American Society of Mammalogists

Conservation Biology of the Cave of Mexico Author(s): Héctor T. Arita Source: Journal of Mammalogy, Vol. 74, No. 3 (Aug., 1993), pp. 693-702 Published by: American Society of Mammalogists Stable URL: http://www.jstor.org/stable/1382291 . Accessed: 01/10/2013 17:30

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This content downloaded from 132.248.49.122 on Tue, 1 Oct 2013 17:30:27 PM All use subject to JSTOR Terms and Conditions CONSERVATIONBIOLOGY OF THE CAVE BATS OF MEXICO

HECTORT. ARITA Departmentof Wildlifeand Range Sciences, University of Florida, Gainesville,FL 32611-0304 Presentaddress: Centro de Ecologia,Universidad Nacional Aut6noma de Mexico, ApartadoPostal 70-275, 04510 M6xico,D.F., Mdxico

The available informationon use of caves by Mexican bats was examined to determinethe effectivenessof a conservationstrategy based on diversity.Diversity was estimatedby species richness, or the numberof species present in a cave. Sixty of the 134 Mexican species of bats regularly roost in caves. Seventeen of these species tend to roost in caves with low species richness (segregationists), 14 tend to roost in caves with high species richness (integrationists), and 29 show no tendency in terms of the species richness of the caves (indifferent).Of the 215 caves includedin this study, 80% supportfew (three or less) species, whereas only 10% harbor six or more species. In general, species that share caves with many species form small or medium-sized colonies, and there is no positive correlation between species richness and total number of individuals in the caves. Few of the fragile and vulnerablespecies of Mexican bats roost in caves with high species richness or with large populations. A conservation plan based solely on diversity is not adequate for the protectionof cave bats in Mexico.

Key words: conservationbiology, cave bats, Mexico

Caves are essentialroosting sites for many cies that requirecaves as roosting sites, and species of bats (Dalquestand Walton, 1970; a catalogue of caves that would be impor- Kunz, 1982a; Tuttle, 1976). Of the 39 chi- tant in a conservation plan for these bats. ropteran species that occur in the United A central objective of this review is to States, at least 18 depend heavily on caves analyze the concept that the protection of for roosting sites, and many others use cav- sites with high diversity or high population erns as occasional refugia (Culver, 1986; levels assures the preservation of endan- Humphrey, 1975; McCracken, 1989). In- gered and threatenedspecies. I test such an formation is less precise for other parts of idea for Mexican cave bats by evaluating the world, but several species are known to the distributionof species of particularcon- rely completely on caves for roosting sites. cern among caves with different diversity An extreme case is Kitti's hog-nosed bat and abundance. If bats of special concern (Craseonycteris thonglongyai), which is frequently occur in caves with rich and known from only six caves in Thailand abundant communities, then the preserva- (Humphreyand Bain, 1990). For such spe- tion of these sites also would protect the cies, destruction of caves would mean in- fragile species. Conversely, if few or none evitable extinction. of the species of particularconcern roost in Mexico has a rich bat faunaof 134 species rich caves, separatestrategies would be nec- (Jones et al., 1988; Ramirez-P. et al., 1986), essary to protect the caves and the bats. many of which are known to use caves as roosts (Tuttle, 1976; Villa-R., 1967). Here, METHODS I review the available literature on use of Froma surveyof theliterature on distribution, caves by Mexican bats, provide a list of spe- naturalhistory, and taxonomyof Mexicanbats

