springs in the vicinity of Cautro Cienegas de Carranza,Coahuila. This unusual region produced a new genusand two new species of amphipod (Holsinger andMinckley, 1971), a new genus and species of stenasellidisopod (Cole and Minckley, 1972), and a newgenus and two new species of cirolanid isopod (Coleand Minckley, 1970).In January 1969 several caves in the Sierra de EIAbra were visited by R. W. Mitchell, James Reddell,and William Russell. A collection made in Cueva Pintaincluded a new species of trichoniscid isopod (Schultz,1970a). In March several caves in the Sierra de Guatemalaand Sierra de EI Abra were visited by R. W.Mitchell, James Reddell, Suzanne Wiley, and others.The most notable collection was of a new species ofpseudoscorpion in Cueva de la Florida, Tamaulipas(Muchmore, 1972c). Aerial reconnaissance of theSierra de EI Abra was conducted by Robert W.Mitchell, Richard O. Albert, and William Russellduring 1969 with the result that numerous new cavescontaining blind fish of the genus Astyanax were located.On July 31, 1969, William Elliott collected thethird species of blind scorpion to be found; it wastaken from Sotano de Yerbaniz, San Luis Potosi, andwas described by Mitchell (1971b) as Typhlochactaselliotti.From June 18 to September 13, 1969, Stewart B.Peck and James H. Peck conducted a field trip tomany parts of Mexico and Guatemala (Peck andPeck, 1973). Their field trip resulted in the collectionof several new species of troglobite. Among areasvisited were Cuesta de Chipinque, Cueva de Chorrosde Agua, and Cueva de la Boca, Nuevo Leon; theSierra de Guatemala and northern Sierra de EI Abra,Tamaulipas; the southern Sierra de EI Abra and theXilitla region, San Luis Potosi; Pinal de Amoles,Queretaro; Paraje Nuevo and Tlilapan, Veracruz; theChiapas higWands; and several areas in Guatemala.Among the many new species discovered were blindspiders (Gertsch, 1971a) and blind millipeds (Shear,1972; 1974).From October 10 to December 9,1969, the AccademiaNazionale dei Lincei of Italy sponsored a zoologicalexpedition to Mexico. It was conducted byValerio Sbordoni, Roberto Argano, and VittorioParisi. They visited 17 caves during the course of theirexpedition, including caves in the Sierra de Guatemala,Tamaulipas; Sierra de EI Abra, San Luis Potosiand Tamaulipas; western Xilitla region, Queretaro;Cacahuamilpa region of Guerrero and Mexico; andcaves near Orizaba and Cordoba, Veracruz. Althoughmost of their collections included only species previouslystudied, several at that time were still undescribedand a few additional <strong>for</strong>ms of interest wereobtained. Many of the results of this expedition werecollected in a volume edited by Sbordoni and Arganoand published in 1972.In March 1970 two German zoologists, HorstWilkens and Jakob Parzefall, visited several caves inthe vicinity of Micos, San Luis Potosi, <strong>for</strong> the primarypurpose of collecting and studying new populationsof blind Astyanax. Among the fauna collectedwere two new species of troglobitic stenasellid isopodin Cueva del Huisache (Magniez, 1972). The Sierra deTamaulipas, Tamaulipas, was visited <strong>for</strong> the first timewhen, on October 31, 1970, William Russell, Gil Ediger,and Jill Ediger explored and made collections intwo caves there. Most of the remaining work of the<strong>Association</strong> <strong>for</strong> <strong>Mexican</strong> <strong>Cave</strong> <strong>Studies</strong> in 1970 wasdevoted to the continuing study of the caves of theSierra de EI Abra.In January 1971 Jerry Cooke, William Elliott,Robert W. Mitchell, James Reddell, Suzanne Wiley,and others from Texas Tech University were joinedby Dr. Masaharu Kawakatsu of Fuji Women's College,Japan, on a trip to the Sierra de Guatemala, Tamaulipas.Among numerous specimens collected were twotroglobitic and one troglophilic flatworm of the genusDugesia, the first to be reported from Mexico (Mitchelland Kawakatsu, 1973a).The Second Zoological Mission to Mexico sponsoredby the Accademia Nazionale dei Lincei wasconducted from January 24 to April 