sity of bats in the tropics. In addition to the insectivorousbats, which are also common in temperate regions,the tropics contain nectarivous, frugivorous,carnivorous, and sanguivorous species. Observationsin tropical caves have revealed that the guano of eachtype of bat is inhabited by its own distinctive faunalassemblage. Few detailed studies have been made ofthe fauna of caves in the New World tropics withrespect to their population size and the relationshipof this to bat guano. R. W. Mitchell (1970a) hasstudied the fauna inhabiting a passage in Cueva de laFlorida, Tamaulipas, which is inhabited by insectivorousbats. H. A. Mitchell (1965) studied the atmosphereof Cueva del Tigre, Sonora, a large cavecontaining several species of bats. In addition to thesignificance of bats to the study of the cave ecosystem,bats are also of importance to human health.Many species of cave-associated bat in Mexico havebeen found to carry rabies. Furthermore, many caveswhich are inhabited by bats harbor Histoplasmacapsulatum Darling, the causative agent of the dangerouslung disease, histoplasmosis. Constantine(1970) has published a detailed review of the significanceof bats to human health.Although there are numerous records of bats in<strong>Mexican</strong> caves, these records are few when the numberof known caves is considered. The difficulty ofobtaining collecting permits and of collecting andproperly preserving bats has left their study largelyto mammalogists working in the field. Many of thepapers published on bats include references to theiroccurrence in caves, but in many instances onlygeneral localities are given without reference towhether or not the bats were obtained from caves.The bat fauna of the caves of Guatemala and Belizeis even less well known, and there are very few caverecords <strong>for</strong> these two countries. A total of 72 speciesand subspecies of bat have been recorded fromthe caves of Mexico; only 20 species from Guatemalaand 8 species from Belize are known fromcaves.It is outside the scope of this review to do morethan briefly summarize the bat fauna associatedwith caves in Mexico, Guatemala, and Belize. Villa R.(1967) has published the most comprehensive reporton the bats of Mexico; this study includes manycave records. The only general review of the bats ofGuatemala is that of Jones (1966). No comprehensivestudy appears to have been published on the batfauna of Belize, but a few records were found in apaper on histoplasmosis in Belize (Quinones et al.,1978) and in general taxonomic studies. Records ofbats in the caves of various <strong>Mexican</strong> states and regionshave been included in the following surveys:Baja Cali<strong>for</strong>nia (Jones et al., 1965), Chihuahua (Anderson,1972), Coahuila (Baker, 1956), Durango(Baker and Greer, 1962), Guerrero (Lukens andDavis, 1957), Hidalgo (Carter and Jones, 1978),Jalisco (Watkins et al., 1972), \lichoacan (Hall andVilla R., 1949), Morelos (Davis and Russell, 1952),Nuevo Leon (Jimenez Guzman, 1968), Oaxaca(Goodwin, 1969), San Luis Potosi (Dalquest, 1953a),the Sierra de El Abra, San Luis Potosi and Tamaulipas,and the Sierra de Guatemala, Tamaulipas (:vIollhagen,1971), Sinaloa (Jones et al., 1972), Sonora(Burt, 1938; Cockrum and Bradshaw, 1963), Tamaulipas(Alvarez, 1963), Valley of Mexico (Villa R.,1952), Veracruz (Hall and Dalquest, 19(3), Yucatan(Pearse and Kellogg, 1938), Yucatan and QuintanaRoo (Hatt and Villa R., 1950; Birney et al., 1974),and the Yucatan Peninsula (Jones et al., 1973).Family EmbaUonuridaeFive species of emballonurid bat have been reportedfrom caves in Mexico and Gutemala. Thomas'ssac-winged bat, Balantiopteryx io Thomas, has beenreported from caves in Chiapas, Oaxaca, Tabasco,and Veracruz, Mexico; and Alta Verapaz and habal,Guatemala. This species seems to prefer to hang fromthe tops of pits and crevices, and may be presentwithin the twilight zone as well as in total darkness.Colonies as large as 1000 have been reported fromcaves (Hall and Dalquest, 1963). Peters' bat, H. plicataplicata Peters, is among the more frequentlycollected bats in the caves of southern Mexico. It isknown from caves from San Luis Potosi south intoGuatemala and from Colima east to Tabasco. Theymay inhabit both large, dark caves and smaller, welllitshelter-like openings. The greater doglike bat,Peropteryx kappleri kappleri Peters, is known fromcaves in Tabasco and Veracruz, Mexico; and Izabal,Guatemala. Most of the records are from small cavesand few bats were present. The lesser doglike bat,P. macrotis macrotis (Wagner), is known from cavesin Tabasco, Quintana Roo, Veracruz, and Yucatan,Mexico. This species has been found in both largecaves and small shallow cliff-face recesses. The greaterwhite-lined bat, Saccopteryx bilineata centralisThomas, is known from caves in Campeche, Jalisco,and Oaxaca, but this bat probably prefers to roost inhollow trees.Family MolossidaeSeven species of molossid bat have been recordedfrom caves in Mexico and one 8pecies in Guatemala.Of these, only species of the genus Tadarida are frequentlyencountered in caves. Tadarida aurispinosa252
