platalina genovensium natural history and ecology

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NATURAL HISTORY OF THE LONG-SNOUTED BAT, Platalina

genovensium (PHYLLOSTOMIDAE: GLOSSOPHAGINAE) IN

SOUTHWESTERN PERU

CATHERINE T. SAHLEyt, Department of BiolOgy, Universityof

Miami, Coral Gables, Florida 33124, USA

lUIS E. BARAYBAR, Museo de Historia Natural, Universidad Nacional de San Agustrn. Arequipa. Peru

ABSTRACTFteld nh",,,'atoon<, fecal rellet " n . I ~ , j , aoo 3" c.roon i,,,t<>pe ,ai,,,.

of toe-muscle tissue Indicate<.! thai lonB·,nouted bats, Pia/alma

genovellsium.depend primarily on the columnar cactul. Webtroaut-

r o « < ' e ~ s , , · t b t r b a u e r l . f o r f Q r > d . 1 3 2 5 0 0 m S l l e i n A r e q u i p a . P = In

laiC 1991 weeslimalcdalolalof87batsalroostslleSQCCUrnngw'lhln

a20km " , " r d r a d i ~ s In 1 ' l 9 2 , d u r ; n g " s e " " r e d r o t l g h l c a u ~ e d b ) ' " nEIN"lot\,enl.caclu, flowtrand fruit producliondtcre•• e < ; l . ~ d i d bat

numbers In late 1?93. aflc' usual raIns resumed. nower nnd fnllt

pmJuct,,,,, i " c , ~ " , - . e J , but bat " u " , b ~ " ",,,re still /iO'/o lower thnn '"

\991 Long-snouted bats may ba"e efTIIsrated from Ar"'1uipa in

response to decreased frull and t ~ o w c r productIOn because of drought

co",hl;on, a,wcmt.u w,th .n 101 N , ~ o u••"t fly ahering CllC1U'

phenology Aod bal abundance, Ei l'"no events may havesif!,,,,ficanl

mTlucnce On lhc ceology of[)at-planc mulualisms in southwestern Pcru

~ l a b i l a l deslructton Aoo harvesting of bals for folkloric medLclnal

pUl'f)Osesmaynegall\",ly,mpactbatpoplllal,ons,nndpocrs'SlerlCcofbal

populalions ,n Arequipa may b<: lhfcalcncd.

KEY WOUlJS· Andcs, bals.Caclaccac, CAM planls, 101 N , ~ o , r.cctar

f'",d"<:l,on c ~ ' r y i " g "'paclly. P ~ ' f u , I'/"/"I",,,geno'

RESUMENObservac,ones de campo. ~ n ~ l i s ' > de mlleSl1""4S f c c a l e ~ y la proporc,6n

del 's;\IOpO C J r b o ~ o I i ' en mueStt,S de l e J i d o ~ mus<:ulures. , n d ' c a r o ~que Plll/aI",,, g e n o ~ O l S l u " , depende pnnc,palmenle de la caclkea

colonmaf Weberbtmemcenu,''''''hrrhtJJien, pam ,,, a l , n , ~ " l a < : i n n e n nn,'IIO de eSlud,,, a los 2S00 msnm en AreqUlpa, Peru II finales de

1991 est,mamo, que el nomero de m u r c i / : I ~ g o s denIm de nuestro roldio

<::e cslod,o de 20 km era de 87 ,nd""d'JOS, EOn 1992,dul'1lnteuna

sequiaca",:ldaf'O,el Fen6me"udel'l N.liu,hubu"nadi,minucionen

la p'oducclon de l1t>res y Frutos en la. caculcea•. y el numem de

murclclagos d,s",mU}o. II finales de 1993. de>!,o/:s de que Se

n Q r m n l ' l : l I r o ~ l o s p a t r o n e s " l u , , " l c s , l a p r o d o c c 1 6 n d c t l o r e s y f r u t o . s er e e ' l ~ b l e e i 6 pcro nuestro, C e ~ S O : ; reve!.r""qoeel numem demurci",lallus era 60% menuren c u m p a r ~ c \ 6 n COn eI n",mern en \991 Q U l ~ , ~ SI'. g e " " ~ e n s , u , , , em'gr6de ArequtpaOO'''Q raspocsl• • Ia d,smmoc.6n

deFlUlosyl1orcscuusadaporlaseond,cionesdcseqo,adlirulllcei

Thelong.snOlllcd bal (l'hylJoslomidac, Glossupltaginac)

is a rore n ~ c ! a r - f e e d i n g bm endctnic 10 lhe arid regions of

\\cs!etll I'cru. This monotypie genus was r i r ~ l deserihcd by

Thomas (1928) rrom fI single individuul collected in the

~ ' ' ' < ; I ' ' ' ' O o , p a , . t a l ' ' ' e < ' ' ~ O o , y c ' ' ' ' . e f v a ' ' ' ' ' ' d . l a~ ' l u r a 1 < > a , Apa""",, ~ ~ 8 , "'<qUII", l'em (Td.fa, 51,H_.(241J2)

