Hydrobiologia 287: 95-104, 1994. D. C. Culver & J. R. Holsinger (eds), Biogeography of Subterranean : the Effects of Different Scales. 95 © 1994 Kluwer Academic Publishers. Printed in Belgium.

Biogeography of subterranean decapods in North and Central America and the Caribbean region (, Astacidea, Brachyura)*

Horton H. Hobbs III Department of Biology, Wittenberg University, P.O. Box 720, Springfield, Ohio 45501-0720, USA

Key words: , biogeography, caves

Abstract

A significant number of decapod crustaceans (81 troglobites and 58 other cavernicoles) has been described from various subterranean waters in North and Central America (United States south to Costa Rica) and from the islands in the western north Atlantic and the Caribbean Sea, posing puzzling questions concerning their evolution and biogeography. Of these troglobitic species, 36 are shrimps (1 procarid, 11 atyids, 2 agostocarids, 15 palaemonids, 2 alpheids, 5 hippolytids), 35 are cambarid , and 10 are crabs (1 grapsid, 7 pseudothelphusids, 2 trichodactylids). They are known to occur in caves, springs, cenotes, blue holes, anchialine environments, and various crevicular habitats in localized areas throughout the region. Many, if not the majority, of the troglobites appear to have arisen independently from epigean progenitors rather than sharing common subterranean precursors.

Introduction tinental drift) and dispersal biogeography as well as isolated dispersal phenomena. This paper represents The decapod crustaceans of North America and an attempt to address briefly such explanations and Mesoamerica (Mexico, Central America, West Indies) to summarize the current knowledge of the distribu­ are an extremely diverse group of of mixed tion of the cavernicolous decapod species of 11 fami­ geographical origin. A significant number of species lies (shrimps, crayfishes, crabs) in North America and has been discovered inhabiting various subterranean Mesoamerica (see Fig. 1 and Table 1). No attempt is waters, the distribution of which often poses puz­ made to treat the possible origins and individual ranges zling questions concerning their evolution and bio­ of all members of these families. geography. In Mesoamerica, caves and other subter­ ranean habitats have provided opportunities for marine List of cavernicolous decapods in North America and stocks and refugia for others that invaded continental Mesoamerica waters. Also there and in North America, representa­ tives of several stocks that became established in fresh The following is a list of shrimps, crayfishes, and crabs waters invaded subterranean waters opportunistically known to occur in caves, springs, cenotes, blue holes, and became obligate cavernicoles. For many, if not anchialine, and crevicular habitats, distributed from the most, of the decapods treated herein, the acquisition United States to Central America and on the islands of of troglomorphic characters associated with obligate the western north Atlantic and Caribbean from Bermu­ subterranean life represents parallel development (con­ da to Bonaire. All species listed are aquatic troglobites vergence). Explanations for the disjunct distribution (stygobionts) unless otherwise indicated. of related species in these crustaceans commonly have involved discussions concerning vicariance (e.g., con-

* Presented at the Symposium on the "Biogeography of Subter­ ranean Crustaceans: the Effects of Different Scales" (June 1992; Charleston, South Caroina, USA). Manuscript prepared for print by J. R. Holsinger and D. C. Culver. ON

United States Bermuda

ATLANTIC OCEAN

** Barbados 9 fc

PACIFIC OCEAN

100 50°

Fig. 1. Map of area treated in text, including parts of North America and Mesoamerica. 97

Table 1. Distribution by family of cavernicolous decapods of North America and Mesoamerica

* CARIDEA (Shrimps): Procarididae (Procaris) - Bermuda, Mexico; (Ascension, Hawaii) (Atya, Palaemonias, , Xiphocaris) Barbados, Jamaica, United States, Mexico, Cuba, Bahamas, Caicos Islands, Bermuda, Barbuda, Isla Mona, Puerto Rico, Dominican Republic; (Ascension, Galapagos) Agostocarididae (Agostocaris) - Mexico, Bahamas, Turks Islands, Caicos Islands Palaemonidae (Cryphiops, Creaseria, Macrobrachium, Neopalaemon, Palaemonetes, Troglocubanus - Mexico, Costa Rica, Jamaica, United States, Bonaire, Curacao, Dominican Republic, Cuba, Puerto Rico, Isla de Pinos Alpheidae (Alpheopsis, Potamalpheops) - Mexico, Bermuda Hippolytidae (Barbouria, Calliasmata, Janicea, Somersiella, Yagerocaris) - Bahamas, Caicos Islands, Cayman Islands, Bermuda, Cuba, Dominican Republic, Antigua, Mexico

* ASTACIDEA (Crayfishes): (Cambarus, Orconectes, Pwcambarus, Troglocambarus) Canada, United States, Mexico, Belize, Guatemala, Cuba

