JAN'
jl%H
COMPOSITION, MICROGEOGRAPHIC VARIATION,
AND SEXUAL DIMORPHISM IN
CAATINGAS AND CERRADO BAT COMMUNITIES
FROM NORTHEAST BRAZIL
MICHAEL R. WILLIG
NUMBER 23 PITTSBURGH, 1983
BULLETIN
o CARNEGIE MUSEUM OF NATURAL HISTORY
COMPOSITION, MICROGEOGRAPHIC VARIATION,
AND SEXUAL DIMORPHISM IN
CAATINGAS AND CERRADO BAT COMMUNITIES
FROM NORTHEAST BRAZIL
MICHAEL R. WILLIG
Pyma uning Labo a o y o Ecology,
Uni e si y o Pi sbu gh,
Lines il/e, Pennsyl ania 16424
(P esen Add ess: Depa men o Biological Sciences,
Texas Tech Uni e si y, Lubbock,
Texas 79409
)
NUMBER 23 PITTSBURGH, 1983
1
BULLETIN OF CARNEGIE MUSEUM OF NATURAL HISTORY
Numbe 23, pages 1-131, 8 igu es, 29 ables
Issued 30 Decembe 1983
P ice $14.00 acopy
Robe M. Wes , Di ec o
Edi o ial S a : Hugh H. Genoways, Edi o ; Duane A. Schli e , Associa e Edi o ; S ephen
L. Williams, Associa e Edi o ; Ma y Ann Schmid , Technical Assis an .
©1983 by he T us ees o Ca negie Ins i u e, all igh s ese ed.
CARNEGIE MUSEUM OF NATURAL HISTORY, 4400 FORBES AVENUE
PITTSBURGH, PENNSYLVANIA 15213
CONTENTS
Abs ac 5
In oduc ion 5
Gene al Desc ip ion o he Caa ingas 6
S udy Si es 9
Ma e ials and Me hods 13
Faunal Composi ion 13
Indi idual, Seconda y Sexual, and Geog aphic Va ia ion 14
Resul s and Discussion 15
Faunal Composi ion 15
Species Accoun s 17
Saccop e yx lep u a 17
Pe op e yx mac o is 17
Noc i/io lepo inas 17
P e ono us da yi
Mic onyc e is mega/o is .
Mic onyc e is minu a . . .
Tona ia bidens
Tonalia b asiliense
Tona ia sil icola
Mimon c enula um
Phyllos omus discolo ...
Phyllos omus has a us . . .
T achops ci hosus
Glossophaga so icina
Lonchophyl/a mo dax ...
Anou a geo oyi
Ca ollia pe spicilla a
S u ni a lilium
U ode ma magni os um
Vampy ops linea us
A libeus concolo
A ibeus jamaicensis
A ibeus li u a us
Desmodus o undas
Diphylla ecauda a
Na alus s amineus
Fu ip e us ba ens
Myo is nig icans
Ep esicus u inalis
Lasiu us bo ealis
Lasiu us ega
Molossops plani os is 1
Molossops emminckii 1
Tada ida la icauda a 1
Neopla mops ma og ossensis 1
Molossus a e 1
Molossus molossus 1
Eumops sp 1
Mo phome ies 1
00'0'CWWW’-NlslslslO'ONO UiUislslslUHOUi««00 0
—
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—
Mic ogeog aphic Va ia ion 124
Seconda y Sexual Va ia ion 125
Acknowledgmen s 127
Li e a u e Ci ed 128
ABSTRACT
Al hough he Sou h Ame ican opics con ains he mos di e se
chi op e an auna in he wo ld, mos in o ma ion conce ning ba s
om his egion is o a axonomic o dis ibu ional na u e. Fu -
he , mos o ou knowledge o Neo opical biology is es ic ed
o locales wi hin mesic p edic able biomes. The B azilian No h-
eas is an ex ensi e opical semia id egion cha ac e ized by
clima ic unp edic abili y and opog aphic he e ogenei y, ye i s
auna has ne e been s udied. An 18 mon h ield s udy o chi-
op e an popula ions in Caa ingas and Ce ado communi ies o
he No heas e ealed indi idual, seconda y sexual, and mic o-
geog aphic a ia ion in he ba auna. S udy si es we e es ic ed
in size o educe he p obabili y o sampling ba s om di e en
communi ies while in ensi e mon hly sampling egimes we e
es ablished o de ine he aunal composi ion o each communi y.
Despi e hei geog aphic p oximi y, he Caa ingas and Ce ado
con ain s ikingly di e en aunas bo h axonomically and eco-
logically. This dispa i y is mos clea ly exempli ied by he insec
-
i o e guilds in each biome. Two hypo heses a e p oposed o
accoun o he g ea e di e si y o he Caa ingas: 1) La ge mesic
encla es (se o es) a e common in he Caa ingas and p obably
insula e he auna om he ad e se e ec s o clima ic icissi udes,
whe eas g ea spa ial he e ogenei y p oduces dis inc habi a s o
di e en ial exploi a ion by di e en ba species, and 2) low insec
abundance and di e si y on he Chapada as well as educed can-
opy complexi y wi hin habi a s diminishes he numbe o insec-
i o e niches a ailable wi hin he Ce ado.
S a is ical analyses based on 14 ex e nal and 16 c anial mea-
su emen s showed ha many species wi h a leas mode a e sam-
ple sizes display sexual dimo phism, whe eas in e demic a ia-
ion be ween Caa ingas and Ce ado popula ions is somewha
a e . The applicabili y o a ious hypo heses conce ning sexual
dimo phism in ba s is e iewed in ligh o he obse a ion ha
emales a e la ge han males in many ba species. Signi ican
geog aphic a ia ion be ween adjacen popula ions o highly mo-
bile species is main ained in many ba species in he B azilian
No heas .
INTRODUCTION
The Ame ican opics con ains he iches mic o-
chi op e an auna in he wo ld (Pa e son and Pas-
cual, 1968). The subo de con ains 28.8% o all gen-
e a and 27.4% o all species o mammals ound in
he Neo opics (Walke , 1975). In he mo e a o -
able pa s o hei ange, ba s may domina e he
mammal auna in e ms o bo h densi y and species
ichness (He shko i z, 1972). Fi y- wo pe cen o
he Cos a Rican mammal auna is composed o ba s
(Robinson, 1971); simila ly, ba s ep esen 46% o
he Panamanian mammal auna (Flandley, 1966).
In sha p con as o he si ua ion in he Neo opics,
ba s ep esen only 12% o Nea c ic mammal species
(Walke , 1964). The dispa i y be ween no he n and
sou he n con inen s, and he high deg ee o ende-
mism in he Neo opical ba auna may be a ibu ed
in pa o biogeog aphic condi ions. Sou h Ame ica,
exis ing as an island om he C e aceous o la e
Cenozoic (Die z and Holden, 1970) would o e g ea
oppo uni y o chi op e an adia ion. The la e
connec ion o Sou h Ame ica wi h No h Ame ica
would accoun in pa o he ecen occupancy o
opical No h Ame ica by ba s, whe eas he sub-
sequen pauci y o mic ochi op e an species migh
be caused, o he mos pa , by he aga ies o em-
pe a e zone clima es.
The axonomic di e si y o Neo opical ba s is
pa alleled by asimila di e si y o eeding o ms.
The yea - ound abundance o ui s, seeds, nec a ,
insec s and o he animals p o ides sus enance o
ba s o di e se die a y specializa ions. Indeed, he
adap i e adia ion o ba s seems o ha e e ol ed
a ound he exploi a ion o many ypes o ood and
he pa i ioning o pa icula ood i ems by size
wi hin gene al ca ego ies (McNab, 1971; Smi h and
Genoways, 1974). Thus, Neo opical ba s occupy a
a ie y o di e en ood niches ( ugi o y, pisci o y,
nec a i o y, ca ni o y, insec i o y and sanguini-
o y), whe eas Nea c ic ba s a e p ima ily insec i -
o ous (McNab, 1971; Fleming e al., 1972).
Simpson (1964) was he i s o quan i y he in-
c ease in he numbe o mammal species as la i ude
dec eases om 60° Nla i ude o he equa o , while
MacA hu (1965) a emp ed o de ine he ac o s
a ec ing species di e si y o accoun o he in-
c eased ichness obse ed in he opics. Mo e e-
cen ly, J. Wilson (1974) has shown ha Simpson’s
o iginal obse a ions a e an a i ac caused by an
exponen ial inc ease in he numbe o ba species
wi h dec easing la i ude (see also Ma es and Ojeda,
1982). Despi e he signi ican con ibu ion o ba s
o Neo opical di e si y, ew wo ks o he han he
in es iga ions o Flemming e al. (1972), Thomas
(1972), and Hei haus e al. (1975) ha e conside ed
di e si y wi hin chi op e an communi ies. None-
heless, communi ies a e he con ex in which species
5
6BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
su i e and e ol e (Whi ake , 1975) and a i m
unde s anding o Neo opical di e si y g adien s is
p edica ed upon an adequa e delinea ion o he au-
nal composi ion o es ic ed a eas wi hin he Neo-
opics. Many su eys o ba aunas ha e been made
h oughou Sou h Ame ica, bu mos ha e deal wi h
la ge geopoli ical uni s (e.g. Husson, 1962; Tu le,
1970; Handley, 1976; Koopman, 1978). The auna
o b oad geog aphic egions is he sum o he a ious
communi ies ha hey con ain; howe e , he di -
e ences be ween communi ies wi hin apa icula
biome canno , in gene al, be asce ained om such
widesp ead collec ing p ocedu es.
Biomes as dis inc i e as he Caa ingas and Ce -
ado would be expec ed o ha bo unique aunas.
Fu he , due o he unp edic able and se e e cli-
ma ic condi ions cha ac e is ic o he Caa ingas, i
would be expec ed o suppo adiminished o de-
paupe a e auna when compa ed o Ce ado habi-
a s. The close p oximi y o hese biomes in he Exu-
C a o egion o B azil would e ec i ely elimina e
dispe sal as a ac o a ec ing he p esence o absence
o apa icula species and hus, clima ic o ecolog-
ical ac o s should emain as impo an limi ing
agen s.
Like he p e iously conside ed s udies o aunal
composi ion, chi op e an sys ema ic s udies usually
deal wi h mo phome ic a ia ion wi hin ela i ely
la ge a eas con aining anumbe o po en ially di -
e en popula ions. I is he e o e di icul o esol e
indi idual and in e demic a ia ion because sam-
ples om pa icula popula ions wi hin an a ea a e
uniden i ied o a e usually small in size and no
amenable o s a is ical analyses. Fu he , ew com-
p ehensi e s a is ical analyses o chi op e an a i-
a ion appea in he li e a u e; he wo ks o Taddei
(1975a, 1975b, 1979) a e an ob ious excep ion o
he Phyllos omidae. This s udy de ines and analyzes
indi idual, seconda y sexual, and geog aphic a i-
a ion o specimens om he B azilian No heas .
The a ia ion he ein epo ed o ba s om bo h
Caa ingas and Ce ado popula ions ep esen s ue
in ademic a ia ion and he s a is ical analyses ( wo-
way analysis o a iance) pe mi he isola ion o
bo h in e demic and seconda y sexual a ia ion, wi h
he subsequen assessmen o s a is ical signi icance
acili a ed o each ac o .
GENERAL DESCRIPTION OF THE CAATINGAS
The No heas o B azil con ains i e majo ege-
a ion zones (see Ma es e al., 1981, o ab ie de-
sc ip ion o each zone). Al hough Iwas able o make
limi ed su eys in h ee o hese (A lan ic Rain o -
es , Palm Fo es , and Caa inga-Ce ado Con ac
Zones), he esea ch conside ed he ein is es ic ed
o he Caa ingas and Ce ado habi a s, whe e Iwas
able o conduc in ensi e ield wo k.