\0,

(""ome"o de 1':1 Ni,'" Cunsiucrom.,,; que Ius camb'OS de fenolngia de

W. web",lu"'eri y la eonsecuente disminuci6n en el n i J r n ~ r o de

mcrcielngos c''''lados por eI fenomeno del N , ~ o pueden lener una

inn""nciaslg"ifiClllivaen In ccologiad. mutualismo,enllemurc,,,_

hg m y plantas ""elsuroeste del Peru Deslrucei6nde h:lbilaly la

exull<:c,6ndemurc,/:laaos confinesdcuoomed,cinalpucdenlef'eruna

inl1ueneianegallvasobre bpoblacKln. por I"que In persnlerlCta dc

poblac,oncsdemurcielagosmArcqu,papuedeseramenazada

PALABRAS CLAVE Andes, CaCtICClle, capacidad de carg».. EI

N , ~ o , mUr<:'dcgos, Peru. plMtas CAM, PlaJl>-

b n a g e n o ~ ~ n s l l < l > ' , produccion de ~ " " t a r

RESUMOO b s e r v a ~ I X : S de campo, amillse de amOSlras feca" e a propof\:.k> de

carvdo Il " em amOSlras de lecldos mWlCulafe< de los digLlos, ",d,cam

que Pluralma 8"novl''''1U1n uepend(' p r i m a r i o m ~ n l e de caetos e o l u ~ a r e s ,Wt/)<Jrbauero<:l'I'VJU$ wtbubOl,erl, para a sua a h m e n l ~ 1 0 . em un s!t,o

en AreqUIpa, Peru No f i ~ a l de 1'191, eSllmamos quc 0 niJmere de

morcegos dcnlro no nosso rai" de e,md" de 20 km era de 87 in-

d"·ld",,s.Em 1991,dufanleuma:;o:cacauSlldapclofen6menol::l,",no,

hou;e uma d , m i n u , ~ ~ o du p r o d u ~ ~ o de 110r'" e de trulas naj cact.ceaseo numero do morcegosd,mmu,u. No rnal dc 1993. dcpo"que sc

normahzaram oS poome. pluv,alS e a p r o d \ l ~ . k > de nOf"eS e frutas Se

rcstam,leccu, os tlOS5O' c... ' w '"vclarAm que" n"mero de mOfcellosera 600.. men", que em c o r n ~ f l l c ) o ao ",:'mero que er.eontrnrno, em

1901 Propornos que a' mudancns de fctlologia c a conseqUentc

d i m j n u l ~ d o do numeID de "''''cego.< caoSldos pel" f " n ~ m c n o T'I N , ~ "podem ler "ma mfloenci" sillnift.:al,va 11K e, ...logia de mulualo.",,,.

cntremorcegoseplanlasno,uooestedoPeru Deslrutcdodeh3bl1.te

3 h l l a ~ i i o de morcegos Com lins de uso mcdlCmal podem ter uma

m f 1 u c n c i a n c g a l l v a s o b r e u p o p u l ~ d o . e a p c r s i s t t r l C i a d e p o p u l " ~ i j e ,de m o r c e g o s < : m A , , ~ u ' j ' l a p O < l e ,cr.m""cad.

PALAVRAS-CIIAVE Ar.des, Cacloc<:ae, EI Niflo, morcegos. Peru,

plantu CAM, " ' a : o l " , , , g t l l o . ' t n s , ~ m _ produ,c ~ de neL1ar

VIDA SILVESTRE NEOTROPICAL 5(2):101·109

deparlmenl of Lima. Collcctiun Im:alitics indude til.:

dcpa'1tnCnlS of Piura. Lima and Arequipa (Aclkn 1965.

Jimcnezalld I'climr 1982.0rti7dclaf'uenle.1951).ns\\cll

as one location castorthc Andcs. in lhellrid vafleysofthc

dep(lrlmcnl ofllu(lnucn (S,unbotll 1936). Recentcnplur(:ol'

long-snoulcdbalsnCarlhceilyofTaena.l'ero(Dcpartmenl



Fillure 1. M ~ d.plctlnll the city ol ......qulp;1l ~ l 1 d surroundlnll

~ r u . L o c ~ t l o n s Of the Ihree con.lslenlly cen.u• •d

roostsit.......hown ...quare.

of Tacna). extends this bw's range to the southemmosttip

of Peru. Collection sites mnge in elevation from near sea

level (Lima) to 2$00 m (Arequipa IIml Huanuco), and all

are known arid habitats.

Nectar-feeding bat species that shaft': part of the long

snouted bat's range include Lom:hophylfo hesperia, Anouru

geoffroyi. and Glossophaga soncma.. LO/'IChophylla hesperiais restricted 10 north""estem Peru; A. geo//ro)"! is not foond

south of the department of Lima west of the Andes; and G.

soricina doe... not occur abmc 140U m (Koopman 1978.

Tunle 1970). Thus. Ihe loog.snouted bat is the onl} neetar

feedingb3t specit;:.>; present at mid-to highclcvlltion sites in

southwestern Pt;:ru lind i.. the only potential bat pollinator

and seed disperser of planL. in this region.

Despite its probable role d!o an important pollinalor ami

seed disperser in arid h a b j t a t ~ lind ils high dei'"«: of

morphological specialization forne\.1ar.feeding(Winkelman

1')71), prior to this study liule was kllO",n about the natural

history, population status, and ecolugical importano.;t;:ofthis

bat.

STUDY AREA AND SPECIES

The stud)' sile is located in the depanment of I\rcquipa.

Peru. 20 km south orlhe cily of r\requipa (16.2rs. 71.]0"

W; Fig. I) on the western slope of the Andes. at ap

proximatel) 2500 m above sea level. At our mllin study

site we ohserved hat visitalion ofcElcti Ilowcrs. Ilnd rccord_

cd Cl1o.;lus phenolog) and dens ity. We recorded addit iona

bat census dald. at various locations within a 20 km radiu

of Arequipa. Within the 1257 I..m' area searched. at leas

314 lm ' is under residential or agricultural use. AIM}

because of heterogeneous IOJ'Ogrdphy, area.. \\iithout cactu

occurred within our search radius. Thus, suitable fnrdging

hahitat for the 10ng-snOl.lted bat was much less than the

actual area searched. Sites that supported bat colonies al

corresponded to the ~ u b t r o p i c a l . low-montane, descn scrub

l ife zone as classified by the Holdridge system (Ol\ERN

1976). In Peru. this l ife zone occur.> from 7"]0' to 1 8 ~ 1 0and can occur in two areas; a long the coast from 500-1000

m. and along the western slopes of the Andes. general!)

