CH 21 Crustacea PDF

rustaceansare one of the most popular invertebrate groups, even
amongnonbiologists,for they include someof the world's most delec-

table gourmet fare,such as lobsters,crabs,and shrimps (Figure 21.1).
Thereare an estimated 70,000describedliving speciesof Crustacea,and
probably five or ten times that number waiting to be discovered and named.
They exhibit an incredible diversity of form, habit, and size. The smallest known
crustaceans are less than 100 pm in length and live on the antennules of cope-
pods. The largest are Japanesespider crabs (Macrocheiraknempferi),withleg spans
of 4m, and giant Tasmanian crabs (Pseudocarcinusgigas)with carapace widths
of 46 cm. The heaviest crustaceans are probablv American lobsters (Homarus
americanus),which, before the present era of overfishing, attained weights in
excess of 20 kilograms. The world's largest land
arthropod by weight (and possibly the largest
land invertebrate) is the coconut crab (Birgus latro),
Classification of The Animal weighing in at up to 4 kg. Crustaceans are found
Kingdom (Metazoa) at all depths in every marine, brackislu and fresh-
water envfuonment on Earth, including in pools at
Non-Bilateria* Lophophorata
(a.k.a. the diploblasts)
6,000m elevation (fairy shrimp and cladocerans in
PHYLUM PHORONIDA
PHYLUM PORIFERA PHYLUM BRYOZOA
northem Chile). Afew have become successful on
PHYLUM PLACOZOA PHYLUM BRACHIOPODA. land, the mostnotablebeing sowbugs and pillbugs
PHYLUM CNIDARIA (the terrestrial isopods). I
EcpYsozoA
PHYLUM CTENOPHORA Nematoida Crustaceans are commonly the dominant or-
PHYLUM NEMATODA ganisms in aquatic subterranean ecosystems, and
Bilateria PHYLUM NEMATOMORPHA
(a.k.a.the triploblasts) new species of these stygobionts continue to be
Scalidophora
PHYLUM XENACOELOMORPHA discovered as new caves are explored. They also
PHYLUM KINORHYNCHA
Protostomia PHYLUM PRIAPULA dominate ephemeral pool habitats, where many
PHYLUM CHAETOGNATHA PHYLUM LORICIFERA undescribed species are known to occur.2 And
SprRmn Panarthropoda crustaceans are the mostwidespread, diverse, and
PHYLUM PLATYHELMINTHES
PHYLUM TARDIGRADA
PHYLUM GASTROTRICHA
PHYLUM ONYCHOPHORA
PHYLUM RHOMBOZOA
This chapter has been revised by Richard C. Bruscaand Joel
PHYLUM ARTHROPODA W. Martin
PHYLUM OBTHONECTIDA
SUBPHYLUMcRusrAcEA" lWhen most people hear the word "shrimp" they think of
PHYLUM NEMERTEA
SUBPHYLUM HEXAPODA edible shrimps, two crustaceangroups nestedwithin the
PHYLUM MOLLUSCA SUBPHYLUM MYRIAPODA order Decar:oda(in the subordersDendrobranchiataand
PHYLUM ANNELIDA
SUBPHYLUM CHELICEFATA Pleocyemati).However, the term "shrimp" is applied to a
PHYLUM ENTOPROCTA number of long-tailed crustaceans,many not closely related
Deuterostomia
PHYLUM CYCLIOPHORA at all to decapods.So,in this generalsense,there are fairy
PHYLUM ECHINODERMATA
Gnathifera PHYLUM HEMICHORDATA
shrimps, tadpole shrimps, mantis shrimps, etc. In much of
PHYLUM GNATHOSTOMULIDA the English-speakingworld, the word "prawn" is used for
PHYLUM CHORDATA
PHYLUM MICROGNATHOZOA the edible shrimps, thus eliminating some of the confusion.
*Paraphyletic group 2Onestudy of ephemeralpools in northern California dis-
PHYLUM ROTIFERA
covered30 nrobable undescribedand unnamed crustacean
species(King et al.7996).
762 ChapterTwenty-One
(A)
PHYLUMARTHROPODACrustacea:Crabs,Shrimps,and Their Kin 763
(o)
Figure 21 .1 Diversity among the Crustacea. (A-D)

Classes Remipedia, Cephalocarida, and Branchiopoda.
(A) Spe/eonectes ondinae; note the remarkably homono- vivesi (Decapoda: Axiidea). (M) The Hawaiian regal lobster,
mous body of this swimming crustacean (Remipedia). Enoplometopus (Decapoda: Achelata). (N,O) Two unusual
(B) A cephalocarid. (C) A tadpole shrimp (Branchiopoda; amphipods (Peracarida: Amphipoda). (N) Cysfisoma, a
Notostraca) from an ephemeral pool. (D) A clam shrimp huge (some exceed 10 cm), transparent, pelagic hyperiid
(Branchiopoda: Diplostraca), carrying eggs. (E-Q) Class amphipod. (O) Cyamus scammoni, a parasitic caprellid
Malacostraca. (E) The fiddler crab, Uca princeps amphipod that lives on the skin of gray whales. (P) A male
(Decapoda: Brachyura). (D The giant Caribbean hermit and female cumacean; note the eggs in the marsupium
crab, Petrochirus diogenes (Decapoda: Anomura). (G) A of the female (Peracarida: Cumacea). (Q) Ligia pacifica
hermit crab (Paragiopagurus fasciatus) removed from its (Peracarida: lsopoda), the rock louse; Lrgra are inhabitants
snail shell (Decapoda: Anomura). (H) The coconut crab, of the high spray zone on rocky shores worldwide. (R) The
Birgus latro (Decapoda: Anomura), climbing a tree. mysid Slriel/a sp. (Peracarida: Mysida) with a parasitic epi-
(l) Euceramus praelongus, the New England olive pit caridean isopod (Peracarida: lsopoda: Dajidae) attached to
porcelain crab (Decapoda: Anomura). (J) The pelagic lob- its carapace (Western Australia). (S,T) Class Thecostraca.
sterette, Pleuroncodes planipes (Decapoda: Anomura). (S) Acorn barnacles, Semibalanus balanoides (Cirripedia:
(K) The cleaner shrimp, Lysmata californica (Decapoda: Thoracica). (l-) Lepas anatifera (Cirripedia: Thoracica), a
Caridea). (L) The unusual rock-boring lobster shrimp, Axrus pelagic barnacle hanging from a floating timber.
(Continued on next page)
Figure 21 .1 (continued) Diversity among the Crustacea.

(U) Class Copepoda; the calanoid copepod Gaussra,
a common planktonic genus. (V) Class Tantulocarida,
Deoterthron, parasitic on other crustaceans. abundant animals inhabiting the world's oceans.The
biomassof one species,the Antarctic krill (Euphausia
superba), has been estimated at 500million tons at any
given time, probably surpassingthe biomass of any
other group of marine animals (and rivaling that of the
BOX 21A Characteristics world's ants).The range of morphological diversity
of the among crustaceansfar exceedsthat of even the insects.
SubphylumCrustacea Many speciesof Crustaceaare threatenedby environ-
1. Body composed of a 6-segmented head, or cepha- mental degradation,over 500 are listed on the IUCN
lon (plus the acron), and a long postcephalic trunk; Red List, and about two dozen are protected by the
trunk divided into two more or less distinct tagmata U.S.Environmental ProtectionAgency. Crustaceans
(e,9., thorax and abdomen) in all but the remipedes
are also the most common invasive invertebrates/and
and ostracods (Figure 21 .2)
well over 100specieshave establishedthemselvesin
2. Cephalon composed of (anteriorto posterior):pre-
marine and estuarinewaters of North America alone.
segmental (indistinguishable)acron, protocerebral
segment (lacking appendages), antennularseg-
Becauseof their taxonomic diversity and numerical
ment, antennal segment, mandibular somite, maxil- abundance,it is often said that crustaceansare the "in-
lulary somite, and maxillarysomite; one or more sectsof the sea."We prefer to think of insectsas "crus-
anterior thoracomeres may fuse with the head in taceansof the land." And indeed, there is now strong
members of some classes (e.9., Remipedia, and phylogenetic evidence that the insectsarose from a
Malacostraca),their appendages forming maxillipeds branch within the Crustacea.
3. Cephalic shield or carapace present (highlyreduced Despitethe enormousmorphological diversity seen
in anostracans, amphipods, and isopods) among crustaceans(Figures21.1" to 21..20),
they display
4. Appendages multiarticulate,uniramous or biramous a suite of fundamental unifying features (Box 21A). In
5. Mandibles usually multiarticulatelimbs that function an effort to introduce both the diversity and the unity
as biting, piercing, or chewing/grinding jaws of this enormousgroup of arthropods,we first present
6. Gas exchange by aqueous diffusion across special- a classificationand synopsesof the major taxa.We then
ized branchial surfaces, either gill-likestructures or discussthe biology of the group as a whole, drawing
specialized regions of the body surface examplesfrom its various members.As you read this
7. Excretion by true nephridial structures (e.9., anten- chapter,we ask that you keep in mind the general ac-
nal glands, maxillaryglands) count of the arthropodspresentedin Chapter20.
B. Both simple ocelli and compound eyes occur in
most taxa (not Remipedia),at least at some stage
of the life cycle; compound eyes often elevated on
stalks Classificationof The Crustacea
9. Gut with digestive ceca
Crustaceans have been known to humans since ancient
10. With nauplius larva (unknown from any other arthro- times and have provided us with sources of both food
pod subphylum); development mixed or direcl
and legend. It is somewhat comforting to carcinolo-
gists (those who study crustaceans) to note that Can-
PHYLUM ARTHROPODA Crustacea:Crabs,Shrimps,and Their Kin 765
(A)
Tail
l--cephal
Carapace
Abdomen---l
dllFt|F"L l t-,l
fanI
Propodus Swimmerets
(= pleopods)
Dactyl
Figure 21 .2 Crustacean external mor-
phology: A crayfish (Malacostraca,
Astacidea). (A) External form. Note the
fully developed carapace covering the
Compound cephalon and thorax. (B) The typical mala-
Antennule eye costracan tail fan (ventral view). Note the.
position of the anus on the telson.
Legs
(= pereopods)
cer, one of the two invertebrates represented in the

zodiac, is a crab (the other, of course, is Scorpio-an-
other arthropod). Our modem view of Crustacea as a
taxon can be traced to Lamarck's scheme in the early
nineteenth century. He recognized most crustaceans
as such, but placed the barnacles and a few others in
separate groups. For many years barnacles were clas-
SUBORDER ONYCHOCAUDATACIam shrimps,
sified with molluscs because of their thick, calcareous
cladocerans(waterfleas),and cyclestherians
outer shell. Crustacean classification as we know it
today was more or less established during the second IN FR A OR D E R S P IN IC A U D A TACI aM shr im os
half of the nineteenth century, although internal revi- (e.9.,Cyzic us, Eulim nadia, Im nadia, M etalimnadia)
sions continue. Martin and Davis (2001)presented an INFRAORDER CLADOCEROMORPHA
Water fleas
overview of crustacean classification, and readers are and cyclestheriids
referred to that publication for a window into the laby-
rinthine history of this subphylum. Our classification TRIBE CYCLESTHERIDAMonotypic: Cycle-
recognizes 1-1- classes,following Ahyong et al. (2011) stheria hislopi
and Martin et al. (201.4). TR IB E C LA D OC E R A W ater f leas ( e. g. ,
Anchistropus, Daphnia, Leptodora, Moina, Poly-
phemus)
CLASSIFICATION OF CRUSTACEA
CLASSMALACOSTRACA
CLASSREMIPEDIA Remipedes.One livingorder, Nec-
tiopoda (e.9., Cryptocorynectes,Godzillius,Lasionectes, SUBCLASSPHYLLOCARIDA
Pleomothra, Spe/eonectes)
ORDERLEPTOSTRACALeotostracansor nebali-
CLASSCEPHALoCARIDACephalocarids(e.9.,Chiltoniella, aceans (e.9., Dahlella,Levinebalia,Nebalia, Neb-
Hampsonellus,Hutchinsoniella,Lightiella,Sandersiella) aliella, Nebaliopsis,Paranebalia)
Branchiooods
CLASSBRANCHIOPODA HOPLOCARIDA
SUBCLASS
oRDERANoSTRAoA Fairyshrimps (e.9.,Artemia, OR D E RS TOMA TOP OD AManti s shr im ps ( e. 9. ,
Branchinecta,Branchinella,Streptocephalus) Echinosquilla,Gonodactylus,Hemisquilla,Squilla)
ORDERNOTOSTRACATadoole shrimps (Lepidu- SUBCLASSEUMALACOSTMCA
rus, Triops)
SUPERORDER
SYNCARIDASyncarids
ORDERDIPLOSTRACA The "bivalved" branchio-
ORDERBATHYNELLACEA(e.9.,Bathynella)
oods
ORDERANASPIDACEA(e.9., Anaspides,Psam-
SUBORDERLAEVICAUDATAF|at-tailed clam
maspides)
shrimps(e.9.,Lynceus)
SUPERORDER
EUCARIDA ORDERLOPHOGASTRTDA (e.9.,
Lophogastrids
ORDER EUPHAUSIACEA or krill(e.9.,
Euphausids, Gnathoph ausia,Lophogaster)
Bentheuphausia,
Euphausia,Meganyctiphanes, ORDERCUMACEACumaceans (e.9.,Campylaspis,
Nyctiphanes) Cumopsis,Diastylis,
Diastylopsis)
ORDERAMPHIONIDACEA Amphionids.
Monotypic: ORDERTANAIDACEATanaids(e.9.,Apseudes,
Amphionidpsreynaudii Heterotanais,Paratanais,Tanais)
ORDER
DECAPODACrabs,shrimps,lobsters,
etc. ORDERMfCTACEAMictaceans(e.9.,Mictocaris,
SUBORDER DENDROBRANCHIATA Penaeid etc.).Some researchers separateout the family
and sergestidshrimps(e.9.,Lucifer,Penaeus, Hirsutiidaeas a distinctorder(Bochusacea,contain-
Sergesfes,Sicyonia) ing Hirsutia,MontucarisandThetispelecaris).
SUBORDER
PLEOCYEMATA ORDERSPELAEOGRIPHACEA Spelaeogripha-
ceans. Four describedlivingspecies(Potiicoara
INFRAORDER CARIDEACaridean and procarid- brazilienses,
Spelaeogriphuslepidops,and two spe-
ean shrimps(e.9.,Alpheus,Crangon,Hippolyte, ciesof Mangkurtu),
and two knownfossilspecies
Lysmata,Macrobrachium,Palaemon,Pandalus, (theCarboniferous
Acadiocarisnovascofbaand the
Pasiphaea) UpperJurassicLiaoningogriphusquadripartitus)
INFRAORDER "Primitive"
PROCARlDOlDA shrimp ORDERTHERMOSBAENACEA Thermosbaena-
(Procaris, Vetericari
s) ceans(e.9.,Halosbaena,
Limnosbaena,Monodella,
Theosbaena,Thermosbaena,
Tulumella)
INFRAORDER STENOPODTDEA Stenopodidean
shrimps(e.9.,Sponglcola,
Stenopus) ORDER ISOPODAlsopods(seaslaters,rock lice,
pillbugs,
sowbugs,roly-polies)
INFRAORDER GLYPHEIDEA,.Primitive,,
Iobsters
(Neoglyphea,Laurentaeglyphea) SUBoRDERANTHURTDEA (e.g., Anthura,
INFRAORDER Colanthura, Cyathura, Mesanthu ra)
POLYCHELIDA Deep-seaslipper
lobsters(e.9.,Polycheles,
Stereomasfis) SUBORDER ASELLOTA(e.9.,Ase//us,Eurycope,
INFRAORDER BRACHYURA "True"crabs(e.9., Jaera,Janira,Microcerberus,
Munna)
Actaea,Callinectes,
Cancer,Cardisoma, Carcinus, SUBORDER (Calabozoa)
CALABOZOIDEA
Grapsus,Hemigrapsus,Maja,Menippe,Ocypode,
Pachygrapsus,Pinnotheres,Polydectus,Portunus, SUBORDERCYMOTHOIDA(e.g., Aega,
Scylla,Uca,Xantho) Bathynomus,
Cymothoa,Cirolana,Rocinela)
INFRAORDER ANOMURAHermitcrabs,galatheid SUBORDER EPICARIDEA(e.9.,Bopyrus,Dajus,

crabs,sand crabs,porcelaincrabs,etc. (e.9., Hemiarthrus, lone, Pseudio ne)
Birgus,Coenobita,Emerita,Galathea,Hippa,Kiwa, SUBORDER (e.9.,Gnathia,Para-
GNATHIIDEA
Lithodes,Lomis, Paguristes,Pagurus,Petrochirus, gnathia)
Petrolisthes,Pleuroncodes,Pylopagurus)
SUBORDER (e.9.,Limnoria,Keu-
LfMNORIDEA
INFMORDERASTACTDEA Crayfishes
andclawed phylia,Hadromastax)
(chelate)lobsters(e.9.,Astacus, Cambarus,
Homarus,Nephrops) SUBORDER MICROCERBERIDEA(e.9,
Atlantase
IIus, Microcerberus)
INFRAORDER ACHELATA(formerlyPalinura).
Palinurid,
spiny,and Spanish(slipper) lobsters SUBORDER oNISCIDEA(e.9.,Armadillidium,
(e.9.,Enoplometopus,Evibacus,lbacus,Jassa, Ligia, Oniscus,Porcellio,Trichoniscus,Tylos,
Jasus,Palinurus,
Panulirus,Scyllarus) Venezillo)
INFRAORDER AXIIDEA Lobster shrimos SUBORDER PHREATOICIDEA (e.9.,Mesam-

(e.9.,Axiopsis,Axius, Calocarides,Calocaris, phisopus,Phreatoicopis,
Phreatoicus)
Callianassa,
iVeaxrus) SUBORDER (Phoratopus)
PHORATOPIDEA
INFRAORDER GEBIIDEAMudandghostshrimps SUBoRDER SPHAEROMATTDEA (e.9.,Ancinus,
(e.9.,Axianassa,
Gebiacantha, na, Upo-
Thalassi Bathycopea,Paracerceis,
Paradella,Sphaeroma,
gebia) Sero/b,Tecticeps)
SUPERORDER
PERACARIDA SUBORDER TAINISOPIDEA(Pygolabis,
oRDERMYSTDAMysidsor opossumshrimps(e.9., Tainisopus\
Acanthomysis,Hemimysis,Mysis, Neomysis) SUBORDER VALVIFERA(e.9.,Arcturus,ldotea,
Saduria)
PHYLUMARTHROPODACrustacea:Crabs,Shrimps,and TheirKin 767
ORDER AMPHIPODA Amphipods-beach hoppers, SUBCLASSNEOCOPEPODA

whalelice,etc.
shrimps,
sandfleas,scuds,skeleton SUPERORDER
GYMNOPLEA
SUBORDER GAMMARIDEA (e.9.,Ampithoe,
ORDERCALANOIDA Calanoids(e.9.,Bathycalanus,
Anisogammarus, Corophium, Eurythenes, Calanus,Diaptomus, Eucalanus,Euchaeta)
Gammarus, Niphargus, Phoxocephalus,
Orchestia,
Talitrus) PODOPLEA
SUPERORDER
SUBORDER HYPERIIDEA (e.9., Cystisoma, oRDERcYcLoPolDA Cyclopoids(e.9.,Cyclopina,

Hyperia,Phronima,Primno,Rhabdosoma,Scrna, Cyclops, Eucyclops, Lernaea, Mesocyclops,
Streetsia,Vibilia) Notodelphys)
SUBORDER CAPRELLIDEA (e.9., Caprella, ORDERGELYELLOIDAGelyelloids(e.9.,Gelyella)

Phtisica,Syncyamus)
Cyamus,Metacaprella, (e.9.,Harp-
ORDERHARPACTICOIDAHarpacticoids
SUBORDER (e.9.,lngolfiella,
INGOLFIELLIDEA acticus, Longipedia, Peltidium, Porcellidium,
Metaingolfiella) P samm us, Sunarisfes,Ilbbe)
andtheirkin
Barnacles
THECOSTRACA
CLASS ORDERMISOPHRIoIDA Misophriods(e.9., Box-
shallia, Misophria)
SUBCLASS FACETOTECTA Monogeneric(Hansenoca-
"y-larvae,"
ris):the mysterious a groupof marinenauplii ORDERMONSTRILLOIDAMonstrilloids(e.9.,Mon-
andcypridsfor whichadultsareunknown strilla,Stilloma)
SUBCLASS ASCOTHORACIDA Two orders (Laurida, OR D E R MOR MON ILLOID A Mo r m onilloids.

Dendrogastrida) of parasiticthecostracans(e.9.,Asco- Monogeneric: Mormonilla
thorax, Dendrog aster,Laura, Synagoga, Zoanthoecus)
OR D E R P OE C ILOS TOMA TOIDPoecilost
A om a-
SUBCLASS the barnacles,
CIRRIPEDIACirripedes, and toids (e.9.,Chondracanthus,Erebonaster,Ergasilus,
theirkin Pseudanthessrus)
SUPERORDERTHORACICA Truebarnacles. Fouror- OR D E R S IP H ON OS TOMA TOIDASiPhO NO SI O -

ders,Lepadiformes (pedunculate or goose barnacles: matoids (e.g., Clavella, Nernesis, Penella,
e.9., Lepas),lbliformes,Scalpelliformes (Pol/atpes, PontoecielIa, Trebius)
Scalpellum), and Sessilia(sessileor acornbarnacles:
CLASSOSTRACODAOstracods
e.9.,Balanus,Chthamalus, Conchoderma, Coronula,
Tetraclita,Verruca) SUBCLASS MYODOCOPA
SUPERORDER ACROTHORACICA "bar-

Burrowing OR D E R MY OD OC OP ID A (e.g. , Cypr idina,
nacles."Twoorders,CryptophialidaandLythoglyptida Euphilomedes, Eusarsiella, Gigantocypris,
(e.9.,Cryptophialus, Trypetesa) Skogsbergia, Vargula)
SUPERORDER RHIZOCEPHALA PATASitiC,,bATNA- OR D E R H A LOC Y P R ID A (e,9., Conchoecia,
cles."Two orders,Kentrogonida and Akentrogonida Polycope)
accus, Lernaeodiscus, Mycetomorpha,
(e.9., Heteros
SUBCLASSPODOCOPA
Peltogaster,SaccuIina, Sylon)
ORDERPODOCOPIDA (e.g., Cypris, Candona,
CLASSTANTULOCARIDA Deepwater,marineparasites
Celtia, Darwinula, Limnocyth re)
(e.9.,BasipodeIIa, Deoterthron, Microdajus)
ORDERPLATCOP|DA (e.9., Cytherella,Sclero-
CLASSBRANCHIURA A singlefam-
Fishlice,or argulids.
cypris)
(e.S.,Argulus,Chonopeltis,Dipteropeltis,
ily (Argulidae)
Dolops) ORDERPALAEOCOPIDA(e.9.,Manawa)
CLASSPENTASTOMIDA Tongueworms.Two orders,nu-
merousfamilies(e.9.,Cephalobaena,
Linguatula,Pentas-
toma, Waddycephalus) Synopses of Crustacean Taxa
cLAsS MYSTAoooARIDAMystacocarids, with a single The following descriptions of major crustacean taxa
family(Derocheilocarididae), and 13 species(e.9.,Cteno- will give you an idea of the range of diversity within
cheiIocaris, DerocheiIocaris) the group and the variety of ways in which these suc-
cessful animals have exploited the basic crustacean
CLASS
COPEPODA body plan.
PROGYMNOPLEA
SUBCLASS
SubphylumCrustacea
ORDER PLATYCOPIOIDA (e.9.,Antri-
Platycopioids
cephalon(pluspre-
Bodycomposedof a 6-segmented
socopia,Platycopia)
segmental acron), or head, and multisegmented post-
768 Cha o te r T w e n tv -On e
cephalic trunk; trunk divided into thorax and abdo- The discovery of living remipedes in 1981 by ]ill
men (except in remipedes and ostracods); segments of Yager, strange vermiform crustaceans first collected
cephalon bear first antennae (antennules), second an- from a cavern in the Bahamas, gave the carcinological
tennae, mandibles, maxillules, and maxillae (see Table world a hrm. The combination of features distinguish-
20.2); one or more anterior thoracomeres may fuse with ing these creatures ispuzzling, for they possess charac-
the head (e.g., Remipedia and Malacostraca), their ap- teristics that are certainly very primitive (e.g., long, ho-
pendages forming'maxillipeds (secondarily modified monomous trunk; double ventral nerve cord; segmental
for feeding); cephalic shield or carapace present (sec- digestive ceca;cephalic shield) as well as some athibutes
ondarily lost in some groups); with antennal glands traditionally recognized as advanced (e.g., maxillipeds;
or maxillary glands (excretory nephridia); both simple nonphyllopodous [though flattened], biramous limbs).
ocelli and compound eyes in most groups, at least at They swim about on their backs as a result of metachro-
some stage of the life cycle; compound eyes stalked in nalbeating of the trunk appendages, similar to anostra-
many groups; with nauplius larval stage (suppressed cans. The remipedes are thus reminiscent of two other
or bypassed in some groups), and often a series of ad- primitive classes,the branchiopods and cephalocarids.
ditional larval stages. There are an estimated 70,000 F{owever, the laterally directed limbs are unlike those of
living species,in over 1,000families.3 any other crustacean/ and the "intemalized" mandibles
and the poison-injecting hypodermic maxillules are
Class Remipedia unique (the complex venom contains neurotoxins, pep-
Body of two regions, a cephalon and an elongate ho- tidases, and chitinases). The presence of the preanten-
monomous trunk of up to 32 segments, each with a nular processes is also puzzling, although similar struc-
pair of flattened limbs. Cephalon with a pair of sen- tures are known to occur in a few other crustaceans.
sory preantennular frontal processes; first antennae Some phylogenetic analyses based on morphological
biramous; trunk limbs laterally directed, biramous, data suggest that remipedes may be the most primitive
paddle-like, but without large epipods; rami of trunk living crustaceans, whereas molecular data remain am-
limbs (exopod and endopod) each of three or more biguous on the subject or, in some cases,ally them with
articles; without a carapace, but with cephalic shield cephalocarids andlor the Hexapoda.
covering head; midgut with serially arranged digestive All of the living remipedes discovered thus far are
ceca; first trunk segment fused with head and bearing found in caves (usually with connections to the sea)
one pair of prehensile maxillipeds; labrum very large, in the Caribbean Basin, Indian Ocean, Canary Islands,
forming a chamber (atrium oris) in which reside the and Australia. The water in these caves is often dis-
"internalized" mandibles; maxillules function as hypo- tinctly stratified, with a layer of fresh water overlying
dermic fangs; last trunk segment partly fused dorsally the denser salt water in which the remipedes swim.
with telson; telsonwith caudal rami; segmental double Remipedes hatch as lecithotrophic naupliar larvae,
ventral nerve cord; eyes absent in living species; male which is also unusual given their habitat (most cave
gonopore on trunk limb 15, female on 8; up to 45 mm crustaceans have direct development). Postnaupliar
in length. The above diagnosis is for the 24 known liv- development is largely anamorphic; juveniles have
ing remipedes (order Nectiopoda); the fossil record is fewer trunk segments than do the adults. Based on the
currentlybased on a single poorly preserved specimen three pairs of raptorial, prehensile cephalic limbs (and
(order Enantiopoda) (Figures 27.7A 27.3D-F, 27.21D, direct observations), it was long thought that remi-
21..22F,and 21.31E,F). pedes were strictly predators. However, studies by
Stefan Koenemann and his colleagues have suggested
they might also be capable of suspension feeding.
3Segments of the thorax are called thoracomeres (regardless of
whether or not any of these segments are fused to the head), Class Cephalocarida
whereas appendages of the thorax are called thoracopods. The
term "oereon" refers to that oortion of the thorax not fused to Head followed by 8-segmented thorax with each seg-
the heid (when such fusion occurs), and the terms "pereonites" ment bearing limbs, a 11-segmented limbless abdomen,
(= pereomeres) and "pereopods" are used for the segments and
appendages, respectiveiy, of the pereon. Hence, on a crustacean and a telson with caudal rami; common gonopore on
with the first thoracic segment (thoracomere 1) fused to the head, protopods of sixth thoracopods; carapace absent but
thoracomere 2 is typically called pereonite 1, the first pair of head covered by cephalic shield; thoracopods 1*7bira-
pereopods represents the second pair of thoracopods, and so on.
Be assured that we are trying to simplify, not confuse, this issue. mous and phyllopodous, with large flattened exopods
Also, we caution you that the homology of the thorax and abdo- and epipods (exites) and stenopodous endopods; tho-
men among the major crustacean lineages is probably more reverie racopods 8 reduced or absent; maxillae resemble thora-
than reality; the segmental homologies of the thorax and abdomen
have not yet been unraveled among the crustacean classes. For copods; no maxillipeds; eyes absen! nauplii with anten-
summaries of naupliar development across the group and iarval nal glands, adults with maxillary glands and (vestigial)
features in all crustaceans, see Martin et al. (2014). In most crusta- antennal glands (Figures 21.1T,21.3A-C, and 27.21A).
cean species the number of body segments is fixed, but in at least
two groups (notostracans and remipedes) the segment number can Cephalocarids are tiny, elongate crustaceans rang-
vary within a given species. ing in length from2 to 4 mm. There are 12 species in
.=-Protopod Figure 21 .3 Anatomy in the classes Remipedia and
Cephalocarida. (A) The cephalocarid H utch i nsoniella (lat-
eral view). (B) SEM of head and thorax of a cephalocarid.
(C) First trunk limb of the cephalocarid, Lightiella. (D) The
remipede, Spe/eonecfes (ventral view). (E) Tenth trunk
limb of the remipede, Lasionecfes. (F) The anterior end of
a remipede (ventral view). In both the cephalocarids and
the remipedes, the trunk is a long, homonomous series of
somites with biramous swimming appendages. In cepha-
locarids the first trunk appendages are like all the others,
which bear large swimming epipods (exites). In remipedes
the first trunk somite is fused to the head, and its append-
ages are maxillipeds.
zone to depths of over 1,500m, from the North and