The Caa ingas (Fig. 1) is an ex ensi e semia id
egion lying be ween 35° and 45° wes longi ude and
3° and 16° sou h la i ude (Reis, 1976). I occupies
app oxima ely 650,000 km 2(F o a-Pessoa e al.,
1971) and is cha ac e ized by ex eme empo al and
spa ial clima ic a iabili y (Ma kham, 1972; Ma k-
ham and McLain, 1977). Annual ain all may each
1,600 mm in some a eas, ye du ing d ough yea s
o he locali ies may no ecei e any p ecipi a ion a
all (Melo, 1956; Ma kham, 1972). Al hough cyclic
cha ac e is ics ha e been sugges ed (Ma kham, 1972,
1974, 1975; and Ma kham and McLain, 1977), mo e
ecen analyses u ilizing highly sophis ica ed s a is-
ical echniques indica e ha he pa e n o ain all
is uly andom. As such, he Caa ingas has been
called he “zone o calami y” (F eise, 1938) and he
“ egion o anomalous d ough ” (Ma kham, 1972).
S eilein (1981) has succinc ly summa ized he cli-
ma ic a ibu es o he Caa ingas. I is su icien he e
o no e ha he a iabili y and dis ibu ion o ain-
all in he Caa ingas is a ec ed by he in e ela ion-
ships be ween he Sou heas T ade Winds (xe ic in-
ducing agen s) and h ee la ge, mobile, mois u e-
laden ai masses— he equi o ial con inen al mass,
he In e opical Con e gence Zone, and he Sou h
A lan ic An icyclone. P ecipi a ion is u he a -
ec ed and diminished by he complex and i egula
pa e n o o og aphic ba ie s (Fig. 2).
Th ee majo geological elemen s accoun o he
ma ked opog aphic elie o he Caa ingas
(Ab’Sabe , 1970). The mos ex ensi e componen
is he basemen laye o p e-Camb ian c ys alline
ock which p oduces ex ensi e la lands and g adual
slopes. Nume ous g ani ic p o usions in he o m
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 7
Fig. 1. —The B azilian Caa ingas (delimi ed by he do ed line) occupies an a ea o o e 650,000 km2and is con ained wi hin nine s a es
o he No heas . Because o i s i egula shape and suscep ibili y o ex ended pe iods o d ough , he egion is known as "o poligono
das secas” ( he polygon o d ough ). Numbe s indica e he loca ion o Caa ingas (1) and Ce ado (2) s udy si es (modi ied om Ma es
e al„ 1981).
o low moun ain anges (se as), small moun ain
idges (se o es), o lowland ou c oppings (lajei os)
punc ua e he o he wise la su ace and app eciably
inc ease habi a complexi y wi hin he Caa ingas. In
a eas una ec ed by pos -C e aceous e osion, he
o iginal sands one subs a e s ill co e s he c ys al-
line basemen and p oduces la ge mesa-like pla eaus
o chapadas (James, 1942; Ab'Sabe , 1970). The
elemen al composi ion o hese sands one chapadas,
in conjunc ion wi h hei unique hyd ological p op-
14 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 1. —Desc ip ion o ex e nal and c anial cha ac e s mea-
su ed on specimens o ba s cap u ed in Exu, Pe nambuco (Caa-
ingas), and C a o, Cea d (Ce ado).
Ex e nal Cha ac e s
To al leng h: g ea es dis ance om he an e io mos po ion o
he snou o he dis al poin o he ail.
Tad leng h: g ea es dis ance om he dis al caudal e eb a o
he angle made by he ail when posi ioned pe pendicula o
he body.
Hind oo leng h: dis ance om he heel o he oo o he ip o
he longes oe including he claw.
Ea leng h: dis ance om he basal no ch o he ea o he u -
he mos poin on he edge o he pinna.
T agus leng h: dis ance om he base o he agus o i s dis al
edge.
Fo ea m leng h: dis ance om he ou side o he w is o he
ou side o he elbow when he wing is olded.
Weigh : weigh o he esh specimen.
Leng h o digi one: leng h om he w is o he dis almos poin
o he i s digi , including he claw.
Leng h o digi h ee: leng h om he w is o he dis al poin on
he phalange o digi h ee when he wing is maximally ex-
ended.
Leng h o digi ou : leng h om he w is o he dis al poin on
he phalange o digi ou when he wing is maximally ex ended.
Leng h o digi i e: leng h om he w is o he dis al poin on
he phalange o digi i e when he wing is maximally ex ended.
Tibia leng h: leng h om he ou e mos poin o he ankle o he
ou e mos poin o he knee.
Calca leng h: leng h om he dis al poin o he calca o he
angle made by he calca when i is posi ioned pe pendicula
o he leg.
Noselea leng h: leng h om he dis almos poin o he noselea
o i s junc u e wi h he os um.
C anial Cha ac e s
G ea es leng h o skull: dis ance om he mos an e io pa o
he os um (excluding ee h) o he pos e io mos poin o he
skull.
Condylobasal leng h: dis ance om he an e io mos edge o he
p emaxillae o he pos e io mos p ojec ion o he occipi al
condyles.
Zygoma ic b ead h: g ea es dis ance be ween he ou e ma gins
o he zygoma ic a ches.
Pos o bi al cons ic ion: leas dis ance ac oss he op o he skull
pos e io o he pos o bi al p ocess.
Mas oid b ead h: g ea es wid h o he skull, including he mas-
oid.
B ead h o he b aincase: g ea es wid h ac oss he b aincase pos-
e io o he zygoma ic a ches.
Ros al b ead h: wid h o he os um a he su u e be ween p e-
maxillae and maxillae.
Heigh o he b aincase: g ea es heigh o he b aincase om a
line pe pendicula o he long axis o he skull (+ 1mm).
B ead h ac oss he uppe mola s: maximum wid h om he ou e
al eolus o one mola o he ou e al eolus o ano he .
B ead h ac oss he uppe canines: wid h om he ou e al eolus
o one canine o he ou e al eolus o he o he canine.
Leng h o maxilla y oo h ow: leng h om he an e io edge o
he al eolus o he i s oo h p esen in he maxillae o he
pos e io edge o he al eolus o he las mola .
Table 1—Con inued.
Leng h o he uppe mola i o m oo h ow: maximum leng h om
he an e io edge o he al eolus o he i s cheek oo h o he
pos e io edge o he al eolus o he las mola .
Wid h o wides mola : wid h o wides mola in he maxilla
exlcuding he al eolus.
G ea es leng h o he mandible: leng h om he an e io mos
poin on he amus (excluding ee h) o he pos e io mos poin
on he co onoid p ocess.
Leng h o mandibula oo h ow: leng h om he an e io edge
o he al eolus o he canine o he pos e io edge o he al eolus
o he las mola in he mandible.
Leng h o co onoidp ocess: dis ance om he pos e io mos poin
on he co onoid p ocess o he base o he amus.
isi ed each mon h. S anda d Japanese mis ne s (10 mby 2m)
we e used o collec specimens. Al hough he posi ion o he ne s
was de e mined by peculia i ies o he e ain and he physiog-
nomy o he ege a ion, Iwas usually able o e ec 10 sec ions
o ne ing pe nigh in he mos equen ed collec ion si es. All
ne s we e posi ioned be o e dusk and checked a 15-min in e als
o soone depending upon he le el o ba ac i i y. Because ac-
i i y usually was qui e high a mos collec ion si es, he ne s
we e, in e ec , moni o ed con inuously. Ne s emained open o
aminimum o 3.5 o 4heach nigh . Ini ial ield wo k indica ed
ha addi ional ne ing was coun e -p oduc i e; o al ac i i y di-
minished d as ically a e 2100-2 130 hou s and he same species
caugh ea lie in he nigh we e caugh du ing la e ime pe iods.
Supplemen al collec ing om oos s ( o example, ca es, ee
hollows, buildings, cul e s, e c.) was also done in o de o e i y
ha he aunal composi ion was no biased by collec ing ech-
niques. Hal o he collec ion is housed in he Ca negie Museum
o Na u al His o y; he o he hal is deposi ed in he Museu de
Zoologia da Uni e sidade de Sao Paulo in B azil.
Indi idual, Seconda y Sexual, and
Geog aphic Va ia ion
When collec ions we e su icien ly la ge, 14 ex e nal cha ac-
e is ics and 16 c anial cha ac e is ics we e de e mined o a sam-
ple o 20 adul males and 20 adul emales om bo h Caa ingas
and Ce ado habi a s. O he wise he en i e collec ion om each
biome was used in subsequen analyses. Table 1lis s and de-
sc ibes all o he cha ac e s used h oughou he s a is ical anal-
yses. Ex e nal cha ac e s we e measu ed o he nea es millime e
u ilizing ame ic ule ; c anial cha ac e s we e measu ed wi h
me ic dial calipe s o he nea es hund ed h o amillime e .
Indi idual a ia ion was de e mined o males and emales
wi hin bo h Caa ingas and Ce ado communi ies. The s anda d
de ia ion (SD) and coe icien o a ia ion we e u ilized as s a-
is ical es ima es o indi idual a ia ion o each mensu al cha -
ac e .
Two-way analysis o a iance was u ilized o asce ain he
exis ence o geog aphic and seconda y sexual a ia ion in species
wi h su icien ly la ge samples o males and emales om bo h
Caa ingas and Ce ado biomes. When samples we e small o
one o bo h biomes, he da a we e combined and seconda y
sexual a ia ion was examined u ilizing one-way analysis o a i-
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 15
ance; his echnique also was u ilized o species ound only in
one biome. Le ene’s es o homogenei y o a iance was pe -
o med on all a iables in o de o de e mine he app op ia eness
o he analysis o a iance.
Sexual dimo phism o size is indica ed when apa icula sex
consis en ly has la ge sample means han he opposi e sex and
many o hose cha ac e s exhibi s a is ically signi ican sexual
a ia ion. I he ac ual di e ences be ween popula ion mean al-
ues is small, la ge samples a e equi ed o de ec s a is ical sig-
ni icance be ween sample means; in such cases whe e samples
a e small he consis en obse a ion o la ge mean alues o a
pa icula sex sugges s sexual dimo phism o size bu is incon-
clusi e un il la ge samples can be ob ained. Sexual dimo phism
o shape occu s when many cha ac e s exhibi s a is ically sig-
ni ican seconda y sexual a ia ion bu ob ious ends o mean
alue ela ions do no exis .
RESULTS AND DISCUSSION
Faunal Composi ion
O e 5,000 ba specimens ep esen ing 38 species,
29 gene a, and eigh amilies we e cap u ed du ing
his s udy; 65% o he mammalian species known
om he Exu-C a o a ea a e membe s o he Chi-
op e a. Table 2lis s he ba species om his s udy
in sys ema ic o de , indica es hei ela i e abun-
dance in Caa ingas o Ce ado biomes, and iden i-
ies hei eeding guild associa ions. Twen y species
a e sha ed be ween biomes; u he , he Caa ingas
con ains 13species no ound in he Ce ado, whe e-
as he Ce ado con ains i e species no ound in he
Caa ingas. I only he non- a e species in each biome
a e conside ed, he dissimila i ies be ween a eas be-
come mo e p onounced —15 o he 24 species (o e
hal o he species pool) occu exclusi ely in one o
he o he o he a eas. Despi e hei geog aphic p ox-
imi y, he Caa ingas and Ce ado habi a s con ain
ma kedly di e en ba aunas.
Con a y o my ea lie p edic ion, he Caa ingas
communi y is mo e species ich han he Ce ado
communi y. Amo e de ailed examina ion o he
dis ibu ion o ba s wi hin each communi y sugges s
an explana ion o his obse a ion. Faunal com-
posi ion and species densi ies a e a he uni o m
h oughou Ce ado habi a s on he Chapada do A -
a ipe. Con e sely, he Caa ingas is qui e he e oge-
neous in his ega d. Caa inga Baixa con ains ew
species o ba s, and hose species p esen occu a
low densi ies. Lajei os con ain a ew addi ional
species (Neopla ymops ma og ossensis and Pe op-
e yx mac o is), bu in gene al, he auna o he Caa-
ingas in low-lying a eas (Caa inga Baixa and la-
jei os) is depaupe a e. Species o oliage gleaning
insec i o e (Mimon c enula um, Mic onyc e is
megalo is, Mic onyc e is minu a, and Tona ia b as-
iliense) each hei highes densi y and occu almos
exclusi ely in Caa inga Al a. Wi h ew excep ions,
howe e , se o es ha bo he bulk o he species
ound in he Caa ingas and i is on se o es o he
adjacen a eas o Caa inga Al a ha mos species
each hei highes densi ies. Thus, he ela i ely
high species ichness o he Caa ingas can be a ib-
u ed in pa o he opog aphic elie and ege a-
ional di e si y o he Caa ingas, bu equally im-
po an om he poin o iew o he Chi op e a,
he nume ous se o es punc ua ing he la landscape
o he Caa ingas p o ide oos ing si es and mesic
e ugia du ing d ough pe iods.