between 2000 and 2900 m. Mean annual rainfall for this

life zonc b 222 l1un (ONERN 1976). Ilclwccn 1991 and1993. however, a drought a ~ s ( ) ( ; i a t e d Wilh an EI Nino even

resulted in a substantial decrease in rainfall in Arequipa (se

Fig. 2).

rhree species of columnar cacti occur at the sludy site

Weberbouerocerew.r web<!rbaueri. Browningia cDlldefllis

and c.:oryocactus brevist)'lus. Of these. only Browning;(

and Weberbauerocereu$ ha\'e long. tubular, nocturna

nowers that can be visited by bats. The most abundan

columnar cactus at the study site is W. weberbaueri. and

around Arequipa (Aragon 1982. pen.. 000.). is visited and

pollinated by the long-snouted b:lt. 1""0 species of hummingbirds, Pli/(lgona giga! and Rhodopis vesper. and

possibly diurnal inw!:!.>; (SaMey 1996). This eactus exhibit

noral morphological chunlckristics cnn ..istent with both ba

and hummingbird pollination syndrome!'! of columnar cacti

For example, flowers arc long, narrow, and luhular, and

nower color ranges from white to red. F l o w e ~ open in the

aftemoon. remain open during the night and close late dIe

following morning. "eclar production is continuous whil

the now!':!" is open (Sahley 1995).

METHODSWe searched all known mines <lnd tUllnels within a20

km radius ofArequipa. We osed topogrophk mtips as wel

as inten-iews "ith local people and biologists LU locate sile

and in 1991 we censused live sites. We consistently

monitored thl\.'C roost sitcs from one to five times a}ea

from 1990 until early 1l)l)4. These three colonies included

lhc two siles with the largest bat aggregations (Charean

tunnel. ata hydroelectric fllcility. and lJatolito mine #..,) as

102 LONG-SNOUTED BAT ECOlOGY IN PERU· SahJey and BltfByNr VIOA SILVESTRE NEOTROPICAL 5(2):1996



§ $ §M

§M M

~ a § ill

Date

-+-Rainfa l l ..• - -Mature fruits - . . - Flowers

90 ,

80 '\

70 \

E 60 \

§.. 50 "

40

30 "20 .

10 -_ . . . . ,

o +--+-__._ ._...;-..:-O"-".."'. - . . . . ~ _ ~ . .-+- •

30

~ 2

T20

15

..

\ 10

'.I5

2. FloW'll' and mature fruit production 01 W e b e r b a ~ f o c e r e u s weberbaueri In relation to rainfall, October 1991· July 1994

OISUliC abandum:d min<:lhat huused a:.mllltcr "roup of

(Poderosa mine). We did not capture any b:us from the

odcroSll mine. nor did I\C emer the mine to conduct

Visual counts at the Cilarcani lunnel .md Balolito

3 were made by either entering the mir.es during the day

.... llitinjt outsIde until bats Cl\ilCd in the C\(:nina. Cap

... ere made using rniSlnets at OIl<: (olony ~ i in 1990

alol ilo mine _3) and allwo sites (CtLarcani and Ba!olilO

.3) in 1991-93. In 1990. we rccordo:d relative age.

orearm length. and r c p r o d o c t i ~ e condition of capllKl.:d

dividuals: in 1Q91-93 \l,e also record..-d body mass. For

ree adult males Ilnd three Ilduh females. "e measured and

corded wingspan and wing area 3ccording to l \ ~ r g and

(1987). Wing loading \.asealeuI3tcd separate!} for

ult males llJ1d females by dh'iding mean body \l,ci&ht b)

wing area.

Phenological data for W. weberbaueri were obtained at

he main study site by counting the numbers of buds.

wers. developing fruits. and mllture fruits. for 40 mllture.

individuals belween OctubL'f 1991 and July 1994C3Cli were loc3lcd ",40 km from Ihe Charcani roost.

IS km !rom the BOlolito mine 113 roost. and <I km from

he l'odero5a mine roost. I 'lants wen: u.:nsused weekly ur

onlhly whenever poss ible when only LEI] wag present

lig, 2 tor censuS dales). MClIn 110\\er and fruit

JU(;tioniplantlmonth and per dOl>' ,",cre cnlculalcd from

enol08Y data. The rc!ati,'c density of adult IJ c(RldI'J"is

d II i licfJ.erbOll.cri cacti ntthe m;lin study SllC \\as Jcb> (flunt'ng the number ofcacti >1 m in height in

ten 600 ml quadrals. From this dab we estimated the

number of adult cacti/ha. Nectar produ(tion of IV. weber-

bauui nowers was measured by bagging 29 nowt:n. on 29

randomly selected plants prior 10 opening lUKl measurinG lhe

amount of nectar produced lit 2-hr ir.lervllis unlillhe n o , ~ e r sclosed the following morning. Sucrose concentration was

measured l15ing a Reicnen-Jung rcfraclomelCT. The valu.?S

obt3ined. in mg sugar/mg solution. ,",erc con,'ened to mg

sugar/ml nectar as det;eribl:d in ROllef' ~ d. (1979) Medii

sugar content was calculated per ilowcr:'night and converted

to kilojoules. We used mean number of nowers p r o J u ~per nightlplanllha 10 estimate neaar produc:tionlha.

