SHALLOW-WATER AND TERRESTRIAL BRACHYURAN CRABS OF ROATAN AND THE SWAN ISLANDS, HONDURAS ,„„.„,> CRUSTACEA LIP"'

DONALD E. KEITH JJJM^HSONLAHJ '1

KEITH, DONALD E. 1985 12 30. Shallow-water and terrestrial brachyuran crabs of Roatan ant SAR5IA the Swan Islands, Honduras. - Sarsia 70:251-278. Bergen. ISSN 0036-4827.

There are relatively few published accounts of Brachuyra from the northwestern Caribbean and none dealing specifically with the Bay Islands and Swan Islands off mainland Honduras. Brachyura were collected from shallow-water and terrestrial habitats at the Swan Islands from April 1973 through June 1976 and at Roatan and other Bay Islands from May 1975 through March 1979. Sixty-seven species are reported including 32 from the Swan Islands and 50 from Roatan. Of these, 21 species were not previously reported from northwestern Caribbean waters. One new species of Potamocarcinus was discovered in a freshwater stream at Port Royal, Roatan. Species diversity was greater at Roatan resulting primarily from the much greater diversity of habitats.

Donald E. Keith, Department of Biological Sciences, Tarleton State University, Tarleton Station, Stephenville, Texas 76402, U.S.A.

Swan. There is no source of freshwater. One INTRODUCTION brackish-water pool is located near Buffalo Point. There are few published accounts of Brachyura from Drinking-water on the Islands is obtained from the western Caribbean and none dealing specifically cisterns constructed by the US. Weather Service. with the Bay Islands or Swan Islands, Honduras. There are no mangroves or mud flats as found on the Major taxonomic publications on west Indian Brach- Bay Islands. The low habitat diversity results in a yura include papers by CHACE & HOBBS (1969) and low diversity of crab species as compared to the Bay RATHBUN (1933). Other broad resources of major Islands. Principal habitat types at the Swan Islands taxonomic significance include four monographs by include coral reefs, Thalassia flats with loose coral RATHBUN (1918, 1925, 1930, 1937) on crabs of rubble, rocky shores, sand beaches, and wooded America and papers by WILLIAMS (1965), GUINOT terrestrial areas. Of the 32 species collected from the (1970), and GARTH (1978). POWERS' (1977) catalog Swan Islands, none was limited to the islands and and bibliography of crabs of the Gulf of Mexico also most ranged throughout the Caribbean. contains Caribbean species. Taxonomic nomencla- The Bay Islands include 65 cays and 8 islands of ture in this study follows primarily that of RATHBUN which Roatan is the largest. Besides Roatan, Saint except where noted. GUINOT (1970) made major Helene, Morat, and Barbaret were also visited. revisions of genera in the family Xanthidae. These Roatan, however, was most extensively collected revisions have been included, and RATHBUN'S older during the study. This island is approximately 30 km nomenclature also cited. GARTH (1978) used Gui- long and has 4 km maximum width. It is the not's revised names as subgenera of the older emergent crest of a narrow ridge which runs along genera. the south side of the Bartlett Trough. An account of The Swan Islands and Bay Islands are located in the geology of the island is given by MCBIRNEY & the northwestern Caribbean off the coast of BASS (1969). Geographical information may be mainland Honduras. The Swan Islands are situated found in DAVIDSON (1974). approximately 160 km northeast of the Honduran Unlike the Swan Islands, Roatan has an extreme coast and the Bay Islands about 50 km north of the diversity of habitats including coral reefs, Thalassia mainland (Fig. 1). flats, cobble and sand beaches, rocky shores, man- Three islands oonstitute the Swan Islands chain: grove swamps, mud flats, estuarine bays, freshwater Great Swan, Little Swan, and Booby Cay. Fringing streams, and wooded terrestrial areas. The increased reefs are developed around the perimeter of the habitat diversity results in an increased crab diversi- islands with the most extensive reef growth occurr- ty. Fifty crab species were identified from Roatan. ing along the northern shores. This paper provides a convenient means for Habitat diversity at the Swan Islands is consider- recognizing most of the Bay Islands and Swan ably less than at the Bay Islands. Maximum eleva- Islands species and includes information important tion on Great Swan is only 14 m and 20 m on Little to the geographic distribution of many Caribbean

251 Dromia erythropus (GEORGE EDWARDS, 1771) Fig. 2A. Distinguishing features. Carapace pilose, wi- der than long with posterior lateral margins converg- ing. Front small and tridentate. Four prominent anterior lateral teeth. Range. From Bermuda and the Bahamas, throughout the Gulf of Mexico and Caribbean to Sao Paulo, Brazil. Habitat. Hard substrates including corals, shells, and rocks. Reported by FEEDER (1973) to be most common at depths of less than 46 m, but known to 364 m from some areas. Remarks. The dorsal surface of the carapace is characteristically covered with sponges or compound ascidians. A single specimen was collected at Fort Cay near the water's edge where it presumably was washed in from the nearby reef.

Dromidia antillensis STIMPSON, 1858 Fig 2B. Distinguishing features. Carapace pubescent, species. Distinguishing features are given for each longer than braod with posterior lateral margins species as an aid for recognizing differences between subparallel. Frontal region longitudinally grooved related crabs and are not intended to be species along middle; front strongly deflexed and 5-toothed descriptions. including supraorbital teeth. Lateral margins of carapace 4-toothed and deflected toward corner of MATERIAL AND METHODS buccal areas. Brachyuran collections were made from terrestrial and Range. From Bermuda, North Carolina, Bahamas shallow-water habitats at the Swan Islands during April and Gulf of Mexico, throughout the Caribbean to 1973, July 1974. January 1975 and June 1976. Crabs were Espirito Santo, Brazil. collected by hand during the day. Night collections were Habitat. Reported from hard bottoms such as also made intertidally and in various terrestrial habitats using lights and nets. Collections from coral reef habitats rock, shells, and coral at depths from shore to 331 m. were made with the aid of SCUBA equipment. Some coral Collected at Roatan in shallow water from Thalassia heads were returned to the lab and pulverized to obtain flats. associated crab species. Shallow-water sediments were Remarks. The dorsal surface is commonly covered screened using a No. 60 standard sieve and the undersur- faces of coral rubble were carefully examined for crabs. by a sponge or compound ascidian. Similar techniques were used to collect brachyurans at Roatan and other Honduran Bay islands with the addition of dredging. Extensive collections were made along the Subsection Oxystomata southern shores of Roatan during March 1979. Less Family Calappidae extensive collections were made at Saint Helene and Barbaret. Collections of brachyurans made by Dr. J. C. Calappa angusta A. MILNE EDWARDS, 1880 Britton, Texas Christian University, during May 1975, May Fig. 2C. 1976, and March 1977 were also examined. May 1975 collections included some crabs taken by dredging in Distinguishing features. Carapace slightly wid- approximately 35 m of water off Roatan. er than long with anterolateral margins finely Drawings of crabs included in this study were made by granulate, with somewhat larger granules at inter- tracing projected transparencies of each species. Fine vals; front bilobed. Tooth at posterolateral angle of details were added by microscopic examination. carapace is largest followed anteriorly by four smaller teeth diminishing in size, and posteriorly by SYSTEMATIC ACCOUNT one small and one or two very minute teeth. Orbit Section Brachyura completely separated from antennular cavity. Cheli- peds with upper margin of palm bearing six to eight Subsection Dromiacea teeth and lateral margins with coarse granules Family Dromiidae interspersed with tubercles.

252 Range. North Carolina, Florida, Gulf of Mexico, and sand bottoms from depths from 13 to 274 m. Yucatan to Grenada and Barbados. Specimens were collected at Roatan by dredging Habitat. Reported from coral, broken shell, gravel over Thalassia flats.

253 Calappa gallus (HERBST, 1803) Habitat. Reported from coral, sand, gravel and Fig. 2D. shell bottoms from depths of 10 to 329 m. At Roatar Distinguishing features.Carapace width slight- this species was collected from dredges over a ly exceeds length with anterior two-thirds covered sand-flat area "at 10 m. with tubercles; posterior one-third bears short gra- Iliacantha sparsa STIMPSON, 1871 nulated ridges. A deep sulcus occurs between the Fig. 3A. gastric and hepatic regions. Longitudinal furrows Distinguishing features. Carapace globular, separate the cardiac and branchial regions. Postero- longer than broad with three spines at posterior end lateral expansions well developed with six teeth, which are widely separated and divergent. Median three anterior and two posterior to posterolateral spine longest and all have upturned tips. Spine also tooth. Prominent rostrum with four subequal blunt present on subhepatic margin. Chelipeds with slen- teeth. der palms. Fingers and palm subequal in length. Range. From Bermuda and Florida Keys, Range. Northwest of Tortugas, Puerto Rico, Bar- throughout the Caribbean to Bahia, Brazil. GARTH bados to Bahia, Brazil. Roatan collections extend (1978) reports this species to occur also in the the distribution to the northwestern Caribbean. eastern Atlantic and Indo-West Pacific but not in the Habitat. Reported from gravel and shell bottoms, eastern Pacific Ocean. on calcareous algae, corals, and sponges at depths Habitat. Occurs on hard substrates including reef from 23 to 73 m. Collected off Bay Islands by flats, coral, sand, shell, and rocks from shallow trawling at approximately 35 m. water to 218 m. Individuals were collected at Roatan from cobble beaches in shallow water. Subsection Oxyrhyncha Family Majidae Cycloes bairdii STIMPSON, 1860 Subfamily Acanthonychinae Fig. 2E. Epialtus dilatatus A. MILNE EDWARDS, 1878 Distinguishing features. Carapace subcircular, Fig. 3B. slightly broader than long, and without a posterolat- Distinguishing features. Eyes without true eral expansion. Surface granulate or tuberculate orbits and with short movable eyestalks; preorbital with tubercles partially arranged in longitudinal tooth distinct. Carapace subpentagonal, nearly rows. Anterolateral margins with beaded edge and smooth, with hepatic lobe longer than branchial five to six small teeth behind the orbit. Posterolat- lobe. Rostrum bilobed. Chelipeds of moderate size eral angle with sharp spine. Carapace widens anterior with four longitudinal crests; fingers short and stout. to spine. Range. North Carolina, Bahamas, throughout the Range. North Carolina, Bermuda, Bahamas, Flori- Caribbean to St. Thomas and the Virgin Islands. da, throughout Caribbean to Barbados in the wes- Habitat. Coral, sand, and broken shell bottoms tern Atlantic and from Cape San Lucas to Equador from depths of 5 to 22 m. Collected at Great Swan in the eastern Pacific. Island from coral rubble in shallow water. Habitat. Reported from coral, rock, sand and shell Remarks. WILLIAMS (1965) points out several bottoms from depths of 3 to 229 m. Collected at variations in carapace shape. The hepatic expansions Roatan by dredging at approximately 35 m. may vary and the rostrum may vary from triangular to suboblong in shape.

