Journal of Natural History

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Journal of Natural History Publication details, including instructions for authors and subscription information: http://www.informaworld.com/smpp/title~content=t713192031 Food habits of the threatened bat Leptonycteris nivalis (Chiroptera: Phyllostomidae) in a mating roost in Mexico

Online Publication Date: 01 January 2007 To cite this Article: Sánchez, Ragde and Medellín, Rodrigo A. (2007) 'Food habits of the threatened bat Leptonycteris nivalis (Chiroptera: Phyllostomidae) in a mating roost in Mexico', Journal of Natural History, 41:25, 1753 - 1764 To link to this article: DOI: 10.1080/00222930701483398 URL: http://dx.doi.org/10.1080/00222930701483398

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The publisher shall not be liable for any loss, actions, claims, proceedings, demand or costs or damages whatsoever or howsoever caused arising directly or indirectly in connection with or arising out of the use of this material. © Taylor and Francis 2007 Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 oeoVle ta.20) lo hr ssm nietifraino t it saresult a as diet, its on information indirect some is there Also, 2004). al. et Moreno-Valdez fisdsrbto (A distribution its of 41,Me 04510, ora fNtrlHistory Natural of Journal O:10.1080/00222930701483398 online DOI: 1464-5262 print/ISSN 0022-2933 ISSN Sa Ragde Correspondence: are young the summer, south- toward of back end migrate bats the young at and (Te roosts; adults Mexico and central maternity disband, depart establish 2005). roosts apparently maternity to females (Simmons weaned, pregnant areas (March), Guatemala and spring northern September early and for between In season, winter). mating Mexico to the that (autumn during suggests south February Mexico evidence south-central preliminary New to in understood, south well remain from not (USA) females ranges is distribution ecology Texas Its migration g. their 24 west Although to up and of mass Mexico body a attaining 1988), Wilkins nivalis Leptonycteris Introduction Keywords: vulnerable a that plants food the on data present and collect to was study bat, nectarivorous this migratory of objective The Abstract ( Mexico D.F., Ecologı de Instituto SA RAGDE Mexico in roost mating a in bat Phyllostomidae) threatened (Chiroptera: the of habits Food dasylirioides arborescens eeeaie.Ltl skonaottede fti a,epcal ntesuhr afo its of half southern the in especially bat, ( this Cactaceae fur families: and of five faeces from diet from found was grains the species Pollen plant about species. beneckei 10 this known to for belonging is Pollen known distribution. roost Little mating examined. only the were is cave This Mexico. aepnclt nlrsecsta r omnyatiue obtpliain hl h te two other insect-pollinated. the primarily while be pollination, to bat considered are to which attributed inflorescences, commonly four the spicate are Of had that sexes. between inflorescences diet paniculate in differences have significant no were There November. cetd2 a 2007) May 29 Accepted uhr aesuidcranapcso h itof diet the of aspects certain studied have Authors ,Bmaaee( Bombacaceae ), xc,DF,Mxc.Eal [email protected] Email: Mexico. D.F., ´xico, ,Fbca ( Fabaceae ), , .horrida A. hrpea osrain odhabits, food conservation, Chiroptera, ´ CE ORG .MEDELLI A. RODRIGO & NCHEZ a nvria ainlAuto Nacional Universidad á, nhz il,Isiuod Ecologı de Instituto Biol., ńchez, le 2001). ´llez ´ Susr,16)i n ftelretncaioosbt Hnlyand (Hensley bats nectarivorous largest the of one is 1860) (Saussure, vrzadGonza and lvarez , .inaequidens A. aladahoustoniana Calliandra eb aesculifolia Ceiba 07 12–8:1753–1764 41(25–28): 2007; , etncei nivalis Leptonycteris # , 07Tyo Francis & Taylor 2007 .salmiana A. lz17;Esel 92 ade 97 el 1979; Hevly 1977; Gardner 1972; Easterla 1970; ´lez , a nvria ainlAuto Nacional Universidad á, suoobxellipticum Pseudobombax sddrn t esnlsa tacv nTepoztlan, in cave a at stay seasonal its during used , , nm eMe de ńoma ahnaungulata Bauhinia etncei nivalis Leptonycteris .Bt hwdtehgetdeaydvriyin diversity dietary highest the showed Bats ). .nivalis L. etncei nivalis Leptonycteris ´ xc,Cua nvriai,Me Universitaria, Ciudad ´xico, N ,adAayldca ( Amaryllidaceae and ), nm eMe de ńoma otyi h otenpart northern the in mostly ,Cnovlca ( Convolvulaceae ), etrvru bats nectarivorous , xc,Cua Universitaria Ciudad ´xico, Agave pce,two species, Stenocereus Ipomoea Agave ´xico, Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 h nlso fti pce nednee pce it sacneuneo recent of Medellı consequence 1994; a Service Wildlife is and lists Fish ( US species bat