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the most poorly known species in the genus, and adiscussion of its affinities must await careful descriptionof the male genitalia. It has been taken in associationwith a possibly undescribed species of troglophilicParacophus.Tohila atelomma HubbellTohila atelomma Hubbell, 1938:191-192, 194-200,204, fig. 1-10, 77; Jeannel, 1943:156; Pearse,1945:175; Chopard, 1947:67; Nicholas, 1962:180; Vandel, 1964:217; Vandel, 1965a:181; Leroy,1967:665, 684, 685, 686, 709; Chopard,1968:262; Delamare-Deboutteville and Botosaneau,1970:70; Reddell, 1971b:55; Hubbell,1972:53; Reddell, 1977b:234, 239, 240, 258,261, 266, 286, 288.Tohila: Hubbell, 1938:191,201,202; Hubbell, 1972:54-56.Type-Iocality.-Chac Mol Cave (=Cueva Chac Mol),Tohil, Yucatan, Mexico.Distribution.-Known from caves in the state ofYucatan. See Fig. 69.Records.-Yucatan: Grutas de Balankanche, CuevaChac Mol, Cueva Luchil, Cueva Oxolodt, Cueva deSan Isidro, and Cueva del Cenote Xtolok.New records.-Yucatan: Cenote Aka Chen, CenoteCalchum, Cenote de Sambula (Motul), Grutas deTzab-Nah, and Actun Xpukil (det. T. H. Hubbell).GUATEMALAFig. 71.-Distribution of troglophilic gryllid crickets of the genera Amphiacusta and Arachnomimus: 1, Amphiacustaazteco; 2, Amphiaculta bolivari; 3, Amphiacusta maya; 4, Amphiacusta yucataTUJ; 5, Arachnomimus cavicola.210
Discussion.-The genus Tohila was originally assignedto the subfamily Pentacentrinae and consideredto be most closely related to the genus Trigonidomimusof the Old World (Hubbell, 1938). Hubbell(1972) has reexamined these genera and feelsnow that Tohila is most closely related to Paracophusand that they are probably both members of the subfamilyPhalangopsinae. Tohila is a monotypic genusand is small and pale with degenerate eyes. It is usuallyfound beneath rocks, but it is also occasionallytaken from moist flowstone or cave walls. It has beencollected from several caves which are also inhabitedby the troglophilic Amphiacusta yucatana. Numerouscollections of crickets from caves in the YucatanPeninsula await study, and some will certainly proveto contain this species.Family RhaphidophoridaeThe camel crickets of the family Rhaphidophoridaeare frequently found in the caves of Mexico andGuatemala. The 12 described species, however, representa small percentage of the fauna to be reportedeventually from caves. The described species belongto the genera Ceuthophilus, Anargyrtes, Leptargyrtes,Exochodrilus, Hypsobadistes, and Phoberopus. Severalother genera await description. The Rhaphidophoridaeappear to be absent from low elevations alongthe eastern flanks of the Sierra Madre Oriental southinto the Yucatan Peninsula. The caves in this regionare instead populated by species of the gryUid genusAmphiacusta. Amphiacusta ill this area inhabits theentrance zone usually occupied by rhaphidophoridsin other regions.Hubbell (1972) erected the genus Anargyrtes fortwo species from the states of Mexico and Guerrero.Anargyrtes annulata (Bilimek) was described fromGrutas de Cacahuamilpa, Guerrero; it is now knownalso from many caves in Guerrero and Mexico. Anargyrtesbolivari Hubbell is known primarily from epigeanlocalities, but it has been found in a lava cavenear San Geronimo, Mexico (see Fig. 72).The genus Ceuthophilus is the principal cavecricket genus inhabiting caves in the southwesternUnited States and northern Mexico. Only two species,Ceuthophilus (Geotettix) cunicularis Hubbell and C.(Ceuthophilus) variegatus Scudder, have been identifiedfrom Mexican caves. The first species is knownonly from Cueva de los Lagos, Coahuila, and the latterhas been collected in several caves in Coahuila andChihuahua (see Fig. 72). Many undescribed species ofCeuthophilus have been collected in caves in northernMexico.Hubbell (1972) described a new genus and twonew species for the camel crickets inhabiting thehigher elevations of the Sierra de Guatemala, Tamaulipas.Exochodrilus caelestis occurs in two caves atRancho del Cielo, and E. forcipatus is known fromcaves at higher elevations (see Fig. 72).The genus Leptargyrtes was erected by Hubbell(1972) to include two new species from Queretaro:L. boneti from caves near El Lobo in eastern Queretaro,and L. tejamanilae from caves to the west nearPinal de Amoles and Tejamanil (see Fig. 72).Hubbell (1977) described the Phoberopes groupto include several species of camel cricket from Guatemalaand Chiapas, Mexico. Phoberopus minor Hubbellis known from Cueva de Chemal n. 1 and Cuevade Santa Eulalia (=Cueva de los Resadores), Huehuetenango,Guatemala. Three species of long-legged,delicate