Annotated Checklist of Decapod Crustaceans of Atlantic Coastal and Continental Shelf Waters of the United States

Home , Anasimus, Armases cinereum, Carpoporus, Chaceon quinquedens, Clythrocerus, Dyspanopeus

PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON 116(1):96—157. 2003. Annotated checklist of decapod crustaceans of Atlantic coastal and continental shelf waters of the United States

Martha S. Nizinski

National Marine Fisheries Service National Systematics Laboratory, Smithsonian Institution, P.O. Box 37012, NHB, MRC-0153, Washington, D.C. 20013-7012, U.S.A., e-mail: [email protected]

Abstract.—The decapod crustacean assemblage inhabiting estuarine, neritic and continental shelf waters (to 190 m) of the temperate eastern United States is diverse, with 391 species reported from Maine to Cape Canaveral, Florida. Three recognized biogeographic provinces (Boreal in part, Virginian and Car- olinian) are included in this region. The assemblage contains 122 shrimp species (28 penaeids, 2 stenopodids, and 92 carideans), 10 thalassinideans, 8 lobsters, 61 anomurans and 190 brachyurans. Since previous compilation of this fauna, 12 additional species have been described, including four carideans, one callianassid, four anomurans, and three brachyurans. Range extensions into the region have been reported for another five species (Parapenaeus americanus, Scyllarides aequinoctialis, Petrolisthes armatus, Dromia erythropus, Clythro- cerus nitidus). One species, Hemigrapsus sanguineus, has been introduced and become established throughout intertidal environments from southern Maine to northern North Carolina. Six species previously recorded from this region are no longer considered to occur there. Two of these species occur south of the Carolinian biogeographic province, three others are now known to occur only in the Pacific Ocean, and one species previously considered as likely to occur in the region has never actually been recorded there. Scientific nomenclature for all species recorded from the region is updated and referenced. Geographic distributions are summarized for each species incorporating recent published information where available.

The decapod crustacean assemblage m) decapods in this region has advanced. (shrimps, lobsters, and crabs) inhabiting es- Numerous changes in taxonomy and/or sys- tuarine, neritic and continental shelf waters tematic placement of species occurring in (to 190 m) of the temperate eastern United this region have also been published in pa- States is diverse and fairly well known. The pers scattered throughout the literature. most recent comprehensive review of this Objectives of this paper are to re-evaluate fauna (Williams 1984) recognized 342 spe- the taxonomic status and to update nomen- cies of decapods and listed 14 extralimital clature for the 342 nominal species of deca- species (species thought to occur inciden- pods treated by Williams (1984). The list of tally in the region). Williams' (1984) mono- species he presented is updated by inclusion graph is widely used and continues to be of all decapods now known to occur in this the best available reference for decapod region based on recently published infor- crustaceans of the east coast of the United mation. Emendations to Williams' original States. However, in the nearly 20 years list include (1) the addition of species in since publication of Williams (1984), this region described after Williams' pub- knowledge regarding shallow water (<190 lication, (2) adding of species newly re- VOLUME 116, NUMBER 1 97 ported from the region as published range teras, North Carolina, to Cape Canaveral, extensions and introductions, and (3) re- Florida). Williams (1984) commented that moval of species no longer considered to inclusion or exclusion of some species from occur in the region based on published his list was subjective. For example, he re- range revisions. The emended list also in- garded some species as extralimital (i.e., cludes 13 species listed in Williams (1984) their center of distribution was outside the as extralimital, as well as 16 other species region) and did not consider them to be part that Williams (1984) did not treat, but of the assemblage despite the fact that they which previously had been recorded from had been recorded from the region. Wil- this region. liams' treatment of extralimital taxa has proved to be problematic and confusing es- Methods pecially in attempts to summarize decapod crustacean diversity of the region. To alle- This compilation largely follows the clas- viate this confusion, the present work does sification and arrangement of Martin & Da- not consider species as extralimital (sensu vis (2001). Based on other recent system- Williams). Based on available distributional atic studies (cited below), some species and information, the present compilation treats genera have been moved to different fami- species as either occurring in the geograph- lies, and other species have been reassigned ic region and thus included in the list, or as to different genera. Where new systematic not occurring in the region and thus ex- information has necessitated changes, up- cluded. dated nomenclature with relevant references is provided, together with a cross-reference Systematic Account to names used in Williams (1984). Symbols preceding species listings highlight the fol- Order Decapoda Latreille, 1802 lowing changes between Williams (1984) Suborder Dendrobranchiata Bate, 1888 and the present compilation: • denotes new Infraorder Penaeidea Rafinesque, 1815 additions to the decapod assemblage; * de- Superfamily Penaeoidea Rafinesque, 1815 notes changes in nomenclature and/or sys- • Family Aristeidae Wood-Mason, 1891 tematic placement. Additionally, distribu- Remarks.—Perez Farfante & Kensley tional information is reported for each spe- (1997) diagnosed the family and genera, cies that occurs in the region with recently presented a key to the genera based on mor- published range revisions incorporated and phological characters, and illustrated im- referenced. Justification explaining changes portant diagnostic characters for the genera in systematic placement of taxa, together included in their study. with appropriate reference sources, are provided in a remarks section when relevant. • Aristaeomorpha foliacea (Risso, 1827) The region of coverage in the present work is the same as that in Williams (1984), Remarks.—Species diagnosis and figures spanning depths from shallow water to the are provided in Perez Farfante (1988) and 100-fathom (ca. 190 m) depth contour in Rodriguez (1993). This species occurs be- three biogeographic provinces. These prov- tween 170-1300 m (Perez Farfante 1988). inces, defined by marine climatic zones Known range.—Western Atlantic: Mas- (Williams 1984, Engle & Summers 1999), sachusetts to Florida, Gulf of Mexico, Ca- are, from north to south, Boreal (represent- ribbean Sea to Venezuela (Perez Farfante & ed by the region from Maine to north of Kensley 1997), and Brazil (D'lncao 1998); Cape Cod, Massachusetts), Virginian (Cape Eastern Atlantic: Bay of Biscay to Western Cod, Massachusetts, to Cape Hatteras, Sahara; Azores; Madeira; Canary Islands; North Carolina) and Carolinian (Cape Hat- Mediterranean Sea; Indo-West Pacific: 98 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

South Africa to Mozambique; Madagascar; Remarks.—This species occurs in shal- Reunion Island; Maldive Islands; Sri Lan- low water to 366 m (Caetano da Costa et ka; Indonesia; Philippines; Taiwan; Japan; al. 2000), but more frequently at depths of Australia; New Zealand; New Caledonia; less than 60 m (D'Incao 1998). Wallis and Futuna Islands; Fiji (Perez Far- Known range.—Western Atlantic: Cape fante & Kensley 1997). Hatteras, North Carolina, to Florida Keys; southern Gulf of Mexico (off Campeche) Family Penaeidae Rafinesque, 1815 and Yucatan; Caribbean Sea to Rio Grande Remarks.—Dall et al. (1990) provided a do Sul, Brazil; Bermuda (Williams 1984, comprehensive account of many aspects of Perez Farfante & Kensley 1997). penaeid systematics, biology and ecology. Perez Farfante & Kensley (1997) diagnosed ^k Farfantepenaeus duorarum the family and genera, presented a key to (Burkenroad, 1939) the genera based on morphological charac- Penaeus (.Farfantepenaeus) duorarum Bur- ters, and illustrated important diagnostic characters for the genera included in their kenroad.—Williams, 1984:28. study. Former subgenera (.Farfantepenaeus Known range.—Western Atlantic: lower and Litopenaeus) of the more inclusive Chesapeake Bay to Florida; Gulf of Mexico Penaeus were raised to full generic status to tip of Yucatan Peninsula; Bermuda (Wil- by Perez Farfante & Kensley (1997). Re- liams 1984, Perez Farfante & Kensley sults of some molecular phylogenies (Bald- 1997). win et al. 1998, Gusmao et al. 2000) have failed to support elevation of these subgen- * Litopenaeus setiferus (Linnaeus, 1767) era to genera, however, an alternative hypothesis of relationships proposed in the Penaeus (Litopenaeus) setiferus (Linnae- molecular phylogeny of Maggioni et al. us).—Williams, 1984:32. (2001) supported conclusions of Perez Far- Known range.—Western Atlantic: New fante & Kensley (1997) derived from their York to St. Lucie Inlet, Florida; Gulf of analysis of morphological characters. Tra- Mexico to Yucatan (Williams 1984, Perez chypenaeus, as previously defined and as Farfante & Kensley 1997). listed in Williams (1984), was shown to be a complex of five genera (Perez Farfante & Metapenaeopsis goodei (Smith, 1885) Kensley 1997). Only one of these five genera, Rimapenaeus Perez Farfante & Ken- Known range.—Western Atlantic: north- sley, 1997, is represented in the region by east of Cape Lookout, North Carolina, a single species. through the Florida Straits to Alabama; off Cape Catoche, Yucatan; Bahamas through * Farfantepenaeus aztecus (Ives, 1891) the Caribbean Sea and along the coasts of Penaeus (.Farfantepenaeus) aztecus Ives.— Central and South America to Espfrito San- Williams, 1984:24. to, Brazil; Bermuda (Williams 1984, Perez Farfante & Kensley 1997, D'Incao 1998). Known range.—Western Atlantic: Mar- tha's Vineyard, Massachusetts, to Florida; • Parapenaeus americanus Rathbun, Gulf of Mexico to Yucatan (Williams 1984, 1901 Perez Farfante & Kensley 1997). Remarks.—Perez Farfante & Kensley * Farfantepenaeus brasiliensis (1997) record this species from off New (Latreille, 1817) England. This species occurs at 37—412 m Penaeus (Farfantepenaeus) brasiliensis La- (Perez Farfante 1977b, D'Incao 1998). treille.—Williams, 1984:28. Known range.—Western Atlantic: off VOLUME 116, NUMBER 1 99

New England; off Ponte Vedra, Florida; Perez Farfante & Kensley 1997, D'Incao Gulf of Mexico; Bahamas; Cuba; Puerto 1998). Rico; St. Lucia; Belize; Brazil (Rio de Ja- neiro to Rio Grande do Sul); northern Uru- Xiphopenaeus kroyeri (Heller, 1862) guay (Perez Farfante 1977b, Perez Farfante & Kensley 1997, D'Incao 1998). Remarks.—This species occurs in shallow water to 70 m (Caetano da Costa et al. Parapenaeus politus Smith, 1881 2000). Known range.—Western Atlantic: Vir- Remarks.—Parapenaeus politus occurs ginia (Maris 1986) to Rio Grande do Sul, over mud and sandy mud sediments at Brazil, including the Gulf of Mexico and depths of 3—752 m, but is usually found be- Caribbean Sea (Williams 1984, Perez Far- tween 65-275 m (Rodriguez 1993). fante & Kensley 1997, Caetano da Costa et Known range.—Western Atlantic: Mar- al. 2000); Eastern Pacific: Punta Piaxtla, tha's Vineyard, Massachusetts, south Sinaloa, Mexico to Paita, Peru (Perez Far- through Gulf of Mexico, Caribbean Sea to fante & Kensley 1997). French Guiana, and Para, Brazil (Williams 1984, Perez Farfante & Kensley 1997, Family Sicyoniidae Ortmann, 1898 D'Incao 1998). Remarks.—Perez Farfante & Kensley • Penaeopsis serrata Bate, 1881 (1997) provided family and generic diagnoses and illustrations of diagnostic char- Remarks.—Species diagnosis, descrip- acters. tion, illustrations, as well as, color and size information were provided in Perez Farfan- te (1980). Penaeopsis serrata occurs at Sicyonia brevirostris Stimpson, 1871 120—750 m, with maximum concentrations Known range.—Western Atlantic: Nor- occurring between 300-450 m (Perez Far- folk, Virginia, to Florida, through the Gulf fante 1980, Rodriguez 1993). of Mexico; Campeche and Yucatan banks; Known range.—Western Atlantic: New Cuba; Bahamas; Eastern Pacific: off south- Jersey to Gulf of Mexico; Caribbean Sea to ern Mexico (Williams 1984, Perez Farfante French Guiana; southern Brazil; Eastern At- & Kensley 1997). lantic: Portugal to northwest coast of Africa (Perez Farfante 1980, Perez Farfante & Sicyonia burkenroadi Cobb, 1971 Kensley 1997). Known range.—Western Atlantic: Cape :k Rimapenaeus constrictus Lookout, North Carolina, to Florida, (Stimpson, 1871) through the Gulf of Mexico; West Indies; Trachypenaeus constrictus (Stimpson).— Caribbean coasts of Central and South Williams, 1984:38. America to Bahia, Brazil (Williams 1984, Perez Farfante & Kensley 1997). Remarks.—Rimapenaeus constrictus is a new combination proposed by Perez Far- Sicyonia dorsalis Kingsley, 1878 fante & Kensley (1997). This species occurs in shallow water to 127 m (Caetano da Known range.—Western Atlantic: Cape Costa et al. 2000). Hatteras, North Carolina, through the Gulf Known range.—Western Atlantic: Nova of Mexico to Texas; Caribbean coasts of Scotia; Chesapeake Bay to Florida Keys, Central and South America to southern Bra- Gulf of Mexico; Bermuda; Caribbean Sea zil (Williams 1984, Perez Farfante & Ken- to Santa Catarina, Brazil (Williams 1984, sley 1997, D'Incao 1998). 100 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Sicyonia laevigata Stimpson, 1871 • Hadropenaeus affinis (Bouvier, 1906)

Known range.—Western Atlantic: Cape Remarks.—Description, affinities, com- Hatteras, North Carolina, to southern Flor- parisons, and illustrations are included in ida; Gulf of Mexico; West Indies; Carib- Perez Farfante (1977a). This species occurs bean coasts of Mexico, Central America between 165-570 m (Perez Farfante and South America to Rio Grande do Sul, 1977a). Brazil (Williams 1984, Perez Farfante & Known range.—Western Atlantic: North Kensley 1997, Caetano da Costa et al. Carolina to Florida; Gulf of Mexico; Carib- 2000); Eastern Pacific: southern Gulf of bean; Eastern Atlantic: Cape Verde Islands California; Costa Rica; Panama (Perez Far- (Perez Farfante 1977a, Perez Farfante & fante & Kensley 1997). Kensley 1997).

• Hadropenaeus modestus (Smith, 1885) Sicyonia parri (Burkenroad, 1934) Remarks.—Description, affinities, com- Known range.—Western Atlantic: Beau- parisons, and illustrations are included in fort, North Carolina, through the Gulf of Perez Farfante (1977a). This species occurs Mexico; West Indies to Sao Paulo, Brazil at depths of 150-550 m (Perez Farfante (Williams 1984, Perez Farfante & Kensley 1977a). 1997, Caetano da Costa et al. 2000). Known range.—Western Atlantic: Dela- ware to Florida, Gulf of Mexico, Bahamas, Sicyonia stimpsoni Bouvier, 1905 Caribbean, and northern Brazil (Perez Far- fante 1977a, Perez Farfante & Kensley Known range.—Western Atlantic: Cape 1997). Hatteras, North Carolina, to Florida through the Gulf of Mexico; West Indies; Caribbean Mesopenaeus tropicalis (Bouvier, 1905) coasts of Mexico, Central and northern Known range.—Western Atlantic: north- South America to Suriname (Williams east of Cape Lookout, North Carolina, to 1984, Perez Farfante & Kensley 1997). southern Brazil, including Gulf of Mexico (Williams 1984, Perez Farfante & Kensley Sicyonia typica (Boeck, 1864) 1997).

Known range.—Western Atlantic: • * Pleoticus robustus (Smith, 1885) Wrightsville Beach, North Carolina, through the Gulf of Mexico; Cuba through Hymenopenaeus robustus Smith.—Wil- the West Indies; Caribbean coasts of Cen- liams, 1984:484. tral and South America to Rio Grande do Remarks.—Williams (1984) considered Sul, Brazil (Williams 1984, Perez Farfante this species to be extralimital. Perez Farfan- & Kensley 1997, D'Incao 1998, Caetano da te (1977a) concluded that Hymenopenaeus Costa et al. 2000). comprised a complex of genera and con- sequently transferred H. robustus and two Family Solenoceridae Wood-Mason, 1891 other species (H. muelleri and H. stein- dachneri) to the genus Pleoticus Bate, Remarks.—Perez Farfante & Kensley 1888. Description, affinities, comparisons, (1997) diagnosed the family and genera, and illustrations of P. robustus were includ- presented a key to the genera based on mor- ed in Perez Farfante (1977a) and Squires phological characters, and illustrated im- (1990). This species occurs at 70-915 m, portant diagnostic characters for the genera but is most abundant between 250-475 m included in their study. (Perez Farfante 1977a, Rodriguez 1993). VOLUME 116, NUMBER 1 101

Known range.—Western Atlantic: Mas- ered. Lucifer typus is usually found offshore sachusetts to Gulf of Mexico; Caribbean (>180 m bottom depth), but occasionally is Sea to French Guiana (Perez Farfante collected in shallower waters (Bowman and 1977a, 1988; Squires 1990; Perez Farfante McCain 1967). Bowman and McCain & Kensley 1997). (1967) provided information on diagnostic characters. Solenocera atlantidis Burkenroad, 1939 Known range.—Western Atlantic: New- foundland; east coast of United States Known range.—Western Atlantic: North (Maine to Florida); Sargasso Sea; Brazil; Carolina to southern Brazil, including Gulf Eastern Atlantic: Mediterranean Sea; Cape of Mexico and West Indies (Williams 1984, of Good Hope; east coast of South Africa; Perez Farfante & Kensley 1997). East-Cental Pacific: Baja California; Gulf of California to north of 4°; Indo-West Pa- Solenocera necopina Burkenroad, 1939 cific: Bay of Bengal; Philippines; Queens- Known range.—Western Atlantic: off land, Australia (Abele and Kim 1986, Perez Oregon Inlet, North Carolina, to Uruguay, Farfante & Kensley 1997, D'Incao 1998). including Gulf of Mexico and Bahamas (Williams 1984, Perez Farfante & Kensley Family Sergestidae Dana, 1852 1997). Remarks.—Perez Farfante & Kensley Solenocera vioscai Burkenroad, 1934 (1997) provided family and generic diagnoses and illustrations of diagnostic char- Known range.—Western Atlantic: south- acters. east of Cape Lookout, North Carolina, to Florida and Gulf of Mexico (Williams Acetes americanus carolinae Hansen, 1984, Perez Farfante & Kensley 1997). 1933

Superfamily Sergestoidea Dana, 1852 Remarks.—Perez Farfante and Kensley Family Luciferidae De Haan, 1849 (1997) considered subspecific designation valid. Remarks.—Perez Farfante & Kensley Known range.—Western Atlantic: lower (1997) provided a family diagnosis and il- Chesapeake Bay through Gulf of Mexico to lustrated important diagnostic characters. Panama; Suriname; French Guiana (Wil- liams 1984, Perez Farfante and Kensley Lucifer faxoni Borradaile, 1915 1997). Known range.—Western Atlantic: Long Island Sound, New York, to Rio Grande do Suborder Pleocyemata Burkenroad, 1963 Sul, Brazil, including Gulf of Mexico, Ca- Infraorder Stenopodidea Claus, 1872 ribbean Sea, and Bermuda (Williams 1984, Family Stenopodidae Claus, 1872 Perez Farfante & Kensley 1997, D'Incao Stenopus hispidus (Olivier, 1811) 1998); Eastern Atlantic: Senegal; Congo Known range.—Western Atlantic: Ber- (Williams 1984, Perez Farfante & Kensley muda; North Carolina to southern Florida, 1997). through the Gulf of Mexico to Fernando de Noronha and Espirito Santo, Brazil (Wil- • Lucifer typus H. Milne Edwards, 1837 liams 1984, Coelho & Ramos-Porto 1998a); Remarks.—Williams (1984) did not in- Central Pacific: Hawaii; Indo-West Pacific: clude this species in his study because Durban, South Africa; Red Sea; Japan; depth of occurrence was centered beyond western Australia; eastern Australia through the limits of bathymetric range he consid- New Caledonia; New Hebrides; Lord Howe 102 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Island; northern New Zealand to Tuamotu Norway to British Isles; Bay of Biscay; Archipelago (Williams 1984). Mediterranean to Adriatic (Williams 1984, Squires 1990). Stenopus scutellatus Rankin, 1898 Superfamily Bresilioidea Caiman, 1896 Known range.—Western Atlantic: Ber- * Family Disciadidae Rathbun, 1902 muda; South Carolina; Gulf of Mexico to Remarks.—Martin & Davis (2001) rec- Fernando de Noronha and Rio Grande do ognized the family Disciadidae. Species of Norte, Brazil (Williams 1984, Coelho & Discias were previously considered part of Ramos-Porto 1998a). the Bresiliidae (Williams 1984).

Infraorder Caridea Dana, 1852 Discias atlanticus Gurney, 1939 Superfamily Pasiphaeoidea Dana, 1852 Remarks.—Criales & Lemaitre (1997) Family Pasiphaeidae Dana, 1852 reported the discovery of this species in- Remarks.—Subgeneric designations habiting tubes of the polychaete Chaetop- within Leptochela are considered valid by terus variopedatus. This represents the first Holthuis (1993). reported occurrence of this symbiotic rela- tionship for these species. These authors Leptochela (.Leptochela) papulata Chace, also discussed morphological sexual dimor- 1976 phism. Known range.—Western Atlantic: Ber- Known range.—Western Atlantic: muda; Savannah, Georgia; Fort Pierce, Georges Bank, off Massachusetts; North Florida; Gulf of Mexico; Guadeloupe; Co- Carolina to Georgia; eastern Gulf of Mex- lombia; Maranhao and Ceara, Brazil (Ken- ico (Williams 1984). sley 1983, Williams 1984, Criales & Le- maitre 1997, Ramos-Porto & Coelho 1998); Leptochela (.Leptochela) serratorbita Bate, Eastern Atlantic: Cape Verde Islands; Ga- 1888 bon (Williams 1984); Indian Ocean: northern Kenya (Williams 1984); Red Sea; Known range.—Western Atlantic: Beau- Western Pacific: Great Barrier Reef, Aus- fort, North Carolina, to South Carolina; tralia (Kensley 1983). western Gulf of Mexico; Florida Keys to Leeward Islands; Amapa to Pernambuco, • Discias vernbergi Boothe & Heard, and Sao Paulo, Brazil (Williams 1984, Ra- 1987 mos-Porto & Coelho 1998). Remarks.—Boothe & Heard (1987:506) described this species and provided a di- Leptochela (Proboloura) carinata agnosis, illustrations, and size and sexual Ortmann, 1893 maturity information. This species occurs at Known range.—Western Atlantic: 54-74 m (Boothe & Heard 1987). Georges Bank, off Massachusetts; South Known range.—Western Atlantic: Geor- Carolina; Gulf of Mexico, through Baha- gia; eastern Gulf of Mexico (west Florida) mas to Alagoas, Brazil (Williams 1984, Ra- (Boothe & Heard 1987). mos-Porto & Coelho 1998). Superfamily Palaemonoidea Rafinesque, 1815 Pasiphaea multidentata Esmark, 1866 • Family Anchistioididae Borradaile, Known range.—Western Atlantic: south- 1915 east of Greenland to Cape Cod, Massachu- Anchistioides antiguensis (Schmitt, 1924) setts, including Gulf of St. Lawrence and Remarks.—This species was listed under Gulf of Maine; Eastern Atlantic: Iceland; family Palaemonidae, subfamily Pontoni- VOLUME 116, NUMBER 1 103 inae, in Williams (1984). Chace (1992) pro- Macrobrachium acanthurus vided evidence for familial separation of (Wiegmann, 1836) Anchistioides\ Holthuis (1993) also adopted Known range.—Western Atlantic: Neuse this arrangement. River estuary, North Carolina, to Rio Known range.—Western Atlantic: Ber- Grande do Sul, Brazil (Williams 1984, Ra- muda; South Carolina; west Florida through mos-Porto & Coelho 1998). West Indies to Maranhao, Pernambuco, and Alagoas, Brazil (Williams 1984, Ramos- Macrobrachium carcinus (Linnaeus, 1758) Porto & Coelho 1998). Known range.—Western Atlantic: St. Family Gnathophyllidae Dana, 1852 Augustine, St. Johns County and Silver Gnathophyllum modestum Hay, 1917 Glen Springs, Marion County, Florida, to Rio Grande do Sul, Brazil, including Gulf Known range.—Western Atlantic: Beau- of Mexico and Caribbean Sea (Williams fort, North Carolina; Biscayne Bay, Florida 1984, Ramos-Porto & Coelho 1998). (Williams 1984). Macrobrachium ohione (Smith, 1874)

Family Palaemonidae Rafinesque, 1815 Known range.—Western Atlantic: James Subfamily Palaemoninae Rafinesque, 1815 River, Hopewell, Virginia, to southern Georgia; coastal Alabama to Aransas Bay, Remarks.—Subgeneric designations Texas; Freshwater: Mississippi River and within Palaemonetes are considered valid tributaries upstream to McCurtain County, by Holthuis (1993). Oklahoma; Fort Smith, Arkansas; St. Louis, Missouri; Washington County, Ohio (Wil- Brachycarpus biunguiculatus liams 1984). (Lucas, 1849) Macrobrachium olfersii (Wiegmann, 1836) Remarks.—This species occurs from shallow water to 105 m (Ramos-Porto & Known range.—Western Atlantic: lower Coelho 1998). Cape Fear River, North Carolina; Florida; Known range.—Western Atlantic: Ber- Louisiana; Texas; Veracruz, Mexico, to Rio muda; Cape Fear, North Carolina; western Grande do Sul, Brazil (Williams 1984, Ra- Gulf of Mexico through West Indies to Cu- mos-Porto & Coelho 1998). ragao and Old Providence Island (Williams 1984); Amapa to Espirito Santo and Fer- Palaemonetes (Palaemonetes) intermedins nando de Noronha Archipelago, Brazil (Ra- Holthuis, 1949 mos-Porto & Coelho 1998); Eastern Atlan- tic: Mediterranean; West Africa; Eastern Known range.—Western Atlantic: Vine- Pacific: west American coast; Indo-Pacific yard Sound, Massachusetts, to Port Aransas (Williams 1984). Texas; Bahia de la Ascension, Quintana Roo, Mexico (Williams 1984).

