Key To The Ostracods 2007

2007. Cohen, Anne C., Dawn E. Peterson, and Rosalie F. Maddocks. Ostracoda, pp.
417-446, In: James T. Carlton, ed., The Light & Smith Manual: Intertidal Invertebrates
from Central California to Oregon. Fourth Edition. University of California Press,
Berkeley and Los Angeles.
GRBQ188-2777G-CH27[411-693].qxd 5/3/07 05:29 PM Page 417 Techbooks (PPG Quark)
Pleopis polyphemoides (Leuckart, 1859) (Podon polyphe- Saunders (1980), and King and Hanner (1998) if you are work-
moides). Number of exopodite setae on thoracopods I–3, II–3, ing with a species of Lepidurus, or Sassaman et al. (1997) if you
III–3, IV–2; Monterey Bay and North American West Coast have a species of Triops.
(Baker 1938, Proc. Cal. Acad. Sci. 23: 311–365).
Pleopis schmackeri (Poppe, 1889). Number of exopodite setae
on thoracopods I–4, II–4, III–4, IV–2; not yet known from the References
North American West Coast (T. Onbé, personal communication).
Belk, D. 1989. Identification of species in the conchostracan genus Eu-
Podon intermedius Liljeborg, 1853. Number of exopodite setae
limnadia by egg shell morphology. Journal of Crustacean Biology 9:
on thoracopods I–2, II–1, III–1, IV–2; also not yet known from the 115–125.
North American West Coast (T. Onbé, personal communication). Dodson, S. I., and D. G. Frey. 2001. Cladocera and other Branchiopoda,
Podon leuckarti Sars, 1862. Number of exopodite setae on pp. 849–913. In: Ecology and classification of North American fresh-
thoracopods I–1, II–1, III–1, IV–2; Northern Bering Sea, water invertebrates. 2nd ed. J. H. Thorp and A. P. Covich, eds. San
Diego: Academic Press.
Chukchi Sea, Bering Strait (Onbé et al. 1996, Proc. NIPR Symp.
Egloff, D. A., P. W. Fofonoff, and T. Onbé. 1997. Reproductive biology
Polar Biol. 9: 141–152). of marine cladocerans. Advances in Marine Biology 31: 79–167.
Pseudevadne tergestina Claus, 1877. Number of exopodite se- Eriksen, C. H., and D. Belk. 1999. Fairy shrimps of California’s puddles,
tae on thoracopods I–2, II–3, III–3, VI–1; Monterey Bay and pools, and playas. Mad River Press, 196 pp.
North American West Coast (Baker 1938, Proc. Cal. Acad. Sci. Kerfoot, W. C., and M. Lynch. 1987. Branchiopoda communities: asso-
ciations with planktivorous fish in space and time, pp. 367–378. In:
23: 311–365, as Evadne tergestina). See Uye and Onbé 1993,
Predation. Direct and indirect impacts on aquatic communities. W. C.
Bull. Plankton Soc. Japan 40: 67–69 (feeding). Kerfoot and A. Sih, eds. Hanover, NH: University Press of New England.
King, J. L., and R. Hanner. 1998. Cryptic species in a “living fossil” lin-
eage: taxonomic and phylogenetic relationships within the genus
Notostraca Lepidurus (Crustacea: Notostraca) in North America. Molecular Phy-
logenetics and Evolution 10: 23–36.
Linder, F. 1952. Contributions to the morphology and taxonomy of the
Notostracans (plate 182C) range in size from about 2 cm–10 cm.
Branchiopoda Notostraca, with special reference to the North Amer-
They have an elongate body covered dorsally at the anterior ican species. Proceedings of the United States National Museum 102:
end of the animal by a broad shieldlike carapace giving them 1–69.
a tadpolelike appearance. The head is incorporated into the Linder, F. 1959. Notostraca, pp. 572–576. In: Fresh-water biology. 2nd
anterior portion of the carapace. Their compound eyes are in- ed. W. T. Edmondson, ed. New York: John Wiley and Sons.
Lynch, J. E. 1966. Lepidurus lemmoni Holmes: a redescription with notes
ternalized and located on top of the head. Such positioning
on variation and distribution. Transactions of the American Micro-
goes along with the generally bottom–dwelling nature of scopical Society 85: 181–192.
these crustaceans. The exoskeleton of the thorax and abdomen Lynch, J. E. 1972. Lepidurus couesii Packard (Notostraca) redescribed with
appears segmented; however, these “segments” are best re- a discussion of specific characters in the genus. Crustaceana 23: 43–49.
ferred to as “body-rings” because they do not always accurately Martin, J. W. 1992. Branchiopoda, pp. 25–224. In: Microscopic anatomy
of invertebrates. Volume 9, Crustacea. F. W. Harrison and A. G.
reflect underlying segmentation (Linder 1952). The first 11 leg-
Humes, eds. New York: Wiley-Liss.
bearing body rings make up the thorax, with the genital open- Martin, J. W., and D. Belk. 1988. Review of the clam shrimp family Lyn-
ings on the eleventh. The abdomen includes both an anterior ceidae Stebbing, 1902 (Branchiopoda: Conchostraca), in the Ameri-
leg-bearing section and a posterior legless section. There is cas. Journal of Crustacean Biology 8: 451–482.
variation within each species in the number of body rings in Martin, J. W., and G. E. Davis. 2001. An Updated Classification of the
Recent Crustacea. Natural History Museum of Los Angeles County,
the abdomen and in the numbers of body rings having and
Science Series No. 39: 1–124.
not having legs. Also, the number of legs per body ring in the Mattox, N. T. 1959. Conchostraca, pp. 577–586. In: Fresh-water biology.
abdomen is variable. The abdomen ends in a telson from 2nd ed. W. T. Edmondson, ed. New York: John Wiley and Sons.
which extends a pair of long, thin, cylindrical, and multiar- Sars, G. O. 1867. Histoire naturelle des Crustacés d’eau douce Norvége.
ticulate caudal rami. This telson identifies membership in the C. Johnson, 145 pp.
Sassaman, C. 1995. Sex determination and evolution of unisexuality in
genus Triops, one of the two genera in the order Notostraca.
the Conchostraca. Hydrobiologia 212: 169–179.
An elongate platelike process extends from the dorsal edge of Sassaman, C., M. A. Simovich, and M. Fugate. 1997. Reproductive
the telson between the caudal rami in members of the other isolation and genetic differentiation in North American species of
genus, Lepidurus. Triops (Crustacea: Branchiopoda: Notostraca). Hydrobiologia 359:
Notostracans are omnivores that readily capture and eat 125–147.
fairy shrimps, small fishes, and other small animals. Notostra- Saunders, J. F. 1980. A redescription of Lepidurus bilobatus Packard (Crus-
tacea: Notostraca). Transactions of the American Microscopical Soci-
can reproduction includes obligate sexual, mixed sexual and ety 99: 179–186.
female self-fertilization, and unisexual modes. Both North Walossek, D. 1993. The Upper Cambrian Rehbachiella and the phy-
American species of Triops, T. longicaudatus Leconte, 1846, and logeny of Branchiopoda and Crustacea. Fossils and Strata 32: 1–202.
T. newberryi (Packard, 1871), represent complexes of bisexual
and unisexual populations (Sassaman et al. 1997).
Tadpole shrimps live in ephemeral inland waters ranging Ostracoda
from fresh to alkaline or even brackish. Temporary rain-pools ANNE C. COHEN, DAWN E. PETERSON, AND ROSALIE F. MADDOCKS
at the San Francisco National Wildlife Refuge and surrounding
(Plates 184–196)
areas on San Francisco Bay are home to Lepidurus packardi Si-
mon, 1886. Lepidurus packardi is protected under the U.S. En- The Ostracoda are a large and important class of small bivalved
dangered Species Act and cannot be collected without a federal crustaceans. Because of their microscopic size (0.20 mm–2 mm,
permit. It is the only notostracan so protected. rarely to 32 mm), they are not conspicuous animals, but they
Identification of notostracans to species is often a difficult are abundant and diverse in most aquatic ecosystems if sampled
process that ends in an uncertain conclusion. Use the key in correctly. There are probably at least 25,000 extant species, of
Linder (1959) and then consult Lynch (1966), Lynch (1972), which roughly 12,000 have been described (3,000 freshwater,
OSTRACODA 417
9,500 marine). Ostracoda have the best fossil record of any especially species of Cytheridae, Hemicytheridae, Lepto-
arthropod group: the carapace is usually well impregnated with cytheridae, Loxoconchidae, Microcytheridae, Paradoxostom-
calcite (calcium carbonate), so dead and molted carapaces ac- atidae and Xestoleberididae. Because of their small size and
cumulate abundantly as fossils in both modern sediments and fragility and difficulties of identification, several of these
ancient sedimentary rocks. families are greatly under-represented in the California liter-
Ostracodes are found worldwide in the ocean from intertidal ature, although doubtless they are abundant on California
to abyssal and hadal depths (7,000 m), in coastal estuaries and shores (see “References”).
marshes, in most fresh waters, and in a few terrestrial habitats. By contrast, the littoral and sublittoral faunas of southern
In tropical coral reef ecosystems and in deep-sea faunas, the California (south of Pt. Conception) include many podocopids
careful collector may expect to find several hundred species, in common with coastal Mexico and Central America. A
whereas high-latitude and stressed environments (e.g., coastal greater research effort off southern California has yielded a
marshes, salt lakes, and ephemeral ponds) may yield only a few more diverse ostracode fauna (littoral and sublittoral) with
species. Diversity in the littoral zone varies with the habitat but many podocopid species in common with coastal Mexico
may be surprisingly high; 58 species of interstitial and meio- and Central America. Many of these warmer water species are
faunal ostracodes were collected from intertidal sandy beaches also well represented in late Cenozoic fossil assemblages of
of Anglesey, North Wales; 40 podocopid species from intertidal California.
habitats of the Kurile Islands; and 20 species in a single sam- Most ostracodes are benthic, crawling over or burrowing be-
ple of intertidal mussels near Seto, Japan (see “References”). neath the sediment surface, through the interstices of shelly
In seasonally freshwater rock pools in the high intertidal to sands and gravels, over rocks and plants, or through microal-
supratidal zone and in the freshwater lens above the halocline gae. Some are demersal plankton, swimming for short dis-
of anchialine (cavern and fracture) systems, certain freshwater tances usually just above the sea floor to feed or mate (often at
(some salt-tolerant) species of Cyprididae may thrive; such night). Fewer are planktonic in the open ocean. Ostracodes in-
populations are usually parthenogenetic and are probably dis- clude detritivores, scavengers, herbivores, suspension-feeders,
persed by wind or birds. On foggy and rainy shores, the col- predators, commensals (of crustaceans, polychaetes, sponges,
lector can inspect water-holding plants, such as mosses and and echinoderms) and a single fish parasite. In turn, some are
bromeliads, and leaf litter for rare semiterrestrial ostracodes. parasitized by even more minute copepods, isopods, nema-
Although ostracodes are sometimes abundant, they are todes, and ciliates.
poorly known from the California and Oregon intertidal, Ostracodes are important food for fish and benthic inverte-
which hosts more species than are reported here. The faunas brates. Their carapace remains are consistently included in in-
of coastal estuaries, deltas, and lagoons of the accretionary ventories of gut contents of fish, although not necessarily as
coasts of North America (especially of the southern Atlantic dominant or selected components of the diet. The biotic in-
and Gulf of Mexico) have been more intensively studied than teractions of most ostracodes with other invertebrates and fish
those of erosional coasts, such as the rocky shores of central are not well documented, and collectors are urged to watch for
and northern California and Oregon. Users of this manual who evidence of such relationships.
collect from algae, mussel, and oyster beds; rocky tide pools; Some myodocopid and unrelated halocyprid ostracodes pro-
pocket beaches; seagrass; silty sand; and other appropriate duce bioluminescence to deter predators. The luciferin and lu-
habitats should expect to discover many additional unre- ciferase are exuded from glands in the upper lip (some
ported, and even some undescribed, species and more than a Cypridinidae) or from epidermal glands around the margin of
few taxonomic tangles. the carapace (some planktonic Halocyprididae). Porichthys no-
This section lists the ostracode species known from the lit- tatus, the California midshipman fish, has been shown to be
toral and immediate sublittoral (to a depth of a few meters) bioluminescent only in the southern part of its range where it
within the geographic range of this handbook (from Pt. Con- can feed on the bioluminescent California myodocopid Vargula
ception through central and northern California and Oregon). tsujii. Otherwise nonbioluminescent midshipman fish from
The list of 32 Podocopa and 10 Myodocopa includes those Puget Sound were able to bioluminesce after being fed biolu-
species reported in previous literature that are considered valid, minescent ostracodes. Additionally, a speciose Caribbean group
as well as new records for this region (specimens identified by of myodocopids has males capable of performing spectacular
the authors). Generic assignments have been updated for many species-specific bioluminescent mating displays somewhat
species, and some species are left in open nomenclature be- similar to those of fireflies.
cause they are new or not yet firmly identified.
Of the 10 myodocopid ostracodes known from our region,
three are also found both north and south of our region, three General Morphology
south only, one is a probable European introduction, and three
have not yet been reported beyond our boundaries. Some of Ostracodes can withdraw all limbs inside the enveloping bi-
the seven additional myodocopids that have been reported valved CARAPACE , the margins of which may lock to form a
from other localities on the West Coast may yet be found tight seal. (There are reports of ostracode survival after passage
within our region, particularly two reported both north and through the guts of fish. A few may survive hours or even days
south of our region (see the “List of Species”). of emersion, which has facilitated geographic dispersal to iso-
Because of the cold California Current, the intertidal lated coastal lagoons and terrestrial waters by hitchhiking on
podocopid ostracodes are closely related to those of British migratory waterfowl.) Ostracode valves are closed by CE NTRAL
Columbia, Alaska, the Aleutian Islands, the Kurile Islands, AD D UCTOR M USCLE S connecting the inner dermis of each valve
and northern Japan. Alaskan faunas are poorly known. Prob- through the enclosed limb-bearing part of the body. The valves
ably most of the 32 species described from Vancouver Island are either nudged apart by appendages or opened hydrauli-
range south into California. The intertidal faunas of rocky cally, as there is no complementary set of opening muscles,
shores at these latitudes are dominated by phytal dwellers, nor is there any functional equivalent to the bivalve mollusk
418 ARTHROPODA
ligament. The ostracode carapace varies from smooth to nal soft anatomy (testes, ovaries, and associated receptacles,
highly ornamented but never has the series of fine concen- reservoirs and pumps), external genitalia (especially complex
tric “growth lines” (indicating previous molts) present on in males), appendages (with sensory, clasping and egg-clean-
the bivalved carapaces of Conchostraca. It is perforated by ing modifications), DOM I CI LI U M (in brooders), and external
numerous, minute N OR MAL P OR E CANALS with chemosensory size, shape and ornament of the carapace (sometimes very di-
and tactile sensilla, which communicate with nerve cells in morphic). The details of sexual dimorphism are essential for
the underlying epidermis. Ecophenotypic variability of cara- identification at all taxonomic levels. One sex is sometimes eas-
pace structure is common in podocopid species populations ier to identify than the other.
of the coastal zone, especially in certain families of orna- Identification of Ostracoda has been difficult for nonspe-
mented Cytheroidea, in response to fluctuations of salinity, cialists. Dissection and examination must be performed under
substrate, organic carbon, and other poorly understood en- a light microscope. Few keys exist, and none should be relied
vironmental parameters. upon as substitutes for consulting the primary taxonomic
The exoskeleton of the soft body consists of a somewhat literature of many countries and languages. Secondary faunal
rigid head capsule (with labrum [upper lip] and MOUTH ) and a compilations exist for a few other regions, but there are none
flexible cuticular integument over the posterior region. The for marine ostracodes of North America (except papers on Myo-
endoskeleton is a loose array of internal chitinous SCLE R ITE S . docopida of Western North Atlantic).
The demarcation between the cephalon and thorax is indis- Considerable information about ostracode specialists and
tinct, and there is argument about whether it has segmental or their research can be found on the Web site of the International
functional significance. The abdomen is either missing or not Research Group on Ostracoda (www.uh.edu/rmaddock/
differentiated from the thorax; the posterior body (TR U N K ) of IRGO/cypris.html; http://userpage.fu-ber/in.de/~palaeont/irgo/
most ostracodes appears under light microscopy to be unseg- ostracoda.html), with links to other sites, such as Cypris, the
mented. In a few taxa, external transverse folds fringed with International Ostracoda Newsletter and Cohen’s tabular key
spines or small setae (most visible with scanning electron mi- (illustrated). A Web search on “Ostracoda” will turn up numer-
crography [SEM]), and sometimes an internal chitinous frame- ous other sites, many beautifully illustrated. Southern California
work, probably indicate vestigial trunk segmentation. Vestiges fauna are listed at www.scamit.org/taxonomic_tools.htm.
