bats as indicators of habitat disturbance_wilson_1996

8/20/2019 Bats as Indicators of Habitat Disturbance_wilson_1996

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ats as lndicators

o

Habitat

Disturbance

B w J a \ ~ , . > Ü ) Prc¡;m•ns, "\ia :ona M : - t ~ n m t o Nacurai - i:1tún

Smich.wr,¡,tn

lmmuucn \VashinKton:

DC

20560 usA

CEsAR F.

AscoRRA

SERGIO

SOLAR

T

D::p¿¡.rwm::>n:o di M a . s w ~ - ; x . : l o . ; J a , M t i i ~ · o ;;k'

Hrswria

Nac:<.mL

U n i t ~ < • n i d o d

:"J.vmna

i\-layor ¿ S1m ,'v1a ::os, Ap. 140434 Un:;; 14. PERli

BSTR CT

B01t

sampks

from

six

:t.rC>lS

in ;:'eru

show

that

the

rebtionship

bcnvecn

thos-e species

thar

an: rotttindy

,Isso

-i:.tted

only

wi:-h

ur.dls:urbcd, p:-im.try f(m:sted h : ~ b í t a t s , and ::nosc

th:<t

: ~ r e mo::c

normrdly associ::ire:d with disrmbcd, secondary g;rowtb hahir:1ts,

m t ~ h t

be

used

to

indiGne the d ~ r e e of cEsturbancc- of J givcn

habitar.

l"JTRODUCT ON

V a ~ 1 0 U S authors have 10ted the differencc m Corcpo::;itíOn ofbat faunas from

differentregions and habitar.s (Fbdlcy

and

Wii:;on,

984,

WiLson, l973 ). Ah:hough

rhe

:elmionship

benveor.

bm

spedes diversity 2.mi c0mposttion, '-'t:'td habitat

Jiversity and compositton

bm been r.mcd by

údJ sctcnnsts for years (Johns. et a

e

1965), quanti:-kmicn

cf

::he relationship has been Compa:-1s0ns of

b::Jt

faunas from distur'be_d and undisrurbcd habitats in Mexko (Fenron et al.,

l992),

suggest a

s t r o r . _ ~

relatiomhip

bet\veen bats

and thelr

habi:.:ats,

\Vith

the current

leve

of mtere;.;t in identifying habirats

that

rcma n

relativdy

unciisturbed witha view towa:ds practicing

m:;:;re

appropriate.conse:-varion

rr.easure;:;

in the futl:te, it would seern uscfu: to 2ocument the rciattonsl-:ip between those

species

of Dnts

tñm

are roctinely

asS()Clareci

only

VitÜt u n d t s t ~ J f b e d ,

prirr.ary-for<::st

habítats, a d those tbn: are more normally

assocuted

w ~ r h diz;turbed, second"

growtD habita:rs,

lf the relar::onships

can

be c s r a b ~ i s h e d for a vanety of ;;p..:ctesl

ir

r:1ighr be

possible ro use the density

of ce:tain

species ro üi.dicate

:he

disturb::mce degree in

a grven habttac The converse shculd h o i d ~ obvioGsly disturbed habítats can be

613



D-on E. W ilson, CésaT F. Ascorra and S2rgio So .ari

expected :o

contain

a:distmct subset cf

t :-te species that

mtght

be available in

undisturbed habitats.

\\lith these hypothe:-:es in minci. we ,:,onpared data sets for areas

in Pent

where

we

have

as.':>embied

considerable knmvled2c

of thc

bar f::lunas.

Four

of

O e ~ e

rcpresent areas \vith signiflcant dnd dJfc.::rent deg.,..ec.s of habttat dist;..¡rh;:¡nce, and

the others are reprcse:1latíve of l •md:srmbeJ. prím0ry lowLmd tropical raínforest.

STUDYSITES

Sucus RI

-n--.e Amazon Cerner for Environmental E2:.Jcanon

<-'.nd

Research (ACEER) is

a Scienrific Station heki by C o : ~ : l e r v a c i o n

la

Naturale-za Am8:onica

dd

Pcru

CONAPAC)

and

ACEER roun.:.lation,

~ m d

iocatcd

r:.ear

the

Sucusari,

a

white

·water tnbutary on thc ieft bank of the Nnpo River, :naín rributa·rv of the Amazor.

River, aSout 70 k:u \lE [quitos, Region ofLoreto. ~ n a z t ) t Ü ; : m Peru (Figure 1). As

in

the f c ~ l l o w i n g

loca iucs

o

Loreto tlle dry

season

bsrs

fron

ju:1e ro

Septcmber

(Spichiger

et

al., 1989). We snmpled

ha::.s

m rhe following h ~ ; _ b i t a t s :

Ter¡ac¿

foresr.

Undis.curbcd

fores;:

wi:h a cJr.opy

of

5 ~ 4 ( 1 m on

wdldratned

s:opcs abo\'C th<:'

t1oodplmrc.

Furesr

Borders of the sctcnuftc station surro:..mded by a

snnli

fnnge early

.sccondary forrst, :-egenerating frcm thc

statkn

clearing.

Y N MONO

Yuna:nono {

1'

16' S, 72-· 54

1

\\/

is an ec::::tourísm lodge operatcd bv E x p l o r a ~

cíones

Ama:onicas

S.A., in thi.'

the Quebrada

Yanamono, a w h i ~ e

\'1.'8.ter

tributary

on

thc

north bank of the Ann::on Rtver,

a:1d

located

60

k:n NE }quitos in :he

Region

ofLoreto, Amazonian

Pcru (figure

l). At

Yanarnono

we s a m p l e c ~ ln the

followíng

habitats:

ld secundar)' {ores

t.-·

A

largc

patcf1

o

20-yertr-otd successionai vegeration surrour.dcd

by continuous

prir:tdry

forest

1

pastures. :J.nd small

f c m i ~ i s r a.g:ricu t'...lral

d c o : ~ n g s .

Forest e d g e s . ~

Borders

of ~ h e

lodge

demí<1g.

lSLA MuYUY

Isla u y u y is an isia::td 1Okm long

:md

5km wide, located in rhe Arnazor. River,

about 20 krr: SE of lquitos, Region of Lo:eto, Amazontan Peru (Fig'...lre ). The

614



water levei

on the i s ~

land is reguiated hy

the sec:sonal floo¿ing

of

tl -.e

Amazo:1.

Riv

e:-.

