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Discussion.-This species is distinct from othermembers of the genus Uncuklbes and in many respectsresembles Aceratophallus. It is interesting that thistroglobite inhabits the same caves as does the lesshighly evolved Strongylodesmus harrisoni. It is veryuncommon in the area, whereas S. harrisoni is abundant.It is possible that S. harrisoni is a more successfulrecent invader.Uncuklbes crispus CauseyPolydesmoid, undescribed genus and species: Reddell,1967d:l06.Rhachodesmidae, undescribed genus and species:Reddell, 1971a:44 (San Luis Potosi records only).Uncuklbes crispus Causey, 1971b:30, 31-32, fig. 1820; Causey, 1973:118; Reddell, 1973a:34, 40;Shear, 1974:264,265,267,268.Uncuklbes versatilis Causey, 1971b:32; Causey, 1973:118; Reddell, 1973a:34, 40; Shear, 1974:264,265,267.Type-Iocality.-Of Uncuklbes erispus: Sotano deHuitzmolotitla, 2 km SW of Tlamaya, which is 10 kmNE of Xilitla, San Luis Potosi, Mexico; of Uncuklbesversatilis: Cueva de Potrerillos, 1.5 km W of Ahuacatlan,San Luis Potosi, Mexico.Distribution.-Known from caves in the Xilitla,Ahuacatlan, and Aquismon regions, San Luis Potosi.See Fig. 60.Records.-QuerE!taro: Sumidero del Llano Conejo;San Luis Potosi: Sotano de Guadalupe, Sotano deHuitzmolotitla; Cueva de los Potrerillos, and Sotanode Tlamaya.Discussion.-This comparatively widespread speciesshows some variation from area to area but notenough for taxonomic recognition. U. crispus isapparently most closely related to U. arganoi to thewest in Queretaro. The two areas are well separatedstratigraphically. Uncuklbes crispus was found inSotano de Huitzmolotitla on mud and silt banks inincredibly large numbers; the collectors reportedthat they literally turned mud banks white withtheir bodies. This species is extremely delicate, andmost preserved specimens are badly broken.Uncuklbes porrensis ShearUncuklbes porrensis Shear, 1974:264,268, fig. 45-49.Type-locality.-Cueva de la Porra, 3 mi north ofXilitla, San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 60.Discussion.-This species is closely related to U.arganoi and U. crispus.Family SphaeriodesmidaeThe family Sphaeriodesmidae is well representedin caves, and seven species are known only from caveswith many specimens still unstudied. Nicholas (1962)listed Cylionus kauanus Chamberlin from ActunKaua, Yucatan, as a troglobite, but it is doubtful ifany sphaeriodesmids can be reliably listed as troglobites.The cave inhabiting species show no significantdifferences from species which inhabit endogean habitats.Shear (1974) described four species from Mexicancaves: Sphaeriodesmus cruzbelem, S. golondrinensis,and S. zontehuitz are known only from .;avesin Chiapas; S. nortoni has been identified from CuevaChica de la Perra, Tamaulipas. Shear (1977a) addedS. trulkltus and S. redondo from caves in Chiapas.Collections of this family from many caves in NuevoLeon, Tamaulipas, Veracruz, Oaxaca, Campeche, andYucatan await study. See Fig. 61 for the distributionof this family in Mexican caves.Family TrichopolydesmidaeThe use of the name Trichopolydesmidae for thesmall millipeds discussed here is uncertain. Shear(1977a) has discussed the status of these species andspeculates that two or more families of small polydesmoidmillipeds now considered trichopolydesmidsmay occur and that the family Trichopolydesmidaemay not be present in Mexico. As presently understood,the family is represented in Mexico by two epigeanspecies in Baja California and by the six describedtroglobites listed here. Berlese collections ofsurface litter have resulted in the collection of additionalundescribed epigean species. Undescribed troglobitesare known from caves in Nuevo Leon, Oaxaca,San Luis Potosi, Veracruz, Campeche, and Yucatan.Caramba delburro ShearCaramba delburro Shear, 1977a:245, 246, 247, 248249, fig. 24-28.Type-Iocality.-Cueva del Burro, Rio Escondido,Lago de Malpaso, Ocozocuatla (=Ocozocoautla), Chiapas,Mexico.Distribution.-Known only from the type-locality.See Fig. 54.Discussion.-The genus Caramba is known onlyfrom the three troglobitic species listed here. It ismore closely related to Tylogoneus tbm to othertrichopolydesmid genera. These two genera are consideredby Shear (1977a) to belong to a group whichincludes the genera Chaetaspis and Antriadesmus ofthe United States.188
Caramba delnegro ShearCaramba delnegro Shear, 1977a:246, 247, 248, 250,251, fig. 30.Type-locality.-Cueva del egro, Bochil, Chiapas,Mexico.Distrihution.-Known only from the type-locality.See Fig. 54.Discussion.-The shape of the gonopods of thisspecies is similar in some respects to that of the speciesof Tylogoneus, but other characters clearly placeit in Caramba.Caramba grandeza ShearCaramba grandeza Shear, 1977a:247, 248, 249, 250,fig. 29.Type-locality.-Grutas de Llano Grande, LlanoGrande, La Grandeza, Huiztla, Chiapas, Mexico.Distribution.-Known only from the type-locality.See Fig. 54.Discussion.-This species, known only from males,is closely related to C. delburro.MEXICO--', /" ,,/, /'/IIII),/(/IGUATEMALAFig. 61.-Distribution of troglophilic millipeds of the family Sphaeriodesmidae: 1, Sphaeriodesmus nortoni; 2, S. redondo;3, S. golondrinensis; 4, S. %ontehuit%; 5, S. cru%belem; 6, S. trullatus; 7, Cylionus kauanus.189
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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
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 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
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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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