CHAPT E R 13

Development
Pedro Gnaspini

T
he life cycle o.f a lwrvestman may be clivi,.led into two r_najor_ phases. cmhry_oni c
and postembryonic, which arc separated by the hatchm g ol tht•t•gg (cd osron ).
Postcmbryomc development is charactcnzcd by three phases: larval. nymph crl.
and adult Ouberthic. 1964 ). The lar va hatdtcs from the egg and molt s to the tirst
nymphal in star. The adu lt slagc. dclincd by the capacity to reprodu ce. is l"l'<rl'h ed
afier a variable number of moll s.
Vander Hammen ( 19 7~) discuSSt.x lthe terms used in postembryonic de\'t'lop·
ment of arachnids a nd suggested that the term 11!/IIIJl/ r should be ust·d only wlwn lht•
number of ju venile instars is constant within an or(lcr. Otherwise. when this nu rnbt•r
varies. as in the case of Opiliones. he su ggc.,"Sied the use of the tt·rm ll!fmp/roicf. l'cm ard
and Stockman ( 199 3) compared life ryd t· patt ern s ohser w d a morr g <lini.·rcnt 1rnkrs
of a rachnids wh ile also all emplin g to standardize the tt•rminology of the di f1l·rt· nt
types of immature in stars occurring. They used irrrmr11rm· jrll'l'rr ilt•:s instt·crd of hrn·w•
and jlll'('lli/t•s instead of 11!/llljJirs. Here r follow tht• ll<lllll' S ;rlre<rtly l'Sicrhlis lll'd Hlll t )ll g
harvestmen specialists. namely. /arl'll. II !fill Jill. and adu/1
There is a wide range of liiC cycles among hcrrvcstmt•n. with some srx:rit'S bd ng
stcnochronc and others elrryl..'hrorrc (sec l'hcrplt•r 7 1. Witlr reg;rrd ltr ltmgt•vity 111' crd1rlt s.
there a rc annual species with two to three months or rxrsh:rnbryonk duration ;urd
epheme ral adults. bie nnial species. and tx·rcnnial S(X't'ics with one yt·;rr of pnstt•mhry-
onic duration and ;rdults that live for more tharrthrce yccrrs trdl-rt'llt't'S irr Tc rhk I ~ . 11
This chapter presents information rq~arding embryonic curd fXIS\ernhrynrric tlt-wlop-
ment: phenology crnd re productive biology arc tre<rle<l in l'lwpl t'I'S I ;r ml I 2. respt·c-
livcly.

455
Tab le 13. 1 summary of embryonic and postembryonic development in harvestmen

Nymphal phase

Taxa
?~~~~~~~J 1 Adr~:n~~5t,ion 1 References

CYPHOPHTHALMI
Sironldae
Cyphophthalmus 60-72 Juberthie. 1964,
duricorius {10°C-17"Cl 1965

Parasiro coiffaiti o.55--o.6 66-72 Juberth ie, 1964,

(10~C-1 7"Cl 196 5

Sirorubens 60-90 4-7 5 24- 48 54-60 Juberth ie, 1960b,

(15°C) (10"C-17. C) (10"C-17"Cl 1964,1965

EUPNOI
Phalangiidae
Eumesosoma roeweri 40(- 18"C) Cokendolpher,
1981b
Homolophus biceps -3 Schmoller,1970
Lacinius ephippiatus 240(20"C) 5 3.5 3.5 Pfeifer,1956;
Winkler,1957~

Lacinius horridus 120(20"C); 3- 3.5 4 Pfeifer,1956;

150 Parisot,1962
Lophopilio palpinalis 180 4 3.5 < 1.5 Pfei fer, 1956
Mitopus mario 42-56 5 Tischler, 1967
{20"C-22"Cl
Odie/Ius gal/icus 0.9 1.08 80(11.5"C); 10-60 S{rarely4) 2.5-3.5 (11.5"C); 2-3{16"Cl Juberthie,1957a.
150(21 "C) 4-6(16"Cl 1960a. 1964.
1965; Naisse,
1959
0/igo/ophus tridens 150-180 15(21"C) 5 3-3.5(21 "C) Gueutal.1943;
(30"Cl Pfeifer, 1956;
Naisse, 1959;
Parisot, 1962
Opilio parietinus 0.53--0.56; 1 0.56--Q.82 180; 5 I Henking, 1886;
0.75--0.8 120-150 Faussek, 1892;
Holm, 1947;
Pfeifer, 1956
Phalangium cinereum I -2-3 I Weed, 1897
Phalangium cornutum I
IFaussek, 1892
(ra;~5l I
45--60 (RD
Phalangium opilio 0.67-{).71 28-42 15 2.5( 15-c); Gueutal, 1943;
(30•C);
40 (20"CJ;
I 2(20•C) Todd, 1949;
Pfei~er, 1956;
30 (30"Cl Montz, 1957,
1959; Winkler,
1957; Naisse,
1959; Parlsot,
1962; Ju-
berthie, 1964;
Bachmann&
Schaefer,
1983a

Plarybunus
bucepha/us
I 0.55 I 0.64
I 45 56 (RD;
(20~C-24 °C)
15-30
(20"C-24°C)
5 2-3
(20"C-2 4"C)
lmmel, 1955;
Parisot. 1962
Plarybunus triangularis 0.56-{).62 28-43; 6 7-8 Todd, 1949;
43-SJ(RD Pfeifer, 1956;
Moritz, 1957 •

5clerosomatldae I
Homatenotus 08 130 (10:-17' CI 6Q-90 6 6-10 Juberthie, 1957a,
quadridentatus 1 (10"C-17"C) 1964, 1965
Le1obunum rotundum - 075 I 15(21 "C) Henking. 1886;
Purcell, 1892;
Naisse, 1959
Le!obunum townsend/ I 40(-18"Cl ICoke ndolpher,
1981b
Le10bunum vittatum -2-3 Weed, 1897

