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nation must await study, but it is probably a troglophile.An undescribed species from two caves in theSierra de EI Abra, San Luis Potosi, and from S6tanode Nogal, Queretaro, is an apparent troglophile. Avery distinctive new species from Cueva del Brinco,Tamaulipas, is unquestionably a higWy adapted troglobite.Family ScolopendridaeScolopendra sumichrasti Saussure is a large speciesfrequently found in the entrance area of caves. It isknown from several caves in the Sierra de El Abra,San Luis Potosi, and the Sierra de Guatemala, Tamaulipas.A large specimen was collected in a remote innerroom of ActUn Loltun, Yucatan.Order ScutigeromorphaFamily PseUiodidaePselliodes sabinorum, described by Chamberlin(1942) from Cueva de Los Sabinos, San Luis Potosi,was listed by Nicholas (1962) as a troglobite. Thisspecies is now considered to be a synonym of thewidespread P. guildingii (Newport) and is presumablya troglophile.GUATEMALAFig. 51.-Distribution of troglophilic and troglobitic centipedes of the families Lithobiidae and Cryptopidae: 1, Newportiapelaezi; 2, Garcibius osorioi; 3, Nuevobius cavicolens; 4, Newportia n. sp. 1; 5, Newportia sabina; 6, Newportia n. sp. 2; 7, eruzobiusatoyacus.172
Family ScutigeridaeScutigera carrizala Chamberlin from Cueva delCarrizal, Nuevo Leon, was listed by Nicholas (1962)as a troglobite; this species shows no signs of caveadaptation and should be considered a troglophile.Scutigera cacahuamilpensis Herrera was describedfrom Grutas de Cacahuamilpa, Guerrero. It is probablya synonym of S. linceci (Wood) which has beenreported from that cave as well as from Grutas deJuxtlahuaca, Guerrero, and Grutas de Atoyac, Veracruz;this species is also a troglophile.Class PauropodaThe only record of pauropods in Mexican cavesis that of a single undetermined specimen collectedin Cueva del Jobo, San Luis Potosi (Bonet, 1953a).Class DiplopodaThe millipeds, better than any other group, characterizethe cave fauna of the region under consideration.Every cave with any appreciable moisture canbe expected to contain at least one and usually severalspecies of milliped. Although 46 troglobitic and90 other species have been reported (see Table 19),the class is still poorly known. Many genera andspecies of both troglobite and troglophile remain undescribedor belong to such poorly known groupsthat specific identifications are not presently possible.Order PolyxenidaeFamily LophoproctidaeThe minute miIlipeds of the order Polyxenida havebeen found in several caves throughout southernMexico, but only one species, Lophoproctinus diversunguisSilvestri, from Grutas de Atoyac, Veracruz,has been identified. The taxonomy of this complexgroup is very different from that of other millipeds,and no systematists are presently working with theNorth American fauna. The polyxenids are usuallyfound in soil and deep litter, and the cavernicolespecies are presumably all troglophiles.Order GlomeridesmidaFamily GlomeridesmidaeThe order Glomeridesmida in North America isrepresented by six species from Panama, one fromGuatemala, and this Mexican troglobite. Shear(1974) reports five undescribed epigean species fromMexico. All of the described forms have been placedin the genus Glomeridesmus, but Shear believes thatat least three genera are represented.Table 19.-Summary of cave inhabiting Diplopoda.Troglobites Other SpeciesPolyxenidaLophoproctidae 0 1GlomeridesmidaGlomeridesmidae 1 0GlomeridaGlomeridae 4 0ChordeumidaCleidogonidae 5 3Trichopetalidae 4 0JulidaJulidae 0 1Paraiulidae 0 1*Polydesmida 23 64PolyzoniidaPolyzoniidae 0 2Siphonophoridae 0 1SpirobolidaAtopetholidae 0 1Messicobolidae 0 2Rhinocricidae 0 4Spirobolellidae 1 0Spirobolidae 0 2SpirostreptidaCambalidae 6 0Spirostreptidae 2 6StemmiulidaStemmiulidae 0 1PlatydesmidaPlatydesmidae 0 1Total 46 90*For summary by families see Table 20.Glomeridesmus sbordonii ShearGlomeridesmus sbordonii Shear, 1974:241,245,246,248, fig. 1-4; Sbordoni et al., 1974: 15,30 (nomennudum); Sbordoni, 1974:366.Type-locality.-Grutas de Cocona, Teapa, Tabasco,Mexico.Distribution.-Known from the type-locality andpossibly one cave in Chiapas. See Fig. 52.Records.-Chiapas: ?Cueva de la Golondrina; Tabasco:Grutas del Cocona.Discussion.-Specimens from Grutas del Coconahave been taken in large numbers from rotten woodnear the end of the cave.Order GlomeridaFamily GlomeridaeThe family Glomeridae is represented in the NewWorld only by two genera: Onomeris contains severalepigean species in the southeastern United States;Glomeroides contains the four troglobitic species includedhere, two described epigean species from173
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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
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 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,
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Vigna Taglianti, 1972:118, 121-127,
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Veracruz) belongs in another subgen
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Troglobacanius sbordonii VomeroTrog
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Ptomaphagus (Adelops) is a group of
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(A.) speLaeus, from Grutas de Cacah
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epresented by many species. Tenebri
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Myodopsylla gentilis Jordan and Rot
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genus Pholeomyia have been collecte
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Wheeler (1938) has reported the pre
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263.264, 265, 267, 270, 273, fig. 5
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Anophthyctys jordani: Stefanelli, 1
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Prietella phreatophila Carranza, 19
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TyphlilJs: Pearse, 1938a:13, 15; Hu
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Family BufonidaeFour species of toa
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Oaxaca; and E. yucatanensis Lynch f
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sity of bats in the tropics. In add
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Three species of short-tailed bats
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abundant in the entrance area of Cu
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Andrews, E. W., IV. 1970. Balankanc
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Bolivar y Pieltain, C. 1940. Explor
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Brignoli, P. M. 1979a. The morpholo
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Conde, B. 1975. Description du prem
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Ediger, G. 1970. The Birthday Passa
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Gonzalez Ochoa, A. 1963b. Relacione
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Hobbs, H. H.,Jr., and T. C. Barr, J
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Jordan, c. B. 1937. Bringing in the
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McKenzie, D. 1965a. Caves of the Si
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Monod, T. 1975. Sur la distribution
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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
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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
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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.
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