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com

CHAPTER 21

Phylum Arthropoda
Subphylum Crustacea:
Crabs, Shrimps, and
Their Kin

Photo by Simon Richards, courtesy of J. Yager

C
rustaceans are one of the most popular invertebrate groups, even among
nonbiologists, for they include some of the world’s most delectable gour-
met fare, such as lobsters, crabs, shrimps,1 and goose barnacles. There
are about 72,000 described living species of Crustacea, and probably five times
that number waiting to be discovered and named. They exhibit an incredible
diversity of form, habit, and size (Figure 21.1). The smallest known crustaceans
are less than 100 μm in length and live on the antennules of copepods. The largest
are Japanese spider crabs (Macrocheira kaempferi) with leg spans of 4 m, and giant
Tasmanian crabs (Pseudocarcinus gigas) with
carapace widths of nearly a half-meter. The
heaviest crustaceans are probably American
lobsters (Homarus americanus) that, before
the present era of overfishing, attained
weights in excess of 20 kg. The world’s larg-
est land arthropod by weight (and possibly
the largest land invertebrate) is the coconut
crab (Birgus latro), weighing in at up to 4 kg,
and the largest freshwater invertebrate is
the Tasmanian giant freshwater crayfish
(Astacopsis gouldi) or lutaralipina in Tasma-
nian Aboriginal language. Crustaceans are
found at all depths in every marine, brack-
ish, and freshwater environment, including
in pools at 6,000 m elevation (fairy shrimp
and cladocerans in northern Chile). A few
have become successful on land, the most
notable being sowbugs and pillbugs (the
terrestrial isopods) and species in several
groups of Brachyura (crabs). Beginning
1
When most people hear the word “shrimp,” they think
of edible shrimps, two crustacean groups nested within
the order Decapoda (in the suborders Dendrobranchiata
and Pleocyemata). However, the term “shrimp” is
applied to a number of long-tailed crustaceans, many
not closely related at all to decapods. So, in this general
sense, there are fairy shrimps, tadpole shrimps, mantis
shrimps, etc. In much of the English-speaking world,
the word “prawn” is used for the edible shrimps, thus
eliminating some of the confusion.

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 3/8/22 1:09 PM
Courtesy of Gary McDonald
Courtesy of P. Fankboner Courtesy of A. Kerstitch
Courtesy of A. Kerstitch
(C)

(H)

Arnstein Rønning/Wikipedia/CC BY 3.0

(K)
© Larry Jon Friesen

(M)
Courtesy of A. Anker

(G)
(B)

(E)

Courtesy of J. Olesen Courtesy of A. Kerstitch

© Vladimir Levantovsky/Alamy Stock Photo

(J)
© Larry Jon Friesen

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660 Chapter 21

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Brusca 4e
(A)

(D)

(L)
(F)

(I)
Photo by D. Williams, courtesy of J. Yager © Larry Jon Friesen
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(N) (O)

Courtesy of Gary McDonald

Courtesy of Gary McDonald
(Q)

(P)

© Larry Jon Friesen

(R)
Courtesy of C. Holliday

Courtesy of A. Anker
(S)
(T)
Courtesy of David McIntyre

Courtesy of A. Kerstitch

FIGURE 21.1 Diversity among the Crustacea.

(A–D) Classes Remipedia, Cephalocarida, and Branchiopoda.
(A) Morlockia ondinae; note the remarkably homonomous
segmentation of this swimming crustacean (Remipedia). (M) A Hawaiian regal lobster, Enoplometopus (Achelata).
(B) Lightiella monniotae, from New Caledonia (Cephalocarida). (N,O) Two unusual amphipods (Amphipoda). (N) Cystisoma,
(C) A tadpole shrimp (Notostraca) from an ephemeral pool. a huge (some exceed 10 cm), transparent, pelagic hyperiid
(D) A clam shrimp (Diplostraca), carrying eggs. (E–Q) Class amphipod. (O) Cyamus scammoni, a parasitic caprelloidean
Malacostraca. (E) A fiddler crab, Uca princeps (Brachyura). amphipod that lives on the skin of gray whales. (P) A male
(F) A giant Caribbean hermit crab, Petrochirus diogenes and female cumacean; note the eggs in the marsupium of
(Anomura). (G) A hermit crab (Paragiopagurus fasciatus) the female (Cumacea). (Q) Ligia (Isopoda), the rock louse;
removed from its snail shell (Anomura). (H) A coconut crab, Ligia are inhabitants of the high spray zone on rocky shores
Birgus latro (Anomura), climbing a tree. (I) Emerita analoga, worldwide. (R) A mysid Siriella sp. (Mysida) with a parasitic
a Pacific mole crab (Anomura). (J) A pelagic lobsterette, epicaridean isopod (Dajidae) attached to its carapace
Pleuroncodes planipes (Anomura). (K) A cleaner shrimp, (Western Australia). (S,T) Class Thecostraca. (S) Acorn barna-
Lysmata californica (Caridea). (L) Alienaxiopsis clypeata, cles, Semibalanus balanoides (Thoracica). (T) Lepas anatifera
aBrusca 4e lobster-shrimp from Bali, Indonesia (Axiidae).
coral reef (Thoracica), a pelagic barnacle hanging from a floating timber.
BB4e_21.[Link] (Continued on next page)
12/20/2021

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 (U) (V)

© Nature Picture Library/Alamy Stock Photo

From G. A. Boxshall and R. J. Lincoln 1987.

Philos Trans R Soc B 315: 267–303
FIGURE 21.1 (continued) Diversity among the Crustacea. (U) Class
Copepoda; the calanoid copepod Gaussia, a common planktonic genus.
(V) Class Tantulocarida, Deoterthron, parasitic on other crustaceans.

early in the twenty-first century, negative effects of Despite the enormous morphological disparity seen
ocean acidification on crustaceans began to be recog- among crustaceans (Figures 21.1–21.20), they display
nized, especially on the larval stages and subadults a suite of fundamental unifying features (Box 21A). In
(see Selected References section). an effort to introduce both the diversity and the unity
Crustaceans are commonly the dominant organisms of this enormous group of arthropods, we first pres-
in aquatic subterranean ecosystems, and new species of ent a classification and synopses of the major taxa. We
Brusca
these 4e
stygobionts continue to be discovered as new
BB4e_21.[Link]
caves are explored. They also dominate ephemeral
pool habitats, where many undescribed species are
12/20/2021 BOX 21A C
 haracteristics of the
known to occur.2 And crustaceans are the most wide- Subphylum Crustacea
spread, diverse, and abundant animals inhabiting
the world’s oceans. The biomass of one species of 1. Body composed of a six-segmented head, or cephalon,
Euphausiacea, the Antarctic krill (Euphausia superba), and a long postcephalic trunk; trunk divided into two
has been estimated at 500 million tons at any given more or less distinct tagmata (e.g., thorax and abdomen)
time, probably surpassing the biomass of any other in all but the remipedes and ostracods (Figure 21.2)
group of marine animals (and rivaling that of the 2. Cephalon composed of (anterior to posterior) ocular or
world’s ants, summing up all their species!). In fact, protocerebral segment (lacking appendages), antennular
krill are the dominant fished species in the Southern segment (deutocerebral), antennal segment (tritocerebral),
mandibular somite, maxillulary somite, and maxillary
Ocean in terms of catch weight. The range of mor-
somite; one or more anterior thoracomeres fused with the
phological disparity among Crustacea far exceeds head in some members of some classes (e.g., Remipedia
that of even the insects. Many species of crustaceans and Malacostraca), their appendages forming maxillipeds
are threatened by environmental degradation; over 3. Cephalic shield or carapace present (highly reduced in
3,000 are listed on the IUCN Red List, and about anostracans, amphipods, and isopods)
two dozen are protected by the U.S. Environmental 4. Appendages multiarticulate, uniramous or biramous
Protection Agency. Crustaceans are also among the
5. Mandibles usually multiarticulate limbs that function
most common invasive invertebrates, and well over as biting, piercing, or chewing/grinding jaws
100 invasive species have established themselves in
6. Gas exchange by aqueous diffusion across specialized
marine and estuarine waters of North America alone. branchial surfaces, either gill-like structures or
Because of their taxonomic diversity and numeri- specialized regions of the body surface
cal abundance, it is often said that crustaceans are the 7. Excretion by structures derived from nephridia
“insects of the sea.” We prefer to think of insects as (e.g., antennal glands, maxillary glands)
“crustaceans of the land.” And indeed, there is now 8. Both simple ocelli and compound eyes in most taxa
very strong phylogenetic evidence that insects arose (not Remipedia), at least at some stage of the life cycle;
from a branch within the Crustacea and this larger compound eyes often elevated on stalks
clade is known as Pancrustacea (or Tetraconata). 9. Gut with digestive ceca
2
One study of ephemeral pools in Northern California discov- 10. With nauplius larva (unknown from any other arthropod
ered 30 probable undescribed/unnamed crustacean species subphylum); development mixed or direct
(King et al. 1996).

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(A)

(B)

FIGURE 21.2 General crustacean external morphology:

a crayfish (Malacostraca: Astacidea). (A) External anatomy. Note
the fully developed carapace covering the cephalon and thorax.
(B) The typical malacostracan tail fan (ventral view). Note the
position of the anus on the telson.

then discuss
Brusca 4e the biology of the group as a whole, draw- we herein recognize as superclasses. Multigene phy-
ing examples from its various members. As you read
BB4e_21.[Link] logenetics also supports the division of Altocrustacea
this chapter, we ask that you keep in mind the general
5/12/2021 into two major clades, which have been named Multi-
account of arthropods presented in Chapter 20. crustacea and Allotriocarida, the latter including Hexa-
poda (insects and allies). See Crustacean Phylogeny at
the end of this chapter for further details.
Classification of the Crustacea
Crustaceans have been known to humans since ancient
times and have provided us with sources of both food CLASSIFICATION OF CRUSTACEA
and legend. It is somewhat comforting to carcinologists SUPERCLASS OLIGOSTRACA
(those who study crustaceans) to note that Cancer, one CLASS OSTRACODA Ostracods
of the two invertebrates represented in the zodiac, is a
crab (the other, of course, is Scorpio—another arthro- SUBCLASS MYODOCOPA
pod). Our modern view of Crustacea as a taxon can ORDER MYODOCOPIDA (e.g., Cypridina,
be traced to Lamarck’s scheme in the early nineteenth Euphilomedes, Eusarsiella, Gigantocypris,
century. He recognized most crustaceans as such but Photeros, Polycope, Skogsbergia, Vargula)
placed the barnacles and a few others in separate
groups. For many years barnacles were classified with ORDER HALOCYPRIDA (e.g., Conchoecia)
molluscs because of their thick, calcareous outer shell. SUBCLASS PODOCOPA
Crustacean classification as we know it today was
more or less established during the second half of the ORDER PODOCOPIDA (e.g., Baffinicythere,
Cypris, Candona, Darwinula, Limnocythre ,
nineteenth century, although internal revisions con-
Loxoconcha, Sclerocypris)
tinue. Martin and Davis (2001) presented an overview
of crustacean classification, and readers are referred ORDER PLATYCOPIDA (e.g., Cytherella)
to that publication for a window into the labyrinthine
ORDER PALAEOCOPIDA Almost entirely extinct;
history of this subphylum. Multigene phylogenetics
living genera are Manawa and Puncia.
strongly supports the division of Crustacea into two
major lineages, Oligostraca and Altocrustacea, which

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 664 Chapter 21

CLASS MYSTACOCARIDA Mystacocarids, with ORDER CUMACEA Cumaceans (e.g., Campy­

a single family (Derocheilocarididae) and 13 species laspis, Cumopsis, Diastylis, Diastylopsis)
(e.g., Ctenocheilocaris, Derocheilocaris)
ORDER TANAIDACEA Tanaidaceans
CLASS BRANCHIURA Fish lice, or argulids. A single (e.g., Apseudes, Heterotanais, Paratanais, Tanais)
family (Argulidae) (e.g., Argulus, Chonopeltis, Dipteropeltis,
ORDER MICTACEA Mictaceans (e.g., Mictocaris,
Dolops)
etc.). Some researchers separate out the family
CLASS PENTASTOMIDA Tongueworms. Two orders, Hirsutiidae as a distinct order (Bochusacea, contain-
numerous families (e.g., Cephalobaena, Linguatula, ing Hirsutia, Montucaris, and Thetispelecaris).
Pentastoma, Waddycephalus). In some recent classifi­
ORDER SPELAEOGRIPHACEA Spelaeogriphace­
cations combined with the Branchiura in the clade
ans. Four described living species (Potiicoara brazil­
Ichthyostraca.
ienses, Spelaeogriphus lepidops, and two species
SUPERCLASS ALTOCRUSTACEA of Mangkurtu), and two known fossil species (the
Carboniferous Acadiocaris novascotica and the
clade MULTICRUSTACEA
Upper Jurassic Liaoningogriphus quadripartitus)
CLASS MALACOSTRACA
ORDER THERMOSBAENACEA Thermosbaena­-
SUBCLASS PHYLLOCARIDA c­eans (e.g., Halosbaena, Limnosbaena, Monodella,
Theosbaena, Thermosbaena, Tulumella)
ORDER LEPTOSTRACA Leptostracans or
nebaliaceans (e.g., Dahlella, Levinebalia, Nebalia, ORDER ISOPODA Isopods (sea slaters, rock lice,
Nebaliella, Nebaliopsis, Paranebalia) pillbugs, sowbugs, roly-polies)
SUBCLASS HOPLOCARIDA SUBORDER ASELLOTA (e.g., Asellus, Eurycope,
Jaera, Janira, Microcerberus, Munna)
ORDER STOMATOPODA Mantis shrimps (e.g.,
Echinosquilla, Gonodactylus, Hemisquilla, Squilla) SUBORDER CALABOZOIDEA (Calabozoa)
SUBCLASS EUMALACOSTRACA SUBORDER CYMOTHOIDA
SUPERORDER SYNCARIDA Syncarids SUPERFAMILY ANTHUROIDEA (e.g.,
Anthura, Colanthura, Cyathura, Mesanthura)
ORDER BATHYNELLACEA (e.g., Bathynella)
SUPERFAMILY CYMOTHOOIDEA
ORDER ANASPIDACEA (e.g., Allanaspides,
(including the families Aegidae, Anuropidae,
Anaspides, Paranaspides, Psammaspides)
Barybrotidae, Cirolanidae, Corallanidae,
SUPERORDER EUCARIDA Cymothoidae, Gnathiidae, Protognathiidae,
and Tridentellidae)
ORDER EUPHAUSIACEA Euphausiaceans,
or krill (e.g., Bentheuphausia, Euphausia, INFRAORDER EPICARIDEA (e.g., Bopyrus,
Meganyctiphanes, Nyctiphanes) Dajus, Hemiarthrus, Ione, Pseudione)
ORDER DECAPODA Crabs, shrimps, lobsters, etc. SUBORDER LIMNORIDEA (e.g., Limnoria,
Keuphylia, Hadromastax)
SUBORDER DENDROBRANCHIATA Penaeid
and sergestid shrimps (e.g., Lucifer, Penaeus, SUBORDER MICROCERBERIDEA
Sergestes, Sicyonia) (e.g, Atlantasellus, Microcerberus)
SUBORDER PLEOCYEMATA All other deca- SUBORDER ONISCIDEA (e.g., Armadillidium,
pods (shrimp, lobster, crabs, etc.; classified in 10 Ligia, Oniscus, Porcellio, Trichoniscus, Tylos,
infraorders: Caridea, Stenopodidea, Brachyura, Venezillo)
Anomura, Astacidea, Achelata, Axiidea,
SUBORDER PHORATOPIDEA (Phoratopus)
Gebiidea, Glypheidea, Polychelida)
SUBORDER PHREATOICIDEA (e.g., Mesam­
SUPERORDER PERACARIDA Ten orders, all of
phisopus, Phreatoicopis, Phreatoicus)
which brood their embryos in a pouch
SUBORDER SPHAEROMATIDEA
ORDER MYSIDA Mysidans or opossum shrimps
(e.g., Acanthomysis, Hemimysis, Mysis, Neomysis) SUPERFAMILY SEROLIDEA (Serolis)
ORDER STYGIOMYSIDA Stygiomysidans SUPERFAMILY SPHAEROMATOIDEA
(e.g., Lepidomysis, Stygiomysis) (Ancinus, Paracerceis, Sphaeroma)
ORDER LOPHOGASTRIDA Lophogastridans SUBORDER TAINISOPIDEA (Pygolabis,
(e.g., Gnathophausia, Lophogaster) Tainisopus)

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SUBORDER VALVIFERA (e.g., Arcturus, Idotea, ORDER POECILOSTOMATOIDA

Saduria) Poecilostomatoids (e.g., Chondracanthus,
Erebonaster, Ergasilus, Pseudanthessius)
ORDER AMPHIPODA Amphipods (beach hop-
pers, sand fleas, scuds, skeleton shrimps, whale ORDER SIPHONOSTOMATOIDA
lice, etc.) Siphonostomatoids (e.g., Clavella, Nemesis,
Penella, Pontoeciella, Trebius)
SUBORDER AMPHILOCHIDEA (e.g., Alicella,
Ampelisca, Byblis, Haustorius, Ipanema, CLASS THECOSTRACA Barnacles and their kin
Kroyera, Leucothoe, Liljeborgia, Lysianassa, (about 367 genera and 2120 species)
Phoxocephalus, Stenothoides)
SUBCLASS FACETOTECTA Monogeneric
SUBORDER COLOMASTIGIDEA (Hansenocaris): the mysterious “y-larvae,” a group
(e.g., Colomastix) of marine nauplii and cyprids for which adults
are unknown
SUBORDER HYPERIIDEA (e.g., Cystisoma,
Hyperia, Phronima, Primno, Themisto, Vibilia) SUBCLASS ASCOTHORACIDA Two orders
(Laurida, Dendrogastrida) of parasitic thecostracans
SUBORDER HYPERIOPSIDEA
(e.g., Ascothorax, Dendrogaster, Laura, Synagoga,
(e.g., Hyperiopsis, Podosirus)
Zoanthoecus)
SUBORDER PSEUDINGOLFIELLIDEA
SUBCLASS CIRRIPEDIA Cirripedes, the barnacles,
(e.g., Pseudingolfiella)
and their kin
SUBORDER SENTICAUDATA (e.g., Ampithoe,
SUPERORDER THORACICA True (stalked and
Caprella, Crangonyx, Cyamus, Elasmopus,
acorn) barnacles, e.g., Balanus, Chthamalus,
Gammarus, Grandidierella, Hadzia, Hyale,
Conchoderma, Coronula, Lepas, Pollicipes,
Hyalella, Maera, Melita, Niphargus, Pontogeneia,
Scalpellum, Tetraclita, Verruca.
Pseudamphithoides, Talitrus)
SUPERORDER ACROTHORACICA Burrowing bar-
ORDER INGOLFIELLIDA Ingolfiellidans
nacles. Two orders, Cryptophialida and Lythoglyptida
(e.g. Ingolfiella)
(e.g., Cryptophialus, Trypetesa)
CLASS COPEPODA
SUPERORDER RHIZOCEPHALA Thirteen families
SUBCLASS PROGYMNOPLEA of highly modified parasitic barnacles (e.g., Hetero­
saccus, Lernaeodiscus, Mycetomorpha, Peltogaster,
ORDER PLATYCOPIOIDA Platycopioids
Sacculina, Sylon)
(e.g., Antrisocopia, Platycopia)
CLASS TANTULOCARIDA Deep water, marine para-
SUBCLASS NEOCOPEPODA
sites (e.g., Basipodella, Deoterthron, Microdajus)
SUPERORDER GYMNOPLEA
clade ALLOTRIOCARIDA
ORDER CALANOIDA Calanoids
CLASS CEPHALOCARIDA Cephalocarids
(e.g., Bathycalanus, Calanus, Diaptomus,
(e.g., Chiltoniella, Hampsonellus, Hutchinsoniella,
Eucalanus, Euchaeta)
Lightiella, Sandersiella)
SUPERORDER PODOPLEA
CLASS BRANCHIOPODA Branchiopods
ORDER CYCLOPOIDA Cyclopoids
SUBCLASS ANOSTRACA Fairy shrimps and brine
(e.g., Cyclopina, Cyclops, Lernaea, Notodelphys)
shrimps (e.g., Artemia, Branchinecta, Branchipus,
ORDER GELYELLOIDA Gelyelloids (e.g., Gelyella) Linderiella, Streptocephalus)
ORDER HARPACTICOIDA Harpacticoids SUBCLASS PHYLLOPODA
(e.g., Harpacticus, Hase, Longipedia, Peltidium,
ORDER NOTOSTRACA Tadpole shrimps
Porcellidium, Psammus, Sunaristes, Tisbe)
(Lepidurus, Triops)
ORDER MISOPHRIOIDA Misophriods
ORDER DIPLOSTRACA The “bivalved”
(e.g., Boxshallia, Misophria)
branchiopods
ORDER MONSTRILLOIDA Monstrilloids
SUBORDER LAEVICAUDATA Flat-tailed clam
(e.g., Monstrilla, Stilloma)
shrimps (e.g., Lynceus)
ORDER MORMONILLOIDA Mormonilloids.
SUBORDER ONYCHOCAUDATA Clam shrimps,
Monogeneric: Mormonilla
cladocerans (water fleas), and cyclestherians

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 666 Chapter 21

INFRAORDER SPINICAUDATA Clam larval stage (suppressed or bypassed in some groups),

shrimps (e.g., Cyzicus, Imnadia, Leptestheria, and often a series of additional larval stages.4
Limnadia, Metalimnadia)
INFRAORDER CLADOCEROMORPHA Superclass Oligostraca
Water fleas and cyclestheriids Class Ostracoda
PARVORDER CYCLESTHERIDA Body segmentation reduced, trunk not clearly divided
Monotypic: Cyclestheria hislopi into thorax and abdomen; trunk with 0 to 3 pairs of
PARVORDER CLADOCERA Water fleas limbs; abdomen greatly reduced; caudal rami (furca)
(e.g., Anchistropus, Daphnia, Diaphonosoma, present; gonopores on lobe anterior to caudal rami; car-
Leptodora, Moina, Polyphemus) apace bivalved, hinged dorsally and closed by a central
adductor muscle, enclosing body and head; carapace
CLASS REMIPEDIA Remipedes. One living order,
highly variable in shape and ornamentation, smooth
Nectiopoda (e.g., Cryptocorynectes, Godzillius,
or with various pits, ridges, spines, etc.; most with 1
Lasionectes, Pleomothra, Speleonectes, Xibalbanus).
simple median naupliar eye and sometimes weakly
Sometimes combined with the Hexapoda as the
clade Labiocarida. stalked compound eyes (e.g., in Myodocopida); adults
with maxillary and (in some) antennal glands; males
with distinct copulatory limbs (Figure 21.3).
Synopses of Crustacean Taxa Ostracods include about 13,000 described living spe-
cies (and nearly 65,000 fossil species) of small bivalved
The following descriptions of major crustacean taxa crustaceans, ranging in length from 0.1 to 2.0 mm,
will give you an idea of the range of diversity within although some giants (e.g., Gigantocypris) reach 32 mm.
the group and the variety of ways in which these suc- They superficially resemble clam shrimps in having the
cessful animals have exploited the basic crustacean entire body enclosed within the valves of the carapace.
body plan. However, ostracod valves lack the concentric growth
rings of clam shrimps, and there are major differences
in the appendages. The carapace is usually penetrated
Subphylum Crustacea by pores, some bearing setae, and is shed with each
Body generally composed of a 6-segmented cepha- molt. A good deal of confusion exists about the nature
lon or head (the ocular segment, plus 5 limb-bearing of ostracod limbs, and homologies with other crustacean
segments)3 and multisegmented postcephalic trunk; taxa (and even within the Ostracoda) are unclear—this
trunk divided into thorax and abdomen (except in confusion is reflected in the variety of names applied
remipedes and ostracods). First segment of cephalon by different authors. We have adopted terms here that
(ocular segment) bears the eyes and protocerebrum; allow the easiest comparison with other taxa.
subsequent segments bear first antennae (antennules), Ostracods possess the fewest limbs of any crustacean
second antennae, mandibles, maxillules, and max- class. The four or five head appendages are followed by
illae (see Table 20.2); one or more anterior thoraco- zero to three trunk appendages. Superficially, the (sec-
meres may fuse with the head (e.g., in Remipedia and ond) maxillae appear to be absent; however, the highly
Malacostraca), their appendages forming maxillipeds modified fifth limbs are in fact these appendages. The
(secondarily modified for feeding); cephalic shield or trunk seldom shows external evidence of segmentation,
carapace present (secondarily lost in some groups); although all 11 postcephalic somites are discernable in
with antennal glands or maxillary glands (excretory
4
nephridia); both simple ocelli and compound eyes in Segments of the thorax are called thoracomeres (regardless of
whether or not any of these segments are fused to the head), whereas
most groups, at least at some stage of the life cycle; appendages of the thorax are called thoracopods. The term “pereon”
compound eyes stalked in many groups. With nauplius refers to that portion of the thorax not fused to the head (when such
fusion occurs), and the terms “pereonites” (= pereomeres) and
“pereopods” are used for the segments and appendages, respectively,
3
For a very long time the arthropod ocular segment, which has no of the pereon. Hence, on a crustacean with the first thoracic segment
appendages but houses the protocerebrum and the eyes, was called (thoracomere 1) fused to the head, thoracomere 2 is typically called
the “acron,” a term borrowed from annelid development when the pereonite 1, the first pair of pereopods represents the second pair of
two phyla were thought to be very closely related (i.e., the Articulata thoracopods, and so on. Be assured that we are trying to simplify,
hypothesis). However, the demise of the Articulata hypothesis, and not confuse, this issue. Also, we caution you that the homology
the realization that arthropods are ecdysozoans (and annelids are of the thorax and abdomen among the major crustacean lineages
spiralians) undermined the concept of acron homology between the is probably more reverie than reality; the segmental homologies of
two phyla. Further, unlike the arthropod ocular segment, the “acron” the thorax and abdomen have not yet been unraveled among the
in annelids (usually called the prostomium) commonly bears both crustacean classes. For summaries of naupliar development across
eyes and antennae, as well as a variety of palps, tentacles, and sense the group and larval features in all crustaceans, see Martin et al.
organs. Thus, the term “ocular segment” (or “opthalmic segment”) (2014). In most crustacean species the number of body segments is
best describes the first, eye-bearing segment of Crustacea and other fixed, but in at least two groups (notostracans and remipedes) the
arthropods. segment number can vary within a given species.

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Shrimps, and Their Kin 667
(A)

Median eye

Furca

Antennule

Zenker’s Organ

Maxillule
Male copulatory
appendage

Sixth limb
(walking leg)

Antenna Seventh limb

(cleaning limb)

Mandibule Fifth limb (male) Fifth limb (female)

(maxilliped/clasper) maxilliped

(B)

Furca
Antennule

Seventh limb

Sixth limb

Maxillule Antenna

Fifth limb Mandible

FIGURE 21.3 Anatomy and diversity in the class Ostracoda. (A) Anatomy of
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(C) Antennula Bellonci organ Right valve FIGURE 21.3 (continued) Anatomy and diversity
in the class Ostracoda. (C) Internal view of
Metapolycope (Myodocopa), left valve removed.
(D) The highly ornate Eusarsiella (Myodocopa); side view
and edge view, showing the ornate shell. (E) Examples
of genera from the major living ostracod groups
(arrows point anteriorly). A: Vargula (Myodocopa,
Myodocopida). B: Polycope (Myodocopa, Halocyprida).
C: Cytherelloidea (Podocopa, Platycopida).
D: Saipanetta (Podocopa, Podocopida). E: Neonesidea
(Podocopa, Podocopida). F: Propontocypris (Podocopa,
Podocopida). G: Macrocypris (Podocopa, Podocopida).
H: Ilyocypris (Podocopa, Podocopida). I: Centrocypris
(Podocopa, Podocopida). J: Candona (Podocopa,
Podocopida) K: Cyprinotus (Podocopa, Podocopida).
Courtesy of A. Cohen

L: Darwinula (Podocopa, Podocopida). M: Baffinicythere

(Podocopa, Podocopida). N: Loxoconcha (Podocopa,
Podocopida). O: Pterygocythereis (Podocopa,
Podocopida). P: Cyprideis (Podocopa, Podocopida).
Q: Semicytherura (Podocopa, Podocopida).
Antenna Maxillule Fifth limb Furca
R: Hemicytherura (Podocopa, Podocopida).
S: Sahnicythere (Podocopa, Podocopida).
T: Terrestricythere (Podocopa, Podocopida).