factors human-related 1985; long-nosed endangered various Mexico al. to In attributable in et mainly 1994). been (Wilson species have Service that this abundance Wildlife in decreases and of Fish inclusion (US The government US the nivalis by and 2006) guild. entire the for scarce are topics oet Hfmn ta.18) h lmt ssbrpclwt oto h rainfall the of most with subtropical is Garcı climate following pattern The climatic 1986). The summer. al. in occurring et (Hoffmann ( temperate forests includes area the in Vegetation whether test season. to Methods mating (2) the and during Mexico, females and Morelos, males Diablo, between del differences Cueva dietary the are in there season their of mating diet of the their document distribution more to particularly and (1) gain were: is composition study 1994). are: latter Service the bat Wildlife The about this and known habitats. Fish of (US is foraging recovery resources little and the alimentary very (caves) for because steps status important important roost most about two information the that suggests (Sa bats nectarivorous in differences female suggested and have studies male Nagy several between A and Indeed, (Kunz periods requirements 2000). such during Entwistle (Te food males and of bat is those it Racey this from periods, differ lactation 1988; for may and diets known reproduction their during roost that increase feasible of bats mating 1987; female diet only of the al. requirements the about energy et information is little the which Eguiarte is in Mexico, there However, 1977; distribution 2000). its Slauson Schaffer of 2000; nivalis and al. part et central Schaffer Arizaga and 1997b; southern (i.e. 1997a, al. studies et Valiente-Banuet system pollination of e iboi ntemncplt fTpzln ntesaeo oeo,Mxc (18 Mexico Morelos, of state the in Tepoztlan, of municipality the in is Diablo del 19.9 temperature annual mean 1754 rprd eln hmwt rnprn alpls.Fea ape eeotie by obtained were samples Faecal polish. nail transparent with them sealing prepared, (Medellı bodies bats’ the scored. (not was males testicles) and post-lactating) enlarged or or pregnant reproductive, reproductive (not females of status reproductive 99 rdn Y S)wsue.Tentwsoe o uigtepro hnbats when period were the animals captured during All h 4 h). 4:00 for and open 0:00 Inc., (between (Avinet was foraging net (precision net after mist weighed The cave m 6 the used. A to was roosted. returned typically USA) individuals NY, 400 a Dryden, about contain where to entrance, (Caballero estimated the 2001 was of gallery December main and the October between in individuals colony 2004). 3000 bat about The of 1986). maximum al. et (Hoffmann ´ vrz20;Rehr ta.20;Tcak 05.Hwvr culdt bu l these all about data actual However, 2005). Tschapka 2003; al. et Riechers 2000; lvarez etncei nivalis Leptonycteris nodrt rvd rca aafrcnevto n eoeyproe,tegaso our of goals the purposes, recovery and conservation for data crucial provide to order In olnwssmldfo h u fcpue as sn e-arnn qae oldover rolled squares gel-safranine using bats, captured of fur the from sampled was Pollen tec ii aswr atrdisd h aei ml ordrlctd20mfrom m 250 located corridor small a in cave the inside captured were bats visit each At u 03 9 hr sn nomto ntede fbt naiigCeadlDal,Morelos, Diablo, del Cueva inhabiting bats of diet the on information no is There . ) ta lvto f18 ...Tecv a 6criosad1 galleries 11 and corridors 26 has cave The a.s.l. m 1883 of elevation an at W), sctgrzda hetndseis(iroOiild aFederacio la de Oficial (Diario species threatened a as categorized is .Sa R. nhz&R .Medellı A. R. & ńchez etncei nivalis Leptonycteris ¡ scniee nagrdb h UN(hrpeaSeils Group Specialist (Chiroptera IUCN the by endangered considered is . ) oer egh(aie,precision: (caliper, length forearm g); 0.5 ne l 93.Praetmcocp ldso ahsml were sample each of slides microscope Permanent 1983). al. et ń u ,adrifl s16. m(NG 03.TeCueva The 2003). (INEGI mm 1463.2 is rainfall and C, eoeyPa U ihadWllf evc 1994) Service Wildlife and Fish (US Plan Recovery ) ń etncei nivalis Leptonycteris Pinus spp., Quercus n t eprlcagsduring changes temporal its and a(93 s(A)Ca(w is (1973) á p. n rpcldeciduous tropical and spp.) le 01.Bcuethe Because 2001). ´llez ¡ n20) h Mexican The 2003). ń . m,sx n the and sex, mm), 0.1 nhzCssand ńchez-Casas n19,2002). 1991, ń 2 )(w)(i u 59 9 9 )g: N, L. L. Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 ( o h itof diet the for arborescens eecletd e odpat eeietfe:Ccaee( 136 Cactaceae of total identified: visited a were was visits samples our plants Diablo faecal During 98 del food and visits. ( fur 123 11 Cueva Ten and Bombacaceae of captured, the collected. were total females) a 2002 20 were and for males February (116 weeks individuals and 2 every 2001 approximately September Between Results their because records discarded, accidental. from were considered pollen grains spores was Pollen fungal of samples sexes. and the between frequencies corn) in use and presence total plant pine Yates’ in (like with in and plants used agaves anemophilous differences spicate were and significant tests paniculate the Chi-square between determine in (1988). deposited to Thomas were following correction, fur sample, and per faeces recorded (UNAM). from the Geologia samples de Gonza and Instituto of 1970; the 1968, Slides (UNAM) of (Palacios 1986). Collection Mexico studies al. Palynological other well Instituto et de as from Quiroz the UNAM) descriptions Autonoma Biologia, 1969; of and de Nacional photographs (Instituto Collection with Herbarium Palynological Universidad MEXU as the the the of in Collection of Palynological samples reference Geologia to de samples our from acetolysis the in 100 purified pollen (A the gel of glycerine subsample pollen of melted a exine mixing with the by technique of prepared components were main the Slides are grains. which pseudochitin or chitin sporopollenin, Bauhinia ooiz(92.Ti diinlse lmntsogncmte rsn ntefaeces the in the present identification matter for organic grains eliminates was pollen sample step prepare faecal additional by to Each proposed (A This modification order h. the 1–2 with (1992). In ca 1966), (Erdtman Horowitz bag. for used bag was paper technique cloth standard labelled acetolysis clean a a in in stored animal captured each keeping of rsn nmr hn7%o h aclsmls l eann oln eefudi less in fur found were On pollens samples. remaining all the samples; faecal of the 30% of 70% than than more in present arborescens ln h lnae cp)dmnt h ape ih5 n 5 epciey while respectively; 45% and 54 with samples the dominate scape) elongated the showed along samples fur dasylirioides the species-specific Agave of of 57% observation of the the and species to four prevented samples due all that primarily Combining genus, slide of characteristics. gel-based level diagnostic each the to in possible debris only abundant was the pollen with of 50%, identification samples than higher frequencies at found were of plants exception remaining the and (93.49%) gv dasylirioides Agave ´ olngan eeietfe sn ih irsoe(lmu,C3;40 CH30; (Olympus, microscope light a using identified were grains Pollen odtriea niiulbtsde,tepeec/bec fple pce was species pollen of presence/absence the diet, bat’s individual an determine To olnfo l ln pce a on nbt u n aclsmls ihteexception the with samples, faecal and fur both in found was species plant all from Pollen h olno orseisof species four of pollen The vrzadGonza and lvarez tncru beneckei Stenocereus 6 n ycmaigtemrhlgclpoete n esrmnso olngrains pollen of measurements and properties morphological the comparing by and ) TbeI). (Table ) ,Fbca ( Fabaceae ), and ahnaungulata Bauhinia etncei nivalis Leptonycteris Agave eb aesculifolia Ceiba , lz17;Hrwt 92 rc n rs 99,wt h xeto of exception the with 1999), Pross and Bruch 1992; Horowitz 1970; ´lez and .horrida A. hc a eoddol ntefrsmls olnof Pollen samples. fur the in only recorded was which , oiae h aclsmlsadwr h nytople groups pollen two only the were and samples faecal the dominated .horrida A. aladahoustoniana Calliandra , .inaequidens A. 48% Fgr 1). (Figure (4.87%) , Agave n w ftegnr r lonw( new also are genera the of two and , siaeifoecne ihfoesarne nrows in arranged flowers with inflorescences (spicate suoobxellipticum Pseudobombax ´ pme arborescens Ipomoea vrzadGonza and lvarez a dniidi aclsmls u ntefur the in but samples, faecal in identified was , .salmiana A. Agave , ahnaungulata Bauhinia itof Diet oln(iue1.I aclsamples, faecal In 1). (Figure pollen lz1970). ´lez Agave .Al1 pce r e records new are species 10 All ). a otfeunl detected frequently most was etncei iai 1755 nivalis Leptonycteris ,Cnovlca ( Convolvulaceae ), oehr 5 ftefaecal the of 75% together, ,adAmaryllidaceae and ), tncru beneckei Stenocereus Stenocereus Ipomoea Ipomoea 6 and and ´lez A. ), Crescentia cujete Crescentia salmiana Agave parryi Agave palmeri Agave neomexicana Agave lecheguilla Agave pce ln aiyCmo aePaeReference Place name Common Family inaequidens Agave horrida Agave havardiana Agave dasylirioides Agave chisosensis Agave Agave plant for Species reported plants food of Species I. Table afeabrachystachya Manfreda Ipomoea 1838 Don, arborescens Ipomoea alata Crescentia pentandra Ceiba Ceiba aesculifolia Ceiba acuminata Ceiba angagigantea Carnegia