J. Mamm., 74(3):693-702, 1993 693

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(Appendix I), I constructeda list of caves har- I developed a list of cave species of special boringpopulations of bats. For each cave, I com- concernfor conservationbased on the list of frag- piled a list of bat species, includingin some cases ile and vulnerablespecies of Mexican bats pre- informationon population size. sented by Medellin and Arita (in press). Addi- I identified the most important caves for the tionally, I included species that are endemic to conservationof Mexicancave bats based on their Middle America (Mexico and CentralAmerica) diversity and abundance. Diversity was esti- and those that virtuallyare endemic to this area, mated by species richness,or the numberof spe- barelyentering the United States(California leaf- cies reported from the cave. Abundance was nosed bat, Macrotuscalifornicus; Mexican long- quantified by the number of individuals of all nosed bat, Leptonycteris nivalis; Mexican species present at a given time in a cave. Caves long-tongued bat, Choeronycterismexicana). I of low abundancewere definedas sites sheltering included endemics in the analysis because the multispeciespopulations of < 1,000 individuals. Mexican government treats them as species of Caves with high and very high abundancewere special concern,along with rare and endangered defined as supportingmultispecies populations forms, in its ecological regulations. of > 1,000 and > 10,000 individuals, respective- The distribution of species among caves was ly. Importantsites were defined as caves with a analyzedusing "incidencefunctions." These are high species richnessand high or very high abun- graphsof the proportionof caves occupied by a dance. given species among categoriesbased on species Mexicanbats were classifiedin four categories richnessof the sites. These graphsare analogous of cave use. The first categoryincluded species to the incidencefunctions first used by Diamond for which caves are the main roost, whereasthe (1974, 1975) to study distribution of birds on second containedbats that regularlyuse caverns archipelagoesin relation to their colonization as refugia, but frequently rely on alternative ability. In caves, incidencefunctions can be used roosts. The third categoryincluded species that to analyze patterns of use by bats at different use caves occasionally,but usuallyroost in other levels of species richness. sites, and the last group comprised species not I classified Mexican species of cave bats in known to use caves as roosts. Species in the first threecategories of incidence(Fig. 1). Speciessuch two groups(caves as main or alternativeroosts) as P. davyi,which tend to occupy caves with high are treated here as cave bats, whereas those in speciesrichness, are "integrationists." Species that the last two categoriesare considered non-cave tend to be found in caves with few species, such bats. I assignedspecies to these categoriesbased as the Mexicanbig-eared bat (Plecotusmexican- on informationfrom sources in Appendix I and us) are "segregationists."Finally, species with no from personal experience. For consistency in apparentpreference for rich or poor caves, such classifyingspecies, I followed specialcriteria. For as the common vampire bat (Desmodus rotun- a few poorly known species, single or few reports dus) are "indifferent."I assignedspecies to these of use of caves in Mexico were sufficientto con- categoriesbased on the shape of their incidence sider them cave forms. For example, the trum- curves. pet-nosed bat (Musonycterisharrisoni) has been from a cave once reported only (Winkelmann, RESULTS 1962), but so little is known about its roosting habits that it was groupedwith species that use The literature survey yielded a total of cavesas alternativerefugia. Similar cases are those 215 Mexican caves with information on of Allen's big-earedbat (Idionycterisphyllotis- bats. In these caves, species richness ranges Czaplewski,1983) and free-tailedbats of the ge- from one to 13 species. The frequencydis- nus Nyctinomops(Carter and Davis, 1961;Jones tribution shows a negative exponential of the United et al., 1972). Conversely, species curve, with several caves supporting few Statesthat use caves as hibernacula,but that use species and few caves harboring many spe- other sites for summer refugia, generally were contain six or not consideredcave species in Mexico. For most cies. Only 10% of the caves of these species there are no reports of hiber- more species, whereas 80% shelter three or naculain Mexico. Examplesinclude several spe- fewer species (Fig. 2). cies of mouse-eared bats (Myotis-Humphrey, Most caves have low abundance; caves 1982; Schowalter, 1980). with high and great abundance are com-

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0.6 100 Pteronotus davyi a 0.5 MEXICANCAVES 80 0.4 n = 215 C 0.3 60 A 0.2 V 40 0.1 SE .:ii.i:i .. 20 ...... 1 2 3 4to6 > 6 i NUMBEROF BAT SPECIES o I+IT f i V, 1 2 3 4 5 6 7 8 9 10 11 12 13 NUMBEROF BAT SPECIES 0.6 Plecotus mexicanus b U ,<0.5 FIG.2. -Species richnessof cavebats in Mex-