of as many as 10 or perhaps 11 segments (plus a telson) occur
in the Platycopida (and fewer in the podocopid Saipanetta). In
the Myodocopa, obvious dorsal trunk segmentation is found How to Collect and Prepare Ostracodes
in a few Cladocopina (up to four segments) and a few Thau- for Identification
matocypridoidea (probably more).
There are five to seven paired limbs (six to eight if copula- Look for ostracodes in most intertidal habitats—even freshwa-
tory limbs included), all more or less different in structure and ter pools in the highest zone. Use the same collecting equip-
function, and their homologies with other Crustacea are diffi- ment and methods that you would use for other tiny
cult to trace. Ostracoda never have six or more serially repeated crustaceans. Collect samples of algae, pitted rocks, oysters, or
pairs of trunk limbs, as in the superficially similar barnacle mussels into buckets or stout plastic bags and then extract the
“cypris” larvae or in Conchostraca. The male and female ex- hidden tiny animals. Examine these samples directly under a
ternal genitalia are thought, at least in some cases (particularly dissecting microscope and remove the ostracodes with fine for-
Myodocopa), to be a modified last (usually eighth) pair of ceps (the flexible ones are good) or an eyedropper (for swim-
limbs. Certain perhaps sensory structures have also been in- ming ones).
terpreted as the remnants of limbs. The body terminates in a For stubbornly hidden ostracodes, a wash may help. Try
pair of caudal FU RCAE , which in Myodocopa is posterior to the adding some fresh water or a little alcohol to the seawater and
anus but in Podocopa is anterior to the anus. shaking or soaking the algae (etc.) until more animals emerge.
Ostracodes have no larval stage and no metamorphosis but You can wash and strain the sample water through a fine-
hatch from the egg as a juvenile with at least three functional meshed net or screen (holes not more than 0.5 mm for myo-
limbs and the enveloping bivalved carapace (less calcified than docopids, as small as 0.02 mm for the tiniest interstitial
adult). Juveniles resemble reduced adults. During a fixed series podocopids) and flush the inverted contents with a squirt bot-
(usually five to eight) of molts, they grow larger and acquire tle into a small dish for observation. Live ones often play pos-
additional limbs; limbs also become more complex, more se- sum at first but are fun to watch when they open their valves
tose, and, in some cases, change function. Carapace details and and move around after a few minutes. Picking and observing
primary and secondary sexual characteristics may begin to take lively ostracodes from a sorting dish may be easier if you slow
shape in late instars but are not fully developed until the adult them down by adding a little anesthetic such as MgCl2 (50%
(terminal) instar. Because molting requires abandonment of 0.36M) for a few minutes; revive them by returning them to
the calcified part of the carapace cuticle, the hinge structures plain seawater.
and marginal details are fully generated only during the final To hunt for ostracodes in bottom sediment and seagrass,
molt of Podocopa. Fossil assemblages in modern and ancient drag a fine-meshed net (or homemade hand dredge) along the
sediments include numerous molted valves and carapaces in surface of the bottom, not digging in more than just a few cen-
addition to adult specimens. Juveniles are difficult to identify, timeters. Cohen’s favorite dredge is one that Todd Oakley made
so collectors should strive to recover large populations with by attaching pantyhose to a frame. Flush seawater through the
complete growth series and adults of both sexes. net (small running seawater hoses work) to remove fine sedi-
Sexes are separate, and fertilization is internal. Partheno- ment before flushing net contents into a sorting dish and pick-
genesis is common in some freshwater lineages but has not ing ostracodes from the coarser sediment (stirring the sediment
been conclusively verified in marine dwellers. Many ingenious and quickly decanting mainly liquid may produce many but
adaptations have evolved to enable copulation in spite of the not all of the ostracodes). Some myodocopids are attracted to
enveloping carapace. Sexual dimorphism occurs in the inter- traps baited with a little fish or a light stick and anchored
OSTRACODA 419
overnight. Some male myodocopids are attracted to a bright some molecular analyses suggest that they may actually not
light suspended from a dark dock, and they then can be form a monophyletic clade. The classification of Podocopa re-
scooped up with nets. Preserve and store your labeled ostracode mains more uneven than that of Myodocopa.
samples in 70% EtOH until you are ready to identify them.
Take field notes.
Start the identification process by examining a sample un- Glossary
der a dissecting microscope and sorting the ostracodes into
groups of similar ones. Gently transfer an ostracode into a lit- This list contains abbreviations used in the plates and gives il-
tle glycerol in a labeled depression slide (use alcohol instead if lustrative examples from among the plates.
you are preparing for SEM or molecular studies) and examine Terms used herein (and in many other ostracode publica-
it under a microscope. Fine flexible forceps, an eyedropper tions) are defined (some with figure references).
(don’t use much liquid), mouth pipette, brush, needles, or tiny
wire loop (using a fine probe in your other hand as a pusher) 1 ST ANTE N NA , ANTE N N U LA , A 1 1st (cephalic) pair of limbs.
are useful. Examine the ostracode under a compound micro- Plates 184, 190A, 191D, 191E.
scope and perhaps sketch it (camera lucida helpful). Use a 2 N D ANTE N NA , ANTE N NA , A 2 2nd (cephalic) pair of limbs.
coverslip for higher powers. With the slide under the dis- Plates 184, 191B, 191D, 191E.
secting microscope, remove one or both carapace valves. Hold 3 R D ( CE PHALIC ) LI M B see “mandible (md).” Plate 184.
the ostracode in place with one tool (try the tiny wire loop) 4 TH LI M B (1 ST MAXI LLA , MAXI LLU LA , 4) 4th (cephalic) limb,
while you slide a sharp tool (fine dissecting needle or single labeled “maxilla” in many papers; use care in reading literature
edged razor blade) between the valve and the body to sever descriptions. Plates 184, 192D, 193F.
the central adductor muscle. Pry the valve off the body and 5 TH LI M B ( 2 N D MAXI LLA , 5 ) 5th limb, labeled “maxilla” in
cut the weak dorsal body connection to the valves. Examine many papers; controversially considered to be and labeled “1st
the specimen again under the compound microscope. You trunk leg,” “1st thoracic limb,” or “maxilliped” by some
may be able to identify the subclass or order of ostracode at podocopid specialists; use care in reading literature descrip-
this point. tions. Plates 184, 191D, 191E, 192F.
If you are unfamiliar with ostracodes, sketch the limb po- 6 TH LI M B (6) 6th limb (and 1st trunk/thoracic limb); contro-
sitions and shapes before removing limbs (with fine needles) versially considered to be and labeled “2nd trunk leg” by some
under the dissecting microscope. Cohen uses insect minuten podocopid specialists; use care in reading descriptions. Plates
pins glued inside the tip of narrow wooden sticks or fine 184, 191D, 191E, 192G.
pipette tips. Retain the valves and body in the glycerin slide, 7 TH LI M B (7) 7th limb (and 2nd trunk/thoracic limb); con-
but with a needle move each limb to a permanent slide; cover troversially considered to be and labeled “3rd trunk limb” by
both slides with a coverslip. Cohen makes permanent slides some podocopid specialists; use care in reading descriptions.
with polyvinyl lactophenol (this is very poisonous, so use Plates 184, 191D, 191E, 192H.
care). When the limb slide is fully dry, the limbs can be ex- 8 TH LI M B (8) see “copulatory limb (CP).”
amined and drawn under high power with the compound AD D UCTOR M USCLE SCAR PATTE R N ( AM S ) rather central pat-
microscope. tern of scars (raised or depressed) indicating where central ad-
The valves of myodocopans will weaken only somewhat ductor muscles attach to valves; not always clearly visible.
over time in glycerin. Both valves and dissected limbs can also Plates 189A–189G, 189I, 189J, 190K, 190L.
be stored in a vial of alcohol stoppered with cotton and placed ARTICLE see “podomere.”
in a larger vial that also contains a permanently inked label. BAS I S distal part of divided protopod (separated from coxa
Cotton stoppered vials should be stored in a larger and well- by suture). Plates 185C, 188A–188D.
sealed jar of alcohol, perhaps with labeled vials of similar os- B E LLONCI ORGAN ( BO ) organ projecting from forehead or
tracodes. However, the valves of podocopans will decalcify in naupliar (medial) eye of most Myodocopa. Plates 184B,
alcohol and glycerin. For storage, they should be dried on pa- 185B.
per micropaleontological slides. SEM reveals many important B RANCH IAL PLATE ( VI B RATORY PLATE , B P ) flat setose epipod,
minute characters (particularly of the carapace and copulatory (at least in Myodocopa) used to circulate water through the
limbs) not otherwise visible (see “References”). Additional domicilium. Plates 184C, 184D, 192D.
voucher specimens (preserved in alcohol) of species used for B ROOD POUCH an expanded region of the female domicil-
SEM, molecular and ecological studies should always be re- ium, usually posterior, for protecting developing eggs and in-
tained and deposited in a museum collection as verification of stars.
your identifications. B R US H - S HAPE D ORGAN S small pair of lobes bearing numer-
ous fine setae; found between the fifth limbs in males of most
Cytherocopina and some Cypridocopina; might represent ves-
Classification of Living Ostracoda tigial appendages. Plates 190F, 194D.
CALCI FI E D I N N E R LAM E LLA ( CI L ) calcified part of the inner
Early classifications of living Ostracoda were based mostly on (medial) lamella of valve margin. Plate 189F.
soft anatomy, whereas classifications of fossils were based on CARAPACE lateral outfolds of dorsal epithelium and cuticle,
carapace features. Older classifications are obsolete, and none bivalved and enveloping the entire body. Plates 184D, 187,
of the newer syntheses is universally regarded as authoritative. 195, 196.
Since the mid-twentieth century, in an effort to reconcile the CAU DAL PROCE SS ( KE E L , K ) posterior extension of carapace.
discrepancies, some paleontologists began to study living fau- Plates 184A, 186G, 186H.
nas, and the advent of SEM revolutionized the description of COPU LATORY LI M B S ( CP ) male limbs modified for copulation,
carapace and appendage structure. Myodocopa and Podocopa at least in most Myodocopa, usually the 8th limb pair (see also
differ noticeably from each other (see tabular key below), and “hemipenes”). Plate 184A.
420 ARTHROPODA
COXA proximal part of divided protopod (separated from I N N E R MARG I N ( I M ) the inner or proximal edge of the calci-
“basis” by suture; see “protopod”). Plates 185C, 188A–188D. fied inner carapace lamella, forming an abrupt line or shelf in
DOM ICI LI U M the volume between the two valves of the cara- dead or fossil valves of Podocopida. Plates 189A, 189H, 190L.
pace and occupied by the soft body and limbs. KE E L ( K ) posterior extension of carapace (also called “caudal
ENDITE medial extension (tooth, lobe, masticatory process) of process”). Plates 184A, 184B, 185A, 186G, 186H.
protopod, generally used in feeding (see also “masticatory LATE RAL EYE ( LE ) paired compound eyes of most Myo-
process”). Plate 188A. docopida. Plates 184A, 184B, 185A, 185F.
E N DOPOD / E N DOPOD ITE ( E N ) in distally biramous crustacean LI N E OF CONCR E SCE NCE ( LC ) the proximal edge of the zone
limb, the medial ramus. Plates 184C, 184D, 185C, 190B. of concrescence. Plates 189I, 190L.
E PI POD ( E P ) extension (usually setose flat lobe) of lateral pro- MAN D I B LE third limb (third head limb). Plates 184A–184D,
topod; in ostracodes (at least in Myodocopa) it often forms a 192E, 193G.
branchial plate used to circulate water. Plate 184A, 184B. MAXI LLU LA , “ MAXI LLA” see “4th and 5th limbs.”
E XOP OD / E XOP OD ITE ( E X ) in distally biramous crustacean MASTICATORY PROCE SS see “endite.”
limb, the lateral ramus. Plates 184, 185C, 190B. NAU PLIAR EYE ( M E D IAL EYE , N E ) single medial anterior eye,
FR E E MARG I N the perimeter of the valves (exclusive of the normally composed of three eye cups, has reflecting tapetal
hinge region) where valves are in contact. layer. Plates 184, 185B.
FU RCA ( E ) ( F ) terminal (may be ventral in position) body ex- NOR MAL POR E CANALS ( N PC ) numerous perforations through
tensions. Plates 184, 185D, 185H, 186F, 188E, 191E. valve with chemosensory and tactile sensilla, which com-
FUS E D ZON E see “zone of concrescence.” municate with nerve cells in the underlying epidermis; may
H E M I PE N E S ( H P ) podocopan term for paired male copulatory be simple (open) or covered with sieve plates. Plate 189I.
appendages. Plates 184C, 191D, 193H. OUTE R LAM E LLA the outer layer of epidermis and cuticle of
H I NG E ( H ) complementary interlocking articulatory struc- the dorsal body fold forming the valve; may be calcified or un-
tures on inner dorsal margin of valves. Basic types mentioned calcified.
in key: PALP term used for a podocopan armlike jointed portion (endo-
AD ONT single element hinge, bar on one valve fits into pod or exopod plus basis) of limb, either directed forward and used
groove of other valve for food manipulation, or directed backward and used for clasping.
MERODONT three-element hinge, with anterior and posterior PODOM E R E ( ARTICLE ) segment of jointed crustacean limb
teeth (or sockets) separated by median bar (or groove) separated from other segments by articulated joint or inflexi-
LOPHODONT a merodont hinge in which all elements are ble suture.
smooth PROTOPOD ( PROTOPOD ITE , PRO ) basal podomere(s) of crus-
PENTODONT a five element hinge, in which the anterior and tacean limb (plate 184B), often divided into two (sometimes
posterior region of the median element are differenti- three) segments: the coxa attaching the limb to the body and
ated, either crenulate or smooth (family Pectocytheridae the more distal basis. Plate 188A, 188B.
only) RADIAL PORE CANALS ( RPC ) modified normal pores located at
HOLOM E RODONT a merodont hinge in which all elements are free margins of valves, housing nerve filaments that lead from tac-
crenulate and negative on one valve, positive on the other tile sensilla to nerve cells in the underlying epidermis. Plate 189H.
ANTI M E RODONT a merodont hinge in which the median el- ROSTR U M anterior projection of valves of many myodoco-
ement is reversed (negative or positive) from the termi- pans, overhangs incisur. Plates 184A, 184B, 186E, 186G.
nal elements S CLE R ITE internal chitinous struts supporting body and
E NTOMODONT a four-element hinge, with the anterior region limbs. Plate 188C.
of the median bar or groove differentiated, usually with S E M scanning electron micrography. Plates 195, 196.
all elements crenulate S PI N N E R ET G LAN D ( SG ) gland (connected to 2nd antenna) se-
GONGYLODONT a crenulate hinge with a systematic increase creting adhesive thread. Plates 184D, 190B, 191D.
in size of teeth and decrease in size of sockets from an- VALVE right or left half of bivalved ostracode carapace (some-
terior to posterior in the right valve, left valve comple- times erroneously referred to as “shell”).
mentary (family Loxoconchidae only) VE STI B U LE ( VE STI B U LU M , V ) in Podocopa, a marginal cavity
AM PH I DONT a modified entomodont hinge in which the between the outer valve lamella and calcified part of the inner
four elements are well differentiated (families Hemi- valve lamella, extending from the line of concrescence to the
cytheridae, Trachyleberididae) inner margin, housing epidermal, glandular, and sometimes re-
HOLAM PH I DONT an amphidont hinge in which all elements productive tissues. Plates 189I, 190L.
are smooth VI B RATORY PLATE ( B RANCH IAL PLATE , B P ) flat plate (with mar-
H E M IAM PH I DONT an amphidont hinge in which the poste- ginal setae) located proximally and laterally on limb (an epi-
rior element is crenulate and the others are smooth pod in Myodocopa, but podocopan homologies uncertain);
SCH I ZODONT entomodont hinge in which the anterior and used to circulate water. Plates 184C, 184D,192D.
anteromedian elements are bifid, and the posterior ele- “ WALK I N G LE G S ” posterior limbs that are long, slender,
ment is bifid or crenulate (Family Schizocytheridae only). jointed, directed ventrally, and used for locomotion. Plates
Plate 191F. 184C, 184D, 191D, 191E.
I NCI SU R ( I NCI SU R E , ROSTRAL I NCI SU R , I ) an anterior inden- Z E N K E R ’ S OR GAN ( Z ) muscular sperm pump (ejaculatory
tation (ranging from slight to deep and even slitlike) of the pump) located dorsally within male body (Cypridocopina) or
valves of the carapace; it acts like an oar-lock for the rowing interlamellar cavity (Sigilliocopina).
antennae of many swimming Myodocopa. Plates 184A, 186G. ZON E OF CONCR E SCE NCE ( FUS E D ZON E , ZC ) a clear marginal
I N N E R LAM E LLA inner layer of epidermis and cuticle of the band extending from the free margin to the line of concrescence,
dorsal body fold forming the valve, entirely uncalcified or cal- formed by a flangelike outgrowth of the outer lamella and tra-
cified only around the free margin. versed by radial pore canals.
OSTRACODA 421
P LATE 18 4 Ostracoda, diagrammatic comparison of Myodocopa and Podocopa. Myodocopida (Cypridinidae) after Morin 1986: A, male;
B, female; Podocopida (Cytheroidea), after Athersuch, et al. 1989: C, male; D, female (redrawn by Ginny Allen).
Key to the Ostracoda — Carapace ovoid, inflated-subtriangular, oblong elongate,

