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oaxacae Hobbs from the Valle Nacional region ofOaxaca. and P. (A.). oaxacae reddelli Hobbs fromcaves in the Atoyae region ofV~racruz and the AcatIan and Valle I\acional regions of Oax aca. These speciesare both closely related to species now inhabitingthe same region.CraLs of the families Pseudothelphusidae andlrichodactyJidae are represented in the caves of;\lhico and Guatemala. Three pseudothelphusid crabsof UIC cave-restricted genus 1'yphlopseudothelphusahave been described, one from Chiapas, '\Iexico, andthe remaining two from Alta Verapaz, Guatemala.1~vphlopseudothelphusais closely related to the epigeangenus Patamacarcinus. The family Trichodactylidaeis represented only by Trichodactylus (Rodriguezia)mensabak Cottarelli and Argano from Cuevade ;\Iicolas Bravo, Chiapas. Its closest relative is anepigean species also known from Chiapas.The fish fauna of '\
Temperate relicts.-A few species of troglobite areknown from Mexican caves which belong to groupsunknown from the surface in this region. These includethe centipede Nuevobius cavieolens Chamberlin,millipeds of the families Cambalidae and Trichopetalidae,and collembolans of the genera Acherontidesand Oneopodura. In our present state of knowledgeof the endogean fauna of tropical and subtropicalregions, we can only speculate that thesegroups will not be found on the surface. It wouldappear, however, that they represent true relicts of afauna once widespread throughout Mexico but nowto be found only in caves.The centipede Nuevobius cavieolens Chamberlinis known only from Cueva de la Boca in the Sierra dela Silla, Nuevo Leon. The only other species in theI!tnus is a trog/obite from Tennessee (U.S.A.).The rnilliped families Cambalidae and Trichopetalidaeare unknown on the surface in Mexico orCentral America. It is likely that these species aresurvivor;; of a once-widespread fauna inhabiting thetemperate forests that once covered parts of thisregion. These forests are now represented only byrelict poekeb at higher elevations of the ChiapasGuatemalan Highlands, Sierra Madre del Sur, andSierra :\1adre Oriental.The family Camhalidae is an important element ofthe cave and endogean fauna of the United States.One aberrant species, larmilka alba Shear, is knownonly from a cave in Belize. The genus lvlexicambalaincludes described species from caves in Oaxaca, SanLuis Potosi, and Tamaulipas. It is probably a relictof a group once widespread in Mexico hut that isnow restricted to caves. Finally, Cambala speobia(Chamberlin) has been collected in Cueva de losLagos, Coahuila. This is a common species in thecaves of Central Texas immediately north of the RioGrande.The family Trichopetalidae is a significant group inthe cave and endogean habitats of the southeasternUnited States. In Mexico it is represented only by thegenus Mexiterpes with four troglobitic species fromcaves in San Luis Potosi and Queretaro. The mosthighly cave-adapted species (M. sabinus Causey)occurs at comparatively low elevations in the Sierrade El Abra. The remaining species, which occur atmuch higher elevations, retain ocelli and are apparentlymore recent troglobites. It would appear thatthese were inhabitants of a temperate forest whichbecame extinct earlier at lower elevations than athigher ones.Two genera of collembolans appear to be temperaterelicts in the caves of Mexico. The hypogastruridgenus Acherontides is known only from two Mexicancaves, A. atoyaeense Bonet in Veracruz and A. potosinusBonet in the Xilitia region of San Luis Potosi.Other species of the genus occur in Rumania, Japan,and Afghanistan. The oncopodurid genus Oncopoduraincludes O. atoyaeense Bonet from Veracruz andO. prietoi Bonet from Nuevo Leon and possiblyTamaulipas. Other species of the genus occur inEurope, Japan, and the United States.There are several other groups which are unknownfrom the epigean habitat in Mexico but include troglobites.Among the more important of these are sixdescribed and several undescribed millipeds of thefamily Trichopolydesmidae, agelenid spiders of thegenus Cieurina, and ptinid beetles of the genus Niptus.It is likely, however, that as the endogean faunaof Mexico becomes better known these groups willalso be found in surface habitats.The family Trichopolydesmidae is known from theepigeum in Mexico only by two species in BajaCalifornia. All of the remaining species are troglobites.These are divided among three genera, Carambawith three species in Chiapas, Speodesmus with onespecies in Tamaulipas, and Tylogoneus with twospecies in San Luis Potosi and Tamaulipas. Each ofthese species is known only from the type-locality.The spider genus Cicurina is a major element ofthe cave and epigean fauna of Texas and other par.tsof the United States. In Mexico three troglobites andone troglophile have been described. The troglobitesare known from caves in Yucatan, Tamaulipas, andCoahuila, and the troglophile from caves in Tamaulipas.With more collections from high elevations inMexieo, this genus will almost certainly be found inendogean habitats.Two species of beetle of the family Ptinidae areknown from ;\1exican caves: Niptus abseonditusSpilman, a possible troglobite from Hidalgo, and N.abstrusus Spilman, a troglophile from caves in Texas(U.S.A.) and Coahuila and Durango, Mexico. Neitherspecies has been collected from the surface, but it islikely that N. abstrusus or other species of the genuswill be found there with more collecting.Species with temperate affinities.-Several groupsof cavernicole are members of genera containing bothepigean and cavernicole species, but which havecloser affinities to temperate species than to speciesinhabiting Mexico. These include trichoniscid isopodsof the genus Braekenridgia, scorpions of the genusTyphloehaetas, pseudoscorpions of the genus Aphrastochthonius,and spiders of the genera Tegenaria,Leptoneta, and Nestieus.The isopods of the genus Brackenridgia are amongthe most commonly collected animals in the caves ofMexico. Although only five species have thus far been63
Page 1:
A REVIEW OF THECAVERNICOLE FAUNA OF
Page 8:
. 1-----,, ,", ,\'"I/I~" )/Fig. I.-
Page 12 and 13:
collected a troglobitic cirolanid i
Page 14 and 15:
C. Bolivar and J. Carranza investig
Page 16 and 17:
springs in the vicinity of Cautro C
Page 18 and 19: Cueva del Diablo near Orizaba, Vera
Page 20 and 21: Those species of troglobite which h
Page 22 and 23: Table 2.-(continued)1970. Eodrilus
Page 25 and 26: CAVE REGIONSOnly the cave regions o
Page 27 and 28: Table 3.-(continued)TroglobitesTrog
Page 29 and 30: y the waters of the Amistad Reservo
Page 31 and 32: Although a few caves in the Norther
Page 33 and 34: 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 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 86 and 87: The genus Helicodiscus is a signifi
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
Page 139 and 140:
Distribution.-Known only from caves
Page 141 and 142:
een collected. It is frequently fou
Page 143 and 144:
Distribution.-Known from two caves
Page 146 and 147:
actually a natural cave artificiall
Page 148 and 149:
Discussion.-This large species has
Page 150 and 151:
and T. patellaris Bryant from caves
Page 152 and 153:
estricted to the Mexican Plateau (s
Page 154 and 155:
maya Chamberlin and Ivie from Campe
Page 156 and 157:
Metagonia chiquita GertschMetagonia
Page 158 and 159:
Metagonia capilla: Mitchell and Kaw
Page 160 and 161:
the cave and in general association
Page 162 and 163:
DiscWlSion.-This is a delicate, eye
Page 164 and 165:
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 216 and 217:
the most poorly known species in th
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
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
Page 304 and 305:
*Gruta Sur de San Bartolo, 9 km S S
Page 306 and 307:
*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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