0.4 ico. O 0.3 0.2 and the cave z _ stramineus), myotis (Myotis 0 0.1 velifer and Myotis peninsularis-Villa-R., Woloszin and Woloszin, a 1 2 3 4to6 > 6 1967; 1982). O NUMBEROF BATSPECIES The list of important sites for the con-

0 servation of Mexican cave bats includes 12 S0.6 --- Desmodus rotundus caves (Table 1). All these sites shelter more Ca -0- .* Macrotuswaterhousii C 0.5 ----.---- Artibeusjamaicensis than seven species of bats and supportlarge 0.4 multispecies populations. 0.3 * * . Sixty (45%)of the 134 species of Mexican 0.2 bats can be consideredcave species.Of these, 0.1------27 primarilyroost in caves, whereas 33 use 2 3 4to6 > 6 caves as alternativerefugia (Table 2). Of the NUMBEROF BATSPECIES remaining species, 18 use caves occasion- FIG.1.-Incidence functionsof cave bats in ally or rarely, and 56 do not use caves at Mexico: a, Pteronotusdavyi (n = 15), an inte- all. grationistspecies; b, Plecotus mexicanus(n = 8), The distributionof cave and non-cave bats a threeindifferent segregationistspecies; c, spe- taxa is not random. All Desmodusrotundus = Macrotuswa- among mormoop- cies, (n 63), and natalids are clas- terhousii (n = 18), and Artibeusjamaicensis (n ids, desmodontines, = 32). sified as cave species, as are the majority of glossophagines (10 of 12 species, binomial test with expectedP = 0.45; P < 0.01). Con- paratively rare. Sites with the largest pop- versely, fewervespertilionids and molossids ulations are refugia for the Mexican free- than expectedby chanceare consideredcave tailed bat (Tadarida brasiliensis),which in dwellers (14 of 43 vespertilionids, four of Mexico forms colonies rangingfrom a few 18 molossids; binomial test, P < 0.05 in thousand to >1 x 106 individuals (Cock- both cases). rum, 1969). Other species that frequently Fifteen fragileand four vulnerablespecies form large colonies (> 1,000 individuals) in regularlyuse caves. There is no demonstra- Mexican caves include some mormoopids ble association between use of caves and (mustached bat, Pteronotus parnellii; na- fragilityor vulnerability;the distributionof ked-backedbat, Pteronotusdavyi; the ghost- species of concern among cave and non- faced bat, Mormoops megalophylla - cave categories does not deviate from ex- Bateman and Vaughan, 1974), some phyl- pected by chance (contingency-tableanal- lostomids (Palla's long-tongued bat, Glos- ysis, x2 = 0.36, P > 0.05, d.f = 1). Eleven sophagasoricina; little long-nosed bat, Lep- of the Mexican cave bats are endemic to tonycteriscurasoae- Villa-R., 1967; Wilson Middle America, and three others barely et al., 1985), the funnel-earedbat (Natalus penetrate into the United States. There is

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TABLE1.- Mexicancaves with high species richness (more than seven bat species)and withhigh abundance(> 1,000 individuals). Fragile Cave Richnessa Abundanceb speciesc Chiapas Cueva de la Trinitaria 8 very high 2 Guerrero Grutade Juxtlahuaca 9 very high 1 Morelos Cueva del Cerro,Tequesquitengo 9 high 2 Cueva del Salitre,Tetecalita 12 high 0 Puebla Cueva de las Vegas 13 high 0 Sonora Mina del Tigre 9 very high 1 Tabasco La Murcielaguera 8 high 2 Cueva de Don Luis 9 high 1 Tamaulipas Cueva de Quintero 9 very high 1 Veracruz Cueva de la LagunaEncantada 9 high 1 Yucatan Gruta de Lol-Tuin 11 high 2 GrutaSpukil 9 high I