or compressed; no rostrum or incisur; valves overlap around
TABULAR KEYS TO MYODOCOPA free margin; 2nd antenna geniculate, pediform; with very
small exopod with no more than two podomeres; much
ANNE C. COHEN
larger propulsive endopod with up to four podomeres; vari-
The three tabular keys on the following pages cover all Myo- able furca anterior to anus; no lateral eyes or Bellonci organ
docopa from subclass to the family level (along with just the (plates 184C, 184D, 189–196) . . . . . . Subclass Podocopa 2
subclass Podocopa). A dichotomous key by Rosalie Maddocks 2. Carapace size, shape diverse; left (sometimes right) valve
further identifies Podocopa from orders to families (along with usually overlaps right valve. Adductor muscle scar pattern
just the subclass Myodocopa). Because of the chance of en- varied (but includes rows) (plates 189A, 190K, 191D); seven
countering unreported species, taxonomic literature should pairs of limbs (including at least one walking leg) (plates
be consulted if identification is attempted beyond the family 184C, 184D, 191D, 191E) plus large male hemipenes
level. (plates 184C, 191D, 192I, 193H, 194H); 2nd antennal ex-
opod greatly reduced (plates 184C, 184D, 190B); 4th limb
(and usually mandible) without lateral setal comb (plates
184C, 184D, 192D, 192E), furcae mostly reduced (plates
Key to Podocopa (and Subclass only of Myodocopa)
184C, 184D, 191E), posterior body only rarely transversely
ROSALIE F. MADDOCKS
ridged (plates 184C, 184D, 189–196) . . . . . . . . . . . . . . . . .
1. Carapace elongate oblong to almost circular; with or with- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Order Podocopida 3
out rostrum and incisur; valves usually not overlapping; — Carapace oblong, laterally compressed, right valve over-
2nd antenna with large muscular protopod; long exopod lapping left all around; adductor muscle scar pattern bise-
with nine podomeres (rarely less) with long setae (often rial; six pairs of limbs (no walking legs) plus male large
used for swimming); smaller (often much smaller) endo- hemipenes; 2nd antenna with large setose exopod (two
pod with one to three podomeres, usually conspicuously podomeres); 4th limb (and mandible) with setal combs;
dimorphic in adults; large lamellar furcae posterior to anus; furcae large, lamellar, with stout claws; posterior body
with or without lateral eyes and Bellonci organ (plates transversely ridged; marine, not intertidal . . . . . . . . . . . . .
184A, 184B, 185–188) . . . . . . . . . . . . Subclass Myodocopa . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Order Platycopida
422 ARTHROPODA
Key to the Suborders of Podocopida identifiable; marine and nonmarine; global, extremely
abundant and diverse in all aquatic habitats, accounting
3. Carapace ovoid to elongate, usually smooth (or punctate); for more than 90% of all living Ostracoda, especially dom-
adductor muscle scars an aggregate or rosette of numerous inant in brackish water, intertidal and shallow sublittoral
scars; calcified inner lamella narrow . . . . . . . . . . . . . . . . 4 faunas (plates 184C, 184D, 190–194, 195C–195G, 195I,
— Carapace form and ornament diverse; adductor muscle scars 195J, 196) . . . . . . . . . . . . . . . . . Suborder Cytherocopina 9
varied, but distinctive arrangements of a smaller number of
discrete scars (see descriptions further in key); calcified in-
ner lamella relatively broad (plate 189F) . . . . . . . . . . . . . . 5 Key to the Superfamilies of Cypridocopina
4. Carapace elongate-ovoid, symmetrical; right valve overlaps
left; anterior end narrower, posterior inflated as brood cham- 7. Carapace elongate; right valve overlaps left; hinge sinu-
ber; adductor muscle scar pattern of six to 12 in compact ous, consisting of five elements, terminal elements
rosette; hinge adont; mandibular palp with lateral setal crenulated; adductor muscle scar pattern a compact cir-
comb; posterior body smooth; furcae generally absent; males cular spot with three discrete upper scars and a lower
rare; freshwater to slightly oligohaline; global . . . . . . . . . . . group of indistinct scars; female 5th limb jointed; 6th
. . . . . . . . . . . . . . . . . . . . . . . . . Suborder Darwinulocopina limb with one long and two shorter terminal claws; testes
— Carapace inflated-ovoid, asymmetrical; left valve strongly and ovaries do not enter interlamellar valve cavity;
overlapping right; adductor muscle scar pattern an ag- Zenker’s organ with numerous chitinous spikes not
gregate of 20–35 scars; hinge merodont; mandible with- arranged in rosettes. One to two balls of sediment visi-
out setal comb; posterior body transversely ridged; large ble in hindgut. Global, marine benthos, represented in
lamellar furca with fairly stout claws; males with spirally all sublittoral and deeper habitats but usually uncom-
muscularized Zenker’s organs; marine, always rare, cir- mon . . . . . . . . . . . . . . . . . . Superfamily Macrocypridoidea
cumtropical in reef habitats, probably cryptic and inter- — Carapace shape and overlap variable; hinge adont; adduc-
stitial . . . . . . . . . . . . . . . . . . . . . . . Suborder Sigilliocopina tor muscle scar pattern of five to six discrete scars (anterior
5. Carapace diverse, usually smooth (or slightly punctate) (plate column of three scars and slightly lower posterior column
195A, 195B); hinge usually adont (otherwise macrocypridid of two to three scars); female 5th limb unjointed, 6th limb
described in couplet 7a); adductor muscle scar pattern of with one terminal claw; ovaries and testes or vas deferens
five to nine discrete scars in a distinctive arrangement (plate partly housed within interlamellar valve cavity; Zenker’s
189); 5th limb endopod a small walking leg, or reduced in organ without chitinous spikes or with spikes arranged in
females and modified as a recurved clasper in males; 7th rosettes; gut contents unrecognizable . . . . . . . . . . . . . . . 8
limb recurved over posterior body, modified distally for 8. Carapace shape and overlap varied; usually smooth; ad-
cleaning; some males with brush-shaped organ; testes and ductor muscle scar pattern of five scars in two columns;
ovaries sometimes extend into interlamellar cavity of valves; mandible with weak simple teeth; 7th limb with three short
Zenker’s organs in dorsal region of male body; many marine terminal setae, none recurved, no pincer; Zenker’s organ
and most freshwater ostracodes, some parthenogenetic without chitinous spikes; global, marine; swimming, crawl-
(plates 189, 195A, 195B) . . . . . . . Suborder Cypridocopina 7 ing and burrowing; represented in all sublittoral and deeper
— Carapace shape varied, often ornate, with complex hinges habitats . . . . . . . . . . . . . . . . Superfamily Pontocypridoidea
and marginal details; adductor muscle scar pattern of four — Carapace varied, usually smooth (but not always; e.g., plate
to nine discrete scars in distinctive arrangement (see next 195A, 195B); left valve usually slightly overlaps right; ad-
couplets); 5th, 6th, and 7th limbs are walking legs (plates ductor muscle scar pattern (plate 189) of one cap (dorsal)
184C, 184D, 191D, 191E); paired brush-shaped organ in scar plus about five scars in two columns; mandible with
males (plate 190F); testes and ovaries not extending into strong trifid teeth; 7th limb with long recurved pectinate
interlamellar cavity of valves; no Zenker’s organs (plates seta and one to two short or long terminal setae or mod-
184C, 184D, 190–194, 195C–195J, 196). . . . . . . . . . . . . . 6 ified as pincer; Zenker’s organ with chitinous spikes usu-
6. Carapace with characteristic bairdioid shape (plate 195H), ally grouped in rosettes; global, highly diverse; marine,
smooth or punctate; left valve overlaps right; adductor brackish and freshwater, found in all aquatic habitats,
muscle scar pattern of three pairs of scars in anterior ver- dominant in terrestrial waters; also well represented in lit-
tical column plus posterior pair of scars; 2nd antennal ex- toral and shallow-sublittoral habitats of tropical regions,
opod a minute scale with up to three small setae; furcae coastal lagoons, and estuaries, and in anchialine cave
small, rodlike with small setae; one to two balls of sedi- faunas (plates 189, 195A, 195B) . . . . . . . . . . . . . . . . . . . . . .
ment usually visible in dorsal region of midgut; marine, . . . . . . . . . . Superfamily Cypridoidea, Family Cyprididae 9
global, especially diverse in reef habitats but represented
in most subtidal faunas from all depths and sediment
types . . . . . . . . . . . . . . . . . . . . . . . Suborder Bairdiocopina Key to the Families of Cytheroidea Known to Be
— Carapace extremely varied; left valve usually slightly over- Present in the California and Oregon Littoral
laps right; adductor muscle scar pattern typically a vertical
row of four (rarely, five) small scars (rarely some of the scars This key is valid only for the species included in the species list
divided) (plates 190K, 190L, 191A, 191D–191F, 192J, 193J, of this section.
193K, 194I–194K); hinge adont, merodont, amphidont,
schizodont (plate 191F), or other specialized condition; 9.Carapace smooth or very faintly pitted (plates 190K, 190L,
sexual dimorphism may be conspicuous; 2nd antennal ex- 192J, 193J, 193K, 194J, 194K) . . . . . . . . . . . . . . . . . . . . . 10
opod modified as a hollow spinning seta connected to — Carapace conspicuously (except Acuminocythere, plate
spinneret gland (plates 184D, 190B, 191D); furcae reduced 196C) pitted, ridged, or otherwise ornamented (plates
to two tiny setae (plate 191E); gut contents not usually 195C–195G, 195I, 195J, 196) . . . . . . . . . . . . . . . . . . . . . 14
OSTRACODA 423
TA B L E 2
Tabular Key to Myodocopa (and Subclass only of Podocopa)
Subclass Subclass
Character Podocopa Myodocopa Order Myodocopida Order Halocyprida
Carapace (valve)
Adult length 0.1 mm–8 mm 0.1 mm–32 mm Most 1 mm–3 (but up to 0.1 mm–3 mm
(approximate) 32) mm
Shape (lateral outline) Ovoid, inflated- Elongate oblong to Elongate-ovoid or ovoid, Varied, elongate sub-
(plates 184–187, subtriangular, oblong or almost circular few rather circular quadrate to almost
189–196) elongate, not circular circular
Anterior rostrum (plates Without (projection in With or without Most Many, but not most
184–187) some Cypridoidea)
Anterior incisur (notch) Without With or without Most with (usually not With (often above mid-
(plates 184, 190–194) above midheight), but height) or without
not obvious on more (more circular valves)
circular valves
Ventral margin Straight, sinuous (partly Usually convex, some
concave) or slightly straight, none concave
convex
Posterior keel (project- n.g. Yes or no Present in many None, except some with
ing caudal process) posterodorsal corner-
(plates 184–186) shaped process or spine
Anterior inner valve n.g. Setose or not With at least 1 seta Without setae
surface
Body (plate 184)
Medial eye With or without With or without Present in most Absent