* BRACHYURA (Crabs): Gecarcinidae (Cardisoma, Gecarcinus) - Cuba, Jamaica, Puerto Rico Grapsidae (Sesarma) - Jamaica, Cuba, Guatemala Pseudothelphusidae (Epilobocera, Isabellagordonia, Odontothelphusa, Phrygiopilus, Potamocarcinus, Pseudothelphusa, Ptychophallus, ?Raddus, Stygothelphusa, Tehuana, Typhlopseudothelphusa, Zilchia) - Mexico, Cuba, Puerto Rico, Guatemala, Belize, Costa Rica Trichodactylidae (Avotrichodactylus, Rodriguezia, Trichodactylus) - Mexico

CLASS CRUSTACEA Superfamily Atyoidea

Order Decapoda Family Atyidae SUBORDER PLEYCYEMATA Atya brachyrhinus Hobbs & Hart - cave - Barbados Infraorder Caridea Atya innocous (Herbst) - trogloxene -caves - Jamaica: Saint Elizabeth and Saint Mary parishes Superfamily Procaridoidea Atya lanipes Holthuis - trogloxene - caves - Jamaica: Saint Mary Parish Family Procarididae Palaemonias alabamae Smalley - caves - USA: Madi­ Procaris chacei Hart & Manning - Green Bay Cave son County, Alabama (anchialine habitat) - Bermuda Palaemonias ganteri Hay - caves - USA: Edmonson, Procaris n.sp. - anchialine habitat - Mexico: Quintana Hart counties, Kentucky Roo Typhlatya campecheae Hobbs & Hobbs - caves - Mex­ Two additional species are known from anchialine lava ico: Campeche pools: P. ascensionis Chace & Manning (1972), from Typhlatya consobrina Botosaneanu & Holthuis - caves Ascension Island in the South Atlantic Ocean and - Cuba: Camaguey, Pinar del Rio provinces P. hawaiana Holthuis (1973) on Maui and Hawaii, Hawaiian Islands 98

Typhlatya garciai Chace - caves, anchialine habitats - Macrobrachium lucifugum Holthuis - troglophile - Cuba: Oriente, Pinar del Rio provinces; Caicos Islands: caves and anchialine habitats - Bonaire, Curacao, Providenciales Dominican Republic, Jamaica: Saint Ann Parish, Typhlatya iliffei Hart & Manning - Tucker's Town Cave Cuba: Oriente, Pinar del Rio provinces; Puerto Rico (anchialine cave) - Bermuda Macrobrachium villalobosi Hobbs - Cueva del Naci- Typhlatya mitchelli Hobbs & Hobbs - caves, cenotes - mento del Rio San Antonio - Mexico: Oaxaca Mexico: Quintana Roo and Yucatan Neopalaemon nahuatlus Hobbs - Cueva del Guano - Typhlatya monae Chace - well, caves - Barbuda, Isla Mexico: Oaxaca Mona, Puerto Rico, Dominican Republic Palaemonetes (Alaocaris) antrorum (Benedict) - wells Typhlatya pearsei Creaser - caves, cenotes - Mexico: and caves - USA: Hays, Uvalde counties,Texas Campeche, Quintana Roo, Yucatan Palaemonetes (Alaocaris) holthuisi (Strenth) - cave - Typhlatya taina Estrada and Gomez - Cueva de la India USA: Hays County, Texas - La Habana, Cuba Palaemonetes (Palaemonetes) cummingi Chace - Typhlatya n.sp. [Kensley 1988:688] - Mexico: Yucatan Squirrel Chimney - USA: Alachua County, Peninsula Palaemonetes (Palaemonetes) paludosus (Gibbes) The genus is represented by two additional species: trogloxene - cave - USA: Leon County, Florida T. galapagensis Monod & Cals (1970), stygobiont Troglocubanus calcis (Rathbun) - caves - Cuba: Cam- from anchialine habitat on Isla Santa Cruz, Galapa­ agiiey and La Habana provinces gos Islands and T. rogersi Chace & Manning (1972) Troglocubanus eigenmanni (Hay) - caves - Cuba: Pinar from saline, anchialine pools on Ascension Island del Rio, La Habana, Matanzas provinces; Isla de Pinos Xiphocaris elongata (Guerin-Meneville) - trogloxene Troglocubanus gibarensis (Chace)-caves - Cuba: Ori­ - caves - Jamaica, Puerto Rica ente and Las Villas provinces Troglocubanus inermis (Chace) - caves - Cuba: Cam- Superfamily Rhynchocinetoidea aggiiey and La Habana provinces Troglocubanus jamaicensis Holthuis - cave - Jamaica: Family Agostocarididae Saint Mary Parish Agostocaris bozanici Kensley - Xcan-ha Cenote - Mex­ Troglocubanus perezfarfanteae Villalobos F. -caves - ico: Quintana Roo (Cozumel Island) Mexico: San Luis Potosf, Tamaulipas Agostocaris williamsi Hart & Manning - blue hole, anchialine caves - Bahamas: Cat and Grand Bahama Superfamily Alpheoidea Islands; Caicos Islands: Providenciales Family Alpheidae Superfamily Palaemonoidea Automate dolichognatha De Man - Tucker's Town Cave (anchialine cave) - Bermuda Family Palaemonidae Potamalpheops stygicola (Hobbs)-Cuevadel Nacimien Cryphiops (Bithynops) luscus (Holthuis) - Grutas del to del Rio San Antonio - Mexico: Oaxaca Arco - Mexico: Chiapas Cryphiops (Bithynops) perspicax (Holthuis) Cenote La Family Hippolytidae Cueva - Mexico: Chiapas Barbouria cubensis (von Martens) - caves, blue holes, Creaseria morleyi (Creaser) - caves - Mexico: Yucatan, anchialine habitats - Bahamas: Abaco Island, San Sal­ Quintana Roo vador Island; Caicos Islands: Providenciales; Cay­ Macrobrachium acherontium Holthuis - caves- Mexi­ man Islands: Cayman Brae; Bermuda; Cuba: Habana, co: Tabasco Matanzas, Oriente provinces Macrobrachium carcinus (Linnaeus) - troglophile- Calliasmata rimolii Chace - anchialine cave - Domini­ caves and/or springs - Costa Rica: Puntarenas can Republic: Puerto Plata Province Province; Jamaica: Manchester and Saint Ann parish­ Janicea antiguensis (Chace) - caves, anchialine caves es; Mexico: Chiapas, San Luis Potosf; USA: Marion - Antigua; Bahamas: Grand Bahama; Bermuda; Mex­ and Putnam counties, Florida ico: Quintana Roo (Cozumel Island) Macrobrachium heterochirus (Wiegmann) - troglophile; Somersiella sterreri Hart & Manning - anchialine caves cave - Jamaica: Saint Ann Parish - Bermuda; Mexico: Quintana Roo (Cozumel Island) 99

