140

Cor Pyloric Dorsal frontalle stomach abdominal Supraesophageal art ry ganglion . . K Ophthalmic / Ostium ® Antennary artery Cardiac / / Heart \ Segmental Optic \ artery i stomach/Cecum / / Hepato- \ artery nerve I / / / / / pancreas\ Hindgut Antennal nerve Rectum

ganglion

Hepatic artery Subesophageal Ventral Midgut ganglion nerve cord

dorsomedial branchiocardiac dorsomedial Antenna Antennule intestinal / urogastric Compound eye posterior margina

hepatic Th°racopods

lateromarginal I inferior \ intercervical parabranchial postcervical B

Uropod Telson

Figure 47 Decapoda: A. Diagrammatic astacidean with gills and musculature removed to show major organ systems; B. Diagrammatic nephropoidean carapace illustrating carapace grooves [after Holthuis, 1974]; C. Phyllosoma larva. ORDER DECAPODA 141 midgut or the other, pierces the ventral nerve cord, and indistinct from the fused ganglia of the mandibles, maxil- then branches anteriorly and posteriorly. The anterior lulae, maxillae, and first 2 pairs of maxillipeds. The gang­ branch, the ventral thoracic artery, supplies blood to the lia of the first 3 pairs of pereopods are segmental; the mouthparts, nerve cord, and 1st 3 pairs of pereopods. ganglia of the 4th and 5th pairs lie very close together. The course of this artery cannot be traced until the Follow the ventral nerve cord into the abdomen and stomach and hepatic cecum have been removed. The identify the abdominal ganglia. posterior branch, the ventral abdominal artery, which Larval development is direct (epimorphic) in all fresh­ also should be traced later, provides blood to the 4th and water taxa; in marine taxa early developmental stages 5th pairs of pereopods, nerve cord, and parts of the ven­ are passed through in the egg and hatching usually occurs tral abdomen. Returning to the heart, follow the dorsal at the my sis stage. Development at this stage corre­ abdominal artery posteriorly through the abdomen. In sponds to similar stages in other decapods. each somite a pair of segmental arteries are given off that supply blood to the midgut, abdominal musculature, and pleopods. The posterior pair supplies the uropods and telson. References Remove the heart and underlying pericardial mem­ Balss, H., 1940-1957. Decapoda. In H. G. Bronn, Klassen brane to expose the gonad. In females the gonad typi­ und Ordnungen des Tierreichs, 5(1) 7: 321—480. Leipzig: cally is Y-shaped, with lobes extending anteriorly on Akad. Verl. either side of the stomach. The male testis is similarly Bott, R., 1950. Die Flusskrebse Europas (Decapoda, As- positioned. If possible trace the oviducts or vas deferens tacidae). Abhandl. Senckenberg. Naturf. Ges., 483: 1—36. from their origins on the gonad ventrally to the gono- Budd, T. W., J. C. Lewis, and M. L. Tracey, 1978. The filter- pores on the coxae of the pereopods (3rd in females, 5th feeding apparatus in crayfish. Canad. J. Zool., 56: 695— 707. in males). Caiman, W. T., 1909. Treatise on zoology. Pt. 7, Appen- If the anterior cephalothoracic muscles have not yet diculata, Fasc. 3, Crustacea. 346 pp. London: Ray Soc. been removed, remove them at this time to expose the Creaser, E. P., 1933. Descriptions of some new and poorly esophagus and stomach. The esophagus is short and known species of North American crayfishes. Occ. Pap. opens into the cardiac portion of the stomach. A constric­ Mus. Zool, Univ. Mich., 275: 1-21. tion marks the boundary between the cardiac and pyloric Crocker, D. W., and D. W. Barr, 1968. Handbook of the parts of the stomach. At the junction of the pyloric crayfishes of Ontario. 158 pp. Toronto: Roy. Ontario Mus., stomach and midgut, a pair of ducts from the hepatopan- Univ. Toronto Press. creas open into the midgut. In addition to the prominent Glaessner, M. E, 1969. Decapoda. In R. C. Moore (ed.), bilobed hepatopancreas, locate the small cecum that Treatise on invertebrate paleontology, Pt. R, Arthropoda 4, projects anteriorly adjacent to the pyloric stomach. Just 2: R399— R566. Lawrence, Kans.: Geol. Soc. America and posterior to this cecum a slight ridge indicates the junc­ Univ. Kansas. tion of the midgut and hindgut; in most crayfish the latter Hobbs, H. H., Jr., 1942a. A generic revision of the crayfishes of extends nearly the entire length of the abdomen. In the the subfamily Cambarinae (Decapoda, Astacidae) with the 6th abdominal somite it usually enlarges to form a rec­ description of a new genus and species. Am. Midi. Natur., tum before terminating at the anus on the ventral sur­ 28:334-357. face of the telson. Remove the stomach and examine the , 1942b. The crayfishes of Florida. Univ. Florida Publ. structure of the gastric mill in the cardiac portion of the Biol. Sci. Ser.,3: 1-179. stomach. The pyloric stomach is made up of plates, bars, , 1967. The current status of the crayfishes listed by and channels. How does this stomach compare with that Girard (1852) in his "A revision of the North American Astaci of the penaeid? Excretion in astacideans is via antennal . . ." Crustaceana 12: 124-132. glands. In contrast to the condition found in penaeoi- , 1968. Crustacea: Malacostraca. In F. K. Parrish, Keys deans, the antennal gland of astacideans is distinct. Lo­ to water quality indicative organisms (Southeastern United cate this gland and trace its duct to the excretory pore at States). Fed. Water Poll. Contr. Admin. Dept. Interior: the base of the antenna. K1-K36. Reissued 1975. , 1969. On the distribution and phylogeny of the The supraesophageal ganglion lies in the midline just crayfish genus Cambarus. In P. C. Holt (ed.), The distribu­ ventral to the ocular peduncles. Locate this ganglion and tional history of the biota of the Southern Appalachians, Part identify the 4 large pairs of nerves radiating from it, the I: Invertebrates. Virginia Polytechnic Inst. Res. Div. antennular, optic, antennal, and tegumental nerves. Monogr. J: 93-178. Ventrally trace the esophageal connective of one side Hobbs, H. H., Jr., and A. Villalobos, 1964. Los Cambarinos de around the esophagus; it passes beneath the endo- Cuba. An. Inst. Biol. Univ. Nal. A. de Mexico 84: 307-366. phragmal shelf before joining the subesophageal ganglion. Huxley, T. H., 1880. The crayfish. Intern. Sci. Series 28. New Portions of this shelf must be removed to expose the York: D. Appleton and Co. 371 pp. (Reprinted 1973, MIT ventral thoracic ganglia. The subesophageal ganglion is Press). 142 CLASS MALACOSTRACA

Kingsley, J. S., 1899. Astacoid and thallassinoid Crustacea. Rhoades, R., 1944. The crayfishes of Kentucky, with notes on Synopses of North American invertebrates. IV. Am. Natur., variation, distribution and descriptions of new species and 33:819-824. subspecies. Am. Midi. Natur. 31: 111— 149. Meredith, W. G., and F. J. Schwartz, 1960. Maryland cray­ Rick, E. E, 1969. The Australian freshwater crayfish (Crus­ fishes. Maryland Dept. Res. Ed. Educational Series 46, 32 tacea: Decapoda: Parastacidae), with descriptions of new pp. species. Aust. J. Zool., 17: 855— 918. Ortmann, A. E., 1906a. Mexican, Central American and , 1971. The freshwater crayfishes of South America. Cuban cambari. Proc. Wash. Acad. Sci., 8: 1—24. Proc. Biol. Soc. Wash., 84: 129- 136. , 1906b. The crawfishes of the state of Pennsylvania. Schwartz, F. J., and W. C. Meredith, 1960. Crayfishes of the Cheat River watershed in West Virginia and Pennsylvania. Mem. Carnegie Mus. 2: 343- 523. Parti. Species and localities. Ohio J. Sci., 60: 1—46. , 1907. Grabende Krebse in Nordamerika. Aus der , 1962. Crayfishes of the Cheat River watershed in West Natur, 1906/07: 705-716. Virginia and Pennsylvania. Part II. Observations upon , 1931. Crawfishes of the southern Appalachians and the ecological factors relating to distribution. Ohio J. Sci., 62: Cumberland Plateau. Ann. Carnegie Mus. 20: 61—160. 260-273.

INFRAORDER AUSTROASTACIDEA Clark, 1936

Recent species Few in one genus, Austroastacus. Size range 10 to ~ 80 mm. Carapace Strongly vaulted posteriorly. Eyes Stalked, compound, well developed. Antennules With 2nd flagellum small or absent. Antennae Inserted beneath antennules; scaphocerite long or short, slen­ der, sometimes with long slender spine. Mandibles (?) With palp. Maxillulae (?) With endopodal palp. Maxillae (?) Biramous. Maxillipeds Epipods of 1st without branchial filaments; 3rd without exo- pods. Thoracic appendages Uniramous; first 3 pairs chelate; 5th pereopods without gills. Abdominal appendages Pleopods biramous; 1st pair absent in both sexes. Uropods biramous, without transverse sutures, well calcified. Telson Entirely calcareous. Tagmata Cephalothorax and abdomen. Somites Head with 5+3 thoracic (maxillipeds); thorax with 5; abdomen with 6, excluding telson. Sexual characters Gonopores on coxae of 3rd pereopods of female; on short, sim­ ple papillae on coxae of 5th pereopods of male. Sexes Presumably separate. Larval development Presumably epimorphic. Fossil record Recent. Feeding types Presumably herbivores. Habitat Burrows in swampy and dry areas. Distribution Australia.

The type species, Austroastacus hemicirratulus (Smith ean genus Engaeus. Clark (1936) removed A. hemicir­ and Schuster), of this small group of semiterrestrial Aus­ ratulus from its former genus and assigned it to his newly tralian crayfishes originally was described in the astacid- established Austroastacus, and at the same time de- ORDER DECAPODA 143 scribed for it the family Austroastacidae. Bowman and sides lack cones; the entrances are indicated only by Abele (in press) have since elevated the family to infra- small round holes, usually beneath fallen logs or stones. order rank. Several characters distinguish the austroastacideans References from other crayfish. The telson and uropods are com­ pletely calcified. The first pair of pleopods are absent in Bowman, T. E., and L. G. Abele, in press. Classification. In L. both sexes, and the abdomen is relatively small, al­ G. Abele (ed.) The biology of Crustacea, vol. 1, Systematics, though larger in females than in males. The carapace is Genetics, Morphology and the Fossil Record. New York: higher than broad and vaulted posteriorly; the cervical Academic Press. and branchiocardiac grooves are strongly impressed and Clark, E., 1936. The freshwater and land crayfishes of Aus­ the areola narrow. The inner flagella of the antennules, tralia. Mem. Natl. Mus. Melbourne, 10: 5-58. when present, are much shorter than the outer. Glaessner, M. E, 1969. Decapoda. In R. C. Moore (ed.), These crayfish live in family groups and dig deep bur­ Treatise on invertebrate paleontology, Part R Arthropoda, 2: rows both in swampy patches and on dry hillsides in R399— R533. Lawrence, Kans.: Geol. Soc. America and heavily timbered areas. The entrances to the burrows Univ. Kansas. dug in swampy ground usually have cones a few inches Healy, A., and J. Yaldwyn, 1970. Australian crustaceans in high with an opening at the top. Burrows dug on hill­ colour. 112 pp. Sydney: A. H. and A. W. Reed.

INFRAORDER PALINURA Latreille, 1803

Recent species Approximately 130. Size range Up to 61 cm. Carapace Cylindrical, subovoid, or dorsoventrally compressed. Eyes Stalked, compound, generally well developed. Antennules Biramous; peduncle with 3 segments. Antennae Peduncle with 5 or fewer segments; exopod reduced; flagella sometimes modified. Mandibles Usually with palp; molar and incisor processes usually not dis­ tinct. Maxillulae With endopodal palp. Maxillae Biramous; with 2 bilobed endites. Maxillipeds Endopod of 1st with 2 or fewer segments; usually with crista dentata. Thoracic appendages Uniramous; 1st pair sometimes chelate; other pairs often sub- chelate. Abdominal appendages Pleopods usually biramous; 1st pair frequently absent; male copulatory structures frequently present. Uropods biramous. Telson Together with broad uropods forms tailfan. Tagmata Cephalothorax and abdomen. Somites Head with 5-1-3 thoracic (maxillipeds); thorax with 5; abdomen with 6, excluding telson. Sexual characters Gonopores on coxae of 3rd pereopods of female, on 5th of male; male frequently with copulatory structure. Sexes Presumably separate. Larval development Metamorphic; phyllosoma —» puerulus (postlarva) [Palinu- roidea]. Fossil record (?) L. Triassic, M. Triassic to Recent. Feeding types Various. Habitat Marine. Distribution Worldwide. 144 CLASS MALACOSTRACA

This infraorder contains three superfamilies; however, , 1940. Decapodes marcheurs. In Faune de France, 37: specimens from only one, the Palinuroidea, can be ob­ 1-399. Paris: Paul Lechevalier. tained relatively easily. It contains, among others, the Caiman, W. T., 1909. Treatise on zoology. Pt. 7, Appen- Palinuridae (spiny lobsters) and the Scyllaridae (slipper diculata, Fasc. 3, Crustacea, 346 pp. London: Ray Soc. or shovel-nosed lobsters). If specimens are available, Corrivault, G. W. and J. L. Tremblay, 1948. Contribution a la representatives of these taxa should be compared with biologie du homard (Homarus americanus Milne-Edwards) the crayfishes and true lobsters (see Figure 46). In scyl- dans la Baie-des-Chaleurs et le golfe Saint-Laurent. Contr. larids, the great modification of the antenna is the most StnBiol. St. Laurent, 19: 1-222. striking character. The antennal flagella have been trans­ Dawson, C. E., 1954. A bibliography of the lobster and the formed into spinose or tuberculate platelike structures; spiny lobster, families Homaridae and Palinuridae. 86 pp. the first segments of the antennular peduncles are im­ Florida State Board of Conservation. movable. In the Palinura the rostrum is not prominent, Firth, R. W., and W. E. Pequegnat, 1971. Deep-sea lobsters of and the carapace is fused to the epistome. In the the Families Polychelidae and Nephropidae (Crustacea De­ Palinuridea, the 5th pereopods of females are the only capoda) in the Gulf of Mexico and Caribbean Sea. 103 pp. chelate appendages; however, chelate appendages usu­ Glaessner, M. R, 1969. Decapoda. In R. C. Moore (ed.), ally are present in the other superfamilies. Treatise on invertebrate paleontology, Pt. R, Arthropoda 4, 2: R399—R566. Lawrence, Kans.: Geol. Soc. America and Larval development is typically metamorphic. Early Univ. Kansas. larval stages are passed through in the egg; the first post- Herrick, F. H., 1895. The American lobster: a study of its embryonic stage in the Palinuroidea is the phyllosoma. habits and development. Bull. U.S. Fish. Comm., 15: This is a very distinctive larval form, with an extremely 1-252. flattened, disk-shaped body and elongate appendages; it Holthuis, L. B., 1946. The Stenopodidae, Nephropsidae, Scyl­ is equivalent to the my sis stage. The postlarval stage in laridae and Palinuridae. The Decapoda Macrura of the this superfamily is referred to as the puerulus, nisto, or "Snellius" Expedition. Temminckia, 7: 1— 178. pseudibacus stage, a benthic form more closely resem­ , 1974. The lobsters of the superfamily Nephropidea of bling the adult. the Atlantic Ocean (Crustacea: Decapoda). Bull. Mar. Sci., 24: 723-884. Lewis, R. D., 1970. A bibliography of the lobsters, genus Ho­ References marus. U.S. Fish Wildl. Serv. Spec. Sci. Rept. Fish., 591: Alcock, A., 1901. A descriptive catalogue of the Indian deep- 1-47. sea Crustacea Decapoda Macrura and Anomala in the In­ Maigret, J., 1976. Contribution a 1'etude des langoustes de la dian Museum. 286 pp. Calcutta: Indian Museum. cote occidentale dAfrique (Crustaces, Decapodes, Palinu­ ridae). 1. Notes sur la biologie et l'ecologie des especes sur Balss, H., 1940-1957. Decapoda. In H. G. Bronn, Klassen les cotes du Sahara. Bull. Inst. Fondam. Afr. Noire, A38: und Ordnungen des Tierreichs 5(1) 7: 321-480. Leipzig: 266-302. Akad. Verl. Bonde, C. von, and J. M. Marchand, 1935. The natural history Stephensen, K., 1923. Decapoda-Macrura excl. Sergestidae. and utilization of the Cape crawfish, Kreef, or spiny lobster, Rept. Danish Oceanogr. Exped. Mediterranean, 2 (D3): Janus (Palinurus) lalandii (Milne Edwards) Ortmann. Fish. 1-85. Bull. Fish Mar. Biol. Sur. S. Afr., 1: 1-55. Wolff, T, 1978. Maximum size of lobsters (Homarus) (De­ Bouvier, E. L., 1925. Les Macroures Marcheurs. Reports of capoda, Nephropidae). Crustaceana, 34: 1— 14. the results of dredging in the Gulf of Mexico (1877- 78), in Zarenkov, N. A., and V. N. Semenov, 1972. Novyi vid roda the Caribbean Sea (1878— 79), and along the Atlantic coast of Nephropides (Decapoda, Macrura) iz yugo-zapadnoi Atlan- the United States (1880) by the U.S. Coast Survey Steamer tiki. Zool. J. Moscow, 51: 599-601. "Blake" . . . Mem. Mus. Comp. Zool. Harvard, 47: 397— Zariquiey Alvarez, R., 1968. Crustaceos Decapodos Ibericos. 472. Invest. Pesqueras Barcelona, 32: 1—510.