F ugi o e guilds in he Caa ingas and Ce ado a e
qui e simila in e ms o species ichness and axo-
nomic composi ion. The main dis inc ion among
he guilds is he g ea e densi y o all ugi o ous
species on he Chapada and he p esence he e o
an addi ional small species, A. concolo . The p es-
ence o L. mo dax in he Caa ingas is he mos
ob ious di e ence be ween he biomes wi h espec
o nec a i o es. Nei he ugi o es no nec a i o es
app eciably a ec he aunal di e ences be ween he
Caa ingas and Ce ado.
The no able absence o sanguini o es and pisci-
o es om among he common Ce ado ba s di-
minishes he di e si y o ha auna. La ge na i e
mammals a e absen om he Chapada and do-
mes ic species a e p ohibi ed by law om en e ing
he Flo es a Nacional A a ipe-Apodi. Thus, i is no
su p ising o ind ampi es a e in Chapada habi-
a s. The absence o s anding bodies o wa e om
he Chapada accoun s o he a eness o pisci o es.
The dispa i y be ween he Caa ingas and Ce ado
ba aunas is mos clea ly mani es ed wi hin he
a ious insec i o e guilds. The Caa ingas ha bo s a
di e se insec i o e auna; i e species o oliage
gleaning insec i o es, wo species o ae ial insec i-
o es, and wo species o molossid insec i o es each
app eciable densi ies he e. In con as , M. ni-
g icans, E. u inalis, and M. molossus a e he only
insec i o ous ba s ha a ain app eciable densi ies
16 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 2.— Sys ema ic lis ing o ba s om Caa ingas (Exu, Pe nambuco) and Ce ado (C a o, Cea /S) biomes; Aindica es abundan , C
indica es common, Rindica es a e, and— indica es absen . Feeding guild abb e ia ions: AERIN, ae ial insec i o e: PISCI, pisci o e;
FOLGL, oliage-gleaning insec i o e: OMNIV, omni o e;NECTA, nec a i o e; FRUGI, ugi o e;SANGU, sanguini o e: MOLOS,
molossid ae ial insec i o e.
Species
P esence
Caa ingas Ce ado Guild
Family Emballonu idae
Saccop e yx lep u a RAERIN
Pe op e yx mac o is c-AERIN
Family Noc ilionidae
Noc ilio lepo inus cRPISCI
Family Mo moopidae
P e ono us da yi RRAERIN
Family
Sub amily
Phyllos omidae
Phyllos ominae
Mic onyc e is megalo is R-C RFOLGL
Mic onyc e is minu a R-C RFOLGL
Tona ia bidens R—FOLGL
Tona ia b asiliense R-C —FOLGL
Tona ia sil icola C—FOLGL
Milmon c enula um R-C —FOLGL
Phyllos omus discolo R-C AOMNIV
Phyllos omus has a us RAOMNIV
T achops ci hosus C-OMNIV
Sub amily Glossophaginae
Glossophaga so icina AANECTA
Lonchophylla mo dax C—NECTA
Anou a geo oyi RCNECTA
Sub amily Ca olliinae
Ca ollia pe spicilla a AAFRUGI
Sub amily S enode ma inae
S u ni a lilium RC-R FRUGI
U ode ma magni os um RRFRUGI
Vampy ops linea us AAFRUGI
A ibeus concolo —C-R FRUGI
A ibeus jamaicensis CAFRUGI
A ibeus li u a us C-R AFRUGI
Sub amily Desmodon inae
Desmodus o undus ARSANGU
Diphylla ecauda a R-SANGU
Family Na alidae
Na alus s amineus _RAERIN
Family Fu ip e idae
Fu ip e us ho ens RAERIN
Family Vespe ilionidae
Myo is nig icans C-A CAERIN
Ep esicus J'u inalis -C-R AERIN
Lasiu us bo ealis —RAERIN
Lasiu us ega RRAERIN
Family Molossidae
Molossops plani os is RMOLOS
Molossops emminckii RRMOLOS
Tada ida la icauda a RRMOLOS
Neopla ymops ma og ossensis C-MOLOS
Molossus a e R—MOLOS
Molossus molossus AAMOLOS
Eumops sp. R-MOLOS
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 17
on he Chapada do A a ipe, and hei p esence is
p ima ily es ic ed o dis u bed a eas con aining
abandoned buildings. None o he common Ce ado
insec i o es glean hei p ey om oliage. The sa-
anna-like cha ac e is ics o he Chapada ( ha is,
open spaces, low canopy, educed e ical s a i i-
ca ion wi hin he canopy) as well as he depaupe a e
na u e o he Ce ado insec auna limi he po en ial
numbe o insec i o e niches a ailable o ba s o
exploi and o a la ge ex en accoun s o he de-
paupe a e na u e o he Ce ado ba auna in gene al.
Species Accoun s
Family Emballonu idae
Saccop e yx lep u a Sch ebe , 1774
This ae ial insec i o e was absen om he Caa-
ingas and was qui e a e on he Chapada do A -
a ipe. Fu he , he speci ic designa ion o S. bili-
nea a was applied o bo h specimens o S. lep u a
om he Chapada do A a ipe by Ma es e al. (1981).
Ve y li le is known abou he biology o S. lep u a
and only isola ed eco ds o mensu al cha ac e s
appea in he li e a u e. S a is ical analyses could
no be pe o med on he sample o wo indi iduals
ob ained om he Chapada do A a ipe; hence only
selec ed indi idual measu emen s (a e Swanepoel
and Genoways, 1979) a e epo ed he e o an adul
emale and an adul male ( o al leng h, 61, 69; ail
leng h, 17, 14; hind oo leng h, 6, 6; ea leng h, 13,
13; g ea es leng h o skull, —,13.3; condylobasal
leng h, —,121; zygoma ic b ead h, —,—;pos o -
bi al cons ic ion, 2.3, 2.3; b ead h o b aincase, 6.8,
6.9; leng h o he maxilla y oo h ow, 5.4, —;b ead h
ac oss he uppe mola s, 6.2, —).
Pe op e yx mac o is (Wagne , 1843)
O he h ee subspecies p esen ly ecognized, only
P. m. mac o is occu s on he mainland o Sou h
Ame ica (Cab e a, 1957). Al hough uncommon in
he Caa ingas, his ae ial insec i o e was ound
oos ing in small agg ega ions o up o 10 indi id-
uals in la ge openings inside ockpiles o cul e s.
O e a6mon h pe iod, he numbe o indi iduals
in each o wo moni o ed oos ing si es emained
cons an . Fu he , each oos con ained only one
adul male. This sugges s ha P. m. mac o is males
migh main ain small ha ems and exhibi esou ce
de ense polygony when app op ia e oos ing con-
di ions a e a ailable. Dis ances be ween neighbo -
ing indi iduals in he oos a ied om 15 o 60
cm. Indi iduals oos ed by ancho ing hei ee and
humbs o he ceiling o he e ea si e; hence, hei
bodies assumed an acu e angle wi h espec o he
o e lying ock s a um. These ba s we e equen ly
obse ed lea ing hei oos s be o e dusk. The ab-
sence o P. m. mac o is om he Chapada do A -
a ipe can mos easily be explained by he lack o
sui able oos ing si es.
The esul s o he s a is ical analyses o 13ex e nal
and 16 c anial cha ac e s on specimens om he
Chapada do A a ipe a e summa ized in Table 3.
Sample means o emales a e la ge han sample
means o males o 10 ex e nal cha ac e s and 15
c anial cha ac e s; s a is ically signi ican di e ences
exis o o al leng h, agus leng h, o ea m leng h,
leng h o digi III, leng h o digi V, condylobasal
leng h, b ead h ac oss he uppe mola s, b ead h
ac oss he uppe canines, and g ea es leng h o he
maxilla y. The sample means o males a e la ge
han hose o emales o h ee ex e nal and one
c anial cha ac e ; none exhibi s a is ically signi i-
can di e ences. Sexual dimo phism wi h emales
la ge han males is clea ly suppo ed by he da a.
Family Noc ilionidae
Noc ilio lepo inus (Linnaeus, 1758)
O he h ee subspecies o his pisci o e cu en ly
ecognized, Noc ilio l. lepo inus is he app op ia e
designa ion o specimens om he No heas o
B azil (Da is, 1973). I was excep ionally a e on
he Chapada do A a ipe; he ew indi iduals caugh
he e we e p obably ansien s om su ounding
habi a s. I was common in he Caa ingas whe e i
oos ed du ing he day in g oups o up o 30 indi-
iduals. The la ge hollow ha dwood ees ypically
ound in Caa inga Al a habi a s p o ided i s p e-
e ed diu nal oos ing si es. Sex a ios did no sig-
ni ican ly di e om one o one based upon o e all
ne ing eco ds o da a om indi idual day oos s
(Binomial Tes , P>.05). App oxima ely 47% o
he 328 cap u ed adul specimens om he Caa in-
gas we e males.
Indi iduals equen ly o aged a small lakes and
ponds in g oups con aining i e o 15ba s. Indi id-
uals seemed o coo dina e eeding ac i i ies and
concen a ed o aging ac i i y on he pe iphe y o
bodies o wa e . Aconc e e b idge which a e sed
asemipe manen s eam con ained anoc u nal
eeding oos ha was used by o e 150 indi iduals.
This oos was in con inual use o he du a ion o
18 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 3. —Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, N) o/Pe op e yx mac o is males and emales om he Caa inga biome. Aone-way analysis o
a iance (Mode! 1) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual a ia ion is indica ed
by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
<3.5 22 <3<3 22 Fac o d MS FSigni icance
To al leng h
Mean 61.33 64.14 Sex 137.673 5.30 .032
SD 1.99 3.81 Wi hin 20 7.110
SE .51 1.44
CV 3.24 5.94
n15 7
Tail leng h
Mean 14.20 14.14 Sex 1.016 .00 .945
SD 1.61 2.12 Wi hin 20 3.163
SE .42 .80
CV 11.34 14.99
n15 7
Hind oo leng h
Mean 6.60 6.57 Sex 1.004 .01 .919
SD .51 .79 Wi hin 20 .366
SE .13 .30
CV 7.73 12.02
n15 7
Ea leng h
Mean 14.20 14.43 Sex 1.249 .49 .491
SD .56 .98 Wi hin 20 .506
SE .15 .37
CV 3.94 6.79
n15 7
T agus leng h
Mean 6.13 6.71 Sex 11.61
1
6.25 .021
SD .52 .49 Wi hin 20 .258
SE .13 .18
CV 8.48 7.30
n15 7
Fo ea m leng h
Mean 42.00 43.57 Sex 111.786 13.31 .002
SD .85 1.13 Wi hin 20 .886
SE .22 .43
CV 2.02 2.59
n15 7
Weigh
Mean 4.23 4.57 Sex 1.546 1.08 .312
SD .59 .93 Wi hin 20 .507
SE .15 .35
CV 13.95 20.35
n15 7
Leng h o digi one
Mean 7.53 7.14 Sex 1.728 1.00 .330
SD .74 1.07 Wi hin 20 .730
SE .19 .40
CV 9.83 14.99
n15 7
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 19
Table 3. —Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 92 <3<5 22 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 64.67 68.00 Sex 153.03 5.91 .025
SD 2.38 4.08 Wi hin 20 8.97
SE 62 1.54
CV 3.68 6.00
n15 7
Leng h o digi ou
Mean 45.80 47.43 Sex 112.658 3.01 .098
SD 1.37 3.10 Wi hin 20 4.206
SE .36 1.17
CV 2.99 6.54
n15 7
Leng h o digi i e
Mean 44.40 46.86 Sex 128.82 12.41 .002
SD 1.35 1.86 Wi hin 20 2.32
SE .35 .71
CV 3.04 3.97
n15 7
Tibia leng h
Mean 18.80 19.14 Sex 1.561 1.21 .284
SD .56 .90 Wi hin 20 .463
SE .15 .34
CV 2.98 4.70
n15 7
Calca leng h
Mean 15.80 16.14 Sex 1.561 .34 .568
SD 1.27 1.35 Wi hin 20 1.660
SE .33 .51
CV 8.04 8.36
n15 7
G ea es leng h o skull
Mean 13.75 13.99 Sex 1.258 3.67 .070
SD .30 .17 Wi hin 20 .070
SE .08 .06
CV 2.18 1.22
n15 7
Condylobasal leng h
Mean 12.74 13.03 Sex 1.397 5.00 .037
SD .25 .35 Wi hin 20 .080
SE .06 .13
CV 1.96 2.69
n15 7
Zygoma ic b ead h
Mean 8.15 8.3! Sex 1.134 3.91 .480
SD .16 .23 Wi hin 20 .034
SE .04 .09
CV 1.96 2.77
n15 7
20 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 3. —Con inued.