DY5PN0 1
I I
Dlcranolasmatidae
D1cranolasma 0.6h 0.45 35- 42(20"C) 6 12-18 24-36 I Gruber, 1996
scaiJrwn (15· c9_?20"Cl I (15"C- 20"C) (15"C-20"Cl

lschyropsahdidae

1 4-4. 5~1SCJ
lsc/Jyropsalls llellwigu 102 (10"C-11 "Cl I Martens, 1965
ISCIJYfOpsalis ltlte!peS 1 0.85 44 {11 .5"CJ 1 6 (ra:ly5) 6-8(11SCJ Juberthie, 1964,
1965; Martens,
1965
lsCIJyropsa/IS noelifera' I 0.9 50(11 .5"Cl 6 I 5-6(1 1.5"C) I 6- 8(1 1.5"C) 1 Juberthie, 1964,
1965; Martens,
1965

(CmllimK'II/
Table 13. 1 continued

Egg size (mm) Embryonic Larval phase Nymphal phase

duration durat ion Number of durat ion Ad ult duration
TaJCa At oviposition At hatching (days) (days) nymphs (months) (m onths) References

lschyropsatis 1.4 58( 11 -S"C) 6 6-7(11 -S"Cl 6-8(1 1.5"Cl Juberthle, 1964.
pyrenaea 1 Martens. 1965

lschyropsalis strandi 1.3 x 1.8 142 2- 211 (S"c-6•c J Jubertl11e. 1965.

15"C-6.5"Cl 1974

/schyropsa/is superba 1 0.85 1.10 6 < 12(11 -S"CJ 6-8(11 .SOC) Juberthle, 1964.
1965

Nemastomatidae
Mitostoma Juberthle, 1964
pyrenaeum 1
Nemastoma 0.85 56-60 30-60 6 >2.5 > 14 lmmel. 1954
quadripunctatum (20"C-24"C) (20"C- 24"C) (20"C-24"Cl (20"C- 24"C)
Paranemastoma 0.65--0.75 27 (16"Cl 6 -2.5{5"C- 21"Cl AVram. 1973
SillII

Nipponopsalididae
Nipponopsalis abeP 6 3 I Miyosi, 1942

sabaconidae
sabacon viscayanus 12( 10"C-1 7"CJ Juberthie, 1964,
1965

Trogulidae
Anelasmocepha/us 0.8-1.1x 47-206 (20"Cl 4 1- 6.5 23-34 Pabst, 1953
cambridge/ 0.3---0.4 (5"C- 17"C) {5"C- 1rCJ
Trogulus nepaeformls 1.3-1.7x 44-226 (20"C) 5 3.5-8.5 12.5-25 Pabst,1953;
0.7-1.0 (5"C-17"Cl (5"C-17"C) Juberthie, 1964
Trogulus tricarinatus 1.1-1.5x 42-207 (20"Cl 5 4-5 (5"C- 17"Cl 15. 5-35.5 Pabst, 1953
0.4-o.s (5"C-17"Cl

LANIATORES
Cosmetidae
Cynortoides cubanus I 0.95 1.1- 1.2 16.5
(25"C);
6 I 2.5
{22"C-24.5"C)
I 30 12o•c-25•cJ I Jubenhie, 1972
27.5(20"C)
Erginulus c/avotibialis I I ,_, 12o·c1;
G~~~~~h~. 1976
1.1 6 4
13(26"Cl I I
Gryne orensis -30(Lab) canals, 1936
Metatibitia - 30 1Lab) canals, 1936
paraguayensis
vonones sayi 20-38 > 36 (5*C-20"C) Cokendolpher &
(S"C-20°C) Jones, 1991

Gonyleptidae
Acanthopachytus 1.02 - 30(Lab) canals, 1936;
acuteatus capocasale &
Bruno· Trezza,
1964
Acutisoma discolor 1.44 1.85 Machado, 2002
Acutisoma longipes 1.44 1.91 45-<>4 Machado &
Oliveira, 1998
Acutisoma 2.01 Ramires &
proxtmum Giaretta, 1994
Acutisoma aff. 1.46 1.89 Machado, 2002
proxtmum
Discocyrtus oliverioi 1.25 1.33 Elpino·Campos
etal., 2001
Discocyrtus S 36; 54 (Lab) Matthiesen, 1985
pectinifemur
Discocyrtus -30 (Labl canals. 1936
prosptcuus
Goniosoma 1.52 1.97 24-60 21-41 Willemart &
albiscriptum Gnaspini, 2004a
Goniosoma 1.58 2.11 Machado, 2002
genicu/atum
Goniosoma spelaeum - 18 Gnaspinl, 1995
Goniosoma sp
2. 1-2.3
1.51 1.91
30-60 > 60 6
'" Machado, 2002
Hernandaria -30(Labl Canals, 1936
scabncula
HOioscotolemon 40 4-5 18(10°C-17"C) Juberthie, 1964,
quenlhaci 1 11 1•c-13•cJ (10"C-17"CJ 1965

Pachylotdes tllOrellt - 30(Lab) cana ls, 1936

Pachylvs 1-1.3 1.6 37 (20"C) 2-10(20°C) 7 4 (18"CJ 36-48 (20"C) Juberthie&

qumamavrdensrs Murioz·Cuevas,
1971;MuriOZ·
Cuevas,
1971a,b.c. 1973

ICou!luurd)
Table 13.1 continued

- ~

Parapachy/oides
fontanensis
Pygophalangodus
canalsi
egg stze(mm)

IAt o~:-ffionl At ha~hing

Embryonic
duration
(days)

-JO(Lab)

-30 (Labl
Larval phase
duration
(days)
Number of
nymphs
Nymphal phase
duration
<months)
Adult duration
(month s)
I Refe<ences
Cana ls, 1936

Canals, 1936

Manaosbiidae
Zygopachylus 15-20 I Rodriguez &
albomarginis Guerrero, 1976

Phalangodidae
Pseudobiantes 0.7 Miyosi, 1941
japonicus
Scotolemon doriae 30(20"Cl 4-6(10"C-17"C) 24-42 Juberthie, 1964,
(10"C-17"Cl 1965
scotolemon lespesi 1 55- 70 5 5 (10"C-1 7"C) 24- 30 Juberthie, 1964.
(11.5"C) (10"C-17"Cl 1965
Scotolemon lucasi 68 (11 "C- 13"Cl 18(10"C-17"Cl Juberthie, 1964

