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A REVIEW OF THECAVERNICOLE FAUNA OF MEXICO,GUATEMALA, AND BELIZEBULLETIN 27 OF THE TEXAS MEMORIAL MUSEUMTHE UNIVERSITY OF TEXAS AT AUSTIby James R. Reddell

A PUBLICATION OF TEXAS l\IEMORIAL MUSEUM, THE UNIVERSITY OF TEXAS AT AUSTINA REVIEW OF THE CAVERNICOLE FAUNAOF MEXICO, GUATEMALA,AND BELIZEby James R. ReddellBULLETIN 27July 1981

. 1-----,, ,", ,\'"I/I~" )/Fig. I.-Physiographic provinces of Mexico, Guatemala, and Belize: 1, Baja California; 2, Buried Ranges; 3, Sierra MadreOccidental; 4, Basins and Ranges; 5, Edwards Plateau; 6, Sierra Madre Oriental; 7, Gulf Coastal Lowlands; 8, Central Mesa;9, Neovolcanic Plateau; 10, Sierra Madre del Sur System; 11, Pacific Coastal Lowlands; 12, Chiapas-Guatemala Highlands;13, Yucatan Peninsula.Arkansas (McDaniel and Smith, 1976), Florida(Warren, 1961; Peck, 1970), Georgia (Holsinger andPeck, 1971), Hawaii (Howarth, 1972, 1973), Illinoisand southeastern Missouri (Peck and Lewis, 1977),Kentucky (Barr, 1967b; Barr and Kuehne, 1971),Missouri (Craig, 1975, 1977), New Jersey (Nicholas,1976), New Mexico (Barr and Reddell, 1967; Welbourn,1978), Oklahoma (Black, 1971), Pennsylvania(Holsinger, 1976), Tennessee (Barr, 1961), Texas(Reddell, 1965b, 1966b, 1967e, 1970a, 1970b,1970c; Reddell and Mitchell, 1969; Mitchell andReddell, 1971; Lundelius and Slaughter, 1971),Virginia (Holsinger, 1963), volcanic caves of westernNorth America (Peck, 1973b), West Virginia (Holsingeret al., 1976), and Canada (Peck and Fenton,1973). Our knowledge of the fauna of tropical andsubtropical America is much more limited (Peck,1974b). Preliminary reports have now been publishedon the cave fauna of Puerto Rico (Peck, 1974a),Jamaica (Peck, 1976), Cuba (Orghidan et al., 1973,1974; Silva Taboada, 1974), and Panama (Peck,1971a). No summaries exist for the cave fauna of theremainder of the islands of the West Indies, thecountries of Central America, nor any part of SouthAmerica.It is safe to assume that far less than half of thecave-inhabiting species of Mexico, Guatemala, andBelize are now known. In Mexico, which has been farmore intensively studied than the other two countries,only a few areas are well known. The Sierra de Guatemalain Tamaulipas, the Sierra de EI Abra in San LuisPotosi and Tamaulipas, and the Yucatan Peninsulaare the only regions in Mexico where a significantpercentage of the fauna is known. Due to the outstandingwork of the Italian biospeleologists much ofChiapas is now well collected, but a majority of theirmaterial remains unstudied. The cavernicole fauna ofwestern, southwestern, and parts of southern Mexicoremain completely unknown. Of the 13 physiographicprovinces (see Fig. 1) into which this regionmay be divided, no collections other than hats or hatassociatedparasites have been made in Baja California,the Buried Ranges, the Central Mesa, or the PacificCoastal Lowlands. The cavernicole fauna of the2

Eric Remington, William Rhodes, Jim Rodemaker,Francis Rose, J. Mark Rowland, Carol Russell, PhillipRussell, Fred Sawin, Terry Sayther, Michael Shawcross,Jim Shepperd, Rex Shepperd, Tommy Shifflett,Mark Shumate, A. Richard Smith, Jim Smith,Richard M. Smith, Carmen Soileau, Reynaldo Solis,Bill Steele, Bill Stone, Peter Strickland, Pierre Strinati,Joe Sumbera, David Sustare, Ron Syme, MillsTandy, Virginia Tipton, William Tozer, Cindy Tracy,Gary Tucker, Merydith Turner, 1. Turpin, GregWalker, Mike Wharton, Jack White, Lisa Wilk, DanWitter, and Steve Zeman. To those whose names havebeen inadvertently omitted I offer my apologies.Numerous systematists have assisted with identificationand description of the material collected fromthe caves of this area. I thank all of the following fortheir kindness in supplying this and much other informationon the cave and related epigean fauna: R. K.Allen, mayflies; D. M. Anderson, beetles; E. W.Baker, mites; T. C. Barr, Jr., beetles; S. W. T. Batra,hymenopterans; R. W. Baumann, embiopterans, mayflies,stoneflies; T. E. Bowman, mysids, isopods; A.Brindle, earwigs; H. R. Burke, beetles; G. W. Byers,flies; M. J. Campbell, beetles: R. W. Carlson, hymenopterans;O. 1. Cartwright, beetles; N. B. Causey,millipeds; F. E. Chace, shrimps; E. A. Chapin, beetles;K. Christiansen, collembolans; A. C. Cole, ants; D. R.Davis, moths; Denis Delorme, ostracods; W. E. Duellman,frogs, salamanders, lizards; Terry Erwin, beetles;R. G. Fennah, homopterans; D. C. Ferguson, moths;O. S. Flint, mayflies, dragonflies, neuropterans, trichopterans;O. F. Francke, scorpions; R. C. Froeschner,hemipterans; R. J. Gagne, flies; G. E. Gates,earthworms; W. J. Gertsch, sCOTpions, spiders, amblypygids,schizomids, ricinuleids; R. Gonzalez R.,diplurans; C. J. and M. 1. Goodnight, opilionids;A. B. Gurney, earwigs, roaches; T. F. Halstead, hemipterans;1. H. Herman, beetles; .J. 1. Herring,hemipterans; H. H. Hobbs, ]r., crayfish, shrimps, andentocytherid ostracods; H. H. Hobbs, III, shrimps,entocytherid ostracods; J. R. Holsinger, amphipods;H. F. Howden, beetles; T. H. Hubbell, crickets; C.Hubbs, fish; Leslie Hubricht, snails; P. D. Hurd, Jr.,bees; W. 1. Jellison, fleas; M. Kawakatsu, planarians;J. E. Keirans, ticks; J. M. Kingsolver, beetles; 1. V.Knutson, flies; G. M. Kohls, ticks; J. P. Kramer,homopterans; R. E. Lewis, fleas; J. D. Lynch, frogs;T. C. Maa, flies; P. M. Marsh, hymenopterans; A. S.Menke, hymenopterans; R. W. Mitchell, planarians,scorpions, fish; E. 1. Mockford, psocids; W. B. Muchmore,pseudoscorpions; C. Mullinex, amblypygids;C. Nelson, lizards; R. Newcomer, salamanders; S. B.Peck, beetles; W. 1. Peters, mayflies; G. B. Rabb,salamanders, D. A. Rossman, snakes, lizards; J. M.Rowland, amblypygids, schizomids; C. W. Sabrosky,flies; G. A. Schultz, isopods; W. A. Shear, millipeds;A. E. Smalley, crabs; R. 1. Smiley, mites; D. R.Smith, termites, ants; M. Soleglad, scorpions; A.Solem, snails; P. J. Spangler, beetles; T. J. Spilman,beetles; G. C. Steyskal, flies; M. B. Stoetzel, homopterans;A. Stone, flies; R. W. Strandtmann, mites;W. R. Suter, beetles; S. Szerlip, hemipterans; M.Tandy, fish; F. G. Thompson, snails; W. Voss, mites;F. W. Wagner, scorpions; D. Wake, salamanders; L. M.Walkley, hymenopterans; R. E. Warner, beetles; A.Weaver, centipeds; J. P. Webb, mites; D. M. Weisman,moths; R. 1. Wenzel, beetles, flies; R. White, beetles;W. W. Wirth, flies; D. 1. Wray, collembolans; P.Wygodzinsky, diplurans, thysanurans; H. C. Yeatman,copepods; C. E. Yunker, mites.I thank the following for their reading of all orpart of the manuscript and for making many usefulsuggestions: T. C. Barr, Jr., T. E. Bowman, N. B.Causey, K. Christiansen, W. R. Elliott, F. W. Fisk,O. F. Francke, R. C. Froeschner, G. E. Gates, W. J.Gertsch, C. J. and M. 1. Goodnight, H. H. Hobbs, Jr.,H. H. Hobbs, III, J. R. Holsinger, 1. Hubricht, E. 1.Mockford, W. B. Muchmore, S. B. Peck, J. M. Rowland,W. H. Russell, G. A. Schultz, W. A. Shear, A. R.Smith, H. M. Smith, T. J. Spilman, j. Sullivan, F. W.Wagner, A. Weaver, P. Wygodzinsky, and H. C. Yeatman.I take full responsibility for all errors and forall interpretations of the data.Jerry Atkinson, David McKenzie, William H.Russell, and Peter Sprouse assisted in the preparationof the locality list. William R. Elliott and Hal Storyprovided invaluable aid in the preparation of themaps.Financial support for some of the field work inMexico was provided by the National GeographicSociety and The Museum, Texas Tech University.Additional assistance was provided by Nell B. Causey,Willis J. Gertsch, and Theodore H. Hubbell.HISTORYMexicoThe first biological investigation of a cave inMexico was conducted by Dominik Bilimek on January14, 1866, when he visited Grutas de Cacahuamilpa,Guerrero (Bilimek, 1867). Among the II speciesreported was one troglobite, Lepisma anophthalma.The next important contribution to ourknowledge of the cavernicole fauna of Mexico wasthat of the distinguished Mexican zoologist, D. A. 1.Herrera (1891). Among species he collected in Grutasde Cacahuamilpa was the same species of troglobite,described as Lepisma cacahuamilpensis. Herrera also5

collected a troglobitic cirolanid isopod from a well inMonterrey, Nuevo Leon, which was described byA. S. Packard (1900). With the exception of a fewscattered records of bats, birds, fishes, and snailsthese are the only publications on Mexican cavefauna prior to 1936.In the summer of 1932 Edwin P. Creaser, F. G.Hall, and A. S. Pearse investigated 35 cenotes andcaves in the state of Yucatan. The volume resultingfrom their studies, published in 1936, has become aclassic in biospeleologicalliterature. Although emphasizingthe taxonomy of the collected fauna, this workincluded one of the first studies ever made on thephysical and chemical nature of karst waters in NorthAmerica (Hall, 1936). Pearse's (1936d) observationson the ecology of cenotes and caves are of someinterest to the student of the evolution of cave faunas.Although about 70 species were reported fromcaves and cenotes in the state of Yucatan, only fourcrustaceans were troglobites. Creaser (1936) describednew species of isopod, mysid, and shrimp.In the summer of 1936 A. S. Pearse continued hisstudies of the subterranean fauna of Mexico with theexploration of 27 caves in various parts of the state ofYucatan. The results of this expedition were collectedin a volume published in 1938 by the Carnegie Institutionof Washinbrl:on. A total of about 300 animalspecies were identified, of which many were newspecies. Ten species of troglobite were added to thecave fauna of Yucatan, of which eight were terrestrial.Two spiders (Chamberlin and Ivie, 1938b), oneisopod (Creaser, 1938), one milliped (Chamberlin,1938), three collembolans (Mills, 1938), one cricket(Hubbell, 1938), and two fish (Hubbs, 1938) wereconsidered to be troglobites.In 1936 Salvador Coronado discovered in CuevaChica, San Luis Potosi, the first known species ofeyeless characin. It was described by C. 1. Hubbs andW. T. Innes (1936) as Anoptichthys jordani. The easewith which this species could be maintained in aquariainspired an expedition in March 1940 by the NewYork Aquarium. The members of the party includedC. M. Breder, Jr., E. B. Gresser, M. B. Bishop, WilliamBridges, S. C. Dunton, and Salvador Coronado. Athorough collection was made in the cave of both fishand invertebrates, and the initial results were publishedby Bridges (1940) and Breder (1942). Theunique situation in which this species was found, itspresence in vast numbers, the ease with which it couldbe raised in the laboratory, and its ability to interbreedwith its surface ancestor Astyanax mexicanus(Filippi) have led to its study by numerous Americanand European workers over the past 43 years. The descriptionof two additional species, AnoptichthysantTobius Alvarez (1946) and Anoptichthys hubbsiAlvarez (1947), inspired still further study of theseremarkable fish. In the 38 years since the originaldescription of A. jordani more than 200 publicationshave been more or less devoted to the study of thespecies.In 11)38 two distinguished Spanish biospeleologists,Federico Bonet and Candido Bolivar y Pieltain,emigrated to Mexico and began an intensive study ofits cave fauna. Their work during the next 20 yearsgreatly expanded our knowledge of the cave biologyof Mexico.Their investigations began with a visit to Grutas deCacahuamilpa on December 13-16, 1939. This expedition,conducted in conjunction with A. Dampf andD. Pelaez, led to the discovery of many of the samespecies reported by Bilimek and of several new species(Bolivar, 1940). A second visit was made onJune 13, 1940, by D. Pelaez.During December 1939 and January 1940, lIortonII. Hobbs, Jr., A. F. Carr, Jr., and others visited acave near Hacienda Potrero Viejo, Veracruz, resultingin the discovery of the first blind crayfish knownfrom Mexico (Hobbs, 1943). On May 7,1940, Ivan T.Sanderson collected ricinuleids in Actun Kaua, Yucatan(Sanderson, 1941). During an expedition for thepurpose of collecting insects and arachnids of medicalimportance, Harry Hoogstraal visited several caves inNuevo Leon. This included a trip on June 16, 1940,to Cueva de la Boca and the discovery of the firsttroglobitic centiped known from Mexico (Chamberlin,1941).In 1941 three trips of some importance were madeby F. Bonet and C. Bolivar y Pieltain. On January 16a visit to the large, Olmec cave, Grutas de Jllxtlahuaca,Guerrero, led to the discovery o[ several troglubites.Trips on May 30 and November 11-13 to Grutasde Atoyac, Veracruz, also resulted in the discoveryof several troglobites.In April 1942 F. Bonet, C. Bolivar y Pieltain, B. F.Osorio Ta[all, D. Pelaez, F. Cardenas, M. Correa, andJ. Alvarez visited Cueva Chica and Cueva de Los Sabinos,San Luis PotOSI (Anonymous, 11)42a). Amongthe more remarkable discoveries in the latter cavewere new species of troglobitic ricinuleid (Bolivar,1946), isopod (Bolivar, 1950), fish (Alvarez, 1947),and phalangid (Goodnight and Goodnight, 1942).Cueva de Los Sabinos was revisited July 19 by C.Bolivar y Pieltain, B. F. Osorio Tafall, and M. cardenas.These two trips resulted in one of the more significantpublications on the aquatic fauna of Mexicancaves (Osorio Tafall, 1943).The research by members of the Escuela _ acionalde Ciencias Biol6gicas de Mexico was expanded into6

Nuevo Leon when, on July 13-19, 1942, C. Bolivar yPieltain, F. Bonet, B. F. Osorio Tafall, D. Pelaez, andothers visited Cueva de la Boca, Grutas de Villa deGarcia, Cueva del Carrizal, Grutas del Palmito, andother caves. A second trip to this area was madeSeptember 15-17, 1942, by C. Bolivar y Pieltain(Anonymous, 1942b). Collected on these two expeditionswere new species of troglobitic dipluran (Wygodzinsky,1944), collembolan (Bonet, 1943), phalangid(Goodnight and Goodnight, 1944), pseudoscorpion(Chamberlin, 1946), and beetle (Bolivar,1942).In September 1943 B. F. Osorio Tafall visited Yucatan.His collections in Actun Sabaca and Grutas deBalankanche resulted in the discovery of two speciesof pseudoscorpion representative of a new family,the Vachoniidae (Chamberlin, 1947). In November ofthe same year, C. Bolivar y Pieltain and B. F. OsorioTafall visited Grutas de Acuitlapan, Guerrero.During 1944 three of Mexico's better known caveswere revisited. On April 30 l Alvarez and C. Tellezvisited Grutas de Atoyac, Veracruz; on May 21 C.Bolivar y Pieltain, F. Bonet, l Alvarez, and C. Tellezvisited Cueva de Los Sabinos, San Luis Potosi; and onJune 17 F. Bonet visited Cueva de la Boca, NuevoLeon.In April 1944 Dr. and l\Jrs. M. W. Stirling of theBureau of American Ethnology of the SmithsonianInstitution and Mr. R. H. Stewart of the NationalGeographic Society visited Cueva del Azufre, Tabasco,and collected specimens of blind fish, now identifiedas Poecilia sphenops.In December 1945 F. Bonet, C. Bolivar y Pieltain,J. Alvarez, and other members-of the Escuela Nacionalde Ciencias Biologicas de Mexico visited Cueva deEl Pachon, Cueva del Abra, and Cueva de los Cuartelesin Tamaulipas and Cueva de El Jobo in San LuisPotosi. The most notable discovery of this expeditionwas the third population of blind fish of the genusAnoptichthys (described as A. antrobius by Alvarezin 1946).In April 1946 B. l Dontzin and E. Ruda revisitedCueva del Azufre, Tabasco, and collected a large seriesof blind and eyed Poecilia sphenops.B. F. Osorio Tafall and M. Cardenas Figueroavisited several caves in Yucatan as part of an expeditionsponsored in March 1947 by the Secretaria deRecursos Hidraulicos (Anonymous, 1947). Their discoveriesincluded specimens of the rare blind fishTyphliasina pearsei in Cenote del Pochote (Solorzano,1953). The infonnation gathered on this expeditionserved as material for an excellent study byCardenas Figueroa (1950) on the hydrobiology ofYucatan.During the course of the David Rockefeller MexicanExpedition of the American Museum of NaturalHistory in the summer of 1947, W. J. Gertsch visitedCueva del Diablo, Chihuahua, and collected an undescribedspecies of troglobitic Psilochorus (Spieth,1950).While conducting studies on the distribution andtaxonomy of opilionids during the summer of 1948,C. l Goodnight visited Grutas del Cocona, Tabasco,and several caves in Yucatan. Among many interestingspecies of arachnid collected were two eyelessspecies of amblypygid, not reported until they weredescribed by Rowland (1973d).In November 1950 Alejandro Villalobos of theInstituto de Biologia de Mexico began a series ofinvestigations into the cavernicole fauna of Mexicowhich was to result in the description of many newspecies of troglobite of great interest. His first expeditionwas to several caves in the vicinity of Comitanand Tuxtla Gutierrez, Chiapas. His visit to Cueva delTio Ticho resulted in the discovery of one of themore remarkable cavernicoles known from Mexico,an eyeless crab described by Enrique Rioja (1953h)as Typhlopseudothelphusa mocinol.In November and December 1950 F. Bonet andothers began the serious study of one of Mexico'sgreatest karst regions, that of Xilitla, San Luis Potosi.This study of the Xilitla region was continuedin January 1952 by F. Bonet in conjunction withR. Ortiz, l V. Flores, M. Camacho, and A. Gonzalez.In addition to collecting extensively they prepareddetailed maps and descriptions of the caves theyvisited (Bonet, 1953a).On November Il, 1951, C. Bolivar y Pieltain andAna Maria Bolivar visited Grutas de Quintero, Tamaulipas.This collection included new species ofcirolanid isopod (Rioja, 1953d) and pseudoscorpion(Beier, 1956).In Decmber 1951 and May 1952 A. Villalobosvisited Cueva del Ojo de Agua Grande, Veracruz. Thefauna discovered included a remarkable species oftroglobitic isopod (Rioja, 1953c).In January 1953 A. Villalobos continued his investigationsof the cavernicole fauna of Mexico witha visit to Cenote de Sambula at Motul, Yucatan.Among the fauna collected was a new species of trichonscidisopod (Rioja, 1958). In July C. J. Goodnightcollected arachnids, including a new troglobiticspider, in Grutas de Atoyac, Veracruz.The first study of the cavemicole fauna of Oaxacawas made in May 1954 by C. Bolivar y Pieltain, lCarranza, and others. A visit to Grutas de Monteflorresulted in the collection of a new species of troglobiticpseudoscorpion (Beier, 1956). In August 19547

C. Bolivar and J. Carranza investigated wells andmines in the vicinity of Musquiz, Coahuila. The mostsignificant discovery made was of the first blind catfishin Mexico (Bolivar and Carranza, 1954); thisspecies was described as Prietella phreatoph'ila byCarranza (1954).In December 1955 Malcolm S. Gordon, R. Deering,and J. Till made the third trip to Cueva delAzufre, Tabasco, and once again collected a series ofPoecilia sphenops.In July 1956 W. J. Gertsch and Vincent Roth revisitedCueva del Diablo, Chihuahua, and visitedCueva de EI Ocote, Hidalgo. In the latter cave theycollected specimens of new species of spider (Gertsch,1971a) and carabid beetle (Hendrichs and Bolivar,1966).Grutas del Cocona, Tabasco, was visited in 1958by A. Villalobos, where he discovered the first troglobiticamphipod known from Mexico and reported thepresence of a troglobitic planarian (Villalobos, 1960).The planarian was later rediscovered and described byMitchell and Kawakatsu (1972) as a new family, genus,and species (Dimarcusidae, Dimarcus villalobosi).On February 15 and July 27, 1958, Grutas del Mogote,Guerrero, was visited by Raymond de Saussure,at which time he collected specimens of a troglobiticrhachodesmid milliped (Causey, 1973).On March 20, 1959, Stanley Kiem visited a cavenear the Hacienda San Bernardo, Yucatan, nowidentified as Actun Xpukil. He discovered a speciesof troglobitic milliped which was described by Loomis(1962) as Orthoporus kiemi. A large series ofliving and preserved troglobitic Poecilia sphenopswas collected in Cueva del Azufre, Tabasco, in March1959 by A. G. Dinkins.Visits were made by A. Villalobos in April andSeptember 1960 to Grutas del Cocona, Tabasco. Theresults of these trips remain largely unpublished.In 1961 the distinguished American ichthyologist,Carl L. Hubbs, visited the marshes and springs in thevicinity of Cuatro Cienegas de Carranza, Coahuila.Among collections made in the Pozos de la Becerraon April 6 were specimens of the troglobitic snailCoahuilix hubbsi, described by Taylor in 1966.In November 1962 C. Bolivar y Pieltain and A.Urquijoi visited Grutas de Acuitlapan, Guerrerro.Their collections included the troglobitic pseudoscorpionAlbiorix bolivari Beier (1963).With the organization in 1962 of the SpeleologicalSurvey of Mexico in Austin, Texas, a new era in thestudy of the cavernicole fauna of Mexico was initiated.In November 1962 the first expedition of thenewly organized group (now called the Associationfor Mexican Cave Studies) was undertaken. A fewsmall collections in caves were made by the fourmembers of this expedition, T. R. Evans, T. W.Raines, ]. R. Reddell, and W. H. Russell. This materialincluded specimens of new genera and species ofisopod from Sotano de Huitzmolotitla, San LuisPotosi (Schultz, 1964), and milliped from Sotano delArroyo, San Luis Potosi (Causey, 1963). Because ofthe interest of this initial small collection, an activeprogram of collection and study of the cavernicolefauna of Mexico was initiated and continued to thepresent. It is impossible to recount in detail here thehistory of the work of the Association for MexicanCave Studies during the last 17 years. More than1000 species have been collected in more than 500caves during this period. The following account willemphasize only the more distinctive discoveries ofthis period.On March 2, 1963, W. H. Russell visited Cueva delCarrizal, Nuevo Leon, collecting among other speciesa new troglobitic hyid pseudoscorpion (Muchmore,1972a). On October 26 of the same year he also visitedCueva de la Porra near Xilitla, San Luis Potosi.This collection included specimens of a new genusand species of cambalid milliped (Causey, 1964a).The activities of the Association for Mexican CaveStudies increased in 1964 with explorations in thekarst areas of Tequila, Veracruz; Sierra de Guatemala,Tamaulipas; and Xilitla, San Luis Potosi. On June 2Terry Raines and William Bell entered a deep pit nearTequila, Sotano del Profesor, which was notorious forhaving been the disposal site for the body of a murderedschool teacher. Among the fauna collectedabout the remains of the unfortunate teacher was anew genus and species of carabid beetle (Barr, 1965).Also in June Larry Manire, David McKenzie, andJames Reddell visited what was to prove to be one ofthe more notable faunal areas in North America, theSierra de Guatemala, Tamaulipas. Among numerousspecies of troglobite were new species of gryllid(Hubbell, 1972), glomeroid milliped (Causey, 1964b),rhachodesmid milliped (Causey, 1971b), and pseudoscorpion(Muchmore, 1972a). In November explorationby Terry Raines and William Bell in one of thedeeper caves in North America, Sotano de Tlamaya,at Xilitla, resulted in the discovery of a new speciesof cavernicole earthworm (Gates, 1967). In additionto these visits to distinct karst regions, a trip by W. H.Russell to Cueva de los Riscos in Durango led to thediscovery of a troglobitic ricinuleid (Gertsch, 1971b)and spiders (Gertsch, 1971a).In January and June of 1965 David McKenzie,Orion Knox, and others continued work in the Sierrade Guatemala, most notably with the mapping andexploration of Sotano de la Joya de Salas. Two spe-8

cies of carabid beetle of great interest were discovered(Barr, 1966a; 1967a). On April 8. 1965, Jorge Urquijoand Antonio Guerrero revisited Sotano del Profesorat Tequila, Veracruz, and obtained additional specimensof the blind carabid Mexisphodrus veraecrucis.In July 1965 William Russell led a reconnaissancetrip to Huautla de Jimenez, Oaxaca. This great karstregion now is known to include some of the deeperknown caves in the Western Hemisphere. Many newspecies were obtained on the trip, but most of theresults remain unpublished. During the same monthJohn Fish, Terry Raines, and James Reddell conducteda reconnaissance trip to several caves in northcentral Mexico, including caves in Coahuila, Chihuahua,and Durango. In August William Bell, John Fish,and James Reddell explored caves in Hidalgo, Guerrero,and Veracruz; among the fauna collected werenew species of troglobitic spider beetle (Spilman,1968) and rhachodesmid milliped (Causey, 1973).In February 1966 William Bell and James Reddellexplored several caves in Coahuila, Chihuahua, andDurango. The most notable find was a second speciesof relict ricinuleid from western Mexico, in Cueva delGuano, Durango (Gertsch, 1971b). In June severalgypsum caves near Galeana, Nuevo Leon, were visitedby David McKenzie, Orion Knox, A. Richard Smith,and James Reddell. New species of catopid (Peck,1973c) and blind spider (Gertsch, 1971a) were discoveredin Resumidero del Pablillo.In August 1966 Francis Abernethy, Robert W.Mitchell, and William Rhodes visited several caves inthe Sierra de Guatemala, Tamaulipas. While exploringCueva de la Mina, Rhodes discovered the first eyelessscorpion in the world. It was' described by Mitchell(1968) as Typhlochactas rhodesi. Also in AugustJohn Fish, David McKenzie, James Reddell, andRichard M. Smith made the first biological collectionsin the gypsum karst area of Matehuala, San Luis Potosi;the karst region of Valle de los Fantasmas, SanLuis Potosi; and the karst region of Pinal de Amoles,Queretaro. In Sotano de Tejamanil, Queretaro, theydiscovered two species of eyeless trechine beetle(Barr, 1967d). In November John Fish and JonathanDavis discovered a third new species of trechine in asecond visit to Valle de los Fantasmas (Barr, 1967d).In December Terry Raines led a trip to the Caftan deHuasteca in Nuevo Leon. He discovered in Grutas deSan Bartolo a new species of troglobitic schizomid(Rowland, 1973a).In June 1967 Francis Abernethy, Robert Mitchell,James Reddell, and Pierre Strinati visited several cavesin the Sierra de Guatemala and Sierra de EI Abra.This resulted in the collection of new species ofcollembolans (Christiansen, 1973), as well as supplementingprevious collections. During the same yearcollections by Robert Mitchell, James Reddell, SuzanneWiley, and other members of the Departmer.tof Biology of Texas Tech University in Lubbock,Texas, were made in numerous caves in these twokarst regions. Discoveries included many new populationsof troglobite and troglophile, including blindfish of the genus Astyanax.In July 1967 John Fish, James Reddell, and PhilipRussell explored caves near Pinal de Amoles, Queretaro;Ciudad de Maiz and Xilitla, San Luis Potosi; andCiudad Victoria, Tamaulipas. Among many troglobitescollected were specimens of the first species ofblind theraphosid tarantula (Gertsch, 1971a) and newspecies of rhaphidophorid and gryllid crickets (Hubbell,1972).T. R. Evans, John Fish, James Reddell, and MillsTandy in August 1967 visited caves near Cordoba andTequila, Veracruz; Huautla de Jimenez, Oaxaca; andat several localities in Chiapas. Numerous troglobitesof considerable interest were obtained, but probablythe most singular discovery was of a second species oftroglobitic scorpion in Cueva del Ojo de Agua de Tlilapan,Veracruz. This species was described by Mitchell(1968) as Typhlochactas reddelli. Other species ofinterest included additional specimens of the blindcrab, Typhlopseudothelphusa mocinoi, in Cueva delTio Ticho, Chiapas; a new species of troglobitic collembolannear Huautla de Jimenez, Oaxaca (Christiansen,1973); and a new species of cavernicolousricinuleid from Chiapas (Gertsch, 1971b). In SeptemberJohn Fish and William Russell made the firstcollections in the caves of the karst region of Aquisman,San Luis Potosi.Collections in 1968 were made primarily in theSierra de Guatemala and Sierra de El Abra by R. W.Mitchell, James Reddell, and their associates at TexasTech University. Cueva de la Capilla, near El Porvenirin the Sierra de Guatemala, was visited for the firsttime on January 28 by John George, R. W. Mitchell,James Reddell, and Francis Rose. Discovered in thiscave were new species of cave-adapted earthworm(Gates, 1971), spider (Gertsch, 1971a), phalangid(Goodnight and Goodnight, 1971), catopid beetle(peck, 1968), and trechine beetle (Barr, 1971).Several caves in the Valle de los Fantasmas regionof San Luis Potosi were visited in November 1968by William Elliott, Jimmy Jarl, and members of theSouthwest Texas Grotto at San Marcos, Texas.Among the fauna collected was a new genus andspecies of troglobitic milliped (Causey, 1969).Between 1966 and 1968 numerous collectionswere made by G. A. Cole, W. 1. Minckley, D. W.Taylor, J. J. Landye, and others in the pozos and9

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 forms 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, for 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 for 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 theAssociation for Mexican Cave Studies 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 Association forMexican Cave Studies 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 Mexicanoconducted 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. Caveswere 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).Caves 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 Association for Mexican CaveStudies was that made from December 22, 1972, toJanuary 8, 1973, by David McKenzie, Martha HelenMcKenzie, Stuart Murphy, and James Reddell (Reddell,1973e). Caves 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 Association for MexicanCave Studies 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

Cueva del Diablo near Orizaba, Veracruz; it remainsundescribed. Collections were made in one cave inCampeche and in about 40 in Yucatan. Attemptswere made to revisit as many of the caves studied byPearse in 1936 as was possible; numerous new cavesof great interest were also located. In addition tomore specimens of all of the troglobites previouslyreported and many new species of troglophile, collectionsincluded new blind spiders (Gertsch 1977b),shrimp (Hobbs and Hobbs, 1976), pseudoscorpion(Muchmore, 1977), and amphipod (Holsinger, 1977).Of particular interest were additional populations ofthe rare blind fish Typhliasina pearsei and Ophisternoninfernale.During July 1973 a visit to the Yucatan Peninsulawas made by Dr. Masaharu Kawakatsu, Robert W.Mite:hell, James Reddell, J. Mark Rowland, andothers. Collections were made in many caves and epigeanlocalities in the state of Yucatan.The Third Zoological 1ission to !\lexico sponsoredby the Accademia Nazionale dei Lincei wasconducted from July 20 to October 11, 1973, hyValerio Sbordoni, Roberto Argano, Aldo Zullini,and Vincenzo Vomero. A total of 45 caves werevisited in several parts of Chiapas and adjacentHudlllrtenango, Guatemala. Among the morenotable rliscoveries of this expedition were newspecies of troglobitic stenasellid isopod (Argano,1(77), amphipod (Ruffo and Vigna TagJianti, ] 977),palat:monid shrimp (Holthuis, 1977), opilionid(Silhavy, 1977), and milliped (Shear, 1977a).In AUhrust 1973 David McKenzie, Roy .I arneson,and others made collections in several caVf:S in theRancho Nuevo region of :'-Juevo Leon and Tamaulipas.Among species of intnest were new, still undescribedspecies of trechine hectic and thr secondspecies of troglohitic catopid known from Mexico(Peck, 1977).In December 1973 Peter Sprouse, Peter Strickland,and others visited several cavrs ill the A1luismon regionof San Luis PotosI. Additional populations ofspveral troglobites and a stillundescrihed new speciesof t>nrlemic earthworm were found.From December 21, 197:1, to January 12, 1974,William Elliott, Bob Harr, Roy Jameson, DavirlMcKenzie, and James Reddell conductt:d biologicalinvestigations in the caves of the ZacapoaxtlaCuetzalan area of Puebla, the Aeatlan region ofOaxaca, and the Soledad At7.0mpa anrl Cofre: de:Perote areas of Veracruz. Each area produced spl':riesof interest. Among many species discovered in theZacapoaxtla-Cuetzalan region were new crayfish(Hobbs, 1975), spiders, and millipeds (Causey,1975b). A return visit to Cueva del Nacimiento delRio San Antonio in Oaxaca resulted in the collectionof a new species of troglobitic cyphophthalmidopiliollid (Shear, 1977b), the first record for thissuborder in :\Iexico. Collections in the SoledadAtzornpa region included specimens of several raretroglobites also known from the Tequila regionbut also new species of spider and milliped. Finally,study of Cueva del Volcancillo, a lava tube on Cofrede Perote, was the first investigation of the biologyof a lava tube in :'.Iexico. Although not a cave wellsuitedfor fauna, Cueva del Volcancillo possessedmany species of interest, including troglohitic opilionids,millipeds, and trechine beetles. 1\']ost of theresults of this expedition remain unpublished.The caves of the Micos region and the Sierra deEl Ahra, San Luis Potosi, were biologically studiedfrom May 15 to June 8, 1974, by William Elliott,John Prentice, Carmen Soileau, and others. Theprimary emphasis was on obtaining specimens andother information on Astyanax jordani.The members of the second expedition were AndyGrubbs, David McKenzie, James Reddell, and SuzanneWiley; Robert W. Mitchell, William Russell, andothers joined the expedition for part of the time.Their studies included the biological investigation offour caves in Tahasco, eight caves in Campeche, 14caves in Quintana Roo, and 13 caves in Yucatan. Inaddition to further delineating the ranges of manytroglobites, the expedition resulted in the discoveryof new species of blind spiders, amblypygid, andmilliped (Grubbs, 1975).The fourth expedition sponsored hy the AccademiaNar,ionale dei Lincei was conducted from July 30to October 20, 1975. The personnel on this expeditionwere Roberto Argano, Valerio Shordoni, VincenzoVomero, and Aldo Zullini. Thirty caves werevisited in Chiapas; studies of the phreatic biotopewere also made in Chiapas, Campeche, Quintana Roo,and Yucatan. Among the more significant discoveriesof this expedition were new species of troglobitieasellid isopod, am phipod, crab, and trechine beetle(Sbordoni et aI., 1977).In August 1975 Andy Grubbs, David \lcKenzie,and Suzanne Wiley made the first biological study ofcaves in the karst region of San Nicolas de los Montes,San Luis Potosi. Thpir most notable discovery was ofa new gfnus and species of troglohitic chactid scorpIOn.An ex pfdition in December 1973 hy DennisBarnes, Tom Byrd, l\lareia Cossey, Andy Grubb;;,Shari Larason, and Terry Sayther to the Acatlanregion, Oaxaca, included biological collfction;; inseveral caves. Their discoveries included a new speciesof blind diplorentrid scorpion, described by Francke12

(1979) as Diplocentrus cueva. A new species of troglobitichomopteran was collected in this same monthby Mike McEachern in Cueva de Cayetano, Oaxaca.Further studies were made in December 1975 andJanuary 1976 by Dennis Barnes, Tom Byrd, MarciaCossey, Andy Grubbs, and Jim Rodemaker in thecaves of the San Nicolas de los Montes region, SanLuis Potosi.December 1975 to January 1976 Peter Sprouse,Peter Strickland, and Carmen Soileau made collectionsin caves in the Sierra el Pino, San Luis Potosiand near Pinalito, Hidalgo.Biological investigations were made of severalcaves in the San Juan area, Queretaro, in January1976, by Roy Jameson and Patty Mothes, and inMarch 1976 by Alexia Cochrane, Gill Ediger, AndyGrubbs, and Roy Jameson.June 8-15, 1976, Roy Jameson and Patty Mothescollected new species of blind planaria, tarantula,and milliped in caves northwest of Xilitla in Queretaro(Jameson, 1977).In November 1976 the Rio Purificacion area ofTamaulipas was the object of intensive study. Collectionsby Andy Grubbs and Peter Sprouse includednew species of planarian, opilionid, and trechinebeetle.Collections in the Huautla de Jimenez region,Oaxaca, December 1976 to January 1977 by RoyJameson and Patty Mothes included blind millipeds,opilionids, carabid beetles, and a new genus andspecies of eyeless scorpion of unknown familialaffinities.An expedition from December 15, 1976, toJanuary 15,1977, by Andy Grubbs, David McKenzie,James Reddell, and Carmen Soileau included visits tothe Sierra de Tamaulipas, Tamaulipas; the Cuetzalanregion, Puebla; the Acatliin region, Oaxaca; and theJalapa, Cuitlahuac, and Atoyac regions, Veracruz. Atotal of 38 caves were studied; collections includednew species of mysid (Bowman, 1977b), crayfish,shrimp, spider, milliped, and homopteran.The first visit to the karst region of Zoquitliin,Puebla, was made by Preston Forsythe, Jim Rodemaker,Peter Strickland, and others in January 1977.Their collections included a new genus and speciesof snail and blind opilionids, millipeds, and carabidbeetles.In January 1977 Robert W. Mitchell and LindaFaulkenberry collected, among other species ofinterest, a new species of cirolanid isopod in the cavesof the San Nicolas de los Montes region, San LuisPotosi.Collections were made by Carmen Soileau inMarch 1977 in caves in the vicinity of Yochib, Chiapas.The study of the Rio Purificacion region, Tamaulipas,intensified in 1977 with expeditions in March,May, November, and December. The principal collectorsin this area were Gill Ediger, David McKenzie,Dale Pate, and Peter Sprouse. Their collections includedtroglobitic stenasellid and trichoniscid isopods,spiders, opilionids, centipeds, and carabidbeetles.During an expedition in April-May 1977 to Huautlade Jimenez, Oaxaca, Tracy Johnson and Bill Steelecollected blind millipeds, opilionids, and carabidbeetles in several caves.In May 1977 Roy Jameson and Patty Mothes continuedtheir study of the caves in the region northwestof Xilitla in Queretaro. Their collections includedblind planaria, isopods, spiders, opilionids,and millipeds.Roy Jameson and Patty Mothes in November andDecember 1977 made the first study of the caves ofthe San Joaquin, Queretaro, region (Mothes, 1978).Among their collections were new species of blindmilliped.December 1977 to January 1978 Jean Jancewicz,Roy Jameson, Dino Lowery, and Steve Zeman collectedtroglobitic amblypygids, spiders, millipeds,and carabid beetles in several caves in the Huautla deJimenez region of Oaxaca.In January 1978 Jerry Atkinson, Preston Forsythe,Jocie Hooper, and Peter Strickland revisited theZoquitliin, Puebla, area (Atkinson, 1978). Theircollections included a new species of blind nesticidspider (Gertsch, in press).Terry Sayther in January 1978 collected blind isopodsand other fauna of interest in several caves inthe Atoyac region, Veracruz.The caves of the Purificacion region of Tamaulipaswere the object of intensive study during 1978, withthe primary emphasis being placed on the SistemaPurificacion, Cueva X, Sumidero de Oyamel, andCueva de los Allarines. The principal collectors duringthis period were William Elliott, Andy Grubbs, DavidMcKenzie, Dale Pate, Terry Raines, Peter Sprouse,Terri Treacy, and Lisa Wilk. Among the more notablefinds were undescribed species of troglobitic planaria,isopod, pseudoscorpion, opilionid, milliped, andcentiped.An expedition to the Huautla de Jimenez, Oaxaca,region in April and May 1978 by Bill Steele, BillStone, Andy Grubbs, Mike McEachern, Jill Dorman,Jim Smith, and others, resulted in the collection oftroglobitic scorpions, opilionids, millipeds, and carabidbeetles.13

Those species of troglobite which have been describedfrom Mexico are listed in Table 2 in thechronological order of their description.BelizeThe cave fauna of Belize has been little studiedand most collections have been made in caves nearAugustine and Caves Branch in Cayo District.Paul Williams studied the phlebotomine sandfliesof four caves near Augustine and one near Millionariobetween January 1964 and May 1969 (Williams,1976c).Marie and Charles Goodnight collected, amongother species, opilionids in one of the Rio Frio caves,Cayo District, onJuly 1, 1971 (Goodnight and Goodnight,1977).A small collection of invertebrates was made byDavid McKenzie on January 19, 1972, in St. Augus.tine Cave, Cayo District.Most of our knowledge of the cave fauna of Belizeis the result of a study by Sand J. Peck during Julyand August 1972. They visited several caves nearAugustine and Caves Branch and obtained severalspecies of unusual interest. Among their collectionswere new species of troglobitic spider (Gertsch,1973b), pseudoscorpion (Muchmore, 1973a), opilionid(Goodnight and Goodnight, 1977), and milliped(Shear, 1973).On July 20, 1972, Charles Goodnight visited acave near Augustine and collected a new species oftroglobitic pseudoscorpion (Muchmore, 1973b).During July and August 1976 Logan McNatt, TomMiller, and Michael Shawcross studied several cavesnear Caves Branch, Cayo District. Most of these collectionsremain unstudied, but a troglobitic opilionidwas described by Goodnight and Goodnight (1977).Other specimens of interest included undescribedspecies of troglobitic milliped and charontid amblypygid.Logan McNatt made additional collections inthis same region in May 1977.The troglobites which have been described fromBelize are listed in chronological order in Table 2.GuatemalaThe first cave inhabiting species to be describedfrom Guatemala was the cricket Arachnomimuscavicola Saussure. This species was collected byGeorge C. Champion during the period of 1879-1881in Grutas de Lanquin, Alta Verapaz (Saussure, 1897).Dr. Giaquinto ,1ira, an Italian physician studyingmalaria and sleeping sickness in Guatemala in 1933,visited Cueva de Sepacuite, Alta Verapaz, and obtaineda series of a new species of catopid beetle,described by J eannel (1936) as Ptomaphagus (A delops)giaquintoi. Additional specimens of the samespecies were obtained in 1948 from Grutas de Lan·quin by R. D. Mitchell (Peck, 1973c).On June 13, 1959, William A. Varnedoe collecteda new species of milliped in Grutas El Silvino, IzabaJ.Also in June he visited and made collections in Grutasde Lanquln, Alta Verapaz; Cueva Jobitzinaj, Peten;and Cueva Caman, Solola.In 1960 and 1961 W. E. Duellman, .f. Knox Jones,Jr., and others collected bats from caves in severalareas of Guatemala (Jones, 1966).Collections were made in Cueva de Tabacal andCueva de los Resadores, Huehuetenango, in November1967 by David McKenzie. In the latter cave hefound a remarkable troglobitic isopod (Schultz,1977).During January and February 1968 the ExplorersClub of New York sponsored an expedition to AltaVerapaz (Gurnee, 1968). The biologist for this expeditionwas Bro. G. Nicholas, and he made collectionsin 12 caves (Nicholas, 1968). Probably the mostnotable find of this expedition was a new genus andspecies of cave inhabiting crab (Smalley, 1970) fromCueva Seamay. Other species of interest included astill undescribed troglobitic planarian in Grutas deLanquln. This is the only cavernicole planarianknown from Guatemala.In August 1969 Stewart B. and James H. Peckvisited eight caves in Izabal and Alta Verapaz (Peckand Peck, 1973). Among the fauna of in terest wereblind amphipods (Ruffo and Vigna Taglianti, 1974),spider (Gertsch, 1973b), and pseudoscorpion (Much.more, 1973b).During the course of the 1971 Italian ZoologicalMission to Mexico, three caves in Peten were visitedby Valerio Sbordoni, Roherto Argano, and AldoZullini (1974).D. Dreux, P. J ouhert, and O. Rubio of the Centred'Etudes et de Recherches Speleologiques en MilieuTropical explored numerous caves in many parts ofGuatemala (Dreux, 1976). Their biological collectionsin Alta Verapaz included, most notably, two new speciesof blind crab, described by Delamare Deboutte·ville (1976) as Typhlopseudothelphusa juberthiei andT. mitchelli.On January 22, 1972, David McKenzie made a col·lection in Cueva Jobitzinaj, Peten. This included anew species of ricinuleid, described by Gertsch(1977a) as Cryptocellus cookei. In May 1972 severalcaves in the Montaiias de Cuilco near La Libertad,Huehuetenango, were visited by David McKenzie andStuart Murphy. Their collections included a new genusand species of rhaphidophorid cricket (Hubbell,14

Table 2.-ChronologicaJ list of troglobites described fromMexico, Guatemala, and Belize.Mhico1867. Lepisma anophthalma Bilimek [=A nelpistina anophthalmal1891. Lepisma cacahuami/pensis Herrera [syn. of Anelpistinaanophthalma]1900. Coni/era stygia Packard1936. Anoptichthys jordani Hubbs and Innes [=Astyanaxjordanil1936. Cirolana anops Creaser [=Creaseriella anopsl1936. Antromysis cenotensis Creaser [=Antromysis (Antromysis)cenotensis]1936. Palaemon morleyi Creaser [=Creaseria morleyiJ1936. Typhlatya pearsei Creaser1938. Porcellio pearsei Creaser [=l'richorhina pearseiJ1938. Wanops coecus Chamberlin and lvie [=Oonopscoecus]1938. Anopsicus pearsei Chamberlin and lvie [=Pholcophorapearseil1938. Gymnostreptus zizicolens Chamberlin [=Orthoporuszizicolens]1938. Trogolaphysa maya Mills [=Troglopodetes mayaI1938. Cyphoderus innominatus Mills1938. Sulcuncus falciferus Mills [=Metasinella falcifera]1938. Tohila atelomma Hubbell1938. l'yphlias pearsei Hubbs [=Typhliasina pearsei]1938. Pluto infernalis Hubbs [=Ophisternon infernale]1941. Nuevobius cavicolens Chamberlin1942. Diaptomus (Microdiaptomus) cokeri Osorio Tafall1942. Protrichoniscus bridgesi Van Name [=BrackenridgiabridgesiJ1942. Serrobunus boneti Goodnight and Goodnight [=Hoplobunusboneti]1942. Garcibius osorioi Chamberlin1942. Newportia sabina Chamberlin [=Newportia (Scolopendrides)sabinal1942. Cavota crucis Chamberlin [=Cleidogona crucis]1942. Bonetesmus verus Chamberlin1942. Ceuthauxus palmitonus Chamberlin1942. Pararachistes amblus Chamberlin [=Pararhachistesamblusl1942. Mexaphaenops prietoi BolIvar1943. Procambarus rodriguezi Hobbs [=Procambarus (A ustrocambarus)rodriguezil1943. Oncopodura atoyacense Bonet1943. Oncopodura prietoi Bonet1944. Chinquipellobunus osorioi Goodnight and Goodnight[=Hoplobunus osorioi]1944. Plusiocampa (Litocampa) atoyacensis Wygodzinsky[=Litocampa atoyacensis]1944. Paratachycampa boneti Wygodzinsky1944. Juxtlacampa juxtlahuacensis Wygodzinsky1944. Parallocampa (Parallocampa) cavernicola Wygodzinsky[=Podocampa cavernicolal1945. Acherontides atoyacense Bonet1945. Spelaeogastrura guerrerense Bonet1946. Leucohya heteropoda Chamberlin1946. Cryptocellus osorioi BolIvar1946. Acherontides potosinus Bonet1946. Anoptichthys antrobius Alvarez [=syn. of Astyanaxjordanil1947. Vachonium boneti Chamberlin1947. Vachonium maya Chamberlin1947. Pararrhopalites anops Bonet and Tellez1947. Anoptiehthys hubbsi Alvarez [syn. of Astyanax jordani]1950. Protriehoniseus villalobosi Rioja [=Braekenridgia villalobosil1950. Cirolana (Speoeirolana) pelaezi BolCvar [=Speocirolanapelaezil1951. Sphaeromieola cirolanae Rioja1951. Protriehoniscus aeostai Rioja [=Braekenridgia aeostail1951. Typhlolepidomysis quinterensis Villalobos [=Spelaeomysisquinterensis11953. 'l'yphlotricholigioides aquatieus Rioja1953. Typhlopseudothelphusa mocinoi Rioja1953. Cirolana (Speoeirolana) bolivari Rioja [=Speocirolanabolivaril1954. Prietella phreatophila Carranza1956. Triehorhina boneti Rioja1956. Cordioniscus laevis Rioja [=Mexiconiscus laevis]1956. Paravaehonium bolivari Beier1956. Troglohya earranzai Beier1958. Cylindroniseus maya Rioja1960. Spherarmadillo cavernieola Mulaik1960. Trichorhina atoyacensis Mulaik1960. Trichorhina yucatanensis Mulaik [syn. of Trichorhinapearsei]1960. Protrichoniscus potosinus Mulaik [syn. of Brackenridgiabridgesil1960. Protrichoniscu.v palmitensis Mulaik [=BrackenridgiapalmitensisJ1960. Antroniseus eavieola Mulaik [=Cylindroniscus cavicolal1960. Antroniscus balamensis Mulaik [syn. of Cylindroniscusmayal1960. Bogidiella tabaseensis Villalobos1962. Orthoporus kiemi Loomis [syn. of Orthoporus zizicolens]1962. "Cave form of Poecilia sphenops Valenciennes" reportedfrom Mhico1963. Albiorix bolivari Beier1963. Mexiterpes sabinus Causey1964. Mexiconiscus tlamayaensis Schultz [syn. of Mexiconiscuslaevis]1964. Glomeroides caecus Causey1964. Glomeroides promiscus Causey1964. Mexicambala russelli Causey1965. Cam bala speobia (Chamberlin)-lst record for M~xico1965. Mexisphodrus veraecrucis Barr [=Platynus (Mexisphodrus)veraecrucis J1966. Coahuilix hubbsi Taylor1966. Speocirolana thermydronis Cole and Minckley1966. lI1exisphodrus profundus Barr [=Platynus (Mexisphodrus)profundus]1967. Antroforceps bolivari Barr1967. Mexaphaenops elegans Barr1967. Mexaphaenopsfishi Barr1967. Paratrechus (Hygroduvalius) pallescens Barr1968. Eodri/us mexicanus Gates1968. Ptomaphagus (Adelops) troglomexicanus Peck1968. Typhlochactas rhodesi Mitchell1968. Typhlochactas reddelli Mitchell1968. Niptus absconditus Spilman1969. Tyrannochthonius troglobius Muchmore1969. Poterpes egeo Causey [=Mexiterpes egeol1969. Poterpes fishi Causey [=Mexiterpes fishil15

Table 2.-(continued)1970. Eodrilus albus Gates [=Eodrilus albidus]1970. Sphaerolana affinis Cole and Minckley1970. Sphaerolana interstitialis Cole and Minckley1970. Cylindroniscus vallesensis Schultz1971. Cyathura sbordonii Argano1971. Mexiweckelia colei Holsinger and Minckley1971. Mexiweckelia particeps Holsinger1971. Typhlochactas elliotti Mitchell1971. Agastoschizomus lucifer Rowland1971. Schizomus cookei Rowland1971. Schizomus mitchelli Rowland1971. Schizomus reddelli Rowland1971. Euagrus cavernicola Gertsch1971. Aphonopelma stygia Gertsch [=Schizopelma stygia]1971. Cicurina coahuila Gertsch [=Cicurina (Cicurella)coahuila]1971. Cicurina mina Gertsch [=Cicurina (Cicurusta) mina]1971. Tegenaria blanda Gertsch1971. Tegenaria caverna Gertsch1971. Leptoneta capilia Gertsch1971. Leptoneta delicata Gertsch1971. Leptoneta isolata Gertsch1971. Leptoneta reclusa Gertsch1971. Nesticus nahuanus Gertsch1971. Metagonia atoyacae Gertsch1971. Metagonia pachona Gertsch1971. Metagonia pura Gertsch1971. Metagonia tlamaya Gertsch1971. Pholcophora exigua Gertsch1971. Pholcophora gruta Gertsch1971. Psilochorus delicatus Gertsch1971. Psilochorus diablo Gertsch1971. Cryptocellus reddelli Gertsch1971. Hoplobunus inops Goodnight and Goodnight [=Troglostygnopsisinops]1971. Rhagidia trisetatus Elliott and Strandtmann [=Rhagi.dia trisetata]1971. Rhagidia weyerensis (Packard)-lst M~xico record1971. Strongylodesmus harrisoni Causey1971. Unculabes crispus Causey1971. Unculabes versatilis Causey [syn. of Unculabes cris·pus]1971. Bonetesmus novenus Causey [syn. of Bonetesmusverus]1971. Mexicambala blandus Causey [=Mexicambala blanda]1971. Mexicambala flShi Causey1971. Mexicambala inopis Causey1971. Mexaphaenops intermedius Barr1972. Opisthobursa mexicana Benazzi1972. Dimarcus villalobosi Mitchell and Kawakatsu [syn. ofOpisthobursa mexicana]19'72. Conasellus pasquinii Argano [=Caecidotea pasquiniil1972. Mexisteruuellus coahuila Cole and Minckley1972. Mexistenasellus parzefalli Magniez1972. Mexistenasellus wilkensi Magniez1972. Vejovis gracilis Gertsch and Soleglad [=Vaejovis gracilis]1972. Aphrastochthonius parous Muchmore1972. Aphrastochthonius russelli Muchmore1972. Leucohya magnifica Muchmore1972. Mexobisium paradoxum Muchmore1972. Paravachonium superbum Muchmore1972. Pachychitra grandis Muchmore1972. Nesticus arganoi Brignoli1972. Cleidogona baroqua Shear1972. Cleidogona pecki Shear1972. Mexiterpes metallicus Shear1972. Paracophus caecus Hubbell1972. Paracophus cladonotus Hubbell1972. Paracophus lippus Hubbell1973. Dugesia barbarae Mitchell and Kawakatsu1973. Dugesia typhlomexicana Mitchell and Kawakatsu1973. Dugesia mckenziei Mitchell and Kawakatsu1973. Sphaeromicola coahuiltecae Hobbs and Hobbs1973. Mexiweckelia mitchelli Holsinger1973. Spelaeomysis olivae Bowman1973. Alpheopsis stygicola Hobbs1973. Macrobrachium villalobosi Hobbs1973. Neopalaemon nahuatlus Hobbs1973. Procambarus (Austrocambarus) oaxacae oaxacaeHobbs1973. Procambarus (Austrocambarus) oaxacae reddelliHobbs1973. Aphrastochthonius major Muchmore1973. Tyrannochthonius pallidus Muchmore1973. Mexobisium maya Muchmore1973. Troglohya mitchelli Muchmore1973. Vachonium kauae Muchmore1973. Schizomus bartolo Rowland1973. Schizomus firstmani Rowland1973. Schizomus pecki Rowland1973. Schizomus lukensi Rowland1973. Tarantula chacmool Rowland [=Paraphrynus chacomool]1973. Tarantula chiztun Rowland [=Paraphrynus chiztunl1973. Euagrus anops Gertsch1973. Schizopelma reddelli Gertsch1973. Theotima pura Gertsch1973. Metagonia martha Gertsch1973. Hoplobunus apoalensis Goodnight and Goodnight1973. Hoplobunus planus Goodnight and Goodnight1973. Glomeroides addititius Causey1973. Acutangulus alius Causey1973. Ceuthauxus constans Causey1973. Tylogoneus minus Causey1973. Tylogoneus rainesi Causey1973. Pseudosinella bonita Christiansen1973. Pseudosinella strinatii Christiansen [=Pseudosinellapetrustrinatiil1973. Cixius orcus Fennah1973. Oeclidius hades Fennah1973. Mexisphodrus urquijoi Hendrichs and Bolfvar [=Plaotynus (Mexisphodrus) urquijoiJ1974. Troglocubanus perezfarfanteae Villalobos1974. Mexistenasellus magniezi Argano1974. Mexicerberus troglodytes Schultz1974. Bogidiella arganoi Ruffo and Vigna Taglianti1974. Bogidiella sbordonii Ruffo and Vigna Taglianti1974. Bithynops luscus Holthuis1974. Leptoneta limpida Gertsch1974. Cryptocellus sbordonii Brignoli1974. Caecoa arganoi Silhavy1974. Hoplobunus queretarius Silhavy1974. Troglostygnopsis anophthalma Silhavy1974. Ortholasma sbordonii Silhavy1974. Glomeridesmus sbordonii Shear1974. Glomeroides pellucidus Shear16

Table 2.-(concluded)1974. Cleidogona felipinna Shear1974. Bonetesmus ojo Shear1974. Aceratophallus scutigeroides Shear1974. Unculabes arganoi Shear1974. Unculabes causeyae Shear1974. Unculabes porrensis Shear1974. Speodesmus pecki Shear1974. Mexanillus .~bordonii Vigna Taglianti1974. Troglobacanius bolivari Vomero1974. Troglobacanius maya Vomero1974. Troglobacanius reddelli Vomero1974. Troglobacanius sbordonii Vomero1975. Mexilana saluposi Bowman1975. Agastoschizomus huitzmolotitlensis Rowland1975. Schizomus pallidus Rowland1975. Paraphrynus baeops Mullinex1975. Paraphrynus velmae Mullinex1975. Reddellobus troglobius Causey1976. Caecidotea chinpas Bowman1976. Typhlatya campecheae Hobbs and Hobbs1976. Typhlatya mitchelli Hobbs and Hobbs1976. Opisthobursa josephinae Benazzi1977. Neogovea mexasca Shear1977. Troglophiloscia laevis Schultz1977. Mayaweckelia cenoticola Holsinger1977. Mayaweckelia yucatanensis Holsinger1977. Antromysis (Antromysis) reddelli Bowman1977. Diplocentrus anophthalmus Francke1977. Diplocentrus mitchelli Francke1977. Vachonium cryptum Muchmore1977. Metagonin torete Gertsch1977. Metagonin chiquita Gertsch1977. Theotima martha Gertsch1977. Cicurina maya Gertsch [=Cicurina (Cicurella) maya)1977. Orthoporus spelaeus Causey1977. Ptomaphagus (Adelops) mckenziei Peck1977. Caecidotea vomeroi Argano1977. Caecidotea zullinii Argano1977. Etlastenasellus mixtecus Argano1977. Bogidiella michaelae Ruffo and Vigna Taglianti1977. Bogidiella niphargoides Ruffo and Vigna Taglianti1977. Bogidiella orchestipes Ruffo and Vigna Taglianti1977. Bogidiella vomeroi Ruffo and Vigna Taglianti1977. Macrobrachium acherontium Holthuis1977). In August 1972 S..and j. Peck visited CuevaNojohnaj Cohultunich, Peten.In January 1973 Henry Frania and Michael Shawcrossdiscovered a new genus and species of troglobiticcarabid beetle in Cueva Seamay, Alta Verapaz(Barr, 1973).The Swiss biospeleologist, Pierre Strinati, visitedcaves in Guatemala during April 1973. His collectionsin Cueva Chirrepeck, AI ta Verapaz, includeda new trogloLitic pseudoscorpion (Beier, 1974) anddipluran (Conde, 1975).On September 30, 1973, Valerio Shordoni, RobertoArgano, and Aldo Zullini visited Cueva de lasCalaveras, Huehuetenango.1977. Trichodactylus (Rodriguezin) mensabak Cottarelli andArgano1977. Hoplobunus zullinii Silhavy1977. Mexotroglinus sbordonii Silhavy1977. Cleidogona hunapu Shear1977. Polylepiscus vomeroi Shear1977. Caramba delburro Shear1977. Caramba delnegro Shear1977. Caram ba grandeza Shear1977. Chinpadytes bolivari Vigna Taglianti1978. Matta mckenziei Shear1978. Diplocentrus cueva Francke1979. Paraphrynus reddelli Mullinex1979. Pseudosinella aera Christiansen1979. Pseudosinella finca Christiansen-1st Mexico recordBelize1973. Mexobisium goodnighti Muchmore1973. Vachonium belizense Muchmore1973. Metagonin jarmila Gertsch1973. Jarmilka alba Shear1977. Stygnomma pecki Goodnight and Goodnight1977. Cynortina misteca Goodnight and GoodnightGuatemala1936. Ptomaphagus (Adelops) ginquintoi Jeanne!1972. Aphrastochthonius verapazanus Muchmore1973. Mexobisium guatemalense Muchmore1973. Telema mayana Gertsch1973. Pseudosinella finca Christiansen1973. Speocolpodes franini Barr1974. Bogidiella holsingeri Ruffo and Vigna Taglianti1974. A ceratophallus scutigeroides Shear1974. Paraliochthonius strinatii Beier1975. Juxtlacampa hauseri Beier1976. Typhlopseudothelphusa juberthiei Delamare Deboutteville1976. Typhlopseudothelphusa mitchelli De!amare Deboutteville1977. Colombophiloscin cavernicola Vandel-1st Guatemalarecord1977. Caecidotea mitchelli Argano1977. Bogidiella pasquinii Ruffo and Vigna Taglianti1977. Tridontomus loomisi Shear1977. Mayaphaenops sbordonii Vigna TagliantiIn September 1975 Roberto Argano, ValerioSbordoni, Vincenzo Vomero, and Aldo Zullinivisited 11 caves in the Altos de los Cuchamatanes,Huehuetenango. Among the more notable discoverieswere new species of bogidiellid amphipod and a newgenus and species of treehine beetle (Sbordoni et aI.,1977).All species of troglobite which have been describedfrom Guatemala are listed in Table 2 in the chronologicalorder of their description.17

CAVE REGIONSOnly the cave regions of Mexico, Guatemala, andBelize which have been studied with respect to theirinvertebrate fauna are discussed. The greatest emphasishas naturally been placed on the better studiedregions, and this discussion is further limited by theincomplete knowledge of the physiography andgeology.The definition of cave regions varies greatly fromarea to area. In general I have attempted to defineregions on the basis of isolation from other regions.This has been simple in some areas, particularly innorthern Mexico where isolated limestone ranges suchas Pico de Carrizal and the Sierra de Gomas are separatedby noncavernous deposits. In other areas, however,the definition of regions has been more arbitrary.An example is in southern San Luis Potosi and adjacentHidalgo and Queretaro. Despite the fact that thisarea is a rather contiguous region of limestone, fiveregions have been defined: Aquismon, Xilitla, LagunaColorado, Xilitla Plateau, and Jalapa. These regionsshow some distinct faunal differences and some differencesin elevation, geology, and vegetation. Onlywith further study and the availability of detailedgeologic and topographic maps can these and manyother regions be defined more naturally.In the description of each region I have commentedbriefly, where possible, on geology, karstmorphology, elevation, and cave type. I have alsoincluded a general discussion of the composition andrelationships of the cave fauna. The locations of thecave regions discussed below are given in Fig. 2.The discussion of physiographic provinces follows,more from convenience than conviction, the systemproposed by Raisz (1964). Other systems whichdeserve consideration include those of West (1964),Alcorta Guerrero (1966), and Russell (1969). Table 3summarizes by physiographic province and caveregion the cavernicole fauna and numher of biologicallyinvestigated caves in .\lexico, Belize, and Guatemala.Sierra Madre OccidentalThe northern portion of the Sierra Madre Occidentalconsists of a series of north-south ranges lyingbetween the ranges of the Sonoran desert and themain plateau of the Sierra. It extends from the UnitedStates into northeastern Sonora. The principalmass of the Sierra Madre Occidental is a broad highplateau formed of Tertiary volcanics extending fromnorthern Chihuahua into northern J alisco where it isbounded on the south by the Rio Santiago. The easternslopes of the Sierra tend to be gentle, but thewestern side is abrupt and cut by deep gorges, includingBarranca del Cobre, with a depth of morethan 2,000 meters. Although most of the area isformed of igneous rock, a few isolated outcrops ofCretaceous limestone have been exposed by erosionof the overlying Tertiary deposits. King (1939) hasdiscussed the geology of the northern Sierra MadreOccidental, and his geologic map shows a few of thelarger of these outcrops.Speleologically, this region is virtually unstudied.Many large shelter and shelter-like caves have beenreported, but only one has been biologically investigated.The limestone region which has been studied isa limited area near the town of Santo Tomas to thewest of Ciudad Guerrero in central Chihuahua. Alarge limestone cave is known from southern Chihuahuanear Guadalupe y Calvo but has not been visitedby speleologists.Santo Tomas, Chihuahua.-This region consists ofan isolated outcrop of Cretaceous limestone of only afew kilometers extent. Water flowing south off a lowrange of igneous hills sinks in several sumidero-typeentrances at the ends of arroyos. The water is reportedto emerge from a spring or springs a fewkilometers away. Only three caves have been investigated,the largest of which is the Socavon de SantoTomas, which attains a depth of 110 meters and alength of about 200 meters. The fauna of the regionis limited and no troglobites are known. Of the 11species thus far identified only spiders of the generaMeioneta (Linyphiidae) and Psilochorus (Pholcidae)appear to be troglophiles.Villa Matamoros. Chihuahua.-The Villa Matamorosregion is an area of Tertiary volcanics southof the city of Parra!' The only cave investigated isCueva del Salitre, a 40-meter-long passage formed inignimbrite and inhabited by a colony of cave swallows.Seven species have been identified from thecave; of these only chernetid pseudoscorpions andspiders of the genus Filistatinella (Filistatidae) aretrue cave associates.Edwards PlateauRaisz (1964) includes this area as part of the GulfCoastal Plain province, but I feel it should be includedwith the Edwards Plateau of Central Texas.It consists of a narrow band of Cretaceous limestoneex tending from a few kilometers ~orthwest of CiudadAcui'ia to the Serranias del Burro. It has been isolatedfrom the Edwards Plateau proper by the incision ofthe Rio Grande.Ciudad Acuna, Coahuila.-The only cave knownfrom this region is Cueva de los Lagos, now inundated19

Table 3.-Number of troglobites, troglophiles, total species, and biologically investigated caves in Mexico,Guatemala, and Belize, arranged by physiographic province, state, and cave region. Numbers following theregion name refer to the location map (Fig. 2); Santo Tomas and Villa Matamoros regions not shown on map.TroglobitesTroglophilesRegion Aquatic Terrestrial Aquatic Terrestrial Total Species CavesSierra Madre OccidentalChihuahuaSanto Tomas 0 0 0 2 3 3Villa Matamoros 0 0 0 1 7 1Edwards PlateauCoahuilaCiudad Acuna (6) 0 2 1 10 17 1Basins and RangesChihuahuaSalaices (2) 0 1 0 5 17 2Santa Elena (1) 0 0 0 2 5 3CoahuilaCuatro Cienegas de Carranza (9) 7 0 1 10 25 25DurangoRancho Descubridora (3) 1 0 1 7 12 1Sierra Madre OrientalCoahuilaHermanas (8) 0 0 0 6 10 2Monclova (10) 0 0 0 0 1 1Sierra de las Animas (11) 0 0 0 7 9 1Sierra Arteaga (21) 0 0 0 8 17 1Sierra de Maynin (12) 0 0 0 5 8 4Sierra de Santa Rosa (7) 1 0 0 5 9 3DurangoCiudad Lerdo (5) 0 0 0 8 13 4Sierra de la India (4) 0 4 0 5 14 2HidalgoJacala (55) 0 3 3 17 39 10Lagunilla (58) 0 1 0 5 15 1Tonoltongo (57) 0 0 0 2 3 1Nuevo LeonCanon Huasteco (19) 0 2 0 14 23 3Canon de Santa Rosa (27) 2 0 0 0 4 3Cerro de la Boca (25) 0 1 0 14 16 1Cuesta de Chipinque (20) 0 2 0 8 12 3GaJeana (26) 0 1 0 16 23 7Pico de CarrizaJ (13) 0 3 0 12 _u ')..1Potrero Redondo (24) 0 1 0 4 6 2Sierra del Fraile (18) 0 6 0 7 14 2Sierra de Garia (17) 0 0 0 0 2 1Sierra de Gomas (15) 0 10 0 20 36 4Sierra de Iguana (14) 0 0 0 1 3 2Sierra de Santa Clara (16) 0 0 0 0 1 1Sierra de la Silla (23) 2 3 0 16 33 2Sierra EI Tarillal (22) 0 0 0 5 5 1Zaragoza (31) 0 0 0 2 3 3Nuevo Leon and TamaulipasPurificacion (32) 4 15 2 46 100 3320

Table 3.-(continued)TroglobitesTroglophilesRegion Aquatic Terrestrial Aquatic Terrestrial Total Species CavesPueblaCuetzalan (60) 1 10 7 43 87 20Xicotepec dejuarez (59) 0 0 0 3 11 4QueretaroAhuacatl

Table 3.-(concluded)TroglobitesTroglophilesRegion Aquatic Terrestrial Aquatic Terrestrial Total Species CavesOaxaca (concluded)San Sebastian de las Grutas (75) 0 3 0 11 21 4Santiago Apoala (73) 0 2 0 13 30 6Valle Nacional (74) 4 4 2 22 42 5PueblaSan Pablo Zoquitlan (71) 0 1 0 10 17 9VeracruzAtoyac (67) 8 15 5 55 111 25Orizaba (68) 0 12 0 49 104 28Tezonapa (69) 0 2 0 9 14 1Chiapas-Guatemala HighlandsChiapasAltamirano (85) 3 1 4 7 16 9Bochil and Soyalo (82) 0 3 1 9 16 6Comitan de Dominguez (87) 1 4 0 11 25 2Ixtacomitan (77) 0 0 0 2 4 3Malpaso (76) 0 2 0 5 16 10Montebello (88) I 4 10 19 49 8Motozintla (89) 1 4 0 3 8 2Palenque (80) 0 0 0 0 2 1Rancho del Cie/ito (86) I 2 0 2 7 2..,San Cristbhal de las Casas (84)I 7 2 21 53 38Silllojovel and Tila (81) 3 I 1 5 17 8Tux tla Gutierrez (83) 0 1 12 39 81 22TabascoTapijulapa (79) 2 1 0 11 20 1Teapa (78) 3 5 1 17 59 12Cayo, BelizeAugustine (100) 0 1 0 16 35 7Caves Branch (101) 0 7 0 19 33 8Alta Verapaz, GuatemalaCohan (96) 2 3 0 8 18 6Lanqu in (97) 1 3 1 17 47 7Raxruja (95) 0 0 4 0 4 3Senahu (98) 1 7 1 4 18 5Huehuetenango, GuatemalaAltos Cuchumatanes (92) 2 2 1 2 13 12Montanas de Cuileo (91) 0 0 0 3 5 5Santa Ana Huista (90) 0 0 0 3 11 3Izabal, GuatemalaLago de Izahal (99) 0 0 0 7 12 4Peten, GuatemalaFlores (94) 0 0 0 11 15 5Yaxchilan (93) 0 0 0 11 13 3Yucatan PeninsulaCarnpecheSierra de Bolonchen (102) 5 5 6 45 94 18Campeche, Quintana Roo, and YucatanCoastal Plain (104) 10 14 33 116 515 157YucatanSierra de Ticul (103) 3 14 3 79 290 4122

y the waters of the Amistad Reservoir. The cave,which consists largely of a steeply sloping passageending in a deep pool, is formed in the Salmon PeakFormation of Cretaceous age (Smith, 1970) at anelevation of 330 meters.The fauna, as is to be expected, is closely relatedto that of Texas immediately across the Rio Grande.Of the 17 species identified, two are troglobites: theagelenid spider Cicurina (Cicurella) coahuila Gertschand the cambalid milliped Cambala speobia (Chamberlin).The latter species is widespread in CentralTexas. Troglophiles include the porcellionid isopodPorcellio gertschi Van Name, the pholcid spiderPhysocyclus enaulus Crosby, a scytodid spider of thegenus Loxosceles, the collembolan Pseudosinellaviolenta (Folsom), and the carabid beetles Rhadinearaizai (Bolivar) and Tachys (Tachys) proximus Say.Basins and RangesThe Basins and Ranges province consists of an areaof folded and faulted mountain ranges separated bywide valleys and basins. The mountains are generallyoriented along a northwest-southeast axis and frequentlyenclose large closed basins (boisones). Themost notable of these is the Bolson de Mapimi inDurango, but the Bolson de Cuatro Cienegas deCarranza in Coahuila is of particular importance becauseof its rich endemic subterranean aquatic fauna.The western ranges are largely igneous, but to theeast they tend to be composed of folded Cretaceoussediments. This province extends from the southwesternUnited States into northern Durango andsouthern Coahuila. It is bounded on the west by theSierra Madre Occidental, on the east by the SierraMadre Oriental, and on the south by the Cross Rangesof the Sierra Madre Oriental. The average elevation ofthe filled basins is about 1,200 meters, with themountains rising up to several thousand meters ahovethe valleys. Russell and Raines (1967) have describedthe general geology of the Laredo-Monterrey areawith special reference to cave origins. Brand (1937)includes a concise account of the physiography andgeology of northwestern Chihuahua.Despite its proximity to the United States, thisarea has been little studied speleologically. Only sixcaves are known for the province outside of the vicinityof Cuatro Cienegas de Carranza. Most work evenin the Cuatro Cienegas de Carranza region has been inthe waters flowing into the basin itself, with the cavesin the mountains surrounding the basin remainingvirtually unknown.Salaices, Chihuahua.-This region includes severallow limestone hills in the vicinity of Salaices, a smalltown on the highway between Jimenez and Parral.The only caves known are Cueva de los Muchachos, avertical sinkhole dropping to a short horizontal passage,and Cueva del Diablo. The latter cave is an extensivemaze containing more than one kilometer ofexplored passage (Reddell, 1977a). The only faunarecorded from Cueva de los Muchachos is a species ofrhaphidophorid cricket. Cueva del Diablo is generallydry but it does contain 16 species, of which one is apossible troglobite. Cirolanid isopods found dead in alow pool remain to be rediscovered. The only apparenttroglobite is the endemic pholcid spider Psilochorusdiablo Gertsch. The troglophile fauna includesthe nesticid spider Eidmannella pallida (Emerton),the pholcid spider Physocyclus enaulus Crosby, andthe psocid Psyllipsocus ramburii Selys-Longchamps.The fauna is typical of the arid regions of northernMexico and adjacent United States.Santa Elena, Chihuahua.-This is a poorly definedregion located south of Santa Elena Canyon on theRio Grande. The only cave to be investigated withrespect to its invertebrate fauna is Sotano de Sauz,a 220-meter-deep cave with an unusually high airtemperature. A massive die-off of the bat Mormoopsmegalophylla megalophylla Peters was reported bySprouse (1977). The only probable troglophile identifiedfrom the cave is a pholcid spider of the genusPsilochorus.Cuatro Cienegas de Carranza, Coahuila.-Thisregion includes the Bolson de Cuatro Cienegas deCarranza and several ranges of mountains which surroundit. The basin is about 40 kilometers wide (eastto west) and 30 kilometers long (north to south) andis formed at an elevation of about 740 meters. Thesurrounding mountains attain elevations of more than3,000 meters. Minckley (1969) has provided an excellentsummary of the geology, hydrology, and physiographyof this region. The deposits in the closedbasin include alluvium, detrital deposits from themountains, and gypsum (mostly in the form of sandor gypsite). Drainage in the basin is almost entirelysubterranean and water rising from deep sourcesalong the bases of the mountains either emerges assprings or flows underground through solution channels.Collapse of the poorly consolidated gypsite andother deposits allows access to these subterraneanstreams through the means of shallow natural wellsor lagunas. The surrounding mountains are formedlargely of Cretaceous limestone.Only five caves have been investigated in thisregion. One of these, Cueva de San Vicente, is locatedin the Sierra de San Vicente to the east of the basin,while the remaining four occur in Canon del Pedregosoin the Sierra de San Marcos to the west of the23

asin. Cueva de San Vicente is a large single-roomcave heavily mined for phosphates. The cave is verydry and the fauna of no particular interest. Of thecaves in Canon del Pedregoso, only Cueva del Pedregosois extensive. It consists of more than' 1,000meters of large paSS

Although a few caves in the Northern Section havebeen well studied, many ranges remain unvisited.Very little is known of the caves and cave biologybetween Monterrey and the Purificacion region orbetween Xilitla and Jalapa. The Low Ranges and theCross Ranges remain virtually unknown.Hennanas, Coahuila.-This is a poorly definedregion located near the small town of Hermanas.Cueva de la Herradura is the only cave with studiedinvertebrate fauna. This small, dry bat cave is locatedin a small outcrop of limestone in the plains in frontof the main ranges of the Sierra Madre Oriental. Notroglobites are known among the 10 species reported.The fauna is typical of that of the arid parts of northernMexico and includes the filistatid spider Filistatoidesn. sp., the pholcid spiders Physocyclus hoogstraaliGertsch and Davis and Psilochorus sp., thedermestid beetle Dermestes carnivorus Fabricius, andthe tenebrionid beetles Alphitobius laevigatus (Fabricius)and Cryptoglossa mexicana mexicana Champion.Monclova, Coahuila.-This region is located to thewest of the town of Monclova. The only cave that hasbeen studied is Bocas del Carmen, a small shelter-likecave located on the cliffs overlooking the highway toCuatro Cienegas de Carranza. The cave is dry andgenerally unproductive biologically. The only speciesso far identified is a camel cricket of the genus Ceuthophilus.Sierra de las Animas, Coahuila.-The Sierra de lasAnimas is an isolated range formed of Cretaceouslimestone east of Monclova. The only cave known isCueva de las Animas at an elevation of 760 meters.It has been extensively mined for phosphates andnow consists of one large dry room with a few smallmoist alcoves. :\0 troglobites have been found in thecave. The troglophile fauna is typical of that of northernMexico and includes three troglophilic spiders:the filistatid Filistatoides n. sp., the pholcid Physocyclushoogstraali Gertsch and Davis, and the scytodidLoxosceles belli Gertsch. The only other nonguano-associatedtroglophile known from the cave isthe gryllid cricket Paracophus subapterus Chopard.Sierra Arteaga, Coahuila.-The Sierra Arteaga islocated east of the town of Arteaga: it comprisesCretaceous limestone of the Aurora and Cupido Formations(de Cserna, 1956). The only cave that hasbeen studied in the region is Gruta de Cuevecillas,which is an elongated chamber subdivided by formationsinto smaller "rooms." The cave is moist only inthe lower sections. Of the 17 species recorded fromthe cave none are troglobitic and several are apparenttroglophiles. These include the agelenid spider Tegenariagertschi Roth, the filistatid spider Filistatinellacrassipalpis Gertsch, the pholcid spider Metagoniacoahuila Gertsch, the collembolan Pseudosinellareddelli Christiansen, the psocid Psyllipsocus ramburiiSelys-Longchamps, and the carabid beetle Rhadinerotgeri (Bolivar and Hendrichs). This fauna istypical of that of the northern High Sierra.Sierra de Mayrim, Coahuila.-The Sierra de Mayninis an east-west trending range of mountains to thesouth of the now-dry Laguna de Mayran. It is formedof Cretaceous limestone of the Aurora Formation (deCserna, 1956). Only four small caves, all at the northernbase of the range at an elevation of about 1,150meters, have been investigated. These caves areformed by the solution of gypsum beds in the limestoneand are very dry. No troglobites are known, buteight species of troglophile and trogloxene have beenidentified. The fauna, which is typical of the aridregions of northern Mexico, includes spiders of thegenera Filistatoides (Filistatidae) and Physocyclus(Pholcidae) and tenebrionid beetles of the generaCentrioptera, Cryptoglossa, and Eleodes.Sierra de Santa Rosa, Coahuila.-The Sierra deSanta Rosa is located east and north of Musquiz.It is a southern extension of the general uplift tenninatingin Mexico in the Sierra del Carmen and Serraniasdel Burro. The geology of the northern part ofthe Sierra de Santa Rosa has been discussed bySmith (1970). Only two caves in this region havebeen investigated with respect to their invertebratefauna. Cueva del Leon is a horizontal bat cave containinga large population of the Mexican freetailbat Tadarida brasiliensis mexicana (Saussure). Itsinvertebrate fauna includes species typical of freetailbat caves. The only troglophile known from the caveis the pholcid spider Physocyclus enaulus Crosby, aspecies abundant in northern Mexico and the southwesternUnited States. The only other cave studiedis the natural well Pozo de El Potrero to the east ofMusquiz. It is inhabited by a spider of the pholcidgenus 1Hetagonia and by the troglobitic ictaluridcatfish Prietella phreatophila Carranza. a speciesmost closely related to blind catfish from CentralTexas.Ciudad Lerdo, Durango.-This is a poorly definedregion which includes several arid mountain ranges tothe south of Ciudad Lerdo. The geology of this regionhas been discussed by Kellum (1936). The principalcavernous deposits are Cretaceous formations ofAptian-Albian limestone. The only cave which hasbeen examined for its invertebrate fauna is Cueva delGuano, located in the Sierra de la Espana. This rangeis near the western limit of the Cross Ranges of theSierra Madre Oriental. Cueva del Guano is a largehorizontal cave inhabited by several species of bat,25

including the Mexican freetail bat Tadarida brasiliensismexicana (Saussure) (Reddell, 1977a). Of the 11species known from the cave, the most interesting isthe ricinuleid Cryptocellus mitchelli Gertsch. As inthe case of C. reddelli Gertsch in Cueva de los Riscosnear Mapimi, this is an obvious relict now isolated inCueva del Guano by the surrounding desert; it doesnot show troglobitic adaptations, however. Much ofthe fauna is typical of that of Mexican freetail batcaves. Troglophiles include spiders of the pholcidgenus Physocyclus and the scytodid genus Loxosceles,the collembolan Pseudosinella violenta (Folsom), thepsocid Psyllipsocus ramburii Selys-Longchamps, andthe spider beetle Niptus abstrusus Spilman. With theexception of the ricinuleid, the fauna is typical ofthat of western Texas and the arid regions of northernMexico.Sierra de la India, Durango.-Sierra de la India islocated to the south of Mapimi and is formed ofCretaceous limestone of the Aurora Formation(Clemons and McLeroy, 1966). One of the two cavesinvestigated in this region, Cueva de la Cucaracha, issmall and of no biological interest. The other, Cuevade los Riscos, is a large horizontal cave still not fullyexplored (Reddell, 1977a). The fauna of this cave isvery interesting in that it contains four troglobitesand 10 other identified species. Of particular interestis the presence in the cave of the highly cave-adaptedricinuleid Cryptocellus reddelli Gertsch. Other speciesof troglobite include the leptonetid spider Leptonetalimpida Gertsch and the pholcid spiders Pholcophoraexigua Gertsch and Psilochorus delicatus Gertsch.With the exception of L. limpida these troglobites aretypical elements of the cavernicole fauna of the SierraMadre Oriental to the east and southeast. Leptonetalimpida, however, has its closest affinities with speciesof the same genus in Central Texas. The troglophilefauna includes pholcid spiders of the genera Modisimusand Physocyclus and the psocid Psyllipsocusramburii Selys-Longchamps. The troglobite fauna presumablyrepresents relict species now restricted to thecavemicole habitat by the surrounding desert.jacala, Hidalgo.-This region extends from theRio Moctezuma in the north to near Zimapan on thesouth. It is bounded on the west by the Rio Moctezumaand on the east by the Rio Amajaque. Thegeology of this region has been discussed by Bodenlos(1956). The principal limestone unit in the area is themassive El Abra (=EI Doctor) Formation. Karstdevelopment is extensive in some areas, but the regionremains virtually unknown speleologically. Tencaves have been biologically investigated but onlyfive have been studied with respect to their invertebratefauna. The largest cave in the region is Cueva deEl Tenango, a small resurgence cave about 300 meterslong. The only other cave of consequence studied isSotano del Hondo de Pinalito, a vertical cave thatdescends in several drops to a depth of 175 meters.Three of the 39 species reported from the caves ofthis region are troglobites. The schizomid Agastoschizomusn. sp. is known from two caves; it isclosely related to A. huitzmolotitlensis Rowland fromthe Xilitla region. The remaining two troglobites arean undescribed species of nicoletiid thysanuran andthe gryllid cricket Paracophus cladonotus Hubbell.The latter species is also known from the Xilitla region.The troglophile fauna contains species generallyknown from the XiIitla and Aquismi)n regions. Speciesof interest include the crayfish Procambarus (Ortmannicus)toltecae Hobbs, the agelenid spider Tegenariarothi Gertsch, the nesticid spiders Gaucelmuscalidus Gertsch and Nesticus hoffmanni Gertsch, thepholcid spiders Coryssocnemis iviei Gertsch and C.simoni O. P.-Cambridge, the scytodid spider Loxoscelestenango Gertsch, the gryllid cricket Paracophusplaconotus Hubbell, and the carabid beetle Platynus(Mexisphodrus) gertschi (Hendrichs and Bolivar).Lagunilla. Hidalgo.-The only cave which has beeninvestigated in the Lagunilla region is the famedGrutas de Xoxafi. This cave is fomled in an isolatedoutcrop of EI Abra (=EI Doctor) Limestone (Segerstrom,1962). The cave has been only partially exploredbut is known to be extensive. The 15 speciesreported from it doubtless represent a slIlall percentageof the fauna. The only troglobite is the ptinidbeetle Niptus absconditus Spilman. The troglophilefauna includes two endemic spiders, Physocyclusreddelli Gertsch and Psilochorus fishi Gertsch, bothof which are closely related to species known fromother parts of the Sierra \Iadre Oriental.Tonoltongo. Hidalgo.-The Tonoltongo regionconsists of an isolated deposit, known as the TonoltongoBank, of EI Abra (= EI Doctor) Limestone(Segerstrom, 1962). The only cave known from theregion is the Grutas de Tonoltongo, from whichemerge the waters of the Rio Tonoltongo. Thewater flowing from the cave is highly mineralizedand quite warm. Only three species have been identifiedfrom the cave: these include two possible trog10phiIic carabid beetles of the genus Schizogenius.Canon Huasteco. Nuevo Leon.-Caiion I1uastecois a spectacular canyon bordered for much of itslength by high, vertical cliffs. The geology of theregion is discussed in Maldonado-Koerdell (1956).The principal limestone units are the Cupido, Cuestadel Cura, and Aurora Formations. Two of the caves(known collectively as Grutas de San Bartolo) arehorizontal passages located at the base of a diff at26

an elevation of about 900 meters. The third cavewhich has been biologically studied is a small, dryvertical pit at an elevation of about 740 meters.Two troglobites are among the 23 species knownfrom this region. Schizomus bartolo Rowland isclosely related to species known from San LuisPotosi and Tamaulipas. An undescribed milliped ofthe family Trichopolydesmidae is of uncertain affinities.The troglophile fauna includes the leptonetidspider Leptoneta pecki Gertsch, the scytodid spiderLoxosceles luteola Gertsch, and the pyrgodesmidmilliped 1Hyrmecodesmus ilymoides (Shear). Gru tai\orte de San Bartolo is inhabited by a large colonyof the Mexican freetail bat Tadarida brasiliensismexicana (Saussure).Calion Santa Rosa, Nuevo Leon.-This ill-definedregion is located west of the town of Iturbide andincludes the Canon San ta Rosa and adjacent mountains.The region is bounded on the west by the gypsumdeposits of the Galeana region. The geology ofthis region is complex: the principal cavernous limestoneis the Cupido Limestone, but the La Pena andother formations are exposed (Padilla y Sanchez,1978). Only two caves have been visited, and the onlyfauna reported are aquatic species, the troglobiticcirolanid isopod Speocirolana n. sp. and its entocytheridostracod associate Sphaeromicola cirolanaellioja. This isopod is known only from this region.Cerro de la Boca, Nuevo Leon.-This range of Cretaceouslimestone is located about 20 kilometerssouthwest of Montemorelos. The only cave known isCueva de Chorros de Agua at an elevation of 700meters. It is essentially horizontal and only about40 meters long. The fauna indudes one troglobite,the leptonetid spider Leptoneta reclusa Gertsch.Among the 15 other species reported from the caveare ctenid spiders of the genus Ctenus, the pholcidspider ilIetagonia placida Gertsch, a collembolan ofthe genus Pseudosinella, the leiodid beetle Dissochaetusaztecus Szymczakowski, and staphylinid beetlesof the genera Belonuchus and Stilicolina. All ofthese species are typical members of the fauna ofthe Sierra Madre Oriental.Cuesta de Chipinque, Nuevo Leon.-Cuesta deChipinque is located immediately to the south ofMonterrey. Several small caves and sinkholes occuron the Cuesta, only two of which have received biologicalstudy. Both are formed in Cretaceous limestoneat an elevation of about 1,500 meters.Twelve species, of which one is a troglobite, havebeen identified from the caves of the region. Thetroglobitic spider Nesticus nahuanus Gertsch is alsorecorded from caves to the east and south of thisregion. The troglophile fauna includes species typicalof the fauna of the Sierra Madre Oriental, such as thecoIlembolan Pseudosinella reddelli Christiansen, theleiodid beetle Ptomaphagus (Adelops) leo Peck, andthe rhachodesmid milliped Ceuthauxus mediatorChamberlin.Galeana, Nuevo Le&n.-The Galeana region is anextensive area of karst developed on Jurassic gypsumof the OIivido Formation. The geology has beenmapped by Moor (1980). Almost all of the drainageis subterranean, with numerous sinkholes ranging insize from a few meters to more than 200 meters indiameter and up to 100 meters in depth. One of thelarger caves, Resumidero de Pablillo, receives thefloodwaters of an extensive area (Russell and Raines,1967). Seven caves, ranging in elevation from 1,740to 2,000 meters, have been studied. Since most ofthe caves are smaIl and probably of comparativelyrecent origin, few troglobites are to be expectedfrom the region.The only cave-adapted species among the 23recorded is the spider Nesticus nahuanus Gertsch,which is widespread in the northern High Sierra. Thetroglophile fauna is closely related to that of thenorthern Sierra Madre Oriental and includes spidersof the agelenid genus Tegenaria and the pholcidgenera Metagonia, Modisimus, Physocyclus, andPsilochorus. Other elements of the fauna include thewidespread collembolan Pseudosinella reddelli Christiansenand the endemic leiodid beetle Ptomaphagus(Adelops) gypsum Peck.Pico de Carrizal, Nuevo Leon.-Pico de Carrizal isthe eroded core of a Tertiary igneous intrusion whichrises more than 1,500 meters above the surroundingplain. A ring of much lower hills is developed onlower Cretaceous limestone (Russell and Raines,1967). The only cave known in this area is Gruta deCarrizal, a largely horizontal cave which has served asa coIlecting point for water falling on the mountain;an active stream emerges from the cave entrance.Three of the 25 species recorded from the cave aretroglobites: the pseudoscorpion Leucohya magnifical\luchmore, an undescribed species of the schizornidgenus Schizomus, and the opiIionid lloplobunusosorioi (Goodnight and Goodnight). The genusLeucohya is known only by this and a second speciesfrom Grutas del Palmito to the south. Hoplobunusosorioi is known from this cave, Grutas del Palmito,and Grutas de Villa de Garcia; the presence of smalleyes and its occurrence in three isolated mountainranges indicate it is a recent troglobite. The undescribedspecies of Schizomus is closely related tospecies occurring in the Sierra Madre Oriental to thesouth. The troglophile fauna also has its closestaffinities to tile fauna of the Sierra Madre Oriental to27

the south and includes the endemic pholcid spiderMetagonia candela Gertsch, the phalangodid harvestmanPellobunus mexicanus Goodnight and Goodnight,and the gryllid cricket Paracophus subapterus Chopard.Potrero Redondo, Nuevo Leon.-This ill-definedregion occurs to the west and southwest of Villa deSantiago. The two caves which have been studied areformed in Cretaceous limestone. Sotano de la Anticlinais a large single-room pit, while Sotano de PotreroRedondo is a vertical cave with some horizontaldevelopment at the bottom.The only troglobite among the six species recordedfrom the region is the nesticid spider Nesticus nahuanusGertsch, which is also known from caves to thenorth. The troglophile fauna includes the endemicoctochaetid earthworm Trigaster albida Gates andmillipeds of the rhachodesmid genus Strongylodesmusand sphaeriodesmid genus Sphaeriodesmus.Sierra del Fraile, Nuevo Leon.-The Sierra delFraile is formed almost entirely of vertical-beddedCupido Limestone of lower Cretaceous age (Russelland Raines, 1967). The only cave which has beenstudied is the spectacular commercial cave, Grutas deVilla de Garcia. It has been much modified by commercialization,and recent attempts at collection havebeen largely unproductive. It is located at an elevationof 1,060 meters. A second cave, Cueva del Rinconde la Virgen, houses a large colony of Mexicanfreetail bats, Tadarida brasiliensis mexicana (Saussure),but has not been studied for its invertebratefauna.Of the 13 species of invertebrate known fromGrutas de Villa de Garcia, six are troglobites: theleptonetid spider Leptoneta isolata Gertsch, theopilionid Hoplobunus osorioi (Goodnight and Goodnight),an undetermined rhagidiid mite, the lithobiidcentipede Garcibius osorioi Chamberlin, and thecampodeid diplurans Parallocampa (Parallocampa)cavemicola Wygodzinsky and Paratachycampa bonetiWygodzinsky. Of these species, L. isolata is clearlymost closely related to species known from the SierraMadre Oriental to the south. Hoplobunus osorioioccurs also in Grutas del Palmito and Gruta del Carrizalto the north. The other species are of uncertainaffinities. The troglophile fauna includes the pholcidspiders Metagonia serena Gertsch, Modisimus pusillusGertsch, and Physocyclus hoogstraali Gertsch andDavis; the scytodid spider Loxosceles bolivari Gertsch;and the carabid beetle Platynus pelaezi (Bolivar andHendrichs).Sierra de Garia, Nuevo Leon.-This range is locatedin western Nuevo Leon near the town of Espinazo.The only cave which has been investigated is Cuevade Constantin, a single-room cave which is heavilydecorated with speleothems. The fauna recordedfrom the cave includes undetermined reduviid hemipteransand crane flies.Sierra de Gomas, Nuevo Leon.-The Sierra deGomas is a north-south range located south of thetown of Bustamante. It is composed to a large extentof the massive Cupido Limestone of Cretaceous age(Fuentes, 1964; Russell and Raines, 1967). Althoughcollections have been made in four caves in the area,the only cave which has been well studied is Grutasdel Palmito. This cave, now commercialized, isamong the larger and more beautiful in Mexico. Thesmall entrance is located about 500 meters above thefloor of the plain. From the large entrance chambera steep breakdown-floored slope descends into achamber about 100 meters wide, 600 meters long,and up to 30 meters high. Total depth of the cave isabout 220 meters, making it one of the deeper cavesin northern Mexico.The fauna of the cave is very rich, with 10 troglobitesbeing represented. The troglobite fauna, withone possible exception, is closely related to the faunaof the Sierra Madre Oriental. Troglobites include thetrichoniscid isopods Brackenridgia palmitensis (Mulaik)and Cylindroniscus cavicolus (Mulaik); the hyidpseudoscorpion Leucohya heteropoda Chamberlin;the opilionid lloplobunus osorioi (Goodnight andGoodnight); the rhachodesmid milliped Ceuthauxuspalmitonus Chamberlin; the trichopolydesmid millipedSpeodesmus sp.; the collembolans Oncopoduraprietoi Bonet and Pararrhopalites anops Bonet andTellez; and the trechine beetle Mexaphaenops prietoiBolivar. This is the northern limit of the range for thegenera Cylindroniscus, Ceuthauxus, Pararrhopalites,and Mexaphaenops. Hoplobunus reaches its northernlimit in Texas, where it is represented only by tworelict troglobites. Brackenridgia and Oncopodurarange from southern Mexico well into the UnitedStates. Speodesmus is apparently a relict of a morenorthern distribution. It is well represented in thecaves of Texas and New Mexico, but only S. peckiShear in the Sierra de El Abra occurs south of Grutasdel Palmito.The troglophile fauna includes widespread species,such as the nesticid spider Gaucelmus augustinusKeyserling which ranges from the United States intosouthern Mexico. A few species are typical of thecave fauna of the Sierra Madre Oriental, such as themysmenid spider Maymena chica Gertsch; the pholcidspider Me tagonia candela Gertsch; the opilionid Pellobunusmexicanus Goodnight and Goodnight; thescolopendrid centipede Newportia pelaezi Chamberlin;the collembolan Acherontiella sabina Bonet; and28

the gryllid cricket Paracophus subapterus Chopard.Two species are temperate in their affinities: thecarabid beetle Rhadine araizai araizai (Bolivar) andthe leiodid beetle Ptomaphagus (Adelops) cavernicolacavernicola Schwarz.Sierra de Iguana, Nuevo Leon.-The Sierra deIguana is a north-south trending range formed of Cretaceouslimestone and located north of the highwayfrom Sabinas Hidalgo to Villaldama. The only cavethat has been investigated in this area is Cueva delDiablo, a dry horizontal cave inhabited by the batMormoops megalophylla megalophylla Peters. Theonly troglophile known is the pholcid spider Physocyclushoogstraali Gertsch and Davis.Sierra de Santa Clara, Nuevo Leon.-The Sierra deSanta Clara is a range of Cretaceous limestone locatedsouth of the Sierra de Iguana (Fuentes, 1964). Onlyone cave is known, Cueva de las Fisuras; this is aseries of dry, fissure-like passages in the northerncliffs. The only species identified from the cave is anundescribed genus and species of rhaphidophoridcricket.Sierra de la Silla, Nuevo Le&n.-The Sierra de laSilla is a mountain range located south-southwest ofMonterrey. The only cave which has been studied inthis region is Cueva de la Boca, at an elevation of 540meters. It is a large horizontal passage developed inthe Cupido Limestone and is now being mined forphosphates (Russell and Raines, 1967). A small mineadit, now partially flooded, located about 40 metersbelow Cueva de la Boca has also been studied.The mine adit was found to contain two species oftroglobitic isopods, the cirolanid Sphaerolana a/finisCole and Minckley and an undescribed species of thestenasellid genus Mexistenasellus. The former speciesis also known from springs and natural wells atCuatro Cienegas de Carranza, Coahuila. The genusMexistenasellus, also known from Cuatro Cienegas deCarranza, ranges into Veracruz. Two of the 32 terrestrialspecies recorded from Cueva de la Boca aretroglobites: the nesticid spider Nesticus nahuanusGertsch and the lithobiid centipede Nuevobiuscavicolens Chamberlin. Nesticus nahuanus is alsoknown from other nearby parts of Nuevo Leon. Thecentipede is of particular interest in that the onlyother species in the genus occurs in a cave in Tennesseein the United States.The troglophile fauna includes principally elementstypical of the Sierra Madre Oriental. Of noteare the pholcid spiders Metagonia placida Gertsch andModisimus rainesi Gertsch; the collem bolans Pseudosinellasp. and Acherontiella sabina Bonet; the carabidbeetle Platynus boneti (Bolivar and Hendrichs); andthe staphylinid beetle Stilicolina n. sp. The leiodidbeetle Ptomaphagus (Adelops) cavernicola aditusPeck is an endemic subspecies belonging to a speciesranging from the United States into Nuevo Leon. Theremaining fauna is closely associated with the guanoof Mexican freetail bats, Tadarida brasiliensis mexicana(Saussure), a large colony of which inhabits thecave.Sierra El Tarillal. Nuevo Leon.-The Sierra £1Tarillal is a range of Cretaceous limestone mountainslocated about 20 kilometers east of Arteaga. Thegeology of this region has been discussed by deCserna (1956). The only cave which has been investigatedis Cueva de las Vigas at an elevation of 2,300meters. This 100-meter-long cave has been muchmodified by mining activities. The fauna is typical ofthat of the northern Sierra Madre Oriental and includesfive species of troglophile: the agelenid spidersCicurina sp. and Tegenaria ?gertschi Roth, the pholcidspiders Coryssocnemis simoni O. P.-Cambridgeand Metagonia coahuila Gertsch, and undeterminedstaphylinid beetles.Zaragoza, Nuevo Leon.-This is an area of Jurassicgypsum deposits located along the western slopes ofthe Sierra Madre Oriental near the town of Zaragoza.Three species, two of which are troglophiles, areknown from the three small caves which have beenvisited. The troglophiles are ptilodactylid beetles ofthe genus Ptilodactyla and staphylinid beetles of thegenus Belonuchus.Purificacion. Nuevo Leon and Tamaulipas.-Thismajor karst region is located west of the town of £1Barretal. Caves are known from massive Cretaceouslimestone of the TamauJipas Formation at elevationsfrom 1,100 to 2,200 meters. Much of the drainage issubterranean with several caves receiving considerablefloodwater. Surface karst features include sinkholes,dolinas, pinnacles, and karren. Many of the caves aresmall single-room pits, but others are quite extensive.The two largest caves in the region are S6tano de lasCalenturas and Sistema Purificacion. The first receivesthe floodwater of a large arroyo and contains largeamounts of organic debris and a small active stream.Sistema Purificacion is a complex, multi-level cavewith several active streams. The cave is the longestin Mexico; more than 35 kilometers have been surveyedto date. It has a vertical range of almost 900meters, the highest entrance being at 1,980 metersand the lowest at 1,100 meters. Pate (1979) andTreacy (1979) have described several of the caves inthe region.About 100 species, of which 19 are troglobites,have been identified from the 33 caves which havebeen studied but much of the material remains undeterminedor undescribed. The aquatic fauna includes29

three undescribed species of isopod: a cirolanidbelonging to the genus Speocirolana, an asellid of thegenus Caecidotea, and a stenasellid of the genusMexistenasellus. The terrestrial troglobite fauna includesan undescribed genus and species of chactidscorpion; a second species of this genus is knownfrom the San Nicolas de los Montes region. Threeundescribed species of pseudoscorpion have beenfound; these belong to the genera Aphrastochchonius,Typhloroncus, and Paravachonium. The troglobiticspider fauna includes a tarantula of the genusSchizopelma and the leptonetid Leptoneta isolataGertsch. The latter species is also known from Gru tasde Villa de Garcia, Nuevo Leon. Other troglobites, allundescribed endemics, include an opilionid of thegenus lloplobunus, a scolopendrid centipede of thegenus Newportia, several as yet undeternlined milli·peds, and three species of carabid beetle of the genusMexaphaenops. The highly cave-adapted leiodidbeetle, Ptomaphagus (Adelops) mckenziei Peck, isknown only from caves in this region.The troglophile fauna includes undescribed spidersof the genera Euagrus, Cicurina, Tegenaria, Nesticus,and Ctenus. The spiders Eidmannella pallida (Emerton),Coryssocnemis abernathyi Gertsch, Modisimusrainesi Gertsch, and M. reddelli Gertsch are abundantin the caves of the region. The troglophile millipedfauna, though not well studied, includes species ofthe genera Cleidogona, Myrmecodesmus, Strongylodesmus,and Sphaeriodesmus. Other troglophiles ofinterest include the collembolans Pseudosinellareddelli Christiansen and Pseudosinella sp., the carabidbeetle Platynus (iHexisphodrus) n. sp., and theleiodid beetle Dissochaetus aztecus Szymczakowski.Both the troglobite and troglophile fauna is typicalof that of the Sierra Madre Oriental.Cuetzalan, Puebla.-This is one of the major karstregions in Mexico but remains poorly known. Theregion is bounded on the south by the Rio Apulcoand on the north by the Rio Tecuantepec. It extendsfrom near Cuetzalan on the east to Zapotitlan on thewest Elevations range frolll about 250 meters tomore than 2,000 meters. Drainage is almost entirelysubterranean with floodwaters entering many caves;active streams running from the higher elevationsfrequently enter sumidero-type cave entrances.Springs are numerous along the banks of the RioTecuantepec. Caves tend generally to be large tnmkchannels containing active streams (Davis, 1974;I{eddell, 1974: Sprouse, 1979). Several caves are long,with one surveyed for more than 10 kilometers.The fauna of the area is not yet well known systematically,and the 87 species determined from the 20investigated caves are only a small percentage ofthose which will eventually be known. The only aquatictroglobite from the region is the widespread cirolanidisopod Speocirolana pelaezi (Bolivar). The onlyterrestrial troglobite thus far described is the endemicspirobolellid milliped Reddellobus troglobius Causey.Other troglobites include an ideoroncid pseudoscorpion,a schizomid of the genus Schizomus, anamblypygid of the genus Paraphrynus, a tarantula ofthe genus Schizopelma, a leptonetid spider of thegenus Leptoneta, a pholcid spider of the genus Metagonia,a glomerid milliped of the genus Glomeroides,an undescribed genus and species of the millipedfamily Trichopolydesmidae, and a collembolan of thegenus Acherontides.The streams in the caves of this region su pport anabundance of troglophilic crayfish, including Procambarus(Paracambarus) ortmanni (Villalobos), P.(Villalobosus) n. sp., and P. (V.) xochitlanae lIobbs.Terrestrial troglophiles of interest include a vaejovidscorpion of the genus Vaejovis, a species of the amblypygidgenus Paraphrynus, an agelenid spider ofthe genus Tegenaria, a ctenid spider of the genusCtenus, a mysmenid spider of the genus Maymena,the nesticid spiders Eidmannella pallida (Emerton)and Gaucelmus calidus Gertsch, pholcid spiders ofthe genera Coryssocnemis and Pholcophora, the uloboridspider Philoponella signatella (Roewer), a cleidogonidmilliped of the genus Cleidogona, a pyrgodesmidmilliped of the genus Myrmecodesmus, arhachodesmid milliped of the genus Strongylodesmus,a sphaeriodesmid milliped of the genus Sphaeriodesmus,a collernbolan of the genus Pseudosinella,the psocid Psyllipsocus ramburii Selys-Longchamps,cixiid homopterans, an undescribed species of carabidbeetle of the genus Platynus (Mexisphodrus), andstaphylinid beetles of the genera Belonuchus andllomaeotarsus. The fauna of the region appears to betypical of that of the Sierra Madre Oriental.Xicotepec. Puebla.-This region is located near thetown of Xicotepec; it is poorly known and defined.Biological records are available for four caves in thisarea, but only two have been visited by AMCS members.The latter caves are small horizontal passages ina cliff face. No troglobites are known from this region,and of the 11 species recorded only three are probabletroglophiles: the amJadiIIid isopod Venezillollamasi Rioja, the nesticid spider Eidmannella pal/ida(Emerton), and the psocid Psyllipsocus ramburiiSelys-Longchamps.Ahuacatlan, Queretaro.-This region is located inone of the western ranges of the Sierra Madre Orientaljust west of the town of Jalpan. All of the cavesknown from the region are formed in the EI Abra(=EI Doctor) Limestone. The geology of the region30

has been Jiscussed by Segerstrom (1961) and Smith(1972). Karst development is represented by largedolinas, sinkholes, and lapiez. The Rio J alpan hasbeen captured by a large cave and now runs through anatural tunnel (known as Puente de Dios) for 1.5kilometers. Most of the caves are vertical pits, withthe Sotanito de Ahuacatlan attaining a depth of320 meters.This region has been little studied biologically;only nine caves have received attention. The onlytroglobite among the 17 species identified is thecixiid homopteran Cixius orcus Fennah. The troglophilefauna includes the diplurid spider Euagrusluteus Gertsch, a ctenid spider of the genus Ctenus,a pholcid spider of the genus Modisimus, a scytodidspider of the genus Loxosceles, a rhachodesmid millipedof the genus Strongylodesmus, campodeid diplurans,the gryllid cricket Paracophus placonotusHubbell, the carabid beetle Platynus acuminatus(Chevrolat), and a leiodid beetle of the genus Ptomaphagus(Adelops). All of these species are typicalof the cave fauna of the Sierra Madre Oriental to theeast.Laguna Colorado, Queretaro.-This region is thewestern extension of the karst region of Xilitla, SanLuis Potosi. All of the caves investigated are formedin the EI Abra (=EI Doctor) Limestone. Karst developmentis not as great as near Xilitla, but large shallowdolinas and numerous sinkholes occur. Bonet (1953a)has discussed the regional geology and physiographyand includes a map of the largest cave in the region,Cueva del Madrofio. The elevation of this cave is1,810 meters. The remaining four caves studied aresmall.The only two troglobites thus far recorded are the()pilionid Hoplobunus queretarius Silhavy and therhachodesmid milliped Unculabes arganoi Shear. Bothare closely related to other species in the Sierra MadreOriental. Although 29 other species have been recordedfrom the region, many are mites identifiedonly to family. The determined troglophile fauna includessuch typical elements of this general area asthe leptonetid spider Leptoneta sp., the pholcidspider Metagonia maximiliani Brignoli, the cleidogonidmilliped Cleidogona sp., the rhachodesmidmilliped Strongylodesmus sp., the collembolan Lepidocyrtussp., the gryllid cricket Paracophus placonotusHubbell, the carabid beetle Platynus (Mexisphodrus)sp., and the Iciodid beetle Ptomaphagus (Adelops)leo Peck.Pinal de Amoles, Queretaro.-This region is locatedimmediately west of the Ahuacatlan region and consistsof an extensive area of Cretaceous limestone ofthe El Abra (=EI Doctor) Formation. A geologicalmap of the region is included in Segerstrom (1961).Karst development includes many large dolinas,sinkholes, and lapiez. Most of the caves investigatedhave been rather small, but Sotano del Buque is1,149 meters long and attains a depth of 506 meters(Jefferys, 1979) Although 21 caves have beenstudied, the region remains largely unknown.A total of 46 species have been identified from thecaves of the region. Of these, six are troglobites: theagelenid spider Tegenaria caverna Gertsch, the leptonetidspider Leptoneta delicata Gertsch, the trichopetalidmilliped Mexiterpes metallicus Shear, therhachodesmid milliped Strongylodesmus sp., andthe carabid beetles Mexaphaenops elegans Barr andParatrechus (Hygroduvalius) pallescens Barr. All ofthese species are most closely related to species knownfrom other parts of the Sierra Madre Oriental. Troglophilesof interest include the diplurid spider Euagrusluteus Gertsch; an agelenid spider of the genus Tegenaria;a ctenid spider of the genus Ctenus; the nesticidspiders Eidmannella pallida (Emerton), Gaucelmusaugustinus Keyserling, and Nesticus vasquezaeGertsch; the pholcid spiders Coryssocnemis ivieiGertsch, C. simoni O. P.-Cambridge, and Physocyclusreddelli Gertsch; the scytodid spider Loxoscelesaranea Gertsch; the phalangodid harvestman Pellobunusmexicanus Goodnight and Goodnight; thecleidogonid millipeds Cleidogona mayapec Shearand C. totonaca Shear; a rhachodesmid milliped ofthe genus Strongylodesmus; the gryllid cricket Paracophusplaconotus Hubbell; the carabid beetlesPlatynus tlamayaensis (Barr) and P. (Mexisphodrus)sp.; and staphylinid beetles of the genera Belonuchusand Philonthus.San Joaquin, Queretaro.-This region is locatedsouth of the Pinal de Amoles region. The geology hasbeen studied by Wilson et al. (1955). All of the cavesknown are formed in the EI Abra (=EI Doctor) Limestone.Karst development includes dolinas, sinkholes,and lapiez. Much of the drainage is internal. The elevationis about 2,400 meters.This region remains essentially unstudied; onlythree caves have been investigated. Of the three specieswhich have been identified, two spiders are troglophiles:the pholcid Coryssocnemis simoni O. P.Cambridge and an undescribed species of nesticid ofthe genus Nesticus.San Juan, Queretaro.-Only seven caves have beenbiologically examined in this region, which is locatednorth of the Ahuacatlan region. Stone and Jameson(1977) have published a comprehensive report on theregion, including descriptions of all caves and a generalstudy of the geology. All of the caves known are31

fonned in Cretaceous limestone of the EI Abra Formation.The large caves are characterized by considerablevertical development with two (Hoya de lasConchas and Sotano de Nogal) being more than 500meters deep. Elevations range from 1,000 to 1,500meters, with most of the caves occurring at about1,400 meters. Drainage is almost entirely internal andmany of the caves receive considerable runoff.Of the 18 species detennined from the caves of theregion, the only troglobite is an unidentified isopodof the family Trichoniscidae. The remainder of thefauna is typical of the Sierra Madre Oriental and includesthe nesticid spider Eidmannella pallida (Emerton),the opilionid Hoplobunus sp., the centipedeNewportia n. sp., the collembolan Pseudosinellareddelli Christiansen, and the gryllid cricket Paracophussp.Tancoyol, Queretaro.-This region is located immediatelywest of the Xilitla Plateau. Little is knownabout the region, but karst development includeslarge dolinas and sinkholes. The largest cave in theregion is the largely vertical EI Socavon, whichattains a depth of about 200 meters.Only five species have been reported from thethree caves that have been biologically investigated.The only troglobite is the opilionid Hoplobunusqueretarius Silhavy, also known from the LagunaColorado region to the south. Troglophiles includean agelenid spider of the genus Tegenaria, a ctenidspider of the genus Ctenus, and a pholcid spider ofthe genus Physocyclus.Xilitla Plateau, Queretaro and San Luis Potosi.ThiS region includes an area of high elevation karstlying north of Xilitla and including parts of the statesof Queretaro and San Luis PotosI. No collectionshave yet been made in the San Luis Potosi part ofthe region. Fish (1978; 1979) has discussed thegeology, physiography, and caves of the region. Thecaves are fonned in the Cretaceous El Abra Limestone,and there is extensive karst development, withdolinas, sinkholes, pinnacles, and lapiez present.Elevations range from about 1,500 to 2,900 meters.Although a few deep caves occur, such as the 559meter-deep Sotano de Trinidad, the three caves whichhave been biologically investigated are relatively horizontal.Much of the fauna from this region awaits study,but 19 species, of which four are troglobites, havebeen identified. The troglobites include trichoniscidisopods, the diplurid spider Euagrus n. sp., the opilionidHoplobunus sp., and the rhachodesmid millipedUnculabes crispus Causey. The last species is alsoknown from the Xilitla region to the south, and theothers have their closest affinities with that region.The troglophile fauna includes flatwonns of thegenus Dugesia; the agelenid spider Tegenaria selvaRoth; the nesticid spiders Eidmannella pallida (Emerton),Gaucelmus augustinus Keyserling, and two undescribedspecies of the genus Nesticus; the opilionidKaros depressus Goodnight and Goodnight; and thecarabid beetle Platynus (Mexisphodrus) sp.Aquismon, San Luis Potosi.-This region is locatedwest of the town of Aquism6n and extends from theXilitla Plateau north for about 20 kilometers; it isbounded on the west by the Rio Santa Maria. Elevationsrange from about 400 to 1,000 meters. Karstdevelopment in the region is spectacular in places,with large deep dolinas, huge sinkholes, karren, andhaystack hills. Most of the drainage is subterraneanand several caves receive considerable floodwater.Caves range from small pits and shallow horizontalpassages to enonnous vertical shafts such as Hoya delas Guaguas and Sotano de las Golondrinas. The principallimestoneunit in the region is the EI Abra (=EIDoctor) Fonnation. Raines (1968) includes a discussionof the geology and physiography.A total of 57 species, of which 10 are troglobites,has been reported from the 15 caves studied. Theonly aquatic troglobites are the cirolanid isopodsSpeocirolana bolivari (Rioja) and S. pelaezi (Bolivar).Terrestrial troglobites, all of which are typical elementsof this general area, include the trichoniscidisopod Mexiconiscus laevis (Rioja); the amblypygidParaphrynus velmae Mullinex; the opilionid Hoplobunusplanus Goodnight and Goodnight; the glomeridmilliped Glomeroides sp.; the trichopetalidmilliped Mexiterpes fishi (Causey); the rhachodesmidmilliped Unculabes criJpus Causey; the cambalidmilliped Mexicambala russelli Causey; and the gryllidcricket Paracophus cladonotus Hubbell. Except forH. planus and M. fishi all of these species are alsoknown from the Xilitla region.The troglophile fauna includes flatwonns of thegenus Dugesia; the crayfish Procambarus (Ortmannicus)toltecae Hobbs; the spiders Ctenus mitchelliGertsch, Gaucelmus sp., and Metagonia sp.; therhachodesmid millipeds Strongylodesmus sp. andUnculabes columbinus Causey; the collembolanPseudosinella sp.; the gryllid cricket Paracophusplaconotus Hubbell; carabid beetles of the generaArdistomis and Tachys; and the leiodid beetles Dissochaetussp. and Proptomaphaginus microps Peck.Cerro de la Cochina, San Luis Potosi.-This is asmall limestone range located west of Matehuala. Theonly cave studied is Cueva del Cochino, a dry horizontalcave located at an elevation of 1,560 meters.No troglobites are known from the region and the32

only apparent troglophile is the psocid Psyllipsocusramburii Selys-Longchamps.La Libertad, San Luis Potosi.-This is an area ofdesert located west of Ciudad del Maiz at an elevationof about 1,000 meters. The four caves investigatedare all shallow, dry sinkholes formed in Jurassicgypsum. No troglobites are known from this regionand only 12 species have been identified. Of these,four are presumed troglophiles: the diplurid spiderEuagrus sp., the pholcid spider Physocyclus reddelliGertsch, the carabid beetle Agonum sp., and thestaphylinid beetle Stilicolina n. sp. The fauna istypical of that of the drier areas of northern Mexico.Matehuala, San Luis Potosi.-This region consistsof an area of karst developed on ] urassic gypsum.Most of the drainage is internal, with runoff enteringthrough shallow gypsum sinks. The two caves investigatedin the region, Sumidero de Matehuala at anelevation of 1,500 meters and Sumidero 552 at anelevation of 1,330 meters, are both small.Eight species have been determined from thisregion, none of which is troglobitic. The troglophilefauna includes the pholcid spider Physocyclus merusGertsch, carabid beetles of the genus Rhadine, andptilodactylid beetles of the genus Ptilodactyla. Thefauna resembles that of other arid regions to the westof the Sierra Madre Oriental.Micos, San Luis Potosi.-This region is locatednear the town of Micos in a series of north-southtrending ranges. The seven caves that have beenvisited are located at the base of the range at an elevationof about 250 meters. Several of these caves receivefloodwater. Mitchell et al. (1977) discuss thegeneral geology and physiography of the region andinclude descriptions of some of the caves.The 30 species known from this region include sixaquatic troglobites: the entocytherid ostracod Sphaeromicolacoahuiltecae Hobbs and Hobbs, the cirolanidisopod Mexilana saluposi Bowman, the stenasellidisopods Mexistenasellus parzeJalii Magniezand M. wilkensi Magniez, the anthurid isopod Cyathuran. sp., and the blind characin Astyanax jordani(Hubbs and Innes). The aquatic fauna is quite differentfrom that of other parts of the general region.With the exception of A. jordani all of these speciesare endemic to this region. Terrestrial troglophiles, onthe other hand, are typical of the Sierra Madre Oriental.Included are ctenid spiders of the genus Ctenus,the nesticid spiders Eidmannella pallida (Emerton)and Gaucelmus augustinus Keyserling, the scytodidspider Loxosceles valdosa Gertsch, the opilionidKaros parous Goodnight and Goodnight, the pyrgodesmidmilliped Myrmecodesmus monasticus (Causey),the gryllid cricket Paracophus placoltotus Hubbell,alleculid heetles of the genus Lystronychus, andstaphy-linid heetles of the genus Homae0 tarsus.Piedra Paloma, San Luis Potosi.-This is a limitedarea of gypsum karst located east of Ciudad del Maiz.The most notable cave known from the region isSumidero de Piedra Paloma, a largely vertical caveinto which much floodwater runs. The only troglophileknown from the region is the theridiid spiderAchaearanea porteri Banks.Puente de Dios, San Luis Potosi.-This is a poorlydefined region located southwest of Ciudad Valles.The Puente de Dios is a cave passage through whichthe Rio Santa Maria runs for a short distance. Theonly cave investigated in the region is a small onelocated along the hanks of the river.1\0 troglohites are known from this region; troglophilesinclude the pholcid spiders Modisimus bonetiGertsch and Physocyclus sp., the scytodid spiderLoxosceles sp., the rhachodesmid milliped Strongylodesmussp., the gryllid cricket Paracophus placonotusHubbell, and the leiodid beetle Ptomaphagus (Adelops)elabra Peck.Rayon, San Luis Potosi.-This ill-defined region islocated near the town of Rayon on the highway fromCiudad Valles to San Luis Potosi. The only cave thathas been studied is a small horizontal resurgenceknown as Cueva del Agua.Ten species, none troglohitic, have been identifiedfrom the cave. These include several troglophiles, alltypical of the Sierra Madre Oriental. The crayfishProcambarus (Ortmannicus) villalobosi Hohhs isknown only from here and from one cave in the SanNicolas de los Montes region. Terrestrial troglophilesinclude spiders of the genera Ctenus and Maymena,the gryllid cricket Paracophus placonotus Huhhell,the leiodid beetles Dissochaetus aztecus Szymczakowskiand D. hetschkoi Reitter, and staphylinidbeetles of the genus Belonuchus.San Nicolas de los Montes, San Luis Potosi.-Thisregion is located northwest of Ciudad Valles near thetown of San Nicolas de los Montes. Elevations rangefrom ahout 600 to 1,700 meters, with most of theknown caves lying at about 900 meters. Although afew arroyos drain part of the region, most drainage isinternal. The water level is generally shallow andseveral small lakes occur, the most notahle of whichis Laguna Grande. Karst development is characterizedhy numerous small sinkholes and shallow depressions.All of the 16 caves studied are small pits or shallowhorizontal caves; many contain water.The aquatic fauna includes three troglobites: theentocytherid ostracod Sphaeromicola cirolanae Riojaand the cirolanid isopods Speocirolana n. sp. and S.pelaezi (Bolivar). The only terrestrial troglohite is an33

undescribed genus and species of chactid scorpion; asecond species of the same genus is known from thePurificacion region to the north. The troglophilefauna is typical of the general region and includes thecrayfish Procambarus (Ortmannicus) villalobosiHobbs, the amblypy¢d Paraphrynus pococki l\Iullinex.the ctenid spider Ctenus mitchelli Gertsch, thenesticid spiders Eidmannella pallida (Emerton) andGaucelmus augustinus Keyserling, the opilionidsKaros gratiosus Goodnight and Goodnight and K.projectus Goodnight and Goodnight, and the gryllidcricket Paracophus placonotus Hubbell.Sierra de Alvarez, San Luis Potosl.-This is one ofthe westernmost ranges of the Sierra Madre Orie~taland is located east of the city of San Luis PotosI.Karst development is considerablt~, with most drainagebeing subterranean. Karst forms include dolinas,sinking streams, lapiez, and pinnacles. The greatestdevelopment is in an area known as Valle de los Fantasmas,where numerous pinnacles brive an eerieappearance during times of fog. The caves arc formedin the El Abra (=El Doctor) Limestone. Cserna andBello-Barradas (1963) have studied the regionalgeology. Brief descriptions of the caves and a checklistof the fauna are in Elliott and Reddell (1973).There have been 26 caves investigated biologically;these range in elevation from 2,100 to 2,350 meters.This is one of the hetter-known regions in Mexicowith respect to its invertebrate cave fauna, with 95species having been identified. Of these, only two aretroglobites: the trichopetalid milliped JIJexiterpesegeo (Causey) and the trechine beetle Mexaphaenopsfishi Barr. Both belong to genera known from otherparts of the Sierra Madre Oriental. The rarity of troglobitesin this high altitude region is surprising, butmay be a reflection of greater aridity and less diverseflora. The troglophile fauna is also typical of theSierra Madre Oriental. It includes the tarantulaSchizopelma elliotti Gertsch; the agelenid spiderTegenaria selva Roth; the mysmenid spider Maymenachica Gertsch; the nesticid spider Eidmannella pallida(Emerton); the pholcid spiders Coryssocnemis abernathyiGertsch, Metagonia punctata Gertsch, andPsilochorus concinnus Gertsch; the opilionids Cynortajamesoni Goodnight and Goodnight and Karosparous Goodnight and Goodnight; millipeds of thegenera Myrmecodesmus, Strongylodesmus, and Tiphallus;the collembolan Pseudosinella reddelliChristiansen: and carabid beetles of the generaBembidion, Platynus, and Platynus (Mexisphodrus).Sierra EI Pino, San Luis Potosi.-This region is amountain range east of Ciudad del Maiz with elevationsfrom about 1,100 to 1,900 meters. The regionremains poorly knvwn, and only six caves have beenvisited. Large sinkholes are known on top of therange but are unvisited. Water falling on the westernside of the range drains into a closed lake or intocaves located in a wide, shallow arroyo. Two of thesecaves and a few on the slopes of the mountain havebeen studied.The fauna of the region includes two aquatic andthree terrestrial troglobites: the entocytherid ostracodSphaeromicola cirolanae Rioja, the cirolanidisopod Speocirolana pelaezi (Bolivar), a trichopolydesmidmilliped, the collembolan Pseudosinellapetrustrinatii Christiansen. and the gryllid cricketParacophus lippus Hubbell. All are typical representativesof the fauna of the Sierra Madre Oriental.The troglophile fauna includes the ctenid spiderCtenus mitchelli Gertsch, the gryllid cricket Paracophusn. sp., the scydmaenid beetle Euconnus(Napochus) sp., the scarabaeid beetle Onthoplwguscuevensis II owden, and staphylinid beetles of thegenus Belonuchus.Xilitla, San Luis Potosi.-The Xilitla region is oneof the great karst regions in ~orth America. l\lassivereef deposits of Cretaceous EI Ahra (=EI Doctor)Limestone are highly cavernous, and numerous majorcaves and pits ex ist in the region. As considered inthis report, the region ex tends west from the easternface of the Sierra Madre Oriental to near the SanLuis Potosi-Queretaro border just past the town ofAhuacatLin. It is bordered on the north by the hill:hXilitla Plateau, on the west by the Laguna Coloradoregion, and on the south by the Rio .\Ioctezuma.Elevations range from about 100 to ahout 1,200meters. Much of the drainage is subterranean, withwater emerging from large springs, sOlTle vauclusian innature, along the east face of the range. Karst developmentis frequently intense with large dolinas, sinkholes,lapiez, and pinnacles present. Although severallarge caves exist, drainage does not appear to he ashighly integrated as in the Purificacion region, and nosystems to rival that of the Sistema Puri ficaciilll arclikely. In addition to numerous vertical shafts, somemore than 100 meters in depth, large chambers andvertical cave systems exist. Among the more importantcaves are Sotano de Huitzmolotitla with morethan three kilometers of passage and Sotano de rlamayawith a vertical depth of about -t.00 meters.Carrasco (1970) has discussed the stratigraphy of thegeneral area and concluded that the name El Ahrashould be applied to the limestone of the area. Ronet(1953a) has discussed the geology and physiography,and includes maps and descriptions of many of thecaves. Russell and Raines (1967) add infonnation ongeology and descnbe additional caves.34

The cave fauna of this region is rich and, comparedto many other parts of Mexico, well known. Collectionshave been made in 31 caves and 163 specieshave been identified, of which 18 are troglobites.The fauna, both troglobitic and troglophilic, is typicalof that of the Sierra Madre Oriental. The only aquatictroglobite known from the region is the endemicacanthodrilid earthworm Eodrilus mexicanus Gates.The terrestrial troglohite fauna includes nine endemicspecies: the squamiferid isopod 1'richorhina bonetiRioja, the chthoniid pseudoscorpion Tyrannochthoniuspallidus J\luchmore, the protoschizomid Agastoschizomushuitzmolotitlensis Rowland. the dipluridspider Euagrus anops Certsch, the pholcid spiderMetagonia tlamaya Gertsch, the glomerid millipedGlomeroides caecus Causey, till:' rhachodesrnid millipedUnculabes porrensis Shear, the trichopolydesmidmilliped Tylogoneus rainesi Causey, and a new speciesof the collembolan genus Smintlwrus. The remainingspecies of troglobite are all restricted to this and contiguousregions. They include the trichoniscid isopodiVlexiconiscus laevis (Rioja), the amblypygid Paraphrynusvelmae J\lullinex, the tarantula Schizopelmastygia (Gertsch), the rhachodesmid milliped Unculabescrispus Causey, the cambalirl milliped Mexicambalarusselli Causey, the collem bolans Pseudosine/lasp. and Acheron tides potosinus Bonet, andthe gryllirl cricket Paracophus cladonotus Hubbell.Among the llIorc notable troglophiles are the crayfishProcambarus (Ortmannicus) toltecae Hobbs: theagelenirl spiders Tegenaria decora Gertsch and T. selvaRoth: the denid spirler Ctenus mitchelli Gertsch: thenesticid spiders Eidmannella pallida (Emerton) andGaucelmus calidus (;crtsch: the. phokid spiders Coryssocnemisiviei Gntseh and .lIe/agonia arnica Gertsch;the opilionirls Aarris dybasi (Goodnight and Goodnight),K. gratio.ms Coodnight and Goodnight, andK. projectus Goodnight anrl (;oodnight: the c1eidogonidlllilliped Cleidopma sp.: the parado"osolllatidlllilliped Oxidus gracilis (I-.:o

(Villalobos). Endemic aquatic troglobites include thediaptomid copepod Diaptomus (Microdiaptomus)cokeri Osorio Tafall, the microcerberid isopod Microcerberusn. sp., and the palaemonid shrimp Troglocubanusperezfarfanteae Villalobos. The terrestrialtroglobite fauna includes the rhagidiid mite Rhagidiaweyerensis (Packard), also known from caves in theUnited States. Other species known from otherregions include the squamiferid isopod Spherarmadillocavernicola Mulaik, the trichoniscid isopodBrackenridgia bridgesi (Van i\arne), the opilionidHoplobunus boneti (Goodnight and Goodnight), thescolopendrid centipede Newportia sabina Chamherlin,and the collembolan Pseudosinella petrustrinatiiChristiansen. Endemic troglobites are the trichoniscidisopod Cylindroniscus vallesensis Schultz, thechactid scorpion Typhlochactas elliotti !\Iitchell, thechthoniid pseudoscorpions Aphrastochthonius pan'usMuchmore and A. russelli \Iuchmore, the vachoniidpseudoscorpions Paravachonium n. sp. and P. bolivariBeier, the protoschizomid Agastoschizomus luciferRowland, the schizomids Schizomus cookei K.owlandand S. mitchelli K.owland. the amhlypygid Paraphryllusbaeops Mullinex, the pholcid spider Metagoniapachona Gertsch, the ricinllleids Cryptocellusn. sp. and C. osorioi Bolivar, the rhagidiid miteRhagidia trisetata Elliott and ~trandtrnann, the trichopetalidmilliped JIexiterpes sabinus Causey, thetrichopolydesmid milliped Speodesmus pecki Shear,an undescribed genus and species of Jlicoletiid thysanuran,the kinnarid homopteran Oeclidius hadesFennah, and the histerid bectle Troglobacanius bolivariVomero.The aquatic troglophile fauna includes several speciesof protozoans, rotifers. ostracods, copepods, thecrayfish Procambarus (Ortmannicus) aeutus cuevachicae(Hobbs), and th~~ characiu Astyanax mexicanus(Phillipi). About 100 terrestrial species arc prohabletroglophiles in the Sierra de EI Ahra. [t is not possiblehere to do more than list some of the more distinct orwidespread species. The arachnid fauna includes theschizomid Schizomus mexicanus I{owland, the amblypygidParaphrynus pococki \Iullinex. the ricinuleidCryptocellus pelaezi Coronado, the opilionidsCynorta jamesoni Goodnight and Goodnight andKaros parous (;oodnight and Goodnight, and the palpigradeKoenenia hanseni Silvestri. Troglophilicspiders include the ctenid Ctenus mitchelli Gertsch;the leptonetid Leptoneta rainesi Gertsch; the mysmenidMaymena chica Gertsch; the nesticids Eidmannellapallida (Emerton) and Eidmannella n. sp.;the pholcids Metagonia pasquinii Brignoli, Metagoniatinaja Gertsch, 1Hodisimus boneti Gertsch, Ph 01cophora elliotti Gertsch, and Physocyclus globosus(Taczanowski); and the scytodid Loxosceles valdosaGertsch. The milliped fauna includes the pyrgodesmidsMyrmecodesmus errabundus (Shear) and M.sabinus (Chamberlin) and the spirostreptid Orthoporusmimus Chamberlin. Apterygote insect troglophilesinclude the campodeid dipluran Campodea(Campodea) chica Wygodzinsky, the japygid dipluranAllojapyx allodontus (Silvestri), amI thecollembolan Acherontiella sabina Bonet. Two speciesof troglophile gryllid cricket are known, Paracophusapterus Chopard and P. placonotus Hubbell. i\umerousspecies of beetle are tro~lophiles in the region.These include the carabid Pachyteles urrutiai Hoi Ivar,the leiodids Dissochaetus aztecus ~zymczakowskiand Ptomaphagus (£ldelops) elabra Peck, scydmaenidsof the genera "Connophron " and Euconnus(Napochus), and staphylinid beetl(~s of the generaBelonuchus, Homaeotarsus, and Stilicolina. Both thetroglophile and troglobite fauna of this region aretypical of the Sierra Madre Oriental, with its closestaffinities lying with the Sierra de Guatemala to thenorth and somewhat less with that of the ranges tothe west and sou th west.Altas Cumbres, Tamaulipas.-This is a poorlydefined region located to the southwest of CiudadVictoria on the highway to Tula. Of the three cavesknown for the region, only two havl~ been investigatedwi th respect to their invertebrate fauna. Bothare small fissure-like caves with a limited fauna. Offour invertebrates identified, th ree are troglophiles:the leptonetid spider /,eptoncta bonita Certsch, thepholeid spider Modisim us reddelli G (~rtsch, and thegryllid cricket Paracophus apterus Chopard. Thesearc typical elements of the cave fauna of the generalregIOn.Sierra de Guatemala, Tamaulipas.-The Sierra deGuatemala is a range located immediat~:ly north ofthe Rio Boqllillas and to the west of the town ofGomez Farias. It is hounded on the north by the RioGuayalejo; on the west it deseends abruptly to theplains east of JaUll1ave. Elevations range from 170 toabout 2,200 meters. Martin (1958) has summarizedthe geology, physiography, vegetation, and dirnateof the region. His study of the reptiles allll amphibiansof the region also includes mudl useful hiogeographicinformation. The principal limestone unitis the EI Abra (= El Doctor) Limestone (Russell andRaines, 1967; Priego de Wit, 1974). Most of thedrainage is internal, with water flowing from twomajor springs, the Nacimiento del Rio Frio to thesouth and the Nacimiento del Rio Sabinas to thenorth. These springs are vauclusian in nature. Cavesare known throughout the area, but most are small.The town of Gomez Farias is built on a small ridge of36

igneous rock immediately east of the Sierra de Guatemala.In the shallow valley separating this ridge andthe main range, several caves receive floodwater viashallow arroyos. These caves are largely vertical andattain depths up to 148 meters below their entrances.Water has been encountered in three of these (Mitchellet al., 1977). Karst development in the Sierra deGuatemala includes numerous dolinas, pinnacles,lapiez, and sinkholes. A few small springs emerge toflow short distances before sinking. Most of the exploredcaves on or near the top of the range are smallsinkholes ranging from a few meters to more than100 meters in depth. Several large chambers and phreaticallyformed passages have been encountered; themost notable of these are Cueva de la Mina, Cueva dela Capilla, and Cueva del InfierniIlo. These welldecoratedcaves usually have permanent pools ofstanding water, occasionally inhabited by aquatictroglobites. Elliott (1973b) has included descriptionsof several caves in the area. Along the northern edgeof the Sierra de Guatemala several significant caveshave been discovered. These include long, horizontalcaves which may represent fossil resurgences. Thelargest cave in the range is Sotano de la Joya de Salas,located on the western side of the range in a largeclosed valley. The cave receives massive amounts ofwet-season floodwater and attains a depth of 376meters. In general, karst development in the Sierrade Guatemala appears to be in a rather early stageand it appears doubtful that a major cave system,such as the Sistema Purificacion, will be found.The Sierra de Guatemala is among the betterknownareas in Mexico with respect to its invertebratefauna. From the 67 caves which have beenstudied, 273 species, including 45 troglobites, areknown. This is largely a reflection of the great rangein elevation and the variety of habitats. The aquatictroglobite fauna includes two flatworms, Dugesiabarbarae Mitchell and Kawakatsu and D. typhlomexicanaMitchell and Kawakatsu. These are closelyrelated to the troglophile D. guatemalensis Mitchelland Kawakatsu, also known from caves in the region.The aquatic earthworm Eodrilus albidus Gates hasbeen found in pools in two caves; it is closely relatedto the troglobitic E. mexicanus Gates from the XiIitiaregion. The microcerberid isopod Mexicerberus troglodytesSchultz is a marine relict known only fromCueva de la Mina. Its affinities are unclear. The remainingaquatic troglobites are also known fromcaves in the Sierra de EI Abra. These are the entocytheridostracod Sphaeromicola cirolanae Rioja, thecirolanid isopods Speocirolana bolivari (Rioja) andS. pelaezi (Bolivar), and the characin Astyanax jordani(Hubbs and Innes). The last species is knownfrom caves at Gomez Farias and from the areaimmediately to the north of Ocampo along thesouthern edge of the range.The terrestrial troglobite fauna is the richest ofany region in Mexico and includes a wide variety ofspecies. Most of the species are endemic, but severalare also known from the Sierra de El Abra and oneoccurs to the north. The collembolan Oncopoduraprietoi Bonet was described from Grutas del Palmitoin the Sierra de Gomas. Six species are also knownfrom caves in the Sierra de El Abra: the squamiferidisopod Spherarmadillo cavernicola Mulaik, the trichoniscidisopod Brackenridgin bridgesi (Van Name), theamblypygid Paraphrynus baeops Mullinex, the opilionidHoplobunus boneti (Goodnight and Goodnight),the centipede Newportia sabina Chamberlin,and the collembolan Pseudosinella petrustrinatiiChristiansen. Of the remaining fauna some speciesare known from both high and low elevations, whileothers may be restricted to either highland or lowlandcaves. Species which are known from all elevationsare the glomerid miIIiped Glomeroides promiscusCausey, the rhachodesmid milliped Strongylodesmusharrisoni Causey, the opilionid Hoplobunusinops Goodnight and Goodnight, and the gryllidcricket Paracophus caecus Hubbell. The cambalidmilliped Mexicambala blanda Causey is known bothfrom medium and low elevations, while its siblingspecies M. inops Causey occurs at higher elevations.Species which are apparently restricted to low elevationsinclude the pseudoscorpion Paravachoniumsuperbum Muchmore, the schizomid Schizomusreddelli Rowland, the ochyroceratid spider Theotimapura Gertsch, the trichopolydesmid milliped Tylogoneusminus Causey, and the histerid beetles Troglobacaniusreddelli Vomero and T. sbordonii Vomero.Of these species all bu t the milliped are probablytropical forms not likely to occur at higher elevations.The nearest relative of the milliped occurs inthe Xilitla region and thus it may also be found athigher elevations.The high-elevation fauna is of particular interest.It includes the chactid scorpion Typhlochactas rhodesiMitchell, the pseudoscorpions Aphrastochthoniusmajor Muchmore and Tyrannochthonius troglobiusMuchmore, the diplurid spider Euagrus cavernicolaGertsch, the agelenid spiders Cicurina (Cicurusta)mina Gertsch and Tegenaria blanda Gertsch, the leptonetidspider Leptoneta capilla Gertsch, the pholcidspider Metagonia pura Gertsch, the palpatorid opilionidOrtholasma sbordonii Silhavy, the cleidogonidmilliped Cleidogona pecki Shear, the rhachodesmidmiIIiped Unculabes causeyae Shear, a trichopolydesmidmilliped possibly belonging to the genus37

Speodesmus, an undescribed homopteran belongingto the family Kinnaridae, the leiodid beetle Ptomaphagus(Adelops) troglol1lexicanus Peck, and thecarabid beetles Antroforceps bolivari Barr, lHexaphaenopsintermedius Barr, and Platynus (Mexisphodrus)profundus (Barr). All of the troglobitesappear to be typical elements of the fauna of theSierra Madre Oriental.The aquatic troglophile fauna includes the dugesiidflatwoml Dugesia guatemalensis, the hyalellidamphipod Hyalella azteca (Saussure), the dytiscidbeetle Hydroporus belfragei Sharp, and the characinAstyanax mexicanus (Hubbs and Innes). The flatwormis endemic to the Sierra de Guatemala, but theremaining species are widespread.The terrestrial troglophile fauna contains an estimated83 species, a number of which are possibly endemicto the Sierra de Guatemala. It is not possibleto do more than mention some of the lllore distinctiveor important species here. Endemic species includethe syarinid pseudoscorpion Pachychitra similisMuchmore; the a~elenid spitler Cicurilla (C'icurusta)iviei Gertsch; the mysmenid spider JIaymena griseaGertsch: the pholcid spiders Coryssocnelllis clarusGertsch, Metagonia capil/a Gertsch, JI. secretaGertsch, 11'1. suzanne Gertsch. and J/. IIlckenzieiGertsch: the c1eidogonid milliped Cleidogona crystal/inaShear; the pyrgodesmid millipeds Myrmecodesmuscomutus (Shear), M. egenus (Causey). M.errabundus (Shear), and M. gelidus (Causey); thesphaeriodesmid milliped Sphaeriodesmus nortoniShear; the gryllid cricket Paracoplzus reddeUi Hubbell:and the staphylinid beetle Stenopholea reddelliHerman.Several troglophiles are known only from theSierra de Guatemala and Sierra de EI Ahra. Theseinclude the schizomid Schizomus mexiCQllUS Rowland,the amblypygid Paraphrynus pococki :'Ilullinex. thescytodid spider Loxosceles valdosa Gertsch, and thecarahid beetle Pachyteles urrutiai Bolivar. Amongmore widespread troglophiles mention may he madeof the agelenid spider Tegenaria selva Roth, thectenid spider Ctenus mitchelli Gertsch, the mysmenidspider 1Haymena chica Gertsch, the nesticid spidersEidmannella pal/ida (Emerton) and Caucelmus augustinusKeyserling, the pholcid spiders Coryssocnemisabernathyi Gertsch and Modisimus reddelli Gertsch,the tetrahlemmid spider Matta sbordonii (Brignoli),the opiJionid Cynorta jamesoni Goodnight and Goodnight,the collemholans Pseudosinella reddelli Christiansenand Acherontiella sabina Bonet, the gryllidcrickets Paracophus apterus Chopard and P. placonotusHubbell, the leiodid beetles Dissochaetus aztecusSzymczakowski and Ptomaphagus (Adelops) elabraPeck, the scarabaeid beetle Onthophagus cuevensisHowden, and staphylinid beetles of the genera Belonuchus,Homaeotarsus, Philonthus, and Stilicolina.Tula, Tamaulipas.-This ill-defined region is locatedsouthwest of Tula and comprises an area ofgypsum karst. The one cave studied in the region isat an elevation of 1,030 meters and is formed in] urassic gypsum. The only two species thus far identifiedare both troglophiles common in this part ofl\lexico: the carabid beetle Rhadine rotgeri (Bolivarand Hendrichs) and the staphylinid heetle Stilicolinan. sp.Villa Hidalgo, Tamaulipas.-This poorly definedregion is located to the north of the Rio Blanco inthe mountain~ west of Villa Hidalgo. The only cavethat has been visited is Cueva de la Virgen. a smalltwo-room cave about 60 meters in length and heavilypoilu ted. ~o troglobites are known from the regionand only three species, all probable troglophiles,have heen identified: the nesticid spider Eidmannellapal/ida (Emerton), the endemic gryllid cricketParacophus sanctorum Hubbell, and the alleculidheetle llymenorus sp.Gulf Coastal LowlandThe G ul I' Coastal Lowland province is a generallylevd plain extending from Texas to the YucatanPeninsula. It is broken in two places, first near Jalapawhere the Neovolcanic Plateau reaches the coast andfarther sou th by the Volcanes de los Tux tlas. AlcortaGuerrero (1966) includes only the area north of thel\l~ovolcanic Plateau in this province and places thesuuthem portion in a separate province. '\[ost of the"ulf Coastal Lowland is covered by non-cavernousdeposits, but one major and several smalll~r range~ ofCretaceous limestone arc exposed. The most extl~nsiveof these is the Sierra de Tamaulipas to the eastof Cilldad Victoria. Other smaller ranges such as theCerro el Aire and the Sierra de San Carlos also oceurin this area. :\[lIir (1936) has discussed the geology ofthis region. With the exception of one cave in theCerro eI Airc and three in the Sierra de Talllalllipas,this province remains unknown with respect to itsinvertebrate cave fauna.Cerro el Aire, Tamaulipas.-The Cerro el Aire islocated in the vicinity of the town of Abasolo immediatelynorth of the Sierra de Tamaulipas. The onlycave which has been visited is a small vertical pit,Sotano de Abasolo. Only two species of spider havebeen identified; one of these, the scytodid /'oxoscelessp., is an apparent troglophile.Sierra de Tamaulipas. Tamaulipas.-This range ofCretaceous limestone extends north-south for about100 kilometers. It is hounded on the Jlorth by the38