Caa inga Ce ado Analysis o a iance
66 92 <5<5 29 Fac o d MS FSigni icance
Pos o bilal cons ic ion
Mean 2.47 2.50 Sex 1.003 .15 .702
SD .15 .14 Wi hin 20 .023
SE .04 .05
CV 6.07 5.60
n15 7
Mas oid b ead h
Mean 7.25 7.27 Sex 1.003 .08 .786
SD .13 .30 Wi hin 20 .039
SE .03 .12
CV 1.79 4.13
n15 7
B ead h o b aincase
Mean 6.49 6.49 Sex 1.000 .00 .989
SD .16 .09 Wi hin 20 .020
SE .04 .03
CV 2.47 1.39
n15 7
Ros al b ead h
Mean 5.97 6.15 Sex 1.134 2.45 .135
SD .23 .24 Wi hin 18 .055
SE .06 .10
CV 3.85 3.90
n 14 6
Heigh o b aincase
Mean 7.45 7.50 Sex 1.009 .16 .692
SD .26 .18 Wi hin 19 .058
SE .07 .07
CV 3.49 2.40
n15 6
B ead h ac oss he uppe mola s
Mean 6.02 6.48 Sex 1.896 13.53 .002
SD .21 .35 Wi hin 18 .066
SE .06 .14
CV 3.49 5.40
n14 6
B ead h ac oss he uppe canines
Mean 3.47 3.67 Sex 1.160 6.52 .020
SD .14 .19 Wi hin 18 .025
SE .04 .08
CV 4.03 5.18
n14 6
Leng h o he maxilla y oo h ow
Mean 5.49 5.53 Sex 1.007 .16 .695
SD .18 .27 Wi hin 19 .043
SE .05 .11
CV 3.28 4.88
n15 6
Leng h o he uppe mola i o m oo h ow
Mean 4.01 4.10 Sex 1.032 1.40 .252
SD .16 .14 Wi hin 19 .023
SE .04 .06
CV 3.99 3.41
n15 6
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 21
Table 3.—Con inued.
Caa inga Ce ado Analysis o a iance
<5(3 99 <5<5 29 Fac o d MS FSigni icance
Wid h o he wides mola
Mean 1.73 1.78 Sex 1.01 12.49 .131
SD .06 .08 Wi hin 19 .004
SE .02 .03
c 3.47 4.49
n15 6
G ea es leng h o he mandible
Mean 9.51 9.91 Sex 1.747 8.03 .01 1
SD .27 .37 Wi hin 19 .093
SE .07 .14
CV 2.84 3.73
n14 7
Leng h o he mandibula oo h ow
Mean 5.64 5.77 Sex 1.077 1.61 .219
SD .17 .29 Wi hin 19 .048
SE .05 .11
CV 3.01 5.03
n14 7
Leng h o he co onoid p ocess
Mean 2.47 2.63 Sex 1.104 2.89 .109
SD .19 .20 Wi hin 16 .036
SE .06 .08
CV 7.69 7.60
n117
he s udy and had p obably been used p e iously
o an ex ended pe iod (based upon he quan i y o
accumula ed eces).
The Caa ingas collec ion ep esen s he la ges
analyzed sample o Noc ilio lepo inus om asingle
eely in e b eeding popula ion (see Da is, 1973).
The esul s o as a is ical analysis o 20 mensu al
mo phological cha ac e s in N. 1. lepo inus a e sum-
ma ized in Table 4. Like he esul s epo ed by
Da is (1973) o specimens om he Paci ic e san
o Chiapas, Mexico (N. 1. mas i us), males end o
be mo e a iable han emales wi h consis en ly
la ge coe icien s o a ia ion. The sample means
o males a e la ge han sample means o emales
o all 13ex e nal cha ac e s analyzed, wi h s a is-
ically signi ican di e ences ound o 10 o he
a iables. Males a e s a is ically signi ican ly di e -
en han, and la ge on he a e age han, emales
o 15 o he 16 c anial cha ac e s. Al hough he
sample mean o emales is la ge han ha o males
o he single cha ac e wid h o he wides mola ,
he di e ence is no s a is ically signi ican . Noc ilio
l. lepo inus exhibi s ex eme sexual dimo phism wi h
males consis en ly la ge on he a e age han e-
males.
Da is (1973) epo ed mo phome ic da a om
h ee locali ies (Uppe Amazon Basin, Lowe Am-
azon Basin, and he Guianas) wi hin he ange o
N. 1. lepo inus. This acili a es compa ison wi h he
Caa ingas popula ion which is he no heas e n-mos
collec ion locale o he species o which ex ensi e
mo phome ic da a a e a ailable. The Caa ingas
popula ion has la ge mean alues o ou o he
i e mo phome ic cha ac e s epo ed by Da is
(1973) and as such ep esen s he mos obus pop-
ula ion wi hin he subspecies. Fu he , among pop-
ula ions o N. I. lepo inus, he Caa ingas specimens
a e mo phome ically mo e simila o specimens
om he Uppe Amazon Basin han o o he pop-
ula ions.
Family Mo moopidae
P e ono us da yi G ay, 1838
This ae ial insec i o e was a e in bo h Caa ingas
and Ce ado biomes. O he h ee subspecies cu -
en ly ecognized, P.d. da yi is he appella ion ap-
p op ia e o specimens om he No heas o B azil
(Smi h, 1972).
Smi h (1972) p esen ed an analysis o sys ema ic
11 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 4. —Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o/’Noc ilio lepo inus males and emales om bo h Caa inga and Ce ado biomes. Due o
small sample sizes, da a om bo h biomes we e combined o de e mine he exis ence o signi ican (P< .050) seconda y sexual a ia ion
ia aone-way analysis o a iance (Model I) wi h eplica ion.
Caa inga Ce ado Analysis o a iance
<5<5 22 <5<5 22 Fac o d MS FSigni icance
To al leng h
Mean 123.35 116.50 120.00 116.75 Sex 1495.1 14 27.46 <.001
SD 3.84 4.57 —5.38 Wi hin 43 18.027
SE .86 1.02 —2.69
CV 3.1 13.92 —4.61
n20 20 14
Tail leng h
Mean 27.60 25.30 28.00 25.75 Sex 156.400 11.52 .002
SD 2.54 2.06 —1.50 Wi hin 43 4.857
SE .57 .46 —.75
CV 9.20 8.14 —5.83
n20 20 14
Hind oo leng h
Mean 28.60 27.25 30.00 27.00 Sex 123.819 15.85 <.001
SD 1.27 1.16 —1.41 Wi hin 43 1.503
SE .29 .26 —.71
CV 4.44 4.26 -5.22
n20 20 14
Ea leng h
Mean 28.45 28.55 28.00 27.50 Sex 1.032 .02 .888
SD 1.57 .95 —.58 Wi hin 43 1.599
SE .35 .21 —.29
CV 5.52 3.33 —2.1
1
n20 20 14
T agus leng h
Mean 8.25 8.10 8.00 7.50 Sex 1.635 1.98 .167
SD .55 .55 —.58 Wi hin 43 .321
SE .12 .12 —.29
CV 6.67 6.79 —7.73
n20 20 14
Fo ea m leng h
Mean 85.70 84.85 84.00 84.75 Sex 16.914 1.71 .199
SD 1.98 2.1
1
—2.06 Wi hin 43 4.053
SE .44 .47 —1.03
CV 2.31 2.49 —2.43
n20 20 14
Weigh
Mean 69.20 61.33 61.50 55.88 Sex 1793.411 11.35 .002
SD 11.45 4.07 —3.75 Wi hin 43 69.895
SE 2.56 .91 —1.88
CV 16.55 6.64 —6.71
n20 20 14
Leng h o digi one
Mean 13.45 12.35 13.00 12.75 Sex 111.468 15.92 <.001
SD 1.00 .75 —.50 Wi hin 43 .720
SE .22 .17 —.25
CV 7.43 6.07 —3.92
n20 20 14
1983 W1LLIG- BRAZILIAN BAT COMMUNITIES 23
Table 4. —Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 99 66 99 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 169.35 164.75 167.00 166.00 Sex 1205.143 10.13 .003
SD 4.57 4.67 —4.08 Wi hin 43 20.250
SE 1.02 1.04 —2.04
CV 2.70 2.83 —2.46
n20 20 14
Leng h o digi ou
Mean 127.1
1
123.05 124.00 124.25 Sex 1153.525 10.76 .002
SD 3.93 3.86 —2.75 Wi hin 43 14.266
SE .88 .86 —1.38
CV 3.09 3.14 —2.21
n20 20 14
Leng h o digi i e
Mean 100.70 98.05 100.00 96.75 Sex 189.91
1
11.11 .002
SD 3.13 2.69 —2.50 Wi hin 43 8.093
SE .70 .60 —1.25
CV 3.11 2.74 —2.58
n20 20 14
Tibia leng h
Mean 41.55 39.80 41.00 40.00 Sex 132.006 17.52 <.001
SD 1.36 1.40 —1.41 Wi hin 43 1.827
SE .30 .31 —.71
CV 3.27 3.52 —3.53
n20 20 14
Calca leng h
Mean 42.85 41.30 43.00 40.00 Sex 135.240 31.99 .001
SD 1.95 1.56 —.82 Wi hin 43 2.940
SE .44 .35 —.41
CV 4.55 3.78 —2.05
n20 20 14
G ea es leng h o skull
Mean 26.77 24.93 24.50 24.43 Sex I36.745 69.69 <.001
SD .81 .47 —.34 Wi hin 43 .527
SE .18 .11 —.17
CV 3.03 1.89 —1.39
n20 20 14
Condylobasal leng h
Mean 24.33 23.16 23.60 23.23 Sex 114.265 103.22 <.001
SD .43 .30 —.29 Wi hin 43 .138
SE .10 .07 —.14
CV 1.77 1.30 —1.25
n20 20 14
Zygoma ic b ead h
Mean 19.63 18.70 19.70 18.60 Sex 110.007 58.18 <.001
SD .54 .31 —.16 Wi hin 43 .172
SE .12 .07 —.08
CV 2.75 1.66 —.86
n20 20 14
30 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 6.— Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o/'Mic onyc e is megalo is males and emales om bo h Caa inga and Ce ado biomes. Due
o small sample sizes, da a om bo h biomes we e combined o de e mine he exis ence o signi ican (P <.050) seconda y sexual
a ia ion ia aone-way analysis o a iance (Model I) wi h eplica ion.