Yagerocaris cozumel Kensley - cenote, anchialine cave Cambarus (Jugicambarus) cryptodytes Hobbs - caves - Mexico: Quintana Roo (Cozumel Island) - USA: Jackson and Washington counties, Florida and Decatur County, Georgia Cambarus (Jugicambarus) setosus Faxon - caves - Infraorder Astacidea USA: Christian, Dade, Greene, Jasper, Lawrence, Newton, and Stone counties, Missouri; Delaware and Superfamily Astacoidea Mayes counties, Oklahoma Cambarus (Jugicambarus) tartarus Hobbs & Cooper Family Cambaridae - Stansberry-January Cave System, - USA: Delaware Subfamily Cambarinae County, Oklahoma Cambarus (Aviticambarus) hamulatus (Cope) -caves - Cambarus (Jugicambarus) zophonastes Hobbs & USA: Alabama, Tennessee Bedinger - Hell Creek Cave - USA: Stone County, Cambarus (Aviticambarus) jonesi Hobbs & Barr - Arkansas caves - USA: northeastern Alabama Cambarus (Puncticambarus) nerterius Hobbs - caves Cambarus {Cambarus) sp. - troglophile? - caves - - USA: Greenbrier and Pocahontas counties, West Vir­ USA: southwestern Virginia; reported by Holsinger ginia (1964:62) as 'Cambarus bartonii ssp.' Cambarus (Puncticambarus) robustus Girard - Cambarus (Cambarus) bartonii bartonii (Fabricius)- troglophile? - surface streams, cave streams - USA: troglophile - surface waters, springs, and caves - USA: New York Alabama, Georgia, Virginia, West Virginia Orconectes (Buannulifictus) meeki brevis Williams - Cambarus (Cambarus) bartonii cavatus Hay - trogloxene? - surface streams, caves - USA: Arkansas, troglophile - surface streams and springs - USA: Geor­ Oklahoma gia, Kentucky, Ohio, Tennessee, Virginia Orconectes (Buannulifictus) palmeri longimanus Cambarus (Depressicambarus) latimanus (LeConte) (Faxon)-trogloxene? - surface lotic habitats; cave, - trogloxene? - surface burrows and streams, cave spring - USA: Pontotoc County, Oklahoma streams - USA: Chatooga, Dade, Walker counties, Orconectes (Crockerinus) propinquus (Girard) - Georgia trogloxene? - surface streams and rocky shores of Cambarus (Depressicambarus) striatus Hay - troglox­ lakes; cave - USA: Lawrence County, Indiana ene? - surface burrows and streams, cave streams - Orconectes (Gremicambarus) immunis (Hagen) - USA: Chatooga, Walker counties, Georgia trogloxene? - surface lentic and sluggish lotic habi­ Cambarus (Erebicambarus) hubbsi Creaser - troglox­ tats; cave - USA: Lawrence County, Indiana; Canada ene? - surface streams, burrows, caves - USA: north­ Orconectes (Gremicambarus) virilis (Hagen) - troglox­ eastern Arkansas and southeastern Missouri ene? - surface lakes and streams - cave - USA: Alabama Cambarus (Erebicambarus) hubrichti Hobbs - springs Orconectes (Orconectes) australis australis (Rhoades) and caves - USA: Missouri - caves - USA: Alabama, Tennessee, Cambarus (Erebicambarus) laevis Faxon - troglophile Orconectes (Orconectes) australis packardi Rhoades - surface streams, springs, and caves - USA: Indiana, caves - USA: Kentucky Kentucky Orconectes (Orconectes) incomptus Hobbs & Barr - Cambarus (Erebicambarus) ornatus Rhoades - caves - USA: Jackson County, Tennessee trogloxene? - intermittent surface streams, springs, Orconectes (Orconectes) inermis inermis Cope - caves cave - USA: Kentucky USA: Indiana, Kentucky Cambarus (Erebicambarus) tenebrosus Hay Orconectes (Orconectes) inermis testii (Hay) - caves troglophile - surface streams, caves and springs - USA: USA: Monroe County, Indiana Alabama, Kentucky, Tennessee Orconectes (Orconectes) pellucidus (Tellkampf) - Cambarus (Jugicambarus) aculabrum Hobbs & caves - USA: Kentucky, Tennessee Brown - caves - USA: Benton Co., Arkansas Orconectes (Procericambarus) luteus (Creaser) - Cambarus (Jugicambarus) dubius Faxon - trogloxene? trogloxene? - surface lotic habitats; cave - USA: Mis­ - surface primary burrower, cave - USA: Russell Coun­ souri ty, Virginia Orconectes (Procericambarus) neglectus neglectus (Faxon) - trogloxene? - surface rocky streams; caves - 100

