a) in
opter
optera)
Chiropter
Movements
Mov
ements of bats (Mammalia, Chir
n Br
emnants in souther
est rremnants
or
Atlantic FFor
azil
Brazil
southern
orest
Gledson V. Bianconi
1, 2
; Sandra B. M ikich 3 & Wagner A. Pedro
2
1
Corresponding author. M ülleriana: Sociedade Fritz M üller de Ciências Naturais. Caixa Postal 19093, 81531-980 Curitiba,
Paraná, Brasil. E-mail: bianconi@terra.com.br
2
Laboratório de Chiroptera, Departamento de Apoio, Produção e Saúde Animal, Universidade Estadual Paulista.
Caixa Postal 341, 16050-680 Araçatuba, São Paulo, Brasil.
3
Laboratório de Ecologia, Embrapa Florestas. Caixa Postal 319, 83411-000 Colombo, Paraná, Brasil.
ABSTRACT. We used mark and recapture techniques to evaluate movements of bats within and between three
brazilian forest remnants. We captured bats with mist-nets in four 1 ha plots representing different degrees of
isolation of riparian (two plots) and submontane (two plots) forests between July 2002 and June 2003. Using
numbered aluminium tags, we marked 635 bats of seven species and 54 individuals of six species were recaptured. Overall, we recaptured Carollia perspicillata (Linnaeus, 1758) (short-tailed fruit bat) most frequently, especially in plots where they were banded in the riparian forest plots. These results suggest that this bat has
restricted feeding areas, which are probably determined by the abundance of Piper Linnaeus (Piperaceae), its
preferred food item. In contrast, species of the genus Artibeus Leach, 1821 exhibited few recaptures, suggesting high
mobility and larger feeding areas. In fact Artibeus seems to use more of the forest remnants in their search for
food, especially Ficus Linnaeus (Moraceae), the preferred food of this bat. Our results suggest that even small
forest isolates are valuable for the maintenance of some bat species because they offer many of the resources
they need or because they are spatially distributed in a pattern that allows use of the entire landscape.
KEY WORDS. Artibeus; Carollia perspicillata; foraging behavior; forest fragmentation; mark/recapture.
Floresta
morce
ceg
(Mammalia, Chir
RESUMO. Deslocamentos de mor
Chiropter
optera)
esta Atlântica no
ce
gos (Mammalia
opter
a) em remanescentes de Flor
sul do Br
asil. Nós utilizamos a técnica de marcação-recaptura para avaliar os deslocamentos de morcegos dentro
Brasil.
e entre três fragmentos florestais do sul do Brasil. Entre julho de 2002 e junho de 2003, os animais foram
capturados com redes-de-neblina instaladas em quatro parcelas de 1 ha, que representavam diferentes graus de
isolamento das subformações florestais, aluvial (duas parcelas) e submontana (duas parcelas). Utilizando anilhas
metálicas numeradas, nós marcamos 635 morcegos de sete espécies e recapturamos 54 indivíduos de seis espécies. A maior freqüência de recaptura foi obtida para Carollia perspicillata (Linnaeus, 1758), especialmente nas
parcelas de origem representadas pela floresta aluvial. Os resultados sugerem que este frugívoro apresenta uma
área restrita de forrageio, determinada provavelmente pela abundância de Piper L. (Piperaceae), planta reconhecida como seu alimento preferencial. Em contraste, espécies do gênero Artibeus Leach, 1821 exibiram uma baixa
freqüência de recaptura, sugerindo alta mobilidade e grande área de forrageio, provavelmente relacionada à
exploração conjunta dos fragmentos da região na busca por recursos, em especial Ficus L. (Moraceae), seu
alimento preferencial. Nossos resultados sugerem que mesmo pequenos fragmentos florestais são valiosos para a
manutenção de algumas espécies de morcegos, seja por oferecerem muitos dos recursos que eles necessitam ou
por estarem espacialmente dispostos de forma a facilitar a utilização conjunta da paisagem.
PALAVRAS-CHAVE. Artibeus; Carollia perspicillata; comportamento alimentar; fragmentação florestal; marcação e recaptura.
Fragm en tation an d isolation of n atu ral h abitats are th e
m ain th reats to region al an d global biod iversity (N OSS & C SUTI
1997) becau se th ey resu lt in several con seq u en ces to th e fau n a,
in clu d in g alteration at th e com m u n ity level an d local to region al extin ction s (LOVEJOY et al. 1984, ANDRÉN 1994, ROBINSON
& ROBINSON 1999, STRATFORD & STO UFFER 1999). In th is con text,
th e m obility of a p articu lar sp ecies h as m ajor im p lication s for
its su rvival (KOZAKIEW ICZ & SZACKI 1995, YABE & M ARQUES 2001),
besid es bein g eco lo gically relevan t becau se seed an d p o llen
d isp ersers can in flu en ce t h e sp at ial d ist ribu t io n an d gen et ic
Revista Brasileira de Zoologia 23 (4): 1199–1206, dezembro 2006
1200
stru ctu re of p lan ts (FENTON et al. 1992, G ARCIA et al. 2000, SLAUSON
2000). Som e p attern s of m ovem en ts rep resen t ecological an d
evolu tion ary resp on ses th at are excellen t tools for th e develop m en t of biological m odels (H EITHAUS & FLEMING 1978, AUGUST 1981,
M ERRIAM 1995, ANDRADE & M ARINI 2001, ESTRADA & C OATES-ESTRADA
2002). Th ese p attern s, wh ich are determ in ed by th e stru ctu re of
th e h abitat as well as by beh avioral p attern s of each sp ecies (G RAHAM 2001), are p articu larly n oteworth y in som e m am m alian
groups. As an exam ple, bats are th e on ly m am m als th at fly, wh ich
en ables th em to exp lore th e h abitat in a com p lex way (FENTON
1992), resu ltin g in regu lar local (sh ort d istan ce) an d season al
(lon g distan ce) m ovem en ts (TRAJANO 1996).
Som e au th ors su ggest th at bat m ovem en ts are closely related to th e availability of h abitat resou rces, esp ecially food an d
roosts (H EITHAUS & FLEMING 1978, FLEMING & H EITHAUS 1986, FLEMING
1991). Sin ce forest fragm en tation an d isolation ten d to redu ce
th e abu n dan ce an d to m odify th e sp atial distribu tion of th ese
resou rces in th e lan dscap e, foragin g p attern s an d h abitat u se by
bat sp ecies are m odified by th ese p rocesses (ESTRADA & C OATESESTRADA 2002, EVELYN & STILES 2003, Q UESADA et al. 2004).
Alth ou gh in th e last d ecad es a n u m ber of p ap ers on th e
m o v e m e n t s o f sm a ll m a m m a ls h a v e b e e n p u b lish e d (v.