a Numberof speciesof batsin thecave. bNumberof individualsof all speciesof batsin thecave: high (> 1,000individuals); very high (> 10,000individuals). c Numberof fragile,vulnerable, or endemicspecies of batsin the cave. no association between cave dwelling and (Chrotopterusauritus), and M. harrisoni.Of endemism among Mexican bats (contingen- the remaining 25 species, only seven form cy-table analysis, x2 = 0.06, P > 0.05, d.f large populations of > 10,000 individuals. = 1). Among these are T. brasiliensis,P. parnellii, No definitive association exists between P. davyi, M. megalophylla,and N. strami- incidence and endangermentamong Mex- neus. There is no clear association between ican cave bats. Five of the 17 integrationist incidence and size of multispecies popula- = species are fragile, vulnerable or endemic. tions (contingency-tableanalysis, x2 3.21, = The figuresfor segregationistand indifferent P > 0.05, d.f 2). species are eight of 14 and 14 of 29, re- spectively. A contingency-table analysis DIscussIoN shows no significantdeviation from a ran- Many current conservation efforts con- dom distribution of species among inci- centrateon the protectionof areaswith high = P > = (Soul6 and Kohm, 1989; Wilson, dence types (x2 2.63, 0.05, d.f. 2). diversity Most species occur in caves in small col- 1988). The premise behind this approachis onies. Of the 60 cave species, 35 never form that sites with unusually high richness not colonies of > 100 individuals. Typical spe- only harbor more species, but also are pre- cies of this class include the spear-nosedbat ferred habitats for several endangered or (Mimon bennetti),the wooly false vampire threatened species. In some cases, the as-

This content downloaded from 132.248.49.122 on Tue, 1 Oct 2013 17:30:27 PM All use subject to JSTOR Terms and Conditions August 1993 ARITA--CONSERVATIONOF MEXICAN CAVE BATS 697 sumption seems reasonable. Tropical rain and the fringe-lippedbat (Trachopscirrho- forests, for example, are ecosystems of high sus). These species almost always occur in diversity that also featurea host of rareand caves by themselves or share the roost with endangeredspecies. In other instances, the few other species. Otherbats of special con- relationshipbetween diversity and the pres- cern, such as T. brasiliensisand M. penin- ence of threatenedspecies is not well estab- sularis, are less segregationist,but seldom lished. The data presented here show the occupy caves with more than five species. diversity approach to the conservation of In Mexican cave bats, diversity is a poor Mexican cave bats to be inadequate. criterion if the intention is to protect bats I have presented evidence for a lack of of special concern. correlationamong the three variables used Few of the fragile species regularlyroost here to assess the suitability of a cave as a in caves with high diversity and high abun- protected area: diversity, size of multispe- dance (Table 3). Entries in the lower right cies populations,and the presenceof fragile, cell of the table (integrationistbats that form vulnerable, and endemic species. For in- colonies of > 100 individuals) are the spe- stance, some caves with high or very high cies that would benefit the most from pro- abundance were excluded from the list of tection of criticalcaves. However, only three important sites because they have low di- species in this cell (the sac-wingedbat, Bal- versity. Several large colonies of the Mex- antiopteryxio, and the two species of Lep- ican free-tailedbat in the states of Durango, tonycteris)are fragile,vulnerable, or endem- Nuevo Le6n, and Queretarowere excluded ic. Conversely,one-half of the species in the from the list because T. brasiliensis is the upper left cell of the table (segregationists only species occupying these caves. Simi- that never form colonies of > 100 individ- larly, the caves in the Sierra de la Laguna uals) are fragile or vulnerable (M. macro- in BajaCalifornia Sur that shelterlarge pop- phyllum, M. bennetti, T. cirrhosus,L phyl- ulations of the endemic Myotispeninsularis lotis, and the fish-eatingbat, ). (Woloszin and Woloszin, 1982) were ex- These species would benefit little from the cluded because of their low species richness. protection of critical caves. Similarly, some of the caves that provide An effective plan for the conservation of shelterfor fragileor vulnerablespecies have Mexican cave bats would require a double low abundance. This happens because strategy: the protection of caves with un- threatenedspecies usually exist at low pop- usuallyhigh diversity and multispeciespop- ulation levels, but it also is due to the fact ulations and the management of cave bats that species of concern usually are not as- of special concern (fragile,vulnerable, and sociated with other species with higherpop- endemic species). Data presentedhere dem- ulations. Species such as M. bennetti, the onstrate that the implementation of a con- long-leggedbat (Macrophyllummacrophyl- servation plan for caves in Table 1 would lum), and M. harrisonialways occur in small not necessarilyprovide protection for frag- groups, but they seldom or never are found ile or vulnerablespecies. Despite this, caves in association with other, more abundant listed in Table 1 are important because the species. The protection of caves with un- greatconcentration of many species and in- usually high abundancewould add little to dividuals constitutes an unusual natural the conservation of these endangered spe- phenomenon that deserves protection. cies. Richness and abundanceby themselves are The majority of fragile species are seg- a sufficient reason for the conservation of regationist or indifferent to other species. caves in Table 1, despite their compara- Typical examples of fragile segregationist tively low contributionto the protection of bats are the endemic P. mexicanus and the endangeredspecies. carnivorousphyllostomids like M. bennetti Special conservation plans need to be de-