DunnfS

the inun

dation pcnods oxbov,,

t a k e ~ n c ~ m a l l y i s o ~

lated from

rhe

main

rivcr, are connectcd

ro the rivc:- and

many

channei;:;

~ u e found

betwcen

tfie small

portions of non-Ho

odt>d

foresc

At

Isla

M ~ ; y t : y

we samplc:J

bat5

tr. the toHm.ving

hahitats:

N o . . ~ J l x l d e d forest

Parches of undis·

turbed pnmary non

fkoded fores::: s u ~

rrounded

bv

lüw

f:oo--

L

[ > n ~ - W O Q

70°

Fll;- i

Scud.v

.mes,

1= Sa::usaT ,

o;;;_ Ymumor.o,

::::;

l5 u ;'v:u)'U) .

4= Q u e ~ r a d a B a r ~ c : J , 5:=: ]é ro Hern:n.

an 6= P:Jlic:.::a

ded vegcwtion

agricu tural clemings and

..

1iHercnt srages of

secordary

growth

Agriculturt: ckarings_

Smal: ag:-:::ul::ure cl..:anngs ..vi::h fr c:ops, 2t:t\\Tc:t-::lparian

f:ooded

vcgeution

and vegt"tc.uon-

QUEBR D BL NCO

Quebrada

Blanco

is

a \vh:re

water

tributary

on

the

north

bar:k of

the

bh:ck

water

Tahc;ayo River,

in the

Corn.munat Reserve

T a m ; ; h i y a c L ~ ~ T a h u a y o ca

70 k

o

SSE of lqutms

Reg:cm

L f

Loreto, Ama:::onían Pcru (ftg•.:re 1).

At Tamshíyacu. 40 km N ; a i n f a ~ l ave-rages 2-,337 :nm per yt?ar (Bodr:ter, 1990).

T: e

vegecatior,

of the

::ueJ has bce:1 ch:sstf:ed

as

upbnd

:10n

::-1o(xied,

terra firme

forest (Bodrner, 199C).

At Quebrada

Blanco we

sampled

b:: :.ts ir.

the

fotlowing

habitats:

Ter mee foresc U ldist:.:rbe:.:l forest

with

a canopy of25 -30 mj on

stcep

ríght

margin

of the stream

and

undisturbed

fore.st wirh a

canopy

35 , 40 m en

the

t1at

left

margin

of rhe stream, <1bout 17

km

from its : n o u ~ h in

the

Tahuayo River.

ScconCw ry

forest. A four h2 pacch of regene:-ating

forcst wi(hifl undistt:rbed and

continuous pnmary

foresr dose to rhe previous s2.mpling site.

Tnis si

te was 3 fruit

farm approxímately 20 years ago.

615



Don E Wilson,

César

F

Ascorra

and Sergio Solan

griculture

clearings.-

A

>

30

1

ha agricultural

clearing

\Vith

fruit

crop

farming, ios:.ated about 8 km

ups.tream from

the mouth of the Tahuayo

River. .

}ENARO HERRERA

The

Centro

de lnvestigaeto "les "jenaro Herrera" (CIJH) (4 55'S, 73 45 W)

is a field

station hdd

by rhe Instituto d.:·

investigaciones

de la Amazonia

Pe:mana

(IIAP), located

aproximatecy 2 5

km east

of

the

Ucaya i

Rtver and 140 km

SSW

of quitos in t ~ e Region of Loreto

1

A m n ~ o n í a :

Peru (Figure 1 .

Rainfal averages 2,521

mm rerycar with

a

drierscason that

normally lasts from

)une to Sc-pternber { S p ~ c h i g c r

er:

al.,

I989);

howcver, in

each

month of the

year

rainfall

is

variable, and several rdatively dry

:nor:ths

occurred i:: the rain

y

seasoE

during

this

StUd)\

Meau

annuai tere.perature

is 26

e

{Lope:rParodi .and

FreltaS,

1990}.

The

average

e t ~ v a t t o : t ts 13 m

(Ríos.

J. u r . p u b L s h e t ~

rhesi::;).

The

vegc-tar:lon has been

cbssifled as

low,.r:errace

b r o a ' - ~ l e a f

tropical rain forcsr

by

Lope::-ParoJ: anU Freitas

(1990), wh.. ..; povided

a

demiled

descríption of

the

various habitats Jvailablc. At CIJH

\Ve

samplet1 bats tn thc foilo .ving habtats:

Tenace fore5t.-

Undísturbed or h g : h ~ i y (_-Lsmrbed

fore;:;t

with a

canory

of

21 ·

JO

::n, 1.3 1.5

km nortP. of thc

CIJH station d ~ a n : t g .

C ean:d ~ p · i p s . -

T

.

vo

30m

wide, lSC m

long

strips,

dearcd in ~ · l a y

1989 (Srri;;

l)

and

October

i989 ( S ~ r i p 2 ,

140m

apq:-;:,

:nsidc primary

forestar

1.3

km.

N of

the :::talion.

ScconJary

RegenerJLio:t fürest

arproximately

18

y e a r ~

old, zJor:g

an

aba:1dor.ed rímbcr

extraction road, apr::oxi:o:tme:y l

km

east of

the

3tation,

Open

a T e a s . ~

Forestry and a g r o f o r e ~ t r y rlar.rations withb S:m of the station and ncar

buitdings

wíth:n

the

sorion

dearmg

PAKiTZA

The

RIOLAT Biologica

Srarior,

at

Pabr:::a

1 t

56'S, 71''

l7 \t )

1

1s

a

fte{J

sratior. O ~ e r a t c - ( i

by the

SmtrJ-..so lian

ln5titution's

B i o d ~ v e r ~ a v in

L1:::in

Ame:-:ca

Program

(BIOLA

T)j locatcd

u:.

~ h e Rescrved Zone

of :he

Yianu Natíonal

Park,

a p p r o x i n ~ a t e l y 65 km

upstream

un

the

Manu R1vcr from 1ts mcurh

ar

the Madre:

de D k \ ~

River,

in the Province

of

: v 1 a n ~ I . D c p a r t m e ~ r

of

:V1adre de

D;os,

I : ~ k a

Region, Peru (Figure

l).

616



1

Bats

as lndicarors

of

Habitat Disturbance

At rhe Cocha Cashu Btologicai S<:ation, 21 ktn upstream from Pakitza, rainfalt

averages 2080

mm, with

a drier season that lasts frorn lv1ay ro

Septembe:. 1v1ean

annua[

temperature is 24.1 C. The elevation ofP'a'kitzJ is 356m. The

vegeratton

has

been

classified as lowland evergreen tropicnl

rain

forest (Kalliola e-ral.; 1987).