KOZAKIEW ICZ & SZACKI 1995), su ch in form ation is u n kn own for
m ost bat sp ecies (M ARQUES 2003). In fact, even wh ere stu d ies
on th e m obility p attern s of Ch irop tera h ave been in p rogress
for m an y years (Costa Rica – H EITHAUS et al. 1975, LAVAL & FITCH
1977, H EITHAUS & FLEMING 1978, FLEMING & H EITHAUS 1986, FLEMING
1988, 1991; Mexico – M ORRISON 1978, ESTRADA et al. 1993, ESTRADA
& C OATES-ESTRADA 2002; Pan am a – BONACCORSO 1979, H ANDLEY et
al. 1991), th ere is still th e n eed for fu rth er in form ation in ord er to im p rove con servation efforts locally.
A sign ifican t p art o f t h e availab le lit erat u re regard in g
foragin g m ovem en ts, h abitat u se, m igration , an d h om e ran ge
refers to Old World bats (JONG 1994, RACEY 1998, W INKELMANN et
al. 2000, H OYLE et al. 2001). In th e New World , th ese stu d ies are
con cen trated in North Am erica (C LARK et al. 1993, BRIGHAM et
al. 1997, BETTS 1998, ESTRADA & C OATES-ESTRADA 2002) an d Cen t ral Am erica (LAVAL & FITCH 1977, H EITHAUS & FLEMING 1978,
H ANDLEY et al. 1991), wh ile in Sou th Am erica th ey h ard ly h ave
been d on e (bu t see LEMKE 1984, RUIZ et al. 1997, C OSSON et al.
1999). In cou n tries like Brazil, wh ich lacks basic in form ation
on th e n atu ral h istory of its bats (M ARINHO -FILHO & SAZIMA 1998),
available in form ation on bat m ovem en ts is even less exten sive. A few n otable excep tion s in clu d e TRAJANO (1996), wh o stu d ied bat m o vem en t s am o n g caves o f so u t h east ern Brazil an d
BERN ARD & FENTO N (2003), wh o in vest igat ed b at m o vem en t s
am o n g n at u ral fo rest fragm en t s in Cen t ral Am azo n ia. Ad d ition ally, th ere are isolated record s of th e d istan ces travelled by
sp ecies in Rio d e Jan eiro State (ESBÉRARD 2003).
Con sid erin g th e lack of in form ation on th e m ovem en ts
of bats am on g n on -Am azon ian forest fragm en ts an d th e n eed
of in form ation on local assem blages d u e to th e d ifferen cial resp on se of sp ecies d ep en d in g on p articu lar ch aracteristics of each
Revista Brasileira de Zoologia 23 (4): 1199–1206, dezembro 2006
G. V. Bianconi et al.
region , we d esign ed th e p resen t stu d y. Herein , we p resen t th e
first resu lts of a lon g-term to be bat ban d in g stu d y th at is bein g
con d u cted in Atlan tic Forest rem n an ts of Paran a State, sou th ern Brazil, in ord er to in vestigate h abitat u se an d m ovem en t
p attern s of sp ecies th at live in h igh ly fragm en ted forest h abit at s.
MATERIAL AND METHODS
Study area
Th e forest fragm en ts we stu d ied are located n ear th e town
of Fên ix, Paran á State, sou th ern Brazil (Fig. 1). Th e clim ate of
th e region is classified as m id -latitu d e h u m id su btrop ical or
Cfa (Köp p en ). Average an n u al tem p eratu re varies between 16
an d 25ºC (M IKICH & O LIVEIRA 2003) an d average an n u al rain fall
ran ges from 1 400 to 1 500 m m . Th e wet season occu rs between Decem ber an d March an d th ere is n o clearly d efin ed d ry
season (M AACK 1981). Th e stu d y region is covered by th e Atlan tic Forest, m ore sp ecifically by Sem id ecid u ou s Season al Forest
(sensu VELOSO et al. 1992), th at can be d ivid ed in to two typ es or
h abitats accord in g to th e d istan ce an d in flu en ce of large rivers. Th e allu vial or rip arian forest (RF) is associated with th e
rivers a n d is sea so n a lly o r o cca sio n a lly flo o d ed , a n d t h e
su bm on tan e forest (SF) occu p ies large areas away (> 300 m )
from th e m ain rivers an d is n ot d irectly in flu en ced by th em .
At p resen t, th e stu d y region , wh ich was on ce covered by
con tin u ou s forest, h old s few forest isolates u p to 800 h a su rrou n d ed by h u ge areas of agricu ltu re (esp ecially corn an d soybean ) an d largely d estroyed rip arian forests. Th e lim its of th e
forest fragm en ts with th e cu ltivated areas th at su rrou n d th em
are very abru p t, with n o bu ffer zon e, an d th e field s th em selves
h ave n o bu sh es or trees. A d etailed d escrip tion of th e gen eral
veget at io n o f t h e st u d y area, as well as sp ecies co m p o sit io n
an d p h en ological d ata, can be fou n d in M IKICH & SILVA (2001).
However, h ere it is im p ortan t to em p h asize th at som e ch aracteristics, as p lan t sp ecies com p osition an d p h en ology, typ e an d
in ten sity of p ast forest exp loration , p ast an d p resen t u se of soil
between th e fragm en ts (m atrix), are all sim ilar am on g th e stu d ied fragm en ts (M IKICH & SILVA 2001, M IKICH & O LIVEIRA 2003).
Sampling plots
Fou r sam p lin g p lots of 1 h a (100 x 100 m ) were establish ed in sid e th ree forest rem n an ts: two in th e Parq u e Estad u al
Vila Rica d o Esp írito San to (F1 an d F2), on e in Fazen d a Cagibi
(FC) an d on e in Fazen d a Gu aju vira (FG) (Fig. 1). Th e n earest
p lots were F1 an d F2, wh ich were 1.2 km ap art, an d th e m ost
d istan t p lots were F1 an d FC, wh ich were 4.9 km ap art.
Parq u e Est ad u al Vila Rica d o Esp írit o San t o (23º55’S,
51º57’W ) is a n at u re reserve su rro u n d ed by cu lt ivat ed areas
an d by th e Ivaí an d Coru m bataí rivers (Fig. 1). Most of its 354
h a are covered by on e of th e old est (m ore th an 370 years) secon d ary forests in Paran á State (C.V. RODERJAN , p ers. com m .) an d
is sim ilar in sp ecies com p osition an d stru ctu re to sligh tly d istu rbed p rim ary forests located in th e sam e region (M IKICH &
Movements of bats in Atlantic Forest remnants...
1201
coverage), 12-14 m h igh an d h as an exten sive n u m ber of lian as an d vin es over th e can op y trees. Th e u n d erstory h as 60%
coverage an d is 2 m h igh .
Capture and banding
Figure 1. Location of the study plots set inside three Atlantic Forest
rem nants at Fênix, Parana State, southern Brazil. Legend: 1. Parque Estadual Vila Rica do Espírito Santo (F1 and F2 plots), 2. Fazenda Cagibi (FC), and 3. Fazenda Guajuvira (FG).