USA: southwestern Missouri; Delaware County, Okla­ {Ortmannicus) franzi Hobbs & Lee - homa caves USA: Marion County, Florida Orconectes {Procericambarus) punctimanus (Creaser) Procambarus {Ortmannicus) horsti Hobbs & Means- trogloxene? - surface lotic habitats; cave - USA: Mis­ springs and caves - USA: Jefferson, Leon, and Wakulla souri counties, Florida Orconectes spp. - two undescribed troglobitic species Procambarus {Ortmannicus) leitheuseri Franz & from caves in northern Alabama Hobbs-caves - USA: Hernando and Pasco counties, Procambarus {Austrocambarus) mirandai Villalobos- Florida troglophile - caves - Mexico: Chiapas Procambarus {Ortmannicus) lucifugus alachua (Hobbs) Procambarus {Austrocambarus) niveus Hobbs & Vil- caves - USA: Alachua, Gilchrist, Levy, and Marion lalobos - Cuevas de Santo Tomas - Cuba: Pinar del counties, Florida Rio Procambarus {Ortmannicus) lucifugus lucifugus Procambarus {Austrocambarus) oaxacae oaxacae (Hobbs) - caves - USA: Citrus, Hernando, Marion Hobbs-Cueva del Guano - Mexico: Oaxaca counties, Florida Procambarus {Austrocambarus) oaxacae reddelli Procambarus {Ortmannicus) orcinus Hobbs & Means- Hobbs-caves - Mexico: Oaxaca, Veracruz caves - USA: Leon and Wakulla counties, Florida Procambarus {Austrocambarus)pilosimanus (Ortmann)- Procambarus {Ortmannicus)pallidus (Hobbs) - caves- trogloxene? - surface waters and caves - Belize, USA: Alachua, Columbia, Suwannee counties, Florida Guatemala, and Mexico: Chiapas and Quintana Roo Procambarus {Ortmannicus) toltecae Hobbs - troglox­ Pwcambarus {Austrocambarus) rodriguezi Hobbs - ene? - caves - Mexico: Hidalgo, San Luis Potosi, Cueva de Ojo de Agua Grande - Mexico: Veracruz Tamaulipas Procambarus {Austrocambarus) sbordonii Hobbs - Procambarus {Ortmannicus) villalobosi Hobbs troglox­ troglophile - Cueva del Nacimiento de Rio S.Domingo ene? - surface lotic habitats; cave - Mexico: San Luis - Mexico: Chiapas Potosi Procambarus {Girardiella) simulans (Faxon) - Procambarus {Ortmannicus) attiguus. Hobbs and troglophile - surface lotic, lentic, burrow habitats; Franz (1992) - cave - USA: Marion County, Florida caves - USA: Greer, Major, Murray, Washita, Wood­ Procambarus {Remoticambarus) pecki Hobbs - caves ward counties, Oklahoma; Burnet County, Texas - USA: Colbert, Lauderdale, and Morgan counties, Procambarus {Leconticambarus) milleri Hobbs - well Alabama in Miami - USA: Dade County, Florida Procambarus {Scapulicambarus) clarkii (Girard) - Procambarus {Lonnbergius) acherontis (Lonnberg)- trogloxene? - surface lentic, lotic, burrow habitats; springs and caves - USA: Orange and Seminole coun­ caves - USA: Texas ties, Florida Procambarus {Scapulicambarus) paeninsulanus (Fax­ Procambarus {Lonnbergius) morrisi Hobbs & Franz- on) - troglophile? - surf ace lotic, lentic, burrow habi­ Devil's Sink - USA: Putnam County, Florida tats; caves - USA: Florida Procambarus {Ortmannicus) acutus acutus (Girard) Procambarus {Scapulicambarus) xilitlae Hobbs & trogloxene? - surface lentic, sluggish lotic habitats; Grubbs - Hoya de las Guaguas - Mexico: San Luis caves - USA: Texas Potosi Procambarus {Ortmannicus) acutus cuevachicae Procambarus {Villalobosus) cuetzalanae Hobbs - (Hobbs)-troglophile - surface waters and caves - Mexi­ troglophile? - caves and springs - Mexico: Puebla co: Hidalgo, Puebla, San Luis Potosi, Tamaulipas, and Procambarus {Villalobosus) xochitlanae Hobbs - caves Veracruz - Mexico: Puebla Procambarus {Ortmannicus) delicatus Hobbs & Troglocambarus maclanei Hobbs - caves - USA: Franz-Alexander Spring and Cave - USA: Lake Coun­ Alachua, Citrus, Hernando, Marion, Suwannee coun­ ty, Florida ties, Florida Procambarus {Ortmannicus) erythrops Relyea & Sutton-caves - USA: Suwannee County, Florida Procambarus {Ortmannicus) fallax (Hagen) - troglox­ ene? - surface lotic, lentic environvents, burrows, cave- USA: Florida 101