Although

smne

selecuve

timber exr:r<.ctior. may have

taken

place

in the

general region prior to the crcarion of

the

par k in

l970,

the amount of disturbance

was

mini

maL Erwin ( 1991) provided a detnded descriptíon of r:he various habitacs

avaitable

at

Pakítza. At

Pakitza

we san:pleci bars ut

the fGilowing

habítats:

Primary forest.-

Undisturbed forest Wtth a canopy of se m N,

ar:.d

E of the Station.

Forest

edges.

Borders o( rhe biological star:on clearing.

f{¡porúm

forcst.-

Vcgetation of cane

(Gyr:eriumsuginaram)

a:-td

ba::nOoo Guadua

Emd E y r r o s t t ~

ch;s), 10 20 m ctrarr a ong

stre2m banks,

Rive-r banks.-

Sandy

beathei::i bt::side

thc

lv1anu river . Somc

nct:> wen: phceci

acoss

the strean:.

METHODS

At Sucusar:

we mlst-nctteJ

Lns :nJune, 1991; July 1992;

Scprer;1bf:r.

l993;

and

December, 1993.

A:though

nets wcre se: :n::1in:y t grOL:nd lcvel,

atour

l/10

o{ thc

effort Wd.S

investr:ci

in canopy ne:s. In Yana mono, bats were misc-nerted in '\Luc:.,

199 ; ]u

1

y, :992:

December. 1992:

onJ March 093. At

Isla Muyuy

we s a m ~ · l e d

in August- Sepcemher,

l992. At

Qt:.ebra.- a Blanco wc sampled bvts :n Octobcr

p.;ovember, 1992. About 1/10 of the

netring

cffort at this

süe

w;Js spen.t in

canopy

~ e t s .

l\

rhese sitt'S w e r ~ sampied as

pC i"t

oi srudoes on h 2 t - p ~ a n t interac:wrrs

and

their

role in the propaga tive success of useful

ar:1azoniar:

pbnts

irl.

order to plan

the sustaí l.able use of

the

forcs::.

Atjenaro Herrera,

\\-'e

m ¡ s t ~ n e t t c d

bats in o v e - r a b e : : ~ e c e m b e r 1988 ar:d from

lviay 1989

to

Ocrober

1991.

J s p ~ . r t

of

s ~ u d y

of nnrural

r e ~ e n e r a t i o n

afrer strip

clcaric.g. Net:ing

f f o ~ t

in

e8ch

habita: each

: r ~ c n t h

cor.sísted

of

1 - 3 mght:::L

f.Aost

ne::s were p accd

at

g r o u n ~ levei {0.5 3

r.1).

a thcug:h in deared s:rips about holf

the nct:tir:g effort was

at

n:id-story and subcannpy ie;tds (5- >20m).

At

Pakin:a,

\Ve r:1ist-nette::l

bats in Ocwbc:r 1987,

Scpcerabcr

1988, Nc>vcmber

1990, and Febmary • fv1arch 1992 as p a ~ t e astudy of rhe toral hídogical diversity

of the area. Netting

effo'i:"t

a: cach haSitat each montfi

co:1:slsted

of 1 10 r:.tghts.

Most

nets

wcre placed nt

ground

level (0.5 3 m), ;rlthough l í l of

nerting

eff;)rt

was

at mld-srorv and sub-canopy eveL., ( 5 -

>

2 m).

617



Don

E.

Wilson,

C¿sar F. Ascor-ra and Sergio Solari

Nets \ven: checked

fo:-

bats every 30 mmutes. ¡-.,¡etted bats were

held

in

doth

b;;;gs, identified,

and

measured (forearm

leng:h and -..veight)_ Heigh:

f

capture

a':

well as habitat

was

recorded fot

each

n1div1duaL Rcproductive t a t ~ S (pregnar.t or

lacmring) was obtair,ed frorn seiecred kmales by exr.ernai c-xammatíon.

Some v o u c h e ~

spec:imens of -each species we:;-e saved anci deposir.eci

:n

the vfuseo

de Historia NaturaL

U n i - v e ~ s i d a d

Nacional Mayor de San tv·1arcos in Ltma, Peru,

and i:1 ~ h e Naticnal N l u : > e ~ I m of

Natural

H i s t o r ~ ' in \XI:d"".mgt>.xt, D.C., U5.A

Otherwise, indiv dl als were re eased

at

r.hc e::td of each net:::ing session.

Thc :-a tío cf caprure:-ínetting scss:on, was used to standardi:c tb.c captures from

each

sin:.

Bat species con-..position in

the

J¡fi.crer.t habitats was compared using the lndex

ofSimilaritv I=2C/A B \VherE" e equals the number cf speci::s sh: red

between

the

two hab¡tars

;:md

A a:1d B are

the

mar.bers of species in eoch

ofthe

twosítes (Krebs.

1985).

Thb

index

~ a n g e s

from

Ü

(no

overlap)

toLO

(complett:

, J v e r l a ~ ) .

For de;:ermmmg the

twphic

structure of comrnu:tities, each ·species wcre

segregated into o::e of the fdlo\vir,g

trophk

gdlds:

Aerial lnst?ctlvorcs.--

Emb-allonurídae, Noctilicniúuc Thyrcpteridae, Furipterkbt , Vespertilionid3e,

and

M u l o s , ~ ; i d ~ e .

Foltage G b ~ n e r s ~

Phyl ostor:linae (lv1acroplr; lur<l -rnacTophyllum J,"v xron Ctt'ris spp., ? v 1 i m o r ~

cre7tt-tla.Lum,

Tonaría.

spp.,

and

Ph:vHosto-rrws

elor.gmus).

Necw-;

Feedcrs.-

Glossophaginac

ond

the

P h v l o s t o ~ n i n e

P ~ \ v l l o s w m u s

discolor, whích we

ofte l

fcund d u s t ~ d

W l t ~

r(JI:en ]c:1a:-o :-lene

m

and :shl

lv1uyuy.

Phyllosron:irte

F n t g i v o r e s . ~

Ph)Uoswmus

ñc:staws and P,

stent)PS- Thc fir::rt

is usuall

y considered an

omnivore,

but wc fcund :na:ialy seeds acd occasional insect remams in thei: feccs. lnspccc:ed

day

roosts

at hunan

chvc lmgs

only

had sceds o:- fruit r e m a t C ~ S ,

Undeoswry Frugit-orc;¡.-Cacolliln'i1e

Car:opy F r u _ g i v m · e s . ~ A r t i b e u s srr-

Othe• F r u g i t i O r e . L ~

The

:-ernair:mg St;;:r.oderminac,

Camivores.---Lmge Phvl ostcrnl:1ae

that

feed on vertebrares

Chrowpterus

auritus

and

Trachops cirrhosHs).

