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The genus Helicodiscus is a significant element of theTexas cave fauna.Family OrthalicidaeAlthough seven species of the family Orthalicidaehave been reported from caves, the only one which isknown with certainty to live in caves is Bulimulusunicolor (Sowerby). This species was reported fromseveral caves in Yucatan (Bequaert and Clench, 1936,1938); it has been collected from several additionalcaves recently.Family SpiraxidaeThis is the most important family of snails withrespect to the cave ecosystem, with species of Euglandina,Spiraxis, and Streptostyla having been found astroglophiles. Euglandina cylindracea (Phillips), Streptostylameridana meridana (MoreIet), and Streptostylaventricosula (Morelet) are frequently collectedin the caves of the Yucatan Peninsula. Streptostylajilitlana Dall is an important species in the high altitudecaves of Queretaro and San Luis PotosI. Eggs,newly hatched young, and adults of Streptostylabartschi (Dall) have been found in the inner rooms ofCueva de la Mina, Tamaulipas (Reddell and Mitchell,1971b). This species is also known from other cavesin San Luis PotosI and Tamaulipas.Family UrocoptidaeBrachypodella dubia Pilsbry and Microceramusconcisus (Morelet) are frequently encountered speciesin the caves of the Yucatan Peninsula. Thompson(1967) reported Brachypodella speluncae (Pfeiffer)from two caves in El Peten, Guatemala. A new speciesof the genus Coelocentrum is known from caves inTamaulipas.Family XanthonychidaeThe only species of this family known from Mexicancaves is an undescribed species of Xanthonyxfrom Cueva de la Capilla, Tamaulipas. It was foundon the wall of the cave near a small upper entrance.Family ZonitidaeSeveral species of zonitid snail are known fromMexican caves. The only one which is common isHawaiia minuscula (Binney), known from caves inCampeche, Veracruz, and Yucatan.Class PelecypodaFew clams have ever been recorded from caves andthen usually from the vicinity of spring entrances. Asmall undetermined species is known from Cueva delNacimiento del Rio San Antonio, Oaxaca. This speciesinhabits the stream passage in large numbers andis pigmentless with a· thin shell allowing the internalorgans to be observed. The determination of itsecological status must await further study.Phylum ArthropodaClass CrustaceaThe class Crustacea is second only to the Arachnidain the number and diversity of troglobites presentin Mexican and Guatemalan caves. Of the eightorders of subterranean crustaceans represented, sixcontain troglobites (see Table 10). Two additionalorders, the Thermosbaenacea and the Bathynellacea,are present in the caves of Texas and other parts ofthe world but are not yet known from Mexico orCentral America. This lack is probably a reflectionon collecting techniques rather than an indication ofthe absence of these groups. In general the planktonTable 10.-Summary of cave inhabiting Crustacea.TroglobitesOther SpeciesBranchiuraArguloidaArgulidae 0 2CopepodaCalanoidaDiaptomidae 1 3CyclopoidaCyclopidae 0 10HarpacticoidaAmeiridae 0 5Canthocamptidae 0 2OstracodaPodocopaCypridae 0 12Darwinulidae 0 1Entocytheridae 2 10BranchipodaCladoceraChydoridae 0 1Daphnidae 0 7Malacostraca*Isopoda 40 16AmphipodaBogidiellidae 9 0Hadziidae 5 0Hyalellidae 0 1Melitidae 0 1MysidaceaLepidomysidae 2 0Mysidae 2 0*Decapoda 17 18Total 78 89*For summary by families see Tables 11-12.80
of Mexican caves is unknown, but recent collectionsin southern Mexico by Dr. Valerio Sbordoni and hiscolleagues have included emphasis on planktonicforms.Subclass BranchiuraOrder ArguloidaFamily ArgulidaeTwo species of argulid have been taken in associationwith fish in the cenotes of Yucatan (Wilson,1(36). Argulus chromidis Kroyer was found in theintestine of Rhamdia guatemalensis depressa Barbourand Cole in Cenote Chica de XanaLa; A. rhamdiaeWilson was found on the skin of R. guatemalensisdepressa in Cenote Scan Yui.Subclass CopepodaCopepods are an important element III the cavefauna of Mexico but remain poorly known. Only onespecies is definitely troglobitic, but others may proveto be so once they are more carefully studied. A totalof 20 species of non-troglobitic copepod arl' nowknown from the underground waters of lVlexico (seeTable 10).Order CalanoidaFamily DiaptomidaeTwo diaptomids have been identified from caves inMexico, one of which is a trogloLite and is discussedbelow. Diaptomus texensis M. S. Wilson has been collectedfrom a small drip pool in Actun Loltun, Yucatan.Diaptomus (Microdiaptomus) cokeri Osorio TafallDiaptomus nueva especie y subgenera: Anonymous,1942a:221.Diaptomus (Microdiaptomus) cokeri Osorio Tafall,1942:206, 207-210, fig. 1-17; Osorio Tafall, 1943:49, 50, 51, 52,53,68,69; Rioja, 1953a:287, 288,292; Wilson, 1959:780; Nicholas, 1962:168; Vandel,1964:131; Vandel, 1965a:l07; McKenzie,1965a:38; Reddell, 1971b: 17; Reddell and I\Iitchell,1971a:141; Reddell and 8lliott, 1973a:I71.Microdiaptomus: Osorio Tafall, 1946: 153.Diaptomus cokeri: Bolivar, 1950:216; Tamayo,1962:vol. III:260.Diaptomus (Microdiaptomus) cokersi: Barrera, 1968:311 (erroneous spelling).Microdiaptomus cokeri: Thines and Tercafs, 1972:66.Type-Iocality.