Caa inga Ce ado Analysis o a iance
<5<5 99 <5<5 99 Fac o d MS FSigni icance
To al leng h
Mean 59.86 60.00 58.67 Sex 1.714 .06 .805
SD 2.91 2.00 5.86 Wi hin 12 11.208
SE 1.10 1.00 3.38
CV 4.86 3.33 9.99
n743
Tail leng h
Mean 13.57 13.00 14.67 Sex 12.314 .96 .346
SD 1.40 1.41 2.08 Wi hin 12 2.408
SE .53 .71 1.20
CV 10.32 10.85 14.17
n743
Hind oo leng h
Mean 8.14 8.00 8.00 Sex 1.029 .12 .737
SD .38 01.00 Wi hin 12 .242
SE .14 0.58
CV 4.67 012.50
n743
Ea leng h
Mean 20.86 21.25 20.33 Sex 1.864 2.14 .169
SD .69 .50 .58 Wi hin 12 .404
SE .26 .25 .33
CV 3.31 2.35 2.85
n743
T agus leng h
Mean 7.57 6.75 8.00 Sex 12.579 10.86 .006
SD .53 .50 0Wi hin 12 .238
SE .20 .25 0
CV 7.13 7.41 0
n743
Fo ea m leng h
Mean 34.29 34.75 34.33 Sex 1.579 1.01 .334
SD .76 .50 1.16 Wi hin 12 .571
SE .29 .25 .67
CV 2.22 1.44 3.38
n743
Weigh
Mean 5.93 5.75 6.67 Sex 1.457 .75 .402
SD .54 .29 1.44 Wi hin 12 .606
SE .20 .14 .83
CV 9.1
1
5.04 21.59
n743
Leng h o digi one
Mean 8.43 8.25 9.00 Sex 1.350 1.33 .271
SD .54 .50 0Wi hin 12 .263
SE .20 .25 0
CV 6.41 6.06 0
n743
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 31
Table 6. —Con inued.
Caa inga Ce ado Analysis o a iance
<5<$ 22 66 22 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 61.00 62.50 63.67 Sex 11.400 .24 .635
SD 2.00 1.00 3.79 Wi hin 12 5.883
SE .76 .50 2.19
CV 3.28 1.60 5.95
n743
Leng h o digi ou
Mean 46.57 46.75 47.00 Sex 1.007 .00 .947
SD 1.40 1.26 1.00 Wi hin 12 1.571
SE .53 .63 .58
CV 3.01 2.70 2.13
n743
Leng h o digi i e
Mean 47.71 48.00 48.33 Sex 1.029 .01 .918
SD 1.80 .82 2.08 Wi hin 12 2.575
SE .68 .41 1.20
CV 3.77 1.71 4.30
n743
Tibia leng h
Mean 16.71 15.75 16.67 Sex 12.579 2.41 .147
SD 1.25 .50 1.16 Wi hin 12 1.071
SE .47 .25 .67
CV 7.48 3.17 6.96
n743
Calca leng h
Mean 11.00 10.75 10.67 Sex 1.064 .10 .756
SD 1.00 .50 .58 Wi hin 12 .638
SE .38 .25 .33
CV 9.09 4.65 5.44
n743
Noselea leng h
Mean 6.42 7.00 7.00 Sex 1.457 .86 .373
SD .79 01.00 Wi hin 12 .533
SE .30 0.58
CV 12.31 014.29
n743
G ea es leng h o skull
Mean 18.18 17.73 18.17 Sex 1.568 2.81 .122
SD .44 .31 .70 Wi hin 1
1
.202
SE .18 .16.41
CV 2.42 1.75 3.85
n643
Condylobasal leng h
Mean 16.17 15.58 16.20 Sex 11.006 4.49 .058
SD .57 .13 .62 Wi hin 1
1
.224
SE .23 .06 .36
CV 3.53 .83 3.83
n643
Zygoma ic b ead h
Mean 8.88 8.67 8.90 Sex 1.106 1.86 .206
SD .33 .12 .17 Wi hin 9.057
SE .15 .07 .10
CV 3.72 1.38 1.91
n533
32 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 6. —Con inued.
Caa inga Ce ado Analysis o a iance
66 29 66 22 Fac o d MS FSigni icance
Pos o bi al cons ic ion
Mean 3.85 3.78 3.77 Sex 1.006 .30 .592
SD .18 .05 .15 Wi hin 10 .020
SE .07 .03 .09
CV 4.68 1.32 3.98
n643
Mas oid b ead h
Mean 8.54 8.33 8.70 Sex 1.202 5.20 .046
SD .17 .13 .30 Wi hin 10 .039
SE .08 .06 .17
CV 1.99 1.56 3.45
n543
B ead h o b aincase
Mean 7.38 7.35 7.63 Sex 1.042 1.03 .334
SD .16 .13 .25 Wi hin 10 .041
SE .07 .07 .15
CV 2.17 1.77 3.28
n543
Ros al b ead h
Mean 3.26 3.23 3.27 Sex 1.004 .14 .711
SD .13 .17 .23 Wi hin 12 .024
SE .05 .09 .13
CV 3.99 5.26 7.03
n743
Heigh o b aincase
Mean 9.48 9.35 9.43 Sex 1.034 .44 .522
SD .34 .24 .31 Wi hin 1
1
.086
SE .14 .12 .18
CV 3.59 2.57 3.29
n643
B ead h ac oss he uppe mola s
Mean 6.02 5.85 5.93 Sex 1.053 3.28 .097
SD .08 .13 .21 Wi hin 11 .016
SE .03 .07 .12
CV 1.33 2.22 3.54
n643
B ead h ac oss he uppe canines
Mean 3.15 3.20 3.17 Sex 1.006 .49 .498
SD .11 .12 .12 Wi hin 1
1
.011
SE .04 .06 .07
CV 3.49 3.75 3.79
n643
Leng h o he maxilla y oo h ow
Mean 6.80 6.60 6.93 Sex 1.165 1.02 .333
SD .55 .08 .21 Wi hin 12 .162
SE .21 .04 .12
CV 8.09 1.21 3.03
n743
Leng h o he uppe mola i o m oo h ow
Mean 5.73 5.50 5.97 Sex 1.179 1.12 .310
SD .54 .10 .15 Wi hin 12 .159
SE .20 .05 .09
CV 9.42 1.82 2.51
n743
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 33
Table 6. —Con inued.
Caa inga Ce ado Analysis o anance
<55 99 55 99 Fac o d MS FSigni icance
Wid h o he wides mola
Mean 1.69 1.55 1.73 Sex 1.064 8.57 .013
SD .09 .10 .06 Wi hin 12 .008
SE .03 .05 .03
CV 5.33 6.45 3.47
n743
G ea es leng h o he mandible
Mean 11.79 11.40 11.73 Sex 1.391 2.93 .113
SD .44 .18 .42 Wi hin 12 .113
SE .17 .09 .24
CV 3.73 1.58 3.58
n743
Leng h o he mandibula oo h ow
Mean 7.50 7.18 7.50 Sex 1.302 5.77 .033
SD .29 .10 .20 Wi hin 12 .052
SE .11 .05 .12
CV 3.87 1.39 2.67
n743
Leng h o he co onoid p ocess
Mean 3.87 3.68 3.93 Sex 1.132 10.13 .008
SD .14 .05 .12 Wi hin 12 .013
SE .05 .03 .07
CV 3.62 1.36 3.05
n743
Table 7.—Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion. SD; s anda d e o o he mean. SE;
coe icien o a ia ion. CV; sample size, n) o/Mic onyc e is minu a males and emales om bo h Caa inga and Ce ado biomes. Due
o small sample sizes, da a om bo h biomes we e combined o de e mine he exis ence o signi ican (V <.050) seconda y sexual
a ia ion ia aone-way analysis o a iance (Model I) wi h eplica ion.
Caa inga Ce ado Analysis o a iance
55 99 55 99 Fac o d MS FSigni icance
To al leng h
Mean 54.75 56.00 63.00 Sex 110.971 .70 .427
SD .50 1.00 1.73 Wi hin 815.679
SE .25 .58 1.00
CV .91 1.79 2.74
n43 3
Tail leng h
Mean 11.25 12.00 12.33 Sex 1.171 .12 .738
SD .96 1.00 1.53 Wi hin 81.429
SE .48 .58 .88
CV 8.53 8.33 12.41
n43 3
Hind oo leng h
Mean 8.50 8.67 8.67 Sex 1.019 .06 .807
SD .58 .58 .58 Wi hin 8.298
SE .29 .33 .33
CV 6.82 6.70 6.69
n433
34 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 1.—Con inued.
Caa inga Ce ado Analysis o a iance
<5.5 92 <5<5 22 Fac o d MS FSigni icance
Ea leng h
Mean 18.50 19.33 26.00 Sex 1.476 .28 .610
SD 1.00 .58 4.58 Wi hin 81.691
SE .50 .33 2.65
CV 5.41 3.00 17.62
n433
T agus leng h
Mean 7.00 7.00 7.00
SD 0 0 0
SE 000
CV 000
n433
Fo ea m leng h
Mean 32.75 34.00 34.00 Sex 11.071 1.15 .314
SD .96 01.00 Wi hin 8.929
SE .48 0.58
CV 2.93 02.94
n433
Weigh
Mean 5.75 6.83 5.67 Sex 12.630 5.85 .042
SD .29 1.26 .29 Wi hin 8.449
SE .14 .73 .17
CV 5.04 18.45 5.1 1
n433
Leng h o digi one
Mean 6.25 7.00 6.67 Sex 1.686 3.20 111
SD .50 0.58 Wi hin 8.214
SE .25 0.33
CV 8.00 08.70
n 4 33
Leng h o digi h ee
Mean 54.50 57.33 56.67 Sex 17.619 2.99 .122
SD .58 2.31 .58 Wi hin 82.548
SE .29 1.33 .33
CV 1.06 4.03 1.02
n433
Leng h o digi ou
Mean 41.50 43.33 42.67 Sex 13.733 1.45 .264
SD 1.29 2.52 .58 Wi hin 82.583
SE .65 1.45 .33
CV 3.1 I5.82 1.36
n 4 3 3
Leng h o digi i e
Mean 41.50 44.00 42.67 Sex 18.400 5.60 .046
SD 1.00 1.73 .58 Wi hin 81.500
SE .50 1.00 .33
CV 3.77 3.93 1.36
n433
Tibia leng h
Mean 15.00 15.33 16.00 Sex 1.019 .02 .896
SD .82 1.16 1.00 Wi hin 81.048
SE .41 .67 .58
CV 5.47 7.57 6.25
n433
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 35
Table 1.—Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 29 66 29 Fac o d MS FSigni icance
Calca leng h
Mean 8.75 9.00 9.67 Sex 1.043 .04 .849
SD .96 1.00 1.16 Wi hin 81.107
SE .48 .58 .67
CV 10.97 11.11 12.00
n433
Noselea leng h
Mean 5.25 5.33 5.33 Sex 1.005 .02 .896
SD .50 .58 .58 Wi hin 8.262
SE .25 .33 .33
CV 9.52 10.88 10.88
n433
G ea es leng h o skull
Mean 16.80 16.77 16.95 Sex 1.014 .12 .735
SD .41 .32 .21 Wi hin 7.112
SE .20 .19 .15
CV 2.44 1.91 1.24
n432
Condylobasal leng h
Mean 14.83 14.93 15.00 Sex 1.005 .10 .763
SD .29 .15 .14 Wi hin 7.051
SE .14 .09 .10
CV 1.96 1.00 .93
n432
Zygoma ic b ead h
Mean 7.93 8.10 8.00 Sex 1.030 4.00 .116
SD .96 .14 —Wi hin 4.008
SE .03 .10 —
CV .76 1.73 —
n3 2 1
Pos o bi al cons ic ion
Mean 3.75 3.90 4.03 Sex 1.002 .05 .835
SD .17 .17 .06 Wi hin 8.037
SE .09 .10 .03
CV 4.53 4.36 1.49
n433
Mas oid b ead h
Mean 8.15 8.18 8.40 Sex 1.009 .19 .673
SD .24 .15 .14 Wi hin 7.046
SE .12 .09 .10
CV 2.94 1.84 1.67
n432
B ead h o b aincase
Mean 7.25 7.23 7.40 Sex 1.005 .18 .685
SD .13 .21 .14 Wi hin 7.028
SE .06 .12 .10
CV 1.79 2.90 1.89
n432
Ros al b ead h
Mean 2.98 2.90 3.03 Sex 1.021 4.20 .075
SD .05 .10 .06 Wi hin 8.005
SE .03 .06 .03
CV 1.68 3.45 1.98
n43 3
36 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 1.—Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 9$ 66 99 Fac o d MS FSigni icance
Heigh o b aincase
Mean 8.90 8.93 8.90 Sex 1.002 .04 .838
SD .29 .21 0Wi hin 7.050
SE .15 .21 0
c 3.26 2.35 0
n432
B ead h ac oss he uppe mola s
Mean 5.03 5.07 5.27 Sex 1.008 .46 .518
SD .05 .06 .12 Wi hin 8.018
SE .03 .03 .07
CV .99 1.18 2.28
n433
B ead h ac oss he uppe canines
Mean 2.73 2.67 2.87 Sex 1.030 2.50 .153
SD .05 .06 .15 Wi hin 8.012
SE .03 .03 .09
CV 1.83 2.25 5.23
n433
Leng h o he maxilla y oo h ow
Mean 5.75 5.97 5.73 Sex 1.015 5.14 .053
SD .06 .25 .12 Wi hin 8.021
SE .03 .15 .07
CV 1.04 4.19 2.09
n43 3
Leng h o he uppe mola o m oo h ow
Mean 4.88 4.90 4.90 Sex 1.000 .03 .863
SD .10 .17 .10 Wi hin 8.014
SE .05 .10 .06
CV 2.05 3.47 2.04
n433
Wid h o he wides mola
Mean 1.38 1.43 1.33 Sex 1.012 4.10 .078
SD .05 .06 .06 Wi hin 8.003
SE .03 .03 .03
CV 3.62 4.20 4.51
n43 3
G ea es leng h o he mandible
Mean 10.43 10.43 10.47 Sex !.000 .02 .896
SD .10 .06 .15 Wi hin 8.011
SE .05 .03 .09
CV .96 .58 1.43
n43 3
Leng h o he mandibula oo h ow
Mean 6.28 6.27 6.05 Sex 1.009 .27 .617
SD .17 .12 .21 Wi hin 7.032
SE .09 .07 .15
CV 2.71 1.91 3.47
n432
Leng h o he co onoid p ocess
Mean 3.05 3.10 3.27 Sex 1.004 .26 .622
SD .06 .10 .06 Wi hin 8.015
SE .03 .06 .03
CV 1.97 3.23 1.83
n433
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 37
an analyzing males and emales sepa a ely in u u e
analyses.