V a m p t r e s . - B i o o d ~ f < : : > e d i : ~ g

ht:s

(Desmodus

rotundus

Dnd Diphyl .a ecaHdcta

618



Bars as Indtcawrs

of

Habita Disturbanc¿

RESULTS

ANO

DISCUSSION

At

Sucusari,

\VC

:ecorded 33 species

in

20

net::if'-g

sessions"

At

Yanamono, we

registercd 36 species in 24

nert::tg

sessions.

On Is a :Vfuyuv. we obmined

19 Sat

species

t

7

nerting

sessions. At

QueSrada

Blanco, ;,ve recorded 30 species in 13

nettíng

sc::;sions. At Jenaro Hc:rrer3. we

re;.::or...:kd

ó2 in 103 netti r:g stsswn.s

(Ascor:-a t aL, 1993), .:md ac P2bc:a wc

rtxord

57 in 61 eetting sesstons

(A.scorra

ct

al., this v,Jlume) (Tablt-

i

).

Alrhough the i ~ i s causcd by the different t éttir:..g eit"orr a

e

eachsite is corree red

by

the of rhe ratlo of Capturc/nz:t:tng effort, tf. .:

d c f f e ~ e n c e s

in species i v e r ~ ¡ t y

recorded at each sitc m::1y be due to rhe u:lequal sampling e l o r ~ s .

L 1

J Herrera.

the a l r a o ~ t

conttnmtl invcntory

of

spectes

t h r o u g ~ l ::tme,

and

the sig:1ifc8nt effca

spenr in

canopy and

subcanopy r:::tst,netting re:;ul:..:d m a more

com.plett

;;arnple

of the bat fauna (Table ;)

In all pbces the ba:: fauna seerr.s ro De r c p r e s e 1 ~ e d Oy a few species with a ~ a r g e

nwnber

of

i n d i v i J u a ~ ~

::md rrunv

with

only

J.

few captures (Ftgure 2).

T:able Con:par:lti<.·;: s::ucturc of ba: c o m t m m i n e ~

tn

tice sudv site5 csíng

(\--.e

r:mc·

captGres/nettíng dfott

(,A,

1-=-

;n;:i?.:

i n ~ e c r i v o r e ~ .

Gí"'

g e ~ m e r nsectivores, CA= clrt1Í\ {_m:s. ~ E

ncctmiv;;res,

UF"' U : 1 : i e r ~ r c r y "n,givores, CF"" canory frugív.xes Arribcm spp.), or: mher

S t < ? a o d e r n m : v ~ . PF"'

PhvJk--,stmnin<:

Íi'ugívorcs

Ph::n&mon:uc; hast(ltus,

P.

srcwJps), VA= v n m ~ ' i : e }.

S e

d.:;;

· · · · · · · ·

· · · ~

Pcrvpt.:r:x kapp .:n

/d

P rut;:,_r<)

rc C<;."Oj::.t'TII

/ ..)

C<'U

ghsr;d-u;.:n;n:ns twsc

Ai c1

C\'7

('i,)t_''-j

e

~ o J

S : J . , x o p : ~ r y x b t,'L.'cr.:.t

A:

o,,,

_ . _ ~ .

0.(':

{

(' ,((\(;

o , . , ~ o - + 0,(('4

S o . n , a p c ~ r y x

U:pnm

.·\i

l :):)j

O,OJ7

o '('6

o

002

Noci n albit\."n:--rs i'\l C.CDI

c.ou

M,,..·roph:1i:tm •ruxwph, iwn

..:_:¡

c_c¡r;

M : : . - r ) • - , - . , · . _ : r ~ , . T i ;

~ w ; h " ' i O Í S Gi

0:2{12

, \ ~ < l T O l l ; ' t : r t ' T l S

n : ~ · . w : : w : t s

(1(

C.

COl ::.C\:6

e

ooz

J

CC6

\krcnyc:cns

ffi :liHG

C'

,

c.o: ·t

O.Cl2

: V : i c " ' - > n : ; . : < ~ n s n;.;:fon

G:

ú . ~ ' O J

0.CC·4 OC24 0.CD1

Míc-onycwú schln«Ü.Jrum

Gl

C,OOC

~ A i r r o ; :

c n : n : . i a t ~ m

e;

C.Cto

O.C26

0.182

O.C'll

C.Oú.f

F'A):lostomus d.:soior

é:E C.105

OC02

f'hy ioswu:us

.:bngma.s

Gt

G.ü20

C,023

0.025

ÜY)42

C.ú07

0,0J5

P h - : i . o ~ w m u s hastuws

PF 0017

o ccv

'::.16',' 0.0 2 0.069

;),025

P h : - ; / 1 o ~ t 0 w . u s

:;¡cn0r>

PF

0.0C2 O.l'C4

Tó:inn.::a i:>dcns

G'

o_co7

O . C C · ~ 1

0 >2D2

O.OC6

T

mt:t tk & r r u l i < ' r l 5 ~

Cl

O.OCl

C.Oü

Tona.ca ccrrr:ka1

0.006

O.C'Ol

Trachops

..:mftosu.;c

,.,

~ r . 0.003 0.C'J7

O.OI:J 0,012

O.C\/1

o : Jl::

Tonmw stlvKola

G: ('.003

0.002

í.\012

();)J4

O.OD

Chro1opreru;, a:.ntu.s

CA

C.;:07

0.002

V a m p ~ w n

s}>. .:rrum

CA C.OOJ

Anoura

c a u d t j ~ ' Y O

NE

0003 0.002

C.COl

J.C'CJ

C h o c ~ o n ; : ; c u s im.;rmcdn;s

NE

::\COl

Choercnr;:cus

mir.or

NE 0.006

619



Don

E

\vi/son,

César F. Ascorra

and

Sergio So/ari

Speóes

Guild

Sucusari

Yan.1mono

l. Muyuy

Q.

