptiles,
but in some countries such as the USA, they are being lost quickly (Dahl, 1990), and so
are the species inhabiting them (Gibbons et al., 2000). Habitat alterations that change the
spatial distribution of wetlands, such as the loss of small isolated wetlands, are believed to
have increased mortality rates for N. e. neglecta. Conservation strategies for vagile wetland
animals inhabiting variable aquatic resources over large spatial scales should take advan-
tage on protecting and restoring large areas with numerous, heterogeneous wetlands (Roe
et al., 2004). Simulations of small wetland loss shows that highly vagile species such as N.
e. neglecta would move longer distances to find suitable habitat, suggesting that protection
and restoration of wetland landscapes are critical for the long-term persistence of many wet-
land-associated species (Roe et al., 2003). Similarly, studies on radiotracking of population
movement of Oocatochus rufodorsatus suggest that this aquatic species also uses terrestrial
habitats within 150 of the core pond habitats, which need to be preserved (Lee et al. 2011)
Many wetland reptiles migrate or disperse at a temporal and spatial scale, and such as-
sociated movements put them at risk especially in areas heavily modified by humans. Con-
sequently, in addition to threats imposed by wetland loss, many wetland reptiles may have a
high vulnerability to dispersal through roads. Despite the high mortality of reptiles on roads,
conservation hardly addresses the threats that roads impose to wetland species. Roe et al.
(2006) used models integrating road maps, traffic volume and snake movements to assess
the mortality of species with their vagility in three regions of the USA. Their findings re-
vealed that the vagile species Nerodia e. neglecta mortality accounted for between 14-21% of
its populations, while the more sedentary species Nerodia sipedon mortality was much lower
(3-5%). They concluded that conservation measures should consider not only the quality of
wetlands and nearby terrestrial habitats, but also terrestrial corridors between wetlands and
safe passage for wildlife.
Climate change is a global problem affecting all species worldwide, however, its effects

583
 33. Conservation

will likely be seen sooner rather than later in marine and freshwater ecosystems and their
associated wildlife. In freshwater ecosystems, drought will impose strong pressures on species
survival. Vogrinc et al (2018) assessed the effect of drought on semi-aquatic snake popula-
tion occupancy in South Carolina before (year 2006) and after the drought (year 2013). They
found that site occupancy decreased dramatically following drought for Nerodia fasciata
(0.95 to 0.69) and Nerodia floridana (0.32 to 0.05), but was relatively unchanged for Semina-
trix pygaea, Farancia abacura and Regina rigida. This study concluded that species lacking
adaptations to adapt to drought may become locally extinct if climate change projections are
correct or if landscapes are degraded or fragmented in ways that prevent drought.
Santos and Llorente (2009) conducted a demographic study to evaluate the relative
densities of snakes in natural lagoons in 1995 in the Ebro Delta (Spain). In 2008, they repeated
the same study in the same area and season. The field work consisted of 29 censuses and
their potential prey. In 1995, they recorded 27 Natrix maura (0.93 animals/ha), occupying
48% of the sites but in 2008 no snakes were encountered. However, prospected rice fields
nearby detected a great number of snakes, with 25% occupancy overall. They attributed such
findings to habitat transformation and degradation, natural predator increment, decrease in
prey populations, pollutants in rice fields, death on roads, and thorough human persecution.
They argue that the recovery of the N. maura population in the rice fields in the region
depends on agricultural management changes, reducing of pollutants, the recovery of snake
prey, and the maintenance of favorable habitats to prevent predation.

Pollution

Seasnakes can be used as bioindicators for the health of the marine environment because
of the uptake of trace elements via the ingestion of prey, and the snake’s accumulation of
trace elements over time. Marine snakes in fishing areas could indicate regions where fish
consumption may have important health implications.
Pollutants are carcinogenic and accumulate in different parts of the snakes´ body, such
as the liver, kidneys, blood, and fatty tissues. Gillet et al (2017) found that several species
in southern Queensland (Australia) suffered from pancreatic, liver and kidney neoplasia,
thought to be derived from toxigenic or viral etiology. In a recent study, Goran et al (2017)
surveys revealed that in the seasnake Emydocephalus annulatus, melanism is more frequent
in urban-industrial sites than in less polluted locations. High trace-element concentrations in
sloughs of sea kraits from close to a river-mouth suggest that melanism may benefit seasnakes
in many areas. Importantly, concentrations of trace elements were higher in darker than in
lighter bands within the same slough.
Growing evidence in recent years suggests increased seasnake mortality caused
primarily by marine debris and plastics on a global scale, but what the actual effects on species
and populations remain largely unknown. The bioaccumulation of marine contaminants
and pollutants and the threshold required for the accountable detrimental effects on snake
populations and the regions of the highest impact are key points for population management.
Pollutants are not only found in the marine environment, and can are traceable in fresh-
water species too. In a recent review of pollutants detected in snake tissues, Quoc Hoang et al
(2021) found that concentrations of polychlorinated biphenyls (organic chlorine compounds)
were found in numerous snake tissues. Within the USA, some reported species were Dia-
mondback Watersnake (Nerodia rhombifer), Blotched Watersnake (Nerodia erythrogaster),
Northern Watersnake (Nerodia sipedon), Cottonmouth (Agkistrodon piscivorus). In Canada;
the Lake Eire Water snake (Nerodia sipedon insularum and N. s. sipedon), and in China, the

584
Aquatic Snakes

Chinese Mudsnake (Myrrophis chinensis). Traces of flame retardants have also been found in
Chinese Mudsnake. Other known pollutants are pesticides, as seen in Lake Erie Watersnake
(Nerodia sipedon insularum).
The mining industry has affected the distribution pattern of the Copperbelly Water-
snake, Nerodia erythrogaster. Populations have been isolated, making them more vulnera-
ble to threats. The major causes for population fragmentation is coal mining, drainage and
damming of wetlands, channelization, diversion of streams and rivers, and development of
upland habitat. Although the impact of the mining industry on freshwater snakes remains
unreported and unknown, it is likely that mining has profound implications for both the
health and the fragmentation of populations. Large mining operations likely cause whole
snake populations to relocate to other areas, affecting niche resource partitioning and creat-
ing competitive displacement between species. The frequent use of water in mining (eg., gold
mining) has devastating effects on natural river drainages, increasing water turbidity, pollu-
tion by heavy metals, and changing water systems for snakes to breed. In addition, the effects
of mining acoustic pollution remain unreported, but is likely to cause habitat shift of snakes
and their prey for much larger areas than the mining regions. Interestingly, Lacki et al. (2005)
found that Copperbelly snake populations abundance was higher after mines had been closed
and the land reclaimed. The use of human made ponds and drainage systems suggested that
populations could fully recover. However, future studies should address how the presence of
pollutants in the reclaim lands might pose a health hazard to the population.
Plastics have not received much attention regarding snakes, but França et al. (2018) re-
moved a plastic balloon from the esophagus and stomach of a Helicops angulatus it was nota-
ble that no other prey items were in the digestive tract.

Fishing bycatch

Seasnakes are vulnerable to trawler fishing techniques and they are an unfortunate by-
catch of these fishing practices. Seasnakes have been commercially exploited since 1934
(Dunson, 1975) in the Philippines, where protection laws have been necessary to avoid their
overexploitation. However, seasnakes have been harvested in several countries (Australia,
Japan, Taiwan China, Thailand, and Vietnam) directly or as a by-catch product of fishing.
Despite some countries (eg. Australia) reporting snake by-catch and having introduced
species licensing requirement to collect seasnakes, many other countries continue to avoid
such documentation and the overall impact remains worryingly unknown. Rasmussen et al
(2011) suggest that the main threat to seasnakes are indifferences to conservation issues in
local communities. Education can solve this problem. In addition, they argue that the limited
knowledge of marine snake natural history and biology in Asia continues to be a problem.
Thus the main threat to seasnakes is the cultural indifference to conservation issues and, the
subsequent commercial exploitation. Gathering more information on species biology is piv-
otal, especially in poorly studied regions such as Asia. Data on breeding cycles, growth rates,
sexual maturity, population estimates, and accurate taxonomy and systematic knowledge are
needed to fully protect species.
The effects of by-catch on the seasnakes are almost unknown – the exceptions are the
Philippines and Australia (Milton 2001; Tonks et al., 2008; Courtney et al., 2010, Wassenberg
et al., 1994). In Australia, between Koolan Island and Cape York (northern Australia) circa
80,000 snakes were reported to have been captured by trawlers in 1990 (Ward 1996), while
Wassenberg et al. (1994) estimated 70,000-165,000 were by-catch from commercial prawn
trawlers in the Gulf of Carpentaria in 1991, with a 60% survival. Later work by Wassenberg et

585
 33. Conservation

al. (2001) estimated that the overall mortality of trawl-caught snakes was 48.5%, mostly from
injuries while trawling. Analysis of the species of conservation interest of seasnake data indi-
cated that for 2003, Queensland trawlers incidental catch an annual maximum of 4,840 (year
2003) an annual minimum of 1,640 (in 2006). Between 2003 and 2008, a total of 17,640 snakes
were reported, with 5% mortality and 2% injury (Courtney et al 2010). Currently, the impact
of prawn fishing on seasnake populations in northern Australia has declined substantially
with the reduction in the size of the fleet and the fishing effort since 2004 (Milton et al. 2008).
Courtney et al (2010) attempted to estimate a by-catch of the Queensland East Coast
Trawl Fishery based sixty-seven trawl fishers in a voluntary study. Their data collected 3,910
snakes belonging to 12 species (Lapemis and Aipysurus mostly) from 8,289 trawls between
2003-2007. Using this data they extrapolated the estimated mean catch at 105,000 per year.
About 8.5% of snakes were reported dead, and about 17.4% dying shortly after, giving a com-
bined incidental mortality rate of 25.9%. Reef-associated fishing for Redspot King Prawns
accounted for most of the seasnake by-catch, with mortality at 58.9% with 84.5 %.
A study of Queensland’s trawl fishery evaluated the effects of three different bycatch re-
duction devices (BRDs) on the catch rate of seasnakes, as well as prawns in 2006. The fisheye
and square mesh codend BRDs significantly reduced the catch rate of seasnakes without in-
curring any significant reduction in targeted (≥ 20 mm carapace length) prawn catch rates.
Seasnake by-catch was approximately 60% lower than control nets. Additionally, the distances
of the BRDs installed from the codent drawstring significantly affect effectiveness of avoiding
bycatch, 50 m is recommended. Reports of by-catch reductions of 87% have been recorded
when the device was installed 70 m from the drawstring (Raudzens, 2007). Similarly, other
studies have concluded that the Yarrow fisheye BRD reduced the weight of small fish bycatch
and seasnakes by a mean of 22% and 43.3%, respectively, with no loss of tiger prawn catch
(Heales et al. 2008). Similar results were observed by Milton et al., (2009), where using a con-
trol versus treatment nets in 1,365 trawls, 43% of seasnake catches were reduced with Fisheye
BRD positioned less than 70 m for the codent. They also concluded that using ‘popeye’ Fish-
box BRD reduced seasnake catch by 85%. They argue that a BRD positioned within 70 meshes
of the codend could reduce the seasnake catch and mortality.
Fishing by-catch of snakes is not only present in marine snakes. Thousands of freshwa-
ter snakes are killed yearly (e.g. Natrix tessellata) in the nets set by poachers, notably gravid
females, raising population viability concerns and causing population declines (Sterijovski et
al., 2014).

Trade

Commercial trade is a major driver of the over-exploitation of marine and freshwater

snakes. Nguyen et al (2014) inspected several squid fishing vessels in the Gulf of Thailand
from 2008-2013. Fishermen trades indicated that Lapemis curtus and Hydrophis cyanocinctus
constituted more than 85% of the total seasnake biomass harvested, and other six species made
up the remaining portion of the catch: Acalyptophis peronii, Aipysurus eydouxii, Hydrophis
atriceps, H. belcheri, H. lamberti, and H. ornatus. In Vietnam, seasnakes are not protected, and
their harvest is legal as the catching of seasnakes is regulated by the United Nations Conven-
tion on the Law of the Sea, Section II, Conservation and Management of the Living Resources
of the High Seas, that is dependent on domestic laws. Domestically, seasnakes are not protect-
ed, and their harvest is completely within the legal framework of Vietnam. Market demand
has increased in recent years, while in the past seasnake bycatch in the province of Quang Ngai
was discarded due to fear of bites and a lack of market. In 2011 their price was US$10–35/kg.
Overall, Nguyen et al (2014) estimated average biomass of seasnakes removed from the Gulf
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Aquatic Snakes

of Thailand via Song Doc and Khanh Hoi at circa 82 tones, about 225,500 specimens per year.
On the South China Sea coast, the total biomass of seasnakes harvested through southern
Vietnam was no less than 100 tons (more than 275,000 specimens) per year.
Seasnake meat is used in products such as soup blood mixed with alcohol to improve
health, various organs are used in elixirs, and dried skins from whole seasnakes are added to
the wine (Nguyen et al., 2014). Seasnakes are also reported to be a source of rheumatism,
used to treat various joint, bone, and mental health conditions (Nguyen et al., 2014). The
causes of declining seasnakes in the Gulf of Thailand since 2009 are not clear but include
overharvesting and anthropogenic disturbances such as trawling and pollution (Nguyen et
al., 2014). Auliya (2011) reported a tannery in West Malaysia harvesting 6,000 individuals
per month.
The United Nations Environment World Conservation Monitoring Centre (UNEP-WC-
MC) (2018) categorized three species under consideration for examination for EU trade
imports (Appendix D) under CITES convention for the period 2007-2016. In this report
Subsessor bocourti was used for skin trade and human consumption and as crocodile food
in Cambodia. Captive breeding farms producing aquatic snake species, including S. bocourti
were reported in the Mekong Delta, with 174 government registered farms and circa 80,000
individual snakes held in farms in Vietnam as of 2014 (Aust, 2015). In addition, this species
is also found in reptile shops in Vietnam.
Homalopsis buccata was listed in Annex D of the EU Wildlife Trade Regulations on
01/06/1997 with the annotation that the listing “applies only to live specimens; whole or sub-
stantially whole, dead specimens; and whole or substantially whole skins, raw or tanned.”
Direct imports of Homalopsis buccata comprised circa 3.5 million skins. Skins were predom-
inantly imported from Thailand (49 per cent), Indonesia (38 per cent) and Singapore (12 per
cent). Import of skins to the EU-28 increased in 2007-2015. Indirect trade to the EU-28 pre-
dominantly comprised 725,405 skins, of which 93 per cent were reported without a purpose
or source code and the remainder were wild-sourced for commercial purposes. Most skins
indirectly imported by the EU-28 originated in Indonesia and were predominantly re-export-
ed via Singapore.
Brooks et al. (2010) commented that, the pattern of demand and how it responds to
changes in supply is poorly known. In his region, mostly Enhydris enhydris, Enhydris longi-
cauda, Homalopsis buccata, Erpeton tentaculatum, Fowlea piscator, Cylindrophis ruffus, Sub-
sessor bocourti are heavily fished (Brooks et al. 2008). In Cambodia, the largest driver of snake
exploitation is the domestic trade in snakes as crocodile food. Brooks et al (2010) estimate
that farmed crocodiles consume between 2.7 and 12.2 million snakes yearly. This over-ex-
ploitation of snakes is the result of a progressive decline in the market price of crocodile skin
since 2003, and preference for snakes helps maintain the farms functioning, and snake food
is highly dependent of other substitute available resources such as fish.
Other Asian countries, such as China sell snakes such as Sinonatrix annularis, some En-
hydris spp. and Homalopsis in large numbers at the markets and restaurants (Zhou & Jiang,
2005). In western central Africa, Grayia ornata plays an important role in local culture food,
as well as in magic and traditional medicine (Pauwels et al., 2002). Similarly, in Brazil, among
other snakes, Eunectes mirunus is commercialized in outdoor markets, and they are used
medicinal and for magic–religious purposes (Alves and Filho, 2007).

Steps forward towards conservation

The status of most snake populations (mainly marine species) remains data deficient.

587
 33. Conservation

Lack of knowledge in most areas, especially in Southeast Asia, suggests that there is no
comparative means to address the status of populations and if species and populations are
under extinction risk. Only through monitoring and controlling by-catch and the commer-
cial catch of seasnakes can a sustainable yield be achieved. Efforts should focus on govern-
mental funding to research the importance of the most harvested biological groups and
assess their populations' status in different regions. Habitat destruction and pollution in
rivers, estuaries, mangroves, and boat traffic are causing a reduction in population in Asia
and Australia. Breeding biology remains data deficient for most species, and breeding areas
and periods are lacking from scientific literature, key points to manage and protect the pop-
ulations. Taxonomical data is similarly deficient, and few studies barcoding and identifying
species. Genetic divergence between populations to assess key lineages worth preserving for
future genetic enrichment of other less genetically rich populations is critical to address the
conservation genetic management of species, especially those more harvested populations.
Genetic diversity combined with taxonomical identification will elucidate pivotal informa-
tion to focus harvesting on the more common species and protect other less common ones.
The most accountable and destructive practice of seasnake mortality is trawling fishing,
and evidence indicates that the distance BRDs are installed from the codent drawstring and
the type of net, significantly reduces bycatch or seasnakes, not affecting market table. The
success of the bycatch reduction is dependent on the area (e.g., reefs) and targeted species
(e.g., prawns) to harvest, and only specific studies conducted in different countries and re-
gions through voluntary work of trawl vessels in different regions can help elucidate the
methods that best suit different fishing industries. For example, knowing the size of adult
seasnakes in fishing grounds can help determine the size of the net mesh, as mortality is
higher in larger seasnakes than in smaller individuals, and increased mesh sizes that do not
compromise target harvest could result in a significant reduction of large seasnake by-catch
that likely constitutes gravid females. Trawling vessels or fishing docs would be advanta-
geous with the presence of expert specialists (Milton et al., 2009) to conduct species counts
and sex ratios as well as estimation of gravid females and breeding seasons (Fry et al., 2001).
Such simple data would estimate population reproductive cycles and areas and could help
establish fishing bands or a reduction in licenses throughout breeding seasons. According to
Elfes et al.´s (2013) study on the distribution of seasnakes, the most critical areas identified
as needing basic research are Southeast Asia and peninsular Malaysia, where the greatest
concentration of Data Deficient species are found. They comment that the Data Deficient
status probably reflects on high species diversity in the region, but extremely limited surveys
have resulted in hardly any information in species ecology, abundance, and distribution.
Future surveys should be conducted in the region to solve this lack of knowledge of these
species.
Local awareness is important to reduce by-catch mortality. For example, in India, fish-
ermen kill seasnakes intentionally when encountered as they are believed to threaten the
local community. A report in 2015 showed that over only two consecutive days between
around 60–80 dead seasnakes lay scattered over 20–30 meter stretch on shore due to such
practice (Jamalabad 2015). At a regional scale, the impact on unnecessary seasnake mortal-
ity could make essential differences to the populations.
Species-based conservation programs are challenged in socio-economic deprived re-
gions where water snakes are regularly harvested such as in Tonle Sap Lake in Cambodia
for the last 2 to 3 decades (Stuart et al. 2000). Brooks et al. (2008) concluded that resource
decline, along with the high and potentially increasing levels of poverty, are the major con-
cerns fishing in the Tonle Sap Lake. They argue that conservation strategies that focus on

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Aquatic Snakes

589
 33. Conservation

strengthening, rather than eroding the adaptive capabilities of resource users to cope with
uncertainty, should be supported by increasing rather than restricting access. The nature of
such markets, consumer preferences, and their drivers suggest that protection of snakes to
their breeding season is likely to be the most effective tools for their conservation (Brooks et
al., 2010)
Addressing effective future conservation measures for freshwater aquatic snakes is chal-
lenging. Their global distribution and the vast amounts of habitats they inhabit (swamps,
marshes, lakes, rivers, estuaries, seas, etc) makes general measures unviable. In addition, the
conservation and protection of freshwater biodiversity face challenges influenced by the to-
pology of the biota, such as upstream drainage networks, surrounding lands, riparian zones
and downstream reaches, making conservation management and delimitation of such areas
complex. The presence of species throughout continents and their countries implies differ-
ent governmental legislations to be implemented and many challenging management actions
with needs for “reconciliation ecology,” thus the adoption of biodiversity protection and fresh-
water ecosystem management. Documented threats to global freshwater biodiversity include
overexploitation, water pollution, flow modification, destruction or degradation of habitat,
and invasion by exotic species (Dudgeon et al., 2006), which all have an impact of aquatic
snake populations in different ways. Unfortunately, because freshwater bodies are experienc-
ing higher declines in biodiversity than most terrestrial ecosystems (Sala et al., 2000), and the
human need for water will only continue to increase, species losses at even greater rates are
likely to be expected.
Knowledge of species´ natural history, ecology, and distribution remains critical to
protecting freshwater and marine snakes. However, data remains insufficient to estimate or
predict population extinctions in many world regions accurately. Identifying major hotspots,
especially for freshwater snakes, is pivotal, and will require joint efforts from non-government
for organizations, research institutions, and scientific societies. Strong conservation efforts
should focus on species-rich areas with high biodiversity, such as the Asian oriental region
(Andaman Sea and Southeast Asia) and the neotropics (Pauwels et al., 2008). The conservation
challenges of freshwater (Dudgeon et al., 2006; Hughes, 2019) and marine biota suggests the
need to protect regions with high biodiversity.

On the previous page. Figure 33.1 The Tonle Sap snake harvest is the largest known snake har-
vest in the world. (A) Is Tonle Sap Lake when it is dry. (B) A fisher middleman who buys snakes from
fishers and sells them to another middleman. (C) A collection of snakes going to market. (D) A fisher
in his canoe with gill nets used to collect snakes, (E) Bags filled with snakes. (F) Snakes being cut into
piecies to feed small crocodiles. (G) Crocodiles are the primary reason for the snake harvest - people
buy the snakes to feed their crocodiles that they are raising for skins and meat. Photography by JCM

The following table was compiled from the IUCN online database

590
Aquatic Snakes

Table 1. A list of aquatic and semi-aquatic snakes and their status givnen by the IUCN
Abbreviations used:
CATEGORY
DD (Data Deficient), NT (Not Threatened), LC (Least Concern), VU (Vulnerable), EN (Endangered),
CR (Critically Endangered).
POPULATION TREND

U (Unknown), S (Stable), D (Decreasing), I, (Increasing), Uns (Unspecified)

NPF, (No population severely fragmented), PF (Population severely fragmented), UPD (Unknown
population fragmentation), NAD (No continuous decline of mature adults), UDM (Unknown
decline of mature adults), MAD (Mature adults decreasing).

CONSERVATION ACTIONS
In-place land/water protection: yes O (yes, over the entire range), U (Unknown)

In place research and monitoring: ARP, (Action Recovery Plan), SMS (Systematic monitoring
scheme)

THREATS
L (Local), N (National), I (International)

RISKS
Agriculture and Aquaculture; AP, (Annual & perennial non-timber crops), LR (Livestock farming
& ranching), MA (Marine & freshwater aquaculture), WP (Wood & pulp plantations), AA
(Agriculture and Aquaculture)

Development; TR (Tourism and recreation areas), HU (Housing and urban areas), CI (Commercial
and industrial areas) Human intrusions & disturbance; RA (Recreational activities)

Natural system modification; DW (Dams & water management/use), OM (Other ecosystem

modifications), F (Fire & fire suppression)
Biological resource use; HT (Hunting & trapping terrestrial animals), FH (Fishing & harvesting
aquatic resources), LW (Logging & wood harvesting)

Pollution; AF (Agricultural & forestry effluents), DU (Domestic & urban waste water), IM
(Industrial & military effluents)

Transportation and service corridors; RA (Roads & railroads), SL (Shipping lanes)

Energy production and mining; OD (Oil & gas drilling), RE (Renewable energy), MQ (Mining &
quarrying)

Climate change and severe weather; SF (Storms & flooding) O (Other impacts), HA (Habitat
shifting & alteration), TE (Temperature extremes), D (Droughts)

Invasive and other problematic species, genes & diseases; IAD (Invasive non-native/alien
species/diseases)

591
Aquatic Snakes

592
 33. Conservation

593
Aquatic Snakes

594
 33. Conservation

595
Aquatic Snakes

596
 33. Conservation

597
Aquatic Snakes

598
 33. Conservation

599
Aquatic Snakes

600
 33. Conservation

601
Aquatic Snakes

602
 33. Conservation

603
Aquatic Snakes

604
 33. Conservation

605
Aquatic Snakes

606
 33. Conservation

607
Aquatic Snakes

608
Aquatic Snakes

Literature Cited

609
Aquatic Snakes

Abegg AD, Santos Jr AP, Costa HC, Battilana J, Graboski R, Vianna FS, Azevedo WS, Fa-
gundes NJ, Castille CM, Prado PC, Bonatto SL. 2022. Increasing taxon sampling sug-
gests a complete taxonomic rearrangement in Echinantherini (Serpentes: Dipsadidae).
Frontiers in Ecology and Evolution. 20;10:969263
Ackley JW, Meylan PA.2010. Watersnake Eden: use of stormwater retention ponds by
mangrove salt marsh snakes (Nerodia clarkii compressicauda) in urban Florida. Herpe-
tological conservation and biology 5(1):17-22.
Afrasiab SR, Al-Ganabi MI, Al-Fartosi K. 2011. Snake species new or rare to the herpeto-
fauna of Iraq. Herpetozoa 24:179–181.
Ahmadzadeh F, Mebert K, Ataei S, Hamidi S, Faghiri A, Böhme W. 2011. Some ecological
and biological aspects of grass snake, Natrix natrix (Linnaeus, 1758) in the southern
coastal area of the Caspian Sea. Acta Herpetologica 6(2):209–221.
Akani GC, Luiselli L. 2001. Ecological studies on a population of the water snake Grayia
smythii in a rainforest swamp of the Niger Delta, Nigeria. Contributions to Zoology,
70:139–146.
Akat E, Çakici Ö, Dinçaslan YE, Arikan H. 2011. Histochemical and histological investiga-
tions on Duvernoy’s gland in Natrix tessellata (Squamata: Colubridae). Kafkas Univ Vet
Fak Derg 17(2):285-89.
Ahmadzadeh F, Mebert K, Ataei S, Rezazadeh E, Goli LA. Bohme W. 2011. Ecological and
biological comparison of three populations of Dice snakes (Natrix tessellata) from the
southern Caspian Sea coast, Iran. Mertensiella 18:403–414.
Ahnelt H, Romanova T, Klinge A, Böhme W, Fritz U, Asztalos M. 2021. The common grass
snake (Natrix natrix) on Sylt: human–mediated colonization of a North Sea island. Sal-
amandra 57(2):285–290.
Ajtic R, Tomovic L, Sterijovski B, Crnobrnja-Isailovic J, Djordjevic S, Djurakic M, Gol-
ubovic A, Simovic A, Arsovski D, Andjelkovic M, Krstic M, Šukalo G, Gvozdenovic S,
Aïdam A, Michel CL, Ballouard JM, Bonnet X. 2013. Unexpected life history traits in a
very dense population of dice snakes. Zoologischer Anzeiger 252(3):350–358.
Alarcos G, F Flechoso, R Codesal, M Lizana. 2012. Comportamiento trepador de Natrix
maura para capturar un ejemplar adulto de Hyla arborea. Boletín de la Asociación Her-
petológica Española 23(2):43–45.
Alarcos G, F Alvarez-Collado, MF Flechoso, J Madrigal, M Lizana. 2009. Peces exoticos de
la familia Centrarchidae, un peligro para Natrix maura. Boletín de la Asociación Her-
petológica Española 20:95-97.
Albadalejo PV. 2008. Caso de melanismo en Natrix natrix en el Parque Nacional de los Pi-
cos de Europa. Boletín de la Asociación Herpetológica Española 19: 38–39.
Albuquerque ND, De Lema T. 2008. Taxonomic revision of the Neotropical water snake
Hydrops triangularis (Serpentes, Colubridae). Zootaxa 1685:55-66.
Alcock AW, Rogers L. 1902. On the toxic properties of the saliva of certain ‘non-poisonous’
Colubrines. Proceedings of the Royal Society of London 70:446-454.
Alcover JA, J Mayol. 1981. Espècies relíquies d’amfibis i de rèptils a les Balears i Pitiüses.
Bolletí de la Societat d’Història Natural de les Balears 25:151-167.
Alencar LR, Nascimento LB. 2014. Natural history data of a common snake suggest Inter-
populational variation and conservatism in life history traits: the case of Erythrolam-
prus poecilogyrus. The Herpetological Journal 24(2):79-85.

610
Aquatic Snakes

Alexander GJ. 2018. Reproductive biology and maternal care of neonates in southern Afri-
can python (Python natalensis). Journal of Zoology 305(3):141-8.
Alfaro ME, Arnold SJ. 2001. Molecular systematics and evolution of Regina and the tham-
nophiine snakes. Molecular Phylogenetics and Evolution 21(3):408-23.
Alfaro ME, Karns DR, Voris HK, Abernathy E, Sellins SL. 2004. Phylogeny of Cerberus (Ser-
pentes: Homalopsinae) and phylogeography of Cerberus rynchops: diversification of a
coastal marine snake in Southeast Asia. Journal of Biogeography 31(8):1277–1292.
Akat E, Çakici Ö, Dinçaslan YE, Arikan H. 2011. Histochemical and histological investiga-
tions on Duvernoy’s gland in Natrix tessellata (Squamata: Colubridae). Kafkas Univ Vet
Fak Derg 17(2):285–289.
Alfaro ME, Karns DR, Voris HK, Brock CD, Stuart BL. 2008. Phylogeny, evolutionary his-
tory, and biogeography of Oriental–Australian rear-fanged water snakes (Colubroidea:
Homalopsidae) inferred from mitochondrial and nuclear DNA sequences. Molecular
Phylogenetics and Evolution 46(2):576–593.
Alfaro ME. 2002. Forward attack modes of aquatic feeding garter snakes. Functional Ecol-
ogy, 16(2):204-15.
Alfaro ME. 2003. Sweeping and striking: a kinematic study of the trunk during prey
capture in three thamnophiine snakes. Journal of Experimental Biology. 2003 Jul
15;206(14):2381-92.
Allemand R, Boistel R, Daghfous G, Blanchet Z, Cornette R, Bardet N, Vincent P & Hous-
saye A. 2017. Comparative morphology of snake (Squamata) endocasts: evidence of
phylogenetic and ecological signals. Journal of Anatomy 231(6):849–868.
Allison A, Bickford D, Richards S, Torr G. 1998. Herpetofauna and Appendix 15. Herpe-
tofauna species accounts. In: A.L. Mack (ed.), A Biological Assessment of the Lakeka-
mu Basin, Papua New Guinea. RAP Working Papers 9, pp. 58-62 and 157-172. Con-
servation International, Washington, DC.
Álvarez BB, Aguirre R, Céspedez JA, Hernando A, Tedesco ME. 2003. Herpetofauna del
Iberá. In: Fauna del Iberá, p. 99- 178.
Álvarez BS, Garduza FP, Ojeda AM, Plata VM, Mondragón SL, Charruau P. 2023. Bird-Her-
petofauna Interactions in the Usumacinta River Basin, Mexico. Revista Latinoamericana
de Herpetología 6(01):131-9.
Alves RRN, Filho GAP. 2007. Commercialization and use of snakes in North and Northeast-
ern Brazil: implications for conservation and management. Biodiversity Conservation
16: 969-985.
Amr ZS, Mebert K, Hamidan N, Baker MA, Disi A. 2011. Ecology and conservation of the
Dice snake (Natrix tessellata) in Jordan. Mertensiella 18:393-401
An S, Li H, Guan B, Zhou C, Wang Z, Deng Z, Zhi Y, Liu Y, Xu C, Fang S, Jiang J. 2007. Chi-
na’s natural wetlands: past problems, current status, and future challenges. AMBIO: A
Journal of the Human Environment 36(4):335-42.
Anderson J.1899. Report on the mammals, reptiles, and batrachians, chiefly from the Mer-
gui Archipelago, collected for the Trustees of the Indian Museum. Zoological Journal of
the Linnean Society 21(133-135):331–350.
Andjelković M, Tomović L, Ivanovic A. 2016. Variation in skull size and shape of two snake
species(Natrix natrix and Natrix tessellata). Zoomorphology 135(2):243-53.
Andrén C, Nilson G. 1981. Gotlands reptile och amfibier. Fauna och Flora 76:105–118.
611
Aquatic Snakes

Annandale N. 1916. Preliminary report on the fauna of the Tale Sap or inland sea of Sing-
gora. Journal of the Natural History of the Siam Society 2:90-103.
Apesteguía S, Zaher H. A 2006. Cretaceous terrestrial snake with robust hindlimbs and a
sacrum. Nature 440(7087):1037-40.
Aprea G., Odierna G., Capriglione T., Caputo V. & Guarino F.M., 2000. Analisi cromosom-
ica in tre spe-cie del genere Natrix Duméril (Reptilia, Squamata). Museo Regionale di
Scienze Naturali, Torino, 1999: 419-424.
Arnett JR., Goodwin M. and Teagarden HR. 1992. Trachyboa Peters in Captivity: An
Overview, In Strimple, P.D. and J.L. Strimple (Eds) Contributions in Herpetology. Gr.
Cinn. Herp. Soc. 91-94.
Arrayago MJ, Bea A. 1988. Atlas de citología e histología del aparato reproductor masculino
de los anfibios y reptiles del País Vasco. Sociedad Aranzadi de Estudios Vascos, Cuader-
nos de Sección, Ciencias Naturales 4: 1–112.
Arruda JA. 2006. Natural history of the South American water snake Helicops leopardi-
nus (Colubridae: Hydropsini) in the Pantanal, central Brazil. Journal of Herpetology
40(2):274-9.
Arteaga A. 2020a Golden-bellied Snake (Erythrolamprus albiventris). In: Arteaga A, Busta-
mante L, Vieira J, Guayasamin JM (Eds) Reptiles of Ecuador: Life in the middle of the
world. Available from: [Link]. DOI: 10.47051/LLLF9348
Arteaga A. 2020b Olive Marsh-Snake (Erythrolamprus fraseri). In: Arteaga A, Bustamante
L, Vieira J, Guayasamin JM (Eds) Reptiles of Ecuador: Life in the middle of the world.
Available from: [Link]. DOI: 10.47051/XXDZ6274
Arteaga A. 2021. Peter’s Watersnake (Helicops petersi). In: Arteaga A, Bustamante L, Vieira
J, Guayasamin JM (Eds) Reptiles of Ecuador: Life in the middle of the world. Available
from: [Link]. DOI: 10.47051/UNSB1612.
Asad S, Wilting A, Siku J, Roedel MO. 2020. Possible spatial separation at macro-habitat
scales between two congeneric Psammodynastes species, including observations of fish-
ing behaviour in Psammodynastes pictus. Salamandra. 56(4):411-5.
Ascenso AC, Costa JC, Prudente AL. 2019. Taxonomic revision of the Erythrolamprus re-
ginae species group, with description of a new species from Guiana Shield (Serpentes:
Xenodontinae). Zootaxa. 4586(1):065-97.
Asztalos M, Ayaz D, Bayrakcı Y, Afsar M, Tok CV, Kindler C, Jablonski D, Fritz U. 2021b.
It takes two to tango – Phylogeography, taxonomy and hybridization in grass snakes
and dice snakes (Serpentes: Natricidae: Natrix natrix, N. tessellata). Vertebrate Zoology
71:813–834.
Asztalos M, Glaw F, Franzen M, Kindler C, Fritz U. 2021a. Transalpine dispersal: Italian
barred grass snakes in southernmost Bavaria – This far but no further! Journal of Zoo-
logical Systematics and Evolutionary Research 59:1136–1148.
Asztalos M, Wielstra B, Struijk RPJH, Ayaz D, Fritz U. 2021c. Aliens in the Netherlands:
Local genetic pollution of barred grass snakes (Squamata: Serpentes: Natricidae). Sala-
mandra 57:174–179.
Asztalos M, Schultze N, Ihlow F, Geniez P, Berroneau M, Delmas C, Guiller G., Legentil-
homme J., Kindler C, Fritz U. 2020. How often do they do it? An in-depth analysis of the
hybrid zone of two grass snake species (Natrix astreptophora, N. helvetica). Biological
Journal of the Linnean Society 131:756–773.
612
Aquatic Snakes

Aubret F, A Mangin. 2014. The snake hiss: potential acoustic mimicry in a viper-colubrid
complex. Biological Journal of the Linnean Society 113:1107–1114.
Aubret F, Michniewicz RJ. 2010. Warming up for cold water: influence of habitat type on
thermoregulatory tactics in a semi-aquatic snake. Amphibia-Reptilia 31: 525-531.
Aubret F, Shine R. 2007. Rapid prey‐induced shift in body size in an isolated snake popu-
lation (Notechis scutatus, Elapidae). Austral Ecology 32(8):889-99.
Auffenberg W. 1980. The herpetofauna of Komodo, with notes on adjacent areas. Bulletin
of the Florida State Museum, Biological Sciences 25:39–146.
Auliya MA. 2003. Taxonomy, Life History and Conservation of Giant Reptiles in West
Kalimantan (Indonesia, Borneo). PhD Dissertation. Mathematisch-Naturwissenschaft-
lichen Fakultät der Rheinischen Friedrich-Wilhelms-Universität Bonn, 513 pp.
Auliya MA. 2006. Taxonomy, life history and conservation of giant reptiles in West Kali-
mantan (Indonesian Borneo). Natur und Tier.
Auliya, M. 2011. Lapemis curtus (Serpentes: Elapidae) harvested in West Malaysia. Sea
Snake Specialist Group Newsletter 6–8
Aust P. 2015. An assessment of the commercial production of CITES‐listed snake species
in Viet Nam and China. AC28 Inf. 1. IUCN SSC Boa & Python Specialist Group, Jo-
hannesburg, South Africa. 1-44 pp
Ávila RW, Ferreira VL, Arruda JA. 2006. Natural history of the South American water
snake Helicops leopardinus (Colubridae: Hydropsini) in the Pantanal, central Brazil.
Journal of Herpetology 40(2):274-80.
Avolio C, Shine R, Pile A. 2006. Sexual dimorphism in scale rugosity in sea snakes (Hy-
drophiidae). Biological Journal of the Linnean Society, 89(2), 343–354. doi:10.1111/
j.1095-8312.2006.00678.x
Avolio C, Shine R, Pile A. 2006. Sexual dimorphism in scale rugosity in sea snakes (Hy-
drophiidae). Biological Journal of the Linnean Society, 89(2), 343–354. doi:10.1111/
j.1095-8312.2006.00678.x
Ayres C. 2012. Natrix maura (Viperine snake). Diet: scavenging. Herpetological Review
43(2):346-347.
Babonis LS, Miller SN, Evans DH. 2011. Renal responses to salinity change in snakes with
and without salt glands. Journal of Experimental Biology 214(13):2140-56.
Baha El Din S. 2011. Distribution and recent range extension of Natrix tessellata in Egypt.
Mertensiella 18:401-03.
Baier F, Sparrow D, Wiedl H–J. 2009. The Amphibians and Reptiles of Cyprus. Edition
Chimaira, Frankfurt, Germany, 362 pp.
Bailey JR, Thomas RA, Silva NJ 2005. A revision of the South American genus Tham-
nodynastes Wagler, 1830 (Serpentes, Colubridae, Tachymenini). I. Two new species
of Thamnodynastes from Central Brazil and adjacent areas, with redefinition of and
neotype designation for Thamnodynastes pallidus (Linnaeus, 1758). Phyllomedusa, 4:
83–101.
Bailey JR. 1967. The synthetic approach to colubrid classification. Herpetologica 23 (2):
155-161.
Bailón S. 1991. Amphibiens et reptiles du Pliocène et du Quaternaire de France et d’Es-
pagne: Mise en place et évolution des faunes. Thèse de doctorat, Univ. Paris VII.
Baker RA, Gawne TJ, Loop MS & Pullman S. 2007. Visual acuity of the midland banded
613
Aquatic Snakes

water snake estimated from evoked telencephalic potentials. Journal of Comparative

Physiology A: Neuroethology, Sensory, Neural, and Behavioral Physiology, 193(8):865–
870.
Bakiev A, Kirillov A, Mebert K. 2011. Diet and parasitic helminths of Dice snakes from the
Volga Basin, Russia. Mertensiella 18:325-30
Ballinger R, Lynch J, Cole PH. 1979. Distribution and Natural History of amphibians and
reptiles in western Nebraska with ecological notes on the herptiles of Arapaho Prairie.
Prairie Nat. 11 :65-74.
Bannikov AG, Darevskii IS, Rustamov AK. 1971. Amphibians and Reptiles of the SSSR.
Publ. Mysl, Moscow [in Russian].
Barata M, Harris DJ, Castilho R. 2008. Comparative phylogeography of northwest African
Natrix maura (Serpentes: Colubridae) inferred from mtDNA sequences. African Zool-
ogy 43(1):1–7.
Barbault R. Les traits quantitatifs du peuplement des Ophidiens. La Terre et la vie.
1970:94-107.
Barbour T, Do Amaral A. Notes on some Central American snakes. Apuntes sobre algunas
serpientes centroamericanas. Occasional Papers. 1924;5:129-32.
Barker DG, Barker TM. 2008. The distribution of the Burmese python, Python molurus
bivittatus. Bulletin of the Chicago Herpetological Society 43(3):33-8.
Barquero MD, Arguedas V. 2019. Biology of snakes of the genus Tretanorhinus: an integra-
tive review. Amphibian & Reptile Conservation 13(2): 227–238.
Barrera Jr. WB, Jaluague JCA, Albaño SDC, Jalbuena GLL. New Distributional Record and
Intra-Specific Variation of Cerberus schneiderii in Iyam River, Lucena City, Quezon,
Philippines. Philippine Journal of Systematic Biology 11(1):29–36.
Barrio-Amorós CL, Manrique R. 2007. Observaciones Sobre Historia Natural de la Ana-
conda (Eunectes murinus, Linnaeus, 1758) en los Llanos de Venezuela. Una Perspectiva
Ecoturística.
Baruah M, Das M, Sengupta S. 2001. Food and feeding of Amphiesma stolatum (Linnaeus
1758). Journal of environmental biology 22(4):315-8.
Bassi N, Kumar MD, Sharma A, Pardha-Saradhi P. 2014. Status of wetlands in India: A
review of extent, ecosystem benefits, threats, and management strategies. Journal of
Hydrology: Regional Studies 21-19.
Batchelor DM. 1963. Protective behavior in snakes. Malayan Nature Journal 14:131–132.
Batista SF, Scartozzoni RR, Marques OA. 2020. Food habits and substrate use by the
South American xenodontine snake Erythrolamprus frenatus, with comments on its
brightly-coloured venter. Herpetology Note 13:405-9.
Bauer AM, DeBoer JC, Taylor DJ. 2017. Atlas of the Reptiles of Libya. Proceedings of the
California Academy of Sciences 64(8):155–318.
Bauer AM, Jackman T. Global diversity of lizards in freshwater (Reptilia: Lacertilia). In:
Freshwater Animal Diversity Assessment 2007 (pp. 581-586). Springer, Dordrecht.
Beadle LC. 1981. The inland waters of tropical Africa: an introduction to tropical limnolo-
gy. Longman Group Limited.
Beddek M. 2017. Déficit de connaissances de la biodiversité et biologie de la conservation
: le cas de l’herpétofaune d’Algérie. Biodiversité et Ecologie. Université Montpellier,
France.