Podoctidae
Leytpodoctis oviger 0.8-1 .1 Martens, 1993b

Stygnopsidae
Hoplobunus boneti 2.0 I Mitchell, 1971

DaShes indicate lack of Information. Species with complete descriptionS of embryOnic development are marked wi th an asterisk. some Gata on the duratiOn of developmental phases
are followed by the temperature In which the eggs and/or nymphs were reared. RT .. room tempera ture (without any explanation in the origiMI data about the exact climatic condi
tlon). "Lab" indicates that the study was carried out In the laboratory, but no temperature was recorded. wtlen not stated, data were obtained from field experiments
1. Jubefthie (1964) noted that the development of this species was primarily the same as that of other studied species
2. Miyosl did not clearly state the number of nymphs, but 6 were deduced on the basis of his descriptions
 Development 461

A BRIEF HISTORICAL ACCOUNT

Balbiani (1872), who studied Plmlmryium comutum (Phalangildac). W<lS the fi rst to
describe the embryology of harvestmen. Un til the middle of the twentieth century.
knowledge of the embryonic development of harvestmen was mainly based on the
studies of Balbiani (1872) a nd Faussck (1892). who studied Oplllo paril'liuus (Pha-
langiidae). and Schimkewitsch (1898). who studied a n uniden tilied phalangiid.
Other arliclcs from the end of the nineteenth century. includin g Henking (1886)
with 0. parietinus and Leiobwwm rotwulwn (Sclcrosomatidac) a nd Purcell (1892)
with L. l!emisp/raericwn. did no t provide a n adequate account of the embryology of
the group, and some of their data should be considered with ca ution. Holm ( 194 7)
conducted a more detailed study of 0. jmril'tinu s and provided ligu res of its main
stages. while Dawydoff (1949) provided an early review on embryology. but lillie
was recorded about harvestmen. From the 1950s to the 1970s scvcnrl studies (e.g ..
Pabst. 1953: Pfeifer. 1956: Morit':.:. 1957. 1959: Winkler, 1957: juberthie, 1964:
Mutio:.:-Cuevas. 197lb. 1973: Avram. 1973) and com parative reviews (D.T. An-
derson. 1973: Yoshikura. 1975) were published that provided the basis for the cur-
rent knowledge on harvestman embryology.
Concerning postembryonic development. the lirst descriptive studies \\"Cre prob-
ably those from Miyosi (1941, 1941). published in Japanese. but mostly neglected.
The study by Gueuta l (194 3) dealt main ly with the duration of each nymph. Pabst
(195 3) and Immel {1954. 1955) present ed more [Link] surveys. but the !irst ex-
tensive reference is the pu blished thesis of ju berthic ( 1964 ). Mtcrw;ml a st.:rics of
mainly descriptive and comparative articles fo llowed (Edgar. 19 7 1: Murloz-Cucvas.
197la.c: Goodnigh t & Goodnight, 1976: Gnaspini. 1995: Gruber. 1996)
The most extensive developmental studies considered species of Eupnoi and Dys-
pnoi from the Nort hern Hemisphere: [Link] tores. mostly from the Neotropical re-
gion. have only recently been studied in more deta il (Table \3. 1 ). Unti l now. approx-
imately 3 5 spcdes (2 Cyphophtha lmi. I I Eupnoi. 12 Dyspnoi. ;md I 0 L;miatores)
have been the subject of ontogenctir studies. This number would increase to at least
76 species (3 Cyphophthalrni. 26 Eupnoi. 19 Dyspnoi. ;md lX J,;rni<1torcs1 if we
added assort ed information l'rorn artil·lcs not dealing spcci lkally \Vith dcvcltlpnK·nt
or from comments within studies on dcvdopmcnt.

EMBRYONIC PHASE

Embryonic development begins just aficr oviposition and ends at the lirnt· of t•do-
sion. when the larva ha tches. Since la rval organogenesis is the lc1st pl1c1St' d1rring em-
bryonic dcvcloprncnt. some a uthors prdcr to nmsidcr thl' last embryonic plwsc as
endin g when th e l<1rva has formcll inside the eggshell ;rnd till' lw;rrt 11l'~ins to hL"al
(Mur'im:-Cucvas, 19 7 J b).
462 Dcvclopmcnt

Eggs
Balhh111i I I X7 2) dt•snilwd I ht• l'ggs or 1'111111111!/illlll I'Orlllll/1111 as [Link] g 1.2
111111 in dioum·h·r. heing sp lwroidal :md light yellowish in eolor in the beginning. t1ml
l;lll'l' lwcolllillf! [.!I'<Jyish lll't'<lll-"L' or pi)!IIICill<l li on of ! Ill' embryo. 1\rcnrding IO Bal -
hi;llli 11 Xi21. tll t'l'gg b -"• •rn••• ••dt•d by two m t•mhr:•nes. an ou ter thin chorion und
an inner [Link] dtdlim· lllt'llllmult'. lluhn I 19-1 7 1 iJiso subsequently dcll't:tcd a
~ln l r lllre re.-;t• mbli11g •••nil'l'tlpy le
1\l:uJy eggs ilrt' sphl'roid;d: others lll<IY be ovoid or dong<~led cmd whitish to
grayis h whit t· inwl11r t luberthie. 1Yh-ll. In somt• species the egg batd1cs arc sur-
roundn1 h~· 11ll ll'tiS t Figurt· 1 ~- L \ 1. hut not in others (Figurt• 1 3. 1B). a nd the eggs