INFRAORDER ANOMURA H. Milne Edwards, 1832

Recent species Approximately 1575. Size range Up to 185 cm (leg span). Carapace Variable in shape; not fused to epistome. Eyes Stalked, compound; generally well developed. Antennules Peduncle with 3 segments; flagella usually paired. Antennae Peduncles sometimes with 5, possibly 6, or fewer segments; exopod usually reduced to acicle; flagellum variable in length. ORDER DECAPODA 145

Mandibles With or without palp; molar and incisor processes usually not distinct. Maxillulae With endopodal palp. Maxillae Biramous; usually with bilobed endites. Maxillipeds Flagella usually present; sometimes absent from 1st; often with crista dentata; usually not operculate. Thoracic appendages First usually, 2nd occasionally, chelate; 4th and 5th usually che­ late or subchelate, one or both frequently reduced. Abdominal appendages Pleopods rarely well developed; often reduced or present only on 1 side; both sexes sometimes with copulatory structures. Uropods often reduced or modified, occasionally absent. Telson Occasionally reduced or absent; sometimes together with uropods forms tailfan. Tagmata Cephalothorax and abdomen. Somites Head with 5+3 thoracic (maxillipeds); thorax with 5; abdomen with 6, excluding telson, although segmentation frequently obscured. Sexual characters Gonopores on coxae of 3rd pereopods of female, on 5th of male; 1st and/or 2nd pleopods often modified as gonopods in both sexes; female sometimes with abdominal brood pouch; male sometimes with sexual tube(s). Sexes Presumably separate. Larval development Metamorphic; zoea—» megalopa (glaucothoe). Fossil record L. Jurassic to Recent. Feeding types Various. Habitat Most frequently marine, but with few semiterrestrial and fresh­ water species. Distribution Worldwide.

Among the decapod crustaceans, the infraorder Anom- Begin your study with an overall examination of the ura presents the greatest diversity in body form; there general morphology of a typical hermit crab. Aside from are five superfamilies. The shrimplike or crayfishlike the membranous abdomen, perhaps the most noticeable elongate body is represented by the Thalassinoidea. character and one that is not common in other crusta­ The abdomen, although often not drastically reduced in ceans is the strong tendency toward asymmetry. De­ length in the Coenobitoidea and Paguroidea, is most fre­ pending upon the species under study, the left or the quently membranous or only weakly calcified and an ex­ right 1st pereopod (cheliped) usually is noticeably en­ trinsic protective covering often is adopted. In certain larged. A second indication of asymmetry is the presence families the familiar crablike body form has been devel­ of pleopods on only the left side of the abdomen; some oped (i.e., Lomidae and Lithodidae). In these families, primitive genera of coenobitoideans have paired pleo­ the abdomen is tucked under the cephalothorax; how­ pods. Usually, unless modified as gonopods, pleopods ever, in contrast to that of brachyurans, the abdomen are absent from the 1st abdominal somite. Sexual mod­ remains relatively soft and membranous or is covered ifications of the 1st pair may occur in both male and fe­ by individual calcified plates. The superfamily Galathe- male; in the male the 2nd also may be modified. In some oidea has representatives with crayfishlike body forms genera of pagurids the vas deferens is markedly ex­ (Galatheidae) and crablike forms (Porcellanidae), while tended from the gonopores to form 1 or a pair of sexual representatives of the Hippoidea exhibit variations be­ tubes. The asymmetrical uropods common to the major­ tween the two extremes. If a number of anomuran types ity of paguroideans are another example of the secondary are available, identify the characters that are possessed asymmetry characteristic of the group. in common. As the paguroidean body form is perhaps In dorsal view, the eyes with their prominent pedun­ the most atypical, a pagurid will be used as a basis for cles usually are obvious. At the base of each ocular discussing anomuran characters (see Figures 48 and 49). peduncle observe the ocular acicle; it may have several 146

Dactyl Fixed finger Palm(propodus) Carpus Merus

Pleopod Anterolateral Developing eggs plate Abdomen Intermediate plate

Figure 48 Decapoda, Anomura: A, B, Paguroidea; C, D, Galatheoidea. A. Typical pagurid (dorsal view); B. Pagurid thorax (ventral view); C. Typical galathoidean (dorsal view), showing regions of carapace; D. Typical porcellanid. 147

Antennule Ocular peduncle —Major cheliped Antenna

Branchio- stegite Dactyl

Antennal. Propodus flagellui

Dactyl

Figure 49 Decapoda, Anomura; A, B, Thalassinoidea; C—K, Hippoidea. A. Typical callianassid (dorsal view); B. 3rd maxilliped; C. Typical hippid (lateral view); D. Hippid telson; E. Hippid 3rd maxilliped; F. Typical albuneid (dorsal view); G. Albuneid 5th pereopod; H. Albuneid 3rd maxilliped; I. Typical Lepidopa (Albuneid); J. Left cheliped of Lepidopa; K. 3rd maxilliped of Lepidopa. 148 CLASS MALACOSTRACA spines, a single spine, or less frequently be unarmed. mit crabs. If specimens of lithodids and porcellanids are The antennules are located ventrally between the ocular available, what characters do you find that relate the peduncles; the 1st segment has a prominent statocyst. former closely with the pagurids and the latter with the The antennae are lateral to the ocular peduncles and galatheids? The superfamily Hippoidea is represented by appear to have, in addition to the typical 5 segments of a relatively few species, many of which are rather the peduncle, a small supernumerary segment between specialized. The common "mole crabs'' Emerita spp. the 3rd and 4th segments dorsolaterally. The antennal have the antennae extremely setose and adapted for filter exopod is reduced to a slender acicle. The pagurid feeding. Some of the Albuneidae have the antennules cephalothorax usually is calcified only in the anterior modified for a similar function. part, referred to as the shield. In some species the bran- The description of major organ systems also is based chiostegites are calcified as well. The shield is separated on the conditions found in pagurids (see Figure 50), al­ from the posterior part of the carapace by the cervical though these are not necessarily always typical of all groove. Usually 3 distinct pairs of lines or grooves can be anomurans. Differences between pagurids and other distinguished on the posterior carapace. In the midline anomurans will be pointed out. Remove the carapace are a pair of elongate sutures, the sulcus cardiobran- very carefully to avoid damaging the heart; the pericar­ chalis; 2 short lines or grooves slightly laterad are re­ dium attaches to the membrane underlying the cara­ ferred to simply as sulcus "a"; a linea anomurica on pace. As the cephalothoracic body cavity of typical pa­ each side of the carapace represents the 3rd pair. If the gurids is narrow, remove the gill chamber and inner branchiostegite is lifted or removed, the gills are ex­ branchial wall from one side. Locate the heart in the posed. Typically the gill number in pagurids varies from 4 posterodorsal region of the cephalothorax. The pericar­ 10 to 13 pairs, depending upon the taxa. What is the gill dium extends from the cervical groove to the 8th thoracic formula for your specimen(s)? somite, but usually will not be easily distinguished from In addition to the asymmetrical chelipeds (except in neighboring tissues. Blood flows from the gills into the the genera Paguristes and Clibanarius, and the family pericardium and into the heart via 3 pairs of ostia, 1 Pylochelidae), the 2nd and 3rd pereopods also may be anterodorsal pair and 2 lateral pairs. Anteriorly the heart asymmetrical and/or dissimilarly armed from right to gives off 3 arteries, the unpaired, median ophthalmic left. The 4th pereopods are usually considerably shorter artery and the paired lateral arteries. Follow the oph­ than the preceding pairs and usually are provided with a thalmic artery; shortly after leaving the heart a cor fron- rasp on the external surface of each propodus; in some tale is formed. After passing over the dorsal surface of taxa they have a preungal process at the base of each the stomach the ophthalmic artery turns ventrally and dactyl (the function of this structure is unknown). The divides into 2 branches that provide blood to the anterior 5th pereopods also are reduced in size and most fre­ cephalic area and supraesophageal ganglion. Returning quently are chelate. They too are equipped with rasps. to the heart trace the path of one of the lateral arteries Although the pagurid abdomen is usually membra­ anterolaterally. Enroute to the cephalic appendages, it nous, it often is possible to distinguish the outline of some gives off branches to the stomach and musculature. Ven­ of the abdominal tergites. The pagurid telson and trally from the heart locate the small hepatic arteries. In uropods are distinctive. Notice that the uropods also are pagurids this pair of arteries no longer supplies blood to provided with rasps. These are used by the animal to the hepatopancreas, which lies almost exclusively in the help in holding the shell in place. abdomen, but terminates instead on the stomach or Before removing the mouthparts, observe the position midgut. From the posterior margin of the heart identify of the 3rd maxillipeds relative to one another. This is a the large, posteriorly directed superior abdominal artery valuable diagnostic character; the two superfamilies are and the ventrally directed sternal artery. The path of the differentiated on the position of these maxillipeds. In the latter will be described, but should not be traced until Coenobitoidea the 3rd maxillipeds are approximate at after the stomach and midgut have been removed. Fol­ their bases; in the Paguroidea the bases are widely sepa­ low the superior abdominal artery posteriorly, cutting rated. The approximate position of the 3rd maxillipeds in open the abdomen if necessary. At the level of the 1st the Thalassinoidea and the widely separated condition in abdominal somite the artery branches. Trace the large the Galatheoidea suggest that the Coenobitoidea may be segmental artery ventrally; at the level of the 3rd somite closely related to the former and the Paguroidea to the it divides into sub- and supramuscular branches. The latter. If such relationships are true, the similarities in former passes ventrally along the ventral nerve cord and body form that exist among representatives of the latter terminates in the 6th somite. The latter provides numer­ two superfamilies would have to have resulted from con­ ous branches to the hepatopancreas and gonads and ter­ vergent evolution. minates with branches to the telson and uropods. The If specimens of galatheoideans or thalassinoideans are superior abdominal artery supplies blood to the gonads available, list the similarities and differences that you and to the pleopods. Follow the sternal artery as it passes observe between representatives of these taxa and her­ first ventrally into the 7th thoracic somite and then turns 149

Superior Heart Cor frontale abdomina Lateral artery artery Cardiac stomach Vas deferens Ophthalmic artery Midgut cecum Cephalic artery Segmental artery Supraesophageal ganglion Antennal gland Testis Esophageal connective

Posterior "hindgut" cecum

Figure 50 Decapoda, Anomura: A. Diagrammatic pagurid with musculature removed to show major organ systems; B. Zoea; C. Megalopa (Glaucothoe.) 150 CLASS MALACOSTRACA horizontally. At the level of the 5th somite it turns ven- abdominal somite. In the abdomen the nerve cord is of trally again and pierces the central ganglionic mass be­ the ladder type with 5 pairs of fused ganglia. As a result tween the nerves of the 2nd and 3rd pereopods. Beneath of the flexure of the abdomen, the nerve cord in pagurids the nerve cord the vessel divides into anterior and pos­ is skewed to the left from the 2nd to the 4th somites. The terior branches. The anterior branch, the inferior thoracic abdominal flexure also has resulted in the atypical devel­ artery, provides blood to the chelipeds, mouthparts, opment of the abdominal musculature. The flexor mus­ renal gland, and ventral region of the stomach. The pos­ cles have become abnormally enlarged and those of the terior branch supplies blood to the remaining pereopods, right side are considerably larger than those of the left. but in contrast to other anomurans, does not enter the Larval development is metamorphic; the first postem- abdomen. bryonic stage is a zoea, with 2 or 3 pairs of maxillipeds The pagurid reproductive system differs not only from present, and a carapace with the posterior margins pro­ other anomurans but from decapods in general in that it duced into 2 lateral spines. The zoeal stage is followed by is located almost exclusively in the abdomen. In the male metamorphosis to a megalopa. In pagurids the zoea is the testes are paired, ovoid structures consisting of long basically symmetrical, but the megalopa (also referred to convoluted tubules, frequently nearly imbedded in as a glaucothoe) may exhibit asymmetry in the uropods tubules of the hepatopancreas. Anteriorly from the and chelipeds. testes, at about the level of the 3rd abdominal somite, trace one of the vas deferens anteriorly to its opening on the coxa of the 5th pereopod. The paired ovaries of the References female lie in approximately the same position; the Alcock, A., 1905. Catalogue of the Indian decapod Crustacea oviducts are distinct and lead from the abdomen, in the collection of the Indian Museum. Part 2, Anomura, through the thorax, to their openings on the coxae of the Fasc. 1. Pagurides. 197 pp. Calcutta: Indian Museum. 3rd pereopods. % Balss, H., 1940-1957. Decapoda. In H. G. Bronn, Klassen The muscular esophagus leads into the large stomach. und Ordnungen des Tierreichs. 5(1) 7: 321—480. Leipzig: The cardiac portion easily is recognized by its baglike Akad. Verl. appearance and membranous dorsal surface. After ex­ Benedict, J. E., 1892. Preliminary descriptions of 37 new amining the remainder of the digestive system, remove species of hermit crabs of the genus Eupagurus in the U.S. the stomach and examine the gastric mill of the cardiac National Museum. Proc. U.S. Natl. Mus., 15: 1-26. stomach. The considerably smaller pyloric stomach is , 1903. Revision of the crustaceans of the genus separated from the former by a cardiopyloric valve. At Lepidopa. Proc. U.S. Natl. Mus., 26: 889-895. the junction of the pyloric stomach and midgut a pair of , 1904. A new genus and two new species of crustaceans small, anteriorly directed ceca usually arise dorsally; of the family Albuneidae from the Pacific Ocean; with re­ ducts of the hepatopancreas enter the midgut at this marks on the probable use of the antennule in Albunea and level too. The midgut is an elongate thin-walled smooth Lepidopa. Proc. U.S. Natl. Mus., 27: 621-625. tube extending almost the full length of the abdomen. X Bliss, D. E., 1968. Transition from water to land in decapod Just anterior to its junction with the hindgut or rectum, a crustaceans. Am. Zool., 8: 355—392. prominent, anteriorly directed cecum arises. It has been 1 Caiman, W. T., 1909. Crustacea. In R. Lankester (ed.), A called the hindgut cecum, but this is a misnomer as it treatise on zoology, pt. 7, Appendiculata, fasc. 3, 346 pp. originates from the midgut. The hindgut terminates in a London: Ray Soc. ventrally directed anus at the terminal end of the telson. Chace, F. A., Jr., 1942. Report on the scientific results of the Excretion in anomurans is via the antennal glands; in Atlantis expeditions to the West Indies, under the joint aus­ pices of the University of Havana and Harvard University. pagurids this usually is supplemented by an unpaired The anomuran Crustacea. I. Galatheidae. Torreia, Havana, abdominal sac or bladder often hidden in the lobes of the no. 11: 1- 106. hepatopancreas. H-'Chace, F. A., Jr., and H. H. Hobbs, Jr., 1969. The freshwater The prominent supraesophageal ganglion can be seen and terrestrial decapod crustaceans of the West Indies with in the midline between the ocular peduncles and above special reference to Dominica. Bull. U.S. Natl. Mus., 292: the epistome. Trace the major nerves radiating from the 1-258. ganglion; that is, the optic, antennal, antennular, and Efford, I. E., 1966. Feeding in the sand crab Emerita analoga. tegumental. Follow one of the esophageal connectives Crustaceana, 10: 167-182. around the esophagus; the swelling is the paraesophageal , 1967. Neoteny in Emerita. Crustaceana, 13: 81— 93. ganglion. The connective terminates in a thoracic Forest, J., and M. de Saint Laurent, 1968. Campagne de la ganglionic mass overlaying the inferior thoracic artery. "Calypso" au large des cotes Atlantiques de lAmerique de Three masses of fused ganglia, separated by constric­ sud (1961—1962). 6. Crustaces Decapodes: Pagurides. Ann. tions, comprise this thoracic mass. The 3rd cluster of Inst. Oceanogr., 45: 47—169. ganglia, which is pierced by the sternal artery, is com­ Gordan, J., 1956. A bibliography of pagurid crabs, exclusive of posed of the ganglia of the 4th and 5th pereopods and 1st Alcock, 1905. Bull. Am. Mus. Nat. Hist, 108: 257-352. ORDER DECAPODA 151