614
Aquatic Snakes

Bedriaga J. von 1882. Die Amphibien und Reptilien Griechlands. Moscow, 195 pp.
Beebe W. 1946. Field notes on the snakes of Kartabo, British Guiana, and Caripito, Venezue-
la. Zoologica 31: 11–52.
Bełcik A, Różycki M, Korpysa-Dzirba W, Marucci G, Fafiński Z, Fafińska P, Karamon J, Ko-
chanowski M, Cencek T, Bilska-Zając E. 2022. Grass snakes (Natrix natrix) as a reservoir
of Alaria alata and other parasites. Pathogens 11:156.
Bellini GP, Arzamendia V, Giraudo AR. 2013. Ecology of Thamnodynastes hypoconia in sub-
tropical–temperate South America. Herpetologica 69(1):67-79.
Bellini GP, Giraudo AR, Arzamendia V. 2014. Comparative ecology of three species of
Thamnodynastes (Serpentes, Dipsadidae) in subtropical-temperate South America. The
Herpetological Journal 24(2):87-96.
Ben Hassine J., Escoriza D. 2014. Bufo spinosus in Tunisia: new data on occurrence, parasit-
ism and tadpole morphology. Herpetological Bulletin, 127: 22-32.
Bendel P. 1997. Zur Physiologie, Morphometrie und Populationsökologie der Würfelnatter
Natrix tessellata am Alpnachersee. MS thesis, Zoological Museum, University of Zurich,
Switzerland.
Bendel P. 2001. Zur Physiologie der Wurfelnatter Natrix tessellata Laurenti 1768 am Alp-
nachersee. Naturforschende Gesellschaft Ob- und Nidwalden (NAGON), Grafenort,
Schweiz:161–75.
Beolens, Bo; Michael Watkins, and Michael Grayson 2011. The Eponym Dictionary of Rep-
tiles. Johns Hopkins University Press, Baltimore, USA.
Berec M, Moravec J, Fric ZF. 2015. Natrix natrix (Linnaeus, 1758) – užovka obojková. Pp.
337–361. In: Fauna ČR. Plazi/Reptilia. Moravec J. (Ed.), Praha, Czech Republic, Aca-
demia, 532 pp.
Bergman RAM. 1951. The anatomy of Homalopsis buccata. Proceeding of the Koninklijke
Nederlandse Akademie van Wetenschappen Series C 54(5):511–524.
Bergman RAM. 1960. The anatomy of some Homalopsinae. Biologisch Jaarboek Uitgegeven
Door Het Koninklijk Natuurwetenschappelijk Genootschap Dodonaea 1960:119-139.
Bernarde PS, Moura-Leite JC, Machado RA, Kokobum MNC. 2000. Diet of the colubrid
snake, Thamnodynastes strigatus (1858) from Paraná state, Brazil, with field notes on an-
uran predation. Rev Bras Biol 60(4): 695-699.
Bérnils RS, Batista MA, Bertelli PW. 2001. Cobras e lagartos do Vale: levantamento das
espécies de Squamata (Reptilia, Lepidosauria) da Bacia do Rio Itajaí, Santa Catarina,
Brasil. Revista de Estudos Ambientais 3(1): 69-79.
Bernstein JB, Murphy JC, Voris HK, Brown RM, Ruane S. 2021. Phylogenetics of Mud
Snakes (Squamata: Serpentes: Homalopsidae): A Paradox of Both Undescribed Diversity
and Taxonomic Inflation. Molecular Phylogenetics and Evolution: 160:107109.
Bernstein JM. A Multifaceted Approach to Investigating Diversity, Systematics, and Bioge-
ography of Old World Mud Snakes (Serpentes: Homalopsidae) (Doctoral dissertation,
Rutgers The State University of New Jersey, Graduate School-Newark).
Bernstein, JM, Voris HK, Stuart BL, Phimmachak S, Seateun S, Sivongxay N, Neang T.,
Karns DR, Andrews HL, Osterhage J. Phipps EA. 2022. Undescribed Diversity in a
Widespread, Common Group of Asian Mud Snakes (Serpentes: Homalopsidae: Hypsis-
copus). Ichthyology & Herpetology, 110(3), pp.561-574.
Bernstein JM, de Souza H, Murphy JC, Voris H, Brown R, Myers E, Harrington S, Shanker

615
Aquatic Snakes

K, Ruane S, 2023a. Phylogenomics of Fresh and Formalin Specimens Resolves the Sys-
tematics of Old World Mud Snakes (Serpentes: Homalopsidae) and Expands Biogeo-
graphic Inference. Bulletin of the Society of Systematic Biologists, 2(1), pp.1-24.
Bernstein JM, Voris HK, Stuart BL, Karns DR, McGuire JA, Iskandar DT, Riyanto A,
Calderón-Acevedo CA, Brown RM, Gehara M, Soto-Centeno JA. 2023b. Integrative
Methods Reveal Multiple Drivers of Diversification in Rice Paddy Snakes. Research
Square
Berry PY, Lim GS. 1967. The breeding pattern of the puff-faced water snake, Homalopsis
buccata Boulenger. Copeia 1967:307–313.
Beshkov V, Dushkov D. 1981. Materials on the batrachophagy and herpetophagy of snakes
in Bulgaria. Ecology 9:43–50.
Beutelspacher-García PM. 2015. Thamnophis proximus. Behavior. Nature Notes. Mesoamer-
ican Herpetology 4:531-532.
Bezuijen MR, Vinn B, Send L. 2009. A collection of amphibians and reptiles from the Me-
kong River, north-eastern Cambodia. Hamadryad 34:135–164.
Bhaisare D, Pelling E. 2015. Trimeresurus malabaricus (Malabar pit viper): Diving behaviour
and underwater apnoea duration. Herpetological Bulletin 134:33-4.
Bhosale H, Joshi D. 2014. Notes on distribution, natural history and habitat use of a Col-
ubridae snake, Rhabdops olivaceus (Beddome, 1863). Russian Journal of Herpetology
21(3):166-8.
Bhosale HS, Thite V. 2019. Death feigning behavior in Large-eyed False Cobra Pseudox-
enodon macrops (Blyth, 1854) (Squamata: Colubridae). Russian Journal of Herpetolo-
gy 20: 190–192.
Bhupathy S, Ramesh C. 2010. Breeding biology of Python molurus molurus in Keoladeo
national park, Bharatpur, India. The Herpetological Journal 20(3):157-63.
Bilcke J, Herrel A, Aerts P. 2007. Effect of prey-and predator size on the capture success of
an aquatic snake. Belgian Journal of Zoology. 2007 Jul 1;137(2):191
Biodiversitas. 2007. Revisão das listas das espécies da flora e da fauna ameaçadas de ex-
tinção do estado de Minas Gerais. Relatório final, vol. 3. Electronic database accessible
at 0p. Belo Horizonte.
Bird MI, Taylor D, Hunt C. 2005. Palaeoenvironments of insular Southeast Asia during the
Last Glacial Period: a savanna corridor in Sundaland?. Quaternary Science Reviews 24:
2228–2242.
Bjelica V, Anđelković M, Lakušić M, Maričić M, Arsovski D, Tomović L, Golubović A. 2023.
A dicey situation: capture behaviours in free-ranging dice snakes. Behavioral Ecology
and Sociobiology 77:48.
Blain H-A, S Bailon, J Agusti. 2008a. Amphibians and squamate reptiles from the latest early
Pleistocene of Cueva Victoria (Murcia, southeastern Spain, SW Mediterranean): paleo-
biogeographic and paleoclimatic implications. Geologica Acta 6:345-361.
Blain H-A, S Bailon, G Cuenca-Bescos. 2008b. The Early-Middle Pleistocene palaeoenviron-
mental change based on the squamate reptile and amphibian proxies at the Gran Dolina
site, Atapuerca, Spain. Palaeogeography, Palaeoclimatology, Palaeoecology 261:177-192.
Blain H-A, Bailon S, Cuenca-Bescos G, Bennasar M, Rofes J, López-García JM, Huguet R,
Arsuaga JL, Bermudez de Castro JM, Carbonell E. 2010. Climate and environment of
the earliest West European hominins inferred from amphibian and squamate reptile
616
Aquatic Snakes

assemblages: Sima del Elefante Lower Red Unit, Atapuerca, Spain. Quaternary Science
Reviews 29:3034-3044.
Blain H-A, CP Gleed-Owen, JM López-García, JS Carrión, R Jennings, G Finlayson, C Fin-
layson, F Giles-Pacheco. 2013. Climatic conditions for the last Neanderthals: Herpeto-
faunal record of Gorham’s Cave, Gibraltar. Journal of Human Evolution 64:289-99.
Blanvillain G, Lorch J, Joudrier N, Bury S, Cuenot T, Franzen,M, Hoyt JR. 2022. Hotspots
for snake fungal disease across Europe are maintained by host and pathogen identity.
bioRxiv 2022-11.
Blais BR, Hefty KL, Dutt NR. 2021. Thamnophis elegans vagrans (Wandering Gartersnake).
Diet. Herpetological Review 52(4):883-884.
Blake DK. 1960. Bite from an Amplorhinus multimaculatus at Umtali. The Journal of the
Herpetological Association of Rhodesia 9(1):17.
Blanford WT. 1881. On a collection of reptiles and frogs chiefly from Singapore. Proceed-
ings of the Zoological Society of London 1881:215–226.
Blanke I, Borgula A, Brandt T. 2008. Verbreitung, Ökologie und Schutz der Ringelnatter (Na-
trix natrix LINNAEUS, 1758). Mertensiella 17:1–312.
Bleeker P. 1857a. Berigt omtrent eenige reptiliën van Sumatra, Borneo, Batjan en Boero.
Natuurkd Tijdschr Nederl Ind 13:470–475.
Bleeker P. 1857b. Opsomming der soorten van Reptiliën, tot dus verre van het eiland Java
beken geworden. Natuurkd Tijdschr Nederl Ind 14:235–246.
Bleeker P. 1860. Over de Reptiliën-Fauna van Amboina. Natuurkundig Tijdschrift voor
Nederlandsch Indië 22:39–43.
Blosat B. 2008. Population status, threats and protection of the Grass Snake, Natrix natrix
cypriaca (Hecht, 1930) on Cyprus. Mertensiella 17:246–271.
Blumgart D, Dolhem J, Raxworthy CJ. 2017. Herpetological diversity across intact and
modified habitats of Nosy Komba Island, Madagascar. Journal of Natural History 51(11-
12):625-42.
Blyth E. 1855. Collections presented by Capt. Beddome and Mr. Theobald. The Journal of
the Asiatic Society of Bengal, Calcutta 24:187.
Boettger O. 1895. Liste der reptilien und batrachier der insel Halmheira nach den sammlu-
ngen Prof. Dr. W. Kükenthals. Zoolgischer Anzeiger 18:129–138.
Bogert CM, Rowley JS. 1968. A new genus and species of dwarf boa from southern Mexi-
co. American Museum Novitates; no. 2354.
Bogdanov OP. 1960. Fauna of the Uzbek SSR. Amphibians and Reptiles. Tashkent, Uzbeki-
stan, vol.1:260 pp.
Bohaty SM, Zachos JC. 2003. Significant Southern Ocean warming event in the late middle
Eocene. Geology. 2003 Nov 1;31(11):1017-20.
Böhme W, Ziegler T. 2009. A review of iguanian and anguimorph lizard genitalia (Squa-
mata: Chamaeleonidae; Varanoidea, Shinisauridae, Xenosauridae, Anguidae) and their
phylogenetic significance: comparisons with molecular data sets. Journal of Zoological
Systematics and Evolutionary Research, 47(2), pp.189-202.
Böhm M, Collen B, Baillie JE, Bowles P, Chanson J, Cox N, Hammerson G, Hoffmann M,
Livingstone SR, Ram M, Rhodin AG. et al. 2013. The conservation status of the world’s
reptiles. Biological Conservation. 157:372-85.
Boi H. 1827. Bemerkungen über Merrem’s Versuch eines Systems der Amphibien. Erste
617
Aquatic Snakes

Lieferung: Ophidier. Isis von Oken 20: 508–566 [columns].

Bolaños R, Cerdas L, Taylor RT. 1975. Color patterns and venom characteristics in Pelamis
platurus. Copeia 1974: 909–91.
Bonaparte CLP. 1845. Specchio generale dei sistemi erpetologico, anfibiologico ed ittiologi-
co. Atti Riunione degli Scienziati Italiani 6:376–378.
Bonnet X, Brischoux F. 2008. Thirsty sea snakes forsake refuge during rainfall. Austral
Ecology 33(7):911–921.
Bonnet X, Brischoux F, Bonnet C, Plichon P, Fauvel T. 2014. Coastal Nurseries and Their
Importance for Conservation of Sea Kraits. PLoS ONE, 9(3), e90246. doi:10.1371/
[Link].0090246.
Bonnet X, Ineich I, Shine R. 2005. Terrestrial locomotion in sea snakes: the effects of sex
and species on cliff-climbing ability in sea kraits (Serpentes, Elapidae, Laticauda). Bi-
ological Journal of the Linnean Society. 2005 Jul 20;85(4):433-41.
Bonnet X, Pearson D, Ladyman M, Lourdais O, Bradshaw D. ‘Heaven’ for serpents? 2002.
A mark–recapture study of tiger snakes (Notechis scutatus) on Carnac Island, Western
Australia. Austral Ecology 27(4):442-50.
Bonnet X., F. Brischoux, D. Pearson, and P. Rivalan. 2009. Beach rock as a keystone habitat
for amphibious sea snakes. Environmental Conservation 36:62–70.
Bons J. 1967. Recherches sur la biogéographie et la biologie des amphibiens et des reptiles
du Maroc. Université de Montpellier.
Bons J, Geniez P. 1996. Anfibios y Reptiles de Marruecos (incluido Sahara Occidental. At-
las biogeográfico. Asociación Herpetológica Española, Barcelona.
Boulenger G.A., 1913. The snakes of Europe. Methusen & Co. Ltd, London, 269 pp.
Boulenger GA. 1890. The Fauna of British India, Including Ceylon and Burma. Reptilia
and Batrachia. Taylor and Francis, London.
Boulenger GA. 1891. Remarks on the herpetological fauna of Mount Kina baloo, North
Borneo. Ann. Mag. nat. Hist. (6) 7:341-345.
Boulenger GA. 1893. Catalogue of the Snakes in the British Museum, Vol. 1–3. Trustees
of the British Museum, London.
Boulenger GA. 1897. A catalogue of the reptiles and batrachians of Celebes with special
reference to the collections made by Drs P & F Sarasin in 1893–1896. Proceedings of
the Zoological Society of London 1897:193–237 [p228–237: Batrachia].
Boulenger GA. 1903. Report on the batrachian and reptiles. Fasciculi Malayenses Zoology
1:164.
Boulenger GA. 1904. Descriptions of new frogs and snakes from Yunnan. The Annals and
Magazine of Natural History; Zoology, Botany, and Geology Series 7, 13 (74):130-134.
Boulenger GA. 1913. The snakes of Europe. Methusen and Co. Ltd, London, 269 pp.
Bourne J, Hammer E, Howland J. 2021. Thamnophis eques megalops. Predation. Herpeto-
logical Review 52(2):435.
Bourret R. 1934. Notes herpétologiques sur l’Indochine Française IV. Sur une collection
d’ophidiens de Cochinchine et du Cambodge. V. Sure Liopeltis major et ses alliés. Bull.
Instr. Pub. Gen. Hanoi Sept. 1934:13–125 (in print 3-20).
Bowley JL, Willemssens KA, Highley LG. 2020. Thamnophis elegans (Terrestrial Garter-
snake). Extreme Habitat Use. Herpetological Review 51(2):360-361.
Brakles P, Mai TTT, Hguyen TV. 2021. Hebius chapaensis (Sapa Keelback Snake). Diet.
618
Aquatic Snakes

Herpetological Review 52(1):159.

Braña F., 1998. Natrix natrix (Linnaeus, (1758). Pp. 454-466.
in: Salvador, A. (Coord.), Ramos M. A. et al. (Eds.), Fauna Iberica vol. 10: Reptiles. Museo
Nacional de Ciencias Naturales-CSIC, Madrid.
Branch WR. 1998. Field Guide to the Snakes and Other Reptiles of Southern Africa. 3rd
ed. Fully Revised and Updated to Include 83 New Species. Ralph Curtis Books (Sanibel
Island, Florida).
Branch WR. 2018. Snakes of Angola: An annotated checklist. Amphibian & Reptile Conser-
vation. 12(2):41-82.
Branson BA, Baker EC. 1974. An ecological study of the queen snake, Regina septemvtttata
(Say) in Kentucky. Tulane Studies in Zoology and Botany. 18:153-171.
Braz HB, Almeida‐Santos SM, Murphy CR, Thompson MB. 2018. Uterine and eggshell
modifications associated with the evolution of viviparity in South American water
snakes (Helicops spp.). Journal of Experimental Zoology Part B: Molecular and Develop-
mental Evolution 330(3):165-80.
Braz HB, Scartozzoni RR. 2020. Thamnodynastes strigatus (Coastal House Snake). Repro-
duction. Herpetological Review 51(4):880.
Braz HB. 2020. Helicops gomesi. Diet and Reproduction. Herpetological Review 51(4):866-
867.
Brecko J, Vervust B, Herrel A, Van Damme R. 2011. Head morphology and Diet in the Dice
snake (Natrix tessellata). Mertensiella 18:20-30.
Breitfeld HT, Hennig-Breitfeld J, BouDagher-Fadel MK, Hall R, Galin T. 2020. Oligo-
cene-Miocene drainage evolution of NW Borneo: Stratigraphy, sedimentology and
provenance of Tatau-Nyalau province sediments. Journal of Asian Earth Sciences
195:104331.
Bressan PM, Kierulf MCM, Sugieda AM. 2009. Fauna Ameaçada de Extinção do Estado
de São Paulo: Vertebrados. São Paulo: Fundação Parque Zoológico de São Paulo / Sec-
retaria do Meio Ambiente. 645 p.
Breuer H, Murphy WC 2010. Pseudolaticauda semifasciata. [Link]. Re-
trieved 12 January 2020.
Bringsøe H, Suthanthangjai M, Suthanthangjai W & Nimnuam K. 2020. Eviscerated alive:
Novel and macabre feeding strategy in Oligodon fasciolatus (Günther, 1864) eating or-
gans of Duttaphrynus melanostictus (Schneider, 1799) in Thailand. Herpetozoa 33:157–
163.
Brischoux F, Bonnet X, Pinaud D. 2009. Fine scale site fidelity in sea kraits: implications for
conservation. Biodiversity Conservation 18:2473–2481.
Brischoux F, Bonnet X. 2009. Life history of sea kraits in New Caledonia. Zoologia Neo-
caledonica, 7, pp.37-51.
Brischoux F, Briand MJ, Billy G & Bonnet X. 2013. Variations of natremia in sea kraits (Lati-
cauda spp.) kept in seawater and fresh water. Comparative Biochemistry and Physiology
- A Molecular and Integrative Physiology 166(2):333–337.
Brischoux F, Kato A, Ropert-Coudert Y & Shine R. 2010. Swimming speed variation in
amphibious seasnakes (Laticaudinae): A search for underlying mechanisms. Journal of
Experimental Marine Biology and Ecology 394(1–2):116–122.
Brischoux F, Kornilev YV, Lillywhite HB. 2017. Physiological and behavioral responses
619
Aquatic Snakes

to salinity in coastal Dice snakes. Comparative Biochemistry and Physiology Part A:

Molecular & Integrative Physiology 214:13-18.
Brischoux F, Kornilev YV. 2014. Hypernatremia in Dice snakes (Natrix tessellata) from a
coastal population: Implications for osmoregulation in marine snake prototypes. PloS
One 9(3): e92617.
Brischoux F & Shine R. 2011. Morphological adaptations to marine life in snakes. Journal
of Morphology 272(5):566–572.
Brischoux F, Tingley R, Shine R, & Lillywhite HB. 2012. Salinity influences the distribution
of marine snakes: implications for evolutionary transitions to marine life. Ecography
35(11):994–1003.
Brischoux F, Tingley R, Shine R, Lillywhite HB. 2013. Behavioral and physiological
correlates of the geographic distributions of amphibious sea kraits (Laticauda spp.).
Journal of Sea Research, 76, 1–4. doi:10.1016/[Link].2012.10.010.
Broadley DG, 1962. The Herpetofauna of the Inyanga National Park. The Journal of the
Herpetological Association of Rhodesia, 17(1):8-10.
Broadley DG, Cotterill FP. 2004. The reptiles of southeast Katanga, an overlooked ‘hot
spot’. African Journal of Herpetology 53(1):35-61.
Broadley DG. 1984. A review of geographical variation in the African python, Python sebae
(Gmelin). British Journal of herpetology. 1984;6(10):359-67.
Broadley, DG. 1983. Fitzsimons’ Snakes of Southern Africa. Delta Books, Johnannesburg.
Broadley DG. 1987. Caudal autotomy in African snakes of the genera Natriciteres Love-
ridge and Psammophis Boie. The Journal of the Herpetological Association of Africa.
1987 Nov 1;33(1):18-9.
Brongersma LD. 1951. Some notes upon the anatomy of Tropidophis and Trachyboa (Ser-
pentes). Zool Meded. 31:107-24.
Brooks SE, Allison EA, Gill AJ, Reynolds JD 2010. Snake process and crocodile appetites:
Aquatic wildlife supply and demand on Tole Sap Lake, Camdobia. Biological Conserva-
tion 143: 2127-2135.
Brooks SE, Allison EH, Gill JA, Reynolds JD. 2009. Reproductive and trophic ecology of
an assemblage of aquatic and semi-aquatic snakes in Tonlé Sap, Cambodia. Copeia
2009(1):7-20.
Brooks SE, Allison EH, Reynolds JD. 2007. Vulnerability of Cambodian water snakes:
Initial assessment of the impact of hunting at Tonle Sap Lake. Biological Conservation
139(3-4):401–414.
Brooks SE, Reynolds JD, Allison EA, Touch B. 2007. The exploitation of homalopsid wa-
ter snakes at Tonle Sap Lake, Cambodia. Pp. 31–37. In: Homalopsid Snakes. Evolu-
tion in the Mud. Murphy JC (ed.). Krieger Publishing Company, Florida.
Brooks SE, Reynolds JD, Allison EH. 2008. Sustained by snakes? Seasonal livelihood strat-
egies and resource conservation by Tonle Sap fishers in Cambodia. Human Ecology
36(6):835–851.
Brown BC. 1950. An annotated check list of the reptiles and amphibians of Texas. Baylor
University Studies, 1-257.
Brown GP, Shine R, 2004. Maternal nest‐site choice and offspring fitness in a tropical
snake (Tropidonophis mairii, Colubridae). Ecology, 85:1627-1634.
Brown GP, Shine R. 2005. Female phenotype, life history, and reproductive success in
620
Aquatic Snakes

free‐ranging snakes (Tropidonophis mairii). Ecology 86(10):2763-70.

Brown GP, Shine R. 2005. Nesting snakes (Tropidonophis mairii, Colubridae) selectively
oviposit in sites that provide evidence of previous successful hatching. Canadian Jour-
nal of Zoology, 83:1134-1137.
Brown GP, Weatherhead PJ. 2000. Thermal ecology and sexual size dimorphism in north-
ern water snakes, Nerodia sipedon. Ecological monographs 70(2):311-30.
Brown WC, Leviton AE. 1961. Discovery of the snake genus Opisthotropis in the Philip-
pine Islands, with description of a new species. Occasional Papers of the Natural His-
tory Museum of Stanford University (8):1-11.
Brust T, Aldecoa M, Semasko A. 2020. Clonophis kirtlandii (Kirtland’s Snake). Activity. Her-
petological Review 51(3):611.
Burbrink FT, Crother BI. 2011. Evolution and taxonomy of snakes, pp. 19 – 53 in Al-
dridge, RA and Sever, DM. (eds). Reproductive Biology and Phylogeny of Snakes.
Boca Raton, CRC Press.
Burbrink FT, Ruane S, Kuhn A, Rabibisoa N, Randriamahatantsoa B, Raselimanana AP,
Andrianarimalala MS, Cadle JE, Lemmon AR, Lemmon EM, Nussbaum RA. 2019.
The origins and diversification of the exceptionally rich gemsnakes (Colubroidea:
Lamprophiidae: Pseudoxyrhophiinae) in Madagascar. Systematic Biology 68(6):918-
36.
Burbrink FT, Grazziotin FG, Pyron RA, Cundall D, Donnellan S, Irish F, Keogh JS, Kraus
F, Murphy, RW, Noonan B, Raxworthy CJ, Ruane S, Lemmon AR, Lemmon EM, Zaher
H. 2020. Interrogating genomic-scale data for Squamata (lizards, snakes, and amphis-
baenians) shows no support for key traditional morphological relationships. Systematic
Biology 69(3):502–520.
Burić I, Baškiera S. 2014. New record of the Dice snake (Natrix tessellata) from Cres island,
Croatia. Herpetological Bulletin 1:18-19.
Burton A. 2021. Life Lines: Bithynian snake bombs. Frontiers in Ecology and the Environ-
ment:197. doi:10.1002/fee.2332
Butler H, Malone B, Clemann N. 2005. Activity patterns and habitat preferences of trans-
located and resident tiger snakes (Notechis scutatus) in a suburban landscape. Wildlife
Research 32(2):157-63.
Buzás B, Farkas B, Gulyás E, Géczy E. 2018. The sea snakes (Elapidae: Hydrophiinae) of
Fujairah. Tribulus Journal of Emirates Natural History Group Volume 26- 2018. ISSN
69-19
Buzás B, Farkas B, Gulyás E, Géczy E. 2018. The sea snakes (Elapidae: Hydrophiinae) of
Fujairah. Tribulus Journal of Emirates Natural History Group Volume 26- 2018. ISSN
69-19
Cabot J, Olea S. 1978. Observación de una culebra viperina, Natrix maura, en agua marina.
Doñana, Acta Vertebrata 5:107.
Cabral MJ, Almeida J, Almeida PR, Dellinger T, Ferrand de Almeida N, Oliveira ME,
Palmeirim JM, Queiroz AI, Rogado L, Santos-Reis M (Eds.) 2005. Livro Vermelho dos
Vertebrados de Portugal. Instituto da Conservação da Natureza. Lisboa.
Cadle JE. 2003. Colubridae, snakes, In: Goodman, S.M. & Benstead, J.P. (Eds.), The Natu-
ral History of Madagascar. The University of Chicago Press, Chicago and London, pp.
997–1004.
621
Aquatic Snakes

Cafuta V. 2011. First report of melanistic Dice snakes (Natrix tessellata) in Slovenia. Mer-
tensiella 18:442-44.
Caldwell MW, Albino AM. 2001. Palaeoenvironment and palaeoecology of three Creta-
ceous snakes: Pachyophis, Pachyrhachis, and Dinilysia. Acta Palaeontologica Polonica
46, 2,203-218.
Caldwell MW, Lee MS. 1997. A snake with legs from the marine Cretaceous of the Middle
East. Nature. 1997 Apr;386(6626):705-9.
Caldwell MW, Nydam RL, Palci A, Apesteguía S. 2015. The oldest known snakes from the
Middle Jurassic-Lower Cretaceous provide insights on snake evolution. Nature com-
munications. 6(1):1-1.
Camargo IC, Veras JC, Ribeiro S, Kawashita-Ribeiro RA, de Fraga R, Junior AP. 2021.
Sexual dimorphism in the South American water snake Helicops polylepis (Serpentes:
Dipsadidae). Phyllomedusa: Journal of Herpetology. 2021 Jun 22;20(1):15-25.
Camera BF, Miranda EB, Ribeiro-Jr RP, Barros M, Draque J, Waller T, Micucci PA, Dam-
bros CS, Strüssmann C. 2019. Historical Assumptions about the Predation Patterns
of Yellow Anacondas (Eunectes notaeus): Are They Infrequent Feeders? Journal of
Herpetology 53(1):47-52.
Camerano L. 1891. Monografia degli Ofidi italiani. Parte II. Colubridi. Mem. R. Accad. Sci.
Torino, ser. II.
Camp CD, Sprewell WD, Powders VN. 1980. Feeding habits of Nerodia taxispilota with
comparative notes on the foods of sympatric congeners in Georgia. Journal of Herpe-
tology 14(3):301-4.
Camp CL. 1923. Classification of the lizards. Bulletin of the American Museum of Natu-
ral History, Volumes 4-86:386.
Campden-Main SM. 1970. A field guide to the snakes of South Vietnam. Smithsonian In-
stitution, Washington, 112 pp.
Cantor T. 1847. Catalogue of reptiles inhabiting the Malayan peninsula and islands. Jour-
nal of the Asiatic Society of Bengal 16(2): 1–157.
Caprette CL, Lee MS, Shine R, Mokany A, Downhower JF. 2004. The origin of snakes
(Serpentes) as seen through eye anatomy. Biological Journal of the Linnean Society
81(4):469-82.
Capula M, Rugiero L, Luiselli L. 1994. Ecological observations on the Sardinian grass
snake, Natrix natrix cetti. Amphibia-Reptilia 15:221–224.
Carlsson M, Karvemo S, Tudor M, Sloboda M, Mihalca AD, Ghira I, Bel L, Modry D.
2011. Monitoring a large population of Dice snakes at Lake Dinoe in Dobrogea, Roma-
nia. Mertensiella 18:237-45.
Carpenter CC. 1952. Comparative ecology of the common garter snake (Thamnophis s. sir-
talis), the ribbon snake (Thamnophis s. sauritus), and Butler’s garter snake (Thamnophis
butleri) in mixed populations. Ecological Monographs 22(4):236-58.
Carranza S, Arnold EN, Mateo JA, López-Jurado LF. 2000. Long-distance colonization and
radiation in gekkonid lizards, Tarentola (Reptilia: Gekkonidae), revealed by mitochon-
drial DNA sequences. Proceedings of the Royal Society of London. Series B: Biological
Sciences 267(1444):637-49.
Carvalho PS, Zaher H, da Silva Jr NJ, Santana DJ. 2020. A morphological and molecular
study of Hydrodynastes gigas (Serpentes, Dipsadidae), a widespread species from South
622
Aquatic Snakes

America. PeerJ. 2020 Nov 25;8:e10073.

Catania KC, Leitch DB & Gauthier D. 2010. Function of the appendages in tentacled
snakes (Erpeton tentaculatus). Journal of Experimental Biology 213(3):359–367.
Catania KC. 2009. Tentacles snakes turn C-starts to their advantage and predict future
prey behavior. Proceedings of the National Academy of Sciences 106(27):11183–11187.
Catania KC. 2010. Born Knowing: Tentacled Snakes Innately Predict Future Prey Behav-
ior. PLoS ONE 5(6): e10953. [Link]
Cathrine C. 2014. Grass Snakes (Natrix natrix) in Scotland. The Glasgow Naturalist
26(1):36–40.
Cavalheri DG, Santos Carvalho P, Santana DJ. 2021. Defensive repertory of Pseudoeryx
plicatilis (Linnaeus, 1758) in the Pantanal, Mato Grosso do Sul, Brazil. Cuadernos de
Herpetología DOI: 10.31017/18039
Cedeno-Vazquez JR, Beutelspacher-Grcia PM, Nieto-Toscano. 2021. Tretanorhinus nig-
roluteus (Orange-bellied Swamp Snake). Diet and intraspecific competition. Herpeto-
logical Review 52(4):884-885.
Cerón-de la Luz NM, Víctor Vásquez-Cruz V, Canseco-Márquez L, Lara-Hernández FA
2020. Depredación del sapo, en peligro crítico, Incilius cristatus (Anura: Bufonidae)
por Thamnophis sumichrasti (Squamata: Natricidae) en Veracruz, México. Revista Lati-
no Americana de Herpetología 3(1):93-101.
Chan KO, Ahmad NO. 2009. Distribution and natural history notes on some poorly
known frogs and snakes from Peninsular Malaysia. Herpetological Review 40(3):294–
301.
Chan-ard T, Parr JK, Nabhitabhata J. 2015. A Field Guide To The Reptiles of Thailand.
Oxford University Press, NY
Chan-ard T, Parr JWK, Nabhitabhata J. 2015. A field guide to the reptiles of Thailand. Ox-
ford University Press, NY
Chandramouli S, Baiju M, Sebastien J, Ganesh S. 2012. Expanded description of Enhydris
dussumierii (Duméril, Bibron & Duméril, 1854) (Reptilia: Colubridae: Homalop-
sinae). Taprobanica: The Journal of Asian Biodiversity 4(1):42–47.
Chaney T, Pauwels OSG, Nagy ZT, Gvozdik V, Kusamba C, Badjedjea G, Masudi FM,
Ernst R. Greenbaum E. 2021. Poster: Phylogenetics and integrative taxonomy African
Water Snakes (Squamata: Colubridae, Grayia).
Chang TH, Fang PW. 1931. A study of the ophidians and chelonians of Nanking. Contri-
butions from the Biology Laboratory of the Science Society of China 7:249–288.
Chapuis L, Kerr CC, Collin SP, Hart NS, Sanders KL. 2019. Underwater hearing in sea
snakes (Hydrophiinae): First evidence of auditory evoked potential thresholds. Journal
of Experimental Biology 222(14):1–7.
Charlton T. 2020. A guide to snakes of Peninsular Malaysia and Singapore. Natural History
Publications (Borneo), Kota Kinabalu, 299 pp.
Cheng YQ, Ren JL, Wang Z, Zhu GX, Li JT. 2021. On The Taxonomy of Atretium yunnan-
ensis Anderson, 1879 (Squamata: Natricidae), With Redescriptions and Natural Histo-
ry Data of the Poorly Known Species. Russian Journal of Herpetology. 2021 Jul 1;28(4).
Chesoh S, Choonpradub C. 2009. Method for analyzing fish assemblage distribution
with application to fishery landings of tropical shallow lake as Songkhla Lake, Thai-
land. Modern Applied Science 3(5):179-92.
623
Aquatic Snakes

Chim CK, Diong CH. 2009. Reproduction in a female population of the dog-faced water
snake Cerberus rynchops (Homalopsidae) from the Sungei Buloh Wetland Reserve, Sin-
gapore. Joint Meeting of Ichthyologists and Herpetologists (JMIH) Abstract Book, Port-
land: Abstract 405.
Chim CK, Diong CH. 2013. A mark-recapture study of a dog-faced water snake Cerberus
schneiderii (Colubridae: Homalopsidae) population in Sungei Buloh wetland reserve,
Singapore. The Raffles Bulletin of Zoology 61(2): 811–825.
Chippaux JP, Jackson K. 2019. Snakes of central and western Africa. John Hopkins Univer-
sity Press.
Chippaux JP. 2006. Les serpents d’Afrique occidentale et centrale. IRD éditions.
Christman BL, Rodriguez IJ, Jennings RD, Wright JA, Tecca JC. 2021. Thamnophis rufipunc-
tatus (Narrow-headed Gartersnake) Diet. Herpetological Review 52(2):435-436.
Chuaynkern Y, Duengkae P, Pongcharoen C, Chuaynkern C, and Horsin L. 2014. Opisth-
otropis spenceri Smith, 1918 (Serpentes: Natricidae): the third and fourth specimens. J.
Wildlife Thailand 21(1):1-14.
Clark MK, Schoenbohm LM, Royden LH, Whipple KX, Burchfiel BC, Zhang X, Tang W,
Wang E, Chen L. 2004. Surface uplift, tectonics, and erosion of eastern Tibet from large-
scale drainage patterns. Tectonics 23:1–20.
Clay WM. 1938. A synopsis of the North American water snakes of the genus Natrix. Copeia
1938(4):173-82.
Clift PD, Blusztajn J. 2005. Reorganization of the western Himalayan river system after five
million years ago. Nature 438:1001–1003
Cogger HG. 1975. Sea snakes of Australia and New Guinea. In: Dunson, W.A., ed. The Biol-
ogy of Sea Snakes. Baltimore: University Park Press.
Cogger HG. 2000. Reptiles and Amphibians of Australia. Chatswood, NSW: Reed Books.
Cogger HG. 2000. Reptiles and Amphibians of Australia. Chatswood, NSW: Reed Books.
Crowe-Riddell, J. M., D’Anastasi, B. R., Nankivell, J. H., Rasmussen, A. R., & Sanders, K.
L. 2019. First records of sea snakes (Elapidae: Hydrophiinae) diving to the mesopelagic
zone (>200 m). Austral Ecology. doi:10.1111/aec.12717
Cogger HG. 2014. Reptiles and Amphibians of Australia, 7th ed. CSIRO Publishing, xxx +
1033 pp.
Cogger HG. 2018. Reptiles and Amphibians of Australia. Updated Seventh Edition. CSIRO
Publishing, Clayton South, Victoria, Australia. xxxii+1060 pp. ISBN 978-1-4863-0969-6.
Cogger HG. 2018. Reptiles and Amphibians of Australia. Updated Seventh Edition. CSIRO
Publishing, Clayton South, Victoria, Australia. xxxii+1060 pp. ISBN 978-1-4863-0969-6.
Cogger HG, Heatwole HF. 2006. Laticauda frontalis (de Vis, 1905) and Laticauda saintgiron-
si [Link]. from Vanuatu and New Caledonia (Serpentes: Elapidae: Laticaudinae)—a new
lineage of sea kraits? Records of the Australian Museum 58(2): 245–256.
Cogger H, Heatwole H, Ishikawa Y, McCoy M, Tamiya N, Teruuchi T. 1987. The status and
natural history of the Rennell Island sea krait, Laticauda crockeri (Serpentes: Laticau-
didae). Journal of Herpetology 21:255-66.
Cogger HG, Lindner DA. 1974. Frogs and reptiles. Pp. 63–107. In: Fauna Survey of the Port
Essington District, Cobourg Peninsula, Northern Territory of Australia. Frith HJ, Calaby
JH (eds.). CISRO Division of Wildlife Research Technical Paper No. 28.

624
Aquatic Snakes

Cogger H, Heatwole H, Ishikawa Y, McCoy M, Tamiya N, Teruuchi T. 1987. The status

and natural history of the Rennell Island sea krait, Laticauda crockeri (Serpentes: Lati-
caudidae). Journal of Herpetology 21:255-66.
Conant R. 1938. The reptiles of Ohio. American Midland Naturalist. 20(1):1-200.
Conant R. 1943. Studies on North American water snakes- I: Natrix kirtlandii (Kennicott).
American Midland Naturalist 29(2):313-341.
Conant R. 1946. Studies on North American water snakes - II. The subspecies of Natrix vali-
da. American Midland Naturalist 35 (1): 250-275.
Conant R. 1953. Three new water snakes of the genus Natrix from Mexico. Natural History
Miscellanea (126): 1-9.
Conant R. 1961. A new water snake from Mexico, with notes on anal plates and apical pits
in Natrix and Thamnophis. American Museum Novitates (2060):1-22.
Conant R. 1963. Semiaquatic snakes of the genus Thamnophis from the isolated drainage
system of the Rio Nazas and adjacent areas in Mexico. Copeia 1963:473-99.
Conant R. 1965. Miscellaneous notes and comments on toads, lizards, and snakes from
Mexico. American Museum Novitates, no. 2205.
Conant R. 1969. A review of the water snakes of the genus Natrix in México. Bulletin of the
American Museum Natural History. 142: 1-140.
Conelli A, Nembrini M, Mebert K, Schmidt BR. 2009. Monitoring of Dice snakes, Natrix
tessellata (Laurenti, 1768) in the Canton of Ticino (Switzerland). Bollettino della Società
Ticinese di Scienze Naturali 97:27-34 (in Italian).
Conelli AE, Nembrini M, Mebert K. 2011. Different habitat use of Dice snakes (Natrix
tessellata) among three populations in Canton Ticino, Switzerland – a radiotelemetry
study. Mertensiella 18:100-17.
Cook TR, Brischoux F. 2014. Why does the only “planktonic tetrapod” dive? Determinants
of diving behaviour in a marine ectotherm. Animal Behaviour 98:113–123.
Cope ED. 1866. An analytical table of the genera of snakes. Proceedings of the American
Philosophical Society 23:479–499.
Cope ED. 1869. On the reptilian orders Pythonomorpha and Streptosauria. Proceedings
of the Boston Society of Natural History 1869, 12: 250-266.
Cornelis J, Lettoof D. 2020. Notechis scutatus occidentalis (Western Tiger Snake). Defensive
Behavior. Herpetological Review 51(3): 622.
Corrêa DN, Quintela FM, Loebmann D. 2016. Feeding ecology of Erythrolamprus jae-
geri jaegeri and E. poecilogyrus sublineatus. Anais da Academia Brasileira de Ciências
88(1):293–308.
Cortés JE. 1982. The herpetofauna of Gibraltar. Status, recent history and current research.
British Journal of Herpetology 6:273-275.
Costa HC, Santana DJ, F. Leal, R. Koroiva, and P. CA Garcia. 2016. A New Species of Heli-
cops (Serpentes: Dipsadidae: Hydropsini) from Southeastern Brail. Herpetologica 72
(2):157-166.
Costanzo JP, 1986. Influences of hibernaculum microenvironment on the winter life his-
tory of the garter snake (Thamnophis sirtalis). Ohio Journal of Science 86 (5): 199-204.
Costanzo JP. 1989a. Conspecific scent trailing by garter snakes (Thamnophis sirtalis)
during autumn Further evidence for the use of pheromones in den location. Journal of

625
Aquatic Snakes

Chemical Ecology 15:2531-8.

Costanzo JP. 1989b. A physiological basis for prolonged submergence in hibernating
garter snakes Thamnophis sirtalis: evidence for an energy-sparing adaptation. Physi-
ological zoology 62(2):580-92.
Costanzo JP. 1989c. Effects of humidity, temperature, and submergence behavior on sur-
vivorship and energy use in hibernating garter snakes, Thamnophis sirtalis. Canadi-
an Journal of Zoology 67(10):2486-92.
Courtney AJ, Schemel, BC, Wallace R, Campbell MJ, Mayer D.G. Young B. 2010. Reducing
the impact of Queensland’s trawl fisheries on protected sea snakes. Fisheries Research
and Development Coorporation (FRDC) Project #2005/053 Final Report.
Cox MJ, Hoover MF, Chanhome L, Thirakhupt K. 2012. The snakes of Thailand. Chu-
lalongkorn University Museum of Natural History.
Cox MJ, Van dijk PP, Nabhitabhata J, Thirakhupt K.1998. A Photographic Guide to Snakes
and Other Reptiles of Peninsular Malaysia, Singapore and Thailand. Ralph Curtis
Books, Sanibel Island, 144 pp.
Cox MJ. 1991. The snakes of Thailand and their husbandry. Krieger Publishing Company,
Malabar, Florida, 526 pp.
Crawshaw PG. 1991. Effects of hunting on the reproduction of the Paraguayan caiman
(Caiman yacare) in the Pantanal of Mato Grosso, Brazil, Pp. 145-153. In: Neotropical
wildlife use and conservation. J. G. Robinson and K. H. Redford (eds.). University of
Chicago Press, Chicago.
Crombie RI, Pregill GK. 1999. A checklist of the herpetofauna of the Palau Islands (Repub-
lic of Belau), Oceania. Herpetological Monographs 13:29–80.
Crowe A. 1992. Muscle Spindles, Tendon Organs, and Joint Receptors. In C. Gans & P. S.
Ulinski (Eds.), Biology of the Reptilia, Vol. 17, Neurology C (pp. 454–495). University
of Chicago Press.
Crowe-Riddell JM, Simões BF, Partridge JC, Hunt DM, Delean, S, Schwerdt JG, Breen J,
Ludington A, Gower DJ, Sanders KL. 2019. Phototactic tails: evolution and molecular
basis of a novel sensory trait in sea snakes. Molecular Ecology, December 2018:1–16.
Crowe-Riddell JM, Snelling EP, Watson AP, Suh AK, Partridge J & Sanders KL. 2016. The
evolution of scale sensilla in the transition from land to sea in elapid snakes. Open Bi-
ology 6:160054.
Crowe-Riddell JM, Williams R, Chapuis L, Sanders KL. 2019. Ultrastructural evidence of a
mechanosensory function of scale organs (sensilla) in sea snakes (Hydrophiinae). Roy-
al Society Open Science 6:182022.
Crowe-Riddell JM., D’Anastasi BR., Nankivell JH, Rasmussen AR, Sanders KL. 2019. First
records of sea snakes (Elapidae: Hydrophiinae) diving to the mesopelagic zone (>200
m). Austral Ecology. doi:10.1111/aec.12717
Cunningham JD. 1959. Reproduction and food of some California snakes. Herpetologica
15(1):17-9.
Cuvier G. 1829. Le régne animal, Vol. 1. Deterville, Paris. 584 pp.
Czaplicki JA, Porter RH. 1974. Visual cues mediating the selection of goldfish (Carassius
auratus) by two species of Natrix. Journal of Herpetology 8(2):129–134.
D’Abreu, EA. 1912 [1913]. Effect of a bite from Schneider’s water snake (Hypsirhina en-
hydris). Journal of the Bombay Natural History Society 22:203.
626
Aquatic Snakes

D’Anastasi BR., van Herwerden L, Hobbs JA., Simpfendorfer, C. A., & Lukoschek, V. 2016.
New range and habitat records for threatened Australian sea snakes raise challenges for
conservation. Biological Conservation, 194, 66–70. doi:10.1016/[Link].2015.11.032
Da Costa FRF, Bezerra LL, Rodrigues VLM, Cassiano-Lima. 2021. Helicops leopardinus
(Leopard Keelback Snake). Mortality. Herpetological Review 52(4):872-873.
Da Frota JG. 2005. New species of Helicops Wagler, 1830 (Serpentes, Colubridae) from
Tapajós river, Amazon region, Brazil. Phyllomedusa: Journal of Herpetology 4(1):61-7.
Da Silva, MAO, Heegaard S, Wang T, Gade JT, Damsgaard C, Bertelsen MF 2017. Mor-
phology of the snake spectacle reflects its evolutionary adaptation and development.
BMC Veterinary Research, 13(1): 1–8.
Dahl TE. 1990. Wetland Losses in the United States 1780’s to 1980’s. United States Depart-
ment of the Interior, Fish and Wildlife Service, Washington, DC, USA
D’Août K, Aerts P. 1999. A kinematic comparison of forward and backward swimming in
the eel Anguilla anguilla. Journal of Experimental Biology 202(11):1511–1521.
Das A, Gower DJ, Deepak V. 2020. Lost and found: rediscovery and systematics of the
Northeast Indian snake Hebius pealii (Sclater, 1891). Vertebrate Zoology 70(3):305-18.
Das I, Breuer H, Shonleben S. 2013. Gerarda prevostiana (Eydoux and Gervais, 1837)
(Squamata: Serpentes: Homalopsidae), a new snake for Borneo. Asian Herpetological
Research 4(1):76–78.
Das I, Lakim M, Lim KK, Hui TH. 2008. New species of Anomochilus from Borneo
(Squamata: Anomochilidae). Journal of Herpetology. 42(3):584-91.
Das I. 1993. Annandale’s sea snake, Kolpophis annandalei (Laidlaw 1901): a new record for
Borneo. Raffles Bull. Zool. 41: 359-361.
Das I. 2010. Field Guide To The Reptiles Of South-East Asia. New Holland Publishers
(UK) Ltd. London. ISBN 978-1-84773-347-4
Das I. 2010. Field Guide To The Reptiles Of South-East Asia. New Holland Publishers
(UK) Ltd. London. ISBN 978-1-84773-347-4
David P, Vogel G. 1996. The Snakes of Sumatra. An annotated checklist and key with
natural history notes. Bücher Kreth, Frankfurt/M.
David P, Pauwels OS, Nguyen TQ, Vogel G. 2015. On the taxonomic status of the Thai
endemic freshwater snake Parahelicops boonsongi, with the erection of a new genus
(Squamata: Natricidae). Zootaxa. 3948(2):203-17.
David P, Vogel G, Nguyen TQ, Orlov NL, Pauwels OSG, Teynié A, Ziegler T. 2021. A
revision of the dark-bellied, stream-dwelling snakes of the genus Hebius (Reptilia:
Squamata: Natricidae) with the description of a new species from China, Vietnam, and
Thailand. Zootaxa 4911: 1–61.
David P. Agarwal I, Athreya R, Mathew R, Vogel G, Mistry VK. 2015. Revalidation of Na-
trix clerki Wall, 1925, an overlooked species in the genus Amphiesma Duméril, Bibron
& Duméril, 1854 (Squamata: Natricidae). Zootaxa 3919 (2): 375–395.
David P., Ineich I. 1999. Les serpents venimeux du monde: systématique et repartition.
Dumerilia 3:1–499.
David, P. & Vogel, G. 1996. The Snakes of Sumatra. An annotated checklist and key with
natural history notes. Bücher Kreth, Frankfurt/M.
Davies WL, Cowing JA, Bowmaker JK, Carvalho LS, Gower DJ, Hunt DM. 2009. Shed-
ding light on serpent sight: the visual pigments of henophidian snakes. Journal of
627
Aquatic Snakes

Neuroscience, 29(23):7519-25.
de Aguiar LF, Di-Bernardo M. 2004. Diet and feeding behavior of Helicops infrataeniatus (Ser-
pentes: Colubridae: Xenodontinae) in southern Brazil. Studies on Neotropical Fauna and En-
vironment 39(1):7-14.
de Carvalho Teixeira C, de Assis Montag LF, dos Santos-Costa MC. 2017. Diet composition and
foraging habitat use by three species of water snakes, Helicops Wagler, 1830,(Serpentes: Dipsa-
didae) in eastern Brazilian Amazonia. Journal of Herpetology 51(2):215-22.
de Jong JK. 1926. Heurnia ventromaculata n.g. n. sp. und Cantoria annulata n. sp., wei neue Sch-
langen von Neu-Guinea. Zoologischer Anzeiger 67:302–304.
de la Quintana P, Rivas JA, Valdivia F, Pacheco LF. 2017. Home range and habitat use of Beni ana-
condas (Eunectes beniensis) in Bolivia. Amphibia-Reptilia. 38(4):547-53.
de Lang R. 2013. The snakes of the Moluccas (Maluku), Indonesia. Edition Chimaira, Frankfurt
am Main. 2013:417.
de Lang R. 2017. The snakes of Java, Bali, and surrounding islands. Frankfurt: Edition Chimaira.
de Lepiney J, F Nemeth. 1945-46-47. Notes zoologiques marocaines. Bulletin de la Société des Sci-
ences Naturelles et Physiques du Maroc 25-27:227-233.
de Lima Moraes LJ, de Fraga R.2015. Climbing behavior in the aquatic snake Helicops hagmanni
Roux 1910 (Serpentes: Dipsadidae). Herpetology Notes 26;8:19-20.
de Oliveira L, Scartozzoni RR, de Almeida-Santos SM, Jared C, Antoniazzi MM, da Graça
Salomão M. 2016. Morphology of Duvernoy’s glands and maxillary teeth and a possible
function of the Duvernoy’s gland secretion in Helicops modestus Günther, 1861 (Serpentes:
Xenodontinae). South American Journal of Herpetology 11(1):54-65.
de Oliveira LCC, Leite AK, Pagel, GS, de ArauJo H, Tinoco MS. 2019 Erythrolamprus miliaris
merremi (watersnake). Diet. Herpetological Review 50(4):800.
de Queiroz A, Lawson R, Lemos-Espinal JA. 2002. Phylogenetic relationships of North American
garter snakes (Thamnophis) based on four mitochondrial genes: how much DNA sequence is
enough? Molecular phylogenetics and evolution 22(2):315-29.
de Queiroz K. 2021. Response to Criticisms of an updated subspecies concept. Herpetological Re-
view 52(4):773-776.
de Rooij N. 1917. The Reptiles of the Indo-Australian Archipelago. II. Ophidia. Leiden: E.J. Brill.
xiv + 334 pp., 117 Figures. (Hydrophis mertoni, new combination, p. 233).
de Solan T, Renoult JP, Geniez P, David P, Crochet PA. 2020. Looking for mimicry in a snake as-
semblage using deep learning. The American Naturalist 196(1):74–86.
de Souza HF. A study on the population dynamics & morphometric variations in Cerberus ryn-
chops (dog-faced water snake) population of Ganjam coast, Odisha. Unpublished Masters the-
sis, Pondicherry University, 64 pp.
Decemson HT, Lalremsanga HT. 2021. Herptoreas xenura (Wall’s Keelback). Diet. Herpetological
Review 52(4):873.
Deepak, V., Lalronunga, S., Lalhmingliani E., Das, A., Narayanan, S., Das, I., Gower, D.J. 2021.
Phylogenetic relationships of xenodermid snakes (Squamata: Serpentes: Xenodermidae), with
the descriptionof a new genus. Vert. Zool. 71: 747-763.
Deepak V, Cooper N, Poyarkov NA, Kraus F, Burin G, Das A, Narayanan S, Streicher JW, Smith
S-J, Gower DJ. 2022. Multilocus phylogeny, natural history traits and classification of natricine
snakes (Serpentes: Natricinae). Zoological Journal of the Linnean Society 195:279–298.
Deepak V, Maddock ST, Williams R, Nagy ZT, Conradie W, Rocha S, Harris DJ, Perera A,

628
Aquatic Snakes

Gvoždík V, Doherty-Bone TM, Kamei RG. 2021. Molecular phylogenetics of sub-Saha-

ran African natricine snakes, and the biogeographic origins of the Seychelles endemic
Lycognathophis seychellensis. Molecular Phylogenetics and Evolution 61:107152.
Deepak, V., Ruane S, Gower D. 2019. A new subfamily of fossorial colubroid snakes from
the Western Ghats of peninsular India. Journal of Natural History. 52:45-46, 2919-2934.
Degenhardt WG, Painter CW, Price AH. 1996. Amphibians and Reptiles of New Mexico.
University of New Mexico Press, Albuquerque, New Mexico.
Del Canto R, Busack SD. 2011. Natrix maura (viperine snake). Non-accidental salt water
activity. Herpetological Review 42:295-296.
Delisle ZJ, Ransom Jr D, Lutterschmidt WI, Delgado-Acevedo J. 2019. Multiscale Spatiotem-
poral Habitat Selection of Northern Cottonmouths. Journal of Herpetology 53(3):187-
95.
Demuth J. 1997. Regina alleni (striped crayfish snake). Size. Herpetological Review 28:90-1.
Department of the Environment 2021. Hydrophis elegans in Species Profile and Threats Da-
tabase, Department of the Environment, Canberra. Available from: https:// [Link]-
[Link]/sprat. Accessed Mon, 27 Sep 2021 11:06:21 +1000.
Deraniyagala PEP. 1955 [1987 reprint]. A colored atlas of some vertebrates from Ceylon
Volume 3. Serpentoid Reptilia. Colombo: Ceylon National Museum Publication.
Deuve J. 1970. Serpents du Laos. Office de la Recherche Scientifique et Technique Outre-
Mer, Paris, Mémoire no. 39, 251 pp.
Díaz-Paniagua C. 1976. Alimentación de la culebra bastarda (Malpolon monspessulanus;
Ophidia, Colubridae) en el S.O. de España. Doñana, Acta Vertebrata 3(2):113–127.
Di Nicola M. R. & Bruni G., 2020. A case of active predation of Natrix helvetica (Serpentes:
Colubridae) on Sturnus vulgaris (Passeriformes: Sturnidae). Herpetology Notes, vol. 13:
461-462.
Di Nicola MR, Cavigioli L, Luiselli L, Andreone F. 2021: Anfibi & Rettili d’Italia. Edizione
aggiornata a cura di Matteo R. Di Nicola. Edizioni Belvedere, Latina, “historia naturae”
(8), 576 pp.
di Pietro DO, Alcalde L, Williams J D. 2014. Nasal cartilages, hyobranchial apparatus,
larynx, and glottal tubes in four species of Hydropsini (Serpentes: Dipsadidae: Xeno-
dontinae); Senckenberg Gesellschaft für Naturforschung; Vertebrate Zoology, 64; 1;
30-4-2014; 103-111.
Díaz E. 2004. Predación de culebras de collar Natrix natrix en colonias de avión zapador Ri-
paria riparia. Boletín de la Asociación Herpetológica Española 15(1):32–33.
Díaz EA, Aguirre-Ramírez KA. 2014. Natural History Notes. Thamnophis pulchrilatus. Her-
petological Review 45:2.
Dimitropoulos A. 1989. The Dice snake and its breeding in captivity. Herptile 14(2):66-67.
Díncaslan YE, Arikan H, Ugurtas IH, Mebert K. 2011. Morphology and blood proteins of
Dice snakes from western Turkey. Mertensiella 18:370-83.
Dirksen L, Böhme W. 2005. Studies on Anacondas. III. A Reappraisal of Eunectes beniensis
Dirksen, 2002, from Bolivia, and a Key to the Species of the Genus Eunectes Wagler,
1830 (Serpentes: Boidae). Russian Journal of Herpetology, 12(3), pp.223-229.
Dirksen L, Henderson RW. 2002. Eunectes deschauenseei. Catalogue of American Amphibi-
ans and Reptiles (CAAR).
Dirksen L. 2002. Anakondas. Münster: Natur und Tier Verlag.