Figure 13.1. (A) Clutch of lporangaia pustulosa (Gonyleptidae) on the undersurface of a leaf in southeastern
Brazil. composed of eggs surrounded by an abundant mucus coating in different embryonic stages. (B) Clutch of
Acurisoma proximum (Gonyleptidae) showing recently laid eggs without mucus coating on a rock in southeastern
Brazil (photos: B.A. Buzatto). {C) Eggs from Coniosoma spelaeum {Gonyleptidae) near hatching on a cave wall in
southeastern Brazil. Note that eyes. legs. and color patching of the body can be observed through the egg chorion.
(D) View of eggs: a larva (l) with its curved and short legs: a larva molting into a first nymph (M), in which the legs
still attached to the exuvium can be seen; and two first nymphs (N) from Coniosoma spelaeum still attached by
the anal lobe to the exuvium (photos: P. Gnaspini). Scale bars • lO mm.
 Development 463

Figure 13.2. Embryonic development of the gonyleptid Pachylus quinamavidensis {Gonyleptidae). Roman
numerals indicate the embryonic developmental phase. and Lindicates the "larval phase." beginning after
the first heartbeat. For the same stage, S.A. V, and D indicate side, anterior, ventral, and dorsal views of
embryo, respectively. Mouthparts are formed on the 21st day. while eyes. heart, and digestive caecum are
formed on the 22nd day. Modified from Muiioz-Cuevas {1971b)

may be laid inside a subst ratum (mostly in soil I or on surl"aees (see Chapter 111. Egg
size varies among groups and species. gcncr<llly between ll. 5 and 1.3 mm. At till' hl'-
ginning of embryonic development th e l'gg.s [Link] in size and weight by absorbing
wa ter from the air and/or !"rom the hygroscopic lllLU.:US that surrounds them. and
they therefore need a high rdatil'C humidity (Henking. I SNh: ]ubcrth k'. 19 h-l.
19h5). The size of eggs. both when laid and <J I"tcr growth by absorbing watn. is rom-
piled in Table I 3.1.

De scription of development
Holm (J 947 ). Moritz (1 1)57 ). and Ju bcrthic [19(1-1 . 19h 'l) USl'i.l different
sc hemes to divide embryonic development into phases. ]uherthie"s sc heme. tlw most
commonly used in recent tk sai ptions. divides embryonic developmen t intn lin'
phases (Figure 1 3.1): ( 1) cleavage: ( 2) form;~tion of tlw germ lmnd: l 3lllll'l<~meriz;~ ­
tion of thcprosoma . when there are imporlilll\ cdlullu· migration s, and ceph;1lir iilld
amd plates and appcnd<Jge buds iiPIX'ar: (4) inversion. when ahtlolllinal somitt· ~.
mouth. und eye-primordium appear. ll·gs liikc till' lin:ll lengt h. the hod~· \\"a ll i~
form ed . and the genital platl' migrat es anteriml bct\\"eentlw lq! coxae: t'll la n ·a l
orgt~nogenesis. including fornwtion of the digcslivl' din'rtindu1n. tlw dilkn· nt i<lli•lll
of eyes a nd heart illld, ilner the lirst he01rtbea t. the l<1r va l period tli1r dt·t;Jil ~ . ~n·
Ch apt er 2). The d ural ion of et~ch phasl" is known only liw tht• fullowi11g tlll'l'l' S [Wl"il'.~:
464 Develol}ment

the phal;u1giid Cldicllu.\ !JIIIIinrs 1h. lJ. ::i. 2 '5, <Jlld 3'5 days. rcspcc.:L ivcly ). th e ischyrop-
s;d idid hdi.I/I"III'SII/i.. /ul!'iJWS (-!.h. '5. 10. a mi I '5 days). a nd the gonyleptid Parl!y/us
!/llilltllllfll"idcmis l '5. ~- 3. H. and 1h dc 1ysl (rd .crenl"cs in Table I 3. 1 ).

Duration
J:mbryo nic tkw lopllll'lll gen era lly l<lkcs around 30 to llO days (Table 1 3. 1). but
grc;J tly dqx·nds on Sl'<lSO /l and tl'lllpenltun:. For ins t;mce. the egg.s or Opilio j)(ll"il'-
lillll:i dit· if thL·y a rt• mai ntainL'"<I <tt ruomtcmpenllurc sim:e they need to pass through
<l cold ll"in ter. <llHI rurthcr ckvclupmcn t is achieved when they arc placed upon th e
ground 1FcJussL·k. I H()2 1. Eggs "r Jn<my SJ>edl'S from temperat e regions hibcrn cJ te.
ll"hirh b the CCJ SL' l"i1r tht• phalangiids J\liWjm ... morio. C>lino/op/russpp.. Opi/io fHirif'lil ms.
;JlHIJ'/w/tiii!JIWII opilio. the srlerusonwtid Uio/nomm m111111111111. the nemastomatid NI'-
IIIIISI/111/tl lu!Julm•. c1nd Sl"\'cra l ischyropsa lidids of the genus /sdJ!Jropsn/is (Kiist ner.
1 ~3'5 h : l'li:ifer. 19::ih: Phillipson . 1959: r.. la rtens. 1965: Bachm ann & Schaefer.
I YH 3a I. In coni r<JSI In 1hese spec ies . trogulid s reproduce yea r-round. Development
may take l"rmn '5 luX \\'l'eks l"or s unHncr eggs (lunt.' -August) to 12 to 1 5 weeks for hi-
ht'rll il ting winter t•ggs (Febr uary ) (Pabst. 195 3 ). However. winter and summer eggs
dt·wlop <lithe s;um· r<1te wlll"n artilidally ma in tained at 20°C.
Obli g; Jtury dh1pcmse induced by chcmgcs in temperature was described in a
l;en nan popul ation of i\ 1. moria in which eggs a rc laid during the late s ummer and
hatchin g occ urs in tht• spring (Tischll•r. 19 6 7). Embryologica l development proceeds
at summer temperatures until it is interrupted a t a genetically predetermined stage
DL•vclupment remains [Link] until low temperatu res trigger completion of the
second phc1 se of embryogenesis. The last phase takes place only with th e advent of
w:lrn) spring weather. Th us the tirst and third phases take place only at higher tem-
pewturcs. wherl'ilS the Sl'<:ond phase demands lower t cmper;o~tures. In this way un-
timely hatching in th e fc1ll and win ter is prevented. and nymph s do not appear until
<Jdcqutlle l'o od is avc1ilable in the sprin g.
The effect of tem perature on egg development has been reported for sevcml
spccil's (sec a lso Table I 3. I ). Development in Odil'/lus gal/icus Iukes 162 days at 6°C,
\46 days at 7°C. <Jnd 84 days at 1 I . S°C: minimum and maximum lethal tempera-
tures are. respct·th•cly. 0 °C and 20°C. but no mortality is detected between 6°C and
I I.5°C (Jubcrthie. \ 960a ). In Par/I!Jius quiuauwvide11sis (Gonyleptidae) development
takes 70 days at I 2°C and 3 7 days a t 20°C: development is abnormal below soc and
above 2 S°C: no development occurs at soc (Mu ii oz~Cuevas. J 97 lb). In PammmrtiS-
romtl sillii (Nemastomatidae) th e hat ching rate is reduced at higher temperatures
l"rom about9S% above u~ oc to O'X, at 20°C-2JOC ( Avr<~m. 1973). Development of
the eggs of /~ Oj1ilio is strongly deluyed at lower temperatures a nd not even possible a t
I 0°( with in 2 50 days. while taking 129 days at J soc. 40 days at 20°C, and 30 days
at 2soe, a nd at ll.8 °C the development was estimated to be zero (Bachmann &
Schaefer. 1983a). Jubert hie (1 964) and Avram {19 73) presented further discussion
about the influence of temperature and humidity on embryonic development.
 Development 465