SILVA 2001). Sin ce th is reserve is bord ered by two large rivers, it
h as exten sive areas of RF, in ad d ition to th e SF. Th ese two forest h abitats h ave a con tact zon e ap p roxim ately 3 km lon g. Th e
RF, wh ere F1 p lot is set, is ap p roxim ately 100 h a an d exten d s
in lan d for alm ost 300 m from th e Ivaí an d th e Coru m bataí
rivers. Th is p lot h as soil with d eficien t d rain age in som e p laces
an d is op en (30% coverage), with a 12 m h igh tree can op y. Th e
m ed iu m (5-6 m ) an d th e lower forest strata (1.2-2 m ) also are
n ot d en se (40 an d 30% coverage, resp ectively). Th e secon d p lot
(F2), set in SF, h as a d en ser (40%) an d h igh er (16-18 m ) can op y
with som e in d ivid u als reach in g 25 m , th e su b-can op y level is
also n ot d en se, with treelets u p to 6 m h igh .
Fazen d a Cagibi (23º55’S, 51º57’W ) con tain s 325 h a of
forest d ivid ed in to two blocks of d ifferen t sh ap es an d sizes (Fig.
1). It is su rrou n d ed by cu ltivated an d p astu re lan d s, an d ad jacen t to th e Ivaí River wh ich ru n s alon g a sm all p art of on e
fo rest b lo ck. Th ere are so m e sign s o f h u m an d ist u rb an ces,
m ain ly d u e to p ast tim ber exp loration (M IKICH & SILVA 2001).
Th e sam p lin g p lot (FC) was set in th e forest block (184 h a)
covered exclu sively by SF. Th is p lot h as a d en ser can op y (13-14
m , u p to 50% coverage) th an th e oth er on es, with several trees
u p to 20 m h igh , th e m ed iu m strata is ap p roxim ately 8 m h igh
wit h 50% co verage, t h e lo wer st rat a, wh ich is co m p o sed o f
bu sh es an d h erbs, ran ges from 0.7 to 1.3 m h igh .
Fazen d a Gu aju vira (23º53’S, 51º57’W ) is located between
th e oth er two stu d y sites an d is covered alm ost exclu sively by
secon d ary or d istu rbed p rim ary RF. It h as 24 h a d ivid ed in to
two p ortion s of sim ilar sh ap es an d sizes an d is bord ered by
cu ltivated lan d , p astu res, an d th e Ivaí River (Fig. 1). Th e forest
p ortion wh ere th e fou rth p lot (FG) was set is 14 h a, an d like
th e RF in th e Parq u e Vila Rica, it also exten d s for ap p roxim ately
300 m from th e Ivaí River. Th is p lot is relatively op en (30-40%
From Ju ly 2002 to Ju n e 2003 we n etted each p lot for two
n igh ts p er m on th . Th e bats were cap tu red u sin g 10 m ist-n ets
(12 x 2.5 m ), set in th e sam e p osition , en d to en d , in th e m id d le
of each p lot alon g a sin gle trail op en ed for th is work. Nets were
op en ed at su n set an d ch ecked every 20 m in u tes for six h ou rs.
Cap tu re effort d id n ot ch an ge am on g p lots, with 43 200 m 2 .h /
p lot (totallin g 172 800 m 2 .h ), calcu lated accord in g to STRAUBE
& BIANCONI (2002), i.e. m u ltip lyin g th e area of each n et by th e
tim e th ey rem ain exp osed in th e field by th e n u m ber of rep etition s by th e n u m ber of n ets. Followin g cap tu re, all bats were
id en tified , th eir forearm s m arked with n u m bered alu m in u m
rin gs (Etiq u etal ®), an d fin ally, released at th e cap tu re site. We
calcu lat ed t h e “m in im u m ” fligh t d ist an ces fo r each sp ecies
based on recap tu res in d ifferen t p lots an d th e lin ear d istan ces
between th em .
Sp ecies id en t ificat io n fo llo wed G O O D W IN & G REEN H ALL
(1961), VIZO TTO & TADDEI (1973), BARQ UEZ et al. (1993), an d
SIMMONS & VOSS (1998), an d th e taxon om ic arran gem en t followed SIMMONS (2005). Sp ecim en s th at cou ld n ot be id en tified
in th e field were collected , p reserved in alcoh ol (PETERSON 1965)
an d d ep o sit ed in t h e Mu seu d e Hist ó ria Nat u ral Cap ão d a
Im bu ia in Cu ritiba, Paran á State, Brazil.
RESULTS
Banding and recapture
We cap tu red an d ban d ed 635 in d ivid u als of seven bat
sp ecies of wh ich 54 in d ivid u als (8.5% of all ban d ed bats) of six
sp ecies were recap tu red . Most ban d ed in d ivid u als belon ged to
two sp ecies: Artibeus lituratus (Olfers, 1818) (great fru it-eatin g
bat) (387 m arked ) an d Carollia perspicillata (Lin n aeu s, 1758)
(sh ort-tailed fru it bat) (114), th e oth ers in clu d ed A. fim briatus
Gray, 1842 (frin ged fru it-eatin g bat), A. jam aicensis Leach , 1821
(jam aican fru it-eatin g bat), Sturnira lilium (E. Geoffroy, 1810)
(little yellow-sh ou ld ered bat), Chrotopterus auritus (Peters, 1856)
(woolly false vam p ire bat) an d Desm odus rotundus (E. Geoffroy,
1810) (com m on vam p ire bat) (Tab. I).
Table I. Banded and recaptured bat species between July 2002 and
June 2003 in four 1 ha Atlantic Forest plots located near the sm all
t ow n of Fênix, Paraná St at e, Brazil. (Al) Art ibeus lit urat us, (Af)
Art ibeus fimbriat us, (Aj) Art ibeus jamaicensis, (Sl) St urnira lilium ,
( Cp ) Ca r o l l i a p er sp i ci l l a t a , ( D r ) D esm o d u s r o t u n d u s, ( Ca)
Chrotopterus auritus.
Al
Banded individuals
Af
Aj
Sl
387 114 52
Cp
40
38
2
2
3
3
2
0
Recaptured individuals
11
30
5
Percentage of recapture (%)
2.8 26.3 9.6
7.5
Dr Ca
7.9 100 0
Revista Brasileira de Zoologia 23 (4): 1199–1206, dezembro 2006
1202
G. V. Bianconi et al.
Th e h igh est recap tu re rates were obtain ed for D. rotundus
(100%, n = 2) an d C. perspicillata (26.3%, n = 30). For th e oth er
sp ecies, excep t C. auritus (wh ich was n ot recap tu red ), valu es
ran ged from 2.8% for A. lituratus to 9.6% for A. fim briatus (Tab.