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TABLE2. - Cave bats that occur in Mexico. Size of Species Cave usea Incidenceb populationc Statusd Emballonuridae Balantiopteryxio Main I L, M F, E B. plicata Main S L, M E Peropteryxkappleri Alt. I L F P. macrotis Main I L, M F Noctilionidae Noctilio leporinus Alt. S L Mormoopidae Mormoopsmegalophylla Main I M, H Pteronotusdavyi Main I M, H P. gymnonotus Main I L F P. parnellii Main I M, H P. personatus Main I M Phyllostomidae Phyllostominae Macrotuscalifornicus Main Ind. M E* M. waterhousii Main Ind. M E Micronycterisbrachyotis Alt. Ind. L F M. megalotis Alt. Ind. L M. sylvestris Alt. Ind. L Lonchorhinaaurita Main I L F Macrophyllummacrophyllum Main S L F Mimon bennetti Alt. S L F Trachopscirrhosus Alt. S L F Chrotopterusauritus Alt. Ind. L F Glossophaginae Glossophagacommissarisi Alt. Ind. L G. leachii Alt. Ind. L G. morenoi Alt. Ind. L E G. soricina Alt. I L, M, H E Hylonycterisunderwoodi Alt. Ind. L Anourageoffroyi Main S L F, E Leptonycteriscurasoae Main I L, M F L. nivalis Main I L, M V, E* Choeronycterismexicana Main Ind. L E* Musonycterisharrisoni Alt. Ind. L V Carolliinae Carolliabrevicauda Alt. Ind. L, M C. perspicillata Alt. Ind. L, M C. subrufa Alt. Ind. L, M Stenodermatinae Dermanuraazteca Main S L E Artibeushirsutus Main Ind. L, M E A. intermedius Alt. I L A. jamaicensis Alt. Ind. L, M A. lituratus Alt. I L Desmodontinae Desmodusrotundus Main Ind. L, M D. youngii Main Ind. L F Diphylla ecaudata Main I L

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TABLE2.- Continued.