Family Leucosiidae Epialtus longirostris STIMPSON, 1860 Iliacantha intermedia MIERS, 1886 Fig. 3C. Fig. 2F. Distinguishing features. Appearance is similar Distinguishing features. Carapace globular to E. dilatatus except the rostrum is narrow and and coarsely granulate. No distinct spine on subhep- truncate. Carapace sides deeply bilobed and nearly atic margin. Three spines at posterior extremity with as wide across the hepatic as branchial regions. marginal spines short, flattened, and triangular. Cheliped with cylindrical arm. Palm long and nar- Chelipeds slender with fingers less than half as long row; fingers thick. as palm and incurved at tip. Range. Western Florida, Cuba, Jamaica, St. Tho- Range. From North Carolina, Florida, Virgin mas and Virgin Islands to northeast Brazil. Roatan Islands, Venezuela to Bahia, Brazil. Roatan collec- collections extend the distribution to the northwest- tions extend the distribution to the northwestern ern Caribbean. Caribbean. Habitat. Collected at Roatan from bottom samples

254 slightly arcuate in dorsal view. She called this form Stenorhynchus seticornis (HERBST, 1788) Epialtus longirostrus forma portoricensis. This form Fig. 3F. was known only from Perto Rican waters. The Distinguishing features. Carapace smooth and specimen from Lime Key showed rostral features somewhat arrow-shaped with a long slender rostrum like those of Rathbun's forma portoricensis. bearing small lateral spines and often exceeding the carapace length. Legs and chelipeds long and slen- Subfamily Inachinae der with small spines; palm considerably longer than fingers. Podochela macrodera STIMPSON, 1860 Range. Cape Hatteras, North Carolina, Bermuda, Fig. 3D. Bahamas, Gulf of Mexico, Caribbean to Santa Distinguishing features. Carapace somewhat Catarina, Brazil. triangular, rostrum short, thick, subtriangular and Habitat. Coral, rock, sand, shell and pebble not hollow beneath. Basal segment of antenna with bottoms, wharf pilings, and sponges. Depth ranges longitudinal sulcus. Propodus of last two legs much from shallow water to 1489 m. This species was longer than dactylus and slightly curved. common at Roatan in crevices and under coral Range. Bahamas, Florida, throughout the Caribbe- ledges at about 5 m. an to Brazil. Remarks. The growth and biology of this species Habitat. On coral, sand, gravel, under rocks, and has been studied by HARTNOLL (1965) and BARR on sponges. Depth usually ranges from shallow (1971, 1975), and commensal relationships with a water to 53 m. COELHO (1971) reported this species Florida anemone have been studied by HERRNKIND, from 53 m off Brazil. Collected from Roatan by STANTON, and CONKLIN (1976). dredging over Thalassia flats. Subfamily Mithracinae Podochela riisei STIMPSON, 1860. Macrocoe}oma trispinosum nodipes (DESBONNE & Fig. 3E. SCHRAMM, 1867) Distinguishing features. Carapace subtriangu- Fig. 4A. lar with a tubercle projecting downward from the Distinguishing features. Basal antennal seg- hepatic region. Rostrum short, hood-shaped and ment broad with spine near inner angle visible hollow beneath. Mature males with narrow gape dorsally. Orbits project beyond general outline of betwen fingers. Walking legs slender with last pair carapace. Carapace subtriangular with middorsal about one and one-half times the carapace length. region elevated and bearing four rounded tubercles. Dactyli of second legs short, propodus two and one Posterolateral angles terminate in broad spines with half times or more the dactylus length. margins continuous with that of the carapace. The Range. North Carolina, Bermuda, Bahamas, rostrum bears horns which are usually subparallel at throughout Caribbean to Rio de Janeiro, Brazil. the base, but great variability is shown in length, Habitat. Reported from corals, sand, shell and direction, and curvature. The subspecies M. t. gravel bottoms; also from algal beds, grasses, and nodipes differs from the typical form, M. trispino- among Sargassum. Depth ranges from shallow water sum (LATREILLE) by having posterolateral projec- to 90 m. At Roatan, this species was collected on the tions which are broad and blunt rather than long and coral Agaricia tenuifolia DANA at a depth of about 5 acute. The subspecies also shows much less constric- m. Individuals had bits of algae attached to the body tion behind the orbits than the typical form. and legs. Range. North Carolina, Bermudas, Florida, Yuca- Remarks. POWERS (1977) cites observations of tan, West Indies to Cape Sao Roque, Brazil. these crabs being covered with other organisms such Habitat. Reported from coral, sand, rock, and as bryozoans, ascidians, and red algae. A closely shell bottoms and from grassy areas, dock pilings, related species, P. sidrteyi RATHBUN, has been mangrove roots, and masses of floating Sargassum. reported from the Gulf of Mexico, Yucatan, and Depth ranges from shore to 48 m. Individuals were northwest coast of Cuba. It's general appearance is collected at Great Swan Island in shallow water near similar to P. riisei but can be distinguished by the the shoreline on the undersides of loose coral rubble shorter and more curved dactyli on the last three in Thalassia beds. walking legs. The dactylus of P. sidneyi is more than one half the length of the propodus. Macrocoeloma diplacanthum (STIMPSON, 1860) Fig. 4B. Distinguishing features. Carapace subtriangu-

5 N 1

Fig. 3. A. Iliacantha sparsa, female. B. Epialtus dilatatus, male. C. Epialtus longirostris, female. D. Podochela macrodera, female. E. Podochela riisei, male. F. Stenorhvnchus seticornis, female.

taken in less than 1 m in an area of interspersed Remarks. RATHBUN (1925) recognized two forms Thalassia, sand, and coral rubble. Depth ranges of the species. Besides the typical E. longirostris, she reported by POWERS (1977) are from 3 m off Cuba to recognized a form in which the rostrum differed 54 m off Brazil. slightly, being a little wider and less thick and the tip

6 Fig. 4. A. Macrocoeloma trispinosum nodipes, female. B. Macrocoeloma diplacanthum, male. C. Microphrys bicornutus, male. D. Mithrax coryphe, male. E. Mithrax forceps, female. F. Mithrax ruber, male. denticulate and gaping. Walking legs spinose and Swan Island collections extend the distribution to covered with hair. the northwestern Caribbean. Range. Northern Cuba to Barbados, Curacao. Habitat. On coral, sand, shell, grass, and mud

7 lar with middorsal regions elevated and bearing four Coast of South America from Trinidad to Sao Paulo, conical tubercles near center of carapace; a smaller Brazil. Roatan and Swan Island collections extend spine-like tubercle is located near the posterior the distribution to the northwestern Caribbean. medial edge. Posterolateral processes bifid, one Habitat. Littoral rocks, sand, and algae, coral spine above the other, with a web-like lamina crevices, sponges, broken shell, grass, and mud between. Rostrum long with slender, acute horns bottoms. Depth ranges from shallow water to 55 m. which diverge slightly. A very abundant species around the Swan Islands, Range. Florida, Bahamas, West Indies and Carib- particularly under coral rubble in Thalassia beds and bean. in crevices of the coral Agaricia agaricites (LINNAE- Habitat. Sand, shallow reefs from shallow water to US). M. coryphe was also collected from shallow 24 m. Individuals were collected at Roatan from a cobble beaches around Roatan and other Honduran shallow sand bottom interspersed with coral rubble Bay Islands. and clumps of Thalassia. Remarks. M. coryphe was the dominant littoral spider crab at Great Swan Island but not at Roatan. Microphrys bicornutus (LATREILLE, 1825) Similar habitats at Roatan contained a much greater Fig. 4C. crab diversity and the most common spider crab was Distinguishing features. Carapace subtriangu- Mithrax sculptus. lar, tuberculate, and moderately hairy with a sharp marginal spine at posterolateral angle. A row of four small tubercles arches upward on intestinal region. Mithrax (Mithraculus) forceps (A. MILNE EDWARDS, Rostral horns long and divergent distally. Eyes are 1875) small. A long spine occurs at the anterolateral Fig. 4E. margin of the basal antennal segment which is visible Distinguishing features. Carapace slightly wid- dorsally. Chelipeds spotted, palm slightly enlarged, er than long and deeply sculptured in young individ- fingers hollowed at tip. Arm with three or four uals. Anterolateral margins with four teeth (not tubercles. rounded lobes) separated by rounded sinuses. First Range. From North Carolina, Bermuda, West tooth is usually shortest, other teeth turned forward Indies to Florianapolis, Brazil. Reported by BRATT- at tip. Three brancial sulci run diagonally across STROM (1980) from Colombia and Panama (pers. carapace from first, second, and fourth sinuses. commn) Collections from Roatan and the Swan Rostrum with V-shaped median notch. Merus of Islands extend the distribution to the northwestern chelipeds with two anterior spines followed by five Caribbean. spines or spine-like tubercles on posterior margin. Habitat. Common on coral reefs from very shallow Anterior margin with two teeth. Carpus with small water to 30 m. Also, on rocks, sand, shell fragments, spine on inner margin. Palm smooth, fingers gaping sponges, Thalassia beds, mud and mangrove roots. in males with hollowed tips. Walking legs spiny and This crab was very common on Great Swan Island hairy. where it was abundant on the undersides of loose Range. Bermuda, North Carolina, Florida, Gulf of coral rubble in shallow Thalassia beds. It was also Mexico, Antilles, to Rio de Janeiro, Brazil. very common in shallow rubble areas around the Habitat. Under stones and in crevices of coral Bay Islands. rubble, on rocky shores, sand, shell and grass Remarks. The crab is often covered with sponges, bottoms, on sponges. Reported at depths from anemones, hydroids, algae or other foreign objects intertidal to 90 m. At Roatan, this species was making it very inconspicuous in its habitat. collected from under large pieces of coral rubble and from sponges taken from 30 m. Mithrax (Mithraculus) coryphe (HERBST, 1801) Fig. 4D. Distinguishing features. Carapace nodose with Mithrax (Mithraculus) ruber (STIMPSON, 1871) smooth, oblique, branchial sulci, and broader than Fig. 4F. long. Anterolateral margins behind orbital angle Distinguishing features. Carapace with bear three prominent rounded lobes. Rostrum very smooth, shallow, branchial sulci and broader than small. Chelipeds strong, palm broad, fingers deflex- long. Three anterolateral lobes behind orbit blunt or ed and gaping with crenulated spoon tips; wrist angular in older individuals but last one sharp or nodose. spine-like in young specimens. Rounded prominen- Range. Bahamas, Florida, Antilles to Barbados, ces occur on branchial regions. Chelipeds strong, Caribbean coast of Panama, Colombia, Curacao. palm with parallel margins, carpus smooth, fingers