(D) (E)

Volume 17, pp. 1–14. Elsevier, Amsterdam; D. J. Horne et al. 2002. In J. A. Holmes and A. R. Chivas (Eds.), The Ostracoda:
From J. Rodriguez-Lazaro and F. Ruiz-Muñoz. 2012. In D. J. Horne et al. (Eds.), Developments in Quaternary Sciences
Applications in Quaternary Research. [Link] © American Geophysical Union
Both courtesy of A. Cohen

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Shrimps, and Their Kin 669

some taxa. The third pair of trunk limbs bears the gono- brooding embryos. And in a remarkable twist of nature,
pores and constitutes the so-called copulatory organ. a number of bioluminescent fishes have been shown to
Ostracods are one of the most successful groups of obtain their luciferase from their ostracod prey—a pro-
crustaceans. They also have the best fossil record of cess called kleptoprotein bioluminescence. Fertilization
any arthropod group, dating from at least the Ordovi- in these luminescent ostracods is internal, with sperm
cian, and an estimated 65,000 fossil species have been being transferred by a spermatophore. The male eighth
described (a bit over 20,000 fossil trilobites have been limb functions as a copulatory organ, which is enlarged
described). Most are benthic crawlers or burrowers, but and modified for grasping. In females the eighth limb is
many have adopted a suspension-feeding planktonic reduced and includes a pair of knobs, probably grasped
lifestyle, and a few are terrestrial in moist habitats. One by the males. The spermatophore appears to act as a mat-
species is known to be parasitic on fish gills—Sheina orri ing plug to block other male copulations. All offspring
(Myodocopida, Cypridinidae). Ostracods are abundant in one brood are thus fertilized by the same male, and
worldwide in all aquatic environments and are known females can store sperm and produce multiple broods
to depths of 7,000 m in the sea. Some are commensal on from one insemination.
echinoderms or other crustaceans. A few podocopans
have invaded supralittoral sandy regions (members of
the family Terrestricytheridae), and members of sev- Class Mystacocarida
eral families inhabit terrestrial mosses and humus. Two Body divided into cephalon and 10-segmented trunk;
principal taxa (ranked as subclasses here) are recog- telson with clawlike caudal rami; cephalon charac-
nized within the Ostracoda: Myodocopa and Podocopa. teristically cleft; antennae and mandibles biramous;
Myodocopans are all marine. Most are benthic, but antennules, maxillules, and maxillae uniramous; first
the group also includes all of the marine planktonic trunk segment bears maxillipeds but is not fused with
ostracods. The largest of all ostracods, the planktonic cephalon; no carapace; gonopores on fourth trunk seg-
Gigantocypris, is a member of this group. Myodocopans ment; trunk segments 2–5 with short, single-segment
include scavengers, detritus feeders, suspension feed- appendages (Figure 21.4A).
ers, and some predators. There are two orders: Myodo- There are only 13 described species of mystacocarids, 8
copida and Halocyprida. in the genus Derocheilocaris and 5 in Ctenocheilocaris. Most
Podocopans include predominantly benthic forms; are less than 0.5 mm long, although D. ingens reaches
although some are capable of temporary swimming, 1 mm. The head is marked by a transverse “cephalic
none are fully planktonic. Their feeding methods constriction” between the origins of the first and second
include suspension feeding, herbivory, detritus feed- antennae, perhaps a remnant of primitive head segmen-
ing, and parasitism. The Podocopa are divided into tation. In addition, the lack of fusion of the cephalon and
three orders: the exclusively marine Platycopida, the maxillipedal trunk segment, the simplicity of the mouth
ubiquitous Podocopida, and the Palaeocopida. The Pal- appendages, and other features have led some workers
aeocopida were diverse and widespread in the Paleo- to propose that the mystacocarids are among the most
zoic but are represented today only by the extremely primitive living crustaceans. These attributes may, how-
rare Punciidae (known from a few living specimens ever, be related to a neotenic origin and specialization for
and from dead valves dredged in the South Pacific). interstitial habitats. Mystacocarids are marine, intersti-
One of the most remarkable examples of biolumines- tial crustaceans that live in littoral and sublittoral sands
cence in the animal kingdom occurs among ostracods throughout the world’s temperate and subtropical seas.
in the family Cypridinidae. Worldwide, over half of the Their rather vermiform body and small size are clearly
known 300 species in this family are luminescent, prob- adaptations to life among sand grains. Mystacocarids are
ably all to deter predators, but one clade (that occurs thought to feed by scraping organic material from the
only in the Caribbean) also uses their luminescence for surfaces of sand grains with their setose mouthparts.
complex courtship displays much the same way fireflies
do, only by ejecting pulses of light into the sea in near
darkness. The luminescent reaction involves cypridinid Class Branchiura
luciferin and luciferase that, together with mucus, are Body compact and oval, head and most of trunk covered
ejected separately through specialized nozzles from by broad carapace; antennules and antennae reduced,
separate secretory cells in the upper lip, and they mix the latter sometimes absent; mouthparts modified for
in the seawater. As a result, light is extracellular, being parasitism; no maxillipeds; thorax reduced to 4 seg-
produced external to the body of the ostracod. Virgin ments, with paired biramous appendages; abdomen
females respond to species-specific male displays by unsegmented, bilobed, limbless, but with minute caudal
swimming to intercept a signaling male in the water rami; female gonopores at bases of fourth thoracic legs,
column, but without luminescing themselves. These male with single gonopore on midventral surface of last
mating arenas are equivalent to underwater leks. Mated thoracic somite; paired, sessile compound eyes and 1 to
females immediately move to the seafloor and begin 3 median simple eyes (Figure 21.4M).

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 670 Chapter 21

(A)

Courtesy of J. Olesen
50 µm

(C) (D) (E)

(B) (F)

(G)

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Shrimps, and Their Kin 671

(H) (I)

Courtesy of E. Peebles
(J)

(L)

From P. H. C. Corgosinho et al. 2018. ZooKeys 766: 1– 38./CC BY 4.0

(K)

100 µm

(M)

(hooked sucker)

Courtesy of Gary McDonald

FIGURE 21.4 Anatomy of the classes Mystacocarida, with egg sacs. (I) A female siphonostomatid copepod
Copepoda, and Branchiura. (A) General anatomy and SEM (Trebius heterodonti, a parasite of horn sharks in California)
of the mystacocarid Derocheilocaris. (B) General anatomy with egg sacs. (J) A siphonostomatid copepod, Clavella
of a cyclopoid copepod. (C–E) General body forms of (C) a adunca, showing extreme body reduction; this species
calanoid, (D) a harpacticoid, and (E) a cyclopoid copepod. attaches to the gills of fishes by its elongate maxillae. (K)
Note the points of body articulation (dark band) and the Notodelphys, a wormlike cyclopoid copepod adapted for
position of the genital segment (shaded segment). Roman endoparasitism in tunicates. (L) The harpacticoid copepod
numerals are thoracic segments; Arabic numerals are Hase talpamorphicus (confocal laser scans). (M) Branchiura:
abdominal segments; T = telson. (F) An elaborately setose Argulus foliaceus (drawing and photograph), a branchiuran
calanoid copepod adapted for flotation. (G) A poecilosto- that parasitizes fishes. Note the powerful hooked suckers
matid copepod, Ergasilus pitalicus, ectoparasitic on cichlid (modified maxillules) on the ventral surface. (G after V. E.
fishes. (H) A female siphonostomatid copepod (Caligus sp.) Thatcher. 1984. J Crust Biol 4: 495–501.)

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 672 Chapter 21

The Branchiura comprise about 230 species of ecto- (lungs, nasal passages, etc.) of their host. Body highly
parasites on marine and freshwater fishes. The anten- modified, wormlike, 2–13 cm in length. Adult append-
nules generally bear hooks or spines for attachment ages reduced to 2 pairs of head appendages, lobelike and
to their host fish. The mandibles are reduced in size with chitinous claws used to cling to host. Body cuticle
and complexity, bear cutting edges, and are housed nonchitinous and highly porous. Body muscles some-
within a styliform “proboscis” apparatus. The maxil- what sheetlike, but clearly segmental and cross-striated.
lules are clawed in Dolops, but they are modified as Mouth lacks jaws; often on end of snoutlike projection;
stalked suckers in the other genera (Argulus, Chono- connected to a muscular pumping pharynx used to
peltis, Dipteropeltis). The uniramous maxillae usually suck blood from host. The combination of the snout
bear attachment hooks. The thoracopods are bira- and the 2 pairs of legs gives the appearance of there
mous and used for swimming when the animal is not being 5 mouths, hence the name (Greek penta, “five”;
attached to a host. Branchiurans feed by piercing the stomida, “mouths”). In many species the appendages are
skin of their hosts and sucking blood or tissue fluids. reduced to no more than the terminal claws. No specific
Once they locate a host, they crawl toward the fish’s gas exchange, circulatory, or excretory organs. Gonocho-
head and anchor in a spot where water flow turbulence ristic; females larger than males. About 130 described
is low (e.g., behind a fin or gill operculum). Members species, including 2 cosmopolitan species that can occa-
of the genus Argulus occur worldwide, and can pose sionally infest humans (Figure 21.4K, 21.5).
a serious problem to aquaculture, but members of the For years it was believed that pentastomids were
other genera have restricted distributions. Chonopeltis allied with the fossil lobopodians, onychophorans, and
is found only in Africa, Dipteropeltis in South America, tardigrades as some kind of segmented, vermiform,
and Dolops in South America, Africa, and Tasmania. proto-arthropod creature. However, molecular phylo-
genetics informs us that pentastomids are highly modi-
fied crustaceans, perhaps derived from the Branchiura.
Class Pentastomida Corroboration has come from studies of sperm and lar-
Obligatory parasites of various amphibians, reptiles, val morphology, nervous system anatomy, and cuticu-
birds, and mammals. Adults inhabit respiratory tract lar fine structure.

(A) (B) Mouth (C) (D)

Legs

(E)

FIGURE 21.5 Anatomy and diversity

in the class Pentastomida. (A) Linguatula
serrata. (B) Cephalobaena tetrapoda.
(C) Internal anatomy of female
Pentastomum. (D) Internal anatomy
of female Waddycephalus teretiusculus,
a parasite of Australian snakes. (E) A gen-
eralized pentastomid primary larva.

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Work on the Swedish Orsten fauna indicates that male pores on eighth thoracomeres. When uropods are
pentastomid-like animals had appeared as early as the present, they are often broad and flat, lying alongside
late Cambrian (500 Ma), long before the land verte- the broad telson to form a tail fan.
brates had evolved. What might the original hosts of Most classification schemes divide the more than
these parasites have been? Conodont fossils are com- 40,200 species of malacostracans into three subclasses,
mon in all the Cambrian localities that have yielded Phyllocarida (leptostracans), Hoplocarida (stomato-
pentastomids, raising the possibility that conodonts pods), and the megadiverse Eumalacostraca. The phyl-
(also long a mystery, but now widely regarded as parts locarids are typically viewed as representing the primi-
of early fishlike vertebrates) may have been at least one tive malacostracan condition (6-8-7 body segments
of the original hosts of these early Pentastomida. plus telson; Figure 21.6). The basic eumalacostracan
body plan, characterized by the 6-8-6 (plus telson)
Superclass Altocrustacea: arrangement of body segments, was recognized in the
Clade Multicrustacea early 1900s by W. T. Calman, who termed the defining
features of the Eumalacostraca “caridoid facies” (Fig-
Class Malacostraca ure 21.7). Much work has been done since Calman’s
Body of 19–20 segments, including 6-segmented day, but the basic elements of his caridoid facies are still
cephalon/head, 8-segmented thorax (anterior-most present in all members of the subclass Eumalacostraca.
somites often fused with head), and 6-segmented
pleon (7-segmented in leptostracans), plus telson; with
or without caudal rami; carapace covering part or all Subclass Phyllocarida
of thorax, or reduced, or absent; 0–3 pairs of maxil- Order Leptostraca With typical malacostracan charac-
lipeds; thoracopods primitively biramous, uniramous teristics, except notable for presence of 7 free pleomeres
in some groups, phyllopodous only in members of the (plus telson), generally taken to represent the primitive
subclass Phyllocarida; antennules and antennae usu- condition for the class. Also, with phyllopodous thora-
ally biramous; abdomen (pleon) usually with 5 pairs copods (all similar to one another); no maxillipeds; large
of biramous pleopods and 1 pair of biramous uropods;
eyes usually present, compound, stalked or sessile.
(B)
Mainly gonochoristic; female gonopores on sixth, and

(A)

(C)

FIGURE 21.6 Anatomy of leptostracans

(class Malacostraca, subclass Phyllocarida).
(A) General anatomy of Nebalia. (B) Phyllopodous
C,D courtesy of T. Haney

(D)
swimming limb of Nebalia. (C) SEM of Nebalia.
(D) Anterior end of an ovigerous Nebalia.

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FIGURE 21.7 The basic eumala­

costracan body plan and the
“caridoid facies.” Note the thick
(muscled) abdomen and the tail fan,
which work in combination to produce
a powerful tail flip escape reaction.

carapace covering thorax and compressed laterally so as but the pleopods are quite different in the two groups.
to form an unhinged bivalved “shell,” with an adductor The tubular, thin, highly branched gills of stomatopods
muscle; cephalon with a movable, articulated rostrum; provide a large surface area for gas exchange in these
pleopods 1–4 similar and biramous, 5–6 uniramous; no active animals.
uropods; paired stalked compound eyes; antennules All 500 or so living hoplocarids are placed in the
biramous; antennae uniramous; adults with both anten- order Stomatopoda, known as mantis shrimps. They
nal and maxillary glands (Figures 21.6 and 21.21C). are relatively large crustaceans, ranging in length from
The subclass Phyllocarida includes about 40 species 2 to 30 cm. Compared with that of most malacostra-
in 10 genera. Most are 5–15 mm long, but Nebaliopsis cans, the muscle-filled abdomen is notably robust.
typica is a giant at nearly 5 cm in length. The leptostracan Most stomatopods are found in shallow tropical or
body form is distinctive, with its loose bivalved cara- subtropical marine environments. Nearly all of them
pace covering the thorax, a protruding rostrum, and an live in burrows excavated in soft sediments or in cracks
elongate abdomen. All leptostracans are marine, and and crevices, among rubble, or in other protected
most are epibenthic from the intertidal zone to a depth places. All species are raptorial carnivores, preying on
of 400 m; Nebaliopsis typica is bathypelagic. Most species fishes, molluscs, cnidarians, and other crustaceans. The
seem to occur in low-oxygen environments. One species, large, distinctive subchelae of the second thoracopods
Dahlella caldariensis, is associated with the hydrothermal act either as crushers or as spears (Figure 21.8C).
vents of the Galapagos and the East Pacific Rise. Speone- Stomatopods crawl about using the posterior thora-
balia cannoni is known only from marine caves. copods and the flaplike pleopods. They also can swim
Most leptostracans suspension feed by stirring up by metachronal beating of the pleopods (the “swim-
bottom sediments. They are also capable of grasping merets”). For these relatively large animals, living in
relatively large bits of food directly with the mandi- narrow burrows requires a high degree of maneuver-
bles. Some are carnivorous scavengers, and some are ability. The short carapace and the flexible, muscular
known to aggregate in areas on the seafloor where abdomen allow these animals to twist double and turn
large amounts of detritus accumulate. In many species around within their tunnels or in other cramped quar-
the antennae or antennules of males are modified to ters. This ability facilitates an escape reaction whereby
hold females during copulation. a mantis shrimp darts into its burrow rapidly head
first, then turns around to face the entrance.
Subclass Hoplocarida
Order Stomatopoda Carapace covering portion of Subclass Eumalacostraca
head and fused with thoracomeres 1–4; head with mov- Head, thorax, and abdomen of 6-8-6 somites respec-
able, articulated rostrum; thoracopods 1–5 uniramous tively (plus telson); with 0, 1, 2, or 3 thoracomeres fused
and subchelate, second pair massive and raptorial (all with head, their respective appendages usually modi-
5 are sometimes called “maxillipeds” or gnathopods fied as maxillipeds; antennules and antennae primi-
because they are involved in feeding); thoracopods tively biramous (but often reduced to uniramous); most
6–8 biramous, ambulatory; pleopods biramous, with with well-developed carapace, secondarily reduced in
dendrobranchiate-like gills on exopods; antennules tri- syncarids and some peracarids; gills primitively as tho-
ramous; antennae biramous, with large, paired, stalked racic epipods; tail fan composed of telson plus paired
compound eyes that are unique in the animal kingdom uropods; abdomen long and muscular. Three super-
(Figures 21.8A–C, 21.27D, and 21.33K). Stomatopods orders: Syncarida, Peracarida, Eucarida, although the
are one of only two groups of malacostracans that pos- monophyly of the last has been contested in morpho-
sess pleopodal gills. Only the isopods share this trait, logical and molecular analyses.

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Shrimps, and Their Kin 675

(A)

(B)

Courtesy of A. Kerstich
(C) Legs 3–5
(gnathopods)
(D)
Pleon Thoracomeres 1–8
Cephalon
Pleotelson
Furcal lobe Antennule

Rudimentary pleopods

Antenna

(E)
Both courtesy of R. Caldwell

(F) FIGURE 21.8 External anatomy in the class

Malacostraca, subclasses Hoplocarida (stomato­
pods) and Eumalacostraca (syncarids). (A) The
spiny Hawaiian stomatopod Echinosquilla guerinii.
(B) External anatomy of the stomatopod Squilla.
(C) “Spearing” claw and “clubbing” claw (second
thoracopod) of stomatopods. (D) A bathynellacean,
Courtesy of Shane Ahyong

Bathynella. (E) An anaspidacean, Stygocarella.

(F) An anaspidacean, Paranaspides lacustris.

Superorder Syncarida Without maxillipeds (Bathy­ There are about 285 described species of syncarids in
nellacea) or with 1 pair of maxillipeds (Anaspidacea, 2 orders, Anaspidacea and Bathynellacea. Syncarids
including Stygocarididae); no carapace; pleon bears might be an ancient relictual taxon now restricted to
telson with or without furcal lobes; at least some thora- refugial habitats. Through studies of the fossil record
copods biramous; pleopods variable; compound eyes and extant members of the order Anaspidacea (e.g.,
present (stalked or sessile) or absent (Figure 21.8D–F). Anaspides), it has been suggested that syncarids may
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encompass the most primitive living eumalacostracan exceed 1,000 animals/m3 (614 g wet weight/m3).5 The
body plan. Bathynellaceans occur worldwide in inter- high concentrations of omega-3 fatty acids in krill make
stitial or groundwater habitats, whereas the anaspi- them popular for use in dietary supplements. Gener-
daceans are strictly Gondwanan in distribution. Many ally, euphausiaceans are suspension feeders, although
Anaspidacea are endemic to Tasmania, where they predation and detritivory also occur (Figures 21.9A,B
inhabit freshwater environments, such as open lake and 21.21E).
surfaces, streams, ponds, and crayfish burrows. No
syncarids are marine. These reclusive eumalacostra- Order Decapoda With over 15,000 named species,
cans show various degrees of characteristics of what the decapods are among the most familiar eumalacos-
some have regarded as paedomorphism, including tracans. They possess a well-developed carapace enclos-
small size (Anaspidacea includes members to 5 cm, ing a branchial chamber, but they differ from other
whereas most others are less than 1 cm long), eyeless- eucarid orders in always possessing 3 pairs of maxilli-
ness, and reduction or loss of pleopods and some pos- peds, leaving 5 pairs of functional uniramous or weakly
terior pereopods. Bathynellaceans are small (1–3 mm biramous pereopods (hence the name Decapoda); 1 or
long), possess 6 or 7 pairs of long, thin swimming legs, more pairs of anterior pereopods are usually clawed
and have a pleotelson formed by the fusion of the tel- (chelate). Adults have antennal glands. In vernacular
son to the last pleonite. terms, nearly every decapod may be recognized as
Syncarids either crawl or swim. Little is known some sort of shrimp, crab, lobster, or crayfish.
about the biology of most species, although some We do not want to belabor the issue of decapod gill
are considered omnivorous. Unlike most other crus- nomenclature. However, the gills play a prominent
taceans, which carry the eggs and developing early role in the taxonomy of this group; thus, we provide
embryos, syncarids lay their eggs or shed them into brief descriptions of the basic types. All decapod gills
the water following copulation. arise as thoracic coxal exites (epipods), but their final
placement varies. Those that remain attached to the
Superorder Eucarida Telson without caudal rami; coxae are podobranchs (= “foot gills”), but others
0, 1, or 3 pairs of maxillipeds; carapace present, cov- eventually become associated with the articular mem-
ering and fused dorsally with head and entire thorax; brane between the coxae and body and are thus called
usually with stalked compound eyes; gills thoracic. arthrobranchs (= “joint gills”). Some actually end up
Although members of this group are highly diverse, on the lateral body wall, or side-surface of the thorax,
they are united by the presence of a complete cara- as pleurobranchs (= “side gills”). The sequence by
pace that is fused with all thoracic segments, forming which some of these gills arise ontogenetically varies.
a characteristic cephalothorax. Most species (several For example, in the Dendrobranchiata and the Steno-
thousand) belong to the order Decapoda. The other podidea, arthrobranchs appear before pleurobranchs,
order is Euphausiacea (krill). The formerly monotypic whereas in members of the Caridea the reverse is true.
order Amphionidacea (Amphionides reynaudii) has been In most of the other decapods the arthrobranchs and
shown to be a larval caridean decapod. pleurobranchs tend to appear simultaneously. These
developmental differences may be minor heterochronic
Order Euphausiacea Euphausiaceans (krill) are dissimilarities and of less phylogenetic importance
distinguished among the eucarids by their shrimplike than actual gill anatomy.
appearance, absence of maxillipeds, exposure of the Among the decapods, the gills can also be one of
thoracic gills external to the carapace, and possession three basic structural types, described as dendrobran-
of biramous pereopods (the last 1 or 2 pairs sometimes chiate, trichobranchiate, and phyllobranchiate (Figure
being reduced). Adults have antennal glands. Most of 21.28B–D). All three of these gill types include a main
them have photophores on the eyestalks, the bases of axis carrying afferent and efferent blood vessels, but
the second and seventh thoracopods, and between the they differ markedly in the nature of the side filaments
first 4 pairs of abdominal limbs.
5
The 86 known species of euphausiaceans are all Where krill densities exceed about 100 g/m3, they are often fished
commercially. Krill schools can extend for tens of miles, contain
pelagic, range in length from 4 to 15 cm, and are known millions of tons of krill, and stain the ocean red with their surface
from all oceanic environments to depths of 5,000 m. swarms in coastal waters. Large baleen whales can eat a ton of krill
Most species are distinctly gregarious, and species that in one mouthful. Seals, fish, squid, and humans also eat krill. Krill
fishing has been banned in most of North America, but it continues
occur in huge schools provide a major source of food
in Japan, where tens of thousands of tons are landed annually and
for larger nektonic animals (baleen whales, squids, used mainly as feed for farmed fish. The largest krill fishery is in
fishes) and even some marine birds. Northern krill has the ocean surrounding Antarctica, where they have been harvested
been a significant fishery in Japan and Canada, but the commercially since the 1970s. In the 1980s, large fleets from the
Soviet Union caught up to 400,000 tons of Antarctic krill annually,
largest commercial stock is from the Southern Ocean, but by 2015 the annual catch was down to 120,000 tons (taken by
where the annual catch can exceed a half-million tons. Argentina, Chile, Japan, Korea, Norway, Poland, the Ukraine, and
Krill densities, particularly for Euphausia superba, often the United States).

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Shrimps, and Their Kin 677

(A) (B)

FIGURE 21.9 Anatomy of Malacostraca, superorder Eucarida—

euphausiaceans. (A) General body form of a euphausiacean,
Meganyctiphanes. (B) Pereopod of Euphausia superba.

or branches. Dendrobranchiate gills bear two princi-

pal branches off the main axis, each of which is divided
into multiple secondary branches. Trichobranchiate
gills bear a series of radiating unbranched tubular
filaments. Phyllobranchiate gills are characterized by
a double series of platelike or leaflike branches from
the axis. Within each gill type, there may be consider-
able variation. The occurrences of these three major gill
types among various taxa are presented below. Suborder Dendrobranchiata This group includes
Close inspection of the proximal parts of the over 500 species of decapods, most of which are
pereopods usually reveals another decapod feature: penaeid (Penaeoidea) and sergestid (Sergestoidea)
in most forms, the basis and ischium are fused (as a shrimps. As the name indicates, these decapods possess
Brusca 4e
basi-ischium), with the point of fusion often indi- dendrobranchiate gills (see Figure 21.28B), a unique
BB4e_21.[Link]
cated by a suture line (see Figure 21.21E-I). Tegumen- synapomorphy of the taxon. One genus, Lucifer, has
tal5/12/2021
glands are also a ubiquitous feature among the secondarily lost the gills completely. The dendrobran-
Decapoda. These glands originate below the epidermal chiate shrimps are further characterized by chelae on
cells and produce a fluid that opens on the surface of the first 3 pereopods, copulatory organs modified from
the cuticle. They have been reported from gills, legs, the first pair of pleopods in males, and ventral expan-
pleopods, and uropods. The roles of tegmental glands sions of the abdominal tergites (called pleural lobes).
are not well known, and they have been suspected to Generally, none of the chelipeds is greatly enlarged. In
be involved in cuticular tanning, the production of addition, females of this group do not brood their eggs.
mucus by the mouthparts, the production of cement Fertilization is external, and the embryos hatch as nau-
substance involved with egg attachment, and possibly plius larvae. Many of these animals are quite large, over
also grooming. 30 cm long. The sergestids are pelagic and all marine,
Decapods occur in all aquatic environments at all whereas the penaeids are pelagic or benthic, and some
depths, and a few spend most of their lives on land. occur in brackish water. Some dendrobranchiates (e.g.,
Many are pelagic, but others have adopted benthic sed- Penaeus, Sergestes, Acetes, Sicyonia) are of major com-
entary, errant, or burrowing lifestyles. Decorating of the mercial importance in the world’s shrimp fisheries,
exoskeleton is frequently seen among the decapods, most of which are now being exploited beyond sustain-
especially in spider crabs (Brachyura: Majoidea), which able levels and often with fishing techniques that are
use Velcro-like hooked setae to attach dead or living highly habitat destructive (Figures 21.10A and 21.33G).
seaweeds and animals; decorating has been shown to
reduce predation through camouflage and/or chemi- Suborder Pleocyemata All of the remaining deca-
cal deterrence. Decapod feeding strategies include sus- pods belong to the suborder Pleocyemata, divided
pension feeding, predation, herbivory, scavenging, and among 10 infraorders (see below). Members of this
more. Two suborders are recognized: Dendrobranchiata taxon never possess dendrobranchiate gills. The
and Pleocyemata. Recent phylogenetic research sup- embryos are brooded on the female’s pleopods and
ports the monophyly of the suborders and infraorders. hatch at some stage later than the nauplius larva.

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 678 Chapter 21

(A) (B)

(C) (E)

(D)

FIGURE 21.10 External anatomy and diversity

in decapod shrimps. (A) A penaeid shrimp (Dendro­
branchiata), Penaeus setiferus. (B) A procarididean
shrimp (Procarididea), Procaris ascensionis. (C) A hippo-
lytid shrimp (Hippolytidae), Lysmata californica. (D) An
alpheid, or snapping shrimp (Alpheidae), Alpheus.
(E) A stenopodid shrimp (Stenopodidea), Stenopus.