crickets were placed in the new genus Hypsobadistes:H. gracilior Hubbell from two caves in Huehuetenango,Guatemala;H. stuarti Hubbell from Cuevadel Rayo de San Felipe, Chiapas; andH. tenuis Hubbellfrom three caves at San Agustin, Chiapas (see Fig. 72).Apparently undescribed species of the generaPristoceuthophilus and Argyrtes have been collectedfrom caves in Coahuila and Hidalgo respectively.Family StenopehnatidaeUnidentified species of the stenopelmatid generaAnabropsis, Glaphyrosoma, and Stenopelmatus havebeen collected from Mexican caves, but they are allprobably accidentals.Family TettigoniidaeA single unidentified tettigoniid of the genusDichopetala was collected below the entrance dropinto Cueva de Taninul n. 4, San Luis Potosi; it iscertainly an accidental.Order EmbiopteraAn undetermined genus and speci~ of the seldomcollected order Embioptera was found in Sotano deEI Triunfo, San Luis Potosi. It is presumably an accidental.Order PsocopteraThe order Psocoptera is occasionally taken fromcaves in Mexico and is doubtless far more abundantthan the few records would indicate. This group includestroglobites in other parts of the world, but allof the species known from Mexican caves are probablytroglophiles or trogloxenes.Family CaeciliidaeAn undetermined species of the genus Caeciliuswas collected in Sotano de El Triunfo, San Luis Potosi.Its ecological status is unknown.211
Page 1:
A REVIEW OF THECAVERNICOLE FAUNA OF
Page 8:
. 1-----,, ,", ,\'"I/I~" )/Fig. I.-
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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
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the gryllid cricket Paracophus suba
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has been Jiscussed by Segerstrom (1
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only apparent troglophile is the ps
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The cave fauna of this region is ri
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igneous rock immediately east of th
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Rio Soto la Marina and terminates a
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caves studied are many large stream
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unusual species is Neogovea mexasca
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pallidus Rowland, the nesticid spid
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troglobites: the trichoniscid isopo
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Cambridge, and Pholcophora bispinos
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of the genus Lutzomyia has not been
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The principal cavernous fonnation i
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ocks of Paleocene to Pliocene age.
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Beach and Supra-tidal Zone, the I\o
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The aquatic fauna of Mexico and Cen
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oaxacae Hobbs from the Valle Nacion
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described, the genus occurs in cave
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large genus with numerous epigean s
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milliped, laniatorid harvestmen, an
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Table 4.-Summary of cave inhabiting
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Discussion.-This large, troglobitic
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any way a part of the true cave fau
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they have been washed in with soil
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part of the aquatic fauna. A very l
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The genus Helicodiscus is a signifi
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associated with true cave waters. P
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Family DaphnidaeWilson (1936) repor
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tul), Cueva de San Isidro, Cueva de
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30 m below entrance of Cueva de la
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underflow of rivers, alluvial grave
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the similarities between the distri
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Yucatan but did not identify the sp
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Protrichoniscus: Reddell, 1967a:24.