8 Fig. 5. A. Mithrax sculptus, male. B. Mithrax acuticornis, i C. Mithrax hispidus, male. D. Mithrax pleuracanth- us, male. E. Mithrax verrucosus, male. F. Thoe puella , female. ly equals width, oblique branchial sulci absent. separated by a deep sinus. Anterolateral margins Carapace covered with flattened, closely set gran- with eight spines occurring in pairs united at bases ules. Cervical suture deep. Rostral horns short and with the anterior spines being smaller. Posterolateral

9 bottoms. Also reported from sponges. Occurs at U-shaped notch between. Anterolateral border with depths from shallow water to 46 m. This species was four spine-like teeth. The species is distinguished by common on living coral around the Swan Islands, the one small tooth located just dorsal to the particularly in crevices of Agaricia agaricites and posterolateral margin and by not having more than individuals were collected at Roatan from the coral three tubercles on the posterolateral region. Cheli- Agaricia tenuifolia. peds large, merus with four or five spines on upper margin and two on inner margin. Carpus and palm Mithrax (Mithraculus) sculptus (LAMARCK, 1818) smooth; fingers with spooned tips. Fig. 5A. Range. Delaware Bay, South Carolina, Bahamas, Distinguishing features. Carapace broader Florida, Texas, Jamaica, Curacao to Sao Paulo, than long with posterior two-thirds nodose. Branchi- Brazil. al areas with oblique sulci. Anterolateral margins Habitat. Holes in littoral rocks, sand, shell and with four rounded lobes never angular or spine-like. stone bottoms; on sponges and occasionally sea Chelipeds large in male. Merus bears two spine-like grass. Reported from shallow water to depths of 65 tubercles in front. Carpus smooth and round. Fin- m. At Roatan this species was dominant on the coral gers gaping with tooth near base and spooned tips. Agaricia tenuifolia collected at depths of 7 m. Color usually greenish and persists in alcohol. Remarks. The collection consisted only of young Range. Bahamas, Florida, throughout the Carib- individuals. They differed from RATHBUN'S (1925) bean to the Abrolhos Islands, Brazil. description by having rostral horns which were not Habitat. On corals, under stones, on sand, shell, significantly wider than the notch which separates and grass bottoms from shallow water to 55 m. This them, and in the development of the anterolateral was the dominant spider crab at Roatan, found spines which were smaller than in RATHBUN'S fig- predominantly under coral rubble in shallow water, ures. and on sand flats with clumps of Thalassia and scattered rubble. Mithrax (Mithrax) pleuracanthus STIMPSON, 1871

Remarks. RATHBUN (1925) reported two males and Fig. 5D. one female from the Swan Islands in 1887. However, Distinguishing features. Carapace slightly this species was not collected at the Swan Islands broader than long and tuberculate, without oblique during this study. branchial sulci. Front wide, rostral horns short (usually shorter than in M. hispidus). Four protuber- Mithrax (Mithrax) acuticornis STIMPSON, 1871 ances on anterolateral border; last is spine-like, Fig. 5B. others tuberculated. Two tubercles, arranged trans- Distinguishing features. Carapace without versely, lead inward from last anterolateral spine. smooth oblique branchial sulci and longer than wide. Chelipeds large, merus with five small spines or Surface covered with spines, particularly in cardiac, tubercles on posterior border and scattered tubercles branchial, and intestinal regions. Rostral horns taper on upper margin. Walking legs dentate and hairy. to point and are variable in length. Four large Fingers with slight gape and spooned tips. anterolateral spines (not counting post orbital) and Range. Bermuda, North Carolina, Florida, Baha- one posterolateral. First spine double; the last three mas, Yucatan, Antilles to Curacao. have a smaller spine in front. Chelipeds of male Habitat. Sand, rock, gravel, shell, and mud bot- about as stout as walking legs; palm not armed toms, from shallow water to 51 m. Reported by above with spines. PEARSE (1934) from sponges. Collected at Roatan by Range. East and west Florida coasts, Yucatan, dredging over a sand flat. * Antilles to Santos, Brazil. Remarks. Closely remembers M. hispidus and M. Habitat. Corals, sand, mud, shell, and rock bot- caribbaeus. It can usually be distinguished from M. toms from 3 to 33 m. This species was collected from hispidus by having a tubercle rather than a spine just living coral heads around the Swan Islands. The above the posterolateral margin and shorter rostral corals were at depths of 3 to 7 m. horns. M. caribbaeus is distinguished by two parallel rows of tubercles and spines which run almost Mithrax (Mithrax) hispidus (HERBST, 1790) transversely across the posterolateral region. Fig. 5C. Distinguishing features. Carapace without ob- Mithrax (Mithrax) verrucosus H. MILNE EDVARDS, lique branchial sulci and broader than long. Low 1832 rounded prominences occur toward outer margins of Fig. 5E. the branchial regions. Rostral horns short with Distinguishing features. Carapace length near-

10 ,5 mm 1 1(5 mm 1 Fig. 6. A. Pitho aculeata, male. B. Pitho mirabilis, male. C. Callinectes exasperatus, male. D. Callinectes marginatus, male. E. Portunus sebae, male. F. Portumts spinicarpus, female. ly. Lateral spine is less than 2.5 times length of walking legs. Their tips are divergent. Gonopod preceding tooth. Gonopods of adult male very short, differences are the easiest means of distinguishing C. falling far short of the suture between the ventral marginatus from C. exasperatus. thoracic segments bearing the first and second Range. North Carolina, Bermuda, Bahamas,

262 spine also present. Chelipeds with six sharp spines Habitat. Sand, shell, coral, grass, and mud bot- on the posterior margin of merus, and four on the toms in shallow water. At Great Swan Island, inner margin. Carpus spinose; palm unarmed, elon- individuals were common in the shallows under coral gate; fingers gaping with large tooth near middle of rubble surrounded by Thalassia. At Roatan speci- gape. Walking legs hairy with spines on merus and mens were collected in shallow water from coral carpus. rubble, from dredges taken in mangrove channels, Range. South Carolina, Bahamas, Florida, Swan and by dredging over Thalassia and sand flats at 10 Islands, Antilles, Curacao to Fernando de Noronha, m. Brazil. Habitat. Reported from shallow water in rock Pitho mirabilis (HERBST, 1794) crevices, and on the coral Pontes porites (PALLAS) at Fig. 6B. Curacao by PEARSE (1932). At Great Swan Island Distinguishing features. Carapace length and the species was collected from living coral (Agaricia width about equal and finely granulated. Granules agaricites) at about 4 m. Individuals were collected with coarse hairs on margins of gastric lobes. Five at Roatan from cobble beaches, under coral rubble, anterolateral teeth which are acute and separated to and in rock crevices. their bases. Chelipeds slender in both sexes. Range. Bahamas, Florida Keys to Guadeloupe. Thoe puella STIMPSON, I860 Swan Island collections extend the distribution to Fig. 5F. the northwestern Caribbean. Distinguishing features. Carapace longer than Habitat. Reported from rock and coral bottoms in wide with anterolateral margins nearly straight; shallow water. The species was collected from Great dorsal surface with lobes and covered with clusters Swan Island under loose coral rubble in shallow of granules. Rostral horns very short and separated water. by fissure; antennal flagella setose. Merus and carpus of chelipeds granulated, with line of tubercles Subsection Brachyrhyncha on upper margin of merus. Palm smooth, fingers Family Portunidae gaping with shallow tooth near base in males. Callinectes exasperatus (GERSTAEKER, 1856) Walking legs with posterior laminate crests on merus Fig. 6C. longer and wider than anterior crests, their upper margins being concave. Distinguishing features.Carapace convex with central trapezoidal (metagastric) area almost half as Range. Bahamas, Florida through West Indies to long as anterior width. Front with four rounded Curacao. Swan Island collections extend the range teeth, lateral pair most advanced. Anterolateral to the northwestern Caribbean. margins arcuate, teeth broad and curved forward. Habitat. Corals and littoral rocks in shallow water. Lateral spine less than twice the length of the At Great Swan Island, individuals were collected in preceding tooth. Gonopods of adult male reach shallow water under loose coral rubble surrounded approximately to suture between ventral thoracic by Thalassia. segments bearing the first and second walking legs, their tips curving mesially. Subfamily Ophthalmiinae Range. Bermuda, Bahamas, Southern Florida, Pitho aculeata (GIBBES, 1850) Yucatan to Panama and Venezuela; Antilles to Fig. 6A. Santa Catariria, Brazil. Distinguishing features. Carapace longer than Habitat. Marine and estuarine water, mangrove broad with outer lobe expanded laterally. Width swamps, and around river mouths from shore to 75 between outer orbital angles two-thirds entire width. m. A very common species ifi the bay, and mangrove Anterolateral margins with five teeth, the second swamps at Roatan. Individuals were also collected in and third teeth joined at their base. Fourth and fifth mangroves around St. Helena Island. teeth are small. First movable segment of antenna is much wider than long. Chelipeds of adult male Callinectes marginatus (A. MILNE EDWARDS, 1861) strong, fingers with wide gape, and tooth near Fig. 6D. proximal end of dactyl. Distinguishing features. Carapace with central Range. Bahamas, Florida Keys throughout the trapezoidal (metagastric) area almost half as long as West Indies to Curacao. Collections from Roatan anterior width. Frontal teeth four, medial pair small. and the Swan Islands extend the distribution to the Anterolateral border slightly arched but much less northwestern Caribbean. than in C. exasperatus, teeth curve slightly anterior-