-Cueva Chica, Ciudad Valles, SanLuis Potosi, Mexico.Distribution.-Known only from two caves nearValles, San Luis Potosi.Records.-San Luis PotOSI: Cueva Chica and Cuevade Los Sabinos.Discussion.-This species of fresh-water origin has,to my knowledge, not been collected since it was firstreported. The subgenus Microdiaptomus includesonly this species.Order CyclopoidaFamily Cyclopidael\ine species of cyclopid copepods have been identifiedfrom Mexican caves. Osorio Tafall (1943) rcportedthe presence of Cyclops (AcanthocycLops) vernalisrobustus Sars and MesocycLops (ThermocycLops)inversus (Kiefer) from caves near Valles, San LuisPotosi. EucycLops serruLatus (Fischer) and Tropocyclopsprasinus (Fischer) are known from caves inSan Luis Potosi (Osorio Tafall, 1(43) and ct~nolt'sand caves in Yucatan (Wilson, 1936; Pearse and Wilson,1(38). MacrocycLops albidus (Jurine) is a frequentinhabitant of caves, and in Mexico it is knownfrom caves in Yucatan (Wilson, 1936; Pearse and Wilson,1938), San Luis Potosi (Osorio Tafall, 1943),Campeche, and Guerrero. A probable new sp(~cies ofMesocycLops related to M. brazilianus Kiefer has beencollected in Grutas del 10gote, Guerrero. Yeatman(1977) redescribed .f\Ilesocyciops ellipticus Kieferfrom Grutas de XtaclImhilxunam, Campeche. Thisspecies, also known from caves in Oaxaca and Yucatan,is probably that reported as iH. leuckarti (Claus)from caves and cenotes in Yucatan (Wilson, 1936;Pearse and Wilson, 1938). Mesocyclops (Thermocyclops)tenuis (Marsh) is known from caves andcenotes in Yucatan (Wilson, 1936; Pearse and Wilson,1938) and from caves in Campechc. Paracyclops Jimbriatus(Fischer) is one of the mort~ frequently encounteredcopepods in North American caves; inMexico it has been found in caves in the states ofSan Luis Potosi (Osorio TafaH, 1(43), Yucatan, andCampeche.Order HarpacticoidaFamily AmeiridaeNitocra sp. was reported by Osorio TafaU (1943)from Cueva Chica and Cueva de Los Sahinos, SanLuis Potosi. Wilson (1936) identified four species ofNitocra from caves and cenotes in YlICatan, but onlyN. pusilla Sars from Cenote de Samhula (Motul) is81
Page 1:
A REVIEW OF THECAVERNICOLE FAUNA OF
Page 8:
. 1-----,, ,", ,\'"I/I~" )/Fig. I.-
Page 12 and 13:
collected a troglobitic cirolanid i
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C. Bolivar and J. Carranza investig
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springs in the vicinity of Cautro C
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Cueva del Diablo near Orizaba, Vera
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Those species of troglobite which h
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Table 2.-(continued)1970. Eodrilus
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CAVE REGIONSOnly the cave regions o
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Table 3.-(continued)TroglobitesTrog
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y the waters of the Amistad Reservo
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Although a few caves in the Norther
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an elevation of about 900 meters. T
Page 35 and 36: the gryllid cricket Paracophus suba
Page 37 and 38: has been Jiscussed by Segerstrom (1
Page 39 and 40: only apparent troglophile is the ps
Page 41 and 42: The cave fauna of this region is ri
Page 43 and 44: igneous rock immediately east of th
Page 45 and 46: Rio Soto la Marina and terminates a
Page 47 and 48: caves studied are many large stream
Page 49 and 50: unusual species is Neogovea mexasca
Page 52 and 53: pallidus Rowland, the nesticid spid
Page 54 and 55: troglobites: the trichoniscid isopo
Page 56 and 57: Cambridge, and Pholcophora bispinos
Page 58 and 59: of the genus Lutzomyia has not been
Page 60 and 61: The principal cavernous fonnation i
Page 62 and 63: ocks of Paleocene to Pliocene age.