Haig, J., 1956. The Galatheidae (Crustacea Anomura) of the capoda, Paguridea) of northwestern North America. Zool. Allan Hancock Expedition with a review of the Porcellanidae Verhandl, no. 130: 1-396. of the western Atlantic. Allan Hancock Pacific Exped.: Makarov, V. V., 1938 (1962). Fauna of USSR, Crustacea. 10, Rept. no. 8:1-44. Anomura. Zool. Inst. Akad. Nauk SSSR. 278 pp. (Translated ,1960. The Porcellanidae (Crustacea Anomura) of the by F. D. Por, for the National Science Foundation and eastern Pacific. Allan Hancock Pacific Exped., 2A: 1—440. Smithsonian Institution). Hay, W. P., and C. A. Shore, 1918. The decapod crustaceans of Man, J. G., de, 1927. A contribution to the knowledge of 21 Beaufort, N.C., and the surrounding region. Bull. Bur. species of the genus Upogebia Leach. Capita Zool, 2: Fish., 35: 369-475. 1-56. Hazlett, B. A., 1966. Social behavior of the Paguridae and Melin, G., 1939. Paguriden und Galatheiden von Prof. Sixten Diogenidae of Curacao. Stud. Fauna Curaqao Carib. Isl., Bock's Expedition nach den Bonin-Inseln 1914. K. Svenska. 23: 1- 143. Vetensk. Akad. Handl. 18:, 1- 119. )r Holthuis, L. B., 1959. The Crustacea Decapoda of Suriname A Miyake, S., 1978. The crustacean Anomura ofSagami Bay. 200 (Dutch Guiana). Zool. Verhandl. no. 44: 1—296. pp. Tokyo: Biol. Lab., Imperial Household (ed.). i Kensley, B., 1977. The South African Museums Meiring Pike, R. B., 1947. Galathea. Liverpool Mar. Biol Comm., Naude cruises. Part 2. Crustacea, Decapoda, Anomura and Brachyura. Ann. S. Afr. Mus., 72: 161-188. Mem. 34:1-179. X Lewinsohn, Ch., 1969. Die Anomuren des Roten Meeres Pilgrim, R. L. C, 1965. The morphology of Lomis hirta and a (Crustacea Decapoda: Paguridea, Galatheidea, Hippidea). discussion of its systematic position and phylogeny. Aust. J. Zool. Verhandl, no. 104: 1-213. Zool, 13: 545-557. , 1978. Bemerkungen zur Taxonomie von Paguritta Schmitt, W. L., 1921. The marine decapod Crustacea of harmsi (Gordon) (Crustacea Decapoda, Anomura) und California. Univ. Calif. Publ. Zool, 23: 1-470. Beschreibung einer neuen Art der gleichen Gattung aus , 1935. Mud shrimps of the Atlantic coast of North Australien. Zool. Meded., 53: 243-252. America. Smiths. Misc. Coll., 93: 1—21. MacDonald, J. D., R. B. Pike, and D. I. Williamson, 1957. Snodgrass, R. E., 1952. The sand crab Emerita talpoida. Larvae of the British species of Diogenes, Pagurus, Anapa- Smiths. Misc. Coll. 117:1-34. gurus and Lithodes (Crustacea, Decapoda). Proc. Zool. Soc. Vinogradov, L. G., 1950. Opredelitel krevetok, rakov i karbov London, 128: 209-257. Dalnego Vostlka. Izvest. Tikhookean. Nauchno-issledovat. X McLaughlin, P. A., 1974. The hermit crabs (Crustacea De­ Inst. Rybiogo Khozyaistva Okean. 33: 179— 358. ^Williams, A. B., 1965. Marine decapod crustaceans of the Carolinas. Fish. Bull, 65: 1-298.

INFRAORDER BRACHYURA Latreille, 1803

Recent species Approximately 4500. Size range Up to 365 cm (leg span). Carapace Progressively shortened and widened; fused to epistome. Eyes Stalked, compound; sometimes reduced. Antennules Peduncles with 3 segments; flagella often reduced or vestigial. Antennae Peduncles usually with 1 or 2 segments; usually without exopod; flagella short, sometimes absent. Mandibles With or without palp; molar and incisor processes usually not distinct. Maxillulae Usually with endopodal palp. Maxillae Biramous; usually with bilobed endites. Maxillipeds Flagella often reduced or absent; 3rd often with ischium and merus flattened, operculate. Thoracic appendages First chelate; 5th, or 4th and 5th, occasionally subchelate; 5th occasionally paddle-shaped. Abdominal appendages In males 1st and 2nd pairs developed as gonopods, 3rd—5th absent; in females 2nd— 5th usually developed, rarely 1st pair also developed; uropods in both sexes usually absent; some­ times present in Dromioidea. Telson Usually reduced. 152 CLASS MALACOSTRACA

Tagmata Cephalothorax and abdomen. Somites Head with 5+3 thoracic (maxillipeds); thorax with 5; abdomen with 6 or fewer (some frequently coalesced), excluding telson. Sexual characters Gonopores on coxae or sternite of 3rd pereopods in female, on 5th in male; 1st and 2nd pairs of pleopods usually modified as gonopods in male. Sexes Presumably separate. Larval development Metamorphic; zoea —» megalopa. Fossil record L. Jurassic to Recent. Feeding types Various. Habitat Marine, freshwater, and semiterrestrial. Distribution Worldwide.

The infraorder Brachyura is divided into five major sec­ about 1 to 5 mm from the dorsolateral margin on each tions, the Dromiacea, Oxystomata, Oxyrhyncha, Can- side and on the anterior margin just behind the orbits. cridea, and Brachyrhyncha. Although considerable vari­ Starting at the posterior margin, carefully begin to sepa­ ation occurs, the basic brachyuran characters can be rec­ rate the carapace from the underlying hypodermis. In ognized. Examine the external morphology of all the the region of the cervical groove, a pair of prominent available taxa, paying particular attention to the regions muscles, the anterior dorsal pyloric muscles, attached to of the carapace, the structure and position of the anten- the carapace will have to be severed. A few additional nules and antennae, and the ocular peduncles and orbits muscle strands probably also will have to be cut to re­ (see Figures 51 and 52). What differences from the struc­ move the carapace. With the removal of the hypodermis, tures in anomurans do you notice? In the oral region, certain muscles, the stomach, heart, and the gills will be observe the complete closure often obtained by the ex­ apparent. In the dorsal midline anteriorly to either side panded and operculate segments of the 3rd maxillipeds. of the stomach identify the median posterior gastric In contrast to the anomurans, the gonopores of muscles. Posteriorly in the midline observe the large brachyurans are not readily discernible, as they are hid­ heart; a pair of prominent ostia is present on the dorsal den by the tightly folded-under abdomen. The sex of the surface. Two additional pairs can be seen laterally if the animal may be determined by the shape of the abdomen. heart is gently pushed to one side. Three arteries, the In males the abdomen is acutely triangular; in females it unpaired ophthalmic and paired antennal, leave the is broadly triangular to semisubcircular. Care must be heart anteriorly. Follow the ophthalmic artery across the exercised in attempting to sex juveniles or young adults dorsal surface of the stomach; anteriorly it turns ventrally as the abdomen frequently is not appreciably expanded and branches to provide blood to the supraesophageal in immature females. If the abdomen is extended, the ganglion, eyes, and antennules. It will be difficult to pairs of modified pleopods that form the gonopods of the trace the path of the antennal arteries until components males or the posterior 4 pairs of female pleopods can be of the digestive system have been examined and re­ observed. moved. Posterolaterally of the heart, behind the bran­ After completing your general examination of the ex­ chial chambers, observe the flattened pericardial sacs. ternal morphology proceed to an examination of the From the posterior margin of the heart the unpaired major organ systems before studying the mouthparts, as superior abdominal artery turns ventrally and gives off a the removal of these appendages often will cause serious pair of branches that provides blood to the dorsal proxi­ damage to the cephalic region of the animal (see Figure mal muscles of the more posterior pairs of pereopods. It 53). The description of the internal anatomy is based on will be difficult to trace these and the ventral hepatic specimens of the Portunidae; however, the information is arteries until some of the other organs are removed. sufficiently general to be applicable to most of the larger, Laterally the gills may be seen to converge to an apex unspecialized taxa. Begin by carefully removing the dor­ abutting a portion of the endoskeleton. Overlying the sal carapace. This can best be done by cutting, with a gills observe the long epipodite of the 1st maxilliped. pair of sharp scissors, first the posterior margin and then What is the gill formula of your specimen? Remove the

Figure 51 Decapoda, Brachyura: A. Typical raninoidean; B. Typical dromioidean; C. Typical caloppoidean; D. Typical majoidean; E—I, General structures from Portunoidea. E. 3rd maxilliped; F. 1st maxilliped; G. Maxilla; H. Maxillule; L Male gonopod (1st pleopod). 153

Endopod (ischium)

Pereopods Dactyl Propodus Cheliped Carpus Merus

'Anterolateral spines

Palp Merus Merus Ischium Ischium Exopod Exopod

Figure 52 Decapoda, Brachyura: A, B, Portunoidea; C, D, Grapsoidea. A. Typical portunid (dorsal view) showing regions of carapace; B. Female portunid (ventral view, with pereopods removed); C. Typical grapsid (dorsal view) showing regions of carapace; D. Male grapsid (ventral view, with pereopods removed). 155

Anterior Lateral adductor stomach Supraesophageal Antennal Ocular muscle of muscle ganglion gland orbit mandible Antenna \ I Antennule Cheliped

Hepatopancreas Cardiac stomach

. Hepatopancreas

Ophthalmic artery

— Muscles of 5th pereopod

Superior abdominal artery Hindgut cecum Midgut Lateral abdominal artery

Postero Pyloric lateral stomach ostium Hepatic Supraesophageal artery ganglion Sternal Superior artery Optic abdominal nerve artery

Lateral abdominal artery

Antennal gland

Esophageal connective

3rd maxilliped

Postion of cheliped

Hepatopancreas

Figure 53 Decapoda, Brachyura: A. Diagrammatic portunid (dorsal view) with carapace removed to show gills and pericardial sac (left) and pereopodal musculature (right); B. Diagrammatic portunid (lateral view) with appendages, gills, and musculature removed to show major organ systems; C. Zoea; D. Megalopa. 156 CLASS MALACOSTRACA gills and locate the smaller but similarly appearing the posterior thoracic appendages often are present, it epipodites of the 2nd and 3rd maxillipeds. Remove the may be considered a later substage than typically seen in stiff membranous floor of the branchial chamber to ex­ anomuran zoeae. The brachyuran megalopal appendages pose the muscles of the cheliped and first 3 pereopods; resemble those of the adult; however, the abdomen is removal of the pericardial sac and underlying membrane large and usually carried in an extended position; the 5 will expose the muscles of the 5th pereopod. pairs of pleopods are used in swimming. In cases of ab­ Lateral to the stomach the mass of tissue filling much breviated development, the megalopal stage may be of the cephalothorax represents lobes of the hepatopan- suppressed. Metamorphosis apparently is suppressed in creas or 1st pair of ceca. If this is removed carefully, 2 most, if not all, freshwater and terrestrial species. mandibular muscles can be distinguished: the large lat­ eral adductor muscle and the smaller external abductor muscle. The latter can be traced from the mandible to a References point on the endoskeleton just anterior to the apex of the Alcock, A., 1901. Catalogue of the Indian decapod Crustacea gills. in the collection of the Indian Museum. Part 1. Brachyura, Ovaries or testes lie slightly posterior to the pyloric Fasc. 1. Introduction and Dromides or Dromiacea (Brachy­ stomach on top of or among the lobes of the hepatopan- ura Primigenia). 80 pp. Calcutta: Indian Museum. creas. The difference in texture of the gonadal tissue will Balss, H., 1940-1957. Decapoda. In H. G. Bronn, Klassen aid in distinguishing it from the tissue of the hepatopan- und Ordnungen des Tierreichs. 5(1) 7: 321—480. Leipzig: creas. The ovaries actually are connected by a cross- Akad. Verl. bridge just behind the pyloric stomach. A short distance ^Bennett, E. W., 1964. The marine fauna of New Zealand: Crus­ behind the bridge are the paired oviducts, but these tacea Brachyura. Mem. IV.Z. Oceanogr. Inst., no. 22: usually are concealed in the ovaries. Near the median 7- 120. A border of each duct is an oval-shaped seminal receptacle Bliss, D. E., 1968. Transition from water to land in decapod that extends ventrally almost to the sternal floor; ducts crustaceans. Am. Zool., 8: 355—392. lead from the receptacles to the gonopores on the 6th A Bliss, D. E., and L. H. Mantel, 1968. Adaptations of Crustacea somite. The testes similarly are connected by a bridge to land. Am. Zool, 8: 673-685. behind the pyloric stomach. The vas deferens leaving the Broekhuysen, G. J., Jr., 1936. On growth and distribution of testes are coiled, and in mature individuals can be sepa­ Carcinidesmaenas (L.). Arch. Neerland. Zool., 2:257— 400. rated into 4 distinct regions related to spermatophore y Bourne, G. C, 1922. The Raninidae; a study in carincology. /. production. Linn. Soc. London (Zool.), 35: 25—79. After tracing the midgut from the pyloric stomach to Caiman, W. T., 1909. Crustacea. In R. Lankester (ed.), A the abdomen, carefully remove the stomach by cutting it treatise on zoology, pt. 7, Appendiculata, fasc. 3, 346 pp. London: Ray Soc. at its junctions with the esophagus and midgut. Posterior and lateral of the stomach a 2nd pair of thin, elongate Chace, F. A., Jr., 1940. Reports on the scientific results of the ceca can be observed. A 3rd, unpaired cecum, arising "Atlantis" expeditions to the West Indies, under the joint auspices of the University of Havana and Harvard Univer­ from the hindgut, is located behind or posterior to the sity. The brachyuran crabs. Torreia Havana, 4: 3— 67. heart. Examine the gastric mill of the cardiac stomach. , 1951. The oceanic crabs of the genera Planes and How does this grinding apparatus compare with those of Pachygrapsus. Proc. U.S. Natl. Mus., 101: 65- 103. the other decapods studied? X Chace, F. A., Jr., and H. H. Hobbs, Jr., 1969. The freshwater The general plan of the nervous system is similar to and terrestrial decapod crustaceans of the West Indies with that of the crayfish but may not be observed as easily. special reference to Dominica. Bull. U.S. Natl. Mus., 292: The major difference between the nervous systems of 1-258. the two is that in the brachyuran the ventral ganglia are X Christensen, A. M., and J. J. McDermott, 1958. Life history fused into a single mass. The supraesophageal ganglion and biology of the oyster crab Pinnotheres ostreum. Biol. lies immediately behind the rostrum beneath the trans­ Bull, 114: 146-179. verse apodeme between the eyes. Trace the major \ Christiansen, M. E., 1969. Crustacea Decapoda Brachyura. nerves radiating from the ganglion; these are the same as Mar. Invert. Scand., 2: 1— 143. have been seen in other decapods. The esophageal con­ * Crane, J., 1937. The Templeton Crocker Expedition. III. nectives are considerably longer than in other decapods Brachygnathous crabs from the Gulf of California and the because the fused ventral ganglia are situated between west coast of lower California. Zoologica, 22: 47— 78. the 4th and 5th thoracic somites. Now remove and , 1975. Fiddler crabs of the world. Ocypodidae: Genus examine the mouthparts. What structural similarities Uca. 736 pp. Princeton, N.J.: Princeton University Press. and differences do you find between the mouthparts of Cronin, L. E., W. A. van Engel, D. G. Cargo, and F. J. Woj- brachyurans and those of other decapods? cik, 1957. A partial bibliography of the genus Callinectes. Larval development is metamorphic. The first post- Spec. Sci. Kept., Virginia Fish. Lab., no. 8, 21 pp. embryonic stage is a zoea; however, since rudiments of \ Felder, D. L., 1973. An annotated key to crabs and lobsters ORDER DECAPODA 157