11, 1971, byValerio Sbordoni, Roberto Argano, and Aldo Zullini.This expedition was devoted entirely to extremesouthern Mexico and adjacent Guatemala. Two caveswere visited in Tabasco and 24 in Chiapas. The resultsof this trip, together with additional studies of speciescollected on the First Zoological Mission to Mexico,were published in 1974 in a volume edited by Sbordoni,Argano, and Zullini. This expedition to southernMexico resulted in the discovery of many speciesof interest, among them the first nematodes to bereported from caves in Chiapas (Zullini, 1974), newspecies of troglobitic amphipods (Ruffo and VignaTaglianti, 1974), shrimp (Holthuis, 1974), opilionid(Silhavy, 1974), milliped (Shear, 1974), anillinebeetle (Vigna Taglianti, 1974), histerid beetle (Vomero,1974), and ricinuleid (Brignoli, 1974c).In September 1971 Terry Raines visited severalcaves in the Canon de Huasteca region of NuevoLeon. In Sotano del Anticlino he discovered a newspecies of endemic earthworm (Gates, 1973). InNovember David McKenzie and William Russellvisited Cueva del Ojo de Agua de Manantiales, SanLuis Potosi, where they collected a new genus andspecies of troglobitic trichopolydesmid milliped(Causey, 1973). They also visited Cueva del Hui-10
sache in the Micos region of San Luis PotosI andfound a new genus and species of troglobitic cirolanidisopod (Bowman, 1975). Also during NovemberDavid McKenzie visited several caves in southernMexico, including Grutas de Atoyac, Veracruz, andGrutas del Cocona, Tabasco.The biological activities of the <strong>Association</strong> <strong>for</strong><strong>Mexican</strong> <strong>Cave</strong> <strong>Studies</strong> reached their highest levelduring 1972. David McKenzie visited several cavesin Yucatan in January. Among collections of interestwere additional specimens of the troglobitic spirostreptidOrthoporus zizicolens.On March 27-30 the Grupo Espeleologico <strong>Mexican</strong>oconducted an expedition to the Huautla de Jimenezregion of Oaxaca, which was directed by Jorge Urquijo.In S6tano de San Agustin, Jesus HernandezLopez collected specimens of a new species of troglobiticcarabid beetle (Hendrichs and Bolivar, 1973).William Elliott, Pam Lynn, Mike McEachern, andRon Ralph continued the investigation of the karstregion of Valle de los Fantasmas, San Luis Potosi,in May 1972 with the exploration of several caves inthat area (Elliott and Reddell, 1973). In June studywas continued in the Pinal de Amoles rel,rion whennumerous new caves were located and explored byWilliam Russell, Terry Ibines, I{oll Ralph, and others.Discoveries included additional populations of blindtrechine beetles, spiders, millipeds, and isopods.Most of these results remain unpublished.Study of the cave fauna of northwestern Mexicowas resumed with a trip to this area on June 10-22,1972, by Ed Alexander, William Elliott, Carl Kunath,Anf.,rie McLaughlin, and James Reddell. <strong>Cave</strong>swere visited in the vieinities of Parral, Chihuahua, andlVlapimi and Picardias, Durango (Reddell, 1973c).Arnon~ the more signifieant diseoveries were additionalspeeimens of both species of reliet ricinuleidfrom Durango, a new species of troglobitic amphipod(Holsinger, 1(73), and a new species of leptonetidspider (Gertsch, 1974).During August 1972 Jerry Cooke, Rexell Mitchell,Robert \V. Mitchell, and William Russell visited eavesin Chiapas, Tabasco, and Yucatan. The fauna collectedincluded a new family, genus, and species oftroglobitic planarian (Dimarcus villalobosi), a newspecies of planarian of the genus Dugesia (l\'1itchelland Kawakatsu, 1973b), a new species of asellidisopod (Bowman, 1976), a new species of troglobiticpseudoscorpion (Muchmore, 1973a), and specimensof the blind crab Typhlopseudothelphusamocinoi from two additional localities.The intensive study of tile caves of the Pinal deAmoles region continued in November 1972 withexploration in several caves by Roy Jameson, PeterStrickland, and others. The area was also visited inDecember by Roy Jameson and others, at whichtime a collection in Cueva de Emilia resulted in thediscovery of the first troglobitic homopteran knownfrom Mexico (Fennah, 1973).