houstoniana Calliandra Calliandra ungulata Bauhinia schotii Agave scabra Agave il obcca eioGrnr1977 Gardner Mexico – Bombacaceae Mill. .Aayldca eioGrnr1977 Gardner Mexico – Amaryllidaceae L. .Cnovlca eioGrnr1977 Gardner Mexico – Convolvulaceae L. .Bignoniaceae L. et.Fbca eioGrnr1977 Gardner Mexico – Fabaceae Benth. nem,17 mrliaeePrysAae eclypviAioaSeDbtadPietHle1985 Peikert-Holle and Dobat See Arizona yapavai Mezcal Agave, Parry’s Amaryllidaceae 1875 Engelm., nem,17 mrliaeeSht gv,Mge puercoe- Maguey Agave, Schott Amaryllidaceae 1875 Engelm., rea 77Aayldca og gv,Mge eioAioaEsel 1972 Easterla Arizona Cenizo Maguey Agave, Rough Amaryllidaceae 1797 Ortega, e.e aoi 84Aayldca eclelMxc hsstudy This Mexico Mexcalmetl Amaryllidaceae 1864 Jacobi, ex Lem. nem,17 mrliaeePle’ etr ln,Maguey Plant, Century Palmer’s Amaryllidaceae 1875 Engelm., ut,11 11]Bgoica Jı Bignoniaceae [1819] 1818 Kunth, toe amDc,15 mrliaeeA Amaryllidaceae 1859 Salm-Dyck, ex Otto . 73Bgoica Jı Bignoniaceae 1753 L., L)Get. 71BmaaeeCia Purı Ceiba, Bombacaceae 1791 Gaertn., (L.) S asn oe 95BmaaeePcoeMxc e oa n ekr-ol 1985 Peikert-Holle and Dobat See Mexico Pochote Bombacaceae 1905 Rose, Watson) (S. .H ul,13 mrliaeeAaeo auyAioaEsel 1972 Easterla Arizona Maguey or Agave Amaryllidaceae 1939 Mull., H. C. or,15 mrliaeeSidge,Mge ehgil rzn atra1972 Easterla Arizona lechuguilla Maguey Shindagger, Amaryllidaceae 1859 Torr., Knh rte n ae . 86BmaaeePcoeMxc hsstudy This Mexico Pochote Bombacaceae 1896 F., Baker and Britten (Kunth) Eg)Bit n oeCcaeeSgaoMexico, Saguaro Cactaceae Rose and Britt. (Eng.) .Kc,16 mrliaeeMge rt eioTi study This Mexico Bruto Maguey Amaryllidaceae 1860 Koch, K. rl,11 mrliaeeHvr’ etr ln,Chisos Plant, Century Havard’s Amaryllidaceae 1911 Trel., . 73Fbca aad eaoMxc hsstudy This Mexico venado de Pata Fabaceae 1753 L., aoiadC .Bouche D. C. and Jacobi otnadSad.Aayldca e eioCnuyPlant, Century Mexico New Amaryllidaceae Standl. and Wooton Hm.adBnl xWld)G. Willd.) ex Bonpl. and (Humb. Ml. tnl,12 aaeePmoaoMxc hsstudy This Mexico Pambotano Fabaceae 1922 Standl., (Mill.) Cv)Rs mrliaeeDcdosAae–CcrmadHyad16,see 1962, Hayward and Cockrum – Agave Deciduous Amaryllidaceae Rose (Cav.) Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 ,16 mrliaeeDslro gv eioTi study This Mexico Agave Dasylirion Amaryllidaceae 1865 ´, etncei nivalis Leptonycteris ovluaeeMrigGoy aaut eioTi study This Mexico Cazahuate Glory, Morning Convolvulaceae nldn hssuyadpeiul ulse records. published previously and study this including ´ aefe gave rx auymanso, Maguey ´rox, etlalcoyote de Tlacamel Mezcal Agave cr eioSeDbtadPietHle1985 Peikert-Holle and Dobat 1985 See Peikert-Holle and Dobat See Mexico Mexico ćaro ćaro spin – eioSeDbtadPietHle1985 Peikert-Holle and Dobat See Mexico ´ rzn e oa n ekr-ol 1985 Peikert-Holle and Dobat See Arizona rzn oku n awr 92 see 1962, Hayward and Cockrum Arizona eioGrnr1977 Gardner Mexico eioTi study This Mexico ea ua 1989 Kuban Texas USA ede l 1985 al. et Reid –

ety15,seDobat see 1955, Beatty n ekr-ol 1985 Peikert-Holle and esla ta.2006 al. et Geiselman esla ta.2006 al. et Geiselman

1756 nhz&R .Medellı A. R. & ´nchez Sa R. ´n avca eioGrnr1977 Gardner 1977 Gardner 2006 al. et Geiselman 1977 See Gardner 2006 al. et 1977 Geiselman Gardner See Mexico 2006 Mexico 2006 al. al. et et Geiselman Geiselman See 2006 See 2006 al. al. – et et Geiselman Geiselman See See Mexico 1977 Gardner – Mexico – – – – – – – – Mexico – – Malvaceae – – Liliaceae – – Leguminosae Lamiaceae – Convolvulaceae Compositae Caesalpiniaceae Boraginaceae Bignoniaceae Asteraceae Apocynaceae Reference Place – – – – name Common – – – – – – Family – thurberi Stenocereus stellatus Stenocereus 1961 beneckei Stenocereus ellipticum Pseudobombax Pseudobombax 1897 chrysacanthus Pilosocereus weberi Pachycereus tetezo Neobuxbaumia mezcalensis 1952 Neobuxbaumia Dawson, Y. E. Schum.) macrocephala Neobuxbaumia Myrtillocactus plant Species Continued. I. Table osl atca eioGrnr1977 Gardner Mexico – Cactaceae Console uadBmaaee–Mxc ade 1977 Gardner Mexico – Bombacaceae Dugand J .Cut)Bce. 90CcaeeCardo Cactaceae 1960 Backeb., Coult.) M. (J. Egl. ub,16 atca ra ieCcu,Pty uc S e esla ta.20,Dbtand Dobat 2006, al. et Geiselman See USA Dulce Pitaya Cactus, Pipe Organ Cactaceae 1961 Buxb., (Engelm.) Pef. icb atca iaa–SeGiemne l 2006 al. et Geiselman See – Pitaya Cactaceae Riccob. (Pfeiff.) Ern. .Bre n Buxb., and Berger A. (Ehrenb.) J .Cut)Bce. 98CcaeeTtco eioVlet-aute l 1997a al. et Valiente-Banuet Mexico Tetechos Cactaceae 1938 Backeb., Coult.) M. (J. Knh uad 93BmaaeeCaeln,PcoeMxc hsstudy This Mexico Pochote Clavellina, Bombacaceae 1943 Dugand, (Kunth) .A .Wbre Schum., ex Weber C. A. F.