12 A B

Fig. 7. A. Portunus ventralis, female. B. Potamocarcinus sp. (new species), female. C. Carpilius corallinus, male. D. Cataleptodius floridanus, male. E. Chlorodiella longimana, male. F. Domecia acanthophora, female. fingers. In males the dark color is continued a short and Swan Island collections extend the distribution distance on the palm. to the northwestern Caribbean. Range. From Bermuda and the Bahamas, Habitat. Reported on coral, Sargassum, sponges, throughout the Antilles to Sao Paulo, Brazil. Roatan sand, mud, and grass bottoms at depths from

13 southern Florida, Gulf of Mexico, Yucatan to sandy bottom with patches of Thalassia and scat- Panama, Antilles to Sao Paulo, Brazil. Senegal to tered coral rubble. Angola in the eastern Atlantic. Habitat. Marine and estuarine, mangrove swamps, Family Pseudothelphusidae around river mouths, sand and mud flats, grass flats, tide pools, and oyster bars. Depth generally ranges Potamocarcinus sp. New species from littoral to 15 m. The species was collected from Fig. 7B. both Swan Islands and Roatan in shallow rocky A new species of freshwater crab was collected at areas and Thalassia flats. night from a spring-fed stream at Port Royal, Roatan. It was identified as an undescribed species Portunus sebae (H. MILNE EDWARDS, 1834) of Potamocarcinus by Dr. Alfred Smalley (personal Fig. 6E. communication). The species has not yet been Distinguishing features. The species can be described. Systematics of freshwater crabs is some- easily recognized by two large round reddish spots what unsettled at the present time. RATHBUN (1905) on the carapace above each posterolateral margin has published the most recent monograph of the and the dark tips on all of the spines except those of group. BOTT (1955) split the family Potamidae into the front and inner orbits. Palm of chela bears a two groups, the Pseudothelphusidae and the Tricho- longitudinal carina just below upper margin. dactylidae. PRETZMANN (1965), BOTT (1968), and Swimming legs with sharp spine at base. SMALLEY (1970) have suggested other schemes for Range. From Bermuda, Florida Keys and Gulf of classification. Most freshwater crabs have restricted Mexico to Brazil. ranges as a result of inhabiting lakes, streams, and Habitat. Sand, gravel, rocky and mud substrates. caves. Also reported from coral and broken shell bottoms. Several juvenile specimens and one large adult This species was abundant at Roatan and was were collected at Roatan. Unfortunately, the large collected frequently from bays and around dock male was dead and in poor condition. The specimens pilings. were sent to Dr. Smalley for description.

Portunus spinicarpus (STIMPSON, 1871) Family Xanthidae Fig. 6F. Carpilius corallinus (HERBST, 1783) Distinguishing features. The most distinguish- Fig. 7C. ing feature is the inner spine on the carpus of the Distinguishing features. Carapace very broad, chela which is longer than the palm. Merus bears 4 convex, and smooth, with strongly converging poste- to 5 curved spines in front and a single distal spine rolateral borders. Front deflexed almost vertically behind. and has three lobes. Legs compressed; chelipeds Range. From North Carolina and Bahamas, Gulf unequal, with two prehensile teeth on immovable of Mexico and Cuba to Sao Paulo, Brazil. finger of large chela and a single tooth on movable Habitat. Open marine waters from c. 9 to 540 m. finger. Color pale brick red with yellowish markings; Substrates include sand, gravel, mud, coral, and fingers with dark brown tips. broken shell. This species was collected off Roatan Range. From Bermuda and the Bahamas, Gulf of during a five-minute bottom trawl haul at about Mexico, throughout the Antilles to Pernambuco, 50 m. Brazil. Habitat. Relatively common in coral reefs on Portunus ventralis (A. MILNE EDWARDS, 1879) sandy and cobble beaches where it lives in burrows. Fig. 7A. Reported at depths from 2 to 46 m. At Roatan Distinguishing features. Absence of spine at specimens were collected from cobble beaches in extremity of outer margin of arm. Sharp spine at shallow water. distal third of upper margin of palm. Front with four blunt teeth of approximately equal length and rounded at their tips. Cataleptodius floridanus (GIBBES, 1850) Range. From Georgia, along the Florida coast, Fig. 7D. Gulf of Mexico. Cuba, Virgin Islands to Rio de Distinguishing features. Carapace suboval Janeiro, Brazil. with five anterolateral teeth (including outer orbital Habitat. Reported from sandy substrates and tide angle); posterior tooth directed obliquely forward. pools. Specimens were collected at Roatan from a Front double-edged in middle. Chelipeds with dark

14 shallow water to 33 m. Also found in tide pools and ing orbital angle). Front four-lobed with small under stones on cobble beaches. This species was median notch. Chelipeds stout and only slightly very common at the Swan Islands, Roatan, and unequal; fingers dark and spoon-tipped; legs spinu- other Bay Islands. Most commonly collected from lose and hairy. Variations in cheliped and rostral the undersides of coral rubble in cobble-beach areas characteristics have been noted in specimens from and Thalassia flats. Colombia, Florida, and Swan Islands by Dr. John Remarks. Listed as Leptodius floridanus by RATH- Garth of the Allan Hancock Foundation (pers. BUN (1930). Changed to Cataleptodius floridanus bv commn). GUINOT (1968). Range. Reported from the Bahamas, Florida Keys, throughout the Greater Antilles, and Colombia. Clorodiella longimana (H. MILNE EDWARDS, 1834) Roatan and Swan Island collections extend the range Fig. 7E. to the northwestern Caribbean. Distinguishing features. Carapace hexagonal Habitat. Coral reefs and shallow-water rocky and microscopically granulate; five anterolateral substrates. Specimens were collected at Swan teeth (including orbital angle) widely separated and Islands from a stabilized sand flat. The species was acute in younger specimens. In older individuals very abundant at Roatan in Thalassia flats among only the last two teeth are acute. Epigastric regions the coral rubble. with smooth, transverse ridges; front double-edged. Remarks. Identified by RATHBUN (1930) as Phy- Chelipeds extremely long; two-thirds of arm project modius maculatus. GUINOT (1969) transferred the beyond carapace. Fingers black with color of im- species to Etisus maculatus making it the sole movable finger continued on palm. representative of this genus in the Atlantic. Other Range. From the Bahamas and east Florida coast, members of the genus are Indo-Pacific. throughout the Antilles to Barbados and Curacao. Swan Island collections extend the distribution to Micropanope nuttingi (RATHBUN, 1898) the northwestern Caribbean. Fig. 8B. Habitat. Reported from coral, sponges, and rocky Distinguishing features. Carapace hexagonal substrates from 5 to 154 m. At Great Swan Island to suboval, with second lateral tooth united with the individuals were collected from the coral Agaricia orbital tooth which is small. The three remaining agaricites. anterolateral teeth are sharp. Front bilobed and separated by a small V-shaped sinus. Chelipeds are Domecia acanthophora (DESBONNE & SCHRAMM, heavy with spines on upper margin. Wrist and palm 1867) tuberculate. Fingers dark and without a gape. Fig. 7F. Range. From Cape Hatteras, North Carolina, Distinguishing features. Carapace narrows Florida, and Gulf of Mexico throughout the Carib- posteriorly, anterolateral border with four to six bean to Rio Grande, Brazil. acute dark spines and several dark spines medial to Habitat. Coral, rock, sand and broken shell anterolateral borders. Chelipeds also armed with bottoms, cobble beaches, and sponges from shore to black spines. 183 m. Several individuals were collected at Great Range. From North Carolina, Florida, Texas coast, Swan Island from living brain coral. Most were Panama and Colombia, throughout the Antilles to juvenile. Pernambuco, Brazil. Remarks. According to GUINOT (1967, 1968) only Habitat. Reported from coral, sponges, and cob- the type species M. sculptipes STIMPSON, and M. lo- ble beaches. The species was collected at Great bifrons A. MILNE EDWARDS belong to the genus Swan Island from living coral heads, particularly Micropanope. The others are divided among four Agaricia. Individuals were collected at Roatan from new genera. The paper, howe\er, omits many of the cobble beaches in shallow water. Depth ranges have species formerly included in Micropanope. LEMAITRE been reported from low tide to 146 m. (1984) and WILLIAMS (1978) point out, regarding REMARKS. Reported as Domecia hispida by RATH- Micropanope pusilla A. MILNE EDWARDS, that be- BUN (1930). Changed to Domecia acanthophora cause of insufficient knowledge of intrageneric varia- acanthophora by GUINOT (1964). tion of gonopods in the Xanthidae a new genus for M. pusilla is not justified at this time. J. Garth (pers. Etisus maculatus (STIMPSON, 1860) commn) suggested that the genus Micropanope be Fig. 8A. retained for M. nuttingi and M. spinipes since they Distinguishing features. Carapace hexagonal had not been assigned to one of Guinofs new and areolated with four anterolateral teeth (exclud- genera.