Leander tenuicornis (Say, 1818) Palaemonetes (Palaemonetes) pugio Holthuis, 1949 Known range.—Western Atlantic: New- foundland Banks to Falkland Islands; trop- Known range.—Western Atlantic: Verte ical and subtropical waters worldwide ex- River, west of St. Modeste, Quebec, cept for west coast of Americas (Williams through Nova Scotia to Corpus Christi, 1984). Texas; Matamoros, Tamaulipas, to Campe- 104 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

che, Mexico (Williams 1984, Squires 1990, ~k Periclimenes (.Periclimenes) iridescens Rodrfguez-Almaraz et al. 2000). Lebour, 1949 Periclimenes iridescens Lebour.—Williams, Palaemonetes (Palaemonetes) vulgaris 1984:85. (Say, 1818) Known range.—Western Atlantic: Ber- Known range.—Western Atlantic: south- muda; off Cape Hatteras, North Carolina; ern Gulf of St. Lawrence to Cameron Coun- southern and northwestern Florida; Tobago; ty, Texas; Rio Champoton and near Progre- Cubagua Island, Venezuela (Williams so, Yucatan, Mexico (Williams 1984, 1984). Squires 1990). 'k Periclimenes (Periclimenes) Subfamily Pontoniinae Kingsley, 1878 longicaudatus (Stimpson, 1860) Remarks.—Subgeneric designations Periclimenes longicaudatus (Stimpson).— within Periclimenes are considered valid by Williams, 1984:86. Holthuis (1993). Known range.—Western Atlantic: Cape Neopontonides beaufortensis Hatteras, North Carolina, to southwestern (Borradaile, 1920) Florida; West Indies to Sao Paulo, Brazil (Williams 1984). Known range.—Western Atlantic: Beau- fort, North Carolina, to Grand Isle, Louisi- 'k Periclimenes (.Periclimenes) pedersoni ana; Caledonia Bay, Panama; Antigua (Wil- Chace, 1958 liams 1984). Periclimenes pedersoni Chace.—Williams, Periclimenaeus schmitti Holthuis, 1951 1984:87. Known range.—Western Atlantic: Bogue Known range.—Western Atlantic: off Sound, Black Rocks off New River and Cape Lookout, North Carolina; off north- Lockwoods Folly River, North Carolina; west Florida (?); Bahamas through West In- Dry Tortugas, Florida (Williams 1984). dies to Bonaire; Belize (Williams 1984).

Periclimenaeus wilsoni (Hay, 1917) Pontonia domestica Gibbes, 1850 Known range.—Western Atlantic: Beau- Known range.—Western Atlantic: Atlan- fort, North Carolina; Sapelo Island, Geor- tic Beach near Beaufort Inlet, North Caro- gia; Loggerhead Key, near Dry Tortugas; lina, through Gulf of Mexico to South Pa- Franklin County, Florida (Williams 1984). dre Island, Texas; Bahamas; Caribbean coast of Costa Rica (Williams 1984, Strenth ^k Periclimenes (Harpilius) americanus & Chace 1995, Vargas & Cortes 1999); (Kingsley, 1878) Eastern Atlantic: Madeira (Williams 1984). Periclimenes americanus (Kingsley).—Wil- • Pontonia manningi Fransen, 2000 liams, 1984:83. Remarks.—Fransen (2000:101) de- Known range.—Western Atlantic: Beau- scribed this species and provided illustra- fort, North Carolina, to western Gulf of tions, size, color, and host information. This Mexico, through West Indies to Aruba; species is found at shallow depths to 80 m Amapa to Pernambuco, Espfrito Santo and (Fransen 2000). Sao Paulo, Brazil (Williams 1984, Ramos- Known range.—Western Atlantic: North Porto & Coelho 1998). Carolina to Caribbean Sea; Gulf of Mexico; VOLUME 116, NUMBER 1 105

Eastern Atlantic: Canary Islands; Cape Cura£ao; Ceara to Parana, Brazil (Christof- Verde Islands (Fransen 2000). fersen 1984, 1998; McClure & Wicksten 1997). Christoffersen (1998) reported that Superfamily Alpheoidea Rafinesque, 1815 his earlier records (Christoffersen 1984) of Family Alpheidae Rafinesque, 1815 A. estuariensis in the Gulf of Mexico are • Alpheus angulosus. McClure, 2002 actually A. angulatus ( = A. angulosus, see above). However, McClure & Wicksten Alpheus angulatus McClure, 1995. (1997) have reported A. estuariensis occur- Remarks.—McClure (1995:85) described ring in the northern Gulf of Mexico. this species and provided a diagnosis, illustrations, color, size, and habitat information. Alpheus formosus Gibbes, 1850 McClure (2002) provided a replacement name (A. angulosus) for this species be- Known range.—Western Atlantic: Ber- cause angulatus had previously been used muda; near Beaufort, North Carolina, by Coutiere (1905) for a subspecies of Al- through Gulf of Mexico and West Indies to pheus (A. strenuus var. angulatus) from the Sao Paulo, Brazil (Williams 1984, Chris- Indo-Pacific. Comparisons of morphologi- toffersen 1998). cal variation among species, including A. angulosus, of the edwardsii group of Al- Alpheus heterochaelis Say, 1818 pheus occurring in the northern Gulf of Remarks.—McClure (1995) redescribed Mexico and northwestern Atlantic were this species and designated a neotype to provided in McClure (1995) and McClure clarify taxonomic confusion surrounding & Wicksten (1997). this species. Comparisons of morphological Known range.—Western Atlantic: Beau- variation among species, including A. het- fort, North Carolina, to Quintana Roo, erochaelis, of the edwardsii group of Al- Mexico, including northern Gulf of Mexi- pheus occurring in northern Gulf of Mexico co; Haiti (McClure 1995). and northwestern Atlantic were provided in McClure (1995) and McClure & Wicksten Alpheus armillatus H. Milne Edwards, (1997). 1837 Known range.—Western Atlantic: lower Known range.—Western Atlantic: Ber- Chesapeake Bay to Aransas County, Texas; muda; North Carolina through Gulf of Bermuda; Cuba; Cura£ao; Suriname; Para Mexico and West Indies to Santa Catarina, to Paraiba, Brazil (Williams 1984, McClure Brazil (Williams 1984, Christoffersen 1995, Christoffersen 1998). 1998). Alpheus normanni Kingsley, 1878 • Alpheus estuariensis Christoffersen, Remarks.—Based on material examined 1984 from both western Atlantic and eastern Pa- Remarks.—Christoffersen (1984:191) cific locations, Kim & Abele (1988) con- described this species and provided illustra- cluded that variation between Pacific and tions, color description, and ecological in- Atlantic forms of A. normanni was suffi- formation. This species occurs from the in- cient to recognize two species. This deci- tertidal region to depths of 22 m (Christof- sion has not gained universal acceptance fersen 1984). since A. normanni continues to be used for Known range.—Western Atlantic: east the Atlantic form. If A. normanni is restrict- coast of Florida (near Jacksonville and In- ed to the eastern Pacific as suggested by dian River region); northern Gulf of Mex- Kim & Abele (1988), then western Atlantic ico; Cuba; Dominican Republic; Trinidad; specimens would be known as A. packardii 106 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Kingsley, 1880 (Christoffersen 1998). Fur- Synalpheus longicarpus (Herrick, 1891) ther research is needed to resolve this tax- Known range.—Western Atlantic: Beau- onomic question. fort, North Carolina, to west Flower Garden Known range.—Western Atlantic: Reef, southeast of Galveston, Texas; Yu- around Cape Charles, Virginia, and lower catan, Mexico through West Indies to Rio Chesapeake Bay through Gulf of Mexico de Janeiro, Brazil (Williams 1984, Chris- and West Indies to Sao Paulo, Brazil; Ber- toffersen 1998). muda; Eastern Pacific: Gulf of California; Panama (Williams 1984, Christoffersen 1998). Synalpheus minus (Say, 1818) Known range.—Western Atlantic: Ber- * Automate dolichognatha De Man, 1888 muda; Cape Hatteras, North Carolina, to Sao Paulo, Brazil (Williams 1984, Chris- Automate gardineri Coutiere.—Williams, toffersen 1998). 1984:100.

Remarks.—Chace (1988) placed A. Synalpheus townsendi Coutiere, 1909 gardineri and three other nominal species in the synonymy of 'the variable and wide- Known range.—Western Atlantic: Ber- ranging' A. dolichognatha. muda; Beaufort, North Carolina, to Rio de Known range.—Western Atlantic: Beau- Janeiro, Brazil; Rocas Atoll, Brazil (Wil- fort Inlet, North Carolina; Yucatan; Virgin liams 1984, Christoffersen 1998); Eastern Islands; Barbados; Rio de Janeiro, Brazil; Pacific: Gulf of California (Williams 1984). Indo-Pacific: Red Sea to Samoa (Williams 1984, Christoffersen 1998). Chace (1988) Family Hippolytidae Dana, 1852 considered the range to be pantropical, ex- • Bythocaris nana Smith, 1885 cept for the eastern Atlantic. Remarks.—This species was considered extralimital by Williams (1984). Abele & Automate evermanni Rathbun, 1901 Martin (1989) redescribed the species, and Known range.—Western Atlantic: Vir- provided illustrations and developmental ginia to Texas; Puerto Rico; Sao Paulo and notes. Bythocaris nana occurs at 79—1175 Rio Grande do Sul, Brazil; Eastern Atlantic: m depth and is not common (Abele & Mar- Cape Verde Islands; Liberia to Nigeria tin 1989). (Williams 1984, Chace 1988, Christoffersen Known range.—Western Atlantic: Mar- 1998). tha's Vineyard, Massachusetts, to southern Florida; northeastern Gulf of Mexico (Abele & Martin 1989). Leptalpheus forceps Williams, 1965

Known range.—Western Atlantic: North • Caridion gordoni (Bate, 1858) Carolina to Sergipe, Brazil (Williams 1984, Christoffersen 1998). Remarks.—Williams (1984) considered this species to be extralimital. Squires (1990) provided a description and illustra- Synalpheus fritzmuelleri Coutiere, 1909 tions. This species occurs at 5—421 m depth Known range.—Western Atlantic: Ber- (Williams & Wigley 1977). muda; Beaufort, North Carolina, to Santa Known range.—Western Atlantic: south- Catarina, Brazil (Williams 1984, Christof- western Newfoundland to Chesapeake Bay; fersen 1998); South Atlantic: St. Helena Is- North Atlantic: Iceland; Eastern Atlantic: land; Eastern Pacific: Baja California (Wil- northern Europe to Bay of Biscay (Williams liams 1984). 1984, Squires 1990). VOLUME 116, NUMBER 1 107

Eualus fabricii (Kr0yer, 1841) Hippolyte coerulescens (Fabricius, 1775)

Known range.—Western Atlantic: Hud- Known range.—Tropical and subtropical son Bay, Foxe Basin, and northwestern Atlantic Ocean, including south of the Greenland to Cape Cod; North Pacific: Grand Banks in the Gulf Stream; and Sar- Chukchi Sea; Bering Sea to British Colum- gasso Sea (Williams 1984, Squires 1990). bia; Western Pacific: Sea of Okhotsk to Sea of Japan (Williams 1984, Squires 1990). * Hippolyte obliquimanus Dana, 1852 Hippolyte curacaoensis Schmitt.—Wil- Eualus gaimardii liams, 1984:117. (H. Milne Edwards, 1837) Remarks.—Udekem d'Acoz (1997) ex- Remarks.—Squires (1990) recognized amined topotypic specimens of Hippolyte two subspecies, E. g. gaimardii and E. g. obliquimanus Dana and H. exilirostratus belcheri, both of which show considerable Dana and determined that these two nomi- variation in diagnostic features and overlap nal species are identical. He also concluded completely in distribution. Williams (1984), that H. curacaoensis Schmitt is conspecific however, believed that observed morpho- with H. obliquimanus. Udekem d'Acoz logical variation was not sufficient to war- (1997) redescribed H. obliquimanus based rant recognition of more than one species on this new information. among this material. Known range.—Western Atlantic: Beau- Known range.—Western Atlantic: fort and Sneads Ferry, North Carolina; Greenland and Baffin Island to Cape Cod; Puerto Rico; West Indies from Cuba to Cu- Eastern Atlantic: Spitsbergen to North Sea; ra£ao; Venezuela to Santa Catarina, Brazil Arctic Ocean: Point Barrow to Siberia; (Williams 1984, Udekem d'Acoz 1997, North Pacific: south to Sitka, Alaska (Wil- Christoffersen 1998). liams 1984, Squires 1990).

Hippolyte pleuracanthus (Stimpson, 1871) Eualus pusiolus (Kr0yer, 1841) Known range.—Western Atlantic: Con- Known range.—Western Atlantic: Gulf necticut to North Carolina (Williams 1984). of St. Lawrence to Cape Henry, Virginia; Northeastern Atlantic: Iceland; Murman Hippolyte zostericola (Smith, 1873) Sea to Channel Islands, southward along Bay of Biscay to Spain; Catalonian coast of Known range.—Western Atlantic: south- Spain; North Pacific: Chukchi and Bering ern Massachusetts; Bermuda; North Caro- seas to British Columbia and Washington; lina to Yucatan; Trinidad; Cura£ao; Ceara, Western Pacific: Sea of Okhotsk and Sea of Brazil (Williams 1984, Christoffersen Japan (Williams 1984, Squires 1990). 1998). Christoffersen (1998) noted that previously reported occurrences of this species Exhippolysmata oplophoroides in Brazil may actually refer to H. obliqui- (Holthuis, 1948) manus.

Remarks.—This species occurs over a Latreutes fucorum (Fabricius, 1798) bathymetric range of 5-45 m (Rodriguez 1993, Caetano da Costa et al. 2000). Known range.—Western Atlantic: New- Known range.—Western Atlantic: Cape foundland to Brazil (Pernambuco to Bahia), Fear, North Carolina, to Port Aransas, Tex- including Gulf of Mexico south to Bahia de as; Guyana to northern Uruguay (Williams la Ascension, Quintana Roo, Mexico; East- 1984, Caetano da Costa et al. 2000). ern Atlantic: Azores; Cape Verde Islands 108 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

(Williams 1984, Christoffersen 1998, Rod- British Columbia; Eastern Atlantic: Arctic riguez-Almaraz et al. 2000). to the Hebrides (Williams 1984, Squires 1990). Latreutes parvulus (Stimpson, 1866) Lysmata rathbunae Chace, 1970 Known range.—Western Atlantic: Beau- fort, North Carolina, to Buenos Aires, Ar- Known range.—Western Atlantic: Ber- gentina; Eastern Atlantic: West Africa (Wil- muda; Cape Fear, North Carolina; east coast liams 1984, Christoffersen 1998). of Florida to Yucatan; Venezuela (Williams 1984). Lebbeus groenlandicus (Fabricius, 1775) Lysmata wurdemanni (Gibbes, 1850) Known range.—Western Atlantic: Hud- Known range.—Western Atlantic: New son Bay to Greenland, southward to Rhode Jersey to Port Aransas, Texas; Suriname; Island; North Pacific: Arctic Canada, south- French Guiana; Ceara and Bahia to Rio ern Chukchi Sea through Bering Sea to Pu- Grande do Sul, Brazil (Williams 1984, get Sound; Western Pacific: Okhotsk Sea Christoffersen 1998). southward to Vladivostok (Williams 1984, Squires 1990). Spirontocaris liljeborgii (Danielssen, 1859) * Lebbeus microceros (Kr0yer, 1841) Known range.—Western Atlantic: Foxe Lebbeus zebra (Leim).—Williams, 1984: Channel and Davis Strait; Greenland, to off 125. Delaware Bay; Northeastern Atlantic: Ice- Remarks.—Holthuis (1947) recognized land; Spitsbergen; Murman coast to south both L. microceros and L. zebra as valid coast of England; west and southwestern species. Couture & Trudel (1968) noted Ireland; Arctic: Alaska (Williams 1984, great similarity between these nominal spe- Squires 1990). cies and commented that they might be syn- onymous, but rarity of material prevented Spirontocaris phippsii (Kr0yer, 1841) an adequate evaluation of these taxa (Wil- Known range.—Circumarctic; Western liams 1984). Squires (1990) placed L. zebra Atlantic: Cornwallis Island; Hudson Bay to into the synonymy of L. microceros without Martha's Vineyard; North Atlantic: Spits- comment or explanation for this action. bergen to southern Norway; Britain; Arctic: Chace (1997) also listed L. zebra in the syn- north of Siberia; Beaufort Sea; North Pa- onymy of L. microceros. cific: Bering Sea to Siberian east coast; Known range.—Western Atlantic: South- Shumagin Islands, Alaska (Williams 1984, ern Greenland; Foxe Basin; Ungava Bay; Squires 1990). Newfoundland to New Brunswick, including Gulf of St. Lawrence to southeast of Spirontocaris spinus (Sowerby, 1805) Isles of Shoals; North Pacific: possibly from Bering Sea to Kamchatka; Eastern Pa- Known range.—Circumarctic; Western Atlantic: Hudson Bay; Foxe Basin; Green- cific: Checleset Bay, Vancouver Island land, southward to Massachusetts Bay; (Williams 1984, Squires 1990). North Atlantic: Spitsbergen to northern North Sea and Irish Sea; North Pacific: Plo- Lebbeus polaris (Sabine, 1824) ver Bay, Siberia; Bering Sea; Shumagin Is- Known range.—Circumarctic; Western lands, Alaska; Puget Sound, Washington; Atlantic: southward to Chesapeake Bay; Western Pacific: Okhotsk Sea; Sea of Japan North Pacific: Okhotsk Sea; Bering Sea to (Williams 1984, Squires 1990). VOLUME 116, NUMBER 1 109

Thor dobkini Chace, 1972 mented on color, interspecific comparisons, and sexual dimorphism. Known range.—Western Atlantic: off Known range.—Western Atlantic: South Shackleford Bank, North Carolina, to Yu- Carolina through the Gulf of Mexico; Ca- catan; Louisiana; north coast of Cuba (Wil- ribbean Sea to Santa Marta, Colombia (Cri- liams 1984). ales 1992).