South Pacific, to the North and South Atlantic, and
the Mediterranean.Hatching is at a slightly advanced
naupliar stage(calleda metanauplius).Most research-
ers agreethat cephalocaridsare very primitive crusta-
ceans,largely becauseof their relatively homonomous
body form, undifferentiated maxillae, long and gradu-
al (anamorphic)larval development, and shapeof the
trunk limbs (from which limbs of all other crustaceans
seemto be easilyderived).
Class Branchiopoda
Number of segmentsand appendageson thorax and
abdomen vary, the latter usually lacking appendages;
5 genera. All are benthic marine detritus feeders. Most carapacepresentor absent;telson usually with caudal
are associated with sediments covered by a layer of rami; body appendagesgenerallyphyllopodous; max-
flocculent organic detritus, although some have been illules and maxillae reduced or absent;no maxillipeds
found in clean sands. They occur from the intertidal (Figures21..IC,D,21.4,21.218,2L37C,and 21.35B).
770 Chapter Twenty-One
(A)
(F)
Digestive cecum
Food shing
Heart
Rostrum
Carapace
Antennule
Brood chamber Labrum
Shell gland
Midgut
Thoracic
appendage
Abdominal
Postabdominal
Processes
Postabdomen
Compound eye
0)
Sensoryfield
Antenna
Rostrum
First thoracopod
(modified as a clasper)
Carapace
Opercular lamellae (she11)
(covering anal somite)
Cercopods
(=caudal rami)
PHYLUM ARTHROPODA Crustacea:Crabs,Shrimps,and TheirKin 771
Compound eyes
Subfrontal plate
Food groove
Row of eftes
Egg pouch of eleventh

hunk appendage
Abdominal phyllopodia
Caudal ramus
Mandible
Anal somiie
Figure 21 .4 Anatomy and diversity in the class

Branchiopoda. (A) An anostracan (Branchinecta) in swim-
ming posture. (B) The anostracan, Branchipus schaefferi
swimming. (C) Trunk limb of an anostracan (Linderiella).
(D,E) The notostracan, Triops: (D) dorsal and (E) ventral
views. (F,G) Two cladocerans: (F) Daphnia and (G) Lep-
todora. (H) The shed carapace, or ephippium, ot Daphnia,
with the embryos enclosed. (l) Two extreme stages in the
seasonal change in head form of Daphnia (cyclomorpho-
sis). (J-L) The clam shrimp (Diplostraca), Lynceus: (J) the
valves are partially open (ventral view); (K) the head (ventral
view); (L) the whole animal, with one valve removed.
(M) The clam shrimp (Cyclestherida), Cyclestheria, SEM
Carapace
photo, with one valve removed (the specimen is slightly
(shell)
distorted; in life the shell is round).
772 Chapter Twentv-One
years or decadesuntil the next adequaterains appear. Triopsidae.Most speciesare 2-10 cm long. The com-
As diverse as the branchiopods might appear,both mon name derives from the generalbody shape:the
morphological and molecular analysesindicate that broad carapaceand narrow "trrtrk" give the animals a
they comprise a monophyletic group. Development superficialtadpole-Iikeappearance.
is remarkably uniform acrossthe group, with naupli- Notostracansinhabit inland waters of all salinities,
ar larvae distinguishableby severalunique features. but none occur in the ocean.Of the two known gen-
Severaltaxonomft nameshave beenproposed to clus- eta, Triops(Figure 21.4D,E)lives only in temporary
ter alleged sister-groupswithin the Branchiopoda,but waters, and its eggsare capableof surviving extended
there is as yet little consensus,and branchiopod rela- dry periods.Most speciesof Lepiduruslive in tempo-
tionships remain unsettled.Nearly 1,000extant species rary ponds,but at leastone species(L. arcticus)inhab-
have been described,and at leastone fossilbranchio- its permanentponds and lakes.However, all species
pod(Rehbachiella) is knownfrom the middle Cambrian, are short-lived, and most completetheir lifecycle in
confirming that they are an ancientgroup. just 30 to 40 days. Triopsis of someeconomicimpor-
tancein that large populations often occur in rice pad-
Order Anostraca Postcephalictrunk divisible into dies and destroy the crop by burrowing into the mud
appendage-bearing thorax of 11segments(17or 19 in and dislodging young plants. Tadpole shrimps mosily
members of the family Polyartemiidae)and abdomen crawl, but they are also capableof swimming for short
of 8 segmentsplus telson with caudal rami; gono- periods by beating the thoracic limbs. They are om-
pores on genital region of abdomen;trunk limbs bira- nivorous, feeding mostly on organic material stirred
mous and phyllopodous; small cephalicshield pres- up from the sediments,although some scavengeor
ent; paired, large, stalked compound eyesand a single prey on other animals,including molluscs,other crus-
median simple (naupliar) eye. taceans,frog eggs,and even frog tadpolesand small
The anostracansare commonly calledfairy shrimps fishes.Somespeciesof notostracansare exclusively
and brine shrimps. They differ from other branchio- gonochoristic,but othersmay include hermaphroditic
pods in lacking a carapace.There arejust over 300liv- populations (often thosepopulations living at high lat-
ing speciesin the order, worldwide, most of which are itudes). Earlier reports of parthenogeneticpopulations
less than 1 cm in length, although a few giants attain have been questioned.
lengths of 10 cm. Theseanimals inhabit ephemeral
ponds (including snowmelt ponds and desertpools), Order Diplostraca The clam shrimps and cladocer-
hypersaline lakes,and marine lagoons.In many areas ans(water fleas)comprisetwo groups of closelyrelated
they are an important food resourcefor water birds. branchiopodsknown as the Diplostraca (Figure21.4J-
Anostracans are sometimesunited with the extinct L). They sharethe feature of a uniquely developed,
order Lipostracaas the subclassSarsostraca. Their fos- large,"bivalved" carapacethat coversall or most of the
sil record datesback to the Devonian. body. Most diplostracansare benthic, but many swim
Anostracansswim ventral side up bv metachronal during reproductive periods. Someare direct suspen-
beating of the trunk appendages.Many use theselimb sion feeders,whereas others stir up detritus from the
movementsfor suspensionfeeding. Someother spe- substratum and feed on suspendedparticles, and oth-
ciesscrapeorganic material from submergedsurfaces, ers scrapepiecesof food from the sediment.
and at leasttwo species(Branchinecta gigas,B. raptor) The speciesformerly lumped together as clam
are specializedas predators on other fairy shrimps. shrimps are now partitioned between the two diplos-
Although most anostracanslive in isolatedponds, their tracan suborders,Laevicaudata and Onychocaudata.
eggsmight be transportedduring passagethrough the Diplostracansshareseveralfeatures:body divided into
gut of predatory divingbeetles (Dytiscidae). cephalonand trunk, the latter with 10-32segments,all
with appendages,and with no regionalizationinto tho-
Order Notostraca Thorax of 11segments,eachwith rax and abdomen;trunk limbs phyllopodous, decreas-
a pair of phyllopodous appendages;abdomen of ing in sizeposteriorly;maleswith trunk limbs 1,or 7-2,
"rings," each formed of more than one true segment; modified for grasping females during mating; trunk
eachanterior ring with severalpairs of appendages; typically terminates in spinous anal somite or telson,
posterior rings lack appendages;telson with long cau- usually with robust caudal rami (cercopods);gono-
dal rami; gonoporeson last thoracomere;broad, shield- pores on eleventhtrunk segment;bivalved carapace
like carapacefused only with head,but extendingto completelyenclosesbody; valvesfolded (Spinicaudata,
loosely cover thorax and part of abdomenipaired, ses- Cyclestherida)or hinged (Laevicaudata)dorsally; usu-
sile compound eyesand a single simple eye near ante- ally with a pair of sessilecompound eyesand a single,
rior midline on carapace. median,simple eye.
Notostracansare often called tadpole shrimps. The Laevicatdata, or flat-tailed clam shrimps, con-
There are only about a dozen living species,in two tains the single family Lynceidae, with close to 40
genera(TriopsandLepidurus)placed in a single farnlly, speciesin three genera,characterizedby a hinged,
globular carapace that encloses the entire animal. The is ectoparasitic on Hydra. Most of the benthic forms
Onychocaudata contains the spiny-tailed or "true" feed by scraping organic material from sediment par-
clam shrimp in the infraorder Spinicaudata, and the ticles or other objects; the planktonic species are sus-
cladocerans (commonly called water fleas) in the pension feeders. Some (e.g., Leptodora,Bythotrephes)are
infraorder Cladoceromorpha. Spinicaudatans in- predators on other cladocerans.
clude the commonly encountered freshwater genera In sexual reproduction, fertllization generally oc-
Limnadia, Eulimnidia, Leptestheria, and Cyzicus. The curs in a brood chamber between the dorsal surface of
Cladoceromorpha contains the well-known cladoc- the trunk and the inside of the carapace. Most species
eran water fleas Daphnia, Moina, Diaphonosoma,and have direct development. In the family Daphnidae the
Leptodora, as well as the unique genus Cyclestheria (in developing embryos are retained by a portion of the
the Cyclestherida). Confusingly, Cyclestheriais also shed carapace, which functions as an egg case called an
commonly called a clam shrimp, though it is more ephippium (Figure 21..4H),whereas in the Chydoridae
closely related to the water fleas). the ephippium remains attached to the entire shed
The common name "clam shrimo" derives from the carapace. Leptodoraexhibits a heterogenous life cycle,
clamlike appearance of the valves, which usually bear altemating between parthenogenesis and sexual repro-
concentric growth lines reminiscent of bivalved mol- duction, the latter of which results in free-living larvae
luscs. The approximately 200 species of clam shrimps (metanauplii hatch from the shed resting eggs).
(including laevicaudatans, spinicaudatans, and Cladoceran life histories are often compared with
Cyclestheria)live primarily in ephemeral freshwater those of animals such as rotifers and aphids. Dwarf
habitats worldwide. Cyclestheriahislopi,the only mem- males occur in many species in all three groups, and
ber of the Cyclestherida, inhabits permanent freshwa- parthenogenesis is common. Members of two cladoc-
ter habitats throughout the world's tropics, and is one eran families that undergo parthenogenesis (Moinidae
of the most widespread animals on Earth. Cyclestheria and Polyphemidae) produce eggs with very little yolk.
is also the only clam shrimp with direct development, In these groups the floor of the brood chamber is lined
the larval and juvenile stages being passed within the with glandular tissue that secretes a fluid rich in nu-
brood chamber, one of the features allying it with the trients, which is absorbed by the developing embryos.
cladocerans. Periods of overcrowding, adverse temperatures/ or
In cladocerans, the carapace is never hinged (only food scarcity can induce parthenogenetic females to
folded dorsally, like a taco) and never covers the entire produce male offspring. Occasional periods of sexual
body, and appendages do not occur on all the trunk reproduction have been shown to occur in most par-
somites. The body segmentation is generally reduced. thenogenetic species. Many planktonic cladocerans
The thorax and abdomen are fused as a "trunk" bear- undergo seasonal changes in body form through suc-
ing 4-6 pairs of appendages anteriorly and terminat- ceeding generations of parthenogenetically produced
ing in a flexed "postabdomen" with clawlike caudal individuals, a phenomenon known as cyclomorphism
rami. Trunk appendages are usually phyllopodous. (Figure 21,.4I).
The carapace usually encloses the entire trunk, but not
the cephalon, serving as a brood chamber (and greatly Class Malacostraca
reduced to this function) in some species; a single me- Body of 19-20 segments, including 6-segmented ceph-
dian compound eye is always present. alon, 8-segmented thorax, and 6-segmented pleon
The cladocerans, or water fleas, include about 400 (7-segmented in leptostracans), plus telson; with or
species of predominantly freshwater crustaceans, al- without caudal rami; carapace covering part or all
though several American marine genera and species of thorax, or reduced, or absent; 0-3 pairs of maxil-
are known (e.9., Eaadne,Podon). Although there are lipeds; thoracopods primitively biramous, uniramous
relatively few species, the group exhibits great morin some groups/ phyllopodous only in members of the
phological and ecological diversity. Most cladocerans subclass Phyllocarida; antennules and antennae usu-
are 0.5-3 mm long, butLeptodorakindtii reaches18 mm ally biramous; abdomen (pleon) usually with 5 pairs
in length. Except for the cephalon and large natatory of biramous pleopods and 1 pair of biramous uropods;
antennae, the body is enclosed by a folded carapace, eyes usually present, compound, stalked or sessile;
which is fused with at least some of the trunk region. mainly gonochoristic; female gonopores on sixth, and
The carapace is greatly reduced in members of the male pores on eighth thoracomeres. \Alhen uropods are
families Polyphemidae and Leptodoridae, in which it present, they are often broad and flat, lying alongside
forms a brood chamber. the broad telson to form a tail fan.
Cladocerans are distributed worldwide in nearly all Most classification schemes divide the more than
inland waters. Most are benthic crawlers or burrowers; 40,200species of malacostracans into three subclasses,
others are planktonic and swim by means of their large Phyllocarida (leptostracans), Hoplocarida (stomato-
antennae. One genus (Scapholeberis)is typically found pods), and the megadiverse Eumalacostraca. The
in the surface film of ponds, and another (Anchistropus) phyllocarids are typically viewed as representing the
(A)
Adductor Ovary Maxillary

palp
venmal
nerve cord
Hindgut
Compound eye
Seventh abdominal
Roshum segment
pleopod
-Fifth
Antennal
gland
Endopod
Antennule First pleopod
Thoracopods
Figure 21 .5 Anatomy in leptostracans (class
Malacostraca, subclass Phyllocarida). (A) General
anatomy of Nebalia. (B) Phyllopodous swimming limb of
Nebalia. (C) SEM ot Nebalia. (D) Anterior end of an oviger-
ous Nebalia.
primitive malacostracancondition (6-8-7body seg-

ments plus telson;Figure 21.5).The basiceumalacos-
tracan body plan, characterizedby the 6-8-6(plus tel-
son) arrangementof body segments,was recognized
in the early 1900sby W.T. Calman, who termed the
defining features of the Eumalacostraca"caridoid fa-
cies" (Figure 27.6).Much work has been done since
Calman's day, but the basicelementsof his caridoid thorax and compressed laterally so as to from an
faciesare still presentin all membersof the subclass unhinged bivalved "sheIl," with an adductor muscle;
Eumalacostraca. cephalon with a movable, articulated rostrum; pleo-
pods 1-4 similar and biramous, 5-6 uniramous; no uro-
Subclass Phyllocarida pods; paired stalked compound eyes; antennules bira-
Order Leptostraca With the typical malacostracan mous; antennae uniramous; adults with both antennal
characteristics,exceptnotable for presenceof seven and maxillary glands (Figures 21.5 and 27.21C).
free plpomeres (plus telson) rather than six, generally The subclass Phyllocarida includes about 40 species
taken to representthe primitive condition for the class. in 10 genera. Most are 5-15 mm long, butNebiliopsis
Also, with phyllopodous thoracopods(all similar to typicais a giant at nearly 5 cm in length. The leptostra-
one another); no maxillipeds; large carapacecovering can body form is distinctive, with its loose bivalved
Cephalothorax
Thoracic
Figure 21 .6 The basic eumalacostran Carapace somites
body plan and the "caridoid facies."
Note the thick (muscled) abdomen and j \i .-, Antennule
the tail fan, which work in combination :ii
:z'tti
to produce a powedul tail flip escape r!:
reaction. Antennal scale

Telson
Mandible
r'fan- x(
Tail Antenna
Pleopods
LY
PHYLUM ARTHROPODA CrustACEA:CrAbS, Shrimps,and TheirKin 775
(B) Compound eye

Carapace
Antennule
Antenna
Antennal
scale
Pleopods Eighth/ Penis / o#".fhs

thoracopod Legs3-5 (secondthoracoPod)
(gnathopods)
Cephalon
Thoracomeres
Antennule
Pleotelson
Rudimentary
pleopods
2-8
Cephalon [-Thoracomeres
Antenna Maxilliped
Figure 21 -7 External anatomy in the class Malacostraca, subclass

Eumalacostraca-stomatopods and 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, Bathynella. (E) An anaspidacean, Stygocarella.
carapacecovering the thorax, a protruding rostrum, Subclass Hoplocarida

and an elongateabdomen.All leptostracansare ma- Order Stomatopoda Carapacecovering portion of
rine, and most are epibenthic from the intertidal zone headand fusedwith thoracomeres1-4;head with mov-
to a depth of 400 m; Nebaliopsistypicais bathypelagic. able,articulated rostrum; thoracopods1-5 uniramous
Most speciesseemto occur in low-oxygen environ- and subchelate,secondpair massiveand raptorial (all
ments.One species,Dahlellacaldariensis, is associated five are sometimescalled "maxillipeds" or gnathopods
with the hydrothermal vents of the Galapagosand becausethey are involved in feeding); thoracopods
the EastPacific Rise.Speonebalia cannoniis known only 6-8 biramous, ambulatory;pleopodsbiramous,with
from marine caves. dendrobranchiate-likegills on exopods;antennulestri-
Most leptostracanssuspensionfeed by stirring up ramous;antennaebiramous,with large,paired, stalked
bottom sediments.They are also capableof grasping compound eyesthat are unique in the animal kingdom
relatively large bits of food directly with the mandi- (Figures21.7A-C, 21.27D,and 21.33K).
bles.Someare carnivorousscavengers,and someare All500 or so living hoplocarids are placed in the
known to aggregatein areason the seafloor where order Stomatopoda,known as mantis shrimps. They
large amounts of detritus accumulate.In many species are relatively large crustaceans,ranging in length from
the antennaeor antennulesof males are modified to 2 to 30 cm. Compared with that of most malacostra-
hold femalesduring copulation. cans,the muscle-filledabdomenis notably robust.
776 Chaoter Twentv-One
Most stomatopods are found in shallow tropical or and extant members of the order Anaspidacea(e.g.,
subtropical marine environments. Nearly all of them Anaspides), it has been suggestedthat syncaridsmay
live inburrows excavated in soft sediments or in cracks encompassthe most primitive living eumalacostra-
and crevices, among rubble, or in other protected can body plan. Bathynellaceansoccur worldwide
spots. All species are raptorial carnivores, preying on in interstitial or groundwater habitats, whereas the
fishes, molluscs, cnidarians, and other crustaceans. The anaspididaceansare strictly Gondwanan in distribu-
large, distinctive sirbchelae of the second thoracopods tion. Many Anaspidaceaare endemic to Tasmania,
act either as crushers or as spears (Figure 21.7C). where they inhabit freshwater environments, such as
Stomatopods crawl about using the posterior thora- open lake surfaces,streams,ponds, and crayfish bur-
copods and the flaplike pleopods. They also can swim rows. No syncarids are marine. Thesereclusive eum-
by metachronal beating of the pleopods (the swimmer- alacostracans show various degreesof what somehave
ets). For these relatively large animals, living in narrow regarded as paedomorphism, including small size
burrows requires a high degree of maneuverability. (Anaspididaeincludesmembersto 5 cm, whereasmost
The short carapace and the flexible, muscular abdomen othersare lessthan 1 cm long), eyelessness, and reduc-
allow these animals to twist double and turn around tion or lossof pleopodsand someposterior pereopods.
within their tunnels or in other cramped quarters. This Bathynellaceansare small (1-3 mm long), possess6 or
ability facilitates an escape reaction whereby a mantis 7 pafusof long, thin swimming legs, and have a pleo-
shrimp darts into its burrow rapidly head first, then telson formed by the fusion of the telson to the last
turns around to face the entrance. pleonite.
Stomatopods are one of orLly two groups of malacos- Syncarids either crawl or swim. Little is known
tracans that possess pleopodal gills. Only the isopods about the biology of most species,although some are
share this trait, but the pleopods are quite different in considered omnivorous. Unlike most other crusta-
the two groups. The tubular, thin, highly branched gills ceans,which carry the eggsand developing early em-
of stomatopods provide a large surface area for gas ex- bryos, syncarids lay their eggs or shed them into the
change in these active animals. water following copulation.
Subclass Eumalacostraca Superorder Eucarida Telsonwithout caudal rami;
Head, thorax, and abdomen of 6-8-6 somites respec- 0,1.,or 3 pairs of maxillipeds; carapacepresent,cov-
tively (plus telson); with 0, 7,2, or 3 thoracomeres fused ering and fused dorsally with head and entire thorax;
with head, their respective appendages usually modi- usually with stalked compound eyes;gills thoracic.
fied as maxillipeds; antennules and antennae primi- Although members of this group are highly diverse,
tively biramous (but often reduced to uniramous); they are united by the presenceof a completecarapace
antennae often with scalelike exopod; most with well that is fused with all thoracicsegments,forming a char-
developed carapace, secondarily reduced in slmcarids acteristiccephalothorax.Most species(severalthou-
and some peracarids; gills primitively as thoracic epi- sand) belong to the order Decapoda.The other two
pods; tail fan composed of telson plus paired uropods; orders are the Euphausiacea(krill), and the monotypic
abdomen long and muscular. Three superorders: S1m- Amphionidacea.
carida, Eucarida, and Peracarida.
Order Euphausiacea Euphausidsare distinguished
Superorder Syncarida Without maxillipeds (Bathy- among the eucaridsby the absenceof maxillipeds, the
nellacea) or with one pair of maxillipeds (Anaspid- exposureof the thoracic gills external to the carapace,
acea); no carapacei pleon bears telson with or with- and the possessionof biramous pereopods(the last 1 or
out furcal lobes; at least some thoracopods biramous, 2pairs sometimesbeing reduced).They are shrimplike
eighth often reduced; pleopods variable; compound in appearance.Adults have antennal glands. Most of
eyes present (stalked or sessile) or absent (Figure them have photophores on the eyestalks,the basesof
21,.7D,E).There are about 285 described species of s;m- the secondand sevenththoracopods,and between the
carids in two orders, Anaspidacea and Bathlnellacea.a first 4 pairs of abdominal limbs.
To many workers, the syncarids represent a key group The 90 or so speciesof euphausidsare allpelagic and
in eumalacostracan evolution, and they may rep- rangein length from 4 to 15cm. The pleopods function
resent an ancient relictual taxon that now inhabits as swimmerets.Euphausidsare known from all ocean-
refugial habitats. Through studies of the fossil record ic environmentsto depths of 5,000m. Most speciesare
distinctly gregarious,and speciesthat occur in huge
schools(krill) provide a major sourceof food for larger
4Until recently a third slmcarid
order was recognized, the nektonic animals (baleenwhales, squids, fishes) and
Stygocaridacea, endemic to the Southern Hemisphere. Most work-
ers now agree that the stygocarids should be reduced to the rank evensomemarine birds. Krill densities,particularly for
of family within the order Anaspidacea. Euphausia superba,often exceed1,000animals/m3(6t+
(A) Carapace
Compound eye
Antennule
Pereopod Branchial I
Pleopod
epipod
Photophore
Antenna
Carapace
Antennule
Propodus
Fifth pleopod
Maxilliped First pleopod
Figure 21 .8 Anatomy in the class Malacostraca, super-