Iateral eye Absent Present or absent Present in most Absent
Bellonci organ Absent Usually present Usually present Usually present
Bifurcate? Bifurcate or not Not bifurcate Many bifurcate
Posterodorsal body Few with some Few with some Body smooth; some A few with some
segmentation segmentation with gills or pouches segmentation; no gills
or pouches
Furca (plates 184, 185D, 185H, 186F, 188E, 191E)
Furcal position Anterior to anus Posterior to anus

Furcal shape Varied, most not plate- Always strongly
like with claws, (no sclerotized flat plate
strongly sclerotized with claws
plates)
No. of Limbs (including 7–8 (plus rudimentary 6–8 8 6–8
male copulatory limb) brush organ in many)
2nd Antenna
Best developed Endopod (plates Exopod (plate 184)

ramus 191D, 191E, 192B
Number of endopod 3–4 1–3 1–3 2–3
articles
Male endopod No clasper, but some Most with clasper Many with clasper Many with hooklike
dimorphic (usually reflexed) clasper
Number of exopod 1–2 (or seta only) 9 (rarely less) 9 (rarely less) 9
articles
TA B L E 2 (continued)
Subclass Subclass
Mandible (plates 185C, 188A–188D, 190G, 192E, 193G, 194B)
Sexual dimorphism n.g. Some Dimorphic in Sarsielloidea Apparently none

Proximal setose Many Podocopida None
branchial plate
Lateral setose comb Platycopida only None
Proximal endite n.g. Usually present, varied Usually present and Present as a cusped
(on coxa) lobelike tooth
Basis endite n.g. Present in some, varied 1 family with lobe, none Many with cusped
with large tooth tooth
Endopod: 1st article n.g. V and/or D setae V setae only D (and sometimes V)
setal position setae
Exopod n.g. Present in some = lobe Present in most Present (1 article) or
distally inserted on (1 article) or absent absent
basis with endopod
4th Limb (plates 184, 190D, 190E, 192D, 193E)
Sexual dimorphism n.g. Present in some Reduced only in Apparently none

Sarsielloidea males
Proximal setose Present in Podocopida, Absent (?except in Possibly bare proxi- Absent
branchial plate absent in Platycopida Cylindroleberididae) mal lobe in Cylindrole-
berididae
Lateral setose comb Platycopida only Present in some Cylindroleberididae only Probably absent
Exopod n.g. Many Usually present 1 Present (1–2 articles)
article or not
Endopod Podocopida jointed Jointed 2 articles 2–3 articles
5th Limb (plates 184, 190H, 191D, 191E, 192F, 193C, 194E)
Sexual dimorphism Often, some males with Some Limb reduced in male Rare
hooklike clasper Sarsielloidea; Cyclas-
teropinae males with
process
Proximal setose Well-developed to Present epipod
branchial plate absent
Limb shape Leglike or reduced Varied Compact feeding limb Leglike
With setose comb No Some Cylindroleberididae only No
Teeth or short claws n.g. Present or absent Many with strong non- Some with terminal
terminal teeth or claws claw (only)
Endopod Present Present Rami unclear Present and jointed
Exopod Absent present in few Rami unclear Polycopina only
6th Limb (plates 184, 190I, 191D, 191E, 193D, 194F)
Limb present Present Present or absent Present Present or absent

Sexual dimorphism Platycopida Some None or slight Some (Conchoecia)
Proximal setose Well-developed to Present in some = Absent Present in some =
branchial plate absent epipod epipod
Limb shape Leglike or reduced Varied Short, flat, poorly Leglike or absent
jointed
(continued)
Subclass Subclass
Endopod Present Present Rami unclear Present

Exopod Absent Present in a few Rami unclear Present in some
(1 article)
7th Limb (plates 190J, 191D, 191E, 192H, 193E, 194G)
Limb present Present or absent Present or absent Usually present Present or absent
Sexual dimorphism n.g. Some Limb reduced or absent None
in some males
Proximal setose Reduced if present Absent
branchial plate
Limb size n.g. Varied Usually long Reduced or absent
Limb shape Leglike, absent, or a Not leglike Long wormlike Lobe with few setae
jointed (but not annu- annulated cleaning or limb absent
lated) cleaning limb limb
Copulatory (8th) Limb or Hemipenis (plates 184, 191D, 192I, 193H, 194H)
Paired or single Paired Paired or single Paired Single

Branches n.g. Branched in many 3 lobes Single or 2 branches
NOTE : Bold major character difference; D dorsal; V ventral; n.g. character not given for Podocopa.
10. Adductor muscle scar pattern of five scars in vertical row date; ventral margin sinuate; exterior smooth; deep vestibules;
(plate 190K, 190L); 1st antenna (plate 190A) with eight hinge lophodont (plate 194K) . . . . . . . . . . Cytheromatidae
podomeres; basal podomere of 5th limb (plate 190H) has — In lateral view, anterior carapace margin obliquely rounded,
small branchial plate with setae (plate 190) . . . . . . . . . . posterior margin angulate to caudate; ventral margin
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Bythocytheridae nearly straight; exterior ribbed and pitted, with subtle ven-
— Adductor muscle scar of four scars in vertical row (plates trolateral alar thickenings, vestibules shallow or absent;
191A, 191D, 191E, 192J, 193J, 193K, 194I–194K); 1st an- hinge merodont (plate 192J) . . . . . . . . . . . . . Cytheruridae
tenna with no more than six podomeres (plate 192A); basal 14. Carapace subrhomboidal to subhexagonal in lateral out-
podomere of 5th limb without branchial plate (plate 193C) line; with lightly pitted ornament (plates 191B, 191C,
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11 196B) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 15
11. In lateral view, anterior end of carapace more narrowly — Carapace subovate or subquadrate in lateral outline; with
rounded than posterior end (plates 191A, 191D, 191E, lightly to heavily pitted to reticulate ornament (plates
193J, 193K, 194I, 194J) . . . . . . . . . . . . . . . . . . . . . . . . . . 12 191F, 195C–195G, 195I, 195J, 196A, 196C–196J) . . . . . 16
— In lateral view, anterior end of carapace usually more 15. Carapace subrhomboidal in lateral outline, with caudal
broadly rounded than posterior end (plates 192J, 194K) process located above midheight; hinge gongylodont;
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13 normal pore canals simple (plate 196B) . . . . . . . . . . . . . . .
12. Carapace fragile, elongate, laterally compressed (plates . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Loxoconchidae
191A, 191D, 191E, 193J, 193K); inconspicuous sexual di- — Carapace subhexagonal in lateral outline, with more or
morphism; hinge lophodont; rather oval frontal scar (plate less distinct posterior angle at about midheight; lightly pit-
191A); normal pore canals inconspicuous, funnel-type; no ted ornament with faint radial ridges; hinge antimerodont;
eye scar; mouth region (plate 193I) extended as cone with sieve-type and simple normal pore canals (plate 191B,
sucking disk; mandibles and 4th limbs greatly reduced and 191C) . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Cytheridae
transformed to hypodermic-like apparatus (plates 191A, 16. Hinge amphidont . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 17
191D, 191E, 193) . . . . . . . . . . . . . . . . Paradoxostomatidae — Hinge merodont or pentodont . . . . . . . . . . . . . . . . . . . . 18
— Carapace strongly calcified, inflated, egg-shaped and ventrally 17. Carapace elongate-oblong in lateral view, with concave pos-
flattened; conspicuous sexual dimorphism; hinge merodont; terodorsal element and slightly angulate posteroventral re-
frontal scar V-, J-, or W-shaped; normal pore canals conspic- gion; hinge hemiamphidont to holamphidont; adductor
uous, sieve-type; eye scar present behind eye spot; mouth muscle scar pattern with upper one to two scars divided, plus
region and mouth parts normal (plate 194I, 194J) . . . . . . . . two to three frontal scars; anterior radial pore canals very
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Xestoleberididae numerous; internal chitinous supports (sclerites) present in
13. In lateral view, anterior carapace margin broadly rounded, knee joints of walking legs (plates 195C–195G, 196E–196J)
posterior margin slightly more narrowly rounded, not cau- . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Hemicytheridae
426 ARTHROPODA
TA B L E 3
Tabular Key to Suborders and Superfamiliies of Order Halocyprida
Suborder Polycopina Suborder Halocypridina

Family
Character Family Polycopidae Thaumatocyprididae Family Halocyprididae
Carapace (valve)
Adult size (length) 0.1 mm–1.1 mm 0.4 mm–3.2 mm 0.4 mm–2.5 mm 0.75 mm–3.3 mm
Shape (outline) Ovoid or almost Elongate-ovate to Almost circular Usually somewhat sub-
circular ovate, few rather quadrate to oblong,
circular less often rather ovate
(Deeveya)
Rostrum Rare (only Ponto- Present in many Present in none Usually conspicuous
polycope)
Incisur Present in none Present in many Absent, but an- Usually conspicuous
(except overhang in teroventral mar- (often above mid-
Pontopolycope) gin straight or height) (slight in
slightly concave Deeveya)
and delimited by
anterior and an-
teroventral
processes
Dorsal margin Convex Convex or straight Convex or straight Usually rather straight,
sometimes convex
Posterodorsal margin Without keel Without keel With or without long Often with spine or
spinelike process corner-shaped
glandular process
Body
Bellonci organ Usually bifurcate and Bifurcate or not, not Not bifurcate; some- Bifurcate or not, often
Formed as two setae formed as setae times absent sexually dimorphic;
sometimes absent
Posterodorsal body Few with some Few with some Some with traces of Smooth
segmentation segmentation segmentation
Furca With short triangu- No triangular Anterior and ven- All claws articu-
lar process between processes (few with tral edges forming lated at base
at least longest a minute blunt peg) right angle, claws (Deeveyinae with
claws between claws on anterior mar- minute glandular peg
gin longer and ar- between two claws)
ticulated at base,
claws on ventral
margin unarticu-
lated
No. of Limbs (in- 6 8 8 8
cluding male copula-
tory limb)
1st Antenna
No. of articles 3–5 (inferred 8) 3–8 7–8 3–8

Bend in limb None Vanes Between first and Distal (Deeveya also
second article bent between first and
second)
Distal filament pad Absent Present in some Absent Euconchoecinae only
(continued)
Suborder Polycopina Suborder Halocypridina

Family
Character Family Polycopidae Thaumatocyprididae Family Halocyprididae
Mandible
Coxal endite Cusped tooth Cusped tooth