Infraorder Brachyura Pseudothelphusa puntarenas Hobbs III - troglophile - cave - Costa Rica: Puntarenas Province Superfamily Grapsidoidea Pseudothelphusa sonorae Rodriguez & Smalley - trogloxene? - cave - Mexico: Sonora Family Gecarcinidae Pseudothelphusa sp. - cave - Mexico: Guerrero; report­ Cardisoma guanhumi Latreille - trogloxene? - cave ed by Hoffman et al. (1986:175,186,213,237) - Cuba: La Habana, Las Villas, Oriente provinces; Ptychophallus montanus (Rathbun) - troglophile - cave Jamaica: Clarendon Parish; Puerto Rico - Costa Rica: Puntarenas Province Gecarcinus ruricola (Linnaeus) - trogloxene - cave ?Raddus sp. - cave - Mexico: Chiapas; reported by - Cuba: Las Villas and Oriente provinces; Jamaica: Rodriguez & Hobbs (1989:397) Clarendon Parish Stygothelphusa lopezformenti Alvarez and Villalobos - cave - Mexico: Oaxaca Family Grapsidae Tehuana complanata Rodriguez - troglophile? - cave - Sesarma (Sesarma) verleyi Rathbun - caves - Jamaica: Mexico: San Luis Potosi, Veracruz Manchester, Saint Ann, Saint Catherine, Saint Eliza­ Typhlopseudothelphusa acanthochela Hobbs - cave - beth, Saint James, Saint Mary parishes Belize Sesarma sp. - trogloxene? - cave - Cuba: Oriente Typhlopseudothelphusa hyba Rodriguez & Hobbs - Province caves - Mexico: Chiapas Sesarma (Holometopus) miersii Rathbun - trogloxene? Typhlopseudothelphusa iuberthiei Deboutteville - cave - cave - Jamaica - Guatemala: Alta Verapaz Province Sesarma {Holometopus) roberti H. Milne Edwards- Typhlopseudothelphusamitchelli Deboutteville - caves trogloxene? - cave - Guatemala - Guatemala: Alta Verapaz Province Sesarma (Sesarma) bidentatum Benedict - trogloxene? Typhlopseudothelphusa mocinoi (Rioja) - caves - Mex­ - cave - Jamaica: Saint Ann, Saint Elizabeth parishes ico: Chiapas Zilchiafalcata Rodriguez & Hobbs - cave - Guatemala Family Pseudothelphusidae Pseudothelphusidae indet. species - troglophile - cave Family Trichodactylidae - Mexico: Veracruz; reported by Rodriguez & Hobbs Avotrichodactylus bidens (Bott) - troglophile - cave - (1989:398) Mexico: Tabasco Epilobocera armata Smith - trogloxene? - cave - Cuba: ?'Avotrichodactylus constrictus (Pearse) - troglophile - Las Villas cave - Mexico: Oaxaca Epilobocera gertraudae Pretzmann - trogloxene? - Rodriguezia mensabak Cottarelli & Argano - cave - cave - Cuba: Las Villas Mexico: Chiapas Epilobocera sinuatifrons (A. Milne-Edwards) - Trichodactylus bidens Bott - cave(s) - Mexico: Tabasco trogloxene? - cave - Puerto Rico 3. Isabellagordonia lonqipes Pretzmann - troglophile? - cave - Guatemala Odontothelphusa monodontis Rodriguez & Hobbs - Biogeographical Considerations: troglophile - cave - Mexico: Tabasco Phrygiopilus acanthophallus Smalley - trogloxene? - The discussions of nearly a century concerning the evo­ cave - Guatemala: Altaverapaz lution and biogeography of cave-dwelling decapods Potamocarcinus aspoekorum (Pretzmann) - troglophile in the Americas and Caribbean region are reviewed - cave - Belize in Hobbs et al. (1977). Some recent discoveries and Potamocarcinus leptomelus Rodriguez & Hobbs - hypotheses treating these crustaceans are summarized troglophile - cave - Mexico: Veracruz by Hart et al. (1985), Mannings al. (1986), Rodriguez Potamocarcinus maanus (Rathbun) - trogloxene - cave (1986), and Hobbs (1988). - Costa Rica Pseudothelphusa mexicana Al varez-Noguera - troglophile - cave - Mexico: Guerrero 102