(peale) is known only from a small cave at EI Salto,San Luis Potosi, and from Cueva del Ahra, Tamaulipas.Two subspecies of the Brazilian free-tailed bathave been found in <strong>Mexican</strong> caves, and one from acave in Guatemala. Tadarida brasiliensis intermediaShamel is known only from Grutas de Zapaluta, Chiapas,Mexico, and a cave near Coban, Alta Verapaz,Guatemala. The <strong>Mexican</strong> free-tailed bat, T. b. mexicana(Saussure), is known from caves throughoutmuch of Mexico. This insectivorous species is frequentlypresent in caves in colonies ranging into themillions. The invertebrate fauna associated with theenormous guano deposits of this species is a distinctiveassemblage of species (Mitchell, 1970b). Cockrum(1969) and Villa R. and Cockrum (1962) havestudied the migratory habits and patterns of thisspecies, while Constantine (1967) has studied its activitypatterns. The broad-tailed bat, T. laticaudataferruginea Goodwin, is known only from two cavesin Tamaulipas.Family MonnoopidaeSix species of mormoopid bat have been recordedfrom caves in Mexico, Guatemala, and Belize. Smith(1972) has revised the family, and his names are usedin the discussion here. Peters' ghost-faced bat (alsoknown as the old man bat), Mormoops megalophyllamegalophylla (Peters), has been found in caves fromTexas south into Guatemala. This species is insectivorousand may occur in colonies of up to severalthousand individuals. Mass die-offs, possibly as a resultof disease, have been recorded <strong>for</strong> this species inCueva del Diablo, Nuevo Leon (Villa R., 1956), andin Sotano de Sauz, Chihuahua (Sprouse, 1977). Theycommonly occur in caves with very high temperaturesand humidities. Davy's naked-backed bat, Pteronotusdavyi fulvus (Thomas), has been found in caves fromSonora and Nuevo Leon, Mexico, south into AltaVerapaz, Guatemala. This species is insectivorous.Parnell's mustached bat, P. parnellii (Gray), is representedin the caves of this region by two subspecies.Pteronotus parnellii mesoamericanus Smith has beentaken in caves in Chiapas, Tabasco, Veracruz, andYucatan, Mexico; the Maya Mountains of Belize; andEI Peten and Alta Verapaz, Guatemala. Pteronotus p.mexicanus (Miller) is known from caves from Tamaulipasand southern Chihuahua into Guerrero. This speciesis frequently taken with other bats, and one individualmay occur in a cluster of another species. Wagner'smustached bat, P. personatus psilotis (Dobson),is known from caves south from San Luis Potosi,Mexico, into Alta Verapaz, Guatemala. This insectivorousspecies is frequently found in large humidcaves in large numbers. The big naked-backed bat,Pteronotus suapurensis (]. A. Allen), is known onlyfrom Cueva Laguna Encantada, Veracruz, and Grutasde Lanquin, Alta Verapaz.Family NatalidaeThe family Natalidae is represented III <strong>Mexican</strong>caves only by the two subspecies of the <strong>Mexican</strong>funnel-eared bat, Natalus stramineus Gray, discussedbelow. Natalus stramineus mexicanus Miller is knownfrom caves in Baja Cali<strong>for</strong>nia Sur and Sonora. Nataluss. saturatus Dalquest and Hall occurs in caves fromNuevo Leon south into Guatemala. This insectivorousspecies may be present in colonies of several hundredindividuals.Family PhyUostomatidaeThe family Phyll08tomatidae is the largest andmost diverse family of bats occurring in the NewWorld tropics. Thirty-three species and subspecies ofthis family have been reported from the caves ofMexico, Guatemala, and Belize. The nomenclatureused belllw follows that of Jones and Carter (1976).Geoffroy's tailless bat, Anoura geoffroyi lasiopyga(Peters), has been found in caves in Chiapas, Guerrero,Mexico, Morelos, and Tamaulipas. Ten speciesand subspecies of the fruit-eating bats of the genusArtibeus have been recorded from caves in this region.Davis (1969, 1970a, 1970b) has reviewed thesystematics of the genus Artibeus in Middle America.Artibeus aztecus aztecus Andersen is known fromcaves in Guerrero, Mexico, San Luis Potosi, andTamaulipas; A. a. minor Davis occurs in caves in Chiapas.Artibeus hirsutus Anderson is known from cavesin Guerrero, Morelos, and Nayarit. The Jamaicanfruit-eating bat, Artibeus jamaicensis Leach, is representedin the caves of this region by four subspecies:A. j. paulus Davis from Chiapas; A. j. richardsoni J. A.Allen from Tabasco, Mexico, and Escuintla and AltaVerapaz, Guatemala; A. j. triomylus Handley fromGuerrero, Jalisco, Morelos, Michoacan, and Oaxaca;and A. j. yucatanicus Allen from Quintana Roo, SanLuis Potosi, Tamaulipas, Veracruz, and Yucatan.This large species may <strong>for</strong>m colonies containing thousandsof individuals. The areas under their roosts frequentlycontain thousands of sprouted seeds, and theinvertebrate fauna in these areas may be unlike thatin other parts of the cave. The big fruit-eating bat,A. lituratus intermedius J. A. Allen, is known fromcaves in Guerrero, Morelos, San Luis Potosi, andTamaulipas. Artibeus phaeotis phaeotis Miller hasbeen recorded from caves in Quintana Roo and Tabasco.Artibeus toltecus toltecus (Saussure) is knownfrom caves in Chiapas, Morelos, Nuevo Leon, andVeracruz, Mexico; and Alta Verapaz, Guatemala.253
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A REVIEW OF THECAVERNICOLE FAUNA OF
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. 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
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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
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Records.-Chiapas: Sumidero de Canad
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Family HeterozerconidaeSpecimens of
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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
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Records.-San Luis PotOSI: Sotano de
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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,
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appears to be most closely related
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this is almost certainly M. russell
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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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- Page 214 and 215: Cave. Paracophus caecus appears to
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- 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 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
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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
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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.