629
Aquatic Snakes

Dissanayake DSB and Wellappuliarachchi. 2016. Amphiesma stolatum (Buff-striped Keel-

back). Diet. Herpetological Review 47(2):307.
Dixon JR 1989. A key and checklist to the neotropical snake genus Liophis with country lists
and maps. Smithsonian Herpetological Information Series 79:1–40.
Dixon JR, Markezich AL. 1992. Taxonomy and geographic variation of Liophis poecilo-
gyrus (Wied) From South America (Serpentes, Colubridae). Texas Journal of Science
44(2):131-66.
Dixon JR, Soini P. 1977. The reptiles of the upper Amazon Basin, Iquitos region, Peru. II.
Crocodilians, turtles, and snakes. Contributions in biology and geology. 1977;12.
Dixon JR. 1983. Taxonomic status of the South American snakes Liophis miliaris, L. amazo-
nicus, L. chrysostomus, L. mossoroensis and L. purpurans (Colubridae: Serpentes). Copeia
1983:791-802.
Dixon JR. 2000. Amphibians and reptiles of Texas, second edition. Texas A&M University
Press, 421 pp.
Dixon-MacCallum GP, Bell KA, Gregory PT. Edge-habitat Use by Northwestern Garter-
snakes (Thamnophis ordinoides) in Saanich, British Columbia. The Canadian Field-Nat-
uralist. 2017 Jul 14;131(1):37-45.
Dixon-MacCallum GP, Bell KA. 2016. Thamnophis ordinoides (Northwestern Gartersnake>
Reproduction. Herpetological Review 47(3):487.
Dobreff M. 1939. Über Hungerfähigkeit bei Schlangen. Zool. Garten Leipzig 10:218–221.
Donini J. 2018. Nerodia fasciata (Southern Watersnake). Diet. Herpetological Review
49(4):757.
Donoso-Barros P. 1966. Reptiles de Chile. Ediciones Universidad de Chile. Santiago Chile.
458 pp.
Dorcas ME, Poppy SM, Ernst CH, Gibbons JW. 2003. Regina alleni. Catalogue of American
Amphibians and Reptiles (CAAR). 2003.
Dorge T, Hofmann S, Wangdwei M, Duoje L, Solhøy T, Miehe G, 2007. The ecological
specialist, Thermophis baileyi (Wall, 1907): new records, distribution, and biogeo-
graphic conclusions. Herpetological Bulletin, 101: 8–12.
Dowling HG, Duellman WE. 1978. Systematic Herpetology: A Synopsis of Families and
Higher Categories. HISS Publications, New York, 118 pp.
Dowling HG. 1950. Studies of the black swamp snake, Seminatrix pygaea (Cope), with de-
scriptions of two new subspecies. Miscellaneous publications, Museum of Zoology, Uni-
versity of Michigan (76): 1-38.
Dowling HG. 1950. Studies of the black swamp snake, Seminatrix pygaea (Cope), with de-
scriptions of two new subspecies. Miscellaneous publications, Museum of Zoology, Uni-
versity of Michigan (76): 1-38.
Dowling HG. 1960. The curious feeding habits of the Java Wart Snake. Animal Kingdom
63:13–15.
Dowling HG. 1974. A provisional classification of snakes. Pp. 167–170. In: HISS Yearbook
of Herpetology. Dowling HG, (ed.). HISS, American Museum of Natural History, New
York.
Drummond HM. 1979. Stimulus control of amphibious predation in the Northern Water
Snake (Nerodia s. sipedon). Zeitschrift Für Tierpsychologie, 50(1):18–44.

630
Aquatic Snakes

Drummond H. 1983. Aquatic Foraging in Garter Snakes: a Comparison of Specialists and

Generalists. Behaviour 86(1-2):1-30.
Drummond H. 1985. The role of vision in the predatory behaviour of natricine snakes. Ani-
mal Behaviour 33(1):206-15.
Duarte MR, Neto DG, Junior TV, Pinheiro MA. 2014. Predation on the sleeper goby,
Guavina guavina (Perciformes, Eleotridae), by the military ground snake, Erythrol-
amprus miliaris orinus (Serpentes, Dipsadidae), in a mangrove area of Southeastern
Brazil. Herpetology Notes, 7:577-80.
Dubach J., Sajewicz A., Pawley R. 1997. Parthenogenesis in the Arafuran file snake, Acro-
chordus arafurae. Herpetol. Nat. Hist. 5, 11–8
Dubey S, Ursenbacher S, Schuerch J, Golay J, Aubert P, Dufresnes C. 2017. A glitch in the
Natrix: cryptic presence of alien grass snakes in Switzerland. Herpetology Notes 10:205–
208.
Dubrovsky YuA. 1967. New finds of reptiles in the steppes of Kazakhstan. Bulletin of the
Moscow Society of Naturalists. Biological Department, 72(1.C):146-147.
Dudgeon D, Arthington AH, Gessner MO, Kawabata ZI, Knowler DJ, Lévêque C, Naiman
RJ, Prieur-Richard AH, Soto D, Stiassny ML, Sullivan CA. 2006. Freshwater biodiversity:
importance, threats, status and conservation challenges. Biological reviews 81(2):163-82.
Duellman WE. 1961. The amphibians and reptiles of Michoacan, Mexico. University of
Kansas Publication Museum. Natural History 15(1):1-148.
Duellman WE. 1978. The biology of an equatorial herpetofauna in Amazonian Ecuador.
Publications of the Museum of Natural History, University of Kansas 65: 1–352.
Duguy R, Saint-Girons H. 1966. Cycle annuel d’activité et reproduction de la couleuvre vi-
perine Natrix maura (L.), d’après les notes manuscrites de Rollinat et des observations
personnelles. La Terre et la Vie 4:423-457.
Duméril AMC, Bibron G, Duméril A. 1854. Erpétologie Générale ou historie naturelle com-
plete des reptiles. Vol. 7 (2 parts) Librarie Encyclopedique de Roret, London.
Dumont M. 1979. Elevage et reproduction de Natrix maura. Bulletin de la Société her-
pétologique de France. 12:7-10.
Dunn ER. 1939. Mainland forms of the snake genus Tretanorhinus. Copeia 1939: 212–
217.
Dunn ER. 1944. A revision of the Colombian snakes of the genera Leimadophis, Lygophis,
Liophis, Rhadinaea, and Pliocercus, with a note on Colombian Coniophanes. Caldasia
1:479-95.
Dunn ER, Conant R. 1936. Notes on anacondas, with descriptions of two new species. Pro-
ceedings of the Academy of Natural Sciences of Philadelphia. 88:503-6.
Dunson W. (ed.) 1975. The Biology of the Sea Snakes. University Park Press, Baltimore,
Maryland 339 pp.
Dunson WA & Dunson MK. 1973. Convergent evolution of sublingual salt-glands in marine
file snake and true sea snakes. Journal of Comparative Physiology 86(3):193–208.
Dunson WA, Dunson MK. 1979. A possible new salt gland in a marine homalopsid snake
(Cerberus rhynchops). Copeia 1979(4):661–672.
Dunson WA, Ehlert GW. 1971. Effects of temperature, salinity, and surface water flow on
distribution of the sea snake pelamis. Limnology and Oceanography, 16(6), 845–853.
doi:10.4319/lo.1971.16.6.0845
631
Aquatic Snakes

Dunson WA, Packer RK & Dunson MK. 1971. Sea snakes: An unusual salt gland under the
tongue. Science 173(3995):437–441.
Dunson WA. 1980. The relation of sodium and water balance to survival in sea water of
estuarine and freshwater races of the snakes Nerodia fasciata, N. sipedon and N. valida.
Copeia. 1980 1:268-80.
Dunson, W.A., 1975. The biology of sea snakes. University of Pennsylvania Press.
Durso AM, Willson JD, Winn C. 2013. Habitat influences diet overlap in aquatic snake as-
semblages. Journal of Zoology 291:185-193.
Durson AM, Laverick A. 2021. Francia erythrogramma (Rainbow Snake). Plastic mesh en-
tanglement. Herpetological Review 52(1):157.
Dutta SK. 1989. On the mass killing of the dog-faced water snake (Cerberus rhynchops) in
the Chilka Lake, Orissa. Hamadryad 14(2):27–[Link] SK. 1989. On the mass killing of
the dog-faced water snake (Cerberus rhynchops) in the Chilka Lake, Orissa. Hamadryad
14(2):27–28.
Dwyer Q, Arteaga A, Barrio-Amorós, CL, Flagle A. 2018. Trachyboa boulengeri: Diet.
Herpetological Review 49(2):359-360.
Dyer B. 1979. Some reptile breeding records. The Journal of the Herpetological Associa-
tion of Africa, 20(1):6-6.
Eckstein H-P. 1993. Untersuchungen zur Ökologie der Ringelnatter (Natrix natrix Linnae-
us, 1758). Abschlussbericht Ringelnatter - Projekt-Wuppertal (1986–1991). Verlag für
Ökologie und Faunistik. Duisburg, Germany.
Edgar PW, Lilley RPH. 1993. Herpetofauna survey of Manusela National Park. Chapter 8.
Pp. 131–141. In: Natural history of Seram, Maluku, Indonesia. Edwards, MacDonalds,
and Proctor (eds.). Intercept Ltd, Andover, UK.
Edelstamm C. 1989. A long term study of snake populations. First World Congress of Her-
petology. University of Kent and Canterbury (Abstracts).
Edwards M, Sheehy CM, Fedler MT, Lillywhite HB. 2021. Thirst and drinking in North
American watersnakes (Nerodia spp.). The Journal of Experimental Biology 224:1–4.
Egan D. 2007. Snakes Of Arabia: A Field Guide To The Snakes Of The Arabian Peninsula
And Its Shores. Motivate Publishing, Dubai, UAE. 208pp. ISBN 978-1-86-63-239-6.
Eisermann K, Avendaño C, Acevedo M, Matías E. 2016. Elevational range extension and
new habitat for Thamnophis fulvus (Bocourt, 1893) (Squamata: Natricidae). Mesoameri-
can Herpetology 3:1094-1097.
Eisfeld A, Vrcibradic D. 2019. Reproductive aspects of the semi-aquatic snake Erythrolam-
prus miliaris (Dipsadidae: Xenodontini) in the state of Rio de Janeiro, southeastern Bra-
zil. Anais da Academia Brasileira de Ciências. 2019 Apr 8;91.
Eisfeld A, Pizzatto L, Vrcibradic D. 2021, Diet of the semiaquatic snake Erythrolamprus mil-
iaris (Dipsadidae, Xenodontinae) in the Brazilian Atlantic Forest. Journal of Herpetolo-
gy 55(4):330-7.
El Din SB. 2006. A guide to the reptiles and amphibians of Egypt. The America University,
Cairo Press, Cairo.
Elfes CT, Livingstone SR, Lane ALV, Sanders KL, Courtney AJ, Gatus JL, Guinea M, Lobo
AS, Milton D, Rasmussen A, Read M, White M-D, Sanciangco J, Alcala A, Heatwole H,
Karns DR, Seminoff JA, Voris HK, Carpenter KE, Murphy JC 2013. Fascinating and for-
gotten, the conservation status of marine elapid snakes. Herpetological Conservation

632
Aquatic Snakes

and Biology, 8 (1). pp. 37-52.

Els, J., Al Johany, A.M.H., Amr, Z.S.S., Orlov, N.L., Shafiei Bafti, S., Werner, Y.L., Disi, A.M.,
Papenfuss, T., Ananjeva, N.B., Borkin, L., Milto, K., Golynsky, E., Rustamov, A, Munkh-
bayar, K., Nuridjanov, D., Dujsebayeva, T., Shestopal, A. & Das, A. 2021. Platyceps
rhodorachis. IUCN Red List of Threatened Species. 2021: e.T164605A1060890.
Emmons ID, Nowak EM, Lauger KK. 2016. Prey availability and foraging events of the
northern Mexican Gartersnake (Thamnophis eques megalops) in north-central Arizona.
Herpetological Review 47:555-561.
Ernst CH, Ernst EM. 2003. Snakes of the United States and Canada. Smithsonian Books.
Ernst CH, Zug GR. 1996. Snakes in question: The Smithsonian answer book. Washington,
DC: Smithsonian Institution Press.
Ernst CH. Regina septemvittata. Catalogue of American Amphibians and Reptiles (CAAR).
2002.
Ersan JS, Halstead BJ, Wildy EL, Casazza ML, Wylie GD. 2020. Giant Gartersnakes (Tham-
nophis gigas) Exploit Abundant Nonnative Prey While Maintaining Their Appetite for
Native Anurans. Herpetologica 76(3):290-6.
Escoriza D. 2018. Patterns of occurrence of semi-aquatic reptiles in highly invaded Mediter-
ranean rivers. NeoBiota 38:23.
Escoriza D, Ben Hassine J. 2017. Niche separation among north-west African semi-aquatic
reptiles. Hydrobiologia 797:47–56.
Escoriza D, Pascual G. 2021. Habitat occupancy by semi‐aquatic reptiles on an aridity gra-
dient in the western Mediterranean. River Research and Applications 37(9):1233–1242.
Eskew EA, Willson JD, Winne CT. 2009. Ambush site selection and ontogenetic shifts in
foraging strategy in a semi‐aquatic pit viper, the Eastern cottonmouth. Journal of Zo-
ology. 277(2):179-86.
Esquerré D, Keogh JS. 2016. Parallel selective pressures drive convergent diversification
of phenotypes in pythons and boas. Ecology Letters 19(7):800-9.
Esquerré D, Keogh JS. 2016. Parallel selective pressures drive convergent diversification of
phenotypes in pythons and boas. Ecology Letters 19:800–809.
Etchepare E, Zaracho V, Semhan R, Aguirre R. 2012. Further notes on the reproduction
of Hydrops caesurus (Serpentes: Colubridae) from Corrientes, Argentina. Herpetology
Notes 5: 169-170.
Eversole CB, Powell RL, Lizarro D, Bravo RC. 2016. Thamnodynastes lanei Diet. Herpeto-
logical Review 47 (1): 155.
Eydoux F, Gervais P. 1837. Voyage de la Favourite. Reptiles. Magasin de Zoologie Journal
destine á établir une correspondence entre les Zoologistes de tous les pays, et á leur faci-
liter les moyens de publiler les espéces nouvelles ou peu connues qu’ils possédent; par F.
E. Guérin, 3, 1–10.
Everitt B, Philipis J. 2019. Nerodia erythrogaster (Plain-bellied Watersnake) and Nerodia fas-
ciata (Southern Watersnake). Death Feigning. Herpetological Review 50 (3):598.
Fabre A-C, Bickford D, Segall M, Herrel A. 2016. The impact of diet, habitat use, and be-
havior on head shape evolution in homalopsid snakes. Biological Journal of the Linnean
Society 118(3):634–647.
Faghiri A, Shiravi A, Hojati V, Kami HG. 2011. Observations on the spermatogenetic cycle
of the grass snake, Natrix natrix (Serpentes: Colubridae) in Northern Iran. Asian Herpe-
633
Aquatic Snakes

tological Research 2:55–59.

Fahd S. 2001. Biogeographie, Morphologie et Ecologie des Ophidiens du Rif (Nord du Ma-
roc. Thès. Doc., Univ. Abdelmalek Essaddi, Tetuán.
Fahd S. 2001. Biogéographie, Morphologie et Ecologie des ofidiens du Rif (Nord du Maroc).
Tesis doctoral, Universidad de Tetuán, Marruecos.
Fantuzzi JA. 2020. Observations of captive roughbacked litter snakes (Xenodermus javani-
cus - Reinhardt, 1836). IRCF, Reptiles & Amphibians. 27 (3): 543545.
Faraone FP, Giacalone G, Lo Valvo, M. 2010. Dati preliminari sulla biometria, il cromatismo
e la dieta di una popolazione di Natrix natrix della Sicilia occidentale. In Atti VIII Con-
gresso Nazionale Societas Herpetologica Italica, p. 247–252.
Farina RK, Dias J, Tozetti AM. 2019. Erythrolamprus miliaris (Military Ground Snake). Diet
and Foraging Behavior. Herpetological Review 50(4):799-800.
Feaver PR. 1976. A population study of the Northern Watersnake, Natrix sipedon. Herpeto-
logical Review 7:81 (Abstr.).
Feder ME. Blood oxygen stores in the file snake, Acrochordus granulatus, and in other ma-
rine snakes. Physiological Zoology. 1980 Oct 1;53(4):394-401.
Feriche M. 1998. Ecología de la reproducción en colúbridos del sureste de la Península
Ibérica. Tes. Doc. Univ. Granada, Granada.
Feriche M, Pleguezuelos JM, Cerro A. 1993. Sexual dimorphism and sexing of mediterra-
nean colubrids based on external characteristics. Journal of Herpetology 27(4):357–362.
Feriche M, JM Pleguezuelos. 1999. Características de la reproducción de la culebra viperina,
Natrix maura (Linnaeus, 1758), en la Depresión de Granada (sureste ibérico). Revista
Española de Herpetología 13:37-48.
Fernandes DS, Germano VJ, Fernandes R, Franco FL. 2002. Taxonomic status and geo-
graphic distribution of the lowland species of the Liophis cobella group, with comments
on the species from the Venezuelan tepuis (Serpentes: Colubridae). Boletim do Museu
Nacional, Nova serie, Zoologia, 481: 1–14.
Ferreira AS, Ferrao M, Dayrell JS. 2020. Anilius scytale (Red Pipesnake). Reproduction and
Neonate Size. Herpetological Review 51(4):858.
Figueroa A., McKelvy AD, Grismer LL, Bell CD, Lailvaux SP. 2016. A species-level phylog-
eny of extant snakes with description of a new colubrid subfamily and genus. PloS One
11(9):e0161070.
Filipek DM. 2016. Phylogeography and historical demography of the Queensnake (Regina
septemvittata). University of Central Arkansas.
Filippakopoulou A, X Santos, M Feriche, JM Pleguezuelos, GA Llorente. 2014. Effect of prey
availability on growth-rate trajectories of an aquatic predator, the viperine snake Natrix
maura. Basic and Applied Herpetology 28:35-50.
Filippi E, Capula M, Luiselli L, Agrimi U. 1996. The prey spectrum of Natrix natrix (Linnae-
us, 1758) and Natrix tessellata (Laurenti, 1768) in sympatric populations ((Squamata:
Serpentes: Colubridae). Herpetozoa 8: 155–164.
Finley Jr RB, Chiszar D, Smith HM. 1994. Field observations of salivary digestion of rodent
tissue by the wandering garter snake, Thamnophis elegans vagrans. Bulletin of the Chica-
go Herpetological Society 29:5-6.
Fischer JG. 1886. Herpetologische Notizen. Abhandlungen der Gesellschaft der Naturwis-
senschaften zu Hamburg 9(1):51–67.
634
Aquatic Snakes

Fitch H. 1965. An ecological study of the garter snake, Thamnophis sirtalis. University of
Kansas Publication Museum of Natural History 15(10):493- 564.
Fitch HS. 1980. Remarks concerning Certain Western Garter Snakes of the Thamnophis ele-
gans complex. Transactions of the Kansas Academy of Science. 83(3): 106–113.
Fitch HS. Thamnophis sirtalis. Catalogue of American Amphibians and Reptiles (CAAR).
1981.
Fitzsimons FW. 1930. Pythons and their ways. GG Harrap & Company Limited.
Flores E, Jadin RC, Orlofske SA. 2010. Helicops polylepis (Norman’s Keelback). Diet and in-
traspecific competition. Herpetological Review 41:93-94.
Flower SS. 1925. Contributions to our Knowledge of the Duration of Life in Vertebrate An-
imals.—III. Reptiles. In Proceedings of the Zoological Society of London 95, No. 3, pp.
911-981. Oxford, UK: Blackwell Publishing Ltd.
Folie A, Mees F, De Putter T, Smith T. Presence of the large aquatic snake Palaeophis afri-
canus in the middle Eocene marine margin of the Congo Basin, Cabinda, Angola. Geo-
bios. 2021 Jul 1;66:45-54.
Ford NB, Ball R. 1977. Clutch size and size of young in the Mexican garter snake, Thamno-
phis melanogaster (Reptilia, Serpentes, Colubridae). Herpetological Review 8:118.
Ford NB, Ford DF. 2002. Notes on the ecology of the South American water snake Helicops
angulatus (Squamata: Colubridae) in Nariva Swamp, Trinidad. Caribbean Journal of Sci-
ence 38(1/2):129-31.
Fortes VB, Lucas EM, Caldart VM 2010. Reptilia, Serpentes, Dipsadidae, Gomesophis
brasiliensis (Gomes, 1918): Distribution extension in state of Santa Catarina, Brazil. –
Checklist, 6: 414– 415.
França FGR, Araújo AFB. 2007. Are there co-occurrence patterns that structure snake
communities in Central Brazil? Brazilian Journal of Biology 67(1): 33-40.
Franca RC, Sampaio ILR, Franca FGR. 2018. Helicops angulatus (Brown-banded Water-
snake). Plastic ingestion. Herpetological Review 49(2):342.
Francis L. Rose, Thomas R. Simpson, James R. Ott, Richard W. Manning, and Julien
Martin 2010. Survival of Western Cottonmouths (Agkistrodon piscivorus leucosto-
ma) in a pulsing environment. The Southwestern Naturalist 55(1), 11-15, (1 March
2010). [Link]
Franco FL, Fernandes DS, Bentim BM. 2007. A new species of Hydrodynastes Fitzinger,
1843 from central Brazil (Serpentes: Colubridae: Xenodontinae). Zootaxa. 1613(1):57-
65.
Franco FL. 2000. Relações filogenéticas entre gêneros da tribo Tachymenini Bailey (1967)
(Serpentes; Colubridae) (Doctoral dissertation).
Frazzetta TH. 1999. Adaptations and significance of the cranial feeding apparatus of the
sunbeam snake (Xenopeltis unicolor): Part I. Anatomy of the skull. Journal of Mor-
phology 239(1):27-43.
Freedman B, Catling PM. 1979. Movements of sympatric species of snakes at Amherstburg,
Ontario. Canadian Field-Naturalist 93:399–404.
Friedlander AM, Arribas J, Ballesteros E, Betz J, Bosserelle P, Brown E, Caselle JE, Cramp
JE, Gataua L, Helagi N, Mayorga J,McAloney D, Myers D, Pasisi B, Richmond’Rex A,
Rose P, Salinas-de-León P, San Félix M, Thompson C, Turchik A, Sala E. 2017. Exploring
the marine ecosystems of Niue and Beveridge Reef. Report to the government of Niue.
635
Aquatic Snakes

National Geographic Pristine Seas.

Frith CB. 1977. A survey of the snakes of Phuket Island and the adjacent mainland areas of
peninsular Thailand. Journal of the Natural History Society of Siam 26:263–316.
Frith CB. 1977. A survey of the snakes of Phuket Island and the adjacent mainland areas of
peninsular Thailand. Journal of the Natural History Society of Siam 26:263–316.
Fritz U., Corti C. & Päckert M., 2012. Mitochondrial DNA sequences suggest unexpected
phylogenetic position of Corso-Sardinian grass snakes (Natrix cetti) and do not sup-
port their species status, with notes on phylogeography and subspecies delineation of
grass snakes. Organisms Diversity and Evolution, 12 (1): 71-80.
Fritz U, and Ihlow F. 2022. Citizen Science, taxonomy and grass snakes: iNaturalist helps
to clarify variation of coloration and pattern in Natrix natrix subspecies. Vertebrate
Zoology 72:533–549. [Link]
Fritz U. and Schmidtler J.F., 2020. The Fifth Labour of Heracles: Cleaning the Linnean sta-
ble of names for grass snakes (Natrix astreptophora, N. helvetica, N. natrix sensu stric-
to). Vertebrate Zoology 70 (4):621-665.
Frotzler N, Davitashvili N, Mebert K. 2011. Distribution of the Dice snake (Natrix tessella-
ta) in Georgia (transcaucasia) and Comparative notes on the Genus Natrix. Mertensi-
ella 18:357-65.
Fry BG, Scheib H, de Azevedo ID, Silva DA, Casewell NR. 2012. Novel transcripts in the
maxillary venom glands of advanced snakes. Toxicon 59(7-8):696-708.
Fry BG. February 24, 2012. “Snakes Venom LD – List of the Available Data and Sorted by
Route of Injection”. [Link].
Fry GC, Milton A, Wassenberg DJT. 2001. The reproductive biology and diet of sea snake
bycatch of prawn trawling in northern Australia: characteristics important for assess-
ing the impacts on populations. Pacific Conservation Biology 7: 55-73.
Fry GC, Milton A, Wassenberg TJ. 2001. The reproductive biology and diet of sea snake
bycatch of prawn trawling in northern Australia: characteristics important for assess-
ing the impacts on populations. Pacific Conservation Biology. 7:55-73.
Fry GC, Milton DA, Wassenberg TJ 2001. The reproductive biology and diet of sea snake
bycatch of prawn trawling in northern Australia: characteristics important for assess-
ing the impacts on populations. Pacific Conservation Biology 7: 55-73
Fuchs AL, Easterling IC, Hengstebeck KC. 2020. Nerodia fasciata (Southern Watersnake).
Diet. Herpetological Revie2w 51(1)149-150.
Fuentes MA, D Escoriza. 2015. Natrix maura (viperine snake) marine foraging. Herpeto-
logical Bulletin 134:31-32.
Fulton AM, Munoz DA. 2018. Thamnophis gigas (Giant Gartersnake). Diet. Herpetological
Review 49(4)764-765.
Galán P. 1988. Segregación ecológica en una comunidad de ofidios. Doñana, Acta Verte-
brata 15:59-78.
Galán P. 2004. Natrix maura (viperine snake). Marine inhabitation. Herpetological Review
35:71.
Galán P. 2012. Natrix maura en el medio marino de las Islas Atlánticas de Galicia. Boletín
de la Asociación Herpetológica Española 23(2):38-43.
Galán P. 2013. Depredación de gato doméstico sobre reptiles en Galicia. Boletín de la Aso-
ciación Herpetológica Española 24(1):103–107.
636
Aquatic Snakes

Galán P, Fernández-Arias G. 1993. Anfibios e Réptiles de Galicia. Xerais, Lugo, Spain.

Galán P, Ferreiro R. 2010. Consumo de una puesta de Alytes obstetricans por Natrix natrix.
Boletín de la Asociación Herpetológica Española 4:14–18.
Gamble T, Daza JD, Colli GR, Vitt LJ, Bauer AM. 2011. A new genus of miniaturized and
pug-nosed gecko from South America (Sphaerodactylidae: Gekkota). Zoological jour-
nal of the Linnean Society. 163(4):1244-66.
Ganança PH, Fraga RD, Vasconcelos Neto LBD, Santos Júnior APD. 2021. A case report
of human intoxication due to a snakebite by the opisthoglyphous dipsadid Tham-
nodynastes lanei Bailey, Thomas & Silva-Jr, 2005. Revista da Sociedade Brasileira de
Medicina Tropical, 54.
Ganesh SR, Chadramouli SR, Sreekar R, Shankar PG. 2013. Reptiles of the central Western
Ghats, India-a reappraisal, and revised checklist, with emphasis on the Agumbe Pla-
teau. Russian Journal of Herpetology 20(3):181-9.
Gans C. 1974. Biomechanics: An approach to vertebrate biology. Lippincott Company, J. B.
Garcia CM, Drummond H. 1990. Population differences in fish-capturing ability of the
Mexican aquatic garter snake (Thamnophis melanogaster). Journal of Herpetology
24(4):412-6.
García G. 1987. Helmintofauna de algunas especies de reptiles de los Pirineos Ibéricos. Te-
sis de Licenciatura, Universidad de Valencia.
García-Antón P, Sánchez-Vialas A, Calvo-Revuelta M. 2017. Un registro destacable de
tamaño en Natrix astreptophora. Boletín de la Asociación Herpetológica Española
28(1):32–34.
García-Cobos D, Crawford AJ, Ramírez-Pinilla MP. 2020. Reproductive phenology in a
Neotropical aquatic snake shows marked seasonality influenced by rainfall patterns.
Journal of Natural History 54(29-30):1845-62.
García-Cobos D, Gómez-Sánchez D, Lynch JD, Molina J. 2018. Characterization, densi-
ty and distribution of mechanoreceptor sensillas on cephalic scales of two species of
snakes inhabiting different aquatic environments. Thesis.
Garcia-Cobos D, Sánchez DA. 2019. Reproductive mode and defensive behaviour of the
South American aquatic snake Helicops pastazae (Serpentes: Dipsadidae). Herpetology
Notes. 12:447-51.
García-Vázquez UO, Mendoza-Hernández AA, Solano-Zavaleta I. 2012. Contribution to
the knowledge of the brood size of Storeria storerioides (Cope, 1865) and Thamnophis
pulchrilatus (Cope, 1884) at Distrito Federal, Mexico. Acta Zoológica Mexicana 28(1):
211-214.
Gardner AS. 2013. The amphibians and reptiles of Oman and the UAE. Frankfurt Contri-
butions of Natural History 58. Edition Chimaira, Frankfurt am M. 480pp.
Gardner CJ, McDonnell N, Ellis C, Jasper LD. 2017. Observations of aquatic behaviour
in Malagasy ground boas Acrantophis madagascariensis (Duméril & Bibron, 1844)
and A. dumerili Jan, 1860. Herpetology Notes. 10:271-3.
Garzón J. 1974. Natrix natrix capturando Leuciscus, Rana y Bufo bufo. Doñana, Acta Ver-
tebrata 1:58.
Gatica-Colima A, Hernandez-Alvarez JC. 2019. Thamnophis marcianus (Checkered Gar-
tersnake) Diet. Herpetological Review 50(3):602-603.
Gaul Jr Rw, Mitchell JC. 2007. The herpetofauna of Dare County, North Carolina: history,
637
Aquatic Snakes

natural history, and biogeography. Journal of the North Carolina Academy of Science.
1:65-109.
Gautschi B, Widmer A, Joshi J, Koella JC. 2002. Increased frequency of scale anomalies and
loss of genetic variation in serially bottlenecked populations of the Dice snake Natrix
tessellata. Conservation Genetics 3:235-45.
Gazzola M, Argentina M, Mahadevan L. 2014. Scaling macroscopic aquatic locomotion.
Nature Physics 10(10):758–761.
Gené J. 1839. Synopsis reptilium Sardiniae indigenoruni. Memorie della Reale Accademia
delle Scienze di Torino, 1: 257-285.
Geniez P. 2015. Serpents d’Europe, d’Afrique du Nord et du Moyen-Orient. Delachaux et
Niestlé, Paris.
Georgalis GL, Smith KT. 2020. Constrictores Oppel, 1811–the available name for the taxo-
nomic group uniting boas and pythons. Vertebrate Zoology 70(3):291-304.
Georgalis G, Villa A, Martin I, Vasilyan D, Delfino M. [Link] amphibians and reptiles
from the Neogene locality of Maramena (Greece), the most diverse European herpeto-
fauna at the Miocene/Pliocene transition boundary. Palaeontologia Electronica 22.3.68
1-99. [Link]
Gezova S, Jablonski D. 2018. Natrix natrix (Grass Snake). Mortality. Herpetological Review
49:348–349.
Gharpurey KG. 1944. The snakes of India and Pakistan. The Popular Book Depot, Bombay,
178 pp.
Gherghel I, Papeş M, Brischoux F, Sahlean T, Strugariu A. A revision of the distribution of
sea kraits (Reptilia, Laticauda) with an updated occurrence dataset for ecological and
conservation research. ZooKeys. 2016(569):135.
Ghira I, Butanescu D, Marosi B. 2009. Feeding behavior of the Dice Snake (Natrix tessella-
ta). Herpetologica Romanica 3:1-7.
Ghizoni Jr IR, TS, Kunz JJ, Cherem, RS Bérnils. 2009. Registros notáveis de répteis de
áreas abertas naturais do planalto e litoral do estado de Santa Catarina, sul do Brasil.
Biotemas 22(3): 129-141.
Ghodke S, Andrews HV. 2002. Recent record of Cantoria violacea (Girard, 1857) from the
North and Middle Andaman Islands, India, with a note on its bite. Hamadryad-Madras
26:371–373.
Ghodke S., Chandi M, Patankar V. 2018. Yellow-banded mangrove snakes (Cantoria vio-
lacea) consume hard-shelled orange signaler crabs (Metaplax elegans). Reptiles & Am-
phibians 25(1):50–51.
Gibbons JW, Dorcas ME. 2004. North American Watersnakes: a Natural History. University
of Oklahoma Press.
Gibbons JW, Scott DE, Ryan TJ, Buhlmann KS, Tuberville TD, Metts BS, Greene JL, Mills T,
Leiden Y, Poppy S, Winne CT. 2000. The global decline of reptiles, de´ja vu amphibians.
BioScience 50:653–666.
Gilbert E, Goodyear J. 2019. Predation of Xenopeltis unicolor (Serpentes: Xenopeltidae) on
Kaloula pulchra (Anura: Microhylidae) in Bangkok, Thailand. Tropical Natural History
19(1):37-8.
Gillett AK, Richard Ploeg, R, Flint, M, Mills PC 2017. Postmortem examination of Austra-
lian sea snakes (Hydrophiinae): Anatomy and common pathologic conditions. Journal
638
Aquatic Snakes

of Veterinary Diagnostic Investigation 2017, Vol. 29(5) 593–611

Gimbel JG, Zahratka JL. 2021. Thamnophis elegans vagrans (Wandering Gartersnake). Diet.
Herpetological Review 52(4):882-83.
Girard C. 1858. Descriptions of some new Reptiles, collected by the US. Exploring Expedi-
tion under the command of Capt. Charles Wilkes, U.S.N. Third Part. Proceedings of the
Academy of Natural Sciences of Philadelphia 9:181–182 [1857].
Giraudo A. 2001. Serpientes de la Selva Paranaense y del Chaco Húmedo. Buenos Aires:
L.O.L.A., 328 p.
Giraudo AR, Arzamendia V, Bellini GP, Bessa CA, Costanzo MB. 2014. Ecology of the large
South American snake, Hydrodynastes gigas (Serpentes: Dipsadidae). Revista mexicana
de biodiversidad 85(4):1206-16.
Giraudo AR, Arzamendia V, Cacciali P. 2006. Geographic variation and taxonomic status of
the southernmost populations of Liophis miliaris (Linnaeus, 1758) (Serpentes: Colubri-
dae). The Herpetological Journal 16(2):213-20.
Giraudo AR, Bellini GP, Arzamendia V. 2014. Comparative ecology of three species of
Thamnodynastes (Serpentes, Dipsadidae) in subtropical-temperate South America.
The Herpetological Journal 24(2):87-96.
Giri VB, Deepak V, Captain A, Das A, Das S, Rajkumar KP, Rathish RL, Gower DJ. 2017.
A new species of Rhabdops Boulenger, 1893 (Serpentes: Natricinae) from the northern
Western Ghats region of India. Zootaxa 4319(1):27-52.
Giri VB.; D J Gower, A. Das, HT Lalremsanga, S. Lalronunga, A. Captain, V. Deepak 2019.
A new genus and species of natricine snake from northeast India. Zootaxa 4603 (2):
241-264.
Glass M, Johansen K. Control of breathing in Acrochordus javanicus, an aquatic snake. Phys-
iological Zoology. 1976 Jul 1;49(3):328-40.
Glauert L. 1950. A handbook of the snakes of Western Australia. Western Australia Natural-
ists Club, Perth, 48 pp.
Glaw F, Nagy ZT, Vences M. 2007. Phylogenetic relationships and classification of the Mala-
gasy pseudoxyrhophiine snake genera Geodipsas and Compsophis based on morphologi-
cal and molecular data. Zootaxa. 2007 Jun 28;1517(1):53-62.
Glaw F, Vences M. 2007. A Fieldguide to the Amphibians and Reptiles of Madagascar.
Third Edition. Vences & Glaw Verlag, Cologne.
Glaw F, Nagy ZT, Köhler J, Franzen M, Vences M. 2009. Phylogenetic relationships of a new
species of pseudoxyrhophiine snake (Reptilia: Lamprophiidae: Thamnosophis) suggest a
biogeographical link between western and northern Madagascar. Organisms Diversity &
Evolution 9(1):13-22.
Gleed-Owen C., 1994. The stomach contents of a grass snake, Natrix natrix, identified from
skeletal remains. British Herpetological Society Bulletin, 50, 34-36.
Glodek GS, Voris HK. 1982. Marine snake diets: prey composition, diversity and overlap.
Copeia, 661-666.
Gloyd HK, Conant R. 1990. Snakes of the Agkistrodon complex. Society for the Study of
Amphibians and Reptiles. Contributions to Herpetology 6:1-614.
Glutz von Blotzheim UN, Bauer KM, Bezzel E. 1971. Handbuch der Vögel Mitteleuropas.
Vol. 4. Aula Verlag, Frankfurt a. M, Germany.
Gmelin JF. 1789. Caroli a Linné Systema Naturae per regna tria naturae secundum classes,
639
Aquatic Snakes

ordines, genera, species, cum characteribus, differentiis, synonymis, locis. Tom I. Pars III
(volume 1 part 3). G.E. Beer, Lipsiae (=Leipzig). 1033–1516 pp.
Gocmen B, Cicek K, Yildiz MZ, Atatur MK, Dincaslan YE, Mebert K. 2011. A preliminary
study on the feeding biology of the Dice snake (Natrix tessellata) in Turkey. Mertensiella
18:365-70.
Gocmen B, Mebert K. 2011. The rediscovery of Natrix tessellata on Cyprus. Mertensiella
18:383-88.
Godinho R, Teixeira J, Rebelo R, Segurado P, Loureiro A, Álvares F, Gomes N, Cardoso P,
Camilo-Alves C, Brito JC. 1999. Atlas of continental Portuguese herpetofauna: an as-
semblage of published and new data. Revista Española de Herpetología 13:61-82.
Goiran C, Bustamante P, Shine R 2017. Industrial Melanism in the Seasnake Emydocephalus
annulatus. Current Biology 27, 2510-2513.
Goldenberg J, D’Alba L, Bisschop K, Vanthournout B, Shawkey MD. 2021. Substrate thermal
properties influence ventral brightness evolution in ectotherms. Communications biolo-
gy 4(1):1-0.
Gomes JF. 1918. Contribuição para o conhecimento dos ofidios do Brasil. III (1). Mem. Inst.
Butantan 1 (1): 57-83.
Gómez JP. 2016, Sistemática do gênero Erythrolamprus Boie 1826 (Serpentes: Dipsadidae)
utilizando dados morfológicos e moleculares (Doctoral dissertation, Universidade de
São Paulo).
Gonzalez RC, Prudente AL, Franco FL. 2014. Morphological variation of Gomesophis
brasiliensis and Ptychophis flavovirgatus (Serpentes, Dipsadidae, Xenodontinae). Sala-
mandra. 50(2):85-98.
González-Fernández A, Manjarrez J, García-Vázquez U, D’Addario M, Sunny A. 2018. Pres-
ent and future ecological niche modeling of garter snake species from the Trans-Mexi-
can Volcanic Belt. PeerJ 6:e4618 [Link]
Gonzalez de la Vega JP, Barnestein JAM, Martínez del Mármol G, Mebert K. 2021. Hybrid-
ization between Natrix astreptophora and Natrix maura: potential cases from Andalusia,
Spain. BAHE [Boletín de la Asociación Herpetológica Española] 32(2):45–53.
Gonzalo A, López P, Martín J. 2008. Avoidance responses to scents of snakes that pose dif-
ferent risks of predation by adult natterjack toads, Bufo calamita. Canadian Journal of
Zoology 86(8):928–932.
Goodyear J, Gilbert E. 2018. First record of male-male combat in Xenopeltis unicolor. Her-
petological Bulletin 2018:143:45.
Gopal B., Krishnamurthy K. (1993) Wetlands of South Asia. In: Whigham D.F., Dyky-
jová D., Hejný S. (eds) Wetlands of the world: Inventory, ecology and management
Volume I. Handbook of vegetation science, vol 15-2. Springer, Dordrecht. [Link]
org/10.1007/978-94-015-8212-4_10
Gopalakrishnakone P, Kochva E. 1990. Venom glands and some associated muscles in sea
snakes. Journal of Morphology 205(1):85–96.
Goris RC, Maeda N. 2004. Guide to the Amphibians and Reptiles of Japan. Krieger, Mala-
bar, 285 pp.
Goris RC. 1971. Geographic variation in color and pattern of Rhabdophis tigrinus (Boie).
Snake. 3(1):57-9.
Gorman G, Licht P, McCollum F. 1981. Annual Reproductive Patterns in Three Species of
640
Aquatic Snakes

Marine Snakes from the Central Philippines. Journal of Herpetology, 15/3: 335-354.
Gove D. 1979. A comparative study of snake and lizard tongue-flicking, with an evolution-
ary hypothesis. Z. Tierpsychol 76:58–76.
Gow GF. 1989. Graeme Gow’s complete guide to Australian snakes. Angus and Robertson,
Sydney, 171 pp.
Graham JB. 1974. Body Temperatures of the Sea Snake Pelamis platurus. Copeia, 1974(2),
531. doi:10.2307/1442545
Gray JE. 1842. Monographic Synopsis of the Water Snakes, or the Family of Hydridae. Zoo-
logical Miscellany 2:59–68.
Gray JE. 1849. Catalogue of the specimens of snakes in the collection of the British Muse-
um. Edward Newman, London, i-xv; 1–125.
Greenbaum E, Portillo F, Jackson K. Kusamba C. 2015. A phylogeny of Central African
Boaedon (Serpentes: Lamprophiidae), with the description of a new cryptic species
from the Albertine Rift. African Journal of Herpetology, 64(1):18-38.
Greenbaum E, Portillo F, Jackson K. Kusamba C. 2015. A phylogeny of Central African
Boaedon (Serpentes: Lamprophiidae), with the description of a new cryptic species from
the Albertine Rift. African Journal of Herpetology, 64(1):18-38.
Greenberg RS, Maldonado JE. 2006. Diversity and endemism in tidal marsh vertebrates.
Vertebrates of Tidal Marshes Studies in Avian Biology No. 32:32-53
Greene BD, Dixon JR, Mueller JM, Whiting MJ, Thornton Jr OW. 1994. Feeding ecolo-
gy of the Concho water snake, Nerodia harteri paucimaculata. Journal of Herpetology
1994:165-72.
Greene BD, Dixon JR, Whiting MJ, Mueller JM. Reproductive ecology of the Concho water
snake, Nerodia harteri paucimaculata. Copeia. 1999 Aug 2:701-9.
Greene HW. 1893. Dietary correlates of the origin and radiation of snakes. American Zoolo-
gist 23(2):431–441.
Gregor J. 1980. Kotázkám termoadaptácie a dynamiky aktivity u dvoch zástupcov rodu Na-
trix (Ophidia: Colubridae). Nepublikovaná rigorozní práce, Bratislava, Slovakia.
Gregory, P.T., Isaac, L.A. (2004): Food habits of the grass snake in southeastern England: is
Natrix natrix a generalist predator? Journal of Herpetology 38: 88–95.
Gressitt JL. 1940. On a collection of amphibians and reptiles made on Hainan Island. Peking
Society Natural History Bulletin 15:175–193.
Gressitt, JL. 1941. Amphibians and reptiles from southeastern China. Philippine Journal of
Science 75:1–58.
Griffin LE. 1909. A list of snakes found in Palawan. Philippine Journal of Science 75: 1–58.
Grillitsch H, Werner YL. 2009. The southern limit of Natrix natrix in the Levant – a detec-
tive story. Herpetozoa 22:65–74.
Gripshover ND, Jayne BC. 2021. Crayfish eating in snakes: Testing how anatomy and
behavior affect prey size and feeding performance. Integrative Organismal Biology
3(1):obab001.
Groot TV, Bruins E, Breeuwer JA. 2003. Molecular genetic evidence for parthenogenesis in
the Burmese python, Python molurus bivittatus. Heredity 90(2):130-5.
Grundler MC, Rabosky DL. 2021. Rapid increase in snake dietary diversity and complexity
following the end-Cretaceous mass extinction. PLoS Biol 19(10): e3001414. [Link]
org/10.1371/[Link].3001414.
641
Aquatic Snakes

Gruschwitz M, Kornacker PM, Podloucky R, Völkl W, Waitzmann M. 1993. Verbreitung,

Ökologie und Schutz der Schlangen Deutschlands und angrenzender Gebiete. Merten-
siella 3, Berlin (Ziegan):7–38.
Gruschwitz M, Lenz S, Mebert K, Laňka V 1999. Natrix tessellata (Laurenti, 1768) –
Würfelnatter. – In: Böhme, W. (Ed.): Handbuch der Reptilien und Amphibien Euro-
pas, Vol. 3/Schlangen II. – AULA-Verlag, Wiesbaden, Germany: 581–644.
Guicking D, Joger U, Wink M. 2008. Molecular phylogeography of the viperine snake
Natrix maura (Serpentes: Colubridae): evidence for strong intraspecific differentia-
tion. Organisms Diversity & Evolution. 8(2):130-45.
Guicking D, Joger U. 2011. Molecular phylogeography of the Dice snake. Mertensiella
18:1-10.
Guicking D, Lawson R, Joger U, Wink M. 2006. Evolution and phylogeny of the genus Na-
trix (Serpentes: Colubridae). Biological Journal of the Linnean Society, 87 (1): 127-143.
Guicking D, RA Griffiths, RD Moore, U Joger, M Wink. 2006a. Introduced alien or perse-
cuted native? Resolving the origin of the viperine snake (Natrix maura) on Mallorca.
Biodiversity and Conservation 15:3045-3054.
Guicking D, R Lawson, U Joger, M Wink. 2006b. Evolution and phylogeny of the genus
Natrix (Serpentes: Colubridae). Biological Journal of the Linnean Society 87:127-143.
Guicking D, U Joger, M Wink. 2002. Molecular phylogeography of the Viperine Snake Na-
trix maura and the Dice Snake Natrix tessalata: first results. Biota 3:49-59.
Guicking D, U Joger, M Wink. 2008. Molecular phylogeography of the viperine snake Na-
trix maura (Serpentes: Colubridae): Evidence for strong intraspecific differentiation.
Organisms Diversity & Evolution 8:130-145.
Guidry EV. 1953. Herpetological notes from southeastern Texas. Herpetologica 1:49-56.
Guinea ML. 1994. Sea snakes of Fiji and Niue. pp. 212-233, In Gopalakrishnakone, P. (ed.).
Sea Snake Toxinology. Venom & Toxin Research Group. National University of Singa-
pore. 251 p.
Guinea ML. 1996. Functions of the cephalic scales of the sea snake Emydocephalus annula-
tus. Journal of Herpetology 30(1):126–128.
Guinea M, Milton D. 2010a. Aipysurus apraefrontalis. The IUCN Red List of Threatened
Species 2010: e.T176770A7301138. [Link] [Link].2010-4.
[Link].
Guinea M, Lukoschek V, Milton D, Courtney T, Fletcher E, White M-D. 2010b. Aipysurus
eydouxii. The IUCN Red List of Threatened Species 2010: e.T176757A7298462. https://
[Link]/ 10.2305/[Link].
Guinea M, Lukoschek V, Milton D, Courtney T. 2010c. Parahydrophis mertoni. The IUCN
Red List of Threatened Species 2010: e.T176772A7301678. [Link]
[Link].
Guinea M, Lukoschek V, Rasmussen A. 2010d. Aipysurus tenuis. The IUCN Red List of
Threatened Species 2010: e.T176760A7299043. [Link]
[Link].
Guinea M, Courtney T. Read M. 2010e. Hydrophis major. The IUCN Red List of Threat-
ened Species 2010: e.T176729A7292011. [Link]
[Link].
Guinea ML, McGrath P. Love B. 1993. Observations of the Port Darwin sea snake Hydre-
642
Aquatic Snakes

laps darwiniensis. Northern Territory Naturalist 14: 28-30.