Thor floridanus Kingsley, 1878 Family Ogyrididae Holthuis, 1955 Known range.—Western Atlantic: Black Ogyrides alphaerostris (Kingsley, 1880) Rocks off New River, North Carolina (?), Known range.—Western Atlantic: Vir- to Yucatan (Williams 1984). ginia to Rio Grande do Sul, Brazil, including Gulf of Mexico (Williams 1984, Chris- Thor manningi Chace, 1972 toffersen 1998). Known range.—Western Atlantic: Beau- fort, North Carolina, to Yucatan, through Ogyrides hayi Williams, 1981 West Indies to Cura9ao; Brazil (Paraiba, Known range.—Western Atlantic: Beau- Bahia, and Sao Paulo); Eastern Pacific: Is- fort, North Carolina, to Sebastian Inlet, las Tres Marias, Mexico (Williams 1984, Florida; northwestern Florida to Mississip- Christoffersen 1998). pi; Puerto Rico; Pernambuco and Sao Pau- lo, Brazil (Williams 1984, Christoffersen Tozeuma carolinense Kingsley, 1878 1998). Known range.—Western Atlantic: Vine- yard Sound, Massachusetts, to Colon, Pan- • Superfamily Processoidea Ortmann, ama, including Gulf of Mexico; through 1890 West Indies to Cura9ao; Paraiba to Alagoas Remarks.—Previously the family Proces- and Sao Paulo, Brazil (Williams 1984, sidae was placed in the superfamily Al- Christoffersen 1998). pheoidea (Williams 1984). Chace (1992) erected the superfamily Processoidea, Tozeuma serratum A. Milne-Edwards, which consists of a single family (Proces- 1881 sidae). Holthuis (1993) also adopted this arrangement. Known range.—Western Atlantic: Non- amesset Island, Massachusetts; off Cape Family Processidae Ortmann, 1890 Hatteras and Cape Lookout, North Caroli- Nikoides schmitti Manning & Chace, 1971 na; Cape Canaveral; extreme southern and northwestern Florida; Barbados; Colombia; Known range.—Western Atlantic: east of off Rio de Janeiro and Sao Paulo, Brazil Cape Lookout, North Carolina; Biscayne (Williams 1984, Christoffersen 1998). Bay and Dry Tortugas, Florida; Guade- loupe; the Guianas (Williams 1984). • Trachycaris rugosa (Bate, 1888) Processa bermudensis (Rankin, 1900) Remarks.—Based on available information, Williams (1984) presumed this species Known range.—Western Atlantic: Ber- to occur beyond the southern limits of the muda; Cape Hatteras, North Carolina, to region. However, Criales (1992) recorded T. northwestern Florida; Veracruz, Mexico; rugosa from southern areas within this re- Cuba; Puerto Rico; Guadeloupe; Venezuela; gion. Criales (1992) provided a redescrip- Bahia, Rio de Janeiro and Parana, Brazil tion of the species, illustrations, and com- (Williams 1984, Christoffersen 1998). 15 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Processa fimbriata Manning & Chace, Remarks.—In his revision of the genus 1971 Pandalus, Komai (1999) transferred Pan- dalus propinqvus to a monotypic genus At- Known range.—Western Atlantic: off lantopandalus; a description was provided. New River, North Carolina, to Rio de Ja- Additionally, Komai (1999) stated that G.O. neiro, Brazil (Williams 1984, Christoffersen Sars consistently spelled the name of this 1998). species as 'propinqvus' in the original description and that this spelling should be Processa guyanae Holthuis, 1959 used. This species occurs at depths of 20- Known range.—Western Atlantic: Cape 2180 m (Komai 1999). Hatteras, North Carolina, to eastern Gulf of Known range.—Western Atlantic: Mexico, including northern coast of Cuba; Greenland; Davis Strait to Delaware Bay, Suriname to Uruguay (Williams 1984, including Gulf of St. Lawrence and Gulf of Christoffersen 1998). Maine; North Atlantic: Iceland; Eastern At- lantic: Norway to the British Isles; Bay of Processa hemphilli Manning & Chace, Biscay (Komai 1999). 1971 Dichelopandalus leptocerus (Smith, 1881) Known range.—Western Atlantic: east of Cape Lookout and Bogue Sound, North Known range.—Western Atlantic: Gulf Carolina; east coast of Florida; northwest of St. Lawrence and St. Mary's Bay, New- Florida; Guadeloupe; Brazil to Buenos Ai- foundland, to off Oregon Inlet, North Car- res (Williams 1984, Christoffersen 1998). olina; North Pacific: Shumagin Bank, Alas- ka (Williams 1984, Squires 1990). Processa profunda Manning & Chace, 1971 • Heterocarpus ensifer A. Milne-Edwards, 1881 Known range.—Western Atlantic: southeast of Cape Hatteras, North Carolina; off Remarks.—Diagnostic characters of this South Carolina; Gulf of Mexico, off south- species were reported in Crosnier & Forest ern and western Florida; Suriname to Uru- (1973), Chace (1985), and Rodriguez guay (Williams 1984, Christoffersen 1998). (1993). This species occurs on muddy bottoms usually at 200-885 m (Chace 1985), Processa vicina Manning & Chace, 1971 but may be found at shallower depths (140 m; Rodriguez 1993). Known range.—Western Atlantic: south- Known range.—Western Atlantic: North east of Cape Lookout, North Carolina; Carolina to the Guianas, including the Gulf northwest Florida; off Isla Margarita, Ve- of Mexico and Caribbean Sea; Brazil (Cha- nezuela (Williams 1984). ce 1985, Rodriguez 1993, Ramos-Porto & Coelho 1998). Superfamily Pandaloidea Haworth, 1825 Remarks.—Christoffersen (1989) pro- Pandalus borealis Kr0yer, 1838 posed a phylogeny of this taxon. Remarks.—The geographic range of Pandalus borealis has been reported as Family Pandalidae Haworth, 1825 Arctic boreal in both Atlantic and Pacific ^k Atlantopandalus propinqvus oceans (Williams 1984). Recent studies (G.O. Sars, 1870) (Squires 1992, Komai 1999) have demon- Pandalus propinquus G.O. Sars.—Wil- strated that morphological differences be- liams, 1984:156. tween Atlantic and Pacific 'varieties' were VOLUME 116, NUMBER 1 ill sufficient to warrant full species rank for Mediterranean Sea; Central Pacific: Hawaii; both. Pandalus borealis is restricted to the Indo-Pacific: East Africa to Japan, includ- Atlantic Ocean and considered a geminate ing Gulf of Aden; New Zealand; southeast- species of P. eous Makarov from Pacific ern Australia (Williams & Wigley 1977). localities (Squires 1992). Known range.—Western Atlantic: west- • Plesionika tenuipes (Smith, 1881) ern Greenland to Gulf of Maine; North At- Remarks.—Williams (1984) considered lantic: Barents Sea to the North Sea (Komai this species to be extralimital. This species 1999). occurs at 159-476 m depth (Williams 1984). Pandalus montagui Leach, 1814 Known range.—Western Atlantic: Rhode Known range.—Arctic-boreal; Western Island to southern Florida; Gulf of Mexico Atlantic: Greenland and Hudson Bay to (Williams 1984). Rhode Island; North Atlantic: Iceland; • :k Plesionika willisi White Sea; Eastern Atlantic: Norway to the (L. H. Pequegnat, 1970) western Baltic; North Sea; British Isles (Williams 1984, Squires 1990). Parapandalus willisi L. H. Pequegnat.— Williams, 1984:484. Pantomus parvulus A. Milne-Edwards, Remarks.—Williams (1984) considered 1883 this species to be extralimital. Chace (1985) Known range.—Western Atlantic: Cape placed Parapandalus into the synonymy of Lookout, North Carolina, to Yucatan; Plesionika. This species occurs at 150-500 Puerto Rico; St. Croix; Suriname; Uruguay m depth (Williams & Wigley 1977). (Williams 1984, Christoffersen 1989). Known range.—Western Atlantic: south of Martha's Vineyard, Massachusetts; Gulf • Plesionika edwardsii (Brandt, 1851) of Mexico to French Guiana (Williams & Wigley 1977). Remarks.—Chan & Yu (1991) provided a diagnosis, color description, and interspe- Superfamily Crangonoidea Haworth, 1825 cific comparisons. This species occurs on Family Crangonidae Haworth, 1825 muddy bottoms at depths of 50-690 m Argis dentata (Rathbun, 1902) (Rodriguez 1993), commonly at 200-400 Remarks.—Komai (1997) evaluated the m (Chan & Yu 1991). systematics of this species and several con- Known range.—Western Atlantic: Vir- geners to clarify their taxonomic status. In ginia to Gulf of Mexico; Eastern Atlantic: this study, he also provided a detailed de- Mediterranean to Angola; Indo-Pacific scription, illustrations, color, size and com- (Chan & Yu 1991). parative information for A. dentata. Known range.—Arctic-boreal; Western • Plesionika martia North Atlantic: northwest Greenland; Hud- (A. Milne-Edwards, 1883) son Bay; Canadian Arctic islands to Nova Remarks.—Williams (1984) considered Scotia; North Pacific: Bering Sea to Sitka, this species to be extralimital. This species Alaska; southeast coast of Kamchatka; occurs at 165-2100 m depth (Williams & northern Okhotsk Sea (Komai 1997). Wigley 1977). Known range.—Western Atlantic: south * Crangon (Crangon) septemspinosa Say, of Nantucket, Massachusetts, to Brazil; 1818 Bermuda; Eastern Atlantic: southwest Ire- Crangon septemspinosa Say.—Williams, land; Bay of Biscay to southern Africa; 1984:159. 112 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—Subgeneric designation con- Sabinea septemcarinata (Sabine, 1824) sidered valid by Holthuis (1993). Known range.—Western Atlantic: Hud- Known range.—Primarily subarctic-bo- son Bay and Greenland to Massachusetts real; Western Atlantic: northern Gulf of St. Bay; Arctic-North Atlantic: Iceland (except Lawrence to east Florida (Williams 1984, south coast), Kara, White and Barents seas; Squires 1990). Squires (1990) restricted the Wrangel Island; Eastern Atlantic: north of geographic range to the north Atlantic. Faroes; Norway (north of 67°N); British Isles; North Pacific: Arctic Canada and * Philocheras gorei (Dardeau, 1980) Alaska to Point Barrow and Chukchi Sea Pontophilus gorei Dardeau.—Williams, (Williams 1984, Squires 1990). 1984:161. Sclerocrangon boreas (Phipps, 1774) Remarks.—Chace (1984) provided morphological evidence for recognizing Phil- Known range.—Western Atlantic: Hud- ocheras as a valid genus and removed it son Bay; east and west Greenland south to from the synonymy of Pontophilus. Chris- Cape Cod; Arctic-North Atlantic: Iceland; toffersen's (1988) phylogenetic analysis Spitsbergen; Kara and White seas; Franz supported this arrangement. Bathymetric Joseph Land; Novaya Zemlya; Eastern At- range was reported as 59—194 m (Christof- lantic: Faroes; Norway (north of approxi- fersen 1998). mately 67°N); Arctic-North Pacific: Arctic Known range.—Western Atlantic: central Canada, north coast of Alaska; Chukchi Sea Georgia; central eastern Florida; Gulf of south to British Columbia (Williams 1984, Mexico (southwestern Florida, Cape San Squires 1990). Bias and Padre Island, Texas); Rio de Ja- Infraorder Astacidea Latreille, 1802 neiro, Brazil; Uruguay (Dardeau & Heard 1983, Williams 1984, Christoffersen 1988). Superfamily Nephropoidea Dana, 1852 Family Nephropidae Dana, 1852 Pontophilus brevirostris Smith, 1881 Subfamily Nephropinae Dana, 1852 Homarus americanus H. Milne Edwards, Known range.—Western Atlantic: Gulf 1837 of Maine to eastern Gulf of Mexico, off Known range.—Western Atlantic: New- Dry Tortugas, Florida; Cuba (Williams foundland to Cape Hatteras, North Caroli- 1984). na; occasionally Wilmington, North Caro- lina and south Florida (Williams 1984, Cof- Pontophilus norvegicus (M. Sars, 1861) er-Shabica & Nielsen 1988, Holthuis 1991). Known range.—Western Atlantic: Williams (1984) noted that this species is Greenland to Maryland; North Atlantic: occasionally found as far south as Wil- Iceland; Spitsbergen; Murman coast; East- mington, North Carolina. One specimen of ern Atlantic: northwestern Europe, includ- H. americanus, however, was collected off ing British Isles to Bay of Biscay; and Bal- southern Florida (Miami Beach, 252 m; earic Islands (Williams 1984, Squires Cofer-Shabica & Nielsen 1988), suggesting 1990). that the range of this species occasionally extends further south, especially in deeper Sabinea sarsii Smith, 1879 waters. Known range.—Western Atlantic: Davis Strait to southeast of Nantucket, Massachu- • Subfamily Thymopinae Holthuis, 1974 setts; North Atlantic: Iceland; Eastern At- • Nephropsis aculeata Smith, 1881 lantic: northern Europe (Williams 1984, Remarks.—Species diagnosis provided in Squires 1990). Hothuis (1991). This species occurs on mud VOLUME 116, NUMBER 1 113 or fine sand sediments at 137-824 m, but Remarks.—In a recent revision of the usually between 200-600 m (Holthuis American Callianassidae, Manning & Feld- 1991). er (1991) concluded that Callianassa was a Known range.—Western Atlantic: Mas- composite of numerous genera. Manning & sachusetts to French Guiana, including Gulf Felder (1991) described the genus Biffarius of Mexico and Caribbean Sea; Bermuda and provided a diagnosis, illustrations, and (Holthuis 1991). comparative information regarding the genera within the subfamily Callianassinae. * Infraorder Thalassinidea Latreille, 1831 Known range.—Western Atlantic: Bass Remarks.—Thalassinidea was considered River, Yarmouth, Nova Scotia; Nantucket a Section under the infraorder Anomura in Sound, Massachusetts; Chesapeake Bay (?); Williams (1984:180). Poore (1994) pro- North Inlet, South Carolina, to Mcintosh posed a phylogeny of the infraorder and County, Georgia; Franklin County, north- provided a new classification, diagnoses, west Florida (Williams 1984). and keys to families and currently recognized genera. * Gilvossius setimanus (De Kay, 1844) Callianassa atlantica Rathbun.—Williams, Superfamily Callianassoidea Dana, 1852 1984:180. Family Callianassidae Dana, 1852 Remarks.—Manning & Felder (1991) re- Remarks.—Manning (1987) addressed stricted this family as a result of their re- the status of Gonodactylus setimanus vision of the American Callianassidae. In a DeKay. He determined this species was val- phylogenetic analysis of generic relation- id as Callianassa setimanus (DeKay) and ships within the family based on 93 adult that it was the senior synonym of Calli- morphological characters, Tudge et al. anassa atlantica Rathbun. Members of Cal- (2000) determined that the Callianassidae lianassa sensu stricto, as restricted by Man- comprised a monophyletic group. Sakai ning & Felder (1991), are not represented (1999) presented very different conclusions in the American fauna. The genus Gilvos- regarding the composition of, and generic sius was described (Manning & Felder, relationships within, the Callianassidae. Sa- 1992) with Gonodactylus setimanus DeKay kai's work, however, was not conducted (= Callianassa atlantica Rathbun) as the within a phylogenetic framework and is type species. widely regarded as controversial. More re- Known range.—Western Atlantic: Bass search is needed to resolve the composition River, Nova Scotia, to Georgia; Franklin and relationships of genera in this and re- County, Florida (Williams 1984). lated families and subfamilies. • Necallianassa berylae Heard & Subfamily Callianassinae Dana, 1852 Manning, 1998 Remarks.—Tudge et al. (2000) conduct- Remarks.—Heard & Manning (1998: ed a phylogenetic analysis of relationships 883-884) described this genus and species within the family Callianassidae based on and provided illustrations and diagnostic adult morphological characters and they de- comparisons with eastern Atlantic conge- termined that the subfamily Callianassinae ners. Necallianassa berylae occurs at was a monophyletic group. depths of 35-75 m (Heard & Manning 1998). * Biffarius biformis (Biffar, 1971) Known range.—Western Atlantic: South Callianassa biformis Biffar.—Williams, Carolina and Georgia (Heard & Manning 1984:182. 1998). 19 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Subfamily Callichirinae Manning & Superfamily Axioidea Huxley, 1879 Felder, 1991 Family Axiidae Huxley, 1879 • Axius armatus Smith, 1881 Remarks.—Tudge et al. (2000) conducted a phylogenetic analysis of relationships Remarks.—Kensley (2001) redescribed within the family Callianassidae utilizing this species, provided illustrations, and adult morphological characters. They deter- compared A. armatus to its sympatric con- mined that the subfamily Callichirinae was gener, A. serratus. Although Sakai & de a paraphyletic group and that more research Saint Laurent (1989) questioned the generic is needed to resolve relationships within placement of this species, Kensley (2001) this group. provided morphological evidence supporting placement of this species in the genus it Callichirus major (Say, 1818) Axius. This apparently rare species occurs at depths of 108-260 m (Kensley 2001). Callianassa major Say.—Williams, 1984: Known range.—Western Atlantic: Mas- 183. sachusetts to South Carolina (Kensley Remarks.—Manning & Felder (1986) re- 2001). defined Callichirus to remove the ambigu- ity surrounding previous concepts of this Axius serratus Stimpson, 1852 genus. Remarks.—Kensley (2001) redescribed, Known range.—Western Atlantic: Beau- illustrated, and compared this species to its fort Inlet, North Carolina, to Cape Canav- Atlantic congeners. This species occurs at eral, Florida; Grand Terre Island to Tim- depths of 19-220 m (Kensley 2001), which balier Island, Louisiana; Sergipe to Santa represents an increase in the maximum re- Catarina, Brazil (Williams 1984, Rodrigues ported depth of occurrence. & Shimizu 1998). Known range.—Western Atlantic: Nova Scotia to Maryland (Kensley 2001). Family Laomediidae Borradaile, 1903 Naushonia crangonoides Kingsley, 1897 * Calaxius jenneri (Williams, 1974) Known range.—Western Atlantic: Bass Axiopsis jenneri (Williams).—Williams, River, Yarmouth, Nova Scotia; Vineyard 1984:185. Sound and Elizabeth Islands, Massachu- setts, to Bogue Sound, North Carolina (Wil- Remarks.—Sakai & de Saint Laurent liams 1984). (1989) described the genus Calaxius in their revision of the family Axiidae. Known range.—Western Atlantic: Cape Family Upogebiidae Borradaile, 1903 Lookout, North Carolina (Williams 1984). Upogebia ajfinis (Say, 1818) Remarks.—Williams (1993) provided an • Family Calocarididae Ortmann, 1891 improved diagnosis and detailed description Remarks.—Kensley (1989) reinstated the of this species and noted that its southern Calocarididae Ortmann, which contained limits of distribution are in Texas. Speci- only Calocaris, and then expanded the fam- mens identified as U. ajfinis from more ily with the addition of several genera. southern locations (West Indies to Sao Pau- lo, Brazil) are actually Upogebia parafftnis Calocaris templemani Squires, 1965 Williams, 1993. Known range.—Western Atlantic: Mas- Known range.—Western Atlantic: New- sachusetts to southern Texas (Williams foundland to the Gulf of Maine; Cape 1993). Lookout, North Carolina (Williams 1984, VOLUME 116, NUMBER 1 115

Kensley 1989, Sakai & de Saint Laurent ribbean Sea (Williams 1984, Holthuis 1989, Squires 1990). The southernmost lo- 1991). cality reported for this species is southeast of Cape Lookout, North Carolina (Williams Family Scyllaridae Latreille, 1825 1984). This appears to be the only recorded Subfamily Arctidinae Holthuis, 1985 occurrence of C. templemani this far south. • Scyllarides aequinoctialis (Lund, 1793)

Remarks.—This species was originally Infraorder Palinura Latreille, 1802 considered to occur outside the region, and Superfamily Palinuroidea Latreille, 1802 therefore was not included in Williams Family Palinuridae Latreille, 1802 (1984). Lyons (1970) provided a detailed Panulirus argus (Latreille, 1804) description. Scyllarides aequinoctialis occurs on sandy or rocky bottoms at depths Remarks.—Based on mtDNA samples of 0-180 m, usually 1-64 m (Lyons 1970, from individuals collected from nine loca- Holthuis 1991). tions between Bermuda and Venezuela, Sil- Known range.—Western Atlantic: Ber- berman et al. (1994) hypothesized that Pan- muda; South Carolina to southern Brazil, ulirus argus was genetically homogenous including the Gulf of Mexico, Caribbean throughout the tropical western Atlantic and Sea, and West Indies (Holthuis 1991). Caribbean. However, three individuals with distinctly different mtDNA haplotypes, collected off Miami, Florida, were identified Scyllarides nodifer (Stimpson, 1866) (Silberman et al. 1994). Sarver et al. (1998) Known range.—Western Atlantic: Ber- compared mtDNA sequences between muda; Cape Lookout, North Carolina, to western Atlantic-Caribbean populations and Yucatan, including Gulf of Mexico (Wil- Brazilian lobsters and found sufficient dif- liams 1984, Holthuis 1991). ferences to suggest that P. argus is a complex of two species or subspecies. In addition to genetic differences, characteristic Subfamily Scyllarinae Latreille, 1825 color patterns were also identified distin- Scyllarus americanus (Smith, 1869) guishing the Brazilian P. argus from the Known range.—Western Atlantic: Bogue Caribbean form (Sarver et al. 1998). They Inlet, North Carolina, to Campeche Banks, recommended provisional recognition of Mexico; Venezuela (Williams 1984). two subspecies until formal taxonomic revision could be completed: Panulirus argus argus representing populations from Ber- Scyllarus chacei Holthuis, 1960 muda to Venezuela and Panulirus argus Known range.—Western Atlantic: Cape westonii representing populations from Bra- Hatteras, North Carolina, to Bahia, Brazil, zil. Sarver et al. (2000) re-examined the ge- including Gulf of Mexico, West Indies, and netically distinct individuals of Silberman Caribbean Sea (Williams 1984, Coelho & et al. (1994) and determined that these un- Ramos-Porto 1998b). usual individuals were the provisionally recognized Brazilian form of P. argus (P. argus westonii). However, until formal re- Scyllarus depressus (Smith, 1881) vision is conducted, the taxonomic status of Known range.—Western Atlantic: Mar- these subspecies remains uncertain. tha's Vineyard, Massachusetts; Cape Hat- Known range.—Western Atlantic: Ber- teras, North Carolina, to Sao Paulo, Brazil, muda; North Carolina to Rio de Janeiro, including Gulf of Mexico and West Indies Brazil, including Gulf of Mexico and Ca- (Williams 1984). 21 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Infraorder Anomura MacLeay, 1838 Munida longipes A. Milne-Edwards, 1880 Superfamily Galatheoidea Samouelle, 1819 Known range.—Western Atlantic: Balti- Family Galatheidae Samouelle, 1819 more Canyon, off the coast of Maryland Galathea rostrata A. Milne-Edwards, (Williams 1988) southward through Gulf of 1880 Mexico to Belize; through West Indies to Curasao (Williams 1984); Sao Paulo to Rio Known range.—Western Atlantic: off Grande do Sul, Brazil (Melo-Filho 1998). Cape Hatteras, North Carolina, to southern Florida; northwestern Florida to Mississippi Munida pusilla Benedict, 1902 River delta; off Cape Catoche, Yucatan (Williams 1984). Known range.—Western Atlantic: off Cape Lookout, North Carolina, to Straits of • Munida forceps A. Milne-Edwards, Florida, through eastern Gulf of Mexico to 1880 Yucatan; Colombia; Trinidad (Williams 1984); Amapa, Brazil (Melo-Filho 1998). Remarks.—Williams (1984) did not include this species in his study because the Munida valida Smith, 1883 known geographic range and depth of occurrence were outside the region. This spe- Known range.—Western Atlantic: off cies is now known from off southern New Nova Scotia to Rio Grande do Sul, Brazil, England and Virginia within the region. including Gulf of Mexico and Caribbean Munida forceps occurs at 80—337 m (Wil- (Williams 1984, Squires 1990, Melo-Filho liams 1988, Melo-Filho & Melo 1992). & Melo 1992, Melo-Filho 1998). Melo-Filho & Melo (1992) provided a description, illustrations, and measurements; Family Porcellanidae Haworth, 1825 Williams (1988) provided a color descrip- Euceramus praelongus Stimpson, 1860 tion and figure of male carapace. Known range.—Western Atlantic: Veatch Known range.—Western Atlantic: Dela- and Lydonia Canyons, off southern New ware Bay to Aransas area, Texas (Williams England (Williams 1988); Virginia; Florida; 1984). Gulf of Mexico; Antilles; Guianas; Brazil (Melo-Filho & Melo 1992). Megalobrachium soriatum (Say, 1818)

Munida iris iris A. Milne-Edwards, 1880 Known range.—Western Atlantic: off Cape Hatteras, North Carolina, to Port Known range.—Western Atlantic: off Aransas, Texas; West Indies to Barbados; Nova Scotia through southeastern Gulf of Contoy Island, Mexico; Bahia Caledonia Mexico to near Cozumel Island, Yucatan; and Galeta Island, Panama (Williams through Caribbean islands to Rio Grande do 1984); Ceara to Sao Paulo, Brazil (Veloso Sul, Brazil (Williams 1984, Squires 1990, 1998). Melo-Filho 1998). Pachycheles pilosus Munida irrasa A. Milne-Edwards, 1880 (H. Milne Edwards, 1837)

Known range.—Western Atlantic: off Known range.—Western Atlantic: Bermuda; off Cape Lookout, North Caro- Charleston, South Carolina; Key West to lina, through eastern Gulf of Mexico; Ca- Sarasota Bay, Florida; through West Indies ribbean Sea to Uruguay (Williams 1984). to Tobago and Aruba (Williams 1984). VOLUME 116, NUMBER 1 117

Pachycheles rugimanus Porcellana sayana (Leach, 1820) A. Milne-Edwards, 1880 Known range.—Western Atlantic: Cape Known range.—Western Atlantic: Cape Hatteras, North Carolina, to Rio Grande do Lookout, North Carolina, through Florida Sul, Brazil, including Gulf of Mexico and to St. Thomas, U.S. Virgin Islands; Contoy Caribbean Sea (Williams 1984, Veloso Island, Mexico (Williams 1984); Amapa to 1998). Para (Veloso 1998) and Pernambuco, Brazil (Williams 1984). Porcellana sigsbeiana A. Milne-Edwards, 1880 • Petrolisthes armatus (Gibbes, 1850) Known range.—Western Atlantic: off Remarks.—This species was recently re- Martha's Vineyard, Massachusetts, to ported in the region (Knott et al. 2000). Al- southwestern Caribbean Sea off Colombia; though Knott et al. (2000) reported the West Indies to Virgin Islands (Williams presence of this crab as an introduction, the 1984); Para and Maranhao, Brazil (Veloso occurrence of P. armatus in Georgia and 1998). South Carolina possibly represents a northern range extension because the original de- Superfamily Hippoidea Latreille, 1825 scription by Gibbes (1850) listed the local- Family Albuneidae Stimpson, 1858 ity of specimens examined as "Florida." Albunea gibbesii Stimpson, 1859 This species is the dominant decapod crustacean on rocky substrates in Georgia and Known range.—Western Atlantic: Ber- is well established on rocky rubble, oyster muda; east of Cape Lookout, North Caro- reefs, and other shallow subtidal and inter- lina, to Texas, through West Indies to Sao tidal habitats throughout Georgia and South Paulo, Brazil (Williams 1984, Calado Carolina (Knott et al. 2000). 1998). Known range.—Western Atlantic: Ber- muda; South Carolina southward through Albunea paretii Guerin-Meneville, 1853 the Gulf of Mexico; Bahamas; West Indies and Caribbean; northern South America to Known range.—Western Atlantic: Beau- Santa Catarina, Brazil; Central Atlantic: As- fort Inlet, North Carolina, to Corpus Christi, cension Island; Eastern Atlantic: tropical Texas, through West Indies to Rio Grande West Africa; Eastern Pacific: Gulf of Cali- do Sul, Brazil (Williams 1984, Calado fornia to Peru (Veloso 1998, Knott et al. 1998); Eastern Atlantic: Cape Verde Is- 2000). lands; Senegal to Ghana (Williams 1984).