Bl-a:1c0

J.Herrera

G os.1opha;;r1 commissan;;l

NE O.OC9 0.025

OJJC6

Glossophaf a sanciHil

NE

0.076

0.056 00L2 0.008 0.013

Loncbphy /a

rr4nbx

NE J.054

O.MI

Lanchophyi n thorrtr ."Í

NE

OJ/22

0,() 8

o.ocs \1005

c,lroll4r l:m.:ticnuda

UF

OJU 0.107 0. 42 C'.lCZ 0.158 0.030

C:::.rol ia cosu:Hk'ü

UF O.lCJ O;J55 0.102 C.026 OJ24

Cc.rollia

persptC1iilrW

UF

0.041)

0.396 0.265 C.036 0.273 0092

Rhinophyilc.

f¡Sz'h::l'o.O

UF

0.00)

0.018 0.0t

Rhínophyila pwml:o

UF

ü.Ol0 0.02$

0.21

0.0ll 0.006

Artibeu:;

arv: t" 'wu

CF

0.020

0.::'63

O. lOS (\002 1l023

ArtJI::cus cinccu.> CF 0.006

0.004

Aníb.:us ::otUolcr

CF 0.007

Amheus Ri.:mcus CF

O.OC6

Anibcus

gnomus

CF

O.Cl2 0.014

C.CV5

Amb..4As J-..awn CF O. COl

Artib..'uS /itwmus CF

:h""53

C.Ol5 O Cl2 0.096 0. 24 0.052

i Ar.:Jxus

ohs<.:u:rus

CF

0.053

C.OC\.f

0.025

C.054

O.CJl

0. 9

A1[1beus p a r u r c s m ~

CF :.l9J

e 2<+

O.OZ4 o056 0.101

Chi:-odtrm::: nibHw: OF e003 0.004 0.002

CLC04

Chrrod.:1w.a trin:wtwn OF

une

(:( ()[

C.(\01

P anrrfjr¡w

Ju:r::;:nus

OF

O.C\'4

PlorPhinus bro .._·h:;cep.fu,:: .:i

OF

O.l\)6

0.('1)6

e

C\'l

C.Cl7

P atyrrhir.m h d c ~ · ,

OF

c.:ot

O.Ol i

O.C04

l ~ . O l 6

Plmy:-rhinus m/uscus OF

C.C04

Sturr;m;¡

iihvm

OF

;;.ct7

O.CJC o.on<

Sa.:mna iwn UF O.CJ6

St:mlÍTG 1TBS.'Hd OF

0.Cú_i

0.C04

Str<:1tin: tJ/:.1:::

CiF

o.oo::

0.CC'J

l

'r:Jdi nrw

~ ; l o b _ ; a w :

C>F

0.01;:

('.L\20

C.Cl2

2

015

0.021

Urodcmw l ' l . : : H , ' l : l r o s ~ n ¡ r , ¡

OF

0.(\)2

0.002

0.016

Vmr.p)'1<'SS4

bid,•rs OF

e

OC('

~ . 0 ( :

Vnmp::r.:s:.c;

i : m ; ~ - ; . ; 1 1 ( f

ec.ot

\/wt,p;n,;va

ma..<omJ;:Ur

OF

e

COC:

o

004

C.Oi

7

V,nnp:'IT<'SSa

n;.'ii.L;l

o:::

c . ~ ~ c 3

0.0:)?

Vmr.¡ryro:d p ~ e 1 i l ~ 1

0"

0 033 O.CO'J

O.CD4 0.COJ

V:rmpyn:rl:s

lG.TOtXIr. l:

OF

0 . ~ Ü 7

0.005

DesrnoJu.¡

rownJus

V

J.(\)J

CJ}:'f

C.C4S

._

1

.CC5

Diplryi'o c:mukuü

VA

0.003

Thyropwrn :\

0.C0l

Tf-.yropta::

:'\.

c . ~ . ' o ~

Q.U6

Fw;pL-rus

h.arL'n.:-: 1 -.i

C.(lúl

C\CiJl

Ln:w·•.ü :.:;a

Al

C.OC

E"pt251C IS / : n t : : , i l i L l \ S i ~

Ai

c ~ o 1

E:.m:ops ltmnue

Al

\'.CQ6

Myons

u 1 ~ s e ; : ; ; s

.AI

0.ü24

0.04

v1you.o rugrrntns Al

o . ~ : ; . : : - ¡

l\004 C.C06

C'.207

C.0C4

1-.h·or;s

nt,trrus

/-.

0.0:3 J.Ot2

(\('04

e012

\(:m;s > i ~ n t s

Al 2

C('6

OC\:3 C.C·ü.:J

Mc ossus uccr

AI

CGül

Md:_•is.ü >Hoimsus

Al

:::'006

::

C OS

0.001

Mdoss:t.S i ' : " . < : t o ~ /d :J.C \.. \2

M c ~ o s ; J 6 ) ) 5

.•\i

O.Jú2

Pr<:"rnops ]"\.i

C.009

..

·

. · · · · · .. · · · ·

NDI\/lDL:A.LS

{7ur;d)

301

46C 162

lb

2229

172

KET SESS DNS (TotaD

~ l . \

24

}

103

51

620



atsas

Indicators of Habita<

Disturbance

n ~ e símilarity in bat spcctes hcv.veen Isla ~ v f u y e y

a:-:d

any

other

si te was very

low ( <0.44

),

while

the

other

.si

tes proved similar, jn bot :jpecies compositíon

(1>,05) (Table 2). The highest values of

similanty

were found

between

Yanarrtono

anJ Sucusan ( l ~ O . 73) and t ~ e lowe:;r

(

. 31) bcrween Sucusari a id Isla Muyuy

íTublc 2).

Aerial insec:,vore:::. J ~ e more

abunJant

ar SucusJri, Ya-namono.

am:l

P a ~ i t z a

(>0,05 captures/nett::ng sessicn), ::;:tes wrth d o s ~ c i < 'r'.d c o n t ü : u m : . . ~ foresL sla

f.,/fuyuy, Quebrada Blanco, and Jenaro Herrera, wíth less than 0.04 captures/netting

:;;ession, Gre

meas with only

fnrest

parches, 0r h:1rge

farms dose

to

f o r e ~ t ,

or

brge

pasrures, and agricultur-a

dear:ngs respectivdy

(T

able l and Figure }.

Foliage

Gleaners

are more abundant in Sucusari, Yc.namono, Quebrada Blar.co,

;:md

Puk tz3 (

>0.05 captures/ncrt ing session), where there lS tall

and

contínuous

forcst, ;:han in Isla : \ ~ f u y u y and Jcnaro Herrera (<0.04 captures/netting se-s.síor,),

where

rhe

forest

is

patchy

or

dtsturC.ed

\Vir.l-..

fc•restTy'

plant::lttOns

(Table

l

and

F'gu:e ).

Carnivorcs and vampires secm be rt1re 0nd less abunJ:mt everywb:re (Table

1

and

Figure 3),

although thc

la.ct<:r are more abundant in Je1::1ro Herrera (Tab f'

1) tha<: in

the

other sncs.

