Anophthyctys jordani: Stefanelli, 1954a:436-438(erroneous spelling).<strong>Cave</strong> tetra: Woods, 1954:4.Astyanax jordani: Barr, 1956:12-13; Bull, 1969:37;Mitchell, 1969a:80; Morris, 1969:26, 27; Popper,1970:552-562; Hubbs, 1971:93; Reddell, 1971b:82-83; Reddell and Mitchell, 1971a: 159; Reddelland Elliott, 1973a:172; Johnson and Heath, 1977:132; Moore and Nicholas, 1978:116-117; Thinesand Weyers, 1978:35-55, fig. 1-2.Anoptichtys jordani: Sadoglu, 1957b:432-439; Berra,1963:44-45 (erroneous spellinl!;).Anoptichtys hubbsi: Sadoglu, 1957b:432-439 (erroneousspelling).Anoptichtys antrobius: Sadoglu, 1957b:432-439 (erroneousspelling).Anoptichtys: Sadoglu, 1957b:433; Parzefall andWilkens, 1972:69 (erroneous spelling).Anopichthys: Woods and Inger, 1957:232 (erroneousspelling).Anaptichthys jordani: Tamayo, 1962:vol. IlI:328(erroneous spelling).Astyanax (Anoptichthys) jordani: Poulson, 1964:751, 754, 755, 756, 758, 760, 761, 762, 763, 764,767.Astyanax (Anoptichthys) jordani hubbsi: Poulson,1964:757.Astyanax (Anoptichthys) jordani jordani: Poulson,1964:757.Astyanax (Anoptichthys) spp.: Poulson, 1964:767.Anoptichthys jordan: Vandel, 1964:477 (erroneousspelling).Astyanax ("Anoptichthys"): Greenwood, 1967:268.Astyanax mexicanus jordani: WaIters and Liu, 1967:437, 441, 442.Astyanax mexicanus, cave derivatives of: Peters andPeters, 1968:213, 219,230,231,235,236; Mitchell,1973:146; Schemmel, 1974a:196-215, fig. 1-7;Besharse and Brandon, 1976:538; Mitchell et aI.,1977:1-89, fig. 1-21.Astyanax hubbsi: Bull, 1969:37; Reddell, 1971b:82;Reddell and Mitchell, 1971a: 159; Schmatolla,1972:555, 556, 558, 559-564, 571, fig. IB, 2-3,4B, 5B; Reddell and Elliott, 1973a: 172; Schmatollaand Erdmann, 1973:705, 708; Voneida,1973:462-463; Egar, 1974:350; Sligar, 1974:467;Fernandez Ruiz, 1976:712, 716; Sligar and Voneida,1976:107-124, pI. l(fig_ 1-4), pI. 2(fig. 5-8),pI. 3(fig. 9-10), pI. 4(fig. 11-12), pI. 5(fig. 13),pI. 6(fig. 14), pI. 7(fig. 15), pI. 8(fig. 16), pI. 9(fig.17); Voneida and Sligar, 1976:89-106, pI. l(fig.4), pI. 2(fig. 8-9), pI. 7(fig. 18-19), pI. 8(fig. 2021); Yew and Yoshihara, 1977:175-180, fig. 1-9.Astyanax antrobius: Bull, 1969:37; Mitchell, 1970b:64; Reddell, 1971b:82; Reddell and Mitchell,1971a:159, fig. 14-15; Reddell and Elliott, 1973a:172; Kawakatsu, 1974:27, fig. 15; Kawakatsu,1976:36, fig. 3.Astyanax mexicanus: Copp, 1969:381; Wilkens,1970b:173-199; Parzefall and Wilkens, 1972:64,71, 77-78; Wilkens, 1972a:l-11; Wilkens andBurns, 1972:263, 266, 267; Avise and Kitto,1973:113-132.Anoptichthys antobius: Kleerekoper, 1969: 127 (erroneousspelling).Astyanax, blind: Mitchell, 1969a:76; Elliott, 1971:3-5,42; Elliott and Mitchell, 1973:172; Vomero,1974:345; Fernandez Ruiz, 1976:712.Astyanax: Mitchell, 1969d:map, letter; Reddell andMitchell, 1971c:2; Kawakatsu, 1973b:255; Reddell,1973a:36.Characins: Thines, 1969 :3-4, 12.Anophtichtys hubbsi: de Lachica B., 1970:62 (erroneousspelling).Anophthycthys hubbsi: de Lachica B., 1970:122 (erroneousspelling).Anoptychtys: Pasquini, 1970:470 (erroneous spelling).Astyanax fasciatus, cave <strong>for</strong>ms of: Wilkens, 1970c:54-75.Astyanax sp.: Mitchell, 1971b:136; Reddell, 1971b:82; Reddell and Mitchell, 