(Decapoda, Reptantia) from the coastal waters of the north­ naturels chez les crustaces decapodes brachyoures. VII. Les western Gulf of Mexico. La. St. Univ. Publ. no. LSU SG- Goneplacidae (suite et fin). Bull. Mus. Natl. Hist. Nat., 41: 73-02: 1-103. 688-724. \ Flipse, H. J., 1930. Die Decapoda Brachyura der Siboga- , 1971. Recherches preliminaires sur les groupements Expedition. Part 6. Oxyrrhyncha: Parthenopidae. Siboga- naturels chez les crustaces decapodes brachyoures. VIII. Exped. Mongr. 39c2: 1—96. Synthese et Bibliographic Bull. Mus. Natl. Hist. Nat., 42: x Garth, J. S., 1940. Some new species of brachyuran crabs from 1063- 1090. Mexico and the Central and South American mainland. Hagen, H. O. von, 1975. Klassifikation und phylogenetische Allan Hancock Pacific Exped., 5: 53-127. Einordnung der Lautausserungen von Ocypodiden und v , 1946. Littoral brachyuran fauna of the Galapagos Ar­ Grapsiden (Crustacea, Brachyura). Z. Zool. Syst. Evol., 13: chipelago. Allan Hancock Pacific Exped., 5: 341- 601. 300-316. , 1958. Brachyura of the Pacific coast of America, Holthuis, L. B., 1959. The Crustacea Decapoda of Suriname Oxyrhyncha. Allan Hancock Pacific Exped., 21, pts. 1, 2: (Dutch Guiana). Zool. Verhandl, no. 44:1-296. 1-499, 500-854. Kensley, B., 1977. The South African Museums Meiring , 1960. Distribution and affinities of the brachyuran Naude cruises. Part 2. Crustacea, Decapoda, Anomura and Crustacea. In The biogeography of Baja California and adja­ Brachyura. Ann. S. Afr. Mus., 72: 161-188. cent seas (Symposium). Syst. Zool., 9: 105—123. Lewinsohn, Ch., 1977. Die Dromiidae des Roten Meeres Garth, J. S., and J. Haig, 1971. Decapod Crustacea (Anomura (Crustacea Decapoda, Brachyura). Zool. Verhandl., no. 151: and Brachyura) of the Peru-Chile Trench. Anton Bruun 1-41. Rept., no. 6:1-20. , 1977. Die Ocypodidae des Roten Meeres (Crustacea Glaessner, M. F., 1969. Decapoda. In R. C. Moore (ed.), Decapoda, Brachyura). Zool. Verhandl., no. 152: 45—84. Treatise on invertebrate paleontology, Part R Arthropoda. 2: R400— R651. Lawrence, Kans.: Geol. Soc. America and Man, J. D., de, 1891. Decapoden des Indischen Archipels. Univ. Kansas. Zool. Ergebn., 2: 265-527. Gordon, I., 1963. On the relationship of Dromiacea, Tymolinae , 1902. ErgebnisseeinerzoologischenForschungsreisein and Rainindae to the Brachyura. In H. B. Whittington and den Molukken und Borneo, im Auftrage der Senckenberg. W. D. I. Rolfe (eds.), Phylogeny and evolution of Crustacea, naturforsch. Gesellschaft ausgefuhrt von Dr. Willy Kiikent- pp. 51-57. Spec. Publ. Mus. Comp. Zool. Harvard. Cam­ hal. Abh. Senckenberg. Naturf. Ges., 25: 467-929. bridge, Mass.: Harvard University Press. Monod, Th., 1956. Hippidea et Brachyura ouest-Africains. Gore, R. H., and L. E. Scotto, 1979. Crabs of the family Par­ Mem. Inst. Fr. Afr. Noire, no. 45:1— 674. thenopidae (Crustacea Brachyura: Oxyrhyncha) with notes Nation, J. D., 1975. The genus Cancer (Crustacea: Brachyura): on specimens from the Indian River region of Florida. systematics, biogeography and fossil record. Nat. Hist. Mus. Mem. Hourglass Cruises 3: 1— 98. L.A. Count. Sci. Bull, no. 23: 1- 104. Ortmann, A., 1892. Die Decapoden-Krebse des Strassburger Griffin, D. J. G., 1976. Spider crabs of the family Majidae Museum. Zool. Jahrb. Syst., 6: 532-588. (Crustacea: Brachyura) from the Philippine Islands. /. Nat. Hist., 10: 179-222. , 1893. Die Decapoden-Krebse des Strassburger Museum. Pts. 6—8 (Brachyura). Zool. Jahrb. Syst., 7: Griffin, D. J. G., and H. A. Tranter, 1974. Spider crabs of the 23-88, 411-495, 683-772. family Majidae (Crustacea: Decapoda: Brachyura) from the Red Sea. Israel J. Zool, 23: 162-198. Pequegnat, W. E., 1970. Deep water brachyuran crabs. In W. Guinot, D., 1967. Recherches preliminaires sur les groupe- E. Pequegnat, and F. A. Chace, Jr. (eds.), Contributions on ments naturels chez les crustaces decapodes brachyoures. I. the biology of the Gulf of Mexico. Texas A & M Univ. Les affinites des genres Aethra, Osachila, Hepatus, Oceanogr. Stud. 1:171-204. Houston, Texas: Gulf Publ. Co. Hepatella etActaeomorpha. Bull. Mus. Natl. Hist. Nat. (2)38: Pequegnat, W. E., L. H. Pequegnat, R. W. Firth, Jr., B. M. 744-762,828-845. James, and T. W. Roberts, 1971. Gulf of Mexico deep-sea , 1968a. Recherches preliminaires sur les groupements fauna. Decapoda and Euphausiacea. 12 pp. Serial Atlas of naturels chez les crustaces decapodes brachyoures. IV. Ob­ the Marine environment, Folio 20. Am. Geogr. Soc. servations sur quelques genres de Xanthidae. Bull. Mus. Rathbun, M. J., 1893. Scientific results of explorations by the Natl. Hist. Nat. 39: 695-727. U.S. Fish Commission steamer "Albatross." XXIV. Descrip­ , 1968b. Recherches preliminaires sur les groupements tions of new genera and species of crabs from the west coast naturels chez les crustaces decapodes brachyoures. VI. Les of North America and the Sandwich Islands. Proc. U.S. Carpilinae. Bull. Mus. Natl. Hist. Nat. 40: 320-334. Natl. Mus., 16: 223-260. , 1969a. Recherches preliminaires sur les groupements , 1905. Fauna of New England. 5. Lists of the Crus­ naturels chez les crustaces decapodes brachyoures. VII. Les tacea. Occ. Pap. Boston Soc. Nat. Hist. Goneplacidae. Bull. Mus. Natl. Hist. Nat., 41: 241-265. , 1925. The spider crabs of America. U.S. Natl. Mus. , 1969b. Recherches preliminaires sur les groupements Bull. 129:1-613. naturels chez les crustaces decapodes brachyoures. VII. Les , 1930. The cancroid crabs of America of the families Goneplacidae (suite). Bull. Mus. Natl. Hist. Nat.,41:507- 528. Euryalidae, Portunidae, Atelecyclidae, Cancridae and , 1969c. Recherches preliminaires sur les groupements Xanthidae. U.S. Natl. Mus. Bull. 152: 1-609. 158 CLASS MALACOSTRACA

Rathbun, M. J., 1937. The oxystomatous and allied crabs of Tesch, J. J., 1918a. The Decapoda Brachyura of the Siboga- America. 17. S. Natl. Mus. Bull. 166:1-278. Expedition. I. Hymenosomidae, Retroplumidae, Ocypodi- Sakai, T., 1965. The crabs of Sagami Bay. 330 pp. Tokyo: dae, Grapsidae, and Gecarcinidae. Siboga-Exped., Mongr. Maruzen Co. Ltd. 39c: 1-148. , 1975. Crabs of Japan and the adjacent seas, vols. 1-3. , 1918b. The Decapoda Brachyura of the Siboga Expe­ 773 pp., 461 pp., 251 pis. Tokyo: Kodansha Ltd. dition. II. Goneplacidae and Pinnotheridae. Siboga- Schmitt, W. L., 1921. The marine decapod Crustacea of Exped., Mongr. 39c; 149-295. California. U. Calif. Publ. Zool. 23: 1-470. Tiirkay, M., 1974. Die Gecarcinidae Asiens und Ozeaniens Serene, R., 1973. A key for the separation of the Indo-Pacific (Crustacea: Decapoda). Senckenberg. Biol. 55: 223— 259. species of Uca Leach 1914. Spl. Publ. Mar. Biol. Assoc. Warner, G. F., 1967. Life history of the mangrove tree crab India, pp. 349-361. Aratus pisoni. J. Zool. London, 153: 321—335. Serene, R., and P. Lohavanijaya, 1973. The Brachyura (Crus­ , 1977. The biology of crabs. 202 pp. New York: Van tacea: Decapoda) collected by the Naga Expedition, includ­ Nostrand Reinhold Co. ing a review of the Homolidae. Naga Kept., 4: 1—146. Williams, A. B., 1965. Marine decapod crustaceans of the Stephensen, K., 1945. The Brachyura of the Iranian Gulf. Carolinas. Fish. Bull, 65: 1-298. Danish Sci. Invest. Iran, Pt. 4: 57— 237. , 1976. Distinction between a Gulf of Mexico and a Stimpson, W., 1907. Report on the Crustacea collected by the Carolinian Atlantic species of the swimming crab Ovalipes North Pacific Exploring Expedition 1853—1856. Smiths. (Decapoda: Portunidae). Proc. Biol. Soc. Wash., 89: 205— Misc. Coll., 49: 1-240. 214. GENERAL CRUSTACEAN REFERENCES

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GLOSSARY OF MORPHOLOGICAL TERMS

Abdomen. Trunk somites (tagma) between thorax and telson; Antennal muscle scar. Impression of antennal muscle on inner somites with or without limbs; syn., pleon. surface of valve, situated in front of adductor muscle scar, Abdominal process. Fingerlike projection^) on dorsal surface generally above (in some podocopans behind) mandibular of abdomen (Cladocera). muscle scar (Ostracoda). Abdominal somite. Any single division of body between thorax Antennal region. See decapod carapace, region a. and telson; syn., pleomere, pleonite. Antennal scale. See scaphocerite. Abreptor. Postabdomen, bent forward from junction with Antennal spine. See decapod carapace, spine a. body, terminating in 2 claws (Cladocera). Antennular fossette. Depression, pit or socket containing basal portion of the antennule. Acanthopod. Appendage (cirrus) with short row of strong sharp spines distally at each articulation of greater curvature Antennular scale. See stylocerite. and few or no spines along lesser curvature (Cirripedia). Antennule. One of pair of appendages of first cephalic somite; Acron. Anteriormost part of body carrying eyes; not consid­ syn., 1st antenna. ered true cephalic somite; syn., ophthalmic somite, preseg- Anterolateral region. See decapod carapace, region b. mental region. Aperture. Posteroventral opening into mantle cavity (Cir­ Adductor muscle. (1) Muscle attached to carapace for pulling it ripedia); cf. orifice. to body or connecting halves of bivalve carapace (e.g., Con- chostraca, Ostracoda, Leptostraca); or (2) Muscle attached to Apex. Upper angle of scutum or tergum (thoracic Cirripedia). scutum for closing aperture (thoracic Cirripedia). Apodeme. Infold of exoskeleton for attachment of muscles. Adductor muscle scar. Impression of adductor muscle on inner Appendix interna. Median process of pleopodal endopod unit­ surface of each valve (Ostracoda); see muscle scar. ing members of each pair; syn., stylamblys. Adductor pit. Depression on inner surface of scutum between Appendix masculina. Complex median process of 2nd pleo­ adductor ridge and occludent margin for attachment of ad­ podal endopod of male Caridea and some other Eucarida ductor muscle (thoracic Cirripedia). used in copulation or spermatophore transfer. Adductor ridge. Linear elevation on inner surface bounding Areola. Area between branchiocardiac grooves and posterior adductor pit on tergal side in sessile barnacles. to cervical groove on dorsal surface of carapace (Astacidea and Austroastacidea). Aesthetasc. Sensory seta covered by delicate cuticle, often projecting from antenna or antennule; syn., olfactory hair, Arthrobranch. Gill attached to articular membrane between ethetasc, esthete.. appendage and body (Decapoda); syn., arthrobranchiata. Afferent channel. Opening through which water passes to gills. Arthrobranchiata. See arthrobranch. Ala (pi. aloe). One of pair of posteriorly directed cephalic- Arthrophragm. See endophragm. shield extensions (Branchiura and some parasitic Copepoda); Article. Subdivision of antennal or antennular flagella or ap­ triangular lateral part of compartmental plate, with or with­ pendage (cf. segment). out radii, overlapped by adjacent compartmental plate Articular furrow. Groove near tergal margin of scutum or (balanomorph Cirripedia). scutal margin of tergum forming part of articulation between Aliform apophyses. Incurved anterior and posterior ex­ the two valves (balanomorph Cirripedia). tremities of growth lines (Conchostraca). Articular ridge. Linear elevation on scutum or tergum border­ Ambulatory leg. Seepereopod. ing articular furrow and with it forming articulation between the two valves (balanomorph Cirripedia). Anal spines. Single row of spines on either side of postabdo­ men (Cladocera) or prominent telsonal spines (Conchos­ Atrium oris. Preoral cavity, bounded ventrally by posteriorly traca). directed labrum, dorsally by ventral surface of cephalon just behind mouth, and laterally by paragnaths and mandibles. Annulus ventralis. Seminal receptacle of female crayfish. Attractor epimeralis muscle. Prominent muscle in many de­ Antenna. One of pair of appendages of second cephalic somite; capods, inserting along line of branchiocardiac groove. syn., 2nd antenna. Basal margin. Lower edge of scutum or tergum or other plate Antennal carina. See decapod carapace, carina a. (thoracic Cirripedia). Antennal gland. One of pair of complex excretory glands in Basicarinal angle. Intersection of basal and carinal margins of many malacostracans with duct opening on antenna; syn., tergum (thoracic Cirripedia). green gland. Basicerite. Second segment of antennal peduncle, bearing Antennal groove. See decapod carapace, groove a. scaphocerite (Caridea). 162 GLOSSARY OF MORPHOLOGICAL TERMS

Basiocchident angle. Intersection of basal and occludent mar­ anterior prominence in complex tooth and socket hingement gins of scutum (thoracic Cirripedia). (Ostracoda). Basiophthalmite. Proximal segment of eyestalk, articulating Carapace. Cuticular, usually calcified, structure arising from with distal segment (podophthalmite) bearing corneal sur­ posterior margin of cephalon, extending anteriorly and pos­ face of eye. teriorly, often covering head and thorax (cf. Eucarida); also Basipod(ite). See basis. fold of integument extending from maxillary segment, form­ Basis (pi. bases). Segment of protopod adjoining coxa and car­ ing bivalved shell of cyprid larvae and of ascothoracicans and rying exopod and endopod distally; also basal calcareous or mantle of other Cirripedia; mantle usually with calcified membranous plate furnishing anchorage to substrate in ses­ plates in thoracicans. sile cirripeds. Carapace angles. Intersection of straight dorsal margin by an­ Basiscutal angle. Intersection of basal and scutal margins of terior rib (a) and posterior rib (/3) (Conchostraca). tergum (thoracic Cirripedia). Carapace carina. Narrow ridge on surface of carapace (cf. dec­ Basitergal angle. Intersection of basal and tergal margins of apod carapace and stomatopod integumental ornamenta­ scutum (thoracic Cirripedia). tion). Biformes. Carapaces reflecting sexual dimorphism (e.g., Con- Carapace costae. Closely spaced radial ridges, grading from chostraca), marked by differing valve proportions for each fine to coarse, not crossing umbo (Conchostraca); syn., radial sex of the same species. lirae. Biramous. Having two branches. Crustacean appendage with Carapace costeUae. Fine radial ridges extending from ventral two rami; also antennule or antenna with two flagellar ele­ margin across umbo (Conchostraca). ments. Carapace groove. Furrow on surface of carapace (cf. decapod Biserial branch. Primary branch of dendrobranchiate gill sub­ carapace and stomatopod integumental ornamentation). divided into two rows or series. Carapace growth line. Peripheral margin of successive mem­ Bopyridum. Postlarva of epicaridean isopod that is attached to branes added to shell during each molt (Conchostraca). permanent host. Carapace lirae. Linear concentric ridges parallel to and be­ Brain. See supraesophageal ganglion. tween growth lines (Conchostraca). Bronchia (pi. branchiae). Thin-walled, fingerlike or leaflike Carapace region. Differentiated portion of carapace surface structure extending outward from appendage or secondarily (cf. decapod carapace). from side of body, functioning in respiration; syn., gill. Carapace spine. Sharp projection from carapace (cf. decapod Branchial carina. See decapod carapace, carina b. carapace). Branchial chamber. Area between body and carapace enclos­ Carapace tooth. Generally blunt projection of carapace, often ing branchiae; syn. gill chamber. broader than spine (cf. decapod carapace and stomatopod integumental ornamentation). Branchial glands. Masses of connective-tissue cells without Cardiac notch or incision. Indentation on posterior margin of ducts surrounding venous channels in branchiae. carapace. Branchial region. See decapod carapace, region c. Cardiac region. See decapod carapace, region d. Branchiocardiac carina. See decapod carapace, carina c. Cardiac tooth. See decapod carapace, teeth a. Branchiocardiac groove. See decapod carapace, groove b. Cardo. Basal segment of maxillule articulating with cephalon. Branchiostegal area. Part of carapace extending laterally and Caridean lobe. External rounded projection on basal part of ventrally over branchiae. exopod of 1st maxilliped (Caridea). Branchiostegal spine. See decapod carapace, spine b. Caridoid fades. Basic group of characters distinguishing eu- Branchiostegite. Expanded dorsal and lateral branchial region malacostracan crustaceans: enclosure of thorax by carapace, of carapace. movable stalked eyes, biramous antennules, antennae with Buccal cavity. Area of cephalon containing mouthparts; scaphocerites, thoracopods with natatory exopods, ventrally bounded by epistome anteriorly and free margins of carapace flexed abdomen, and tailfan. laterally (Malacostraca). Carina. Any keellike structure; any well-defined projecting Buccal frame. Structural region of cephalon enclosing mouth- ridge on outer surface of carapace (podocopan Ostracoda); parts (Brachyura). unpaired posterodorsal plate of thoracic Cirripedia (in Buccal groove. See decapod carapace, groove c. lepadomorphs 1 of up to 4 unpaired plates of capitulum; in Bullate. Inflated, blisterlike. verrucomorphs compartmental plate between rostrum and Calceolus (pi. calceoli). Complex sensory filaments on anten- fixed tergum; in balanomorphs compartmental plate, with nules (Amphipoda). alae on each side, opposite rostrum). Calyptopis stage. Third larval stage, characterized by differ­ Carinal lotus. See latus. entiation of abdomen and appearance of compound eyes Carinal margin. Edge of any plate adjacent to carina; occlud­ (Euphausiacea); see also zoea. ing with carinal margin of opposed tergum (thoracic Cir­ Capitulum. Portion of carapace enclosing body, commonly ripedia). protected by calcareous plates (lepadomorph cirripeds); or Carinate. Valve bearing rib(s) (Conchostraca). GLOSSARY OF MORPHOLOGICAL TERMS 163