Anomalous development
Anomalous specimens have been recorded and experiment ally produced severn!
times. Gynandromorphs a nd intersexual specimens were recorded in i\11itojms mario
{Cokendolpher & Sissom. 1988) and Plwlangimu opi/io (Blaszak. 1968) and in the
sclerosomatids Gagrellula ferruginnl (Suzuki. 1980b). G. mo111mm (Tsurusaki.
1982b). Leiobw111111 g/oboswn (Suzuki, 1980a), and J\lklmWJHI yrmulis (Tsur usaki.
1982b). Andromorphy in fema les is a rarer phenomenon and W<IS reported only for
NI'IIWSlOIIW deuligerum (Chcmini. 19M4 ). Teratological sped mens ht1ve been
recorded and experi mentally produced in Otlidlus uallit'us tsec Figures 2.6e.f in
Chapter 2).

LARVAL PHASE

Close to hatchin g. in cases where the chorion is tnmsparcnL. it is possiblt• to see the
developin g larva (Figure 13.2). The larva is the li rst frl'e inslilr afier cdosiun. pre·
sen ling several incomplete fea tu res if compart.'<l with the li rst nymph s tscc lli:\l.·us·
sion in Ju berthie. 1965). The legs typic<llly bend around the body in onkr to tit insid<.·
the rou nd eggs during development (Figure I 3.2) . In Eupnni. Dyspnoi. and Lani<1 ·
torcs the larva hatches in a more mature stag<.• than in l'yp hophthulmi Hnd quickly
molts into the lirst nymph (Table I 3.1 ). In contra:-;\. in l'yphnphtln ilmi lunching or·
curs earl ier in larval forma tion. a nd the hatch ling :-;pends fi1ur to .st'\'t'll day:-; as a
larva before molt ing into the lirst nymph tlu!x·rthic. I lJ(1()b. I I.Jh5J.
The larvae of Cyphophthalmi. Eupnni. wul l)yspnoi ar<.' l'l[Uipped wit h Olll.' or
two fronta lly located egg teet h (<II thl' median anterior margin of the prosnm<~ . be-
tween the chelicerae) tlmt hl'ip in bre<~kin~ the eggshel l. Egg tt•t•th are i<lt'king in
nymphs. Balbiani (I M72J was the !irst to rct'ord un egg tooth in <1 l;~ te h;wvt•:-;tnwn
embryo. Larvae of Eupnoi (e.g.. Astmlwnus !lrtlllator. !\lt'!/11/mmts sliwfmw. (Jdicl/us ual-
lkus. 0/inolopfm ... lridl'us. l'aroli!lolop/111.~ 11!/I'I'Siis. 1'/wftlll!fiwu OJii/io. 1'1111.11111!111/S lrillll·
!llllnris. Homalmmm; tflllldricfmltiW.~. and /,d,l/1111111111 spp.: JuhtTihk. l lJ57a. llH1-I
Nclissc. 19 59: Edgar. I 97 J ). and most Dyspnoi (e.g .. Ptmllll'IIIIISIOIIW sil/ii and /)iarl·
110/asmu sm/Jrum: 1\Vr<llll , l lJ73: Crubcr. llJl)()) lwvc om· qe.g totllh. l)ysp1H1i 11f th t•

ftlmil y lsc hyropsa lidid ac. such <IS /sdi,IJrtiJISrdis lwcipcs. I. uorl(li·m. /. Jl!f/'1'11/l!'tl. ;11u\ I
lu·llwiuii lonmll'i. have two egg teeth (julx·rt h il'. 1 ~Jfl+l. Tlw lc1r\'a of the siro11id .\'iro
ru/Jt•us also has two egg lt,"Cih !lulx·rthit•. llJhOb. llJh-IJ. No t•gg ll'l'lh tk.'t'Ur in thl'
cosmctids Er!!imllus dm·oli!Jittlis (Coodnight & t:oodnight. llJ i' h ! <llld \iuuuws SII!Ji
(Cokcndolphcr & Iones. l lJ~ I). or in I Ill' gonyleplill /'lldi!Jiu .~ (/llillmlltll'irlmsis
(Mtuio:t.-Cucvas. \ 1)7) b). /\bscncc ol' egg teet h nwy be the ruk in l.<ll1i<l!t1rt'S. <~I k·;~st
among Grassatorcs.
466 Develol)ment

NYMPHAL PHASE

Till' numlwr of ny mph s varies <llllong the diiTercn l harvestmclll species studied
tsonwtinws t'WII within tlw Silll'll' spcries). being six in most n1ses. bu t ra nging from
li>ur to l'ight in others tT<J bk l ~ .I ) . 'l'lu· last ny mphal stage is generally called the
··suhadull ." 'l'hL·numbt·r of nymphs is imkpcndcnt of the lim1l size of the aduh har-
wstma n tju herthil·. l lJh-ll.

