platalina genovensium natural history and ecology
TRANSCRIPT
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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
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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
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§ $ §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
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-"......... 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
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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.
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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
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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' $Mt:ey 81Id S . ~ y H r 107
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c ' l ; p a i m e n L ~ nn IV. ll'cberbcueri cacti showed a decrease in
bat-pollinated flowers in 1993 relath,e to 1991 (Sahley
1996). Th;$ 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