Guinea ML, McMorrow L, Peerzada N. 1992. Yield and molecular weight of the Duvernoy’s
gland secretions of the dog-faced water snake Cerberus rynchops (Serpentes: Colubri-
dae: Homalopsinae). Toxicon 30:516.
Guinea ML, McMorrow L, Peerzada N. 1992. Yield and molecular weight of the Duvernoy’s
gland secretions of the dog-faced water snake Cerberus rynchops (Serpentes: Colubridae:
Homalopsinae). Toxicon 30:516.
Guinea M, Sanders K, Lobo A. 2010b. Hydrophis gracilis. The IUCN Red List of Threatened
Species 2010: e.T176765A7299914. [Link]
[Link].
Guinea ML, Tamiya N, Cogger HG. 1983. The neurotoxins of the sea snake Laticauda schis-
torhynchus. Biochemical Journal, 213:39–41. doi:10.1042/bj2130039
Guinea ML, Whiting SD. 2005. Insights into the distribution and abundance of sea snakes at
Ashmore Reef. The Beagle: Records of the Museums and Art Galleries of the Northern
Territory, Supplement 1, 199–206.
Günther ACLG. 1864. Reptiles of British India. Ray Society, London.
Günther R, Völkl W. 1996. Ringelnatter - Natrix natrix (LINNAEUS, 1758). In: Günther R.
(Ed.): Die Amphibien und Reptilien Deutschlands. Gustav Fischer Verlag, Jena, Germa-
ny:666–684.
Guo P, Liu Q, Xu Y, Jiang K, Hou M, Ding L, Pyron RA, Burbrink FT. 2012. Out of Asia:
natricine snakes support the Cenozoic Beringian dispersal hypothesis. Molecular phy-
logenetics and evolution 63(3):825-33.
Guo P, Liu Q, Xu Y, Jiang K, Hou M, Ding L, Pyron RA, Burbrink FT. 2012. Out of Asia:
natricine snakes support the Cenozoic Beringian dispersal hypothesis. Molecular phylo-
genetics and evolution 63(3):825-33.
Guo P, Liu SY, Feng JC, He M. 2008. The description of a new species of Thermophis (Ser-
pentes: Colubridae). Sichuan Journal of Zoology 27(3):321.
Guo P, Zhu F, Liu Q, Zhang L, Li JX, Huang YY, Pyron RA. 2014. A taxonomic revision
of the Asian keelback snakes, genus Amphiesma (Serpentes: Colubridae: Natricinae),
with description of a new species. Zootaxa 3873(4):425-40.
Gvozdenović S, Schweiger M. 2014. Melanism in Natrix natrix and Natrix tessellata (Ser-
pentes: Colubridae) from Montenegro. Ecologica Montenegrina 1:231–233.
Gygax, P. 1968. Die Entwicklung der Giftdrüse bei Natrix tessellata. – Rev. Suisse Zool. 75:
549–557.
Gygax, P. 1971. Entwicklung, Bau und Funktion der Giftdrüse (Duvernoy‘s gland) von
Natrix tessellata. – Acta trop. 28: 226–274.
Gyi KK. 1970. A revision of colubrid snakes of the sub-family Homalopsinae. University of
Kansas Publications, Museum of Natural History, 20(2):44–223.
Haagner GV, Carpenter G. 1988. Notes on the reproduction of captive forest cobras, Naja
melanoleuca (Serpentes: Elapidae). The Journal of the Herpetological Association of
Africa 34(1):35-7.
Haagner GV. 1994. A checklist and biogeographic appraisal of the herpetofauna of the Sod-
wana Bay National Park and adjacent lakes area. Durban Museum Novitates. 1994 Jan
1;19(1):30-40.
Hagen B. 1890. Die Pflanzen-und Thierwelt von Deli auf der Ostküste Sumatras. Tijdschrift
643
Aquatic Snakes

Koninklijke Nederlandsche Aadrijksk Genootsch 2:1–240.

Hagen B. 1890. Die Pflanzen-und Thierwelt von Deli auf der Ostküste Sumatras. Tijd-
schrift Koninklijke Nederlandsche Aadrijksk Genootsch 2:1–240.
Hailey A, Davies PMC, Pulford E. 1982. Lifestyle and thermal ecology of natricine snakes.
British Herpetological Journal 6:261-268.
Hailey A, Davies PMC. 1986b. Selection of prey from groups: water snakes and fish. Her-
petological Journal 1:71–77.
Hailey A, PMC Davies. 1986a. Effects of size, sex, temperature, and condition on activity
metabolism and defence behaviour of the viperine snake, Natrix maura. Journal of Zo-
ology, London 208:541-558.
Hailey A, PMC Davies. 1986b. Lifestyle, latitude and activity metabolism of natricine
snakes. Journal of Zoology, London 209:461-476.
Hailey A, PMC Davies. 1986c. Diet and foraging behaviour of Natrix maura. Herpetologi-
cal Journal 1:53-61.
Hailey A, PMC Davies. 1987a. Digestion, specific dynamic action, and ecological energet-
ics of Natrix maura. Herpetological Journal 1:159-166.
Hailey A, PMC Davies. 1987b. Maturity, mating and age-specific reproductive effort of the
snake Natrix maura. Journal of Zoology, London 211:573-587.
Hailey A, Davies PMC. 1987c. Activity and thermoregulation of the snake Natrix maura 1.
R and K thermoregulation. Journal of Zoology, London 213:71–80.
Hailey A, PMC Davies. 1987d. Growth, movement and population dynamics of Natrix
maura in a drying river. Herpetological Journal 1:185-194.
Hall PM, Meier AJ. 1993. Reproduction and behavior of western mud snakes (Farancia
abacura reinwardtii) in American alligator nests. Copeia 1993(1):219-22.
Hall R. 2009. Southeast Asia’s changing palaeogeography. Blumea-Biodiversity, Evolution
and Biogeography of Plants 54(1-2):148–161.
Hall R. 2013. The palaeogeography of Sundaland and Wallacea since the Late Jurassic.
Journal of Limnology 72:1–17
Hall RJ. 1969. Ecological observations on Graham’s watersnake (Regina grahami Baird and
Girard). American Midland Naturalist. 1969 Jan 1:156-63.
Hallas JM, Parchman TL, Feldman CR. 2021. The influence of history, geography, and en-
vironment on patterns of diversification in the western terrestrial garter snake. Journal
of Biogeography 48(1) DOI: 10.1111/jbi.14146 Jun.
Hallas JM, Parchman TL, Feldman CR. 2022. Phylogenomic analyses resolve relationships
among garter snakes (Thamnophis: Natricinae: Colubridae) and elucidate biogeo-
graphic history and morphological evolution. Molecular Phylogenetics and Evolution
167:107374.
Halpern M, Halpern J, Erichsen E & Borghjid S. 1997. The role of nasal chemical senses in
Garter Snake response to airborne odor cues from prey. Journal of Comparative Psy-
chology 111(3):251–260.
Hamidy, A., Zakky, Q., Fitriyana, N., and Endarwin, W. 2023. A new species of water snake
genus Hypsiscopus (Serpentes: Homalopsidae) from Sulawesi, Indonesia. Treubia
50(1): 21-38.
Harding SF, Becker CG, Yates JR, Crump P, Forstner MR, Mullin SJ, Rodriguez D. 2022.
Comparative host–pathogen associations of Snake Fungal Disease in sympatric spe-

644
Aquatic Snakes

cies of water snakes (Nerodia). Scientific reports 12(1):1-2.

Hampton PM. 2011. Comparison of cranial form and function in association with
diet in natricine snakes. Journal of Morphology, 272:1435–1443. doi:10.1002/
jmor.10995PMID:2178015857.
Hampton PM. 2011. Ventral and sub-caudal scale counts are associated with macrohabitat
use and tail specialization in viperid snakes. Evolutionary Ecology 25(2):531-46.
Hanebuth T, Stattegger K, Grootes PM. 2000. Rapid Flooding of the Sunda Shelf: A
Late-Glacial Sea-Level Record. Science 288: 1033–1035.
Hansknecht KA, Burghardt GM. 2010. Stimulus control of lingual predatory luring and
related foraging tactics of Mangrove Saltmarsh snakes (Nerodia clarkii compressicauda).
Journal of Comparative Psychology 124(2):159–165.
Hansknecht KA. 2008. Lingual luring by mangrove saltmarsh snakes (Nerodia clarkii com-
pressicauda). Journal of Herpetology 42(1):9–15.
Harmer AC. 2019. Thamnophis elegans terrestris (Coast Gartersnake). Predation. Herpeto-
logical Review 50(4):814.
Harper F. 1940. Some works of Bartram, Daudin, Latreille, and Sonnini, and their bearing
upon North American herpetological nomenclature. The American Midland Naturalist
23(3):692-723.
Harrington SM, Haan de JM, Shapiro L, Ruane S. 2018. Habits and characteristics of arbore-
al snakes worldwide: arboreality constrains body size but does not affect lineage diversi-
fication. Biological Journal of the Linnean Society 125:61–71.
Harrington SM, Reeder TW. 2017. Phylogenetic inference and divergence dating of snakes
using molecules, morphology and fossils: new insights into convergent evolution
of feeding morphology and limb reduction. Biological Journal of the Linnean Soci-
ety 121(2):379–394.
Hart KM, Cherkiss MS, Smith BJ, Mazzotti FJ, Fujisaki I, Snow RW, Dorcas ME. 2015.
Home range, habitat use, and movement patterns of non-native Burmese pythons in
Everglades National Park, Florida, USA. Animal Biotelemetry 2015 (1):8.
Hart KM, Schofield PJ, Gregoire DR. 2012. Experimentally derived salinity tolerance of
hatchling Burmese pythons (Python molurus bivittatus) from the Everglades, Florida
(USA). Journal of Experimental Marine Biology and Ecology. 10:413:56-9.
Hart NS, Coimbra JP, Collin SP & Westhoff G. 2012. Photoreceptor types, visual pigments,
and topographic specializations in the retinas of hydrophiid sea snakes. Journal of Com-
parative Neurology 520(6):1246–1261.
Harzhauser M, Tempfer PM. 2004. Late Pannonian wetland ecology of the Vienna Basin
based on molluscs and lower vertebrate assemblages (Late Miocene, MN 9, Austria).
Courier-Forschungsinstitut Senckenberg. 2004:55-68.
Hauzman E, Bonci DMO, Suárez-Villota EY, Neitz M & Ventura DF. 2017. Daily activity
patterns influence retinal morphology, signatures of selection, and spectral tuning of op-
sin genes in colubrid snakes. BMC Evolutionary Biology 17(1):1–14.
Head, Jason J. 2005. Snakes of the Siwalik Group (Miocene of Pakistan): Systematics and
Relationship to Environmental Change, Palaeontologia Electronica Vol. 8, Issue 1;
18A:33p, 2.5MB; [Link] /paleo/2005_1/head18/iue1_05.htm
Head JJ. 2015. Fossil calibration dates for mlecular phylogenetic analysis of snakes 1:
Serpentes, Alethinophidia, Boidae, Pythonidae. Palaeontologia Electronica 18.1.6FC;
645
Aquatic Snakes

1-17.
Head JJ, Bloch JI, Hasting AK, Bourque JR, Cadena EA, Herrera FA, Polly PD, Jaramillo
CA. 2009. Giant boid snake from the Paleocene neotropics reveals hotter past equato-
rial temperatures. Nature 457: 715–718.
Head, J. J., Mohabey, D. M., & Wilson, J. A. (2007). Acrochordus Hornstedt (Serpentes,
Caenophidia) from the Miocene of Gujarat, Western India: temporal constraints on disper-
sal of a derived snake. Journal of Vertebrate Paleontology, 27(3), 720–723. doi:10.1671/027
2-4634(2007)27[720:ahscft][Link];2
Healy K, Carbone C & Jackson AL. 2019. Snake venom potency and yield are associated
with prey-evolution, predator metabolism and habitat structure. Ecology Letters 22:527–
537.
Heatwole H, Busack S, Cogger H. 2005. Geographic Variation in Sea Kraits of the Laticauda
colubrina Complex (Serpentes: Elapidae: Hydrophiinae: Laticaudini). Herpetological
Monographs, 19: 1-136.
Heatwole H, Guinea M. 1993. Fauna Of Australia Family Laticuadidae. AGPS Canberra,
37: 1-8.
Heatwole H, Seymour RS. 1975. Pulmonary and cutaneous oxygen uptake in sea snakes
and a file snake. Comparative Biochemistry and Physiology A, Comparative Physiology
51:399–405.
Heatwole H. 1977. Voluntary submergence time and breathing rhythm in the homalopsine
snake, Cerberus rhynchops. Australian Zoology 19(2):155–167.
Heatwole H. 1999. Sea snakes. University of New South Wales Press Ltd.
Heatwole, H. And Guinea, M. 1993. Fauna Of Australia Family Laticuadidae. AGPS Can-
berra, 37: 1-8.
Hebard WB. 1951. Notes on the life history of the Puget Sound garter snake, Thamnophis
ordinoides. Herpetologica 7(4):177-9.
Hecht G. 1930. Systematik, Ausbreitungsgeschichte und Ökologie der europäischen Arten
der Gattung Tropidonotus (Kuhl) H. Boie. Mitteilungen aus dem Zoologischen Museum
in Berlin, 16: 244-393.
Hediger 1971. Die Ringelnatter. In: Grzimeks Tierleben VII, Kriechtiere. Deutscher Taschen-
buch Verlag (DTV), Munich, Germany.
Henderson RW, Hoevers LG. The seasonal incidence of snakes at a locality in northern Be-
lize. Copeia. 1977 May 25:349-55.
Henderson RW, Nickerson MA, Ketcham S. Short term movements of the snakes Chironius
carinatus, Helicops angulatus and Bothrops atrox in 1976. Peru. Herpetologica 304-10.
Henderson RW, Powell R, editors. 2007. Biology of the Boas and Pythons. Eagle Mountain
Publishing.
Hendrickson KE. The waters of the Brazos: a history of the Brazos River Authority, 1929-
1979. Texian Press; 1981.
Hendrickson, KE. 2001. Handbook of Texas Online, Brazos River. The Texas State Histori-
cal Association, 1997-2001. [Link]
BB/[Link]
Hennig J, Breitfeld HT, Gough A, Hall R, Long TV, Kim VM., Quang SD. 2018. U-PB
Zircon Ages and Provenance of Upper Cenozoic sediments from the Da Lat Zone, SE
Vietnam: Implications for an Intra-Miocene unconformity and paleo-drainage of the
646
Aquatic Snakes

Proto–Mekong River. Journal of Sedimentary Research 88:495–515.

Hennig-Breitfeld J, Breitfeld HT, Hall R, BouDagher-Fadel M, Thirlwall M. 2019. A new
upper Paleogene to Neogene stratigraphy for Sarawak and Labuan in northwestern Bor-
neo: Paleogeography of the eastern Sundaland margin. Earth-Science Reviews 190:1–32.
Heo JH, Lee HJ, Kim IH, Fong JJ, Kim JK, Jeong S, Park D. 2014. Can an Invasive Prey
Species Induce Morphological and Behavioral Changes in an Endemic Predator? Ev-
idence from a South Korean Snake (Oocatochus rufodorsatus). Asian Herpetological
Research 5(4):245-54.
Herault J, Segall M, Clement E, Xie X, Lebastard V & Boyer F. (under review). Stability
mechanisms of aquatic snakes on water: from the biological aspects to the bio-inspired
robot. Bioinspiration and Biomimetics.
Hernández-Camacho J.I, Álvarez-León R, Renjifo-Rey JM. 2006. Pelagic sea snake Pelamis
platurus (Linnaeus, 1766) (Reptilia: Serpentes: Hydrophiidae) is found on the Caribbean
Coast of Colombia.
Herrel A, Vincent SE, Alfaro ME, Van Wassenbergh S, Vanhooydonck B, Irschick DJ. 2008.
Morphological convergence as a consequence of extreme functional demands: examples
from the feeding system of natricine snakes. Journal of Evolutionary Biology 21(5):1438-
48
Hibbitts TJ, Painter CW, Holycross AT. 2009. Ecology of a population of the narrow-head-
ed garter snake (Thamnophis rufipunctatus) in New Mexico: catastrophic decline of a
river specialist. The Southwestern Naturalist, 54(4):461-8.
Hierink F, Bolon I, Durso AM, Ruiz de Castañeda R, Zambrana-Torrelio C, Eskew EA, Ray
N. (2020). Forty-four years of global trade in CITES-listed snakes: Trends and implica-
tions for conservation and public health. Biological Conservation, Volume 248, 108601.
Hill JG, Beaupre SJ. 2008. Body size, growth, and reproduction in a population of Western
Cottonmouths (Agkistrodon piscivorus leucostoma) in the Ozark Mountains of northwest
Arkansas. Copeia 2008(1):105-14.
Hoang AQ, Tu MB, Takahashi S, Kunisue T, Tanabe S. Snakes as bimonitors of environ-
mental pollution: A review on organic contaminants. Science of The Total Environment.
2021 Jan 20:144672.
Hoesel JKP. 1959. Ophidia Javanica. Kebun Raya, Indonesia: Museum Zoologicum Bo-
goriense.
Hoffstetter R, Rage JC. 1977. Le gisement de vertébrés miocènes de La Venta (Colombie)
et sa faune de serpents. Annales de Paléontologie (Vertébrés). 63: 161–190.
Hoffstetter R. 1939. Contribution à l’étude des Elapidæ actuels et fossiles et de l’ostéologie
des Ophidiens. Archives du Muséum d’Histoire Naturelle de Lyon 15:1–78.
Holm O. 1934. Vanliga snokens utbredning I Norland. Fauna och Flora 29:241–257.
Holman JA. 2000. Fossil snakes of North America: origin, evolution, distribution, paleoecol-
ogy. Indiana University Press.
Hoogerwerf A. 1954. Rapport over een naar Komodo, Padar en Rintja gemachte dienstreis
van 21 Mei–6 Juli, 1953. Unpublished manuscript, Museum Bororiense (Bogor, Java).
[Not seen, cited in Auffenberg.]
Hoogmoed MS. 1982. Snakes of the Guianan region. Memórias do Instituto Butantan
46:219-54.
Hoogmoed MS. 1985. Xenodon werneri Eiselt, a poorly known snake from Guiana, with
647
Aquatic Snakes

notes on Waglerophis merremii (Wagler)(Reptilia: Serpentes: Colubridae). Notes on the

herpetofauna of Surinam IX. Zoologische Mededelingen 59(8):79-88.
Hoorn C, Wesselingh FP, Hovikoski J, Guerrero J. 2010. The development of the Amazo-
nian mega-wetland (Miocene; Brazil, Colombia, Peru, Bolivia). Amazonia, Landscape
and Species Evolution: A Look into the Past. Blackwell-Wiley, Hoboken, 123-142.
Hoorn C, Wesselingh FP, Hovikoski J, Guerrero J. 2010. The development of the Amazonian
mega-wetland (Miocene; Brazil, Colombia, Peru, Bolivia). Amazonia, Landscape and
Species Evolution: A Look into the Past. Blackwell-Wiley, Hoboken, 123-142.
Hoser RT. A new species of Raclitia Gray (Serpentes, Homalopsidae) from Peninsular Ma-
laysia. Australasian Journal of Herpetology 49:24–25.
Houssaye A, Fish FE. 2016. Functional (secondary) adaptation to an aquatic life in ver-
tebrates: an introduction to the symposium. Integrative and comparative biology
56(6):1266-70.
Houssaye A, Herrel A, Boistel R, Rage JC. 2019. Adaptation of the vertebral inner struc-
ture to an aquatic life in snakes: Pachyophiid peculiarities in comparison to extant and
extinct forms. Comptes Rendus Palevol 18(7):783-99.
Houssaye A. 2012. Bone histology of aquatic reptiles: what does it tell us about secondary
adaptation to an aquatic life? Biological Journal of the Linnean Society 108(1):3-21.
Houston D, Shine R. 1994. Low growth rates and delayed maturation in Arafura File-
snakes (Serpentes: Acrochordidae) in tropical Australia. Copeia 1994: 726-731.
Houston D, Shine R. Movements and activity patterns of Arafura filesnakes (Serpentes:
Acrochordidae) in tropical Australia. Herpetologica. 1994 Sep 1:349-57.
Hsiang AY, Field DJ, Webster TH, Behlke AD, Davis MB, Racicot RA, Gauthier JA. 2015.
The origin of snakes: revealing the ecology, behavior, and evolutionary history of early
snakes using genomics, phenomics, and the fossil record. BMC Evolutionary Biolo-
gy, 15, 87. doi:10.1186/s12862-015-0358-5
Hsiou AS, Albino AM. 2009. Presence of the genus Eunectes (Serpentes, Boidae) in the
Neogene of Southwestern Amazonia, Brazil. Journal of Herpetology. 43 (4): 612–619.
doi:10.1670/08-295.1.
Hugall AF, Foster R, Lee MS. 2007. Calibration choice, rate smoothing, and the pattern of
tetrapod diversification according to the long nuclear gene RAG-1. Systematic Biolo-
gy. 56(4):543-63.
Hughes J.M.R. Freshwater Ecology and Conservation: Approaches and Techniques (2019).
Publisher: Oxford University Press. ISBN: 978-0-19-876642-1
Hughes TP., AH Baird, DR Bellwood, M. Card, SR Connolly, C Folke, R Grosberg, O Hoegh
Guldberg, JBC Jackson, J. Kleypas, et al. 2003. Climate change, human impacts, and the
resilience of coral reefs. Science 301 (5635):929–933
Hutchison CS. 1989. Geological Evolution of South-east Asia. Claredon Press, Oxford,
UK.
Hutinec BJ, Mebert K. 2011. Ecological partitioning between dice snakes (Natrix tessella-
ta) and grass snakes (Natrix natrix) in southern Croatia. Mertensiella. 18:225-33.
Igan D. 2008. Snakes of Arabia: a field guide to snakes of the Arabian Peninsula and its
shores, Motivate Publishing, ISBN 1860632394
Inam A, Clift PD, Giosan L, Tabrez AR, Tahir M, Rabbani MM, Danish M. 2007. The geo-
graphic, geological and oceanographic setting of the Indus River. Pp. 333–345. In: Large
648
Aquatic Snakes

rivers: geomorphology and management. Gupta A (ed). John Wiley and Sons, Chiches-
ter, UK.
Ineich I, Laboute P. 2002. Sea Snakes Of New Caledonia. Paris, France. IRD Orstom
Ineich I. 1996. - Geographic Distribution. Serpentes. Hydrophis torquatus diadema (West
Coast Blackhead Sea Snake). Herpetological Review, 27(3): 154.
Ioannidis Y, Mebert K. 2011. Habitat preferences of Natrix tessellata at Strofylia, NW Pelo-
ponnese, and comparison to syntopic N. natrix. Mertensiella 18:302–310.
Iskandar DT. 1979. A second specimen of the Matanna water snake, Enhydris matannen-
sis (Blgr.), from Raha, Muna Island, Indonesia. British Journal of Herpetology 5(12):
849–850.
Iskandar DT. 1987. The occurrence of Enhydris alternans at Java The Snake 19:72–73.
Iskandar DT. 2004. The amphibians and reptiles of Malinau Region, Bulungan Research
Forest, East Kalimantan: annonated checklist with notes on the ecological presence of
the species and local utilization. Center for International Forestry Research, Jakarta, 27
pp.
Islam SN. 2010. Threatened wetlands and ecologically sensitive ecosystems management
in Bangladesh. Frontiers of Earth Science in China, 4(4): 438-448.
IUCN 2020. The IUCN Red List of Threatened Species. Version 2020-3. [Link]
[Link].
Ivanov M. 2000. Snakes of the lower/middle Miocene transition at Vieux Collonges (Rhône,
France), with comments on the colonisation of western Europe by colubroids. Geodiver-
sitas 22(4):559-88.
Ivanov M. 2002. The oldest known Miocene snake fauna from Central Europe: Merkur−
North locality, Czech Republic. Acta Palaeontologica Polonica 47 (3): 513–534.
Ivanov M., 2001. Changes in the composition of the European snake fauna during the Early
Miocene and at the Early/ Middle Miocene transition. Paläontologische Zeitschrift, 74:
563-573.
Jablonski D., Kautman J. 2017. Melanism in Natrix tessellata (Serpentes: Colubridae) from
Slovakia. Herpetology Notes 10: 173-75.
Jablonski D, Trapp B, Tzoras E, Mebert K. 2022. Erythrism in the Eastern Grass Snake, Na-
trix natrix (Linnaeus, 1758). Herpetozoa 35: 213–217.
Jablonski D, Asztalos M, Yılmaz C, Avcı A. 2023. The range-wide mitochondrial lineage of
Natrix natrix scutata (Pallas, 1771) presented in the northern Zagros Mountains. Evolu-
tionary Systematics 7: 67–71.
Jablonski D, Khalili F, Masroor R. 2023b. The herpetofaunal diversity of Takhar Province,
Afghanistan. Herpetozoa 36:73–90.
Jablonski D, Mebert K, Masroor R, Simonov E, Kukushkin O, Abduraupov T, Hofmann S.
2023a. The Silk roads: phylogeography of Central Asian dice snakes (Serpentes: Natrici-
dae) shaped by rivers in desert and mountain valleys. Current Zoology: online version at
[Link]
Jablonski D., Tzoras E., Panagiotopoulos A., Asztalos M., Fritz U. 2023c. Genotyping the
phenotypic diversity in Aegean Natrix natrix moreotica (Bedriaga, 1882) (Reptilia, Ser-
pentes, Natricidae). ZooKeys 1169:87–94. [Link] /zookeys.1169.104594
Jackson K, Zassi-Boulou AG, Mavoungou LB, Pangou S. 2007. Amphibians and reptiles of
the Lac Télé community reserve, Likouala region, Republic of Congo (Brazzaville). Her-
649
Aquatic Snakes

petological Conservation and Biology (2):75-86.

Jaén MJ, V Pérez-Mellado. 1989. Temperaturas corporales y ritmos de actividad en una po-
blación de Natrix maura (L.) del Sistema Central. Doñana, Acta Vertebrata 16:203-217.
Jan G. 1861. Uber die familie Homalopsidae. Archiv für Naturgeschichte 27:87–95.
Jan G. 1863. Elenco sistematico degli ofidi, descritti e disegnati per l’iconografia générale.
Tipografia di A. Lombardi, Milan.
Jan G. 1864. Iconographie générale des ophidiens. Atlas. Georges Jan and 622 Ferdinand
Sordelli, Paris, France.
Jan G. 1865. Enumerazione sistematica degli ofidi appartenenti ai gruppo Potamophilidae.
Archive per la Zoologia, l’Anatomia et la Fisiologia 3 (2):201–265.
Janecka MJ, Janecka JE, Haines AM, Michaels A, Criscione CD. Post-delisting genetic mon-
itoring reveals population subdivision along river and reservoir localities of the endemic
Concho water snake (Nerodia harteri paucimaculata). Conservation Genetics. 2021
Dec;22(6):1005-21.
Janev Hutinec B, Mebert K. 2011. Ecological partitioning between Dice snakes (Natrix tes-
sellata) and Grass snakes (Natrix natrix) in southern Croatia. Mertensiella 18:225-34.
Jansen DW. 1987. The myonecrotic effect of Duvernoy’s gland secretion of the snake Tham-
nophis elegans vagrans. Journal of Herpetology 21(1):81-3.
Jansen KP, Mushinsky HR, Karl SA. 2008. Population genetics of the mangrove salt marsh
snake, Nerodia clarkii compressicauda, in a linear, fragmented habitat. Conservation Ge-
netics 9(2):401-10.
Janssen I, Völkl W. 2008. Gibt es räumlich und zeitlich getrennte Teilhabitate bei der Ringel-
natter (Natrix natrix LINNAEUS, 1758). Mertensiella 16:162–172.
Jovanovic O, Vences M, Safarek G, Rabemananjara FC, Dolch R. 2009. Predation upon
Mantella aurantiaca in the Torotorofotsy wetlands, central-eastern Madagascar. Herpe-
tology Notes. 2(1):95-7.
Jayne BC, Voris HK, Heang KB. 1988. Diet, feeding behavior, growth, and numbers of a
population of Cerberus rynchops (Serpentes: Homalopsinae) in Malaysia: a contribution
in celebration of the distinguished scholarship of Robert F. Inger on the occasion of his
sixty-fifth birthday. Fieldiana. Zoology 50:1–15.
Jayne BC, Voris HK, Ng PK. 2002. Snake circumvents constraints on prey size. Nature
418(6894):143.
Jayne BC, Voris HK, Ng PK. 2018. How big is too big? Using crustacean-eating snakes
(Homalopsidae) to test how anatomy and behaviour affect prey size and feeding perfor-
mance. Biological Journal of the Linnean Society 123(3):636–650.
Jayne BC, Ward TJ, Voris HK. 1995. Morphology, reproduction, and diet of the marine
homalopsine snake Bitia hydroides in peninsular Malaysia. Copeia 21:800–808.
Jayne BC, Ward TJ, Voris HK. 1995. Morphology, reproduction, and diet of the marine
homalopsine snake Bitia hydroides in peninsular Malaysia. Copeia 21:800–808.
Jayne BC. 1985. Swimming in constricting (Elaphe g. guttata) and nonconstricting (Nerodia
fasciata pictiventris) colubrid snakes. Copeia 1985(1):195–208.
Jelić, Dusan & Suvad Lelo 2011. Distribution and status Quo of Natrix tessellata in Croatia,
and Bosnia and Herzegovina. Mertensiella 18:217-24.
Jelic D, Lauš B. 2011. The gull Larus cachinnans (Pallas, 1811) as natural predator of Natrix
tessellata (Laurenti, 1768). Mertensiella 18:451–452.
650
Aquatic Snakes

Jelić D, Lelo S. 2011. Distribution and status Quo of Natrix tessellata in Croatia, and Bosnia
and Herzegovina. Mertensiella 18:217–224.
Jha S. 2008. Status and conservation of lowland Terai wetlands in Nepal. Our Nature.
2008;6(1):67-77.
Johnson S, Francois K. 2018. First record of an Eastern Ribbonsnake (Thamnophis sauritus)
from Paradise Island, The Bahamas. Reptiles & Amphibians 25(3):214.
Junqueira-de-Azevedo IL, Campos PF, Ching AT, Mackessy SP. 2016. Colubrid venom
composition: an-omics perspective. Toxins 8(8):230.
Juszczyk W. 1974. Plazy I gady krajowe. PWN, Warsaw, Poland, 727 pp
Kabisch K. 1978. Die Ringelnatter. Ziemsen-Verlag (Neue Brehm Bücherei), Wittenberg,
Germany.
Kabisch K. 1999. Natrix natrix (Linnaeus, 1758) - Ringelnatter. Pp.482–815 In: Böhme W.
(Ed. Handbuch der Reptilien und Amphibien Europas, Band 3/IIA: Schlangen (Serpentes)
II: Colubridae 2 (Boiginae, Natricinae). Aula-Verlag, Wiebelsheim, Germany.
Kalboussi M, Achour H. 2018. Modelling the spatial distribution of snake species in north-
western Tunisia using maximum entropy (Maxent) and Geographic Information System
(GIS). Journal of Forestry Research 29:233–245.
Kalki Y, Morrill DP, Schramer TD, West TR, Feng CY, Wylie DB. 2018. A dietary synopsis of
Nerodia rhombifer, including a novel prey item. Bulletin of the Chicago Herpetological
Society 53(12):249-52.
Kammel W, Mebert K. 2011. Effects of rehabilitation of the polluted river system Mur in
Styria in Austria and construction of hydroelectric power plants on fish fauna and distri-
bution of the Dice snake. Mertensiella 18:188-97.
Kammerer JC. 1987. Largest river in the United States. USGS U.S Geological Survey, Water
Resources Division MS 419, National Center Reston, Virginia 22092.
Karns DR, Lukoschek V, Osterhage J, Murphy JC, Voris HK. 2010a. Phylogeny and biogeog-
raphy of the Enhydris clade (Serpentes: Homalopsidae). Zootaxa 2452(1):18–30.
Karns DR, Murphy JC, Voris HK, Suddeth JS. 2005. Comparison of semi-aquatic snake
communities associated with the Khorat Basin, Thailand. Tropical Natural History
5(2):73-90.
Karns DR, Murphy JC, Voris HK. 2010. Semi-aquatic snake communities of the central
plain region of Thailand. Tropical Natural History 10(1):1-25.
Karns DR, Voris HK, Chan-ard T, Goodwin JC, Murphy JC. 2000. The spatial ecology of the
rainbow water snake, Enhydris enhydris (Homalopsinae) in southern Thailand. Herpeto-
logical Natural History 7(2):97-115.
Karns DR, Voris HK, Goodwin TG. 2002. Ecology of Oriental-Australian rear-fanged
water snakes (Colubridae: Homalopsinae) in the Pasir Ris Park mangrove forest, Sin-
gapore. Raffles Bulletin of Zoology 50(2):487-98.
Karsen SJ, Lau MW, Bogadek A. 1986. Hong Kong amphibians and reptiles. Urban Council,
Hong Kong, 136 pp.
Karthikeyan R, Balasubramanian T. 2007. Species Diversity of Sea Snake (Hydrophiidae)
Distributed in the Coramantal Coast (East Coast of India). International Journal of Zoo-
logical Research, 3: 107-131.
Karunarathna S, Surasinghe T, Botejue M, Madawala M. 2018. Gerarda prevostiana (Ser-
pentes: Homalopsidae) in Sri Lanka: distribution and behaviour. The Herpetological
651
Aquatic Snakes

Bulletin 145:8–13.
Kawashita-Ribeiro RA, Ávila RW, Morais DH. 2013. A new snake of the genus Helicops
Wagler, 1830 (Dipsadidae, Xenodontinae) from Brazil. Herpetologica 69(1):80-90.
Kelly CM, Branch WR, Broadley DG, Barker NP, Villet MH. 2011. Molecular systematics
of the African snake family Lamprophiidae Fitzinger, 1843 (Serpentes: Elapoidea), with
particular focus on the genera Lamprophis Fitzinger 1843 and Mehelya Csiki 1903.
Molecular Phylogenetics and Evolution. 58(3):415-26.
Keogh JS, Branch WR, Shine R. 2000. Feeding ecology, reproduction and sexual dimor-
phism in the colubrid snake Crotaphopeltis hotamboeia in southern Africa. African
Journal of Herpetology 49(2):129-37.
Kerford, M. 2005. The Ecology of the Bar Bellied Sea Snake (Hydrophis elegans) In Shark
Bay, Western Australia. PhD diss., Biological Sciences Department-Simon Fraser Uni-
versity, 2005.
Keswall, [sic]. 1886. Waters of Western India. Part 2. Journal of the Bombay Natural Histo-
ry Society 1:153–175.
Key KN, Banta JA, Gluesenkamp AG, Hertweck KL, Placyk JS. 2019. Conservation biology
of a southwestern endemic, the Texas garter snake, Thamnophis sirtalis annectens: an
integrative perspective. The Southwestern Naturalist 64(1):8-19.
Khan AA, Arshad SA. Wetlands of Pakistan: distribution, degradation, and management.
Pakistan Geographical Review. 2014;69(1):28-45.
Khan MS. 2002. A guide to the snakes of Pakistan. Edition Chimaira, Frankfurt am Main,
141 pp.
Kharin VE 2012. Sea Snakes of the Genus Chitulia Gray, 1849 (Serpentes: Hydrophiidae).
The Taxonomic Status of Intrageneric Complexes. Russian Journal of Marine Biolog 38
(6): 456-458
Kharin VE, Cheblukov VP. 2007. On first reliable record of the sea snake Chitulia belcheri
(Gray, 1849) from Australian waters, with notes on species composition and taxonomic
status of the genus Chitulia (Serpentes, Hydrophiidae). Russian Journal of Marine Biol-
ogy, 33(3), 161– 165. doi:10.1134/s1063074007030030
Kharin VE, Hallermann J. 2010. A remote new distribution record and a redescription of a
rare and little-known sea snake, Leioselasma czeblukovi Kharin, 1984 (Hydrophiidae).
Herpetology Notes 3: 315-319.
Kharin VE. 1984. Sea snakes of the genus Hydrophis sensu lato (Serpentes, Hydrophiidae)
on taxonomic status of the New Guinea H. obscurus [in Russian]. Zoologicheskii Zhur-
nal 63 (4) : 630-632.
Kharin VE. 1985. A new species of sea snakes of the genus Enhydrina (Serpentes, Hy-
drophiidae) from waters of New Guinea. [in Russian]. Zoologicheskii Zhurnal 64 (5):
785-787.
Kharin VE. 2004. Review of sea snakes of the genus Hydrophis sensu stricto (Serpentes:
Hydrophiidae). Russian Journal of Marine Biology, 30(6), 387–394. doi:10.1007/
s11179-005-0004-y
Kharin VE. 2005. On new findings of a rare sea snake Leioselasma czeblukovi Kharin,
1984, with remarks on species composition and taxonomical position of the genus
Leioselasma Lacepede, 1804, (Serpentes: Hydrophiidae). Russian Journal of Marine
Biology 31: 269-272.
652
Aquatic Snakes

Kieffer Merki M-L, Bolzern H, Gemsch J, Strickler P. 2018. Die Reptilien des Kantons
Luzern. Bau-, Umwelt- und Wirtschaftsdepartement, Landwirtschaft und Wald (lawa),
Sursee, Switzerland, pp. 27.
Kindler C, Fritz U. 2018. Phylogeography and taxonomy of the barred grass snake (Natrix
helvetica), with a discussion of the subspecies category in zoology. Vertebrate Zoology
68(3):253–267.
Kindler C, Böhme W, Corti C, Gvoždík V, Jablonski D, Jandzik D, Metallinou M, Široký
P, Fritz U. 2013. Mitochondrial phylogeography, contact zones and taxonomy of grass
snakes (Natrix natrix, N. megalocephala). Zoologica Scripta 42:458–472.
Kindler C, Bringsøe H, Fritz U. 2014. Phylogeography of grass snakes (Natrix natrix) all
around the Baltic Sea: implications for the Holocene colonization of Fennoscandia. Am-
phibia-Reptilia 35(4):413-24.
Kindler C, Chèvre M, Ursenbacher S, Böhme W, Hille A, Jablonski D, Vamberger M, Fritz
U. 2017. Hybridization patterns in two contact zones of grass snakes reveal a new Cen-
tral European snake species. Scientific Reports 7:7378.
Kindler C, De Pous P, Carranza S, Beddek M, Geniez P, Fritz U. 2018a. Phylogeography of
the Ibero-Maghrebian red-eyed grass snake (Natrix astreptophora). Organisms Diversity
and Evolution 18:143–150.
Kindler C, Graciá E, Fritz U. 2018b. Extra-Mediterranean glacial refuges in barred and com-
mon grass snakes (Natrix helvetica, N. natrix). Scientific Reports 8:1821.
King RB. 1986. Population ecology of the Lake Erie Water Snake, Nerodia sipedon insula-
rum. Copeia 1986: 757–772.
Kinghorn JR. 1929. The Snakes of Australia. Angus and Robertson, Sydney.
Kirillov AA., Kirillova NYu. 2019. Comparative analysis of the helminth fauna of Natrix
natrix and Natrix tessellata (Reptilia, Colubridae) in the Samarskaya Luka National Park
(Russia). Nature Conservation Research. Conservation Science 4(4): 12–25.
Kishida T & Hikida T. 2010. Degeneration patterns of the olfactory receptor genes in sea
snakes. Journal of Evolutionary Biology 23(2):302–310.
Kishida T, Go Y, Tatsumoto S, Tatsumi K, Kuraku S & Toda M. 2019. Loss of olfaction in sea
snakes provides new perspectives on the aquatic adaptation of amniotes. Proceedings of
the Royal Society B: Biological Sciences 286(1910).
Kishida T, Toda M, Go Y, Tatsumoto S, Sasai T, Hikida T. 2021. Population history and
genomic admixture of sea snakes of the genus Laticauda in the West Pacific. Molecular
Phylogenetics and Evolution 155:107005.
Knight A., Mindell DP. 1994. On the phylogenetic relationship of Colubrinae, Elapidae,
and Viperidae and the evolution of front-fanged venom systems in snakes. Copeia
1994(1):1–9.
Kofron CP. 1978. Foods and habitats of aquatic snakes (Reptilia, Serpentes) in a Louisiana
swamp. Journal of Herpetology 30:543-54.
Köhler G, Khaing KPP, Than NL, Baranski D, Schell T, Greve C, Janke A, Pauls SE. 2021. A
new genus and species of mud snake from Myanmar (Reptilia, Squamata, Homalopsi-
dae). Zootaxa 4915 (3):301–325.
Kojima Y, Mori A. 2014. Home range and movements of Rhabdophis tigrinus in a mountain
habitat of Kyoto, Japan. Current herpetology 33(1):8-20.
Koller N, S. Ursenbacher. 1999. Estimation de l’effectif de Couleuvres vipérines (Natrix
653
Aquatic Snakes

maura) et de Couleuvres tesselées (N. tessellata) dans le Lavaux, “in: Annales du Col-
loque “”Découvrir le Léman”” (eds. Bertola C., Goumand C., Rubin J.-F.), Musée du
Léman & Editions Slatkine, Genève”
Koo KS, Chang MH, Song JY. 2019. Report on the group mating of Asian keelback snake
Hebius vibakari Boie 1826 (Squamata; Colubridae) in South Korea. Korean Journal
of Environmental Biology 37(1):88-92.
Kopstein F. 1926. V. Reptilien von der Molukken und den benachbarten Inseln. Zoolo-
gische Mededelingen, Leiden 9:71–122.
Kopstein F. 1931. Fordonia leucobalia Schegel und Cerberus rhynchops Schneider. Treubia
13(1):1–4.
Kotagama SW, Bambaradeniya CN. 2006. An overview of the wetlands of Sri Lanka and
their conservation significance. National Wetlands Directory of Sri Lanka :7-16.
Kotenko TI, Shaitan SV, Starkov VG, Zinenko OI. 2011. The northern range limit of the
Dice snake (Natrix tessellata) in Ukraine and the Don River Basin in Russia. Mertensi-
ella 18:311-25.
Kramer E., 1970. Revalidierte und neue Rassen der europäischen Schlangenfauna. Lavori
Soc. ital. Biogeogr., (n.s.) 1 (1970): 667-676.
Kramer E, Schnurrenberger H. 1963. Systematik, Verbreitung und Ökologie der Libyschen
Schlangen. Revue Suisse de Zoologie. 70(3):453–568
Kreiner G., 2007. The Snakes of Europe. Edition Chimaira, Frankfurt, 317 pp.
Kuhl H, van Hasselt JC. 1822. Uittreksels uit brieven van den heeren Kuhl en van Hasselt
aan de Heeren C.J. Temminck, Th. Van Swinderen en W. de Haan. Algemeene Kunsten
Letter Bode voor het jaar 1822. 1:99–104.
Kumar AB, Captain A. 2011. Recent records of the endemic Kerala mud snake, Enhydris
dussumieri (Duméril, Bibron & Duméril, 1854) from India. Current Science 100:928–
931.
Kumar AB, Sanders KL, George S, Murphy JC. The status of Eurostus dussumierii and Hyp-
sirhina chinensis (Reptilia, Squamata, Serpentes): with comments on the origin of salt
tolerance in homalopsid snakes. Systematics and Biodiversity 10(4):479–489.
Kuntz RE. 1963. Snakes of Taiwan. United States Navy Medical Research Unit No. 2, Tai-
pei, Taiwan, 80 pp.
Kutsuma R, Sasai T & Kishida T. 2018. How snakes find prey underwater: Sea Snakes use
visual and chemical cues for foraging. Zoological Science 35(6):483–486.
Kuśmierek N, Pyrka E, Popiołek M. 2020. Diversity of helminths in polish reptiles: A re-
view. Biologia 75:733–739.
Kyi SW, Zug GR. 2003. Unusual Foraging Behavior of Naja kaouthia at the Moyingue Wet-
lands Birds Sanctuary, Myanmar. Hamadryad 27:265-266.
Kyle KJ, Alexander GJ, Du Preez LH. 2021. Habitat associations, reproduction, and geo-
graphical distribution of Lycodonomorphus obscuriventris (Serpentes: Lamprophiidae).
Herpetology Notes 14:865-7.
Lacépède B. G. E. de la Ville sur Illon (Compte), 1788-1789. Histoire générale et particu-
lière des Quadrupèdes ovipare set des Serpents, pour faire suite à l’Histoire naturelle
des animaux de Buffon. Imprimerie du Roi, Hôtel de Thou, Paris, 2 volumes., Vol.1
(1788): XVII + 1 + 651 + 1 pp.; Vol. 2 (1789): 9 + 20 + 144 +527 pp.
Lacépède BGH. 1800. Sur un nouveau genre de serpent. Bulletin de la Sociéte Philo-
654
Aquatic Snakes

mathique de Paris 2 (46):169.