Shrimps: species resulted in isolation of these small gene pools. The genus Typhlatya is one of the most succesful of The diverse "cave" shrimp faunas found in the region all troglobitic shrimps and the current distribution of covered in this discussion are polyphyletic in origin. cavernicolous species is widely disjunct. The subter­ Some are truly fresh water organisms while others are ranean species live in cave, anchialine, or crevicular capable of tolerating wide variations in salinity, such as habitats in Mexico, the Caribbean, western Atlantic, those species found in coastal anchialine and crevicular as well as on the Galapagos Island of Santa Cruz and habitats. Certain genera and species are known from Ascension Island. Manning et al. (1986) suggest that widely separated anchialine localities and pose puz­ the ancestors of extant Typhlatya may have frequented zling questions concerning their biogeography. Hart crevicular habitats along the Mid-Atlantic Ridge pri­ et al. (1985) and Manning et al. (1986) address some or to major plate tectonic activity and argued against of these questions and summarize similar distribution the derivation of today's species from possible pelagic patterns for other groups of crustaceans. They present larvae of ancestral forms. evidence to support the hypotheses that shrimps (and The family Agostocarididae is represented by a others) may have invaded crevicular and anchialine single genus and two disjunct species in the western habitats in Bermuda and the Caribbean before the open­ Atlantic and the western Caribbean Sea. Hart and Man­ ing of the North Atlantic in the Jurassic and thus these ning (1986) and Kensley (1988) point out similarities habitats may have served as Mesozoic refugia for the with the bresiliids and atyids but the phytogeny of this ancestors of many extant species. Also they suggested group is unclear. that subterranean environments could have provided a Palaemonid shrimps have invaded caves in the continuum of habitats from the abyssal zone to above United States, Mexico, Central America, and in sea level fresh waters and that these could have been many Caribbean islands. It is likely that the genera entered at virtually any level (depth) via marine crevic­ Cryphiops, Creaseria, Neopalaemon (all inhabiting ular habitats. caves and anchialine habitats in southern Mexico), and The procarid shrimps are represented by a single Troglocubanus (caves in Cuba, Jamaica, and Mexi­ genus and four species that are disjunctly distributed in co) were derived from various marine ancestors. The anchialine habitats in the western Atlantic Ocean, the species of Palaemonetes inhabiting fresh waters were western Caribbean Sea, and the eastern Pacific Ocean. derived from a widespread Mesozoic or early Ceno- This is one of the most primitive of all caridean shrimps zoic ancestor that gave rise to P. cummingi in the east and because of low variability among the species, Hart and to P. antrorum and P. holthuisi to the west. Mac- and Manning (1986) suggest that the genus demon­ robrachium is believed to have moved inland more strates an extremely slow rate of evolution. Procaris, recently and probably through competitive exclusion or its predecessors, may have been distributed widely expelled Palaemonetes from the middle latitudes by throughout the oceans and survives today in cryptic the Miocene. Five species of Macrobrachium have habitats where environmental rigors are minimal. invaded subterranean waters in this area of study, pri­ The Aty idae is a successful group that is widespread marily in Mexico (see Strenth 1976 and Hobbs et al. in caves in the Atlantic, the Bahamas and West Indies, 1977). in North and Middle America, and in the southeastern The family Alpheidae is represented by two gen­ Pacific Ocean. The genus Atya is found in caves only in era and two species that are cavernicolous in Mexico the West Indies and only A. brachyrhinus demonstrates (fresh water) and Bermuda (anchialine habitat). Auto­ any recognized troglomorphic adaptations; the two mate dolichognatha is widely distributed in the west­ additional species depict surface populations that have ern Atlantic and the Indo-West Pacific regions while extended their distributions into subterranean waters. Potamalpheops stygicola is known only from a single Xiphocaris is represented by a single species that is cave in Oaxaca, Mexico, and while sharing affinities epigean and occasionally enters caves in the Greater with members of the widespread genus Alpheopsis, its Antilles. The genus Palaemonias contains two disjunct congeners occur in West Africa. species that are troglobitic in the southeastern United Five genera and species constitute the hippolytid States. It is likely that they shared a common ancestor shrimps inhabiting caves and anchialine habitats in that was widely distributed in epigean aquatic systems. the area of study. All are of marine origin with Yage- Populations entered hypogean waters in different parts rocaris cozumel and Calliasmata rimolii having the of the range and subsequent extinction of the surface most restricted ranges (Cozumel Island and Domini- 103 can Republic, respectively) and Barbouria cuben- most part, allopatric and these crayfishes have retained sis demonstrating a markedly broader distribution primitive characteristics (plesiomorphy). (Bermuda, Bahamas, Caicos Islands, and West Indies). The cambaroid stock spread mainly to the north and Like B. cubensis, Janicea antiguensis and Somersiella east [e.g., ancestor of C. (P.) nerterius entered caves in sterreri exhibit disjunct distributions and are derived West Virginia] but also extended south and westward from widely ranging marine stocks. where different epigean ancestors entered subterranean waters in the Tennessee Valley, the Florida Panhandle Crayfishes: region, and in the Ozarks.