Page 64 and 65: Beach and Supra-tidal Zone, the I\o
Page 66 and 67: The aquatic fauna of Mexico and Cen
Page 68 and 69: oaxacae Hobbs from the Valle Nacion
Page 70 and 71: described, the genus occurs in cave
Page 72 and 73: large genus with numerous epigean s
Page 74 and 75: milliped, laniatorid harvestmen, an
Page 76 and 77: Table 4.-Summary of cave inhabiting
Page 78 and 79: Discussion.-This large, troglobitic
Page 80 and 81: any way a part of the true cave fau
Page 82 and 83: they have been washed in with soil
Page 84 and 85: part of the aquatic fauna. A very l
Page 88 and 89: associated with true cave waters. P
Page 90 and 91: Family DaphnidaeWilson (1936) repor
Page 92 and 93: tul), Cueva de San Isidro, Cueva de
Page 94 and 95: 30 m below entrance of Cueva de la
Page 96 and 97: underflow of rivers, alluvial grave
Page 98 and 99: the similarities between the distri
Page 100 and 101: Yucatan but did not identify the sp
Page 102 and 103: Protrichoniscus: Reddell, 1967a:24.
Page 104 and 105: It indudes five spccies, of which t
Page 106 and 107: Type-locality.-Pozzo, Etla, Oaxaca,
Page 108 and 109: Records.-Campeche: Volc
Page 110 and 111: 164; Pearse, 1945:167, 168, fig. 13
Page 112 and 113: Family AtyidaeThe family Atyidae is
Page 115 and 116: Creaseria morleyi (Creaser)Blind sh
Page 117 and 118: the pond which is fed by the cave s
Page 119 and 120: philes. Pseudothelphusa (Pseudothel
Page 121 and 122: Typhlochactas elliotti MitchellTyph
Page 123 and 124: Distribution.-Known only from the t
Page 125 and 126: (Dumitresco and Orghidan, 1977); th
Page 127 and 128: Leucohya magnifica MuchmoreUndescri
Page 129 and 130: Discussion.-This is the largest spe
Page 131 and 132: Type-Iocality.-S6tano de Huitzmolot
Page 133 and 134: Bartolo is a name applied to two ne
Page 135 and 136: Rowland, 1975b:19, 20; Dumitresco,
Page 137 and 138:
Type-Iocality.-Actlin Kaua, Yucatan
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Distribution.-Known only from caves
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een collected. It is frequently fou
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Distribution.-Known from two caves
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actually a natural cave artificiall
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Discussion.-This large species has
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and T. patellaris Bryant from caves
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estricted to the Mexican Plateau (s
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maya Chamberlin and Ivie from Campe
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Metagonia chiquita GertschMetagonia
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Metagonia capilla: Mitchell and Kaw
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the cave and in general association
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DiscWlSion.-This is a delicate, eye
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Type-Iocality.-Cueva de Los Sabinos
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family. Clarence and Marie Goodnigh
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n. 2, San Luis Potosi; and K. rugos
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Type-locality.-Cueva de Apoala, San
Page 172 and 173:
Records.-Chiapas: Sumidero de Canad
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Family HeterozerconidaeSpecimens of
Page 176 and 177:
only one species. All known epigean
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nation must await study, but it is
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Mexico and Guatemala, and several u
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Type-locality.-Gruta de Atoyac, Ver
Page 184 and 185:
Records.-San Luis PotOSI: Sotano de
Page 186 and 187:
Records.-Veracruz: Grutas de Atoyac
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the genus Synoptura and removed the
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known from caves all contain troglo
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las Perlas, Cueva Chica de la Perra
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Discussion.-This species is distinc