Growth
During posh:mbryonk dc\'l'lnplllL'nt cm·h molt results in a larger a nimal. Since
body sizL' may dL')X'JHI on thL' amoun t of food previously ingested. length and width
measurL'IllL'rl iS arL' ge nerally not appropriate for d(x:umenting growt h. In con trast.
thL· apJX'JUia ges <lfl' adctlll<lte strtteturcs for dtx:umenti ng growth and have been used
inmostcomp<ll'illiw studies. The pe rcenl<lgc growth of some struct ures between con-
or
SL'l' Utivl' inStill'S ol' various species harvestmen is given in Table 13.2. The three
trogulids studied by P<Jbst 11 Y53) show a change in body length of approximately
I 30'Y.,;Jt eadl moh. being a lillie largL•rdurin g the molt from larva to first nymph. Ad-
diliomdly. it .sho uld be mentioned that at least in Goniosonw SJH.'/r/('11/11 (Gonyleptidae).
there is no d ear allometric growth for the <!ppcndage segments. that is. they keep the
S<HllL' proportion through the successive nymphal and adult st<lges (Gnaspini. J 995).
Juberthie ( JlJ6-l). refe rring to a "hlrV<Jl step." noted especially rapid growt h in
phalangiids. sderosomatids. ;md ischyropsalidids as a whole. The growth rate de-
dines aflt'rward ttl less than half the larva/ tirst nymph growth rate. In the fi na l molt
to adulthood . he also noted that fema les of H011w/(' IIOII!S lJIIac/riclrlllall!s (Scleroso-
lll<lli(I<Je) lla\'c the same growth rate a s nymphs. but males have un increased growth
rate. These c harm:tcristics c;m be observed in other species (Table 1 3.2). The differ-
Ctlli<ll nHllc/f\..' tmdc growth tll e~1ch insl<lr in G. SpC'irlw ll! can be observed in Figure
I ~.3. Another interesting dnmlcleristic of th is species is that the growth rate from
lirst to second nymph is mu ch larger than the subseq uen t ones.

Morphological and morphometric nymphal changes during growth

l'\'hmy structu res develop or inc rease in size and/or com plexity during postem-
bryonic development. as the fo ll owing examples illustrate. Among Eu pnoi and Dys-
pnoi the setae of the legs generally increase in number and size and may change
s hape. The shape of the body and ocularium may a lso change with age (A. MUller,
1915: Miyosi. 1942: )uberthic. I 957a. 1964). The nymphal s tages of Phalangium
opilio can be separated from one another by the number of tarsal joints (Bachmann
& Schaefer. 198 Ja). In Ptmmcmastoma si/lii the pseudoarticulalion of femora in-
creases (Avram. 1973). and in Dil:rmwlasma sca/mm1 (Dicranolasmatidae) the hood
progressively grows (Gruber. 1996). In Gonyleptidae and other Lanlatores the arma-
ture of pedipalps and legs. as well as the number of tarsomercs. increases {Muiioz-
Cuevas.l97la:Gnaspini. 1995).
Secondary sexual c haracters arc highly variable within harvestmen and may in-
 Development 467

Table 13.2 Examples of percentage growth of different struct ures with age among harvestmen

...., port toN1 toN2 toN3 toN4 toNS t ON6 to Adult Reference

Phalangium opilio Body 163 122 158 137 119 135F, 119M Naisse. 1959
Homalenotus 300 90 90 90 90 90 90F, 130M Juberthie,
quadridentatus 1964
lll 500 80 80 80 80 80 80F, 130M
l lll 300 70 70 70 70 70 ?OF, 180M
LIV 400 80 80 80 80 80 80F, 140M
Pedipatp 200 80 80 80 80 80 80F, 100M
Leiobunum calcar LIV 156 146F, 143M Edgar, 1971
Leiobunum longipes LIV 158 174 169 156 175F, 155M Edgar, 1971
Leiobunum politum LIV 161 150F, 144M Edgar, 1971
Leiobunum rotundum Body 150 117 164 135 113 171F,129M Naisse. 1959
Leiobunum vittarum 150 156 161F, 150M Edgar, 1971
Oicranolasma scabrum 171 124 123 132 133 126F,133M 137F,142M Gruber, 1996
Lll 229 125 125 126 134 128F, 135M 135F, 141M
l lll 173 123 126 133 135 128F,133M 138F, 143M
LIV 196 128 126 131 138 130F, 134M 141F, 147M
Body 136 127 130 132 134F, 133M 145F, 143M
Nemastoma 204 131 124 135 123 131 tmmel,1954
quadripunctatum
lll 244 127 124 137 110 105 153

llll 189 127 133 129 131 140

LIV 240 125 135 143 104 115 142
Body 91 152 118 131 139 115
Paranemastoma sillii 219 131 134 104 138 169 133 Avram, 1973
lll 255 151 120 109 136 152 145
llll 226 126 140 101 145 133
LIV 230 122 138 109 141 163 138
Body 100 138 120 117 116 150 170
Nipponopsa/is abei 1 Body 143 155 129 140 Miyosi, 1942
Goniosoma spe/aeum 139 142 126 131 102 Gnaspini
1995
160 137 120 128 106
LIJI 170 142 135 124 129 104
LIV 170 137 135 121 125 105
Pedipalp 169 133 139 129 129 109

Aquestionmarlc:(?)indicates in formationnotprovided
Ntto N6lndlcate nymphal stages from the first to t11e sixth . Ll to UV indicate legs from t to IV.
1. Miyosi did not clearly state the number of nympt1s, but si x r1ympt1swere deduced on tile basrs ollusclesu rpt rons He (fl(f not pi OVI(fc ti.H. II<H til< , , u,~:, •d
thi rd nymph; thus the percentage growth w as added in th e table from tile seconcl to the fourth nymph
468 Developmem