Included in this suborder are several kinds of shrimps, Processa, Syncaris, Thor, Typton, Typhlocaris, Vetericaris).
Brusca 4e crayfish, lobsters, and a host of less familiar
the crabs, Swimming decapods with phyllobranchiate gills. The
BB4e_21.[Link]
forms. De Grave et al. (2009) divide the Pleocyemata first 1 or 2 pairs of pereopods may be chelate and variably
5/12/2021
into 10 infraorders (below). One older approach divid- enlarged. The second abdominal pleurae (side walls) are
ed decapods into 2 large groups, called the Natantia distinctly enlarged to overlap both the first and third
and Reptantia—the swimming and walking decapods, pleurae (Figures 21.1K, 21.10C,D, 21.24E, and 21.31D).
respectively. Although these terms have largely been Procaris and Vetericaris are known from anchialine habi-
abandoned as formal taxa, recent molecular phyloge- tats—inland pools with brackish water on top and seawa-
netic work finds support for a monophyletic Reptantia ter below, with connections to the ocean (Figure 21.10B).
(pleocyemates with dorsoventrally flattened pleons).
In any case, the terms still serve a useful descriptive Infraorder Stenopodidea Stenopodidean shrimps;
purpose, and one continues to see references to natant about 70 species (e.g., Odontozona, Spongicola,
decapods and reptant decapods. Stenopus). The first 3 pairs of pereopods are chelate,
and the third pair is significantly larger than the oth-
Infraorder Caridea Caridean and procarididean ers. The gills are trichobranchiate. The second abdomi-
shrimps; about 3,270 species (e.g., Alpheus, Ambidexter, nal pleurae are not expanded as they are in carideans
Atya, Betaeus, Crangon, Hippolyte, Hymenocera, (Figures 21.10E and 21.31B).
Hymenodora, Leander, Lysmata, Macrobrachium, Ogyrides, These colorful shrimps are usually only a few cen-
Palaemon, Pandalus, Pasiphaea, Periclimenes, Procaris, timeters long (2–7 cm). Most species are tropical and

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Shrimps, and Their Kin 679

associated with shallow benthic environments, espe- but exceptions occur. First pereopods chelate, usually
cially with coral reefs; some are known from the deep enlarged and larger in males (usually used in contest-
sea. Many are commensal, and the group includes the ing mating rights with other males). Pereopods 2 to 5
cleaner shrimps (e.g., Stenopus) of tropical reefs, which typically simple stenopodous walking legs. Eyes posi-
are known to remove parasites from fishes. Stenopo- tioned lateral to the antennae. Males lack pleopods 3
dids often occur as male-female couples. Perhaps the to 5. Always gonochoristic. The distinctive larval stage
most noted example of this bonding is associated with is called a zoea; its carapace is spherical and bears a
the glass sponge (Euplectella) shrimp, Spongicola venusta: ventrally directed rostral spine (or no spine). Common
a young male and female shrimp enter the atrium of brachyuran genera include Calappa, Callinectes, Cancer,
a host sponge, eventually growing too large to escape Cardisoma, Dromia, Ebalia, Epialtus, Eriphia, Fabia,
and thus spending the rest of their days together. Gecarcinus, Geryon, Goneplax, Grapsus, Hepatus, Herbstia,
Libinia, Loxorhynchus, Maja, Menippe, Microphrys,
Infraorder Brachyura The so-called “true crabs” Ocypode, Ozius, Panopeus, Percnon, Pinnixa, Portunus,
(about 7,000 species). Abdomen symmetrical but highly Pugettia, Thoe, Trapezia, Uca, and Xanthias (Figures 21.1E,
reduced and flexed beneath the thorax. Body hidden 21.11, 21.27H, 21.28F,G, 21.29C, 21.32, and 21.33H,I).
beneath well-developed carapace and distinctly flat- Brachyuran crabs are mostly marine, but freshwa-
tened dorsoventrally. Gills typically phyllobranchiate, ter, semiterrestrial, and terrestrial species occur in the

(A) (C)

(D)
(B)

(E)

FIGURE 21.11 Anatomy and diversity of the “true,” or

brachyuran, crabs (Decapoda: Brachyura). (A,B) General crab
anatomy: frontal and ventral views of a swimming crab (family
Portunidae). (C) A spider crab (family Majidae), Loxorhynchus. (D) An
arrow crab (family Majidae), Stenorhynchus. (E) A cancer crab (family
Cancridae), Cancer.
(Continued on next page)

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 680 Chapter 21

(F)

(G)

(H) (I)

FIGURE 21.11 (continued) Anatomy and diversity of the

“true,” or brachyuran, crabs (Decapoda: Brachyura).
(F) A grapsid crab (family Grapsidae), Pachygrapsus. (G) A pin-
notherid or pea crab (family Pinnotheridae), Parapinnixa.

Courtesy of E. Spivak
(H) A dromiid crab (family Dromiidae), Hypoconcha (anterior
view). Members of the Dromiidae carry bivalve mollusc shells
(or other objects) on their backs. (I) Ventral views of a female
(upper photo) and male (lower photo) Cyrtograpsus angulatus.

tropics. The land crabs (certain species in the families porcelain crabs, mole crabs, and sand crabs. The abdo-
Gecarcinidae, Ocypodidae, Grapsidae, etc.) are still men may be soft and asymmetrically twisted (as in her-
dependent on the ocean for breeding and larval devel- mit crabs) or symmetrical, short, and flexed beneath
opment. The surprisingly large number of freshwater the thorax (as in porcelain crabs and others). Those
crabs (about 3,000 species, classified into about a dozen with twisted abdomens typically inhabit gastropod
families) all have direct development, incubate their shells or other empty “houses” not of their own mak-
embryos, and are independent of seawater. Some fresh- ing. Carapace shape and gill structure vary among the
water crabs are intermediate hosts of Paragonimus, a cos- Anomura. The first pereopods are chelate; the third
motropical parasitic human lung fluke, and others are pereopods are never chelate. The second, fourth, and
obligate phoretic hosts of larval black flies (Simulium), fifth pairs are usually simple, but occasionally they are
the vector for Onchocerca volvulus (the causative agent chelate or subchelate. The fifth pereopods (and some-
of river blindness). A number of crab species carry other times the fourth) are generally much reduced and do not
invertebrates on their carapace (e.g., sponges, tunicates) function as walking limbs; the fifth pereopods function
or on their claws (e.g., anemones); these associations as gill cleaners and often are not visible externally. The
are generally thought to be mutualistic, providing cam- pleopods are reduced or absent. The eyes are positioned
ouflage or predator deterrence for the crab while their medial to the antennae. The zoea larva is similar to that
partner is moved about in the environment and may of the true crabs but is typically longer than broad, with
feed off debris from their host’s feeding activities. In the the rostral spine directed anteriorly. Most anomurans
northeast Pacific, megalopae and juveniles of the crab are marine, but a few freshwater and semiterrestrial
Metacarcinus gracilis ride (and feed) on the bell of certain species are known. The so-called “yeti crab” (Kiwa hir-
jellyfish; individuals of Phacellophora camtschatica (Scy- suta) was discovered in 2005 at a depth of 2,200 m in
phozoa) have been found with hundreds of M. gracilis hydrothermal vents south of Easter Island. It is remark-
megalopae. Evidence suggests Brachyura and Anomura able for its “garden” of filamentous bacteria that grow
are sister groups (a clade known as Meiura). on the long setae of the exoskeleton; the bacteria are het-
erotrophic, utilizing sulfides in the deep environment
Infraorder Anomura This group includes about 2,500 (the precise role of the bacteria, of several species, in the
species of hermit crabs, galatheid crabs, king crabs, life history of the yeti crab is not yet well understood).
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Shrimps, and Their Kin 681

Several additional Kiwa species have been described terminates in a strong tail fan (Figure 21.2). The gills
since 2011. Common anomuran genera include Aegla, are trichobranchiate. The first 3 pairs of pereopods are
Birgus, Blepharipoda, Calcinus, Clibanarius, Coenobita, always chelate, and the first pair is greatly enlarged.
Dardanus, Diogenes, Emerita, Hapalogaster, Hippa, Kiwa, Homarus americanus, the American or Maine lobster, is
Lepidopa, Lithodes, Lomis, Pagurus, Petrocheles, Petrochirus, strictly marine and is the largest living crustacean by
and Pleuroncodes (Figures 21.1F–J, 21.12C–G, 21.24A–C, weight (the record weight being over 20 kg). Most cray-
21.31A, and 21.33I). fish live in fresh water, but a few species live in damp
soil, where they may excavate extensive and complex
Infraorder Astacidea Freshwater crayfish and clawed burrow systems. The 670+ species of freshwater cray-
lobsters; 685 species (e.g., Astacus, Cambarus, Cherax, fish (in 5 families) comprise a monophyletic group that
Faxonella, Homarus, Nephrops, Pacifastacus). As in most is sister group to clawed lobsters, the crayfish originat-
other decapods, the dorsoventrally flattened abdomen ing around 330 million years ago. Over 425 species of

(A) (C)

Walking legs Chela

(D)
(B)

(E) (F) (G)

FIGURE 21.12 External anatomy and diversity

in some other reptant decapods (Eumalacostraca,
Eucarida). (A) A mud shrimp, Callianassa (Gebiidea).
(B) A spiny lobster, Panulirus (Achelata). (C) A hermit
crab, Paguristes, in its shell (Anomura). (D) A hermit crab,
Pagurus, removed from its shell to expose the soft abdo-
men. (E) A porcelain crab, Petrolisthes (Anomura), with the
reduced posterior pereopods extended. (F) A sand crab,
Emerita (Anomura). (G) The umbrella crab Cryptolithodes
(Anomura), in ventral view.

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 682 Chapter 21

crayfish occur in North America alone, where they show articulated accessory process in adults, between molar
high levels of endemicity to particular regions or river and incisor process, called the lacinia mobilis; carapace,
drainages (Figures 21.27E,G and 21.29B). Many of these when present, not fused with posterior pereonites and
are among the most invasive freshwater crustaceans. usually reduced in size; gills thoracic or abdominal; with
unique, thinly flattened thoracic coxal endites, called
Infraorder Achelata This group includes 140 species oostegites, that form a ventral brood pouch or marsupi-
of spiny and slipper lobsters, many of economic impor- um in females of all species except members of the order
tance (e.g., Arctides, Evibacus, Ibacus, Panulirus). The Thermosbaenacea (the latter using the carapace to brood
name Achelata comes from the fact that they lack chelae embryos); young hatch as mancas, a prejuvenile stage
on all pereopods as adults (except for a small groom- lacking the last pair of thoracopods (no free-living larvae
ing claw on pereopod 5 in some females). The flattened occur among the Peracarida) (Figures 21.13–21.16).
abdomen bears a tail fan; the carapace may be cylindri- The roughly 25,000 species of peracarids are divided
cal or flattened dorsoventrally; the gills are trichobran- among ten orders. The peracarids are an extremely suc-
chiate. The large, flattened larvae, called phyllosomas cessful group of malacostracan crustaceans known from
because of their leaflike appearance, are unique and dis- many habitats. Although most are marine, many also
tinctive. All species are marine, and they are found in occur on land and in fresh water, and several species live
a variety of habitats throughout the tropics. Many spe- in hot springs at temperatures of 30°C–50°C. Aquatic
cies produce sounds by rubbing a process (the plectrum) forms include planktonic as well as benthic species at all
at the base of the antennae against a “file” on the head depths. The group includes the most successful terres-
(Figures 21.1M, 21.12B, 21.30A,C, and 21.33L). trial crustaceans—the pillbugs and sowbugs of the order
Isopoda—and a few amphipods that have invaded land
Infraorders Axiidea and Gebiidea The lobster and live in damp forest leaf litter or gardens. Peracarids
shrimps (Axiidea, 423 species, e.g., Axius, Callianassa, range in size from tiny interstitial forms only a few mil-
Neotrypaea, Upogebia) and mud/ghost shrimps limeters long to planktonic amphipods over 12 cm long
(Gebiidea, 192 species, e.g., Axianassa, Naushonia). These (Cystisoma), deep-sea necrophagous amphipods exceed-
2 infraorders, formerly combined as Thalassinidea, ing 34 cm (Alicella gigantea), and benthic isopods grow-
have recently been recognized as distinct (the ver- ing to 50 cm in length (Bathynomus giganteus; Cirolani-
nacular term “thalassinid” is still sometimes used to dae). These animals exhibit all sorts of feeding strategies;
refer to them together) though they are very similar in a number of them, especially isopods and amphipods,
appearance. The phylogenetic relationships of these are commensals or parasites.
shrimps remain unsettled. The symmetrical abdomen
is flattened dorsoventrally and extends posteriorly as Order Mysida Carapace well developed, cover-
a well-developed tail fan. The carapace is somewhat ing most of thorax, but never fused with more than 4
compressed laterally, and the gills are trichobranchiate. anterior thoracic segments; maxillipeds (1–2 pairs) not
The first 2 pairs of pereopods are chelate, and the first associated with cephalic appendages; thoracomere 1
pair is generally much enlarged. Most of these animals separated from head by internal skeletal bar; abdomen
are marine burrowers or live in coral rubble. They gen- with well-developed tail fan; pereopods biramous,
erally have a rather thin, lightly sclerotized cuticle, but except last pair, which are sometimes reduced; pleo-
some (e.g., members of the family Axiidae) have thick- pods reduced or, in males, modified; compound eyes
er skeletons and are more lobsterlike in appearance. stalked, sometimes reduced; gills absent; usually with
Gebiideans often occur in huge colonies on tidal flats, a statocyst in each uropodal endopod; adults with
where their burrow holes form characteristic patterns antennal glands (Figures 21.13A,B, 21.30B, and 21.33C).
on the sediment surface (Figures 21.1L and 21.12A). There are more than 1,050 species of mysidans, rang-
ing in length from about 2 mm to 8 cm, in two families:
Infraorders Glypheidea and Polychelida The gly- Mysidae and Petalophthalmidae. Most swim by action of
pheids are something of a relict group, represented the thoracic exopods. These are shrimplike crustaceans
by 2 living genera (Neoglyphea and Laurentaeglyphea), that are often confused with the superficially similar
each with a single species, of a formerly diverse group euphausiaceans (which lack oostegites and uropodal
known from the fossil record. Polychelids are a group statocysts). Mysidans are pelagic or demersal and are
of about 38 blind deep-sea lobsters (e.g., Polycheles), known from all ocean depths; a few species occur in fresh
notable for having chelae on all of their pereopods and water. Some species are intertidal and burrow into the
unusual, large, globate larvae (called eryoneicus lar- sand during low tides. Most are omnivorous suspension
vae) unique among the decapods. feeders, eating algae, zooplankton, and suspended detri-
tus. In the past, mysidans were combined with lophogas-
Superorder Peracarida Telson without caudal rami; tridans, stygiomysidans, and the extinct Pygocephalo-
1 pair (rarely 2–3) of maxillipeds; maxilliped basis typi- morpha as the “Mysidacea.” Phylogenetic studies show
cally produced as a bladelike endite; mandibles with these groups to be very closely related to one another.

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 for more ebook/ testbank/ solution manualsArthropoda
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Shrimps, and Their Kin 683

(A) (B)
Courtesy of E. Peebles

(C)

(D)

(E)

(F)

Courtesy of J. Corbera
FIGURE 21.13 Anatomy and diversity in some peracarid
(G) crustaceans (Eumalacostraca, Peracarida)—mysidans,
lophogastrids, cumaceans, and tanaidaceans. (A) A mysid,
Chlamydopleon dissimile. (B) Anatomy of a generalized mysid.
Courtesy of E. Peebles

(C) Anatomy of a lophogastrid, Gnathophausia. (D) Second

pereopod of Gnathophausia. (E) A cumacean, Diastylis, in its
typical partially buried position. The arrows indicate the feed-
ing and ventilation current. (F) A cumacean. (G) A tanaidacean.
(H) Anatomy of a generalized tanaidacean.

(H)

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 684 Chapter 21

(A)

Courtesy of G. Giribet
(B) (C)

Courtesy of J. Olesen
(D)

NOAA Ocean Explorer/Peter Parks/Wikipedia

(E)

FIGURE 21.14 More peracarids. (A) The spelaeogriphacean,

Spelaeogriphus lepidops (color photo), and a drawing of generalized
spelaeogriphacean anatomy. (B) The thermosbaenacean, Monodella.
(C) A thermosbaenacean. (D,E) The mictacean, Mictocaris. (E after
T. E. Bowman and T. M. Iliffe. 1985. J Crust Biol 5: 58–73.)
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Shrimps, and Their Kin 685

(A) (B)

(C)
(D)

(E)
(F) (G)

(H)

FIGURE 21.15 More peracarids: the order Isopoda. (E) An asellote, Joeropsis (family Joeropsididae). (F) An
(A) Excorallana (Cymothooidea: Corallanidae). (B) A flabel- anthurid, Mesanthura (family Anthuridae). (G) A gnathiide-
liferan, Codonophilus (family Cymothoidae). Members of an, Gnathia (family Gnathiidae). Note the grossly enlarged
this genus are parasites that attach to the tongues of vari- mandibles characteristic of male Gnathiidae. (H) An onisci-
ous marine fishes. (C) A flabelliferan, Heteroserolis (family dean, Ligia (the common seashore “rock louse”).
Serolidae). (D) A valviferan, Idotea (family Idoteidae).

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 686 Chapter 21
(A) Sessile compound eye
Ped
Key Head unc
le
Urosome

Fl
ag
Pleon

el
lu
n

m
me
Coxal plates

do
1 Antenna

Ab
Pereon Antennule
2
Pleopads Gnathopods
Telson 4 3
5
3 2 1 7 6

Uprods
Pereopods
Antennule

Sessile
compound eye
(B) (C)
en Peduncle Lateral gill
d om
Ab Pereopod 3
7 6 5 4 321 Medial gill
3
2
1
7 6 5 43 2 1
Telson 2-3
1
2
3 2 3 Pereopods
1

Uprods Pleopods
Pereopods
Pene
Antenna Flagellum

Antennule
Abdomen

(D) (E)
Courtesy of E. Peebles

Courtesy of E. Peebles

FIGURE 21.16 And still more peracarids: amphipods and (F)

ingolfiellidans. (A) General anatomy of an amphipod. (B) General
anatomy of a hyperiidean amphipod. (C) General anatomy of
a cyamid amphipod (Cyamus monodontis). (D) A senticaudatan
Courtesy of T. Haney

amphipod, Melita. (E) A caprellid amphipod. (F) A cyamid amphi-

pod, Cyamus erraticus, parasitic on right whales. (G,H) Two
senticaudatan amphipods: (G) Hyale, a beach hopper, and
(H) Heterophlias, an unusual, dorsoventrally flattened species.
(I–K) Three hyperiidean amphipods: (I) Primno, (J) Leptocottis,
and (K) a hyperiid on its host medusa. (L) A free-living caprellid,
Brusca 4e
Caprella. (M) The ingolfiellidan Ingolfiella.
BB4e_21.[Link]
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Shrimps, and Their Kin 687

(G) (H)

(J) (K)
(I)

(L)

(M)

Order Stygiomysida Similar to mysidans. With bira- maxillary glands; without uropodal statocysts; all 7
mous male and female pleopods, reduced to 1-segmented pairs of pereopods well developed and similar (except
endopods and 3-segmented exopods; elongated uropod among members of the family Eucopiidae, in which
protopodites; uropods lacking statocysts; lack of podo- their structure varies) (Figures 21.13C,D and 21.21G).
branchs; thoracopods 2–4 modified as gnathopods; mar- There are three families and about 55 known spe-
supium formed of 4 pairs of oostegites arising from tho- cies of lophogastridans, most of which are 1–8 cm long,
racopods 3–6. Found primarily in subterranean waters although the giant Neognathophausia ingens reaches 35
with a marine influence, including anchialine caves, land cm. All are pelagic swimmers, and the group has a cos-
crab burrows, and shrimp culture fields. Endemic spe- mopolitan oceanic distribution. Lophogastridans are
Brusca
cies are4e
known from anchialine caves in Caribbean Sea primarily predators on zooplankton.
BB4e_21.[Link]
and Italy. Two families, fewer than 20 described species.
5/14/2021 Order Cumacea Carapace present, covering and
Order Lophogastrida Similar to mysidans, except fused to first 3 thoracic segments, whose appendages
for the following: maxillipeds (1 pair) associated with are modified as maxillipeds, the first with modified
the cephalic appendages; thoracomere 1 not separat- branchial apparatus associated with branchial cavity
ed from head by internal skeletal bar; pleopods well formed by carapace; pereopods 1–5 ambulatory and
developed; gills present; adults with both antennal and simple, 1–4 may be biramous; pleopods usually absent

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 688 Chapter 21

in females and present in males; telson sometimes specimens have been recovered and some have been
fused with sixth pleonite, forming pleotelson; uropods studied alive. It is pelagic in cave waters and swims by
styliform; compound eyes absent, or sessile and usu- using its pereopodal exopods. Some workers recognize
ally fused (Figures 21.1P and 21.13E,F). the family Hirsutiidae (containing Hirsutia, Montucaris,
Cumaceans are small, odd-looking crustaceans with and Thetispelecaris) as a separate order, the Bochusacea
a large, bulbous anterior end and a long, slender poste- (male pleopods biramous).
rior—resembling a horizontal comma! The great carci-
nologist Waldo Schmitt referred to them as “little won- Order Spelaeogriphacea Carapace short, fused with
ders and queer blunders.” They occur worldwide and first thoracomere; 1 pair of maxillipeds; pereopods 1–7
include about 1,500 species, most of which are between simple, biramous, with shortened exopods; exopods
0.1 and 2 cm long, though some species in cold waters on legs 1–3 modified for producing currents, on legs
reach 3 cm in length. Most are marine, although a few 4–7 as gills; pleopods 1–4 biramous, natatory; pleopod
brackish-water species are known. They live in associa- 5 reduced; tail fan well developed; compound eyes
tion with bottom sediments but are capable of swim- nonfunctional or absent, but eyestalks persist (Figures
ming and probably leave the bottom to breed. Most are 21.14A and 21.21H). The order Spelaeogriphacea is
deposit feeders or predators on the meiofauna; others currently known from only four living species. These
eat the organic film on sand grains. The oldest known rare, small (less than 1 cm) peracarids were long
fossil cumacean, Eobodotria muisca, is from an exquisite, known only from a single species living in a freshwater
mid-Cretaceous (90–95 Ma), sedimentary deposit in stream in Bat Cave on Table Mountain, South Africa
tropical Colombia, and it has modern familial affinities. (Spelaeogriphus lepidops; Figure 21.14A). A second spe-
cies is known from a freshwater cave in Brazil, and
Order Tanaidacea Carapace present and fused with a third and fourth species were described from an
first 2 thoracic segments, forming a cephalothorax; aquifer in Australia. Little is known about the biology
thoracopods 1–2 are maxillipeds, the second being of these animals, but they are suspected to be detri-
chelate; thoracopods 3–8 are simple, ambulatory pereo- tus feeders. Like thermosbaenaceans, spelaeogriph-
pods; pleopods present or absent; uropods biramous aceans are thought to be relicts of a more widespread
or uniramous; telson and last 1 or 2 pleonites fused as shallow-water marine Tethyan fauna stranded in inter-
pleotelson; adults with maxillary and (vestigial) anten- stitial and ground-water environments during periods
nal glands; compound eyes absent, or present and of marine regression.
on cephalic lobes. Members of this order are known
worldwide from benthic marine habitats; a few live in Order Thermosbaenacea Carapace present, fused
brackish or nearly fresh water. Most of the 1,500 or so with first thoracomere and extending back over 2–3
species are small, ranging from 0.5 to 2 cm in length. additional segments; 1 pair of maxillipeds; pereopods
They often live in burrows or tubes and are known biramous, simple, lacking epipods and oostegites;
from all ocean depths. Many are suspension feeders, carapace forms dorsal brood pouch (unlike all other
others are detritivores, and still others are predators peracarids, which form the brood pouch from ventral
(Figure 21.13G,H). oostegites); 2 pairs of uniramous pleopods; uropods
biramous; telson free or forming pleotelson with last
Order Mictacea Without a carapace, but with a pleonite; eyes absent (Figure 21.14B,C). About 35 spe-
well-developed head shield fused with first thoraco- cies of thermosbaenaceans are recognized in 7 genera.
mere and produced laterally over bases of mouthparts; Thermosbaena mirabilis is known from freshwater hot
1 pair of maxillipeds; pereopods simple, 1–5 or 2–6 springs in North Africa, where it lives at temperatures
biramous, exopods natatory; gills absent; pleopods in excess of 40°C. Several species in other genera occur
reduced, uniramous or biramous; uropods biramous, in much cooler fresh waters, typically in groundwater
with 2–5 segmented rami; telson not fused with ple- or in caves. Other species are marine or inhabit under-
onites; stalked eyes present (Mictocaris) but lacking ground anchialine pools. Limited data suggest that
any evidence of visual elements, or absent (Hirsutia) thermosbaenaceans feed on plant detritus.
(Figure 21.14D–E).
The order was erected to accommodate two species Order Isopoda Carapace absent; first thoracomere
of unusual crustaceans: Mictocaris halope (from marine fused with head; 1 pair of maxillipeds; 7 pairs of uni-
caves in Bermuda) and Hirsutia bathyalis (from a benthic ramous pereopods, the first of which is sometimes
sample 1,000 m deep in the Guyana Basin off northeast- subchelate, others usually simple (gnathiids have
ern South America). A third species of Mictacea was only 5 pairs of pereopods, as thoracopod 2 is a max-
described in 1988 from Australia, and a fourth from illipedal “pylopod” and thoracopod 8 is missing);
the Bahamas in 1992; there are now six species known. pereopods variable, modified as ambulatory, pre-
Mictaceans are small, 2–3.5 mm in length. Mictocaris hensile, or swimming; in the more derived suborders
halope is the best known of these species because many pereopodal coxae are expanded as lateral side plates

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Shrimps, and Their Kin 689

(coxal plates); pleopods biramous and well developed, uropods; telson free or fused with last urosomite; other
natatory and for gas exchange (functioning as gills in urosomites sometimes fused; compound eyes sessile,
aquatic taxa, and with air sacs called pseudotrachea in absent in some, huge in many (but not all) members of
most terrestrial Oniscidea); adults with maxillary and the suborder Hyperiidea (Figures 21.1N,O, 21.16, 21.23,
(vestigial) antennal glands; telson fused with 1 to 6 21.27F, and 21.29D). Our classification follows Lowry
pleonites, forming pleotelson; eyes usually sessile and and Myers (2013, 2017) who recognize 6 suborders.
compound, absent from some, pedunculate in most The roughly 10,000 species of amphipods range in
Gnathiidea; with biphasic molting (posterior region length from tiny 1 mm forms to giant deep-sea ben-
molts before anterior region) (Figures 21.1Q, 21.15, thic species reaching 30 cm, and some planktonic
21.21I, 21.28H,I, and 21.33M). forms exceed 10 cm. They have invaded most marine
The isopods comprise about 10,000 marine, freshwa- and freshwater habitats and often constitute a large
ter, and terrestrial species, ranging in length from 0.5 to portion of the biomass in many areas. Some are com-
500 mm, the largest being species of the benthic genus mon in subterranean groundwater ecosystems, the
Bathynomus (Cirolanidae). They are common inhabit- majority being stygobionts (e.g., Niphargus, Stygobro-
ants of nearly all environments, and some groups are mus)—obligatory groundwater species characterized
exclusively (e.g., Bopyridae, Cymothoidae) or partly by reduction or loss of eyes, pigmentation, and occa-
(e.g., Gnathiidae) parasitic. sionally appendages. There are many intertidal species,
The suborder Oniscidea includes about 3,600 spe- and a great many of these live in association with other
cies that have invaded land (pillbugs and sowbugs); invertebrates and with algae. Domicolous gammarid-
they are the most successful terrestrial crustaceans. ean amphipods in at least three families spin silk from
Their direct development, osmoregulatory capabili- their legs that is used for consolidating the walls of
ties, thickened cuticle, and aerial gas exchange organs their tube or shelter. The largely terrestrial superfamily
(pseudotrachea) allow most oniscideans to live com- Talitroidea alone comprises 117 genera, some of which
pletely divorced from aquatic environments. However, are common in moist gardens and greenhouses (e.g.,
recent molecular phylogenies suggest Oniscidea is not Arcitalitrus sylvaticus). The largest known amphipod is
a monophyletic group, invasion of land from the sea Alicella gigantea, a cosmopolitan marine species living
happened more than once, and the first land isopods at depths of up to 7,000 m.
appeared about 290 million years ago (coincident with The suborder Hyperiidea includes exclusively
the formation of Pangaea and the diversification of pelagic amphipods that have escaped the confines of
vascular plants on land). Males of several species of benthic life by becoming associated with other plank-
Oniscidea are reported to have tegumental glands; the ters, particularly gelatinous zooplankton such as
glands comprise secretory cells and ducts to the cuticle medusae, ctenophores, and salps. The hyperiideans
surface, and they are suspected to produce peptides or are usually characterized by transparent bodies and
glycoproteins involved in mating. huge eyes (and a few other inconsistent features), but
Isopod feeding habits are extremely diverse. Many several groups bear eyes no larger than those of most
are herbivorous or omnivorous scavengers, but direct other amphipods. The Hyperiidea are almost certainly
plant feeders, detritivores, and predators are also com- a polyphyletic group, and it is thought that several lin-
mon. The wood-boring marine gribbles, Limnoria (Onis- eages are derived independently from various ances-
cidea), comprise one of the few animal groups able to tors, although a modern phylogenetic analysis has yet
survive on a diet of cellulose without relying on resi- to be attempted. The precise nature of the relation-
dent gut microbiota. Some isopods are parasites (e.g., ships between hyperiideans and their zooplankton
on fishes or on other crustaceans) that feed on the tissue hosts remains controversial. Some appear to eat host
fluids of their hosts. Overall, grinding mandibles and tissue, others may kill the host to fashion a floating
herbivory seem to represent the primitive state, with “home,” and still others may utilize the host merely
slicing or piercing mandibles and predation appearing for transport or as a nursery for newly hatched young.
later in the evolution of several isopod clades. Specimens of the scyphozoan Phacellophora camtschatica
have been found with nearly 500 Hyperia medusarum
Order Amphipoda Carapace absent; first thoraco- riding (and feeding) on them. Species of Phronima are
mere fused to head; 1 pair of maxillipeds; 7 pairs of uni- famous for their habit of hollowing out the bodies of
ramous pereopods, with first, second, and sometimes salps (pelagic tunicates) to reside inside, propelling
others frequently modified as chelae or subchelae; themselves around like miniature submersible vehi-
pereopodal coxae expanded as lateral side plates (coxal cles. Their extraordinarily large eyes are said to have
plates); gills thoracic (medial pereopodal epipods); inspired the look of the creature in the legendary sci-
adults with antennal glands; abdomen “divided” into ence fiction film Alien.
2 regions of 3 segments each, an anterior “pleon” and The 300 or so species of caprelloidean amphipods,
posterior urosome, with anterior appendages as typi- or “skeleton shrimp” (a clade within the suborder Sen-
cal pleopods and urosomal appendages modified as ticaudata), are highly modified for clinging to other