Petrolisthes galathinus (Bosc, 1802) Lepidopa websteri Benedict, 1903 Known range.—Western Atlantic: Cape Known range.—Western Atlantic: mouth Hatteras, North Carolina, through Gulf of of Chesapeake Bay; Drum Inlet, North Car- Mexico and Caribbean Sea to Rio Grande olina, to east central Florida; Tampa Bay, do Sul, Brazil; Eastern Pacific: Isla San Lu- Florida; Petit Bois Island, Mississippi (Wil- cas, Costa Rica, to off La Libertad, Ecuador liams 1984, Manning 1988). (Williams 1984, Veloso 1998). Family Hippidae Latreille, 1825 Poly onyx gibbesi Haig, 1956 Emerita benedicti Schmitt, 1935 Known range.—Western Atlantic: Woods Known range.—Western Atlantic: Hole, Massachusetts, to Uruguay (Williams Charleston County, South Carolina, to Ve- 1984). racruz, Mexico (Williams 1984). 118 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Emerita talpoida (Say, 1817) sas, Texas; through West Indies to Argen- tina (Williams 1984, Rieger 1998). Known range.—Western Atlantic: Har- wich, Massachusetts, to Horn Island, Mis- Paguristes hummi Wass, 1955 sissippi; Progreso, Yucatan, Mexico (Wil- liams 1984). Known range.—Western Atlantic: New- port River, North Carolina, to off Sapelo Superfamily Coenobitoidea Dana, 1851 Island, Georgia; southwestern Florida, in- Remarks.—Williams (1984) recognized cluding Tampa Bay, to off Isles Dernieres, two superfamilies (Coenobitoidea and Pa- Louisiana; Caribbean coast of Colombia guroidea) for hermit crabs. McLaughlin (Williams 1984, Campos & Sanchez 1995, (1983) had recommended that the super- Strasser & Price 1999). family Coenobitoidea be suppressed and that a single superfamily of hermit crabs Paguristes lymani A. Milne-Edwards & should be recognized. Martin & Davis Bouvier, 1893 (2001) accepted the arrangement proposed Known range.—Western Atlantic: south- by McLaughlin (1983). However, Mc- east of Cape Lookout, North Carolina; Flor- Laughlin & Lemaitre (2001a) recently pro- ida Keys to Swan Island, off Honduras; vided evidence to support reinstatement of through West Indies to Guyana (Williams the Coenobitoidea. Based on their results, 1984). two distinct superfamilies should be recognized with respect to hermit crab higher Paguristes moorei Benedict, 1901 classification. Known range.—Western Atlantic: off Family Diogenidae Ortmann, 1892 Cape Lookout, North Carolina; Florida Cancellus ornatus Benedict, 1901 Straits; Puerto Rico (Williams 1984). Known range.—Western Atlantic: Cape Paguristes sericeus A. Milne-Edwards, Fear, North Carolina, through eastern Gulf of Mexico; Greater and Lesser Antilles, to 1880 Bahia, Brazil (Williams 1984, Rieger Known range.—Western Atlantic: off 1998). Cape Lookout, North Carolina; West Flow- er Garden Bank, northwestern Gulf of Mex- Clibanarius vittatus (Bosc, 1802) ico, to Virgin Islands (Williams 1984). Known range.—Western Atlantic: Poto- mac River, Gunston, Virginia, to Santa Ca- Paguristes spinipes A. Milne-Edwards, tarina, Brazil, including the Gulf of Mexico 1880 (Williams 1984). Known range.—Western Atlantic: Gulf Stream south of Cape Lookout, North Car- Dardanus fucosus Biffar & Provenzano, olina; off Cape Canaveral to Florida Straits; 1972 Sarasota, Florida; Barbado3 to Pernambuco Known range.—Western Atlantic: near and Alagoas, Brazil (Williams 1984, Rieger Cape Hatteras, North Carolina, to Para, 1998). Brazil (Williams 1984, Rieger 1998). Paguristes tortugae Schmitt, 1933 Dardanus insignis (de Saussure, 1858) Known range.—Western Atlantic: reefs Known range.—Western Atlantic: off off Beaufort, North Carolina, to southern Oregon Inlet, North Carolina, to Port Aran- and southwestern Florida; Gulf of Mexico; VOLUME 116, NUMBER 1 119 through West Indies to Sao Paulo, Brazil consisting of sand, mud and shell fragments (Williams 1984, Rieger 1998). (Rieger 1998). Known range.—Western Atlantic: 40°N, Paguristes triangulatus A. Milne-Edwards southward to Rio de Janeiro, Brazil, includ- & Bouvier, 1893 ing the Caribbean Sea (Williams & Wigley 1977). Known range.—Western Atlantic: off Oregon Inlet, North Carolina, to Tortugas, ir Goreopagurus piercei (Wass, 1963) Florida; Barbados; Trinidad (Williams 1984). Pagurus piercei Wass.—Williams, 1984: 218. Petrochirus diogenes (Linnaeus, 1758) Remarks.—McLaughlin (1988) exam- Known range.—Western Atlantic: off ined a large sample of P. piercei and ob- Cape Lookout, North Carolina, to Uruguay, served several characters that excluded this including Gulf of Mexico and West Indies species from the genus Pagurus. A new ge- (Williams 1984, Rieger 1998). nus (Goreopagurus) was erected and described (McLaughlin 1988) and G. piercei was rediagnosed and illustrated. Discovery Superfamily Paguroidea Latreille, 1802 of an additional species belonging to this Family Lithodidae Samouelle, 1819 genus required McLaughlin & Haig (1995) Lithodes maja (Linnaeus, 1758) to emend and expand the original generic Known range.—Western Atlantic: West diagnosis. Greenland to Baltimore Canyon, off the Known range.—Western Atlantic: Mid- coast of Maryland; North Atlantic: East Atlantic Bight, off eastern United States, to Greenland; Iceland; Northeastern Atlantic: southeastern Florida; northern Gulf of Mex- Spitsbergen to the British Isles; the Neth- ico (McLaughlin 1988). erlands (Williams 1984, 1988; Squires 1990). • it Hemipagurus gracilis Smith, 1881 Catapagurus gracilis (Smith).—Williams, Family Paguridae Latreille, 1802 1984:484. • Anisopagurus hopkinsi Lemaitre & McLaughlin, 1996 Remarks.—Asakura (2001) removed Hemipagurus from the synonymy of Cata- Remarks.—Lemaitre & McLaughlin pagurus, rediagnosed Hemipagurus, and re- (1996:101) described and illustrated this described H. gracilis, the type species. Wil- species and discussed congeneric affinities. liams (1984) considered this species to be This species occurs at depths of 91-165 m extralimital. This species occurs at 73-418 (Lemaitre & McLaughlin 1996). m depth (Williams & Wigley 1977). Known range.—Western Atlantic: off Known range.—Western Atlantic: Mas- Georgia; Gulf of Mexico (Lemaitre & sachusetts to Barbados (Williams & Wigley McLaughlin 1996). 1977).

• Catapagurus sharreri • Iridopagurus reticulatus Garcfa-Gomez. A. Milne-Edwards, 1880 1983 Remarks.—Williams (1984) considered Remarks.—Garcia-Gomez (1983:37) de- this species to be extralimital. Asakura scribed this species and provided a species (2001) rediagnosed this genus and species. diagnosis, color description, and behavioral This species occurs at 60-882 m depth observations. This species occurs at depths (Williams & Wigley 1977) on sediments of 1—38 m (Garcfa-Gomez 1983). 120 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Known range.—Western Atlantic: North Pagurus arcuatus Squires, 1964 Carolina to Florida; Bahamas; Jamaica; Do- minican Republic; U.S. Virgin Islands; Lee- Known range.—Western Atlantic: ward Islands; Bonaire; Curasao; Belize; Greenland to off Virginia Capes (Williams Colombia; Venezuela; Suriname; French 1984). Guiana (Garcfa-Gomez 1983). • Pagurus brevidactylus (Stimpson, 1859)

ic Manucomplanus ungulatus Remarks.—Lemaitre et al. (1982) provid- (Studer, 1883) ed a species diagnosis and commented on interspecific comparisons. This species oc- Manucomplanus corallinus (Benedict).— curs from the intertidal to 50 m (Lemaitre Williams, 1984:224. et al. 1982). Remarks.—Lemaitre & McLaughlin Known range.—Western Atlantic: Ber- (1996) determined that the range of varia- muda; northeastern Florida; Bahamas to tion of specimens of the nominal species M. Brazil; Gulf of Mexico; Caribbean (Lemai- corallinus and M. ungulatus was such that tre et al. 1982). the two taxa could not be separated. Con- sequently, they placed M. corallinus in the Pagurus carolinensis McLaughlin, 1975 synonymy of M. ungulatus. The authors provided a species diagnosis and illustra- Remarks.—Lemaitre et al. (1982) provid- tions. ed species diagnosis and interspecific comparisons. Known range.—Western Atlantic: off Known range.—Western Atlantic: North Cape Hatteras, North Carolina, to Florida; Carolina to southeastern Florida (Lemaitre Gulf of Mexico; Straits of Florida; Baha- et al. 1982); Tampa Bay, Florida (Strasser mas; Eastern Atlantic: South and West Af- & Price 1999). rica (Williams 1984, Lemaitre & Mc- Laughlin 1996). Pagurus defensus (Benedict, 1892)

Pagurus acadianus Benedict, 1901 Known range.—Western Atlantic: Cape Hatteras, North Carolina, to Georgia; Dry Known range.—Western Atlantic: Straits Tortugas, Florida, to Alabama (Williams of Belle Island and Notre Dame Bay, New- 1984). foundland; Gulf of St. Lawrence to mouth of Chesapeake Bay (Williams 1984, Squires Pagurus impressus (Benedict, 1892) 1990). Known range.—Western Atlantic: off Di- Pagurus annulipes (Stimpson, 1860) amond Shoals, North Carolina, to near Cape Canaveral, Florida; Florida Bay (near Remarks.—Lemaitre et al. (1982) report- Flamingo), north to Pensacola, Florida; Port ed the depth distribution for this species as Aransas, Texas (Williams 1984). subtidal to 90 m, which represented an increase in the maximum reported depth of Pagurus longicarpus Say, 1817 occurrence. Known range.—Western Atlantic: Vine- Known range.—Western Atlantic: Minas yard Sound, Massachusetts, to south central Basin and Chignecto Bay to Hutchinson Is- Florida (Williams 1984, Lemaitre et al. land, Florida; southwestern Florida to Texas 1982). (Williams 1984, Squires 1990). VOLUME 116, NUMBER 1 121

• Pagurus maclaughlinae Garcfa-Gomez, Wass are problematic according to Lemaitre 1982 et al. (1982) who were unable to verify the accuracy of these collection data. Remarks.—Garcia-Gomez (1982:647) Known range.—Western Atlantic: North recognized and described this species. Le- Carolina to Florida; Gulf of Mexico; Carib- maitre et al. (1982) reported this species in bean coast of South America (Lemaitre et the region and provided a species diagnosis. al. 1982). Pagurus maclaughlinae occurs from the subtidal to 5 m (Garcfa-Gomez 1982). Phimochirus holthuisi (Provenzano, 1961) Known range.—Western Atlantic: Was- saw, Georgia, to Florida; Gulf of Mexico; Remarks.—McLaughlin (1981) reported Caribbean (Lemaitre et al. 1982). the bathymetric range for this species as 1— 210 m. Pagurus politus (Smith, 1882) Known range.—Western Atlantic: North Carolina to Florida; Straits of Florida and Known range.—Western Atlantic: Nova Bahama Islands; Gulf of Mexico from Flor- Scotia to off Dry Tortugas, Florida (Wil- ida to Texas; Caribbean and northern South liams 1984, Squires 1990). America from Colombia to Bahia, Brazil (McLaughlin 1981, Williams 1984, Rieger Pagurus pollicaris Say, 1817 1998). Known range.—Western Atlantic: Grand Manan, New Brunswick, to northeastern Pylopagurus discoidalis Florida; Key West, Florida, to Texas (Wil- (A. Milne-Edwards, 1880) liams 1984). Remarks.—McLaughlin & Lemaitre (2001b) rediagnosed the genus Pylopagurus Pagurus pubescens Kr0yer, 1838 and diagnosed, illustrated, and discussed Known range.—Western Atlantic: West morphological variation of all species with- Greenland; Foxe Basin; Hudson Bay to in this genus. Cape Hatteras, North Carolina; Arctic- Known range.—Western Atlantic: North North Atlantic: East Greenland; Iceland; Carolina to central Brazil, including Gulf of Spitsbergen; Barents Sea; Novaya Zemlya; Mexico and Caribbean (Williams 1984, Eastern Atlantic: Faroes and the British McLaughlin & Lemaitre 2001b). Isles (Williams 1984, Squires 1990). Rhodochirus rosaceus ir Pagurus stimpsoni (A. Milne-Edwards & Bouvier, 1893) (A. Milne-Edwards & Bouvier, 1893) Known range.—Western Atlantic: south Pagurus hendersoni Wass.—Williams, of Cape Lookout, North Carolina, to Key 1984:214. West, Florida; northwestern Gulf of Mexi- co; Grenada; Suriname; Sao Paulo and Rio Remarks.—Lemaitre et al. (1982), in Grande do Sul, Brazil (McLaughlin 1981, comparing type material of P. hendersoni Williams 1984, Rieger 1998). with specimens of P. stimpsoni, determined that the two species were conspecific, and Tomopaguropsis problematica placed P. hendersoni in the synonymy of P. (A. Milne-Edwards & Bouvier, 1893) stimpsoni. Pagurus stimpsoni occurs from the subtidal purportedly to 512 m (Wass Known range.—Western Atlantic: north- 1963), usually from the subtidal to 73 m. east of Cape Lookout, North Carolina; Depths of occurrence reported for types of southern Florida and Bahamas; Barbados; P. hendersoni (228 and 347-512 m) by off Honduras (Williams 1984). 122 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

• Tomopagurus cokeri (Hay, 1917) Infraorder Brachyura Latreille, 1802 Superfamily Dromioidea De Haan, 1833 Remarks.—McLaughlin (1981) rede- Family Dromiidae De Hann, 1833 scribed the genus and provided a species ic Cryptodromiopsis antillensis diagnosis and illustrations. This species oc- (Stimpson, 1858) curs at 44-302 m (McLaughlin 1981). Known range.—Western Atlantic: south Dromidia antillensis Stimpson.—Williams, of Cape Lookout, North Carolina; off 1984:255. northeastern Florida; Gulf of Mexico; Ca- Remarks.—McLay (1993) redefined the ribbean; off French Guiana (McLaughlin genera within the family Dromiidae. As a 1981). result, Dromidia antillensis was reassigned to the genus Cryptodromiopsis. • Tomopagurus wassi McLaughlin, 1981 Known range.—Western Atlantic: Ber- Remarks.—McLaughlin (1981) rede- muda; off Cape Hatteras, North Carolina, to scribed the genus, described this species, Rio Grande do Sul, Brazil, including Gulf and provided illustrations and comparative of Mexico and Caribbean Sea; Central At- information. This species occurs at 75-360 lantic: Saint Helena (Williams 1984, Melo m (McLaughlin 1981). et al. 1998). Known range.—Western Atlantic: south- • Dromia erythropus eastern United States; Straits of Florida; (George Edwards, 1771) Gulf of Mexico; Caribbean to northern Bra- zil (McLaughlin 1981). Remarks.—Previously, the geographic range of this species was considered to be • Family Parapaguridae Smith, 1882 outside the region, however, Williams (un- • Parapagurus pilosimanus Smith, 1879 published data) records this species from off North Carolina. Dromia erythropus in- Remarks.—Lemaitre (1989) provided a habits various types of sediments from the description, illustrations, and information intertidal to 360 m (Laughlin et al. 1982, on symbiotic associations and affinities. Melo et al. 1998) and is usually found with This species occurs at 102-3864 m, but is sponges and ascidians on its carapace most frequently found in depths of 400- (Melo et al. 1998). 1400 m (Lemaitre 1989). Known range.—Western Atlantic: Ber- Known range.—North Atlantic: south- muda; Florida to Sao Paulo, Brazil (Laugh- west of Iceland and the Faroe Islands to lin et al. 1982, Melo et al. 1998). Williams west of Ireland; Western Atlantic: Nova (unpublished) noted a specimen "Taken off Scotia to Guyana; Eastern Atlantic: Bay of North Carolina (33°48'06"N, 76°34'24"W), Biscay to Gulf of Guinea; South Atlantic: 105 m, 14 May 1981 (USNM 202800)." Tristan da Cunha (Lemaitre 1989). Hypoconcha arcuata Stimpson, 1858 • Sympagurus pictus Smith, 1883 Known range.—Western Atlantic: off Remarks.—Lemaitre (1989) provided a Cape Lookout, North Carolina, to west description, illustrations, and information Florida; St. Thomas, U.S. Virgin Islands; on symbiotic associations and affinities. Suriname to Sao Paulo, Brazil (Williams This species occurs at 180-2322 m, but is 1984, Melo et al. 1998). most frequently found at depths of 200 to 800 m (Lemaitre 1989). * Hypoconcha parasitica Known range.—Western Atlantic: off (Linnaeus, 1763) Long Island, New York to off French Gui- Hypoconcha sabulosa (Herbst).—Williams, ana (Lemaitre 1989). 1984:258. VOLUME 116, NUMBER 1 123

Remarks.—Holthuis and Manning (1987) Superfamily Raninoidea De Haan, 1841 concluded that Hypoconclia parasitica was Family Raninidae De Haan, 1841 the oldest available name for the species Remarks.—Guinot (1993) subdivided the formerly known as H. sabulosa. family Raninidae into six subfamilies: Ran- Known range.—Western Atlantic: off ininae De Haan, 1841, Notopodinae Serene Cape Hatteras, North Carolina, through & Umali, 1972, Symethinae Goeke, 1981. Gulf of Mexico to Sao Paulo, Brazil (Wil- Raninoidinae Lorenthey & Beurlen, 1929, liams 1984, Melo et al. 1998). Lyreidinae Guinot, 1993, and Cyrtorhinae Guinot, 1993. Three subfamilies (Ranini- Hypoconcha spinosissima Rathbun, 1933 nae, Notopodinae, and Symethinae) were recognized in Williams (1984). Known range.—Western Atlantic: off Cape Hatteras, North Carolina, to Gulf of Mexico, off Mississippi River delta; Yuca- * Subfamily Lyreidinae Guinot, 1993 • Lysirude nitidus tan; Jamaica (Williams 1984). (A. Milne-Edwards, 1880)

Superfamily Homoloidea De Haan, 1839 Remarks.—Diagnostic characteristics Family Homolidae De Haan, 1839 and congeneric comparisons are provided in • Homola minima Guinot & Richer de Goeke (1980, 1985) and Tucker (1998). Forges, 1995 This species is found on soft mud bottoms at depths of 119-823 m (Powers 1977, as Homola barbata (Fabricius).—Williams, Lyreidus bairdii). 1984:261 (in part). Known range.—Western Atlantic: Mas- sachusetts; Gulf of Mexico; Greater Antil- Remarks.—Guinot & Richer de Forges les to Venezuela; Suriname (Goeke 1980). (1995:326) described this species, provided photographs, and discussed congeneric ^r Subfamily Notopodinae Serene & comparisons. Homola minima occurs at Umali, 1972 depths of 55-690 m (Guinot & Richer de Forges 1995, Martin & Zimmerman 2001). Remarks.—Placement of Ranilia in sub- Homola barbata, formerly considered to be family Notopodinae was supported by a wide-ranging species (Williams 1984), is Tucker's (1998) phylogenetic study. now known to occur only in the Mediter- ranean Sea and also possibly in the adjoin- Ranilia constricta ing eastern Atlantic (Guinot & Richer de (A. Milne-Edwards, 1880) Forges 1995). Known range.—Western Atlantic: south- Known range.—Western Atlantic: Mar- east of Cape Fear, North Carolina; Palm tha's Vineyard, Massachusetts, to Rio Beach, Florida, to Florida Straits and Yu- Grande do Sul, Brazil; Bahamas (Guinot & catan Channel; Cuba; off Barbados (Wil- Richer de Forges 1995, Martin & Zimmer- liams 1984); Amapa, Rio de Janeiro to Rio man 2001). Grande do Sul, Brazil (Melo et al. 1998); Central Atlantic: Ascension Island; Eastern Family Latreilliidae Stimpson, 1858 Atlantic: Senegal to Congo; Annobon Is- Latreillia manningi Williams, 1982 land (Williams 1984).

Known range.—Western Atlantic: Nan- Ranilia muricata H. Milne Edwards, 1837 tucket Shoals, off Massachusetts, to off Ha- vana, Cuba; Venezuela; Central Atlantic: Known range.—Western Atlantic: off Ascension Island (Williams 1984). Cape Lookout, North Carolina, to north- 124 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON western Gulf of Mexico; Colombia (Wil- vares 1991a, 1993, 1996). Tavares (1996) liams 1984); Pernambuco, Brazil (Melo et conducted a revision of the family Cyclo- al. 1998). dorippidae.

* Subfamily Raninoidinae Lorenthey & Clythrocerus granulatus (Rathbun, 1898) Beurlen, 1929 Known range.—Western Atlantic: south- Remarks.—Placement of Raninoides in east of Cape Lookout, North Carolina; subfamily Raninoidinae was supported by Honduras; southern Florida, through Antil- Tucker's (1998) phylogenetic study. les, to Venezuela and Trinidad (Williams 1984), and southward to Rio Grande do Raninoides loevis (Latreille, 1825) Sul, Brazil (Melo et al. 1998).