60 140
Leg I Leg II
50
I o'
120

100 hi
~·
40

o'~·
80
30
o• 60
20

e 10
40

20 .
g
1 2 3 4 5 sA A 2 3 4 5 sA A
£Oil
Figure 13.3. Growt h of
.,c::
....l
80 Leg III 120 Leg IV
appendages du ring the
• Q' 100
development of Goniosoma 60 10'
0' ~·
spelaeum (Gonyleptidae) for males 80

o• ~·
(light gray) and females (dark
gray). White boxes represent
nymphsinwhichsexisnot 40 60
recognizable externally. Each box
represents the average z standard
0 40
deviation,whilet helinesrepresent 20
the upper and lower observed
intervals. The numbers 1- 5 =first
20 0

to fifth nymphal instars:sA =

[Link]; A= adult. Modified from
Gnaspini (1995) 2 3 4 5 sA A 2 3 4 5 sA A

dude color and the shape of body. chelicerae. pedipalps. genital opening. anal plate.
sternites. tergitc.:s. and legs. among ot hers (sec Chapters 4 and 12). In Eupnoi and
Dyspnoi these sex ua l charact ers commonly appear in the penultimate instar (or
subadult ) a nd seldom in the ant epenultimate instar (Juberthie, 1964). In Laniatores
secondary sex ua l dwracters me generally conspicuous and appear in the ante-
penultimate instar (rvlulioi:-Cucvas. 19 7 1a: Gnaspin i. 1995).
Although most Opiliones have a single claw on each leg. Laniatores have com-
plex claws on legs Ill and IV (see Chapter 4). Among lnsidiatores (Travunioidea)
legs Ill and IV have a complex claw. which is generally three-branched. but may
have more side branches with a si ngle insertion at the tarsus. A transformation se-
ries from a three-bra nched sta te to a complex peltonychium was discussed by Hunt
and Hick man (199 3) on the basis of species of Lomanel/a (Triaenonychidae). Among
the Grassatores legs Ill and IV have two single claws independently inserted into the
tarsus, although the claws may fu se at the base. Accessory structures between the
 Development 469

two tarsal claws that differ in the nymphal and the adult phases arc present in Gras·
satores. According to .Mulioz-Cuevas ( 19 7lc). in all nymphal stages except the
subadu lt. two typical "juvenile structures" arc found ventrally in both third and
fourth leg tarsi: one projection similar to a third tars."ll claw. called a tJSI'IIIIcmwhium.
and a fleshy projection. called an tlrolium (Figures I 3.4t\.B: see also Chapter 21.
These structures grow during nymphal development but arc not observed in either
the last nympha l stage (the subadult ) or in the adult (sec r\'llllloz-Cucvas. 197 la .c:
Gnaspini. 1995: Figures 13.4C.D). Conversely. a third stru ctu re in most Gonylcp-
toidca (Cosmetidac. Cran aidae. Gonylcptidae, Manaosbiidm·. and Stygnid;u,·). l'a lk·d

Figure 13.4. Apex of tarsus IV of the cosmetid Cosmetus variolosus {ph01os: P. Gnaspini). (A- B) nymph
lateral and frontal views. (C-O) Adult: lateral and dorsolrontal views. AR = arolium: PS = pseudonychium:TC
= tarsal claw; TP = tarsal process.
470 Development

<I !Ursa/ pro!'1'SS. llliiY gn>11· graduully wilh each molt. but develops fully only in adults
! \lu(wz - i."UL'\ " <1~. 19 / I;Lct. i\ dors;~l projeL·tion on the apex of the last tarsomcrc
gL"IWr<dly he<1rs a long <IIHillllliSU<il se1<1 iFigure 1 ~.4C,lJ) . These differences during
gn>l\"lh haw lKTII \\"ell dorumentt·d only in l'[Link] 1/llillllllllll'idi'IISis (MUJioz-
Cu L'\"<! S. I 971 <II illld (;olliiiSOII/11 spclacwn l(;nuspini. \1.)1.)5). Alth ough not spccili-
cally studying dcvdr>pmL'Ill. 1\l<irlens ami Schwendingcr ( IIJ9t)) illustrated on-
n>pl>did species in \\"hich 1he juwnile structures <Ire Jacking in the adults, whereas a
\\"l'll-de\·eloped <JroliUIIl Gill be observed in the leg IV or a ju venile. In addition
,\ liyosi l 1LJ-t I recorded ··;~ strut·ture simih1r to what is cal led an arolium·· in Psl'!ldo-
1

l>ianl l'S jr1poni!"11S t Eped<ulid;~e). lienee tht• jul'enilc structures probably occur in all
Crass<1torcs. \\"hcn:as the tars;il pron.-ss only Ot"l'Urs in ccrt<lin fam ilies. t\mong ln-
sidiutort·s di!Tcrenn:s in t;u·s<JI structures <1ppear <II difl"erent ages. that is the tarsus
;d sodwngcs through ontogeny te.g .. J\ lu1ioz-Cucvas& Vachon. 19 79). However. the
strucwres invol1·ed nwy be diiTcrent. since the juvenile structures just discussed for
l;r<ISSiilOrt·s nwy be l<~cking illl10ng lnsidiutorcs (e.g.. Briggs. 1969. 1971a. for
nymph s with <llld without <~rnlium. respectively ).