The most successful cambarid crayfishes in the North­ Crabs: ern Hemisphere are those belonging to the subfamily Cambarinae and they have undergone an adaptive radi­ The present distribution of freshwater crabs (Tri- ation that surpasses that of any other group chodactylidae, Pseudothelphusidae) can be summa­ (Hobbs 1988). Specifically, the cavernicolous cam- rized in terms of vicariance, dispersal, and "vari­ barines represent a polyphyletic group that has under­ ous dispersion events" (Rodriguez 1986). The wide gone multiple colonizations of subterranean waters by ranging, polyphyletic trichodactylids and other rela­ ancestral epigean species of the genera Cambarus, tives (originally from Africa) generally occupy large Orconectes, and Procambarus. rivers at low elevations, demonstrate plesiomorphy, Hobbs, Jr. (1988) and Hobbs III (1991) summa­ and have comparatively fewer adaptations to terrestrial rized the arguments concerning the origin and disper­ environments. The narrow ranging pseudothelphusids sal of cambarid decapods and an abbreviation of those and other groups (originally from West Africa) and discussion is presented here (see also Hobbs et al. India are usually found in small streams at higher alti­ 1977). The ancestral cambarine stock entered fresh tudes and show progressive adaptations to air breath­ waters of southeastern North America no later than ing. The Neotropical/Paleotropical disjunct distribu­ the Cretaceous or and perhaps earlier in the Mesozoic tion of crabs sharing synapomorphies of the buccal Era. Through adaptive radiation this ancestral Procam­ area and branchial chamber implies that pseudothel- barus stock underwent differentiation into the genus phusid freshwater crabs are a monophyletic group that Procambarus and two additional stocks, the orconec- was separated by a vicariant event (the break up of toid and cambaroid crayfishes. This highly variable Pangaea) during the mid-Cretaceous. The pseudothel- Procambarus stock extended its range primarily to the phusid morphocline in southern Mexico, based on the west and south and during the early Cenozoic became structure of the gonopod, follows a general westward widespread in the southern part of the United States direction and suggests a series of allopatric speciation and Mexico. During the Miocene the crayfish stock of primitive demes, encompasing a progressive immi­ from which the Mexican and Cuban troglobites arose gration along the westward geographical axis. Chance was isolated by a vicariance event. Probably compet­ dispersal after the Miocene played a role in the dis­ itive exclusion played a role in the resulting separa­ junct distribution of species in the Lesser Antilles and tion and slight overlap within the ranges of the pseu- Guianas. Independent invasion of the spelean environ­ dothelphusid crabs and the cambarid crayfishes (see ment by trichodactylid and pseudothelphusid ancestors Rodriguez 1986). of extant cavernicolous species occurred in Belize, The monotypic Troglocambarus probably shared a Costa Rica, Cuba, Guatemala, Mexico, and Puerto common ancestry with several members of the sub­ Rico. genus Ortmannicus (genus Procambarus), the ranges The two species of gecarcinids inhabiting caves in of which it overlaps (see Hobbs and Franz 1986). Cuba and Jamaica are trogloxenes and are far more fre­ The orconectoid stock moved primarily to the north quently found in burrows in epigean habitats. Ances­ and west, giving rise predominantly to stream-dwelling tors of both undoubtedly were marine forms that invad­ species of the genus Orconectes (other genera also). ed fresh waters. The ancestors of the extant troglobitic members of