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 690 Chapter 21

organisms, including filamentous algae and hydroids. Harpacticoida, and Cyclopoida, although even some of
In most species the body and appendages are very nar- these are parasitic. We focus here on these three groups
row and elongated. In one family of Caprelloidea, the and then briefly discuss some of the other, smaller
Cyamidae (with 28 species), individuals are obligate orders and their modifications for parasitism.
symbionts on cetaceans (whales, dolphins, and por- The calanoids are characterized by a point of major
poises) and have flattened bodies and prehensile legs. body flexure between the metasome and the urosome,
In addition to parasitism, amphipods exhibit a vast marked by a distinct narrowing of the body. They
array of feeding strategies, including scavenging, her- possess greatly elongate antennules. Most of the cala-
bivory, carnivory, and suspension feeding. noids are planktonic, and as a group they are extremely
important as primary consumers in freshwater and
Order Ingolfiellida Head with vestigial pedunculate marine food webs. The point of body flexure in the
eyes; gnathopods 1–2 eucarpochelate; pleosome with 6 orders Harpacticoida and Cyclopoida is between the
relatively undifferentiated segments, without epimera, last two (fifth and sixth) metasomal segments. (Note:
with reduced pleopods and uropods. Formerly placed Some authors define the urosome in harpacticoids and
within Amphipoda; 45 species in 2 families distributed cyclopoids as that region of the body posterior to this
worldwide, from deep waters to underground fresh point of flexure.) Harpacticoids are generally rather
waters in all sorts of habitats. vermiform, with the posterior segments not much nar-
rower than the anterior; cyclopoids generally narrow
abruptly at the major body flexure. Both the antennules
Class Copepoda and the antennae are quite short in harpacticoids, but
Without a carapace, but with a well-developed cephalic the latter are moderately long in cyclopoids (although
shield; single, median, simple maxillopodan eye (some- never as long as the antennules of calanoids). Most
times lacking); 1 or more thoracomeres fused to head; harpacticoids are benthic, and those that have adapted
thorax of 6 segments, the first always fused to the head to a planktonic lifestyle show modified body shapes.
and with maxillipeds; abdomen of 5 segments, includ- Harpacticoids occur in all aquatic environments;
ing anal somite (= telson); well-developed caudal rami; encystment is known to occur in at least a few fresh-
abdomen without appendages, except an occasional water and marine species. Cyclopoids are known from
reduced pair on the first segment, associated with the fresh and salt water, and most are planktonic.
gonopores; point of main body flexure varies among The nonparasitic copepods move by crawling or
major groups; antennules uniramous, antennae unira- swimming, using some or all of the thoracic limbs.
mous or biramous; 4–5 pairs of natatory thoracopods, Many of the planktonic forms have very setose append-
most locked together for swimming; posterior thoraco- ages, offering a high resistance to sinking. Calanoids
pods always biramous; many species with myelin-like are predominantly planktonic feeders. Benthic harpac-
sheaths on some nerves (Figures 21.1U, 21.4B–L, ticoids are often reported as detritus feeders, but many
21.27A, 21.30D, and 21.33D). feed predominantly on microorganisms living on the
There are more than 14,500 described species of cope- surface of detritus or sediment particles (e.g., diatoms,
pods. They can be incredibly abundant in the world’s bacteria, and protists).
seas, and also in some lakes, and they compete with Of the seven remaining orders, the Mormonilloida
euphausiaceans and ants for the title of most numer- are planktonic; the Misophrioida are known from
ous of all animals on Earth. Some estimates suggest deep-sea epibenthic habitats as well as anchialine caves
that copepods account for 80% of all mesozooplankton in both the Pacific and Atlantic; and the Monstrilloida
biomass in the ocean. Most are small, 0.5–10 mm long, are planktonic as adults, but the larval stages are endo-
but some free-living forms exceed 1.5 cm in length, and parasites of certain gastropods, polychaetes, and occa-
certain highly modified parasites may reach 25 cm. sionally echinoderms. Members of the orders Ergasi-
The bodies of most copepods are distinctly divided lida and Siphonostomatoida are exclusively parasitic
into three tagmata, the names of which vary among and often have modified bodies; some form galls on
authors. The first region includes the fused head seg- their hosts (e.g., lamippids on sea pens). Siphonosto-
ments and one or two additional fused thoracic somites; matoids are endo- or ectoparasites of various inverte-
it is called a cephalosome (= cephalothorax) and bears brates as well as marine and freshwater fishes; they are
the usual head appendages and maxillipeds. All of the often very tiny and show a reduction or loss of body
other limbs arise on the remaining thoracic segments, segmentation. Ergasilidans parasitize invertebrates
which together constitute the metasome. The abdomen, and marine fishes and may also show a reduced num-
or urosome, bears no limbs. The appendage-bearing ber of body segments. The Platycopioida are benthic
regions of the body (cephalosome and metasome) are forms known primarily from marine caves; the Gely-
frequently collectively called the prosome. The major- elloida are known only from European groundwaters.
ity of the free-living copepods, and those most fre- Huys and Boxshall (1991) estimated that nearly half of
quently encountered, belong to the orders Calanoida, all known copepod species are parasites.

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Shrimps, and Their Kin 691

Class Thecostraca The 1,300 or so described cirripede species are mostly

This group includes the barnacles, parasitic asco- free-living barnacles, but this group also includes some
thoracidans, and the mysterious Facetotecta (so far, strange parasitic “barnacles” rarely seen except by spe-
known only from their larvae). The thecostracan clade cialists. The common acorn and goose barnacles belong
is defined by several rather subtle synapomorphies of to the superorder Thoracica. Some acorn barnacles are
cuticular fine structure, including cephalic chemosen- known to transform from their oval-shaped aperture to
sory structures known as lattice organs. The group is a narrow or bent aperture when exposed to predatory
also supported by molecular phylogenetic analyses. All snails, making them better able to withstand preda-
taxa have pelagic larvae, the terminal instar of which is tor assaults. The superorder Acrothoracica consists of
a unique larva, the cypris larva (or cyprid), with pre- minute animals that burrow into calcareous substrata,
hensile antennules specialized for locating and attach- including corals and mollusc shells (Figure 21.17G).
ing to the substratum of the sessile adult state. The superorder Rhizocephala are exceptionally modi-
fied parasites of other crustaceans, especially decapods
Subclass Facetotecta Monogeneric (Hansenocaris): (Figure 21.17H).
The “y-larvae,” a half dozen small (250–620 mm) In cyprids the carapace is always present and
marine nauplii and cyprids (Figure 21.17I). Although “bivalved,” the two sides being held by a transverse
known since Hansen’s original description in 1899, the cypris adductor muscle; in adults the carapace may
adult stage of these animals has still not been identi- be lost (Rhizocephala) or modified as a membranous,
fied. However, a stage subsequent to the cypris larva, saclike mantle (thoracicans and acrothoracicans). In the
the sluglike ypsigon stage, has been induced by treat- barnacles (Thoracica), it is this mantle that produces the
ing y-larvae with molting hormones. The prehensile familiar, hard, calcareous plates that enclose the body.
antennules and hooked labrum of the y-cyprids and Cyprids and adult acrothoracicans share a unique tripar-
the degenerative nature of the ypsigon suggest that the tite crystalline cone structure in the compound eye, a fea-
adults are parasitic in yet-to-be-identified hosts. ture not known from any other crustacean group. Most
species of barnacles are hermaphrodites, although sepa-
Subclass Ascothoracida About 125 described species rate sexes are the rule in acrothoracicans and rhizocepha-
of parasites of cnidarians and echinoderms. Although lans, and some androgonochoristic species (males + her-
greatly modified, they retain a bivalved carapace and maphrodites; e.g., Scalpellum) have also been reported.
the full complement of thoracic (11 segments) and Locomotion in barnacles is generally confined to the
abdominal (5 segments) somites. Ascothoracidans larval stages, although adults of a few species are specifi-
generally have mouthparts modified for piercing and cally adapted to live attached to floating objects (e.g., sea-
sucking body fluids, but some live inside other animals weeds, pumice, logs) or nektonic marine animals (e.g.,
and absorb the host’s tissue fluids. In at least one spe- whales, sea turtles). Cyprids can both swim and walk
cies, Synagoga mira, males retain the ability to swim (on the antennules) to locate a suitable site for settlement,
throughout their lives, attaching only temporarily whereupon they cement themselves to the substratum;
while feeding on corals (Figure 21.17F). the paired cement glands constitute a synapomorphy for
cirripedes (ascothoracidan cyprids attach by mechanical
Subclass Cirripedia Primitively with tagmata as in means). Others are often found on the shells and exo-
the class, but in most groups the adult body is modi- skeletons of various errant invertebrates (e.g., crabs and
fied for sessile or parasitic life. Thorax of 6 segments gastropods), which inadvertently provide a means of
with paired biramous appendages; abdomen strongly transportation from one place to another. Thoracican and
reduced or absent; telson absent in most, although cau- acrothoracican barnacles use their feathery thoracopods
dal rami persist on abdomen in some; unique nauplius (called cirri) to suspension feed. Barnacles in the family
larva with frontolateral horns (which house nerves and Coronulidae are suspension feeders that attach to whales
large unicellular glands). All begin life with a series of and turtles (e.g., Chelonibia, Platylepas, Stomatolepas, Coro-
free-swimming naupliar instars (planktotrophic or nula, Xenobalanus). The 300 or so known species of rhizo-
lecithotrophic) terminating in a unique, “bivalved,” cephalans are all endoparasitic in other Crustacea, and
nonfeeding cypris larva (modified for settlement and they are the most highly modified of all cirripedes. They
subsequent metamorphosis into an adult); cyprid with mainly inhabit decapod crustaceans, but a few are known
articulating telson bearing 2 caudal (furcal) rami. Adult from isopods, cumaceans, stomatopods, and even tho-
carapace “bivalved” (folded) or forming fleshy mantle; racican barnacles. Some even parasitize freshwater and
first thoracomere often fused with cephalon and bear- terrestrial crabs. The body consists of a reproductive part
ing maxilliped-like oral appendages; female gonopores (the externa) positioned outside the host’s body, and an
near bases of first thoracic limbs, male gonopore on internal, ramifying, nutrient-absorbing part (the interna).
median penis on last thoracic or first abdominal seg- Most of the externa is female and contains the ovary as
ment; compound eyes lost in adults (Figures 21.1S,T, well as the male “organs” in the form of one or several
21.17A–E, 21.25, 21.26, 21.27B,C, 21.32E, and 21.33F). dwarf males that are implanted into the virginal female.

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 692 Chapter 21

(A) (D)

Baptiste François/Wikipedia/CC BY 4.0

© Larry Jon Friesen
(C)
(B)

(E)

(F)

(G)

(H) (I)
Both courtesy of J. T. Høeg

Brusca 4e
BB4e_21.[Link]
12/30/2021

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 for more ebook/ testbank/ solution manualsArthropoda
Subphylum requests: emailCrabs,
Crustacea: 960126734@[Link]
Shrimps, and Their Kin 693

◀ FIGURE 21.17 Anatomy and diversity in the class 70 to 150 μm (comparable in size to some protists). The
Thecostraca—barnacles and their kin. (A–E) Thoracican unique tantulus stage is adapted to both free swimming
barnacles. (A) A sessile (acorn) barnacle with its cirri extend- (through sediments) and an attached parasitic mode
ed for feeding. (B) Plate terminology in a balanomorph
of life. In the free-swimming phase there is a cepha-
(acorn) barnacle. (C) The lepadomorph (stalked) barnacle
Pollicipes polymerus. (D) Verruca, the “wart” barnacle. lon, 6-segmented thorax, and abdomen with up to 7
(E) Two thoracican barnacles that live in association with each segments. The cephalon lacks appendages (except for
other and with whales. The stalked barnacle Conchoderma unsegmented antennules in sexual females; absent in
attaches to the sessile barnacle Coronula, which in turn larvae) but has an internal median stylet. Natatory tho-
attaches to the skin of certain whales. (F) Anatomy of the racopods 1–5 are biramous, 6 is uniramous; the abdomen
ascothoracidan Ascothorax ophioctenis, a parasite that feeds lacks appendages but bears caudal rami. Paired bundles
periodically on echinoderms (longitudinal section). (G) An
of longitudinal muscles run along the midline in the
acrothoracican barnacle, Trypetesa. Note the highly modi-
fied female and the tiny attached male. This species bores thoracic somites and partly into the abdomen. There is
into calcareous substrata such as coral skeletons. (H) A crab emerging molecular and morphological evidence that
(Carcinus) infected with the rhizocephalan Sacculina carcini. tantulocarids are closely related to thecostracans, and
The crab’s right side is shown as transparent, exposing the they might one day be moved into that class.
ramifying body of the parasite. (I) A cypris y-larva, in lateral Tantulocarids have a most remarkable life cycle.
and dorsal views. (E after P. A. Meglitsch. 1972. Invertebrate Atypically, the tantulus is able to switch its lifestyle
Zoology. Oxford University Press, Oxford.)
from one of mobility to that of a sessile parasite without
further molting. Thus, this one stage represents adapta-
All stages, including the nauplii and cyprids, lack any tions for both motile and attached/sessile lifestyles, and
trace of an alimentary canal, and the parasitic stages lack the transformation from one to the other involves dra-
any segmentation or appendages.6 matic changes in the animal’s physiology and anatomy.
When the free-swimming tantulus finds a proper host, a
unique cuticular organ, the proboscis (sometimes called
Class Tantulocarida the “funnel-shaped organ”), is everted from an oral disc
A small group of about 3 dozen bizarre parasites of to establish an attachment, likely using a type of glue.
mostly deep-water crustaceans, although some are The proboscis is then retracted back into the tantulus,
known from the intertidal zone
and from anchialine pools and
hydrothermal vents (Figures 21.1V (A)
and 21.18). Tantulocarids are the (B)
smallest living crustaceans, at least
in the juvenile or “larval” stage (the
tantulus), which ranges from just
6
Rhizocephalans of the family Sacculinidae
infest only decapod crustaceans and have
been suggested as biological control agents
for invasive exotics such as the green crab
(Carcinus maenas), which are upsetting
coastal ecosystems worldwide. Sacculinds
have the ability to take control over such
major host functions as molting and repro-
duction, and also to compromise the host’s
immune system.
(C) (D) [Link]
From G. A. Boxshall et al. Syst Parasitol 14: 17–30.

FIGURE 21.18 Anatomy in the class

Tantulocarida. (A) An adult Basipodella
atlantica. Note the absence of an
abdomen and the modifications for
Courtesy of G. Boxshall

para­sitic life. (B) Basipodella attached

to the antenna of a copepod host.
(C,D) Microdajus pectinatus on a crus-
tacean host, adult (C), and juvenile (D)
(SEM). (A,B after G. A. Boxshall and
R. J. Lincoln. 1983. J Crust Biol 3: 1–16.)

Brusca 4e
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 694 Chapter 21

and a permanent attachment is made by the oral disc. maxillules and maxillae reduced or absent; no maxilli-
To access the host’s nutrients, the tantulus pierces the peds (Figures 21.1C,D, 21.20, 21.21B, 21.31C, and 21.35B).
victim’s cuticle with a stylet (which has protractor and The branchiopods are difficult to describe in a gen-
retractor muscles), which is subsequently withdrawn eral way. The majority are small freshwater forms with
back into the larval head. The tip of the stylet is solid, leaflike legs and minimal body tagmosis. Most are
and the organ apparently does not inject anything into short-lived, and those inhabiting ephemeral waters
the host. The tiny hole left from this puncture serves as complete their life cycle in just a few weeks. Because
an entrance for a unique rootlet system that grows into of their short life cycle and predilection for tempo-
the host to absorb nutrients. Once attached to a host, rary pools (e.g., vernal pools), many groups produce
the tantulus loses its attachment organs, including the drought-resistant eggs or zygotes, called cysts, which
cement glands, and all of the musculature degenerates can survive years or decades until the next adequate
and is resorbed. It then either sheds the whole trunk and rains appear. As diverse as the branchiopods might
forms a cuticular sac containing a developing partheno- appear, both morphological and molecular analyses
genetic tantulus or female, or retains the trunk somites indicate that they comprise a monophyletic group.
but expands to form a cuticular sac in which a male Development is remarkably uniform across the group,
develops. Thus, the tantulus either enters a partheno- with naupliar larvae distinguishable by several unique
genetic phase (becoming a so-called parthenogenetic features. A few groups have conquered the world oceans
female, which produces more tantuli) or develops into a secondarily (e.g., marine cladocerans). Most species are
sexually reproducing adult. In both cases, the next stage filter feeders, but predators are known both within
develops inside the cuticular sac of the tantulus itself. Anostraca and Cladocera. About 1,130 extant species
Progeny are released into the environment. have been described (around half of which are cladocer-
ans), and at least one fossil branchiopod (Rehbachiella) is
Superclass Altocrustacea: known from the middle Cambrian, confirming that they
Clade Allotriocarida are an ancient group. Although branchiopod taxonomy
is still in a bit of flux, the group is generally divided
Class Cephalocarida into two subclasses, Anostraca (the carapaceless fairy
Head followed by 8-segmented thorax with each seg- shrimps) and Phyllopoda (the carapaced branchiopods).
ment bearing limbs, an 11-segmented limbless abdo-
men, and a telson with caudal rami; common gono-
pore on protopods of sixth thoracopods; carapace Subclass Anostraca
absent but head covered by cephalic shield; thoraco- Postcephalic trunk divisible into appendage-bearing tho-
pods 1–7 biramous and phyllopodous, with large flat- rax of 11 segments (or, rarely, 10, 17, or 19) and abdomen
tened exopods and epipods (exites) and stenopodous of 8 segments plus telson with caudal rami; gonopores
endopods; thoracopods 8 reduced or absent; maxillae on genital region of abdomen; trunk limbs biramous and
resemble thoracopods; no maxillipeds; eyes absent; phyllopodous; small cephalic shield present, though fully
nauplii with antennal glands, adults with maxillary developed carapace lacking; paired, large, stalked com-
glands and (vestigial) antennal glands; with long and pound eyes and a single median simple (naupliar) eye.
gradual (anamorphic) larval development (Figures Anostracans are primitive-looking crustaceans com-
21.1B 21.19A–C, and 21.21A). monly called fairy shrimps and brine shrimps that dif-
Cephalocarids are tiny, elongate crustaceans ranging fer from all other branchiopods in lacking a carapace.
in length from 2 to 4 mm. There are 12 species in five There are just over 300 living species, worldwide, most
genera. All are benthic marine detritus feeders, although of which are less than 1 cm in length, although a few
many seem to have commensal relationships with sed- giants attain lengths of 10 cm. Anostracans inhabit
entary polychaetes. Most are associated with sediments ephemeral ponds (including snowmelt ponds and
covered by a layer of flocculent organic detritus, although desert pools), hypersaline lakes, and marine lagoons.
some have been found in clean sands. They occur from In many areas they are an important food resource for
the intertidal zone to depths of over 1,500 m, from the water birds. Anostracans are sometimes united with
North and South Pacific to the North and South Atlan- the extinct Lipostraca as the “Sarsostraca.” Their fossil
tic and in the Mediterranean. Hatching is at a slightly record dates back to the Devonian.
advanced naupliar stage (called a metanauplius). Anostracans swim ventral side up by metachronal
beating of the trunk appendages. Many use these limb
movements for suspension feeding. Some other species
Class Branchiopoda scrape organic material from submerged surfaces, and
Numbers of segments and appendages on thorax and at least two species (Branchinecta gigas, B. raptor) are spe-
abdomen vary, the latter usually lacking appendages; cialized as predators on other fairy shrimps. Although
carapace present or absent; telson usually with cau- most anostracans live in isolated ponds, their eggs are
dal rami; body appendages generally phyllopodous; transported on the feathers of birds and feet of mammals

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 for more ebook/ testbank/ solution manuals requests: email 960126734@[Link]

(A) (B)

Courtesy of J. Olesen
(C)

(D)

Courtesy of F. Schram
(E)

(F)

Courtesy of F. Schram

FIGURE 21.19 Anatomy in the classes Remipedia the cephalocarids and the remipedes, the trunk is a long,
and Cephalocarida. (A) The cephalocarid Hutchinsoniella homonomous series of somites with biramous swimming
macracantha (lateral view). (B) SEM of head and thorax appendages. In cephalocarids the first trunk appendages
of a cephalocarid. (C) First trunk limb of the cephalocarid are like all the others, which bear large swimming epipods
Lightiella.
Brusca (D)4e
The remipede Morlockia ondinae (ventral (exites). In remipedes the first trunk somite is fused to the
view). (E) Tenth trunk limb of the remipede Lasionectes. head, and its appendages are maxillipeds. See chapter
BB4e_21.[Link]
(F) The anterior end of a remipede (ventral view). In both opener photo of Xibalbanus tulumensis.
12/20/2021

and might also be transported during passage through Order Notostraca Thorax of 11 segments, each
the gut of predatory diving beetles (Dytiscidae). with 1 pair of phyllopodous appendages; abdomen
of “rings,” each formed of more than 1 true segment;
each anterior ring with several pairs of appendages;
Subclass Phyllopoda posterior rings lack appendages; telson with long cau-
The tadpole shrimps (Notostraca) and “bivalved” dal rami; gonopores on last thoracomere; broad, shield-
branchiopods (Diplostraca). like carapace fused only with head, but extending to

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 696 Chapter 21

(A) (C)

(B)

Martin Pelanek/Shutterstock

(F)
Antenna
(G)
Compound eye
Food
string

Rostrum
Heart
Antennule
Labrum
Carapace
Dieter Ebert/Wikipedia/CC BY-SA 4.0

Brood Thoracic
chamber appendage

Midgut

Anus
Postabdomen

(J) (K) From J. W. Martin et al. 1986. Zool Scripta 15: 221–232

Brusca 4e
BB4e_21.[Link]
12/20/2021

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 for more ebook/ testbank/ solution manualsArthropoda
Subphylum requests: emailCrabs,
Crustacea: 960126734@[Link]
Shrimps, and Their Kin 697

(D) (E)

(H) (I)

(M)
(L)

Courtesy of J. Olesen
From J. W. Martin et al. 1986. Zool Scripta 15: 221–232

FIGURE 21.20 Anatomy and diversity in the class

Branchiopoda. (A) An anostracan (Branchinecta) in
swimming posture. (B) An anostracan, Branchipus
schaefferi, swimming. (C) Trunk limb of an anostracan
(Linderiella). (D,E) A notostracan, Triops: (D) dorsal and
(E) ventral views. (F,G) Two cladocerans: (F) Daphnia and
(G) Leptodora. (H) The shed carapace, or ephippium, of
Daphnia, with the embryos enclosed. (I) Two extreme
stages in the seasonal change in head form of Daphnia
(cyclomorphosis). (J–L) A clam shrimp (Laevicaudata),
Lynceus: (J) the valves are partially open (ventral view);
(K) the head (ventral view); (L) the whole animal, with
one valve removed. (M) A cyclestherid, Cyclestheria.
SEM photo, with one valve removed (the specimen is
slightly distorted; in life the shell is round). (C after A.
Thiery and A. Champeau. 1988. J Crust Biol 8: 70–78.)

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 698 Chapter 21

loosely cover thorax and part of abdomen; paired, ses- resemblance to cladocerans and ostracods, and lack of
sile compound eyes and a single simple eye near ante- obvious growth lines on the carapace quickly distin-
rior midline on carapace. guishes them from Spinicaudata and Cyclestherida.
There are only about a dozen living species of tadpole The suborder Onychocaudata contains the spiny-tailed
shrimps, in two genera (Triops and Lepidurus) placed in or “true” clam shrimp (Spinicaudata), the monotypic
a single family, Triopsidae. Most species are 2–10 cm Cyclestherida (Cyclestheria hislopi), and the cladocerans
long. The common name derives from the general body (the water fleas). Cyclestheria is also sometimes called
shape: the broad carapace and narrow “trunk” give the a clam shrimp, though it is more closely related to the
animals a superficial tadpolelike appearance. Cladocera than to Spinicaudata. Despite the confused
Notostracans inhabit inland waters of all salinities, taxonomy of Daphnia (only a quarter of the >360 named
but none occur in the ocean. Of the two known genera, species are valid), it has become a model genus for eco-
Triops (Figure 21.20D,E) lives only in temporary waters, logical, toxicological, and evolutionary studies.
and its eggs are capable of surviving extended dry peri- The common name “clam shrimp” derives from the
ods. Most species of Lepidurus live in temporary ponds, clamlike appearance of the valves, which usually bear
but at least one species (L. arcticus) inhabits permanent concentric growth lines reminiscent of bivalved mol-
ponds and lakes. However, all species are short-lived, luscs. The approximately 200 species of clam shrimps
and most complete their life cycle in just 30 to 40 days. (including laevicaudatans, spinicaudatans, and Cycl-
Triops is of some economic importance in that large estheria) live primarily in ephemeral freshwater habi-
populations often occur in rice paddies and destroy the tats worldwide. Cyclestheria hislopi inhabits permanent
crop by burrowing into the mud and dislodging young freshwater habitats throughout the world’s tropics and
plants. Tadpole shrimps mostly crawl, but they are also is one of the most widespread animals on Earth. Cycles-
capable of swimming for short periods by beating the theria is also the only clam shrimp with direct develop-
thoracic limbs. They are omnivorous, feeding mostly on ment, the larval and juvenile stages being passed within
organic material stirred up from the sediments, although the brood chamber, one of the features allying it with
many species scavenge or prey on other animals, includ- the cladocerans in the infraorder Cladoceromorpha.
ing molluscs, other crustaceans, frog eggs, and even frog In cladocerans, the carapace is never hinged (only
tadpoles and small fishes. Some species of notostracans folded dorsally, like a taco) and never covers the entire
are exclusively gonochoristic, but others may include body, and appendages do not occur on all the trunk
hermaphroditic populations (often those populations somites. The body segmentation is generally reduced.
living at high latitudes). Earlier reports of parthenoge- The thorax and abdomen are fused as a “trunk” bear-
netic populations have been questioned. ing 4–6 pairs of appendages anteriorly and terminating
in a flexed “postabdomen” with clawlike caudal rami.
Order Diplostraca The “bivalved” branchiopods: The trunk appendages are usually phyllopodous. The
clam shrimps and cladocerans (water fleas) (Figure carapace typically encloses the entire trunk, but not
21.20J–L). Large “bivalved” carapace covers all or most the cephalon, serving as a brood chamber (and greatly
of the body; body divided into cephalon and trunk, reduced to this function) in some species; a single
the latter with 10–32 segments, all with appendages, median compound eye is always present.
and with no regionalization into thorax and abdomen; The cladocerans, or water fleas, include about 400
trunk limbs phyllopodous, decreasing in size poste- species of predominantly freshwater crustaceans. Sev-
riorly; males with trunk limbs 1, or 1–2, modified for eral American marine genera and species are known
grasping females during mating; trunk typically ter- (e.g., Evadne, Podon). Although there are relatively few
minates in spinous anal somite or telson, usually with species, the group exhibits great morphological and
robust caudal rami (cercopods); gonopores on eleventh ecological diversity. Most cladocerans are 0.5–3 mm
trunk segment; bivalved carapace completely enclos- long, but Leptodora kindtii reaches 18 mm in length.
es body; valves folded (Onychocaudata) or hinged Except for the cephalon and large natatory antennae,
(Laevicaudata) dorsally; usually with a pair of sessile the body is enclosed by a folded carapace, which is
compound eyes and a single, median, simple eye. fused with at least some of the trunk region. The car-
Most diplostracans are benthic, but many swim dur- apace is greatly reduced in members of the families
ing reproductive periods. Some are direct suspension Polyphemidae and Leptodoridae, in which it forms a
feeders, whereas others stir up detritus from the sub- brood chamber.
stratum and feed on suspended particles, and others Cladocerans are distributed worldwide in nearly all
scrape pieces of food from the sediment. There are two inland waters. Most are benthic crawlers or burrow-
suborders, Laevicaudata and Onychocaudata. ers, while others are planktonic and swim by means of
The suborder Laevicaudata, or flat-tailed clam their large antennae. One genus (Scapholeberis) is typi-
shrimps, contains the single family Lynceidae with cally found in the surface film of ponds, and another
close to 40 freshwater species. They are character- (Anchistropus) is ectoparasitic on Hydra. Most of the
ized by a hinged, globular carapace that encloses the benthic forms feed by scraping organic material from
entire animal. This carapace gives them a superficial sediment particles or other objects; the planktonic