Remarks.—This species was listed under • Clythrocerus nitidus subfamily Ranininae in Williams (1984). (A. Milne-Edwards, 1880) Known range.—Western Atlantic: south of Cape Hatteras, North Carolina, to Sao Remarks.—Tavares (1996) included a de- Paulo, Brazil, including Gulf of Mexico, scription and illustrations, and discussed southern Caribbean Sea and Leeward Is- morphological variation for this species. lands (Williams 1984, Melo et al. 1998). This species occurs at 12-531 m (Tavares 1996). * Family Symethidae Goeke, 1981 Known range.—Western Atlantic: South Carolina; Florida; Barbados; Grenada (Ta- Remarks.—This taxon was previously vares 1996). considered a subfamily of the Raninidae (e.g., Williams 1984, Guinot 1993). Tucker * Deilocerus perpusillus (Rathbun, 1901) (1998) considered the subfamily Symethi- nae sufficiently distinct to warrant elevation Clythrocerus perpusillus Rathbun.—Wil- of the subfamily to the rank of family. liams, 1984:260. Remarks.—As a result of his review of Symethis variolosa (Fabricius, 1793) the New World cyclodorippoid crabs, Ta- Known range.—Western Atlantic: south- vares (1993) identified and described four east of Cape Lookout, North Carolina, new genera, including Deilocerus. Clythro- through western Gulf of Mexico to Sao cerus perpusillus was designated the type Paulo, Brazil, and Fernando de Noronha species of this new genus. (Williams 1984, Melo et al. 1998). Known range.—Western Atlantic: North Carolina to Georgia; Bahama Banks; Gulf * Superfamily Cyclodorippoidea of Mexico; Puerto Rico; Barbados; Amapa Ortmann, 1892 to Rio Grande do Sul, Brazil (Williams 1984, Tavares 1996, Melo et al. 1998). Remarks.—Tavares (1991a) proposed the use of the superfamilial name Cyclodorip- Superfamily Dorippoidea MacLeay, 1838 poidea to replace Tymoloidea. Martin & Family Dorippidae MacLeay, 1838 Davis (2001) accepted this arrangement. Subfamily Ethusinae Guinot, 1977 * Ethusa americana A. Milne-Edwards, * Family Cyclodorippidae Ortmann, 1892 1880 Family Tymolidae.—Williams, 1984:259. Ethusa mascarone americana A. Milne-Ed- wards.—Williams, 1984:269. Remarks.—Family Tymolidae is placed in the synonymy of Cyclodorippidae (Ta- Remarks.—Hendrickx (1989) elevated VOLUME 116, NUMBER 1 125 the subspecies E. m. americana to full spe- mann (2000) supported the conclusions of cies status. Ethusa americana occurs only Bellwood (1996). in the western Atlantic; references of this species at eastern Pacific localities refer to Acanthocarpus alexandri Stimpson, 1871 E. panamensis Finnegan. Ethusa mascaro- Known range.—Western Atlantic: ne (Herbst) occurs in the Mediterranean Georges Bank, off Massachusetts, to west (Manning & Holthuis 1981). coast of Florida; Puerto Rico to Grenadines; Known range.—Western Atlantic: south Rio de Janeiro (Williams 1984) to Rio of Cape Lookout, North Carolina, to Gulf Grande do Sul, Brazil (Melo et al. 1998). of Mexico and West Indies; Maranhao to Rio de Janeiro, Brazil (Williams 1984, Calappa flammea (Herbst, 1794) Melo et al. 1998). Known range.—Western Atlantic: Woods Ethusa microphthalma Smith, 1881 Hole region, Massachusetts, to Florida Keys; Gulf coast of United States and Mex- Known range.—Western Atlantic: off ico; Bahamas; Bermuda (Williams 1984). Martha's Vineyard, Massachusetts, to Cuba; throughout Gulf of Mexico (Williams Calappa ocellata Holthuis, 1958 1984); Sao Paulo, Brazil (Melo et al. 1998). Known range.—Western Atlantic: Ber- Ethusa tenuipes Rathbun, 1897 muda; Cape Hatteras, North Carolina, to Rio de Janeiro, Brazil (Williams 1984). Known range.—Western Atlantic: off Cape Lookout, North Carolina; East Florida Calappa sulcata Rathbun, 1898 to Gulf of Mexico, east of Mississippi River delta; Cuba (Williams 1984); Rio de Janeiro Remarks.—This species occurs on sand, to Sao Paulo, Brazil (Melo et al. 1998). mud, and calcareous algal bottoms from shallow depths to 200 m (Melo et al. 1998). Known range.—Western Atlantic: Cape Superfamily Calappoidea H. Milne Hatteras, North Carolina, through Gulf of Edwards, 1837 Mexico to Parana, Brazil (Williams 1984, Family Calappidae H. Milne Edwards, Melo et al. 1998). 1837 Remarks.—Bell wood (1996) evaluated * Calappa tortugae Rathbun, 1933 phylogenetic relationships of four subfam- Calappa angusta A. Milne-Edwards.—Wil- ilies within the Calappidae (Calappinae, liams, 1984:273 (in part). Matutinae, Orithyiinae, and Hepatinae) using cladistic analysis. She rejected the Remarks.—Williams & Child (1989) de- monophyly of an expanded Calappidae, but termined that "Calappa angustaas pre- demonstrated support for the monophyly of viously understood, was poorly defined and each component taxon and proposed ele- actually comprised a complex of species. vating the four subfamilies to family status. The next available name for species of Ca- Additionally, Bellwood (1996) reassigned lappa in the western Atlantic, Calappa these families to different superfamilies, saussurei tortugae Rathbun, was removed with Calappidae and Hepatidae remaining from synonymy and elevated to full species in the superfamily Calappoidea, Matutidae rank (Williams & Child 1989). Species di- placed in the superfamily Leucosioidea, and agnosis, illustrations and measurements Orithyiidae placed in the superfamily Do- were provided in Williams & Child (1989). rippoidea. Based on fossil evidence (cara- This species occurs at 13-238 m (Williams pace morphology), Schweitzer and Feld- & Child 1989). 126 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Known range.—Western Atlantic: North tions, measurements and color description. Carolina to off Venezuela, including Gulf This species occurs at 31-188 m depth, of Mexico, Caribbean, and Leeward Islands rarely to 640 m (Williams & Child 1989). (Williams & Child 1989). Known range.—Western Atlantic: off Cape Lookout, North Carolina, through Ba- ^k Cryptosoma balguerii hamas; eastern Gulf of Mexico; and Suri- (Desbonne, 1867) name (Williams & Child 1989). Cycloes bairdii (Stimpson).—Williams, 1984:278. * Family Hepatidae Stimpson, 1871 Remarks.—Galil & Clark (1996) report- Remarks.—Species in the genera Hepa- ed that populations of C. bairdii occurred tus and Osachila were previously assigned exclusively in the eastern Pacific (Baja Cal- to the subfamily Matutinae (Williams ifornia to Costa Rica); previous records of 1984). Based on results of a phylogenetic C. bairdii from Costa Rica to Ecuador were study by Bellwood (1996) and corroborat- actually those of a new Cryptosoma spe- ing fossil evidence (carapace morphology) cies, whereas records reported as C. bairdii of Schweitzer & Feldmann (2000), these from Atlantic localities pertained to C. bal- genera are now placed in the family He- guerii (a nominal species previously con- patidae. sidered to be a subjective synonym of Cy- cloes bairdii). Cryptosoma balguerii occurs Hepatus epheliticus (Linnaeus, 1763) in shallow waters to 230 m (Galil & Clark Known range.—Western Atlantic: Ches- 1996). apeake Bay to western Gulf of Campeche, Known range.—Western Atlantic: North Mexico; Cuba; Jamaica; Dominican Repub- Carolina and Bermuda to Espirito Santo, lic (Williams 1984). Brazil (Galil & Clark 1996). Hepatus pudibundus (Herbst, 1785) • it Cyclozodion angustum (A. Milne-Edwards, 1880) Known range.—Western Atlantic: Geor- gia to Rio Grande do Sul, Brazil (Williams Calappa angusta A. Milne-Edwards.—Wil- 1984, Melo et al. 1998). liams, 1984:273 (in part). Remarks.—Williams & Child (1989) Osachila semilevis Rathbun, 1916 concluded that Calappa angusta, as de- Known range.—Western Atlantic: off scribed by A. Milne-Edwards, was generi- Beaufort, North Carolina, to northwest cally misplaced. They described the genus Florida (Williams 1984). Cyclozodion, of which C. angustum (A. Milne-Edwards) is the type species, and Osachila tuberosa Stimpson, 1871 provided illustrations, measurements, and comparative information. This species oc- Known range.—Western Atlantic: off curs at 95-421 m (Williams & Child 1989). Cape Hatteras, North Carolina, to northwest Known range.—Western Atlantic: off Florida and Yucatan Channel (Williams Cape Canaveral, Florida, to Isla Providen- 1984); Rio de Janeiro and Rio Grande do cia, Colombia; Guyana (Williams & Child Sul, Brazil (Melo et al. 1998). 1989). Superfamily Leucosioidea Samouelle, • Cyclozodion tuberatum Williams & 1819 Child, 1989 Family Leucosiidae Samouelle, 1819 Remarks.—Williams & Child (1989:112) Remarks.—Leucosiidae was previously described this species and provided illustra- assigned to the superfamily Calappoidea VOLUME 116, NUMBER 1 127

(Williams 1984). Based on results of a phy- Remarks.—Galil (2000) removed I. in- logenetic study of the Calappidae, Bell- termedia from the genus Iliacantha and wood (1996) placed the family Leucosiidae placed it in the newly erected genus Acan- in the superfamily Leucosioidea. thilia Galil; generic description and species redescription are provided. This species oc- Subfamily Ebaliinae Stimpson, 1871 curs at depths of 10-329 m. Ebalia cariosa (Stimpson, 1860) Known range.—Western Atlantic: North Known range.—Western Atlantic: Bogue Carolina to Brazil (Galil 2000). Sound near Beaufort, North Carolina, to west Florida; western Gulf of Mexico; Ja- Callidactylus asper Stimpson, 1871 maica; northeastern South America to Sao Known range.—Western Atlantic: south Paulo, Brazil (Williams 1984). of Cape Lookout, North Carolina, through southeastern Gulf of Mexico to Panama, Ebalia stimpsonii A. Milne-Edwards, 1880 and southeastward to Alagoas, Brazil (Wil- Known range.—Western Atlantic: south- liams 1984). east of Cape Lookout, North Carolina; west Florida to Barbados (Williams 1984); Ama- Iliacantha subglobosa Stimpson, 1871 pa to Sao Paulo, Brazil (Melo et al. 1998). Known range.—Western Atlantic: off Speloeophorus nodosus (Bell, 1855) Cape Hatteras, North Carolina, to northwest Known range.—Western Atlantic: Flori- Florida; through eastern Gulf of Mexico and Caribbean Sea, south to Alagoas, Brazil da; West Indies (Williams 1984); Maranhao (Williams 1984). to Rio de Janeiro, Brazil (Melo et al. 1998).

Speloeophorus pontifer (Stimpson, 1871) Superfamily Majoidea Samouelle, 1819 Known range.—Western Atlantic: south- Remarks.—Recent investigations of in- east of Cape Lookout and off Beaufort, trarelationships within the Majoidea include North Carolina, to west Florida; West In- those of Drach & Guinot (1983), Griffin & dies to Barbados (Williams 1984). Tranter (1986), Clark & Webber (1991), Guinot & Richer de Forges (1997), Guinot Subfamily Iliinae Stimpson, 1871 & Bouchard (1998), and Pohle & Marques Myropsis quinquespinosa Stimpson, 1871 (2000). Although all subfamilies within the Known range.—Western Atlantic: south superfamily have not been considered in a of Martha's Vineyard, Massachusetts, to phylogenetic framework, previously recog- Suriname, including Gulf of Mexico and nized subfamilies within the family Majidae Caribbean Sea (Williams 1984). have been elevated to the level of family (Hendrickx 1995). Martin & Davis (2001) Persephona mediterranea (Herbst, 1794) also adopted this arrangement. However, results of a recent phylogenetic analysis Known range.—Western Atlantic: New based on larval characters (Pohle & Jersey to Rio Grande do Sul, Brazil, in- Marques 2000) supported monophyly only cluding Gulf of Mexico and Caribbean Sea of the Oregoniidae, Majidae and Inachidae. (Williams 1984, Melo et al. 1998).

Subfamily Leucosiinae Samouelle, 1819 * Family Epialtidae MacLeay, 1838 :k Acanthilia intermedia (Miers, 1886) Remarks.—All species listed below were Iliacantha intermedia Miers.—Williams, previously placed in subfamily Epialtinae 1984:290. of the Majidae in Williams (1984). 128 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Epialtus dilatatus A. Milne-Edwards, 1878 Podochela gracilipes Stimpson, 1871

Known range.—Western Atlantic: off Known range.—Western Atlantic: off Beaufort Inlet and New River, North Car- Cape Lookout, North Carolina, to Rio olina; southwest Florida; Yucatan; Bahamas Grande do Sul, including Gulf of Mexico to St. Thomas (Williams 1984). and Caribbean Sea (Williams 1984, Melo 1998). Sphenocarcinus corrosus A. Milne-Edwards, 1875 Podochela riisei Stimpson, 1860 Known range.—Western Atlantic: Ber- Remarks.—Tavares (1991b) provided a muda; North Carolina to Campeche, Mex- generic revision and diagnosis. ico; Antilles; Rio de Janeiro, Brazil (Wil- Known range.—Western Atlantic: off liams 1984, Melo 1998). Cape Lookout, North Carolina; Gulf of Mexico to Barbados (Williams 1984). Podochela sidneyi Rathbun, 1924

* Family Inachidae MacLeay, 1838 Known range.—Western Atlantic: off Cape Hatteras, North Carolina, to Veracruz, Remarks.—Elevation of subfamily Ina- Mexico; northwestern Cuba; Yucatan Chan- chinae to family level was supported by the nel (Williams 1984). cladistic analysis of Clark & Webber (1991). Other morphological evidence Rochinia crassa (Guinot & Bouchard 1998) supported this (A. Milne-Edwards, 1879) conclusion. All species of the genera An- omalothir, Metoporhaphis, Podochela, and Known range.—Western Atlantic: Nan- Stenorhynchus (listed below) were previ- tucket Shoals, Massachusetts, to Gulf of ously placed in subfamily Inachinae of the Mexico, off southern Texas; northern Cuba; Majidae in Williams (1984). The genus Ro- west of Cabo de la Vela, Colombia; off chinia was previously placed in the subfam- French Guiana (Williams 1984). Recently, ily Pisinae of the Majidae (Williams 1984, a male measuring 89.2 mm carapace width Griffin & Tranter 1986). Cladistic relation- and 89.5 mm carapace length was caught ships hypothesized by Clark & Webber off Lunenburg, Nova Scotia, at 243 m (Mo- (1991) indicated that Rochinia should be riyasu et al. 2001). This first recorded oc- placed in the Inachidae. Tavares (1991b) currence of this species in Canadian waters conducted a generic revision of Rochinia. constitutes a northern range extension from the previous northernmost occurrence on Nantucket Shoals, Massachusetts (Williams Anomalothir furcillatus (Stimpson, 1871) 1984). Moriyasu et al. (2001) considered Known range.—Western Atlantic: off this as a stray occurrence of this species in Cape Lookout, North Carolina, through Canadian waters. eastern Gulf of Mexico; West Indies to Gre- nada (Williams 1984). Rochinia tanneri (Smith, 1883) Known range.—Western Atlantic: off Metoporhaphis calcarata (Say, 1818) Martha's Vineyard, Massachusetts, to Straits of Florida (Williams 1984). Known range.—Western Atlantic: off Cape Hatteras, North Carolina, to Rio de Rochinia umbonata (Stimpson, 1871) Janeiro, Brazil, including Gulf of Mexico and Caribbean Sea (Williams 1984, Melo Known range.—Western Atlantic: south- 1998). east of Cape Lookout, North Carolina, VOLUME 116, NUMBER 1 129 through eastern and northern Gulf of Mex- Banks to Sao Paulo, Brazil, and Fernando ico to northeast of Nicaragua; through West de Noronha Archipelago and Rocas Atoll, Indies to St. Vincent (Williams 1984). Brazil (Williams 1984, Melo 1998).

Stenorhynchus seticornis (Herbst, 1788) Anasimus latus Rathbun, 1894 Remarks.—Goeke (1989) determined Known range.—Western Atlantic: off that two co-occurring species were con- Cape Lookout, North Carolina, to Amapa, fused under the name S. seticornis. He re- Brazil, including Gulf of Mexico and An- stricted the specific description of S. seti- tilles (Williams 1984, Melo 1998). cornis, redescribed the species, and selected a neotype. This species occurs at 1—366 m Arachnopsis filipes Stimpson, 1871 (Goeke 1989). Known range.—Western Atlantic: Ber- Known range.—Western Atlantic: south- muda; Cape Fear, North Carolina, to Rio east of Cape Hatteras, North Carolina; Gulf Grande do Sul, Brazil, including Gulf of of Mexico, off northwest Florida, through Mexico, Antilles, and northern South Antilles to Rio Grande do Norte, Brazil America (Williams 1984, Goeke 1989, (Williams 1984, Melo 1998). Melo 1998). Batrachonotus fragosus Stimpson, 1871 • Stenorhynchus yangi Goeke, 1989 Known range.—Western Atlantic: Cape Remarks.—Goeke (1989:631) described Hatteras, North Carolina, to southern and this species and provided a diagnosis, illus- western Florida; West Indies to Barbados trations, color description, and discussed (Williams 1984). morphological variation. This species occurs at 31-365 m (Goeke 1989). Collodes robustus Smith, 1883 Known range.—Western Atlantic: off Known range.—Western Atlantic: Cape Martha's Vineyard, Massachusetts, to Suri- Cod, Massachusetts, to southeast of Cape name, including Gulf of Mexico (Goeke Lookout, North Carolina (Williams 1984). 1989). Collodes trispinosus Stimpson, 1871 * Family Inachoididae Dana, 1851 Known range.—Western Atlantic: near Remarks.—Drach & Guinot (1983) pro- Cape Hatteras, North Carolina, to southern posed elevating the Inachoidinae to family and western Florida (Williams 1984); Ama- level. This decision was corroborated with pa, Rio de Janeiro, and Sao Paulo, Brazil morphological evidence (Guinot & Richer (Melo 1998). de Forges 1997). The genera included within this family are based on the recommen- Euprognatha rastellifera Stimpson, 1871 dation of Guinot & Richer de Forges (1997). All species listed below were pre- Known range.—Western Atlantic: off viously placed in the subfamily Inachinae Georges Bank to Uruguay, including Antil- of the Majidae in Williams (1984). les (Williams 1984, Melo 1998 as E. acuta).

Aepinus septemspinosus Inachoides forceps A. Milne-Edwards, (A. Milne-Edwards, 1879) 1879 Known range.—Western Atlantic: south Known range.—Western Atlantic: south- of Cape Lookout, North Carolina; southeast of Cape Lookout, North Carolina; west west of Cape San Bias, Florida; Bahama coast of Florida to Rio de Janeiro, Brazil, 130 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON including Antilles; the Guianas (Williams ern Florida to Alligator Harbor, Florida 1984, Melo 1998). (Williams 1984); Amapa to Maranhao, Bra- zil (Melo 1998). • Pyromaia arachna Rathbun, 1924 Remarks.—Lemaitre et al. (2001) provid- Macrocoeloma eutheca (Stimpson, 1871) ed information on distinguishing characteristics and congeneric comparisons. This Known range.—Western Atlantic: south- species occurs on mud, mud-sand, and east of Cape Lookout, North Carolina; off mud-shell sediments at 183-324 m depth northwest Florida through Bahama Banks (Powers 1977). and West Indies; Central America; Maran- Known range.—Western Atlantic: South hao to Espirito Santo, Brazil (Williams Carolina to Gulf of Mexico (Powers 1977, 1984, Melo 1998). Lemaitre et al. 2001). Macrocoeloma trispinosum Pyromaia cuspidata Stimpson, 1871 (Latreille, 1825) Known range.—Western Atlantic: off Macrocoeloma trispinosum, variety of Cape Lookout, North Carolina, to west Rathbun, 1925 Florida; Cuba; Yucatan Channel to Nicara- Macrocoeloma trispinosum nodipes gua (Williams 1984). (Desbonne, 1867)

ir Family Mithracidae Balss, 1929 Remarks.—Williams (1984:328) listed these nominal taxa together in one species Remarks.—All species of Hemus, Ma- account. No further investigation of these crocoeloma, Microphrys, Mithrax, Mithra- taxa has been conducted, and further revi- culus, and Stenocionops (listed below) were sion is needed. Until further results are previously placed in the subfamily Mithra- available, these taxa are considered as they cinae of the Majidae in Williams (1984). were in Williams (1984). Based on results of his investigation of Known range (for M. trispinosum (La- western Atlantic Mithrax crabs, Wagner treille, 1825).—Western Atlantic: Beaufort, (1990) considered the morphological evi- North Carolina, to Alligator Harbor, Flori- dence sufficient to warrant recognition of da; Yucatan; West Indies to Sao Paulo, Bra- distinct genera instead of two subgenera zil, and Fernando de Noronha Archipelago within Mithrax', Mithraculus was elevated (Williams 1984, Melo 1998). to full generic status.

Hemus cristulipes A. Milne-Edwards, Microphrys antillensis Rathbun, 1920 1875 Known range.—Western Atlantic: Cape Known range.—Western Atlantic: off Hatteras to Cape Fear, North Carolina; An- Cape Lookout, North Carolina; South Car- tilles; Paraiba to Rio de Janeiro, Brazil olina; northwest Gulf of Mexico and Yu- (Williams 1984, Melo 1998). catan, through West Indies to Rio de Janei- ro, Brazil, and Fernando de Noronha Ar- Microphrys bicornutus (Latreille, 1825) chipelago (Williams 1984, Melo 1998). Known range.—Western Atlantic: Ber- Macrocoeloma camptocerum muda; near Beaufort, North Carolina, (Stimpson, 1871) through Gulf of Mexico to Rio Grande do Known range.—Western Atlantic: Beau- Sul, Brazil; Fernando de Noronha Archi- fort Harbor, North Carolina; around south- pelago, Brazil (Williams 1984, Melo 1998). VOLUME 116, NUMBER 1 131

* Mithraculus forceps Known range.—Western Atlantic: North (A. Milne-Edwards, 1875) (?) and South Carolina to Nicaragua, Mithrax (Mithraculus) forceps (Milne Ed- through West Indies to Barbados; Venezue- la (Williams 1984, Wagner 1990). wards).—Williams, 1984:337.

Known range.—Western Atlantic: Ber- * Mithrax verrucosus H. Milne Edwards, muda; Cape Hatteras, North Carolina, 1832 through Gulf of Mexico and Antilles to Sao Paulo, Brazil; Fernando de Noronha Archi- Mithrax (Mithrax) verrucosus H. Milne Ed- pelago and Rocas Atoll, Brazil (Williams wards.—Williams, 1984:336. 1984, Wagner 1990, Melo 1998). Known range.—Western Atlantic: Charleston, South Carolina; Florida; Cam- * Mithrax cornutus de Saussure, 1857 peche Banks; Curacao; Venezuela; West In- Mithrax (Mithrax) acuticomis Stimpson.— dies to Fernando de Noronha Archipelago Williams, 1984:332. and Rocas Atoll, Brazil (Williams 1984. Remarks.—Wagner (1990) determined Wagner 1990, Melo 1998). that M. coniutus and M. acuticomis were conspecific. Mithrax coniutus was original- Stenocionops furcata coelata ly regarded as extralimital by Williams (A. Milne-Edwards, 1878) (1984:484). This species occurs between Known range.—Western Atlantic: Beau- 20-458 m (Wagner 1990). fort, North Carolina, to northwest Florida Known range.—Western Atlantic: Ber- and Alabama; Yucatan Channel; West In- muda; off Cape Lookout, North Carolina; dies to Barbados (Williams 1984). east and west coasts of Florida, through Gulf of Mexico, Yucatan Channel, and West Indies to Rio de-Janeiro, Brazil (Wil- • Stenocionops furcata flircata liams 1984, Wagner 1990, Melo 1998). (Olivier, 1791) Remarks.—This subspecies was only ref- * Mithrax hispidus (Herbst, 1790) erenced in the remarks section of S. f coe- Mithrax {Mithrax) hispidus (Herbst).—Wil- lata in Williams (1984). It is unclear why liams, 1984:333. S. f. fiircata was not given a full account Mithrax (.Mithrax) pleuracanthus Stimp- since its geographic range and depth distri- son.—Williams, 1984:334. bution (shallow water to 64 m) were clearly within the limits of coverage. Much con- Remarks.—Wagner (1990) placed M. fusion suiTounds identification of members pleuracanthus in the synonymy of M. his- of this species group, and more research is pidus. needed (D. Felder, pers. comm.). Until such Known range.—Western Atlantic: Dela- time, both subspecies are considered valid ware Bay; Bermuda; Beaufort, North Car- as in Williams (1984). olina; Charleston, South Carolina; Georgia to Pensacola, Florida; northwestern Gulf of Known range.—Western Atlantic: Geor- Mexico to Yucatan Channel; Bahamas and gia to Rio Grande do Sul. Brazil, including Florida Keys through West Indies; Colom- Antilles and Colombia (Williams 1984. bia to Sao Paulo, Brazil (Williams 1984, Melo 1998). Wagner 1990, Melo 1998). Stenocionops spinimana (Rathbun. 1892) * Mithrax spinosissimus (Lamarck, 1818) Known range.—Western Atlantic: off Mithrax (Mithrax) spinosissimus (La- Cape Hatteras. North Carolina, to Florida marck).—Williams, 1984:335. Straits; Gulf of Mexico, off Mobile Bay, 132 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Alabama, and east of Chandeleur Island, off Atlantic: between Greenland and Iceland; Mississippi (Williams 1984); Sao Paulo, Spitsbergen; Kara Sea; Eastern Atlantic: Brazil (Melo 1998). through British Isles and northwest France (Williams 1984, Squires 1990). • Stenocionops spinosissima (de Saussure, 1857) Hyas coarctatus coarctatus Leach, 1815 Remarks.—This species was considered extralimital by Williams (unpublished), but Known range.—Western Atlantic: Hud- reported occurrence off North Carolina son Bay and Greenland to North Carolina; (Powers 1977) indicated that this species Arctic-Eastern Atlantic: Murman Sea to should be included. This species occurs at Iceland and the British Isles (Williams depths of 46-480 m on mud and sand sed- 1984, Squires 1990). iments (Powers 1977, Melo 1998), with its center of distribution in the Gulf of Mexico * Family Pisidae Dana, 1851 at 110-183 m (Powers 1977). Known range.—Western Atlantic: off Remarks.—The following six species North Carolina; south and southwest Flor- (listed below) were previously placed in the ida; off Texas and east coast of Mexico; subfamily Pisinae of the Majidae in Wil- north coast of Cuba; Haiti; Guadeloupe; liams (1984). Dominica; Rio Grande do Norte to Rio Grande do Sul, Brazil; Fernando de Noro- Coelocerus spinosus A. Milne-Edwards, nha Archipelago, Brazil (Powers 1977, 1875 Melo 1998). Known range.—Western Atlantic: off * Family Oregoniidae Garth, 1958 Cape Fear, North Carolina, to Cape Can- averal, Florida; west Florida to east of Mis- Remarks.—Family level status was sup- sissippi River delta (Williams 1984). ported by a cladistic analysis (Clark & Webber 1991) and morphological evidence (Guinot & Bouchard 1998). In their classi- Libinia dubia H. Milne Edwards, 1834 fication, Martin & Davis (2001) recognized Known range.—Western Atlantic: Cape this taxon at the subfamily level. All species Cod, Massachusetts, to southern Texas; Ba- listed below were previously placed in the hamas; Cuba (Williams 1984). subfamily Oregoniinae of the Majidae in Williams (1984). Libinia emarginata Leach, 1815 Chionoecetes opilio opilio (O. Fabricius, 1788) Known range.—Western Atlantic: Prince Edward Island and Nova Scotia to western Known range.—Western Atlantic: Green- Gulf of Mexico (Williams 1984, Squires land south to St. Lawrence estuary and Gulf 1990). of Maine; Arctic-North Pacific: Point Barrow, Alaska, and northeastern Siberia through Ber- Nibilia antilocapra (Stimpson, 1871) ing Strait to Alaskan Peninsula and Aleutian chain; Eastern Pacific: Kamchatka; Okhotsk Known range.—Western Atlantic: off Sea southward to Japan (Williams 1984). Cape Hatteras, North Carolina, to Gulf of Mexico just east of Mississippi River delta; Hyas araneus (Linnaeus, 1758) Gulf of Campeche; Antilles; off Guyana; Known range.—Western Atlantic: West Rio Grande do Norte and Rio Grande do Greenland to Rhode Island; Arctic-North Sul, Brazil (Williams 1984, Melo 1998). VOLUME 116, NUMBER 1 133