Lacki M, Hummer J, Fitzgerald J. 2005. Population Patterns of Copperbelly Water
Snakes (Nerodia erythrogaster neglecta) in a Riparian Corridor Impacted by Mining
and Reclamation. The American Midland Naturalist, 153(2): 357-369.
Laidlaw FF. 1901. List of a collection of snakes, crocodiles, and chelonians from the Malay
Peninsula, made by members of the “Skeat Expedition.” 1899–1900. Proceedings of the
Zoological Society of London 2:575–583.
Lamb JY, Ljustina O. 2018. Nerodia fasciata confluens (Broad-banded Watersnake). Diet.
Herpetological Review 2018(4):757-758.
Lamonica RD, Abrahao-Charles H, Loguercio MF, Rocha-Barbosa O. 2007. Crecimiento,
Muda y Consumo de Alimento en Cautiverio de la Eunectes murinus (Linnaeus, 1758)
(Serpentes: Boidae). International Journal of Morphology 25(1):103-8.
Lane A, Gatus, J. 2010. Laticauda semifasciata. The IUCN Red List of Threatened Species
2010: e.T176721A7290432. [Link]
Lane A, Guinea M, Lobo A, Gatus J. 2010. Laticauda laticaudata. The IUCN Red List of
Threatened Species 2010: e.T176771A7301306. [Link]
[Link].
Lane A, Guinea M. 2010a. Laticauda crockeri. The IUCN Red List of Threatened Spe-
cies 2010: e.T11369A3273817. [Link]
[Link].
Lane A, Guinea M. 2010b. Laticauda schistorhynchus. The IUCN Red List of Threatened
Species 2010: e.T176740A7294814. [Link]
[Link].
Lane A, Guinea M. 2010c. Laticauda frontalis. The IUCN Red List of Threatened Spe-
cies 2010: e.T176736A7293312. [Link]
[Link].
Lane A, Guinea M. 2010d. Laticauda guineai. The IUCN Red List of Threatened Spe-
cies 2010: e.T176739A7294503. [Link]
[Link].
Lane A, Guinea M. 2010d. Laticauda guineai. The IUCN Red List of Threatened Spe-
cies 2010: e.T176739A7294503. [Link]
[Link].
Lane A, Shine R. 2011. Phylogenetic relationships within laticaudine sea snakes (Elapidae).
Molecular Phylogenetics and Evolution 59: 567–577. doi: 10.1016/[Link].2011.03.005
Lane A. Guinea M, Lobo A. Gatus J. 2010. Laticauda laticaudata. The IUCN Red List
of Threatened Species 2010:e.T176771A7301306. [Link]
[Link].
Lang R de, Vogel G. 2005. The snakes of Sulawesi, A field guide to the land snakes of Su-
lawesi with identification keys. Edition Chimaira, Frankfurt am Main, 312 pp.
Lang R de. 2013. The snakes of the Moluccas (Maluku), Indonesia. Edition Chimaira,
Frankfurt am Main, 417 pp.
Lang, R de 2017. The Snakes of Java, Bali, and Surrounding Islands. Edition Chimaira,
Frankfurt am Main, Germany, 435 pp.
Lanza B. 1983. Guide per il Riconoscimento delle Specie Animali nelle Acque Interne Ital-
iane. 27. Anfibi, Rettili (Amphibia, Reptilia. Roma: Consiglio Nazionale delle Richer-
655
Aquatic Snakes

che 196 pp.

Lanza B, Nistri A. 2005. Somali Boidae (genus Eryx Daudin 1803) and Pythonidae (genus
Python Daudin 1803)(Reptilia Serpentes). Tropical Zoology 8(1):[Link] (1956).
Larsen KW, Gregory PT. 1988. Amphibians and reptiles in the Northwest Territories. The
Natural History of Canada’s North: Current Research. Occasional Papers of the Prince of
Wales Northern Heritage Centre (Yellowknife) No. 1988 3:31-52.
Laurenti JN. 1768. Specimen medicum, exhibens synopin reptilium emendatam cum exper-
imentis circa venena et antidota reptilium Austriacorum. Trattner.
Law IS, Neo X, Chan DN, Shivaram R. 2020. Rediscovery of the Selangor mud snake, Raclit-
ia indica, in Singapore. Singapore Biodiversity Records 2020:165–167.
Lawson R. 1987. Molecular studies of thamnophiine snakes: 1. The phylogeny of the genus
Nerodia. Journal of Herpetology. 21:140-57.
Lee JC. 1996. The amphibians and reptiles of the Yucatán Peninsula. Comstock, Cornell
University Press, Ithaca, 500 pp.
Lee HJ, Lee JH, Park D. 2011. Habitat use and movement patterns of the viviparous aquat-
ic snake, Oocatochus rufodorsatus, from Northeast Asia. Zoological Science 28(8):593-
9.
Lee JC. 1996. The amphibians and reptiles of the Yucatán Peninsula. Comstock, Cornell
University Press, Ithaca, 500 pp.
Lee J, R. Calderón Mandujano, MA Lopez-Luna, 2013. Coniophanes quinquevittatus. IUCN
Red List of Threatened Species. 2013: e.T63755A3128807
Lee JL, Vogel G, Miller AH, Zug GR. 2018. Rediscovery of Xenochrophis bellulus (Stoliczka,
1871) from Myanmar (Serpentes: Natricinae) with comments on its taxonomic status.
Proceedings of the Biological Society of Washington 131: 19–35.
Leelawathanagoon P, Lheknim V, Marshall AG. 1997. Bio-ecology of the Khu Khut Bird
Sanctuary, lake Songkhla, southern Thailand I. An introduction to study sites and meth-
ods used. Warasan Songkhlanakharin.
Lema, T. 2002. Os Répteis do Rio Grande do Sul - atuais e fósseis, biogeografia, ofidismo.
Porto Alegre. EDIPUCRS. 264 pp.
Lenz N. 2012. Von Schmetterlingen und Donnerdrachen - Natur und Kultur in Bhutan.
Karlsruher Naturhefte 4, Naturkundemuseum Karlsruhe, 124 pp.
Lenz S, Schmidt A. 2011. Ergebnisse eines bundesweiten Projektes zur Forderung der
Wurfelnatter-Populationen und ihrer Lebensraume. Mertensiella 18: 30-39.
Lethaby M, Gray BS. 2016. Thamnophis brachystoma (Short-headed Gartersnake). Move-
ments and Site Fidelity. Herpetological Review 47(3): 485.
Leuenberger W, Davis AG, Mckenzie JM, Drayer AN, Price SJ. 2019. Evaluating Snake Den-
sity Using Passive Integrated Transponder (PIT) Telemetry and Spatial Capture–Recap-
ture Analyses for Linear Habitats. Journal of Herpetology 53(4):272-81.
Leviton AE. 1977. On the status of Natrix dubbiosi, Scortecci. Journal of the Saudi Arabian
Natural History Society 19:26–29.
Leviton AE, Brown RM, Siler CD. 2014. The dangerously venomous snakes of the Philippine
Archipelago. in The Coral Triangle: The 2011 Hearst Biodiversity Philippine Expedi-
tion (GC. Williams and TM. Gosliner, eds.). California Academy of Sciences, USA, Pp
473–530.
Leviton AE, Siler CD, Weinell JL, Brown RM. 2018. Synopsis of the Snakes of the Philip-
656
Aquatic Snakes

pines. Proceedings of the California Academy of Science 64 (14): 399-568.

Leviton AE, Wogan GOU, Koo MS, Zug GR, Lucas RS, Vindum JV 2003. The Dangerous-
ly Venomous Snakes of Myanmar Illustrated Checklist with Keys. Proceedings of the
California Academy of Science (24): 407–462.
Leviton AE. Guinevere OU. Wogan, Koo MS; Zug GR; S, Lucas RS, Vindum JV 2003. The
Dangerously Venomous Snakes of Myanmar Illustrated Checklist with Keys. Proc. Cal.
Acad. Sci. 54 (24): 407–462
Leviton, Alan E.; Guinevere O.U. Wogan; Michelle S. Koo; George R. Zug; Rhonda S. Lucas
and Jens V. Vindum (2003). The Dangerously Venomous Snakes of Myanmar Illustrated
Checklist with Keys. Proc. Cal. Acad. Sci. 54 (24): 407–462.
Lewin J. 1992. Parasites of water snake, Natrix natrix L., in Poland. Acta Parasitologica
37:195–199.
Lheknim V. 1998. The bio-ecology of the Khu Khut Bird Sanctuary, Lake Songkhla, south
Thailand II. Ichthyofauna and introduction to conservation and management. Song-
klanakarin Journal of Science and Technology 19(2):131-40.
Li M, Fry BG & Kini RM. 2005. Eggs-only diet: Its implications for the toxin profile changes
and ecology of the marbled sea snake (Aipysurus eydouxii). Journal of Molecular Evolu-
tion 60(1):81–89.
Lillywhite, H. B., 1991. The biology and conservation of Acrochordid snakes. Hamadryad,
16(1–2): 1–9.
Lillywhite HB. 1989. Unusual shedding behaviors in an aquatic snake, Acrochordus granula-
tus. Copeia 1989(3):768-70.
Lillywhite HB. 1996. Husbandry of the little file snake, Acrochordus granulatus. Zoo Biology
15(3):315–327.
Lillywhite HB. 2014. How snakes work: structure, function and behavior of the world’s
snakes. Oxford University Press.
Lillywhite HB & Ellis TM. 1994. Ecophysiological aspects of the coastal-estuarine distribu-
tion of acrochordid snakes. Estuaries 17(1):53–61.
Lillywhite HB, Albert JS, Sheehy CMI & Seymour RS. 2012. Gravity and the evolution of
cardiopulmonary morphology in snakes. Comparative Biochemistry and Physiology - A
Molecular and Integrative Physiology 161(2):230–242.
Lillywhite HB, Babonis LS, Sheehy CM & Tu MC. 2008. Sea snakes (Laticauda spp.) require
fresh drinking water: Implication for the distribution and persistence of populations.
Physiological and Biochemical Zoology 81(6):785–796.
Lillywhite HB, Brischoux F, Sheehy III CM, Pfaller JB. 2012. Dehydration and drinking
responses in a pelagic sea snake. Integrative and Comparative Biology 52(2): 227–234
Lillywhite HB, Heatwole H, Sheehy III CM. 2013. Dehydration and drinking behavior of
the marine file snake Acrochordus granulatus. Physiological and Biochemical Zoolo-
gy. 2013 Sep 30;87(1):46-55.
Lillywhite HB, Heatwole H, Sheehy III CM. 2015. Dehydration and drinking behav-
ior in true sea snakes (Elapidae: Hydrophiinae: Hydrophiini). Journal of Zoology
296(4):261-9.
Lillywhite HB, Menon JG, Menon GK, Sheehy CM, Tu MC. 2009. Water exchange and per-
meability properties of the skin in three species of amphibious sea snakes (Laticauda
spp.). Journal of Experimental Biology 212(12):1921–1929.
657
Aquatic Snakes

Lillywhite HB, Sheehy CM, Sandfoss MR, Crowe-Riddell J & Grech A. 2019. Drinking by
sea snakes from oceanic freshwater lenses at first rainfall ending seasonal drought. PLoS
ONE 14(2):1–11.
Lillywhite HB, Sheehy III CM, Brischoux F, Grech A. 2014. Pelagic sea snakes dehydrate
at sea. Proceedings of the Royal Society B: Biological Sciences. 281(1782):20140119.
Lillywhite HB, Sheehy III CM, Zaidan III F. 2008. Pitviper scavenging at the intertidal zone:
an evolutionary scenario for invasion of the sea. BioScience 58(10):947-55.
Lillywhite HB, Tu MC. 2011. Abundance of sea kraits correlates with precipitation. PLoS
ONE 6(12):12–15.
Lillywhite, Harvey B.; III, Coleman M. Sheehy; Brischoux, François; Pfaller, Joseph B.
(2015). On the Abundance of a Pelagic Sea Snake. Journal of Herpetology, 49(2), 184–
189. doi:10.1670/14-004
Lim BL. 1963. Comments on some rare snakes. Federated Museum Journal (Natural Histo-
ry) 8:60–64.
Lim BL. 1964. Notes on the elephant’s trunk snake and the puff-faced water snake in Kuala
Lumpur. Malayan Nature Journal 18:179–183.
Lim BL, 1991. Poisonous snakes of Peninsular Malaysia. Third edition. Malayan Nature So-
ciety and Institute for Medical Research, Kuala Lumpur, 74 pp
Lim BL, Kamarudin MS Bin. 1975. Notes on new locality records of some rare snakes in
Peninsular Malaysia. Malayan Nature Journal 29(1):23–27.
Lim BL, Ratnam L, Hussein NA. 1995. Snakes examined from the Sungai Singgor area of
Temengor, Hulu Perak, Malaysia. Malayan Nature Journal 48:357–364.
Limpus CJ. 1975. Coastal sea snakes of subtropical Queensland waters (23° to 28° South
Latitude). In: Dunson, W. A., ed. The Biology of Sea Snakes. Page(s) 173-182. Baltimore:
University Park Press.
Lind AJ, Welsh HH. 1990. Predation by Thamnophis couchii on Dicamptodon ensatus. Jour-
nal of Herpetology 24(1):104-6.
Lind AJ, Welsh Jr HH. 1994. Ontogenetic changes in foraging behaviour and habitat
use by the Oregon garter snake, Thamnophis atratus hydrophilus. Animal Behaviour,
48(6):1261-73.
Liner EA. 1992. Thamnophis exsul. Catalogue of American Amphibians and Reptiles 549:
1-2 Liner E A 1992.
Linnaeus C., 1758. Sistema Naturae per Regna tria Naturae, secondum Classes, Ordines,
Genera, Species cum Caracteribus, Differentiis, Sinonymis, Locis. Editio X riformata.
Tomus I. Impensis Direct. Laurentii Salvii, Holmiae (Stoccolma); (IV) + 823 + (1) pp.
Litvinov N, Bakiev A, Mebert K. 2011. Thermobiology and microclimate of the Dice snake
at its northern range limit in Russia. Mertensiella 18: 330-37.
Liu W, Cai B, Xia QR, Wu YQ. 2021. The complete mitochondrial genome of the Chinese
mountain keelback (Natricidae: Opisthotropis kuatunensis), comparative and phyloge-
netic analysis. Mitochondrial DNA Part B.6(12):3350-1.
Liu Y, Ding L, Lei J, Zhao E & Tang Y. 2012. Eye size variation reflects habitat and daily ac-
tivity patterns in colubrid snakes. Journal of Morphology 273(8):883–893.
Liu Y, Mebert K, Shi L. 2011. Notes on distribution and morphology of the Dice snake (Na-
trix tessellata) in China. Mertensiella 18:430.

658
Aquatic Snakes

Li-Wei Chung, Shih-Ping Chou, Si-Min Lin. 2016. Python molurus bivittaus (Burmese Py-
thon). Marine Dispersal. Herpetological Review 47(1)153.
Lizarro D, Powell RL, Eversole CB, Grerminger MA. Erythrolamprus poecilogyrus (Yel-
low-bellied Liophis). Diet. Herpetological Review 52:155-156.
Llorente GA, N Garriga, A Montori, A Richter-Boix, X Santos. 2005. Incidència de les car-
reteres sobre els amfibis i rèptils de Catalunya. Memoria inédita. Departament de Políti-
ca Territorial i Obres Públiques. Generalitat de Catalunya.
Lluch J, P Navarro. 1986. Rhabdias fuscovenosa (Railliet, 1899) Goodey, 1924, parásito pul-
monar de culebras de agua. Revista Ibérica de Parasitología 43:63-65.
Lobo A, Rasmussen A. 2010. Hydrophis mamillaris. The IUCN Red List of Threatened Spe-
cies 2010: e.T176724A7291020. [Link]
[Link].
López MS, Giraudo A. 2004. Diet of the large water snake Hydrodynastes gigas (Colubridae)
from northeast Argentina. Amphibia-Reptilia 25(2):178-84.
López MS, Giraudo AR. 2008. Ecology of the snake Philodryas patagoniensis (Serpentes,
Colubridae) from northeast Argentina. Journal of Herpetology 42(3):474-80.
López Seoane V. 1885. Identidad de Lacerta schreiberi (Bedriaga) y Lacerta viridis var.
gadovi (Boulenger) e investigaciones herpetológicas de Galicia, A Coruña: Impr. V.
Abad.
López-Hurtado Y, García-Padrón LY, González A, Díaz LM, Rodríguez-Cabrera TM. 2020.
Notes on the feeding habits of the Caribbean Watersnake, Tretanorhinus variabilis (Dip-
sadidae). Reptiles & Amphibians. 17;27(2):147-53.
Loughman ZJ, Meyers G, Glon MG, Adams SB. 2020. Regina rigida sinicola (= Lyodytes
rigida sinicola) (Gulf Glossy Snake Crayfish Snake). Diet and Habitat. Herpetological
Review 51(4):877-878
Loveridge A. 1931. On two amphibious snakes of the Central African Lake Region. Anti-
venin Institute of America; 114:7-15.
Loveridge AR. 1955. On a second collection of reptiles and amphibians taken in Tangan-
yika Territory by CJP Ionides, Esq. Journal of the East African Natural History Society
22:168-98.
Loveridge, A. 1958. Revision of five African snake genera. Bulletin of the Museum of Com-
parative Zoology 119:1-198.
Ludeking EA. 1867. Natuur en geneeskundige topographie van Agam (Westkust van Suma-
tra). Martinus Nijhoff, ’s-Gravenhage, 162 pp.
Luiselli L. 1996. Individual success in mating balls of the grass snake, Natrix natrix: size is
important. Journal of Zoology 239:731–740.
Luiselli L, Akani GC. 2002. Is thermoregulation really unimportant for tropical reptiles?
Comparative study of four sympatric snake species from Africa. Acta Oecologica
23(2):59-68.
Luiselli L, Akani GC, Angelici FM, Politano E, Ude L, Wariboko SM. 2003. Diet of the semi‐
aquatic snake, Afronatrix anoscopus (Colubridae) in southern Nigeria. African Journal
of Herpetology, 2003 52(2): 123-126.
Luiselli L, Akani GC, Angelici FM, Ude L, Wariboko SM. 2005. Seasonal variation in hab-
itat use in sympatric Afrotropical semi-aquatic snakes, Grayia smythii and Afronatrix
anoscopus (Colubridae). Amphibia Reptilia 26(3):372.
659
Aquatic Snakes

Luiselli L., Capula,M., & Shine R., 1997. Food habits, growth rates, and reproductive biol-
ogy of grass snakes, Natrix natrix (Colubridae) in the Italian Alps. Journal of Zoology,
241(2): 371-380.
Luiselli L, Capizzi D, Filippi E, Anibaldi C, Rugiero L, M. 2007. Comparative diets of three
populations of an aquatic snake (Natrix tessellata, Colubridae) from Mediterranean
streams with different hydric regimes. Copeia 2007(2):426–35.
Luiselli L, Madsen T, Capizzi D, Rugiero L, Pacini N, Capula M. 2011. Long‐term popula-
tion dynamics in a Mediterranean aquatic snake. Ecological research 26(4):745-53.
Luiselli L, Rugiero L., 1991. Food niche partitioning by water snakes (genus Natrix) at a
freshwater environment in central Italy. Journal of Freshwater Ecology 6: 439–444.
Luiselli L., Filippi E., Capula M., 2005: Geographic variation in diet composition of the
grass snake (Natrix natrix) along the mainland and an island of Italy: the effects of hab-
itat type and interference with potential competitors. The Herpetological Journal 15:
221–230.
Lukoschek V, Beger M, Ceccarelli D, Richards Z, Pratchett M. 2013. Enigmatic declines of
Australia’s sea snakes from a biodiversity hotspot. Biological Conservation. 166:191-202.
Lukoschek V, Guinea M, Milton D. 2010. Ephalophis greyae. The IUCN Red List of Threat-
ened Species 2010: e.T176755A7298070. [Link] [Link].2010-4.
[Link].
Lukoschek V., Osterhage JL, Karns DR, Murphy JC, Voris HK. 2011. Phylogeography of the
Mekong mud snake (Enhydris subtaeniata): the biogeographic importance of dynamic
river drainages and fluctuating sea levels for semiaquatic taxa in Indochina. Ecology and
Evolution 1(3):330–342.
Lukoschek, V. & Guinea, M. 2010. Aipysurus foliosquama. The IUCN Red List of Threat-
ened Species 2010: e.T176714A7288783. [Link] [Link].2010-4.
[Link].
Lukoschek, V., Courtney, T., Milton, D. & Guinea, M. 2010e. Aipysurus laevis. The IUCN
Red List of Threatened Species 2010: e.T176704A7286736. [Link]
[Link].
Lukoschek, V., Guinea, M., & Rasmussen, A. 2010d. Aipysurus fuscus. The IUCN Red List
of Threatened Species 2010: e.T176763A7299535. [Link] IUCN.
[Link].
Lukoschek, V., Rasmussen, A.R., Sanders, K., Lobo, A.S. & Courtney, T. 2010. Hydrophis
peronii. The IUCN Red List of Threatened Species 2010: e.T176756A7298218. https://
[Link]/10.2305/[Link]. Accessed on 13 May
2022.
Lyle JM, GJ Timms 1987. Predation on Aquatic Snakes by Sharks from Northern Austra-
lia. Copeia 1987, (3):802-03. doi:10.2307/1445681.
Macartney JM, Larsen KW, Gregory PT. 1989 Body temperatures and movements of hiber-
nating snakes (Crotalus and Thamnophis) and thermal gradients of natural hibernacula.
Canadian Journal of Zoology 67(1):108-14.
MacNae W. 1968. A general account of the fauna and flora of mangrove swamps and forests
in the Indo-West-Pacific region. Advances in Marine Biology 6:73–270.
Macrì S, Savriama Y, Khan I & Di-Poï N. 2019. Comparative analysis of squamate brains
unveils multi-level variation in cerebellar architecture associated with locomotor spe-
660
Aquatic Snakes

cialization. Nature Communications 10(1):5560.

Madsen T. 1983. Growth rates, maturation and sexual size dimorphism in a population of
Grass Snakes, Natrix natrix, in southern Sweden. Oikos 40: 277–282.
Madsen T. 1984. Movements, home range size and habitat use of radio–tracked grass snakes
(Natrix natrix) in southern Sweden. Copeia 1984:707–713.
Madsen T. 1997. Are juvenile grass snakes, Natrix natrix, aposematically coloured? Oikos
48:265–267.
Madsen T, Osterkamp M. 1982. Notes on the biology of the fish-eating snake Lycodonomor-
phus bicolor in Lake Tanganyika. Journal of Herpetology 16(2):185-8.
Madsen T, Shine R. 2000a. Rain, fish and snakes: climatically driven population dynamics of
Arafura filesnakes in tropical Australia. Oecologia. 124(2):208-15.
Madsen T, Shine R. 2000b. Energy versus risk: costs of reproduction in free‐ranging py-
thons in tropical Australia. Austral Ecology. 25(6):670-5
Madsen T, Ujvari B, Shine R, Olsson M. 2006. Rain, rats and pythons: climate driven popu-
lation dynamics of predators and prey in tropical Australia. Austral Ecology 31:30–37.
Mahendra BC. 1938. Some remarks on the phylogeny of the Ophidia. Anatomischer An-
zeiger 86: 347– 356.
Maia DC, Silva AG, Travaglia-Cardoso SR. 2015. Helicops modestus (Ophidia: Dipsadidae):
reproduction. Acta zoológica Mexicana 31(2):333-4.
Malkmus R. 1996. Die “gestreifte” form der Vipernatter (Natrix maura) aus Portugal. Sala-
mandra 32:285-286.
Malkmus R. 1997. Slangen van Portugal (9): de Spaanse Ringslang (Natrix natrix astrepto-
phora). Lacerta 56 (1):21–23.
Malkmus R. 2008. Winteraktive Schlangen in Portugal (Erganzende Bemerkungen.
Zeitschrift fur Feldherpetologie 15(1):97-98.
Malkmus R, Brühl C. 2002. Amphibians & Reptiles of Mount Kinabalu (North Borneo).
Ruggell: ARG Gantner.
Malkmus R, Sauer H. 2013. Totstellen bei Iberischen Ringelnattern. Terraria/Elaphe 6:72–
73.
Malnate EV. 1953. The taxonomic status of the Tibetan colubrid snake Natrix baileyi. Co-
peia, 1953(2):92-96.
Malnate EV. 1960. Systematic division and evolution of the colubrid snake genus Natrix,
with comments on the subfamily Natricinae. Proceedings of the Academy of Natural
Sciences of Philadelphia 112:41-71.
Malnate EV. 1966. Amphiesma platyceps (Blyth), and Amphiesma sieboldii (Guenther): sib-
ling species (Reptilia: Serpentes). Journal of Bombay Natural History Society 63:1-7.
Malnate EV, Underwood G. 1988. Australasian Natricine snakes of the genus Tropidonophis.
Proceedings of the Academy of Natural Sciences of Philadelphia 140(1): 59-201.
Malnate EV, Underwood G. 1988. Australasian natricine snakes of the genus Tropidonophis.
Proceedings of the Academy of Natural Sciences of Philadelphia. 1988:59-201.
Manjarrez J, Drummond H. 1996. Temperature‐limited activity in the garter snake Tham-
nophis melanogaster (Colubridae). Ethology 102(1):146-56.
Mamonekene V, Lavoué S, Pauwels OSG, Mve Beh JH, Mackayah J-E, Tchignoumba L.
2006. Fish diversity at Rabi andGamba, Ogooué-Maritime Province, Gabon. Pp. 285-
296. In: A. Alonso, M. E. Lee, P. Campbell, O. S. G. Pauwels and F. Dallmeier,editors,
661
Aquatic Snakes

Gamba, Gabon: Biodiversité d’une forêt équatoriale africaine / Gamba, Gabon: Biodiver-
sity of an equatorial African [Link] of the Biological Society of Washington
(12).
Manthey U, Grossmann W. 1997. Amphibien & Reptilien Südostasiens. Natur und Tier
Verlag (Münster), 512 pp.
Manuilova O, Pisatets’ AM. 2008. Preliminary data on karyology of the water snake, Natrix
tessellata, from Ukraine. Studies of the Ukrainian Herpetological Society 1:39–42.
Mao SH. 1970. Food of the common venomous snakes of Taiwan. Herpetologica 26(1):45–
48.
Mao J-J. 1998. Phylogenetic relationships of natricinae snakes in Taiwan. Master Thesis, Na-
tional Sun Yat-sen University, Kaohsiung City, Taiwan, 74 pp. (In Chinese)
Mao SH, Chen BY. 1980. The Sea Snakes of Taiwan. NSC Special Publications, No. 4. Na-
tional Science Council, Taipei.
Mao J-J, Chen W-H, Ger R, Hsieh T-H, Liu L-X. 2012. Ophidian fauna and species richness
along an elevational gradient in Yilan. Ilan University Journal of Bioresources 8(2):39-
49. (In Chinese).
Mao J-J, Norval G, Kung W-B, Chang H-N. 2006. Bungarus multicinctus multicinctus diet.
Herpetological Review 37 (3): 350.
Mao J-J, Norval G, Hsu C-L, Chen W-H, Ger R. 2010. Observations and comments on the
diet of the many-banded krait (Bungarus multicinctus multicinctus) in Taiwan. IRCF
Reptiles & Amphibians 17(2):73-76.
Mao J-J, Norval. G. 2013. Does rotavator use in Taiwanese rice paddies expose reptiles to
increased opportunistic avian predation? IRCF Reptiles & Amphibians 20(2):41-43.
Mario-Da-Rosa C, Abegg AD, Malta-Borges L, Righi AF, Bernarde PS, Cechin SZ , Dos
Santos TG. 2020. A fisherman’s tale: Activity, habitat use and the first evidence of lingual
lure behavior in a South American snake. Salamandra 56(February):39–47
Marques OAV 1996. Sordellina punctata. Diet. Herpetological Review 27:147–147.
Marques OA, Sazima I. 2008. Winding to and fro: constriction in the snake Anilius scytale.
Herpetological Bulletin. 2008:103.
Marques OAV, Souza VC. 1993. Nota sobre a atividade alimentar de Liophis miliaris no am

biente marinho (Serpentes, Colubridae). Revista Brasileira de Biologia 53, 645–648.

Marsh H, Corkeron PJ, Limpus, CJ, Shaughnessy PD, Ward TM. 1993. Conserving marine
mammals and reptiles in Australia and Oceania. In: C. Moritz and J. Kikkawa (eds),
Conservation Biology in Australia and Oceania, pp. 225-244. Surrey, Beatty & Sons,
Chipping Norton.
Martens H. 1996. The rediscovery of the grass snake Natrix natrix (L.) in the Levant. Zoolo-
gy in the Middle East 12:59–64.
Martínez A, Nguyen D, Basson M, Medina J, Irschick D, Baeckens S. 2021. Quantifying sur-
face topography of biological systems from 3D scans. Methods in Ecology and Evolution
2021(October 2020):1–12.
Martínez del Marmol G, Harris DJ, Geniez P, de Pous P, Salvi D. 2019. Amphibians and
Reptiles of Morocco. Ed. Chimaira. Pp. 478.
Martínez-Freiría F, Argaz H, Fahd S, Brito JC. 2013. Climate change is predicted to nega-
tively
662
Aquatic Snakes

influence Moroccan endemic reptile richness. Implications for conservation in protected

areas. Naturwissenschaften 100(9):877–889.
Martinez J, Behler JL. 1988. Erpeton tentaculatum (Tentacled Snake). Reproduction. Herpe-
tological Review 19:35.
Martins M, Oliveira ME. 1998. Natural history of snakes in forests of the Manaus region,
Central Amazonia, Brazil. Herpetological Natural History 6(2):78-150.
Martorell E. 1990. Natrix maura: ¿Posibilidad de aprendizaje? Boletin de la Asociación Her-
petológica Española 1: 24.
Marx H. 1958. Catalogue of type specimens of reptiles and amphibians in Chicago Natural
History Museum. Fieldiana, Zoology 36(4):409-496.
Maschio GF, Prudente AL, Rodrigues FD, Hoogmoed MS. 2010. Food habits of Anilius scy-
tale (Serpentes: Aniliidae) in the Brazilian Amazonia. Zoologia (Curitiba) 27(2):184-90.
Maslin TP. 1959. An annotated check list of the amphibians and reptiles of Colorado. Univ.
Colorado Studies, Ser. Bio. 6:vi + 98 p.
Maslin TP. Beidleman RG, Lowe CH, 1958. The status of the lizard Cnemidophorus perplex-
us Baird and Girard (Teiidae). Proc. United States Nat. Mus. 108:331-345.
Masseti M. 2014. Herpetological enigmas from the Arabian seas, with particular reference
to the Sarso island racer, Platyceps insularis Mertens 1965 (Frasan archipelago, Saudi
Arabia. Scripts Herpetologica. Studies on Amphibians and Reptiles in honour of Bene-
detto Lanza, Pages 99-116.
Masunaga G, Nagai Y, Tanase H, and Ota H. 2005. A record of the black-headed sea snake,
Hydrophis melanocephalus (Reptilia: Elapidae), from Wakayama Prefecture, Japan. Cur-
rent herpetology, 24(1):37-41.
Mateo Miras JA, M Cheylan, MS Nouira, U Joger, P Sá-Sousa, V Pérez-Mellado, B Schmidt,
A Meyer, R Sindaco, A Romano, I Martínez-Solano. 2009. Natrix maura. En: IUCN Red
List of Threatened Species. Version 2009.2. <[Link]>.
Maumary L, Duperrex H, Cloutier J, Vallotton A. 2013. Première nidification du Circaèt
Jean-le-Blanc (Circaetus gallicus) en Suisse - Observations sur la biologie de reproduction,
en particulier le régime alimentaire. Nos Oiseaux 60:3–24.
Mazza G, Monney JC, Ursenbacher S. 2011. Structural habitat partitioning of Natrix tessel-
lata and Natrix maura at Lake Geneva, Switzerland. Mertensiella 18: 80-86

McAllister CT. 1985. Nerodia rhombifera. Catalogue of American Amphibians and Reptiles
(CAAR). 1985.
McAllister CT. 2021. Nerodia erythrogaster (Plain-bellied Watersnake). Diet. Herpetological
Review 52(2)427.
McCosker, J.E. 1975. Feeding behaviour of Indo-Australian Hydrophiidae. In: Dunson, W.
A., ed. The Biology of Sea Snakes. Page(s) 217-232. Baltimore: University Park Press.
McCoy CJ. 1969. Snakes of the genus Coniophanes (Colubridae) from the Yucatan Peninsu-
la, México. Copeia 1969 (4): 847-849.
McCoy CJ. 1984. Ecological and zoogeographic relationships of amphibians and reptiles
of the Cuatro Ciénegas Basin. Journal of the Arizona-Nevada Academy of Science
1:49-59.
McCranie JR. 1983. Nerodia taxispilota. Catalogue of American Amphibians and Reptiles
(CAAR). 1983.
663
Aquatic Snakes

McCranie JR. 2011. The snakes of Honduras. SSAR, Salt Lake City, 725 pp.
McCranie JR. 2015. A checklist of the amphibians and reptiles of Honduras, with addi-
tions, comments on taxonomy, some recent taxonomic decisions, and areas of further
studies needed. Zootaxa 3931 (3): 352–386.
McCranie JR; McAllister CT 1988. Nerodia valida (Kennicott). Mexican west coast water
snake. Catalogue of American Amphibians and Reptiles 431: 1-3.
McDowell SB. 1979. A catalogue of the snakes of New Guinea and the Solomons, with
special reference to those in the Bernice P. Bishop Museum. Part III. Boinae and Acro-
chordoidea (Reptilia, Serpentes). Journal of Herpetology. 26:1-92.
McDowell SB. 1987. Systematics in Snakes. In Seigel, R. A.; Collina, J. T.; Novak, S. S.
(eds.). Systematics in Snakes: Ecology and Evolutionary Biology. New York: Macmil-
lan Publishing Company.
McVay JD, Flores-Villela O, Carstens B. 2015. Diversification of North American natricine
snakes. Biological Journal of the Linnean Society 116(1):1-2.
Mead JI, Steadman DW. 2017. Late Pleistocene snakes (Squamata: Serpentes) from Abaco,
The Bahamas. Geobios. 50(5-6):431-40.
Mebert K. 1993. Untersuchungen zur Morphologie und Taxonomie der Würfelnatter
Natrix tessellata (Laurenti 1768) in der Schweiz und im südlichen Alpenraum. – M.S.
thesis, Zoological Museum, University of Zürich, Switzerland.
Mebert K. 2007. The Dice Snake at Lake Brienz, in Annual Volume 2007, Shore-Conser-
vation of the Lakes Brienz and Thun. UTB Publishing, Thun, Switzerland:169-180 (in
German)
Mebert K . 2004. Summary on Hybridization between Nerodia sipedon and N. fasciata. Pp.
104-108 in Gibbons JW, Dorcas ME, North American Watersnakes: a Natural History,
University of Oklahoma Press..
Mebert K. 2005. Extensive hybridization between the two taxa Nerodia sipedon and N. fas-
ciata in North Carolina. NC Herps 28(2):13-16.
Mebert K. 2010. Massive Hybridization and Species Concepts, Insights from Watersnakes.
– VDM Verlag, Saarbrücken, Germany.
Mebert K. 2011a. The Dice snake, Natrix tessellata: Biology, Distribution and Conservation
of a Palaearctic species. Deutsche Gesellschaft für Herpetologie und Terrarienkunde
(DGHT) e. V., Rheinbach Production: Andreas Mendt, DGHT-office. ISSN 0934-6643
ISBN 978-3-9812565-4.
Mebert K. (ed.) 2011a. The Dice Snake, Natrix tessellata: Biology, Distribution and Conser-
vation of a Palaearctic Species. Mertensiella 18.
Mebert K. 2011b. Geographic variation of morphological characters in the Dice snake
(Natrix tessellata). Mertensiella 18: 11-20.
Mebert K. 2011c. Sexual dimorphism in the Dice snake (Natrix tessellata) from the Central
Alps. Mertensiella 18: 94-100.
Mebert K. 2011d. Introduced and indigenous populations of the Dice snake (Natrix tessel-
lata) in the Central Alps – microgeographic variation and effect of inbreeding. Merten-
siella 18: 71-80.
Mebert K. 2011e. Terrestrial Dice snakes: how far from water a semiaquatic snakes ven-
tures out? Mertensiella 18: 453-55.
Mebert K. 2011f. Unsuitable food for Dice snakes (Natrix tessellata)! Mertensiella 18: 448-

664
Aquatic Snakes

49.
Mebert K, B Trapp, A Dall’Asta, P Velensky, W Böhme. 2011. Hybrids between Natrix tessel-
lata and N. natrix/maura. Mertensiella 18:154-156.
Mebert K, Conelli AE, Nembrini M, Schmidt BR. 2011a. Monitoring and assessment of the
distribution of the Dice snake in Ticino, southern Switzerland. Mertensiella 18:117-31.
Mebert K, Masroor R, Chaudhry MJI. 2013. The Dice snake, Natrix tessellata (Serpentes:
Colubridae) in Pakistan: analysis of its range limited to few valleys in the Western Kara-
koram. Pakistan Journal of Zoology 45(2):395-410.
Mebert K, Masroor R. 2013. Dice snakes in the western Himalayas: discussion of potential
expansion routes of Natrix tessellata after its rediscovery in Pakistan. Salamandra 49(4):
229-33.
Mebert K, Ott T. 2011. Mating aggregations in Natrix tessellata. Mertensiella 18:437-39.
Mebert K, Polzer W. 2011. Fatal hunting accidents: killing of Dice snakes (Natrix tessellata)
by Bullheads (Cottus gobio). Mertensiella 18: 145-47.
Mebert K, Trapp B, Dall’Asta A, Velensky P, Bohme W. 2011. Hybrids between Natrix tessel-
lata and N. natrix/maura. – Mertensiella 18:154-57.
Mecham JS. 1983. Nerodia harteri. Catalogue of American Amphibians and Reptiles (CAAR).
1983.
Meek R. 2009. Patterns of reptile road-kills in the Vendee region of western France. Herpe-
tological Journal 19:135-142.
Meijide MW, R Salas. 1989. Observaciones sobre el comportamiento depredativo de algunos
colúbridos ibéricos en estado salvaje. Doñana, Acta Vertbrata 16:329-332.
Meijide MW. 1989. Observaciones sobre el comportamiento depredador de algunos colúbri-
dos ibéricos en estado salvaje. Doñana, Acta Vertebrata 16(2):329–332.
Meijide MW, Pérez-Melero J. 1994. Nuevos casos de melanismo en Coronella austriaca y
Natrix natrix (Ophidia, Colubridae) en el norte de Iberia. Boletín de la Asociación Her-
petológica Española 5:33–36.
Meister B, Ursenbacher S, Baur B. 2012. Grass snake population differentiation over differ-
ent geographic scales. Herpetologica 68:134–145.
Mell R. 1922. Beiträge zur fauna sinica. Die Vertebraten Südchinas: feldlisten und feldnoten
der säuger. vögel, reptilien, Batrachier. Archiv für Naturgeschichte 88A:122.
Mell R. 1928. Beobachtungen über das Sinnesleben chinesis cher Reptilien, insbesondere
Schlangen. Zeitschrift für Morphologie und Ökologie der Tiere (Zeitschrift Wissen-
schaftlich Biologie A) 11(5):539–569.
Mertens R. 1969. Zur Synonymie und Variabilität der Würfelnatter (Natrix tessellata).
Senckenbergiana Biologica 50: 125–131.
Mertens D. 1994. Some aspects of thermoregulation and activity in free-ranging grass
snakes (Natrix natrix L.). Amphibia-Reptilia 15:322–326.
Mertens D. 1995. Population Structure and Abundance of Grass Snakes, Natrix natrix, in
Central Germany. Journal of Herpetology 29(3):454–456.
Mertens D. 2008. Untersuchungen zur Ökologie der Ringelnatter - Ergebnisse einer radiote-
lemetrischen Freilandstudie. Mertensiella 17:151–161.
Mertens R. 1930. Die Amphibien und Reptilien der Inseln Bali, Lombok, Sumbawa und
Flores (Beiträge zur Fauna der Kleinen Sunda Inseln, I). Frankfurt am Main: Abhand-
lungen der Senckenbergischen Naturforschenden Gesellschaft 42(3):115-344.
665
Aquatic Snakes

Mertens R. 1959. Eine neue Wassernatter aus West-Pakistan. Senckenbergiana Biologica

40:117–120.
Mertens R. 1969. Zur Synonymie und Variabilität der Würfelnatter (Natrix tessellata).
Senckenbergiana Biologica 50: 125-31.
Messenger KR, Wang Y. 2015. Notes on the natural history and morphology of the Ning-
shan Lined Snake (Stichophanes ningshaanensis Yuen, 1983; Ophidia: Colubridae) and
its distribution in the Shennongjia National Nature Reserve, China. Amphibian & Rep-
tile Conservation 9:(2):111-9.
Metzger C, Christe P, Ursenbacher S. 2011. Diet variability of two convergent natricine
colubrids in an invasive-native interaction. Mertensiella 18:86-93.
Metzger C, Ursenbacher S, Christe P. 2009. Testing the competitive exclusion principle us-
ing various niche parameters in a native (Natrix maura) and an introduced (N. tessella-
ta) colubrid. Amphibia-Reptilia 30:523-531.
Meyer AB. 1874. Eine Mittheilung von Hern. Dr. Adolf Berhard Meyer über die von ihm
auf Neu-Guinea und den Inseln Jobi, Mysore und Mafoor im Jahre 1873 gesammelten
Amphibien. Pp. 128–140. In: Monatsberichte der Königlichen Preussische Akademie
des Wissenschaften zu, Königliche Akademie der Wissenschaften, Berlin.
Meyer AB. 1874. Eine Mittheilung von Hern. Dr. Adolf Berhard Meyer über die von ihm
auf Neu-Guinea und den Inseln Jobi, Mysore und Mafoor im Jahre 1873 gesammelten
Amphibien. Pp. 128–140. In: Monatsberichte der Königlichen Preussische Akademie
des Wissenschaften zu, Königliche Akademie der Wissenschaften, Berlin.
Michaud EJ, Dixon JR. 1989. Prey items of 20 species of the Colubridae snake genus Lio-
phis. Herpetological Review 20(2): 39–41.
Michniewicz R, Aubret F. 2010. Warming up for cold water: influence of habitat type on
thermoregulatory tactics in a semi-aquatic snake. Amphibia-Reptilia 31(4):525-31.
Michniewicz R, Aubret F. 2010. Warming up for cold water: influence of habitat type on
thermoregulatory tactics in a semi-aquatic snake. Amphibia-Reptilia 31(4):525-31.
Mihalca AD. 2011. Parasitism in the Dice snake (Natrix tessellata) – a literature review.
Mertensiella 18:255-72.
Milko DA, Trotchenko NV, Knyazeva VS. 2021. The first record of grass snake Natrix
natrix (Colubridae) in Kyrgyzstan. [in Russian] Исследование живой природы
Кыргызстана [Wildlife Research in Kyrgyzstan] (1):32–34.
Miller AH, Davis HR, Luong AM, Do QH, Pham CT, Ziegler T, Lee JL, De Queiroz K,
Reynolds RG, Nguyen TQ. 2020. Discovery of a New Species of Enigmatic Odd-Scaled
Snake (Serpentes: Xenodermidae: Achalinus) from Ha Giang Province, Vietnam. Co-
peia 108(4):796-808.
Miller DE, Mushinsky HR. 1990. Foraging ecology and prey size in the mangrove water
snake, Nerodia fasciata compressicauda. Copeia 1990 (4):1099-106.
Mills MS. 2002. Ecology and life history of the brown water snake (Nerodia taxispilo-
ta) (Doctoral dissertation, University of Georgia).
Milton D, Courtney T, Guinea M, Lukoschek V, Fletcher E. 2010b. Hydrophis kingii. The
IUCN Red List of Threatened Species 2010: e.T176761A7299189. [Link]
org/10.2305/ [Link].
Milton D, Courtney T, Guinea M, Lukoschek V. 2010a. Hydrophis elegans. The IUCN Red
List of Threatened Species 2010: e.T176734A7292879. [Link]
666
Aquatic Snakes

[Link].
Milton DA, Fry GC, Dell Q 2009. Reducing impacts of trawling on protected sea snakes:
by-catch reduction devices improve escapement and survival. Marine and Freshwater
Research 60, 824-832.
Milton DA, Zhou S, Fry G, Dell Q. 2008. Risk assessment and mitigation for sea snakes
caught in the Northern Prawn Fishery, Queensland Government, Queensland Depart-
ment of Primary Industries and Fisheries, Fisheries Research and Development Cor-
poration (FRDC) Final Report 2005/051, Brisbane
Milton DA. 2001. Assessing the susceptibility to fishing of populations of rare trawl by-
catch: sea snakes caught by Australia´s Northern Prawn Fishery. Biological Conserva-
tion. 281-290.
Milton DA. 2001. Assessing the susceptibility to fishing of populations of rare trawl by-
catch: sea snakes caught by Australia’s Northern Prawn Fishery. Biological Conserva-
tion 101:281-290.
Milton DA. 2001. Assessing the susceptibility to fishing of populations of rare trawl by-
catch: sea snakes caught by Australia’s Northern Prawn Fishery. Biological Conserva-
tion 101:281-290.
Milton, D., Courtney, T., Guinea, M. Lukoschek, V. 2010. Hydrophis pacificus. The IUCN
Red List of Threatened Species 2010: e.T176712A7288420. [Link]
[Link]. Accessed on 20 April 2022.
Minton SA, Heatwole H. 1975. Sea snakes from three reefs of the Sahul Shelf. Pp. 141–144
In The Biology of Sea Snakes. Dunson, W.A. (Ed). University Park Press, Baltimore,
Maryland, USA
Minton SA. 1966a. A contribution to the herpetology of West Pakistan. Bulletin of the
American Museum of Natural History 134:27–184.
Minton SA. 1966b. A contribution to the herpetology of West Pakistan. Bulletin of the
American Museum of Natural History 134:27–184.
Minton SA. 1980. Thamnophis butleri. Catalogue of American Amphibians and Reptiles
(CAAR).
Minton SA. 1983. Lethal toxicity of venoms of snakes from the Coral Sea 21(6), 901– 902.
doi:10.1016/0041-0101(83)90082-x
Miralles A, Marin J, Markus D, Herrel A, Hedges SB, Vidal N. 2018. Molecular evidence
for the paraphyly of Scolecophidia and its evolutionary implications. Journal of evolu-
tionary biology 31(12):1782-93.
Miranda BH, Fernandes DS. 2012. Morphological variation of Sordellina punctata (Peters,
1880) with notes on the phylogenetic position of the genus within the family Dipsadi-
dae (Serpentes). International Journal of Zoology. 2012.
Mitchell JC. 1982. Farancia erytrogramma. Catalogue of American Amphibians and Rep-
tiles (CAAR).
Mitchell JC. 1994. The Reptiles of Virginia. Smithsonian Institution Press, Washington,
D.C.
Mittleman MB, Goris RC. 1974. Envenomation from the bite of the Japanese colubrid
snake Rhabdophis tigrinus (Boie). Herpetologica. 30:113-9.
Mittleman MB, Goris RC. 1978. Death caused by the bite of the Japanese colubrid snake
Rhabdophis tigrinus (Boie) (Reptilia, Serpentes, Colubridae). Journal of Herpetology.
667
Aquatic Snakes