Rio Soto la Marina and terminates a short distancesouth of Aldama. Although narrow in the northernpart it widens to about 60 kilometers in the south.The highest peaks approach 1,400 meters in elevation.The principal cavernous rock unit is the TamaulipasLimestone (Muir, 1936). Martin et aI. (1954)have discussed the Sierra de Tamaulipas with respectto geology, climate, vegetation, and vertebrate fauna.The only caves that have been studied in the region arelocated at low elevations along the eastern side of therange. Nothing is known of the karst features or cavepotential of high elevations, although a few caveshave been reported by vertebrate biologists. Twocaves were visited about 50 kilometers southwest ofSoto la Marina, approximately midway in the range.One of these is a resurgence cave, but neither is verylarge. The third known cave is Cueva de los Cuartelesnear Aldanla in the extreme southern part of therange. This is a large, horizontal cave with manyentrances and more than one kilometer of passage.One part of the cave contains a large bat population,while another area consists of a single major passageinterrupted along its length by numerous large skylightentrances. A sizable spring is located near thecave and it is likely that Cueva de los Cuarteles is afossil resurgence.Despite our general paucity of knowledge of theinvertebrate fauna of the region 45 species, of whichfour are troglobites, have been thus far identified.The troglobitic fauna includes the cirolanid isopodSpeocirolana pelaezi (Bolivar) and its entocytheridostracod associate, Sphaeromicola cirolanae Rioja.These species are also known from caves in SanLuis Potosi and Tamaulipas in the Sierra MadreOriental. The remaining tw~ troglobites are schizomids,Schizomus lukensi Rowland and an undescribedspecies of Schizomus. Both are closelyrelated to species in the Sierra de EI Abra and Sierrade Guatemala and presumably represent relicts of awidespread species ancestral to species in all threeregions. The terrestrial troglophile fauna is alsoclosely related to the fauna of the Sierra MadreOriental, with some species occurring in both regions.Three arachnids are endemic to the caves of theSierra de Tamaulipas: the pholcid spiders Pholcophorabolivari Gertsch and P. mitchelli Gertschand the opilionid Stygnomma tuberculata Goodnightand Goodnight. Other species of interest include theamblypygid Paraphrynus pococki Mullinex, the ctenidspider Ctenus mitchelli Gertsch, the mysmenidspider ,vJaymena chica Gertsch, the nesticid spiderGaucelmus augustinus Keyserling, the pholcid spiderModisimus texanus Banks, the scytodid spider Loxoscelesdevia Gertsch and !Vlulaik, and the collembolansPseudosinella violenta (Folsom) and Acherontiellasabina Bonet.Neovolcanic PlateauThe Neovolcanic Plateau forms a great band acrosscentral Mexico from the Pacific coast in J alisco to theGulf of Mexico east of Jalapa, Veracruz. Hundreds ofvolcanos, most now extinct, rise from a comparativelylevel plain formed by ash and lava deposits.Two volcanos, J orullo and Paricutin, have erupted inrecent times and an eruption south of Mexico Citycovered prehistoric settlements in the Pedregal de SanAngel. A chain of large peaks, including Tancitaro,Toluca, Popocatepetl, Malinche, Orizaba, and Cofrede Perote, dominates the landscape. The average elevationin this province is above 2,500 meters, withOrizaba rising to 5,747 meters above sea level. In afew areas erosion has exposed older sedimentaryrocks, some of which are known to be cavernous.. 'one of these inliers have been investigated for caves.Lava tubes are abundant throughout much of thearea, but they have been little studied speleologically.Caves on the slopes of Cerro Xictle to the south ofMexico City have been studied with respect to theirbat and associated ectoparasitic fauna, but generalinvertebrate collections have not been made. Onecave on Cerro de la Estrella to the east of MexicoCity has been visited and others are known to exist.Two caves on the slopes of Cofre de Perote have beenexamined with collections made in one. A third cavenear Jalapa has been studied, and three caves near thetown of Buena Vista have been visited. The presenceof troglohites in the caves on Cerro de la Estrella andCofre de Perote indicates the existence of a potentiallyrich cavernicole fauna on the Neovolcanic Plateau.Cerro de la Estrella, Distrito Federal.~This smallpeak is located south of Tapijutapa on the easternedge of Mexico City. It is formed of Pliocene volcanicdeposits (Schlaepfer, 1968). One cave has beenbiologically investigated, but other larger ones arereported. This cave is generally dry and heavily pollutedwith trash and human feces. Of eight speciesidentified from the cave, an undetermined genus andspecies of nicoletiid thysanuran is the only possibletroglobite. The troglophilic fauna includes the nesticidspider Eidmannella pallida (Emerton) and anundescribed species of the pholcid genus Coryssocnemis.Buena Vista, Veracruz.-This is an ill-definedregion in the immediate vicinity of the town ofBuena Vista. Three caves, none very well-suited forcavernicoles, have been studied. One cave is small,horizontal, and of little interest; the second, Cuevade Cantil Blanco, consists of a single 80.meter.long39

horizontal passage with a small stream issuing fromit; the third is a large, poorly-explored bat cave,Cueva de Camposantos, which consists of one ormore large chambers. The stream in Cueva de CantilBlanco was inhabited by the snail Melanoides maculata(Born) and an undescribed species of palaemonidshrimp of the genus Macrobrachium. The terrestrialtroglophile fauna consists only of the pholcid spiderPhysoeyclus globosus (Taczanowski) and an undescribedspecies of the scytodid spider genus Loxosceles.The remainder of the fauna consists of accidentalsor bat guano associates.Jalapa, VeracRlz.-This poorly defined and studiedregion consists of the volcanic regions in the generalvicinity of Jalapa, including Cofre de Perote to theeast. Only two caves have been investigated withrespect to their invertebrate fauna. One, Cueva delInfiernillo, is a lava cave entered by a 10-meter deepsinkhole. A single large passage about 200 meters longis now accessible. The cave contains much guano andis not suitable for troglobites. The second cave, Cuevadel Volcancillo, is located on the slopes of Cofre dePerote. The entrance opens onto the wall of thecrater of a subsidiary cone, EI Volcancillo, and extendsfor more than 1,000 meters as a large tube.Numerous caves occur on Cofre de Perote and thearea should prove rich biologically. Two troglobiteshave been identified from Cueva del Volcancillo, anundescribed species of the cleidogonid milliped genusCleidogona and an undescribed carabid beetle of thegenus Para tree hus. The troglophile fauna of theregion includes the amblypygid Paraphrynus aztecus(pocock), a scorpion of the genus Vaejovis, agelenidspiders of the genus Tegenaria, the nesticid spiderEidmannella pallida (Emerton), the pholcid spiderModisimus n. sp., the scytodid spider Loxoscelesn. sp., the cleidogonid milliped Cleidogona n. sp.,campodeid diplurans, and the collembolan Schae!feria sp.Sierra Madre del Sur SystemAccording to Raisz (1964) the Sierra Madre delSur System includes all of the area between theNeovolcanic Plateau and the Isthmus of Tehuantepec,with the exception of a narrow strip along the PacificOcean and the Gulf Coastal Lowland to the north. Hesubdivides this province into five sections: the BalsasMexcala Basin, the Oaxaca Upland, the NortheastFolded Ranges, the Southern Slope, and the NorthernSection. The Southern Slope is a mountainous areaalong the Pacific coast; the Northern Section is thecoastal area in Colima and immediate vicinity. Neitherof these regions has been studied speleologicallyand will not be discussed here.The Balsas-Mexcala Basin is a structurally complexregion largely drained by the Tepalcatepec, Balsas,and Mexcala rivers. It is bounded on the north by theNeovolcanic Plateau, on the northwest by the NorthernSection, on the south by the Southern Slope andon the east by the Oaxaca Upland. The rivers flow atelevations of 300 to 600 meters, while to the souththe land rises to the heights of the Sierra Madre delSur at 2,000 to 3,000 meters. To the east the basinwidens to form a broad plateau dissected by severalrivers. Part of the area is covered by Tertiary volcanics,but Cretaceous limestone is exposed in manyplaces. Along the northern edge of the region, erosionhas removed the overlying volcanic rocks to exposeheavily karsted Cretaceous deposits. This is mostnotably seen in the Cacahuamilpa Region, as discussedbelow. Bonet (1971) has discussed the physiographyand geology of the Cacahuamilpa region, while Fries(1956) has reported on the geology along the highwaybetween Mexico, D. F., and Acapulco, Guerrero.With the exception of the Cacahuamilpa regionthe Balsas-Mexcala Basin has been little studiedspeleologically. Only 17 caves have been examinedfor their invertebrate fauna, but reports on bats inmany parts of the area indicate that it will prove tobe of the greatest interest speleologically. Extensivelimestone deposits, considerable local relief, and highrainfall have contributed to the development of manyextensive cave systems.The Oaxaca Upland is a higher area than the basinto the west and has an overall plateau-like character.To the north it is much dissected and in the centerlies the Valley of Oaxaca. The region is geologicallycomplex with granites, metamorphic rocks, and sedimentarydeposits exposed. Cretaceous limestonecrops out particularly in the southern part of the upland.The area has been little studied speleologicallyand only ten caves have been visited. These are locatedat 1,800 to 2,000 meters elevation in the vicinitiesof San Sebastian and Santiago Apoala.The Northeast Folded Ranges is a disjunct part ofthe Sierra Madre Oriental extending from Cordoba,Veracruz, to Tehuantepec, Oaxaca. It consists largelyof Cretaceous limestone, but volcanic deposits insome areas cover the limestone. The geology of thenorthern part of the area has been discussed by ContrerasV. (1956). This part of Mexico is among themore speleologically important areas in the country.Heavy rainfall, high relief, and massive limestonedeposits have contribute~ to the development ofsome of the more signir;tant caves in Mexico. It isalso one of the better known parts of Mexico withrespect to its invertebrate cave fauna. Among the 10840

caves studied are many large stream caves (particularlyin the Acatlan region of Oaxaca) and some ofthe deeper caves in the world. Extensive karst developmentsouth of Orizaba, Veracruz, and in the Huautlade Jimenez region of Oaxaca has resulted in nearlycomplete internal drainage. Deep, incised river canyonsat the base of these plateau-like regions has providedthe relief necessary for the development ofdeep vertical cave systems, one of which is more than1,000 meters deep. Knowledge of the cave fauna ofthe area, however, is limited to the northern part ofthe mountains. Extensive cave systems have beenreported from the southern part of the ranges butremain unstudied.Colotlipa. Guerrero.-This region is located in easternGuerrero in the vicinity of the town of Colotlipa.The general geology of this part of Guerrero is describedby Fries (1956). The principal limestone unitis the Morelos Formation of Cretaceous age. TheColotlipa region is poorly known and defined andonly Grutas de Juxtlahuaca has been studied. Thislarge cave, now partially developed as a tourist attraction,is famous for its Olmec burials and paintings.The cave is essentially horizontal and contains a smallstream and deep pools. With more than five kilometersof surveyed passage, it is among the longer cavesin Mexico (Roy, 1974).Despite the fact that 35 species, of which four aretroglobites, have been identified from the cave, thefauna remains poorly studied. Several rooms are inhabitedby a large bat population, ten species ofwhich have been identified. The terrestrial troglobitefauna includes the pholcid spider Pholcophora grutaGertsch, the rhachodesmid milliped Pararhachistesamblus Chamberlin, the campodeid dipluran Juxtlacampajuxtlahuacensis Wygodzinsky, and a collembolanof the genus Trogolaphysa. All of these speciesare endemic to Grutas de Juxtlahuaca and are allclosely related to other species in southern Mexicoand Guatemala. Several troglophiles are endemic tothe cave; these include the armadillid isopods Venezilloarticulatus (Mulaik) and V. boneti (Mulaik),the tridenthchoniid pseudoscorpion Tridenchthoniusjuxtlahuaca Chamberlin and Chamberlin, the pholcidspider Physocyclus bicornis Gertsch, the pygrodesmidmilliped llJyrmecodesmus colotlipa (Chamberlin), andthe spirostreptid milliped Orthoporus guerreronus(Chamberlin). Other troglophiles include the annadillidisopod Venezillo cacahuamilpensis (Bilimek),the amblypygid Paraphrynus mexicanus (Bilimek),the scutigerid centipede Scutigera linceci (Wood),the nicoletiid thysanuran Anelpistina boneti (Wygodzinsky),and the carabid beetle Platynus segregatus(Bates). These are all typical elements of the fauna ofsouthern Mexico and many are also reported fromGrutas de Cacahuamilpa.Taxco. Guerrero.-This a poorly defined regionnear the city of Taxco in northern Guerrero. Theprincipal cave-forming limestones in the region arethe Cretaceous Mexcala and Morelos fonnations(Fries, 1956). Although numerous caves are knownfrom reports by zoologists studying bats, only twohave been examined with respect to their invertebratefauna, and neither of these is well known. Spiderswere collected from a small cave on the outskirts ofTaxco, but nothing is known of the cave. The onlyother cave examined is Pozo Melendez, a vertical cavesystem to the south of Taxco. Of eleven species ofinvertebrate reported from the area the only troglophilesof note are the agelenid spider Tegenaria mexicanaRoth, the scytodid spider Loxosceles mistecaGertsch, and the paradoxosomatid milliped Oxidusgracilis (Koch).Cacahuamilpa. Guerrero and Mexico.-This regionis located in northern Guerrero and adjacent Mexicoin Valle de Ixtapan immediately south of the slopesof the volcano Nevada de Toluca. The region isflanked on the east and west by plains of volcanicdeposits and on the south by the Sierra de Tenerias.The geology, speleologenesis, and caves of the areaare discussed in detail by Bonet (1971). This is amongthe more interesting karst regions in Mexico, withseveral large caves known, the most famous of whichis Grutas de Cacahuamilpa. This large horizontalcommercial cave has been known for hundreds ofyears and is described in numerous popular and scientificarticles. Other well-known caves includeGrutas de la Estrella and the two large river caves(Grutas del Rio Chontalcoatlan and Grutas del RioSan Jeronimo) which emerge from two enonnousentrances, collectively called the Dos Bocas, belowGrutas de Cacahuamilpa. The caves of the region areformed in Cretaceous limestone, principally of theMorelos and Mezcala fonnations. Most of the areawas covered by volcanic deposits, but as erosionexposed the underlying limestone karst topographyseveral major streams invaded preexisting cave systemsand now run for several kilometers throughunderground channels.Fourteen caves have been investigated with respectto their invertebrate fauna, but most of the wellknowncaves are large stream passages and not wellsuited to cavernicoles. Additional work in the smaller,drier caves is needed to further elucidate the natureof the fauna of the area. Although 91 species havebeen recorded, only six are troglobites. This paucitymay be due to our limited knowledge of the regionor to the possibility that filling or burial of the caves41

y volcanic deposits prevented the development of arich troglobitic fauna. The troglobites known fromthe region include the pseudoscorpion Albiorix bolivariBeier, an undescribed species of the schizomidgenus Schizomus, the opiIionid Caecoa arganoi Silhav)',the rhachodesmid milliped Ceuthauxus constansCausey, the thysanuran Anelpistina anophthalma(Bilimek), and the collembolan Spelaeogastruraguerrerense Bonet. Two copepods, Macrocyclopsalbidus (J urine) and Mesocyclops sp. ne. brazilianusKiefer, are the only probable aquatic troglophiles.The terrestrial troglophile fauna is closely allied withthe cavemicole fauna of southern Mexico, but severalspecies are known only from the caves of this region.These include the pseudoscorpion Lechytia cavicolaMuchmore; the pholcid spiders Physocyclus modestusGertsch, Psilochorus cordatus (Bilimek), and Psilochorustellezi Gertsch; the ricinuleid Cryptocellusboneti Bolivar; the opilionid Cynortina minutus(Goodnight and Goodnight); the pyrgodesmid millipedMyrmecodesmus acona (Shear); and the carahidbeetIe Tachys (Tacltyura) un is triatus (Bilimek).Other troglophiles of interest include the armadillidisopods Venezillo cacaltuamilpensis (Bilimek) andV. osorioi (Mulaik), the amblypygid Paraphrynusmexicanus (Bilimek), the agelenid spider Tegenariamexicana Roth, the mysmenid spider Maymenamistica Gertsch, the scytodid spiders Loxoscelesboneti Gertsch and L. misteca Gertsch, the carabidbeetles Paratrechus (Paratrecltus) tepoztlanensis Bolivarand Platynus (Stenoplatynus) umbripennis(Casey), the leiodid beetle Ptomapltagus (A delops)spelaeus (BiIimek), and the scarabaeid beetle Onthophagusvespertilio Howden, Cartwright, and Halffter.Acatlan, Oaxaca.-This region includes a series ofmountain ranges located near the town of Acatlimin ex treme northern Oaxaca. The principal cavernousformation is the Escamela Limestone of Cretaceousage (Lozano Romen, 1955). The mountainsarise abruptly from an alluvial plain on which sugarcaneis grown. No caves have yet been visited on thehigh slopes or tops of the ranges where pits andsumideros are reported to occur.. ;umerous cavesexist on the slopes of the mountains, and severalsizable streams emerge from cave entrances at thebase of the ranges. One stream now utilizes themain passage in Cueva de Juan Sanchez for morethan a kilometer (Byrd, 1976). Other notable cavesin the area include Cueva de la Finca, Cueva deLaguna Verde, Cueva de las Maravillas, and Cuevadel Nacimiento del Rio San Antonio, each of whichcontains several kilometers of passage.The cavemicole fauna of this still poorly studiedregion is among the richer in Mexico. From the 21caves biologically investigated, 91 species, of which25 are troglobites, have thus far been identified. Theregion is particularly rich in aquatic troglobites withseven species known: the mysids Spelaeomysis olivaeBowman and Antromysis (Antromysis) reddelliBowman, the a1pheid shrimp Alpheopsis stygicolaHobbs, the palaemonid shrimps iYlacrobrachium n. sp.and M. villalobosi Hobbs, the crayfish Procambarus(Austrocambarus) oaxacae reddelli Hobbs, and anundescribed species of catfish of the genus Rhamdia.The mysids and alpheid shrimp are presumablymarine relicts, while the other species probably havea freshwater origin. With the exception of A. stygicola,which is otherwise unknown from fresh water inthe New World, the remaining species are all closelyrelated to the fauna of southern lVlexico. The terrestrialtroglobite fauna includes the diplocentrid scorpionDiplocentrus cueva Francke, an undescribedspecies of pseudoscorpion of the genus Tyrannochthonius,the schizomid Schizomus ?firstmani Rowland,an amblypygid of the genus Paraphrynus, thetarantula Schizopelma reddelli Gertsch, the pholcidspiders Metagonia martha Gertsch and Pholcophoran. spp., the cyphophthallllid opilionid Neogoveamexasca Shear, an undescribed opilionid of the genusHoplobunus, undescribed millipeds of the familiesOniscodesmidae and Trichopolydeslllidae, an undescribedthysanuran of the family \icoletiidae,collembolans of the subfamily I'aronellinae and thegenera Pseudosinella and .-lcherontides, and an undescribedcixiid hOlllopteran.The troglophile fauna has not, for the most part,undergone systematic study, bu t a few species havebeen identified. These include the schizolllid Schizomusportoricensis (Chamberlin). the amblypygidParaphrynus aztecus (Pocock), dl'~ rnysmenid spidersJ-Jaymena delicata Gertsch and M. mayana (Chamberlinand Ivie), and the leiodid beetles Dissochaetllshetschkoi Reitter and Ptomaphagus (A delops) reddelliPeck. In addition to these species many undeterminedor undescribed troglophiles are known. Among themore significant of these are scorpions of the genusVaejovis, pseudoscorpions of the genera Albiorixand Tyrannochtltonius, diplurid spiders of the genusEuagrus, ctenid spiders of the genus Ctenus, pholcidspiders of the genus Pholcophora, scytodid spiders ofthe genus Loxosceles, pyrgodesmid millipeds of thegenus IHyrmecodesmus, eampodeid diplurans, leiodidbeetles of the genus Proptomaphaginus, scydmaenidbeetles of the genera Euconnus (Napochus) and Scydmaenus,and staphylinid beetles of the genus Belonuchus.Both the troglophile and troglobi te faunas ofthe Acatlan region are typical of the cavernicolefauna of other parts of southern Mexico. The only42

unusual species is Neogovea mexasca; this is the onlytroglobitic cyphophthalmid in the !\ew World and theonly species recorded for lVlexico or Central America.Huautla de Jimenez, Oaxaca.-This region is a highplateau located near the town of Huautla de Jimenezin northern Oaxaca near the Puebla border. Thisplateau, at an elevation of about 2,000 meters, isbounded on the south by the valley of the Rio SantoDomingo at an elevation of less than 350 meters, onthe north and west by the valley of the Rio Coyomeapanand the Rio del Camaron, and on the northeastby the Miguel Aleman Reservoir at an devation ofless than 100 meters (A nonymous, 1965). The greatestdevelopment of karst has been in the hil!her partsof the plateau to the cast and southeast of Huautlade Jimenez. Most of the drainage on the plateau isinternal. the water emerging from springs in the rivervalleys more than 1,500 meters below. Karst formspresent on the plateau include lapiez, pinnacles, andlarge dohnas. :Vlany of these dohnas contain pits, andthe larger-such as that in which Sotano de San Agustinis found-are more than 100 meters deep and onekilometer lonl!. The maturity of the karst, the greatlocal relief, anrl massivl~ Crdacl~oUS limestone haveprovided the facLors necessary for the developmentof extensive integrated caVI~ syst(~ms. The larger cavesinclude Sotano de A~ua de Carrizo. La Grieta, Sotanodel Rio Iglesia, and Si,tano de .::lan Agustin. The lastcave has a depth of more than 1,000 nwters and isalllong the deeper in the world. ,\dive streams flowthrough tlw deep caVI~S. Russell (1965), Fish andI{ussell (1966), Boon (1969). ~tone (1977, 1979a).and Steel (1978) describe SOUle of the caves. Inaddition to caves on the platcllu, several caves in thevalley of the Rio Coyomeapan have heen studied.These are generally small, horizontal caves with arich fauna.The invertehrate fauna of the rqrion. thou!!h stillpoorly known includes -l6 identifil~d species. ofwh idl f~iO"htr:':'an~ trfwl ohi tes. Tlw tnwlobi tic fauna~ ~inrludes a scorpion of Iincertain affinities. an undescribedtarantula of tht:: geluls Schi::opelma, anundescribed phokid spider of the /!enu~ Pholcophora,the ckidogonid millipl~d Cll'idof!ona baroquaShear, the l:ambalid Inillipl'dllexicam bala fishiCallsey, the collemholan J'seudosinel/a bonita Christiansen,an IIndescrihed i'l'llIIS and ,;pecies of nicoletiidthysanllran. and L1", carahid beetle Platynus(Mexisphodrus) llrqllijoi (Ilendrichs and Hoi ivar). Thetruglobites all hdon/! to i'enera known from theSierra \Iadre Oriental as well as ()ax aca and Puehla.The tro/,Iophile fallna. tholl/!h poorly known taxonomically,bdol1l!s. as ooes the troglouite fauna, togroups common both in southern \Iexico and theSierra Madre Oriental. Species included are a schizomidof the genus Schizomus; an amblypygid of thegenus Paraphrynus; ctenid spiders of the genus Ctenus;mysmenid spiders of the genus ]l,Jaymena; thenesticid spiders Eidmannel/a pallida (Emerton) andGaucelmus calidus Gertsch; pholcid spiders of thegenera Coryssocnemis, Metagonia, and Modisimus;opilionids of the genera lloplobunus and Karos;the rhachodesmid milliped Rhachodesmus digitatusCausey; sphaeriodesmid millipeds of the genusSphaeriodesmus; campodeid diplurans; collembolansof the genus Pseudosinella; carabid beetles of thegenus Platynus; and staphylinid beetles of the genusBelonuchus.San Sebastian de las Grutas, Oaxaca.~This regionis located in the Oaxaca Upland to the north of Solade Vega. It comprises an ill-defined area of Cretaceouslimestone at elevations of 1.800 to 2,100meters. Karst development consists principally oflarge shallow dohnas. Two of the caves studied aresmall pits and a third is a small horizontal cave. Thefourth cave, Grutas de San Sebastian, is a large,essentially horizontal cave with a large stream in alower level.The troglobitic fauna includes an undescribedmilliped of the family Triehopolydesmidae, an undescribednicoletiid thysanuran, and a collembolanof the genus Acherontides. The troglophile faunaincludes the endodontid snail lIelicodiscus singleyanus(Pilshry), a schizomid of the genus Schizomus,the Inysmenid spider Maymena delicata Gertsch, thenesticid spider Caucelmus augustinus Keyserling, apholcid spider of the genus Physocyclus, the opilionidHoplobunus oaxacensis Goodnight and Goodnight,and a pyrgodesmid milliped of the genus Myrmecodesmus.These species all helong to genera commonto the eaves of southern Mexico.Santiago Apoala, Oaxaca.-This is a little-studiedarea of limestone karst located about 20 kilometersnortheast of Asuncion \ochixtlan in northern Oaxacaon a high plateau at an e1f~vation of about 2,240meters. Karst development consists of numerouslarge, shallow rlolinas. from a few of which pitsdescend. Drainage appears to be almost entirely internal,but no large integrated systems have beendiscovered. Of the six caves visited only two are ofany great extent. One of these, Comedor del Diablo,is a deep vertical system still poorly explored; theother, Cueva de Apoala, is loeatt~d more than 200meters below the edge of the plateau and consistsof a large passage from which a stream emerges. It isalso poorly explored.Tht~ cave fauna of this rather isolated region ispotentially of considerable interest, but it has been43

little studied. Of 30 species identified, only two aretroglobites, the endemic opilionid Hoplobunus apoalensisGoodnight and Goodnight and an undescribedrollembolan of the subfamily Paronellinae. The troglophilefauna includes the nesticid spiders Gaucelmusaugustinus Keyserling and Nesticus n. sp., the pholcidspiders Coryssocnemis sp. and Psilochorus murphyiGertsch, a cleidogonid milliped of the genus CleidogOM,a collembolan of the genus Pseudasinella, andthe carabid beetle Mexitrechus coarctatus (Bates).VaUe Nacional, Oaxaca.-This region is an area ofCretaceous limestone located near the town of ValleNacional in northern Oaxaca. All of the caves studiedhave been near Valle Nacional at an elevation of lessthan 100 meters, but the mountains to the south riseto an elevation of more than 2,000 meters. Karst inthe form of large dolinas occurs at higher elevationsand this area deserves study. Two of the five cavesvisited are small and of little interest. Grutas deMonteflor is a well-decorated, horizontal cave withseveral hundred meters of passage. Cueva del Guanohas a large bat population in its upper levels, but alower-level stream passage contains good habitatsfor both aquatic and terrestrial invertebrates. Thelargest cave in the area, Cueva del Guayabo, is a long,horizontal cave which has not been well explored;pools in the lower part of the cave are apparentlyleft from seasonal artesian flooding.This region, still poorly known biologically, contains42 identified species of cavernicole, of whicheight are troglobites. The aquatic troglobitic faunaincludes the endemic palaemonid shrimp Neopalaemannahuatlus Hobbs, an undescribed stenasellidisopod, and the crayfishes Procambarus (Austracambarus)oaxacae oaxacae Hobbs and P. (A.) oaxacae?reddelli Hobbs. The latter subspecies is separatedfrom the other by the Rio Valle Nacional and isotherwise known from caves in the Acatlan andAtoyac regions. The terrestrial troglobite fauna includesthe hyid pseudoscorpion Troglohya carranzaiBeier, an undescribed genus and species of trichopolydesmidmilliped, an undescribed nicoletiid thysanuran,and a collembolan of the subfamily Paronellinae.The terrestrial troglophile fauna includes severalendemic species: the schizomids Schizomus moisiiRowland and S. stewarti Rowland, the opilionidHoplobunus spinooculorum Goodnight and Goodnight,the pyrgodesmid milliped Myrmecodesmusamplus Causey, and the leiodid beetle Ptomaphagus(Adelops) reddelli Peck. Other troglophiles, all typicalrepresentatives of the cavernicole fauna of southernMexico, include the amblypygid Paraphrynus aztecus(Pocock), ctenid spiders of the genus Ctenus, themysmenid spider Maymena mayana (Chamberlin andIvie), the nesticid spider Eidmannella pallida (Emerton),the tetrablemmid spider Matta sbordonii (Brignoli),cleidogonid millipeds of the genus CleidogOlla,campodeid diplurans, the collembolan Acherontiellasabina Bonet, scydmaenid beetles of the genus Scydmaenus,and staphylinid beetles of the genus Belonuchus.San Pablo Zoquitlim, Puebla.-This region islocated in extreme soutlleastern Puebla and comprisesan area of Cretaceous limestone with muchinternal drainage. Although nine caves have beenbiologically investigated, the region remains poorlvknown. Cave development in the area appears to belargely vertical and several caves attain considerabledepth. The most notable of these are Sotano deCoyomeapan which terminates in a sump at 166me ters (Atkinson, 1978) and Cueva de Xocotlatwhich has been explored to a depth of 339 meters,but remains largely unexplored (Atkinson and Forsythe,1979). These two caves have very large streamsand exploration is difficult and hazardous.Only 17 species have been identified, of which theonly troglobite is an eyeless nesticid spider of thegenus Nesticus. The troglophile fauna includes theamblypygid Paraphrynus mexicanus (Bilimek), agelenidspiders of the genera Cicurina and Tegenaria,the pholcid spider Coryssocnemis abernathyi Gertsch,an opilionid of the genus Hoplobunus, nicoletiidthysanurans, and staphylinid heetles of the genusStilicolina.Atoyac, Veracruz.-This region is located east ofCordoba and comprises Cretaceous limestone mountainsex tending from the vicinity of Cuitlahuac northwestfor about 30 kilometers. The exact ex tent ofthe region is not known but includes the town ofAtoyac and the higher mountains north of Potrero.Elevations range from less than 300 meters to morethan 1,000 meters. At higher elevations near thevillage of Man:taniIlo, karst development is moderatewith numerous shallow dolinas, many small pits,lapiez, and pinnacles. 1uch of the drainage is internal,water exiting from large springs at much lowerelevations. No large caves are known from the higherelevations and most were found to be short, horizontaltunnels or small pits. At and near the base of therange, however, several significant caves were found.The most notable of these is the famous Gru tas deAtoyac, a horizontal cave of several hundred metersin length, located on the cliffs overlooking the canyonof the Rio Atoyac. Other interesting caves includeCueva de Sala de Agua north of Cuitlahuac andCueva del Ojo de Agua Grande north of PotreroViejo, from both of which large streams emerge.Despite the fact that 25 caves are known for the44

pallidus Rowland, the nesticid spider Nesticus arganoiBrignoli, the oniscodesmid milliped Bonetesmus ojoShear, the rhachodesmid milliped Acutangulus aliusCausey, an undescribed species of the milliped familyTrichopolydesmidae, a collembolan of the genusPseudosinella, a kinnarid homopteran, and the carabidbeetle Plat.ynus (Mexisphodrus) veraecruclS(Barr). With the exception of the two millipeds, allof the fauna belongs to groups widespread in southernand eastern Mexico. Bonetesmus and Acutangulusare primarily restricted to southern Mexico.The troglophile fauna includes several endemicspecies, among which are the schizomid Schizomuslanceolatus Rowland; the pholcid spiders Coryssocnemisplacidus Gertsch and Modisimus beneficusGertsch; and the opilionid Karos brignolii Silhavy.Other troglophiles of interest include the amblypygidParaphrynus aztecus (Pocock); ctenid spidersof the genus Ctenus; the mysmenid spiders Maymenacascada Gertsch and M. mayana (Chamberlin andIvie); the nesticid spiders Eidmannella pallida (Emerton)and Gaucelmus calidus Gertsch; pholcid spidersof the genera Coryssocnemis and Psilochorus; thetetrablemmid spider Matta sbordonii (Brignoli); theopilionids lloplobunus robustus Goodnight andGoodnight and Karos rugosus Goodnight and Goodnight;a rhagidiid mite; cleidogonid milIipeds of thegenus Cleidogona; sphaeriodesmid millipeds of thegenus Sphaeriodesmus; campodeid diplurans; collembolansof the genus Pseudosinella; the psocidPsyllipsocus ramburii Selys-Longchamps; the carabidbeetles Paratrechus (Paratrechus) mexicanus Putzeysand Platynus tlamayaensis (Barr); scydmaenid beetlesof the genus Euconnus; and staphylinid beetles of thegenus Belonuchus. This fauna is also typical of that ofsouthern and eastern Mexico.Tezonapa, Veracruz.-This poorly defined regionis located about 20 kilometers west-southwest of thetown of Tezonapa. The only cave visited, Cueva deUngurria, is located now on an island in the RioTonto. The lower parts of the cave are flooded as aresult of raised water levels from the Miguel AlemanReservoir. The cave itself now has limited access, butconsiderable karst development on the island indicatesthat the area is promising.The fauna of Cueva de Ungurria includes 14 species,of which two are troglobites. The troglobites arethe glomerid milliped Glomeroides addititius Causeyand a collembolan of the subfamily Paronellinae. Thetroglophile fauna includes a species of the schizomidgenus Schizomus, the amblypygid Paraphrynus aztecus(Pocock), a scytodid spider of the genus Loxosceles,campodeid diplurans, and a scydmaenid beetleof the genus Scydmaenus.Chiapas-Guatemala HighlandsThe Chiapas-Guatemala Highlands province is atitle applied by Raisz (1964) to the mountainousregions of Guatemala and Belize and the mountainsof Mexico east of the Isthmus of Tehuantepec. Thearea has been concisely described by West (1964),and the discussion below relies heavily upon hisaccount. A series of arcuate ranges and depressions beginsin Chiapas, extends generally northwest-southeastinto Guatemala where it continues more nearly westeast.The first major range is a series of igneousmountains, the northern part of which is the SierraMadre de Chiapas, rising from the Pacific coastalplain. To the north of this is a major depressionknown as the Chiapa Depression in Mexico and theMotagua Valley in Guatemala. Rising from this is asecond major series of mountains. The southern partof the range is a high, plateau-like region that beginsas the Sierra de San Cristobal in Chiapas, becomes theAltos Cuchumatanes and highlands of Alta Verapazin Guatemala, and finally the Sierra de Santa Cruznear the Caribbean coast. Along the northern edgeof this plateau lies a series of folded ranges. Beginningin Mexico as the Sierra de los Lacandones, theranges descend to form low, knob-like mountains inthe southern Peten of Guatemala, and then rise toform the Maya Mountains of Belize.The geology of the Chiapas-Guatemala Highlandsis complex and not very well known. Gutierrez Gil.(1956) and Olivas (1956) have summarized thegeology of Chiapas. The geology of southeasternMexico and northern Guatemala is discussed byLopez Ramos (1969), while Vinson (1962), Bonis(1969), and Nagle et al. (1977) briefly summarizeGuatemalan geology. Dixon (1957) gives an accountof the geology of southern Belize.The Sierra Madre de Chiapas is a batholith ofPaleozoic age composed mainly of granite and dioritepartly covered in the southern part by Cretaceousrocks. The Guatemalan continuation is largely mantledby lava and ash deposits. Most of the area is, ofcourse, noncavernous, but two caves in the Motozintlaregion have been studied.The Chiapa Depression is a wide basin drained bythe Rio Grijalva and its tributaries. The basin risesgradually from about 700 meters along the Guatemalanborder to about 900 meters in the northwest.The Rio Grijalva cuts through the basin, its canyongradually increasing in depth. It finally dissects theSierra de San Cristobal in the great gorge known asEl Sumidero. The principal rock units in the area areof Cretaceous age, but some Eocene and Oligocenedeposits occur. The Guatemalan portion of thisdepression is drained by the Rio Motagua and is46

generally more dissected than in Chiapas. Numerouscaves are known in the Chiapa Depression, mostly inthe vicinities of Tuxtla Gutierrez, Ocozocoautla, andMalpaso.The central plateau of Chiapas is among the greatkarst regions in Mexico. The plateau rises steeplyfrom the Chiapa Depression to an elevation of about2,300 meters. Parts of the plateau, such as near SanCristobal de las Casas, are rolling plains with numerousshallow dolinas. In other areas there is the greatestdevelopment of karst in Mexico, with sinkholes,dolinas, poljes, and sinking streams occurring everywhereon the Cretaceous limestone surface. This isthe best-studied part of Chiapas; 48 caves have beenstudied biologically. This surface continues on alesser scale into the Altos Cuchumatanes of Guatemalawhich attains an altitude of more than 3,000meters. Farther east in Alta Verapaz the topographyis more rugged; and in the vicinity of Coban, it isbroken into three parallel mountain ranges separatedby basins. Karst development in this area is notable,with large caves, dolinas, sinking streams, and othersolution features being abundant.The northern side of the plateau is composed of aseries of folded ranges in Cretaceous and Tertiaryrocks. As the ranges pass into Guatemala, theydecrease in heigh l until they are hills in the sou themPeten. In Belize the land rises in the Maya Mountains,where intrusive rocks and Pennsylvanian and olderdeposits are exposed. Limestones of Cretaceous toEocene age are exposed to the north, south, and westof the Maya \10untains and support a mature karsttopography.Altamirano, Chiapas.-This region is located nearthe towns of Altamirano and Ocosingo in the foldedmountains north of the high plateau. Eight caves havebeen studied, seven of which are described by Sbordoniet al. (1977). The caves are generally horizontaland at least one, Sumidero de Panteon, receives considerablefloodwater.The caves appear to be quite rich biologically, butlittle has been published on them to date. Of 16species thus far reported, four are troglobites. Threeof the troglobites are aquatic and include the asellidisopod Caecidotea vomeroi Argano and the bogidiellidamphipods Bogidiella sbordonii Ruffo and VignaTaglianti and B. tabascensis Villalobos. The last speciesis also known from Grutas del Cocona nearTeapa, Tabasco. The only terrestrial troglobite is anundescribed roach of the genus Nelipophygus. This isthe only tro~lobitic roach in Mexico; the genus isotherwise known from Jamaica where a troglobiticspecies also occurs. The troglophile fauna includes theamphipod Hyalella azteca Saussure, the ochyroceratidspider Ochyrocera fagei Brignoli, the oonopid spiderOonops chickering; Brignoli, the pholcid spiderCoryssocnemis pecki Gertsch, and the roach Nesomylacrislateralis Fisk. Except for the Widespreadamphipod H. azteca and the spider C. pecki, all ofthe troglophiles are endemic to this region. The latterspecies is also known from caves in the San Cristobalde las Casas region.Bochil and Soyalo, Chiapas.-This region is locatedto the northwest of San Cristobal de las Casas. Thesix caves studied occur at elevations ranging from1,250 to 1,600 meters and are described by Sbordoniet aI. (1974, 1977). Several of the caves are ratherlarge and generally horizontal. Sumidero del Naranjoreceives some floodwater, and Cueva del Nacimientodel Rio Santo Domingo contains a large stream.Although most of the cave fauna remains to bereported on, 16 species have been identified, of whichthree are troglobites. The troglobites are the opilionidTroglostygnopsis anophthalma Silhavy, the glomeridesmidmilliped Glomeridesmus ?sbordonii Shear,and the trichopolydesmid milliped Caramba delnegroShear. The opilionid and trichopolydesmid millipedare endemic to the region, but G. sbordonii is alsoknown from Grutas del Cocona, Tabasco. The troglophilefauna includes the crayfish Procambarus (Austrocambarus)sbordonii Hobbs, the schizomid Schizomusarganoi Brignoli, the mysmenid spider Maymenamayana (Chamberlin and Ivie), the nesticidspider Eidmannella pallida (Emerton), the pyrgodesmidmilliped Synoptura italolegata (Shear), thesphaeriodesmid millipeds Sphaeriodesmus golondrinensisShear and S. redondo Shear, the roach Aglaopteryxchiapas Fisk, and a species of leiodid beetleof the genus Ptomaphagus (Adelops). With the exceptionof the two spiders and the milliped S. italolegataall of these species are endemic to the region. The spidersare widespread in southern Mexico and S. italolegatais also known from other parts of Chiapas eastinto the Yucatan Peninsula.Comitim de Dominguez, Chiapas.-This region islocated in the vicnity of the town of Comitan deDominguez in eastern Chiapas. The principal cavernousformation in the region is the Sierra MadreLimestone of Cretaceous age (Gu tierrez Gil, 1956).Only two caves have been studied in this area, thelargest of which is Cueva del Tio Ticho (Sbordoniet al., 1974). This cave is a sinkhole about 60 metersdeep, with a stream at the bottom. The other cave issmall and of little interest. Both caves are located atan altitude of about 1,700 meters.The fauna of the region is of considerable interestand further study should greatly increase the numberof species known. Of the 25 species recorded, five are47

troglobites: the trichoniscid isopod Brackenridgiaacostai (Rioja), the crab Typhlopseudothelphusamocinoi Rioja, the syarinid pseudoscorpion Pachychitragrandis Muchmore, an undescribed species ofschizomid of the genus Schizomus, and the carabidbeetle Mexanillus sbordonii Vigna Taglianti. Brackenridgiaacostai is also known from the Montebelloregion, and T. mocinoi has been found in the SanCristobal de las Casas region. Troglophiles of interestinclude the leptonetid spider Archoleptoneta obscuraGertsch, the nesticid spiders }

ites, have been reported. The large number of troglobitesindicates that additional discoveries of interestawait study. The only aquatic troglobite known fromthe region is the bogidiellid amphipod Bogidiella sp.ef. vomeroi Ruffo and Vigna Taglianti. Bogidiellavomeroi is known from the Simojovel and Tila region.The terrestrial troglobites are the opilionid Hoplabunuszullinii Silhavy, the trichopolydesmid millipedsCaramba grandeza Shear and Tylogoneus sp.,and the leiodid beetle Ptomaphagus (Adelops) sp.The troglophiles are the pyrgodesmid millipedsMyrmecodesmus inornatus Shear and Synopturazullinii (Shear) and the sphaeriodesmid millipedSphaeriodesmus trullatus Shear. The genus Carambais known only from caves in Chiapas; otherwise thefauna is typical of that of southern and easternMexico.Palenque. Chiapas.-This is a poorly definedregion located south of Palenque and includingSaito de Agua. The only cave so far visited is Cuevadel Salta de Agua, a large seasonal resurgence locatednear the town of Salta de Agua. Two species, anundetermined rhachodesmid milliped and an ant ofthe genusPachycondyla, have been identified.Rancho del Cielito, Chiapas.-This region is locatednear the village of Colonia Galeana in westernChiapas. The only caves studied in the area are Cuevade las Canicas at an elevation of 1.350 meters andCueva del Cerro Brujo at 1,320 meters. The first caveis a 20-meter-deep sinkhole leading to a small passage;the latter cave is complex and only partially ex plored.Both caves are described by Sbordoni et al. (1974).The fauna of this region remains poorly studiedand only seven species, of which three are troglobites,have been reported. The troglobites are the trichoniscidisopod Brackenridgia sp., the bogidiellid amphipodBogidiella sbordonii Ruffo and Vigna Taglianti,and the ricinuleid Cryptocellus sbordonii Brignoli.All are typical elements of the Chiapas fauna. Thetroglophiles are the mysmenid spider lHaymenasbordonii Brignoli and the nesticid spider Eidmannellapallida (Emerton). Maymena sbordonii isknown only from this region, while E. pallida is awidespread species throughout North America.San Cristobal de las Casas, Chiapas.~ This region,located on the plateau-like Sierra de San Cristobal,extends from San Cristobal de las Casas to the vicinityof Comitan de Dominguez. Karst developmentis considerable and includes areas of dolinas withlittle relief. Other areas, however, contain tlle greatestdevelopment of karst in North America, with hugesinkholes, sinking streams, poljes, and pinnacles. Thegeology along the highway from San Cristobal de lasCasas to Comitan de Dominguez is discussed byGutierrez Gil (1956). The caves, many of which aredescribed by Sbordoni et al. (1974, 1977), areprobably formed in the Sierra Madre Limestone andoccur at elevations from 1,820 to 2,520 meters. Thecaves range from shallow pits with minimal horizontalex tent to large horizontal caves and stream systems.Grutas de Rancho Nuevo is located about 10 kilometersfrom San Cristobal de las Casas on the highwayto Comitan de Dominguez and is an extensive horizontalsystem with more than two kilometers of surveyedpassage (Thompson, 1972). Several major cavesare known in the vicinity of Huixtan. The largest ofthese is Sumidero Yochib with a depth of 213 metersand a surveyed length of 3,316 meters (Van Note,1977). Sumidero Yochib contains one of the largerunderground streams in Mexico and is very dangerous.Other major stream caves in the Huixtan area areCueva Mapachero and Salida de Cruz Pilal.The San Cristobal de las Casas region is one of thebetter studied in Chiapas with respect to its invertebratecave fauna despite the fact that much of thematerial collected by the Italian biospeleologistsremains unstudied. The 38 investigated caves contain53 identified species, of which 14 are troglobites.The aquatic fauna is particularly interesting and includesseven troglobites: the dimarcusid flatwormOpisthobursa josephinae Benazzi; the dugesiid flatwormDugesia mckenziei Mitchell and Kawakatsu; theasellid isopod Caecidotea chiapas Bowman; the bogidiellidamphipods Bogidiella orchestipes Ruffo andVigna Taglianti, B. sbordonii Ruffo and Vigna Taglianti,and B. tabascensis Villalobos; and the crabTyphlopseudothelphusa mocinoi Rioja. With theexception of B. tabascensis (known also from Grutasdel Cocona, Tabasco) and T. mocinoi (known alsofrom the Comitan de Dominguez region) all of thesespecies are endemic to this region. The terrestrialtroglobites are an undescribed species of schizomid ofthe genus Schizomus, the opilionid Troglostygnopsisanophthalma Silhavy, a glomerid milliped of thegenus Glomeroides, the cleidogonid milliped CleidogonaJelipiana Shear, a collembolan of the subfamilyParonellinae, and the carabid beetle Chiapadytesbolivari Vigna Taglianti. The last species is the onlytroglobitic trechine beetle in Chiapas and is endemicto this region.The only aquatic troglophile in the region is theubiquitous amphipod Hyalella azteca (Saussure).Terrestrial troglophiles of interest are the agelenidspider Tegenaria florea Brignoli; the leptonetidspider Archoleptoneta arganoi (Brignoli); the nesticidspiders Eidrnannella pallida (Emerton) and Gaucelmuscalidus Gertsch; the pholcid spiders Coryssocnemispecki Gertsch, Modisimus propinquus O. P.-49

Cambridge, and Pholcophora bispinosa Gertsch; thericinuleid Cryptocellus bolivari Gertsch; the pyrgodesmidmillipeds Myrmecodesmus fissus (Causey)and Synoptura tioticho (Shear); the sphaeriodesmidmillipeds Sphaeriodesmus cruzbelem Shear and S.zontehuitz Shear; the roach Aglaopteryx chiapasFisk; and the histerid beetle Anapleus wenzeli V0mero. All of these species are typical representativesof the cavernicole fauna of eastern and southernMexico.Simojovel and Tila, Chiapas.-This region islocated in north-central Chiapas in the vicinity of thetowns of Simojovel and Tila. The eight caves studiedin the region are described by Sbordoni et aI. (1977)and are located at elevations of 810 to 1,790 meters.The caves are generally large and most contain eitherstanding or running water. Several remain largely unexploredand may prove to be quite extensive.Of the 17 identified species, four are troglobites.The aquatic fauna includes the asellid isopod Caecidoteazullinii Argano, the bogidiellid amphipod Bogidie/lavomeroi Ruffo and Vigna Taglianti, and the trichodactylidcrab Trichodactylus (Rodriguezia) mensabakCottarelli and Argano. The last species is theonly troglobite in the family Trichodactylidae. Thetwo crustaceans are known with certainty only fromthis region, but are closely related to species occurringelsewhere in Chiapas, Oaxaca, and Veracruz. Theonly terrestrial troglobite is the euryurid millipedPolylepiscus vomeroi Shear. This species is of specialinterest in that it is the only troglobite in the familyEuryuridae. The troglophile fauna includes the nesticidspider Gaucelmus calidus Gertsch and the pyrgodesmidmilliped Synoptura italolegata (Shear); bothare widespread in southern Mexico.Tuxua Gutierrez, Chiapas.-This region is locatedin the Chiapa Depression and includes caves in thevicinity of Tuxtla Gutierrez, Berriozabal, and Ocozocoautla.Most of the caves are described by Sbordoniet aI. (1974, 1977) and are located at elevationsranging from 460 to 950 meters. The region is characterizedby relatively low relief and numerous smallmesa-like hills. Gutierrez Gil (1956) and Olivas(1956) have discussed the geology of the region.Cretaceous and Eocene limestone crops out in thearea. The principal Cretaceous formations are theOcozocoautla and Sierra Madre Limestones, andmost of the caves probably are formed in one of theseformations. Cueva Cerro Hueco, a horizontal cavefrom which a stream emerges, may be formed in anunnamed Eocene limestone. Cave development inthe area is generally horizontal, although a few sink.holes up to 75 meters in depth are known. Many ofthe caves visited are small, but some are large and ofgreater interest. Hoyo de Don icho to the west ofOcozocoautla contains a stream passage severalhundred meters long at the bottom of a 20-meterdeepsinkhole. The largest cave in the area is Cuevadel Chorreadero. This is a major stream cave 345meters deep and with more than three kilometers ofsurveyed passage (Thompson, 1972; Shawcross et aI.,1974).The Tuxtla Gutierrez region is the best-studiedpart of Chiapas, with 81 species having been identifiedfrom the 22 investigated caves. The fauna, however,includes only one possible troglobite, an undescribedmilliped of the family Trichopolydesmidae.The aquatic fauna includes, in addition to severalspecies of nematode, the crayfish Procambarus(Austrocambarus) mirandai Villalobos and the catfishRhamdia guatemalensis GUnther. Both endemicand more widely distributed species are representedin the terrestrial troglophile fauna. The endemicspecies are the isopod Trichorhina vandeli Rioja; thepholcid spiders Coryssocnemis facetus Gertsch, Metagoniamcnatti Gertsch, and .Modisimus tzotzileBrignoli; the scytodid spider Loxosceles tehuanaGertsch; the opilionid Akdalima vomeroi Silhavy;the pyrgodesmid milliped Synoptura rodriguezi(Shear); the roach Pseudomops nigrimaculis Fisk;and the scydmaenid beetle Euconnus (illadagassoconus)arganoi Franz. Other troglophiles of interestare the schizomid Schizomus portoricensis (Chamberlin),the amblypygid Paraphrynus aztecus (Pocock),the mysmenid spider llrIaymena mayana(Chamberlin and Ivie), the nesticid spiders Eidmannellapal/ida (Emerton) and Gaucelmus calidusGertsch, the pholcid spiders Modisimus propinquusO. P.-Cambridge and Physocyclus globosus (Taczanowski),the opilionid Arganotus macrochelis (Goodnightand Goodnight), the pyrgodesmid millipedSynoptura italolegata (Shear), the carabid beetlePlatynus colibor Whitehead, and the leiodid beetleDissochaetus curtus Portevin. All of these species aretypical representatives of the fauna of southernMexico.Tapijulapa, Tabasco.-This region is located nearthe town of Tapijulapa in south-central Tabasco nearthe Chiapas border. The only cave studied in the areais Cueva del Azufre a few kilometers south of Tapijulapaat an altitude of 50 meters. The geology ofthis region has been examined by Salas and LopezRamos (1951) who report the presence of Cretaceouslimestone of the Guayal Formation in the area ofTapijulapa. A stream of sulphurous water flows fromthe cave, which is about 500 meters long and containsa large bat colony in one part.50

The fauna of the cave remains somewhat imperfectlyknown, but 20 species, of which two are troglobites,have been identified. The two troglobites areflatworms of the genus Dugesia and a blind populationof the fish Poecilia sphenops Valenciennes.Troglophiles recorded from the cave include thetrichodactylid crab Trichodactylus (Rodriguezia)bidens Bott, the chernetid pseudoscorpion Lustrochernes?minor Chamberlin, an undescribed amblypygidof the genus Phrynus, an araneid spider of thegenus Tetragnatha, the mysmenid spider Maymenamayana (Chamberlin and Ivie), the nesticid spidersEidmannella pallida (Emerton) and Gaucelmus calidusGertsch, and a hemipteran of the genus Belostoma.Teapa, Tabasco.-This region is located near thetown of Teapa in south-central Tabasco along theborder of Chiapas. Contreras V. (1958) has discussedthe geology of this region and reported thepresence of the Sierra :\1adre Limestone. The karstin this area is well developed with lapiez, pinnacles,towers, and other forms occurring in abundance.The known caves all occur at elevations of about40 meters, but immediately to the south the mainmass of the mountains of Chiapas rises to more than2,700 meters. Many caves are known to exist in thisregion from which numerous species of bat have beenreported. Only five caves, however, have been studiedwith respect to their invertebrate fauna. With the exceptionof the large, semi-commercial Grutas delCocona, all are small and of limited interest. Grutasdel Cocona, however, is an extensive horizontal cavewith almost one kilometer of passage. The cave is afossil resurgence and deep lakes occur there withoccasional running water in the back portions of thecave.Although 59 species have been reported from thecaves of the Teapa region, 23 of these are bats. Theremaining fauna is particularly rich and includeseight troglobites. Three aquatic troglobites areknown: the dimarcusid flatworm Opisthobursa mexicanaBenazzi, the bogidiellid amphipod Bogidiellatabascensis Villalobos, and the palaemonid shrimpMacrobrachium acherontium Holthuis. The shrimpand flatworm are endemic to this region, and theamphipod has been tentatively identified from cavesin Chiapas. The terrestrial troglobites are the hyidpseudoscorpion Mexobisium maya Muchmore, theschizomid Schizomus pecki Rowland, the amblypygidParaphrynus chiztun (Rowland), the glomeridesmidmilliped Glomeridesmus sbordonii Shear,and the histerid beetle Troglobacanius maya V0mero. With the possible exception of G. sbordoniiwhich has been tentatively identified from a cave inChiapas, all of these species are endemic to the Teaparegion. All are also closely related to other speciesfrom eastern and southern Mexico. Glomeridesmussbordonii is of interest in being the only troglobiticmember of the order Glomeridesmida in the NewWorld. The genus Troglobacanius is otherwise knownonly from caves in San Luis Potosi and Tamaulipas.The troglophile fauna includes a schizomid Schizomustrilobatus Rowland, an amblypygid of thegenus Phrynus, the mysmenid spider Maymenamayana (Chamberlin and Ivie), the oonopid spiderTriaeris patellaris Bryant, the pholcid spider Modisimusiviei Gertsch, the pyrgodesmid milliped Cry~tyma cocona Shear, nicoletiid thysanurans, the leiodidbeetle Ptomaphagus (Adelops) tabascensis Sbordoni,and the scydmaenid beetle Scydmaenus teapanusFranz. These species all belong to generacommon to southern and eastern Mexico.Augustine, Cayo, Belize.-This region is located inwest-central Belize on an outcrop of CretaceousEocene limestone to the west of the Maya Mountainsand south of the Rio On. Elevations in this regionrange from 400 to 500 meters. Very little work hasbeen done and there is confusion about how manycaves have been studied. Seven names have beenapplied to caves in the area, but some of these areprobably synonymous. The caves for which evenfragmentary descriptions are available are associatedwith streams, the most notable of which is the RioFrio which passes through the region. The geologyof the general area has been briefly discussed byDixon (1957).The fauna of the Augustine region is poorly knownand of 35 species reported, only 23 are closely associatedwith caves. The only troglobite known is ahyid pseudoscorpion, Mexobisium goodnight Muchmore.Other troglophiles include the mysmenidspider Maymena mayana (Chamberlin and Ivie), thescytodid spider Loxosceles yucatana Chamberlinand Ivie, the opilionids Erginulus serratipes (Cambridge)and Cynortina acanthotibialis Goodnight andGoodnight, the pyrgodesmid miIlipeds Myrmecodesmusbrevis Shear and Rettenmeyeria cryptymoidesShear, and the leiodid beetles Dossochaetus hetschkoiReitter and Ptomaphagus (Adelops) barbaraePeck. Of particular interest is the presence in thecaves of large numbers of psychodid flies, some ofwhich are apparently troglophilic. The most abundantof these are Lutzomyia beltrani (Vargas andDiaz Najera), 1. deleoni (Fairchild and Hertig), 1.shannoni (Dyar), and 1. trinidadensis (Newst.). Thefauna of this region is typical of that of southernMexico, although the presence of psychodid flies51

of the genus Lutzomyia has not been reported out·side of southern Belize. The miJIiped R. cryptymoidesis also the only cavernicole recorded for that genus.Caves Branch, Cayo, Belize.-This region is locatedin central Belize north of the Maya Mountains. Theterrain is characterized by well-developed cockpitkarst, with hundreds of steep hills separated by enclosedvalleys. Drainage is almost entirely subterraneanand numerous major caves are found in theregion. The caves are formed in Cretaceous limestoneand occur at elevations from 40 to 120 meters(Dixon, 1957; Graham et al., 1980). Caves Branch isa stream flowing off the Paleozoic rocks of the MayaMountains, across the karst surface eventually toenter the Sibun River. For about three kilometers ina direct line, and much more underground, it passesthrough a major cave, the Caves Branch Cave System.Most of the caves in the area have numerous ent·rances, known locally as breakou ts, where collapsehas opened the underground stream passage to thesurface. Footprint Cave, a major cave, has only oneentrance but more than seven kilometers of surveyedpassage and includes both an upper-level dry area anda lower active stream passage (Graham et al., 1980).The Caves Branch region is still inadequatelystudied bIOlogically, but 33 species, including seventroglobites, have been identified. The troglobitesinclude the vachoniid pseudoscorpion Vachoniumbelizense Muchmore, an undescribed schizomid ofthe genus Schizomus, an undescribed species ofcharontid amblypygid, the pholcid spider Melagoniajarmila Gertsch, the opilionids Cynortina misticaGoodnight and Goodnight and Stygnomma peckiGoodnight and Goodnight, and the cambalid miJIipediarmilka alba Shear. This fauna is particularly inte·resting in that it contains several quite distinctiveelements. The charontid amblypygid is the only trog·lobitic representative of this family in North America;the opilionid genera Cynortina and Stygnomma arenot otherwise known to have troglobites on mainlandNorth America; and iarmilka alba is an aberrantmember of a family not otherwise known south ofOaxaca, Mexico. The pseudoscorpion genus Vachoniumis known only from caves in the Yucatim Penin·sula and Belize. The remaining troglobites, as well asthe troglophiles, are all typical members of the caver·nicole fauna of southern Mexico. Troglophiles ofinterest include the spiraxid snail Streptostyla meri·dana meridana (Morelet), the amblypygid Paraphry·nus raptator (Pocock), the mysmenid spider May·menu mayana (Chamberlin and Ivie), ochyroceratidspiders of the genera Ochyrocera and Theotima, anopilionid of the genus Stygnomma, the pyrgodesmidmilliped Myrmecodesmus unicorn Shear, and theleiodid beetle Ptomaphagus (Adelops) barbarae Peck.Cobin, Alta Verapaz. Guatemala.-This region islocated in the vicinity of the town of. Coban insouthwestern Alta Verapaz. The terrain in much ofthe region is characterized by a mature karst withnumerous uvalas, dolinas, and solution valleys and byalmost complete internal drainage. The principallimestone unit in the area was mapped by Walper(1960) as undifferentiated Ixcoy and Coban Formations.Only four caves have been studied with respectto their invertebrate fauna. One, Cueva Chirrepeck, islocated to the south of Coban; two, Grotte C3 andGrotte G3, are in the Sierra de Pampur to the west ofCoban; and the third, Cueva Chiacam, is in the Sierrade Chama to the northeast of Coban. Nothing isknown of the nature of Cueva Chirrepeck, but theremaining caves are described in Delamare Deboutteville(1976) as being small and containing water.Most of the collecting in this region was done bySwiss and French biospeleologists and reports onmost of their material have not yet been published.The eighteen species which have been identified includefive troglobites and several troglophiles ofinterest. The troglobites are the pseudothelphusidcrabs TyphLopseudothelphusa juberthiei DelamareDeboutteville and T. mitchelli Delamare Deboutteville,the chthoniid pseudoscorpion Paraliochthoniusstrinatii Beier, the trichopolydesmid milliped Chirrepeckialyncilecta Hoffman, and the campodeid dipluraniuxtlacampa hauseri Conde. The crab genusTyphlopseudothelphusa is otherwise known only bya species in Chiapas and the dipluran genus iuxtlacampaby a species in Guerrero. Chirrpeckia is amonotypic genus and P. strinatii is the only troglobitein the genus in North America. Troglophiles ofinterest include the neobisiid pseudoscorpion Ideobisiumsimile Balzan, the mysmenid spider Maymenamayana (Chamberlin and Ivie), the nesticid spiderGaucelmus strinutii Brignoli, the opilionid Arganotusstrinatii Silhavy, and the pyrgodesmid millipedSynoptura petrinus (Hoffman). The last three speciesare endemic to this region, but otherwise the fauna istypical of southern Mexico.Lanquin. Alta Yerapaz. Guatemala.-This regionis located in east-central Alta Verapaz in the vicinityof the town of Lanquin. The geology, with specialreference to karst morphology and speleogenesis, hasbeen discussed by Smith (1968). The principal cavebearingformations in the area are the Campur Formationof Cretaceous age and the Chochal Formation,probably of Permian age. The largest cave in theregion, Grutas de Lanquin, is formed in the CampurFormation. The terrain is typical of many tropical52

egions and is referred to by Smith (1968) as kegelkarst.This is very similar to karst forms also referredto as mogote karst and cockpit karst. It is characterizedby isolated conical hills separated by funnelshapeddolinas, usually floored with red soil. In otherparts of the region karst development is less matureand is characterized by shallower dolinas with moregentle slopes. Elevations range from about 300 tomore than 1,500 meters. Most of the caves are rathersmall and generally horizontal, but Grutas de Lanquinis extensive. This commercial cave, a map of which isin Gurnee (1968), contains more than a kilometer ofsurveyed passage. A large stream flowing through partof the cave emerges below the entrance to becomethe Rio Lanquin.The Lanquin region is the best-studied part ofGuatemala, with 47 species, four of which are troglohites,having been identified to date. The only aquatictroglobite is an undetermined flatworm of the familyDimarcusidae. The terrestrial troglobites are the hyidpseudoscorpion Mexobisiumguatemalense Muchmore,the collembolan Pseudosinella Jinca Christiansen, andthe leiodid beetle Ptomaphagus (Adelops) giaquintoiJ eannel. The flatworm and IH. guatemalense areknown only from this region, but the other speciesare also found in the adjacent Senahu region. Endemictroglophiles include the pseudothelphusid crabIsabellagordonia (Isabellagordonia) longipes Pretzmann,the amblypygid Paraphrynus emaciatus Mullinex,the pholcid spider Modisimus gracilipes Gertsch,the scytodid spider Loxosceles guatemala Gertsch,and the gryllid cricket Arachnomimus cavicolaSaussure. More widespread troglophiles include theoonopid spider Triaeris patellaris Bryant, the pholcidspider Metagonia blanda Gertsch, the pyrgodesmidmilliped Cryptyma guatemala Shear, and scydmaenidbeetles of the genera Euconnus (Napochus) andScydmaenus.Raxruja, Alta Verapaz, Guatemala.-This region,located in the vicinity of the town of Raxruja innorthern Alta Verapaz, has been studied by Frenchspeleologists. The geology, physiography, and speleogenesisof the region have been discussed by Courbonand Dreux (1976). The principal formations in theregion are the Sepur, Campur, and Coban Formationsof Cretaceous age. Karst development, which is typicalof that known as kegelkarst, is primarily formedon the Campur Formation. Of the three caves studiedin the area, two are quite small. The third is the Sistemadel Rio Candelaria which represents the undergroundcourse of the Rio Candelaria and actuallyconsists of several isolated caverns having a total ofabout 17 kilometers of passage. A map of the cave isin Delamare Deboutteville and Juberthie (1976).Essentially nothing has yet been published on thecave fauna of this region and only four species haveso far been identIfied. Two species of shrimp otthe genus Macrobrachium have been recorded forSistema del Rio Candelaria. The crayfish Procambarus(Austrocambarus) pilosimanus (Ortmann) is atroglophile in the other caves of the region.Senahu. Alta Verapaz. Guatemala.-This region islocated to the southeast of the Lanquin region inextreme eastern Alta Verapaz. The geology, physiography,and speleogenesis of the region are discussedby Smith (1968). The principal cave-forming limestonein the vicinity of Senahu is the Chochal Formationof Permian age. Karst development is less welldeveloped than in the Lanquin region, but numerouswidely spaced hills surrounding sinks occur. At higherelevations the terrain is characterized by shallow dolinas.Of the five caves investigated in this region, allare rather small with the exception of the Sistema deSeamay-Sejul which contains a stream and more than1.5 kilometers of surveyed passage.Of 18 species identified from the region, eight aretroglobites. The only aquatic troglobite is the bogidiellidamphipod Bogidiella holsingeri Ruffo and VignaTaglianti. Two species of terrestrial troglobite, thecollembolan Pseudosinella Jinca Christiansen and theleiodid beetle Ptomaphagus (Adelops) giaquintoiJeannel, are shared with the Lanquin region. Therhachodesmid milliped Aceratophallus scutigeroidesShear has been tentatively identified from a cave inChiapas. The remaining species are all endemic tothis region. These include the chthoniid pseudoscorpionAphrastochthonius verapazanus Muchmore,the telemid spider Telema mayana Gertsch, the tridontomidmilliped Tridontomus loomisi Shear, andthe carabid beetle Speocolpodes franiai Barr. Tridontomusloomisi belongs to a family of millipedsknown only from this region. Speocolpodes is anendemic genus but is closely allied to Platynus(Mexisphodrus) in southern and eastern Mexico.The remaining species are both typical representativesof the cavernicole fauna of southern Mexico.The troglophile fauna includes the pseudothelphusidcrab Isabellagordonia (Phrygiopilus) acanthophallus(Smalley), a diplurid spider of the genus Euagrus,the pholcid spider Metagonia blanda Gertsch, andthe pyrgodesmid milliped Cryptyma guatemala Shear.Altos Cuchumatanes. Huehuetenango. Guatemala.-The Altos Cuchumatanes is a range of mountainsextending across much of southern Alta Verapaz.The top of the range is plateau-like with extensivedevelopment of dolina karst. The geology of part ofthe region is shown on a geologic map published bythe Guatemala Instituto Geografico Nacional (1968b).53

The principal cavernous fonnation in the region isthe Ixcoy Limestone of Cretaceous age. Elevationson the plateau range from 2,500 to 3,300 meters.Most of the caves art~ small and many take the flow ofsmall streams. Sbordoni et al. (1977) described thecaves which have been studied.The Italian biospeleologists who studied thisregion have yet to publish most of the results of theirwork and only 13 species have been identified todate. The aquatic fauna includes two endemic species,the asellid isopod Caecidotea mitchelli Argano andthe bogidiellid amphipod Bogidiella pasquinii Ruffoand Vigna Taglianti. The terrestrial troglobites arethe philosciid isopod Colombophiloscia cavernicolaVandel (also known from Venezuelan caves) and thecarabid beetle Mayaphaenops sbordonii Vigna Taglianti.TroglophiIes of interest include the hyalellidamphipod Hyalella azteca (Saussure), a pyrgodesmidmiIliped of the genus Synoptura, and a carabidbeetle of the genus Paratrechus.Montafias de Cuilco, Huehuetenango, Guatemala.This region is located in southwestern Alta Verapazsouth and southwest of the town of La Libertad. Thetop of the Montanas de Cuileo forms a plateau atan average elevation of about 3,000 meters. The fivecaves visited occur at elevations of 2,880 to 3,120meters. The geology of the northern part of theregion has been studied by Boyd (1966), Davis(1966), and Anderson (1967). A geologic map of theregion has been published by the Guatemala InstitutoGeogritfico !\acional (1967, 1968a). The principalcavernous fonnation in the area is the Ixcoy Limestone(sometimes not differentiated from the CobanFormation) of Cretaceous age. Karst development onthe plateau is primarily dolina karst with almostcomplete internal drainage. This region has been littlestudied speleologically and few caves are known. Allof the caves which have been biologically investigatedare small, shallow pits with little horizontal extent.The fauna of this promising region is essentiallyunknown and only five species have been identified.No troglobites are known, and the only probabletroglophiles are an undetermined species of millipedof the family Peridontodesmidae, a rhachodesmidmilliped of the genus Aceratophallus, and a staphylinidbeetle of the genus Philonthus. The rhaphidophoridcricket Hypsobadistes gracilior Hubbell, becauseof its extremely long legs and delicate body, isprobablj' restricted to the use of caves, though stilldependent on the surface for food.Santa Ana Huista, Huehuetenango, Guatemala.This region is located in west-central Huehuetenangonear the Chiapas, Mexico, border. The investigatedkarst region is in the vicinity of the village of El Taba-cal about six kilometers west-northwest of the townof Santa Ana Huista. There has been little study ofthe area and only three caves have been examined fortheir invertebrate fauna. The principal limestone unitsin the region are the Ixcoy and Coban Fornlations ofCretaceous age. The Rio Huista sinks in a large caveknown as El Sumidero, but sumps after about 400meters. The stream re-emerges from a sump in thelargest cave in the region, Cueva de Agua Escondida;this large, treacherous stream has been surveyed foralmost four kilometers (Broughton and Boon, 1975).The only other cave for which information is availableis Cueva de las Calaveras, a small, little exploredpit about 10 meters deep (Sbordoni et al., 1977).Very little is known of the cave fauna of thisregiun and only 11 species, none troglobitic, havebeen reported. Troglophiles include the nesticidspiders Gaucelmus augustinus Keyserling and G. calidusGertsch and the endemic pyrgodesmid millipedSynoptura shawcrossi (Shear).Lago de Izabal, Izabal, Guatemala.-This region islocated in eastern Guatemala near the Gulf of Honduras.The principal city in the area is the port ofPuerto Barrios. There has been little study of thearea, with only two caves having been explored. One,Cueva de la Coche, is located near the Rio Dulceabout 2.5 kilometers from the Gulf. It is reported byPeck and Peck (1973) as being small and only about83 meters long. The larger cave, Gruta EI Silvino, islocated 34 kilometers west of Puerto Barrios to thesouth of Lago de Izabal. This cave, a map of which isin Gurnee (1962), contains more than 0.5 kilometerof dry passage, where an unexplored stream passageis encountered.Only 12 species, none troglobitic, have been reportedfor this region. Troglophiles include thegrapsid crab Sesarma (Holometopus) roberti H.Milne Edwards, the schizomid Schizomus silvinoRowland and Reddell, the mysmenid spider Maymenamayana (Chamberlin and Ivie), the ochyroceratidspider Ochyrocera formosa Gertsch, the pholcidspiders Metagonia blanda Gertsch and Ph 01cophora quieta Gertsch, the pyrgodesmid millipedCalymmodesmus inquinatus Causey, and scydmaenidbeetles of tl1e genus Scydmaenus. This fauna is typicalof Guatemala and southern Mexico.Flores, Petim, Guatemala.-This region is locatedin central Peten south of the town of Flores. Theterrain consists of the low, rolling hills which constitutethe lowland area of the Chiapas-GuatemalanHighlands province. Although five caves have beenbiologically studied, the only one for which informationis available is Cueva Jobitzinaj, located south ofFlores. A partial map of this poorly explored cave54

shows more than 0.5 kilometer of generally horizontalpassage (Gurnee, 1962).Only 15 species, none troglobitic, have been reportedfrom the caves of the region. The most interestingmember of the troglophile fauna is the ricinuleidCryptocellus cooki Gertsch, a species closelyrelated to C. pearsei Chamberlin from Yucatan. Theurocoptid snail Brachypodella speluncae (Pfeiffer) isalso a probable troglophile restricted to the Peb~n.The leiodid beetle Ptomaphagus (AdelopsJ barbaraePeck, tentatively identified from the region, is otherwiseknown as a troglophile in caves in Belize. Theremaining species are all also known as troglophilesin the caves of the Yucatan Peninsula and other partsof southern Mexico and adjacent Guatemala. Theseinclude the amblypygid Paraphrynus raptator (Pocock),the mysmenid spider Maymena mayana(Chamberlin and Ivie), the scytodid spider Loxoscelesyucatana Chamberlin and Ivie, the paradoxosomatidmilliped Orthomorpha coarctata (Saussure),and a pyrgodesmid milliped of the genus Synoptura.Yaxchilim, Peten, Guatemala.-This region islocated in the Peten east of the Rio Usumacinta andacross the river from the ruins of Yaxchilan in Chiapas,Mexico. Salas (1962) has studied the geology ofthe region; the caves are probably formed in theCretaceous Repasto Limestone. The three caves thathave been biologically investigated are located at elevationsof 100 to 240 meters. The work in this regionhas been done by Italian biospeleologists and most oftheir results remain unpublished. The caves aredescribed by Sbordoni et al. (1974). Two of the cavesare small, but Cueva de Juan Flores (=Cueva de Yaxchilan)contains about 840 meters of surveyed passage(McEachern, 1974).Only 13 species, none troglobitic, have thus farbeen reported from the caves of this region. Two ofthe troglophiIes, the diplurid spider Euagrus lynceusBrignoli and the oonopid spider Triaeris lacandonaBrignoli, are endemic to the region. The pyrgodesmidmilliped Calymmodesmus inquinatus Causey is alsoknown from caves in other parts of Guatemala. Theremaining troglophiles are closely allied with thefauna of other parts of Guatemala and southernMexico. These include a squamiferid isopod of thegenus Trich orhina, the schizomid Schizomus porto·ricensis (Chamherlin), the mysmenid spider Maymenamayana (Chamberlin and Ivie), the pholcid spiderPholcophora ?speophila (Chamberlin and Ivie), thescytodid spider Loxosceles yucatana Chamberlin andIvie, and a pyrgodesmid milliped of the genus Myrmecodesmus.Yucatan PeninsulaThe Yucatan Peninsula is a limestone platformprojecting northward from Central America. It isdivided politically into the Mexican states of Campeche,Quintana Roo, Yucatan, and eastern Tabasco;the Peten of Guatemala north of Flores; and Belizenorth of the Maya Mountains. Reddell (1977b) hassummarized the geology, physiography, and karstfeatures of the Mexican portion of the Peninsula.Northern Yucatan is generally low, with elevationsgradually increasing from sea level to about 30 metersin the south. The Sierra de Ticul rises abruptly fromthe southwestern Yucatan plains to elevations of 70to 100 meters. The Sierra de Bolonchen lies south ofthe Sierra de Ticul in the state of Campeche. Thenorthern part of the Sierra de Bolonchen is a distinctrange, but to the south it is broken into numerouslow hills separated by valleys, some containing pondsand lakes. This hilly region continues south into thePeten of Guatemala. The peninsular part of Belize isalso low, rolling terrain. North of the Rio Hondo,which serves as the boundary between Belize andMexico, essentially all drainage is internal. Except forthe Rio Champoton in Campeehe no streams of anyconsequence flow in the Mexican part of the Peninsula.Small lakes and ponds occur in the hilly regionsof Campeche and saline lakes are found in southernQuintana Roo, but otherwise there is not muchstanding water. The eastern Peten and Belize haveseveral sizable streams flowing into the Rio Hondo orCaribbean Sea. Parts of the Peten contain large closedbasins, some with permanent water but others, nearlyfilled with alluvium, form broad marshes during thewet season.With the exception of a few minor deposits ofgypsum and marl cropping out in southern QuintanaRoo and Campeche and alluvium on the floors of thebroader valleys in the Campeche hill district, all ofthe Mexican part of the Yucatan Peninsula is onlimestone. Limestone is also the principal rock typein the peninsular part of Belize and in the Peten ofGuatemala. It is in places covered by extensive alluvialdeposits, especially in the closed basins in thePeten, along the river valleys of the Peten and Belize,and along the Caribbean coast. Butterlin and Bonet(1963) mapped the stratigraphy of the Mexican partof the Peninsula including the extreme northern edgeof Belize and the Peten. The 1:500,000 geologic mapof Guatemala published by the Guatemala InstitutoGeografico Nacional (1975) includes the generalgeology of Belize and the Peten.Pleistocene and Holocene deposits crop out ineastern Campeche and along the northern coast ofYucatan, but most of the Peninsula is covered by55

ocks of Paleocene to Pliocene age. A few Cretaceousoutcrops occur in the Peten and Belize but are notextensive. The Pliocene and Miocene formationsgenerally crop out along the coast of the Peninsula,with Eocene and older rocks inland. The Sierra deTicul and most of the hill district of Campeche areformed of rocks of Eocene or Paleocene age.The Mexican part of the Yucatim Peninsula is thebest-studied area in Mexico or Central America withrespect to its invertebrate cave fauna. No caves haveyet been visited in either the Peten or northern Belize,although they are known to exist. A total of 216caves, cenotes, and other subterranean habitats havebeen biologically investigated in Campeche, QuintanaRoo, and Yucatan. Reddell (1977b) includes descriptionsof 191 of these localities. Study has been madeboth of large open-air cenotes and true caves. Theterm "cenote," though generally thought to apply toa deep, well-like cavity, may refer to either pond-likewater-floored sinkholes, deep water-floored shafts,or true caves with bodies of water in total darkness.The more open cenotes generally do not containtroglobites but may harbor species also found astroglophiles in the cavernous cenotes. Of approximately565 species known from the caves and cenotesof the Yucatan Peninsula, US are known only fromopen-air cenotes. The fauna of the Peninsula includes12 aquatic and 28 terrestrial troglobites. A generalsummary of the cavernicole fauna is in Reddell(1977b).Sierra de Bolonchen, Campeche.-The Sierra deBolonchen is located in northern Campeche immediatelysouth of the Sierra de Ticul. The region designatedby this name also includes all of the hill districtof Campeche. In much of this area there are numerousdistinct rounded hills, ranging in elevation from100 to 300 meters, and separated by large, flatbottomedvalleys up to five kilometers wide. Most ofthese valleys are dry, but some contain shallow lakes.The only prominent karst features in addition to theclosed basins are sumideros-caves which receive theseasonal flow of arroyos. Four limestone units cropout in this region: the Icaiche Formation, the Pisteand Xcabal Members of the Chichen Itza Formation,and undifferentiated rocks of Paleocene or Eoceneage. The Icaiche Formation is Paleocene or Eocenein age and is exposed only in southeastern Campecheand adjacent Guatemala. The only cave of notebelieved to be formed in the Icaiche Formation isVolcan de los Murcielagos, a large, tunnel-like cavemore than one kilometer long. Undifferentiated Eoceneor Paleocene rocks crop out in much of Campeche.Although large caves occur in these rocks inthe Sierra de Ticul, no caves of consequence havebeen found in them in Campeche. The Chichen ItzaFormation is Eocene. The Piste Member crops out inthe higher parts of the Sierra de Bolonchen south ofthe Sierra de Ticul and-continues south and east alongthe border of Campeche, Yucatan, and QuintanaRoo. The largest caves in Campeche, including largesumideros such as Grutas de Xtacumbilxunam andGrutas de San Antonio, are formed in this rock unit.The Xcabal Member crops out only in a small areabetween Champoton and Escarcega in western Campeche.One large cave, Grutas de Monte Bravo, isknown from this unit.The caves in this region are among the larger in thePeninsula and attain much greater depths than cavesin either the Sierra de Ticul or Coastal Plain. Grutasde Xtacumbilxunam with a depth of at least 105meters is the deepest cave in the Peninsula. Several ofthe caves contain deep standing pools, and at leastone (Grutas de San Jose) contains a flowing stream.The Sierra de Bolonchen is the least-known part ofthe Peninsula, but 94 species, of which 10 are troglobites,have been identified from the 18 caves studied.The troglobite fauna includes species known onlyfrom this region and species shared by other parts ofthe Peninsula. The endemic fauna includes the amphipodMayaweckelia yucatanensis Holsinger, the atyidshrimp Typhlatya campecheae Hobbs and Hobbs, thediplocentrid scorpion Diplocentrus mitchelli Francke,and the tetrablemmid spider Matta mckenziei Shear.Two species, the atyid shrimp Typhlatya pearseiCreaser and the palaemonid shrimp Creaseria morleyi(Creaser), are found throughout the Peninsula. Theremaining troglobites are known only from theCoastal Plain and the Sierra de Bolonchen. These arethe amphipod Mayaweckelia cenoticola Holsinger,the oonopid spider Oonops coecus (Chamberlin andIvie), and the pholcid spider Metagonia torete Gertsch.The only other troglobite known from the Sierra deBolonchen is an undetermined species of miIliped ofthe family Trichopolydesmidae.The troglophile fauna is generally shared by otherparts of the Yucatan Peninsula. Aquatic troglophilesinclude the physid snail Stenophysa sp., the copepodsMesocyclops ellipticus Kiefer and Paracyclops fimbriatus(Fischer), and the amphipod Hyalella azteca(Saussure). Troglophiles include the spiraxid snailsStreptostyla meridana meridana (Morelet) and S.ventricosula (Morelet); the schizomid Schizomusportoricensis (Chamberlin); the amblypygid Paraphrynusraptator (Pocock); the amaurobiid spiderGoeldia tizamina (Chamberlin and Ivie); the mysmenidspider Maymena mayana (Chamberlin andIvie); the nesticid spider Eidmannella pallida (Emerton);the oonopid spider Triaeris patellaris Bryant;56

the pholcid spiders Metagonia maya Chamberlin andIvie, M. yucatana Chamberlin and Ivie, and Pholcophoraspeophila (Chamberlin and Ivie); the scytodidspider Loxosceles yucatana Chamberlin and Ivie;the opilionid Erginulus bimaculata Goodnight andGoodnight; pyrgodesmid millipeds of the generaCalymmodesmus and Synoptura; rhachodesmid millipedsof the genus Aceratophallus; the spirostreptidmilliped Orthoporus solicolens Chamberlin; theleiodid beetles Dissochaetus hetschkoi Reitter andPtomaphagus (Adelops) tabascensis Sbordoni; scydmaenidbeetles of the genus Euconnus (Napochus);and the ant Paratrechina pearsei (Wheeler).Sierra de Ticul, Yucatan.-The Sierra de Ticul risesabruptly from the coastal plain in southern Yucatannear its border with Carnpeche. Elevations range from70 meters in the north to about 100 meters in thesouth. The only rocks cropping out in the region areundifferentiated limestone units of Eocene or Paleoceneage. Karst development is limited to caves andminor surface features, such as shallow solutionalweathering. The Sierra de Ticul is made up of twonarrow ridges separated by a shallow valley. Caves inthe first ridge receive no significant run-off and tendto have large collapse sinkholes dropping into largerooms. Some of the larger caves, such as Actun Xpukil,contain several large rooms connected by narrowpassages. Others, such as Actun Loltlin and ActlinSabaca, have long, linear passages. Caves in the secondridge tend to be entered by small vertical or nearverticalpits, some dropping more than 30 meters.Larger caves, such as ActUn Chac, usually consist ofsingle passages gradually descending to water. A fewof the caves in this part of the range receive considerablefloodwater.The fauna of the Sierra de Ticul is very wellknown with 290 species, including 17 troglobites,identified from the 41 caves studied. The aquatictroglobite fauna includes the cirolanid isopod Creaseriellaanops (Creaser), the atyid shrimp Typhlatyapearsei Creaser, and the palaemonid shrimp Creaseriamorleyi (Creaser). The two shrimp are known fromall parts of the Peninsula, while the isopod is alsoknown from the Coastal Plain. The terrestrial troglobitefauna includes species endemic to the Sierra deTicul and species found in other parts of the Peninsula.The endemic species are the philosciid isopodTroglophiloscia laevis Schultz, the scorpion Diplocentrusanophthalmus Francke, the vachoniid pseudoscorpionsVachonium boneti Chamberlin and V.cryptum Muchmore, the amblypygid Paraphrynusreddelli Mullinex, the agelenid spider Cicurina (Cicurella)maya Gertsch, the spirostrcptid milliped Orthoporuszizicolens (Chamberlin), and the collembolanMetasinella falcifera (Mills). Species shared with theCoastal Plain are the squamiferid isopod Trichorhinapearsei (Creaser), the amblypygid Paraphrynus chacmool(Rowland), the collembolans Cyphoderusinnominatus Mills and Troglopodetes maya (Mills),and the gryllid cricket Tohila atelomma Hubbell. Theonly other troglobite is an undetermined species ofmilliped of the family Trichopolydesmidae.The only aquatic troglophiles are the copepodsMesocyclops ellipticus Kiefer, Paracyclops fimbriatus(Fischer), and Diaptomus texensis M. S. Wilson.The terrestrial troglophile fauna contains about 79species, of which some are endemic to the Sierra deTicul. These include the diplocentrid scorpion Diplocentrusreddelli Francke, the syarinid pseudoscorpionPachychitra maya Chamberlin, the oonopid spiderOonops mitchelli Gertsch, the pholcid spider Pholcophoramaria Gertsch, and the rhachodesmid millipedsAceratophallus calcehtokanus Chamberlin andA. oxkutzcabus Chamberlin. Troglophiles restrictedto the Yucatan Peninsula are the acanthodrilid earthwormsBalanteodrilus pearsei Pickford and Eodrilusoxkutzcabensis Pickford; the c1ubionid spider Tixcocobamaya Gertsch; the oonopid spider Oonopsreddelli Gertsch; the pholcid spiders Metagonia ivieiGertsch, M. maya Chamberlin and Ivie, and M. yucatanaChamberlin and Ivie; the ricinuleid Cryptocelluspearsei Chamberlin and Ivie; the chelodesmid millipedChondrodesmus sabachanus Chamberlin; the pyrgodesmidmilliped Calymmodesmus viabilis (Chamberlin):the spirostreptid milliped Orthoporus solicolensChamberlin; the collembolan Lepidocyrtus pearseiMills; the ant lion Eremeleon longior Banks; and theant Paratrechina pearsei (Wheeler). Among the moreinteresting troglophiles with a wider distribution arethe cyclophorid snail Neocyclotus dysoni berendti(Pfeiffer); the pomatiid snail Choanapoma largillierti(Pfeiffer); the spiraxid snails Euglandina cylindracea(Phillips). Streptostyla meridana meridana (Morelet),and S. ventricosula (Morelet); the schizomid Schizomusportoricensis (Chamberlin); the amblypygidParaphrynus raptator (Pocock); the mysmenid spiderMaymena mayana (Chamberlin and Ivie); the nesticidspider Eidmannella pallida (Emerton); the scytodidspider Loxosceles yucatana Chamberlin and Ivie; thetetrablemmid spider Matta sbordonii (Brignoli); andthe leiodid beetles Dissochaetus hetschkoi Reitter andPtomaphagus (Adelops) tabascensis Sbordoni.Coastal Plain, Campeche, Quintana Roo, andYucatim.-The Coastal Plain of the Yucatan Peninsulaincludes all of the state of Quintana Roo, all of Yucatanexcept for the Sierra de Ticul, and a narrow stripalong the northwest coast of Campeche. Reddell(1977b) recognizes four subdivisions: the Coastal57

Beach and Supra-tidal Zone, the I\orthwesternCoastal Plain, the Northeastern Coastal Plain, and theEastern Block Fault District. The first and last ofthese are of limited speleological interest. The o~hertwo subdivisions exhibit quite different morphology,but are so similar faunally that they are here treatedas one unit. The northwestern Coastal Plain is characterizedby very low relief (usually less than twometers locally), little soil, Hat-lying limestont' anddolomite, and the absence of deep caves and cenotes.The water level is ~enerally less than 10 mcters helowthe surface. The Northeastern Coastal Plain has muchgreater relief and ex tensive karst development. Theterrain is largely characterized by the presence ofnumerous cenotes, some more than 30 meters deep.This is best seen in the vicinity of Chichen Itza. Inextreme northwestern Yucatan and adjacent QuintanaRoo, the terrain is more rolling with large shallowdolinas and few cenotes. At the highest part ofthe plain, near Coba in Quintana Roo. cenotes an:generally absent and several lakes may be found.Along the Caribbean coast in ()uintana Hoo cenotesare generally shallow and water occurs less than 10meters below the surfa

from the Coastal Plain are the ostracods Cypridopsisinaudita Furtos, C. mexicana Furtos, and C. yucatanensisFurtos; the pimelodid catfishes Rhamdia guatemalensisdecolor Hubbs, R. guatemalensis depressaBarbour and Cole, and R. guatemalensis stygaeaHubbs; and the cichlid fish Cichlasoma urophthalmusericymba Hubbs. Among the more abundant aquaticspecies with wider distributions are the copepodsEucyclops serrulatus (Fischer), Macrocyclops albidus(Jurine), Mesocyclops ellipticus Kiefer, 1I-lesocyclopsleuckarti (Claus), Mesocyclops (Thermocyclops)tenuis (Marsh), Paracyclops fimbriatus (Fischer),Tropocyclops prasinus (Fischer), and Diaptomus(Leptodiaptomus) novamexicanus Herrick; the melitidamphipod Quadrivisio lutzi (Shoemaker); and thelimnesiid mite Limnesia paucispina Wolcott.Terrestrial troglophiles known only from theCoastal Plain are the oniscid isopod Hoctunus vespertilioMulaik; the trichoniscid isopod Trichoniscushoctuni Mulaik; the chernetid pseudoscorpion Parazaonacavicola Chamberlin; the laelapid mite llaemolaelapsglasgowi (Ewing); the uropodid mite Uropodapearsei Wharton; the erythraeid mite Erythraeusbisetosa Wharton; the galumnid mite Calumna jacotiWharton; the oribatellid mite Oribatellu monospicusWharton; the oribatulid mite Scheloribates luchiliWharton; the pyrgodesmid millipeds Calymmodesmusalienus (Chamberlin), C. isidricus (Chamberlin),C. hoctunanus (Causey), and C. muruztunicus (Chamberlin);the rhachodesmid milliped Aceratophallushoctunanus Chamberlin; the sphaeriodesmid millipedCylionus kauanus Chamberlin; the psocid Psyllipsocusyucatan Gurney; and the ant Brachymyrmex cavernicolaWheeler. Troglophiles restricted to the YucatanPeninsula are the acanthodrilid earthworms Balanteodriluspearsei Pickford and Eodrilus oxkutzcabensisPickford, the amaurobiid spider Coldia tizamina(Chamberlin and lvie), the c1ubionid spider Tixcocobamaya Gertsch, the oonopid spider Oonopsreddelli Gertsch, the pholcid spiders Metagonia ivieiGertsch and M. maya Chamberlin and lvie; the ricinuleidCryptocellus pearsei Chamberlin and lvie; theopilionid Erginulus bimaculata Goodnight and Goodnight;the pyrgodesmid milliped Calymmodesmusviabilis (Chamberlin), the spirostreptid miIlipedOrthoporus solicolens Chamberlin, the collem bolanLepidocyrtus pearsei Mills, the ant lion Eremeleonlongior Banks, and the ant Paratrechina pearsei(Wheeler). Some of the more widely distribu tedtroglophiles are the cyclophorid snail Neocyclotusdysoni berendti (Pfeiffer); the pomatiid snail Chounopomalargillierti (Pfeiffer); the spiraxid snailsEuglandina cylindrucea (Phillips), Streptostyla meridanameridana (\10relet), and S. ventricosula (Morelet);the urocoptid snail Brachypodella dubia Pilsbry;the schizomid Schizomus portoricensis (Chamberlin);the amblypygid Paraphrynus raptator (Pocock);the mysmenid spider Maymena mayana(Chamberlin and Ivie); the nesticid spider Eidmannellapallida (Emerton); the pholcid spiders Modisimusiviei Gertsch, Pholcophora speophila (Chamberlinand Ivie), and Physocyclus globosus (Taczanowski);the scytodid spider Loxosceles yucatanaChamberlin and Ivie; the pyrgodesmid miIlipedSynoptura italolegata (Shear); the leiodid beetlesDissochaetus hetschkoi Reitter and Ptomaphagus(Adelops) tabascensis Sbordoni; and the ant Erebomyrmaurichi (Wheeler).DISTRIBUTIONThe distribution and relationships of the cavern icole fauna of Mexico, Guatemala, and Belize remainpoorly known. In few instances can we delineate therange of any species of troglobite. It possible, nevertheless,to perceive some general distributional patternsand to draw a few conclusions with respect tothe distribution and relationships of the cavernicolefauna of these three countries. Although it is temptingto speculate on the zoogeography of the cavernicolefauna as well, I feel it premature to do so at thistime. Far too little is known of the past climatichistory of the region and of the faunal distributionsand relationships to confidently analyze the zoogeographyof the fauna in any but the most general sense.It is beyond the scope of this study to discuss thegeologic, hydrographic, and other factors which areimportant in obtaining a general understanding of thedistribution and evolu tion of the cavernicoles ofMexico and Central America. This region is amongthe more complex in the world, and even a discussionof the general features of its geology, hydrography,climate, and vegetation would take far more spacethan can be devoted to it here. The regional geologyhas been discussed by Schuchert (1935) in a pioneeringwork. Maldonado-Koerdell (1964) and West (1964)have provided useful summaries of the geology ofCentral America. The geology of Mexico has beensummarized by Garfias and Chapin (1949). Thehydrography of Mexico has been discussed by de laO. de Carreno (1951, 1954) and that of MiddleAmerica by Tamayo and West (1964). Vivo Escoto(1964) has summarized the climate of Mexico andCentral America. A good introduction to the vegetationof the region is that of Wagner (1964). A fewsignificant studies of specific areas have been citedin the section on Cave Regions above.59

The aquatic fauna of Mexico and Central Americaincludes cavernicole species derived from both freshwaterand marine ancestors; the terrestrial faunaincludes species with both temperate and tropicalaffinities. The purpose of the present discussion is tooutline briefly the distribution of some speciesbelonging to each category.Aquatic TroglobitesMarine relicts.-Species which are apparentlyderived from marine ancestors include triclad turbellarians,isopods, amphipods, mysids, shrimp, andfish.The flatworm family Dimarcusidae is known onlyfrom Mexican caves and is apparently a member ofthe marine suborder Maricola and without obviousclose relatives. Two species of this family, Ophistobursamexicana Benazzi and O. josephinae Benazzi,occur in caves in Tabasco and Chiapas respectively.Both have been found in association with bogidiellidamphipods, another group of marine relicts.The isopod fauna includes representatives of thefamilies Cirolanidae, Stenasellidae, Anthuridae, andMicrocerberidae. The distribution of the Cirolanidaeis of particular interest. Most of the species areknown from one or more caves in the same karstregion. This is true of Speocirolana thermydronisCole and Minckley, Mexilana saluposi Bowman, andSphaerolana interstitialis Cole and Minckley. Speocirolanabolivari (Rioja) has been found only in theSierra de El Abra, Sierra de Guatemala, and Aquismonregions, contiguous karst regions along the easternface of the Sierra Madre Oriental in San LuisPotosi and Tamaulipas. Sphaerolana affinis Cole andMinckley occurs both in the Cuatro Cienegas deCarranza region, Coahuila, and Sierra de la Silla,Nuevo Leon. Creaseriella anops (Creaser) has beenfound both in the Coastal Plain and Sierra de Ticul ofthe Yucatan Peninsula. This is a continuous limestonearea, and subterranean dispersal is likely. Of someinterest is the distribution of Speocirolana pelaezi(Bolivar). This species is the most widely distributedaquatic troglobite in Mexico and has been found inthe Sierra de Tamaulipas and Sierra de Guatemala,Tamaulipas; the Sierra El Pino, Sierra de El Abra, andAquismon regions, San Luis Potosi; and the CuetzaIan region, Puebla. It is likely, therefore, that itsrange extends the length of the Sierra Madre Oriental.It has an altitudinal range of about 1200 meters.The stenasellid isopods of Mexico range fromCoahuila to Veracruz. The genus Mexistenasellus isrepresented by species in the Cuatro Cienegas deCarranza, Micas, and Atoyac regions. Each species isrestricted to subterranean habitats in its respectiveregion. The monotypic genus Etlastenasellus hasbeen collected only from a well in Oaxaca and ispresumably a phreatobite.The family Anthuridae contains two Mexicantroglobites, one each in a cave in the Atoyac regionof Veracruz and in the Micos region of San LuisPotosi.The only described microcerberid isopod knownfrom Mexico is Mexicerberus troglodytes Schultz,which appears to be endemic to the Sierra de Guatemala,Tamaulipas. Other species of this family occuras inhabitants of the interstitial zone in Cuba.Two families of amphipod, both apparently ofmarine origin, have contributed to the cavernicolefauna of Mexico and Guatemala. The Bogidiellidae,with the single genus Bogidiella, is represented bynine species in extreme southern Mexico (Oaxaca,Chiapas, Veracruz, Tabasco) and Guatemala (Huehuetenango,Alta Verapaz). Possibly undescribed specieshave been taken in southern Puebla and Campeche.Several of the species are known only from wells andmay be phreatobites, but others are clearly caveassociated. Two genera of the family Hadziidae havebeen described from Mexican caves (Mayaweckeliawith two species in Yucatan and adjacent Campecheand Mexiweckelia with two species in the CualroCienegas de Carranza region, Coahuila, and one speciesin the Rancho Descubridora region, Durango.The order Mysidacea includes four Mexican troglobitesin two families. The Lepidomysidae is representedby Spelaeomysis olivae Bowman from a cavein the Acatlan region of Oaxaca and by S. quinterensis(Villalobos) from caves in the Sierra de El Abraand Sierra de Guatemala. Two species of the familyMysidae are troglobites in Mexico: Antromysis (A.)cenotensis Creaser from the Coastal Plain of theYucatan Peninsula and A. (A.) reddelli Bowman froma cave in the Acatlan region of Oaxaca.Most of the troglobitic shrimp inhabiting the cavesof Mexico are probably derived from marine ancestors.This speculation is greatly strengthened by thefact that they are found only in the areas borderingthe Gulf of Mexico and Caribbean Sea. Three familieshave been found in Mexican caves; the Alpheidae,Atyidae, and Palaemonidae.The Alpheidae is represented only by Alpheopsisstygicola Hobbs from caves in the Acatlan region ofOaxaca. With the exception of two Old World species,all members of this family are marine.The Atyidae includes three species of the genusTyphlatya in the Yucatan Peninsula. Although thefamily Atyidae includes both freshwater and marinespecies, the distribution of the genus Typhlatya(Cuba, Puerto Rico, Dominican Republic, Ascension60

Island, Galapagos Islands, Yucatan Peninsula) indicatesthat the Mexican species are derived from oneor more ancestral marine species.The family Palaemonidae is generally well representedin subterranean waters and six species areapparent troglobites in Mexico. Troglocubanus perezfarfanteaeVillalobos is the only Mexican member ofa genus also known from caves in Cuba. The generaCreaseria and Neopalaemon are monotypic andknown only from species inhabiting caves in theYucatan Peninsula and the Valle Nacional region ofOaxaca, respectively. Both are probably of marineorigin.The blind fish Typhliasina pearsei (Hubbs) isknown only from caves in the Yucatan Peninsula andis clearly a marine relict having as its closest relative aspecies now inhabiting marine littoral waters.Most of the marine relicts inhabiting caves inMexico are found in the vicinity of the Gulf ofMexico or Caribbean Sea. The Yucatan species aredistributed along the ancient Pliocene shoreline andhave not yet been found significantly farther inland.Other species inhabit the Sierra Madre del Sur,Chiapas-Guatemalan Highlands, and Sierra MadreOriental along their faces bordering the Gulf ofMexico. It is likely that these species have all beenderived from ancestral species inhabiting shallowwaters of the Cretaceous sea. Some may have undergonea preliminary epigean freshwater stage beforeentering caves, but this is impossible to determine.Some of the phreatobitic species probably inhabitedan interstitial habitat grading from salt through brackishto fresh water. Species of amphipod and isopodfound in the subterranean habitats of the regions ofRancho Descubridora, Durango, and Cuatro Cienegasde Carranza, Coahuila, are certainly derived fromspecies inhabiting the Cretaceous embayment thatcovered the region. Holsinger (1977a) postulates alate Cretaceous or early Tertiary age for the invasionof cave habitats in this general area.Most of the species of marine relict known fromMexico and Guatemala occur at comparatively lowelevations, but a few (such as Opisthobursa josephinae,Mexicerberus troglodytes, and various speciesof Bogidiella) are found at higher elevations. In someinstances these distributions may reflect an originalpattern of invasion as the seas receded, but in othersit seems possible that migration occurred from lowerelevations via a flooded interstitial or cavernous zone.Speocirolana pelaezi is of interest in that it rangesfrom Tamaulipas to Puebla and inland from the Sierrade Tamaulipas to the Sierra El Pino along the westernface of the Sierra Madre Oriental. Dispersal amongthese regions is now not possible and the assumptionis justifiable that its distribution reflects multipleinvasions by its ancestral species and a subsequentlack of divergence.Freshwater derivatives.-Troglobites which haveapparently evolved from freshwater ancestors includediaptomid copepods, triclad turbellarians, asellid isopods,shrimp, crayfish, crabs, and fish. All of thesespecies are closely related to species now inhabitingepigean habitats in the same general area.Three troglobitic flatworms of the family Dugesiidaeare known from caves in Mexico, and other speciesawait description. All belong to the widespreadgenus Dugesia which is found in epigean habitatsthroughout Mexico. Two species (D. typhlomexicanaMitchell and Kawakatsu and D. barbarae Mitchell andKawakatsu) are known only from caves in the Sierrade Guatemala and are closely related to the troglophilicD. guatemalensis Mitchell and Kawakatsu fromthe same region. The third species (D. mckenzieiMitchell and Kawakatsu) is known only from caves inthe San Cristobal de las Casas region, Chiapas. All ofthe described species of troglobitic Dugesia areknown from high elevation caves, but possible troglobitesare known from lower elevations.The diaptomid copepod Diaptomus (Microdiaptomus)cokeri Osorio Tafall is known only from cavesin the Sierra de El Abra, San Luis Potosi. Althoughthe subgenus is monotypic, the genus Diaptomusranges widely throughout North America.Isopods of the family Asellidae are represented inthe caves of Mexico and Guatemala by five describedand several undescribed species of the genus Caecidotea.This genus ranges throughout North America,although only one epigean species, C. communis(Say), is known from Mexico. The described troglobitesare from caves in Huehuetenango, Guatemala,and Chiapas and Veracruz, Mexico.Three troglobitic shrimp from Mexico are probablyof freshwater origin. Macrobrachium villalobosiHobbs from the Acatlan region of Oaxaca and M.acherontium Holthuis from a cave in Tabasco arealmost certainly derived directly from a freshwaterancestor, albeit one which may itself have been partlydependent on marine waters for the completion of itslife cycle as are other species of the genus. The genusBithynops is known only from one troglobite and onetroglophile in caves of the Montebello region of Chiapas.It is likely that the troglobitic B. luscus Holthuishas evolved quite recently and possibly from the troglophilicB. perspicax Holthuis, which inhabits caves ofthe same region.Three troglobitic crayfish have been describedfrom Mexican caves: Procambarus (Austrocambarus)rodriguezi Hobbs from Veracruz, P. (A.) oaxacae61

oaxacae Hobbs from the Valle Nacional region ofOaxaca. and P. (A.). oaxacae reddelli Hobbs fromcaves in the Atoyae region ofV~racruz and the AcatIan and Valle I\acional regions of Oax aca. These speciesare both closely related to species now inhabitingthe same region.CraLs of the families Pseudothelphusidae andlrichodactyJidae are represented in the caves of;\lhico and Guatemala. Three pseudothelphusid crabsof UIC cave-restricted genus 1'yphlopseudothelphusahave been described, one from Chiapas, '\Iexico, andthe remaining two from Alta Verapaz, Guatemala.1~vphlopseudothelphusais closely related to the epigeangenus Patamacarcinus. The family Trichodactylidaeis represented only by Trichodactylus (Rodriguezia)mensabak Cottarelli and Argano from Cuevade ;\Iicolas Bravo, Chiapas. Its closest relative is anepigean species also known from Chiapas.The fish fauna of '\

Temperate relicts.-A few species of troglobite areknown from Mexican caves which belong to groupsunknown from the surface in this region. These includethe centipede Nuevobius cavieolens Chamberlin,millipeds of the families Cambalidae and Trichopetalidae,and collembolans of the genera Acherontidesand Oneopodura. In our present state of knowledgeof the endogean fauna of tropical and subtropicalregions, we can only speculate that thesegroups will not be found on the surface. It wouldappear, however, that they represent true relicts of afauna once widespread throughout Mexico but nowto be found only in caves.The centipede Nuevobius cavieolens Chamberlinis known only from Cueva de la Boca in the Sierra dela Silla, Nuevo Leon. The only other species in theI!tnus is a trog/obite from Tennessee (U.S.A.).The rnilliped families Cambalidae and Trichopetalidaeare unknown on the surface in Mexico orCentral America. It is likely that these species aresurvivor;; of a once-widespread fauna inhabiting thetemperate forests that once covered parts of thisregion. These forests are now represented only byrelict poekeb at higher elevations of the ChiapasGuatemalan Highlands, Sierra Madre del Sur, andSierra :\1adre Oriental.The family Camhalidae is an important element ofthe cave and endogean fauna of the United States.One aberrant species, larmilka alba Shear, is knownonly from a cave in Belize. The genus lvlexicambalaincludes described species from caves in Oaxaca, SanLuis Potosi, and Tamaulipas. It is probably a relictof a group once widespread in Mexico hut that isnow restricted to caves. Finally, Cambala speobia(Chamberlin) has been collected in Cueva de losLagos, Coahuila. This is a common species in thecaves of Central Texas immediately north of the RioGrande.The family Trichopetalidae is a significant group inthe cave and endogean habitats of the southeasternUnited States. In Mexico it is represented only by thegenus Mexiterpes with four troglobitic species fromcaves in San Luis Potosi and Queretaro. The mosthighly cave-adapted species (M. sabinus Causey)occurs at comparatively low elevations in the Sierrade El Abra. The remaining species, which occur atmuch higher elevations, retain ocelli and are apparentlymore recent troglobites. It would appear thatthese were inhabitants of a temperate forest whichbecame extinct earlier at lower elevations than athigher ones.Two genera of collembolans appear to be temperaterelicts in the caves of Mexico. The hypogastruridgenus Acherontides is known only from two Mexicancaves, A. atoyaeense Bonet in Veracruz and A. potosinusBonet in the Xilitia region of San Luis Potosi.Other species of the genus occur in Rumania, Japan,and Afghanistan. The oncopodurid genus Oncopoduraincludes O. atoyaeense Bonet from Veracruz andO. prietoi Bonet from Nuevo Leon and possiblyTamaulipas. Other species of the genus occur inEurope, Japan, and the United States.There are several other groups which are unknownfrom the epigean habitat in Mexico but include troglobites.Among the more important of these are sixdescribed and several undescribed millipeds of thefamily Trichopolydesmidae, agelenid spiders of thegenus Cieurina, and ptinid beetles of the genus Niptus.It is likely, however, that as the endogean faunaof Mexico becomes better known these groups willalso be found in surface habitats.The family Trichopolydesmidae is known from theepigeum in Mexico only by two species in BajaCalifornia. All of the remaining species are troglobites.These are divided among three genera, Carambawith three species in Chiapas, Speodesmus with onespecies in Tamaulipas, and Tylogoneus with twospecies in San Luis Potosi and Tamaulipas. Each ofthese species is known only from the type-locality.The spider genus Cicurina is a major element ofthe cave and epigean fauna of Texas and other par.tsof the United States. In Mexico three troglobites andone troglophile have been described. The troglobitesare known from caves in Yucatan, Tamaulipas, andCoahuila, and the troglophile from caves in Tamaulipas.With more collections from high elevations inMexieo, this genus will almost certainly be found inendogean habitats.Two species of beetle of the family Ptinidae areknown from ;\1exican caves: Niptus abseonditusSpilman, a possible troglobite from Hidalgo, and N.abstrusus Spilman, a troglophile from caves in Texas(U.S.A.) and Coahuila and Durango, Mexico. Neitherspecies has been collected from the surface, but it islikely that N. abstrusus or other species of the genuswill be found there with more collecting.Species with temperate affinities.-Several groupsof cavernicole are members of genera containing bothepigean and cavernicole species, but which havecloser affinities to temperate species than to speciesinhabiting Mexico. These include trichoniscid isopodsof the genus Braekenridgia, scorpions of the genusTyphloehaetas, pseudoscorpions of the genus Aphrastochthonius,and spiders of the genera Tegenaria,Leptoneta, and Nestieus.The isopods of the genus Brackenridgia are amongthe most commonly collected animals in the caves ofMexico. Although only five species have thus far been63

described, the genus occurs in caves from Nuevo Leoninto Guatemala The known distributions of the speciesare quite limited but will doubtless be found tobe much wider with study of unidentified material.All species of the genus known from Mexico are troglobites,as are two species found in Texas caves. Theonly epigean species of the genus is B. heroldi (Arcangeli)from caves and surface localities in California.The troglobitic species occur from low to high elevationsand presumably represent a relict of a widespreadfauna that ranged thoughout the western andsouthwestern United States into Central America buthas now become extinct on the surface, except in themountains of California.Another group which is probably a relict of aonce-widespread temperate fauna is the scorpiongenus Typhlochactas. This genus includes three describedtroglobites from caves in Veracruz, San LuisPotosi, and Tamaulipas, and a litter-inhabitingspecies from high elevation forests in Oaxaca. As insome other groups, the most highly cave-adaptedspecies occurs in low elevations. The evolution anddistribution of Typhlochactas are discussed in moredetail in the Systematic Review below.Knowledge of the pseudoscorpion fauna of Mexico,both cave and epigean, is very limited, but the chthoniidgenus Aphrastochthonius would appear to be amember of a group with significant temperate affinities.Four troglobites in this genus have been describedfrom this region, each known only from thetype-locality. These occur in Tamaulipas, San LuisPotosi, and Alta Verapaz, Guatemala. Other troglobitesin the genus have been found in Alabama, NewMexico, and Cuba. One epigean species has beendescribed from leaf litter in Chiapas.Agelenid spiders of the genus Tegenaria are animportant element of the cavernicole fauna of Mexico.This genus is frequently collected on the surface inthe United States, but appears to be rare in caves outsideof Mexico, where two troglobites and seven troglophileshave been described.The family Leptonetidae is a largely Nearcticgroup containing numerous troglobites and troglophilesin the caves of the United States and Mexico.With the exception of a few epigean species, mostof the records for this family south of the UnitedStates are from caves. This may reflect a bias forcave collections, but nevertheless the group as awhole is certainly aligned with the Nearctic fauna.Troglobites have been collected from caves in theSierra Madre Oriental from Tamaulipas, Queretaro,and Nuevo Leon. One species, Leptoneta limpidaGertsch from Cueva de los Riscos, Durango, is mostclosely related to species from Texas caves and isnow a relict isolated by the surrounding desert.Although some species of the spider family Nesticidaerange widely throughout North America wellinto Central America; the family has reached itsgreatest degree of radiation in Nearctic regions. Thegenus Nesticus is unknown from the surface inMexico and includes troglobites from Veracruz,Tamaulipas, Nuevo Leon, and Puebta.Species with tropical affinities.-The vast majorityof the cavernicole fauna of Mexico is made up ofNeotropical elements. In some instances all or mostof the species in these groups known from Mexico,Guatemala, and Belize are trog[ohites, but in mostcases there are closely relakd taxa both on the surfaceand underground. It i~ not possible to do morehere than to mention SOIlIl' of the more significantspecies and briefly ou tlin.: di~lribll tiona} patterns asthey are presently known.Two families of lIIilliped which include troglobitesreach their northern distributional limit in southernMexico. The Glomeridesmidal' ranges from Panamainto Mexico south of the Isthmus of Tehuantepec.The only Mexican species is the troglohite Clomeridesmussbordonii Shear from caws in Chiapas andTabasco. With the exception of one .:pii(ean species inTabasco, the family OniscodeslIIidat· is represented inMexico only by two troglobites in YCraITIlZ: Bonetesmusojo Shear from the Orizaba region and B. verusChamberlin from the Atoyac region.Two species of philosciid isopod are troglobitesin Mexican and Guatemalan caves; both helong togenera known only from the tropics. Colombophilosciacavernicola Vandel was described frolll caves inVenezuela and has since been reported by Schultz(1977) from a cave in Huehuetenango, Guatemala.Even if this species proves to be an undescribed one,it is obviously a tropical element and one of the fewhaving close affinities with the South Americanfauna. The only other species of the genus occur inEcuador and the Galapagos Islands. The other troglobiticphilosciid known from this region is Troglophiloscialaevis Schultz from a cave in Yucatan; theonly other species of the genus is a Cuban troglobite.Several tropical groups include species of troglobitein more northern Mexico, but which are otherwisenot found on the surface outside of southernMexico. Included among these is the sphaeroniscidisopod Spherarmadillo cavernicola Mulaik from cavesin the Sierra de El Abra and Sierra de Guatemala.This genus and its relatives are not found on the surfacenorth of Veracruz. The squamiferid isopodgenus Trichorhina includes epigean and cavernicolespecies in Mexico and Cuba. Three troglobites andone troglophile are known. The troglophile has been64

found in Chiapas, while the troglobites occur inYucatan, Veracruz, and San Luis Potosi. The troglobiteT. boneti Rioja from higher elevations in theXilitla region of San Luis Potosi is the only speciesto occur north of Veracruz. A third genus of isopod,Cylindroniscus, also has a similar distributionalpattern. Endogean species are k~lOwn from Cuba andYucatan, while troglobites occur in Yucatan, NuevoLeon, and San Luis Potosi.Two families of pseudoscorpion include representativesin both the Old World and New World tropics.The Vachoniidae contains one African genus andtwo Central American genera. The Central Americanspecies are all troglobites divided between twogenera: Vachonium with one species in Belize andfour in Yucatan and Paravachonium with two speciesin Tamaulipas. The family Hyidae contains severalepigean species from Asia and epigean and troglobiticspecies from the New World. Two of the genera,Leucohya with two species in isolated mountainranges of Nuevo Leon and Trogloh.ya with one specieseach in Chiapas and Oaxaca, contain only troglobites.The third genus, Mexobisium, contains troglobites inBelize, Guatemala, and Tabasco and Veracruz,Mexico. Epigean species have been described fromCuba and southern Mexico.One of the more interesting species known fromMexican caves is the cyphophthalmid Neogoveamexasca Shear from caves in the Acatlan region ofOaxaca. This is the only species of the suborderknown from Mexico and is a member of a genusotherwise found only in Brazil and Guyana.The milliped order Glomerida is largely tropical,although some species do occur in the United States.The genus Glomeroides, however, includes epigeanspecies in Mexico, Guatemala, and California (U.S.A.).The only species known from the surface in Mexicoare restricted to the extreme southern part of thecountry; troglobites have been taken from caves atlow elevations in Veracruz and from higher elevationsin the Sierra Madre Oriental of San Luis Potosi andTamaulipas.The carabid beetle tribe Trechini includes numerousspecies in the genera Paratrechus and Mexitrechusfrom high-elevation, endogean habitats insouthern Mexico and Central America. The northernmostspecies of Paratrechus is P. (Hygroduvalius)pallescens Barr, a troglobite from caves in Queretaro.Three trechine genera, probably derived from thesame lineage as Para trechus, are known only fromcaves: Chiapadytes with one species in the SanCristobal de las Casas region of Chiapas, Mayaphaenopswith one species from Huehuetenango, Guatemala,and Mexaphaenops with four high-elevationspecies in Queretaro, San Luis Potosi, and Tamaulipas.The latter genus also includes a species fromGrutas del Palmito, Nuevo Leon, which has apparentlybeen isolated by the surrounding desert.Most of the remaining species of troglobite knownfrom caves in this region are members of groupspresent both on the surface and in caves throughoutthe region, although some may not occur at higherelevations. It is not possible here to do more thanmention a few examples to illustrate various aspectsof their general distribution.Three arachnid orders which are almost entirelytropical in their distribution are the Schizomida,Amblypygida, and Ricinulei. These are all found insurface habitats throughout tropical and subtropicalAmerica and all contain both troglobite and troglophilerepresentatives.The order Schizomida ranges from South Americanorth into the southern United States and is animportant element of the cavernicole fauna of Mexico,Guatemala, and Belize. The family Protoschizomidaeincludes only four species, two of which are epigeanspecies from Colima and Tamaulipas. The genusAgastoschizomus includes two troglobitic species: A.lucifer Rowland from caves in the Sierra de EI Abra,San Luis Potosi; and A. huitzmolotitlensis Rowlandfrom the Xilitla region of San Luis Potosi. The latterspecies is the only described species in the order fromhigher elevations in Mexico. The family Schizomidaeincludes numerous troglophilic and troglobitic speciesfrom Guatemala into Nuevo Leon, Mexico. The eightpresumed troglobites are from Veracruz, Tabasco,San Luis Potosi, Oaxaca, Tamaulipas, and NuevoLeon. The more northern species appear to be isolatedin caves by the surrounding desert.Numerous species of amblypygid of the genusParaphrynus have been reported from caves. Speciesfrom Tamaulipas, Yucatan, Tabasco, and San LuisPotosi are troglobitic. All but one of these are lowlandtroglobites; P. velmae Mullinex occurs at higherelevations in the Xilitla and Aquismon regions of SanLuis Potosi.The order Ricinulei, once thought to be incrediblyrare and still seldom collected, has been found to be asignificant component of the cavernicole fauna ofparts of Mexico and Guatemala. Most of the describedspecies are epigean or troglophilic, but threeappear to be cave restricted: Cryptocellus sbordoniiBrignoli from Chiapas, C. osorioi Bolivar from SanLuis Potosi and Tamaulipas, and C. reddelli Gertschfrom the Sierra de la India of Durango. The last speciesis apparently isolated by the surrounding desert.The phalangodid harvestman genus Hoplobunusranges from Central America into Texas. This is a65

large genus with numerous epigean species, mostknown only from the southern part of its range.Troglobitic species have been described from Chiapas,Oaxaca, Queretaro, San Luis Potosi, Tamaulipas,and Nuevo Leon. With the exception of H. boneti(Goodnight and Goodnight) from caves of the Sierrade EI Abra and Sierra de Guatemala and H. osorioi(Goodnight and Goodnight) from three isolatedmountain ranges in Nuevo Leon, all of the species arerestricted to high elevations. Two additional high.elevation relicts, both assigned by Silhav), (1974) tothe genus Troglostygnopsis, may also belong in Hoplobunus.Two troglobitic species of Hoplobunus inTexas are apparently tropical relicts in a temperateregIon.The milliped family Rhachodesmidae ranges fromCentral America into Nuevo Leon and has been amajor contributor to the troglobite and troglophilefauna of this region. Six genera of this family containtroglobites, all but one of which also containepigean species. Aceratophallus ranges north intoChiapas and Yucatan, with several troglophiles knownfrom Yucatim. The only troglobite is A. scutigeroidesShear from Alta Verapaz, Guatemala, and possiblyChiapas. Acutangulus includes five species from Veracruz,but the only troglobite is A. alius Causey fromthe Orizaba region. Two troglobitic species of Ceuthauxushave been described, one from the Cacahuamilparegion of Guerrero and the other from Grutasdel Palmito, Nuevo Leon. This genus ranges fromMorelos and Veracruz into Coahuila. The genusPararhachistes includes two epigean and one troglobiticspecies in Guerrero. Several species of Strongylodesmushave been described from Mexico, but theonly troglobite is S. harrisoni Causey from the Sierrade Guatemala This is a fairly recent troglobite andmay now be restricted to caves at higher elevations.The most important genus of rhachodesmid millipedin Mexican caves is Unculabes. This genus includesonly five species, all known only from caves in SanLuis Potosi, Queretaro, and Tamaulipas. All but oneof these are troglobitic.Mexico contains a rich truglobitic gryllid cricketfauna, with four species in two genera described.Tohila atelomma Hubbell is a species known onlyfrom caves in the Yucatan Peninsula. It is apparentlymost closely related to the genus Paracophus.The latter genus is known only from the SierraMadre Oriental, where three troglobitic and fivetroglophilic species are found. The three troglobites-Pocaecus Hubbell from the Sierra de Guatemala,P. cladonotus Hubbell from southern SanLuis Potosi and adjacent Hidalgo, and P. lippusHubbell from tlle Sierra EI Pino region of San LuisPotosi-are restricted to high elevations.The last group to warrant mention here includesspecies belonging to genera which are widespread inboili tropical and temperate regions. Among the moreimportant of these groups are spiders of the familiesDipluridae, Theraphosidae, and Pholcidae; millipedsof ilie families Cleidogonidae and Spirostreptidae;entomobryid collembolans of the genus Pseudosinella;and beetles of the family Leiodidae.The spider suborder Mygalomorphae tends to bepoorly represented in caves, but two families includetroglobites in Mexico. The Dipluridae includes twodescribed troglobites, Euagrus anops Gertsch fromthe Xilitla region and E. cavernicola Gertsch from theSierra de Guatemala; both occur only at high elevations.The family Theraphosidae is represented inMexican caves by two troglobites: Schizopelmareddelli Gertsch from caves in the Acatlan region ofOaxaca and S. stygia (Gertsch) from the Xilitla region.The first species occurs at low elevations, whilethe latter is found at higher ones.One of the most characteristic features of thecavernicole fauna of this region is the presence incaves of numerous spider species belonging to thefamilv Pholcidae. Troglobitic species are found inthe genera JUetagonia, Pholcophora, and Psilochorus.lIetaf!oniaincludes lowland troglobites in Veracruz.Oaxaca, Tamaulipas, the Yucatan Peninsula, andBelize. Highland troglobites are found in the Sierra deGuatemala and Xilitla regions. Two of the three troglobiticspecies of Pholcophora in Mexico are knownfrom lowland eavt·s in Guerrero and the YucatimPeninsula. The third species of Pholcophora and thetwo troglohitic Psilochorus species are found in thecaves of Chihuahua and Durango where they are isolatedhy the surrounding desert.The cleidogonid millipt~d genus Cleidogona is ofinterest in that the most highly cave-adapted speciesin the genus, C. crucis (Chamberlin) from the Atoyacregion of Veracruz, is found at the lowest elevation ofany of the troglobites. The remaining troglohiticspecies of the genus are found at higher elevations inthe states of Oaxaca, Chiapas, and Tamaulipas. Thegenus Cleidogona ranges from Central America intothe United States; Shear (1972) speculates that thegenus originated in the Mexican highlands.Although only two species of the spirostreptidmilliped genus Orthoporus have become troglobites,they are of particular interest in that they are theonly troglobites in the genus. The species of Orthoporustend to have very wide ranges and to be highlyvagile. Orthoporus spelaeus Causey has been found66

only in one cave in the Coastal Plain of Yucatan; O.zizicolens (Chamberlin) appears to be restricted tocaves in the Sierra de Ticul, Yucatan.The collembolan family Entomobryidae includesseveral genera with cave representatives, but the mostimportant by far is Pseudosinella. This widely distributedgenus contains troglobitic species in Alta Verapaz,Guatemala, and Oaxaca and Tamaulipas, Mexico.The genus is now under revision, and more meaningfuldata on its distribution in Mexico and CentralAmerica should be forthcoming.One of the more important beetle families to inhabitcaves in North America is the Leiodidae. Numerousspecies of this family have invaded Mexican andCentral American caves, but only two troglobiteshave been described. These two species, Ptomaphagus(Adelops) troglomexicanus Peck from the Sierra deGuatemala and P. (A.) mckenziei Peck from the Purificacionregion of Tamaulipas and Nuevo Leon, areboth restricted to high elevations.As is apparent from the above, the terrestrial troglobitefauna of Mexico and Central America includesan assortment of species derived from ancestors witha variety of affinities. Perhaps one of the moreinteresting aspects of the fauna of this region is theadmixture of temperate and tropical derivatives inthe same caves. This alone is a good indication of thecomplexity of the past history of the region. Speciesderived from ancestors adapted for colder climatesnow coexist in the same caves with species derivedfrom those adapted for wamler climates.Also of no slight interest is the existence of numeroustroglobites in caves in areas of lowland tropicalforests. It has been one of the axioms of modembiospeleology (Vandel, 1964; \Iitchell, 1969a) thatterrestrial troglobites are rare in tropical regions. Thepresence of a rich troglobitic fauna at high elevationscan be readily explained by the model of Barr (I968a)in which past climatic changes extirpated the epigeanfauna, allowing for divergence to occur in the cavemicolepopulations of that fauna. Climatic fluctuationsduring the Pleistocene, though presumably less severein Mexico and Central America, certainly were feltat higher elevations and may have allowed for theextinction of the surface ancestors of the present-daytroglobites. The existence of a rich troglobitic faunaat lower elevations in tropical regions is now uncontested.Not only do we have a rich fauna in lowlandparts of Mexico and Central America, but studies byHowarth (1973) in Hawaii, and by Peck (1976) inJamaica and other islands of the West Indies provethe existence of a rich tropical troglobite fauna inother areas as well. Only with much additional study,both of existing collections and in under-investigatedareas, can a really comprehensive model be proposedfor the evolution of this fauna. It is possible, however,that the Pleistocene climatic fluctuations werefelt sufficiently at lower elevations to cause the extinctionof surface populations of many groups insome karst regions. The more vagile species, such asbeetles, were able to repopulate the areas beforedivergence occurred, while groups such as millipeds,isopods, and the smaller arachnids had time to becomegenetically isolated from the re-invading stock.The preponderance of the smaller, more delicatespecies in lowland tropical caves would seem to givecredence to this hypothesis.Terrestrial TroglophilesThe terrestrial troglophile fauna of Mexico, Guatemala,and Belize contains both species which areabundant on the surface and species known onlyfrom caves. Some of the latter may eventually proveto be cave restricted, although they do not yet exhibitthe reduction or loss of eyes and pigmentation tobe found in the troglobites. Other species known onlyfrom caves belong to groups of endogean forms inwhich eyes are absent and pigmentation reduced. Thepresent brief discussion of Mexican troglophiles willoutline the various groups of troglophiles withrespect to their general distribution. For furtherdetails on the distrihu tion of the more importantspecies, see the Systematic Review section whichfollows.A few species which are troglophilic in caves inthis region have extremely wide ranges. Among these,mention may be made of the nesticid spiders Gaucelmusaugustinus Keyserling and Eidmannella pallida(Emerton). These two species are important membersof the cavemicole communities of the United States,Mexico, and Central America. The preponderance ofrecords for the two species in caves in many areasreflects both a predilection for the cavernicole habitatand a bias towards cave collections. One other exampleof wide distribution should suffice here. Thisis that of the leiodid beetles of the genus Dissochaetus.Some species of this genus range from SouthAmerica into northern Mexico. Although cave recordsfor the southern part of the range are rare (as arecollections), numerous troglophile populations occurin Mexico, Guatemala, and Belize.Many species with more limited ranges will befound in caves wherever they are available. This isespecially true of the more mesic-adapted forms,such as many millipeds, isopods, arachnids, and softbodiedinsects. Examples include various species ofarmadillid and porcellionid isopod, rhachodesmid67

milliped, laniatorid harvestmen, and collembolan.Two specific examples may be cited, both belongingto comparatively rare arachnid orders. The schizomidSchizomus mexicanus Rowland and the ricinuleidCryptocellus pelaezi Coronado are abundant, both incaves and on the surface in the caves of the Sierra deEl Abra. During the wet season both species may befrequently collected under rocks in this area, as wellas in caves; but with the onset of the dry season bothspecies retreat into the soil and may be found only incaves. Another interesting species with respect to itsdistribution is Schizomus portoricensis (Chamberlin).This species is known throughout many parts oftropical America. Sexual populations are knownfrom the surface in mainland Mexico and from cavesin Chiapas; peripheral populations are apparentlyparthenogenetic. An analysis of cavernicole populationsin Yucatan indicate that facultative parthenogenesishas been a significant factor in the abundantcolonization of the caves of that region (Rowlandand Reddell, 1977).In some arid regions of northern Mexico speciesoccur only in caves, while being found on the surfacein more mesic areas of their ranges. Examples of thistype of distribution are to be found in the pholcidspiders, and most notably Physocyclus enaulusCrosby and P. hoogstraali Gertsch and Davis. Althoughit would he premature to do more than speculate,it is possible that cave populations of these andother species may now be isolated and will eventuallybecome genetically distinct from the parental stock.Numerous species of troglophile in the caves ofthis region are known only from caves. This is particularlytrue with respect to many arachnids andmillipeds. There can be little doubt but that many ofthese will eventually be found in endogean situations;a bias both for cave collections and the study ofcavernicoles by taxonomists has emphasized thecavemicole fauna at the expense of the related endogeanforms. Among the groups in which this occurrenceis most notable are schizomids, pseudoscorpions,opilionids, the smaller spiders, and the moredelicate millipeds. These groups tend to seek a moist,dark habitat of a sort frequently overlooked by thecasual collector. Collecting during drier periods inareas with marked dry and wet seasons will also failto turn up the more mesic-adapted forms. Thecavernicole habitat, with its nearly constant con·ditions of temperature and humidity, provides astable environment at almost all times and thus thechances are greater for finding these forms. Also inmany caves food is scarce in comparison with that onthe surface and even the casual visitor to a cave canquickly locate troglophile and troglobite speciesbecause of their concentration in a few areas whereorganic matter has accumulated. In the case of thesmaller, more secretive forms, such as the pyrgodesmidmillipeds and most pseudoscorpions, only intensivesearch, including the use of special extractiontechniques, will result in their discovery on thesurface.68

Systematic ReviewThe following is a review of the known cavernicolefauna of Mexico, Guatemala, and Belize, with specialemphasis on those species which are of unusual interestor which make a significant contribution to theecology of the cave habitat. This is in no sense a completelist of species recorded from the caves of thisregion. The tables which accompany each major sectionrecord the number of species of each familyknown from caves in this area; comparatively fewspecies are referred to by name in the text. No completelist of the known cave fauna of these countriesis available, although Reddell (1971b) includes allpublished records for Mexico up to 1971.The checklist of cave-adapted species is as completeas possible. As has been discussed above, the useof the term "troglobite" is arbitrary, and, since wecannot in many instances know if a species is restrictedto the cave habitat, I include in the checklistpart of this review all of those species which appearsignificantly more cave adapted than other relatedspecies. All references to each species are given withas accurate page citations as is possible. A questionmarkfollowing the year in the citation indicates thatthe paper in question is known to refer to that particularspecies but that I have not seen the paper. Typelocalitydata are given as listed in the original description;the correct name of the cave or locality is includedin parentheses. Cave names are those currentlyused by the Association for Mexican Cave Studies.Locations and alternate names of the caves are givenin the locality list in the Appendix.Phylum ProtozoaWith the exception of the study by Osorio Tafall(1943) of two caves in the Sierra de EI Abra, San LuisPotosi, there are no published records of free-livingprotozoans from caves in Mexico (see Table 4). Thisis a group worthy of study as shown by preliminarystudies of Protozoa in the United States and Europe.Class RhizopodeaOrder AmoebidaFamily AmoebidaeThe only species in this family reported fromMexican caves is an undetermined species of Amoebafrom Cueva Chica, San Luis Potosi.Class ActinopodeaOrder HeliozoidaFamily ActinophryidaeAn undetermined species of Actinophrys was reportedfrom Cueva Chica, San Luis Potosi.Order ArceUinidaFamity ArceUidaeArcella vulgaris Ehrenberg was found in Cueva deLos Sabinos, San Luis Potosi.Family DifflugiidaeCentropyxis aculeatus Ehrenberg was taken inboth Cueva Chica and Cueva de Los Sabinos, San LuisPotosi.69

Table 4.-Summary of cave inhabiting Protozoa.TroglobitesOther SpeciesRhizopodeaAmoebidaAmoebidae 0 1ActinopodeaHeliozoidaActinophryidae 0 1ArcellinidaArcellidae 0 1Difflugiidae 0 1ZoomastigophoreaKinetoplastidaTrypanosomatidae 0 3CiliateaGymnostomatidaColepidae 0 1PeritrichidaVorticellidae 0 1Total 0 9Class CiliateaOrder GymnostomatidaFamily ColepidaeColeps sp. d. hirtus (Muller) was collected fromCueva Chica, San Luis Potosi.Order PeritrichidaFamily VorticeUidaeVorticella sp. cf. microstoma Ehrenberg was collectedin Cueva Chica, San Luis Potosi.Phylum PoriferaClass DemospongiaeOrder HaplosderinaFamily SpongiUidaePenney and Racek (1968) described Spongillacenota from Cenote Grande de Xanaba and CenoteXtolok, Yucatan. These two cenotes contain largeopen-air bodies of water, and the sponge recoveredfrom them is certainly not a part of the true cavefauna.Phylum CnidariaClass HydrozoaOrder HydroideaFamily HydridaeOsorio Tafall (1943) reported an unidentified speciesof Hydra from Cueva de Los Sabinos, San LuisPotosi. This is surely a form washed into the cavefrom the surface.Phylum PlatyhelminthesThe phylum Platyhelminthes is represented incaves both by free-living members of the class Turbellariaand by numerous species of parasite of theclasses Trematoda and Cestoda (see Table 5). Theparasites have been found in bats, fishes, and amphibians;they are not discussed further since they arenot a part of the true cave fauna.Class TurbellariaOrder TricladidaThe triclad turbellarian fauna of Mexican caves ispoorly known in comparison to that of the UnitedStates. Many collections have been made recently inMexican and Guatemalan caves, and once this materialhas been studied we will have a far better conceptof the nature and relationships of the Mexicanand Central American troglobitic and troglophilicplanarian fauna.Suborder MaricolaFamily DimarcusidaeOne of the more exciting events in recent years inMexican cave biology was the description of the firstmarine relict troglobitic flatworms. Unfortunately,the first species was discovered and described inde-Table 5.-Summary of cave inhabiting Platyhelminthes.TroglobitesOther SpeciesTurbellariaTricladidaMaricolaDimarcusidae 2 0PaludicolaDugesiidae 3 1TerricolaBipaliidae 0 1Geoplanidae 0 1Rhynchodemidae 0 2TrematodaDigeneaAcanthostomidae 0 1Allocreadidae 0 IAnenterotrematidae 0 2Clinostomidae 0 1Hemiuridae 0 1Lecithodendriidae 0 4Plagiorchiidae 0 1Strigeidae 0 1Urotrematidae 0 1CestodaCyclophyllideaAnoplocephalidae 0 1PseudophyllideaBothriocephalidae 0 1Total 5 2070

pendently by an Italian and by American and Japanesespecialists. Benazzi (1972) published the descriptionof Opisthobursa mexicana in a very briefand uncertain fashion, while a complete descriptionby Mitchell and Kawakatsu (1972) of Dimarcus villalobosiappeared very shortly afterwards. Mitchell andKawakatsu also erected the family Dimarcusidae intheir paper to include this remarkable species. Benazziand Giannini (1974) described the familyOpisthobursidae for the same species. Although thenomenclatorial status of the species is still in doubt,it appears that Benazzi's name has priority. Thefamily Dimarcusidae clearly has priority and is thusthe correct name for the family, with the familyOpisthobursidae falling as a junior synonym to it.Opisthobursa josephinae BenazziOpisthobursa josephinae Benazzi, 1976:533-536, pI. I(fig. 1-2), pI. 2(fig. 3-4); Ball, 1977a:151, 153;Ball, 1977b:27; Pasquini, 1977:3.Opisthobursa (part): Ball, 1977a:154; Ball, 1977b:27,29, fig. 1,3.Type-Iocality.-Pozza Casa Bell, S. Cristobal de lasCasas, Chiapas, Mexico.Distribution.-Known only from the type-locality.See Fig. 3.MEXICOGUATEMALA--~,/Fig. 3.-Distribution of troglobitic and troglophilic planarians: 1, Dugesia barbarae and D. guatemaleruis; 2, D. typh/o.mexicana; 3, Opisthobursa mexicana; 4, O. josephinae; 5, D. mckenzie; 6, Dugesia spp.71

Discussion.-This large, troglobitic flatworm issimilar to O. mexicana in many respects but is morethan twice as long. Since the dimarcusid planariansbelong to the Maricola, a primarily marine group, thediscovery of this relict species at 2,115 meters in elevationis very interesting. Marine relict species at altitudesin excess of 1,000 meters are rare, but in Mexicothey include Mexicerberus troglodytes Schultzand Speocirolana pelaezi Bolivar in the caves of theSierra de Guatemala, Tamaulipas.Opisthobursa mexicana BenazziTurbelarido cavemicola: Villalobos, 1960:332.Opisthobursa mexicana Benazzi, 1972:405; Mitchelland Kawakatsu, 1972:16; Benazzi, 1973:133-134;Benazzi and Giannini, 1974:47-54, fig. 3, pI. 1-4;Sbordoni, 1974:366; Sbordoni et al., 1974:14, 15;Benazzi, 1976:533, 535; Ball, 1977a:150, 153,154; Ball, 1977b:27, 29, 32.Dimarcus villalobosi Mitchell and Kawakatsu, 1972:2-15, fig. 1-13; Benazzi, 1973:133, 134; Mitchelland Kawakatsu, 1973a:640, 642; Mitchell and Kawakatsu,1973b:168, 169; Reddell, 1973a:32, 37;Benazzi and Giannini, 1974:54; Kawakatsu, 1975:7, fig. 2; Benazzi, 1976:533; Kawakatsu, 1976:36.Opisthobursa: Ball, 1976:649.Opisthobursa (part): Ball, 1977a:154; Ball, 1977b:27,29, fig. 1,3.Type-locality.-Of Opisthobursa mexicana: Grutasde Cocona, Tabasco, Mexico; of Dimarcus villalobosi:Las Grutas de Cocona, Teapa, Tabasco, Mexico.Distribution.-Known only from the type-locality.See Fig. 3.Discussion.-This species is believed by Mitchelland Kawakatsu (1972) to be a true marine relict. Itsrelationship to other species of Maricola is obscureand so distant as to have necessitated the erection ofa new family to include it. Ball (1977a; 1977b) hasdiscussed the status and relationships of the familyand considers it to be a very primitive group. Thespecies has been found in small travertine pools.Suborder PaludicolaFamily DugesiidaeThe family Dugesiidae is represented in the cavesof Mexico by three described troglobites and onetroglophile (see Fig. 3). In addition, troglophilic andtroglobitic Dugesia have been found in caves in SanLuis Potosi, Tabasco, Tamaulipas, Veracruz, and Yucatan.This material, together with many related epigeanspecimens, is now under study by M. Kawakatsuand R. W. Mitchell. It is interesting that the Mexicanpaludicolan flatworms belong to the Dugesiidae ratherthan to the Planariidae or Kenkiidae. The lattertwo families predominate in the cave fauna of theUnited States and include numerous troglobites. NoDugesia is known as a troglobite in the United Statesalthough a few populations are troglophiles in Texasand probably elsewhere.Dugesia barbarae Mitchell and KawakatsuDugesia sp. II: Reddell and Mitchell, 1971b: 182;Reddell and Elliott, 1973b:182.Dugesia barbarae Mitchell and Kawakatsu, 1973a:641, 646-649, 662, 663, 661-671, 673, 675, fig.4-5, 10, 15, 28-30; Mitchell and Kawakatsu,1973b:168-169; Reddell and Elliott, 1973b:181,182; Kenk, 1974:22,28; Kenk, 1975:113; Kawakatsu,1976:36.Dugesia (part): Reddell, 1973a:32.Type-locality.-La Cueva de la Capilla, Municipiode Jaumave, Tamaulipas, Mexico.Distribution.-Known only from the type-locality.See Fig. 3.Discussion.-This species is closely related to D.typhlomexicana. It is known only from small drippools in the terminal room of this cave. A troglophilicflatworm, probably D. guatemalensis Mitchell andKawakatsu, inhabits larger silt-floored pools in thesame room. The evolution of D. barbarae has beendiscussed by Mitchell and Kawakatsu (1973a). Theypostulate that the ancestor of this species and D.typhlomexicana invaded caves in the Sierra de Guatemalaearly in the Pleistocene. With the colder climateof the glacials, the surface ancestor was eliminatedfrom higher elevations and the cave species evolved.The presence in these caves of the troglophilic D. guatemalensisis explained by postulating a re-invasion ofthe area by Dugesia from the north.Dugesia mckenziei Mitchell and KawakatsuDugesia mckenziei Mitchell and Kawakatsu, 1973b:165-170, fig. 1-10; Kenk, 1974:26; Kenk, 1975:113; Kawakatsu; 1976:36.Dugesia mackenziei: Kawakatsu, 1977:18 (erroneousspelling).Type-Iocality.-La Cueva de Los Llanos, 15 kmESE San Cristobal de las Casas, Chiapas, Mexico.Distribution.-Known only from two caves at LosLlanos, Chiapas. See Fig. 3.Records.-Chiapas: Cueva de Los Llanos andCueva de los MurcieIagos.72

Discussion.-This is a minute-eyed, scarcely pigmentedspecies apparently most closely related to D.guatemalensis, known only from the Sierra de Guatemala,Tamaulipas. It is presumably a recent troglobite.It inhabits a slow moving stream which probablyruns through both caves from which the species isknown.Dugesia typhlomexicana Mitchell and KawakatsuDugesia sp. I: Reddell and Mitchell, 1971b:182, fig.1-2; Reddell and Elliott, 1973b:182.Dugesia typhlomexicana Mitchell and Kawakatsu1973a:641, 642-646, 647, 649, 662, 663, 666671, 673, fig. 1-3, 9, 13-14, 22, 24-27; Mitchelland Kawakatsu, 1973b:168, 169; Reddell and Elliott,1973b:181, 182; Kenk, 1974:22, 28; Kawakatsu,1975:7, fig. 2; Kenk, 1975:113; Kawakatsu,1976:36; Kawakatsu, 1977:17, fig. 3A.Dugesia (part): Reddell, 1973a:32.Dugesia, troglobia: Fernandez Ruiz, 1976:717.Type-locality.-Cueva de la Mina, Municipio deGomez Farias, Tamaulipas, Mexico.Distribution.-Known only from the type-locality.See Fig. 3.Discussion.-This species is closely related to D.barbarae. It has been found only in a small drip poolmid-way down a flowstone-breakdown slope. Thispool was also inhabited by the troglobitic trichoniscidisopod Brackenridgia bridgesi (VanName).Suborder TerricolaFamily BipaliidaeThe only record of this family in Mexico is of asingle specimen of Bipalium, probably referable tothe widespread species B. kewense Moseley. It wasfound in Sotano de Huitzmolotitla, San Luis Potosi.Family GeoplanidaeHyman (1938) described Geoplana multipunctataFuhrmann from Actun Loltlin, Yucatan. A specimenpossibly referable to the genus Geoplana has recentlybeen collected in Actun Sabaca, Yucatan.Family RhynchodemidaeDiporodemus yucatani was described by Hyman(1938) from Actlin Sabaca, Yucatan. Specimensprobably belonging to this species have recently beenfound among leaf litter in the entrance sink of ActlinSabaca. Specimens possibly belonging to the genusRhynchodemus have been collected from Cueva delLencho Virgen, Oaxaca, and So13no del Pozo, SanLuis Potosi.Table 6.-Summary of cave inhabiting Rotifera.TroglobitesOther SpeciesBdelloideaBdelloidaPhilodinidae 0MonogonataFlosculariaceaFlosculariidae 0 1PloimaBrachionidae 0 2Lecanidae 0 2Total 0 6Phylum RotiferaThe only records of rotifers in Mexican caves arethe six species included in the study by Osorio Tafal!(1943) of two caves in the Sierra de EI Abra, San LuisPotosi (see Table 6). Their ecological status is unknown.Class BdelloideaOrder BdelloidaFamily PhilodinidaePhilodina roseola Ehrenberg was collected InCueva de Los Sabinos, San Luis Potosi.Class MonogonontaOrder PloimaFamily BrachionidaeTwo species of this family, Lepadella patella (MUller)and Platyias patulus (Muller), were found inCueva Chica, San Luis Potosi.Family LecanidaeMonostyla closterocerca Schmarda was found inCueva de Los Sabinos, San Luis Potosi. Monostylaquadridentata Ehrenberg was collected in both CuevaChica and Cueva de Los Sabinas, San Luis Potosi.Order FlosculariaceaFamily FlosculariidaeThe only species reported for this family in Mexicancaves is Sinantherina socialis (Linnaeus); it wasfound both in Cueva Chica and Cueva de Los Sabinos,San Luis Potosi.Phylum NematodaAlthough numerous species of nematodes havebeen reported as parasites from the cave inhabitingbats and fishes of Mexico, these do not constitute in73

any way a part of the true cave fauna and are notfurther discussed here (see Table 7). Zullini (1974)has, however, produced an excellent paper on theNematoda of the soil and water of Chiapas and adjacentGuatemala. His report includes records of sevenaquatic and seven soil species of nematodes fromcaves in Chiapas and Guatemala. A second study ofthe aquatic nematode fauna of southern Mexico includednumerous records of species from wells, cenotes,and caves (Zullini, 1977).Class AdenophoreaOrder AraeolaimidaFamily PlectidaePlectus cirratus Bastian was found in the mudfrom the floor of pools in Cueva del Panteon andGrutas del Rancho Nuevo, Chiapas. This widespreadspecies was also found in stream sediment near Huistin,Chiapas (Zullini, 1974).Order MonhysteridaFamily MonhysteridaeMonhystera paludicow de Man was collected fromthe bottom of the stream in Cueva Chorreadero, Chiapas.It was also found in surface streams (Zullini,1974).Order ChromadoridaFamily CyatholaimidaeProdesmodora circuwta (Micoletzky) was foundonly in the mud of Cueva Chorreadero, Chiapas (Zullini,1974).Order EnoplidaFamily AlaimidaeAlaimus primitivus de l\'lan, a common species inEurope and America, was collected from mold inCueva Chorreadero, Chiapas (Zullini, 1974). An undeterminedspecies of Amphidelus was found indamp, guano-rich mud in Grutas de Zapaluta, Chiapas(Zullini, 1977).Family AxonchiidaeAxonchium sbordonii Zullini was described fromslime in Sima del Ojito, Chiapas (Zullini, 1974); it isknown only from this cave.Family IronidaeIronus ignavus Bastian was collected from water inCueva Chorreadero, Chiapas (Zullini, 1974), and fromwater and bottom mud in a cenote at Yokdzonot,Yucatan (Zullini, 1977). Ironus longicaudatus de Manwas found in slime in Sima del Ojito, Chiapas (Zullini,1974), and in water and bottom sediment in CenoteXtojil, Yucatan (Zullirri, 1977).Family LongidoridaeXiphinema index Thorne and Allen was reportedby Zullini (1974) from mud in Cueva del Tepesquintle,Huehuetenango, Guatemala, and from SimaTable 7.-Summary of cave inhabiting Nematoda.TroglobitesOther SpeciesAdenophoreaAraeolaimidaPlectidae 0 1MonhysteridaMonhysteridae 0 1ChromadoridaCyatholaimidae 0 1EnoplidaAlaimidae 0 2Axonchiidae 0 1Ironidae 0 2Longidoridae 0 2Mononchidae 0 5Tripylidae 0 3DorylaimidaDorylaimidae 0 6SecementeaAscarididaAscaridiidae 0 1Heterocheilidae 0 2FilariideaDipetalonematidae 0 1Filariidae 0 1OxyurideaCruziidae 0 1axyuridae 0 3SpiruridaRhabdochonidae 0 1Rictulariidae 0 1Seuratidae 0 1StrongylidaTrichostrongylidae 0 6TrichurideaTrichuridae 0 2TylenchidaTylenchidae 0 1Total 0 45del Ojito, Chiapas. This is a common species in Europe,America, and Africa. Zullini (1977) reportedX. basiri Siddiqi from wells in Merida and Teya, Yucatan.Family MononchidaeMononchus longicaudatus Cobb was reported frommud in Cueva del Panteon, Chiapas (Zullini, 1974); it74

is a cosmopolitan species. Clarkus venezolanus(Loo!), previously known only from Venezuela, wascollected from slime in Sima del Ojito, Chiapas (Zullini,1974). Mononchus sp. and Mylonchulus sigmaturus(Cobb) were found in Grutas de Zapaluta, Chiapas(Zullini, 1977). Mylonchulus sigmaturellus Mulveywas taken in Cenote Xtojil, Yucatan, and Mylonchuluslacustris (Cobb) was found in a well at Tamhek,Yucatan (Zullini, 1977).Family TripylidaeTwo species of this family were reported fromcaves in Chiapas by Zullini (1974). Trischistoma monhystera(de Man), previously known only from Italy,Germany, and Sumatra, was collected in Cueva delPanteon, Chiapas. Tripyla tenuis Brzeski was found insediment in Cueva Chorreadero, Chiapas. Zullini(1977) reported Trischistoma arenicola (de Man)from Cenote Xtojil, Yucatan.Order DorylaimidaFamily DorylaimidaeChitwood (1938) described Dorylaimus yucatanensisfrom Cueva Luchil, Yucatan. This species hassince been placed in the synonymy of Eudorylaimusgranuliferus (Cobb). Zullini (1974) reported Mesodorylaimusbastiani (Biitschli) from mud in Cueva delPanteon, Chiapas. Zullini (1977) reported four additionalspecies of this family from wells and cenotes inYucatan and Tabasco. He also reported Mesodorylaimussp. and Aporcelaimellus sp. from guano-rich mudin Grutas de Zapaluta, Chiapas.Phylum EctoproctaClass PhylactolaemataOrder PhylactolaemataFamily PlumatellidaeLacourt (1968) reported two species of the bryozoangenus Plumatella from cenotes in Yucatan: P.emarginata Allman from Cenote Xtolok and P. evelinae(Marcus) from Cenote del Country Club. Bothcenotes are large and open, and these species certainlyare not a part of the true cave fauna.Phylum AnnelidaClass CliteUataOrder BranchiobdellidaFamily BranchiobdellidaeThe branchiobdellid worms are commensals ofcrayfishes and isopods. Two species are known fromMexican caves (Holt, 1973). Cambarincola acudentatusHolt is of special interest in that it has been foundonly on the troglobitic cirolanid isopods Speocirolanapelaezi (Bolivar) and S. bolivari (Rioja) in Grutas deQuintero, Tamaulipas. The second species, C. susanaeHolt, is known from the crayfish Procambarus acutuscuevachicae (Hobbs) in Cueva Chica, San Luis Potosi.Table 8.-Summary of cave inhabiting Annelida.TroglobitesOther SpeciesClitellataBranchiobdellidaBranchiobdellidae 0 2HaplotaxidaHaplotaxinaTubificidae 0 1LumbricinaAcanthodrilidae 2 3Glossoscolecidae 0 1Lumbricidae 0 3Megascolecidae 0 2Ocnerodrilidae 0 1Octochaetidae 0 5HirudineaArhynchobdellaeErpobdellidae 0 1RhynchobdellaeGlossiphoniidae 0 2Ichthyobdellidae 0 1Piscicolidae 0 1Incertae SedisAelosomatidae 0 1Total 2 24Order HaplotaxidaSuborder HaplotaxinaFamily TubificidaeThe family Tubificidae is represented in Mexicancaves only by a record of Aulophorus sp. in Cueva deLos Sabinos, San Luis Potosi (Osorio Tafall, 1943).Numerous collections of small aquatic oligochaetesremain unstudied. Specimens from caves in the Sierrade Guatemala, Tamaulipas, and from Cueva de la Siquita,Durango, were found in unusual situations(drip pools and deep spring-fed pools) and in directassociation with aquatic troglobites. The extremetransparency of these specimens and the circumstancesin which they were found strongly indicatethat they are troglobitic.Suborder LumbricinaEarthworms are not uncommon in caves but arefrequently in very poor condition, indicating that75

they have been washed in with soil and will not survive.In many caves they abound and obviously reproduce.It is noteworthy that of only 17 earthwormsendemic to Mexico, seven are known solely fromcaves. Of the epigean species all are rare and seldomcollected from caves. Gates (1971) speculates that thenative earthworm fauna has been largely eliminatedby the hardy exotic species which now are widespreadin Mexico. The abundance of endemic speciesin caves may well reflect the ability of these wormsto survive in that environment. In addition to theseven endemic earthworms, at least II exotic specieshave been collected from Mexican caves (see Table8).Family AcanthodrilidaeFive species belonging to the family Acanthodri·lidae have been found in Mexican caves. Of the fourspecies which have be"en positively determined, all areknown only from the cave habitat and two appear tobe troglobites. Undetermined specimens of a probablyendemic species of Diplocardin have been foundin caves in San Luis Potosi and Tamaulipas. Pickford(1938) described Eodrilus oxkutzcabensis and Balanteodriluspearsei from caves in Yucatan. The latterspecies has recently been collected from a moist siltyarea in Actun Loltun, Yucatan (Gates, 1977). The remainingtwo species are troglobites and discussed below.GUATEMALA,.."".0-r,."'/.//Fig. 4.-Distribution of troglobitic and troglophilic earthworms of the family Acanthodrilidae: 1, Eodrilus albidus; 2, E.mexicanu8; 3, E. oxkut%cabensi8; 4, Balanteodrilwpear8ei; 5, E. oxkutzcabensi8 and B. pearsei.76

Eodrilus albidus GatesEodrilus albus Gates, 1970:267-269 (nom. preocup.);Reddell, 1971b:13; Reddell and Mitcnell, 1971b:183, fig. 7; Mitchell and Kawakatsu, 1973a:675,676; Gates, 1973:21; Reddell, 1973a:37; Reynoldsand Cook 1976:67.Eodrilus sp.: Gates, 1971:7-8; Gates, 1973:21.Eodrilus albidus Gates, 1973:21 (nom. nov.); Reynoldsand Cook, 1976:67.Large earthworm, probably Eodrilus albus: Mitchelland Kawakatsu, 1973a:657.Eodrilus (part)· Reddell, 1973a:32.Type-Iocality.-Cueva de la Perra (=Capilla), LaPerra (=EI Porvenir), 15 miles northwest of GomezFarias, Tamaulipas, Mexico.Distribution.-Known from the type-locality andpossibly one other cave in the Sierra de Guatemala,Tamaulipas. See Fig. 4.Records.-Tamaulipas: Cueva de la Capilia and?Cueva de las Perlas.Discussion.-This species is included as a troglobitestrictly on the basis of its habitat and the transparencyof the body wall. It is only known to inhabitmud-floored drip pools in darkness in the two abovecaves. The integument is thin and pigmentation isgreatly reduced. Other members of the genus areknown only from tropical lowland areas. Specimensfrom Cueva de las Perlas were poorly preserved butprobably belong to this species.Eodrilus mexicanus GatesEodrilus, undescribed: Reddell, 1967d: 106.Eodrilus mexicanus Gates, 1968:63-70; Gates, 1970:267, 268, 269; Gates, 1971:7-8; Reddell, 1971b:183; Gates, 1972:26; Reddell, 1973a:37; Reynoldsand Cook, 1976:136.Eodrilus: Peck and Peck, 1973:67.Eodrilus (part): Reddell, 1973a:32.Type.locality.-Sotano de Tlamaya, San Luis Potosi,Mexico.Distribution.-Known only from caves in the vicinityof Xilitla, San Luis Potosi. See Fig. 4.Records.-San Luis POtOSI: Cueva de la Parra,Cueva del Salitre, and S6tano de Tlamaya.Discussion.-This species is known only from drippools in each of the above caves. It is essentially pigmentlessand, when alive, is so transparent that internalorgans are easily visible. For this reason, and becausethe genus is otherwise one of a strictly lowlandtropical distribution, this species is considered a possibletroglobite. A record of this species from Sotanode San Agustin, Oaxaca (Gates, 1972), is doubtlessan error in identification or curation.Family GlossoscolecidaeThis family is represented in Mexican caves onlyby specimens of Pontoscolex sp. d. corethrurus(Mueller) from Sotano de Huitzmolotitla, San LuisPotosi.Family LumbricidaeThree species, all exotics, are known from Mexicancaves. Dendrodrilus rubidus (Savigny) has beencollected in caves in San Luis Potosi, Tamaulipas, andVeracruz (Gates, 1973, 1977). Octolasion tyrtaeum(Savigny) is known from several caves in Queretaroand San Luis Potosi (Gates, 1971, 1973). Specimensof Aporrectodea rosea (Savigny) have been foundonly in S6tano de la loya de Salas, Tamaulipas(Gates, 1971).Family MegascolecidaeTwo species of the family l\legascolecidae havebeen collected in Mexican caves. Pheretima diffringens(Baird) is frequently found in stream caves andhas been taken from mud and beneath rotting woodin Oaxaca, Puebla, San Luis Potosi, and Veracfllz(Gates, 1971, 1977). Pheretima hawayana (Rosa)was Gollected from wet mud under rotting wood inSumidero de Cuetzeltemanes, Puebla (Gates, 1977).Family OcnerodrilidaeEukerria saltensis (Beddard) was collected fromwet guano and silt in Cueva de la Florida. Tamaulipas(Gates, 1971). this is the only species of this familyreported from Mexican caves.Family OctochaetidaeFive species of the family Octochaetidae havebeen found in Mexican caves. Pickford (1938) reportedDichogaster affinis (Michaelsen) and D. bolaui(Michaelsen) from caves in Yucatan. Undeterminedspecimens of Dichogaster have been found incaves in San Luis Potosi and Tamaulipas. The genusTrigaster is represented by three cave species (presumablytroglophiles) in San Luis Potosi and NuevoLeon. Trigaster reddelli and T. vallesensis were describedby Gates (1971) from Sotano de Yerbaniz,San Luis Potosi; T. albidus Gates is known onlyfrom So13no del Anticlino, uevo Leon (Gates,1973). These three species are extremely abundantand obviously thriving in deep mud in these twocaves.Class HirudineaLeeches are only rarely collected in caves, but inat least several localities they constitute a significant77

part of the aquatic fauna. A very large population ofan undetermined species is present in Grutas de SanJose, Campeche. In addition to the free-living speciesothers are known to be parasites of snails and fishes.Order ArhynchobdellaeFamily ErpobdeUidaeThe predaceous leech Erpobdella triannulataMoore was reported from Cenote Xtolok, Yucatan,by Moore (1936).Order RhynchobdellaeFamily GlossiphoniidaeGlossiphonia magnidiscus was described by Moore(1938) from Cenote de Sambula, Motul, Yucatan. Itwas a parasite of the pimelodid catfish Rhamdia guatemalensisdecolor Hubbs.Family IchthyobdeUidaeCystobranchus sp. was reported by Moore (1936)to be a parasite of R. guatemalensis decolor in Cenotede Sambula. It is also known from open-air cenotes innorthern Yucatan.Family PiscicolidaePiscicola platense Cordero is a parasite of R. guatemalensisdecolor in Cenote de Sambula (Motul) andCenote de Sambula (Merida), Yucatan (Moore, 1938).Phylum MolluscaClass GastropodaThe snail fauna of Mexican caves is poorly knowndespite the fact that 82 species have been identified(see Table 9). Many of these are known only frompaleontological and archeological excavations, anddoubtless were washed or carried into the caves.Other species are known only from open-air cenotesin Yucatan (Bequaert and Clench, 1933, 1936, 1938)and are not a part of the true cave fauna. The onlypossible troglobite is an aquatic species of the familyHydrobiidae. The absence of terrestrial troglohiticsnails from caves in Mexico is not unexpected. Noterrestrial snails are troglobites in caves in Texas orother states of the southwestern United States. Terrestrialsnails are also rare in the troglobitic fauna oftropical caves in general, although species of the genusOpeas with reduced eyes have been describedfrom caves in Assam and Malaya.Order ArchaeogastropodaFamily HelicinidaeThis is the only family of archaeogastropod snailrepresented in the Mexican cave fauna. Ceres nelsoniDall and Helicina chrysocheila Binney are knownfrom caves in the Xilitla and Aquismon regions, SanLuis Potosi. The former specip.s is a conspicuous elementof the entrance area fauna and is presumably atrogloxene. Oligyra arenicola (Morelet) has been collectedfrom several caves in Yucatan. The other speciesof this family are known only from single caveseach and are of uncertain status.Order MesogastropodaFamily CyclophoridaeAlthough several species of this family have beenfound in caves, most were dead. The only importantspecies is the troglophile Neocyclotus dysoni berendti(Pfeiffer), which is known from numerous cenotesand caves in Yucatan (Bequaert and Clench, 1936,1938).Family HydrobiidaeThis family of aquatic snails includes two speciesassociated with the subterranean habitat. Bequaertand Clench (1936, 1938) reported the presence ofPyrgophorus coronatus (Pfeiffer) in several caves inTable 9.-Summary of cave inhabiting Gastropoda.ArchaeogastropodaHelicinidaeMesogastro podaChondropomidaeCyclophoridaeHydrobiidaePilidaePomatiidaePotamididaeThiaridaeBasommatophoraAncylidaePhysidaePlanorbidaeStylommatophoraAchatinidaeCharopidaeEndodontidaeFerussaciidaeFruticicolidaeHelminthoglyptidaeOrthalicidaePolygyridaePupillidaeSpiraxidaeUrocoptidaeXanthonychidaeZonitidaeTotalTroglobitesooo1oooooooooooooooooooo1Other Species81311313145912113712126148178

Yucatan. Specimens of Pyrgophorus are known fromGrutas de San Antonio, Campeche. The probabletroglobite Coahuilix hubbsi Taylor is discussed below.Coahuilix hubbsi TaylorCoahuilix hubbsi Taylor, 1966: 156, 163, 180-181,fig. 8-13; Minckley, 1969:39; Reddell, 1971b:77.Coahuilix: Taylor, 1966:175, 180.Type-Iocality.-Northernmost pool of Pozos de laBecerra, 14 km southwest of Cuatro Cienegas, Coahuila,Mexico.Distribution.~Known only from the type-locality.Discussion.-This species is known only by shells,but it is included here because it apparently waswashed from the subterranean habitat by the springsfeeding Pozos de la Becerra. This interesting snail is amember of the tribe Horatiini, a group known fromthe vicinity of the Mediterranean and from subterraneanwaters in Texas and Alabama. Horatia (Hauffenia)micra (Pilsbry and Ferris) has been reported fromshells in the Guadalupe River, Texas, where it hasprobably been washed from one of the numerouscaves or springs along the river. Undescribed troglobiticspecies of Horatia (Hauffenia) are known fromthe artesian well at San Marcos, Hays County, andfrom Salamander Cave, Travis County, Texas. A recordof H. (H.) micra from a cave in Alabama isprobably of an undescribed form. The distributionof the Horatiini corresponds to the general patternexemplified also by the isopod families Cirolanidaeand Stenasellidae.Family PomatiidaeBequaert and Clench (1936, 1938) reported Choanopomalargillierti (Pfeiffer) from caves and cenotesin Yucatan. This species has recently been collectedfrom several additional Yucatan caves.Family ThiaridaeA large population of Melanoides maculata (Born)was found in Cueva de Cantil Blanco, Veracruz. Thisis a species introduced from southeast Asia.Order BasommatophoraFamily PhysidaeThree genera of physid snails are closely associatedwith the cave habitat in Mexico. Aplexa spiculata abbreviata(Fischer and Crosse) was found in Grutas deBalankanche, Yucatan (Bequaert and Clench, 1938).Species of the genus Physa are known from severalcaves in Coahuila, San Luis Potosi, and Tamaulipas,but remain unstudied. Stenophysa sp. has proven tobe a significant troglophile in several caves in Campeche.Family PlanorbidaeAlthough several planorbid snails have been reportedfrom archeological sites in Yucatan, noneappears to be inhabiting the caves. Helisoma trivolvis(Say), however, is a possible troglophile in Sotano dela Joya de Salas, Tamaulipas.Order StylommatophoraThis is doubtless the most important group ofsnails inhabiting caves in Mexico. The order is representedby about 50 species in 13 families. Only afew of the more significant species are discussedhere.Family AchatinidaeThis is among the better represented families to befound in caves, with nine species in five genera havingbeen determined to date. The genus Lamellaxis is oneof the more frequently encountered groups, withthree determined species, and many specimens fromseveral areas in Mexico await study. Lamellaxis gracilis(Hutton) is an abundant troglophile in the caves ofOaxaca, San Luis Potosi, and Yucatan. Lamellaxismartensi (Pfeiffer) was reported from Grutas de Balankanche,Yucatan, by Bequaert and Clench (1936).Lamellaxis micra (d 'Orbigny) is present in numerouscaves in Yucatan. Other species of achatinid snailknown from Mexican caves include: Leptinaria mexicana(Pfeiffer) from San Luis Potosi and Puebla;Opeas pyrgula (Schmacker and Boettger) from Grutasde Cacahuamilpa, Guerrero; O. yucatanense Pilsbryand Subulina octona (Brugiere) from Yucatan;and S. porrecta von Martens from Grutas de JuxtlalUilaca,Guerrero. All of these species are apparenttroglophiles.Family CharopidaeThe only record for this family in Mexican caves isof an undescribed genus and species found in Cuevadel Rio Texocotla, Puebla. It is presumably a troglophile.Family EndodontidaeHelicodiscus singleyanus (Pilsbry) is commonlypresent on cave walls and floors in large numbers. Ithas been found in caves in Oaxaca and Tamaulipas.79

The genus Helicodiscus is a significant element of theTexas cave fauna.Family OrthalicidaeAlthough seven species of the family Orthalicidaehave been reported from caves, the only one which isknown with certainty to live in caves is Bulimulusunicolor (Sowerby). This species was reported fromseveral caves in Yucatan (Bequaert and Clench, 1936,1938); it has been collected from several additionalcaves recently.Family SpiraxidaeThis is the most important family of snails withrespect to the cave ecosystem, with species of Euglandina,Spiraxis, and Streptostyla having been found astroglophiles. Euglandina cylindracea (Phillips), Streptostylameridana meridana (MoreIet), and Streptostylaventricosula (Morelet) are frequently collectedin the caves of the Yucatan Peninsula. Streptostylajilitlana Dall is an important species in the high altitudecaves of Queretaro and San Luis PotosI. Eggs,newly hatched young, and adults of Streptostylabartschi (Dall) have been found in the inner rooms ofCueva de la Mina, Tamaulipas (Reddell and Mitchell,1971b). This species is also known from other cavesin San Luis PotosI and Tamaulipas.Family UrocoptidaeBrachypodella dubia Pilsbry and Microceramusconcisus (Morelet) are frequently encountered speciesin the caves of the Yucatan Peninsula. Thompson(1967) reported Brachypodella speluncae (Pfeiffer)from two caves in El Peten, Guatemala. A new speciesof the genus Coelocentrum is known from caves inTamaulipas.Family XanthonychidaeThe only species of this family known from Mexicancaves is an undescribed species of Xanthonyxfrom Cueva de la Capilla, Tamaulipas. It was foundon the wall of the cave near a small upper entrance.Family ZonitidaeSeveral species of zonitid snail are known fromMexican caves. The only one which is common isHawaiia minuscula (Binney), known from caves inCampeche, Veracruz, and Yucatan.Class PelecypodaFew clams have ever been recorded from caves andthen usually from the vicinity of spring entrances. Asmall undetermined species is known from Cueva delNacimiento del Rio San Antonio, Oaxaca. This speciesinhabits the stream passage in large numbers andis pigmentless with a· thin shell allowing the internalorgans to be observed. The determination of itsecological status must await further study.Phylum ArthropodaClass CrustaceaThe class Crustacea is second only to the Arachnidain the number and diversity of troglobites presentin Mexican and Guatemalan caves. Of the eightorders of subterranean crustaceans represented, sixcontain troglobites (see Table 10). Two additionalorders, the Thermosbaenacea and the Bathynellacea,are present in the caves of Texas and other parts ofthe world but are not yet known from Mexico orCentral America. This lack is probably a reflectionon collecting techniques rather than an indication ofthe absence of these groups. In general the planktonTable 10.-Summary of cave inhabiting Crustacea.TroglobitesOther SpeciesBranchiuraArguloidaArgulidae 0 2CopepodaCalanoidaDiaptomidae 1 3CyclopoidaCyclopidae 0 10HarpacticoidaAmeiridae 0 5Canthocamptidae 0 2OstracodaPodocopaCypridae 0 12Darwinulidae 0 1Entocytheridae 2 10BranchipodaCladoceraChydoridae 0 1Daphnidae 0 7Malacostraca*Isopoda 40 16AmphipodaBogidiellidae 9 0Hadziidae 5 0Hyalellidae 0 1Melitidae 0 1MysidaceaLepidomysidae 2 0Mysidae 2 0*Decapoda 17 18Total 78 89*For summary by families see Tables 11-12.80

of Mexican caves is unknown, but recent collectionsin southern Mexico by Dr. Valerio Sbordoni and hiscolleagues have included emphasis on planktonicforms.Subclass BranchiuraOrder ArguloidaFamily ArgulidaeTwo species of argulid have been taken in associationwith fish in the cenotes of Yucatan (Wilson,1(36). Argulus chromidis Kroyer was found in theintestine of Rhamdia guatemalensis depressa Barbourand Cole in Cenote Chica de XanaLa; A. rhamdiaeWilson was found on the skin of R. guatemalensisdepressa in Cenote Scan Yui.Subclass CopepodaCopepods are an important element III the cavefauna of Mexico but remain poorly known. Only onespecies is definitely troglobitic, but others may proveto be so once they are more carefully studied. A totalof 20 species of non-troglobitic copepod arl' nowknown from the underground waters of lVlexico (seeTable 10).Order CalanoidaFamily DiaptomidaeTwo diaptomids have been identified from caves inMexico, one of which is a trogloLite and is discussedbelow. Diaptomus texensis M. S. Wilson has been collectedfrom a small drip pool in Actun Loltun, Yucatan.Diaptomus (Microdiaptomus) cokeri Osorio TafallDiaptomus nueva especie y subgenera: Anonymous,1942a:221.Diaptomus (Microdiaptomus) cokeri Osorio Tafall,1942:206, 207-210, fig. 1-17; Osorio Tafall, 1943:49, 50, 51, 52,53,68,69; Rioja, 1953a:287, 288,292; Wilson, 1959:780; Nicholas, 1962:168; Vandel,1964:131; Vandel, 1965a:l07; McKenzie,1965a:38; Reddell, 1971b: 17; Reddell and I\Iitchell,1971a:141; Reddell and 8lliott, 1973a:I71.Microdiaptomus: Osorio Tafall, 1946: 153.Diaptomus cokeri: Bolivar, 1950:216; Tamayo,1962:vol. III:260.Diaptomus (Microdiaptomus) cokersi: Barrera, 1968:311 (erroneous spelling).Microdiaptomus cokeri: Thines and Tercafs, 1972:66.Type-Iocality.-Cueva Chica, Ciudad Valles, SanLuis Potosi, Mexico.Distribution.-Known only from two caves nearValles, San Luis Potosi.Records.-San Luis PotOSI: Cueva Chica and Cuevade Los Sabinos.Discussion.-This species of fresh-water origin has,to my knowledge, not been collected since it was firstreported. The subgenus Microdiaptomus includesonly this species.Order CyclopoidaFamily Cyclopidael\ine species of cyclopid copepods have been identifiedfrom Mexican caves. Osorio Tafall (1943) rcportedthe presence of Cyclops (AcanthocycLops) vernalisrobustus Sars and MesocycLops (ThermocycLops)inversus (Kiefer) from caves near Valles, San LuisPotosi. EucycLops serruLatus (Fischer) and Tropocyclopsprasinus (Fischer) are known from caves inSan Luis Potosi (Osorio Tafall, 1(43) and ct~nolt'sand caves in Yucatan (Wilson, 1936; Pearse and Wilson,1(38). MacrocycLops albidus (Jurine) is a frequentinhabitant of caves, and in Mexico it is knownfrom caves in Yucatan (Wilson, 1936; Pearse and Wilson,1938), San Luis Potosi (Osorio Tafall, 1943),Campeche, and Guerrero. A probable new sp(~cies ofMesocycLops related to M. brazilianus Kiefer has beencollected in Grutas del 10gote, Guerrero. Yeatman(1977) redescribed .f\Ilesocyciops ellipticus Kieferfrom Grutas de XtaclImhilxunam, Campeche. Thisspecies, also known from caves in Oaxaca and Yucatan,is probably that reported as iH. leuckarti (Claus)from caves and cenotes in Yucatan (Wilson, 1936;Pearse and Wilson, 1938). Mesocyclops (Thermocyclops)tenuis (Marsh) is known from caves andcenotes in Yucatan (Wilson, 1936; Pearse and Wilson,1938) and from caves in Campechc. Paracyclops Jimbriatus(Fischer) is one of the mort~ frequently encounteredcopepods in North American caves; inMexico it has been found in caves in the states ofSan Luis Potosi (Osorio TafaH, 1(43), Yucatan, andCampeche.Order HarpacticoidaFamily AmeiridaeNitocra sp. was reported by Osorio TafaU (1943)from Cueva Chica and Cueva de Los Sahinos, SanLuis Potosi. Wilson (1936) identified four species ofNitocra from caves and cenotes in YlICatan, but onlyN. pusilla Sars from Cenote de Samhula (Motul) is81

associated with true cave waters. Pearse and Wilson(1938) reported Nitocrella subterranea (Chappuis)from Grutas de Balankanche, Yucatan, but this UnitedStates species may bc misidentified.Family CanthocamptidaeTwo genera of canthocamptid copepod have beenrecorded from \Iexican caves. Canthocamptus sp. wasidentified from Cueva de Los Sabinos, San Luis Potosi,by Osorio Tafall (1943). Attheyella pilosaChappuis was reported by Wilson (1936) from CuevaLuchil, Yucatilll, and is listed by Nicholas (1962) asa troglobite. This species is a commensal of crayfishesin Kentucky and Indiana and furthermore occurs oncrayfish in an epigean locality in Kentucky (Bowmanet aI., 1968). Nicholas (1962) also lists Moraria cristataChappuis as a troglobite from "unnamed cave atCienga, near Progreso, Yucatan." This species wastaken from a cienega (a large open pond) and not acave.Subclass OstracodaOrder PodocopaThe ostracods of J\lexican caves are very poorlyknown. A total of 25 species have been identified,but eight are known only from open-air cenotes inYucatan and 12 are commensals on crayfish and isopods(see Table 10). The remaining five species surelyconstitute a very small percentage of the free-livingostracods inhabiting caves in ;\'Iexico.Family CypridaeOsorio Tafall (1943) reported Candona sp. andCypris sp. from caves near Valles, San Luis Potosi.Furtos (1936, 1938) described Cypridopsis inaudita,C. mexieana, and C. yucatanensis from caves in Yucatan.Chlamydotheea areuata (Sars) has recently beencollected from Cueva de EI Pachon, Tamaulipas.None of these show any signs of adaptation for a caveexistence.Family DarwinulidaeThe only species of this family reported from subterraneanwaters is Darwinula stevensoni (Brady andRobertson); it was found in Cenote Xlaka, Yucatan(Furtos, 1936).Family EntocytheridaeTwelve species of entocytherid ostracod, two ofwhich are troglobites and commensals of cirolanidisopods, have been collected from Mexican caves. Theremaining species are all associated with crayfish. Hartand Hart (1974) in a review of the family have includedall synonymies and records. Ankylocytherebidentata (Rioja) is known from caves in Chiapas,Oaxaca, and Veracruz; A. maya Hobbs from caves inOaxaca; A. sinuosa (Rioja) from Cueva Chica, SanLuis Potosi; A. tolteeae Hobbs from caves in Hidalgoand San Luis Potosi; A. villalobosi Hobbs from Grutasde Zapaluta, Chiapas; and Entocythere claytonhoffiRioja from caves in Chiapas, Oaxaca, and SanLuis Potosi. Entocythere mexicana Rioja, Uncinoeytherebieuspida (Rioja), U. euadrieuspida (Rioja),and U. dobbinae (Rioja) have all recently been foundin caves in the Cuetzalan region of Puehla.Sphaeromieola cirolanae RiojaSphaeromieola cirolanae Rioja, 1951a: 170-179, fig.1-16; Rioja, 1953a:289-290, 291, 292; Levinson,1959:259; Nicholas, 1962:168; Hart, 1962:122,139; Vandel, 196-1-:303; Vandel, 1965a:254; Reddell,1967b:82; Reddell, 1971b:lB: Hobbs, 1971:2,3,7,8-9,45-47, fig. 30-31; Reddell and l\litchell,1971a: l-I-2; Hobbs and Hobbs, 1973:39-41,fig. la, b; Reddell and Elliott, 11)73a:17]-172;Reddell and Elliott, 1973b:181, 18:3; 1I01t, 11)73:4; Hart and Hart, 1974:15,164-165, pI. XLIU(fig.8-14), LX]; Danielpol, 1977:22,34-36,39, fig. 10;Hart, 1978:724,725,727,728,729, fig. 1,8-9.Sphaeromicola sp.: Reddell, 1967b:82.Type-locality.-Cueva de los Sabinos, San LuisPotosi, Mexico.Distribution.-Known from caves in Nuevo Leon,San Luis Potosi, and Tamalllipas. See Fig. 5.Records.-Nuevo Leon: Cueva La Chorrera; SanLuis Potosi: Sotano del Arroyo, Cueva de la Curva,Cueva de La Lagunita, Sotano de I\latapalma, Sotanitode Montecillos, Sotano de Pichijumo, Sotano delas Piedras, Cueva de Los Sabinos, Sotano del Tigre,and Sotano de Yerbaniz; Tamaulipas: Bee Cave,Cueva de la Florida, Sotano del Molino, Cueva deEI Pachon, Grutas de Quintero, and Sotano de EIVenadito.New records.-San Luis Potosi: Cueva de las Lagunitas;Tamaulipas: Cueva del Agua de Simon Salinas(det. H. H. Hobbs, Jr.).Discussion.-Sphaeromicola cirolanae is a commensalof isopods of the family Cirolanidae. It hasbeen taken from Speocirolana bolivari (Rioja) inGrutas de Quintero, Tamaulipas; from Speocirolanan. sp. in Cueva La Chorrera, Nuevo Leon; andfrom Speocirolana pelaezi (Bolivar) in all of theabove caves except Cueva La Chorrera. There arenine described species of Sphaeromicola, only twoof which occur in Mexico. Six species are found in82

the Mediterranean region: three are troglobites, oneis marine, and two are epigean fresh-water species.Sphaeromicola moria Hart is a troglobite knownfrom Cirolanides texensis Ulrich in Rambie's Cave,Uvalde County, Texas (Hart, 1978).Sphaeromicola coahuiltecae Hobbs and HobbsSphaeromicola coahuiltecae Hobbs and Hobbs, 1973:41, fig. lc-g; Hart and Hart, 1974: 15, 165, pI.XLIlI(fig. 15-18), LXI; Hart, 1978:724, 725, 728,729, fig. I.Type-locality.-Cueva del Huisache, 4 km NW Micos,San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 5.Discussion.-The host of this species is not knownwith certainty. It was taken from sediment in a jarcontaining four species of aquatic isopod: MexistenasellusparzeJalii Magniez, M. wilkensi Magniez,Mexilana saluposi Bowman, and an undescribed speciesof Cyathura. Since all other Mexican records forSphaeromicola are from cirolanids, it is probablethat Mexilana saluposi is the host for this species.Subclass BranchipodaOrder CladoceraFamily ChydoridaeThe only species of this family known from Mexicancaves is Alona sp.; it was found in Cueva de LosSabinos, San Luis Potosi (Osorio Tafall, 1943).1 ,.,.~( "I ,MEXICOGUATEMALA/.//r\,.... r'-"Fig. 5.-Distribution of troglobitic ostracods of the family Entocytheridae: 1, Sphaeromicola cirolanae; 2, S. coahuiltecae.83