I). Th e r e w e r e o n ly t w o d o u b le r e ca p t u r e s, b o t h fo r C.
perspicillata an d in th e sam e ban d in g p lot (FG an d F1).
Recaptures per plot
Carollia perspicillata exh ibited th e h igh est p ercen tage of
in d ivid u als recap tu red in th e sam e p lot wh ere ban d in g took
p lace (18.7%), esp ecially in p lots located in th e RF (F1 an d FG)
(Tab. II). Oth er sp ecies exh ibited lower recap tu re rates for th e
p lots in wh ich in d ivid u als were first cap tu red .
Movements among plots
Seven teen m ovem en ts am on g p lots were recorded for five
sp ecies: A. lituratus (n = 5), A. fim briatus (n = 2), A. jam aicensis (n
= 1), C. perspicillata (n = 7) an d D. rotundus (n = 2). Fou r of th ese
m ovem en ts occu rred between th e two p lots located in side th e
Parq u e Vila Rica (F1 an d F2, A. fim briatus, A. jam aicensis, C.
perspicillata, D. rotundus) an d 13 am on g th e differen t forest fragm en ts, excep t between th e Fazen d a Cagibi an d th e Fazen d a
Gu aju vira (all sp ecies excep t D. rotundus). Based on th e lin ear
distan ces between p lots, we obtain ed th e distan ces travelled by
th e five recap tu red sp ecies. Artibeus lituratus sh owed th e h igh est
valu e (4.9 km ), followed by C. perspicillata an d A. fim briatus (3.7
km ), D. rotundus (1.6 km ) an d A. jam aicensis (1.2 km ).
DISCUSSION
We observed elevated recaptu re rates with in an d between
ou r sam p le areas for a few sp ecies du rin g on e year of sam p lin g.
Accordin g to FLEMING (1988), elevated recaptu re rates su ggest lim ited feedin g areas an d h igh site fidelity. Two species, D. rotundus
an d C. perspicillata, h ad h igh recap tu re rates in th e forest fragm en ts we stu died. Even th ou gh D. rotundus h ad th e h igh est rate,
we did n ot u se it in ou r com p arison s becau se of th e low n u m ber
of ban ded in dividu als (n = 2). Carollia perspicillata h ad th e h igh est recap tu re rate, esp ecially in th e ban din g p lots of th e rip arian
forest, su ggestin g th e h igh est fidelity to th e sam p le area of all
oth er recap tu red sp ecies. Given th at it was th e secon d m ost com m on sp ecies (114 in dividu als m arked), an d th e on e recap tu red
m ost (30 recap tu res), C. perspicillata seem s to be able to u se relatively sm all an d n arrow forest fragm en ts, like th e sam p led p ortion (14 h a) of th e Fazen da Gu aju vira.
Carollia perspicillata feed s m ain ly in th e m id level of th e
forest (“u n d erstory fru givores”, sensu BONACCORSO 1979) (FLEMING
1988, C OSSON et al. 1999, STOCKW ELL 2001) wh ere fru it-bearin g
sh ru bs su ch as th e gen u s Piper L. (Pip eraceae), its p refered food
item (BONACCORSO 1979, FLEMING 1988, PALMEIRIM et al. 1989, M ELLO
et al. 2004) are u su ally co m m o n . In fact , at least n in e Piper
sp ecies occu r in th e stu d y sites, an d six of th em are com m on
or abu n d an t in th e u n d erstory. Fu rth erm ore, d ifferen t sp ecies
of Piper in ou r stu d y area bear fru it seq u en tially so th at th is
resou rce is available all year rou n d (M IKICH & SILVA 2001). Based
Revista Brasileira de Zoologia 23 (4): 1199–1206, dezembro 2006
on th is in form ation , we sp ecu late th at th is bat sp ecies in ou r
stu d y area d oes n ot n eed to m ove far to fin d food , an d th erefore h as sm all feed in g areas.
Th is stron g association between th e feed in g beh avior of
C. perspicillata an d th e d istribu tion of food h as been exam in ed
in p revio u s st u d ies in Co st a Rica (H EITHAUS & FLEMING 1978,
FLEMING & H EITHAUS 1986, FLEMING 1991). Th ese au th ors su ggested
th at fru it availability is th e m ain factor in flu en cin g th e foragin g beh avior of th is bat sp ecies based on h igh freq u en cies of
recap tu re at th e sam e p laces. Becau se th e d iet of C. perspicillata
in clu d es several fru it sp ecies, esp ecially th ose of th e gen u s Piper,
t h ere is a co n t in u o u s rep lacem en t o f availabilit y an d abu n d an ce of fru it sp ecies locally (H EITHAUS & FLEMING 1978).
Even t h o u gh we kn o w t h at t h e fo rest fragm en t s h ave
n in e Piper sp ecies com m on ly d istribu ted alon g in n er road s an d
trails, as well as alon g forest ed ges (M IKICH & SILVA 2001), we d o
n ot yet kn ow wh eth er th e RF p lots h ad a h igh er d en sity of
th ese food resou rce com p ared to SF p lots. Con seq u en tly, we
can n ot con firm th at th e su p p osed p referen ce an d h igh er fid elity to th e RF by C. perspicillata is d u e to h igh er d en sities of
Piper. Fu tu re stu d ies m u st in clu d e both q u alitative an d q u an titative in ven tories of p oten tial food sp ecies in each forest typ e,
as well as th e collection of ad d ition al biological in form ation
on C. perspicillata in ord er to ch eck th is h yp oth esis.
For fru givores, if elevated recap tu re rates su ggest fid elity
to local areas (FLEMING 1988, PEDRO & TADDEI 1997), th e op p osite
sh ou ld be tru e wh en low rates are record ed . Com p ared to C.
perspicillata, all oth er recap tu red fru givore sp ecies h ad lower
recap tu res su ggestin g a p ossible h igh er m obility an d a larger
feed in g area. Artibeus lituratus, fo r exam p le, h ad t h e largest
n u m ber of ban d ed in d ivid u als (n = 387), bu t th e lowest recap tu re rate (2.8%), esp ecially in th e ban d in g p lots (1.6%). Con gen eric sp ecies, A. fim briatus an d A. jam aicensis, also exh ibited
low recap tu re rates in p lots in wh ich in d ivid u als were first cap tu red (Tab. II).
