Three species of short-tailed bats of the genusCarollin have been recorded from the caves of thisregion. The nomenclature <strong>for</strong> this genus follows therevision of Pine (1972). Carollin brevicauda (Schinz)is known from caves in Chiapas, Puebla, QuintanaRoo, San Luis Potosi, Tabasco, Tamaulipas, andVeracruz, Mexico; and the Maya Mountains of Belize.Carollin perspicillata azteca Saussure has been foundin caves in Chiapas, Veracruz, and Yucatan, l\lexico;the Maya Mountains of Belize; and El Peten and AltaVerapaz, Guatemala. Carollia subrufa (Hahn) isknown only from caves in Tabasco, Mexico; and AltaVerapaz, Guatemala.The nectar-feeding <strong>Mexican</strong> long-tongued bat,Choeronycteris mexicana Tschudi, has been found incaves in Mexico from Coahuila and Sonora south toMichoacan. Peters' false vampire bat, Chrotopterusauritus auritus (Peters), has been found in caves inChiapas, Oaxaca, Veracruz, Yucatan, and QuintanaRoo. This is apparently a carnivorous species.Three species of vampire bat occur in Mexico andCentral America, and all are known from caves.Villa R. (1953a) has discussed the systematics of thesubfamily Desmodontinae. Diaemus youngi (Jentink)is a rare species known only from caves by specimenscollected in Grutas del Cocona, Tabasco. The hairyleggedvampire bat, Diphylla ecaudata centralisThomas, has been reported from caves in Chiapas,Oaxaca, Quintana Roo, San Luis Potosi, Tamaulipas,Veracruz, and Yucatan. This species is reported toleave only dry, brown stains beneath its roosts (Halland Dalquest, 1963). The most common vampire bat,and probably the most studied of all <strong>Mexican</strong> bats, isDesmodus rotundus murinus Wagner. Various aspectsof the behavior of a colony of this species inhabitingCueva de Don Luis, Tabasco, has been studied byWimsatt (1969). Intensive investigations of this specieswere conducted in southern Mexico by the UnitedStates Bureau of Sport Fisheries and Wildlife(1970, 1971). This species is known from caves fromNuevo Leon into Belize and Guatemala. It usuallyroosts in small domes or crevices in the ceiling of thecave in clusters ranging from a few individuals tomore than 100. The presence of this species in a cavecan be immediately determined by pools of black,tarry guano beneath their roosts. These pools harbora distinctive invertebrate fauna, including severalfamilies of fly, and histerid and leiodid beetles.Pallas' long-tongued bat, Glossophaga soricinaleachii (Gray), is among the more frequently encounteredbats in the caves of Mexico, Guatemala,and Belize. This species feeds on nectar and fruit,and roosts deep in caves. It is known from caves fromTamaulipas and Durango south into Guatemala andBelize.Underwood's long-tongued bat, Hylonycteris underwoodiThomas, is a rare species known from cavesin Tabasco and Veracruz. In one cave pits of joboplums were found beneath their roost, indicating theyhad taken the fruit into the caves to eat (Hall andDalquest, 1963). Another species which is seldomcollected from caves is Tornes' long-eared bat, Lonchorhinaaurita aurita Tomes, known only from cavesin Oaxaca, Quintana Roo, and Tabasco. In QuintanaRoo this species was found to roost in clusters ofabout 10 individuals in depressions in the ceiling(Jones et aI., 1973).Two species of long-nosed bat of the genus Leptanycterisare known from caves in Mexico. This genusis known from caves from Texas south into Guerrero.The ranges of the two species are broadly sympatricthroughout most, if not all, of Mexico, and there hasbeen much confusion as to their correct identity.Hoffmeister (1957) and Ramirez-Pulido and Alvarez(1972) have discussed the taxonomy of the genus.Leptonycteris nivalis (Saussure) is known from cavesfrom Tamaulipas into Guerrero. Leptonycteris sanbomiHoffmeister has been taken in caves from Sonoraand Chihuahua south into Chiapas and Guerrero.Both species have been taken from the same caves onseveral occasions. These species are nectar-feeders andmay occur in colonies containing several hundred individuals.The only cave record <strong>for</strong> the long-legged bat, ,Wacrophyllummacrophyllum Schinz, is that of a cavenear Teapa, Tabasco. This is the northern limit of therange <strong>for</strong> this species.Two species of the leaf-nosed bats of the genusMacrotus are known from caves in Mexico. The systematicsof these insectivorous bats have been studiedby Anderson and Nelson (1965). Macrotus califomicusBaird is known from caves in Baja Cali<strong>for</strong>nia Surand Sonora. Macrotus waterhousii Gray is representedin the caves of Mexico by two subspecies: U. w. bulleriH. Allen from Hidalgo, Jalisco, and Nuevo Leon;and M. w. mexicanus Saussure from Colima, Guerrero,Mexico, and Morelos.Three species of small-eared bats of the genusMicronycteris have been recorded from caves inMexico and Belize. Micronycteris megalotis mexicanaMiller has been found in caves in Guerrero, Morelos,San Luis Potosi, Tabasco, Tamaulipas, and Yucatan.This is apparently an insectivorous species and isusually present in small numbers. Micronycteris sylvestris(Thomas) has been recorded only from cavesin Jalisco and Veracruz. Micronycteris brachyotis(Dobson) has been recorded from caves only fromthe Maya fountains of Belize.254