Family DaphnidaeWilson (1936) reported seven species of cladoceransof the genera Ceriodaphnia, Daphnia, Moina, andSimocephalus from cenotes in Yucatan. One of these,C. cornuta Sars, has recently been collected in CenoteBolchen, Campeche. Ceriodaphnia lacustris Birge wasreported from Cueva de Los Sabinos, San Luis Potosi(Osorio Tafall, 194.3).Subclass MalacostracaOrder IsopodaThe order Isopoda is the most abundant group tobe found in the waters of Mexican caves and is amongthe more important inhabiting the terrestrial environment.It is also one of the few groups in which thetroglobites outnumber the troglophiles. To date, atotal of 48 species of isopod has been identified fromthe caves of Mexico, Guatemala, and Belize (see Table11). The 32 troglobites are distributed through allfive suborders represented; three of these suborders(Flabellifera, Anthuridea, and Microcerberidea) areknown from Mexico only by troglobites.Table ll.-Summary of cave inhabiting Isopoda.TroglobitesOther SpeciesFlabelliferaCirolanidae 10 0AsellotaAsellidae 5 1Stenasellidae 6 0AnthurideaAnthuridae 2 0Microcerberid eaMicrocerberidae 2 0OniscoideaArmadillidae 0 9Oniscidae 0 1Philosciidae 2 0Porcellionidae 0 3Sphaeroniscidae 1 0Squamiferidae 3 1Trichoniscidae 9 1Total 40 16Suborder FlabelliferaFamily CirolanidaeThe family Cirolanidae is primarily marine, butcavernicolous species are now known from the Mediterraneanregion, East Africa, the greater Caribbeanregion, and Texas and Virginia (U.S.A.). In Mexico itranges from near the United States border into theYucatan Peninsula.Conilera stygia PackardEyeless asellid crustacean: Packard, 1894:732.Conilera stygia Packard, 1900:300-301; Packard,1901:228; Richardson, 1905:116, 120; Banta,1907:78; Chappuis, 1927:72; Van Name, 1936:31, 426-427; Jeannel, 1943:255-256; Bolivar,1950:211, 217, fig. 11; Rioja, 1953a:294; Rioja,1953d:147, 168; Rioja, 1957:447; l\icholas,1962:170; Rioja, 1962:40: Bowman, 1964:233,234, 236, fig. 61; Vandel, 1964:150; Vandel,1965a:120; Cole and Minckley, 1966:17, 21;Reddell, 1967a:24; Straskraba, 1969:22; Reddell,1971b:20; Bowman, 1975: 1."Conilera" stygia: Cole and Minckley, 1972:321.Conilera: Chappuis, 1927: 142.Conilera (part): Bowman, 1975:5,6, fig. 4b.Conylera stygia: Rioja, 1953a:286 (erroneous spelling).Comilera stygia: Rioja, 1953a:293 (erroneous spelling).Type-Iocality.-Well at Monterey (=Monterrey),Nuevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 6.Discussion.-This species was too poorly describedto be accurately identified. The genus Conilera is exclusivelymarine; when rediscovered, this species willprobably prove to belong in the genus Speocirolana.Creaseriella anops (Creaser)Cirolanid isopods: Pearse, 1933:110.Cirolana anops Creaser, 1936:117-120, 131, fig. 1-12;Pearse, 1936d:24; Van Name, 1936:523: Creaser,1938:159-160, 164; Van Name, 1940:134, fig. 26;Jeannel, 1943:255; Osorio Tafall, 1943:55; Pearse,1945:165; Anonymous, 1947:128; Bolivar, 1950:211, 217, 218, fig. 11; Cardenas Figueroa, 1950:156; Villalobos, 1951:215; Bonet, 1953b:241;Rioja, 1953a:286, 293, 294; Rioja, 1953d:147;Delamare Deboutteville, 1960:648.Cirolana: Creaser, 1938:159; Pearse, 1938a:13, 15;Pearse, 1945:111, 166; Cardenas Figueroa, 1950:157.Cirolana (part): Rioja et al., 1961:315.Cirolana ampos: Carreno, 1959:24 (erroneous spelling).Creaseriella anops: Rioja, 1953d:162, 164-166, 168,fig. 36-58; Nicholas, 1962:170; Rioja, 1962:40;Bowman, 1964:236, fig. 61; Vandel, 1964:151;Vandel, 1965a:121; Cole and Minckley, 1970:76;Reddell, 1971b:21; Reddell, 1977b:229, 239,240,251,253,255,258,260,262,263,264,266,84

267, 268, 273, 274, 277, 278, 283, 291; Ruffoand Vigna Taglianti, 1977: 167.Craeseriella: Straskraba, 1969:22; Juberthie, 1974:81(erroneous spelling).Craeseriella anops: Rioja, 1962:40 (erroneous spelling).Creaseriella: Rioja, 1953d:147, 161; Rioja, 1957:447; Bowman, 1964:234; Straskraba, 1969:4,28;Cole and Minckley, 1970:77-78; Cole and Minckley,1972:321; Bowman, 1975:5,6,7, fig. 4c.Cresseriella anops: Rioja, 1971:509 (erroneous spelling).Type-Iocality.-San Bulha Cave (=Cenote de Sambula)at Motul, Yucatan, Mexico.Distribution.-Known from many caves in QuintanaRoo and Yucatan and a well in Campeche. SeeFig. 6.Records.-Campeche: Well (Campeche); QuintanaRoo: Cenote de San Martin and Cenote de Tos Virlol;Yucatan: Cenote de las Abejas, Cueva Amil, CenoteCalchuhuim, Cueva Chac Mol, Cenote de la Culebra,ActUn Chac, ActUn Gongora, Cenote de HoctUn,ActUn Kaua, Cenote de Kankirixche, ActUn Okobichen,Cenote del Pochote, Cenote de Sambula (Mo-\ .. '\."-., .,\)\'.,MEXICOGUATEMALAr\,.. r·..-'/.//Fig. 6.-Distribution of troglobitic isopods of the family Cirolanidae: 1, Sphaerolana affinis, Sphaerolana interstitialis, andSpeocirolana thermydronis; 2, Conilera stygia; 3, Sphaerolana affinis; 4, Speocirolana n. sp. 1; 5, Speocirolana n. sp. 2; 6,Speocirolana pelaezi; 7, Speocirolana pelaezi and Speocirolana bolivari; 8, Mexllana saluposi; 9, Creaseriella anops.85

tul), Cueva de San Isidro, Cueva de Santa Elena, Pozode Santa Elena, Cueva Xconsacab, and Actun Xpukil.Discussion.-This is one of three genera of cirolanidscapable of rolling into a ball, one being Sphaerolana(discussed below) and the other Faucheriafrom France. Faucheria and Creaseriella have beencombined into the subfamily Faucheriinae, but thisis probably an artificial grouping. The ability to rollinto a ball has probably evolved more than once inthe cirolanids (Cole and Minckley, 1970). Relationshipto Sphaerola1la is quite remote. Creaseriella is amonotypic genus. Creaseriella anops is usually foundcrawling along the bottom or sides of pools. In atleast one instance it was found clinging to the undersideof a small floating piece of "cave ice" on the surfaceof the pool. When disturbed, this species quicklyrolls into a tight ball and falls to the bottom of thepool.Mexilana saluposi BowmanCirolanidae, undescribed member of: Hobbs andHobbs, 1973:39,41; Hart and Hart, 1974:165.Speocirolana: Magniez, 1973:163 (misidentification).Mexilana saluposi Bowman, 1975:2-6,7, fig. 1-3.Mexila1la: Bowman, 1975:1-2,5,6,7, fig. 4d.Type-Iocality.-Cueva del Huisache, San Luis Potosi,Mexico.Distribution.-Known only from the type-locality.See Fig. 6.Discussion.-This species is the only member ofthe genus Mexilana. It has been found in associationwith an undescribed anthurid isopod of the genusCyathura and with the stenasellid isopods Mexistenasellusparzefalli and M. wilkensi. The fauna of Cuevadel Huisache is very different from that of the nearbySierra de El Abra. Our knowledge of the area inwhich this cave is located is still too poor to allowany speculations as to why the faunas of the two regionsare so unlike.Speocirolana bolivari (Rioja)Cirolana sp.: Rioja, 1951a: 170, 178; Rioja, 1953a:290,291,293.Cirolana, otra especie: Villalobos, 1951:215.Cirola1la (part): Bonet, 1953b:242; Rioja et al.,1961:315.Cirolana (Spaeocirolana) nueva especie: Rioja, 1953a:289 (erroneous spelling).Cirolana (Speocirolana) bolivari Rioja, 1953d:147,148, 149-160, fig. 1-3, 5-8, 10-32, 34; Rioja,1957:455; Nicholas, 1962:170; Rioja, 1962:40;Vandel, 1964:150; Vandel, 1965a:119.Cirolana bolivari: Rioja, 1953e:11, fig. 5; Paclt,1971:424; Rioja, 1971:509; Sbordoni and Argano,1972:9.Speocirolana (part): Bowman, 1964:233,.234; Straskraba,1969:23; Argano, 1972b:33; Bowman,1975:5,6,7, fig. 4c.Speocirolana bolivari: Bowman, 1964:236, fig. 61;Reddell, 1965a:19; Cole and Minckley, 1966: 18,19, 20, 21; Reddell, 1967b:82; Minckley andCole, 1968:3,4; Edwards, 1971:3; Elliott, 1971:1,10, 17, 19,20,27-28,34,37-38,39,40,44, fig.12-13; Hobbs, 1971:3, 46; Reddell, 1971b:21;Reddell and Mitchell, 1971a: 142; Reddell andMitchell, 1971b:183, 184; Elliott, 1973a:31;Elliott and Mitchell, 1973:171, 175, 176, 179,181-182, 184, 185, 186-187, 188, fig. 5; Hobbsand Hobbs, 1973:41; Holt, 1973:4, 13; lVlagniez,1973:163; Reddell and Elliott, 1973a:171, 172;Reddell and Elliott, 1973b:181, 183; Hart andHart, 1974:164.Speocirolano bolivari: Holt, 1973: 11 (erroneousspelling).Blind isopods: Reddell, 1973b:77.Type-Iocality.-Cueva (=Grutas) de Quintero, km560 de la carreterra de Valles al Mante, Tamaulipas,Mexico.Distribution.-Known from one cave each in theSierra de El Abra and Sierra de Guatemala, Tarnaulipas;and the Aquismon region, San Luis Potosi. SeeFig. 6.Records.-Tamaulipas: Bee Cave and Grutas deQuintero.New record.-San Luis PotOSI: Spring at La Laja(det. T. E. Bowman).Discussion.-This species is abundant in Grutas deQuintero, where it cohabits standing pools with S.pelaezi. Although it presumably ranges throughoutthe Sierra de EI Abra, it is obviously very rare. Thegenus Speocirolana is represented only by the threetroglobites listed here and by undescribed speciesfrom near Linares, Nuevo Leon, and San Nicolas, SanLuis Potosi. Elliott and Mitchell (1973) examinedthe temperature preference responses of this speciesand found that it still retains a temperature preferendum.They speculate that the species is the descendantof a marine species which directly colonizedcaves along the front of the Sierra de EI Abra, perhapsas early as the Cretaceous.Speocirolana pelaezi (Bolivar)Cirolanidos: Anonymous, 1942a:221.Cirolana nueva especie: Osorio Tafall, 1942:210;Osorio TafaU, 1943:46,50,51; Rioja, 1953e:ll.86

Cirolana (part): Osorio TafaU, 1946:153; Bonet,1953b:242; Rioja et al., 1961:315.Cirolana pelaezi: Osorio Tafall, 1943:54-55,57,67,69 (nomen nudum); Bonet, 1953b:241; Rioja,1953a:288; Tamayo, 1962:vol. III:260; Vandel,1964:303; Vandel, 1965a:254; Sbordoni and Argano,1972:9.Cirolana (Speocirolana) pelaezi Bolivar, 1950:211,212-216, 217, 218, fig. 1-12; Rioja, 1951a:170,178; Villalobos, 1951:215; Rioja, 1953d:147,148, 149, 151, 152, 159, fig. 4, 9, 33, 35; Rioja,1957:455; Hart, 1962:139; Nicholas, 1962: 170;Rioja, 1962:40; Vandel, 1964:150; Schultz, 1965:101,102; Vandel, 1965a:119; Barrera, 1968:309.Cirolana (Spaeocirolana) pelaezi: Rioja, 1953a:288,289, -290, 291, 293 (erroneous spelling).Conilera stygia Packard: Dearolf, 1953:226 (misidentification)_Speocirolana (part): Bowman, 1964:233-234; Straskraba,1969:23; Argano, 1972b:33; Bowman,1975:5,6,7, fig. 4c.Speocirolana pelaezi: Bowman, 1964:236, fig. 61;Reddell, 1965a:19; Reddell, 1965d:49; Cole andMinckley, 1966:18, 19, 20, 21; Reddell, 1967b:82; Minckley and Cole, 1968:3,4; Elliott, 1971:10; Hobbs, 1971:3, 46; Reddell, 1971b:21; Reddelland Mitchell, 1971a:141-143, fig. 10, 27;Reddell and Mitchell, 1971b: 183; Reddell andMitchell, 1971d:cover; Elliott and Mitchell, 1973:175; Hobbs and Hobbs, 1973:39, 40, 41; Holt,1973:4, 13; Magniez, 1973:163; Mitchell andKawakatsu, 1973a:671; Reddell and Elliott,1973a:171, 172; Reddell and Elliott, 1973b:181,183; Hart and Hart, 1974:164; Fernandez Ruiz,1976:717; Danielpol, 1977:35.Speocirolano pelaezi: Holt, 1973: 11 (erroneous spelling).Type-locality.-Cueva de Los Sabinos, cerca deValles, San Luis Potosi, Mexico.Distribution.-Known from caves in Puebla, SanLuis Potosi, and Tamaulipas. See Fig. 6.Records.-San Luis Potosi: Sotano del Arroyo,Sotano de Coatimundi, Cueva de la Curva, CuevaChica, Cueva de La Lagunita, Sotano de Matapalma,Sotanito de Montecillos, Sotano de Pichijumo, SOtanode las Piedras, Cueva de Los Sabinos, Sotano delTigre, Sotano de la Tinaja, and Sotano de Yerbaniz;Tamaulipas: Cueva de la Florida, Cueva de la Mina,Sotano del Molino, Cueva de EI Pachon, Grutas deQuintero, Cueva de Ojo de Agua de Manantiales, andSotano de Vasquez.New records.-Puebla: Grutas de Atepolihuit; SanLuis Potosi: Cueva del Agua, Cueva de Agua Vendita,Cueva de Berna Be B, Cueva de Las Lagunitas, Springat La Laja, Cueva de la Luz, Small cave near HoyaQuital, and Hoya Quital; Tamaulipas: Cueva del Aguade Simon Salinas and Bee Cave (det. T. E. Bowman).Discussion.-This species is remarkable because ofits great range, occurring in the Sierra de Tamaulipas,Sierra de EI Abra, Sierra de Guatemala, Aquismon region,near Ciudad de Maiz in the Sierra el Pino, and inthe Cuetzalan region. In addition it has an altitudinalrange of at least 1200 meters. It usually inhabits stillpools and lakes floored with silt or guano. It has beentaken in direct association with S. bolivari in Grutasde Quintero, Bee Cave, and the spring at La Laja.Speocirolana thermydronis Cole and MinckleySpeocirolana thermydronis Cole and Minckley, 1966:17-22, fig. 1-21; Minckley and Cole, 1968:2-4;Minckley, 1969:24-25; Cole and Minckley, 1970:71, 80; Holsinger and Minckley, 1971:441; Reddell,1971b:21; Cole and Minckley, 1972:321.Speocirolana: Taylor and Minckley, 1966:21.Speocirolana (part): Straskraba, 1969:23; Argano,1972b:33; Bowman, 1975:5,6, 7, fig. 4c.Type-locality.-Posos de la Becerra, 9.8 miles SSWCuatro Cienegas, Coahuila, Mexico.Distribution.-Known only from spring-fed "posos"southwest of Cuatro Cienegas de Carranza, Coahuila.See Fig. 6.Records.-Coahuila: Posos de la Becerra, EI Mojarral,and an unnamed poso (12 km SW Cuatro Cienegas).Discussion.-Speocirolana thermydronis has beentaken from interstices in blocks of porous travertinealong the sides of natural wells (posos). The habitatin which these isopods has been found is well describedby Minckley (1969). Cole and Minckley(1966) speculate that the genus is of pre-Tertiaryorigin; they also discuss the evolution of the genus.Sphaerolana affinis Cole and MinckleyAberrant cirolanid (part): Minckley, 1969:25.Sphaerolana affinis Cole and Minckley, 1970:75-76,78-79, 80, fig. 28-40; Holsinger and Minckley,1971:441; Reddell, 1971b:21; Peck, 1977c:67.Sphaerolana (part): Cole and Minckley, 1970:71-72;Cole and Minckley, 1972:321.Type-locality.-Small poso ca. 20.3 km south and5.5 km east of Cuatro Cienegas de Carranza, Coahuila,Mexico.Distribution.-Known from spring-fed wells nearCuatro Cienegas, Coahuila; and from a flooded minenear Villa Santiago, Nuevo Leon. See Fig. 6.Records.~oahuila: Small poso about 20.3 kmSand 5.5 km E of Cuatro Cienegas; other posos nearCuatro Cienegas; Nuevo Leon: Flooded mine about87

30 m below entrance of Cueva de la Boca, near VillaSantiago.Discussion.-Sphaero/ana is one of three genera ofcirolanids capable of rolling into a ball, the other twobeing Faucheria and Creaseriella (see under Creaseriellaanops above). It appears to be only slightly relatedto either genus. The two species of the genus arequite secretive during the day but roam in the springmouths and in mud about plant roots in the bottomof the wells at night. Cole and Minckley (1970) discussthe evolution of the genus.Sphaerolana interstitia/is Cole and MinckleyAberrant cirolanid (part): Minckley, 1969:25.Sphaerolana interstitia/is Cole and Minckley, 1970:72-75, 76, 78-79, 80, fig. 1-27,40; Holsinger andMinckley, 1971:441; Reddell, 1971b:21; Brown,1974:44,45.Sphaerolana (part): Cole and Minckley, 1970:71-72;Cole and Minckley, 1972:32l.Type-Iocality.-Small, unnamed pit ca. 8.2 kmsouth and 4.7 km west of Cuatro Cienegas de Carranza,Coahuila, Mexico.Distribution.-Small springs and posos near CuatroCienegas, Coahuila. See Fig. 6.Records.-Coahuila: Small unnamed poso about8.2 km Sand 4.7 km W of Cuatro Cienegas; threesmall springs within 1 km W of this poso.Discussion.-One specimen of this species wastaken from the stomach contents of the aquatic boxturtle T~rrapene coahuila Schmidt and Owens(Brown, 1974). It has been taken in association withS. affinis in one poso (Cole and Minckley, 1970).Suborder AsellotaFamily AsellidaeThe family Asellidae is an extremely abundantpart of the cavernicole fauna of the temperate regionsof the United States, but until recently was notknown from Mexico. Five species of asellid, four ofwhich are troglobites, have been described from wellsand caves in southern Mexico: a single troglobitic speciesis known from a cave in Guatemala.Caecidotea chiapas BowmanCaecidotea chiapas Bowman, 1976:340-345, 354,355, fig. 1-39; Argano, 1977:103-105, 1l0, 112,114, 116; Ruffo and Vigna Taglianti, 1977:134,141; Sbordoni et aI., 1977:74, pI. lia.AseUi troglobi: Sbordoni et al" 1977:56.Type-Iocality.-Cueva de los Iurcielagos, 15 kmESE of San Cristobal de las Casas, Chiapas, Mexico.Distribution.-Known from four caves and a wellnear San Cristobal de las Casas, Chiapas. See Fig. 7.Records.-Chiapas: Well in casa Bell (San Cristobalde las Casas), Cueva de Los Llanos, Cueva- de los Murcielagos,Cueva de la Planta n. 2, and Cueva de laPlanta n. 3.Discussion.-This species is most closely related toC. pasquinii. It was taken from a small stream IIICueva de Los Llanos and Cueva de los Murcielagos.Caecidotea mitchel/i ArganoCaecidotea mitchelli Argano, 1977:112-116, fig. 5-6;Ruffo and Vigna Taglianti, 1977:146.Type-Iocality.-Cueva de los Resadores, Santa Eulalia,Huehuetenango, Guatemala.Distribution.-Known only from the type-locality.See Fig. 7.Discussion.-This species was taken from pools inassociation with Bogidiel/a pasquinii. It is a memberof the chiapas species group and appears to be closelyrelated to C. pasquinii.Caecidotea pasquinii (Argano)Conasel/us pasquinii Argano, 1972a:35-42, fig. 1-3;Bowman, 1976:339.Caecidotea pasquinii: Bowman, 1976:344, fig. 39;Argano, 1977:103, 112.Type-Iocality.-Well in the pueblo of San Joan(=Juan) de.la Punta, on the road between Veracruzand Cordoba, Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 7.Discussion.-This species is closely related to C.communis (Say), a widespread species known, inMexico, from the states of Mexico, Puebla, and Veracruz.Argano (1972a) considers that C. pasquinii is afairly recent troglobite, probably evolved from C.communis or a common ancestor. Creaser (1938)reported the presence in Gru tas de Balankanche,Yucatan, of an undescribable species of Caecidotea.Caecidotea vomeroi ArganoCaecidotea vomeroi Argano, 1977:110-112, fig. 4;Ruffo and Vigna Taglianti, 1977:13l.Type-Iocality.-Cueva de Chita! no. 2, Ococingo(=Ocosingo), Chiapas, Mexico.Distribution.-Known only from the type-locality.See Fig. 7.Discussion.-This species belongs to the chiI.Jpasspecies group and appears to be most closely relatedto C. zul/inii. It was taken i~ association with theamphipod Bogidie/la tabascl!tl.sis and troglobitic planaria.88

Caecidotea zullinii ArganoCaecidotea zullmii Argano, 1977:105.110, 112, 114,fig. 1-3; Ruffo and Vigna Taglianti, 1977:151.Type-Iocality.-Cueva de Chanchaniptic, Sitala,Chiapas, Mexico.Distrihution.-Known only from the type-locality.See Fig. 7.Discussion.-This species is a member of the chiapasspecies group and appears to be closely related toC. vomeroi. It was taken in association with Bogidiellavomeroi and troglobitic planaria.Family StenasellidaeThe family Stenasellidae is found in the periMediterranean area, east-central Asia, tropical Africa,and Mexico. The appearance of the family in theCuatro Cienegas basin implies a pre-Tertiary originfor the family and provides a further link betweenthe peri-Mediterranean and greater Caribbean regions(Cole and Minckley, 1972). Magniez (1974a) has discussedthe evolution and ecology of the family. Hecontends that the stenasellids had their origin in thelittoral gravels, from which they migrated into the(//.//('GUATEMALA~'" r',--Fig. 7.-Distribution of troglobitic isopods of the suborder Asellota: 1,Mexistenasellus coahuila; 2,Mexistenasellus n. sp.l;3, Mexistenasellus n. sp. 2; 4, M. parzeJalli and M. wilkensi; 5, M. magniezi; 6, Caecidotea pasquinii; 7, Etlastenasellus n. sp.;8,E. mixtecus; 9, C. chiapas; 10, C. zullinii; 11, C. vomeroi; 12, C. mitchelli; 13, Caecidotea sp.89

underflow of rivers, alluvial gravels, and finally thekarstic waters.Etlastenasellus mixtecus ArganoEtlastenasellus mixtecus Argano, 1977:117-121, fig.7-8; Ruffo and Vigna Taglianti, 1977:166.Type.locality.-Pozzo, villaggio di Etla, Oaxaca,Oaxaca, Mexico.Distribution.-Known only from the type-locality.See Fig. 7.Discussion.-Etlastenasellus is a monotypic genuswhich shows some affinities both to the MexicanMexistenasellus and to the African Magniezia. It wastaken from phreatic waters in association with Bogidiellaarganoi, B. niphargoides, and B. michaelae.Mexistenasellus coahuila Cole and MinckleyStenasellinae, undescribed genus: Minckley, 1969:25; Holsinger and Minckley, 1971:44l.Mexistenasellus coahuila Cole and lVlinckley, 1972:314, 315-319, fig. 1-31; Magniez, 1972:19, 20,23, 29; Holsinger, 1973:11; Magniez, 1973:164,168; Argano, 1974:102; Magniez, 1974a:50, 51,56,57,58,59, fig. llC, 13; Magniez, 1974b:100;Argano,1977:116.Mexistenasellus coahuili: Magniez, 1974a:57 (erroneousspelling).Mexistenasellus (part): Husson et aI., 1973: 122; Magniez,1973:181; Magniez, 1974a:7, 9, 25, SO-51,56-57,64,69, fig. 1; Bowman, 1976:339.Type-Iocality.-Eastern end of Laguna Juan Santos,8.5 km south and 8.0 km west of Cuatro Cienegasde Carranza, Coahuila, Mexico.Distribution.-Known only from springs andspring-fed pools southwest of Cuatro Cienegas, Coahuila.See Fig. 7.Records.-Coahuila: Pozo Barbado, Laguna Escobeda,Laguna Juan Santos, Small unnamed spring (8.2km S, 8.4 km W Cuatro Cienegas), Small unnamedspring (8.8 km S, 4.0 km W Cuatro Cienegas), andSmall unnamed seep near Pozo Barbado.Discussion.-The genus Mexistenasellus is knownonly from the four subterranean fOTIns discussed hereand from undescribed species from Nuevo Leon andTamaulipas.Mexistenasellus magniezi ArganoMexistenasellus magniezi Argano, 1974:98-102, fig.1-3; Magniez, 1974a:58, 59, fig. 13; Ruffo andVigna Taglianti, 1974:120; Argano, 1977:116.Mexistenasellus (part): Magniez, 1974a:7, 9, 25, 50,51,56-57,64,69, fig. 1; Bowman, 1976:339.Type-locality.-Well on the outskirts of the villageof Paraje Nuevo, near Cordoba, Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 7.Discussion.-This species is very similar to JH. coahuilain its dimensions and general body form. Sinceboth of these species inhabit a similar (phreatic) biotope,this is not surprising.Mexistenasellus parzeJalii MagniezMexistenasellus parzeJalii Magniez, 1972:19-24, 30,pI. 1-3; Hobbs and Hobbs, 1973:39; Husson et al.,1973:122; Magniez, 1973:163-170, pI. 1-2; Argano,1974:102; Magniez, 1974a:50, 51, 57-58,59, fig. llC, 13; Magniez, 1974b:122, 123, 125,126,128; Bowman, 1975:7; Argano, 1977:116.Mexistenasellus (part): Magniez, 1973:181; Magniez,1974a:7, 9, 25, 50-51, 56-57, 64,69, fig. 1; Bowman,1976:339.Type-locality.-Cueva del Huizache (=Huisache),Ciudad Valles, San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 7.Discussion.-This species is more closely related toJ{ wilkensi than to the other two species of the genus.It is very different from 11-1. coahuila and M. magnieziwhich inhabit phreatic waters rather than theopen waters of caves. It was found crawling on thebottom of a small pool OVi'r whil'h bats roosted.Mexistenasellus u;ilkensi \la~lliezMexistenasellus u;ilkensi :\lagniez, 1972: 19, 20, 23,24-29, :.lO, pI. 4-6; Hobbs and Hobbs, 1973:39;Husson et al., 1973:122; Magniez, 1973:163,164; Argano, 1974:102; lagniez, 1974a:1O-ll,57,58,59, fig. 2C, 13; Magniez, 1974b:115, 123,126; Bowman, 1975:7; Argano, 1977:116.Mexistenasellus (part): Magniez, 1973:181; Magniez,1974a:7, 9, 25, 50-51, 56-57, 64, 69, fig. 1; Bowman,1976:339.Type-Iocality.-Cueva del Huizache (=Huisache),Ciudad Valles, San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 7.Discussion.-This species is most closely related toM. parzeJalii. It was found in direct association withM. parzeJalli, the cirolanid Mexilana saluposi, and anundescribed anthurid of the genus Cyathura.Suborder AnthurideaFamily AnthuridaeCyathura sbordonii ArganoCyathura sbordonii Argano, 1971:304, fig. 1-4; Argano,1972b:26-33, fig. 1-3; Peck and Peck, 1973:68.90

Cyathura new species: Sbordoni and Argano, 1972:19.Type-Iocality.-Cueva del Ojo de Agua Grande, ParajeNuevo, Cordoba, Veracruz, Mexico.Distribution.-Known only from two caves inVeracruz. See Fig. 8.New record.-Veracruz: Cueva de la Sala de Agua(Ejido Colonia) (det. T. E. Bowman).Discussion.-The genus Cyathura consists primarilyof marine species, but four freshwater forms havebeen described. One of these, C. milloti Chappuis,Delamare, and Paulian, is known only from an interstitialhabitat on Reunion Island; it is not closely relatedto the other three freshwater species. Thesethree species constitute the specus group (Argano,1972b). Cyathura specus Bowman is known from acave in Cuba, C. currassavica Storck from a spring inthe island of Cura~ao, and C. sbordonii from a cavein Veracruz. Cyathura sbordonii is most closely relatedto C. specus. The distribution of the subterraneanCyathura is similar to that of the Cirolanidae,Stenasellidae, Bogidiellidae, and Hadziidae. Argano(1972b) speculates that the three subterranean speciesevolved from a common interstitial ancestorwidely distributed throughout the Caribbean region.The recent discovery of an undescribed species nearValles in San Luis Potosi only serves to emphasize,.".\\\"-\\i-V>-" "\" \ ~ \,\"-~j ,,)\I ,I\I(I,\~JMEXICO//'/I ~.: I;, I~"'- -L. .---t-f....·1~ I I'\._iI'-i.------_._'//i\./I....." i\..., Ii IIIIIGUATEMALAFig. 8.-Distribution of troglobitic isopods of the suborders Anthuridea and Microcerberidea: 1, Mexicerberus troglodytes;2, Cyathura sp.; 3, C. sbordonii.91

the similarities between the distribution of thisgroup and that of the others.Suborder MicrocerberideaFamily MicrocerberidaeThe family Microcerberidae in North America isrepresented only by marine interstitial forms of thegenera Microcerberus and Yvesia and by the troglobitediscussed below. Although some species ofMicrocerberus have been collected from caves in Europe,the species discussed below was the first caverecord for the suborder in the New Worid. An undescribedspecies of Microcerberus is known fromCueva Pinta, San Luis Potosi.Mexicerberus troglodytes SchultzMexicerberus troglodytes Schultz, 1974:309, 311,fig. 1-7.Type-Iocality.-Cueva de la Mina, 7 km northwestof Gomez Farias, Municipio de Gomez Farias,Tamaulipas, Mexico.Distribution.-Known only from the type-locality.See Fig. 8.Discussion.-This species was collected in a smallshallow drip pool near the end of Cueva de la Mina.That it is a true cave form and not merely an interstitialspecies is indicated by its appearance in a caveat high altitude in association with other undisputedtroglobites and by its great size (2.5 mm as opposedto 1.6 mm in previously known species).Suborder OniscoideaThe largest group of isopods inhabiting caves inMexico is the primarily terrestrial Oniscoidea. Ofseven families represented in Mexican caves four containtroglobites. The terrestrial isopods of Mexico,Guatemala, and Belize are now under study by Dr.George A. Schultz, so that any list of species must beconsidered premature. At the present time 15 speciesof oniscoid isopod are known to be troglobites,whereas 15 are considered to be troglophiles. Some ofthe troglobites listed below may prove to be identicalwith forms inhabiting the endogean zone.Family ArrnadillidaeThe family Armadillidae is the principal contributorto the troglophile fauna of Mexican caves. Eightspecies of the genus Venezillo and one species ofCubaris have been identified from caves in Mexico(Mulaik, 1960). The following species are consideredto be troglophiles: Cubaris mirandai Rioja fromCueva del Ojo de Agua Grande, Veracruz (Rioja,1954); Venezillo nrticulatus (Mulaik) and V. boneti(Mulaik) from Grutas de Juxtlahuaca, Guerrero(Mulaik, 1960); V. cacahuamilpensis (Bilimek) fromcaves in Guerrero and Mexico (Bilimek, 1867; Mulaik,1960; Rioja, 1954); V. chiapensis Rioja from Grutasde Zapaluta, Chiapas (Rioja, 1955a); V. llamasi Riojafrom Cueva de Patla, Puebla (Rioja, 1954); V. osorioi(Mulaik) from caves in Guerrero and Nuevo Leon(Mulaik, 1960); V. pleogoniophorus (Rioja) fromCueva de Los Sabinos, San Luis Potosi (Rioja, 1952);and V. tanneri (Mulaik and Mulaik) from Grutas delPalmito, Nuevo Leon (Schultz, 1965). These isopodsare usually found on bat guano or organic debris,and none show any adaptations for a cave existence.Family OniscidaeOne species of the family Oniseidae has been reportedfrom Mexican caves. Hoctunus vespertilio lVlulaikwas described from Cenote de Hoctun, Yucatan(Mulaik, 1960). This species was listed by Nicholas(1962) as a troglobite, but it should be considered atroglophile. It was found on bat guano.Family PhilosciidaeColombophiloscia cavernicola \" allddColombophiloscia cavernicola Valid,·!. I ')()Bh:lJ8,102-104,156,162, fig. 30-:J2:Schullz. llJ77:12,13, fig. 19-24.Type-Iocality.-Cueva de Quijano, a Caripe, di~tritoMonaguas, dans la region Jlord-orientalt- duVenezuela; or Cueva del Guacharo, pres de Caripe,distrito Monaguas.Distribution.-Known from two caves in DistritoMonaguas, Venezuela, and one cave in Huchuetenango,Guatemala. See Fig. 9.Guatemala record.-Huehuetenango: Cueva de losResadores.Discussion.-Vandel (1968b) does not indicatewhich of the two Venezuelan caves is the typelocality.The distribution of this blind species defiesexplanation. Although Schultz (1977) found no differencesbetween Vandel's description of the speciesand the specimen from Cueva de los Resadores, it isconceivable that study of the type series might revealspecific differences. Two other species of thegenus Colombophiloscia are known. C. alticola Vandelwas described from humus in a cave at Banos,Ecuador, but it has normal eyes. C. naevigesta Vandelwas described from a deep crevasse on Isla SantaCruz in the Galapagos Islands and is eyelpss (Vandel,1968b).92

C \{Troglophiloscw laevis SchultzTroglophiloscw laevis Schultz, 1977:9-12, 13, fig.1-18.Troglophiloscia levis: Reddell, 1977b:229, 239,240,273 (erroneous spelling).Type-locality.-ActUn Xpukil, 3 km S of Calcehtok,Yucatan, Mexico.Distribution.-Known only from the type-locality.See Fig. 9.Discussion.-This species is known only from asingle male. The only other species in the genus is T.silvestrii Brian, a troglobite described from Grutas deBellamar, near Matanzos, Cuba.Family PorceUionidaeThe family Porcellionidae is represented in Mexicancaves by two species of the genus Porcellio andby one of Porcellionides. Porcellio gertschi VanName was collected in Cueva de los Lagos, Coahuila(Schultz, 1965). Porcellio laevis Latreille was identifiedfrom Grutas de Cacahuamilpa, Guerrero, andfrom Cenote lxii, Yucatan. Creaser (1938) reportedthe genus Porcellionides from numerous caves in,~,,•MEXICOGUATEMALA.--,.. ./r·\r"//Fig. 9.-Distribution of troglobitic and troglophilic isopods of the families Philosciidae, Sphaerioniscidae, and Squamiferidae:1, Spherarmadillo cavernicola; 2, Trichorhina boneti; 3, T. atoyacensis; 4, T. vandeli; 5, T. pearsei; 6, Troglophiloscialaevis; 7. Colombophiloscia cavernicola.93

Yucatan but did not identify the species; he indicatedthat it was probably a troglobite. Numerousrecent collections from the Yucatan Peninsula containspecimens probably applicable to that reportedby Creaser, but remain unstudied.Family SphaeroniscidaeSpherarmadillo cavernicola MulaikSpherarmadillo cavernicola Mulaik, 1960:84, 179,180-181, fig. 334-341; Nicholas, 1962:170;Schultz, 1970b:125, fig. 11-17; Reddell, 1971b:22; Reddell and Mitchell, 1971a:143; Reddell andMitchell, 1971b: 183; Reddell and Elliott, 1973a:171; Reddell and Elliott, 1973b:181; FernandezRuiz, 1976:717.Type.locality.-Cueva de los Sabinos, San Luis PotosI,Mexico.Distribution.-Known from one cave each in theSierra de El Abra, San Luis PotosI, and the Sierra deGuatemala, Tamaulipas. See Fi~. ~.Records.-San Luis Potosi: Cueva de Los Sabinos;Tamaulipas: Cave (Rancho del Cielo).Discussion.-A record of this species from Huatusco,Veracruz (Mulaik, 1960) is doubtlcss an errorand is presumably applicable to S. huatuscensis .Vlulaik.The species belonging to the genus Spherarmadilioare only very poorly separated from those ofthe genus Sphaeroniscus. Spherarmadillo cavemicolais more closely related to S. huatuscensis from Veracruzthan to the only other species of the genus, S.schwarzi Richardson, from Guatemala. Schultz (pers.comm.) expresses the opinion that this spccies maybe an endogean form, but it is apparently still knownonly from the cave habitat. It is, thereforc, rctainedin this list with some reservations.Family SquamiferidaeThe family Squamifcridae is represented in :\'lcxicancaves by four species of the genus Trichorhina,three of which are anophthalmic and included here.A fourth species, T. vandeli Rioja, is known onlyfrom Cueva Cerro Hueco, Chiapas, and possessesreduced eyes; it is probably a troglophile. The remainingfour species are epigean forms. Vandel(1964) indicated that all of these species are prob.ably endogeans, and he may be correct. The epigeanisopod fauna of Mexico is only poorly known, anduntil careful collections of the endogean habitat havebeen made, any speculations on the relationships ofthese species are premature.Trichorhina atoyacensis MulaikTrichorhina atoyacensis Mulaik, 1960:83, 140, 141,fig. 127-130; Nicholas, 1962:171; Lemos de Castro,1964:1; Reddell, 1971b:22.Type.locality.-Cueva (=Grutas) de Atoyac, Atoyac,Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 9.Discussion.-Schultz (pers. comm.) has indicatedthat this species may be synonymous with T. pearsei(Creaser) and T. boneti Rioja. If these species provcto be identical, they are almost ccrtainly not troglohites,since these cave regions are so widely separatet]and the troglobitic faunas so distinct.Trichorhina boneti RiojaOniscidae sp. D: Bonet, 1953a:24, 65.La forma aGn al genera Troglophiloscia: Bond,1953a:27, 30, 33.Trichorhina boneti Rioja, 1956:451-456, fig. J5-27;:\lulaik, 1960:83, 140, 1-1,4-147, fig. 618-630;\icholas, 1962:171; Lcmos de Castro, 1964:J;Reddell, 19C>7d:l06; Reddell, 197111:22.Oniscidac, unidentified genus and species: Redddl,1971b:2J (Cueva del Ahuate n. 2 record only).Type.locality.-Cueva de Ahuate numero dos,pucrto de Ahuate, al SO. de Xilitla, San Luis PotosI,.\:Iexico.Distribution.---Known only from the tnw-Iocality.See Fig. 9.Discussion.~This is the most northern record forthe genus in Mcxieo. Schultz (pers. comll1.) has indicatedthat this species may be synonymous with T.pearsei.Trichorhina pearsei (Creaser)Porcellio: Pearse, EI:38a: 13, 15; Pearse, 1945:Ill,166.Porcellio pearsei Creaser, 1938: L60, 162, 163, fig.1-8; Pearse, 1945:166; Rioja, 1953a:286, 295;\Iulaik, 1960:166, 170-171; Nicholas, 1962:171.Porcellio pearsi: Vandel, 1950:206, 207 (erroneousspelling).Trichorhina yucatanensis Mulaik, 1960:83, 140,142, 144, fig. 146-150; Lemos de Castro, 1964:J,2.Trichorhina yucatensis: l'\icholas, 1962:171 (erroneousspelling).Trichorhina pearsei: Lemos de Castro, 1964:1, 2;Reddell, 1971b:22; Reddell, 1977b:229, 239,240,268,270,273,274,276,285.94

Type-Iocality.-Of Porcellio pearsei: Balaam CancheCave (=Grutas de Balankanche), near ChichenItza, Yucatan, Mexico; of Trichorhina yucatanensis:Cueva de Gorgosa (=Actun Gongora), Oxkutzcab,Yucatan, Mexico.Distribution.-Known from seven caves in Yucatan.See Fig. 9.Records.-Yucatan: Gru tas de Balankanche, ActlInGongora, Actun Puz, Cenote de Sambula (l\lotul),Cueva Primera del Camino a San Roque, Actun Sazich,and Actun Xpukil.Discussion.--This is an eyeless species hut mayprove to be an endogean form. One eyeless epigeanspecies, T. xoltumae Mulaik, is known from Xoltum,Yucatan (Mulaik, 1960). :'iumerous collections ofcave and epigean isopods from the Yucatan Peninsulaare now under study by Dr. George A. Schultz.Further speculation on the ecologic status of thisspecies is premature.Family TrichoniscidaeBrackenridgia acostai (Rioja)Protrichoniscus acostai Rioja, 195Ih:181-189, fig. 123; Rioja, 1953a:290, 291, 295: Rioja, 195::3b:217; Rioja, ] 955b:49-53, 61. fig. 19-26; '\lulaik,1960:82, 100, 102-105, fig. 552-570; l\icholas,1962: 172; Vandel, 1965c:352, :l5~, fig. I.Protrichoniscus (part): Rioja, 1953e: 11.Protrichonischus acostai: Rioja, 1955a:208 (erron(~ousspelling).Brackenridgia acostai: Keddell, 1971b :23.Type-Iocality.-Cueva de toma de ai,rtla para lapoblacion de Comitan (=Cueva del Tio Ticho), Chiapas,Mexico.Distribution.-Known from two caves near Corn itan, Chiapas. See Fig. 10.Records.-Chiapas: Cueva dcl Tio Ticho and Gmtasde Zapaluta.Discussion.-The genus Brackenridgia is a memherof the First Division of the subfamily Trichoniscinae.This division is considered by Vandel (1965c) to bethe most primitive. Members of it are found in Europeand North America. Brackenridgia includeseight species, seven of which are troglobites. Theeighth, B. heroldi (Arcangeli), is known only fromcaves and epigean localities in California (U.S.A.).Two species, B. cavernarum Ulrich and B. reddelli(Vandel), are known only from caves in Texas.Brackenridgia bridgesi (Van Name)Protrichoniscus bridgesi Van Name, 1942:299,302304, fig. 4-5; Bolivar, 1950:211; Rioja, 1950:127-137, 138, 140, 141, 143, fig. 1-20; Vandel,1950:206, 207; Rioja, 1951b:183, 184, 187,188; Villalobos, 1951:215; Bonet, 1953a:27,30;Rioja, 1953a: 287, 289, 291, 295; Vandel, 1953:371, 372; Rioja, 1955b:39-44, 46, 53, 61, fig.1-10; Mulaik, 1960:100, 101-102, 103, 104, 105,106, 107, 108, 109, 110, fig. 27-38; Nicholas,1962:172; Reddell, 1965a:19; Vandel, 1965c:353, 354, 360-364, 366, fig. 1, 5-6; Reddell,1967b:82; Schultz, 1968:256-257, fig. 3, 6, 9,12,15,18,21,26-27; Schultz, 1970a:412.Protrichoniscus (part): Rioja, 1953e:ll, fig. 7.Protrichoniscus potosinus Mulaik, 1960:100-10],fig. 21-26; Nicholas, 1962:172; Bowman, 1965:212; McKenzie, 1965a:38; Vandel, 1965c:353,354,360.Brackenridgia bridgesi: Mitchell and Reddell, 1971:59; Reddell, 1971h:23; Reddell and Mitchell,1971a:143, fig. 9; Reddell and Mitchell, 1971b:183, fig. 1, 15; Mitchell and Kawakatsu, 1973a:671, 673, 675; Reddell and Elliott, 1973a:171;Reddell and Elliott, 1973b:181; Fernandez Ruiz,1976:716, 717; Johnson and Heath, 1977:131135.Type-Iocality.-Of Protrichoniscus bridgesi: Caveat El Pujal (=Cueva Chica), San Luis Potosi, Mexico;of Protrichoniscus potosinus: Cueva Chica, El Pujal,San Luis Potosi, Mexico.DistributiolJ.-Known from caves in the Sierra deEl Abra, San Luis Potosi and Tamaulipas; and theSierra de Guatemala, Tamaulipas. See Fig. 10.Records.-San Luis Potosi: Boca del Ahra, Sotanodel Arroyo, Cueva Chica, Ventana Jahali, Cueva deLos Sabinos, Sotano del Tigre, and Sotano de laTinaja; Tamaulipas: ?Cueva de la Capilla, Cueva dela Mina, Cueva de El Paehon, and Grutas de Quintero.DiscussiolJ.-Boca del Abra is an unidentified localitybut may be the same cave as Cueva de Valdosa.This widespread species may also be found in a varietyof habitats, both terrestrial and aquatic. Individualshave been seen to walk though pools severalinches deep and even to be seen remaining underwater for long periods of time. It is usually collectedon small pieces of rotten wood, among bits of organicdebris washed into caves, or about small pools. Johnsonand Heath (1977) estimated the population sizein Cueva Chiea to be about 533 individuals. This speciesshows strong affinities with B. reddelli of CentralTexas (Vandel, 1965c).Brackenridgia palmitensis (Mulaik)Protrichoniscus palmitensis Mulaik, 1960:100, 102,fig. 39-48; Nicholas, 1962: 172; Bowman, 1965:212; Vandel, 1965c:353, 354; Reddell, 1967a:24.95

Protrichoniscus: Reddell, 1967a:24.Brackenridgia palmitensis: Reddell, 1971b:23.Type-Iocality.-Cueva (=Grutas) del Palmito, Bustamante,uevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 10.Discussion.-The description of this species is inadequateto allow speculation on its relationship toother members of the genus. A single small isopod,probably of this species, has been collected from organicdebris in a small side passage near the entranceto Grutas del Palmito.Brackenridgia villalobosi (Rioja)Protrichoniscus villalobosi Rioja, 1950: 127, 128,129, 132, 137-145, fig. 21-49; Rioja, 1951b:183,184, 187, 188; Rioja, 1953a:289, 291, 295: Rioja,1953c:227; Rioja, 1955b:44-49, 52, 53, 61, fig.11-18; Mulaik, 1960:82, 100, 103, 104, 105-110,fig. 527-551; Nicholas, 1962:172; Vanclcl, 1965IJ:502; Vandel, 1965c:354, 362,367-368, fig. L 8;Argano,1972b:31.Protrichoniscus (part): Rioja, 1953e: 11, fig. 7.Brackenridgia villalobosi: Reddell, 1971b:23."~"J/.j 1/ .\',-MEXICOGUATEMALAFig. 10.-Distribution of troglobitic trichoniscid isopods of the genera Brackenridgia, Mexiconiscus, and Typhlotricholigioides:1, Brackenridgia palmitensis; 2, B. bridgesi; 3, Mexiconiscw laevis; 4, B. villalobosi; 5, B. villalobosi and Typhlotricholigioidesaquaticus; 6, B. acostai.96

Type-Iocality.-Cueva del Ojo de Agua Grande,Cordoba, Veracruz, Mexico.Distribution.-Known only from caves near Cordobaand Orizaba, Veracruz. See Fig. 10.Records.-Veracruz: ?Grutas de Atoyac, Cueva delCarbon, and Cueva del Ojo de Agua Grande.Discussion.-This species was collected from organicdebris and from the vicinity of and in small pools.It is most closely related to B. acostai. The record ofB. villalobosi from Grutas de Atoyac is tentative. Mulaik(1960) originally reported the population fromGrutas de Atoyac as Protrichoniscus bridgesi; this iscertainly a misidentification.Cylindroniscus cavicolus (Mulaik)Antroniscus cavicola Mulaik, 1960:82,117,118-119,fig. 50-64; Nicholas, 1962:171; Schultz, 1970a:407-408.Cylindroniscus cavicolus: Schultz, 1970a:409, 411412; Reddell, 1971b:23.Type-Iocality.-Gruta del Palmito, Bustamente("'Bustamante), Nuevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 11.Discussion.-The genus Cylindroniscus is a memberof the First Division of the subfamily Trichoniscinae.MEXICOI//), /'v//GUATEMALA_r-,IIIIII ~.: i'.., _1-""'-- ~......./1--,-,-,--, I~ // ......,,! I'1--" \.._, \'-',I'"\Ii..., I/\,/)-----, LI/~,--/.//('\r'Fig. n.-Distribution of troglobitic trichoniscid isopods of the genus Cylindroniscus: 1, Cylindroniscus cavicolus; 2, C.va/lesensis; 3, C. maya.97

It indudes five spccies, of which thn:e are troglooites.Cylindroniscus seurati .\rcangcli is an endo~can ~pecicsfrolll Cuha: C. yucatanensis C\lulaik) is knownonlY frorn ~anta \laria, Yucatan, and is probablyalso all endogean. C~vlindroniscus cavicolus is apparentlyvcry closely related to C. vallesensis Schultz(Schultz. IlJ70a).Cylindroniscus maya I{iojaCylindronisch us: I{ioja, ] lJS5b: 61 (erroncous spellln~).Cylilldrolliscus maya I{ioja, 1l)58:~69-277, fii!. 1-28;\luliJik, 1%0:B2, lJ2-11(l. ] 18, fii!. 571-Sl.JB;:\icllOlas. 1%2: 172: Schultz, ]970a:407, ·108·WlJ; J{t:ddell, ]l.J710::2:3; I{l:dddl, 1977b::22l.J,240, 2(J8, :2B5.AntrolliscllS balamensis l\lulaik, ]960:B2, 117, IHl,fig. 56-S8; 0iidlOlas, 1%2:171; Schultz, 1970a:W7·40B.Type-Iocality.-Of CylindroniscLls maya: Ct:notede SiJlnlJUlha (=SarnhullOlJ-4J2,fig. 1-1lJ; I{eddell, llJ71 b ::23: Heddell and \Iitchell,LlJ7]a: 143; l\cddcll, ]lJ7:la:32, :37; Rl:ddellJnu Elliott. llJ7:IJ:171, 172.Cylindroniscus 51'. nL vallesensis: I{.eddcll and Elliott,1973a: 172.Type-Iocality.-Cucva Pinta, 8 lIIi :\1

1~yphlotricholigidioidesaqunticus: Vande!, 1%4:332:Vandel, 11)65a:283 (erroneous spelling).Typhlotricholigioides: Vandel. ll)58 :;~..J., fi~. 1: Vandd,I1J60: 13B: Vanckl. IIJ6:ie::l,j1.7:yphlotricholigiodps aquaticus: \r~ano. llJ72b:2:l(nrOlwous spellin~).Type-locality.-Clleva d,·1 (ljo d"-\~lIa (;ran,!c. I'araje\IIl'VO, C'·lfl!ol.a. \·,·ranuz. \I,:"ico.Oistribution.-KnowlI only froln lllP lyp,--Iocalil}.See Fig. 10.Oiscussion.-This sp,~ci,:s was fOllnd in lravl'I'lilll'pools filled wilh walt'r Il'fl from noodin~ fronl d,'('pwilhin the cave. Yand('1 (11)();ih) docs nol !wli,'v,' lhalil ever had lnreslriaJ alll·"slon;. II" eonl"lllb lhal it ison(~ of lh,~ mon' primilive living (lniscoidea and thalit is most c!os,'h f{·lalt:d lo LlIl' trol!Jobilic isopodfrom :'pain, CantabwnisclIs fJrimitivlIs Vandd. II"ciles LlIl' similarily of LI)(' lwo sp,'Cie:.; lo show llll'dfeds of conlinental drift.Order AmphipodaAllhough lIlon' lhan 100 sp"cil's of alnphipod havebeC'n found in LIlt:' can's or lh,· hlil('(l .'lal'·s. lill' orderis poorly repn':i\·nlr·d in \I"·\.ico and C"nlral\nll'rica. (lnly 16 spcci('s of alllphipod ha\(' 11(','n collectedin lhl' cave:.; of lhis f{'~ion. \\ilh lwo ""I"'plionsall are lroglohilic (s"(' Tald,' 10).Family Bo~idiellidac'1'1)(' fa,nil~ 11o~idi"'lidal' is f{'pfl'~,'nl ...J ill llll' cav('Sof \Ic"il'o and (;lIat"1I1ala 11\ nin,' ,Jr.scrilll'd sp('('i('s inthe genlls lJogidiel/a. This ~('nIlS i~ known fronl 1':11rop'-, Isr'lI'l. Central\sia. Japan. \frica. IJrazil. ILirGilda, :'ainl \Iarlin. \I"'"ico. and (;lIal"lIlaJa. \1thongh IIlosl sp,'

Type-locality.-Pozzo, Etla, Oaxaca, Mexico.Distribution.-Known only from the type-locality.See Fig. 12.Discllssion.-This is a phreatic species withoutclear affinities to other described bogidiellids. It wastaken in association with Bogidiella sp. d. arganoi, B.michaelae, and the stenasellid isopod Etlastenasellusmixtecus Argano. A single, incomplete specimen froma well in Tehuacan, Puehla, may also prove to belongto this species (Ruffo and Vigna Taglianti, 1977).Bogidiella orchestipes Ruffo and Vigna TagliantiBogidiella orchestipes Ruffo and Vigna Taglianti,1977:135-141,148,149,150,151,168,169,170,fig. 3-6, 21.Bogidiella sbordonii s. I.: Argano, 1977: 105.Type-locality.-Pozzo in casa bell, S. Cristobal delas Casas, Chiapas, Mexico.Distribution.-Known only from the type-locality.See Fig. 12.MEXICOGUATEMALA......-"/.,/rr·\rFig. 12.-Distribution of troglobitic amphipods of the families Bogidiellidae and Hadziidae: 1, Mexiweckelia mitchelli;2, Mexiweckelia colei and Mexiweckelia particeps; 3, Mayaweckelia yucatanensis; 4, Mayaweckelia cenoticola; 5, Bogidiellaarsonoi; 6, B. arganoi, B. michaelae, and B. niphargoides; 7, B. tabascensis; 8, B. sbordonii; 9, B. orchestipes; 10, B. vomeroi;11, B. pasquinii; 12. B. holsingeri; 13, Bogidiella sp.100

Discussion.-This is a phreatic species most closelyrelated to B. holsingeri.Bogidiella pasquinii Ruffo and Vigna TagliantiBogidiella pasquinii Ruffo and Vigna Taglianti, 1977:141-146, 148, 149, 150, 151, 168, 169, 170, fig.7-9,21; Argano, 1977: 116.Type-locality.-Cueva de los Resadores, S. Eulalia,Huehuetenango, Guatemala.Distribution.-Known only from the type-locali ty.See Fig. 12.Discussion.-This species was taken in associationwith Caecidotea mitchelli and planaria. It is related toB. holsingeri.Bogidiella sbordonii Ruffo and Vigna TagliantiBogidiella sbordonii Ruffo and Vigna Taglianti, 1974:107-111, 11:'>-115, 121, 129, 130-131, 132, fig.2-7,14; Sbordoni et al., 197-1-:29: Holsinger, 1977:15; Ruffo and Vigna Taglianti, 1977:126, 1311:'>4, 141, 16~ J68, 170, 171, fi~2.1,21.Bogidiella: Sbordoni et al., 1977:5-;-: Vigna Taglianti,1977:332.Bogidiella cfr. sbordonii: SLordoni Pi aI., 1977:74,pI. llc.Type-Iocality.-Cueva de Cerro 13rujo, Kancho delCie/ito, Ocozocoautla, Chiapas, .\lexico.Distribution.-Known from three caves in Chiapas.See Fig. 12.Records.-Chiapas: Cueva de Cf'rro 13rujo, Cuevade los Chivos, and Cueva de la Planta n. 3.Discussion.-This is a typical cavernicole speciesapparently most closely related to B. tabascensis. Itwas found in rimstone pools at the type-locality.Bogidiella tabascensis VillalobosBogidiella tabascensis Villalobos, 1960:317-334, fig.1-36; Nicholas, 1962: 172; Dancau and Serban,1965:341; Holsinger and Peck, 1968:250; Holsingerand Minckley, 1971:425; Reddell, 1971b:24; Mitchell and Kawakatsu, 1972: 14; Holsinger,1973:1; Mitchell and Kawakatsu, 1973a:640;Benazzi and Giannini, 1974:52; Ruffo and Vi b '11aTaglianti, 1974: 106-107, 115, 121, 129, 130-131,132, fig. 1, 14; Sbordoni, 1974:366; Sbordoni etal., 1974: 14,15,34; Argano, 1977: 112; Holsinger,1977a:15; Ruffo and Vigna Taglianti, 1977:126,127-131, 134, 141, 168, 170, 171, fig. 1,21.Type-locality.-Grutas del Cocona, 3 km. N.:'IE.de Teapa, Tabasco, Mexico.Distribution.-Known from one cave in Tabascoand two in Chiapas. See Fig. 12.Records.-Chlapas: Cueva de Chital n. 2 and Grutasdel Rancho Nuevo; Tabasco: Grutas del Cocom1.Discussion.-This distinctive troglobite is apparentlymost closely related to B. sbordonii. It hasbeen taken in association with the troglobitic planarianOpisthobursa mexicana at the type-locality. Bogidiellatabascensis has been collected from rimstonepools in Grutas del Rancho Nuevo and from guanorichpools in Grutas del Cocona.BogidielLa vomeroi Ruffo and Vigna TagliantiBogidiella vomeroi Ruffo and Vigna Taglianti, 1977:141, 146-151,152,153,168,169,170, fig. 10-11,21.Bogidiella cfr. vomeroi: Ruffo and Vigna Taglianti,1977:151-153, 16B, fig. 12,21.Type-Iocality.-Cueva de Chanchaniptic, RanchoS. Antonio, Sitala, Chiapas, .\lexico.Distribution.-Known only from the type.localityand possibly one other cave in Chiapas. See Fig. 12.Records.-Chiapas: ?Sumidero de Canada andCueva de Chanchaniptic.Discussion.--This species is similar to B. holsingeri.The specimens from Sumidero de Canada are onlytentatively referred to this species by Ruffo andVigna Taglianti (1977). Typical specimens were takenfrom a stream also inhabited by planaria and the asellidisopod Caecidotea zullinii.Family HadziidaeThe family Hadziidae illustrates very well the distributionalpattern characterized by a concentrationof species in the reri.\lediterranean and Caribbeanregions. The Mexican troglobitic species all belong toa relatively closely related group of genera knownonly from Texas, l\lexico, and the Antilles. Two additionalgenera occur in Europe, while three related generaare known only from Reunion Island, and frommarine waters off California and in the Caroline Islands.Holsinger and Peck (1968), Holsinger andMinckley (1971), Holsinger (1973), and Stock (1977)have discussed the evolu tion and zoogeography of thefamily. A complex of new species and genera fromthe artesian well in Sail ~arcos, Hays County, Texas,is under study by Dr. John R. Holsinger and shouldhelp to elucidate the relationships of the Mexicanspecies.Mayaweckelia cenoticola HolsingerMayaweckelia cenoticola Holsinger, 1977a:19, 21-24,25, fig. 4-7; Reddell, 1977b:230, 240,244,251,252,253,277,281,290,293.New genus (part): Holsinger, 1977b:268.Type-locality.-Cenote XtacaLiha, Yucatan, Mexico.Distribution.-Known from nine caves in the YucatanPeninsula. See Fig. 12.101

Records.-Campeche: Volc

Distribution.-Known only from the type-locality.See Fig. 12.Discussion.-This species was taken in direct associationwith M. colei. Holsinger (in press) now considersthis species to belong to a separate, distinctgenus, more primitive in a number of characters thanMexiweckelia.Family HyaleUidaeThe only species in the family Hyalellidae knownto inhabit caves in Mexico is Hyalella azteca (Saussure).This ubiquitous species ranges through much ofthe United States south to Peru. It has been collectedin cenotes and caves in Yucatan (Creaser, 1936) andfrom caves in Campeche, Chiapas, ;vlichoacan, SanLuis Potosi, and Tamaulipas.Family MelitidaeQuadravisio lutzi (Shoemaker) is the only speciesin the family lVlelitidae known from ;\lexican caves. Itwas collected from brackish water in Cenote de Tulumand Cueva de Tancah, Quintana Roo. Otherwise,it is known only from brackish and freshwater localitiesalong the northwest coastal area of Venezuela,Guayana, and Brazil.Order MysidaceaThe order Mysidacea is represented Il1 subterraneanwaters by at least 17 species. Two of these areknown only from crao burrows, hut the remainingspecies are probable tro/llohites. They are knownfrom Zam:ibar, Italy, India, the Canary Islands, Herzegovina,the An tilles, Costa Rica, Colombia, Peril,and Mexico. All are apparently of marine origin (Vandel,1964). Four species in two families have heencollected from caves in \l(~xico and are all troglobites.Family LepidomysidaeSpelaeomysis olivae BowmanSpelaeomysis oiivae Bowman, 1973:14, 17, fig. 1-11,13-19; Hobbs, 1973L:73; Bowman, 1977a:149150; Hobbs et al., 1977:52,67.Type-Iocality.-Cueva del Nacimiento del Rio SanAntonio, 10 km SSW Acatlan, Oaxaca, Mexico.Distribution.-Known only from the type-locality.See Fig. 13.Discussion.-The genus Spelaeomysis includesseven species, all of which are known from subterraneanhabitats. These have been reviewed by Bowman(1973). Speiaeomysis olivae is most closely related toS. quinterensis. It was collected from clear, siltflooredpools in association with blind shrimps (Aipheopsisstygicoia and J;Jacrobrachium villaiobosi).Spelaeomysis quinterensis (Villalobos)Typhlolepidomysis quinterensis Villalobos, 1951:191-218, fig. 1-14; Rioja, 1953a:290, 291, 293,294; Rioja, 1953e:11, fig. 1; Gordan, 1957:337;Delamare Deboutteville, 1960:644; Gordon, 1960:299; Rioja et al_, 1961:315; Nicholas, 1962:168;Rioja, 1962:40; Vandel, 1964:147; Reddell,1965a:19; Vandel, 1965a:118; Straskraba, 1969:24; Nath and Pillai, 1971:285; Paclt, 1971:424;Sbordoni and Argano, 1972:9; Bowman, 1973:13.Typhlolepidomysis: Martin and Martin, 1954:584;Gordon, 1960:297, 301, 304, 305, 307, 308,309; Pillai and Mariamma, 1964:113; Nath andPillai, 1971:285,286,287.Spelaeomysis quinterensis: Gordon, 1960:308; Pillaiand Mariamma, 1964:113, 122, 123; Bacescu andOrghidan, 1971:225,229,230; Rioja, 1971:503;Bowman, 1973:14, 17, fig. 12, 20; Reddell andElliott, 1973a:I71, 173; Reddell and Elliott,1973b:181, 183; Bowman, 1977a:149-150.Speleomysis quinterensis: Reddell, 1967b:82; Elliott,1971:10; Reddell, 1971b:24; Reddell and Mitchell,1971a:143, fig. 11; Reddell and Mitchell,1971b: 184 (erroneous spelling).Lepidomysis quinterensis: Nath and Pillai, 1971:286,287; ath et al., 1972:53.Blind mysidaceans: Reddell, 1973b:77.Type-Iocality.-Grutas de Quintero, Tamaulipas,\Iexico.Distribution.--Caves in the Sierra de El Abra, SanLuis Potosi and Tamaulipas. and the Sierra de Guatemala,Tamaulipas. See Fig. Vl.Records.-San Luis Potosi: Si>tano de la Tinaja;Tamaulipas: Bee Cave and Grutas de Quintero.New record.-San Luis Potosi: S6tano del Toro(del. T. E. Bowman).Discussion.-In addition to the two troglobiticspecies of the genus listed here, a related species, S.nllniezi Bacescu and Orghidan, is known from cavesin CuLa. This species is certainly more abundant inthe caves of the Sierra Je ~I Abra and Sierra de Guatemalathan the four above records indicate. In Gmtasde Quintero hundreds of inJividuais have beenseen to cluster abou t small pieces of rotten wood inlarge travertine pools fed by a seasonal spring. Onlyone individual has been collected from each of theother caves.Family MysidaeAntromysis (Antromysis) cenotensis CreaserSchizopods: Pearse, 1933:110.Antromysis cenotensis Creaser, 1936:121-123, 131,fig. 13-24; Pearse, 1936c:24; Creaser, 1938:159,103

164; Pearse, 1945:167, 168, fig. 13-24; Anonymous,1947:128; Cardenas Figueroa, 1950:156;Carreno, 1950:24; Tattersall, 1951:4, 229, 230,232, 233, 234, 265; Villalobos, 1951:213, 215;Banner, 1953:3; Rioja, 1953a:286, 290, 293;Gordan, 1957:340; Delamare Deboutteville, 1960:644,648; Gordon, 1960:299; Nicholas, 1962:168;Rioja, 1962:38, 40; Vandel, 1964:147; Vandel,1965a:1l8; Straskraba, 1969:24; Bacescu andOrghidan, 1971:225, 227; Reddell, 1971b:24;Rioja, 1971:503; Bowman, 1973:19; Bowman,1977b:26, 34; Reddell, 1977b:230, 239, 240,251, 252, 253, 255, 256, 257, 258, 260, 261,262, 263, 266, 267, 268, 269, 273, 277, 279,281,283,285,288,289,290,291,293.Antromysis: Creaser, 1938:159; Pearse, 1938a:13,15; Pearse, 1945:111, 167; Cardenas Figueroa,1950:157; Tattersall, 1951:229-230; Argano,1972b:33.Antromysis coenotensis: Bacescu and Orghidan,1971:228 (erroneous spelling).Antromysis (Antromysis) cenotensis: Bowman, 1977b:27-31,34,36,37, fig. 1-4,8.Anthromysis:spelling).Holthuis, 1977:180, 181 (erroneousGUATEMALAFig. l3.-Distribution of troglobitic species of the order Mysidacea: 1, Spelaeomysis quinterensis; 2, S. olivae and Antro·mysis reddelli; 3, A. cenotensis.104

Type-Iocality.-Balaam Canche Cave (=Grotas deBalankanche), three miles east, half mile south ofChichen Itza, Yucatan, Mexico.Distribution.-Known from many caVf~S and wellsin Quintana Roo and Yucatan. See Fig. 13.Records.-Quintana Roo: Actun Ha, Cenote deJuan Coh, Cenote de Las Ruinas, Cenote de San Martin,and Cenote de Santo Domingo; Yucatan: Well(Hacienda Calcehtok), Well (iron mill at Oxkutzcab),Cenote de las Abejas, Cenote Aka Chen, Grotas deBalankanche, Cenote Calchuhuim, Cenote Calchum,ActUn Chac, Cueva Chac Mol, Cenote Chen Mul, Cenotede la Culebra, Cenote G, Cenote de Hoctun,Cenote de Kankirixche, Actun Kaua, Cueva Luchil,Cenote Nohchen, Cenote de Orizaba, Cueva de Orizaba,Cueva Oxolodt, Cenote del Pochote, Cenote deSambula (Motul), Cenote de San Diego, Cueva de SanIsidro, Cenote de San Jose, Pozo de Santa Elena, Cenotede Sihunchen, Cenote de Sodzil, Grotas de TzabNah, Cueva Xconsacab, Cenote Xkeken, Cenot Xtacabiha,Cenote Yunchen, and Cenote Zaci.Discussion.-This species is most closely related tothe troglobitic A. cubanica Hacescu and Orghidanfrom Cuba and A. peckorum Bowman from Jamaica.Antromysis cenotensis is extremely abundant in almostevery cave containing groundwater in the coastalplain of Yucatilll.Antromysis (Antromysis) reddelli BowmanAntromysis (Antromysis) reddelli Bowman, 1977b:34,35,36,37, fig. 7-8.Type-Iocality.-Cueva de las Maravillas, 6 km SAcatlan, Oaxaca, Mexico.Distribution.-Known only from the type-locality.See Fig. 13.Discussion.-This species was taken from a rockand silt-floored stream passage in which also werefound the blind shrimp Alpheopsis stygicola, theblind crayfish Procambarus oaxacae reddelli, andundescribed troglobitic catfish of the genus Rhamdia.This species is most closely related to A. peckorum,A. cubanica, and A. cenotensis.Order DecapodaThe order Decapoda is well represented III thecaves of Mexico by both troglophiles and troglobites.A total of 33 species of shrimp, crayfish, and crabhave been found in Mexican and Guatemalan caves(see Table 12). Many more species probably awaitdiscovery, especially in the caves of Belize and Guatemala.Table 12.-Summary of cave inhabiting Decapoda.TroglobitesOther SpeciesNatantiaAlpheidae 1 0Atyidae 3 0Palaemonidae 6 3ReptantiaCambaridae 3 9Grapsidae 0 1Pseudothelphusidae 3 4Triehodaetylidae 1 1Total 17 18Suborder NatantiaThe caves of Mexico are particularly rich in troglobiticshrimps with ten described species. This is insharp contrast with the United States where only fourtroglobitic shrimps have been described.Family AlpheidaeAlpheopsis stygicola HobbsAlpheopsis stygicola Hobbs, 1973b:73, 76-77, fig.1-2; Hobbs et al., 1977:6, 8, 11, 12, 16, 24, 52,67,68,69, fig. 31-32; Holthuis, 1977:174.Alpheopsis: Hobbs et al., 1977:5,66-67.Type-Iocality.-Cueva del Nacimiento del Rio SanAntonio, 10 km SSW Acatlan, Oaxaca, lexico.Distribution.-Known only from three caves nearAcatlan, Oaxaca, See Fig. 14.New records.-Oaxaca: Cueva de la Finca andCueva de Las Maravillas.Discussion.-This species is of special interest inbeing the only known subterranean member of thefamily and only the third species of the family to bereported from fresh water. The other freshwater speciesare known from a lake in the Ogooue Basin inGabon and from a stream in the vicinity of Manoka,Cameroon. The genus Alpheopsis is known from marinehabitats throughout much of the world, includingAntigua Island and Puerto Rico. The relationshipsof this species to other members of the genusare obscure and probably cannot be clarified until acomparative study of many poorly described speciesis made (Hobbs, 1973b). The species is unquestionablya marine relict. It is an inhabitant of lakes andpools in the caves in which it has been collected. Onespecimen was disgorged by an undescribed blind catfishof the genus Rhamdia when the fish was preserved.105

Family AtyidaeThe family Atyidae is entirely freshwater with theexception of two species of Typhlatya which inhabitmarine or brackish water on Ascension Island andIsla de Santa Cruz, Galapagos Islands (Chace andManning, 1972). Troglobitic species of atyid areknown from Europe, Japan, Africa, Madagascar,Australia, Fiji, the United States, Cuba, Puerto Rico,and Mexico.Typhlatya campecheae Hobbs and HobbsTyphlatya campecheae Hobbs and Hobbs, 1976:6-10,14, 15, 16, 20, 21, fig. 3-4; Hobbs et al., 1977:6,8, 11, 12, 24, 32, 33, fig. 7-8; Holthuis, 1977:174; Reddell, 1977b:230, 240, 241, 243, 249;Hobbs, 1979:618, 619-620, 621, 624, 625, 626,629, fig. 1.Typhlatya (part): Hobbs et al., 1977:5, 13, 32.Type-Iocality.-Grutas de Xtacumbilxunam, Bolonchenticul,Campeche, Mexico.Distribution.-Known only from two caves inCampeche. See Fig. 14.Records.-Campeche: Cenote de Cantemo andGrutas de Xtacumbilxunam.Discussion.-This species was found to be extremelyabundant in guano-floored pools at the type-MEXICOI,lGUATEMALAFig. 14.-Distribution of troglobitic shrimps of the families Alphaeidae and Atyidae: 1, Alpheopsis stygicola; 2, Typhlatyacampecheae; 3, T. pearsei; 4, T. mitchelli; 5, T. pearsei and T. mitchelli.106

locality and in association with ostracods and amphipods(Mayaweckelia yucatanensis). They were observedto rest on the floor of the pool and when disturbedwould swim straight up or away at an angle.In Cenote de Cantemo they inhabited a small poolcontaining much organic debris and polluted by oil.Typhlatya campecheae is most closely related to T.pearsei.Typhlatya mitchelli Hobbs and HobbsTyphlatya mitchelli Hobbs and Hobbs, 1976:2-6, 10,14, 15, 16,20,21, fig. 1-2; Hobbs et al., 1977:6,8, 11, 12, 25, 32, 39, 40, 41, 45, fig. 12-13; Holthuis,1977:174, 175-181, fig. la-i, 2; Reddell,1977b:230, 239, 240, 251, 255, 256, 260, 262,263, 268, 279, 281, 289, 291,292,293; Hobbs,1979:618,620-625,626,629, fig. 1-2.Typhlatya (part): Hobbs et al., 1977:5, 13,32.Type-iocality.-Cenote Kabahchen, Mani, Yucatan,Mexico.Distribution.-Known from caves in Quintana Rooand Yucatan. See Fig. 14.Records.-Quintana Roo: Cueva del Fermin andActun Ha; Yucatan: Cenote Aka Chen, Cenote Calchuhuim,Cenote Chen Mul, Cenote Chun Kapoc,Cenote de la Culebra, Cenote Kabahchen, ActunKaua, Cenote de Orizaba, Cenote de la Paca, Cenotedel Pochote, Cenote de Sodzil, Grutas de Tzab-Nah,Cenote Xkeken, Cenote Xtacabiha, and Cenote Zaci.Discussion.-This species is s)'mpatric with T.pearsei in several caves. It possesses some characterswhich make it intermediate between the Cuban T.garciai Chace and the Puerto Rican T. monae Chaceand the Galapagos Islands T. galapagensis Monod andCals. In most of the above caves it was taken fromsmall pools.Typhlatya pearsei CreaserBlind shrimps (part): Pearse, 1933:110.Typhlatya pearsei Creaser, 1936: 128-130, 131, fig.31-41; Pearse, 1936c:24; Creaser, 1938:162-163,164; Chace, 1942:100; Chace, 1943:30, 32;Pearse, 1945:169, 170, fig. 31-41; Cardenas Figueroa,1950:156; Villalobos, 1951:215; Rioja,1953a:286, 292; Chace, 1954:319, 323; Balss,1955:1310; Holthuis, 1955:26; Holthuis, 1956:52; Nicholas, 1962:173; Chace and Hobbs, 1969:20-21; Straskraba, 1969 :25; Botosaneanu andHolthuis, 1970:122, 123, 127; Monod and Cals,1970:69, 73, 78, 82, 84, 85, 93, 94, 100, fig. 67;Reddell, 1971b:25; Rioja, 1971:524; Chace, 1972:15; Chace and Manning, 1972:17; Croizat et al.,1974:275, fig. 2; Holthuis, 1974:141; Silva Taboada,1974:54; Monod, 1975:99, fig. 1; Hobbs andHobbs, 1976:1, 10-14, 15, 16, 20, 21, fig. 5-6;Hobbs et al., 1977:6, 7, 9, 12, 15,24,32,39,4345, fig. 3, 15-16; Holthuis, 1977:174-175, 176,180, 181, fig. Ij; Reddell, 1977b:230, 239, 240,241, 243, 247, 251, 252, 253, 258, 260, 262,266,277,278,281,283,285,293;Hobbs,1979:618,621, 624, 625~27,628, 629,630,631,fi~1, 3A, 3D.Typhlatya: Hubbs, 1936:168; Creaser, 1938:159;Pearse, 1938a:13, 15; Pearse, 1945:111, 167;Argano,1972b:33.Typhlata: Cardenas Figueroa, 1950: 157 (erroneousspelling).Typhlatya pearsi: Rioja, 1953a:293; Delamare Deboutteville,1960:648; Vandel, 1964:178; Vandel,1965a:139; Peck, 1974a:21; Juberthie-Jupeau,1977:108 (erroneous spelling).Typhlatya pearsii: Rioja, 1971:1150 (erroneous spelling).Typhlatya (part): Croizat et al., 1974:276; Juberthie,1974:81; Monod and Cals, 1970:69,72,73,85,87,91,97,99, 101, 102; Rosen, 1976:441, fig. 5E.Type-locality.-Balam Canche Cave (=Grutas deBalankanche), 4.8 kilometers east, 0.8 kilometerssouth Chichen ltza, Yucatan, Mexico.Distribution.-Known from caves in Campeche,Quintana Roo, and Yucatan. See Fig. 14.Records.-Campeche: Grutas de Monte Bravo andGrutas de San Antonio; Quintana Roo: Cueva Coop,Cueva del Fermin, Cenote de Juan Coh, Pozo de SanMartin, Cenote de Santo Domingo, and Cenote deTos Vidol; Yucatan: Cenote de las Abejas, Grutas deBalankanche, Cenote Calchum, Actun Chac, Cenotede Hoctun, Cenote Kabahchen, ActUn Kaua, ActunOkobichen, Cenote del Pochote, Cueva de San Isidro,Cueva de Santa Elena, Grutas de Tzab-Nah, and CenoteXtacabiha.Discussion.-The genus Typhlatya is known fromAscension Island, the Galapagos Islands, Cuba, DominicanRepublic, Puerto Rico, and the Yucatan Peninsulaof Mexico. This species is most closely related toT. mitchelli. The record by Nicholas (1962) of thisspecies in "Cueva del Ponte" is probably an error forCenote del Pochote.Family PalaemonidaeSix species of palaemonid shrimp are troglobites inthe caves of Mexico. An additional three species havebeen found in caves as either troglophiles or trogloxenes.Other troglobitic palaemonids are known fromAssam, the Mediterranean region, the southern UnitedStates, and Cuba.107

Creaseria morleyi (Creaser)Blind shrimps (part): Pearse, 1933: 110.Palaemon morleyi Creaser, 1936:126-128, 131, fig.25-30; Pearse, 1936c:24; Creaser, 1938:163-164;Chace, 1943:31,33; Pearse, 1945:169, fig. 25-30;Anonymous, 1947:128; Cardenas Figueroa, 1950:156; Carreno, 1950:24; Villalobos, 1951:215;Rioja, 1953a:29:3, 294; Balss, 1955:1:Hl; Holthuis,1955:44; Hobbs and Hobbs, 1976:1; Hobbset aI., 1977:7,9; Hobbs, 1979:618.Palaemon: Pearse, 1938a: 13, 15; Pearse, 1945:111,167; Cardenas Figueroa, 1950:157.Creaseria morleyi: Holthuis, 1950:6 (by implication);Holthuis, 1952:153-154, 356-357, pI. 40; Chace,1954:323; Holthuis, 1955:44 (by implication),fig. 22a; Holthuis, 1956:56-57; i\laccagno and Cucchiari,1957:207 (by implication); Nicholas, 1962:174; Rioja, 1962:38,40; Vandel, 1965:179; Rodriguezde la Cruz, 1965:76, 97-98, pI. 4(fig. A);Vandel, 1965a:140; Andrews, 1970:4; Reddell,1971b:25; Rioja, 1971:522; Parzefall and Wilkens,1972:66; Wilkens, 1973a:327, 328, 330, fig. 3;Wilkens, 1973b:205; Wilkens, 1973c:50-54, 56,58, 59, fig. 1-3; Burukovskii, 1974:fig. 96a; Holthuis,1974:141; Hobbs and Hobbs, 1976:1, 1619, 20, 21, fig. 7-8; Strenth, 1976:13; Hobbs etal., 1977:6,9, 12,25,46-50, fig. 18-19; Holthuis,1977:186-187, fig. 5; Reddell, 1977b:230, 240,241,243,247,251,252,253,255,258,262,263,264,266,267-268,273,274,277,281,283,285,291, 292, 293; Hobbs, 1979:618,619,621,624,625,627-632, fig. 1, 4B-C.Creaseria: Holthuis, 1952:2, 152-153; Holthuis,1956:69; Balss, 1957:1549; Delamare Deboutteville,1960:646; Argano, 1972b:33; Chace, 1972:17; Hobbs et aI., 1977:5, 14, 16,46.Palaemon morley: Rioja, 1953a:286 (erroneous spelling).Creaseria morlayi: Delamare Deboutteville, 1960:648(erroneous spelling).?Shrimp: Lee, 1966:88.Creaseria morelyi: Hobbs and Hobbs, 1976:1 (erroneousspelling).Type-locality.-San Isidro Cave (=Cueva de San Isidro),Salar Colony, Yucatan, Mexico.Distribution.-Known from many caves in Campeche,Quintana Roo, and Yucatan. See Fig. 15.Records.-Campeche: Cenote Bolchen, Grutas deMonte Bravo, and Grutas de San Antonio; QuintanaRoo: Cueva Coop, Cueva del Fermin, Cenote de JuanCoh, Cenote de Las Ruinas, Pozo de San Martin, Cenotede Santo Domingo, and Cenote de Tos Virlol;Yucatan: Cenote de las Abejas, Cueva Amil, Grutasde Balankanche, Cenote Calchuhuim, ActUn Chac,Cueva Chac Mol, Cenote de la Culebra, ActUn Gongora,Cenote de HoctUn, Cenote Kabahchen, Cenotede la Paca, Cenote del Pochote, Cenote de Sambula(Motul), Cueva de San Isidro, Pozo de Santa Elena,Cueva de Sodzil, Grutas de Tzab-Nah, ?Cueva Xconsacab,Cenote X-ebiz, ActUn Xpukil, and CenoteYunchen.Discussion.-This large, distinctive species is anever-present element of the fauna of pools and lakesin caves in the Yucatan Peninsula. It is frequentlyfound in large numbers, especially crawling about thefloors of pools over which bats roost. Many individualsare seen which have lost one cheliped, presumablyin fights with other members of the same species.Wilkens (1973c) in a study of the structure of the eyeremnants in this species reports that the eye stalkconsists only of medulla interna and medulla terminalis.The record for Cueva Xconsacab was a sightrecord only and should be verified by specimens.iHacrobrachium acherontium HolthuisMacrobrachium acherontium Holthuis, 1977:174,188-193,194, fig. 6-7.Macrobrachium new species: Hobbs et al., 1977:5.Type-locality.-Grutas del Cocona, 3 km NE ofTeapa, Tabasco, Mexico.Distribution.-Known only from two caves nearTeapa, Tabasco. See Fig. 15.Records.-Tabasco: Grutas del Cocona and Resumiderodel Cocona.Discussion.-This species is most closely relatedto M. villalobosi. The cornea is very reduced and theanimals are transparent brownish. This species ispresumably a rather recent troglobite. In Grutas delCocona it primarily inhabits a large, shallow pond.The animals usually rest quietly or walk slowly abouton the silt floor. In addition to the two troglobitesdiscussed here, two other species are known fromcaves in Mexico and Guatemala. The river prawn,IvI. carcinus (Linnaeus), has been collected in CuevaChica, San Luis Potosi, and Cueva del Saito de Agua,Chiapas. In hath instances this species has enteredfrom rivers directly associated with the caves. Macrobrachiumacanthurus (Wiegmann) has heen collectedin Cueva de la Coche, Izahal, Guatemala, and inCueva de Abispa, Quintana Roo, Mexico.Macrobrachium villalobosi HobbsMacrobrachium villalobosi Hobbs, 1973b:77, 79-80,fig. 3; Strenth, 1976:1:3; Hobbs et al., 1977:6,8,11, 12, 16, 22, 23, 25, 50-52, 67, 118, fig. 20;Holthuis, 1977:188, 192-193.109

Shrimp: Reddell, 1973d:89.Macrobrachium (part): Hobbs et al., 1977:5, 13, 15,16,50.Type-Iocality.-Cueva del Nacimiento del Rio SanAntonio, 10 km SSW Acathln, Oaxaca, Mexico.Distribution.-Known only from two caves nearAcatlan. Oaxaca. See Fig. 15.New record.-Oaxaca: Cueva de la Finca (det. H.H. Hobbs, Jr.).Discussion.-This species is most closely related toM. acherontium. It was taken from pools in associationwith Spelaeomysis olivae, Alpheopsis stygicola,Procambarus (Austrocambarus) oaxacae reddelli, andan undescribed blind catfish of the genus Rhamdia.Neopalaemon nahuatlus Hobbs.Veopalaemon nahuatlus Hobbs, 1973a:25, 26-27, 29,fig. 1-2; Holthuis, 1974: 142; Hobbs et al., 1977:6,8, 11, 12,25,52,53,118, fig. 21; Holthuis, 1977:181.Large blind shrimp: Reddell, 1973d:90.Neopalaemon: Hobbs et aI., 1977:5, l-t, 16,52.Type-Iocality.-Cueva del Guano, 10 km NE Valle, acional, Oaxaca, l\lexico.Distribution.-Known only from the type-locality.See Fig. 15.Discussion.-This remarkable troglobite is the onlyknown species in the genus Neopalaemon. Neopalaemonis possibly most closely related to Macrobrachium.The eye pigment and cornea are absent in thisspecies. These large shrimps were found in abundancein a deep permanent stream which runs beneath alarge bat colony. They have been found in associationwith Procambarus (Austrocambarus) oaxacae oaxacae.Troglocubanus perezfarfanteae VillalobosTroglocubanus sp.: Reddell, 1967d:82; Reddell,1971b:26; Reddell and Mitchell, 1971a:143-144;Reddell and Elliott, 1973a: 171.Troglocubanus: Reddell, 1973a:32.Troglocubanus, undescribed species: Holthuis, 1974:142.Troglocubanus perezfarfanteae Villalobos, 1974:1-5,fig. 1-23; Strenth, 1976:13; Hobbs et al., 1977:6,8, 11, 25, 64-66, fig. 30; Holthuis, 1977:181;Mitchell et aI., 1977:49.Troglocubanus (part): Hobbs et al., 1977:5,14,58.Type-Iocality.-Sotano de Tinaja, 11.7 km E.NE.Valles, San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 15.Discussion.-This remarkably delicate species hasextremely elongate appendages. The only known specimenwas obtained from near the surface of a deeplake in the deepest part of Sotano de la Tinaja. Itsextreme transparency has probably aided it in escapingnotice during the ex tensive explorations ofcaves in the Sierra de EI Abra. The genus Troglocubanusis otherwise known only from caves in Cuba.This species, therefore, is another of a series ofgroups of marine relicts shared both by the Mexicanmainland and the greater Antilles.Suborder ReptantiaSection MaeruraFamily CambaridaeAlthough crayfishes are frequently found in Mexicancaves and a total of 12 species are recorded, onlythree closely related forms have become troglobitic(see Fig. 16). This is a surprisingly low figure when itis compared with the crayfish fauna of the easternUnited States where more than 20 species and subspecieshave been described as troglobites (Hobbs andBarr, 1972). In addition to the three troglobites listedand discussed below, the following crayfish species,all presumably troglophiles, are known from caves inMexico. Procambarus (Austrocambarus) mirandaiVillalobos inhabits a long, meandering stream passagein Cueva Cerro Hueco, Chiapas (Villalobos, 1954); ithas also been found in Cueva del Arco and Cueva dePaso Burro, Chiapas (Hobbs, 1977). Procambarus (A.)pilosimanus (Ortmann) is a widely distributed speciesin southern Mexico; it is known from a pool in Grutasde Zapaluta, Chiapas, which connects directly with asurface pool (Villalobos, 1955), and from two cavesin Alta Verapaz, Guatemala (Delamare Debouttevilleand Juberthie, 1976). Procambarus (A.) sbordoniiHobbs was described from Cueva del Nacimiooto delRio Santo Domingo, Chiapas (Hobbs, 1977). Procambarus(Ortmannicus) acutus cuevachicae (Hobbs)is found in abundance in Cueva Chica, San Luis Potosi(Hobbs, 1941); it is a widely distributed formthroughout east-central Mexico, but it is not definitelyknown from other caves in the Sierra de EIAbra. Procambarus (0.) toltecae Hobbs is seldom collectedon the surface but has been taken from cavesin the Xilitla and Aquismon regions of San Luis Potosiand fmm Cueva de EI Tenango, Hidalgo. InCueva de EI Tenango it was abundant in the longstream passage which runs through the cave. Procambarus(0.) villalobosi Hobbs is known only fromCueva del Agua, near Rayon, San Luis Potosi (Hobbs,1969b), and from Pozita de Ojo de Agua, San LuisPotosi. This species inhabits both the cave stream anduo

the pond which is fed by the cave stream in Cueva delAgua. Procambarus (Villalobosus) xochitlanae Hobbsis known from Cueva de los Camarones, Puebla(Hobbs, 1975). A new species ofProcambarus (Villalobosus)from caves in the Cuetzalan region of Pueblaawaits description. Procambarus (Paracambarus) ortmanni(Villalobos) was recently collected from astream in Grutas de Olivares, Puebla.Procambarus (Austrocambarus) oaxacae oaxacaeHobbsProcambarus (A ustrocambarus) oaxacae oaxacaeHobbs, 1973a:29, 32-33,38, fig. 3-5; Hobbs et al.,1977:6, 8, 11, 12, 17,26,52,116-118, 152, fig.52.Blind crayfish: Reddell, 1973d:90.GUATEMALAFig. 16.-Distribution of troglobitic and troglophilic decapods of the families Cambaridae, Grapsidae, Pseudothelphusidae,and Trichodactylidae: 1, Procambarus (Ortmannicus) villalobosi; 2, Procambarus (0.) acutus cuevachicae; 3, Procambarus (0.)toltecae; 4, Procambarus (Villalobosus) xochitlanae; 5, Procambarus (V.) xochitlanae and Procambarus (Paracambarus) ortmanni;6, Procambarus (Austrocambarus) rodriguezi and Pseudothelphusa (Tehuana) cordobensis; 7, Procambarus (A.) oaxacaereddelli; 8, Procambarus (A.) oaxacae oaxacae; 9, Trichodactylus (Rodriguezia) bidens; 10, Trichodactylus (R.) mensabak;11, Procambarus (A.) sbordonii; 12, Procambarus (A.) mirandai; 13, Procambarus (A.) pilosimanus; 14, Typhlopseudothelphusamocinoi; 15, Typhlopseudothelphusa mitchelli; 16, Typhlopseudothelphusa juberthiei; 17, lsabellagordonia (Phrygiopilus)longipes; 18,1. (P.) acanthophallus; 19, Sesarma (Holometopus) roberti.111

Type-Iocality.-Cueva del Guano, 10 km NE ValleNacional. Oaxaca, Mexico.Distribution.-Known only from the type-locality.See Fig. 16.Discussion.-This attractive crayfish has the eyesgreatly reduced. It was found in association with Neopalaemonnahuatlus. Procambarus oaxacae is consideredto be most closely related to P. rodrigueziHobbs and may even prove to be only subspecificallydistinct when collections from intermediate areas aremade. The crayfish from this cave are used frelluentlyby local inhabitants of the area for food. It is extremelyeasy to catch, but fortunately much of thestream it inhabits is quite deep and other parts ofthe stream are very difficult of access because of lowclearance between the cave ceiling and the water. Thestream contains much guano, an important factor inexplaining the large population of both species ofdecapod in this cave.Procambarus (A ustrocambarus) oaxacae reddelliHobbsProcambarus (Austrocambarus) oaxacae reddelliHobbs, 1973a:33, 35,37-38, fig. 6·8; Hobbs et al.,1977:6,8,11,12,17,26,52,67,118,119,152,fig. 53.Troglobitic crayfish: Reddell, 1973d:90.Procambarus oaxacae reddelli: Hobbs, 1973b:73.Type-Iocality.-Cueva del Nacimiento del Rio SanAntonio, 10 km SSW Acatlan, Oaxaca, Mexico.Distribution.-Known from four caves in Oaxacaand one in Veracruz. See Fig. 16.Records.-Oaxaca: ?Cueva del Guayabo and Cuevadel Nacimiento del Rio San Antonio.New records.-Oaxaca: Cueva de la Finca andCueva de Las Maravillas; Veracruz: Cueva de Corralde Piedra (det. H. H. Hobbs, Jr.).Discussion.-Specimens trom Cueva del Guayaboare only tentatively assigned to this SUbspecies. Theyshow some in termediate characters between P. o.oaxacae and P. o. reddelli. The crayfish in Cueva delGuayabo were collected from small pools apparentlyleft from a rise in a postulated lower stream. In Cuevadel Nacimiento del Rio San Antonio crayfish wereextremely abundant throughout the length of thislarge cave. They tended to congregate especiallyabout small areas of guano left by bats which roostedin small groups on the ceiling. The crayfish in thiscave are also used for food by local inhabitants. InCueva de Corral de Piedra crayfish were found onlyin a side passage which led to an apparent sealed entranceinto which organic debris had washed.Procambarus (Austrocambarus) rodriguezi HobbsProcambarus rodriguezi Hobbs, 1943: 198, 203-206,fig. 15-26; Villalobos, 1948:182; Rioja, 1953a:288, 289, 292; Villalobos, 1953:346, 356, 364,365, 372; Villalobos, 1954:303, 306, 321-323,363; Villalobos, 1955:2, 20, 160, 161, 176-178,218; Nicholas, 1962:173; Hobbs and Villalobos,1964:313; Hobbs, 1966:71; Hobbs, 1969a:161;Hobbs, 1971:3, 12, 30-31; Reddell, 1971a:~17,219; Reddell, 1971b:25; Rioja, 1971:530; Hobbsand Barr, 1972:1, 4; Hart and Hart, 1974:22;lIohLJs et al., 1977:7,9.Paracambarus rodriguezi: Rioja, 1953a:293 (erroneOilS combination).Procambarus (AustrC'cambarus) rodriguezi: Hobbs,IY72a:-Hl, 152, fig. 22c; Hobbs, 1973a:25, 38;lI(Jbh~, 11)74:45,123, fig. 174; Hobbs, 1975:276;IIl)bLJ~ et al., 1977:6, 12, 17, 26. 118, 120-121,152, fig. 54.Procambarus (Austrocambarus) rodiguezi: Hobbs,1972b:6 (erroneous spelling).Troglobitic procambarid: Caine, 11)74:488.Type-Iocality.-Cave, 4 kilometers west northwestof the hacienda at Potrero Viejo, Paraje Nuevo, Veracruz,Mexico.Distribution.-Known definitely only from thetype-locality. See Fig. 16.Records.-Veracruz: Cave 4 km WNW of PotreroViejo and ?Cueva del Ojo de Agua Grande.Discussion.-The type·locality of this species hasbeen stated to be "Cueva del Agua" and "Cueva delOjo de Agua Grande," but the brief description ofthe type-locality as given by Hobbs (1943) does notfit Cueva del Ojo de Agua Grande. Furthermore,Cueva del Ojo de Agua Grande has no habitat suitablefor crayfish. It contains only a single passage and thiscontains a rapid rock-floored stream. A single juvenilecrayfish was seen trapped in a tiny drip pool in thelatter cave where flood water apparently isolated it.It must be assumed that the type-locality has notbeen rediscovered.Section BrachyuraFamily GrapsidaeThe only species of grapsid crab known from thecaves of Central America is Sesarma (Holometopus)roberti H. Milne Edwards. It was recently collected inCueva de la Coche, Izabal, Guatemala.Family PseudothelphusidaeIn addition to the three remarkable troglobiticcrabs of the genus Typhlopseudothelphusa, fourpseudothelphusid crabs are known as apparent troglo-112

philes. Pseudothelphusa (Pseudothelphusa) sonoraeRodriguez and Smalley was described from a streamin Mina La Aduana, Sonora (Rodriguez and Smalley,1972); P. (Tehuana) cordobensis Rodriguez andSmalley is known only from Cueva de Ojo de AguaGrande, Veracruz (Rodriguez and Smalley, 1972);Isabellagordonia (Phrygiopilus) acanthophallus (Smalley)is known from Cueva Seamay, Alta Verapaz,Guatemala (Smalley, 1970); and 1. (P.) 10ngipesPretzmann is known only from Grutas de Lanquin,Alta Verapaz, Guatemala (Pretzmann, 1972).Typhlopseudothelphusa juberthiei DelamareDebouttevilleTyphlopseudothelphusa juberthiei Delamare Deboutteville,1976:837, 838, 839, fig. G-1; DelamareDebou tteville and Juberthie, 1976: 19:Delamare Deboutteville, 1977:115, 119, 122, 123,125-126,127,128, fig. 6-7,14-15.Typhlopseudotelphusa juberthiei: Cottarelli and\rgano,1977:212 (erroneous spelling).Type-Iocality.-Grotte de Chicam, Sierra de Chama,Alta Verapaz, Guatemala.Distribution.-Known only from the type-locality.See Fig. 16.Discussion.-Delamare Deboutteville (1977) hasdiscussed at leng:th the habitat and relationships ofthis and the following species.Typhlopseudothelphusa mitchelli DelamareDebou ttevilleTyphlopseudothelphusa mitchelli Delamare Deboutteville,1976:8:17, 838, 8:39, fig. D-F; DelamareDeboutteville and J uberthie, 1976:23: DelamareDeboutteville, 1977:115, 119, 121, 124,125,126,127,128,129, fig. 5,9-1:3.Typhlopseudotelphusa mitchelli: Cottarelli and Argano,1977:212 (erroneous spelling).Type-Iocality.-Grottes C3 et (;3, sierra de Pampurau Guatemala, Alta Verapaz, Guatemala.Distribution.-Known only from the two caveslisted as the type-locality. See Fig. 16.Discussion.-lt is not known which of the twocaves listed above is that from which the holotypewas taken. Delamare Deboutteville (1977) reportedthat a specimen of T. mitchelli collected on 29 April1976 carried 48 egb'S.Typhlopseudothelphusa mocinoi RiojaTyphlopseudothelphusa n. g., n. sp.: Rioja, 1953a:291,292,293 (nomen nudum).Typhlopseudothelphusa mociiloi Rioja, 1953b:218225, fig. 1-12; Rioja, 1971:548; Hobbs et al.,1977:7, 10.Cangrejo ciego: Rioja, 1953e:11, fig. 2.Typhloseudothelphusa mocifioi: Rioja et al., 1961:313; Hartnoll, 1964b:78 (erroneous spelling).Typhlopseudothelphusa mocinoi: Nicholas, 1962:174; Hartnoll, 1964a:I64; Hartn011, 1964b:79;Holthuis, 1964:65; Thines and Tercafs, 1972:68;Delamare Deboutteville, 1976:837,838, 839, fig.A-C; Delamare Deboutteville and Juberthie, 1976:23, fig. 3; Delamare Deboutteville, 1977:115, 116,119,120,124,126, fig. 1,4.Typhlopseudotelphusa mocinoi: Vandel, 1964: 183;Vandel, 1965a:142; Delamare Deboutteville,1971:60; Cottarelli and Argano, 1977:212 (erroneousspelling).Potamocarcinus (Typhlopseudothelphusa) mocinoi:Pretzmann, 1965:2 (by implication); Smalley,1970:102; Reddell, 1971a:217, 219; Reddell,1971b:26; Reddell and Mitchell, 1971c:2; Pretzmann,1972:7, 64 (by implication), 81, fig. 19,730-732; Sbordoni et al., 1974:9; Hobbs et al.,1977:6, 12,28, 144-145, fig. 67.?Cave crab: Barr, 1968a:82.Potamocarcinus (Typhlopseudothelphusa): Smalley,1970:103,105.Potamocarcinus (Typhlopseudothelphusa) mocifioi:Rodriguez and Smalley, 1972:72, 92, fig. 3, 25,26.Potamocarcinus (Typhlopseudotelphusa) mocifioi:Rodriguez and Smalley, 1972:92 (erroneousspelling).Potamocarcinus (Typhlopseudotehlphusa) mocifioi:Rodriguez and Smalley, 1972:92 (erroneous spelling).Potamocarcinus mocinoi: Sbordoni et al., 1974:26.Typhlopseudothelphusa: Hobbs et al., 1977:143-144.Potamocarcinus (part): Hobbs et al., 1977:5, 142-143.Type-Iocality.-Cueva del Tio Ticho, Comitan,Chiapas, l\lexico.Distribution.-Known only from three caves inChiapas. See Fig. 16.Records.-Chiapas: Cueva de Los Llanos, Cueva delos MurcieIagos, and Cueva del Tio Ticho.Discussion.-1n Cueva del Tio Ticho this species inhabitsthe lowest level of the cave. A female with 75eggs and another with 57 young were collected on21 August 1967 on a mud slope above the cavestream. In Cueva de Los Llanos and Cueva de los MurcieIagosthe crabs inhabited a small stream passageover which bats roosted, and in association with blindasellid isopods (Caecidotea chiapas) and planarians(Dugesia mckenziei).113

Family TrichodactylidaeTwo species of the family Trichodactylidae havebeen reported from Mexican caves. Bott (1969) describedTrichodactylus (Rodriguezia) bidens fromCueva del Azufre, Tabasco. This is an eyed speciesand presumably a troglophile. The other species is atroglobite and is discussed below.Trichodactylus (Rodriguezia) mensabak Cottarelliand ArganoTrichodactylus (Rodriguezia) mensabak Cottarelliand Argano, 1977:207-212, fig. 1·2; Sbordoni etaI., 1977:74, pI. lIB (nomen nudum).Trichodactylus mensabak: Sbordoni et al., 1977:37(nomen nudum).Type-Iocality.-Cueva de Nicolas Bravo, 'fila, Chiapas,Mexico.Distribution.-Known only from the type-locality_See Fig. 16.Discussion.-This is the only known troglobiticcrab in the family Trichodactylidae. It is most closelyrelated to T. (R.) vil/alobosi Rodriguez and Manrique,an epigean species from Chiapas.Class ArachnidaThe class Arachnida has been the most successfulof all groups which have invaded the cave habitat inMexico and Central America. A total of 122 speciesof troglobite is known from Mexi,;o, Guatemala, andBelize, and another 420 species have been iden tifiedas troglophiles, trogloxenes, or accidentals. Mexico isunique in that all 11 orders of arachnid have been collectedin caves (see Table 13)_ The orders Scorpiones,Pseudoscorpionida, Schizomida, Amblypygida, Araneae,Ricinulei, Opilionida, and Acarina contain troglobites.Order ScorpionesThe order Scorpiones is generally poorly representedin the fauna of caves. The European l1elisaurusxambeui Simon is frequently found in caves but alsoinhabits deep soil and litter. Uroctonus sequoiaGertsch and Soleglad and Vaejovis iviei Gertsch andSoleglad, both from California, and V. reddelliGertsch and Soleglad from Texas are troglophileswhich show little or no adaptation to a cavernicoleexistence. Only eight described species are consideredto be significantly cave adapted, of which seven occurin Mexico. Uroctonus grahami Gertsch and Solegladfrom Samwell Cave, California, is probably a troglobitealthough it retains vestiges of eyes (Gertsch andTable 13.-Summary of cave inhabiting Arachnida.TroglobitesOther SpeciesScorpionesUnknown family 1 0Buthidae 0 2Chactidae 5 1Diplocentridae 3 2Vaejovidae 1 2*Pseudoscorpionida 24 17UropygidaThelyphonidae 0 1SchizomidaeProtoschizomidae 3 0Schizomidae 12 12AmblypygidaCharontidae 1 0Phrynidae 5 7*Araneae 47 252RicinuleiRicinoididae 3 6*Opilionida 15 35PalpigradidaEukoeninlid ae 0 ISolpugida 0 1*Acarina 2 81Total 122 420*-For summary by families see Tables 14-17.Soleglad, 1972). The troglobitic scorpion fauna ofMexico includes three described and two undescribedchactids, three diplocentrids, one vaejovid, and a remarkableeyeless species from Oaxaca of unknownfamilial relationship.Family ButhidaeCentruroides yucatanus (Chamberlin and Ivie) wasdescribed from Actun Loltun, Yucatan (Chamberlinand lvie, 1938a), but is a junior synonym of C. ochraceousPocock, which has been collected from underrocks in the entrance area of two caves and fromdarkness in two other caves in Yucatan (Wagner,1977). Centruroides gracilis (Latreille) has beenfound in Cueva del Salitre, San Luis PotosI, andCueva de los Cuarteles, Tamaulipas. In the latter cavea single large specimen was collected from the ceilingof thc cave in total darkness.Family ChactidaeThe family Chactidae is represented in Mexicancaves by the three troglobites of the genus Typhlochactasdiscussed below, by an undescribed genuswith two new troglobitic species from caves in SanLuis Potosi and Tamaulipas, and by a single speciesof possible troglophile. The last species, Megacormusgertschi Diaz Najera, has been found in two caves inQueretaro.114

Typhlochactas elliotti MitchellTyphlochactas elliotti Mitchell, 1971b:135-148, fig.1-17, 26-27; Reddell and Mitchell, 1971a:144,fig. 1; Reddell, 1973a:33, 37; Reddell and Elliott,1973a:171; Gonzalez Sponga, 1974:56; Soleglad,1976:253,254; Mitchell and Peck, 1978:159, 164,167,168.Troglobitic scorpion: Elliott, 1972:130.Typhlochactas: Vomero, 1974:353 [Sotano de Jerbaniz(= Yer1aniz) record only]; Mitchell et al., 1977:56.Typlochactas (part): Diaz Najera, 1975:3,35 (erroneousspelling).Typlochactas elliotti: Diaz Najera, 1975 :4, 30 (erroneousspelling).Type-locality. -Sotano de Yer1aniz, located approximately21 km north of Ciudad Valles, San LuisPotosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 17.Discussion.-This is the most highly cave adaptedof the three known troglobitic scorpions in the genus.It appears to have its closest affinities with T. rhodesiMitchell, the geographically nearest species (Mitchell,1971b). Mitchell expresses surprise at the discoveryof a highly evolved troglobite in the Sierra de EIAbra, but this is perhaps a logical development in theevolution of the troglobitic Typhlochactas. The recentdiscovery of the eyeless T. sylvestris Mitchelland Peck from forest litter at 1220 meters in Oaxacagives us an important clue to the evolution of thisgroup. It may be assumed that the ancestral blindscorpions originally in habited much of Mexico asinhabitants of deep forest litter. With the rise oftemperature in the lowland tropics following thecolder climate of the glacials, they survived in cavesat low elevations and the surface populations wereeliminated. The surface populations still existed inthe cooler forested highlands for longer periods oftime, but eventually they, too, became extinct, isolatingthe ancestors of T. rhodesi and T. reddelli incaves. The comparatively robust endogean populationscontinue to live in Oaxaca and probably elsewherein Mexico in moist, heavily forested highlandsituations. The distribution pattern of Typhlochactasis very similar to that of the millipeds of the Cleidogonacrucis group in Oaxaca and Veracruz, and isprobably duplicated by the distribution of the millipedsof the genus Mexiterpes in San Luis Potosi andQueretaro. Typhlochactas elliotti has been collectedfrom the pants leg of the original collector and frombare rock on the cave ceiling and walls.Typhlochactas reddelli MitchellTyphlochactas reddelli Mitchell, 1968:762-776, fig.22-41; Reddell, 1971a:217; Reddell, 1971b:26;Mitchell, 19711:135, 138, 143, 145-146, fig. 2123, 25, 27; Muchmore, 1972a:275; Kawakatsu,1973b:255; Reddell, 1973a:32, 37; Vachon,1974:930,931,934, fig. 167,183,199-201; Soleglad,1976:253-254; Mitchell and Peck, 1978:159,162, 164, 167, 168.Typhlochactas reddeli: Gonzalez Sponga, 1974:56(erroneous spelling).Typhlochactas: Vomero, 1974:353 (Cueva del Ojo deAgua de Tlilapan record only).Typlochactas (part): Diaz Najera, 1975:3,35 (erroneousspelling).Typlochactas redelli: Diaz Najera, 1975:4, 33 (erroneousspelling).Type-Iocality.-Cueva del Ojo de Agua de Tlilapan,in the village of Tlilapan, Municipio de Tlilapan, approximatelyfive km sou th of Orizaba, Veracruz,Mexico.Distribution.-Known only from the type-locality.See Fig. 17.Discussion.-The single known specimen of thisspecies was collected from beneath a rock in the firstbat room of the cave.Typhlochactas rhodesi MitchellTyphlochactas rhodesi Mitchell, 1968:756-765,768776, fig. 1-21, 41, pI.; Delamare Deboutteville,1971:49; Reddell, 1971b:3, 27, cover; Mitchell,1971b:135, 138, 143, 145-146, fig. 18-20,24,27;Reddell and Mitchell, 1971b:181, 184, fig. 9; Reddelland Mitchell, 1971d:foll. table of contents;Sbordoni and Argano, 1972:8; Mitchell and Kawakatsu,1973a:671; Reddell, 1973a:32, 37; Reddelland Elliott, 1973b:181; Gonzalez Sponga, 1974:56; Kawakatsu, 1976:35; Soleglad, 1976:253,254; Mitchell and Peck, 1978:159,164,165,167,168.Typhlochactas: Vomero, 1974:353 [Cueva Chica(=Cueva de la Mina) record only].Typlochactas (part): Diaz Najera, 1975:3, 35 (erroneousspelling).Typlochactas rhodesi: Diaz Jajera, 1975:4,32 (erroneousspelling).Type-Iocality.-La Cueva de la Mina, Sierra deGuatemala, Municipio de Gomez Farias, Tamaulipas,Mexico.Distribution.-Known only from the type-locality.See Fig. 17.Discussion.-This species is known only by a fewspecimens taken from a single small upper level roomin this cave.us

Family DiplocentridaeDiplocentrus anophthalmus FranckeDiplocentrus anophthalmus Francke, 1977:56, 58-61,fig. 40-52, 54; Reddell, 1977b:230, 231,240,241,263, fig. 6; Francke, 1978:44-45.Eyeless diplocentrid scorpion: Mitchell and Peck,1978:167.Type-Iocality.-Acttin Chukum, Yucatan, Mexico.Distribution.-Known only from the type-locality.See Fig. 17.Discussion.-This is the most highly cave-adaptedspecies known in the family Diplocentridae. The me-dian eyes are absent and the lateral eyes are reducedto vestiges. It was found under rocks in the innerroom of Actun ChukuID. In addition to the two otherspecies of troglobite listed below, the family Diplor.entridaeis represented in caves by D. reddelliFrancke, known only from Aclun Xpukil, Yucatan;and by an undescribed species of Cazierius from ActUnLoltun, Yucatan.Diplocentrus cueva FranckeDiplocentrus cueva Francke, 1978:39-45, fig. 1-12.Type-Iocality.-Cueva Desapareciendo, 2 km W.Acatlan, Oaxaca, Mexico.MEXICO/II,,, /\///IIIIII ~GUATEMALAI .I I l\ •., ...J............ - ~· .....I_;"!--._.-._._. I[ // .....~! i\/ \.-! \ii\..'I\,.'-.,Ii/'-"1..JiL/r·\r- . r'.-'//Fig. 17.-Distribution of troglobitic scorpions: 1, Chactidae gen. et sp. 1; 2, Chactidae gen. et sp. 2; 3, Typhlochactasrhodesi; 4, T. elliotti; 5, T. reddelli; 6, Vaejovis gracilis; 7, Family incertae sedis; 8, Diplocentrus cueva; 9, D. mitchelli; 10, D.anophthalmus.116

Distribution.-Known only from the type-locality.See Fig. 17.Discussion.-This species, known only from theholotype male, is an attenuate form with reducedmedial eyes, pigmentation, and carinae. It is the leastcave adapted of the three troglobitic species of thegenus.Diplocentrus mitchelli FranckeDiplocentrus mitchelli Francke, 1977:56-58, 60, fig.29-39, 53; Reddell, 1977b:~30, 231, 2--10, 241,2--15, fig. 5; Francke, 1978:4--1-45.Type-Iocality.-Aclun Halmensura, Campeche,Mexico.Distribution.-Known only from the type-locality.See Fig. 17.Discussion.-This species is known only from theimmature holotype. It is an attenuate form with vestigialmedial and reduced lateral eyes and with reducedcarinae. The holotype was collected from undera small rock in the inner chamber of Accin Hal·mensura.Family VaejovidaeThree species of the family Vaejovidae have beenidentified from i\Jexican caves. Vaejovis gracilisGertsch and Soleglad is a probable troglobite and isdiscussed below. Williams (1968) described V. minckleyifrom a cave near Cuatro Cienegas, Coahuila.Vaejovis nigrescens Pocock was found in the entrancearea of Cueva de las Rusias, San Luis Potosi. Manyundescribed or immature specimens of Vaejovis havebeen collected from caves in Durango, Nuevo Leon,San Luis Potosi, and Tamaulipas.Vaejovis gracilis Gertsch and SolegladVejovis gracilis Gertsch and Soleglad, 1972:556,557,563, 593, 602, 603-604, 606, fig. 24-25, 80-83,147-149; Reddell, 1973a:37; Soleglad, 1975:107120, fig. 1-32.Type-locality.-Cueva de Atoyac, Atoyac, Veracruz,Mexico.Distribution.-Known from two caves in Veracruz.See Fig. 17.New record.-Veracruz: Solano de las Golondrinas(det. O. F. Francke).Discussion.-The holotype of this species is a veryimmature fonn. Soleglad (1975) reported V. gracilisfrom Cueva del Lencho Virgen, Oaxaca; Cueva de laTable l4.-Summary of cave-inhabiting Pseudoscorpionida.TroglobitesOther SpeciesHeterosphyronidaChthoniidae 7 3Tridenchthoniidae 0 1DiplosphyronidalIyidae 8 0ldeoroncidae 1 1Syarinidae 1 2Vachoniidae 7 0MonosphyronidaCheiridiidae 0 1Cheliferidae 0 1Chernetidae 0 8Total 24 17Barranca and Grutas de ] onotla, Puebla; and Cuevadel Vokancillo, Veracruz. He discussed the variationin the species and, based on the new specimens, concludedthat the species was not a troglobite. A largeadult was collected from Sotano de las Golondrinasin the same mountain range that contains Grutas deAtoyac. The specimen corresponds well with theholotype of V. gracilis and is almost certainly theadult of that species. Furthermore, it is not conspecificwith the specimens reported by Soleglad(Francke, pers. comm.). The Sotano de las Golondrinasadult is highly depigmented and attenuate andis a probable cave-adapted form.Order PseudoscorpionidaThe order Pseudoscorpionida is well represented inthe caves of Mexico, Guatemala, and Belize, with24 troglobitic and 17 troglophilic species having beenidentified to date (see Table 14). Many species, especiallyamong the troglophilic forms, await study_Suborder HeterosphyronidaFamily ChthoniidaeIn addition to the troglobitic species of Aphrastochthonius,Paraliochthonius, and Tyrannochthonius,the family Ch thoniidae includes two troglophileswhich are known from caves (see Fig. 18).Lechytia cavicola Muchmore is known only from batguano in the terminal room of Grutas de Cacahuamilpa,Guerrero. 11'fundochthonius mexicanus Muchmorewas described from leaf litter on Cuesta deChipinque, Nuevo Leon; it is also known from leaflitter near Teopisca, Chiapas, and from rat droppings117

in Crystal Cave, Tamaulipas. The genus Tyrannochthoniusalso includes at least one and probably severalundescribed species of troglophile from caves in SanLuis Potosi and Tamaulipas.Aphrastochthonius major MuchmoreAphrastochthonius new species: Reddell, 1973a:33.Aphrastochthonius major Muchmore, 1973a:47-48,fig. 1; Reddell and Elliott, 1973b:181, 183; Muchmore,1976:364.Type.locality.-Cueva de la Capilla, 13.5 km NWGomez Farias, Tamaulipas, Mexico.Distribution.-Known only from the type-locality.See Fig. 18.Discussion.-This large species is known only froma single female. The genus Aphrastochthonius appearsto be largely restricted to caves. Aphrastochthoniustenax Chamberlin and A. pecki Muchmore are knownonly from caves in Alabama (Muchmore, 1972c); A.pachysetus Muchmore is known only from Doc BritoCave, Eddy County, New Mexico (Muchmore, 1976);A. alteriae Muchmore is known only from litter onthe surface at Ruinas de Palenque, Chiapas (Muchmore,1977); A cubanus Dumitresco and Orghidanwas described from fissures on Isla de Pinos, CubaGUATEMALA/r/r"\,.. . r"...:-Fig. 18.-Distribution of troglobitic pseudoscorpions of the family Chthoniidae: 1, Aphrastochthonius n. sp.; 2, A. major;3, Tyrannochthonius troglobius; 4, A. parous; 5, A. TUsse/li; 6, T. pa/lidus; 7, Tyrannochthonius n. sp.; 8, Paraliochthoniwstrinatii; 9, A. verapazanus.118

(Dumitresco and Orghidan, 1977); the remammgfour species are troglobites from Mexico and Guatemalaand are listed below. An undescribed troglobitehas been recently collected in Sumidero de Oyamel,Tamaulipas.Aphrastochthonius parvus MuchmoreAphrastochthonius sp.: Reddell and Mitchell, 1971a:144.Aphrastochthonius parvus Muchmore, 1972c :437438, 440, 441, 442, fig. 4-7; Reddell, 1973a:33,37; Reddell and Elliott, 1973a:171, 173; Muchmore,1976:364; Dumitresco and Orghidan, 1977:100, 10l.Type-locality.~La Cueva de la Florida, 15 kilometersS.S.W. of Mante, Tamaulipas, Mexico.Distribution.~Known only from the type-locality.See Fig. 18.Discussion.~This is the smallest species in the genusand is known only from a single female collectedfrom the cave wall.Aphrastochthonius russelli MuchmoreAphrastochthonius russelli Muchmore, 1972c:440441,442, fig. 10-12; Reddell, 1973a:33, 37; Reddelland Elliott, 1973a:171, 173; Muchmore,1976:364.Type-locality.-La Cueva Pinta, about 12 kilometersN.E. of Valles, San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 18.Discussion.-This species is known only from asingle tritonymph. The species is probably mostclosely related to A. parvus, a species occurring inthe northern Sierra de EI Abra. Until adults of A.russelli are available, the relationships of this speciesmust remain uncertain.Aphrastochthonius verapazanus MuchmoreAphrastochthonius verapazanus Muchmore, 1972c:438, 440, 442, fig. 8-9; Muchmore, 1973a:48;Muchmore, 1976:364; Dumitresco and Orghidan,1977:101; Muchmore, 1977:63.Type.locality.-La Cueva Sepacuite No.2, Senahu,Finca Sepacuite, Alta Verapaz, Guatemala.Distribution.-Known only from the type-locality.See Fig. 18.Discussion.~This very attenuate species is knownonly from a single female.Paraliochthonius strinatii BeierParaliochthonius strinatii Beier, 1974:101-102, fig. 1;Strinati, 1977:388.Type-locality.-Cueva Chirrepeck, Alta Verapaz,Guatemala.Distribution.-Known only from the type-locality.See Fig. 18.Discussion.-In addition to this troglobite, the genusParaliochthonius is known by species from theeastern Mediterranean, Madeira, the Canary Islands,Jamaica, Puerto Rico, and Baja California and J alisco,Mexico (Muchmore, 1972b). Muchmore (pers.comm.) indicates that this species is probably misplacedand may belong in Tyrannochthonius.Tyrannochthonius pallidus MuchmoreTyrannochthonius pallidus Muchmore, 1973c:81-82,fig. 1-2.Type-locality.-Cueva de EI Jobo, 5 km NE Xilitla,San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 18.Discussion.-This species is known only from asingle male collected from under a rock. It is mostclosely related to T. troglobius, but it is smaller withless attenuate appendages. Troglobitic members ofthis genus are also known from caves in Alabama. Anundescribed troglobitic species has recently been collectedin Cueva de Las Maravillas, Oaxaca.Tyrannochthonius troglobius MuchmoreTyrannochthonius troglobius Muchmore, 1969:3132, fig. 1-3; Reddell, 1971b:27; Reddell andMitchell, 1971b:184; Mitchell and Kawakatsu,1973a:671; Muchmore, 1973c:81-82; Reddell,1973a:33, 37; Reddell and Elliott, 1973b:18l.Type-locality.-Mine Cave (=Cueva de la Mina),Rancho del Cielo, Tamaulipas, Mexico.Distribution.~Known only from the type-locality.See Fig. 18.Discussion.~This species is most closely related toT. pallidus. Both troglophilic and epigean species ofthe genus are known from the vicinity of Cueva de laMina, but all remain undescribed.Family TridenchthoniidaeA single species of the family Tridenchthoniidaehas been reported from Mexican caves. Tridenchthoniusjuxtlahuaca was described from bat guano inGrutas de Juxtlahuaca, Guerrero, and is presumably atroglophile (Chamberlin and Chamberlin, 1945) (seeFig. 18).119

Suborder DiplosphyronidaFamily HyidaeLeucohya heteropoda ChamberlinPseudoscorpion troglobio: Anonymous, 1942b:26S.Pseudoscorpiones nueva subfamilia: Bolivar, 1944:26.Leucohya heteropoda Chamberlin, 1946:7, 8-10,pI. 1; Chamberlin and Malcolm, 1960:114; Muchmore,1969:32; Reddell, 1971b:27; Muchmore,1972a:261, 269, 272,273; Muchmore, 1973a:S1,53, fig. 15-17.Leucohya heterodonta: Reddell, 1967a:24 (erroneousspelling).Type-locality.-Gruta del Palmito, Bustamante,Nuevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 19.Discussion.-This species is closely related to L.magnifica Muchmore, but the relationships of thisgenus to other neobisioid genera remain uncertain. Itis apparently related to Mexobisium, Apohya, andmore distantly to Paravachonium. An editorial addendumto Muchmore (1969) reports this species as beingknown from caves in Yucatan; this is an obviouserror. Topotypic specimens collected recently werefound under rocks in the entrance room of the cave.GUATEMALA,...-"r,.. \/.//Fig. 19.-Distribution of troglobitic pseudoscorpions of the family Hyidae: 1, Leucohya magnifica; 2, 1. heteropoda;3, Mexobisium paradoxum; 4, Troglohya carranzai; 5, M. maya; 6, T. mitchelli; 7, M. guatemalense; 8, M. goodnighti.120

Leucohya magnifica MuchmoreUndescrihed genus and species of pseudoscorpion:Reddell, 1967a:24.Hyidae, undescrihed genus and species: Reddell,1971h:27.Leucohya magnifica Muchmore, 1972a:271-272,fig. 12-13; Muchmore, 1973a:51; Reddell, 1973a:33,37.Type-locality.-Grutas del Carrizal, Nuevo Leon,Mexico.Distribution.-Known only from the type-locality.See Fig. 19.Discussion.-This species is known only from asingle female. Both species of Leucohya are knownfrom single caves in isolated limestone ranges separatedby desert terrain.Mexobisium goodnighti l\luchmoreMexobisium goodnighti Muchmore, 1973b:69-71, fig.22-25; Muchmore, 1977:71.Type-locality.-Cave near Augustine, Belize.Distribution.-Known only from the type-locality.See Fig. 19.Discussion.-This large species with attenuated appendagesis the only species of the genus known fromBelize. It is very distinct and may eventually provegenerically distinct from other species of Mexobisium.The genus Mexobisium includes seven speciesof fairly disparate morphology. Epigean species areknown from Cuba and from Chiapas and Oaxaca,Mexico. Troglohites occur in Guatemala, Belize, andTabasco and Veracruz, Mexico.}Uexobisium guatemalensf! 1\IuchmoreMexobisium guatemalense Muchmore, 1973h:67, 69,71, fig. 18-21; Muchmore, 1977:71.Type-locality.-Cueva Lanquin, Alta Verapaz,Guatemala.Distribution.-Known only from the type-locality.See Fig. 19.Discussion.-This is the only species of the genusknown to occur in Guatemala.Jlexobisium maya \luchmoreMexobisium maya l\luchmore, 1973b:67, 69, 71, fig.12-17; Muchmore, 1977:71.Type-locality.-Grutas de Cocona, 3 km NE Teapa,Tahasco, Mexico.Distribution.-Known only froUl the type-locality.See Fig. 19.Discussion.-This species is known from a singlemale, two females, and two tritonymphs. Its relationshipto other memhers of the genus is not clear althoughit is similar to M. pecki Muchmore, an epigeanspecies, in its general confonnation.Mexobisium paradoxum MuchmoreMexobisium paradoxum Muchmore, 1972a:273-275,fig. 14-19; Reddell, 1973a:37; Muchmore, 1973h:63,71-72.Mexobisium garadoxum: Reddell, 1973a:33 (erroneousspelling).Type-locality.-Cueva del Ojo del Agua de Tlilapan,Tlilapan, Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 19.Discussion.-This species is known only from theholotype female collected from a flowstone slope inthe Formation Room at the type-locality. J'1. paradoxumis unique in many respects, and its affinitiesto other memhers of the genus are ohscure.Troglohya carranzai BeierTroglohya carranzai Beier, 1956:83, 84-85, fig. 2;Chamherlin and Malcolm, 1960: 114; Barrera,1968:313; Muchmore, 1969:32; Reddell, 1971h:27; Muchmore, 1972a:261, 272; Muchmore,1973a:54,55.Type-locality.-Cueva de Monteflor, cerca de ValleNacional, N. Oaxaca, Mexico.Distribution.-Known only from the type-locality.See Fig. 19.Discussion.-This species is known only from asingle tritonymph. It is related to M. mitchelli Muchmore,but until further material is availahle, the degreeof relationship is unknown. The genus Troglohyamay prove to he more closely related to Vachoniumthan to Leucohya.Troglohya mitchelli MuchmoreTroglohya mitchelli Muchmore, 1973a:55-56, fig. 2431; Iuchmore, 1977:71.Type-Iocality.-Grutas de Zapaluta, 6.5 kill SE LaTrinitaria, Chiapas, Mexico.Distribution.-Known only from the type-locality.See Fig. 19.Discussion.-This species, known from a single female,is larger and more attenuate than T. carranzai.Family IdeoroncidaeAlbiorix bolivari BeierAlbiorix bolivari Beier, 1963: 133-134, fig. 1; Barrera,1968:313; Muchmore, 1969:32; Reddell, 1971b:27; Muchmore, 1972a:261.121

Type-locality.-Gruta de Acuitlapan, Guerrero,Mexico.Distrihution.-Known only from the type-locality.See Fig. 20.Discussion.-This species is most closely related toA. magnus Hoff. Undescribed troglobites belonging tothe genus Albiorix are known from Cueva del Diablo,Veracruz, and Grutas de Atepolihuit, Puebla. An undescribedtroglophile is known from Cueva de LasMaravillas, Oaxaca.Family SyarinidaePachychitra grandis MuchmorePachychitra sp.: Reddell, 1971a:219.Pachychitra grandis Muchmore, 1972a:266, 267, fig.6-7; Reddell, 1973a:33, 37; i\luchmore, 1977:70.Type-locality.-Cueva del 'fio Ticho, one milesouth of Comitan, Chiapas, Mexico.Distribution.-Known only from the ty pe-Iocality.See Fig. 20.MEXICOGUATEMALAr\rr-..-"///Fig. 20.-Distrihution of troglobitic and troglophilic pseudoscorpions of the families Ideoroncidae, Syarinidae, and Vachoniidae:1, Paravachonium n. sp.; 2, Paravachonium $uperbum; 3, Paravachonium bolivari; 4, PachychitTa similis; 5, Albiorixn. sp. 1; 6, Albiorix n. sp. 2; 7, A. bolivari; 8, Albiorix n. sp. 3; 9, Pachychitra grandis; 10, Vachonium cryptum; 11, Vachoniumsp. 1; 12, PachychitTa maya; 13, Vachonium sp. 2; 14, V. boneti; 15, V. maya; 16, V. kauae; 17, V. belizeme.122

Discussion.-This is the largest species of the genus.The greater size and more slender appendagessuggest that it is restricted to the cave habitat. Thegenus is known from Florida (U.S.A.), Jamaica,Puerto Rico, Cura/iao, Tamaulipas, Chiapas, and Yucatan.Two additional species are known only fromcaves (P. maya Chamberlin from Yucatan and P. similisMuchmore from Tamaulipas), but are smaller andless attenuate than P. grandis. Nicholas (1962) listsP. maya as a troglobite, but it is probably best consideredto be a troglophile.Family VachoniidaeParavachonium bolivari BeierParavachonium bolivari Beier, 1956:81, 82-83, 85,fig. 1; Chamberlin and Malcolm, 1960:114; Reddell,1965a:19; Barrera, 1968:313; Muchmore,1969:32; Reddell, 1971b:27; Reddell and Mitchell,1971a:144; Muchmore, 1972a:261, 267, 268,269; Muchmore, 1973a:57, fig. 32; Reddell andElliott, 1973a:I71, 173.Paravachonium: Reddell, 1967c:55.Type-locality.-Cueva (=Grutas) de Quintero, Tamaulipas,Mexico.Distribution.-Known only from the type-locality.See Fig. 20.Discussion.- This species is known only from theholotype female and a deutonymph. Although Beierconsidered this species as belonbring to the familyVachoniidae, Muchmore (1973a) considers the affinitiesof the genus to be obscure. Paravachonium isknown only from these two species and an undescribedtroglobite from Sumidero de Oyamel, Tamaulipas.Paravachonium superbum l\luchmoreParavachonium sp.: Reddell, 1967c:55; Reddell,1971b:27; Reddell and Mitchell, 1971b:184-185;Vomero, 1974:350.Paravachonium superbum Muchmore, 1972a:268,269, 271, fig. 8-11; Reddell, 1973a:33, 37; Reddelland Elliott, 1973b:181, 183.Type.locality.-Sotano de Gomez Farias, justsouth of the village of Gomez Farias, Tamaulipas,Mexico.Distrihution.-Known only from the type-locality.See Fig. 20.Discussion.-This species is known from a single femalecollected beneath a rock at the 25-meter level ofthe cave. This large, attenuate species is closely relatedto P. bolivari.Vachonium belizense MuchmoreVachonium belizense Muchmore, 1973a:58, fig. 3638; Muchmore, 1977:72.Type.locality.-Mountain Cow Cave, Caves Branch,Belize.Distribution.-Known only from the type-locality.See Fig. 20.Discussion.-This is the largest species of the genusand the only one known outside of the Mexican stateof Yucatan. It is known only from a single tritonymph.Vachonium boneti ChamberlinVachonium boneti Chamberlin, 1947:4,5,6-7,8,9,fig. 1-23; Chamberlin and Malcolm, 1960:114;Muchmore, 1969:32; Reddell, 1971b:27; Muchmore,1972a:261, 267; Muchmore, 1973a:57;Muchmore, 1977:72; Reddell, 1977b:230, 240,282.Type.locality.-Cueva de Sabaca (=Actun Sabaca),Yucatan, Mexico.Distribution.-Known only from the type-locality.See Fig. 20.Discussion.-This species is known only from asingle female. The genus Vachonium is possiblyclosely related to Troglohya. Vachonium containsonly the f; ie troglobites listed here. An editorial addendumto l\luchmore (1969) lists this species asoccurring in "Ia Cueva de Palmito, 'uevo Leon";this is an obvious error. A single tritonymph fromActUn Xpukil, Yucatan, is this or a closely relatedspecies (Muchmore, 1977).Vachonium cryptum MuchmoreVachonium cryptum Muchmore, 1977:72, 73, fig.20-21; Reddell, 1977b:230, 240, 271.Type-locality.-ActUn Xkyc, 1 km S Calcehtok,Yucatan, Mexico.Distribution.-Known only from the type-locality.See Fig. 20.Discussion.-This species, known only from asingle female, is closely related to V. kauae and V.maya. The only specimen was collected from theunderside of a Mayan stone water dish.Vachonium kauae MuchmoreVachonium kauae Muchmore, 1973a:57-58, fig. 3335; Muchmore, 1977:72; Reddell, 1977b:230,239, 240, 260.Type-locality.-Cueva de Kaua (=ActUn Kaua),1 km S Kaua, Yucatan, Mexico.123

Distrihution.-Known only from the type-locality.See Fig. 20.Discussion.-This species is known only from asingle female. It appears to be most closely related toV. maya but is less attenuate.Vachonium maya ChamberlinVaehonium maya Chamberlin, 1947:5, 8-9, fig. 2436; Chamberlin and Malcolm, 1960: 114; Muchmore,1969:32; Reddell, 1971b:28; Muchmore,1972a:261, 267; Muchmore, 1973a:57; Muchmore,1977:72; Reddell, 1977b:230, 239, 240,285.Type-locaIity.-Cueva de Balaam Canche (=Grutasde Balankanche), Chichen Itza, Yucatan, Mexico.Distribution.-Known only from the type-locality.See Fig. 20.Discussion.-This species appears to be mostclosely related to the geographically near species V.kauae. It has recently been collected from beneathrocks at the type-locality.Suborder MonosphyronidaFamily CheiridiidaeThe family Cheiridiidae is known from undeterminedspecimens recently collected from two caves inthe Sierra de Ticul, Yucatan.Family CheliferidaeA single species of the· family Cheliferidae, Mexieheliferreddelli Muchmore, has been collected froma cave in Mexico. It was found in the entrance area ofCueva de las Carnicerias, San Luis Potosi. Althoughits ecological status is uncertain, it is probably a troglophile.Family ChernetidaeThe family Chernetidat is the best represented ofall families of pseudoscorpions in Mexican caves, althoughapparently contributing no troglobites to thefauna. J. C. Chamberlin (1938) described two speciesof this family from caves in Yucatan: Lustroehernesminor and Parazaona eavieola. Nicholas (1962) listedthese as troglobites, but they should be considered astroglophiles. Specimens from caves in all parts ofMexico, Guatemala, and Belize that have been visitedare now under study by Dr. W. B. Muchmore. He hastentatively identified the genera Dinoeheirus, Hesperoehernes,Lustroehernes, Neoalloehernes, and Semeioehernesfrom caves in this area. Chernetids are commonlyfound in bat guano, where they may be presentin vast numbers.Order UropygidaFamily ThelyphonidaeThe order Uropygida is primarily tropical in distribution,although one species ranges north into subtropicalregions. A single species of whipscorpion,Mastigoproetus giganteus Lucas, is known from thecaves of Mexico. It has been found on walls near theentrance of eleven caves in the states of Coahuila,Hidalgo, San Luis Potosi, and Tamaulipas.Order SchizomidaThe order Schizomida is an important and, untilrecently, a neglected element of the soil and litterfauna of Mexico and Central America. It is worldwidein distribution but rare in temperate regions. In NorthAmerica schizomids are known from Panama into thesouthern United States. The order in Mexico appearsto be limited to southern Mexico and along the eastternslopes of the Sierra Madre Oriental to Monterrey.Relict populations have been found in caves andother mesic habitats north and east into south Texas.Epigean schizornids are commonly found beneathrocks along hillsides, but they also may be collectedin leaf litter. Cavernicole schizomids are usuallyfound on the underside of rotting wood, in organicdebris, and beneath rocks. Some are found runningover silt banks and in bat guano.All North American schizomids lack true eyes, althoughsome retain small "eyespots." Epigean speciestend to he fairly robust, orange-brown to green, andmost have distinct eyespots. The cavernicole speciesrange from very dark,' robust forms to elongate, paleforms. Ten of the described species appear to be sufficientlypale, delicate, and poorly sclerotized towarrant inclusion in this list as cave-adapted forms.Family ProtoschizomidaeThe family Protoschizomidae contains two genera,each with two described species. Protosehizomuspachypalpus (Rowland) is known only from epigeanlocalities in Tamaulipas, while P. oeeidentalis Rowlandis known only from a single epigean site in Colima.The genus Agastosehizomus contains twodescribed and one undescribed species of troglobite.Agastosehizomus huitzmolotitlensis RowlandSehizomus sp.: Reddell, 1967d: 106; Reddell, 1971b:28 (S6tano de Huitzmolotitla record only).Agastosehizomus huitzmolotitlensis Rowland, 1975b:6,8-10, fig. 3; Rowland, 1975a:28, 44,45,46-47,48-49,50, 167-168, fig. 17; Rowland and Reddell,1977:80, 81-82, fig. I; Rowland and Reddell,1979a:162, 167, 169, 170, fig. 4, 7.124

Type-Iocality.-S6tano de Huitzmolotitla, 2 kmSW Tlamaya, and approximately 10 km N Xilitla, SanLuis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 21.Discussion.-This species is smaller and has relativelylonger appendages than does A. lucifer. Knownonly by a single male, it is unique in being the onlydescribed Mexican schizomid from high elevations(about 700 meters). An undescribed species of Agastoschizomushas been found at about this same elevationin caves in Hidalgo.Agastoschizomus lucifer RowlandAgastoschizomus lucifer Rowland, 1971a:15, 17, fig.1-8; Reddell and Mitchell, 1971a:145, fig. 3-4;Dumitresco, 1973:282; Reddell, 1973a:33, 38;Reddell and Elliott, 1973a:171; Rowland, 1973a:10; Rowland, 1973b:197, 202, fig. 2, 4; Rowland,1973c:136; Brignoli, 1974a:150; Rowland, 1975a:8-9, 14-15, 27,28,43-45,46-47,48-49,50, 167168, 181,211, fig. 1,8, 13,16; Rowland, 1975b:8, 9, 10, fig. 4; Rowland and Reddell, 1977:80,81, 82, 85, 96, fig. 1; Rowland and Reddell,1979a:162, 167, 168, 169-170, fig. 4, 7.\.""- ,"-\'~\'", -., ;I-J-.......·_·-." "\'>-,r "\\ I"- \-.~,I~\\\\-~~~-MEXICO(/___-J ....... '-.r!L lI,rJ)"t,../;r/i(,ItGUATEMALAFig. 21.-Distribution of troglobitic schizomids of the family Protoschizomidae: 1, Agastoschizomus lucifer; 2, A. huitzmolotitlensis;3, Agastoschizomus n. sp.125

Agastochizomus lucifer: Rowland, 1971a: 13 (erroneousspelling).Type-locality.-Sotano de la Tinaja, 10 km ~ECiudad Valles, San Luis Potosi, Mexico.Distribution.-Known only from three caves in theSierra de El Abra, near Valles, San Luis Potosi. SeeFig. 21.Records.-San Luis Potosi: Sotano de Matapalma,Sotano de la Tinaja, and Sotano de Yerbaniz.Discussion.-This is the largest species in the orderSchizomida. It is abundant on silt and along walls inSotano de la Tinaja. Although Brignoli (1974a) hasreservations about considering any schizomid to be atroglobite, the habitat and general facies of this andthe preceding species leave little doubt but that theyare restricted to the cave habitat.Famity SchizomidaeThe family Schizomidae includes 24 species whichhave been reported from the cave habitat in ~lexico,Guatemala, and Belize. Twelve of these are consideredto be troglobites; eight have been describedand are discussed below. The remaining species showno indication of adaptation for a cave existence, andsome are also known from epigean collections.Rowland and Reddell (1979) have subdivided theNew World schizomids of the genus Schizomus intoseven species groups. The dumitrescoae group is representedonly in the Antilles and Costa Rica. Althoughseveral species are known only from caves,their ecological status is uncertain. The simonisgroup ranges from northern South America to CostaRica; none are known from caves. The briggsi groupis known only from Arizona and California in theUnited States, with one species, S. shoshonensis(Briggs and Hom), being an apparent troglobile inUpper Shoshone Cave, Inyo County, California.The brasiliensis group ranges from Brazil tosouthern Mexico. Two species of this group haveinvaded caves. Schizomus stewarti Rowland is knownonly from Cueva del Guayabo, Oaxaca. Schizomustrilobatus Rowland is a dark species with distincteyespots and is known only from Grutas del Cocona,Tabasco. The only other brasiliensis group speciesknown from 1exico is S. lacandonus Rowland fromLas Ruinas de Palenque, Chiapas.With the exception of one species, the mexicanusgroup is restricted to Mexico, Belize, and Guatemala,where it is abundantly represented in caves. Schizomusportoricensis (Chamberlin) has been collec tedfrom Ecuador, the Galapagos Islands, southeasternMexico, the Antilles, and southern Florida. Peripheralpopulations are all parthenogenetic; parthenogeneticpopulations also occur in caves in the Yucatan Peninsula(Rowland and Reddell, 1977). Six species of themexicanus group are probably cave-adapted formsand are discussed below. Schizomus mexicanus Rowlandis known both from caves and the surface in theSierra de EI Ahra and in the lowland Sierra de Guatemala.Schizomus moisii Rowland is a dark specieswith distinct eyespots; it is known only from Grutasde Monteflor, Oaxaca. An undescribed species fromGrutas de Cacahuamilpa, Guerrero, lacks eyespotsand may be a troglobite.The pecki group is known only from sou themMexico, Guatemala, and Belize. Schizomus firstmaniand S. pecki are apparently troglobites and are discussedbelow. Schizomus sbordonii Brignoli is a trog10phiJe described from Grutas de Atoyac, Vera(;ruz.Undescribed species from caves ncar Comilan, Chiapas,and from Grutas de :\lontef!or, Oaxaca, areapparently troglophiles. An undescribed species fromS1. Herm an's Cave, Belize. lacks eyespots and is probablya troglobile.Four species have been described in the goodnightorumgroup. One species from Yucatan and asecond from Chiapas are known only from the surface.The remaining species are represented only incave collections but show no modifications for cavelife. Schizomus lanceolatus Rowland is known onlyfrom Cueva del Diablo, Veracruz: S. silvino Rowlandand Reddell has been collected only in Gruta de Silvino,habal, Guatemala. See Fig. 22 for the distributionof troglophili(; Schizomus in J\lexico and Guatemala.Schizomus bartolo RowlandSchizomus sp.: Reddell, 1967a:25; Reddell, 1971b:28 (Grutas de San Bartolo record only).Schizomus bartolo Rowland, 1973a:13-16, 18, fig.11-13; Rowland, 1973c:135, 137; Rowland,1975a:34, 166, 169, 173-174, 176,177,214-215,218-219, 222-223, 228-229, 366-367, 368-369,394, fig. 158, 174, 193, 292; DUlIlitreseo, 1977:157: Rowland and Reddell, 1977:80, 83, 84, fig.2; Rowlano and Reddell, 1979a:163.Type-locality.-Gruta de San Bartolo, 16 km SSWMonterrey, Nuevo Leem, Mexico.Distribution.-Known only from Grutas de SanBartolo. See Fig. 23.Records.-Nuevo Leon: Grutas de San Bartolo andGruta Sur de San Bartolo.Discussion.-Grutas de San Bartolo is located in adesert ref,rion, and S. bartolo is apparently an isolatedpopulation now restricted to the cave habitat. It ismost closely related to S. firstmani. Grutas de San126

Bartolo is a name applied to two nearby caves (designatedas Sur and Norte); it is not known from whichcave the type series was collected.Schizomus cookei RowlandSchizomus cookei Rowland, 1971b:122·123, fig. 1012, 19; Reddell and Mitchell, 1971a:145; Dumitresco,1973:291; Reddell, 1973a:38; Rowland,1973c:135; BrignoIi, 1974a:146; Rowland, 1975a:34, 166, 167·168, 169, 181, 182, 210-211, 212,213,216-217,218-219, 220-221, 228,231,366367, 368-369, 395, fig. 165, 180, 183-184,292;Rowland and Reddell, 1977:80, 84, 85, fig. 2;Rowland and Reddell, 1979a: 163.Type-Iocality.-Sotano de la Tinaja, 10.5 km NEof Ciudad Valles, San Luis Potosi, Mexico.Distribution.-Known only from two caves northof Valles in the Sierra de EI Abra, San Luis Potosi.See Fig. 23.Records.-San Luis POtOSI: Sotano de la Tinajaand Sotano de Yerbaniz.Discussion.-This elongate species is sympatric inboth of these caves with Agastoschizomus lucifer; italso occurs with S. mexicanus in Sotano de la Tinaja.Extensive epigean collections have failed to includeS. cookei, although S. mexicanus is abundant on thesurface throughout the Sierra de EI Abra.GUATEMALA,. //\.... r.--/Fig. 22.-Distrihution of troglophilic schizomids of the family Schizomidae: 1, Schizomus mexicanus; 2, S. lanceolatus;3, S. sbordonii; 4, S. moisii; 5, S. stewarti; 6, S. trilobatus; 7, S. arganoi; 8, S. portoricensis.127

Schizomus firstmani RowlandSchizomus sp.: Reddell, 1971a:219 (Grutas de Atoyacrecord only).Schizomus firstmani Rowland, 1973a:16-19, fig. 1416; Rowland, 1973c:136; Rowland, 1975a:34,167-168, 232, 234, 235-237, 238, 239-240, 243,246-247, 248-249, 252-253, fig. 217, 219-220,226-227; Rowland and Reddell, 1977:80,84,98,99, fig. 2; Rowland and Reddell, 1979a:163.Type-Iocality.-Grutas de Atoyac, 2 km E Atoyac,Veracruz, Mexico.Distribution.-Known only from the type-localityand possibly caves near Acatlan, Oaxaca. See Fig. 23.Records.-Oaxaca: ?Cueva Desapareciendo, ?Cuevade la Finca, and ?Cueva del Nacimiento del Rio SanAntonio; Veracruz: Grutas de Atoyac.Discussion.-This species is most closely related toS. bartolo, and is sufficiently delicate and elongate tobe considered a probable troglobite. The Oaxacanrecords for this species are based on females andshould be verified by collection of males.Schizomus lukensi RowlandSchizomus lukensi Rowland, 1973c: 136-137, fig. 1,4; Rowland, 1975a:34, 166, 169, 174, 175-176,177, 214-215, 218-219, 222-223, 228-229, 366367, 368-369, 394, fig. 160, 173, 191-192,292;MEXICOGUATEMALAFig. 23.-Distribution of troglobitic schizomids of the family Schizomidae: 1, Schizomus bart%; 2, S. /ukensi; 3, S.reddeUi; 4, S. mitche/li; 5, S. cookei; 6, S. flTstmani; 7, S. pallidus; 8, S. pecki; 9, Schizomus spp.128

Rowland, 1975b:19, 20; Dumitresco, 1977:157;Rowland and Reddell, 1977:80, 83-85, 98, fig. 2;Rowland and Reddell, 1979a: 163.Type-Iocality.-Cueva del Agua (de Simon Salinas),50 km SW Soto la Marina, Tamaulipas, Mexico.Distrihution.-Known only from two caves in theSierra de Tamaulipas, Tamaulipas. See Fig. 23.Records.-Tamaulipas: Cueva del Agua de SimonSalinas and Cueva de la Virgen de Guadalupe.Discussion.-This pale species with long first legsis apparently isolated in its moist cave habitat by thesemi-arid environment of the Sierra de Tamaulipas. Itappears to be most closely related to S. mexicanus.Rowland (1975a) tentatively reported females fromCueva de los Cuarteles, Tamaulipas, as belonging tothis species. The recent discovery of males from thispopulation indicates that it is an undescribed speciesvery closely related to S. lukensi.Schizomus mitchelli RowlandSchizomus sp.: Mitchell, 1970b:65.Schizomus mitchelli Rowland, 1971b:121-122, fig. 79, 18; Reddell and Mitchell, 1971a:145; Brignoli,1973a:6; Reddell, 1973a:38; Rowland, 1973c:135; Brignoli, 1974a:145-146, fig. la, 2e; Rowland,1975a:34, 166, 167-168, 169, 181, 182,211-213, 216-217, 228, 231,366-367,368-369,395, fig. 164, 182, 188, 212-215, 292; Dumi·tresco, 1977:157; Rowland and Reddell, 1977:80,84, 85, 96, fig. 2; Rowland and Reddell, 1979a:163.Schizomids: Rowland, 1972a:73.Type-Iocality.-Cueva de El Pachon, 7.5 km NE ofAntigua Morelos, Tamaulipas, Mexico.Distribution.-Known only from three caves in thenorthern Sierra de El Abra, Tamaulipas. See Fig. 23.Records.-Tamaulipas: Cueva de la Florida, Cuevade El Pachon, and Grutas de Quintero.Discussion.-This species possesses slender elongateappendages and is almost certainly a troglobite. Collectionsoutside of the entrance of Cueva de la Floridahave included only S. mexicanus. A female ofthis species was observed carrying eggs. This is un·usual behavior since previous accounts of schizomidsindicated that they built cocoons and the female remainedin the cocoon until the eggs hatched. Thismay be an adaptation to the cavernicole existenceand its comparatively low number of predators.Schizomus pallidus RowlandSchizomus pallidus Rowland, 1975b:7, 13-15, fig. 6;Rowland, 1975a:34, 166, 167-168, 169,184-186,188, 216-217, 218·219, 222-223, 228, 230, 280,366-367, 368-369, 396, fig. 166, 179, 196,292;Rowland and Reddell, 1977:80,84,87,88, fig. 2;Rowland and Reddell, 1979a:163.Type-Iocality.-Cueva Macinga, Tlilapan, Veracruz,Mexico.Distribution.-Known only from the type-locality.See Fig. 23.Discussion.-This pale species with elongate appendagesappears to be related to S. mexicanus but isquite distinct in some respects. It was found on siltbeneath rocks in a dry side passage of the cave.Schizomus pecki RowlandSchizomus pecki Rowland, 1973a:19-23, fig. 17-19;Rowland, 1973c:136; Sbordoni et al., 1974:1415; Rowland, 1975a:34, 135, 167-168, 188,209,232, 234, 239-241, 242, 243, 244, 246-247, fig.216,218,221,231; Dumitresco, 1977:157; Rowlandand Reddell, 1977:80, 83, 84, 96, 98-99,fig. 2; Rowland and Reddell, 1979a:163; Rowlandand Reddell, 1979b:l04.Type-Iocality.-Las Grutas de Cocona, 2 mi NETeapa, Tabasco, Mexico.Distribution.-Known only from two caves nearTeapa, Tabasco. See Fig. 23.Records.-Tabasco: Grutas del Cocona and Resumideradel Cocona.Discussion.-This large, pale species is abundant onsilt under rotten wood in the more remote sectionsof Grutas del Cocona. A dark robust species, S. trilobatus,inhabits litter near the cave entrance.Schizomus reddelli RowlandSchizomus reddelli Rowland, 1971b:123, 124, 126,fig. 13-15; Reddell and Mitchell, 1971b:185; Reddell,1973a:38; Rowland, 1973a:21; Rowland,1973c:135; Brignoli, 1974a:147, 149; Rowland,1975a:34, 166, 169, 177, 178-179,181,214-215,218-219, 220-221, 228-230, 366-367, 368-369,394, fig. 159, 175, 189-190; Rowland and Reddell,1977:80, 84, 85, fig. 2; Rowland and Reddell,1979a:163.Schizomus mexicanus Rowland: Reddell and Mitchell,1971b:185 (misidentification); Vomero, 1974:341 (misidentification).Schizomus reddeli: Dumitresco, 1977: 157 (erroneousspelling).Type-Iocality.-Cueva de Tres Manatiales (=Cuevadel Ojo de Agua de Manantiales), 23 km NW Limon,Tamaulipas, Mexico.Distribution.-Known from two caves in the Sierrade Guatemala, Tamaulipas. See Fig. 23.129

Records.-Tamaulipas: Cueva del Ojo de Agua deManantiales and Cueva de los Vampiros.Discussion.-This species is very closely related toS. mexicanus. Rowland (1975a) speculates that thismay represent a high altitude relict of a species ancestralto it and S. mexicanus.Order AmblypygidaThe order Amblypygida is world-wide in distributionbut, like the Schizomida, barely reaches northinto temperate regions. In North America it rangesthroughout Central America into the southern UnitedStates. Epigean amblypygids are found under rocks,on cliff-faces at night, on the walls of buildings, andin culverts and beneath bridges. Cavernicole amblypygidsare seen, frequently in large numbers, alongcave walls, on flowstone mounds, and beneath rocks.Family CharontidaeThe family Charontidae is known from caves onlyby an undescribed troglobite of uncertain generic affinities.It has recently been collected in FootprintCave, Belize.Family PhrynidaeThe described species of amblypygid known fromMexico and Central America include five troglobiticand six troglophilic species belonging to the familyPhrynidae. Several undescribed species, including atleast one troglobite, are known from caves in thisregion.The large, distinctive species Acanthophrynuscoronatus Koch is known from an unidentified limestonecave at Ixtlahutan, Colima. Its ecological statusis unknown.The genus Phrynus is represented in Mexican cavesby a probably undescribed species from Grutas delCocona, Tabasco, and by undetermined specimensfrom several other caves. This genus is much rarer incaves than is Paraphrynus.Mullinex (1975) has revised the genus Paraphrynusfor North America. In addition to the four troglobiteslisted below, she included six species as probable troglophiles.A fifth species of troglobite has since beendescribed from Mexico (Mullinex, 1979). Paraphrynusraptator (pocock) is an extremely abundantspecies in the caves of the Yucatan Peninsula. It isknown from many caves in Campeche, Quintana Roo,and Yucatan, Mexico; and in caves in EI Peten, Guatemala,and Belize. See Fig. 24 for the distribution oftroglophilic Paraphrynus in Mexico, Guatemala, andBelize.Paraphrynus williamsi Mullinex and P. emaciatusMullinex are rare species known only from Grutas deZapaluta, Chiapas, and Cemetery Cave, Alta Verapaz,Guatemala, respectively. The latter species has elongateappendages but is otherwise not adapted forcave life.Paraphrynus mexicanus (Bilimek) is a highly variablespecies with three distinct morphological formsseparated by considerable distributional gaps. Thesethree forms may be eventually given specific recognition.The so-called Cacahuamilpan form has beenfound in caves in Guerrero, Mexico, Oaxaca, andPuebla, Mexico. The other two forms (Arizonan andCuban) have not been found in caves and occur respectivelyin northern Mexico and the southwesternUnited States and in Cuba.Paraphrynus pococki Mullinex is a somewhat variablespecies known only from cave and epigean habitatsin San Luis Potosi and Tamaulipas. This is thecommonly occurring species in the caves of the Sierrade EI Abra.Paraphrynus azteca (Pocock) is represented by twodistinct forms. The Isthmus form is known from Chiapas,Tabasco, Oaxaca, and southern Veracruz,Mexico. The Atoyac fonn occurs in more northernVeracruz in the vicinity of Cordoba and Orizaba.Both forms are known from caves. This or a relatedspecies is known from caves in the Cuetzalan area ofPuebla.Paraphrynus baeops MullinexParaphrynus baeops Mullinex, 1975:1,9,26,28-29,43, fig. 27-28, 39.Type.locality.-Sotano de Vasquez, 10 km SEOcampo, Tamaulipas, Mexico.Distribution.-Known from four caves in the Sierrade Guatemala and Sierra de EI Abra, Tamaulipas. SeeFig. 25.Records.-Tamaulipas: Cueva de la Florida, Grutasdel Puente, and Sotano de Vasquez.New record.-Tamaulipas: Sotano de Santa Elena(det. C. Mullinex).Discussion.-This species possesses median eyeswhich are greatly reduced in size and lacking in pigment;furthermore, the median ocular tubercle isabsent.Paraphrynus chacmool (Rowland)Tarantula chacmool Rowland, 1973d:123, 125-126,128, fig. 1,3,5,7,9,11,13,15,17.Paraphrynus chacmool: Mullinex, 1975:43; Reddell,1977b:230, 239, 240, 256, 258, 260, 263, 267,273,276,279,281,282,285,289,293.130

Type-Iocality.-Actlin Kaua, Yucatan, Mexico.Distribution.-Known from caves in Quintana Rooand Yucatan. See Fig. 25.Records.-Yucauin: Cenote de Aka Chen, Grutasde Balankanche, Cenote Calchum, Cenote Chen Mul,Actlin Chukum, Cenote de Hoctun, Actlin Kaua,ActUn Sabaca, Cenote de Sihunchen, Cueva de Tecoh,Grutas de Tzab-Nah, Actun Xpukil, Cenote Xtacabiha,and Actlin Ziz.New records.-Quintana Roo: Cenote de Las Ruinas;Yucatan: Cenote de Catzin and Cueva Escondida(det. C. Mullinex).Discussion.-Most of these caves are also inhabitedby P. raptator. The eyes of this species are greatly reducedin size and are depigmented; the ocular tubercleis nearly obsolete; the body is generally depigmented;and the appendages are elongated.Paraphrynus chiztun (Rowland)Tarantula chiztun Rowland, 1973d:125, 126, 128,fig. 4, 6, 8, 10, 12,14,16,18.Paraphrynus chiztun: Mullinex, 1975:43.Type-Iocality.-Las Grutas de Cocona, Teapa, Tabasco,Mexico.Distribution.-Known only from the type-locality.See Fig. 25.Discussion.-This species is apparently mostclosely related to P. chacmool, but its relationship toMEXICOGUATEMALA.s //(/,..'.;..-'r'"'Fig. 24.-Distribution of troglophilic amblypygids of the family Phrynidae: 1, Paraphrynus pococki; 2, Paraphrynus mexicanus;3, Paraphrynus azteca; 4, Paraphrynus williamsi; 5, Paraphrynus emaciatus; 6, Paraphrynus raptator; 7. Phrynus sp.131

other members of the genus are unclear. Grutas delCocona is also inhabited by a probably undescribedspecies of the genus Phrynus. Paraphrynus chiztunhas been observed running across the walls of Grutasdel Cocona near the end of the cave. Phrynus sp. wastaken from flowstone and walls and beneath rocksmuch closer to the cave entrance. Paraphrynus chiztunhas reduced and depigmented eyes, reduced oculartubercle, depigmented body, and elongated appendages.Paraphrynus reddelli MullinexParaphrynus reddelli Mullinex, 1979 :267-269, fig. 1-7.Type-locality.-Act6n Loltun, 7 km SSW Oxkutzcab,Yucatan, Mexico.Distribution.-Known only from the type-locality.See Fig. 25.Discussion.-This is a completely eyeless species;it is very different in many respects from other describedspecies of Paraphrynus and its relationshipsare uncertain.Paraphrynus velmae MullinexParaphrynus velmae Mullinex, 1975:1,9,26-28,40,65,66,67, 78, fig. 25-26, 38.Type-locality.-S6tano de Tlamaya, San Luis Potosi,Mexico.MEXICOGUATEMALA\r'r-'...;/./rr'Fig. 25.-Distrihution of troglobitic amblypygids of the families Charontidae and Phrynidae: 1, Paraphrynus baeops;2, P. velmae; 3, P. chiztun; 4, P. chacmool; 5, P. reddelli; 6, Paraphrynus sp.; 7, Charontidae gen. et sp.132