order Eucarida-euphausids and amphionidaceans.
(A) General body form of the euphausid, Meganyctiphanes.
(B) Pereopod of Euphausia superba. (C) Amphionides
reynaudii (female), the only living species of the
Amphionidacea.
first pair in femalesis uniramous and greatly enlarged,
perhaps functioning to form a brood pouch extend-
g wet weight/m3).s Generally,euphausidsare suspen- ing under the thorax. Femaleshave a reduced gut and
sion feeders,although predation and detritivory also apparently do not feed.Amphionidesis a worldwide
occff (Figures21.8,4',8arrd2L.2lB). member of marine oceanicplankton and occurs to a
depth of 1.,700 m (Figure21.8C).
Order Amphionidacea The singleknown speciesof
the order Amphionid acea,Amphionidesreynaudii,p os- Order Decapoda The decapodsare among the most
sessesan enlargedcephalothoraxcoveredby a thin, familiar eumalacostracans. They possessa well devel-
almost membranous carapacethat extendsto enclose oped carapaceenclosinga branchial chamber,but they
the thoracopods.The thoracopods are biramous with differ from other eucarid ordersin always possessing3
short exopods. The first pair is modified as maxil- pairs of maxillipeds,leaving 5 pairs of functional unira-
lipeds, and the last pair is absentin females.Someof mous or weaklybiramous pereopods(hencethe name,
the mouthparts are highly reduced in females.The Decapoda);one (or more) pairs of anteriorpereopods
pleopods are biramous and natatory,exceptthat the are usually clawed (chelate).Adults have antennal
glands.Rearrangementof the subtaxawithin this order
sWhere krill densities exceed about 100 grams per cubic meter, is a popular carcinologicalpastime (seeMartin and
they are often fished commercially. Krill schools can extend for Davis 2001for an entry into the vast literature on deca-
tens of miles, contain millions of tons of krill, and stain the ocean
pod classification).In vernacularterms,nearly every
red with their surface swarms in coastal waters. Large baleen
whales can eat a ton of krill in one mouth-ful. Seals, fish, squid, decapodmay be recognizedas some sort of shrimp,
and humans also eat krill. Krill fishine has been banned in most crab,lobster,or crayfish.
of North America, but it continues inJapan where tens of thou-
We do not want to belabor the issue of decapodgill
sands of tons are landed annually and used mainly as feed for
farmed fish. The largest krill fishery is in the ocean surrounding nomenclature.However, the gills play a prominent role
Antarctica, where they have been harvested commercially since in the taxonomy of this group; thus, we provide brief
the 1970s. In the 1980s, large fleets from the Soviet Union caught
descriptionsof the basic types. All decapod gills arise
up to 400,000 tons of Antarctic krill annuallv, but the current
citch is now down to about 120,000 tons (taken by Japan, Korea, as thoracic coxal exites(epipods),but their final place-
Norway, Poland, the Ukraine, and the U.S.). ment varies. Thosethat remain attachedto the coxaeare
778 Ch a o te r T w e n tv -On e
podobranchs (= "foot gills"), but others eventually be- Suborder Dendrobranchiata This group includes
come associated with the articular membrane between over 500 speci es of decapods, most of which ar e
the coxae and body and are thus called arthrobranchs (= penaeid and sergestid shrimps. As the name indicates,
'Joint gills"). Some actually end up on the lateral body these decapods possess dendrobranchiate gills (Figure
wall, or side-surface of the thorax, as pleurobranchs 21,.288),a unique synapomorphy of the taxon. One
(= "side gills"). The sequenceby which some of these genus/ Lucifer, has secondarily lost the gills completely.
gills arise ontogdnetically varies. For example, in the The dendrobranchiate shrimps are further character-
Dendrobranchiata and the Stenopodidea, arthrobranchs izedby chelae on the first three pereopods, copula-
appear before pleurobranchs, whereas in members of tory organs modified from the first pair of pleopods
the Caridea the reverse is true. In most of the other deca- in males, and ventral expansions of the abdominal ter-
pods the arthrobranchs and pleurobranchs tend to ap- gites (called pleural lobes). Generally, none of the che-
pear simultaneously. These developmental differences lipeds is greatly enlarged. In addition, females of this
may be minor heterochronic dissimilarities and of less group do not brood their eggs. Fertilization is external,
phylogenetic importance than actual gill anatomy. and the embryos hatch as nauplius larvae (see the sec-
Among the decapods, the gills can also be one of tion on Reproduction and Development later in this
three basic structural types, described as dendrobran- chapter). Many of these animals are quite large, over
chiate, trichobranchiate, and phyllobranchiate (Figure 30 cm long. The sergestids are pelagic and all marine,
21.28F.-D). All three of these gill types include a main whereas the penaeids are pelagic or benthic, and some
axis carrying afferent and efferent blood vessels, but occur in brackish water. Some dendrobranchiates
they differ markedly in the nature of the side filaments (e.9., Penaeus,Sergestes, Acetes,Sicyonia) are of major
or branches. Dendrobranchiate gills bear two principal commercial importance in the world's shrimp fisher-
branches off the main axis, each of which is divided ies, most of which are now being exploited beyond
into multiple secondary branches. Trichobranchiate sustainable levels and often with fishing techniques
gills bear a series of radiating unbranched tubular fila- that are highly habitat-destructive (Figures 27.9Aand
ments. Phyllobranchiate gills are characterizedby a 2r.33G).
double series of platelike or leaflike branches from the
axis. Within each gill type, there may be considerable Suborder Pleocyemata All of the remaining deca-
variation. The occurrences of these three major gill pods belong to the suborder Pleocyemata. Members
types among various taxa are presented below. of this taxon never possess dendrobranchiate gills.
Close inspection of the proximal parts of the pe- The embryos are brooded on the female's pleopods
reopods usually reveals another decapod feature: in and hatch at some stage later than the nauplius
most forms, the basis and ischium are fused (as a basi- larva. Included in this suborder are several kinds of
ischium), with the point of fusion often indicated by a shrimps, the crabs, crayfish, lobsters, and a host of
suture line. Tegumental glands are also a ubiquitous less familiar forms. Most current workers now rec-
feature among the Decapoda. These glands originate ognize 11 infraorders within the Pleocyemata, as we
below the epidermal cells and produce a fluid that have done below, but a number of other schemes have
opens on the surface of the cuticle. They have been been proposed and persist in the literature. One older
reported from gills, legs, pleopods, and uropods. The approach divided decapods into two large groups,
roles of tegmental glands are not well known, and they called the Natantia and Reptantia-the swimming
have been suspected to be involved in cuticular tan- and walking decapods, respectively. Although these
ning, the production of mucus by the mouthparts, the terms have largely been abandoned as formal taxa,
production of cement substance involved with egg at- they still serve a useful descriptive purpose, and one
tachment, and possibly also grooming. continues to see references to natant decapods and
The 18,000or so living species of decapods comprise reptant decapods.
a highly diverse group. They occur in all aquatic envi-
ronments at all depths, and a few spend most of their Infraorder Procarididea The procarids consist of a sin-
lives on land. Many are pelagic, butbthers have adopt- gle family containing two genera of shrimp, Procaris and
ed benthic sedentary, errant, or burrowing lifestyles. Vetericaris,that have been called "primitive shrimps."
Decorating of the exoskeleton is frequently seen among Like caridean shrimp (below) and most other pleocye-
the decapods, especially in spider crabs (Brachyura: mates, they bear flattened, platelike (nondendrobranch)
Majoidea), which use Velcro-like hooked setae to at- gills, but they lack claws on any of their legs, have a very
tach dead or living plants and animals; decorating has leglike (pediform) third maxilliped, and have epipods
been shown to reduce predation through camouflage on all of the maxillipeds and pereopods, assumed to be
and/ or chemical deterrence. Decapod feeding strate- an ancestral condition because most decapod groups no
gies include suspension feeding, predation, herbivory, longer bear the full complement. Th"y are known from
scavenging, and more. Most workers recognize two anchialine habitats-inland pools with connections to
suborders: Dendrobranchiata and Pleocvemata. the sea (Figure 21.98).
PHYLUM ARTHROPODA Crustacea:Crabs, Shrimps,and TheirKin 779
(B)
Compound Secondpleomere
Antennule
Antenna
Antennal
scale
Third
maxillipeds
Pereopods----J Pleopods
Figure 21 .9 External anatomy and diver-

sity in shrimps (Decapoda). (A) A penaeid shrimp
(Dendrobranchiata), Penaeus seflferus. (B) A procarid
shrimp (Procarididea), Procaris ascenslonis. (C) A hippo-
lytid sh rimp (Caridea, H ippolytidae), Lysmata califo rn ica.
(D) An alpheid, or snapping shrimp (Caridea,Alpheidae),
Alpheus. (E) A stenopodid shrimp (Stenopodidea),
Stenopus.
InfraorderCaridea The nearly 3,500livingspeciesin associatedwith shallow benthic environments, espe-

this infraorder are generallyreferredto as the caridean cially with coral reefs;othersare known from the deep
shrimps. Theseswimming decapodshave phyllobran- sea.Many are commensal,and the group includes
chiate gills. The first 1 or 2 pairs of pereopodsare che- the cleanershrimps (e.g.,Stenopas) of tropical reefs,
late and variably enlarged.The secondabdominalpleu- which are known to remove parasitesfrom local fish-
ra (side walls) are distinctly enlarged to overlap both es.Stenopodidsoften occur as male-femalecouples.
the first and third pleura. The first pleopods are gener- Perhapsthe most noted example of this bonding is
ally somewhatreduced,but not much modified, in the associatedwith the glasssponge(Euplectella) shrimp,
males(Figures21.1K,21.9C,D,2L24D, and 21.31D). Spongicola aenusta.A young male and female shrimp
enter the atrium of a host sponge,eventually growing
InfraorderStenopodidea The 70 or so speciesin this too large to escapeand thus spending the rest of their
infraorder belong to three families. The first 3 pairs of days together.
pereopodsare chelate,and the third pair is significant-
ly larger than the others.The gills are trichobranchiate. Infraorder Brachyura Theseare the so-called"true
The first pleopodsareuniramous in malesand females, crabs." The abdomen is symmetrical but highly
but are not strikingly modified. The secondabdomi- reduced and flexed beneath the thorax, and uropods
nal pleura are not expandedas they are in carideans are usually absent.The body, hidden beneatha well-
(Figures2L.9Eand 27.318). developed carapace,is distinctly flattened dorso-
Thesecolorful shrimps are usually only a few cen- ventrally and often expandedlaterally. The gills are
timeters long (2-7 cm). Most speciesare tropical and typically phyllobranchiate, but exceptionsoccur.
(A) Figure 21 .10 Anatomy and diversity of the "true," or brachy-

Compound uran, crabs (Decapoda: Brachyura). (A,B) General crab anatomy:
Palp of maxilliped Antennule frontal and ventral views of a swimming crab (family Portunidae).
eye
Antenna (C) A spider crab (family Majidae), Loxorhynchus. (D) A kelp crab
Carapace Carpus
/orat (Majidae), Pugettia. (E) An arrow crab (Majidae), Stenorhynchus.
(F) A cancer crab (family Cancridae), Cancer. (G) A grapsid crab
(family Grapsidae), Pachygrapsus. (H) A pinnotherid or pea crab
(family Pinnotheridae), Parapinnixa. (l) A xanthid crab (family
Xanthidae), Trapezia. Many members of this genus are obligate
commensals in scleractinian corals. (J) A dromiid crab (fam-
Third maxilliped ily Dromiidae), Hypoconcha (anterior view). Members of the
Dromiidae carry bivalve mollusc shells (or other objects) on their
backs. (K) A calappid crab (family Calappidae), Hepatus (ante-
rior view). (L) Ventral views of a female (upper photo) and (lower
photo) male Hemigrapsus sexdentatus.
(c)
Claw of cheliped
Basi-ischium
Propodus
Dactyl
Carpus Coxa
Thoracic sterna
Merus Basis
Abdomen
Fifth pereopod
about a dozen families) all have direct development,

incubate their embryos, and are independent of sea
water. Somefreshwater crabs are intermediate hosts
of Paragonimus, a cosmotropicalparasitic human lung
fluke, and others are obligate phoretic hosts of larval
black flies (Simulium),the vector for Onchocerca
aolaulus
(the causativeagent of river blindness). A number of
speciescarry other invertebrateson their carapace(e.g.,
sponges,tunicates)or on their claws (e.g.,anemones);
these associationsare generally thought to be mutu-
(K)
alistic, providing camouflage or predator deterrence
for the crab while their partner is moved about in the
environment and may feed off debris from their host's
feeding activities. The opener photo for this chapter
shows the tropical Pacific teddy-bear crab,Polydectus
cupulifer,which is densely coveredwith setaeand fre-
quently carries a sea anemone on each cheliped. In
the northeast Pacific, megalopaeand juveniles of the
crabCancergracilisride (and feed) on the bell of cer-
tain jellyfish; individuals of Phacellophora
camtschatica
(Scyphozoa)have been found with hundreds of C. gra-
ciils megalopae.There are about 7,000described spe-
ciesof Brachyura.
Infraorder Anomura This group includes hermit

crabs, galatheid crabs, king crabs, porcelain crabs,
mole crabs, and sand crabs. The abdomen may be
soft and asymmetrically twisted (as in hermit crabs)
or symmetrical, short, and flexed beneath the thorax
(asin porcelain crabsand others).Thosewith twisted
abdomenstypically inhabit gastropod shells or other
empty "houses" not of their own making. King crabs
(Lithodidae and Hapalogastridae)probably evolved
out of hermit crab-like ancestors.Carapaceshape
and gill structure vary. The first pereopodsare che-
late; the third pereopodsare never chelate.The sec-
ond, fourth, and fifth pairs are usually simple, but
occasionallythey are chelateor subchelate.The fifth
pereopods (and sometimes the fourth) are generally
much reduced and do not function as walking limbs;
the fifth pereopodsfunction as gill cleanersand often
The first pereopods are chelateand usually enlarged. are not visible externally.The pleopods are reduced or
Pereopods2 to 5 are typically simple, stenopodous absent.The eyes are positioned medial to the anten-
walking legs,although in somegroups/the fifth pereo- nae. The zoealarva is similar to that of the true crabs
pods are also chelate.The eyesare positioned lateral to but is typically longer than broad, with the rostral
the antennae.Males lack pleopods 3 to 5. The distinc- spine directed anteriorly.Most anomuransare marine,
tive larval stageis called a zoea;its carapaceis spheri- but a few freshwater and semi-terrestrial speciesare
cal and bears a ventrally directed rostral spine (or no known. The so-called"yeti crab" (Kiwa hiisuta) was
spine) (Figures 21.18, 21.10,21,.27H,2L.28F,G,21..29 C, discovered in 2005 from2,200 m-deep hydrothermal
21.32,and21,.33H,I). vents south of EasterIsland. It is remarkable for its
Brachyuran crabs are mostly marine, but freshwa- " gatden" of filamentous bacteria that grow on the long
ter, semi-terrestrial,and moist terrestrial speciesoccur setaeof the exoskeleton;the bacteriaareheterotrophic,
in the tropics. The land crabs (certain speciesin the utilizing sulfides in the deep environment (the pre-
families Gecarcinidae,Ocypodidae,Grapsidae,etc.) ciserole of the bacteria,of severalspecies,in the life
are still dependent on the oceanfor breeding and lar- history of the yeti crab is not yet well understood). A
val development. The surprisingly large number of secondspeciesof Kiwa was describedin 2011(Figures
freshwater crabs (about 3,000species,classifiedinto 21.IF-L 2L.11C-G,21.24A-C, 21.3IA, and 21.33I).
(A)
CepholothoraVcarapace Major cheliped
Walking legs
Antenna Antennule
Cheliped.
| |
Lpereooods r-5 I (Pereopod-.
1)
'::--:\-l-
i:-;:t-l
:- L - t.
:) rfuP\,\
v_\L--ri-v
_--t 2
-J L
Figure 21 .1 1 External anatomy and diversity in some
other reptant decapods (Eumalacostraca, Eucarida).
(A) A mud shrimp, Callianassa (Gebiidea). (B) A spiny lob-
ster, Panulirus (Achelata). (C) A hermit crab, Paguristes, in
its shell (Anomura). (D) A hermit crab, Pagurus, removed
from its shell to expose the soft abdomen. (E) A porcelain
crab, Petrolisthes (Anomura), with the reduced posterior
pereopods extended. (F) A sand crab, Emerita (Anomura).
(Anomura),
umbrellacrabCryptolithodes in ventral
Infraorder Astacidea The 650+ species of crayfish l?Jl"
and clawed lobsters comprise some of the most famil-
iar decapods (Figure 21.2).As in most other decapods,
the dorsoventrally flattened abdomen terminates in a of crayfish occur in North America alone, where they
strong tail fan. The gills are trichobranchiate. The first 3 show high levels of endemicity to particular regionsor
pairs of pereopods are always chelate, and the first pair river drainages.(Figures21.27E,Gand 21.298).
is greatly enlarged. Homarus americanus,the American
or Maine lobster, is strictly marine and is the largest liv- Infraorder Achelata This group includes the coral
ing crustacean by weight (the record weight being over and spiny lobsters (family Palinuridae) and slipper
20 kilograms). Most crayfish live in fresh water, but a lobsters(family Scyllaridae).The nameAchelatacomes
few species live in damp soif where they may excavate from the fact that they lack chelaeon all pereopodsas
extensive and complex burrow systems. The 600+ spe- adults (exceptfor a small grooming claw on pereo-
cies of freshwater crayfish comprise a monophyletic pod 5 in some females).The flattened abdomen bears
group that is sister to clawed lobsters. Over 425 species atailfan; the carapacemay be cylindrical or flattened
dorsoventrally; the gills are trichobranchiate. The large, prejuvenile stagelacking the last pair of thoracopods
flattened larvae, called phyllosomas becauseof their (no free-living larvae occur in this group) (Figures
leaf{ike appearance,are unique and distinctive. All 21.72-21.15).
speciesare marine, and they are found in a variety of The roughly 25,000speciesofperacaridsare divided
habitats throughout the tropics.Ma.y speciesproduce among nine orders. The peracarids are an extremely
soundsby rubbinga process(the plectrum) at the base successfulgroup of malacostracancrustaceansand
of the antennae against i, "file" on the head (Figures are known from many habitats. Although most are
21.1M,21.11B, 21.30A,C, and 21.33L). marine, many also occur on land and in fresh water,
and severalspecieslive in hot springs at temperatures
Infraorders Gebiidea and Axiidea Thesetwo infra- of 30-50'C. Aquatic forms include planktonic as well
orders, traditionally referred to collectively as the asbenthic speciesat all depths.The group includes the
Thalassinidea,have recentlybeen recognizedas dis- most successfulterrestrial crustaceans-the pillbugs
tinct. The vernacular term "thalassinid" is still some- and sowbugs of the order Isopoda-and a few amphi-
times used to refer to them together. The mud and pods that have invaded land and live in damp forest
ghost shrimps are particularly difficult to placewithin leaf litter or gardens.Peracaridsrange in sizefrom tiny
the decapods.Sometimesthey are depicted as related interstitial forms only a few millimeters long to plank-
to the crayfish and chelatelobsters(Astacidea),and tonic amphipods over 12 cm long (Cystisoma), deepsea
sometimesthey are grouped with the hermit crabs necrophagousamphipods exceeding34 cm (Alicellagi-
and their relatives (Anomura). Thesedecapodshave a gantea),and benthic isopodsgrowing to 50 cm in length
symmetrical abdomen that is flattened dorsoventrally (Bathynomus giganteus).Theseanimals exhibit all sorts
and extendsposteriorly as a well-developedtailfan. of feeding strategies;a number of them, especiallyiso-
The carapaceis somewhatcompressedlaterally,and pods and amphipods,are commensalsor parasites.
the gills are trichobranchiate.The first 2pairs of pereo-
pods are chelate,and the first pair is generally much Order Mysida Carapacewell developed,covering
enlarged.Most of theseanimals are marine burrowers most of thorax, but never fused with more than four
or live in coral rubble. They generally have a rather anterior thoracic segments;maxillipeds (1-2 pairs) not
thin, lightly sclerotizedcuticle,but some (e.g.,mem- associatedwith cephalic appendages;thoracomere
bers of the family Axiidae) have thicker skeletonsand 1 separatedfrom head by internal skeletalbar; abdo-
are more lobster-like in appearance.Gebiideans(par- men with well developed tail fan; pereopods bira-
ticularly Upogebia, Callinnassa,
and relatedgenera)often mous/ except last pair, which are sometimesreduced;
occur in huge colonies on tidal flats, where their bur- pleopods reducedor, in males,modified; compound
row holes form characteristicpatterns on the sediment eyesstalked, sometimesreduced;gills absent;usually
surface(Figures21.1Land 21,.11, A). with a statocystin eachuropodal endopod; adults with
antennalglands(Figures21.72A,8,21.308, and 21.33C).
lnfraordersGlypheideaand Polychelida The gly- There are more than 1,050speciesof mysids, rang-
pheids are something of a relict group, representedby ing in length from about 2 mm to 8 cm. Most swim by
two living genera (Neoglypheaand Laurentaglyphea), action of the thoracic exopods.Mysids are shrimplike
each with a single species,of a formerly diverse crustaceansthat are often confused with the superfi-
group known from the fossil record. Polychelidsare cially similar euphausids(which lack oostegitei and
a small group of blind deep-sealobsters,notable for uropodal statocysts).Mysids are pelagic or demer-
having chelaeon all of their pereopods and unusual, sal and are known from all oceandepths; a few spe-
large, globatelarvae (called eryoneicuslarvae) unique cies occur in freshwater.Somespeciesare intertidal
among the decapods. and burrow in the sand during low tides. Most are
omnivorous suspensionfeeders,eating algae,zoo-
Superorder Peracarida Telsonwithout caudal planktorg and suspendeddetritus. In the past, mysids
rami; 1 (rarely 2-3) pair of maxillipeds; maxilliped were combined with lophogastrids and the extinct
basis typically produced into an anteriorly directed, Pygocephalomorphaasthe "Mysidacea."
bladelike endite; mandibles with articulated acces-
sory processesin adults, between molar and incisor Order Lophogastrida Similar to mysids, exceptfor
processes/called the lacinia mobilis; carapace,when the following: maxillipeds (1 pair) are associated^with
present, not fused with posterior pereonitesand usu- the cephalic appendages;thoracomere 1 not separat-
ally reduced insize; gills thoracic or abdominal; with ed from head by internal skeletalbar; pleopods well
unique, thinly flattened thoracic coxal endites, called developed; gills present; adults with both antennal
oostegites,that form a ventral brood pouch or mar- and maxillary glands;without statocysts;all 7 pairs of
supium in femalesall speciesexceptmembersof the pereopodswell developed and similar (exceptamong
order Thermosbaenacea(the latter using the cara- members of the family Eucopiidae, in which their
pace to brood embryos);young hatch as mancas,a structure varies) (Figures 21.I2C,D and 21.21G).
(B) Compound
Carapace
Antennule
Antennal
scale
Antenna
Mandible
Pleopods
(c) Statocyst
Pereopods
Roshum Antennule
Telson Uropod
Coxa
Mandible
Exopod
Endopod
Antenna
Pereopods
Antennule
\ /
-Pseudorostrum
Figure 21 .12 Anatomy and diversity in some peracarid crus-

taceans (Eumalacostraca; Peracarida)-mysids, lophogastrids,
cumaceans, and tanaids. (A) A mysid, Bowmanella braziliensis.
(B) Anatomy of a generalized mysid (Mysida). (C) Anatomy of a
lophogastrid, Gnathophausla. (D) Second pereopod of Gnatho-
phausia. (E) A cumacean, Diastylis, in its typical partially buried
position. The arrows indicate the feeding and ventilation current.
(F) A cumacean. (G) A tanaid. (H) Anatomy of a generalized tanaid.
(H)
Antennule
Antenna
Pleopods
Gnathopod Pereopods
(secondthoracopods)
There are about 60 known speciesof lophogastrids, two speciesof unusual crustaceansMictocarishalope
most of which are1-8 cm long, although the $antGnatho- (from marine cavesin Bermuda) and Hirsutia bathya-
phausiaingensreaches35 cm. All arepelagicswinuners, lls (from a benthic sample 1,000m deep in the Guyana
and the group has a cosmopolitanoceanicdistribution. Basin off northeasternSouth America). A third spe-
Lophogastridsareprimarily predatorson zooplankton. ciesof Mictaceawas describedin 1988from Australia,
and a fourth from the Bahamasrn7992;there are now
Order Cumacea Carapacepresen! coveringand fused 6 speciesknown. Mictaceansare small,2-3.5 mm in
to first three thoracic segments,whose appendagesare length.Mictocarishalopeis the best known of thesespe-
modified as maxillipeds, the first with modified bran- ciesbecausemany specimenshave beenrecoveredand
chial apparatusassociatedwithbranchial cavity formed somehave been studied alive. It is pelagic in cavewa-
by carapace;pereopods1-5 ambulatory,simple, 1-4 ters and swims by using its pereopodal exopods.The
may be biramous; pleopods usually absentin females statusof the Mictaceaas a monophyletic grouping and
and presentin males;telson sometimesfused with sixth its relationships to other peracarid orders is a subject
pleonite, forming pleotelson;uropods styliform; com- of ongoing debate.Someworkers recognize the fam-
pound eyesabsent,or sessileand usually fused (Figures ily Hirsutiidae (containingHirsutia,Montucaris,and
21.7Pand21.12E,F). Thetispelecaris) as a separateorder, the Bochusacea
Cumaceansare small, odd{ooking crustaceanswith (malepleopodsbiramous).
a large,bulbous anterior end and a long, slenderposte-
rior-resembling horizontal commas!The greatcarcin- Order Spelaeogriphacea Carapaceshort, fused with
ologist Waldo Schmitt referred to them as "little won- first thoracomere;1 pair of maxillipeds; pereopods1-Z
ders and queer blunders." They occur worldwide and simple,biramous,with shortenedexopods;exopods
include about 1,500species,most of which arebetween on legs 1-3 modified for producing currents, on legs
0.1 and 2 cm long, though some speciesin cold water 4-7 as gills; pleopods 1-4 biramous, natatory; pleopod
reach 3 cm in length. Most are marine, although a few 5 reduced; tail fan well developed; compound eyes
brackish-waterspeciesare known. They live in associa- nonfunctional or absen! but eyestalkspersist (Figures
tion with bottom sediments,but are capableof swim- 21.13Aand 21.21H).The order Spelaeogriphacea is cur-
ming and probably leavethe bottom to breed.Most are rently known from only four living species.Theserare,
deposit feedersor predators on the meiofauna,others small (lessthan 1 cm) peracarids were long known
eat the organic film on sand grains. only from a single speciesliving in a freshwater stream
in Bat Cave on TableMountain, South Africa. A sec-
Order Tanaidacea Carapacepresentand fused with ond speciesis known from a freshwatercavein Brazil,
first two thoracicsegments;thoracopods1.-2aremaxil- and a third and fourth specieswere describedfrom an
lipeds, the secondbeing chelate;thoracopods 3-8 are aquifer in Australia. Little is known about the biology
simple, ambulatory pereopods;pleopods presentor of theseanimals,but they are suspectedto be detritus
absent;uropods biramous or uniramous; telson and feeders.Like thermosbaenaceans, spelaeogriphaceans
last one or two pleonites fused as pleotelson;adults are thought to be relictsof a more widespreadshallow-
with maxillary and (vestigial) antennal glands; com- water marine Tethyan fauna stranded in interstitial and
pound eyes absent,or present and on cephalic lobes. ground-water environmentsduring periods of marine
Membersof this order areknown worldwide from ben- regression.
thic marine habitats; a few live in brackish or nearly
fresh water. Most of the 1,500or so speciesare small, OrderThermosbaenacea Carapacepresent,fused
ranging from 0.5 to 2 cm in length. They often live in with first thoracomereand extending back over 2-3
burrows or tubesand areknown from all oceandepths. additional segments;1 pair of maxillipeds; pereopods
Many are suspensionfeeders,others are detritivores, biramous,simple,lacking epipodsand oostegites;cara-
and still othersarepredators(Figure 21.12G,H). pace forms dorsal brood pouch (unlike all other pera-
carids,which form the brood pouch from ventral ooste-
Order Mictacea Without a carapace, but with a well gites);2 pairs of uniramous pleopods;uropods bira-
developed head shield fused with first thoracomere mous; telson free or forming pleotelson with last ple-
and produced laterally over basesof mouthparts;1 pair onite; eyesabsent(Figure 21.138,C).About 34 species
of maxillipeds; pereopodssimple, 1-5 or 2-6biramous, of thermosbaenaceans are recognizedin sevengenera.
exopodsnatatory; gills absent;pleopods reduced,uni- Thermosbaena mirabilisis known from freshwaterhot
ramous or biramous; uropods biramous, with 2-5 seg- springs in North Africa, where it lives at temperatures
mented rami; telson not fused with pleonites; stalked in excessof 40'C. Severalspeciesin other generaoccur
eyespresent(Mictocaris)but lacking any evidenceof in much cooler fresh waters, typically in groundwater
visual elements,or absent(Hirsutin)(Figure21.13D-E). or in caves.Other speciesare marine or inhabit under-
Mictaceais the most recently (1985)establishedper- ground anchialine pools. Limited data suggestthat
acaridanorder. The order was erectedto accommodate thermosbaenaceans feed on plant detritus.
Ocular lobe
(A) Antennule
Pereopods 1-7
Carapace
Pleopods (B) Thoracomeres3-8

Antennule
Pereopods1-7
Endopod
Pleopods
Antennule
Exopod Endopod
LPereopod 5r
Figure 21 .13 More peracarids. (4 The spelaeogriphacean,

Spelaeogriphus. (B) The thermosbaenacean, Monodella. (C'1A
thermosbaenacean. (D,E) The mictacean, Mictocaris.
pleopods biramous and well developed, natatory and

Order lsopoda Carapaceabsent;first thoracomere for gas exchange (functioning as gills in aquatic taxa,
fused with head; 1 pair of maxillipeds;7 pairs of uni- and with air sacs called pseudotrachea in most ter-
ramous pereopods, the first of which is sometimes restrial Oniscidea);adults with maxillary and (ves-
subchelate,othersusually simple (gnathiidshave only tigial) antennal glands; telson fused with one to six
five pairs of pereopods,as thoracopod2 is a maxilli- pleonites,forming pleotelson;eyesusually sessileand
pedal "pylopod" and thoracopod8 is missing);pereo- compound, absentfrom some,pedunculate in most
pods variable, modified as ambulatory, prehensile,or Gnathiidea;with biphasic molting (posterior region
swimming; in the more derived suborderspereopodal molts before anterior region) (Figures 27.7Q,2L.L4,
coxaeare expandedas lateral side plates (coxalplates); 2I.21I,2L.28H,I,and 21.33M).
PHYLUM ARTHROPODA Crustacea:Crabs,Shrimps,and TheirKin 787
Antennule (E
Compound eye
Antenna
Pereonite one
(= thoracomere two)
Coxal plates
(E)
Pleotelson
Protopod
-l
Exopod UroPod
Endopod
_l
(G)
Figure 21 .14 More peracarids:

members of the order lsopoda.
(A) A flabelliferan, Excorallana
(family Corallanidae). (B) A fla-
belliferan, Paracerceis (family
Sphaeromatidae). Male mem-
bers of this genus possess
greatly enlarged uropods.
(C) A flabelliferan, Codonophilus
(family Cymothoidae). Members
of this genus are parasites that
attach to the tongues of various marine fishes. (D) A fla-
The isopods comprise about 10,500marine, freshwa- belliferan, Heteroserolis (family Serolidae). (E) A valviferan,
ter, and terrestrial species, ranging in length from 0.5 to ldotea (tamily ldoteidae). (F) An asellote, Joeropsis (tam-
ily Joeropsididae). (G) An anthurid, Mesanthura (family
500 mm, the largest being species of the benthic genus
Anthuridae). (H) A gnathiidean, Gnathia (family Gnathiidae).
Bathynomus(Cirolanidae). They are common inhabit- Note the grossly enlarged mandibles characteristic of
ants of nearly all environmenti, and some groups are male Gnathiidea. (l) An oniscidean, Ligia (the common
exclusively (e.9., Bopyridae, Cymothoidae) or partly seashore "rock louse").
788 Chapter TwentY-One
(A)
Key
$-ffi utoto-"
ffi H"o.t
ffi Coxalplates
P"."o"
F,--II] Antennule
Gnathopods
! lleopads
Pereopods Anterrrule
Sessile
(c) Lateral gili
Pereopod3
Medial gill
Pereopods
Figure 21 .15 And still more peracarids: amphipod diversity'