rv,13 atca ounrcciMxc aineBne ta.1997a al. et Valiente-Banuet Mexico cacti Columnar Cactaceae Downloaded By: 1938 Bravo, [Sánchez, Ragde] At: 22:02 30 August 2007 F .C ee xK. ex Weber C. A. (F. atca atsMxc hsstudy This Mexico Cactus Cactaceae Golden- Man, Old Golden Cactaceae O Cactaceae pnd atsViejita, Cactus spined, nEpns,Cnear eioVlet-aute l 1997a al. et Valiente-Banuet Mexico Candelabro Espinoso, ´n ´ gnsggne eioVlet-aute l 1997a al. et Valiente-Banuet Mexico gigantes rganos eioVlet-aute l 1997a al. et Valiente-Banuet Mexico

ekr-ol 1985 Peikert-Holle

itof Diet etncei iai 1757 nivalis Leptonycteris Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 ro oAiaadHmhe 18)ms ecniee aeul.Tkn noaccount into Taking carefully. considered be must (1988) Humphrey and Arita to prior curasoae osiue 98ad1.8.Feunyo pct n aiuaeseiswere species paniculate peduncles) and lateral spicate long on of clusters Frequency umbellate 19.38%. ( large different and significantly in 39.8 flowers constituted with inflorescences, inaequidens Species Tepoztlan. Diablo, del Cueva the of samples fur and houstoniana faeces Calliandra in a, (%) I. frequencies code: species Plant 1. Figure 1758 h te eia pce ftegenus, for the information of reporting of species habits studies Mexican dietary the other about the information little is There Discussion Calliandra obndi .sp,bcuei a mosbet ifrnit nfrsamples. fur in differentiate to impossible was it because spp., A. in combined oebr(iue2,wihcrepne omnh ihtehgetfo species was food diet the highest and frequencies the in with decrease a months showed by species to dominated many corresponded December, In which diversity. 2), (Figure November ( months all types, in frequencies high particularly rqece fmn olntpswr ih u nfee,only faeces, in but high, were types pollen many mean of not does frequencies faeces the other in insects. and them the scales finding but ate leave fur; were bat insects bat scales the to Flower-visiting stick that Butterfly analysis. easily the identification. which them, in allow on included to remains not chewed but finely found, too also were parts Insect samples. P eae 1 n 3 epciey.Teeoe o oprsno h rqec of frequency randomly no were samples, the fur males the of from In species: males. samples plant comparison and for 13 females found of a and numbers were comparable 16 for differences have sexes, to Therefore, order between in items selected, respectively). food 13, of occurrence and (16 females and . h bevdmnhyfeunisdfee ewe apetps(iue2:o u,the fur, on 2): (Figure types sample between differed frequencies monthly observed The fur in 34.14% and samples faeces the of 28.57% in found were parts exoskeleton Insect oesmlswr bandfo ae 17fo eae 5fo acs hnfrom than faeces) from 85 pelage, from (107 males from obtained were samples More .5.I acs ape eeaudn nuhol for only enough abundant were samples faeces, In 0.05). Agave ahnaungulata Bauhinia AiaadHmhe 98.Bcueo hscnuin nomto generated information confusion, this of Because 1988). Humphrey and (Arita pme arborescens Ipomoea .Sa R. ( ( x x and 2 2 5 5 nhz&R .Medellı A. R. & ńchez pme arborescens Ipomoea 0.0476, 0.45, ;B.u, .salmiana A. P x ahnaungulata Bauhinia . 2 .spp., A. ; P 5 .5;n ifrne eefound. were differences no 0.05); . hwdlwfeunisadwsol on nOtbrand October in found only was and frequencies low showed 22.9, 0.05), pnclt pce hrceie yoe paniculate open by characterized species (paniculate Agave P . , Bauhinia .nivalis L. 0.001). ń ,S.b, p. .a, C. spp.; Ipomoea tncru beneckei Stenocereus .curasoae L. ( . x eeatal ae nosrain on observations on based actually were 2 0 nmnhysmls.I ohsample both In samples). monthly in 80% 5 eb aesculifolia Ceiba 0.0555, ( x 2 5 .nivalis L. 0.0370, Mle,10) paety many Apparently, 1900). (Miller, P rqec ftefu gv pce are species agave four the of Frequency . . ;P.e, Ipomoea .5 riscs( insects or 0.05) P uhmr skonfor known is more Much . . suoobxellipticum Pseudobombax 0.05), pme arborescens Ipomoea ( x 2 5 Agave 0.166, x , 2 Ceiba 5 P 1.5625, . ;C.h, 0.05) and had L. Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 oeoVle ta.(04 on httesaoa rsneo ooyo nestimated an of colony a of presence seasonal example, the For that 2004). al. found et (2004) Moreno-Valdez of al. 2004; et species Fleming Moreno-Valdez 2004; has both al. behaviour et feeding for (Bogan Opportunistic distributions documented cave. Tepoztlan well the inhabits been bat this while resources, oee,a pce ee,orsuyad 0nwfo lnst h nw ito this of diet known the to plants of food new diet 10 the adds of study bat. part our endangered be level, rare, species to a reported at previously I). However, (Table been families had 10 paper and nivalis genera, this 13 in species, report 21 of consist diet its in reported plants nytepbihdrprsta r nw orfrto refer to known are that reports published the only Tepoztlan. Diablo, del Cueva of samples faeces and fur on species plant a, of I. (%) code: Species frequencies Monthly 2. Figure h, aladahoustoniana Calliandra h itof diet The ( genera two of exception the With (A ´ vrzadGonza and lvarez pme arborescens Ipomoea .nivalis L. ;B.u, ahnaungulata Bauhinia rbbytak h ptotmoa viaiiyo oa floral local of availability spatio-temporal the tracks probably lz17;Grnr17;DbtadPietHle1985). Peikert-Holle and Dobat 1977; Gardner 1970; ´lez .spp., A. ; Agave Stenocereus ,S.b, p. .a, C. spp.; tncru beneckei Stenocereus Leptonycteris and eb aesculifolia Ceiba .nivalis L. itof Diet Bauhinia ntenrhr at ftheir of parts northern the in etncei iai 1759 nivalis Leptonycteris . ihcrany h ubrof number the certainty, with ,alpatgnr htwe that genera plant all ), ;P.e, suoobxellipticum Pseudobombax Leptonycteris ;C. Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 eonzd(lmn 04,sc eainhpwsntpeiul nw for known previously not was relationship a such 2004), (Fleming recognized eaoimpat,like plants, metabolism of of regions diet forest deciduous the tropical the (Ramı throughout occurs Mexico and range Mountain Tepoztlan flowering Ipomoea nearby of availability 4000 ( hs lnsavr trciersuc o h as utemr,i h aeof case the in Furthermore, bats. the for resource makes attractive range Mountain very Tepoztlan the dasylirioides a in also agaves plants may spicate this of these that abundance that high possible The species is Diablo. agave it del spicate needed, insects the is by with research pollination happened further diurnal have Although favour 1996). that al. characteristics et of flower (Fleming arrival developed have asynchronous of change may and blooming this plants unpredictable to zones, the relation shift temperate by in In evolutionary explained bats 1977). be recent could Schaffer a entomophily and experienced toward (Schaffer and pollination pollinated insect bat towards originally were characteristics 1979). Hevly 1977; Schaffer and Schaffer 1962; Hayward itiuin hs asaefeigo C on feeding are Te bats these 1993; distribution, al. et (Fleming oten(A eaoimpat,like plants, metabolism (CAM northern ie cafradShfe 97 oeoVle ta.20) principally 2004), al. et Moreno-Valdez 1977; Schaffer and Schaffer (i.e. erto Hwl 97 etas18;Toa 98 err n Martı and Herrera 1988; nectar and Thomas flower open 1982; of timing Heithaus the as and well 1977; time, as feeding flowers (Howell of the pollination length by secretion body, produced the animal’s grains the pollen on of increases deposition amount efficiency of turn position the feeding in time, transit maximizing which by 1983) bats Eguiarte species. both attract 1977; for probability might Schaffer that and (Schaffer patches Garcı flower-density (A. high clusters dense forming such bats of diet the in pollen other agave However, spicate agaves. of spicate presence by as the dominated documented be also would have winter authors the during Mexico Central hel ybt,wiesiaeaae seiswt nrnhdsae n ihflowers with and scapes unbranched like with shoot, main (species the agaves on spicate directly while forming bats, like by clusters, chiefly umbellate pollen large only in Cave), flowers Diablo El two of to N August) km 2 (670.23 and Zacatecas July of (17 state visits of two the during in obtained colonies samples maternity fur 41 and samples faecal cafr17;Feige l 93.Tu,w i o xett idple fspicate of pollen find to and expect not (Schaffer did mechanisms we from Thus, entomophilic samples 1993). our of al. in et predominance agaves Fleming a 1977; suggest Schaffer that syndromes 1760 98.Teefcoscudcnrbt otedfeecsi ln rqece ewe the between frequencies plant in differences the to contribute could factors These 1998). Agave h onn lr reo aaut ( cazahuate or tree Glory Morning The thsbe ugse htsiaeseisof species spicate that suggested been has It h essec fple risi u n acsi nlecdb ayfcos gut factors: many by influenced is faeces and fur in grains pollen of persistence The eea uhr aesgetdta aiuaeaae seiswt rnhdsae and scapes branched with (species agaves paniculate that suggested have authors Several .curasoae L. Agave .nivalis L. .Tedfeec sta ntenorth the in that is difference The ). sfo eore for resources food as a detected. was .Sa R. hs lnsfeunl rwnx opnclt gvslike agaves paniculate to next grow frequently plants these , .nivalis L. and rz14) ti hrfr o upiigta hsaudn ln dominates plant abundant this that surprising not therefore is It 1944). ´rez nhz&R .Medellı A. R. & ńchez nnrhr eiofo pi oOtbrwshgl orltdwith correlated highly was October to April from Mexico northern in heoytrsmexicana Choeronycteris nalmnh.Tepooe odsitof shift food proposed The months. all in Ipomoea le 01.Orrslsso hti h otenpr fthe of part southern the in that show results Our 2001). ´llez etncei nivalis Leptonycteris .curasoae L. ,hsbe eosrtdb sn abniooeanalysis isotope carbon using by demonstrated been has ), Agave Agave a esnlcmuiain.Teecutr constitute clusters These communication). personal á, ń AiaadHmhe 98 luo 00.Thus, 2000). Slauson 1988; Humphrey and (Arita lhuhteiprac fBmaaeeand Bombacaceae of importance the Although . 