the Except for the troglobitic Sesarma (Sesarma) ver- genus found refuge from Pleistocene rigors in subter­ leyi the remaining grapsid cavernicolous crabs are ranean waters on the Cumberland Plateau, the Penny­ trogloxenes found in Cuba, Guatemala, and Jamaica royal Plateau, and on the Mitchell Plain. The current and all have marine ancestors that invaded freshwater ranges of these blind, depigmented forms are, for the environments. Although not known, it is likely that the 104 larva of the troglobitic S. (S.) verleyi may complete its Hobbs, H. H. III. 1991. Decapoda. In J. H. Thorp & A. P. Covich metamorphosis in fresh water. (eds), Ecology and Classification of North American Freshwater Invertebrates. Academic Press, New York: 823-858. Hoffman, A., J. G. Palacios-Vargas & J. B. Morales-Malacara, 1986. Manual de Bioespeleologia (Con nuevas aportaciones de Morelos References y Guerrero Mex.). Univ. Nac. Autonoma Mex., Mexico. Holsinger, J. R., 1964. The biology of Virginia Caves. In H. H. Chace, R. A. Jr. & R. B. Manning, 1972. Two new caridean shrimps, Douglas (ed.), Caves of Virginia. Falls Church (Va.): 57-74. one representing a new family, from marine pools on Ascension Holthuis, L. B., 1973. Caridean Shrimps found in land-locked salt­ Island (Crustacea: Decapoda: Natantia). Smithson. Contr. Zool. water pools at four Indo-West pacific localities (Sinai Peninsula, 131: 1-18. Funafuti Atoll, Maui and Hawaii Islands), with the description of Hart, C. W., Jr. & R. B. Manning, 1986. Two new shrimps (Pro- one new genus and four new species. Zool. Verhand. (Leiden): carididae and Agostocarididae, new family) from marine caves 128: 1-48. of the western north Atlantic. J. Crust. Biol. 6: 408^116. Kensley, B. 1988. New species and records of cave shrimps from Hart, C. W„ Jr. R. B. Manning & T. M. Iliffe, 1985. The fauna of the Yucatan peninsula (Decapoda: Agostocarididae and Hippoly- Atlantic marine caves: evidence of dispersal by sea floor spread­ tidae). J. Crust. Biol. 8: 688-699. ing while maintaining ties to deep waters. Proc. Biol. Soc. Wash. Manning, R. B„ C. W. Hart, Jr. & T. M. Iliffe, 1986. Mesozoic relicts 98: 288-292. in marine caves of Bermuda. Stygologia 2: 156-166. Hobbs, H. H., Jr., 1988. Crayfish distribution, adaptive radiation and Monod, T. & P. Cals, 1970. Sur une espece noubelle de crevette cav- evolution. In D. M. Holdich and R. R. Lowery (eds), Freshwater ernicole: Typhlatya alapagensis (Decapoda Natantia; Atyidae). Crayfish. Biology, Management and Exploitation. Croom Helm, Mis. Zool. Bel. lies Galapagos Ecuador 2: 57-103. London: 52-82. Rodriguez, G., 1986. Centers of radiation of freshwater crabs in Hobbs, H. H., Jr. & R. Franz, 1986. New troglobitic crayfish with the Neotropics. In R. H. Gore & K. L. Heck (eds), comments on its relationship to epigean and other hypogean cray­ Biogeography. Balkema, Rotterdam: 51-67. fishes of Florida. J. Crust. Biol. 6: 509-519. Rodriguez, G. & H. H. Hobbs, Jr., 1989. Freshwater crabs associated Hobbs, H. H., Jr. & R. Franz, (1992) Pwcambarus (Ortmanni- with caves in southern Mexico and Belize, with descriptions of cus) attiquus, a new toglobitic crayfish (Decapoda: Cambaridae) three new species (Crustacea: Decapoda). Proc. Biol. Soc. Wash. from the St. Johns River Basin, Florida. Proc. Biol. Soc. Wash, 102: 394-400. 105: 359-365. Strenth, N. E., 1976. A review of the systematics and zoogeography Hobbs, H. H, Jr., H. H. Hobbs HI & M. A. Daniel, 1977. A review of of the freshwater species of Palaemonetes Heller of North Amer­ the troglobitic decapod crustaceans of the Americas. Smithson. ica (Crustacea: Decapoda). Smithson. Contr. Zool. 228: 1-27. Contr. Zool. 244: 1—183.