Estimates of(iII'rying Cllpacit) b:tSed on neclar availability

per heclare for baLS , ~ e r ( obtnined from Nag)"s (19871

equations for field metabolic rates of eutheriWl mammals

and passerine birth. Elootimales for mlllllmais and birds were

bOlh used because bats may have field metabolic I"Jtes

approaching those of birds (Pctit 1995. Petit and Pors 19%.

von Hclvcrscn and Ue)'cr 1984). The equuliuns (lloocd to

(uleuiute energetic requirements \liere: 3) kJrcquired/individual/d J.3Sxeffcctivcmllss.O lll (cuthcrian

mammals): and h) kJ re1:luired/individlllll/d - 888

t : f f L ' C l i ~ e mas{'" (passer ine birds). Effective muss wa.s

dclined as the mean mass of adult. non·rcproductiH:

individuuls {19.7 g),

TlI lIbtain II preliminary assessment ofthcdietllry hllhits

oflong-snOllted bats. in 1992 l I e collected 1111 fecal pellets

in II 0.5 m' plot undernealh an area whcn: II cluster of bats

rcgularlyroost....,J in Batolitllmin.:iG. T h c p c l k : ~ " ...re8ir

SILVESTRE NEOTROPlCAL 5(2) '996 lONG-SNQUTED BAT ECOlOGY IN PERU· ~ h l e r . n d S.reyb., 103



-"......... I"._......_.. ....; . l -P ' . . . . ' •• ,

Subaduhs Atlult females Adult males

Variable 1- SD n x SO n SD n

T.ble 1. M. . . . mOfPhologiellm...u...m.ntsforP.". . .o n"illmelptu,.d betwe.n 1991 .nd t t t3 . Mann.Wh.t ., U

c o m ~ r h l o n offor.. rm length.nd body m u . be_"lIdultm.I. . .ndf.m . . . . .

Cactus pulh:n was Ihe mosl frequ.:nl item found in feca

samples (99%). followed by unidenlitied insect par

(5').11%). seeds andlor frtlit pulp (I7.M/,,). anu atltherpar

(14.7%). Of 102 pellels examined, \\oe found nine \\ho

II'. webllfbaueri seeds. and 6 large picees of W. webcrbat<

ri S<..""Cd h u ~ k s . We found a 1000al of 52 it'" K eberbout:

seeds in the same area where feo.;al pclletS\Verecolleclc

Presumably tht'Se separated from f«al pellets durin

defccativn or upon striling lhe ground.

Diet

The long-snouted bat is a rdalivel) ' large neelar-feeding

bal, third in size 10 I.tpumycreru c u r o s ~ and L. n/l'aJi

(Table I; sec Norberg and Rayner 1981 lOr comparJth

morpholoekal data on glu!>SOphagine species discussed ithis paper). Adult females ha\(: a slightly longer forearm

( U - I . ' i ~ 1 . n-g5. 1'-0.037) than adull males. The long

snouted bat has a l ~ e wingspan and fommn. sa:und to L

niHiis. Wing area of PIOlalno is Ihe largest of an

g1ossophllgine recorded. although wing Brt:ll measurement

fur the largest nectar-feeding bat. L. ni"oIi1. are unavailabl

for comparison. The long-snouted bal has a 10\\ win

loading relative to most other glossophagine hat" lor whic

measurL"lllt'nts are available. especially when compared t

glossophllgines with a similar body mass (Fig. 3).

RESULTS

Morphology

eSlimale significant differences belween adulls of both

sexes.

Flgu ... ). W,ng Ioadloga of glosaopl ' ll l lPn. b.ts. Oata from

NOfberg.ndR.yner(19S1I.ndSlhlly.r . , .( t .U) .

L.LIc:l!onyel.....t. C.·Choer'Ol'lisc:us. G.·GIOlI.toplle".,

A._AIlour•• Ch._Choeronyelen.t. P.·Plele/",•••nd

L.LtoptonTC"';lI. Forpurpo... ofc.omperleon.mea"

wing Ioadiog fo ' P ge<10 ....nll;um W. . ulc.ulated from

m•• " wing m u r ~ m e n l $ lind bod., m .n of both

..... combl d.

19S IJ 4 SOO I i i 52

OlS(, 1

OOlO1 l.. ,

1_9 H 41

:105' 12 "01<1 }

OOIU 1

104 }

dried \Ind Slored in a plastic bag. One hundred and two

p e l 1 e L ~ w ~ r ~ randoml) sel«ted. placed in separate petri

dishes. and reh)'dralcdwilhwatcrpriorloobservlItion with

aslcreomicroscope. PellelSwereleas.,Jupartwithdis<;cct

ine probes. and frequency uf occurrence of pollcn. fruit

pulp. seeds. and insecls per pcllct wasscorcd. Pollen. fruit

pulp and o;eeds found in fecal samples were c o m p ~ r t : d to

reference samples of W. weberbl1U£ri o;ol1eo;led in the field.

Carbon isotope analysis of bat t o e · m u ~ l e tissuc was

performed to delermine this speein' dependence on CAM

cacti for food. I3ceause columnar cacti use crussulacean

acid metabolism (CAM) as a photosyntho:tic adaptation to

arid conditions (Gimoon and Nobel 19f(6).the proponion of

"c ,,1."C (Oil ralio) lhal is fixed into cnctU5 t i ~ u e s \aries

suMtantiall) from that of plantS that undel)!u the morecommon C

Jphotosynthetic path\\ilY. When l I n i m a l ~ leed

from CAM plantS or on hcrbhores thai feed on CAM

plant:., lhe carbon isotopic composition of the CAM plant

t i ~ u e is consened and fixed into animal tisstle. Then:fore.

lInal)'sis of the carbon isotopic tomposition ofanimallissue

can offer imponant insights inlo animal dicts. Muscle

tissue from one toe per individual was obtained from three

live bats captured in January 1'}91 and from thrt:t: ~ e " " ' e dspecimcns in the colll,';l;tiun ufthe Universidad '1ocionnl de

San A g u ~ t l n (no tolhxtion dale was nvuilnblc for these).