Size of Species Cave usea Incidenceb populationc Statusd Natalidae Natalus stramineus Main I M, H Myotis californicus Alt. Ind. L M. keaysi Main I L, M M. lucifugus Alt. Ind. L M. nigricans Alt. I L M. peninsularis Main Ind. M, H F, E M. thysanodes Alt. S L M. velifer Main Ind. M, H M. vivesi Alt. S L V, E Pipistrellussubflavus Alt. Ind. L Eptesicusfuscus Alt. I L Plecotus mexicanus Main S L, M E Plecotustownsendii Main S L, M Idionycterisphyllotis Main S L Antrozouspallidus Alt. S L, M Molossidae Tadaridabrasiliensis Main S M, H V Nyctinomopsaurispinosus Alt. Ind. L Nyctinomopsfemorosaccus Alt. Ind. L Nyctinomopslaticaudatus Alt. Ind. L a Cave use: Main, caves are the main roost for the species;Alt. (alternative),other types of roosts regularlyare used. b Incidence:I, integrationist;S, segregationist;Ind., indifferent. c Size of population:L, low (usually < 100 individuals);M, moderate(100-10,000 individuals);H, high (> 10,000 individuals). Status:F, fragile;V, vulnerable;E, endemic to Middle America;E*, restrictedto Middle Americaand a small portionof the southwesternUnited States. veloped for fragile,vulnerable, and endemic nrimerode especies de murci6lagospresen- species of cave bats. On-site researchis re- tes en una cueva. Sesentade las 134 especies quired to verify that these species still occur de murcielagosmexicanos utilizan las cue- in the caves where they have been recorded. vas regularmente.Diecisiete de estas espe- More detailed studies, focused on the de- cies son "segregacionistas"y comparten el mographyand ecology of some of the most refugiocon pocas especies; 14 son "integra- vulnerablebats, would identifythe key caves cionistas"que estainpresentes en cuevas con for each species. A complete conservation varias otrasespecies y 29 son "indiferentes" plan for Mexican cave bats should integrate the protection of caves in Table 1 with the TABLE3. -Relationship betweenincidence and preservationof caves that are needed by the size of populations among cave bats of Mexico. species of special concern. Valuesin cells are the numberof species in each category. RESUMEN Incidence Conservaci6nde los caver- murcidlagos Segrega- Indif- Integra- nicolas de Mdxico. En este trabajose revisa Size tionist ferent tionist Total la informaci6n existe sobre los murci&- que 10 de M6xico. Se analiza Lowa 18 7 35 lagos cavernicolas la Highb 4 11 10 25 eficacia de una estrategia de conservaci6n Total 14 29 17 60 basada en la diversidad, fue estimada que "Populations always are <100 individuals. usando la riqueza de especies, es decir el b Populationsusually are > 100 individuals.