Distribution.-Known only from caves in the Xilitlaand Aquismon regions, San Luis Potosi. See Fig.25.Records.-San Luis POtOSI; Sotano de Huitzmolotitlaand Sotano de Tlamaya.New record.-San Luis PotOSI; Cueva de San Rafael(det. C. Mullinex).Discussion.-This species has the median oculartubercle and median eyes completely missing.Order AraneaeThe spider fauna of the caves of Mexico is one ofthe richer in the world, both in number of species andin number of troglobites. The few collections of spidersfrom caves in Guatemala and Belize also givepromise of many troglobites. A total of 47 species oftroglobite has been collected from the caves of thisregion; an additional 252 species of troglophile, trogloxene,and accidental have been recorded from thecaves of these three countries (see Table 15).Suborder MygalomorphaeThe suborder Mygalomorphae is generally poorlyrepresented in the cave faunas of the world. Gertsch(1973b) lists only seven species of cave-adapted mygalomorph.It is especially interesting that four ofthese species are from caves in Mexico.Family BarychelidaeAt least one unidentified species of the barychelidgenus Zygapelma has been collected from caves inMexico and Guatemala. Chamberlin and Ivie (1938b)described Zygopelma meridana from Cueva de SanIsidro, Yucatan. The ecological status of the cavernicoleZygopelma is unknown. A single barychelid species,Traglothele coeca Fage, is a troglobite; it isknown from Grutas de Bellamar, Cuba.Family CtenizidaeThe trap-door spiders of the family Ctenizidae includea single species which has been found in cavesin Mexico. Several specimens of Cyclosamin (Chari·zaps) laricata (C. Koch) were obtained from burrowsbelow the entrance drop of Sotano de los Guacamayos,Tamaulipas; a single female was collected as itran along a mud bank in Sotano del Tigre, San LuisPotosi (Gertsch and Platnick, 1975).Family DipluridaeThe family Dipluridae includes, in addition to thetwo troglobites listed below, two other troglobiticdiplurids: Accola caeca Simon from caves in theTable I5.-Summary of cave inhabiting Araneae.MygalomorphaeBarychelidaeCtenizidaeDipluridaeTheraphosidaeAraneomorphaeAgelenidaeAmaurobiidaeAnyphaenidaeAraneidaeCaponiidaeClubionidaeCtenidaeDictynidaeDysderidaeFilistatidaeGnaphosidaeHahniidaeLeptonetidaeLinyphiidaeLycosidaeMysmenidaeNesticidaeOchyroceratidaeOecobiidaeOonopidaePholcidaePisauridaePlectreuridaeSalticidaeScytodidaeSegestriidaeSelenopidaeTelemidaeTengellidaeTetrablemmidaeTheridiidaeThomisidaeUloboridaeZodariidaeTotalTroglobiteso355o oooooooooo6ooo42o119oooooo1o1oooo47Other Species11221221151122114415113810416751242011o1128341252Philippine Islands and Trogladiplura lawryi Mainfrom Roaches Rest Cave, Australia.Euagrus anops GertschEuagrus anops Gertsch, 1973b:141, 142, 144, 145,fig. 4; Brignoli, 1974b:199, 200; Hoffmann,1976:16.Type-Iocality.-Cueva de la Porra, 5 km N Xilitla,San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 26.Discussion.-This is a pale, eyeless species knownonly from a single female. In addition to the twotroglobites listed here, several other species of Euagrusare known from caves in Mexico, only one of133

which has been described. Euagrus luteus Gertsch isabundant in caveS in the vicinity of Jalpan and Pinalde Amoles, Queretaro. It has small eyes and slightlyattenuate appendages and is probably a troglophile.Brignoli (1974b) described E. lynceus from Cuevadel Diablo, Huehuetenango, Guatemala; it is alsoa troglophiie. A species of troglobite remains to bedescribed from a cave in San Luis Potosi.Euagrus cavernicola GertschEuagrus cavernicola Gertsch, 1971a:47, 48-49; Reddelland Mitchell, 1971b:186, fig. 11; Brignoli,1973b:327; Gertsch, 1973b:142, 145; Mitchelland Kawakatsu, 1973a:671, 673; Reddell, 1973a:33, 38; Reddell and Elliott, 1973b:181; Brignoli,1974b: 199; Hoffmann, 1976:16.Type-locality.-Cueva de la Capilla, El Porvenir,13.5 km NW of Gomez Farias, Tamaulipas, Mexico.Distribution.-Known from three caves in theSierra de Guatemala, Tamaulipas. See Fig. 26.Records.-Tamaulipas: Cueva de la Capilla, HarrisonSinkhole, and Cueva de la Mina.Discussion.-This pale, eyeless species is comparativelycommon in the caves of the Sierra de Guatemala,although no fully mature specimens have yetMEXICOGUATEMALA/.//r·\,.... r".-'Fig. 26.-Distribution of troglobitic and troglophilic spiders of the suborder Mygalomorphae: 1, Euagrus cavernico1a;2, Schizopelma elliotti; 3, Euagrus n. sp.; 4, S. stygia; 5, E. anops; 6, E. luteus; 7, Schizopelma n. sp. 1; 8, S. reddelli; 9, Schizo·pelma n. sp. 2; 10, E. Iynceus.134

een collected. It is frequently found walking acrossmoist flowstone.Family TheraphosidaeTwo species of blind Schizopelma have been describedfrom Mexican caves and are discussed below.Large dark tarantulas are frequently found near caveentrances, but are probably only accidentals ortrogloxenes. Until the taxonomy of this difficultgroup is better understood they await study.Schizopelma reddelli GertschSchizopelma reddelli Gertsch, 197:3b:141, 142, 143144,146, fig. 1, 2b; Hoffmann, 1976:19.Type-Iocality.-Cueva del Nacimiento del Rio SanAntonio, Oaxaca, Mexico.Distribution.-Known only from two caves nearAcatl

Cicurina (Cicurelln) maya GertschCicurina maya Gertsch, 1977b:127-128, 129, fig. 86;Reddell, 1977b:230, 239,240,241, 269.Cicurina, eyeless species: Gertsch, 1977b:l03.Type-Iocality.-Actlin Tucil, 2 km S Muna, Yucatan,Mexico.Distrihution.-Known only from the type-locality.See Fig. 27.Discussion.-This small eyeless species is related toC. buwata of Texas. It is the only species of the genusknown south of Tamaulipas.Cicurina (Cicurusta) mina GertschCicurina mina Gertsch, 1971a:48, 108, 110, fig. 166167; Reddell and Mitchell, 1971b:185; Mitchelland Kawakatsu, 1973a:673; Reddell, 1973a:33,38; Hoffmann, ]976>~6.Cicurina (Cicurusta) mina: Hrignoli, 1972: 149-150,151-152, fig. 5(4J): Reddell and Elliott, 1973b:]81,184.Circurina mina: \Iitchell and KawakatslI, lCJ73a:671(erroneous spelling).Type-locality.-Cueva de la Capilla. 13.5 km NWof Comez Farias, Tamaulipas, \Iexico.I•-.... ,.\,. ///\~/(/IGUATEMALAI.... .., 'I i;-i- l ........ _!--.- --_.-. I( /- '" I \L,_,/ "\..,i ~....., i'-.. i----j ,-IIII .... u....../i\./)/'.//r\r'" ..--Fig. 27.-Distribution of troglobitic and troglophilic spiders of the genus CiclJnna: 1, CiclJrina coahlJila; 2, C. mina; 3, C.iviei; 4, C. maya; 5, CiclJnna spp.136

Distribution.-Known from two caves in the Sierrade Guatemala, Tamaulipas. See Fig. 27.Records.-Tamaulipas: Cueva de la Capilla andCueva de la Mina.Discussion.-This species is very closely related toC. iviei Gertsch, a troglophile known from Cueva dela Mina and Harrison Sinkhole, Tamaulipas. Brignoli(1972) found few differences, other than eyelessness,between the two species and is apparently not convincedthat they are specifically distinct.TegenariaTegenaria blanda Gertschblanda Gertsch, 1971a:105-106; Reddelland Mitchell, 1971b:185; Reddell, 1973a:38;Brignoli, 1974b:230;Hoffmann, 1976:47.Type-locality.-Cueva de la Capilla, 13.5 km NWof Gomez Farias, Tamaulipas, Mexico.Distrihution.-Known only from the type-locality.See Fig. 28.Discussion.-This is a pale orange, long-legged specieswith reduced eyes.Tegenaria caverna GertschTegenaria caverna Gertsch, 1971a:105, 106·107, fig.158-161; Roth and Brame, 1972:3; Reddell,1973a:38; Brignoli, 1974b:230; Hoffmann, 1976:47.MEXICO,,II,1_.....,\ ///'\ //, /'../IIIIGUATEMALA" I'-\.._i'-...... '\'-..i ,,.._._-_..-"ii\../III:iL.j .....i_._.-.-. IiIiiiL,... ,..."'.--//.( ./Fig. 28.-Distribution of troglobitic and troglophilic spiders of the genus Tegenaria: 1, Tegenaria gertschi; 2, T. blanda;3, T. selva; 4, T. decora; 5, T. caverna; 6, T. TOthi; 7, T. tlaxcala; 8, T. mexicana; 9, T. florea.137

Type-Iocality.-Cueva del Puerto del Leon, 6.5 kmSE of Rio Blanco, Queretaro, Mexico.Distribution.-Known only from three caves inQueretaro. See Fig. 28.New records.-Quenitaro: Cueva de los Otates andCueva de Penalta (det. W.]. Gertsch).Discussion.-This pale, orange-brown species hasthe eyes greatly reduced in size. It is apparentlyclosely related to T. blanda.Family AmaurobiidaeThe family Amaurobiidae includes one specieswhich has been identified from a cave in Mexico.Nicholas (1962) lists Titanoeca tizamina Chamberlinand Ivie from MuruztUn Cave, Yucatan, as a troglobite.This species, now known as Goeldia tizamina,shows no modifications for a cave existence andshould be considered a troglophile. It is widespreadin caves in Yucatan. An undescribed species of Titanoecais known from Cueva Languin, Alta Verapaz,Guatemala.Family AraneidaeAlthough 15 species of the family Araneidae havebeen recorded from caves, none appears to be particularlyadapted for cave life, and most are probablyaccidentals. Nicholas (1962) has included Leucageloltuna Chamberlin and Ivie from Actun LoltUn, Yucatan,as a troglobite; this species, however, is probablya troglophile. The genus Azilia is frequentlyfound in Mexican caves. Azilia affinis O. P.Cambridge has been collected in caves in Chiapas andVeracruz, while A. vagepicta Simon is common incaves of the Sierra de Guatemala, Tamaulipas. Otherundetermined specimens belonging to Azilia havebeen collected from caves in Oaxaca, San Luis Potosi,and Veracruz.Family ClubionidaeMembers of the family Clubionidae are frequentlytaken in caves, but few species have yet been identified.The genera most commonly found are Corinnaand Phrurotimpus. Corinna saga F. P.-Cambridge hasbeen found in several caves in Yucatan. Tixcocobamaya Gertsch is known from several caves in northernYucatan and from epigean sites in Campeche andYucatan (Gertsch, 1977b).Family CtenidaeThe family Ctenidae is an abundant element of thecave fauna of much of Mexico. Only one species hasbeen described, but several others doubtless occur.Ctenus mitchelli Gertsch is a large, interesting speciesfrom San Luis Potosi and Tamaulipas. It is found inall parts of caves but may be seen most often on thewalls of dark rooms near the cave entrance. It probablyshould be classified as a troglophile.Family FilistatidaeFive species of the family Filistatidae are knownto occur in Mexican caves, but only one is of particularinterest. An undescribed species of Filistatoideshas been found in numerous caves in Nuevo Leon andCoahuila. It is probably a troglophile.Family GnaphosidaeNicholas (1962) listed the gnaphosid Zelotes mayanusChamberlin and lvie from Actun Sabaca, Yucatan,as a troglobite. This species shows no signs ofadaptation for a cave existence. Other gnaphosid speciesknown from caves in !\lexico are Z. rusticus (L.Koch) from Cueva del Leon, Coahuila, and Drassodespallidipalpis (Bilimek) from Grutas de Cacahuamilpa,Guerrero.Family LeptonetidaeThe family Leptonetidae has contributed fivetroglobitic and five troglophilic species to the cavefauna of Mexico (see Fig. 29). One of the troglophiles,Archoleptoneta obscura Gertsch, from Cuevadel Tio Ticho, Chiapas, is a primitive leptonetid mostclosely related to species from California and Texas(Gertsch, 1974). Brignoli (1974b) described Seoleptonetaarganoi from Cueva Grande de San A~lIstln,Chiapas; this species has heen removed to Archoleptonetaby Brignoli (1977). The remaining troglophilicleptonetds all belong in lilt: genus Leptoneta.Leptoneta bonita Gertsch was described from CuevaBonita, Tamaulipas. Leptoneta pecki Gerlsch isknown only from Grutas de San Barlolo, 0lllevoLeon. Leptoneta rainesi Gertsch is a species knownfrom caves in the Sierra de El Abra, Tamaulipas.Leptoneta capilla GertschLeptoneta capilla Gertsch, 1971a:48, 52, 53, fig. 1-2;Reddell and Mitchell, 1971b: 186; Brignoli, 1972:134; Mitchell and Kawakatsu, 1973a:673; Reddelland Elliott, 1973b:181, 184; Gertsch, 1974:145,147, 177, 178, 179, 180, fig. 82,84,92,115-116,119; Hoffmann, 1976:81.Leptoneta sp.: Reddell and Mitchell, 1971b: 186(Cueva de la Mina record only).Neoleptoneta capilla: Brignoli, 1972:134, 135-136,137, 139, fig. 1(6-10); Reddell, 1973a:33, 38;Hoffmann, 1976:21, 81; Brignoli, 1977:216;Brignoli, 1979a:233, fig. 4.Type-Iocality.-Cueva de la Capilla, 13.5 km i\Wof Gomez Fari~s, Tamaulipas, Mexico.138

actually a natural cave artificially enlarged by mercury(not iron) miners.Leptoneta isolata GertschLeptoneta: Reddell, 1967c:55.Leptoneta isolata Gertsch, 1971a:48, 51, 52, 53, fig.9-11; Gertsch, 1974:145, 178,179,180-181, fig.94,107-108,123; Hoffmann, 1976:8l.Neoleptoneta isolata: Brignoli, 1972: 137; Reddell,1973a:33, 38; Hoffmann, 1976:21, 81; Brignoli,1977:216.Type.locality.-Grutas de Garcia (=Grutas de Villade Garcia), Nuevo Leon, Mexico.Distribution.-Known only from caves in NuevoLeon and Tamaulipas. See Fig. 29.New record.-Tamaulipas: Cueva de California (det.W. J. Gertsch).Discussion.-This is a pale, essentially eyeless spe·cies related to the troglophile 1. pecki.Leptoneta limpida GertschLeptoneta limpida Gertsch, 1974:166, 174-175, fig.60,79; Hoffmann, 1976:113; Reddell, 1977a:90.Neoleptoneta limpida: Brignoli, 1977:216; Brignoli,1979:437.Type-Iocality.-Cueva de los Riscos, 4 km S Mapimi, Durango, i\'lexico.Distribution.-Known only from the type-locality.See Fig. 29.Discussion.-This pale, long-legged cavernicole hasreduced eyes. It is more closely related to the caveinhabiting species of the genus from Central Texasthan to the Mexican cave species. It was found insmall webs at the junction of wall and floor in a smallside room near the end of the cave.Leptoneta reclusa GertschLeptoneta reclusa Gertsch, 1971a:53-54, fig. 12-13;Gertsch, 1974:178, 179, 184, fig. 86, 95, 111-112,117; Hoffmann, 1976:82.Neoleptoneta reclusa: Brignoli, 1972:137; Reddell,1973a:38; Hoffmann, 1976:21, 82; Brignoli,1977:215.Type-Iocality.-Cueva de Chorros de Agua, 20 kmW of Montemorelos, Nuevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 29.Discussion.-This pale species with reduced eyes ismost closely related to 1. capil/a.Family LinyphiidaeAlthough nine species of linyphiid have been collectedfrom caves in this area, none show any signs ofadaptation for a cave existence. Erigone tamazunchalensisGertsch and Davis has been found in severalcaves in San Luis Potosi and Puebla; Jalapyphantesis represented by at least two species from caves inMexico and Guatemala; and the genus Meioneta includesseveral species in caves throughout Mexico.Family MysmenidaeThe family Mysmenidae is represented in Mexicancaves by seven species of the genus Maymena(Gertsch, 1960, 1971a, 1973a; Brignoli, 1974) (seeFig. 30). One of these, M. mayana (Chamberlin andIvie), ranges south from Oaxaca into Belize and Guatemala,where it also inhabits caves. Some specimensfrom caves have reduced eyes. The remaining six speciesare known only from caves in 'Mexico: Jl'1. cascadaGertsch from Oaxaca and Veracruz; M. chica Gertschfrom Nuevo Leon, San Luis Potosi, and Tamaulipas;M. delicata Gertsch from Oaxaca and Veracruz; M.grisea Gertsch from Tamaulipas; IH. misteca Certschfrom Guerrero and Oaxaca; and M. sbordonii Hrignolifrom Chiapas. Undescribed species are known fromcaves in Campeche, Oaxaca, Puehla, and Veracruz.Although Nicholas (1962) lists M. misteca and M.mayana as troglobites, no species of the genus showsextreme adaptations for a cave existence and allshould probably be considered troglophiles.Family NesticidaeAmong the more important elements in the cavefauna of Mexico are the spiders of the family Nesticidae.Species of the genera Nesticus, Eidmannel/a,and Gaucelmus are frequently found in caves in thisregion. Gertsch (in press) has revised the family inNorth America, Central America, and the West Indies.Two species of described Nesticus are considered hereto be troglobites, and an additional two species aretroglophiles: Nesticus hoffmanni Gertsch from cavesin Hidalgo and N. vazquezi Gertsch from caves inQueretaro (see Fig. 31). Five troglophilic speciesfrom caves in Queretaro, Oaxaca, and Tamaulipas remainundescribed. Eidmannel/a pal/ida (Emerton)ranges south from the United States into Guatemalaand has been recorded from essentially every state inMexico for which we have cave collections (see Fig.32). An undescribed troglophile of the genus isknown only from Cueva de El Pachon, Tamaulipas.The genus Gaucelmus is represented by three species:G. calidus Gertsch ranges north from Huehuetenango,Guatemala, into Hidalgo, Mexico; G. augustinus Keyserlingis a troglophile in caves from Guatemala intothe United States; G. strinatii Brignoli was recentlydescribed from Cueva Chirrepeck, Alta Verapaz,Guatemala (Brignoli, 1979b).140

Nesticus arganoi BrignoliNesticus arganoi Brignoli, 1972: 148-149, fig. 5(40.42); Hoffmann, 1976:39.Type-locality.-Cueva de Ojo de Agua de Tlilapanno. 2 (=Cueva Macinga), Orizaha, Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 3l.Discussion.-This is the only described eyelessNesticus from Mexico. It appears to be most closelyrelated to N. nahuanus. A second eyeless species ofNesticus has heen collected recently in a cave nearZoquitlan, Puehla.Nesticus nahuanus GertschNesticus nahuanus Gertsch, 1971a:99-100, fig. 151,156; Brignoli, 1972:149; Reddell, 1973a:38;Hoffmann, 1976:39.Nesticus nahnanus: Gueorguiev, 1974:682 (erroneousspelling).Type-locality.-Cueva de la Boca, 6 km SE ofSantiago, Nuevo Leon, Mexico.Distribution.-Known from caves in Nuevo Leon.See Fig. 3l.Records.-Nuevo Leon: Small caves (Cuesta deChipinque), Cueva de la Boca, and Resumidero delPahlillo.(/GUATEMALAr/.//r\r'.-'--'Fig. 30.-Distribution of troglophilic spiders of the family Mysmenidae: 1, Maymena chica; 2, M. grisea; 3, M. misteca;4,M. delicata; S,M. mayana; 6,M. cascada; 7,M. sbordonii.141

Discussion.-This large species has the eyes reducedto small, evanescent spots in most individuals.Although not as highly cave adapted as N. arganoi,this species is almost certainly committed to a caveexistence.Family OchyroceratidaeThe tropical family Ochyroceratidae includes twoapparent troglobitic and three troglophilic species(see Fig. 34). Ochyrocera fagei Brignoli is knownfrom Cueva del Panteon, Chiapas; O. formosa Gertschwas described from Gruta del Silvino, Izabal, Guatemala;and an undescribed species of Ochyrocera hasrecently been collected in Grutas de Balankanche,Yucatan. In addition to the two troglobites discussedbelow, cave-adapted species of this family are knownfrom Jamaica and the Hawaiian Islands.Theotima martha GertschTheotima martha Gertsch, 1977b:126; Reddell,1977b:232, 239, 240, 291. Brignoli, 1979c:597.Type-locaJity.-Cueva Sodzil, 3 km W Sucopo,Yucatan, Mexico.Distribution.-Known only from the type.localityand possibly one cave in Quintana Roo. Sec Fig. 34.GUATEMALA//.r\r·-- ..-"/Fig. 31.-Distribution of troglohitic and troglophilic spiders of the genera Eidmannella and Nesticus: 1, Nesticus nahuanus;2, Nesticus n. sp. 1; 3, Eidmannella n. sp.; 4, Nesticus n. sp. 2-3; 5, N. hoffmanni; 6, N. vasquezi; 7, Nesticus n. sp. 4; 8, N.arganoi; 9,Nesticus n. sp. 5; 10, Nesticus n. sp. 6.142

New record.-Quintana Roo: ?Cueva de la Abispa(det. W. J. Gertsch).Discussion.-This is a pale yellow species with longlegs and reduced eyes. It is closely related to T. ruinaGertsch from Las Ruinas de Palenque, Chiapas.Theotima pura GertschTheotima pum Gertsch, 1973b:141, 155; Reddell andElliott, 1973b: 181, 184; Hoffmann, 1976:24;Shear, 1976:249; 'Gertsch, 1977b:125; Brignoli,1978:15,16. Brignoli, 1979c:597.Type-Iocality.-Cueva de los Vampiros, 20 km NWEI Limon, Tamaulipas, Mexico.Distribution.-Known only from the type-locality.See Fig. 34.Discussion.-This species is known only by a singleeyeless female. It is the first, and only eyeless, speciesof the family known from Mexican caves.Family OonopidaeOne troglobitic oonopid, Oonops coecus (Chamberlinand Ivie), is known from Mexico. Other cavedwellingoonopids of interest include O. chickeringiBrignoli from Cueva del Panteon, Chiapas; O. mitchelliGertsch from ActUn Xpukil, Yucatan; O. reddelliGertsch from ActUn Tucil, Yucatan; Triaeris lacandonaBrignoli from Cueva de Yaxchilan, Guatemala:,~/,GUATEMALA•..:.,.. ././/(',.' "Fig. 32.-Distribution of the troglophilic nesticid spider EidmannelIa pal/ida.143

and T. patellaris Bryant from caves in Campeche,Oaxaca, Quintana Roo, Tabasco, San Luis Potosi,and Veracruz, Mexico. and Alta Verapaz, Guatemala.Oonops coecus (Chamberlin and Ivie)Wanops coecus Chamberlin and Ivie, 1938b: 123, 125,fig. 1-2; Pearse, 1945:155; Nicholas, 1962:181;Gertsch, 1971a:47, 48, 56; Reddell, 1971b:31;Brignoli, 1972:153; Brignoli, 1973b:328;Gertsch,1973a:164; Hoffmann, 1976:21; Gertsch, 1977b:103,120; Reddell, 1977b:232.Oonops coecus: Gertsch, 1977b: 120, 121, fig. 64-66;Reddell, 1977b:232, 240, 241, 245, 246, 253,259,260,279,281,285.Type-Iocality.-Balaam Canche Cave (=Grutas deBalankanche), Chichen Itza, Yucatan, Mexico.Distribution.-Known from caves in the YucatanPeninsula, Mexico. See Fig. 35.Records.-Campeche: Actun Huachap and Grutasde Xkalumkin; Quintana Roo: Cenote de Santo Domingo;Yucatan: Grutas de Balankanche and CuevaEscondida.New record.-Quintana Roo: Cenote de Tos Virlol(det. W. J. Gertsch).Discussion.-This eyeless species is closely relatedto O. mitchelli and O. reddelli. It is usually found beneathsmall rocks on silt.., ., .,\)\\.'-MEXICOGUATEMALAFig. 33.-Distribution of troglophilic nesticid spiders of the genus Gauce/mus: 1, Gauce/mus augustinus; 2, G. co/idus; 3, G.augustinus and G. ca/idus; 4, G. strinotii.144

Family PholcidaeWithout doubt, the most important family thathas invaded caves in Mexico and Central America isthe Pholcidae. A total of 94 species of pholcid isalready known to inhabit caves in this region; 19 ofthese are troglobites.Seven described species of the genus Coryssocnemishave been identified from caves in Mexico(Gertsch, 1971a, 1973b; Brignoli, 1974b): C. abernathyiGertsch from San Luis Potosi, Tamaulipas,Oaxaca, and Puebla; C. clarus Gertsch from Tamaulipas;C. facetus Gertsch and C. pecki Gertsch fromChiapas; C. iviei Gertsch from Hidalgo, Queretaro,and San Luis Potosi; C. placidus Gertsch from Veracruz;and C. simoni O. P.-Cambridge from Coahuila,Hidalgo, Nuevo Leon, Oaxaca, and Queretaro. Severalundescribed species have been collected recently.These spiders are usually found hanging in webs,usually in the vicinity of the cave entrance. See Fig.36 for the distribution of the species of this genusin Mexican caves.The genus Physocyclus is known from cavesthroughout much of Mexico (Gertsch, 1971a, 1973b;Brignoli, 1974b), although many species appear to be1 2••MEXICOGUATEMALAFig. 34.-Distribution of troglophilic and troglobitic spiders of the families Ochyroceratidae and Telemidae: I, Theotimapura; 2, Matta sbordonii; 3, Ochyrocera fagei; 4, M. mckenziei; 5, Theotima n. sp.; 6, Ochyrocera n. sp.; 7, Theotima martha;8, Telema mayana; 9, o. formosa.145

estricted to the Mexican Plateau (see Fig. 37). Nicholas(1962) lists P. hoogstrlUlli Gertsch and Davis as atroglobite, but it like all species of the genus is presumablya troglophile. Physocyclus is also usuallyfound hanging in webs in the vicinity of the cave entrance.Although its distribution overlaps that ofCoryssocnemis, the genus seems to be more commonlyfound in caves in the arid regions of Mexico. Elevenspecies have been reported from Mexican caves: P.bicornis Gertsch and P. modestus Gertsch from Guerrero;P. enaulus Crosby from Chihuahua and Coahuila;P. globosus (Taczanowski) from Chiapas, SanLuis Potosi, Veracruz, and Yucatan; P. hoogstraaliGertsch and Davis from Coahuila and Nuevo Leon; P.lautus Gertsch and P. validus Gertsch from Colima; P.merus Gertsch from San Luis Potosi; P. pedregosusGertsch from Coahuila; P. reddelli Gertsch from Hidalgo,Queretaro, and San Luis Potosi; and P. tanneriGertsch from Sonora.The genus Modisimus includes 12 described specieswhich have been identified from caves in Mexico andGuatemala (Gertsch, 1971a, 1973b; Brignoli, 1974b):M. beneficus Gertsch from Veracruz; M. bonetiGertsch and M. texanus Banks from San Luis Potosiand Tamaulipas; M. iviei Gertsch from Quintana Roo,Tabasco, and Yucatan; M. mckenziei Gertsch, M.mitchelli Gertsch, and M. reddelli Gertsch from Tamaulipas;M. propinquus O. P.-Cambridge and M.MEXICOGUATEMALA/r·\,....".-'//Fig. 35.-Distribution of troglophilic and troglobitic spiders of the family Oonopidae: 1, Triaeris patellaris; 2, Oonopschi£keringij 3, O. coeCUSj 4, O. mitchellij 5, O. reddelli; 6, T. lacandonus.146

tzotzile Brignoli from Chiapas; M. pusillus Gertschfrom Nuevo Leon; M. rainesi Gertsch from NuevoLeon and Tamaulipas; and M. gracilipes Gertsch fromAlta Verapaz, Guatemala. Three undescribed speciesare known from caves in Veracruz. All of the cavernicolespecies of the genus are presumably troglophiles.These spiders are found in all parts of the caves,where they hang in delicate webs along the walls oramong speleothems. See Fig. 38 for the distributionof this genus in the caves of the region.The best-represented genus of spider in the cavesof Mexico, Guatemala, and Belize is Metagonia (seeFigs. 39-41). Eight species are considered to he troglobitesand are discussed below. The remaining 17species are troglophiles (Chamberlin and Ivie, 1938b;Gertsch, 1971a, 1973h, 1977h; Brignoli, 1972,1974b): M. amica Gertsch, M. pasquinii Brignoli, andM. punctata Gertsch from San Luis Potosi; M. candelaGertsch and M. serena Gertsch from NuevoLeon; M. capilla Gertsch, M. secreta Gertsch, and M.suzanne Gertsch from Tamaulipas; M. coahuilaGertsch from Coahuila; M. iviei Gertsch from QuintanaRoo and Yucatan; M. mcnatti Gertsch fromChiapas; M. maximiliani Brignoli from Queretaro; M........ " .. ././"~ /I--~"~ "I\I$,\ ""'-v ~I$/,/'.2,--4'..J--'-' .2'_ .3MEXICO~, //I/), /\/IIII\\, ././GUATEMALAI/i(,)I• i....-'-.'-."'\...,ii: Iu· ..../-'-'-' IIIii,... ..-'i1-/.//r\r'Fig. 36.~Distribution of troglophilic pholcid spiders of the genus Coryssocnemis: 1, Coryssocnemis simoni; 2, C. abernathyi;3, C. c1arus; 4, C. iviei; 5, C. iviei and C. simoni; 6, Coryssocnemis n. sp. 1; 7, Coryssocnemis n. sp. 2; 8, Coryssocnemisn. sp. 3; 9, C. placidus; 10, Coryssocnemis n. sp. 4; 11, C. facetus; 12, C. pecki.147

maya Chamberlin and Ivie from Campeche, QuintanaRoo, and Yucatan; M. placida Gertsch from NuevoLeon and Tamaulipas; M. tinaja Gertsch from SanLuis Potosi and Tamaulipas; M. yucatana Chamberlinand Ivie from Campeche and Yucatan; and At. blandaGertsch from Izabal and Alta Verapaz, Guatemala.Undescribed troglobites and troglophiles are knownfrom caves in Puebta and Veracruz.The genus Pholcophora is also well represented inthe caves of Mexico and Guatemala (Chamberlin andIvie, 1938b; Gertsch, 1971a, 1977b) (see Fig. 42).Three species are considered to be troglobites and arediscussed below. The remaining nine species are troglophiles:P. bispinosa Gertsch and P. evansi Gertschfrom Chiapas; P. bolivari Gertsch and P. mitchelliGertsch from Tamaulipas; P. elliotti Gertsch from SanLuis Potosi; P. maria Gertsch from Yucatan; P. speophila(Chamberlin and Ivie) from Campeche andYucatan; P. troglodyta Gertsch from Veracruz; andP. quieta Gertsl:h from Izabal, Guatemala. Undescribedspecies of the genus, including four troglobites,are known from caves in Oaxaca, Puebla, Veracruz,and Campeche.The genus Psilochorus is known from caves inMexico from near the border of the United States toGuerrero and Oaxaca (see Fig. 43). Two species aretentatively considered to be troglobites and are discussedbelow. The remaining six species are probableGUATEMALA..--'-'-'-'I/i\.I),.. ..-"/.//r\r·Fig. 37.-Distribution of troglophilic pholcid spiders of the genus Physocyclus: 1, Physocyclus enaulus; 2, P. hoogstTaali;3,P. pedregosus; 4,P. merus; 5,P. reddelli; 6,P. globosus; 7, P. lautus and P. validus; 8, P. modestus; 9, P. bicomis.148

troglophiles: P. concinnus Gertsch from San LuisPotosi; P. cordatus (Bilimek) and P. tellezi Gertschfrom Guerrero; P. fishi Gertsch from Hidalgo; P.murphyi Gertsch from Oaxaca; andP. russelli Gertschfrom Coahuila. An undescribed species is a probabletroglophile in the caves of Veracruz.Metagonia atoyacae GertschMetagonia atoyacae Gertsch, 1971a:48, 87-88, fig.123, 129; Gertsch, 1973b:152; Reddell, 1973a:33,38; Hoffmann, 1976:25.Type-Iocality.-Grutas de Atoyac, Atoyac, Veracruz,Mexico.Distribution.-Known only from two caves inVeracruz. See Fig. 39.New record.-Veracruz: Grutas de Abejas (det.W. J. Gertsch).Discussion.-This pale, eyeless species is related toM. tlamaya Gertsch, M. pura Gertsch, and othermembers of the tinaja group. All of the troglobiticspecies of Metagonia form a single close-knit groupof species. It is interesting to note that six of thetroglobites are from lowland tropical caves.I,r,,.\ I'' ......./ ;GUATEMALA,.\.... ,..--/.17 ,/\I

Metagonia chiquita GertschMetagonia chiquita Gertsch, 1977b:l03, 105, fig. 3132; Reddell, 1977b:232, 239, 240, 279.Type-Iocality.-Cenote Chen Mul, Ruinas de Mayapan,Yucatan, Mexico.Distribution.-Known only from the type-locality.See Fig. 39.Discussion.-This small, eyeless species was takenfrom beneath a rock on bat guano in an inner chamberof Cenole Chen Mul.Metagonia jarmila GertschMetagonia jarmila Gertsch, 1973b:141, 147, 154, fig.24-26.Type-Iocality.-Bucks Bypass Cave, Caves Branch,Belize.Distribution.-Known only from two caves ncarCaves Branch, Belize. See Fig. 39.Records.-BELIZE: Cayo District: Bucks BypassCave and St. Hermans Cave.Discussion.-This is a pale, essentially eyeless speciesrelated to 11-1. tinaja. It is the only troglobitic spiderknown from Belize.Metagonia martha GertschMetagonia martha Gertsch, 1973b:141, 147, 153·154, fig. 27; HoHmann, 1976:25.MEXICOI /t\.//GUATEMALAr\..... r·.,.:././/Fig. 39.-Distribution of troglobitic pholcid spiders of the genus Metagonia: 1, Metagonia pura; 2, M. pachona; 3, M. tlomaya;4, Metagonia n. sp. 1;5, M. atoyacae; 6, Metagonia n. sp. 2; 7, M. martha; 8, M. torete; 9,M. chiquita; 10, M. jarmila.150

Type-locality.~Cueva del Nacimiento del Rio SanAntonio, 10 km SSW Acathin, Oaxaca, Mexico.Distribution.-Known only from the type-locality.See Fig. 39.Discussion.-This pale species with rudimentaryeyes is related to M. tinaja.Metagonia pachona GertschMetagonia pachona Gertsch, 1971a:88, fig. 117, 124;Reddell and Mitchell, 1971a: 147; Reddell, 1973a:38; Hoffmann, 1976:25.Type-locality.-Cueva de EI Pachon, 7.5 km NE ofAntiguo Morelos, Tamaulipas, Mexico.Distribution.~Known only from the type-locality.See Fig. 39.Discussion.-This pale, small-eyed species is tentativelyconsidered to be a troglobite. It is related to M.tinaja.Metagonia pum GertschMetagonia pum Gertsch, 1971a:48, 87, fig. 108.109,122; Reddell and Mitchell, 1971b:187; Brignoli,1972:137, 138-139, fig. 2(16-18); Reddell, 1973a:33, 39; Reddell and Elliott, 1973b:184; Hoffmann,1976:25.MEXICO/'/GUATEMALA;.10 ,..II i>,_I~ ~l-----Lf/. I\.-' !I/i\. .t)'-.'-."i..,-----'IiiL/..//r\r·r-..-"Fig. 40.-Distribution of 10 species of troglophilic pholcid spider of the genus Metagonia: 1, Metagonia candela; 2, M.serena; 3, M. placida; 4, M. capilla;.5, M. secreta; 6, M. tinaja; 7, M. punctata; 8, M. maximiliani; 9, M. amica; 1O,M. iviei.151

Metagonia capilla: Mitchell and Kawakatsu, 1973a:673 (error for M. pura); Reddell and Elliott,1973b:181 (error for M. pura).Type-Iocality.-Cueva de la Capilla, 13.5 km NWof Gomez Farias, Tamaulipas, Mexico.Distribution.-Known only from the type-locality.See Fig. 39.Discussion.-This pale, eyeless species is related toM. tlamaya.Metagonia tlamaya GertschMetagonia sp.: Reddell, 1967d:106; Reddell, 1971b:31 (Sotano de Tlamaya record only).Metagonia tlamaya Gertsch, 1971a:48, 83-84, 87,fig. liO-lil, 125; Reddell, 1973a:33, 39; Hoffmann,1976:25.Type.locality.~Sotano de Tlamaya, San Luis Potosi,Mexico.Distribution.-Known only from the type-locality.See Fig. 39.Discussion.-This pale, eyeless species is related toM. tinaja.Metagonia torete GertschMetagonia torete Gertsch, 1977b:105, 107,108-109,fig. 9, 25-26, 29-30; Reddell, 1977b:232, 240,241, 281, 292, 293.MEXICO4..4GUATEMALAii\i)/""_._._._.\r·,.....--,e/.//rFig. 41.-Distrihution of eight species of troglophilic pholcid spider of the genus Metagonia: 1, Metagonia coahuila; 2, M.suzanne; 3, M. pasquinii; 4, M. mcnatti; 5, M. maya; 6, M. yucatana; 7, Metagonia n. sp. 3; 8, M. blanda.152

Type-locality.-Cueva Sodzil, 5 km W Sucopo,Yucatan, Mexico.Distribution.-Known from caves in Campeche,Quintana Roo, and Yucatan. See Fig. 39.Records.-Yucatan: Cenote de Orizaba, Cueva deSodzil, Grutas de Tzab-Nah, and Cenote Xtacabiha.New records.-Campeche: Grutas de San Antonio;Quintana Roo: Cueva de Tancah (det. W. J. Gertsch).Discussion.-This is a small species with evanescenteyes and elongated legs.Pholcophora exigua GertschPholcophora exigua Gertsch, 1971a:80, 82, fig. 99100; Reddell, 1973a:39; Hoffmann, 1976:26;Reddell, 1977a:90.Type-locality.-Cueva de los Riscos, Sierra de laIndia, 6.5 km S of Mapimi, Durango, Mexico.Distribution.-Known only from the type-locality.See Fig. 42.Discussion.-This small, white species has the anteriormedian eyes obsolete and the lateral eyes reduced.It was collected from the remotest parts ofMEXICOGUATEMALAlb~//('\..... r".;.Fig. 42.-Distribution of troglobitic and troglophilic pholcid spiders of the genus Pholcophora: 1, Pholcophora exigua;2, P. mitchelli; 3, P. bolivari; 4, P. elliotti; 5, Pholcophora n. sp. 1; 6, Pholcophora n. sp. 2 and P. troglodyta; 7, Pholcophoran. spp. 3-4; 8, Pholcophora n. spp. 5-6; 9, Pholcophora n. sp. 7; 10, P. gruta; 11, P. bispinoso; 12, P. evansi; 13, Pholcophoran. sp. 8; 14, P. speophila; 15, P. pearsei; 16, P. quieta.153

the cave and in general association with Leptonetalimpida, Psilochorus delicatus, and Cryptocellus reddelli.Nicholas (1962) listed Spermophila (sic) speophilaChamberlin and Ivie from caves in Yucatan asa troglobite. This species, now placed in Pholcophora,should be considered a troglophile.Pholcophora gruta GertschPholcophora gruta Gertsch, 1971a:48, 78, fig. 95-96,137-138; Reddell, 1973a:33, 39; Brignoli, 1974b:223; Hoffmann, 1976:26.Type-Iocality.-Grutas de Juxtlahuaca, 6.5 km Nof Colotlipa, Guerrero, Mexico.Distribution.-Known only from the type-locality.See Fig. 42.Discussion.-This is a pale, eyeless species.Pholcophora pearsei (Chamberlin and Ivie)Anopsicus pearsei Chamberlin and Ivie, 1938b:123,130, 132, fig. 17-18; Pearse, 1945:156; Nicholas,1962:181; Gertsch, 1971a:47, 76; Reddell,1971b:31; Hoffmann, 1976:84; Gertsch, 1977b:103, ll2.Anopsicus: Pearse, 1945:155; Brignoli, 1972: 137,138,139; Brignoli, 1974b:219, 223.Pholcophora (Anopsicus) pearsei: Gertsch, 1971a:48.Pholcophora pearsei: Gertsch, 1971a:77; Brignoli,1974b:223; Hoffmann, 1976:26, 84; Gertsch,1977b:ll5, ll6, fig. 47-49; Reddell, 1977b:232,239,240,253,259,260,261.Type-Iocality.-Oxolodt Cave (=Cueva Oxolodt),Kaua, Yucatan, Mexico.Distribution.-Known from caves in Quintana Rooand Yucatan. See Fig. 42.Records.-Quintana Roo: Cenote de Las Ruinasand Cenote de Santo Domingo; Yucatan: Cueva Escondida,Actlin Kaua, and Cueva Oxolodt.Discussion.-This small, eyeless species is usuallyfound beneath rocks in moist areas.Psilochorus delicatus GertschPsilochorus delicatus Gertsch, 1971a:76, fig. 90-91;Hoffmann, 1976:27; Reddell, 1977a:90.Type-Iocality.-Cueva de los Riscos, Sierra deIndia, 6.5 km S of Mapimi, Durango, Mexico.Distribution.-Known only from the type-locality.See Fig. 43.Discussion.-This small pale species with reducedeyes is known only from a single female and two immaturespecimens. It was collected from a small sideroom near the end of the cave. Nicholas (1962)listed Pholcus cordatus Bilimek from Grutas de Cacahuamilpa,Guerrero, as a troglobite. This species,now placed in Psilochorus, is a troglophile.Psilochorus diablo GertschPsilochorus new species: Spieth, 1950:31.Psilochorus sp.: Reddell, 1971b:32 (Cueva del Diablorecord only).Psilochorus diablo Gertsch, 1971a:72, 73, fig. 77-81;Reddell, 1973a:39; Hoffmann, 1976:27; Reddell,1977a:85.Type-locality.-Cueva del Diablo, 1 km W of Salaices,35 km N of Parral, Chihuahua, Mexico.Distribution.-Known only from the type-locality.See Fig. 43.Discussion.-This small, pale species has reducedeyes and is probably a troglobite restricted to thecave habitat by the surrounding desert.Family ScytodidaeThe spiders of the scytodid genus Loxoscelesabound in the caves of Mexico, Guatemala, andBelize (Chamberlin and Ivie, 1938b; Gertsch, 1958,1973b) (see Fig. 44). A total of 13 species of thisgenus is known from caves: L. aranea Gertsch fromQueretaro; L. aurea Gertsch from Durango; L. belliGertsch from Coahuila; L. bolivari Gertsch and L.luteola Gertsch from Nuevo Leon; L. colima Gertschfrom Colima; L. devia Gertsch and l\Iulaik from Tamaulipas;L. misteca Gertsch from Guerrero; L. tehuanaGertsch from Chiapas; L. tenango Gertschfrom Hidalgo; L. valdosa Gertsch from San Luis PotOSIand Tamaulipas; L. yucatana Chamberlin andIvie from Campeche, Quintana Roo, and Yucatan,Mexico; Cayo District, Belize; and EI Peten, Guatemala;and L. guatemala Gertsch from Alta Verapaz,Guatemala. These spiders are usually found in drierareas of the cave. They live in small tangled websamong loose dry breakdown and are frequently presentin large numbers. Although several species"of thegenus Scytodes are known from Mexican caves, noneshow signs of cave adaptation. Nicholas (1962) liststwo Yucatan cave species, Scytodes itzana Chamberlinand Ivie and S. meridana Chamberlin and Ivie astroglobites; these species are probably troglophiles.Scytodes fusca Walckenaer is also frequently found incaves in Yucatan. Brignoli (1976) has reviewed theworld scytodid fauna.Family TelemidaeThe family Telemidae is known from southernEurope, Africa, the western United States, and Guatemala.A single species, Telema mayana Gertsch, is154

a troglohite in Guatemalan caves (see Fig. 34). Telemagracilis (Keyserling) from Alabaster Cave, California,is a troglophile.Telema mayana GertschTelema mayana Gertsch, 1973b: 141, 154-155; Shear,1976:249.Type-locality.-Cueva Sepacuite n. 2, Finca Sepacuite,Senahu, Alta Verapaz, Guatemala.Distribution.-Known only from the type-locality.See Fig. 34.Discussion.-The eyes in this species are reducedto vestiges.,.,"., .,,Family TetrablemmidaeMatta mckenziei ShearTetrablemma n. sp.: Reddell, 1977b:232, 240, 241,245,247.Matta mckenziei Shear, 1978:14, 16, 18, 19,20, fig.37-39.Type-locality.-Grutas de San Ignacio, Chencedro,15 km. N of Bolonchenticul, Campeche, Mexico.Distribution.-Known only from two caves innorthwestern Campeche. See Fig. 35.Records.-Campeche: ActUn Halmensura and Grutasde San Ignacio.)\,."-,,·.r--"' __ II,.MEXICOGUATEMALA/r\~, r//Fig. 43.-Distribution of troglobitic and troglophilic pholcid spiders of the genus Psilochorus: 1, PsilochoTUs diablo; 2, P.russelli; 3, P. delicatus; 4, P. concinnus; S,P. Fuhi; 6, P. cordatus and P. te/lezi; 7, Psilochorus n. sp.; 8, P. murphyi.155

DiscWlSion.-This is a delicate, eyeless species withlong legs. The genus Matta ranges from San Luis Potosi,Mexico, into Brazil. The only other species ofMatta known from caves is M. sboTdonii, which wasdescribed from an eyeless population in Cueva delOjo de Agua de Tlilapan, Veracruz. In a revision ofthe genus, Shear (1978) studied numerous specimensof M. sboTdonii from epigean and cave collectionsand found no significant differences in the genitaliabetween cave and surface populations. Some epigeanspecimens possess reduced lateral eyes. Matta SbOTdoniiis much darker and more robust than is M.mckenziei. In addition to the type-locality M.sbOTdonii is known from caves In Oaxaca, Tamaulipas,and Yucatan.Family TheridiidaeAlthough at least 24 species of theridiid spidershave been reported from caves in Mexico, most havebeen recorded only once and are probably accidentals.Nicholas (1962) lists Spelobion spukilum Chamberlinand Ivie as a troglobite. This species, nowplaced in the genus Thymoites, is probably a troglophile.The genus Achaearanea includes five specieswhich are frequently found in caves and which maybe troglophiles. They are found hanging in webs inMEXICO•GUATEMALA12•,....r·"'.-'"/.//r·Fig. 44.-Distrihution of troglophilic scytodid spiders of the genus Loxosceles: 1, Loxosceles belli; 2, L. bolivari; 3, L.aurea; 4, L. luteola; 5, L. valdosa; 6, L. devia; 7, L. aranea; 8, L. tenango; 9, L. colima; 10, L. misteca; 11, L. tehuana; 12, L.guatemala; unnumbered circles, L. yucatana.156

darkness along the cave wall. One of these is A.porteri Banks, a common troglophile in the caves ofCentral Texas. Several species of Theridion, Thymoites,and Tidarren have been occasionally collected,but their ecological status is not known.Family UIoboridaeThe genus Philoponelw is known from many cavesthroughout Mexico. Philoponella signatelw (Roewer)is an apparent troglophile in caves in Oaxaca, Puebla,and Yucatan. Philoponella semiplumosa (Simon) isknown from caves in Tamaulipas and Veracruz.Philoponelw vicina (0. P.-Cambridge) has been takenin several caves in Tabasco and Veracruz.Order RicinuleiFamily RicinoididaeThe order Ricinulei includes only one family andtwo living genera, Ricinoides in Africa and Cryptocellusin the New World. Cryptocellus ranges northfrom South America into southern Texas. In southernMexico it has been found in Chiapas, Guerrero, andYucatan; it occurs north along the Sierra Madre Orientalfrom Veracruz to Nuevo Leon, with isolatedcave populations in Durango. A single species, C.dorotheae Gertsch and Mulaik, is known from Texas.One cave species, C. cookei Gertsch, is known fromCueva Jobitzinaj, near Flores, EI Peten, Guatemala(Gertsch, 1977a). Ten Mexican species have beendescribed, of which two (c. spinotibialis Goodnightand Goodnight from Chiapas and C. gertschi Marquezand Conconi from Veracruz) a.re exclusively epigean.One species (c. pelaezi Coronado) is known fromboth cave and epigean sites in the Sierra de EI Abra,San Luis Potosi and Tamaulipas, and from epigeansites in the Sierra de Guatemala, Tamaulipas. Theremaining seven species are all known only fromcaves (see Fig. 45). Three of these are considered tobe troglobites and are listed below. The remainingfive species are either troglophiles or of uncertainecological status. An epigean species from NuevoLeon and a troglobitic species from Cueva de Valdosa,San Luis Potosi, await description.Chamberlin and Ivie (1938a) described the firstMexican cavernicole species, C. pearsei, from Grutasde Balankanche and Cueva Oxolodt, Yucatan. Thisspecies has since been found in nine additionalYucatan caves (Gertsch, 1977a). Cryptocellus bonetiwas described by Bolivar (1941) from Grutas deCacahuamilpa, Guerrero; it is also known fromGrutas de Acuitlapan, Guerrero (Coronado, 1970).Cryptocellus pewezi was described by Coronado(1970) from Cueva de Taninul n. 1, San Luis Potosi.This species has become the best known of allricin'uleids, due to the outstanding work of Dr.Robert W. Mitchell and his students at Texas TechUniversity. Pittard and Mitchell (1972) studied theextemal morphology of all life stages; Cooke (1971)studied the mating behavior; and Mitchell (1970)studied various aspects of the population dynamicsin Cueva de la Florida, Tamaulipas. Other studies,including the life cycle and the development of thethird leg, are in preparation. Gertsch (1971b) describedthree additional species of cavernicole ricinuleid.In addition to the troglobite, C. reddelli, headded C. mitchelli from Cueva del Guano, Durango,and C. bolivari from Sumidero del Camino andGrutas de Zapaluta, Chiapas. Brignoli (1974c) describedC. sbordonii from Chiapas and tentativelyreported C. bolivari from Cueva del Tio Ticho,Chiapas.Epigean ricinuleids are usually found beneathrocks along hillsides or under permanent groundcover. The cavemicolous species have been foundon guano, silt, or rarely beneath rocks. The presenceof C. pearse~ C. osorioi, and C. pewezi in vast numbershas proven that the order is not the incrediblyrare group of animals that it was once thought to be.Cryptocellus osorioi BolivarCryptocellus osorioi Bolivar, 1946:24-28, fig. 1-7;Bolivar, 1952:296; McKenzie, 1965a:36; Reddell,1967b:82; Reddell, 1967f:99; Barrera, 1968:309;Beck and Schubart, 1968:68,69,70,74; Kaestner,1968:207; Mitchell, 19691:137; Mitchell, 1969d:letter; Woolley, 1969: 151; Coronado, 1970:48,61; Mitchell, 1970b:64, 71-72, 73, 74; Pittard,1970:3,6, 10; Cooke, 1971:9, 17,18,19,20,21,22; Gertsch, 1971b:127; Reddell, 1971b:3, 34,frontispiece; Reddell and Mitchell, 1971a:148,fig. 8; Reddell and Mitchell, 197c:2; Pittard andMitchell, 1972:4, 6; Sbordoni and Argano, 1972:9, 11; Kawakatsu, 1973b:255; Reddell, 1973a:34;Reddell and Elliott, 1973a:174; Brignoli, 1974c:154, 155, 156, 157, 159,160,166,167-168,169,fig. lC; Marquez and Conconi, 1974:82, 83, fig.20; Vomero, 1974:345; Dumitresco and JuvaraBals, 1976:163; Mitchell et al., 1977:53.Cryptocoellus osorioi: Anonymous, 1942a:221 (erroneousspelling).Cryptocellus osorici: Dumitresco and Juvara-Bals,1976: 163 (erroneous spelling).Cryptocellus ossorioi: Dumitresco and J uvara-Bals,1976:164, 178 (erroneous spelling).157

Type-Iocality.-Cueva de Los Sabinos, Valles, SanLuis Potosi, Mexico.Distribution.-Known only from caves in theSierra de El Abra, San Luis Potosi and Tamaulipas.See Fig. 45.Records.-San Luis Potosi: Cueva de Los Sabinos,Cueva de Taninul n. 1, and Satano del Tigre; Tamaulipas:Cueva de la Florida, Grutas de Quintero, andSatano de El Venadito.New record.-Tamaulipas: Cueva de Diamante(det. W. J. Gertsch).Discussion.-This large species possesses attenuateappendages and is a probable troglobite. Unlike itscompanion species in some of these caves, C. pelaezi,it has never been collected from the surface despiteintensive collecting throughout the Sierra de El Abra.Brignoli (1974c) places it with a group of speciesadapted for cave existence but refrains from designatingany as troglobites. Cryptocellus osorioi is usuallyrare, but in Satano del Tigre it is present in enormousnumbers. In this cave it was found living onguano in an extremely humid, warm passage severalhundred meters long. In Cueva de la Florida it inhabitsparts of the cave not utilized by C. pelaezi andis only rarely seen.l"-J"------·-o,r '\......r-.' "\\\.,,,\r-"-Al.., "\ \" c. "\1'-'- .......-.("""'\.r,--_',.88••s""\,8., 8/\/IIII'2_6•'---L.__._,-" U',·I IiI"\.._1 .9 I'-i.... " i7 , .., I..7. ' Iri\/)iII:IGUATEMALAFig. 45.-Distribution of troglobitic and troglophilic species of the order Ricinulei: 1, Cryptocellus reddelli: 2, C. mitchelli:3, C. osorioi and C. pelaezi: 4, C. osorioi: 5, C. boneti: 6, C. sbordonii: 7, C. bolivari: 8, C. pearsei: 9, C. cookei.158

Cryptocellus reddelli GertschCryptocellus new species: Reddell, 1967f:99-100.Cryptocellus sp.: Reddell, 1971b:34 (Cueva de losRiscos record only).Cryptocellus reddelli Gertsch, 1971b: 127, 130, 132,fig. 8-12; Cooke and Shadab, 1973:6; Reddell,1973a:34, 39; Reddell, 1973c:53; Brignoli,1974c:160, 167, 168, 169; [\-larquez and Conconi,1974:81, 82, 83, fig. 20; Dumitresco andJuvara-Bals, 1976:173; Reddell, 1977a:90.Type-locality.-Cueva de los Riscos, Sierra de laIndia, four miles south of Mapimi, Durango, Mexico.Distribution.-Known only from the type-locality.See Fig. 45.Discussion.-This is the only unquestionable troglobitein the order. The extreme elongation of appendages,paleness, and habitat within the cave all pointto a degree of cave adaptation not found among otherricinuleids. The species has been taken only from thecomplex of small rooms at the limit of exploration inthe cave. The few specimens which have been foundwere taken from clay or moist rock. The only otherspecies of ricinuleid known from the desert regionsof Mexico is C. mitchelli from Cueva del Guano,Durango. It is a typical guanophile and is relativelyabundant on guano in the inner rooms of this largebat cave.Cryptocellus sbordonii BrignoliCryptocellus sbordonii Brignoli, 1974c: 159-160, 164,167, 169, fig. lB, 3; Sbordoni et aI., 1974:30,45,pI. 4(10) (nomen nudum).Cryptocellus prope sbordonii: Brignoli, 1974c:155,160, 162, fig. 10; Sbordoni et aI., 1974:29; Dumitrescoand Juvara-Bals, 1976:163, 164, 178.Type-Iocality.-Cueva de las Canicas, Rancho delCielito, Ocozocoautla, Chiapas, Mexico.Distribution.-Known only from the type-localityand possibly one other cave at Rancho del Cielito,Chiapas. See Fig. 45.Records.-Chiapas: Cueva de las Can leas and?Cueva del Cerro Brujo.Discussion.-The holotype is a male, whereas thesingle specimen known from Cueva de Cerro Brujois a female; they are probably conspecific. This specieshas the most elongate legs in the genus and is,therefore, presumed to be a troglobite.Order OpilionidaThe order Opilionida includes a total of 50 specieswhich have been reported from the caves of the areaTable 16.-Summary of cave inhabiting Opilionida.TroglobitesOther SpeciesCyphophthalmiSironidae 1 0LaniatoresCosmetidae 0 7Phalangodidae 13 25PalpatoresNemastomatidae 1 0Phalangiidae 0 3Total 15 35under consideration. Many groups of opilionids havenot yet been studied, so this list will eventually bemuch longer. In particular, many species of thefamily Phalangiidae remain to be identified. SeeTable 16.Suborder CyphophthalmiFamily SironidaeNeogovea mexasca ShearNeogovea mexasca Shear, 1977b:165, 166, 167, 168,172-175, fig. 16-24.Type-Iocality.-Cueva del Nacimiento del Rio SanAntonio, 10 km SSW of Acatlan, Oaxaca, Mexico.Distribution.-Known only from two caves nearAcatlan, Oaxaca. See Fig. 46.New record.-Oaxaca: Cueva de la Finca (det.W. A. Shear).Discussion.-This species, the only troglobiticcyphophthalmid known in the Western Hemisphere,is the only species of the suborder known betweenFlorida and Guyana. The genus Neogovea includesthree other species, one from Brazil and two fromGuyana. Neogovea mexasca was described from females;the study of recently discovered males mayresult in a new generic assignment, possibly Siro.Other troglobitic cyphophthalmids are known fromSouth Africa and central Europe. The type specimenswere found on bat guano deposited on red clay approximately2 km from the entrance to the cave.Specimens collected in Cueva de la Finca were takenfrom a similar situation about 1 km from the caveentrance. This species has reduced sclerotization andmuch lengthened appendages, and is pale red.Suborder LaniatoresThe suborder Laniatores is well represented in thecaves of Mexico and Belize. Thirteen species of troglobiteand 31 other species are known. The taxonomyof this group is unfortunately in a state of flux. Silhavy(1974, 1977) recognizes the family Stygnopsidaeand places several species of laniatorids in this159

family. Clarence and Marie Goodnight, however,place all of these species,in the family Phalangodidae.The latter family is used in this report, but the mostrecent generic allocations are used until a thoroughrevisionary study is made of this group.Family CosmetidaeThe family Cosmetidae includes no troglobites,but a total of seven cave inhabiting species are knownfrom Mexico and Belize (see Fig. 46). Goodnight andGoodnight (1973) described Cynorta guadalupensisfrom one cave in San Luis Potosi and C. jamesonifrom numerous caves in San Luis Potosi and Tamaulipas.Goodnight and Goodnight (1977) reported thepresence of Vonones compressus (Cambridge) andErginulus roeweri (Goodnight and Goodnight) fromcaves in Yucatan; E. bimaculata Goodnight andGoodnight from caves in Campeche and Yucatan;and E. serratipes (Cambridge) from Rio Frio Cave,Belize.Family PhalangodidaeThe most frequently found group of opilionids intropical American caves are those belonging to thefamily Phalangodidae (including the Stygnopsidae).Thirteen species of phalangodids are believed to be\"-" "-"-\)\",,-i,}-/I'\'\\\I" \.)." ~\\\..~, ",~-- -_.;,MEXICOr f -- \..'-- /-- \ )I,' .....1-- -'-IiI'. I,,- I 7"- I-'- '\'., I, !I/GUATEMALAIII\I.~/r\r"/Fig. 46.-Distribution of troglobitic and troglophilic opilionids of the families Cosmetidae and Sironidae: 1, Cynortajamesoni; 2, C. guadalupensi3; 3, Neogovea mexasca; 4, Erginulus bimaculata; 5, E. roeweri; 6, Vonones compressus; 7, E.serratipes.160

troglobites in the caves of Mexico and Belize. Theseare currently placed in the genera Caecoa, Cynortina,Hoplobunus, Mexotroglinus, Stygnomma, and Troglostygnopsisand are discussed below.Silhavy (1977) described three genera for speciescollected in caves in Chiapas (see Fig. 47). Arganotuswas described to include Phalangodinus macrochelisGoodnight and Goodnight; this species, previouslyknown from Ocosingo, Chiapas, was collected inCueva del Sabin, Chiapas. Akdalima vomeroi was describedfrom Cueva del Sabin, Chiapas. SbordoniLJwas described to include Paramitraceras paroulaGoodnight and Goodnight, an epigean species fromChiapas, and SbordoniLJ armigera n. sp. from Sotanode Malpaso, Chiapas. Silhary (1979) described Arganotusstrinatii from Cueva Chirrepeck, Alta Verapaz,Guatemala.Goodnight and Goodnight (1971) described fivenew species of the genus Karos from Mexican caves:K. depressus from Sumidero del Llano Conejo, Queretaro;K. gratiosus from caves in the Xilitla andAgua Buena regions of San Luis Potosi and from acave near Huautla de Jimenez, Oaxaca; K. parousfrom caves in the Sierra de EI Abra, Tamaulipas, andthe Micos and Valle de los Fantasmas regions of SanLuis Potosi; K. projectus from Cueva del AhuateMEXICOGUATEMALA/.//rr·'-.".-'Fig. 47.-Distribution of troglophilic phalangodid opilionids of the genera Pe/lobunus, Paramitraceros, Sbordonia, Arganotus,and Akda/ina: 1, Pe/lobunus mexicanus; 2, Paramitraceros granulotus; 3, Paramitraceros femora/is and Sbordoniaarmigera; 4, Arganotus macroche/is and Akdalina vomeroi; 5, Paramitraceros hispidu/us; 6, Arganotus strinatii.161

n. 2, San Luis Potosi; and K. rugosus from Cueva deOjo de Agua de Tlilapan, Veracruz. Silhavy (1974)reported the presence of K. rugosus in Cueva del Ojode Agua Grande, Veracruz, and described K. brignoliifrom Cueva Tlilapan, Veracruz. Karos dybasi (Goodnightand Goodnight) was reported from Cueva deEI Jobo, San Luis Potosi, by Goodnight and Goodnight(1973). See Fig. 48 for the distribution ofKaros in Mexican caves.Three species of the genus Paramitraceras havebeen identified from Mexican caves (see Fig. 47).Goodnight and Goodnight (1973) reported P. granulatusCambridge from Cueva del Guayabo, Oaxaca.Silhary (1974) identified P. hispidulus Cambridgefrom Cueva de la Golondrina, Chiapas. Paramitracerasfemoralis Goodnight and Goodnight was reoported by Silhary (1977) from Sotano de Malpaso,Chiapas. Pellobunus mericanus Goodnight and Goodnightis known from caves in Nuevo Leon and Queretaro(Goodnight and Goodnight, 1971, 1973) (seeFig. 47). Goodnight and Goodnight (1971) describedHoplobunus robustus from caves in the Tequila regionof Veracruz. This species is also known fromcaves in the Potrero region of Veracruz (Goodnightand Goodnight, 1973) (see Fig. 50). Silhavy (1974)removed this species to the genus Stygnopsis. This isa large distinctive species which is frequently abundanton cave walls and rocky floor areas.GUATEMALA/r\r'~ ...;.//Fig. 48.-Distribution of troglophilic phalangodid opilionids of the genus Karos: 1, Karos parow; 2, K. gratiosus; 3, K.depre8sus; 4, K. dybasi, K. gratio8Us, and K. projectus; 5, K. rugosw; 6, K. brignolii and K. rugosw.162

Caecoa arganoi Silhav-yCaecoa arganoi Silhav-y, 1974:176, 189, 191, fig. 6(38-41), 7(42).Type-locality.-Cueva de Coatepec, Coatepec Harinas,Mexico, Mexico.Distribution.-Known only from the type-locality.See Fig. 49.Discussion.-This eyeless species is the type andonly known species of the genus.Cynortina misteca Goodnight and GoodnightCynortina misteca Goodnight and Goodnight, 1977:142, 143, fig. 3a,3b.Type-locality.-Footprint Cave, 4 km southwest ofCaves Branch, Cayo District, Belize.Distribution.-Known only from the type-locality.See Fig. 49.Discussion.-This pale, eyeless species is mostclosely related to C. acanthotibialis Goodnight andGoodnight from Chiapas, Guatemala, and Belize.Cynortina minutus (Goodnight and Goodnight) wasdescribed from Resumidero del Rio San Geronimo,Guerrero.Hoplobunus apoalensis Goodnight and GoodnightHoplobunus apoalensis Goodnight and Goodnight,1973:84,85,86-87, fig 4-5 ..I -J........._._.,I '\..... ,r '\)-.. \\l".\( ',;>-.., ). \~ \\"-~ '"'---. '-",-,.....;...r-,er,:-../'!'; I) ,... " ' ,- I .-(~ :,-.,/" -,~ L-....-J-.JMEXICO,.1',~".......... '.3 _J' 7.... '~-/-:GUATEMALAI\.-'"-......"i......,iI .•'-------//\I\ i'./'-'-'-"-,I,IIiI/r,... ." r'.-'//Fig. 49.-Distribution of troglobitic opilionids of the family Nemastomatidae and of troglophilic and troglobitic phalangodidopilionids of the genera Stygnomma, Caecoa, Cynortina, and Mexotroglinus: 1, Ortholasma sbordonii; 2, Stygnommatuberculata; 3, Caecoa arganoi; 4, Cynortina minutus; 5, Mexotroglinus sbordonii; 6, Cynortina misteca and S. pecki.163

Type-locality.-Cueva de Apoala, Santiago Apoala,20 km N Asuncion Nochixtlan, Oaxaca, Mexico.Distrihution.-Known only from the type-locality.See Fig. 50.Discussion.-This species has very small eyes whichappear to have lost the retinas. It was collected fromwalls and silt hanks in a moist room above the streampassage.Hoplobunus boneti (Goodnight and Goodnight)Serrobunus boneti Goodnight and Goodnight, 1942:2-3, fig. 7-11; Goodnight and Goodnight, 1945:3-4; Bonet, 1946a:115.Opilion, troglobio: Anonymous, 1942a:221.Serrobunus: Goodnight and Goodnight, 1953:20.Hoplobunus: Goodnight and Goodnight, 1953:20.Hoplobunus boneti: Goodnight and Goodnight,1971:38,40, fig. 18-19; Mitchell, 1971a:392-394;Reddell, 1971b:35; Reddell and Mitchell, 1971a:149, fig. 21-22; Goodnight and Goodnight, 1973:87; Reddell and Elliott, 1973a:I71, 174; Reddelland Elliott, 1973b:181, 184; Silhary, 1974:176,178,182, fig. 1(5-8); Vomero, 1974:345;FernandezRuiz, 1976:716, 717; Goodnight and Goodnight,1976:662.MEXICO////II,,GUATEMALA, , , /, /'/ IIIIII: i\",.-, ~-"--L. ·_·-·LJ·I......·/[ ,,- ~ t 1,./ \..-' \12 '-. I• .12 '-', i'-"1iI L./r'r\r'.... '.--r/Fig. SO.-Distribution of troglobitic and troglophilic phalangodid opilionids of the genera Hoplobunus, Troglostygnopsis,and Stygnopsis: 1, Hoplobunus osorioi; 2, Troglostygnopsis inops; 3, H. mexicanus and T. inops; 4, H. boneti; 5, H. planus;6, T. queretarius; 7, Stygnopsis robustus; 8, H. mexicanus; 9, H. spinooculorum; 10, H. apoalensis; 11, H. oaxacensis; 12, T.anophthalma; 13, H. %ullinii.164

Hoplobonus boneti: Sbordoni et al., 1974:37 (erroneousspelling).Type-Iocality.-Cueva de Los Sabinos, San LuisPotosi, Mexico.Distribution.-Known from many caves in theSierra de El Abra, San Luis Potosi and Tamaulipas;and possibly from one cave in the Sierra de Guatemala,Tamaulipas. See Fig. 50.Records.-San Luis Potosi: S6tano de Coatimundi,Cueva de la Curva, S6tano de Matapalma, Cueva delos Monos, S6tano de Pichijumo, Cueva Pinta, S6tanode la Pipa, Cueva de Los Sabinos, S6tano de Soyate,Cueva de Tanchipa, S6tano del Tigre, S6tano de laTinaja, Cueva de Valdosa, and S6tano de Yerbaniz;Tamaulipas: ?Cueva del Remolino and S6tano de ElVenadito.New records.-San Luis Potosi: Cueva de los Cuatesand S6tano de las Piedras (det. C. J. Goodnight).Discussion.-This species is one of the more obviousspecies in the cave fauna of the Sierra de ElAbra. When present, it is usually abundant. It hasnever been found in the caves of the northern Sierrade El Abra, apparently reaching its northern limit ofdistribution in S6tano de El Venadito, immediatelynorth of the San Luis Potosi-Tamaulipas state line.The single record from a high elevation cave in theSierra de Guatemala (Cueva del Remolino) is a possibleerror in identification or curation; additionalcollection is needed to verify this record. Individualsof this species are occasionally found guarding eggsor young (Mitchell, 1971a). Although some specimenspossess eyes, others are eyeless, and in all theeyes are reduced and probably non-functional.Hoplobunus osorioi (Goodnight and Goodnight)Chinquipellobunus osorioi Goodnight and Goodnight,1944:1, 3, fig. 4-9; Bolivar, 1944:26; Goodnightand Goodnight, 1945:3; Reddell, 1967a:24.Chipinquelobunus osorioi: Bonet, 1946a: 115 (erroneousspelling).Chinquepellobunus: Goodnight and Goodnight,1953:20.Hoplobunus: Goodnight and Goodnight, 1953:20.Hoplobunus osorioi: Reddell, 1971b:35.Type-locality.-Gruta del Carrizal, Nuevo Le6n,Mexico.Distribution.-Known only from three caves inNuevo Le6n. See Fig. 50.Records.-Nuevo Leon: Cueva del Carrizal, Grutasdel Palmito, and Grutas de Villa de Garcia.Discussion.-This species has greatly reduced,probably non-functional eyes. Each of the three cavesfrom which it is recorded is located in an isolatedlimestone range in the Northern Basin and RangeProvince. The lack of divergence and the retentionof eye remnants imply that this is a comparativelyrecent troglobite. This species was found in largenumbers along the stream passage in Cueva del Carrizal.Geographically close members of the genus areH. russelli Goodnight and Goodnight and H. madlaeGoodnight and Goodnight, both of which are troglobitesfrom Central Texas.Hoplobunus planus Goodnight and GoodnightHoplobunus planus Goodnight and Goodnight, 1973:88, 90, fig. 9.Type-Iocality.-Cueva de San Nicolas, 10 km SWAquism6n, San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 50.Discussion.-This eyeless species is apparentlymost closely related to Troglostygnopsis inops and, ifthe genus Troglostygnopsis is valid, this species maybelong to it. The genus Hoplobunus includes threeother cave-associated species, all of which are probablytroglophiles. Hoplobunus mexicanus (Roewer)has been found in one cave each in Oaxaca and Tamaulipas;H. spinoocularum Goodnight and Goodnightis a distinctive species found in abundance on thewalls and floor of Cueva del Guyabo, Oaxaca; and H.oaxacensis Goodnight and Goodnight was collectedfrom the walls and rocky floor areas of Cueva delLlano Grande and S6tano de los Arboles, Oaxaca.Hoplobunus queretarius SilhavyHoplobunus queretarius Silhary, 1974:176,179,180,182, fig. 2; Silhary, 1977:223.Type-Iocality.-Cueva del Madroiio, El Lobo, Queretaro,Mexico.Distribution.-Known from two caves in Queretaro.See Fig. 50.New record.-Queretaro: El Socav6n (det. C. J.Goodnight).Discussion.-This yellowish brown species apparentlyhas the eyes greatly reduced and probablylacking the retina. It is most closely related to H.zullinii from Chiapas.Hoplobunus zullinii SilharyHoplobunus zullinii Silhavy, 1977:219, 220-223, fig.1-9.Type-Iocality.-Grutas de Llano Grande, LlanoGrande, La Grandezza, Huixtla, Chiapas, Mexico.Distribution.-Known from two caves in Chiapas.See Fig. 50.165

Records.-Chiapas: Sumidero de Canada and Grutasde Llano Grande.Discussion.-This small-eyed presumed troglobiteis most closely related to H. queretarius.Mexotroglinus sbordonii SilhavyMexotroglinus sbordonii Silhavy, 1977:220, 231233, fig. 24-28.Type·locality.-Cueva del Perro de Agua, Rioegro, Lago de 1alpaso, Ocozocoautla, Chiapas,Mexico.Distribution.-Known only from the type-locality.See Fig. 49.Discussion.-The monotypic genus Mexotroglinusis most closely related to Monterella. This species iseyeless and depigmented with a reduced eyemound.Stygnomma pecki Goodnight and GoodnightStygnomma pecki Goodnight and Goodnight, 1977:148, 150, fig. 11.Type.locality.-St. Herman's Cave, Caves Branch,Belize.Distribution.-Known only from two caves atCaves Branch, Belize. See Fig. 49.Records.-BELIZE: Cayo District: Mountain CowCave and St. Herman's Cave.Discussion.-This is the only troglobitic species ofthe genus Stygnomma known from mainland NorthAmerica. Rambla (1969) described S. fiskei from acave in Jamaica; it is also eyeless. Stygnomma peckiis eyeless and light yellow. Stygnomma tuberculataGoodnight and Goodnight is a troglophile in Cuevade los Cuarteles, Tamaulipas.Troglostygnopsis anophthalma SilhavyTroglostygnopsis anophthalma Silhavy, 1974:176,182, 184, fig. 3(14-20); Sbordoni et al., 1974:32,34,45, pI. 5(fig. 13) (nomen nudum).Type.locality.-Grutas de Rancho Nuevo, S. Cristobalde las Casas, Chiapas, Mexico.Distribution.-Known from three caves in centralChiapas. See Fig. 50.Records.-Chiapas: Cueva de la Golondrina, Cuevadel Nacimiento del Rio Santo Domingo, and Grutasde Rancho Nuevo.Discussion.-This species is most closely related toT. inops. Eyes are absent. Like T. inops and H. planus,T. anophthalma is known only from high elevationcaves.Troglostygnopsis inops (Goodnight and Goodnight)Hoplobunus inops Goodnight and Goodnight, 1971:40-41, fig. 17; Reddell and Mitchell, 1971b:188,fig. 13; Goodnight and Goodnight, 1973:87, 90,fig. 7; Mitchell and Kawakatsu, 1973a:671;Mitchelland Reddell, 1973b:cover; Reddell, 1973a:34,39; Reddell and Elliott, 1973b:181, 184.Troglostygnopsis inops: Silhavy, 1974:176, 184,185,fig. 4(21).Type.locality.-Sotano de la loya de Salas, 21 kmNW of Gomez Farias, Tamaulipas, Mexico.Distribution.-Known only from caves in theSierra de Guatemala, Tamaulipas. See Fig. 50.Records.-Tamaulipas: Cueva de la Capilla, Sumiderode EI lineo, Sotano de la loya de Salas, Cuevade la Mina, Cueva del Ojo de Agua de Manantiales,and Sotano de Vasquez.Discussion.~This eyeless species is considered bySilhavy (1974) to be most closely related to T.anophthalma. A record of T. inops from Cueva delNacimiento del Rio San Antonio, Oaxaca (Goodnightand Goodnight, 1973), is doubtless in error. The tworegions share no other cave species in common, andany resemblance is probably due to convergence. TheOaxacan species is presumably an undescribed troglobite.Suborder PalpatoresFamily NemastomatidaeOrtholasma sbordonii SilhavyOrtholasma sbordonii Silhavy, 1974:176, 191, 193,fig. 7(43-49).Type-Iocality.-Cueva de la Perra (=Cueva de laCapilla), Sierra de Guatemala (Gomez Farias), Tamaulipas,Mexico.Distribution.-Known only from two caves in theSierra de Guatemala, Tamaulipas. See Fig. 49.Records.-Tamaulipas: Cueva de la Capilla andCueva de la Mina.Discussion.-Ortholasma has been erroneouslyplaced in the Trogulidae in the past; it clearly belongsin the Nemastomatidae, though probably in a newsubfamily also including Dendrolasma of Oregon andWashington and Cladolasma of Japan. Epigean speciesof Ortholasma are known from Oregon south tosouthern California, and from central Mexico, wherethe genus has been described under the synonym Trilasma(W. A. Shear, pers. comm.). The function ofthe peculiar cuticular sculpture of Ortholasma speciesappears to be the accumulation of dirt for purposesof camoflage. In O. sbordonii the dirt also covers thedepigrnented eyes.166

Family PhalangiidaeThe family Phalangiidae is well represented incaves throughout Mexico, but only three species haveyet been identified. Ceaya yucatana Goodnight andGoodnight is known from caves in Campeche andQuintana Roo and will probably be eventually recordedfrom many caves in the Yucatan Peninsula.Leiobunum metallicum Roewer has been found incaves in Queretaro and San Luis Potosi. Leiobunumviridorsum Goodnight and Goodnight is known froma single cave in San Luis Potosi. Leiobunum townsendiWeed is the common trogloxenic harvestmenin the caves of Central Texas and doubtless will beidentified from many caves in northern Mexico.Order PalpigradidaThe order Palpigradida is widespread in NorthAmerica, but it is seldom collected because of its minutesize and secretive habits. It is known to rangenorth from southern i\:lexico into California andTexas. The only known Mexican species, Koeneniahanseni Silvestri, has been reported by Remy (1948)from surface localities in Guerrero, Jalisco, NuevoLeon, Puebla, Quintana Roo, San Luis Potosi, Tabasco,Tamaulipas, Veracruz, and Yucatan. It hasbeen taken in abundance in detritus in darkness inSotano de la Tinaja, San Luis PotosI. Remy (1948)reported Koenenia sp. from Cuevas de Bellamar,Cuba.Order SolpugidaA single unidentified specimen of solpugid hasbeen collected from below the upper entrance toCueva del Abra, Tamaulipas. It doubtless fell into theentrance and should not be considered a part of thetrue cave fauna. No species of solpugid is known tobe associated with caves.Order AcarinaThe order Acarina is the most commonly encounteredgroup of arachnids in the caves of Mexico;it is also the least studied. The accompanying list offamilies (see Table 17) includes a minimum of 83 species,but many of these represent only family identifications.Other species included in this list are aquaticmites taken from open-air cenotes in Yucatan.The only part of Mexico which has seen any systematicstudy of its mite fauna is the state of Yucatan.Wharton (1938) studied several species of interestfrom the caves of that state.It is impossible here to go into detail on theknown mite fauna of Mexico, but a few families ofinterest are discussed below.Table 17.-Summary of cave inhabiting Acarina.TroglobitesOther SpeciesMesostigmataAscidae 0 1Heterozerconidae 0 2Laelapidae 0 1Macrochelidae 0 1Paramegistidae 0 1Parasitidae 0 1Polyaspidae 0 1Prodinychidae 0 1Spelaeorhynchidae 0 1Spintumicidae 0 3Trachytidae 0 1Trematurellidae 0 1Uropodidae 0 1MetastigmataArgasidae 0 12Ixodidae 0 3ProstigmataArrenuridae 0 1Cheyletidae 0 2Cunaxidae 0 1Ereynetidae 0 1Erythraeidae 0 1Hydrachnidae 0 1Hydrodromidae 0 1Limnesiidae 0 1Mideopsidae 0 1Myobiidae 0 1Pionidae 0 1Pterygosomidae 0 1Rhagidiidae 2 1Smaridiidae 0 1Trombiculidae 0 14Trombidiidae 0 3Unionicolidae 0 1AstigmataAcaridae 0 4Anoetidae 0 1Glycyphagidae 0 1Rosensteiniidae 0 1CryptostigmataAchipteriidae 0 1Belbidae 0 1Carabodidae 0 1Eremaeidae 0 1Galumnidae 0 1Hermanniidae 0 1Hypochthoniidae 0 1Oribatellidae 0 1Oribatulidae 0 1Phthiracaridae 0 1Total 2 81Suborder MesostigmataFamily AscidaeThe only species of ascid mite reported from cavesIn Mexico is an undetermined member of the genusMelichares, taken in Hoyo de Don Nicho, Chiapas. Itwas found in stream detritus.167

Family HeterozerconidaeSpecimens of Discozercon sp. were removed froma large centipede collected in Cueva de Taninul n. I,San Luis PotosI. Undetermined specimens probablybelonging to the genus Heterozercon were found inthe sediment of jars from Cueva de la Mina, Tamaulipas,and Cueva de EI Ocote, Hidalgo. The hosts wereprobably rhachodesmid millipeds.Family LaelapidaeBonet (1953a) reported the presence of undeterminedlaelapids of the subfamily Laelapinae in sixcaves of the Xilitla region, San Luis Potosi. Haemolaelapsglasgowi (Ewing) was reported by Wharton(1938) from the carcass of a paca in Grutas de Balankanche,Yucatan; it has recently been collected fromswallow guano in Actun Xpukil, Yu~atan.Family MacrochelidaeMacrocheles coprophila Womersley is a guano inhabitingspecies known from Cueva de la Hoya, SanLuis Potosi (Bonet, 1953a). Undetermined specimensof the same genus were reported from Cuevadel Jobo, San Luis Potosi (Bonet, 1953a).Family PolyaspidaeA single species of polyaspid mite, Dipolyaspissp., was reported from three caves in the Xilitla region,San Luis Potosi, by Bonet (1953a).Family SpelaeorhynchidaeThe unusual bat parasite, Spelaeorhynchus praecursorNeumann, was reported from caves in SanLuis Potosi and Tabasco by Hoffmann and de Barrera(1970).Family SpinturnicidaeThe spinturnicid mites are important parasites ofbats and are frequently present on the animals inlarge numbers. In areas of large bat deposits thisgroup of mites may be extremely abundant on theguano. Rudnick (1960) has revised the family. Threespecies have been found on bats in caves: Periglischrusiheringi Oudemans in Yucatan; P. vargasi Hoffmannin Chiapas and Tabasco; and Spinturnix carloshoffmanniHoffmann in Tamaulipas.Family TrachytidaeThis little-known family is represented in Mexicancaves only by specimens of the genus Uroseius fromCueva de Los Sabinos, San Luis Potosi.Family TrematureUidaeBonet (1953a) reported the presence of Trematurellasp. in one cave in Queretaro and three in SanLuis Potosi.Family UropodidaeThe family Uropodidae is a frequently collectedfamily in Mexican caves. Uropoda pearsei Whartonwas described from Cenote de Sambula (Motul), Yucatan(Wharton, 1938). Although Nicholas (1962)lists this species as a troglobite, it is certainly a troglophilicguanophile. It has been recently collected atthe type-locality from guano in a small crawlway atthe end of the cave.Suborder MetastigmataFamily ArgasidaeTwelve species of argasid tick have been found inthe caves of Mexico, but many are only accidentalvisitors. Species of the genera Antricola, Ornithodoros,and Nothoaspis are frequently found and areof some importance to the cave ecosystem. Antricolacoprophilus (McIntosh), A. marginatus (Banks), andA. mexicanus Hoffmann have all been found to beabundant in guano in bat caves in several parts ofMexico. The immature stages are parasites of bats,but the adults appear to be free-living in the guano.At times A. mexicanus is present in vast numbers.At least five species of the genus Ornithodoros occurin Mexican caves, but these are usually not found inlarge numbers and probably have merely dropped offthe bats or other host mammal. A unique new genusand species of argasid tick, Nothoaspis reddelli, wasrecently described from Grutas de Xtacumbilxunam,Campeche, by Keirans and Clifford (1975). It hasalso been found in Actun Xpukil, Yucatan, andCueva del Azufre, Tabasco (Keirans et al., 1977). Itis probably a parasite of the old man bat, Mormoopsmegalophylla.Family IxodidaeThree species of ixodid tick of the genus Amblyommahave been reported from caves in several statesof Mexico. All probably were parasites of small mammalsusing the cave entrance area for shelter.Suborder ProstigmataThe suborder Prostigmata is represented in Mexicancaves by 17 families and includes the most interestingspecies in the order with respect to the cavefauna. Marshall (1936) reported specimens of thefamilies Arrenuridae, Hydrachnidae, Hydrodromidae,168

Limnesiidae, Mideopsidae, Pionidae, and Unionicolidaefrom caves and cenotes in Yucatan. With theexception of two species reported from caves thesefamilies will not be further discussed.Family CheyletidaeTwo species of cheyletid mite have been collectedfrom caves in Mexico. Cheyletus cacahuamilpensisBaker, described from Grutas de Cacahuamilpa,Guerrero (Baker, 1949), is also known from caves inYucatan. Cheyletus malaccensis Oudemans has beencollected recently from caves in Tamaulipas and Veracruz.Family EreynetidaeBaker (1945) described Ereynetes sabinensisfrom Cueva de Los Sabinos, San Luis Potosi. Thispredaceous mite probably inhabits washed-in organicdebris in caves.Family ErythraeidaeOne species of this family, Erythraeus bisetosaWharton, was described from Cueva de San Isidro,Yucatan. ;\licholas (1962) lists it as a troglobite, butit is at best a troglophile. The larvae of erythraeidsare parasitic on insects.Family HydrodromidaeHydrodroma despiciens (l\liiller) is a troglophile inhabitingpools in Grutas de Balankanche, Yucatan(Marshall,1936).Family LimnesiidaeMarshall (1936) reported Limnesia paucispinaWolcott from five open-air cenotes and two caves inYucatan. It is presumably a troglophile in Gru tas deBalankanche and Cueva de Santa Elena.Family MyobiidaeThe only species of myobiid mite reported fromMexican caves is famesonia arganoi Vomero. Thisspecies was described from the vampire bat Desmodusrotundus murinus Wagner in Cueva de Los Sabinos,San Luis Potosi (V omero, 1972).Family PterygosomidaeCunliffe (1952) reported Hirstiella trombidiformes(Berlese) from Cueva del Carrizal, Nuevo Leon. Thisis a frequent parasite of lizards; the host in this caveis unknown.Family RhagidiidaeThe family Rhagidiidae is frequently collected incaves, and many species in Europe and the easternUnited States have been tentatively considered to betroglobites. Two described species are possible troglobitesand discussed below. An undescribed species ofprobable troglophile is known from Cueva del Diablo,Veracruz. Undetermined material is also known fromGrutas del Palmito and Grutas de Villa de Garcia,Nuevo Leon.Rhagidia trisetata Elliott and StrandtmannRhagidia sp.: Reddell and Mitchell, 1971a:149.Rhagidia trisetatus Elliott and Strandtmann, 1971:473-474, fig. 1-3, 8-16; Reddell, 1973a:34, 39;Reddell and Elliott, 1973a: 174.Rhagidia trisetata: Elliott, 1976: 14.Type-Iocality.-El Sotano de la Tinaja, San LuisPotosi, Mexico.Distribution.-Known only from the type-locality.Discussion.-This species is known only from onefemale. It was found in association with R. weyerensis.These two species are the only members of thefamily which have been described from Mexico.Rhagidia weyerensis (Packard)Bryiobia? (or Penthaleus?) weyerensis Packard, 1888:42, pl.ll(fig. 1, la, Ib).Bryiobia weyerensis: Banks, 1907:598.Rhagidia cavicola (part): Banta, 1907:68.Rhagidia weyerensis: Holsinger, 1965:654, 655-658,661, fig. 1-6; Elliott and Strandtmann, 1971:469473, fig. 4-7, 17-24; Reddell and Mitchell, 1971a:149; Reddell and Elliott, 1973a:174.Type-Iocality.-Weyer's Cave (=Grand Caverns),17 miles north of Staunton, Augusta County, Virgima.Distribution.-Known from Grand Caverns, Virginia;Old Spanish Cave, Stone County, Missouri; CarlsbadCaverns, Eddy County, New Mexico; and Sotanode la Tinaja, San Luis Potosi, Mexico.Mexico record.-San Luis Potosi: Sotano de la TinaJ3.Discussion.-The distribu tion of this mite defiesexplanation at this time. No known species of troglobiteis known to have so great and disjunct a distribution;yet no collections of this species are knownfrom the surface, and much of the intervening terrainis hardly suitable for these humus inhabitinganimals. Elliott and Strandtmann (1971) found nodifferences between specimens from Virginia andMexico and were confident that they dealt with169

only one species. All known epigean rhagidiidspossess eyespots and are either red, orange, or yellow.This species lacks eyespots and pigment and givesevery indication of being a troglobite. It is, therefore,listed here with reservations.Family SmaridiidaeSpecimens of Fessonin sp. were collected fromcave swallow guano in Cueva del Salitre, Chihuahua.This group of mites is parasitic on insects as larvae,but predaceous as adults.Family TrombiculidaeThe family Trombiculidae is a significant elementof the cave fauna of Mexico. The larvae of this familyare parasites of vertebrates, and many are known onlyfrom bats. The adults of many trombiculids havebeen collected from caves in Mexico, but the presentsystematics of the family is based on larval charactersso that identification of the free-living adults is practicallyimpossible. Wharton (1938) described oneadult species, Trombicula camilla, from caves in Yucatan,but it cannot now be identified with any ofthe described larval species. Either this species orothers are frequently present as adults on guano inmany caves in Yucatan and are certainly importantto the ecology of the caves. Nicholas (1962) lists T.camilla and Hannemanin (=Whartonin) nudosetosaWharton as troglobites, but these are obviously notcave adapted. Hoffmann (1970) has begun a revisionof the trombiculids of Mexico.Family TrombidiidaeThe trombidiid mites are parasites of arthropodsin the larval stages. Nicholas (1962) lists two Yucatanspecies, Platyseta yucatanicus Wharton and Monunguisstreblida Wharton, as troglobites; neither speciesis cave adapted. Robaux et al. (1977) described aninteresting trombidiid, Ceuthothrombium cavaticum,from camel crickets (Rhaphidophoridae) from cavesin Texas, New Mexico, and Mexico (Cueva del Ojode Agua, Dulces Nombres, uevo Leon).Suborder AstigmataFamily AcaridaeBonet (1953a) reported the presence of Acarus sp.in Cueva de la Hoya, San Luis Potosi. Nesbitt (1949)described three species of the genus Caloglyphus fromMexican caves: C. armipes longisetosus from Cueva dela Calera, Michoacan; C. longipilus from Grutas deJuxtlahuaca, Guerrero, and Cueva del Carrizal, NuevoLeon; and C. paranomalus from Grutas de Atoyac,Veracruz, Cueva de la Calera, and Cueva del Carrizal.Species of Caloglyphus have been reported to feed ondecaying insects.Family AnoetidaeHistiostoma sp. has been recently collected in SOtanode Yerbaniz, San Luis Potosi. The anoetids areusually found on decaying organic matter.Family GlycyphagidaeBonet (1953a) reported the presence of undeterminedspecimens of this family in caves in Queretaroand San Luis Potosi. Glycyphagus domesticus(DeGeer) has been collected in Cueva del Tempisque,Chiapas, and Cueva del Pedregoso, Coahuila. This isa cosmopolitan species usually found on organic matter;when present in large numbers it can causedermatitis.Family RosensteiniidaeNycteriglyphus sp. was found in Cueva de LosSabinos, San Luis Potosi, and Cueva de la Mina, Tamaulipas.This genus of mite is commonly associatedwith caves and is present at times on bat guano inenormous numbers.Suborder CryptostigmataThe oribatid mites certainly abound in the caves ofMexico, but few have been identified. None show anysigns of adaptation to the cave environment, but theyare doubtless of considerable importance to the caveecosystem.Family BelbidaeBonet (1953a) reported undetermined species ofthe family to be present in six caves of the Xilitla region,San Luis Potosi. Belba sp. has been identifiedfrom Cueva de La Lagunita, San Luis Potosi.Family GalumnidaePearse (1936c) reported Galumna sp. from twocenotes in Yucatan. Galumna jacoti Wharton wasdescribed from Cenote de Sambula (Motul), Yucatan(Wharton, 1938).Family OribateUidaeBonet (1953a) reported the presence of this familyin caves in Queretaro and San Luis Potosi. Oribatellamonospicus Wharton was described from Cenote deSambula (Motul), Yucatan (Wharton, 1938).170

Family OribatulidaeScheloribates luchili was described by Wharton(1938) from Cueva Luchil, Yucatan. Bonet (1953a)reported the presence of this family in Cueva delJobo, San Luis Potosi.Class ChilopodaThe cavernicole fauna of Mexico is rich in numbersand diversity of centipedes, but this group remainsessentially unstudied. Only 13 species of centipedehave been identified from caves in Mexico and onlyfive of these have been reported from more than onecave (see Table 18).Order GeophilomorphaFamily GeophilidaeThe order Geophilomorpha is frequently found incaves in Mexico, but only the geophilid Pachymeriumferrugineum (C. 1. Koch) has been identified. It isknown only from Cueva del Ojo de Agua Grande andCueva del Ojo de Agua de Tlilapan, Veracruz.Order LithobiomorphaFamily LithobiidaeGarcibius osorioi ChamberlinGarcibius osorioi Chamberlin, 1942:5-6; Chamberlin,1943a:27; Bonet, 1946a:111; Nicholas, 1962:175;Reddell, 1967a:24; Reddell, 1971b:42.Type-Iocality.-Grutas de Garcia (=Grutas de Villade Garcia), Nueva (=Nuevo) Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 51.Discussion.-In this species the ocelli are rudimentary,the antennae are very long, and the animal ispale. In the present state of North American chilopodtaxonomy it is impossible to say anything about theaffinities of this species.Nuevobius cavicolens ChamberlinNuevobius cavicolens Chamberlin, 1941:188; Crabill,1960:121-123,127; Reddell, 1971b:42.Type-locality.-Bat Cave (=?Cueva de la Boca),Villa Santiago, Hacienda Vista Hermosa-HorsetailFalls, Nuevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 51.Discussion.-Nuevobius cavicolens is a pale specieswith long, slender legs and antennae. The only othermember of the genus is N. cottus Crabill, known onlyfrom Tuckaleechee Caverns, Blount County, Tennessee.This remarkable distributional discontinuity isTable 18.-Summary of cave inhabiting Chilopoda.TroglobitesOther SpeciesGeophilomorphaGeophilidae 0 1LithobiomorphaLithobiidae 2 0Watobiidae 0 1ScolopendromorphaCryptopidae 2 2Seolopendridae 0 1SeutigeromorphaPselliodidae 0 1Seutigeridae 0 3Total 4 9similar in some respects to that of the milliped familyTrichopetalidae. The identity of the type-locality issomewhat uncertain, but it is probably Cueva de laBoca.Family WatobiidaeOne species of troglophile, Cruzobius atoyacusChamberlin, has been described in this family. It isknown only from Grutas de Atoyac, Veracruz, butdoes not appear to be cave adapted (see Fig. 51).Order ScolopendromorphaFamily CryptopidaeNewportia (Scolopendrides) sabina ChamberlinNewportia sabina Chamberlin, 1942:4; Chamberlin,1943a:5, 7, 9; Bonet, 1946a:l11; Bucherl, 1959:230, 231; Nicholas, 1962:175; Reddell, 1971b:43; Reddell and Mitchell, 1971a: 150.Newportia (Scolopendrides) sabina: Bucherl, 1959:234,235.Type-locality.-Cueva de los Sabinos, San Luis Potosi,Mexico.Distribution.-Known from three caves in theSierra de EI Abra, San Luis Potosi, and possibly onecave in the Sierra de Guatemala, Tamaulipas. SeeFig. 51.New records.-San Luis Potosi: Sotano de la Tinajaand Sotano de Yerbaniz; Tamaulipas: ?Bee Cave(det. A. Weaver).Discussion.-A single specimen from Bee Cavelacks the ultimate legs, but in other characters appearsto be identical to typical N. sabina. Three otherspecies of the genus Newportia are known from Mexicancaves. Newportia (Scolopendrides) pelaezi Chamberlinis a small species described from Grutas del Palmito,Nuevo Leon, and known only from the holotype(Chamberlin, 1942). Its correct ecological desig-171

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

Mexico and Guatemala, and several undescribed epigeanspecies from Mexico and Belize. One isolatedspecies of the genus occurs in California.Glomeroides addititius CauseyGlomeroides addititius Causey, 1973:107, fig. 1-2.Type-Iocality.-Cueva de Ungurria, about 20 kmWSW Tezonapa, Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 52.Discussion.-This is an eyeless species and in thisrespect most closely resembles G. caecus Causey. Itwas found on mud in a lower-level room of the cave.Glomeroides caecus CauseyGlomeroides caecus Causey, 1964b:63, 65, 66, fig.1-5; Reddell, 1967d:l06; Loomis, 1968:7; Causey,1971b:30; Reddell, 1971b:43; Causey, 1973:107;Reddell, 1973a:34, 40; Shear, 1974:248,249.Type-!ocality.-Sotano de Huitzmolotitla, about2 km southwest of Tlamaya, which is 10 km northeastof Xilitla, San Luis Potosi, Mexico.Distribution.-Known only from two caves in theXilitla region, San Luis Potosi. See Fig. 52.Records.-San Luis Potosi: Sotano de Huitzmolotitlaand Sotano de Tlamaya.GUATEMALA/.//r"\r"..:- ""Fig. 52.-Distrihution of troglobitic millipeds of the families Glomeridae and Glomeridesmidae: 1, Glomeroides promiscus;2, Glomeroides caecus; 3, Glomeroides n. sp. 1; 4, Glomeroides pellucidus; 5, Glomeroides addititius; 6, Glomeroides n. sp. 2;7, Glomeridesmus sbordonii.174

Discussion.-This eyeless species is apparently restrictedin its distribution to caves near Xilitla. Bothof the caves from which it is recorded are deep wetsystems.Glomeroides pellucidus ShearGlomeroides pellucidus Shear, 1974:248-249, fig.5-8.Type-Iocality.-Cueva del Ojo de Agua Grande,Paraje Nueva (=Nuevo), Cordoba, Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 52.Discussion.-This species possesses cuticular remnantsof ocelli. A single specimen was collected recentlyfrom silt in an upper level.Glomeroides promiscus CauseyGlomeroides promiscus Causey, 1964b:63, 65-66,fig. 6-8; Reddell, 1966a:13; Reddell, 1967c:55;Loomis, 1968:7; Reddell, 1971b;43; Reddell andMitchell, 1971b:189; Reddell, 1973a:34, 40;Reddell and Elliott, 1973b:181, 185; Shear, 1974;248,249; Vomero, 1974:350.Type-Iocality.-Sotano de Gomez Farias, 5 kmeast of Gomez Farias, Tamaulipas, Mexico.Distribution.-Known only from caves in theSierra de Guatemala, Tamaulipas. See Fig. 52.Records.-Tamaulipas: Sotano de Gomez Farias,?Cueva de la Mina, Sotano del Naranjo, Cueva delOjo de Agua de Manantiales, ?Cueva del· Rancho delCielo n. 3, and ?Cueva del Rancho del Cielo n. 7.Discussion.-This species possesses a few ocelli butis doubtless a troglobite. The records indicated by aquestion mark are represented only by females andare thus tentative. Immature specimens taken inCueva del Nacimiento del Rio Frio, Tamaulipas,probably also belong to this species. This milliped isusually found on moist silt and frequently in directassociation with other species of milliped.Order ChordeumidaThe order Chordeumida is the dominant group inthe milliped fauna of the caves of the eastern UnitedStates. It has not been nearly so successful in thecaves of Mexico and Central America; few specieshave invaded caves and fewer yet have become troglobites.Only two families in this order are representedin the cave fauna of Mexico.Family CleidogonidaeShear (1972) has monographed the family Cleidogonidaeand reviewed the order Chordeumida inNorth America. He reported six species of Cleidogonafrom caves in Mexico; two additional species havebeen added since (Shear, 1974, 1977a). None of thesespecies shows the high degree of adaptation to thecave environment which is present in some othermilliped families in Mexico, but five appear to beadapted to cave life and are included here (see Fig.53). Of the remaining three, one (c. crystallina Shear)is known only from caves and may now be isolated inthe cave environment. The two other species are alsoknown from epigean localities. Cleidogona mayapecShear has been identified from Cueva del Judio, Queretaro,and C. totonaca Shear has been found in SOtanodel Gobernador, Queretaro. Other collections ofthis family from caves in Tamaulipas and Veracruzawait study.Cleidogona baroqua ShearCleidogona baroqua Shear, 1972:198,207,209-210,fig. 197-200.Type-Iocality.-Sotano de San Augustin (=Agustin),Huautla de Jimenez, Oaxaca, Mexico.Distribution.-Known only from the type-locality.See Fig. 53.Discussion.-This depigmented species possessesocelli and is only tentatively considered to be a troglobite.It is a member of the crucis group. This speciesgroup includes the troglobitic C. crucis (Chamberlin)from Veracruz and three epigean species fromOaxaca. Shear (1972) considers this to be the mostprimitive group in the genus. Because of the presenceof ocelli, this species would have to be considered avery recent troglobite. The epigean species of thecrucis group are known from comparatively highelevations (1700 to 2500 meters), whereas C. baroquais found at about 1400 meters. The more highlyevolved troglobite C. crucis is found at 500 meters.Although only much additional collecting on the surfaceand in caves can clarify the situation, it is possiblethat the epigean ancestor of C. crucis becameextinct in the Veracruz lowlands much earlier thanin the Huautla area and that it or its closest relativesstill exist at high elevations in Oaxaca. This may be asimilar situation to that in the scorpion genus Typhlochactasand the milliped genus Mexiterpes.Cleidogona crucis (Chamberlin)Cavota crucis Chamberlin, 1942:8-9, fig. 3-7; Chamberlin,1943b;5, 36; Bonet, 1946a:112; Nicholas,1962:177; Loomis, 1968:66; Reddell, 1971a:222;Reddell, 1971b;44.Cleidogona crucis: Shear, 1972:207, 209, fig. 193196; Shear, 1974:245,259.175

Type-locality.-Gruta de Atoyac, Veracruz, Mexico.Distribution.-Known only from two caves nearCordoba, Veracruz. See Fig. 53.Records.-Veracruz: Grutas de Atoyac and Cuevadel Ojo de Agua Grande.Discussion.-This species lacks pigment, and theocelli are reduced to 10. It is the most highly caveadaptedmember of the genus in Mexico.Cleidogona felipiana ShearCleidogonafelipiana Shear, 1974:254-255, fig. 9-13.Type-locality.-Cueva del Rayo de San Felipe, SanCristobal de las Casas, Chiapas, Mexico.Distribution.-Known only from the type-locality.See Fig. 53.Discussion.-This probable troglobite is lacking inpigment and has fewer ocelli than most epigean species.This species is most closely related to C. decurvaShear, an epigean species from Chiapas. It is a memberof the stolli group which, in addition to the twoChiapas species, includes species from El Salvador andGuatemala.MEXICO///)I\ /'v/IIIIII:/GUATEMALAil" /............... _ L..J ..../- _r,- f-.-.-._._' I[\_// "'-... I i,,_I'-.....I/i\./)'"'-.,/----'IiL///r\rr-'.-"Fig. 53.-Distrihution of troglobitic and troglophilic millipeds of the families Cleidogonidae and Trichopetalidae; 1, Cleidogonacrystallina and C. pecki; 2, Mexiterpes sabinus; 3, M. egeo; 4, M. FlShi; 5, C. mayapec, C. totonaca, and M. metallicus;6, C. crucis; 7, C. baroqua; 8, C. hunapu; 9, C. jelipiana.176

Cleidogona hunapu ShearCleidogona hunapu Shear, 1977a:236-238, fig. 1-8.Type-locality.-Cueva del Burro, Lago de Malpaso,Rio Encajonado, Chiapas, Mexico.Distribution.-Known only from the type-locality.See Fig. 53.Discussion.~This depigmented species with only14 ocelli is closely related to C. forficula Shear, anepigean species from San Cristobal de las Casas, Chiapas.These are the only species known in the forficulagroup.Cleidogona pecki ShearCleidogona sp.: Reddell and Mitchell, 1971b:188(Cueva de la Capilla and Cueva de la Mina recordsonly).Cleidogona pecki Shear, 1972:207, 213, fig. 221225; Reddell and Elliott, 1973b:185; Shear,1974:254.Type-locality.-Cueva de la Mina, 6 mi northwestof Gomez Farias, Tamaulipas, Mexico.Distribution.-Known from two caves in the Sierrade Guatemala, Tamaulipas. See Fig. 53.Records.-Tamaulipas: Cueva de la Capilla andCueva de la Mina.Discussion.~ This probable troglobite is unpigmentedand has the ocelli reduced to 16-20. It is amember of the crystallina group. The only other speciesin this group is C. crystallina Shear, which isknown only from four caves in the Sierra de Guatemala,including Cueva de la Capilla. Cleidogona crystallinaalso has a slightly reduced number of ocelli,and some individuals have reduced pigmentation.The crystallina group is intermediate in structurebetween the crucis group and the large, apparentlyactively speciating maculata group to the north andsouthwest. The crystallina group may represent apair of relict species now limited to caves.Family TrichopetalidaeMexiterpes egeo (Causey)Poterpes egeo Causey, 1969:46-47, 49, fig. 5,7-10;Reddell, 1971b:44; Shear, 1972:247; Elliott andReddell, 1973:196; Reddell, 1973a:34, 40.Mexiterpes egeo: Shear, 1974:255,256.Type-Iocality.-Cueva de EI (=la) Puente, 13 milessouth of San Francisco, San Luis Potosi, Mexico.Distribution.-Known from caves in the vicinity ofSan Francisco, San Luis Potosi. See Fig. 53.Records.-San Luis Potosi: ?Sotano del Puerto delos Lobos and Cueva de la Puente.Discussion.-Only a single female is known fromSotano del Puerto de los Lobos so this record must beconsidered tentative. Although eyes are present, theyare reduced. The family Trichopetalidae is representedin Mexico only by the four troglobitic specieslisted here. Although Shear (1974) considersPoterpesto be a synonym ofMexiterpes, Causey (pers. comm.)does not feel this position to be justified. The discoveryof additional species should help to clarify thegeneric status of the species ofPoterpes.Mexiterpes fishi (Causey)Poterpes fishi Causey, 1969:46, 47-49, fig. 6, 11-16;Reddell, 1971b:44; Shear, 1972:247; Reddell,1973a:34, 40.Mexiterpes fishi: Shear, 1974:256.Type-locality.-Cueva de la Luz, 20 miles west ofAquismon, San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 53.Discussion.-This species is most closely related toM. egeo. The ocelli are reduced in number and thespecies is certainly a troglobite.Mexiterpes metallicus ShearMexiterpes metallicus Shear, 1972:247, 275, 280,281, fig. 532-538; Shear, 1974:256; Peck, 1977c:67.Type-locality.-1.2 miles east of Pinal de Amoles,Queretaro, Mexico, in an iron mine at roadside.Distribution.-Known only from the type-locality.See Fig. 53.Discussion.-This species possesses four smallocelli. It is most closely related to M. fishi and M.egeo. The "iron mine" is apparently a natural caveenlarged during the mining of mercury rather thanIron.Mexiterpes sabinus CauseyMexiterpes sabinus Causey, 1963:235-237, fig. 1-3;Reddell, 1965c:7; Reddell, 1967b:82; Causey,1968:124; Loomis, 1968:69; Causey, 1969:4546, 49; Reddell, 1971b:44; Reddell and Mitchell,1971a:151; Shear, 1972:280, 281, fig. 529-531;Reddell, 1973a:34, 40; Reddell and Elliott,1973a:I72; Shear, 1974:255,256.Type-Iocality.-Sotano del Arroyo, near village ofLos Sabinos and 8 miles north of Valles, San Luis Potosi,Mexico.Distribution.-Known from four caves in theSierra de EI Abra near Valles, San Luis Potosi. SeeFig. 53.177

Records.-San Luis PotOSI: Sotano del Arroyo,?Cueva Pinta, Cueva de Los Sabinos, and ?Sotano dela Tinaja.Discussion.-The localities marked with a questionmark are not known by adult males but almost certainlybelong to this species. Shear (1974) stated thatspecimens from Cueva de Los Sabinos are "topotypes,"but this is in error since the type-locality isSotano del Arroyo. Specimens from Cueva de losMonos, San Luis Potosi, are very immature, but willprobably prove to belong to this species. Mexiterpessabinus is the most highly evolved troglobite in thegenus. This is of special interest since it is a lowlandtroglobite. Since the Trichopetalidae is a group oftemperate relicts in Mexico, it is not surprising thatthe epigean ancestors survived longer at more temperatehigher elevations. The existence of species withocelli in caves at high elevations suggests that theyhave become more recently restricted to caves. It isimportant to make intensive surface collections inthe Sierra Madre Oriental to determine whetherfully eyed species still survive in the epigean environment.The situation with these millipeds is probablyanalogous to that already discussed for Typhlochactasand for the Cleidogona crucis group.Order JulidaFamily JulidaeA single introduced species in the family Julidae,Diploiulus latistrintus (Curtis), has been taken inMexican caves. This species was collected in Cuevade las Cuatas, San Luis Potosi, and is probably anaccidental.Family ParaiulidaeAn apparently undescribed species of the genusParaiulus has been collected from three caves at Vallede los Fantasmas, San Luis Potosi. All were takennear the entrance or in areas where flood debris hadaccumulated, indicating that this is an accidental.Order PolydesmidaThe order Polydesmida has been a remarkablysuccessful group in colonizing the caves of Mexicoand Central America. Although 23 troglobites and 64other species are known from caves in this area (seeTable 20), probably as many more species await descriptionor identification. Each visit to a new caveregion produces additional new species of polydesmoids,including at least one or two troglobites, whilethe most carefully studied regions continue to producenew species with almost every investigation.Table 20.-Summary of cave-inhabiting Polydesmida.TroglobitesOther SpeciesChelodesmidae 0 1Euryuridae 1 1Oniscodesmidae 2 0Paradoxosomatidae 0 2Peridontodesmidae 1 2Pyrgodesmidae 0 34Rhachodesmidae 10 14Sphaeriodesmidae 0 8Trichopolydesmidae 8 0Tridontomidae 1 0Xystodesmidae 0 2Total 23 64Family ChelodesmidaeOnly one species of the family Chelodesmidae hasbeen identified from caves in Mexico. Chondrodesmussabachanus Chamberlin was described from ActUnSabaca, Yucatan. It or a closely related specieshas since been collected from several cave and surfacelocalities in northern Yucatan and eastern Campeche.Although Nicholas (1962) lists it as a troglobite, it isprobably a troglophile or trogloxene.Family EuryuridaeUndetermined species of the genus Pseudamplinusare known from caves in Chiapas, Oaxaca, San LuisPotosi, and Veracruz. The ecological status of themembers of this genus is unknown, but some may betroglophiles. The only troglobite in the family isPolylepiscus vomeroi Shear, which is discussed below.Polylepiscus vomeroi ShearPolylepiscus vomeroi Shear, 1977a:242, 244, 245,fig. 19-23.Type-locality.-Gruta II Finca Santa Anita, Simoyovelde Allende, Chiapas, Mexico.Distribution.-Known only from the type-locality.See Fig. 54.Discussion.-This depigmented and weakly sclerotizedspecies is the only trogiobite in the family. SpeciesofPolylepiscus are known only from Chiapas andGuatemala.Family OniscodesmidaeBonetesmus ojo ShearBonetesmus ojo Shear, 1974:260, fig. 29-31; Shear,1977a:260, fig. 44.Bonetesmus: Shear, 1977a:253.178

Type-locality.-Cueva del Ojo de Agua de Tlilapan,Tlilapan, near Orizaba, Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 54.Discussion.-The Oniscodesmidae reaches itsnorthern distributional limit in Veracruz where it isrepresented by the two troglobitic species listed here.The only other Mexican species is an epigean species,Ligiodesmus pusillus Pocock, from Tabasco.Bonetesmus verus ChamberlinBonetesmus verus Chamberlin, 1942: 11-12, fig. 1820; Chamberlin, 1943b:6, 62; Bonet, 1946a:113;Loomis, 1968:111; Causey, 1971b:30; Reddell,1971b:44; Shear, 1974:256, 257, 259-260, 274,fig. 20-28.Bonestmus verus: Nicholas, 1962:177 (erroneousspelling).Bonetesmus novenus Causey, 1971b:30; Reddell1973a:40; Shear, 1974:257,259.Type-locality.-Of Bonetesmus verus: Grutas deAtoyac, Veracruz, Mexico; of Bonetesmus novenus:Cueva del Ojo de Agua Grande, 12 km NE Cordoba,Veracruz, Mexico.Distrihution.-Known from three caves in the vicinityof Cordoba, Veracruz. See Fig. 54.MEXICOGUATEMALA\r",..""--//rr"Fig. 54.-Distribution of troglobitic millipeds of the families Trichopolydesmidae, Oniscodesmidae, Euryuridae, and Tridontomidae:1, Ty/ogoneus minus; 2, Speodesmus pecki; 3, Ty/ogoneus rainesi; 4, Bonetesmus verus; 5, B. ojo; 6, Carambade/burro; 7, Po/y/episcus vomeroi; 8, C. de/negro; 9, C. grandeza; 10, Tridontomus loomisi.179

Records.-Veracruz: Grutas de Atoyac, Cueva delOjo de Agua Grande, and Cueva de Sala de AguaGrande.Discus&on.-This species was abundant on andnear rotting wood in the wetter areas of each of thethree above caves.Family ParadoxosomatidaeTwo species of the family Paradoxosomatidae havebeen found in caves in Mexico, Guatemala, and Belize.Both are tropicopolitan in distribution and nowappear almost everywhere man is found. Oxidus gracilis(Koch), the hothouse milliped, has been collectedin 14 caves in Nuevo Leon, San Luis Potosi,Queretaro, Hidalgo, Veracruz, and Guerrero, Mexico;and from one cave in Belize. Orthomorpha coarctataSaussure has been found in Cueva Jobitzinaj, El Peten,Guatemala; and in caves in Oaxaca and Yucatan.Most of the caves inhabited by these two species areeither frequented by man or are in or near towns.Family PeridontodesmidaeThe peridontodesmid genera Hexodontin andPeridontodesmus have been collected from caves inSan Luis Potosi and Tamaulipas; their ecologicalstatus is unknown. A possibly undescribed genus andspecies of troglobite is known from two caves inChiapas.Family PyrgodesmidaeNo family which has invaded caves in Mexico andCentral America has been so prolific in the number ofspecies which have become established in caves as thePyrgodesmidae (=Stylodesmidae). Although 34 speciesin this family have been reported from caves andmost are known only from caves, this is only a smallpercentage of the number which eventually will bedescribed from the cave habitat. Numerous collectionsawait study, and despite the recent contributionof Shear (1977a), systematic problems still preventmany of these from correct identification orgeneric placement.Many epigean forms are poorly pigmented so thatit is presently not possible to list any of the speciesas restricted to caves. Some members of the familyhave certainly become sufficently distinct as to beisolated now in the cave environment, but only muchadditional study and a better understanding of thesystematics of the family will allow us to say whichspecies are troglobites and which troglophiles. Nicholas(1962) listed seven species from Mexico and onefrom Guatemala as troglobites; none of these appearto possess unusual modifications restricting them tocaves.These small millipeds are usually black, gray, orcream colored, and many species are covered with afine layer of dirt adhering to the numerous smallknobs and projections on the body. They are foundin a variety of habitats in caves, ranging from moistflowstone, guano, and silt to the underside of smallrocks and rotting wood.The genus Myrmecodesmus has recently beenexpanded by Shear (1977a) to include species previouslyplaced in the genera Bolivaresmus, Ceratesmus,Gibberdesmus, and Stenotodesmus. The followingis a list of the 18 species of cavernicole Myrmecodesmusknown from this area (see Figs. 55-56).Myrmecodesmus aconus (Shear, 1974)-Gruta de laEstrella, Guerrero, and Cueva de Coatepec, Mexico,Mexico.M. amarus (Causey, 1971b)-Sotano de Tlamaya, SanLuis Potosi, Mexico.M. amplus (Causey, 1973)-three caves near Valle Nacional,Oaxaca, Mexico.M. brevis Shear, 1977a-Rio Frio Cave A, Belize.M. clarus (Chamberlin, 1942)-Grutas de Atoyac andLas Tres Cuevas, Veracruz, Mexico.M. colotlipa (Chamberlin, 1942)-Grutas de J uxtlahuaca,Guerrero, Mexico.M. cornutus (Shear, 1974)-Cueva del acimiento delRio Frio, Tamaulipas, Mexico.M. egenus (Causey, 1971b)-caves in Tamaulipas,Mexico.M. errabundus (Shear, 1974)-caves in the northernSierra de EI Abra, Tamaulipas, Mexico.M. fissus (Causey, 1977a)-Sumidero del Camino,Chiapas-NEW COMBINATION.M. fuscus (Causey, 1977a)-Grutas de Atoyac andCueva de la Sala de Agua Grande, Veracruz, Mexico-NEWCOMBINATION.M. gelidus (Causey, 1971b)-caves in Tamaulipas,Mexico-NEW COMBINATION.M. ilymoides (Shear, 1974)-Grutas de San Bartolo,Nuevo Leon, Mexico.M. inornatus Shear, 1977a-Grutas de Llano Grande,Chiapas, Mexico.M. monasticus (Causey, 1971b)-Cueva de Llanura,San Luis Potosi, Mexico.M. potosinus (Shear, 1974)-Cueva de la Porra, SanLuis Potosi, Mexico.M. sabinus (Chamberlin, 1942)-caves in the southernSierra de EI Abra, San Luis Potosi, Mexico.M. unicorn Shear, 1977a-S1. Hermans Cave andMountain Cow Cave, Cayo District, Belize.The genus Calymmodesmus includes one speciesfrom Guatemala and five from Yucatan (see Fig. 57).180

Calymmodesmus inquinatus Causey was describedfrom Gruta del Silvino, Izabal, Guatemala (Causey,1960); it is also known from Cueva de Yaxchilan,EI Peten, Guatemala. Calymmodesmus alienus (Chamberlin),C. isidricus (Chamberlin), C. muruztunicus(Chamberlin), C. viabilis (Chamberlin), and C.hoctunanus (Causey) are all known only from cavesin Yucatan (Chamberlin, 1938; Causey, 1971b).Shear (1974) described two species of Cryptymafrom caves in Mexico and Guatemala (see Fig. 57).Cryptyma cocona is known only from Grutas delCocona, Tabasco; and C. guatemala has been foundin three caves in Alta Verapaz, Guatemala.The genus Rettenmeyeria is represented in cavesonly by R. cryptymoides Shear. It was described byShear (1977a) from Rio Frio Cave A, Belize (seeFig. 57). Other species of this genus are myrmecophiles.Lophodesmus includes six species known fromcaves in Chiapas and Guatemala (see Fig. 57). Shear(1974) described 1. italolegatus from Cueva de laGolondrina, Chiapas; it is now known from two additionalcaves in Chiapas and from four in Yucatan.Lophodesmus tioticho Shear is known from threecaves in Chiapas (Shear, 1974). Causey (1977) consideredthe American "Lophodesmus" to belong toGUATEMALA.....-'/.//r\,"Fig. 55.-Distribution of eight species of troglophilic pyrgodesmid milliped of the genus Myrmecodesmus: 1, Myrmecodesmusilymoides; 2, M. egenus; 3, M. cornutus and M. errabundus; 4, M. errabundus; 5, M. monasticus; 6, M. amarus; 7, M.aconus; 8, M. clarus.181

the genus Synoptura and removed the above speciesto that genus. Hoffman (1976) described L. petrinusfrom Cueva Chirripeck, Alta Verapaz, Guatemala.Shear (1977a) added L. rodriguezi from Sima I delTempisque (=Hoyo de Don Nicho), Chiapas; L.shawcrossi from Cueva de Agua Escondida, Huehuetenango,Guatemala; and L. zuUinii from Grotas deLlano Grande, Chiapas.Two species apparently spread by commerce havebeen found in caves in the Yucatan Peninsula. Poratioidesdisparatus Loomis, known from Florida to ElSalvador, has been found in Cueva Luchil, Yucatan.Prosopodesmus jacobsoni Silvestri, a species describedfrom Java and ranging from Florida into CentralAmerica, is known from seven caves in Campeche andYucatan.Family RhachodesmidaeThe family Rhachodesmidae is among the moreimportant elements in the milliped fauna of caves inthis area. Ten species of troglobite and 14 other specieshave been identified from caves; many other species,especially of troglobite, await description.Nicholas (1962) lists five species of this family as(/GUATEMALA,...,..--/r/"'/Fig. 56.-Distribution of nine species of troglophilic pyrgodesmid milliped of the genus Myrmecodesmus: 1, Myrmecodesmusgelidus; 2, M. sabinus; 3, M. potosinus; 4, M. fuscus; 5, M. amplus; 6, M. colotlipa; 7, M. {usus; 8, M. inornatus; 9, M.unicorn.182

troglobites, only two of which are unquestionablysuch. The remaining three, Aceratophallus calcehtokanus,A. hoctunanus, and A. oxkutzcabus were describedby Chamberlin (1938) from caves in Yucatan(see Fig. 58). These three species are of uncertainecological status. The only unquestioned troglobite inthis genus, A. scutigeroides Shear, is listed below.The genus Rhachodesmus is represented in Mexicancaves by two species (Causey, 1973): R. digitatusCausey is an apparent troglophile from caves nearHuautla de Jimenez, Oaxaca; R. viridis (Saussure) ispresumably an accidental in Cueva del Ojo de AguaGrande, Veracruz (see Fig. 58).Two other genera of rhachodesmid are representedin Mexican caves by species which have probablybeen accidentally introduced into the caves. Rhachidomorphaadunca (Saussure and Humbert) is knownonly from Sotano del Relicario, Veracruz; and Tiphallusfrivolus Causey has been collected only inCuevacita de Nopales, San Luis PotosI (Causey,1973).The remaining four genera of rhachodesmidsMEXICOGUATEMALA·~~2 ,. 11~/(\....."...;0Fig. 57.-Distribution of troglophilic pyrgodesmid millipeds of the genera Cryptyma, Lophodesmus, Calymmodesmus, andRettenmeyeria: 1, Cryptyma eoeona; 2, Lophodesmus italolegatus; 3, L. italolegatus and L. rodriguez:i; 4, L. tiotieho; 5, L.z:ullinii; 6, Calymmodesmus isidricus and Calymmodesmus viabilis; 7, Calymmodesmus viabilis; 8, Calymmodesmus hoetunanus;9, Calymmodesmus alienus; 10, Calymmodesmus mUTUz:tunieus; 11, Calymmodesmus inquinatus; 12, Cryptyma guatemala;13, Rettenmeyeria eryptymoides.183

known from caves all contain troglobites and are discussedbelow.Aceratophallus scutigeroides ShearAceratophallus scutigeroides Shear, 1974:263, 269,271, fig. 50-54.Type-locality.-Cueva Sepacuite n. 1 and CuevaSepacuite n. 2, Finca Sepacuite, Senahli, Alta Verapaz,Guatemala.Distribution.~Known from two caves in AltaVerapaz, Guatemala, and possibly one in Chiapas,Mexico. See Fig. 58.Records.-GUATEMALA: Alta Verapaz: CuevaSepacuite n. 1 and Cueva Sepacuite n. 2; MEXICO:Chinpas: ?Cueva del Arco.Discussion.-The genus Aceratophallus rangesnorth into Chiapas and Yucatan. Several cave specieshave been described from Yucatan, but none are obvioustroglobites. The single female from Chiapas iseither A. scutigeroides or a closely related undescribedspecies. This species is in some ways similar toUnculabes arganoi Shear, and it may eventually benecessary to rearrange the species in several generaof the Rhachodesmidae.GUATEMALA/.//..... r r ,.-"Fig. S8.-Distribution of troglobitic and troglophilic rhachodesmid millipeds of the genera Ceuthauxus, Pararhachistes,Acutangulus, Rhachodesmus, and Aceratophallus: 1, Ceuthauxus palmitonus; 2, C. mediator; 3, C. galeanae; 4, C. constans;5, Pararhachistes amblus; 6, Acutangulus alius; 7, Acutangulus pictus; 8, Rhachodesmus viridis; 9, R. digitatus; 10, Aceratophallusscutigeroides; n, Aceratophallus calcehtokanus; 12, Aceratophallus oxkutzcabus; 13, Aceratophallus hoctunanus.184

Acutangulus alius CauseyAcutangulus alius Causey, 1973:108, 109, fig. 3-6.Type-locaIity.-Cueva del Ojo de Agua de Tlilapan,Tlilapan, Veracruz, Mexico.Distrihution.-Known only from the type-locality.See Fig. 58.Discussion.-This is the only troglobite in the genusAcutangulus. There are presently five describedspecies in this genus from Veracruz and two undescribedspecies from Oaxaca (Causey, 1973). Acutanguluspictus Causey has been collected in Sotano deBotella Chica, Veracruz, but is probably an accidental.Acutangulus alius was found on silt and breakdownalong the edges of the first bat room in thetype-locality. It appeared to avoid the main guanodeposits.Ceuthauxus constans CauseyCeuthauxus constans Causey, 1973:111-112, fig. 910.Type-locaIity.-Grutas del Mogote, El Mogote, 15km NNE Taxco, Guerrero, Mexico.Distrihution.-Known only from two caves in theDolina de El Mogote, Guerrero. See Fig. 58.Records.-Guerrero: Cueva Chica del Mogote andGrutas del Mogote.Discussion.-This abundant troglobite is thesouthernmost species of the genus, which rangesnorth to Nuevo Leon. Other species are known fromMorelos, Veracruz, Oaxaca, Tamaulipas, and Coahuila.This species is apparently closely related to C.palmitonus Chamberlin. In Grutas del Mogote it waspresent in large numbers on silt banks. Ceuthauxusgaleanae (Chamberlin) has been taken from caves nearGaleana, Nuevo Leon, while C. mediator Chamberlinhas been found in small caves on Cuesta de Chipinquenear Monterrey, Nuevo Leon. Both species are probablyaccidentals.Ceuthauxus palmitonus ChamberlinCeuthauxus palmitonus Chamberlin, 1942:12; Chamberlin,1943b:5, 52; Bonet, 1946a:112; Chamberlin,1947:32; Nicholas, 1962:176; Causey, 1973:111.Ceuthauxius palmitonus: Bolivar, 1944:26 (erroneousspelling).Ceuthauxus palmitensis: Reddell, 1967a:24 (erroneousspelling).Strongylodesmus palmitonus: Loomis, 1968:41; Reddell,1971b:45.Type-locaIity.-Gruta del Palmito, Bustamente(=Bustamante), Nuevo Leon, Mexico.Distrihution.-Known only from the type-locality.See Fig. 58.Discussion.-This is the northernmost describedspecies of the family Rhachodesmidae. It is apparentlya relict species now restricted to its cave habitatby the surrounding desert. Other relict speciesinclude C. nuevus Chamberlin which is known onlyfrom the vicinity of the spring, Ojo de Agua, atSabinas Hidalgo, Nuevo Leon; and an undescribedepigean species taken in the entrance area of Cuevadel Carrizal, Nuevo Leon. Specimens have been collectedfrom Grutas del Palmito in the main entranceroom in moist, silted areas, from rotting wood, andfrom an oatmeal trail.Pararhachistes amblus ChamberlinPararachistes amblus Chamberlin, 1942: 13, fig. 2122; Chamberlin, 1943b:5, 53; Bonet, 1946a:112;Nicholas, 1962: 176.Pararhachistes amblus: Loomis, 1968:39; Reddell,1971b:45.Type-locaIity.-Cueva de Jutxlahuaca (=Grutas deJuxtlahuaca), Colotlipa, Guerrero, Mexico.Distribution.-Known only from the type-locality.See Fig. 58.Discussion.-The genus Pararhachistes, in additionto this troglobite, includes two epigean species fromGuerrero. Pararhachistes amblus has been collectedrecently from silt banks and rotten wood.Strongylodesmus harrisoni CauseyMilliped, new genus and species: Reddell, 1966a: 13;Reddell, 1967c:55.Rhachodesmidae, undescribed genus and species:Reddell, 1971b:44 (Tamaulipas records only).Strongylodesmus harrisoni Causey, 1971b:31; Reddelland Mitchell, 1971b:189, fig. 14; Causey,1973:113, 114, 116, fig. 17-18; Mitchell andKawakatsu, 1973a:671, 673; Reddell, 1973a:34,40; Reddell and Elliott, 1973b:181, 185; Shear,1974:261, 263, fig. 32-34; Fernandez Ruiz, 1976:717.Type-locaIity.-Cueva del Rancho del Cielo No.7,5 km NW of Gomez Farias, Tamaulipas, Mexico.Distribution.-Known from caves in the Sierrade Guatemala, Tamaulipas. See Fig. 59.Records.-Tamaulipas: ?"Sinkhole" at Ranchodel Cielo, Cueva de la Capilla, Crystal Cave, ?Sotanode Gomez Farias, Harrison Sinkhole, Cueva del Infiernillo,Sumidero de EI Jineo, ?Sotano de la Joyade Salas, Sotano del Leon, Resumidero de los Mangos,Cueva de la Mina, Sotano del Molino, Cueva de185

las Perlas, Cueva Chica de la Perra, ?Cueva del Ranchodel Cielo n. 3, Cueva del Rancho del Cielo n. 7, Cuevadel Remolino, Sotano de Tres Cerritos, Cueva del Ojode Agua de Manantiales, 2,000 Meter Cave, Sotano deVasquez, and ?Wet Cave.Discussion.-This large, delicate species ranges incolor from a pale blue-green to pure white. It is amajor element of the cave fauna of the Sierra de Guatemala.Retention of pigment in some individuals indicatesthat it is a fairly recent troglobite. It is frequentlyabundant on silt and moist flowstone. A large,epigean species, S. conspicuus Causey, is abundant onthe surface and is occasionally collected in the entranceareas of caves in the Sierra de Guatemala andSierra de El Abra. A second epigean species, S. potosianus(Chamberlin), is known from Sumidero de Fantasmas,San Luis Potosi. The genus Strongylodesmusranges from Veracruz to Tamaulipas and containseight species.Unculabes arganoi ShearUnculabes arganoi Shear, 1974:264, 265, 267,268,269, fig. 39-44.Type-locality.-Cueva del Madroiio, El Lolo (=EILobo), Queretaro, Mexico.MEXICO~,\\: I',;-"\_1- '--~. __._._.G;'.//--....... I IL\.r/ '-\._i I\"'\"..'\ i\'...,II_____' I-I//,iIjrGUATEMALAFig. 59.-Distribution ottroglobitic and troglophilic rhachodesmid miIlipeds of the genus Strongylodesmus: 1, Strongylodesmwharrisonij 2, S. harrisoni and S. conspicuus; 3, S. conspicuusj 4, S. potosianus.186

Distribution.-Known only from the type-locality.See Fig. 60.Discussion.-This species is apparently mostclosely related to U. crispus Causey, its geographicallyclosest relative. The genus Unculabes includes fivespecies, all known from caves. Four of these are certainlytroglobites. A fifth species, U. columbinusCausey, from Sotano de las Golondrinas, San LuisPotosi (Causey, 1973), is red and probably should beclassified as a troglophile. Unculabes includes severalrather divergent species, and some may eventually beremoved to other genera.Unculabes causeyae ShearUnculabes sp.: Causey, 1973:118; Reddell and Elliott,1973b:181, 185.Unculabes causeyae Shear, 1974:263, 264-265, 267,fig. 35-38.Type-locality.-Cueva Chica de la Perra, 8 mlnorthwest of Gomez Farias, Tamaulipas, Mexico.Distribution.-Known only from two caves in theSierra de Guatemala, Tamaulipas. See Fig. 60.Records.-Tamaulipas: ?Cueva de la Mina andCueva Chica de la Perra.\\\,'" '~, ,- .... j,'..--.r-,,-! "I\;(r,\ .''- -"'J ~I,----,j/r(r(/c,IMEXICO'","".jI.. ,-'"( r-"'/'_'}... _ ........ v\\'.I..~ II j~ 1_-" >,..J '-~ ._._L~'·I"--, ~ l './//-.......! j'/1 \..-1 IGUATEMALA//\")'-,,-,,\ i\..., i.-'- -----' ,-III/r'r\r',...'..;r/Fig. 60.-Distribution of troglobitic and troglophilic rhachodesmid millipeds of the genus Unculabes: 1, Unculabes causeyae;2, U. columbinus and U. crispus; 3, U. crispus and U. porrensis; 4, U. crispus; 5, U. arganoi.187

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

Speodesmus pecki ShearPolydesmidae, undescribed genus and species: Reddelland Mitchell, 1971a:150.Speodesmus pecki Shear, 1974:241, 272, 273-274,fig. 55-58; Shear, 1977a:246, 247.Type-Iocality.-Cueva del Pachon, 12 mi south ofCiudad Mante, Tamaulipas, Mexico.Distribution.-Known only from the type-locality.See Fig. 54.Discussion.-Shear (1974) placed this species inSpeodesmus with the admission that it may be representativeof an undescribed genus. Speodesmus isknown only by troglobitic species in Central Texasand southern New Mexico. An undescribed speciesoccurs in Grutas del Palmito, Nuevo Leon.Tylogoneus minus CauseyTylogoneus minus Causey, 1973:121, fig. 32; Shear,1977a:246.Type-Iocality.-Cueva de Tres Manantiales (=Cuevadel Ojo de Agua de Manantiales), Sierra de Guatemala,17 km SW Gomez Farias, Tamaulipas, Mexico.Distribution.-Known only from the type-locality.See Fig. 54.Discussion.-This species is known only by a singlemale and two fragmentary specimens of each sex. Thegenus Tylogoneus is known only by the two troglobiteslisted here. Shear (1977a) reported a female ofthis genus from Sumidero de Canada, Chiapas.Tylogoneus rainesi CauseyTylogoneus rainesi Causey, 1973:121, fig. 27-31;Shear, 1977a:246, 247.Type-Iocality.-Cueva de Poca Ventana (=Cuevadel Ahuate n. 2), Xilitla, San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 54.Discussion.-This species is known only from themale holotype.Family TridontomidaeTridontomus loomisi ShearTridontomus loomisi Shear, 1977a:241-243, fig. 1418.Type-locality.-Cueva Seamay, near Senahu, AltaVerapaz, Guatemala.Distribution.-Known only from the type-locality.See Fig. 54.Discussion.-This species is known only from themale holotype. The family Tridontomidae includesonly two species in addition to this troglobite. Theother two species, Tridontomus procerus Loomis andHoffman and Aenigmopus alatus Loomis and Hoffmanare known from Sepacuite and Seamay, AltaVerapaz, Guatemala, respectively. All three specieshave very long legs and antennae and weakly sclerotizedcuticles. Shear (1977a) speculates that the twospecies described by Loomis and Hoffman (1962)are also from caves since well known caves occur atboth Sepacuite (Cueva Sepacuite) and Seamay (CuevaSeamay).Family XystodesmidaeThe family Xystodesmidae is among the more conspicuouselements in the forest fauna of Mexico. Thisfamily has, however, not been successful in the colonizationof caves, and none are known to be troglobitesor troglophiles. The genera Cruzodesmus andRhysodesmus have both been collected from manycaves in the forested, mountain regions of Mexico.These millipeds all appear either to have entered cavesaccidentally with washed-in debris, to have fallen intovertical cave entrances, or to have utilized the moistcave entrances for temporary shelter.Order PolyzoniidaFamily PolyzoniidaeThis family is represented in Mexican caves onlyby an undetermined species of the genus Siphonotusin Cueva de la Mina, Tamaulipas. It is presumably anaccidental.Family SiphonophoridaeOne species of siphonophorid, Siphonophora sabachanaChamberlin, has been described from Mexicancaves. It was found in Actun Sabaca, Yucatan, and islisted by Nicholas (1962) as a troglobite, but isclearly an accidental or trogloxene. It has recentlybeen collected in Cueva Luchil, Yucatan. This or arelated species has also been taken from three otherYucatlin caves.Order SpirobolidaFamily AtopetholidaeThe only species of atopetholid identified fromcaves in Mexico is Aztecolus nigrior (Chamberlin).This species was found below the entrance drop intoChevy Sink, Nuevo Leon.Family MessicobolidaeThis family is represented only by undeterminedspecies of the genera Anelus and Messicobolus from190

several caves in Mexico. Their ecological status is unknown,but none are troglobites.Family RhinocricidaeTwo species of this family were described fromcaves in Yucatan by Chamberlin (1938) and are listedby Nicholas (1962) as troglobites. Rhinocricus motuiensisis known only from Cenote de Sambula, at Motul,and Yucatobolus spukilensis was described fromActun Xpukil. Both are apparently accidentals.Family SpiroboUelidaeReddellobus troglobius CauseyReddellobus troglobius Causey, 1975b:333, 334-336,337,338, fig. 3-5.Type-Iocality.-Grutas de Jonotla, 3700 ft el, 7 kmSW of Cuetzalan, Puebla, Mexico.Distribution.-Known from four caves near Cuetzalan,Puebla. See Fig. 62.Records.-Puebla: Cueva de la Barranca, Grutas deJonotla, Cueva Octimaxal SUI n. 2, and Cueva Xochitl.Discussion.-This remarkable species is a memberof the subfamily Typhlobolellinae, which is representedonly by four or possibly five described species,all of which occur in Mexico. Reddellobus troglobiusMEXICOGUATEMALAr,.. \,.....--/.//Fig. 62.-Distribution of troglobitic millipeds of the families Cambalidae and Spirobolellidae: 1, Cambala speobia; 2, Mexi·cambala inopis; 3, M. blanda; 4, M. russelli; 5, Reddellobus troglobius; 6, M. [lShi; 7, Jarmilka alba.191

appears to be most closely related to Ergene setosusChamberlin, an epigean species known from San LuisPotosi and Tamaulipas. Other typhlobollelids includetwo species of Typhlobolellus from Veracruz andpossibly Morelene from Morelos. Reddellobus troglobiuswas found in areas of organic debris and batguano in each of the recorded caves. Although alltypWobollelids lack eyes, this species has reducedpigmentation, slender body, and attenuate appendages,indicating that it should be considered to be atroglobite.Family SpirobolidaeTwo species of this family have been collectedfrom caves in Mexico. Hiltonius carpinus carpinusChamberlin was found in the entrance area of Grutasdel Mogote and Cueva Chica del Mogote, Guerrero.An undetermined genus and species has been collectedin a single cave in Nuevo Leon.Order SpirostreptidaFamily CambalidaeThe family Cambalidae is represented in the cavesof Mexico and Belize only by troglobitic species. Thisfamily is well represented in the caves of the southernUnited States but also abounds in forested areas. Nocollections of the family from the epigean environmentare known for Mexico, Guatemala, or Belize.Cambala speobia (Chamberlin)Eclytus (not Holmgren 1855) speobius Chamberlin,1952b:1O.Eclomus speobius: Chamberlin, 1952a:71; Chamberlinand Hoffman, 1958:175.Cambala caeca Loomis, 1953:417, fig. 1-3.Cambala captiosa Causey, 1959:67-71, fig. 1-3.Cambala speobia: Causey, 1964a:243-244; Causey,1971a:272, 273, fig. la, 2a; Shear, 1974:249.Cambala speobia new subspecies: Reddell, 1965b:164-165; Loomis, 1968:95.Cambala speobia subspecies: Causey, 1965:63; Mitchelland Reddell, 1971:64; Reddell, 1971b:46.Cambala: Causey, 1968:124.Type-Iocality.-Of Eclytus speobius: Wyatt Cave,Sutton (=Edwards) County, Texas, U.S.A.; of Cambalacaeca: Wyatt Cave, Edwards County, Texas,U.S.A.; of Cambala captiosa: Beck's Ranch Cave,Williamson County, Texas, U.S.A.Distribution.-Known from many caves in CentralTexas (U.S.A.) and one cave in northern Coahuila,Mexico. See Fig. 62.Mexico record.-Coahuila: Cueva de los Lagos.Discussion.-This is the common cave milliped ofCentral Texas. Its occurrence in a cave 2 km south ofthe Rio Grande in Coahuila is not surprising. Thelimestones of ValVerde County, Texas, are identicalto those in adjacent Coahuila. Specimens from Coahuilaappear to exhibit slight differences from thoseacross the Rio Grande, and, at first, these appearedto be sufficient to justify subspecific recognition forthe Mexican population. Careful study of all materialfrom Central Texas has shown that these differencesfall within the range of variation within the speciesin Texas. The species is closely related to Cambalareddelli reddelli Causey in northwestern Texas andadjacent New Mexico and C. reddelli inomatusCausey in northwest Texas. Both of these forms arealso cavernicolous. Cambala speobia was collectedfrom silt and guano in a short horizontal passage inthe upper levels of Cueva de los Lagos. This cave hasnow been inundated by the waters of the AmistadReservoir.Jarmilka alba ShearJarmilka alba Shear, 1973:44-45, fig. 1-3.Type-locality -Mountain Cow Cave, Caves Branch,Belize.Distribution.-Known only from the type-locality.See Fig. 62.Discussion.-The ecological status of this form is indoubt, and Shear (1973) expresses the opinion that itmay be a facultative troglophile also occurring in theendogean habitat. It is included here because of itslack of eyes and pigment and because all knownMexican cambalids appear to be restricted to caves.This is the smallest species in the family and the mostsouthern representative of the Cambalidae in NorthAmerica.Mexicambala blanda CauseyMexicambala russelli undescribed subspecies: Reddell,1966a:13; Reddell, 1967c:55 (all records exceptSotano de la Joya de Salas and Cueva de los Leones);Reddell, 1971b:47 (all records except Sotanode la J oya de Salas and Cueva de los Leones).Mexicambala sp.: Reddell and Mitchell, 1971b:190(all records except Cueva de la Capilla, Sotano dela Joya de Salas, and Cueva de los Leones).Mexicambala blandus Causey, 1971a:272, 275, 276,278-279, fig. lc, 2b, 3; Reddell, 1973a:35, 40.Mexicambala: Reddell, 1973b:77.Mexicambala blanda: Causey, 1973:121; Reddell andElliott, 1973b:182, 185; Shear, 1974:250.Mexicambala russelli: Vomero, 1974:350 (Sotano deGomez Farias record only).192

Type-locality.-Harrison's Sinkhole, Rancho delCielo, Tamaulipas, Mexico.Distribution.-Caves in the southern Sierra de Guatemala,Tamaulipas. See Fig. 62.Records.-Tamaulipas: Bee Cave, Crystal Cave,Sotano de Gomez Farias, Harrison Sinkhole, Sumiderode EI lineo, Sotano del Leon, Sotano del Molino,Cueva del Nacimiento del Rio Frio, Cueva dela Paloma, Grutas de EI Puente, Cueva del Remolino,Sotano de Tres Cerritos, and Sotano de Vasquez.Discussion.-The genus Mexicambala is composedonly of the four troglobitic species listed here.Causey (1971a) considers this species, together withM. inopis and M. russelli, to comprise the russellisuperspecies. They have contiguous ranges and arevery closely related.Mexicambala fishi CauseyMexicambala fishi Causey, 1971a:272, 275, 280,fig. Ie, 2d, 4; Reddell, 1973a:35, 40.Type-locality.-Cueva Arriba de Rio Iglesia (=SOtanodel Rio Iglesia), 4 mi. E Huautla, Oaxaca,Mexico.Distribution.-Known only from the type-locality.See Fig. 62.Discussion.-This species is most closely related toIVI. russelli. With the exception of the aberrant }armilkaalba, this represents the southern limit of distributionof the Cambalidae in 1exico.Mexicambala inopis CauseyMexicambala russelli undescribed subspecies: Reddell,1967c:55 (Sotano de la loya de Salas and Cuevade los Leones records only); Reddell, 1971b:47(Sotano de la loya de Salas and Cueva de los Leonesrecords only).Mexicambala sp.: Reddell and Mitchell, 1971b:190(Cueva de la Capilla, Sotano de la loya de Salas,and Cueva de los Leones records only); Mitchelland Kawakatsu, 1973a:673.Mexicambala inopis Causey, 1971a:272, 275, 276,278, 279, fig. Ib; Causey, 1973:122; Reddell,1973a:35, 40; Reddell and Elliott, 1973b:182,185; Shear, 1974:250.Type-locality.-Sotano de la loya de Salas, W ofRancho del Cielo about 6 mi., Tamaulipas, Mexico.Distribution.-Known from caves in the northernSierra de Guatemala, Tamaulipas. See Fig. 62.Records.-Tamaulipas: Cueva de la Capilla, Sotanode la loya de Salas, Cueva de los Leones, and Cuevade la Mina.Discussion.-This species occurs to the north ofand at higher elevations than M. blanda. The evolutionin the same area of two species as closely relatedas are M. blanda and M. inopis is somewhat difficultto explain. The epigean ancestor may have becomeextinct in the higher, more northern part of theSierra de Guatemala before it did so in the lower,southern area. The higher areas are generally pineforest and are drier than the cloud forest at the lowerelevations of the range of M. blanda. In the UnitedStates the Cambalidae generally prefer wet litter habitats,and we would assume the epigean ancestor ofMexicambala to have had a similar preference. Causey(1971a) speculates that Mexicambala moved southalong the eastern slopes of the Sierra Madre Orient:

this is almost certainly M. russelli. Specimens havebeen taken from moist cave walls with a fine layerof washed-in silt, from wet flowstone with smallamounts of bat guano on it, and from silt floors.Family SpirostreptidaeThe only genus of spirostreptid known from cavesin this area is Orthoporus. In addition to the twotroglobites discussed below, this genus is representedin caves in Mexico by five species (see Fig. 63).Orthoporus fraternus (Saussure) is a guanophile inGrutas de Zapaluta and Sumidero del Camino, Chiapas;O. guerreronus (Chamberlin) has been collectedin Grutas de Juxtlahuaca, Guerrero; O. solicolensChamberlin is an abundant troglophile in the caves ofCampeche and Yucatan; O. yucatanensis Causey israrely collected in caves in Campeche and Yucatanand is probably an accidental; and O. mimus Chamberlinis a troglophile in caves in San Luis Potosi andTamaulipas (Causey, 1975a). Orthoporus disc rim inuns Chamberlin has been collected in Grutas de Lanquin,Alta Verapaz, Guatemala (Causey, 1960).Nicholas (1962) lists O. guerreronus as a troglobite,but this species is probably a guanophile. Numerouscollections of this genus from many caves throughoutMexico await study.Orthoporus spelaeus CauseyOrthoporus spelaeus Causey, 1977:167, 176, 178,180, fig. 16-19; Reddell, 1977b:233, 239, 240,256.Type-Iocality.-Cenote de Catzin, Catzin, nr.Valladolid, Yucat

Class InsectaThe class Insecta is, of course, a dominant elementIn the cavemicole fauna of the world, just as it is inthe epigean fauna. Troglobites are, however, surprisinglysparse in all tropical areas, and only theApterygota have been very successful in adapting toa cave environment. Twenty-five orders of insect havebeen collected from caves in Mexico, Guatemala, andBelize. Of these only seven-Collembola, Diplura,Thysanura, Blattodea, Saltatoria, Homoptera, andColeoptera-are represented by troglobites (see Table21).Order CoUembolaCollembolans are present in essentially every cavein Mexico and Central America. A total of 16 troglobitesand 21 other species have been identified (seeTable 22), but large collections from numerous cavesin all parts of this area remain unstudied. Althoughcollembolans are most commonly found on organicdebris, animal droppings, and bat guano, they areoccasionally taken from moist flowstone, the surfaceof pools, and on cave walls.MEXICO•l , I~)GUATEMALAFig. 63.-Distrihution of troglobitic and troglophilic millipeds of the family Spirostreptidae: 1, Orthoporus mimus; 2, O.guerreronus; 3, O. fraternus; 4, O. zizicolens; 5, O. spelaeus and O. solieolens; 6, O. diseriminans; unnumbered circles, O.solieolens.195

Table 21.-Summary of cave inhabiting Insecta.Troglobites Other Species*Collembola 16 21DipluraCampodeidae 5 1Japygidae 0 4Parajapygidae 0 1ArchaeognathaMachilidae 0 1ThysanuraLepismatidae 0 2Nicoletiidae 1 2OdonataAeschnidae 0 1Coenagrionidae 0 6Cordulegastridae 0 1Libellulidae 0 13EphemeropteraBaetidae 0 1Leptophlebiidae 0 2BlattodeaBlaberidae 0 6Blattellidae 1 6Blattidae 0 1Polyphagidae 0 4IsopteraTermitidae 0 1MantodeaMantidae 0 1DermapteraForficulidae 0 2Labiduridae 0 1Labiidae 0 1PlecopteraPerlidae 0 1*Saltatoria 4 37Embioptera 0 1PsocopteraCaeciliidae 0 1Epipsocidae 0 1Liposcelidae 0 1Pachytroctidae 0 1Psocidae 0 1Psyllipsocidae 0 2Troctopsocidae 0 1MallophagaMenoponidae 0 1Philopteridae 0 1*Hemiptera 0 63*Homoptera 2 10MegalopteraCorydalidae 0 1NeuropteraMyrmeleontidae 0 1*Coleoptera 25 246SiphonapteraHystrichopsyllid ae 0 1lschnopsyllidae 0 3Pulicidae 0 3*Diptera 0 97TrichopteraCalamoceratidae 0 1Hydropsychidae 0 1PhilopotamidaePolycentropidae*Lepidoptera*HymenopteraTotalTable 21.-(Continued)Family EntomobryidaeThe best-represented family in this area is the Entomobryidae.Nine species of troglobite have beenidentified, but others doubtless await determination.Cave-adapted forms have been described in the generaCyphoderus, Metasinella, Pseudosinella, and Troglopodetes.These troglobites and related troglophilesare discussed below. At least two species of troglobiticDicranocentruga from caves in Oaxaca andVeracruz are presen tly under study. Mills (1938)reported lsotomus sp., Lepidocyrtus pearsei Mills,and Proisotoma centralis Denis from caves in Yucatan.These are all troglophiles. Bonet (1953a) reportedLepidocyrtus sp. from caves in the Xilitlaregion of San Luis Potosi and Queretaro.Cyphoderus innominatus MillsCyphoderus innominatus Mills, 1938:183, 186, 188,fig. 17-19; Pearse, 1945:174; Delamare Deboutteville,1948a:349, 350, 412; Nicholas, 1962:178;Salmon, 1964:537; Reddell, 1971b:49; Reddell,1977b:234,239,240,268,274,276,289.Type-Iocality.-San Bulha Cenote (=Cenote deSambula), Motul, Yucatan, Mexico.Distribution.-Known from four caves in Yucatan.See Fig. 64.Records.-Yucatan: ActUn Gongora, Cueva MuruztUn,Cenote de Sambula (Motul), and Cueva Segundadel Camino a San Roque.Discussion.-This is a white, eyeless species mostclosely related to the African termitophile C. limboxiphiusBomer.ooooTable 22.-Summary of cave inhabiting Collembola.Troglobites111581653Other SpeciesEntomobryidae 9 5Hypogastruridae 2 3lsotomidae 0 3Neelidae 0 3Oncopoduridae 2 0Onychiuridae 0 3Poduridae 1 3Sminthuridae 2 ITotal 16 2154*For summary by families see Tables 22-29.196

Metasinella falcifera (Mills)Sulcuncus falciferus Mills, 1938:183, 188, 190, fig.25-27; Ahsolon and Kseneman, 1942:22, fig. 20;Pearse, 1945:175; Salmon, 1964:483; Reddell,1971b:49; Massoud and Gruia, 1973:335.Metasinella falcifera: Bonet, 1944a:20, 22, 23; DelamareDeboutteville, 1949:119; Vandel, 1964:200-201; Vandel, 1965a:168; Massoud and Gruia,1973:338, 339; Reddell, 1977b:234, 239, 240,241,270.Sulcuncus: Pearse, 1945:174; Salmon, 1964:131,483.Sulcunus falciferus:spelling).Nicholas, 1962:179 (erroneousType-locaIity.-Sazich Cave (=Acfun Sazich), Calcehtok,Yucatan, Mexico.Distribution.-Known only from the type-locality.See Fig. 64.Discussion.-This is a white, eyeless species. Metasinellaalso includes four cavernicole species fromCuba.Pseudosinella bonita ChristiansenPseudosinella bonita Christiansen, 1973: 129-130,131, fig. 1..r._r-....._._.,r .,"'\r-.,r 1·"\/ \\"\(-,~.l-, '\\ \" "'~(., \\ ...-.~_.-MEXICO(/GUATEMALA.;5 ///.r\r ,......;Fig. 64.-Distribution of troglobitic and troglophilic collembolans of the families Entomobryidae and lsotomidae: 1, Pseudosinellaviolenta; 2, P. reddelli; 3, P. reddelli and P. petrustrinatii; 4, P. petrustrinatii; 5, P. finca; 6, P. bonita; 7, Cyphoderusinnominatus and Troglopedetes maya; 8, Metasinella falcifera; 9, Proisotoma centralis.197

Type-Iocality.-Cueva Bonita del Presidente, 2 kmN Huautla, Oaxaca, Mexico.Distribution.-Known only from the type-locality.See Fig. 64.Discussion.-Pseudosinella bonita is white and eyeless.It is the most highly evolved troglobite in thegenus in Mexico. Five species of Pseudosinella areknown from Mexican caves. Of the two troglophiles,one, P. violenta (Folsom), is abundant in caves inCentral Texas; it is also known from caves in Coahuilaand Durango. The other troglophile, P. reddelliChristiansen, is a widespread, variable species rangingfrom Coahuila south to southern San Luis PotosI. Anundescribed troglobite, reported by Bonet (1953a)as Pseudosinella sp., is known from caves in theXilitla region, San Luis PotosI.Pseudosinella Jinca ChristiansenPseudosinella Jinca Christiansen, 1973:129, 130-131,fig. 4.Type-Iocality.-Cueva Sepacuite, Alta Verapaz,Guatemala.Distribution.-Known from three caves in AltaVerapaz, Guatemala, and one in Oaxaca, Mexico. SeeFig. 64.Records.-GUATEMALA: Alta Verapaz: Grutasde Lanquin, Cueva Seamay, and Cueva Sepacuite.New record.-MEXICO: Oaxaca: Cueva del Nacimientodel Rio San Antonio (det. K. Christiansen).Discussion.~This species is eyeless and pigmentless.It appears to be most closely related to P. petrustrinatii.Pseudosinella petrustrinatii Christiansen-New NamePseudosinella strinatii Christiansen, 1973:130, 131132, fig. 2 (not P. strinatii Gisin, 1951); Reddelland Elliott, 1973a:175; Reddell and Elliott,1973b:182,185.Type-Iocality.-Crystal Cave, Rancho del Cielo,5 km WGomez Farias, Tamaulipas, Mexico.Distribution.-Known from caves in the Sierra deGuatemala, Tamaulipas; Sierra de EI Abra, San LuisPotosi and Tamaulipas; and one cave in the Sierra elPino, San Luis Potosi. See Fig. 64.Records.~an Luis Potosi: Cueva Chica, Cueva deLa Lagunita, Cueva de los Monos, Cueva Pinta, Cuevade Taninul n. 1; Sotano del Tigre, and Sotano de Yerbaniz;Tamaulipas: Crystal Cave, Cueva de la Florida,Cueva de EI Pachon, Cueva de la Paloma, and Cuevade San Rafael de los Castros.Discussion.-Dr. Kenneth Christiansen has suggestedthat the new name, Pseudosinella petrustrinatii,be used for P. strinatii. "Due to an oversight,P. strinatii Christiansen (1973) is a junior homonymof P. strinatii Gisin (1951). I propose the new namePseudosinella petrustrinatii to replace P. strinatiiChristiansen" (Christiansen, in litt.). Pseudosinellapetrustrinatii is pigmentless and eyeless, and has beendivided by Christiansen into three forms, which mayeventually prove to be distinct species. The typelocality,Crystal Cave, is the only locality for Type C.Type A occurs only in the Sierra de EI Abra; Type Bis known from the Sierra de EI Abra and from Cuevade La Lagunita.Troglopedetes maya (Mills)Trogolaphysa maya Mills, 1938:183, 184, 186, fig.13-16; Arle, 1939:29; Absolon and Kseneman,1942:22, fig. 21; Bonet, 1942:59; Pearse, 1945:175; Delamare Deboutteville, 1951:293; Nicholas,1962:179.Troglopedetes maya: Bonet, 1942:59; Salmon, 1964:531; Reddell, 1971b:50; Massoud and Gruia,1973:339; Gueorguiev, 1974:683; Reddell,1977b:234, 239, 240, 271, 285-286.Trogolaphysa: Delamare Deboutteville, 1951:293,294,295, fig. 65; Salmon, 1964:134.Type-Iocality.-Balaam Canche Cave (=Grutas deBalankanche), Chichen Itza, Yucatan, Mexico.Distribution.-Known from two caves in Yucatan.See Fig. 64.Records.-Yucatan: Grutas de Balankanche andActUn Xkyc.Discussion.-This white, eyeless species is closelyrelated to T. delamarei Massoud and Gruia from acave in Cuba. Undescribed troglobites in this genusare known from caves in Guerrero and Veracruz.Family HypogastruridaeTwo species of the family Hypogastruridae (Acherontidesatoyacense and A. potosinus) are troglobites.In addition, three species of troglophile have beenidentified from Mexican caves (see Fig. 65). Acherontiellasabina Bonet is a frequently collected speciesin caves in San Luis Potosi, Tamaulipas, and NuevoLeon (Bonet, 1945; 1946b). Brachystomella parvula(Schaeffer) was identified by Bonet (1953a) fromCueva de la Hoya, San Luis Potosi. Bonet (1945)described Willemia persimillis bulbosa from Cueva deLos Sabinos, San Luis Potosi.Acherontides atoyacense BonetAcherontides atoyacense Bonet, 1945:28-31, fig. 2431; Bonet, 1946a:114; Bonet, 1946b:52, 55, 56,57, 62; Bonet, 1953a:23, 30; Stach, 1959:435;Nicholas, 1962:179; Salmon, 1964:189; Gruia,198

\"1970:197, 198, 199; Reddell, 1971b:50; Hiither,1975:287.Acherontides atoyacence: Delamare Deboutteville,1948b:49, 55; Thibaud, 1963:288, 295-296,fig. 3C (erroneous spelling).Type.locaIity.-Cueva (ooGrutas) de Atoyac, Atoyac,Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 65.Discussion.-The genus Acherontides includesseven species, of which the two troglobites listed hereare the only Mexican species. The other five speciesare known from Rumania, Japan, and Afghanistan.Acherontides atoyacense is eyeless and pigmentless.Acherontides potosinus BonetAcherontides potosinus Bonet, 1946b:52, 53-57, 62,fig. 1-2; Delamare Deboutteville, 1948b:49, 55;Bonet, 1953a:23, 24, 30, 33, 41, 48, 54, 55, 71,72,85,86; Stach, 1959:434; Nicholas, 1962:179;Thibaud, 1963:288; Salmon, 1964:189; Reddell,1967d:l06; Gruia, 1970:197, 198, 199; Moller,1970:5; Reddell, 1971b:50; Hiither, 1975:286,287.'\ , ,'- "-\)\"- '- I\)-/'\'I\\\-~" 1" \~ "-').1 I \,'l.~./MEXICO-,GUATEMALAII, ItII• I,,'-....'\i....,t," I_._.ll_r....·1/.//r\rr~IIIiIiFig. 65.-Distribution of troglobitic and troglophilic collembolans of the family Hypogastruridae: 1, Acherontiella sabina;2, Acherontides potosinus; 3, Acherontides atoyacense.199