The spear-nosed bat, Mimon cozumelae Goldman,has been taken from caves in Oaxaca, Tabasco, Veracruz,and Yucatan. These large bats have been reportedto feed on very ripe fruit or else on insectsfeeding on the fruit (Hall and Dalquest, 1963). Theyusually occur in caves only in small numbers.The yellow-shouldered bat, Stumira lilium parvidensGoldman, has been recorded from caves inPuebla, Quintana Roo, and Tabasco, Mexico, and inAlta Verapaz, Guatemala. The fringe-lipped bat,Trachops cirrhosus coffini Goldman, is known fromcaves in Oaxaca and Veracruz, Mexico; and El Peten,Guatemala. This species may occur in caves in coloniesof up to 50 or more individuals; it is carnivorousand its droppings are white and resemble the feces ofbirds more than those of bats (Hall and Dalquest,1963).Famity VespertilionidaeNineteen species of vespertilionid bat have beenrecorded from the caves of Mexico, Guatemala, andBelize. Some of the records are of species which donot usually inhabit caves and so are not discussedhere.The pallid oat, Antrozous pallidus pallidus (LeConte), is known from caves in Chihuahua and Durango.This species is primarily insectivorous but mayoccasionally eat small lizards.Two species of the genus Eptesicus have oeen recordedfrom caves in :'"lexico and Guatemala. TheCentral American species of this genus have beenstudied by Davis (1965). The tropical brown bat,Eptesicus furinalis gaumeri (.T. A. Allen), is knownfrom caves in Morelos and Yucatan. The big brownbat, E. fuscus (Palisot de Beauvois), is represented incaves by two subspecies: E. fuscus miradorensis H.Allen from Puebla, Tamaulipas, and Veracruz,Mexico, and Alta Verapaz, Guatemala; and E. f. pallidusYoung from one cave in Coahuila. These bats areinsectivorous.Five species of the genus Myotis are known fromcaves in Mexico, Guatemala, and Belize. The CentralAmerican species of the genus have been studied byLaVal (1973). The Cali<strong>for</strong>nia myotis, M. califomicusmexicanus (Saussure), is known only from a cave inTlaxcala. Keays'myotis, M. keaysi pilosatibialis La Val,is known from caves in Quintana Roo, Tabasco, Tamaulipas,Veracruz, and Yucatan, Mexico; the MayaMountains of Belize; and El Peten, Guatemala. Thisinsectivorous species is frequently present in caves inlarge numbers. The black myotis, M. nigricans nigricans(Schinz), is known from caves in Oaxaca, Tamaulipas,and Veracruz, Mexico, and Escuintla, Guatemala.The fringed myotis, Myotis thysanodes Miller,is represented in <strong>Mexican</strong> caves by two subspecies:M. t. aztecus Miller from the state of Mexico; andM. t. thysanodes Miller from Chihuahua and Durango.Three subspecies of the cave myotis, M. velifer J. A.Allen, have been reported from <strong>Mexican</strong> caves. Myotisvelifer incautus J. A. Allen is known from caves inCoahuila and Durango; this is the common subspeciesinhabiting caves in Texas. It is frequently present incolonies containing several thousand individuals.Myotis velifer velifer J. A. AJlen occurs in caves fromSonora and Durango south through the Sierra MadreOccidental and into southern Mexico. Myotis veliferpeninsularis Miller is known only from caves in BajaCali<strong>for</strong>nia Sur (Jones et al., 1965). All species ofMyotis are insectivorous.The only pipistrelle known from <strong>Mexican</strong> caves isPipistrellus subflavus veraecrucis (Ward). It has onlybeen reported from caves on Cofre de Perote, Veracruz.Two species of the genus Plecotus are known from<strong>Mexican</strong> caves. The systematicsof this genus have beenstudied by Handley (1959). The <strong>Mexican</strong> big-eared bat,P. mexicanus (G. M. Allen), is known from caves fromsouthern Chihuahua south to iVlichoacan and east toYucatan. Townsend's big-eared bat, P. townsendiiaustralis Handley, occurs in caves from Chihuahuaand Coahuila south to Michoacan.Order RodentiaTen species of rodent have been reported from<strong>Mexican</strong> caves, but other species utilize caves at leastoccasionally. A few of the species are more closelyassociated with caves than others, and these are brieflydiscussed below.Family CricetidaeThe Tamaulipan wood rat, Neotoma angustapalataBaker, has been reported from caves in the Sierra deGuatemala and Sierra de El Abra, Tamaulipas (Hooper,1953). This species builds its nests in the caves. Signsof the IVlexican wood rat, Neotoma mexicana torquataWard, were found in caves in Veracruz (Hall and Dalquest,1963). The big-eared climbing rat, Ototylomysphyllotis phyllotis Merriam, is known from caves andcenotes in Yucatan (Pearse and Kellogg, 1938; Hatt,1938), and Alta Verapaz, Guatemala (Lawlor, 1969).Two species of Peromyscus have been reported from<strong>Mexican</strong> caves. Hall and Dalquest (1963) reported P.mexicanus (Saussure) from "the gloom of caves" inVeracruz. Dalquest and Roth (1970) reported thatthe white-ankled mouse, P. pectoralis Osgood, was255
- Page 1:
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
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Order DipluraFamily CampodeidaeAlth
- Page 210 and 211: of small soil inhabiting japygoids
- Page 212 and 213: (Linnaeus) has been collected from
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- 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 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
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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.