Tona ia bidens (Spix, 1823)
T. b. bidens is he only ex an subspecies cu en ly
ecognized (Jones and Ca e , 1976). This oliage
gleaning insec i o e was a e in he Caa ingas whe e
i was only cap u ed on se o es; i was appa en ly
absen om he Chapada do A a ipe.
Few mensu al da a a e epo ed in he li e a u e
o T. bidens. Dobson (1879), Lima (1926), Sanbo n
(1936), Cunha Viei a (1942), and Goodwin (1942)
epo ed measu emen s om B azilian specimens.
Only selec ed measu emen s (a e Swanepoel and
Genoways, 1979) a e epo ed he e o he wo adul
males and one adul emale cap u ed in he Caa in-
gas ( o al leng h, 85, 96, 93; ail leng h, 18, 19, 18;
hind oo leng h, 14, 14, 13; ea leng h, 32, 30, 33;
g ea es leng h o skull, 27.4, 27.3, 27.1; condylo-
basal leng h, 24.1, 23.1, 23.6; zygoma ic b ead h,
14.3, 14.0, 14.0; pos o bi al cons ic ion, 5.7, 5.3,
5.2; b ead h o b aincase, 11.0, 10.7, 10.7; leng h
o he maxilla y oo h ow, 9.8, 9.8, 9.5; b ead h
ac oss he uppe mola s, 8.6, 8.8, 8.5).
Tona ia b asiliense (Pe e s, 1866)
Jones and Ca e (1976) conside T. b asiliense
o be mono ypic; howe e , i one includes speci-
mens o T. enezuelae, T. minu a, and T. nica a-
guae wi hin his axon as is equen ly done (Ga d-
ne , 1976; Koopman, 1978), hen he species is, no
doub , poly ypic. I was uncommon in he Caa ingas
bu was consis en ly cap u ed on se o es o in Caa-
inga Al a habi a s. T. b asiliense was absen om
all habi a s on Chapada do A a ipe.
Mos o he eco ded measu emen s o T. b as-
iliense we e de i ed om Pe e s (1866) o iginal de-
sc ip ion o he holo ype. Goodwin (1942) p esen ed
da a on wo mo e specimens om B azil and Ga d-
ne (1976) p esen ed da a on wo addi ional speci-
mens om Pe u. Samples om he Caa ingas a e
oo small o allow s a is ical analyses, he e o e da a
a e epo ed he e on aselec ed se o mensu al cha -
ac e s (a e Swanepoel and Genoways, 1979). The
i s igu e ep esen s he mean alue de i ed om
six adul emales, whe eas he second igu e ep e-
sen s he mean alue de i ed om wo adul males
( o al leng h, 69.8, 75.0; ail leng h, 9.2, 11.0; hind-
oo leng h, 9.8, 10.0; ea leng h, 23.8, 22.0; g ea es
leng h o skull, 20.44, 19.95; condylobasal leng h,
17.86, 17.40; zygoma ic b ead h, 10.30, 9.95; pos -
o bi al cons ic ion, 3.23, 3.25; b ead h o b ain-
case, 8.42, 8.25; leng h o maxilla y oo h ow, 8. 18,
8.10; b ead h ac oss he uppe mola s, 6.62, 6.45).
Tona ia sil icola (D’O bigny, 1836)
Two subspecies a e cu en ly ecognized in his
axon; T. s. laepho is is he designa ion app op ia e
o specimens om no heas e n B azil (Jones and
Ca e , 1976). This oliage gleaning insec i o e is
absen om he Chapada do A a ipe, and i s oc-
cu ence in he Caa ingas was p ima ily es ic ed
o se o es. All bu h ee o he specimens om he
No heas o B azil lis ed by Ma es e al. (1981) as
T. bidens we e ac ually T. sil icola. Based upon ne -
ing eco ds o adul specimens om he du a ion
o his s udy, he sex a io was no equal in his
species (Bionomial es , P<.05); app oxima ely
30% o he 47 cap u ed adul indi iduals we e males.
Swanepoel and Genoways (1979) summa ized he
sou ces o eco ded measu emen s o T. sil icola;
Pe e s (1865), Dobson (1878), Cunha Viei a (1942),
and Goodwin (1942) a e no ewo hy in ha lis ing
because hey ha e included B azilian specimens in
hei wo k. S a is ical analyses o indi idual a ia-
ion ha e no been done no has he exis ence o
sexual dimo phism been conside ed in he li e a-
u e. Samples a e su icien ly la ge om he Caa in-
gas o de e mine indi idual a ia ion and asce ain
wi h con idence he deg ee o sexual dimo phism
p esen in apopula ion o T. sil icola. The esul s
o he s a is ical analysis a e p esen ed in Table 8.
The sample means o males a e la ge han he
sample means o emales o 24 o he 30 cha ac e s
examined. S a is ically signi ican seconda y sexual
a ia ion is e ealed o wo ex e nal cha ac e s (ea
leng h and weigh ) and six c anial cha ac e s (g ea -
es leng h o skull, condylobasal leng h, zygoma ic
b ead h, mas oid b ead h, heigh o he b aincase,
and leng h in he mandible om he canine o he
las mola ). S a is ical signi icance is no ound in
he a iables ha had la ge sample mean alues
o emales. T. sil icola is clea ly dimo phic o c a-
nial cha ac e s wi h males la ge han emales.
Min on c enula um (E. Geo oy, 1819)
Jones and Ca e (1976) ecognize ou subspecies
in his axon, whe eas Koopman (1978) includes a
i h o m. In ei he case, M. c. c enula um is he
designa ion p ope ly applied o specimens om
no heas e n B azil. This oliage gleaning insec i-
o e was ound exclusi ely in Caa inga Al a habi-
a s.
Apauci y o da a exis s conce ning he mo pho-
me ies o M. c enula um (see Swanepoel and Gen-
38 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 8. —Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample sice, n) o Tona ia sil icola males and emales om he Caa inga biome. Aone-way analysis o
a iance (Model I) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual a ia ion is indica ed
by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
36 29 <5<$ 22 Fac o d MS FSigni icance
To al leng h
Mean 97.85 97.58 Sex 1.660 .07 .799
SD 3.34 3.12 Wi hin 47 10.095
SE .93 .52
CV 3.41 3.20
n13 36
Tail leng h
Mean 17.69 17.69 Sex 1.000 .00 .998
SD 2.50 2.04 Wi hin 47 4.690
SE .69 .34
CV 14.13 11.53
n13 36
Hmd oo leng h
Mean 14.62 14.69 Sex 1.060 .11 .747
SD .51 .82 Wi hin 47 .568
SE .14 .14
CV 3.49 5.58
n13 36
Ea leng h
Mean 29.62 28.81 Sex 16.264 7.60 .008
SD 1.12 .82 Wi hin 47 .824
SE .31 .14
CV 3.78 2.85
n13 36
T agus leng h
Mean 11.69 11.86 Sex 1.272 .28 .597
SD 1.38 .80 Wi hin 47 .959
SE .38 .13
CV 11.80 6.74
n13 36
Fo ea m leng h
Mean 58.23 57.58 Sex 14.004 3.30 .076
SD 1.42 .97 Wi hin 47 1.214
SE .40 .16
CV 2.44 1.68
n13 36
Weigh
Mean 33.85 31.79 Sex 140.314 5.06 .029
SD 3.07 2.73 Wi hin 47 7.966
SE .85 .46
CV 9.07 8.59
n13 36
Leng h o digi one
Mean 15.08 14.94 Sex 1.168 .23 .637
SD .76 .89 Wi hin 47 .74!
SE .21 .15
CV 5.04 5.96
n13 36
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 39
Table 8. —Con inued.