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It indudes five spccies, of which t
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Type-locality.-Pozzo, Etla, Oaxaca,
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Records.-Campeche: Volc
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164; Pearse, 1945:167, 168, fig. 13
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Family AtyidaeThe family Atyidae is
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Creaseria morleyi (Creaser)Blind sh
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the pond which is fed by the cave s
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philes. Pseudothelphusa (Pseudothel
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Typhlochactas elliotti MitchellTyph
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Distribution.-Known only from the t
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(Dumitresco and Orghidan, 1977); th
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Leucohya magnifica MuchmoreUndescri
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Discussion.-This is the largest spe
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Type-Iocality.-S6tano de Huitzmolot
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Bartolo is a name applied to two ne
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Rowland, 1975b:19, 20; Dumitresco,
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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
Page 166 and 167: family. Clarence and Marie Goodnigh
Page 168 and 169: n. 2, San Luis Potosi; and K. rugos
Page 170 and 171: Type-locality.-Cueva de Apoala, San
Page 172 and 173: Records.-Chiapas: Sumidero de Canad
Page 174 and 175: Family HeterozerconidaeSpecimens of
Page 176 and 177: only one species. All known epigean
Page 178 and 179: nation must await study, but it is
Page 180 and 181: Mexico and Guatemala, and several u
Page 182 and 183: 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
Page 188 and 189: the genus Synoptura and removed the
Page 190 and 191: known from caves all contain troglo
Page 192 and 193: las Perlas, Cueva Chica de la Perra
Page 194 and 195: Discussion.-This species is distinc
Page 196 and 197: Speodesmus pecki ShearPolydesmidae,
Page 198 and 199: appears to be most closely related
Page 200 and 201: this is almost certainly M. russell
Page 202 and 203: Table 21.-Summary of cave inhabitin
Page 204 and 205: Type-Iocality.-Cueva Bonita del Pre
Page 206 and 207: Type-Iocality.-Cueva de El Jobo, El
Page 208 and 209: Order DipluraFamily CampodeidaeAlth
Page 210 and 211: of small soil inhabiting japygoids
Page 212 and 213: (Linnaeus) has been collected from
Page 214 and 215: Cave. Paracophus caecus appears to
Page 218 and 219: Family EpipsocidaeThe family Epipso
Page 220 and 221: citata Van Tyne in Cenote de Piste,
Page 222 and 223: een found in caves and mines in Yuc
Page 224 and 225: 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
Page 256 and 257: 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
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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.
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Reddell, J. R., and W. R. Elliott.
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Sanborn, C. C. 1936. Records and me
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Stone, B. 1979. The 1979 San Agusti
Page 292 and 293:
Villa R., B. 1956. Una extrana y se
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Wimsatt, W. A. 1969. Transient beha
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+*Cueva de los Chivos, 5 km E Altam
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Sumidero Yochib, Yochib, 8 km NE Te
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Serrania del Burro RegionCave, Haci
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+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
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Cueva de los Riscos, 2 km S Rancho
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Cueva de El Coy-see Cueva del Nacun
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+*Cueva del Aire, 1.8 km at S85°W
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*Sistema Purificacion, Conrado Cast
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*Cueva del Nacimiento del Rio Frio,
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Sotano de Humo, 5 km NW San Andres,
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and Bohor, 1975:25-27, 26m; Reddell
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*Actun Xpukil, 3 km S Calcehtok, Mp
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Appendix 2LIST OF TROGLOBITES BY ST
Page 327 and 328:
Paravachonium superbum-I'seudoscorp
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SUPPLEMENTWhile this manuscript was
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Cambala speobia (Chamberlin) (p. 19
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mida (Arachnida) in the New World.
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