Pelia mutica (Gibbes, 1850) (Chiong & Ng 1998). Generic comparisons, species redescription and illustrations were Known range.—Western Atlantic: Buz- also provided (Chiong & Ng 1998). zards Bay and Vineyard Sound, Massachu- Known range.—Western Atlantic: south- setts, to Texas; Cuba; Puerto Rico; St. east of Cape Lookout, North Carolina; east Thomas, U.S. Virgin Islands (Williams central Florida; south of Cape San Bias, 1984). Florida, to St. Thomas, U.S. Virgin Islands; through Antilles; Maranhao to Rio de Ja- * Family Tychidae Dana, 1851 neiro, Brazil (Williams 1984, Melo 1998). Remarks.—All species listed below were previously placed in the subfamily Tychi- Heterocrypta granulata (Gibbes, 1850) nae of the Majidae in Williams (1984). Known range.—Western Atlantic: from Nantucket Sound, Massachusetts, around Pitho Iherminieri (Schramm, 1867) peninsular Florida to southern Texas; Known range.—Western Atlantic: off through West Indies to Trinidad; Ceara to Beaufort Inlet, North Carolina, to west Parana, Brazil (Williams 1984, Melo 1998). Florida; Veracruz, Mexico; West Indies to Sao Paulo, Brazil (Williams 1984). Mesorhoea sexspinosa Stimpson, 1871

Tyche emarginata White, 1847 Known range.—Western Atlantic: southeast of Cape Lookout, North Carolina; off Known range.—Western Atlantic: off northwest Florida to Flanagan Passage, Vir- Beaufort Inlet, North Carolina, through Ba- gin Islands (Williams 1984). hamas to west coast of Florida (Williams 1984); Antilles; Rio Grande do Norte, Bra- * Parthenope agona (Stimpson, 1871) zil (Melo 1998). Parthenope (.Parthenope) agona (Stimp- Superfamily Parthenopoidea MacLeay, son).—Williams, 1984:342. 1838 Known range.—Western Atlantic: off Family Parthenopidae MacLeay, 1838 Cape Hatteras and Cape Lookout, North Subfamily Parthenopinae MacLeay, 1838 Carolina; east central Florida; Gulf of Mex- Remarks.—Ng & Rodriguez (1986) did ico (Pensacola, Florida to near Ft. Myers, not recognize subgenera within the genus Florida), through Florida Straits, West In- Parthenope', Platylambrus was assigned dies and Caribbean Sea to Parana, Brazil full generic status. (Williams 1984, Melo 1998).

* Celatopesia concava (Stimpson, 1871) * Platylambrus fraterculus (Stimpson, 1871) Cryptopodia concava Stimpson.—Wil- liams, 1984:346. Parthenope (Platylambrus) fraterculus (Stimpson).—Williams, 1984:343. Remarks.—Chiong & Ng (1998) determined that the American species of the ge- Known range.—Western Atlantic: off nus Cryptopodia, including C. concava, dif- Cape Fear, North Carolina; east central fered markedly in carapace appearance Florida southward; Gulf of Mexico (off from that of the Indo-West Pacific species. Cape San Bias, Florida) to Florida Straits; Celatopesia was described, and the Amer- off Cape Catoche, Yucatan, Mexico; ican species formerly included in Crypto- through West Indies to Rio Grande do Sul, podia were referred to the new genus Brazil (Williams 1984, Melo 1998). 134 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

it Platylambrus granulata Edwards, 1862, Cancer, sensu stricto (Kingsley, 1879) Linnaeus, 1758) within the genus Cancer. Williams (1984) adopted this classification. Parthenope (Platylambrus) granulata Schweitzer & Feldmann (2000) reevaluat- (Kingsley).—Williams, 1984:344. ed the Cancridae and elevated the subgen- Known range.—Western Atlantic: Ber- era of Nations (1975) to full generic status, muda; off Cape Hatteras, North Carolina, again basing their conclusions only on fos- southward around Florida into Gulf of sil evidence. This classification has not Mexico to Louisiana; Bahfa Honda, Cuba gained acceptance with researchers working (?); St. Thomas, U.S. Virgin Islands (Wil- on extant species. Until more thorough in- liams 1984). vestigation, incorporating a broader range of characters, is conducted, the classifica- it Platylambrus pourtalesii tion of Williams (1984) will be followed. (Stimpson, 1871) Williams & Wahle (1992) summarized and illustrated the differences between juvenile Parthenope (Platylambrus) pourtalesii Cancer borealis and C. irroratus. (Stimpson).—Williams, 1984:345.

Known range.—Western Atlantic: off Cancer (.Metacarcinus) borealis Stimpson, Martha's Vineyard, Massachusetts; New 1859 Jersey to southern Florida; Gulf of Mexico through West Indies to Grenada (Williams Known range.—Western Atlantic: Nova 1984, Abele & Kim 1986). Scotia to south of Dry Tortugas, Florida (Williams 1984). Solenolambrus tenellus Stimpson, 1871 Cancer {Cancer) irroratus Say, 1817 Known range.—Western Atlantic: off Cape Lookout, North Carolina; east central Known range.—Western Atlantic: Lab- Florida southward, including the Florida rador to off Miami, Florida (Williams Keys, into eastern Gulf of Mexico to near 1984). Cape St. George, Florida; Bahamas; Bar- bados (Williams 1984, Abele & Kim 1986). Superfamily Portunoidea Rafinesque, 1815 Family Geryonidae Colosi, 1923 Solenolambrus typicus Stimpson, 1871 • * Chaceon quinquedens (Smith, 1879) Known range.—Western Atlantic: southeast of Cape Lookout, North Carolina; Geryon quinquedens Smith.—Williams, western Gulf of Mexico off Corpus Christi, 1984:485. Texas, and north of Yucatan; Nicaragua Remarks.—Williams (1984) considered Shelf; southern Florida through West Indies this species to be extralimital. Manning & to Suriname and Rio de Janeiro, Brazil Holthuis (1989) described the new genus (Williams 1984, Melo 1998). Chaceon for 21 species including C. quinquedens. This species occurs at 40—2155 m Superfamily Cancroidea Latreille, 1802 depth, but is usually found at the shelf edge Family Cancridae Latreille, 1802 or on the continental slope (Wigley et al. Remarks.—Based on fossil evidence (pri- 1975, Williams 1984). marily characters of the carapace and che- Known range.—Western Atlantic: Nova lipeds) Nations (1975) recognized four sub- Scotia southward to Gulf of Mexico; Cuba; genera (Glebocarcinus Nations, 1975, Rom- Brazil and Argentina (Wigley et al. 1975, aleon Gistl, 1848, Metacarcinus A. Milne- Williams & Wigley 1977, Williams 1984). VOLUME 116, NUMBER 1 135

Family Portunidae Rafinesque, 1815 Ovalipes stephensoni Williams, 1976 Subfamily Carcininae Alcock, 1899 Known range.—Western Atlantic: south- Carcinus maenas (Linnaeus, 1758) ern New Jersey to Biscayne Bay, Florida Remarks.—Behrens Yamada & Hauck (Williams 1984, Stehlik et al. 1991). (2001) provided extensive information on distinguishing characteristics of this species Subfamily Portuninae Rafinesque, 1815 and evaluated the usefulness of these char- Arenaeus cribrarius (Lamarck, 1818) acters in making field identifications. Remarks.—Juveniles of this species were Known range.—Western Atlantic (intro- collected at 6-10 m on sand and mud-sand duced): Northumberland Strait and Cape sediments (Scelzo 2001). Breton to Virginia; Eastern Atlantic: Ice- Known range.—Western Atlantic: Vine- land; Norway, including southwestern and yard Sound, Massachusetts, to Mar del Pla- rarely southern Baltic Sea, through North ta, Argentina (Williams 1984, Scelzo 2001). Sea and British Isles to Mauritania; northwest Africa; also introduced to South Af- Callinectes bocourti A. Milne-Edwards, rica; Madagascar; Red Sea; Myanmar; In- 1879 dia; Ceylon; Japan; Australia; Tasmania (Williams 1984, Squires 1990, Behrens Ya- Known range.—Western Atlantic: occa- mada & Hauck 2001); multiple eastern Pa- sionally North Carolina, Florida, and Mis- cific localities, including sites in California, sissippi; otherwise Jamaica; Belize to Santa Oregon, Washington, and Vancouver Is- Catarina, Brazil, including Antilles and land, British Colombia (Cohen et al. 1995, northern coast of South America (Williams Grosholz & Ruiz 1996, Behrens Yamada et 1984, Melo 1998). al. 2000). Callinectes danae Smith, 1869 Subfamily Polybiinae Ortmann, 1893 Known range.—Western Atlantic: Ber- • Bathynectes longispina Stimpson, 1871 muda; New Hanover County, North Caro- Remarks.—Williams (1984) considered lina (near Cape Fear); southern Florida; this species to be extralimital. This species eastern side of Yucatan Peninsula to Rio has been captured and observed recently at Grande do Sul, Brazil, including Antilles depths somewhat shallower (124—152 m; V. and northern coast of South America (Wil- Guida, pers. comm.) than previously re- liams 1984, Melo 1998). ported (100-1455 m, commonly >200 m; Williams & Wigley 1977). Manning & Hol- Callinectes exasperatus thuis (1981) reported that western Atlantic (Gerstaecker, 1856) specimens identified as B. superbus (Costa, Known range.—Western Atlantic: Ber- 1853) are actually B. longispina. muda; Duval County, east of Jacksonville, Known range.—Western Atlantic: Mar- Florida, to Santa Catarina, Brazil, including tha's Vineyard, Massachusetts, to off Mis- Antilles and Venezuela; extreme southern sissippi River delta southward to Goajara Texas; Veracruz, Mexico (Williams 1984, Peninsula, Colombia; Bermuda (Williams Melo 1998). 1984). Callinectes larvatus Ordway, 1863 Ovalipes ocellatus (Herbst, 1799) Known range.—Western Atlantic: Ber- Known range.—Western Atlantic: North- muda; Beaufort, North Carolina, through umberland Strait, Prince Edward Island, to Caribbean Sea to Sao Paulo, Brazil, includ- Georgia (Williams 1984). ing Antilles (Williams 1984, Melo 1998). 136 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Callinectes ornatus Ordway, 1863 Portunus depressifrons (Stimpson, 1859) Known range.—Western Atlantic: Vir- Known range.—Western Atlantic: Ber- ginia through southern Florida; Bermuda; muda; Fort Macon, North Carolina, through northwestern Yucatan to Rio Grande do northwest Florida to Gulf of Campeche and Sul, Brazil, including Antilles (Williams Caribbean Sea (Williams 1984). 1984, Melo 1998). Portunus floridanus Rathbun, 1930 Callinectes sapidus Rathbun, 1896 Known range.—Western Atlantic: east of Known range.—Western Atlantic: Cape Cape Lookout, North Carolina, to Nicara- Cod, Massachusetts, to northern Argentina, gua; through West Indies and northern including Bermuda, Antilles, Central Amer- South America to Suriname (Williams ica, and Venezuela (occasionally north of 1984). Cape Cod to Maine and Nova Scotia, in favorably warm periods); Eastern Atlantic: Portunus gibbesii (Stimpson, 1859) 0resund, Denmark; the Netherlands and ad- Known range.—Western Atlantic: south- jacent North Sea; northwest and southwest ern Massachusetts through Gulf of Mexico France; Mediterranean Sea, including along coast to French Guiana (Williams northern Adriatic, Aegean, and western 1984); Bahia, Brazil (Melo 1998). Black seas; Eastern Pacific: Japan (Wil- liams 1984, Squires 1990, Melo 1998). Portunus ordwayi (Stimpson, 1860) Callinectes similis Williams, 1966 Known range.—Western Atlantic: Vine- yard Sound, Massachusetts; Bermuda; Known range.—Western Atlantic: off North Carolina through Gulf of Mexico, Delaware Bay to Key West, Florida; north- West Indies and Caribbean Sea to Rio western Florida around Gulf of Mexico to Grande do Sul, Brazil; Fernando de Noro- off Campeche, Yucatan; Isla de Providen- nha, Brazil (Williams 1984, Melo 1998). cia, Colombia (Williams 1984). Portunus sayi (Gibbes, 1850) Cronius ruber (Lamarck, 1818) Known range.—Western Atlantic: off the Known range.—Western Atlantic: Little Grand Banks through Gulf of Mexico to the Egg Inlet, New Jersey; Rehoboth Bay, Del- Guianas; Bermuda; Eastern Atlantic: Ca- aware; Virginia; South Carolina to Rio nary Islands; Morocco (Williams 1984, Grande do Sul, Brazil, including Central Squires 1990). America, Antilles, and northern South America; Eastern Atlantic: West Africa Portunus spinicarpus (Stimpson, 1871) from Mauritania to Angola; Cape Verde, Principe, Sao Tome and Annobon islands; Known range.—Western Atlantic: off Eastern Pacific: Baja California to Peru; Oregon Inlet, North Carolina, to Santa Ca- Clipperton Island; Galapagos Islands (Wil- tarina, Brazil, including Antilles and north- liams 1984, Melo 1998). ern South America (Williams 1984, Melo 1998). Portunus anceps (de Saussure, 1858) Portunus spinimanus Latreille, 1819 Known range.—Western Atlantic: Ber- muda; Cape Hatteras, North Carolina, to Known range.—Western Atlantic: New Rio de Janeiro, Brazil, including Antilles Jersey to Rio Grande do Sul, Brazil, in- (Williams 1984, Melo 1998). cluding Gulf of Mexico, West Indies and VOLUME 116, NUMBER 1 137 northern South America; Bermuda (Wil- Goneplax sigsbei liams 1984, Melo 1998). (A. Milne-Edwards, 1880) Known range.—Western Atlantic: east of Superfamily Xanthoidea MacLeay, 1838 Cape Fear, North Carolina; Grenada (Wil- Remarks.—Guinot (1978) concluded that liams 1984). Xanthidae actually represented eight different families (Carpiliidae Ortmann. 1893, Speocarcinus carolinensis Stimpson, 1859 Menippidae Ortmann. 1893, Platyxanthidae Known range.—Western Atlantic: south Guinot, 1977, Xanthidae McLeay, 1838, of Cape Hatteras, North Carolina, to Rio Panopeidae Ortmann, 1893, Pilumnidae Sa- Grande do Sul, Brazil, including West In- mouelle. 1819, Trapeziidae Miers, 1886, dies (Williams 1984, Melo 1998). and Geryonidae Colosi, 1924) which should be recognized under the superfamily * Family Menippidae Ortmann, 1893 Xanthoidea. Williams (1984) elected not to adopt this classification. Instead, he placed Remarks.—Williams (1984) listed these the majority of xanthid taxa (exclusive of taxa under the family Xanthidae. the Goneplacidae) under the single family Xanthidae. In the intervening years, Guin- Eriphia gonagra (Fabricius, 1781) ot's classification, with minor modification, Known range.—Western Atlantic: Ber- has gained acceptance by the majority of muda; North Carolina to Patagonia, includ- crustacean researchers (Schubart, Neigel, & ing Central America, Antilles, and northern Felder 2000, Martin & Davis 2001). South America (Williams 1984, Melo 1998). Family Goneplacidae MacLeay, 1838 Menippe mercenaria (Say, 1818) Remarks.—The Goneplacidae has long been recognized as containing heterogenous Remarks.—Williams & Felder (1986) groups of genera (Hendrickx 1998, and ref- recognized two morphologically distinct erences therein). Because revisionary stud- populations of stone crab in the Gulf of ies of the Goneplacidae are still in progress, Mexico and determined that these popula- formal subdivision into subfamilies is not tions represented distinct species. Range of attempted here. Furthermore, after further Menippe mercenaria was restricted; M. ad- revision of this family, it is likely that new ina Williams & Felder, which ranges from families will be added and subfamilies el- northwestern Florida around the Gulf of evated to full family status (Guinot 1978, Mexico to Tamaulipas State, Mexico, was Williams 1984, Sternberg, pers. comm.). described (Williams & Felder 1986). Known range.—Western Atlantic: Cape Euryplax nitida Stimpson, 1859 Lookout, North Carolina, through peninsular Florida; Bahamas and Greater Antilles Known range.—Western Atlantic: Ber- to Yucatan Peninsula, Mexico; and Belize muda; off Beaufort, North Carolina, to (Williams & Felder 1986). Heald Bank, Texas; Antilles to Santa Ca- tarina, Brazil (Williams 1984, Melo 1998). * Family Panopeidae Ortmann, 1893

Frevillea hirsuta (Borradaile, 1916) Remarks.—Williams (1984) listed these taxa under the family Xanthidae. Genera in- Known range.—Western Atlantic: North cluded in this family were based on the Carolina to Rio Grande do Sul, Brazil (Wil- conclusions of Guinot (1978) and subse- liams 1984. Melo 1998). quent acceptance of generic placements by 138 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Martin & Abele (1986). Schubart, Neigel, Known range.—Western Atlantic: Cape & Felder (2000) presented a molecular phy- Hatteras, North Carolina, to Gulf of Mexico logeny of western Atlantic Panopeidae. and Yucatan Channel (Williams 1984).

it Dyspanopeus sayi (Smith, 1869) Hexapanopeus angustifrons Neopanope sayi (Smith).—Williams, 1984: (Benedict & Rathbun, 1891) 409. Known range.—Western Atlantic: Vine- Remarks.—Martin & Abele (1986) de- yard Sound, Massachusetts, to Port Aran- scribed the new genus Dyspanopeus for D. sas, Texas; West Indies; Pernambuco to sayi (occurring in the western Atlantic) and Santa Catarina, Brazil (Williams 1984, D. texanus (occurring in the Gulf of Mex- Melo 1998). ico). Known range.—Western Atlantic: south- Hexapanopeus paulensis Rathbun, 1930 ern Gulf of St. Lawrence to Florida Keys; Known range.—Western Atlantic: South Eastern Atlantic (introduced): Bristol Chan- Carolina to Uruguay, including Gulf of nel, United Kingdom (Williams 1984, Mexico (Williams 1984). Squires 1990).

Eurypanopeus abbreviatus Panopeus herbstii H. Milne Edwards, (Stimpson, 1860) 1834 Known range.—Western Atlantic: South Remarks.—Williams (1983) examined Carolina to Rio Grande do Sul, Brazil, in- the Panopeus herbstii species complex and cluding Gulf of Mexico, Antilles, and determined that 'forms' previously recog- northern South America (Williams 1984, nized by Rathbun (1930) were sufficiently Melo 1998). different and represented distinct species. As a result, P. herbstii was better defined, Eurypanopeus depressus (Smith, 1869) and its range determined to be the shallow intertidal and subtidal waters of the eastern Known range.—Western Atlantic: Mas- United States. sachusetts Bay through Florida to southern Known range.—Western Atlantic: Bos- Texas; Dutch West Indies; Uruguay; Ber- ton Harbor, Massachusetts, to Indian River muda (Williams 1984). County, Florida (Williams 1983).

Eurytium limosum (Say, 1818) • Panopeus obesus Smith, 1869 Known range.—Western Atlantic: Ber- Remarks.—This species was recognized muda; South Carolina; Louisiana to Santa previously as a form of Panopeus herbstii Catarina, Brazil, including West Indies and (Rathbun 1930). Williams (1983) rediag- Caribbean Sea (Williams 1984, Melo nosed this form and elevated it to full spe- 1998). cies rank; diagnosis, measurements, and color description of this species were also Glyptoplax smithii A. Milne-Edwards, provided. Panopeus obesus is found in 1880 marsh edge, and shallow intertidal and sub- Remarks.—This species previously was tidal waters of the Carolinian province. placed in the family Goneplacidae (Wil- Known range.—Western Atlantic: Beau- liams 1984). Guinot (1978) transferred this fort, North Carolina, to northeastern Flori- species to the Panopeidae; Martin & Abele da; Sarasota County, Florida, to Louisiana; (1986) adopted this arrangement. Texas; northern Mexico (Williams 1983). VOLUME 116, NUMBER 1 139

Panopeus occidentalis de Saussure, 1857 Pilumnus floridanus Stimpson, 1871

Known range.—Western Atlantic: Ber- Known range.—Western Atlantic: off muda; North Carolina to Santa Catarina, Cape Lookout, North Carolina, to Bahia, Brazil, including Central America, Antilles Brazil, including Gulf of Mexico, Yucatan and northern South America (Williams Channel, Central America, Venezuela, and 1984, Melo 1998). West Indies (Williams 1984, Melo 1998).

Panoplax depressa Stimpson, 1871 Pilumnus lacteus Stimpson, 1871

Remarks.—This species was placed pre- Known range.—Western Atlantic: near viously in the family Goneplacidae (Wil- Beaufort, North Carolina, to Florida; Cuba liams 1984). Guinot (1978) transferred this (Williams 1984). species to the Panopeidae; Martin & Abele (1986) adopted this arrangement. Pilumnus pannosus Rathbun, 1896 Known range.—Western Atlantic: southeast of Cape Lookout, North Carolina; off Known range.—Western Atlantic: Bogue Jacksonville and Cape San Bias, Florida; Sound, off Beaufort, North Carolina, to Port through Antilles; Amapa to Pernambuco, Aransas, Texas; West Indies to Virgin Is- Brazil (Williams 1984, Melo 1998). lands (Williams 1984).