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Shrimps, and Their Kin 699

species are suspension feeders. Some, such as Leptodora from a cavern in the Bahamas, gave the carcinological
and Bythotrephes, are predators on other cladocerans. world a turn. The combination of features distinguish-
In sexual reproduction, fertilization generally occurs ing these creatures is puzzling, for they possess charac-
in a brood chamber between the dorsal surface of the teristics long thought to be primitive (e.g., long, homon-
trunk and the inside of the carapace. Most species omous trunk; double ventral nerve cord; segmental
have direct development. In the family Daphniidae the digestive ceca; cephalic shield) as well as some attributes
developing embryos are retained by a portion of the traditionally recognized as advanced (e.g., maxillipeds;
shed carapace, which functions as an egg case called nonphyllopodous [though flattened], biramous limbs).
an ephippium (Figure 21.20H), whereas in the Chy- They swim about on their backs as a result of metachro-
doridae the ephippium remains attached to the entire nal beating of the trunk appendages, similar to anostra-
shed carapace. Leptodora exhibits a heterogenous life cans. The remipedes are thus reminiscent of two other
cycle, alternating between parthenogenesis and sexual classes, the branchiopods and cephalocarids. However,
reproduction, the latter of which results in free-living the laterally directed limbs are unlike those of any other
larvae (metanauplii hatch from the shed resting eggs). crustacean, and the “internalized” mandibles and the
Cladoceran life histories are often compared with poison-injecting hypodermic maxillules are unique (the
those of animals such as rotifers and aphids. Dwarf complex venom contains neurotoxins, peptidases, and
males occur in many species in all three groups, and chitinases). The presence of the preantennular processes
parthenogenesis is common. Members of two cladoc- is also puzzling, although similar structures are known
eran families that undergo parthenogenesis (Moinidae to occur in a few other crustaceans. Some phylogenetic
and Polyphemidae) produce eggs with very little yolk. analyses based on morphological data suggest that
In these groups the floor of the brood chamber is lined remipedes may be the most primitive living crustaceans,
with glandular tissue that secretes a fluid rich in nutri- whereas phylogenomic data place them unambiguously
ents, which is absorbed by the developing embryos. as the sister group of the Hexapoda.
Periods of overcrowding, adverse temperatures, or food All of the living remipedes discovered thus far are
scarcity can induce parthenogenetic females to produce troglobitic, found in caves, cenotes, lava tubes, and
male offspring. Occasional periods of sexual reproduc- sinkholes (usually with connections to the sea) in the
tion have been shown to occur in most parthenogenetic Caribbean Basin, Canary Islands, and Western Austra-
species. Many planktonic cladocerans undergo seasonal lia. The water in these systems is often distinctly strati-
changes in body form through succeeding generations fied, with a layer of fresh water overlying the denser
of parthenogenetically produced individuals (Figure salt water in which the remipedes swim. Remipedes
21.20I), a phenomenon known as cyclomorphism. hatch as lecithotrophic naupliar larvae, which is also
unusual given their habitat (most cave crustaceans
have direct development). Postnaupliar development
Class Remipedia is anamorphic; juveniles have fewer trunk segments
Body of 2 regions, a short cephalon and an elongate than do the adults. Based on the three pairs of raptorial,
homonomous trunk of up to 32 segments, each with a prehensile cephalic limbs (and direct observations), it
pair of flattened limbs. Cephalon with a pair of sensory was long thought that remipedes were strictly preda-
preantennular frontal processes; first antennae (anten- tors. However, studies by Stefan Koenemann and his
nules) biramous; trunk limbs laterally directed, biramous, colleagues have suggested they might also be capable
paddlelike, but without large epipods; rami of trunk of suspension feeding.
limbs (exopod and endopod) each of 3 or more articles;
without a carapace, but with cephalic shield covering
head; midgut with serially arranged digestive ceca; first
The Crustacean Body Plan
trunk segment fused with head and bearing 1 pair of pre- We realize that the above synopses are rather exten-
hensile maxillipeds; labrum very large, forming a cham- sive, but the diversity of crustaceans demands empha-
ber (atrium oris) in which reside the “internalized” man- sis before we attempt to generalize about their biology.
dibles; maxillules function as hypodermic fangs, injecting The evolutionary success of crustaceans, like that of
venom from associated venom glands; last trunk seg- other arthropods, has been closely tied to modifications
ment partly fused dorsally with telson; telson with caudal of the jointed exoskeleton and appendages, the latter
rami; segmental double ventral nerve cord; eyes absent in having an extensive range of modifications for a great
living species; male gonopore on trunk limb 15, female on variety of functions.
8; up to 45 mm in length. The above diagnosis is for the The most basic crustacean body plan is a head (cepha-
27 known living remipedes (order Nectiopoda); the fos- lon) followed by a long body (trunk) with many simi-
sil record is currently based on a single poorly preserved lar appendages, as seen in the class Remipedia (Figures
specimen (order Enantiopoda) (chapter opener and Fig- 21.1A and 21.19D,E). In the other crustacean classes, how-
ures 21.1A, 21.19D–F, 21.21D, 21.22F, and 21.31E,F). ever, various degrees of tagmosis occur, and the cephalon
The discovery of living remipedes in 1981 by Jill is typically followed by a trunk that is usually divided
Yager, strange vermiform crustaceans first collected into two distinct regions, a thorax and an abdomen. The

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 700 Chapter 21

first segment of the body (ocular segment) and the last Uniformity within the subphylum Crustacea is dem-
segment of the body (telson) are without appendages. onstrated particularly by the consistency of elements of
Many crustacean groups have a cephalic shield (head the cephalon and the presence of the nauplius larva.
shield) or a carapace. The cephalic shield results from the Except for a few cases of secondary reduction, the head
fusion of the dorsal head tergites to form a solid cuticular of all crustaceans has five pairs of appendages. From
plate, often with ventrolateral folds (pleural folds) on the anterior to posterior, these are the antennules (first
sides. Head shields are found in ancient Cambrian fossil antennae), antennae (second antennae), mandibles,
crustaceans (e.g., from the Orsten fauna), and they are maxillules (first maxillae), and maxillae (second max-
characteristic of the classes Remipedia and Cephalocar- illae). The presence of two pairs of antennae is, among
ida; they also occur in a few other groups. The carapace is arthropods, unique to the Crustacea (as is the nauplius
a more expansive structure, composed of the head shield larva, although similar “head larvae” are known from
and a large fold of the exoskeleton that probably arises other arthropod groups in the fossil record).7 Although
(primitively) from the maxillary somite. The carapace the eyes of some crustaceans are simple, most possess
may extend over the body dorsally and laterally as well a pair of well-developed compound eyes, either set
as posteriorly, and it often fuses to one or more thoracic directly on the head (sessile eyes) or borne on distinct
segments, thereby producing a cephalothorax (Figure movable stalks (stalked eyes).
21.2A). Occasionally, the carapace may grow forward In many crustaceans, from one to three anterior
beyond the head as a narrow rostrum. thoracic segments (thoracomeres) are fused with the
Most of the differences among the major groups of cephalon. The appendages of these fused segments
crustaceans, and the basis for much of their classification, are typically incorporated into the head as additional
arise from variations in the number of somites in the tho- mouthparts called maxillipeds. In some cases, anterior
rax and abdomen, the form of their appendages, and the thoracopods may not fuse with the cephalon during
size and shape of the carapace. A brief skimming of the 7
Cambrian crustacean-like larvae in the fossil record termed “head
synopses (above) and the corresponding figures will give larvae” may be precursors to the distinctive nauplius stage seen in
you some idea of the range of variation in these features. many modern crustaceans. See Martin et al. (2014).

TABLE 21.1 Comparison of Distinguishing Features among the 11 Crustacean Classes

(and in the Subclasses of Class Eumalacostraca)
Body Tagmata and Number
Carapace or of Segments in Each
Taxon Cephalic Shield (excluding telson) Thoracopods

Class Ostracoda Carapace, bivalved Subdivisions not clear; 6–8 pairs of limbs Not phyllopodous, reduced
Class Mystacocarida Cephalic shield Cephalon (6), trunk (10) Not phyllopodous
Class Branchiura Carapace broad, Cephalon (6), thorax (6), abdomen (4?) Not phyllopodous (but
covering head all natatory)
and trunk
Class Pentastomida None Not distinguishable; body vermiform None
Class Malacostraca, Large, folded carapace Cephalon (6), thorax (8), abdomen (7) Phyllopodous
subclass Phyllocarida covers thorax
Class Malacostraca, Well-developed Cephalon (6), thorax (8), abdomen (6) Not phyllopodous
subclass Hoplocarida carapace covers
thorax
Class Malacostraca, Carapace well developed Cephalon (6), thorax (8), abdomen (6) Not phyllopodous,
subclass Eumalacostraca or secondarily reduced uniramous in many
or lost
Class Copepoda Cephalic shield Cephalon (6), thorax (6), abdomen (5) Not phyllopodous, natatory,
often reduced
Class Thecostraca Carapace bivalved (at Cephalon (6), thorax (6), abdomen (4) Not phyllopodous,
some stage), often often reduced
modified as a mantle
Class Tantulocarida Cephalic shield Cephalon (6), thorax (6), abdomen Not phyllopodous,
(up to 7) greatly reduced
Class Cephalocarida Cephalic shield Cephalon (6), thorax (8), abdomen (11) Phyllopodous
Class Branchiopoda Cephalic shield or Cephalon (6), thorax (usually 10–32), Phyllopodous
carapace abdomen (8 to many)
Class Remipedia Cephalic shield Cephalon (6), trunk (up to 32) Not phyllopodous

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development, but still become modified to move food The abdomen, called a pleon in malacostracans, is
to the mouth; these, too, are usually called maxillipeds. composed of several segments, or pleonites (= pleo-
In the class Malacostraca, the remaining free thoraco- meres), followed by a “postsegmental” plate or lobe,
meres are together termed the pereon. Each segment of the telson or anal somite, bearing the anus (Figure
the pereon is called a pereonite (= pereomere), and the 21.2B). In some crustaceans this anal somite bears a
appendages on those segments are called pereopods. pair of appendage-like or spinelike processes conven-
The pereopods may be specialized for walking, swim- tionally called caudal rami. In the Eumalacostraca, the
ming, gas exchange, feeding, and/or defense. Crusta- anal somite lacks caudal rami and is followed by a flat-
cean thoracic (and pleonal) appendages might be prim- tened flap; this flap is often referred to as the telson.
itively biramous, although the uniramous condition is In general, distinctive abdominal appendages (pleo-
seen in a variety of taxa. The general crustacean limb is pods) occur only in the malacostracans. These append-
composed of a basal protopod (= sympod), from which ages are almost always biramous, and often they are
may arise medial endites (e.g., gnathobases), lateral flaplike and used for swimming (e.g., Figures 21.10–
exites (e.g., epipods), and two rami, the endopod and 21.16). The posteriorly directed last pairs of abdominal
exopod. Members of the classes Remipedia, Cephalo- appendages are usually different from the other pleo-
carida, and Branchiopoda and some members of the pods and are called uropods. Together with the telson,
class Ostracoda possess appendages with uniarticulate the uropods form a distinct tail fan in many malacos-
(single-segment) protopods; the remaining classes usu- tracans (Figure 21.2B).
ally have appendages with multiarticulate protopods Crustaceans produce a characteristic larval stage
(Table 21.1).8 called the nauplius (Figures 21.25B,C and 21.33D).
Even upper Cambrian fossil nauplii (Bredocaris admi-
8
The term “peduncle” is a general name often applied to the basal rabilis, Rehbachiella kinnekullensis) are nearly indistin-
portion of certain appendages; it is occasionally (but not always)
used in a way that is synonymous with “protopod.” As noted in
guishable from modern-day nauplii. In their simplest
Chapter 20, the exopod might be no more than a highly modified form, nauplii are oval or pear-shaped, lack external
exite that evolved from an ancestral uniramous condition segmentation, and have three pairs of appendages

Maxillipeds Antennules Antennae Compound Eyes Abdominal Appendages

None Uniramous Biramous Absent (typically) None

1 pair Uniramous Biramous Absent None
None Uniramous, Biramous, reduced Present None
reduced

None None None Absent None

None Biramous Uniramous Present Pleopods
(posteriorly reduced)
5 pairs of thoracopods Triramous Biramous Present, well developed Well developed pleopods
referred to as with gills, 1 pair of
maxillipeds uropods
0–3 pairs Uniramous or Uniramous or biramous Present, well developed Usually 5 pairs of pleopods,
biramous 1 pair of uropods

Usually 1 pair Uniramous Uniramous or biramous Absent None

None Uniramous Biramous Absent (in adults) None

None None (paired in None Absent None

one stage)
None Uniramous Biramous Absent None
None Uniramous Uniramous, biramous, or Present None, or present but often
vestigial posteriorly reduced
1 pair Biramous Biramous Absent All trunk appendages similar

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 702 Chapter 21

that will become the adult antennules, antennae, and nauplius eye (= naupliar eye) that may persist into
mandibles. The antennules are uniramous and primar- adulthood in some taxa. The nauplius eye is actually a
ily sensory. The antennae and mandibles are almost cluster of three or four median eyes. The body is often
always biramous and usually natatory and feed- covered with a dorsal cephalic (naupliar) shield, some-
ing limbs. There is usually a single, anterior eye, the times called a head shield, that commonly expands

TABLE 21.2 Summary of Crustacean Reproductive Features in Major Groups

Development Type, or Hermaphroditic (at Parthenogenesis (in at
Taxon Larval Type at Hatching Least Some Species) Gonochoristic Least Some Species)

Class Ostracoda Direct development, or No Yes Yes

with bivalved nauplius/
metanauplius with
anamorphic development
Class Mystacocarida Metanauplius (partly No Yes No
anamorphic)
Class Branchiura Metanauplius-like (partly No Yes No
anamorphic), or with
direct development
Class Malacostraca, Direct development No Yes No
subclass Phyllocarida
Class Malacostraca, Zoea larva (“antizoea” No Yes No
subclass Hoplocarida or “pseudozoea”)
(metamorphic)

Class Malacostraca, Direct development No Yes No

subclass Eumalacostraca,
superorder Syncarida
Class Malacostraca, Direct development No Yes No
subclass Eumalacostraca,
superorder Peracarida

Class Malacostraca, Nauplius (metamorphic) No Yes No

subclass Eumalacostraca,
superorder Eucarida,
order Euphausiacea
Class Malacostraca, Prezoea or zoea larva, Rare Yes No
subclass Eumalacostraca, and with a nauplius
superorder Eucarida, in Dendrobranchiata
order Decapoda (metamorphic), or direct
development
Class Copepoda Nauplius (metamorphic) No Yes No

Class Thecostraca ? Yes Yes No

Class Tantulocarida Nauplius? + tantulus No Yes ?
(metamorphic)

Class Cephalocarida Metanauplius (anamorphic) Yes No No

Class Branchiopoda Nauplius or metanauplius Yes/no Yes Yes

(anamorphic), or direct
development

Class Remipedia Nauplius (anamorphic) Yes No No

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over the thorax of the adult as a carapace. Behind the with larval hatching taking place at some postnaupliar
head segments is a growth zone and the telson. stage (Table 21.2). Often other larval stages follow the
In many groups (e.g., Peracarida, most Decapoda), nauplius (or other hatching stage) as the individual
the free-living nauplius larva is absent or suppressed. In passes through a series of molts, during which segments
such cases, development is either fully direct or mixed, and appendages are gradually added. A recent compi-
lation of all crustacean larval forms (Martin et al.
2014) includes keys to the distinctive naupliar lar-
vae in all groups that hatch as a nauplius as well as
synopses of larval development in all crustaceans.
Comments

Embryos usually deposited directly on substratum; many Locomotion

myodocopans and some podocopans brood embryos
between valves until hatching as reduced adults; no Crustaceans move about primarily by use of
metamorphosis; up to 8 preadult instars. their limbs (Figure 21.21), and lateral body undu-
Little studied, but apparently the eggs are laid free and up to lations are not known. They crawl or swim, or
7 preadult stages may occur. more rarely burrow, “hitchhike,” or jump. Many
Embryos are deposited; only Argulus is known to hatch as metanauplii; of the ectoparasitic forms (e.g., branchiurans,
others have direct development and hatch as juveniles. certain isopods and copepods) are largely sed-
entary on their hosts, and most cirripedes are
Undergo direct development in the female brood pouch, fully sessile.
emerging as a postlarval/prejuvenile stage called a “manca.”
Swimming is usually accomplished by a row-
Eggs brooded or deposited in burrow; hatch late as clawed ing action of the limbs. Archetypical swimming is
pseudozoea larvae, or earlier as unclawed antizoea larvae.
Both molt into distinct ericthus (and some into alima) larvae
exemplified by crustaceans with relatively undif-
before settling on bottom as postlarvae or juveniles. ferentiated trunks and high numbers of similar
All free larval stages have been lost; eggs deposited on substratum.
biramous appendages (e.g., remipedes, anos-
tracans, notostracans). In general, these animals
swim by posterior to anterior metachronal beat-
Embryos brooded in marsupium of female, typically formed from ing of the trunk limbs (Figure 21.22 and Chapter
ventral coxal plates called oostegites; usually released as mancas 20). The appendages of such crustaceans are often
(subjuveniles with incompletely developed eighth thoracopods). broad and flattened, and they usually bear fringes
Brood pouch (marsupium) in Thermosbaenacea formed by dorsal
carapace chamber. of setae that increase the effectiveness of the power
stroke. On the recovery stroke the limbs are flexed,
Embryos shed or briefly brooded; typically undergo nauplius-
metanauplius-calyptopis-furcilia-juvenile developmental series. and the setae may collapse, reducing resistance.
In members of some groups (e.g., cephalocarids,
branchiopods, leptostracans), large exites or epi-
Dendrobranchiata shed embryos to hatch in water as nauplii or pods arise from the base of the leg, producing
prezoea larvae; all others brood embryos (on pleopods), which broad, “leafy” limbs called phyllopodia. These
do not hatch until a prezoeal or zoeal stage (or later).
flaplike structures aid in locomotion and may
also serve as osmoregulatory (branchiopods) or
Usually with 6 naupliar stages leading to a second series of 5 “larval”
gas exchange (cephalocarids and leptostracans)
stages called copepodites. surfaces (Figure 21.21A–C). Although such epi-
Six naupliar stages followed by a unique larval form called a cypris larva. pods increase the surface area on the power stroke,
they also are hinged so that they collapse on the
A benthic nonfeeding stage, possibly a nauplius, has been reported
but not formally described. Development entails complex recovery stroke, reducing resistance. Metachro-
metamorphosis with infective “tantulus larva.” nal limb movements are retained in many of the
One or 2 eggs at a time are fertilized and carried on genital “higher” swimming crustaceans, but they tend
processes of the first pleonites. to be restricted to selected appendages (e.g., the
Anostraca: embryos usually shed from ovisac early in development; pleopods of shrimps, stomatopods, amphipods,
resistant (cryptobiotic) fertilized eggs accommodate unfavorable and isopods; the pereopods of euphausiaceans
conditions. and mysidans). In swimming euphausiaceans and
Notostraca: eggs briefly brooded, then deposited on substratum;
mysidans the thoracopods beat in a metachronal
resistant (cryptobiotic) fertilized eggs accommodate
unfavorable conditions. rowing fashion. The movements and neuromus-
Diplostraca: most cladocerans undergo direct development cular coordination of crustacean limbs are deceiv-
(Leptodora hatches as nauplii or metanauplii). Clam shrimps carry ingly complex. In the common lophogastridan
developing embryos on the thoracopods prior to releasing them Neognathophausia ingens, for example, 12 separate
as nauplii or metanauplii.
muscles power the thoracic exopod alone (3 that
Eight naupliar stages (2 orthonauplii and 6 metanauplii), followed
are extrinsic to the exopod, 5 in the limb peduncle,
by a “prejuvenile” stage, have been reported.
and 4 in the exopodal flagellum).

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 704 Chapter 21

(A) (B) (C)

(D) (E) (F)

(G)

(H)
(I)

Brusca 4e
BB4e_21.[Link]
5/14/2021

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◀ FIGURE 21.21 Generalized thoracic appendages of that in four groups of crustaceans (cephalocarids, bran-
various crustaceans. (A–C) Biramous, phyllopodous thora- chiopods, phyllocarids, and remipedes) the protopod is
copods. (A) Cephalocarida. (B) Branchiopoda. Dashed composed of a single article. And in branchiopods and
lines indicate fold or “hinge” lines. (C) Leptostraca leptostracans the articles of the endopod are not clearly
(Phyllocarida). (D) A biramous, flattened, but nonphyllo­ separated from one another. In other crustaceans
podous thoracopod (Remipedia). (E–I) Stenopodous (Malacostraca and former “maxillopodans”) the protopod
thoracopods. (E) Euphausiacea. (F) Caridea (Decapoda). comprises two or three separate articles, although in most
(G) Lophogastrida (Peracarida). (H) Spelaeogriphacea former maxillopodans these may be reduced and not easi-
(Peracarida). (I) Isopoda (Peracarida). Because of the pres- ly observed. In the lophogastridans (G), the large marsupi-
ence of large epipods on the legs of the cephalocarids, al oostegite characteristic of most female peracarids is
branchiopods, and phyllocarids, some authors refer to shown arising from the coxa. In two groups (amphipods
them as “triramous” appendages. However, smaller and isopods) all traces of the exopods have disappeared,
epipods also occur on many typical “biramous” legs, so and only the endopod remains as a long, powerful, unira-
this distinction seems unwarranted (and confusing). Note mous walking leg.

Recall from our discussions in Chapters 3 and 20 approaching 1, do setose appendages (e.g., the feed-
that at the low Reynolds numbers at which small ing cirri of barnacles) begin to act as filters, or rakes,
crustaceans (such as copepods or larvae) swim about, as the surrounding water acts less viscous and the
the netlike setal appendages act not as a filtering net, boundary layer is relatively thinner. Of course, the
but as a paddle, pushing water in front of them and closer together the setae and setules are placed, the
dragging the surrounding water along with them more likely it is that their individual boundary layers
due to the thick boundary layer adhering to the limb. will overlap; thus densely setose appendages are more
Only in larger organisms, with Reynolds numbers likely to act as paddles.

(A) (B)

(C)

(D)

(F)

(E) Photo by D. Williams, courtesy of J. Yager

FIGURE 21.22 Some aspects of locomotion (and feeding) in three crustaceans (also
see Chapter 20). (A,B) Generation of swimming and feeding currents in an anostracan.
(A) An anostracan swimming on its back by metachronal beating of the trunk limbs. The
limbs are hinged to fold on the recovery stroke, thereby reducing resistance. (B) Water is
drawn from anterior to posterior along the midline and into the interlimb spaces, and food
particles are trapped on the medial sides of the endites; excess water is pressed out later-
ally, and the trapped food is moved anteriorly to the mouth. (C–E) Locomotion in the post-
larva of Panulirus argus. (C) Normal swimming posture when moving forward slowly.
(D) Sinking posture with appendages flared to reduce sinking rate. (E) A quick retreat by
rapid tail flexure (the “caridoid escape reaction”), a method commonly employed by crusta-
ceans with well-developed abdomens and tail fans. (F) A swimming remipede, Lasionectes.
Note the metachronal waves of appendage movement. (C,E after M. D. Calinski and W. G.
Lyons. 1983. J Crust Biol 3: 329–335.)

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 706 Chapter 21

Not all swimming crustaceans move by metachro- over 1,000 km upriver from the sea—testimony to their
nal waves of limb action. Certain planktonic copepods, superb locomotory ability. Not unexpectedly, E. sinensis
for example, move haltingly and depend on their long is also an important (and destructive) invasive species
antennules and dense setation for flotation between in North America and Europe. It has been accidentally
movements (Figure 21.4F). Watch living calanoid cope- introduced into the Great Lakes several times but has
pods, and you will notice that they may move slowly, not yet been able to establish a permanent population.
by use of the antennae and other appendages, or in Recent work suggests that stomatopods have the ability
short jerky increments, often sinking slightly between to accurately return to their “home” burrow using navi-
these movements. The latter type of motion results from gation similar to that seen in the terrestrial environment
an extremely rapid and condensed metachronal wave by social hymenopterans, spiders, rodents, fiddler crabs,
of power strokes along the trunk limbs. Although the etc.—through a process known as “path integration.”
long antennae may appear to be acting as paddles, they During path integration an animal navigates back on a
actually collapse against the body an instant prior to the path previously taken by monitoring the position of the
beating of the limbs, thus reducing resistance to forward sun and overhead polarization patterns in combination
motion. Some other planktonic copepods create swim- with an internal “odometer.”
ming currents by rapid vibrations of cephalic append- Most walking crustaceans can also reverse the
ages, by which the body moves smoothly through the direction of leg action and move backward, and most
water. Some copepods execute rapid escape responses, brachyuran crabs can walk sideways. Brachyuran
or jumps, when they encounter “upstream” movement crabs are perhaps the most agile of all crustaceans.
that may indicate an approaching predator. These escape The extreme reduction of the abdomen in this group
responses accelerate them to 200 body lengths per sec- allows for very rapid movement because adjacent
ond within milliseconds and, in part, are due to the pres- limbs can move in directions that avoid interference
ence of myelin-like sheaths on key nerve fibers (notably with one another (and much the same thing has hap-
in the antennules). “Rowing” occurs in the swimming pened, independently, in many anomurans with
crabs (family Portunidae) and some deep-sea asellote reduced abdomens). Brachyuran crab legs are hinged
isopods (e.g., family Eurycopidae), both of which use in such a way that most of their motion involves lat-
paddle-shaped posterior thoracopods to scull about. eral extension (abduction) and medial flexion (adduc-
Most eumalacostracans with well-developed abdo- tion) rather than rotation frontward and backward. As
mens exhibit a form of temporary, or “burst,” swimming a crab moves, its limbs move in various sequences, as
that serves as an escape reaction (e.g., mysidans, synca- in normal crawling, but those on the leading side exert
rids, euphausiaceans, shrimps, lobsters, and crayfish). their force by flexing and pulling the body toward the
By rapidly contracting the ventral abdominal (flexor) limb tips, while the opposite, trailing, legs exert pro-
muscles, such animals shoot quickly backward, the pulsive force as they extend and push the body away
bent-down, spread tail fan providing a large propulsive from the tips. Still, this motion is simply a mechani-
surface (Figure 21.22C–E). This behavior is sometimes cal variation on the common arthropodan walking
called a tail flip, or “caridoid escape reaction.” behavior. Many crustaceans move into mollusc shells
Surface crawling by crustaceans is accomplished by or other objects, carrying these about as added protec-
the same general sorts of leg movements seen in insects tion. In most cases, the exoskeleton of the crustacean is
and other arthropods: by flexion and extension of the reduced, especially the abdomen (e.g., hermit crabs).
limbs to pull or push the animal forward. Walking Of course, crustaceans grow in size as they go through
limbs are typically composed of relatively stout, more their molts, so these mobile-home-carrying crustaceans
or less cylindrical articles (i.e., stenopodous limbs), as must continually find larger shells to inhabit. Many
opposed to the broader, often phyllopodous limbs of hermit crabs assemble in congregations to exchange
swimmers (see Figure 21.21 for a comparison of crus- shells in a sort of group “passing of the shells” as they
tacean limb types). Walking limbs are lifted from the move into vacated larger abodes.
substratum and moved forward during their recovery In addition to these two basic locomotor methods
strokes; then they are placed against the substratum, (“typical” walking and swimming by metachronal beat-
which provides purchase as they move posteriorly ing of limbs), many crustaceans move by other special-
through their power strokes, pulling and then push- ized means. Ostracods, cladocerans, and clam shrimps
ing the animal forward. Like many other arthropods, (Diplostraca), most of which are largely enclosed by
crustaceans generally lack lateral flexibility at the body their carapaces (Figures 21.3 and 21.20F,G,J,L,M), swim
joints, so turning is accomplished by reducing the stride by rowing with the antennae. Mystacocarids crawl
length or movement frequency on one side of the body, in interstitial water using various head appendages.
toward which the animal turns (like a tractor or tank Most semiterrestrial amphipods known as “beach
slowing one tread). Many crustaceans migrate; perhaps hoppers” (e.g., Orchestia and Orchestoidea) execute dra-
the most famous is the Chinese mitten crab (Eriocheir matic jumps by rapidly extending the urosome and its
sinensis), which spends most of its life in fresh water but appendages (uropods), reminiscent of the jumping of
returns to the sea to breed. These crabs have been found springtails described in Chapter 22. Most caprelloidean

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amphipods (Figure 21.16E) move about in inchworm exhibit various migratory behaviors, employing their
fashion, using their subchelate appendages for cling- locomotor skills to avoid stressful situations or to remain
ing. There are also a number of crustacean burrowers, where conditions are optimal. A number of planktonic
and even some that build their own tubes or “homes” crustaceans undertake daily vertical migrations, typi-
from materials in their surroundings. Many benthic cally moving upward at night and to greater depths
amphipods, for example, spin silk-lined mud burrows during the day. Such vertical migrators include various
in which they reside. At least one species, Pseudam- copepods, cladocerans, ostracods, and hyperiid amphi-
phithoides incurvaria, constructs and lives in an unusual pods (the latter may make their migrations by riding on
“bivalved pod” cut from the thin blades of the same their gelatinous hosts). Such movements place the ani-
alga on which it feeds (Figure 21.23A). Another amphi- mals in their near-surface feeding grounds during the
pod, Photis conchicola, actually uses empty gastropod dark hours, when there is probably less danger of being
shells in a fashion similar to that of hermit crabs (Fig- detected by visual predators. In the daytime, they move
ure 21.23B). “Hitchhiking” (phoresis) occurs in various to deeper, perhaps safer, waters. These crustaceans can
ectosymbiotic crustaceans, including isopods that par- form enormous shoals that contribute to the deep scat-
asitize fishes or shrimps and hyperiidean amphipods tering layer seen on ships’ sonar. Many intertidal crus-
that ride on gelatinous drifting plankters. taceans use their locomotor abilities to change their
In addition to simply getting from one place to another behaviors with the tides. Crab larvae in particular are
in their usual day-to-day activities, many crustaceans known to migrate upward or downward according to

(A) (B)

NO67/Wikipedia/CC BY-SA 4.0

(C)

FIGURE 21.23 Amphipod “houses.” (A) The complex sequence

of steps in the construction of a “bivalve pod” from the brown alga
Dictyota by the Caribbean amphipod Pseudamphithoides incurvaria:
(1) Initiation of cut and notch; the upper flap of the alga forms the
first “valve.” (2) Continuation of the cut across the algal thallus.
(3) Measuring and clearing algal hairs off the second branch tip.
(4) Cutting of the second valve. (5) Completed “pod” with valves
attached along margins by threadlike secretions. (B) The gammarid-
ean amphipod Grandiderella produces silk-lined tubes in which the
male and female reside. (C) Photis conchicola, a temperate eastern
Pacific amphipod that spins its silken tube inside a minute snail shell,
which is then carried about in the style of hermit crabs. (C from
J. W. Carter. 1982. J Crust Biol 2: 328–341.)