Carbon turnovcr in g ~ r b i l muscle tissue has a half-life of

a p p r o x i m l l t e l ~ 21.6 days (rieLsen el al. 1983). thus l l n a l y ~ i ~of Cllrbon slll.blc isotope c ( ) m p O ~ i l i o l l of bal m u ~ c l e tissue

probably integrated isotopic composition ofdicllll')' carbon

ingested duringlhe preceding year. Analysis of tissues Wlll>

performed aco;ordingto methods outlined in Herrera et aJ.

(1993) lind Buchanon ami Corcoran (19.'i9). Tissuc samples

were not collceted in 19'12 through 1994 because of 10\\ bat

numbersStatistical analys is was perfunned using SYSTAT

\ersion .'i.3 (Wilkinson 1991). \ \c conducted Mnnn

Wtlitney L t e S L ~ on foramn lengths and bod)' mass to

M'I ' t J) 11>91)1 nF ~ " " ' I " " I ~ Y l O ~ ) 11

W,_• . , . . , l_ l

W'B....... S( . .")

W ' B . I o U , ~ " C ' ' ' ' ' ' )

1()<l LONG-SNOUTED BAT ECOlOGY IN PERU' &hley and Bal1lybarVIDA SILVESTRE NEOTROPICAL 5(2).19



Bat Populations

Table 2. Census dlla and roost eltOl forlong_snouled batt; und

toestimetetotelpopulatlonsizewilhinourstudyradlua

In lale1991 and early 1992.

Figure 4. a) Census data fo r Poderosa mine. b) Census.nd

se1 composition dala lo r Chareani tunnel. 8a ...

repre..nt number of IndiwidUlls eeptured b s e end

age. DiamondS indicate "isuel counts. Mon ths withonly "isual counts lilCk s e composition data. c)

Census and sex composit;on data for 8atoli tomine 1/3

from 1990·1993.

In 1990. prior to the El ~ i i i o drought, groups containing

males, females, and subadults were present during ever)'

c c n s u ~ period at Batolito mine 10 (only mine ccnsused in

1990). Subsequent censuses of Batolilo mine #3, Charcani

tunnel, and Poderosa mine show¢<! thutlong-snl!uted bilt

abundance declined suhstantially during Ihe onset of Ihe

s e v e r e . ~ t part of thc EI Nitlo drought in April 1992 (Figs.

4a. b, and c). In late 19'H and early 1992 we found a total

of65 bntsduring our censlls ofthcsc three mines, hut in

Junu:uy 1993 we found only two bats. In late 1993 after

the drought cnded. bat numbers increased slightly but did

not approach pre-drought levels. Only 24 individuals were

fuundat Ihe two largest colony sitcs(Charcani and Balolito

1i3) in October 1993. OJ. 60% decrease compared with late

December 1991 and early January' 1992 whcn 60 individu_

!lIs were found. Numbcrs remained low in 1994 relative to

1991. although two rainy . ~ e a s o n s had occurred since the

drought (Fig. 4). No bats were found in lhc I'oderosamine

w

,,W

"

ApJlII990

JanuJlyT9'l2

Oecember 1991

Jnnu:uyI992

Janu.ryl992

De<e,"bcll'l9l

M o r c h l ~

BaloIl10mi",,02

Batnlllomi""iJ

1'<X!c'osam,nc

RC",.lcminc

Coric",nchammc

Charconi lun",,1

BOIOlllo luonel ~ (flooded 1991)

WeobtaincdOIiCratiosof-1O.29.-1O.52.-l0.90.-9,40.

and -10.48 from tissue samples of five long-snouted bats.

A previous study found tissues of CAM eaetusand agave

plants from Mexico to hllVC'O"C values of-12.6 whereas

bat-visited Cjplants hav.: a mean O"C of-26,4 (flt:ming

1'1 ai. 1993). These data, along with our observations,indicated that in Arequipa long-snouted b a l ~ fecd almost

exclusively from cactus flowcr:>. fruits and insccts that fccd

on CAM plants. Estimates of dcnsity indicate that IV

weberbaueri (98 plant5/ha) is much more abundant than B.

candcfaris (4 plants/ha) at our site. Observations of flower

visitalion behavior and of bat droppings collected at day

rooSlS indicated bats feed from flowers and fruits of W

weberbaueri. Phenology dala indicated year-round fruit and

flower production of W. weberbmleri (see below)

Seven roost sites c.on1aining long-snou1ed bats were

loealed Crahle 2). All potential r o o ~ t sites ~ e a r c h e d con

tained bats. Four sites wcre loeatcd in 1990 and three

additional sites in 1991 (Table 2). All occurred between

2200 and 2600 m above sell level. and all were located

within WeberbauerocerclI.S habitat. Five roost sites were

abandoned copper mines and two were man-made tunnels

e:>a.:avaled into ridges. Both tunnels were unused by

humans and had only one opening. Visual counLS of bats

indicated that colon)' sizes ranged from a minimum of one

toamaximumof5Uindividuals; prior to the 1992 drought.

the median colony "i7e was five. Within a roost. groups 01

5-1 bats were found at different roost mierosilcs. In late

1991, a census of live (If scven roost silcs revca1cd the

prcscneeofapproximUlel)' 77 individuals. Due 10 logisticul

constraints in late 1991 to 1994, we did not census the

remaining two sites (Ilatolito mine 112 )' tunnel #1) we had

fuund in 1990. If we assume these two sites contained a

10taloflO individuals(themaximulll numberofbatsfound

there in 1990), wc estimate a total number of87 individuals

presenl in tunnels and mines within a20 km radius of the

cil)' of Arequipa in late 1991.

VIDA SILVESTRE NEOTROPICAL 5(2)1995 LONG-SNOUTED BAT ECOLOGY IN PERU • S 8 h l ~ y and Barllybar 105



Reproduction

D. . .

Figure 5. Age Ind au compolitlon of tlpturad bats II Chlrclni

roost Ind bIlge an d IU compo,ltlon ofclplu,..d

blIt••t8ll toll tomln,/II l .

aller April 1992. The Cr.lIrcani tunnel. . .i t t of the largest

colony that contained SO bats in 1991. contained only three

indi\iduals In \lan:h 1994.