Sp ecies of th e gen u s Artibeus Leach , 1821 are con sidered
feedin g sp ecialists at large fru itin g trees (“can op y fru givores”,
sensu BONACCORSO 1979), esp ecially th e gen u s Ficus L. (Moraceae)
(PALMEIRIM et al. 1989, H ANDLEY et al. 1991, G ALETTI & M ORELLATO
1994). In fact, accordin g to M IKICH (2002), th ese fru its are largely
con su m ed by Artibeus sp p ., wh ich seem s to visit isolated Ficus
trees with abu n dan t fru it in th e stu dy area. Th e stu died fragm en ts h ave six sp ecies of Ficus, m ost rare or in term ediate, i.e.
rep resen ted by on e or few in d ivid u als accord in g to M IKICH &
SILVA (2001). So, if we con sider th at Ficus are u su ally low den sity
trees with low in ter-tree syn ch ron y in th e p rodu ction of large,
sh ort-lived fru it crop s (M ORRISON 1978, FLEMING & H EITHAUS 1981,
BONACCORSO & G USH 1987, C OSSON et al. 1999), it is likely th at th e
observed m ovem en ts of Artibeus sp p . are directly related to food
location s in differen t forest fragm en ts. In ou r stu dy site th ese
bats p robably in clu de differen t forest fragm en ts or h abitats in
th eir daily feedin g areas, in relation to th e p attern of den sity
an d distribu tion of fru its, as observed in oth er stu dies (ESTRADA et
Movements of bats in Atlantic Forest remnants...
1203
Table II. Percentage (and num ber) of individuals/ species recaptured in the sam e plot of banding between July 2002 and June 2003 in
Atlantic Forest fragm ents located near the small town of Fênix, Paraná State, Brazil. (Al) Artibeus lituratus, (Af) Artibeus fimbriatus, (Aj)
Artibeus jamaicensis, (Sl) Sturnira lilium , (Cp) Carollia perspicillata .
Recapture plot
(habitat type)
F1 (Riparian forest)
Bat species
Al
Af
Aj
Sl
Cp
1.1 (n = 1)
6.2 (n = 1)
10.0 (n = 1)
0 (n = 0)
24.2 (n = 8)
F2 (Submontane forest)
0.9 (n = 1)
7.1 (n = 1)
9.1 (n = 1)
0 (n = 0)
10.3 (n = 3)
FC (Subm ontane forest)
1.7 (n = 2)
7.1 (n = 1)
0 (n = 0)
14.3 (n = 2)
11.8 (n = 2)
FG (Riparian forest)
2.6 (n = 2)
0 (n = 0)
0 (n = 0)
25.0 (n = 1)
28.6 (n = 10)
1.6
5.1
4.8
9.8
18.7
M ean
al. 1993, SCHULZE et al. 2000, ESTRADA & C OATES-ESTRADA 2002, BER& FENTON 2003). M ORRISON (1978) rep orted an in crease in
th e distan ces travelled by A. jam aicensis on Barro Colorado Islan d wh en ever th e availability of m atu re Ficus fru its was low.
Alth ou gh S. lilium h ad th e secon d h igh est recap tu re rate
at th e ban d in g p lot, su ggestin g a p ossible fid elity to th e FG
p lot (wh ich was covered by RF, Tab. II), th e sm all n u m ber of
recap t u res d o es n o t allo w u s t o co n firm t h is h yp o t h esis. As
discu ssed for C. perspicillata, th e con tin u ation of th is stu dy cou ld
p rovid e fu rth er in form ation on th is su p p osed m ovem en t p attern an d / or h abitat p referen ce.
Regard in g th e “m in im u m ” recap tu re d istan ces rep orted
h ere, we em p h asize t h at su ch valu es d o n o t rep resen t d aily
m ovem en t p attern s (wh ich can on ly be obtain ed by rad iotelem et ry) o r t h e m axim u m d ist an ce a sin gle bat can fly, bu t
th ey d o con tribu te to th e kn owled ge of th e biology of th ese
bats in to th is an d oth er fragm en ted region s, as well as em p h asize th eir beh avioral flexibility in fragm en ted lan d scap es (oth er
region s: ESTRADA et al. 1993, W ILSO N et al. 1996, ESTRADA & C OATESESTRADA 2002, ou r region : BIANCONI et al. 2004). Exam in in g available d ata on th e lin ear d istan ces flown by d ifferen t bat sp ecies
(H EITHAUS & FLEMING 1978, TRAJANO 1996, C O SSO N et al. 1999,
ESBÉRARD 2003) as well as telem etry d ata (BERNARD & FENTON 2003),
we fou n d a wid e ran ge in p u blish ed d istan cies, wh ich is p robably related to d ifferen tial h abitat u se, in clu d in g sp atial an d
tem p oral d istribu tion of resou rces (BERNARD & FENTON 2003). As
an exam p le, we can m en tion th e valu es obtain ed in Brazil by
som e au th ors, su ch as ESBÉRARD (2003) for Rio d e Jan eiro State:
A. lituratus-21 km , A. fim briatus-25 km , D. rotundus-4.5 km an d
S. lilium -1.5 km , BERNARD & FENTON (2003) for n atu ral forest fragm en ts of Cen tral Am azon ia: C. perspicillata-1.6 to 2.5 km , an d
TRAJANO (1996) for São Pau lo State: D. rotundus-2.0 to 3.0 km .
Th is in form ation rein forces th e d yn am ic n atu re of th e exp loratory beh avior of bats, wh ich is th e p rod u ct of d ifferen t ecological an d evolu tion ary p ressu res.
In d istu rbed lan d scap es forest fragm en ts can be u sed by
bats as sh elters, as food sou rces or even as step p in g ston es to
reach n ew h abitats (ESTRADA & C OATES-ESTRADA 2002). In ou r stu d y
area, alth ou gh biologically im p overish ed if com p ared to in tact
fo rest s, t h e resu lt s su ggest t h at even sm all fo rest fragm en t s,
NARD
su ch as th e Fazen d a Gu aju vira, with 24 h a, are valu able for th e
m ain ten an ce of som e bat sp ecies. Th ese fragm en ts m ay offer
m an y of th e resou rces th e bats n eed or becau se th ey are sp atially d istribu ted in a p attern th at allows th e u se of th e en tire
lan d scap e, th ey m ay con tribu te to th e m ain ten an ce of local
an d region al biological d iversity.
ACKNOWLEDGEMENTS
We are in d ebted to th e FEMA/ IAP/ SEMA (con tract n º 024/
02) an d th e Mater Natu ra: In stitu to d e Estu d os Am bien tais for
th eir fin an tial su p p ort, to th e CNPq an d CAPES for gran ts given
to W.A.P. an d G.V.B. To th e FAPESP (p rocess 98/ 12556-1 an d
9 8 / 0 8 9 4 0 -0 ), t h e Em b rap a Flo rest as, t h e em p lo yees o f t h e
Parq u e Vila Rica d o Esp írit o San t o , an d t h e o wn ers o f t h e
Fazen d a Cagibi an d th e Fazen d a Gu aju vira, to Fabian a Roch a
Men d es, Dan iel C. Carn eiro, Carlos E. Con te, Arth u r Â. Bisp o
for field assistan ce; to Fábio Fogaça, Mich el Miretzki, Marlon
Zortéa, Lílian Casatti, Eleon ora Trajan o, an d Erica Sam p aio for
criticism an d su ggestion s on earlier d rafts. Also to Alberto Urben
Filh o for d rawin g th e m ap , an d to Bu rton K. Lim an d Don E.