Brusca 4e
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 708 Chapter 21

daily rhythms, taking advantage of incoming or outgo- crustaceans live in a world of low Reynolds numbers,
ing tides to move in and out of estuaries. Some anomu- a world dominated by viscosity rather than by inertia.
ran and brachyuran crabs simply move in and out with Thus, the setose mouth appendages behave more like
the tide, or seek shelter beneath rocks when the tide is paddles than like sieves, with a water layer near the
out, thus avoiding the problems of air exposure. One of limb adhering to it and forming part of the “paddle.”
the most interesting locomotor behaviors among crusta- As the maxillae move apart, parcels of water contain-
ceans is the mass migration of the spiny lobster, Panuli- ing food are drawn into the interlimb space. As the
rus argus, in the Gulf of Mexico and northern Caribbean. maxillae press together, the “parcel” is moved forward
Each autumn, lobsters queue up in single file and march to the endites of the maxillules, which push it into the
in long lines across the seafloor for several days. They mouth. Thus, food particles are actually not filtered
move from shallow areas to the edges of deeper oceanic from the water, but captured in small parcels of water.
channels. This behavior is apparently triggered by win- High-speed cinematography indicates that copepods
ter storm fronts moving into the area, and it may be a are capable of capturing individual algal cells, one at
means of avoiding rough water conditions in the shal- a time, by this “hydraulic vacuum” method. In fact,
lows. At the beginning of the wet season, the endemic copepods may be fairly selective about what they con-
Christmas Island red crabs (Gecarcoidea natalis) under- sume, which includes everything from protistan micro-
take their annual mass migration from the forest to shore plankton (e.g., diatoms) to other small crustaceans.
(up to 8 km) to deposit their eggs in the sea. Sessile thoracican barnacles feed by using their long,
feathery, biramous thoracopods, called cirri, to filter
feed on suspended material from the surrounding water
Feeding (Figure 21.17A,C,E). Studies indicate that barnacles are
With the exception of ciliary mechanisms, crustaceans capable of trapping food particles ranging from 2 μm to
have exploited virtually every feeding strategy imag- 1 mm, including detritus, bacteria, algae, and various
inable (and some “unimaginable”). Even without cilia, zooplankters. Many barnacles are also capable of prey-
many crustaceans generate water currents and engage ing on larger planktonic animals by coiling a single cir-
in various types of suspension feeding. We have selected rus around the prey, in tentacle fashion. In slow-moving
a few examples to demonstrate the range of feeding or very quiet water, most barnacles feed actively by
mechanisms that occur in this group. In most cases, extending the last three pairs of cirri in a fanlike man-
suspension-feeding Crustacea capture and consume par- ner and sweeping them rhythmically through the water.
ticles in the <100 μm size (unicellular algae, invertebrate The setae on adjacent limbs and limb rami overlap to
eggs and larvae, bacteria, suspended organic detritus). form an effective filtering net. The first three pairs of
In some crustaceans the action of the thoracic cirri serve to remove trapped food from the posterior
limbs simultaneously creates the swimming and cirri and pass it to the mouthparts. In areas of high water
suspension-feeding currents. As the metachronal movement, such as wave-swept rocky shores, barnacles
wave of appendage motion passes along the body, often extend their cirri into the backwash of waves,
adjacent limb pairs are alternately moved apart and allowing the moving water to simply run through the
then pressed together, thus changing the size of each “filter,” rather than actively moving the cirri through the
interlimb space (Figure 21.22A,B; see also Chapters 3 water. In such areas you will often see clusters of bar-
and 20). Surrounding water is drawn into an interlimb nacles in which all the individuals are oriented similarly,
space as the adjacent appendages move away from taking advantage of this labor-saving device.
one another, and waterborne particles are trapped Most krill (euphausiaceans) feed in a fashion similar
by setae on the endites as the appendages then press to that of barnacles, but while swimming. The thora-
together. From here, the trapped particles are moved to copods form a “feeding basket” that expands as the
a midventral food groove and then anteriorly, toward legs move outward, sucking food-laden water in from
the head. This mechanism of forming a boxlike “filter the front. Once inside the basket, particles are retained
press” with setose phyllopodous limbs is the typical on the setae of the legs as the water is squeezed out
suspension-feeding strategy of cephalocarids, most laterally. Other setae comb the food particles out of the
branchiopods, and many malacostracans. “trap” setae, while yet another set brushes them for-
Planktonic copepods were long thought to “filter” ward to the mouth region.
feed by generating lateral feeding gyres or currents by Anomuran sand crabs (e.g., Emerita, Lepidopa, Bleph-
movements of the antennae and mouth appendages. It aripoda) use their long, setose antennae in a fashion
was believed that these gyres swept in small particles similar to that of barnacle cirri that “passively” strain
that were directly filtered by the maxillae. This classic wave backwash (Figure 21.24; see also Chapter 3).
idea of maxillary filtration, built on work by H. G. Can- Sand crabs are adapted to living on wave-swept sand
non in the 1920s, has been questioned by recent work, beaches. Their compact oval shape and strong append-
but the model persists and is still commonly presented ages facilitate burrowing in the unstable substratum.
in general works. As mentioned in Chapter 3, we They burrow posterior end first in the area of wave
now know that copepods and other small planktonic wash, with the anterior end facing upward. Following

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Shrimps, and Their Kin 709
(A)
Antennae

Antennules

(C) (D)

(B)

© Larry Jon Friesen

FIGURE 21.24 Some crustacean feeding mechanisms

(see also Figure 21.22). (A) Suspension feeding in the sand
crab Emerita. The arrows point seaward and indicate the
(E)
direction of water movement as waves recede. The anten-
nules direct water through branchial chambers. The antennae
remove food particles from the water and then brush them
onto the mouthparts. (B) The sand crab Blepharipoda occi­
dentalis suspension feeding with its feathery antennae; the
antennules are interlocked with stiff setae that form a chan-
nel between them for water to circulate down to the gills.
(C,D) The suspension-feeding hermit crabs (C) Lophopagurus
(Australeremus) cooki and (D) Areopaguristes pilosus twirl their
antennae, in either a circle or a figure eight, to create water
currents that pull food particles to the mouth region. (E) The
predatory shrimp Procaris ascensionis (Caridea) is shown here
munching on another shrimp (Typhlatya) as it holds the prey
in a “cage” formed by the pereo­podal endopods. (E from
L. G. Abele and B. E. Felgenhauer. 1985. J Crust Biol 5: 15–24.)

a breaking wave, as the water rushes seaward down

the beach, Emerita unfurls its antennae into the mov-
ing water along the surface of the sand. The fine setose
mesh traps protists and phytoplankton from the water,
and the antennae then brush the collected food onto
the mouthparts. Many porcelain and hermit crabs also
engage in suspension feeding. By twirling their anten-
nae in various patterns, these anomurans create spiral- feed within their burrows. They drive water through
ing currents that bring food-laden water toward the the burrow by beating the pleopods, and the first two
mouth
Brusca (Figure
4e 21.24B,C). Food particles then become pairs of pereopods remove food with medially directed
entangled on setae of the mouth appendages and are setal brushes. The maxillipeds then comb the captured
BB4e_21.[Link]
brushed into the mouth by the endopods of the third particles forward to the mouth.
12/21/2021
maxillipeds. Many of these animals also feed on detri- Most other crustacean feeding mechanisms are
tus by simply picking up particles with their chelipeds. less complicated than suspension feeding and usually
Some mud, ghost, and lobster shrimps, such as Cal- involve direct manipulation of food by the mouthparts
lianassa and Upogebia (Axiidea, Gebiidea), suspension and sometimes the pereopods, especially chelate or

21_Brusca4e_CH21.indd 709 3/8/22 1:09 PM

 710 Chapter 21

subchelate anterior legs. However, even in these cases injection of venom from the hypodermic maxillules. It
of direct manipulation, the sheer number of crusta- is suspected that tissues are then sucked out of the prey
cean appendages adapted for feeding makes “simple” by action of a mandibular mill and muscular foregut.
feeding a surprisingly complex affair, with the various Remipedes are probably also facultative suspension
mouthparts taking on tasks that include tasting (via sen- feeders and scavengers.
sory setae), holding, chewing, scraping, and macerating. Another fascinating adaptation for predation can
Many small crustaceans may be classified as microph- be seen in many species of Alpheidae (the snapping
agous selective deposit feeders, employing various shrimps, e.g., Alpheus, Synalpheus) (Figure 21.10D). In
methods of removing food from the sediments in which such species, one of the chelipeds is much larger than
they live. Mystacocarids, many harpacticoid copepods, the other, and the movable finger is hinged in such
and some cumaceans and gammaridean amphipods are a way that it can be held open under muscle tension
referred to as “sand grazers” or “sand lickers.” By vari- and then snapped closed quickly; this forceful clos-
ous methods these animals remove detritus, diatoms, ing produces a loud popping sound and a pressure
and other microorganisms from the surfaces of sediment or “shock” wave in the surrounding water. Some spe-
particles. Interstitial mystacocarids, for example, sim- cies appear to use this mechanism in ambushing prey
ply brush sand grains with their setose mouthparts. On (although most alpheids are probably omnivores and
the other hand, some cumaceans pick up an individual even include algae in their diet). When feeding in a
sand grain with their first pereopods and pass it to the predatory mode, the shrimp sits at its burrow entrance
maxillipeds, which in turn rotate and tumble the particle with the antennae extended. When a potential prey
against the margins of the maxillules and mandibles. approaches (usually a small fish, crustacean, or anne-
The maxillules brush and the mandibles scrape, remov- lid), the shrimp “pops” its cheliped, and the resulting
ing organic material. Some sand-dwelling isopods may pressure wave stuns the victim, which is then quickly
employ a similar feeding behavior. pulled into the burrow and consumed. These shrimps
Predatory crustaceans include stomatopods, remi- typically live in male-female pairs within the burrow,
pedes, and most lophogastridans, as well as many spe- and prey captured by one individual is shared with
cies of anostracans, cladocerans, copepods, ostracods, its partner. Two mechanisms have been proposed for
cirripedes, anaspidaceans, euphausiaceans, decapods, the production of the “pop” and associated shock
tanaidaceans, isopods, and amphipods. Predation wave. In some species the pop seems to be created by
typically involves grasping the prey with chelate or mechanical impact of the dactylus hammering into the
subchelate pereopods (or sometimes directly with the propodus. A second mechanism recently proposed is
mouth appendages), or even with the antennae in the the collapse of cavitation bubbles, which are created
case of predatory anostracans, followed by tearing, by the rapid closure of the claw (in excess of 100 km/
grinding, or shearing with various mouthparts, partic- sec). Cavitation occurs in liquids when bubbles form
ularly the mandibles. Perhaps the most highly adapted and then implode around an object (it is a well-known
predatory specialists are the stomatopods (Figure 21.8), phenomenon damaging ship propellers). Both mech-
which possess greatly enlarged, raptorial subchelate anisms create shock waves. Cavitation shocks are
limbs, which they use to stab or to club and smash prey. also used by some stomatopods while feeding. Some
Some species search out prey, but many sit in ambush boring (endolithic) alpheids even use the claw snap
at their burrow entrance. The actual attack generally to break off pieces of the rock into which they are
follows visual detection of a potential prey item, which digging. In some areas of the world, populations of
may be another crustacean, a mollusc, or even a small snapping shrimps are so large that their noise dis-
fish. Once captured and stunned or killed by the rapto- turbs underwater communication. In the Caribbean,
rial claws, the prey is held against the mouthparts and colonies of alpheids have been likened to those of
shredded into ingestible pieces. social insects (e.g., Synalpheus regalis colonies in Belize
Although the cave-dwelling and often presumed inhabit sponges and contain up to 350 sibling males
“primitive” shrimp Procaris is omnivorous, its predatory and females, with a single dominant breeding female).
behavior is particularly interesting. Its prey includes Many crustaceans emerge from the benthos under
other crustaceans, particularly amphipods and shrimps. cover of darkness to feed or mate in the water column.
After Procaris locates a potential victim (probably by che- Many predatory isopods emerge at night to feed on
moreception), it moves quickly to the prey and grasps invertebrates or fish, particularly weak or diseased fish
it within a “cage” formed by the pereopodal endopods (or fish caught in fishing nets).
(Figure 21.24E). Once captured, the prey is eaten while Macrophagous herbivorous and scavenging crusta-
the shrimp swims about. Apparently the third maxil- ceans generally feed by simply hanging onto the food
lipeds press the prey against the mandibles, which bite source and biting off bits with the mandibles (a feed-
off chunks and pass them to the mouth. ing technique similar to that of grasshoppers and other
The remipedes capture prey with their rapto- insects). Notostracans, some ostracods, and many
rial mouth appendages (Figures 21.1A, 21.19D–F, decapods, isopods, and amphipods are scavengers and
and 21.22F) and then immobilize the victim with an herbivores. Isopods in several families bore into wood,

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and those in the family Limnoriidae (the gribbles) the kentrogon forms a hollow cuticular structure, the
produce their own cellulase, which they use to digest stylet, which injects a motile, multicellular, vermiform
woody tissues (the presence of blood hemocyanin in stage called the vermigon into the host. The vermigon
the hindgut appears to enhance the digestibility of the is the active infection stage. It has a thin cuticle and
cellulose). Some species of Sphaeromatidae bore into epidermis, several types of cells, and the anlage of an
the aerial roots of mangrove trees, and their activities ovary. It invades the host’s hemocoel by sending out
often result in root breakage followed by new multiple long, branching, hollow rootlets that penetrate most of
root initiation, creating the stiltlike appearance charac- the host’s body and draw nutrients directly from the
teristic of red mangroves (Rhizophora), although at an hemocoel. So profound is the intrusion by the rootlets
overall growth-productivity cost to the trees. A number that the parasite takes over nearly complete control of
of crustaceans are full-time or part-time detritivores; the host’s body, altering its morphology, physiology, and
many scavenge directly on detritus, but others (e.g., behavior. Once the parasite invades the host’s gonads,
cephalocarids) stir up the sediments in order to remove parasitic castration may result (i.e., the gonads of para-
organic particles by suspension feeding. sitized crabs never produce mature gametes). Thus the
A great many crustacean species, and entire lin- host is transformed into a slave that serves the needs of
eages are symbiotic with other organisms. Many are its master. The internal root system, or interna, even-
parasites, but others are commensals or mutualists. tually develops an external reproductive body (the
Some decapod mutualists are simply cases in which externa), where egg production occurs. A male cyprid
the crustacean defends the host against predators and/ settles on the externa, transforms into a minute sexu-
or removes detritus and potential pathogens from the ally mature instar called a trichogon, and moves into the
host’s surface. Among the true crabs (Brachyura), six ovary-filled externa to take up residence, where its sole
families consist exclusively of obligate symbionts, and function is to produce sperm. It takes only one or two
obligate symbionts are also found in at least four addi- trichogons to stimulate the female ovaries to mature and
tional brachyuran families (and facultative symbionts begin releasing eggs into the chamber of the externa. An
occur in numerous other families). For example, the gall externa that fails to obtain male trichogons eventually
crabs (Cryptochiridae) comprise over 50 species of obli- dies. The males are thus parasitic on the female (which
gate symbionts of scleractinian corals. There is a range is itself parasitic on the crustacean host), and kentrogo-
of “gall” formation, from simple pits or chambers to nids are gonochoristic. A mature externa, usually aris-
enclosures formed by the coral as it grows over the crab ing from the host’s abdomen, will produce a succession
(presumably induced by the respiratory currents and of larval broods, molting after each larval release (it is
scratching of the crabs on the coral). Gall crab females the only part of the rhizocephalan body that molts). The
always inhabit pits or chambers and rarely leave; males larvae are lecithotrophic and develop through several
live in separate chambers or free on the coral surface nauplius stages to the cyprid (Figures 21.25 and 21.26).
and must move into the females’ chambers for copula- The akentrogonid rhizocephalans lack a kentrogon
tion. Members of the brachyuran families Domeciidae stage in their life cycle. Instead of injecting a vermi-
and Tetraliidae are also all symbionts of zooxanthellate gon, the female cyprid has long, slender antennules
corals, Trapeziidae are obligate symbionts of corals and that it uses to attach to the abdomen of the host, one of
other colonial cnidarians and sponges, and the subfam- which actually penetrates the host’s cuticle, becomes
ily Eumedoninae (Pilumnidae) are obligate symbionts hollow, and serves for the passage of embryonic cells
of Indo–West Pacific echinoderms. from cypris larva to host. Male cyprids somehow find
Several groups of crustaceans have adopted vari- infected hosts and penetrate them in the same fashion
ous degrees of parasitism. These animals range from as the females, releasing their sperm in such a way that
ectoparasites with mouthparts modified for piercing or they actually enter the body of the female parasite.
tearing and sucking body fluids (e.g., many copepods, The akentrogonid life cycle is similar to that of kent-
branchiurans, tantulocarids, several isopod families, rogonid species, although in some cases more than one
and at least one species of ostracod), to the highly mod- individual parasite might infect a single host, leading to
ified and fully parasitic rhizocephalans, whose bodies multiple externas. And in at least one genus (Thompso-
ramify throughout the host tissue and absorb nutrients nia), multiple externas can develop from a single infec-
directly (Figures 21.25 and 21.26), and pentastomids. tion. The anatomy of the externa is more variable than
Rhizocephalans, which are cirripeds that have been in kentrogonids, and the embryos develop directly into
highly modified to become internal parasites of other cypris larvae—there are no free-swimming nauplius
crustaceans, are some of the most bizarre organisms in stages. The extreme sexual size dimorphism of kent-
the animal kingdom. The most complex rhizocephalan rogonids does not occur in akentrogonid species, and
life cycle is that of the kentrogonid forms, in which a no migrating trichogon stage has been observed. These
settled female cypris larva undergoes an internal reor- two life cycle types used to be the basis for dividing
ganization that rivals that of caterpillar pupae in scope, the Rhizocephala into two orders, Kentrogonida and
developing an infective stage, called the kentrogon, Akentrogonida, but an elegant phylogenetic analysis
beneath the cyprid exoskeleton. Once fully developed, by Høeg et al. (2020) showed these two groups to be

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 712 Chapter 21

(A) (K)
Hermit crab (host)

Courtesy of J. T Høeg
Successive broods Externa

♂
♀
(J)

(B) (C) Immature female

♀ ♂
Trichogon produces

♂
Nauplius larvae sperm in the receptacles

♀
(I)
Male cyprid transforms into
trichogon which invades
receptacles (see Fig. 21.26I)

♀ ♂
(D)

♀
Cypris larvae
(H)
Virginal female
Receptacles empty

♂
♀
Female cyprid settles on host

(E) Parasite penetrates host abdomen

Reproductive body is developed

from root system within host ♀
Female develops into kentrogon
larva and penetrates host

(F) Kentrogon larva

Kentrogon injects (G)

vermigon larva into host
(see Fig. 21.26G)

Root system is developed within host

Vermigon larva
(see Fig. 21.26H)

Brusca 4e
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 ◀ FIGUREfor more The
21.25 ebook/ testbank/life
remarkable solution
cycle ofmanuals requests:
the rhizo­ email
Inside the host, 960126734@[Link]
the vermigon grows with rootlets that rami-
cephalan cirripede Peltogaster paguri, a kentrogonid fy throughout much of the host’s body; it is now called the
parasite of hermit crabs. (A) The mature reproductive interna (G). Eventually the female parasite emerges on the
portion of the parasite (externa) produces numerous abdomen of the host as a virginal externa (H). When the
broods of male and female larvae, which are released as externa acquires a mantle pore, or aperture, it becomes
nauplii (B,C) and eventually metamorphose into cypris lar- attractive to male cyprids (I). Male cyprids settle within the
vae (D). Female cyprids settle on the thorax and limbs of aperture, transform into a trichogon form, and implant part
host crabs (E) and undergo a major internal metamorphosis of their body contents in the female’s receptacles (J). The
into the kentrogon form (F), which is provided with a pair deposit proceeds to differentiate into spermatozoa, which
of antennules and an injection stylet. The kentrogon’s vis- fertilize the eggs of the female. (K) The dissected externa,
cera metamorphoses into an infective stage, the vermigon, with its rootlets, of Peltogaster, removed from its host.
which is transferred to the host through the hollow stylet. Note the mantle aperture.

(A) (B)

From H. Glenner J. T. Høeg. 1995. Nature

377: 147–150. [Link]
66: 1–46. © 1985 The Royal Swedish
From J. T. Høeg. 1985. Acta Zool
Academy of Sciences

articles/377147a0
(C) Epidermis Epidermal
Caudal inclusions
filaments Cement gland
Posterior (of cypris)
mantle cavity
Carapace

Antennule glands

Thoracopods Cerebral
ganglion
50 µm Cuticular reinforcement Anterior mantle cavity
on antennular article 1 Cement gland canal
Antennule article 2 Antennule article 3

(D)

Inclusion-filled
epidermis

Remnant
of anterior
mantle cavity

Cement
FIGURE 21.26 Various stages in the life cycle
of parasitic rhizocephalans. (See Figure 21.25 for
a full description of a rhizocephalan [kentrogonid] life cycle.) Invasion cell Cement gland Gill filament
(A–D) Larvae of Lernaeodiscus porcellanae: (A) A live cyprid. of kentrogon of host
(B) A cyprid (lateral view; SEM). (C,D) Diagrams of cyprid larvae
before and after settlement (right side of carapace removed; naupliar
eye omitted). The dotted line in the second antennular article indicates
the primordial kentrogon cuticle, and the placement of muscle fibers in
the cyprid is indicated by arrows; the muscles are hypothesized to effect
formation of the kentrogon and separation of the old cyprid from the
kentrogon. In D, kentrogon formation is complete. (Continued on next page)

21_Brusca4e_CH21.indd 713 3/8/22 1:09 PM

 714 Chapter 21

(E) (F)
© 1985 The Royal Swedish Academy of Sciences
From J. T. Høeg. 1985. Acta Zool 66: 1–46.

(H)

From H. Glenner J. T. Høeg. 1995. Nature

377: 147–150. [Link]
(G)

Courtesy of J. T. Høeg

articles/377147a0
(I)
Digestive System
The digestive system of crustaceans includes the usual
arthropod foregut, midgut, and hindgut. The foregut
and hindgut are made from ectodermally derived tis-
Courtesy of J. T. Høeg

sue and are thus lined with a cuticle that is continuous

with the exoskeleton and molted with it. The stomo-
deal foregut is modified in different groups but usu-
ally includes a relatively short pharynx-esophagus
region followed by a stomach. The stomach often has
FIGURE 21.26 (continued) Various stages in the life chambers or specialized regions for storage, grinding,
cycle of parasitic rhizocephalans. (E) A whole gill (SEM) and sorting; these structures are best developed in the
of a host4ecrab, Petrolisthes cabrilloi (Anomura), with sev-
Brusca malacostracans (Figure 21.27G). The midgut forms a
eral attached kentrogons (arrows). (F) A 2-hour-old kentro- short or long intestine—the length depending mainly
BB4e_21.[Link]
gon (sagittal section). (G) A kentrogon injecting a vermi- on overall body shape and size—and bears variably
12/21/2021
gon via the stylet. (H) A vermigon. (I) A trichogon.
placed digestive ceca. The ceca are serially arranged
only in the remipedes. In some malacostracans, such as
crabs, the ceca fuse to form a solid glandular mass, best
nonmonophyletic. The kentrogonid mode of host inva- called a digestive gland but sometimes called a mid-
sion is likely the original, whereas the akentrogonid gut gland or hepatopancreas, within which are many
mode has been derived at least twice. branched, blind tubules. The digestive gland in some
Isopods of the family Cymothoidae use modified crustaceans has been shown to store lipids. The hind-
mouthparts to suck the body fluids of their fish hosts. gut is usually short, and the anus is generally borne
They attach to their host’s skin or gills or, in some gen- on the anal somite or telson, or on the last segment of
era, to the tongue. One species, Cymothoa exigua, sucks the abdomen (when the anal somite is reduced or lost).
so much blood out of its host’s tongue (the spotted rose Examples of some crustacean digestive tracts are
snapper, in the Gulf of California) that the tongue com- shown in Figure 21.27. Generally, expansion of the
pletely degenerates. But, remarkably, the fish doesn’t esophagus and foregut creates a suction that draws
die; the isopod remains attached with its clawed legs food into the mouth and esophagus. Foregut con-
to the basal muscles of the missing tongue, and the fish tractions push the food into the midgut, where most
uses the crustacean as a “replacement tongue” to con- breakdown and absorption occur. The foregut is typi-
tinue to feed. This is one of the few known cases of a cally elaborate with folds and ridges and can even be
parasite functionally replacing a host organ it destroys. armed with calcareous ossicles (e.g., many decapods).

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 for more ebook/ testbank/ solution manualsArthropoda
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Shrimps, and Their Kin 715

(A) Ovary
Gut
Heart
Accessory Oviduct
photoreceptor
X-organ
Frontal horn
(B) Nauplius eye Anus
Brain Ventral Gonopore
Second antenna nerve cord Seminal
receptacle
Mouth
First antenna

(C)

(D)

FIGURE 21.27 Internal anatomy of representative crustaceans.

(A) A calanoid copepod. (B) A lepadomorph barnacle. (C) A balanomorph
barnacle. (D) A stomatopod. (A after S. M. Marshall and A. P. Orr. 1955.
The Biology of a Marine Copepod. Springer Verlag, New York, [Link]
[Link]/chapter/10.1007/978-3-662-13138-1_2 [and see credits].). (Continued on next page)
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 716 Chapter 21

Posterior Anterior
(E) aorta aorta
Midgut Cor Antennal
Midgut frontale artery
Abdominal segmental artery cecum
gland Optic artery
Hindgut
Heart Supraesophageal
ganglion

Antennal nerve

Antennal gland
Ventral thoracic artery

Sternal artery

Ventral abdominal Midgut

Ventral nerve cord
artery Oviduct
Cardiac stomach
Pyloric stomach

(F)

(G)

FIGURE 21.27 (continued) Internal anatomy of

representative crustaceans. (E) A crayfish. (F) An amphipod.
(G) The stomach of a crayfish. (H) A brachyuran crab, dorsal
view with carapace removed.