12:109-11.
Mociño-Deloya E, Setser K, Reguera S, Feriche M, Pleguezuelos JM, D 2009b. Thamnophis
scaliger (Mesa Central Blotched Garter Snake). Maximum Elevation. Herpetological
Review 40(4):443.
Modahl CM, Mukherjee AK, Mackessy SP. 2016. An analysis of venom ontogeny and
prey-specific toxicity in the Monocled Cobra (Naja kaouthia). Toxicon 119:8-20.
Moen DS, Winne CT, Reed RN. 2005. Habitat-mediated shifts and plasticity in the evap-
orative water loss rates of two congeneric pit vipers (Squamata, Viperidae, Agkistro-
don). Evolutionary Ecology Research 7(5):759-66.
Moon BR, Penning DA, Segall M, Herrel A. 2019. Feeding in Snakes: Form, Function,
and Evolution of the Feeding System. Pages 527-574 in: Feeding in Vertebrates.
Springer, Cham.
Moore MK, Townsend Jr VR. 1998. The interaction of temperature, dissolved oxygen
and predation pressure in an aquatic predator-prey system. Oikos 1:329-36.
Moraes-DA-Silva A, Amaro RC, Nunes PM, Rodrigues MT, Curcio FF. 2021. Long known,
brand new, and possibly threatened: a new species of watersnake of the genus Helicops
Wagler, 1828 (Serpentes; Xenodontinae) from the Tocantins-Araguaia River Basin,
Brazil. Zootaxa. 4903(2):zootaxa-4903.
Mori A. 1998. Prey-handling behavior of three species of homalopsine snakes: features
associated with piscivory and Duvernoy’s glands. Journal of Herpetology 32(1):40–
50.
Morice A. 1875a. Coup d’oeil sur la faune de la Cochinchine française. H. Georg, Lyon,
101 pp.
Morice A. 1875b. Note sur l’Herpeton tentaculatum. Annales des Sciences Naturelles 6th
Series 2(5):1–11.
Morice A. 1875c. Sur les habitudes d’un remarquable serpent de la Cochinchine: l’Herpe-
ton tentaculatum. Comptes Rendus Hebdomadaires Des Seances De L’Acadamie Des
Science 80:128–129.
Muansanga L, Lalremsanga HT. 2020. Feeding behavior of the Mizo Rainsnake, Smithophis
atemporalis (Squamata: Natricidae), in Mizoram, India. IRCF Reptiles & Amphibians
27 (3): 487-488.
Muchtar M, Ilahuge AG. 2001. Mamberamo River Estuary, Irian Jaya. LOICZ Biogeo-
chemical Modeling Node.
Muchtar M, Ilahuge AG. 2001. Mamberamo River Estuary, Irian Jaya. LOICZ Biogeo-
chemical Modeling Node.
Mullin SJ, Mushinsky HR. 1995. Foraging ecology of the mangrove salt marsh snake, Nero-
dia clarkii compressicauda: effects of vegetational density. Amphibia-Reptilia. 1995 Jan
1;16(2):167-75.
Müllner A, Eduard Linsenmair K. 2007. Nesting behavior and breeding success of Hoa-
tzins. Journal of Field Ornithology 78(4):352-61.
Muñoz A, Felicísimo ÁM, Santos X. 2021. Analysing how pre-fire habitat legacy and post-
fire management influence the resilience of reptiles to fire. Forests 12(11):1487.
Murphy J, Cox J, Voris HK 1999. A key to the sea snakes in the Gulf of Thailand. Natural
History Bulletin of the Siam Society. 47:.
Murphy JC, Chan-ard T, Mekchai S, Cota M, Voris HK. 2008. The rediscovery of Angel’s
668
Aquatic Snakes

stream snake, Paratapinophis praemaxillaris Angel, 1929 (Reptilia: Serpentes: Natri-

cidae). Tropical Natural History 8(2):169-83.
Murphy JC, Henderson RW. 1997. Tales of giant snakes: a historical natural history of ana-
condas and pythons. USA: Krieger Publishing Company.
Murphy JC, Mumpuni, de Lang R, Gower DJ, Sanders KL. 2012a. The Moluccan short-
tailed snakes of the genus Brachyorrhos Kuhl (Squamata: Serpentes: Homalopsidae),
and the status of Calamophis Meyer. The Raffles Bulletin of Zoology 60(2): 501–514.
Murphy JC, Mumpuni, Sanders KL. 2011. First molecular evidence for the phylogenetic
placement of the enigmatic snake genus Brachyorrhos (Serpentes: Caenophidia). Mo-
lecular Phylogenetics and Evolution 61(3):953–957.
Murphy JC, Mumpuni, Sanders KL. 2011. First molecular evidence for the phylogenetic
placement of the enigmatic snake genus Brachyorrhos (Serpentes: Caenophidia). Mo-
lecular Phylogenetics and Evolution 61(3):953–957.
Murphy JC, Muñoz-Mérida A, Auguste RJ, Lasso-Alcala O, Rivas GA, Jowers MJ. 2020.
Evidence for cryptic diversity in the Neotropical water snake, Helicops angulatus (Lin-
naeus, 1758) (Dipsadidae, Hydropsini), with comments on its ecology, facultative re-
productive mode, and conservation. Amphibian & Reptile Conservation 14(3):138-55.
Murphy JC, Voris HK, Auliya M. 2005. A new species of Enhydris (Serpentes: Colubridae:
Homalopsinae) from the Kapuas river system, West Kalimantan, Indonesia. Raffles
Bulletin of Zoology 53(2):115–119.
Murphy JC, Voris HK, Karns DR, Chan-ard T, Suvunrat K. 1999. The ecology of the water
snakes of Ban Tha Hin, Songkhla Province, Thailand. Natural History Bulletin of the
Siam Society 47:129–147.
Murphy JC, Voris HK, Karns DR. 2012b. The dog-faced water snakes, a revision of the ge-
nus Cerberus Cuvier, (Squamata, Serpentes, Homalopsidae), with the description of a
new species. Zootaxa 3484:1–34.
Murphy JC, Voris HK, Murthy BHCK, Traub J, Cumberbatch C. 2012c. The masked water
snakes of the genus Homalopsis Kuhl & van Hasselt, 1822 (Squamata, Serpentes, Hom-
alopsidae), with the description of a new species. Zootaxa 3208:1–26.
Murphy JC, Voris HK. 2005. A new Thai Enhydris (Serpentes: Colubridae: Homalopsinae).
Raffles Bulletin of Zooogy 53(1):143–147.
Murphy JC, Voris HK. 2013. An unusual, fangless short-tailed snake (Squamata, Ser-
pentes, Homalopsidae) from Sumatra, Indonesia. Asian Herpetological Research
4(2):140–146
Murphy JC, Voris HK. 2014. A checklist and key to the homalopsid snakes (Reptilia, Squa-
mata, Serpentes), with the description of new genera. Fieldiana Life and Earth Scienc-
es 2014(8):1–43.
Murphy JC, Voris HK. 2021. A new species of Brachyorrhos from Seram, Indonesia and
notes on fangless homalopsids (Squamata, Serpentes). Philippine Journal of Systematic
Biology 14(2):1–8, i–ii.
Murphy JC. Crutchfield T. 2019. Giant Snakes: A Natural History. Privately Published.
Murphy JC. 1997 Amphibians and reptiles of Trinidad and Tobago. Krieger Publishing,
Malabar, FL.
Murphy JC. 2007a. Homalopsid snakes, evolution in the mud. Krieger Publishing, Mala-
bar, 249 pp.
669
Aquatic Snakes

Murphy JC. 2007b. A review of Enhydris maculosa (Blanford, 1879) and the description of a
related species (Serpentes, Homalopsidae). Hamadryad 31(2):281–287.
Murphy JC. 2011. The nomenclature and systematics of some Australasian homalopsid
snakes (Squamata: Serpentes: Homalopsidae). Raffles Bulletin of Zoology 59(2):229–
236.
Murphy JC. 2011. The nomenclature and systematics of some Australasian homalopsid
snakes (Squamata: Serpentes: Homalopsidae). Raffles Bulletin of Zoology 59(2):229–
236.
Murphy JC. 2012. Marine invasions by non-sea snakes, with thoughts on terrestrial-aquatic-
marine transitions. Integrative and Comparative Biology 52(2):217–226.
Murphy JC. 2012. Synonymized and forgotten, the bird’s head stout-tailed snakes, Calam-
ophis Meyer (Squamata: Serpentes: Homalopsidae). The Raffles Bulletin of Zoology
60(2):515–523.
Murphy JC. Cox MJ. Voris HK. 1999. A key to the sea snakes in the gulf of Thailand. Natu-
ral History Bulletin of the Siam Society 47: 95-108.
Murta-Fonseca RA, Franco FL, Fernandes DS. 2015. Taxonomic status and morphological
variation of Hydrodynastes bicinctus (Hermann, 1804)(Serpentes: Dipsadidae). Zootaxa
4007(1):63-81.
Murthy TSN, Rao KVR. 1986. Observations on some snake-eating birds of the Chilka La-
goon, Orissa. Journal of the Bombay Natural History Society 85:620.
Murthy TSN. 1987. Herpetofauna of the Chilka Lagoon, Orissa, India. British Journal of
Herpetology 21:8–10.
Murthy TSN. 1987. Herpetofauna of the Chilka Lagoon, Orissa, India. British Journal of
Herpetology 21:8–10.
Mushinsky HR, Hebrard JJ, Walley MG. 1980. The role of temperature on the behavioral
and ecological associations of sympatric water snakes. Copeia. 1980:744-54.
Musilová R, Zavadil V. 2011. The Dice snake in the Czech Republic. Zoo Report Brno, De-
cember, special suppl.:1-4.
Myers, C. W. 1969. Snakes of the genus Coniophanes in Pantamá. American Museum No-
vitates 2372: 1- 28.
Nafis G. 2020. California Herps - A Guide to the Amphibians and Reptiles of California.
Available at: [Link] (accessed 10 June 2021).
Nagy ZT, Gvoždík V, Meirte D, Collet MA, Pauwels OS. 2014. New data on the morpholo-
gy and distribution of the enigmatic Schouteden’s sun snake, Helophis schoutedeni (de
Witte, 1922) from the Congo Basin. Zootaxa. 3755(1):096-100.
Nahuat-Cervera PE, Barrao-Nobrga JAL 2021. Coniophanes bipunctatus (Two-spotted
Snake). Diet. Herpetological Review 52(1):151.
Nandi NC. 1984. Bandicoot rat seizing a snake. Journal of the Bombay Natural History So-
ciety 81:183.
Nandi NC. 1984. Bandicoot rat seizing a snake. Journal of the Bombay Natural History So-
ciety 81:183.
Natusch DJ, Lyons JA, Riyanto A, Shine R. 2019. Interspecific divergence in biological
attributes of short-tailed pythons (Python breitensteini and P. brongersmai) from Kali-
mantan and Sumatra. Australian Journal of Zoology 66(4):272-8.
Naulleau G. 1964. Premières observations sur le comportement de chasse et de capture chez
670
Aquatic Snakes

les vipères et les couleuvres. La Terre et la Vie 1:54-76.

Naulleau G, B Schätti. 1997. Natrix maura (Linnaeus, 1758. Pp. 368-369. In: Gasc JP et al.
(Eds). Atlas of amphibians and reptiles in Europe. Societas Europaea Herpetologica y
Muséum National d’Histoire Naturelle, París.
Naumov B, Tzankov N, Popgeorgiev G, Stojanov A, Kornilev Y. 2011. The Dice snake (Na-
trix tessellata) in Bulgaria: distribution and morphology. Mertensiella 18:288-98.
Navarro P, J Lluch, V Roca. 1987. Contribución al conocimiento de la helmintofauna de
los herpetos ibéricos. VI. Parásitos de Natrix maura (Linnaeus, 1758) (Reptilia: Colub-
ridae). Revista Ibérica de Parasitología 47:65-70.
Navarro P, J Lluch. 1993. Strongyloides natricis (Strongyloididae), un nouveaua nematode
parasite de Natrix maura (L. 1758) (Colubridae) en Espagne. Annales d Parasitologie
Humaine et Comparée 68:136-138.
Navarro P, Lluch J, Roca V. 1987. Contribución al conocimiento de la helmintofauna de los
herpetos ibéricos. VI. Parásitos de Natrix maura (Linnaeus, 1758) (Reptilia: Colubri-
dae). Revista Ibérica de Parasitología 47(1):65–70.
Neaves J, Robert SH, Longosoa TH, Andreone F, Rosa GM. 2019. Dietary observations of
the lamprophiid snake Madagascarophis meridionalis (Domergue, 1987) in southeast-
ern Madagascar. Herpetology Notes 12:75-8.
Neill WT. 1958. The occurrence of amphibians and reptiles in saltwater areas, and a bib-
liography. Bulletin of Marine Science of the Gulf and Caribbean 8: 1–97.
Neill WT. 1964. Taxonomy, natural history, and zoogeography of the Rainbow Snake,
Farancia erytrogramma (Palisot de Beauvois). American Midland Naturalist 1964
1:257-95.
Neill WT. 1965. Notes on aquatic snakes, Natrix and Tretanorhinus, in Cuba. Herpeto-
logica 21: 172–181.
Neill WT, Allen ER. 1956. Secondarily ingested food items in snakes. Herpetologica
12(3):172–174.
Neill WT, Allen R. 1962. Parturient anaconda, Eunectes gigas Latreille, eating own abortive
eggs and foetal membranes. Quarterly Journal of the Florida Academy of Sciences.
1;25(1):73-5.
Neuman C, Mebert K. 2011. Migration behavior of endangered Dice snakes (Natrix tessel-
lata) at the river Nahe, Germany. Mertensiella 18:39-49.
Neuman-Lee LA, McGaugh SE, Pfrender M, Janzen FJ. 2011. Using mitochondrial DNA to
determine the identity and origin of a gartersnake found in Alaska. Journal of Herpe-
tology 45(1):63-5.
Ng TH. 2011. A record of the banded file snake, Acrochordus granulatus (Reptilia: Squa-
mata: Acrochordidae) in a freshwater habitat in Singapore. Nature in Singapore 4:91-3.
Nguyen QT, David P, Tran TT, Luu QV. Le KQ, Ziegler T. 2010. Amphiesmoides ornati-
ceps (Werner, 1924), an addition to the snake fauna of Vietnam, with a redescription
and comments on the genus Amphiesmoides Malnate, 1961 (Squamata: Natricidae).
Revue Suisse de Zoologie 117 (1): 45-56.
Nguyen TQ, Thy N, Sanders K, Murphy J, Lobo AS, Gatus J, Diesmos AC, Gonzalez,
JC, Hamilton A, Allison A, Tallowin O, O’Shea M, Parker F, Gillespie G, Stubbs A,
Maglangit EP, Delima EM and Pitogo K. 2022. Acrochordus granulatus (amend-
ed version of 2021 assessment). The IUCN Red List of Threatened Species 2022:
671
Aquatic Snakes

e.T176769A211491360. [Link]
[Link].
Nguyen TV, Brakels P, Maury N, Sudavanh S, Pawangkhanant P, Idiiatullina S, Lorphengsy
S, Inkhavilay K, Suwannapoom C, Poyarkov NA. 2020. New herpetofaunal observations
from Laos based on photo records. Amphibian & Reptile Conservation 14(2):218–249.
Nguyen TV, Brakels P, Maury N, Sudavanh S, Pawangkhanant P, Idiiatullina S, Lorphengsy
S, Inkhavilay K, Suwannapoom C, Poyarkov NA. 2020. New herpetofaunal observations
from Laos based on photo records. Amphibian & Reptile Conservation 14(2):218–249.
Nicholson E. 1874. Indian snakes. An elementary treatise on ophiology, with a descriptive
catalogue of the snakes found in India and the adjoining countries. Higginbotham and
Co., Madras, 186 pp.
Nilsen TH. 1978. Lower Tertiary laterite on the Iceland–Faeroe ridge and the Thulean land
bridge. Nature 274(5673):786-8.
Nobbs M, Blamires SJ. 2004. Fordonia leucobalia (yellowbellied mangrove snake) and Myron
richardsonii (Richardson’s mangrove snake). Diurnal feeding and prey type. Herpetolog-
ical Review 35:67–68.
Nobuhiro M, Tu AT 1988. Isolation and primary structure of the major toxin from sea
snake, Acalyptophis peronii, venom. 260(1), 0–17. doi:10.1016/0003-9861(88)90418-3
Noonloy T, Kunya K, Chanhome L, Sumontha M, Chomngam N, Pauwels OS. 2018. Cr-
ab-ripping: an unusual feeding behavior newly recorded in freshwater snakes. Bulletin
of the Chicago Herpetological Society 53(3):53-6.
Norval G, Mao J-J, Chu H-P. 2005. Amphiesma stolatum (Striped Keelback). Prey size. Her-
petological Review 36 (2):187.
Norval G, Huang S-C, Mao J-J. 2007. Amphiesma stolatum (Striped keelback): Predation of
Polypedates megacephalus (Java Treefrog). Herpetological Bulletin 101:39-40.
Norval G, Mao J-J, Huang S-C. 2008. Schlüpflinge der Gestreiften Kieltückennatter (Am-
phiesma stolatum [Linnaeus, 1758]) fressen giftige Schwarznarbenkröten (Bufo melanos-
ticus Schneider, 1799), mit Anmerkungen zum Futterspektrum von A. stolatum. Sauria,
Berlin 30(2):51-54.
Norval G, Mao J-J, Huang S-C, Hou C, Lee J. 2010. Xenchrophis piscator (Checkered Keel-
back): Predation. Herpetological Bulletin 112:39-41.
Noss RF, Platt WJ, Sorrie BA, Weakley AS, Means DB, Costanza J, Peet RK. 2015. How glob-
al biodiversity hotspots may go unrecognized: lessons from the North American Coastal
Plain. Diversity and Distributions 21(2):236-44.
Nowak EM. 2006. Monitoring surveys and radio-telemetry of narrow-headed gartersnakes
(Thamnophis rufipunctatus) in Oak Creek, Arizona. Unpublished report to Arizona
Game and Fish Department, State Wildlife Grant, Narrow-headed Gartersnake Plan-
ning.
Oberg K, Shelton JM, Gardiner N, Jackson PR. 2009. Discharge and other hydraulic mea-
surements for characterizing the hydraulics of lower Congo River, Proceedings of the
international association for hydraulic research congress 33:8.
Obst FJ, Strasser P. 2011. Das sachsische Vorkommen der Wurfelnatter im elbtal bei
Meißen. Mertensiella 18:58-71.
Oeffner J, Lauder GV. 2012. The hydrodynamic function of shark skin and two biomimetic
applications. Journal of Experimental Biology 215(5):785–795.
672
Aquatic Snakes

Oliveira JL, Michela M, Marques OAV. 2003. Goesophis brasiliensis (NCN) Reproduction.
Herpetological Review 34:251–252
Oliveira L, Scartozzoni RR, de Almeida-Santos SM, Jared C, Antoniazzi MM, da Graça
Salomão M. 2016. Morphology of Duvernoy’s glands and maxillary teeth and a pos-
sible function of the Duvernoy’s gland secretion in Helicops modestus Günther, 1861
(Serpentes: Xenodontinae). South American Journal of Herpetology 11(1):54-65.
OmPraba G, Chapeaurouge A, Doley R, Devi KR, Padmanaban P, Venkatraman C, Velm-
urugan D, Lin Q, Kini RM. 2010. Identification of a novel family of snake venom pro-
teins Veficolins from Cerberus rynchops using a venom gland transcriptomics and pro-
teomics approach. Journal of Proteome Research 9(4):1882–1893.
Onn CK, Ahmad NO. 2009. Distribution and natural history notes on some poorly known
frogs and snakes from Peninsular Malaysia. Herpetological Review 40(3):294-301.
Onofre N, Sampaio L. 2020. Feeding Ecology of Short-Toed Snake-Eagle (Circaetus gallicus
[Gmelin, 1788]) in the Montados of Iberian Peninsula. Silva Lusitana 28(2):155–179.
Orlov NL. 2000. Distribution, biology, and comparative morphology of the snakes of Xe-
nopeltis genus (Serpentes: Macrostomata: Xenopeltidae) in Vietnam. Soot. J. Herpetol. 7
(2): 103-11.
Ortega-Andrade HM, Bermingham J, Aulestia C, Paucar C. 2010. Herpetofauna of the Bilsa
Biological Station, province of Esmeraldas, Ecuador. Check List. 6(1):119-54.
O’Shea M. 1986. Snakes of the Homalopsinae (Serpentes: Colubridae) from the southern
Trans-Fly, Papua New Guinea. The Herptile 11(4):155–163.
O’Shea M. 1996. A Guide to the Snakes of Papua New Guinea. Independent Publishing,
Port Moresby, 239 pp.
O’Shea M. 2005. Venomous Snakes of the World, Princeton University Press
O’Shea M. 2018. The Book of Snakes: A life-size guide to six hundred species from around
the world. University of Chicago Press, Chicago.
O’Shea M, Kaiser H. 2016. The first female specimen of the poorly known Arfak Stout-tailed
Snake, Calamophis sharonbrooksae Murphy, 2012, from the Vogelkop Peninsula of West
New Guinea, with comments on the taxonomic history of primitive homalopsids. Am-
phibian & Reptile Conservation 10(2): 1–10 (e122).
Ota H, Chen SL, Lin JT, Toriba M. 1999. Taxonomic Status of the Taiwanese Populations of
Rhabdophis tigrinus (Squamata: Colubridae) Morphological and Karyological Assess-
ment. Japanese Journal of Herpetology 18(1):1-6.
Ota H, Mori A. 1985. Notes on the fourth collected specimen of Rhabdophis tigrinus formo-
sanus and the morphological features characteristic of this subspecies. Japanese Journal
of Herpetology 11:41-45
Ovadia M. 1984. Embryonic Development of Duvernoy’s Gland in the Snake Natrix tessella-
ta (Colubridae). Copeia 1984(2):516-21.
Pakenham RH. The reptiles and amphibians of Zanzibar and Pemba Islands (with a note on
the freshwater fishes). Journal of East African Natural History. Society National Muse-
um. 1983(177):40.
Palci A, Hutchinson MN, Caldwell MC, Lee MSY. 2017. The Morphology of the Inner Ear of
Squamate Reptiles and Its Bearing on the Origin of Snakes. Royal Society Open Science
4: 170685. [Link]
Palci A, Hutchinson MN, Caldwell MW, Lee MSY. 2017. The morphology of the inner ear of
673
Aquatic Snakes

squamate reptiles and its bearing on the origin of snakes. Royal Society Open Science
4(170685):1-15.
Palci A, Seymour RS, Van Nguyen C, Hutchinson MN, Lee MSY, Sanders KL. 2019.
Novel vascular plexus in the head of a sea snake (Elapidae, Hydrophiinae) revealed
by high-resolution computed tomography and histology. Royal Society Open Sci-
ence 6, 191099. doi:10.1098/rsos.191099.
Palcu DV. 2018. The Dire Straits of Paratethys: Dating, matching, and modeling connectiv-
ity between the Miocene seas of Eurasia. UU Dept. of Earth Sciences.
Palcu DV, Patina IS, Șandric I, Lazarev S, Vasiliev I, Stoica M, Krijgsman W. 2021. Late
Miocene megalake regressions in Eurasia. Scientific reports 11(1):1-2.
Pallas PS. 1771. Reise durch verschiedene Provinzen des russischen Reichs. St. Petersburg,
1771-1776. 3 vol. Eng. abridged trans. in J. Trusler, The Habitable World Described.
London, 1790.
Pallas PS. 1814. Zoographia Rossa-Asiatica, sistens omnium anmalium in extenso Impe-
rio Rossico et adjacentibus maribus observatorum recensionem, domicilia, mores, et
descriptiones anatomen atque icones plurimorum. Vol. 3. Petropoli: Academia Scien-
tiarum, 422 pp.
Palomares MLD, Pauly D. Editors. 2021. SeaLifeBase. World Wide Web electronic publica-
tion. http:// [Link], version (04/2021). Accessed on 6/15/2021.
Pandolfi J.M., R.H. Bradbury, E. Sala, T.P. Hughes, K.A. Bjorndal, R.G. Cooke, D. McArdle,
L. McClenachan, M.J.H. Newman, G. Paredes, et al. 2003. Global trajectories of the
long-term decline of coral reef ecosystems. Science 301:955–958
Panzera A, Maneyro R. 2014. Feeding Biology of Lygophis anomalus (Dipsadidae, Xeno-
dontinae). South American Journal of Herpetology 9(2):75-82.
Parameswaran KN. 1954. On the viviparous habit of the fresh-water snake, Enhydris dus-
sumieri (Smith). Current Science 23(1):27–28.
Parameswaran KN. 1962. The foetal membranes and placentation of Enhydris dussumi-
eri (Smith). Proceedings of the Indian Academy of Science B 56:302–327.
Parker F. 1982 [1983]. The snakes of Western Province. Wildlife in Papua New Guinea.
Konedobu: Division of Wildlife 82:1–78.
Parker MR, Young BA & Kardong KV. 2008. The forked tongue and edge detection in
snakes (Crotalus oreganus): an experimental test. Journal of Comparative Psychology
122(1):35–40.
Passos P, Fernandes DS. 2005. Variation and taxonomic status of the aquatic coral snake
Micrurus surinamensis (Cuvier, 1817) (Serpentes: Elapidae). Zootaxa 953(1):1-8.
Pattishall A & Cundall D. 2008. Dynamic changes in body form during swimming in the
water snake Nerodia sipedon. Zoology 111(1):48–61.
Pauwels OS, Albert JL, Arrowood H, Mvele C, Casanova M, Dodane JB, Morgan J, Pri-
mault L, Thepenier L, Fenner JN. 2017. Miscellanea Herpetologica Gabonica X. Bul-
letin of the Chicago Herpetological Society 52(8):133-8.
Pauwels OS, Carlino P, Chirio L, Huijbregts B, Leuteritz T, Rousseaux D, Tobi E, Vigna
C, Van Neer W. Miscellanea herpetologica gabonica VII. The Bulletin of the Chicago
Herpetological Society. 2017;52(1):1-7.
Pauwels OS, G. Lenglet, J.-F. Trape, A. Dobois. 2000. Grayia smithii (Leach, 1818)

674
Aquatic Snakes

Smith’s African Water Snake Africa Herp News 2000 (31):7-8.

Pauwels OS, Sumontha M. 2016. Taxonomic identity of two enigmatic aquatic snake popu-
lations (Squamata: Homalopsidae: Cerberus and Homalopsis) from southern Thailand.
Zootaxa 4107(2):293–300.
Pauwels OS, Toham AK, Mamonekene V. 2002. Ethnozoology of the dibomina (Serpentes:
Colubridae: Grayia ornata) in the Massif du Chaillu, Gabon. Hamadryad 27:136-41.
Pauwels OS, Wallach V, David P. 2008. Global diversity of snakes (Serpentes; Reptilia) in
freshwater. Hydrobiologia 595(1):599-605.
Pauwels OSG, Kadeyeva M, Kovshar V, Sakharbayev A, Sarayev FA, Sarsengaliyev S, Ukhov
S, Yerbulekov S. 2020. Colonization of artificial islands in the Kazakh sector of the Cas-
pian Sea by the aquatic snake Natrix tessellata (Squamata: Natricidae). Bulletin of the
Chicago Herpetological Society 55(7):133-140.
Peng L, Lu C, Huang S, Guo P, Zhang Y. 2014. A new species of the genus Thermophis
(Serpentes: Colubridae) from Shangri-La, Northern Yunnan, China, with a proposal
for an eclectic rule for species delimitation. Asian Herpetol Res. 5(4):228-39.
Pereira DN, Stender-Oliveira FE, Rodrigues MG, Bérnils RS. 2007. Distribution and habitat
use of Sordellina punctata (Serpentes, Colubridae), with a new record from State of São
Paulo, Brazil. Herpetological Bulletin. 100:18-22.
Pérez i de Lanuza G. 2011. Depredación de Anguilla anguilla por Natrix maura en la desem-
bocadura del río Ebro. Boletín de la Asociciación Herpetológica Española 22:51-53.
Pérez-Bote, JL, R Roso. 2009. Depredacion de Natrix maura por Sander lucioperca en el SO
de la Peninsula Iberica. Boletín de la Asociciación Herpetológica Española 20:117-119
Perkins MW, Eason PK. 2017a. Nerodia erythrogaster. (Plain-bellied Watersnake). Diet.
Herpetological Review 48:215-216.
Perkins MW, Eason PK. 2017b. Nerodia rhombifera (Diamondback Watersnake). Diet. Her-
petological Review 48:216.
Perkins S. 2021. The rise and fall of the world’s largest lake. Science 372 (6547).
Perthuis A. 2008. Situation du Circaète en Loir–et–Cher. La Plume du Circaète 6:7–9.
Petch S. 1991. Notes on a case of double-clutching in the viperine snake, Natrix maura.
Herptile 16:118-119.
Petermann H, Gauthier JA. 2018. Fingerprinting snakes: paleontological and paleoecologi-
cal implications of zygantral growth rings in Serpentes. PeerJ, 6, e4819.
Peters WCH. 1863. Über die von Hrn. Dr. F. Jagor in Siam gesammelten Amphibien. Pp.
245–247. In: Monatsberichte der Königlichen Preussische Akademie des Wissen-
schaften zu, Königliche Akademie der Wissenschaften, Berlin.
Peters WCH. 1871. Uber neue Reptilien aus Ostafrika und Sarawak (Borneo), vorzüglich
aus der Sammlung des Hrn Marquis J. Doria zu Genua. Pp. 566–581. In: Monats-
berichte der Königlichen Preussische Akademie des Wissenschaften zu, Königliche
Akademie der Wissenschaften, Berlin.
Pettus D. 1958. Water relationships in Natrix sipedon. Copeia. 1958(3):207-11.
Petzold HG. 1967. Some remarks on the breeding biology and the keeping of Tretanorhi-
nus variabilis, a water snake of Cuba. Herpetologica. 1967 Sep 20:242-6.
Pickwell GV, Vick JA, Shipman WH, Grenan MM. 1972. Production, toxicity, and prelimi-
nary pharmacology of venom from the sea snake, Pelamis platurus. In: Worthen LR (Ed)
Food-drugs from the sea. Marine Technology Society, Washington, D.C., 247–265.
675
Aquatic Snakes

Pienaar, U. de V. 1966. The reptiles of the Kruger National Park, National Parks Board Pre-
toria 223 pp.
Pinheiro RT, Silva PH, Oliveira MS, Dias MT, Campos CE. 2020. Predation on the Am-
azon water snake Hydrops martii (Squamata: Dipsadidae) by the redeye piranha Ser-
rasalmus rhombeus (Serrasalmidae). Herpetology Notes 13:553-4.
Pinto RR, Fernandes R. 2004. Reproductive biology and diet of Liophis poecilogyrus poe-
cilogyrus (Serpentes, Colubridae) from southeastern Brazil. Phyllomedusa: Journal of
Herpetology. 2004 3(1):9-14.
Pizzato L, Marques OAV. 2006. Interpopulational variation in sexual dimorphism, re-
productive output, and parasitism of Liophis miliaris (Colubridae) in the Atlantic
Forest of Brazil. Amphibia-Reptilia 27: 37–46.
Pizzatto L, Marques OAV. 2007. Reproductive ecology of boine snakes with emphasis on
Brazilian species and a comparison to pythons”. South American Journal of Herpe-
tology 2:107–122. doi:10.2994/1808-9798(2007)2[107:reobsw][Link];2.
Pizzatto L, Almeida-Santos SM, Shine R. 2007. Life‐history adaptations to arboreality in
snakes. Ecology. 88(2):359-66.
Pizzatto L, Jordão RS, Marques OA. 2008. Overview of reproductive strategies in Xeno-
dontini (Serpentes: Colubridae: Xenodontinae) with new data for Xenodon neuwiedii
and Waglerophis merremii. Journal of Herpetology 42(1):153-62.
Pizzatto L, Madsen T, Brown GP, Shine R. 2009. Spatial ecology of hatchling water py-
thons (Liasis fuscus) in tropical Australia. Journal of Tropical Ecology 25(2):181-91.
Platt SG, Rainwater TR, Gibbons JW. 2020. Nerodia rhombifera (Diamondback Water-
snake) and Nerodia erythrogaster (Plain-bellied Watersnake). Kleptoparasitism. Herpe-
tological Review 51(4):871-871.
Pleguezuelos JM. 2018. Culebra de collar mediterránea – Natrix astreptophora. In: Enciclo-
pedia Virtual de los Vertebrados Españoles. Sanz JJ, Martínez-Freiría F. (Eds.). Museo
Nacional de Ciencias Naturales, Madrid.
Pleguezuelos JM. 2016. Culebra de collar mediterránea – Natrix astreptophora. En: Enci-
clopedia Virtual de los Vertebrados Españoles. Salvador, A., Marco, A. (Eds.). Museo
Nacional de Ciencias Naturales, Madrid. [Link]
Pleguezuelos JM, M Feriche. 2003. Anfibios y reptiles. Los Libros de la Estrella nº 18.
Granada, Guías de la Naturaleza. Diputación de Granada, Granada.
Pleguezuelos JM, M Moreno. 1989. Alimentación primaveral de Natrix maura (Linné,
1758) (Ophidia, Colubridae) en el SE de la Península Ibérica. Revista Española de
Herpetología 3:221-236.
Pleguezuelos JM, Villafranca C. 1997. La distribución altitudinal de los anfibios y reptiles
ibéricos. En: Pleguezuelos, J. M. (Ed). Distribución y Biogeografía de los anfibios y rep-
tiles de España y Portugal. Monografías Revista Española de Herpetología 3:321-341.
Pleguezuelos JM. 2018. Culebra de collar mediterránea – Natrix astreptophora. In: Enciclo-
pedia Virtual de los Vertebrados Españoles. Sanz JJ, Martínez-Freiría F. (Eds.). Museo
Nacional de Ciencias Naturales, Madrid. [Link]
Pleguezuelos JM, Ontiveros D. 2010. Diet and prey selection by the Short-toed Eagle,
Circaetus gallicus. Pp. 219–225. In: Zuberogoitia I, Martínez JE. (Eds). Forest-dwelling
raptors. Conservation, ecology, behaviour and management implications, Agriculture
Department, Diputación Foral de Bizkaia. Bizkaia, Spain.
676
Aquatic Snakes

Pleguezuelos JM, Brito JC, Fahd S, Feriche M, Mateo JA, Moreno-Rueda G, Reques R, San-
tos X. 2010. Setting conservation priorities for the Moroccan herpetofauna: the utility of
regional red lists. Oryx 44(4):501–508.
Pokrant F, Kindler C, Ivanov M, Cheylan M, Geniez P, Böhme W, Fritz U. 2016. Integra-
tive taxonomy provides evidence for the species status of the Ibero-Maghrebian grass
snake Natrix astreptophora. Biological Journal of the Linnean Society, 118(4):873-88.
Polhemus D. 2007. Tectonic geology of Papua. Pp. 137–164. In: The ecology of Papua. Mar-
shall A, Beehler B (eds.). Periplus Editions, Singapore.
Pongcharoen C, Voris HK, Seelanan T, Pradatsundarasar A-O, Thirakhupt K (2016). Diet,
female reproduction and conservation of Jagor’s water snake, Enhydris jagorii in Bung
Ka Loh wetland, Uttaradit province, Thailand. Agriculture and Natural Resources, Vol-
ume 50, Issue 3, Pages 204-210
Pongcharoen C, Voris HK, Seelanan T, Pradatsundarasar AO, Thirakhupt K. 2016. Diet,
female reproduction and conservation of Jagor’s water snake, Enhydris jagorii in Bung
Ka Loh wetland, Uttaradit province, Thailand. Agriculture and Natural Resources
50(3):204–210.
Poo S, M-R, Devan-Song A, 2016. Xenochrophis flavipunctatus (Yellow-spotted Keelback
Watersnake). Diet. Herpetological Review 47(2):319.
Pope CH, Granger W. 1929. Notes on reptiles from Fukien and other Chinese provinces.
Bulletin of the American Museum of Natural History 58:335–487.
Pope CH. 1935. The reptiles of China. The Natural History of Central Asia. American Muse-
um of Natural History 1935:604.
Porto M, Caramaschi U. Notes on taxonomic status, biology and distribution of Ptychophis
flavovirgatus Gomes, 1915 (Ophidia, Colubridae). Anais da Academia Brasileira de
Ciências. 1988;60:471-5.
Pough FH. 1973. Heart rate, breathing and voluntary diving of the elephant trunk snake,
Acrochordus javanicus. Comparative Biochemistry and Physiology Part A: Physiology
44(1):183-9.
Povel DE & Van Der Kooij J. 1997. Scale sensillae of the file snake (Serpentes: Acrochor-
didae) and some other aquatic and burrowing snakes. Netherlands Journal of Zoology
47(4):443–456.
Powell R, Conant R, Collins JT. 2016. Peterson field guide to reptiles and amphibians of
eastern and central North America. Houghton Mifflin Harcourt; 2016 Apr 12.
Pratt KL & Franklin CE. 2010. Temperature independence of aquatic oxygen uptake in an
air-breathing ectotherm and the implications for dive duration. Comparative Biochem-
istry and Physiology - A Molecular and Integrative Physiology 156(1):42–45.
Pratt KL, Campbell HA, Watts ME, Franklin CE. 2010. Environmental and ecological fac-
tors influencing dive behaviour in the freshwater snake Acrochordus arafurae: a field-
based telemetric study. Marine and Freshwater Research 61(5):560-7.
Pratt, K., 2009. Metabolic rate, respiratory partitioning and the implications for dive dura-
tion in the bimodally respiring Arafura filesnake, Acrochordus arafurae. PhD disserta-
tion, School of Biological Sciences, The University of Queensland.
Preston DL, Johnson PT. 2012. Importance of native amphibians in the diet and distribution
of the Aquatic Gartersnake (Thamnophis atratus) in the San Francisco Bay area of Cali-
fornia. Journal of Herpetology 46(2):221-8.
677
Aquatic Snakes

Prieto YA, Giraudo AR, López MS. 2012. Diet and sexual dimorphism of Liophis poecilo-
gyrus (Serpentes, Dipsadidae) from the wet regions of Northeast Argentina. Journal of
Herpetology 46(3):40.
Procter JB. 1923. XXII.—On a new genus and species of colubrine snake from SE Brazil.
Journal of Natural History 11(62):227-30.
Prodon R, Geniez P, Cheylan M, Besnard A. 2020. Amphibian and reptile phenology:
the end of the warming hiatus and the influence of the NAO in the North Mediterra-
nean. International Journal of Biometeorology 64:423–432.
Proenca HC, Lourenco-de-Moraes R, Zawadzki CH. 2015. Thamnodynastes strigatus
(Corredeira) Diet. Herpetological Review 46(4)651-652.
Prudente, A.L.C. and R. Brandão. 1998. Gomesophis brasiliensis. Herpetological Review
29(2): 112.
Punay EY 1975. Commercial sea snake fisheries in the Philippines. In: Dunson WA, ed.
The Biology of Sea Snakes. Baltimore, London & Tokyo. pp 417–462.
Purkayastha J, KALITA J, Brahma RK, Doley R, DAS M. 2018. A review of the relation-
ships of Xenochrophis cerasogaster Cantor, 1839 (Serpentes: Colubridae) to its conge-
ners. Zootaxa 4514 (1):126-136, DOI: 10.11646/zootaxa.4514.1.10.
Pyron RA & Burbrink FT. 2014. Early origin of viviparity and multiple reversions to ovi-
parity in squamate reptiles. Ecology Letters 17(1):13–21.
Pyron RA, Burbrink FT, Colli GR, de Oca AN, Vitt LJ, Kuczynski CA, Wiens JJ. 2011.
The phylogeny of advanced snakes (Colubroidea), with discovery of a new subfamily
and comparison of support methods for likelihood trees. Molecular Phylogenetics
and Evolution. 58 (2): 329–342. doi:10.1016/[Link].2010.11.006.
Pyron RA, Burbrink FT, Wiens JJ. 2013. A phylogeny and revised classification of Squama-
ta, including 4161 species of lizards and snakes. BMC Evolutionary Biology 13(1):1–54.
Pyron RA, Burbrink FT. 2012. Extinction, ecological opportunity, and the origins of global
snake diversity. Evolution: International Journal of Organic Evolution 66(1):163-78.
Pyron, R. A., Kandambi, H. K. D. K., Hendry, C. R., Pushpamal, V., Burbrink, F. T., So-
maweera, R. (2013). Genus-level phylogeny of snakes reveals the origins of species
richness in Sri Lanka. Molecular Phylogenetics and Evolution, 66 (3): 969-978.
Quah ES, Grismer LL, Wood PL, Thura MK, Zin T, Kyaw H, Lwin N, Grismer MS, Mur-
doch ML. 2017. A new species of Mud Snake (Serpentes, Homalopsidae, Gyiophis
Murphy & Voris, 2014) from Myanmar with a first molecular phylogenetic assess-
ment of the genus. Zootaxa 4238(4):571–582.
Quah ES, Grismer LL, Wood PL, Thura MK, Zin T, Kyaw H, Lwin N, Grismer MS, Mur-
doch ML. 2017. A new species of Mud Snake (Serpentes, Homalopsidae, Gyiophis
Murphy & Voris, 2014) from Myanmar with a first molecular phylogenetic assessment
of the genus. Zootaxa 4238(4):571–582.
Quah ES, Wood PL Jr., Grismer L, Sah SA. 2018. On the taxonomy and phylogeny of the
rare Selangor Mud Snake (Raclitia indica) Gray (Serpentes, Homalopsidae) from Pen-
insular Malaysia. Zootaxa 4514(1):53–64.
Queiroz AD, Lawson R. 2008. A peninsula as an island: multiple forms of evidence for
overwater colonization of Baja California by the gartersnake Thamnophis validus. Bio-
logical Journal of the Linnean Society 95(2):409-24.

678
Aquatic Snakes

Quillon J, Lamouille M. 1978. A propos d’éclosions chez les couleuvres Natrix natrix et Na-
trix maura. Bulletin de la Société herpetologique de France 8:18-28.
Quintela FM 2013. Liophis poecilogyrus (Yellow-bellied Liophis): Copulation. Herpetologi-
cal Bulletin (123) 2-406.
Quintela FM. 2010. Liophis poecilogyrus sublineatus (Serpentes: Dipsadidae) bite and
symptoms of envenomation. Herpetology Notes, 3:309-311.
Radhakrishnan C. 1999. Lizards and Snakes of four conservation areas in the Idukki dis-
trict, Kerala state. Records of the Zoological Survey of India 97(2):155.
Rage J. 1983. Palaeophis-colossaeus nov-sp (the largest snake known) from the Eocene of
Mali, with remarks on the genus problem within the Palaeopheinae. Comptes rendus
de l academie des sciences serie iii-sciences de la vie-life sciences, 296(22):1029-32.
Rage J.-C., Auge M., 1993. Squamates from the Cainozoic of the western part of Europe. A
Review. Revue de Paléobiologie, 7: 199-216.
Rage JC, Bajpai S, Thewissen JG, Tiwari BN. 2003. Early Eocene snakes from Kutch, West-
ern India, with a review of the Palaeophiidae. Geodiversitas 25(4):695-716.
Rage JC, Escuilié F. 2003. Le Cénomanien: étage des serpents bipèdes. Carnets de Geolo-
gie, (A01-fr), pp.1-11.
Rahadian R, Das I. 2012. A new record of Pseudoxenodon inornatus (Boie In: Boie, 1827)
from Gunung Gedeh National Park, West Java, Indonesia (Squamata: Pseudoxeno-
dontidae), Hamadryad 36:174–177.
Rahman SC, Reza AA. 2013. Algal fouling on dog-faced water snake (Cerberus rynchops,
Serpentes: Homalopsidae). Herpetology Notes 6:373–374.
Rainboth WJ. 1996. Fishes of the Cambodian Mekong. Food and Agriculture Organiza-
tion of the United Nations, Rome.
Rajabizadeh M, Javanmardi S, Rastegar-Pouyani N, Karamiani R, Yusefi M, Salehi H,
Joger U, Mebert K, Esmaeili H, Parsa H, Gholi Kami H, Rastegar-Pouyani E. 2011.
Geographic variation, distribution, and habitat of Natrix tessellata in Iran. Mertensiella
18:414-30.
Ramberg L, Hancock P, Lindholm M, Meyer T, Ringrose S, Sliva J, Van As J, Vander Post
C. Species diversity of the Okavango delta, Botswana. Aquatic sciences. 2006 Oct
1;68(3):310-37.
Rasmussem A. 2010. Hydrophis torquatus. The IUCN Red List of Threatened Species 2010:
e.T176723A7290809. [Link]
[Link]. Accessed on 28 May 2022.
Rasmussen A, Lobo A. 2010. Hydrophis stricticollis. The IUCN Red List of Threatened
Species 2010: e.T176733A7292717. [Link]
[Link].
Rasmussen A, Gatus J. 2010. Hydrophis lamberti. The IUCN Red List of Threatened Spe-
cies 2010: e.T176754A7297862. [Link]
[Link].
Rasmussen A, Lobo A, Sanders K. 2010e. Hydrophis caerulescens. The IUCN Red List
of Threatened Species 2010: e.T176715A7288947. [Link]
[Link].
Rasmussen A, Lobo A. 2010c. Hydrophis fasciatus. The IUCN Red List of Threatened Spe-
cies 2010: e.T176744A7295660. [Link]
679
Aquatic Snakes

[Link].
Rasmussen A, Lobo A. 2010d. Hydrophis nigrocinctus. The IUCN Red List of Threatened
Species 2010: e.T176722A7290670. [Link]
[Link].
Rasmussen A, Lukoschek V, Lobo A. 2010h. Hydrophis ornatus. The IUCN Red List of
Threatened Species 2010: e.T176737A7293481. [Link] IUCN.
[Link].
Rasmussen A, Lukoschek V. 2010. Hydrophis czeblukovi. The IUCN Red List of Threatened
Species 2010: e.T176730A7292260. [Link]
[Link].
Rasmussen A, Sanders K, Lobo A, Gatus J. 2010j. Hydrophis spiralis. The IUCN Red List
of Threatened Species 2010: e.T176717A7289307. [Link]
[Link].
Rasmussen A, Sanders K, Lobo A. 2010f. Hydrophis cyanocinctus. The IUCN Red List
of Threatened Species 2010: e.T176726A7291386. [Link]
[Link].
Rasmussen A, Sanders K, Lobo A. 2010g. Hydrophis brookii. The IUCN Red List of Threat-
ened Species 2010: e.T176709A7287652. [Link] [Link].2010-4.
[Link].
Rasmussen A, Sanders K, Lobo A. 2010i. Hydrophis lapemoides (errata version published in
2016). The IUCN Red List of Threatened Species 2010: e.T176741A98102816. https://
[Link]/10.2305/[Link].
Rasmussen A, Sanders K. 2010. Hydrophis belcheri. The IUCN Red List of Threatened Spe-
cies 2010: e.T176751A7297371. [Link]
[Link].
Rasmussen A, Sanders K., Lobo A. 2018. Hydrophis viperinus (amended ver-
sion of 2010 assessment). The IUCN Red List of Threatened Species 2018:
e.T176711A136259318. [Link]
[Link].
Rasmussen A. Lobo A. 2010e. Hydrophis obscurus. The IUCN Red List of Threatened Spe-
cies 2010: e.T176752A7297560. [Link]
[Link].
Rasmussen AR, Lobo A. 2010a. Hydrophis bituberculatus. The IUCN Red List of Threatened
Species 2010: e.T176728A7291867. [Link] [Link].
[Link].
Rasmussen AR, Auliya M, Böhme W. 2001. A new species of the sea snake genus Hy-
drophis (Serpentes: Elapidae) from a river in West Kalimantan (Indonesia, Borneo).
Herpetologica 1:23-32.
Rasmussen AR, Boesl-Riquier M. Böhme, W. 2001. A new species of the sea snake Genus
Hydrophis (Serpentes: Elapidae) from a river in West Kalimantan (Indonesia, Borneo).
Herpetologica, 57, 23-32.
Rasmussen AR, Crowe-Riddell JM, Courtney T, Sanders K. 2021. Hydrophis curtus. The
IUCN Red List of Threatened Species 2021: e.T176746A132780885. [Link]
10.2305/[Link].
Rasmussen AR, Elmberg J, Gravlund P, Ineich I. 2011b. Sea snakes (Serpentes: subfamilies
680
Aquatic Snakes