Wilson for ed itorial assistan ce an d ad d ition al su ggestion s th at
im p roved th e m an u scrip t.
REFERENCES
ANDRADE, R.D. & M.A. M ARINI. 2001. Bird m ovem en t between
n atu ral forest p atch es in sou th east Brazil, p . 125-136. In :
J.L.B. ALBUQ UEQ UE; J.F. C ÂNDIDO JR; F.C. STRAUBE & A.L. ROOS
(Eds). Orn ito lo gia e co n servação : da ciên cia às estratégias. Tu barão, Ed itora Un isu l, 341p .
ANDRÉN , H. 1994. Effects of h abitat fragm en tation on birds an d
m am m als in lan dscapes with differen t proportion s of su itable
h abitat: a review. Oiko s, Copen h agen , 71 (3): 355-366.
AUGUST, P.V. 1981. Fig fru it con su m p tion an d seed d isp ersal by
Artibeus jam aicensis in th e Llan os of Ven ezu ela. Bio tro pica,
Wash in gton , 13 (1): 70-76.
BARQUEZ, R.M.; N.P. G IANNINI & M.A. M ARES. 1993. Guide to th e
bats o f Argen tin a. Guia de lo s m urcielago s de Argen tin a.
Oklah om a, Norm an , Oklah om a Mu seu m of Natu ral History,
VIII+119p .
Revista Brasileira de Zoologia 23 (4): 1199–1206, dezembro 2006
1204
BERNARD , E. & M.B. FENTON . 2003. Bat m obility an d roosts in a
fragm en ted lan d scap e in cen tral Am azon ia, Brazil. Bio tro pica, Wash in gton , 35 (2): 262-277.
BETTS, B.J. 1998. Roosts u sed by m atern ity colon ies of silverh aired bats in n orth eastern Oregon . Jo un al o f Mam m alo gy ,
Lawren ce, 79 (2): 643-650.
BIANCONI, G.V.; S.B. M IKICH & W.A. P EDRO . 2004. Diversid ad e d e
m orcegos (Mam m alia, Ch irop tera) em rem an escen tes florestais d o m u n icíp io d e Fên ix, n oroeste d o Paran á, Brasil.
Revista Brasileira de Zo o lo gia, Cu ritiba, 21 (4): 943-954.
BONACCORSO , F.J. 1979. Foragin g an d rep rod u ctive ecology in a
p an am an ian bat com m u n ity. Bulletin o f th e Flo rida State
Museum , Bio lo gical Scien ces, Gain esville, 24 (4): 359-408.
BONACCORSO , F.J. & T.J. G USH . 1987. Feed in g beh avior foragin g
strategies of cap tive p h yllostom id fru it bats: an exp erim en tal stu d y. Jo urn al o f An im al Eco lo gy , Oxford , 56 : 907-920.
BRIGHAM , R.M; M.J. VO NHO FF; R.M.R. BARCLAY & J.C. G W ILLIAM .
1997. Roostin g beh avior an d roost-site p referen ces of forestd wellen g Callifo rn ia bat s (Myotis californicus). Jo urn al o f
Mam m alo gy , Lawren ce, 78 (4): 1231-1239.
C LARK, B.S.; D.M. LESLIE JR & T.S. C ARTER. 1993. Foragin g activity
of adu lt fem ale big-eared bats (Plecotus townsendii) in Sum m er.
Journ al o f Mam m alo gy, Lawren ce, 74 (2): 422-427.
C O SSO N , J.; J. P O N S & D . M ASSO N . 1 9 9 9 . Effe c t s o f fo r e st
fragm en t at io n o n fru givo ro u s an d n ect arivo ro u s b at s in
Fren ch Gu ian a. Jo urn al o f Tro pical Eco lo gy , Cam brid ge,
15 (4): 515-534.
ESBÉRARD , C. 2003. Marcação e d eslo cam en t o s em m o rcego s.
Divulgaçõ es do Museu de Ciên cias e Tecn o lo gia UBEA/
PUCRS, Porto Alegre, 2 : 23-24.
ESTRADA, A. & R. C OATES-ESTRADA. 2002. Bats in con tin u os forest,
forest fragm en ts an d in a agricu ltu ral m osaic h abitat-islan d
at Los Tu xtlas, Mexico. Bio lo gical Co n servatio n , Essex, 103:
237-245.
ESTRADA, A.; R. C OATES-ESTRADA & D. M ERITT JR. 1993. Bats sp ecies
rich n ess an d abu n d an ce in trop ical rain forest fragm en ts
a n d in a gr ic u lt u r a l h a b it a t s a t Lo s Tu x t la s, M e x ic o .
Eco graph y , Cop en h agen , 16: 309-318.
EVELYN , M.J. & D.A. STILES. 2003. Roostin g req u irem en ts of two
fru givorou s bats (Sturnira lilium an d Artibeus interm edius) in
fragm en ted n eotrop ical forest. Bio tro pica, Wash in gton , 35
(3): 405-418.
FENTON , M.B. 1992. Bats. New york, Facts On File, XVI+207p .
FENTON , M.B.; L. ACHARYA; D. AUDET; M.B.C. H ICKEY; C. M ERRIMAN ;
M.K. O BRIST & D.M. SYME. 1992. Ph yllostom id bats (Ch iroptera:
Ph yllostom id ae) as in d icators of h abitat d isru p tion in th e
Neotrop ics. Bio tro pica, Wash in gton , 24 (3): 440-446.
FLEMING , T.H. 1988. Th e sh o rt-tailed fruit bat: a study in plan tan im al in teractio n s. Ch icago, Un iversity of Ch icago Press,
XIII+365p .
FLEMING , T.H. 1991. Th e relation sh ip between body size, diet, an d
h abitat u se in fru givorou s bats, gen u s Carollia (Ph yllostom atidae). Journ al o f Mam m alo gy, Lawren ce, 72 (3): 493-501.
Revista Brasileira de Zoologia 23 (4): 1199–1206, dezembro 2006
G. V. Bianconi et al.
FLEM IN G , T.H. & E.R. H EITH AUS. 1981. Fru givo ro u s b at s, seed
sh ad ows, an d th e stru ctu re of a trop ical forest. Bio tro pica,
Wash in gton , 13 (2): 45-53.
FLEMING , T.H. & E.R. H EITHAUS. 1986. Season al foran gin g beh avior
o f t h e fru givo ro u s b at Carollia perspicillata. Jo u rn al o f
Mam m alo gy , Lawren ce, 67 (4): 660-671.
G ALETTI, M. & L.P.C. M O RELLATO . 1994. Diet of th e large fru iteatin g bat Artibeus lituratus in a forest fragm en t in Brazil.
Mam m alia, Paris, 58 (4): 661-665.