Type-Iocality.-Cueva de El Jobo, El Jobo, SanLuis Potosi, Mexico.Distribution.-Known from caves in the Xilitlaregion, San Luis Potosi. See Fig. 65.Records.-San Luis PotoSI: Cueva del Aire, Cuevade los Cuchos, Cueva de la Roya, Cueva del J obo, andCueva del Salitre.Discussion.-This is an eyeless, pigmentless speciesclosely related to A. atoyacense.Family IsotomidaeTwo troglophilic species of the family Isotomidae(Folsomia sp. and Folsomina onychiurina Denis) wereidentified by Bonet (1953a) from caves in the Xilitlaregion, San Luis Potosi. Proisotoma sp. has been collectedin Sotano de Yerbaniz, San Luis Potosi; andP. centralis Denis was reported by Mills (1938) fromCenote Yunchen, Yucatan.Family NeelidaeBonet (1947) in his monograph on the familyNeelidae reported Neelus murinus Folsom fromCueva de Los Sabinos, San Luis Potosi; N. murinusbolivari Bonet from Cueva de San ta Elena, Yucatan;and Megalothorax minimus Willem from caves InNuevo Leon and Yucatan. All are troglophiles ortrogloxenes.Family OncopoduridaeOncopodura atoyacense BonetOncopodura atoyacense Bonet, 1943:133, 140, 145147, fig. 14-16; Bonet, 1946a:114; Nicholas,1962:179; Salmon, 1964:297; Reddell, 1971b:51.Oncopodura atoyacensis: Gueorguiev, 1974:683(erroneous spelling).Type-Iocality.-Cueva (=Grutas) de Atoyac, Atoyac,Veracruz, Mexico.Distribution.-Known only from the type.locality.See Fig. 66.Discussion.-The genus Oncopodura is representedby several species in Europe, one in Japan, one troglobitein Montana (U.S.A.), and the two Mexican troglobiteslisted here.Oncopodura prietoi BonetOnchopodura: Anonymous, 1942b:265 (erroneousspelling).Oncopodura prietoi Bonet, 1943:133, 141, 147-150,fig. 17·22; Bolivar, 1944:26; Bonet, 1946a:114;Nicholas, 1962:179; Salmon, 1964:298; Reddell,1971b:51; Reddell and Elliott, 1973b:182·186.Oncopodura sp.: Reddell and Mitchell, 1971b:191.Oncopodura prietori: Gueorguiev, 1974:683 (erroneousspelling).Type-Iocality.-Gruta del Palmi to, Bustamante,Nuevo Leon, Mexico.Distribution.-Known from the type-locality andpossibly one cave in the Sierra de Guatemala, Tamau·lipas. See Fig. 66.Records.-Nuevo Leon: Grutas del Palmito; Tamaulipas:?Bee Cave.Discussion.-This eyeless species is closely relatedto several European species. The record of this speciesfrom Bee Cave is based on a single specimen. Additionalmaterial is needed to verify this identification.Family OnychiuridaeAt least three species of the family Onychiuridaehave been identified from Mexican caves. Mesaphorurakrausbaueri Borner and Onychiurus sp. were reportedby Bonet (1953a) from caves in the Xilitlaregion, San Luis PotosI. Onychiurus fimetarius(Linnaeus) has been collected in Resumidero dePablillo, Nuevo Leon. Mesaphorura foveata Bonetwas reported from Cueva Chica, San Luis Potosi(Bonet, 1944b).Family PoduridaeIn addition to the troglobitic Spelaeogastruraguerrerense Bonet, three other podurids have beenidentified from Mexican caves (see Fig. 66). Mills(1938) described the troglophile Xenylla yucatanafrom Cenote de Sambula (Motul), Yucatan. Recentcollections have included Paranura caeca Folsomfrom Grutas del Mogote, Guerrero, and Tafallia sp.from Cueva del Huisache, San Luis PotosI.Spelaeogastrura guerrerense BonetSpelaeogastrura guerrerense Bonet, 1945:22-27, fig.1-10, 17-23; Bonet, 1946a:114; Nicholas, 1962:179; Salmon, 1964:239; Reddell, 1971b:51.Speleogastrura: Delamare Deboutteville, 1951:272(erroneous spelling).Spelaeogastrura: Salmon, 1964:108,239.Type-locality.-Gruta de Cacahuamilpa, Cacahuamilpa,Guerrero, Mexico.Distribution.-Known only from the type-locality.See Fig. 66.Discussion.-This eyeless, pigmentless species is theonly member of the genus. Spelaeogastrura is possiblymost closely related to the genus Schafferia.200

Family SminthuridaeThe family Sminthuridae includes one describedtroglobite, Pararrhopalites anops, in Mexico. A speciesof the genus Sminthurus was reported by Bonet(1953a) as a troglobite in the caves of the Xilitla region,San Luis Potosi. Temeritas sp. has been recentlycollected in Sotano de la Tinaja, San Luis Potosi; itsecological status is unknown.Pararrhopalites anops Bonet and TellezPararrhopalites anops Bonet and Tellez, 1947:194198, 202, fig. 1-16; Cassagnau and DelamareDehoutteville, 1953:143; Stach, 1956:62; Nicholas,1962:179; Salmon, 1964:576; Reddell,1967a:24; Reddell, 1971h:51.Type-locality.-Cueva (=Grutas) del Palmito, Bustamante,Nuevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 66.Discussion.-Pararrhopalites includes only the troglobitelisted here and P. oculatus Bonet from Isla dela Roqueta, Acapulco, Guerrero, Mexico.MEXICOGUATEMALA///r\r·'-..-"Fig. 66.-Distribution of troglobitic and troglophilic collembolans of the families Oncopoduridae, Poduridae, and Sminthuridae:1, Oncopodura prietoi and Pararrhopalites anops; 2, O. prietoi; 3, Spelaeogastrura guerrerense; 4, O. atoyacense;5,Xeny/la yucatana.201

Order DipluraFamily CampodeidaeAlthough campodeids may be found in almostevery moist cave in Mexico and Central America, veryfew species have been described. In addition to thefive troglobites discussed below, Wygodzinsky (1944)described Campodea (Campodea) chica from CuevaChica, San Luis Potosi (see Fig. 67). This is presumablya troglophile. Large collections of campodeidsremain unstudied. Campodeids are usually found insoil, under rocks, and in debris; troglobites may befound on cave walls or running among rocks on thecave floor.luxtlacampa hauseri Condeluxtlacampa hauseri Conde, 1975:421-424, fig. 1-2;Strinati, 1977:388.Type-locaIity.-Cueva Chirrepeck, Alta Verapaz,Guatemala.Distribution.-Known only from the type-locality.See Fig. 67.Discussion.-This species is closely related to 1.juxtlahuacensis.luxtlacampa juxtlahuacensis Wygodzinskyluxtlacampa juxtlahuacensis Wygodzinsky, 1944:368, 374-377, fig. 4; Bonet, 1946a:113; Conde,MEXICO~ \./,'GUATEMALAFig. 67.-Distribution of troglobitic and troglophilic diplurans of the family Campodeidae: 1, Podocampa cavernicola andParatachycampa boneti; 2, Campodea (Campodea) chica; 3, Litocampa atoyacensis; 4, Juxtlacampa juxtlahuacensis; 5, J.hauleri.202

1955:40, 60, 62, 64, 170, 187; Padt, 1957:33;Nicholas, 1962:180; Reddell, 1971b:48; Conde,1975:421,424.luxtlacampa: Conde, 1955:31, 48, 51, 94, 166,178; Padt, 1957:4; Vandel, 1964:203; Vandel,1965a: 170; Delamare Deboutteville, 1971:62.Type-locality.-Cueva (=Grutas) de Juxtlahuaca,Colotlipa, Guerrero, Mexico.Distribution.-Known only from the type-locality.See Fig. 67.Discussion.-The genus luxtlacampa indudes onlythe two species listed here, although Padt (1957)considered leannelicampa stygin Conde from Algeriato properly belong in luxtlacampa. This is dearlyincorrect.Litocampa atoyacensis (Wygodzinsky)Plusiocampa (Litocampa) atoyacensis Wygodzinsky,1944:368-370, fig. 1; Conde, 1949: 137; Conde,1955:124; Nicholas, 1962:180; Reddell, 1971b:49.Plusiocampa atoyacensis: Conde, 1955:130, 170,187.Litocampa (Cocytocampa) atoyacensis: Padt, 1957:27.Litocampa atoyacensis: Conde, 1975:421.Type-locality.-Cueva (=Grutas) de Atoyac, Atoyac,Veracruz, Mexico.Distribution.-Known only from the type-locality.See Fig. 67.Discussion.-This troglobite has been placed byPadt (1957) in the subgenus Cocytocampa. This subgenusis a heterogeneous assemblage of species fromEurope, South America, and Mexico. It is certainlynot a natural grouping of species. I am, therefore,following the usage of Conde (1975).Paratachycampa boneti WygodzinskyParatachycampa boneti Wygodzinsky, 1944:368,371-374, fig. 2-3; Conde, 1955:42,45, 72-73, 77,170,187; Nicholas, 1962:180; Reddell, 1967a:24;Reddell, 1971b:49; Conde, 1975:421.Plusiocampa boneti: Bonet, 1946a:113; Padt, 1957:4.Plasiocampa (Stygiocampa) boneti: Paclt, 1957:45.Paratachycampa: Conde, 1955:31, 42, 50, 94, 166,173; Vandel, 1964:203; Vandel, 1965a:170;Delamare Deboutteville, 1971:62.Type-locality.-Grutas de Garcia (=Grutas deVilla de Garcia), Garcia, Nuevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 67.Discussion.-Paratachycampa is a monotypic genus.Padt (1957) synonymized the genus with Plusiocampaand placed P. boneti in the subgenus Stygiocampaalong with species from Bulgaria, Yugoslavia,and Australia. This is certainly not a natural assemblageof species and does not reflect the true relationshipsof this distinctive troglobite; therefore, at thesuggestion of Dr. Pedro Wygodzinsky (pers. comm.),I am retaining the original classification.Podocampa cavernicola (Wygodzinsky)Parallocampa (Parallocampa) cavernicola Wygodzinsky,1944:368,380-382, fig. 7; Conde, 1955:166,170,187; Nicholas, 1962:180; Reddell, 1971b:48.Parallocampa cavernicola: Bonet, 1946a: 113; Reddell,1967a:24.Meiocampa (Remycampa) cavernicola: Paclt, 1957:31.Parallocampa: Vandel, 1964:203; Vandel, 1965a:170; Delamare Deboutteville, 1971:62.Podocampa cavernicola: Conde, 1975:421.Type-locality.-Grutas de Garcia (=Grutas deVilla de Garcia), Garcia, Nuevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 67.Discussion.-This species was originally placed inParallocampa, a complex of North American species.Paclt (1957) has removed some of these species, includingP. cavernicola, and placed them in the Moroccansubgenus Remycampa. This is certainly an artificialgrouping. I am here following the usage ofConde (1975) in placing this species in the genusPodocampa.Family JapygidaeOur knowledge of the japygids of Mexico is extremelylimited. Only one species has been positivelyidentified. Allojapyx allodontus Silvestri, an epigeanspecies described from southern Mexico, has beenreported from Sotano de la Tinaja, San Luis Potosi.Careful study of the holotype of this poorly knownspecies may reveal that the San Luis Potosi specimensbelong to an undescribed species. Other generaknown from Mexico include Metajapyx and Mixojapyx.Several undescribed species of troglophile areknown. Japygids are usually found under rocks, inleaf litter, and in rotting wood. In some Mexicancaves, however, large specimens have been collectedfrom moist silt and clay banks.Family ParajapygidaeSilvestri (1948) reported Parajapyx mexicanusSilvestri from Cueva de Carroza, Yucatan. This family203

of small soil inhabiting japygoids has been found inseveral caves in Mexico and Guatemala, but the specimensall remain unstudied.Order ArchaeognathaFamily MachilidaeMachilids are seldom collected in caves, and onlyone species has been found in a Mexican cave. An unidentifiedspecies was collected below the entrancedrop of Sotano de Gomez Farias, Tamaulipas. Carefulstudy of the litter in the vicinity of entrances shouldreveal additional machilids.Order ThysanuraFamily LepismatidaeTwo species of the family Lepismatidae have beenidentified from caves in Mexico. Allacrotelsa spinulata(Packard) was collected from cave swallow guanoin the entrance of Cueva del Salitre, Chihuahua. Wygodzinsky(1972) reported Ctenolepisma ciliata (Dufour)from the entrance area of Cueva del Diablo,Chihuahua. Both species are probably trogloxenes oraccidentals.Family NicoletiidaeThe family Nicoletiidae is a frequently encounteredgroup in the caves of Mexico and Central America.Numerous species of troglobite and troglophilehave been collected from all parts of this area, butwith one exception they await description. Troglobiticthysanurans are usually found running alongclay or silt banks or on the cave walls. The troglophilicspecies are more commonly found on thecave floor under and among debris and rocks. A recordof the troglobitic species, Texoreddellia texensis (Ulrich),from Grutas de Quintero, Tamaulipas (Paclt,1971) is in error. Texoreddellia is a genus describedby Wygodzinsky (1973) for a highly evolved troglobitefrom Central Texas. Although known fromnumerous caves in Texas, it has not been reliablyreported from Mexico. Wygodzinsky (1973) reportsthat he has considerable material from Grutas deQuintero, none of which is Texoreddellia. The speciesprobably occurs in northern Coahuila. A single individual,which could not be captured, was seen inCueva de los Lagos, Coahuila, immediately across theilio Grande from caves inhabited by T. texensis.Unfortunately, this cave is now inundated by thewaters of the Amistad Reservoir.Anelpistina anophthalma (Bilimek)Lepisma anophthalma Bilimek, 1867:905; Packard,1894:732; Silvestri, 1912:204; Wolf, 1934-1938:vol. 2 :484, vol. 3: 143; Jeannel, 1943: 144; Vandel,1964:205; Vandel, 1965a:171.Lepisma anophthalmum: Packard, 1888:88 (erroneousspelling).Lepisma cacahuamilpensis: Herrera, 1891:219, pI. II(fig. 1b); Packard, 1894:732.Lepisma anopthalma: MacGillivray, 1891:270 (erroneousspelling).Lepisma (?) anophtlalma: Silvestri, 1912:221 (erroneousspelling).Nicoletia anophthalma: Escherich, 1904:133, 140,155.Nicoletia: Bolivar, 1940:126.Nicoletia (Anelpistina) anophtalma: Wygodzinsky,1946:15-17, 19, 20, 21, fig. 1-2 (erroneous spelling).Nicoletia (Anelpistina) anophthalma: Paclt, 1963:47.Anelpistina: Wygodzinsky, 1967:519,523.Anelpistina anophthalma: Reddell, 1971b:48.Type-Iocality.-Of Lepisma anophthalma: Hohle(=Grutas de) Cacahuamilpa in Mexiko; of Lepismacacahuamilpensis: Caverna (=Grutas) de Cacahuamilpa,Guerrero, Mexico.Distribution.-Known only from the type-locality.See Fig. 68.Discussion.-This species is a member of a genusranging from southern Mexico into the southern UnitedStates, with one species in Peru (Wygodzinsky,1967). A closely related species, A. boneti (Wygodzinsky),is a troglophile in Grutas de Juxtlahuaca,Guerrero. Anelpistina is a frequent inhabitant ofcaves in Mexico, as both troglophiles and troglobites,but most of the species remain undescribed.Order EphemeropteraFamity BaetidaeThis family is represented in Mexican caves onlyby an unidentified genus and species collected inGrutas de Olivares, Puebla.Family LeptophlebiidaeTwo species of this family have been collected inMexican caves. Choroterpes (Neochoroterpes) mexicanusAllen was taken in Cueva de la Puente, San LuisPotosi (Allen, 1974). Nymphs of Thraulodes litotesAllen were found in a stream in total darkness inCueva del Brinco, TamauJipas. Both of these speciesare certainly troglophiles.Order OdonataEighteen species of dragonflies and damselflieswere reported from cenotes in Yucatan (Williamson,204

1936). Although these doubtless utilize the cenotesas places in which to deposit their eggs, they are certainlynot part of the true cave fauna. Nymphs of afew additional species have been collected from caveswhich receive floodwaters, but none should he consideredmore than accidentals.Order BlattodeaRoaches are frequently found in caves, both in theentrance area and in total darkness, and are certainlyimportant aspects of the cavernicole fauna. Large collectionsfrom all parts of Mexico await study, and thebrief summary of the families recorded from caveswill doubtless be considerably expanded when thismaterial is studied.Family BlaberidaeThe most conspicuous roaches which inhabit cavesare the larger species of the family Blaberidae. Blaberusatropos (Stoll) and B. craniifer Burmeister havebeen reported from caves in Yucatan (Pearse, 1938b).The latter species is also extraordinarily abundant inGrutas de Juxtlahuaca, Guerrero, where it nearlycovers the walls and floor of the principal bat room.Fisk (1977) also reported this species from Cueva delRancho San Miguel, Chiapas. Blaberus giganteusGUATEMALA/, ./r,.... ,...--Fig. 68.-Distribution of troglobitic and troglophilic thysanurans of the family Nicoletiidae: 1, Undescribed troglobiticNicoletiidae; 2, Anelpistina anophthalma; 3, A. boneti.205

(Linnaeus) has been collected from Cueva de SanRafael de los Castros, Tamaulipas. Pycnoscelus surinamensis(Linnaeus) was found in the entrance room ofBee Cave, Tamaulipas; it has also been reported byFisk (1977) from caves in Chiapas, Mexico, and Huehuetenango,Guatemala; and by Fisk (pers. comm.)from Cueva de Taninul n. 1, San Luis Potosi, andCueva de EI Pachon, Tamaulipas. An undescribedspecies of Panchlora was reported from Las TresCuevas (=Cueva de Sala de Agua Grande), Veracruz(Fisk, 1977).Family BlatteUidaeUndetermined species of the genera Chorisoneura,Ischnoptera, and Latiblatella have been collectedfrom caves in several parts of Mexico. Fisk (1977)reported undetermined species of A naplecta, Euthlastoblatta,and Ischnoptera from caves in Chiapas.Agalopteryx chiapas was described by Fisk (1977)from five caves in Chiapas. Nesomylacris reddelli wasdescribed by Fisk and Gurney (1972) from Cuevade EI Pachon, Tamaulipas. Fisk (1977) described N.lateralis from Cueva de Chital n. 1, Chiapas. Pseudomopsseptentrionalis Hebard has been collected fromcaves in San Luis Potosi and Tamaulipas. Fisk (1977)described P. nigrimaculis from Cueva del Aguacero,Chiapas. The most remarkable roach known fromMexican caves is a delicate, eyeless species of thegenus Nelipophygus, unfortunately known onlyfrom one nymph collected in Cueva de Chital n. 2,Chiapas (Fisk, 1977). This is the first species of thegenus known from mainland North America. An undescribedtroglobite of the genus is known fromcaves on Jamaica. Other troglobitic roaches areknown from Hawaiian caves.Family BlattidaeThis family is known from Mexican caves only byan undetermined species of Periplaneta from SanLuis Potosi and Tamaulipas.Family PolyphagidaeThe family Polyphagidae is well represented inMexican caves. The genus Arenivaga is frequentlycollected from the dry entrance areas of caves inCoahuila and Durango. Holocompsa azteca Saussurewas reported from Cueva del Venado, Chiapas (Fisk,1977). Holocompsa zapoteca Saussure was found inmany caves in Yucatan (Pearse, 1938b) and in Cuevadel Sabin, Chiapas (Fisk, 1977). Homoeogamia mexicanaBurmeister has heen found in caves in Guerrero,Hidalgo, Mexico, Michoacan, and San Luis Potosi.Table 23.-Summary of cave inhabiting Saltatoria.TroglobitesOther SpeciesAcrididae 0 1Gryllidae 4 14Rhaphidophoridae 0 21StenopelJuatidae 0 3Tettigoniidae 0 1Total 4 40Order IsopteraFamily TennitidaeTermites are rarely collected from rotten woodwashed into caves. The only Mexican record is of afew workers of the family Termitidae collected fromCenote Amil, Yucatan.Order MantodeaFamily MantidaeImmature and unidentified mantids have beencollected rarely from the entrance area of caves.Pearse (1938b) reported a specimen from AccinSabaca, Yucatan.Order DennapteraFamily ForficulidaeTwo species of this family of earwigs have beencollected from caves in Mexico: Ancistogaster sp. d.toltecus (Scudder) from Cueva de la Puente, San LuisPotosi; and A. impennis Bormas from Cueva de Carnicerias,San Luis Potosi. Both species are probablytrogloxenes.Family LabiduridaeThe only species of this family recorded fromMexican caves is Euborellia annulipes (Lucas) fromGrutas de Juxtlahuaca, Guerrero. Its ecological statusis unknown.Family LabiidaeThis family is represented in Mexican caves onlyby Labia rotundata Scudder from Pozo Melendez,Guerrero. It is probably an accidental.Order PlecopteraFamily PerlidaeThe only stone fly recorded from Mexican cavesis Anacroneuria sp., collected from the entrance areaof Sima Esteban, Puehla. It is prohably an accidental.206

Order SaltatoriaThe cricket fauna of Mexico and Central Americais poorly known, and the 41 species identified to daterepresent only a small percentage of the actual numberof species which inhabit caves in this region (seeTable 23). Extensive collections await study.Family AcrididaeA single species of the family Acrididae, Necaxacrissp. d. micans (Hebard), has been collected inSotano del Profesor, Veracruz. It is certainly an accidental.Family GryUidaeThe family Gryllidae is well represented in thecaves of Mexico and Central America. The most commonlycollected genera are Paracophus and Amphiacusta.Paracophus includes three troglobitic and fivedescribed troglophilic species in northeastern Mexico(Hubbell, 1972). Additional collections include undescribedspecies of troglophile. Paracophus apterusChopard is known from many caves in San Luis Potosiand Tamaulipas; P. placonotus Hubbell is abundantin the caves of the Xilitla region in San LuisPotosi and Queretaro; P. reddelli Hubbell is knownfrom two caves at low elevations in the Sierra deGuatemala, Tamaulipas; P. sanctorum Hubbell isknown only from Cueva de la Virgen, Tamaulipas;and P. subapterus Chopard occurs in caves in NuevoLeon and Coahuila. The crickets of the genus Paracophusare usually found in darkness under rocksand on silt banks. Troglobitic species are more frequentlyseen on the walls and on speleothems.Paracophus apterus and P. placonotus are the onlyspecies which have been collected from epigeanforest habitats. The only other troglobitic gryllidin Mexico is Tohila atelomma from Yucatan. SeeFigs. 69-70 for localities of Paracophus and Tohilain Mexico.The genus Amphiacusta ranges from Belize andGuatemala north into southern Tamaulipas, Mexico.It is frequently found near cave entrances where itmay be heard stridulating. Four species have beenidentified from caves in Mexico; many other speciesfrom Belize, Guatemala, and Mexico remain to bedescribed. Chopard (1947) reported A. azteca Saussurefrom a cave at Tepoztlan, Morelos; A. bolivariChopard from Grutas de Atoyac, Veracruz; and A.maya Hubbell from Cueva de Berriozabal, Chiapas.Hubbell (1938) described A. yucatana from numerouscaves in Yucatan; it is also now known fromcaves in Campeche. Amphiacusta bolivari has beenreported by Hubbell (1972) from several additionalcaves near Orizaba and Cordoba, Veracruz. Saussure(1897) described Arachnomimu8 cavicola fromGrutas de Lanquin, Alta Verapaz, Guatemala. Thecorrect generic assignment of this species is still indoubt. See Fig. 71 for the localities of Amphiacustaand Arachnomimus in the caves of Mexico and Guatemala.Undetermined species of Cycloptilum,Gryllus, Miogryllus, and Nemobius have been collectedfrom various caves in Mexico; all are probablyaccidentals.Paracophus caecus HubbellParacophus undescribed species: Reddell, 1966a:14.Paracophus sp.: Reddell, 1971b:55 (Harrison Sinkhole,Sotano de la Joya de Salas, Sotano del Molino,Cueva del Nacimiento del Rio Frio, andSotano de las Piftas (=Pinos) records only]; Reddelland Mitchell, 1971b:191, fig. 16.Paracophus sp. A: Reddell, 1967c:55.Paracophus sp. B: Reddell, 1967c:55.Paracophus caecus Hubbell, 1972:51,52,57,58,67,70, 76, 77, 80-84, 86, 87, fig. 1(4), 2(17), 3(28),5(37); Mitchell and Kawakatsu, 1973a:671; Mitchelland Reddell, 1973b:frontispiece; Reddell,1973a:35, 40; Reddell and Elliott, 1973b:182,186; Vomero, 1974:341,350.Paracophus, troglobio: Fernandez Ruiz, 1976:717.Type-Iocality.-Sotano de la Joya de Salas, Tamaulipas,Mexico.Distribution.-Known from caves in the Sierra deGuatemala, Tamaulipas. See Fig. 69.Records.-Tamaulipas: Cave at Rancho del Cielo,Bee Cave, Cueva de la Capilla, Crystal Cave, Sotanode Gomez Farias, Harrison Sinkhole, Cueva del Infiernillo,Sotano de la Joya de Salas, Cueva de laMina, Sotano del Molino, Cueva del Nacimiento delRio Frio, Sotano de los Pinos, Cueva del Ojo de Aguade Manantiales, Cueva de los Vampiros, and WetCave.Discussion.-This is the only eyeless gryllid in theNew World. It is also wingless and very pale withelongate appendages. The genus Paracophus is mostclosely related to the Yucatan cave genus Tohila. Allof the troglobitic species apparently evolved in thehigher mountains and, with the exception of sixpopulations of P. caecus, are known only from highelevations. It is interesting that some of the lowlandpopulations of P. caecus are also somewhat less paleand delicate, indicating that they have maintainedcontact with the parent species longer. Paracophuscaecus has been taken in association with P. apterusin Sotano de Gomez Farias, Cueva del Nacimiento delRio Frio, Cueva del Ojo de Agua de Manantiales, andCueva de los Vampiros; and with P. reddelli in Bee207

Cave. Paracophus caecus appears to have descendedfrom a common ancestor from which P. apterus andP. placonotus and their allies also later developed(Hubbell, 1972). Paracophus caecus is found underrocks, but it is also often collected from cave wallsand from among stalactites. It has been observedcopulating in several caves while sitting on stalagmitesor stalactites.Paracophus cladonotus HubbellParacophus sp.: Reddell, 1967d:106; Reddell, 1971b:55 (Sotano del Pozo and Sotano de Tlamayarecords only).Paracophus cladonotus Hubbell, 1972:51,52,57,58,74, 76, 77, 80, 84-87, fig. 1(8),2(9-10, 16),3(18,22,26-27),5(36); Reddell, 1973a:35, 40.Type-locaIity.-Sotano de Tlamaya, San Luis Potosi,Mexico.Distribution.-Known from caves in southern SanLuis Potosi and adjacent Hidalgo. See Fig. 69.Records.-Hidalgo: Cueva de Piedra Ancha andCueva de EI Tenango; San Luis Potosi: Sotano deGuadalupe, Cueva de la Luz, Cueva de los Potrerillos,Sotano del Pozo, Cueva de San Nicolas, and Sotanode Tlamaya.GUATEMALA,... ...;./, ./rr'"'Fig. 69.-Distribution of troglobitic crickets of the family Gryllidae: 1, Paracophw caecus; 2, P. lippus; 3, P. cladonotus;4, Tohila atelomma.208

Discussion.-This species is known from caves inthe Xilitla and Aquism6n regions, San Luis Potosi,and from the geologically contiguous parts of Hidalgo.The eyes of P. cladonotus are very small and theappendages elongate, but not to the extent that theyare in P. caecus. This species probably also evolvedfrom the ancestral stock which gave rise to P. caecus.Paracophus cladonotus has been taken with P. placonotusin Cueva de El Tenango, Hidalgo, and Cueva delos Potrerillos and S6tano de Tlamaya, San Luis Potosi.Paracophus lippus HubbenParacophus lippus Hubbell, 1972:51, 52, 57, 58, 74,77-78, 80, 82, fig. 1(3), 2(15), 3(25), 4(34); Redden,1973a:35, 40.Type-Iocality.-Cueva de La Lagunita, BlaggRanch, 15 mi. NE of Ciudad del Maiz, San Luis Potosi,Mexico.Distribution.-Known only from the type-locality.See Fig. 69.Discussion.-This species also has very small eyes,reduced pigmentation, and elongate appendages. It isGUATEMALA -----,/r.... r· "'..;./ ../rFig. 70.-Distrihution of troglophilic gryllid crickets of the genus Paracophus: 1, Paracophus subapterus; 2, P. sanctorum;3, P. apterus; 4, P. apterus, P. reddelli, and P. placonotus; 5, P. apterus and P. placonotus; 6, P. placonotus.209

the most poorly known species in the genus, and adiscussion of its affinities must await careful descriptionof the male genitalia. It has been taken in associationwith a possibly undescribed species of troglophilicParacophus.Tohila atelomma HubbellTohila atelomma Hubbell, 1938:191-192, 194-200,204, fig. 1-10, 77; Jeannel, 1943:156; Pearse,1945:175; Chopard, 1947:67; Nicholas, 1962:180; Vandel, 1964:217; Vandel, 1965a:181; Leroy,1967:665, 684, 685, 686, 709; Chopard,1968:262; Delamare-Deboutteville and Botosaneau,1970:70; Reddell, 1971b:55; Hubbell,1972:53; Reddell, 1977b:234, 239, 240, 258,261, 266, 286, 288.Tohila: Hubbell, 1938:191,201,202; Hubbell, 1972:54-56.Type-Iocality.-Chac Mol Cave (=Cueva Chac Mol),Tohil, Yucatan, Mexico.Distribution.-Known from caves in the state ofYucatan. See Fig. 69.Records.-Yucatan: Grutas de Balankanche, CuevaChac Mol, Cueva Luchil, Cueva Oxolodt, Cueva deSan Isidro, and Cueva del Cenote Xtolok.New records.-Yucatan: Cenote Aka Chen, CenoteCalchum, Cenote de Sambula (Motul), Grutas deTzab-Nah, and Actun Xpukil (det. T. H. Hubbell).GUATEMALAFig. 71.-Distribution of troglophilic gryllid crickets of the genera Amphiacusta and Arachnomimus: 1, Amphiacustaazteco; 2, Amphiaculta bolivari; 3, Amphiacusta maya; 4, Amphiacusta yucataTUJ; 5, Arachnomimus cavicola.210

Discussion.-The genus Tohila was originally assignedto the subfamily Pentacentrinae and consideredto be most closely related to the genus Trigonidomimusof the Old World (Hubbell, 1938). Hubbell(1972) has reexamined these genera and feelsnow that Tohila is most closely related to Paracophusand that they are probably both members of the subfamilyPhalangopsinae. Tohila is a monotypic genusand is small and pale with degenerate eyes. It is usuallyfound beneath rocks, but it is also occasionallytaken from moist flowstone or cave walls. It has beencollected from several caves which are also inhabitedby the troglophilic Amphiacusta yucatana. Numerouscollections of crickets from caves in the YucatanPeninsula await study, and some will certainly proveto contain this species.Family RhaphidophoridaeThe camel crickets of the family Rhaphidophoridaeare frequently found in the caves of Mexico andGuatemala. The 12 described species, however, representa small percentage of the fauna to be reportedeventually from caves. The described species belongto the genera Ceuthophilus, Anargyrtes, Leptargyrtes,Exochodrilus, Hypsobadistes, and Phoberopus. Severalother genera await description. The Rhaphidophoridaeappear to be absent from low elevations alongthe eastern flanks of the Sierra Madre Oriental southinto the Yucatan Peninsula. The caves in this regionare instead populated by species of the gryUid genusAmphiacusta. Amphiacusta ill this area inhabits theentrance zone usually occupied by rhaphidophoridsin other regions.Hubbell (1972) erected the genus Anargyrtes fortwo species from the states of Mexico and Guerrero.Anargyrtes annulata (Bilimek) was described fromGrutas de Cacahuamilpa, Guerrero; it is now knownalso from many caves in Guerrero and Mexico. Anargyrtesbolivari Hubbell is known primarily from epigeanlocalities, but it has been found in a lava cavenear San Geronimo, Mexico (see Fig. 72).The genus Ceuthophilus is the principal cavecricket genus inhabiting caves in the southwesternUnited States and northern Mexico. Only two species,Ceuthophilus (Geotettix) cunicularis Hubbell and C.(Ceuthophilus) variegatus Scudder, have been identifiedfrom Mexican caves. The first species is knownonly from Cueva de los Lagos, Coahuila, and the latterhas been collected in several caves in Coahuila andChihuahua (see Fig. 72). Many undescribed species ofCeuthophilus have been collected in caves in northernMexico.Hubbell (1972) described a new genus and twonew species for the camel crickets inhabiting thehigher elevations of the Sierra de Guatemala, Tamaulipas.Exochodrilus caelestis occurs in two caves atRancho del Cielo, and E. forcipatus is known fromcaves at higher elevations (see Fig. 72).The genus Leptargyrtes was erected by Hubbell(1972) to include two new species from Queretaro:L. boneti from caves near El Lobo in eastern Queretaro,and L. tejamanilae from caves to the west nearPinal de Amoles and Tejamanil (see Fig. 72).Hubbell (1977) described the Phoberopes groupto include several species of camel cricket from Guatemalaand Chiapas, Mexico. Phoberopus minor Hubbellis known from Cueva de Chemal n. 1 and Cuevade Santa Eulalia (=Cueva de los Resadores), Huehuetenango,Guatemala. Three species of long-legged,delicate crickets were placed in the new genus Hypsobadistes:H. gracilior Hubbell from two caves in Huehuetenango,Guatemala;H. stuarti Hubbell from Cuevadel Rayo de San Felipe, Chiapas; andH. tenuis Hubbellfrom three caves at San Agustin, Chiapas (see Fig. 72).Apparently undescribed species of the generaPristoceuthophilus and Argyrtes have been collectedfrom caves in Coahuila and Hidalgo respectively.Family StenopehnatidaeUnidentified species of the stenopelmatid generaAnabropsis, Glaphyrosoma, and Stenopelmatus havebeen collected from Mexican caves, but they are allprobably accidentals.Family TettigoniidaeA single unidentified tettigoniid of the genusDichopetala was collected below the entrance dropinto Cueva de Taninul n. 4, San Luis Potosi; it iscertainly an accidental.Order EmbiopteraAn undetermined genus and speci~ of the seldomcollected order Embioptera was found in Sotano deEI Triunfo, San Luis Potosi. It is presumably an accidental.Order PsocopteraThe order Psocoptera is occasionally taken fromcaves in Mexico and is doubtless far more abundantthan the few records would indicate. This group includestroglobites in other parts of the world, but allof the species known from Mexican caves are probablytroglophiles or trogloxenes.Family CaeciliidaeAn undetermined species of the genus Caeciliuswas collected in Sotano de El Triunfo, San Luis Potosi.Its ecological status is unknown.211

Family EpipsocidaeThe family Epipsocidae is represented in Mexicancaves only by undetennined species of the genus Epipsocuscollected in caves in Chiapas, Puebla, andVeracruz.Family Lipo8celidaeUndetennined species of the genus Liposcelis havebeen taken in Grutas de San Bartolo, Nuevo Leon,and in Grutas de Atoyac, Veracruz.Family PachytroctidaeThe family Pachytroctidae is represented in Mexicancaves only by an undetennined species ofPachytroctestaken from cave swallow guano in ActUnTucil, Yucatan.Family PsocidaeThis family is represented in Mexican caves byCerastipsocus sp. prob. trifasciatus (Prov.) from Grutade Cuevacillas, Coahuila. It is probably a trogloxene.GUATEMALA/./rr\..... r.--Fig. 72.-Distribution of cavernicole crickets of the family Rhaphidophoridae: 1, Ceuthophilus (Geotettix) cunicularis;2, C. (Ceuthophilu,) variegatu,; 3, Exochodrilus !orcipatu,; 4, E. caelestis; 5, Leptargyrtes boneti; 6, L. tejamanilae; 7, Anargyrtyesbolivari; 8, A. annulata; 9, Hyp,obadistes ,tuarti; 10, H. tenuis; 11, Phoberopus minor; 12, H. gracilior.212

Family PsyllipsocidaeTwo species of the family Psyllipsocidae areknown from Mexican caves (see Fig. 73). Psyllipsocusramburii Selys is a species found in caves in manyparts of the world. In Mexico it is known from cavesin Coahuila, Durango, Chihuahua, Nuevo Leon, SanLuis Potosi, Tamaulipas, Puebla, Veracruz, and Chi·apas. This species is usually found in dry areas, bothin total darkness and the twilight zone. It is fre·quently found on rotting wood or bits of paper.Psyllipsocus yucatan Gurney was described fromCueva del Cenote Xtolok, Yucatan (Gurney, 1943).Family TroctopsocidaeMockford (1967) has described the remarkablepsocid Protroctopsocus enigmaticus from leaf litterin the entrances of caves on Cuesta de Chipinque,Nuevo Leon. This species is also known from leaflitter in Durango (E. L. Mockford, pers. comm.).Order MallophagaFamily MenoponidaeThe only record of this family from Mexican cavesis of Myrsidea sp., taken from Petrochelidon fulvaMEXICOGUATEMALAFig. 73.-Distribution of troglophilic p60copterans of the family Psyllipsocidae: 1, Psyllipsocus ramburii; 2, P. yucatan.213

citata Van Tyne in Cenote de Piste, Yucatan (Klaas,1968).Family PhilopteridaeThis family of lice is represented in Mexican cavesonly hy Philopterus excisus Nitzch from Petrochelidonfulva citata in Cenote de Piste, Yucatan (Klaas,1968).Order HemipteraThe order Hemiptera is represented in Mexicancaves hy 63 species in 25 families. Many of these speciesare known only from open·air cenotes in Yucatanor from entrance areas and will not be discussed here.Although troglobitic hemipterans are known fromcaves in Hawaii, no troglohitic species have heenfound in North or Central America. Several, however,are troglophiles, and at least one emesine reduviidshows some signs of hecoming adapted to cave existence.Suborder HydrocorizaeFamily BelostomatidaeHungerford (1936) reported two species of thegenus Belostoma and one of Lethocerus from openaircenotes in Yucatan. Abedus immensus Menke is atroglophile in Cueva de la Puente and Cueva de laLaguna, San Luis Potosi. Abedus signoreti Mayr isknown from Sotano del Molino, Tamaulipas; its ecologicalstatus is unknown. An undetermined speciesof the genus Belostoma is extremely ahundant inCueva del Azufre, Tabasco; numerous individualscarrying eggs were observed and it is certainly atroglophile. Belostoma subspinosum Beauvois isprobably a troglophile in Sotano del Lienzo, San LuisPotosI.Suborder AmphibicorizaeFamily GerridaeHungerford (1936) reported four species of gerridsfrom open-air cenotes in Yucatan. Two species havebeen collected from Mexican caves and are apparentlytroglophiles: Gerris remigis Say from Sotano de SanFrancisco n. 2, San Luis Potosi; and Trepobates pictus(Herrich-Schaeffer) from Cenote Poxil, Yucatan.An undetennined species of Trepobates was found inCenote Aka Chen, Quintana Roo.Family VeliidaeHungerford (1936) reported two species of Microveliafrom open-air cenotes in Yucatan. Numerous undetenninedspecies of this genus have been recentlycollected from many caves in Mexico. Rhagoveliavaripes Champion is a troglophile in caves in HidalgoTable 24.-Summary of cave inhabiting Hemiptera.TroglobitesOther SpeciesHydrocorizaeBelostomatidae 0 6Corixidae 0 1Gelastocoridae 0 1Naucoridae 0 2Nepidae 0 2Notonectidae 0 3Ochteridae 0 1AmphibicorizaeGerridae 0 5Hebridae 0 1Hydrometridae 0 1Mesoveliidae 0 1Veliidae 0 5GeocorizaeCeratocombidae 0 1Cimicidae 0 4Cydnidae 0 7Dipsocoridae 0 1Enicocephalidae 0 1Lygaeidae 0 5Miridae 0 2Pyrrhocoridae 0 3Reduviidae 0 6Rhopalidae 0 1Schizopteridae 0 1Thyreocoridae 0 1Tingidae 0 1Total 0 63and San Luis Potosi. Velia sp. is an apparent troglophilein Grutas del Mogote, Guerrero.Suborder GeocorizaeFamily CimicidaeFour species of bedbug of the family Cimicidaehave heen collected from Mexican caves. Cimexhemipterus Fahricius was reported hy Pearse (1938b)from caves in Yucatan. Hoffmann (1972) reportedCimex pilosellus (Horvath) from Cuevas del Cerro deXictle, Distrito Federal. The rare hat hedhug Primicimexcavernis Barher, has heen taken from Cueva deJanitzio, Michoacan, and Cueva de Valladolid, Yucatan(Ueshima, 1968). Primicimex cavernis and Cimexincrassatus Usinger and Ueshima are known from ahat cave at Chocoyos, Chimaltenango, Guatemala(Sailer, 1950; Usinger, 1966). Primicimex cavernis isotherwise known only from Ney Cave and Fern Cave,Texas (U.S.A.). This species lives in crevices in thewalls of the chamhers inhabited by bats and duringthe day emerges to feed on the resting bats.Family CydnidaeSeven species of the family Cydnidae have beenfound in Mexican caves (see Fig. 74). All are probably214

troglophiles usually found in guano. Amnestus pusio(Stal) was taken in caves in Oaxaca and Yucatan; A.subferrugineus (Westwood) has been found in caves inOaxaca and San Luis Potosi, and in "cave earth atCacao, Trece Aguas, Guatemala" (Froeschner, 1960).The most frequently encountered genus of cydnid inMexican caves isPangaeus. Pangaeus (pangaeus) aethiops(Fabricius) was found in caves in San Luis Potosiand Veracruz; P. (P.) docilis (Walker) has been foundin caves in Campeche, Oaxaca, Quintana Roo, SanLuis Potosi, Tamaulipas, Veracruz, and Yucatan; P.(P.) moestus (Stal) has been found only in ActunHuachap, Campeche; P. (p.) piceatus Stal is knownfrom caves in Chiapas, Oaxaca, San Luis Potosi, andYucatan. Tominotus unisetosus Froeschner has beenfound in caves in Guerrero and Yucatan.Family ReduviidaeThe family Reduviidae is frequently found in thevicinity of cave entrances. This family is of specialinterest since many members of it are capable oftransmitting Chagas disease. Several specimens of thesubfamily Reduviinae have been collected from thedry entrance area of caves in Coahuila, Nuevo Leon,and Durango. Opisthacidius mexicanus (pelaez) hasMEXICOGUATEMALA-'",--r,..-"/.//Fig. 74.-Distribution of troglophilic hemipterans of the family Cydnidae: 1, Pangaeus (Pangaeus) docilis; 2, P. (P.) dodlis,P. (P.) piceatus, and P. (P.) aethiops; 3, P. (P.) dodlis and P. (P.) piceatus; 4, P. (P.) dodlis; 5,P. (P.) piceatus; 6, P. (P.) dodlisand P. (P.) aethiops; 7, Tominotus unisetosus; 8, P. (P.) moestus.215

een found in caves and mines in Yucatan. Two speciesof reduviids in Yucatan and Quintana Roo havebeen found to be vectors of Trypanosoma cruzi Chagas,the protozoan which is responsible for Chagasdisease (Gonzalez-Angulo and Ryckmen, 1967). Triatomadimidiata (Latreille) is known from caves inAlta Verapaz, Guatemala; Cayo District, Belize; andYucatan, Mexico. T. hegneri Mazzotti was collectedfrom a cave on EI Rancho Santa Rita, Quintana Roo.Lent and Wygodzinsky (1979) reported T. longipennisUsinger from a bat cave in Sinaloa. The mostinteresting group of reduviids inhabiting Mexicancaves are the thread-legged bugs of the subfamilyEmesinae. Wygodzinsky (1966) described Ploiariamaya from an unidentified cave in Yucatan. Thissmall-eyed species has been taken from cave wallsin total darkness in several additional caves in Yucatan.Immature emesines, some belonging to the genusPloiaria, have been found in caves in San Luis Potosiand Veracruz.Table 25.-Summary of cave inhabiting Homoptera.TroglobitesOther SpeciesAuchenorrhynchaCicadellidae 0 6Cixiidae 1 1Fulgoridae 0 1Kinnaridae 1 0Membracidae 0 1StemorrhynchaOrtheziidae 0 1Total 2 10Type-locaIity.-Cueva de Emilia, 14 km. NNWAhuacatlan, Queretaro, Mexico.Distribution.-Known only from the type-locality.See Fig. 75.Discussion.-This species has the eyes and ocelliabsent, has the tegmina and wings reduced, and ispale and poorly scierotized.Order HomopteraTwelve species of the order Homoptera have beenidentified from caves in Mexico. Although most homopteransare accidentals or trogloxenes, at leasttwo are troglobites (see Table 25).Family CicadellidaeLeafhoppers of the family Cicadellidae are occasionallycollected in the entrance area of caves. Balcluthasp., Empoasca sp., Hortensia similis (Walker),Planicephalus flavicosta (Stal), and Xestocephalussp. have all been found in the heavily vegetated entranceareas of caves in Yucatan. Gyponana germari(Stal) was taken at the bottom of the drop into S6tano de las Golondrinas, San Luis Potosi.Family CixiidaeCixiid fulgoroids are not infrequently collectedfrom caves, and one described species is a troglobitein Mexico. Pearse (1938b) reported Cixius sp. fromActUn Gongora, Yucatan. An undetermined speciesof this genus has recently been collected from Cuevade los Camarones, Puebla. Undetermined cixiids, includingtwo probable troglobites, have been found incaves in Oaxaca, Puebla, Veracruz, and QuintanaRoo. The only other troglobitic cixiids are in the genusOliarus and are known from lava caves in Hawaii.Cixius orcus FennahCixius orcus Fennah, 1973:439,440-442, fig. 1-10.Family KinnaridaeOeclidius hades FennahOeclidius hades Fennah, 1973:439,442-444, fig. 1117.Type-locaIity.-Cueva de Valdosa, 8 mi. E Valles,San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 75.Discussion.-This is the only described troglobitein the family Kinnaridae. An apparent new speciesof troglobite is known from Cueva de la Mina, Tamaulipas,and a possible troglobite is known fromCueva Macinga, Veracruz. The eyes and ocelli in O.hades are absent, the body is pale yellowish brown,and the wings are reduced to scales.Order MegaIopteraFamily CorydalidaeThe large aquatic larvae of the dobsonfly CorydaIus sp. have been found in Guerrero (Sbordoni andArgano, 1972) and Veracruz. They are presumablyaccidentals.Order NeuropteraFamily MyrrneleontidaeAn antlion, Eremeleon longior Banks, was describedfrom caves in Yucatan (Banks, 1938). Adults216

and larvae of this species have been collected recentlyin many caves in Campeche, Quintana Roo, and Yucatan.This is apparently a troglophile. The adults areusually seen resting on the cave walls near entrances,while the larvae inhabit silty areas in dry parts of thecaves.Order ColeopteraThe order Coleoptera is well represented in thecaves of Mexico and Central America. Althoughabout 270 species have been identified from caves inthis region, only 25 are known to be troglobites (seeTable 26). This is an amazingly low number of troglobitesfor so vast and diverse an area, but it has alreadybeen noted many times that the beetle faunaof tropical regions is very poorly represented bytroglobites (Vandel, 1964). The reasons for this arenot clear, and only additional collecting and a betterunderstanding of tropical faunas in general will helpto explain the paucity of troglobitic beetles in thetropics. The following discussion of the beetle faunaof this region can, of course, only emphasize thosefamilies which have made a significant contributionto the cavemicole fauna. Because of their frequentGUATEMALAr"'\,... .,..'.-'/./Fig. 75.-Distribution of troglobitic and troglophilic homopterans of the families Cixiidae and Kinnaridae: 1, Kinnaridaegen. et sp.; 2, Oeclidius hades; 3, Cixius orcus; 4, Cixiidae gen. et sp.217

ground dwelling habit, many beetles are washed intocaves with organic debris; most of these constitute inno way a part of the true cave fauna.Suborder AdephagaFamily CarabidaeThe family Carabidae is naturally the largest contributorto the beetle fauna of Mexican caves, as it isin most of the world. More than 70 species have beenidentified from the caves in this region, of which 17are troglobites and discussed below. Many of the carabidgenera of significance to the cavernicole ecologyare in serious need of revision, and identifications arenot presently possible. Agonum, Amara, Anisotarsus,Apenes, Ardistomis, Bembidion, Clivina, Platynus,Schizogenius, Selenophorus, and Taehys are all frequentlycollected in caves. A few species in some ofthese and in a few other genera have been identifiedand are briefly discussed here.Among the agonine carabids collected in Mexicancaves are many species of Platynus (s. lat.). Most ofthese remain unidentified, but several are known.Platynus aeuminatus (Chevrolat) has been found incaves in San Luis Potosi and Queretaro; P. melanoenemis(Chaudoir) has been collected in Cueva dePuente Fierro, Oaxaca; P. segregatus (Bates) has beenfound in Grutas de Juxtlahuaca, Guerrero; and P.strictieollis (Bates) has been taken in Sotano de Tlamaya,San Luis Potosi. Platynus eolibor Whitehead(=Colpodes bie%r Chaudoir) is a distinctive speciesknown from several caves in Chiapas and Guerrero.Platynus (Stenoplatynus) umbripennis (Casey) is atroglophile in caves in the states of Guerrero and Mexico(Bolivar and Hendrichs, 1965; Barr et al., 1968).Many species of the genus Rhadine are known astroglobites in the caves of Central Texas (Barr, 1974),but all of the known Mexican cavernicole species aretroglophiles. Rhadine araizai araizai (Bolivar) was describedfrom Grutas del Palmito, Nuevo Leon (Bolivar,1944). It has since been found in Cueva del Pedregoso,Coahuila. A possible new subspecies of R.araizai has been collected from Cueva de los Lagos,Coahuila. Bolivar and Hendrichs (1964) describedfour species of Rhadine from Mexican caves. Rhadinemedellini is known only from Cueva de Carnicero,San Luis Potosi. Rhadine rotgeri was described fromGruta de Cuevacillas, Coahuila; it has recently beencollected in Cueva Abaja de Carreterra, Tamaulipas,and Sumidero de Matehuala, San Luis Potosi. Twospecies assigned to Rhadine by Bolivar and Hendrichs(1964) were placed by Whitehead (1973) inthe genus Platynus. One of these, Platynus boneti,is known only from Cueva de la Boca, Nuevo Leon;the other, P. pelaezi, is known only from Grutas deVilla de Garcia, Nuevo Leon. An undescribed speciesof Rhadine is known only from Sumidero 552, SanLuis Potosi. See Fig. 76 for localities from whichMexican cavernicole Rhadine are known.Among other carabids identified from Mexicancaves are Apenes obseura Chaudoir from Bee Cave,Tamaulipas; JlrJasoreus sp. from Grutas de Xtacumbilxunam,Campeche; Tachys (Taehys) proximusSay from Cueva de los Lagos, Coahuila; T. (Tachyuralunistriatus (Bilimek) from Grutas de Cacahuamilpa,Guerrero, and Grutas de la Estrella, Mexico;and an undescribed species of Pterostichus (IthytoIus) from caves in the Sierra de El Abra, San LuisPotosi (see Fig. 77). Paehyteles urrutiai Bolivar isan unusual species known only from caves in theSierra de El Abra, San Luis Potosi and Tamaulipas(Bolivar, 1952) (see Fig. 77).The remaining genera of cavernicole carabids areall represented by troglobites and are discussed indetail below.Antroforceps bolivari BarrAntroforeeps bolivari Barr, 1967a:66-70, fig. 1; Reddell,1971b:60; Reddell and Mitchell, 1971b:193;Whitehead, 1972:196-197; Reddell, 1973a:35, 41;Table 26.-Summary of cave inhabiting Coleoptera.TroglobitesOther SpeciesArchostomataMicromalthidae 0 1AdephagaCarabidae 17 56Dytiscidae 0 7Gyrinidae 0 1PolyphagaAderidae 0 1Alleculidae 0 3Anthicidae 0 1Bruchidae 0 2Cantharidae 0 2Chrysomelidae 0 6Cleridae 0 1Coccinellidae 0 3Colydiidae 0 1Curculionidae 0 2Dascyllidae 0 1Dermestidae 0 3Dryopidae 0 1Elateridae 0 4Elmidae 0 1Endomychidae 0 2Euglenidae 0 1Helodidae 0 1Histeridae 4 14Hydrobiidae 0 1Hydrophilidae 0 5218

Table 26.-(Continued)TroglobitesOther SpeciesLampyridae 0 1Leiodidae 3 14Limnichidae 0 1Lycidae 0 1Meloidae 0 1Melyridae 0 1Mordellidae 0 2Mycetophagidae 0 1Nitidulidae 0 4Oedomeridae 0 1Passalidae 0 2Platypodidae 0 1Pselaphidae 0 4Ptilidae 0 1Ptilodactylidae 0 1Ptinidae 1 1Rhizophagidae 0 1Scaphidiidae 0 1Scarabaeidae 0 18Scolytidae 0 1Scydmaenidae 0 7Silphidae 0 2Silvanidae 0 1Staphylinidae 0 34Tenebrionidae 0 22Trogidae 0 1Total 25 246Reddell and Elliott, 1973b:182; Vomero, 1974:351; Erwin et al., 1977:1l.Scaritini: Barr, 1968b: 184-185.Type-Iocality.-Sotano de la loya de Salas, 25kilometers west of Encino, Tamaulipas, Mexico.Distribution.-Known only from the type-locality.See Fig. 77.Discussion.-Antroforceps is a monotypic genusbelonging to the tribe Scaritini. Only two other speciesof this tribe (both from Europe) are known tobe troglobites. These are very different from Antroforceps,however, and apparently not closely related.Antroforceps bolivari is eyeless, rufotestaceous, andhas very elongate appendages. Unfortunately, it isknown from a single female. Vomero (1974) hassuggested that it is an endogean, but its occurrencein the lower levels of the cave indicates that it is atroglobite.Chiapadytes bolivari Vigna TagliantiTrecchino troglobio: Sbordoni et al., 1977:56.Chiapadytes bolivari Vigna Taglianti, 1977:327,328333, fig. 1-2.Type-locality.-Cueva de la Planta n. 2, Las Piedrecitas,S. Cristobal de las Casas, Chiapas, Mexico.Distribution.-Known only from two caves at LasPiedrecitas, Chiapas. See Fig. 78.Records.-Chiapas: Cueva de la Planta n. 1 andCueva de la Planta n. 2.Discussion.-The record for Cueva de la Planta n. 1is based only on a pair of elytra. This large species haselongated appendages and the eyes reduced to small,pale areolae. This monotypic genus appears to beclosely related to Paratrechus (Hygroduvalius).Mayaphaenops sbordonii Vigna TagliantiTrechino troglobio: Sbordoni et al., 1977:70.Mayaphaenops sbordonii Vigna Taglianti, 1977:333,334-339, fig. 3-4.Type-locality.-Resumidero Chico, La Capellania,Huehuetenango, Guatemala.Distribution.-Known only from the type-locality.See Fig. 78.Discussion.-This is a small, slender eyeless speciesknown only from the holotype male. Although themonotypic genus Mayaphaenops clearly belongs tothe Paratrechus group of genera, it is very distinct inmany respects.Mexanillus sbordonii Vigna TagliantiAnillini, troglobite: Vigna Taglianti, 1973:153-154.Mexanillus sbordonii Vigna Taglianti. 1974:309,317,318, 320-323, fig. 5-6; Sbordoni et al., 1974:26(nomen nudum); Vomero, 1974:353.Type-Iocality.-Cueva de Tio Ticho, Comitan,Chiapas, Mexico.Distribution.-Known only from the type-locality.See Fig. 77.Discussion.-This troglobite is one of only two speciesof the tribe Anillini described from Mexico. Theother species, Geocharidius zullinii Vigna Taglianti, isan endogean from Comit

Type-Iocality.-Sotano de Tejamanil, 4.5 Km westof Pinal de Amoles, Queretaro, Mexico.Distribution.-Known from three caves in Queretaro.See Fig. 78.New records.-Queretaro: Sotano de la Escuelaand Cueva del Judio (det. T. C. Barr, Jr.).Discussion.-The genus Mexaphaenops is knownonly from caves at comparatively high elevations. Inaddition to the four species listed here, three speciesfrom Tamaulipas and Nuevo Leon await description.Barr (1967c) postulates that the genus has evolvedfrom a Paratrechus-like ancestor. Mexaphaenops eleganshas been taken from flows tone and from rottenwood. It was rather abundant among the debris leftfrom a campfire built in Cueva del Judio.Mexaphaenops fishi BarrTrechine (part): Barr, 1967a:66.Mexaphaenops fishi Barr, 1967d:162-164, 165, fig. 3,6; Barr, 1971:115-116; Laneyrie, 1971:192; Reddell,1971b:61; Vigna Taglianti, 1972:118, 126,127, fig. 5; Elliott and Reddell, 1973:192, 197;GUATEMALA///r\r~ ..-"Fig. 76.-Distribution of troglobitic and troglophilic carabid beetles of the genera Rhadine, Platynus, and Speocolpodes:I, Rhadine araizai; 2, R. pelaezi; 3, Platynus boneti; 4, R. rotgeri; 5, P. medellini; 6, Platynus (Merisphodrus) n. sp. 1;7, P. (M.) profundus; 8, Platynus (Merisphodros) n. sp. 2; 9, P. tlamayaensis and Platynus (Merisphodros) n. sp. 3; 10, Platynus(Merisphodros) n. sp. 3; 11, P. (M.) gertschi; 12, Platynw (Merisphodros) n. spp. 4-5; 13,P. (M.) veraecrocis; 14,P. (M.)urquijoi; 15,P. (Stenoplatynus) umbripennis; 16, Speocolpodes franiai.220

Reddell, 1973a:3S, 41; Vigna Taglianti, 1977:326,333,338.Mexaphaenops new species: Barr, 1968b:183.Mexaphaenops fiski: Erwin et aI., 1977:24 (erroneousspelling).Type-locality.-SmaIl cave in VaIle de los Fantasmas,45 Km east of San Luis Potosi on the highwayto Valles, San Luis Potosi, Mexico.Distribution.-Known only from two caves atVaIle de los Fantasmas, San Luis Potosi. See Fig. 78.Records.-San Luis Potos{, Cave (Valle de los Fantasmas)and Sotano de la Golondrina.Discussion.-Mexaphaenops fishi is probably mostclosely related to M. prietoi. It is known from twocaves at 2,800 meters in elevation. It is interestingthat only two troglobites are known from the Vallede los Fantasmas region and that both are quite rare.It is possible that this area has been too cold and dryat times to support large numbers of species of thelitter-associated fauna which appears to have suppliedmost of the troglobites known from Mexican caves.Mexaphaenops intermedius BarrMexaphaenops intermedius Barr, 1971:113, 115, fig.1-2; Reddell and Mi!Ch.~ll, 1971b:193, fig. 19;GUATEMALA,......;,/r/,./Fig. 77.-Distribution of troglobitic and troglophilic carabid beetles of the genera Tachy., Antroforcep., Mexani/lu., Pochytele.,and Pterostichu.: 1, Tachy. (Tachy.) proximu.; 2,Antroforcep. bolivari; 3, Pachytele. urrutiai; 4,Pachytele. urrutiai andPtero.tichu. (Ithytolus) n.sp.; 5, Ptero.tichu. (Ithytolu.) n. sp.; 6, Tachy. (Tachyura) uniltriatul; 7. Mexani/lu••bordonii.221

Vigna Taglianti, 1972:118, 121-127, fig. 3-5;Reddell, 1973a:35, 41; Reddell and Elliott,1973b:182, 186; Vomero, 1974:351,353; Erwinet al., 1977:24; Vigna Taglianti, 1977:326, 333,338; Rossi and Cesari Rossi, 1977:374,376.Mexaphaenops: Sbordoni and Argano, 1972:8.Mexaphenops intermedius: Mitchell and Kawakatsu,1973a:673 (erroneous spelling).Paratrechus: Peck, 1973c:129 (misidentification).Type-locality.-Cueva de la Capilla, 13.5 km NWof Gomez Farias, Tamaulipas, Mexico.Distribution.-Known from two caves 10 theSierra de Guatemala, Tamaulipas. See Fig. 78.Records.-Tamaulipas: Cueva de la Capilla andCueva de la Mina.Discussion.-This species has been taken frommoist flowstone and among rotten wood. Barr (1971)considers this species to be intermediate in severalrespects between the more generalized M. elegans andthe highly specialized M. prietoi. It has been taken ingeneral association with Platynus (Mexisphodrus) profundusand Ptomaphagus (Adelops) troglomexicanus.GUATEMALA///r'\.... r'.--Fig. 78.-Distrihution of troglobitic and troglophilic carabid beetles of the tribe Trechini: 1, Mexaphaenops prietoi; 2,Mexaphaenops n. sp. 1; 3, Mexaphaenops n. spp. 1-3; 4, Mexaphaenops intermedius; 5, Mexaphaenops jlShi; 6, Mexaphaenopselegans and Paratrechus (Hygroduvalius) pallescens; 7, P. (Paratrechus) tepoztlanensis; 8, Paratrechus (Paratrechus) n. sp.;9, P. (P.) me:ricanu.r; 10, Mexitrechus coarctatus; 11, Chiapadytes bolivari; 12, Mayaphaenops sbordonii.222

Rossi and Cesari Rossi (1977) described the ascomycetefungus Laboulbenia sbordonii as a parasite of M.intermedius.Mexaphaenops prietoi BolivarMexaphaenops prietoi Bolivar, 1942:351, 352, 353354, fig. 1-6; Bolivar, 1944:26; Barr, 1967a:65-66;Barr, 1967d:161, 162,164, 165; Reddell, 1967a:24; Barr, 1968b:182-183; Barrera, 1968:309;Barr, 1971:113, 116; Laneyrie, 1971:192; Reddell,1971b:61; Vigna Taglianti, 1972:118, 124,125, 126, 127, fig. 5; Erwin et al., 1977:24; VignaTaglianti, 1977:326,333.Mexicaphaenops prietoi: Anonymous, 1942b:265(erroneous spelling).Mexaphaenops: Barr, 1966b:97; Vandel, 1964:237;Vandel, 1965a:199.Type-locality.-Grutas del Palmito, Bustamante,Nuevo Leon, Mexico.Distribution.-Known only from the type-locality.See Fig. 78.Discussion.-This is the rarest species of the genusand is known only from a few specimens. Barr(1967d) considers this to be the most highly apomorphicspecies in the genus.Paratrechus (Hygroduvalius) pallescens BarrTrechine (part): Barr, 1967a:66.Paratrechus (Hygroduvalius) pallescens Barr, 1967d:161-162, 165, fig. 1,4; Reddell, 1971b:61; VignaTaglianti, 1972:118; Reddell, 1973a:41; Mateu,1977:192, 194; Vigna Taglianti, 1977:326, 332Trechini: Reddell, 1967d:l06.Paratrechus new species: Barr, 1968b:183.Paratrechus (Hygroduvalius) pallescans: Reddell,1973a:35 (erroneous spelling).Paratrechus pallescens: Erwin et al., 1977:24.Paratrechus (Hygroduvalius) pallens: Mateu, 1977:200,206 (erroneous spelling).Type-locality.-Sotano de Tejamanil, 4.5 km westof Pinal de Amoles, Queretaro, Mexico.Distribution.-Known from four caves in Queretaro.See Fig. 78.New records.-Quenitaro: Sotano del Buque, SOtanode la Escuela, and Cueva del Judio (det. T. C.Barr, Jr.).Discussion.-This species is most closely related toParatrechus (Hygroduvalius) sylvaticus Bolivar, aneyed species from Morelos and the Distrito Federal.Paratrechus (H.) pallescens may represent a furtherlink in the series running from the robust, large-eyedspecies of Paratrechus (s. str.) to the elongate, anophthalmicspecies of Mexaphaenops (Barr, 1967d). Thisspecies has been taken from moist flowstone andfrom under rotten wood. An apparently troglobiticspecies of Paratrechus (s. str.) has recently been collectedfrom Cueva del Volcancillo, Veracruz, butremains undescribed. Two other species of Paratrechus(s. str.) have been taken from caves in Mexico(see Fig. 78): P. mexicanus Putzeys from Cueva de laCascada, Veracruz and P. tepoztlanensis Bolivar fromGrutas del Mogote, Guerrero, and Cueva de Coatepec,Mexico. Rossi and Cesari Rossi (1977) reported thatP. tepoztlanensis was parasitized by the ascomycetefungus Rhachomyces quetzalcoatl Balazuc in Cuevade Coatepec. They also report this same fungus as aparasite of an undetermined species ofParatrechus inResumidero Grande and Resumidero de Chemal,Huehuetenango, Guatemala. Mateu (1977) describedthe genus Mexitrechus to include P. coarctatus Bates.This species has been recently found in Sotano de lasBellotas and Comedor del Diablo, Oaxaca (see Fig.78).Platynus (Mexisphodrus) profundus (Barr)Mexisphodrus profundus Barr, 1966a:113-U4, US;Barr, 1967a:66; Reddell, 1967c:55; Barr, 1968b:184; Reddell, 1971b:61; Reddell and Mitchell,1971b:193, fig. 18; Vigna Taglinati, 1972:125;Hendrichs and Bolivar, 1973:38, 41; Mitchell andKawakatsu, 1973a:671; Reddell, 1973a:36, 41;Reddell and Elliott, 1973b:182; Vomero, 1974:353.Mexisphodrus sp.: Hendrichs and Bolivar, 1966:7(Tamaulipas record only).Mexisphodrus new species: Reddell, 1966a:13.Mexisphodrus: Sbordoni and Argano, 1972:8 [Cuevade la Perra (=Capilla) record only]; Peck, 1973c:129.Platynus profundus: Whitehead, 1973: 185, 207; Erwinet al., 1977:31.Type-locality.-Sotano de la Joya de Salas, 25 kmwest of Encino, Tamaulipas, Mexico.Distribution.-Known from caves in the Sierra deGuatemala, Tamaulipas. See Fig. 76.Records.-Tamaulipas: Sinkhole at Rancho delCielo, Cueva de la Capilla, Crystal Cave, HarrisonSinkhole, Sotano de la Joya de Salas, and Cueva de laMina.Discussion.~Five species have been described inMexisphodrus, all known only from caves. Of these,three are presumably troglobites and are includedhere. Of the remaining species, one (M. tlama.yaensisBarr from caves in Queretaro, San Luis Potosi, and223

Veracruz) belongs in another subgenus (Barr, pers.comm.), and the other (M. gertschi Hendrichs andBolivar) is a dark, eyed troglophile from Cueva deEI Ocote, Hidalgo. Barr (1965) originally consideredMexisphodrus to be a member of the tribe Sphodrini,but with recent collections and further study he feelsit to be more closely related to Platynus and its alliesin the Agonini (Barr, pers. comm.). Whitehead (1973)considers Mexisphodrus to be no more than a subgenusof Platynus. Platynus (M.) profundus is frequentlytaken from guano and from organically richareas. Undescribed species of the subgenus are knownfrom caves in Nuevo Leon, Puebla, Queretaro, SanLuis Potosi, and Tamaulipas.Platynus (Mexisphodrus) urquijoi (Hendrichs andBolivar)-NEW COMBINATIONMexisphodrus, undescribed species: Barr, 1967a:66(Oaxaca record only); Barr, 1968b:184.Mexisphodrus sp.: Reddell, 1971b:61 (Sotano de SanAgustin record only).Mexisphodrus urquijoi Hendrichs and Bolivar, 1973:38-41, fig. 1·2.Type-Iocality.-Sotano de San Agustin, cerca deHuautla de Jimenez, norte del estado de Oaxaca,Mexico.Distribution.-Known only from the type-locality.See Fig. 76.Discussion.-This is the most highly cave-adaptedspecies in the subgenus; it is the only Mexisphodruscompletely lacking eyes. Although known only fromthe type-locality, this species may be represented byspecimens from other caves in the Huautla region.This species is placed in the genus Platynus in accordancewith the evidence offered by Whitehead(1973).Platynus (Mexisphodrus) veraecrucis (Barr)Mexisphodrus veraecrucis Barr, 1965:66-71, fig. 1;Barr, 1966a:112, 113, 114,115; Barr, 1967a:66;Barr, 1968b:184; Reddell, 1971a:226; Reddell,1971b:61; Hendrichs and Bolivar, 1973:38, 41;Reddell, 1973a:36, 41.Mexisphodrus veracrucis: Hendrichs and Bolivar,1966:7,8,10 (erroneous spelling).Platynus (Mexisphodrus) veraecrucis: Whitehead,1973: 182, 214.Platynus veraecrucis: Erwin et al., 1977:31.Type-Iocality.-Sotano del Profesor, near Tequila,Veracruz, Mexico.Distribution.-Known from caves in the vicinitiesof Tequila and Soledad Atzompa, Veracruz. See Fig.76.Records.-Veracruz: Sotano del Profesor and Sotanode Sphodrini.New record.-Veracruz: Sotano Itamo (det. T. C.Barr, Jr.).Discussion.-This species, like P. (M.) profundus,has an elongate body, small eyes, and vestigial metathoracicwings. In Sotano del Profesor it was takenfrom under rocks near the body of a murderedschoolteacher who had been thrown into the pit. InSotano de Sphodrini specimens were taken from thecave walls.Speocolpodes franiai BarrSpeocolpodes franiai Barr, 1973:273,275-276, fig. 1;Mateu,1978:22.Eyeless carabid beetle: Syme and Peck, 1974:146.Platynus franiai: Erwin et al., 1977:30.Speocolpodes: Mateu, 1978:22,26,27.Type-Iocality.-Seamay Cave near Senahu, AltaVerapaz, Guatemala.Distribution.-Known only from the type-locality.See Fig. 76.Discussion.-This monotypic genus is most closelyrelated to Platynus. Erwin et al. (1977) place it inPlatynus, but without offering any evidence for thiscombination. With the exception of the Venezuelantroglobite Speleodesmoides raveloi Mateu, S. franiaiis the most southern troglobitic beetle in the NewWorld. The species is known only from two femalescollected on flowstone and from under a rock nearpools.Family DytiscidaeDarlington (1936) reported several species of dytiscidbeetle from open-air cenotes in Yucatan. Thermonectessp. was reported by Pearse (1938b) fromCenote de Sambula (Motul), Yucatan. It may be atroglophile. Two additional species of possible troglophileare known from Mexican caves: Agabus amencanusAube from Cueva de la Cascada, Veracruz, andHydroporus belfragei Sharp from Cueva de la Capilla,Tamaulipas. The latter species is abundant in siltflooreddrip pools.Suborder PolyphagaFamily AUeculidaeSeveral species of the family Alleculidae are probablytroglophiles in Mexican caves. Species of thegenera Hymenorus and Lystronychus have heen foundin caves in Coahuila, San Luis Potosi, and Tamaulipas.Lobopoda subcuneata Casey has been identifiedfrom Sotano del Pozo, San Luis Potosi.224

Family CantharidaeThe cantharids of caves are of uncertain ecologicalstatus, but the presence of larvae in many caves indicatesthat they may be troglophiles. Undeterminedlarvae have been found in caves in Hidalgo, Queretaro,Tamaulipas, and Veracruz, Mexico; and in Huehuetenango,Guatemala. Cantharis sp. was found inSumidero de Matehuala, San Luis Potosi, and Discodonsp. was taken in Cueva de la Puente, San LuisPotosI.Family DennestidaeDermestid beetles are frequently found in cavesinhabited by bats. Three species of this family havebeen identified from caves in Mexico. The most commonis Dermestes carnivorus Fabricius; it was collectedfrom caves in Coahuila, Durango, Guerrero,and San Luis Potosi. In the larger bat caves it may bepresent in vast numbers. The other two species foundin Mexican caves are Apsectus araneorum Beal fromCueva de la Boca, Nuevo Leon, and Dermestes maculatusDe Geer from Pozo Melendez, Guerrero.Family HisteridaeHister beetles are present in large numbers in manycaves throughout Mexico and Central America, butfew have yet been determined. Pearse (1938b) reportedthe presence of Gnathoncus sp., Oxarthriusn. sp. nr. hematicallis Sharp, Phelister sp., Platysomasp., Plegaderus sp., and Saprinus spp. 1 and 2 fromcaves in Yucatan. Recent collections have includedundetermined species of Epierus and Euspilotus fromcaves in San Luis Potosi and Tamaulipas, of Euspilotus(Hesperosaprinus) from Cueva de la Boca, NuevoLeon, and of Euspilotus (Neosaprinus) from caves inOaxaca and Yucatan. Paromalus luderti Marseul hasbeen identified from Cueva del Ojo de Agua de Tlilapan,Veracruz. Vomero (1977) described the troglophileAnapleus wenzeli from Cueva de la Canada n. 1,Chiapas (see Fig. 79). The four troglobitic speciesknown from the family in North America all belongto the distinctive genus Troglobacanius and are discussedbelow.Troglobacanius bolivari VomeroTroglobacanius bolivari Vomero, 1974:328, 340,341-345, 349, 350, 353-355, 356-358, fig. 6;Vomero, 1973:364 (nomen nudum); Decou andTherond, 1977:404.Type-Iocality.-Sotano del Tigre, 10 miles NE ofValles, San Luis Potosi, Mexico.Distribution.-Known only from the type-locality.See Fig. 79.Discussion.-Although ten blind species of histerbeetle are known, only Speleacritus anophthalmusJeannel and the four Mexican species included hereare troglobites (Vomero, 1973). The genus Troglobacaniusincludes only the troglobites listed here.It is most closely related to the genus Bacanius.Troglobacanius bolivari is closely allied to T. mayaand T. reddelli.Troglobacanius maya VomeroTroglobacanius maya Vomero, 1974:328, 329,331333,335-336,338,340,342,343,344,350,353355,356-358, fig. 1-3; Vomero, 1973:364 (nomennudum); Sbordoni, 1974:366; Sbordoni et al.,1974:15 (nomen nudum); Decou and Therond,1977:404.Type.locality.-Grutas del Cocona, Teapa, Tabasco,Mexico.Distribution.-Known only from the type-locality.See Fig. 79.Discussion.-This species is very closely allied to T.reddelli. It is the most highly evolved troglobite in thegenus and is from the most tropical region.Troglobacanius reddelli VomeroTroglobacanius reddelli Vomero, 1974:328, 336,338-341, 342, 343, 344, 350, 353-355, 356-358,fig. 4-5; Vomero, 1973:364 (nomen nudum); Decouand Therond, 1977:404.Type.locality.-Grutas de El Puente, 5 miles SE ofOcampo, Tamaulipas, Mexico.Distribution.-Known from two caves in theSierra de Guatemala, Tamaulipas. See Fig. 79.Records.-Tamaulipas: Grutas de El Puente andCueva de los Vampiros.Discussion.-This species is very closely allied toT. maya and somewhat more distantly to T. bolivari.Together these three species form the maya speciesgroup. It is probably significant that no species oftroglobitic histerid has been found at high elevationsin the Sierra de Guatemala. All of the species whichhave become cave adapted are known from lowlandtropical caves. Although some of the species whichhave been found in lowland caves of the Sierra deGuatemala are also known from higher elevationcaves and presumably evolved there and moveddownward through subterranean routes to the lowerelevations, it must be assumed that this is not thecase with Troglobacanius but that all of the speciesof this genus have evolved in the lowlands.225

Troglobacanius sbordonii VomeroTroglobacanius sbordonii Vomero, 1974:328, 345,347, 349-350, 353-355, 356-358, fig. 7-8; V0mero, 1973:364 (nomen nudum); Decou andTherond, 1977:404.Type-Iocality.-Sotano de Gomez Farias, Tamaulipas,Mexico.Distribution.-Known only from the type-locality.See Fig. 79.Discussion.-This species is unique III many respectsand has been placed by Vomero (1974) in themonotypic sbordonii species group.Family HydrophilidaeDarlington (1936) reported three species of hydrophilidbeetle from open-air cenotes in Yucatan. Tropisternusapicipalpis Chevrolat was recorded by Pearse(1938b) from Cueva Yunchen, Yucatan. The onlyother record of this family from the cave habitat inMexico is of T. (Cyphostethus) chalybeus Laportefrom Sotano del Arroyo, San Luis PotosI.Family LeiodidaeThe family Leiodidae is represented in Mexicancaves by many species, including two troglobites. Arecent collection of the leiodine Aglyptinus sp. fromHarrison Sinkhole, Tamaulipas, is the first record ofGUATEMALAr\.... r'...:-///Fig. 79.-Distrihution of troglobitic and troglophilic beetles of the family Histeridae: 1, Troglobacanius sbordonii; 2, T.reddelli; 3, T. bolivari; 4, T. maya; 5, Anapleus wenzeli.226

this genus in caves in mainland North America, althoughit is common in Antillean caves. Peck (1973a,1977a) has summarized the catopine beetle fauna ofthe caves of Mexico, Belize, and Guatemala. Threespecies of Dissochaetus have been identified fromcaves in southem and eastern Mexico and from Belize.Dissochaetus aztecus Szymcakowski has been reportedfrom caves in Nuevo Leon, San Luis Potosi,and Tamaulipas; D. curtus Portevin has been collectedin Cueva Cerro Hueco, Chiapas; D. hetschkoiReitter, a widespread species ranging north fromBrazil into Nuevo Leon, Mexico, has been reportedfrom caves in Campeche, Oaxaca, San Luis Potosi,Veracruz, and Yucatan, Mexico, and from Rio FrioCave B, Belize. A species with reduced eyes, Proptomaphaginusmicrops Peck, is known from caves inthe Xilitla and Aquismon regions of San Luis Potosi;Peck (1973a) considers this to be an endogean form.An undescribed species of Proptomaphaginus has recentlybeen collected in Cueva de la Finca, Oaxaca.See Fig. 80 for the distribution of cavemicole Dissochaetusand Proptomaphaginus in Mexico and Belize.MEXICOGUATEMALA/.//r\r"r- 0o~Fig. BO.-Distribution of troglophilic leiodid beetles of the genera Dissochaetus and Proptomaphaginus: 1, Dissochaetusaztecus; 2, D. aztecus and D. hetschkoi; 3, Proptomaphaginus microps; 4, P. microps and D. aztecus; 5, D. hetschkoi; 6, Proptomaphaginusn. sp.; 7, D. curtus.227

Ptomaphagus (Adelops) is a group of beetles representedin this region by eight troglophilic and threetroglobitic species. Peck (1973a, 1977a) has placedthe Mexican and Central American cavernicoles intotwo species groups. The consobrinus group rangesfrom the southern United States into Central America.Three cave inhabiting species from southernMexico, Belize, and Guatemala belong to this group:P. (A.) reddelli Peck from caves in the Valle Nacionaland Acatlan regions of Oaxaca; and P. (A.) barbaraePeck from caves in Belize and possibly in EI Peten,Guatemala, are troglophiles. Ptomaphagus (A.) giaquintoiJeannel from three caves in Alta Verapaz,Guatemala, is a probable troglobite. The remainingnine species all belong to the cavernicola group. Thisgroup ranges from the United States into Guatemala.Ptomaphagus (A.) cavernicola cavernicola Schwarz isa common troglophile in the United States; in Mexicoit is known only from Grutas del Palmito, NuevoLeon, where it is abundant on human feces throughoutthis heavily polluted cave. Ptomaphagus (A.)cavernicola aditus Peck is known only from Cueva dela Boca, Nuevo Leon; P. (A.) gypsum Peck, describedfrom Resumidero del Pablillo, Nuevo Leon, and originallythought to be a troglobite (Peck, 1973a), hasrecently been found in nearby epigean localities(peck, 1977a); P. (A.) elabra Peck is known frommany caves in the Sierra de EI Abra and lowlandSierra de Guatemala; P. (A.) leo Peck has been foundin caves in Nuevo Leon and Queretaro; P. (A.) spelaeus(Bilimek) is known from caves in Guerrero andMexico, as well as nearby epigean localities; P. (A.)tabascensis, described by Sbordoni (1974) fromGrutas del Cocona, Tabasco, has been collected alsoin caves in Campeche, Quintana Roo, and Yucatan,and from forested habitats in Campeche. The remainingtwo species of the cavernicola group, P. (A.)mckenziei and P. (A.) troglomexicanus, are troglobitesand discussed below. Sbordoni et al. (1977)reported undetermined troglophilic and troglobiticspecies ofPtomaphagus from several caves in Chiapas.See Fig. 81 for localities from which cavernicolePtomaphagus (Adelops) are known.Ptomaphagus (A delops) giaquintoi JeannelPtomaphagus (Adelops) giaquintoi Jeannel, 1936:93;Jeannel, 1949:98-99; Nicholas, 1962:180; Nicholas,1968:168.Ptomaphagus giaquintoi: Barr, 1973:276; Peck,1973a:l00, 101; Peck, 1973c:38, 40, 42, 43, 44,45, 61, 85, 86, 92, 93, 94, 95, 98, 99, 102, 103,104, 105, 111-112, 146, 147, 154, fig. 49, 93,123, 175,204;Strinati, 1977:387.Ptomaphagus: Peck and Peck, 1973:70.Type-Iocality.-Cueva Sepacuite (=Cueva Sepacuiten. 2), near Panzos, Alta Verapaz, Guatemala.Distribution.-Known from three caves in AltaVerapaz, Guatemala. See Fig. 81.Records.-GUATEMALA: Alta Verapaz: Grutasde Lanquin, Cueva de Seamay, and Cueva Sepacuiten.2.Discussion.-This species is the only troglobiticleiodid known from Guatemala. Its ecological statusis uncertain since it still retains functional flight wingsand pigmented eyes. The fact that the eyes are reduced,the body is depigmented, and the appendagesare elongated suggests that it is now restricted to thecave habitat (Peck, 1973a). The beetles have beenfound in association with the guano of insectivorousbats.Ptomaphagus (Adelops) mckenziei PeckPtomaphagus (Adelops) mckenziei Peck, 1977a:196,199,200,205,212, fi~48-59, 112.Type-Iocality.-Cueva de California, 4 mi. NERancho Nuevo, 46 road miles SW of EI Barretal (inTamps.), Nuevo Leon (=Tamaulipas), Mexico.Distribution.-Known only from two caves in thePurificacion region, Tamaulipas. See Fig. 81.Records.-Tamaulipas: Cueva del Brinco andCueva de California.Discussion.-This species possesses reduced and depigmentedeyes, elongated antennae, and reducedflight wings. It is very closely related to P. (A.) cavernicola,from which it may be directly descended(peck, 1977a). The cavernicola group includes, inaddition to the two troglobites listed here, a thirdform which shows some adaptations to the cave environment:P. (A.) cavernicola aditus from Cueva dela Boca, Nuevo Leon.Ptomaphagus (Adelops) troglomexicanus PeckPtomaphagus (Adelops) troglomexicanus Peck, 1968:92-97, fig. 1-5; Reddell, 1971b:63; Reddell andMitchell, 1971b:194, fig. 21; Reddell, 1973a:41;Reddell and Elliott, 1973b:182, 187; Peck, 1977a:193, 199,206, 207,212, fig. 19,58, 130.Ptomaphagus troglomexicanus: Peck, 1971b: 12;Sbordoni and Argano, 1972:8; Vigna Taglianti,1972:125; Mitchell and Kawakatsu, 1973a:671,673; Peck, 1973a:l03, fig. 1, 12; Peck, 1973c:38,40, 42, 44, 45, 60, Ill, 128, 129-130, 154, fig.60, 131, 190; Reddell, 1973a:36; Sbordoni andCobolli-Sbordoni, 1973a:340-346, fig. 1; Sbordoniand Cobolli-Sbordoni, 1973b:150; Sbordoni,228

1974:366; Vomero, 1974:353; Peck, 1977b:81,82.Type-Iocality.-Cueva de la Perra (=Capilla), LaPerra (=El Porvenir), 15 miles northwest of GomezFarias, Tamaulipas, Mexico.Distribution.-Known from four caves in the Sierrade Guatemala, Tamaulipas. See Fig. 8l.Records.-Tamaulipas: Cueva de la Capilla, Cuevade la Mina, Cueva de las Perlas, and Cueva Chica de laPerra.Discussion.-This species has greatly reduced eyesand very elongated antennal segments. It is a memberof the cavernicola group and is the most highly caveadaptedmember of the group. It was found on flowstoneand silt in association with Mexaphaenops intermediusin Cueva de la Capilla. Peck (1968) has discussedthe evolution of this species. Sbordoni andCobolli-Sbordoni (1973a) have considered the morphologicaldifferences between this "temperate" speciesand a typical "tropical" troglophilic species, P.GUATEMALA\r,...'...;/,./r'Fig. 81.-Distribution of troglobitic and troglophilic leiodid beetles of the genusPtomaphagus: 1,Ptomaphagw (Adelops)cavernicola cavernicola; 2, P. (A.) leo; 3, P. (A.) cavernicola aditus; 4, P. (A.) gypsum; S,P. (A.) mckenziei; 6,P. (A.) troglamexicanus;7, P. (A.) elabra; 8, P. (A.) spelaeus; 9, P. (A.) reddelli; 10, P. (A.) tabascemu; 11, P. (A.) barbarae; 12, P. (A.)giaquintoi.229

(A.) speLaeus, from Grutas de Cacahuamilpa, Guerrero.They concluded that these differences are a resultof the degree of energy input into the respectivecaves. Peck (1977a) has discussed alternatives to theirconclusions.Family LimnichidaeSpecimens of this family have been collected fromseveral caves in San Luis Potosi and Tabasco and apparentlybelong to at least two species of troglophile.They remain unstudied.Family PselaphidaeThe family Pselaphidae is among the principalgroups contributing to the troglobitic and troglophilicfauna of the United States, but it is rarely collectedin Mexican caves. A few species, including membersof the genera Cerocerus and Reichenbachia, havebeen collected, but all await study.Family PtilodactylidaeThe family Ptilodactylidae is represented in cavesin Mexico by at least one probable new species of thegenus Ptilodactyla. This troglophile has been found incaves in Guerrero, Nuevo Leon, Oaxaca, San Luis Potosi,Tamaulipas, and Veracruz.Family PtinidaeTwo species of spider beetle have been found inthe caves of Mexico, one of which is a possible troglobite(see Fig. 82). The other species, Niptus abstrususSpilman, is known only from caves in Texas (U.S.A.)and Coahuila and Durango, Mexico (Ashworth,1973). The eyes in this species are somewhat reducedbut less so than in N. absconditus. In addition to thetwo cavernicole species from Mexico and Texas, thegenus Niptus includes three species in orth America.One of these is a nearly cosmopolitan stored productpest, one is an inhabitant of mammal nests in thesouthwestern United States, and the third (N. abditusBrown) is a cavernicole known only from SpiderCave, Utah.Niptus absconditus SpilmanNiptus absconditus Spilman, 1968:194, 196, 197198, fig. 1-3; Reddell, 1971b:65; Reddell, 1973a:36,41.Type-Iocality.-Grutas de Xoxafi, Hidalgo, MeXICO.Distribution.-Known only from the type-locality.See Fig. 82.Discussion.-This species possesses the smallesteyes in the genus and is known only from the mostremote sections of Grutas de Xoxafi. Whether it is atroglobite or not is speculative, but it is included herebecause the species is doubtless in the process of becomingadapted to the cave environment.Family ScarabaeidaeSeveral species of scarab beetle have been collectedfrom caves, but most are probably accidentals. Thegenus Onthophagus, however, is a frequent contributorto the cave fauna of Mexico. These beetles areusually found in bat guano and are probably troglophiles.Onthophagus incensus Say was found in Sotanode Tlamaya, San Luis Potosi; O. cuevensis wasrecently described from caves in San Luis Potosi andTamaulipas (Howden, 1973); O. Landolti Harold hasbeen found in Cueva del Ojo de Agua Grande, Veracruz;and O. vespertilio was described by Howden,Cartwright, and Halffter (1956) from Gru ta de Acuitlapan,Guerrero, and has also been found in Gru tasdel Mogote, Guerrero. See Fig. 82 for the distributionof Onthophagus in Mexican caves.Family ScydmaenidaeThe family Scydmaenidae is well represented inthe caves of Mexico and Guatemala where it apparentlyreplaces the Pselaphidae as a major element ofthe small beetle fauna. Franz (1977) described twocavernicole species of scydmaenid from Mexicancaves: Scydmaenus teapanus from Grutas del Cocona,Tabasco; and Euconnus (Madagassoconnus) arganoifrom Cueva del Muju, Chiapas (see Fig. 83). Bothappear to be troglophiles. Species of the genus Scydmaenus(s. str.) have been collected in caves in Izabaland Alta Verapaz, Guatemala, and in Veracruz andOaxaca, Mexico. Euconnus (? Drastophus) n. sp. hasbeen collected from bat guano in Cueva del Ojo deAgua de Tlilapan, Veracruz. Several species of Euconnus(Napochus) have been collected in caves inAlta Verapaz, Guatemala, and in Campeche, Oaxaca,Tamaulipas, San Luis Potosi, and Veracruz, Mexico.Their ecological status is uncertain, but at least somespecies are presumed troglophiles. An undescribedspecies of Euconnus, possibly belonging to a newsubgenus, has been collected in Sotano del Leon,Tamaulipas. It is a highly attenuate form and maybe a troglobite.Famity StaphylinidaeThe family Staphylinidae has made a major contributionto the Mexican cave fauna, and at least 34species have been found. Many of these are known230

only by generic identifications. The subfamily Neocharinaeis frequently collected, but the taxonomyof this group is too poorly understood to allow evengeneric identifications. The genera Anotylus, Belonuchus,Carpelimus, Erchomus, Homaeotarsus, Medon,Philonthus, Scopaeus, Stamnoderus, and Stenusare frequently collected in almost every part of Mexicoand Central America; species identifications inthese genera must await revisionary studies. Stilicolinacondei Jarrige is a species described from a cavein Central Texas (U.S.A.) and reported by Herman(1970) from many caves in Texas and northernMexico. Recent study, however, indicates that theMexican populations belong to an undescribed spe-cies (H. Frania, pers. comm.). This species rangessouth from near the Texas border into the Aquismonregion of San Luis Potosi (See Fig. 83). Stenopholeareddelli Herman was originally described as atroglobite from Cueva de la Mina, Tamaulipas (Herman,1969), but it has recently been found to beabundant in the endogean fauna of the high mountainregions of the Sierra Madre Oriental (Herman,pers. comm.) (see Fig. 83).Family TenebrionidaeThe family Tenebrionidae is widely distributed incaves in the western United States and Mexico and is\(~GUATEMALA/.//rr ,,.. ..--Fig. 82.-Distribution of troglobitic and troglophilic beetles of the families Ptinidae and Scarabaeidae: 1, Niptus absconditus;2, Onthophagus cuevensis; 3, O. incensus; 4, N. abstrusus; 5, O. vespertilio; 6, O. landolti.231

epresented by many species. Tenebrionids are usuallyfound in the drier parts of caves, frequently asguanophiles. Many species have been identified andsome await description. Large collections remainundetermined, and the number of identified tenebrionidspecies will doubtless increase with furtherstudy. Alphitobius laevigatus (Fabricius) is a commonguanophile in caves in Coahuila, San Luis Potosi,and Tamaulipas. Cryptoglossa mexicana mexicanaChampion is abundant in the caves of northernand western Mexico. It is known from dry, dustycaves in Chihuahua, Coahuila, Durango, and NuevoLeon. The genus Eleodes is the most commonly collectedgenus of tenebrionid in Mexico. Althoughusually found as guanophiles, they are also takenin dry dusty areas of many caves not inhabited bybats. Most collections of this genus await study, buta few species have been identified. Eleodes glabricollisChampion has been found in gypsum cavessouth of Galeana, Nuevo Leon. Eleodes rugosa Perboseis frequently found in the drier area of caves inthe Sierra de EI Ahra, San Luis Potosi and Tamaulipas.Eleodes sallei Champion has been found in manycaves at higher elevations in the Sierra Madre Orientalin San Luis Potosi and Queretaro and in Oaxaca. Eleodeshispilabris (Say), a common guanophile in Texascaves, has been collected from Cueva de los Lagos,Coahuila. Eleodes spinolai Solier has been collected inGUATEMALAFig. S3.-Distribution of troglophilic beetles of the families Staphylinidae and Scydmaenidae: 1, Stilieolina n. sp.; 2, Stilieolinan. sp. and Stenopholea reddelli; 3, Seydmaenul teapanw; 4, Eueonnul (MadagtUloeonnul) arganoi.232

Grutas de Cacahuamilpa and Grutas del Mogote, Guerrero.Liodema sp. nr. kirschi Bates is a frequentlycollected species in the caves of the Sierra de El Abra,San Luis Potosi and Tamaulipas. Rhinandrus ?elongatusHom was reported by Pearse (1938b) from severalcaves in Yucatan. Zophobas atratus (Fabricius),known from caves in the Sierra de El Abra, San LuisPotosi, is otherwise known only from a cave in Venezue1a.For the cave localities of several species oftenebrionid in Mexico see Fig. 84.Family TrogidaeThis family of beetles is frequently collected asguanophiles m Central Texas. The only record forMexican caves is of Omorgus carinatus (Loomis) fromCueva de la Siquita, Durango. It is probably a guanophilein this large bat cave.Order SiphonapteraFamily HystrichopsyllidaeThe only species of this family reported fromMexican caves is the bat flea Anomiopsyllus traubiBarrera from Cueva de Belen, Hidalgo (Barrera,1951).Family IschnopsyllidaeThree species of bat flea of the family Ischnopsyllidaehave been taken from guano in Mexican caves.GUATEMALA/ ..//r,.... r"'.-"Fig. 84.-Distribution of cavemicole beetles of the family Tenebrionidae: 1, Eleodes hupilabris; 2, Cryptoglossa mexicanamexicana; 3, E. glabricollis; 4, E. rugosa; 5, E. sallei; 6, E. spinolai.233

Myodopsylla gentilis Jordan and Rothschild wasfound in Cueva del Cochino, San Luis Potosi. A singlespecimen of M. globata Holland was collected fromGrutas del Cocom!, Tabasco; this is only the secondspecimen of this species, previously known from SanCristobal de las Casas, Chiapas. Sternopsyllus distinctatexana (C. Fox) was extremely abundant inguano in Grutas de San Bartolo and Cueva de la Boca,Nuevo Leon.Family PulicidaeThe family Pulicidae is represented in Mexicancaves by three species. Pulex irritans Linnaeus isknown from Cenote Sabacah (Sucopo), Yucatan, andCueva del Cochino, San Luis Potosi; it is usually associatedwith man and large carnivores. The specimensfrom Cueva del Cochino were taken from thedry entrance area of the cave. Pulex porcinus Jordanand Rothschild is known from Grutas de MonteBravo and Grutas de Xtacumbilxunam, Campeche,and from Cenote Amil, Yucatan. This species isusually associated with peccaries; its host in theabove caves is unknown. Pearse (1938b) reportedthe plague £lea, Xenopsylla cheopis (Rothschild) fromGrutas de Balankanche, Yucatan.Order DipteraAlmost 100 species of £ly have been collected inMexican caves (see Table 27). In addition, numerouscollections await study. Many £lies utilize theentrance areas as daytime retreats; others are attractedby water or carrion. Some species apparently aretroglophiles and complete their life cycle underground.No species from Mexico or Central Americais troglobitic. The following is a brief summary ofsome of the more significant species with respect tothe cave habitat.Suborder NematoceraFamily CeratopogonidaeTwo genera of this family have been identifiedfrom caves. The presence of larvae in caves indicatesthat they may possibly be troglophiles. Pearse(1938b) reported Dasyhelea sp. from Actun Gongora,Yucatan. Forcipomyia sp. has been reported fromCueva Chac Mol, Yucatan (Pearse, 1938b); it has alsobeen collected in Actun Xpukil, Yucatan.Family ChironomidaeSeveral species of midge of the family Chironomidaehave been collected from caves and cenotes inYucatan (pearse, 1936b, 1938b). The larvae of thisfamily are usually aquatic, and presumably, theTable 27.-Summary of cave inhabiting Diptera.NematoceraCecidomyiidaeCeratopogonidaeChironomidaeCulicidaeMycetophilidaePsychodidaeSciaridaeTipulidaeBrachyceraAsilidaeDolichopodidaeEmpidiidaeRhagionidaeScenopinidaeStratiomyidaeTabanidaeTherevidaeCyclorrhaphaAgriomyzidaeCalliphoridaeChloropidaeDrosophilidaeLonchaeidaeMetopiidaeMilichiidaeMuscidaePhoridaeSciomyzidaeSphaeroceridaeStreblidaeTachinidaeTotalTroglobitesooooooooooooooooooooooooooooooOther Speciesadults' presence in this habitat is a result of theabsence of surface water in most of northern Yucatan.Tendipes fulvipilus Rempel is an apparent troglophilein Cueva del Azufre, Tabasco (Gordon andRosen, 1962). It has been taken from guano depositsin large numbers.Family Culicidae225831625Eight species of mosquito of the family Culicidaehave been collected from caves and cenotes in Yucatan(Pearse, 1938b). Mosquitoes frequently utilizethe entrance of caves for shelter, but in Yucatanwhere surface water is almost nonexistent, manyspecies are dependent upon caves and cenotes asplaces in which to deposit their eggs. Pearse (1938b)reported Aedes angustivittatus Dyar and Knab, A.euplocamus Dyar, A. taeniorhynchus (Wiedemann),Culex sp., Haemagogus sp., Isostomyia sp., Mochlonyxsp., and Psorophora sp. from caves in northernYucatan.I2III2I2I221115441217297234

Family MycetophilidaeMycetophilid flies are occasionally found in thecavemicole habitat, and three genera are known fromMexican caves: Keraplatus sp. from Sotano Encantado,Queretaro; Mycetophila sp. from Cueva de lasPerlas, Tamaulipas; and Rhymosia sp. from caves inthe Valle de los Fantasmas region, San Luis Potosi.Family PsychodidaeThree genera of psychodid flies have been found inMexican caves. Nem.opalpus sp. was reported fromCueva Segunda del Camino a San Roque, Yucatan(Pearse, 1938b). Breder (1942) reported Psychodasp. from Cueva Chica, San Luis Potosi. Telmatoscopusalbipunctatus (Williston) was reported to be veryabundant in Grutas del Cocona, Tabasco (Ryckman,1956). The last species, at least, is a troglophile.Williams (1976a, 1976b, 1976c) has studied indetail the biology of the phlebotomine sandflies inhabitingcaves in Belize. During the course of hisstudies he collected 15,041 specimens from fourcaves in the Cayo District (Millionario Cave, AugustineCave 1, Augustine Cave 2, and San AntonioCave). Thirteen species were identified, two of whichbelonged to the genus Brumptomyia and 11 to thegenus Lutzomyia. Some of the species were quiterare and may have followed the collectors into thecaves. A few, however, apparently complete theirlife cycle in the caves, utilizing bats as their sourceof blood meals. The most common species in all ofthe caves was L. beltrani (Vargas and Diaz Najera).The second most abundant species in all but SanAntonio Cave was L. deleoni (Fairchild and Hertig).Lutzomyia trinidadensis (Newstead) was the secondmost abundant species in the San Antonio Cave,but was comparatively rare in the other caves. Theonly other species present in all of the caves was L.shannoni (Dyar). The remaining species were all veryrare and probably not of significance to a study ofthe cave fauna. Williams (1976a) reported the presenceof trypanosomatid flagellates and filaria wormsin L. trinidadensis from the Belize caves. The vertebratehost of the trypanosomes infecting this speciesis not known. An undetermined trypanosome speciesalso infected L. beltrani; this species is probably a batparasite.Family SciaridaeThe small gnats of the family Sciaridae are frequentlyabundant in Mexican caves. Sciara sp. hasbeen found in caves in Yucatan (Pearse, 1938b) andTamaulipas. Bradysia coprophila (Lintner) is knownfrom Sotano de Caballo Moro, Tamaulipas. Undeterminedspecies of Bradysia are known from caves inCampeche, Durango, Nuevo Leon, Oaxaca, San LuisPotosi, and Tamaulipas. These gnats are apparentlytroglophiles and may be so abundant as to be extremelyannoying.Family TipulidaeCrane flies are common inhabitants of caves,where they are frequently found resting on cave wallsin the vicinity of the entrance. No species have yetbeen identified, but the following genera are knownto occur in Mexican caves: Epiphragma, Helius, Limnophila,Limonia, and Teucholabis.Suborder BrachyceraFamily DolichopodidaeTwo species of this family have been found inMexican caves: Chrysotus sp. from Salamander Cave,Tamaulipas, and Peloreopeodes cornutus Van Durzeefrom Cueva de la Puente, San Luis Potosi. Some speciesof dolichopodid fly are known to be troglophiles,but the ecologic status of the above species is unknown.Family TherevidaeTwo species of the family Therevidae have beenfound in Mexican caves. Henicomyia hubbardi Coquillettis known from Cueva de los Pajaws, Tamaulipas,and Psilocephala sp. is abundant in the caves ofYucatan (pearse, 1938b).Suborder CyclorrhaphaFamily DrosophilidaeThe family Drosophilidae is frequently collected incaves in Mexico. Drosophila repleta Wollaston hasbeen identified from Cueva Xconsacab, Yucatan(Pearse, 1938b). Undetermined species of Drosophilahave been taken from caves in Nuevo Leon, Puebla,San Luis Potosi, Tamaulipas, and Yucatan. In Cuevade las Perlas, Tamaulipas, enormous swarms were observedon the ceiling of the cave. Thousands of individualswere collected in dung-baited traps in Cuevade Chorros de Agua, Nuevo Leon.Family MilichiidaeThe small flies of the family Milichiidae are frequentlyfound in caves. Milichia sp. is abundant inthe caves of Yucatan (Pearse, 1938b). Pearse (1938b)also reported the presence of Desmometopa sp fromtwo caves in Yucatan. Undetermined species of the235

genus Pholeomyia have been collected from caves inChiapas, Puebla, Tamaulipas, and Veracruz. Threespecies of Pholeomyia have been reported fromMexican and Guatemalan caves: P. dampfi Sabroskyfrom Grutas del Cocona, Tabasco, Mexico, and Cuevade Jobitzinaj, El Peten, Guatemala (Sabrosky, 1959);P. indecora Loew from Cueva Chica, San Luis Potosi(Breder, 1942); and P. leucozona Bilimek from Grutasde Cacahuamilpa, Guerrero (Bilimek, 1867), andCueva Chirripeck, Alta Verapaz, Guatemala (Papp,1978). These species are all presumed troglophiles.Family PhoridaeThe humpbacked flies of the family Phoridae areoccasionally collected in Mexican caves. Megaseliascalaris Loew has been found in several caves in Yucatan(Pearse, 1938b). Conicera dauci Meigen wasfound in Sumidero del Jineo, Tamaulipas. Undeterminedspecies of the genera Conicera, Dohrniphora,and Puliciphora have been taken from caves throughoutMexico.Family SphaeroceridaeThe small dung flies of the family Sphaeroceridaeare frequently taken from bat guano in caves. Archiborborusmexicanus Steyskal was described from SOtanode El Porvenir, Tamaulipas (Steyskal, 1973).This is the most northern record for the genus. Undeterminedspecies of Leptocera have been collectedfrom caves in Hidalgo, Nuevo Leon, San Luis Potosi,Tamaulipas, and Yucatan.Family StreblidaeSeventeen species of streblid fly have been identifiedfrom caves in Mexico. These flies are all parasitesof bats and are frequently abundant in caves, wherethey may be found on cave walls or on guano. Theirbite is annoying, but not usually painful. This familyin Mexico has been reviewed by Hoffmann (1953).Among the more frequently collected species are Euctenodesmirabilis Waterhouse from caves in Guerrero,San Luis Potosi, and Yucatan; Megistopoda araneae(Coquillett) from caves in San Luis Potosi and Yucatan;Nycterophilia coxata Ferris from caves throughoutMexico; Trichobius adamsi Augustson from cavesin Baja California Sur, Colima, and Guerrero; T. caecusEdwards from caves in Guerrero, San Luis Potosi,Tamaulipas, and Yucatan; T. intermedius Petersonand Hurka from caves in Yucatan (Peterson andHurka, 1974); T. parasiticus Gervais from caves inChiapas, Guerrero, San Luis Potosi, Tabasco, andYucatan; and T. sphaeronotus Jobling from caves inBaja California Sur, Chiapas, Nuevo Leon, and Sonora.Table 28.-Summary of cave inhabiting Lepidoptera.TroglobitesOther SpeciesAcrolophidae 0 1Arctiidae 0 1Blastobasidae 0 1Gelechiidae 0 1Noctuidae 0 1Phalaenidae 0 1Pterophoridae 0 1Pyralidae 0 2Tineidae 0 6Total 0 15Order TrichopteraTrichoptera larvae are occasionally taken fromstreams in caves, and a few species may completetheir life cycle underground.Family CalamoceratidaeSpecimens of the genus Phylloicus have been takenIn Cueva de la Puente and Sotano de Tlamaya, SanLuis Potosi. These are probably troglophiles.Family PhilopotamidaeApparently troglophilic species of the genus Wormaldiahave been collected from the streams in Cuevade la Puente, San Luis Potosi, and Cueva del Brinco,Tamaulipas.Family PolycentropidaePolycentropus picana Ross was found III thestream passage in Cueva de la Puente, San Luis Potosi.An undetermined species of Polycentropus is anapparent troglophile in Cueva del Brinco, Tamaulipas.Order LepidopteraAlthough nine families of moth have been collectedin Mexican caves, only two are of unusualsignificance to the study of the cavernicole fauna(see Table 28). Most collections of moths have beenin alcohol and, therefore, are unsuitable for descriptionor specific determination.Family NoctuidaeNumerous specimens of noctuid moth have beencollected from Mexican caves, most from the entrancearea. Latebraria amphipyroides Guenee is frequentlyfound in caves throughout much of Mexico.This large moth is to be seen resting during the daytimeon the cave wall. It was reported from Yucatancaves by Pearse (1938b) and from caves in the Xilitlaregion of San Luis Potosi by Bonet (1953a).236

Table 29.-Summary of cave inhabiting Hymenoptera.TroglobitesOther SpeciesApidae 0 4Bethylidae 0 3Braconidae 0 1Chalcididae 0 2Cynipidae 0 1Diapriidae 0 3Figitidae 0 1Formicidae 0 52Halictidae 0 2Ichneumonidae 0 3Mutilidae 0 1Pompilidae 0 2Scelionidae 0 1Sphecidae 0 4Vespidae 0 1Total 0 81Family TineidaeSeveral, probably undescribed, species of the familyTineidae are known from caves in Mexico. Aprobable new genus and species has recently beencollected from bat guano in Cueva de Tasalolpan,Puebla. Amydria spp. have been collected from cavesin Nuevo Leon, Puebla, Tamaulipas, and Veracruz.Decardarchis sp. is known from Cenote de Sambuhi(Motul), Yucatan, and Grutas de Xtacumbilxunam,Campeche. Episcardia sp. has been taken from cavesin Nuevo Leon and Tamaulipas. Tinea sp. is knownfrom caves in San Luis Potosi and Yucatan. Monopisimpresipenella (Bilimek) is a poorly known speciesrecorded from Grutas de Cacahuamilpa, Guerrero(Bilimek, 1867).Order HymenopteraThe order Hymenoptera is well represented incaves, and 81 species have been identified from thishabitat in Mexico (see Table 29). Many species areaccidentals, but others, such as bees and wasps, buildtheir hives and nests within the shelter of the caveentrance. Some of the species are very aggressive andmay inflict severe stings to the cave explorer. Beesalso are found occasionally around moist flowstonein the entrance rooms of caves.Family ApidaeFour species of the family Apidae have been collectedfrom caves in Mexico. Apis melli/era Linnaeusis known from the entrance sinks of caves in Campecheand Yucatan. This species usually builds itshives in crevices on the vertical walls of the sinkholes.Local inhabitants frequently build elaborate scaffoldingsup the cave walls to reach the hives. Three additionalspecies of apid bees have been collected fromthe entrance area of caves in Mexico: Partamona cupiraorizabensis (Str.) from Cueva de los Pajaros,Tamaulipas; Trigona atrolutea Moure from ActUnXpukil, Yucatan; and T. testacea cupira Sm. fromCenote Sucila, Yucatan. The latter two species aremuch more common in caves in Yucatan than thesetwo records indicate.Family BethylidaeThree genera of wasp of the family Bethylidaehave been found in Yucatan caves. Pearse (1938b)reported Apanesia n. sp. from ActUn Kaua, Yucatan.Holepyris sp. has been found in Actun Xkyc,Yucatan, and Rhabdepyris sp. has been collectedin Actun Kaua.Family FormicidaeThe only group of hymenopterans which may beconsidered to be truly associated with the cave habitatis the family Formicidae. Ants are not uncommonin cave entrances and occasionally are found throughoutthe cave. Most of these species are accidentals ortrogloxenes, but a few are apparently permanent inhabitantsof caves.Among species commonly taken in caves but obviouslynot in any way adapted for a cave existenceis Pachychondyla harpax montezumia F. Smith. Thisspecies has been found in several caves in San LuisPotosi and Yucatan. Another species commonlytaken in caves is Aeromyrmex octospinosus (Reich),which was reported from caves in Yucatan (Wheeler,1937). This species is frequently found building itsnest in the twilight zone of caves. Solenopsis geminata(Fabricius) has been found in caves in Oaxaca,Puebla, San Luis Potosi, Tabasco, Veracruz, Campeche,and Yucatan. In some caves this species hasbeen present in thousands, both in twilight and intotal darkness. In Sotano de Guadalupe, San LuisPotosi, a large nest at the entrance allowed immediateaccess by the ants into the cave, and individualswere observed carrying out numerous animals,including some troglobites. Other ants collectedfrequently in Mexican caves are: Hypoponera punctatissima(Roger) from Campeche, San Luis Potosi,and Yucatan; Labidus coecus (Latreille) from Oaxacaand Yucatan; L. praedator (F. Smith) fromSan Luis Potosi and Yucatan; Leptogenys spp.from Oaxaca, Campeche, Yucatan, and QuintanaRoo; Pachycondyla apicalis (Latreille) from Campeehe,Quintana Roo, and Yucatan; and P. villosa(F. Smith) from caves in Quintana Roo, Tamaulipas,and Yucatan.237

Wheeler (1938) has reported the presence in cavesin Yucatan of three species that at various times havebeen reported as troglobites. Erebomyrma urichi(Wheeler) was originally described from a cave onTrinidad and is known also from Cenote de Sambula(Motul), Yucatan. It is a pale species now known alsofrom endogean localities. Two other species areknown only from caves in the Yucatan Peninsula:Brachymyrmex cavernicola Wheeler from Grutas deBalankanche, Yucatan; and Paratrechina pearsei(Wheeler) from caves in Campeche, Quintana Roo,and Yucatan. Both of these species are pale yellowwith minute eyes. The latter species has been foundto occur only in darkness and is frequently takenfrom moist flowstone in the most remote parts ofthe caves. Wilson (1962) has discussed the ecologicstatus of these species and has convincingly arguedthe case against the existence of troglobitic ants.Family PompilidaeTwo genera of the spider wasps of the familyPompilidae have been collected from caves in Yucatan.Auplopus sp. was collected from Actun Chukum.Pepsis sp. was taken from total darkness in Cueva deAguacate while it was in battle with a theraphosidtarantula. The same genus was found in total darknessin ActUn Kaua, where the wasps flew at thecollectors, apparently attracted by their lights.Family VespidaeThe vespid wasp Polybia diguetana Buysson wascollected from nests in the entrance areas of Cenotede Ek Bis, Campeche, and Actun Kaua, Yucatan.Phylum ChordataClass TeleosteiThe blind fish of the caves of Mexico have excitedgreater interest than any other group of animals to befound in the caves of North America. The genusAstyanax is certainly the best studied of all cavernicoles.In addition to the five species of troglobiticfish discussed in detail below, 16 other species offish have been recorded from Mexican caves (seeTable 31). Some of these are troglophiles, whileothers are probably accidentals or trogloxenes.Order CypriniforrnesFamily CharacidaeAstyanax jordani (Hubbs and Innes)Anoptichthys jordani Hubbs and Innes, 1936:5-7,pI. 1; Hykes, 1937:108-109; Innes, 1937:200-202; Jordan, 1937:203-204; Greenberg, 1938:80;Hubbs, 1938:261, 264, 271, 275; Bridges, 1940:74-97, map; De Buen, 1940:8; Gresser and Breder,1940:113-116, pI. 1-3; Breder and Gresser, 1941b:123-130, fig. 1-2, pI. 1-4; Anonymous, 1942a:221;Breder, 1942:7, 11-14, fig. 2, 5; Osorio Tafall,1942:206;Walls, 1942:210, 387; Breder, 1943a:2630; Breder, 1943b:82; Breder, 1943c:168-176;Breder and Rasquin, 1943:169-200, fig. 1-9, pI.1-3; Bridges, 1943:82-90; Hubbs, 1943: 121; OsorioTafall, 1943:44, 55, 57-67, 68, 69; Anonymous,1945:70; Barbour, 1945a:77;Barbour, 1945b:189;Benn, 1945:12; Breder, 1945:95·100, fig. 1-2, pI.1; Alvarez,l946:263, 264, 265, 267, 268, 270, 271,272,273,274-275,276,280,281, fig. 2-8; De Buen,1946:127; Jordan, 1946:198; Pavan, 1946:358;Rasquin, 1946:578-579; Alvarez, 1947:219; Brederand Rasquin, 1947a:323-351, fig. 1-9; Brederand Rasquin, 1947b:29-33; Nigrelli, 1947:105,108, pI. \11(fig. 38); Rasquin, 1945:35-42, fig. 1,pI. 1; Schlagel and Breder, 1947:17-27, fig. 2;Allee et al., 1949:612, 674, 675, fig. 247B-C;Rasquin, 1949a:499-531, fig. 4-5, 7, pI. 20-21,23; Rasquin, 1949b:131, 133; Alvarez, 1950:Table 30.-Summary of cave inhabiting Chordata.TroglobitesOther Species*Teleostei 5 16AmphibiaUrodelaAmbystomatidae 0 1Plethodontidae 0 9AnuraBufonidae 0 4Hylidae 0 6Leptodactylidae 0 17Ranidae 0 1Rhynophrynidae 0 1ReptiliaCheloniaKinostemidae 0 3SquamataBoidae 0 1Colubridae 0 5Crotalidae 0 2Gekkonidae 0 219uanidae 0 5Teiidae 0 1Xanthusiidae 0 3AvesFalconifonnesCathartidae 0 1PsittacifonnesPsittacidae 0 2PodicipitifonnesPodicipitidae 0 1238

Table 30.-(Continued)Troglobites Other SpeciesStrigiformesStrigidae 0 1Tytonidae 0 1ApodiformesApodidae 0 2Trochilidae 0 2CoraciifonnesMomitidae 0 2PasseriformesHirundinidae 0 2Troglodytidae 0 1Turdidae 0 2MammaliaMarsupialiaDidelphidae 0 2ChiropteraEmballonuridae 0 5Molossidae 0 7Mormoopidae 0 6Natalidae 0 2Phyllostomatidae 0 33Vespertilionidae 0 19RodentiaCricetidae 0 8Dasyproctidae 0 1Erithizontidae 0 1CarnivoraCanidae 0 1Felidae 0 1Mustelidae 0 1Procyonidae 0 1Total 5 182*For summary by family see Table 31.27-28; Atz, 1950:87, 89-90; Heuts, 1951:223;Ladiges, 1951a:258-260; Ladiges, 1951b:169;Rasquin and Hafter, 1951a:398; Rasquin andHafter, 1951b:163; Meder, 1952:171-174; Bonet,1953b:241; Breder, 1953:180; Luling, 1953a:314-318; Luling, 1953b:62-65; Luling, 1953c:289-299; Rioja, 1953e:l0; Angel, 1954:85-86,foIl. 96; Breder, 1954:13-16, pI. 1; Kauffeld,1954:140-141; Kuhn and Kiihling, 1954:385-398,fig. 1-2; Luling, 1954a:197-203; Luling, 1954b:942; Olivereau and Herlant, 1954:52; Rasquin andRosenbloom, 1954:406; Stefanelli, 1954b:277282, fig. 1-2; Thines, 1954:35-58; Axelrod andVorderwinkler, 1955:61; Bridges, 1955:256-268;Luling, 1955a:138-143, fig. 1-6; Luling, 1955b:401-477; Luling, 1955c:138-143; Quaghebeur,1955:?, Thines, 1955:9,41,68-70, 108-109,115,123; Olivereau and Francotte-Henry, 1955-1956:143; Bonet, 1956b:96; Bonet, 1956c:238; Dear01£, 1956:202, 210; Pfisterer, 1956:11-12; Sadoglu,1956:113-114; Thines, 1956:79; Gordon,1957:466; Hahn, 1957:1-81; John, 1957:129,130; Kiihling, 1957:?; Luling, 1957:145-150;Table 31.-Summary of cave inhabiting Teleostei.TroglobitesOther SpeciesCypriniformesCharacidae 1 1Ictaluridae 1 0Pimelodidae 1 4CyprinodontiformesPoeciliidae 1 4PerciformesBrotulidae 1 0Cichlidae 0 6SynbranchiformesSynbranchidae 1 0Total 5 16Pickford and Atz, 1957:77, 159, 324-325, 327;Quaghebeur, 1957:?; Thines and Kiihling, 1957:150-160; Bertin, 1958b:2661-2662; Cahn, 1958:76-107, pI. 14-15, 22-24, 26-27; Grobbel andHahn, 1958:249-266; Marshall and Thines, 1958:441, 446, 447, 449, 450, 453; Pelz, 1958:172175; Stolk, 1958:382-394; Alvarez, 1959:79;Breder, 1959:423, 424, 425, 426, 436, 437,446,447, fig. 4-6; Frank, 1960:39-41; Hahn, 1960:611-612; Humbach, 1960a:1-88; Humbach,1960b:551-552; Kosswig, 1960a:24; Kuhn, 1960:481-483; Strotkoetter, 1960:611-612; Thines,1960:44, 45, 46; Frank, 1961:347-365; Heuts,1961: 196; Kiihling, 1961:439-451; Poulson, 1961:143, 171-174; Axelrod et aI., 1962:C-35.00,F-38.00, F-141.00; Bath, 1962:242,281-285,287,fig. 35-36; Franck, 1962:?; Franck-Krahe, 1962:?;Goettert, 1962:56-58; Goodrick, 1962:10; Gordonand Rosen, 1962:365-366; Luling, 1962:139-145; Nicholas, 1962:165, 184; Tamayo,1962:vol. 1II:260; Bonet, 1963b:95; Burgers et aI.,1963: 189-195; Durand, 1963: 183; Pfeiffer, 1963:112, 116, 117, 123, 125, 128, 131, 132, 139, 141;Poulson, 1963:286, 287, 288; Sterba, 1963:144,145, 146-147, fig. 173; Steven, 1963:219, 221,230; Berra, 1964:12-19; Franck, 1964:96; Kosswig,1964:73-74; Moore and Nicholas, 1965:9495; Vandel, 1964:52,262-264,361,404,405,414,458-459,462,463,466,472,473,476,483,509,570,571, fig. 59; Boucquey et aI., 1965:88,91,92; Brittan and Bohlke, 1965:2; Glaser, 1965:?;Kosswig, 1965:246-250, fig. 9; McKenzie, 1965a:38, 39; Post, 1965:56, 57, 64, 89; Thines et aI.,1965:65, 77, 92, 101, 102; Vandel, 1965a:39,222,223,344,351,387,392,395,401,403,404,409, 429, 483, fig. 59; Breder and Rosen, 1966:155-156; Innes, 1966:161; Peters and Peters, 1966:393-414; Pfeiffer, 1966:97-105; Reed, 1966:7980, 83, 85; Barr, 1967c:486; Greenwood, 1967:239

263.264, 265, 267, 270, 273, fig. 58, pI. 40A;Johnson, 1967:6-7, 9-11, 25, 27,28,30,32,34,36,37,38, 39,40,44,46,48,49, fig.7,9;Kos~wig, 1967:441-450; Kuhn and Strotkoetter, 1967:I-57; Mattheij and van Oordt, 1967:472; Pfeiffer,1967b:218, 224, 225; Popper and Tavolga, 1967:791; Reddell, 1967b:82; Russell and Raines,1967:79, 84; Schemmel, 1967:255-316; Thineset aI., 1967:438,442; Barr, 1968a:77, 89, 90,91;Frank, 1968:299-391; Glaser, 1968:648-658; Mattheij,1968a:588-595; Mattheij, 1968b:542-553;Wilkens, 1968:454-464; Campos, 1969:255,256,259, 263, 265, 266, 277-281, 292, 298, fig. 1, 4,6,9; Frank, 1969:11-13; Fujii, 1969:324; Kleerekoper,1969:123; Mattheij, 1969:588-597; Mattheijand Sprangers, 1969:411-419; Mattheij et aI.,1969:519-520; Sadoglu and McKee, 1969:10-14;Thines, 1969:frontispiece, 10, 20, 24-25, 27, 55,57-62, 235, 247, 304, 305, 335, 336, 344, 347349, 352-354, 356-357, 360, fig. 13-14,97, 113;Alvarez, 1970:44; Delamare-Deboutteville andBotosaneau, 1970:75; Gertychowa, 1970:9-69;Love, 1970: 186, fig. 74; Mattheij, 1970a:lxix;Mattheij, 1970b:91-106; Walker, 1970:6-7,44-50;Weiss and Martini, 1970:34-37, fig. 1; Whitt andMaeda, 1970:728-731,739-740, fig. 3, 9a; Wilkens,1970a:I-47; Wilkens, 1970b:173; Wilkens, 1970c:56, 59; Bauer, 1971:56, 121; Gertychowa, 1971:66-69; Glydenholm and Scheel, 1971 :480; Hara,1971:93, 94, 108; Heuts and Thines, 1971:147;Reddell, 1971b:l; Avise and Selander, 1972:2;De Buen, 1972:119,121; Durand, 1972:621,622,690, 693; Herald, 1972:87, 103; Schmatolla,1972:556; Thines and Tercafs, 1972:46, 55, fig.11; Thines and Wissocq, 1972:140, 143-153,167,fig. 2-5; Wilkens, 1972a:l; Wilkens, 1972b:411431; Wilkens and Bums, 1972:263,264,266,268;Zaccone, 1972:106-115; Denton, 1973:99; Thinesand Durand, 1973:275, 276, 277, 278, fig. 5,pI. I(fig. a, e, f), pI. II(fig. a); Thines and Legrain,1973:291-297; Zeitlin, 1973:461462; Zeitlin andMcDevitt, 1973:357a; Pfeiffer, 1975:708; Thinesand Capon, 1975:209,216; Vives, 1975:119-123;Migdalski and Fichter, 1976:138, 139; Durand,1977:150; Kirby et aI., 1977:578; Mitchell et aI.,1977:4, 5; Miyashita et aI., 1977:?; Wilkens,1977a:143, 146; Zaccone, 1977:31-38, fig. 1-2;Thines and Piquemal, 1978: 197; Thines andWeyers, 1978:36, 39; Coss, 1979:341; Coss andGlobus, 1979:348.Anoptichthys: Anonymous, 1940:221; Breder andGresser, 1941a:112; Breder and Gresser, 1941c:289-296, pI. 1; Anonymous, 1942a:221; OsorioTafall, 1943:45, 46, 49, 52, 55, 57-67, 68, 69;Walls, 1942:387; Osorio Tafall, 1946:153; Pavan,1946:343; Alvarez, 1950:13, 26; Bonet, 1953b:238,241,242; Rioja, 1953a:287; Thines, 1955:27,119, 121, 123; Olivereau and Francotte-Henry,1955-1956:144; John, 1957:123-132; Sadoglu,1957a:394; Bertin, 1958a:1918, 1919, 1920,1921, 1922, 1923; Bertin, 1958b:2661; Alvarez,1959:78; Breder, 1959:420, 422, 426,429,431,fig. 7; Kosswig, 1959:50,55-56; Kosswig, 1960b:495,496,500,504,505-506,507,508-510; Kuhn,1960:418-483; Thines, 1960:46, 47, 48, 49;Frank, 1961:360; Poulson, 1961:14, 171, 175,177; Kosswig, 1963:205-239; Pfeiffer, 1963:123,127, 128, 129, 142, 144, 145, 146; Poulson,1963:285,286; Steven, 1963:212,227,228,229,230,231; Kosswig, 1964:72-74, fig. 11-13; Vandel,1964:19, 52,382, 412,413, 420,459,462, 474,495, 505, 506, pI. 10; Fingerman, 1965 :323; Kosswig,1965:233-234,246-256, fig. 11-12; McKenzie,1965a:34; McKenzie and Russell, 1965:30; Reddell,1965c:6; Vandel, 1965a:11, 39, 222, 350,356, 387, 402, 418, 428, 430, 434, 483, pI. 10;Greenwood, 1967:266,269,271; Husmann, 1967:417; Johnson, 1967:6-9, 23, 26, 45,47,48; Kosswig,1967:441-450; Kosswig and Peters, 1967:828-835; Pfeiffer, 1967a:365-378; Pfeiffer, 1967b:218-245; Reddell, 1967c:55; Thines et aI., 1967:437-448, pI. 73-76; Barr, 1968a:77, 78, 88-89;Durand, 1968:352; Peck, 1968:96; Peters andPeters, 1968:212,220,231,233,234,238; Campos,1969:259,265,284,286,287,288,289,293,294, 295, 296, 297; Fujii, 1969:335,343; Kleerekoper,1969:127; Thines, 1969:5, 15,55-57,69,214, 223-225, 233-235, 239-241, 273, 276, 300,301, 304-307, 317, 322, 327, 329, 332-336,338-344, fig. 100, 114-115; Alvarez, 1970:43,44,45, fig. 47; Fenwick, 1970:98; Strinati, 1970:2;Vandel and Durand, 1970:2699; Culver and Poulson,1971:83; Delamare-Deboutteville, 1971:69;Gertychowa, 1971:67; Heuts and Thines, 1971:113, 117, 140, 141, 158, 159, 160, 161; Rouch,1971:108; Durand, 1972:693, 760; Thines andTercafs, 1972:55, 64, 135, 136, 137, 143-144,147; Thines and Wissocq, 1972:139, 140, 142,154, 156, 157, 158, 159, 160, 161, 162, 163-167,168; Wilkens and Burns, 1972:263-269; Petersand Peters, 1973a: 187-201; Peters and Peters,1973b:301-313; Thines and Durand, 1973:275,277, 280; Juberthie, 1974:81, 82; Peters et aI.,1975:110, 116, 123; Pfeiffer, 1975:706-711;Thines and Capon, 1975:209, 221, 225, 227;Durand, 1976:181, 219, fig. I; Durand, 1977:149-161, pI. 1-4; Mitchell et aI., 1977:5; Yew andYoshihara, 1977:175; Thines and Piquemal, 1978:240

196,197,198,202-203; Thines and Weyers, 1978:36, 39; Durand, 1979:270-274, pl.(fig. 3-5).Anoptichthys n. subsp.: Osorio TafaU, 1942:210.Anoptichthys n. sp.: Breder, 1943c: 169,175.Blind characins: Bridges, 1943:90; Heuts, 1953b:300.Anoptichthys sp.: Breder and Rasquin, 1943:169-200, pI. 1-3; Alvarez, 1946:265, 267, 268, 270,271,272,274,275-278,280,281, fig. 9-15; Bolivar,1946:27; De Buen, 1946:127; Rasquin, 1946:578-579; Breder and Rasquin, 1947a:323-351;Cahn, 1958:76-107, pI. 16-20, 22-24, 26-27;Tamayo, 1962:vol. III:260; Walters and Walters,1965:22; Reddell, 1967c:55; Schemmel, 1967:255-316. Rance and Baker, 1979:65,66.Blind fish: Breder, 1944:131-143, pI. 1; Mannix,1947:59; Mohr, 1950:5; Heuts, 1953a:391-392;Warner and Beer, 1957:17; Longsworth, 1959:117, 119; Lanham, 1962:93, pI. 23; McKenzieand Russell, 1965:31, 32; l\lohr and Poulson,1966:181, 182; Reddell, 1967b:82; Russell andRaines, 1967:53, 69; Bauer, 1971:120; Elliott,1972:129, 130, 131; Walsh, 1972:15, 23; Kawakatsu,1973a:?; Oguri and Omura, 1973:419,421,424,434, fig. XV-10; Reddell, 1973b:77-78.Anoptichthys antrobius Alvarez, 1946:265,267,268,270, 271, 272, 274, 276, 278-281, fig. 16-23;Alvarez, 1947:219; Breder and Rasquin, 1947a:323-351; Alvarez, 1950:28; Breder and Atz, 1952:262; Bonet, 1953b:241; Thines, 1954:36; Bridges,1955:268; Thines, 1955:9, 41, 70-71, 108-109;Bonet, 1956a:83; Bonet, 1956d:250; Dearolf,1956:202; Sadoglu, 1956:113-114; Hahn, 1960:611; Thines, 1960:44; Heuts, 1961:196, 197;Poulson, 1961:172, 174; Nicholas, 1962:184;Bonet, 1963a:99; Kosswig, 1963:220, fig. 2;Poulson, 1963:286, 287, 288; Franck, 1964:95107; Kosswig, 1964:73; Vandel, 1964:262,474;Boucquey et al., 1965:79-103; Kosswig, 1965:251, fig. 10; Reddell, 1965a:20; Thines and WolffVan Ermengem, 1965:585; Thines et aI., 1965:61115; Vandel, 1965a:222, 402; Vandenbussche etaI., 1965:586; Peters and Peters, 1966:393-414;Pfeiffer, 1966:97-105; Johnson, 1967:6-7, 12-13,27,28,30,32,34,36,37,38,39,40,41,44,46,48, 49, fig. 7, 9; Pfeiffer, 1967a:366, 376, 377;Pfeiffer, 1967b:218, 219, 224, 225, 226, 232,242, 243; Reddell, 1967b:82; Russell and Raines,1967:58; Sadoglu, 1967:541-549; Schemmel,1967:255-316; Thines et aI., 1967:438; Wilkens,1968:454-464; Sadoglu and McKee, 1969:10-14; Thines, 1969:10, 25, 27, 64-66,344-345,348, 352, 356, 360-363, fig. 100, 123; Alvarez,1970:45; Gertychowa, 1970:9,10; Strinati, 1970:3, fig. 1; Wilkens, 1970a:1-47; Wilkens, 1970b:173; Wilkens, 197Oc:54-75; Gertychowa, 1971:66,67; Wilkens, 1971:530-543; Avise and Selander,1972:3; Thines and Tercafs, 1972:55; Wilkens,1972a:l; Wilkens, 1972b:412, 414, 417, 421;Wilkens and Burns, 1972:263, 266; Schemmel,1973:215; Mitchell et al., 1977:4,5,57; Wilkens,1977a:139.Anoptichthys hubbsi Alvarez, 1947:215-219, fig. 1;Rasquin, 1949a:499-531; Rasquin, 194%:131,133; Alvarez, 1950:28; Bolivar, 1950:216; Heuts,1951:223; Bolivar, 1952:295,296; Bonet, 1953b:241; Olivereau and Herlant, 1954:52; Thines,1954:36; Bridges, 1955:268; Thines, 1955:9,41,70, 108-109; Olivereau and Francotte-Henry,1955-1956:143; Bonet, 1956a:86; Bonet, 1956d:247; Dearolf, 1956:202; Sadoglu, 1956:113-114;Gordon, 1957:467; John, 1957:123-132; Pickfordand Atz, 1957:159; Alvarez, 1959:79; Breder,1959:423,424,425,436,437,446,447, fig. 4-6;Hahn, 1960:611; Humbach, 1960a:1-88; Humbach,1960b:551-552; Thines, 1960:44; Heuts,1961:196; Poulson, 1961:172, 173, 174; Nicholas,1962:184; Bonet, 1963a:98; Poulson, 1963:286,287, 288; Moore and Nicholas, 1964:94; Vandel,1964:262, 414,466, 474, 482; Boucquey et aI.,1965:88; Thines et aI., 1965:65; Vandel, 1965a:222, 351, 395, 402, 407; Peters and Peters, 1966:393-414; Johnson, 1967:6-7, 11-12, 28, 30, 32,34, 36, 37, 38, 39, 40, 41, 44, 46, 50, fig. 7,9;Pfeiffer, 1967b:218, 225; Reddell, 1967b:82;Schemmel, 1967:255-316; Thinesetal., 1967:438;Wilkens, 1968:454-464; Sadoglu and McKee,1969:10-14; Thines, 1969:10,20,25,27,62-68,335, 348, 356, fig. 97; Alvarez, 1970:45; Gertychowa,1970:9,65; Wilkens, 1970a:1-47; Wilkens,1970b:173; Wilkens, 1970c:54-75; Gertychowa,1971:67; Wilkens, 1971:530-543; Avise and Selander,1972:2; Thines and Tercafs, 1972:55;Wilkens, 1972a:l; Wilkens, 1972b:412, 414, 416,421; Wilkens and Burns, 1972:263,266; Mitchellet al., 1977:4,5,51; Wilkens, 1977a:139.Anopitchthys hubbsi: Alvarez, 1947:215 (erroneousspelling).Characinidae of Mexican caves: Allee et al., 1949:643.Anoptichthys undescribed species: Rasquin, 1947:3542, pl. 1.Anopthychtys: Rioja, 1953a:294 (erroneous spelling).Anopichthys antrobius: Martin and Martin, 1954:584 (erroneous spelling).241

Anophthyctys jordani: Stefanelli, 1954a:436-438(erroneous spelling).Cave 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 forms 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, Mexican 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 form: Kosswig,1973:13; Schemmel, 1973:209-219; Wilkens,1973c:55, 57-58; Durand, 1976:18; Parzefall,1976:29; Thines and Piquemal, 1978:197.Mexican 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 form: 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 form 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 Cave,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 for 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 environmentfor A. mexicanus and by the epigean environment foriI. 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

Prietella phreatophila Carranza, 1954:129-136, fig. 1,pI. I; Smith, 1956:15-16, 41, 42, 43, 44, pI. 27;Suttkus, 1961:55, fig. 5; Nicholas, 1962:184; Vandel,1965a:224; Husmann, 1967:421; ContrerasBalderas, 1969:294; Taylor, 1969:1, 14, 17, 18,220, fig. 5; Thines, 1969:28,90-95,215, fig. 29;Alvarez, 1970:72; Benjamin, 1970:18322; Hubbs,1971:91; Reddell, 1971b:83; Reddell and Mitchell,1971c:2; Thines and Tercafs, 1972:56; Thinesand Durand, 1973:276; United States Office ofEndangered Species and International Activities,1974:5; Longley and Karnei, 1979a:7, fig. 3;Longley and Karnei, 1979b:8, fig. 4; Moore andNicholas, 1978:114.Prietella: Smith, 1956:33-34, fig. 1; Suttkus, 1961:57,63,64; Taylor, 1969:5, 10, 12, 13, 15, 16, 18,24, 217, 310; Thines, 1969:263,267,322; Alvarez,1970:71, 72; Juberthie, 1974:81.Prietella phreaticola:spelling).Vandel, 1964:264 (erroneousPriatella phreatophila: Cole and Minckley, 1966:22(erroneous spelling).Type.locality.~Pozo EI Potrero de Dona Mariana,Municipio de Musquiz, Coahuila, Mexico.Distribution.-Known only from the type-locality.See Fig. 85.///II\\;..... r-.,r'..J' .......·-·-rI•'""." \\I'" \(-"\.~I l., "\\ ~",J "(,\\'""'\ ......-...'-'\.r" '--_ ,.: I;",_1 "--L. ._.IU; .....t~ I" i :"'-;::i......" i

Discussion.-Prietella phreatophila is one of onlythree troglobitic catfishes described from North America.The other two species are Satan eurystomusHubbs and Bailey and Trogloglanis pattersoni Eigenmann,both known only from deep artesian wells inand near San Antonio, Bexar County, Texas (U.S.A.).It is notable that all three species of blind ictaluridsare found only in phreatic waters. Prietella is mostclosely related to the genus Noturus, which rangesnortheast from Texas into Canada. Prietella phreatophilahas been collected only in an artificially enlargednatural well fed by a deep crevice. Following!:teavy rains, water gushes from the well, and manyfish are reported to be washed out at that time. Thewell is frequently cleaned with algaecides, and fishmay be found only sporadically in the wells. Thisspecies has been placed on the list of endangeredforeign species.Family PimelodidaeThe pimelodid catfishes of the genus Rhamdiafrom the caves and cenotes of Yucatan have beenstudied by Hubbs (1936, 1938). His revision of thisgroup includes four subspecies associated with subterraneanwaters: R. guatemalensis decolor Hubbs,R. g. depressa Barbour and Cole, R. g. sacrificiiBarbour and Cole, and R. g. stygaea Hubbs. Rhamdiag. decolor is somewhat depigmented, and R. g. stygaeahas somewhat smaller eyes than the remainingsubspecies. Both of these subspecies are known onlyfrom caves. A troglobitic species of Rhamdia fromcaves in the Acatlan region of Oaxaca is presentlybeing described.Order CyprinodontifonnesFamily PoeciliidaeThe family Poeciliidae is represented in Mexicancaves by the troglobitic population of Poecilia sphenopsdiscussed below and by two other species.Hubbs (1936) reported the presence of P. sphenopsaltissima (Hubbs), P. velifera (Regan), and Gambusiayucatana Regan in the cenotes of Yucatan. Theseshould not be considered a part of the true cavefauna.Poecilia sphenops ValenciennesPoecilia sphenops Valenciennes (in Cuvier and Valenciennes,1846):130.Poecilia sphenops, cave population: Gordon and Rosen,1962:360-368, fig. 1-8; Rosen and Bailey,1963:49-53; Kosswig, 1964:74; Moore and Nicholas,1964:95; Walters and Walters, 1965:214-223;Barr, 1968a:89; Peters and Peters, 1968:211-244;Reddell, 1971b:84; Zeiske, 1971:387-393; Parzefalland Wilkens, 1972:64-65, 76; Wilkens andBurns, 1972:269; Kosswig, 1973:5, 10; Parzefall,1973a:177-183, fig. 1-3; Parzefall, 1973b:283290; Peters, 1973:315-326; Peters et al., 1973:417-436, fig. 2-5; Juberthie, 1974:82; Peters et aI.,1975:111; Thines and Capon, 1975:226; Besharseand Brandon, 1976:538; Parzefall, 1979:399-422,fig. 1-10.Mollienesia sphenops, cave population: Poulson,1964:756; Zeiske, 1968:190-222, fig. 1-13; Parzefall,1969:1-37; ParzefaII, 1970:323-342; Wilkens,1970c:65.Mollienesia: Poulson, 1964:755.Discussion.-Poecilia sphenops has been reportedas a cave-adapted fonn only in Cueva del Azufre,Tabasco. The troglobitic population has never beengiven taxonomic recognition. The situation of P.sphenops in Cueva del Azufre is remarkably similarto that of Astyanax jordani and A. mexicanus inCueva Chica, San Luis Potosi. The stream emergingfrom Cueva del Azufre has permitted epigean fish tore-invade the cave habitat, and a limited amount ofhybridization with the derivative troglobitic fish isnow occurring. It is not possible to traverse a greatdistance into Cueva del Azufre and, therefore, theeyeless fish population remains unstudied. It is ofgreat importance to explore this region extensivelyin an attempt to locate the source of the eyeless fishnow hybridizing with the eyed P. sphenops. Waltersand Walters (1965) have made some laboratory observationson this species, and the European workersN. and G. Peters, J. Parzefall, H. Wilkens, and E.Zeiske have studied the morphology and geneticsof the cave-adapted form of this species. NormaleyedP. sphenops have been collected in Cueva delRio Subterraneo and in a well near Cueva Chica,San Luis Potosi.Order PercifonnesFamily BrotulidaeTyphliasina pearsei (Hubbs)Typhlias pearsei Hubbs, 1938:261, 263, 265, 267,269, 290, 291, pI. 3; Chitwood, 1938:62; DeBuen, 1940:66; Anonymous, 1947:128; Pearse,1945:112, 113, 151,214; Alvarez, 1950:15,109,110; Cardenas Figueroa, 1950:156; Robles Ramos,1950:64; Ladiges, 1951b:168; Dearolf, 1956:203;Barrera, 1962:84; Tamayo, 1962:vol. III:326; Polland Leleup, 1965:466, 467; Breder and Rosen,1966:385; Greenwood, 1967:267, 270, fig. 58;De Buen, 1972:269; Bravo-Hollis and CaballeroDeloya, 1973:74,99.245

TyphlilJs: Pearse, 1938a:13, 15; Hubbs, 1938:261,287-290; Walls, 1942:388;Osorio Tafall, 1946:156;Pearse, 1945:135, 214;Alvarez, 1950:109;Bolivar,1950:216; Cardenas Figueroa, 1950:157; Whitley,1950:67; Poll and Leleup, 1965:466.Typhlias pearsi: Osorio Tafall, 1943:68-69; Thines,1955:42, 82, 108-109; Thines, 1960:45, 50, 52,pI. 2; Tamayo, 1962:vol. III:261; Vandel, 1964:271, 300; Vandel, 1965a:230, 252; Greenwood,1967:264, fig. 56B; Delamare Deboutteville,1971:68 (erroneous spelling).Tiphlias pearsei: Carreno, 1950:24 (erroneous spelling).Typhliasina pearsei: Whitley, 1950:67; Solorzano,1953:286; Nicholas, 1962:185; Rioja, 1962:38;Alvarez, 1970: 150; Andrews, 1970:4; Reddell,1971b:84; Wilkens, 1973a:329-330, fig. 1; Wilkens,1973b:204, 205; Wilkens, 1973c:50, 54-55,56, 57,58, fig. 4; Reddell, 1977b:236, 239, 240,258,263,268,281,285.Typhliasina: Whitley, 1950:67; Rioja, 1962:39; Cohenand Robins, 1970:134, 135; Juberthie, 1974:8l.Blind brotula: Woods, 1954:4.Typhlia: Bertin, 1958a: 1919 (erroneous spelling).Blind Hohlenfische: Wilkens, 1960b:499.Blind brotulid of Mexico: Norman, 1963: 194_Yucatan Brotulidae: Norman, 1963:196.Typhliasina pearsi: Vandel, 1964:271; Vandel,1965a:230; Thines, 1969:29, 135,172-174,250,fig. 73-74; Delamare Deboutteville and Botosaneau,1970:75; Thines and Tercafs, 1972:63, 138,fig. 23 (erroneous spelling).Typhlyasina: Argano, 1972b:33 (erroneous spelling).Typhliasina pearcei: Wilkens, 1973b:204 (erroneousspelling).Type-Iocality.-Balaam Canche Cave (=Grutas deBalankanche), near Chichen Itza, Yucatan, Mexico.Distribution.-Known only from five caves in Yucatan.See Fig. 85.Records.-Yucatan: Gru tas de Balankanche, Cenotede Calchuhuim, Cenote de Hoctun, Cenote delPochote, and Grutas de Tzab-Nah.Discussion.-Typhliasina pearsei is eyeless and depigmented.It was collected from a deep lake in theinner passage in Grutas de Tzab-Nah and from shallowpools connected to the groundwater in the othercaves. Wilkens (1973a, 1973b) contends that this speciesis a very old troglobite, based at least in part onthe degree of reduction of the eyes. Chitwood (1938)has reported the presence of the nematode Rhabdochonakidderi Pearse in the intestine of this species_Four other freshwater brotulids, all troglobites, areknown: Caecogilbia galapagosensis Poll and Leleupfrom deep crevices on Isla de Santa Cruz, GalapagosIslands; Lucifuga (5tygicola) dentatus (Poey) and1. (Lucifuga) subterraneus Poey from Cuba; and1. (5.) spelaeotes Cohen and Robins from '\lew Providence,Bahamas. Cohen and Robins (1970) believethat l'yphliasina is more closely related to the marinegenera Dinematichthys and Ogilbia than to Lucifuga.Vandel (1964) speculates that the troglohitic Lrotulidshave been derived from marine littoral specieswhich inhabited cavities in coral reefs.Family CichlidaeTwo species of cichlids, Cichlasoma meeki (Brind)and C. urophthalmus (Gunther), have been reportedfrom the subterranean waters of Yucatan (HubLs,1936, 1(38). Cichlasoma meeki is known only froman artificial cenote and is probably a species introducedby man. Cichlasoma urophthalmus containsfour subspecies associated with caves and cenotes:C. u. conchitae Hubbs, C. u. ericymba Hubbs, C. u.mayorum Hubbs, and C. u. zebra Hubbs. The onlyone of thesc which seems to be dosely associatedwith the cave habitat is C. u. ericymba, which isknown only from Cenote de Sambula (Merida).Cichlasoma cyanoguttatum (Baird and Girard) hasbeen collected in Cueva del Carrizal, Nuevo Leon.Order SynbranchiformesFamily SynbranchidaeOphisternon infernale (Hubbs)Pluto infernalis Hubbs, 1938:261, 263, 265, 269,270,292-293, pI. 4; De Buen, 1940:6; Osorio Tafall,1943:68-69; Anonymous, 1947:128; Pearse,1945:112,114,208; Alvarez, 1950:15, 23;CardenasFigueroa, 1950:156, fig. 4; Carreno, 1950:24;Robles Ramos, 1950:64; Ladiges, 1951b:168;rhines, 1955:41, 71,108-109; Dearolf, 1956:203;Barrera, 1962:84; Mees, 1962:27,28,31,32; Tamayo,1962:vol. III:261, 320; Vandel, 1964:266;Vandel, 1965a:226; Andrews, 1970:4; DelamareDeboutteville and Botosaneau, 1970:75.Pluto: Hubbs, 1938:261, 291-292; Osorio Tafall,1946:156; Pearse, 1945:111, 135, 207; Alvarez,1950:23; Bolivar, 1950:216; Whitley, 1950:67;Bertin, 1958b:2661; Vandel, 1964:272; Vandel,1965a:231.Furmastix infernalis: Whitley, 1950:67; Nicholas,1962:184; Vandel, 1964:266; Vandel, 1965a:226;Alvarez, 1970:130; de Lachica 8., 1970:65, 122;Herald, 1972: 139; Parzefall and Wilkens, 1972:6379, fig. 2-3, 4h, 5b; Rosen and Rumney, 1972:1,246

4, 11, 27, 28, 29, 40, 41, 42, 43, 44, fig. 13-14,28, 31; Wilkens, 1973a:329, fig. 2; Wilkens,1973b:204, 205; Wilkens, 1973c:55, 56,57; Rosenand Greenwood, 1976:6; Durand, 1977:17.Furmastix: Whitley, 1950:67; Mees, 1962:32.Synbranchus infemalis: Mees, 1962:27, 28, 31, 32;Thines, 1969:18, 23-24, 29, 182-183, 261, 271272, fig. 77; Thines and Tercafs, 1972:63.Symbranchus infemalis: Reddell, 1971b:85; Thinesand Durand, 1973:275.Furmastik infemalis: De Buen, 1972:197 (erroneousspelling).Ophistemon infemale: Rosen and Greenwood, 1976:8, 10, 13, 18,22, 23, 25, 28, 34, 36, 37,42,43,45, 47,52,60,61,62,63,64,fig. 7,8,31,45,66,67; Reddell, 1977b:236, 237,239,24~258,268,281, 285, fig. 7-8; Reddell, 1977c:cover, insidetitle page.Ophistemon: Rosen, 1976:434,435,436,442,457,458, fig. 1,2, 2l.Type-locality.-Hoctun Cave (=Cenote de Hoccin),at Hocrun, between Merida and Chichen Itza, Yucatan,Mexico.Distribution.-Known only from four caves in Yucatanand possibly two caves in Quintana Roo. SeeFig. 85.Records.-Quintana Roo: ?Actun Ha and ?Cenotede Santo Domingo; Yucatan: Grutas de Balankanche,Cenote de Hoccin, Cenote del Pochote, and Grutasde Tzab- ah.Discussion.-The genus Ophistemon, as revised byRosen and Greenwood (1976), includes a diversegroup of species from both the Old and New Worlds.The only New World species, other than O. infemale,is O. enigmaticum Rosen and Greenwood, whichranges from northern South America into southernMexico. One other species, O. candidum (Mees) fromWestern Australia, is a troglobite. The zoogeographyof the genus has been discussed by Rosen (1976).Ophistemon infemale has been found resting on theguano-covered floor of a large pool near the end ofCenote de Hoccin, buried partially in mud in a smallpool below the inner skylight entrance to Grutas deTzab-Nah, and swimming along the silt floor of apool in Grutas de Balankanche. The two questionablerecords from Quintana Roo are based on sightings ofsingle individuals partially buried in silt and rocks insmall shallow pools in each cave.Class AmphibiaOrder UrodelaAlthough several species of salamander in the UnitedStates and one in Europe have become sufficientlyadapted to cave life to be classified as trogl

Family BufonidaeFour species of toad of the genus Bufo have beenfound in Mexican caves: Bufo debilis Girard wastaken in Cueva de Dos Cuartos, San Luis Potosi; B.occidentalis Camerano was found in S6tano del Gobernador,Queretaro; B. valliceps Wiegemann wasfound in caves and cenotes in Yucatan (Gaige, 1936);and B. marinus (Linnaeus), reported from cenotes inYucatan by Barbour and Cole (1906) and Gaige(1936), has been collected in Hoyo de Don Nicho,Chiapas, and Cueva de la Virgen de Guadalupe,Tamaulipas.Family HylidaeSix species of hylid frog have been found in cavesand cenotes. Agalychnis callidryas (Cope) was reportedfrom Cenote de Thompson, Yucatan (Gaige,1936). Triprion petasatus (Cope), described fromCenote Taamanche, Yucatan (Cope, 1865), has beenfound in Cenote de Santa Elena, Yucatan (Kellogg,1932). Four other species have been taken fromMexican caves: Hyla staufferi staufferi Cope fromHoyo de Don Nicho, Chiapas; H. taeniopus Guntherfrom Sima Esteban, Puebla; Plectrohyla sp. d. sagorumHartweg from the streams in Chen SibilmutMEXICO(/C" ,/GUATEMALAr\,-"ro-"~///Fig. 86.-Distribution of cavernicole salamanders of the family Plethodontidae: 1, Chiropterotriton multidentata; 2, C.multidentata and Psudoeurycea scandens; 3, C. chondrostega, C. multidentata, P. belli, and P. scandens; 4, C. multidentata andThorius sp.; 5, C. arborea and C. magnipes; 6, C. magnipes; 7, C. arborea; 8, C. mosaueri; 9, Bolitoglossa yucatana.248

and Cueva de Mapachero, Chiapas; and Smiliscabaudinii (Oumeril and Bibron) from three caves inthe Sierra de EI Abra, San Luis Potosi.Family LeptodactylidaeThe family Leptodactylidae is closely associatedwith caves in Central America, the Antilles, andTexas (U.S.A.). Five genera have been found inMexican caves, the most important of which areEleutherodactylus and Syrrhophus (see Fig. 87).Six species of Eleutherodactylus have been foundin Mexican caves: E. alfredi (Boulenger) from Satanode los Perros, Veracruz; E. guerreroensis Lynch fromCueva del Nacimiento del Rio San Antonio, Oaxaca;E. decoratus decoratus Taylor from caves in Hidalgo,Queretaro, San Luis Potosi, Tamaulipas, and Veracruz;E. decoratus purpurus Lynch from caves inOaxaca and Tamaulipas (Lynch, 1967); E. rhodopis(Cope) from Grutas de Atepolihuit, Puebla; E. spatulatusSmith from caves near Huautla de Jimenez,\.," ., ","' ) \'.,MEXICOGUATEMALAii\./)(._._._.-/.//r·\..... r·.-'Fig. 87.-Distrihution of cavernicole leptodactylid frogs of the genera Eleutherodactylus and Syrrhophus: 1, Syrrhophuscystignathoides and S. longipes; 2, S. dennisi; 3, S. longipes; 4, Eleutherodactylus decoratus decoratus, E. decoratus purpurus,and S. longipes; 5, E. decoratus pUrpUTUS and S. longipes; 6, S. dennisi and S. longipes; 7, S. cyst;gnathoides; 8, S. guttilatus;9, E. decoratus decoratus and S. cystignathoides; 10, E. decoratus decoratus and S. longipes; 11, E. decoratus decoratus;12, S. verrucipes; 13, E. rhodopis; 14, E. alfredi; 15, E. decoratus purpurus and E. guerreroensis; 16, E. spatulatus; 17, E.yucatanensis.249

Oaxaca; and E. yucatanensis Lynch from a cave atPueblo Nuevo X-Can, Quintana Roo (Lynch, 1965).Five species of the genus Syrrhophus have beenfound in Mexican caves (Lynch, 1970): S. cystignathoides(Cope) from caves in Nuevo Leon, SanLuis Potosi, and Tamaulipas; S. dennisi Lynch fromcaves in the Sierra de EI Abra, San Luis Potosi andTamaulipas; S. guttilatus (Cope) from Cueva de DosCuartos, San Luis Potosi; S. longipes (Baird) fromcaves in Nuevo Leon, Queretaro, San Luis Potosi,and Tamaulipas; and S. verrucipes Cope from CuevaGrillo de la Mona, Puebla.Four other species of leptodactylid have beentaken from Mexican caves: Hylactophryne augustiaugusti (Duges) from caves in San Luis Potosi andTamaulipas; H. a. cactorum Taylor from caves inPuebla; Leptodactylus labialis (Cope) from ActUnChen, Campeche; L. melanonotus (Hallowell) fromCueva de Sala de Agua Grande, Veracruz; and Tomodactylusnitidus nitidus (peters) from Grutas delMogote, Guerrero.Family RanidaeThe leopard frog, Rana pipiens (Schreber), isknown from caves in Hidalgo, Quintana Roo, SanLuis Potosi, Tamaulipas, and Yucatan.Class ReptiliaOrder CheloniaFamily KinosternidaeTwo species of turtle, Kinosternon creaseri Hartwegand K. cruentatum Dumeril and Bibron, havebeen reported from large open-aired cenotes in Yucatan(Duellman , 1965). A third species, K. integrumLe Conte, was reported from a cave at Raboso,Puebla, by Smith and Van Gelder (1955).Order SquamataFamily BoidaeA boa, Constrictor constrictor imperator (Daudin),was collected from Cueva de Tezoapa, Guerrero,where it presumably was preying on bats (Villa R.and Lopez-Forment, 1966).Family ColubridaeVilla R. and Lopez-Forment (1966) reported thepresence of Elaphe flavirufa flavirufa (Cope) in Cuevade la Sepultura, Tamaulipas; it was probably preyingon bats. Three other species of colubrid snake havebeen found in Mexican caves: Rhadinaea crassa Smithfrom caves in the Sierra de Guatemala, Tamaulipas;Storeria dekayi (Holbrook) from Sotano de Tlamaya,San Luis Potosi; and Tropidodipsas sartorii sartoriiCope from a cave at Pueblo Nuevo X-Can, QuintanaRoo (Duellman, 1965), and ActUn LoltUn, Yucatan(Gaige, 1938).Family CrotalidaeThe fer-de-lance, Bothrops atrox asper (Garman),is frequently found in the entrance area of caves inthe Sierra de EI Abra, San Luis Potosi. It is alsoknown from caves in Veracruz and Yucatan. A rattlesnake,Crotalus durissus tzabcan Klauber, has beenfound in the entrance area of caves in Campeche andYucatan.Family GekkonidaeTwo gekkos, Coleonyx elegans elegans Gray andThecadactylus rapicaudus (Houttuyn), have beenfound in many caves in Yucatan (Gaige, 1938). Theyare frequently seen on the walls near the cave entrances.Family IguanidaeThree species of iguanid lizard have been found incaves and cenotes in Yucatan: Anolis lemurinus bougeaeiBocourt, Basiliscus vittatus Wiegmann, andCtenosaura similis (Gray). All three species probablyonly utilize the cave entrance area for shelter.Family XantusiidaeThe lizards of the family Xantusiidae are closelyassociated with the cave habitat, and three Mexicanspecies have been found in caves. Lepidophyma micropholisWalker inhabits caves in the Sierra de ElAbra, San Luis Potosi and Tamaulipas (Walker,1955b). It has been taken both from the twilightzone and in total darkness. Smith and del Toro (1977)described L. lipetzi from a cave north of Cintalapa,Chiapas. Mautz and Lopez-Forment (1978) discussedthe cavernicolous habits of L. smithii Bocourt incaves near Puerto Marquez, Guerrero.Class AvesSixteen species of bird have been recorded fromcavernicolous habitats in Mexico (see Table 30).Many of these utilize the cave entrances for temporaryshelter, but others build their nests in the protectedentrance rooms of caves. A few of the moresignificant species are briefly discussed below.Order FalconifonnesFamily CathartidaeThe vulture Coragyps atratus (Bechstein) has beenobserved nesting in three caves at the ruins of Ake,Yucatan.250