Caa inga Ce ado Analysis o a iance
<33 99 <5.3 99 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 106.54 106.00 Sex 12.769 .65 .423
SD 2.63 1.82 Wi hin 47 4.239
SE .73 .30
CV 2.47 1.72
n13 36
Leng h o digi ou
Mean 80.46 79.8! Sex 14.1 10 1.26 .267
SD 2.37 1.56 Wi hin 47 3.253
SE .66 .26
CV 2.95 1.95
n13 36
Leng h o digi i e
Mean 81.39 80.47 Sex 17.951 3.06 .087
SD 2.40 1.23 Wi hin 47 2.597
SE .67 .21
CV 2.95 1.53
n13 36
Tibia leng h
Mean 27.08 27.25 Sex 1.286 .25 .6 1
9
SD 1.66 .77 Wi hin 47 1.142
SE .46 .13
CV 6.13 2.83
n13 36
Calca leng h
Mean 18.54 17.78 Sex 15.527 3.35 .073
SD 1.33 1.27 Wi hin 47 1.648
SE .37 .21
CV 7.17 7.14
n13 36
Noselea leng h
Mean 6.54 6.33 Sex 1.402 1.68 .201
SD .52 .48 Wi hin 47 .239
SE .14 .08
CV 7.95 7.58
n13 36
G ea es leng h o skull
Mean 27.80 27.32 Sex 12.180 7.93 .007
SD .54 .52 Wi hin 47 .275
SE .15 .09
CV 1.94 1.90
n13 36
Condylobasal leng h
Mean 24.31 23.84 Sex 12.050 114.56 <.001
SD .50 .32 Wi hin 47 .141
SE .14 .05
CV 2.06 1.34
n13 36
Zygoma ic b ead h
Mean 13.68 13.47 Sex 1.412 6.82 .012
SD .24 .25 Wi hin 45 .060
SE .07 .04
CV 1.75 1.86
n13 34
46 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 10.— Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o/’Phyllos omus discolo males and emales om bo h Caa inga and Ce ado biomes. A
wo-way analysis o a iance (Model I) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual o
geog aphic a ia ion is indica ed by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
<36 9$ 66 $9 Fac o d MS FSigni icance
To al leng h
Mean 96.30 98.31 98.50 98.70 A ea 125.505 2.28 .136
SD 4.08 3.14 3.58 2.83 Sex 118.648 1.67 .201
SE 1.29 .78 .80 .63 AXS112.515 1.12 .294
CV 4.24 3.19 3.63 2.87 E o 62 11.173
n10 16 20 20
Tail leng h
Mean 14.90 15.13 14.65 14.05 A ea 17.736 3.91 .052
SD 1.66 1.31 1.47 1.28 Sex 1.288 .15 .704
SE .53 .33 .33 .29 AXS12.002 1.01 .318
CV 11.14 18.66 10.10 9.11 E o 62 1.977
n10 16 20 20
Hind oo leng h
Mean 13.20 13.19 13.70 13.45 A ea 12.215 4.14 .046
SD .79 .66 .73 .76 Sex 1.263 .49 .486
SE .25 .16 .16 .17 AXS1.215 .40 .529
CV 5.98 5.00 5.33 5.65 E o 62 .535
n10 16 20 20
Ea leng h
Mean 21.60 21.13 20.80 21.80 A ea 1.060 .03 .863
SD 1.35 1.15 1.15 1.80 Sex 11.050 .53 .469
SE .43 .29 .26 .40 AXSI8.288 4.19 .045
CV 6.25 5.44 5.53 8.26 E o 62 1.977
n10 16 20 20
T agus leng h
Mean 8.90 8.81 9.05 9.00 A ea 1.434 .78 .379
SD .74 .91 .61 .73 Sex 1.072 .13 .719
SE .23 .23 .14 .16 AXS1.005 .01 .922
CV 8.31 10.33 6.74 8.1
1
E o 62 .553
n10 16 20 20
Fo ea m leng h
Mean 61.60 59.81 60.20 59.70 A ea 18.715 3.14 .081
SD 1.84 1.52 1.80 1.56 Sex 119.934 7.18 .009
SE .59 .38 .40 .35 AXS16.315 2.27 .137
CV 2.99 2.54 2.99 2.61 E o 62 2.778
n10 16 20 20
Weigh
Mean 37.50 40.06 38.85 36.68 A ea 115.815 1.49 .227
SD 3.27 3.29 2.16 4.05 Sex 1.572 .05 .817
SE 1.03 .82 .48 .91 AXS185.501 8.05 .006
CV 8.72 8.21 5.56 11.04 E o 62 10.615
n10 16 20 20
Leng h o digi one
Mean 11.30 11.13 11.75 11.35 A ea 11.736 4.46 .039
SD .48 .50 .64 .75 Sex 11.260 3.23 .077
SE .15 .13 .14 .17 AXS1.193 .50 .484
CV 4.25 4.49 5.45 6.61 E o 62 .390
n10 16 20 20
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 47
Table 10. —Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 99 66 99 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 109.10 105.00 105.55 105.50 A ea 135.438 3.33 .073
SD 3.60 2.68 2.46 4.10 Sex 165.610 6.17 .016
SE 1.14 .67 .55 .92 AXS162.486 5.88 .018
CV 3.30 2.55 2.33 3.89 E o 62 10.627
n10 16 20 20
Leng h o digi ou
Mean 80.70 78.31 78.65 78.15 A ea 118.648 2.50 .119
SD 2.83 1.89 2.06 3.68 Sex 131.763 4.26 .043
SE .90 .47 .46 .82 AXS113.572 1.82 .182
CV 3.51 2.41 2.62 4.71 E o 62 7.462
n10 16 20 20
Leng h o digi i e
Mean 73.40 71.63 71.85 71.35 A ea 112.688 2.28 .136
SD 2.01 2.06 1.98 3.00 Sex 119.717 3.54 .065
SE .64 .52 .44 .67 AXS16.193 1.11 .296
CV 2.74 2.88 2.76 4.20 E o 62 5.569
n10 16 20 20
Tibia leng h
Mean 24.70 24.88 25.45 24.45 A ea 1.402 .26 .615
SD 1.83 .89 1.32 1.10 Sex 12.593 1.64 .205
SE .58 .22 .29 .25 AXS15.260 3.34 .073
CV 7.41 3.58 5.19 4.50 E o 62 1.577
n10 16 20 20
Calca leng h
Mean 9.80 9.75 9.55 10.05 A ea 1.010 .01 .917
SD .92 1.13 .95 .76 Sex 1.771 .88 .353
SE .29 .28 .21 .17 AXS11.152 1.31 .257
CV 9.38 11.59 9.95 7.56 E o 62 .879
n10 16 20 20
Noselea leng h
Mean 6.70 6.19 6.50 6.40 A ea 1.001 .00 .972
SD .48 .91 .51 .75 Sex 11.429 2.92 .092
SE .15 .23 .12 .17 AXS1.648 1.32 .254
CV 7.16 14.70 7.85 11.72 E o 62 .489
n10 16 20 20
G ea es leng h o skull
Mean 29.22 29.18 29.44 28.65 A ea 1.345 1.06 .307
SD .52 .61 .65 .47 Sex 12.562 7.87 .007
SE .17 .15 .15 .1
1
AXS12.016 6.19 .016
CV 1.78 2.09 2.21 1.64 E o 61 .326
n9 16 20 20
Condylobasal leng h
Mean 26.34 26.22 26.6! 25.7! A ea 1.234 .80 .375
SD .62 .51 .48 .58 Sex 13.845 13.10 .001
SE .21 .13 .11 .13 AXS12.191 7.47 .008
CV 2.35 1.95 1.80 2.26 E o 61 .294
n916 20 20
Zygoma ic b ead h
Mean 15.23 15.56 15.55 14.94 A ea 1.342 2.07 .156
SD .46 .37 .39 .43 Sex 1.288 1.74 .192
SE .15 .09 .09 .10 AXS13.224 19.48 <.001
CV 3.02 2.38 2.51 2.88 E o 6
1
.166
n 9 16 20 20
48 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 10 .—Con inued.
Caa inga Ce ado Analysis o a iance
<56 92 6<5 99 Fac o d MS FSigni icance
Pos o bi al cons ic ion
Mean 6.47 6.38 6.38 6.37 A ea 1.035 .63 .432
SD .15 .16 .26 .28 Sex 1.033 .59 .444
SE .05 .04 .06 .06 AXS1.021 .37 .545
CV 2.32 2.51 4.08 4.40 E o 61 .056
n916 20 20
Mas oid b ead h
Mean 14.42 14.78 14.91 14.19 A ea 1.042 .38 .538
SD .24 .38 .37 .32 Sex 1.493 4.50 .038
SE .08 .08 .08 .07 AXS14.207 38.41 <.001
CV 1.66 2.23 2.48 2.26 E o 61 .110
n916 20 20
B ead h o b aincase
Mean 12.13 12.03 12.04 11.82 A ea 1.339 5.10 .028
SD .20 .18 .30 .29 Sex 1.379 5.70 .020
SE .07 .04 .07 .07 AXS1.051 .76 .385
CV 1.65 1.50 2.49 2.45 E o 61 .067
n9 16 20 20
Ros al b ead h
Mean 6.74 6.96 7.07 6.67 A ea 1.003 .06 .815
SD .29 .19 .23 .25 Sex 1.129 2.27 .137
SE .10 .05 .05 .06 AXS1.368 24.03 <.001
CV 4.30 2.73 3.25 3.75 E o 61 .057
n916 20 20
Heigh o b aincase
Mean 13.18 13.33 13.33 12.99 A ea 1.122 .94 .336
SD .43 .27 .30 .44 Sex 1.136 1.05 .311
SE .14 .07 .07 .10 AXS1.868 6.68 .012
CV 3.26 2.03 2.25 3.39 E o 61 .130
n916 20 20
B ead h ac oss he uppe mola s
Mean 9.86 9.76 9.98 9.73 A ea 1.028 .32 .573
SD .24 .32 .25 .34 Sex 1.446 5.04 .028
SE .08 .08 .06 .08 AXS1.083 .94 .337
CV 2.43 3.29 2.51 3.49 E o 61 .089
n916 20 20
B ead h ac oss he uppe canines
Mean 6.88 7.21 7.27 6.75 A ea 1.018 .42 .518
SD .15 .22 .21 .22 Sex 1.126 2.94 .092
SE .05 .05 .05 .05 AXS12.670 62.52 <.001
CV 2.18 3.05 2.89 3.26 E o 61 .043
n916 20 20
Leng h o he maxilla y oo h ow
Mean 9.53 9.53 9.69 9.37 A ea 1.000 .01 .922
SD .19 .18 .15 .22 Sex 1.391 11.16 .001
SE .06 .04 .03 .05 AXS1.381 10.87 002
CV 1.99 1.89 1.55 2.35 E o 61 .035
n916 20 20
Leng h o he uppe nola i o m oo h ow
Mean 7.59 7.43 7.60 7.47 A ea 1.009 .24 .624
SD .16 .23 .17 .20 Sex 1.302 8.07 .006
SE .05 .06 .04 .04 AXS1.003 .07 .786
CV 2.1 13.10 2.24 2.68 E o 61 .038
n9 16 20 20
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 49
Tabic 10. —Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 92 66 29 Fac o d MS FSigni icance
Wid h o he wides mola
Mean 2.12 2.06 2.13 2.08 A ea 1.00! .08 .781
SD .11 .14 .09 .08 Sex 1.044 4.02 .050
SE .04 .04 .02 .02 AXS1.000 .03 .860
CV 5.19 6.80 4.23 3.85 E o 6
1
.011
n916 20 20
G ea es leng h o he mandible
Mean 19.31 19.30 19.43 18.76 A ea 1.648 3.82 .055
SD .38 .39 .40 .45 Sex 11.696 9.98 .003
SE .13 .10 .09 .10 AXS11.587 9.34 .003
CV 1.97 2.02 2.06 2.40 E o 6
1
.170
n916 20 20
Leng h o he mandibula oo h ow
Mean 10.34 10.59 10.75 10.39 A ea 1.142 1.98 .165
SD .27 .29 .31 .19 Sex 1.049 .68 .412
SE .09 .07 .07 .04 AXS11.379 19.27 <.001
CV 2.61 2.74 2.88 1.83 E o 6
1
.072
n916 20 20
Leng h o he co onoid p ocess
Mean 6.38 6.74 7.03 6.79 A ea 11.789 16.44 <.001
SD .29 .29 .34 .37 Sex 1.048 .44 .509
SE .10 .07 .08 .08 AXS11.337 12.28 <.001
CV 4.55 4.30 4.84 5.45 E o 6
1
.109
n916 20 20
o he uppe mola i o m oo h ow, wid h o he
wides mola , and g ea es leng h o he maxilla)
exhibi s a is ically signi ican seconda y sexual
a ia ion. In he only case whe e emales ha e a
la ge sample mean alues han males ( o al leng h),
s a is ical signi icance is no obse ed.
Clea ly, when samples a e om a es ic ed lo-
cali y, as in he wo k o Taddei (1975#) o when
geog aphic di e ences be ween sampling locali ies
can be s a is ically con olled, as in he analyses o
he Caa ingas and Ce ado, s ong sexual dimo -
phism has been de ec ed by uni a ia e analyses wi h
males consis en ly la ge han emales. Pe haps he
inclusion o specimens om dispa a e locali ies has
con ounded he esul s in he wo k o Valdez (1970).
Simila ly, he la ge mo phome ic he e ogenei y a -
ibu able o geog aphic di e ences in he samples
used by Powe and Tamsi (1973) may ha e p e-
en ed asimple se o unipola size ec o s om
accoun ing o he di e ences be ween he sexes
shown by disc iminan unc ion analysis.