G ARCIA, Q.S.; J.L.P. REZENDE & L.M.S. AGUIAR. 2000. Seed d isp ersal
by bats in a d istu rbed area of Sou th eastern Brazil. Revista
de Bio lo gia Tro pical, San Jose, 48 (1): 125-128.
G OODW IN , G.G. & A.M. G REENHALL. 1961. A review of th e bats of
Trin id ad an d To bago : d escrip t io n s, rabbies in fect io n an d
eco lo gy. Bu lletin o f th e Am erican Mu seu m o f Natu ral
Histo ry, New York, 122 (3): 187-302.
G RAHAM , C.H. 2001. Factors in flu en cin g m ovem en t p attern s of
Kell-billed Tou can s in a fragm en ted trop ical lan d scap e in
Sou th ern Mexico. Co n servatio n Bio lo gy, Boston , 15 (6):
1789-1798.
H ANDLEY JR, C.O.; D.E. W ILSON & A.L. G ARDNER. 1991. Dem ograp h y
a n d n a t u ra l h ist o r y o f t h e co m m o n fru it b a t , A rtibeus
jam aicensis, on Barro Colorado Islan d, Pan am á. Sm ith so n ian
Co n tributio n s to Zo o lo gy¸ Wash in gton , 511 : 1-173.
H EITHAUS, E.R & T.H. FLEMING . 1978. Foragin g m ovem en ts of a
fru givo ro u s b at , Carollia perspicillata (Ph yllo st o m id ae).
Eco lo gical Mo n o graph s, Lawren ce, 48 (2): 127-143.
H EITHAUS, E.R.; T.H. FLEMING & P.A. O PLER. 1975. Foragin g p attern s
an d resou rce u tilization in seven sp ecies of bats in a season al
Trop ical Forest. Eco lo gy , Du rh an , 56 (4): 841-854.
H OYLE, S.D.; A.R. P O PLE & G.J. TOOP. 2001. Mark-recap tu re m ay
reveal m ore abou t ecology th en abou t p op u lation tren d s:
d em o grap h y o f t h reat en ed gh o st b at (Macroderm a gigas)
p op u lation . Austral Eco lo gy , Carlton , 26 (1): 80-92.
JONG , J. 1994. Habitat u se, h om e ran ge an d activity p attern of
th e North ern Bat, Eptesicus nilssoni, in a h em iboreal forest.
Mam m alia, Paris, 58 (4): 535-548.
KOZAKIEW ICZ, M. & J. SZACKI. 1995. Movem en ts of sm all m am m als
in a lan d scap e: p atch restriction or n om ad ism ?, p . 78-94.
In : W.Z. LI D I C KER J R (Ed ). La n d s c a p e a p p ro a c h e s i n
m am m alian e c o l o g y an d c o n se rv atio n . M in n eap o lis,
Un iversity of Min n esota Press, IX+215p .
LAVAL, R.K. & H .S. FITC H . 1977. St ru ct u re, m o vem en t s an d
r e p r o d u ct io n in t h r e e C o st a Rica n b a t co m m u n it ie s.
Occasio n al Papers Museum o f Natural Histo ry, Un iversity
o f Kan sas, Lawren ce, 69: 1-27.
LEM KE, T.O . 1 9 8 4 . Fo ragin g eco lo gy o f t h e lo n g-n o sed b at ,
Glossophaga soricina, wit h resp ect t o resou rce availabilit y.
Eco lo gy , Tem p e, 65 (2): 538-548.
LOVEJOY, T.E.; J.M. RANKIN ; R.O. BIERREGAARD JR; K.S. BROW N JR; L.H.
EMMO NS & M.E. VAN DER VO O RT. 1984. Eco syst em d ecay o f
Am azon forest rem n an ts, p . 295-325. In: M.H. N ITECKI (Ed .).
Ex tin ctio n . Ch icago, Un iversity of Ch icago Press, 340p .
Movements of bats in Atlantic Forest remnants...
M AACK, R. 1981. Geo grafia física do Estado do Paran á. Rio d e
Jan eiro, J. Olym p io, XLIII+442p .
M ARIN H O -FILH O , J.S. & I. SAZIM A. 1 9 9 8 . Br a zilia n b a t s a n d
con servation biology: a first su rvey, p . 282-294. In: T.H. KUNZ
& P.A. RACEY (Ed s). Bat bio lo gy an d co n servatio n . Wash in gton , Sm ith son ian In stitu tion Press, XIV+365p .
M ARQUES, R.V. 2003. Migração d e m orcegos e o caso d e Tadarida
brasiliensis n o su l d o Brasil. Divulgaçõ es do Museu de Ciên cias e Tecn o lo gia UBEA/ PUCRS, Porto Alegre, 2 : 19-21.
M ELLO , M.A.R; G.M. SCHITTINI; P. SELIG & H.G. BERGALLO . 2004.
Season al variation in th e d iet of th e bat Carollia perspicillata
(Ch iro p t era: Ph yllo st o m id ae) in a At lan t ic Fo rest area in
sou th eastern Brazil. Mam m alia, Paris, 68 (1): 49-55.
M ERRIAM , G. 1995. Movem en t in sp atially d ivid ed p op u lation s:
resp on ses to lan d scap e stru ctu re., p . 64-77. In: W.Z. LIDICKER
JR (Ed). Lan dscape appro ach es in m am m alian eco lo gy an d
co n servatio n . Min n eap olis, Un iversity of Min n esota Press,
IX+215p .
M IKICH , S.B. & S.M. SILVA. 2001. Com p osição florística e fen ologia
d a s e sp é cie s zo o có r ica s d e r e m a n e sce n t e s d e Flo r e st a
Estacion al Sem id ecid u al n o cen tro-oeste d o Paran á, Brasil.
Acta Bo tân ica Brasilica, São Pau lo, 15 (1): 89-113.
M IKICH , S.B. 2002. A d ieta d os m orcegos fru gívoros (Mam m alia,
Ch irop tera, Ph yllostom id ae) d e u m p eq u en o rem an escen te
d e Floresta Estacion al Sem id ecid u al d o su l d o Brasil. Revista Brasileira de Zo o lo gia, Cu ritiba, 19 (1): 239-249.
M IKICH , S.B. & K.L. O LIVEIRA. 2003. Revisão do Plan o de Man ejo do Parque Estadual Vila Rica do Espírito San to . Cu ritiba,
Mater Natu ra/ FNMA, XXIII+452p .
M O RRISO N , D.W. 1978. In flu en ce o f h abit at o n t h e fo ran gin g
d istan ces of th e fru it bat, Artibeus jam aicensis. Jo urn al o f
Mam m alo gy , Lawren ce, 59 (3): 622-624.
N OSS, R.F. & B. C SUTI. 1997. Habitat fragm en tation , p . 269-304.
In: G.K. M EFFE & R.C. C ARROLL (Ed s). Prin ciples o f co n servatio n bio lo gy. Su n d erlan d , Sin au er Associates, 2 n d ed ., 729p .