265 Micropanope spinipes (A. MILNE EDWARDS, 1880) collections extend the range to the northwestern Fig. 8C. Caribbean. Distinguishing features. Carapace hexagonal, Habitat. Reported from rocks, sand, shell and dorsal surface rough to granular. Five anterolateral gravel bottoms; among sponges, ascidians, seaweed, teeth; first pair small but not fused with orbital and pier pilings. Depth ranges from shore to 18 m. tooth. Third and fourth teeth larger than others. All individuals collected at Roatan were taken from Palms smooth, fingers dark but dark color does not canals through the mangrove swamps on mud extend onto palms. bottoms. Range. From Bermuda Islands, Florida Keys, Gulf Remarks. RATHBUN (1930) lists two environmental of Mexico, Curacao to the Abrolhos Islands, Brazil. forms in addition to the typical. Roatan individuals Habitat. Reported from coral, sand substrate, and conform most closely to Rathbun's forma serrata sponges at depths from low tide to 55 m. Specimens since the third to fifth lateral teeth are prominent were collected at Roatan from the coral Agaricia and widely separated. The Roatan species closely tenuifolia at approximately 8 m. matches young males from Caledonia Bay, Atlantic Panama in the Allan Hancock Foundation collec- Neopanope packardii? (KINGSLEY, 1879) tions but according to Dr. John Garth (pers. Distinguishing features. Carapace subhexago- commn) neither the Caledonia Bay species or the nal, high and sloping on sides and narrow posteri- Roatan species agree completely with larger males orly. Five anterolateral teeth with fusion of first and from Trinidad and Florida. second; last three are spiniform with tips widely separated. Front advanced and arcuate. Finger of Paractaea rufopunctata nodosa (STIMPSON, 1860) large cheliped with large basal tooth. Fig. 8E. Range. Florida, Bahamas, west to Louisiana and Distinguishing features. Carapace bears high south to the northern coast of Cuba. nodules which are granulose and widely separated. Habitat. Reported from a variety of substrates Depressions between nodules are hairy. Most ante- including rocks, coral, sand, gravel, mud, shell, and rior nodules not fused with frontal margin. Edges of grass beds from low tide to 74 m. At Roatan, one legs fringed with coarse hair. Fingers hollowed at specimen was taken from a Thalassia flat and one tips. from a bottom trawl at 10 m. Range. From North Carolina, Bahamas, and Flori- Remarks. The two individuals found at Roatan da, throughout Antilles, Curacao to Cape Frio, were both juvenile females making identification Brazil. Also reported from Ascension Island in the extremely difficult. Dr. John Garth of the Allan south Atlantic. Collections from the Swan Islands Hancock Foundation compared the individuals with extend the range to the northwestern Caribbean. like size and sex of other Neopanope packardii and Habitat. Reported from coral, stone, broken shell concluded that they were possibly the same. Because and sand bottoms, and from sponges at depths of 5 of the lack of mature specimens and uncertainty of to 212 m. Specimens were collected at Great Swan the identification this species is not illustrated. See Island near shore from the undersides of coral RATHBUN (1930, plate 168) for photograph. If the rubble in Thalassia beds. individuals are N. packardii it extends the range of Remarks. Listed as Actaea rufopunctata nodosa by this species to the northwestern Caribbean. RATHBUN, 1930; changed to Paractaea rufopunctata nodosa by GUINOT (1969). Panopeus occidentalis SAUSSURE, 1857 Fig. 8D. Paraliornera dispar (STIMPSON, 1871) Distinguishing features. Carapace wider than Fig. 8F. long and crossed by broken, transverse, granulated Distinguishing features. Carapace smooth and lines. Five anterolateral teeth with first and second shiny and about three-fourths as long as broad. separated by shallow sinus. Third and fourth teeth Anterolateral margins without spines or teeth; re- larger with convex posterior margins; fifth tooth gions not well defined. Front about one-third width shorter. Carapace narrows posteriorly; posterolater- of carapace. Chelipeds unequal, but more equal al border longer than anterolateral border. Fingers than in P. longimana. Minor chela about half the pointed, dark color of immovable finger of cheliped width of major. Fingers dark with color extending not continued on palm. slightly on palm. Range. From North Carolina, Bermuda, Bahamas Range. From Bermuda, Florida Keys, Antilles, through Florida and the Antilles to Santa Catarina, Curacao to Colombia. Swan Island collections ex- Brazil. Previous reports also from Panama. Roatan tend the range to the northwestern Carribbean.

266 2mm '

Fig. 8. A. Etisus maculatus, male. B. Micropanope nuttingi, female. C. Micropanope spinipes, female. D. Panopeus occidentalis, female. E.Paractaea rufopunctata nodosa, female. F. Paraliomera dispar, female.

Habitat. Reported from coral reefs, sand, shell and Swan Island the species was collected from living mud bottoms, and rocky and grassy areas at depths coral heads at depths of approximately 6 m. ranging from shallow water to 154 m. At Great

267 Paraliomera longimana (A. MILNE EDWARDS, 1865) Platyactaea setigera (H. MILNE EDWARDS, 1834) Fig. 9A. Fig. 9D. Distinguishing features. Carapace about three- Distinguishing features. Carapace wide, uni- fifths as long as broad with gastric region plainly formly granulose and covered with short hairs. visible. Anterolateral margins are faintly four-lobed. Nodules low and separated by narrow furrows. Front with transverse fringe of hair. Chelipeds Margins of the carapace are lobiform and the unequal in both sexes and smooth. Palms elongate posterolateral borders concave. Chelipeds hairy and with larger one heavy and smaller one very slender. granulose with black fingers. Legs smooth and hairy. Range. Bermuda, Bahamas, Florida Keys, and Dry Range. From Florida Keys to Barbados. Reported Tortugas; throughout Antilles to Trinidad; Curacao also from Veracruz, Mexico and Curacao. and Caribbean coast of Colombia. Collections from Habitat. Rocky and grassy substrates and coral the Swan Islands extend the distribution to the reefs from shallow water to 154 m. At Roatan northwestern part of the Caribbean. individuals were identified from substrate samples Habitat. Reported in shallow water from coral collected on a Thalassia flat. reefs and rocky substrates. This species was collec- ted at Great Swan Island from the undersides of Pilumnus holosericus (RATHBUN, 1898) coral rubble in Thalassia beds, and from crevices in Fig. 9B. the coral Agaricia.

Distinguishing features. Carapace covered Remark's. Identified as Actaea setigera by RATH- with hairs which more or less conceal the dorsal BUN (1930). The species was placed in the genus tubercles or areoles. Front separated by a median Platyactaea by GUINOT (1967). V-shaped sinus. Anterolateral margins with four teeth; first tooth (outer orbital) is smaller. Wrist and Platypodiella spectabilis (HERBST, 1794) palm pubescent like carapace. Fingers short and Fig. 9E. dark. Legs covered with short hairs and fringed with Distinguishing features. Carapace granulate, coarse hairs. relatively smooth with low lobules; front deflexed. Range. From the Bahamas south to Trinidad and Anterolateral margins thin, crested and slightly Curacao. Swan Island collections extend the range lobed; posterolateral borders concave. Upper bor- to the northwestern Caribbean. ders of chelipeds and legs crested; fingers pointed Habitat. Reported from shallow water under and dark. stones in rocky and coral reef areas. At Great Swan Range. Distributed throughout the Caribbean and Island, individuals were collected in rock crevices Gulf of Mexico from Bermuda and the Bahamas as and under coral rubble in Thalassia beds near shore. far south as Fernando de Noronha, Brazil. Habitat. Reported from coral reefs, under stones Pilumnus longleyi RATHBUN, 1930 and occasionally from sponges from 4 to 13 m. Fig. 9C. Specimens were collected at Great Swan Island from Distinguishing features. Carapace covered living coral heads. with hairs which are shorter posteriorly. Anterolate- Remarks. This species was identified by RATHBUN ral margins with six spines, the third and fourth (1930) as Platypodia spectabilis. GUINOT (1967) being joined near base. Chelipeds and legs are long transferred it to the new genus Platypodiella. and hairy. Manus of both chelae rough and hairy on outer margins. RATHBUN (1930) points out the ease Xantho denticulatus WHITE, 1848 of confusing this species with P. caribaeus DESBON- Fig. 9F. NE & SCHRAMM and P. sayi RATHBUN because of the Distinguishing features. Carapace more or less similarities in general appearance. P. longleyi, how- oval and anterolateral borders bear nine to ten teeth ever, can be distinguished by the six spines on the (not including orbital angle) and continue behind anterolateral margins and the greater number of widest part of the carapace. Front with small, spines on the dorsal surface of the anterolateral shallow median notch. Upper wrist surface and palm region. rugose. Inner margin of wrist bears two teeth. Range . Reported from the Bahamas, Florida Keys, Fingers pointed and dark. and Dry Tortugas. Collections from Roatan extend Range . Widely distributed, ranging from Bermuda, the range to the northwestern Caribbean. Florida Keys, throughout Antilles to Trinidad and Habitat. Rocky and coralline bottoms in shallow from Panama to the Abrolhos Islands, Brazil with water. At Roatan, this species was collected from range extensions to the Gulf of Guinea and west rocky shores. coast of Africa.