Carinolateral. One of pair of compartmental plates typically Chela (pi. chelae). Distal part of appendage; pincerlike, with overlapping carina on each side, with radius on carinal side opposed movable and immovable fingers; occasionally both and ala on lateral side, sometimes absent; homologous with fingers movable. lepadomorph carinal latus (balanomorph Cirripedia). Chelate. Pincerlike. Carpocerite. Distal (5th) segment of antennal peduncle. Cheliped. Any thoracopod bearing chela. Carpopod(ite). See carpus. Cincinnuhts. See retinaculum. Carpus. Antepenultimate segment of thoracopod or pereopod; Cirrus (pi. cirri). Thoracic, usually biramous, multiarticulate syn., carpopod. appendage generally flattened laterally and curled anteriorly, Caudal appendage. One of terminal, multiarticulate or sim­ with food-gathering function; anterior and posterior margins ple, uniramous paired appendages homologous with caudal designated lesser and greater curvature respectively (Cir­ furca (Cirripedia). ripedia, except Ascothoracica). Caudal fan. Powerful swimming structure formed of laterally Clasper. Appendage, including antenna, modified for holding expanded uropods and telson; syn. tailfan. female during copulation; or an organ for fixation in para­ Caudal furca. Paired caudal rami of terminal abdominal seg­ sites. ment or telson; syn., furca. Clypeus. Part of cephalon carrying labrum; plate anterome- Caudal process. Posterior projection of valve border generally dially on head formed by fusion of basal segments of anten­ above midheight, or posteroventral, directed upward (Os- nae (Anostraca). tracoda). Colleteric gland. Single or paired gland in female or hermaph­ Caudal ramus. One of paired appendages constituting caudal rodite producing viscid material for binding eggs together furca, usually rodlike or bladelike, sometimes filamentous (Rhizocephala). and multiarticular; syn., caudal filament, caudal style, cer- Comb collar. Retractable membrane supporting row of uni­ cus, cercopod, furcal ramus, stylet. form setae, at superior angle of aperture (Acrothoracica). Caudal siphon. Posteroventral opening in valve border; some­ Compartmental plate. One of several rigidly articulated plates times produced as tubular structure (Ostracoda). forming wall in sessile cirripeds. Caudal style. See caudal ramus. Compound eyes. Paired array of contiguous ommatidia having Cement gland. Special concentration of cells in dermal cover common optic nerve trunks. that secretes calcareous substances of valves (Cirripedia). Compound rostrum. Compartmental plate formed by fusion of Cephalic constriction. Constriction delimiting anterior anten- rostrolaterals with rostrum or of rostrolaterals, with rostrum nulary part of head from posterior part (Mystacocarida). missing (balanomorph Cirripedia). Cephalic flexure. Forward, or sometimes upward, deflection of Copepodid. Postnaupliar developmental stage (Copepoda). anterior sterna (Decapoda). Cor frontale. Special pulsating structure or accessory heart Cephalic shield. Chitinous or more or less calcified covering of formed from enlargement of blood vessel; contraction caused the head region, formed of fused tergites of cephalic somites, by outer tangential muscles or internal muscles derived from commonly having pleura. muscles having other functions. Cephalic somite. One unit of cephalon bearing distinctive pair Cormopod(ite). See thoracopod. of appendages; syn., cephalomere. Cornea. Transparent cuticle covering ommatidia of compound Cephahmere. See cephalic somite. eye. Cephalon. Anteriormost tagma, bearing eyes, mouth, two Corpus mandibulae. See mandible body. pairs of antennae, and three pairs of mouthparts; syn., head. Coxa. Segment of appendage adjoining sternite, except in forms having precoxa; syn., coxopodite. Cephalosome. Head region with one or two fused thoracic so­ mites (Copepoda). Coxal endite. Lobe produced from inner margin of coxa. Cephalothorax. Anterior part of body composed of fused Coxal exite. Lobe produced from outer margin of coxa; syn., cephalic and thoracic somites; latter with appendages mod­ coxepipod. ified as mouthparts, sometimes also with relatively unmod­ Coxal plate. Lateral expansion of coxa broadly joined to lateral ified appendages. margin of tergite. Cercopod. See caudal ramus. Coxepipodfite). See coxal exite. Cercus (pi. cerci). See caudal ramus. Crista dentata. Toothed crest on ischium of 3rd maxilliped (Decapoda). Cervical furrow. See decapod carapace, groove d. Cryptoniscus. Planktonic larval stage of epicaridean isopod Cervical groove. See decapod carapace or stomatopod in- with pereopods modified as holdfasts. Stage at which larva tegumental ornamentation. seeks permanent host. Cervical notch or incision. Strong indentation of carapace at Ctenopod. Appendage (cirrus) usually with long paired setae level of cervical groove. on segments of lesser curvature and few setae distally on Cervical sinus. Rounded to angular indentation anteriorly each articulation of greater curvature. along dorsal edge of carapace (Cladocera). Cycladiformes. Carapace with dorsal margin of valve forming Cervical suture. See cervical groove. obtuse angle with posterior margin (Conchostraca). 164 GLOSSARY OF MORPHOLOGICAL TERMS

Cyclops stage. Post-metanaupliar stage (Copepoda). (j) rostral. Continuous with lateral margin of rostrum. Cyprid. Bivalve larval stage (Cirripedia). (k) subhepatic. Extending posteriorly from branchiostegal Cyrtopia. Formerly considered 5th larval stage with antennae spine. no longer used in locomotion; currently included in furcilia (1) submedian. On either side of, and parallel to, postros­ stage (Euphausiacea). tral carina, sometimes joining rostral carina. Dacty1(us). Ultimate segment of thoracopod; syn., dactylo- (m) supraorbital. See decapod carapace, carina d. podite. Grooves: Dactylopodite. See dactyl. (a) antennal. Extending posteriorly from vicinity of anten­ Decapod carapace. Weakly or strongly calcified integuments nal spine. covering cephalothorax; variously subdivided marked, and (b) branchiocardiac. Oblique groove approximately in mid­ armed; named parts: dle of posterior half on each side of carapace, separating Regions: branchial and cardiac regions and reaching dorsomedian (a) antennal. Anterior marginal part bordering orbital re­ part well posterior to cervical or postcervical grooves; gion laterally, adjoining hepatic, pterygostomial, and oc­ sometimes longitudinal, connecting cervical and post- casionally also frontal regions. cervical grooves or extending posteriorly from subme­ (b) anterolateral. Lateral part bordering subhepatic or dian point on postcervical groove. hepatic regions. (c) buccal. Transverse groove crossing mandibular eleva­ (c) branchial. Lateral part posterior to ptergostomial re­ tion behind antennal spine, connecting gastroorbital and gion, overlying branchiae; epibranchial, mesobranchial, antennal grooves (Nephropidae). and metabranchial lobes or areas sometimes distin­ (d) cervical. Transverse groove medially between gastric guished. and cardiac regions, curving toward antennal spine; (d) cardiac. Median part posterior to cervical groove be­ syn., cervical suture, cervical furrow. tween urogastric and intestinal regions. (e) dorsomedian. Longitudinal groove extending from tip (e) frontal. Anteromedian part including rostrum and re­ of rostrum to posterior carapace margin dorsomedially gion behind it. (Nephropidae). (f) gastric. Median part anterior to cervical groove and (f) gastroorbital. Short, longitudinal groove branching posterior to frontal region; sometimes epigastric, from cervical groove at level of orbit and directed to­ mesogastric, metagastric, protogastric, and urogastric ward it. (genital) areas distinguished. (g) hepatic. Short, longitudinal groove connecting cervical (g) hepatic. Part adjoining antennal, cardiac, and pterygo­ with postcervical and branchiocardiac grooves, more or stomial regions. less continuous with antennal groove. (h) intestinal. Short transverse part posterior to cardiac re­ (h) inferior. Extending from junction of hepatic and cervi­ gion; sometimes referred to as posterior cardiac lobe. cal grooves toward lateral margin, more or less continu­ (i) jugal. See pterygostomial region. ous with cervical groove. (j) orbital. Part posterior to eyes bordered by frontal and (i) intercervical. Oblique groove connecting postcervical antennal regions. and cervical grooves (Nephropidae). (k) pterygostomial. Anterolateral part on ventral surface (j) intestinal. Short, transverse groove in median part of located on opposite sides of buccal cavity; syn., jugal posterior carapace, interrupted by intestinal tubercle region, pterygostome. (Nephropidae). (1) subhepatic. Part on ventral surface below hepatic re­ (k) marginal. Close to, and parallel with, posterior margin. gion, bounded by pterygostomial and suborbital re­ (1) parabranchial. Groove below, behind and almost paral­ gions. lel with branchiocardiac and postcervical grooves, join­ (m) suborbital. Part on anteroventral surface beneath orbit. ing latter in lower part (Nephropidae). Carinae: (m) postcephalic. One of three transverse furrows on (a) antennal. Extending posteriorly from antennal spine. carapace of many fossils. (b) branchial. Extending posteriorly from orbit over bran­ (n) postcervical. Posterior to, and parallel with, cervical chial region. groove, bisecting cardiac region. (c) branchiocardiac. Dividing branchial from cardiac re­ (o) sellar. Short transverse groove dorsally anterior to cer­ gion. vical groove (Nephropidae). (d) gastroorbital. Extending posteriorly from supraorbital (p) submedian. Longitudinal groove in submedian dorsal spine; syn., supraorbital. part, contiguous with postrostral carina. (e) lateral. On lateral margin of carapace. (q) urogastric. Short transverse groove in median or sub- median region posterior to postcervical groove, some­ (f) orbital. On margin of orbit. times joining upper part of postcervical groove (Neph­ (g) posterior. Transverse ridge anterior to marginal groove. ropidae). (h) postorbital. Slightly posterior and parallel to orbital margin. Spines: (i) postrostral. Posterior to rostrum along dorsal midline. (a) antennal. On anterior margin slightly below orbit. GLOSSARY OF MORPHOLOGICAL TERMS 165

(b) branchiostegal. On or close to anterior margin medially peripheral part of inner lamina in contact with, or separated between antennal and pterygostomial spines. by vestibule from, outer lamina, generally narrow, some­ (c) hepatic. Below and posterior to lower branch of cervical times extensive (Ostracoda). groove. Ecdysis. Act of molting the integument. (d) infraorbital. On lower angle of orbit. Efferent channels. Passageways through which water moves (e) postorbital. At moderate distance posterior to middle of away from gills and out of branchial region. orbit. Endite. Inwardly (medially) directed lobe of precoxa, coxa, (f) postrostral. Immediately posterior to rostrum. basis, or ischium. (g) pterygostomial. On anterolateral angle. Endognath. Endopod of maxilliped. (h) suborbital. Slightly below and posterior to middle of Endophragm. Septum formed by cephalic and thoracic apodemes; syn., arthrophragm. orbit. (i) supraorbital. At moderate distance obliquely behind Endophragmal skeleton. Complex internal structure formed and above orbit; sometimes on postorbital carina. by fusion of apodemes, providing framework for muscle at­ tachment. Teeth: Endophurite. Lateral apodeme of endoskeleton (Decapoda). (a) cardiac. On midline of carapace just posterior to cervi­ Endopod(ite). Inner ramus of biramous appendage. cal groove. Endosternite. Mesodermal tendonous plate below anterior (b) gastric. On midline of carapace immediately anterior to part of alimentary canal (Notostraca); also firm calcareous cervical groove. plate between nerve cord and alimentary canal anteriorly in (c) lateral. On lateral margin of carapace; anterolateral, thorax of some decapods. mediolateral, and posterolateral teeth distinguished. Endostome. Palatelike part of buccal frame in some (d) orbital. On orbital margin. brachyuran decapods; syn., palate. (e) posterior. On midline of carapace between posterior Ephippium (pi. ephippia). Semielliptical modification of cuti­ margin and marginal groove. cle in dorsal region of carapace valves forming encasement (f) pregastric. On midline of carapace between gastric for eggs, capable of withstanding dessication after being shed tooth and rostrum. (Cladocera). (g) rostral. On rostrum, either single or multiple; upper, Epibranchial lobe. See decapod carapace, region c. lower, and lateral teeth distinguished. Epicaridum. First larval stage of epicaridean isopod; syn., mi- Deflected front. Broadly downturned front margin of carapace cronicus. in some decapods. Epigastric area. See decapod carapace, region/. Dendrobranch(ia). Type of gill with lamellae divided into ar­ Epimeral fold. Steep fold of endopleurites connected with borescent bundles. branchiostegite to form branchial chamber in some dec­ Depressor muscle. Muscle inserted at basicarinal angle of ter- apods. gum (balanomorph Cirripedia). Epimere. Each lateral part of integument of somite; syn., Depressor muscle crests. Elevated denticles or ridges on inner epimeron, pleurepimere, pleurite, pleuron (pi. pleura), surface of tergum near basicarinal angle for attachment of pleura (pi. pleurae), pleural lobe, tergal fold. depressor muscles (balanomorph Cirripedia). Epimeron. See epimere. Dermal gland. Cell or concentration of cells in epidermis Epipod(ite). Laterally directed exite of protopod, usually traversed by canals communicating with surface through fine branchial in function. ducts. Episternum. Posterolateral projection of various sterna Deuterocerebrum. See mesocerebrum. (Decapoda). Diaeresis. Transverse groove on posterior part of exopod (also Epistome. Plate of varying shape between labrum and bases of rarely of endopod) of uropod; sometimes dividing exopod antennae; also sternum of antennal somite. into 2 movable parts. Esthestasc. See aesthetasc. Distal. Part of structure farthest from midline of body or base Esthete. See aesthetasc. of attachment; opposed to proximal. Exhalant passage. Anterior to gill chamber leading to large Dorsal organ. Thickened glandular area of hypoderm on dor­ anterior opening, with scaphognathite for driving water out­ sal surface, usually in posterior part of cephalon, sometimes ward. in anterior part; not homologous in all taxa. Exite. Laterally directed lobe arising from external margin of Dorsal plate. Spindle-shaped division of carapace intercalated protopodal segment. with median suture in some decapods. Exognath. Exopod of maxilliped. Dorsomedian groove. See decapod carapace, groove e. Exopodfite). Outer ramus of biramous appendage. Dorsoventralis posterior. Prominent muscle connecting head Exoskeleton. Chitinous or calcified outer integument of crusta­ apodemes with inner surface of carapace posterior to cervical ceans. groove in many decapods. Eyestalk. See ocular peduncle. Doublure. Reflected margin of carapace (Stomatopoda). Falcate. Sickle-shaped or hooked. Duplicature. That part of border of shell with calcareous 166 GLOSSARY OF MORPHOLOGICAL TERMS