(A) General anatomy of a gammaridean amphipod. (B) General anat-
omy of a hyperiidean amphipod. (C) General anatomy of a cyamid
amphipod (Cyamus monodontis). (D) A gammaridean, Melita' (E) A
caprellid amphipod. (F) A cyamid amphipod, Cyamus erraticus, para-
sitic on right whales. (G,H) Two gammaridean amphipods: (G) Hyale,
a beach hopper, and (H) Heterophlias, an unusual, dorsoventrally
flattened species. (l-K) Three hyperiidean amphipods: (l) Primno'
(J) Leptocottis, and (K) a hyperiid on its host medusa' (L) A free-living
caprellid, Capretta. (M) An ingolfiellid amphipod, lngolfiella'
PHYLUMARTHROPODA Crustacea:Crabs, Shrimps,and Their Kin 789
(L)
(e.9.,Gnathiidae)parasitic.The suborderOniscideain- Order Amphipoda Carapaceabsent;first thoraco-

cludesabout 5,000speciesthat have invaded land (pill- mere fused to head; 1,pair of maxillipeds; 7 pairs of
bugs and sowbugs);they are the most successfulterres- uniramous pereopods,with first, second,and some-
trial crustaceans.Their direct development, flattened times others frequently modified as chelae or sub-
shape,osmoregulatory capabilities,thickened cuticle, chelae;pereopodalcoxaeexpanded as lateral side
and aerial gasexchangeorgans (pseudotrachea)allow plates (coxalplates);gills thoracic (medial pereopo-
most oniscideansto live completely divorced from dal epipods);adults with antennalglands; abdomen
aquatic environments. Fossilsas old as 325 million "divided" into fwo regions of three segmentseach,an
years(Carboniferous)have beenreported. anterior "pleon" and posterior urosome,with anterior
Isopod feeding habits are extremely diverse.Many appendagesastypical pleopodsand urosomalappend-
are herbivorous or omnivorous scavengers,but direct agesmodified asuropods;telsonfreeor fused with last
plant feeders,detritivores, and predatorsare also com- urosomite; other urosomitessometimesfused; com-
mon. Some are parasites(e.g.,on fishes or on other pound eyessessile,absentin some,huge in many (but
crustaceans)that feed on the tissuefluids of their hosts. not all) members of the suborder Hyperiidea (Figures
Overall, grinding mandibles and herbivory seemto 21.1NO, 21.15,21.23,21.27F,and 21.29D).
represent the primitive state,with slicing or piercing Isopods and amphipods sharemany featuresand
mandibles and predation appearing later in the evolu- are often said to be closely related. Earlier workers
tion of severalisopod clades. recognizedthesesimilarities (e.g.,sessilecompound
eyes, loss of carapace, and presence of coxal plates) contains only about 40 species, most of which live in
and classified them together as the "Edriopthalma" subterranean fresh and brackish waters, although a few
or "Acarida." Flowever, recent work suggests that are marine and interstitial. Little is known about their
many similarities between these two taxa may be con- biology. The 300 or so species of caprellid amphipods
vergences or parallelisms. The roughly 11,000species ("skeleton shrimp") are highly modified for clinging to
of amphipods range inlength from tiny 1mm forms other organisms, including filamentous algae and hy-
to giant deep-sea'benthic species reaching 29 cm, and droids. In most species the body and appendages are
a group of planktonic forms exceeds 10 cm. They very narrow and elongated. In one family of caprellids,
have invaded most marine and freshwater habitats the Cyamidae (with 28 species),individuals are obligate
and often constitute a large portion of the biomass in symbionts on cetaceans(whales, dolphins, and porpois-
many areas. The largest known amphipod is Alicelkt es) and have flattened bodies and prehensile legs.
gigantea, a cosmopolitan marine species living at In addition to parasitism, amphipods exhibit a vast
depths up to 7,000m. array of feeding strategies, including scavenging, her-
The principal suborder is Gammaridea. A few gam- bivory, carnivory, and suspension feeding.
marideans are semiterrestrial in moist forest leaf litter
or on supralittoral sandy beaches (e.9., beach hop- " Maxi l l opodans" The fol l ow i ng se ven classes-
pers); a few others live in moist gardens and green- Thecostraca, Tantulocarida, Branchiura, Pentastomida,
houses (e.g.,Talitrus sylaaticus and T. pncificus).They Mystacocarida, Copepoda, and Ostracoda-histori-
are common in subterranean groundwater ecosystems cally were united in a group called the Maxillopoda
of caves, the majority being stygobionts-obligatory that is now known to be artificial (nonmonophyletic).
groundwater species characterized by reduction or loss Howeveq, many of these "maxillopodan" groups share
of eyes, pigmentation, and occasionally appendages. some basic characteristics. These include a body of five
About 900 species of stygobiontic amphipods have cephalic, six thoracic, and four abdominal somites, plus
been described, including the diverse genercNiphargus a telson, but reductions of this basic 5-6-4body plan are
(in Europe) and Stygobromus(in North America), each common, and different specialists sometimes interpret
with over 100 described species. However, most of the the nature of these tagmata in different ways, leading
gammaridean amphipods are marine benthic species, to some confusion. Additional characters that most of
and a few have adopted a pelagic lifestyle, usually in these classeshave in common are: thoracomeres vari-
deep oceanic waters. There are many intertidal spe- ously fused with cephalon; usually with caudal rami;
cies, and a great many of these live in association with thoracic segments with biramous (sometimes unira-
other invertebrates and with algae. Domicolous gam- mous) limbs,lacking epipods (except in many ostra-
maridean amphipods in at least three families spin silk cods); abdominal segments lack typical appendages;
from their legs that is used for consolidating the walls carapace present or reduced; with both simple and
of their tube or shelter. compound eyes, the latter being unique, with three
The suborder Hyperiidea includes exclusively pe- cups, each with tapetal cells (an affangement still often
lagic amphipods that have apparently escaped the referred to as the maxillopodan eye).
confines of benthic life by becoming associated with Most of these former "maxillopodans" are small
other plankters, particularly gelatinous zooplankton crustaceans, barnacles being a notable exception. They
such as medusae, ctenophores, and salps. The hype- are generally recognizable by their shortened bodies,
riideans are usually characterized by huge eyes (and especially the reduced abdomen, and by the absence
a few other inconsistent features), but several groups of a full complement of legs. The reductions in body
bear eyes no larger than those of most gammarideans. size and leg number, emphasis on the naupliar eye,
The Hyperiidea are almost certainly a polyphyletic minimal appendage specialization, and certain other
group, and it is thought that several lineages are de- features have led biologists to hypothesize that pae-
rived independently from various gammaridean an- domorphosis (progenesis) played a role in the origin
cestors, although a modern phylogenetic analysis has of some of these classes.That is, in many ways, they
yet to be attempted. The precise nature of the relation- resemble early postlarval forms that evolved sexual
ships between hyperiideans and their zooplankton maturity before attaining all the adult features. Over
hosts remains controversial. Some appear to eat host 26,000 species have been described within the seven
tissue, others may kill the host to fashion a floating classes.
"home," and still others may utilize the host merely
for transport or as a nursery for newly hatched young. Class Thecostraca
Specimens of the scyphozoarr Phacellophoracamtschtica This group includes the bamacles, parasitic ascothorac-
have been found with nearly 500 Hyperia medusarum ids, and mysterious "y-lalae." The thecostracanclade
riding (and feeding) on it. is defined by several rather subtle synapomorphies of
There are two other small amphipod suborders: cuticular fine structure, including cephalic chemosen-
Ingolfiellidea and Caprellidea. The first suborder sory structures known as lattice organs. The group is
also supportedby molecularphylogeneticanalyses.All crystalline cone structure in the compound eye, a fea-
taxa have pelagic larvae, the terminal instar of which ture not known from any other crustaceangroup and
possessesprehensileantennulesand is specializedfor perhaps a vestigeof the ancestralthecostracanbody
locating and attaching to the substratum of the sessile plan. Most speciesof barnaclesare hermaphrodites,
adult state. whereasseparatesexesare the rule in acrothoracicans
and rhizocephalans,and someandrogonochoristicspe-
Subclass Ascothiracidd About 125describedspe- cies (males+ hermaphrodites;e.g.,Scalpellum)have
cies of parasitesof anthozoansand echinoderms. alsobeenreported.
Although greatly modified, they retain a bivalved Locomotion in barnaclesis generally confined to
carapaceand the full complement of thoracic and the larval stages,although adults of a few speciesare
abdominal segments(factsthat suggestthey might be specifically adapted to live attached to floating ob-
the most primitive living thecostracans).
Ascothoracids jects (e.9.,seaweeds,pumice, logs) or nektonic marine
generally have mouthparts modified for piercing and animals (e.g.,whales, sea turtles). Others are often
sucking body fluids, but some live inside other ani- found on the shells and exoskeletonsof various er-
mals and absorbthe host's tissuefluids. In at leastone rant invertebrates(e.9.,crabsand gastropods),which
species,Synagoga mira, malesretain the ability to swim inadvertently provide a meansof transportation from
throughout their lives, attaching only temporarily one place to another.Thoracicanand acrothoraci-
while feeding on corals(Figure21.16F). can barnaclesuse their feathery thoracopods (cirri) to
suspensionfeed. Barnaclesin the family Coronulidae
Subclass Cirripedia Primitively with tagmataas in are suspensionfeedersthat attach to whales and tur-
the class,but in most groups the adultbody is modified tles (e.g., Chelonibia,Platylepas,St omatoIepas,Coronula,
for sessileor parasitic life; thorax of six segmentswith Xenobqlanus). The 265 or so known speciesof rhizo-
paired biramous appendages;abdomenwithout limbs; cephalansare all endoparasitic in other Crustacea,
telson absentin most, although caudal rami persist on and are the most highly modified of all cirripedes.
abdomen in some;nauplius larva with frontolateral They mainly inhabit decapodcrustaceans,but a few
horns; unique, "bivalved" cypris larva; adult carapace are known from isopods/ cumaceans,stomatopods,
"bivalved" (folded) or forming fleshy mantle; first tho- and even thoracican barnacles.Someeven parasitize
racomereoften fused with cephalonand bearing max- freshwaterand terrestrialcrabs.The body consistsof
illiped-like oral appendages;female gonoporesnear a reproductive part (the externa)positioned outside
basesof first thoraciclimbs, male gonoporeon median the host's body, and an internal, ramifying, nutrient-
penis on last thoracicor first abdominal segment;com- absorbingpart (the interna).6
pound eyeslost in adults (Figures21.15,T,21.16A-E,
21.25,2L.26,21..278,C, 21.32E,and 21.33F). Subcfass Facetotecta Monogeneric(Hansenocaris):
The 1,285or so describedcirripede speciesaremost- The "y-larvae," a half-dozen small (250-620mm)
ly free-living barnacles,but this group also includes marine nauplii and cyprids (Figure 21.16I).Although
some strange parasitic "barrlacles" rarely seenexcept known sinceHansen'soriginal description in 1899,the
by specialists.The common acorn and goosebarnacles adult stageof these animals has still not been identi-
belong to the superorder Thoracica.The superorder fied. However, a stage subsequentto the y-larva, the
Acrothoracica consistsof minute animals that burrow slug-like ypsigon stage,has been induced by treat-
into calcareoussubstrata,including coralsand mollusc ing y-larvae with molting hormones.The prehensile
shells (Figure21.16G).The superorderRhizocephala antennulesand hooked labrum of the y-cyprids and
are exceptionallymodified parasitesof other crusta- the degenerativenature of the ypsigon suggestthat the
ceans/especiallydecapods(Figure 21.16H). adults areparasiticin yet-to-be-identifiedhosts.
If the body plan of the cirripedes is derived from
something similar to what is seenin other "maxillopo- Class Tantulocarida
dan" classes,then it has been so extensivelymodified Bizarceparasitesof deep-watercrustaceans.juveniles
that its basic features are nearly unrecognizablein with cephalon,6-segmented thorax, and abdomenof
adults of this subclass.The abdomenis greatly reduced up to7 segments;cephalonlacking appendages(other
in adults, and also in most cypris larvae. In cyprids than antennulesin one known stageonly) but with an
(cypris larvae) the carapaceis always present and "bi- internal median stylet; thoracopods1-5 biramous, 6
vaIved," the two sidesbeing held by a transversecypris
adductor muscle;in adults the carapacemay be lost oRhizocephalans of the family Sacculinidae
(Rhizocephala)or modified as a membranous,saclike infest only decapod
crustaceans and have been suggested as biological control agents
mantle (thoracicansand acrothoracicans). In the bar- for invasive exotics such as the green crab (Carcinus maenas),
nacles(Thoracica),it is this mantle that producesthe fa- which are upsetting coastal ecosystems worldwide. Sacculinds
have the ability to take control over such maior host functions
miliar calcareousplatesthat enclosethe body. Cyprids as molting and reproduction, and also to compromise the host's
and adult acrothoracicansshare a unique tripartite lmmune system.
(A) E
Mantle
opening
Attached
male
Maxilla
{ Figure 21 .16 Anatomy and diversity in the class The tiny tantulocarids are less than 0.5 mm long.
Thecostraca-barnacles and their kin. (A-E) Thoracican They attach to their hosts by penetrating the body with
barnacles. (A) A sessile (acorn) barnacle with its cirri
extended for feeding. (B) Plate terminology in a balano-
a protruding cephalicstylet. The young bear natatory
morph (acorn) barnacle. (C) The lepadomorph (stalked) thoracopods.About three dozen speciesin 22 genera
barnacle Pollicipes polymerus. (D) Verruca, the "wart" bar- have been described(Figures21.1Vand 21.77).They
nacle. (E) Two thorapican barnacles that live in association are known from abyssaldepths to the intertidal zone,
with each other and'with whales. The stalked barnacle from polar to tropical waters, and from anchialhe pools
Conchoderma attaches to the sessile barnacle Coronula, and hydrothermal vents, always as parasiteson other
which in turn attaches to the skin of certain whales. (F)
crustaceans.Until recently, members of this group had
The ascothoracican Ascothorax ophiocentenis, a para-
site that feeds periodically on echinoderms (longitudinal
beenassignedto various parasiticgroups of Copepoda
section). (G) An acrothoracican, Alcippe. Note the highly and Cirripedia.In 1983GeoffreyBoxshalland Roger
modified female and the tiny attached male. This species Lincoln proposed the new classTantulocarida. Some
bores into calcareous substrata such as coral skeletons. early work on the group supported a view of theseani-
(H) A crab (Carcinus) infected with the rhizocephalan mals asmaxillopodans,although the presenceof six or
Sacculina carcini. The crab's right side is shown as trans- sevenabdominal segmentsin juveniles of somespecies
parent, exposing the ramifying body of the parasite. (l) A
is inconsistent with this view; they are now believed to
cypris y-larva, in lateral and dorsal views.
be allied to the Thecostraca.The life rycle is unique and
includes a larva called the tantulus.
uniramous; abdomen without appendagesbut with Class Branchiura

caudalrami; adults highlymodified, with "unsegment- Body compact and oval, head and most of trunk cov-
ed" sacciformthorax and a reduced abdomenbearing ered by broad carapace;antennulesand antennaere-
a uniramous penis on the first segment;female gono- duced, the latter sometimesabsent;mouthparts modi-
pores on fifth thoracic segment. fied for parasitism; no maxillipeds; thorax reduced
Figure 21 .17 Anatomy in the

class Tantulocarida. (A) An adult
Basipodella atlantica. Note the
absence of an abdomen and the
modifications for parasitic life.
(B) Basipodella attached to the
antenna of a copepod host.
(C,D) Microdajus pectinatus on
a crustacean host, adult, and
juvenile (SEM).
(A)
Antennule
Antenna
Mandible
Maxillule
Maxilla
Maxilliped Third thoracic
appendage
First abdominal
segment
udal ramus
Figure 21 .18 Anatomy of the classes Mystacocarida,

Copepoda, and Branchiura. (A) General anatomy and SEM
of the mystacocarid Derocheilocaris. (B) General anatomy of
Antennule a cyclopoid copepod. (C-E) General body forms of (C) a cala-
noid, (D) a harpacticoid, and (E) a cyclopoid copepod. Note
the points of body articulation (dark band) and the position
of the genital segment (shaded segment). Roman numerals
are thoracic segments; Arabic numerals are abdominal seg-
ments; T = telson. (F) An elaborately setose calanoid copepod
adapted for flotation. (G) A poecilostomatid copepod, Ergasilus
pitalicus, ectoparasitic on cichlid fishes. (H) A female sipho-
nostomatid copepod (Caligus sp.) with egg sacs. (l) A female
siphonostomatid copepod (Trebius heterodont, a parasite of
horn sharks in California) with egg sacs. (J) A siphonostomatid
copepod, Clavella adunca, showing extreme body reduction;
this species attaches to the gills of fishes by its elongate max-
illae. (K) Notodelphys, a wormlike cyclopoid copepod adapted
for endoparasitism in tunicates. (L) Argulus foliaceus, a
branchiuran that parasitizes fishes. Note the powerful hooked
suckers (modified maxillules) on the ventral surface.
midventral surfaceof last thoracic somite;paired, ses-

to four segments, with paired biramous appendages; sile compound eyes and one to three median simple
abdomen unsegmented, bilobed, limbless, but with eyes(Figure21.18L).
minute caudal rami; female gonopores at bases of The Branchiuracomprise about 230 speciesof ec-
fourth thoracic legs, male with single gonopore on toparasiteson marine and freshwater fishes. The
Maxillule (hookedsucker)
First
thoracic limb
(A) (B)
Hooks Claw glands
Penehation stvlet
Figure 21 .19 Anatomy and diversity in

the class Pentastomida. (A) Linguatula
se rrata. (B) Cep hal obae n a tet rapod a.
(C) lnternal anatomy of female Penta-
stomum. (D) Internal anatomy of female
Waddycephalus teretiusculus, a parasite
of Australian snakes. (E) A generalized
pentastomid primary larva.
antennules generally bear hooks or spines for attach- agesreduced to 2 pairs of head appendages,lobelike
ment to their host fish. The mandibles are reduced and with chitinous claws used to cling to host. Body
in size and complexity, bear cutting edges,and are cuticle nonchitinous and highly porous. Body muscles
housed within a styliform "proboscis" apparatus.The somewhat sheetlike,but clearly segmentaland cross-
maxillules are clawed in Dolops,but they are modi- striated. Mouth lacks jaws; often on end of snoutlike
fied as stalked suckersin the other genera(Argulus, projection;connectedto a muscular pumping pharlmx
Chonopeltis, Dipteropeltis).
The uniramous maxillaeusu- used to suck blood from host. The combination of the
ally bear attachmenthooks. The thoracopodsare bira- snout and the 2 pairs of legs gives the appearanceof
mous and used for swimming when the animal is not there being five mouths, hencethe name (Greekpenta,
attached to a host. Branchiurans feed by piercing the "fiye"; stomida,"mouths"). In many speciesthe ap-
skin of their hosts and sucking blood or tissue fluids. pendagesare reduced to no more than the terminal
Once they locate a host, they crawl toward the fish's claws. No specificgas exchange,circulatory, or excre-
head and anchorin a spot where water flow turbulence tory organs.Gonochoristic;femaleslarger than males.
is low (e.g.,behind a fin or gill operculum). About 130describedspecies,including two cosmopoli-
Members of the genus Argulus occur worldwide, tan speciesthat can occasionallyinfest humans (Figure
and can pose a seriousproblem to aquaculture,but 21,.1e).
members of the other generahave restricted distribu- For yearsit was believedthat pentastomidswere al-
tions. Chonopeltisis found only in Africa, Dipteropeltis lied with the fossil lobopodians, onychophorans,and
in SouthAmerica, and Dolopsin SouthAmerica,Africa, tardigradesas some kind of segmented,vermiform,
and Tasmania. proto-arthropod creature-and some workers still
entertain this idea. However, independent molecular
Class Pentastomida studies suggestthe pentastomidsare highly modified
Obligatory parasites of various amphibians, reptiles, crustaceans,perhaps derived from the Branchiura.
birds, and mammals. Adults inhabit respiratory tracts Corroboration has come from cladistic analysesof
(lungs, nasalpassages,etc.)of their hosts.Body highly sperm and larval morphology, nervous systemanato-
modified, wormlike, 2-73 cmin length. Adult append- my, and cuticular fine strucfure.
Work on the Swedish Orsten fauna indicates that There are more than 12,500 described species of
pentastomid-like animals had appeared as early as copepods. They can be incredibly abundant in the
the late Cambrian (500 Ma), long before the land ver- world's seas,and also in some lakes-by one estimate,
tebrates had evolved. \zVhatmight the original hosts of they outnumber all other multicellular forms of life
these parasites have been? Conodont fossils are com- on Earth. Most are small,0.5-10 mm long, but some
mon in all the Cambrian localities that have yielded free-living forms exceed 1.5 cm in length, and certain
pentastomids, raising the possibility that conodonts highly modified parasites may reach 25 cm. The bod-
(also long a mystery, but now widely regarded as parts ies of most copepods are distinctly divided into three
of early fishlike vertebrates) may have been at least one tagmata, the names of which vary among authors.
of the original hosts of these early Pentastomida. The first region includes the five fused head segments
and one or two additional fused thoracic somites; it
Class Mystacocarida is called a cephalosome (= ggphrlothorax) and bears
Body divided into cephalon and 1O-segmented trunk; the usual head appendages and maxillipeds. All of
telson with clawlike caudal rami; cephalon character- the other limbs arise on the remaining thoracic seg-
istically cleft; all cephalic appendages nearly identical, ments, which together constitute the metasome. The
antennae and mandibles biramous; antennules, max- abdomen, or urosome, bears no limbs. The append-
illules, and maxillae uniramous; first trunk segment age-bearing regions of the body (cephalosome and
bears maxillipeds but is not fused with cephalon; no metasome) are frequently collectively called the pro-
carapace; gonopores on fourth trunk segment; trunk some. The majority of the free-living copepods, and
segments 2-5 with short, single-segment appendages those most frequently encountered, belong to the
(Figure 21.i8A). orders Calanoida, Harpacticoida, and Cyclopoida,
There are only 13 described species of mystaco- although even some of these are parasitic. We focus
carids, eight in the genus Derocheilocqrisand five in here on these three groups and then briefly discuss
Ctenocheilocaris.Most are less than 0.5 mm long, al- some of the other, smaller orders and their modifica-
though D. ingensreaches 1 mm. The head is marked by tions for parasitism. The calanoids are characterized
a transverse "cephalic constriction" between the origins by a point of major body flexure between the meta-
of the first and second anterurae, perhaps a remnant of some and the urosome, marked by a distinct nar-
primitive head segmentation. In addition, the lack of fu- rowing of the body. They possess greatly elongate
sion of the cephalon and maxillipedal tmnk segment the antennules. Most of the calanoids are planktonic, and
simplicity of the mouth appendages, and other features as a group they are extremely important as primary
have led some workers to propose that the mystacoca- consumers in freshwater and marine food webs. The
rids are among the most primitive living crustaceans. point of body flexure in the orders Harpacticoida and
These attributes may, however, simply be related to a Cyclopoida is between the last two (fifth and sixth)
neotenic origin and specialization for interstitial habitats. metasomal segments. (Note: Some authors define
Mystacocarids are marine, interstitial crustaceans the urosome in harpacticoids and cyclopoids as that
that live in littoral and sublittoral sands throughout the region of the body posterior to this point of flexure.)
world's temperate and subtropical seas. Their rather Harpacticoids are generally rather vermiform, with
vermiform body and small size are clearly adaptations the posterior segments not much narrower than the
to life among sand grains. Mystacocarids are thought anterior; cyclopoids generally narrow abruptly at the
to feed by scraping organic material from the surfaces major body flexure. Both the antennules and the an-
of sand grains with their setose mouthparts. tennae are quite short in harpacticoids, but the latter
are moderately long in cyclopoids (although never
Class Copepoda as long as the antennules of calanoids). The anten-
Without a catapace, but with a well developed cephal- nae are uniramous in cyclopoids but biramous in the
ic shield; single, median, simple maxillopodan eye other two groups. Most harpacticoids are benthic,
(sometimes lacking); one or more thoracomeres fused and those that have adapted to a planktonic lifestyle
to head; thorax of six segments, the first always fused show modified body shapes. Harpacticoids occur in
to the head and with maxillipeds; abdomen of five seg- all aquatic environments; encystment is known to
ments, including anal somite (= telson); well developed occur in at least a few freshwater and marine species.
caudal rami; abdomen without appendages, except an Cyclopoids are known from fresh and salt water, and
occasional reduced pair on the first segment, associ- most are planktonic.
ated with the gonopores; point of main body flexure The nonparasitic copepods move by crawling or
varies among major groupsi antennules uniramous, swimming, using some or all of the thoracic limbs.
antennae uniramous or biramous ; 4-5 pafus of natatory Many of the planktonic forms have very setoseappend-
thoracopods, most locked together for swimming; pos- ages, offering a high resistance to sinking. Calanoids
terior thoracopods always biramous (Figures 27.7U, are predominantly planktonic feeders. Benthic harpac-
21.18B-K, 21.27A, 27.30D,and 21.33D). ticoids are often reported as detritus feeders, but many
feed predominantly on microorganisms living on the modified fifth limbs are in fact these appendages. The
surface of detritus or sediment particles (e.g., diatoms, trunk seldom shows external evidence of segmenta-
bacteria, and protists). tion, although all 11 postcephalic somites are discern-
Of the seven remaining orders, the Mormonilloida able in some taxa. The trunk limbs vary in structure
are planktonic; the Misophrioida are known from among taxa and on individuals. The third pair of trunk
deep-sea epibenthic habitats as well as anchialine caves limbs bears the gonopores and constitutes the so-called
in both the Pacific and Atlantic; and the Monstrilloida copulatory organ.
are planktonic as adults, but the larval stages are en- Ostracods are one of the most successful groups of
doparasites of certain gastropods, polychaetes, and crustaceans. They also have the best fossil record of any
occasionally echinoderms. Members of the orders arthropod group, dating from at least the Ordovician,
Poecilostomatoida and Siphonostomatoida are ex- and an estimated 65,000 fossil species have been de-
clusively parasitic and often have modified bodies. scribed. Most are benthic crawlers or burrowers, but
Siphonostomatoids are endo- or ectoparasites of vari- many have adopted a suspension-feeding planktonic
ous invertebrates as well as marine and freshwater lifesfyle, and a few are terrestrial in moist habitats. One
fishes; they are often very tiny and show a reduc- species is known to be parasitic on fish gills-Sheina
tion or loss of body segmentation. Poecilostomatoids orri (Myodocopida, Cypridinidae). Ostracods are
parasitize invertebrates and marine fishes, and may abundant worldwide in all aquatic environments and
also show a reduced number of body segments. The are known to depths of 7,000 m in the sea. Some are
Platycopioida are benthic forms known primarily from commensal on echinoderms or other crustaceans. A
marine cavesi the Gelyelloida are known only from few podocopans have invaded supralittoral sandy re,
European groundwaters. gions (members of the family Terrestricytheridae), and
members of several families inhabit terrestrial mosses
Class Ostracoda and humus. Two principal taxa (ranked as subclasses
Body segmentation reduced, trunk not clearly divided here) are recognized within the Ostracoda: Myodocopa
into thorax and abdomen, with 6 to 8 pairs of limbs and Podocopa.
(including the male copulatory limb); trunk with 1 to 3 Myodocopans are all marine. Most are benthic, but
pairs of limbs, variable in structure; caudal rami pres- the group also includes all of the marine planktonic
ent; gonopores on lobe anterior to caudal rami; cara- ostracods. The largest of all ostracods, the planktonic
pace bivalved, hinged dorsally and closed by a central Gigantocypris,is a member of this group. Myodocopans
adductor muscle, enclosing body and head; carapace include scavengers, detritus feeders, suspension
highly variable in shape and ornamentation, smooth feeders, and some predators. There are two orders:
or with various pits, ridges, spines, etc.; most with one Myodocopida and Halocyprida.
simple mediannaupliar eye (often called a "maxillopo- Podocopans include predominantly benthic forms;
dan eye") and sometimes weakly stalked compound although some are capable of temporary swimming,
eyes (in Myodocopida); adults with maxillary and (in none are fully planktonic. Their feeding methods in-
some) antennal glands; males with distinct copulatory clude suspension feeding, herbivory, and detritus
limbs; caudal rami (furca) present (Figure 27.20). feeding. The Podocopa are divided into three orders:
O s t r ac o d s i n c l u d e a b o u t 3 0 ,0 0 0 descri bed l i v- the exclusively marine Platycopida, the ubiquitous
ing species of small bivalved crustaceans, ranging Podocopida, and the Palaeocopida. The Palaeocopida
in length from 0.1 to 2.0 mm, although some giants were diverse and widespread in the Paleozoic, but
(e.g., Gigantocypris) reach 32 mm. They superficially are represented today only by the extremely rare
resemble clam shrimps in having the entire body en- Punciidae (known from a few living specimens, and
closed within the valves of the carapace. However, from dead valves dredged in the South Pacific).
ostracod valves lack the concentric growth rings of
clam shrimps, and there are major differences in the
appendages. The shell is usually penetrated by pores,
some bearing setae, and is shed with each molt. A good
The Crustacean Body Plan
deal of confusion exists about the nature of ostracod We realize that the above synopses are rather exten-
limbs, and homologies with other crustacean taxa (and sive, but the diversity of crustaceans demands empha-
even within the Ostracoda) are unclear-this confusion sis before we attempt to generalize about their biology.
is reflected in the variety of names applied by differ- The evolutionary success of crustaceans, like that of
ent authors. We have adopted terms here that allow the other arthropods, has been closely tied to modifica-
easiest comparison with other taxa. tions of the jointed exoskeleton and appendages, the
Ostracods possess the fewest limbs of any crusta- latter having an extensive range of modifications for a
cean class. The four or five head appendages are fol- great variety of functions.
lowed by 1-3 trunk appendages. Superficially, the (sec- The most basic crustacean body plan is a head
ond) maxillae appear to be absen| however, the highly (cephalon) followed by a long body (trunk) with many
(A) Figure 21 .2O Anatomy and diversity in the class