Order PsittacifonnesFamily PsittacidaeMany species of parrot inhabit caves in Mexico,but very few "records are available. The green parakeet,Aratinga holochlora (Sclater), has been reportednesting in the entrance of Hoya de las Guaguas andother deep pits in the Xilitla region of San Luis Potosi(Ukrain, 1979; Whitacre, 1979); they also havebeen observed nesting in the entrance passage atCueva de los Pajaros, Tamaulipas. The beautiful militarymacaw, Ara militaris Linnaeus, has been observednesting in Sotano de los Guayacamos, Tamaulipas.Order StrigifonnesFamily StrigidaeThe mottled wood-owl, Ciccaba virgata tamaulipensis(Phillips), has been reported from caves in theSierra de Guatemala (Harrell, 1951) and Sierra deTamaulipas (Martin et aI., 1954), Tamaulipas.Family TytonidaeThe barn owl, Tyto alba (Scopoli), nests in manycaves. This species has been reported from caves inOaxaca (Mones, 1968), Puebla (Flannery, 1967),Sonora (Bradshaw and Hayward, 1960), Tamaulipas(Holman, 1970); and Alta Verapaz, Guatemala (Land,1968); it is also known from caves in San Luis Potosiand Yucatan, Mexico. This species is doubtless farmore frequent a cave inhabitant than the few recordsindicate.Order ApodifonnesFamily ApodidaeVaux swift, Chaetura vauxi tamaulipensis Sutton,has been reported from several caves at Rancho delCielo in the Sierra de Guatemala, Tamaulipas (Harrell,1951). The white-collared swift, Streptoprocne zonaris(Shaw), has been reported to nest in large numbersin pits in Chiapas (Sbordoni et al., 1977) and SanLuis Potosi (Ukrain, 1979).Family TrochilidaeTwo species of hummingbird have been reportedby Harrell (1951) from caves at Rancho del Cielo,Tamaulipas: the wedge-tailed sabrewing, Campylopteruscurvipennis curvipennis (Lichtenstein), and theamethyst-throated hummingbird, Lampornis amethestinusamethestinus Swainson.Order CoraciifonnesFamily MomotidaeThe turquoise-browed motmot, Eumomota superciliosasuperciliosa (Swainson), is an almost everpresentsight in the caves and cenotes of Yucatan.Their haunting cry is heard in the entrance to almostevery cave, and many caves were found by followingtheir call. Harrell (1951) reported the blue-crownedmotmot, Momotus momota coeruliceps (Gould),from caves at Rancho del Cielo, Tamaulipas.Order PasserifonnesFamily HinmdinidaeTwo species of swallow are common inhabitantsof caves in the Yucatan Peninsula. The cave swallow,Petrochelidon fulva citata Van Tyne, builds its distinctivenests in the large entrance rooms of manycaves. Ridgway's swallow, Stelgidopteryx ruficollisridgwayi Nelson, is also seen in almost every cavewhich possesses a large entrance. The latter specieswas reported by Land (1968) from Cueva Seamay,Alta Verapaz, Guatemala.Family TroglodytidaeHarrell (1951) has reported the presence of thecanyon wren, Catherpes mexicanus (Swainson), IIIthe caves of Rancho del Cielo, Tamaulipas.Family TurdidaeTwo species of thrush have been reported fromcaves at Rancho del Cielo, Tamaulipas (Harrell,1951): the black-headed nightingale thrush, Catharusmexicanus mexicanus (Bonaparte), and the brownbackedsolitaire, Myadestes obscurus obscurus Lafresnaye.Class MammaliaOrder MarsupialiaFamily DidelphidaeTwo species of opossom have been reported fromcaves in Mexico. Villa R. (1967) reported Didelphismarsupialis Linnaeus from Cueva Cerro Hueco, Chiapas,and Pearse and Kellogg (1938) reported D. virginianayucatanensis j. A. Allen from ActUn Ebizt,Yucatan.Order ChiropteraThe importance of bats to the cave ecosystem cannotbe overemphasized. The amount and type ofenergy input provided, to a certain extent, determinethe population size and composition of the invertebratefauna. One reason for the greater diversity ofthe cave fauna in tropical regions is the greater diver-251

sity of bats in the tropics. In addition to the insectivorousbats, which are also common in temperate regions,the tropics contain nectarivous, frugivorous,carnivorous, and sanguivorous species. Observationsin tropical caves have revealed that the guano of eachtype of bat is inhabited by its own distinctive faunalassemblage. Few detailed studies have been made ofthe fauna of caves in the New World tropics withrespect to their population size and the relationshipof this to bat guano. R. W. Mitchell (1970a) hasstudied the fauna inhabiting a passage in Cueva de laFlorida, Tamaulipas, which is inhabited by insectivorousbats. H. A. Mitchell (1965) studied the atmosphereof Cueva del Tigre, Sonora, a large cavecontaining several species of bats. In addition to thesignificance of bats to the study of the cave ecosystem,bats are also of importance to human health.Many species of cave-associated bat in Mexico havebeen found to carry rabies. Furthermore, many caveswhich are inhabited by bats harbor Histoplasmacapsulatum Darling, the causative agent of the dangerouslung disease, histoplasmosis. Constantine(1970) has published a detailed review of the significanceof bats to human health.Although there are numerous records of bats inMexican caves, these records are few when the numberof known caves is considered. The difficulty ofobtaining collecting permits and of collecting andproperly preserving bats has left their study largelyto mammalogists working in the field. Many of thepapers published on bats include references to theiroccurrence in caves, but in many instances onlygeneral localities are given without reference towhether or not the bats were obtained from caves.The bat fauna of the caves of Guatemala and Belizeis even less well known, and there are very few caverecords for these two countries. A total of 72 speciesand subspecies of bat have been recorded fromthe caves of Mexico; only 20 species from Guatemalaand 8 species from Belize are known fromcaves.It is outside the scope of this review to do morethan briefly summarize the bat fauna associatedwith caves in Mexico, Guatemala, and Belize. Villa R.(1967) has published the most comprehensive reporton the bats of Mexico; this study includes manycave records. The only general review of the bats ofGuatemala is that of Jones (1966). No comprehensivestudy appears to have been published on the batfauna of Belize, but a few records were found in apaper on histoplasmosis in Belize (Quinones et al.,1978) and in general taxonomic studies. Records ofbats in the caves of various Mexican states and regionshave been included in the following surveys:Baja California (Jones et al., 1965), Chihuahua (Anderson,1972), Coahuila (Baker, 1956), Durango(Baker and Greer, 1962), Guerrero (Lukens andDavis, 1957), Hidalgo (Carter and Jones, 1978),Jalisco (Watkins et al., 1972), \lichoacan (Hall andVilla R., 1949), Morelos (Davis and Russell, 1952),Nuevo Leon (Jimenez Guzman, 1968), Oaxaca(Goodwin, 1969), San Luis Potosi (Dalquest, 1953a),the Sierra de El Abra, San Luis Potosi and Tamaulipas,and the Sierra de Guatemala, Tamaulipas (:vIollhagen,1971), Sinaloa (Jones et al., 1972), Sonora(Burt, 1938; Cockrum and Bradshaw, 1963), Tamaulipas(Alvarez, 1963), Valley of Mexico (Villa R.,1952), Veracruz (Hall and Dalquest, 19(3), Yucatan(Pearse and Kellogg, 1938), Yucatan and QuintanaRoo (Hatt and Villa R., 1950; Birney et al., 1974),and the Yucatan Peninsula (Jones et al., 1973).Family EmbaUonuridaeFive species of emballonurid bat have been reportedfrom caves in Mexico and Gutemala. Thomas'ssac-winged bat, Balantiopteryx io Thomas, has beenreported from caves in Chiapas, Oaxaca, Tabasco,and Veracruz, Mexico; and Alta Verapaz and habal,Guatemala. This species seems to prefer to hang fromthe tops of pits and crevices, and may be presentwithin the twilight zone as well as in total darkness.Colonies as large as 1000 have been reported fromcaves (Hall and Dalquest, 1963). Peters' bat, H. plicataplicata Peters, is among the more frequentlycollected bats in the caves of southern Mexico. It isknown from caves from San Luis Potosi south intoGuatemala and from Colima east to Tabasco. Theymay inhabit both large, dark caves and smaller, welllitshelter-like openings. The greater doglike bat,Peropteryx kappleri kappleri Peters, is known fromcaves in Tabasco and Veracruz, Mexico; and Izabal,Guatemala. Most of the records are from small cavesand few bats were present. The lesser doglike bat,P. macrotis macrotis (Wagner), is known from cavesin Tabasco, Quintana Roo, Veracruz, and Yucatan,Mexico. This species has been found in both largecaves and small shallow cliff-face recesses. The greaterwhite-lined bat, Saccopteryx bilineata centralisThomas, is known from caves in Campeche, Jalisco,and Oaxaca, but this bat probably prefers to roost inhollow trees.Family MolossidaeSeven species of molossid bat have been recordedfrom caves in Mexico and one 8pecies in Guatemala.Of these, only species of the genus Tadarida are frequentlyencountered in caves. Tadarida aurispinosa252

(peale) is known only from a small cave at EI Salto,San Luis Potosi, and from Cueva del Ahra, Tamaulipas.Two subspecies of the Brazilian free-tailed bathave been found in Mexican caves, and one from acave in Guatemala. Tadarida brasiliensis intermediaShamel is known only from Grutas de Zapaluta, Chiapas,Mexico, and a cave near Coban, Alta Verapaz,Guatemala. The Mexican free-tailed bat, T. b. mexicana(Saussure), is known from caves throughoutmuch of Mexico. This insectivorous species is frequentlypresent in caves in colonies ranging into themillions. The invertebrate fauna associated with theenormous guano deposits of this species is a distinctiveassemblage of species (Mitchell, 1970b). Cockrum(1969) and Villa R. and Cockrum (1962) havestudied the migratory habits and patterns of thisspecies, while Constantine (1967) has studied its activitypatterns. The broad-tailed bat, T. laticaudataferruginea Goodwin, is known only from two cavesin Tamaulipas.Family MonnoopidaeSix species of mormoopid bat have been recordedfrom caves in Mexico, Guatemala, and Belize. Smith(1972) has revised the family, and his names are usedin the discussion here. Peters' ghost-faced bat (alsoknown as the old man bat), Mormoops megalophyllamegalophylla (Peters), has been found in caves fromTexas south into Guatemala. This species is insectivorousand may occur in colonies of up to severalthousand individuals. Mass die-offs, possibly as a resultof disease, have been recorded for this species inCueva del Diablo, Nuevo Leon (Villa R., 1956), andin Sotano de Sauz, Chihuahua (Sprouse, 1977). Theycommonly occur in caves with very high temperaturesand humidities. Davy's naked-backed bat, Pteronotusdavyi fulvus (Thomas), has been found in caves fromSonora and Nuevo Leon, Mexico, south into AltaVerapaz, Guatemala. This species is insectivorous.Parnell's mustached bat, P. parnellii (Gray), is representedin the caves of this region by two subspecies.Pteronotus parnellii mesoamericanus Smith has beentaken in caves in Chiapas, Tabasco, Veracruz, andYucatan, Mexico; the Maya Mountains of Belize; andEI Peten and Alta Verapaz, Guatemala. Pteronotus p.mexicanus (Miller) is known from caves from Tamaulipasand southern Chihuahua into Guerrero. This speciesis frequently taken with other bats, and one individualmay occur in a cluster of another species. Wagner'smustached bat, P. personatus psilotis (Dobson),is known from caves south from San Luis Potosi,Mexico, into Alta Verapaz, Guatemala. This insectivorousspecies is frequently found in large humidcaves in large numbers. The big naked-backed bat,Pteronotus suapurensis (]. A. Allen), is known onlyfrom Cueva Laguna Encantada, Veracruz, and Grutasde Lanquin, Alta Verapaz.Family NatalidaeThe family Natalidae is represented III Mexicancaves only by the two subspecies of the Mexicanfunnel-eared bat, Natalus stramineus Gray, discussedbelow. Natalus stramineus mexicanus Miller is knownfrom caves in Baja California Sur and Sonora. Nataluss. saturatus Dalquest and Hall occurs in caves fromNuevo Leon south into Guatemala. This insectivorousspecies may be present in colonies of several hundredindividuals.Family PhyUostomatidaeThe family Phyll08tomatidae is the largest andmost diverse family of bats occurring in the NewWorld tropics. Thirty-three species and subspecies ofthis family have been reported from the caves ofMexico, Guatemala, and Belize. The nomenclatureused belllw follows that of Jones and Carter (1976).Geoffroy's tailless bat, Anoura geoffroyi lasiopyga(Peters), has been found in caves in Chiapas, Guerrero,Mexico, Morelos, and Tamaulipas. Ten speciesand subspecies of the fruit-eating bats of the genusArtibeus have been recorded from caves in this region.Davis (1969, 1970a, 1970b) has reviewed thesystematics of the genus Artibeus in Middle America.Artibeus aztecus aztecus Andersen is known fromcaves in Guerrero, Mexico, San Luis Potosi, andTamaulipas; A. a. minor Davis occurs in caves in Chiapas.Artibeus hirsutus Anderson is known from cavesin Guerrero, Morelos, and Nayarit. The Jamaicanfruit-eating bat, Artibeus jamaicensis Leach, is representedin the caves of this region by four subspecies:A. j. paulus Davis from Chiapas; A. j. richardsoni J. A.Allen from Tabasco, Mexico, and Escuintla and AltaVerapaz, Guatemala; A. j. triomylus Handley fromGuerrero, Jalisco, Morelos, Michoacan, and Oaxaca;and A. j. yucatanicus Allen from Quintana Roo, SanLuis Potosi, Tamaulipas, Veracruz, and Yucatan.This large species may form colonies containing thousandsof individuals. The areas under their roosts frequentlycontain thousands of sprouted seeds, and theinvertebrate fauna in these areas may be unlike thatin other parts of the cave. The big fruit-eating bat,A. lituratus intermedius J. A. Allen, is known fromcaves in Guerrero, Morelos, San Luis Potosi, andTamaulipas. Artibeus phaeotis phaeotis Miller hasbeen recorded from caves in Quintana Roo and Tabasco.Artibeus toltecus toltecus (Saussure) is knownfrom caves in Chiapas, Morelos, Nuevo Leon, andVeracruz, Mexico; and Alta Verapaz, Guatemala.253

Three species of short-tailed bats of the genusCarollin have been recorded from the caves of thisregion. The nomenclature for 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 Mexican 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 Mexican 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 for the long-legged bat, ,Wacrophyllummacrophyllum Schinz, is that of a cavenear Teapa, Tabasco. This is the northern limit of therange for 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 California 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 California 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 Mexican 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 Mexican 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 BajaCalifornia Sur (Jones et al., 1965). All species ofMyotis are insectivorous.The only pipistrelle known from Mexican caves isPipistrellus subflavus veraecrucis (Ward). It has onlybeen reported from caves on Cofre de Perote, Veracruz.Two species of the genus Plecotus are known fromMexican caves. The systematicsof this genus have beenstudied by Handley (1959). The Mexican 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 fromMexican 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 fromMexican 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

abundant in the entrance area of Cueva del Abra,Tamaulipas. Peters' climbing rat, Tylomys nudicaudusgymnurus Villa R., has been reported from cavesin Guerrero and Puebla (Ramirez-Pulido and SanchezHernandez, 1971).Family DasyproctidaeRemains of the paca, Agouti paca nelsoni Goldman,have been found in caves in Yucatan (Pearse andKellogg, 1938; Jones et aI., 1974). Local inhabitantsreport that pacas frequently are shot in the large entrancesinks of the caves of Yucatan; they doubtlessventure into darkness in some caves in search ofwater.Family ErithizontidaeThe Mexican porcupine, Coendou mexicanus yucataniaeThomas, has been reported from caves in Yucatan(Pearse and Kellogg, 1938; Jones et aI., ]974).Order CarnivoraFamily CanidaeThe gray fox, Urocyon cinereoargenteus nigrirostris(Lichtenstein), was reported from Cueva de TiaJuana, Guerrero, by Villa R. (1967). It was presumablypreying on bats.Family FelidaeThe jaguar, Felis onca veraecrucis Nelson andGoldman, has been reported from Cueva de Los Sabinos(Hall and Kelson, 1959) and Sotano del Tigre,San Luis Potosi (Harris, 1971). Local hunters in theSierra de EI Abra of San Luis Potosi and Tamaulipasreport that jaguars are frequently seen in caves.Family MustelidaeThe western spotted skunk, Spilogale gracilis Merriam,was reported from Cueva del Tigre, Sonora(Villa R., 1958), where it was presumably preying onbats.Family ProcyonidaeThe coati, Nasua narica (Linnaeus), was reportedby Villa R. (1967) from a cave ncar lfuajintLin, MoreIos. This species is probably a frequent predator ofbats in the entrance area of caves.256

Literature CitedAnonymous. 1940. Expedicion para recoger peces ciegos enMexico. Ciencia, 1\I1'xico, 1:221.Anonymous. 1942a. Exploraciones biospcologicas en la regionde Valles (San Luis Potosi. i\lexico). Ciencia, Mexico,3:221.Anonymous. 1942b. Exploraciones biospeologicas en \uevoLeon (\Iexico). Ciencia. i\'lexico, 3:265.Anonymous. 1945. Notes on the blind cave tetra. Aquarium,Philadelphia, 14: 70.Anonymous, 1947. Expedicion cient ifica a Yucatan. Ciencia,Mexico, 8:128-129.Anonymous. 1965, Ai\ICS area report. Huautla. Assoc. :\'IexicanCave Stud. i\'ews., 1:60-61.Anonymous. 1968, l\ews: Baleones. Texas Caver, 13:11-12.Anonymous. 1974a, 1973 Belizean caving summarized.Inside Earlh, 3:7, 34,43.Anonymous. 1974b. News and not,:s, Assoc, i\lexiean CaveStud. "ews., 4:112,125.Anonymous. 1974,c, "ews and notes. Assoc, :VI ex ican CaveStud. i\'ews., 5:2-4.Anonymous. 1978. Cave notes. 1301. Asoc. ;Vlexieana Espeleol.,1:11-17.Absolon, K., and M. Kseneman. 1942. TrogJopedetini. VergleichendeStudie iiber eine altertiimliche hohlenbewohnendeKollembolengruppe aus den dinarischen Karstgebieten.Stud. Geb. Karstforsch, H, no, 16:1-57,Aguirre Pequeno, E. 1959. Aislamiento de Histoplasma cap·sulatum del guano de murcieJago en cuevas del noresle deMexico. Gac. Mcd. Mexieo, 89:243·253.Albert, D. P., and B. MacLeod. 1971. Caving in British Hon.duras. Natl. Speleol. Soc. News, 29:6-9.Alcorta Guerrero, R. 1966. Esquema geografico de Mexico.Pp. 1-8 in Caminos de Mexico, 2nd ed. Mexico, D. F.:Compania Hulera Euzkadi.Allee, W. C., A. E. Emerson, 0, Park, T. Park, and K. P,Schmidt. 1949. Principles of animal ecology. Philadel.phia: W. B, Saunders Co, xii + 837 pp.Allen, G. M. 1942. Hylonycteris underwoodi in Mexico. ].Mammal., 23:97.Allen, R. K. 1974. Neochoroterpes, a new subgenus of ChoroterpesEaton from North America (Ephemeroptera:Leptophlebiidae). Canadian Entomol., 106:161·168.Alley, K., R. Uinas, and D. E. Hillman. 1973. Neuronaland synaptic morphology in the optic tectum of a "blind"cavcfish. (Abstr.). Ana!. Rec., 175 :263.Alvarez, J, 1946. Revision del genero Anoptichthys con des·cripcion de una especie nueva (Pisc., Characidae). Anal.Esc. Nac. Cienc. BioI., 4:263·282.Alvarcz, .J. 1947. Descripeion de Anoptichthys hubbsi cara·cinido ciego de la Cueva de los Sabinos, S. L. P. Rev. Soc.Mexieana His!. Nat., 8:215·219.Alvarez, .J, 1950. Claves para la determinacion de especics enlos peces de las aguas continentales mexicanas. Mexico,Sec. Marina Direc. Gral. Pesca. 143 pp.Alvarez, .J. 1959. Nota preliminar sobre la ictiofauna delestado de San Luis Potosi. Acta Cient. Potosina, 3:71-88.Alvarez del Villar, J. 1970. Peces mexicanos (claves). lnst.Nac. Investigaciones BioI. Pesqueras, Ser. InvestigacionPesquera, Est., 1. 166 pp.Alvarez, T. 1963. The recent mammals of Tamaulipas,:\-Iexico. Univ. Kansas Publ., Mus. Nat. His!., 14: 363·473.Alvarez, T. 1968. Notas sobre una coleccion de mamiferos dela region costera del Rio Balsas entrt; Michoacan y Gue.rrero. Rev, Soc. Mexicana Hist. Nat., 29:21-35.Anderson, S. 1972. Mammals of Chihuahua. Taxonomy anddistribution. Bull. American Mus. Nat. Hist., 148: 149410.Anderson, S., and C. E. Nelson. 1965. A systematic revisionof Macrotus (Chiroptera). American Mus. oV., 2212.39 pp.Anderson, T. H. 1967. Geology of the middle third of LaDemocracia Quadrangle, Guatemala. M. A. Thesis. Austin:Univ. Texas at Austin. x + 81 pp., 1 pI.257

Andrews, E. W., IV. 1970. Balankanche, throne of the tigerpriest. Tulane Univ. Middle American Res. Inst. Pub!.,32. xi + 182 pp., phonograph record.Angel, F. 1954. Atlas des Poissons. IV. Poissons de eauxdouces. Especes exotiques & d'omement aquariophilie.3rd ed., rev. by]. Gery. Paris: Editions N. Boubee et Cie.181 pp., 12 pIsArgano, R. 1971. Cyathura sbordonii, nuova specie cavernicoladel Messico sudorientale. Diagnosi preliminare (Crustacea,Isopoda, Anthuridae). Fragmenta Entomol., 7:303305.Argano, R. 1972a. An asellid of the subterranean waters ofVeracruz, Mexico (Crustacea, Isopoda). Quad. Accad.Naz. Lincei, Probl. Att. Sci. Cult., 17(1 ):35-42.Argano, R. 1972b. On a troglobitic Cyathura from subterraneanwaters of Mexico (Crustacea, lsopoda). Quad.Accad. Naz. Lincei, Probl. Att. Sci. Cult., 171(1):23-34.Argano, R. 1974. 1Hexistenasellus magniezi n. sp., a blindaquatic isopod from Veracruz Mexico (Crustacea). Quad.Accad. Naz. Lincei, Probl. Att. Sci. Cult., 171(2):97-103.Argano, R. 1977. Asellota del Messico meridionale e Guatemala(Crustacea, Isopoda). Quad. Accad. Naz. Lincei,Probl. Att. Sci. Cult., 171(3):101-124.Arle, R. 1939. Collemboles nouveaux de Rio de Janeiro.Acad. Brasileira Cienc., Rio de Janeiro, Ann., 11 :25-32,pis. 1-4.Arnold, D. E., and B. F. Bohor. 1975. Attapulgite and Mayablue. Archaeology, 28:23-29.Ashworth, A. C. 1973. Fossil beetles from a fossil wood ratmidden in western Texas. Coleopterists Bull., 27:139-140.Atkinson, G., and P. Forsythe. 1979. Cueva de Xocotlat.Assoc. Mexican Cave Stud. Activities News., 10:76-81,map.Atkinson, J. 1978. Zoquitlan. Return to the river caves.Assoc. Mexican Cave Stud. Activities News., 8:46-50,map.Atz, J. W. 1950. Fishes from deserts and caves. Anim. King.,53:87-90.Avise,J. C., and G. B. Kitto. 1973. Phosphoglucose isomerasegene duplication in the bony fishes: An evolutionaryhistory. Biochem. Gen.,8:113.132.Avise, J. c., and R. K. Selander. 1972. Evolutionary geneticsof cave-dwelling fishes of the genus Astyanax. Evolution,26:1-19.Axelrod, H. R., C. Emmens, D. Sculthorpe, W. Vorderwinkler,R. Socolof, and N. Pronek. 1962. Exotic tropical fishes.New York: Sterling Publ. Co.Axelrod, H. R., and W. Vorderwinkler, eds. 1955. Colorguide to tropical fish. New York: Sterling Publ. Co. 160 pp.Bacescu, M., and T. Orghidan. 1971. Antromysis eubanican. sp. et Spelaeomysis nuniezi n. sp., mysis cavernicolesnouvelles de Cuba. Rev. Roumaine BioI., Zool., 16:225231.Baker, E. W. 1945. Five mites of the family Ereynetidae fromMexico. J. Washington Acad. Sci., 35: 16.19.Baker, E. W. 1949. A review of the mites of the family Cheyletidaein the United States National Museum. Proc.United States Natl. Mus., 99:267-320, pIs. 6-17.Baker, F. C. 1895. A naturalist in Mexico, being a visit toCuba, northern Yucatan and Mexico. Chicago: David Oli·phant. 145 pp., map.Baker, R. H. 1956. Mammals of Coahuila, Mexico. Univ.Kansas Publ., Mus. Nat. Hist., 9: 125-335.Baker, R. H. 1960. Mammals of the Guadiana Lava Field,Durango, Mexico. Pub!. Mus. Michigan State Univ., BioI.Ser., 1: 303-328.Baker, R. H., and J. K. Greer. 1960. Notes on Oaxacan mammals.J. Mammal., 41:413-415.Baker, R. H., and J. K. Greer. 1962. Mammals of the Mexicanstate of Durango. Pub!. Mus. Michigan State Univ., BioI.Ser., 2:25-154.Ball, I. R. 1976. Les caryotypes de trois planaires marinesnord-americaines: Contribution a la phylogenie et a laclassification du groupe (Platyhelminthes, Turbellaria,Tricladida). Canadian J. Zool., 54:644-651.Ball, I. R. 1977a. A new and primitive retrobursal planarianfrom Australian fresh waters (Platyhelminthes, Turbellaria,Tricladida). Bijdragen tot de Dierkunde, 47:149155.Ball, I. R. 1977b. On the phylogenetic classification of aquaticplanarians. Acta Zoo!' Fennica, 154:21-35.Balss, H. 1955. Decapoda, VI: Okologie. Pp. 1285-1367 inH. G. Bronn's Klassen und Ordnungen des Tierreichs,vo!. 5, pt. 1, book 7, no. 10.Banks, N. 1907. A catalogue of the Acarina, or mites, of theUnited States. Proc. United States Natl. Mus., 32:595625.Banks, N. 1938. A new myrmeleonid from Yucatan. CarnegieInst. Washington Pub!., 491 :235.Banner, A. H. 1953. On a new genus and species of mysidfrom southern Louisiana (Crustacea, Malacostraca). TulaneStud. Zool., 1:3-8.Banta, A. M. 1907. The fauna of Mayfield's Cave. CarnegieInst. Washington Publ., 67.114 pp.Barbour, T. 1945a. Cave hunting. Atlantic Monthly, 176(4):76-79.Barbour, T. 1945b. A naturalist in Cuba. Boston: Little,Brown and Co. 317 pp.Barbour, T., and 1. C. Cole. 1906. Vertebrata from Yucatan.4. Reptilia, Amphibia, and Pisces. Bull. Mus. Compo Zool.,50: 146-159, pis. 1-2.B[arlow], R. H. 1946. Cerro de San Lorenzo, Coahuila: Dossitios arquel6gicos. Rev. Mexicana Estud. Antropol.,8: 266.267, pI. 1.Barr, T. C., Jr. 1956. Note on cave blindfish. NetherworldNews, 4(6). Reprinted in: Speleo Digest, 1956(2):12-13.Barr, T. C., Jr. 1961. Caves of Tennessee. Tennessee Dept.Conservation, Commerce, Div. Geol. Bull., 64. 567 pp.,map.Barr, T. C., Jr. 1965. A new cavernicolous sphodrinc fromVeracruz, Mexico (Coleoptera: Carabidae). CoIeopteristsBull., 19:65-72.Barr, T. C., Jr. 1966a. New species of Mexisphodrus fromMexican caves (Coleoptera: Carabidae). Psyche, 73:112.115.Barr, T. C., Jr. 1966b. A progress report on study of the cavebeetles of Texas and Mexico. (Abstr.). Bull. Natl. Speleol.Soc., 28:97.Barr, T. C., Jr. 1967a.Antrojorceps, an eyeless cave scaritinefrom Mexico (Coleoptera: Carabidae). Coleopterists Bull.,21:65-70.Barr, T. C., Jr. 1967b. Ecological studies in the MammothCave System of Kentucky. I. The biota. Intcrnatl. J.Speleol., 3:147-204, pIs. 37-64.Barr, T. C., Jr. 1967c. Observations on the ecology of caves.American Nat., 101:475-492.Barr, T. C., Jr. 1967d. Three new cave trechines from Mexico(Coleopt.: Carab.). Ciencia, Mexico, 25 :161-166.258

Barr, T. c., Jr. 1968a. Cave ecology and the evolution oftroglobites. Evolutionary Biology, 2:35-102.Barr, T. C., Jr. 1968b. Mexican cave beetles of the familyCarabidae. Assoc. Mexican Cave Stud. News., 2:182-185.Barr, T. C., Jr. 1971. A new species of Mexaphaenops fromTamaulipas, Mexico (Coleoptera: Carabidae). Assoc.Mexican Cave Stud. Bull., 4:113-116.Barr, T. c., Jr. 1973. Speocolpodes, a new genus of troglobiticbeetles from Guatemala (Coleoptera: Carabidae).Psyche, 80:271-276.Barr, T. C., J r. 1974. Revision of Rhadine LeConte (Coleoptcra,Carabidae). I. The subterranea group. American Mus.Nov., 2539.30 pp.Barr, T. C., Jr., C. Bolivar Pieltain, and J. Hendrichs. 1968.Nota sinonimica sobre Agonum (Platynus) bilimekiBolivar y Hendrichs (Col., Carab.). Ciencia, Mexico,26:107-108.Barr, T. C., Jr., and R. A. Kuehne. 1971. Ecological studiesin the Mammoth Cave System of Kentucky. II. The ecosystem.Ann. Speleol., 26:47·96.Barr, T. C., Jr., and J. R. Reddell. 1967. The arthropod cavefauna of the Carlsbad Caverns Region, New Mexico.Southwestern Nat., 12:253-27:1.Barrera, A. 1951. Notas sobre sifonaptermi, II.-Deseripcion deAnomiopsyllus traubi nov. sp. (Siph., Hystrichops.).Ciencia, Mexico, 11: 197-200.Barrera, A. 1958. Insectos parasitos de mamiferos salvajes deOmiltemi, Gro., y descripcion de un nuevo Sifonaptero:Pleochaetis soberoni nov. sp. Anal. Esc. Nac. Cienc. BioI.,9:89-96.Barrera, A. 1962. La Peninsula de Yucatan como provinciabiotica. Rev. Soc. Mcxicana Hist. Nat., 23:7]-105.Barrera, A. 1968. La revista Ciencin y la obra entomologicadel Dr. Candido Bolivar y Pieltain. I{ev. Soc. MexicanaHist. Nat., 29:307-314.Bartholomew, R. 1973. Belize caving. Texas Caver, 18:259261.Bartsch, P. 1906. The urocoptid mollusks from the mainlandof America in the collection of the United States NationalMuseum. Proc. United States Natl. Mus., 31:109-160,pis. 3-5.Bateman, G. C., and T. A. Vaughan. 1974. Nightly activitiesof mormoopid bats. .T. :Vlammal., 55 :45-65.Bath, H. 1962. Vergleichende biologisch-anatomische Unlersuchungenuber die Leistungsfiihigkeit der SinnesorganefUr den Nahrungserwerb, ihre gegenseitige Abhiingigkeitund ihre Beziehungen zum Bau des Gehirns bei verschiedenenKnoehenfischarten. Z. Wiss. Zool., 167:238-290.Bauer, E. 1971. The mysterious world of caves. London:Collins Publ. 129 pp.Beck, L., and H. Schubart. 1968. Revision der Gattung Cryptoce/lusWestwood 1874 (Arachnida: Ricinulei). SenckenbergianaBioI., 49:67-78.Beier, M. 1956. Neue troglobionte Pseudoscorpione ausMexico. Ciencia, Mexico, 16:81-85.Beier, M. 1963. Eine neue Art der PseudoscorpionidenGattung Albiorix aus Hohle Acuitlapan, Gro., Mexico(Arach.). Ciencia, Mexico, 22:133·134.Beier, M. 1974. Ein neuer ParaliochtllOnius aus Guatemala.Rev. Suisse Zool., 81: 101-102.Benazzi, S. M. 1972. Notizie prelirninari sulle planarie raccoltenella seconda spedizione Lincea in Messico. Rend.Accad. Naz. Lincei, ser. 8, 52:403-405.Benazzi, S. M. 1973. Priority of the name Opisthobursa mexicanaBenazzi, 1972 over Dimarcus villalobosi Mitchell andKawakatsu, 1972. Rend. Aecad. Naz. Lineei, ser. 8, 54:533-536, pis. 1-2.Benazzi, S. M. 1976. Opisthobursa josephinae, a new troglobitieplanarian from Chiapas, Mexico. Rend. Acead. Naz.Lincei, ser. 8,59:533-536, pis. 1-2.Benazzi, S. M., and E. Giannini. 1974. A remarkable caveplanarian: Opisthobursa mexicana Benazzi, 1972. Quad.Acead. 'hz. Lincei, Probl. Att. Sci. Cult., 171(2):47-54,pis. 1-4.Benjamin, S. 1970. List of endangered foreign fish and wildlife.Fed. Reg., 35: 18319-18322.Benn, J. H. 1945. Composite observations on cave life (withspecial rderence to blind fishes). Bull. Natl. Speleol. Soc.,7:9·13.Bequaert, .T. C., and W. .T. Clench. 1933. The non-marinemollusks of Yucatan. Carnegie Inst. Washington Publ.,431:525-545, pI. 68.Bequaert, J. C., and W. J. Clench. 1936. A second contributionto the molluscan fauna of Yucatan. Carnegie Inst.Washington Publ., 457:61-75, pis. 1-2.Bequaert, J. C., and W. J. Clench. 1938. A third contributionto the molluscan fauna of Yucatan. Carnegie Inst.Washington Publ., 491 :257-260.Berra, T. 1963. A study of A noptichtys jordani. St. LouisUniv. Grotto News., 2:44-45. Reprinted in: SpeleoDigest, 1963(2):125-126.Berra, T. 1964. Obstacle avoidance in the blind Mexican cavefish, characin. St. Louis Univ. Grotto News., 3:12-19.Keprinted in: Speleo Digest, 1964(2):63-69.Bertin, L. 1958a. Eeologie. Traite de Zoologie, 13:18851933.Bertin, L. 1958b. Poissons cavernieoles. Traite de Zoologie,13: 2660-2662.Besharse, J. C., and R. A. Brandon. 1976. Effects of continuouslight and darkness on the eyes of the troglobiticsalamander Typhlotriton speweus. .J. Morphol., 149:527545.Bhatnagar, K. P. 1978. Breech presentation in the hairyleggedvampire, Diphylla ecaudata. J. Mammal., 59:864·866.Bilimek, D. 1867. Fauna der Grotte Cacahuamilpa in Mexiko.Verhandl. ZooI. Bot. Ges. Wein, 17:901-908.Birney, E. C.,.J. B. Bowles, R. M. Timm, and S. L. Williams.1974. Mammalian distributional records in Yucatan andQuintana Roo, with comments on reproduction, structure,and status of peninsular populations. Occ. PapersBell Mus. Nat. Hist., Univ. Minnesota, 13. 25 pp.Bittinger, C. 1972. Trip report: Spring 1971. Assoc. MexicanCave Stud. News., 3:56-57.Bittinger, C. 1973. Trips: November 24-December 2, 1972.Texas Caver, 18 :86.Bittinger, S. 1975. Sotano Hondo de Pinalito. Assoc. MexicanCave Stud. Activities News., 3:13·14.Black, .J. H. 1971. The cavc life of Oklahoma. A preliminarystudy (excluding Chiroptera). Oklahoma Underground,4:2-53.Bodenlos, A. .l- 1956. Notas sobre la geologia de la SierraMadre en la seccion Zimapan-Tamazunchale. Pp. 293-309in M. Maldonado-Koerdell, ed., Estratigrafia del Cenozoicoy del Mesozoico a 10 largo de la carretera entreReynosa, Tamps. y Mexico, D. F. Tectonica de la SierraMadre Oriental. Vulanismo en eJ Valle de Mexico. Congr.Geol. Internac., 20th Sesion, Excursiones A-14 y C-6.259

Bolivar y Pieltain, C. 1940. Exploracion de la Caverna deCacahuamilpa (Cuerrero, Mexico). Ciencia, Mexico, 1:125-126.Bolivar y Pieltain, C. ) 941. Estudio de un ricinulideo de laCaverna de Cacahuamilpa, Guerrero, Mex. (Arachnida).Rev. Soc. Mexicana Hist. Nat., 2:197-209, pl. 11.Bolivar y Picltain, C. 1942. Estudio del primer Trechinaeciego hallado en cavernas de Mexico (Col. Carab.). Ciencia,Mexico, 3:349-354.Bolivar y Pieltain, C. 1944. Descubrirniento de un Rhadineafenopsiano en el estado de l\uevo Leon, Mexico (Col.Carab.). Ciencia, Mexico, 5:25-28.Bolivar y Pieltain, C. 1946. Hallazgo de un nuevo rieinulideoen el Mexico central (Arach.). Ciencia, Mexico, 7:24-28.Bolivar y Pieltain, C. 1950. Estudio de una Cirolana cavernicolanueva de la region de Valles, San Luis Potosi, Mexico(Isop. Cirolanidae). Ciencia, Mexico, 10:211-218.Bolivar y Pieltain, C. 1952. Un Ozaeninae troglofilo de laCueva de los Sabinos, Valles, S. L. Potosi (Mexico) (Col.Carab.)_ Ciencia, Mexico, 11:295-296.Bolivar y Pieltain, C., and J. Carranza. 1954. Hallazgo enMexico de un pez ciego y depigmcntado de la familiaAmeiuridae, que vive en aguas freaticas del estado deCoahuila. Ciencia, Mexico, 14:115.Bolivar y Pieltain, c., and J. Hendrichs. 1964. Agoninoscavernicolas nuevos del genero Rhadine de Nuevo Leon,Coahuila y San Luis Potosi (Mexico). (Col. Carab.).Ciencia, Mexico, 23:5-J 6, pI. 1.Bolivar y Pieltain, C., and J. Hendrichs. 1965. Los Carahidaede la Gruta de Cacahuamilpa (Mexico) con descripcion deAgonum (Platynus) bilimeki n. sp. y de su larva (Ins.,Col.). Ciencia, Mexico, 23:225-232, pI. 5.Bonet, r. 1942. Notas sinonimicas sobre eI orden colembolos.Ciencia, Mexico, 3:56-59.Bonet, F. 1943. Sobrc la c1asificacion de los Oncopoduridae(Collembola), con descripcion de especies nuevas. Anal.Esc. Nac. Cienc. Biol., Mexico, 3:127-153.Bonet, F. 1944a. Sobre el genero Metasinella Denis, y algunosotros colemholos cavernicolas de Cuba. Ciencia, Mexico,5:17-24.Bonet, F. 1944b. Tullberginos de Mexico (Collembola). Rev.Soc. Mexicana '-list. Nat., 5 :51-72.Bonet, F. 1945. Nuevos generas y especies de hipogastruridosde Mexico (CoUembola). Rev. Soc. Mexicana Hist. Nat.,6:13-45, pIs. 2-6.Bonet, r. 1946a. Laboratorio de Zoologia. Bol. InformacionEsc. Nac. Cienc. BioI., \1exico, 4:105-117.Bonet, F. 1946b. Mas hipogastruridos anoftalmos de Mexico(CoUembola). Rev. Soc. Mexicana Hist. Nat., 7:51-62.Bonet, F. 1947. Monografia de la familia Neelidae (CoUemhola).Rev. Soc. Mcxicana Hist. Nat., 8:131-192.Bonet, F. 1953a. Cuevas de la Sierra Madre Oriental en laregion de Xilitla. Univ. Nae. Auton. Mexieo, Inst. Geol.,Bol., 57. vi + 96 pp., II pIs.Bonet, F. 1953b. Datos sabre las cavernas y otros fenomenoserosivos de las calizas de la Sierra de EI Abra. Mem.Congr. Cient. Mexicana, 5:238-273.Bonet, F. 1956a. Excursion A-14 (sentido norte-sur). ItinerarioCiudad Victoria, Tamps. Taninul. San Luis PotosI.Pp. 69-91 in M. MaJdonado-Koerdell, ed., Estratigrafia delCenozoico y del Mesozoico a 10 largo de la earreterraentre Reynosa, Tamps. y Mexico, D. F. Teetoniea de laSierra Madre Oriental. Vulcanismo en el VaJle de Mexico.Congr. Ceol. Intemac., 20th Sesion, Excursiones A-14 yC-6.Bonet, F. 1956b. Excursion C-6 (sentido sur-norte). ltine.rario Tamazunchale-Taninul. Pp. 217-240 iT! M. MaJdonado.Koerdell,ed., Estratigrafia del Cenozoico y delMesozoico a 10 largo de la carreterra entre Reynosa,Tamps. y Mexico, D. F. Tectonica de la Sierra MadreOriental. Vulcanismo en cl Valle de Mexico. Congr. Ceol.Internac., 20th Sesion, Excursiones A-14 y C-6.Bonet, F. 1956c. Excursion C-6 (sentido sur-norte). hinerarioTamazunchaJe·Taninul. Pp. 217·240 in M. MaJdo.nado-Koerdell, cd., Estratigrafia del Cenozoico y del\1esozoico a 10 largo dr la carreterra entre Reynosa,Tamps. y \Iexico, D. F. Tectonica de la Sierra J\'ladreOriental. Vulcanismo en el Valle de Mexico. Congr. Ceol.Internac., 20th Scsion, Excursiones A-14 y C-6.Bonet, F. 1956d. Excursion C-6 (senlido sur-norte). HinerarioTaninul, S. L. P.-Ciudad Victoria, Tamps. Pp. 241261 in M. Maldonado-Koerdell, ed., Estratigrafia delCcnozoieo y del J\lesozoico a \0 largo de la carreterraentre Reynosa, Tamps. y \-Iexico, D. F. Tectoniea de laSierra Madre Oriental. Vuleanismo en el Valle de Mexico.Congr. Ceol. Internac., 20th Sesion, Excursiones A-14 yCob.Bonet, F. 1963a. Road log. Ciudad Valles - Ciudad Victoria.\'Iay 26, 1963. Corpus Christi Geol. Soc. Ann. Field Trip,May 23·26, 1963, pp. 97-107.Bonet, F. 1963b. Road log. Taninul - TantobaJ' May 25,1963. Corpus Christi Ceo!. Soc. Ann. Field Trip, May 2326, 1963, pp. 91-96.Bonet, F. 1971. Espelcologia de la region de Cacahuamilpa,Gro. LIniv. Nae. Auton. Mexico, Inst. Ceol., Bol., 90: 1-98,pis. 1-26.Bonet, r., and C. Tellez. 1947. Un nuevo genero de esmin·turidos (Collembola). Rev. Soc. Mexicana Hist. Nat.,8:) 93-203, pIs. 22-23.Bonis, S. H. 1969. A summary of the geology of Guatemala.Publ. Geol. Inst. Centroamericano Investigacion y Teeno!.IndustriaJ,2:76-80.Boon, M. 1969. Caving in Mexico. Canadian Caver, no.1 :32-40.Boon, M. 1974. Cruz Pilal again. Canadian Caver, 6(2):11-12_Boon, M. 1975. Return to Yoehih-Part 2. Canadian Caver,7(1 ):6.11,15-16.Botosaneanu, L., and L. B. Holthuis. 1970. Subterraneanshrimps from Cuba (Crustacea Deeapoda Natantia). Trav.Inst. Speol. "Emile Racovitza", 9:121-133.Bott, R. 1969. Die Siisswasserkrahben Siid-Amerikas und ihreStammesgeschichte. Eine Revision der TrichodactyIidaeund der Pseudothelphusidae ostlieh der Anden (Crustacea,Deeapoda). Abhandl. Senckenbergischen Naturforsch.Ges., 518: 1-94.Boucquey, C., G. Thines, and C. Van dcr Borght. 1965.Etude comparative de la capacite photopathique et deI'activite chez Ie poisson cavernieole Anoptichthys antrobius,chez la forme epigce ancestrale Astyanax mexicanus,et chez les hybrides F 1 (Astyanax X Anoptichthys) et r 2'Pp. 79-103 in La distribution temporeUc des activites animaleset humaines. Paris: Masson et Cie.Bousfield, E. L. 1977. A new look at the systematies of gammaroideanamphipods of the world. Crustaceana Suppl.,4:282-316.Bowman, T. E. 1964. Antrolana lira, a new genus and speciesof troglobitic cirolanid isopod from Madison Cave, Vir!/;inia. Internatl. J. Speleol., 1:229-236, pIs. 50-57.260

Bowman, T. E. 1965. Xilitloniscus, a new genus for the Mexi·can troglobitic isopod, Cordioniscus laevis Rioja (Oniscoidea:Trichoniscidae). Proc. BioI. Soc. Washington,78:209-215.Bowman, T. E. 1973. Two new American species of Spelaeomysis(Crustacea: Mysidacea) from a Mexican cave andland crab burrows. Assoc. Mexican Cave Stud. Bull.,5:13-20.Bowman, T. E. 1975. A new genus and species of troglobiticcirolanid isopod from San Luis Potosi, Mexico. Occ.Papers Mus. Texas Tech Univ., 27. 7 pp.Bowman, T. E. 1976. Three new troglobitic asellids fromwestern North America (Crustacea: Isopoda: Asellidae).Internatl. J. Speleol., 7:339-356.Bowman, T. E. 1977a: Mysidacea. Pp. 149-150 in S. H. Hurlbert,ed., Biota acquatica de Sudamerica Austral. SanDiego, California: San Diego State Univ.Bowman, T. E. 1977b. A review of the genus Antromysis(Crustacea: Mysidacea), including new species fromJamaica and Oaxaca, Mexico, and a redescription andnew records for A. cenotensis. Assoc. Mexican CaveStud. Bull., 6:27-38.Bowman, T. E., R. Prins, and B. F. Morris. 1968. Notes onthe harpacticoid copepods Attheyella pilosa and A. caralineTlsis,associates of crayfishes in the eastern UnitedStates. Proc. BioI. Soc. Washington, 81:571-585.Boyd, A. 1966. Geology of the western third of La DemocraciaQuadrangle, Guatemala. M. A. Thesis. Austin: Univ.Texas at Austin. ix + 79 pp., 1 pI.Bradshaw, G. V. R., and B. Hayward. 1960. Mammal skullsrecovered from owl pellets in Sonora, Mexico. 1. Mammal.,41 :282-283.Brand, D. D. 1937. The natural landscape of northwesternChihuahua. Univ. i\ew Mexico Bull., Geol. Ser., 5(2).74 pp., 10 pis.Bravo-Hollis, M., and J. Caballero Deloya. 1973. Catalogo dela collecion helmintologica del Instituto de Biologia.Univ. Auton. Mexico, Inst. BioI., Publ. Especiales, 2.138 pp.Breder, C. M., Jr. 1942. Descriptive ecology of La CuevaChica, with especial reference to the blind fish, Anoptichthys.Zoologica, New York, 27:7·15, pis. 1-3.Breder, C. M., Jr. 1943a. Apparent changes in phenotypicratios of the characins at the type locality of AnoptichthysjordaniHubbs and Innes. Copeia, 1943:26-30.Breder, C. M., J r. 1943b. A note on erratic viciousness inAstyaTlax mexicanus (Phillipi). Copeia, 1943:82-84.Breder, C. M., Jr. 1943c. Problems in the behavior andevolution of a species of blind cave fish. Trans. New YorkAcad. Sci.,ser. 2, 5:168-176.Breder, C. M., Jr. 1944. Ocular anatomy and light sensitivitystudies on the blind fish from Cueva de los Sabinos,Mexico. Zoologica, New York, 29:131-144, pI. 1.Breder, C. M., Jr. 1945. Compensating reactions to the lossof the lower jaw in a cave fish. Zoologica, New York,30:95-100, pI. 1.Breder, C. M., Jr. 1953. Cave fish evolution. Evolution, 7:179-181.Breder, C. M., Jr. 1954. A second case of survival by a teleostwithout a lower jaw. Zoologica, New York, 39:13-16,pI. 1.Breder, C. M., Jr. 1959. Studies on social groupings in fishes.Bull. American Mus. Nat. Hist., 117:393.481, pis. 70-80.Breder, C. M., .Jr., and E. H. Atz. 1952. Conditioned restrictionsof movement in fishes, fancied and real. Copeia,1952:261-265.I:Ireder, C. M., Jr., and E. B. Gresser. 1941a. Behavior ofMexican cave characins in reference to light and caveentry. (Abstr.). Anat. Rec., Suppl., 81:112.Breder, C. M., Jr., and E. B. Gresser. 1941b. Correlations betweenstructural eye defects and behavior in the Mexicanblind characin. Zoologica, New York, 26: 123-131, pis.1-4.Breder, C. M., Jr., and E. B. Gresser. 1941c. Further studieson the light sensitivity and behavior of the Mexican blindcharacin. Zoologica, New York, 26:289-296, pI. 1.Breder, C. M., .Jr., and P. Rasquin. 1943. Chemical sensoryreactions in the Mexican blind characins. Zoologica, NewYork, 28:169-200, pIs. 1-3.Breder, C. M., Jr., and P. Rasquin. 1947a. Comparative studiesin the light sensitivity of blind characins from a seriesof Mexican caves. Bull. American Mus. Nat. Hist., 89:319351.Breder, C. M., Jr., and P. Rasquin. 1947b. Evidence for thelack of a growth principle in the optic cyst of Mexicancave fish. Zoologica, New York, 32:29-33.Breder, C. M., Jr., and D. E. Rosen. 1966. Modes of reproductionin fishes. Garden City, New York: Natural HistoryPress. 951 pp.Bridgemon, R. 1974. Mollusks found in caves near Tlamaya,San Luis Potosi. Assoc. Mexican Cave Stud. News., 4:141,143.Bridges, W. 1940. The blind fish of La Cueva Chica. Bull.New York Zool. Soc., 43:74-97, map.Bridges, W. 1943. What we .have learned about blind cavefish. Anim. King., 46:82, 87-90.Bridges, W. 1955. No eyes in the darkness. Pp. 256-268 inC. E. Mohr and H. N. Sloane, eds., Celebrated Americancaves. New Brunswick, New Jersey: Rugers Univ. Press.Brignoli, P. M. 1972. Some cavernicolous spiders fromMexico (Araneae). Quad. Accad. Naz. Lincei, Probl. Alt.Sci. Cult., 171(l ):129-155.Brignoli, P. M. 1973a. Note sulla morfologia dei genitali deglischizomidi e diagnosi preliminari di due nuove specie delMessico (Arachnida, Schizomida). Fragmenta Entomol.,9:1-9.Brignoli, P. M. 1973b. II popolamento di ragni nelle grottetropicali (Araneae). Internatl. J. Speleol., 5: 325-336.Brignoli, P. M. 1974a. A contribution to the knowledge ofthe Schizomida of Mexico and Guatemala (Arachnida,Schizomida). Quad. Accad. Naz. Lincei, Probl. Alt. Sci.Cult., 171(2):143-152.Brignoli, P.M. 1974b. Notes on spiders, mainly cave-dwelling,of southern Mexico and Guatemala (Araneae). Quad.Aecad. Naz. Lincei, Probl. Alt. Sci. Cult., 171(2):195238.Brignoli, P. M. 1974c. On some Ricinulei of Mexico withnotes of the morphology of the female genital apparatus(Arachnida, Ricinulei). Quad. Accad. Naz. Lincei, Probl.Alt. Sci. Cult., 171(2):153-174.Brignoli, P. M. 1976. Beitriige zur Kenntnis der Scytodidae(Araneae). Rev. Suisse Zool., 83:125-191.Brignoli, P. M. 1977. Spiders from Mexico. III. A new leptonetidfrom Oaxaca (Araneae, Leptonetidae). Quad. Accad.Naz. Lincei, Probl. Alt. Sci. Cult., 171(3):213-218.Brignoli, P. M. 1978. Spinnen aus Brasilien. II. Vier neueOchyroceratidae aus Amazonas nebst Bemerkungen liberandere Amerikanische Arten (Arachnida: Araneae). Stud.Neotrop. Fauna & Environ., 13:11-21.261

Brignoli, P. M. 1979a. The morphology and the relationshipsof the Leptonetidae (Arachnida, Araneae). 1. Arachnol.,7:231-236.Brignoli, P. M. 1979b. On some cave spiders from Guatemalaand United States (Araneae). Rev. Suisse Zool., 86:435443.Brignoli, P. M. 1979c. Ragni delle Filippine III. Su alcuniOchyroceratidae (Araneae)_ Rev. Suisse Zool., 86:595604.Brittan, M. R., and 1. E. Bohlke. 1965. A new blind characidfish from southeastern Brazil. Notulae Naturae, 380. 4 pp.Broughton, P. L. 1973. Exploration of Cueva de Agua Escondida,Huehuetenango, Guatemala. Canadian Caver, 5(1):52-59.Broughton, P., and M. Boon. 1975. The Rio Huista karst cavesurvey: Western Guatemala. Expl. 1.,53(1):8-15.Broussard, D. 1975. Taneoyol, Qro. Assoc. Mexican CaveStud. Activities News., 1:2-3.Brown, W. S. 1974. Ecology of thc aquatic box turtle, TerrapenecoahlJila (Chelonia, Emydidae) in northern Mexico.Bull. Florida State Mus., BioI. Sci., 19:1-67.Bucher!, W. 1959. Kritische Untersuchungen der NewportiaArten (Chilopoda, Scolopendromorpha, Cryptopidae).Beitr. Neotrop. Fauna, 1 :229-242.Bull, E. G. 1969. Relative humidity and temperature responsesin two troglobitic millipedes-Cambala speobiaand SpeodesmlJs bicornolJrtls. M. S. Thesis. Lubbock:Texas Tech. CoIl. 54 pp.Burgers, A. C.l.,P. 1. H. Bennink, and G. 1. van Oordt. 1963.Investigations into the regulation of the pigmentary systemof the blind Mexican cave fish, Anoptichthys jordani.Proc. K. Nederlandse Akad. Wet. Amsterdam, ser. C, 66:189·195, pI.Burns, R. J. 1970. Twin vampire bats born in captivity. 1.Mammal., 51:391-392.Burt, W. H. 1938. Faunal relationships and geographic distributionof mammals in Sonora, Mexico. Misc. Publ. Mus.Zool., Univ.Miehigan, 39:5-77.Burt, W. H., and E. T. Hooper. 1941. Notes on mammalsfrom Sonora and Chihuahua, Mexico. Occ. Papers Mus.Zool., Univ. Michigan, 430.7 pp.Burukovskii, R. N. 1974. Keys to the shrimps, spiny lobstersand homarids. Moskow: Pischevaya promyshiennost,"Ford Industry." 126 pp.Bustamante, M. E. 1964. Symposium sobre histoplasmosispulmonar primaria. II. Epidemiologia. Gac. Med. Mexico,94:509-518.Butterlin, 1., and F. Bonet. 1963. Mapas geologicos de laPeninsula de Yucatan. I.-Las formaciones Cenozoicas dela parte mexicana de la Peninsula de Yucatan. IngenieriaHidraulica en Mexico, 17(1):63-71, map.Byrd, T. 1976. Christmas in Acatlan. Assoc. Mexican CaveStud. Activities News., 4:23-24.Caballero y C., E. 1942. Deseripcion de la segunda especie deCapillaria encontrada en los murcielagos de America delNorte. III (Nematoda: Trichuridae). Anal. Inst. BioI.,Mexico, 13:tM,9-654.Caballero y C., E. 1943. Algunas especies de trematodos delos murcielagos de la region de Izucar de Matamoros, Pue.V. Anal. Inst. BioI., Mexico, 14:423-430.Cahn, P. H. 1958. Comparative optic development in Astya·nax mexicanlJs and in two of its blind cave derivatives.Bull. American Mus. Nat. Hist., 115:69-112, pis. 11-27.Caine, E. A. 1974. Zoogeography of the Floridian troglobiticcrayfishes, genus ProcambarlJs. American MidI. Nat., 92:487-492.Campos, H. 1969. Die Geschmacksknospen im Vorderdarmvon Susswasserfischen, Zahl, Verteilung und Entwicklung(PhoxinlJs phoxinlJs L., GasterostelJs aClJleatIJs L., HemigrammlJscalJdovittatlJs Ahl, Anoptichthys jordani Hubbset Innes und Salmo gairdneri Rich.). Z. wiss. Zool., 179:253·299.Cardenas Figueroa, M. 1950. Los recursos naturales de Yucatan.IV .-Informe hidrobiologico y faunistico de Yucatan.Bol. Soc. Mexicana Geogr. Estadist., 69:135-159.Carranza, 1. 1954. Descripcion del primer bagre anoftalmo ydepigmentado encontrado en aguas mexicanas. Ciencia,Mexico, 14:129-136, pI. 1.Carrasco, B. 1970. La Formacion EI Abra (Formacion EIDoctor) en La Plataforma Valles-San Luis Potosi. Rev.Inst. Mexieano Petrol., 2(3):97-99.Carreno, A. de la O. 1950. Los reeursos naturales de Yucatan.Preambulo. Bol. Soc. Mexicana Geogr. Estadist., 69:21·26.Carreno, A. de la O. 1951. Las provincias geohidrologicas deMexico (primera parte). Univ. Nac. Aut. Mexico, Inst.Geol., Bol., 56(1). xiii + 98 pp., 19 figs., 2 maps, 41 tables.Carreno, A. de la O. 1954. Las provineias geohidrologicas deMexico (segunda parte). Univ. Nac. Aut. Mexico, lnst.Gcol., Bol., 56(2). viii + 166 pp., 8 pis., 2 figs., 5 tables.Carter, D. C., and 1. K. Jones, 1r. 1978. Bats from the Mexicanstate of Hidalgo. Occ. Papers Mus. Texas Tech Univ.,54.12 pp.Cassagnau, P., and C. Delamare Deboutteville. 1953. LesArrhopalites et Pararrhopalites d'Europe [Collembolessymphyleones cavernicolesl. Notes Biospeol., 8:133-147,pI. 1.Causey, N. B. 1959. Two new troglodytic millipeds fromTexas. Proe. BioI. Soc. Washington, 72:69-73.Causey, N. B. 1960. Millipeds collected in Guatemalan caves,including Calymmodesmus inquinatus, n. sp. (Stylodesmidae:Polydesmida). Proc. BioI. Soc. Washington, 73:275-279.Causey, N. B. 1963. Mexiterpes sabinus, new genus and newspecies, a Mexican troglobite (Diplopoda: Trichopetali.dae). Psyche, 70:235~239.Causey, N. B. 1964a. New cavernieolous millipeds of thefamily Cambalidae (Cambalidea: Spirostreptida) fromTexas (U. S. A.) and Mexico. Internatl. J. Speleol., 1:237246, pIs. 58-59.Causey, N. B. 1964b. Two new troglobitic millipeds of thegenus Glomeroides from Mexico (G lomeridae: Glomerida).Proc. Louisiana Acad. Sci., 27:63-66.Causey, N. B. 1965. The millipeds in the caves of Texas andadjacent states. (Abstr.). Bull. Natl. Speleol. Soc., 27:63.Causey, N. B. 1968. The millipedes in the caves of Mexicoand Guatemala. Assoc. Mexican Cave Stud. News., 2:124125.Causey, N. B. 1969. New trichopetalid (Chordeumidea: Chordeumida)millipedes from caves in North and CentralAmerica. Proc. Louisiana Acad. Sci., 32:43-49.Causey, N. B. 1971a. The Cambalidae in Mexican caves, withdescriptions of three new species of Mexicambala (Diplopoda:Cambalida). Proc. BioI. Soc. Washington, 84:271281.Causey, N. B. 1971b. Millipedes in the collection of the Associationfor Mexican Cave Studies (Diplopoda). Assoc.Mexican Cave Stud. Bull., 4:23-32.Causey, N. B. 1973. Millipedes in the collection of the Associationfor Mexican Cave Studies. II. Keys and additional262

ecords and descriptions (Diplopoda). Assoc. MexicanCave Stud. Bull., 5 :107-122.Causey, N. B. 1975a. Desert millipedes (Spirostreptidae:Spirostreptida) of the southwestern United States andadjacent Mexico. Occ. Papers Mus. Texas Tech Univ., 35.12 pp.Causey, N. B. 1975b. Millipedes in the collection of theAMCS. HI. Reddellobus troglobius, n. gen., n. sp., an unusualtroglobite from Puebla, Mexico, and other recordsof the family Spirobolellidae (Order Spirobolida, ClassDiplopoda). lnternatl. J. Speleol., 6:333-338.Causey, N. B. 1977. Millipedes in the collection of the Associationfor Mexican Cave Studies IV. New records anddescriptions chiefly from the northern Yucatan Peninsula,Mexico (Diplopoda). Assoc. Mexican Cave Stud. Bull.,6:167-183.Chabert, C. 1973. Jalons mexicains. Grottes et Gouffres.Bull. periodique du Speleo-Club de Paris, 50: 15-26.Chace, F. A., Jr. 1942. A new cave shrimp from Cuba. Proc.New England Zool. Club, 19:99-102, pI. 29.Chace, F. A., J r. 1943. Two new blind prawns from Cubawith a synopsis of the subterranean Caridea of America.Proc. New England ZooI. Club, 22:25.40, pIs. 5-7.Chace, F. A., Jr. 1954. Two new subterranean shrimps (Deca.poda: Caridea) from Florida and the West Indies, with arevised key to the American species. J. Washington Acad.Sci., 44:318-324.Chace, F. A., J r. 1972. The shrimps of the SmithsonianBredin Caribbean Expeditions with a summary of theWest Indian shallow-water species (Crustacea: Decapoda:Natantia). Smithsonian Contr. Zool., 98. 179 pp.Chace, F. A., Jr., and H. H. Hobbs,Jr. 1969. The freshwaterand terrestrial decapod crustaceans of the West Indieswith special reference to Dominica. Bull. United StatesNatl. Mus., 292. 258 pp.Chace, F. A., Jr., and R. B. Manning. 1972. Two new Carideanshrimps, one representing a new family, from marinepools on Ascension Island (Crustacea: Decapoda: Natantia).Smithsonian Contr. Zool., 131. 18 pp.Chakraborty, R., and M. Nei. 1974. Dynamics of gene differentiationbetween incompletely isolated populations ofunequal sizes. Theor. Popul. BioI., 5:460-469.Chamberlin, J. C. 1938. A new genus and three new speciesof false scorpions from Yucatan caves (ArachnidaChelonethida). Carnegie lnst. Washington Publ., 491:109121.Chamberlin,.J. C. 1946. The genera and species of the Hyidae.A family of the arachnid order Chelonethida. Bull. Univ.Utah, BioI. Ser., 9(6).16 pp.Chamberlin, J. C. 1947. The Vachoniidae-A new family offalse scorpions. Two new species from caves in Yucatan.Bull. Univ. Utah, BioI. Ser., 10(4). 15 pp.Chamberlin, J. C., and R. V. Chamberlin. 1945. The generaand species of the Tridenchthoniidae (Dithidae). A familyof the arachnid order Chelonethida. Bull. Univ. Utah,BioI. Ser., 9(2). 67 pp.Chamberlin, J. C., and D. R. Malcolm. 1960. The occurrenceof false scorpions in caves with special reference to cavernicolousadaptation and to cave species in the NorthAmerican fauna (Arachnida-Chelonethida). AmericanMidI. Nat., 64: 105-115.Chamberlin, R. V. 1938. Diplopoda from Yucatan. Carnegielnst. Washington Publ., 491 :165-182.Chamberlin, R. V. 1941. New chilopods from Mexico. PanPacific Entomol., 17:184-188.Chamberlin, R. V. 1942. On centipeds and millipeds fromMexican caves. Bull. Univ. Utah, BioI. Ser., 7(2).19 pp.Chamberlin, R. V. 1943a. On Mexican centipeds. Bull. Univ.Utah, BioI. Ser., 7(3). 55 pp.Chamberlin, R. V. 1943b. On Mexican millipeds. Bull. Univ.Utah, BioI. Ser., 8(3).103 pp.Chamberlin, R. V. 1947. Some records and descriptions ofdiplopods chiefly in the collection of the Academy. Proc.Acad. Nat. Sci. Philadelphia, 99:21-58.Chamberlin, R. V. 1952a. Eclomus nom. nov. (Diplopoda).Entomol. News, 63:71.Chamberlin, R. V. 1952b. Three cave-dwelling millipeds.Entomol. News, 63:10-12.Chamberlin, R. V., and R. L. Hoffmann. 1958. Checklist ofthe millipeds of North America. United States Natl. Mus.Bull., 212:1·236.Chamberlin, R. V., and W. lvie. 1938a. Arachnida of theorders Pedipalpida, Scorpionida and Ricinulida. Carnegielnst. Washington Publ., 491:101-107.Chamberlin, R. V., and W. Ivie. 1938b. Araneida from Yuca·tan. Carnegie lnst. Washington Publ., 491:123.136.Chappuis, P. A. 1927. Die Tierwelt der unterirdischen Gewasser.Die Binnengewiisser, 3. 175 pp.Chtwood, B. G. 1938. Some nematodes from the caves ofYucatan. Carnegie lnst. Washington Publ., 491:51-66.Chopard, L. 1947. Note sur les orthopteres cavernicoles duMexique. Ciencia, Mexico, 8:67-70.Chopard, L. 1968. Gryllides. Fam. Gryllidae: Subfam. Mogoplistinae,Myrmecophilinae, Scleropterinae, Cachoplistinae,Pteroplistinae, Pentacentrinae, Phalangopsinae, Trigonidiinae,Eneopterinae; Fam. Oecanthidae, Gryllotalpidae.Orthopterorum Catalogus, 12:215·500.Christiansen, K. 1973. The genus Pseudosinella in Mesoamericancaves. Assoc. Mexican Cave Stud. Bull., 5:129134.Clemons, R. E., and D. F. McLeroy. 1966. Hoya Torreonl3R-l(l) con Resumen de la geologia de la Hoya Torreon,Estados de Coahuila y Durango. Univ. Nac. Aut. Mexico,lnst. Geol., Carta Geologica de Mexico, Serie de 1:100,000,no. 12.Cockrum, E. L. 1969. Migration in the guano bat, Tadaridabrasiliensis. Univ. Kansas Mus. Nat. Hist. Misc. Publ.,51:303-336.Cockrum, E. L., and G. van R. Bradshaw. 1963. Notes onmammals from Sonora, Mexico. American Mus. Nov.,2138.9 pp.Cohen, D. M., and C. R. Robins. 1970. A new ophidioid fish(genus Lucifuga) from a limestone sink, New ProvidenceIsland, Bahamas. Proc. Bio!. Soc. Washington, 83:133-144.Cole, G. A., and W. L. Minckley. 1966. Speocirolana thermydronis,a new species of cirolanid isopod crustacean fromcentral Coahuila, Mexico. Tulane Stud. Zool., 13:17-22.Cole, G. A., and W. L. Minckley. 1970. Sphaerolana, a newgenus of cirolanid isopod from northern Mexico, with de.scription of two new species. Southwestern Nat., 15:71-81.Cole, G. A., and W. L. Minckley. 1972. Stenasellid isopodcrustaceans in the Western Hemisphere-a new genusand species from Mexico-with a review of other NorthAmerican freshwater isopod genera. Proc. BioI. Soc. Washington,84: 313-326.Conde, B. 1949. Campodeides cavernicoles de la region desAppalaches. Notes Biospeo!., 4:125-137.Conde, B. 1955. Materiaux pour une monographie des diplourescampodCides. Mem. Mus. Nat!. Hist. Nat., n. ser.,ser. A, Zool., 12: 1·202.263

Conde, B. 1975. Description du premier campodeide cavernicoledu Guatemala. Rev. Suisse Zool., 82:421-424.Constantine, D. G. 1966. New bat locality records from Oaxaca,Arizona and Colorado. J. Mammal., 47: 125-126.Constantine, D. G. 1967. Activity patterns of the Mexicanfree-tailed bat. Univ. New Mexico Pub\. BioI., 7.79 pp.Constantine, D. G. 1970. Bats in relation to the health, welfareand economy of man. Pp. 319-449 in W. A. Wimsatt,ed., Biology of bats, Vol. n. New York: Academic Press.Contreras-Balderas, S. 1969_ Perspectivas de la ictiofauna enlas zonas aridas del norte de Mexico. Internatl. CenterArid Semi-Arid Land Stud. Publ., 3:293-304.Cooke, J. A. L., and M. U. Shadab. 1973. New and littleknown ricinuleids of the genus Cryptocellus (Arachnida,Ricinulei). American Mus. Nov., 2530.25 pp.Cooke, J. W. 1971. Mating behavior and the functionalmorphology of the male copulatory apparatus in Cryptocelluspelaezi (Arachnida, Ricinulei). M. S. Thesis. Lubbock:Texas Tech Univ. 50 pp.Coons, D. 1974. Trip report: February 1974. Assoc. MexicanCave Stud. News., 5:15,17.Coons, D. 1976. The river caves. Canadian Caver, 8(1):34-41.Coons, D. 1977. Cueva de la Laguna Verde, Oaxaca. CanadianCaver, 8(2):18-21.Cope, E. D. 18b5. Third contribution to the herpetology oftropical America. Proc. Acad. Nat. Sci. Philadelphia,17:185-198.Copp, LJ. H. 1969. The ultimobranchial glands and calciumregulation. Pp. 377-398 in W. S. Hoar and D. J. Randall,eds., Fish physiology, volume II: The endocrine system_New York: Academic Press.Coronado Gutierrez, L. 1970. Estudio de un Cryptocellusde cavernas de Mexico (Arachn., Ricin.). Ciencia, Mexico,27:47-62.Coss, R. G. 1979. Delayed plasticity of an instinct: Recognitionand avoidance of 2 facing eyes by the jewel fish.Develop. Psychobiol., 12:335-345.Coss, R. G., and A. Globus. 1979. Social experience affectsthe development of dendritic spines and branches on tee·tal interneurons in the jewel fish. Develop. Psychobiol.,12:347-358.Cottarelli, V., and R. Argano. 1977. Trichodactylus (Rodriguezia)mensabak n. sp. (Crustacea, Decapoda, Brachyura),granchio cieco delle acque sotterranee del Chiapas (Messico).Quad. Accad. Naz. Lincei, Probl. Att. Sci. Cult.,171(3):207-212.Contreras V., H. 1956. Resefia de la geologia del surestede Mexico. Pp. 39-122 in M. Maldonado-Koerdell,ed., Geologia general de la Sierra Madre Oriental entreMexico, D. F. y Cordoba, Ver. Depositos continentalesy volcamcos del Cenozoico Superior y sedimentosmarinos del Mesozoico y Cenozoico. Campos petro-leros de la Cuenca de Veracruz. Obras hidraulicas delRio Papaloapan. Campos petroleros y azufreros delIstrno de Tehuantepec. Geomorfologia de la Peninsulade Yucatan. Visitas a las zonas arqueologicasMayas. Congr. Geol. Internac., 20th Sesion, ExcursionC-7.Contreras V., H. 1958. Resumen de la geologia de la partemedia del estado de Tahasco y el norte del estado de Chiapas.Bol. Asoc. Mexicana Geol. Petrol., 10:193-210,fig. 1-7.Courbon, P., and D. Dreux. 1976. Candelaria. Etude dureseau hydrospeleologique de Candelaria, Alta Verapaz.Spelunca, 1976, no. 3, suppl., pp. 12-17.264Crabill, R. E., Jr. 1960. A new Nuevobius, with review of thegenus (Chilopoda: Lithobiomorpha: Lithobiidae). Bull.Brooklyn Entomol. Soc., 55 :121-133.Craig, 1. L. 1975. A checklist of the invertebrate speciesrecorded from Missouri subterranean habitats. MissouriSpeleol., 15(2). 10 pp.Craig, 1. L. 1977. Invertebrate faunas of caves to be inundatedby the Meramec Park Lake in eastern Missouri. Nat!.Speleol. Soc. Bull., 39:80.g9.Creaser, E. P. 1936. Crustaceans from Yucatan. Carnegie Inst.Washington Publ., 457:117-132.Creaser, E. P. 1938. Larger cave Crustacea of the YucatanPeninsula. Carnegie Inst. Washington Publ., 491:159-164.Croizat, L., G. Nelson, and D. E. Rosen. 1974. Centers oforigin and related concepts. Syst. Zool., 23:265-287.Cserna, E. G., and A. Bello-Barradas. 1963. Geologia de laparte central de la Sierra de Alvarez, Municipio de Zaragoza,estado de San Luis PotosI. Univ. Nac. Aut. Mexico,Inst. Geol., Bol., 71:23-62,12 pis.Culver, D. C., and T. L. Poulson. 1971. Oxygen consumptionand activity in elosely related amphipod populations fromcave and surface habitats. American MidI. Nat., 85:74.g4.Cunliffe, F. 1952. Biology of the cockroach parasite, Pimelia·philus podapolipophagus Tragardh, with a discussion ofthe genera Pimeliaphilus and Hirstiella (Acarina, Pterygosomidae).Proc. Entomol. Soc. Washington, 54:153·169.Cuvier, M. Ie B., and M. A. Valenciennes. 1846. Histoirenaturelle des poissons, vol. 18. Paris: P. Bertrand. 505 pp.Dalquest, W. W. 1953a. Mammals of the Mexican state of SanLuis Potosi. Louisiana State Univ. Stud., BioI. Sci. Ser.,1:1-229.Dalquest, W. W. 1953b. Mexican bats of the genus Artibeus.Proc. BioI. Soc. Washington, 66:61-65.Dalquest, W. W., and E. R. Hall. 1949. Five bats new to theknown fauna of Mexico. J. Mammal., 30:424-427.Dalquest, W. W., and E. Roth. 1970. Late Pleistocene mammalsfrom a cave in Tamaulipas, Mexico. SouthwesternNat., 15:217·230.Dancau, D., and E. Serban. 1965. La presence de Bogidiellaalbertimagni Hertzog 1 ~Jj3 en Roumanie et quelquesremarques sur les especes europeennes du genre. Internatl.J. Speleol., 1:339-348, pis. 71-75.Danielpol, D. L. 1977. Recherches sur les Ostracodes Entocytheridae.Donnees sur Sphaeromicola cebennica juberthieinov. ssp. et Sphaeromicola cirolanae Rioja. Internatl.J. Speleol., 9:21-41.Darlington, P. J. 1936. Aquatic Coleoptera from Yucatan.Carnegie Inst. Washington Publ., 457:153-155.Davis, G. H. 1966. Geology of the eastern third of La DemocraciaQuadrangle, Guatemala. M. A. Thesis. Austin: Dniv.Texas at Austin. ix + 78 pp., 2 pIs.Davis, N. W. 1974. A preliminary report on the ZacapoaxtlaCuetzalan Area, northern Puebla, Mexico. Assoc. MexicanCave Stud. News., 4:153-184, pIs. 1-4.Davis, W. B. 1944. Notes on Mexican mammals. J. Mammal.,25: 370-403.Davis, W. B. 1965. Review of the Eptesicus brasiliensis complexin Middle America with the description of a newsubspecies from Costa Rica. J_Mammal., 46:229-240.Davis, W. B. 1969. A review of the small fruit bats (genusArtibeus) of Middle America. Southwestern Nat., 14:1529.Davis, W. B. 1970a. The large fruit bats (genus Artibeus) ofMiddle America, with a review of the Artibeus jamaicensiscomplex. J. Mammal., 51:1 05·122.

Davis, W. B. 1970b. A review of the small fruit bats (genusArtibeus) of Middle America. Part n. Southwestern Nat.,14:389-402.Davis, W. B., and D. C. Carter. 1962. Notes on Central Americanbats with description of a new subspecies of Mormoops.Southwestern at., 7:64-74.Davis, W. B., D. C. Carter, and R. H. Pine. 1964. Noteworthyrecords of Mexican and Central American bats. J. Mam·mal.,45:375-387.Davis, W. B., and R. J. Russell, Jr. 1952. Bats of the Mexicanstate of Morclos. J. Mammal., 33:234-239.Davis, W. B., and R. J. Russell. 195,t Mammals of the Mexicanstate of Morclos. J. Mammal., 35 :63-S0.Dearolf, K. 1953. The invertebrates of 75 caves in the UnitedStates. Proc. Pennsylvania Acad. Sci., 27:225-241.Dearolf, K. 1956. Survey of North American cave vertebrates.Proc. Pennsylvania Aead. Sci., 30:201-210.De Buen, F. 1940. Lista de peces de agua dulce de Mexico.En preparacion de su catalogo. Trab. Est. Limnol. Patzcuaro,2:1·66.De Buen, F. 1946. Ictiogeografla continental mexicana (I,II, y III). Rev. Soc. Mexicana Hist. Nat., 7:87-138.De Buen, F., 1972. Clase V. Los peces teleostomos (Teleostomi).Pp. 55-332 in 1. Cendrero, ed., Zoologia hispanoamericana.Vertebrados. Mexico, D. F.: Editorial Parma.Decou, V. G., and J. Therond. 1977. Histerid es hypoges deCuba. Resultats des Expeditions Biospeologiques CubanoRoumaines aCuba, 2:403-405.de Cserna, Z. 1956. Tectoniea de Ja Sierra Madre Oriental deMexico, entre Torreon y Monterrey. CongI. Geol. Inter.nac., 20th Sesion. 87 pp., 7 pIs.de Cserna, Z. 1965. Reconocimiento geologico en la SierraMadre del Sur de Mexico, entre Chilpancingo y Acapulco,estado de Guerrero. Univ. Nac. Aut. Mexico, lnst. Geol.,Bol., 62. 64 pp.Delamare Deboutteville, C. 1948a. Recherches sur les collembolestermitophilcs et myrmecophiles (Ecologie, Ethologie,Systematique). Arch. Zool. Exper. Gen., 85:261-425.Delamare Deboutteville, C. 1948b. Sur la presence du genreAcherontiella Absolon dans les grottes de l'Ardeche et duGard. Notes Biospeol., 3:49-56.Delamare DcboutteviJIe, C.1949. Collemboles cavernicoles duTennessee et de I'Alabama. Notes Biospeol., 4: 117-124.Delamare DeboutteviUe, C. 1951. Microfaune du sol des paystemperes et tropicaux. Paris: Hermann. 360 pp.Delamare Deboutteville, C. ] 960. Biologie des eaux souterraineslittorales et continentales. Paris: Hermann. 750 pp.,1 pI.Delamare Deboutteville, C. 1971. La vie dans les grottes.Paris: Presses Universitaires de France. 126 pp.Delamare Deboutteville, C. 1976. Interet biologique et eco]ogique des crabes cavernicolcs du Guatemala et duMexique appartenant au genre TyphlopseudothelphusaRioja. Compt. Rend. Acad. Sci. Paris, ser. 0, 283:837-S40.Delamare Deboutteville, C. 1977. Sur la radiation evoJutivedes crabes du genre Typhlopseudothelphusa au Guatemalaet au Mexique avec description d'espeees nouvelles. Ann.Speleol.,31:115-129.Delamare Deboutteville, C., and L. Botosaneanu. 1970.Formes primitives vivantes. Paris: Hermann. 232 pp.Delamare DeboutteviUe, C., and C. Juberthie. 1976. Guatemala.Recherches biospeleologiques. Spelunca, Supplement,3:18-19,23-24.de la Torre, L. 1955. Bats from Guerrero, J alisco and Oaxaca,Mexico. FieIdiana: Zool., 37:695-701, pis. 30·31.Deane, B_ 1977. Trip report: 23-25 December 1969. Assoc.Mexican Cave Stud. News., 5:55-56.Denton, T. E. 1973. Fish chromosome methodology. Spring.field, Illinois: Charles C. Thomas. vii + 166 pp.Diaz N3jera, A. 1975. Listas y datos de distribueion geograficade los alacranes de Mexico (Scorpionida). Rev. Inv.Salud Publica (Mexico), 35: 1-36.Dixon, C. G. 1957. Geology of southern British Hondur88with notes on adjacent areas. Belize, British Honduras:Govt. Printer. 85 pp.Donovan, J. 1975. Caving in the Huixtan Area, Chiapas_Chen-Ven·Sil-Mut. Canadian Caver, 7(1):21-22, 23, 25-27.Drake, R. J. 1951. Humboldtiana tay/ori, new species, fromnorthern Coahuila. Rev. Soc. Malacologica, 8:93-96,pI. 13.Dreux, D. 1974. Recherches du C.E.R.S.M.T. au Guatemala.Trav. Inst. Speol. "Emile Racovitza", 13:205-211.Duellman, W. E. 1965. Amphibians and reptiles from theYucatan Peninsula, Mexico. Univ. Kansas Mus. Nat. Hist.Publ.,15:577-614.Dumitresco, M. 1973. Deux especes nouvelles du genreSchizomus (Schizomida), trouvees a Cuba. Resultats desExpeditions Biospeologiques Cubano-Roumaines aCuba,1:279-292.Dumitresco, M. 1977. Autres nouvelles especes du genreSchizomus des grottes de Cuba. Resultats des ExpeditionsBiospeologiques Cubano-Roumaines a Cuba, 2:147-158,1 pI.Dumitresco, M., and I. Juvara·Bals. 1971. Cryptocel/us cubanicusn. sp. (Arachnida-Ricinulei). Premier representant de lafam. Ricinuleidae de Cuba. Resultats des ExpeditionsBiospeologiques Cubano·Roumaines a Cuba, 1:259-275.Dumitresco, M., and I. Juvara-Bals. 1976. Position systematiquede Heteroricinoides bordoni n. g. n. sp. dans la familleRicinuleididae (Arachnida). Bol. Soc. VenezolanaEspeleol., 7: 147-180.Dumitresco, M., and T. N. Orghidan. 1977. Pseudoseorpionsde Cuba. Resultats des Expeditions BiospeologiquesCubano-Roumaines aCuba, 2:99-122, 2 pis.Dunn, E. R. 1926. The salamanders of the family Plethodontidae.Northampton, Massachusetts: Smith College. viii +441 pp.Durand, J. P. 1963. Developpement larvaire de l'oeil chez Ieprotee (Batracien cavernicole). Spelunca, 4th Ser., Mem.,3:181-184. ,Durand, J. P. 1968. Etude des poissons recoltes dans laGrotte de Umayalanta (Bolivie), Trichomycterus chabertisp. n. Ann. Speleol., 23:343-353.Durand, J. P.1972. Recherches sur I'appareil visuel du protee,Proteus anguinus Laurenti, urodele hypoge. Ann. SpeJeol.,26: 497-S24.Durand, J. P. 1976. Rudimentation des yeux chez les poissonset urodeIes souterrains. Bull. Soc. Zool. France, 101,suppl. 1: 13-21.Durand, J. P. 1977. Sur la structure oculaire, l'ultrastructuredes muscles extrinseques des enveloppes de l'oeil et deI'epithelium pigmentaire retinien de I'Anoptichthys(forme aveugle de I'Astyanax mexicanus-Characidae,Pisces). Ann. Speleol., 31: 149-161.Durand, J. P. 1979. Phenomenes de convergences tissulaireset cytologiques, lies aux processus degeneratifs qui affee.tent l'oeil, chez deux teleosteens cavernicoles Astyanax(Anoptichthys) mexicanus (Characidae) et Lucifuga(Stygicola) dentatus (Ophidiidae). Bull. Soc. Zool. France,103:269-274, pI.265

Ediger, G. 1970. The Birthday Passage. Texas Caver, 15:3-7.Edwards, C. L. 1971. Relative humidity and temperaturepreference responses of the ricinuleid Cryptoce/luspelaezi (Arachnida). M. S. Thesis. Lubbock: Texas TechUniv. 50 pp.Egar, M. W.1974. An ultrastructural study of the optic nerveof the blind cave fish. (Abstr.). Anat. Rec., 178:350.Elliott, W. R. 1970. El Sotano de Soyate. Texas Caver,15:63-66.Elliott, W. R. 1971. Temperature preferences of aquatic,cave-adapted crustaceans from Central Texas and Mexico.M. S. Thesis. Lubbock: Texas Tech Univ. 47 pp.Elliott, W. R. 1972. Trip report: July-August 1969. Pp. 129131 in M. Walsh, ed., Mexican caving of the SouthwestTexas Grotto: 1966-1971. San Marcos: Southwest TexasGrotto.Elliott, W. R. 1973a. Temperature preference responses ofsome aquatic, troglobitic crustaceans from Central Texasand Mexico. (Abstr.). Bull. Natl. Speleol. Soc., 35:31-32.Elliott, W. R. 1973b. Trip report: 9-16 January 1971. Assoc.Mexican Cave Stud. News., 4:79.g5.Elliott, W. R. 1974. Trip report: 15 May - 8 June 1974.Assoc. Mexican Cave Stud. News., 5:19-21.Elliott, W. R. 1976. New cavernicolous Rhagidiidae fromIdaho, Washington, and Utah (Prostigmata: Acari: Arachnida).Occ. Papers Mus. Texas Tech Univ., 43.15 pp.Elliott, W. R. 1979. Sierra de Guatemala. Assoc. MexicanCave Stud. Activities News., 10:16-20.Elliott, W. R., and R. W. Mitchell. 1973. Temperature responsesof some aquatic, cave-adapted crustaceans fromCentral Texas and northeastern Mexico. Internatl. J.Speleol., 5 :171-189.Elliott, W. R., and J. R. Reddell. 1973. A checklist of thecave fauna of Mexico. VI. Valle de los Fantasmas Region,San Luis PotosI. Assoc. Mexican Cave Stud. Bull., 5:191·201.Elliott, W. R., and R. W. Strandtmann. 1971. New localityrecords for Rhagidia from Mexican and American caves.J. Kansas Entomol. Soc., 44:468-475.Erwin, T., D. R. Whitehead, and G. E. Ball. 1977. Checklistof the beetles of Canada, United States, Mexico, CentralAmerica, and the West Indies (Yellow Version). Part 1.The tiger beetles, ground beetles, water beetles, and relatedgroups (Families 1-9). Family 4. Carabidae, theground beetles_ Kinderhook, New York: World DigestPub1. 68 pp.Escherich, K. 1905. Das System der Lepismatiden. Zoologica,Stuttgart, 18:1-164, pls.I-4.Fennah, R. G. 1973. Three new cavernicolous species of Fulgoroidea(Homoptera) from Mexico and Western Australia.Proc. BioI. Soc. Washington, 86:439-446.Fenwick, .1. C. 1970. The pineal organ. Pp. 91-108 in W. S.Hoar and D. J. Randall, eds., Fish physiology, Volume IV.The nervous system, circulation, and respiration. NewYork: Academic Press.Fernandez Ruiz, G. 1976. Montaiias y cavernas. Donde lavida parece irnposible. Rev. Geogr. Universal, 1:6Q6-718.Finch, W. A., Jr. 1965. The karst landscape of Yucatan.Ph.D. Diss. Urbana: Univ. Illinois. Ann Arbor, Michigan:University Microfilms. x + 168 pp.Fingerrnan, M. 1965. Chromatophores. Physiol. Rev., 45:296-339.Finn, E.1971. Mexico '71. Canadian Caver, no. 4:45-54.Fish, J. 1965. Trip report: July 18-28, 1965. Assoc. MexicanCave Stud. News., 1:65-68.Fish, J. 1968. Caves of Guerrero and Morelos. Assoc. MexicanCave Stud. News., 2:111-123.Fish, J. 1970. Exploration of Sotano de San Agustin, Oaxaca,Mexico. Canadian Caver, no. 3:3-7, map.Fish, J. 1974. La Sistema de Los Sabinos. Mexico's longestcave. Canadian Caver, 6(1 ):3-20.Fish, J. 1978a. EI Sotano de J apones. Canadian Caver, 10(1):1-2, map.Fish, J. 1978b. The Xilitla Plateau & Sotano de Trinidad.Canadian Caver, 10(2):39-53.Fish, J. 1979. The Xilitla Plateau. Part 2. Canadian Caver,11(1):3-23.Fish, J. E. 1977. Karst hydrogeology and geomorphology ofthe Sierra de El Abra and the Valles-San Luis Potosiregion, Mexico. Ph.D. Dissertation. Hamilton, Ontario:McMaster Univ. xvii + 469 pp.Fish, J., and J. Reddell. 1965. Trip report: August 18-26_Assoc. Mexican Cave Stud. News., 1:73-77.Fish, J., and J. Reddell. 1967. Trip report: 2-19 August 1966.Assoc. Mexican Cave Stud. News., 2:82.g7.Fish, J., and W. Russell. 1966. Preliminary report on thecaves of Huautla de Jirnenez. Assoc. lexican Cave' Stud.News., 2:59-67.Fisk, F. W. 1977. Notes on cockroaches (Blattaria) fromcaves in Chiapas, Mexico and environs with descriptionsof three new species. Quad. Accad. Naz. Lincei, Probl.Att. Sci. Cult., 171(3):267·274, pI. 1.Fisk, F. W., and A. B. Gurney. 1972. Synopsis of the neotropicalcockroaches of the genus Nesomylacris (Dictyoptera:Blattaria: BlattelJidae). Proc. Entomol. Soc. Washington,74:196-206.Flannery, K. V. 1967. Vertebrate fauna and hunting patterns.Pp. 132-177 in D. S. Byers, ed., The prehistory of the TehuacanValley. Volume One. Environment and subsistence.Austin: Univ. Texas Press.Flores Crespo, R., R. J. Burns, and S. B. Linhart. 1970. Loadliftingcapacity of the vampire bat. J. Mammal., 51:627629.Franck, A. 1962. Mexicanische Hohlencharaciniden im Ver·g1eich zu ihren oberirdischen Vorfahren. Straatsexamenarbeitder Universitiit Hamburg.Franck, A. 1964. Vergleichende Untersuchungen am HohlenfischAnoptichthys antrobius und seinem oberirdischenVorfahren Astyanax mexicanus. Zool. Anz., 172:95-107.Franck·Krahe, C. 1962. Mexikanische Hohlencharaciniden irnVergleich zu ihren oberirdischen Vorfahren. StaatsexamaensarbeitUniversitiit Hamburg.Francke, O. F. 1977. The genus Dip/ocentrus in the YucatanPeninsula with description of two new troglobites (Scorpionida,Diplocentridae). Assoc. Mexican Cave Stud. Bull.,6:49·61.Francke, O. F. 1978. New troglobite scorpion of genus Dip/ocentrus(Scorpionida: Diplocentridae). Entomol. News,89:39-45.Frank, S. 1960. Anoptichthys jordani Hubbs a Innes 1936.Akvar. terar., Praha, 3(3):39-41.Frank, S. 1961. A morphological study about blind cave fishAnoptichthys jordani. Vest. Ceskosl. Zool. Spolec., 25:347-365.Frank, S. 1968. Der Augenlose. Der blinde Hohlensalmlerkommt nicht blind zur Welt. Aquarien Mag., 7:299-301.Frank, S. 1969. Blind geboren? Entwicklung und Riickbil·dung des Auges bei einem blinden Hohlensalmler. Mikrokosmos,1:11-13.Franz, H. 1977. Scydmaeniden aus Mexico (mit einter Re-

vision der von Schaufuss aus Zentralamerika beschriebenenArten). Quad. Accad. Naz. Lincei, Prob!. Att. Sci.Cult., 171(3):349·372.Fries, C., Jr. 1956. Bosquejo geologico de la region entreMexico, D. F. y Acapulco, Gro. Pp. 7-53 in Maldonado·Koerdell, ed., Geologia a 10 largo de la carretera entreMexico, D. F. y Acapulco, Gro., via Taxco, Gro. y Chilpancingo,Gro. Geologia de los alrededores de Acapulco,Gro. Los yacimientos de dolomita de EI Ocotito, Gro.Congr. Geo!. Internac., 20th Sesion, Excursiones A·9 yC·12.Froeschner, R. C. 1960. Cydnidae of the Western Hemisphere.Proc. United States Nat!. Mus., 111:337.680, pls.l-l3.Fromen, C. 1965. Trip report: August 1964; ThanksgivingDay 1964; 14-19 April 1965. Assoc. Mexican Cave Stud.News., 1:44-45.Fuentes, R. P. 1964. Stratigraphy of Sierra Santa Clara andSierra Gomas, Nuevo Leon, Mexico. M. A. Thesis. Austin:Univ. Texas at Austin. x + 217 pp.Fujii, R. 1969. Chromatophores and pigments. Pp. 307-353in W. S. Hoar and D. J. Randall, eds., Fish physiology,Volume HI: Reproduction and growth, bioluminescence,pigments, and poisons. New York: Academic Press.Furtos, N. C. 1936. On the Ostracoda from the cenotes ofYucatan and vicinity. Carnegie Inst. Washington Pub!.,457:89-115.Furtos, N. C. 1938. A new species of Cypridopsis from Yuca·tan. Carnegie Inst. Washington Pub!., 491 :155-157.Gaige, H. T. 1936. Some reptiles and amphibians from Yucatanand Campeche, Mexico. Carnegie Inst. WashingtonPubl.,457:289-304.Gaige, H. T. 1938. Some reptilian records from caves ofYucatan. Carnegie Inst. Washington Pub!., 491:297.298.Garcia Lozano, G. 1939a. Deposito de guano en la Cueva LosCuarteles, del Municipio de Almada, Tamps. Rev. Industria,4(20):5-7.Garcia Lozano, G. 1939b. EI guano en los estados de Coahuilay Nuevo Leon. Rev. Industria, 3(18):5-28.Garfias, V. R., and T. C. Chapin. 1949. Geologia de Mexico.Mexico, D. F.: Editorial Jus. 202 pp., 6 figs.Gascoyne, M., and B. Pratt. 1975. The MUCC expedition toMexico 1974-5. Canadian Caver, 7(1 ):34-46.Gates, G. E. 1968. On a new species of earthworm from aMexican cave. Internatl. J. Speleo!., 3:63-70.Gates, G. E. 1970. On a new species of earthworm fromanother Mexican cave. Southwestern Nat., 15:267·269.Gates, G. E. 1971. On some earthworms from Mexican caves.Assoc. Mexican Cave Stud. Bull., 4:~.Gates, G. E. 1972. Contributions to North American earth·worms (Annelida). No.5. On variation in another anthro·pochorous species of the oriental earthworm genus Phere·tima Kinberg 1866 (Megascolecidae). Bull. Tall TimbersRes. Sta., 13:18·44.Gates, G. E. 1973. On more earthworms from Mexican cavesAssoc. Mexican Cave Stud. Bull., 5:21-24.Gates, G. E. 1977. On some earthworms from North Ameri·can caves. Assoc. Mexican Cave Stud. Bull., 6:1.4.Genoways, H. H., and J. K. Jones, Jr. 1968. Notes on batsfrom the Mexican state of Zacatecas. J. Mamma!., 49:743-745.Gertsch, W. J. 1958. The spider genus Loxosceles in NorthAmerica, Central America, and the West Indies. AmericanMus. Nov., 1907.46 pp.Gertsch, W. J. 1960. Descriptions of American spiders of thefamily Symphytognathidae. American Mus. Nov., 1981.40 pp.Gertsch, W. J. 1971a. AJ'eport on some Mexican cave spiders.Assoc. Mexican Cave Stud. Bull., 4:47-111.Gertsch, W. J. 1971b. Three new ricinuleids from Mexicancaves (Arachnida, Ricinulei). Assoc. Mexican Cave Stud.Bull., 4:127·135.Gertsch, W. J. 1973a. The cavernicolous fauna of Hawaiianlava tubes, 3. Araneae (spiders). Pacific Insects, 15:163·180.Gertsch, W. J. 1973b. A report on cave spiders from Mexicoand Central America. Assoc. Mexican Cave Stud. Bull.,5:141-163.Gertsch, W. J. 1974. The spiderfamily Leptonetidae in NorthAmerica. J. Arachno!., 1:145-203.Gertsch, W. J. 1977a. On two ricinuleids from the YucatanPeninsula (Arachnida: Ricinulei). Assoc. Mexican 'CaveStud. Bull., 6: 133·138.Gertsch, W. J. 1977b. Report on cavernicole and epigeanspiders from the Yucatan Peninsula. Assoc. Mexican CaveStud. Bull., 6:103-131.Gertsch, W.J. In press. The spider family Nesticidae (Araneae)in North America, Central America and the West Indies.Bull. Texas Mem. Mus.Gertsch, W. J., and N. I. Platnick. 1975. A revision of thetrapdoor spider genus Cyclocosmia (Araneae, Ctenizidae).American Mus. Nov., 2580. 20 pp.Gertsch, W. J., and M. Soleglad. 1972. Studies of NorthAmerican scorpions of the genera Uroctonus and Vejovis(Scorpionida, Vejovidae). Bull. American Mus. Nat. Hist.,148:547·608.Gertychowa, R. 1970. Studies on the ethology and spaceorientation of the blind cave fish Anoptichthys jordaniHubbs et Innes, 1936 (Characidae). Folia BioI., Krakow,18:9-69.Gertychowa, R. 1971. Heliotaktyzm mlodych rybek jaskiniowychAnoptichthys jordani Hubbs et Innes. PrzegladZool.,15:66-69.Gisin, H. 1951. Collemboles recoltes dans des grottes duMoyen Atlas. Bull. Soc. Sci. Nat. Maroc, 31:53·56.Giuseffi, S., T. C. Kane, and W. F. Duggleby. 1978. Geneticvariability in the Kentucky cave beetle Neaphaenops tell·kampfii (Coleoptera: Carabidae). Evolution, 32:679·681.Glaser, D. 1965. Untersuchungen tiber die absoluten Geschmacksschwellenvon Fischen (Phoxinus phoxinus L., Gasterostusaculeatus 1., Hemigrammus caudovittatus Ahlund Anoptichthysjordani Hubbs et Innes). Diss. Giessen.Glaser, D. 1968. Zum Verhalten blinder Fische. Z. Tierpsycho!.,25:648-658.Glydenholm, A. 0., and J. J. Scheel. 1971. Chromosomenumbers of fishes. I. J. Fish Bio!., 3:479-486.Goettert, L. 1962. Orientierungsmoglichkeiten beim augen·losen Hohlenfisch (Anoptichthys jordani). Naturwiss.Rdsch., Stuttgart, 15:56-58.Goldman, E. A. 1951. Biological investigations in Mexico.Smithsonian Misc. Coli., 115. 476 pp., 70 pis"Gonzalez-Angulo, W., and R. E. Ryckman. 1967. Epizootiologyof Trypanosoma CMJzi in southwestern NorthAmerica. Part IX: An investigation to determine the incidenceof Trypanosoma CMJzi infections in Triatominaeand man on the Yucatan Peninsula of Mexico. J. Med.Entomo!.,4:44-47.Gonzalez Ochoa, A. 1963a. Epidemiologia de la histoplasmosisprimaria en Mexico. Rev. Inst. Salubr. Enferro.Trop., Mexico, 23:65-80.267

Gonzalez Ochoa, A. 1963b. Relaciones entre el habitat delmurcielago y el Histoplasma capsulatum. Rev. Inst. Salubr.Enfenn. Trop., Mexico, 23:81-86.Gonzalez Ochoa, A. 1964. Realizaciones de la investigacioncientifica en Mexico para la salud publica. III. Histoplas.mosis. Gac. Med. Mexico, 94:981-986.Gonzalez Sponga, M. A. ]974. Dos nuevas especies de alacranesdel genero Tityus, en las cuevas venezolanas (Scorpionida:Buthidae). Bol. Soc. Venezolana Espeleol., 5:5572.Goodnight, C. J., and M. L. Goodnight. 1942. Phalangidafrom Mexico. American r.lus. Nov., 1211. 18 pp.Goodnight, C. J., and M. L. Goodnight. ]944. More Phalangidafrom Mexico. American Mus. Nov., 1249. 13 pp.Goodnight, C. J., and M. L. Goodnight. 1945. AdditionalPhalangida from Mexico. American Mus. Nov., 1281.17 pp.Goodnight, C. J., and M. L. Goodnight. 1953. The opilionidfauna of Chiapas, Mexico, and adjacent areas (Arachnoi.dea, Opiliones). American Mus. Nov., 1610. 81 pp.Goodnight, C. J., and M. L. Goodnight. 1971. Opilionids(Phalangida) of the family Phalangodidae from Mexicancaves. Assoc. Mexican Cave Stud. Bull., 4:33-45.Goodnight, C. J., and M. L. Goodnight. 1973. Opilionids(Phalangida) from Mexican caves. Assoc. Mexican CaveStud. Bull., 5:83-96.Goodnight, C. J., and M. L. Goodnight. 1977. Laniatores(Opiliones) of the Yucatan Peninsula and Belize (BritishHonduras). Assoc. Mexican Cave Stud. Bull., 6:139-166.Goodnight, M. L., and C. J. Goodnight. 1976. Observationson the systematics, development, and habits of Erginulusclavotibinlis (Opiliones: Cosmetidae). Trans. AmericanMieroseop. Soc., 95:654-664.Goodrick, C. L. 1962. Differential adaptation of activity inthe blind cave fish, Anoptichthys jordani. Perceptual andMotor Skills, 14:10.Goodwin, G. G. 1934. Mammals collected by A. W. Anthonyin Guatemala, 1924-1928. Bull. American Mus. Nat. Hist.,68:1-60, pis. 1-5.Goodwin, G. G. 1955. Mammals from Guatemala, with thedescription of a new little brown bat. American Mus.Nov., 1744. 5 pp.Goodwin, G. G. ]969. Mammals from the state of Oaxaca,Mexico, in the American Museum of Natural History.Bull. American Mus. Nat. Hist., 141: 1-270, pis. 1-40.Gordan, J. 1957. A bibliography of the order Mysidaeea.Bull. American Mus. Nat. Hist., 112:279-393.Gordon, I. 1960. On a Stygiomysis from the West Indies,with a note on Spelaeogriphus (Crustacea, Peracarida).Bull. British Mus. (Nat. Hist.), Zool., 6:285-324, pis. 3-4.Gordon, M. 1957. Physiological genetics of fishes. Pp. 431501 in M. E. Brown, ed., The physiology of fishes, VolumeIll: Behavior. New York: Academic Press.Gordon, M. S., and D. E. Rosen. 1962. A eavernicolous formof the poeciliid fish, Poecilin sphenops from Tabasco,Mexico. Copeia, 1962:360-368.Gould, R. 1968. Mayan cave discoveries. Archeology. ExplorersJ.,46:164·167.Graham, E., L. McNatt, and M. A. Gutchen. 1980. Excavationsin Footprint Cave, Caves Branch, Belize. J. FieldArchaeol.,7:153-172.Greenberg, A. 1938. Strange spawning of the blind characin.Aquarium, Philadelphia, 7(5):80.Greenwood, P. H. 1967. Blind cave fishes. Stud. Speleol.,1:262-274, pI. 40.Greer, J. W. 1974. Petroglyphs at Sotano de los Monos,Sierra de El Abra, San Luis Potosi, Mexico. Assoc. MexicanCave Stud. News., 5:23-30.Greer, J. W. 1977. Archeological notes on Hoya de Higueron,Sierra de EI Abra, S. L. P. Assoc. Mexican Cave Stud.News., 5:72-73.Greer, J. W. 1979. Archeological reconnaissance in a limestonekarst region in northern Queretaro. Assoc. MexicanCave Stud. ews., 5: 106-123.Gresser, E. B., and C. M. Breder, Jr. 1940. The histology ofthe eye of the cave charaein, Anoptichthys. Zoologica,New York, 25:113-116, pis. 1-3.Grobbel, G., and G. Hahn. 1958. Morphologie und Histologieder ;';cill'norgane des augenlosen Hohlenfisches, Anoptichth)sj()rJ"ni im Vergleich zu anderen Teleosteern.I.. \lorl'h. Oko/. Tine, 47:249·266.Grubb,..\. 1'J7.'5. 1'J7.'5 Yueatan Expedition. Assoc. MexicanCavl' :-:tud. ,\ctivitil's \l'ws.. :\:2-,1-(;ruia, \'1. I'J70. Con,irlt'ralioll' 'lIr dl'IlX 1'51'('(":5 avcugles deCollemboles eavernieoll" ,k HOllmanil': ·ld,erontidesspelaea (I o 11l'SeO ) f'l I. trl/UJs"ch ia/! «(; no ia). Tra v. Inst.Speo/. "Emile l{al·ovill.a·'. '): 1 )7-.")1).Grummon, It. \ .. and \. \oviek. 1')(,;\. ()hsladl' avoidaneein the bat. .II"a"/lIs "",xicaIlIlS. I'hysio/. I.oo/.. :\(" :\61369.Guatemala. Institulo Ccogr:lfim \al'iona/. ]'J(,7. \\al'a geologico,1:50,000. lIoya 1862 IV (;. La lkmocracia.Guatemala. Institulo Gcografieo i\aeional. ]9(,8a. \lapageologico, l:50,OOO. Hoya 1%2 III G. Chiantla.Guatemala. Instituto GeogrHico :\acional. 1968L. \lapageologico, 1:50,000. Hoya 1862 III (;. Cuilco.Guatemala. Instituto GeogrHico :\aeionaJ. 1975. ''lapageologico de la Republica de (;uatemala. J'J70. Eseala1:500,000. 2nd Impresion.Gueorguiev, V. H. 1974. La Laurasi.. ct la formation de lafaune troglobie terrestre dans la I'(,nin,"k Ilalkanique.Compl. Rend. Acad. Bulgarc Sci., 27:IJBJ-(JB:I.Gurnee, R. II. 1962. The caves of GlIall"mala. Nat/. Spelcol.Soc. Bull., 24:25-30.Gurnee, R. II. 1968. Mayan cave discoveries. Description ofthe expedition. Explorers J., 46: 148-160.Gurney, A. B. 1943. A synsopsis of the psocids of the tribePsyllipsoeini, including the description of an unusual newgenus from Arizona (Corrodentia: Empheriidae: Empheriinae).Ann. Entomol. Soc. America, 36: 195-220.Gutierrez Gil, R. 1956. Bosquejo geologico del estado deChiapas. Pp. 9-60, figs. 1-13 in M. MaJdonado-Koerdell,ed., Geologia del Mesozoico y estratigrafia Permica delestado de Chiapas. Congr. Geol. [nternae., 20th Sesion,Excursion C-15.Hahn, G. 1957. Ferntastsinn und Stromungssinn beim augenlosenHohlenfisch, Anoptichthysjordani lIubbsund Innes,und das Problem der Rezeptionsorte von Liehtreizen.Dissertation, Koln, 81 pp.Hahn, G. 1960. Ferntastsinn und Stromungssinn beim augenlosenHohlenfisch Anoptichthys jordani Hubbs und Innesim Vergleieh zu einigen andercn Teleosteern. (Abstr.).aturwissensehaften,47:611-612.Halffter, G. 1950. 11\ Conveneion :Ylundial de Espeleologos.Cieneia, Mexico, 10: 165-166.Hall, E. R., and W. W. Dalquest. 1963. The mammals of Vera·cruz. Univ. Kansas Publ., Mus. Nat. Hist., 14:165-362.Hall, E. R., and K. R. Kelson. 1959. The mammals of NorthAmerica. New York: Ronald Press Co. 2 vols.Hall, E. R., and B. Villa R. 1949. An annotated check list of268

the mammals of Michoacan, Mexico. Univ. Kansas Publ.,Mus. Nat. Hist., I :431-472, pis. 4-5.Hall, F. G. 1936. Physical and chemical survey of cenotes ofYucatan. Carnegie Inst. Washington Publ., 457:5-16.Handley, C. 0., Jr. 1959. A revision of American bats of thegenera Euderma and Plecotus. Proc. United States Natl.Mus., 1l0:95-246.Handley, C. 0., Jr. 1966. Descriptions of new bats (Chuo·derma and Artibeus) from Mexico. Anal. Inst. Bio!.,Mexico, 36:297-301.Hara, T. J. 1971. Chemoreception. Pp. 79-120 in W. S. Hoarand D. J. Randall, eds., Fish physiology, Volume V:Sensory systems and electric organs. New York: AcademicPress.Harden, S. 1971. Trips: 24 December 1970·4 January 1971.Texas Caver, 16:36-37.Harmon, R. 1979. Trip report: 3-12 April 1968. Assoc.Mexican Cave Stud. News., 5 :101-103.Harrell, B. E. 1951. The birds of Rancho del Cielo. An ecologicalinvestigation in the oak-sweet gum forests of Tamaulipas,Mexico. M. A. Thesis. Univ. Minnesota. 283 pp.Harris, A. H. 1971. Use of Texas caves by terrestrial mammals.Pp. 117-121 in E. L. Lundelius, Jr., and B. H.Slaughter, eds., Natural history of Texas caves. Dallas:Gulf Nat. Hist.Hart, C. W., Jr. 1962. A revision of the ostracods of the familyEntocytheridae. Proc. Acad. Nat. Sci. Philadelphia,114:121-147.Hart, C. W., Jr. 1978. A new species of the genus Sphaeromi·cola (Ostracoda: Entocytheridae: Sphaeromicolinae) fromTexas, with notes on relationships between European andNorth American species. Proc. BioI. Soc. Washington,91:724·730.Hart, D. G., and C. W. Hart, Jr. 1974. The ostracod familyEntocytheridae. Acad. Nat. Sci. Philadelphia Monogr.,18. ix + 239 pp.Hartnoll, R. G. 1964a. The freshwater grapsid crabs of J amaica.Proc. Linn. Soc. London, 175:145-169.Hartnoll, R. G. 1964b. Two cavernicolous decapods fromJamaica. Crustaceana, 7:78-79.Hau, R. T. 1938. Notes concerning mammals collected inYucatan. J. Mammal., 19:333-337.HaU, R. T. 1953. Faunal and archeological researches inYucatan caves. 1. Introduction. Cranbrook Inst. Sci.Bull., 33:1-42, pis. 1·6.HaU, R. T., and B. Villa R. 1950. Observaciones sobre algunosmamiferos de Yucatan y Quintana Roo. Anal. Inst.BioI., Mexico, 21:215-240.Heim, A. 1940. The front ranges of the Sierra Madre Oriental,Mexico, from Ciudad Victoria to Tamazunchale.Eco!. Geol. Helvetiae, 33:313-362.Hendrichs, J., and C. Bolivar y Pieltain. 1966. Hallazgo de unnuevo Mexisphodrus cavernicola en el estado de Hidalgo(Mexico): M. gertschi nov. sp. Ciencia, Mexico, 25:7-10,pI. 1.Hendrichs, J., and C. Bolivar y Pieltain. 1973. Un nuevo esfodrinociego del Sotano de San Agustin, Oaxaca, Mexico(Coleopt., Carab.). Ciencia, Mexico, 28:37-41.Herald, E. S. 1972. Fishes of North America. New York:Doubleday and Co. 254 pp.Herman, L., Jr. 1969. A troglobitic staphylinid from Mexico(Coleoptera, Staphylinidae, Paederinae). American Mus.Nov., 2367. 9 pp.Herman, L., Jr. 1970. The ecology, phylogeny, and taxonomyofStilicolina (Coleoptera, Staphylinidae, Paederinae).American Mus. Nov., 2412. 26 pp.Herrera, A. L. 1891. Fauna cavernicola. Mem. Soc. Cient."Antonio Alzate", 5 :218-221, pis. 2-3.Herrera, A. L. 1911. Nota adicional. Naturaleza, 3rd ser.,1(2):4-6.Hershberger, B. 1967. Trip report: 22 July 1966. Assoc.Mexican Cave Stu.d. News., 4:88-90.Heuts, B., and G. Thines. 1971. Behavioural changes in astock of Barbus conchonius (Hamilton, Pisces, Cyprinidae)after accidentally induced partial depigmentationand blinding. Z. Tierpsychol., 28: 113-163.Heuts, M. J. 1951. Ecology, variation and adaptation of theblind African cave fish, Caecobarbus geertsii Blgr. Ann.Soc. Roy. Zoo!' Belgique, 82:155-230.Heuts, M. J. 1953a. Comment on "Cave fish evolution."Evolution, 7:391-392.Heuts, M. J. 1953b. Regressive evolution in cave animals.Symp. Soc. Exp. BioI., 7:290-309.Heuts, M. J. 1961. Cave biology. Pp. 195-197 in P. Gray, ed.,The encyclopedia of the biological sciences. New York:Reinhold Book Corp.Hobbs, H. H., Jr. 1941. A new crayfish from San Luis Potosi,Mexico. (Decapoda, Astacidae). Zoologica, New York,26:1-4.Hobbs, H. H., Jr. 1943. Two new crayfishes of the genus Procambarusfrom Mexico (Decapoda, Astacidae). Lloydia,6:198-206.Hobbs, H. H., Jr. 1966. An illustrated key to the species ofthe genus Ankylocythere with a description of a new speciesfrom Louisiana (Ostracoda, Entocytheridae). Proc.Louisiana Acad. Sci., 29:67-75.Hobbs, H. H., Jr. 1969a. On the distribution and phylogenyof the crayfish genus Cambarus. Pp. 93-178 in P. C. Holt,ed., The distributional history of the biota of the southernAppalachians. Part I: Invertebrates. Virginia PolytechnicInst., Res. Div. Monogr., 1.Hobbs, H. H., Jr. 1969b. Procambarus villalobosi, un nuevocambarino de San Luis Potosi, Mexico (Decapoda, Astacidae).Anal. Inst. BioI., Mexico, 38, Ser. Cienc. del Mar yLimnol., (I):41-46.Hobbs, H. H., Jr. 1971. The entocytherid ostracods of Mexicoand Cuba. Smithsonian Contr. Zoo!', 81. 55 pp.Hobbs, H. H., Jr. 1972a. Crayfishes (Astacidae) of North andMiddle America. Biota of Freshwater Ecosystems IdentificationManual, 9. 173 pp.Hobbs, H. H., Jr. 1972b. The subgenera of the crayfish genusProcam barus (Decapoda: Astacidae). Smithsonian Contr.Zool., 117. 22 pp.Hobbs, H. H., Jr. 1973a. Three new troglobitic decapod crustaceansfrom Oaxaca, Mexico. Assoc. Mexican Cave Stud.Bull., 5:25-38.:fobbs, H. H., Jr. 1973b. Two new trogiobitic shrimps (Decapoda:Alpheidae and Palaemonidae) from Oaxaca, Mexico.Assoc. Mexican Cave Stud. Bull., 5 :73-80.Hobbs, H. H., Jr. 1974. A checklist of the North and MiddleAmerican crayfishes (Decapoda: Astacidae and Cambaridae).Smithsonian Contr. Zool., 166. 161 pp.Hobbs, H. H., Jr. 1975. New crayfishes (Decapoda: Cambaridae)from the southern United States and Mexico. SmithsonianContr. Zoo!., 201. 34 pp.Hobbs, H. H., Jr. 1977. Cave-inhabiting crayfishes of Chiapas,Mexico (Decapoda: Cambaridae). Quad. Accad. Naz.Lincei, Probl. AU. Sci. Cult., 171(3):197-206.269

Hobbs, H. H.,Jr., and T. C. Barr, Jr. 1972. Origins and affini·ties of the troglobitic crayfishes of North America (Decapoda:Astacidae). II. Genus Orconectes. SmithsonianContr. Zool., 105. 84 pp.Hobbs, H. H., Jr., and H. H. Hobbs 111. 1973. The genusSphaeromicola (Ostracoda, Entocytheridae) in Mexico.Assoc. Mexican Cave Stud. Bull., 5 :39·42.Hobbs, H. H., Jr., H. H. Hobbs III, and M. A. Daniel. 1977. Areview of the troglobitic decapod crustaceans of theAmericas. Smithsonian Contr. Zool., 244. 183 pp.Hobbs, H. H., Jr., and A. Villalobos. 1964. Los cambarinosde Cuba. Anal. Inst. BioI., Mexico, 34:307-366.Hobbs, H. H., III. 1979. Additional notes on cave shrimps(Crustacea: Atyidae and Palaemonidae) from the YucatanPeninsula, Mexico. Proc. BioI. Soc. Washington, 92:618633.Hobbs, H. H., III, and H. H. Hobbs, Jr. 1976. On the troglobiticshrimps of the Yucatan Peninsula, Mexico (Decapoda:Atyidae and Palaemonidae). Smithsonian Contr.Zool., 240. 23 pp.Hoffman, R. L. 1976. A new lophodesmid milliped from aGuatemalan cave, with notes on related forms (Polydesmida:Pyrgodesmidae). Rev. Suisse Zool., 83:307-316.Hoffmann, A. 1944. Los ectoparasitos de los murcielagosmexicanos. Tesis. Universidad Nacional de Mexico.150 pp.Hoffmann, A. 1953. Estado actual del conocimiento de losestreblidos mexicanos. (Diptera: Pupipara). Mem. Congr.Cient. Mexicano, 7:175-193.Hoffmann, A. 1962. Monografia de los Ixodoidea de Mexico.I Parte. Rev. Soc. Mexicana Hist. Nat., 23:191-307.Hoffmann, A. 1970. Estudio monografico de los trombiculi·dos de Mexico (Acarina: Trombiculidae).-Primera parte.An. Esc. Nac. Cienc. BioI., Mexico, 18:191-263.Hoffmann, A. 1972. Dos ectoparasitos de murcielagos pocofrecuentes en Mexico. Rev. Latino-Americano Microbiol.,14:191-195.Hoffmann, A. 1976. Relacion bibliografica preliminar de lasaranasde Mexico (Arachnida: Araneae). Univ. Nac. AutonMexico, Inst. BioI., Publ. Esp., 3. 117 pp.Hoffmann, A., I. E. Canales, G. Ibarra, R. Lira, T. M. Perez,M. T. Quintero, and M. 1. Zamudio. 1978. Estudios eco·logicos de una cueva en Valle de Bravo, Mexico. (Abstr.).Folia Entomol. Mexicana, 39·40:41.Hoffmann, A., and I. B. de Barrera. 1970. Acaros de la familiaSpelaeorhynchidae. Rev. Latino-Americano MicrobioI.,12:145·149.Hoffmeister, D. F. 1957. Review of the long-nosed bats ofthe genus Leptonycteris. J. Mammal., 38:454-461.Holman, J. A. 1970. A small Pleistocene herpetofauna fromTamaulipas. Quart.J. Florida Acad. Sci., 32:153-158.Holsinger, J. R. 1963. Annotated checklist of the macroscopictroglobites of Virginia with notes on their geographicdistribution. Bull. Natl. Speleol. Soc., 25:23-36.Holsinger, J. R. 1965. Redescriptions of two poorly knownspecies of cavernicolous rhagidiid mites (Acarina: Trom·bidiformes) from Virginia and Kentucky. Acarologia,7:654-662.Holsinger, J. R. 1973. Two new species of the subterraneanamphipod genus Mexiweckelia (Gammaridae) fromMexico and Texas, with notes on the origin and distributionof the genus. Assoc. Mexican Cave Stud. Bull.,5:1-12.Holsinger, J. R. 1974a. Comments on the newly proposedgammaridean amphipod families Crangonycidae andMelitidae. Crustaceana, 26:316-318.Holsinger, J. R. 1974b. Zoogeography of the subterraneanamphipod crustaceans (Gammaridae, Hadzia group) ofthe Greater Caribbean region. (Abstr.). Virginia J. Sci.,25(2):64.Holsinger, J. R. 1976. The cave fauna of Pennsylvania. Pp.72-S7 in W. B. White, ed., Geology and biology of Pennsylvaniacaves. Pennsylvania Geol. Surv. (4th Ser.), Gen.Geol. Rept., 66.Holsinger, J. R. 1977a. A new genus and two new species ofsubterranean amphipod crustaceans (Gammaridae s. lat.)from the Yucatan Peninsula in Mexico. Assoc. MexicanCave Stud. Bull., 6:15-25.Holsinger, J. R. 1977b. Some observations on the taxonomyof the Cuban subterranean amphipod genus Weckelia(Gammaridae). Resultats des Expeditions BiospeologiquesCubano-Roumaines aCuba, 2:267·269.Holsinger, J. R. In press. Paramexiweckelia, a new genus ofsubterranean amphipod crustacean (Hadziidae) fromnorthern Mexico. Internatl. J. Spelcol.Holsinger, J. R., R. A. Baroody, and D. C. Culver. 1976. Theinvertebrate cave fauna of West Virginia. West VirginiaSpeleol. Surv. Bull., 7. 82 pp.Holsinger, J. R., and W. 1. Minckley.1971. A new genus andtwo new species of subterranean amphipod crustaceans(Gammaridae) from northern Mexico. Proc. BioI. Soc.Washington, 83:425·443.Holsinger, J. R., and S. B. Peck. 1968. A new genus and speciesof subterranean amphipod (Gammaridae) fromPuerto Rico, with notes on its ecology, evolution andrelationship to other Caribbean amphipods. Crustaceana,15: 249-262.Holsinger, J. R., and S. B. Peck. 1971. The invertebrate cavefauna of Georgia. Bull. Natl. Speleol. Soc., 33:23-44.Holt, P. C. 1973. A summary of the branchiobdellid (Annelida:Clitellata) fauna of Mesoamerica. Smithsonian Contr.Zool., 142.40 pp.Holthuis, L. B. 1950. The Palaemonidae collected by the Sibogaand Snellius Expeditions, with remarks on otherspecies. I. Subfamily Palaemoninae. The Deeapoda of theSiboga Expedition. Part X. Siboga-Expeditie, 39a9:1.268.Holthuis, L. B. 1952. A general revision of the Palaemonidae(Crustacea Decapoda Natantia) of the Americas. II. Thesubfamily Palaemoninae. Occ. Papers Allan HancockFoundation, 12. 396 pp.Holthuis,1. B. 1955. The recent genera of the caridean andstenopodidean shrimps (Class Crustacea, Order Decapoda,Superseetion Natantia) with keys for their determination.Zool. Verh., Leiden, 26: 1-157.Holthuis, L. B. 1956. An enumeration of the Crustacea Deca·poda Natantia inhabiting subterranean waters. Vie etMilieu, 7:43·76.Holthuis, L. B. 1964. Sesarma (Sesarma) cerberus, a newcavernicolous crab from Amboina. Zool. Mededelingen,40:65-72.Holthuis, 1. B. 1974. Bithynops luscus, a new genus and spe.cies of eavernicolous shrimp from Mexico (CrustaceaDecapoda, Palaemonidae). Quad. Accad. az. Lincei,Probl. Att. Sci. Cult., 171(2):135-142.Holthuis, 1. B. 1977. Cave shrimps (Crustacea Decapoda,Natantia) from Mexico. Quad. Accad. Naz. Lincei, Probl.Att. Sci. Cult., 171(3):173-195.270

Hooper, E. T. 1947. Noteson Mexican mammals. J. Mammal.,28:40-57.Hooper, E. T. 1953. Notes on mammals of Tamaulipas, Mexico.Occ. Papers Mus. Zool., Univ. Michigan, 544.12 pp.Hooper, E. T. 1961. Notes on mammals from western andsouthern Mexico. J. Mammal., 42:120-122.Horst, R. 1972. Bats as primary producers in an ecosystem.Bull. NatI. Speleol. Soc., 34:49-54.Horst, R., and M. Langworthy. 1972. Rabies in a colony ofvampire bats. J. Mammal., 53:903-905.Howarth, F. G. 1972. Cavernicoles in lava tubes on the islandof Hawaii. Science, 175:325-326.Howarth, F. G. 1973. The cavernicolous fauna of Hawaiianlava tubes, 1. Introduction. Pacific Insects, 15: 139-151.Howden, H. F. 1973. Four new species of Onthophagus fromMexico and the United States (Coleoptera; Scarabaeidae).Proc. Entomol. Soc. Washington, 75: 329-337.Howden, H. F., O. L. Cartwright, and G. Halffter. 1956.Descripcion de una nueva especie mexicana de Onthophaguscon anotaciones ecologicas sobre especies asociadas anidos de animales y a cuevas. Acta Zool. Mexicana, 1(9):1-16.Hubbell, T. H. 1938. New cave-crickets from Yucatan, with areview of the Pentacentrinae, and studies on the genusAmphiacusta (Orthoptera, Gryllidae). Carnegie Inst.Washington Publ., 491:191-233.Hubbell, T. H.1972. Records of cave Orthoptera from northeasternand central Mexico, with a revision of the gryllidgenus Paracophus and descriptions of three new genera ofRhaphidophoridae. Quad. Accad. Naz. Lincei, l'robl. Au.Sci. Cult., 171(1):47.115, pI. 1.Hubbell, T. H.1977. Middle American cave-crickets and allies.r. The group Phoberopodes, with a review of the phallicstructures of the Ceuthophilinae (Orthoptera Saltatoria:Ensifera: Rhaphidophondae). Quad. Accad. Naz. Lincei,Prob!. AU. Sci. Cult., 171(3):275-324.Hubbs, C. 1971. Texas cave fishes. Pp. 91-93 in E. L. Lundelius,Jr., and B. H. Slaughter, eds., Natural history ofTexas caves. Dallas: Gulf Nat. Hist.Hubbs, C. L. 1936. Fishes of the Yucatan Peninsula. CarnegieInst. Washington Publ., 457: 157-287, pis. 1-15.Hubbs, C. L. 1938. Fishes from the caves of Yucatan. CarnegieInst. Washington Publ., 491:261-295, pis. 1-4.Hubbs, C. L. 1943. Criteria for subspecies, species andgenera, as determined by researches on fishes. Ann. NewYork Acad. Sci., 44:109-121.Hubbs, C. L., and W. T. Innes. 1936. The first known blindfish of the family Characidae: A new genus from Mexico.Occ. Papers Mus. Zool. Univ. Michigan, 342. 7 pp., 1 pI.HiUher, W. 1975. Ein neuer Acherontides aus Peru. SenckenbergianaBioI., 56:283-288.Humbach, I. 1960a. Geruch und Geschmack bei den augenlosenHohlenfischen Anoptichthys jordani, Hubbs undInnes und Anoptichthys hubbsi, Alvarez. Inaug. Diss.Koln. 88 pp.Humbach, r. 1960b. Geruch und Geschmack bei den augenlosenHohlenfischen Anoptichthys lordani, Hubbs undInnes und Anoptichthys Hubbsi, Alvarez. (Abstr.).Naturwissenschaften,47:551-552.Hungerford, H. B. 1936. Aquatic and semiaquatic Hemipteracollected in Yucatan and Campeche. Carnegie Inst. WashingtonPubl., 457:145-150.Husmann, S. 1967. Die okologische Stellung der Hohlen- undSpaitengewasser innerhalb der subterranaquatilen Lebensbereiche.Internatl. J. Speleol., 2:409-436.Husson, R.,F. Graf,J. P. Henry,G. Magniez, and C.Marvillet.1973. Lea recherches biospeleologiques pou1'5uivies au Laboratoirede Biologie Animale et Generale de la Faculte desSciences de Dijon. Pp. 113-133 in Livre du cinquantenairede I'Institut de Speologie "Emile Racovitza". Bucuresti:Editura Academiei Republicii Socialiste Romania.Hykes, O. V. 1937. Anoptichthys jordani, Hubbs and Innes.Akvar. listy, 11:108-109.Hyman, L. H. 1938. Land planarians from Yucatan. CarnegieInst. Washington Publ. 491:23-32.Ingles, L. G. 1959. Notas acerca de los mamiferos mexicanos.Anal. Inst. Bio!., Mexico, 29:379-408.Innes, W. T. 1937. A cavern characin, Anoptichthys jordani,Hubbs & Innes. Aquarium, Philadelphla,5:200-202.Innes, W. T. 1966. Exotic aquarium fishes, 19th ed. Philadelphia:Innes Publ. Co. 541 pp.Jameson, R. 1977. Xilitla Plateau. June 8-15. Assoc. MexicanCave Stud. Activities News., 5:7-11.Jeannel, R. 1936. Monographie des Catopidae. Mem. Nus.Natl. Hist. Nat., Paris, n. ser., 1:1-433.Jeannel, R. 1943. Les fossiles vivants des cavernes, 3rd ed.Paris: Gallimard. 321 pp., 12 pis.Jeannel, R. 1949. Les coleopteres cavernicoles de la regiondes Appalaches. Etude systematique. Notes Biospeol.,4:37-104.Jefferys, R. 1979. Sotano del Buque. Assoc. Mexican CaveStud. Activities News., 10:70-74, map.Jimenez Guzman, A. 1968. Nuevos registros de murcielagospara Nuevo Leon, Mexico. Ana!. Inst. Bio!' Univ. Nal.Auton., Mexico, 39, Ser. Zoo!., (1):133--144.John, K. R. 1957. Observations on the behavior of blind andblinded fishes. Copeia, 1957:123-132.Johnson, H. N.1948. Derriengue: Vampire bat rabies inMexico. American J. Hyg., 47:189-204.Johnson, K. W. 1967. Temperature responses of the Mexicanblind cave-fishes of the genus Anoptichthys. M. S. Thesis.Lubbock: Texas Tech. Coil. 53 pp.Johnson, K.W., and L.M. Heath. 1977. Densityestirnates oftwocavernicoles in a Mexican cave. Ann. Spelt~o!., 31:131-135.Jones, J. K., Jr. 1964. Additional records of mammals fromDurango, Mexico. Trans. Kansas Acad. Sci., 66:750-753.Jones, J. K., Jr. 1966. Bats from Guatemala. Univ. KansasPub!., Mus. Nat. Hist., 16:439-472.Jones, J. K., Jr., and T. Alvarez. 1964. Additional records ofmammals from the Mexican state of San Luis Potosi. J.Mammal., 45: 302-303.Jones, J. K.,Jr., and D. C. Carter. 1976. Annotated checklist,with keys to subfamilies and genera. Pp. 7-38 in R. J.Baker, J. K. Jones, Jr., and D. C. Carter, oos., Biology ofbats of the New World family Phyllostomatidae. Part r.Spec. Pub!. Mus. Texas Tech Univ., 10.Jones, J. K., Jr., J. R. Choate, and A. Cadena. 1972. Mammalsfrom the Mexican state of Sinaloa. II. Chiroptera.Occ. Papers Mus. at. Hist., Univ. Kansas, 6: 1-29.Jones, J. K., Jr., H. H. Genoways, and T. E. Lawlor. 1974.Annotated checklist of mammals of the Yucatan Peninsula.II. Rodentia. Occ. Papers Mus. Texas Tech Univ.,22.24 pp.Jones, J. K., Jr., J. D. Smith, and T. Alvarez. 1965. Notes onbats from the Cape region of Baja California. Trans. SanDiego Soc. Nat. Hist., 14:53-56.Jones, J. K., Jr., J. D. Smith, and H. H. Genoways. 1973.Annotated checklist of mammals of the Yucatan Peninsula,Mexico. r. Chiroptera. Occ. Papers Mus. Texas TechUniv., 13.31 pp.271

Jordan, c. B. 1937. Bringing in the new cave fish, Anoptich.thys jordani Hubbs & Innes. Aquarium, Philadelphia,5:203·204.Jordan, c. B. 1946. Anoptichthys X Astyanax hybrids.Aquarium, Philadelphia, 15: 198.Juberthie, C. 1974. Speleologie. EncycL Int. Sci. Tech., 9:79-33.Juberthie.Jupeau, L. 1977. Sur Ie systc~me neurosecreteur dupedoncule oculaire d'un decapode souterrain micropthalme,Typhlatya garciai Chace. Ann. Speleol., 31: 107114.Judd, F. W. 1967. Notes on some mammals from Big BendNational Park. Southwestern Nat., 12: 192·194.Kiihling, J. 1957. Progressive und regressive Augenentwick.lung bei dem Hohlenfisch Anoptichthys jordani Hubbsund Innes und das Problem der Rezeptionsorte von Lichtreizen.Dissertation, Koln.Kiihling, J. 1961. Untersuchungen uber den Lichtsinn unddessen Lokalisation bei dem Hohlenfisch Anoptichthysjordani Hubbs & Innes (Characidae). Bio\. Zentr., 80:439451.Kaestner, A. 1968. Invertebrate zoology, Volume II. Trans·lated and adapted from the German by H. W. Levi andL. R. Levi. New York: Interscience Pub\. 472 pp.Kauffeld, J. E. 1954. Ctenobrycon spilurus fathers strangeyoung. Aquarium, Philadelphia, 23:140·141.Kawakatsu, M. 1973a. Animals from the Mexican caves.Hokkaido/Shinbun, Oct. 4·5, 1973. (In Japanese).Kawakatsu, M. 1973b. Dr. Robert W. Mitchell, an Americanbiospeologist, and "The Journal of Arachnology." Collectingand Breeding, 35( 11 ):255. (In Japanese).Kawakatsu, M. 1974. A report on caves of the United Statesand Mexico. Japan Caving, 6:23-29. (In Japanese)Kawakatsu, M. 1975. Planarians from the Mexican caves.T6sho K6ko-tsushin Seibutsu, 138:6·7. (In Japanese)Kawakatsu, M. ]976. Mexico: Its nature and landscape.Heredity (Iden), 30(3):34·45. (In Japanese)Kawakatsu, M. 1977. Ecology and distribution of freshwaterplanarians. Heredity (lden), 31(10):13·24. (In Japanese)Keirans, J. E., and C. M. Clifford. 1975. Nothoaspis reddelli,new genus and new species (Ixodoidea: Argasidae), from abat cave in Mexico. Ann. EntomoL Soc. America, 68:81·85.Keirans, J. E., C. M. Clifford, and J. R. ReddelL 1977. De.scription of the immature stages of Nothoaspi.< reddelli(Ixodoidea: Argasidae) from bat caves in Mexico. Ann.EntomoL Soc. America, 70:591-595.Kellogg, H.. 1932. Mexican tailless amphibians in the UnitedStates National Museum. Hull. United States NatL Mus.,160.224 pp., 1 pLKellum, L. B. 1936. Evolution of the Coahuila Peninsula,Mexico. Part Ill. Geology of the mountains west of theLaguna District. Bull. Geol. Soc. America, 47:1039·1090, 14 pis.Kenk, R. 1974. Index of the genera and species of the freshwatertriclads (Turbellaria) of the world. SmithsonianContr. Zool., 183. 90 pp.Kenk, R. 1975. Freshwater triclads (Turbellaria) of NorthAmerica. VIII. Dugesia arizonensis, new species. Proc.BioL Soc. Washington, 88: 113-120.King, R. E. 1939. Geological reconnaissance in northernSierra Madre Occidental of Mexico. BulL Geol. Soc.America, 50:1625·1722, pis. 1·9.Kirby, R. F., K. W. Thompson, and C. Hubbs. 1977. Karyo.typic similarities between the Mexican and blind tetras.Copeia, 1977:578·580.Kirpichnikov, V. S. 1973. Biochemical polymorphism andmicroevolution processes in fish. Pp. 223·241 in J. 1-1.Schroder, ed., Genetics and mutagenesis of fish. NewYork: Springer-Verlag.Klaas, E. E. 1968. Summer birds from the Yucatan Penin·sula, Mexico. Univ. Kansas Pub\., Mus. Nat. Hist., 17:579·611.Kleerekoper, H. 1969. Olfaction in fishes. Bloomington: Indi·ana lJniv. Press. viii + 222 pp.Knobloch, I. W. 1942. Notes on a collection of mammalsfrom the Sierra Madres of Chihuahua, Mexico. J. Mam·mal.,23:297.298.Koopman, K. F., and P. S. Martin. 1959. Subfossil mammalsfrom the Gomez Farias region and the tropical gradient ofeastern Mexico. J. Mammal., 40:1-12.Kosswig, C. 1960a. Darwin und die degenerative Evolution.AbhandL VerhandL Naturwiss. Ver. Hamburg, N. F.,4:21-42, pL 1.Kosswig, C. 1960b. Genetische Analyse stammesgeschicht·licher Einheiten. Verh. Deutsche Zoo\. Ges., 1959:42·73.Kosswig, C. 1960c. Zur Phylogenese sogenannter Anpas.sungsmcrkmale bei Hohlentieren. Int. Rev. Ges. Hydro.biol.,45:493-512.Kosswig, C. 1963. Genetische Analyse konstruktiver unddegenerativer Evolutionsprozesse. Z. ZooL Syst. Evolu·tionsforsch., 1:205·239.Kosswig, C. 1964. Problems of polymorphism in fishes.Copeia, 1964:65·75.Kosswig, C. 1965. Gcnetique et evolution regressive. Rev.Quest. Sci., 136:227·257.Kosswig, C. 1967. Uber das Tempo evolutorischer Prozesse.ZooI. Reitr., 13:441·450.Kosswig, C. 1973. The role of fish in research on genetics andevolution. Pp. 3-16 in ./. H. Schroder, cd., Genetics andmutagenesis of fish. New York: Springer.Verlag.Kosswig, C., and N. Peters. 1967. Die Evolution der Hohlentiere.Bild der Wissenschaft, 1967:828-335.Kuhn, O. 1960. Ausseroptische Lichtwirkungen bci tier·ischen Organismen. Stud. Gen., 13:477·491.Kuhn, 0., and J. Kiihling. 1954. Augenruckbildung undLichtsinn bei Anoptichthys jordani Hubbs und Innes.Experientia, 10:385·388.Kuhn, 0., and E. Strotkoetter. 1967. Untersuchungen vonDrucken auf den tierischen Korper. I. Druckreceptionbei Fischen und ihre Mitwirkung bei der Orientierungim Raum. Forschungsberichte des Landes Nordrhein·Westfalen, 1857. 57 pp.Lacourt, A. W.1968. A monograph of the freshwater Hryo7.0a.Phylactolaemata. Zool. Verhandl., 93.159 pp.Ladiges, W. 1951a. Hlinde Hohlenfische im Aquarium. DieAqu. U. Terr. Z., 4:258-260.Ladiges, W. 1951b. Der Fisch in der Landschaft: Beitriigc zurOkologie der Susswasserfische, 2nd ed. Braunschweig:Verlag Gustav Wenzel und Sohn. 182 pp.Land, H. 1968. Mayan cave discoveries. Birds. Explorers J.,46: 180·182.Laneyrie, R. 1971. Catalogue des Trechinae cryptiques. Ann.SpeleoL, 26:189·194.Lanham, U. 1962. The fishes. New York: Columbia Univ.Press. 116 pp., 24 pis.Larson,B.1975.Caving in the I-Iuixtan Area, Chiapas. The Huix·Resurgence (Mapachero). Canadian Caver, 7(1 ):28·30.272

LaVal, R. K. 1973. A revision of the neotropical bats of thegenus Myotis. Nat. Hist. Mus. Los Angeles Co. Sci. Bull.,15.54 pp.Lay, D. M. 1962. Seis mam{feros nuevos para la fauna deMexico. Anal. Inst. BioI., Mexico, 33:373-377.Lee, R. 0.1966. Through Quintana Roo 1965. ExplorersJ.,44:83-93.Lemos de Castro, A. 1964. Trichorhina heterophthalma,nueva especie de iwpodo terrestre cavernicola de Cuba.Poeyana, ser. A, 2. 7 pp.Lent, H., and P. Wygodzinsky. 1979. Revision of the Triatominae(Hemiptera, Reduviidae), and their significance asvectors of Chagas' disease. Bull. American Mus. Nat. Hist.,163:123-520.Leroy, Y. 1967. Gryllides et gryllacrides cavernicoles. Ann.Speleol.,22:659-722.Levinson, S. A. 1959. Bibliography and index to new generaand species of Ostracoda for 1957. Micropaleontology,5:241-260.Li, W.-H., and M. Nei. 1977. Persistence of common allelesin two related populations or species. Genetics, 86:901914.Long, C. A., and C. J . .Jones. 1966. Variation and frequenceof occurrence of the baculum in a population of Mexicanfree-tailed bats. Southwestern Nat., 11 :290-295.Longley, G., and II. Karnei,.J r. 1979a. Status of Trogloglanispattersoni Eigenmann, the toothless blind cat. UnitedStates Fish and Wildlife Service, Endangered SpeciesReport, 5( 1). vi + 54 pp.Longley, G., and H. Karnei, .Jr. 1979b. Status of Satan eurystomusHubbs and Bailey, the widemouth blindcat. Uni·ted States Fish and Wildlife Service, Endangered SpeciesReport, 5(2). vi + 48 pp.Longsworth, P. 1959. Exploring caves. New York: ThomasY. Crowell Co. 175 pp.Loomis, H. F. 1953. New millipeds of the western states andLower California. J. Washington Acad. Sci.,43:417-422.Loomis, H. F. 1962. Two unusual Central American spirostreptidmilliped species. Proc. BioI. Soc. Washington,75:47·51.Loomis, H. F. 1968. A checklist of the millipeds of Mexico andCentral America. Bull. United States NatI.Mus., 266.137pp.Loomis, H. F., and R. L. Hoffman. 1962. A remarkable newfamily of spined polydesmoid Diplopoda, including aspecies lacking gonopods in the male sex. Proc. BioI. Soc.Washington, 75: H5-158.L[opezl-Forrnent, W., U. Schmidt, and A. M. Greenhall. 1971.Movement and population studies of the vampire bat(Desmodus rotundus) in Mexico. J. Mammal., 52:227-228.Lopez Ortiz, R. 1962. Geologia y posibilidades petroleras delos sedimentos Cretacicos en la parte sureste del frente dela Sierra Madre de Chiapas. 1301. Asoc. Mexicana Geol.Petrol., 14: 135·151, 13 figs.Lopez Ramos, E. 1969. Geologia del sureste de Mexico ynorte de Guatemala. Publ. Geol. Inst. CentroamericanoInvestigacion y Tecnol. Industrial, 2:57-67.Lord, P. 1974. Sumidero Yochip, e'en ulis, and some others.Canadian Caver, 6(2):13.17.Louch, B., and J. Fish. 1970. Retreat to Orizaba. CanadianCaver, no. 3:19-23.Love, R. M. 1970. The chemical biology of fishes. London:Academic Press. xv + 547 pp.Lozano Romen, F. 1955. Bosquejo geologico de la Provinciadel Papaloapi"m, estado de Veracruz, Mexico. Bol. Asoc.Mexicana Geol. Petrol., 7:1-68,8 figs.Luling, K. H. 1953a. Die Heirnat und die Entdeckung unseresBlindfisches Anoptichthys jordani. Aquar. Terrar. Z.,6:314-318.Luling, K. H. 1953b. Uber das Sehenjugendlicher Anoptichthysjordani (Hubbs und Innes). Die Aqu. u. Terr. Z.,6:62-65.Luling, K. H. 1953c. Uber die fortschreitende Augendegenerationdes Anoptichthys jordani; Hubbs und Innes (Characidae).Zool. Anz., 151:289·299.Luling, K. H. 1954a. Untersuchungen am Blindfisch Anoptichthysjordani Hubbs und Innes (Characidae). EinigeBeobachtungen uber das Verhalten des BlindfischesAnoptichthys jordani beim Laichen. Naturwiss. Rdsch.Stuttgart, 7:197-203.Luling, K. H. 1954b. Untersuchungen am Blindfisch Anop·tichthys jordani Hubbs und Innes (Characidae). II. Beo·bachtungen und Experirnente an Anoptichthys jordanizur Prufung der Einstellung zum Futter, zum Licht undzur Wasserturbulenz. Zool. Jb. (Abt. Allgemeine Zool.Physiol. Tiere), 65 :9-42.Luling, K. H. 1955a. On the subject of eye reduction foundin the blind fish Anoptichthys Jordani (Hubbs and Innes)from the Mexican caves. Photogr. u. Forsch., 6: 138-143.Luling, K. H. 1955b. Untersuchungen am Blindfisch Anoptichthysjordani Hubbs und Innes (Characidae). III. Vergleichendanatomisch-histologische Studien an den Augendes Anopticthys jordani. Zool. Jb. (Abt. Anal. u.Ontog.),74:401-477.Luling, K. H. 1955c. Zur Augenreduktion des aus mexikanischenHohlen stammenden blinden Salmlers Anop·tichthys jordani (Hubbs und Innes). Photogr. u. Forsch.,6:138-143.Luling, K. H. 1957. Caecobarbus geertsi Boulenger-seineEntdeckung, Verbreitung und Lebensweise-und nochmalsAnoptichthys jordani Hubbs und Innes. Aquar.Terrar. Z., 10:145·150.Luling, K. H. 1962. Untersuchungen am Blindfisch Anoptichthysjordani Hubbs und Innes (Characidae). IV. Bemerkungenzur Okologie und Tiergeographie. BonnerZool. Beitr., 13:139-145.Lukens, P. W., .Jr., and W. B. Davis. 1957. Bats of the Mexicanstate of Guerrero. J. MammaL, 38: 1-14.Lundelius, E. 1., and B. H. Slaughter, eds. 1971. Naturalhistory of Texas caves. Dallas: Gulf Nat. Hist. 174 pp.Lynch, .J. D. 1965. Two new species of Eleutherodactylusfrom Mexico (Amphibia: Leptodactylidae). Herpetologica,20:246-252.Lynch, J. D. 1967. Synonymy, distribution and variation inEleutherodactylus decoratus of Mexico (Amphibia: Leptodacylidae).Trans. Illinois State Acad. Sci., 60:299-304.Lynch, .J. D. 1970. A taxonomic revision of the leptodactylidfrog genus Syrrhophus Cope. Univ. Kansas Mus. Nat. Hist.Publ.,20:1-45.Maccagno, T. P., and B. Cucchiari. 1957. Revisione dellePalaemoninae del Museo di Torino. Boll. Inst. Mus. Zool.Univ. Torino, 5 :201-369.McDaniel, V. R., and K. 1. Smith. 1976. Cave fauna ofArkansas: Selected invertebrate taxa. Arkansas Acad. Sci.Proc., 30:57-60.McEachern, M. 1974. Another histoplasmosis cave. Assoc.Mexican Cave Stud. News., 4: 141-142.MacGillivray, A. D. 1891. A catalogue of the Thysanoura ofNorth America. Canadian Entomol., 23:267-276.?73

McKenzie, D. 1965a. Caves of the Sierra de El Abra. Part III.Tamuin and EI Pujal, S. L. P. Assoc. Mexican Cave Stud.News., 1:34-41, 2 maps.McKenzie, D. 1965b. Trip report: 19-25 January 1965.Assoc. Mexican Cave Stud. News., 1:23-27, map.McKenzie, D., and B. Russell. 1965. Caves of the Sierra de ElAbra. Part II. Los Sabinos, S. L. P. Assoc. Mexican CaveStud. News., 1:28-33, map.Magniez, G. 1972. Deux Stenasellidae cavernicoles nouveauxde l'Amerique centrale: Mexistenasellus parzefalli n. sp. etMexistenasellus wilkensi n. sp. (Crustacea Isopoda Asellota).Internat!. J. Speleo!., 4:19-31.Magniez, G. 1973. Description du male de Mexistenasellusparzefalli (Crustacea Isopoda Asellota cavernicole duMexique) et observations sur cette espece. Internatl. J.Speleol., 5 :163-170.Magniez, G. 1974a. Donnees faunistiques et ecologiques surles Stenasellidae (Crustacea Isopoda Asellota des eauxsouterraines). Internatl. J. Speleol., 6:1-80.Magniez, G. 1974b. Observations sur Stenasellus virei dans sesbiotopes naturels (Crustacea Isopoda Asellota des eauxsouterraines). Internatl. J. Speleol., 6: 115-171.Magniez, G. 1979. Les stenasellides de France (Crustaceslsopodes Asellotes souterrains): Faune ancienne et peuplementsrecents. Bull. Soc. ZooI. France, 103:255-262.Malaga Alba, A., and B. Villa R. 1957. Algunas notas acercade la distribucion de los murcielagos de America delNorte relacionados con el problema de la rabia. Anal. Inst.BioI., Mexico, 27:529-569.Maldonado.Koerdell, M., ed. 1956. Estratigrafia del CenOzoicoy del Mesozoico a 10 largo de la carretera entreReynosa, Tamps. y Mexico, D. F. Tectonica de la SierraMadre Oriental. Vulcanismo en el Valle de Mexico. Congr.Geol. Internac., 20th Sesion, Excursiones A-14 y C-6.325 pp., 15 figs., 9 tables.Maldonado-Koerdell, M.1964. Geohistory and paleogeog.raphy of Middle America. Pp. 3-32 in R. C. West, ed.,Handbook of Middle American Indians, Volume One:Natural environment and early cultures. Austin: Univ.Texas Press.Mannix, D. P. 1947. Monsters in minature. Collier's, 119(Jan. 11. 1947):26-28,59.Marden, L. 1959. Dzibilchaltun: Up from the well of time.Natl. Geogr. Mag., 115:110-129.Marquez Mayaudon, C., and J. Ramos Elorduy de Conconi.1974. Un nuevo ricinulideo del genero Cryptocellus Westwoodpara la fauna de Mexico (Arthropoda, Arachnida).J. Arachnol., 1:73-84.Marshall, N. B., and G. L. Thines. 1958. Studies of the brain,sense organs and light sensitivity of a blind cave fish(Typhlogarra widdowsoni) from Iraq. Proc. Zool. Soc.London.. 131:441-456, pI.Marshall, R. 1936. Hydracarina from Yucatan. Carnegie Inst.Washington Publ., 457:133-137.Martin, M., and P. S. Martin. 1954. Notes on the capture oftropical bats at Cuevo el Pachon, Tamaulipas, Mexico.J. Mammal., 35:584-585.Martin, P. S. 1958. A biogeography of reptiles and amphibiansin the Gomez Farias region, Tamaulipas, Mexico. Mus.Zool., Univ. Michigan, Misc. Publ., 101. 102 pp., 7 pIs.Martin, P. S., C. R. Robins, and W. B. Heed. 1954. Birds andbiogeography of the Sierra de Tamaulipas, an isolatedpine-oak habitat. Wilson Bull., 66:38-57.Martinez, L. 1941. Tercera contribucion acerca de la hemato·metria de los murcielagos mexicanos. Anal. Inst. BioI.,Mexico, 12:1-5.Martinez, L., and B. Villa R. 1940. Segunda contribucion alconocimiento de los murcielagos mexicanos. ll.-Estadode Guerrero. Anal. Inst. BioI., Mexico, 11:291-361.Martinez del Rio, P. 1956. Investigaciones anteriores y extensionde la cultura. Pp. 16-24 in L. Aveleyra Arroyo deAnda, M. Maldonado.Koerdell, and P. Martinez del Rio,Cueva de la Candelaria, Volume 1. Mem. Inst. Nac. Antro·pol. Hist., Mexico, 5.Massoud, Z., and M. Gruia. 1973. Collemboles Arthropleonesde Cuba recoltes en 1969 par la mission cubano·roumaine.Resultats des Expeditions Biospeologiques Cubano.Roumaines aCuba, 1: 327·343.Mateu, J. 1977. Sobre algunos linajes de carabidos borco.montanos de Mexico y sus relaciones con el poblamientoentomologico del Sistema Volcanico Transversal. Rev.Soc. Mexicana Hist. Nat., 35:181-224.Mateu, J. 1978. Speleodesmoides rave/oi, nuevo genero yespecie de carahido troglobio en una cueva de Venezuela(Coleoptera: Carabidae). Bol. Soc. Venezolana Espeleol.,9(17):21·28.Mattheij, J. A. M. 1968a. The ACTH cells in the adenohypophysisof the Mexican cave fish Anoptichthys jordani,as identified by metopirone (SU 4885) treatment.Z. Zellforsch., 92:588·595.Mattheij,.T. A. M. 1968b. The cell types in the adenohypophysisof the blind Mexican cave fish, Anoptichthys jordani(Hubbs and Innes). Z. Zellforsch., 90:542·553.Mattheij, J. A. M. 1969. The thyrotropin secreting basophilsin the adenohypophysis of Anoptichthys jordani. Z.Zellforsch., 101:588·597.Mattheij, J. A. M. 1970a. The function of the basophilic cellsin the meso-adenohypophysis of the blind Mexican cavefish Anoptichthys jordani. (Abstr.). J. Endocrinol., 48:lxix.Mattheij, J. A. M. 1970b. The gonadotropic cells in theadenohypophysis of the blind Mexican cave fish, Anoptichthysjordani. Z. Zellforsch., 105 :91·106.Mattheij, J. A. M., and J. A. P. Sprangers. 1969. The site ofprolactin secretion in the adenohypophysis of the steno·haline teleost Anoptichthys jordani, and the effects ofthis hormone on mucous cells. Z. Zellforsch., 99:411-419.Mattheij, J. A. M., J. A. P. Sprangers, and P. G. W. J. vanOordt. 1969. The site of prolactin synthesis in the pituitarygland of Anoptichthys jordani and the influence ofthis hormone on mucous cells. (Abstr.). Gen. CompoEndocrinol., 13:519·520.Mattheij, J. A. M., and P. G. W. J. van Oordt. 1967. The celltypes in the adenohypophysis of the blind Mexican cavefish Anoptichthys jordani. (Abstr.). Gen. Compo Endocrinol.,9:472.Mautz, W. J., and W. Lopez-Forment. 1978. Observations onthe activity and diet of the cavernicolous lizard Lepidophymasmithii (Sauria: Xantusiidae). Herpetologica,34:311-313.Mazzotti, L. 1940. Ornithodoros coprophilus Mc.Intosh, enel estado de Chiapas, Mexico. Ciencia, Mexico, 1:405-406.Mazzotti, L. 1941. Ornithodoros dyeri en Mexico. Medicina,21:313·314.Meder, E. 1952. Uber den blinden Hohlenfisch Anoptichthysjordani Hubbs & Innes. Aus der Arbeitsgemeinschaft:Naturziichtung im VDA. Die Aqu. U. Terr. Z., 5:171-174.Mees, G. F. 1962. The subterranean freshwater fauna ofYardie Creek Station, North West Cape, Western Australia.J. Roy. Soc. Western Australia, 45 :24·32.274

Mercer, H. C. 1896. The hill-caves of Yucatan. Philadelphia:J. B. Lippincott Co. 183 pp.Migdalski, E. C., and G. S. Fichter. 1976. The fresh & saltwater fishes of the world. New York: Alfred A. Knopf.316 pp.Mills, H. B. 1938. Collembola from Yucatan caves. CarnegieInst. Washington Pub!., 491:183-190.Minckley, W. L. 1969. Environments of the Bolson of CuatroCienegas, Coahuila, Mexico, with special reference to theaquatic biota. Univ. Texas at EI Paso Sci. Ser., 2.65 pp.Minckley, W. L., and G. A. Cole. 1968. Speocirolana thermydronis(Crustacea: Isopoda) from northeast Mexico, rediscovery,habitat, and supplemental description. TulaneStud. Zool. Bot., 15:2-4.Minton, M. 1978. The Diamante Story. Four years under theEI Abra. Assoc. Mexican Cave Stud. Activities News.,8:6-15, map.Mitchell, H. A. 1965. Investigations of the cave atmosphereof a Mexican bat colony. J. Mamma!., 45:568-577.Mitchell, R. W. 1968. Typhlochactas, a new genus of eyelesscave scorpion from Mexico (Scorpionida, Chactidae).Ann. Speleol., 23:753-777,1 pI.Mitchell, R. W. 1969a. A comparison of temperate and tropicalcave communities. Southwestern Nat., 14:73-88.Mitchell, R. W. 1969b. The cover illustration, Cryptocellusosorioi, (Arachnida; Ricinulei). Southwestern Nat., 14:136-138.Mitchell, R. W. 1969c. La region de la Sierra de El Abra. Mapand Letter. Privately printed.Mitchell, R. W. 1970a. Population size and dispersion andspecies associations of a Mexican cavernicole ricinuleid(Arachn.). Ciencia, Mexico, 27:63-74.Mitchell, R. W. 1970b. Total number and density estimatesof some species of cavernicoles inhabiting Fern Cave,Texas. Ann. SpeIeo!., 25 :73-90.Mitchell, R. W. 1971a. Egg and young guarding by a Mexicancave-dwelling harvestman, Hoplobunus boneti (Arachnida).Southwestern Nat., 15 :392-394.Mitchell, R. W. 1971b. Typhlochactas elliotti, a new eyelesseave scorpion from Mexico (Scorpionida, Chactidae).Ann. Speleol., 26:135-148.Mitchell, R. W. 1973. Introgression between the Mexicaneyeless characin fishes and their epigean ancestor. (Abstr.).P. 146 in V. Panos, ed., International Speleology 1973.Abstracts of papers. 6th Internatl. Congr. Speleol., 010moue, Czechoslovakia.Mitchell, R. W. 1975. Distributional studies of the Mexicaneyeless characin fishes. American Phil. Soc. Year Book,1974:360-361.Mitchell, R. W. 1977. Introgression between the Mexican eyelesscharacin fishes and their epigean ancestor, Astyanaxmexicanus. Proc. 6th Internat!. Congr. Speleol., 5:171173.Mitchell, R. W., and J. W. Cooke. 1973. Preliminary morphometriccomparisons of several populations of Mexicaneyeless characin fishes. (Abstr.). P. 147 in V. Panos, ed.,International Speleology 1973. Abstracts of papers. 6thInternat!' Congr. Speleo!., Olomouc, Czechoslovakia.Mitchell, R. W., and J. W. Cooke. 1977. Preliminary morphometriccomparisons of several populations of Mexicaneyeless characin fishes of the genus Astyanax. Proc. 6thInternat!. Congr. Speleol., 5:175-178.Mitchell, R. W., and W. R. Elliott. 1973. The cave habitats ofthe Mexican eyeless characin fishes. (Abstr.). P.147 in V.Panos, ed., International Speleology 1973. Abstracts ofpapers. 6th Internat!' Congr. Speleo!., Olomouc, Czechoslovakia.Mitchell, R. W., and W. R. Elliott. 1977. The habitats of theeyeless characin fishes of the genus Astyanax. Proc. 6thInternat!' Congr. Speleo!., 5:179·184.Mitchell, R. W., and M. Kawakatsu. 1972. A new family,genus, and species of cave-adapted planarian from Mexico(Turbellaria, Tricladida, Maricola). Occ. Papers Mus.Texas Tech Univ., 8.16 pp.Mitchell, R. W., and M. Kawakatsu. 1973a. Freshwater cavernicoleplanarians from Mexico: New troglobitic and troglophilicDugesin from caves of the Sierra de Guatemala.Ann. Speleol., 27:639-681.Mitchell, R. W., and M. Kawakatsu. 1973b. A new caveadaptedplanarian (Tricladida, Paludicola, Planariidae)from Chiapas, Mexico. Assoc. Mexican Cave Stud. Bull.,5:165-170.Mitchell, R. W., and S. B. Peck. 1978. Typhlochactas sylvestris,a new eyeless scorpion from montane forest litter inMexico (Scorpionida, Chactidae, Typhlochactinae). J.Arachno!., 5: 159-168.Mitchell, R. W., and J. R. Reddell. 1971. The invertebratefauna of Texas caves. Pp. 35-90 in E. L. Lundelius, Jr.,and B. H. Slaughter, eds., Natural history of Texas caves.Dallas: Gulf Nat. Hist.Mitchell, R. W., and J. R. Reddell, eds. 1973. Studies on thecavernicole fauna of Mexico and adjacent regions. Assoc.Mexican Cave Stud. Bull., 5.201 pp.Mitchell, R. W., and W. H. Russell. 1973. Physiography andgeology of the Huastecan Province of Mexico. Proc. 6thInternat!' Congr. Speleo!., 2:261-268.Mitchell, R. W., and W. H. Russel!. 1976a. Stream capture inthe Huastecan Province of Mexico. Proc. 6th Internat!.Congr. Speleo!., 4: 169-172.Mitchell, R. W., and W. H. Russell. 1976b. The subsurfacewaters of the Sierra de EI Abra of Mexico. Proc. 6thInternat!' Congr. Speleol., 4:163-167.Mitchell, R. W., W. H. Russell, and W. R. Elliott. 1977. Mexicaneyeless characin fishes, genus Astyanax: Environment,distribution, and evolution. Spec. Pub!. Mus. Texas TechUniv., 12. 89 pp., map.Miyashita, M., H. Tanaka, A. Shirasaka, and K. Ichirnori.1977. Studies on the biological control on an intermediatehost of Trematoda by tropical fishes. Japanese J.Sanitary Zoo!., 28:291-300.Mockford, E. L. 1967. The electrentomoid psocids (Psocoptera).Psyche, 74: 118-165.Mohr, C. E. 1950. Mexican blind cave fish subject of importantresearch on evolution and cancer. Nat!' Speleo!.Soc. News, 8(4):5.Mohr, C. E. 1968. Mayan cave discoveries. Bats. ExplorersJ.,46:172-179.Mohr, C. E., and T. L. Poulson. 1966. The life of the cave.New York:McGraw-Hill Book Co. 232 pp.Moller, H. 1970. El interminable territorio subterraneo. Finde Semana, Oct. 2, 1970, p. 5.Mollhagen, T. 1971. Checklist of bats in caves in the regionsof the Sierra de Guatemala and Sierra de El Abra, north.eastern Mexico. Assoc. Mexican Cave Stud. Bull., 4:19·22.Mollhagen, T. 1977. Sierra del Burro roadlog. Assoc. MexicanCave Stud. News., 5:79~1.Mones, A. 1971. Restos oseos de mamfferos contenidos enregurgitaciones de lechuza del estado de Oaxaca, Mexico.Anal. Inst. Bio!. Univ. Na!. Auton., Mexico, 39, Ser.Zoo!., (1):169-171.275