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Speodesmus pecki ShearPolydesmidae,
Page 198 and 199:
appears to be most closely related
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this is almost certainly M. russell
Page 202 and 203:
Table 21.-Summary of cave inhabitin
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Type-Iocality.-Cueva Bonita del Pre
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Type-Iocality.-Cueva de El Jobo, El
Page 208 and 209:
Order DipluraFamily CampodeidaeAlth
Page 210 and 211:
of small soil inhabiting japygoids
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(Linnaeus) has been collected from
Page 214 and 215:
Cave. Paracophus caecus appears to
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the most poorly known species in th
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Family EpipsocidaeThe family Epipso
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citata Van Tyne in Cenote de Piste,
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een found in caves and mines in Yuc
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ground dwelling habit, many beetles
Page 226 and 227:
Type-Iocality.-Sotano de Tejamanil,
Page 228 and 229:
Vigna Taglianti, 1972:118, 121-127,
Page 230 and 231:
Veracruz) belongs in another subgen
Page 232 and 233:
Troglobacanius sbordonii VomeroTrog
Page 234 and 235:
Ptomaphagus (Adelops) is a group of
Page 236 and 237:
(A.) speLaeus, from Grutas de Cacah
Page 238 and 239:
epresented by many species. Tenebri
Page 240 and 241:
Myodopsylla gentilis Jordan and Rot
Page 242 and 243:
genus Pholeomyia have been collecte
Page 244 and 245:
Wheeler (1938) has reported the pre
Page 246 and 247:
263.264, 265, 267, 270, 273, fig. 5
Page 248 and 249:
Anophthyctys jordani: Stefanelli, 1
Page 250 and 251:
Prietella phreatophila Carranza, 19
Page 252 and 253:
TyphlilJs: Pearse, 1938a:13, 15; Hu
Page 254 and 255:
Family BufonidaeFour species of toa
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Oaxaca; and E. yucatanensis Lynch f
Page 258 and 259:
sity of bats in the tropics. In add
Page 260 and 261:
Three species of short-tailed bats
Page 262 and 263:
abundant in the entrance area of Cu
Page 264 and 265:
Andrews, E. W., IV. 1970. Balankanc
Page 266 and 267:
Bolivar y Pieltain, C. 1940. Explor
Page 268 and 269:
Brignoli, P. M. 1979a. The morpholo
Page 270 and 271:
Conde, B. 1975. Description du prem
Page 272 and 273:
Ediger, G. 1970. The Birthday Passa
Page 274 and 275:
Gonzalez Ochoa, A. 1963b. Relacione
Page 276 and 277:
Hobbs, H. H.,Jr., and T. C. Barr, J
Page 278 and 279:
Jordan, c. B. 1937. Bringing in the
Page 280 and 281:
McKenzie, D. 1965a. Caves of the Si
Page 282 and 283:
Monod, T. 1975. Sur la distribution
Page 284 and 285:
Peck, S. B., and J. J. Lewis. 1977.
Page 286 and 287:
Reddell, J. R., and W. R. Elliott.
Page 288 and 289:
Sanborn, C. C. 1936. Records and me
Page 290 and 291:
Stone, B. 1979. The 1979 San Agusti
Page 292 and 293:
Villa R., B. 1956. Una extrana y se
Page 294 and 295:
Wimsatt, W. A. 1969. Transient beha
Page 296 and 297:
+*Cueva de los Chivos, 5 km E Altam
Page 298 and 299:
Sumidero Yochib, Yochib, 8 km NE Te
Page 300 and 301:
Serrania del Burro RegionCave, Haci
Page 302 and 303:
+Cueva de Tezoapa, 1.5 km E Acahuiz
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*Gruta Sur de San Bartolo, 9 km S S
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*Cueva del Puente de Fierro, 1.5 km
Page 308:
Cueva de los Riscos, 2 km S Rancho
Page 311 and 312:
Cueva de El Coy-see Cueva del Nacun
Page 313 and 314:
+*Cueva del Aire, 1.8 km at S85°W
Page 315 and 316:
*Sistema Purificacion, Conrado Cast
Page 317 and 318:
*Cueva del Nacimiento del Rio Frio,
Page 319 and 320:
Sotano de Humo, 5 km NW San Andres,
Page 321 and 322:
and Bohor, 1975:25-27, 26m; Reddell
Page 323 and 324:
*Actun Xpukil, 3 km S Calcehtok, Mp
Page 325 and 326:
Appendix 2LIST OF TROGLOBITES BY ST
Page 327 and 328:
Paravachonium superbum-I'seudoscorp
Page 329 and 330:
SUPPLEMENTWhile this manuscript was
Page 331 and 332:
Cambala speobia (Chamberlin) (p. 19
Page 333:
mida (Arachnida) in the New World.
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