This content downloaded from 132.248.49.122 on Tue, 1 Oct 2013 17:30:27 PM All use subject to JSTOR Terms and Conditions 700 JOURNALOF MAMMALOGY Vol.74, No. 3 que se encuentrantanto en cuevas con baja BAKER, R. H. 1956. Mammalsof Coahuila,Mexico. como con alta de El 80% Universityof KansasPublications, Museum of Nat- riqueza especies. ural History, 9:125-335. de las 215 cuevas incluidas en este trabajo BARBOUR,R. W., ANDW. H. DAVIS. 1969. Bats of tiene poblaciones de 3 especies, mientras America. University Press of Kentucky,Lexington, solamente el 10%_sirve de 286 pp. que refugio para BATEMAN,G. C., ANDT. A. VAUGHAN.1974. Nightly >6 especies. En general, las especies inte- activities of mormoopid bats. Journalof - gracionistas forman colonias pequefias o ogy, 55:45-65. corre- BIRNEY, E. C., J. B. BOWLES,R. M. TIMM, AND S. L. medianas, de manera que hay poca WILLIAMS. 1974. Mammaliandistributional rec- laci6n entre la riqueza de especies y el nii- ords of Yucatanand QuintanaRoo, with comments mero total de individuos en las cuevas. De on reproduction,structure, and status of peninsular las se populations. Occasional Papers, Bell Museum of especies frigiles y vulnerables, pocas Natural History, 13:1-25. refugian en cuevas con alta riqueza de es- CARTER, D. C., AND W. B. DAVIS. 1961. Tadarida pecies o alta abundancia.Los planes de con- aurispinosa(Peale) (Chiroptera: Molossidae) in North servaci6n basados solamente en la diver- America. Proceedingsof the Biological Society of Washington,74:161-166. sidad resultan inadecuados para la COCKRUM,E. L. 1969. Migrationin the guano bat, protecci6n de los cavernicolas Tadaridabrasiliensis. Pp. 303-336, in Contributions murci61lagos in mammalogy(J. K. Jones, Jr., ed.). Miscellaneous de M6xico. Publications,Museum of Natural History, Univer- sity of Kansas, 51:1-428. ACKNOWLEDGMENTS COCKRUM, E. L.,AND G. R. BRADSHAW. 1963. Notes I thank S. for his continued en- on from Sonora, Mexico. AmericanMu- Humphrey seum 2138:1-9. and Novitates, couragement generous support during my CULVER, D. C. 1986. Cave faunas. Pp. 427-443, in years at The University of Florida. This manu- Conservationbiology, the science of scarcityand di- script benefitedfrom the comments and sugges- versity(M. E. Soul6,ed.). Sinauer,Sunderland, Mas- tions of J. Eisenberg,P. Feinsinger,T. Fleming, sachusetts,584 pp. and Reiskind. This CZAPLEWSKI,N.J. 1983. Idionycterisphyllotis. Mam- R. Medellin, K. Redford, J. malian 208:1-4. at The Species, study is part of my Ph.D. dissertation DALQUEST, W. W. 1953. Mammals of the Mexican University of Florida and was made possible by state of San Luis Potosi. LouisianaState University scholarshipsfrom Consejo Nacional de Ciencia Studies, BiologicalSciences Series, 1:1-229. W. AND D. 1970. Di- y Tecnologia(Mexico), Universidad Nacional DALQUEST, W., W. WALTON. the Conser- urnalretreats of bats.Pp. 162-187,in Aboutbats Aut6nomade Mexico,and Tropical (B. H. Slaughterand D. W. Walton,eds.). Southern vationand DevelopmentProgram of The Uni- Methodist University Press, Dallas, Texas, 339 pp. versityof Florida.Financial support was pro- DIAMOND, J. 1974. Colonizationof explodedvol- vided The University of Florida's Program canic islands by birds:the supertrampstrategy. Sci- by e for Studies in Conservation, ence, 184:803-806. Tropical by Sigma 1975. Assemblyof species communities.Pp. Xi, and by a grant-in-aidfrom The American 342-444, in Ecologyand evolution of communities Society of Mammalogists.This is a contribution (M. L. Cody and J. M. Diamond, eds.). Harvard of the publicationseries of the Programfor Stud- UniversityPress, Cambridge, Massachusetts, 545 pp. ies in Conservation,The University of FENTON,M. B., ANDR. M. R. BARCLAY.1980. Myotis Tropical lucifugus.Mammalian Species, 142:1-8. Florida. HALL,E. R., ANDW. W. DALQUEST.1963. The mam- mals of Veracruz.University of KansasPublications, LITERATURECITED Museum of Natural History, 14:165-362. ALVAREZ,T. 1963. The Recent mammalsof Tamau- HARRISON,D. L. 1975. Macrophyllummacrophyl- lipas, Mexico. University of Kansas Publications, lum. MammalianSpecies, 62:1-3. Museum of NaturalHistory, 14:363-473. HERMANSON, J. W., AND T. J. O'SHEA. 1983. Antro- ALVAREZ,T., ANDL. GONZALEZ-Q.1970. Andlisis zous pallidus. MammalianSpecies, 213:1-8. roosts and com- polinico del contenidogistrico de murci61agosGlos- HUMPHREY,S. R. 1975. Nursery sophaginae de Mexico. Anales de la Escuela Na- munity diversity of Nearctic bats. Journalof Mam- cional de CienciasBiol6gicas (Mexico), 18:137-165. malogy, 56:321-346. ALVAREZ,T., ANDJ. RAMIREz-P.1972. Notas acerca 1982. Bats,Vespertilionidae and Molossidae. de murci61agosmexicanos. Anales de la EscuelaNa- _-.?Pp. 52-70, in Wild mammalsof North America, cional de CienciasBiol6gicas (Mexico), 19:167-178. biology, management,economics (J. A. 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