Basis endite No tooth Cusped tooth
Exopod Present (one article) Absent
Endopod—number of Two articles Jointed (number of
articles articles unclear)
Fourth Limb
Exopod Present (one to two Absent

articles)
Endopod Two to three articles Two articles
Fifth Limb
Endopod—number of One article Jointed (number of

articles articles unclear)
Exopod One article None
Sixth Limb: presence
Absent Present
Endopod NA Jointed
Exopod NA Present in few (one Present (one article— Present only in
article), usually absent except Danielopolina) Deeveyinae (one article)
Seventh Limb: presence
Absent Reduced (or absent), Unjointed One to two articles

with one to three setae
Male Copulatory Limb
Branched or not Two parts One to two parts Two parts One to two parts
NOTE : Key differences are indicated in boldface.
— Carapace (plates 191F, 196C, 196D) elongate-ovate to sub- List of Species

triangular in lateral view, without concave or angulate el-
ements; hinge schizodont (plate 191F); adductor muscle MYODOCOPA
scar pattern with no divided scars (plate 191F); few ante-
ANNE C. COHEN
rior radial pore canals; knee joints of walking legs without
internal supports . . . . . . . . . . . . . . . . . . . Schizocytheridae Myodocopa are mostly subtidal and few have yet been collected
18. Carapace elongate-subquadrate in lateral view; coarsely pit- intertidally within the geographic range of this book; most listed
ted to reticulate; simple frontal scar; few radial pore canals; below have been reported slightly subtidal near shore. AC
deep vestibules; hinge pentodont; egg care not known; Anne C. Cohen; RFM Rosalie F. Maddocks; DP Dawn Pe-
male 6th and 7th limbs symmetrical (plate 196A) . . . . . . terson; identified specimens of many of the following species are
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . Pectocytheridae in the California Academy of Sciences (San Francisco).
— Carapace elongate-oval in lateral view; surface smooth to
weakly pitted; frontal scar V- to J-shaped; numerous radial MYODOCOPIDA
pore canals; vestibules shallow or absent; females brood
young in domicilium; male right and left 6th and 7th limbs To identify juvenile instars see Kornicker and Harrison-Nelson
asymmetrical (plate 195I, 195J) . . . . . . . . . . Cytherideidae 1999, Smithsonian Cont. Zool. 602: 1–55 (see pp. 32–36) and
428 ARTHROPODA
TA B L E 4
Tabular Key to Families of the Order Myodocopida
Subfamily Subfamily Subfamily

Cypridinidoidea Cylindroleberoidea Sarsielloidea
Character Cypridinidae Cylindroleberididae Philomedidae Rutidermatidae Sarsiellidae
Carapace (valves]
Outline round None (plates 184A, None (plates 185E–G; None (plate 186A, None (plate 187B, Female and
(except for keel) 184B, 185A) 187A) 186B, 186D, 186E) 187C) juvenile Sarsielli-
nae round or
round except for
keel (plate 187G,
187H)
Incisur (anterior Deep (1 exception) Shallow (plate 187A) Minute to deep (plate Shallow (plate 187B, Minute (plate 186H) to
slot or (plate 184A, 184B) to deep (plate 185E) 186A, 186B, 186D, 187C) shallow (plate 186G)
indentation) or slitlike (Cylin 186E) (Pleoschisma
droleberidinae) none)
(plate 185F, 185G)
Smooth at low Usually Cylindroleberidinae, Some None completely None completely
magnification? Cyclasteropinae smooth smooth
Ornamented with Few Asteropteroninae Many Most All
nodes, ridges, etc. only
Posterior vertical row None except male All males (except None None None
of external hairs Codonocera Asteropteroninae)
Male more elongate? Some Some Yes Yes Yes
Body
Bellonci organ Short or long Usually long Long Long Long

Upper lip with Present in all Absent Present (few Absent Absent
valvular nozzles? (many nozzles) nozzles) or
(plate 185B) absent
Posterodorsal body Most smooth (plate Usually 7–8 Smooth Smooth Smooth
184A, 184B); (some fewer) dor-
some males with sal over-lapping
row of large lobes flat leaf-like gills
Furca
lst claw with Yes (plates 184A, Yes (plate 185H) Yes (plate 186F) Yes No (plate 188E)
basal suture 184B, 185D)
Other claws without Sometimes No No No Sometimes
basal suture
1st Antenna
3rd and 4th articles Separated by suture Usually separated Separated by suture Sometimes fused Fused
by complete
suture
Male 5th article Not reduced Usually little Wedged ventrally Wedged ventrally Wedged ventrally
reduction between 4th and between 4th and between 4th and 6th
6th 6th
Male seta of 5th No or slight (some Cyclasteropinae, Yes Yes Yes
article with bush with bushes on most Cylindrole-
of long filaments? setae of terminal berididae and
article) Asteropteroninae
(continued)

Males with Yes No No No No

terminal filaments
with suckers?
7th article with None All Cylindroleberidi- None None None
clawlike a-seta nae, most Cyclas-
teropinae, some
Asteropteroninae
2nd Antenna
Protopod with seta? With minute distal Most with minute No seta No seta No seta
seta distal seta
Male endopod a Not in most, but Always Always Always In most
clasper? in some
Mandible
Coxal endite Spiny lobe, rarely Long, scythe- Spiny lobe, some Spiny lobe (plate Absent or small spiny
shape absent shaped branched, long, reduced in 188A), reduced in lobe, reduced in
spiny male male (plate 188B) male
Basis endite Absent Lobe with setae Absent Absent Absent
Exopod Short, 2 setae Short, 2 setae Short, 2 setae Absent or reduced, Absent or reduced,
0–?2 setae 0–1 setae
Endopodial claws Terminal group Terminal group Terminal group Pincer in females 3-pronged rake in
form (plate 185C) (plate 186C) and juveniles female and juve-
(except nile Sarsiellinae
Metaschismatinae) (plate 188C)
Endopodial All terminal: (2-) 3 All terminal: (1-) 3 All terminal: Not all terminal in Not all terminal in
claws—number relatively short & relatively long & females & juve females & juve- females & juve-
and position all on 3rd article all on 3rd article niles with (2-) 3 niles: claws form niles: usually
(plate 185C) relatively long pincers (plate 188A) with 1 (Dantyi-
claws (plate 186C); (except nae 0–3) claw on
males with 1–3 Metaschisma), each of 3 endopo-
claws and all with 1 (-2) claw dial articles
on 3rd article on 2nd article and (plate 188C); males
1 claw on 3rd; with 1–3 terminal
males with 1 claws only (or plus
terminal claw weaker proximal
(plate 188B) claws) (plate 188D)
4th Limb (plate 184A, 184B)
Reduced and weak No No Yes Yes Yes

in males?
With setal comb No Yes, on protopod No No No
With basal epipod? No Bare lobe ?= epipod No No No
Endopod—claws At least 2 claws, Reduced; no claws Females and Females and juve- Females and juve-
on 2nd article several setae, juveniles with 2 niles with at least 2 niles with termi-
(terminal) many pectinate claws and various claws, several/few nal row of 5 stout
setae setae; Metaschisma claws (3–5 triangu-
with 3 lar) and 1 subtermi-
nal long thin seta on
each side of claw row
(continued)

Reduced, weak in No No Yes Yes Yes

males?
With setal comb No Yes No No No
With teeth or claws Row of claws No teeth or claws 1 big squarish or 1 big squarish Sarsiellinae none;
(usually 6) elongate tooth on tooth on 2nd Dantyinae with
short peg on 2nd article, smaller article, smaller big squarish
1st exopodial teeth on 1st article teeth on 1st tooth on 2nd
article article article, small
tooth on 1st
article
With well deve- 4 All reduced 4 4 1–4

loped endites?
Reduced in males? No No In a few No Usually

Terminus Peg or tooth Opposing combs Peg(s) opposite Peg(s) or comb Small opposing
opposite comb comb opposite comb combs
Male 8th Limb (plate 184A)
Copulatory limb Big, complex, 3 Small in most Short to elongate, 3 Small elongate with Small elongate with
lobes (1 jointed) males, 3 lobes, lobes, some/all 3 lobes 3 lobes, with distal
with internal unjointed, no with sclerites sclerotized hook
sclerites sclerites
NOTE : Key differences are indicated in boldface.
Kornicker 1992, Smithsoman Cont. Zool. 531: 1–243. For in- Vargula tsujii Kornicker and Baker 1977, Proc. Biol. Soc.
formation on seven Northwestern American coastal species Washington 90: 218–231 (Vargula americana [Müller, 1890] of
beyond range of this book, see Juday 1907, Univ. Calif. Publ. Hobson and Chess 1976, Fisheries Bull. 74: 567–598). Benthic
Zool. 3: 135–156; Lie, 1968, Fiskedirektoratets Skrifter, serie in sand, planktonic, 3 m–931 m, Baja California to south of
HavUnders¢kelser, Bergen 14: 229–556 (identifications un- Monterey Bay, not reported further north. Planktonic occur-
verified, specimens not extant); Lie and Evans 1973, Marine rence probably only as nocturnal demersal plankton: see Ham-
Biol. 21: 122–126 (identifications unverified, specimens not mer 1981, Mar. Biol. 62: 275–280; Hobson and Chess 1976
extant); Poulsen 1965, Dana Report 65: 1–484; Kornicker (also fish predation); Chess and Hobson 1997, NOAA Tech.
and Myers 1981, Smithsonian Cont. Zool. 334: 1–35; Chess Memorandum NMFSSWFSC-243, U.S. Dept. Com. (also fish
and Hobson 1997, NOAA Tech. Memorandum NMFS-SWFSC- predation); Stepien and Brusca 1985, Mar. Ecol. Prog. Ser. 25:
243, U.S. Dept. Com.; and website of SCAMIT (South. Calif. 91–105 (also fish predation). Bioluminescence in midshipman
Assoc. Mar. Invert. Taxonomists): http://www.scamit.org/ fish is related to predation on V. tsujii: see Thompson, Naf-
taxonomic_tools. htm or http://www. scamit.org/index.htm) paktitis, and Tsuji 1988, Mar. Biol. 98: 7–13; 1988, Comp.
(plates 184A, 184B, 185–188). Biochem. Physiol. A, 89: 203–210; Thompson and Tsuji 1989,
Mar. Biol. 102: 161–165; Thompson et al. 1998, J. Exp. Biol.
CYPRIDINIDAE 137: 39–52. For more on bioluminescence see Huvard 1990,
Acta Zoologica 71: 217–223; 1993, Comp. Biochem. Physiol.
For summary and phylogenetic analysis see Cohen, A. C. and A, 104: 333–338, J. Morph. 218: 181–193 (plate 185A–185D).
J. Morin 2003. Sexual morphology, reproduction and biolumi-
nescence in Ostracoda. In Bridging the Gap: Trends in the Os- CYLINDROLEBERIDIDAE
tracode Biological and Geological Sciences. L. E. Park and A. J.
Smith, eds. The Paleontological Society Papers 9. New Haven: For keys see Kornicker 1981, Smithsonian Cont. Zool. 319: 1–548
Yale University Press. (subfamilies of Cylindroleberididae, genera of Cylasteropinae,
OSTRACODA 431
Asteropteroninae; also general morphology of Myodocopida); on S. Calif. shelf); Oliver et al. 1980, Fish. Bull. 78: 437–454
Kornicker 1986, Smithsonian Cont. Zool. 425: 1–139 (some gen- (physical ecology, Monterey Bay) (plate 186A–186C).
era of Cylindroleberidinae). To identify juvenile instars see Ko- Euphilomedes longiseta (Juday, 1907) (Philomedes longiseta
rnicker 1992, Smithsonian Cont. Zool. 531: 1–243. Juday 1907, Univ. Calif. Publ. Zool. 3: 135–156; P. longiseta,
of Lucas, 1952, Contr. Can. Biol. Fish., Stud. Biol. Stat. Canada,
n.s. 6: 398–416). Surface plankton tows off San Diego (Juday,
ASTEROPTERONINAE 1907); benthic sand, 6 m –9 m off Monterey Bay (Oliver et al.
1980, Fish. Bull. 78: 437–454); 20 m–90 m, near Vancouver Is-
Asteropella slatteryi Kornicker, 1981. Benthic in sand mixed land (Lucas, 1952).
with silt and clay in harbor (often in kelp bed) at Half Moon Euphilomedes morini Kornicker and Harrison-Nelson, 1997
Bay; Moss Landing, Monterey Bay and Pilar Point Harbor, Half (Philomedes longiseta of Tuel et al. 1976, Biol. Survey Pillar
Moon Bay; 1.8 m–37 m. See Kornicker and Harrison-Nelson Point Harbor). Benthic in 1.8 m on sand, Pillar Point Harbor,
1997, Smithsonian Cont. Zool. 593: 1–53 (description, dis- Half Moon Bay. See Kornicker and Harrison-Nelson 1997,
tribution, ecology); Chess and Hobson 1997, NOAA Tech. Smithsonian Cont. Zool. 593: 1–53 (description, distribution,
Memorandum NMFS-SWFSC-243 (Catalina Is., benthic in ecology) (plate 186D–186F).
sediment by day, nocturnal plankton, fish predation) (plate Zeugophilomedes oblonga (Juday, 1907) (Euphilomedes ob-
187A). longa, of Oliver et al. 1980, Fishery Bull. 78: 437–454). Surface
plankton off San Diego Bay and San Pedro (Juday 1907, Univ.
Calif. Publ. Zool. 3: 135–156); benthic, 9 m–14 m, subtidal high
CYCLASTEROPINAE
energy beach, Monterey Bay (Oliver et al. 1980); see Kornicker
and Harrison-Nelson 1997, Smithsonian Cont. Zool. 593: 1–53.
Leuroleberis sharpei Kornicker, 1981 (Cylindroleberis lo-
bianci Müller, 1894 of Sharpe 1908, Proc. U.S. Nat. Mus. 35:
399-430 [not C. lobianci Müller]; Cycloleberis lobiancoi of
RUTIDERMATIDAE
Hobson and Chess, 1976). Depth 2 m–146 m, planktonic
(probably only as demersal plankton at night) and benthic
For bibliography giving synonymy, distribution, biology for all
from Gulf of California to Monterey Bay, possibly to Alaska;
Rutidermatidae as of 1986, see Cohen and Kornicker 1986,
in Monterey Bay off Sunset Beach, Kaiser Trestle, Watsonville
Smithsonian Cont. Zool. 449: 1–11. For keys see Kornicker and
outfall, and Pajaro River, and also off South Jetty and north-
Myers 1981, Smithsonian Cont. Zool. 334: 1–35 (S. Calif.);
west of harbor entrance at Moss Landing. For fish predation,
Kornicker 1983, Smithsonian Cont. Zool. 371: 1–86 (part of Ru-
benthic and planktonic habitats at Catalina Id., see Hobson
tidermatidae); Kornicker 1992, Smithsonian Cont. Zool. 531:
and Chess 1976, Fish. Bull. 74: 567–598; Chess and Hobson
1–243 (juvenile instars of Rutiderma). For subfamily Metaschis-
1997, NOAA Tech. Memorandum NMFS-SWFSC-243 (plate
matinae see Kornicker 1994, Smithsonian Cont. Zool. 553:
185E).
1–200.
Rutiderma apex Kornicker and Nelson, 1997 (Rutiderma sp.
CYLINDROLEBERIDINAE of Tuel et al. 1976, Biol. Survey Pillar Point Harbor, El Granada
CA, Rep. Marine Ecol. Inst.). Benthic on sand, sandy silt, and
Postasterope barnesi (Baker, 1978) (Parasterope barnesi Baker silty clay, 1.8 m–11 m, Pillar Point Harbor, Half Moon Bay,
1978, Crustaceana 35: 139–141; Parasterope sp. of Tuel et al. Tomales Bay, and Dark Gulch, Mendocino County; also benthic
1976, Biol. Survey Pillar Point Harbor, El Granada, Calif., Rep. on silty sea grass near Spud Pt. Marina, Bodega Harbor (males
Mar. Ecol. Inst., Redwood City, CA). Benthic in sand, scooped from bottom, depth 1 m, and at night in subtidal
1.8 m–210 m; San Diego to Point Conception; Pillar Point Har- benthic light trap) CAS, AC. See Kornicker and Harrison-Nelson
bor, Half Moon Bay; also males collected at night in subtidal 1997, Smithsonian Cont. Zool. 593: 1–53 (description, distri-
benthic light trap on silty sea grass near Spud Pt. Marina bution, ecology) (plates 187B, 187C, 188A, 188B).
Bodega Harbor, CAS, AC. See Kornicker and Harrison-Nelson,
1997, Smithsonian Cont. Zool. 593: 1–53 (description, distri- SARSIELLIDAE
bution, ecology) (plate 185F–185H)