Rhithropanopeus harrisii (Gould, 1841) Pilumnus sayi Rathbun, 1897

Known range.—Western Atlantic: south- Known range.—Western Atlantic: North western Gulf of St. Lawrence to Veracruz, Carolina to Cura£ao, including Gulf of Mexico; Eastern Atlantic (introduced): Mexico and West Indies (Williams 1984). parts of Europe; Eastern Pacific (introduced): west coast of United States (Wil- * Family Pseudorhombilidae Alcock, liams 1984, Squires 1990). 1900

* Family Pilumnidae Samouelle, 1819 Remarks.—This taxon was previously considered a subfamily (Pseudorhombili- Remarks.—Williams (1984) listed these nae) within the Goneplacidae (Williams taxa under the family Xanthidae. 1984). Hendrickx (1998) proposed the family designation to accommodate six genera. Lobopilumnus agassizii (Stimpson, 1871) Nanoplax xanthiformis Known range.—Western Atlantic: Ber- (A. Milne-Edwards, 1881) muda; North Carolina; eastern Gulf of Mexico; Yucatan; Cuba; Venezuela; Trini- Known range.—Western Atlantic: Cape dad (Williams 1984). Hatteras, North Carolina, to Rio de Janeiro, Brazil, including Gulf of Mexico, Yucatan, Piliimnus dasypodus Kingsley, 1879 West Indies, and northern South America (Williams 1984, Melo 1998). Known range.—Western Atlantic: off Cape Hatteras, North Carolina, to Santa Ca- * Family Trapeziidae Miers, 1886 tarina, Brazil, including Gulf of Mexico, Caribbean Sea, and West Indies (Williams Remarks.—Williams (1984) listed this 1984, Melo 1998). taxon under the family Xanthidae. 140 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Domecia acanthophora acanthophora Micropanope sculptipes Stimpson, 1871 (Desbonne & Schramm, 1867) Known range.—Western Atlantic: south- Known range.—Western Atlantic: Ber- east of Cape Lookout, North Carolina, to muda; Cape Lookout Shoals, North Caro- Port Aransas, Texas; West Indies to Bar- lina; northwestern Gulf of Mexico to Ala- bados; Amapa to Rio de Janeiro, Brazil goas, Brazil, including West Indies and Ca- (Williams 1984, Melo 1998). ribbean Sea (Williams 1984, Melo 1998). Micropanope urinator Family Xanthidae MacLeay, 1838 (A. Milne-Edwards, 1881) Allactaea lithostrota Williams, 1974 Known range.—Western Atlantic: off Known range.—Western Atlantic: Ber- Cape Hatteras and Cape Lookout, North muda; off Cape Lookout, North Carolina; Carolina; Florida Keys to St. Croix (Wil- Florida Straits; off Cape Catoche, Yucatan; liams 1984); Antilles; Para to Maranhao, off Venezuela and Suriname (Williams Brazil (Melo 1998). 1984); Antilles; off Rio de Janeiro to Rio Grande do Sul, Brazil (Melo 1998). Paractaea rufopunctata nodosa Carpoporus papulosus Stimpson, 1871 (Stimpson, 1860)

Known range.—Western Atlantic: be- Known range.—Western Atlantic: south- tween Cape Hatteras and Cape Lookout, east of Cape Lookout, North Carolina; off North Carolina; Gulf of Mexico, off Mobile Mississippi River delta, through West In- Bay southeastward; Cape Catoche, Yucatan dies to Uruguay; Central Atlantic: Ascen- (Williams 1984). sion Island (Williams 1984, Melo 1998).

Glyptoxanthus erosus (Stimpson, 1859) Pseudomedaeus agassizii (A. Milne-Edwards, 1880) Known range.—Western Atlantic: Cape Lookout, North Carolina, southward around Known range.—Western Atlantic: Cape Florida into Gulf of Mexico to off Grand Hatteras, North Carolina, to southern Texas Isle, Louisiana; Yucatan; West Indies to (Williams 1984). Guadeloupe (Williams 1984).

Pseudomedaeus distinctus (Rathbun, 1898) Melybia thalamita Stimpson, 1871 Known range.—Western Atlantic: off Known range.—Western Atlantic: off Cape Hatteras, North Carolina, through Cape Lookout, North Carolina, southwest Straits of Florida to northwest of Dry Tor- of Mississippi River delta to Sao Paulo, tugas; Puerto Rico; Barbados (Williams Brazil, including West Indies and northern 1984). South America (Williams 1984, Melo 1998). Tetraxanthus rathbunae Chace, 1939 Micropanope nuttingi (Rathbun, 1898) Known range.—Western Atlantic: off Known range.—Western Atlantic: Cape Cape Lookout, North Carolina, to Rio Hatteras, North Carolina, to Sao Paulo, Bra- Grande do Sul, Brazil, including Gulf of zil, including Gulf of Mexico and West In- Mexico and Antilles (Williams 1984, Melo dies (Williams 1984, Melo 1998). 1998). VOLUME 116, NUMBER 1 141

Superfamily Pinnotheroidea De Haan, new genus Gemmotheres for P. chamae', il- 1833 lustrations were also provided. Family Pinnotheridae De Haan, 1833 Known range.—Western Atlantic: North Carolina coast (Williams 1984, Campos Remarks.—Marques & Pohle (1995) 1996). conducted a phylogenetic analysis of this family using larval characters and demon- • Parapinnixa beaufortensis Rathbun, strated that several taxa are paraphyletic 1918 taxa and that further analysis will be needed to resolve relationships within this family. Remarks.—Williams (1984) considered this species to be extralimital. Subfamily Pinnotherinae De Haan, 1833 Known range.—Western Atlantic: off Beaufort, North Carolina (Williams 1984). Remarks.—Griffith (1987) presented a hypothesis of phylogenetic relationships Parapinnixa bouvieri Rathbun, 1918 within the genus Dissodactylus based on Known range.—Western Atlantic: off adult morphology. Marques & Pohle (1995) Charleston, South Carolina; south of Dry conducted a phylogenetic analysis of mem- Tortugas, Florida; off Cape Catoche, Yu- bers of this genus using larval characters catan; Puerto Rico; Amapa, Brazil (Wil- and produced results that only partially cor- liams 1984, Melo 1998). roborated the relationships proposed by Griffith (1987). In a separate analysis, both Parapinnixa hendersoni Rathbun, 1918 data sets (adult morphology and larval morphology) were combined (Marques & Pohle Known range.—Western Atlantic: south- 1995). Results of this analysis provided a east of Cape Lookout, North Carolina; off more robust hypothesis of relationships Tampa Bay, Florida, through West Indies to than when either character set was analyzed Curasao; Venezuela; Maranhao to Espirito independently. Santo, Brazil (Williams 1984, Melo 1998).

Dissodactylus crinitichelis Moreira, 1901 Pinnaxodes floridensis Wells and Wells, 1961 Known range.—Western Atlantic: southeast of Cape Lookout, North Carolina; off Known range.—Western Atlantic: off northwest Florida; Caribbean Sea and South North Carolina to Georgia; northwest Flor- America to Rio de la Plata, Argentina (Wil- ida (Williams 1984). liams 1984). ^k Tumidotheres maculatus (Say, 1818) Dissodactylus mellitae (Rathbun, 1900) Pinnotheres maculatus Say.—Williams, 1984:441. Known range.—Western Atlantic: western Vineyard Sound, Massachusetts, to Remarks.—Campos (1989) described the Charleston, South Carolina; Hutchinson Is- new genus Tumidotheres and discussed ge- land, east Florida; western Florida; off Gal- neric relationships and life history traits. veston, Texas (Williams 1984). Known range.—Western Atlantic: off Martha's Vineyard, Massachusetts, to Golfo ^k Gemmotheres chamae (Roberts, 1975) San Matfas, Argentina (Williams 1984).

Pinnotheres chamae Roberts.—Williams, ik Zaops ostreum (Say, 1817) 1984:440. Pinnotheres ostreum Say.—Williams, 1984: Remarks.—Campos (1996) described the 444. 142 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Remarks.—Manning (1993) concluded Carolina; Georgia; off Mississippi River that Zaops Rathbun, 1900 should be re- delta; Seven and One-Half Fathom Reef, moved from the synonymy of Pinnotheres off Texas (Williams 1984). and recognized as a distinct genus. Addi- tionally, other pinnotherid genera formerly Pinnixa retinens Rathbun, 1918 placed in the synonymy of Pinnotheres were diagnosed, figured and differentiated Known range.—Western Atlantic: Dela- from Pinnotheres sensu Manning (1993). ware Bay; Chesapeake Bay; Little River In- let, South Carolina; Alligator Harbor, Flor- Known range.—Western Atlantic: Salem, ida; Aransas area of Texas coast (Williams Massachusetts, to Santa Catarina, Brazil 1984). (Williams 1984).

Subfamily Pinnothereliinae Alcock, 1900 Pinnixa say ana Stimpson, 1860 it Austinixa cristata (Rathbun, 1900) Known range.—Western Atlantic: Vine- Pinnixa cristata Rathbun.—Williams, yard Sound, Massachusetts, to Beaufort, 1984:453. North Carolina; Hutchinson Island, east central Florida; Sarasota Bay, Florida to Remarks.—Heard & Manning (1997) Grand Isle, Louisiana; Amapa to Rio recognized and described the new genus Grande do Sul, Brazil (Williams 1984, Austinixa for Pinnixa cristata and six other Melo 1998). species formerly assigned to Pinnixa. Known range.—Western Atlantic: Beau- Superfamily Ocypodoidea Rafinesque, fort, North Carolina, to Miami, Florida; 1815 northern and southwestern Gulf of Mexico Family Ocypodidae Rafinesque, 1815 (Manning & Felder 1989); Central Ameri- Subfamily Ocypodinae Rafinesque, 1815 ca; Brazil (Melo 1998). Remarks.—Rosenberg (2001) conducted Pinnixa chaetopterana Stimpson, 1860 a phylogenetic analysis of the genus Uca using 236 discrete morphological charac- Known range.—Western Atlantic: Well- ters. Although many scientists ignore sub- fleet, Massachusetts, to Rio Grande do Sul, generic designations, Rosenberg (2001) Brazil (Williams 1984). considered subgenera within Uca to be valid. The three species of Uca occurring in Pinnixa cylindrica (Say, 1818) the region are considered to be members of Known range.—Western Atlantic: North the subgenus Minuca (Rosenberg 2001). Falmouth, Massachusetts, to Pensacola, Florida, including Dry Tortugas (Williams Ocypode quadrata (Fabricius, 1787) 1984). Known range.—Western Atlantic: Block Island, Rhode Island, to Rio Grande do Sul, Pinnixa floridana Rathbun, 1918 Brazil; Fernando de Noronha Archipelago, Known range.—Western Atlantic: south- Brazil; Bermuda (Williams 1984, Melo east of Cape Lookout, North Carolina; 1998). Hutchinson Island, east Florida; west coast of Florida (Williams 1984). Uca minax (LeConte, 1855) Remarks.—Felder & Staton (1994) ana- Pinnixa lunzi Glassell, 1937 lyzed electrophoretic allozyme assays and Known range.—Western Atlantic: off the observed slight differentiation between eastern shore of Virginia; North and South Gulf of Mexico and Atlantic populations. VOLUME 116, NUMBER 1 143

Known range.—Western Atlantic: Buz- Palicus sica (A. Milne-Edwards, 1880) zards Bay, Cape Cod, Massachusetts, to Known range.—Western Atlantic: off Daytona Beach, Florida; Yankeetown, Charleston, South Carolina, to Cape Can- northwest Florida, to Matagorda Bay, Texas averal, Florida; west coast of Florida to Rio (Williams 1984, Barnwell & Thurman Grande do Sul, Brazil, including West In- 1984). dies (Williams 1984, Melo 1998). Uca pugilator (Bosc, 1802) Superfamily Grapsoidea MacLeay, 1838 Known range.—Western Atlantic: Cape Family Grapsidae MacLeay, 1838 Cod, Massachusetts, southward around the tip of peninsular Florida and westward to Remarks.—Previously, family Grapsidae Pensacola Beach, Florida; possible rare oc- was considered to be comprised of four currences in Bahamas and western Gulf of subfamilies. Based on a cladistic study by Mexico (Barnwell & Thurman 1984). Sternberg & Cumberlidge (1998) the Grap- sidae were redefined and restricted to in- Uca pugnax (Smith, 1870) clude all genera previously placed in the subfamily Grapsinae. Molecular data also Known range.—Western Atlantic: Prov- supported elevation of grapsid subfamilies incetown, Massachusetts, to Daytona to full family status (Schubart, Cuesta, Die- Beach, Florida (Williams 1984, Barnwell & sel, & Felder 2000). Thurman 1984). Pachygrapsus transversus (Gibbes, 1850) Family Palicidae Bouvier, 1898 Palicus alternatus Rathbun, 1897 Known range.—Western Atlantic: Ber- muda; Cape Lookout, North Carolina, to Known range.—Western Atlantic: Cape Montevideo, Uruguay; Eastern Atlantic: Hatteras to southeast-of Cape Fear, North Mediterranean Sea to northern Angola; Carolina; Gulf of Mexico along west coast Eastern Pacific: California to Peru; Gala- of Florida from Cape San Bias to Key West pagos Islands (Williams 1984). (Williams 1984).

Palicus faxoni Rathbun, 1897 Planes minutus (Linnaeus, 1758) Known range.—Western Atlantic: off Known range.—Western Atlantic: off the Cape Hatteras, North Carolina, to near Grand Banks of Newfoundland south to Cape Canaveral, Florida; off Yucatan, Mex- 11°N, exclusive of Gulf of Mexico; Eastern ico, near Quita Sueno Banks; southwest of Atlantic: southern North Sea south to 11°N, St. Christopher; Rio Grande do Norte to including Mediterranean Sea (Williams Rio de Janeiro, Brazil (Williams 1984, 1984, Squires 1990). Melo 1998). it Family Plagusiidae Dana, 1851 • Palicus gracilis (Smith, 1883) Remarks.—This taxon was previously Remarks.—Williams (1984) considered considered a subfamily of the Grapsidae this species to be extralimital. This species (e.g., Williams 1984, Guinot & Bouchard occurs at 183-512 m (Williams & Wigley 1998). Based on results of a cladistic study, 1977). Sternberg & Cumberlidge (1998) concluded Known range.—Western Atlantic: Mar- that the two genera (Percnon and Plagusia) tha's Vineyard, Massachusetts, to Cura9ao, placed in the subfamily Plagusiinae formed including Gulf of Mexico (Williams & a monophyletic taxon; subfamily Plagusi- Wigley 1977). inae was redefined and elevated to family 144 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON status (Plagusiidae). Based on results of a Plagusia depressa (Fabricius, 1775) molecular phylogeny, however, Schubart, Known range.—Western Atlantic: Ber- Cuesta, Diesel, & Felder (2000) questioned muda; Beaufort, North Carolina, to Bahia, taxonomic placement of Percnon in this Brazil, including Gulf of Mexico and West family. Sternberg & Cumberlidge (1998) Indies; Fernando de Noronha Archipelago, had placed Euchirograpsus in the Varuni- Rocas Atoll, Saint Paul Rocks, and Trin- nae. However, based on morphological and dade Island, Brazil; Eastern Atlantic: molecular evidence (Schubart, Cuesta, Die- Azores; Madeira; Morocco to northern An- sel, & Felder 2000), Euchirograpsus be- gola; Central Atlantic: St. Helena Island longs in the Plagusiidae. (Williams 1984, Melo 1998).

Euchirograpsus americanus * Family Sesarmidae Dana, 1851 A. Milne-Edwards, 1880 Remarks.—This taxon was previously Remarks.—This species was previously considered a subfamily of the Grapsidae considered a member of the Varuninae (e.g., Williams 1984, Guinot & Bouchard (Williams 1984). Based on molecular evi- 1998). Based on results of a cladistic study, dence and larval morphology, Schubart, Sternberg & Cumberlidge (1998) concluded Cuesta, Diesel, & Felder (2000) proposed that taxa previously included in the subfam- inclusion of this species in the family Pla- ily Sesarminae formed a polyphyletic gusiidae. group. However, one clade comprising the Known range.—Western Atlantic: majority of Sesarma-like genera, including Oceanographer Canyon, edge of Georges Sesarma and Armases, was redefined as the Bank; off Oregon Inlet, North Carolina; family Sesarmidae (Sternberg & Cumber- Florida to Venezuela, including West In- lidge 1998). dies; Rio Grande do Sul, Brazil (Williams 1984, 1988; Melo 1998). it Armases cinereum (Bosc, 1802) Sesarma (Chiromantes) cinereum (Bosc).— Percnon gibbesi Williams, 1984:465. (H. Milne Edwards, 1853) Remarks.—Abele (1992) described the Remarks.—Based on molecular evi- new genus Armases for American species dence, Schubart, Cuesta, Diesel, & Felder of Sesarma assigned to the subgenus Chi- (2000) suggested that this species does not romantes; generic diagnosis, species de- belong in the family Plagusiidae. Those au- scription, and illustrations are provided. thors reported that the taxonomic position The phylogeny of Armases proposed by of Percnon gibbesi was uncertain and re- Niem (1996) supported establishment of quired further investigation. Until more de- Armases Abele, 1992. finitive results are available, placement of Known range.—Western Atlantic: Ma- this species will remain in the Plagusiidae. gothy River, Chesapeake Bay, Maryland, to Known range.—Western Atlantic: Ber- Palm Beach, east Florida; Collier County, muda; Fort Macon, North Carolina; southwest Florida, to Veracruz, Mexico (Wil- ern Florida and Bahamas to Fernando de liams 1984). Noronha Archipelago, Brazil, including Antilles; Eastern Atlantic: Azores to An- it Sesarma reticulatum (Say, 1817) gola; Mediterranean Sea (Pipitone et al. Sesarma (Sesarma) reticulatum (Say).— 2001); Eastern Pacific: Cape San Lucas, Williams, 1984:466. lower California, to Chile; Galapagos Is- lands (Williams 1984, Melo 1998). Remarks.—Subgenera were not recog- VOLUME 116, NUMBER 1 145 nized within the genus Sesarma by Abele O'Connor 2001) observed higher abun- (1992). Based on electrophoretic results, dances in the middle and lower intertidal Felder & Staton (1994) concluded that al- zone of sampling locations in eastern Long lozyme divergence between Gulf of Mexico Island Sound and southeastern New Eng- and Atlantic populations was comparable to land. Crab abundance increased with in- levels previously reported for speciated creased rock cover (Ledesma & O'Connor populations, suggesting that Sesarma reti- 2001). culatum represented a species complex. Known range.—Western Atlantic: Ap- Known range.—Western Atlantic: Woods pledore Island, Isles of Shoals, Maine (J. Hole, Massachusetts, to Volusia County, Morin, pers. comm.) and New Hampshire east Florida; Sarasota, west Florida, to Bar- (McDermott 2000) to Oregon Inlet, North ra del Tordo, Tamaulipas, Mexico (Williams Carolina (McDermott 1998); Eastern Atlan- 1984, Felder & Staton 1994). tic: Le Havre, France; "Oosterschelde", Netherlands (Breton et al. 2002); Western * Family Varunidae H. Milne Edwards, Pacific: Sakhalin, Korea; north China to 1853 Hong Kong; all coasts of Japan from Hok- kaido to Okinawa (Williams & McDermott Remarks.—This taxon was previously 1990). considered a subfamily of the Grapsidae (e.g., Williams 1984, Guinot & Bouchard 1998). Cladistic analysis revealed that the Discussion Varuninae is an artificial group in need of The decapod crustacean assemblage oc- re-examination (Sternberg & Cumberlidge curring in shallow waters (^190 m) of the 1998). It is possible that this group, pres- temperate eastern United States from Maine ently recognized at the family level (Schu- to Cape Canaveral, Florida, totals 391 spe- bart, Cuesta, Diesel, & Felder 2000, Martin cies. This assemblage includes 122 shrimp & Davis 2001) may prove to represent sev- species (28 penaeids, 2 stenopodids, and 92 eral different families (Sternberg & Cum- carideans), 10 thalassinideans, 8 lobsters, berlidge 1998). Molecular data also support 61 anomurans, and 190 brachyurans. By this conclusion (Schubart, Cuesta, Diesel, comparison, the previous comprehensive & Felder 2000). review of this assemblage by Williams (1984) recognized 103 species of shrimps • Hemigrapsus sanguineus (21 penaeids, 2 stenopodids, and 80 cari- (De Haan, 1853) deans), 8 thalassinideans (including calli- Remarks.—The Asian shore crab was anassids, upogebiids and axiids), 6 lobsters, most likely introduced in the western At- 51 anomurans, and 174 brachyurans (total lantic via ballast water discharged some- decapods = 342). time in the early 1980's (McDermott 1998). Since publication of Williams (1984), 51 The crab was first discovered in 1988 in species are new to this checklist. Thirteen southern New Jersey (Williams & Mc- species (Table 1) previously considered ex- Dermott 1990). Hemigrapsus sanguineus tralimital by Williams (1984:484) because was the most abundant brachyuran at the their centers of distribution or abundance intertidal monitoring site in southern New occur beyond the boundaries of the region Jersey, some areas of Long Island Sound are now incorporated into the updated (McDermott 1998) and Narragansett Bay, checklist because their geographic and RI (pers. obs.). McDermott (1998) reported bathymetric ranges are within the limits set this crab occurring in the upper to middle for the region under consideration. An ad- intertidal zone of New Jersey, whereas oth- ditional 16 species (Table 1), most likely ers (Lohrer & Whitlatch 1997, Ledesma & excluded (i.e., not even considered as ex- 146 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 1.—Western Atlantic decapod Crustacea occurring in regions that were considered 'extralimital' or excluded from consideration by Williams (1984) because their geographic or bathymetric range centered beyond the boundaries of regions of consideration.

'Extralimital' Excluded

Pleoticus robustus Aristaeomorpha foliacea Bythocaris nana Hadropenaeus ajfinis Caridion gordoni Hadropenaeus modestus Plesionika martia Penaeopsis serrata Plesionika tenuipes Lucifer typus Plesionika willisi Heterocarpus ensifer Catapagurus sharreri Plesionika edwardsii Heniipagurus gracilis Nephropsis aculeata Mithrax cor nut us Tomopagurus cokeri Bathynectes longispina Parapagurus pilosimanus Chaceon quinquedens Sympagurus pictus Parapinnixa beaufortensis Munida forceps Palicus gracilis Lysirude nitidus Cyclozodion angustum Pyromaia arachna Stenocionops furcata furcata tralimital) from consideration by Williams be members of this assemblage (Williams because their bathymetric distributions 1984) are removed from the checklist be- were centered beyond 190 m or their geo- cause recent revisions and investigations in- graphic distributions were centered south of volving these species (Garcfa-Gomez 1983, Cape Canaveral, Florida, are included in the Lemaitre & McLaughlin 1996, Fransen present checklist for this same reason. 2000) clearly indicate that they do not occur Twelve species, discovered and described in the region. One eastern Pacific species during the past 20 years, were also added (Notolopas lamellatus) was erroneously re- to the regional assemblage (Table 2). Four ported from the region by Rathbun (Wil- species (Trachycaris rugosa, Axius arma- liams 1984). Additionally, Leptochela ber- tus, Pagurus brevidactylus, Panopeus obe- mudensis was previously included in the sus) were added through a refined under- checklist because it was considered as like- standing of the systematics of species com- ly to occur in the region by Williams plexes with improved recognition and de- (1984). Since this species has never actually lineation of component species. One been recorded from the region, it has been nonindigenous species (Hemigrapsus san- removed from the checklist. guineus) was introduced and has become Better understanding of species concepts established in the region. An additional five and interspecific variation has necessitated species (Parapenaeus americanus, Scyllar- placement of some nominal species previ- ides aequinoctialis, Petrolisthes armatus, ously considered part of this regional as- Dromia erythropus, Clythrocerus nitidus) semblage into synonymy. For decapod have also become part of the regional as- crustaceans of the eastern United States, semblage through northward extension of eleven nominal species (Table 4) have been their geographic ranges. re-evaluated and determined not to repre- Six species have been removed from sent distinct species. Additionally, one fam- Williams (1984) checklist (Table 3). Four ily and one genus have also been placed species (.Pontonia margarita, Anisopagurus into synonymy (Table 4). pygmaeus, Iridopagurus caribbensis, and A significant proportion of systematic re- Hyas coarctatus alutaceus) once thought to search on western Atlantic decapod crusta- VOLUME 116, NUMBER 1 147

Table 2.—Summary of more recently described western Atlantic decapod Crustacea not previously listed in Williams (1984).

Year of descrip- Depth of Family Genus and species tion Geographic range occurrence

Disciadidae Discias vernbergi 1987 Georgia-west Florida 54—74 m Palaemonidae Pontonia manningi 2000 North Carolina-Gulf of Mexico, Ca- shallow-80 m ribbean Sea, eastern Atlantic Alpheidae Alpheus angulosus 2002 North Carolina-Gulf of Mexico, intertidal, shallow Mexico, Haiti Alpheidae Alpheus estuariensis 1984 Florida, Gulf of Mexico, Caribbean intertidal-22 m Sea, Brazil Callianassidae Necallianassa berylae 1998 South Carolina-Georgia 35-75 m Paguridae Anisopagurus hopkinsi 1996 Georgia-Gulf of Mexico 91-165 m Paguridae Iridopagurus reticulatus 1983 North Carolina-Florida, Bahamas, 1-38 m Greater and Lesser Antilles, northern South America Paguridae Pagurus maclaughlinae 1982 Georgia-Florida, Gulf of Mexico, 1-5 m Caribbean Paguridae Tomopagurus wassi 1981 Southeastern United States, Florida 75-360 m Straits, Gulf of Mexico, Caribbe- an-northern Brazil Homolidae Homola minima 1995 Massachussetts-Brazil 55-690 m Calappidae Cyclozodion tuberatum 1988 North Carolina-Bahamas-eastern 31-188 m Gulf of Mexico, Suriname Inachidae Stenorhynchus yangi 1989 Massachussetts-Gulf of Mexico-Su- 31-365 m riname

Table 3.—Summary of western Atlantic decapod crustacean species previously (Williams 1984) considered to be part of the fauna off the eastern United States but now known not to occur in the region. These species were removed from the checklist.