SocIal Structure

Roosts <:onsistcd of onl) males or mixed-sex aggrega

tions, and "e.x composition of the roosts \llned o\er timl:

(Figs. 5a and b). For t . ~ l I I l 1 p l c . 81 the Charcani IUllne! (Fig.

Sa),which contained approximutdy 5 0 j n d j \ ' i d u l l l ~ in 1991,

wecaplurcd only adult and subadult males from 1991 until

latc 1993. bUllhereaflcf also captured l ~ m a l e s . Suhadult

males ocwTTed in all male and mixed-sex colonies. bUI

suooduh females occurred only in mixed sex aggregations

(Figs. Sa and b). We ne\er found all-female colonies and

pregnant ferules ... ere always found in mixed- colonies.

Althoui!heaplure data indicated that owrall the adult male

and female sex ratiO was almost c.xaetly '.1 .....ecaplurro

more subadult males than subadult females.

Aner March 1992. during lhe height of the droughl ....

caught no subaclults or juveniles. We c;:aught no pn:gnan

females from January 1991 through September 1993. Thi

two-ycar span corresponds \\Ith the drought period and 10\

frui: and flower production by W ....eberlxn/eri. In 1993

three out of eight females captured in October \\cre prt:&

nant or post-!;tctating. AoJuh male"'. however. were foundyear-round in Arequipa. although at ..ery lu\\ numbers, i

all yeani from 1990 through 1993 (Fig. 5).

Cactus Phenology and Carrying Capacity Estimate

for Bats

Mean density for adult (>Im) IV wf'berbuuer ; cacti pc:

sampling quadrant was 4.5 cacti (n=IO. 5D"'4.1, 9.5%

C.1.-2.9) which allowed us to cSlimate adult I,;actus densit)at 9.8/ha. Flower dcnsit)· lit lhe main study site droppe

from 8(>lhafd in October 1991 to 9.3/hafd by April 1C)lJ2

Mature fruit production decreased from 19.3/ha1d to1.4/hlld during this same time pc:riud. althoogh nnwCf and

fruit production ne..er stopped completely. After rain

returned in IW3. fruit and nowerproduction in If. w e ~ r b aueri i n c r e ~ d 10 a level compornble to thai observed in

late lQ91 (Fig. 2). Because nowers produced an aVCfllg

2.I73kJ o f n \ - " ( ; l a r ~ r n i g h t , c n e r g ) from nectar availabl

to the bats dccreased 89"10 between Octuber 1991 and Apri

1992, from 186.8 kJlhalnight to 20.2 kJ/hainight. Daily

energy requiromcnts for non-reproductive long-snouted buts

as calculated from NagJ's (1987) equation for cutherian

mammals. indicate a nccdof 37.11

kJ1dper

individualThus. c ~ t i m a t e d can}'ing capacily for bats based on nectaavailability ranged from 4.93 halv'ha in October 1991 to

053 bltslha in April 1992. E ~ 1 . i m < i l e s ubtained from Ih

cqualion for passerine birds indicate bllS ma:- need up 1

82.78 kJ:d. If this were the case. cltrT}ing capacity wouldhave ranced from 2.6 butsIha in October 1991 to 0.24

bals/hain ,\prill992.

DISCUSSION

Morphology

The long-snoutcd hat has a large wing area and wing

"'pan relative to most other glossophuginc bats for which

measurements are available. Since air density dccrcase

with ahitude above sea level, power required for hmerin

and forward flighl also increases wilh altilude (Nurber

1990). 1\ large ....ingspan and low wing loading, as found

in long-snoult'IJ bats. will reduce power required for fligh

at high altitudes and m:w also increase flight m:tn

euvernbilit) (Norberg and Rayner 1987). Wing lood:ng o

the long-snouted hat is Iuwer than thai oftwo other relati..e

106 L O N G ~ S N O l J T E D BAT ECOLOGY IN PERU· Sahley.nd BBrayb.r VIDA SILVESTRE NEOTROPICAL 5(2):199



large glossophagines that inhabit arid UJrlCS. (hoe-

lJ.7 'lim l• Nurb\:fJJ: and Ra)"n\:f 19!17)

l . tpmnl'ctulJ (J-l'awne t12.8 ~ ! m J . Sahley el 0/. 199Jl.If f 01. (1979) found that mean wing disc looding

"ith increasing c1evation fOf hummingbirJs

n ~ on the eastern Andean slopes of Peru. They

ggested that low wing loading found in h u m m i n g h i r d ~ at

h elcvations is a potential lldapl<ltion to reduce c:rlI:rgetic

during 00\ ering and forwartl night. Wingof long-snouted bats may also T q l ~ n t

10 flight in montane desen habitats. alihoogh

on: detailed information on its altitudinal range and

a l comparisons with other glossophagine b ~ l I sith similar altitudinal rongcs arc necessary to tcst this

Our dala indkalC lhallhe long-snouh:d bat is a cactus

pecilliist at our study sitc in sOlllhwcstcm Pcru. Flower

tlltionobservations. !1owera\ailability. analysis of fecalllets and SLlC values indicatc lhat at Arequipa, the long·

outed bat subsists almost entirely on CAM plants and

se"ls thal f\:l:d from CAM plants. Our observiltions and

of fecal pellet! suggest that in our study ~ r e a . thet ~ o . ! bat obLains most of i l nourishment from ) j

Pollinator c\c1usion e ~ ~ r i m e n t s cono.!ucted

W M,tfHrbaui?ri 1I00\ers indicated that the 10nS-snootcd

3t is an important pollinator of this caclus (Silhlc) 1')%).