For keys to subfamilies of Sarsiellidae, genera of Sarsiellinae, see
Kornicker 1986, Smithsonian Cont. Zool. 415: 1–217; Kor-
PHILOMEDIDAE nicker 1991, Smithsonian Cont. Zool. 505: 1–140. Kornicker,
L. S. and F. E. Caraion. For key to genera of subfamily Dantyi-
For keys, see Kornicker 1981, Smithsonian Cont. Zool. 332: nae see Kornicker and Caraion 1980, Smithsonian Cont. Zool.
1–16 (most genera of Philomedinae); Kornicker 1989, Smith- 309: 1–27.
sonian Cont. Zool. 467: 1–134 (genera of Pseudophilomedinae).
Euphilomedes carcharodonta (Smith 1952) (Philomedes car- SARSIELLINAE
charodonta Smith 1952, J. Fish. Res. Board Canada 9: 16–41;
“myodocopid” in Tomales Bay, of Kornicker 1977, in Aspects Eusarsiella zostericola (Cushman, 1906) (Sarsiella zostericola
Ecol. Zoogeog. Recent Fossil Ostracoda: 159–173 [pers. comm. Cushman, 1906; Sarsiella tricostata Jones, 1958). Benthic at
Kornicker]). Benthic in 5 m–180 m from La Jolla to Ganges Har- 1 m–3 m, off Pt. Richmond, San Francisco Bay; shallow
bor, British Columbia, including Monterey Bay, Half Moon Bay, shores of Atlantic bays from Maine to Chesapeake Bay; also
Tomales Bay, and Bodega Bay. See Kornicker and Harrison- coastal lagoons of Texas, and near Essex, England. The species
Nelson 1997, Smithsonian Cont. Zool. 593: 1–53 (description, was introduced to San Francisco Bay, the Gulf of Mexico, and
distribution, ecology); Poulsen 1962, Dana Report 57 (de- England with oysters (Crassostrea virginica) transplanted from
scription); Baker 1974, Texas J, Science 25: 131–132 (abundant the Atlantic coast. See Kornicker and Wise 1962, Crustaceana
432 ARTHROPODA
P LATE 185 Myodocopida, Cypridinidae and Cylindroleberididae; Cypridinidae: Vargula tsujii, after Kornicker and Baker, 1977: A, fe-
male showing deep anterior incisur, interior brooded embryos; B, anterior of body showing Bellonci organ attached to naupliar eye,
upper lip (with nozzles and tusk); C, male mandible (medial view; coxal endite not shown, but showing relative shortness of termi-
nal claws); D, left male furca showing 2nd claw without basal suture, other claws with sutures; Cylindroleberididae: E. Leuroleberis
sharpei after Kornicker, 1981: female, showing circular shape and deep anterior incisur; Postasterope barnesi after Kornicker and Harri-
son-Nelson 1997: F, male, showing deep slitlike anterior incisur, long male setae of 1st antennae, male posterior hair row on valve;
G, female, showing slitlike incisur, interior embryos; H, left furca showing all claws with basal suture (all but E, redrawn by Ginny
Allen; E by A. Cohen).
P LATE 18 6 Myodocopida, Philomedidae, Sarsiellidae; Philomedidae: Euphilomedes carcharodonta, after Poulsen 1962: A, male
(more elongate valve, larger more visible lateral eye than female); B, female; C, female mandible tip, showing long terminal
claws; Euphilomedes morini, after Kornicker and Harrison-Nelson, 1997: D, male; E, female; F, male left furca; Sarsiellidae: Eusar-
siella zostericola, after Kornicker 1967, anterior of valves to left: G, male (with overhanging anterior rostrum); H, female (with
mostly circular shape, no rostrum) (all redrawn by Ginny Allen).
P LATE 187 Myodocopida, Cylindroleberididae and Rutidermatidae, showing ornate valves. Cylindroleberididae: A, Asteropella slatteryi fe-
male, from Kornicker, 1981, showing shallow incisur; Rutidermatidae: Rutiderma apex, from Kornicker and Harrison-Nelson, 1997: B, female;
C, male (more elongate and less ornate than female) (drawn by Carolyn Gast, loaned by Smithsonian Institution).
4: 59–74 (description); Kornicker 1975, Bull. Amer. Paleont. 65: Bodega Bay, RFM. This parthenogenetic species is abundant
129–139 (spread with oysters); Kornicker and Harrison- worldwide in permanent and temporary fresh waters. See Sars
Nelson 1997, Smithsonian Cont. Zool. 593: 1–53 (distribution, 1925, An Account Crustacea Norway (Ostracoda) 9: 1–277;
and references therein); Bamber 1987, J. Micropaleo. 6: 57–62 Tressler, 1959, in Fresh-water Biology, Ward, Whipple and Ed-
(life history in Great Britain) (plates 186G, 186H, 188C–188E). monson, eds.: 657–734); Delorme 1970, Can. J. Zool. 48: 253–266)
(plate 189C–189E).
PODOCOPA Herpetocypris reptans (Baird, 1835). Females in high inter-
ROSALIE F. MADDOCKS AND DAWN PETERSON tidal/supratidal tide pools with freshwater influx, Bodega
Bay, RFM. Parthenogenetic, common in fresh waters of Eu-
PODOCOPIDA rope and North America. See above references (plate
189F–189H).
BAIRDIOCOPINA
Heterocypris salina (Brady, 1868) (Cyprinotus salinus).
Empty valves in roots of eelgrass in brackish water inlet, depth
BAIRDIIDAE
10 cm, Richardson Bay, Marin County, CA, DP. Females and
juveniles in high intertidal/supratidal pools with freshwater
Neonesidea sp. In sand dredged from shore, Whaler’s Cove,
influx, Bodega Bay, RFM. This parthenogenetic, halophile,
Point Lobos (plate 195H).
somewhat variable species is very widely reported in brackish
CYPRIDOCOPINA waters of Europe, Russia, the Caspian Sea and North America.
See above references and Delorme 1970, Can. J. Zool. 48:
CYPRIDIDAE 153–168; Swain (1999, Fossil Nonmarine Ostracoda U.S.). See
also Ganning 1967, Helgoländer wiss. Meeresunters. 15: 27–40
Cypridopsis vidua (O. F. Müller, 1786). Females and juveniles (ecology of rockpool populations of Scandinavia) (plate 189A,
in high intertidal/supratidal tide pools with freshwater influx, 189B, 189I–189L).
OSTRACODA 435
P LATE 18 8 Myodocopida, Rutidermatidae and Sarsiellidae; Rutidermatidae: Rutiderma apex, after Kornicker and Harrison-Nelson,
1997: A, female mandible (showing terminal pincer-claws), medial view; B, male mandible (no pincer), medial view; Sarsiellidae:
Eusarsiella zostericola: C, female mandible (showing one long ventral claw on each endopodial podomere forming rake), medial
view; D, male mandible (long claw only on terminal podomere), medial view; E, male right furca showing 1st (longest) claw with-
out basal suture, other claws with sutures (C, D, after Kornicker 1967; E, after Kornicker and Wise 1962; redrawn by Ginny Allen).
Sarscypridopsis aculeata (Costa, 1847) (Cypridopsis aculeata). CYTHEROCOPINA

Empty valves in roots of eelgrass in brackish water inlet, depth
10 cm, Richardson Bay, Marin County, DP. It has been widely BYTHOCYTHERIDAE
reported from Europe, Iceland, the Americas, central Asia and
Africa. See above references and Delorme 1970, Can. J. Zool. Sclerochilus sp. On intertidal mud and fine sand flat near
48: 253–266; Forester and Brouwers 1985, J. Paleont. 59: Sandpiper Restaurant, northeast Bodega Harbor, RFM (plate
344–369 (plate 195A, 195B). 190A–190L).
436 ARTHROPODA
P LATE 18 9 Podocopida: Freshwater Cyprididae: Heterocypris salina (Brady, 1868), female specimen RFM 3413F
(A, B, I–L): A, B, exteriors of right and left valves with traces of ovaries, X60; I, exterior of left valve, normal pore canals and
granular micro-ornament indicated as dots and ovals, X105; J, muscle scar pattern of right valve, X105; K, anterior margin of
right valve with radial pore canals, X105; L, anteroventral margin of right valve with minute tuberculae, X175; Cypridopsis
vidua (O. F. Müller, 1786), female specimen RFM 3411F (C–E): C, muscle scar pattern of right valve, X105; D, E, exteriors of
right and left valve, X60; Herpetocypris reptans (Baird, 1835), female specimen RFM 3412F (F–H): F, G, exteriors of right and
left valves, X35; H, anterior margin of left valve with flange, radial pore canals, outer list, striate inner list, and inner margin,
X175 (drawn by R. Maddocks).
CYTHERIDAE
20: 257–352). On intertidal mud and fine sand flat in Bodega
Cythere alveolivalva Smith, 1952 (Cythere uncifalcata Smith, Harbor, RFM. Valentine (1976) reported range in offshore wa-
1952; Cythere sp. A of Valentine, 1976). This species is common ters as Point Reyes to southern Vancouver Island; also fossil
in coastal regions of the northern Kurile Islands, Kamchatka (Pleistocene of Cape Blanco, Oregon).
Peninsula, and Alaska to California; also fossil (Pleistocene and Cythere sp. of Tsukagoshi and Ikeya, 1987 (Cythere alveolivalva
Holocene) in Oregon and California (Tsukagoshi and Ikeya (part) of Swain 1969, in Neale (ed.), Taxon., Morph. Ecol. Rec.
1987) (plate 191B, 191C). Ostracoda, and Swain and Gilby 1974, Micropaleo. 20: 257–352;
Cythere valentinei Tsukagoshi and Ikeya, 1987 (Cythere sp. Cythere sp. aff. C. lutea of Swain and Gilby 1974; Cythere maia of
B of Valentine, 1976; Cythere cf. C. lutea and Cythere alveolivalva Valentine, 1976). Tide pool at Point Piedras Blancas, CA. Valen-
(part) of Swain 1969, in Neale (ed.), Taxon. Morph. Ecol. Rec. tine (1976) reported range in offshore waters as Baja California
Ostracoda; Cythere lutea of Swain and Gilby 1974, Micropaleo. to Point Piedras Blancas, CA; also fossil (Pliocene-Pleistocene).
OSTRACODA 437
P LATE 19 0 Podocopida: Bythocytheridae: Sclerochilus sp.: female specimen RFM 3408F (A–E, G–L): A, first antenna; B, second antenna
with spinnaret gland; C, mouth region with forehead, upper and lower lip; D, E, palp and masticatory process of maxillula (fourth limb);
G, mandible; H–J, fifth, sixth and seventh limbs; all X430; K, L, exteriors of right and left valves, X120; F, male specimen RFM 3403M,
brush-shaped organ, X430 (drawn by R. Maddocks).
CYTHERIDEIDAE marshes along the Pacific coast from southern Chile to British
Columbia, also Midway and Oahu Islands in the North Pacific;
Cyprideis beaconensis (LeRoy, 1943). Shallow water among algae probably dispersed on migratory waterfowl (Sandberg 1964,
bloom, salt marsh, Skaggs Island Naval Station, Sonoma Co., CA, Stockholm Contribs. Geology 7: 1–178; Sandberg and Plusquel-
DP; Lake Merritt, Oakland; widely distributed in estuaries and lec 1974, Geoscience and Man 6: 1–26) (plate 195I, 195J).
438 ARTHROPODA
P LATE 191 Podocopida: Cytheridae, Paradoxostomatidae, Schizocytheridae. Paradoxostomatidae: A, Acetabulostoma californica, exterior of
male left valve, X80, after Watling, 1972. Cytheridae: B, C,, Cythere alveolivalva Smith, 1952, exteriors of male right and left valves, X35, af-
ter Smith, 1952; Paradoxostomatidae: Paradoxostoma striungulum Smith, 1952 (D, E): D, entire male specimen RFM 3400M, as seen from left
side in transmitted light; E, entire female specimen RFM 3401F, as seen from left side in transmitted light; with postmortem extrusion of
body from carapace; all X175; Schizocytheridae: F, Spinileberis hyalina, interior of right valve, X65, redrawn from Watling 1970 (drawn by
R. Maddocks).
CYTHEROMATIDAE HEMICYTHERIDAE
Paracytheroma similis Skogsberg, 1950 (Calif. Acad. Sci., Proc. Ambostracon glaucum (Skogsberg, 1928) (Cythereis glaucum
26: 483–505). On sand and rocks, 15 m in Monterey Bay, near Skogsberg 1928, Calif. Acad. Sci., Occ. Pap. 15: 1–143). On
Pacific Grove (plate 194K). holdfasts of algae in Carmel Bay; also on calcareous algae in
tide pool just outside Hopkins Marine Station, Pacific Grove;
CYTHERURIDAE Swain and Gilby (1974) reported it from 12 m in Bodega Bay;
from 23 m in Morro Bay, and from 11 m–26 m in Coos Bay
Howeina sp. aff. H. camptocytheroidea Hanai, 1957. On inter- (Micropaleo. 20: 257–352). Valentine (1976) gave the range
tidal mud and fine sand flat in Bodega Harbor, RFM (plate in offshore waters as Baja California to Pt. Piedras Blancas
192A–192J). (Micropaleo. 20: 257–352) (plate 195C, 195D).
OSTRACODA 439
P LATE 192 Podocopida: Cytheruridae: Howeina sp. aff. H. camptocytheroidea Hanai, 1957, male specimen RFM 3403M: A, first antenna; B, sec-
ond antenna; C, spinnaret gland; D, maxillula (fourth limb); E, mandible; F–H, fifth, sixth, and seventh limbs; I, hemipenis, all X430; J, exte-
rior of left valve, X175 (drawn by R. Maddocks).
Aurila laeviculoidea Swain and Gilby, 1974. In sand in 10 m ters of Japan, Korea, and the South Kurile Islands (Schornikov
off Crescent City. and Tsareva 1995, Mitt. Hamburg. Zoolog. Mus. Inst. 92:
Aurila lincolnensis (LeRoy, 1943). Among shore rocks at Rock- 237–253) (plate 196I, 196J).
away Beach, San Mateo Co., DP; also on intertidal mud and Aurila sp. of Swain and Gilby, 1974 (Micropaleo. 20: 257–
fine sand flats in Bodega Harbor; Swain and Gilby (1974) re- 352). In sand in 10 m in Bodega Bay. It is likely that addi-
ported it from sand in 12 m in Bodega Bay, RFM, and in 10 m tional species of Aurila and related genera occur in the in-
off Crescent City (Micropaleo. 20: 257–352). The species is tertidal zone within the range of this manual (see Valentine
widely reported off southern California and Mexico. Valentine 1976).
(1976) gave the range in offshore waters as Baja California to Radimella aurita (Skogsberg, 1928) (Cythereis aurita
Cape Flattery (plate 196E, 196F). Skogsberg 1928, Calif. Acad. Sci., Occ. Pap. 15: 1–143). On
Aurila montereyensis (Skogsberg, 1928) (Cythereis mon- calcareous algae in tide pool and on holdfasts of Macrocys-
tereyensis, Skogsberg, 1928). On calcareous algae, among roots tis, outside Hopkins Marine Station on Monterey Bay, near
of eelgrass, and on holdfasts of Macrocystis in 2 m, in tide pools Pacific Grove; also on kelp near shore and on eelgrass in
at Carmel Bay; Bodega Harbor, in fairly dense seagrass, silt with Carmel Bay. Valentine (1976) reported the range in offshore
fine sand, RFM. Valentine (1976) gave the latitudinal range in waters as Baja California to north of Santa Cruz); also fossil
offshore waters as north of Pt. Conception to near Cape Alava, (plate 195E, 195F).
Washington (plate 195G). Radimella? pacifica (Skogsberg, 1928) (Cythereis pacifica
Aurila sp. aff. A. corniculata (Okubo, 1980). McNears Beach, Skogsberg 1928, Calif. Acad. Sci., Occ. Pap. 15: 1–143). On cal-
San Francisco Bay, A. corniculata is reported from coastal wa- careous algae in tide pool and on holdfasts of Macrocystis in 2
440 ARTHROPODA
P LATE 193 Podocopida: Paradoxostomatidae: Paradoxostoma sp., male specimen RFM 3407M: A, first antenna; B, second an-
tenna; C–E, fifth, sixth, and seventh limbs; F, maxillula (fourth limb); G, mandible; H, hemipenis; I, mouth region with fore-
head, upper lip, oral sucking disk, and mandible within esophagus; all X430; J, K, exteriors of right and left valves, X175
(drawn by R. Maddocks).
m, outside Hopkins Marine Station near Pacific Grove; also on LOXOCONCHIDAE