Molting
The posture assumed during molting varies with the grou p. In most groups, es-
pecicdly the long-legged species. the cmimals hang from a substratum for molting.
Eupnoi species lwng by all four pairs of legs. while La niatores usc only those with an
<1rolium (pt~irs Ill tmd IV). which enhcmec substrat e adherence, even on glass (Ju-
berthie. \LJ/2). During molting the individual d isplays periodic body con tract ions
t~nd extends the legs while hanging from the exuvium. to which th ey are li nked by
the <111<1! operculum (PtJrisot. 1962: Edgar. 1971: Gnaspini. 1995: Figure 13.5).
When the process is complete. the animal hangs from the ceiling by ils legs and de-
tarhcs from thl' cxuvium. Afterward many species (except Pha lan giidae) take t he ex-
uvium in their pcdipalps a nd manipulate it with t heir mouthparts. wh ich probably
allows recovery of water from the exuvia l fluid. The larva a nd nymphs of Dicm-
1/o/mmra sca/Jmm may han g. bu t generally rema in on the ground. moving upward.
releasing their legs vertically. <tnd ··silting" on the anal lobe. which detaches last
from the cxuvi um : fim1lly t he animal moves forward . standing on its legs (Gruber.
1996). Nymphs of Siro rub1•ns bu ild a molting c hamber made w ithin a fi ssure of t he
substratum. which they block vvith debris (Juberthic, 1960b).

Duration
The duration of postembryonic development in different species is summarized in
Table 13.1. It shou ld be noted that the duration of the nymphal phase is not directly
proportional to the linal size of lhe adult Uuberthie. 1964). The postembryonic phase
generally occurs within several months. a lthough lengthier periods (over two to three
years) have been recorded in Cyphophthalmi. Dicranolasmatidae. a nd some Gonylep-
tidae. not to mention specialized cave species Uuberthie. 1 964: Gnaspini. 1995:
Gru ber. 1996).
 Development 471

Figure 13.5. (A) Nymphs of Acutisoma proximum (Gonyleptidae) and (B)jussara sp. (Sclerosomatidae)
molting on vegetation in southeastern Brazil (photos: B.A. Buzatto). Scale bars= 10 mm

Tempera ture a lso play~ a rok· in till' duration of postembryonic tk·wlnpnll'nt . For
in sw ncc. Bachmann and S~.·h;.Jder ti9X ~a I sho\\'ed that no indi\'itl u tl l ~~r 1'/mlml_rJiwu
opilio re;xhcd adu lthood at 10 °(. Tlll'y also l'l'l'Orded tlminimum nf 7~ ti;Jys at 1 ~ 0 ('
and a minimum of 5X d<l)'S ill 20° (. Wl't'd 11 HY71 l'l'l'Orded l. !'io/mmuu l't'lltrko'SIIIIJ
hibcnwting as nymphs (ont·-t hird th1.· l"ull-!,t~'0\\' 11 size I.

ADULT PHASE

It is widely considered that <Jdult lHI!"\'eSIIlll'll do not molt. Indeed. thi s is t h~.· c<~~l' liw
most groups wi1h a genital O[X'I'l'Ulum. in which thl' opl'l'ndum gl'ner;dly iippl'iii'S
{<J)rcmly open) only in adults (l'.g .. Julwrthie. llJ~/al. llmt'l'l'er. [Link]-t'lll'l'ilS
11 ")71<1) a nd Gn aspini ( l lJlJ~)lwtk"ed tlwt in so me < ; rass:~ton·s species the gcnit:~l
opercu lum. wh ich was fu s1.·d tot h1.· ilO(Iy during tht• early nymph:~! ~tagt'S. bn:onws
fn.'C in the su lmduiL sugl!estin g that the suh<ttlul! is ;tlrt·<~dy s~.·xuall _t · lli<Hun·.
Cnaspini et al. (201>41 rccnnk-tl the presl'tll'l' of ;rcurnpk•lt' pcnb in ~uh;rduh l;r;l~­
satnrcs and observed a subadult mak·cupulirling. Tngl'thl'r 1\'ilh lltl'lllorpholoi!ka l
study of severa l spcdes. (;naspini e t al. 11111141 proposl'd that Cril~~atorl·~ h<JI'l' a
unique life t.:ycle mnong ilradmids wit h l \\'O instilrs in thl' adult phaSI:: I he su hadult
and till' atlu lt. ;\ simih1r lift• cyd1.· is kno\\'11 orrly inmaylli~,·s !Ephl'IIH'fuph·ra l. 11 hich
47 2 Development

also reprmlt tn· as suhad ul ts. This hytx>thcsis deserves further testin g. a nd detailed
studiL'S of <I nH JI"t' indu sivc llltlllbt•r of st>edes :t re net:essary.
'l'hl' duratiun ol' the <tdult phase l'or difti:rent spcdes is li sted in Table \ 3. 1. In
summ ;u·y. Eupnoi gt·twndly hol\'c a short lit!: span or a few month s. whereas other
~roups. c..'S ]X'c..·i:t Jiy Cyphuplllh<thni and Laniatorcs. may live for severa l years as
adult s. t'\ 't'll those specic..·s s ubjcctL"<Ito ha rs h winter conditions.

CONCLUDING REMARKS

JJc..·spitc th t• few spt•dcs studied . developmen t of harvestmen appears to be fairly ho·

mogencou s in different taxa . Al thou gh morphology and egg size va ry greatly. em·
bryologka J dc\'clopnwnt generally takes one to two months. or a little longer in the
l'itSl' (Jr tt·ntpc..·ntle species !h it\ piiSS through win ter egg diapause. The postembryonic
phase. induding it lar va. it varied number or nymph s. and one or two reproductive
Slitgt·[Link] Iitke from a few mo nth s to scwral years.

ACKNOWLEDGMENTS

Theilu th or is 11 rec ipient of a rcsc t~rch grcmt fro m FAPES P (00/04686A) and of a re·
search li:llowship from CNPq ( 30032()/ 94·7). Dr. A. A. Ribeiro a nd E. Mattos a ].
lowed t~nd facilitated the usc or the electron m it"roscopc (LNIE·IBUSP). I am deeply
indebted to C. E Le rche. who helped with the tran slation of papers in Germa n . to
i\·1. R. I-I <Ira . who hciJX>tl with the translation of papers in Japanese. to B. A. Buzatto
ror providin g photogn1phs. and to S. Hoencn . G. Machado. and two reviewers for
their critical review or th e manuscript