3 Agave gv salmiana Agave lns(mostly plants Tcui 84 nsuhr S Ccrmand (Cockrum USA southern in 1844) (Tschudi, ntesuhr ra fisdsrbto a been has distribution its of areas southern the in .dasylirioides A. pme arborescens) Ipomoea .nivalis L. Agave n atca)adsuhcnr oss(C roosts south-centre and Cactaceae) and uhls httede of diet the that less much , etncei nivalis Leptonycteris paetyol ed nCMplants CAM on feeds only apparently htdslycranchiropterophilous certain display that Ipomoea and and .inaequidens A. .horrida A. sadmnn ln nthe in plant dominant a is ,adas nCMplants CAM on also and ), Leptonycteris ed nna Cueva near on feeds hwpollination show ) gv inaequidens Agave r pollinated are ) Agave nzdlRı del ńez .nivalis L. ewe its between .nivalis. L. .In24 A. in ó 3 , Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 neist h lrsi rvne fSer ar cietladSer ar e Sur, del Madre Sierra and Occidental Madre Sierra (Garcı of provinces respectively floristic the to endemics htpoeto falkonrot n rtcino oaighbttaetemost the are habitat foraging of of recovery protection the and for suggested was considered roosts it be reason, known to this For all aspects foraging species. important of the of to modification protection threats and greatest destruction that the recognizes as 1994) habitat roosting Service of and Wildlife population and the Fish (Medellı in activities (US human declines to related severe probably documented are have studies many nivalis 1970s the Since 1988). Nagy physiological and the maternity (Kunz implications and sexes Conservation a the stages forming between earliest not vary its not but in may mating time only were that was at bats development requirements feeding Tepoztlan, embryo their in Thus, in stay results. colony. bats different their capturing somewhat then During yield case, Diablo. might before the roost, just indeed eaten their was is in and this to cave, opposed If capture the as around roost. the be area grounds, the might the before faeces to in faeces the immediately abundant returned the in most bats foraging pollen is that of that food of type possible plant detected high period food is frequently the the most short it the rather Given 2003) Perhaps faeces 1988). and a Winter scales. (Thomas fur and only time Both (Helversen roost. reflect different the bats examined over to nectar-feeding back probably in heading but before intake had information, groomed, bats diet cave the the meal provide to last the return to bats related When night. the no of almost beginning be the of would flowers at there the them example, visit be For bats should bats. the of but diet simultaneously the used beneckei on be data Stenocereus should collecting when methods separately both considered Therefore types. sample two ealdifraino hsseis iti h otenprso h ag rdrn the during or range the of parts for southern Plan data, the Recovery our in to the diet According of season. species’ mating points this major on information the detailed of one nivalis addresses Leptonycteris research Our 1994). Service nueaeut upiso odt asa ela h ucsflpliaino plants. we of this, pollination Mexico. doing successful Central ( By the tree as both. Glory well protect Morning as to bats need to food we of bats supplies and adequate plants ensure these between of relationships are bat this of diet Federacio the la de in Oficial found (Diario species we plant Mexico. species in plant concern Some species. conservation this 2004). that of (Fleming area recovery foraging production of and purpose types the vegetation to for the us preserved of allows be relevance information the to This about need 2004). perspective al. wider et a Moreno-Valdez of gain 2004; species (Fleming several species distribution this the of for north the in and year of plant food important u itdt i o eelsxa ifrne ntede of diet the in differences sexual reveal not did data diet Our be might fur and faeces the in types pollen certain of dominance the in Differences ute tde hudeaut h oeo asi h olnto fpat uha the as such plants of pollination the in bats of role the evaluate should studies Further on rely to seem plants of species Many Esel 92.Tecue fti eraehv o encmltl dniid but identified, completely been not have decrease this of causes The 1972). (Easterla aMnoaadGalva and á-Mendoza pneryi h vnn,a prxmtl 83 Gnr 92,so 1982), (Gentry h 18:30 approximately at evening, the in early open U ihadWllf evc 94.Bfr u td,teewsno was there study, our Before 1994). Service Wildlife and Fish (US pme arborescens Ipomoea .nivalis L. .beneckei S. erti rtclbedn iei eio tohrtmsof times other At Mexico. in site breeding critical this near gv dasylirioides Agave olngan ntefur. the in grains pollen pme arborescens Ipomoea ,a ela h pct n aiuaeaae in agaves paniculate and spicate the as well as ), n19) eas fteptnilsymbiotic potential the of Because 1995). ń .nivalis L. n20) while 2002), ń n20) h eoeypa o hsspecies this for plan recovery The 2003). ń n te etrfeigbt o fruit for bats nectar-feeding other and slse satraee n endemic and threatened a as listed is itof Diet Agave .nivalis L. MrigGoyte)i h most the is tree) Glory (Morning .inaequidens A. etncei iai 1761 nivalis Leptonycteris r h ansuc ffood of source main the are .nivalis L. U ihadWildlife and Fish (US and nCeadel Cueva in .horrida A. are L. Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 n awr o epu omns hsi otiuinN.1 fteWllf Trust Wildlife the References of 15 No. Contribution Bogan thank is We A.C. This field. the BIOCONCIENCIA, comments. in and help helpful Alliance for for on Rubio comments Ricardo Hayward and for and Caballero reviewers Luis two and and drafts; Rodriguez-Herrera, earlier Bernal Tschapka, Marco Arroyo, olnietfcto,adfrfclttn h nrnet h ayooia Collection Palynological Herbarium the MEXU in to facilities Martı the entrance Biologı all Enrique for de the German INE, work; (Instituto Teresa facilitating this UNAM); Service, Geologia, for of de (Instituto Wildlife and support the identification, and Conservacio for pollen Fish la Museum, Desert para US Sonora Mexicano NFWF, Fondo Foundation, SEMARNAT, Packard CONACYT, oa A hri ,CynP,WieD nan ,AtnahJ.20.Suiso ognsdbats Long-nosed of Studies 2004. JS. Altenbach A, Engalnd D, Weise PM, Cryan R, Sherwin MA, Bogan aalr A 04 bevcoe or acnut erdciade reproductiva conducta la editors. NP, sobre Rowe TP, Observaciones Jones In: 2004. coal. and LA. lignite peat, Caballero from extraction Palynomorph 1999. J. Pross AA, Bruch Thesis, – Science of Master (Saussure). nivalis Leptonycteris Bat Lognose the of Autecology 1955. LD. Beatty Ramı E, Peters E, Ezcurra S, Arizaga Revisio 1988. SR. Humphrey HT, Arita A Conservacio la para Programa thank We Acknowledgements 1762 iroOiild aFederacio la de Oficial Diario Federacio 71:39–44. la History de Natural [online]. bats. Oficial species Hummingbird threatened 1962. Diario of BJ. list Hayward red EL, IUCN Cockrum 2006 In: nivalis. Leptonycteris 2006. Group. Specialist Chiroptera atraD 92 ttsof Status 1972. Blu D. 1985. Easterla T. Peikert-Holle K, Dobat rta .16.Ple opooyadpattxnm:agoprs e ok anrPublishing Hafner Nun GP, Nabhan TH, Fleming In: York: Bats. 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Company. de flores las a visitantes 17:287–292. Naturalist silvestres—categorı ..[israin.Mxc:UiesddNcoa Auto Nacional Universidad Mexico: [dissertation]. D.F. Leptonycteris + 1.Ui.Arizona. Univ. 51S. .Sa R. n exclusio ń, lzL 90 Ana 1970. L. ´lez e A aSler ,SenegL 93 esnlcagsi h it fmgatnectarivorous migrant of diets the in changes Seasonal 1993. L. Sternberg L, Silveira da RA, ˜ez nhz&R .Medellı A. R. & ńchez nNwMxc asrc] 4hAna ot mrcnSmoimo a eerh 2004 Research; Bat on Symposium American North Annual 34th [abstract]. Mexico New in ) ´xico. nzdlRı del ńez nocmi—it eepce nreg.SgnaSeccio Segunda riesgo. en especies de cambio—lista o ń a ereg.Secretarı riesgo. de ás a NM;Cro Nun Carlos UNAM); á, afoa de floral á etncei nivalis Leptonycteris n 02 om fca eiaaNM09EO-01 Proteccio NOM-059-ECOL-2001, Mexicana Oficial Norma 2002. ń. xc efoayfuasilvestres—categorı fauna y flora de ´xico oC rt .18.E ne El 1987. H. 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Waldemar Verlag Main: am Frankfurt üse. ca lplncm euss lpplecolo papel el recursos: como polen el y ćtar e o epwt emnn lds Joaquı slides; permanent with help for ˜ez a Mexico. á, nm eMe de ńoma Cv)Rs ne ergld a A San de Pedregal el en Rose (Cav.) src emurcie de ´strico nd aNtrlz,adArizona and Naturaleza, la de ń lgsmgeeo e ge del magueyeros ´lagos lgsMxcns(CM,BCI, (PCMM), Mexicanos ´lagos a ereg seiiainspr su para especificaciones y riesgo de ás ´xico. n Secretarı ń. lgsGospaia eMe de Glossophaginae ´lagos etncei nivalis Leptonycteris ad ei miney Ambiente Medio de á xc efoayfauna y flora de ´xico nzfrhl with help for ńez gv macroacantha Agave ńero nambiental— ń ´ glMe ngel (Chiroptera: Leptonycteris gc elos de ´gico nm del ńoma ´xico. ´xico, ń Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 etasE.18.Ceouinbtenbt n lns n uzT dtr clg fbt.NwYork: pollinators. New Martı and LG, plants bats. 1988. 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Morelos ń, ´fica. nr lsohg (Mammalia: Glossophaga ńero irncei brachyotis Micronycteris nm eMe de ńoma xc.Bltnd la de Boletin ´xico. a3:1–186. á xc,Isiuode Instituto ´xico, a ei Zoologı Serie á, MxcnLong- (Mexican ńero Leptonycteris (Dobson) Bauhinia ´lagos. re2 ´rie á Downloaded By: [Sánchez, Ragde] At: 22:02 30 August 2007 aineBne ,Rojas-Martı A, Valiente-Banuet Recuperacio de Rojas-Martı Plan A, Valiente-Banuet 1994. Service. Wildlife and Fish u US Untersuchuungen 1844–45. JJ. Tschudi bat the methods of behavioral Reproduction and Ecological 2005. editor. M. T, Kunz Tschapka In: bats. plant-visiting of diets of Analysis 1988. DW. Thomas luo A 00 olnto ilg ftociotrpiosaae nAioa mrcnJunlo Botany of Journal world. American the Arizona. of in agaves species Te chiropterophilous Mammal two editors. of DM, biology Pollination Reeder 2000. DE, LA. Wilson Slauson In: Chiroptera. four Order in production 2005. nectar and NB. Pollen Simmons I. Agavaceae: of biology reproductive The 1977. MV. 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Mexicana ´gica nd o murcie los de ń nzA rzed C Da MC, Arizmendi A, ńez nzA aa ,AimniM,Da MC, Arizmendi A, Casas A, ńez and ebxamamacrocephala Neobuxbaumia nm eMe de ńoma brdeFuaPraa at –.S aln(wteln) 6 p. 262 (Switzerland), Gallen St 1–6. Parts Peruana. Fauna die ¨ber lgshccds( ´lagos-hocicudos ń lsohg commissarisi Glossophaga ´xico. vl .19a olnto ilg ftoclma cacti columnar two of biology Pollination 1997a. P. ´vila ndlMurcie del ń Leptonycteris nteThaa aly eta Mexico. central Valley, Tehuacan the in ) lgsdlnrsed Me de noreste del ´lagos vl .19b olnto ilg ftwo of biology Pollination 1997b. P. ´vila Pylsoia:Gospaia)i the in Glossophaginae) (Phyllostomidae: ne tro el en ) lg auyr ( magueyero ´lago xc.Junlo rdEnvironment Arid of Journal ´xico. pc eiao[dissertation]. mexicano ´pico xc,cnualsade lista una con ´xico, etncei nivalis Leptonycteris ).