kelp near shore and on eelgrass in Carmel Bay. Valentine (1976)
reported the range in offshore waters as Baja California to Cytheromorpha sp. Empty valves in brackish water on medium-
south of Cape Vizcaino; also fossil. grained sand and rocks, in less than 1 m, McNears Beach, Marin
Robustaurila jollaensis (LeRoy, 1943) (Aurila jollaensis, of Co.; in brackish water on muddy sand, 10 cm, Richardson Bay,
Swain and Gilby 1974, Micropaleo. 20: 257–352). Bodega Bay Marin Co.; in brackish water on mudflat among floating algae,
in 12 m and off Crescent City in 10 m. Valentine (1976) re- in 5 cm, Bolinas Bay, Bodega Bay, DP.
ported the range in offshore waters as Baja California to Cape Loxoconcha lenticulata LeRoy, 1943. Empty valves in brack-
Flattery; also fossil (plate 196G, 196H). ish water on medium-grained sand and rocks, 1 m, McNears
OSTRACODA 441
P LATE 19 4 Podocopida: Cytheromatidae and Xestoleberididae. Xestoleberididae: A–I, Xestoleberis sp., male specimen RFM 3402M
(A–H); A, first antenna; B, mandibular base; C, antenna; D, brush-shaped appendage; E–G, fifth, sixth, and seventh limbs; H,
hemipenis; all X430; I, left exterior of female specimen RFM 3410F, with developing eggs in domiciliary brood pouch, pigmenta-
tion and patch pattern indicated by dots, X175; J, Xestoleberis hopkinsi male left valve exterior, X120; Cytheromatidae: K, Para-
cytheroma similis male left valve exterior, X100 ( J, redrawn from Skogsberg, 1950, pl. 29, fig. 1; K, redrawn from Skogsberg, 1950;
drawn by R. Maddocks).
Beach, Marin Co., DP. Valentine (1976) reported the range in Paradoxostoma striungulum Smith, 1952. Empty valves in
offshore waters as northernmost Baja California to Pt. Con- brackish water on medium-grained sand and rocks, less than 1
ception; also fossil (plate 196B). m, McNears Beach, Marin Co., DP; living on intertidal fouled
eelgrass (Zostera marina) blades in the South Slough National
PARADOXOSTOMATIDAE Estuarine Research Reserve, Coos Bay (J. T. Carlton, collector),
CAS, RFM. Described from Departure Bay, British Colombia, on
Acetabulostoma californica Watling l972 (Proc. Biol. Soc. Obelia near water surface (Smith 1952, J. Fish. Res. Board
Washington 85: 481–488). Low intertidal on exposed side of Canada 9: 16–41) (plate 191D, 191E).
Tomales Point, Marin Co., in the zone of the red alga Corallina Paradoxostoma sp. One male living on mud and fine sand flat
gracilis. Most species of Acetabulostoma are parasitic on gam- exposed at low tide, Bodega Harbor, RFM. It is likely that ad-
marid amphipods, but the host (if any) of A. californica is un- ditional species of Paradoxostoma and related genera occur
known (plate 191A). within the range of this book (plate 193A–193K).
442 ARTHROPODA
P LATE 195 Podocopida: Bairdiidae, Cyprididae, Cytherideidae, Hemicytheridae. A, B, Cyprididae: Sarscypridopsis aculeata (Costa,
1847), CAS 121631, right and left exteriors of female carapaces, X67; Hemicytheridae: Ambostracon glaucum (Skogsberg, 1928), CAS
121629 (C, D): C, right exterior of female; D, left exterior of male, X67; E, F, Radimella aurita (Skogsberg, 1928), CAS 121627: E, ex-
terior of male (?) right valve; F, exterior of female (?) left valve; both X67; G, Aurila montereyensis (Skogsberg, 1928), CAS 121622,
exterior of female (?) left valve, X67; Bairdiidae: H, Neonesidea sp., CAS 120522, left exterior of carapace, X67; Cytherideidae: I, J:
Cyprideis beaconensis (LeRoy, 1943), CAS 121619: I, right exterior of male carapace; J, left exterior of female carapace with heavy
coating of microbial slime; both X67 (SEMs by D. Peterson).
P LATE 19 6 Podocopida: Hemicytheridae, Loxoconchidae, Pectocytheridae, Schizocytheridae. Pectocytheridae: A, Pectocythere park-