Filamentary appendage. Membranous process developed on Gastroorbital carina. See decapod carapace, carina d. body in some cirripeds commonly at base of cirrus (Lepa- Gastroorbital groove. See decapod carapace, groove f domorpha, Ascothoracica). Geniculate. Bent; having upper part of filament forming more Filter chamber. Space beneath thorax enclosed by ventral or less obtuse angle with lower, e.g., antennule. body wall and moving thoracopods used for food-gathering. Genital region. See decapod carapace, region f(urogastric). First antenna. See antennule. Gill. See branchia. First maxilla. See maxillule. Gill chamber. See branchial chamber. Fixed finger. Immovable distal part of propodus of chela; syn., Glaucothoe. Stage in larval development of hermit crabs, pollex. equivalent to megalopa. FlabeUum. Thin distal exite (Branchiopoda); or epipodite of Gnathal lobe. Masticatory endite of ihandible; syn., mastica­ thoracopod. tory process. Flagellum (pi. flageUa). Multiarticular^ distal portion of an­ Gnathobases. Paired endites used to manipulate or move tennule, antenna, or exopod. food. Flange. Ridge along valve margin formed by projection of Gnathopod. Prehensile maxilliped; also 1st 2 prehensile outer lamella as narrow brim (Ostracoda). pereopods of amphipods, either chelate or subchelate. Foregut. See stomodeum. Gnathothorax. Cephalothorax with all appendages of fused Fornix. Ridge in lateral part of cephalon above insertion of antennal muscles (Cladocera). thoracic somites modified as maxillipeds. Gonad. Reproductive organ of either sex, communicating with Free edge. Line of contact between closed valves except along hinge line marking distal limit of contact margin, sometimes pair of efferent ducts. inside free margin (Ostracoda). Gonapophysis. Median process from base of 1st or 2nd Free margin. All parts of margin except hingement (Os­ pleopods of male (Syncarida). tracoda). Gonochoristic. Sexes separate; producing distinct males and Frena. Tegumentary folds holding eggs (Cirripedia). females. Frontal appendages. Paired filaments arising from bases of an­ Gonopod. Pleopod modified for reproductive purposes. tennae but independent of them (Anostraca). Gonopore. Outlet for genital products; syn., sexual pore. Frontal band. Glandular organ of adhesion in frontal region of Green gland. See antennal gland. various parasitic copepods used for attachment to host. Haft organ. See frontal organ. Frontal eye. See frontal organ. Head apodeme. Fused endopleurite and endosternite forming Frontal organ. Sensory cells on anterior surface of cephalon; place for muscle attachment at anterior end of skeleton (As- syn., haft organ or frontal eye in non-malacostracans; not tacidea). homologous among taxa. Hemocoel. Lacunar system extending throughout much of Frontal plate. Modified rostrum with downward projecting body, filled with blood. process united with epistome (brachyuran Decapoda). Hemocyanin. Copper-containing respiratory pigment in blood Frontal region. See decapod carapace, region e. (Malacostraca). Frontolateral horn. One pair of tubular frontolateral ex­ Hemoglobin. Oxygen-carrying protein colored substance of tensions of cuticle of cirriped nauplius (except Asco red plasma in blood of some crustaceans. thoracica). Hepatic caecum (pi. caeca). See hepatic cecum. Furca. See caudal furca. Hepatic cecum (pi. ceca). Pouchlike diverticulum generally Furcal ramus. See caudal ramus. connected with mesenteron, with liver function; also see Furcilia stage. Last larval stage marked by movable compound hepatopancreas. eyes projecting beyond margin of carapace; antenna not used for locomotion in later substages (Euphausiacea). Hepatic groove. See decapod carapace, groove g. Galea. Outer distal hoodlike lobe of second segment of maxil­ Hepatic region. See decapod carapace, region g. lule. Hepatic spine. See decapod carapace, spine c. Gastric groove. See stomatopod integumental ornamentation. Hepatopancreas. Digestive gland consisting of tubules ramify­ ing through cephalothorax, with both liver and pancreas Gastric mill. Apparatus in cardiac stomach (stomodeum) with functions. framework of movably articulated ossicles developed as Hermaphrodite. Organism with both male and female repro­ thickened and calcified parts of stomodeal lining used to ductive organs. break up food, mostly highly specialized in decapods; also chitinous triturating apparatus in foregut of some ac- Heterochelate. Chelae of left and right chelipeds differing in rothoracicans (Cirripedia). shape and size. Heteromorph. Adult female, inferred by carapace structure, in Gastric region. See decapod carapace, region f dimorphic genera (Ostracoda). Gastric tooth. See decapod carapace, teeth b. Hindgut. See proctodeum. Gastrolith. Discoid calcareous nodule, common in stomo­ Hinge line. Middorsal line of junction of two valves of deum (Decapoda). carapace, permitting movement between them. GLOSSARY OF MORPHOLOGICAL TERMS 167

Hingement. Collective term for structures comprising articula­ Lateral. One of pair of compartmental plates typically located tion of valves (Ostracoda). between carino- and rostrolaterals, with radius on carinal Hinge nodes. Localized thickened parts of right valve hinge side and ala on rostral side, sometimes between carina and (Phyllocarida). rostrolateral or compound rostrum; homologous with Hinge selvage. Structure of hinge area corresponding to, and lepadomorph median latus (balanomorph Cirripedia). sometimes continuous with, selvage of contact margin (Os­ Lateral bar. Pair of external chitinous thickenings extending tracoda). from apertural thickenings medially down each side of man­ Hypobranchial space. Area of gill chamber below gills. tle sac (Acrothoracica). Hypopharynx. See metastoma. Lateral carina. See decapod carapace, carina e, or stomatopod integumental ornamentation. Hypostoma. See metastoma. Lateral denticle. See stomatopod integumental ornamenta­ Hypostome. See metastoma. tion. Incisor process. Biting portion of gnathal lobe of mandible; Lateral depressor pit. Small depression near basitergal angle syn., pars incisiva. of scutum for attachment of lateral depressor muscle Inferior groove. See decapod carapace, groove h. (balanomorph Cirripedia). Inframedian latus. See lotus. Lateral cups. Paired element of nauplius eye of nonmalacos- Infraorbital spine. See decapod carapace, spine d. tracan crustaceans. Inner lamina. Inner shell layer of compartmental plates sepa­ Lateral gastrocardiac markings. Insertions of attractor epi- rated from outer lamina by longitudinal tubes (balanomorph meralis muscle in brachyuran decapods lacking branchio- Cirripedia). cardiac groove. Interantennular septum. Plate separating antennular cavities Lateral tooth. See decapod carapace, teeth c, or stomatopod in some malacostracans; syn., proepistome. integumental ornamentation. Intercervical groove. See decapod carapace, groove i. Latus (pi. latera). Any of lepadomorph capitular plates except Interlaminate figure. Simple or arborescent lines extending paired scuta and terga and unpaired rostrum, carina, subros- between epicuticle of outer lamina through longitudinal trum, and subcarina. Smaller plates in basal whorls below septa into inner lamina in sections parallel to base in some paired latera referred to as lower latera; sometimes all or balanomorph Cirripedia. some absent. Types of paired plates: Intermediate carina. See stomatopod integumental ornamen­ (a) carinal. Plate on each side of carina (cf. carinolateral). tation. (b) inframedian. Plate below upper latus. Intermediate denticle. See stomatopod integumental ornamen­ (c) median. Plate between rostral and carinal latera in tation. forms with paired latera in one whorl (cf. lateral). Intermediate tooth. See stomatopod integumental ornamenta­ (d) rostral. Plate on each side of rostrum or below scutum tion. (cf. rostrolateral). Intestinal groove. See decapod carapace, groove j. (e) upper. Plate in upper whorl between scutum and ter- Intestinal region. See decapod carapace, region h. gum or carina. Intraparies (pi. intraparietes). Secondary lateral margin of Limadiiformes. Carapace with recurvature of posterior margin carina in some lepadomorph cirripeds. near dorsal line (Conchorstraca). Ischiocerite. Third segment of antennal peduncle. Linea (pi. lineae). Linear marking on carapace. Ischiopodfite). See ischium. Linea anomurica. Longitudinal groove or uncalcified line on Ischium. Third segment of pereopod or 1st segment of en- carapace of many anomuran decapods. dopod articulating with basis; syn., ischiopod(ite). Linea branchiostegalis. Longitudinal groove or uncalcified Jugal region. See decapod carapace, region i. line extending posteriorly from anterior margin of carapace, Kentrogon. Undifferentiated cells following cyprid (Rhizo- slightly above branchiostegal spine, to or slightly beyond cephala). hepatic spine in some caridean decapods. Labium. See metastoma. Linea dromica. Longitudinal groove or uncalcified line on Labrum. Unpaired outgrowth arising just in front of mouth dromioidean decapods comparable with linea thalassinica; and often more or less covering it; syn., upper lip. syn., linea dromiidica. Lacinia. Inner distal spiny lobe of second segment of maxil- Linea dromiidica. See linea dromica. lule. Linea homolica. Longitudinal groove or uncalcified line on Lacinia mobilis. Small, generally toothed, process articulated homoloidean decapods comparable with or equivalent to with incisor process of mandible (Peracarida). linea thalassinica. Lappet. Ventrally projecting subdivisions of pleura (Mysi- Linea lateralis. Longitudinal groove or uncalcified line extend­ dacea). ing posteriorly from frontal margin of carapace below orbit Lasiopod. Appendage (cirrus) with setae in transverse row at sometimes to posterior extremity of carapace in some each articulation (Cirripedia, limited usage). penaeoideans. Latera. See latus. Linea thalassinica. Longitudinal groove or uncalcified line on 168 GLOSSARY OF MORPHOLOGICAL TERMS

dorsal part of carapace extending from anterior margin below Median carina. See stomatopod integumental ornamentation. antennal spine to posterior margin in most thalassinoideans. Median dorsal plate. Elongate plate separating carapace List. Ridge on proximal side of selvage on contact margin (Os- valves posterodorsally in some phyllocarids. tracoda). Median eye. See nauplius eye. Longitudinal canal. See longitudinal tube. Median lotus. See lotus. Longitudinal septum (pi. septa). Partition disposed normal to Megalopa. First postlarval stage in development of many inner and outer laminae of compartmental plate in some Eucarida. balanomorph cirripeds, resulting in longitudinal tubes; syn., parietal tubes. Mereopod(ite). See merus. Longitudinal tube. Canal formed in compartmental plate of Merus. Fourth segment (distally from body), articulating prox- some balanomorph cirripeds between longitudinal septa and imally with ischium and distally with carpus; syn., mero- inner and outer lamina; syn., longitudinal canal; parietal pod(ite). tube, parietal pore. Mesenteron. Midportion of alimentary tract, of endodermal origin, with surface commonly increased by pouchlike exten­ Lower lip. See metastoma. sions serving as digestive glands; syn., midgut. Lunule. Attachment disc at base of antennule in some parasitic Mesobranchial lobe or area. See decapod carapace, region c. copepods. Mesocerebrum. Ganglion of antennular somite; syn., deu- Male-cell receptacle. Pocket or pair of pockets within mantle terocerebrum. cavity of female receiving cells of male cyprid, later differ­ entiated into "testes" (rhizocephalan Cirripedia). Mesogastric lobe or area. See decapod carapace, region/. Mesosome. Collective term for all free thoracic somites behind Manca. Young of some Peracarida lacking last thoracopod at head (not common usage). time of release from marsupium. Mesosternum. Median plate of sternum in some brachyuran Mancoid stage. Postlarval stage with rudimentary 4th pleopod decapods. (Phyllocarida). Metabranchial lobe or area. See decapod carapace, region c. Mandible. One of 3rd pair of cephalic appendages, used to masticate food. Metacerebrum. Ganglion of antennal somite; syn., tritocere- brum. Mandible body. Inflated base of mandible for attachment of mandibular muscles; syn., corpus mandibulae. Metagastric lobe or area. See decapod carapace, region/. Mandibular foramen. Relatively large opening in mandibular Metanauplius. Postnaupliar larva with generally same body body for passage of transverse adductor muscle. and body shape as nauplius, but with additional appendages. Mandibular palp. Distal articulated part of mandible used in Metasoma. See metasome. feeding or cleaning; also nonsegmented part of mandible in Metasome. Part of prosome, consisting of free thoracic somites Acrothoracica; attached to labrum in thoracic Cirripedia. anterior to major articulation (Copepoda); or first three ab­ Mantle. Membranous covering of body, often strengthened by dominal somites (Amphipoda, not general usage); syn., calcareous plates in thoracic Cirripedia (cf. carapace). metasoma. Mantle cavity. Space occupied by body, with aperture (Cir­ Metastoma. Lower lip posterior to mandibles, usually cleft ripedia). into pair of lobes (paragnaths); syn., hypostoma, hypostome, hypopharynx, labium. Manus. Broad proximal part of propodal cheliped; syn., palm. Marginal carina. See stomatopod integumental ornamenta­ Metazoea. Last zoeal substage of larval brachyurans. tion. Metapon. Entire preoral area, including part of mandibular Marginal groove. See decapod carapace, groove k. somite (Decapoda). Marsupium. Brood pouch. Micronicus. See epicaridum. Masticatory process. See gnathal lobe. Midgut. See mesenteron. Mastigobranch. Slender respiratory process at base of epipod; Molar process. Grinding portion of gnathal lobe of mandible; syn., mastigobranchia. syn., pars molaris. Mastigobranchia (pi. mastigobranchiae). See mastigobranch. Mouth cirri. First pair of cirri considerably modified (ac- Mastigopus stage. Larval stage in development of some de­ rothoracican Cirripedia). capods equivalent to adult of Leucifer. Movable finger. Dactyl of chela. Maxilla (pi. maxillae). Paired appendage of 5th cephalic so­ Mucro (pi. mucrones). Spine on inferoposteal angle of mite, used in feeding, often also in respiration; syn., 2nd carapace in some Cladocera. maxilla. Muscle scar. Mark on interior of valve or carapace indicating Maxillary gland. Excretory organ in maxillary somite with position of muscle attachment, generally distinguishable by duct opening on maxilla; syn., shell gland. localized difference in surface texture, elevation, depression, MaxUUpeds. Paired appendages modified for feeding on 1st, or delimiting narrow groove. up to 3rd, thoracic somites, usually fused to cephalon. Mysis stage. Larval stage in penaeoidean development equiva­ Maxillule. One of pair of 4th cephalic appendages, usually lent to zoeal stage of Nephropoidea; cf. zoea; syn., schizopod serving as mouthpart; syn., 1st maxilla. larva. GLOSSARY OF MORPHOLOGICAL TERMS 169

Nauplius (pi. nauplii). Early larval stage with only 3 pairs of Orbital tooth. See decapod carapace, teeth d. appendages, antennules, antennae, and mandibles. Orifice. Opening in sessile barnacle wall occupied by oper­ Nauplius eye. Unpaired median eye, common in nauplii and culum (cf. aperture). many adult crustaceans; structure simple, consisting of 1 to Ostium (pi. ostia). Valved opening in heart for return of few light-sensitive cells; syn., median eye. blood. Neck organ. See nuchal organ. Oviduct. Passageway from ovary to genital aperture. Nephropore. Elevated opening of antennal gland on ventral Ovigerous frena (pi. frenae). Fleshy ridge or flap on inner surface of coxa of antenna. surface of mantle adhering to and holding egg masses in Nisto. Postlarval stage of some palinurans; syn., pseudihacus, place in some lepadomorph Cirripedia. puerulus. Ovigerous lamella. Adherent egg masses forming one or more Notum. Posterior part of dorsal carapace in shrimplike deca­ lamellae within mantle cavity, in some lepadomorph Cir­ pods (limited usage). ripedia, held in place by ovigerous frena. Nuchal organ. Sense organ on upper side of cephalon (Bran- Palate. See endostome. chiopoda); syn., neck organ. Palm. See manus. Occipital notch. Angulated indentation at rear of cephalon Palp. Usually one ramus (endopod), sometimes both, and (Conchostraca). basis, reduced distally to 1 to 3 segments, associated with Occludent margin. Margin of scutum and tergum bordering mouthparts; also nonsegmented setose structure attached to orifice (thoracic Cirripedia). mandible (Acrothoracica) and to lateral margin of labium in Occludent teeth. Small projections on occludent scutal mar­ thoracic Cirripedia. gin, formed by extensions of external growth ridges, inter- Palp foramen. Small opening in mandibular body com­ digitating with teeth on margin of opposed scutum (balano- municating with mandibular palp. morph Cirripedia). Parabranchial groove. See decapod carapace, groove I. Ocellus. Simple single eye or eye spot; common in some bran- Paracopulatory organ. Specialized endopod of pleopod in chiopods and copepods (cf. nauplius eye). some isopods; used in copulation. Ocular bulla. Knob on inner surface of carapace connecting Paragnath(s). See metastoma. lower and upper orbital margins with basal segment of an­ Paries (pi. parietes). Median triangular part of each com- tenna, serving to protect eye. partmental plate in sessile barnacles. Ocular papilla. Anterior projection on eyestalk (Mysidacea Parietal pore. See longitudinal tube. and one Anaspidacea). Parietal septum. See longitudinal septum. Ocular peduncle. Peduncle movably articulated with cepha­ Parietal tube. See longitudinal tube. lon, with compound eye at distal end, sometimes with two or three segments, sometimes retractable; syn., eyestalk. Pars ampuUaris. Bottle-shaped diverticulum at entrance of ceca into pyloric chamber of stomach (Hoplocarida, Anas­ Olfactory hair. See aesthetasc. pidacea). Ommatidium (pi. ommatidia). Cylindrical or prismoidal visual Pars incisiva. See incisor process. constituent of compound eye covered by transparent cuticle (cornea). Pars molar is. See molar process. Oostegite. Inner, medially directed lamella arising from coxa of Parva stage. First postlarval stage in development of caridean pereopod in female, part of midventral marsupium (Pera- decapods. carida). Pedigerous. Bearing footlike appendages. Opercular membrane. Thin, flexible membrane attaching Peduncle. Stalk in lepadomorph cirripeds, supporting opercular valves to sheath, periodically shed during ecdysis, capitulum, attached to substrate by opposite end, commonly in balanomorph Cirripedia; represented by membranous armed with calcareous scales. hinge in verrucomorph Cirripedia. PeniciUa (pi. peniciUae). Dentate setae on mandible (Stygo- Opercular valve. One of 2 or 4 movable plates occluding aper­ caridacea). ture in sessile barnacles. Penicillus (pi. penicilli). Tufts of fine hair resembling small Operculum. Scuta and terga (balanomorph) or movable brush. scutum and tergum (verrucomorph) and associated mem­ Penis (pi. penes). Male copulatory organ; probosciform in brane, forming apparatus occluding aperture. hermaphroditic thoracic Cirripedia and greatly distensible. Ophthalmic somite. See acron. Peraeon. Seepereon. Optic lobe. Ganglion of brain or nervous system for innerva­ tion of eye. Peraeonite. See pereonite. Orbit. Circular to rectangular opening in anterior face of Peraeopod. See pereopod. carapace supporting ocular peduncle in some decapods. Pereion. See pereon. Orbital carina. See decapod carapace, carina f. Pereionite. See pereonite. Orbital hiatus. Gap or slit in orbital margin of carapace. Pereiopod. See pereopod. Orbital region. See decapod carapace, region j. Pereon. Anterior portion of trunk with thoracopods, exclu- 170 GLOSSARY OF MORPHOLOGICAL TERMS