After ingestion, the food material is usually handled chemical digestion. For example, the complex foregut
mechanically by the foregut. This may involve simply of decapods (Figure 21.27G) is divided into an anterior
transporting the food to the midgut or, more com- cardiac stomach and a posterior pyloric stomach. Food
monly, processing the food in various ways prior to is stored in the enlarged portion of the cardiac stomach
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Shrimps, and Their Kin 717

(H)

and then moved a bit at a time to a region containing nutrients, as well as waste products and oxygen (via
a gastric mill, which usually bears heavily sclerotized hemocyanin or hemoglobin). Hemolymph also contrib-
teeth. Special muscles associated with the stomach wall utes to wound healing through the clotting of hemo-
move the teeth, grinding the food into smaller parti- cytes and other cells.
cles. The macerated material then moves into the back The primitive heart structure in crustaceans is a long
part of the pyloric stomach, where sets of filtering setae tube with segmental ostia, a condition retained in part
prevent large particles from entering the midgut. This in cephalocarids and in some branchiopods, leptostra-
type of foregut arrangement is best developed in mac- cans, and stomatopods. However, the general shape of
rophagous decapods (scavengers, predators, and some the heart and the number of ostia are closely related to
herbivores). Thus the food can be taken in quickly, in body form and the location of gas exchange structures.
big bites, and mechanically processed afterward. The heart may be relatively long and extend through
much of the postcephalic region of the body, as it does
in the remipedes, anostracans, and leptostracans, or
Circulation and Gas Exchange it may tend toward a globular or box shape and be
The basic crustacean circulatory system usually con- restricted to the thorax (e.g., as in cladocerans), where
sists of an elongate, dorsal, ostiate heart within a peri- it may be associated with the thoracic gills (e.g., as in
cardial cavity and variously developed vessels emp- decapods). The intimate coevolution of the circula-
tying into an open hemocoel (Figure 21.28). But the tory system with body form and gill placement is best
open hemocoel, which in the past led to descriptions exemplified when closely related groups are compared.
of crustaceans as having an open circulatory system, Although isopods and amphipods, for instance, are
has become highly modified in many groups, and in both peracarids, their hearts are located largely in the
most there is a complex and intricate arrangement of pleon and in the pereon, respectively, corresponding to
true vessels (arteries) and lacunar canals that has only the pleopodal and pereopodal gill locations.
recently begun to be recognized. A distinct heart is The number and length of blood vessels and the
absent in most ostracods, many copepods, and many presence of accessory pumping organs are related to
cirripedes. In some groups the heart is replaced or sup- body size and to the extent of the heart itself. In most
plemented by accessory pumping structures derived nonmalacostracans, for example, there are no arterial
from muscular vessels. But in all cases, the organs and vessels at all; the heart pumps blood directly into the
tissues are exposed to the hemolymph, which fills the hemocoel from both ends. These animals tend to have
body cavities. The circulating hemolymph helps regu- short bodies, long hearts, or both, an arrangement that
late pH and inorganic ions, and it transports amino facilitates circulation of the blood to all body parts. Ses-
acids, proteins, carbohydrates and lipids, and other sile forms, such as most cirripedes, have lost the heart

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 718 Chapter 21

(A) (B) (C) (D)

(E) (F)

(I)

(G) (H)

FIGURE 21.28 Gas exchange structures. (A) Maxilla of of (E) a shrimp and (F) a brachyuran crab. (G) The branchial
the shrimp Pandalus. Note the setose scaphagnathite chamber (cross section) of a brachyuran crab, showing
used to generate the ventilating current. (B–D) Cross sec- the position of a single phyllobranchiate podobranch.
tions of types of decapod gills: (B) Dendrobranchiate. (H,I) A pleopod of the terrestrial isopod Porcellio (surface
(C) Trichobranchiate. (D) Phyllobranchiate. (E,F) Paths of view and section). Note the pseudotracheae. (B–D after
ventilating currents through the left branchial chambers Meglitsch 1972.)

altogether, although it is replaced by a pumping vessel to dissolved hemoglobin. Most malacostracans pos-
in the thoracicans. Large malacostracans tend to have sess hemocyanin in solution (although some contain
well-developed vessel systems (Figure 21.27D,E). Stud- hemoglobin within tissues). Hemoglobin uses iron as
ies using corrosion casting techniques and microCT the oxygen-binding site, whereas hemocyanin uses cop-
scanning have shown how complex these vessel systems per. The latter can give a bluish color to the hemolymph;
can be and have even called into question the paradigm carotenoid pigments frequently give hemolymph a pale
of “open” vs. “closed” circulatory systems in inverte- brown or orange color. Oxygen-binding pigments are
brates. Large or active crustaceans may also possess an never carried in corpuscles, as they are in the vertebrates.
anterior accessory pump called the cor frontale, which We have mentioned the form and position of gas
helps maintain blood pressure, and often a venous sys- exchange organs (gills) for some groups of crustaceans
tem for returning blood to the pericardial chamber. in the taxonomic synopses. Some small forms (e.g., cope-
Crustacean blood contains a variety of cell types, pods, some ostracods) lack distinct gills and rely on cuta-
including phagocytic and granular amebocytes and neous exchange, which is facilitated by their relatively
special wandering explosive cells that release a clotting thin cuticles and high surface-area-to-volume ratio. In
agent at sites of injury or autotomy. In nonmalacostra- the small forms of other groups, a thin, membranous
cans, oxygen is either carried in solution or attached inner lining of the carapace serves this purpose (e.g.,
Brusca 4e
BB4e_21.[Link]
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 for more ebook/ testbank/ solution manualsArthropoda
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Shrimps, and Their Kin 719

Cladocera, Cirripedia, Leptostraca, Cumacea, Mysida, diffusion of respiratory gases commonly continues even
Spinicaudata, and even some members of the Decapoda). during low tides. Some decapods have even invaded
Most crustaceans, however, possess distinct gills land, especially certain crayfish and the anomuran and
of some sort (Figure 21.28). These structures are com- brachyuran crabs known as land crabs (e.g., certain
monly derived from epipods (exites) on the thoracic Gecarcinidae, the hermit crab Coenobita, the coconut
legs that have been modified in various ways to pro- crab Birgus; Figure 21.1H). In these semiterrestrial spe-
vide a large surface area. The inner hollow chambers cies the gills are typically reduced in size. In Birgus the
of these gills are confluent with the hemocoel or its original gills are very small, and the vascularized cuticu-
vessels. Although the gills vary considerably in struc- lar surface of the gill chamber is used for gas exchange.
ture (recall the various decapod gills described earlier), Although young Birgus may carry shells (or coconuts)
they all operate on the basic principles of gas exchange to protect the soft abdomen, adults do not and the abdo-
organs addressed in Chapter 3 and throughout this men is well sclerotized and hardened. Some land crabs
text: the circulatory fluid is brought close to the oxy- in the genus Gecarcinus, which have been reported living
gen source in an organ with a relatively high surface at up to 3,000 m elevation (in Jamaica), have nephritic
area. The gills provide a thin, moist, permeable sur- pads onto which the urine is deposited; microbes living
face between the internal and external environments. on the pads cleanse the urine of its nitrogenous waste
The gills of stomatopods and isopods (Figure 21.28H,I) compounds before the fluid is reabsorbed by the crab.
are formed from the abdominal pleopods. In the first Interestingly, at least three species of Drosophila have
case they are branched processes off the bases of the adapted to this habitat, where they feed on the microbes
pleopods, but in the isopods the flattened pleopods and spend their entire lives in association with their crab
themselves are vascularized and provide the necessary hosts. Another striking decapod adaptation to life in air
surface area for exchange. Stomatopods also have epip- is displayed by the sand-bubbler crabs of the Indo-Pacific
odal gills on the thoracopods, but these are not large. region (family Dotillidae: Scopimera, Dotilla). These crabs
For gills to be efficient, a flow of water must be main- possess membranous discs on their legs or sternites that
tained across them. In stomatopods and aquatic iso- were once thought to be auditory organs but are now
pods a current is generated by the beating of the pleo- thought to function as gas exchange surfaces.
pods. Similarly, the pereopodal gills of euphausiaceans The most successful crustaceans on land are not
are constantly flushed by water as the animals swim. the decapods, however, but the familiar sowbugs
In many crustaceans, however, the gills are concealed and pillbugs. The success of these oniscidean isopods
to various degrees and require special mechanisms in (e.g., Porcellio) is due in part to the presence of aerial
order to produce the ventilating currents. In most deca- gas exchange organs called pseudotrachea (Figure
pods, for example, the gills are contained in branchial 21.28H,I). These organs are inwardly directed, moder-
chambers formed between the carapace and the body ately branched, thin-walled, blind sacs located in some
wall (Figure 21.28). Thus the delicate gills, while still of the pleopodal exopods, connected to the outside via
technically outside the body, appear to be (and are pro- small pores (similar to the tracheae and spiracles of
tected as though they were) internal organs. While such other terrestrial arthropods, such as some arachnids,
an arrangement provides protection from damage to the myriapods, and hexapods). Air circulates through these
fragile gill filaments, the openings to the chambers are sacs, and gases are exchanged with the blood in the
generally small, restricting the passive flow of water. pleopods. Thus, in these animals the original aquatic
Not surprisingly, the solution to this dilemma comes pleopodal gills have been refashioned for air breath-
once again from the evolutionary plasticity of crustacean ing by moving the exchange surfaces inside, where
appendages. Most decapods have elongate exopods on they remain moist. The superficially similar tracheal
the maxillae, called gill bailers or scaphognathites, that systems of isopods, insects, myriapods, and arachnids
vibrate to create ventilating currents through the bran- evolved independently, by convergence, in association
chial chambers (Figure 21.28A). These currents typically with other adaptations to life on land.
enter from the sides and rear through small openings
around the coxae of the pereopods (called in crabs
Milne-Edwards openings, after their discoverer), and Excretion and Osmoregulation
they exit anteriorly from under the carapace in the vicin- Like other fundamentally aquatic invertebrates, crusta-
ity of the mouth field (and antennal glands). They can be ceans are ammonotelic, whether in fresh water or sea-
easily seen by observing a crab or lobster in quiet water. water or on land. They release ammonia both through
The flow rate of the currents can be altered, depending nephridia and by way of the gills. As discussed in
on environmental factors, and can also be reversed, thus Chapter 20, most crustaceans possess nephridial excre-
allowing certain decapods to burrow in sand or mud tory organs in the form of either antennal glands or
with only their front ends exposed to the water. maxillary glands (Figures 21.6A and 21.27). These are
The positioning of the gills in branchial chambers pro- serially homologous structures, constructed similarly
tects them from desiccation during exposure to air and but differing in the position of their associated pores (at
thus enables many crustaceans to live in littoral habitats; the base of the second antennae or the second maxillae,

21_Brusca4e_CH21.indd 719 3/8/22 1:09 PM

 720 Chapter 21

respectively). The inner blind end is a coelomic rem- metabolic wastes, but also are extremely important in
nant of the nephridium called the sacculus, which water and ion balance, particularly in freshwater and
leads through a variably coiled duct to the pore. The terrestrial crustaceans.
duct may bear an enlarged bladder near the opening. The excretion and osmoregulation carried out by
Antennal glands are sometimes called “green glands.” antennal and maxillary gland activity are supplemented
Most crustaceans have only one pair of these nephrid- by other mechanisms. The cuticle itself acts as a barrier
ial organs, but lophogastridans and mysidans have both to exchange between the internal and external environ-
antennal and maxillary glands, and a few others (ceph- ments and is especially important in preventing water
alocarids, and a few tanaidaceans and isopods) have loss on land or excessive uptake of water in fresh water.
well-developed maxillary and rudimentary antennal Moreover, thin areas of the cuticle (especially the gill sur-
glands. Most nonmalacostracans have maxillary glands, faces) serve as sites of waste loss and ionic exchange. The
as do stomatopods, cumaceans, and most tanaidaceans epipods on the legs of Branchiopoda were long assumed
and isopods. Adult ostracods have maxillary glands, but to function in gas exchange (as “gills”), but they are now
antennal glands also occur in freshwater species. All of known to serve primarily as sites of osmoregulation
the other malacostracans have antennal glands. (hence, the taxonomic name Branchiopoda, meaning
Blood-filled channels of the hemocoel intermingle “gill-footed,” is a misnomer!). Phagocytic blood cells
with branched extensions of the sacculus epithelium, and certain regions of the midgut are also thought to
creating a large surface area across which filtration accumulate wastes. In some terrestrial isopods, ammo-
occurs. The cells of the sacculus wall also actively take nia actually diffuses from the body in gaseous form.
up and secrete material from the blood into the lumen
of the excretory organ. These processes of filtration and
secretion are to some degree selective, but most of the Nervous System and Sense Organs
regulation of urine composition is accomplished by The central nervous system of crustaceans is constructed
active exchange between the blood and the excretory in concert with the segmented body structure, along the
tubule. These activities not only regulate the loss of same lines as seen in other arthropods (Figure 21.29). In

(A) (B) (D)

(C)

Pleon ganglia

FIGURE 21.29 Central nervous systems of four brachyuran crab, wherein all thoracic ganglia have fused
crustaceans. (A) The ladderlike system of an anostracan. and the abdominal ganglia are reduced. (D) Nervous sys-
Note the absence of well-developed ganglia in the poste- tem of a hyperiid amphipod. Note the loss of the urosomal
rior, apodous, portion of the trunk. (B) Elongate metameric ganglia typical of all amphipods. (D after G. J. Brusca.
system of a crayfish. (C) Highly compacted system of a 1981. J Crust Biol 1: 358–375.)

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Shrimps, and Their Kin 721

many groups the nervous system is ladderlike, the seg- located on the antennules. In decapods, hundreds of
mental ganglia being largely separate and linked by trans- neurons can innervate each aesthetasc. Thermorecep-
verse commissures and longitudinal connectives (Figure tors probably occur in many crustaceans (those liv-
21.29A). The crustacean brain is composed of three fused ing near hydrothermal vents in the deep sea would
ganglia, or neuromeres, the two anterior being the dorsal seem likely candidates) but are not yet documented.
(supraesophageal) protocerebrum and deutocerebrum, However, behaviors related to thermal avoidance and
which are thought to be preoral in origin. From the pro- temperature preferences have been shown. A pair of
tocerebrum, optic nerves innervate the eyes. Mushroom unique sensory structures whose function is unknown,
bodies have been identified in many crustacean brains. called frontal processes, occurs on the head of remi-
From the deutocerebrum, antennulary nerves run to the pedes. Many crustaceans have dorsal organs, poorly
antennules, while smaller nerves innervate the eyestalk understood glandular-sensory structures on the head,
musculature. The third ganglion of the brain is the trito- which actually constitute several different types of sen-
cerebrum, which presumably represents the first postoral sory structures that may or may not be homologous.
somite ganglion. The tritocerebrum forms a pair of circu- Like all arthropods, crustaceans contain well-­
menteric connectives that extend around the esophagus developed proprioceptors that provide information
to a subesophageal or subenteric ganglion and link the about body and appendage position and movement dur-
brain with the ventral nerve cord bearing the segmen- ing locomotion. A few taxa within the class Malacostraca
tal body ganglia. From the tritocerebrum also arise the possess statocysts, which either are fully closed and con-
antennary nerves as well as certain sensory nerves from tain a secreted statolith (e.g., mysidans, some anthurid
the anterior region of the head. In tantulocarids, bizarre isopods) or are open to the outside through a small pore
minute parasites on other crustaceans, the central ner- and contain a statolith formed of sand grains (e.g., many
vous system has a dorsal brain and ventral nerve cord decapods and some asellote isopods) (Figure 21.30B,C).
that runs through the thorax. The brain is formed of very In the case of the decapods, the statocyst not only serves
small, tightly packed cells, and a neuropil is present (con- as a georeceptor, but also detects the angular and lin-
stituting almost half of the brain volume). The neuropil ear acceleration of the body relative to the surrounding
extends inside the ventral nerve cord. water as well as the movement of water past the animal
The nature of the ventral nerve cord often clearly (i.e., the statolith is rheotactic). And in some shrimps the
reflects the influence of body tagmosis. In crustaceans statocyst is apparently also engaged in hearing.
with relatively homonomous segmentation (e.g., remi- There are two types of rhabdomeric photoreceptors
pedes, cephalocarids, and anostracan and notostracan among crustaceans, median simple eyes and lateral
branchiopods), the ganglia associated with each postan- compound eyes, and both are innervated by the pro-
tennary segment remain separate along the ventral nerve tocerebrum. Many species possess both kinds of eyes,
cord. In more heteronomous forms, however, a single either simultaneously or at different stages of develop-
large subenteric ganglionic mass is formed by the fusion ment. The compound eyes may be sessile or stalked.
of ganglia associated with the postoral cephalic segments Stalked compound eyes occur in the Anostraca, many
(e.g., those of the mandibles, maxillules, maxillae, and, Malacostraca, and perhaps some Cumacea (and in
when present, maxillipeds). The ganglia of the thorax anomalocaridids, and perhaps also some trilobites).
and abdomen may also be variably fused, depending on The median eye generally first appears during the
segment fusion and body compaction. For example, in nauplius larval stage, and for that reason it is often
most long-bodied decapods (lobsters and crayfish), the called a naupliar eye. Like the nauplius larva itself,
thoracic and abdominal ganglia are largely fused across the median eye is thought to be an ancestral (defining)
the body midline but remain separate from one another feature of the Crustacea. Median eyes are in a sense
longitudinally (Figure 21.29B). However, in short-bodied “compound” in that they are composed of more than
decapods (e.g., crabs), all of the thoracic segmental gan- one photoreceptor unit (Figure 21.30D). There are typi-
glia are fused to form a large ventral nerve plate, and the cally three such units in the median eyes of nauplii and
abdominal ganglia are much reduced (Figure 21.29C). up to seven in the eyes of adults in which they persist.
Most crustaceans have a variety of sensory recep- Except for their basic rhabdomeric nature, however,
tors that transmit information to the central nervous the structure of median eye units is unlike that of the
system in spite of the imposition of the exoskeleton (as ommatidia of true compound eyes. The former are
explained in Chapter 20) (Figure 21.30). Among the inverse pigment cups, each with relatively few retinu-
most obvious of these sensory structures are the many lar (photoreceptor) cells. Cuticular lenses are present
innervated setae or sensilla that cover various regions over the median eyes of most ostracods and some
of the body and appendages (Figure 21.31). Studies of copepods. Simple crustacean eyes probably function
these structures suggest most function as both mecha- only to detect light direction and intensity. Such infor-
noreceptors (sensing touch and currents) and chemore- mation is of particular value as a means of orientation
ceptors. Most crustaceans also possess special chemo- in planktonic forms without compound eyes, such as
receptors in the form of clumps or rows of soft, tubular, nauplius larvae, many copepods, etc. In some bran-
cuticular processes called aesthetascs (Figure 21.30A) chiopods, a space above the median eye is connected

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 722 Chapter 21

(A) (C)

(D)

(B)

(E)

FIGURE 21.30 Some crustacean sense organs.

(A) Aesthetascs on the antenna of the lobster Panulirus.
(B) Closed statocysts in the uropodal endopods of a mysidan.
(C) Open statocyst in the antennules of a lobster (cutaway

© Larry Jon Friesen

view). (D) A median simple eye of the copepod Eucalanus
(surface view). (E) Complex compound eyes of the stomato-
pod Hemisquilla californiensis.

to the external environment by a small pore, perhaps light wavelengths ranging from the blue-green region
indicating an invagination of the eye at some point in to the ultraviolet and far-red spectra, at least to 470–570
the distant
Brusca 4e past, although the nature and function of nm). Stomatopods, in particular, have been shown to
the pore is not known.
BB4e_21.[Link] have extremely complex and sensitive eyes with a
The structure
12/21/2021 and function of compound eyes wide range of capabilities. Running through the center
(ommatidia) were reviewed in Chapter 20. In terms of of each stomatopod eye is a distinct band comprising
visual capacity, much more work has been done on the six rows of ommatidia—which in most species contain
eyes of insects than on those of crustaceans, and we are four rows that make up a color vision system with 12
left with a good deal of speculation in terms of what visual pigments—as well as two rows that analyze both
crustaceans actually see. Although most probably lack linearly and circularly polarized light (Figure 21.30E).
the visual acuity of many insects, it has been shown The ommatidia that deal with color have a three-tier
that many crustaceans can discern shapes, patterns, and structure: the top tier contains an ultraviolet-sensitive
movement; color vision has been demonstrated in some pigment, and below this are two tiers with different
species (various species have been shown to respond to pigments sensitive to wavelengths in the human visible

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 for more ebook/ testbank/ solution manualsArthropoda
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Shrimps, and Their Kin 723

(A) (B)

(C) (D)

Micrographs courtesy of B. Felgenhauer

(E) (F)

FIGURE 21.31 Sensilla of selected crustaceans. (A) Serrate seta (mecha-

noreceptor) of the second maxilliped of the anomuran Petrolisthes armatus
(×428). (B) Sockets of serrate setae of the third maxilliped of the cleaning
shrimp Stenopus hispidus (×228). (C) Current receptor from the anterior trunk
limbs of the notostracan Triops. (D) Chemosensory seta from the first pereo-
pod of the freshwater shrimp Atya (×5,700); note the characteristic apical
pore. (E) A dual receptor (mechanoreceptor and chemoreceptor) seta from
the maxilla of the remipede Xibalbanus tulumensis (×4,560). (F) Cross section
of a dual receptor (micrograph and interpretive drawing). Note the micro­
tubules within the dendrites and the dendritic sheath that attaches to the
cuticle of the setal shaft (×17,100).

spectrum. The upper and lower halves of each eye also

have overlapping visual fields, so each eye can act as a
stereoscopic range finder. The 12 different photoreceptor
types each sample a narrow set of wavelengths ranging
from deep UV to far red (300 to 720 nm).
Although both insects and crustaceans have tetra- light has a more restricted angular distribution, a lower
partite ommatidia, there are certain structural differ- intensity, and a narrower range of wavelengths than
ences between the compound eyes of insects and those it does in air. Contrast is also somewhat reduced in
of crustaceans, probably as a result of adaptation to water. All of these factors place a premium on enhanc-
the requirements of aerial and aquatic vision. In water, ing the sensitivity and contrast perception of the eyes
Brusca 4e
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 724 Chapter 21

of aquatic creatures. Mounting the eyes on stalks is one it has also been reported in certain myodocopan ostra-
dramatic way in which many crustaceans increase the cods, hyperiid amphipods, and copepod larvae. Some
amount of information available to the eyes, by increas- copepods (e.g., male Sapphirina metallina) display an irri-
ing the field of view and binocular range. Eyestalks are descence, which is created by a multilayered-membrane
complex structural features with a dozen or so muscles structure in the epidermal cells of the dorsal integument
controlling their movement. (see discussion of bioluminescent ostracods below).
Typical tetrapartite compound eyes are lacking in
the small crustaceans formerly combined as “maxil-
lopodans” (copepods, barnacles, etc.), but various forms Reproduction and Development
of “compound eyes” do occur among the Branchiura, Reproduction We have often mentioned the relation-
Ostracoda (Cypridinacea), and Cirripedia. Eyes in the ships between an animal’s reproductive and develop-
first two taxa most closely resemble those of other crus- mental pattern and its lifestyle and overall survival
taceans in general structure and may be homologous strategy. With the exception of purely vegetative pro-
with them. In the Cirripedia, the median eye and two cesses such as asexual budding, the crustaceans have
lateral eyes are all derived from a single tripartite ocellar managed to exploit virtually every life history scheme
eye of the nauplius larva, which splits into its three com- imaginable. The sexes are usually separate, although
ponents, each forming an adult photoreceptor following hermaphroditism is the rule in remipedes, cephaloca-
metamorphosis of the nauplius into a cypris larva. All rids, most cirripedes, and a few decapods. Sequential
three of these eyes thus appear to be composed of simple hermaphroditism is not uncommon and usually is
ocelli, although the lateral eyes have three photorecep- expressed as protandry (individuals first mature as
tor cells and for this reason are often called “compound males, then later become females), although proto­
eyes.” Rhizocephalan nauplii also have a tripartite nau- gyny occurs in a few species (e.g., the marine isopod
plius eye, which persists into the cypris larval stage. Gnorimosphaeroma oregonense [Sphaeromatidae]). In
Compound eyes are lacking altogether in many crus- addition, parthenogenesis is known in some bran-
tacean taxa (e.g., Copepoda, Mystacocarida, Cephalo- chiopods and certain ostracods. In one species of
carida, Tantulocarida, Pentastomida, Remipedia, and clam shrimp (Eulimnadia texana) a rare type of mixed
some Ostracoda). Members of some other groups pos- mating system exists, called androgonochorism
sess compound eyes only in late larval stages and lose (i.e., androdioecy in plants), in which males coexist
them at metamorphosis (e.g., some cirripedes). Reduc- with hermaphrodites, but there are no true females.
tion or loss of eyes is also common in many deep-sea Androgonochorism is rare, but it is also known in the
species, burrowers, cave dwellers, and parasites. nematode Caenorhabditis elegans, some thoracican bar-
Crustaceans have complex endocrine and neurose- nacles (e.g., Balanus galeatus, Scalpellum scalpellum), and
cretory systems, although our understanding of these several other branchiopod crustaceans.
systems is far from complete. In general, the phenom- The reproductive systems of crustaceans are gener-
ena of molting (see Chapter 20), chromatophore activ- ally quite simple (Figure 21.27). The gonads are derived
ity, and various aspects of reproduction are under hor- from coelomic remnants and lie as paired elongate
monal and neurosecretory control. Interesting recent structures in various regions of the trunk. In many cir-
work indicates that juvenile hormone–like compounds, ripedes, however, the gonads lie in the cephalic region.
long thought to occur only in insects, may also occur In some cases the paired gonads are partially or wholly
in at least some crustaceans. (Juvenile hormones are a fused into a single mass. A pair of gonoducts extends
family of compounds that regulate adult metamorpho- from the gonads to genital pores located on one of the
sis and gametogenesis in insects.) Crustaceans are mas- trunk segments, either on a sternite, on the arthrodial
ters of autotomy, or the dropping of limbs at specific membrane between the sternite and leg protopods, or
breakage planes to escape potentially lethal threats. on the protopods themselves. In many crustaceans the
The process occurs via an intrinsic neuromuscular paired penes are fused into a single median penis (e.g.,
mechanism (i.e., it is a nonreflexive process), and the in tantulocarids, cirripedes, and some isopods). The
break point is capable of rapid sealing. In most crus- female system sometimes includes seminal receptacles.
taceans, loss of a limb stimulates rapid regeneration, The position of the gonopores varies among the classes.
usually accompanied by an initial molt. In snapping The curious phenomenon of intersex—having
shrimps (Alpheidae) and some others with unequal both male and female secondary sexual characteris-
chelae, autotomy of a cheliped commonly results in tics—is widespread among crustaceans. Intersexual
claw reversal during the induced molt, transforming development is often associated with the presence
the larger claw into a small one, and vice versa—a of endoparasites such as bacteria and microsporidi­
remarkable process that involves changes in morphol- ans, and it has also been correlated with the pres-
ogy, underlying muscle types, and external setae. ence of endocrine-disrupting environmental pollut-
Bioluminescence also occurs in several crustacean ants that seem to induce opposing secondary sexual
groups—it is common among pelagic decapods, and characteristics.