Ostracoda. (A) Anatomy ot Sclerocypris (Podocopa).
Median eye (B) Anatomy of Thaumatoconcha (Myodocopa).
Sixth limb
(walking leg)
Fifth limb (male)

(maxilliped/clasper)
Seventh limb

Figure 21 .20 (continued) Anatomy and diver-

sity in the class Ostracoda. (C) Internal view of
(C) Antennula 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
ostracod groups (scale bar = 1.0 mm). A: Vargula
(Myodocopa, Myodocopida). B: Eusarsiella
(Myodocopa, Myodocopida). C: Polycope
(Myodocopa, Halocyprida). D: Cytherelloidea
(Podocopa, Platycopida). Et Propontocypris
(Podocopa, Podocopida). F: Macrocypris (Podocopa,
Podocopida). Gi Saipanetta (Podocopa, Podocopida).
H: Neonesldea (Podocopa, Podocopida). l: Triebelina
(Podocopa, Podocopida). J: Candona (Podocopa,
Podocopida). K: llyocypris (Podocopa, Podocopida).
L'. Cyprinotus (Podocopa, Podocopida).
M: Potamocypris (Podocopa, Podocopida).
N: Hemicytherura (Podocopa, Podocopida).
O'.Acanthocythereis (Podocopa, Podocopida).
P: Celtia (Podocopa, Podocopida). Q: Limnocythere
(Podocopa, Podocopida). R: Sahn i cythere (Podocopa,
Podocopida). S'. Darwinula (Podocopa, Podocopida).
Maxillula Fifth limb
All arrows point anteriorlv.
PHYLUMARTHROPODACrustacea;Crabs,Shrimps,and Their Kin 801
similar appendages, as seen in the class Remipedia together termed the pereon. Each segment of the pe-
(Figures 21.1,A.and 21,.3D,E).In the other crustacean reon is called a pereonite (= pereomere), and their ap-
classes,however, various degrees of tagmosis occur/ pendages are called pereopods. The pereopods may
and the cephalon is typically followed by a trunk that be specialized for walking, swimming, gas exchange,
is divided into two distinct regions, a thorax and an feeding, and/or defense. Crustacean thoracic (and
abdomen. All cruslaceans possess,at least primitive- pleonal) appendages might be primitively biramous,
ly, a cephalic shield (head shield) or a carapace. The although the uniramous condition is seen in a variety
cephalic shield results from the fusion of the dorsal of taxa. The general crustacean limb is composed of
head tergites to form a solid cuticular plate, often with a basal protopod (= sympod), from which may arise
ventrolateral folds (pleural folds) on the sides. Head medial endites (e.g., gnathobases),lateral exites (e.g.,
shields are found in ancient Cambrian fossil crusta- epipods), and two rami, the endopod and exopod.
ceans (e.9., from the Orsten fauna), and they are char- Members of the classes Remipedia, Cephalocarida,
acteristic of the classes Remipedia and Cephalocarida; Branchiopoda, and some ostracods possess append-
they also occur in some former maxillopodan groups ages with uniarticulate (single-segment) protopods;
and in malacostracans. The carapace is a more expan- the remaining classesusually have appendages with
sive structure, composed of the head shield and a large multiarLiculate protopods (Table 21.1).8
fold of the exoskeleton that probably arises (primi- The abdomen, called a pleon in malacostracans, is
tively) from the maxillary somite. The carapace may composed of several segments, or pleonites (= pleo-
extend over the body dorsally and laterally as well as meres), followed by a postsegmental plate or lobe, the
posteriorly, and it often fuses to one or more thoracic anal somite or telson, bearing the anus (Figure 27.28).
segments, thereby producing a cephalothorax (Figure In primitive crustaceans this anal somite bears a pair
21,.2A).Occasionally, the carapace may grow forward of appendage-like or spinelike processes convention-
beyond the head as a narrow rostrum. ally called caudal rami. In the Eumalacostraca, the anal
Most of the differences among the major groups of somite lacks caudal rami and is followed by a dorsal
crustaceans, and the basis for much of their classifica- flattened cuticular flap; this flap is sometimes referred
tion, arise from variations in the number of somites in to as the telson.
the thorax and abdomery the form of their appendages, In general, distinctive abdominal appendages
and the size and shape of the carapace. A brief skim- (pleopods) occur only in the malacostracans. These
ming of the synopses (above) and the corresponding appendages are almost always biramous, and often
figures will give you some idea of the range of varia- they are flaplike and used for swimming (e.9., Figures
tion in these characteristics. 27.9-27.15). The posteriorly directed last pair(s) of ab-
Uniformity within the subphylum Crustacea is dem- dominal appendages is usually different from the other
onstrated particularly by the consistency of elements pleopods, and are called uropods. Together with the
of the cephalon and the presence of a nauplius larva. telson, the uropods form a distinct tail fan in many
Except for a few casesof secondary reductiory the head malacostracans (Figure 21,.28).
of all crustaceans has 5 pairs of appendages. From an- Crustaceans produce a characteristic, and unique,
terior to posterior, these are the antennules (first anten- larval stage called the nauplius (Figures 21.258,C and
nae), antennae (second antennae), mandibles, maxil- 21.33D), which bears a median simple (naupliar) eye
lules (first maxillae), and maxillae (second maxillae). and 3 pairs of setose, functional head appendages-
The presence of 2 pairs of antennae is, among arthro- destined to become the antennules, antennae, and
pods, unique to the Crustacea (as is the nauplius larva, mandibles. Behind the head segments is a growth-zone
although similar "head lalae" are known from other and the telson. In many groups (e.g., Peracarida, and
arthropod groups in the fossil record).7 Although the most of Decapoda), however, the free-living nauplius
eyes of some crustaceans are simple, most possess a larva is absent or suppressed. In such cases,develop-
pair of well developed compound eyes, either set di- ment is either fully direct or mixed, with larval hatch-
rectly on the head (sessile eyes) or borne on distinct ing taking place at some postnaupliar stage (Table
movable stalks (stalked eyes). 21..2).Oftenother larval stages follow the nauplius (or
In many crustaceans, fro- one to three anterior other hatching stage) as the individual passes through
thoracic segments (thoracomeres) are fused with a series of molts, during which segments and append-
the cephalon. The appendages of these fused seg- ages are gradually added. A recent compilation of all
ments are typically incorporated into the head as ad- crustacean larval forms (Martin et al. 2014) includes
ditional mouthparts called maxillipeds. In the class
Malacostraca, the remaining free thoracomeres are
8The term "peduncle" is a general name often applied to the basal
portion of certain appendages; it is occasionally (but not always)
'Cambrian crustacean-like larvae in the fossil record termed "head used in a way that is s;rnonymous with protopod. As noted in
larvae" may be precursors to the distinctive nauplius stage seen in Chapter 20, the exopod might be no more than a highly modified
many modern crustaceans. See Martin et al. (2074) exite that evolved from an ancestral uniramous condition.
802 ChapterTwentv-One
TABLE 21.1 Comparison of Distinguishing Features among the 11 Crustacean Classes

(and in Orders of the Class Branchiopoda and Subclasses of the Glass Eumalacostraca)
Body tagmata and
Carapace or number of segments in
cephalic shield each(excludingtelson) Thoracopods Maxi l l i pe ds
ClassRemipedia Cephalicshield Cephalon (6) Not phyllopodous 1 pair

Trunk (up to 32)
ClassCephalocarida Cephalic shield Cephalon (6)Thorax (B) Phyllopodous None

Abdomen (11)
ClassBranchiopod4 Cephalicshield Cephalon (6) D L .,1 1 ^ - ^ I^ ..^
r l yr u Pu u u u D None
OrderAnostraca Thorax (usually 11)
Abdomen (usually B)
ClassBranchiopod4 Carapace Cephalon (6), Phyllopodous None

Order Notostraca Thorax(11),Abdomen
{m r n \/ cp ( ) m p n fcl
ClassBranchiopod4 Carapace(bivalved, Cephalon 6,Trunk 10-32, Phyllopodous None

Order Diplostraca hinged or folded) or obscured)
ClassMalacostrac4 Large,folded carapace Cephalon(6),Thorax(8), Phyllopodous None

SubclassPhyllocarida coversthorax Abdomen (7)
ClassMalacostraca, Well developed Cephalon(6),Thorax(8), Not phyllopodous Five paris of thoracopods

SubclassHoplocarida caraPacecovers Abdomen (6) referred to as
thorax maxillipeds
ClassMalacostrac4 Carapacewell Cephalon (6),Thorax (8), Not phyllopodous; 0 to 3 pairs
Subclass dorrplnnorj nr Abdomen (6) uniramous in many
Eumalacostraca secondarilyreduced
or lost
ClassThecostraca Carapacebivalved (at Cephalon (6),Thorax (6), Not phyllopodous, often None
some stage),often Abdomen (4) reduced
modified as a mantle
ClassThntulocarida Cephaiic shield Cephalon (6),Thorax (6), Not phyliopodous, None

Abdomen (up to 7) qeeflrr redrrccd
ClassBranchiura Carapacebroad, Cephalon (6),Thorax (6), Not phyllopodous None

eorrerino head Abdomen (4?) /L,.r ^ll
\uuL 4rr l-^r^r^*.\
raraLvry,/
and trunk
ClassPentastomida None Not distinguishable;body None None
vermiform
ClassMystacocarida Cephalic shield Cephalon (6),Trunk (10) Not phyllopodous l pair
ClassCopepoda Cephalicshield Cephalon (6),Thorax (6), Not phyllopodous; Usually 1 pair

Abdomen (5) natatory,often reduced
ClassOstracoda Carapace,bivalved Subdivisionsnot clear;6-8 Not phyllopodous, None
pairs of limbs reduced
keys to the distinctive naupliar larvae in all groups that Swimming is usually accomplishedby a rowing ac-
hatch as a nauplius as well as synopses of larval devel- tion of the limbs. Archetypical swimming is exempli-
opment in all crustaceans. fied by crustaceanswith relatively undifferentiated
trunks and high numbers of similar biramous append-
Locomotion ages(e.g./remipedes,anostracans/notostracans).In
Crustaceans move about primarily by use of their general,theseanimals swim by posterior to anterior
limbs (Figure 21..21),and lateral body undulations are metachronalbeating of the trunk limbs (Figurc21.22
unknown. They crawl or swim/ or more rarely burrow, and Chapter 20).The appendagesof such crustaceans
"hitchhike," or jump. Many of the ectoparasitic forms are often broad and flattened,and they usually bear
(e.9.,branchiurans, certain isopods, and copepods) are fringes of setaethat increasethe effectivenessof the
largely sedentary on their hosts, and most cirripedes power stroke. On the recovery stroke the limbs are
are fully sessile. flexed,and the setaemay collapse,reducing resistance.
Compound Abdominal
Antennules Antennae eyes appendages Gonopore location
Biramous I Biramous Absent A,11

trunk appendages dr-protopodsof trunk segment15;
similar 9 +rrrnL cemenf R
Uniramous Biramous Absent None Common poreson protopodsof

lhnrarnnndc 6
Uniramous Uniramous Present None Q: on segment 1211,3

or 20121,
Uniramous Vestigial Present Present (posteriorly Thoracomere11 (both sexes)

reduced)
Uniramous Biramous Present A11trunk appendages Variable,on trunk segment 9 or 11,

similar, or posterior or on apodous posteriorregion
segmentslimbless (some Cladocera)
Biramous Uniramous Present Pleopods (posteriorly d: coxaeof thoracopods 8;

reduced) Q, co*aeof thoracopods 6
Tiiramous Biramous Present,well Well developed C/: coxaeof thoracopods 8;
developed pleopodswith gills; Q: coxaeof thoracopods 6
1 pair of uropods
Uniramous or Uniramous or Present,well Usually 5 paris of d: coxaeof thoracopods 8 or sternum

biramous biramous developed nlpnnndc 1 nair
of thoracomereB;
nf rrrnnorlc
"_ *_"r.-*" Q: coxaeof thoracopods 6 or sternum
of thoracomere 6
Uniramous Biramous Absent (in None Variable;Cf openingsusuallyon trunk
adults) selmPnt4orT:9nn14 or7
None (paired in one None Absent None Q openings on trunk segment 5

stage)
Uniramous, reduced Biramous,reduced Present None d^singleopeningon thoracomere6;

V gonoporesat baseof thoracopod4
None None Absent None Common gonopore,location variable
Uniramous Biramous Absent None C/ and ? openings on trunk segment 4
Uniramous Uniramous or Absent None Gonopores usually on urosomal

biramous segment1
Uniramous Biramous Absent None Gonoporeson lobe anteriorto
(typically) caudalrami
In members of some groups (e.9., Cephalocarida, to selected appendages (e.9.,the pleopods of shrimps,
Branchiopoda, Leptostraca), large exites or epipods stomatopods, amphipods, and isopods; the pereopods
arise from the base of the le& producing broad, "leafy" of euphausids and mysids). In swimming euphau-
limbs called phyllopodia. These flaplike structures aid sids and mysids the thoracopods beat in a metachro-
in locomotion and may also serve as osmoregulatory nal rowing fashion, with the exopod and setal fan ex-
(branchiopods) or gas exchange (cephalocardis and tended on the power stroke and flexed on the recovery
leptostracans) surfaces (Figure 27.27A-C). Although sfuoke. The movements and nervous-muscular coordi-
such epipods increase the surface area on the power nation of crustacean limbs are deceivingly complex. In
stroke, they also are hinged so that they collapse on the comrnon mysid Gnathophausiaingens, for example,
the recovery stroke, reducing resistance. Metachronal twelve separate muscles power the thoracic exopod
limb movements are retained in many of the "higher" alone (three that are extrinsic to the exopod, five in the
swimming crustaceans, but they tend to be restricted limb peduncle, and four in the exopodal flagellum).
(A)
Carpo-propodus
0)
Coxa -
{ Figure 21 .21 Generalized thoracic appendages of in the four primitive groups of crustaceans (cephalocarids,
various crustaceans. (A-C) Biramous, phyllopodous branchiopods, phyllocarids, and remipedes) the protopod
thoracopods. (A) Cephalocarida. (B) Branchiopoda. is composed of a single article. And in branchiopods and
Dashed lines indicate fold or "hinge" lines. (C) Leptostraca leptostracans, the articles of the endopod are not cleady
(Phyllocarida). (D) A biramous, flattened, but nonphyl- separated from one another. In the higher crustaceans
lopodous thoracopod: Remipedia. (E-l) Stenopodous (Malacostraca and former "maxillopodans") the protopod
thoracopods. (E) Eupfausiacea. (F) Caridea (Decapoda). comprises two or three separate articles, although in most
(G) Lophogastrida (P€racarida). (H) Spelaeogri phacea former maxillopodans these may be reduced and not eas-
(Peracarida). (l) lsopoda (Peracarida). Because of the pres- ily observed. In the lophogastrid (G), the large marsupial
ence of large epipods on the legs of the cephalocarids, oostegite characteristic of female peracarids is shown
branchiopods, and phyllocarids, some authors refer to arising from the coxa. In two groups (amphipods and
them as "triramous" appendages. However, smaller epi- isopods) all traces of the exopods have disappeared, and
pods also occur on many typical "biramous" legs, so this only the endopod remains as a long, powerful, uniramous
distinction seems unwarranted (and confusing). Note that walking leg.
(B)
Water in To head Median space
(midline)
Epipods/ Exopods
Water leaves
under epipods
Endite with
'filter" setae
g;5;Interlimb "filter" setae

@ sPace
(E)
\y
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 reduc-
ing 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 laterally, and the trapped food is moved
anteriorly to the mouth. (C-E) Locomotion in the postlarva 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 crus-
taceans with well developed abdomens and tail fans. (F) A swimming remipede,
Lasionectes. Note the metachronal waves of appendage movement.
TABLE21.2 Summary of CrustaceanReproductiveFeaturesa

Development type, or larval Hermaphroditic (at Parthenogenesis
type at hatching least some species) Gonochoristic (in at least some species)
ClassRemipedia Nauplius (anamorphic) Yes No No
ClassCephalocarida Metanauplius (anamorphic) Yes No No

-
ClassBranchiopod4 order Nauplius or metanauplius No Yes Yes

Anostraca /^- ^* ^--Li^\
\4r rau rurPr rrL./
ClassBranchiopod4 order Nauplius or metanauplius Yes Yes Yes

Notostraca (anamorphic),or direct
development
ClassBranchiopod4 order Nauplius (partly anamorphic) No Yes Yes
Diplostraca
ClassOstracoda Direct, orwith bivalved nauplius/ No Yes Yes

metanauplius with anamorphic
development
ClassMystacocarida Metanauplius (partiy anamorphic) No Yes No
ClassCopepoda Nauplius (metamorphic) No Yes No
ClassBranchiura Metanauplius-like (partly No Yes No

anamorphic), or with direct
development
CIassThecostraca ? Yes Yes No
ClassTantulocarida Nauplius? + tanfulus No Yes 2

(mefamnrnhic)
ClassMalacostraca,subclass Direct development No Yes No

Phyllocarida
ClassMalacostraca,subclass Zoealawa (" antizoea" or No Yes No

Hoplocarida "pseudozoea")(metamorphic)
ClassMalacostrac4 Direct development No Yes No

subclassEumalacostraca,
superorder S1'ncarida
ClassMalacostraca, Direct development No Yes No

subclassEumalacostrac4
superorderPeracarida
ClassMalacostrac4 Nauplius (metamorphic) No Yes No

subclassEumalacostrac4
superorderEucarid4 order
Frrnherrcienee
ClassMalacostraca, Amphion (modified zoea) (weakly No Yes No

subclassEumalacostrac4 m o +e m n r n h i n \
superorderEucarida,order
Amnhinnidacpr
ClassMalacostraca, Pre-zoeaor zoeatand with a Rare Yes No

subclassEumalacostraca, nauplius in Dendrobranchiata
superorderEucarida,order (metamorphic),or direct
Decapoda rlorrolnnnon+
oAlsoseeMartinet al.2014,Table55.1
PHYLUM ARTHROPODA Crustacea:Crabs,Shrimps,and Their Kin 807
Recallfrom our discussionsin Chapters4 and 20

Comments that at the low Relmolds numbers at which small
crustaceans(such as copepods or larvae) swim
Eight naupliar stages(2 orthonaupiii and 6 metanauplii), follo.rzedby
about, the netlike setalappendagesact not as a fil-
a"pre-juvenile"stage,havebeenreported.
tering net, but as a paddle, pushing water in front
One or tvvoeggs at a time are fertilized and carried on genital of them and dragging the surrounding water along
Drocesses of the firshoieonites.
with them due to the thick boundary layer adher-
E*bryor rrrrrulryrn"a t o- ovisacearly in developmen! resistant ing to the limb. Only in larger organisms,with
(cryptobiotic) fertilized eggsaccommodateunfavorableconditions.
Reynoldsnumbers approachingl,, do setoseap-
Eggs briefly brooded, then deposited on substrata;resistant pendages(e.9. the feeding cirri of barnacles)begin
(cryptobiotic) fertilized eggsaccommodateunfavorableconditions. to act as filters, or rakes,as the surrounding water
acts lessviscous and the boundary layer thinner.
Most cladoceransundergo direct development (Leptodorahatches Of course,the closertogetherthe setaeand setules
as nauplii or metanauplii). Clam shrimps carry developing are placed, the more likely it is that their individual
embryos on the thoracopodsprior to releasingthem as nauplii or
m ot r na, r nlii
boundary layers will overlap; thus densely setose
appendagesare more likely to act aspaddles.
Embryos usually deposited directly on substrata;many Not all swimming crustaceansmove by typical
myodocopansand some podocopansbrood embryosbetween
valves until hatching as a reduced adult; no metamolphosis; up to metachronalwaves of limb action. Certain plank-
8 preadult instars. tonic copepods,for example, move haltingly and
depend on their long antennulesand dense se-
Little studied, but apparently the eggsare laid free and up to 7
preadult stagesmay occur. tation for flotation between movements (Figure
21.18F).Watch living calanoidcopepodsand you
Usually with 6 naupliar stagesleading to a secondseriesot5"Iawal"
stagescalledcopepodites.
will notice that they may move slowly by use of
the antennaeand other appendages,or in short
Embryos are deposlted; only Argulus is known to hatch as jerky increments/ often sinking slightly between
metanauplii, others have direct development and hatch as
juveniles. thesemovements.The latter type of motion results
from an extremelyrapid and condensedmetachro-
a cypris
Six naupliar stagesfollowed by a unique iarval form ca11ed
nal wave of power strokes along the trunk limbs.
rarva.
Although the long antennaemay appearto be act-
A benthic non-feeding stage,possibly a nauplius, has been reported ing as paddles, they actually collapse against the
but not formally described. Development entails complex
metamorphosis with infective"tantulus larva."
body an instant prior to the beating of the limbs,
thus reducing resistanceto forward motion. Some
Undergo direct development in the female brood pouch, emerging other planktonic copepodscreateswimming cur-
as a postlarval/prejuvenilestagecalled a"manca."
rents by rapid vibrations of cephalic appendages/
Eggsbrooded or deposited in burrow; hatch late as clawed by which the body moves smoothly through the
pseudozoealarvae,or earlier as unclawed antizoea lawae. Both
water. "Rowing" doesoccur in the swimrning crabs
molt into distinct ericthus (and some into alima) larvae before
settling on bottom as post-larvae or juveniles. (family Portunidae) and some deep-seaasellote
isopods(e.g.,family Eurycopidae),both of which
A11free larval stageshave been los! eggsdeposited on substratum.
use paddle-shapedposterior thoracopods to scull
about.
Most eumalacostracanswith well-developed
Embryos brooded in marsupium of female typically formed from
ventral coxal plates ca1ledoostegites;usually releasedas mancas abdomensexhibit a form of temporary, or "burst,"
(subjuvenileswith incompletely developed8th thoracopods). swimming that servesas an escapereaction (e.g.,
Brood pouch (marsupium) inThermosbaenaceaformed by dorsal mysids, syncarids,euphausids,shrimps, lobsters,
caraPacecnamDer.
and crayfish). By rapidly contracting the ventral
Embryos shed or briefly brooded; typically undergo nauplius- abdominal (flexor) muscles,such animals shoot
metanauplius-ca\ptopis -fu rcilia-juvenile developmentalseries. quickly backward, the spread tail fan providing a
large propulsive surf ace (Figure 21.22C-E).This
behavior is sometimescalled a tail-flip, or " cari-
Eggs apparently brooded in female brood pouch; hatching likely doid escapereaction."
occursas eggsfloat upward after release;hatching probably as an
amphion larva (a zoea with only one pair of thoracopodsmodified Surfacecrawling by crustaceans is accomplished
as mafllipeds), with perhaps 20 stagesthat metamorphose by the samegeneralsorts of leg movements de-
gradually to a subadult. scribed in the preceding chaptersfor insects and
Dendrobranchiata shed embryos to hatch in water as nauplii or pre- other arthropods: by flexion and extension of the
zoea; all others brood embryos (on pleopods),which do not hatch limbs to pull or push the animal forward. Walking
until a pre-zoealor zoealstage(or later).
limbs are typically composedof relatively stout,
more or lesscylindrical articles (i.e.,stenopodous
limbs) as opposed to the broader, often phyllopodous shrimp (Diplostraca), most of which are largely en-
limbs of swimmers (see Figure 27.2I for a comparison closed by their carapaces (Figures 2L.4F,G,LL,M
of crustacean limb types). Walking limbs are lifted and 27.20), swim by rowing with the antennae.
from the substratum and moved forward during their Mystacocarids crawl in interstitial water using vari-
recovery strokes; then they are placed against the sub- ous head appendages. Most semi-terrestrial amphi-
stratum, which provides purchase as they move pos- pods known as "beach hoppers" (e.g., Orchestia and
teriorly through their pbwer strokes, pulling and then Orchestoidea)execute dramatic jumps by rapidly ex-
pushing the animal forward. Like many other arthro- tending the urosome and its appendages (uropods),
pods, crustaceans generally lack lateral flexibility at the reminiscent of the jumping of springtails described in
body joints, so turning is accomplished by reducing Chapter 22. Most caprellid amphipods (Figure 21.15E)
the stride length or movement frequency on one side move about in inchworm fashion, using their subche-
of the body, toward which the animal tums (like a trac- late appendages for clinging. There are also a number
tor or tank slowing one tread). Many crustaceans mi- of crustacean burrowers, and even some that build
grate; perhaps the most famous is the Chinese mitten their own tubes or "homes" from materials in their
crab (Eriocheir sinensis),which spends most of its life in surroundings. Many benthic amphipods, for example,
fresh water, but returns to the sea to breed. These crabs spin silk-lined mud burrows in which they reside.
have been found over 1000 km upriver from the sea- One species, Pseudamphithoidesincura arin (suborder
testimony to their superb locomotory ability. Perhaps Gammaridea), constructs and lives in an unusual"bi-
not unexpectedly, E. sinensisis also an important (and valved pod" cut from the thin blades of the same alga
destructive) invasive species in North America and on which it feeds (Figure 21.23A). Another gammarid-
Europe. It has been accidently introduced into the ean amphipod, Photis conchicola,actually uses empty
Great Lakes several times, but has not yet been able to gastropod shells in a fashion similar to that of hermit
establish a permanent population. crabs (Figur e 21.23C). "Hitchhiking" (phoresis) occurs
Most walking crustaceans can also reverse the di- in various ectosymbiotic crustaceans, including iso-
rection of leg action and move backward, and most pods that parasitize fishes or shrimps and hyperiidean
brachyuran crabs can walk sideways. Brachyuran amphipods that ride on gelatinous drifting plankters.
crabs are perhaps the most agile of all crustaceans. In addition to simply getting from one place to an-
The extreme reduction of the abdomen in this group other in their usual day-to-day activities, many crusta-
allows for very rapid movement because adjacent ceans exhibit various migratory behaviors, employing
limbs can move in directions that avoid interference their locomotor skills to avoid stressful situations or
with one another (and much the same thing has hap- to remain where conditions are optimal. A number of
pened, independently, in many anomurans with re- planktonic crustaceans undertake daily vertical migra-
duced abdomens). Brachyuran crab legs are hinged in tions, typically moving upward at night and to greater
such a way that most of their motion involves lateral depths during the day. Such vertical migrators include
extension (abduction) and medial flexion (adduction) various copepods, cladocerans, ostracods, and hype-
rather than rotation frontward and backward. As a riid amphipods (the latter may make their migrations
crab moves, its limbs move in various sequences, as in by riding on their gelatinous hosts). Such movements
normal crawling, but those on the leading side exert place the animals in theirnear-surface feeding grounds
their force by flexing and pulling the body toward the during the dark hours, when there is probably less
limb tips, while the opposite, trailing, legs exert pro- danger of being detected by visual predators. In the
pulsive force as they extend and push the body away daytime, they move to deeper, perhaps safer, water.
from the tips. Still, this motion is simply a mechani- These crustaceans can form enormous shoals that
cal variation on the common arthropodan walking contribute to the deep scattering layer seen on ship's
behavior. Many crustaceansmove into mollusc shells sonar. Many intertidal crustaceans use their locomotor
or other objects, carrying these about as added protec- abilities to change their behaviors with the tides. Crab
tion. In most cases,the exoskeleton of the crustacean larvae in particular are known to migrate upward or
is reduced, especially the abdomen (e.g., hermit crabs). downward according to daily rhythms, taking advan-
Of course, crustaceans grow in size as they go through tage of incoming or outgoing tides to move in and out
their molts, so these mobile-home-carrying crustaceans of estuaries. Some anomuran and brachyuran crabs
must continually find larger shells to inhabit. Many simply move in and out with the tide, or seek shelter
hermit crabs assemble in congregations to exchange beneath rocks when the tide is out, thus avoiding the
shells in a sort of group "passing of the shells" as they problems of air exposure. One of the most interesting
move into vacated larger abodes. locomotor behaviors among crustaceans is the mass
In addition to these two basic locomotor methods migration of the spiny lobster, Panulirus nrgus, in the
("typical" walking and swimming by metachronal Gulf of Mexico and northern Caribbean. Each autumn,
beating of limbs), many crustaceans move by other lobsters queue up in single file and march in long lines
specialized means. Ostracods, cladocerans, and clam across the seafloor for several davs. Thev move from
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 gammaridean amphipod Grandiderella (male and female)
in its silk-lined tube. The antennules and antennae are protruding
from the pod entrance on the right. (C) Photis conchicola, a tem-
perate eastern Pacific amphipod that spins its silken tube inside
a minute snail shell, which is then carried about in the style of
hermit crabs.
shallow areas to the edges of deeper oceanic channels.