~ e c m : - 1 v o r e s reach the highcst va u es ír. si tes wtth fruir crops

el

ese to :he forest

sucn tl

Yan8.mono,

Is a Muyuy, and

Quebf.\,_'ld;:t Bbnco,

\Vhere rhey Jccnunt for

16

1

'

'

¡¡

'

-

o

( )

.o

E

"

1-2 3-5 6-11

12-23 24-47 48-95

>9

Number of lndividuals

1 : Sucusari - yanamon¿ ~ - ~ M ~ y ~ y - - l

Q.Bianco f f ~ J Herrera

t%'1:

f'akitza i

~ ~ · ~ · ~ · ~ · ~ · ~ · · ~

621



Don E. Wilson

César

F. Ascorra

an.d

St:ig O

Solar:

Al

~

Gl

C

Sucusari

Q.B:anco

VA NE

UF

Trophic Guílds

· ~ · ~ ~

Yanamono LMuyuy

B

J ~ H e r r e r a

B

Pakítza

F:g. 3. Ahmt.L:mce nf lxn s{Jccu cs e< srHd)' <;:tn

b · f,'la ds.

: \. Pn:dttron . Af= aerir.d C l . S ~ . c t i m r s ,

GI= fo&¡gc CA=

c::r;l,t'(JT<:>.

VA= n:rr:pm•:i, Pf-::.:.

Pfr;liJ;'iu_mt:r.i

f r ; . ¡ g : v : ; n : ~ . UF=-

tm.,bs-

t;:;ry fn-¡g:wres,

CF ::::

curwpy

_ t Í 1 ~ i t < J r t . i ,

OF::o- o r h ~ _ : > r s:enr

d:__-'nl' PlC

,\'E:::: r.e::tam'Ore•)

>0.08 capture:o;/nettingsess;or: Table l ~ m Figure 3 ), Undcrstory frugtVores

r e ~ K h

the towest va ues ( <0.3 c8p ::u1'ts/nc::ting s e s ~ i o n ) Ü:l undisn:rbcd J.reas such

Sucu::mri ar:d Pak t::a,

wt:tle

the co:1verse S found in ateZls \Vlth

nearby

and

considcr;1ble secon.Jary grov.·th (

>0.4

capt;_¡res/nettmgscssion) sueh as

Yanamono,

lsla

Mt:y;._¡y,

Quebrada Bbr.co, and Jenaro Hcrrerca

c:·ab c

1 and figure 3}.

Canopy frug1vcres

m

the g:cnus A r r i b e t ~ s Zi.re 1::wre abendan:: at Sucusari (C.5

captures/netring

e s : s i o ~ )

Table l anC F:gure 3 ). Thc presence of son: e salt licb

m rhe area and the ...lrinking

bchavtcr

by Asc;_:;rra and \ X : ~ s o n ( 1992),

m1ght

be affectmg

the rcprescn::ation uf rhcse b < : ~ t s in

s2mp es.

The remarkable

c ~ b u n d a n c e

of

othcr

S'::t:nodcrmir:e frug:vores

in Jenaro

He<Ter8

carmresí

netting se:ssio:l)

might h:

a rcsult of the

higher nen¡ng etTorc

::;penl in d:e

c a r ) ; 1 ~

py in tftls arca ( T ~ 1 b l e l an"i Ftgure 3 ).

In the

six

sitcs,

frugivc•:"es

are (he domi-nanr group, but Unders:ory Frugivores

Ca-rolllínae) <:lre :non:- ;;,bundant m Y a n a ~ m o r : o , :sh> j\.·fuyuy, Quebrada B anco

and Jena-;o Herre-ra, proba-bly

dL:c

m ~ h e high ¿¡vai-hbility of their main :-ood

source

pioneer

and secord growth p ant sp-ecies), thcm at Suct:sarí

and

Pabtz3

where maturc foresr vegetar¡on is predomir-.ant.

Od::er diEc:rences berween

si

tes appear

für the

other

g;:m¡ps

vf

bats (Figure 3 ).

lr.sectivores are :ncre ~ 1 b u n d a n t ín n : : m ~ d i s m r b e d areas and this \S probably

due to

622



1

Table 2 . ~ N u m b e r

of

b(lt

spe:iei neued

in

t:ach x

anJ

mdex of m t i l . a m : ~ Oetween ~ n b ú : a c s

Sinüíariry

t v r ~ h :

Site N Yanan:ono

L:.byuv

Q.B;aoco

Sc:cuszd

Y:·onammw

J.

~ L y u y

Q.

BbW.))

_ . Hener<t

Pakinn

3J 0.73

)6

19

30

6Z

05 :

o 55

C>

7

J.Herrecn

Paktt:a

che

high

pL:mt diversitv thc.t also supporr.s

high

divt:r.sitv

of ::1.:3ects.

In high y

distc.rbed a::eas rhe h_-,\ver plant Giversity ar.d

n b u ~ t ~ b n c e of

a fe:w pioneer e c o n d ~

growrh phmr species mighr be a:-rc.:tlng

the

qua.:-t::ty or qualí:-y of

the

msect

resource. Howevt:;-,lt mav also be casiedor

lnsectivorou;)

p e c i e ~

witit their highly

devcloped

echolocation

svste:n,

to

d<:tect

thc

m ~ s t ntL'> ir.

rr.ore oren

areas.

The

w e l l ~ d e v e l o p e d echotoz:ation

s y s ~ c m s of

:nost i n ~ e c t i v o r o u ; ; bats

ma:kes

ít very

díffk::J::

to

s ; _ u n p ~ e

du:m

i:1

any

unbiased way

u;:;tng

rníst ncts,

The phyllostomtf'.C frugivore Phy k;swmm lws:.uws (Table l ), most abuadant

in disturbed areas, rnight be m2ícating thc at>:..;ndn_r:ce of m: food source

:n

this i r ~ d

ofhabttat. F e c u ~

5amrle

analysis shmved

thc

prefercnce cf this speCics for fruits of

Cecropia

s;_Jp., < 1Il

oi:

which

are pionee::- specics, as wel

~ ; s

for

the

loc:Jl fruit crop

Po<- TOnma cecropiaefolia, Po lcn grains ? v : ~ s a jla;·adisiaca ctnd Ochroma p;:ramidale

hsve becn r e ~ c : : : w e r e d from dusted f¿¡ces a::tó fur of

r ~ 1 b

bnc

The

abundance of a ~ n p í r e : at

Jeacno Herrc:-il :s

obvi(l;_¡sty

duc

tocan

k (lensity

ir_ the area, wherec1s domestic l vestock :s rorallv

L-KKtng at

?aktt:;c. Othcr gcot pS

such as necnrivores, rnore com:non in s e , ~ : : : i : l ~ h t r y b:í.'St

or

;'-g:-cCllltural

h a h r a ~ s .

might

be reflc:cr:ing

thc

::d::un&.;nce

of their :-ooJ

sources

in . ~ : K ~

hab¡tats.