P ALM ERIM , J.M .; D.L. G O RC H O V & S. STO LESO N . 1 9 8 9 . Tro p h ic
st ru ct u re o f a n eo t ro p ical fru givo re co m m u n it y: is t h ere
com p etition between bird s an d bats? Oeco lo gia, Berlin , 79
(3): 403-411.
PEDRO , W.A. & V.A. TADDEI. 1997. Taxon om ic assem blage of bats
from Pan ga Reserve, sou th eastern Brazil: abu n d an ce p attern s
an d trop h ic relation s in th e Ph yllostom id ae (Ch irop tera).
Bo letim do Museu de Bio lo gia Pro fesso r Mello Leitão ,
No va Série, San ta Teresa, 6 : 3-21.
PETERSON , B.L. 1965. Co llectin g bat specim en s fo r scien tific
purpo ses. Toron to, Un iversity of Toron to, 8p .
Q UESADA, M.; K.E. STONER; J.A. LOBO ; Y. H ERRERÍAS-D IEGO ; C. PALACIOSG UEVARA; M.A. M UNGUÍA-RO SAS; K.A. O -SALAZARL & V. RO SASG UERRERO . 2004. Effects of forest fragm en tation on p ollin ator
act ivit y an d co n seq u en ces fo r p lan t rep ro d u ct ive su ccess
an d m atin g p attern s in bat-p ollin ated bom bacaceou s trees.
Bio tro pica, Wash in gton , 36 (2):131-138.
RACEY, P.A. 1998. Ecology of eu rop ean bats in relation to th eir
1205
con servation , p . 249-261. In: T.H. KUNZ & P.A. RACEY (Eds).
Bat bio lo gy an d co n servatio n . Wash in gton , Sm ith son ian
In stitu tion Press, XVI+362p .
ROBINSON , W.D. & S.K. ROBINSON . 1999. Effects of selective loggin g
o n fo rest b ird p o p u lat io n s in a fragm en t ed lan d scap e.
Co n servatio n Bio lo gy , Boston , 13 (1): 58-66.
RUIZ, A.; M. SANTOS; P.J. SORIANO ; J. C AVELIER & A. C ADENA. 1997.
Relacion es m u tu alísticas en tre el m u rciélago Glossophaga longirostris y las cactáceas colu m n ares en la Zon a Arida de La
Tatacoa, Colom bia. Bio tro pica, Wash in gton , 29 (4): 469-479.
SCHULZE, M.D.; N.E. SEAVY & D.F. W HITACRE. 2000. A com p arison
o f t h e p h y llo st o m id b a t a sse m b la ge s in u n d ist u r b e d
n eotrop ical forest an d in forest fragm en ts of a Slash -an d Bu rn farm in g m o saic in Pet én , Gu at em ala. Bio tro p ic a ,
Wash in gton , 32 (1): 174-184.
SIMMONS, N.B. 2005. Ord er Ch irop tera, p . 312-529. In: D.E. W ILSON & D.M. REEDER (Ed s). Mam m al species o f th e Wo rld : a
tax o n o m ic an d geo graph ic referen ce. Balt im o re, Jo h n s
Hop kin s Un iversity Press, 3 rd ed ., 2142p .
SIMMO NS, N.B. & R.S. VO SS. 1998. Th e m am m als o f Paraco u ,
Fren ch Gu ian a: a Neotrop ical lowd lan d rain forest fau n a. Part
1. Bat s. Bu lletin o f th e Am erican Mu seu m o f Natu ral
Histo ry, New York, 237: 1-219.
SLAUSON , L.A. 2000. Pollin ation biology of two ch irop terop h ilou s
agaves in Arizon a. Am erican Jo urn al o f Bo tan y, Colu m bu s,
87 (6): 825-836.
STOCKW ELL, E.F. 2001. Morp h ology an d fligh t m an oeu vrability
in n ew world leaft-n osed bats (Ch irop tera: Ph yllostom id ae).
Jo urn al o f Zo o lo gy , Lon d on , 254 : 505-514.
STRATFO RD , J.A. & P.C . STO U FFER. 1 9 9 9 . Lo ca l ex t in ct io n s o f
terrestrial in sectivorou s bird s in a fragm en ted lan d scap e n ear
Man au s, Brazil. Co n servatio n Bio lo gy , Boston , 13 (6): 14161423.
STRAUBE, F.C. & G.V. BIANCONI. 2002. Sobre a gran d eza e a u n id ad e u tilizad a p ara estim ar esforço d e cap tu ra com u tilização
d e red es-d e-n eblin a. Ch iro ptera Neo tro pical, Brasília, 8 (12): 150-152.
TRAJANO , E. 1996. Movem en ts of cave bats in sou th eastern Brazil,
with em p h asis on th e p op u lation ecology of th e com m on
vam p ire bat, Desm odus rotundus (Ch irop tera). Bio tro pica,
Wash in gton , 28 (1): 121-129.
VELOSO , H.P.; L.C. O LIVEIRA-FILHO ; A.M.S.F. VAZ; M.P.M. LIMA; R.
M ARQ UETE & J.E.M. BRAZÃO . 1992. Man ual técn ico da vegetação brasileira. Rio d e Jan eiro, Secretaria d e Plan ejam en to, Orçam en to e Coord en ação, IBGE, 93p .
VIZO TTO , L.D. & V.A. TADDEI. 1973. Ch ave para determ in ação
de quiró ptero s brasileiro s. São José d o Rio Preto, Gráfica
Fran cal, 72p .
W ILSO N , D.E.; C.F. ASCORRA & S. SO LARI. 1996. Bats as in d icators
o f h ab it at d ist u rb an ce, p . 613-625. In: D.E. W ILSO N & A.
SANDOVAL (Ed s). Man u: Th e bio diversidade o f so uth eastern
Peru. Wash in gton , Sm ith son ian In stitu tion , 365p .
W INKELMANN , J.R.; F.J. BO NACCO RSO & T. STRICKLER. 2000. Ho m e
Revista Brasileira de Zoologia 23 (4): 1199–1206, dezembro 2006
1206
ran ge of th e Sou th ern blossom bat, Syconycteris australis, in
Pap u a New Gu in ea. Jo urn al o f Mam m alo gy, Paris, 81 (2):
408-414.
YABE, R.S. & E.J. M ARQ UES. 2001. Deslocam en to d e aves en tre
Received in 20.IV.2006; accepted in 17.XI.2006.
Revista Brasileira de Zoologia 23 (4): 1199–1206, dezembro 2006
G. V. Bianconi et al.
cap ões n o Pan tan al Mato-grossen se e su a relação com a d ieta, p . 103-123. In : J.L.B. ALBUQ UEQ UE; J.F. C ÂNDIDO JR; F.C.
STRAUBE & A.L. ROOS (Eds). Orn ito lo gia e co n servação : da
ciên cia às estratégias. Tu barão, Ed itora Un isu l, 341p .