erae Swain and Gilby, 1974, CAS 121626, exterior of left valve, X100; Loxoconchidae: B, Loxoconcha lenticulata (LeRoy, 1943),
UMPC 12220, exterior of male left valve, X100; Schizocytheridae: C, Acuminocythere crescentensis Swain and Gilby, 1974, UMPC
12096, right exterior of female carapace, X100; D, Acuminocythere sp. of Swain and Gilby, 1974, UMPC 12080, exterior of left
valve, X100; Hemicytheridae: E, F, Aurila lincolnensis (LeRoy, 1943), CAS 121620, right and left exteriors of carapaces, X100; G, H,
Robustaurila jollaensis (LeRoy, 1943), CAS 1212630, right and left exteriors of carapaces, X100; I, J, Aurila aff. A. corniculata (Okubo,
1980), CAS 121628, exteriors of right and left valves, X100 (SEMs by D. Peterson).
PECTOCYTHERIDAE taceans: the protopod, epipod, exopod and endopod. Lethaia 31:
251–265.
Pectocythere parkerae Swain and Gilby, 1974 (Munseyella sp. Cohen, A. C., and J. G. Morin. 1990. Patterns of reproduction in ostra-
codes; a review. Journal of Crustacean Biology 10: 84–211.
B of Valentine, 1976). On sand in 11 m–36 m in Bodega Bay,
Ellis and Messina Catalogue of Ostracoda. Micropaleontology Press,
off Crescent City, and in Coos Bay. Valentine (1976) reported American Museum of Natural History. For online access go to:
the range in offshore waters as San Diego to Cape Lookout, OR http://www.micropress.org.
(plate 196A). Hartmann, G. 1966, 1967, 1968, 1975, 1989. Ostracoda. In Dr. H. G.
Pectocythere tomalensis Watling, 1970 (Crustaceana 19: 251– Bronns Klassen und Ordnungen des Tierreichs, Fünfter Band: Arthro-
poda, I. Abteilung, 2. Buch, IV. Teil, Lieferungen 1–5: 1–1067.
263). In 6 m on sandy bottom, White Gulch, Tomales Bay; also
Hartmann, G., and H. S. Puri 1974. Summary of neontological and pa-
on sand in 12 m in Bodega Bay (Swain and Gilby 1974, Mi- leontological classification of Ostracoda. Mitteilungen aus dem Ham-
cropaleo. 20: 257–352); Valentine (1976) reported the range in burgischen Zoologischen Museum und Institut 70: 7–73.
offshore waters as Monterey Bay to Cape Flattery. Horne, D., A. C. Cohen, and K. Martens. 2002. Taxonomy, morphology
and biology of Quarternary and living Ostracoda. In The Ostracoda:
Applications in Quaternary Research, J. Holmes and A. Chivas, eds.
SCHIZOCYTHERIDAE AGU Geophysical Monograph Series 131: 5–36.
Kaesler, R. L. 1987. Superclass Crustacea. In Fossil invertebrates: 241-
Acuminocythere crescentensis Swain and Gilby, 1974 (“Pai- 258. R. S. Boardman, A. H. Cheetham, and A. J. Rowell, eds. London:
Blackwell Scientific Publs.
jenborchella” sp. A of Valentine, 1976). Morro Bay and off Cres- Kempf, E. K. 1980. Index and Bibliography of Nonmarine Ostracoda 1,
cent City and Coos Bay in 10 m–26 m. Valentine (1976) Index A. Geologisches Institut der Universitaet zu Koeln, Son-
reported the range in offshore waters as north of Point Con- derveroeffentlichungen, no. 35 (and later volumes in this series).
ception to Cape Flattery (plate 196C). Kempf, E. K. 1986. Index and Bibliography of Marine Ostracoda, Vol. 1
Acuminocythere sp. of Swain and Gilby, 1974 (“Paijen- Index A. Geologisches Institut der Universitaet zu Koeln, Son-
derveroeffentlichungen, no. 50 (and later volumes in this series).
borchella” sp. B of Valentine, 1976). Empty carapace in mud
Maddocks, R. F. 1982. Ostracoda. In The biology of crustacea, vol. I: sys-
and fine sand, flat exposed at low tide, in Bodega Harbor, RFM. tematics, the fossil record and biogeography: 221–239, L. G. Abele,
Valentine (1976) reported the range in offshore waters as Pt. ed. New York: Academic Press.
Buchon to Little River, CA (plate 196D). Maddocks, R. F. 1992. Ostracoda. In Microscopic anatomy of inverte-
Spinileberis hyalina Watling, 1970. On bottom silt and clay, 2 brates 9: crustacea. F. W. Harrison and A. G. Humes, eds. 415–442.
New York: Wiley-Liss, Inc.
m, in Tomales Bay. See Watling 1970, Crustaceana 19: 251–263
Martens, K., ed. 1998. Sex and Parthenogenesis. Evolutionary ecology
(plate 191F). of reproductive modes in non-marine ostracods. Leiden: Backhuys
Publ., 336 pp.
Moore, R. C., ed. 1961. Treatise on Invertebrate Paleontology, Part Q
XESTOLEBERIDIDAE
Arthropoda 3 Crustacea Ostracoda. Lawrence: Geological Society of
America and University of Kansas Press, 442 pp.
Xestoleberis hopkinsi Skogsberg, 1950. On holdfasts of algae Morin, J. G., and A. C. Cohen. 1991. Bioluminescent displays, courtship,
in rocky tide pool full of brown algae, just outside the Hopkins and reproduction in ostracodes. In Crustacean Sexual Biology, R.
Marine Station, Monterey Bay (plate 194J). Bauer and J. Martin eds: 1–16. New York: Columbia University Press.
Morkhoven, F. P. C. M. 1962, 1963. Post-Palaeozoic Ostracoda, Their
Xestoleberis sp. On mud and fine sand flat exposed at low tide, Morphology, Taxonomy and Economic Use 1, 2. Amsterdam: Else-
Bodega Harbor, RFM. It is likely that additional species of vier, 204 and 478 pp.
Xestoleberis occur within our range (plate 194A–194I). Oakley, T., and C. Cunningham. 2002. Molecular phylogenetic evidence
for the independent evolutionary origin of an arthropod compound
eye. Proceedings of the National Academy of Science 99: 1426–1430.
Smith, R. J., T. Kamiya, and D. J. Horne. 2006. Living males of the “an-
ACKNOWLEDGMENTS
cient asexual” Darwinulidae (Ostracoda: Crustacea). Proceedings of
the Royal Society B: 10 pp.
We thank Todd Oakley, Jeff Spees, and James T. Carlton for as- Spears, T., and Abele, L. G. 1998. Crustacean phylogeny inferred from
sistance and for providing new collections of Ostracoda. We 18S rDNA. In Arthropod relationships, systematics association spe-
also thank Frederick M. Swain for the loan of specimens de- cial volume series 55, Fortey, R. A. and R. H. Thomas, eds. 169–187.
London: Chapman & Hall.
scribed by Swain and Gilby (1974) from the University of Min-
Wingstrand, K. G. 1988. Comparative spermatology of the Crustacea
nesota Paleontological Collections; Louis Kornicker and Entomostraca. 2. Subclass Ostracoda. Biologiske Skrifter 32: 1–149.
Elizabeth Nelson for literature and information regarding my-
odocopans and for the loan of the half-tone figures (drawn by Some Additional References (but not cited in Faunal List):
Carolyn Gast) in plate 187; Ginny Allen for redrawing many
myodocopid figures; and the Bodega Marine Laboratory for
OSTRACODA OF NORTHWEST AMERICA
providing lab space for Cohen and Oakley during collections
and for other assistance.
Benson, R. H. 1959. Ecology of recent ostracodes of the Todos Santos
Bay region, Baja California, Mexico. University of Kansas Paleonto-
logical Contributions, Arthropoda, Article 1: 1–80.
Benson, R. H., and R. L. Kaesler. 1963. Recent marine and lagoonal os-
References tracodes from the Estero de Tastiota region, Sonora, Mexico (north-
eastern Gulf of California). University of Kansas Paleontological
GENERAL REFERENCES ON OSTRACODA Contributions, Arthropoda, Article 3: 1–34.
Brouwers, E. M. 1983. Occurrence and distribution chart of ostracodes
Athersuch, J., D. J. Horne, and J. E. Whittaker. 1989. Marine and from the northeastern Gulf of Alaska. U.S. Geological Survey, Mis-
Brackish Water Ostracods, Synopses of the British Fauna (New Se- cellaneous Field Studies Map MF-1518, Pamphlet, pp. 1–13.
ries) (D. M. Kermack and R. S. K. Barnes, eds.) 43: 1–343. New York: Brouwers, E. M. 1988. Paleobathymetry on the continental shelf based
E. J. Brill. on examples using ostracods from the Gulf of Alaska. In Ostracoda
Cohen, A. C., J. W. Martin, and L. S. Kornicker. 1998. Homology of in the earth sciences. P. De Deckker, J.-P. Colin and J.-P. Peypouquet,
Holocene ostracode biramous appendages with those of other crus- eds.: pp. 55–76. Amsterdam: Elsevier.
OSTRACODA 445
Brouwers, E. M. 1990. Systematic paleontology of Quaternary ostracode of their life history stages. Benthic and epibenthic harpacticoid
assemblages from the Gulf of Alaska, Part 1. Families Cytherellidae, copepods often dominate the diets of flatfish and several
Bairdiidae, Cytheridae, Leptocytheridae, Limnocytheridae, Eucytheri-
species of Pacific salmon during their early life histories.
dae, Krithidae, Cushmanideidae. U.S. Geological Survey Professional
Paper 1510: 1–43. Given their importance in nearshore and estuarine food
Brouwers, E. M. 1993. Systematic paleontology of Quaternary ostracode webs, one would expect to find a number of ecological and tax-
assemblages from the Gulf of Alaska, part 2. Families Trachyleberididae, onomic studies of copepods of coastal California and Oregon.
Hemicytheridae, Loxoconchidae, Paracytherideidae. U.S. Geological However, this is not the case. Even though coastal and estuar-
Survey Professional Paper 1531: 1–47.
ine ecosystems have undergone and continue to undergo rapid
Ishizaki, K., and F. J. Gunther. 1974. Ostracoda of the Family Cytheruri-
dae from the Gulf of Panama. Science Reports of the Tohoku Uni- changes and increasing stress due to land use, urbanization, and
versity, Sendai, Second Series (Geology) 45: 1–50. introduced species, there have been no long-term studies of
Ishizaki, K., and F. J. Gunther. 1976. Ostracoda of the Family Loxo- plankton or meiobenthic dynamics in this region. Likewise,
conchidae from the Gulf of Panama. Science Reports of the Tohoku taxonomic compendia of copepods for the Pacific coast are few.
University, Sendai, Second Series (Geology) 46: 11–26.
Esterly (1905, 1906, 1911, 1924) reported on marine plankton
McKenzie, K. G. 1965. Myodocopid Ostracoda (Cypridinacea) from
Scammon Lagoon, Baja California. Crustaceana 9: 57–70.
of San Francisco Bay and the San Diego region, Dawson and
McKenzie, K. G., and F. M. Swain. 1967. Recent Ostracoda from Scam- Knatz (1980) published a list and keys of the planktonic cope-
mon Lagoon, Baja California. Journal of Paleontology 41: pods of San Pedro Bay, and Gardner and Szabo (1982) give keys
281–305. and an annotated bibliography for pelagic marine copepods of
Swain, F. M., and J. M. Gilby. 1967. Recent Ostracoda from Corinto Bay, British Columbia. Lang (1965) produced what remains one of
western Nicaragua, and their relationship to some other assemblages
of the Pacific Coast. Journal of Paleontology 41: 306–334.
the finest treatments of a local fauna for harpacticoid copepods
Valentine, P. C. 1976. Zoogeography of Holocene Ostracoda off western from interstitial waters and tide pools near the Hopkins Marine
North America and paleoclimatic implications. United States Geo- Station in Pacific Grove and the now-gone Pacific Marine Sta-
logical Survey Professional Paper 916, 47 pp. tion (formerly located in Dillon Beach). He described 98 species
of harpacticoids, of which 81 were new.
Thus the taxonomy of the copepod faunas of many Califor-
EXAMPLES OF DIVERSITY IN LITTORAL ZONES
nia nearshore habitats remains little-studied, including those
from salt marshes, estuaries, seagrass beds, sandy beaches,
Schornikov, E. I. 1974. Kizucheniuo ostrakod (Crustacea, Ostracoda)
litorali Kuril’ckix ostrovov: 137–214. Rastitel’n’ii i Zhivotn’ii Mir mudflats, rocky shores, fouling communities, and the shallow
Litorali Kuril’skix Ostrovov (in Russian). subtidal. Many species remain undescribed, and an unknown
Schornikov, E. I. 1975. Ostracod fauna of the intertidal zone in the number of introduced taxa remain undetected. Great care must
vicinity of the Seto Marine Biological Laboratory. Publications Seto be exercised in attempting identification of copepods from
Marine Biological Laboratory 22: 1–30.
these habitats. Unfortunately, the primary taxonomic litera-
Williams, R. 1969. Ecology of the Ostracoda from selected marine in-
tertidal localities on the coast of Anglesey. In The taxonomy, mor-
ture usually must be used to identify the most common inter-
phology and ecology of recent ostracoda, J. W. Neale, ed.: pp. tidal copepods, the Harpacticoida, to genus or species levels.
299–329. Edinburgh: Oliver and Boyd. However, there are several publications that have keys to
higher taxonomic levels. Huys et al. (1996) provide a key to
world harpacticoid families, and Boxshall and Halsey’s An In-
PREPARATION OF SPECIMENS troduction to Copepod Diversity (2004) contains keys to families
of marine planktonic, marine benthic, and fish parasite cope-
Cohen, A. C., and J. G. Morin. 1986. Three new luminescent ostracodes
pods, and to genera of many copepod families. The latter book
of the genus Vargula from the San Blas region of Panama. Contribu-
tions in Science, Natural History Museum of Los Angeles County, is also an excellent starting point for anyone wanting to study
373: 1–23 (anesthetics, etc.). copepod classification.
Cohen, A. C., and J. G. Morin. 1997. External Anatomy of the Female
Genital (Eighth) Limbs and the Setose Openings in Myodocopid Os-
tracodes (Cypridinidae). Acta Zoologica 78: 85–96 (SEM, and refer-
EXTERNAL STRUCTURE OF THREE ORDERS
ences therein).
OF FREE-LIVING COPEPODA
Of the nine copepod orders currently recognized, three—

Copepoda
Calanoida, Cyclopoida, and Harpacticoida—contain most of
JEFFERY R. CORDELL the free-living individuals likely to be found in nearshore habi-
tats. A fourth order, the Poecilostomatoida, has been placed
Free-Living Copepoda (Orders Calanoida, Cyclopoida, into the Cyclopoida (Boxshall and Halsey 2004).
and Harpacticoida) Copepods have developed two basic body plans, gymnoplean
and podoplean, which are defined by the position of the ma-
(Plates 197–206)
jor body articulation between prosome and urosome (plate
Copepods have been compared to insects, because, like their 197). In the gymnoplean plan (Order Calanoida), this articula-
terrestrial counterparts, they have successfully occupied an as- tion is behind the fifth pedigerous somite (plate 197A), and in
tounding diversity of habitats and modes of life (see Huys and the podoplean plan (Orders Cyclopoida and Harpacticoida), it
Boxshall 1991 for a summary of copepod habitats). In terms of is between the fourth and fifth pedigerous segments (plate
their importance in marine food webs, the role of copepods 197B). The prosome consists of the cephalosome that bears the
cannot be overstated. By way of their conversion of detritus first six pairs of appendages together with three or four free pro-
and phytoplankton into animal biomass, copepods often form somites, which are sometimes referred to as the metasome and
the first link between primary and secondary consumers. Many bear four or five pairs of swimming legs (plate 197A, 197B). In
commercially important fish, including herring, anchovies, most harpacticoids, the somite bearing the first pair of legs is
and rockfish feed on planktonic copepods during some or all also fused to the cephalosome and together they form the
446 ARTHROPODA

Key To The Ostracods 2007

Uploaded by

Document Information

Original Title

Copyright

Available Formats

Share this document

Share or Embed Document

Sharing Options

Did you find this document useful?

Is this content inappropriate?

Copyright:

Available Formats

Key To The Ostracods 2007

Uploaded by

Copyright:

Available Formats

2007. Cohen, Anne C., Dawn E. Peterson, and Rosalie F. Maddocks. Ostracoda, pp.

Key to the Ostracoda — Carapace ovoid, inflated-subtriangular, oblong elongate,

Tabular Key to Myodocopa (and Subclass only of Podocopa)

Body (plate 184)

Medial eye With or without With or without Present in most Absent

Furca (plates 184, 185D, 185H, 186F, 188E, 191E)

Furcal position Anterior to anus Posterior to anus

Best developed Endopod (plates Exopod (plate 184)

Mandible (plates 185C, 188A–188D, 190G, 192E, 193G, 194B)

Sexual dimorphism n.g. Some Dimorphic in Sarsielloidea Apparently none

4th Limb (plates 184, 190D, 190E, 192D, 193E)

Sexual dimorphism n.g. Present in some Reduced only in Apparently none

6th Limb (plates 184, 190I, 191D, 191E, 193D, 194F)

Limb present Present Present or absent Present Present or absent

Endopod Present Present Rami unclear Present

7th Limb (plates 190J, 191D, 191E, 192H, 193E, 194G)

Paired or single Paired Paired or single Paired Single

Tabular Key to Suborders and Superfamiliies of Order Halocyprida

Suborder Polycopina Suborder Halocypridina

No. of articles 3–5 (inferred 8) 3–8 7–8 3–8

Suborder Polycopina Suborder Halocypridina

Coxal endite Cusped tooth Cusped tooth

Exopod Present (one to two Absent

Endopod—number of One article Jointed (number of

Sixth Limb: presence

Seventh Limb: presence

Absent Reduced (or absent), Unjointed One to two articles

Male Copulatory Limb

NOTE : Key differences are indicated in boldface.

— Carapace (plates 191F, 196C, 196D) elongate-ovate to sub- List of Species

Tabular Key to Families of the Order Myodocopida

Subfamily Subfamily Subfamily

Character Cypridinidae Cylindroleberididae Philomedidae Rutidermatidae Sarsiellidae

Bellonci organ Short or long Usually long Long Long Long

Subfamily Subfamily Subfamily

Character Cypridinidae Cylindroleberididae Philomedidae Rutidermatidae Sarsiellidae

Males with Yes No No No No

4th Limb (plate 184A, 184B)

Reduced and weak No No Yes Yes Yes

Subfamily Subfamily Subfamily

Character Cypridinidae Cylindroleberididae Philomedidae Rutidermatidae Sarsiellidae

5th Limb (plate 184A, 184B)

Reduced, weak in No No Yes Yes Yes

6th Limb (plate 184A, 184B)

With well deve- 4 All reduced 4 4 1–4

7th Limb (plate 184A, 184B)

Reduced in males? No No In a few No Usually

Male 8th Limb (plate 184A)

NOTE : Key differences are indicated in boldface.

bution, ecology) (plate 185F–185H)

Sarscypridopsis aculeata (Costa, 1847) (Cypridopsis aculeata). CYTHEROCOPINA

m, outside Hopkins Marine Station near Pacific Grove; also on LOXOCONCHIDAE

P LATE 19 6 Podocopida: Hemicytheridae, Loxoconchidae, Pectocytheridae, Schizocytheridae. Pectocytheridae: A, Pectocythere park-

Of the nine copepod orders currently recognized, three—

You might also like