Hydrophiinae and Laticaudinae) in Vietnam: a comprehensive checklist and an updated

identification key. Zootaxa 2894: 1–20
Rasmussen AR, Elmberg J. 2009. Head for my tail: A new hypothesis to explain how ven-
omous sea snakes avoid becoming prey. Marine Ecology, 30(4), 385–390. doi:10.1111/
j.1439-0485.2009.00318.x .
Rasmussen AR, Guinea M. 2010. Hydrophis vorisi. The IUCN Red List of Threatened Spe-
cies 2010: e.T176768A7300648. [Link] [Link].
[Link].
Rasmussen AR, Ineich I, Elmberg J, McCarthy C. 2011c. Status of the Asiatic Sea Snakes of
the Hydrophis nigrocinctus group (H. nigrocinctus, H. hendersoni, and H. walli; Elapidae,
Hydrophiinae). Amphibia-Reptilia 32: 459-464.
Rasmussen AR, Ineich I. 2000. Sea snakes of New Caledonia and surrounding waters (Ser-
pentes: Elapidae): first report on the occurrence of Lapemis curtus and description of
new species from the genus Hydrophis. Hamadryad 25 (2): 91-99.
Rasmussen AR, Lobo A, Sanders K, Guinea M. 2010a. Hydrophis atriceps. The IUCN Red
List of Threatened Species 2010: e.T176735A7293104. [Link] IUCN.
UK.2010 [Link].
Rasmussen AR, Lobo A, Sanders K. 2010d. Hydrophis cantoris. The IUCN Red List of
Threatened Species 2010: e.T176713A7288602. [Link] IUCN.
[Link].
Rasmussen AR, Lobo A. 2010b. Kerilia jerdoni. The IUCN Red List of Threatened Spe-
cies 2010: e.T176767A7300378. [Link]
[Link].
Rasmussen AR, Murphy JC, Ompi M, J. Gibbons W, Uetz P. 2011. Marine Reptiles. PlosOne
6e27373.
Rasmussen AR, Sanders K, Guinea M, Amey A 2014. Sea snakes in Australian waters (Ser-
pentes: subfamilies Hydrophiinae and Laticaudinae)—a review with an updated identifi-
cation key. Zootaxa 3869 (4): 351–371.
Rasmussen AR, Sanders K, Lobo A, Courtney T. 2018a. Hydrophis schistosus (amend-
ed version of 2010 assessment). The IUCN Red List of Threatened Species 2018:
e.T176719A136258180. [Link]
[Link].
Rasmussen AR, Sanders K, Lobo A. 2010b. Kolpophis annandalei. The IUCN Red List
of Threatened Species 2010: e.T176731A7292396. [Link]
[Link].
Rasmussen AR, Sanders K, Lobo A. 2018b. Hydrophis viperinus (amended version of 2010
assessment). The IUCN Red List of Threatened Species 2018: e.T176711A136259318.
[Link] [Link].
Rasmussen AR. 1989. An analysis of Hydrophis ornatus (Gray), H. lamberti Smith, and H.
inornatus (Gray) (Hydrophiidae, Serpentes) based on samples from various localities,
with remarks on feeding and breeding biology of H. ornatus. Amphibia-Reptilia. 10:397-
417.
Rasmussen AR. 1992. Rediscovery and Redescription of Hydrophis bituberculatus Peters,
1872 (Serpentes: Hydrophidae). Herpetologica, vol. 48, no. 1, pp. 85–97. JSTOR, www.
[Link]/stable/3892922.
681
Aquatic Snakes

Rasmussen AR. 2010. Hydrophis inornatus. The IUCN Red List of Threatened Species
2010: e.T176732A7292599. [Link]
[Link].
Rasmussen AR. 2010b. Hydrophis laboutei. The IUCN Red List of Threatened Species
2010: e.T176758A7298709. [Link]
[Link].
Rasmussen AR. 2010c. Hydrophis melanocephalus. The IUCN Red List of Threatened Spe-
cies 2010: e.T176762A7299369. [Link]
[Link].
Rasmussen AR. 2018. Hydrophis zweifeli (amended version of 2010 assessment). The IUCN
Red List of Threatened Species 2018: e.T176716A134229775. [Link]
[Link]. [Link]
Rasmussen AR. Elmberg J, Gravlund P, Ineich I. 2011b. Sea snakes (Serpentes: subfamilies
Hydrophiinae and Laticaudinae) in Vietnam: a comprehensive checklist and an updated
identification key. Zootaxa 2894: 1–20.
Rasmussen JB. 2004. A review of Whyte’s Water‐snake, Lycodonomorphus whytii (Ser-
pentes: Colubridae: Lamprophiinae). African Journal of Herpetology 53(2):155-62.
Ratnikov V, Mebert K. 2011. Fossil remains of Natrix tessellata from the late cenozoic de-
posits of the east European plain. Mertensiella 18:337-43.
Raudzens E. 2007. At sea testing of The Popeye Fishbox bycatch reduction device onboard
the FV Adelaide Pearl for approval in Australia’s Northern Prawn Fishery. Australian
Fisheries Management Authority, Canberra..
Rautsaw RM, Schramer TS, Acuña R, Arick LN, DiMeo M, Mercier KP, Schrum M, Mason
AJ, Margres MJ, Strickland JL, Parkinson CL. 2021. Genomic Adaptations to Salinity
Resist Gene Flow in the Evolution of Floridian Watersnakes. Molecular Biology and
Evolution 38(3):745–760. [Link]
Raw LR.1973. A review of the dusky-bellied water snake, Lycodonomorphus laevissimus
(Gunther), with descriptions o two new subspecies. Annals of the Natal Museum. 1973
Sep 1;21(3):713-8.
Raw LR, Deacon R. 2021. Two additional specimens of Naja (Boulengerina) nana (Ser-
pentes: Elapidae) with notes on captive husbandry and behaviour. Occasional Papers
in Zoology 9: 1–8.
Ray JM. 2020. Snakes of Panama, a field guide to all species. The Snake Team, Panama.
Reading CJ, Davies JL. 1996. Predation by grass snakes (Natrix natrix) at a site in southern
England. Journal of Zoology 239: 73–82.
Rebelato MM, Ferri VY, Dalmolin DA, Tozetti AM, Verrastro L. 2021. Envenomation by
opisthoglyphous snake Thamnodynastes hypoconia (Cope, 1860) (Dipsadinae: Tachyme-
nini) in southern Brazil. Toxicon 189:1-6.
Reed RN, Rodda GH. 2009. Giant constrictors: biological and management profiles and
an establishment risk assessment for nine large species of pythons, anacondas, and the
boa constrictor: U.S. Geological Survey Open-File Report 2009-1202, U.S. Geological
Survey, Reston, Virginia, 2009, xviii + 302 pp
Remruatpui L, Lalremsanga HT. 2020. Smithophis atemporalis (mizo rain snake). Reproduc-
tion and elevation. Herpetological Review 51: 156-157.
Ren JL, Wang K, Nguyen TT, Hoang VC, Zhong GH, Jiang K, Guo PE, Li JT. 2018. Tax-

682
Aquatic Snakes

onomic re-evaluation of the monotypic genus Pararhabdophis Bourret, 1934 (Squa-

mata: Colubridae: Natricinae) with discovery of its type species, P. chapaensis, from
China. Zootaxa 4486(1):031-56.
Ren JL, Yan C, Peng ZL, Li JT. 2022. Sichuan hot-spring snakes imperiled: reason, situa-
tion, and protection. Zoological research 43(1):95.
Reuss A. 1834. Zoologische Miscellen. Reptilien, Ophidier. [Coluber albiventris, Echis
pavo]. Mus. Senckenbergiana, Frankfurt/M., 1:129–162.
Reynolds RG, Henderson RW. 2018. Boas of the world (Superfamily Booidae): a check-
list with systematic, taxonomic, and conservation assessments. Bulletin of the Muse-
um of Comparative Zoology, 162: 1–58.
Reynolds RG, Niemiller ML, Revell LJ. 2014. Toward a Tree-of-Life for the boas and py-
thons: Multilocus species-level phylogeny with unprecedented taxon sampling. Molec-
ular Phylogenetics and Evolution 71:201-13.
Rezaie-Atagholipour M, Ghezellou P, Hesni MA, Dakhteh SMH, Ahmadian H, Vidal N
2016. Sea snakes (Elapidae, Hydrophiinae) in their westernmost extent: an updated
and illustrated checklist and key to the species in the Persian Gulf and Gulf of Oman.
ZooKeys 622: 129–164, doi: 10.3897/zookeys.622.9939
Rezaie-Atagholipour M, Riyahi-Bakhtiari A, Sajjadi M. 2013. Feeding Habits of the Annu-
lated Sea Snake, Hydrophis cyanocinctus, in the Persian Gulf. Journal of Herpetology,
47(2):328–330. doi:10.1670/11-150
Rezaie-Atagholipour, M. 2012. Lapemis curtus (short sea snake) diet. Herpetological Re-
view 43: 494.
Rezaie-Atagholipour, M., Riyahi-Bakhtiari, A., & Sajjadi, M. 2013. Feeding Habits of the
Annulated Sea Snake, Hydrophis cyanocinctus, in the Persian Gulf. Journal of Herpetol-
ogy, 47(2), 328–330. doi:10.1670/11-150
Rhoda D, Polly PD, Raxworthy C & Segall M. 2020. Morphological integration and
modularity in the hyperkinetic feeding system of aquatic foraging snakes. Evolution
75(1):56–72.
Richards SJ, Suryadi S. 2002. A biodiversity assessment of Yongsu-Cyclops mountains and
the southern Mamberamo basin, Papua, Indonesia. RAP Bulletin of Biological Assess-
ment No. 25. Conservation International, Washington, DC, 189 pp.
Rivas JA, Burghardt GM. 2001. Understanding sexual size dimorphism in snakes: wear-
ing the snake’s shoes. Animal Behaviour. 62 (3): F1–F6. doi:10.1006/anbe.2001.1755.
Rivas JA, Burghardt GM. 2005. Snake mating systems, behavior, and evolution: The revi-
sionary implications of recent findings. Journal of Comparative Psychology. 119 (4):
447–454. doi:10.1037/0735-7036.119.4.447.
Rivas JA, Molina CR, Corey SJ, Burghardt GM. 2016. Natural history of neonatal green an-
acondas (Eunectes murinus): a chip off the old block. Copeia. 2016 104(2):402-10.
Rivas JA, Muñoz MC, Burghardt GM, Thorbjarnarson JB. 2007. Sexual size dimorphism
and the mating system of the green anaconda (Eunectes murinus), p.461–473. In: Bi-
ology of Boas, Pythons, and Related Taxa. RW Henderson and R Powell (Eds.). Eagle
Mountain Publishing, LC, Utah.
Rivas JA. 1999. The life history of the green anaconda (Eunectes murinus), with emphasis
on its reproductive biology. The University of Tennessee.
Rivero-Blanco C, Dixon JR. 1979. Origin and distribution of the herpetofauna of the dry
683
Aquatic Snakes

lowland regions of northern South America Pages 281-298 In W. E. Duellman (ed) The
South American Herpetofauna its origin, evolution, and dispersal. Monograph of the
Museum of Natural History, The University of Kansas (Number 7).
Roe JH, Kingsbury BA, Herbert NR. 2003. Wetland and upland use patterns in
semi-aquatic snakes: implications for wetland conservation. Wetlands 23, 1003–1014.
Rorabaugh JC, Turner D, van Devender TR, Hugo-Cabrera V, Maynard RJ, Van Devender
RW, Villa RA, Hamilton P, Hale SF, Aguilar-Morales C, Blanco-Gutiérrez A. 2019.
Herpetofauna of the Mesa Tres Ríos area in the Northern Sierra Madre Occidental of
Sonora, Mexico. Herpetological Review, 50(2), 251-259.
Rosa, G.M., Bergò, P.E., Andreone, F., Crottini, A., Jasper, L., Gardner, C.J. 2016. Insights
into the feeding behaviour of Madagascarophis meridionalis Domergue, 1987, from
Isalo, Madagascar. Herpetozoa 29: 103–107.
Rosen PC and Hollycross A. 2011. The Narrow-Headed Garter Snake in the United States
(Thamnophis rufipunctatus): a report on its status, natural history, and threats. Center
for Biological Diversity 28 pages.
Rosen PC, Schwalbe CR. 1988. Status of the Mexican and narrow-headed garter snakes
(Thamnophis eques megalops and Thamnophis rufipunctatus) in Arizona. Arizona
Game and Fish Department, unpublished report submitted to US Fish and Wildlife
Service, Albuquerque, New Mexico. 1988
Ross RA, Marzec G. 1990. The reproductive husbandry of pythons and boas. Privately
Published.
Rossi JV, Rossi R. 1995. Snakes of the United States and Canada: keeping them healthy in
captivity. Volume 2. Western Area. Krieger Publishing Company.
Rossman DA. 1966. Evidence for conspecificity of the Mexican Garter Snakes Thamnophis
phenax (Cope) and Thamnophis sumichrasti (Cope). Herpetologica, 22(4): 303–305.
Rossman DA. 1975. Redescription of the South American Colubrid Snake Helicops hag-
manni Roux. Herpetologica 31(4): 414-418.
Rossman DA, 1970. Thamnophis proximus. Catalogue of American Amphibians and Rep-
tiles (CAAR). 1970.
Rossman DA, 1971. Systemics of the neotropical populations of Thamnophis marcianus
(Serpentes: Colubridae). Occasional Papers of the Museum of Natural Science, Louisi-
ana State University (41) Article 1. DOI: 10.31390/opmns.041.
Rossman DA, Burbrink FT. (2005) Species limits within the Mexican garter snakes of the
Thamnophis goldmani complex. Occasional Papers of the Museum of Natural Science,
Louisiana State University : No. 79, Article 1. DOI: 10.31390/opmns.079
Rossman DA, Eberle WG. 1977. Partition of the genus Natrix, with preliminary observa-
tions on evolutionary trends in natricine snakes. Herpetologica 33:34-43.
Rossman DA, Ford NB, Seigel RA. 1996. The Garter Snakes: Evolution and Ecology (Ani-
mal Nat). University of Oklahoma Press.
Rossman DA and Powell R, 1985. Clonophis, C. kirtlandii. Catalogue of American Am-
phibians and Reptiles (CAAR).
Rossman DR, Wallach V. 1987. Adelophis Duges, Mountain Meadow Snakes. Catalogue of
American Amphibians and Reptiles 408.1-2.
Roux J. 1910. Reptilien und amphibian der Aru-und Kei Inseln. Abhandlungen der Senck-
enbergischen Naturforschenden Gesellschaft 33:211–247.
684
Aquatic Snakes

Rugiero L, Capula M, Persichetti D, Luiselli L, Angelici FM. 2000. Life-history and diet of
two populations of Natrix maura (Reptilia, Colubridae) from contrasted habitats in
Sardinia. Miscelania Zoologica 23:41-51.
Ruiz-Garcia JA, Romero VCAL, Natalini MB. 2020. Helicops leopardinus (Leopard Keel-
back) and Philodryas bolfersii latirostris (Lichternstein’s Green Racer). Predation. Her-
petological Review 51(1)145-146.
Ryerson W, Schwenk K. 2014. Why Snakes Flick Their Tongues: A Fluid Dynamics Ap-
proach. In University of Connecticut.
Sagar U. 2018. Some observations on the Malabar Pit Viper Trimeresurus malabaricus in
central Western Ghats, India. Newsletter of the South Asian Reptile Network 18:36.
Saha BK. 1984. More on the dog-face. Hamadryad 8(2):4–5.
Said-Aliev SA. 1979. Amphibians and Reptiles of Tajikistan. Donish, Dushanbe, Tajikistan,
Academy of Sciences of the Tajikistan SSR. 145 pp. [in Russian].
Saint Girons H. 1956. Les serpents du Maroc. Variétés scientifiques recueillies par la So-
ciété des sciences naturelles et physiques du Maroc 8 :1-29.
Saint Girons H. 1972. Les serpents du Cambodge. Mémoires du Muséum National d’His-
toire Naturelle, Serie A 74:1–170.
Saint Girons H, Pfeffer P. 1972. Notes sur l’ecologie des serpents du Cambodge. Zoolo-
gische Mededelingen 47:65–86. [English Summary].
Sala, O. E., Chapin, F. S., Armesto, J. J., Berlow, R., Bloomfield, J., Dirzo, R., Huber-San-
wald, E., Huenneke, L. F., Jackson, R. B., Kinzig, A., Leemans, R., Lodge, D., Mooney,
H. A., Oesterheld, M., Poff, N. L., Sykes, M. T., Walker, B. H., Walker, M. & Wall, D. H.
(2000). Global biodiversity scenarios for the year 2100. Science 287, 1770–1774
Salas-Gismondi R, Flynn JJ, Baby P, Tejada-Lara JV, Wesselingh FP, Antoine PO. 2015.
A Miocene hyperdiverse crocodylian community reveals peculiar trophic dynamics
in proto-Amazonian mega-wetlands. Proceedings of the Royal Society of London B:
Biological Sciences, 282(1804), 20142490
Sanchiz B. 1991. Algunas herpetofaunas de yacimientos del Pleistoceno medio ibérico. Re-
vista Española de Herpetología 5:9-13.
Sanders KL. 2010b. Hydrophis melanosoma. The IUCN Red List of Threatened Species
2010: e.T176759A7298829. [Link]
[Link].
Sanders KL. 2010b. Hydrophis melanosoma. The IUCN Red List of Threatened Species
2010: e.T176759A7298829. [Link]
[Link].
Sanders KL, Lee MS, Leys R, Foster R, Scott Keogh J. 2008. Molecular phylogeny and
divergence dates for Australasian elapids and sea snakes (Hydrophiinae): evidence
from seven genes for rapid evolutionary radiations. Journal of Evolutionary Biology
21(3):682-95.
Sanders, K., Mumpuni, Hamidy, A., Head, J. J., & Gower, D. J. 2010. Phylogeny and diver-
gence times of filesnakes (Acrochordus): Inferences from morphology, fossils and three
molecular loci. Molecular Phylogenetics and Evolution, 56(3), 857-867. doi:10.1016/j.
ympev. 2010.04.031
Sanders KL, Rasmussen A. 2010. Hydrophis klossi. The IUCN Red List of Threatened Spe-
cies 2010: e.T176753A7297716. [Link]

685
Aquatic Snakes

[Link].
Sanders KL, Rasmussen AR, Elmberg J, Mumpuni, Guinea M, Blias P, Lee MSY & Fry BG.
2012. Aipysurus mosaicus, a new species of egg-eating sea snake (Elapidae: Hydrophi-
inae), with a redescription of Aipysurus eydouxii (Gray, 1849). Zootaxa 3431: 1–18.
Sanders KL, Rasmussen AR, Elmberg J. 2012. Independent innovation in the evolution of
paddle-shaped tails in viviparous sea snakes (Elapidae: Hydrophiinae). Integrative and
Comparative Biology, 52: 311–320,
Sanders KL, Schroeder T, Guinea ML, Rasmussen AR. 2015. Molecules and Morphology
Reveal Overlooked Populations of Two Presumed Extinct Australian Sea Snakes (Aipy-
surus: Hydrophiinae). PLOS ONE, 10(2), e0115679. doi:10.1371/[Link].0115679.
Sanders KL, White M-D, Courtney T. Lukoschek V. 2018. Hydrophis stokesii (amended
version of 2010 assessment). The IUCN Red List of Threatened Species 2018:
e.T176708A136257093. [Link]
[Link]
Sandfoss MR, Sheehy III CM, Lillywhite HB. 2018. The collapse of a unique insular bird–
snake relationship. Journal of Zoology 304(4):276-83.
Sanmartín I, Enghoff H, Ronquist F. 2001. Patterns of animal dispersal, vicariance and di-
versification in the Holarctic. Biological Journal of the Linnean Society 73(4):345-90.
Santos X. 2000. Ecología de la culebra viperina, Natrix maura (Linnaeus, 1758), en el Delta
del Ebro. Tesis Doctoral. Universitat de Barcelona.
Santos X. 2008. Natrix maura (Linnaeus, 1758). Pp. 176-177. In: Loureiro A, N Ferrand de
Almeida, MA Carretero, OS Paulo (Eds). Atlas dos Anfíbios e Répteis de Portugal. Insti-
tuto da Conservaçao da Natureza e da Biodiversidade, Lisboa. 257 pp.
Santos X. 2014. Natrix maura (Linnaeus, 1758. In: Reptiles. Salvador, A. (Coordinador).
Fauna Ibérica. vol. 10. Ramos MA et al. (Eds). Segunda edición. Museo Nacional de
Ciencias Naturales. CSIC. Madrid.
Santos X. 2015. Culebra viperina - Natrix maura. En: Enciclopedia Virtual de los Verteb-
rados Españoles. Salvador, A, A Marco (Eds). Museo Nacional de Ciencias Naturales,
Madrid. [Link] [Link]/
Santos X, Llorente GA. 2001b. Seasonal Variation in Reproductive Traits of the Oviparous
Water Snake, Natrix maura, in the Ebro Delta of Northeastern Spain. Journal of Herpe-
tology 35:653-660.
Santos X, C Arenas, GA Llorente, X Ruiz. 2007. Exploring the origin of egg protein in an
oviparous water snake (Natrix maura). Comparative Biochemistry and Physiology Part
A: Molecular & Integrative Physiology 147(1):165-172.
Santos X, Azor JS, Cortés S, Rodríguez E, Larios J, Pleguezuelos JM. 2018. Ecological signif-
icance of dorsal polymorphism in a Batesian mimic snake. Current Zoology, 64 (6):745–
753, [Link]
Santos X, Feriche M, León R, Filippakopoulou A, Vidal-García M, Llorente GA, Pleguezue-
los JM. 2011. Tail breakage frequency as an indicator of predation risk for the aquatic
snake Natrix maura. Amphibia-Reptilia 32:375-383.
Santos X. 2008. Natrix natrix Linnaeus, (1758) Cobra-de-agua-de-colar. Pp. 178–179. In:
Loureiro A, Ferrand de Almeida N, Carretero MA, Paulo O (Eds.). Atlas dos Anfíbios e
Répteis de Portugal. Instituto da Conservação da Natureza e da Biodiversidade, Lisboa.
Santos X, Llorente GA, Montori A, Carretero MA. 2002. Natrix natrix Linnaeus (1758) Cul-
686
Aquatic Snakes

ebra de collar. Pp. 293–295. in: Pleguezuelos JM, Márquez R, Lizana M. (Eds.). Atlas y
Libro Rojo de los Anfibios y Reptiles de España. Ministerio de Medio Ambiente, Madrid.
Santos X, García-Cardenete L. 2006. Introducción de peces en ríos de la cuenca mediter-
ránea: una amenaza para sus depredadores. Boletín de la Asociación Herpetológica
Española 16(1-2):50-51.
Santos X, González-Solís J, Llorente GA. 2000. Variation in the diet of the Viperine Snake,
Natrix maura, in relation to prey availability. Ecography 23:185-192.
Santos X, Llorente GA. 1997. Actividad de Natrix maura en el Delta del Ebro, analizada
mediante técnicas de telemetría. Revista Española de Herpetología 11:63-70.
Santos X, Llorente GA. 1998. Sexual and size-related differences in the diet of the snake
Natrix maura from the Ebro Delta, Spain. Herpetolgical Journal 8:161-165.
Santos X, Llorente GA. 2001a. Actividad estacional y diaria en una población de culebra
viperina, Natrix maura, del Delta del Ebro. Boletín de la Asociación Herpetológica Es-
pañola 12:22-24.
Santos X, Llorente GA. 2004. Lipid dynamics in the viperine snake Natrix maura from the
Ebro Delta. Oikos 105:132-140.
Santos X, Llorente GA. 2009. Decline of a common reptile: case study of the viperine snake
Natrix maura in a Mediterranean wetland. Acta Herpetologica 4:161-169.
Santos X, Llorente GA. 2009. Decline of a common reptile: case study of the viperine snake
Natrix maura in a Mediterranean wetland. Acta Herpetologica 4:161–169.
Santos X, Llorente GA, Feriche M, Pleguezuelo JM, Casals F, de Sostoa A. 2005. Food
availability induces geographic variation in reproductive timing of an aquatic ovipa-
rous snake (Natrix maura). Amphibia-Reptilia 26(2):183-91.
Santos X, Llorente GA, Montori A, Carretero MA. 1997. Natrix maura (Linnaeus, 1758)
Culebra viperina. In: Pleguezuelos JM. (Ed.. Distribución y biogeografía de los anfibios
y reptiles en España y Portugal. Monografías de la Revista Española de Herpetología
3:279-281.
Santos X, Llorente GA, Montori A, Carretero MA. 2002. Natrix maura (Linnaeus, 1758.
Pp. 289-291. In: Pleguezuelos JM, R Márquez, M Lizana (Eds). Atlas y libro rojo de los
anfibios y reptiles de España. Dirección General de Conservación de la Naturaleza,
Asociación Herpetológica Española, Madrid.
Santos X, Pastor D, Llorente GA, Albaigés J. 1999. Organochlorine levels in viperine snake
Natrix maura carcasses from the Ebro Delta (NE Spain): sexual and size-related differ-
ences. Chemosphere 39:2641-2650.
Santos X, Pleguezuelos JM, Chergui B, Geniez P, Cheylan M. 2022. Citizen-science data
shows long-term decline of snakes in southwestern Europe. Biodiversity and Conser-
vation 31(5-6):1609–1625.
Santos X, Vacher J-P. 2010. La couleuvre vipérine. Natrix maura (Linnaeus, 1758). Les
Reptiles de France, Belgique, Luxembourg et Suisse. Pp: 456-463. Biotope Éditions.
Paris, France.
Santos X, Vilardebo E, Casals F, Llorente GA, Vinyoles D, de Sostoa A. 2006. Wide food
availability favours intraspecific trophic segregation in predators: the case of a water
snake in a Mediterranean river. Animal Biology 56:299-309.
Sanz JL, FB Sanchiz. 1980. Nota preliminar sobre los Squamata (Reptilia) del Pleistoceno
Medio de Áridos-I (Arganda, Madrid. In: Ocupaciones Achelenses en el valle del Jara-
687
Aquatic Snakes

ma, Santonja, M. Diputación de Madrid, Madrid: 127-138.

Sauvage HE. 1879. Sur un Rhabdosomien de genre nouveau provenant de Ternate. Bulletin
de la Sociéte Philomathique de Paris (7)3:61–63.
Savage JM. 2002. The amphibians and reptiles of Costa Rica: a herpetofauna between
two continents, between two seas. University of Chicago Press.
Savitzky AH. 1983. Coadapted character complexes among snakes: fossoriality, piscivory,
and durophagy. American Zoologist 23(2):397–409.
Savitzky BA, Burghardt GM. 2000. Ontogeny of Predatory Behavior in the Aquatic Spe-
cialist Snake, Nerodia rhombifer, during the first year of life. Herpetological Mono-
graphs 2000 1:401-19.
Sazima I, Strüssmann C. 1990. Necrofagia em serpentes brasileiras: exemplos e pre-
visões. Revista Brasileira de Biologia 50(2): 463–468.
Said-Aliev SA. 1979. Amphibians and Reptiles of Tajikistan. Donish, Dushanbe, Tajikistan,
Academy of Sciences of the Tajikistan SSR. 145 pp. [in Russian].
Scali S. 2011. Ecological comparison of the Dice snake (Natrix tessellata) and the Viperine
snake (Natrix maura) in northern Italy. Mertensiella 18:131–145.
Scali S. 2011. Ecological comparison of the Dice snake (Natrix tessellata) and the Viperine
snake (Natrix maura) in northern Italy. Mertensiella 18:131-45.
Scali S., Gentilli A., Lanza B., 2011. Natrix natrix. In Corti C., Capula M., Luiselli L., Sin-
daco R., Razzetti E., 2011. Fauna d’Italia, vol. XLV, Reptilia. Calderini, Bologna, XII +
869 pp.
Scartozzoni RR, Marques OA. 2004. Sexual dimorphism, reproductive cycle, and fecundity
of the water snake Ptychophis flavovirgatus (Serpentes, Colubridae). Phyllomedusa:
Journal of Herpetology 3(1):69-71.
Schaeffel F, de Queiroz A. 1990. Alternative mechanisms of enhanced underwater vision in
the garter snakes Thamnophis melanogaster and T. couchii. Copeia 1990(1):50–58.
Schaeffel F, de Queiroz A. 1990. Alternative Mechanisms of Enhanced Underwater Vision
in the Garter Snakes Thamnophis melanogaster and T. couchii. Copeia, 1990(1):50–58.
Schaeffel F, Mathis U. 1991. Underwater vision in semi-aquatic European snakes. Natur-
wissenschaften, 78:373–375.
Schargel WE, Rivas Fuenmayor G, Barros TR, Péfaur JE, Navarrete LF. 2007. A new aquat-
ic snake (Colubridae: Pseudoeryx) from the lake Maracaibo basin, northwestern Vene-
zuela: a relic of the past course of the Orinoco River. Herpetologica 63(2):236-44.
Schätti B, McCarthy C. 2004. Saharo-Arabian racers of the Platyceps rhodorhachis com-
plex - description of a new species (Reptilia: Squamata: Colubrinae). Revue Suisse de
Zoologie 111:691-705
Schätti B. 1982. Bemerkungen zur ökologie, verbreitung und intraspezifischen variation
der vipernatter, Natrix maura (Linné, 1758) (Reptilia, Serpentes. Revue Suisse de Zool-
ogie 89:521-542.
Schätti B. 1999. Natrix maura (Linnaeus, 1758) – Vipernatter. In: Böhme, W. (Ed.): Hand-
buch der Reptilien und Amphibien Europas 3/Schlangen II. AULA-Verlag, Wiesbaden,
Germany: 483–503.
Schlegel H. 1837. Essai sur la Physionomie des serpens, Vol. 1 and 2. M. H. Schonekat,
Amsterdam.
Schleich HH, W Kästle, K Kabisch. 1996. Amphibians and Reptiles of North Africa. Koeltz
688
Aquatic Snakes

Science Books, Koenigstein.

Schmidt KP. 1923. Contributions to the herpetology of the Belgian Congo based on the
collection of the American Museum Congo Expedition, 1909-1915. Part II. Snakes.
With field notes by Herbert Lang and James P. Chapini. Bulletin of the American Mu-
seum of Natural History XLIX: 1- 146.
Schneider JG. 1799. Historiae amphibiorum naturalis et literariae. Fasciculus primus, con-
tinens Ranas, Clamitas, Bufones, Salamandras et Hydros in genera et species descriptos
notisque suis distinctos. Friederici Frommann, Jenae, 264 pp.
Schöneberg Y, Köhler G. 2022. Distribution and identification of the species in the genus
Helicops Wagler, 1830 (Serpentes, Colubridae, Xenodontinae). ARPHA Preprints.
1:e67869.
Schöneberg Y, Winter S, Arribas O, Di Nicola MR, Master M, Owens JB, Rovatsos M,
Wüster W, Janke A, Fritz U. 2023. Genomics reveals broad hybridization in deeply di-
vergent Palearctic grass and water snakes (Natrix spp.). Molecular Phylogenetics and
Evolution 184 (2023) 107787.
Schultze N, Spitzweg C, Corti C, Delaugerre M, Di Nicola MR, Geniez P, Lapini L, Liuzzi
C, Lunghi E, Novarini N, Picariello O, Razzetti E, Sperone E, Stellati L, Vignoli L, Asz-
talos M, Kindler C, Vamberger M, Fritz U. 2020. Mitochondrial ghost lineages blur
phylogeography and taxonomy of Natrix helvetica and N. natrix in Italy and Corsica.
Zoologica Scripta, 49: 395-411.
Scortecci G. 1932. Rettili dello Yemen. Atti della Società Italiana di Scienze Naturali e del
Museo Civico di Storia Naturale di Milano 71:39-49.
Scott DA. A directory of Asian wetlands. IUCN, The World Conservation Union; 1989.
Scott NJ, Maxwell TC, Thornton Jr OW, Fitzgerald LA, Flury JW. 1989. Distribution, habi-
tat, and future of Harter’s water snake, Nerodia harteri, in Texas. Journal of Herpetology.
1989:373-89.
Scrocchi GJ, Ferreira VL, Giraudo AR, Ávila RW, Motte M. A new species of Hydrops
(Serpentes: Colubridae: Hydropsini) from Argentina, Brazil and Paraguay. Herpeto-
logica. 2005 Dec;61(4):468-77.
Schweizer R. 1911. Schreckstellung der Ringelnatter. Lacerta:21–22.
Secoy DM. 1976 Distribution and population of Saskatchewan’s amphibians and reptiles.
Report to the Saskatchewan Department of the Environment, Regina.
Segall M, Cornette R, Fabre A-C, Godoy-Diana R & Herrel A. 2016. Does aquatic foraging
impact head shape evolution in snakes? Proceedings of the Royal Society of London, B
Biological Sciences 283:20161645.
Segall M, Cornette R, Godoy-Diana R & Herrel A. 2020. Exploring the functional meaning
of head shape disparity in aquatic snakes. Ecology and Evolution 10(14):6993–7005.
Segall M, Cornette R, Rasmussen AR & Raxworthy C. 2021. Inside the head of snakes:
influence of size, phylogeny, and sensory ecology on endocranium morphology. Brain
Structure and Function. 1:15.
Segall M, Herrel A & Godoy-Diana R. 2019. Hydrodynamics of frontal striking in aquatic
snakes: drag, added mass, and the possible consequences for prey capture success. Bio-
inspiration & Biomimetics 14(036005):1–9.

689
Aquatic Snakes

Segura C, Feriche M, Pleguezuelos JM, Santos X. 2007. Specialist and generalist species for
habitat use: implications for conservation assessment in snakes. Journal of Natural His-
tory 41:2765–2774.
Seigel RA, JW Gibbons, TK Lynch. 1995. Temporal Changes in Reptile Populations: Ef-
fects of a Severe Drought on Aquatic Snakes. Herpetologica 51: 424-34.
Semlitsch RD, Pechmann JHK, Gibbons JW. 1988. Annual Emergence of Juvenile Mud
Snakes (Farancia abacura) at Aquatic habitats. Copeia, 1988(1): 243–245.
Senter PJ, Senter JM. 2020. Nerodia sipedon (Northern Watersnake). Courtship and Mating.
Herpetological Review 51(4):2020.
Seoane [ López Seoane] V. 1884. Lacerta viridis, var. Gadovii (Boulenger) é investigaciones
herpetológicas de Galicia. La Coruña, Vicente Abad, 19 pp.
Sergo D, Shrine R. 2015. Snakes for lunch: bird predation on reptiles in a tropical floodplain.
Australian Zoologist 37(3):311–320.
Sergo D, Shrine R. 2015. Snakes for lunch: bird predation on reptiles in a tropical floodplain.
Australian Zoologist 37(3):311–320.
Setiadi MI, Hamidi A. 2006. Jenis-jenis herpetofauna di pulau Halmahera. Report Universi-
tas Indonesia, Depok and Museum Zoologicum Bogoriense, Cibinong.
Sexton OJ, Drda WJ, Sexton KG, Bramble JE. 2007. The effects of flooding upon the snake
fauna of an isolated refuge. Natural Areas Journal 27(2):133-45.
Sharma RC. 2003. Handbook – Indian Snakes. Director- Zoological Survey of India, Kolk-
ata, 292 pp.
Shehab AH, Al Masri A, Amr ZS. 2011. The Dice Snake, Natrix tessellata in Syria: distribu-
tion, trade and conservation. Mertensiella, 18:388-92.
Sherratt E, Coutts FJ, Rasmussen AR, Sanders KL. 2019. Vertebral Evolution and Ontoge-
netic Allometry: The Developmental Basis of Extreme Body Shape Divergence in Micro-
cephalic Sea Snakes. Evolution & Development, no. 2019: 1–10. [Link]
ede.12284.
Sherratt E, Rasmussen AR, Sanders KL. 2018. Trophic specialization drives morpholog-
ical evolution in sea snakes. Royal Society Open Science, 5(3), 172141. doi:10.1098/
rsos.172141
doi:10.1098/rsos.172141
Shetty S, Shine R. 2002a. Philopatry and Homing Behavior of Sea Snakes (Laticauda col-
ubrina) from Two Adjacent Islands in Fiji. Conservation Biology, 16(5), 1422–1426.
doi:10. 1046/j. 1523-1739.2002.00515.x.
Shetty S, Shine R. 2002b. Sexual divergence in diets and morphology in Fijian sea snakes
Laticauda colubrina (Laticaudinae). Austral Ecology, 27(1), 77–84. doi:10.1046/j.1442-
9993.2002.01161.x
Shetty S, Shine R. 2002c. The Mating System of Yellow-Lipped Sea Kraits (Laticauda colubri-
na: Laticaudidae). Herpetologica, 58(2):170–180. doi:10.1655/0018-0831(2002)058[0170
:tmsoys][Link];2
Shibata H, Shuichi S; Hirano Y; Nitasaka E, Sakabe A. 2017. Facultative parthenogenesis
validated by DNA analyses in the green anaconda (Eunectes murinus). PLoS ONE. 12
(12): e0189654. doi:10.1371/[Link].0189654. PMC 5728508. PMID 29236745.
Shine R, Bonnet X, Elphick MJ & Barrott EG. 2004. A novel foraging mode in snakes:
browsing by the sea snake Emydocephalus annulatus (Serpentes, Hydrophiidae). Func-
690
Aquatic Snakes

tional Ecology 18(1):16–24.

Shine R, Brown GP, Elphick MJ. 2004. Field experiments on foraging in free-ranging water
snakes Enhydris polylepis (Homalopsinae). Animal Behaviour 68(6):1313–1324.
Shine R, Cogger HG, Reed RR, Shetty S, Bonnet X. 2003. Aquatic and terrestrial loco-
motor speeds of amphibious sea-snakes (Serpentes, Laticaudidae). Journal of Zoolo-
gy 259(3):261-8.
Shine R, Goiran C, Shilton C, Meiri S, Brown GP. The life aquatic: an association be-
tween habitat type and skin thickness in snakes. Biological Journal of the Linnean
Society. 2019 Dec 6;128(4):975-86.
Shine R, Grigg GC, Shine TG, Harlow P. 1981. Mating and male combat in Australian
blacksnakes, Pseudechis porphyriacus. Journal of Herpetology 15:101-7.
Shine R, Harlow P, Keogh JS, Boeadi. 1995. Biology and commercial utilization of acro-
chordid snakes, with special reference to Karung (Acrochordus javanicus). Journal of
Herpetology 1:352-60.
Shine R, Houston D. 1993. Family Acrochordidae. In Fauna of Australia, 322–24.
Shine R, Keogh JS. 1996. Food habits and reproductive biology of the endemic Melanesian
elapids: are tropical snakes really different? Journal of Herpetology 30:238-47.
Shine R, Lambeck R. A 1985. radiotelemetric study of movements, thermoregulation
and habitat utilization of Arafura filesnakes (Serpentes: Acrochordidae). Herpetolog-
ica. 1985 Sep 1:351-61.
Shine R, Madsen T. 1996. Is thermoregulation unimportant for most reptiles? An exam-
ple using water pythons (Liasis fuscus) in tropical Australia. Physiological Zoology
69(2):252-69.
Shine R, Madsen T. 1997. Prey abundance and predator reproduction: rats and pythons
on a tropical Australian floodplain. Ecology. 78(4):1078-86.
Shine R, Madsen TR, Elphick MJ, Harlow PS. 1997. The influence of nest tempera-
tures and maternal brooding on hatchling phenotypes in water pythons. Ecology.
78(6):1713-21.
Shine R, Olsson MM, Moore IT, LeMaster MP, Mason RT. 1999. Why do male snakes
have longer tails than females? Proceedings of the Royal Society of London. Series B:
Biological Sciences. 266(1434):2147-51.
Shine R, Schwaner T. 1985. Prey constriction by venomous snakes: a review, and new data
on Australian species. Copeia 1985(4):1067–1071.
Shine R, Shetty S. 2001. Moving in two worlds: aquatic and terrestrial locomotion in sea
snakes (Laticauda colubrina, Laticaudidae). Journal of Evolutionary Biology, 14(2),
338–346. doi:10.1046/j.1420-9101.2001.00265.x
Shine R, Wall M. 2008. Interactions between locomotion, feeding, and bodily elongation
during the evolution of snakes. Biological Journal of the Linnean Society. 95(2):293-
304.
Shine R. 1977. Reproduction in Australian elapid snakes II. Female reproductive cycles.
Australian Journal of Zoology 25(4):655-66.
Shine R. 1986. Ecology of a low-energy specialist: food habits and reproductive biology of
the Arafura filesnake (Acrochordidae). Copeia. 1986:424-37.
Shine R. 1987b. Intraspecific variation in thermoregulation, movements and habitat use
by Australian blacksnakes, Pseudechis porphyriacus (Elapidae). Journal of Herpetology
691
Aquatic Snakes

21:165-77.
Shine R. 1988. Constraints on reproductive investment: a comparison between aquatic
and terrestrial snakes. Evolution. 1988 Jan;42(1):17-27.
Shine R. 1991. Strangers in a strange land: ecology of the Australian colubrid snakes. Co-
peia 1991(1):120–131.
Shine R. 2005. All at sea: Aquatic life modifies mate-recognition modalities in sea snakes
(Emydocephalus annulatus, Hydrophiidae). Behavioral Ecology and Sociobiology
57(6):591–598.
Shine R. Brischoux F, Pile AJ. 2010. A seasnake’s colour affects its susceptibility to algal
fouling. Proceedings of the Royal Society B: Biological Sciences, 2010 DOI: 10.1098/
rspb.2010.0255.
Shine, R., 1987a. Food habits and reproductive biology of Australian snakes of the genus
Hemiaspis (Elapidae). Journal of Herpetology, 21(1), pp.71-74.
Shiravi A, Hojati V, Faghiri A. 2012. The reproductive cycle in the Grass Snake, Natrix na-
trix (Serpentes: Colubridae) in Iran. Russian Journal of Herpetology 19(3):217–220.
Shuntov VP. 1965. Sea snakes. NAVAL Oceanographic Office Nstl Station Ms; 1965 Jan 1.
Sidorovich VE, Sidorovich AA, Ivanovskij VV, Pikulik MM, Shinkevich EP. 2008. The
structure of vertebrate predator community in north-eastern Belarus before and after
naturalization of the American mink and raccoon dog. Folia Zoological 57:373–391.
Siegel DS, Sever DM, Rheubert JL, Gribbins KM. 2009. Reproductive biology of Agkistro-
don piscivorus Lacépede (Squamata, Serpentes, Viperidae, Crotalinae). Herpetological
monographs 23(1):74-107.
Silva FM, Prudente AL, Machado FA, Santos MM, Zaher H, Hingst‐Zaher E. 2018.
Aquatic adaptations in a Neotropical coral snake: A study of morphological conver-
gence. Journal of Zoological Systematics and Evolutionary Research 56(3):382-94.
Simões BF, Gower DJ, Rasmussen AR, Sarker MAR, Fry GC, Casewell NR, Harrison RA,
Hart NS, Partridge JC, Hunt DM, Chang BS, Pisani D & Sanders KL. 2020. Spectral di-
versification and trans-species allelic polymorphism during the land-to-sea transition
in snakes. Current Biology 1–8.
Simões BF, Sampaio FL, Jared C, Antoniazzi MM, Loew ER, Bowmaker JK, Rodriguez A,
Hart NS, Hunt DM, Partridge JC & Gower DJ. 2015. Visual system evolution and the
nature of the ancestral snake. Journal of Evolutionary Biology 28(7):1309–1320.
Sindaco R, Venchi A, Grieco C. 2013. The Reptiles of the Western Palearctic. 2. Annotated
checklist and distributional atlas of the snakes of Europe, North Africa, Middle East
and Central Asia, with an Update to Volume 1. Edizioni Belvedere, 543 pp.
Slowinski JP. 1994. The diet of kraits (Elapidae: Bungarus). Herpetological Review
25(2):51–52.
Smith HM, Brown BC. 1946. The identity of certain specific names in Thamnophis. Herpe-
tologica 3(3):71-2.
Smith HM. 1942. The synonymy of the garter snakes (Thamnophis), with notes on Mex-
ican and Central American species. Zoologica, Scientific Contributions of the New
York Zoological Society 27 (3-4): 97-123.
Smith LA. 1981. A revision of the Liasis olivaceus species-group (Serpentes: Boidae) in
Western Australia. Records of the Western Australian Museum. 9(2):227-33.
Smith MA. 1914a. The snakes of Bangkok. Part 1. Journal of the Natural History Society of
692
Aquatic Snakes

Siam 1:5–18.
Smith MA. 1914b. The snakes of Bangkok. Part 3. Journal of the Natural History Society of
Siam 1(2):93–104.
Smith MA. 1921. Two new batrachians and a new snake from Borneo and the Malay Penin-
sula. Journal of the Federated States of Malay State Museum. Singapore 10:197-199.
Smith MA. 1926. Monograph of the Sea-Snakes (Hydrophiidae). Trustees of the British Mu-
seum, London, xvii + 130 pp., 35 figs., 2 pis.
Smith MA. 1943. The Fauna of British India, Ceylon and Burma, Including the Whole of
the Indo-Chinese Sub-Region. Reptilia and Amphibia. 3 (Serpentes). Taylor and Francis,
London. 583 pp.
Smith SN, Jones MD, Marshall BM, Waengsothorn S, Gale GA, Strine CT. 2021. Native Bur-
mese pythons exhibit site fidelity and preference for aquatic habitats in an agricultural
mosaic. Scientific Reports 11(1):1-3.
Smith TL, Povel GDE, Kardong KV. 2002. Predatory strike of the tentacled snake (Erpeton
tentaculatum). Journal of Zoology 256(2):233–242.
Smole-Wiener AK. 2011. Verbreitung, Habitatnutzung und Gefahrdung der Wurfelnatter
(Natrix tessellata) in Karnten, Osterreich. Mertensiella 18:197-207.
Somaweera R, Ukuwela K, Alagoda T. 2006. A note on specimens of Gerarda prevostiana
(Colubridae: Serpentes) collected from Sri Lanka. Ceylon Journal of Science (Biological
Sciences) 35(1):91–93.
Somaweera R. 2009. An Overview of Sri Lankan Sea Snakes with an Annotated Checklist
and a Field Key. Taprobanica. 1: 43–54.
Souza V, Siefert JL, Escalante AE, Elser JJ, Eguiarte LE. 2012. The Cuatro Ciénegas Basin in
Coahuila, Mexico: an astrobiological Precambrian Park. Astrobiology 12 (7): 641–7.
Spellerberg IF. 1977. Reptile body temperatures. 2. Examples of reptile voluntary tempera-
ture levels. The Herptile 2:20-37.
Srinivasulu C, Srinivasulu B, Molur S. 2014. The status and distribution of reptiles in the
Western Ghats, India. Conservation Assessment and Management Plan (CAMP). Wild-
life Information Liaison Development Society, Coimbatore, Tamil Nadu, 148 pp.
Srivastava DP, Gautam S, Singh M, Sirohi U, Mukherjee D, Khudsar FA. 2020. First resight-
ing since the 1940s of Siebold’s Watersnake, Ferania sieboldii (Schlegel 1837), in Delhi,
India. Reptiles & Amphibians 27(3):525–527.
Starin ED, Burghardt GM. 1992. African rock pythons (Python sebae) in the Gambia: obser-
vations on natural history and interactions with primates. The Snake 24(1):50-62.
Stayton CT. 2011. Biomechanics on the half shell: functional performance influences pat-
terns of morphological variation in the emydid turtle carapace. Zoology 114(4):213–223.
Steen DA, Stevenson DJ, Beane JC, Willson JD, Aresco MJ, Godwin JC, Graham SP, Smith
LL, Howze JM, Rudolph DC, Pierce JB. 2013. Terrestrial movements of the red-bellied
mudsnake (Farancia abacura) and rainbow snake (F. erytrogramma). Herpetological Re-
view 44 (2): 208-213.
Stein B.A. Kutner L.S. Adams J.S. 2000. Precious Heritage: The Status of Biodiversity in the
United States. Oxford University Press; Oxford.
Steinhoff S. 2018. LD50 of venomous snakes. Available from: [Link]
Stejneger LH. 1907. Herpetology of Japan and adjacent territory. Bull. US Natl. Mus. 58:
xx, 1-577
693
Aquatic Snakes

Sterijovski B, Ajtic R, Tomovic L, Bonnet X. 2014. Conservation threats to Dice snakes (Na-
trix tessellata) in Golem Grad Island (FYR of Macedonia). Herpetological Conservation
and Biology 9(3):468-74.
Sterijovski B, Ajtic R, Tomovic L, Djordjević S, Djurakic M, Golubović A, Crnobrnja-Isa-
ilovic J, Ballouard JM, Groumpf F, Bonnet X. 2011. Natrix tessellata on Golem Grad, FYr
of Macedonia: a natural fortress shelters a prosperous snake Population. Mertensiella
18:298-302.
Storr GM, Smith LA, Johnstone RE. 1986. Snakes of Western Australia. Western Australian
Museum, Perth, 254pp.
Storr GM., Smith L.A, Johnstone RE. 1986. Snakes of Western Australia. Perth, WA: West-
ern Australian Museum 254pp
Stothers RB. 2004. Ancient scientific basis of the “Great Serpent” from historical evidence.
Isis 95(2):220-38.
Strasser P, Peters T. 2014. Zur Situation der Würfelnatter (Natrix tessellata) im Wiederan-
siedlungsgebiet „Knorre bei Meißen“ nach der erneuten Jahrhundertflut 2013. Jahress-
chrift zur Feldherpetologie und Ichthyofaunistik Sachsen (15).
Strugariu A, Gherghel I, Ghira I, Covaciu-Marcov SD, Mebert K. 2011. Distribution, habitat
preferences and conservation of the Dice snake (Natrix tessellata) in Romania. Mertensi-
ella 18:272-88.
Sturaro MJ, Gomes JO. 2008. Feeding behavior of the Amazonian water snake Helicops
hagmanni Roux, 1910 (Reptilia: Squamata: Colubridae: Hydropsini). Boletim do Museu
Paraense Emílio Goeldi. Ciências Naturais, Belém 3(3):225-228
Strüssmann C, de Brito ES, Marques OA. 2013. What do water snakes eat? First report of
predation by a Neotropical Hydropsini snake on giant earthworms (Glossoscolecidae).
Salamandra 49(1):48-50.
Strüssmann C. Uma Tatica De Caça 1990. Da Serpente Hydrodynastes gigas no Pantanal,
Mato Grosso. Mem. Inst. Butantan 52(2):57-61.
Stuart BL, Smith J, Davey K, Din P, Platt SG. 2000. Homalopsine watersnakes: the harvest
and trade from Tonle Sap, Cambodia. Traffic Bulletin-Cambridge-Traffic International
18(3):115–24.
Stuart BL, Sok K, Neang T. 2006. A collection of amphibians and reptiles from hilly eastern
Cambodia. Raffles Bulletin of Zoology 5:129–155.
Stuart BL. 2004. The harvest and trade of reptiles at U Minh Thuong National Park, south-
ern Viet Nam. Traffic Bulletin. 20(1):25-34.
Stuebing RB, Inger RF. 1999. Field guide to the snakes of Borneo. Natural History Publica-
tions (Borneo), Sabah, Malaysia, 254 pp.
Stuebing RB, Inger RF. B. Lardner. 2014. Field guide to the snakes of Borneo. Second Edi-
tion. Natural History Publications (Borneo), Sabah, Malaysia, 309 pp.
Su Y, Fong S-C, Tu M-C. 2005. Food habits of the sea snake, Laticauda semifasciata. Zoolog-
ical Studies 44(3):403-408.
Su G, Logez M, Xu J, Tao S, Villéger S, Brosse S. 2021. Human impacts on global freshwater
fish biodiversity. Science. 371(6531):835-8.
Suarez-Varon G. et al. 2016. Thamnophis eques (Mexican Gartersnake). Maximum size. Her-
petological Review 47(3):485.
Subaraj S, Law IS, Groenewoud D, Law IT. 2019. Blackwater mud snake at Upper Seletar.
694
Aquatic Snakes

Singapore Biodiversity Records 2019:2–3.