This behavior is apparently triggered by winter storm
fronts moving into the area, and it may be a means of
avoiding rough water conditions in the shallows. water is drawn into an interlimb space as the adjacent
appendages move away from one another, and water-
Feeding borne particles are trapped by setae on the endites as
With the exception of ciliary mechanisms, crustaceans the appendages then close. From here, the trapped
have exploited virtually every feeding strategy imagin- particles are moved to a midventral food groove and
able (and some "unimaginable"). Even without cilia, then anteriorly, toward the head. This mechanism
many crustaceans generate water currents and engage of forming a boxlike "filter press" with setose phyl-
in various types of suspension feeding. We have select- lopodous limbs is the typical suspension feeding strat-
ed a few examples to demonstrate the range of feeding egy of cephalocarids, most branchiopods, and many
mechanisms that occur in this group. malacostracans.
In some crustaceans the action of the thoracic limbs Planktonic copepods were long thought to "fllter"
simultaneously creates the swimming and suspension feed by generating lateral feeding gyres or currents by
feeding currents. As the metachronal wave of append- movements of the antennae and mouth appendages. It
age motion passes along the body, adjacent Iimb pairs was believed that these gyres swept in small particles
are alternately moved apart and then pressed together, that were directly filtered by the maxillae. This clas-
thus changing the size of each interlimb space (Figure sic idea of maxillary filtration, built on work by H. G.
27.22A,8; see also Chapters 4 and 20). Surrounding Cannon in the I920s, has been questioned by recent
810 C h a p te r T w e n tv -O n e
workers, but the model persists and is still commonly (Figure 27.24A; see also Chapter 4). Emerita are adapted
presented in general works. As mentioned in Chapter to living on wave-swept sand beaches. Their compact
4, we now know that copepods and other small plank- oval shape and strong appendages facilitate burrowing
tonic crustaceans live in a world of low Relmolds num- in the unstable substratum. They burrow posterior end
bers, a world dominated by viscosity rather than by first in the area of shallow wave wash, with the ante-
inertia. Thus, the setose mouth appendages behave rior end facing upward. Following a breaking wave, as
more like paddfrs than like sieves, with a water layer the water rushes seaward, Emeritaunfurls its antennae
near the limb adhering to it and formingpart of the into the moving water along the surface of the sand.
"paddle." As the maxillae move apart, parcels of water The fine setose mesh traps protists and phytoplankton
containing food are drawn into the interlimb space. from the water, and the antennae then brush the col-
As the maxillae press together, the "patcel" is moved lected food onto the mouthparts. Many porcelain and
forward to the endites of the maxillules, which push hermit crabs also engage in suspension feeding. By
it into the mouth. Thus, food particles are not actually twirling their antennae in various patterns these ano-
filtered from the water, but are captured in small par- murans create spiraling currents that bring food-laden
cels of water. High-speed cinemitography indicites water toward the mouth (Figure 27.248,C). Food par-
that copepods are capable of capturing individual algal ticles then become entangled on setae of the mouth ap-
cells, one at a time, by this "hydraulic vacuum" meth- pendages and are brushed into the mouth by the endo-
od. In fact, copepods are probably fairly selective about pods of the third maxillipeds. Many of these animals
what they consume, which includes everything from also feed on detritus by simply picking up particles
protistan microplankton (e.g., diatoms) to other small with their chelipeds.
crustaceans. Some mud, ghost, and lobster shrimps, such as
Sessilethoracican barnacles feed by using their long, Callianassaand Upogebia(Axiidea, Gebiidea), sus-
feathery, biramous thoracopods, called cirri, to fil- pension feed within their burrows. They drive water
ter feed on suspended material from the surrounding through the burrow by beating the pleopods, and the
water (Figure 27.16A,C,8). Studies indicate that bar- first two pairs of pereopods remove food with medi-
nacles are capable of trapping food particles ranging ally directed setal brushes. The maxillipeds then comb
from2 trrmto 1 mm, including detritus, bacteria, algae, the captured particles forward to the mouth.
and various zooplankters. Many barnacles are also ca- Most other crustacean feeding mechanisms are less
pable of preying on larger planktonic animals by coil- complicated than suspension feeding and usually in-
ing a single cirrus around the prey, in tentacle fashion. volve direct manipulation of food by the mouthparts
In slow-moving or very quiet water, most barnacles and sometimes the pereopods, especially chelate or
feed actively by extending the last three pairs of cirri subchelate anterior legs. However, even in these cases
in a fanlike manner and sweeping them rhythmically of direct manipulation, the sheer number of crusta-
through the water. The setae on adjacent limbs and cean appendages adapted for feeding makes "simple"
limb rami overlap to form an effective filtering net. The feeding a surprisingly complex affair, with the vari-
first three pairs of cirri serve to remove trapped food ous mouthparts taking on tasks that include tasting
from the posterior cirri and pass it to the mouthparts. (via sensory setae), holding, chewing, scraping, and
In areas of high water movement, such as wave-swept macerating.
rocky shores, barnacles often extend their cirri into the Many small crustaceans may be classified as mi-
backwash of waves, allowing the moving water to sim- crophagous selective deposit feeders, employing vari-
ply run through the "filter," rather than moving the ous methods of removing food from the sediments in
cirri through the water. In such areas you will often which they live. Mystacocarids, many harpacticoid
see clusters of barnacles in which all the individuals copepods, and some cumaceans and gammaridean
are oriented similarly, taking advantage of this labor- amphipods are referred to as "sand grazerc" or "sand
saving device. Iickers." By various methods these animals remove de-
Most krill (euphausids) feed in a fashion similar tritus, diatoms, and other microorganisms from the sur-
to barnacles, but while swimming. The thoracopods faces of sediment particles. Interstitial mystacocarids,
form a "feeding basket" that expands as the legs move for example, simply brush sand grains with their setose
outward, sucking food-laden water in from the front. mouthparts. On the other hand, some cumaceans pick
Once inside the basket, particles are retained on the up an individual sand grain with their first pereopods
setae of the legs as the water is squeezed out laterally. and pass it to the maxillipeds, which in tum rotate and
Other setae comb the food particles out of the "trap" tumble the particle against the margins of the maxillules
setae, while yet another set brushes them forward to and mandibles. The maxillules brush and the mandibles
the mouth region. scrape/ removing organic material. Some sand-dwelling
Sand crabs of the genus Emerita (Anomura) use isopods may employ a similar feedingbehavior.
their long, setose antennae in a fashion similar to that Predatory crustaceans include stomatopods, remi-
of bamacle cirri that "passively" strain wave backwash pedes, and most lophogastrids, as well as many species
Antennae
Figure 21 .24 Some crustacean feeding mechanisms feeding hermit crabs (B) Australeremus cooki and
(see afso Figure 21 .22). ( ) Suspension feeding in the (C) Paguristes p/osus twirl their antennae, either in a circle
sand crab Emerita. The arrows point seaward and indicate or a figure eight, to create water currents that pull food
the direction of water movement as waves recede. The particles to the mouth region. (D) The predatory shrimp
antennules direct water through branchial chambers. The Procaris ascensionis (Caridea) is shown here munching on
antennae remove food particles from the water and then another shrimp (Typhlatya) as it holds the prey in a "cage"
brush them onto the mouthparts. (B,C) The suspension formed by the pereopodal endopods.
of anostracans, cladocerans,copepods,ostracods,cir- atory behavior is particularly interesting. Its prey in-

ripedes,anaspidaceans, euphausids,decapods,ta- cludes other crustaceans,particularly amphipods and
naids, isopods, and amphipods. Predation typically shrimps. After Procarlslocatesa potential victim (prob-
involves grasping the prey with chelateor subchelate ablyby chemoreception),it moves quickly to the prey
pereopods (or sometimesdirectly with the mouth ap- and graspsit within a " cage"formed by the pereopodal
pendages),or even with the antennaein the caseof endopods (Figure 21,.24D). Once captured, the prey is
predatory anostracans,followed by tearing, grinding, eatenwhile the shrimp swims about. Apparently the
or shearingwith various mouthparts, particularly the third maxillipeds pressthe prey againstthe mandibles,
mandibles.Perhapsthe most highly adaptedpredatory which bite off chunks and passthem to the mouth.
specialistsare the stomatopods(Figure 2L.7),which The remipedes capture prey with their rapto-
possessgreatly enlarged,raptorial subchelatelimbs, rial mouth appendages(Figures21..1.A,21.3D-F, and
which they useto stabor to club and smashprey. Some 2L.22F),then immobilize the victim with an injection
speciessearchout prey, but many sit in ambush at from the hypodermic maxillules.It is suspectedthat tis-
their burrow entrance.The actual attack generally fol- suesare then suckedout of the preyby actionof a man-
lows visual detectionof a potential prey item, which dibular mill and muscular foregut. They are probably
may be another crustacean,a mollusc, or even a small alsofacultativesuspensionfeedersand scavengers.
fish. Oncecaptured and stunned or killed by the rapto- Another fascinatingadaptation for predation can
rial claws, the prey is held againstthe mouthparts and be seenin many speciesof Alpheidae (the snapping
shreddedinto ingestiblepieces. shrimps, e.g.,Alpheus,Synalpheus) (Figure 21.9D).In
Although the cave-dwelling and often presumed such species,one of the chelipedsis much larger than
"primitive" shrimp Procarisis omnivorous, its pred- the other, and the movable finger is hinged in such
a way that it can be held open under muscle tension Figure 21 .25 The remarkable life cycle of the rhizo-
>
and then snapped quickly closed; this forceful clos- cephafan cirripede Peltogaster paguri, a kentrogonid
ing produces a loud popping sound and a pressure parasite of hermit crabs. (A) The mature reproductive
portion of the parasite (externa) produces numerous
or "shock" wave in the surrounding water. Some spe-
broods of male and female larvae, which are releaseo as
cies appear to use this mechanism in ambushing prey
nauplii (B,C) and eventually metamorphose into cypris lar-
(although most alpheids are probably omnivores and vae (D). Female cyprids settle on the thorax and limbs of
even include algae in their diet). When feeding in a host crabs (E) and undergo a major internal metamorpho-
predatory mode, the shrimp sits at its burrow entrance sis into the kentrogon form (F), which is provided with a
with the antennae extended. When a potential prey pair of antennules and an injection stylet. The kentrogon,s
approaches (usually a small fish, crustacean, or anne- viscera metamorphoses into an infective stage, the vermi-
gon, which is transferred to the host through the hollow
lid), the shrimp "pop"" its cheliped, and the resulting
stylet. lnside the host, the vermigon grows with rooflets
pressure wave stuns the victim, which is then quickly
that ramify throughout much of the host's body; it is
pulled into the burrow and consumed. These shrimps now called the interna (G). Eventually the female parasite
typically live in male-female pairs within the burrow, emerges on the abdomen of the host as a virginal externa
and prey captured by one individual is shared with (H). When the externa acquires a mantle pore, or aperture,
its partner. Two mechanisms have been proposed it becomes attractive to male cyprids (l). Male cyprids
for the production of the "pop" and associated shock settle within the aperture, transform into a trichogon form,
and implant part of their body contents in the female,s
wave. In some species the pop seems to be created by
receptacles (J). The deposit proceeds to differentiate into
mechanical impact of the dactylus hammering into the
spermatozoa, which fertilize the eggs of the female.
propodus. A second mechanism recently proposed is (K) The dissected externa, with its rootlets, of Peltogaster,
the collapse of cavitation bubbles, which are created removed from its host. Note the mantle aoedure.
by the rapid closure of the claw (in excessof 100 km/
sec). Captivation occurs in liquids when bubbles form
and then implode around an object (it is a well-known
phenomenon damaging ship propellers). Both mecha- range from ectoparasites with mouthparts modified
nisms create shock waves. Some boring (endolithic) for piercing or tearing and sucking body fluids (e.g.,
alpheids even use the claw snap to break off pieces of many copepods, branchiurans, tantulocarids/ several
the rock into which they are digging. In some areas of isopod families, and at least one species of ostracod) to
the world, populations of snapping shrimp are so large the highly modified and fully parasitic rhizocephalans,
that their noise disturbs underwater communication. whose bodies ramify throughout the host tissue and
In the Caribbean, colonies of alpheids have been lik- absorb nutrients directly (Figures 21.25 and2I.26).
ened to those of social insects (e.g., Synalpheusregalis Rhizocephalans, which are cirripeds that have been
colonies in Belize inhabit sponges and contain up to highly modified to become internal parasites of other
350 sibling males and females, and a single dominant crustaceans, are some of the most bizarre organisms in
breeding female). the animal kingdom. They may have typical cirripede
Many crustaceans emerge from the benthos under nauplius and cypris larvae, but in this group the cyprid
cover of darkness to feed or mate in the water column. will settle only upon another crustacean, selected to be
Many predatory isopods emerge at night to feed on in- the unfortunate host.
vertebrates or fish, particularly weak or diseased fish The most complex rhizocephalan life cycle is that of
(or fish caught in fishing nets). the suborder Kentrogonida, which are obligate para-
Macrophagous herbivorous and scavenging crusta- sites of decapods. Le this group, a settled female cypris
ceans generally feed by simply hanging onto the food larva undergoes an internal reorganization that rivals
source and biting off bits with the mandibles (a feeding that of caterpillar pupae in scope, developing an infec-
technique similar to that of grasshoppers and other in- tive stage, called the kentrogon, beneath the cyprid exo-
sects).Notostracans/ some ostracods, and many deca- skeleton. Once fully developed, the kentrogon forms
pods, isopods, and amphipods are scavengers and her- a hollow cuticular structure, the stylet, which injects a
bivores. Certain isopods in the family Sphaeromatidae motile, multicellular, vermiform creature called the ver-
bore into the aerial roots of mangrorr" ir""r. Their ac- migon into the host. The vermigon is the active infec-
tivities often result in root breakage followed by new tion stage. It has a thin cuticle and epidermis, several
multiple root initiation, creating the stiltlike appear- [,pes of cells, and the anlagen of an ovary. It invades the
ance characteristic of red mangroves (Rhizophora).A host's hemocoel by sending out long, branching, hollow
number of crustaceans are full-time or part-time detri- rootlets that penetrate most of the host's body and draw
tivores; many scavenge directly on detritus, but others nutrients directly from the hemocoel. So profound is
(e.g., cephalocarids) stir up the sediments in order to the intrusion by the rootlets that the parasite takes over
remove organic particles by suspension feeding. nearly complete control of the host's body, altering its
Finally, several groups of crustaceans have ad- morphology, physiology, and behavior. Once the para-
opted various degrees of parasitism. These animals site invades the host's gonads, parasitic castration may
\Er,"r.ru
Immature female
Trichogon produces
sperm in the receptacles
Virginal female
Receptaclesempty
Parasite
$;
+ I
penetrates
hostabdomen
I
Reproductivebody is developed
from root systemwithin host
o
Femaledevelopsinto kentrogon
larva and penetrateshost rl/T'|--
\
(F) (mb Kentrogon larvea
\4/-
*\
Kentrogon injects
vermison larva into host
i;F]?:;it;.;\ mlll
\-u /) l -
< Root systemis developedwithin host
Vermigon larvae
(see Fig. 21.26H)
(A)
Epidermal
inclusions
Cement gland
(of cypris)
Carapace
Antennule glands
Thoracopods
Cerebral ganglion
50 pm
Cuticular reinf orcement
on antennular article 1 Cernent gland
canal
Antennule
article 2
(D)
Remnant of anterior
mantle cavifz
Invasion cell Gill filament

Cement gland
of host
of kentrogon
Figure 21 .26 Various stages in the life

Kentrogon
cement gland cycle of parasitic rhizocephalans. (See
Figure 21 .25 tor a full description of a rhi-
zocephalan (kentrogonid) life cycle.) (A-D)
20pm Larvae ol Lernaeodiscus porcellanae: (A) A
live cyprid. (B) A cyprid (lateral view; SEM).
(C,D) Diagrams of cyprid larvae before and
Cement canal
after settlement (right side of carapace
Antennule removed; naupliar eye omitted). The dot-
5 ted line in the second antennular article
indicates the primordial kentrogon cuticle,
and the placement of muscle fibers in the
cyprid are indicated by arrows; the mus-
Invasion cell Anterior muscles Cement canal pore cles are hypothesized to effect formation
of the kentrogon and separation of the old
cyprid from the kentrogon. In D, kentro-
gon formation is complete. (E) A whole gill
(SEM) of a host crab, Petrolisthes cabiolli'
(Anomura), with several attached kentro-
gons (arrows). (F) A 2-hour-old kentrogon
(sagittal section). (G) A kentrogen injecting
a vermigon via the stylet. (H) A vermigon.
(l) A trichogon.
result (i.e.,the gonads of parasitized crabsnever pro- Members of the rhizocephalan order Akentrogonida
duce mature gametes).Thus the host is transformed parasitize a much wider range of crustaceanhosts
into a slave that servesthe needsof its master.The in- and do not have a kentrogon stage in their life cycle.
ternal root system,or interna, eventually develops an Instead of injecting a vermigon/ the female cyprid has
external reproductive body (the externa), where egg long, slender antennules that it usesto attach to the ab-
production occurs.A male cyprid settleson *te extema, domen of the host, one of which actually penetratesthe
transforms into a minute sexually mature instar called host's cuticle,becomeshollow, and servesfor the pas-
a trichogon, and moves into the ovary-filled extema to sageof embryonic cells from cyprid larva to host. Male
take up residence,where its sole function is to produce cyprids somehow find infected hosts and penetrate
sperm. It takes only one or two trichogons to stimulate them in the samefashion as the females,releasing their
the female ovaries to mature and begin releasingeggs sperm in such a way that they actually enter the body
into the chamber of the externa. Externa that fail to ob- of the femaleparasite.
tain male trichogons eventually die. The males are thus The akentrogonid life cycle is similar to that of the
parasitic on the female (which is itself parasitic on the Kentrogonida although, in somecases,more than one
crustaceanhost), and kentrogonids are gonochoris- individual parasite might infect a single host,lead-
tic. A mature externa, usually arising from the host's ing to multiple externas.And in at least one'genus
abdomen,will produce a successionof larval broods, (Thompsonia), multiple externacan develop from a sin-
molting after eachlarval release(it is the only part of gle infection. The anatomy of the externa is more vari-
the rhizocephalan body that molts). The larvae are leci- able than inkentrogonids, and the embryos develop di-
thotrophic and develop through severalnauplius stag- rectly into cypris larvae-there are no free-swimming
esto the cyprid (Figures 21,.25and21.26). nauplius stages.The extreme sexual size dimorphism
of kentrogonids does not occur in Akentrogonida, and is best developed in macrophagous decapods (scaven-
no migrating trichogon stage has been observed. gers, predators, and some herbivores). Thus the food
Isopods of the family Cymothoidae use modi- can be taken in quickly, in big bites, and mechanically
fied mouthparts to suck the body fluids of their fish processed afterward.
hosts. They attach to their host's skin or gills or, in
some genera, to the tongue. One species, Cymothoa Circulation and Gas Exchange
exigua, sucks so niuch blood out of its host's tongue The basic crustacean circulatory system usually con-
(the spotted rose snapper, in the Gulf of California) sists of a dorsal ostiate heart within a pericardial cav-
that the tongue completely degenerates.But, remark- ity and variously developed vessels emptying into an
ably, the fish doesn't die; the isopod remains attached open hemocoel (Figure 27.27). But the open hemocoel,
with its clawed legs to the basal muscles of the missing which in the past led to descriptions of crustaceans as
tongue, and the fish use the crustacean as a "replace- having an open circulatory system, has become highly
ment tongue" to continue to feed. This is one of the modified in many groups, and in decapods at least
few known cases of a parasite functionally replacing a there is a complex and intricate arrangement of true
host organ it destroys. vessels that has only recently begun to be recognized.
The heart is absent in most ostracods, many copepods,
Digestive System and many cirripedes. In some groups the heart is re-
The digestive system of crustaceans includes the usual placed or supplemented by accessory pumping struc-
arthropod foregut, midgut, and hindgut. The foregut tures derived from muscular vessels.
and hindgut are lined with a cuticle that is continu- The primitive heart structure in crustaceans is a long
ous with the exoskeleton and molted with it (i.e., it is tube with segmental ostia, a condition retained in part
ectodermally derived). The stomodeal foregut is modi- in cephalocarids and in some branchiopods, leptostra-
fied in different groups, but usually includes a rela- cans, and stomatopods. Flowever, the general shape of
tively short pharynx-esophagus region followed by a the heart and the number of ostia are also closely relat-
stomach. The stomach often has chambers or special- ed to body form and the location of gas exchange struc-
ized regions for storage, grinding, and sorting; these tures. The heart may be relatively long and tubular
structures are best developed in the malacostracans and extend through much of the postcephalic region
(Figure 21.27G). The midgut forms a short or long in- of the body, as it does in the remipedes, anostracans,
testine-the length depending mainly on overall body and leptostracans, or it may tend toward a globular
shape and size-and bears variably placed digestive or box shape and be restricted to the thorax (e.g., as in
ceca. The ceca are serially arranged only in the remi- cladocerans), where it may be associated with the tho-
pedes. In some malacostracans, such as crabs, the ceca racic gills (e.g., as in decapods). The intimate coevolu-
fuse to form a solid glandular mass, best called a di- tion of the circulatory system with body form and gill
gestive gland but sometimes called a midgut gland placement is best exemplified when comparing closely
or hepatopancreas, within which are many branched, related groups. Although isopods and amphipods, for
blind tubules. The digestive gland in some crustaceans instance, are both peracarids, their hearts are located
has been shown to store lipids. The hindgut is usually largely in the pleon and in the pereon, respectively,
short, and the anus is generally borne on the anal so- corresponding to the pleopodal and pereopodal gill
mite or telson, or on the last segment of the abdomen locations.
(when the anal somite or telson is reduced or lost). The number and length of blood vessels and the
Examples of some crustacean digestive tracts are presence of accessory pumping organs are related to
shown in Figure 21.27. After ingestion, the food ma- body size and to the extent of the heart itself. In most
terial is usually handled mechanically by the foregut. non-malacostracans, for example, there are no arte-
This may involve simply transporting the food to the rial vessels at all; the heart pumps blood directly into
midgut or, more commonly, processing the food in the hemocoel from both ends. These animals tend to
various ways prior to chemical digestion. For example, have short bodies, long hearts, or both, an arrange-
the complex foregut of decapods (Figure 21.27G) is di- ment that facilitates circulation of the blood to all body
vided into an anterior cardiac stomach and a posterior parts. Sessile forms, such as most cirripedes, have lost
pyloric stomach. Food is stored in the enlarged por- the heart altogether, although it is replaced by a vessel
tion of the cardiac stomach and then moved a bit at a pump in the thoracicans. Large malacostracans tend to
time to a region containing a gastric mill, which usually have well developed vessel systems, thus ensuring that
bears heavily sclerotized teeth. Special muscles associ- blood flows throughout the body and hemocoel and to
ated with the stomach wall move the teeth, grinding the gas exchange structures (Figure 21.27D,8). Recent
the food into smaller particles. The macerated material studies using corrosion casting techniques have shown
then moves into the back part of the pyloric stomach, just how complex these vessel systems can be and have
where sets of filtering setae prevent large particles from even called into question the paradigm of "open" vs.
entering the midgut. This type of foregut arrangement "closed" circulatory systems in invertebrates. Large or
Figure 21 .27 lnternal anatomy of representative

crustaceans. (A) A calanoid copepod. (B) A lepado-
morph barnacle. (C) A balanomorph barnacle.
(D) A stomatopod. (Continued on next page)
(B) Tergum
Cirri
Scu
Foregut
Maxill
Mandibular palp
(mandible Supraesophageal Maxillary
ganglion gland
Adductor muscle
Adductormuscle Ventral
Labrum of scutum nerve mass
Esophagus Rostal
Supraesophageal blood
ganglion sinus
Maxilla Depressor
Midgut
muscle
Digestive ceca
of tergum
Stomach Testis Penis
Oviduct Anus
Stomach
Ovary (peduncle) Egg mass
Mantle
in mantle
cavity
cavity
Sperm duct
Testis
Antennule Antennule
Accessory Supraesophageal
(D) Anterior gland
Heart ganglion
aorta
Optic artery
Sperm duct
Optic nerve
Midgut
Compound
Midgut gland
Antennal artery
Supraesophageal
ganglion Maxillary
gland
Esophagus
Subesophageal
ganglion

(E) Anterior Cardiac Pyloric Posterior

Cor frontale lateral Anterior stomach stomach
artery aorta
Optic artery O-stium_g""r1
Hindgut
Supraesophageal
ganglion
Antennal artery Ganglion
Antennal nerve Venhal
Antennal gland Midgut abdominal artery
Midgut
gland
Sternal
Midgut artery
Subesophageal cecum Oviduct
ganglion Venhal
thoracic artery
Anterior midgut cecum
Ostia Midgut
(F) Anterior / Heart
Testis
_ aorta
)upraesopnageal
ganglion Sperm duct
Antennular Penial
papilla
Posterior
aorta
Foregut
Posterior
midgut cecum
Antennal
gland
Circumesophageal
connective
Hindgut
Midgut
glands
(G) Lateral tooth Dorsal tooth Subesophageal

of gastric mill of gastric mill ganglion
Cardiac stomach Pyloric stomach

Fused ganglia
Straining setae Ventral nerve cord
Intestine 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 cara-
pace removed.
Shaining setae
active crustaceans may also possess an anterior acces- Crustacean blood contains a variety of cell types,
sory pump called the cor frontale, which helps main- including phagocytic and granular amebocytes
tain blood pressure/ and often a venous system for re- and special wandering explosive cells that release
turningblood to the pericardial chamber. a clotting agent at sites of injury or autotomy. In
(H) Supraesophageal
oanolinn
Anterior
stomach
Antenna
Compound eye muscle Lateral adductor
muscle of mandible
Branchial chamber: Cheliped