<strong>Cave</strong>s in the Sierra de EI Abra, Tamaulipas, andAquismon and XiJitia regions of San Luis Potosiwere visited November 21-29, 1972, by Jan Lewis,James Reddell, Terry Raines, Jack White, and others.The collections included new species of opilionid(Goodnight and Goodnight, 1973), pseudoscorpion(Muchmore, 1973c), and milliped (Causey, 1973).One of the more successful expeditions conductedby members of the <strong>Association</strong> <strong>for</strong> <strong>Mexican</strong> <strong>Cave</strong><strong>Studies</strong> was that made from December 22, 1972, toJanuary 8, 1973, by David McKenzie, Martha HelenMcKenzie, Stuart Murphy, and James Reddell (Reddell,1973e). <strong>Cave</strong>s were visited in the Sierra deTamauJipas, Tamaulipas; the Tezonapa region, Veracruz;and the Acatlan, Valle Nacional, San Sebastiande las Grutas, and Apoala regions of Oaxaca. Althoughmany undescribed species were found in allof the regions visited, the most productive were theTezonapa, Acatlan, and Valle Nacional regions. AtTezonapa collections included a new species of glomeridmilliped (Causey, 1973). The spectacular cavenear Acatlan, Cueva del Nacimiento del Rio SanAntonio, yielded specimens of numerous troglobites,including the first cavernicolous alpheid shrimpknown (Hobbs, 1973b), a palaemonid shrimp (Hobbs,1973b), the second troglobitic crayfish from Mexico(Hobbs, 1973a), a mysid (Bowman, 1973), a still undescribedcatfish, and the second species of blindtheraphosid known (Gertsch, 1973). The Valle Nacionalregion proved to be almost as biologicallyamazing. The two most productive caves were Cuevadel Guano and Cueva del Guayabo. New troglobiticspecies included crayfish and palaemonid shrimp(Hobbs, 1973b), pseudoscorpions, and isopods.In February 1973 a second extensive cave was discoverednear Acatlan. This cave, Cueva de LagunaVerde, has been little studied biologically, bu t collectionsby Peter Strickland and James Rodemakerincluded specimens of blind crayfish and shrimp.Serious study of the cave fauna of the YucatanPeninsula by members of the <strong>Association</strong> <strong>for</strong> <strong>Mexican</strong><strong>Cave</strong> <strong>Studies</strong> began in 1973. In a trip from February16 to May 14, partly supported by The Museum,Texas Tech University, Mary Butterwick,David McKenzie, Martha Helen McKenzie, StuartMurphy, and James Reddell visited numerous cavesin Veracruz, Oaxaca, Campeche, and Yucatan. Themost significant find in the non-Yucatan areas wasof a new species of troglobitic pseudoscorpion m11
- Page 1: A REVIEW OF THECAVERNICOLE FAUNA OF
- Page 8: . 1-----,, ,", ,\'"I/I~" )/Fig. I.-
- Page 12 and 13: collected a troglobitic cirolanid i
- Page 14 and 15: C. Bolivar and J. Carranza investig
- Page 18 and 19: Cueva del Diablo near Orizaba, Vera
- Page 20 and 21: Those species of troglobite which h
- Page 22 and 23: Table 2.-(continued)1970. Eodrilus
- Page 25 and 26: CAVE REGIONSOnly the cave regions o
- Page 27 and 28: Table 3.-(continued)TroglobitesTrog
- Page 29 and 30: y the waters of the Amistad Reservo
- Page 31 and 32: Although a few caves in the Norther
- Page 33 and 34: an elevation of about 900 meters. T
- Page 35 and 36: the gryllid cricket Paracophus suba
- Page 37 and 38: has been Jiscussed by Segerstrom (1
- Page 39 and 40: only apparent troglophile is the ps
- Page 41 and 42: The cave fauna of this region is ri
- Page 43 and 44: igneous rock immediately east of th
- Page 45 and 46: Rio Soto la Marina and terminates a
- Page 47 and 48: caves studied are many large stream
- Page 49 and 50: unusual species is Neogovea mexasca
- Page 52 and 53: pallidus Rowland, the nesticid spid
- Page 54 and 55: troglobites: the trichoniscid isopo
- Page 56 and 57: Cambridge, and Pholcophora bispinos
- Page 58 and 59: of the genus Lutzomyia has not been
- Page 60 and 61: The principal cavernous fonnation i
- Page 62 and 63: ocks of Paleocene to Pliocene age.