1971a:143, 158; Reddelland Mitchell, 1971b:196; Reddell and Elliott,1973a: 178; Reddell and Elliott, 1973b: 182, 188,190.Anotichthys jordani: Reddell and Mitchell, 1971a:137 (erroneous spelling).Astyanax mexicanus, blind characin: Tavolga, 1971:171.Astyanax, eyeless: Wiley and Mitchell, 1971:231-239;Mitchell, 1977:171-173; Mitchell and Cooke,1977:175-178; Mitchell and Elliott, 1977:179184; Mitchell and Russell, 1973:261; Mitchell andRussell, 1976a: 169-172; Mitchell and Russell,1976b:163-167.Astyanax mexicanus, troglobitic populations of:Avise and Selander, 1972: 1-16.Astyanax (=Anoptichthys): Sbordoni and Argano,1972: 10-11; Mitchell, 1975:360-361.Astyanax (=Anoptichthys) jordani: Shordoni andArgano, 1972: 12.Astyanax, <strong>Mexican</strong> cavefish: Alley et a!., 1973:263.Astyanax, cave relatives of: Kirpichnikov, 1973:224.Astyanax mexicanus, cave populations of: Kosswig,1973:4-5,9, 10, 13; Parzefall, 1973h:284; Scholl,1973:291,292,296; Chakraborty and Nei, 1974:460-461,467-469; Schemmel, 1974h:193-201, fig.242
1-3; Peters et al., 1975:110-124, fig. 1-4; Li andNei, 1977:913; Wilkens, 1977a:137-148, fig. 1-8;Wilkens, 1977b:604; Giuseffi et al., 1978:679.Astyanax mexicanus, cavernicole <strong>for</strong>m: Kosswig,1973:13; Schemmel, 1973:209-219; Wilkens,1973c:55, 57-58; Durand, 1976:18; Parzefall,1976:29; Thines and Piquemal, 1978:197.<strong>Mexican</strong> eyeless characin fishes: Mitchell and Cooke,1973:147; Mitchell and Elliott, 1973:147.Astyanax mexicanus (blind cave fish): Oguri andOmura, 1973:414.Astyanax mexicanus {=Anoptichthys}: Peters andPeters, 1973a:187-201, fig. 1-5: Peters et al.,1973:428-430.Astianax: Sbordoni and Cobolli-Sbordoni, 1973b:150 (erroneous spelling).Astyanax, hypogean <strong>for</strong>m: Schemmel, 1973:209219.Astyanax complex, blind and depigmented derivativesof: Schroder, 1973:vi.Astyanax mexicanus, blind cave varieties: Sadoglu,1975:419-426, fig. 1-9.Blind tetras: Kirby et al., 1977:578-579, fig. lC.Anopthichthy's: Thines and Piquemal, 1978:197(erroneous spelling).Astyanax mexicanus or Anoptichthys. blind <strong>for</strong>m of:Durand, 1979:269.Type-Iocality.-Of Anoptichthys jordani: Subterraneanstream southwest of Valles (=Cueva Chica),San Luis Potosi, Mexico; of Anoptichthys antrobius:Cueva de EI Pach(lll, Antip;uo \lorelos, Tamaulipas,Mexico; of Anoptichthys hubbsi: Cueva de Los Sabinos,Ciudad Valles, San Luis Potosi, Mexico.Distribution.-Known from caves in the Sierra deEI Abra, Tamaulipas and San Luis Potosi: the Sierrade Guatemala, Tamaulipas: and the J\licos region, SanLuis Potosi. See Fig. 85.Records.-San Luis Potosi: Sotano del Arroyo, EICuate Este, EI Cuate Oeste, Cueva Chica, Cueva de laCurva, Sotano de Japones, Sotano de Jos, Cueva delLienzo, S()tano de i\latapalma, Sistema de \lontecillos,Cueva del ()tate, S()tano de Palma Seea, Siltanode las Piedras, Cueva del Rio Snbterraneo. Sotano dela Roca, Cueva de Los Sabinos, S()tano de Soyate,Sotano del Tigre, S()tano de la Tinaja, S()tano delToro, and S()tano de Yerbaniz: Tamaulipas: Bee <strong>Cave</strong>,Sotano del Caballo Moro, Sotano Escondido, Sumiderodel .lineo, Sotano del \lolino, Cueva de EIPachon, Siltano de Vasquez, and Sotano de EI Venadito.Discussion.