:though this bat probably visi ts BrowlllnS?io c o n d t / ( T i ~the Other co-occurring nocturnally flowering

mnar cactu.s. this plant occurs at utranel) 10" densit

es and flowers only 2 months of the year whereas IIproducn li t leasl some flowas and fruits )ear

(Fii. 2). ThUs. B, comJcfflis llowl:TS and fruits

obably comprio;e onl) a fmction of thc diet of the long

uted bal. r\o other night-bloominl! plants such as ~ a v e snear lilly ofthc roosts we c c n ~ u ' l i : : d . nor did any

ccur at our main study site.

Despile ils exceptionally clongatcdskull and itshypothe

high degree: of specialization lOr nectar.feeding

1971),thelong-snoutcd bat feeds on insects

fruits as well as nectar and pollt:n. Jhe low b·le

founo.! in bal muscJe tissue lind thc fact that insects

"ilh pollen and anlher parts in fl:Cltl pellets

lhl: long-snouted bal probably f e ~ - d s on insectsund within cnclUsnowers,most Iikel) smallcolcopteTlllls

to the lamil)' Nitidulidae. Other studies ha_e

..'n that nectllr-fecding bats eat insects and cactus fruit

as well as nectar and pollen. fo r ""ample, in ~ o n o r a ,LI!PlorryC/l!r/s curmoOt! feeds from both flowers

f r u i t ~ nf three species of coillmnllr cactus f l. I I

peTS. o:omm.) ami in CUnM;30 also consumes

(f'ctil 19(1). In arid VcnOIlChlO hllbitats, G!os-

sophag%ngiros/r;s consumes cactus f r u i l ~ . inse"ts. flowernectar. and pollen l':iorio.no 1'/ 0/. 1991. Sosa 1991)

Census Data

Tm;le (1970) referred 10 the long-snouted bat lIS ~ e ; ' ( t r e m c:ly r n n : ~ and oor data agree. Long-snouted bat abunJanteis low at Arequipa; collection records for the dCl"1rtmenl

of Arequipa indicate this species has never been collected

at altitudes below 2200 m. even though i s o l a t ~ ' d ,,;actuscommunities e,iq ncar the cnaSl We estimate that prior to

the drought. a lotal of 87 bats w",n: present" ithin a 20

kilometer rndiu\ of ArequiJ'lll. HowevCf. we may ha\'e

m i some mines or small tunnels, becaul>': "C used

t O J l ( ) g r u p h i c m u p s . \ ' i S u a l s e a r c h e ' l a n d i n t c r v i e \ l ~ \ l i t h l o c a lpeople to locate potential colooy sites. Beciluse no naturul

caves occur wilhin our stud) area. haL" may be adapted to

roosting in rock)' crevices or shallow depressions along

ridges. We never observed hats emerging from thesct)'pes

ofrooslS,although we walked transccts near the main study

siteat

SUliset and sunrise, llnd "he"kcd shallow creviceswhen possible for bm p r e ~ e n c c . If long-snouted hilts arc

using crevices as roost sites. then our " e n s u s e ~ may huveunderestimaLed bat numbers. B u t ~ may have shifted thcirroosting habits since the advent of mining in Arcquipa wld

at present primarily usc abandoned mines.

Ine pattern of low bat density we found contrasts \ \i th

that of the Mid-d .. ",lIing LCPIQIJ)'C1CriS CUrasDOe. II highl)·

:IDCiaL rclati\dy ahund3nt, and widesprrnd nectar fec:ding

bat (Aril<l 1993. Cockrum lind Pctfyszyn 199L Howell

1979) and \ \ilh the common GfOSJiophaga fongiroslris and

G. soric:ina.1Arita 1993. Soriuoo el uJ. 1991). Th", l o n ~ snouted bat's h::tbit of roosting in relatively small groupsrna} be most similar to Illat of the arid-dwelling CllOe-

ronyclefis Ittil!xfcala (Arilll 1(93). Unfortunately, little

comparnti_c informalion exists on the abundance 01 other

species of arid-dwelling nectar-feeding bats.

Our data indicate that the long-snouted bat "as present

) car-muno.! when eensu:;ed in 1990, but populations decreas

ed during a ~ e v c r c drought from 1991·1993. Ihis drought

included i I 17·monlh period with no rainfall (rig. 2).

Allhough in 1992 and 1991 Wi: regularly eensused only

thrce of lh", original seven colony sites, we believc our

censuses r c p r C ~ c n l l l . C 1 U a l decreases in bllt p o p u l Q t i o n ~ Qnd

notlocalrOOSlshills.lflocalroostshiflsfrorncensust:dlU

uncensuseo.! sites weTC occulTing, we probably would have

noticed temporal varialion of bat numbers over time ilt

r o o ~ t s WI: c e n ~ u s e d . as \JI.:currcd in 1990 for llalOlilo mine

#3 (Fig. .1). It i unlikely that roost disturbal\i:e caused hyour censuses n : ~ u l l e d in Ihe long-term. persistent decline ofhats we observed, We never eought bots at the Podcro!t8

mine and infrcquentlycensusedandforcaptun:J b3tS at the

othrr two roost sites. Some of our censuses oecurrcJ o\cr

8-9 month intcTvals (t'ig. 31- Finall)'. pollinator exclusion

SllVl:STRE NEOTROP1CA!.5f211996 LONG-SNOUTED BAT ECOLOGY IN PERU' $Mt:ey 81Id S . ~ y H r 107



c ' l ; p a i m e n L ~ nn IV. ll'cberbcueri cacti showed a decrease in

bat-pollinated flowers in 1993 relath,e to 1991 (Sahley

1996). Th;$ decrease in bat pollination " , a ~ statistically

sicnificant. thoueh the sitc '" here e'l;periments took place

was '" ithin 10 lm of all uncmsu$ed rooS1S, \ ell within

foraging range of the lone-snouted bat.