sive of maxillipedal somites and appendages; syn., peraeon, Posterior carina. See decapod carapace, region h. pereion. Posterior gastric pit. One of two small depressions dorsally Pereonite. Somite of pereon; syn., peraeonite, pereionite. near midline on exterior of carapace marking point of inser­ Pereopod. Thoracic appendage used in locomotion; syn., tion of stomach muscle (Decapoda). peraeopod, pereiopod, ambulatory leg, walking leg. Posterior tooth. See decapod carapace, teeth e. Pericardium. Blood sinus surrounding heart and communicat­ Postlarval stage. Developmental stage reached after comple­ ing with it by paired ostia. tion of megalopal or equivalent metamorphosis; marked by Peritrophic membrane. Chitinous sheath secreted around initial appearance of adult characters. feces. Postorbital carina. See decapod carapace, carina h. Petasma. Abdominal structure developed from male pleopods, Postorbital spine. See decapod carapace, spine e. used in copulation. Postrostral carina. See decapod carapace, carina i. P-4 structure (type A). Structure on lateral face of propodus of Postrostral spine. See decapod carapace, spine f. 4th pereopod in some pagurids; presumably sensory in func­ Postsegmental region. See telson. tion. Precoxa. Segment of protopod proximal to coxa, rarely pres­ Photophore. Luminous organ. ent; syn., pleuropod. Phyllobranch. Gill with leaflike filaments; syn., phyllobranchia. Pre-epipod(ite). Laterally directed lobe of coxa or from coxal Phyllobranchia. See phyllobranch. position. PhyUopodium (pi. phyUopodia). Leaflike thoracic appendages. Pregastric tooth. See decapod carapace, teeth f. PhyUosoma. Larval stage in development of palinuroidean Preischium. Segment of endopod between protopod and is­ decapods. chium, rarely present. Pleomere. See abdominal somite. Prelateral lobe. Proximalmost lateromarginal lobe of telson Pleon. See abdomen; also first 3 abdominal somites of am- (Stomatopoda). phipods. Preoral sting. Retractile piercing mechanism with poison Pleonite. See abdominal somite. gland (Branchiura). Pleopod. Paired appendage of any of first 5 abdominal somites Presegmental region. See acron. (rarely 6) in Malacostraca, adapted for swimming; syn., Preungal process. Structure at base of dactyl of 4th pereopod swimmeret. in many pagurids; function unknown but presumed sensory. Pleotehon. Structure formed by fusion of one or more abdom­ Prezoea. Just-hatched postnaupliar larva still covered by em­ inal somites with telson. bryonic cuticle. Pleura (pi. pleurae). See epimere. Primordial valve. One of 5 chitinous plates (scuta, terga, and Pleural lobe. See epimere. carina) in cyprid larvae of lepadomorphs and verrucomorphs, Pleural suture. Line of separation of carapace in molting. site of calcification during metamorphosis, sometimes visible at umbones of these plates in adults. Pleurepimere. See epimere. Pleurite. See epimere. Proctodaeum. See proctodeum. Pleurobranch. Gill attached directly to body wall (Decapoda); Proctodeum. Posterior part of alimentary canal lined with syn., pleurobranchia. cuticle of ectodermal origin; syn., hindgut, proctodaeum. Proepistome. See interantennular septum. Pleurobranchia. See pleurobranch. Pleuron (pi. pleura). See epimere. Propodus. Penultimate segment of pereopod (thoracopod). Pleuropod. See precoxa. Prosartema. Scale implanted on inner margin of basal segment of antennular peduncle (Penaeoidea). Podobranch. Gill arising from coxa of thoracopod; syn., podobranchia. Prosoma. See prosome. Podobranchia. See podobranch. Prosome. Anterior region of body, commonly limited pos­ teriorly by major articulation. In cirripeds, large saclike body Podomere. See segment. in position of "head" in front of, and rostral to, thoracic ap­ Podophthalmite. One of 2 segments of eyestalk (when seg­ pendages, supporting trophi and usually 1st cirri; syn., pro­ mented), bearing cornea. soma. PoUex. See fixed finger. Protandry. Hermaphroditic condition with male elements Pore canal. Minute tubular passageway extending through maturing and being released before maturation of female shell (Ostracoda). elements. Postabdomen. Terminal part of body (Cladocera). Protocephalon. See acron. Postcephalic groove. See decapod carapace, groove m. Protocerebrum. Ganglion of preantennulary region. Postcervical groove. See decapod carapace groove n. Protogastric lobe or area. See decapod carapace, region/. Postcervical notch or incision. Strong indentation of carapace Protogyny. Hermaphroditic condition with female elements at level of postcervical groove. maturing and being released before maturation of male ele­ Posterior cardiac lobe. See decapod carapace, region h. ments. GLOSSARY OF MORPHOLOGICAL TERMS 171

Protopod(ite). Proximal part of appendage, consisting of coxa sometimes fused with rostrum, laterally or to each other; and basis or less frequently of precoxa, coxa, and basis, some­ homologous with lepadomorph rostral latus (balanomorph times fused; syn., sympod. Cirripedia). Protozoea. First two or three postnaupliar substages in devel­ Rostrum. (1) Anteriorly projecting, unpaired, usually rigid opment of Penaeiodea and Euphausiacea (cf. calyptopis in median extension of carapace between eyes or ocular pedun­ latter). cles; (2) anterior beaklike projection of valve margins over­ Proventriculus. Elaborated anterior part of alimentary canal in hanging incisure or notch (Ostracoda); (3) unpaired antero- some crustaceans. ventral plate of thoracic Cirripedia; in lepadomorphs, 1 of up Proximal. Part of structure nearest midline of body or base of to 4 unpaired plates of capitulum; in verrucomorphs, com­ partmental plate between carina and fixed scutum; in attachment; opposed to distal. balanomorphs, compartmental plate overlapping adjacent Pseudepipod(ite). Lateral lobe arising from distal part of basis plates, sometimes compound, either fused with rostrolater- or proximal part of exopod. als or missing and formed of fused rostrolaterals; simple or Pseudibacus. Postlarval stage of some palinurans; syn., nisto, compound with or without radii on each side. puerulus. Saw bristles. Heavy setae in row on gnathal lobe of mandible Pseudorostrum. Anterior part of carapace formed by pair of between molar and incisor processes. forward projecting plates (Cumacea). Scale. Small calcareous plates on peduncle of lepadomorph Cirripedia; see also scaphocerite. Pseudotrachea. Respiratory structure developed in pleopods Scaphocerite. Exopod of antenna; syn., scale, squama. of some terrestrial isopods for air-breathing. Scaphognathite. Exopod of maxilla, often used to produce re­ Pterygostome. See pterygostomial region. spiratory current in gill chamber. Pterygostomial region. See decapod carapace, region k. Schizopod larva. See mysis stage. Pterygostomial spine. See decapod carapace, spine g. Scutal margin. Edge of tergum articulating with scutum or Puerulus. See pseudibacus. edge of any other adjacent to scutum (thoracic Cirripedia). Punctum (pi. puncta). Small pitlike depression^) in valve sur­ Scutum (pi. scuta). Paired plate or valve of thoracic Cirripedia; face (Ostracoda). in lepadomorphs, 1 on each side of occludent margin of Pustula (pi. pustulae). Small protuberance(s) on valve surface capitulum. In verrucomorphs, of two types: fixed scutum, 1 with pore at summit (Ostracoda). Radial lirae. See carapace costae. of 4 compartmental plates; movable scutum, 1 of 2 opercular Radius (pi. radii). Lateral part of compartmental plate when plates. In balanomorphs, 1 of 4 opercular plates. marked off from paries by change in direction of growth Second antenna. See antenna. lines; overlaps ala of adjoining compartmental plate. Second maxilla. See maxilla. Ramus. Branch of appendage or other structure (e.g., flagel- Segment. Individual component of crustacean appendage con­ lum). nected by movable articulation with adjoining segments; Raptorial claw. Toothed dactyl, generally strong, curved syn., podomere (cf. article). backward on propodus, modified for quick motion in catch­ Seller groove. See decapod carapace, groove o. ing prey. Selvage. Middle (principal) ridge of contact margin sealing Rasp. One or several rows of chitinous plates or scales on sur­ valves closed (Ostracoda). face of pereopodal or uropodal segments. Seminal receptacle. Diverticulum of oviduct or external pouch Receptaculum seminalis. See seminal receptacle. for storing spermatozoa delivered by male; syn., recep­ Retinaculum (pi. retinacula). Small hook at tip of appendix taculum seminalis, spermaiheca. interna, one of many serving to interlock right and left Seminal vesicle. Sac in male, independent of testes, for storage pleopods; syn., cincinnulus. of spermatozoa; syn., vesicula seminalis. Rostral angle. Intersection of basal and occludent margins of Serration. Irregular saw-toothed outline, e.g., on conchostra- scutum in lepadomorph Cirripedia. can valves. Rostral carina. See decapod carapace, carina j. Seta (pi. setae). Hairlike process of cuticle with which it articu­ Rostral incisure. Gap below rostrum in anterior margin of lates or through which it protrudes. valve for protrusion of antenna (Ostracoda); syn., rostral Sexual pore. See gonopore. notch. Sheath. Cylindrical structure inside shell, consisting of alae Rostral latus. See latus. and thickened upper part of parietes, greatly strengthening Rostral notch. See rostral incisure. orifice and providing for attachment of opercular membrane (balanomorph Cirripedia). Rostral plate. Anteriorly projecting, unpaired, movably articu­ lated, median extension of carapace (Phyllocarida); anten- Shell fold. Part of carapace behind cephalon. nulary portion of cephalon (Mystacocarida). Shell gland. See maxillary gland. Rostral tooth. See decapod carapace, teeth g. Shield. Anterior part of cephalothorax in hermit crabs. Rostrolateral. One of pair of compartmental plates typically Simple velum. Velate structure having simple flangelike form overlapping rostrum on each side, with radii on both sides, or forming ridge (Ostracoda). 172 GLOSSARY OF MORPHOLOGICAL TERMS

Sinus gland. Storage release site for molt-inhibiting and other (b) lateral. Small projection(s) at medial base of each lateral hormones produced by x-organ and other sites of central tooth on telson. nervous system. (c) submedian. Small projection(s) just laterad of midline Skeletal duplicature. Outer chitinous body cover shed during on terminal margin of telson (medial to submedian ecdysis (Conchorstraca, Notostraca, Cladocera). teeth). Somite. Division of body, including exoskeleton, usually with Grooves: pair of appendages. (a) cervical. Distinct transverse depression or groove in Spermatheca. See seminal receptacle. posterior half of carapace. Spermatophore. Packet or capsule of spermatozoa. (b) gastric. Longitudinal pair of grooves, extending from Spur. Pendent projection from basal margin of tergum (sessile base of rostrum to posterior margin of carapace. Cirripedia); also velate structure modified as flattened Teeth: spinelike projection in some dimorphic genera (Ostracoda). (a) intermediate. Strong spinelike or blunt projection at Spur fasciole. Nearly level, slightly depression on outer sur­ distolateral angle of telson, between submedian and lat­ face of tergum extending to apex in line with spur; usually eral teeth. delimited on 1 or both sides by narrow groove (balanomorph (b) lateral. Strong spinelike or blunt projection on lateral Cirripedia). margin of telson distally, between intermediate tooth Spur furrow. Groove on outer surface of tergum extending to and prelateral lobe. apex in line with spur, with sides sometimes folded in (sessile (c) submedian. Strong spinelike or blunt projection just Cirripedia). laterad of midline on terminal margin of telson. Squama. See scaphocerite. Stomodaeum. See stomodeum. Statocyst. Diminutive organ providing sense of balance. Stomodeum. Anterior part of alimentary tract, ectodermal in Stenopodium. Slender, elongate appendage, composed of rod­ origin and lined with cuticle; cast with molt; syn., foregut. like segments. Stridulating organ. Structure producing sound by 2 parts of Sternum canal. Internal skeletal structure of some crabs exoskeleton being rubbed together; surface with ridge or formed by meeting of sternal apodemes of opposite sides tubercles or cross-ridges surface and opposing surface with above nerve cord. single transverse ridge or tubercle. Sternal plastron. See sternum. Stylamblys. See appendix interna. Sternal process. Projection from midsection of sternite Style. See telson. (Mysidacea). Stylet. See caudal ramus. Sternite. Sclerotized ventral surface of body somite. Stylocerite. Rounded or spiniform process on outer part of Sternum (pi. sterna). Structure comprised of ventral portions proximal segment of antennular peduncle; syn., antennular of somites taken together; syn., sternal plastron. scale. Stipe. Stemlike basal part of appendage with exopod some­ Subcarina. Unpaired plate below carina (lepadomorph Cir­ times squamate. ripedia). Stomatopod integumental ornamentation. Armature and or­ Subchela. Distal end of appendage developed as prehensile namentation of carapace, thoracic somites, abdomen, and structure by folding back of dactyl against propodus or some telson; named types include: broadened part of it; also may result from propodus folded Carinae: back against carpus; e.g., gnathopod. (a) intermediate. Between submedian and lateral carinae Subchelate. Provided with subchela. on each side of carapace. Subesophageal ganglion. Nerve plexus below esophagus. (b) lateral. Between intermediate and marginal carinae on Subhepatic carina. See decapod carapace, carina k. each side of carapace and abdomen; extending pos­ Subhepatic region. See decapod carapace, region I. teriorly from near anterolateral angle nearly to margin on carapace. Submedian carina. See decapod carapace, carina I or stomatopod integumental ornamentation. (c) marginal. Carinae of lateral margins on abdomen; ex­ tending along each lateral margin of carapace, often Submedian denticle. See stomatopod integumental ornamen­ curving upward posteriorly to loop around lateral tation. carinae. Submedian groove. See decapod carapace, groove p. (d) median. Middorsal carina, sometimes bifurcate an­ Submedian tooth. See stomatopod integumental ornamenta­ teriorly on either side of cervical groove. tion. (e) submedian. Slightly laterad of midline and median Suborbital region. Narrow region bordering lower margin of carina in abdominal region, not present on carapace. orbit, sometimes denned, sometimes indistinguishable Denticles: (Brachyura); see decapod carapace, region m. (a) intermediate. Row of small projections between inter­ Suborbital spine. See decapod carapace, spine h. mediate and submedian teeth on lateroterminal margin Subrostrum. Unpaired plate below rostrum (lepadomorph of telson. Cirripedia). GLOSSARY OF MORPHOLOGICAL TERMS 173

Suctorial structures. Mouthparts of ectoparasites modified for Tritocerebrum. See metacerebrum. piercing bodywall of host and for sucking out body fluids. Trophi. All mouthparts of cirripeds. Sulcus. Groove or furrow. Trunk. Postcephalic portion of body. Supra-anal plate. Tongue-shaped, spatulate, or rounded plate Umbo. Apical portion of either valve of bivalved crustaceans; produced posteriorly on dorsal side of telson (Notostraca). also location on plate from which successive growth incre­ Supraesophageal ganglion. Nerve plexus above esophagus, ments extend in lepadomorph Cirripedia. equivalent to brain. Umbonal spine. Hollow, minute to large, curved, looped, or Supraorbital carina. See decapod carapace, carina m. nodular spinose projection, sometimes involving entire Sutural edge. Margin of compartmental plate along suture umbo (Conchostraca). (Cirripedia). Upcurved growth lines. Upwardly bent growth lines covering tear in shell margin at site of injury (Conchostraca). Suture. Line or seam at juncture of 2 compartmental plates (Cirripedia); also weakly calcified areas of integument Upper latus. See lotus. separating at ecdysis. Upper lip. See lahrum. Swimmeret. See pleopod. Urogastric groove. See decapod carapace, groove q. Sympod(ite). See protopod. Urogastric lobe or area. See decapod carapace, region f. Tagma (pi. tagmata). Major division of body (e.g., head, Uropod(ite). Appendage of 6th abdominal somite of Malacos- thorax, abdomen), each composed of varying number of so­ traca, generally fanlike, sometimes reduced or modified; also last 3 pairs of abdominal appendages in amphipods. mites. Tailfan. See caudal fan. Urosoma. See urosome. Telopod. Part of appendage distal to coxa (limited usage). Urosome. Part of body behind major articulation (Copepoda); also last 3 abdominal somites bearing modified appendages Telson. Terminal portion of body (not considered to be true somite), usually bearing anus, sometimes with caudal furca; (Amphipoda). syn., postsegmental region, style. Valve. Lateral part of divided carapace, commonly joined to opposite part by hingement along dorsal midline; also any Tergal fold. See epimere. one of opercular elements in sessile Cirripedia. Tergal margin. Edge of scutum adjacent to tergum, or edge of Vas deferens. Duct in male for passage of sperm from testis to any plate bordering tergum (Cirripedia). penis. Tergite. Sclerotized dorsal surface of single body somite. Ventral comb. Row of setae or bristles on posteroventral mar­ Tergum (pi. terga). Paired plate or valve of thoracic Cirripedia; gin of last abdominal somite or telson (Cephalocarida). in lepadomorphs, 1 on each side of occludent margin at apex Ventral cup. Element of nauplius eye in nonmalacostracans. of capitulum. In verrucomorphs, of two types: fixed tergum, Ventral frontal organ. Paired sensory structure associated with 1 of 4 compartment plates, movable tergum, 1 of 2 opercular the nauplius eye of nonmalacostracan crustaceans. plates. In balanomorphs, 1 of 4 opercular plates. Ventral nerve chain. See ventral nerve cord. Terminal cirri. Cirri, except first pair, located at posterior end Ventral nerve cord. Ganglia or connectives in somites joined of thorax (Ascothoracica). by single or double tract of nerve fibers running longitudi­ Terminal claw spines. Toothlike projections of varying size at nally beneath alimentary canal. concave end of postabdomen (Cladocera). Vertex. Top part of head or cephalon (Cladocera). Thelycum. External pocket on ventral side of thorax in pe- Vesicula seminalis. See seminal vesicle. naeid female serving as seminal receptacle. Vestibule. Space between duplicature and outer lamella (Os- Thoracic appendage. Any appendage attached to somite of tracoda). thorax (cf. thoracopod). Walking leg. See pereopod. Thoracomere. Somite of thorax. X-organ. Site of secretion of molt-inhibiting and other hor­ Thoracopodfite). Appendage of any thoracic somite; see mones. Located in eyestalks of most decapods or cephalon of thoracic appendage, phyllopod, maxilliped, pereopod, syn., sessile-eyed crustaceans. Also referred to as paired frontal cormopod. organ in Anostraca. Thorax. Tagma between cephalon and abdomen, i.e., anterior Y-organ. Site of secretion of molting hormone in decapods. part of trunk. Zoea. Larval stage characterized by natatory exopods on some Transverse septum. Thin wall normal to longitudinal septum or all thoracic appendages, pleopods absent or rudimentary; and parallel to basis, dividing longitudinal tubes into series syn., mysis, phyllosome, protozoea, schizopod larva. of cells (balanomorph Cirripedia). Trichobranchfia). Gill with filamentous structure of hairlike projections from axis.