Cardiac stomach
Opthalmic artery
Gilt
Digestive gland
Epipod of Pereooodalmuscles
first maxilliped
Anterior dorsal
pyloric muscle
Ovary
Pyloric stomach
Pericardial sac
Muscles of
fifth pereopod
Hindgut cecum
Superior abdominal
artery
Lateral abdominal
artery
non-malacostracans, oxygen is either caffied in solu- an organ with a relatively high surface area. The gills
tion or attached to dissolved hemoglobin. Most mala- provide a thin, moist, permeable surface between the
costracans possesshemocyanin in solution (although internal and external environments. The gills of sto-
some contain hemoglobin within tissues). Hemoglobin matopods and isopods (Figure 21.28H,I) are formed
uses iron as the oxygen-binding site, whereas hemo- from the abdominal pleopods. In the first case they are
cyanin uses copper. The latter can give a bluish color to branched processes off the base of the pleopods, but in
the hemolymph; carotenoid pigments frequently give the isopods the flattened pleopods themselves are vas-
hemolymph a pale brown or orange color. Oxygen- cularized and provide the necessary surface area for
binding pigments are never carried in corpuscles as exchange. Stomatopods also have epipodal gills on the
they are in the vertebrates. thoracopods, but these are highly reduced.
We have mentioned the form and position of gas ex- For gills to be efficient, a flow of water must be
change organs (gills) for some groups of crustaceans in maintained across them. In stomatopods and aquat-
the taxonomic synopses. Some small forms (e.g., cope- ic isopods a current is generated by the beating of
pods, some ostracods) lack distinct gills and rely on cu- the pleopods. Similarly, the pereopodal gills of eu-
taneous exchange, which is facilitated by their relative- phausids are constantly flushed by water as the ani-
ly thin cuticles and high surface area-to-volume ratio. mal swims. In many crustaceans, however, the gills
In the small forms of other groups a thin, membra- are concealed to various degrees and require special
nous inner lining of the carapace serves this purpose mechanisms in order to produce the ventilating cur-
(e.g., Cladocera, Cirripedia, Leptostraca, Cumacea, rents. In most decapods, for example, the gills are
Mysida, clam shrimps, and even some members of the contained in branchial chambers formed between the
Decapoda). carapace and the body wall (Figure 2L.28). Thus the
Most crustaceans, however, possess distinct gills of delicate gills, while still technically outside the body,
some sort (Figure 21,.28).These structures are common- appear to be (and are protected as though they were)
ly derived from epipods (exites) on the thoracic legs internal organs. While such an arrangement provides
that have been modified in various ways to provide a protection from damage to the fragile gill filaments,
large surface area. The inner hollow chambers of these the openings to the chambers are generally small, re-
gills are confluent with the hemocoel or their vessels. stricting the passive flow of water. Not surprisingly,
Although their structure varies considerably (recall the the solution to this dilemma comes once again from
various decapod gills described earlier), they all oper- the evolutionary plasticity of crustacean appendages.
ate on the basic principles of gas exchange organs ad- Most decapods have elongate exopods on the maxil-
dressed in Chapter 4 and throughout this text: the cir- lae, called gill bailers or scaphognathites, that vibrate
culatory fluid is brought close to the oxygen source in to create ventilating currents through the branchial
Blood vessels
(c)
Baseof first leg
(c) Branchial (H) Attachment point

chamber Inner carapacewall to body wall
Epipod of
first maxilliped Protopod
Body wall
Pseudotracheae
Epipod of .::.r:-.'.':::
second maxilliped
l- Merus
Sternum Coxa Basi-ischium
Figure 21 .28 Gas exchange structures. (A) Maxilla of

the shrimp Pandalus. Note the setose scaphagnathite
used to generate the ventilating current. (B-D) Cross sec- sand or mud with only their front ends exposed to the
tions of types of decapod gills: (B) Dendrobranchiate. water.
(C) Trichobranchiate. (D) Phyllobranchiate. (E,F) Paths of
The positioning of the gills in branchial chambers
ventilating currents through the left branchial chambers of
(E) a shrimp and (F) a brachyuran crab. (G) The branchial
protects them from desiccation during low tides and
chamber (cross section) of a brachyuran crab, showing thus enables many crustaceans to live in littoral habi-
the position of a single phyllobranchiatepodobranch. (H,l) tats; diffusion of respiratory gases commonly contin-
A pfeopod of the terrestrial isopod Porcellio (sur.faceview ues even during low tides. Some decapods have even
and section). Note the pseudotracheae. invaded land, especially certain crayfish and the ano-
muran and brachyuran crabs known as land crabs (e.g.,
the hermit crab Coenobita andthe coconut crab Birgus;
chambers (Figure 21.28A). These currents typically Figure 2I.IH).In these semi-terrestrial species the gills
enter from the sides and rear through small openings are typically reduced in size. InBirgus the original gills
around the coxae of the pereopods (called in crabs are very small, and the vascularized cuticular surface
Milne-Edwards openings, after their discoverer), and of the gill chamber is used for gas exchange. Although
exit anteriorly from under the carapace in the vicin- young Birgus may carry shells (or coconuts) to protect
ity of the mouth field (and antennal glands). They can their soft abdomen, adults do not and the abdomen
be easily seen by observing a crab or lobster in quiet is hardened. Another striking decapod adaptation
water. The flow rate of the currents can be altered, de- to life in air is displayed by the sand-bubbler crabs of
pending on envirorunental factors, and can also be re- the Indo-Pacific region (family Dotillidae: Scopimera,
versed, thus allowing certain decapods to burrow in Dotilla). These crabs possess membranous discs on
their legs or sternites that were once thought to be au- tubule. These activities not only regulate the loss of
ditory organs (tympana), but are now thought to func- metabolic wastes but are also extremely important in
tion as gas exchange surfaces. water and ion balance, particularly in freshwater and
The most successful crustaceans on land are not terrestrial crustaceans.
the decapods, however, but the familiar sowbugs The excretion and osmoregulation carried out by an-
and pillbugs. The successof these oniscidean isopods tennal and maxillary gland activity are supplemented
(e.g., Porcelllo) is due in part to the presence of aerial by other mechanisms. The cuticle itself acts as a barrier
gas exchange organs called pseudotrachea (Figure to exchange between the internal and extemal environ-
21.28Hj). These organs are inwardly directed, moder- ments and, as we have mentioned, is especially im-
ately branched, thin-walled, blind sacs located in some portant in preventing water loss on land or excessive
of the pleopodal exopods, connected to the outside via uptake of water in fresh water. Moreover, thin areas of
small pores (similar to insect tracheae and spiracles). the cuticle, especially the gill surfaces, serve as sites of
Air circulates through these sacs, and gases are ex- waste loss and ionic exchange. The epipods on the legs
changed with the blood in the pleopods. Thus, in these of Branchiopoda were long assumed to function in gas
animals the original aquatic pleopodal gills have been exchange (as "gills") but they are now known to serve
refashioned for air breathing by moving the exchange primarily as sites of osmoregulation (hence, the taxo-
surfaces inside, where they remain moist. The superfi- nomic name Branchiopoda, meaning gill-footed, is a
cially similar tracheal systems of isopods, insects, and misnomer!). Phagocytic blood cells and certain regions
arachnids evolved independently, by convergence, in of the midgut are also thought to accumulate wastes.
association with other adaptations to life on land. In some terrestrial isopods, ammonia acfually diffuses
from the body in gaseous form.
Excretion and Osmoregulation
Like other fundamentally aquatic invertebrates, crusta- Nervous System and Sense Organs
ceans are ammonotelic, whether in fresh water or sea- The central nervous system of crustaceans is con-
water or on land. They release ammonia both through structed in concert with the segmented body structure,
nephridia and by way of the gills. As discussed in along the same lines as seen in other arthropods (Fig-
Chapter 20, most crustaceans possess nephridial ex- ure 21.29).In the more primitive condition it is lad-
cretory organs in the form of either antennal glands or derlike, the segmental ganglia being largely separate
maxillary glands (Figures 21.54 and 21.27). These are and linked by transverse commissures and longitudi-
serially homologous structures, constructed similarly nal connectives (Figure 21.294). The crustacean brain
but differing in the position of their associated pores is composed of three fused ganglia, the two anterior
(at the base of the second antennae or the second max- being the dorsal (supraesophageal) protocerebrum and
illae, respectively). The inner blind end is a coelomic deutocerebrum, which are thought to be preoral in ori-
remnant of the nephridium called the sacculus, which gin. From the protocerebrum, optic nerves innervate
leads through a variably coiled duct to the pore. The the eyes. From the deutocerebrum, antennulary nerves
duct may bear an enlarged bladder near the opening. run to the antennules, while smaller nerves innervate
Antennal glands are sometimes called "gteerrglands." the eyestalk musculature. The third ganglion of the
Most crustaceans have only one pair of these ne- brain is the posterior tritocerebrum, which presumably
phridial organs/ but lophogastrids and mysids have represents the first postoral somite ganglion. The tri-
both antennal and maxillary glands, and a few others tocerebrum forms a pair of circumenteric connectives
(cephalocarids and a few tanaids and isopods) have that extend around the esophagus to a subesophageal
well developed maxillary and rudimentary anten- or subenteric ganglion and link the brain with the ven-
nal glands. Most non-malacostracans have maxillary tral nerve cord bearing the segmental body ganglia.
glands, as do stomatopods, cumaceans/ and most ta- From the tritocerebrum also arise the antennary nerves
naids and isopods. Adult ostracods have maxillary as well as certain sensory nerves from the anterior re-
glands, but antennal glands also occur in freshwater gion of the head.
species. All of the other malacostracans have antennal The nature of the ventral nerve cord often clearly
glands. reflects the influence of body tagmosis. In crustaceans
Blood-filled channels of the hemocoel intermingle with relatively homonomous bodies (e.g., remipedes,
with branched extensions of the sacculus epithelium, cephalocarids, and anostracan and notostracan bran-
creating a large surface area across which filtration oc- chiopods), the ganglia associated with each postanten-
curs. The cells of the sacculus wall also actively take up nary segment remain separate along the ventral nerve
and secrete material from the blood into the lumen of cord. In more heteronomous forms, however, a single
the excretory organ. These processes of filtration and large subenteric ganglionic mass is formed by the fu-
secretion are to some degree selective, but most of the sion of ganglia associated with the postoral cephalic
regulation of urine composition is accomplished by segments (e.g., those of the mandibles, maxillules,
active exchange between the blood and the excretory maxillae, and, when present, maxillipeds). The ganglia
(A) (B)
Brain Brain
Esophagus
r
I
I
Thoracic
(c)
Antennulary
ganglia nerve
II Antennary
nerve
Optic nerve
I
L-
rI Esophagus
Mandibular _
nerve
I
Abdominal
ganglia I
*s \j.:
Thorlcic 5,
I newe plate Aj|
I
I
Pleon ganglia
Figure 21.29 Central nervous systems of four crusta- brachyuran crab, wherein all thoracic ganglia have fused
ceans. (A)The ladderlikesystem of an anostracan.Note and the abdominal ganglia are reduced. (D) Nervous sys-
the absenceof well developedgangliain the posterior, tem of a hyperiid amphipod. Note the loss of the urosomal
apodous,portion of the trunk. (B) Elongatemetameric ganglia typical of all amphipods.
system of a crayfish.(C) Highlycompactedsystem of a
of the thorax and abdomen may also be variably fused, In decapods, hundreds of neurons can innervate each
depending on segment fusion and body compaction. aesthetasc.Thermoreceptorsprobably occur in many
For example, in most long-bodied decapods (lobsters crustaceans(thoseliving near hydrothermal vents in
and crayfish), the thoracic and abdominal ganglia are the deep seawould seemlikely candidates)but are not
largely fused across the body midline, but remain sepa- yet documented.However, behaviors related to ther-
rate from one another longitudinally (Figure Z1,29B). mal avoidanceand temperaturepreferenceshave been
However, in short-bodied decapods (e.g., crabs), all shown, with thermosensitivities reported from 0.2 to
of the thoracic segmental ganglia are fused to form a 2.0' C. A pair of unique sensorystructureswhose func-
large ventral nerve plate, and the abdominal ganglia tion is unknown, called frontal processes,occurson
are much reduced (Figure 21.29C). the head of remipedes.Many crustaceanshave dorsal
Most crustaceans have a variety of sensory recep- organs,poorly understood glandular-sensorystruc-
tors that transmit information to the central nervous tures on the head,which actually constituteseveraldif-
system in spite of the imposition of the exoskeleton ferent types of sensorystructuresthat may or may not
(as previously explained for arthropods in general) behomologous.
(Figure 21..30).Among the most obvious of these senso- Like all arthropods,crustaceanscontain well-devel-
ry structures are the many innervated setae or sensilla oped proprioceptorsthat provide information about
that cover various regions of the body and appendages body and appendageposition and movement during
(Figure 21.3I). Studies of these structures suggest most locomotion. A few taxa within the classMalacostraca
function as both mechanoreceptors (sensing touch and possessstatocysts,which either are fully closedand
currents) and chemoreceptors. Most crustaceans also contain a secretedstatolith (e.g.,mysids, some an-
possess special chemoreceptors in the form of clumps thurid isopods)or open to the outside through a small
or rows of soft, fubular, cuticular processes called aes- pore and contain a statolith formed of sand grains
thetascs (Figure 21.30,4.)located on the first antennae. (e.9.,many decapods)(Figure 21.308,C).Inthe latter
case the statocyst not only serves as a georeceptor, but stage is suppressed. Median eyes are in a sense "com-
also detects the angular and linear acceleration of the pound" in that they are composed of more than one
body relative to the surrounding water as well as the photoreceptor unit (Figure 21.30D). There are typical-
movement of water past the animal (i.e., the statolith ly three such units in the median eyes of nauplii and
is rheotactic). And, in some shrimp, the statocyst is ap- up to seven in the eyes of adults in which they persist.
parently also engaged in hearing. Except for their basic rhabdomeric nature, however,
There are two types of rhabdomeric photoreceptors the structure of median eye units is unlike that of the
among crustaceans, median simple eyes and lateral ommatidia of true compound eyes. The former are in-
compound eyes; both are innervated by the protoce- verse pigment cups, each with relatively few retinu-
rebrum. Many species possessboth kinds of eyes, ei- lar (photoreceptor) cells. Cuticular lenses are present
ther simultaneously or at different stages of develop- over the median eyes of most ostracods and some
ment. The compound eyes may be sessile or stalked. copepods. Simple crustacean eyes probably function
Stalked compound eyes occur in the Anostraca, many only to detect light direction and intensity. Such infor-
Malacostraca, and perhaps some Cumacea (and per- mation is of particular value as a means of orientatiori
haps also some trilobites). These are the only examples in planktonic forms without compound eyes, such as
of moveable stalked compound eyes in the animal nauplius larvae, many copepods, etc. In some bran-
kingdom. chiopods, a space above the median eye is connected
The median eye generally first appears during the to the external environment by a small pore, perhaps
nauplius larval stage, and for that reason it is often indicating an invagination of the eye at some point in
called a naupliar eye. Like the nauplius larva itself, the distant past, although the nature and function of
the median eye is thought to be an ancestral (defin- the pore is not known.
ing) feature of the Crustacea; it is secondarily reduced The structure and function of compound eyes (om-
or lost in many taxa in which the corresponding larval matidia) were reviewed in Chapter 20.In terms of
(c) Edge of
Thread cutaway Opening to
hairs cuticle statocyst
rd setae
Outer row of
free hairs
Large free Convex side of pigment

Statolith anterolateralhairs cup of ventral ocellus
Aesthetascs
(chemosensorysetae) Inner rows of hair
Frontal nerve
supporting statolith
Suspensory
band
Retinal cell
Retinal cell
Connective
tissue cover
Pigment cup
(at base of endopod)
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
mysid. (C) Open statocyst in the antennules of a lobster
(cutaway view). (D) A median simple eye of the copepod
Telson Eucalanus (surface view).
Figure 21 .31 Sensilla of selected crustaceans. (A) Serrate seta (mecha-

noreceptor) of the second maxilliped of the anomuran petrolisthes armatus
$42Q. (B) Sockets of serrate setae of the third maxilliped of the cleaning
shrimp Sfenopus hispidus (x228). (C) Current receptor from the anterior
trunk limbs of the notostracan Triops. (D) chemosensory seta from the first
pereopod of the freshwater shrimp Atya (xSZOO);note the characteristic
apical pore. (E) A dual receptor (mechanoreceptor and chemoreceptor)
seta from the maxilla of the remipede Spe/eonectes tulumensis (x4560).
(F) Cross section of a dual receptor (micrograph and interpretive drawing;.
Note the microtubules within the dendrites and the dendritic sheath that
dttaches to the cuticle of the setal shaft (x17,100).
visual capacity, much more work has been done on the

eyes of insects than on those of crustaceans/ and we are Dendritic sheath Microtubules
left with a good deal of speculation in terms of what
crustaceans actually see. Although most probably lack
the visual acuity of many insects, it has been shown the ultraviolet and far-red spectra,at least to 470-570
that many crustaceans can discern shapes, patterns, nm). Stomatopods,in particular, have been shown
and movement; color vision has been demonstrated to have extremely complex and sensitive eyeswith a
in some species (various species have been shown to wide range of capabilities. Running through the center
respond to light waves from the blue-green region to of eachstomatopod eye is a midband of 6 ommatidial
rows-which in most species contain 4 rows that make systems is far from complete. In general, the phenom-
up a color vision system with 12 visual pigments-as ena of molting (see Chapter 20), chromatophore activ-
well as 2 rows that analyze both linearly and circularly ity, and various aspects of reproduction are under hor-
polarized light. The ommatidia that deal with color monal and neurosecretory control. Interesting recent
have a three-tier structure: the top tier contains an ul- work indicates that juvenile hormone-like compounds,
traviolet-sensitive pigment, and below this are 2 tiers long thought to occur only in insects, may also occur
with different pigments sensitive to wavelengths in the in at least some crustaceans. (]uvenile hormones are
human visible spectrum. The upper and lower halves a family of compounds that regulate adult metamor-
of each eye also have overlapping visual fields, so that phosis and gametogenesis in insects.) Bioluminescence
each eye can act as a stereoscopic rangefinder. The 12 also occurs in several crustacean groups. It is common
different photoreceptor types each sample a narrow set among pelagic decapods, and it has also been reported
of wavelengths ranging from deep UV to far red (300 in certain myodocopan ostracods, hyperiid amphi-
to 720 nm). pods, and copepod larvae.
Although both insects and crustaceans have tetra-
partite ommatidia, there are certain structural differ- Reproduction and Development
ences between the compound eyes of insects and those Reproduction We have often mentioned the relation-
of crustaceans, probably as a result of adaptation to ships between an animal's reproductive and develop-
the requirements of aerial and aquatic vision. ln water, mental pattern and its lifestyle and overall survival
light has a more restricted angular distributior; a lower strategy. With the exception of purely vegetative pro-
intensity, and a narrower range of wavelengths than cesses such as asexual budding, the crustaceans have
it does in air. Contrast is also somewhat reduced in managed to exploit virtually every life history scheme
water. All of these factors place a premium on enhanc- imaginable. The sexes are usually separate, although
ing the sensitivity and contrast perception of the eyes hermaphroditism is the rule in remipedes, cephaloca-
of aquatic creatures. Mounting the eyes on stalks is one rids, most cirripedes, and a few decapods. Sequential
dramatic way in which many crustaceans increase the hermaphroditism is not uncommon and usually is
amount of information available to the eyes, by increas- expressed as protandry (individuals first mature as
ing the field of view and binocular range. Eyestalks are males, then later become females), although protog-
complex structural features with a dozen or so muscles yny occurs in a few species (e.g., the marine isopod
controlling their movement. Gnorimosphaeroma orgonense).Irr addition, parthenogen-
Typical tetrapartite compound eyes are lacking in esis is known in some branchiopods and certain ostra-
the small crustaceans formerly combined as "maxillop- cods. ln one species of clam shrimp (Eulimnndia texana)
odans" (copepods, barnacles, etc.), but various forms a rare type of mixed mating system exists, called and-
of "compound eyes" do occur among the Branchiura, rogonochorism (i.e., androdioecy in plants), in which
Ostracoda (Cypridinacea), and Cirripedia. Eyes in the males coexist with hermaphrodites, but there are no
first fwo taxa most closely resemble those of other crus- true females. Androgonochorism is rare, but is also
taceans in general structure and may be homologous known in the nematode Caenorhabditiselegans,sorne
with them. In the Cirripedia, the median eye and two thoracican b arnacles (e.g., Balanus g aleatus,Scalpellum
lateral eyes are all derived from a single tripartite ocel- scalpellum), and from several other branchiopod
lar eye of the nauplius larva, which splits into its three crustaceans.
components, each forming an adult photoreceptor fol- The reproductive systems of crustaceans are gener-
lowing metamorphosis of the nauplius into a cypris ally quite simple (Figure 27.27).The gonads are derived
larva. All three of these eyes thus appear to be com- from coelomic remnants and lie as paired elongate
posed of simple ocelli, although the lateral eyes have structures in various regions of the trunk. In many cir-
three photoreceptor cells and for this reason are often ripedes, however, the gonads lie in the cephalic region.
called "compound eyes." Rhizocephalan nauplii also In some casesthe paired gonads are partially or wholly
have a tripartite nauplius eye, which persist into the fused into a single mass. A pair of gonoducts extends
cyprid larval stage. from the gonads to genital pores located on one of the
Compound eyes are lacking altogether in many crus- trunk segments, either on a sternite, on the arthrodial
tacean taxa (e.g., Copepoda, Mystacocarida, Cepha- membrane between the sternite and leg protopods, or
locarida, Tantulocarida, Pentastomida, Remipedia, on the protopods themselves. Lr many crustaceans the
and some Ostracoda). Members of some other groups paired penes are fused into a single median penir (".9.,
possess compound eyes only in late larval stages and in tantulocarids, cirripedes, and some isopods). The fe-
lose them at metamorphosis (e.g., some cirripedes). male system sometimes includes seminal receptacles.
Reduction or loss of eyes is also common in many deep- The position of the gonopores varies among the classes
sea species,burrowers, cave dwellers, and parasites. (Table21.1).
Crustaceans have complex endocrine and neurose- The curious phenomenon of intersex-having both
cretory systems, although our understanding of these male and female secondary sexual characteristics-is
Figure 21 .32 Reproduction in Crustacea. (A-D) Mating )

widespread among crustaceans.Intersexual develop-
behaviors of the fiddler crab Uca. (A) Two males in ritual-
ment is associated directly with the presence of endo- ized combat for the favor of a female, while she watches
parasites such as bacteria and microsporidia, and it has (B). (C) A single male waving his enlarged cheliped to
also been correlated with the presence of endocrine- attract a female. (D) A male fiddler crab engaged in claw-
disrupting pollutants that seem to induce opposing waving behavior to attract a female. (E) A balanomorph
secondary sexual characterisitcs. barnacle, with cirri and groping penis extended, impreg-
Most crustaceans copulate, and many have evolved nating a neighbor. The advantage of a long penis in ses-
sile animals is made obvious by this illustration.(F) Ventral
courtship behaviors, the most elaborate and well
views of a male and female brachyuran crab, Cancer
known of which occur among the decapods. Although magister, showing the modified pleopods (setose append-
many crustaceans are gregarious (e.9., certain plank- ages to retain eggs in female; modified as gonopod in
tonic species, barnacles, many isopods and amphi- male). (G) A copulating pair ot Hemigrapsus sexdentafus.
pods), most decapods live singly except during the (H) The planktonic copepod Sapphirina, with egg sacs.
mating season. More or less permanent, or at least sea-
sonal, pairing is known among many crustaceans (e.g.,
stenopodid shrimps; certain parasitic and commensal
isopods; pinnotherid "pea" ctabs, which often live as A good deal of work has been done on fiddler crabs
pairs in the mantle cavities of bivalve molluscs or in of the genus Uca (family Ocypodidae). [r these species,
burrows of thalassinid shrimps). males engage in dramatic cheliped waving (of their
Even the parasitic pentastomids copulate (within greatly enlarged chela, or major claw, which can ac-
the host's respiratory system) and have internal fertil- count for more than 50% of the male's total mass-more
ization, relying on a transfer of sperm to the female's than the largest rack of a male elk!) to atlract females and
vagina by way of the male's cirri (penes). Pentastomid repel competing males (Figure 27.32A-D).In addition,
early embryos metamorphose into a so-called primary males produce sounds by stridulation and substratum
larva with two pairs of double-clawed legs and one or thumping, which are thought to attract potential mates.
more piercing stylets (Figure 21.19F,).The larvae may Mating generally takes place once the male has enticed
be autoinfective in the primary host, or they may mi- the female into his burrow. Males of some fiddler crab
grate to the host's gut and pass out with the feces. In species build sand structures at the entrance of their bur-
the latter case, an intermediate host is required, which row, which have been shown to athact females.
may be almost any kind of vertebrate. The larvae bore Among many crustaceans/ the external sexual char-
through the gut wall of the intermediate host, where acteristics are associated with the actual mating pro-
they undergo further development to the infective cess. In some males, particular appendages, such as
stage. Once the intermediate host is consumed by a de- the antennae of anostracans and some cladocerans, os-
finitive host (usually a predator), the parasite makes its tracods, and copepods, are modified for grasping the
way from the new host's stomach up the esophagus, or female. Additionall!,rrrany males bear special sperm
bores through the intestinal wall, eventually settling in transfer structures, in the form of either modified ap-
the respiratory system. pendages or special penes such as those of the thoraci-
Mating in nonpaired crustaceans requires mecha- can barnacles (Figure 27.32E), anostracans, and ostra-
nisms that facilitate location and recognition of part- cods. Examples of modified appendages include the
ners. Among decapods, and perhaps many other last trunk limbs of copepods and the anterior pleopods
ctustaceans, scattered individuals apparently find one of most male malacostracans (called gonopods in most
another either by distance chemoreception (phero- malacostracans, or petasma in Dendrobranchiata)
mones) or through synchronized migrations associated (Figure 27.32F). Sperm are transferred either loose in
with lunar periodicity, tidal movements, or some other seminal fluid or (in many malacostracans and in cope-
environmental cue. Contact sex pheromones are also pods) packaged in spermatophores. Motile flagellated
utilized. Males of some marine myodocopan ostracods sperm occur only in some of the former maxillopodan
(some Halocyprididae, some Cypridinidae) produce groupsi in other crustaceans the sperm are nonmotile.
complex bioluminescent displays, similar to those of Crustacean sperm are highly variable in shape, even
fireflies, to attract females. Once prospective mates are bizate in many instances, often being large round or
near each other, recognition of conspecifics of the op- stellate cells that move by pseudopods or are, seem-
posite sex may involve several mechanisms. Over 60 ingly, nonmotile.e Sperm are deposited directly into
speceis of cypridinid ostracods engage in luminescent the oviduct or into a seminal receptacle in or near the
courtship behavior in the Caribbean alone. Apparently, female reproductive system. In some crustaceans
most decapods employ chemotactic cues requiring
actual contact. Vision is known to be important in the
eThe sperm of freshwater ostracods
stenopodid shrimps (most of which live in pairs), in are the longest in the animal
kingdom, relative to body size (up to 10x body"length). Even
certain anomurans (e.9., the family Porcellanidae), and though their sperm are aflagellate, they are filiform and range
inbrachyurans (many grapsids and ocypodids). from several hundred microns to millimeters in leneth.
PHYLUMARTHROPODA Crustacea:Crabs, Shrimps, and Their Kin 827
''La
La
,/A\ 1b (B) Dorsal
Mandibular Antennal
ectoderm ectoderm
Antennular
Postnaupliar ectoderm
ectoderm I
I
2d Protocerebral
I
I ectoderm
i
I
Midgut 3c aa
I
(entoderm) 2b I
(G)
First
Telson
Figure 21 .33 Cleavage and posthatching stages among crustaceans.