- Page 64 and 65: Beach and Supra-tidal Zone, the I\o
- Page 66 and 67:
The aquatic fauna of Mexico and Cen
- Page 68 and 69:
oaxacae Hobbs from the Valle Nacion
- Page 70 and 71:
described, the genus occurs in cave
- Page 72 and 73:
large genus with numerous epigean s
- Page 74 and 75:
milliped, laniatorid harvestmen, an
- Page 76 and 77:
Table 4.-Summary of cave inhabiting
- Page 78 and 79:
Discussion.-This large, troglobitic
- Page 80 and 81:
any way a part of the true cave fau
- Page 82 and 83:
they have been washed in with soil
- Page 84 and 85:
part of the aquatic fauna. A very l
- Page 86 and 87:
The genus Helicodiscus is a signifi
- Page 88 and 89:
associated with true cave waters. P
- Page 90 and 91:
Family DaphnidaeWilson (1936) repor
- Page 92 and 93:
tul), Cueva de San Isidro, Cueva de
- Page 94 and 95:
30 m below entrance of Cueva de la
- Page 96 and 97:
underflow of rivers, alluvial grave
- Page 98 and 99:
the similarities between the distri
- Page 100 and 101:
Yucatan but did not identify the sp
- Page 102 and 103:
Protrichoniscus: Reddell, 1967a:24.
- Page 104 and 105:
It indudes five spccies, of which t
- Page 106 and 107:
Type-locality.-Pozzo, Etla, Oaxaca,
- Page 108 and 109:
Records.-Campeche: Volc
- Page 110 and 111:
164; Pearse, 1945:167, 168, fig. 13
- Page 112 and 113:
Family AtyidaeThe family Atyidae is
- Page 115 and 116:
Creaseria morleyi (Creaser)Blind sh
- Page 117 and 118:
the pond which is fed by the cave s
- Page 119 and 120:
philes. Pseudothelphusa (Pseudothel
- Page 121 and 122:
Typhlochactas elliotti MitchellTyph
- Page 123 and 124:
Distribution.-Known only from the t
- Page 125 and 126:
(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 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 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,
- 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 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
- Page 308:
Cueva de los Riscos, 2 km S Rancho
- Page 311 and 312:
Cueva de El Coy-see Cueva del Nacun
- Page 313 and 314:
+*Cueva del Aire, 1.8 km at S85°W
- Page 315 and 316:
*Sistema Purificacion, Conrado Cast
- Page 317 and 318:
*Cueva del Nacimiento del Rio Frio,
- Page 319 and 320:
Sotano de Humo, 5 km NW San Andres,
- Page 321 and 322:
and Bohor, 1975:25-27, 26m; Reddell
- Page 323 and 324:
*Actun Xpukil, 3 km S Calcehtok, Mp
- Page 325 and 326:
Appendix 2LIST OF TROGLOBITES BY ST
- Page 327 and 328:
Paravachonium superbum-I'seudoscorp
- Page 329 and 330:
SUPPLEMENTWhile this manuscript was
- Page 331 and 332:
Cambala speobia (Chamberlin) (p. 19
- Page 333:
mida (Arachnida) in the New World.