INDEX

Pages on which illustrations appear are indicated in italic.

Acron, 64 Body rings, 8, 9 Order Decapoda, 126—127 Acrothoracica, 3, 44- 46,45 Bopyridae, 107 Clibanarius, 148 Ala(e), 38, 41, 52 Boreomysis, 83 Clypeus, 12 Albuneidae, 148 Brachyrhyncha, 126, 152 Coenobitoidea, 126, 145, 148 Allanaspides, 71, 73 Brachyura, 126, 151-158,153,154,155 Complemental male, 47, 54 Alpheidae, 134, 136 Branchinecta, 12 Conchostraca, 3, 16- 19,17 Alpheoidea, 126 B. gigas, 12 Copepoda, 3, 32-40, 35, 36, 39 Amphionidacea, 3, 123-124,125 Branchiopoda, 3, 8 Corfrontale, 104, 122, 136, 138, 148 Amphionides, 124 Branchiura, 3, 41-43, 42 Coronida, 69 A. reynaudii, 123, 124 Bresilioidea, 126 Corystoidea, 127 Amphipoda, 3, 109-117, 111, 113 Brood pouch, 20, 60, 124, 145. See also Coxal plate, 104, 112 Anaspidacea, 3, 70-74, 72, 73 Oostegites Crangonidea, 126, 136 Anaspides, 71, 73 Cumacea, 3, 88- 91,89 A. tasmaniae, 71, 73 Caenestheriella, 16 Cyclomorphosis, 19 Anaspididae, 71 Calanoida, 3, 33, 34, 36- 37 Cyclopoida, 3, 33, 34, 36-37 Annulus ventralis, 138 Calanus, 36, 37 Cymothoidae, 106 Anomura, 126, 144-151,146,147,149 Calappoidea, 126 Cypridinidae, 24, 26- 27 Anostraca, 3, 11-15,13,14 Calmanostraca, 3 Cyzicidae, 16 Antennal glands, 14, 36, 83, 114, 122, Cancridea, 126, 152 131, 136, 141, 150 Caprellidea, 3, 110, 111, 114-115 Daphnia, 20-22 Anthuridea, 96, 100, 104 Caridea, 126, 133-137, 135 Decapoda, 3, 126-157 Appendix interna, 60, 66, 124, 136 Caudal appendage, 48 Dendrobranchiata, 126, 127-132,128, Appendix masculina, 104, 129, 133, 136, Caudal furca, 24, 28, 30, 55, 60, 75. See 129, 138 138 also Caudal rami Derocheilocaris, 28 Apus. See Triops Caudal rami, 6, 9, 12, 18, 20, 34, 38, 43, Digestive system Argulus, 41 78 Amphipoda, 113-114 Ascothoracica, 3, 54- 56, 57 Cement gland, 15, 46, 51 Anaspidacea, 71, 73 Ascothorax ophioctenis, 55 Cephalic shield, 6, 34 Anostraca, 14 Asellota, 98 Cephalocarida, 3, 4-8, 5, 7 Astacidea, 141 Astacidea, 126, 137-142, 139,140 Cephalosome, 34 Brachyura, 156 Astacoidea, 126, 138 Chiltoniella, 4, 6 Cirripedia, 51 Austroastacidae, 143 C. elongata, 4 Cladocera, 20 Austroastacidea, 126, 142-143 Chonopeltis, 41 Copepoda, 36 Austroastacus, 142 Chthamalus, 52 Dendrobranchiata, 131 A. hemicirratulus, 142 Circulatory system Euphausiacea, 122 Amphipoda, 112-113 Isopoda, 105-106 Balanomorpha, 52— 54, 53 Anaspidacea, 71 Leptostraca, 62 Balanus, 52 Anostraca, 12 Mysidacea, 83 Basic characters Astacidea, 138, 141 Ostracoda, 26 Branchiopoda, 8 Brachyura, 152 Paguroidea, 150 Crustacea, 2 Caridea, 136 Stomatopoda, 67 Decapoda, 126 Cirripedia, 51 Dikonophora, 92, 94 Eucarida, 117 Cladocera, 20 Diplostraca, 3 Eumalacostraca, 70 Copepoda, 34, 36 Dipteropeltis, 41 Malacostraca, 59 Dendrobranchiata, 129—131 Dolops, 41 Peracarida, 79 Euphausiacea, 122 Dorippoidea, 127 Syncarida, 70 Isopoda, 104-105 Dorsal organ, 9 Thoracica, 46— 47 Leptostraca, 62 Dorsal shield, 8, 9, 18 Bathynellacea, 75-78, 76 Mysidacea, 82- 83 Dromiacea, 126 Bathynellidae, 77 Ostracoda, 24, 26 Dromioidea, 126, 151, 152 Bathynomus, 104, 105, 106 Paguroidea, 148, 150 Duplicature, 24 B. giganteus, 100 Stomatopoda, 67 Bellioidea, 127 Cirri, 46, 47, 48, 49 Emerita, 148 Bentheuphausia, 117, 118, 120 Cirripedia, 3, 44-58,49, 50, 53, 57 Engaeus, 142 Bivalve carapace, 16, 20, 24, 47, 55, 60 Cladocera, 3, 19-22,21 Ephippium (ephippia), 22 Bodotriidae, 90 Classification Epicaridea, 106 Body flexure, 34 Superclass Crustacea, 3 Eryonoidea, 126 176 INDEX

Eucarida, 3, 117 Harpacticoida, 3, 33, 34, 36-37 Limnadiidae, 18 Eucopia, 81 Hippoidea, 126, 145, 148 Limnobaena, 84 Eucopiidae, 81 Hippolyte, 136 Lithodidae, 145 Eumalacostraca, 3, 70 Homoloidea, 127 Lomidae, 145 Euphausia, 120 Hoplocarida, 3, 63 Lophogaster, 81 Euphausiacea, 3, 117- 123,119,121 Hutchinsoniella, 4, 6 Lophogastrida, 79, 81 Euphausiidae, 118 H. macrocanthus, 6 Loxoconcha, 24 External morphology Hyperiidea, 110, 112 Luminescent organs, 118 Acrothoracica, 45—46 Lunule, 38 Amphionidacea, 124 Idotea, 105 Lynceidae, 16 Amphipoda, 110, 112, 115 Ingolfiellidea, 110, 114 Lysiosquilla, 69 Anaspidacea, 71 Isopoda, 3, 95-109,101,102,103, Anomura, 145, 148 Majoidea, 127 Anostraca, 12 Kalliapseudidae, 94 Malacostraca, 3, 59 Ascothoracica, 55 Kentrogon, 58 Maxillary gland, 9, 14, 36, 66, 73, 83, Astacidea, 138 Koonuga, 71 106 Austroastacidea, 143 Koonugidae, 71 Maxillary hooks, 38 Bathynellacea, 77- 78 Maxilliped somite, 30 Brachyura, 152 Lacinia mobilis, 79, 81, 85, 88, 91, Meganyctiphanes, 120 Branchiura, 41, 43 104, 112, 115 Metasome, 34 Caridea, 134, 136 Larval development Micraspides, 71 Cephalocarida, 4, 6 Acrothoracica, 45 Mimilambroidea, 127 Cladocera, 20 Amphipoda, 115—116 Misophrioida, 3, 34 Conchostraca, 16, 18 Anaspidacea, 73 Monodella, 84, 85 Copepoda, 34, 38 Anomura, 150 Monodellidae, 85 Cumacea, 90- 91 Anostraca, 15 Monokonophora, 92, 94 Dendrobranchiata, 127, 129 Astacidea, 141 Monstrilloida, 3, 37 Euphausiacea, 118 Bathynellacea, 78 Myodocopa, 3, 23-27 Isopoda, 100, 104, 107 Brachyura, 156 Mysida, 79, 81 Leptostraca, 60, 62 Branchiura, 43 Mysidacea, 3, 79- 84, 80 Mysidacea, 81-82 Caridea, 136 Mysis, 83 Mystacocarida, 28, 30 Cephalocarida, 6 Mystacocarida, 3, 28-31, 29 Notostraca, 9 Cladocera, 22 Ostracoda, 24 Conchostraca, 18 Nannastacidae, 90 Rhizocephala, 56 Copepoda, 37 Nebalia, 59, 60, 62 Spelaeogriphacea, 88 Cumacea, 91 N. typhlops, 59 Stenopodidea, 133 Dendrobranchiata, 131 Nebaliacea, 60 Stomatopoda, 64, 66- 67 Euphausiacea, 122-123 Nebaliella, 59, 60, 62 Stygocaridacea, 75 Isopoda, 106, 107 Nebaliidae, 60 Tanaidacea, 92, 94 Leptostraca, 60 Nebaliopsis, 59, 60, 62 Thermosbaenacea, 85 Mysidacea, 83 N. typicus, 60 Thoracica, 48, 50, 52 Mystacocarida, 30 Nematobrachion, 120 Notostraca, 9 Nematoscelis, 120 Filamentary appendages, 48 Ostracoda, 27 Nephropidae, 126, 138 Flabellifera, 96-97, 106 Rhizocephala, 56, 58 Nephropoidea, 126, 138 Food basket, 118 Stenopodidea, 133 Nervous system Food groove, 9, 12 Stomatopoda, 69 Amphipoda, 114 4-celled sense organ, 71 Stygocaridacea, 75 Anaspidacea, 73 Frontal organ, 15, 18, 20, 24 Tanaidacea, 94 Anostraca, 15 Frontal plate. See Clypeus Thermosbaenacea, 85 Astacidea, 141 Thoracica, 47 Brachyura, 156 Galatheidae, 145 Lepadidae, 48 Cirripedia, 51 Galatheoidea, 126, 145, 148 Lepadomorpha, 47- 51,49, 50 Cladocera, 20, 22 Gammaridea, 109- 110, 112 Lepas, 48, 50 Copepoda, 36 Gastric mill, 131, 141, 150, 156 Lepidurus, 8, 9 Dendrobranchiata, 131 Gills (or branchial apparatus), 66, 81, 88, Leptobathynellidae, 77 Euphausiacea, 122 90, 104, 115, 118,126, 129, 133, Leptochelia, 92 Isopoda, 106 138, 152, 156 Leptodora, 19, 20, 22 Leptostraca, 62— 63 Glypheoidea, 126 Leptodoridae, 20 Mysidacea, 83 Gnathiidea, 99-100, 104, 106 Leptostraca, 3, 59- 63, 61, 62 Ostracoda, 26 Gnathophausia, 81 Leuconidae, 90 Paguroidea, 150 Gnathopod, 101, 112, 115 Lifting spines, 85. See also Setal row Stomatopoda, 67—68 Gonopophyses, 70 Lightiella, 4, 6 Notodelphyoida, 3, 37 Grapsidoidea, 127 L. floridana, 4 Notostraca, 3, 8-11,10 L. incisa, 4, 6 Nuchal organ, 12. See also Dorsal Halobaena, 84 L. serendipita, 6 organ Hapalocarcinoidea, 127 Ligia, 105 Nyctiphanes, 120 INDEX 177

Ocypodoidea, 127 Poecilostomatoida, 3, 38 Scyllaridae, 144 Oniscoidea, 100, 104 Pollicipes, 48 Sergestidae, 126 Oostegites, 79, 81, 83, 85, 88, 91, 94, 104, Polyartemia, 11, 12 Sergestoidea, 126, 127, 134 106, 107, 112, 115 Polyartemiella, 12 Setal row, 91, 104, 112, 129 Oplophoridea, 126 Polyartemiidae, 12 Sexual tube, 145 Oral pyramid, 55 Polyphemidae, 20 Siphonostomatoida, 3, 38 Ostracoda, 3, 23-27,25, 26 Porcellanidae, 145 Spelaeogriphacea, 3, 87- 88, 89 Ovisac, 9, 12, 120 Portunidae, 152 Squillidae, 64 Oxyrhyncha, 127, 152 Portunoidea, 127 Statocyst, 71, 75, 82, 96, 129, 148 Oxystomata, 127, 152 Potamoidea, 127 Stenopodidae, 126 Precoxa, 24, 66 Stenopodidea, 126, 132-133,126, 134 Pagurapseudes, 92, 94 Preischium, 71, 81 Stenopus, 133 Paguristes, 148 Preoral sting, 43 S. hispidus, 133 Paguroidea, 126, 145, 148 Preparatory stage, 94 Stomatopoda, 3, 63- 69, 65, 68 Palaemonoidea, 126 Presoma, 44 Stygocaridacea, 3, 74— 75, 76 Palinura, 126,139, 143-144 Preungual process, 148 Stygocaris, 74, 75 Palinuridae, 144 Procaridoidea, 126 j Stylocheiron, 118, 120 Palinuridea, 144 Psalidopodoidea, 126 Stylodactyloidea, 126 Palinuroidea, 126, 144 Pseudoeuphausia, 120 Supernumerary segment, 148 Pandalidae, 136 Pseudorostrum, 90 Supra anal plate, 9, 30 Pandaloidea, 126 Pylochelidae, 148 Syncarida, 3, 70 Pandalus, 136 Parabathynellidae, 77 Raninoidea, 127 Tanaidacea, 3, 91-95, 93 Paranaspides, 71 Raptorial claw, 66 Tanais, 94 Paranebalia, 59, 60, 62 Reproductive system Telsonic comb, 6, 28 Parastacoidea, 126, 138 Amphipoda, 113 Tessarabrachion, 120 Parastygocaris, 74, 75 Anaspidacea, 73 Thalassinoidea, 126, 145, 148 Pars ampullaris, 67, 71 Anostraca, 15 Thaumastochelidae, 126 Parthenopoidea, 127 Astacidea, 141 Thelycum, 129, 138 Pasiphaeoidea, 126 Brachyura, 156 Thermosbaena, 84, 85 Peltogastrella, 56 Cirripedia, 51 Thermosbaenacea, 3, 84— 87, 86 Penaeidae, 126, 129, 136 Cladocera, 22 Thermosbaenidae, 85 Penaeoidea, 126, 127, 134 Copepoda, 36- 37 Thoracica, 3, 46- 54 Penaeus, 127, 131 Dendrobranchiata, 131 Thysanoessa, 120 Penicillae, 75 Euphausiacea, 122 Thysanopoda, 118, 120 Penis (penes), 12, 27, 48, 51, 52, 66, 88, Isopoda, 105 Toothed furrow, 30 112, 113, 114, 115 Leptostraca, 62 Triops, 8, 9 Peracarida, 3, 79 Mysidacea, 83 Trophi, 48, 52 Petasma, 66, 75, 129 Ostracoda, 27 Phreatoicidea, 98- 99, 100, 104 Paguroidea, 150 Urosome, 34, 112, 115 Phyllocarida, 3, 59 Stomatopoda, 67 Physetocaridoidea, 126 Rhizocephala, 3, 56— 58, 57 Valvifera, 95-96, 104 Pinnotheroidea, 127 Rostral plate, 60, 64, 75 Verruca, 51 Pleocyemata, 126, 132, 138 Verrucomorpha, 51, 53 Pleotelson, 66, 77, 85, 90, 94, 104, 106 Sandersiella, 4, 5 Pleuron (pleura), 6, 81, 88, 129, 134, 138 Sarsostraca, 3 Xanthoidea, 127 Podocopa, 3, 23-27 Scalpellidae, 48 X-organ, 15 LOS ANGELES COUNTY MUSEUM OF NATURAL HISTORY PROPERTY OF REGINA WETZER