Compamttve

a:c1aly:us

ofspectes

o m p o s : t ¡ o ~

< . m ~ abundano:':

bctv/een d:c

s-:u'-+{

sit.es (Tablc i), s h c ~ w ~ 3 norevv\)rthy rela::wnshtp benvecn the abum_--lancE' (or

s c a r c \ ~ . : y ) cenain

species and

the

degrce of disturb2ncc

(c)f

:1un-Jistu:bance) of

thc:

h<.1b: t.

Spccies su

eh

as

Ca:

oltia

pe ·stJinilo:w,

PñyUoswmus

, f u . 1 5 t d t u . s f ~ m d

Dcsmodu5

mnmdHs ::m:: more abunclant m

h1ghly

JiswrbcC l:c1bírats. \Y/e suggest

that

rhe

abundance of

these sp.:c;es

m ghr

be used as ;a: ind catior:

vf h i g ~ 1 O a h i ~ a t

dismrbar.ce.

Taxa such as E m b : t l l ~ m u r k b e insecttvorous

I _ : : ; h y : o s t o r r ~ : n a e ,

T h y ~ ; : - p t e r i c i a e Furipleridae, a:td V e ~ p e r t i t i o n i c b e (lab e

l

are :norc Jbui:dacu:

in

undisturbeJ

üun

in disturbed habitats

and the b ~ n d a n c e of

thcse

p e d c s

~ e

used as

indica

non of i t t ~ e or no habit:::t d:.1rurbance.

Although

many

bat

species

-Seem

re be abur.dan: at any

~ ~ t e ,

thcy

probal::ty

are

using

m i . c r o h a ~ 1 i t a t : s

dtffercntta iy at

each

sin:. Fcr many s:tes, more intensive

mvenrcry work

is

nec-Jed ro c o n ~ p l n c l y

:::ssess

species cornposition.

In

g e n e r ~ d we wouid suggest t h a ~

a::t

abund:.mc; of n1er:'.bcrs ~ - l ;:he subfar:lily

C::.rclliirHe

is

a

gocJ

tndtcatior:

of distcrbcJ habmns, a::tci ~ m abundance

o[

phyUos:::omme insecti\·cres indicmes u n d 1 s t ~ ~ r 6 e d foresL Perhaps the best single

índicator of Jtstc.rba :1ce is the common v ~ w 1 p i ; e Dat, Desmodus r o ~ u n d u s

which

ts

abund:::mt o n ~ y in areas

that ha

ve

becn

n:odified to support dom.estic li\-'eswck. :t

is

more

diffícu:t to

síng e

out

an;· particular species as being uníforrr:ly

gooJ

indÍcJ.tors

of undiscurbed forest, <lS aH such srccies naturrtl y occur

at

lmv

623



Don E. W lson,

César

F. Ascorm and Se-;gio Solari

popularion de:tsities,

Howeverj cur

data dearly shc'>v rhat, wirh the

exception of

Phy[/.ostomus

hastatus, members of :he subfamíly P h y : : o s t o m i n a e - 3 . r ~ far more líkdy

to occur undis<.:urbed foresr hJbitars

RESUMEN

Muestreos

de

murcíélagos en se:s n r e : ~ s d(' b c ~ m a z o n f a peru,_¡n,_¡_

demucsrmn

que

la relación

entre

las es¡Jecies a s o c i J . í . ~ ; . l s con bosques prirnar:::.s, no ir.tervenidos, y

aquéllds otras asociadas cun a:-nbientcs intervcnidu8 y for:nackmes secundari:Ls

podría

ser

usGd·a para í:1Jicar el

graGo

de lnte::"\'C:1ClÓn de

un hábitc.r determi:1ado.

ACKNOWLEDGEMENTS

Thc

research

ot

Sucusan and

Yaneil'<:lono

wns

SUIJpOrted by COJ.\ APAC

and

ACEER Fou:-:dation, Grant ::_¡ C. Ascorra. Specia c h a n ~ s

to

Peter

Jcnson

cf

Exploraoones A:nmonicas fo:

res::::arch

f8cbtit:s

at s : r e s . The

:esearch a e l s ~ a

Muyuy

arrd Quebmz-b Bhmco, and the fina: data. aLa:vsis ~ 1 n d c:Ot:lputer

work

was

f ~ m d c d by \X/orld Wíldhfc FunJ s Bíoóive:--sir.y Surport Program Grant No. í564

to

C.

Ascorw and

D.

\Vibon. \Ve thank

thc

lnsrituro Veterínariz> de

l n v e : : ; t i g a ~

cior:cs

T

fO['ÍCah:s

y

de

/\l::ura (C;-IVITA)

of rhe l.Jniversi¿ad

1'\ac10nal

M a ver de

San Mar.:os for pemits and f : J d l i u e ~

Isb

Muy;.;y and

Quebrada

Blanco

Bíologica[ S t a n o n ~ .

The at

Jenaro He:-rcra W dS supported u r d e ~ Grar.t

No.

7

228,

F n : : : g r a ~ 1

in

Seter:ce

a:¡.J

Technology

Ccoperation,

Officc

of Stier.ce

Advtsor, lLS.

Agency for

Intcrnc..tional

Developr:.e:it

l} Gorchov,

F.

Cornejo,

andJ. Terborgh. Resc:Jrch;_1t

Pakit::a \Vass:Jpported

l::y the Smtthsoni<:tr.ln;,títunon's

Biedivers ty in

L:tin

America Programs RIOLAT)

Grant

ro C. A::;corra \Ve

thank

the

Offke

of Fellowship < ~ n d G r a n : . : ~ t.>f rhe

Srnuhsoman

I n s t i t u c i o : ~ . for

surporting Asc,::rra's work in rhe a t i o n a l l v ~ u s e L a n cfNat.Jr:::tl History Collections.

\Ve thank

the

Instí'::uro

~ a c t o n a l

de Re;;;:urso:::. r..;arurale:::- INRENA e x ~ D t r e c d o n

General

de Forestal y ~ a u n a

a:--td

the Progr2ma de Parques Nacio :l::tles

of

the

: V h n : s n . : : : - i ~

de A g : - ~ c u l t u r : : t a n c ~ ~ h e

Dm:·ccto:-1

R e g ~ o n a i <.:e

Recusos N<J:uralcs

y

~ v e d i o A.-nbiente

of the:

G(:.hierno

Reg1cnal

de

Lo::ew.

for collecting re:-;rtír:s. We

thank

::hr

Insr:itL;W

de

fr:.ve:::tii;ac;ones

de

t<:i

Am<.tzoma

Peruam1

(IIAP)

and

J

Lope.:-Parcdi a:1d lvl. Isu:::a fcr use and fac:licies of the Centro de lnves:tgacioncs

1

'}ena::-o

Herrcr:J"_

\X/e Ü:ank ; u ~ l r .