-Astyanax jordani has been more intensivelystudied than any other species of cavernicolein the world. Essentially every aspect of itsmorphology, behavior, and genetics has been examinedat least once. Despite this intense examinationconsiderable confusion remains about the origin ofthe species and its relationships to its parent species,Astyanax mexicanus (Filippi). The species is presentlyunder continuing study by several Europeanzoologists, including H. Wilkens, J. Parzefall, G. andN. Peters, G. Thines, and C. Schemmel. The habitats,evolution, and various aspects of morphology havebeen recently studied in considerable detail by Mitchellet al. (1977). It is hoped that their clear enunciationof the relationship of this fish to the geology andphysiobrraphy of the region in which it occurs and ofits evolution will allow others to appraise more intelligentlythe results of their own studies. Three specieswere originally described from rather isolated parts ofthe range of the species as we know it now. With thediscovery of populations in 30 caves, a far better understandingof the hydrology of this part of Mexico,and the genetic studies of Breder, Sadoglu, Wilkens,and their colleagues, there is no justification <strong>for</strong> recognizingmore than one species of eyeless characinin Mexico. Breder and his colleagues, in particular,have studied the genetics and habitat of this speciesin the type-locality of A. jordani, Cueva Chica, SanLuis Potosi. They discovered that the fish was hybridizingwith A. mexicanus to a certain extent in thecave. Furthermore, it has been demonstrated that thespecies will readily hybridize with A. mexicanus underlaboratory conditions. It is significant, however,that the continual introduction of eyed Astyanaxinto Cueva Chica has not led to an exclusively hybridpopulation even there, and, furthermore, it has beendemonstrated that the eyed fish when raised in darknessfrequently exhibit both morphological andphysiological abnormalities. Based solely on thephysical barriers presented by the cave environment<strong>for</strong> A. mexicanus and by the epigean environment <strong>for</strong>iI. jordani, I feel that it is justified and useful to treatthe two as distinct species. It is almost certain that A.mexicanus can never successfully compete with thecave-adapted A. jordani in the cave, nor that the blindA. jordani could hope to survive in the epigean environment.Although the two species have not divergedenough to lead to complete genetic isolation, theyhave diverged enough to prevent the significant survivalof one species in the environment of the other.Family IctaluridaePrietelLa phreatophila CarranzaAmeiuridae, ciego y depigmentado: Bolivar and Carranza,1954: 115.243
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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,
- 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
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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 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
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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.