Subaraj S, Rajathurai S. 2018. Rainbow water snake in saltwater at Sungei Buloh. Singapore
Biodiversity Records 2018:35.
Subasli DR. 2020. Rediscovery of the Halmahera Keelback, Tropidonophis punctiventris
(Boettger. 1895)(Serpentes: Colubridae) After 116 Years. Zoo Indonesia 28(1)59-63.
Sung YH and Wan JPH. (2011). Opisthotropis kuatunensis. Diet. Herpetological Review,
42(2), 298.
Sworder GH. 1922. A list of the snakes of Singapore Island. Singapore Naturalist 2:55–73.
Takeuchi H, Savitzky AH, Ding L, de Silva A, Das I, Nguyen TT, Tsai TS, Jono T, Zhu GX,
Mahaulpatha D, Tang Y. 2018. Evolution of nuchal glands, unusual defensive organs of
Asian natricine snakes (Serpentes: Colubridae), inferred from a molecular phylogeny.
Ecology and evolution. 8(20):10219-32.
Tallowin O, Parker F, O’Shea M. 2017. Tropidonophis doriae. The IUCN Red List of
Threatened Species 2017: e.T42492548A42492555. [Link]
[Link].
Tamiya N, Puffer H. 1974. Lethality of sea snake venoms. Toxicon, 12(1), 85–87.
doi:10.1016/0041-0101(74)90104-4
Tan CH, Wong KY, Tan KY, Tan NH. 2017. Venom proteome of the yellow-lipped sea
krait, Laticauda colubrina from Bali: Insights into subvenomic diversity, venom an-
tigenicity and cross-neutralization by antivenom. Journal of Proteomics, 166, 48–58.
doi:10.1016/[Link].2017.07.002
Tan HH, Li TJ, Lim WH. 2014. A food item of the blackwater mud snake. Singapore Biodi-
versity Records 2014:311–312
Tanaka K. 2002. Foraging behavior of Rhabdophis tigrinus (Serpentes: Colubridae) in a
gutter with a dense aggregation of tadpoles. Current herpetology, 21(1):1-8.
Tangella K. (n.d.). Pacific Sea snake bite. DoveMed. Retrieved May 20, 2022, from https://
[Link]/diseases-conditions/pacific-sea-snake-bite/
Tanner WW. 1990. Thamnophis rufipunctatus. Catalogue of American Amphibians and
Reptiles (CAAR).
Tashmuradov MYa. 1976. Reptiles of the Floodplains of the Middle Reaches of the River.
Angren, MS, Tashkent State University, Tashkent, Uzbekistan, 69 pp.
Taub AM. 1967. Comparative histological studies on Duvernoy’s gland of colubrid
snakes. Bulletin of the American Museum of Natural History 138:1–150.
Tavares-Pinheiro R, Melo FS, de Figueiredo VA, Sanches PR, Anaissi JS, Santana MM,
Oliveira-Souza AE, dos Santos Reis T, Rebelo GL, Melo FT, Costa-Campos CE. 2021.
“In living color”: predation by the coralsnake Micrurus surinamensis (Cuvier, 1816)
(Serpentes: Elapidae) on a knifefish in the eastern Amazon, Brazil. Herpetology Notes
14:755-8.
Taylor C, Mayes K. 2019. Impact of Hydrologic Alteration on Brazos River Vertebrates.
Final Performance Report 09/01/2016–12/30/2018 As Required By The Endangered
Species Program Texas.
Taylor EH. 1922. The snakes of the Philippine Islands. Manila (Bureau of Printing or Sci-
ence), Monograph 16: 312 pp.
Taylor EH. 1965. The serpents of Thailand and adjacent waters. University of Kansas Sci-
ence Bulletin 45:609–1096.
695
Aquatic Snakes

Taylor JA. 1979. The foods and feeding habits of subadult Crocodylus porosus Schneider in
Northern Australia. Australian Wildlife Research 6(3):347–359.
Taylor P. 1970. An observation on the feeding habits of Lycodonomorphus rufulus. The
Journal of the Herpetological Association of Africa. 6(1):19-20.
Teixeira VH, Quintela FM, Loebmann D. 2020. Reproductive biology of Erythrolamprus
jaegeri coralliventris (Serpentes: Dipsadidae) in the Brazilian Coastal Pampa. Anais da
Academia Brasileira de Ciências 12:92.
Tennant, A. 1984. Snakes of North America - Eastern and Central Regions. Lone Star
Books, 605 pp.
Thakur S, Watve A. 2009. Occurrence of Enhydris sieboldii (Schlegel, 1837) in Bhopal,
Madhya Pradesh State of India. Russian Journal of Herpetology 16(2):159–160.
Theobald W. 1868. Catalogue of the reptiles of British Birma, embracing the provinces of
Pegu, Martaban, and Tenasserim; with descriptions of new or little-known species. Zo-
ological Journal of the Linnean Society 10(41):4–67.
Thomas DW, Adams CS, Schalk CM. 2020. Farancia abacura (Red-bellied Mudsnake).
Overland movement. Herpetological Review 51(1)143.
Thomas N, Li TJ, Lim WH, Cai Y. 2014. New record of the blackwater mud snake in Singa-
pore. Singapore Biodiversity Records 2014:309–310.
Thorpe R.S., 1973. Intraspecific variation of the ringed snake, Natrix natrix (L.) Ph. D. dis-
sert C.N.N.A., [1] + 116 pp., app. 1-10; pp. [117-317], 70+36 ff.
Thorpe RS, 1975a. Biometric analysis of incipient speciation in the ringed snake Natrix na-
trix (L.). Experientia, 31: 180-182.
Thorpe RS, 1975b. Quantitative handling of characters useful in snake systematics with
reference to intraspedific variation in the ringed snake Natrix natrix (L.). Biological
Journal of the Linnean Society, 7 (1): 27-43.
Thorpe RS, 1979. Multivariate analysis of the population systematics of the ringed snake
Natrix natrix (L.). Proceedings of the Royal Society of Edinburgh, 78 (B): 1-62.
Thorpe RS, 1980. A comparative study of ordination techniques in numerical taxonomy in
relation to racial variation in the ringed snake Natrix natrix (L.). Biological Journal of
the Linnean Society, 13: 7-40.
Thorpe RS, 1989. Pattern and function of sexual dimorphism: a biometric study of charac-
ter variation in the Grass Snake (Natrix natrix, Colubridae) due to sex and its interac-
tion with geography. Copeia 1989: 53-63.
Thorpe RS. 1980e. Microevolution and taxonomy of European reptiles with particular
reference to the grass snake Natrix natrix and the wall lizards Podarcis sicula, P. melisel-
lensis. Biological Journal of Linnean Society, 14: 215-233.
Tingle JL, Gartner GE, Jayne BC, Garland T. 2017. Ecological and phylogenetic variabili-
ty in the spinalis muscle of snakes. Journal of evolutionary biology. 30(11):2031-43.
Tonks ML, Griffiths SP, Heales DS, Brewer DT, Dell Q. 2008. Species composition and
temporal variation of prawn trawl bycatch in the Joseph Bonaparte Gulf, northwestern
Australia. Fisheries Research 89: 276–293.
Torres-Carvajal O, Hinojosa KC. 2020. Hidden diversity in two widespread snake species
(Serpentes: Xenodontini: Erythrolamprus) from South America. Molecular Phyloge-
netics and Evolution 146:106772.
Torres-Porras J. 2008. Apareamiento grupal de la culebra viperina (Natrix maura). Boletín
696
Aquatic Snakes

de la Asociación Herpetológica Española 19:47-49.

Tpantiza-Tuguminago L, Medrano-Vizcino P, Arguello P, Auqui E. 2019. Erythrolamprus
miliaris crysostomus (Military Ground Snake) Diet. Herpetological Review 50(4): 800.
Thorpe RS. 1980a. A comparative study of ordination techniques in numerical taxonomy
in relation to racial variation in the ringed snake Natrix natrix (L.). Biological Journal
of the Linnean Society 13:7–40.
Thorpe RS. 1980b. Microevolution and taxonomy of European reptiles with particular
reference to the grass snake Natrix natrix and the wall lizards Podarcis sicula, P. melisel-
lensis. Biological Journal of Linnean Society 14:215–233.
Thorpe RS. 1984. Primary and secondary transition zones in speciation and population
differentiation: a phylogenetic analysis of range expansion. Evolution 38:233–243.
Thorpe RS. 1989. Pattern and function of sexual dimorphism: a biometric study of charac-
ter variation in the Grass Snake (Natrix natrix, Colubridae) due to sex and its interac-
tion with geography. Copeia 1:53–63.
Trapp B, Mebert K. 2011. Upward position of eyes and nostrils of the Dice Snake to break
the water surface? Mertensiella, 18:440.
Trauth SE. 1991. Distribution, scutellation, and reproduction in the Queen Snake, Regina
septemvittata (Serpentes: Colubridae), from Arkansas. Journal of the Arkansas Acade-
my of Science 45(1):103-6.
Trobisch and Gläßer-Trobisch 2011. studied hunting behavior in captive juvenile Natrix
tessellata and report them hunting from branches over water.
Trobisch D and A. Gläßer-Trobisch. 2011. The Rearing of Dice Snakes: Part of a Concept
for the Sustainable Conservation of Endangered and Isolated Populations in Western
Germany. Mertensiella 18:49-57.
Tu MC, Fong SC, Lue KY. 1990. Reproductive Biology of the Sea Snake, Laticauda semi-
fasciata, in Taiwan. Journal of Herpetology, 24(2), 119. doi:10.2307/1564218
Tucker JK. 1977. Notes on the food habits of Kirtland’s water snake, Clonophis kirtlandi.
Bulletin of the Maryland Herpetological Society 13(3): 193-195.
Tucker JK. 1995. Notes on road-killed snakes and their implications on habitat modifica-
tion due to summer flooding on the Mississippi River in west central Illinois. Trans-
actions of the Illinois State Academy of Science 88(1/2):61-71.
Tumlison R, Roberts KG. 2018. Prey-handling behavior in the Gulf Crayfish Snake (Li-
odytes rigida). Herpetological Conservation and Biology 13(3):617-21.
Tuner FB. 1977. The dynamics of populations of squamates and crocodilians. In: Biology
of the Reptilia 7 (Gans C, Tinkle, D.W. Eds.) Academic Press, New York: 157–264.
Tuniyev B, Tuniyev S, Kirschey T, Mebert K. 2011. Notes on the Dice snake (Natrix tessel-
lata) from the Caucasian Isthmus. Mertensiella 18:343-56.
Tuttle KN, Gregory PT. 2009. Food habits of the Plains Garter Snake (Thamnophis radix)
at the northern limit of its range. Journal of Herpetology 43:65–73.
Tuttle KN, Gregory PT. 2012. Growth and maturity of a terrestrial ectotherm near its
northern distributional limit: Does latitude matter? Canadian Journal of Zoology
90:758–765.
Tuttle KN, Gregory PT. 2014. Reproduction of the plains garter snake, Thamnophis
radix, near its northern range limit: more evidence for a “fast” life history Copeia
2014(1):130-5.
697
Aquatic Snakes

Tweedie MWF. 1946. A new snake from the Malay Peninsula. Annals and Magazine of
Natural History 13(98):142–144.
Tweedie MWF. 1983. The Snakes of Malaya. Singapore National Printers, Singapore, 167
pp.
Udyawer V, Barnes P, Bonnet X, Brischoux F, Crowe-Riddell JM, D’Anastasi B, Fry BG,
Gillett A, Goiran C, Guinea ML, Heatwole H, Heupel MR, Hourston M, Kangas M,
Kendrick A, Koefoed I, Lillywhite HB, Lobo AS, Lukoschek V, McAuley R, Nitschke
C, Rasmussen AR, Sanders KL, Sheehy C III, Shine R, Somaweera R, Sweet SS and Vo-
ris HK (2018) Future Directions in the Research and Management of Marine Snakes.
Frontiers in Marine Science 5:399. doi: 10.3389/fmars.2018.0039
Udyawer V, Goiran C, Chateau O & Shine R. 2020. Swim with the tide: Tactics to maxi-
mize prey detection by a specialist predator, the greater sea snake (Hydrophis major).
PLoS ONE 15(10 October):1–13.
Udyawer V, Read M, Hamann M, Heupel MR, Simpfendorfer CA. 2016. Importance of
Shallow Tidal Habitats as Refugia from Trawl Fishing for Sea Snakes. Journal of Herpe-
tology, 50(4), 527–533. doi:10.1670/15-026
Udyawer V, Read M, Hamann M, Heupel MR, Simpfendorfer CA. 2016. Importance of
Shallow Tidal Habitats as Refugia from Trawl Fishing for Sea Snakes. Journal of Herpe-
tology, 50(4), 527–533. doi:10.1670/15-026
Udyawer V, Simpfendorfer CA, Heupel MR & Clark TD. 2016. Coming up for air: ther-
mal-dependence of dive behaviours and metabolism in sea snakes. The Journal of Ex-
perimental Biology September, jeb.146571.
Uetz P, Freed P, Hošek J. 2020. The Reptile Database, [Link] ac-
cessed 21 January 2021
Uetz, P. (editor), 2023. The Reptile Database, [Link] accessed
December 03, 2023.
Ujvari B, Brown G, Shine R, Madsen T. 2016. Floods and famine: climate‐induced collapse
of a tropical predator‐prey community. Functional ecology 30(3):453-8.
Ukuwela KBD, Sanders KL, Fry BG 2012. Hydrophis donaldi (Elapidae, Hydrophiinae),
a highly distinctive new species of sea snake from northern Australia. Zootaxa 3201:
45–57
Ukuwela KD, De Silva A, Sanders KL. 2017. Further specimens of the mud snake, Gerarda
prevostiana (Homalopsidae) from Sri Lanka with insights from molecular phylogenet-
ics. Raffles Bulletin of Zoology 65:29–34. doi:10.1016/[Link].2012.09.031
Ukuwela KDB, de Silva A, Mumpuni, Fry BG. Lee MSY, Sanders KL. 2013. Molecular ev-
idence that the deadliest sea snake Enhydrina schistosa (Elapidae: Hydrophiinae) con-
sists of two convergent species. Molecular Phylogenetics and Evolution, 66: 262–269.
doi:10.1016/[Link].2012.09.031
Ultsch GR. 1989. Ecology and Physiology of Hibernation and Overwintering
Among Freshwater Fishes, Turtles, and Snakes. Biological Reviews, 64: 435–515.
doi: 10.1111/j.1469-185X.1989.tb00683.x
Underwood G. 1967. A contribution to the classification of snakes. Publication 635. Trust-
ees of the British Museum (Natural History), London.
Underwood G. 1999. Morphological evidence on the affinities of vipers. Kaupia 8:3–8.

698
Aquatic Snakes

UNEP World Conservation. Largest lake in the Pacific. Protected Areas Programme.
United Nations. Archived from the original on 18 July 2008. Retrieved 12 December
2020.
Vacher J-P, M Geniez. 2010. Les reptiles de France, Belgique, Luxembourg et Suisse. Bio-
tope, Méze (Cllection Parthénope. Muséum national d’Histoire naturelle, Paris, 544 p.
Valverde JA. 1974. N. natrix pescando Phoxinus sp. Doñana, Acta Vertebrata 1: 58–59.
Van Cao N, Thien Tao N, Moore A, Montoya Am Rasmussen RA, Broad K, Voris HK,
Takacs Z. 2014. Sea Snake Harvest in the Gulf of Thailand. Conservation biology. Vol-
ume 28, No. 6, 1677–1687
Van Der Kooij J, Povel D. 1996. Scale sensillae of the file snake (Serpentes: Acrochordi-
dae) and some other aquatic and burrowing snakes. Netherlands Journal of Zoology.
47(4):443-56.
Van der Meijden A, Chiari Y. 2006. Natural history notes. Natrix tessellata (Dice snake)
[sic]. Marine habitat. Herpetological Review 37(1):94.
Vanek JP, Eastin DM, Call JN, Glowacki GA, Rutter AU. 2021. Thamnophis sirtalis semifas-
ciatus (Chicago Gartersnake). Scavenging and Diet.
Vanni S, R. Cimmaruta. 2010. Natrix cetti. In Corti C., Capula M., Luiselli L., Sindaco R.,
Razzetti E., Fauna d’Italia, vol. XLV, Reptilia. Calderini, Bologna, XII + 869 pp.
Vanni S, R. Cimmaruta. 2011. Natrix cetti, in Corti C., Capula M., Luiselli L., Sindaco R.,
Razzetti E., 2011. Fauna d’Italia, vol. XLV, Reptilia. Calderini, Bologna, XII + 869 pp.
Vashetko EV. 1981. Population number and biomass of some reptile species in natural and
anthropogenic conditions of Jizak Province, Uzbekistan. Problems of Herpetology,
Leningrad, 29 [in Russian].
Vásquez-Restrepo JD. 2020. Validating the Two Records of Mocquard’s Swampsnake,
Tretanorhinus mocquardi Bocourt 1891 (Dipsadidae), in Colombia, with Comments
on the Taxonomy of Mainland Swampsnakes. Reptiles & Amphibians. 2020 Nov
30;27(2):314-7.
Velensky M, Velensky P, Mebert K. 2011. Ecology and ethology of Dice snakes (Natrix tes-
sellata) in the City District Troja, Prague. Mertensiella 18:157-77.
Velikov I. 2011. Dice snake feeds on spiny invasive fish. Mertensiella 18:430.
Vericad JR, A Escarré. 1976. Datos de alimentación de ofidios en el Levante sur ibérico.
Mediterránea 1:3-32.
Vidal N, Azvolinsky A, Cruaud C, Hedges SB. 2008. Origin of tropical American burrow-
ing reptiles by transatlantic rafting. Biology letters, 4(1):115-118.
Vidal N, Branch WR, Pauwels OS, Hedges SB, Broadley DG, Wink M, Cruaud C, Joger
U, Nagy ZT. 2008. Dissecting the major African snake radiation: a molecular phylog-
eny of the Lamprophiidae Fitzinger (Serpentes, Caenophidia). Zootaxa 1945(1):51-
66.
van Riemsdijk I, Struijk RPJH, Pel E, Janssen IAW, Wielstra B. 2020. Hybridisation compli-
cates the conservation of Natrix snakes in the Netherlands. Salamandra 56(1):78–82.
Vanni S, Cimmaruta R. 2011. Natrix cetti. In: Corti C, Capula M, Luiselli L, Sindaco R,
Razzetti E. Fauna d’Italia, vol. XLV, Reptilia. Calderini, Bologna, Italy, 869 pp.
Vericad JR, Escarré A. 1976. Datos de alimentación de ofidios en el Levante sur ibérico.
Mediterránea 1:5–33.
Vidal N, Delmas AS, David P, Cruaud C, Couloux A, Hedges SB. 2007. The phylogeny
699
Aquatic Snakes

and classification of caenophidian snakes inferred from seven nuclear protein-coding

genes. Comptes rendus biologies 1;330(2):182-7.
Vidal N, Dewynter M, Gower DJ. 2010. Dissecting the major American snake radiation: a
molecular phylogeny of the Dipsadidae Bonaparte (Serpentes, Caenophidia). Comptes
rendus biologies 333(1):48-55.
Vidal N, Hedges SB. 2004. Molecular evidence for a terrestrial origin of snakes. Proceed-
ings of the Royal Society of London. Series B: Biological Sciences. 271(suppl 4):S226-
9.
Vidal NRJ, Couloux A, Hedges SB. 2009 Snakes (Serpentes). In: Hedges, S.B. and Kumar, S.
(eds.), The Timetree of life. Oxford University Press, pp. 390-397.
Villa J. 1969. Two new insular subspecies of the natricid snake Tretanorhinus nigroluteus
Cope from Honduras and Nicaragua. Journal of Herpetology 3: 145–150.
Villa JD. 1990. Hydromorphus, H. concolor, H. dunni. Catalogue of American Amphibians
and Reptiles (CAAR).
Vincent SE, Brandley MC, Herrel A, Alfaro ME. 2009. Convergence in trophic mor-
phology and feeding performance among piscivorous natricine snakes. Journal
of Evolutionary Biology 22: 1203–1211. doi:10.1111/[Link]-
MID:1938915356.
Vincent SE, MC Brandley, A. Herrel, ME Alfaro. 2009. Convergence in trophic morphology
and feeding performance among piscivorous natricine snakes. Journal of Evolutionary
Biology 22(6):1203–1211.
Vincent SE, Moon BR, Shine R, Herrel A. 2006. The functional meaning of “prey size” in
water snakes (Nerodia fasciata, Colubridae). Oecologia 147(2):204-11.
Vincent, SE., R Shine, GP Brown. 2005. Does foraging mode influence sensory modalities
for prey detection in male and female filesnakes, Acrochordus arafurae? Animal Be-
haviour, 70: 715–721.
Vincent, SE., Shine R, Brown GP. 2005. Does Foraging Mode Influence Sensory Modalities
for Prey Detection in Male and Female Filesnakes, Acrochordus arafurae? Animal Be-
haviour 70: 715–21. [Link]
Visser J. 1967. Color varieties, brood size, and food of South African Pelamis platurus
(Ophidia: Hydrophiidae). Copeia 1967:219.
Vitt LJ, Caldwell JP. 2009. Herpetology: An Introductory Biology of Amphibians and Rep-
tiles. Third Edition, Academic Press.
Vlcek P, Jablonski D, Kudlacek M, Mebert K. 2015. Rediscovery of the Dice Snake Natrix
tessellata (Laurenti, 1768), from the Island of Krk, Croatia. Herpetozoa 27(3/4): 191-96.
Vlcek P, Zavadil V, Jablonski D, Mebert K. 2011. Dice snake (Natrix tessellata) in the Baltic
Sea Drainage Basin (Karvinsko District in Silesia, Czech Republic). Mertensiella 18:177-
88.
Vogel S. 1994. Life in moving fluids: the physical biology of flow. Princeton University Press.
Vogel G, David P. 2006. On the taxonomy of the Xenochrophis piscator complex (Serpentes,
Natricidae). In Proceedings of the 13th Congress of the Societas Europaea Herpetologi-
ca 2006 (Vol. 241, p. 246).
Vogel G, David P. 2012. A revision of the species group of Xenochrophis piscator (Schneider,
1799) (Squamata: Natricidae). Zootaxa 3473(1):1-60.
Vogrinc PN, Durso AM, Winne CT, Willson JD. 2018. Landscape-Scale Effects of Su-
700
Aquatic Snakes

pra-Seasonal Drought on Semi-Aquatic Snake Assemblages. Wetlands 38: 667-676.

von May R, de Albuquerque NR, Braz HB, Santa-Cruz R, Biggi E, Tomasinelli F, Ra-
bosky DL. 2019. Distribution of the Neotropical water snakes Hydrops caesurus, H.
martii, and H. triangularis in South America, with new records from Peru and Brazil.
Amphibian & Reptile Conservation 13(1) [General Section]: 122–142 (e173).
Voris HK, Alfaro ME, Karns DR, Starnes GL, Thompson E, Murphy JC. 2002. Phyloge-
netic relationships of the Oriental-Australian rear-fanged water snakes (Colubridae:
Homalopsinae) based on mitochondrial DNA sequences. Copeia 2002(4):906–915.
Voris HK, Glodek GS. 1980. Habitat, diet, and reproduction of the file snake, Acrochordus
granulatus, in the Straits of Malacca. Journal of Herpetology. 1980 14(1):108-11.
Voris HK, Jeffries WB. 1995. Predation on marine snakes: a case for decapods supported
by new observations from Thailand. Journal of Tropical Ecology 11:569–576.
Voris HK, Karns DR. 1996. Habitat utilization, movements, and activity patterns of En-
hydris plumbeus (Serpentes: Homalopsinae) in a rice paddy wetland in Borneo. Herpe-
tological Natural History 4(2):111–126.
Voris HK, Murphy JC. 2002. The prey and predators of homalopsine snakes. Journal of
Natural History 36(13):1621–1632.
Voris HK, Voris HH & Liat LB. 1978. The food and feeding behavior of a marine snake,
Enhydrina schistosa (Hydrophiidae). Copeia 1978(1):134–146.
Voris HK, Voris HH. 1983. Feeding strategies in marine snakes: an analysis of evolution-
ary, morphological, behavioural and ecological relationships. American Zoologist.
23(2):411-425.
Voris HK, Voris HH. 1995. Commuting on the tropical tides: the life of the yel-
low-lipped sea krait Laticauda colubrina. Ocean Realm, April 1995:56–61.
Voris HK. 1972. The role of sea snakes (Hydrophiidae) in the trophic structure of coastal
ocean communities. Journal of the Marine Biological Association of India 14(2): 429-
442.
Vyas R. 2011. Preliminary Survey on Reptiles of Jassore Wildlife Sanctuary, Gujarat State,
India. Russian Journal of Herpetology. 2011;18(3):210-4.
Vyas R. 2013. Snake diversity and voluntary rescue practice in the cities of Gujarat State,
India: an evaluation. Reptile Rap 15:27-39.
Vyas RV, Murphy JC, Voris HK. 2013. The dog-faced water snake (Cerberus rynchops)
and Gerard’s mud snake (Gerarda prevostiana) at the western edge of their distribu-
tion. Herpetological Review 44(1):34–36.
Wagner P, Bauer AM, Leviton AE, Wilms TM, Böhme W. 2016. A checklist of the amphi-
bians and reptiles of Afghanistan. Exploring herpetodiversity using biodiversity archi-
ves. Proceedings of the Californian Academy of Sciences 63:457–565.
Walker CF. 1955. A new gartersnake (Thamnophis) from Tamaulipas. Copeia 1955 (2):
110-113.
Wall F. 1897. Notes on two specimens of Hypsirhina sieboldii. Journal of the Bombay
Natural History Society 11:732–734.
Wall F. 1907. Some new Asian snakes. Journal of Bombay Natural History Society 17:612-
618.
Wall F. 1908. Notes on a gravid female of Siebold’s watersnake (Hypsirhina sieboldi). Jour-
nal of the Bombay Natural History Society 18:920.
701
Aquatic Snakes

Wall F. 1918. A popular treatise on the common Indian snakes. Part 26. [Cerberus rhyn-
chops, Enhydris curtus]. Journal of the Bombay Natural History Society 23:166.
Wall F. 1919. Notes on a collection of snakes made in the Nilgiri Hills and the adjacent
Wynaad. Journal of the Bombay Natural History Society 26: 552–584.
Wall F. 1921. Ophidia Taprobanica: Or, The Snakes of Ceylon. H.R. Cottle, Government
Press, Colombo, 581 pp.
Wall F. 1924. A hand–list of the snakes of the Indian Empire. Part 3. Journal of the Bombay
Natural History Society 29:864–878.
Wall F. 1925. Notes on snakes collected in Burma in 1924. Journal of the Bombay Natural
History Society 30(4): 805-821.
Wallach V, Williams KL, Boundy J. 2014. Snakes of the World. A Catalogue of Living and
Extinct Species. CRC Press Boca Raton Florida. ISBN: 978-1-4822-08474
Wallach V, Williams KL, Boundy J. 2014. Snakes of the World. A Catalogue of Living and
Extinct Species. CRC Press Boca Raton Florida. ISBN: 978-1-4822-08474.
Wallach V, Wüster W, Broadley DG (2009). “In praise of subgenera: taxonomic status of
cobras of the genus Naja Laurenti (Serpentes: Elapidae)”. Zootaxa 2236: 26–36.
Waller T, Micucci PA, Alvarenga E. 2007. Conservation biology of the yellow anaconda
(Eunectes notaeus) in Northeastern Argentina. p. 340-362 In: Biology of the Boas and
Pythons 2007 R. W. Henderson and R. Powell (Eds.). Eagle Mountain Publishing, LC.
Utah. 438 pp.
Walley HD, Wusterbarth TL, Stanford KM. 2003. Thamnophis radix. CAAR.
Walls GL. 1940. Ophthalmological implications for the early history of the snakes. Co-
peia 1940: 1-8.
Walters TM, Mazzotti FJ, Fitz HC. 2016. Habitat selection by the invasive species Bur-
mese python in Southern Florida. Journal of Herpetology 50(1):50-6.
Walters TM, Mazzotti FJ, Fitz HC. 2016. Habitat selection by the invasive species Burmese
python in Southern Florida. Journal of Herpetology 50(1):50-6.
Walztöhny D, Ziswiler V. 1979. Vergleichend - morphologische Untersuchungen an den
Hautsinnesorganen der Blindschleiche Anguis fragilis (Anguidae) und der Würfel-
natter, Natrix tessellata (Colubridae). – Rev. Suisse Zool. 86 (3):705–712.
Wang J, Lyu Z-T, Zeng Z-C, Lin C-Y, Yang J-H, Nguyen TQ, Le MD, Ziegler T. and
Wang Y-Y. 2020. Re-examination of the Chinese record of Opisthotropis maculosa
(Squamata, Natricidae), resulting in the first national record of O. haihaensis and de-
scription of a new species. ZooKeys 913: 141-159.
Wang K, Jiang KE, Jin J, Liu XU, Che J. Confirmation of Trachischium guentheri (Ser-
pentes: Colubridae) from Tibet, China, with description of Tibetan T. monticola.
Zootaxa. 2019 Oct 21;4688(1):101-10.
Wang S., Lillywhite, H. B., & Tu, M.-C. (2013). Locomotor performance of three sym-
patric species of sea kraits (Laticauda spp.) from Orchid Island, Taiwan. Zoological
Studies, 52(1), 43. doi:10.1186/1810-522x-52-43.
Wang XH, Messenger K. 2014. Reclassification of Oligodon ningshaanensis Yuan, 1983
(Ophidia: Colubridae) into a New Genus, Stichophanes gen. nov with Description on
Its malacophagous Behavior. Asian Herpetological Research, 5(3):137-149
Wang YY, Guo Q, Liu ZY, Lyu ZT, Wang J, Luo L, Sun YJ, Zhang YW. 2017. Revisions
of two poorly known species of Opisthotropis Günther, 1872 (Squamata: Colubridae:
702
Aquatic Snakes

Natricinae) with description of a new species from China. Zootaxa. 4247(4):391-412.

Ward TM 1996. Sea snake by-catch of prawn trawlers on the northern Australian conti-
nental shelf. Marine and Freshwater Research 47: 631–635.
Ward TM. [Link] affecting the catch rates and relative abundance of sea snakes in
the by-catch of trawlers targeting tiger and endeavour prawns on the northern Austra-
lian continental shelf. Marine and Freshwater Research 51(2):155-64.
Ward TM. 2001. Age structures and reproductive patterns of two species of sea snake, La-
pemis hardwickii Grey (1836) and Hydrophis elegans (Grey 1842), incidentally captured
by prawn trawlers in northern Australia. Marine and Freshwater Research 52(2):193-
203.
Warren LV, Quaglio F, Simoes MG, Freitas BT, Assine ML, Riccomini C. 2014. Underneath
the pantanal wetland: a deep-time history of Gondwana assembly, climate change, and
the dawn of metazoan life. Pages 1-21 in Dynamics of the Pantanal wetland in South
America Springer, Cham.
Wassenberg TJ, Milton DA and Burridge CY 2001. Survival rates of sea snakes caught by
demersal trawlers in northern and eastern Australia, Biological Conservation 100, pp.
271–80
Wassenberg TJ, Salini JP, Heatwole H, Kerr JD (1994) Incidental capture of sea-snakes
(Hydrophiidae) by prawn trawlers in the Gulf of Carpentaria, Australia Australian
Journal of Marine and Freshwater Research 45: 429–443.
Waters RM, Burghardt GM. 2005. The interaction of food motivation and experience in
the ontogeny of chemoreception in crayfish snakes. Animal Behaviour 69(2):363–374.
Waters RM, Burghardt GM. 2013. Prey availability influences the ontogeny and timing of
chemoreception-based prey shifting in the striped crayfish snake, Regina alleni. Journal
of Comparative Psychology 127(1):49–55.
Weatherhead PJ, Brown GP, Prosser MR, Kissner KJ. 1999. Factors affecting neonate size
variation in northern water snakes, Nerodia sipedon. Journal of Herpetology 1999:577-
89.
Webb RG. Thamnophis cyrtopsis. Catalogue of American Amphibians and Reptiles
(CAAR). 1980.
Weinstein SA, Schmidt JJ, Smith LA. 1991. Lethal toxins and cross-neutralization of
venoms from the African water cobras, Boulengerina annulata annulata and Boulen-
gerina christyi. Toxicon. 29 (11): 1315–1327.
Weiperth A, Gaebele T, Potyó I, Puky M. 2014. A global overview on the diet of the Dice
snake (Natrix tessellata) from a geographical perspective: foraging in atypical habitats
and feeding spectrum widening helps colonization and survival under suboptimal con-
ditions for a piscivorous snake. Zoological Studies 53:42.
Welsh HH, Lind AJ. 2000. Evidence of lingual-luring by an aquatic snake. Journal of Her-
petology 34(1):67–74
Werner F. 1923. Neue Schlangen des Naturhistorischen Museum in Wien. Annalen des
Naturhistorischen Museum in Wien 36:160–166.
Werner YL, Shapira T. 2011. A brief review of morphological variation in Natrix tessellata
in Israel: between sides, among individuals, between sexes, and among regions. Turk-
ish Journal of Zoology 35(4):451–466. [Link]
Wesselingh FP. 2008. Molluscan radiations and landscape evolution in Miocene Amazonia.
703
Aquatic Snakes

Turun Yliopiston Julkaisuja Annales Universitates Turkuensis 232:3-41.

Westhoff G, Fry BG, Bleckmann H. 2005. Sea snakes (Lapemis curtus) are sensitive to
low-amplitude water motions. Zoology 108(3):195–200.
Whitaker R, 1978. Common Indian Snakes: A Field Guide. Macmillan India Ltd, p. 154
Whitaker R, Captain A, Ahmed F. 2004. Snakes of India: The Field Guide. Draco Books,
Chennai, 481 pp.
Whitaker R. 1969. The dog-faced watersnake (Cerberus rhynchops) in the Bombay area and
notes on its habits. Journal of the Bombay Natural History Society 66:386.
Whitaker R. 1993. Population status of the Indian Python (Python molurus) on the Indian
subcontinent. Herpetological Natural History 5(1):87-89.
White DR, Mitchell JC, Woolcott WS. 1982. Reproductive cycle and embryonic develop-
ment of Nerodia taxispilota (Serpentes: Colubridae) at the northeastern edge of its range.
Copeia. 1982:646-52.
Whiting MJ, Dixon JR, Greene BD, Mueller JM, Thornton OW, Hatfield JS, Nichols JD,
Hines JE. 2008. Population dynamics of the Concho water snake in rivers and reser-
voirs. Copeia 2008(2):438-45.
Wikramanayake ED, Dinerstein E, Loucks CJ. 2002. Terrestrial ecoregions of the Indo-Pa-
cific: a conservation assessment (Vol. 3). Island Press, Washington, DC, 589 pp.
Wilcox TP, Zwickl DJ, Heath TA, Hillis DM. 2002. Phylogenetic relationships of the dwarf
boas and a comparison of Bayesian and bootstrap measures of phylogenetic support.
Molecular phylogenetics and evolution, 25(2):361-371.
Wilkinson C. 2008. Executive summary. Pp 1–19 In Status of Coral Reefs of the World:
2008 Global Coral Reef Monitoring Network and Reef and Rainforest Research Centre.
Wilkinson, C. (Ed.). Townsville, Queensland, Australia.
Williamson JA, Fenner PJ. Burnett JW. Rifkin JF. 1996. Venomous and Poisonous Marine
Animals: A Medical and Biological Handbook, UNSW Press, p. 403, ISBN 978-0-86840-
279-6
Willson JD, Hopkins WA. 2011. Prey morphology constrains the feeding ecology of an
aquatic generalist predator. Ecology 92(3):744-54.
Willson JD, Snow RW, Reed RN, Dorcas ME. 2014. Python molurus bivittatus (Burmese Py-
thon). Minimum size at maturity. Herpetological Review 45(2):343-4.
Willson JD, Winne CT, Dorcas ME, Gibbons JW. 2002. Post-drought responses of
semi-aquatic snakes inhabiting an isolated wetland: insights on different strategies for
persistence in a dynamic habitat. Wetlands 26(4):1071-8.
Wilson EO, Brown WL. 1953. The subspecies concept and its taxonomic application. Sys-
tematic zoology 2(3):97-111.
Wilson S, Swan G. 2003. Reptiles of Australia. Princeton Field Guide Series. Princeton Uni-
versity Press, Princeton.
Wilson SK, Swan G. 2013 A complete guide to reptiles of Australia / Steve Wilson, Gerry
Swan New Holland Publishers Chatswood, N.S.W
Winne CT, Dorcas MD, Poppy SM. 2005. Population structure, body size, and seasonal ac-
tivity of Black Swamp Snakes (Seminatrix pygaea). Southeastern Naturalist 4:1-14.
Winne CT, Willson JD, Gibbons JW. 2006. Income breeding allows an aquatic snake Sem-
inatrix pygaea to reproduce normally following prolonged drought-induced aestiva-
tion. Journal of Animal Ecology, 75(6):1352-60.
704
Aquatic Snakes

Winokur RM. The integumentary tentacles of the snake Erpeton tentaculatum: structure,
function, evolution. Herpetologica. 1977 Jun 1:247-53.
Wisler C, Hofer U, Arlettaz R. 2008. Snakes and monocultures: habitat selection and
movements of female grass snakes (Natrix natrix L.) in an agricultural landscape. Jour-
nal of Herpetology 42:337–346.
Wogan G, Vogel G. 2020. Ahaetulla fronticincta (amended version of 2012 assessment).
The IUCN Red List of Threatened Species 2020: e.T192058A175775547. [Link]
org/10.2305/[Link].
Wood DA, Emmons ID, Nowak EM, Christman BL, Holycross AT, Jennings RD, Vander-
gast AG. 2018. Conservation genomics of the Mogollon Narrow-headed gartersnake
(Thamnophis rufipunctatus) and Northern Mexican gartersnake (Thamnophis eques
megalops): U.S. Geological Survey Open-File Report 2018-1141, 47 pp.
Woodruff DS. 2010. Biogeography and conservation in Southeast Asia: how 2.7 million
years of repeated environmental fluctuations affect today’s patterns and the future of
the remaining refugial-phase biodiversity. Biodiversity and Conservation 19(4):919–
941.
Woolrich-Piña GA, Ponce-Campos P, Loc-Barragán J, Ramírez-Silva JP, Mata-Silva V,
Johnson JD, García-Padilla E, Wilson LD. 2016. The herpetofauna of Nayarit, Mexi-
co: composition, distribution, and conservation status. Mesoamerican Herpetology
3(2):376-448.
Worrell E. 1963. Reptiles of Australia. Angus and Robertson, Sydney, 207 pp.
Wright E. 2011. Laticauda colubrina (On-line), Animal Diversity Web. Accessed Decem-
ber 22, 2020 at [Link]
Wu SY. 2004. Behavioral and physiological responses of the snake predators, Dinodon ru-
fozonatum and Macropisthodon rudis to Bufo bankorensis. Master Thesis, National Sun
Yat-sen University, Kaohsiung City, Taiwan, 50 pp.
Wüster W, Peppin L, Pook CE, Walker DE. A nesting of vipers: phylogeny and historical
biogeography of the Viperidae (Squamata: Serpentes). Molecular Phylogenetics and
Evolution. 2008 Nov 1;49(2):445-59.
Wüster W, Chirio L, Trape JF, Ineich I, Jackson K, Greenbaum E, Barron C, Kusamba
C, Nagy ZT, Storey R, Hall C. 2018. Integration of nuclear and mitochondrial gene
sequences and morphology reveals unexpected diversity in the forest cobra (Naja
melanoleuca) species complex in Central and West Africa (Serpentes: Elapidae) Zoo-
taxa 4455(1):68-98.
Yagi AR, Planck RJ, Yagi KT, Tattersall GJ. 2020. A long-term study on massasaugas (Sis-
trurus catenatus) inhabiting a partially mined peatland: A standardized method to
characterize snake overwintering habitat. Journal of Herpetology 54(2):235-44.
Yakovleva ID. 1964. The Reptiles of Kirgisia. Frunse. 272 pp. [in Russian].
Yamasaki Y, Mori Y. 2017. Seasonal activity pattern of a nocturnal fossorial snake, Achali-
nus spinalis (Serpentes: Xenodermidae). Current Herpetology. 36(1): 28-36
Yang JH, Sung YH, Chan BP. 2013.A new species of the genus Opisthotropis Günther,
1872 (Squamata: Colubridae: Natricinae) from Guangdong Province, China. Zootaxa
3646(3):289-96.
Yi H, Norell MA. 2015. The burrowing origin of modern snakes. Science advances, 1(10),
p.e1500743.
705
Aquatic Snakes

Yildirimhan HS, Bursey C, Goldberg SR. 2007 Helminth parasites of the Grass Snake, Na-
trix natrix, and the Dice Snake, Natrix tessellata (Serpentes: Colubridae), from Turkey.
Comparative Parasitology 74:343–354.
Yossefi MR, Nikzad R, Nikzad M, Mousapour A, Ramazanpour S, Rahimi MT. 2014. High
helmintic infection of the European grass snake, Natrix natrix and the dice snake, Natrix
tessellate (Serpentes: Colubridae) from Iran. Asian Pacific Journal of Tropical Disease 4
(Supplement 1):236–S267.
Young BA, Aguiar A, Lillywhite H. 2008. Foraging cues used by insular Florida cotton-
mouths, Agkistrodon piscivorus conanti. South American Journal of Herpetology
3(2):135-44.
Young BA. 1991. The influences of the aquatic medium on the prey capture system of
snakes. Journal of Natural History 25(2):519-31.
Young BA. 2009. The influences of the aquatic medium on the prey capture system of
snakes. Journal of Natural History. 25(2):519-31.
Zagar A, Krofel M, Govedic M, Mebert K. 2011. Distribution and habitat use of Dice snakes
(Natrix tessellata) in Slovenia. Mertensiella 18:207-16.
Zaher H. 1999. Hemipenial morphology of the South American xenodontine snakes: with
a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemi-
penes. Bulletin of the American Museum of Natural History 240, 168 pp.
Zaher H. 1999. Hemipenial morphology of the South American xenodontine snakes: with
a proposal for a monophyletic Xenodontinae and a reappraisal of colubroid hemipenes.
Bulletin of the American Museum of Natural History 240, 168 pp.
Zaher H, Murphy RW, Arredondo JC, Graboski R, Machado-Filho PR, et al. 2019. Large-
scale molecular phylogeny, morphology, divergence-time estimation, and the fossil
record of advanced caenophidian snakes (Squamata: Serpentes). PLOS ONE 14(5):
e0217959. [Link]
Zając B, Mołoniewicz L, Solecki A, Antoł W. 2019. New observation of dice snake (Natrix
tessellata) in Poland. Conference Poster: II Polskie Sympozjum Herpetologiczne Novem-
ber 2019.
Zebe V. 1936. Zur Biologie des Schlangenadlers (Circaetus gallicus (GM)). Berichte des
Vereins Schlesischer Ornithologen 21:33–82.
Zhao EM, Jiang YM. 1981. Studies on amphibians and reptiles of Mt. Gongga Shan, Sich-
uan, China. I. A new species and a new subspecies of snakes from Sichuan. Acta Herpe-
tologica Sinica 5.7 (1981): 53-58.
Zhao EM, Huang MH, Zong Y. 1998. Fauna Sinica, Reptilia Vol. 3 Squamata, Serpentes. Sci-
ence Press, Beijing, China, 522 pp. (In Chinensis)
Zheng Y, Wiens, JJ 2016. Combining phylogenomic and supermatrix approaches, and
a time-calibrated phylogeny for squamate reptiles (lizards and snakes) based on 52
genes and 4162 species. Molecular Phylogenetics and Evolution, 94: 537–547.
Zhong, GH., Chen, WD, Liu Q, Zhu F, Peng PH, Guo P, 2015. Valid or not? Yunnan moun-
tain snake Plagiopholis unipostocularis (Serpentes: Colubridae:Pseudoxenodontinae).
Zootaxa 4020 (2): 390–396.
Zhou Z. and Jiang Z. 2005. Identifying snake species threatened by economic exploitation
and international trade in China. Biodiversity and Conservation 14: 3525–3536.
Ziegler T, David P, Miralles A., Doan VK, and Nguyen QT. 2008. A new species of the
706
Aquatic Snakes

snake genus Fimbrios from Phong Nha–Ke Bang National Park, Truong Son, central
Vietnam (Squamata: Xenodermatidae). Zootaxa, 1729, 37–48.
Ziegler T, Ngo HN, Pham AV, Nguyen TT. Le, M.D. and Nguyen, T.Q. 2018. A new spe-
cies of Parafimbrios from northern Vietnam (Squamata: Xenodermatidae). Zootaxa,
4527 (2), 269–[Link]://[Link]/10.11646/zootaxa.4527.2
Ziegler T, Truong Quang Nguyen, Cuong The Pham, Tao Thien Nguyen, Anh Van Pham,
Mona Van Schingen, Tham Thi Nguyen, Minh Duc Le. 2019. Three new species of the
snake genus Achalinus from Vietnam (Squamata: Xenodermatidae). Zootaxa 4590 (2):
249-269
Zöckler C, Aung C. 2019. The Mangroves of Myanmar. In Sabkha Ecosystems 2019 (pp.
253-268). Springer, Cham.
Zotos S, Stamatiou M, Naziri A, Meletiou S, Demosthenous S, Perikleous K, Erotokritou E,
Xenophontos M, Zavrou D, Michael K, Sergides L. 2021. New evidence on the distribu-
tion of the highly endangered Natrix natrix cypriaca and implications for its conserva-
tion. Animals 11:1077.
Zuri I and Halpern M. 2003. Differential effects of lesions of the vomeronasal and olfacto-
ry nerves on garter snake (Thamnophis sirtalis) responses to airborne chemical stimuli.
Behavioral Neuroscience 117(1):169–183.

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 Literature Cited

708
Aquatic Snakes

Endnotes
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