Monod, T. 1975. Sur la distribution de quelques crustacesmalacostraces d'eau douce ou Saumatre. Mem. Mus. I\atl.Hist. Nat., ser. A (Zool.), 88:98·105.Monod, T., and P. Cals. 1970. Sur une espece nouvelle decrevette cavemicole: Typhlatya galapagensis (DecapodaNatantia; Atyidae)..1ission Zoologique BeIge aus liesGalapagos et en Ecuador, 2:57·103.Moor, A. 1980. Stratigraphy and structure of Potosi Anti·cline, Nuevo Leon, Mexico. M. A. Thesis. Austin: Univ.Texas at Austin. x + 116 pp., 1 pI.Moore, G. W., and G. Nicholas. 1964. Speleology: The studyof caves. Boston: D. C. Heath and Co. vii + 120 pp.Moore, G. W., and G. Nicholas. 1978. Speleology: The studyof caves, rev. 2nd ed. Teaneck, New Jersey: ZephyrusPress. xiii + 150 pp.Moore, J. P. 1936. Hirudinea from Yucatan. Carnegie Inst.Washington Publ., 457:41-43.Moore, J. P. 1938. Leeches (Hirudinea) from Yucatan caves.Carnegie Inst. Washington Publ., 491:67.70.Morris, N. 1971. Trip report: 13·19 March, 1971. Assoc.Mexican Cave Stud. News., 3:44·45.Morris, R. A. 1969. Blind cave fishes. Anim. King., 72(3):26·28.Mothes, P. 1978. San Joaquin: A cumulaitive report. Assoc.Mexican Cave Stud. Activities :\ews., 8 :63-67.Muchmore, W. B. 1969. A cavernicolous Tyrannochthoniusfrom Mexico (Arachn., Chelon., Chthon.). Ciencia,Mexico, 27:31·32.Muchmore, W. B. 1972a. :\lew diplosphyronid pseudoscor.pions, mainly cavernicolous, from :Vlexico (Arachnida,Pseudoscorpionida). Trans. American ;\licroscop. Soc.,91 :261-276.Muchmore, W. B. 1972b. The pseudoscorpion genus Para·liochthonius (Arachnida, Pseudoscorpionida, Chthonii·dae). Entomol. News., 83:248·256.Muchmore, W. B. 1972c. The unique, cave·restricted genusAphrastochthonius (Pseudoscorpionida, Chthoniidae).Proc. BioI. Soc. Washington, 85:433-444.Muchmore, W. B. 1973a. New and little known pseudo.scorpions, mainly from caves in Mexico (Arachnida,Pseudoscorpionida). Assoc. Mexican Cave Stud. Bull.,5:47-62.Muchmore, W. B. 1973b. The pseudoscorpion genus Mexo·bisium in Middle America (Arachnida, Pseudoscorpionida).Assoc. Mexican Cave Stud. Bull., 5 :63-72.Muchmore, W. B. 1973c. A second troglobitic Tyrannochthoniusfrom Mexico (Arachnida, Pseudoscorpionida,Chthoniidae). Assoc. Mexican Cave Stud. Bull., 5 :81-S2.Muchmore, W. B. 1976. Aphrastochthonius pachysetus, anew cavernicolous species from New Mexico (Pseudo·scorpionida, Chthoniidae). Proc. BioI. Soc. Washington,89:361-364.Muchmore, W. B. 1977. Preliminary list of the pseudoscor·pions of the Yucatan Peninsula and adjacent regions, withdescriptions of some new species (Arachnida: Pseudoscorpionida).Assoc. Mexican Cave Stud. Bull., 6:63-78.Muir, J. M. 1936. Geology of the Tampico Region, Mexico.London: Thomas Murby & Co. xix + 280 pp., 15 pis.Mulaik, S. B. 1960. Contribucion aI conocimiento de losisopodos terrestres de Mexico (Isopoda, Oniscoidea). Rev.Soc. Mexicana His!. Nat., 21 :79·292.MuIlinex, C. L. 1975. Revision of Paraphrynus Moreno(Amblypygida: Phrynidae) for North America and theAntilles. Occ. Papers California Acad. Sci., 116.80 pp.MuIlinex, C. L. 1979. A new Paraphrynus from Yucatan(Amblypygida, Tarantulidae). J. Arachnol., 7 :267·269.Nagle, F., J. Rosenfeld, and J. J. Stipp. 1977. Guatemala,where plates collide. A reconnaissance guide to Guate.malan geology. Miami, Florida: Miami Geol. Soc. 72 pp.,map.Nath, C. N., and N. K. Pillai. 1971. Studies on the genusLepidomysis Clarke. Crustacea: Mysidacea. Hydrobiologica,37:285·300.Nath, C. N., D. M. Thampy, and N. K. Pillai. 1972. Opticregression in a subterranean mysid (Crustacea, Mysi·dacea). Internat!. J. Speleol., 4:51-54.Neill, W. T., and R. Allen. 1962. Reptiles of the CambridgeExpedition to British Hondruas, 1959-60. Herpetologica,18:79-91.Nelson, E. W. 1899. Mammals of the Tres Marias Islands.North American Fauna, 14:15·19.Nesbitt, 1-1. H. J. 1949. Six new Mexican mites of the subfamilyRhizoglyphinae (Acarina). Pan-Pacific Entomol.,25:57·70.Nicholas, G. 1960. Checklist of macroscopic troglobiticorganisms of the United States. American MidI. ;\jat.,64:123-160.Nicholas, G. 1962. Checklist of troglobitic organisms ofMiddle America. American MidI. Nat., 68:165-188.Nicholas, G. 1968. Mayan cave discoveries. Fauna. ExplorersJ., 46: 168·171.Nicholas, G. 1976. Cave biology. Pp. 42·44 in R. F. Dalton,Caves of ew Jersey, New Jersey Bur. Ccol. Topogr.Bul!.,70.Nigrelli, R. F. 1947. Spontaneous neoplasms in fishes. III.Lymphosarcoma in Astyanax and Esox. Zoologica, NewYork, 32:101-108, pis. 1-11.Norman, J. R. 1963. A history of fishes, 2nd ed. by P. II.Greenwood. New York: Hill and Wang. xxxi + 398 pp.Oguri, M., and Y. Omura. 1973. Ultrastructure and functionalsignificance of the pineal organ of teleost. Pp. 412·434in W. Chavin, ed., Responses of fish to environmentalchanges. Springfield, Illinois: Charles C. Thomas.Olivas, M. 1956. Geologia a 10 largo dc la carretera CristobalColon, entre Tuxtla Gutierrez, Chis. y Oaxaca, Oax.Pp. 3-53, figs. 1-9 in M. Maldonado.Kocrdcll, cd., Geolo·gia a 10 largo de la carrelera entre Tuxtla Gutierrez, Chis.y Mexico, D. F., y visita a monumentos precoloniales deOaxaca, Oax. Congl. Geol. Internac., 20th Sesion, ExcursionC·15B.Olivereau, M., and M. Francotte-Henry. 1955. Etude histologiqueet biometrique de la glande thyroide de Caeca·barbus geertsi Blgr. Ann. Soc. Roy. Zool. Belgique, 86:129-150.Olivereau, M., and M. Herlant. 1954. Etude histologique del'hypophyse de Caeeobarbus geertsii Blgr. Acad. Roy.Belgique Bull., Classe Sci., sel. 5,40:50-57.Orghidan, T., A. unez Jimenez, L. Botosaneanu, V. Decou,S. Negrea, and N. Villa Bayes, eds. 1973. Resultats desExpeditions Biospeologiques Cubano-Roumaines a Cuba,1. Bucuresti: Editura Academiei Republicii SocialisteRomania. 424 pp., 28 pIs.Orghidan, T., A. Nunez Jimenez, V. Decou, S. Negrea, N.Villa Bayes, eds. 1977. Resultats des Expeditions Biospeo.logiques Cubano·Roumaines aCuba, 2. Bucuresti: EdituraAcademiei Republicii Socialiste Romania. 420 pp., 15 pis.Osorio Tafall, B. F. 1942. Diaptomus (Mierodiaptomus)eokeri, nuevos subgenero y especie de diaptomido de las276

cuevas de la region de Valles (San Luis Potosi, Mexico).(Copep., Calan.). Ciencia, Mexico, 3:206-210.Osorio Tafall, B. F. 1943. Observaciones sobre la faunaacuatica de las cuevas de la region de Valles, San LuisPotosi (Mexico). Rev. Soc. Mexicana Hist. Nat., 4:43-71.Osorio Tafall, B. F. 1946. Anotaciones sobre algunos aspec·tos de la hidrologia mexicana. Rev. Soc. Mexicana Hist.Nat., 7:139-165.Pace, N. 1977. The survey of Salida Cruz Pila!. CanadianCaver, 9(2):7-10.Packard, A. S. 1888. The cave fauna of North America, withremarks on the anatomy of the brain and origin of theblind species. Mem. Nat!. Acad. Sci., 4. 156 pp., 27 pis.Packard, A. S. 1894. On the origin of the subterranean faunaof North America. American Nat., 28:727·751.Packard, A. S. 1900a. A new eyeless isopQd crustacean fromMexico. Science, n. ser., 12:300-301.Packard, A. S. 1900b. A new eyeless isopod crustacean fromMexico. Proc. American Assoc. Adv. Sci., 49:228.Paclt, J. 1957. Diplura. Genera Insectorum, 212e. 123 pp.Paclt, J.1963. Thysanura. Fam. Nicoletiidae. Genera Insectorum,216e. 58 pp.Paclt, J. 1971. Occurrence of a new genus of troglobiticNicoletiidae (Ins., Thysanura) in Mexico. Internat!. J.Speleo!., 3:423-424, p!. 127.Padilla y Sanchez, R. J. 1978. Bosquejo geologico-estructuralde la Sierra Madre Oriental en el area Linares-Galeana-SanRoberto, estado de Nuevo Leon. Univ. Nac. Aut. Mexico,Inst. Geo!., Rev., 2(1 ):45-54.Papp, L. 1978. Some cavernicolous Diptera of the GenevaMuseum. Rev. Suisse Zoo!., 85 :99-106.Parzefall, J. 1969. Zur vergleichenden Ethologie verschiede.ner Mollienesia-Arten einschliesslich einer HohlenfonnvonM. sphenops. Behaviour, 33:1·37.Parzefall, J. 1970. Morphologische Untersuchungen an einerHohlenform von Mol/ienesia sphenops (Pisces, Poeciliidae).Z. Morph. Tiere, 68:323-342.Parzefall, J. 1973a. Attraction and sexual cycle of poeciliids.Pp. 177-183 in J. H. Schroder, ed., Genetics and muta·genesis of fish. New York: Springer-Verlag.Parzefall, J. 1973b. Modifications du comportement de Poeciliasphenops (Pisces, Poeciliidae) comicutives a sonpassage a la vie cavernicole. Ann. SpeIeo!., 28:283·290.Parzefall, J. 1976. Die Rolle der chemischen Information imVerhalten des Grottenolms Proteus anguineus Laur. (Protcidae,Urodela). Z. Tierpsycho!., 42:29-49.Parzefall, J. 1979. Zur Genetik und biologischen Bedeutungdes Agressionsverhaltens von Poecilia sphenops (Pisces,Poeciliidae). Untersuchungen an Bastarden ober- undunterirdisch Iebender Populationen. Z. Tierpsycho!., 50:399·422.Parzefall, J., and H. Wilkens. 1972. Artbildung bei Hohlenfischen.Vergleichende Untersuchungen an zwei amerikanischenSynbranchiden (Pisces, Teleostei). Z. Morph.Tiere, 73:63-79.Pasquini, P. 1970. Relazioni e conferenze. Rend. Accad. Naz.Lincei, ser. 8, 48:469-471.Pasquini, P. 1977. Subterranean fauna of Mexico. Part Ill.Presentazione. Quad. Accad. Naz. Lincei, Prob!. Att. Sci.Cult., 171(3):3-4.Pate, D. 1979. Sistema Purificacion. 1979 Spring Project.Assoc. Mexican Cave Stud. Activities News., 10:82-101,map.Pavan, C. 1946. Observations and experiments on the cavefish Pimelodella kronei and its relatives. American Nat.,80: 343-361.Pearse, A. S. 1933. Biological research in inland waters ofYucatan. Carnegie Inst. Washington Year Book, 32:109Ill.Pearse, A. S. 1936a. The cenotes of Yucatan. Introduction.Carnegie Inst. Washington Pub!., 457: 1·4.Pearse, A. S. 1936b. Chironomid larvae from Yucatan. CarnegieInst. Washington Pub!., 457:151.Pearse, A. S. 1936c. Parasites from Yucatan. Carnegie Inst.Washington Pub!., 457:45-59.Pearse, A. S. 1936d. Results of survey of the cenotes in Yucatan.Carnegie Inst. Washington Pub!., 457: 17.28, pis. 1-2.Pearse, A. S. 1938a. Fauna of the caves of Yucatan. Introduction.Carnegie Inst. Washington Pub!., 491 :1-17.Pearse, A. S. 1938b. Insects from Yucatan caves. CarnegieInst. Washington Pub!., 491:237-249.Pearse, A. S. 1945. La fauna. Enciclopedia yucatanense,1:109-271.Pearse, A. S., and R. Kellogg. 1938. Mammalia from Yucatancaves. Carnegie Inst. Washington Pub!., 491:301-304.Pearse, A. S., and C. B. Wilson. 1938. Copepoda from Yuca·tan caves. Carnegie Inst. Washington Pub!., 491:153·154.Peck, S. B. 1968. A new cave catopid beetle from Mexico,with a discussion of its evolution. Psyche, 75:91-98.Peck, S. B. 1970. The terrestrial arthropod fauna of Floridacaves. Florida Entomo!., 53:203·207.Peck, S. B. 1971a. The invertebrate fauna of tropical Ameri·can caves, Part I: Chilibrillo Cave, Panama. Ann. Speleo!.,26:423-437.Peck, S. B. 1971b. New and poorly known Ptomaphagusfrom Mexican caves (Coleoptera; Leiodidae; Catopinae).Assoc. Mexican Cave Stud. Bull., 4:9-12.Peck, S. B. 1973a. A review of the cavernicolous Catopinae(Coleoptera; Leiodidae) of Mexico, Belize, and Guate.mala. Assoc. Mexican Cave Stud. Bull., 5:97-106.Peck, S. B. 1973b. A review of the invertebrate fauna of volcaniccaves in western North America. Bull. Nat!. Speleo!.Soc., 35 :99-107.Peck, S. B. 1973c. A systematic revision and the evolutionarybiology of the Ptomaphagus (Adelops) beetles of NorthAmerica (Coleoptera; Leiodidae; Catopinae), with emphasison cave-inhabiting species. Bull. Mus. Compo Zoo!.,145:29·162.Peck, S. B. 1974a. The invertebrate fauna of tropical Americancaves, Part II: Puerto Rico, an ecological and zoogeographicanalysis. Biotropica, 6: 14-3l.Peck, S. B. 1974b. Recent studies on the invertebrate faunaand ecology of sub·tropical and tropical American caves.Assoc. Mexican Cave Stud. News., 5:30-38.Peck, S. B. 1976. The invertebrate fauna of tropical Americancaves, Part III: Jamaica, an introduction. Internat!. J.SpeleO!.,7:303·326.Peck, S. B. 1977a. The subterranean and epigean Catopinaeof Mexico (Coleoptera: Leiodidae). Assoc. Mexican CaveStud. Bull., 6:185-213.Peck, S. B. 1977b. An unusual sense receptor in internalantennal vesicles of Ptomaphagus (Coleoptera: Leiodidae).Canadian Entomo!., 109:81-86.Peck, S. B. 1977c. Visits to some eave and karst biologylocalities in EI Salvador and Mexico in 1971. Assoc. Mexi·can Cave Stud. News., 5:66-67.Peck, S. B., and M. B. Fenton. 1973. The fauna of Canadiancaves. Canadian Caver, 5(2):18·23.277

Peck, S. B., and J. J. Lewis. 1977. Zoogeography and evolutionof the subterranean invertebrate faunas of Illinois andsoutheastern Missouri. Natl. Speleo1. Soc. Bull., 40:39-63.Peck, S. B., and J. H. Peck. 1967. Preliminary survey of theterrestrial invertebrate cave fauna of Alabama. J. AlabamaAcad. Sci., 38:216-217.Peck, S. B., and J. H. Peck. 1973. 1969 Mexico-Guatemalacave biology field trip report. Assoc. Mexican Cave Stud.News., 4:63-70.Pelz, H. W. 1958. Einiges uber den blinden HohlensalmlerAnoptichthys jordani Hubbs und Innes. Aquar. Terrar. Z.,5:172-175.Penney, J. T., and A. A. Racek. 1968. Comprehensive revisionof a worldwide collection of freshwater sponges(Porifera: Spongillidae). United States Nat1. Mus. Bull.,272.184 pp.Peters, G. 1973. Caracteres degeneratifs et constructifs chezune forme cavernicole phylogeniquement recente dePoecilia sphenops (Pisces, Poeciliidae). Ann. Speleo1.,28:315-326.Peters, ., and G. Peters. 1966. Das Auge zweier Hohlenformenvon Astyanax mexicanus (Philippi) (Characinidae,Pisces). Wilhelm Roux Arch. Entwick1.-Mech. Org., 157:393-414.Peters, N., and G. Peters. 1968. Zur genetischen Interpretationmorphologischer Gesetzmiissigkeiten der degernati·ven Evolution. Untersuchungen am Auge einer Hohlenformvon Poecilia sphenops (Poeciliidae, Pisces). Z. Morph.Tiere,62:211-244.Peters, N., and G. Peters. 1973a. Genetic problems in the regressiveevolution of cavernicolous fish. Pp. 187-201 inJ. H. Schroder, ed., Genetics and mutagenesis of fish.New York: Springer-Verlag.Peters, N., and G. Peters. 1973b. Problemes genetiques del'evolution regressive des cavernicoles. Ann. Speleol.,28:301-313.Peters, N., G. Peters, J. Parzefall, and H. Wilkens. 1973. Uberdegenerative und konstruktive Merkmale bei einer phylo.genetisch jungen Hohlenform von Poecilia sphenops(Pisces, Poeciliidae). rnt. Rev. ges. Hydrobio1., 58:417436.Peters, N., A. Scholl, and H. Wilkens. 1975. DerMicos-Fisch,Hohlenfisch in statu nascendi oder Bastard? Ein Beitragzur Evolution der Hohlentiere. Z. Zoo1. Syst. Evolutionsf.,13:110·124.Peterson, B. V., and K. Hurka. 1974. Ten new species of batflies of the genus rrichobius (Diptera: Streblidae). CanadianEntomol., 106: 1049-1066.Pfeiffer, W. 1963. Vergleichende Untersuchungen uber dieSchrekreaktion und den Schreckstoff der Ostariophysen.Z. Verg1. Physio1., 47:111-147.Pfeiffer, W. 1966. Uber die Vererbung der Schrekreaktion beiAstyanax (Char~cidae,Pisces). Z. Vererbungsl., 98:97-105.Pfeiffer, W. 1967a. Die Korrelation von Augengrosse undMittelhirngrosse bei Hybriden aus Astyanax X Anoptich.thys (Characidae, Pisces). Wilhelm Roux Arch. Entwickl.Mech. Org., 159:365-378.Pfeiffer, W. 1967b. Die Korrelation von Korperliinge, Augen.,Linsen· und Pupillengrosse bei Hybriden aus Astyanax XAnoptichthys (Characidae, Pisces). Wilhelm Roux Arch.Entwickl.-Mech. Org., 158:218-245.Pfeiffer, W. 1975. Uber fluoreszierende Pterine aus der Hautvon Cypriniformes (Pisces) und ilire Beziehung zumSchreckstoff. Rev. Suisse Zool., 82:705-711.Pfisterer, R. 1956. Blind cavers. Netherworld News, 4(3).Reprinted in: Speleo Digest, 1956(2):11-12.Pickford, G. E. 1938. Earthworms in Yucatan caves. Carnegielnst. Washington Publ., 491 :71-100, pis. 1-3.Pickford, G. E., and J. W. Atz. 1957. The physiology of thepituitary gland of fishes. New York: New York Zool.Soc. xxiii + 613 pp.Pillai, N. K., and T. Mariamma. 1964. On a new lepidomysidfrom India. Crustaceana, 7: 113-124.Pine, R. H. 1972. The bats of the genus Carollia. Texas A&MUniv., Texas Agric. Expt. Sta. Tech. Monogr., 8.125 pp.Pittard, S. K. 1970. Comparative external morphology of thelife stages of Cryptocellus pelaezi (Arachnida, Ricinulei).M. S. Thesis. Lubbock: Texas Tech Univ. 122 pp.Pittard, S. K., and R. W. Mitchell. 1972. Comparative morphologyof the life stages of Cryptocellus pelaezi (Arachnida,Ricinulei). Grad. Stud. Texas Tech Univ., 1. 77 pp.Poll, M., and N. Leleup. 1965. Un poisson aveugle nouveau dela famille des Brotulidae provenant des lies Galapagos. Bull.Acad. Roy. Belgique (Sci.), 5th sel., 41:464-474, pis. 1-2.Popper, A. N. 1970. Auditory capacities of the Mexicanblind cave fish (Astyanax jordani) and its eyed ancestor(Astyanax mexicanus). Anim. Behav., 18:552-562.Popper, A. N., and W. N. Tavolga. 1967. Hearing thresholdsin the Mexican blind cavefish. (Abstr.). American Zool.,7:791.Post, A. 1965. Vergleichende Untersuchungen der Chromosomenzahlenbei Susswasser-Teleosteern. Z. Zool. Syst.Evolutionsf.,3:47·93.Poulson, T. L. 1961. Cave adaptation in amblyopsid fishes.Ph.D. Diss. Ann Arbor: Univ. Michigan. Ann Arbor:University Microfilms. 191 pp.Poulson, T. L. 1963. Cave adaptation in amblyopsid fishes.American MidI. Nat., 70:257-290.Poulson, T. L. 1964. Animals in aquatic environments: Animalsin caves. Pp. 749-771 in D. B. Dill, ed., Handbookof physiology, sec. 4, Adaptation to the environment.Washington: American Physio1. Soc.Pretzmann, G. 1965. Vorliiufiger Bericht uber die FamiliePseudothelphusidae. Anz. Osterreichischcn Akad. Wiss.Math.-Natur. Klasse, 1965:1-10.Pretzmann, G. 1972. Die Pseudothelphusidae (CrustaceaBrachyura). Zoologica, Stuttgart, 42. iv + 182 pp.Priego de Wit, M. 1974. Geologia y yacimientos minerales delarea de Uera, Tamps. Mexico, D. F.: Consejo de RecursosNaturales no Re!!ovables. [19 pp."jQuaghebeur, M. 1955. Onderzoekingen over blinde grotvissenin verband met de reductie in de ontwikkeling van hunogen. These. Univ. Louvain. 78 pp.Quaghebeur, M. 1957. Onderzoek over de reductie van deogen, van de oogkas en van het tectum opticum bij blindegrotvissen. These. Univ. Louvain. 120 pp.Quinones, F., J. P. Koplan, L. Pike, F. Staine, and L. AjelJo.1978. Histoplasmosis in Belize, Central America. Ameri·can J. Trop. Med. Hyg., 27 :558·561.Rabb, G. B. 1965. A new salamander of the genus Chirop·terotriton (Caudata: Plethodontidae) from Mexico.Breviora, 235. 8 pp.Radovsky, F. J., J. K. Jones, Jr., and C. J. Phillips. 1971.Three new species of Radfordiella (Acarina: Macronyssi·dae) parasitic in the mouth of phyllostomatid bats. J.Med. Entomo1., 8:737-746.Raines, T. W. 1964. EI Sotano del Profesor. Texas Caver,9:113-114.Raines, T. W., ed. 1968a. Sotano de las Golondrinas. Assoc.Mexican Cave Stud. Bull., 2. 20 pp., map, 8 pis.278

Raines, T. W. 1968b. Trip report: 2-4 December 1966. Assoc.Mexican Cave Stud. News., 2:140-145.Raines, T. W. 1972a. Sotanito de Ahuacathin; Sierra MadreOriental; Jalpan; Ahuacatlan. Assoc. Mexican Cave Stud.Cave Rept. Ser., 1. 20 pp., map.Raines, T. W. 1972b. Trip report: 1-3 October 1971. Assoc.Mexican Cave Stud. News., 3:96.Raines, T. W. 1973. Ten years of Mexican caving. Assoc.Mexican Cave Stud. News., 4:4-6.Raisz, E. 1964. Landforms of Mexico. 2nd ed. Cambridge,Massachusetts.Ralph, R. 1979. Stranded on the bottom of Hoya de lasGuaguas. Assoc. Mexican Cave Stud. Activities News.,9:64-71, map.Rambla, M. 1969. Cave harvestmen from Jamaica (Opiliones:Phalangodidae). Psyche, 76:390-406.Ramirez.Pulido, J., and T. Alvarez. 1972. Notas sobre losmurcielagos del genero Leptonycteris en Mexico, con ladesignacion del lectotipo de L. yerbabuenae Martinez yVilla, 1940. Southwestern Nat., 16:249-259.Ramirez-Pulido, J., and C. Sanchez-Hernandez. 1971. Tylomysnudicaudus from the Mexican states of Puebla andGuerrero. J. Mammal., 52:481.Rance, T., and B. J. Baker. 1979. The teleost melaninconcentratinghormone-A pituitary hormone of hypothalamicorigin. Gen. Compo Endocrinol., 37:64-73.Rasquin, P. 1946. Progressive pigmentary regression in fishesassociated with cave environments. (Abstr.). Anat. Rec.,96:578-579.Rasquin, P. 1947. Progressive pigmentary regression in fishesassociated with cave environments. Zoologica, New York,32:35-42, pI. 1.Rasquin, P. 1949a. The influence of light and darkness onthyroid and pituitary activity of the characin Astyanaxmexicanus and its cave derivatives. Bull. American Mus.Nat. Hist., 93:497-531, pis. 20-25.Rasquin, P. 1949b. Regeneration of the optic nerve aftersection with return of vision in the characin Astyanaxmexicanus. Physiol. Zoo!., 22:131-135, pI. 1.Rasquin, P., and E. Hafter. 1951a. Age changes in the testisof the teleost, Astyanax mexicanus. .T. Morpho!., 89:397407.Rasquin, P., and E. I-Iafter. 1951b. Response of a spontaneousfish lymphosarcoma to mammalian ACTH. Zoologica,New York, 36:163-169, pI. 1.Rasquin, P., and L. Rosenbloom. 1954. Endocrine imbalanceand tissue hyperplasia in teleosts maintained in darkness.Bull. American Mus. Nat. Hist., 104:359·425, pIs. 4·23.Reddell, J. R. 1965a. Biology of the caves of the northern EIAbra range. Assoc. \1exican Cave BioI. News., 1: 19-21.Reddell, J. R. 1965b. A checklist of the cave fauna of Texas.I. The Invertebrata (exclusive of Insecta). Texas J. Sci.,17:143-187.Reddell, J. R. 1965c. The status of Mexican cave biology.Assoc. Mexican Cave Stud. News., 1:6-7.Reddell,.T. R. 1965d. Trip report: 14-19 April 1965. Assoc.Mexican Cave Stud. News., 1:48-51.Reddell, J. R. 1966a. The biology of the caves of Rancho delCielo. Assoc. Mexican Cave Stud. News., 2: 12-15.Reddell, J. R. 1966b. A checklist of the cave fauna of Texas.II. Insecta. Texas J. Sci., 18:25·56.Reddell,.T. R.1966c. Trip report: 20 February-2 March 1966.Assoc. Mexican Cave Stud. News., 2:5-8.Reddell, J .. R. 1966d. Trip report: 1-7 June 1966. Assoc.Mexican Cave Stud. News., 2:49-54.Reddell, J. R. 1967a. Cave biology of the Monterrey area.Assoc. Mexican Cave Stud. Bull., 1:24-25.Reddell, J. R. 1967b. Cave biology of the Sierra de EI Abra.Assoc. Mexican Cave Stud. Bull., 1:82-83.Reddell, J. R. 1967c. Cave biology of the Sierra de Guatemala.Assoc. Mexican Cave Stud. Bull., 1:55-56.Reddell, J. R. 1967d. Cave biology of the Xilitla region.Assoc. Mexican Cave Stud. Bull., 1:106-107.Reddell, J. R. 1967e. A checklist of the cave fauna of Texas.Ill. Vertebrata. Texas J. Sci., 19:184-226.Reddell, J. R. 1967f. The order Ricinulei in Mexican caves.Assoc. Mexican Cave Stud. News., 2:99-101.Reddell, J. R. 1970a. A checklist of the cave fauna of Texas.IV. Additional records of Invertebrata (exclusive of Insecta).TexasJ. Sci., 21:389-415.Reddell, J. R. 1970b. A checklist of the cave fauna of Texas.V. Additional records of Insecta. TexasJ. Sci., 22:47-65.Reddell, J. R. 1970c. A checklist of the cave fauna of Texas.VI. Additional records of Vertebrata. Texas J. Sci., 22:139-158.Reddell, J. R.1971a. A checklist of the cave fauna of Mexico.III. New records from southern Mexico. Assoc. MexicanCave Stud. Bull., 4:217·230.Reddell, J. R. 1971b. A preliminary bibliography of Mexicancave biology with a checklist of published records. Assoc.Mexican Cave Stud. Bull., 3. 184 pp.Reddell, J. R. 1973a. Ten years of Mexican cave biology.Assoc. Mexican Cave Stud. ews., 4: 31-43.Reddell, J. R. 1973b. Trip report: 25 January to 3 February1968. Assoc. Mexican Cave Stud. News., 4:77·78.Reddell, J. R. 1973c. Trip report: 10-22 June 1972. Assoc.Mexican Cave Stud. News., 4:52-54.Reddell, J. R. 1973d. Trip report: 21-29 November 1972.Assoc. Mexican Cave Stud. News., 4:80,87.Reddell, J. R. 1973e. Trip report: 22 December 1972-8 J anuary1973. Assoc. Mexican Cave Stud. News., 4:87-91.Reddell,.T. R. 1973f. Trip report: 16 February-14 May 1973.Assoc. Mexican Cave Stud. News., 4:92-95.Reddell, J. R. 1973g. Trip report: 19-27 July 1973. Assoc.Mexican Cave Stud. News., 4:96.Reddell, J. R. 1974. A preliminary report on the ZacapoaxtlaCuetzalan Area, northern Puebla, Mexico. Supplement.Assoc. Mexican Cave Stud. News., 4:185-190, pIs. 5-6.Reddell, J. R. 1977a. The caves of Chihuahua and Durango.Assoc. Mexican Cave Stud. News., 5:84-93, pIs. 1-4.Reddell, J. R. 1977b. A preliminary survey of the caves ofthe Yucatan Peninsula. Assoc. Mexican Cave Stud. Bull.,6:215-296.Reddell, J. R., ed. 1977c. Studies on the caves and cave faunaof the Yucatan Peninsula. Assoc. Mexican Cave Stud.Bull., 6.296 pp.Reddell, J. R. 1979. Trip report: 6-18 July 1967. Assoc.Mexican Cave Stud. News., 5 :99-101.Reddell, J. R. and W. R. Elliott. 1973a. A checklist ofthe cave fauna of Mexico. IV. Additional recordsfrom the Sierra de EI Abra, Tamaulipas and San LuisPotosi. Assoc. Mexican Cave Stud. Bull,. 5:171180.Reddell, J. R., and W. R. Elliott. 1973b. A checklist of thecave fauna of Mexico. V. Additional records from theSierra de Guatemala, Tamaulipas. Assoc. Mexican Ca~eStud. Bull., 5:181-190.279

Reddell, J. R., and W. R. Elliott. 1974. Trip report: 21 De·cember 1973-12 January 1974. Assoc. Mexican CaveStud. News., 5:7-13, pI. 1.Reddell, J. R., and R. W. Mitchell. 1969. A checklist andannotated bibliography of the subterranean aquatic faunaof Texas. Internatl. Center Arid Semiarid Land Stud.,Spec. Rept., 24. 48 pp.Reddell, J. R., and R. W. Mitchell. 1971a. A checklist of thecave fauna of Mexico. I. Sierra de EI Abra, Tamaulipasand San Luis PotosI. Assoc. Mexican Cave Stud. Bull.,4:137-180.Reddell, J. R., and R. W. Mitchell. 1971b. A checklist of thecavefauna of Mexico. II. Sierra de Guatemala, Tamaulipas.Assoc. Mexican Cave Stud. Bull., 4:181-215.Reddell, J. R., and R. W. Mitchell. 1971c. Preface. Assoc.Mexican Cave Stud. Bull., 4:1-2.Reddell, J. R., and R. W. Mitchell, eds. 1971d. Studies on thecavernicole fauna of Mexico. Assoc. Mexican Cave Stud.Bull., 4. 239 pp.Reed, M. 1966. Raising and breeding the blind cave characin.Trop. Fish Hobbyist, 14:79-85.Remy, P. 1948. Palpigrades du Mexique et de Cuba. Ciencia,Mexico, 9:33-36.Reynolds, J. W., and D. G. Cook. 1976. omenclatura oligochaetologica.A catalogue of names, descriptions and typespecimens of the Oligochaeta. Fredericton: Univ. NewBrunswick. x + 217 pp.Richardson, H. 1905. A monograph on the isopods of NorthAmerica. Bull. United States Natl. Mus., 54.727 pp.Rioja, E. 1950. Estudios carcinologicos. XXII. Los triconiscidoscavernicolas de Mexico del genero Protrichoniscusy descripcion de una nueva especie del mismo. Anal. lnst.BioI., Mexico, 21: 127-146.Rioja, E. 1951a. Estudios carcinologicos. XXV. EI hallazgodel genero Sphaeromicola en America (ostracodos, citeridos)y descripcion de una nueva especie. Anal. Inst.BioI., Mexico, 22: 169-179.Rioja, E. 1951b. Esudios carcinologicos. XXVI. Descripcionde Protrichoniscus acostai n. sp. (crust. ioopodo) deComitan, Chiapas. Anal. Inst. BioI., Mexico, 22:181-189.Rioja, E. 1952. Estudios carcinologicos. XXVII. Descripcionde una nueva esp~cie del genero Cubaris (isopodo,cubarido) de la Cueva de Los Sabinos (San Luis Potosi).Anal. Inst. BioI., Mexico, 22:517-524.Rioja, E. 1953a. Los crustaceos cavernicolas de Mexico.Mem. Congr. Cient. Mexicano, 7:285-298.Rioja, E. 1953b. Estudios carcinologicos. XXVIII. Descripcionde un nuevo genero de potamonidos cavernicolas yciegos de la Cueva del Tio Ticho, Comitan, Chis. Anal.Inst. BioI., Mexico, 23:217-225.Rioja, E. 1953c. Estudios carcinologicos. XXIX. Un nuevogenero de isopodo triconfscido de la Cueva de Ojo deAgua Grande, Paraje Nuevo, Cordoba, Ver. Anal. Inst.BioI., Mexico, 23:227-241.Rioja, E. 1953d. Estudios carcinologicos. XXX. Observacionessobre los cirolanidos cavernicolas de Mexico (crustaceos,isopodos). Anal. Inst. BioI., Mexico, 24:147-170.Rioja, E. 1953e. Los extrafios pobladores del mundo subterraneo.Investigaciones mexicanas. Rev. Univ. Mexico,8(2):9-11.Rioja, E. 1954. Estudios carcinologicos. XXXI. Algunas especiesde armadilidos de las cuevas de Mexico (ioopodosterrestres). Anal. Inst. BioI., Mexico, 25:275-288.Rioja, E. 1955a. Estudios carcinologicos. XXXIII. Observa-ciones acerca de dos nuevas especies de isOpodos cavernicolasde Chiapas. Anal. Inst. BioI., Mexico, 26:199-209.Rioja, E. 1955b. Trichoniscidae cavernicolas de Mexico. Rev.Soc. Mexicana Entomol., 1:39-62.Rioja, E. 1956. Estudios carcinologicos. XXXIV. Dos nuevosisopodos cavernicolas de la Sierra Madre Oriental (regionde Xilitla), Mexico. Anal. Inst. BioI., Mexico, 26:447-457.Rioja, E. 1957. Estudios carcinologicos. XXXV. Datos sobrealgunos isopodos cavernicolas de la isla de Cuba. Anal.Inst. BioI., Mexico, 27:437-462.Rioja, E. 1958. Estudios carcinologicos XXXVI. Descripciony estudio de una especie nueva del genero Cylindroniscus(isopodo triconiscido) de Yucatan. Anal. Inst. BioI.,Mexico, 28:267-278.Rioja, E. 1962. Caracteres biogeograficos de Mexico y deCentro America. Rev. Soc. Mexicana Hist. Nat., 23:27-50.Rioja, E. 1971. Clase IV. Los crustaceos (Crustacea). Pp. 470554 in L. Cendrero, eds., Zoologia hispanoamericana.Invertebrados. Biblioteca de Ciencias Biologicas y Geologicas.Mexico, D. F.: Editorial Porrua.Rioja Lo Bianco, E., M. Ruiz Oronoz, and I. Larios Rodriguez.1961. Tratado elemental de zoologia, 5th ed.Mexico, D. F.: Editorial Porma. xvi + 739 pp.Robaux, P., J. P. Webb, Jr., and G. D. Campbell. 1977. Uneforme nouvelle de Thrombidiidae (Acari) parasite surplusieurs especes d'orthopteres cavernicoles du genreCeuthophilus (Orthoptera, Rhaphidophoridae). Ann.Speleol.,31:213-218.Robles Ramos, R. 1950. Los recursos naturales de Yucatan.I.-Apuntes sobre la morfologia de Yucatan. Bol. Soc.Mexicano Geogr. Estad., 69:27-106.Rodriguez, G., and A. E. Smalley. 1972. Los cangrejos deagua dulce de Mexico de la familia Pseudothelphusidae(Crustacea, Brachyura). Anal. Inst. BioI. Univ. Nal.Auton. Mexico, 40, Ser. Cienc. Mar Limnol., (1 ):69-112.Rodriguez Cabo, J. 1953. Algunos yacimientos de fertilizantesde Mexico. Mem. Congr. Cient. Mexicano, 3:317368.Rodriguez de la Cruz R., M. C. 1965. I: Contribucion al conocimicntode los palemonidos de Mexico. II. Palemoni·dos de Atlantico y Vertiente Oriental de Mexico condescripcion de dos especies nuevas. Anal. Inst. Nac. Invest.Biol.-Pesqueras, 1 :73-112,8 pis.Rosen, D. E. 1976. A vicariance model of Caribbean biogeography.Syst. Zool., 24:431-464.Rosen, D. E., amI R. M. Bailey. 1963. The poeciliid fishes(Cyprinodontiformes), their structure, zoogeography, andsystematics. Bull. American Mus. Nat. Hist., 126:1-176,pis. 1-2.Rosen, D. E., and P. H. Greenwood. 1976. A fourth neotropicalspecies of synbranchid eel and the phylogeny andsystematics of synbranchiform fishes. Bull. AmericanMus. Nat. Hist., 157:1-69.Rosen, D. E., and A. Rumney. 1972. Evidence of a secondspecies of SynbranchlJs (Pisces, Teleostei) in SouthAmerica. American Mus. Nov., 2497. 45 pp.Rossi, W., and M. G. Cesari Rossi. 1977. SuUe Laboulbeniali(Ascomycetes) parasite dei Trechinae del Messico (Coleoptera,Carabidae). Quad. Accad. Naz. Lincei, Probl.Att. Sci. Cult., 171(3):373-376, pI. 1.Roth, V. D. 1968. The spider genus Tegenaria in the WesternHemisphere (Agelenidae). American Mus. Nov., 2323.33 pp.Roth, V. D., and P. 1. Brame. 1972. Nearctic genera of the280

spider family Agelenidae (Arachnida, Araneida). AmericanMus. Nov., 2505. 52 pp.Rouch, R. 1971. Recherches sur les eaux souterraines. 14.Peuplement par les harpacticides d'un drain situe dans lazone de circulation permanente. Ann. Speleol., 26:107133.Rowland, J. M. 1971a. Agastochizomus lucifer, a new genusand species of cavernicole schizomid (Arachnida, Schizomida)from Mexico. Assoc. Mexican Cave Stud. Bull.,4:13-17.Rowland, J. M. 1971b. New species of schizomids (Arachnida,Schizomida) from Mexican caves. Assoc. Mexican CaveStud. Bull., 4: 117-126.Rowland, J. M. 1972. Brooding habits and early developmentof Trithyreus penmpeltis (Arachnida: Schizomida).Entomol. News, 83:69·74.Rowland, J. M. 1973a. A new genus and several new speciesof Mexican schizomids (Schizomida: Arachnida). Occ.Papers Mus. Texas Tech Univ., 11. 23 pp.Rowland, J. M. 1973b. Revision of the Schizomida (Arachnida).J. New York Entomol. Soc., 80:195-204.Rowland, J. M. 1973c. Three new Schizomida of the genusSchizomus from Mexican caves (Arachnida). Assoc. MexicanCave Stud. Bull., 5:135·140.Rowland, J. M. 1973d. Two new troglobitic Amblypygida ofthe genus Tarantula from Mexican caves (Arachnida).Assoc. Mexican Cave Stud. Bull., 5: 123-128.Rowland, J. M. 1975a. Classification, phylogeny and zoogeographyof the American arachnids of the order Schizo.i mida. Ph.D. Diss. Lubbock: Texas Tech Univ. ix + 415 pp.Rowland, J. M. 1975b. A partial revision of Schizomida(Arachnida), with descriptions of new species, genus, andfamily. Occ. Papers Mus. Texas Tech Univ., 31. 21 pp.Rowland, J. M., and J. R. Reddell. 1977. A review of the cavemicoleSchizomida (Arachnida) of Mexico, Guatemala,and Belize. Assoc. Mexican Cave Stud. Bull., 6:79·102.Rowland, J. M., and J. R. Reddell. 1979a. The order Schizomida(Arachnida) in the New World. I. Protoschizomidaeand dumitTescoae group (Schizomidae: Schizomus).J. Arachnol., 6: 161-196.Rowland, J. M., and J. R. Reddell. 1979b. The order Schizomida(Arachnida) in the New World. II. Simonis andbrasiliensis groups (Schizomidae: Schizomus). J. Arachnol.,7:89-119.Roy, S. 1974. Las Grutas de Juxtlahuaca. Assoc. MexicanCave Stud. News., 5:39·42, pI.Rudnick, A. 1960. A revision of the mites of the family Spinturnicidae(Acarina). Univ. California Publ. Entomol.,17:157·283.Ruffo, S., and A. Vigna Taglianti. 1974. Three new subterraneanBogidiella from Mexico and Guatemala (Crustacea,Amphipoda). Quad. Accad. Naz. Lincei, Probl. Alt. Sci.Cult., 171(2):105-133.Ruffo, S., and A. Vigna Taglianti. 1977. Secondo contributoalla conoscenza del genere Bogidiella in Messico e Guatemala(Crustacea, Amphipoda,Gammaridae). Quad. Accad.Naz. Lincei, ProbL Alt. Sci. Cult., 171(3):125-172.Russell, W. 1973a. On the precipice. Texas Caver, 18:363366.Russell, W. 1973b. Trips: May 19-20, 1973. Texas Caver,18:280.Russell, W. H. 1965. Trip report: July 1,1965. Assoc. MexicanCave Stud. News., 1:61-63.Russell, W. H. 1969. Physiographic divisions of Mexico.Assoc. Mexican Cave Stud. News., 3:27-31.Russell, W. H. 1972a. Alphabetical listing of caves of theSierra de El Abra, EI Abra list number one, July 1,1972. Assoc. Mexican Cave Stud. News., 3:129-132.Russell, W. H. 1972b. Corrections and additions to themap of "La Region de la Sierra de El Abra." Assoc.Mexican Cave Stud. News., 3:126-127.Russell, W. H. 1972c. Geographical listing of the caves ofthe Sierra de EI Abra. Assoc. Mexican Cave Stud. News.,3:133-142.Russell, W. H. 1972d. Trip report: 15 August 1971. Assoc.Mexican Cave Stud. News., 3:70-71.Russell, W. H. 1973a. Trip report: Christmas-New Year'svacation 1968·69. Assoc. Mexican Cave Stud. News.,4:47-49.Russell, W. H. 1973b. Trip report: Easter vacation 1969.Assoc. Mexican Cave Stud. News., 4:49--50.Russell, W. H. 1974. Trip report: Christmas 1973. Assoc.Mexican Cave Stud. News., 4:120·121.Russell, W. H. 1977. School bus scoops Zoquithin area.Assoc. Mexican Cave Stud. Activities News., 5:20.Russell, W. H., and T. W. Raines, eds. 1967. Caves of theInter-American Highway, uevo Laredo, Tamaulipas, toTamazunchale, San Luis PotosI. Assoc. Mexican CaveStud. Bull., 1. 126 pp.Rzedowski, J. 1978. Vegetacion de Mexico. Mexico, D. F.:Editorial Lirnusa. 432 pp.Sabrosky, C. W. 1959. A revision of the genus Pholeomyiain North America (Diptera, Milichiidae). Ann. Entomol.Soc. America, 52:316-331.Sadoglu, P. 1956. A preliminary report on the genetics of theMexican cave characins. Copeia, 1956:113-114.Sadoglu, P. 1957. A Mendelian gene for albinism in naturalcave fish. Experientia, 13:394.Sadoglu, P. 1958. Mendelian inheritance in the hybrids betweenthe Mexican blind cave fishes and their overgroundancestor. Verh. Deuthschen ZooL Ges. Graz, 1957:432439.Sadoglu, P. 1967. The selective value of eye and pigmentloss in Mexican cave fish. Evolution, 21:541-549.Sadoglu, P. 1975. Genetic paths leading to blindness inAstyanax mexicanus. Pp. 419-426 in M. A. Ali, eds.,Vision in fishes. New York: Plenum Press.Sadoglu, P., and A. McKee. 1969. A second gene that affectseye and body color in Mexican blind cave fish. J. Hered.,60:10-14.Sailer, R. I. 1950. The second record for Pimicimex cavernisBarber (Heteroptera, Cimicidae). Proc. Entomol. Soc.Washington, 52:308.Salas, G. P. 1962. Estudio general fotogeologico de la partenor-oeste del departamento del Peten, Republica deGuatemala. Bol. Asoc. Mexicana PetroL, 14:183-202, pIs.5·9,1 map.Salas, G. P., and E. Lopez Ramos. 1951. Geologia y tectonicade la region de Macuspana, Tabasco y parte norte de Chiapas.BoL Asoc. Mexicana GeoL PetroL, 3:3-56, 5 pis.,2 tables.Salinas, S. 1960. Caracteristicas estratigraJicas de la region LaPerla-Orizaba-Tequila, Edo. de Veracruz. Bol. Asoc. MexicanaGeol. Petrol., 12:145·199, 3 maps. .Salmon, J. T. 1964. An index to the Collembola. Bull. Roy.Soc. New Zealand, 7. 644 pp.281

Sanborn, C. C. 1936. Records and measurements of neotropicalbats. Field Mus. Nat. Hist., Zoo!' Ser., 20:93-106.Sanderson, I. T. 1941. Living treasure. New York: VikingPress. 290 pp.Santiago Acevedo, J. 1962. Estructuras de la porcion occidentaldel frente dc la Sierra Madre de Chiapas. Bol. Asoc.Mexicana Geo!. Petro!., 14:111-134, figs. 1-12.Saussure, H. de. 1897. Fam. Gryllidae. Pp. 198-284, pis. 1113 in Biologia Centrali-Americana, Zoo!., Orthoptera,Vo!. I, 1893-1899. London [Gryllidae 1894-18971.Sbordoni, V. 1974. A new cave dwelling Ptomaphagus(Co!. Catopidae) from Tabasco, Mexico. Quad. Accad.Naz. Lincei, Prob!. Att. Sci. Cult., 171(2):363-367.Sbordoni, V., and R. Argano. 1972. Introduction; cavesstudied during the 1st mission to Mexico (1969). Quad.Accad. Naz. Lincei, Probl. Att. Sci. Cult., 171(1):5-21,pis. 1-8.Sbordoni, V., R. Argano, and A. ZulIini. 1974. Biologicalinvestigations on the caves of Chiapas (Mexico) and adjacentcountries: Introduction. Quad. Accad. Naz. Lincei,Prob!. Att. Sci. Cult., 171(2):5-45, pis. 1-6, 3 maps.Sbordoni, V., R. Argano, V. Vomero, and A. Zullini. 1977.Richerche sulla fauna cavernicola del Chiapas (Messico)e delle regioni limitrofe: GroUe esplorate nel 1973 e nel1975. Criteri per una classificazione biospeleologica dellegroUe. Quad. Accad. Naz. Lincei, Prob!. Att. Sci. Cult.,171(3):5-74, pis. 1-9, map.Sbordoni, V., and M. Cobolli-Sbordoni. 1973a. Aspetti ecologicied evolutivi del popolamento di grotte temperate etropicali: Osservazioni sui ciclo biologico di aIcune speciedi Ptomaphagus (Coleoptera Catopidae). Internat!. J.Speleo!., 5 :337·347.Sbordoni, V., and M. Cobolli-Sbordoni. 1973b. Evolutionaryrates in tropical and temperate cave communities: Lifecycle and cave adaptation in Mexican species of Ptoma·phagus (Coleoptera Catopidae). (Abstr.). P. 150 in V.Panos, ed., International Speleology 1973. Abstracts ofPapers. 6th Internatl. Congr. Speleol., Olomouc, Czechoslovakia.Schaldach, W. J., Jr. 1965. Notas breves sobre algunos mamiferosdel sur de Mexico. Anal. Inst. Bio!., Mexico,35:129-137.Schaldach, W. J., Jr. 1966. New forms of mammals fromsouthern Oaxaca, Mexico, with notes on some mammalsof the coastal range. Siiugetierk. Mitt., 13-14(4):286297.Schemmel, C. 1967. Vergleichende Untersuchungen an denHautsinnesorganen ober- und unterirdisch lebender Astyanax-Formen.Ein Beitrag zur Evolution der Cavernicolen.Z. Morph. Tiere, 61:255-316.Schemmel, C. 1973. Les organes sensorieIs cutanes du genreAstyanax (Pisces, Characidae) chez les formes occupantdes biotopes souterrains. Ann. SpeIeo!., 28:209-219.Schemmel, C. 1974a. Genetische Untersuchungen zur Evolutiondes Geschmacksapparates bei cavernicolen Fischen.Z. Zool. Syst. Evolutionsf., 12:196-215.Schemmel, C. 1974b. 1st die cavernicole Micos-Populationvon Astyanax mexicanus (Characidae, Pisces) hybridenUrsprungs? Mitt. Hamburg. Zoo!' Mus. lnst., 71: 193-201.Schlaepfer, C. .J. 1968. Hoya Mexico 14Q.h(5) con Resumende la geologia de la Hoya Mexico, Distrito Federal y estadosde Mexico y Morelos. Univ. Nac. Aut. Mexico, Inst.Geo!.. Carta Geologica de Mexico, Serie de 1:100,000,no. 12.Schlagel, S. R., and C. M. Breder, Jr. 1947. A study ofthe oxygen consumption of blind and eyed cave characinsin light and in darkness. Zoologica, New York, 32;17-27.Schmatolla, E. 1972. Dependence of tectal neuron differentiationon optic innervation in teleost fish. J. Embryo!.Exp. Morphol., 27:555-576.Schmatolla, E., and G. Erdmann. 1973. Influence of retinotectalinnervation on cell proliferation and cell migrationin the embryonic teleost tectum. J. Embryo\. Exp.Morpho!.,29:697-712.Scholl, A. 1973. Biochemical evolution in the genus Xiphophorus(Poeciliidae: Teleostei). Pp. 277-299 in J. H.Schroder, cd., Genetics and mutagenesis of fish. NewYork: Springer-Verlag.Schroder,.J. H. 1973. Preface. Pp. v-vii in J. H. Schroder, ed.,Genetics and mutagenesis of fish. New York: SpringerVerlag.Schuchert, C. 1935. Historical geology of the AntilleanCaribbean region. New York: Hafner Pub!. Co. 811 pp.Schultz, G. A. 1964. Mexiconiseus tlamayaensis, a new genusand species of terrestrial cave isopod from San Luis Potosi,Mexico. Trans. American Microscop. Soc., 83:376-380.Schultz, G. A. 1965. Terrestrial isopods from caves and minesin Texas and northern Mexico with a description of Vene·zillo tanneri (Mulaik and Mulaik) allotype. Texas J. Sci.,17:101-109.Schultz, G. A. 1968. Xilitloniscus Bowman a synonym ofMexiconiscus Schultz with notes on the species involved(Isopoda, Oniscoidea). Crustaceana, 14:255-258.Schultz, G. A. 1970a. Cylindroniscus vallesensis sp. nov.:Description with review of genus (lsopoda, Trichoniscidae).Trans. American Microscop. Soc., 89:407-412.Schultz, G. A. 1970b. Disposition of species of terrestrialisopod crustaceans of the genera Synuropus, Spherarmadillo,Spheroniscus and Scleropactes (Oniscoidea, Sphaeroniscidae).Proc. Bio!. Soc. Washington, 83:123-131.Schultz, G. A. 1974. Mexicerberus troglodytes n. gen., n. sp.from a cave in Mexico, with notes on isopod crustaceansof the Microcerberidea from the New World. Crustaceana,26: 308-312.Schultz, G. A. 1977. Two blind species, one new, of terrestrialisopod crustaceans (Oniscoidea: Philosciidae) fromYucatan and Guatemala. Assoc. Mexican Cave Stud.Bull., 6:9-13.Segerstrom, K. 1961. Geology of the Bemal-Jalpan area,estado de Queretaro, Mexico. United States Geo!. Surv.Bull., 1104:19-86, pI. 2.Segerstrom, K. 1962. Geology of south-central Hidalgo andnortheastern Mexico, Mexico. United States Geoi. Surv.Bull., 1104:87-162, pI. 3Shawcross, M. 1971. Some notes on caves in Guatemala andBritish Honduras. Canadian Caver, no. 4:63-71.Shawcross, M. 1978. Some "Hallucination"! Canadian Caver,10(1):3-4.Shawcross, M., B. Pratt, and G. Tracey. 1974. Mexico andGuatemala. Canadian Caver, 6(1):60-72.Shear, W. A. 1969. A synopsis of the eave millipeds of theUnited States, with an illustrated key to genera. Psyche,76: 126-143.Shear, W. A. 1972. Studies in the milliped order Chordeumida(Diplopoda): A revision of the family Cleidogonidaeand a reclassification of the order Chordeumida in theNew World. Bull. Mus. Compo Zoo!., 144:151-352.282

Shear, W. A. 1973. Jarmilka alba, n. gen., n. sp. (Diplopoda:Spirostreptida: Cambalidae), a new milliped from a cavein Belize. Assoc. Mexican Cave Stud. Bull., 5:43-45.Shear, W. A. 1974. Millipeds (Diplopoda) from Mexican andGuatemalan caves. Quad. Accad. Naz. Lincei, Probl. Att.Sci. Cult., 171(2):239·305.Shear, W. A. 1976. The spider family Ochyroceratidae new tothe United States. Bull. British Arachnol. Soc., 3:249-250.Shear, W. A. 1977a. Millipeds (Diplopoda) from caves inMexico, Belize and Guatemala. IIi. Quad. Accad. Naz.Lincei, Probl. Au. Sci. Cult., 171(3):235.265.Shear, W. A. 1977b. The opilionid genus Neogovea Hinton,with a description of the first troglobitic cyphophthalmidfrom the Western Hemisphere (Opiliones, Cyphophthal.mi). J. Arachnol., 3:165·] 75.Shear, W. A. 1978. Taxonomic notes on the armored spidersof the families Tetrablemmidae and Pacullidae. AmericanMus. Nov., 2650. 46 pp.Silhavy, V. 1974. Cavernicolous opilionids from Mexico(Arachnida, Opiliones). Quad. Accad. Naz. Lincei, Prohl.AU. Sci. Cult., 171(2):175·194.Silhavy, V. 1977. Further cavernicolous opilionids fromMexico. Quad. Accad. :"Jaz. Lincei, Probl. Au. Sci. Cult.,171(3):219·233.Silhavy, V. 1979. Opilionids of the suborder Gonylepto.morphi from the American caves, collected by Dr. PierreStrinati. Rev. Suisse Zool., 86:321·334.Silva Taboada, G. 1974. Sinopsis de la espeleofauna cubana.Acad. Cienc. Cuba, Ser. Espeleol. Carsol., 43. 65 pp.Silvestri, F. 1912. Tisanuri finora noti del Messico. Bol.Lab. Zool. Gen. Agr. Portici, 6:204·22l.Silvestri, F. 1948. Specie di Japygidae (Insecta Diplura)linora raccolti nel Messico. Bol. Lab. Entomol. Agr. Lincei,8:297·320.Sligar, C. 1974. An investigation of tectal efferents in theblind cave fish, A. Hubbsi. (Abstr.). Anat. Rec., 178:467.Sligar, C. M., and T. J. Voneida. 1976. Tectal efferents in theblind cave fish Astyanax hubbsi. J. Compo eurol.,165:107·124.Smalley, A. E. 1970. A new genus of freshwater crabs fromGuatemala, with a key to the Middle American genera(Crustacea Decapoda, Pseudothelphusidae). AmericanMidI. Nat., 83:96·106.Smith, A. R. 1968. Mayan cave discoveries. Geology. Explor·ers J., 46:160·163.Smith, C. I. 1970. Lower Cretaceous stratigraphy, northernCoahuila, Mexico. Univ. Texas Bur. Econ. Geol. Rept.Inv., 65. 101 pp., 15 pis.Smith, H. G.1956. A comparative study of the neurocraniumand weherian apparatus of the ictalurid (ameiurid) cat·fishes. M. S. Thesis. Tulane Univ. 77 pp.Smith, H. M., and M. Alvarez del Toro. 1977. A new troglo.dytic lizard (Reptilia, Lacertilia, Xantusiidae) fromMexico. J. Herpetol., 11:37-40.Smith, H. M., and R. G. Van Gelder. 1955. New and note·worthy amphibians and reptiles from Sinaloa and Puebla,Mexico. Herpetologica, 11:145·149.Smith, .T. D. 1972. Systematics of the chiropteran familyMormoopidae. Univ. Kansas Mus. Nat. Hist. Misc. Publ.,56.132 pp.Smith, R. E. 1954. Cenote exploration at Mayapan and Tel·chaquillo. Carnegie Inst. Washington Dept. Archaeol.,Current Repts., 12:222·233.Smith, R. M. 1972. Geology of the Ahuacatlan Area. Pp. 215 in T. W. Raines, Sotanito de Ahuacatlan. Sierra MadreOriental; Jalpan; Ahuacatlan. Assoc. Mexican Cave Stud.Cave Rept. Ser., l.Soleglad, M. E. 1975. A redescription of Vejovis gracilisGertsch & Soleglad based on the adult (Scorpionida:Vejovidae). WasmannJ. BioI., 33:107·120.Soleglad, M. E. 1976. A revision of the scorpion subfamilyMegacorminae (Scorpionida: Chactidae). Wasmann J.Biol.,34:251.303.Solonano, A. 1953. Variacion en Typhliasinapearsei (Hubbs).Ciencia, Mexico, 12:286.Spieth, H. T. 1950. The David Rockefeller Mexican Expeditionof the American Museum of Natural History. Intro·ductory account. American Mus. Nov., 1454. 67 pp.Spilman, T. J. 1968. Two new species of Niptusfrom NorthAmerican caves (Coleoptera: Ptinidae). SouthwesternNat., 13:193-200.Sprouse, P. 1974. Trip report: 29 December 1973·5 January1974. Assoc. Mexican Cave Stud. News., 4:122·124.Sprouse, P. 1977. Sotano de Sauz. Assoc. Mexican Cave Stud.News., 5:82-83, pI. 5.Sprouse, P. 1979. Discovery in Cuetzalan. Assoc. MexicanCave Stud. Activities News., 10:60-63,2 maps.Stach, J. 1956. The apterygotan fauna of Poland in relationto the world·fauna of this group of insects. Family: Smin·thuridae. Krakow: Polska Akademia Nauk. ]nstytut Zoologiczny.287 pp., 33 pis.Stach, J. 1959. On the three species of the genus Acheronti·des Bon. (Collembola) from the Afghanistan caves and therelationshiD of this genus with some other similar !Ienera.Acta Zool. Cracoviensia, 4:425·445.Steele, B. 1977a. Deeper in Yochib. Canadian Caver, 8(2):3·9.Steele, B. 1977b. Sumidero Yochib. Assoc. Mexican CaveStud. Activities News., 6:11·14.Steele, B. 1978. Expedition report. Christmas 1977 Huautlaproject. Assoc. Mexican Cave Stud. Activities News.,8:24·35,3 maps.Stefanelli, A. 1954a. The differentiation of optic lobes neu·rons in a blind cave teleost. Experientia, 10:436·438.Stefanelli, A. 1954b. II tetto oUico di pesci ciechi caverni·coli nei riguardi del differenziamento specifieo dei neu·roni. Rend. Accad. Naz. Lincei, 16:277·282.Stephens, J. L. 1843. Incidents of travel in Yucatan. Harperand Bros.: New York. 2 vols.Sterba, G. 1963. Freshwater fishes of the world. Tr. and rev.by D. W. Tucker. New York: Viking Press. 878 pp.Steven, D. M. 1963. The dermal light sense. BioI. Rev., 38:204-240.Steyskal, G. C. 1973. A new species of the genus Archibor·borus Duda from Mexico (Diptera: Sphaeroceridae). J.Kansas Entomol. Soc., 46:154-157.Stock, J. H. 1977. The taxonomy and zoogeography of thehadziid Amphipoda, with emphasis on the West Indiantaxa. Stud. Fauna Cura~ao and other Caribbean Islands,177:1·130.Stolk, A. 1958. Tumours of fishes. XXIV. Ocular melanomain the characid Anoptichthys jordani Hubbs et Innes.Proc. K. Nederlandse Akad. Wet. Amsterdam, ser. C, 61:382·394.Stone, B. 1977. Ten days in La Grieta. AMCS Huautla Expe.dition, May 1977. Assoc. Mexican Cave Stud. ActivitiesNews., 7:7.17, map.283

Stone, B. 1979. The 1979 San Agustin Expedition. Assoc.Mexican Cave Stud. Activities News., 10:33-58,3 maps.Stone, B., and R. Jameson. 1977. Caves of the San JuanPlateau. Assoc. Mexican Cave Stud. Bull., 7. 59 pp.,7 maps.Straskraba, M. 1969. Lista de 105 crustaceos dulceacuicolasde Cuba y sus relaciones zoogeograficas. Acad. Cienc.Cuba, Ser. BioI., 8. 37 pp.Strenth, N. E. 1976. A review of the systematics and zoogeographyof the freshwater species of PaIaemonetesHeller of North America (Crustacea: Decapoda). SmithsonianContr. Zool., 228. 27 pp.Strinati, P. 1968. Expeditions biospeologiques en Ameriquelatine. Stalactite, 1: [4 pp.].Strinati, P. 1970. Les vertebres cavernicoles. Mus. Geneve,108.4 pp.Strinati, P. 1977. Recherches sur les invertebres cavernicolesde Guatemala. Proc. 7th Internat\. Speleo\. Congr., Sheffield,England, p. 387-389.Stromsvik, G. 1956. Exploration of the cave of Dzab-Nah,Tecoh, Yucatan. Carnegie Inst. Washington Dept. Archaeo\.Current Repts., 35:463-470.Strotkoetter, E, 1960. Rezeption hydrostatischer Drucke beiFischen und ihre Mitwirkung an der Orientierung imRaum. Naturwissenschaften, 47:611-612.Sumhera, J. 1972. Trip report: June 1-6, 1967. Pp. 118-119in M. Walsh, ed., Mexican caving of the Southwest TexasGrotto: 1966-1971. San Marcos: Southwest Texas Grotto.Sumichrast, F. 1882. Enumeracion de las especies de mamiferos,aves, reptiles y batracios observados en la parte centraly meridional de la Republica Mexicana. Naturaleza,5:199-213,322-328.Sustare, D. 1966. Trip report: 30 May-4 June 1966. Assoc.Mexican Cave Stud. News., 2:54-56.Suttkus, R. D. 1961. Additional information about blind catfishesfrom Texas. Southwestern Nat., 6:55-64.Syme, R., and S. B. Peck. 1974. Notes on caves in Chiapas,Mexico, and Guatemala. Assoc. Mexican Cave Stud.News., 4:144-146.Tamayo, J. L. 1962. Geografia general de Mexico. Geografiabiologica y humana. Torno Ill. Mexico, D. F.: InstitutoMexicano de Investigaciones Economicas. 633 pp.Tamayo, J. L., and R. C. West. 1964. The hydrography ofMiddle America. Pp. 84-121 in R. C. West, ed., Handbookof Middle American Indians, Volume 1: Natural environmentand early cultures. Austin: Univ. Texas Press.Tattersall, W. M. 1951. A review of the Mysidacea of theUnited States National Museum. Bull. United States Nat\.Mus., 201. x + 292 pp.Tavolga, W. N. 1971. Sound production and detection. Pp.135-205 in W. S. Hoar and D. J. Randall, eds., Fishphysiology, Volume V: Sensory systems and electric organs.New York: Academic Press. Reprinted in part in:W. N. Tavolga, ed., 1976. Sound reception in fishes, pp.3-36. Stroudsburg, Pennsylvania: Dowden, Hutchinsonand Ross, Inc.Taylor, D. W. 1966. A remarkable snail fauna from Coahuila,Mexico. Veliger, 9:152-228.Taylor, D. W" and W. L. Minckley. 1966. New world forbiologists. Pacific Discovery, 19(5):18-22.Taylor, W. R. 1969. A revision of the catfish genus NoturusRafinesque with an analysis of higher groups in the Ictaluridae.Bull. United States Nat\. Mus., 282. 315 pp.Taylor, W. W. 1956. Some implications of the carbon-14dates from a cave in Coahuila, Mexico. Bull. Texas Archeo\.Soc., 27:215-234.Tellez Giron, A. 1944. EI vampiro portador de virus delderriengue. Rev. Soc. Mexicana Hist. Nat., 5:35-42.Thibaud, J .-M. 1963. Documents sur Ie genre AcherontielIaAbsolon (Collembole) et description d'une espece de Coted'Ivoire. Ann. Speleo\., 18:287-297.Thines, G. 1954. Etude comparative de la photosensibilitedes poissons aveugles Caecobarbus geertsii Blgr. et Anoptichthysjordani Hubbs & Innes. Ann. Soc. Roy. Zool.Belgique, 85:35-58.Thines, G. 1955. Les poissons aveugles (1) Origine-taxonomie-repartitiongeographique-comportement. Ann. Soc.Roy. Zoo\. Belgique, 86:1-128_Thines, G. 1956. A note on the light sensitivity of blind cavefishes. (Abstr.). Proc. Assoc. Study Anim. Behav., 4:79.Thines, G. 1960. Sensory degeneration and survival in cavefishes. Symp. Zoo\. Soc. London, 3:39-51, pIs. 1-2.Thines, G. 1969. L'Evolution regressive des poissons cavernicoleset abyssaux. Paris: Masson et Cie. 394 pp.Thines, G., and D. Capon. 1975. L'aveuglement experimentalde poissons epiges et la cecite genetique des poissonscavernicoles: Effets sur Ie comportement alimentaire.Spelunca, Mem., 8:209-228.Thines, G., and J .-P. Durand. 1973. Connaissances actuellessur l'appareil sensoriel de la ligne laterale chez les vertebrescavernicoles aquatiques. Ann. Speleo\., 28:271-282.Thines, G., and J. Kiihling. 1957. Untersuchungen iiber dieFarbempfindlichkeit des Hohlenfisches Anoptichthys jordaniHubbs und Innes (Characidae). Z. Biol., 109: 150-160.Thines, G., and J.-M. Legrain. 1973. Effets de la substanced'alarrne sur Ie comportement des poissons cavernicolesAnoptichthys jordani (Characidae) et Caecobarbus geertsi(Cyprinidae). Ann. Speleo\., 28:291·297.Thines, G., and M. Piquemal. 1978. Observations sur les comportementsde Lucifuga subterranea Poey (Pisces, Ophidiidae),poisson cavernicole de Cuba. Internat\. J. Speleo\.,10:195-203.Thines, G., M. Soffie, and E. Vandenbussche. 1967. Analysedu comportement alimentaire du poisson cavernicoleaveugle Anoptichthys Gen. et d'hybrides F 1 (Astyanax XAnoptichthys) et F2' Internatl. J. Speleo\., 2:437-448,pIs. 73-76.Thines, G., and E. Tercafs. 1972. Atlas de Ie vie souterraine.Les animaux cavernicoles. Paris: Editions N. Boubee.161 pp.Thines, G., and M. Weyers. 1978. Reponses locomotrices dupoisson cavernicole Astyanax jordani (Pisces, Characidae)ades signaux periodiques et aperiodiques de lumiere et detemperature. Internat\. J. Speleo\., 10:35-55.Thines, G., and N. Wissocq. 1972. Etude comparee du comportementalimentaire de deux poissons cavernicoles(Anoptichthys jordani Hubbs et Innes et Caecobarbusgeertsi Blgr.). Internat\. J. Speleo\., 4:139-169.Thines, G., and F. Wolff-Van Ermengem_ 1965. Activitypatterns in an epigean Characidae and its degeneratedcave-relative. Anim. Behav., 23:585.Thines, G., F. Wolff, C. Boucquey, and M. Soffie. 1965.Etude comparative de I'activite du poisson cavernicoleAnoptichthys anrrobius Alvarez et de son ancetre epigeAstyanax mexicanus (Filippi). Ann. Soc. Roy. Zoo\.Belgique, 96:61-115.Thompson, E. H. 1897. Cave of Loltun, Yucatan. Mem.Peabody Mus. American Archaeo\. Ethnol., HarvardUniv., 1(2):1-24, pis. I-&.-284

Thompson, F. G. 1967. The land and freshwater snails ofCampeche. Bull. Florida State Mus., BioI. Sci., 11:221256.Thompson, F. G. 1968. Some Mexican land snails of thefamily Urocoptidae. Bull. Florida State Mus., BioI. Sci.,12:125-183.Thompson, P. 1970. Caving in Guerrero, Mexico. CanadianCaver, no. 2:58-62.Thompson, 1'. 1'J72. Caving in Chiapas. Canadian Caver,4(1):9-21.Tozzer, A. M. 1957. Chichen Itza and its Cenote of Sacrifice;a comparative study of contemporaneous Maya and Toltec.Mem. Peabody Mus. Archaeol. Ethnol., HarvardUniv., 11-12. viii + 316 pp., 709 figs.Treacy, T. 1979. Spring activities in the Purificacion area.Assoc. Mexican Cave Stud. Activities News., 9:8-31,2 maps.Turner, C. 1972. Joya de Salas revisited. Canadian Caver,4(1):3-7.Tuttle, M. D. 1968. Feeding habits of Artibeus jamaicensis.J. Mammal., 49:787.Ueshima, N. 1968. Cytology and bionomics of Primicimexcavemis Barber (Cimicidae: Hemiptera). Pan-PacificEntomol.,44:145-152.Ukrain, D. 1979. Swift studies take researchers far afield.Chihuabuan Desert Discovery, 4:6-7.United States. Bureau of Sport Fisheries and Wildlife. DenverWildlife Research Center. Palo Alto, Mexico, Field Station.1970. Vampire bats: Rabies transmission and livestockproduction in Latin America. Ann. Rept., 1969.24 pp.United States. Bureau of Sport Fisheries and Wildlife. DenverWildlife Research Center. Palo Alto, Mexico, Field Station.1971. Vampire bats: Rabies transmission and livestockproduction in Latin America. Ann. Rept., 1970.32 pp.United States. Office of Endangered Species and InternationalActivities. 1974. United States list of endangeredfauna. Washington, D. C.: United States GovernmentPrinting Office. 22 pp.Usinger, R. L. 1966. Monograph of Cimicidae (HemipteraHeteroptera). The Thomas Say Foundation, Volume VII.College Park, Maryland: Entomol. Soc. America. 585 pp.Vachon, M. 1974. Etude des caracteres utilises pour classerles families et les genres de scorpions (arachnides). 1. Latrichobothriotaxie en arachnologie. Sigles trichobothriauxet types de trichobothriotaxie chez les scorpions. Bull.Mus. Nat!' Hist. Nat., ser. 3, 140:857-958.Vandel, A. 1950. Campagne speologique de C. Bolivar et R.Jeannel dans I'Amerique du Nord (1928). 14. Isopodesterrestres recueillis par C. Bolivar et R. Jeannel (1928) etIe Dr. Henrot (1946). Arch. Zool. Exper. Gen., 87:183210.Vandel, A. 1953. Remarques systematiques, morphologiqueset biogeographiques sur un groupe de TrichoniscidaeNord-A tlantiques. (crustaces; isopodes terrestres). Bull.Mus. Nat!. Hist. Nat., ser. 2, 25:368-375.Vandel, A. 1958. La repartition des cavernicoles et la paleogeographie.Actes Deuxieme Congr. Internat!. SpeIeo!.,2(3):31-43.Vandel, A. 1960. Faune de France. 64. Isopodes terrestres(premiere partie). Paris: Editions Paul Lechevalier. 416 pp.Vandel, A. 1964. Biospeologie. La biologie des animauxcavernicoles. Paris: Gauthier-Villars. xviii + 619 pp.Vandel, A. 1965a. Biospeleology. The biology of cavernicolousanimals. Tr. by B. E. Freeman. New York: PergamonPress. xxiv + 524 pp.Vandel, A. 1965b. Sur l'existence d'Oniscoides tres primitifsmenant une vie aquatique et sur Ie polyphyletisme desisopodes terrestres. Ann. SpeIeol., 20:489-518.Vandel, A. 1965c. Les Trichoniscidae cavernicoles (Isopodaterrestria; Crustacea) de I'Amerique du Nord. Ann.Speleol.,20:347-389.Vandel, A. 1968a. Description d'un nouveau representant dugenre Cordioniscus (Crustacea, Isopoda, Oniscoidea, Styloniscidae)suivie de considerations sur les voies de migrationde certaines lignees d'isopodes terrestres. Ann.Speleol.,23:621-632.Vandel, A. 1968b. Isopodes terrestres. Pp. 35-168 in Missionzoologique beige aux fles Galapagos et en Ecuador (N. etJ. Leleup, 1964-1965), 1.Vandel, A. 1970. Un troisieme oniscoi"de cavemicole menantune vie aquatique: Mexiconiscus laevis (Rioja). Ann.SpeIeo!., 25:161-177.Vandel, A., and J.-P. Durand. 1970. Le cycle vital du Protee,Proteus anguinus Laurenti (batraciens, urodeles). Compt.Rend. Acad. Sci. Paris, 270:2699-2701.Vandenbussche, E., H. Wijffels, and G. Thines. 1965. Markingsmall tropical fish. Anim. Behav., 13:585-586.Van Name, W. G. 1936. The American land and fresh-waterisopod Crustacea. Bull. American Mus. Nat. Hist., 71.535 pp.Van Name, W. G. 1940. A supplement to the American landand fresh-water isopod Crustacea. Bull. American Mus.Nat. Hist., 77: 109-142.Van Name, W. G. 1942. A second supplement to the Americanland and fresh-water isopod Crustacea. Bull. AmericanMus. Nat. Hist., 80:299-329.Van Note, M. 1977. To the bottom in Yochib. CanadianCaver, 9(2):3-5.Vazquez-Yanes, c., A. Orozco, G. Fran

Villa R., B. 1956. Una extrana y severa mortandad de murcieIagosMormoops megalophylla en el norte de Mexico.Anal. Inst. BioI., Mexico, 26:547-552.Villa R., B. 1958. Pteronotus davyi fulvus. EI murcielago deespaldas desnudas en el norte de Sonora, Mexico. Anal.Inst. BioI., Mexico, 29:375-378.Villa R., B. 1963. Reflexiones acerca de la posicion taxonomicade los murcielagos siricoteros de Mexico, generoGlossophaga. Anal. Inst. BioI., Mexico, 34:381-391.Villa R., B. 1967. Los murcielagos de Mexico. Mexico, D. F.:Inst. BioI., Univ. Nac. Auton. Mexico. xvi + 491 Pi'.Villa R., B., and E. L. Cockrum. 1962. Migration in theguano bat Tadarida brasiliensis mexicana (Saussure). J.Mammal., 43:43-64.Villa R., B., and A. Jimenez G. 1961. Acerca de la posiciontaxonomica de Mormoops megalophyila senicula Rehn, yla presencia de virus rabico en estos murcielagos insectivoros.Anal. Inst. BioI., Mexico, 31:501-509.Villa R., B., and A. Jimenez G. 1962. Tres casos mas de rabiaen los murcielagos de Mexico. Anal. Inst. BioI., Mexico,32:391-395.Villa R., B., and W. Lopez-Forment C. 1966. Cinco casos dedepredacion de pequenos vertebrados en murcielagos deMexico. Anal. Inst. BioI., Mexico, 37:187-193.Villa R., B., G. Meza Ruiz, B. Ortiz Bonilla, and B. Villa Cornejo.1967. Rabia en dos especies de murcielagos insectivorosgenero PteronotIJs, en condiciones naturales, colcctadosen Jalisco, Mexico. Anal. Inst. BioI., lIniv. Nac.Auton. Mexico, 38, Ser. Zool., (1 ):9-16.Villada, M. M. 1910. Reseiia descriptiva y geologica de laGruta de Tonaltongo, del Mineral del Cardonal, que sehalla al paso, y del camino que a aquella conduce, situadosen el estado de Hidalgo. Naturaleza, ser. 3, 1(1 ):2544, pis. 3-10.Villada, M. M. 1911. Breve noticia de un viaje de exploraciona diversos lugares del estado de Veracruz. Naturaleza,ser. 3, 1(2):53-92. pis. 12-21.Villalobos F., A. lY48. Estudios de los cambarinos mexicanos.VII. Descripcion de una nueva especie del generoProcambarus, Procambarus acanthophorus n. sp. Anal.Inst. BioI., Mexico, 19:175-182.Villalobos F., A. 1951. Un nuevo misidaceo de las Grutas deQuintero en el estado de Tamaulipas. Anal. Inst. BioI.,Mexico, 22:191-218.Villalobos F., A. 1953. Distribucion geografica y notas ecologicasde los cambarinos mexicanos. Mem. Congr. Cient.Mexicano,7:343-374.Villalobos F., A. 1954. Estudios de los cambarinos mexicanos.XII, Parte 1. Revision de las especies afines a Procambarusmexicanus (Erichson), con descripcion de nuevasform as. Anal. Inst. BioI., Mexico, 25:299-379.Villalobos F., A. 1955. Cambarinos de la fauna mexicana(Crustacea Decapoda). Tesis. Universidad Nacional Autonomade Mexico. 290 pp.Villalobos F., A. 1960. Un anfipodo cavernicola nuevo deMexico: Bogidiella tabascensis n. sp. Anal. Inst. BioI.,Mexico, 31: 317-334.Villalobos Figueroa, A. 1974. Una nueva especie de Troglocubanus(Crustacea, Decapoda, Palaemonidae), de SanLuis Potosi, Mexico. Anal. Inst. BioI. Univ. Nac. Auton.,Mexico, 42, Ser. Cienc. Mar Limnol., (l): 1-6.Vinson, G. L. 1962. Upper Cretaceous and Tertiary stratigra·phy of Guatemala. Bull. American Assoc. Petrol. Geol..46:425-456.Vives, E. 1975. Un interessant peix cavemicola. Sec. Invest.Subt. Centre Excurs. (Terrassa), 4: 119-123.Vivo Escoto, .J. A. 1964. Weather and climate of Mexico andCentral America. Pp. 187-215 in R. C. West, ed., Handbookof Middle American Indians, Volume 1: Naturalenvironment and early cultures. Austin: Univ. Texas Press.Vomero, V. 1972. A new species of Jamesonia Dusbabek1967 (Acarina, Trombidiformes, Myobiidae) parasitic onthe vampire bat Desmodus rotundus, with a description ofthe nymphal and larval stages. Quad. Accad. Naz. Lincei,Probl. Att. Sci. Cult., 171(1 ):157-172, pI. 1.Vomero, V. 1973. Stato attuale delle conoscenze sugli Histeridaeipogei.lnternatI.J. Speleol., 5:361-367.Vomero, V. 1974. Troglobacanius n. gen. with four new species,a line of cave-adapted Mexican J-Jisteridae (Coleoptera).Quad. Accad. Naz. Lincei, Probl. Au. Sci. Cult.,171(2):325-361.Vomero, V. 1977. Anapleus wenzeli, una nuova specie diDendrophilinae (Col. Histeridae) proveniente da unagroUa del Messico meridionale. Quad. Accad. Naz. Lincei,Probl. Att. Sci. Cult., 171(3):341-348.Voneida, T. J. 1973. A comparative study of retinotectal projectionsin the blind cave characin, Astyanax hubbsi, andits sighted ancestor, Astyanax mexicanus. (Abstr.). Anal.Rec.,175:462-463.Voneida, T. J., and C. !VI. Sligar. 1976. A comparative neuroanatomicstudy of retinal projections in two fishes: Astya·nax hubbsi (the blind cave fish), and Astyanax mexicanus.J. Compo Neurol., 165:89-105.Wagner, F. W. 1977. Descriptions of Centruroides Marx fromthe Yucatan Peninsula (Arachnida, Scorpionida, Buthidae).Assoc. Mexican Cave Stud. Bull., 6:39-4,7.Wagner, P. L. 1964. Natural vegetation of Middle America.Pp. 216-263 in R. C. West, cd., Handbook of MiddleAmerican Indians, Volume 1: Natural environment andearly cultures. Austin: Univ. Texas Press.Walker, B. 1970. A fish without eyes. Aquarium. Philadelphia,3(5):6-7,44-46,48-50.Walker, C. F. 1955a. A new salamander of the genus Pseudoeuryceafrom Tamaulipas. Occ. Papers Mus. Zool. Univ.Michigan, 567. 8 pp., 1 pI.Walker, C. F. 1955b. Two new lizards of the genus T,epidophymafrom Tamaulipas. Occ. Papers Mus. Zoo!' Univ.Michigan, 564. 10 pp.Wall, J. R., G. E. Murray, and T. Diaz. 1961. Geology of theMonterrey area, N. L., Mexico. Trans. Gulf Coast Assoc.Geol. Soc., 11 :57-71.Walls, G. L. 1942. The vertebrate eye and its adaptive radiation.Cranbrook Inst. Sci. Bull., 19. xiv + 785 pp.Walper, .T. 1. 1960. Geology of Coban-Purulha area, AltaVerapaz, Guatemala. Bull. American Assoc. Petrol. Geol.,44:1273-1315.Walsh, M., cd. 1972. Mexican caving of the Southwest TexasGrotto: 1966-1971. San Marcos: Southwest Texas Grotto.146 pp.Walsh, M. 1973. La Gruta del Precipicio. Assoc. MexicanCave Stud. News., 4:100-101.Walters, 1. H., and V. Walters. 1965. Laboratory observationson a cavernicolous poeciliid from Tabasco, 1\1exico.Copeia, 1965:214-223.Walters, V., and R. K. Liu. 1967. Hydrodynamics of navigationby fishes in terms of the mucus-water "interface."Pp. 437-446 in P. H. Cahn, cd., Lateral line detectors.Bloomington: Indiana Univ. Press.286

Ward, H. L. 1891. Descriptions of three new species of Mexicanbats. American Nat., 25:743-753.Ward, H. 1. 1904. A study in the variations of proportionsin bats, with brief notes on some of the species mentioned.Trans. Wisconsin Acad. Sci., Arts, Letters, 14:630-654, pis. 50-55.Warner, D. W., and J. R. Beer. 1957. Birds and mammals ofthe Mesa de San Diego, Puebla, Mexico. Acta Zoo!' Mexicana,2(4-5):1.21, table 1.Warren, R. D. 1961. The obligative cavernicoles of Florida.Spec. Papers Florida Speleol. Soc., 1. 10 pp.Watkins, L. c., J. K. Jones, Jr., and H. H. Genoways. 1972.Bats of Jalisco, Mexico. Spec. Pub!. Mus. Texas TechUniv., 1. 44 pp.Weiss, B. A., and J. L. Martini. 1970. Lateral-line sensitivityin the blind cavefish (Anoptichthys jordani). J. CompoPhysio!. Psycho!., 71:34-37.Welbourn, W. C. 1978. Biology of Ogle Cave with a list of thecave fauna of Slaughter Canyon. Natl. Speleo!. Soc. Bull.,40:27-34.West, R. C. 1964. Surface configuration and associated geologyof Middle America. Pp. 33-83 in' R. C. West, ed.,Handbook of Middle American Indians, Volume 1:Natural environment and early cultures. Austin: Univ.Texas Press.Wharton, G. W. 1933. Acarina of Yucatan caves. CarnegieInst. Washington Publ., 491:137-152.Wheeler, W. M. 1937. Taxonomic notes on Aeromyrmexoctospinosus Reich. Pp. 69-74 in W. M. Wheeler, Mosaicsand other anomalies among ants. Cambridge, Massachusetts:Harvard Univ. Press.Wheeler, W. M. 1938. Ants from the caves of Yucatan. CarnegieInst. Washington PubL 491:251·255.Whitacre, 0.1979. Huastecan memories. Chihuahuan DesertDiscovery, 4:7-8.Whitehead, D. R. 1972. Classification, phylogeny, and zoogeographyof Schizogenius Putzeys (Coleoptera: Carabidae:Searitini). Quaest. Entomo!., 8:131-348.Whitl!head, D. R. 1973. Annotated key to Platynus, ineludingMexisphodrus and most "Colpodes", so far describedfrom North America ineluding Mexico (Coleoptera:Carabidae: Agonini). Quest. Entomol., 9:173-217.Whitley, G. P. 1950. New fish names and records. Proc. Roy.Zoo!' Soc. New South Wales, 1949/50:61-68.Whitt, G. S., and F. S. Maeda. 1970. Lactate dehydrogenasegene function in the blind cave fish, Anoptichthys jordani,and other characins. Biochem. Gen., 4:727-741.Wiley, S., and H.. W. Mitchell. 1971. A bibliography of theMexican eyeless characin fishes of the genus Astyanax.Assoc. Mexican Cave Stud. Bull., 't:231-239.Wilkens, H. 1963. Beitrag zur Degeneration des Auges beiCavernicolen, Genzahl und Manifestationsart (Untersuchungenan mexikanischen Hohlenfischen). ZooI. Anz.,180:454-464.Wilkens, H. 1970a. Beitrage zur Degeneration des Auges beiCavcmicolen, Genzahl und Manifestationsart. Untersuch.ungen an mexikanischen I-Iohlenfischen. Z. Zool. Syst.Evolutionsf. 8:1--1,7.Wilkens, H. 1970b. Beitrage zur Degeneration des Melaninpigmentsbei cavernicolen Sippen des Astyanax mexicanus(Filippi) (Characidae, Pisces). Z. Zool. Syst. Evolutionsf.,3:173-199.Wilkens, H. 1970c. Der Bau des Auges cavernicoler Sippenvon Astyanax /asciatus (Characidae, Pisces). Beitrag zurProblematik degenerativer Evolutionsprozesse. WilhelmRoux Arch. Entwickl.-Mech. Org., 166:54-75.Wilkens, H. 1971. Genetic interpretation of regressive evolutionaryprocesses: Studies on hybrid eyes of two Astyanaxcave populations (Characidae, Pisces). Evolution,25:530-544.Wilkens, H. 1972a. Uber Praadaptationen fUr das Hohlenleben,untersucht am Laichverhalten ober- und unterirdischerPopulationen des Astyanax mexicanus (Pisces).Zool. Anz., 188:1-11.Wilkens, H. 1972b. Zur phylogenetischen Ruckbildung desAuges Cavernicoler: Untersuchungen an A noptichthys jordani(=Astyanax mexicanus), Characidae, Pisces. Ann.Speleol.,27:411-432.Wilkens, H. 1973a. Anciennete phylogenique et degres dereduction chez les animaux cavernicoles. Ann. Speleol.,28:327-330.Wilkens, H. 1973b. Phylogenetic age and degree of reductionof cave animals. Pp. 203-206 in J. H. Schroder, eds.,Genetics and mutagenesis of fish. New York: SpringerVerlag.Wilkens,H. 1973c. Uberdas phylogenetische Alter von Hohlentieren.Untersuchungen tiber die cavernicole SiisswasserfaunaYucatans. Z. Zool. Syst. Evolutions., 11 :49-60.Wilkens, H. 1977a. Genotypic and phenotypic variability incave animals. Studies on a phylogenetically young cavepopulation of Astyanax mexicanus (Filippi) (Characidae,Pisces). Ann. Spel

Wimsatt, W. A. 1969. Transient behavior, nocturnal activitypatterns, and feeding efficiency of vampire bats (Desmo.dus rotundusl under natural conditions. J. Mammal., 50:233·244.Winkelmann, J. R. 1962. Additional records of Mimon cozu·melae. J. Mammal., 43: 112.Wolf, B. 1934-1938. Animalium cavernarum catalogus.Gravenhage: W. Junk. 3 vols.Woodall, H. T. 1941. A new Mexican salamander of the genusOedipus. Occ. Papers Mus. Zoo!' Univ. Michigan, 444.4 pp.Woods, L. P. 1954. Mysterious fishes found in cave pools andstreams. Bull. Chicago Nat. Hist. Mus., 25(11 ):4·5; 25(12):6-7. Reprinted under the title, "Blind fishes found in cavepools and streams," in: Bull. Natl. Speleo!. Soc., 18:24-29.Woods, L. P., and R. F. Inger. 1957. The cave, spring, andswamp fishes of the family Amblyopsidae of central andeastern United States. American Mid!. Nat., 58:232-256.Woolley, T. A. 1969. Ricinuleida, an odd group of chelicerates.Proc. North Central Branch-Entomol. Soc. Ame·rica, 24:150-151.Wygodzinsky, P. 1944. Contribu

Appendix 1LIST OF COLLECTING LOCALITIESThe following is a list of all caves, wells, springs,and other localities in Mexico, Belize, and Guatemalafrom which cave-associated species have been recorded.The localities are listed by state or districtand then by physiographic region within each state.The name of the locality is followed whenever possibleby an approximate location with respect to anearby town, the municipio, elevation, bibliographicreference, and the topographic map on which thelocality occurs. The references cited may include adescription of the locality or, if no description isavailable, the source of the information for the locality.If a map of the cave has been published an"m" is attached to the page number on which themap is printed. Where no reference is given, theinformation is from the files of the Association forMexican Cave Studies (AMCS). An asterisk (*) precedingthe locality name indicates that cave-adaptedspecies are known from it. A plus sign (+) precedingthe locality name indicates that the locality has notbeen visited by members of the AMCS. The namesused are those accepted by the AMCS. Alternatenames are included with a cross-reference to theaccepted name. The approximate locations of allcollecting localities may he found on Fig. 2. Thetopographic maps used for Mexico are those of the1:50,000 series issued by the Direcci6n General deEstudios del Territorio Nacional; the maps used forBelize are those of the 1:50,000 series issued by theBritish Government's Ministry of Overseas Development.Unfortunately, Guatemalan and many Mexicanmaps were unavailable.BAJA CALIFORNIA SUR+Cave 0.8 km S Las Cuevas, Mpo. de Santiago (Jones etal., 1965:54, 56)+Cave 1 km S Las Cuevas, Mpo. de Santiago (Jones et al.,1965:56)+Cuevas de Santiago, Mpo. de Santiago (Hoffmann, 1944:41,108, 1l0, 117)CAMPECHECoastal Plain+*Well, Calle Victoria 49a, Campeche, Mpo. de Campeche(Ruffo and Vigna Taglianti, 1977:167)Cenote de Bocchen-see Cenote de Bolchen*Cenote de Bolchen, 3 km S San Antonio Sacabchen,Mpo. de Calkini (Reddell, 1977b:241)Cueva de la Iglesia, 2 km NW Becabchen, Mpo. deCalkiniCentral Hill DistrictArtificial cave, Cumpich, Mpo. de HecelchakanQuarried cave, 20 km N Champoton, Mpo. de Champoton(Reddell, 1977b:241-243)*Cenote de Cantemo, 1 km N Cantemo, Mpo. deChampoton (Reddell, 1977b:243)Actun Chen, Cumpich, Mpo. de Hecelchakan (Reddell,1977b:244-245)Cueva de Chuina, Chuina, Mpo. de Champoton (Reddell,1977b:243)Cenote Ek Bis, 4 km E Cumpich, Mpo. de HecelchakanCenote Espiritu, 10 km NNW Bolonchenticul, Mpo.de Hopelchen (Reddell, 1977b:245-246)*Aetun Halmensura, 5 km E Cumpich, Mpo. de Hecel·chakan (Reddell, 1977b:245)CHIAPAS*Actun Huachap, 14 km NNW Bolonchenticul, Mpo.de Hopelchen (Reddell, 1977b:246)Grutas de Ixtacumbilxunam-see Grutas de Xtacumbilxunam*Grutas de Monte Bravo, 10 km NW Cantemo, Mpo.de Champoton (Reddell, 1977b:243-244)*Volcan de los Murcielagos, 11 km E Conhuas, Mpo.de Champoton (Reddell, 1977b:244)*Grutas de San Antonio, 10 km NNE Bolonchenticul,Mpo. de Hopelchen (Reddell, 1977b:246-247)*Grutas de San Ignacio, 15 km N Bolonchenticul,Mpo. de Hopelchen (Reddell, 1977b:247)Grutas de San Jose, 15 km NE Bolonchenticul, Mpo.de Hopelchen (Reddell, 1977b:247)Cueva Sascab:i, Ruinas de Chicanna, Mpo. de Champoton(Reddell, 1977b:244)*Grutas de Xkalumkin, 5 km W Cumpich, Mpo. deHecelchakan (Reddell, 1977b:245)*Grutas de Xtacumbilxunam, 2 km SW Bolonchenticul,Mpo. de Hopelchcn (Stephens, 1843:97-104;Reddell, 1977b:247.249)Cueva del Yeso, 4 km W Ruinas de Chicanna, Mpo.de Champoton (Reddell, 1977b:244)Altamirano Region+Small cave near the Pyramid Tzajalalchih, Ocosingo,1130 m, Mpo. de Ocosingo (Goodnight and Goodnight,1953:10)+Cueva de Chital n. J, Rancho Chital, Ocosingo,1390 m, Mpo. de Ocosingo (Sbordoni et al., 1977:46)+*Cueva de Chilal n. 2, Rancho Chital, Ocosingo,1390 m, Mpo. de Ocosingo (Sbordoni et aI., 1977:43m,45)289

+*Cueva de los Chivos, 5 km E Altamirano, Nuevo SantaAna, 1400 m, Mpo. de Altamirano (Sbordoni etal.,1977:49-50)+Cueva presso Cuxulja n. 1, Cuxulja, Ocosingo,1435 m, Mpo. de Ocosingo (Sbordoni et aI., 1977:47)+Cueva presso Cuxulja n. 2, Cuxulja, Ocosingo,1435 m, Mpo. de Ocosingo (Sbordoni et a!', 1977:47,48m)+Cueva de Monte Vidal n. 1, Monte Vidal, Altamirano,1400 m, Mpo. de Altamirano (Sbordoni et aI.,1977:50)+Sumidero del Panteon, Altamirano, 1350 m, Mpo. deAltamirano (Sbordoni et aI., 1977:47)Bochil and Soyalo Region+Cave near Bochi!, Mpo. de Bochi! (Villa R., 1967:224,226)+*Cueva de la Golondrina, 4 km E Bochi!, 1440 m,Mpo. de Bochi! (Sbordoni et a!', 1974:30,32)+*Cueva del Nacimiento del Rio Santo Domingo, FincaSanto Domingo, near Bochi!, 1250 m, Mpo. de Bochi!(Sbordoni et a!., 1974:32)+Sumidero del Naranjo, EI Naranjo, Soyalo, 1540 m,Mpo. de SoyaIo (Sbordoni et aI., 1977:43-44,43m)+*Cueva del Negro, EI Naranjo, Soyalo, 1590 m, Mpo.de Soyalo (Sbordoni et a!., 1977:44m, 45)+Cueva del Puente Redondo, Puente Redondo, Jitotol,1600 m, Mpo. de Jitolol (Sbordoni et aI.,1977:41, 42m, 43)Comiun de Dominguez RegionCueva Chica de Hun Chabin, 1.5 km N Comitan deDominguez, 1700 m, Mpo. de Comitan de DominguezCueva Chica de Hunchenbien-see Cueva Chica deHun Chabin*Cueva del Tio Ticho, 3 km S Comitan de Dominguez,1700 m, Mpo. de Comitan de Dominguez(Sbordoni et a!', 1974:26)Cueva de la Toma de Agua-see Cueva del Tio Ticholxtacomiun Region+Cave 3 km W Ixhuatan, Mpo. de Ixhuatan (Tuttle,1968:787)Cueva de la Frontera-see TABASCO"La Gruta," Ejido Ignacio Allende-see TABASCO+Piccola Grotta in Loc. Malpaso, Malpaso, lxtacomitan,180 m, Mpo. de Ixtacomitan (Sbordoni et aI.,1977:21-22)+Sotano de Malpaso, Malpaso, 2.5 km NE Ixtacomitan,280 m, Mpo. de Ixtacomitan (Sbordoni et a!.,1977:22,22m)MaIpaso Region+Cave in canyon of Rio de la Venta, Lago de Malpaso,600 m, Mpo. de Ocozocoautia (Smith and Alvarezdel Toro, 1977:37)+Cueva del Agua Purificada, Rio Negro, OcozocoautIa,130 m and 115 m, Mpo. de Ocozocoautla (Sbordoniet aI., 1977:26m, 27)+*Cueva del Burro, Lago de Malpaso, Ocozocoautla,125 m, Mpo. de Ocozocoautla (Sbordoni et aI.,1977:23.24,24m)+La Cueva, Lago de Malpaso, Ocozocoautla, 120 m,Mpo. de Ocozocoautla (Sbordoni et a!., 1977:23)+Cueva de las Dos Bocas, at the confluence of the Riode la Venta with the Rio 'egro, Ocozocoautla,120 m, Mpo. de Ocozocoautla (Sbordoni et aI.,1977:25)+Grutas de los Indios, Rio egro, Ocozocoautla,130 m, Mpo. de Cintalapa (Sbordoni et aI., 1977:27)+Cueva de la Mariposa, Rio Negro, Ocozocoautla,125 m, Mpo. de Cintalapa (Sbordoni et aI., 1977:28)+Cueva del Perro de Agua, Rio Negro, near its confluencewith the Rio de la Venta, Ocozocoautla,1I5 m, Mpo. de Ocozocoautla (Sbordoni et a!.,1977:25·26)+Piccola grotta sui Rio Venta, at the confluence of theRio de la Venta with the Rio Negro, Ocozocoautla,145 m, Mpo. de Ocozocoautla (Sbordoniet a!', 1977:25)+Cueva del Venado, Rio Negro, Ocozocoautla, 130 m,Mpo. de Cintalapa (Sbordoni et a!., 1977:27-28)Montebello Region+*Cueva del Arco, San Rafael del Arco, La Trinitaria,1470 m, Mpo. de La Trinitaria (Sbordoni et aI.,1974:23.24,25m)+Cueva de Chinkultic n. 1, near Chinkultic, 30 km SEComitan de Dominguez, 1500 m, Mpo. de La Trinitaria(Sbordoni et aI., 1974:24)+Cueva de Chinkultie n. 2, near Chinkultic, 30 km SEComitan de Dominguez, 1500 m, Mpo. de La Trinitaria(Sbordoni et a!', 1974:24)+Cenote La Cueva, Tziscao, La Trinitaria, 1480 m.Mpo. de La Trinitaria (Sborooni et aI., 1977:52)+Cenote de las Golondrinas, Tziscao. La Trinitaria,1490 m, Mpo. de La Trinitaria (Sbordoni et a!',1977:51·52)+Cueva Leon, 4 km ENE San Lucas, Mpo. de FronteraComalapa (Smith, 1972:117)Grutas de San Francisco-see Grutas de ZapalutaCueva del San Jose del Arco, Lagunas de Montebello,Comitan de Dominguez, Mpo. de La Trinitaria(Peek and Peck, 1973:69)Cueva de La Trinitaria-see Grutas de Zapaluta*Grutas de Zapaluta, 7 km SE La Trinitaria, 1600 m,Mpo. de La Trinitaria (Thompson, 1972:9, 18m;Sbordoni et a!., 1977:51)Motozintia Region+*Sumidero de Canada, Canada, EI Porvenir, 2560 m,Mpo. de EI Porvenir (Sbordoni et a!', 1977:64)+*Grutas de Llano Grande, Llano Grande, La Grandeza,2140 m, Mpo. de La Grandeza (Sbordoni et aI.,1977:63-64)Palenque RegionCueva del Saito de Agua, 15 km SE Palenque, Mpo.de La Libertad (Coons, 1974:17)Rancho del Cielito Region+Cave 3.5 km NNW Galeana, 43.5 km E Altamirano,900 m, Mpo. de Las Margaritas (Pine, 1972:42)+*Cueva de las Canicas, Rancho del Cielito, 12 km fromColonia Galeana, Ocozocoautla, 1350 m, Mpo. deOcozocoautia (Sbordoni et aI., 1974:22m, 29-30)290

+"Cueva del Cerro Brujo, Rancho del Cielito, 12 kmfrom Colonia Galeana, Ocozocoautla, 1320 m,Mpo. de Ocozocoautla (Sbordoni et aI., 1974:2829)San Cristobal de las Casas Region+Cave 2 km W Teopisca, 2120 m, Mpo. de Teopisca(Davis et al., 1964:385)+Cave 5 km from Teopisca, Mpo. de Teopisca (Rudnick,1960:200)+"Well in Casa Bell, San Cristobal de las Casas, Mpo. deSan Cristobal de las Casas (Ruffo and Vigna Taglianti,1977:135)Cueva del Arcotete, 6 km S San Cristobal de lasCasas, Mpo. de San Cristobal de las Casas (Peck andPeck,1973:68-69)+Piccole cavita dell'Arcotete, La Quinta, San Cristobalde las Casas, 2250 m, Mpo. de San Cristobal de lasCasas (Shordoni et aI., 1977:61)+Cueva de las Calaveras, Sa~ Cristobal de las Casas,2265 m, Mpo. de San Cristobal de las Casas (Sbordoniet al., 1977:48m, 55)Sumidero del Camino, 16 km NE Comitan de Dominguez,Mpo. de Comitan de Dominguez+Cueva de la Canada n. I, La Quinta, San Cristohal delas Casas, 2270 m, Mpo. de San Cristobal de lasCasas (Gascoyne and Pratt, 1975:34; Sbordoniet aI., 1977:57-58)+Cueva de la Canada n. 2, La Quinta, San Cristohal delas Casas, 2270 m, Mpo. de San Cristohal de lasCasas (Sbordoni et aI., 1977:58)*Sotano de Cancuc, Cancuc, 11 km NE Tenejapa,Mpo. de Sitala+Sumidero de Casa Clark, San Cristobal de las Casas,2240 m, Mpo. de San Cristobal de las Casas (Sbordoniet aI., 1977:53-54)+Cueva Clara, Finca San Nicolas, San Cristobal de lasCasas, 2330 m, Mpo. de San Cristobal de las Casas(Sbordoni et aI., 1977:59-61, 60m)+Cueva del Coyote, Rancho Nuevo, San Cristobal delas Casas, 2520 m, Mpo. de San Cristobal de lasCasas (Sbordoni et aI., 1977:61-62, 62m)+Cueva de la Cruz Belen, Finca San Antonio de Padua,Comitan de Dominguez, 2210 m, Mpo. de Comitande Dominguez (Sbordoni et aI., 1974:39m, 40-41)Salida de Cruz Pilal, Cruz Pilal, 14 km NNE Tenejapa,Mpo. de Tenejapa (Boon, 1974:11, 12m; Pace,1977:7-8, 9m, 10)Entrance D, Rio Quinta Valley-see Cueva de laCanada n. I+Cueva Encantada, San Cristobal de las Casas, 2300 m,Mpo. de San Cristobal de las Casas (Sbordoni et aI.,1977:54m, 55)+*Cueva de las Florecillas, Colonia San Francisco, Comitande Dominguez, 2265 m, Mpo. de Comit:inde Dominguez (Sbordoni et aI., 1974:36m, 40)Huistan Resurll:ence-see Cueva Mapachero"Cueva de Los Llanos, 15 km ESE San Cristobal de lasCasas, 2295 m, Mpo. de San Cristobal de las CasasCueva Mapachero, 3 km N HUistan, Mpo. de Huistan(Larson, 1975:28, 29m, 30)"Cueva de los Murcielagos, 15 km ESE San Cristobalde las Casas, 2295 m, Mpo. de San Cristobal de lasCasas+Cueva Obscura, Finca San Nicolas, San Cristobal delas Casas, 2300 m, Mpo. de San Cristobal de lasCasas (Sbordoni et aI., 1977:58-59, 59m)+Cueva del Panteon, 2 km WNW Teopisca, 1820 m,Mpo. de Teopisca (Sbordoni et al., 1974:26-28,27m)+*Cueva de la Planta n. I, Las 'Piedrecitas, 6.5 km NSan Cristobal de las Casas, 2180 m, Mpo. de SanCristobal de las Casas (Sbordoni et aI., 1977:55-56)+~·Cueva de la Planta n. 2, Las Piedrecitas, 6.5 km NSan Cristobal de las Casas, 2180 m, Mpo. de SanCristobal de las Casas (Sbordoni et aI., 1977:56)+"Cueva de la Planta n. 3, Las Piedrecitas, 6.5 km NSan Cristobal de las Casas, 2180 m, Mpo. de SanCristobal de las Casas (Sbordoni et aI., 1977:57)Risorgenza de la Planta n. 3-see Cueva de la Plantan.3"Grutas de Rancho Nuevo, 10 km SE San Cristobal delas Casas, 2275 m, Mpo. de San Cristobal de lasCasas (Thompson, 1972:10, 17m; Sbordoni et al.,1974:33-35, map; Shawcross, 1978: 3-4)+Cueva del Rayo de San Felipe, 3.8 km W San Cristobalde las Casas, 2190 m, Mpo. de San Cristobal delas Casas (Sbordoni et aI., 1974:35)+Cueva del Rayo de San Francisco, 4 km from ColoniaSan Francisco, Comitan de Dominguez, 2250 m,Mpo. de Comifan de Dominguez (Sbordoni et aI.,1974:38, 39m, 40)+Cueva de Saclamanton n. 1, Saclamanton, 4 km at100° from San Juan Chamula, 2430 m, Mpo. deSan Juan Chamula (Sbordoni et aI., 1977:48m, 5253)+Cueva de Saclamanton n. 2, Saclamanton, 4 km at100° from San Juan Chamula, 2430 m, Mpo. deSan Juan Chamula (Sbordoni et aI., 1977:53)+Cueva de Saclamanton n. 3, Saclamanton, 4 km at100° from San Juan Chamula, 2430 m, Mpo. deSan JUafl Chamula (Sbordoni et aI., 1977:48m, 53)+Cueva Chica de San Agostin, Rancho de San Agostin,52 km from San Cristobal de las Casas on road toComitan de Dominguez, 2380 m, Mpo. de Comitande Dominguez (Sbordoni et al., 1974:31m, 37)+Cueva Grande de San Agostin, Rancho de San Agostin,51.7 km from San Cristobal de las Casas onroad to Comitan de Dominguez, 2320 m, Mpo. deComitan de Dominguez (Sbordoni et aI., 1974:31m, 38)+Sotano de San Agostin, Rancho de San Agostin, 51.7km from San Cristobal de las Casas on road toComitan de Dominguez, 2320 m (Comitan deDominguez)(Sbordoni et aI., 1974:31m, 38)Pozzo San Agostino-Sotano de San AgostinGruta de San Cristobal-see Grutas de Rancho NuevoChen Sibilmut, 4 km N Huistan, Mpo. de Huistan(Donavan, 1975: 21-22,23 m, 25-27)Chen Senvilmut-see Chen SibilmutChen Ven Sil Mut-see Chen Sibilmut+Cueva de Teopisca, 4.5 km SSW Teopisca, 1700 m,Mpo. de Teopisca (Villa R., 1967:242)tCueva de Tulanca n. 1, 2 km W Rancho de Tulanca,44 km SE San Cristobal de las Casas, 2200 m, Mpo.de Amatenango (Sbordoni et aI., 1974: 35-36,36m)291

Sumidero Yochib, Yochib, 8 km NE Tenejapa, Mpo.de Tenejapa (Lord, 1974:14-15, 17m; Boon, 1975:6-9, 10-lIm, 15-16; Sleele, 1977a:3-9; Sleele,1977b:11-14; Van Note, 1977:3-4, 5m)Simojovel and Tila Region+Cueva del Barrio Francisco Madero n. 1,Tila, 1160 m,Mpo. de Tila (Sbordoni el aI., 1977:37)+Cueva del Barrio Francisco Madero n. 2, Tila, 1160 m,Mpo. de Tila (Sbordoni ct aI., 1977:37-38)+*Cucva de Chanchaniplic, Sitala, 1200 m, Mpo. de SiIala (Sbordoni el aI., 1977:39, 40m, 41)Cueva de Colonia Rincon, near Rincon Chamula,1790 m, Mpo. de Pueblo Nuevo Solistahuacan+Grula de Finca Sanla Anila n. 1, Finca Santa Anila,Simojovel de Allcnde, 830 m, Mpo. de Simojovelde Allende (Sbordoni el aI., 1977:38-39, 38m)+*Gruta dc Finca Santa Anila n. 2, Finca Sanla Anita,Simojovel de Allende, 810 m, Mpo. de Simojovelde Allende (Sbordoni et aI., 1977:39)+Cueva de Nicolas Bravo, Tila, 1160 m, Mpo. de Tila(Sbordoni et aI..1977:36-37, 36m)+Cueva Tehabo oet-oet, Tila, 1050 m, Mpo. de Tila,(Sbordoni et aI., 1977:35-36)Tuxtla Gutierrez RegionCave 2.5 km NE Balneario Chorreadero, 16 km ETuxtla Gutierrez, Mpo. de Chiapa de CorzoCave 2 km SW Las Pililas, \1po. de OcozocoaullaCueva del Aguacero-see Cueva del Eneanlo+Cueva de Berriozabal, Berriozabal, Mpo. de Herriozabal(Chopard, 1947:68)Cueva Chica del Camino, 21 km E Tuxtla Gutierrez,Mpo. de Ixtapa+Cueva del Camino del Aguaeero, Aguacero, Oeozo·eoautla, 630 m, Mpo. de Oeozocoautla (Sbordoniet aI., 1977:30)+Cueva de la Cepona, Tuxtla GUlierrez, 800 m, Mpo.de TuxtJa Gutierrez (Sbordoni et aI., 1974:20-21)Cueva Cerro Hueco, 4 km SE Tuxtla GUlierrez,730 m, Mpo. de Tuxlla Gutierrez, (Sbordoni el aI.,1974:18-19, fig. 5m)+Cueva de la Chepa, 2 km N. 4 km N. or 4 km NETuxtla Gutierrez, 760 m, Mpo. de Tuxtla Gulierrez(Mazzotti, 1940:405)Cueva del Chorreadero, Balneario Chorreadero12 km E Tuxtla Gutierrez, 650 m, Mpo. de Chiap~de Corzo (Thompson, 1972:11-12, 13m, 15-16;Sbordoni et aI., 1974:16-18, 17m; Shawcross et aI.,1974:60, 62m)+Cueva de la Cotorra, Piedra Parada, Ocozocoautla,700 m, Mpo. de OcozocoautJa (Sbordoni et aI.,1977:29)Hoyo de Don Nicho, Rancho del Tempisque, 13 kmW OcozocoautJa, 710 m, Mpo. de OcozocoautJa(Sbordoni el aI., 1974:21; Sbordoni et aI., 1977:31)+Cueva del Encanto, Aguacero, Ocozocoautla, 590 m,Mpo. de OcozocoautJa (Sbordoni et aI., 1977:30;Gascoyne and Pratt, 1975:36m, 37)+Cueva del Muju, Rancho de las Pilitas, OcozocoautJa,680 m, Mpo. de Ocozoeoaulla (Sbordoni et aI.,197Z:32)+Sima del Ojito, Colonia Galeana, 30 km S OcozocoautJa,950 m, Mpo. de Ocozocoautla (Sbordoniet aI., 1974:21-22, 22m)+Cueva de Paso Burro, l3erriozabal, 900 m, Mpo. deBerriozabal (Sbordoni et aI., 1977: 33)Cueva de las Pinas Ramas-scc Cueva de los PinosRamasCueva de los Pinos Ramas, 21 km E Tuxlla Gulierrez,Mpo. de lxtapa+Cucva de la Ramillette., 7 km from the outskirls ofTuxtla Gutierrez in the direction :HO°. 815 m,\Ipo. de Tuxlla Gutierrez (Sbordoni et aI., 1974:19·20)+Cucva del Rancho San Miguel n. 1, Rancho San Miguel,Chiapa de Corzo, 460 m, \1po. de Chiapa deCorzo (SbordorU el aI., 1977:34-35)+Cueva del Rancho San Miguel n. 2, Rancho San Miguel,Chiapa de Corzo, 460 m, Mpo. de Chiapa deCorzo (Sbordoni et aI., 1977:34-35)+Gruta dc Roberlo, Sirada Tuxtla Gutierrez-San Cristobalde las Casas, Chiapa de Corzo, 810 m, \lpo.de Chiapa de Corzo (Sbordoni el aI., 1977::.13-34,34m)+Cueva del Sabin, Rancho del Sabin, Ocozocoautla,890 m, Mpo. de Ocozocoaulla (Sbordoni ct aI.,1977:28-29,29m)Cueva del Tempisque, Rancho del Tempisque, 13 kmW Ocozocoautla, 690 m, i\-lpo. de Ocozocoautla(Sbordoni el aI., 1977:3]-32)Sima del Tempisque n. I-see Hoyo de Don i\iehoSima del Tempisque n. 2-see Cueva del TempisqueCHIHUAHUACarimechi Region+Cave near Carimechi, Mpo. de 'ilJruachic (Anderson,1972:236)+Cavcrn in a canyon near Carimechi, !'Ilpo. dc '!lImachic(Anderson, 1972:239)Magurichic Region+Cave at Magurichic, Mpo. de Magurichic (Knobloch,1942:297)Cave at Mojarachic-sce Cave at MagurichicSalaices Region*Cueva del Diablo, 4 km W Salaices, Mpo. de VillaLopez (Reddell, 1977a:85, pI. 1m)Cueva de los Muehachos, 2 km S Salaices, "Ipo. deVilla Lopez (Reddell, 1977a:85)Santa Elena Region+Cave in Fern Canyon, 2] km NW Santa Elena, Mpo.de Ojinaga (.Judd, 1967:192)+Cave on south side of Canon de Santa Elena, Mpo. deOjinaga (ViUa R., 1967: 383)Sotano de Sauz, 15 km N Manuel Benavides, 1000 m,Mpo. de Ojinaga (Sprouse, 1977:79-80, pI. Vm)Manuel Benavides, H13D44)Santo Tomas RegionSocavon de Santo Tomas, 8 km NNW Santo Tomas,Mpo. de ?Cd. Guerrero (Reddell. 197ia:87)Solano Segundo de Santo Tomas, 8 km ~NW SantoTomas, Mpo. de ?Cd. GuerreroSotano Suciedad de Santo Tomas, 8 km "NW SantoTomas, Mpo. de ?Cd. GuerreroVilla Matamoros RegionCueva del Salitre, 13 km W Villa Matamoros, '\'lpo. deVilla Matamoros (Reddell, 1977a:85, 87)292

COAHUILAAllende Region+Cueva de Allende, Mpo. de ?Allende (Malaga Albaand Villa R., 1957:548,559)El Chiflon Region+Cave at EI Chiflon, 35 km W Saltillo, Mpo. de Saltillo(Baker, 1956:173)Ciudad Acuna Region*Cueva de los Lagos, 24 km W Ciudad Acuna, Mpo. deCiudad AcunaCuatro Cienegas de Carranza Region+Cave 5.3 km NW Cuatro Cienegas de Carranza, Mpo.de Cuatro Cienegas de Carranza (Williams, 1968:24)+*Smalilaguna 7.92 km W, 9.42 km S Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas de Carranza(Holsinger and Minckley, 1971:430) (CuatroCienegas, G13B59)+*Pozo 8.2 km S, 4.7 km W Cuatro Cienegas de Carranza,Mpo. de Cuatro Cienegas de Carranza (Cole andMinckley, 1970:74 (Cuatro Cienegas, G13B59)+*Pozo 20.3 km S, 5.5 km E Cuatro Cienegas deCarranza, Mpo. de Cuatro Cienegas de Carranza(Cole and Minckley, 1970:76 (Cuatro Cienegas,G13B59)+*Pozo 12 km SW Cuatro Cienegas de Carranza, Mpo.de Cuatro Cienegas de Carranza (Minckley andCole, 1968:2) (Cuatro Cienegas, G13B59)+*Small seep near Pozo Barbado, 9.4 km S, 7.9 km WCuatro Cienegas de Carranza, Mpo. de CuatroCienegas de Carranza (Cole and Minckley, 1972:315) (Cuatro Cienegas, G13B59)+*Seep 7.45 km S, 5.50 km W Cuatro Cienegas deCarranza, Mpo. de Cuatro Cienegas de Carranza(Holsinger and Minckley, 1971:430) (CuatroCienegas, G13B59)+*Small spring 7.45 km S, 5.42 km W Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas de Carranza(Holsinger and Minckley, 1971:430) (Cuatro Cienegas,G13B59)+*Small spring 8.2 km S, 8.4 km W Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas de Carranza(Cole and Minckley, 1972:315) (Cuatro Cienegas,G13B59)+*Small spring 8.8 km S, 4.0 km W Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas de Carranza(Cole and Minckley, 1972:315) (Cuatro Cienegas,G13B59)+*Spring 8.15 km S, 2.29 km W Cuatro Cienegas deCarranza, Mpo. de Cuatro Cienegas de Carranza(Holsinger and Minckley, 1971:430) (CuatroCienegas, G13B59)+*Spring-pool, 8.84 km S, 3.96 km W Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas deCarranza (Holsinger and Minckley, 1971:430)(Cuatro Cienegas, G13B59)Sumidero de Alicantre, 16 km W Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas de Carranza+*Pozo Barbado, 9.4 km S, 7.9 km W Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas de Carranza(Cole and Minckley, 1972:320) (Cuatro Cienegas,G13B59)+*Poza de la Becerra, 13.7 km SSW Cuatro Cienegas deCarranza, Mpo. de Cuatro Cienegas de Carranza(Taylor, 1966:162-163) (Cuatro Cienegas, G13B59)+*Pozos Bonitos, Laguna Churince, Churince, 14.73 kmS, 7.05 km W Cuatro Cienegas de Carranza, Mpo.de Cuatro Cienegas de Carranza (Holsinger andMinckley, 1971:430) (Cuatro Cienegas, G13B59)+*Laguna Escobeda, 9.8 km S, 2.3 km W Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas de Carranza(Taylor, 1966:163) (Cuatro Cienegas,G13B59)+Frightful Cave, Site 68 of Taylor's survey, CuatroCienegas basin, Mpo. de Cuatro Cienegas de Carranza(Taylor, 1956:215-234)+*Laguna Juan Santos, 8.5 km S, 8.0 km W Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas deCarranza (Cole and Minckley, 1972:319) (CuatroCienegas, G13B59)+*West Laguna of Mojarral, 11 km SW Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas de Carranza(Cole and Minckley, 1972:319) (Cuatro Cienegas,G13B59)Cueva del Pedregoso, 32 km SE Cuatro Cienegas deCarranza, Mpo. de Cuatro Cienegas de Carranza(Anonymous, 1968:11-12)Pedregoso Circle Cave, 30 km SE Cuatro Cienegas deCarranza, Mpo. de Cuatro Cienegas de CarranzaPedregoso Hidden Cave, 30 km SE Cuatro Cienegasde Carranza, Mpo. de Cuatro Cienegas de CarranzaPedregoso Pipe Cave, 30 km SE Cuatro Cienegas deCarranza, Mpo. de Cuatro Cienegas de CarranzaCueva de San Vicente, 6.5 km S Boquilla, 16 km SECuatro Cienegas de Carranza, Mpo. de CuatroCienegas de Carranza (Garcia Lozano, 1939b:l013, 12m; Rodriguez Cabo, 1953:354-355; Fish,1965:66)Hennanas Region+Cave 10 km E Hennanas, 360 m, Mpo. de Escobedo(Baker, 1956:171)Cueva de la Herradura, 8 km E Hermanas, Mpo. deEscobedo (Fish, 1965:66)Monclova RegionBocas del Cannen, 40 km W Monclova, Mpo. de?SacramentoOjo Caliente Region+Cueva del Socavon del Volcan de Djo Caliente, 2 kmESE Djo Caliente, 16 km ENE Ramos Arizpe,1455 m, Mpo. de Ramos Arizpe (Garcia Lozano,1939b:9; Rodriguez Cabo, 1953:350-351) (RamosArizpe, G14C24)Rancho Guadalupe Region+Cueva de Don Jesus, Rancho Guadalupe, 92 km N,43.5 km W Saltillo, Mpo. de Ramos Arizpe (Baker,1956:173-174)Cueva de San Jesus-see Cueva de Don J erosSan Buenaventura Region+Cave below summit of south face of Canon del RioSalado, 14.5 km W, 6.5 km S San Buenaventura,550 m, Mpo. de San Buenaventura (Baker, 1956:173,183,187)293

Serrania del Burro RegionCave, Hacienda Las Pilas, 100 km SE Boquillas delCarmen, :vJpo. de Zaragoza (Mollhagen, 1977:80)+Spring Cave, C96 of Taylor's survey, in Media LunaCanyon, 26 km S Boquillas del Carmen, \1po. deOcampo (Drake, 1951 :94)Sierra Arteaga RegionGruta de Cuevecillas, 1 km N Cuevecillas, 9 km ENEArteaga, Mpo. de Arteaga (Fish, 1965:68) (Arte.aga, G14C34)Sierra de las Animas RegionCueva de las Animas, 21 km WNW Candela, 760 m,Mpo. de Monclova (Garcia Lozano, 1939b:l0m;Rodriguez Cabo, 1953:352-354, 366m; Reddell,1966c:5-6) (Candela, F14A54)Sierra de Maynin RegionCueva de Empalme, 1.5 km E Entronque la Cuchilla,Mpo. de Viesca (Maynin, G13D27)Cueva de los Escarabajos, 1.5 km E Entronque laCuchilla, Mpo. de Viesca (Mayran, G13027)Cueva de los Grillos, 1.5 km E Entronque la Cuchilla,Mpo. de Viesca (Mayran, G13027)Cueva de Yeoo, 1.5 km E Entronque la CuchiJla,Mpo. de Viesca (\1ayran, G13D27)Sierra de Santa Rosa Region+Cave, north side of Puerto de Santa Ana, 7 km SNacimiento de los Indios, 850 m, \Ipo. de \Iel·chor Musquiz (Baker. 1956:171)Cueva del Leon, Canon de la Alameda, 52 km WNW'\1elchor :Vlusquiz, '.Ipo. de :Vlelchor de Musquiz(Garcia Lozano, 1939b:]7-18; Rodriguez Cabo,1953:356·358)*Pozo de EI Potrero, 8.5 km SSE Melchor \'!usquiz,600 m, Mpo. de !\tlelchor Musquiz (\lelchor Mus·quiz, G14A12)Sierra San Lorenzo Region+Cueva del Buen Abrigo, Buen Abrigo, 19 km N Mata·moros, Mpo. de Matamoros (l3arlow, 1946:266·267) (Matamoros, G] 3026)+Cueva de San Lorenzo, 2 km from Hacienda de SanLorenzo de la Lcgua, 23 km NNE Matamoros,Mpo. de Matamoros (Martinez del Rio, 1956:20)(Matamoros, G13026)Sierra EI Tarillal RegionCueva de las Vigas-see NUEVO LEONSierra Zapaliname Region+Caves 2 km Sand 8 km W Bella Union, 2100 m,Mpo. de Saltillo (Baker, 1956:]87) (Arteaga,G14C34)Torreon Region+Cave 51 km SW San Pedro de las Colonias, llOO m,Mpo. de Matamoros (Baker, 1956:173)+Cave SE of Torreon (Baker, 1956:190)+Cueva de Laguna Seca, 8 km NW :'IIazareno, Mpo. deTorreon (Constantine, 1967:19)Unplaced Cave+Cueva de la Bandurria, 1.5 km N K612 Saltillo.Torreon Highway, 1200 m (Villa R., 1967:370)COLIMA+Cave, Ixtlahuacan, l\1po. de Ixtlahuacan+Cave, Manzanillo, \1po. de Manzanillo (Ingles, 1959:381)+Cueva de la Filirica, 5 km W Coquimatlan, 'VIpo. de Co·quitmatlin (Mazzotti, 1941:313)+Cueva de la Finca, CoquitmatIan, Mpo. de Coquimatlan(Hoffmann, 1953:185,187)DISTRITO FEDERAL+Caves near Ixtapalapa (Goldman, 1951:137)+Lava "blow hole" near San Geronimo (Hubbell, 1972:105)Cueva del Cerro de la Estrella, 2 km S Ixtapalapa (Reddell,1973f:92-93)Cave EI Feuhtle-see Cuevas de Teutli+Cuevas de Teutli (Hoffmeister, 1957:459)+Cueva de Xlctli, Tlalpan (Caballero y C., 1942:652)DURANGOCampana Balcones Region+Cueva de los Indios, 11 km N Campana Ralcones,1140 m, Mpo. de TlahualiJo (Baker and Greer,1962:75)Ciudad Lerdo Region+Cave 16 km S Ciudad Lerdo, 1500 m, Mpo. de Ciu·dad Lerdo (Villa R., 1967:435)+Cueva de Espana, 2 km S, II km W Nazareno, Mpo.de Ciudad Lerdo (Villa R., 1967:59, 67, 442)Cueva del Guano, 20 km S Ciudad Lerdo, Mpo. deCiudad Lerdo (Reddell, 1977a:89-90, pI. 11m)Cueva del Indio-see Cueva de Espana+Cueva de la .Ioya de Lerdo, Ciudad Lcrdo, ;\'1 po. deCiudad Lerdo (Bustamante, 1964:509,513-515)La Pila Region+Fissures 7 km SW La Pila, Mpo. de ?Ourango (Baker,1960:309)Rancho Descubridora Regionde la Siquita, 40 km WNW Mapimi, Mpo. deMapimi (Reddell, 1977a:90-91, pI. IVm)~'CuevaSanta Ana Region+Cave at Santa Ana, 395 m, Mpo. de Coloma (J ones,1964:751)Sierra de la India RegionCueva de la Cucaracha, 7 km S Mapimi, Mpo. de Mapimi(Reddell, ] 977a:89)*Cueva de los Riscos, 7 km S Mapimi, 'I'lpo. de l\lapimi(Reddell, 1977a:90, pI. IIIm)Vicente Guerrero RegionGUERRERO+Cave on Rancho Las Margaritas, SW Vicente Guerrero,2270 m, Mpo. de Vicente Guerrero (Bakerand Greer, 1962:73)+Small caves near Acahuizotla, 850 m, Mpo. de Chilpancingode los Bravos (Lukens and Davis, 1957:3, 4, 11)+Cave 8 km NW Acapulco de Juarez, Mpo. de Acapulcode Juarez (de la Torre, 1955:696)294

+Large crevice, 4 km S Almolonga, 1700 m, Mpo. deTixtIa de Guerrero (Lukens and Davis, 1957:3)+Small cave 2 km SSE Almolonga, l."lpo. de Tixtla deGuerrero (Fish, 1968: 117)+Cave at Atlala, 2 km NE Mexicapan, Mpo. de Teloloapan(Martinez and Villa R., 1940:303)+Large cave 5 km W Mezcala, 600 m, Mpo. de Zumpangodel Rio (Lukens and Davis, 1957:2,4,5,11)+Cave 19 km S Mezcala, Mpo. de Zumpango del Rio (dela Torre, 1955:696)+Sinkhole 3 km W Omiltemi, 2390 m, Mpo. de Chilpancingode los Bravos (Lukens and Davis, 1957:10-11)+Cave, Papayo, 8 m, Mpo. de Coyuca de Benitez (Lukensand Davis, 1957: 10)+Cave system 2 km W Puerto Marquez, Mpo. de Acapulcode Juarez (Mautz and Lopez-Forment, 1978: 311)+Cave on outskirts of Taxeo, Mpo. de Taxeo (Roth,1968:2]) (Taxco, E14A68)+Cave near Cueva de la Laguna Honda, near Yerbabuena,12 km NW Teloloapan, Mpo. de ?Icapulzaco (Martinezand Villa R., 1940:302-303) (NOTE: This is oneof the Cuevas de Yerbabuena)+Cave near Puente de Dios, 1 km NW Yerbabuena, 12 kmNW Teloloapan, 1800 m, Mpo. de 'Ilcapulzaco (Martinezand Villa R., 1940:301) (NOTE: This is one of theCuevas de Yerbabuena)+Cueva de Aeahuizotla, Aeahuizotla, Mpo. de Chilpaneingode los Bravos (Villa R., 1967:97)*Grutas de Aeuitlapan, 2 km ENE Cupango, 12 km NETaxeo, 1470 m, Mpo. de Tctipae (Bonet, 1971:75-78,mapa 15)(Taxeo, E14A68)+Cueva de Agua Brava, Hoya Coralleyo, 12 km NE Taxeo,1180 m, Mpo. de Tetipae (Bonet, 1971:74-75, mapa14; Sbordoni and Argano, 1972:15) (Taxco, E14A68)+Cueva Alpixafia, 7 km NNE Teloloapan, Mpo. de Ixcateopan(Villa R., 1952:325, 327, 328)+Cucva de los Amerieanos, 10 km SSW Teloloapan, Mpo.de Teloloapan (Villa R., 1967:305)+Cueva del Borrego, E of Omiltemi on highway to Amojileea,:VI po. de Chilpaneingo de los Bravos (Barrera,1958:90,91)*Grutas de Caeahuamilpa, 16 km NE Taxeo, UOO m,:Vlpo. de Tetipae (Bonet, ]971:61-71, mapa 12; Sbordoniand Argano, 1972:16-17; Coons, 1976:35.36,39m) (Taxeo, E14A68)+Grieta de las Caeanieuas, 6 km W San Miguel Totolapan,Mpo. de San Miguel Totolapan (Villa R., 1967: 153)+Gruta de Calieanto, 8 km SE Teloloapan, 1400 m, Mpo.de Teloloapan (Martinez and Villa R., 1940:297-298)Gruta de Cal y Canto-see Gruta de Calicanto+Cueva del Canon del Zopilote, 12.5 km S Mezcala,450 m, 'VI po. de Zumpango del Rio (Villa R. and JimenezG., 1962:392)+Cucva de Carlos Pacheco, 400 m SSE Grutas de Cacahuamilpa,15 km NE Taxco, 1060 m, Mpo. de Tetipac(Bonet, 1971:71-74, mapa 13) (Taxeo, E14A68)+Cueva del Coyote, 21 km N Taxeo, 1600 m, Mpo. de Pilcaya(Bonet, 1971:47)Gruta de Cuetzala-see Grutas de Cuetzala del Progreso+Grutas de Cuetzala del Progreso, 6 km N Cuetzala delProgreso, 904 m, Mpo. de Cuetzala del Progreso (Martinezand Villa R., 1940:298-300)+Cueva del Diablo, 0.5 km SE San Miguel Acuitlapan,9 km NE Taxeo, 460 m, Mpo. de Taxeo (Bonet, 1971:79-80, mapa 16)*Grutas de la Estrella, 21 km N Taxco, 1580 m, Mpo. dePileaya (Sbordoni and Argano, 1972:14-15; Bonet,1971 :52-57, mapa 8) (Taxeo, E14A68)+Cueva el Fraile, Teloloapan, Mpo. de Teloloapan (Horstand Langworthy, 1972:903)Grutas de las Granadas, 4.5 km SE San Miguel Aeuitla.pan, 10 km ENE Taxeo, 1300 m, Mpo. de Taxco(Harmon, 1979:102-103m) (Taxco, E14A68)+Cueva del Huarache, 14 km S Chilpaneingo de los Bravos,Mpo. de Chilpaneingo de los Bravos (Villa R.,1967:202-203)+Cueva de las Juntas, 2 km E Aeahuizotla, Mpo. de Chilopancingo de los Bravos (Villa R., 1967:90,91,191)~'Grutas de J uxtlahuaca, 6 km NNW Colotlipa, 765 m,Mpo. de CuautotitIan (Roy, 1974:39-42, map)+Cueva de la Laguna Honda, Yerbabuena, 12 km NW Teloloapan,1800 m, Mpo. de ?Ixcapuzaleo (RamirezPulido and Alvarez, 1972:251, 258) (NOTE: This isone of the Cuevas de Yerbabuena)Cueva de la Mariposa-see Grutas de EI MogotePozo Melendez, Puente Campuzano, 12 km S Taxeo,Mpo. de ?Taxeo (Thompson, 1970:58-62, 59m)(Iguala, E14A78)+Cueva de la Mesa de las Choas, 5.3 km E San MiguelTotolapan, 280 m (Villa R., 1967:178,202,205)*Cueva Chiea de EI \1ogote, 0.5 km E EI Mogote, 10 kmNNE Taxeo, 1480 m , lpo. de Tetipae (Fish andReddell, 1965:76) (Taxco, E14A68)*Grutas de EI Mogote, 0.5 km E EI Mogote, 10 km NNETaxeo, 1480 m, Mpo. de Tetipae (Bonet, 1971 :59-61,mapa 10)(Taxeo, E14A68)+Cueva Ojo de Agua de Chapa, 7 km SSE Teloloapan,1400 m, Mpo. de Teloloapan (Villa R., 1967:333)+Cueva de OSlotilan, \1po. de Teloloapan (Villa R., 1967:63)Grutas de Pacheco-see Gruta de Carlos PachecoCueva del Palo I:llaneo-see Cueva del Huarache+Cueva de la Pedrera de Cajeles, near Aeahuizotla, 30 kmS Chilpancingo de los Bravos, Mpo. de Chilpaneingo delos Bravos (Ramirez-Pulido and Sanchez-Hernandez,1971:481)+Cueva de la Penita, 1.6 km S Palo Blanco, Mpo. de Chilpaneingode los Bravos (Fish, 1968:] 23)+Cueva del Puente de Dios, 1 km NW Yerbabuena, 12 kmNW Teloloapan, 1700 m, Mpo. de ?Ixeapuzaleo (Martinezand Villa R., 1940:301) (NOTE: This is one ofthe Cuevas de Yerbabuena)Gruta "EI Resuello"-see Grutas del Rio Chontalcoatlan+Cueva del Rincon, near Rincon, 790 m, Mpo. de Telolo·apan(Lukensand Davis, 1957:5)+Grulas del Rio Chontaleoatlan, 12 km NE Taxeo,1180 m, Mpo. de Tetipae (Sbordoni and Argano,1972:16; Coons, 1976:37-38, 39m) (Taxeo, E14A68)+Grutas del Rio San Jeronimo, 16 km NNE Taxeo,1200 m, Mpo. de Telipae (Coons, 1976:37-38, 39m)(Taxeo, E14A68)+Cueva de EI Salitre, 12 km N Zaeatula, Mpo. de LaUnion (Alvarez, 1968:24)+Cueva de San Ignacio, near Aeahuizotla, Mpo. de Chilpancingode los Bravos (Lopez-Forment et aI., 1971:227)+Cueva de Suanehe, Piedreas Negras, 17 km N Taxeo,1280 m, Mpo. de Pileaya (Bonet, 1971:57-59, mapa 9)(Taxeo, E14A68)+Cueva Teeabra, Ahuaeatitlan, 1400 m (Villa R., 1967:430,431)295

+Cueva de Tezoapa, 1.5 km E Acahuizotla, 650 m, Mpo.de Chilpancingo de los Bravos (Villa R. and LopezForment,1966:192)+Cueva de Tia Juana, 1.5 km SSW Yerbabuena, 12 kmNW Teloloapan, 1840 m, Mpo. de ?Ixcapuzalco(Villa R., 1963:390)+Cueva de Tlachalitla, 1.5 km SE Apetlanca, 50 m, Mpo.de Cuetzala del Progreso (Malaga Alba and Villa R.,1957:538)+Cueva de la Tranca de Ixcapaneca, 1 km NW Yerbabuena,12 km NW Teloloapan, Mpo. de ?Ixcapuzalco(Martinez and Villa R., 1940:302) (NOTE: This is oneof the Cuevas de Yerbabuena)+Cuevas de Yerbabuena, near Yerbabuena, 12 km NWTeloloapan, 1800 m, Mpo. de ?Ixcapuzalco (Martinezand Villa R., 1940:301-303) (INCLUDES: Cueva dePuente de Dios, Cueva de la Laguna Honda, Cueva dela Tranca de Ixcapaneca, and three unnamed caves)HIDALGOJacala Region+Cave, Durango, Mpo. de Zimapan (Woodall, 1941:1)+Cave W of Pinalito, Colonia Santa Maria, Mpo. deJacala (Villa R., 1967:208)+Cueva de Belen, 500 m N Encarnacion, 2000 m,Mpo. de Zimapan (Barrera, 1951:200)*Sotano del Hondo de Pinalito, near Pinalito, Mpo. deJacala (Bittinger, 1975:13, 14m)+Cueva de la Mariposa, J acala, 1415 m, Mpo. de J acala(Villa R., 1967:327)Cueva de El Ocote, 1.5 km N Palomas, 1600 m, Mpo.de Chapulhuacan (Hendrichs and Bolivar, 1966:9)*Cueva Piedra Ancha, Puerto Obscuro, 19 km by roadSW Chapulhuacan, Mpo. de Chapulhuacan (Fishand Reddell, 1965:74)Cueva del Puerto de la Zorra, Puerto de la Zorra,10 km NE J acala, Mpo. de J acalaCueva de Santa Ana, Santa Ana, Mpo. de ChapulhuacanCueva de El Tenango, Rancho de El Tenango, 3 kmNW Chapulhuacan, Mpo. de Chapulhuacan (Fishand Reddell, 1965:73)Lagunillas Region*Grutas de Xoxafi, 5.6 km N Lagunillas, 1833 m,Mpo. de Santiago de Anaya (Fish and Reddell,1965:74)Tonoltongo RegionGrutas de Tonoltongo, 16 km NE Cardonal, Mpo. deCardonal(Villada, 1910:31-33, pI. VlII·X)Zacualtipan RegionJALISCO+Cave 3 km S Zacualtipan, Mpo. de Zacualtipan (Carterand Jones, 1978:8)+Small cave 2 km NNW Barra de Navidad, Mpo. de Cihuatlan(Watkins et aI., 1972:7,19)+Large cave 3 km E Bolanos, Mpo. de Bolanos (Watkins etal., 1972:8, 12)+Cave, Cuetzmala, Mpo. de La Huerta (Watkins et "al.,1972:8)+Cave, Rancho del Preciado, near El Grullo, Mpo. de EIGrullo (Johnson, 1948:191)+Limestone sink 9.5 km SW and 10.5 km E Pihuamo,Mpo. de Pihuamo (Thompson, 1968:160)+Cave 3 km ESE Plan de Barrancas, Mpo. de Hostotipa·quillo (Watkins et al., 1972:8)+Cueva "El Chico," 5 km NW Tuxcacuesco, Mpo. de Tuxcacuesco(Hoffmann, 1962:222)+Cueva D. C. B., 7.5 km W Jamay, Mpo. de Jamay (VillaR. et aI., 1967:10)+Cueva de las Garrochas (three caves), 17 km NNW Soyat·Ian del Oro, Mpo. de Atengo (Watkins et aI., 1972:30,32)+Cueva de Gines, Etzatlan, Mpo. de EtzatIan (Watkins etaI., 1972:23)+Cueva Hedionda, 10 km S Huascato, Mpo. de DegolladoLa Rivera (Watkins et aI., 1972:14)MEXICO+Cave in Paso OyamecaIco, 25 km N Coatepec Harinas,Mpo. de Coatepec Harinas (Villa R., 1967:428-429)+Cave near Zacazonapan, Mpo. de Zacazonapan (Burns,1970:391)+Cueva de la Barranca de los ldolos, west face of Barrancade los Idolos, 32 km SW or 35 km W Ciudad deMexico (Villa R., 1967:309,367)+Cueva de Coatepec Harinas, 1 km SE Coatepec Harinas,Mpo. de Coatepec Harinas (Bonet, 1971:50-52,mapa 7; Sbordoni and Argano, 1972:14)+Cueva del Diablo, La Pefia, Valle de Bravo, 1800 m,Mpo. de Valle de Bravo (Villa R., 1967:430, 433)Grutas de la Estrella-see GUERRERO+Cueva de la Pena Blanca, Valle de Bravo, Mpo. de Vallede Bravo (Hoffmann et aI., 1978:41)MICHOACAN+Caves on ranches near arroyo of Aguililla River, NEAguililla, Mpo. de Aguililla (Johnson, 1948:191)+Cave at Apatzingan, 315 m, Mpo. de Apatzingan (Halland Villa R., 1949:440)+Cave on limestone hill at eastern edge of Coalcoman deMatamoros, Mpo. de Coalcoman de Matamoros(Hooper, 1961:121)+Cave at El Guyabo, 34 km S Uruapan, Mpo. de ?Uruapan(Hall and Villa R., 1949:441)+Cave 3 km W Patzcuaro, 2330 m, Mpo. de Patzcuaro(Hall and Villa R., 1949:441)+Cave 3 km N Patzcuaro, 2150 m, Mpo. de Patzcuaro(Handley, 1959:149)Cave 30 km W Zamora, 1900 m, Mpo. de Villa MarLas Grutas, 8 km W Ciudad Hidalgo, Mpo. de CiudadHidalgo+Cueva de la Arena, 5 km SW Jacona, 1550 m, Mpo. deJ acona (Villa R., 1967:327, 430, 433)+Cueva de las Caleras, Ciudad Hidalgo, Mpo. de CiudadHidalgo (Nesbitt, 1949:64, 66)+Cueva del Cerro del Borrego, 10 km from Hacienda SanJose de Chila, Mpo. de Apatzingan (Tellez Giron,1944:37-38)+Cueva de la Colmena, Cerro La Beata, 3 km NW Tangancicuaro,Mpo. de Tangancicuaro (Villa R., 1967:327)+Cueva de la Estancia, near Los Bancos, Mpo. de ?Paracuaro(Tellez Giron, 1944:37)Cueva de la Isla J anitzio-see Cueva de J anitzio296

+Cueva de Janitzio, Lago de Patzeuaro, 2200 m, Mpo. dePatzeuaro (Ueshima, 1968:145,146)+Cueva de los Monos, 10 km NW Aguililla, Mpo. de Aguililla(Tellez Giron, 1944:38)+Cueva Prieta, Rancho Agua Fda, SE of Aguililla, Mpo.de Aguililla (Tellez Giron, 1944:38)Grutas de Pujido, Mpo. de ChineuilaMORELOS+Small cave 8 km NE Alpuyeea, 18 km SE Cuernavaea,Mpo. de Xoehitepee (Davis and Russell, 1954:69)(Cuemavaea, E14A59)+Cave at Amaeuzae, 20 km ENE Taxeo, 900 m, Mpo. deAmaeuzae (Davis and Russell, 1954:67 (Taxeo,E14A68)+Cave on Cerro Frio, 21 km SW Jojutla de Juarez, Mpo.de Puente de Ixtla (Ward, 1904:654 (Tilzapotla,EI4A79)(NOTE: May be in Guerrero)+Cave near Cuautla, Mpo. de Cuautla (U. S. Bureau ofSport Fisheries and Wildlife, 1970:15)+Cave at southern edge of Cuernavaea, Mpo. de Cuernavaea(Flores Crespo et aI., 1970:627) (Cuernavaea,E14A59)+Shallow cave, Cuernavaea, Mpo. de Cuernavaea (Roth,1968 :20) (Cuernavaea, E14A59)+Cave, Las Fuentes de Chapultepee, 8 km SE Cuernavaea,Mpo. de Cuernavaea (Herrera, 1911:4) (Cuernavaea,E14A59)+Large deep cave 3 km S Jonaeatepee, Mpo. de J onaeatepee(Davis and Russell, 1954:68)+Cave near Temixeo, 6 km S Cuernavaea, Mpo. de Temixco(Barbour, 1945b:80) (Cuernavaea, E14A59)+Small eave at Tepoztlan, 15 km NE Cuernavaea, Mpo. deTepoztlan (Chopard, 1947:67) (?Cuernavaea, E14A59)+Cave near Laguna Tequesquitengo, 8 km W Jojutla deJuarez, Mpo. de Jojutla de Juarez (Grummon and Novick,1963:363) (Jojutla de Juarez, E14A69)+Small cavern near Zaeoalpan de Amilpas, Mpo. de Zaeoalpande Amilpas (Ward, 1904:645)+Cueva del Amate, near Cuernavaea, Mpo. de ?Cuernavaea(U. S. Bureau of Sport Fisheries and Wildlife, 1970:19) (Cuernavaea, E14A59)+Cueva de la Barranca de Apatlaeo, near Xoehitepee,16 km S Cuernavaea, Mpo. de Xoehitepee (MalagaAlba and Villa R., 1957:536)+Cueva del Cerro, 1 or 3.4 km S Laguna Tequesquitengo,11 km SW Jojutla de Juarez, 840 m, Mpo. de Jojutlade Juarez (Malaga Alba and Villa R., 1957:335) (Jojutlade Juarez, E14A69)+Cueva 18 de Julio, 9 km SSW Temixeo, 15 km SSWCuernavaea, Mpo. de Temixeo (Villa R., 1967:327,333) (Cuernavaea, E14A59)+Cueva del Gallo, Tieuman, 21 km SW Cuernavaea, Mpo.de Tlaltizapan (Villa R., 1967:153) (Cuernavaea,E14A59)+Cuevas de Huajintlan, 0.5 km NW Huajintlan, 19 kmENE Taxeo, Mpo. de Amaeuzae (Davis and Russell,1954:67, 68)(Taxeo, E14A68)+Cueva del Idolo, 1 km S Tequesquitengo, 9 km WSWJojutia de Juarez, 950 m, Mpo. de Jojutla de Juarez(Villa R., 1963:384, 387, 390 (Jojutla de Juarez,E14A69)+Cueva de Leona, District of Juarez, Mpo. de ?Jojutla deJuarez (Ward, 1904:653)+Cueva de Miehapa, in Canada de Miehapa, Mpo. de Coeoyotla(Bonet, 1971:48) (?Tenaneingo, E14A58)+Cueva de los Muiieeos, near Cuemavaea, Mpo. de ?Cuer·navaea (U. S. Bureau of Sport Fisheries and Wildlife,1970:19) (Cuemavaea, E14A59)+Cueva del Mureielago, 6.5 km NW Tres Cumbres,3400 m, Mpo. de Huitzilae (Villa R., 1967:367, 368,369,428,430)Cueva Oeho de Julio-see Cueva 18 de Julio+Cueva de Oxtoyahualoleo, 6 km NNE Tepoztlan,2000 m, Mpo. de Tepoztlan (Malaga Alba and Villa R.,1957:542)+Cueva Palmira III, 6 km S Cuernavaea, Mpo. de Cuernavaea(U. S. Bureau of Sport Fisheries and Wildlife,1971:16) (Cuemavaea, E14A59)+Cueva de Palo Bolero, Palo Bolero, 17 km S Cuernavaea,Mpo. de Xoehitepee (Villa R., 1967:49)+Cueva Poza de Moetezuma, 1 km E, 2 km E, or 2 km WOaxtepee, Mpo. de ?Yautepee (Villa R., 1967:67,69,176,177,178,180,203,457)Cueva de la Presa de Moetezuma-see Cueva Poza deMoetezuma+Cueva del Rio Palmira, S of Cuernavaea, Mpo. de Cuernavaea(U. S. Bureau of Sport Fisheries and Wildlife,1971:16) (Cuernavaea, E14A59)+Cueva del Rio Tetlama, 12 km NNW Cuernavaea, Mpo.de Temixeo (Villa R., 1967:156) (Cuernavaea,E14A59)+Cueva del Salitre, Tequesquitengo, 8 km W Jojutla deJuarez, Mpo. de Jojutla de Juarez (Handley, 1966:300) (Jojutla de Juarez, E14A69)+Cueva del Salitre, 3 or 5 km S Tetaealita, 15 km N Jojutlade Juarez, 1152 m, Mpo. de Emiliano Zapata(Malaga Alba and Villa R., 1957:536, 537, 542)(?Jojutla de Juarez, E14A69)+Cueva del Salitre, 10 km NNW Tieuman, 14 km SE Cuernavaea,Mpo. de Tieuman (Villa R. and Jimenez G.,1961 :504) (Cuernavaea, E14A59)+Cueva del Salitre, 4 km W Xoehimanea, 17 km SE Cuernavaea,1600 m, Mpo. de Tieuman (Villa R., 1967:245)Cueva de Xoehitepee-see Cueva de la Barranca de ApatlaeoNAYARIT+Caves on Isla Maria Madre (Nelson, 1899:18)+Cueva del Fuerte de San Bias, on cliff facing San Bias,Mpo. de San BIas (Villa R., 1967:231)+Cueva del Tesoro, 50 km SSE Tepie, Mpo. de SantaMaria del Oro (Villa R., 1967:305)+Cueva de Ventanas, 2 km S Manzanillo (Villa R., 1967:152)NUEVO LEONCalion Huasteeo RegionGruta Aguila del Oro, 2 km S Santa Catarina, 1740 m,Mpo. de Santa Catarina (Fish and Reddell, 1967:82; Sumbera, 1972:118-119 (Garza Garcia,G14C25)Grutas de San Bartolo-see Gruta Sur de San Bartoloand Gruta Norte de San BartoloGruta Norte de San Bartolo, 9 km S Santa Catarina,900 m, Mpo. de Santa Catarina (Russell andRaines, 1967:21, 22m) (Garza Garda, G14C25)297

*Gruta Sur de San Bartolo, 9 km S Santa Catarina,900 m, Mpo. de Santa Catarina (Raines, 1968b:140-141,143) (Garza Garcia, G14C25)Cueva de Tio Bartolo-see Gruta Norte de San Bartoloand Gruta Sur de San BartoloCanon de Iturbide Region+*Cueva La Charrera, 27.3 km SW Linares, Mpo. deLinaresCueva del Ehanito-see Cueva del Nacimiento de ElEhanito+Cueva de Guadalupe, Canon de Itiurbide, 25 km SWLinares, Mpo.de Linares (Malaga Alba and Villa R.,1957:539)*Cueva del Nacimiento de El Ebanito, El Ebanito,10 km ENE Iturbide, Mpo. de Linares (Russell,1973b:280) (lturbide, G14C67)Cerro de la Boca Region*Cueva de Chorros de Agua, 23 km SW Montemorelos,700 m, Mpo. de Montemorelos (Russell and Raines,1967:29,38) (Montemorelos, G14C47)Cerro Potosi Region+Cave, Lorna La Cueva, 1.5 km SW San Jose La Boya,27 km WNW Galeana, 2720 m, Mpo. de Galeana(Radovsky et a!., 1971:738) (Galcana, G14C56)Cuesta de Chipinque RegionSmall caves on Cuesta de Chipinque-see Caves 1-3*Cave No.1, Cuesta de Chipinque, Monterrey, 1500 m,Mpo. de Monterrey (Peck and Peck, 1973:64)(Garza Garcia, G14C25)Cave No.2, Cuesta de Chipinque, Monterrey, 1500 m,Mpo. de Monterrey (Peck and Peck, 1973:64)(Garza Garcia, G14C25)Cave No.3, Cuesta de Chipinque, Monterrey, 1500 m,Mpo. de Monterrey (Peck and Peck, 1973:64)(Garza Garcia, G14C25)Galeana RegionChevy Sink, 7 km SW Galeana, 1780 m, Mpo. deGaleana (Galeana, G14C56)Pozo de Gavilan, 7 km SW Galeana, 1780 m, Mpo. deGaleana (Russell and Raines, 1967:31, 38, 40m)(Galeana, G14C56)Cueva del Limon, 1.2 km E San Marcos, 15 km SGaleana, Mpo. de Galeana (San Jose de Raices,G14C66)*Resumidero de Pablillo, 4 km NNE Pahlillo, 26 kmSE Galeana, 2000 m, Mpo. de Galeana (Russelland Raines, 1967:31,38, 41m) (Iturhide, G14C67)Sotano de la Pina-see Sotano del PinoSotano del Pino, 6 km SSW Galeana, 1700 m, Mpo.de Galeana (San Jose de Raices, G14C66)Cueva Sur del SOlano de Dos Cuevas, 1.5 km S SantaFe, 15.5 km ESE Galeana, 1800 m, Mpo. de Galeana(Reddell, 1966d:49) (San Jose de Raices,G14C66)Cueva de Yeso Blanco, 6 km SW Galeana, 1740 m,Mpo. de Galeana (Galeana, G14C56)Monterrey Region+Well, Monterrey, Mpo. de Monterrey (Packard, 1894:732)Pico de Carrizal Region*Gruta de Carrizal, 10 km SW El Candela, Mpo. deLampazos de Naranjo (Russell and Raines, 1967:13,14m)Potrero Redondo RegionSotano de la Anticlina, 9 km SSW Villa de Santiago,Mpo. de Villa de Santiago (Raines, 1972b:96)(Allende, G14C36)Sotano de Potrero Redondo, 17 km S Villa de Santiago,Mpo. de Villa de Santiago (Allende,G14C36)Redondo Pit Cave-see Sotano de Potrero RedondoPurificacion RegionCueva del Borrego-see TAMAULIP ASCueva del Brinco--see Sistema Purificacion, TAMAlJ-LIPASCueva de California-see TAMAULIPASCueva del Camino-see TAMAULIPASCueva de Chuparosa-see TAMAULIPASSotano de.Jesus-see TAM AU LIP ASCueva del Ojo de Agua, 4 km WSW Dulces Nomhres,Mpo. de Aramherri (Casas Reales, Fl4A18)*Cueva Chica del Ojo de Agua, 4 km WSW DulcesNombres, Mpo. de Aramherri (Casas Reales,Fl4A18)Cueva de los Parajitos, 2 km N Tinajas (=Los Tanques),7 km SE Dulces Nombres, Mpo. de Aramberri(Casas Reales, F14A18)Pozo de los Penuelos-see Sotano de las Penue!as*Sotano de las Penue!as, 3 km SW Dulces Nombrcs,2010 m, Mpo. de Aramberri (Casas Reales,F14A18)Sotano de Rancho Nuevo n. I-see TAMAULIPASSotano de Rancho Nuevo n. 2-see TAMA ULIP ASSotano de Rancho Nuevo n. 3-see TAMAULIPASSotano de Rancho Nuevo n. 4-see TAMAULIPASCueva de Rancho Revilla-see Cueva de Revilla,TAMAULIPASCueva del Tecolote-see Cueva de \)esmontes, TAMAULIPASCueva Vrincon-see Sistema Purificacion, TAMA ULIPASSierra Las Animas Region+Puente de Dios, 7 km NNW Galeana, 1460 m, Mpo.de Galeana (Russell and Raines, 1967: 38) (Galeana,G14C56)Sierra del Fraile RegionGrutas de Garcia-see Grutas de Villa de Garcia+Cueva del Rincon de la Virgen, 6.8 km N Villa deGarcia, 1200 m, Mpo. de Villa de Garcia (MalagaAlha and Villa R., 1957:559,564,566) (Hidalgo,G14C15),x'Grutas de Villa de Garcia, 7 km NW Villa de Garcia,1060 m, Mpo. de Villa de Garcia (Russell andRaines, 1967:19, 20m)(Hidalgo, G14C15)Sierra de Garia RegionCueva de Constantin, 8 km S Espinazo, Mpo. deMina (Bittinger, 1972:56, 57m) (Reata, G14A83)Sierra de Gomas RegionCave 7 km SSW Bustamante, Mpo. de Bustamante298

Pit 7 km SSW Bustamante, Mpo. de Bustamante(Fromen, 1965:44-45 )*Grutas del Palmito, 7 km SSW Bustamante, Mpo. deBustamante (Russell and Raines, 1967:16, 1718m; Ediger, 1970:3-7, 5m)Cueva del Precipicio, 9 km SW Bustamante, Mpo. deBustamante (Russell, 1973a:363-365; Walsh, 1973:100-101)Sierra de Iguana Region+Cueva del Diablo, 7 km by road W Sabinas Hidalgo,Mpo. de Sabinas Hidalgo (Hershberger, 1967:8889,90m)+Cueva Envenenada, 7 km by road W Sabinas Hidalgo,Mpo. de Sabinas Hidalgo (Aguirre Pequeno, 1959:245,246)Sierra de Santa Clara RegionCueva del Camino-see Cueva de las FisurasCueva de las Fisuras, 16 km by road W Sabinas Hidalgo,Mpo. de ?Sabinas Hidalgo (Russell and Raines,1967:21)Sierra de la Silla Region*Flooded mine below Cueva de la Boca, 3 km E Villade Santiago, 500 m, Mpo. de Villa de Santiago(Peck and Peck, 1973:63) (Allende, G14C36)*Cueva de la Boca, 3 km E Villa de Santiago, 540 m,Mpo. de Villa de Santiago (Russell and Raines,1967:26,35, 36m, 37) (Allende, G14C36)Cueva de la Villa de Santiago-see Cueva de la BocaSierra EI Tarillal RegionCueva de las Vigas, EI Carmen (=Las Vigas), 22 km EArteaga, 2300 m, Mpo. de Santa Catarina (Reddell,1966d:53) (San Antonio de las Alazanas, G14C35)Zaragoza RegionOAXACACueva de Cuesta Blanca, 3 km W Zaragoza, Mpo. deZaragoza (Russell and Raines, 1967: 31, 38) (Zaragoza,Fl4A17)Sotano de Cuesta Blanca, 3 km W Zaragoza, Mpo. deZaragoza (Russell and Raines, 1967:38) (Zaragoza,Fl4A17)Dolina Cave, 4 km N Zaragoza, Mpo. de Zaragoza(Zaragoza, F14A17)Acatlan Region+Cave on Cerro Pinon, 12 km S Acathin, Mpo. deAcatlan (Constantine, 1966:126)Cave near Los Corrales, 11 km SSW Acatlan, Mpo. deAcatlan+Cave 1.5 km NW Los Limones, 18 km S Acatlan,Mpo. de Acatlan (Constantine, 1966:125)+Cave 5 km W Los Limones, 18 km S Acatlan, Mpo.de Acatlan (Constantine, 1966:125)Cueva de Campo Chico, 9 km SSW Acatlan, Mpo. deAcatlan*Cueva de Cayetano, 10 km SW Acatlan, Mpo. deAcatlan*Cueva de Los Corrales, 2 km W Los Corrales, 11 kmSSW Acatlan, Mpo. de Acatlan*Cueva de la Culebra, 10 km SW Acatlan, Mpo. deAcatlan*Cueva Desapareciendo, 2 km SW Acatlan, Mpo. deAcatIanSotano Desapareciendo, 2 km SW AcatJan, Mpo. deAcatlan*Cueva de la Finca, 10 km SW Acatlan, Mpo. de AcatIan*Cueva de Juan Sanchez, 10 km NW Acatlan, Mpo. deAcatlan (Byrd, 1976:23, 24m)*Cueva Chica de Juan Sanchez, 12 km NW Acatlan,Mpo. de AcatlanCueva de la Junta, 5 km SW Acatlan, Mpo. de AcatIan (Reddell, 1973e:88-S9)*Cueva de la Laguna Verde, 10 km SW Acatlan, Mpo.de Acatlan (Coons, 1977:18-20, 21m)*Cueva del Lencho Virgen, 9 km SSW Acatlan, Mpo.de Acatlan (Reddell and Elliott, 1974:9)*Cueva de las Maravillas, 6 km SSW Acatlan, Mpo. deAcatJan (Reddell, 1973f:95)+Cueva del Nacimiento, above Vista Hermosa, Acatlan,Mpo. de Acatlan (Villa R., 1967:224)*Cueva del Nacimiento del Rio San Antonio, 10 kmSSW Acatlan, Mpo. de Acatlan (Reddell, 1973e:89; Reddell, 1973f:95; Reddell and Elliott, 1974:9)Cueva Sin Fin, 2 km SW Acatlan, Mpo. de AcatlanSotano Sin Hondo, 5 km SSW Acatlan, Mpo. deAcatlanHuautla de Jimenez Region*Cave near Puente de Fierro, 1.5 km N Huautla deJimenez, Mpo. de Huautla de JimenezSink near Huautla de Jimenez, Mpo. de Huautla deJimenezCave No.3, near Huautla de Jimenez, Mpo. deHuautla de JimenezPit No.2, near Huautla de Jimenez, Mpo. de Huautlade Jimenez*Sotano de Agua de Carrizo, 5 km ESE Huautla deJimenez, Mpo. de Huautla de Jimenez (Stone,1979a:33-52,2 maps)Cueva del Arroyo Mazateca, San Miguel Dolina, SanMiguel, 5 km SE Huautla de Jimenez, Mpo. deHuautla de JimenezCueva del Camino, Puente de Fierro, 1.5 km NHuautla de Jimenez, Mpo. de Huautla de Jimenez*Cueva del Escorpion, San Miguel Dolina, San Miguel,5 km SE Huautla de Jimenez, Mpo. de Huautla deJimenezThe Fissure-see La Grieta*La Grieta, 2 km E San Andres, 5 km E Huautla deJimenez, Mpo. de Huautla de Jimenez (Stone,1977:7-17, map; Steele, 1978:26-29, map)*Milliped Cave, 5 km SE Huautla de Jimenez, Mpo. deHuautla de Jimenez (Fish and Russell, 1966:63,66m)*Milliped Cave, 1.5 km N Huautla de Jimenez, Mpo.de Huautla de Jimenez (Russell, 1965 :62)*Cueva del Molino de Carne, 5 km ESE Huautla deJimenez, Mpo. de Huautla de Jimenez*Cueva de los Pajaros, 5 km SE Huautla de Jimenez,Mpo. de Huautla de JimenezCueva Arriba del Presidente, 1 km N Huautla deJimenez, Mpo. de Huautla de Jimenez*Cueva Bonita del Presidente, 1 km N Huautla deJimenez, Mpo. de Huautla de Jimenez299