F. L o j a ~ A l e : 1 1 , 1 n . S. Sevick, 2r.d E. Srudler for

their

valuable

assistance

ir.

r:he fteld work at Sucuse ri ar.d Yana:nono.

Lccai

L::trmers

Gonzalo T o r r e ~ anG

Juan Huanaquirí

gave us VJ u:1'::le Cele assísrancc ;)n

Isla :t -.·1uyuy anJ

Qucbnd:::r

Blanco respecttvely. R. A r a n a ~ C a r d o D.L Gon:hov, V.

Pacheco, and J, Pair.1einm hdped wnh che fteld work during c:crr:am m o n t h ~ a::

Jenaro Her;em. f. Dallmeter ::mJ M. V. T..:nícela gave vab.wble ~ e l p wit:h fíeld

work ciurir:.g certaln :nonrhs ru Pokírza. 'We

e s p e c i a ; ~ y

rhar:k

J;;tmes

Penn ]::. and

rhe local Commur:ity of Cbr,o in rhc

Comn':unal

Re:;erve T a m ~ h i y a c a · Ta"nuayo

for

thetr involvcment

in our

resea.rch nnd fo:

opplying

so:ne

of

our

results

in

the

6Z4



Bacs as

indicawrs

o Hc•biwt

DisturbaJ

1

..

ce

managcment

of

their

::omt;f al reserve. W e thank F. E n c , 1 r : . ; a c i o r ~ . M.B.

Fenton,

D.L

Gorcfiov,

andan anonymous

revtewer for.thelr

comwents and

suggestions

that

i;nprove¿ this pa.per.

This

is C o n t r l b u t ~ ~ m ~ o 44

of

the

S:nithsonian

[nstitutinn's

Biodtvcrsity

:n

L1tm

America

P:-ogratns

(BIOLA

T) anJ Cor:tnbutioE

No. 2 of Prog:-;;m¿ de Evoluacion Per:n;:1ncnte de l,t BmJiversidad Ar::.a::oníc:J

BIOAMAZ).

A ~ C C l l T ; l ,

e

F an·J D E. w \;:on_ 092

83r

frugtvnry and seed ,Jgrenal ·n

rhe AmAzü:1,

:.Netd, PenL PuSl.kJ.::;nnes ,;ld ~ - f u s e c l :ie Hi5to

rla Na:L:a de

la

L.'nive:-:>:dd NJc:nn,Ü :ViayN

cic

S n M a : w ~ Ser:e .'\.

43: L-6.

i\sc:)rr.1. C

..

D. E Wi s:m, z n ~ l Jvl_ Rnr.n.

199:.

Ltsw

¡:¡nnt<<dJ ,Jc

k>> Q t _ m 6 p t e r n ~ ,]el

PJh¡ne

Na..;mnal :-..bn,t, Peru. P u 0 ~ t c K - t . > n e ; , Jd

t v f u ~ · : o de Hbte>rt.l

:\"atwa:,

L : a ; v ~ . ; r : : t J : - J d ;...;,,<;10·

:-tai

}..;;¡)'¡; Jc Sdn

:Vbr;:n;:;, Serie,:-.,,

,.',2:L- J..

Ascmm. C F.,

D.

L.

G()r,:.hov.

< H 1 ~ 1 F.

C'rt'.tJ-C'· J •)93

The

h:ns frnrn

Her::t' i\.

L'r.;;:;'>, Peru.

\ümmJ:L1,

r ~ . , . : ; m e r . 0-. E :

9'1('- R:·sp;' 1SCJ

~ ; : l H \ ~ . I ; C ) ( t ; )

S<.,>:lsnn.Jl ::hJfhl:.H:ons in

rhe AmJ;nnl 1n l o c • J t ~ ; . ; n _

Jnurnal·:i

TwrtcJl

b:l9l-:Ct

Erwin, T. L l9CJ1

>-l;;rur.ll h . ~ w r v 0f

the

caraó1d b e e t k ~ 3( tht BIOL:\T BH>k .,;;;;;ai

S:;Hlcl:. Ri0 \ian·J, PJktc:::;. ~ ~ e n .. Rcv:s(,< [\>

rwm;¡ de Enromo ngi.>, JJ·l . : : ; ~

F ; : n n ~ n . ;,.t 3 . L D . 1 . t k ~ : : - t , \LB.C

Hkkey. C. V.errim<Jn. .; :1:-or.or.

, e n ~ ~

D.\1

S:;me. \992. P h . y l o ~ t ; _ ; . m ¡ J b:ns

~ ' h v ·

\lo.:,tor;:.,)ae)

ns

m,JH:ator5 ot·

h.thL1t ~ ~ : : n t ; - m o n

tn

~ h . e Nené"L'TK5 Bí(•rrnptc:t. 24:, '-H.:A-16.

L TERATURE CITED

: : ; m , ~ : e y ,

J.

S.,

,,n,l

D. E Wii5Pn. 19S4l'>rc

b a ~ ~

nre

x.

rrork>l¡

: \ r T i c a ~ . Sinnc,ptc;<, 15·299-303

Jnh:v. :\. [ ._ R H. Fine, and 8. E \X'¡h;;m.

L9S5. Ram fnre5t } : . ; ; r , ~ a n c : n c ~ r : a í n future. Bat

;\:.;wo,

F:nn;;;

: : : n ~ l

Flnru

Pn;)t:tV/:t:nn

s.)déty,

Londnn. ) : 4 ~ 5

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t'.awnl

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en b

Are

c·ccrht 'en;m' l:

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u c e ~ i n n

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~ l : :

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d<:: s;-;n

'vbr

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lS:

L-102.

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icc:cr. phic

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niit>tcJred wctl:Jt'.d:l m the

l'cnn:<Jn

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nr.'n

Fc::est E c > l n ~ y

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s ~ ~ t c h : 2 C C L - Vcro:, :?.: \. L . t : e a u . & L

.;:_. t ~ . l e llrtez,L C t . , n t n S t t C t < ~ : <

:1

L: fl,--,ra

e L; ."\¡;,J,;,'1 'L:J

[ - \ . : ~ . ; : > : . < 1 - L t ~ A r f - . n l e ~ d

Ht:ttera..

V,,;_

i. Cn;:;se¡var<me

\'E Jar;__lm L ~ t G(;'nt:\T. S w i r : . . . : r b c ~ 1

\\\l-;(>t, D

E. 1 9 ~ 3 .

Be: faurus. A

r r c - · ~ + i c

C'11Hf'ctrh'fL S•:>tcmattc ?.2: i

4-

29.

625

bats as indicators of habitat disturbance_wilson_1996

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