Species Range Reason for removal from checklist

Leptochela bermudensis Bermuda, Puerto Rico Previously considered likely to occur in Gurney, 1939 through the Lesser Antilles the region (Williams 1984). but has never been reported. Pontonia margarita Eastern Pacific Formerly thought to be a widespread spe- Smith, 1869 cies; with resolution of species complex now known to occur only in the Pacific Ocean (Fransen 2000). Anisopagurus pygmaeus Florida Keys, Cuba to Curasao Resolution of species systematics indicated (Bouvier, 1918) geographic range occurs beyond limits of region (Lemaitre & McLaughlin 1996). Iridopagurus caribbensis Miami, Florida to Colombia Resolution of species systematics indicated (A. Milne-Edwards & Bou- geographic range occurs beyond limits vier, 1893) of region (Garcfa-Gomez 1983). Hyas coarctatus alutaceus Eastern Pacific Resolution of species systematics indicated Brandt, 1851 geographic range occurs beyond limits of region (Squires 1990). Notolopas lamellatus Eastern Pacific Erroneously reported from the region Stimpson, 1871 (Williams 1984). 148 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

Table 4.—Summary of nominal taxa of western Atlantic decapod crustaceans placed in synonymy since publication of Williams (1984).

Nominal taxa appearing in Williams (1984) Senior synonym

Automate gardineri Automate dolichognatha Hippolyte curacaoensis Hippolyte obliquimanus Lebbeus zebra Lebbeus microceros Callianassa atlantica Gilvossius setimanus Manucomplanus corallinus Manucomplanus ungulatus Pagurus hendersoni Pagurus stimpsoni Hypoconcha sabulosa Hypoconcha parasitica Homola barbata (in part) Homola minima Cycloes bairdii Cryptosoma balguerii Mithrax pleuracanthus Mithrax hispidus Mithrax acuticornis Mithrax cornutus Family Tymolidae Family Cyclodorippidae Genus Parapandalus Genus Plesionika ceans is being conducted on higher-level re- of genera to accommodate western Atlantic lationships. Better understanding of rela- decapod species. Fifteen new genera have tionships based on identification of mono- been described since publication of Wil- phyletic groups has required reorganization liams' (1984) monograph, including one of taxa at higher levels (e.g., within super- genus each in the Penaeidae, Pandalidae, families, families, and genera) as well as Axiidae, and Paguridae, two in Callianas- elevating subfamilies and subgenera to fam- sidae, and nine brachyuran genera placed in ily and generic levels, respectively. In the seven different families (Table 5). Twenty- nearly 20-year period since Williams one species of decapods occurring in the (1984), one new superfamily (Processoi- region have been reassigned to genera other dea) and one new family (Anchistioididae) than those listed in Williams (1984). are recognized in the infraorder Caridea, Knowledge concerning well-known fau- and two subgenera have been elevated to nas, such as that of the decapod crustaceans genera {Farfantepenaeus and Litopenaeus) of the western Atlantic, is not static, and in the family Penaeidae, for shrimps occur- new discoveries, additional collecting, and ring in the region. Thalassinideans, previ- better understanding of systematic relation- ously placed in the section Thalassinidea ships will continue to improve our under- within infraorder Anomura, are now rec- standing of regional biodiversity. New ev- ognized at the level of infraorder. Within idence from adult morphology, fossils, lar- brachyuran taxa, three new subfamilies val development, and molecular genetics within the Raninidae are now recognized, has led to the reinterpretation of classical 17 former subfamilies (seven in Majidae, views of decapod crustacean relationships. four in Xanthidae, three in Grapsidae, one Hypotheses of phylogenetic relationships each in Raninidae, Calappidae, and Gone- are being proposed at a relatively fast rate placidae) have been elevated to family, two compared with previous time periods and subgenera elevated to genus {Mithraculus the taxonomic status of species continues to in family Majidae and Platylambrus in fam- be re-evaluated. Information gained from ily Parthenopidae), and one subspecies (of these systematic studies will undoubtedly Ethusa) elevated to species. Considerable result in better understanding of the species, changes in generic concepts involving west- provide refined hypotheses of relationships ern Atlantic decapod crustaceans have also among these taxa, and subsequently will necessitated the recognition and redefinition continue to improve our knowledge regard- VOLUME 116, NUMBER 1 149

Table 5.—Summary of western Atlantic decapod crustacean species assigned to new genera or reassigned to established genera since publication of Williams (1984).

Species name as appears in Species name as appears in Newly-described genus? Williams (1984) present compilation (Y/N)

Hymenopenaeus robustus Pleoticus robustus N Trachypenaeus constrictus Rimapenaeus constrictus Y Pandalus propinquus Atlantopandalus propinqvus Y Parapandalus willisi Plesionika willisi N Pontophilus gorei Philocheras gorei N Callianassa biformis Biffarius biformis Y Callianassa atlantica Gilvossius setimanus Y Callianassa major Callichirus major N Axiopsis jenneri Calaxius jenneri Y Pagurus piercei Goreopagurus piercei Y Dromidia antillensis Cryptodromiopsis antillensis N Clythrocerus perpusillus Deilocerus perpusillus Y Iliacantha intermedia Acanthilia intermedia Y Cryptopodia concava Celatopesia concava Y Geryon quinquedens Chaceon quinquedens Y Neopanope sayi Dyspanopeus sayi Y Pinnotheres chamae Gemmotheres chamae Y Pinnotheres maculatus Tumidotheres maculatus Y Pinnotheres ostreum Zaops ostreum N Pinnixa cristata Austinixa cristata Y Sesarma cinereum Armases cinereum Y

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{Hemigrapsus sanguineus: Grapsidae), in its Catalogue of Crustacea of Brazil. Museu Na- new habitat along the Atlantic coast of the Unit- cional, Rio de Janeiro. (Serie Livros n. 6). ed States: geographic distribution and ecolo- Melo-Filho, G. A. S. 1998. Malacostraca-Eucarida. gy.—ICES Journal of Marine Science 55:289- Chyrostilidae and Galatheidae. Pp. 393-397 in 298. P. S. Young, ed., Catalogue of Crustacea of Bra- . 2000. Natural history and biology of the zil. Museu Nacional, Rio de Janeiro. (Serie Liv- Asian shore crab Hemigrapsus sanguineus in ros n. 6). the western Atlantic: a review, with new infor- , & G. A. S. Melo. 1992. Reexamination of the mation. Pp. 193-199 in J. Pederson, ed., Marine material of Munida Leach (Crustacea: Anomu- bioinvasions: Proceedings of the First National ra: Galatheidae) collected by the H.M.S. Chal- Conference, January 24-27, 1999. MIT, MIT lenger (1872-1876) along the Brazilian Sea Grant College Program, Massachusetts. coast.—Proceedings of the Biological Society McLaughlin, P. A. 1981. Revision of Pylopagurus and of Washington 105:760-774. Tomopagurus (Crustacea: Decapoda: Paguri- Moriyasu, M., H. J. Squires, R. Campbell, & K. Ben- dae), with the descriptions of new genera and halima. 2001. Northern range extensions for species. Part I. Ten new genera of the Paguridae two decapod crustaceans, the inflated spiny crab and a redescription of Tomopagurus A. Milne Rochinia crassa (A. Milne-Edwards, 1879) and Edwards and Bouvier.—Bulletin of Marine Sci- the purplehead gamba prawn Aristeus antillen- ence 31:1-30. sis A. Milne-Edwards & Bouvier, 1909, in the . 1983. Hermit crabs-are they really polyphy- northwestern Atlantic.—Crustaceana 74:255- letic?—Journal of Crustacean Biology 3:608- 260. 621. Nations, J. D. 1975. The genus Cancer (Crustacea: . 1988. The rediscovery of Ceratopagurus Yo- Brachyura): systematics, biogeography and fos- koya and a new genus for Pagurus piercei Wass sil record.—Natural History Museum of Los (Decapoda, Paguroidea, Paguridae).—Crusta- Angeles County Science Bulletin 23:1-104. ceana 55:257-267. Ng, P. K. L„ & G. Rodriguez. 1986. New records of , & J. Haig. 1995. A new species of Goreo- Mimilambrus wileyi Williams, 1979 (Crustacea: pagurus McLaughlin (Decapoda: Anomura: Pa- Decapoda: Brachyura), with notes on the sys- guridae) from the Pacific, and a comparison tematics of the Mimilambridae Williams, 1979, with its Atlantic counterpart.—Proceedings of and Parthenopidae MacLeay, 1838, sensu Guin- the Biological Society of Washington 108:68- ot, 1978.—Proceedings of the Biological Soci- 75. ety of Washington 99:88-99. , & R. Lemaitre. 2001a. A new family for a Niem, V. H. 1996. Phylogenetic relationships among new genus and new species of hermit crab of American species of Sesarma (subgenus Ar- the superfamily Paguroidea (Decapoda: Ano- mases) (Brachyura, Grapsidae).—Crustaceana mura) and its phylogenetic implications.—Jour- 69:330-348. nal of Crustacean Biology 21:1062-1076. Perez Farfante, I. 1977a. American solenocerid , & . 2001b. Revision of Pylopagurus shrimps of the genera Hymenopenaeus, Hali- and Tomopagurus (Crustacea: Decapoda: Pa- poroides, Pleoticus, Hadropenaeus new genus, guridae), with descriptions of new genera and and Mesopenaeus new genus.—Fishery Bulle- species. Part VI. Pylopagurus A. Milne-Ed- tin 75:261-346. wards & Bouvier, 1891, Haigia McLaughlin, . 1977b. Range extensions of the shrimps So- 1981, and Pylopaguridium, a new genus.—Pro- lenocera necopina Burkenroad and Parapen- ceedings of the Biological Society of Washing- aeus americanus Rathbun (Crustacea, Decapo- ton 114:444-483. da, Penaeoidea).—Proceedings of the Biologi- McLay, C. L. 1993. Crustacea Decapoda: The sponge cal Society of Washington 90:597-599. crabs (Dromiidae) of New Caledonia and the . 1980. Revision of the penaeid shrimp genus Philippines with a review of the genera.—Me- Penaeopsis (Crustacea: Decapoda).—Fishery mories du Museum National d'Histoire Natu- Bulletin 77:721-763. relle 156:111-251. . 1988. Illustrated key to penaeoid shrimps of Melo, G. A. S. 1998. Malacostraca-Eucarida. Brach- commerce in the Americas.—NOAA Technical yura. Oxyrhyncha and Brachyrhyncha. Pp. 455- Report, National Marine Fisheries Service, 64: 515 in P. S. Young, ed., Catalogue of Crustacea 1-32. of Brazil. Museu Nacional, Rio de Janeiro. (Ser- , & B. F. Kensley. 1997. Penaeoid and serges- ie Livros n. 6). toid shrimps and prawns of the world. Keys and , M. F. A. Torres, & O. Campos, Jr. 1998. Ma- diagnoses for the families and genera.—Me- lacostraca-Eucarida. Brachyura. Dromiacea and mories du Museum National d'Histoire Natu- Oxystomata. Pp. 439-454 in P. S. Young, ed., relle 175:1-233. VOLUME 116, NUMBER 1 155

Pipitone, C., F. Badalamenti, & A. Sparrow. 2001. capoda: Thalassinidea).—Zoologische Verhan- Contribution to the knowledge of Percnon gib- delingen, Leiden 326:1-152. besi (Decapoda, Grapsidae), an exotic species , & M. de Saint Laurent. 1989. A check list of spreading rapidly in Sicilian waters.—Crusta- Axiidae (Decapoda, Crustacea, Thalassinidea, ceana 74:1009-1017. Anomura), with remarks and in addition de- Pohle, G., & F. Marques. 2000. Larval stages of Par- scriptions of one new subfamily, eleven new adasygyius depressus (Bell, 1835) (Crustacea: genera and two new species.—Naturalists 3:1- Decapoda: Brachyura: Majidae) and a phylo- 104. genetic analysis for 21 genera of Majidae.— Sarver, S. K., D. W. Freshwater, & P. J. Walsh. 2000. Proceedings of the Biological Society of Wash- The occurrence of the provisional Brazilian sub- ington 113:739-760. species of spiny lobster (Panulirus argus wes- Poore, G. C. B. 1994. A phylogeny of the families of tonii) in Florida waters.—Fishery Bulletin 98: Thalassinidea (Crustacea: Decapoda) with keys 870-873. to families and genera.—Memoirs of the Mu- , J. D. Silberman, & P. J. Walsh. 1998. Mito- seum of Victoria 54:79-120. chondrial DNA sequence evidence supporting Powers, L. W. 1977. A catalogue & bibliography to recognition of two subspecies or species of the the crabs (Brachyura) of the Gulf of Mexico.— Florida spiny lobster Panulirus argus.—Journal Contributions in Marine Science 20 (supple- of Crustacean Biology 18:177-186. ment): 1-190. Scelzo, M. A. 2001. First record of the portunid crab Ramos-Porto, M., & P. A. Coelho. 1998. Malacostraca. Arenaeus cribrarius (Lamarck, 1818) (Crusta- Eucarida. Caridea (Alpheoidea excluded). Pp. cea: Brachyura: Portunidae) in marine waters of 325-350 in P. S. Young, ed., Catalogue of Crus- Argentina.—Proceedings of the Biological So- tacea in Brazil. Museu Nacional, Rio de Janeiro. ciety of Washington 114:605-610. (Serie Livros n. 6). Schubart, C. D„ J. A. Cuesta, R. Diesel, & D. L. Feld- Rathbun, M. J. 1925. The spider crabs of America.— er. 2000. Molecular phylogeny, taxonomy, and United States National Museum Bulletin 129: evolution of nonmarine lineages within the 1-613. American grapsoid crabs (Crustacea: Brachyu- . 1930. The cancroid crabs of America of the ra).—Molecular Phylogenetics and Evolution families Euryalidae, Portunidae, Atelecyclidae, 15:179-190. Cancridae and Xanthidae.—United States Na- , J. E. Neigel, & D. L. Felder. 2000. Molecular tional Museum Bulletin 152:1-609. phylogeny of mud crabs (Brachyura: Panopei- Rieger, P. J. 1998. Malacostraca-Eucarida. Paguroidea. dae) from the northwestern Atlantic and the role Pp. 413-429 in P. S. Young, ed., Catalogue of Crustacea in Brazil. Museu Nacional, Rio de of morphological stasis and convergence.—Ma- Janeiro. (Serie Livros n. 6). rine Biology 137:11-18. Rodrigues, S. de A., & R. M. Shimizu. 1998. Mala- Schweitzer, C. E., & R. M. Feldmann. 2000. New spe- costraca-Eucarida. Thalassinidea. Pp. 379-385 cies of calappid crabs from western North in P. S. Young, ed., Catalogue of Crustacea in America and reconsideration of the Calappidae Brazil. Museu Nacional, Rio de Janeiro. (Serie sensu lato.—Journal of Paleontology 74:230- Livros n. 6). 246. Rodriguez, B. 1993. Shrimps. Pp. 107-131 in K. Car- Silberman, J. D„ S. K. Sarver, & P. J. Walsh. 1994. penter, W. Fischer, and L. Garibaldi, eds., Field Mitochondrial DNA variation and population guide to the commercial marine and brackish- structure in the spiny lobster Panulirus argus.— water resources of the northern coast of South Marine Biology 120:601-608. America. FAO, Rome. Squires, H. J. 1990. Decapod Crustacea of the Atlantic Rodriguez-Almaraz, G., A. Leija-Tristan, & R. Men- coast of Canada.—Canadian Bulletin of Fish- doza. 2000. Records of caridean shrimps (Crus- eries and Aquatic Sciences 221:1-532. tacea: Decapoda) from the coasts of the Mexi- . 1992. Recognition of Pandalus eons Maka- can Pacific Ocean, Gulf of Mexico and Mexican rov, 1935, as a Pacific species not a variety of Caribbean.—Bulletin of Marine Science 67: the Atlantic Pandalus borealis Kr0yer, 1838 857-867. (Decapoda, Caridea).—Crustaceana 63:257- Rosenberg, M. S. 2001. The systematics and taxonomy 262. of fiddler crabs: A phylogeny of the genus Stehlik, L. L„ C. L. MacKenzie, Jr., & W. W. Morse. Uca.—Journal of Crustacean Biology 21:839- 1991. Distribution and abundance of four brach- 869. yuran crabs on the northwest Atlantic shelf.— Sakai, K. 1999. Synopsis of the family Callianassidae, Fishery Bulletin 89:473-492. with keys to subfamilies, genera and species, Sternberg, R. von, & N. Cumberlidge. 1998. Taxic re- and the description of new taxa (Crustacea: De- lationships within the Grapsidae MacLeay, 156 PROCEEDINGS OF THE BIOLOGICAL SOCIETY OF WASHINGTON

1838 (Crustacea: Decapoda: Eubrachyura).— Wagner, H. P. 1990. The genera Mithrax Latreille, Journal of Comparative Biology 3:1 15-136. 1818 and Mithraculus White, 1847 (Crustacea: Strasser, K. M., & W. W. Price. 1999. An annotated Brachyura: Majidae) in the western Atlantic checklist and key to hermit crabs of Tampa Bay, Ocean.—Zoologische Verhandelingen (Leiden) Florida, and surrounding waters.—Gulf Re- 264:1-65. search Reports 11:35-50. Wass, M. L. 1963. New species of hermit crabs (De- Strenth, N. E., & F. A. Chace, Jr. 1995. New records capoda, Paguridae) from the western Atlan- of natant decapods (Crustacea: Palaemonidae) tic.—Crustaceana 6:133-157. from the south Texas coast.—Texas Journal of Wigley, R. L„ R. B. Theroux, & H. E. Murray. 1975. Science 47:315-318. Deep-sea red crab, Geryon quinquedens, survey Tavares, M. 1991a [1990]. Especes nouvelles de Cy- off northeastern United States.—Marine Fish- eries Review 37:1—21. clodorippoidea Ortmann et remarques sur les Williams, A. B. 1965. Marine decapod crustaceans of genres Tymolus Stimpson et Cyclodorippe A. the Carolinas.—Fishery Bulletin 65:1-298. Milne Edwards (Crustacea, Decapoda, Brach- . 1983. The mud crab, Panopeus herbstii, s.l. yura).—Bulletin du Museum National partition into six species (Decapoda: Xanthi- d'Histoire Naturelle, Paris, 4e series, 12, Section dae).—Fishery Bulletin 81:863-882. A, nos 3-4:623-648. . 1984. Shrimps, lobsters, and crabs of the At- . 1991b. Redefinition des genres Rochinia A. lantic coast of the eastern United States, Maine Milne Edwards, Sphenocarcinus A. Milne Ed- to Florida. Smithsonian Institution Press, Wash- wards et Oxypleurodon Miers, et etablissement ington, D.C., 550 pp. du genre Nasutocarcimis gen. nov. (Crustacea, . 1988. Notes on decapod and euphausiid crus- Brachyura, Majidae).—Bulletin du Museum taceans, continental margin, western Atlantic, e National d'Histoire Naturelle, Paris, 4 series, Georges Bank to western Florida, USA.—Fish- os 13, Section A, n 1-2:159-179. ery Bulletin 86:67-76. . 1993. Description preliminaire de quatre nou- . 1993. Mud shrimps, Upogebiidae, from the veaux genres et trois nouvelles especes de Cy- western Atlantic (Crustacea: Decapoda: Thal- clodorippoidea Americains (Crustacea, Deca- assinidea).—Smithsonian Contributions to Zo- poda, Brachyura).—Vie Milieu 43:137-144. ology 544:1-77. . 1996. Revision systematique des Cyclodor- , & C. A. Child. 1989. Comparison of some ippidae americains (Crustacea, Decapoda, genera and species of box crabs (Brachyura: Brachyura).—Bulletin du Museum National Calappidae), southwestern North Atlantic, with d'Histoire Naturelle, Paris, 4e series, 18, Section description of a new genus and species.—Fish- A, nos 1-2:233-295. ery Bulletin 87:105-121. Tucker, A. B. 1998. Systematics of the Raninidae , & D. L. Felder. 1986. Analysis of stone crabs: (Crustacea: Decapoda: Brachyura), with ac- Menippe mercenaria (Say), restricted, and a counts of three new genera and two new spe- previously unrecognized species described (De- cies.—Proceedings of the Biological Society of capoda: Xanthidae).—Proceedings of the Bio- Washington 111:320-371. logical Society of Washington 99:517-543. Tudge, C. C., G. C. B. Poore, & R. Lemaitre. 2000. , & J. J. McDermott. 1990. An eastern United Preliminary phylogenetic analysis of generic re- States record for the western Indo-Pacific crab, lationships within the Callianassidae and Cten- Hemigrapsus sanguineus (Crustacea: Decapo- ochelidae (Decapoda: Thalassinidea: Callianas- da: Grapsidae).—Proceedings of the Biological soidea).—Journal of Crustacean Biology 20 Society of Washington 103:108-109. (Special no. 2): 129-149. , & R. Wahle. 1992. Distinguishing juvenile stages of Jonah and Atlantic rock crabs, Cancer Udekem d'Acoz, C. d'. 1997. Redescription of Hip- borealis and C. irroratus (Decapoda: Cancri- polyte ohliquimanus Dana, 1852, and compari- dae).—Journal of Crustacean Biology 12:464- son with Hippolyte williamsi Schmitt, 1924 466. (Decapoda, Caridea).—Crustaceana 70:469- , & R. L. Wigley. 1977. Distribution of deca- 479. pod Crustacea off northeastern United States Vargas, R., & J. Cortes. 1999. Biodiversidad marina based on specimens at the Northeast Fisheries de Costa Rica: Crustacea: Decapoda (Penaeo- Center, Woods Hole, Massachusetts.—NOAA idea, Sergestoidea, Stenopodidea, Caridea, Technical Report NMFS Circular 407:1-44. Thalassinidea, Palinura) del Caribe.—Revista de Biologia Tropical 47:877-885. Note Added in Proof Veloso, V. G. 1998. Malacostraca-Eucarida. Porcellan- idae. Pp. 399-405 in P. S. Young, ed. Catalogue Recent revision of the Albuneidae (Boy- of Crustacea in Brazil. Museu Nacional, Rio de ko 2002) provided new information for Janeiro. (Serie Livros n. 6). three species included in the present check- VOLUME 116, NUMBER 1 157 list. Western Atlantic specimens identified ger considered part of the regional decapod as Albunea paretii Guerin-Meneville, 1853 assemblage. in Williams (1984) included two species, A. paretii and A. catherinae Boyko, 2002. Of Family Albuneidae Stimpson, 1858 these, only A. catherinae occurs in the re- • Albunea catherinae Boyko, 2002 gion (see account below). Albunea paretii Remarks.—Boyko (2002:343) provided a ranges from the Florida Keys southward description, diagnosis, illustrations, and size through the Caribbean to Rio Grande do information for this species. Albunea cath- Sul, Brazil, and should be removed from erinae occurs at depths less than 64 m. the checklist. According to Boyko (2002), Known range.—Western Atlantic: Vir- all eastern Atlantic references to A. paretii ginia to Palm Beach County, Florida; Col- refer to A. elegans. Boyko (2002) included lier County, Florida, through the Gulf of additional distributional information for Mexico, to southern Texas; absent from the Lepidopa websteri that extends this species Florida Keys (Boyko 2002). range to Texas. Based on new data for A. paretii and A. Literature Cited catherinae, 52 species are new to the Boyko, C. B. 2002. A worldwide revision of the recent checklist. Thirteen of these have been dis- and fossil sand crabs of the Albuneidae Stimp- covered and described since publication of son and Blepharipodidae. new family (Crusta- cea: Decapoda: Anomura: Hippoidea).—Bulle- Williams (1984), and seven species previ- tin of the American Museum of Natural History ously recorded from the region are no lon- 272:1-396.