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 for more ebook/ testbank/ solution manualsArthropoda
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Shrimps, and Their Kin 725

Most crustaceans copulate, and many have evolved takes place once the male has enticed the female into
courtship behaviors, the most elaborate and well his burrow. Males of some fiddler crab species build
known of which occur among the decapods. Although sand structures at the entrance of their burrow, which
many crustaceans are gregarious (e.g., certain plank- have been shown to attract females.
tonic species, barnacles, many isopods and amphi- Among many crustaceans, the external sexual char-
pods), most decapods live singly except during the acteristics are associated with the actual mating pro-
mating season. More or less permanent, or at least sea- cess. For example, the antennae of male anostracans
sonal, pairing is known among many crustaceans (e.g., and some cladocerans, ostracods, and copepods are
stenopodid shrimps; certain parasitic and commensal modified for grasping the female. Additionally, many
isopods; pinnotherid “pea” crabs, which often live as males bear special sperm transfer structures, in the form
pairs in the mantle cavities of bivalve molluscs or in of either modified appendages or special penes such as
burrows of axiid/lobster shrimps). those of the thoracican barnacles (Figure 21.32E), anos-
Even the parasitic pentastomids copulate (within tracans, and ostracods. Examples of modified append-
the host’s respiratory system) and have internal fertil- ages include the last trunk limbs of copepods and the
ization, relying on a transfer of sperm to the female’s anterior pleopods of most male malacostracans (called
vagina by way of the male’s cirri (penes). Pentastomid gonopods in most malacostracans, or petasma in Den-
early embryos metamorphose into a so-called primary drobranchiata) (Figure 21.32F). Sperm are transferred
larva with two pairs of double-clawed legs and one or either loose in seminal fluid or (in copepods, and many
more piercing stylets (Figure 21.5E). The larvae may be malacostracans) packaged in spermatophores. Motile
autoinfective in the primary host, or they may migrate flagellated sperm occur only in a few groups; in most
to the host’s gut and pass out with the feces. In the the sperm are nonmotile. Crustacean sperm are highly
latter case, an intermediate host is required, which variable in shape, even bizarre in many instances,
may be almost any kind of vertebrate. The larvae bore often being large round or stellate cells that move by
through the gut wall of the intermediate host, where pseudopods or are, seemingly, nonmotile.9 Sperm are
they undergo further development to the infective deposited directly into the oviduct or into a seminal
stage. Once the intermediate host is consumed by a receptacle in or near the female reproductive system.
definitive host (usually a predator), the parasite makes In some crustaceans females can store sperm for long
its way from the new host’s stomach up the esophagus, periods (e.g., several years in the lobster Homarus), thus
or bores through the intestinal wall, eventually settling facilitating multiple broods from a single insemination.
in the respiratory system. The great majority of crustaceans brood their eggs
Mating in nonpaired crustaceans requires mecha- until hatching occurs, and a variety of brooding strat-
nisms that facilitate location and recognition of partners. egies have evolved. Peracarids brood the developing
Among decapods, and perhaps many other crustaceans, embryos in a marsupium, a ventral brood pouch
scattered individuals apparently find one another either formed from inwardly directed plates of the leg coxae
by distance chemoreception (pheromones) or through called oostegites (thermosbaenaceans are an excep-
synchronized migrations associated with lunar periodic- tion among the Peracarida and use the carapace as a
ity, tidal movements, or some other environmental cue. brood chamber). Other crustaceans attach the embryos
Contact sex pheromones are also utilized. In one, unique to endites on the bases of the legs or to the pleopods
lineage of myodocopan ostracods in the Caribbean, (Figure 21.32H), usually using mucus secreted by
males produce complex bioluminescent displays similar specialized glands. In some cladocerans, brooding
to those of fireflies to attract females. Once prospective takes place in a dorsal brood chamber formed by the
mates are near each other, recognition of conspecifics carapace. However, the syncarids, almost all den-
of the opposite sex may involve several mechanisms. drobranchiate shrimps, and most euphausiaceans
Vision is known to be important in the stenopodid shed the zygotes directly into the water. A few others
shrimps (most of which live in pairs), in certain anomu- deposit their fertilized eggs in the environment, usu-
rans (e.g., the family Porcellanidae), and in brachyurans ally attaching them to some object (e.g., branchiurans,
(many grapsids and ocypodids). some ostracods, many stomatopods). These deposited
A good deal of work has been done on fiddler crabs embryos may be abandoned or, as is the case in sto-
(Ocypodidae: Uca), of which over 100 species have matopods, carefully tended by the female. Nonethe-
been described. In these species, males engage in dra- less, parental protection of the embryos until they
matic cheliped waving (of their greatly enlarged chela, hatch as larvae or juveniles is typical in crustaceans.
or major claw, which can account for more than 50% of Thus, crustaceans usually engage in mixed or direct
the male’s total mass—more than the largest rack of a life histories (Table 21.2).
male elk!) to attract females and repel competing males 9
The sperm of freshwater ostracods are the longest in the animal
(Figure 21.32A–D). In addition, males produce sounds kingdom, relative to body size (up to 10× body length). Even though
by stridulation and substratum thumping, which are their sperm are aflagellate, they are filiform and range from several
thought to attract potential mates. Mating generally hundred microns to millimeters in length.

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 3/8/22 1:09 PM
© Larry Jon Friesen
Groping penis

Both courtesy of C. Holliday

Nature Picture Library/
Alamy Stock Photo
(C)
(B)

(H)
(E)

Courtesy of C. McLay
© Ivan Kuzmin/Alamy Stock Photo

(I)

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◀ FIGURE 21.32 Reproduction in Crustacea. (A–D) Mating male brachyuran crab, Metacarcinus magister, showing the
behaviors of the fiddler crab Uca. (A) Two males in ritualized modified pleopods (setose appendages to retain eggs in
combat for the favor of a female, while she watches (B). female; modified as gonopod in male). (G) A copulating pair
(C) A single male waving his enlarged cheliped to attract of Hemigrapsus sexdentatus. (H) Gravid female cancer crab,
a female. (D) A male fiddler crab engaged in claw-waving Metacarcinus gracilis, with clutch of developing eggs/
behavior to attract a female. (E) A balanomorph barnacle, embryos. (I) The planktonic copepod Sapphirina, with egg
with cirri and groping penis extended, impregnating a neigh- sacs. (E after G. Gerlach and C. Hinz. 2012. In C. Brönmark
bor. The advantage of a long penis in sessile animals is made and L.-A. Hansson [Eds.], Chemical Ecology in Aquatic
obvious by this illustration. (F) Ventral views of a female and Systems. Oxford University Press, Oxford, UK.)

Development Although crustaceans are among the As discussed above, crustaceans typically share a
most widespread animals on Earth, we know surpris- characteristic larval stage known as the nauplius larva,
ingly little about their embryogeny, which is extraordi- denoted by the appearance of the first three pairs of
narily diverse. The eggs are centrolecithal, with various appendage-bearing somites (Figure 21.33D).10 In those
amounts of yolk. The amount of yolk greatly influences groups having little yolk in their eggs, the nauplius is
the type of early cleavage and is often related to the generally free-living. In those species with yolky eggs,
time of hatching (see Chapter 4). As far as is known, the nauplius stage is commonly passed through as part
the zygotes of most nonmalacostracans undergo some of a longer period of embryonic development (or a long
form of holoblastic cleavage, as do those of syncarids, brood period), and it is sometimes referred to as an egg
euphausiaceans, penaeids, amphipods, and parasitic nauplius. Free-living nauplii are usually planktotro-
isopods. However, cleavage patterns are extremely phic, and their release corresponds to the depletion of
variable, ranging from equal to unequal and from stored yolk. However, in a few groups of crustaceans
radial-like to unique. Early reports of modified spiral (e.g., euphausiaceans and dendrobranchiate shrimps),
cleavage (see Figure 21.33A) in some crustaceans had the nauplius exhibits lecithotrophy.
once been viewed as evidence of close ties between Crustacean development is either direct (the embryos
the crustaceans and other spiralian groups such as the hatch as juveniles that resemble miniature adults) or
annelids. However, the generality of spiral-like cleav- indirect (embryos hatch as a distinct larval stage) (Figure
age among crustaceans has been called into question. 21.32H). In direct development, the embryos are typi-
Furthermore, evidence from molecular phylogenetics cally brooded. With indirect development, the embryos
indicates arthropods belong to the Ecdysozoa clade are typically released into the water (or in some cases
of Protostomia, not the Spiralia (where annelids are deposited on the substratum). Some cases of indirect
nested). In fact, the cell lineages and germ layer ori- development can be viewed as “mixed development,”
gins in Crustacea are quite different from those of spi- in that embryos are brooded for a brief or prolonged
rally cleaving embryos. For example, in barnacles the period of time and early larval stages are passed within
mesodermal germ layer arises from the 3A, 3B, and 3C the brood. Larvae may pass through several phases,
cells, and the 4d cell contributes to ectoderm (Figure with each phase containing slightly different morpho-
21.33B)—whereas typical spiral cleavage involves a logical steps called stages, before the adult condition is
4d origin of mesoderm. Euphausiaceans were long achieved. Direct development occurs in some cladocer-
thought to have spiral cleavage, but recent work indi- ans and branchiurans, and in all syncarids and peraca-
cates they do not—only the oblique angle of the mito­ rids. Ostracods also are typically viewed as having direct
tic spindles during the transition from the two- to the development. However, some ostracod species hatch
four-cell stage superficially resembles spiral cleavage. with only the first three pairs of appendages present, a
Meroblastic cleavage is the rule among many mala- characteristic feature of nauplii, and thus some workers
costracans. Here again, the exact pattern varies, but it consider these to be naupliar larvae (even though they
generally involves intralecithal nuclear divisions fol- are in a bivalved carapace and add limbs gradually).
lowed by nuclear migration to the periphery of the The larval stages that have been recognized in the differ-
embryo and subsequent partitioning of the nuclei into ent crustacean groups have been assigned a plethora of
a cell layer around a central yolky mass (Figure 21.33C). names, and the homologies among these forms are not
The form of the blastula and the method of gastrula- always understood. The more commonly encountered
tion are dependent primarily on the preceding cleavage developmental forms are summarized below (also see
pattern and hence ultimately on the amount of yolk. Table 21.2 and Figure 21.33), but we do not attempt to
Holoblastic cleavage may lead to a coeloblastula that describe them all.
undergoes invagination (as in syncarids) or ingression
(as in many copepods and some cladocerans and anos- 10
It was not until J. V. Thompson discovered the nauplius larvae of
tracans). Other crustaceans form a stereoblastula fol- barnacles in the nineteenth century that this group was finally classi-
fied as Crustacea, a discovery that also marked the first use of larval
lowed by epibolic gastrulation (e.g., cirripedes). Most features in understanding the phylogeny of marine invertebrates. An
cases of meroblastic cleavage result in a periblastula and excellent atlas of crustacean larvae (Martin et al. 2014) includes his-
the subsequent formation of germinal centers. torical notes on larval development in all major crustacean groups.

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 728 Chapter 21

(A) (B) (C)

(F)

(D)

(G)

(H)

(E)

FIGURE 21.33 Cleavage and posthatching stages among crustaceans.

(A) Modified holoblastic cleavage has produced a 28-cell embryo of the cir-
ripede Tetraclita. The cells are labeled as in Wilson’s coding system. (B) A fate
map of a cirripede blastula (right-side view). (C) Intralecithal nuclear divisions
in the early cleavage of a mysidan. (D) Newly hatched copepod nauplius larva.
© Larry Jon Friesen

(E) Nauplius of a copepod. (F) Settling and metamorphosis of a cypris larva of a

lepadomorph barnacle. (G) The zoea (“mysis”) stage larva of the dendrobranchi-
ate shrimp Penaeus. (H) Zoea larva of the brachyuran crab Callinectes sapidus.
(I) Zoea larva of a porcelain crab. (J) Megalopa larva of the xanthid crab Menippe
adina. (K) The characteristic antizoea larva of a stomatopod. (L) The translucent,
paper-thin phyllosoma larva of the lobster Jasus. (M) Cryptoniscus stage (not a
true larva) of the epicaridean isopod Probopyrus bithynis. (N) Cyprid larva of a
barnacle. (M after W. E. Dale and G. Anderson. 1982. J Crust Biol 2: 392–409.)

Crustacean development is sometimes described terms “indirect” and “direct” may be preferable, and
as being epimorphic, anamorphic, or metamorphic, less ambiguous, until we have a better understanding
a nomenclature more often used in myriapods (see of these phenomena.
Chapter 23). However, we caution readers that a clear Anamorphic development is indirect, with larval
evolutionary and functional understanding of crusta- stages, and the number of segments increases with
cean developmental stages is still lacking, and thus the each molt. Epimorphic development is direct, and in
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(I)
(L)

(J) (K)

(M)

(N)
© Larry Jon Friesen

only slightly from the one preceding it.11 Regardless of

name, the zoea is characterized by the presence of nata-
tory exopods on some or all of the thoracic appendages
and by the pleopods being absent (or rudimentary).
Use of the term “postlarva” is unfortunate, as these
crustaceans it is thought to result from a delay in the stages differ dramatically from the preceding larvae as
hatching of the embryo, which causes the nauplius well as from the adults; they represent unique transi-
(and any other possible larval stages) to be suppressed tional stages (both morphologically and ecologically).
or absent. In epimorphic development the hatchlings Examples in the Decapoda include the megalopa of
have a full complement of segments and appendages. true crabs and the puerulus larva of spiny lobsters. In
Metamorphic development is the type of extreme this phase of development, the role of swimming has
development seen among the Eucarida; it includes switched from the thoracopodal limbs to the append-
dramatic transitions in body form from one life his- ages of the abdomen.
tory stage to another. In general, up to five distinct pre- Anamorphic development is a less extreme type of
adult, or larval, phases may be recognized among crus- indirect development in which the embryo hatches as a
taceans: nauplius, metanauplius, protozoea, zoea, and nauplius (or later) larva, but the adult form is achieved
“postlarva.” The zoeal phase shows the greatest diver- through a series of gradual changes in body morphol-
sity in form among the various taxa, and especially in ogy as new segments and appendages are added. It is
decapods it has been given different names in differ- similar in many ways to hemimetabolous development
ent groups (e.g., acanthosoma, antizoea, mysis, phyl- 11
The zoea larvae of palinurid lobsters (phyllosoma larvae) are large,
losoma, pseudozoea). It is common for the zoeal phase bizarre-appearing creatures (Figure 21.33L), which can occur in such
to contain
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in insects in that the postnaupliar stages gradually take as thoracican barnacles, and by groups whose resources
on the adult form with succeeding molts; the classic may not permit production of huge quantities of yolk,
example of anamorphic development is often said to be such as the copepods. At the other end of this adaptive
the Anostraca. But it differs from insect development in spectrum is the direct development of peracarids—a
that insects are born with the full complement of seg- major factor allowing the invasion of land by certain iso-
ments and appendages. Cephalocarida, Remipedia, pod lineages. Between these extremes we see all degrees
many Branchiopoda, and Mystacocarida are anamor- of mixed life histories, with larvae being released at vari-
phic—the nauplius larva grows by a series of molts that ous stages following brooding and care. Second, because
add new segments and appendages gradually as the developmental stages also evolve, an analysis of devel-
adult morphology appears. In many groups hatching opmental sequences can sometimes provide information
is somewhat delayed, and the emergent nauplius larva about the radiation of the principal crustacean lineages.
is termed a metanauplius. The basic nauplius possesses For example, the evolution of oostegites and of direct
only three body somites, while the metanauplius has development combine as a unique synapomorphy of
a few more; however, both possess only three pairs of the Peracarida. Similarly, the addition of a unique larval
similar-appearing appendages (which become the adult form, such as the cypris larva that follows the nauplius
antennules, antennae, and mandibles). The end of the in the cirripedes, can be viewed as a unique specializa-
naupliar/metanaupliar stage is defined by the appear- tion that demarcates that group (Cirripedia). The cyprid
ance of the fourth pair of functional limbs, the maxil- either hatches as the only free-living larva or is the final
lules. In copepods a postnaupliar stage called a cope­ larval stage after a series of lecithotrophic or planktotro-
podite (simply a small juvenile) is often recognized. phic nauplius larval stages.
The most extreme forms of metamorphic, or mixed, It should also be noted that the branchiopods and
development occur in the malacostracan superorder some freshwater ostracods have evolved specialized
Eucarida. The most complex developmental sequences ways of coping with the harsh conditions of many fresh-
are seen among the dendrobranchiate shrimps, which water environments. Parthenogenesis, for example, is
hatch as a typical nauplius larva that eventually under- common in freshwater ostracods. Other adaptations
goes a metamorphic molt to become a protozoea larva, include production of special overwintering forms, usu-
with sessile compound eyes and a full complement of ally eggs or zygotes that can survive extreme cold, lack
head appendages. The protozoea, after several molts, of water, or anoxic conditions. Perhaps most remarkable
becomes a zoea larva, with stalked eyes and three pairs in this respect are the large-bodied branchiopods (fairy
of thoracopods (as maxillipeds). The zoea eventually shrimps, tadpole shrimps, and clam shrimps) whose
yields a juvenile stage (the “postlarva,” a better term encysted embryos are capable of an extreme state of
for which is “decapodid”) that resembles a miniature anaerobic quiescence, or diapause. During these resis-
adult but is not sexually mature. In some other eucarid tant stages, the metabolic rate of the embryos may drop
groups (Caridea and Brachyura) the postlarva is called to less than 10% of their normal rate. Interestingly, there
a megalopa, and in the Anomura it is often called a is a large group of Ediacaran microfossils, called acri-
glaucothoe; in both cases there are setose natatory tarchs, that look remarkably similar to branchiopod
pleopods on some or all of the abdominal somites. In resting stages, although further research is needed to
other eucarids, some (or all) of these stages are absent. confirm the identity of these fossils.
Various other terms have been coined for different Many crustaceans have indeterminate growth,
(or similar) developmental stages. For example, the that is, they continue to molt throughout their life. In
modified zoeal stages of some stomatopods are called contrast, other species have determinate growth and
antizoea and pseudozoea larvae, and the advanced cease molting following puberty (this life history stage
zoeal stage of many other malacostracans is often is sometimes referred to as the terminal molt, or ter-
called a mysis larva. In euphausiaceans, the nauplius minal anecdysis). In some species, the terminal molt is
is followed by two stages, the calyptopis and the fur- sex specific; for example, in American blue crabs (Cal-
cilia, which roughly correspond to protozoea and zoea linectes sapidus) only females have a terminal molt.
stages, before the juvenile morphology is attained.
We’ve only just skimmed the surface of postembry-
onic larval, or developmental stage names; some 140
Crustacean Phylogeny
different larval names have been used for decapods Countless phylogenetic studies have been published on
alone! But, from this wealth of terms and diversity of the topic of crustacean evolution, but recent large-scale
developmental sequences, we can draw two important phylogenomic studies of the Pancrustacea have largely
generalizations concerning the biology and evolution agreed on the higher-level structure of the group’s phy-
of the crustaceans. First, different developmental strate- logeny. Thus our tree (Figure 21.34A) depicts Pancrus-
gies reflect adaptations to different lifestyles. In spite of tacea comprising two large sister groups, Oligostraca
many exceptions, we can cite the early release of disper- and Altocrustacea. Oligostraca houses the Ostracoda,
sal larvae by groups with limited adult mobility, such Mystacocarida, Branchiura, and Pentastomida (though

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FIGURE 21.34 (A) Phylogeny of the Pancrustacea

Crustacean phylogeny.
(A) A current view of relation- Chelicerata
ships among the Pancrustacea. Myriapoda
(B) A phylo­geny of the class
Malacostraca. Oligostraca
(Ostracoda, Mystacocarida,
Branchiura, Pentastomida)
Mandibulata
Thecostraca

Multicrustacea
(Barnacles and their kin)
Pancrustacea Copepoda

Altocrustacea
Malacostraca

Cephalocarida

Allotriocarida
Branchiopoda

Labiocarida
Remipedia

Hexapoda

(B) Phylogeny of the class Malacostraca

Leptostraca

Hoplocarida (Stomatopods)

Mysida

Peracarida

Anaspidacea

Euphaysiacea

Dendrobranchiata

Eumalacostraca
(Penaid and sergestid shrimp)
the last has not yet been included in a phy- Stenopodidea
logenomic analysis). The clade Oligostraca
is a group of short-bodied crustaceans Caridea
initially suggested by Zrzavý et al. (1998)

Decapoda
Achelata

Pleocyemata
based primarily on gene sequence data. The remain-
der of the Crustacea (Altocrustacea) houses two sis- Axiidea
Reptantia

ter groups, one sometimes called the Allotriocarida Brachyura

(the “leaf-footed” crustacean lineages: Cephalocarida,
Branchiopoda, and Remipedia, all of which have grad- Anomura
ual, or anamorphic, development, plus Hexapoda), the Astacidea
other sometimes called Multicrustacea (Copepoda, (Clawed lobsters)
Thecostraca, Malacostraca). Analyses support the
monophyly of most traditionally recognized taxa, such
as Malacostraca, Peracarida, Dendrobranchiata, Decap- in hexapods). The former groups known as “Maxil-
oda, Pleocyemata, Reptantia, Thecostraca, Copepoda, lopoda” and “Entomostraca” are not supported by
Branchiopoda, Cephalocarida, Remipedia, and Hexa- molecular analyses.
poda. Relationships within the Branchiopoda reflect Within the class Malacostraca there are two clades:
current thinking based largely on morphology. Rela- Leptostraca, which have phyllopodous limbs and
tionships among the major decapod clades are not yet seven abdominal somites, and Eumalacostraca, which
fully resolved, and the precise position of Copepoda lack phyllopodous limbs, have six abdominal seg-
(within the Multicrustacea) is still uncertain. Hexapoda ments, and have the sixth abdominal appendages mod-
is likely a sister group to the Remipedia, the two taxa ified as uropods (which can work in conjunction with
together called Labiocarida (based on the likely homol- the telson as a tail fan). Relationships among the main
ogy of the “lower lip” in remipedes and the “labium” eumalacostracan lines are shown in Figure 21.34B.
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The class Branchiopoda is monophyletic in molecular upon, based on a wide array of evidence that includes
analyses, but it is difficult to define on the basis of unique molecular sequence data, neuroanatomy, and other
synapomorphies because it shows such great morpho- information. This realization renders the group called
logical variation. The extreme reduction of adult maxil- “Crustacea” paraphyletic. The monophyletic clade con-
lae, as well as certain naupliar characters (e.g., reduced taining both crustaceans and insects is referred to as the
and tubular antennules, uniramous mandibles) strongly Pancrustacea, or Tetraconata (a name that recognizes
support monophyly, as do almost all studies employing the unique presence and anatomy of four crystalline
molecular data. Apparently some branchiopods have cone cells, known as Semper cells, in the ommatidia).
secondarily lost the carapace, and others have second- Like the arthropods in general, crustaceans exhibit
arily lost most or all of the abdominal appendages. high levels of evolutionary parallelism and conver-
That a group of crustaceans gave rise to the mega- gence and many apparent reversals of character states.
diverse group of arthropods called the Hexapoda This genetic flexibility is no doubt due in part to the
(insects and their kin) is now almost universally agreed nature of the segmented body, the serially homolo-
gous appendages, and the flexibility of developmental
genes, which, as we have stressed, provide enormous
(A)
opportunity for evolutionary experimentation. Any
conceivable cladogram of crustacean phylogeny will
require the acceptance of considerable homoplasy.
Fossil data (including the Cambrian Orsten fauna,
Figure 21.35) seem to favor phyllopodous limbs as the
primitive condition. However, developmental stud-
ies following the expression of Distal-less and other

(B)

Courtesy of D. Waloszek

(C)
Courtesy of D. Waloszek

(D)
100 µm
Courtesy of D. Waloszek

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Shrimps, and Their Kin 733

developmental genes suggest that the early embryogeny for example, is Skara (Figure 21.35C), a cephalocarid-
of limbs is very similar among crustaceans. For example, or mystacocarid-like crustacean for which both nau-
trunk limbs always emerge as ventral, subdivided limb pliar larvae and adults have been recovered (the nau-
buds. In phyllopodous limbs, the subdivisions of these plius larvae are only a couple hundred microns long;
limb buds grow to become the endites and the endopod adults are about 1 mm in length). Skara and many other
of the natatory/filtratory adult limbs. In stenopodous Orsten crustaceans were probably meiofaunal animals
limbs, the same limb bud subdivisions end up develop- not unlike modern marine meiofaunal forms (e.g., the
ing into the actual segments of the adult limb. Hence, the mystacocarids). Dozens of Orsten microcrustaceans
endites of phyllopodous limbs appear to be homologous have so far been described (Figure 21.35). In 2001, a
to the segments of the stenopodous limbs. This discov- beautifully preserved fossil, Ercaia minuscula, was
ery supports an emerging view of developmental plas- described from the middle Cambrian (520 Ma; Series 2)
ticity in arthropod limbs, and it suggests that relatively of South China. It has an untagmatized, 13-segmented
simple genetic “switches” can account for major differ- trunk with serially repeated biramous appendages
ences in adult morphologies. Thus, it is quite plausible and a head with stalked eyes and five pairs of head
that stenopodous limbs have evolved multiple times appendages, including two pairs of antennae. Ercaia
from phyllopodous ancestors, and this is the scenario minuscula is only 2–4 mm long (hence the species
depicted in the tree in Figure 21.34B. name) and bears resemblances to both cephalocarids
Work by Klaus Müller and Dieter Waloszek on and “maxillopodans” (Figure 21.35F).
three-dimensionally preserved microscopic arthro- Studies on Cambrian fossils from the Swedish
pods from the middle Cambrian Orsten (about 510 Ma) Orsten fauna (510 Ma), Burgess Shale–like deposits
deposits of Sweden has documented a diverse fauna from around the world (520 Ma), and the Chengjiang
of minute crustaceans and their larvae. Among them, fossils from China (530 Ma) have shown that Cambrian

(E)

Both courtesy of D. Waloszek

30 µm

(F) FIGURE 21.35 (A–E) Examples of upper Cambrian (~510 Ma),

probably meiofaunal crustacean fossils from the spectacular Swedish
Orsten deposits. This ancient crustacean fauna possessed the key
attributes of modern Crustacea, including compound eyes, head
shields/carapaces, naupliar larvae (with locomotory first antennae),
and biramous appendages on the second and third head segments
(the second antennae and mandibles). (A) Bredocaris. (B) Rehbachiella,
an early branchiopod, lateral (drawing) and ventral (SEM) views.
(C) Skara, drawing and SEM. (D) Cambropachycope clarksoni, a bizarre
species with an expanded head and two pairs of enlarged thoracopods,
drawing and SEM. (E) Martinssonia elongata, SEMs of first larva and
postlarval stage. (F) Ercaia minuscula, an early Cambrian (520 Ma)
crustacean from South China.

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 734 Chapter 21

crustaceans had all the attributes of modern ones, fossils from those strata are viewed as Crustacea. The
such as compound eyes, an ocular segment, distinct earliest undisputed crustacean larval stage is a fossil
head and trunk tagmata, at least four head append- of a slightly advanced nauplius (called a metanau-
ages, a carapace (or head shield), naupliar (or “head”) plius) from the Terreneuvian (525 Ma) of China that in
larvae (with locomotory first antennae), and biramous many ways resembles the naupliar larva of modern cir-
appendages on the second and third head somites ripedes. This is remarkable, as it confirms not only the
(the second antennae and mandibles). We thus know great age of the Crustacea as a whole but also the age
that the crustaceans are an ancient group. Their fossil of groups within the Crustacea, such as the cirripedes,
record dates back to the early Cambrian (the Fortunian that were already becoming distinct.
stage), or likely the Ediacaran period if some arthropod

Chapter Summary
The subphylum Crustacea is a paraphyletic group, active predation, and parasitism. The body is a hemo-
because it excludes the Hexapoda (insects and their kin), coel, but a dorsal ostiate (pumping) heart is usually
which evolved from within Crustacea close to 500 mil- present, as are small vessels that distribute the hemo-
lion years ago. The sister group of Hexapoda may be coel. Excretion relies on various nephridially-derived
Remipedia. Phylogenetically, Crustacea and Hexa­poda structures and diffusion across the gills. The central
comprise the clade Pancrustacea (sometimes called nervous system follows the general arthropod plan,
Te­traconata). As with the insects on land, crustaceans with a brain composed of three fused pairs of ganglia
have been hugely successful in the aquatic world. Close (protocerebrum, deutocerebrum, tritocerebrum), the
to 60,000 living crustacean species have been described, tritocerebrum forming a pair of circumenteric connec-
but five or ten times that number probably await discov- tives around the esophagus to a subenteric ganglion
ery. As with insects, the morphological disparity within that links to the ventral nerve cord formed of paired
Crustacea is staggering, with body sizes ranging from segmental ganglia. Most crustaceans are gonochoristic,
less than 100 µm in length to over 4 meters in width. although hermaphroditism is not uncommon (usually
Given that most of Earth is covered with water, and the expressed as protandry or protogyny). Parthenogenesis
great depths of world’s seas, crustaceans are the most is known in some groups. Cleavage patterns in embryos
widespread animals on the planet! Crustaceans are well are highly diverse and largely dependent on the amount
understood, in part, because of their economic impor- of yolk present, ranging from equal to unequal. Crusta-
tance, especially as seafood. A number of groups, nota- cea have a characteristic first larval stage called the nau-
bly isopods and amphipods, have successfully invaded plius larva (suppressed or bypassed in some groups),
the terrestrial world. The crustacean body is divided denoted by three pairs of appendage-bearing somites.
into a head (bearing 5 pairs of appendages) and trunk, In some groups, various other postnaupliar larvae occur
the latter being subdivided into a thorax and abdomen (e.g., metanauplius, protozoea, zoea, megalopa). A rich
in all but the class Remipedia. The head, and often some record of Cambrian fossil Crustacea has been described
of the thoracic segments, are commonly covered by a from Sweden, China, and elsewhere. We now know
dorsal shield called the carapace. Paralleling their mor- that Pancrustacea likely comprises three great clades:
phological diversity, crustaceans exhibit a wide range Oligostraca (Ostracoda, Mystacocarida, Branchiura,
of locomotory strategies that include swimming, walk- Pentastomida), Multicrustacea (Thecostraca, Copepoda,
ing, and hopping. They also have numerous modes of Malacostraca), and Allotriocarida (Cephalocarida, Bran-
feeding, including suspension feeding, deposit feeding, chiopoda, Remipedia, Hexapoda).

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