(A) Modified holoblastic spiral cleavage has produced a 28-cell embryo of the
cirripede 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 divi-
sions in the early cleavage of a mysid. (D) Newly hatched copepod nauplius
larva. (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
dendrobranchiate shrimp Penaeus. (H) Zoea larva of the brachyuran crab
Callinectes sapidus. (l) Zoea larva of a porcelain crab. (J) Megalopa larvae of
the xanthid crab Menippe adina. (K) The characteristic antizoea larva of a sto-
matopod. (L) The translucent, paper-thin phyllosoma larva of the lobster Jassa.
(M) Cryptoniscus stage (not a true larva) of the epicaridean isopod probopyrus
bithynis. (N) Cyprid larva of a barnacle.
femalescan storespermfor long periods (e.g.,several embryos in a marsupium, a ventral brood pouch
yearsin the lobsterHomarus),thus facilitating multiple formed from inwardly directed plates of the leg coxae
broods from singleinseminations. called oostegites (thermosbaenaceans are an excep-
The great majority of crustaceansbrood their eggs tion among the Peracarida and use the carapace as a
until hatching occurs,and a variety of brooding strat- brood chamber). Other crustaceans attach the embryos
egieshas evolved. Peracaridsbrood the developing to endites on the bases of the legs or to the pleopods
Antennule
Antenna
First thoracopod
Secondthoracopod
Pleotelson
little about their embryology. The eggs are centro-

lecithal, with various amounts of yolk. The amount of
yolk greatly influences the type of early cleavage and
is often related to the time of hatching (Chapter 4). As
(Figure 21..32F),usually using mucus secreted by spe- far as is known, the zygotes of most non-malacostra-
cialized glands. In some cladocerans, brooding takes cans undergo some form of holoblastic cleavage, as do
place in a dorsal brood chamber formed by the cara- those of slmcarids, euphausids, penaeids, amphipods,
pace. However, the slmcarids, almost all dendrobran- and parasitic isopods. However, cleavage patterns are
chiate shrimps, and most euphausids shed the zygotes extremely variable, ranging from equal to unequal
directly into the water. A few others deposit their fertil- and from radial-like to spiral-like. The occurrence of
ized eggs in the environment, usually attaching them modified spiral cleavage (Figure 21.33A) reported from
to some object (e.g., branchiurans, some ostracods, some crustaceans had generally viewed as evidence
many stomatopods). These deposited embryos may be of close ties between the crustaceans and other spira-
abandoned or, as is the case in stomatopods, carefully lian groups such as the annelids. However, the gen-
tended by the female. Nonetheless, parental protection erality of spiral-like cleavage among crustaceans has
of the embryos until they hatch as larvae or juveniles been called into question with evidence from molecu-
is typical in crustaceans. Thus, crustaceans usually en- lar phylogenetics that indicates arthropods belong to
gage in mixed or direct life histories (Table21..2). the Ecdysozoa clade of Protostomia, not the Spriralia
(where annelids are nested). And in some crustacean
Development Although crustaceans are the most groups, the cell lineages and germ layer origins are
widespread animals on Earth, we know surprisingly quite different from those of typical spirally cleaving
830 ChaoterTwentv-One
embryos. For example, in barnacles the mesodermal subsequent phases, with each phase containing slightly
germ layer arises from the 3A, 3B, and 3C cells, and different morphological steps called stages, before the
the 4d cell contributes to ectoderm (Figure 21'338)- adult condition is achieved. Direct development oc-
whereas typical spiral cleavage involves a 4d origin of curs in some cladocerans and branchiurans, and in
mesoderm. Euphausids were long thought to have spi- all ostracods, phyllocarids, slmcarids, and peracarids.
ral cleavage, but recent work indicates they do not- Ostracods are tlpically viewed as having direct devel-
only the oblique ahgle of the mitotic spindles during opment, and they lack a distinct larval stage. However,
the transition from the 2- to the 4-cell stage resembles some ostracod species hatch with only the first three
spiral cleavage. There are also differences between pairs of appendages present, and they are thus true
various crustacean and other arthropod taxa involving nauplii, even though they are in a bivalved carapace
the positions of the presumptive germ layers relative to and add limbs gradually (the juvenile instars resemble
one another, especially the endoderm and mesoderm. miniature adults). All other crustaceans have some
We want to emphasize, however, that such variations form of mixed development. The larval stages that
are not surprising in such a diverse and ancient taxon have been recognized in crustacean groups that under-
and do not negate the fundamental similarities that go mixed development have been assigned a plethora
unite the Arthropoda. of names, and the homologies among these forms are
Meroblastic cleavage is the rule among many mala- not always understood. The more commonly encoun-
costracans.Here again, the exact pattern varies, but it tered developmental forms are summarized below
generally involves intralecithal nuclear divisions fol- (also see Table 21.2 and Figure 21..33),but we do not
lowed by nuclear migration to the periphery of the em- attempt to describe them all
bryo and subsequent partitioning of the nuclei into a Crustacean development is sometimes described as
cell layer around a central yolky mass (Figure 21'.33C). being either epimorphic, metamorphic, or anamorphic.
The form of the blastula and the method of gastrula- Flowever, we caution readers that a clear evolution-
tion are dependent primarily on the preceding cleav- ary and functional understanding of crustacean de-
age pattern and hence ultimately on the amount of velopmental stages is still lacking, and thus the terms
yolk. Holoblastic cleavage may lead to a coeloblastula "rrtixed" and "direct" may be preferable, and less am-
that undergoes invagination (as in syncarids) or in- biguous, until we have a better understanding of this
gression (as in many copepods and some cladocerans phenomenon.
and anostracans). Other crustaceans form a stereoblas- Epimorphic development is direct; in crustaceans it
tula followed by epibolic gastrulation (e.g., cirripedes). is thought to result from a delay in the hatching of the
Most cases of meroblastic cleavage result in a periblas- embryo, which causes the nauplius (and any other pos-
tula and the subsequent formation of germinal centers. sible larval stages) to be suppressed or absent.
Crustaceans share a characteristic larval stage known Metamorphic development is the type of extreme
as the nauplius larva, denoted by the appearance of three mixed development seen among the Eucarida; it in-
pairs of appendage-bearing somites (Figure 21.33D).10 cludes dramatic transitions in body form from one life
In those groups having little yolk in their eggs, the nau- history stage to another. (This pattem is similar to holo-
plius is generally free living. In those species with yolky metabolous development in insects-for example, the
eggs, the nauplius stage is generally passed through as transformation of a caterpillar into a butterfly.) In gen-
part of a longer period of embryonic development (or eral, up to five distinct preadult, or larval, phases may
a long brood period), and it is sometimes referred to as be recognized among crustaceans: nauplius, meta-
an egg nauplius. Free-living nauplii are usually plank- nauplius, protozoea, zoea, and "postlarva." The zoeal
totrophic, and their release corresponds to the depletion phase shows the greatest diversity in form among the
of stored yolk. However, in a few groups of crustaceans various taxa, and especially in decapods it has been
(e.g., euphausids and dendrobranchiate shrimps), the given different names in different grouPs (e.g., acan-
nauplius exhibits lecithotrophy. thosoma, antizoea, mysis, phyllosoma, pseudozoea). It
Crustacean development is either direct, with the is common for the zoeal phase to contain a large num-
embryos hatching as juveniles that resemble miniature ber of stages, each differing only slightly from the one
adults, or mixed, with embryos brooded for a brief or preceding it.1l Regardless of name, zoea ate character-
prolonged period and then hatching as a distinct lar- ized by the presence of natatory exopods on some or all
val form. These larval forms may pass through several of the thoracic appendages and by the pleopods being
absent (or rudimentary). Use of the term "postlarva"
10It was not until is unfortunate, as these stages differ dramatically from
J. V. Thompson discovered the nauplius lawae
of barnacles in the nineteenth century that this group was finally the preceding larvae as well as from the adults; they
classified as Crustacea, a discovery that also marked the first
use of iarval features in understanding the phyiogeny of marine
invertebrates. A recent atlas of crustacean larvae (Martin et al. 11Thezoea larvae of panulirid lobsters (phyllosoma larvae) are
2014) includes historical notes on larval development in all major lar ge, bizart e-appearing creatures (Figure 21.33L), which can
crustacean groups. occur in such large numbers as to be a favorite food of tuna.
represent unique transitional stages (both morphologi- From this wealth of terms and diversity of develop-
cally and ecologically). Examples in the Decapoda in- mental sequences/ we can draw two important gener-
clude the megalopa of true crabs and the puerulus lar- alizations concerning the biology and evolution of the
vae of spiny lobsters. In this phase of development, the crustaceans. First, different developmental strategies
role of swimming has switched from the thoracopodal reflect adaptations to different lifestyles. In spite of
limbs to the appendages of the abdomen. many exceptions, we can cite the early release of disper-
Anamorphic development is a less extreme type of sal larvae by groups with limited adult mobility, such
indirect development in which the embryo hatches as a as thoracican barnacles, and by those whose resources
nauplius larva, but the adult form is achieved through may not permit production of huge quantities of yolk,
a series of gradual changes in body morphology as such as the copepods. At the other end of this adaptive
new segments and appendages are added (it is similar spectrum is the direct development of peracarids-a
in many ways to hemimetabolous development in in- major factor allowing the invasion of land by certain
sects).In other words, the postnaupliar stages gradu- isopod lineages. Between these extremes we see all de-
ally take on the adult form with succeeding molts; the grees of mixed life histories, with larvae being released
classic example of anamorphic development is often at various stages following brooding and care. Second,
said to be the Anostraca. Cephalocarida, Remipedia, because developmental stages also evolve, an analysis
many Branchiopoda, and Mystacocarida are anamor- of developmental sequences can sometimes provide
phic-the nauplius larva grows by a series of molts information about the radiation of the principal crusta-
that add new segments and appendages gradually as cean lineages. For example, the evolution of oostegites
the adult morphology appears. In many groups hatch- and of direct development combine as a unique syn-
ing is somewhat delayed, and the emergent nauplius apomorphy of the Peracarida. Similarly, the addition
larva is termed a metanauplius. The basic nauplius pos- of a unique larval form, such as the cypris larva that
sessesonly three body somites, while the metanauplius follows the nauplius in the cirripedes, can be viewed
has a few more; however, both possessonly three pairs as a unique specialization that demarcates that group
of similar-appearing appendages (which become the (Cirripedia). The cyprid either hatches as the only free-
adult antennules, antennae, and mandibles). The end of living larva, or it is the final larval stage after a series of
the naupliar/metanaupliar stage is defined by the ap- lecithotrophic or planktotrophic nauplius larval stages.
pearance of the foffth pair of functional limbs, the max- It should also be noted that the branchiopods and
illules. In copepods a postnaupliar stage called a copep- some freshwater ostracods have evolved specialized
odite (simply a small juvenile) is often recognized. ways of coping with the harsh conditions of many fresh-
The most extreme forms of metamorphic, or mixed, water environments. Parthenogenesis, for example, is
development occur in the malacostracan superorder conunon in freshwater ostracods. Other adaptations in-
Eucarida. The most complex developmental sequences clude production of special overwintering forms, usu-
are seen among the dendrobranchiate shrimps, which ally eggs or zygotes that can survive extreme cold, lack
hatch as a typical nauplius larva that eventually under- of water, or anoxic conditions. Perhaps most remark-
goes a metamorphic molt to become a protozoea larva, able in this respect are the large-bodied branchiopods
with sessile compound eyes and a full complement of (fairy shrimp, tadpole shrimp, and clam shrimp) whose
head appendages. The protozoea, after several molts, encysted embryos are capable of an extreme state of an-
becomes azoea larva, with stalked eyes and three pairs aerobic quiescence, or diapause. During these resistant
of thoracopods (as maxillipeds). The zoea eventually stages, the metabolic rate of the embryos may drop to
yields a juvenile stage (the "postlawa," a better term less than 10% of their normal rate.
for which is "decapodid") that resembles a miniature Many crustaceans have indeterminate growth,
adult, but is not sexually mature. In some other eucarid that is, they continue to molt throughout their life. In
groups (Caridea and Brachyura) the postlarva is called contrast, other species have determinate growth and
a megalopa, and in the Anomura it is often called a ceasemolting following puberty (this life history stage
glaucothoe; in both cases there are setose natatory is sometimes referred to as the terminal molt, or ter-
pleopods on some or all of the abdominal somites. In minal anecdysis). In some species, the terminal molt
other eucarids, some (or all) of these stages are absent. is sex specific; for example, in American blue crabs
Various other terms have been coined for different (Callinectessapidus)only females have a terminal molt.
(or similar) developmental stages. For example, the
modified zoeal stages of some stomatopods are called
antizoea and pseudozoea larvae, and the advanced
zoeal stage of many other malacostracans is often
Crustacean Phylogeny
called a mysis larva. In euphausids, the nauplius is fol- Countless phylogenetic studies have been published
lowed by two stages, the calyptopis and the furcilia, on the topic of crustacean evolution. General agree-
which roughly correspond to protozoea and zoea stag- ment has been reached in some areas, but despite a
es, before the juvenile morphology is attained. great deal of effort, many fundamental mysteries re-
main unsolved, including the broad structure of the Thecostraca (barnacles and their kin), and Mala-
Pancrustaca tree. The use of molecular gene sequence costraca, but challenged others, and proposed sev-
data in crustacean phylogenetics is only just beginning. eral new names and clades (e.g., Oligostraca and
\ /hile in some casesit has resolved longstanding issues Vericrustacea) that have yet to withstand the test of
or confirmed previous hypotheses, in other casesit has time.
led to still more qlrestions. There are several particu- In the 1950s,Russian biologist W. N. Beklemischev
larly problematic i3sues. What are the basalmost living and Swedish carcinologist E. Dahl independently pro-
crustaceans, what sort of body did they have, and what posed that the copepods and several related classes
are the relationships among them? What are the rela- constitute a monophyletic clade. Dahl proposed the
tionships of, and among, the taxa that were once united class Maxillopoda for these taxa, and that term has
as "maxillopodans" (the copepods, branchiurans, the- been employed often since then. Characters shared by
costracans, tantulocarids, mystacocarids, and pentas- these small taxa include the shortening of the thorax to
tomids)? Where do the Ostracoda, Cephalocarida, and six or fewer segments and of the abdomen to four or
Remipedia fit into crustacean phylogeny? \A/hat are the fewer segments, reduction of the carapace (or, in the
relationships among the Peracarida, especially of the case of ostracods and cirripedes, extreme modifica-
many orders and families of Isopoda and Amphipoda? tion of the carapace), loss of abdominal appendages,
What are the major decapod lineages and how are they and other associated changes, all thought to be tied to
related to one another? What group of crustaceans is early paedomorphic events during the larval (or post-
represented by the mysterious "y-larvae" (the Faceto- larval) stage of this lineage as it began to radiate (an
tecta), beyond the subsequent sluglike ypsigon stage idea first proposed in\942 by R. Gurney). However,
that is clearly still not an adult? And, of course, what is molecular phylogenetic studies since then have shown
the crustacean sister group to the Hexapoda? Maxillopda to be a nonmonophyletic grouping, and
Debates on crustacean phylogeny often center on the taxon has been abandoned by most modern work-
whether paddle{egged Crustacea are ancestral or de- ers. Yet two of those groups, the copepods and thecos-
rived. The paddle-legged ancestry hypothesis holds tracans, have since been reunited in one study (Oakley
that the first crustaceans had leaflike (phyllopodous) et al. 2013) that posits a clade (the Hexanauplia) based
thoracic legs that were used both for swimming and on the same number (6) of naupliar larval stages in
for suspension feeding, as seen in the living cephalo- these groups.
carids, leptostracans, and many branchiopods. Or that The monophyletic nature of the class Malacostraca
the first crustaceans had simple, paddle-like legs that has rarely been questioned. Within the Malacostraca
were used for swimming, but not for feeding; instead, are two clades: Leptostraca, which have phyllopo-
the tasks of feeding were undertaken by the cephalic dous l i mbs and seven abdomi nal som it es, and
appendages-a plan perhaps best represented among Eumalacostraca, which lack phyllopodous limbs
living crustaceans by the remipedes. However, the op- and have six abdominal segments. Whether the
posing view, that paddle-legged crustaceans are more Hoplocarida are members of the Eumalacostraca or
derived, is supported by the recent multigene studies. deserving of their own subclass of the Malacostraca is
A large molecular study suggested a highly-derived still a subject of debate, but we consider them a sepa-
sister group relationship between the two most many- rate subclass here. Hoolocarids and eumalacostracans
segmented and "wormlike" groups, the cephalocarids also have the sixth abdbminal appendages modified as
and remipedes, both being paddle-legged groups, unit- uropods (which work in conjunction with the telson as
ing them in a proposed clade (called Xenocarida) that a tail fan). Relationships among the three main eumala-
would be the sister to the hexapods (Regier et aI.2010; costracan lines (syncarids, peracarids, and eucarids),
Figure 21,.348).Another recent phylogenomic study and even within the Eucarida, are far from settled and
concluded that Remipedia are the closest extant rela- have provided zoologists with many generations of
tives of insects (Misof ela!.201.4),and some compara- lively debate.
tive neurological work also supports this view. In con- The class Branchiopoda is usually monophyletic in
trast, a traditional, morphology-based phylogeny of molecular analyses, but it is difficult to define on the
the Crustacea hypothesizes a paddleJegged ancestry, basis of unique synapomorphies because it shows such
but places the Remipedia at the base of the crustacean great morphological variation. But larval (naupliar)
tree (Figure 2I.34A). Comparative spermatological characters, such as the reduced and tubular first anten-
studies, on the other hand, seem to ally the remipedes na and uniramous mandibles, strongly support their
with certain former maxillipodans. Clearly, we have a monophyly, as do almost all studies employing molec-
good way to go before we will deeply understand the ular data. As is true with so many crustacean groups,
phylogeny of Crustacea. our early classifications greatly underappreciated their
The Regier et al. (2010) study, summarized here diversity, and relationships among the constituent
in Figure 27.348, strongly supported some tradition- groups are far more complex than was first thought.
ally recognized groupings, such as Branchiopoda, Apparently some branchiopods have secondarily lost
(A) . |9 |p
t\'
f)- 'b' .nY
.nlY .ti' .o
:c-
^t\ r q\ ^*oS -.{o"-a{o"
v' ,o? .o,
|
|
|
.Ae";9"
{\-{C'
'<P' "$$
""'-"
Ostracoda
11
10 Branchiura, Pentastomida, Mystacocarida
9
8 Branchiopoda
7
6 Copepoda
Thecostraca (barnacles and their kin)
Malacostraca
.'\
Lepnalocarrdal
Figure 21 .34 Two competing views of evolution within -l
the Crustacea. (A) A traditional view of crustacean rela- Xenocarida
tionships, with paddle-legged Crustacea at the base. The I
Remipedia )
morphological synapomorphies depicted on the clado-
gram are as follows (1-5 represent synapomorphies of
the subphylum "Crustacea"): 1: head composed of 4 or 5
fused segments (plus acron) with 2 pairs of antennae and
2 or 3 pairs of mouth appendages; 2: biramous second
antennae; 3: nauplius lawa; 4: phyllopodous body limbs with maxillopodan naupliar eye; 23'. thorax of 6 or fewer
(with large epipods); 5: with head shield or small cara- segments; 24: abdomen of 4 or fewer segments; 25:
pace; 6: raptorial mouth appendages; 7: mouth append- genital appendages on the first abdominal somite (associ-
ages situated in posteriorly directed atrium; 8: anterior ated with male gonopores); 26: B-segmented thorax and
thoracopods (one or more pairs) modified as maxillipeds 7-segmented abdomen (plus telson); 27: male gonopores
(a highly variable trait that occurs in remipedes, malacos- fixed on thoracomere 8/females on thoracomere 6; 28:
tracans, and some former "maxillopodans"); 9: loss of carapace forms large "folded" structure enclosing most of
phyllopodous condition on trunk appendages; '10: trunk body; 29: abdomen reduced to 6 segments (plus telson);
appendages oriented laterally;11: maxillulesfunction as 30: last abdominal appendages modified as uropods and
hypodermic fangs; .12: postcephalic trunk regionalized as forming tail fan with telson; 31: caridoid tail flip locomo-
thorax and abdomen; 13: loss of internal organ homon- tion (escape reaction); 32: thoracopods with stenopodous
omy (e.9., segmental gut ceca); '14:reduction in number endopods; 33: replacement of thoracic suspension feed-
of body segments; 15: reduction of abdomen (to 11 seg- ing and phyllopodous thoracic limbs with cephalic feeding
ments); 16: fully developed carapace (reduced in several and nonphyllopodous thoracic limbs. Note that loss of
subsequent lineages); 17: reduction of abdomen to fewer the phyllopodous trunk limbs (character 9) has occurred
than 9 segments; 18: reduction (or loss) of abdominal several times, in the Remipedia, Eumalacostraca, some
appendages; 19: first and second maxillae reduced or Iineages of Branchiopoda, and most of the former "maxil-
lost; 20: thorax shortened to fewer than 11 segments; lopodan" groups. (B) Phylogeny of the Pancrustacea clade
21: abdomen shodened to fewer than 8 seomentsi 22i as derived by Regier et al. 20'10.
the carapace, and others have secondarily lost most or realizationrendersthe group called Crustaceapara-
all of the abdominal appendages. phyletic. The monophyletic clade containing both
That a group of crustaceans gave rise to the mega- crustaceansand insectsis most often referred to as the
diverse group of arthropods called the Hexapoda (in- Pancrustacea(seeChapter 20),but it is also sometimes
sects and their kin) is now almost universally agreed called the Tetracorrata,a name that recognizesthe
upon, based on a wide array of evidence that includes squareshapeof the ommatidia of many species.Earlier
molecular sequence data and neuroanatomy. This work suggestedthat Branchiopodawas the likely sister
group to Hexapoda, but recent researchsupports the opportunity for evolutionary experimentation.Any
origin of the Hexapoda from either the Remipedia conceivablecladogram of crustaceanphylogeny will
or a remipede-cephalocaridclade-the later group- require the acceptanceof considerablehomoplasy.
ing (Remipedia-Cephalocarida-Hexapoda) has been Fossil data (including that of the Orsten fauna,
named Miracrustacea,meaning "surprising crusta- Figure 2L.35)seemsto favor phyllopodous limbs asthe
ceans." Like the arthropods in general,crustaceans primitive condition. However, developmental studies
exhibit high levels of evolutionary parallelism and following the expressionof Distal-Iessandother devel-
convergenceand many apparentreversalsof character opmental genessuggestthat the early embryogeny of
states.This genetic flexibility is no doubt due in part to limbs is very similar among crustaceans.For example,
the nature of the segmentedbody, the serially homolo- trunk limbs always emerge as ventral, subdivided
gous appendages,and the flexibility of developmental limb buds. In phyllopodous limbs, the subdivisions
genes/which, as we have stressed,provide enormous of theselimb buds grow to becomethe endites and
Figure 21 .35 (A-E) Examples of Upper Cambrian (-510 Ma), prob-

ably 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 seg-
ments (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) Maftinssonia elongata,
SEMs of first larva and postlarval stage. (F) Ercaia minuscula, an early
Cambrian (520 Ma) crustacean from South China.
larvae are only a couple hundred microns long; adults

are about 1 mm in length). Sknraand many other Orsten
Crustacea were probably meiofaunal animals not un-
like modern marine meiofaunal crustaceans (e.g., the
mystacocarids). Dozens of Orsten microcrustacea have
so far been described (Figure 21.35).Recently, a beauti-
fully preserved fossil crustacean, Ercain minuscl;Ia, was
described from the middle Cambrian (520 Ma) of South
China. It has an untagmatized, L3-segmented trunk
with serially repeated biramous appendages and a
head with stalked eyes and five pairs of head append-
the endopod of the natatory / filtratory adult limbs. In ages, including two pairs of antennae. Ercaia minuscu-
stenopodouslimbs, the samelimb bud subdivisions la is only 24rrrm long (hence the species name), and
end up developing into the actual segmentsof the bears resemblances to both cephalocarids and maxil-
adult limb. Hence,the endites of phyllopodous limbs lopodans (Figure 21.35F).
appear to be homologousto the segmentsof the steno- Studies on the Swedish Orsten fauna (510 Ma), the
podous limbs. This discovery supports an emerging Middle Cambrian (520 Ma) Burgess Shale-like deposits
view of developmental plasticity in arthropod limbs, from around the world, and the Lower Cambrian (530
and it suggeststhat relatively simple genetic "switch- Ma) Chengjiang fossils from China have shown that
es" can account for major differencesin adult mor- Cambrian Crustacea had all the attributes of modern
phologies.Thus, it is quite plausible that stenopodous crustaceans, such as compound eyes/ an acron, distinct
limbs have evolved multiple times from phyllopodous head and trunk tagmata, at least four head append-
ancestors,and this is the scenariodepictedin the clado- ages, a carapace (or head shield), naupliar (or "head")
gram in Figure 21.344. larvae (with locomotory first antennae), and biramous
Work by Klaus Miiller and Dieter Waloszek on appendages on the second and third head somites
three-dimensionallypreservedmicroscopicarthropods (the second antennae and mandibles). We now,know
from the middle Cambrian Orsten (around 510Ma) de- that the crustaceans are an ancient group. Their fossil
posits of Swedenhas documenteda diversefauna of record dates back to the early Cambrian, or likely the
minute crustaceansand their larvae.Among them, for Ediacaran period if some arthropod fossils from ihose
example, is Skara(Figure 21..35C),a cephalocarid- or strata are viewed as Crustacea. The earliest known
mystacocarid-likecrustaceanfor which both naupliar crustacean larval stage is a fossil of a slightly ad-
larvae and adults have been recovered(the nauplius vanced nauplius (called a metanauplius) from the early
Cambrian (525 Ma) of China that in many ways re- within the Crustacea, such as the cirripedes, that were
sembles the naupliar larvae of modern cirripedes. This already becoming distinct. Depending on one's defini-
is remarkable, as it confirms not only of the great age tion of "CrtJslacea," it may even be that the first arthro-
of the Crustacea as a whole but also the age of groups pods were themselves crustaceans.
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rustaceansare one of the most popular invertebrate groups, even

amongnonbiologists,for they include someof the world's most delec-

Figure 21 .1 Diversity among the Crustacea. (A-D)

Figure 21 .1 (continued) Diversity among the Crustacea.

cer, one of the two invertebrates represented in the

INFRAORDER ANOMURAHermitcrabs,galatheid SUBORDER EPICARIDEA(e.9.,Bopyrus,Dajus,

INFRAORDER AXIIDEA Lobster shrimos SUBORDER PHREATOICIDEA (e.9.,Mesam-

ORDER AMPHIPODA Amphipods-beach hoppers, SUBCLASSNEOCOPEPODA

SUBORDER HYPERIIDEA (e.9., Cystisoma, oRDERcYcLoPolDA Cyclopoids(e.9.,Cyclopina,

SUBORDER CAPRELLIDEA (e.9., Caprella, ORDERGELYELLOIDAGelyelloids(e.9.,Gelyella)

SUBCLASS ASCOTHORACIDA Two orders (Laurida, OR D E R MOR MON ILLOID A Mo r m onilloids.

SUPERORDERTHORACICA Truebarnacles. Fouror- OR D E R S IP H ON OS TOMA TOIDASiPhO NO SI O -

SUPERORDER ACROTHORACICA "bar-

zone to depths of over 1,500m, from the North and

Egg pouch of eleventh

Figure 21 .4 Anatomy and diversity in the class

Adductor Ovary Maxillary

Antennule First pleopod

primitive malacostracancondition (6-8-7body seg-

reaction. Antennal scale

(B) Compound eye

Pleopods Eighth/ Penis / o#".fhs

Figure 21 -7 External anatomy in the class Malacostraca, subclass

carapacecovering the thorax, a protruding rostrum, Subclass Hoplocarida

Figure 21 .8 Anatomy in the class Malacostraca, super-

Figure 21 .9 External anatomy and diver-

InfraorderCaridea The nearly 3,500livingspeciesin associatedwith shallow benthic environments, espe-

(A) Figure 21 .10 Anatomy and diversity of the "true," or brachy-

about a dozen families) all have direct development,

Infraorder Anomura This group includes hermit

Figure 21 .12 Anatomy and diversity in some peracarid crus-

Pleopods (B) Thoracomeres3-8

Figure 21 .13 More peracarids. (4 The spelaeogriphacean,

pleopods biramous and well developed, natatory and

Figure 21 .14 More peracarids:

Figure 21 .15 And still more peracarids: amphipod diversity'

(e.9.,Gnathiidae)parasitic.The suborderOniscideain- Order Amphipoda Carapaceabsent;first thoraco-

uniramous; abdomen without appendagesbut with Class Branchiura

Figure 21 .17 Anatomy in the

Figure 21 .18 Anatomy of the classes Mystacocarida,

midventral surfaceof last thoracic somite;paired, ses-

Figure 21 .19 Anatomy and diversity in

(A) Figure 21 .2O Anatomy and diversity in the class

Fifth limb (male)

(Continued on next page)

Figure 21 .20 (continued) Anatomy and diver-

TABLE 21.1 Comparison of Distinguishing Features among the 11 Crustacean Classes

ClassRemipedia Cephalicshield Cephalon (6) Not phyllopodous 1 pair

ClassCephalocarida Cephalic shield Cephalon (6)Thorax (B) Phyllopodous None

ClassBranchiopod4 Carapace Cephalon (6), Phyllopodous None

ClassBranchiopod4 Carapace(bivalved, Cephalon 6,Trunk 10-32, Phyllopodous None

ClassMalacostrac4 Large,folded carapace Cephalon(6),Thorax(8), Phyllopodous None

ClassMalacostraca, Well developed Cephalon(6),Thorax(8), Not phyllopodous Five paris of thoracopods

ClassThntulocarida Cephaiic shield Cephalon (6),Thorax (6), Not phyliopodous, None

ClassBranchiura Carapacebroad, Cephalon (6),Thorax (6), Not phyllopodous None

ClassCopepoda Cephalicshield Cephalon (6),Thorax (6), Not phyllopodous; Usually 1 pair

Biramous I Biramous Absent A,11

Uniramous Biramous Absent None Common poreson protopodsof

Uniramous Uniramous Present None Q: on segment 1211,3