268 Fig. 9. A. Paraliomera longimana, male. B. Pilumnus holosericus, male. C. Pilumnus longleyi, male. D. Platyactaea setigera, male. E. Platypodiella spectabilis, male. F. Xantho denticulatus, male.

Habitat. Reported from coral reefs, under stones, Thalassia. Specimens were also obtained from brain and in tide pools. The species was abundant at Great coral and coralline sand. Swan Island under coral rubble and in beds of Remarks. The species was originally described

269 under the genus Xantho but was transferred by sharp tooth at the posterior orbital angle; front RATHBUN (1930) to Xanthodius. Most carcinologists, sharply deflexed. Third maxillipeds gape widely however, continue to use the older genus. GUINOT exposing the mandibles and they do not bear an (1968) raised some questions about differences oblique hairy ridge. Fossa between second and third between the South American and African species. walking legs fringed with hair. Chelipeds subequal with inner margins of merus expanded and denticu- Family Pinnotheridae late; palm of chela tuberculate dorsally. The species is often recognized by its bright colors which vary Dissodactylus borradailei RATHBUN, 1918 from yellow-brown to red. There is often purple Fig. 10A. markings and spots along the lateral margins and Distinguishing features. Carapace smooth and upper parts of legs. very convex with concave posterolateral surfaces. A Range. From Bermuda and Bahamas, Gulf of ridge runs dorsally from the lateral angles halfway to Mexico, throughout Antilles to Rio de Janeiro, median line. Outer maxillary palp 3-jointed. Brazil. Reported also from the eastern Atlanic from Range. Reported from Florida and Jamaica. Roa- Senegal to northern Angola. tan collections extend their distribution to the Habitat. Frequents mangrove swamps and is often northwestern Caribbean. seen among mangrove roots near the water's edge. Habitat. RATHBUN (1918) reported individuals This species was extremely common in the mangrove from fine white sand taken in dredges from 49 to 55 swamps at Roatan and other Honduran Bay Islands. m. At Roatan individuals were collected from The crab has also been reported from wet muddy dredges through Thalassia flats. They were found to marine shores, along inlets and estuaries. HARTNOLL be commensal with the sea biscuit, Meoma ventrico- (1965) observed small individuals sheltering under sa (LAMARCK). Most of the urchins had several crabs leaves and debris beneath the Jamaican mangroves. clinging to the spines on their oral side. G. cruentata does not construct burrows but may take refuge in the burrows of other crabs such as Family Grapsidae Ucides cordatus. Subfamily Grapsinae

Geograpsus lividus (H. MILNE EDWARDS, 1837) Grapsus grapsus (LINNAEUS, 1758) Fig. 10B. Fig. 10D. Distinguishing features. Carapace subrectan- Distinguishing features. Carapace subcircular gular with lateral margins well defined. Transverse in adults with distinct transverse striations in bran- striations occur laterally and anteriorly. Upper chial regions; front sharply deflexed. A sharp tooth margin of front bears four tubercles; edge nearly occurs just posterior to outer orbital angle. Merus of straight. Tooth located on the lateral margins poste- legs with transverse striations. Chelipeds short and rior to the orbital angle. Chelipeds subequal; wrist subequal with longitudinal ribs on sides of palms. with tooth on inner angle. Inner arm margin Upper margin of palms tuberculate; wrist with spine expanded and denticulate proximally. Palm tubercu- on inner margin. Fingers with spooned tips. late dorsally; fingers pointed. Range. This species has an extremely wide distribu- Range. A widely distributed species ranging from tion from the Bermudas, Florida, throughout the Bermuda, Florida Keys throughout Antilles to Trini- Antilles to Estado de Pernambuco, Brazil. It is also dad and from the Caribbean coast of Colombia to distributed throughout the Caribbean from Curacao Sao Paulo, Brazil. It also occurs in the eastern to the Gulf of Mexico. The range extends to the Pacific from Baja California to Chile with extensions eastern Atlantic from southern Portugal to northern to the Clipperton, Galapagos, and Hawaiian Islands. Angola and in the eastern Pacific from central Baja Habitat. Frequents rocky shores and cobble beach- California to central Chile and to the Galapagos and es from midlittoral to supralittoral zones. Speci- Clipperton Islands. mens were collected at Great Swan Island from the Habitat. A frequent inhabitant of littoral and undersides of coral rubble in shallow water and on supralittoral rocky shores. Also, reported from rocks in the splash zone. cobble beaches and on cliff walls. The species was extremely common at the Swan Islands and Bay Goniopsis cruentata (Latreille, 1803) Islands from rocky areas where they were particular- Fig. 10C. ly abundant in the splash zones. At the Swan Distinguishing features. Carapace subquadrate Islands, they were also abundant on drifts of with distinct striations both laterally and anteriorly; Thalassia at the shoreline.

270 A B

Fig. 10. A. Dissodactylus borradailei, female. B. Geograpsus lividus, male. C. Goniopsis cruentata, male. D. Grapsus grapsus, male. E. Pachygrapsus transversus, male. F. Percnon gibbesi, male.

about one-third broader than long, and covered by Pachygrapsus transversus (GIBBES, 1850) transverse striations anteriorly and laterally. Sides Fig. 10E. convergent posteriorly and armed with a sharp tooth Distinguishing features. Carapace trapezoidal, behind the outer orbital tooth. Front sinuose and 271 slightly more than half the greatest width. Chelipeds Antilles to Pernambuco, Brazil; and in the eastern subequal and bear a laminate expansion on the inner Atlantic from the Azores to Angola. margin of the arm. Inner wrist margin with blunt Habitat. Found intertidally among rocks, on jetties tooth; fingers with spooned tips. and in tide pools. Individuals were collected from Range. Extremely wide range from North Caro- Great Swan Island under coral rubble and at Roatan lina, Bermuda, Bahamas, Florida, throughout the from a cobble-beach habitat in shallow water. Antilles and along the Brazilian coast to Uruguay. Also occurs throughout the Caribbean and Gulf of Mexico. The range extends to the eastern Atlantic Subfamily Sesarminae from the Mediterranean Sea to Angola, and in the eastern Pacific from central California to Peru. The Aratus pisonii (II. MILNE EDWARDS, 1853) species is also reported from the Galapagos and Fig. 11B. Easter Islands. Distinguishing features. Carapace trapezoidal Habitat. Common beneath stones near the tide with lateral margins striated and converging posteri- line. Also, reported from rocks, pilings, and whar- orly. Front strongly deflexed and about seven-tenths ves. Occasional reports on mangrove roots and sand as wide as carapace. Antennae excluded from orbit. beaches. Individuals were collected at Roatan under Merus of third maxillipeds with oblique hairy ridge rocks at the shoreline and at Great Swan Island and gaping, exposing mandibles. Chelipeds sub- under coral rubble in beds of Thalassia. equal, palm with dorsal tubercles; fingers with spooned tips. Walking legs broad and flattened with subterminal tooth on anterior margin. Dactyls of Percnon gibbesi (H. MILNE EDWARDS, 1853) legs very short. Fig. 10F Range . From the Bahamas and Florida throughout Distinguishing features. Carapace longer than the Antilles to Estado de Sao Paulo, Brazil. In the wide, ovate and flattened, with four anterolateral eastern Pacific from Nicaragua to Peru. spines including the outer orbital spine; dorsal Habitat. Very common among mangroves and tubercles flattened. Front spinose and narrow be- along shores of estuaries. Also reported from rock tween antennules. Arm and wrist of chelipeds piles and wharves. It is present on most of the spinose, palm smooth, fingers short and hollowed at Caribbean Islands, especially in mangrove swamps tips. Merus of walking legs with spines on upper where it actively climbs among the mangroves close margins. to their seaward margins. At Roatan this species was Range. Widely distributed from North Carolina, most commonly observed inhabiting the roots of the Bermuda, Bahamas, Florida, throughout Antilles to red mangrove, Rhizophom mangle LINNAEUS, above Brazil. Range extends to the eastern Atlantic from and just below the water line. Larger crabs were the Azores to the Cape of Good Hope and to the reported by HARTNOLL (1965) to climb to consid- eastern Pacific from Baja California to Chile. erable heights to reach mangrove leaves which were Habitat. Found in lower rocky intertidal areas and their principal diet. The crabs are active both during on rock and pebble bottoms. The species was the days and night and when pursued, escape by collected at Great Swan Island from coral rubble in climbing or dropping off into the water where they the splash zone at night. swim to the nearest root and ascend.

Plagusia depressa (FABRICIUS, 1775) Fig. 11A. Cyclograpsus integer H. MILNE EDWARDS, 1837 Distinguishing features. Carapace subcircular, Fig. 11C. slightly wider than long, tuberculate, and partially Distinguishing features. Carapace subrectan- covered with short hairs forming a scale-like pattern. gular, wider than long, and nearly smooth. Lateral Front divided by deep lobes exposing antennules margins smooth and subparallel. Third maxilliped dorsally. Anterolateral borders with four teeth with oblique hairy ridge across merus and gaping including outer orbital angle. Palm of chelipeds with widely, exposing mandibles. Chelipeds subequal, dorsal tubercles arranged in row and separated by fingers pointed. Merus of walking legs with a blunt setose furrows. Merus of walking legs with terminal subterminal tooth on anterior margin. and subterminal spines on anterior border. Last Range. From Bermuda, Florida, and Bahamas, three leg segments fringed with dense hair. throughout the Caribbean and Gulf of Mexico to Range. From North Carolina, Bermuda, Florida, Pernambuco, Brazil. Ranges in the eastern Atlantic Gulf of Mexico, Panama, Colombia, throughout the from Senegal to Zaire.