After flo",ering and fruiting of IV wcberbaueri increased

in 1993, long-snouted bat populat;onsremwned 10'" rclath'c

to 1991. We hypothesize three factors rna}'h a ~ e

pl(l)ed animponant role: in the bat population decline o b o ; c r ~ e d , First.

oots mlly h:1\e emigrated from the study area. Data on

rcduced flower and fruit prududiull in 1992, and thus a

sull'itantial decrease in carrying capacity, suggest that

em i ~ r a t i o n from the sllldy site is plausible. Second, drought

and/or migratiun rna} ha\e caused greater monality in 1992

and 1993. and third. oot collecting for medicinal purposes

might ha\-e pla)ed an imporwnt rule in observed declines.

Although we had no c\idence of increased monatity in

long-snouted bat colonies because of drought, it is ptJltsible:

that a combination of 31 three faclon mllY h3\'e been

responsible.

Resultl> of pollinator exclusion experiment" indicate 1 0 \ ~batahu1d:lnce in 1993 caused significantly lo\\er fruit set

from bat-pollinated floweD uf "", wl'berbaueri in 1993

compared with ,q91 (Sllhley 1996). Wc propose that lucal

decreases in long-snouted bat abundance, probably triggered

by droughts that cause reductions in tlowcr and fruit

produetion, may huveconsiderable intluence on the ecology

of this bat.cal:!Us·hurnrningbird mutualism.

Conservation

Thrcats to continued persistcnceoflhe long·snouted bat

around An:quipa includc rOOSI disturbance, colleclion of

bats for folkluric medicinal purposes. and habital destruc

tion. Bats are belie\ 'oo to cun: vdridy ofailments ranging

from e p i l e p s ~ ' to heart attacks. Bats can be obtained from

h e : a l e r s . o r ~ e u r a n d e r o s . R a n d a t m e d i c i n u l s l 3 f l d s " " ' i t h i n l : i t ymmeLlt lnfunnation"e nbtained from informal intel"\'iews

in the central mme! in downtown Arequipa suggests that

se\-eral hundred hats might be l>Uld annually at this market

alone. Oals sold at mcdicinal sumds are purchased from

intennediarics, and markel vendors do not din'erentiah:

among species wilh respect to sell ing price or medicinalu ~ e . In 1992 we .. isited seven medicinal stands al the

central market in Arequipa and found verspertilionid and

molossid bats brought from thc coast and rtIinforest areas,

but no long·snouted bats. In 1995, during a visit to the

same milflet we found 10 l o n g - ~ n o u t e d bats. priccd from

S2AO to S5.00 c;lch. One of the colonies "e monilored

(DalOlito mine f;3) ""liS \isitcd occasionally hy medicinal

collectors. Unfortunately. since Ihe locations ofmosl m i n ~are well known by local inhwilanls. long-Sflouted bats are

especially \-ulnerable to ovcr·hal"-esting. Howevcr, th

Charcani roost \\aslocat<.'d un the p r e l l l i s c s o f a h ~ d r o c l e c tric facilit}: thus access to this site was restricted and n

hal"\'csting of balS by humans occurred thcre.

To adequatcly assess whcther hW"'<C5ting of bats fo

medicinal purposes is suSl.ainable o\er Ihe long tenn

infonnation on harvest rates. )earl) ju\mile recn.titmen

rates. and hat immigration from outside the .study radius

nel.'l:SSiU)·, Nevertheless, ", e conclude that during thdrought, harvesting of any long-snouted bats '" ilhin ou

study radius would probahly h a ~ e been unsustainabl

because of the extremely 10\\ hat numhers, nhsence 0

juveniles (i.e, less than 2·3 months) or pregnant females

<Ind appan:llilack or immigration frum uutside our stud

radius.

An additional Ihn.."lIt lU lung-ltnUuh:d bat pupuilltion

occurring ncar the city of Arequipa is rapid urban growt

and exp3nsiOfl of resident ia l areas into cactus habitat. A

present, most DeW high density urbanization occurs nOr1h o

the city where areas of extensi\e caclus habitat occu

Ullfortunately, during tho: procelts of JiCttJement and ur

banization. nati\-e plants including columnar cact i ar

removed

Currently. no data exislS on thc population status an

ecology of long-snouted batS throughout the rest of it

range. Obtaining .soch infurmlltion is especilllly itnponan

because of this bat's role as a pollinator of columnar cact

the 10\\ numbers of bats found in Art:4Uip<l, and putt:lltia

over-harvesting of bats lor mcdicinal purposes.

ACKNOWLEDGEMENTS

We thunk M. DoneUy. T. Flcming. D. Janos, K. Lips. S

Petit. K. Waddington. R. Winkclman. P. Feinsinger and a

anonymous individual who reviewal thilt manuscript an

flmvided \aluablc discU5..\ion, assistance and comments

Laboratory facilities for carbon isotope anaJ)'sis wer

provided by l . SternbeTg. Imponanl logistical advic

gi\en to the authors by P. Jimmez made our collaboralio

puSJoible. Permission was graciously gi\-en hy P. rlech

Oaldiuan to conduct research on his property, funds fo

this research wcn: provided by grants from The Wildlif

Cunserv:ltion 5ocicty. a World Wildlife ~ u n d · G a r d e n Clu

of America Scholarship in Tropical Botany, Bat Consel"-'tion I n t ~ r n a t i u n a l . and a Maytllg Graduale Fellowship fromthe Cullege of Arts and Sciences. Uni\'crsity of Miam

(awarued to C, Sahley). ThiJ> llrticle is respectfully ded

caled In the memor)" of Mariscla del C. Sosa.

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A s$odaJe Editor: Don W,/son

Received: May 18, 1996

At't'f!I"ed' January 10. 1997

platalina genovensium natural history and ecology

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