*Cueva del Puente de Fierro, 1.5 km N Huautla deJimenez, Mpo. de Huautla de Jimenez (Russell,1965:62)Cueva de Puente Fierron-see Cueva del Puente deFierro*Gruta de Regadura, 2 km N Huautla de Jimenez,Mpo. de ?Huautla de Jimenez (Russell, 1965:62)Cueva Arriba del Rio Iglesia-see Sotano del RioIglesia*Sotano del Rio Iglesia, 5 km SE Huautla de Jimenez,Mpo. de Huautla de Jimenez (Boon, 1969:32,35m, 39)*Cueva de San Agustin, San Agustin, 5 km SE Huautlade Jimenez, Mpo. de Huautla de Jimenez (Boon,1969:33m, 39-40; Finn, 1971:46, 47m, 49)*Sotano de San Agustin, San Agustin, 5 km SEHuautla de Jimenez, Mpo. de Huautla de Jimenez(Fish, 1970:3-7, map; Steele, 1978:map; Stone,1979b:33-58, 3 maps)Water·Trough Cave, 5 km SE Huautla de Jimenez,Mpo. de Huautla de Jimenez (Fish and Russell,1966:62, 66m)Isthmo de Tehuantepec Region+Cave 2 km S Tollosa, 100 m (Baker and Greer,1960:414,415)+Cave near Montebello, 24 km N Matias Romero,Mpo. de San Juan Guichicovi (Villa R., 1967:157)+Cave 29 km N Matias Romero, Mpo. de ?San JuanGuichicovi (Schaldach, 1965:133)+Cave 38.4 km N Matias Romero, Mpo. de MatiasRomero (Schaldach, 1965:131)+Cave at Santo Domingo Tehuantepec, Mpo. de SantoDomingo Tehuantepec (de la Torre, 1955:696)+Las Cuevas, "bat caves" in the bed of the Rio Tehuantepec,16 k"m NW Santo Domingo Tehuantepee,Mpo. de Jalapa del Marques (Goodwin, 1969:260)+Cueva del Convento, 6 'km NE Magdalena Tequisistlan,Mpo. de Jalapa del Marques (Mones, 1971:169)Cueva Diana Liesa-see Cueva Lisa+Cueva Lisa, 1 km W Santo Domingo Tehuantepec,50 m, Mpo. de Santo Domingo Tehuantepec(Villa R., 1967:153)+Cueva Pma del Colorado, 1 km S Magdalena Tequisistlan,Mpo. de Magdalena Tequisistlan (Villa R.,1967:153)+Cuevas de Santo Domingo, in the center of Istmo deTehuantepec (Sumichrast, 1882:203)Mitla Region+Cave 8 km NW Mitla, Mpo. de ?San Luis Quiavini(Ingles, 1959:384)Oaxaca Region+*Well, Etla, Oaxaca (Argano, 1977:117)San Gabriel Mixtepec Region+Cave at Km 183, 36.5 km N San Gabriel Mixtepec,1700 m, Mpo. de Juquila (Schaldach, 1966:292)San Juan Bautista Cuicatlan Region+Fissure at San Juan Bautista Cuicatlan, Mpo. de SanJuan Bautista Cuicatlan (Villa R., 1967:423)+Cave 4.5 km N San Juan Bautista CuicatIan, Mpo. deSan Juan Bautista Cuicatlan (Villa R., 1967:205)San Sebastian de las Gruw RegionSotano de los Arboles, 1 km S San Vicente Lachixio,Mpo. de San Vicente Lachixio (Reddell, 1973e:91)Cueva de Llano Grande, 1 km S San Vicente Lachixio,2010 m, Mpo. de San Vicente Lachixio (Reddell,1973e:90)Sotano de los Nmos, near Grutas de San Sebastian,3 km N San Sebastian de las Grutas, Mpo. de SantaMaria Sola*Grutas de San Sebastian, 3 km N San Sebastian de lasGrutas, 1820 m, Mpo. de Santa Marfa Sola (Russell,1972d:70-71; Reddell, 1973e:90-91Santiago Apoala Region*Cueva de Apoala, Santiago Apoala, 2000 m, Mpo. deSantiago Apoala (Harden, 1971: 36; Reddell,1973e:91)Sotano de las Bellotas, 5 km NW Santiago Apoala,2240 m, Mpo. de Santiago Apoala*Comedor del Diablo, 5 km NW Santiago Apoala,2240 m, Mpo. de Santiago Apoala (Harden, 1971:36; Reddell, 1973e:91)Cueva de la Laguna-see Cueva de ApoalaSotano de la Milpa Agua, near Santiago Apoala, Mpo.de Santiago ApoalaCueva de Santa Catarina, 12 km NW Apoala, Mpo. deSanta Catarina Ocotlan (Reddell, 1973e:91)Sotano de Un Grillo, 5 km NW Apoala, 2240 m,Mpo. de Santiago ApoalaValle Nacional RegionCueva del Brujo, 12 km N Valle Nacional, Mpo. deValle Nacional(Reddell, 1973e:90)*Cueva del Guano, 8 km N Valle Nacional, Mpo. deValle Nacional (Reddell, 1973e:89-90)*Cueva del Guayabo, 12 km N Valle Nacional, Mpo.de Valle Nacional (Reddell, 1973e:90)Cueva de Lorna del Carmen, 15 km N Valle Nacional,Mpo. de Valle Nacional (Reddell, 1973e:89)*Grutas de Monteflor, Monteflor, 6 km N Valle Nacional,Mpo. de Valle Nacional (Reddell, 1973e:90)Unplaced LocalitiesPUEBLA+Well, Lambityeco (Ruffo and Vigna Taglianti, 1977:166)+Cueva de Dominguillo, Dominguillo (Villa R. andJimenez G., 1961:504)Cuetzalan Region*Grutas de Ateno, 2 km W Xochitlan, Mpo. deXochitIan (Reddell, 1974:185-186, pI. 5m)*Grutas de Atepolihuit, 2 km W Cuetzalan, Mpo. deCuetzalan (Davis, 1974:179, 180m)*Cueva de la Barranca, 7 km SW Cuetzalan, Mpo. deCuetzalan (Reddell, 1974:188)Sima de los Bueyes, 2 km S Cuetzalan, Mpo. deCuetzalan (Sprouse, 1979:61, 63m)Cueva de los Camarones, 2.5 km NW Xochitlan,Mpo. de XochitIan (Reddell, 1974:186)Sima de Cohuatichan-see Sima de los Bueyes300

*Sumidero de Cohuatichan, 2 km S Cuetzalan, Mpo.de Cuetzalan (Davis, 1974:182-183, pI. 4m)Sumidero de Cuetzeitemanes, 1 km W XochitIan,Mpo. de Xochitlan (Reddell, 1974:186)Sima Esteban, 7 km SW Cuetzalan, Mpo. de Cuetzalan(Reddell,1974:187, pI. 6m)*Grutas de Jonotla, 1.5 km SSW Jonotla, Mpo. deJonotla(Davis,1974:171)Cueva de la Milpa, 7 km SW Cuetzalan, Mpo. deCuetzalan (Reddell, 1974:185)Sima Octimaxal Norte-see Sima Octimaxal Sur n. 2Sima Octimaxal Sur n. 1, 3 km SSW Cuetzalan, Mpo.de Cuetzalan (Davis, 1974:177, 178m, 179)Sima Octimaxal Sur n. 2, 3 km SSW Cuetzalan, Mpo.de Cuetzalan (Anonymous, 1973:12-13, 15m;Davis, 1974:179)Grutas de Olivares, 7 km SW Cuetzalan, Mpo. deCuetzalan (Reddell, 1974:187, 189m)*Cueva de Tasalolpan, 2 km W Cuetzalan, Mpo. deCuetzalan (Sprouse, 1979:62, map)*Grutas de Tenextepec, 6 km SW Cuetzalan, Mpo.de Cuetzalan (Davis, 1974:166·167, 168m)Cueva de la Vibora, 1 km NW Xochitlan, Mpo. deXochitlan (Reddell, 1974:186)*Cueva Xochitl, 1 km SE Xochitlan, Mpo. de Xochit·Ian (Davis, 1974:162-163)Sima Chica de Xochitlan, Xochitlan, Mpo. de XochitIanCueva de Xocoyolo, 5.5 km SW Cuetzalan, Mpo. deCuetzalan (Davis, 1974:174-175, pI. 2m)Cueva Murcielago de Xocoyolo, 5.5 km SW Cuetza·Ian, Mpo. de Cuetzalan (Davis, 1974:175)Sumidero de Yohuatichan-see Sumidero de Cohua·tichanHueytemalco Region+Cave 10 km W Hueytemalco, Mpo. de Hueyapan(Ramirez-Pulido and Sanchez-Hernandez, 1971:481)Izucar de Matamoros Region+Cave near San Juan Raboso, Mpo. de Izucar de Matamoros(Gonzalez Ochoa, 1964:983)+Cave at spring 3 km E San Juan Raboso, Mpo. de Izucarde Matamoros (Smith and Van Gelder, 1955:147)Cuevas de Matamoros-see Cuevas del Rio Nexapa+Cuevas del Rio Nexapa, on left side of Rio Nexapanear Izucar de Matamoros, Mpo. de Izucar de Matamoros(Caballero y C., 1943:426)+Cueva del Saito de Alcececa, 9 km S Atzalan (=?Abala),Mpo. de ?Chietla (Villa R., 1967:367)Mesa de San Diego Region+Talus cave on Mesa de San Diego, Mpo. de VenustianoCarranza (Warner and Beer, 1957:17)Tehuacan Region+Caves in the El Riego cliffs south of Tehuacan, Mpo.de Tehuacan (Flannery, 1967:155)+*Well, Tehuacan, Mpo. de Tehuacan (Ruffo and VignaTaglianti,1977:166)Tlapacoya Region+Cave near Tlapacoyan (=Tlapacoya), Mpo. de Tlapacoya(Bhatnagar, 1978:865)Xicotepec Region+Cueva del Azufre, near Xicotepec, Mpo. de Xicotepec(Villa R., 1967:267)Cueva Grillo de la Mona, 3 km W Xicotepec, Mpo.de Xicotepec (Reddell, 1973f:92)Cueva Vampiros de la Mona, 3 km W Xicotepec,Mpo. de Xicotepec (Reddell, 1973f:92)+Cueva de Patla, Planta Hidroelectrica de Tepexi,8 km SW Xicotepec, Mpo. de Xicotepec (Rioja,1954:287)San Pablo Zoquitlan RegionSotano of log-filled sink, 10 km E San Pablo ZoquitIan, Mpo. de San Pablo ZoquitlanCueva del Caballo n. 1, 10 km E San Pablo ZoquitIan, Mpo. de San Pablo ZoquitlanCueva del Caballo n. 2, 10 km E San Pablo Zoquitlan,Mpo. de San Pablo ZoquitlanCueva del Caballo n. 3, 10 km E San Pablo ZoquitIan, Mpo. de San Pablo Zoquitlan*Sotano de Coyomeapan, 12 km ESE San Pablo Zoquitlan,Mpo. de San Pablo Zoquitlan (Atkinson,1978:47-50, map)*Horizontal Cave, 12 km ESE San Pablo Zoquitlan,Mpo. de San Pablo ZoquitlanOverflow Cave-see Cueva de XocotlatPreston's Overflow Cave-see Cueva de XocotlatCueva del Rio Texocotla-see Sumidero de XocotlatSecond River Cave-see Sotano de Coyomeapan*Cueva del Terrible, 10 km E San Pablo Zoquitlan,Mpo. de San Pablo ZoquitlanCueva de Xocotlat, 2 km SSE San Pablo Zoquitlan,Mpo. de San Pablo Zoquitlan (Atkinson and For·sythe, 1979:76-81, map)*Sumidero de Xocotlat, 10 km E San Pablo Zoquitlan,Mpo. de San Pablo Zoquitlan (Russell, 1977:20)QUERETAROAhuacatIan RegionPit 0.5 km S Agua Fda, 22 km NW Jalpan, Mpo. deJalpanPit in pass, La Florida, 14 km NNW Ahuacatlan,Mpo. de Pinal de AmolesSotano de Aguacatilla, 14 km NNW Ahuacatlan,Mpo. de Pinal de AmolesSotanito de Ahuacatlan, 2 km NNW Ahuacatlan,7 km W ] alpan, Mpo. de Pinal de Amoles (Raines,1972a:17-20, map)*Cueva de Emilia, 14 km NNW Ahuacatlan, Mpo. dePinal de AmolesCueva de la Milpa, Agua Fda, 22 km NW Jalpan,Mpo. de JalpanSotano del Potrero-see Xilitla Region, SAN LUISPOTOSICueva de Puente de Dios-see Cueva de Puente deDios del Rio J alpanCueva de Puente de Dios del Rio JaJpan, 4 km SPuerto Animas, 7 km SW Jalpan, Mpo. de .Talpan(Fish and Reddell, 1967:85)Cueva del Puente Natural, 4 km S Puerto Animas,7 km SW ]alpan, Mpo. de Jalpan (Reddell, 1979:100)301

Cueva de los Riscos, 2 km S Rancho Huichale, 8 kmSW Jalpan, Mpo. de J alpan (Fish and Reddell,1967:85-86,87m)Laguna Colorada RegionSinkhole, Laguna Colorada, 23 km W Xilitla, Mpo. deLanda de MatamorosCueva de Camposantos-see Sotano de CamposantosSotano de Camposantos, 5 km W El Lobo, 22 km WXilitla, Mpo. de Landa de Matamoros (Fish andReddell,1967:86)+*Cueva del Madrono, 1.5 km S El Madrono, 19 km WXilitla, 1810 m, Mpo. de Landa de Matamoros(Bonet, 1953a:92, 94-96, 93m; Sbordoni andArgano, 1972: 12-13)"Cueva del Nino,S km W El Lobo, 22 km W Xilitla,Mpo. de Landa de Matamoros (Sbordoni and Argano,1972:13)*Cueva de las Tablas, 5 km W El Lobo, 22 km WXilitla, Mpo. de Landa de MatamorosPinal de Amoles Region*Iron (7) mine, 2 km E Pinal de Amoles, Mpo. de Pinalde Amoles (Peck and Peck, 1973:67)ll5-ft. blind pit, San Gaspar, 10 km NW Pinal deAmoles, Mpo. de Pinal de AmolesCave No.1, 20 km N Pinal de Amoles, Mpo. de Pinalde Amoles (Greer, 1979:109)Cave No. 14, 20 km N Pinal de Amoles, Mpo. dePinal de Amoles (Greer, 1979:112-113, 112m)Cave No. 29-see Cave No. 14Cave No. 33-see Cave No.1*Sotano del Buque, 20 km N Pinal de Amoles, Mpo.de Pinal de Amoles (Jefferys, 1979:70-74, map)Cueva de Chevron, 3 km E Pinal de Amoles, Mpo. dePinal de Amoles (Fish and Reddell, 1967:85)Sotano de Dos Arafias Grandes, 1 km ENE RioBlanco, 2000 m, Mpo. de Pena Miller (Reddell,1979:99)(Pena Miller, F14C47)Cueva Encantada, 3 km W La Cienega, 20 km NNEPinal de Amo)es, Mpo. de Pinal de Amoles (Greer,1979:114,115m)Sotano Encantado-see Cueva Encantada"Sotano de la Escuela, Llano de San Francisco, 10 kmNW Pinal de Amoles, Mpo. de Pinal de Amoles(Bittinger, 1973:86)Sotano del Gobernador, 2 km S Pinal de Amoles,Mpo. de Pinal de Amoles (Reddell, 1979:100)"Cueva del Judio, 2 km S Pinal de Amoles, Mpo. dePinal de Amoles (Reddell, 1979:100)Sotano de La Lagunita, 2 km NE Rio Blanco,2100 m, Mpo. de Pena Miller (Reddell, 1979:99)(Pena Miller, Fl4C47)Cueva del Mercurio, Tejamanil, 3 km SW Pinal deAmoles, Mpo. de Pinal de Amoles (Reddell, 1979:100)"Cueva de los Otates, Los Pinos, 15 km NW Pinal deAmoles, Mpo. de Pinal de Amoles (Bittinger,1973:86)Sotano de la Paila, 20 km N Pinal de Amoles, Mpo.de Pinal de Amoles*Cueva de Penalta, 10 km NW Pinal de Amoles, Mpo.de Pinal de AmolesSotano del Pino, 20 km NNE Pinal de Amoles, Mpo.de Pinal de Amoles*Cueva del Puerto del Leon, 4 km E Rio Blanco,Mpo. de Pinal de Amoles (Reddell, 1979:99)(Pena Miller, Fl4C47)Cueva de Tejamanil, 0.5 km NE Tejamanil, 2.5 kmSW Pinal de Amoles, Mpo. de Pinal de Amoles(Fish and Reddell, 1967:85)*Sotano de Tejamanil, 0.5 km NE Tejamanil, 2.5 kmSW Pinal de Amoles, Mpo. de Pinal de Amoles(Fish and Reddell, 1967:85)Sotano de El Tigre, 24 km by road SW Jalpan, Mpo.de Landa de Matamoros (Reddell, 1979:100)San Joaquin RegionSotano de Herreras-see Grutas de San JoaquinCueva del Rincon, La Lagunita, 11 km SSW SanJoaquin, Mpo. de San Joaquin (San Joaquin,Fl4C58)Sotano del Rincon, La Lagunita, 11 km SSW SanJoaquin, Mpo. de San Joaquin (Mothes, 1978:63)(SanJoaquin,F14C58)Grutas de San Joaquin, 2 km SSW San Joaquin,2440 m, Mpo. de San Joaquin (Mothes, 1978:64-65) (San Joaquin, Fl4(58)San Juan RegionCave of the Bedding, 2 km N San Juan, Mpo. deArroyo Seco*Hoya de las Conchas, 3 km N San Juan, Mpo. deArroyo Seco (Stone and Jameson, 1977:28m, 2933, map)Sotano de Conchas-see Hoya de las ConchasCueva de Guayavitos, 1 km SW San Juan, Mpo. deArroyo Seco (Stone and Jameson, 1977:42)Cueva de la Milpa, 2 km W San Juan, Mpo. de ArroyoSeco*Sotano de Nogal, 5 km NW San Juan, Mpo. deArroyo Seco (Stone and Jameson, 1977:34m,35-39, map)Sotano de Pedregal, 1 km NE San Juan, Mpo. deArroyo Seco (Stone and Jameson, 1977:44m)El Sotanito, 1.5 km N San Juan, Ylpo. de ArroyoSeco (Stone and Jameson, 1977:20m, 21, map)Tancoyol RegionCueva del Agua del Rancho Ojo de Agua, 20 km NEJalpan, Mpo. de J alpanSotano de Lagunita Seca, Tancoyol, 20 km NE Jalpan,Mpo. de Jalpan*EI Socavon, Ojo de Agua, 20 km NE Jalpan, Mpo.de J alpan (Broussard, 1975:2m)Xilitla Plateau RegionCueva de Campamiento-see Sumidero del LlanoConejoCueva de los Grillos, 3 km N Cerro de la Luz, 13 kmW Xilitla, Mpo. de Llanda de MatamorosCueva de Llano de los Chiquitos-see Sumidero delLlano Chiquito*Sumidero del Llano Chiquito, 12 km W Xilitla, Mpo.de Landa de Matamoros (Fish, 1978b:41, 42m;Fish,1979:7)Cueva del Llano del Conejo-see Sumidero del LlanoConejo*Sumidero del Llano Conejo, 10 km ENE Xilitla,Mpo. de Landa de Matamoros (Fish, 1979:12,13m)302

Matehuala RegionSumidero 552, at Km. 552, about 45 km SSE Matehuala,1330 m, Mpo. de ?Villa de Guadalupe (?EIMilagro de Guadalupe, F14A45)Sumidero de Matehuala, 3 km E Matehuala, 1500 m,Mpo. de Matehuala (Fish and Reddell, 1967:82)(Matehuala, Fl4A25)Micos Region+Cave 3 km W Micos, Mpo. de Ciudad Valles (Wilsonand Findley, 1971:420, 421) (Damian Carmona,Fl4A89)*Cueva del Huisache, 7 km NW Micos, 26 km NW CiudadValles, Mpo. de Ciudad Valles (Damian Carmona,F14A89)Cueva de la Libertad, 3.5 km WNW Micos, 23 km NWCiudad Valles, 250 m, Mpo. de Ciudad Valles(Damian Carmona, Fl4A89)*Cueva del Lienzo, 9 km S Micos, 17 km WNW CiudadValles, 236 m, Mpo. de Ciudad Valles (Mitchellet aI., 1977:59) (Damian Carmona, F14A89)Sotano del Lienzo-see Cueva del LienzoCueva de Llanura, 3.5 km WNW Micos, 23 km NWCiudad Valles, 250 m, Mpo. de Ciudad Valles(Damian Carmona, F14A89)Micos Cave-see Cueva del Rio Subterraneo*Cueva de Otates, 9 km S Micos, 17 km WNW CiudadValles, 239 m, Mpo. de Ciudad Valles (Mitchellet aI., 1977:59) (Damian Carmona, F14A89)Cueva de Puente Morita, 3.5 km WNW Micos, 23 kmNW Ciudad Valles, 250 m, Mpo. de Ciudad Valles(Reddell, 1965d:50) (Damian Carmona, Fl4A89)*Cueva del Rio Subterraneo, 9 km S Micos, 17 kmWNW Ciudad Valles, 230 m, Mpo. de Ciudad Valles(Wilkens and Bums, 1972:264m, 265; Mitchellet aI., 1977:58)(Damian Carmona, F14A89)Piedra Paloma RegionCueva de Piedra Paloma n. 1, 10 km NNE Ciudaddel Maiz, Mpo. de Ciudad del Maiz (Ciudad delMaiz, F14A78)Sumidero de Piedra Paloma, 10 km NNE Ciudad delMaiz, Mpo. de Ciudad del Maiz (Reddell, 1965d:49) (Ciudad del Maiz, F14A78)Puente de Dios RegionCueva de Puente de Dios, 30 km SSW Ciudad Valles,Mpo. de Ciudad VallesRayon RegionCueva del Agua, 5 km NNE Rayon, 10 km SSE Cardenas,Mpo. de Rayon (Fish and Reddell, 1967:83)Salto del Agua Region+Cave under EI Salto, 2 km E Saito del Agua, 500 m,Mpo. de Ciudad del Maiz (Jones and Alvarez,1964:303) (Saito del Agua, Fl4A68)San Nicolas de los Montes RegionCueva de la Abanzaba, 2.5 km S San Nicolas de losMontes, 13 km N Agua Buena, Mpo. de Tamasopo(Alaquines, Fl4A88)Cueva Arriba de Agua Buena, 2 km NW Agua Buena,Mpo. de Tamasopo (Fish and Reddell, 1967:84)*S6tano de Bema Be B., 7 km SSE San Nicolas delos Montes, 13 km N Agua Buena, Mpo. de Tamasopo(Alaquines, Fl4A88)Sotano de Bomba, 6 km SSE San Nicolas de los Mon·tes, 13 km N Agua Buena, Mpo. de Tamasopo(Alaquines, Fl4A88)*Cueva de la Bonita, near San Nicolas de los Montes,13 km N Agua Buena, Mpo. de Tamasopo (Ala.quines, Fl4A88)Sotano del Camino, near San Nicolas de los Montes,13 km N Agua Buena, Mpo. de Tamasopo (Alaquines,Fl4A88)Sotano de Chamal, near San Nicolas de los Montes,13 km N Agua Buena, Mpo. de Tamasopo (Alaquines,Fl4A88)*Cueva de Elias, 5.5 km S San Nicolas de los Montes,13 km N Agua Buena, 910 m, Mpo. de Tamasopo(Alaquines, Fl4A88)Sotano de Elias-see Cueva de EliasCueva de Laguna Grande, 8 km SSW San Nicolas delos Montes, 13 km N Agua Buena, Mpo. de Tamasopo(Alaquines, Fl4A88)*Cueva de las Lagunitas, 3 km S San Nicolas de losMontes, 13 km N Agua Buena, Mpo. de Tamasopo(Alaquines, Fl4A88)Cueva del Nacimiento del Quince, EI Quince, 21 kmNW Micos, 35 km NW Ciudad Valles, Mpo. deTamasopo (Elliott, 1974:21) (Alaquines, Fl4A88)Cueva de Ojita de Agua, 3 km SSE San Nicolas de losMontes, 13 km N Agua Buena, Mpo. de Tamasopo(Alaquines, F14A88)Pozita del Ojo de Agua, near San Nicolas de los Montes,13 km N Agua Buena, Mpo. de Tamasopo(Alaquines, Fl4A88)Cueva de Pizzara, 8 km N Agua Buena, Mpo. deTamasopo (Alaquines, Fl4A88)Sotano de Pozita, 4 km S San Nicolas de los Montes,13 km N Agua Buena, Mpo. de Tamasopo (Alaquines,F14A88)Cueva de la Verranza, near San Nicolas de los Montes,Mpo. de Tamasopo (Alaquines, Fl4A88)Sierra de EI Abra RegionWell near Cueva Chica-see Pozo de EI Pujal+*Boca del Abra, 11 km E Ciudad Valles, Mpo. de ?Ciu·dad Valles (Rioja, 1955b:44) (NOTE: May be iden·tical to Cueva de Valdosa)Cueva del Agua-see Cueva de la Curva*Sotano del Arroyo, 12 km NNE Ciudad Valles,195 m, Mpo. de Ciudad Valles (Fish, 1974:3-4,5m, 10, 13-14; Mitchell et aI., 1977:49-51)Sahano de Attoya-see Sotano del ArroyoCueva de Ceiba, 24 km NE Ciudad Valles, Mpo. deTamuin (Russell, 1972c:136)*Cueva Chica, 16 km SE Ciudad Valles, 49 m, Mpo.de Ciudad Valles (Breder, 1942:7-15, fig. 1m;Mitchell et aI., 1977:37-42)Cueva de EI Choy-see Cueva del Nacimiento delRio Choy*Sotano de Coatimundi, 36 km NNE Ciudad Valles,Mpo. de Ciudad Valles+Cueva de Corinto, 15 km by road NNW Tamexin(=?Tamuin), Mpo. de ?Tamuin (Villa R., 1967:17,300)304

Cueva de El Coy-see Cueva del Nacuniento del RioCoyCueva de los Cuates--see Cueva de las Cuatas Esteand Cueva de las Cuatas Oeste*Cueva de las Cuatas Este, 16 km SE Ciudad Valles,59.5 m, Mpo. de Ciudad Valles (Mitchell et aI.,1977:42)*Cueva de las Cuatas Oeste, 16 km SE Ciudad Valles,61.5 m, Mpo. de Ciudad Valles (Mitchell et aI.,1977:42)Cueva de la Cuesta-see Sotano de la CuestaSotano de la Cuesta, 36 km NNE Ciudad Valles,Mpo. de Ciudad Valles (Russell, 1974:120)*Cueva de la Curva, 8 km E Ciudad Valles, 131.5 m,Mpo. de Ciudad Valles (Mitchell et aI., 1977:43·44)Cueva de Diamante-see TAMAUUPASCueva Escondida-see Sotano EscondidoSotano Escondido, 12 km NE Ciudad Valles, Mpo. deCiudad Valles (Russell, 1972c:138;WaIsh, 1972:32)Sotano de la Estrella, 36 km NNE Ciudad Valles,Mpo. de Ciudad VallesCueva de Fer-4e-Lance-see Sotano de Fer-de-LanceSotano de Fer-4e-Lance, 16 km SE Ciudad Valles,Mpo. de Ciudad VallesSotano de Ferrocarril-see Cueva de la CurvaCueva del Fraiser-see Cueva de CeibaCueva Grande, 11 km SE Ciudad Valles, Mpo. deCiudad Valles (Russell and Raines, 1967:78,81m)Hoya de Higueron, 15 km NE Ciudad Valles, Mpo. deCiudad Valles (Greer, 1977:72, 13m)Joya de Higueron-see Hoya de HigueronJoya de Igrion-see Hoya de Higueron*Ventana Jabali, 20 km NE Ciudad Valles, Mpo. deTamuin (Russell and Raines, 1967:75, 77m)*Sotano de Japones, 19.5 km NNE Ciudad Valles,243 m, Mpo. de Ciudad Valles (Mitchell et aI.,1977:53; Fish, 1978a:I-2, map)Sotano de Jerbaniz-see Sotano de Yerbaniz*Sotano de Jos, 8 km NE Ciudad Valles, 176 m,Mpo. de Ciudad Valles (Mitchell et aI., 1977:45)Cueva del Leon, 9 km NE Ciudad Valles, Mpo. deCiudad Valles (Russell, 1972c:139)Sotano del Loro, 24 km NW Ciudad Valles, Mpo. deCiudad Valles (Morris, 1971:44; Russell, 1972c:139)Sotano de los Loros-see Sotano del Loro*Cueva de El Mante, 16 km SE Ciudad Valles, Mpo.de Ciudad Valles (Russell, 1972c:141)Sotano del Mante-see Cueva de El ManteSotano de Manuel, 16 km SE Ciudad Valles, Mpo. deCiudad Valles (Russell and Raines, 1967:79)*Sotano de Matapalma, 21 km NNE Ciudad Valles,242 m, Mpo. de Ciudad Valles (Walsh, 1972:15,16m, 17; Mitchell et aI., 1977:53-55)Cueva de los Monos-see Sotano de los Monos*Sotano de los Monos, 20 km NW Ciudad Valles,Mpo. de Ciudad Valles (Walsh, 1972:31; Russell,1972c: 140; Greer, 1974:23, 24m)*Sistema de Montecillos, 8 km NE Ciudad Valles,190 m (Sotanito de Montecillos entrance) and157.5 m (Sotano de Pichijumo entrance), Mpo. deCiudad Valles (Russell and Raines, 1967:74-75,76m; Mitchell et aI., 1977:46-47)Sotanito de Montecillos--see Sistema de MontecillosSotano de Montecillos--see Sistema de MontecillosCueva del Nacimiento del Rio Choy, 19 km E CiudadValles, Mpo. de Tamuin (Bonet, 1953b:241, 248,257; Russell, 1972c:140)+Cueva del Nacimiento del Rio Coy, 29 km S CiudadValles, Mpo. de ?P. Antonio Santos (Bonet, 1953b:245, 265m; Russell, 1972c:142)+Cueva de El Nilo, 20 km SSE Ciudad Valles, Mpo. deCiudad Valles (Russell, 1972c:141)*Sotano de Palma Seca, 7 km NE Ciudad Valles,151.5 m, Mpo. de Ciudad Valles (Mitchell et aI.,1977:44)Sotano de Pichijumo-see Sistema de Montecillos*Sotano de las Piedras, 7.5 km NE Ciudad Valles,145 m, Mpo. de Ciudad Valles (Mitchell et aI.,1977:44-45)*Cueva Pinta, 13 km NE Ciudad Valles, Mpo. deCiudad Valles (WaIsh, 1972:31)*Sotano de la Pipa, 35 km NNE Ciudad Valles, Mpo.de Ciudad VallesPujal Cave n. I-see Cueva Chica+Pozo de El Pujal, 17 km SE Ciudad Valles, Mpo. deCiudad Valles (Breder, 1942:8; Russell, 1972c:141)Cueva del Rancho Viejo, 14 km E Ciudad Valles,Mpo. de Tamuin (Mitchell et aI., 1977:72)Cueva de la Ranita, 16 km SE Ciudad Valles, Mpo.de Ciudad Valles*Sotano de la Roca, 13 km NNE ClUdad Valles,240.5 m, Mpo. de Ciudad Valles (Mitchell et aI.,1977:51)*Cueva de Los Sabinos, 13 km NNE Ciudad Valles,239.5 m, Mpo. de Ciudad Valles (Fish, 1974:9-10,15m; Mitchell et aI., 1977:51-52)Cuevacita del Sotanito, 8 km NE Ciudad Valles, Mpo.de Ciudad Valles*Sotano de Soyate, 12.5 km NE Ciudad Valles, 293 m,Mpo. de Ciudad Valles (Elliott, 1970:63-66, 65m;Mitchell et aI., 1977:47)*Cueva de Tanchipa, 35 km NNE Ciudad Valles, Mpo.de Ciudad Valles (Russell, 1972c:136)*Cueva de Taninul n. 1, 13 km SE Ciudad Valles,Mpo. de Tamuin (Bonet, 1953b:263m; Russelland Raines, 1967:78; Sbordoni and Argano, 1972:11)Cueva de Taninul n. 4, 13 km ESE Ciudad Valles,Mpo. de Tamuin (Russell and Raines, 1967:78,80m)Cueva de Tantobal, 21 km SSE Ciudad Valles, Mpo.de ?Tanlajas (Russell, 1972c:141)Sotano de la Tenaja-see Sotano de la Tinaja*Sotano del Tigre, 14.5 km NNE Ciudad Valles,245.5 m, Mpo. de Ciudad Valles (Mitchell et aI.,1977:52-53)Cueva de la Tinaja-see Sotano de la Tinaja*Sotano de la Tinaja, 10.5 km NE Ciudad Valles,165.5 m, Mpo. de Ciudad Valles (Fish, 1974:4,8m, 9,14,17; Mitchell et aI., 1977:47-49)*Sotano del Toro, 11 km SE Ciudad Valles, 92 m,Mpo. de Ciudad Valles (Mitchell et aI., 1977:42-43)Mina de Ultima Tiron-see Cueva de Valdosa*Cueva de Valdosa, 11 km E Ciudad Valles, Mpo. deCiudad Valles (Russell, 1972c:140-141)305

*S6tano de Yerbaniz, 21.5 km NNE Ciudad Valles,241.5 m, Mpo. de Ciudad Valles (Mitchell et al.,1977:55-56)*Hoya de Zimapan, 22 km NE Ciudad Valles, Mpo. deCiudad Valles (Morris, 1971:45; Russell, 1972c:139-140)Sierra de Alvarez RegionCave near San Francisco, Mpo. de Zaragoza (Elliottand Reddell, 1973:194) (Santa Catarina, F14A85)*Cave in Valle de los Fantasmas, 4 km NW San Francisco,Mpo. de Zaragoza (Elliott and Reddell,1973:194)(Santa Catarina, F14A85)S6tano in Valle de los Fantasmas, 4 km NW SanFrancisco, Mpo. de Zaragoza (Elliott and Reddell,1973:194) (Santa Catarina, F14A85)S6tano de Abernathy, Los S6tanos Unidos, 6 kmWNW San Francisco, Mpo. de Zaragoza (Elliott andReddell, 1973:192) (Santa Catarina, F14A85)S6tano de Araiia-see S6tano de las AraiiasSotano de las Araiias, Los S6tanos Unidos, 6 kmWNW San Francisco, Mpo. de Zaragoza (Elliott andReddell, 1973:192) (Santa Catarina, Fl4A85)Cueva de los Caballos, Los S6tanos Unidos, 6 kmWNW San Francisco, Mpo. de Zaragoza (Elliott andReddell, 1973:192 (Santa Catarina, F14A85)S6tano de Carlos, 2 km N San Francisco, Mpo. deZaragoza (Elliott and Reddell, 1973:192) (SantaCatarina, F14A85)Cueva de Carnicerias, NW San Francisco, Mpo. deZaragoza (Elliott and Reddell, 1973:192) (SantaCatarina, F14A85)Cueva de Cinquenta y Ocho, I km W Cinquenta yOcho and 4 km S San Francisco, Mpo. de Zaragoza(Elliott and Reddell, 1973:192) (Santa Catarina,Fl4A85)Cave of Dead Cow, 4 km S San Francisco, Mpo. deZaragoza (Santa Catarina, F14A85)Cueva de la Entrada Chica, Valle de los Fantasmas,4 km NW San Francisco, Mpo. de Zaragoza (Elliottand Reddell, 1973:192) (Santa Catarina, F14A85)Sumidero de Fantasmas, Valle de los Fantasmas,4 km NW San Francisco, Mpo. de Zaragoza (Elliottand Reddell, 1973:192) (Santa Catarina, F14A85)*SOtano de la Golondrina, Valle de los Fantasmas,4 km NW San Francisco, Mpo. de Zaragoza (Walsh,1972:50m, 70; Elliott and Reddell, 1973:192)(Santa Catarina, F14A85)S6tano de las Golondrinas-see S6tano de la GolondrinaCueva de la Iglesia, 1.6 km W San Francisco, Mpo. deZaragoza (Walsh, 1972:51m, 72; Elliott and Reddell,1973:192)(Santa Catarina, F14A85)Cueva de la Laguna, 6.5 km NNW San Francisco,Mpo. de Zaragoza (Elliott and Reddell, 1973:192193) (Santa Catarina, F14A85)Cueva de las Moscas, 1.5 km S San Francisco, Mpo.de Zaragoza (Elliott and Reddell, 1973:193)(Santa Catarina, F14A85)Cuevacita de Nopales, 1 km SW Cinquenta y Ochoand 5 km S San Francisco, Mpo. de Zaragoza(Santa Catarina, F14A85)Sotano de Nopales, 1 km SW Cinquenta y Ocho and5 km S San Francisco, Mpo. de Zaragoza (Walsh,1972:55m, 68; Elliott and Reddell, 1973:193)(Santa Catarina, F14A85)SOtano de Ojo de Agua, 9 km ESE San Francisco,Mpo. de Zaragoza (Walsh, 1972:54m, 67; Elliottand Reddell, 1973:193) (Santa Catarina, F14A85)Sotano de Pajaro, Los Sotanos Unidos, 6 km WNWSan Francisco, Mpo. de Zaragoza (Elliott andReddell, 1973:193)(Santa Catarina, Fl4A85)*Cueva de la Puente, 17 km SSE San Francisco, Mpo.de Zaragoza (Walsh, 1972:60-61, 63m; Elliott andReddell, 1973:193)(La Salitrera, Fl4C15)*S6tano de Puerto de los Lobos, 1.5 km S San Francisco,2340 m, Mpo. de Zaragoza (Walsh, 1972:56-57, 59m; Elliott and Reddell, 1973:193) (SantaCatarina, Fl4A85)Cueva de las Rusias, Las Rusias, 5 km E San Francisco,Mpo. de Zaragoza (Fish and Reddell, 1967:83) (Santa Catarina, Fl4A85)S6tano de San Francisco, San Francisco, 2200 m,Mpo. de Zaragoza (Elliott and Reddell, 1973:194;Anonymous, 1974c:4m) (Santa Catarina, F14A85)*S6tano de San Francisco n. 2, 0.5 km S San Francisco,2300 m, Mpo. de Zaragoza (Elliott andReddell, 1973:194) (Santa Catarina, Fl4A85)S6tano de Super-Macho, Los Sotanos Unidos, 6 kmWNW San Francisco, Mpo. de Zaragoza (Elliottand Reddell, 1973:194) (Santa Catarina, Fl4A85)Sierra EI Pino RegionCueva de Los Avalos, Ejido de Los Avalos, 14 kmENE Ciudad del Maiz, 1140 m, Mpo. de Ciudad delMaiz (Reddell, 1965d:49) (Ciudad del Maiz,Fl4A78)Cueva Seca de Los Avalos, Ejido de Los Avalos,14 km ENE Ciudad del Maiz, 1180 m, Mpo. deCiudad del Mafz (Reddell, 1965d:49) (Ciudad delMaiz, F14A78)*Cueva de La Lagunita, La Lagunita, 16 km NE Ciudaddel Maiz, 1150 m, Mpo. de Ciudad del Maiz(Reddell, 1965d:49; Reddell, 1973g:96) (Ciudaddel Maiz, Fl4A78)Cueva de Mosca, La Lagunita, 16 km NE Ciudad delMafz, 1150 m, Mpo. de Ciudad del Mafz (Reddell,1979:101) (Ciudad del Maiz, F14A78)Cueva de los Murcielagos, 15 km NE Ciudad delMaiz, Mpo. de Ciudad del Maiz (Ciudad del Maiz,F14A78)Cueva de los Muros, 15 km NE Ciudad del Maiz, Mpo.de Ciudad del Maiz (Ciudad del Mafz, Fl4A78)Xilitla RegionS6tano near Tlamaya, 2.5 km NNW Xilitla, Mpo. deXilitla+Cave 4 km W on road to Xilitla, Mpo. de Xilitla(Pine, 1972:41)+Cave near road between Pan American highway andXilitla, Mpo. de Xilitla (Zweifel, 1956:16-17)(NOTE: This is probably Cueva de El Jobo)+Large cave 9 km NNE Xilitla, Mpo. de Xilitla (Dalquest,195'3a:25, 30)+Cueva del Ahuate n. 1, Puerto del Ahuate, 2 km SWXilitla, 721 m, Mpo. de Xilitla (Bonet, 1953a:57,pl.5m)*Cueva del Ahuate n. 2, Puerto del Ahuate, 2 km SWXilitla, 721 m, Mpo. de Xilitla (Bonet, 1953a:5868, pI. 5m)306

+*Cueva del Aire, 1.8 km at S85°W from church ofHuichihuayan, 190 m, Mpo. de Xilitla (Bonet,1953a:38-42,39m)+*Cueva de la Barranca, near La Barranca, 1 km NEAhuacatIan, 969 m, Mpo. de Xilitla (Bonet, 1953a:72-76, pi. 7m)Cuevita del Camino, Xilitlilla, 5 km E Ahuacatlan,Mpo. de XilitlaCueva de Campamiento-see Sumidero del LlanoConejo, QUERETAROCueva de la Cisterna, on trail to La Silleta fromTlamaya, NW Xilitla, 1330 m, Mpo. de XilitlaCueva de Cristian, 4 km E Xilitla, Mpo. de Xilitla+*Cueva de los Cuchos, near Plan de Juarez, 2 km SXilitla, 615 m, Mpo. de Xilitla (Bonet, 1953a:6872,69m)Cueva de la Gorra-see Cueva de la PorraCueva de los Grillos-see QUERETARO+*Cueva de la Hoya, 1 km NE Ahuacatlan, 938 m,Mpo. de Xilitla (Bonet, 1953a:83-86, pi. 9m)Sotano de las Hoyas, 1 km NW Ahuacatlan, Mpo. deXilitla (Walsh, 1972:109)Cueva de Huichihuayan-see Cueva del Nacimientodel Rio Huichihuayan*Sotano de Huitzmolotitla, 1 km ESE Tlamaya, 2 kmNNW Xilitla, Mpo_ de Xilitla (Russell and Raines,1967:90)Cueva de la Iglesia, AhuacatIan, Mpo. de Xilitla (Fishand Reddell, 1967:84)*Cueva de EI Jobo, 5 km E XilitIa, 445 m, Mpo. deXilitIa (Bonet, 1953a:44-50, 46m)*Cueva de la Laja, 0.75 km NE AhuacatIan, 870 m,Mpo. de Xilitla (Bonet, 1953a:76-82, 17m)Cueva de Llano de los Chiquitos-see Sumidero delLlano Chiquito, QUERETAROSumidero del Llano Chiquito-see QUERETAROCueva del Llano del Conejo-see Sumidero del LlanoConejo, QUERETAROSumidero del Llano Conejo-see QUERETAROCueva del Lobo-see Cueva de EIJobo+Cueva de la Mujer del Agua, 1.8 km at S85° W fromchurch of Huichihuayan, 245 m, Mpo. de XilitIa(Bonet, 1953a:42-44, 43m)Cueva del Nacimiento del Rio Huichihuayan, 2 kmSW Huichihuayan, lIO m, Mpo. de Xilitla (Bonet,1953a:36-38)*S6tano de la Navidad, SW of La Silleta, NW Xilitla,1970 m, Mpo. de XilitlaCueva de la Parra-see Cueva de la PorraCueva de Poca Ventana-see Cueva del Ahuate n. 2*Cueva de la Porra, 1 km SW Tlamaya, 2 km NNWXilitIa, Mpo. de Xilitla (Russell and Raines, 1967:94)Sotano de la Porra, 1 km SW Tlamaya, 2 km NNWXilitla, Mpo. de Xilitla (Bridgemon, 1974:111,143m)*Cueva de Potrerillos, Rancho de Potrerillos, 2 kmWSW Ahuacatlan, 1250 m, Mpo. de Xilitla (Bonet,1953a:86-91, pi. 10m)SOtano de Potrerillos, Rancho de Potrerillos, 2 km WAhuacatlan, Mpo. de Xilitla (Reddell, 1979:100)*SOtano de Potrero, 1 km N Ahuacatlan, Mpo. deXilitla*SOtano del Pozo, 1 km W AhuacatIan, Mpo. de Xilitla(Russell and Raines, 1967:99, 102m)Sotano del Rancho de la Barranca, 5 km NNE Ahuacatlan,Mpo. de Xilitla (Reddell, 1973d:87)*Cueva del Salitre, 0.5 km NNE Xilitla, 460 m, Mpo.de Xilitla (Bonet, 1953a:50, 52-55, 51m; Russelland Raines, 1967:95, 98m)*Cueva de Ia Selva, 2 km SW Xilitla, Mpo. de Xilitla(Russell and Raines, 1967:99, 101m)*Sotano de Tlamaya, Tlamaya, 2.5 km NNW Xilitla,Mpo. de Xilitla (Russell and Raines, 1967:90, 92m,94)Cueva del Xobo-see Cueva de EI JoboUnplaced CavesSINALOA+Cave near San Luis Potosi (Gonzalez-Ochoa, 1963a:67)+Cueva del Platanito (Villa R., 1967:5)+Large cavern system 1.5 km SW Panuco, Mpo. de Concordia(Bateman and Vaughan, 1974:45,52)+Small cave near La Cueva Chinacatera, 23 km W Pericos,Mpo. de Mocorito (Constantine, 1967:18)+Shallow caves 16 km NNW Los Mochis, Mpo. de LosMochis (Jones et a1., 1972:5)+Cave, Santa Lucia, Mpo. de Concordia (Jones et a1.,1972:7)+Cave 2.5 km NW Topolobampo, Mpo. de Los Mochis(.Jones et aI., 1972:5)+Cave 1.5 km N Zaragoza, Mpo. de Los Mochis (Jones etaI., 1972:13)+Cueva de la Chinacatera, 23 km W Pericos, Mpo. dc Mocorito(Constantine, 1967:18)+Cueva de Don Cristino, 20 km E Mazatlan, Mpo. deMazatlan (Villa R., 1967:152)Cueva de Monte Largo-see Cueva de la ChinacateraSONORA+Large cave, Chinobampo, Mpo. dc Navojoa (Burt, 1938:19)+Shallow caves, Chinobampo, Mpo. de Navojoa (Burt,1938:20)+Cave 24 km NW Guaymas, Mpo. de Guaymas (Burt,1938:19)+Cave, Guiricoba, Mpo. de Alamos (Burt, 1938:21)+Cave 1.5 km W Mayterrena (=Mayorena), 15 km Empalme,Mpo. de Empalme (Bradshaw and Hayward,1960:282)+Cave, PHares de Nacozari, Mpo. de Nacozari (Burt,1938:21)+Cave near San Bernardo, Mpo. de Alamos (Burt andHooper, 1941:2)+Cave below Santa Maria Mine, near EI Tigre, Mpo. de?Naocozari (Burt, 1938:21)+Mina La Aduana, 8 km W Alamos, Mpo. de Alamos(Horst, 1972:49-52, 51m)Cueva de Carbo-see Cueva del TigreLas Minas Cucaracha-see Mina La Aduana+Cueva de la Higuera, San Miguel de Horcasitas, Mpo.de San Miguel de Horcasitas (Roth, 1968:28)+Cueva del Tigre, 22 km SSE Carbo, Mpo. de Carbo(Mitchell,1965:568-577)307

TABASCO+Cave 2 km E Teapa, Mpo. de Teapa (Winkelmann,1962:112)+Cave 3.5 km by road NNE Teapa, Mpo. de Teapa (VillaR.,1967:210)+Cave 4.4 km SE Teapa, Mpo. de Teapa (Lay, 1962:374,375)+Well 22 km SE Villahennosa, Mpo. de Villahennosa(Zullini,1977:76)*Cueva del Azufre, 3.5 km S Tapijulapa, 50 m, Mpo. deTatotalpa (Gordon and Rosen, 1962:360-362, 361m;Sbordoni et al., 1974:15-16)Cueva de El Balneario, 3.5 km NE Teapa, 40 m, Mpo. deTeapa*Grutas del Cocomi, 3 km NE Teapa, 40 m, Mpo. deTeapa (Sbordoni et al., 1974:14-15)*Resumidero del Cocona, 3 km NE Teapa, 40 m, Mpo. deTeapa+Cueva de Don Luis, Finca Rosarita, 3.3 km NE, 3 km E,or SE Teapa, Mpo. de Teapa (Wimsatt, 1969:234)+Cueva del Escorpion, 5 km NNE Teapa, Mpo. de Teapa(Villa R., 1967:149)+Cueva de la Frontera, 1 km ESE El Azufre, on boundarybetween Tabasco and Chiapas, 40 m, Mpo. de Teapa(Sbordoni et al., 1977:21, 22m)+"La Gruta," Ejido Ignacio Allende, 20 0NE El Azufre onboundary between Tabasco and Chiapas, Mpo. deTeapa (Villa R., 1967:157.178)+Cueva La Murcielaguera, 5.3 km by road NNE Teapa,Mpo. de Teapa (Handley, 1966:300)+Cueva del Rancho El Azufre, 8 km W Teapa, Mpo. deTeapa (Villa R., 1967:327)TAMAULIPASAltas Cumbres RegionCueva Aire Mal, 15 km SSW Ciudad Victoria, 1350 m,Mpo. de Ciudad Vict9ria (Chabert, 1973:21, 23m,24) (Ciudad Victoria, F14A29)Sotano de Altas Cumbres n. I-see Cueva BonitaSotano de Altas Cumbres n. 2-see Cueva Aire MalCueva Bonita, 15 km SSW Ciudad Victoria, 1350 m,Mpo. de Ciudad Victoria (Chabert, 1973:21, 22m)(Ciudad Victoria, F14A29)+Cueva La Mula, 10 km W Joya Verde, 28 km SWCiudad Victoria, 800 m, Mpo. de Ciudad Victoria(Alvarez, 1963:339) (Ciudad Victoria, Fl4A29)Cerro El Aire RegionSotano de Abasolo, Abasolo, Mpo. de AbasoloCerro Gordo Region+Cave 3 km WNW EI Carrizo, 14 km SW Llera deCanales, Mpo. de Llera de Canales (Alvarez, 1963:405)(Gomez Farias, FI4A49)La Libertad RegionCueva de "El Murcielago"-see Cueva de Los TronconesCueva de la Presa-see Cueva de Los Troncones+Cueva de la Sepultura, 6 km NW Ciudad Victoria,740 m, Mpo. de Ciudad Victoria (Malaga Alba andVilla R., 1957:539, 543, 545, 567 (Guemez,Fl4A19)+Cueva de Los Troncones, near La Libertad, 8 km NWCiudad Victoria, Mpo. de Ciudad Victoria (RodriguezCabo, 1953:361-362) (Guemez, Fl4A19)Purificacion Region+Cave, El Chihue, 31 km NW Ciudad Victoria, 1800 m,Mpo. de Villa Hidalgo (Walker, 1955a:2) (CasasReales, Fl4A18)*Small spring 0.5 km W Comado Castillo, 41 km NWCiudad Victoria, 2100 m, Mpo. de Villa Hidalgo(Casas Reales, F14A18)*Small spring W of Coorado Castillo, 41 km NW CiudadVictoria, 2100 m, Mpo. de Villa Hidalgo(Casas Reales, Fl4A18)Sumidero de los Abelos-see Sumidero de Oyamel*Agua de los Allarines, 0.8 km N Coorado Castillo,42 km NW Ciudad Victoria, 1960 m, Mpo. de VillaHidalgo (Casas Reales, Fl4A18)*Cueva de los Allarines, 0.8 km N Comado Castillo,42 km NW Ciudad Victoria, 1900 m, Mpo. de VillaHidalgo (Treacy, 1979:16) (Casas Reales, F14A18)+Cueva de la Boca, 5 km SW Canon de la Boca, 22 kmNNW Ciudad Victoria, Mpo. de ?Guemez (Villa R.,1967:231) (?Guemez, Fl4A19)*Cueva del Borrego, 0.5 km S Comado Castillo, 41 kmNW Ciudad Victoria, 1980 m, Mpo. de Villa Hidago(Casas Reales, Fl4A18)Cueva del Brinco-see Sistema Purificacion*Sotano de las Calenturas, 0.7 km S Yerbabuena,34 km NW Ciudad Victoria, 1460 m, Mpo. de VillaHidalgo (Pate, 1979:86-87) (Casas Reales, Fl4A18)*Cueva de la California, 2 km N Rancho Nuevo,2560 m, Mpo. de Villa Hidalgo (Casas Reales,F14A18)*Cueva del Camino, 1 km W Rancho Nuevo, 34 kmWNW Ciudad Victoria, 2500 m, Mpo. de VillaHidalgo (Casas Reales, F14A18)*Cueva de Coral, near summit of Cerro Zapatero,1 km N Comado Castillo, 41 km NW Ciudad Victoria,2100 m, Mpo. de Villa Hidalgo (Pate, 1979:86)(Casas Reales, F14A18)Cueva de Chuparosa, Rancho Revilla, Mpo. de VillaHidalgo (Casas Reales, Fl4A18)*Cueva de Desmontes, Comado Castillo, 41 km NWCiudad Victoria, 1920 m, Mpo. de Villa Hidalgo(Casas Reales, Fl4A18)+Cueva La Esperanza, 6 km SW Rancho Santa Rosa,32 km NW Ciudad Victoria, 360 m, Mpo. de Guemez(Alvarez, 1963:405) (?Casas Reales, F14A18)*Grieta de las Flores, Conrado Castillo, 41 km NWCiudad Victoria, 2000 m, Mpo. de Villa Hidalgo(Casas Reales, Fl4A18)Cueva de Infiemillo-see Sistema Purificacion*Sotano de Jestis, 1 km S Rancho Revilla, 38 km NWCiudad Victoria, 2210 m, Mpo. de Villa Hidalgo(Casas Reales, Fl4A18)Pozo de Juan Fuentes Perez-see Grieta de las FloresCueva de Musgo, 0.75 km E Rancho Revilla, 38 kmNW Ciudad Victoria, 2100 m, Mpo. de Villa Hidalgo(Casas Reales, F14A18)Cueva del Oso-see Sistema PurificacionCueva de Oyamel-see Sumidero de Oyamel*Sumidero de Oyamel, 1 km N Comado Castillo41 km NW Ciudad Victoria, 1950 m, Mpo. de villiHidalgo (Treacy, 1979:16) (Casas Reales, F14AI8)308

*Sistema Purificacion, Conrado Castillo, 41 km NWCiudad Victoria, 1980 m (Cueva del 080 entrance).1900 m (Cueva del Brinco entrance), 1I00 m(Cueva de Infiemillo entrance), Mpo. de VillaHidalgo (Pate, 1979:82-101, map; Treacy, 1979:8-31,2 maps) (Casas Reales, F14A18)*Sotano de Rancho Nuevo n. 1, Rancho Nuevo,34 km WNW Ciudad Victoria, 2600 m, Mpo. deVilla Hidalgo (Casas Reales, Fl4A18)*Sotano de Rancho Nuevo n. 2, Rancho Nuevo,34 km WNW Ciudad Victoria, 2600 m, Mpo. deVilla Hidalgo (Casas Reales, Fl4A18)*Sotano de Rancho Nuevo n. 3, Rancho Nuevo,34 km WNW Ciudad Victoria, 2600 m, Mpo. deVilla Hidalgo (Casas Reales, F14A18)*Sotano de Rancho Nuevo n. 4, Rancho Nuevo,34 km WNW Ciudad Victoria, 2600 m, Mpo. deVilla Hidalgo (Casas Reales, Fl4A18)Cueva de Rancho Revilla-see Cueva de Revilla*Pozo de Rancho Viejo, 2 km SW Rancho Nuevo,34 km .WNW Ciudad Victoria, 2560 m, Mpo. deVilla Hidalgo (Casas Reales, F14A18)*Cueva de Revilla, Rancho Revilla, 40 km NW CiudadVictoria, 2300 m, Mpo. de Villa Hidalgo (CasasReales, F14A18)Cueva del Tecolote-see Cueva de Oesmontes*Cueva de 105 Tres Bobos, near Conrado Castillo,41 km NW Ciudad Victoria. Mpo. de Villa Hidalgo(Casas Reales, F14A18)Entrada del Viento Alta, Conrado Castillo, 41 kmNW Ciudad Victoria, 2000 m, Mpo. de Villa Hidalgo(Treacy, 1979:17)(Casas Reales, Fl4A18)*Entrada del Viento Baja, Conrado Castillo, 41 kmNW Ciudad Victoria, 2000 m, Mpo. de Villa Hidalgo(Pate, 1979:86; Treacy, 1979:17) (Casas Reales,Fl4A18)Cueva Vrincon-see Sistema Purificacion*Cueva X, Conrado Castillo, 41 km NW Ciudad Victoria,1950 m, Mpo. de Villa Hidalgo (Treacy,1979:16-17) (Casas Reales, F14A18)Santander Jimenez Region+Cueva del Guano, 1I km Nand 21 km W Santanderjimenez, 380 m, Mpo. de Santander jimenez(Aguirre Pequeno, 1959:245-256; Alvarez, 1963:401)Sierra de EI Abra RegionCueva del Abra, 1 km S EI Abra, 9 km NE AntiguoMorelos, 300 m, Mpo. de Antiguo Morelos (Russelland Raines, 1967:65, 67m)(Quintero, Fl4A69)Cueva Chica del Arroyo Seco, 15 km ESE AntiguoMorelos, Mpo. de Antiguo Morelos (Russell,1972c:135)Cueva Grande del Arroyo Seco, 15 km ESE AntiguoMorelos, Mpo. de Antiguo Morelos (Russell,1972c:135)Sotano del Arroyo Seco-see Cueva Grande delArroyo SecoCueva de El Choveno, EI Choveno, 9 km NNE AntiguoMorelos,Mpo. de Antiguo Morelos (Quintero,F14A69)*Cueva de Oiamante, 23 km SE Antiguo Morelos,Mpo. de Antiguo Morelos (Minton, 1978:7-15,map)*Cueva de la Florida, 1 km NNW Praxedis Guerrero(=EI Pachon), 8 km NE Antiguo Morelos, 260 m,Mpo. de Antiguo Morelos (Mitchell, 1970:64m;Russell, 1972c:134-135) (Quintero, Fl4A69)Cueva de EI Mante n. I-see Cueva del Nacimientodel Rio Mante+Cueva del Nacimiento del Rio Mante, 10 km SWCiudad Mante, 100 m, Mpo. de Ciudad Mante(Russell and Raines, 1967:64) (Quintero, F14A69)*Cueva de EI Pachon, Praxedis Guerrero (=EI Pachon),7 km NE Antiguo Morelos, 210.5 m, Mpo. de AntiguoMorelos (Russell and Raines, 1967:65, 68m;Mitchell et aI., 1977:57-58) (Quintero, Fl4A69)Cueva de 105 Pajaros, 1 km NNW Praxedis Guerrero(=EI Pachon), 8 km NE Antiguo Morelos, 260 m,Mpo. de Antiguo Morelos (Russell, 1972c:135)(Quintero, F14A69)*Grutas de Quintero, 1.5 km S Quintero, 13 km SWCiudad Mante, 200 m, Mpo. de Ciudad Mante(Russell and Raines, 1967:64, 66m; Russell,1972c:134) (Quintero, F14A69)Cueva de San Rafael de los Castro8-see Sotano deSan Rafael de los Castros*Sotano de San Rafael de los Castros, San Rafael delos Castros, 13 km ENE Ciudad Mante, 100 m,Mpo. de Ciudad Mante (Russell, 1972c:133)(Lorna Alta, Fl4A59)*Cueva de Santa Elena, 14 km SE Antiguo Morelos,Mpo. de Antiguo Morelos (Russell, 1972c:135136)(Sierra La Colmena, Fl4A79)Sotano de Santa Elena-see Cueva de Santa Elena*Sotano de EI Venadito, 16 km SE Antiguo Morelos,312 m, Mpo. de Antiguo Morelos(Walsh, 1972:12,14m; Mitchell et aI., 1977:56-57 (Sierra La Colmena,Fl4A79)Sierra de Guatemala Region+Cave at Agua Linda, 1I km WNW Gomez Farias,1800 m, Mpo. de Gomez Farias (Martin, 1958:43) (Gomez Farias, F14A49)+Cave 1 km S Aserradero del Paraiso, 13 km NNWChamal, 420 m, Mpo. de Ocampo (Koopman andMartin, 1959:2-3)(Loma Alta, Fl4A59)+Cave, Casa Piedras, 8.5 km WNW Gomez Farias,1500 m, Mpo. de jamauve (Martin, 1958:44)(Gomez Farias, F14A49)+Cave, Ojo de Agua, 3 km SE Gomez Farias, 100 m,Mpo. de Gomez Farias (Alvarez, 1963:398)(Gomez Farias, F14A49)+Caves, Rancho del Cielo, 7 km NNW Gomez Farias,1I60 m, Mpo. de Gomez Farias (Hooper, 1953:9) (Gomez Farias, Fl4A49)+Bell-shaped sink, Rancho del Cielo, 7 km NNWGomez Farias, 1I60 m, Mpo. de Gomez Farias(Martin, 1958:47) (Gomez Farias, F14A49)+Ory sinkhole, Rancho del Cielo, 7 km NNW GomezFarias, 1I60 m, Mpo. de Gomez Farias (Martin,1958:56) (Gomez Farias, Fl4A49)+Deep open sink, Rancho del Cielo, 7 km NNW GomezFarias, 1I60 m, Mpo. de Gomez Farias (Martin,1958:43) (Gomez Farias, Fl4A49)+Sinkhole, 12 m deep (No.1), Rancho del Cielo, 7 kmNNW Gomez Farias, 1I60 m, Mpo. de Gomez Farias(Martin, 1958:47) (Gomez Farias, Fl4A49)309

+Sinkhole, 12 m deep (No.2), Rancho del Cielo,7 km NNW Gomez Farias, 1160 m, Mpo. de GomezFarias (Martin, 1958:47) (Gomez Farias,F14A49)+Cave n. 6, Rancho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (Harrell,1951:118, 119, 123, 182) (Gomez Farias,F14A49)+Cave n. 11, Rancho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (Harrell,1951:135) (Gomez Farias, Fl4A49)+Cave n. 13, Rancho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (Harrell,1951:120, 135, 182, 186) (Gomez Farias,F14A49)+Cave n. 14, Rancho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (Harrell,1951:119) (Gomez Farias, Fl4A49)+Cave n. 17, Rancho del Cie!o, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (Harrell,1951:135) (Gomez Farias, Fl4A49)+Cave n. 19, Rancho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (Harrell,1951:118,119) (Gomez Farias, Fl4A49)+Cave n. 20, Rancho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (Harrell,1951:169, 186) (Gomez Farias, F14A49)35 meter deep pit, Rancho del Cielo, 7 km NNWGomez Farias, 1160 m, Mpo. de Gomez Farias(Gomez Farias, F14A49)*Sinkhole, Raneho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (GomezFarias, F14A49)Small eave near EI Refugio, 17 km N Ocampo,1100 m, Mpo. de Ocampo (?Ocampo, Fl4A58)*Cave near San J ore, 8 km W Gomez Farias, 1300 m,Mpo. de Gomez Farias (Gomez Farias, F14A49)Cueva de las Abejas-see Bee CaveCueva del Agua, 3 km S Manantiales, 15 km SWGomez Farias, Mpo. de ?Gomez Farias (LornaAlta, F14A59)*Bee Cave, 8.5 km NE Adolfo Lopez Mateos (=ChamalNuevo), 19 km NE Ocampo, 248.5 m, Mpo. deOcampo (Mitchell et aI., 1977:60·61) (Lorna Alta,F14A59)Cave C-5, 13 km NW Gomez Farias, Mpo. de GomezFarias (Gomez Farias, F14A49)'''Sotano del Caballo Mora, 7.8 km NW Adolfo LopezMateos (=Chamal Nuevo), 11 km NE Ocampo,Mpo. de Ocampo (Mitchell et aI., 1977:61-62)(Lorna Alta, F14A59)'''Cueva de la Capilla, near EI Porvenir (=La Perra),11 km NW Gomez Farias, 2040 m, Mpo. de Jamauve(Sbordoni and Argano, 1972:8; Elliott,1973b:79, 82m; Mitchell and Kawakatsu, 1973a:673-675, 674m) (Gomez Farias, F14A49)Cueva Capilla de la Perra-see Cueva de la Capilla*Cueva de EI Charco de la Cabeza, 6 km NW EI Encino,12 km N Gomez Farias, Mpo. de Llera deCanales (Elliott, 1979:17-18) (Gomez Farias,F14A49)*Crystal Cave, Rancho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (Elliott,1973b:79, 83m) (Gomez Farias, Fl4A49)Dry Cave, Rancho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de Gomez Farias (GomezFarias, Fl4A49)Cueva del Ejido de Ia Libertad, 10 km N GomezFarias, Mpo. de Llera de Canales (Gomez Farias,F14A49)SOtano del Embudo de EI Mirador, EI Mirador,11 km NE Gomez Farias, Mpo. de Jamauve (GomezFarias, F14A49)Sotanito Escondido, Rancho del Cielo, 7 km NNWGomez Farias, 1160 m, Mpo. de Gomez Farias(Gomez Farias, Fl4A49)"'Sotano Escondido, 1 km SW Gomez Farias, 302.5 m,Mpo. de Gomez Farias (Mitchell et al., 1977:62)(Gomez Farias, F14A49)Cuevita de la Escuela-see Cueva de la EscuelaCueva de la Escuela, J oya de Salas, 15 km NW GomezFarias, 1600 m, Mpo. de J amauve (McKenzie,1965b:27) (Gomez Farias, Fl4A49)*Sotano de Gomez Farias, 3 km ESE Gomez Farias,300 m, Mpo. de Gomez Farias (Walsh, 1972:22)(Gomez Farias, F14A49)Sotano de los Guacamayos, EI Refugio, 17 km NOcampo, 1060 m, Mpo. de Ocampo (Deane, 1977:55, 56m) (Ocampo, F14A58)*Harrison Sinkhole, Rancho del Cielo, 7 km NNWGomez Farias, 1160 m, Mpo. de Gomez Farias(Elliott, 1973b:79, 84m)(Gomez Farias, F14A49)Sotano de Harrison, 1 km N Rancho del Cielo, 8 kmNNW Gomez Farias, 1200 m, Mpo. de GomezFarias (Gomez Farias, F14A69)Indian Springs Cave-see Cueva del Nacimiento delIndio*Cueva del Infiernillo, near San J ore, 8 km W GomezFarias, 1300 m, Mpo. de Jamauve (Russell andRaines, 1967:52; Sustare, 1966:54, 55m) (GomezFarias, F14A49)Cueva de EI Infiemo-see Cueva del Infiernillo*SOtano de J ineo, 1 km NW Gomez Farias 292 mMpo. de Gomez Farias (Mitchell et al., 1977:63:64) (Gomez Farias, Fl4A49)Sumidero de EI Jineo-see Sotano de Jineo*Sotano de la .foya de Salas, Joya de Salas, 15 kmNW Gomez Farias, 1560 m, Mpo. de J amauve(Turner, 1972:3-4, 5m, 7; Walsh, 1972:10,21-22)(Gomez Farias, F14A49)*Sotano del Leon, 2 km S Gomez Farias, Mpo. deGomez Farias (Gomez Farias, Fl4A49)*Cueva de los Leones, J oya de Salas, 15 km NW GomezFarias, 1600 m, Mpo. de Jamauve (Russelland Raines, 1967:51) (Gomez Farias, Fl4A49)*Resumidero de los Mangos, 1.5 km W Gomez FariasMpo. de Gomez Farias (Gomez Farias, F14A49) ,*Cueva de la Mina, 9 km NW Gomez Farias 1527 mMpo. de Gomez Farias (Elliott, 1973b:79, 81m;Mitchell and Kawakatsu, 1973a:671-673, 672m)(Gomez Farias, Fl4A49)*Sotano del Molino, 1 km NW Gomez Farias, 268.5 m,Mpo. de Gomez Farias (Walsh, 1972:22-23; Mitehellet aI., 1977:63) (Gomez Farias, Fl4A49)*Cueva del Nacimiento del Indio, 8 km NW GomezFarias, Mpo. de Gomez Farias (Gomez Farias,F14A49)310

*Cueva del Nacimiento del Rio Frio, 7 km S GomezFarias, 170 m, Mpo. de Gomez Farias (Russell andRaines, 1967:53, 54m) (Lorna Alta, F14A59)+Cueva del Nacimiento del Rio Sabinas, 6 km NW ElEncino, 12 km N Gomez Farias, 180 m, Mpo. deLlera de Canales (Alvarez, 1963:398) (GomezFarias, F14A49)*Sotano del Naral\io, 1.5 km W Gomez Farias, Mpo.de Gomez Farias (Gomez Farias, F14A49)Hoya de Nubas, 10 km NW Gomez Farias, Mpo. deGomez Farias (Gomez Farias, F14A49)*Cueva de Ojo de Agua de Manantiales, 14 km NNEOcampo, 450 m, Mpo. de Ocampo (Elliott, 1979:16-20) (Lorna Alta, F14A59)*Cueva de la Paloma, 1 km NNE Gomez Farias, Mpo.de Gomez Farias (Gomez Farias, F14A49)*Cueva de las Perlas, near El Porvenir (=La Perra),11 km NW Gomez Farias, 2000 m, Mpo. deJamauve (Elliott, 1973b:80, 86m; Mitchell andKawakatsu, 1973a:675-676, 675m) (Gomez Farias,F14A49)Cueva de la Perra-see Cueva de la Capilla*Cueva Chica de la Perra, near El Porvenir (=La Perra),11 km NW Gomez Farias, 2040 m, Mpo. de Jamauve(EUiott, 1973b:80) (Gomez Farias, F14A49)SOtano de las Pinas-see Sotano de los Pinos*Sotano de los Pinos, near J oya de Salas, 15 km NWGomez Farias, Mpo. de J amauve (Russell andRaines, 1967:51) (Gomez Farias, F14A49)Sotano de El Porvenir, 11 km NW Gomez Farias,Mpo. de J amauve (Anonymous, 1974b:112, 125m)(Gomez Farias, F14A49)*Grutas de EI Puente, 11 km SE Ocampo, 400 m,Mpo. de Ocampo (Reddell, 1979:100) (LornaAlta, F14A59)Cueva del Rancho del Cielo n. 3-see SalamanderCave*Cueva del Rancho del Cielo n. 7, Rancho del Cielo,7 km NNW Gomez Farias, 1160 m, Mpo. de GomezFarias (Russell and Raines, 1967:52) (GomezFarias, Fl4A49)Sotano de EI Refugio-see S6tano de los Guacamayos*Cueva del Remolino, near San Jose, 8 km W G6mezFarias, Mpo. de Jamauve (Gomez Farias, F14A49)Cueva del Rio Frio-see Cueva del Nacimiento delRio Frio*Salamander Cave, Rancho del Cielo, 7 km NNW G6mez Farias, 1160 m, Mpo. de Gomez Farias(Russell and Raines, 1967:52) (Gomez Farias,F14A49)*Sotano de Tres Cerritos, 20 km NNW G6mez Farias,Mpo. de Jamauve (G6mez Farias, Fl4A49)S6tano de Tres Manantiales-see Cueva de Ojo deAgua de ManantialesSotano de EI Triunfo, G6mez Farias, Mpo. de GomezFarias (G6mez Farias, Fl4A49)*2,000 Meter Cave, near Joya de Salas, 15 km NWGomez Farias, Mpo. de Jamauve (McKenzie,1965b:25) (Gomez Farias, Fl4A49)*Cueva de los Vampiros, 10 km NE Adolfo LopezMateos (=Chamal Nuevo), 20 km E Ocampo,Mpo. de Ocampo (Lorna Alta, F14A59)*S6tano de Vasquez, 6 km ESE Adolfo Lopez Mateos(=Chamal Nuevo), 20 km E Ocampo, 422 m, Mpo.de Ocampo (Mitchell et aI., 1977:59-60) (LornaAlta, Fl4A59)*Wet Cave, Rancho del Cielo, 7 km NNW GomezFarias, 1160 m, Mpo. de G6mez Farias (Russelland Raines, 1967:52) (Gomez Farias, F14A49)Sierra de Tamaulipas Region+Cave 7 km NW Acuna, Mpo. de ?Casas (Martin et al.,1954:46)+Cave 5 km S, 22 km W Piedra, Mpo. de Soto laMarina (Alvarez, 1963:407)+Cave 5 km S, 26 km W Piedra, Mpo. de Soto laMarina (Alvarez, 1963:405)+Cave 0.5 km from cave 5 km S, 26 km W Piedra,Mpo. de Soto la Marina (Alvarez, 1963:405)*Cueva del Agua de Simon Salinas, 48 km SW Soto laMarina, Mpo. de Soto la Marina*Cueva de los Cuarteles, 10 km SW Aldama, Mpo. deAldama (Garcia Lozano, 1939a:5-7, 6m; Reddell,1973e:87-S8)*Cueva del Virgen de Guadalupe, 48 km SW Soto laMarina, Mpo. de Soto la MarinaTula RegionCueva Abajo de Carreterra, 35 km SW Tula, 1030 m,Mpo. de Tula (Reddell, 1979:101) (Tula, F14A57)Villa Hidalgo RegionTLAXCALACueva del Virgen, EI Chorrito, 18 km W Villa Hidalgo,745 m, Mpo. de Villa Hidalgo (Russell and Raines,1967:28,38) (Villa Hidalgo, Gl4C88)+Cave 13 km NE Tlaxcala, 2360 m (Davis, 1944:377)VERACRUZAtoyac Region+Cave at origin of Rio Atoyac (Mullinex, 1975:37)+Well on outskirts of Paraje Nuevo, 500 m, Mpo. deAmatlan (Argano, 1972a:41)+Cave at edge of rain forest, Potrero Viejo, 5~0 m,Mpo. de Amatloin (Hooper, 1947:43)+Cave on hillside 5 km N Potrero, Mpo. de Amatlin(Hall and Dalquest, 1963:235)+Cave 4 km WNW Hacienda at Potrero Viejo, neartrail to La Caldera, Mpo. de Amatlin (Hobbs,1943:206)+Cave 8 km NW Potrero, Mpo. de Amatlin (Hall andDalquest,1963:235)+Well, San Juan de la Punta, Mpo. de Cuitlahuac orCuichapa (Argano, 1972a:35)+Cave along Rio Blanco, 7 km SE San Juan de laPunta, 120 m, Mpo. de Cuichapa (Hall and Dalquest,1963:214)*Grutas de las Abejas, ?N of Potrero*Grutas de Atoyac, 2 km E Atoyac, 500 m, Mpo. deAtoyac (Villada, 1911:61-67, pI. 15-18)*Cueva de la Charca I, Ejido Charca, N of Potrero,Mpo. de Atoyac*Cueva de Corral de Piedra, 3 km SSE Corral de Piedra,Mpo. de Cuitlahuac or Atoyac*Sotano de las Golondrinas, Manzanilla, 11 km NPotrero, Mpo. de Atoyac311

Sotano de El Maguey, 5 km N Cuitlahuac, Mpo. deCuitlahuac*Cueva del Nacimiento del Rancho Nuevo, 14 km NPotrero, Mpo. de AtoyacCueva del Nacimiento Grande-see Cueva del Ojo deAgua GrandeCueva del Ojo de Agua-see Cave 4 km WNW Haciendaat Potrero Viejo*Cueva del Ojo de Agua Grande, 5 km N PotreroViejo, 550 m, Mpo. de AmatIan (Fish and Reddell,1965:75; Sbordoni and Argano, 1972:18-19)*Sotano de la Palma, 14 km N Potrero, Mpo. deAtoyacSotano de la Pastura, 7 km N Potrero, Mpo. deAtoyacSotano de los Perros, Manzanilla, 10 km N Potrero,Mpo. de Atoyac+Cueva de la Pesca, Potrero, 650 m (Malaga Alba andVilla R., 1957:542)Cueva del Rancho Santa Maria, 6 km N Potrero,Mpo. de AtoyacCueva del Rio Atoyac, 3 km E Atoyac, Mpo. deAtoyac*Cueva de la Sala de Agua, Ejido Colonia, N of Potrero,Mpo. de AtoyacCueva de la Sala de Agua Grande, 5 km N Cuitlahuac,Mpo. de Cuitlahuac (Hall and Dalquest, 1963:234-235)*Cueva de la Sala Seca, 5 km N Cuitlahuac, Mpo. deCuitlahuacLos Tres Cuevas, Cuitlahuac-see Cueva de Sala deAgua GrandeBuena Vista RegionCueva de Camposanto, 2 km W Buena Vista, Mpo. deActopanCueva de Cantil Blanco, 1 km N Buena Vista, Mpo.de ActopanCueva de los Vampiros,. 2 km W Buena Vista, Mpo.de ActopanFortin Region+Cave with sloping entrance 4 km WNW Fortin, Mpo.de Fortin (Hall and Dalquest, 1963:231)+Cavr. with vampires, 4 km WNW Fortin, Mpo. de Fortin(Hall and Dalquest, 1963:231)+Small cave 4 km WNW Fortin, Mpo. de Fortin (Halland Dalquest, 1963:227)+Cave at Metlac, 3 km N Fortin, Mpo. de Fortin(Allen, 1942:97)Jalapa Region+Cave at Cofre de Perote, 2800 m, Mpo. de ?Perote(Long and Jones, 1966:290)+Caves16 km WJalapa (Roth, 1968:25)+Cave 9 km NNW Jalapa, Mpo. de Banderilla (Villa R.,1967:387)+Caves 3 km W Limon, Mpo. de Perote (Hall and Dalquest,1963:317)+Cavem at Plan del Rio, 300 m, Mpo. de Dos Rios(Davis,1944:378)+Cave 5 km ESE Las Vigas, Mpo. de Las Vigas (Davisand Carter, 1962:72)+Caves near Las Vigas, Mpo. de Las Vigas (Ward,1891:743-744)+Cave 5 km E Las Vigas, 2400 m, Mpo. de Las Vigas(Davis, ~944:381)+Caves on Volcancillo, 4 km E Las Vigas, Mpo. deLas Vigas (Hall and Dalquest, 1963:245)Cueva del Infiemillo, 1 km E Paso del Toro, Mpo.de Naolinco (Ward, 1904:634.636)+Cueva de Tilapa, near Orizaba (Sumichrast, 1882:202,203)+Cueva de Tuxpango, near Orizaba, Mpo. de Naranjal(Sumichrast, 1882:201)*Cueva del Volcancillo, Volcancillo, 5 km SE LasVigas, Mpo. de Las Vigas (Reddell and Elliott,1974:12-13, pI. 1m)+Grotte de Xilapa, near Orizaba (Villa R., 1953b:153)+Cueva de Zatiopan, 9 km N Jalap'!, 1700 m, Mpo. deBanderilla (LaVal, 1973: 25)Jesus Carranza Region+Cave 10 m long, 35 km SE JesUs Carranza, Mpo. deJesus Carranza (Hall and Dalquest, 1963:225)+Cave 35 km SE JesUs Carranza, Mpo. de Jesus Carranza(Hall and Dalquest, 1963:224)+Small cave with water 38 km SE JesUs Carranza,Mpo. de JesUs Carranza (Hall and Dalquest, 1963:223)+Shallow recess in cliff 38 km SE JesUs Carranza,Mpo. de JesUs Carranza (Hall and Dalquest, 1963:213)+Long, tubelike cave, 38 km SE JesUs Carranza, Mpo.de Jesus Carranza (Dalquest and Hall, 1949:424)+Small cave formed by fallen rocks 38 km SE JesUsCarranza, Mpo. de Jesus Carranza (Dalquest andHall,1949:424)+Small cave 10 m from above cave, 38 km SE JesUsCarranza, Mpo. de Jesus Carranza (Hall and Dalquest,1963:213)+Large cave 38 km SE JesUs Carranza, Mpo. de JesusCarranza (Dalquest and Hall, 1949:425)+Small caves and crevices, 38 km SF, JesUs Carranza,Mpo. de JesUs Carranza (Hall and Dalquest, 1963:215)+Small roomlike cave 38 km SE JesUs Carranza,Mpo. de JesUs Carranza (Hall and Dalquest, 1963:224)+Long, deep cave, 38 km SE JesUs Carranza, Mpo. deJesUs Carranza (Hall and Dalquest, 1963:224-225)+High narrow cave 38 km SE JesUs Carranza, Mpo. deJesUs Carranza (Hall and Dalquest, 1963:225)Orizaba RegionCueva del Agua de Tlilapan-see Cueva MacingaSotano de la Botella Chica, 5 km NW Tequila, Mpo.de Tequila*Cueva del Carbon, near San Martin, W of Tequila,Mpo. de Tequila*Cueva de la Cascada, 2 km E Tequila, Mpo. de Tequila(Louch and Fish, 1970:23)Cueva de Cerro-see Sotano ltamo+Cueva de Cuautlapa, near Orizaba (Sumichrast, 1882:201)*Cueva del Diablo, 3 km SSW Ciudad Mendoza, Mpo.de Ciudad Mendoza (Reddell, 1973f:95)Sotano de las Espadas, Tequila, Mpo. de TequilaSotano del Hombre Invincible, Soledad Atzompa,Mpo. de Soledad Atzompa312

Sotano de Humo, 5 km NW San Andres, Mpo. deTenejapa*Sotano Itamo, near Soledad Atzompa, Mpo. de SoledadAtzompaSotano Ituro-see Sotano ItamoSotano de Lomapa, near Soledad Atzompa, Mpo. deSoledad Atzompa (Reddell and Elliott, 1974:10)*Cueva Macinga, 2 km E Tlilapan, 1150 m, Mpo. d~Tlilapan (Sbordoni and Argano, 1972:18; Reddell,1973f:93)Cueva de Mazatopa, near Soledad Atzompa, Mpo. deSoledad Atzompa (Reddell and Elliott, 1974:11)Sotano de Milpa, 5 km SW San Andres, Mpo. deTenejapa (Louch and Fish, 1970:19, 21m)*Cueva del Djo de Agua de Tlilapan, Tlilapan, 1200 m,Mpo. de Tlilapan (Sbordoni and Argano, 1972:1718; Reddell, 1973f:93, 94m)Cueva del Djo de Agua de Tlilapan n. I-see Cuevadel Djo de Agua de TlilapanCueva del Djo de Agua de Tlilapan n. 2-see CuevaMacingaCueva de Dpilionida, 2 km N Tequila, Mpo. de TequilaSotano de Dztauatliltaloa, Tequila, Mpo. de Tequila+Cueva de la Presa, 4 km E Drizaba, Mpo. de ?Drizaba(Villa R. and Jimenez G., 1961:504)*Sotano del Profesor, Tequila, Mpo. de Tequila(Raines, 1964: 113-114)Sotano de las Ranas, 5 km SW San Andres, Mpo. deTenejapa (Louch and Fish, 1970:19, 22m)Sotano del Relicario, 3 km N Tequila, Mpo. de TequilaResumidero de San Martin, San Martin, W of Tequila,Mpo. de TequilaCueva del Soldado, Orizaba, Mpo. de Orizaba*Sotano de Sphodrini, Tequila, Mpo. de Tequila*Szontecomostoc,5 km N Tequila, Mpo. de Tequila*Sotano de Teanacan, Soledad Atzompa, Mpo. deSoledad Atzompa (Reddell and Elliott, 1974:10)Cueva de las Trozas, near San Martin, W of Tequila,Mpo. de TequilaCueva de los Vampiros, 3 km SSW Ciudad Mendoza,Mpo. de Ciudad Mendoza (Reddell, 1973f:93)Sotano la Y Griega, Tequila, Mpo. de TequilaPresidio Repon+Deep cave 3 km N Presidio, 500 m, Mpo. de IxhuatIan (Hall and Dalquest, 1963:223)Puente National Region+Series of small caves downstream from Puente Nacional,Mpo. de Puente Nacional (Davis, 1944:376)Tezonapa Region*Cueva de Ungurria, 20 km WSW Tezonapa on islandin the Rio Tonto, Mpo. de Tezonapa (Reddell,1973e:88)Tlacotepec Region+Cave 15 km ENE Tlacotepec, 500 m, Mpo. de PuenteNacional (Hall and Dalquest, 1963:229)Tlapaeoyan Region+Cueva de los Murcielagos, Tlapacoyan, Mpo. de Tlapacoyan(Villa R., 1967:327)TotuJa Region+Cave 9 km E Totutla, 750 m, Mpo. de ?Tlatetela(Hall and Dalquest, 1963:222)Tuxpin Region+Caves (3), Tuxpilla, 6 km SE Puerto de Tuxpan,Mpo. de Villa de Cazones (Martinez, 1941:1)+La Gruta I, near Tuxpan, 20 m, Mpo. de Tuxpan(D. S. Bureau of Sport Fisheries and Wildlife,1970:17)+La Gruta II, near Tuxpan, 20 m, Mpo. de Tuxpan(U. S. Bureau of Sport Fisheries and Wildlife,1970:17)+Cueva del Tigre, 5 km W Tuxpan, Mpo. de Tuxpan(Handley, 1966:300)Volcan de Tuxtla Region+Cueva EI Boqueron, Laguna Eneantada, 3 km E SanAndres Tuxtla, 300 m, Mpo. de San Andres Tuxtla(Hall and Dalquest, 1963:216-217)Cueva de Laguna Encantada-see Cueva EI Boquer6n+Cueva Puente de Piedra, 7 km W La Palma on Voleande Tuxtla (Vazquez-Yanes et al., 1975:73)Zacualpilla RegionYUCATAN+Gave 6 km WSW Zacualpilla, 2000 m, Mpo. de Zacualpan (Hall and Dalquest, 1963:253)€oastal Plain+*Well, Calcehtok, Mpo. de Dpichen (Reddell, 1977b:269)+Cave near ChoehoIa, Mpo. de ChochoIa (Reddell,1977b:257)+Cenote, 3.5 km NW Dzitya, Mpo. de Merida (Reddell,1977b:264)+Well, Finca San Pedro, Merida, Mpo. de Merida (Zullini,1977:76)+Cave 6 km Sand 5 km W Kinehil, Mpo. de Kinchil(Reddell,1977b:262)Artificial cave, Calle 24 x 19 in Merida, Mpo. deMerida (Reddell, 1977b:264)+Cenote at No. 97, Calle 24, Merida, Mpo. de Merida(Reddell,1977b:264)+Well, Calle 38, Merida, Mpo. de Merida (Zullini,1977:76)+*Well, iron mill in Dxkutzcab, Mpo. de Dxkutzcab(Reddell,1977b:273)+Well, Tahmek, Mpo. de Tahmek (Zullini, 1977:76)+Well, Teya, 13 km E Merida, Mpo. de Kanasin (Zullini,1977:76)+Well, Ticopo, Mpo. de Acanceh (Zullini, 1977:76)+Cenote 8 km NE Tixpehual, Mpo. de Tixpehual(Reddell, 1977b:288)+Cave 6 km N Tizimin, Mpo. de Tizimin (Reddell,1977b:288)Mine 5 km N Valladolid, Mpo. de Valladolid (Reddell,1977b:291)+Cenote, Yokdzonot, Mpo. de Tunkas (Zullini, 1977:76)*Cenote de las Abejas, 2 km N Kopoma, Mpo. de Kopoma(Reddell, 1977b:262)Cenote Acaneeh-see Cenote Chun Kapoc313

*Cenote Aka Chen, 1 km E Tixcancal, Mpo. de Tizimin(Reddell, 1977b:289)Cenote Amil, 6 km SAbala, Mpo. de AbaIa (Reddell,1977b:253)+*Cueva Ami!, 14 km SE and 2 km E Merida, Mpo. deMerida (Reddell, 1977b:264)+Cenote Anicabil, near the road to Yaxcaba, Mpo. deYaxcaba (Finch, 1965:113-115)+Cueva de Arena, 0.5 km SSW Santa Rosa, Mpo. dePeto (Villa R., 1953a:318)Cueva de Balaam Canche-see Grutas de Balankanche*Grutas de Balankanche, 4 km E Chichen Itza, Mpo.de ?Tinum (Andrews, 1970: 1-4, 2-3m; Reddell,1977b:284-286)Cueva de Belaam Canche-see Grutas de BalankancheCueva Bolonchen-see Grutas de BalankancheAcmn Caha Chen-see Cenote Kabahchen*Cenote Calchuhuim, 2 km E Hacienda San Bernardo,Mpo. de Maxcanu (Reddell, 1977b:263)*Cenote Calchum, 1 km E Hacienda San Bernardo,Mpo. de Maxcanu (Reddell, 1977b:263)Cenote Calchuntunil-see Cenote CalchuhuimCenote de los Camarones-see Cueva de Santa ElenaCueva de los Camarones-see Cueva de Santa Elena+Cueva de Carroza, Hocmn, Mpo. de Hocmn (Reddell,1977b:257)Banos Catacumbas-see Cenote de San Isidro*Cenote de Catzin, Catzin, Mpo. de Chemax (Reddell,1977b:256)Cueva de Caxcuy-see Cenote del Pochote+*Cueva Chac Mol, Tohil, Mpo. de Kaua (Reddell,1977b:258)Cenote de Chac Sikiin (Norte), Ruinas de Mayapan,Mpo. de Tecoh (Reddell, 1977b:278-279)Cenote de Chac Sikiin (Sur), Ruinas de Mayapan,Mpo. de Tecoh(Reddell, 1977b:279)+Cenote Chapultepec, Merida, Mpo. de Merida (Reddell,1977b:264)*Cenote Chen Mul, Ru.inas de Mayapan, Mpo. deTecoh (Smith, 1954:223-225, 230m; Reddell,1977b:279)+Cenote Chiuoh, Hacienda Mukuyche, Mpo. de Abala(Reddell, 1977b:25 3.254)Cenote Chiuok-see Cenote Chiuoh+Cenote Chotch, 3 km NE Piste, Mpo. de Tinum (Reddell,1977b:286)*Cenote Chun Kapoc, Acanceh, Mpo. de Acanceh(Reddell, 1977b:256)+Cenote Ciruak, 3 km N Piste, Mpo. de Tinum (Reddell,1977b:286)Cenote de Cocoyol, Cocoyol, Mpo. de Chemax (Reddell,1977b:256-257)+Cenote Conchita, Merida, Mpo. de Merida (Reddell,1977b:264-265)+Cenote del Country Club, Merida, Mpo. de Merida(Reddell, 1977b:265)*Cenote de la Culebra, 12 km N Muna, Mpo. de AbaIa(Reddell, 1977b:255)Cenote D, Ruinas de Ake, Mpo. de Tixkokob (Reddell,1977b:288)Cueva de las Derrumbes, 1 km S Tixcancal, Mpo. deTizimin (Reddell, 1977b:289)+Cenote Dhiba, near the road to Yaxcaha, Mpo. deYaxcaha (Finch, 1965: 110)Grutas de Dzab-Nah-see Grutas de Tzab-NahAguada Dzadz-see Cenote Dzadz+Cenote Dzadz, 10 km SW Chichen Itza, Mpo. deKaua (Reddell, 1977b:259)+Cenote de la Escuela Carlos Morales, Merida, Mpo. deMerida (Reddell, 1977b:265)*Cueva Escondida, 3.5 km S Kaua, Mpo. de Kaua(Reddell, 1977b:259)*Cenote G, Ruinas de Ake, Mpo. de Tixkokob (Reddell,1977b:288)+Cenote Geiser, Merida, Mpo. de Merida (Reddell,1977b:265)Cenote Geyser-see Cenote GeiserGreat Cenote-see Cenote Xtolok+Cenote Halal, 14 km and 2 km E Merida, Mpo.de Merida (Reddell, 1977b:265)Cueva de Hoctum-see Cenote de HoctUn*Cenote de Hoctun, 1 km W Hoctun, Mpo. de Hoctun(Pearse, 1938a:10, 11m; Robles Ramos, 1950:6366, 64m; Reddell, 1977b:257-258)Cueva de Hoctun-see Cenote de HoctUn+Cenote Hotun, 1.5 km SW Piste, Mpo. de Tinum(Reddell, 1977b:286)Cenote Hunto Chac (Cueva de Marney), 4 km SValladolid, Mpo. de Valladolid (Reddell, 1977b:291-292)Cenote Hunto Chac (Cueva del Pozo), 4 km S Valladolid,Mpo. de Valladolid (Reddell, 1977b:292)+Cenote Huntun, 4.8 km W Piste, Mpo. de Tinum(Reddell, 1977b:286)+Cenote de Hunucma, 2 km N Hunucma, Mpo. deHunucma (Reddell, 1977b:258)Cenote Iki!-see Cenote Ixi!+Cenote Itzmal Chen, Ruinas de Mayapan, Mpo. deTecoh (Finch, 1965:102m)+Cenote lxi!, 4.8 km SE Chichen Itza, Mpo. de Kaua(Reddell, 1977b:259)*Cenote Kahahchen, Mani, Mpo. de :VIani (Mercer,1896:140-141, 140m; Reddell, 1977b:262)+Cenote Kachak, 16 km W Hunucma, Mpo. de Hunucma(Finch, 1965:99, 100m)Cenote Kankirixche, 10 km NW Muna, Mpo. deMuna (Reddell, 1977b:268)*Actun Kaua, 1 km S Kaua, Mpo. de Kaua (Reddell,1977b:259-261)+Cenote de Kaua, Kaua, Mpo. de Kaua (Reddell,1977b:261)+Cenote de Kiki!, 8 km N Tizimin, Mpo. de Tizimin(Reddell, 1977b:289)Cenote Luchil-see Cueva Luchi!*Cueva Luchi!, 8 km SSE Merida, Mpo. de Kanasin(Reddell, 1977b:258)+Cenote Manzanilla, Merida, Mpo. de Merida (Reddell,1977b:265)+Cenote de Miramar, 1 km SW Telchac Puerto, Mpo.de Telchac Puerto (Reddell, 1977b:283)Motul Cave-see Cenote de Sambula, Motul+Cenote Mukuyche, Yuncu, Mpo. de Abala, Reddell,1977b:255)Caverna Murustun-see Cueva Muruztun+*Cueva Muruztun, 5 km S Tizimin, Mpo. de Tizimin(Reddell, 1977b:289)+Cenote Niagara, Merida, Mpo. de Merida (Reddell,1977b:265)Cenote Niagra-see Cenote Niagara*Cenote Nohchen, Sacalum, Mpo. de Sacalum (Arnold314

and Bohor, 1975:25-27, 26m; Reddell, 1977b:276)+Cenote Olivut, Merida, Mpo. de Merida (Reddell,1977b:265-266)*Cenote de Orizaba, 8 km S Buenaventura, Mpo. deTizimin (Reddell, 1977b:289)*Cueva de Orizaba, 8 km S Buenaventura, Mpo. deTizimin (Reddell, 1977b:290)Cavema Oxloth-see Cueva Oxolodt+*Cueva Oxolodt, Kaua, Mpo. de Kaua (Reddell,1977b:261)Cenote de la Paca, 7 km E Tikuch, Mpo. de Valladolid(Reddell, 1977b:292)Cenote Pamanche-see Cenote TaamancheCenote de los Pinos, 7 km S Buenaventura, Mpo. deTizimin (Reddell, 1977b:290)+Cenote de Piste, Piste, Mpo. de Tinum (Reddell,1977b:286)El Pochote Cave-see Cenote del Pochote*Cenote del Pochote, 10 km NW Muna, Mpo. deMuna (Reddell, 1977b:268)Cueva del Ponte-Cenote del PochoteCenote Poxil, 7 km SE Chemax, Mpo. de Chemax(Reddell, 1977b:257)Cenote de El Retiro-see Cenote de San IsidroCenote Sabacah, Sucopo, Mpo. de Tizimin (Reddell,1977b:290)Sacrificial Cenote-see Cenote Sagrado+Cenote Sagrado, Chichen Itza, Mpo. de Tinum (Tozzer,1957:191-192, fig. 706-707m; Reddell,1977b:286-287)Cenote Salud, 12 km S Valladolid, Mpo. de Tekom(Reddell, 1977b:282-283)*Cenote de Sambula, Kopoma, Mpo. de Kopoma(Reddell, 1977b:262)+Cenote de Sambul:i, Merida, Mpo. de Merida (Reddell,1977b;266)*Cenote de Sambula, Motul, Mpo. de Motul (Pearse,1938a:10, 11m; Reddell, 1977b:267-268)Cueva Sambul:i-see Cenote de Sambul:i, MeridaCenote San BuIha-see Cenote de Sambula, MeridaCueva San BuIha-see Cenote de Sambula, Meridaor Cenote de Sambula, MotulCenote San Cosme-see Cenote OlivutCenote de San Diego, 2 km W Cocoyol, Mpo. deChemax (Reddell, 1977b:257)*Cenote de San Isidro, Merida, Merida (Pearse, 1938a:10, 11m; Reddell, 1977b:266)Cueva de San Isidro-see Cenote de San Isidro*Cenote de San Jose, Merida, Mpo. de Merida (Reddell,1977b:266)Cenote de San Luis, 7 km S Tixcancal, Mpo. de Tizimin(Reddell, 1977b:290)+Cenote de Santa Ana, Valladolid, Mpo. de Valladolid(Reddell,1977b:292)*Cueva de Santa Elena, 5 km S Telchae Puerto, Mpo.de Telehac Puerto (Robles Ramos, 1950:67m,68-69; Reddell, 1977b:283)*Pozo de Santa Elena, 5 km S Telchac Puerto, Mpo.de Telchae Puerto (Reddell, 1977b:283)+Cenote Scan Yui, 3 km E Chiehen Itza, Mpo. deTinum (Hall, 1936:9m; Reddell, 1977b:287)+eenote Seeo, Chiehen Itza, Mpo. de Tinum (Reddell,1977b:287)+Cenote Shkolak, 3 km from Sitilpeeh, Mpo. de Izamal(Baker, 1895:23)"Cenote de Sihunehen, Sihunehen, Mpo. de Abal:i(Reddell,1977b:255-256)+Cenote Sisal, Valladolid, Mpo. de Valladolid (Reddell,1977b:292)+Cenote Skashek, 3 km from Sitilpeeh, Mpo. de Izamal(Baker, 1895:23)+Cenote Sodzil, 8 km Nand 1.6 km E Merida, Mpo.de Merida (Reddell, 1977b:266)*Cenote Sodzil, 5 km W Sueopo, Mpo. de Tizimin(Reddell, 1977b:290-291)"Cueva SodziI, 5 km W Sueopo, Mpo. de Tizimin(Reddell,1977b:291)Cenote SueiIa, 8 km E Hunku, Mpo. de Temozon(Reddell, 1977b:283)Cenote Sueopo, Sueopo, Mpo. de Tizimin (Reddell,1977b:291)Cenote SueuiI-see Cenote Sueila+Cenote Taamanehe, Taamanehe, Mpo. de Merida(Reddell, 1977b:267)*Cueva de Teeoh, Merida, Mpo. de Merida (Reddell,1977b:267)Cenote Tekom, Tekom, Mpo. de Tekom (Reddell,1977b:283)Cenote de Telehaquillo, Telehaquillo, Mpo. de Teeoh(Reddell, 1977b:279-280)+Cenote de Thompson, 2.5 km E Chichen Itza, Mpo.de Tinum (Reddell, 1977b:287)Cenote Tieimul-see Cenote Tikimul+Cenote Tikimul, 7 km S Chiehen Itza, Mpo. de Kaua(Reddell,1977b:261)Cenote Tixeaneal, Tixcaneal, Mpo. de Tizimin (Reddell,1977b:291)Acmn Tue-ie-see Aemn Tuz-ieCenote Tuxetue-see Cenote de San Luis+Aemn Tuz-ic, 5 km ENE Caleehtok, Mpo. de Opiehen(Reddell, 1977b:270)"Grutas de Tzab-Nah, 2 km S Teeoh, Mpo. de Teeoh(Stromsvik, 1956:463-465, 469m; Reddell, 1977b:281)Uki Cave-see Cenote Uki+Cenote Uki, 3 km NW Motul, Mpo. de Motul (Reddell,1977b:268)+Cueva Valladolid, Valladolid, Mpo. de Valladolid(Reddell,1977b:292)+Cenote Xal, near Chiehen Itza, Mpo. de Tinum (Reddell,1977b:287)Cenote XaIau-see Cenote Xtaeabiha+Cenote Chiea de Xanaba, 6.5 km SW Chiehen Itza,Mpo. de Kaua (Reddell, 1977b:261)+Cenote Grande de Xanaba, 8 km SW Chiehen Itza,Mpo. de Kaua (Reddell, 1977b:261)Cenote Xcan Yui-see Cenote Sean Yui+"Cueva Xeonsaeab, near Tizimin, Mpo. de Tizimin(Reddell, 1977b:291)+Cenote X-ebiz, Hocmn, Mpo. de Hoemn (Reddell,1977b:258)+Cenote Xix, Valladolid, Mpo. de Valladolid (Reddell,1977b:292)*Cenote Xkeken, 3 km N Dzit-Nup, Mpo. de Valladolid(Reddell, 1977b:293)Cenote Xlaeah-see Cenote Xlaka+Cenote Xlaka, Ruinas de Dzibilehaltun, Mpo. deMerida (Marden, 1959:110-129, 112-113m; Reddell,1977b:267)315

*Cenote Xtacabiha, 9 km NNE Tikuch, Mpo. deValladolid (Reddell, 1977b: 293)+Cenote X-Tojil, Libre Union, Mpo. de Yaxcaba (lullini,1977:76)Cenote Xtolok, Chichen Itza, Mpo. de Tinum (Hall,1936:9m;Reddell,1977b:287)*Cueva del Cenote Xtolok, Chichen Itza, Mpo. deTinum (Reddell, 1977b:287-288)+*Cenote Yunchen, Libre Union, Mpo. de Yaxcaba(Reddell,1977b:293)Cueva Yunchen-see Cenote Yunchen+Cenote Yuncu, Yuncu, Mpo. de ?Abala (Reddell,1977b:256)+*Cenote laci, 36 km ENE Tizimin, Mpo. de Tizimin(Holthuis, 1977:175)Cueva de lopilote Negro, Ruinas de Ake, Mpo. deTixkokob (Reddell, 1977b:288)Sierra de TieulCave on the Hacienda San Bernardo-see Actun Xpukil+Cave, Ticul, Mpo. de Tieul (Reddell, 1977b: 284)+Cave, Yocat, Mpo. de Ticul (Reddell, 1977b:284)Cueva de Aguacate, 2 km S Maxcanu, Mpo. de Maxcanu(Reddell, 1977b:262-263)Actun Ankah-see Actun ChacaljasGrutas de Calcehtok-see Actun Xpukil*Actun Chac, 15 km ESE Santa Elena, Mpo. deSanta Elena (Mercer, 1896:91-93, 92m; Reddell,1977b:277)Actun Chacaljas, 3 km S Calcehtok, Mpo. de Opichen(Reddell,1977b:269)+Cueva Chakxix, near Tekax, Mpo. de Tekax (Reddell,1977b:282)Actun Chen, 3 km W Kiuick, Mpo. de Oxkutzcab(Reddell,1977b:273)Actun Chom, 1 km S Calcehtok, Mpo. de Opichen(Reddell,1977b:269)*Actun Chukum, 2 km S Maxcanu, Mpo. de Maxcanu(Reddell,1977b:263-264)Actun Chunup, 2 km SW Maxeanu, Mpo. de Maxcanu(Reddell, 1977b:264)Cueva del Cinco de Mayas-see Cueva del Cinco deMayo+Cueva del Cinco de Mayo, 1 km SW Tekax, Mpo. deTekax (Finch, 1965:132m, 133; Reddell, 1977b:281-282)Actun Coch Leb, 3 km S Calcehtok, Mpo. de Opichen(Reddell, 1977b:269-270)+Actun Coyok, 6.5 km SW Oxkutzcab, Mpo. de Oxkutzcab(Hatt, 1953:15m; Reddell, 1977b:273)+Actun Ebizt, near Oxkutzcab, Mpo. de Oxkutzcab(Pearse, 1938a:l0, 11m; Reddell, 1977b:273)+*Actun Gongora, 1.5 km S or 3 km E Oxkutzcab,Mpo. de Oxkutzcab (Pearse, 1938a:10, 11m;Reddell,1977b:273-274)Actun Gongurrah-see Actun GongoraCueva de Gorgosa-see Actun Gongora+Actun Has, 6.5 km S Yocat, Mpo. de Ticul (Reddell,1977b:284)Actun Hi-see Actun J ihActun Ix-kix-see Aemn XkycActun Jih, 3 km W Tieul, Mpo. de Ticul (Reddell,1977b:284)Actun Kiuick, Ruinas de Kiuick, Mpo. de Oxkutz·cab (Reddell, 1977b:274)+Aetun Lara, between Yocat and Ticul, Mpo. de Ticul(Mercer, 1896:83m, 84; Hatt, 1953:17-18, 17m;Reddell,1977b:284)*Actun Loltun, 7 km SSW Oxkutzcab, Mpo. de Oxkutzcab(Mercer, 1896:98-125, 99m; Thompson,1897:6-22, 8.10m; Finch, 1965:140-147, 141m,143m; Reddell, 1977b:274-275)+Manhole Cave, 3.2 km S Yocat, Mpo. de Ticul (Reddell,1977b:284)Actun Nohcacab, Santa Elena, Mpo. de Santa Elena(Reddell,1977b:277-278)*Actun Okobichen, 8 km SW Santa Elena, Mpo. deSanta Elena (Reddell, 1977b:278)+Actun On, 4 km SW Oxkutzcab, Mpo. de Oxkutzcab(Finch, 1965:136-140, 137m)+Actun Oxkintok, 2 km SW Calcehtok, Mpo. de Opichen(Mercer, 1896:44m, 45-46; Hatt, 1953:18-20,19m; Reddell, 1977b:270)Cueva de Oxkintok-see also Actun Xpukil+Gruta de Pichjabin, near Muna, Mpo. de ?Muna(Gonzalez·Ochoa, 1963b:84)Caverna P'us-see Actun Puz+* Actun Puz, near Oxkutzcab, Mpo. de Oxkutzcab(Pearse, 1938a:l0, 11m; Reddell, 1977b:275-276)*Actun Sabaca, 6 km SW Tekax, Mpo. de Tekax(Mercer, 1896:146-149, 147m; Reddell, 1977b:282)Actun Sabacha-sec Actun SabacaActun Sabaka-see Actun SabacaCueva Sabre~see Actun SabacaCueva de San Roque-see Cueva Primera del Caminoa San RoqueSan Roque Road Cave-sec Cueva Primera del Caminoa San Roque+Cueva Primera del Camino a San Roque, on roadfrom Oxkutzcab to San Roque, Mpo. de Oxkutzcab(Reddell, 1977b:276)+Cueva Segunda del Camino a San Roque, on roadfrom Oxkutzcab to San Roque, Mpo. de Oxkutzcab(Pearse, 1938a:l0, 11m; Reddell, 1977b:276)Caverna Sasich-see Actun Sazich+Actun Sazich, near Calcehtok, Mpo. de Opichen(Reddell,1977b:270)Cueva de Sebaca-see Actun SahacaAcmn Silil, 3 km S Calcehtok, Mpo. de Opichen(Reddell,1977b:270)Actun Sitz-see Actun lizActun Spukil~see Actun Xpukil*Actun Tucil, 2 km S Muna, Mpo. de Muna (Reddell,1977b:268-269)Actun Tusil-see Actun TucilActun Xcoch-see Actun XkochCaverna Xkic-see Actun XkyeActun Xkoch, north of Santa Elena, Mpo. de SantaElena (Reddell, 1977b:278)Actun X-Koyoc-see Actun Coyok*Actun Xkyc, 2 km S Calcehtok, Mpo. de Opichen(Reddell,1977b:270-271)+Actun Xmahit, near Tekax, Mpo. de Tekax (Reddell,1977b:282)Actun Xpek, 1 km S Muna, Mpo. de Muna (Reddell,1977b:269)Caverna Xpueil-see Actun Xpukil316

*Actun Xpukil, 3 km S Calcehtok, Mpo. de Opichen(Hatt, 1953:20.24, 22m; Reddell, 1977b:271-273)Acron Ziizha, 2 km S Muna, Mpo. de Muna (Reddell,1977b:269)*Actun Ziz, Oxkutzcab, Mpo. de Oxkutzcab (Mercer,1896:128·129, 129m; Reddell, 1977b:276)ZACATECAS+Large cave 8.8 km S Moyahua, Mpo. de Moyahua (Geno·ways and Jones, 1968:744)+Cave 21 km W Valparaiso, Mpo. de Valparaiso (Handley,1959:149-150)BELIZE DISTRICTBELIZE+Cave under Gracy Rock, 8.5 km SSW Hattieville (Neilland Allen, 1962:84)(Sheet 21)CAYO DISTRICTFissure near Mountain Cow Cave, 2 km SE Caves Branch(Sheet 25)+Augustine Cave n. 1, Augustine, 22 km SSE San Ignacio(Williams, 1976c:602) ('!Sheet 28) (NOTE: This isprobably one of the Rio Frio Caves)+Augustine Cave n. 2, Augustine, 22 km SSE San Ignacio(Williams, 1976c:602) (?Sheet 28) (NOTE: This isprobably one of the Rio Frio Caves)+Augustine Cave n. 3, Augustine, 22 km SSE San Ignacio(Williams, 1976c:602) (?Sheet 28) (NOTE: This isprobably one of the Rio Frio Caves)*Buck's Bypass-St. Herman's Cave System, 2 km E CavesBranch, 120 m (Anonymous, 1974a:7; Albert andMcLeod, 1971:29)(Sheet 25)Caves Branch Cave System, 5 km NNE Caves Branch,40 m (Bartholomew, 1973:260-261) (Sheet 25)*Footprint Cave, 4 km SW Caves Branch, 120 m (Sheet25)+Millionario Cave, near Millionario, 45 km SSE San Ignacio(Williams, 1976c:602 (?Sheet 28)*Mountain Cow Cave, 2 km SE Caves Branch, 160 m(Sheet 25)Petroglyph-Satibe Cave System, 2.5 km S Caves Branch,160 m (Sheet 25)Rio Frio Cave A, near Augustine, 22 km SSE San Ignacio,450 m (?Sheet 28)Rio Frio Cave B, near Augustine, 22 km SSE San Ignacio,450 m (?Sheet 28)+Rio Frio Cave D, near Augustine, 22 km SSE San Ignacio,450 m (LaVal, 1973:25)(?Sheet 28)St. Augustine Cave, 0.5 km E Rio Frio, near Augustine,22 km SSE San Ignacio (?Sheet 28)(NOTE: This maybe one of the Rio Frio Caves)St. Herman's Cave-see Buck's Bypass-St. Herman's CaveSystem+San Antonio Cave, San Antonio, 10 km SSE San Ignacio(Williams, 1976c:602) (Sheet 23)Satibe Cave-see Petroglyph-Satibe Cave SystemTop Breakout Cave, near Caves Branch (Sheet 25)*Waterfall Cave, 5 km SSW Cavcs Branch, 160 m (Sheet25)COROZAL DISTRICT+San Antonio Cave, San Antonio, 2 km ENE Corozal(Quinones et aI., 1978:559) (Sheet 2)ALTA VERAPAZGUATEMALA+Cave, Cacao, Trece Aguas (Froeschner, 1960:661(NOTE: This is probably Sistema de Seamay-Sejul)+Cave, Finca Chicoyou, 1 km W Coban, 1300 m (Jones,1966:456)+Cave near Finca Chimoxan (Goodwin, 1934:8)+Caves, Chipoc (Goodwin, 1934:11)+Cave 14.5 km S Coban, 1360 m (Jones, 1966:464, 465)+*Grotte C3, Sierra de Pampur, W of Coban and S of theRio Chixoy Onegro (Delamare Deboutteville, 1976:127m)+*Grotte G3, Sierra de Pampur, W of Coban and S of theRio Chixoy Onegro (Delamare Deboutteville, 1976:126m)+Grotte de Bombil Pee, NE of Chisec (Dreux, 1974:205,210m)+Cueva del Cementerio de los Mayas, Lanquin (Gurnee,1968:150m)+Cueva del Cerrito, on the route from Raxjura to Coban,10 km from Raxjura (Delamare Deboutteville andJ uberthie, 1976:23)+*Cueva Chiacam, Sierra de Chama, near Finca Chiacam,NE of Coban(Delamare Deboutteville, 1976:128m)+Cueva Chiantejau, 8 km ENE Lanquin on Cahab6n road(Mohr, 1968:172)+*Cueva Chirrepeck, S of Coban, 1400 m (Beier, 1974:101,102)+Cueva Golondrinas, Finca Arenal, 10 km S Lanquin(Gurnee, 1968:157)+J ul'Pec Beneack Yaj, 10 km from Raxjura on road toCoban (Delamare Deboutteville and Juberthie, 1976:23)+Cueva .lui Seluc, 2 km S Lanquin (Gurnee, 1968:154m)*Grutas de Lanquin, 1 km NW Lanquin (Gurnee, 1962:26-27, 27m; Gurnee, 1968:151m, 152)+Sistema del Rio Candelaria, Municipio de Chisec (DelamareDeboutteville and Juberthie, 1976:19, 21m, 23)+Cueva Sakalkunte, near Senahu, 1800 m (Bartsch, 1906:117)Cueva Seamay-see Sistema de Seamay-Sejul*Sistema de Seamay-Sejul, 2 km SE Senahu, 26 km SELanquin (Gurnee, 1968:158-160, 158m; Smith, 1968:162, 163m; Shawcross, 1971 :63-64, 69m, 71)Cueva Sejul-see Sistema de Seamay-Sejul+Cueva Seanay, 2 km S Lanquin (Gurnee, 1968:152,154m)*Cueva Sepacuite n. 2, Finca Sepacuite, Senahu (Peckand Peck, 1973:70)Cueva Sepacuite n. 3, Finca Sepacuite, Scnahu (Peck andPeck,1973:70)+Siguan, 1.5 km W Lanquin (Gurnee, 1968:152)CHIMALTENANGO+Cave near Chocoyos, 7 km NW Patzun, 1520 m (Usinger,1966:298,360)+Cave, Santa Elena (Sanborn, 1936:99)317

ESCUINTLA+Cueva de los Ladrones, Finca Los Arcos, 210 m (LaVal,1973: 10)HUEHUETENANGOCaves, Km 130, plateau N of Huehuetenango-see Sumiderode Chemal n. 1 and Sumidero de Chemal n. 2125 ft. pit, plateau N of Huehuetanango-see Cueva deCuI+Cueva de Agua Escondida, 2 km SW Agua Escondida,10 km WNW Santa Ana Huista (Broughton, 1973:52-54, 55m, 57-59; Broughton and Boon, 1975:8-15,14m)+Cueva de las Calaveras, Tabacal, 5 km WNW Santa AnaHuista, 750 m (Sbordoni et a!., 1977:64-65, 65m)+Piccolo Pozzo di Chemal, Llanos de Chemal, 14 kmNNW La Capellania, 3200 m (Sbordoni et aI., 1977:69)Sumidero de Chemal n. 1, 14 km NNW La Capellania,3180 m (Shawcross et aI., 1974:63-64m, 66, 71;Sbordoni et aI., 1977:68)Sumidero de Chemal n. 2, 16 km NNW La Capellania,3290 m (Shawcross et aI., 1974:66, 67m)+Resumidero Chico, La Capellania, 3040 m (Sbordoniet aI., 1977:65m, 70)Cueva de CuI, 12 km NNW La Capellania (Shawcrosset aI., 1974:66, 69m)Cueva del Diablo-see PETEN+Cueva Esculike Alta, Sibila, 2940 m (Sbordoni ct aI.,1977:67m, 70.71)+Cueva Esculike Baja, Sibila, 2930 m (Sbordoni et aI.,1977:71)+Resumidero Grande, 1.5 km W La Capellania, 3030 m(Sbordoni et aI., 1977:69)Sima de los Grillos, 4 km N EI Retiro, 19 km SW Libertad,3120 m+Cueva del Madron n. 1, EI Madron, 3 km at 215 0SSWSan Juan lxcoy, 2750 m (Sbordoni et al., 1977:66,68)+Cueva del Madron n. 2, El Madron, 3 km at 215 0SSWSan Juan Ixcoy, 2740 m (Sbordoni et aI., 1977:68)+La Mina, La Capellania, 3030 m (Sbordoni et aI., 1977:70)'''Cueva de los Resadores, Santa Eulalia, 2500 m (Sbor·doni et aI., 1977:66, 67m)Sima de EI Retiro n. 1, EI Retiro, 16 km SW Libertad,2940 mSima de EI Retiro n. 2, EI Retiro, 16 km SW Libertad,2880 mSima de El Retiro n. 6, EI Rctiro, 16 km SW Libertad,2880 mCueva del Rodeo, 13 km SE Libertad, 3030 m+Resumidero de San Miguel, Llanos de San Miguel, SanJuan Ixcoy, 3300 m (Sbordoni et aI., 1977:69)Cueva de Santa Eulalia-see Cueva de los ResadoresCueva de Tabacal, Tabacal, 5 km WNW Santa AnaHuistaCueva del Tepescuintle-see PETENCueva de Yaxchilan-see Cueva de Juan Flores, PETENIZABAL+Crevice cave, Escobas, near San Tomas (Sanborn, 1936:95)+Talus cave, Escobas, near San Tomas (Sanborn, 1936:94)Cueva de la Coche, 2.5 km W Livingston (Peck and Peck,1973:69)Gruta EI Silvino, 34 km W Puerto Barrios (Gurnee,1962:29,30m)JUTIAPAPETEN+Cave 2 km W Tineal (=Tiucal) (Vercammen-Grandjean,1964:305,307)+Cave 12 km NNW Chinaja (Jones, 1966:443, 447, 452,456,463)+Second cave 12 km NNW Chinaja (Jones, 1966:443,463)+Cave 15 km NW Chinaja (.1ones, 1966:447)+Cave 3 km S Flores (Goodwin, 1955:1,2)+Cave, Toocog, 15 km SE La Libertad, 160 m (Jones,1966:447,456)+Cubixinal Cave, S of Flores (Thompson, 1967:252)+Cueva del Diablo, E side of Rio Usumacinta across fromYaxchilan, Chiapas, Mexico, 230 m (Sbordoni et al.,1974:11-12, 12m)Cueva Jobitzinaj, 7 km S Flores (Gurnee, 1962:28-29,29m)+Cueva de Juan Flores, E side of Rio Usumacinta acrossfrom Yaxchilan, Chiapas, Mexico, 100 m (McEachern,1974:141, 142m;Sbordoni et aI., 1974:10-11)Cueva Najohnaj Coholtunich, 14 km SW Flores+Cueva del Tepescuintle, E side of Rio Usumacinta acrossfrom Yaxchilan, Chiapas, Mexico, 240 m (Sbordoniet aI., 1974:13-14, 13m)Cueva de Yaxchilan-see Cueva de Juan FloresELPROGRESO+Cave at EI Progreso (Goodwin, 1934:9, 14)SOLOLA+Cucva Caman, near Lago de Atitlan (Causey, 1960:275,278)318

Appendix 2LIST OF TROGLOBITES BY STATEMexicoCampecheCreaseriella anops-Isopoda, CiroIanidaeMayaweckelia cenoticola-Amphipoda, HadziidaeMayaweckelia yucatanensis-Amphipoda, HadziidaeTyphlatya campecheae-Decapoda, AtyidaeTyphlatya pearsei-Decapoda, AtyidaeCreaseria morleyi-Decapoda, PalaemonidaeDiplocentrus mitchelli-Scorpiones, DiplocentridaeOonops coecus-Araneae, OonopidaeMetagonia torete-Araneae, PhoJcidaeMatta mckenziei-Araneae, TetrablemmidaeTohila atelomma-Saltatoria, GryllidaeChiapasOpisthobursa josephinae- Tricladida, DimarcusidaeDugesia mckenziei-Tricladida, DugesiidaeCaecidotea chiapas-Isopoda, AsellidaeCaecidotea vomeroi-Isopoda, AsellidaeCaecidotea zullinii-Isopoda, AsellidaeBrackenridgia acostai-Isopoda, TrichoniscidaeBogidiella orchestipes-Amphipoda, BogidiellidaeBogidiella sbordonii-Amphipoda, BogidiellidaeBogidiella tabascensis-Amphipoda, BogidiellidaeBogidiella vomeroi-Amphipoda, BogidiellidaeBithynops luscus-Decapoda, PalaemonidaeTyphlopseudothelphusa mocinoi-Decapoda, PseudothelphusidacTrichodactylus (Rodriguezia) mensabak-Decapoda,TrichodactylidaeTroglohya mitchelli-Pseudoscorpionida, HyidaePachychitra grandis-Pseudoscorpionida, SyarinidacCryptocellus sbordonii-Ricinulei, RicinoididaeHoplobunus zullinii-Opilionida, PhalangorlidaeMexotroglinus sbordonii-Opilionida, PhalangodidaeTroglostygnopsis anophthalma-Opilionida, Phalangodidae?Glomeridesmus sbordonii-Glomeridesmida, GlomeridesmidaeCleidogona felipiana-Chordeumida, CleidogonidaeCleidogona hunapu-Chordeumida, CleidogonidaePolylepiscus vomeroi-Polydesmida, Euryuridae?Aceratophallus scutigeroides-Polydesmida, RhachodesmidaeCaramba delburro-Polydesmida, TrichopolydesmidaeCaramba delnegro-Polydesmida, TrichopolydesmidaeCaramba grandeza-Polydesmida, TrichopolydesmidaeChiapadytes bolivari-Coleoptera, CarabidaeMexanillus sbordonii-Coleoptera, CarabidaeChihuahuaPsi/ochorus diablo-Araneae, PholcidaeCoahuilaCoahuilix hubbsi-Mesogastropoda, HydrobiidaeSpeocirolana thermydronis-lsopoda, CirolanidaeSphaerolana affinis-lsopoda, CirolanidaeSphaerolana interstitialis-lsopoda, CirolanidaeMexistenasellus coahuila-Isopoda, StenasellidaeMexiweckelia colei-Amphipoda, HadziidaeMexiweckelia particeps-Amphipoda, HadziidaeCicurina (Cicurella) coahuila-Araneae, AgelenidaeCambala speobia-Spirostreptida, CambalidaePrietella phreatophila-Cypriniformes, lctaluridaeDurangoMexiweckelia mitchelli-Amphipoda, HadziidaeLeptoneta limpida-Araneae, LeptonetidaePholcophora exigua-Araneae, PholcidaePsilochorus delicatus-Araneae, PholcidaeCryptocellus reddelli-Ricinulei, RicinoididaeGuerreroAlbiorix bolivari-Psueoscorpionida, IdeoroncidaePholcophora gruta-Araneae, PholcidaeCeuthauxus constans-Polydesmida, RhachodesmidaePararhachistes amblus-Polydesmida, RhachodesmidaeSpelaeogastrura guerrerense-Collembola, PoduridaeJuxtlacampa juxtlahuacensis-Diplura, CampodeidaeAnelpistina anophthalma-Thysanura, NicoletiidaeHidalgoParacophus cladonotus-Saltatoria, GryllidaeNiptus absconditus-Coleoptera, PtinidaeMexicoCaecoa arganoi-Opilionida, PhalangodidaeNuevo LeonSphaeromicola cirolanae-Padocopa, EntocytheridaeConi/era stygia-lsopoda, CirolanidaeSphaerolana affinis-lsopoda, CirolanidaeBrackenridgia palmitensis-lsopoda, TrichoniscidaeCylindroniscus cavicolus-Isopoda, TrichoniscidaeLeucohya heteropoda-Pseudoscorpionida, HyidaeLeucohya magnifica--Pseudoscorpionida, HyidaeSchizomus bartolo-Schizomida, SchizomidaeLeptoneta isolata-Araneae, LeptonetidaeLeptoneta reclusa-Araneae, LeptonetidaeNesticus nahuanus-Araneae, NesticidaeHoplobunus osorioi-Opilionida, PhalangodidaeGarcibius osorioi-Lithobiomorpha, LithobiidaeNuevobius cavicolens-Lithobiomorpha, LithobiidaeCeuthauxus palmitonus-Polydesmida, RhachodesmidaeOncopodura prietoi-Collembola, OncopoduridaePararrhopalites anops-Collembola, SminthuridaeParatachycampa boneti-Diplura, CampodeidaePodocampa cavernicola-Diplura, CampodeidaeMexaphaenops prietoi-Coleoptera, CarabidaeOaxacaEtlastenasellus mixtecus-lsopoda, Stenasellidae?Bogidiella arganoi-Amphipoda, BogidiellidaeBogidiella michaelae-Amphipoda, BogidiellidaeBogidiella niphargoides-Amphipoda, BogidiellidaeSpelaeomysis olivae-Mysidacea, LepidomysidaeAntromysis (Antromysis) reddelli-Mysidacea, MysidaeAlpheopsis stygicola-Decapoda, AlphaeidaeMacrobrachium villalobosi-Decapoda, PalaemonidaeNeoplaemon nahuatl~s-Decapoda,PalaemonidaeProcambarus (Austrocambarus) oaxacae oaxacae-Decapoda,Cambaridae319

Procambarus (Austrocambarus) oaxacae reddelli-Decapoda,CambaridaeDiplocentrus cueva-Scorpiones, Diplocentridael'roglohya carranzai-Pseudoscorpionida, Hyidae?Schizomus firstmani-Schizomida, SchizomidaeSchizopelma reddelli-Araneae, TheraphosidaeMetagonia martha-Araneae, PholcidaeNeogovea mexasca-Opilionida, SironidaeHoplobunus apoalensis-Opilionida, PhalangodidaeCleidogona baroqua-Chordeumida, CleidogonidaeMexicambala fishi-Spirostreptida, CambalidaePseudosinella bonita-Collembola, EntomobryidaePseudosinella finca-Collembola, EntomobryidaePlatynus (Mexisphodrus) urquijoi-Coleoptera, CarabidaePuehlaSpeocirolana pelaezi-Isopoda, CirolanidaeReddellobus troglobius-Spirobolida, SpirobollelidaeQueretaroTegenaria caverna-Araneae, AgelenidaeLeptoneta delicata-Arancae, LeptonetidaeHoplobunus queretarius-Opilionida, PhalangodidaeMexiterpes metallicus-Chordcumida, TrichopetalidacUnculabes arganoi-Polydesmida, RhachodesmidaeCixius orcus-Homoptera, CixiidaeMexaphaenops elegans-Colcoptera, CarahidaeParatrechus (Hygroduvalius) pallescens-Coleoptera,CarabidaeQuintana RooCrea.veriella anops-Isopoda, CirolanidacMayaweckelia cenoticola-Amphipoda, HadziidacAntromysis (Antromysis) cenotensis-Mysidacca, MysidacTyphlatya mitchelli-Decapoda, AtyidacTyphlatya pearsei-Dccapoda, AtyidacCreaseria morleyi-Decapoda, PalacmonidaeParaphrynus chacmool-Amblypygida, Phrynidac?Theotima martha-Araneae, OchyroceratidacOonops coecus-Araneac, OonopidacMetagonia torete-Arancac, PholcidacPholcophora pearsei-Arancac, Pholcidac?Ophisternon infernale-Synbranchiformes, SynbranchidacSan Luis PotosiEodrilus mexicanus-Haplotaxida, AcanthodrilidacDiaptomus (Microdiaptomus) cokeri-Calanoida, DiaptomidacSphaeromicola cirolanae-Podocopa, EntocythcridacSphaeromicola coahuiltecae-Podocopa, EntocythcridacMexilana saluposi-Isopoda, CirolanidaeSpeocirolana bolivari-Isopoda, CirolanidaeSpeocirolana pelaezi-lsopoda, CirolanidacMexistenasellus parzefalli-lsopoda, StenasellidaeMexistenasellus wilkensi-lsopoda, StcnascllidacSpherarmadillo cavernicola-lsopoda, SphacroniscidacTrichorhina boneti-lsopoda, SquamiferidaeBrackenridgia bridgesi-Isopoda, TrichoniscidacCylindroniscus vallesensis-lsopoda, TrichoniscidacMexiconiscus laevis-lsopoda, TrichoniscidacSpelaeomysis quinterensis-Mysidacca, LepidomysidaeTroglocubanus perezfarfanteae-Decapoda, PalacmonidaeTyphlochactas elliotti-Scorpioncs, ChactidacAphrastochthonius russelli-Pscudoscorpionida, ChthoniidaeTyrannochthonius pallidus-Pseudoscorpionida, ChthoniidaeAgastoschizomus huitzmolotitlensis-Schizomida, ProtoschizomidaeAgastoschizomus lucifer-Schizomida, ProtoschizomidaeSchizomus cookei-Schizomida, SchizomidaeParaphrynus velmae-AmbIypygida, PhrynidaeEuagrus anops-Araneae, DipIuridaeSchizopelma stygia-Araneae, TheraphosidaeMetagonia tlamaya-Araneae, PhoIcidaeCryptocellus osorioi-Ricinulei, RicinoididaeHoplobunus boneti-Opilionida, PhalangodidaeHoplobunus planus-Opilionida, PhalangodidaeRhagidia trisetata-Acarina, RhagidiidaeRhagidia weyerensis-Acarina, RhagidiidaeNewportia (Scolopendrides) sabina-Scolopendromorpha,CryptopidaeGlomeroides caecus-Glomerida, GlomeridaeMexiterpes egeo-Chordeumida, TrichopetalidaeMexiterpes .flShi-Chordeumida, TrichopetalidaeMexiterpes sabinus-Chordeumida, TrichopetalidaeUnculabes crispus-Polydesmida, RhachodcsmidaeUnculabes porrensis-Polydesmida, RhachodesmidaeTylogoneus rainesi-Polydesmida, TrichopolydesmidaeMexicambala russelli-Spirostreptida, CambalidaePseudosinella petrustrinatii-Collembola, EntomobryidaeAcherontides potosinus-Collembola, HypogastruridaeParacophus cladonotus-Saltatoria, GryllidaeParacophus lippus-Saltatoria, GryllidaeOeclidius hades-Homoptera, KinnaridaeMexaphaenops fish i-Coleoptera, CarabidaeTroglobacanius bolivari-Coleoptera, HisteridaeAstyanax jordani-Cypriniformes, CharacidaeTahascoOpisthobursa mexicana-Tricladida, DimarcusidaeBogidiella tabascensis-Amphipoda, BogidiellidaeMacrobrachium acherontium-Decapoda, PalaemonidaeMexobisium maya-Pscudoscorpionida, HyidaeSchizomus pecki-Schizomida, SchizomidaeParaphrynus chiztun-Amblypygida, PhrynidaeGlomeridesmus sbordonii-Glomeridesmida, GlomeridesmidaeTroglobacanius maya-Coleoptera, HisteridaePoecilia sphenops-Cyprinodontiformes, PoeciliidaeTamaulipasDugesia barbarae-Tricladida, DugesiidaeDugesia typhlomexicana-Tricladida, DugesiidaeEodrilus albidus-Haplotaxida, AcanthodrilidaeSphaeromicola cirolanae-Podocopa, EntocytheridaeSpeocirolana bolivari-lsopoda, CirolanidacSpeocirolana pelaezi-lsopoda, CirolanidaeMexicerberus troglodytes-lsopoda, MicrocerberidaeSpherarmadillo cavernicola-lsopoda, SphaeroniscidaeBrackenridgia bridgesi-lsopoda, TrichoniscidaeSpelaeomysis quinterensis-Mysidacea, LepidomysidaeTyphlochactas rhodesi-Scorpiones, ChactidaeAphrastochthonius major-Pseudoscorpionida, ChthoniidaeAphrastochthonius parvus-Pscudoscorpionida, Chthoniidacl'yrannochthonius troglobius-Pseudoscorpionida, ChthoniidaeParavachonium bolivari-Pseudoscorpionida, Vachoniidae320

Paravachonium superbum-I'seudoscorpionida, VachoniidaeSchizomus /ukensi-Schizomida, SchizomidaeSchizomus mitchelli-Schizomida, SchizomidaeSchizomus reddelli-Schizomida, SchizomidaeParaphrynus baeops-Amblypygida, PhrynidaeEuagrus cavernicola-Araneae, DipluridaeCicurina (Cicurusta) mina-Araneae, AgelenidaeTegenariG blanda-Araneae, AgelenidaeLeptoneta cnpilla-Araneae, LeptonetidaeTheotima pura-Araneae, OchyroceratidaeMetagonia pachona-Araneae, PholcidaeMetagonia pura-Araneae, PholcidaeCryptocellus osorioi-Ricinulei, RicinoididaeHoplobunus boneti-Opilionida, PhalangodidaeTrog/ostygnopsis inops-Opilionida, PhalangodidaeOrtholasma sbordonii-Opilionida, Nemastomatidae? Newportia (Scolopendrides) sabina-Scolopendromorpha,CryptopidaeG/omeroides promiscus-Glomerida, GlomeridaeC/eidogona pecki-Chordeumida, CleidogonidaeStrongy/odesmus harrisoni-Polydesmida, RhachodesmidaeUnculabes causeyae-Polydesmida, RhachodesmidaeSpeodesmus pecki-Polydesmida, TrichopolydesmidaeTy/ogoneus minus-Polydesmida, TrichopolydesmidaeMexicambala blanda-Spirostreptida, CambalidaeMexicambala inopis-Spirostreptida, CambalidaePseudosinella petrustrinatii-Collembola, Entomobryidae? Oncopodura prietoi-Collembola, OncopoduridaeParacophus caecus-Saltatoria, GryllidaeAntroforceps bolivari-Coleoptera, CarabidaeMexaphaenops intermedius-Coleoptera, CarabidaePlatynus (Mexisphodrus) profundus-Coleoptera, CarabidaeTrog/obacanius reddelli-Coleoptera, HisteridaeTrog/obacanius sbordonii-Coleoptera, HisteridaePtomaphagus (Ade/ops) mckenziei-Coleoptera, LeiodidaePtomaphagus (Ade/ops) trog/omexicanus-Coleoptera,LeiodidaeAstyanax jordani-Cypriniformes, CharacidaeVeracruzCaecidotea pasquinii-Isopoda, AsellidaeMexistenasellus magniezi-Isopoda, StenasellidaeCyathura sbordonii-Isopoda, AnthuridaeTrichorhina atoyacensis-Isopoda, SquamiferidaeBrackenridgia vi/Ia/obosi-Isopoda, TrichoniscidaeTyph/otricho/igioides aquaticus-Isopoda, TrichoniscidaeBogidie/Ia arganoi-Amphipoda, BogidiellidaeProcambarus (Austrocambarus) oaxacae reddelli-Decapoda,CambaridaeProcambarus (Austrocambarus) rodriguezi-Decapoda,CambaridaeTyph/ochactas reddelli-Scorpiones, ChactidaeVaejovis gracilis-Scorpiones, VaejovidaeMexobisium paradoxum-Pseudoscorpionida, HyidaeSchizomus firstrnani-Schizomida, SchizomidaeSchizomus pallidus-Schizomida, SchizomidaeNesticus arganoi-Araneae, NesticidaeMetagonia atoyacae-Araneae, PholcidaeG/omeroides addititius-Glomerida, GlomeridaeG/omeroides pellucidus-Glomerida, GlomeridaeC/eidogona crucis-Chordeumida, CleidogonidaeBonetesmus ojo-Polydesmida, OniscodesmidaeBonetesmus verus-Polydesmida, OniscodesmidaeAcutangu/us alius-Polydesmida, RhachodesmidaeAcherontides atoyacense-Collembola, HypogastruridaeOncopodrua atoyacense-Collembola, OncopoduridaeLitocampa atoyacensis-Diplura, CampodeidaePlatynus (Mexisphodrus) veraecrucis-Coleoptera, CarabidaeYucatanCreaseriella anops-Isopoda, CirolanidaeTrog/ophi/oscia laevis-Isopoda, PhilosciidaeTrichorhina pearsei-Isopoda, SquamiferidaeCylindroniscus maya-Isopoda, TrichoniscidaeJl'Iayawecke/ia cenoticola-Amphipoda, HadziidaeAntromysis (Antromysis) cenotensis-Mysidacea, MysidaeTyphlatya mitchelli-Decapoda, AtyidaeTyphlatya pearsei-Decapoda, AtyidaeCreaseria mor/eyi-Decapoda, PalaemonidaeDip/ocentrus anophthalmus-Scorpiones, DiplocentridaeVachonium boneti-Pseudoscorpionida, VachoniidaeVachonium cryptum-Pseudoscorpionida, VachoniidaeVachonium kauae-Pseudoscorpionida, VachoniidaeVachonium maya-Pseudoscorpionida, VachoniidaeParaphrynus chacmool-Amblypygida, PhrynidaeParaphrynus reddelli-Amblypygida, PhrynidaeCicurina (Cicure/Ia) maya-Araneae, AgelenidaeTheotima martha-Araneae, OchyroceratidaeOonops coecus-Araneae, OonopidaeMetagonia chiquita-Araneae, PholcidaeMetagonia torete-Araneae, PholcidaePho/cophora pearsei-Araneae, PholcidaeOrthoporus spelaeus-Spirostreptida, SpirostreptidaeOrthoporus zizico/ens-Spirostreptida, SpirostreptidaeCyphoderus innominatus-Collembola, EntomobryidaeMetasinella falcifera-Collembola, EntomobryidaeTrog/opodetes maya-Collembola, EntomobryidaeTohila ate/omma-Saltatoria, GryllidaeTyph/iasina pearsei-Perciformes, BrotulidaeOphisternon inferna/e-Synbranchiformes, SynbranchidaeGuatemalaAlta VerapazBogidie/Ia holsingeri-Amphipoda, BogidiellidaeTyph/opseudothelphusa juberthiei-Decapoda, PseudothelphusidaeTyph/opseudothe/phusa mitchelli-Dccapoda, PseudothelphusidaeAphrastochthonius verapazanus-Pselldoscorpionida,ChthoniidaeParaliochthonius strinatii-Pselldoscorpionida, ChthoniidaeMexobisium guatema/ense-Pseudoscorpionida, HyidaeTe/ema mayana-Araneae, TelemidaeAceratophallus scutigeroides-Polydesmida, RhachodesmidaePseudosine/Ia finca-Collembola, EntomobryidaeJuxtlacampa hauseri-Diplura, CampodeidaeSpeoco/podes franiai-Coleoptera, CarabidaeHuehuetenangoCaecidotea mitchelli-Isopoda, AsellidaeColombophi/oscia cavernicola-Isopoda, PhilosciidaeBogidie/Ia pasquinii-Amphipoda, BogidiellidaeMayaphaenops sbordonii-Coleoptera, Carabidae321

BelizeCayoMexobisilJm goodnighti-Pseudoscorpionida, HyidaeVachonilJm belizell.fe-Pseudoscorpionida, VachoniidaeMetagonia jarmila-Araneae, PholcidaeCynortina misteca-Opilionida, PhalangodidaeStygnomma pecki-Opilionida, PhalangodidaeJarmilka alba-Spirostreptida, Cambalidae322

SUPPLEMENTWhile this manuscript was in press several additionalpublications have appeared or been brought tomy attention which include infonnation on troglobitesfrom this region. The following citations followthe style used in the systematic review. The pagereferences in parentheses following the species namerefers to the page on which the species is mentioned.Bibliographic references follow the systematic citations.Diaptomus (Microdiaptomus) cokeri Osorio Tafall(p.81)Diaptomus cokeri: Bowman, 1979:225.Conilera stygia Packard (p. 84)Conilera stygia: Richardson, 1904:6.Conilera: Birstein, 1964:25.Creaseriella anops (Creaser) (p. 84)Creaseriella anops: Wilkens, 1979:265, 266, 267,fig. 2.Speocirolana bolivari (Rioja) (p. 86)Cirolana (part): Birstein, 1964:25.Speocirolana bolivari: Holt, 1973:223.Speocirolana peIaezi (Bolivar) (p. 86)Cirolana (part): Birstein, 1964:25.Speocirolana peIaezi: Holt, 1973:223.Caecidotea chiapas (Bowman) (p. 88)Caecidotea chiapas: Lewis, 1980:321,324.Caecidotea mitchelli Argano (p. 88)Caecidotea mitchelli: Lewis, 1980:324.Caecidotea pasquinii Argano (p. 88)Caecidotea pasquinii: Lewis, 1980:324.Caecidotea vomeroi Argano (p. 88)Caecidotea vomeroi: Lewis, 1980:324.Caecidotea zullinii Argano (p. 89)Caecidotea iullini: Lewis, 1980:324 (erroneousspelling).Mexistenasellus coahuila Cole and Minckley (p. 90)Mexistenasellus coahuila: Magniez, 1977:130.Mexistenasellus magniezi Argano (p. 90)Mexistenasellus sp. nov.: Magniez, 1977:130.Mexistenasellus parzefalli Magniez (p. 90)Mexistenasellus parzefalIi: Magniez, 1977:130.Mexistenasellus wilkensi Magniez (p. 90)Mexistenasellus wilkensi: Magniez, 1977:130.Brackenridgia bridgesi (Van Name) (p. 95)Protrichoniscus: Strouhal, 1953:175.Mayaweckelia cenoticoia Holsinger (p. 101)Mayaweckelia cenoticola: Wilkens, 1979:265, fig.2.Mayaweckelia spp. (part): Wilkens, 1979:266.Mayaweckelia yucatanensis Holsinger (p. 102)Mayaweckelia yucatanensis: Wilkens, 1979:265,266, fig. 2.Mayaweckelia spp. (part): Wilkens, 1979:266.Mexiweckelia colei Holsinger and Minckley (p. 102)Mexiweckelia (part): Holsinger and Longley, 1980:9, 10, 27, 51.Mexiweckelia mitchelli Holsinger (p. 102)Mexiweckelia (part): Holsinger and Longley, 1980:9, 10, 27, 51.Mexiweckelia particeps Holsinger (p. 102)Mexiweckelia particeps: Holsinger and Longley,1980:9, 10.Antromysis (Antromysis) cenotens&i Creaser (p. 103)Antromysis cenotensis: Wilkens, 1979:265, 266,fig. 2.Typhlatya campecheae Hobbs and Hobbs (p. 106)Typhlatya campecheae: Buden and Felder, 1977:114; Wilkens, 1979:265,266, fig. 2.Typhlatya (part): Wilkens, 1979:268,269.Typhlatya mitchelli Hobbs and Hobbs (p. 107)Typhlatya mitchelli: Buden and Felder, 1977:114; Wilkens, 1979:265,266,268, fig. 2.Typhlatya (part): Wilkens, 1979:268,269.Typhlatya pearsei (Creaser) (p. 107)Typhlatya pearsei: Buden and Felder, 1977:112; Wilkens, 1979:265,266,268,269, fig. 2.Typhlatya (part): Wilkens, 1979:268,269.Bithynops Iuscus Holthuis (p. 108)Bithynops: Strenth, 1978:67.323

Creaseria morleyi (Creaser) (p. 109)Creaseria morleyi: Wilkens, 1979:265, 266, 267,269, fig. 2.Macrobrachium villalobosi Hobbs (p. 109)Macrobrachium (part): Strenth, 1978:67,68.Neopalaemon nahuatlus Hobbs (p. 110)Neopalaemon: Strenth, 1978:67.Troglocubanus perezfarfanteae Villalobos (p. 110)Troglocubanus perezfarfanteae: Strenth, 1978:6768.Troglocubanus: Strenth, 1978:67,68.Mexobisium goodnighti Muchmore (p. 121)Mexobisium goodnighti: Muchmore, 1980:125.Albiorix bolivari Beier (p. 121)A lbiorix bolivari: Muchmore, 1979:317.Agastoschizomus lucifer Rowland (p. 125)Agastoschizomus lucifer: Rowland and Reddell,1980:21.Schizomus bartolo Rowland (p. 126)Schizomus bartolo: Row land and Reddell, 1980:2,4,5,6,7,8,9,10,15,19, fig. 1,5,21,40.Schizomus cookei Rowland (p. 127)Schizomus cookei: Rowland and Reddell, 1980:2,4,5,11, 15,16, 17,21,22, fig. 1, 12, 27,3031.Schizomus firstmani Rowland (p. 128)Schizomus firstmani: Rowland and Reddell, 1980:23-25, 26, 27, 28, 29, 30, fig. 63, 65, 67-68,74-75.Schizomus lukensi Rowland (p. 128)Schizomus lukensi: Rowland and Reddell, 1980:2,4,5,7,8-9,10,15,19, fig. 1,7,20,38.Schizomus mitchelli Rowland (p. 129)Schizomus mitchelli: Rowland and Reddell, 1980:2, 4, 5, 11, 15, 16, 17,21-22, fig. 1, 11,29,35,59-62.Schizomus pallidus Rowland (p. 129)Schizomus pallidus: Rowland and Reddell, 1980:2, 4, 5, 10, 11, 13, 14, 15, 16, 19, fig. 1, 13,26,43.Schizomus pecki Rowland (p. 129)Schizomus pecki: Rowland and Reddell, 1980: 16,23, 24, 25, 26, 29-30, 31, 32, fig. 63·64, 66,69,79.Schizomus reddelli Rowland (p. 129)Schizomus reddelli: Rowland and Reddell, 1980:2, 4, 5, 7, 10, 12, 13, 15, 17, fig. 1, 10,22,3637.Nesticus arganoi Brignoli (p. 141)Nesticus arganoi: Brignoli, 1979:919.Nesticus nahuanus Gertsch (p. 141)Nesticus nahuanus: Brignoli, 1979:919.Cryptocellus osorioi Bolivar (p. 157)Pseudocellus osorioi: Platnick, 1980:352.Discussion.-Platnick (1980) erected the newgenus Pseudocellus to include all of the describedMex iean cave and epigean rieinuleids previouslyassigned to Cryptocellus.Cryptocellus reddelli Gertsch (p. 159)Pseudocellus reddelli: Platnick, 1980:352.Cryptocellus sbordonii Brignoli (p. 159)Pseudocellus sbordonii: Platnick, 1980:352.Neogovea mexasca Shear (p. 159)Neogovea mexasca: Shear, 1979:238; Shear, 1980:2,4, 14, 15-17, 18, fig. 21-25.Discussion.-Shear (1980) erected the familyNeogoveidae to include the genus Neogovea andrelated genera.Hoplobunus boneti (Goodnight and Goodnight)(p. 164)Hoplobunus bonet: Elliott, 1979:66, 80 (erroneousspelling).Chirrepeckia lyncilecta Hoffman (insert on p. 189)Chirrepeckia lyncilecta Hoffman, 1976:744.746,fig. 1-3.Type-Iocality.-Cueva Chirrepeck (1400 m),south of Coban, Dept. Alta Verapaz, Guatemala.Distribution.-Known only from the typelocality.Discussion.-This is a minute pigmentless speciesknown only from the male holotype. Thegenus Chirrepeckia is monotypic and is withoutknown close relatives.324

Cambala speobia (Chamberlin) (p. 192)Cambala speobia: Shelley, 1979:551, 552, 553,556,557,564,566,567,568,569, fig. 9, 13.Cambala reddelli reddelli Causey, 1964:239-241,243, 246, pI. 58(fig. 1-4); Shelley, 1979:552,564.Cambala reddelli inornatus Causey, 1964:239,241-242, 246, pI. 58(fig. 5); Shelley, 1979:552,564.Cambala reddelli Causey, 1964:237, 239, 240;Shelley, 1979:566, 567.Type-Iocalities.-Of C. reddelli reddelli: BorderCave, Culbertson (:=Culberson) County, Texas,U.S.A.; of C. reddelli inornatus: Big Mouth Cave,2 miles north of Shamrock, Wheeler County,Texas, U.S.A.Discussion.-Shelley (1979) synonymized C.reddelli with C. speobia. This increases the rangeof this species west into New Mexico. Causey(1964) reported specimens intermediate betweenC. r. reddelli and C. r. inornatus from epigeanlocalities in Colfax County, New Mexico; and oftypical C. r. reddelli from an epigean locality inSan Miguel County, New Mexico. The populationsfrom west and northwest Texas and NewMexico have ocelli; regardless of the correctness ofShelley's synonymy, the populations of this speciesfrom caves in the Edwards Plateau and adjacentCoahuila are doubtless cave restricted.Mexicambala blanda Causey (p. 192)Mexicambala (part): Shelley, 1979:533.Mexicambala fishi Causey (p. 193)Mexicambala (part): Shelley, 1979:533.Mexicambala inopis Causey (p. 193)Mexicambala (part): Shelley, 1979:533.Mexicambala russelli Causey (p. 193)Mexicambala (part): Shelley, 1979:533.Oncopodura prietoi Bonet (p. 200)Oncopodura prietoi: Christiansen and Bellinger,1980:1010, 1011, 1016-1017, fig. 832B,831.Discussion.-Christiansen and Bellinger (1980)consider the family Oncopoduridae to be a subfamilyof the Entomobryidae. They report a specimenof this species from a cave in Lincoln County,New Mexico, U.S.A.Litocampa atoyacensis Wygodzinsky (p. 203)Plusiocampa (Litocampa) atoyacensis: Conde,1948:45-46.Oeclidius hades Fennah (p. 216)Oeclidius hades: Fennah, 1980:686.Astyanax jordani (Hubbs and Innes) (p. 238)Anoptichthys jordani: Breder and Roemhild,1947:33,34,35,40, fig. 1; Allee and Schmidt,1951:643; Coates and Atz, 1954:1468-1470;Keenleyside, 1955:205-206; Stolk, 1958:397;Friedman, 1962:75-76; Zeitlen, 1973:461-462;Herwig, 1976:297, 299; Olivereau et al., 1980:390.Anoptichthys: De Buen, 1941:5; Breder and Halpern,1946:155, 170, 174-175; Friedman,1962:73; Keenleyside, 1979:55; Olivereau etal.,1980:401.Anoptichthys sp.: Breder and Roemhild, 1947:33,34, 35, fig. 1.Blind cave fish: Breder and Roemhild, 1947:32,39.Astyanax jordani: Quinn, 1980:123-127.Astyanax mexicanus, cave derivatives of: Rasquin,1958:9.Astyanax hubbsi: Durand, 1979:198.Astyanax, eyeless: Wiley and Mitchell, 1971:1-12.Astyanax mexicanus, cave populations of: Herwig,1976:297-324, fig. 1-2, 3b-c, 4, 7-10, lla-c,12a-b, 13b; Dickson et al., 1979:120; Wilkenset aI., 1979:128,129-130,131-134,136, fig. 5,7, 9-10; Wilkens, 1980a:l03.111, fig. 1-3; Wilkens,1980b:232-238, fig. 1.Astyanax mexicanus, blind: Omura, 1975:99-112,fig. 1-9; Sadoglu, 1979:369-371.Astyanax mexicanus, Mexican cave fish: Beck,1978:23.Astyanax mexicanus, troglophilic: Dickson et aI.,1979:124.Anoptichthys (blind form ofAst)'anax mexicanus):Durand, 1979:196-205, fig. 1-14."Anoptichthys": Wilkens, 1979:269; Schemmel,1980:9, 10, 20, 21.Astyanax mexicanus f. anoptichthys: Schemmel,1980:9-22, fig. 1-4.Poecilia sphenops Valenciennes (p. 245)Poecilia sphenops, cave population: Wilkens et aI.,1979:127,128,131,135-136, fig. 11.Typhliasina pearsei (Hubbs) (p. 245)Typhlias pearsei: Yamaguti, 1961:67.325

Typhlias: De Buen, 1941 :9.Typhliasina pearsei: Miller, 1966:778, 800;Schemmel, 1977:191-202, fig. 1-8; Wilkens etaI., 1979:129; Wilkens, 1979:264, 266, 267,268, fig. l.Typhliasina: Miller, 1966:780.Ogilbia pearsei: Cohen and Nielsen, 1978:?; Wilkens,1979:266.Ophisternon infernale (Hubbs) (p. 246)Pluto: De Buen, 1941:8.Furmastix infernalis: Miller, 1966:778, 794.Furmastix: Miller, 1966 :780.Ophisternon infernale: Wilkens, 1979:264, 267,268, 269, fig. 1; Wilkens et aI., 1979:129,fig. 6.LITERATURE CITEDAllee, W. C., and K. P. Schmidt. 1951. Ecological animalgeography. 2nd ed. Rewritten and revised edition basedon Tiergeographie auf oekologischer Grundlage, byRichard Hesse. New York: John Wiley and Sons. 715 pp.Beck, L. 1978. Das Vivarium. Fische-Amphibien-ReptiJien-Niedere Tiere. Karlsruhe: Landessamlunger fur Naturkunde.48 pp.Birstein, Y. A. 1964. Fauna of U.S.S.R. Crustacea, Vol. VII,No.5. Freshwater isopods (Asellota). Zool. Inst. Akad.SSSR, n. ser., 47. Tr. from Russian. Jerusalem: IsraelProg. Sci. Transl.Bowman, T. E. 1979. Notodiaptomus caperatus, a new calanoidcopepod from phreatic groundwater in Bubuda(Crustacea: Diaptomidae). Bijdragen tot de Dierkunde,49:219·226.Breder, C. M., Jr., and F. Halpern. 1946. Innate and acquiredbehavior affecting the aggregation of fishes. Physiol.Zool.,19:154.190.Breder, C. M., Jr., and J. Roemhild. 1947. Comparative behaviorof various fishes under differing conditions ofaggregation. Copeia, 1947:29-40.Brignoli, P. M. 1979. Ragni del Brasile V. Due nuovi generi equattro' nuove specie dello stato di Santa Catarina (Araneae).Rev. Suisse Zool., 86:913·924.Buden., D. W., and D. L. Felder. 1977. Cave shrimps in theCaicos Islands. Proc. Biol. Soc. Washington, 90:108-115.Christiansen, K., and P. Bellinger. 1980. The Collembola ofNorth America north of the Rio Grande. Part 3. FamilyEntomobryidae. Grinnell, Iowa: Grinnell College. Pages785-1042.Coates, C. W., and J. W. Atz. 1954. Fishes of the world,pp. 1391.1640. In F. Drimmer, ed., The animal kingdom.New York: Doubleday. 3 vols.Cohen, D. M., and J. G. Nielsen. 1978. Guide to the identifi·cation of the genera of the fish order Ophidiiformes witha tentative classification of the order. United States Dept.Commerce NOAA Tech. Rept., NMFS Circ., 417.Conde, B. 1948. Contribution a la connaissance des campodeidescavemicoles de France. Notes Biospeol., 2:35·48.De Buen., F. 1941. Notas sobre ictiologia de aguas dulces deMexico. III. La lista de peces y la moderna clasificacion.Inv. Est. Limnol. Patzcuaro, 11. 9 pp.Dickson, G. W., J. C. Patton, J. R. Holsinger, and J. C. Avise.1979. Genetic variation in cave-dwelling and deep-seaorganisms, with emphasis on Crangonyx antennatus (Crustacea:Amphipoda) in Virginia. Brimleyana, 2: 119·130.Durand, J. P. 1979. Aspects ultrastructureaux des me canis·mes de la rudimentation retinienne chez I'Anoptichthysadulte, forme cavemicole aveugle de I'Astyanax mexica·nus (Characidae, Pisces). Canadian J. Zool., 57: 196·205.Elliott, W. R. 1979. A daddy-long-legs is not a spider. TexasCaver, 24:66,78-80.Fennah, R. G. 1980. New and little-known neotropicalKinnaridae (Homoptera: Fulgoroidea). Proc. BioI. Soc.Washington, 93:674-696.Friedman, L. R. 1962. A study of normal and malignantthymus tissue of the teleost Astyanax mexicanus in tissueculture. Bull. American Mus. Nat. Hist., 124:69-100,pI. 39-51.Herwig, H. J. 1976. Comparative ultrastructural investigationsof the pineal organ of the blind cave fish, Anoptichthysjordani, and its ancestor, the eyed river fish,Astyanax mexicanus. Cell Tiss. Res., 167:297-324.Hoffman, R. L. 1976. A new genus and species of cavedwellingmilliped from Guatemala (Diplopoda: Polydesmida).Rev. Suisse Zool., 83:743-746_Holsinger, J. R., and G. Longley. 1980. The subterraneanamphipod crustacean fauna of an artesian well in Texas.Smithsonian Contr. Zool., 308. 62 pp.Keenleyside, M. H. A. 1955. Some aspects of the schoolingbehaviour of fish. Behaviour, 8:183-248.Kennleyside, M. H. A. 1979. Diversity and adaptation in fishbehaviour. Zoophysiology, vol. 11. New York: SpringerVerlag. xiii + 208 pp.Magniez, G. 1977. Observations sur la biologie des Stenasellidae(Crustacea Isopoda Asellota des eaux souterraines).Actes 6e. Congr. Internatl. Speleol., Olomouc-CSSR,5:129-134.Miller, R. R. 1966. Geographical distribution of CentralAmerican freshwater fishes. Copeia, 1966:773-802.Muchmore; W. B. 1979. Pseudoscorpions from Florida andthe Carribbean area. 9. Typhloroncus, a new genus fromthe Virgin Islands (Ideoroncidae). Florida Entomol.,62:317-320.Muchmore, W. B. 1980. Pseudoscorpions from Florida andthe Caribbean area. 10. New Mexobisium species fromCuba. Florida Entomol., 63: 123-127.Olivereau, M., C. Aimar, and J.-M. Olivereau. 1980. Responsesof the teleost pituitary (goldfish, eel) to deionizedwater. Cell Tiss. Res., 208:389-404.Omura, Y. 1975. Influence of light and darkness on the ultrastructureof the pineal organ in the blind cave fish, Astya·nax mexicanus. Cell Tiss. Res., 160:99-112.Platnick, N. I. 1980. On the phylogeny of Ricinulei. Proc. 8.Internatl. Arachnol.-Kongr., Wien, 1980:349-353.Quinn, T. P. 1980. Locomotor responses of juvenile blindcave fish, Astyanax jordani, to the odors of conspecifics.Behav. Neural BioI., 29:123-127.Rasquin, P. 1958. Studies in the control of pigment cells andlight reactions in recent teleost fishes. Part 1. Morphologyof the pineal region. Part 2. Reactions of the pigmentarysystem to hormonal stimulation. Bull. American Mus.Nat. Hist., 115:1-68, pI. 1-10.Richardson, H. 1904. Contributions to the natural history ofthe Isopoda. Proc. United States Natl. Mus., 27:1-89.Rowland, J. M., and .T. R. Reddell. 1980. The order Schizo-326

mida (Arachnida) in the New World. III. Mexicanu8 andpecki groups (Schizomidae, Schizomus). J. Arachnol.,8:1-34.Sadoglu, P. 1979. A breeding method for blind Astyanaxmexicanus based on annual spawning patterns. Copeia,1979:369-371.Schemmel, C. 1977. Zur Morphologie und Funktion derSinnesorgane von Typhliasina pearsei (Hubbs) (Ophidioidea,Teleostei). Zoomorphologie, 87:191-202.Schemmel, C. 1980. Studies on the genetics of feeding behaviourin the cave fish Astyanax mexicanus f. anoptichthys.An example of apparent monofactorial inheritance bypolygenes. Z. Tierpsychol., 53:9-22.Shear, W. A. 1979. Huitaca ventralis, n. gen., n. sp., with adescription of a gland complex new to cyphophthalmids(Opiliones: Cyphophthalmi). J. Arachnol., 7:237-242.Shear, W. A. 1980. A review of the Cyphophthalmi of theUnited States and Mexico, with a proposed reclassificationof the suborder (Arachnida, Opiliones). AmericanMus. Novitates, 2705. 34 pp.Shelley, R. M. 1979. A synopsis of the milliped genus Camba·la, with a description of C. minor Bollman (Spirostreptida:Cambalidae). Proc. BioI. Soc. Washington, 92:551571.Stolk, A. 1958. Special condition of the interphase nucleus innormal and tumorous cells of fishes, amphibians and reptiles.Proc. Koninklijke Nederlandse Akad. Van Weten·schappen, ser. C, 61:395-416.Strenth, N. E. 1978. Zoogeographical significance of NorthAmerican troglobitic palaemonids (Crustacea, Decapoda).Texas Caver, 23:67-69.Strouhal, H. 1953. Neue Trichoniscinen aus tiirkischenHohlen. (2. Beitrag zur Kinntnis der tiirkinischen Isopoden.)Notes Biospeol., 8:167-183.Wiley, S., and R. W. Mitchell. 1971. A bibliography of theMexican eyeless characin fishes of the genus Astyanax.Preliminary compilation. Lubbock, Texas. Privatelyprinted. 12 pp.Wilkens, H. 1979. Reduktionsgrad und phylogenetischesAlter: Ein Beitrag zur Besiedlungsgeschichte der LimnofaunaYukatans. Z. Zool. Syst. Evolut..forsch., 17:262272.Wilkens, H. 1980a. Prinzipien der Manifestation polygenerSysteme. Z. Zool. Syst. Evolut..forsch., 18:103-111.Wilkens, H. 1980b. Zur Problematik der Rudimentation,untersucht an der Ontogenie des Auges von Hohlenfischen(Astyallax mexicanus). Z. Zool. Syst. Evolut.forsch., 18:232-238.Wilkens, H., N. Peters, and C. Schemmel. 1979. Gesetzmassigkeitender regressiven Evolution. Verh. Deutsch Zool.Ges., 1979:123-140.Yamaguti, S. 1961. Systema helminthum. Volume Ill. Thenematodes of vertebrates. New York: Interscience Publ.2 vols.Zeitlen, S. M. 1973. Hormonal induction of ovulation andspawning in the blind cave fish, Anoptichthys jordaniwith the use of human chorionic gonadotropin. Experientia,29:461-462.327

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