272 A B

I—r^ 1

Fig. 11. A. Plagusia depressa, male. B. Aratus pisonii, male. C. Cyclograpsus integer, male. D. Sesarma ricordi, male. E. Sesarma roberti, male. F. Cardisoma guanhumi, male.

Habitat. Reported from splash areas along rocky Roatan individuals were collected from under small beaches. Burrows in marshy marine areas. At stones along the shoreline.

273 Sesarma ricordi H. MILNE EDWARDS, 1853 There has been some confusion regarding the Fig. 1 ID. taxonomic status of S. roberti and S. angustipes Distinguishing features. Carapace subquadra- DANA. HARTNOLL (1965) used S. angustipes as a te, lightly striate laterally, and slightly wider posteri- senior synonym for S. roberti after comparing a male orly. No tooth present posterior to outer orbital syntype of S. roberti with specimens of S. angustipes angle. Front strongly deflexed at postfrontal lobes used by RATHBUN (1918). CHASE & HOBBS (1969) and broadens distally. Distance between outer orbit- agreed with HARTNOLL'S finding regarding RATH- al angles subequal to maximum carapace width. BUN'S species but was not convinced that S. roberti Third maxillipeds with oblique hairy ridge on surface was a junior synonym. CHASE & HOBBS maintained of merus, and gaping widely. Chelipeds subequal, that until a specimen of the Caribbean species with fingers with wide aperture at base, pointed and distinct frontal and pleopodal features is found on holllowed out. the northeastern and eastern coasts of South Ameri- Range. From Bermuda, Bahamas, and Florida, ca the two should be considered distinct species. throughout Antilles to Trinidad; Curacao, Yucatan. ABEI.E (1972) in his summary of the taxonomic status Habitat. Among rocks and debris along the sho- of several sesarmids agreed with CHASE & HOBBS. reline from the littoral zone to about 50 m inland. Also, reported from the edges of mangrove swamps Family Gecarcinidae and in grassy areas above sandy beaches. HARTNOLL Cardisoma guanhumi LATREILLE, 1825 (1965) reported that some individuals occasion- Fig. 1 IF. ally traveled up to 100 m from the sea, and Distinguishing features. Carapace broad, oval, sometimes shared burrows with Cardisoma guan- and smooth. Distance between outer orbital angles humi. S. ricordi was extremely common at Roatan, more than half the carapace width. Third maxilliped particularly around mangrove swamps and under gaping, exopod visible and hairy. Dactyls of walking diiftwood along the shoreline. They were also found legs with four rows of spines. Color varies from pale under rocks and short distances inland under logs blue or lavender to ashy grey or tan; chelipeds and leaf litter. smooth and unequal in both sexes, but much more so in males; fingers stout and meet only at tips. Sesarma roberti H. MILNE EDWARDS, 1853. Range. Includes the Gulf of Mexico, eastern Fig. HE. Mexico to Colombia and the western Atlantic from Distinguishing features. Carapace subquad- the Bermuda Islands to Sao Paulo, Brazil. rate, striate laterally, with subparallel margins which Habitat. This is one of the most common semiter- converge slightly anteriorly. No tooth present poste- restrial land crabs occurring throughout the Carib- rior to outer orbital angle. Distance between outer bean. C. guanhumi inhabits low-lying coastal areas, orbital angles slightly less than maximum carapace mangrove swamps, river banks, and drainage dit- width. Front strongly deflexed at postfrontal lobes ches. It is primarily nocturnal except in heavily and does not broaden distally. Third maxillipeds shaded areas or on very cloudy days. This species with oblique hairy ridge on surface of merus, and was collected at Roatan from low-lying forested gaping widely. Chelipeds subequal; fingers spoon- areas and mangrove mud flats. Younger crabs tipped. inhabited burrows nearer the shore overlapping with Range. From Cuba throughout Antilles to Venezu- Uca rapax. Larger members of the species tended to ela and from Vera Cruz, Mexico to Nicaragua. inhabit supralittoral areas. At Great Swan Island Habitat. Frequents a wide range of brackish and this species was not as abundant as the other freshwater habitats including rivers, streams, bays, gecarcinid species but was frequently observed along and mangrove swamps. Reported to burrow along the shore and short distances inland at night. muddy slopes of river banks. This species was Remarks. This crab is harvested commercially as collected at Roatan from a freshwater stream at Port food on some Caribbean Islands such as Puerto Royal. They were found submerged among rocks in Rico. POWERS (1977) reports this crab to be an the shallow flowing spring about 100 m from the agricultural pest in southern Florida because of seashore. Individuals were also collected along the extensive burrowing in the fields and consumption of muddy shores of some mangrove swamps. plant shoots. Remarks. Sesarma roberti closely resembles S. ri- cordi but can easily be distinguished by the front. In Gecarcinus lateralis (FREMINVILLE, 1835) S. ricordi the front widens distally, whereas in Fig. 12A. S. roberti the lateral margins of the front are Distinguishing features. Carapace widest at subparallel. anterolateral angles. Distance between outer orbital

274 30 mm Fig. 12. A. Gecarcinus lateralis, male. B. Gecarcinus ruricola, female. C. Ocypode quadrata, male. D. Uca burgersi, male. E. Uca rapax, male. F. Ucides cordatus, male.

angles half or slightly less than half greatest carapace males but usually subequal in young individuals. width. Merus of third maxilliped bears shallow Dactyls of legs armed with four rows of spines. marginal notch distally. Chelipeds unequal in older Carapace with dark reddish-brown central area.

275 Range. Widespread throughout the Caribbean and Caribbean. Ranges from Florida along the Atlantic Gulf of Mexico, southeast Florida, Bahamas, and coast north to Rhode Island and as far south as Santa Bermuda Islands. Catarina, Brazil. Habitat. Burrows in dry, sandy areas further Habitat. Constructs burrows in supralittoral sand inshore than Cardisoma. It has been reported from beaches from the high-water mark to back-dune dune ridges and back dune areas of Florida and from areas. Older individuals have been observed to wooded areas under logs and leaf litter. At Great construct burrows further inland than young crabs. Swan Island, it was very abundant in wooded areas. The species was a common inhabitant of the Swan The species was also observed at Roatan and Islands and the Bay Islands on sandy beaches. Barbaret from inland wooded areas. G. lateralis is Remarks. This is the only species of the genus most frequently observed during the evening or on Ocypode which has been described from the Gulf of cloudy days after a rain. Mexico and the Caribbean. It was described as Ocypode albicans Bosc. by RATHBUN (1918) and Gecarcinus ruricola (LINNAEUS, 1758) others prior to revision by HOETHUIS (1959). Fig. 12B. Distinguishing features. Carapace with bran- Uca burgersi HOETHUIS, 1967 chial regions greatly swollen. Distance between Fig. 12D. outer orbital angles about two-fifths maximum cara- Distinguishing features. Carapace subtrape- pace width in adults, but may approach half the zoidal to hexagonal and broader than long. Distance carapace width in half-grown individuals. Chelipeds between outer orbital angles equal to maximum subequal with dentate fingers narrowly gaping. carapace width. Chelipeds unequal in males and Merus of third maxilliped rounded distally without a subequal in females. Fingers longer than palm in marginal notch. Dactyls of legs bear six rows of large chela of males and gaping. Second, third, and spines. Color is extremely variable. fourth walking legs bear hairs on outer margins of Range. From the Bahamas, southeast Florida to the carpus and propodus. First pleopod (gonopod) Barbados. Also reported from the Cayman Islands, of male has a laterally directed tip (posterior view), Swan Islands, Nicaragua, and Curacao. the lateral lobe being much wider than medial. Habitat. Frequently observed in low and marshy Cornea of eyestalk occupies about two-fifths of its ground of the savannas of the West Indies. The extensor surface. species was very abundant in the wooded areas at Range. From the Florida east coast and Bahamas, Great Swan Island. It was observed mostly at night throughout Antilles to Trinidad. Curacao and Carib- and in evenings on rainy days. In southeast Florida bean coast of Panama to the eastern coast of the species occasionally occurs among dense popula- Yucatan. Range exends along South America from tions of the smaller gecarcinid, G. lateralis. Venezuela to Rio de Janeiro, Brazil. Remarks. The species is reported to migrate down Habitat. Mud flats around river mouths, marshes, from the hills in some areas of the West Indies in and in mangrove mud flats and thickets. This was the swarms during the rainy season in May. The females only species of Uca found on Great Swan Island. It enter the sea allowing the young to hatch. After a was collected from a flat, near a brackish-water few weeks they migrate back into the hills (GREEN pool. 1963). Uca rapax (SMITH, 1870) Family Ocypodidae Fig. 12E. Distinguishing features. This species is similar Ocypode quadrata (FABRICIUS, 1787) in appearance to U. burgersi but distinguished (CHA- Fig. 12C. SE & HOBBS, 1969) by the cornea and male gonopods. Distinguishing features. Carapace nearly sub- The cornea of U. burgersi typically occupies less quadrate in adults with a fine granular surface. Front than two-fifths of extensor surface of distal eyestalk narrow and deflexed. Distance between outer orbi- segment, whereas in U. rapax it occupies about tal angles nine-tenths carapace width. Specialized two-fifths of the surface. The first denuded pleopod hair-fringed openings occur between the basal joints (gonopod) of U. rapax terminates (posterior view) of the second and third pairs of walking legs in all in two projections with the lateral lobe only slightly members of this family. Chelipeds similar and broader than the medial. In U. burgersi the lateral unequal in both sexes. Palm of largest chela bears a lobe is very broad and the medial lobe is inconspi- stridulating ridge on its medial surface. cuous. Range . Occurs throughout the Gulf of Mexico and Range . Florida east coast and Bahamas throughout

276 the Antilles to Santa Cantarina, Brazil and from ACKNOWLEDGMENTS Mexico and northeast Yucatan to Panama. Appreciation is expressed to Tarleton State University lor Habitat. Mud and sandy mud flats, mud banks of the organized research grant which made this study streams, salt marshes, and edges of mangroves. This possible, and to the Texas Christian University Research Foundation who sponsored trips to the Swan Islands. I species is often sympatric with U. burgersi but thank Mr. Frank Roulstone of the U.S. Weather Service U. rapax usually occurs nearer the open sea and, for his assistance in coordiriating trips to the Swan Islands, according to CRANE (1975), in more saline condi- and Dr. J. C. Britton, Texas Christian University, for making brachyuran collections from the Swan Islands and tions. At Roatan and other Bay Islands the species Roatan available for study. I especially thank Dr. John S. was very abundant in mud flats and near the edge of Garth of the Allan Hancock Foundation, University of mangrove swamps ranging from the shoreward Southern California, for his help in confirming many of the mangrove fringe and extending inland some dis- brachyuran species and for his review of this manuscript. tance. In Jamaica it primarily inhabits the Laguncu- laria and Avicennia zones and was characterized as REFERENCES an upper-swamp species by WARNER (1969). Abele, L.G. 1972. The status of Sesarma angustipes Dana, Remarks. This is a widespread and very abundant 1852, S. trapezium Dana, 1852, and S. miersil species of Uca in the Caribbean. U. rapax was listed Rathbun, 1897 (Crustacea: Decapoda: Grapsidae) in the western Atlantic. - Caribbean Journal of as a subspecies of U. pugnax (SMITH) by RATHBUN Science 12:165-170. (1918) until a revision by TASHIAN & VERNBERG Barr, L. 1971. Observations on the biology of the arrow (1958) separated U. rapax out as a separate species. crab, Stenorhynchus seticornis (Herbst) in Lames- hur Bay, St. John, Virgin Islands. - Pp. 213-220 in: Miller, J., J. van Derwalker & R. Walkers (eds). Ucides cordatus (LINNAEUS, 1763) Scientists-in-the-sea. Department of Interior, Was- Fig. 12F. hington. D.C. Distinguishing features. Carapace of male — 1975. Biology and behavior of the arrow crab, about one and one-fourth times wider than long. Stenorhynchus seticornis (Herbst), in Lameshur Bay, St. John, Virgin Islands. - Pp. 47-56 in: Distance between outer orbital angles no more than Results of the Tektite Program. Bulletin of the two-thirds maximum carapace width. Chelipeds Museum of Natural History, Los Angeles County unequal in both sexes; palm concave along lower 20. margin. Walking legs of male bear long hairs on the Bott, R. 1955. Die Siisswasserkrabben von Afrika (Crust., Decap.) und ihre Stammegeschichte. - Annates du ventral margins; first pair are longer and last pair Musee du Congo Beige. Serie in Quarto, Zoologie shorter than the others. The crab is relatively large 1:209-352. with a bluish-gray carapace reaching up to 90 mm in — 1968. Fluss-Krabben aus dem ostlichen Mittel- width. Females differ by having a narrower cara- Amerika und von dem Grossen Antillen (Crusta- cea, Decapoda). - Senckenbergiana Biologica pace, shorter legs and chelipeds, and fewer hairs on 49:39-49. the legs. Brattstrom, H. 1980. Rocky-shore zonation in the Santa Range. From the Bahamas and northeastern Mexi- Marta area, Colombia. - Sarsia 65:163-226. co, throughout the Caribbean to Santa Catarina, Chace Jr, F A. & H.H. Hobbs Jr. 1969. The freshwater and terrestrial decapod crustaceans of the West Brazil. Indies with special reference to Dominica. - Bulle- Habitat. Found around river mouths and in areas tin, United States National Museum 292:1-258. of standing brackish water. Also among mangrove Coelho, P.A. 1971. Nota previa sobre os Majidae do norte roots and mud flats. At Roatan this species bur- e nordeste do Brasil. - Archivos Museu National Rio de Janeiro 54:137-146. rowed in mangrove mud flats. These areas were also Crane, J. 1975. Fiddler Crabs of the World. Ocypodidae: occupied by Uca rapax, Goniopsus cruentata, and Genus Uca. Princeton Univ. Press, New Jersey. young Cardisoma guanhumi. 736 pp. Remarks. CHACE & HOBBS (1969) transferred Davidson, W.V. 1974. Historical geography of the Bay Islands, Honduras. - Southern Univ. Press, Bir- Ucides from the family Gecarcinidae to Ocypodidae. mingham, Ala. 199 pp. Felder, D.L. 1973. An annotated key to crabs and lobsters (Decapoda, Reptantia) from coastal waters of the northwestern Gulf of Mexico. - Publ. No. LSU- SG-73-02 of the Center for Wetland Resources, Louisiana State University, Baton Rouge. 103 pp. Garth, J.S. 1978. Marine biological investigations in the Bahamas 19. Decapoda Brachyura. - Sarsia. 63:317-333. Green, J. 1963. A biology of Crustacea. H.F. and G. Witherby Ltd., London. 180 pp. Guinot, D. 1964. Les trois especes du genre Domecia (Decapoda, Brachyura): D. hispida Eydoux and

277 Souleyet, D. glabra Alcock, et D. acanthophora Powers, L.W. 1977. A catalog and bibliography to the (Dcsbonne and Schramm). - Crustaceana 7:267- crabs (Brachyura) of the Gulf of Mexico. - Contri- 283. butions to Marine Science, University of Texas, — 1967. Recherches preliminaires sur les groupements Supplement to Vol. 20. 190 pp. naturels chez les Crustaces Decapodes Brachyoures Pretzmann, G. 1965. Vorlaufiger Bericht iiber die Familie III. A propos des affinities des genres Dairoides Pseudothelphusidae. - Anzeiger der Osterreich- Stebbing et Daira de Haan. - Bulletin du Museum ischen Akademie der Wissenschaften 1965:1-10. National d'Histoire Naturelle, Paris 39:540-563. Rathbun, M.J. 1905. Les crabes d'eau douce (Potamon- — 1968. Recherches preliminaires sur les groupements idae), Pt. 2. - Archives du Museum National naturels chez les Crustaces Decapodes Brachyou- d'Historie, Paris, Ser. 4, 7:159-321. res. IV. Observations sur quelques genres de — 1918. The grapsoid crabs of America. - Bulletin, Xanthidae. - Bulletin du Museum National d'Histoi- United States National Museum 97:1-461. re Naturelle, Paris 39:695-727. — 1925. The spider crabs of America. - Bulletin, — 1969. Sur divers Xanthidae notamment sur Actaea United States National Museum 129:1-613. de Haan et Paractaea gen. nov. (Crustacea Decapo- — 1930. The cancroid crabs of America of the families da, Brachyura). - Cahiers du Pacifique 13:223-285. Euryalidae, Portunidae, Atelecyclidae, Cancridae — 1970 (1971). Recherches preliminaires sur les and Xanthidae. - Bulletin, United States National groupements naturels chez les Crustaces Decapodes Museum 152:1-609. Brachyoures VIII. Synthese et bibliographie. - — 1933. Brachyuran crabs of Porto Rico and the Virgin Bulletin du Museum National d'Histoire Naturelle, Islands. - Scientific survey of Porto Rico and the Paris Ser. 2, 42:1063-1090. Virgin Islands 15:1-121. New York Academy of Hartnoll, R.G. 1965. Notes on the marine grapsoid crabs of Sciences. Jamaica. - Proceedings of the Linnean Societv of — 1937. The oxystomatous and allied crabs of Ameri- London 176:113-147. ca. - Bulletin, United States National Museum Herrnkind, W., G. Stanton & E. Conklin 1976. Initial 166:1-278. characterization of the commensal complex associ- Smalley, A.E. 1970. A new genus of freshwater crab from ated with the anemone, Lebrunia danae, at Grand Guatemala, with a key to the middle American Bahama. - Bulletin of Marine Science 26:65-71. genera (Crustacea Decapoda, Pseudothelphus- Holthuis, L.B. 1959. The Crustacea Decapoda of Suriname idae). - American Midland Naturalist 83:96-106. (Dutch Guiana). - Zoologische Verhandelingen, Tashian, R.E. & F.J. Vernberg 1958. The specific distinct- Leiden 44:1-296. ness of the fiddler crabs Uca pugnax (Smith) and Lemaitre, R. 1984. Decapod crustaceans from Cay Sal Uca rapax (Smith) at their zone of overlap in Bank, Bahamas, with notes on their zoogeographic northwestern Florida. - Zoologica 43:89-92. affinities. - Journal of Crustacean Biologv 4:425- Warner, G.F. 1969. The occurrence and distribution of 447. crabs in a Jamaican mangrove swamp. - Journal of McBirney, A.R., & M.N. Bass 1969. Geology of Bay Animal Ecology 38:379-389. Islands, Gulf of Honduras. In: McBirney, A.R. Williams, A.B. 1965. Marine decapod crustaceans of the (ed.) Tectonic relations of northern Central Ameri- Carolinas. - Fishery Bulletin 65:1-298. ca and the western Caribbean - The Bonaca — 1978. Transfer to Pseudomedaeus of the xanthid Expedition. - American Association of Petrolium crab Micropanope distinctus (Rathbun). - Procee- Geologists, Memoir 11:229-243. dings of the Biological Society of Washington Pearse, A.S. 1932. Observations on the parasites and 91:546-557. commensals found associated with crustaceans and fishes at Drv Tortugas, Fla. - Papers, Tortugas Accepted 14 May 1985. Laboratory 28:103-115. — 1934. Inhabitants of certain sponges at Dry Tortu- gas. - Papers, Tortugas Laboratory 28:117—124.

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