Phyllos omus has a us (Pallas, 1767)
P. h. has a us is he p esen ly ecognized subspe-
cies designa ion o popula ions o his species in
he No heas o B azil (Jones and Ca e , 1976).
This la ge omni o e was a e in he Caa ingas bu
widesp ead and common in bo h Ce ado and Ce -
adao habi a s on he Chapada do A a ipe. Acolony
o mo e han 100 indi iduals oos ed in asmall
a ic-like space (6 mby .5 mby .5 m) in he en ance
a ch o he Colegio Ag icola de C a o; he oos was
also occupied by ala ge colony o Molossus mo-
lossus (>300 indi iduals). O he 108 indi iduals
o P. h. has a us e en ually cap u ed om he oos ,
48 we e males. This s ongly sugges s an equal sex
a io (Binomial Tes , P>.05) and he absence o
ha em g oups wi hin he colony (con a y o he
si ua ion epo ed by B adbu y, 1977).
Swanepoel and Genoways (1979) ci e au ho s who
ha e published mo phome ic da a on P. has a us.
G. M. Allen (1908), Lima (1926), Cunha Viei a
(1942), and Taddei (1975#) a e he only wo ke s
who ha e included B azilian specimens in hei ac-
coun s. The wo k o Taddei (1975#) is unique be-
cause he pe o med s a is ical analyses on la ge sam-
ples. He de ec ed s a is ically signi ican seconda y
sexual a ia ion in all 17 o he c anial cha ac e s
examined and o eigh o he 17ex e nal cha ac e s.
Males a e la ge on he a e age han emales in all
50 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
cases. The esul s om he No heas o B azil a e
compa able (Table 11). Males ha e la ge sample
means han emales o all bu wo o he 30 a i-
ables. S a is ically signi ican seconda y sexual a i-
a ion is de ec ed in 15o he 16c anial cha ac e s
and in eigh o he 14 ex e nal cha ac e s. Clea ly,
P. has a us is dimo phic wi h males la ge han e-
males.
T achops ci hosus (Spix, 1823)
O he h ee nominal subspecies cu en ly in use,
T. c. ci hosus is he appella ion app op ia e o
No heas B azilian popula ions o his omni o e
because i s ype locali y is in Pe nambuco (Jones
and Ca e , 1976). The dis ibu ion o T. c. ci hosus
in he Caa ingas was es ic ed o se o es o a eas
con aining ock ou c oppings and i was absen om
bo h Ce ado and Ce adao habi a s on he Chapada
do A a ipe. Six een o 35 indi iduals cap u ed om
he Caa ingas we e males, hence he sex a io ap-
pea s o be equal (Binomial Tes , P>.05).
Pe e s (1865), Lima (1926), Cunha Viei a (1942),
and Fel en (1956) a e he only au ho s among he
g oup ci ed by Swanepoel and Genoways (1979) o
include measu emen s o T. ci hosus om B azil.
In o ma ion conce ning indi idual a ia ion is con-
spicuously absen om he li e a u e and seconda y
sexual a ia ion has no been examined in his
species. The esul s o as a is ical analysis o indi-
idual and seconda y sexual a ia ion o T. c. ci -
hosus a e gi en in Table 12. Only asingle cha ac e
exhibi s s a is ically signi ican seconda y sexual
a ia ion (b ead h ac oss he uppe canines). Males
ha e la ge sample means han emales o all o he
c anial cha ac e s and ou o he ex e nal cha ac e s
while emales ha e la ge sample means han males
o 10 ex e nal cha ac e s. Because o he ela i ely
la ge sample sizes, low le els o signi icance asso-
cia ed wi h he analyses o a iance and mino di -
e ences encoun e ed be ween male and emale mean
alues, i seems p uden o sugges ha T. ci hosus
does no exhibi sexual dimo phism.
Sub amily Glossophaginae
Glossophaga so icina (Pallas, 1766)
This nec a i o e was abundan and ubiqui ous in
Ce ado, Ce adao, and mos habi a s o he Caa-
ingas; i is pa icula ly dominan in dis u bed o
second g ow h a eas. Abandoned man-made s uc-
u es and ca es we e p e e ed oos ing si es and
colony size may exceed 2,000 indi iduals. Agg e-
ga ions o Ca ollia pe spicilla a con aining up o 20
indi iduals we e equen ly ound oos ing in as-
socia ion wi h G. so icina h oughou he No heas .
Small g oups o ba s we e equen ly obse ed o -
aging nea ui ing plan s. Solanum psnicula um was
he p edominan ui in he die o Caa ingas spec-
imens whe eas he ui o Vismia was p e e ed in
he Ce ado (al hough Solanum was also con-
sumed). Males and emales occu ed in s a is ically
indis inguishable p opo ions in he Caa ingas (Bi-
nomial Tes , P>.05); 46% o he 421 cap u ed adul
specimens we e males. On he Chapada do A a ipe,
56% o he 494 cap u ed adul specimens we e males.
Al hough he p opo ion o males and emales did
di e signi ican ly (Binomial Tes , P<.01), he
magni ude o he di e ence was small and p obably
e lec s he aga ies o sampling by mis ne a he
han cha ac e is ics o he social s uc u e in his
species.
In he Caa ingas, some sexual seg ega ion oc-
cu ed wi hin he la ge oos s ha occu ed in aban-
doned buildings. Females ended o cong ega e as a
cen al uni ; males occupied mo e dispe sed soli a y
loca ions a he pe iphe y o he oos . This phe-
nomenon became pa icula ly e iden as he ime
o pa u i ion app oached.
O he ou gene ally ecognized subspecies, Glos-
sophaga s. so icina is he designa ion applied o B a-
zilian specimens (Jones and Ca e , 1976). Many
au ho s epo mo phome ic da a on G. so icina
(see Swanepoel and Genoways, 1979), bu only Mil-
le (1913), Lima (1926), Cunha Viei a (1942), and
Taddei (19756) include specimens om B azil. Tad-
dei (19756) is unique among hese au ho s in wo k-
ing wi h la ge samples o G. so icina and pe o ming
s a is ical analyses on nume ous mo phome ic
cha ac e s. He inds s a is ically signi ican sexual
dimo phism in ou o 17ex e nal cha ac e s (head-
body leng h, o ea m leng h, leng h o he i h me a-
ca pal and leng h o he ou h me aca pal); in all
ou cases males we e la ge , on he a e age, han
emales. Among he 15c anial cha ac e s analyzed,
s a is ically signi ican seconda y sexual a ia ion is
appa en in se en cha ac e s (leng h o mola , man-
dibula oo h ow leng h, b ead h ac oss he canines,
zygoma ic b ead h, b aincase b ead h, mas oid
b ead h, and dep h o he c anium); emales a e la g-
e on he a e age o he i s wo cha ac e s while
males a e la ge on he a e age o he la e i e
cha ac e s.
The esul s o s a is ical analyses on he specimens
o G. s. so icina om he No heas o B azil a e
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 51
Table 11.—Summa y s a is ics o ex e na! and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE
;
coe icien o a ia ion, CV; sample size, n) o/'Phyllos omus has a us males and emales om bo h he Caa inga and Ce ado biomes.
Due o small sample sizes, da a om bo h biomes we e combined o de e mine he exis ence o signi ican (P <0.50) seconda y sexual
a ia ion ia aone-way analysis o a iance (Model I) wi h eplica ion.
Caa inga Ce ado Analysis o a iance
<5<5 92 <5<5 99 Fac o d MS FSigni icance
To al leng h
Mean 123.00 119.43 127.25 119.75 Sex 1643.527 34.46 <.001
SD —1.81 4.19 5.07 Wi hin 46 18.673
SE —.69 .94 1.13
CV —1.52 3.29 4.23
n1720 20
Tail leng h
Mean 19.00 17.86 19.15 18.50 Sex 17.741 1.60 .212
SD —2.04 1.93 2.57 Wi hin 46 4.839
SE —.77 .43 .57
CV —11.42 10.08 13.89
n1720 20
Hind oo leng h
Mean 17.00 17.57 17.75 17.05 Sex 13.307 2.80 .101
SD —.98 1.07 1.15 Wi hin 46 1.182
SE —.37 .24 .26
CV —5.58 6.03 6.74
n1720 20
Ea leng h
Mean 28.00 28.14 28.90 27.75 Sex 1 1 1.938 7.23 .010
SD —1.57 .97 1.48 Wi hin 46 1.652
SE —.60 .22 .33
CV —5.58 3.36 5.33
n1720 20
T agus leng h
Mean 13.00 13.08 13.20 13.20 Sex 1.021 .06 .810
SD —.58 .62 .62 Wi hin 46 .062
SE —.22 .14 .14
CV —4.46 4.70 4.70
n1720 20
Fo ea m leng h
Mean 81.00 81.29 84.30 81.55 Sex 183.667 20.33 <.001
SD —2.50 1.63 2.19 Wi hin 46 4.1 16
SE —.94 .36 .49
CV —3.08 1.93 2.69
n1720 20
Weigh
Mean 94.50 84.64 93.43 81.68 Sex 11,437.574 21.46 <.001
SD —8.94 8.58 7.80 Wi hin 46 66.987
SE —3.38 1.92 1.74
CV —10.56 9.18 9.55
nI720 20
Leng h o digi one
Mean 13.00 14.14 14.05 14.25 Sex 1.583 .55 .462
SD —1.07 1.28 .72 Wi hin 46 1.058
SE —.40 .29 .16
CV —7.57 9.1
1
5.05
n1720 20
52 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 11 .—Con inued.
Caa mga Ce ado Analysis o a iance
(36 99 66 99 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 145.00 145.86 147.35 143.95 Sex 192.191 5.64 .022
SD —3.80 3.42 4.86 Wi hin 46 16.358
SE —1.16 .77 1.09
CV —2.1
1
2.32 3.38
n1720 20
Leng h o digi ou
Mean 108.00 108.00 110.70 107.85 Sex 185.003 5.60 .022
SD —1.83 3.91 4.48 Wi hin 46 15.170
SE —.69 .87 1.00
CV —1.69 3.53 4.15
n1720 20
Leng h o digi i e
Mean 98.00 99.14 102.15 98.15 Sex 1148.446 14.42 <.001
SD —1.68 3.20 3.65 Wi hin 46 10.293
SE —.63 .69 .82
CV —1.69 3.03 3.72
n1720 20
Tibia leng h
Mean 32.00 32.57 33.65 32.75 Sex 18.894 10.03 .003
SD —.54 .67 1.21 Wi hin 46 .886
SE —.20 .15 .27
CV —1.66 1.99 3.69
n1720 20
Calca leng h
Mean 19.00 21.29 21.35 20.40 Sex 14.373 1.93 .171
SD —1.11 1.66 1.35 Wi hin 46 2.263
SE —.42 .37 .30
CV —5.21 7.78 6.62
n1720 20
Noselea leng h
Mean 9.00 9.14 8.95 9.30 Sex 11.112 3.62 .063
SD —.38 .39 .73 Wi hin 46 .307
SE —.14 .09 .16
CV —4.16 4.36 7.85
n1720 20
G ea es leng h o skull
Mean 36.20 35.04 36.93 35.29 Sex 132.917 63.35 <.001
SD —.26 .96 .51 Wi hin 46 .520
SE —.10 .21 .12
CV —.74 2.60 1.45
n1720 20
Cond lobasal leng h
Mean 32.20 31.16 32.70 31.16 Sex 127.01
1
92.12 <.001
SD —.38 .65 .48 Wi hin 46 .293
SE —.14 .15 .11
CV —1.22 1.99 1.54
n1720 20
Zygoma ic b ead h
Mean 20.90 19.54 20.93 19.70 Sex 119.000 68.79 <.001
SD —.47 .67 .39 Wi hin 46 .276
SE —.18 .15 .09
CV —2.41 3.20 1.98
n1720 20
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 53
Table 11. —Con inued.
Caa inga Ce ado Analysis o a iance
92 55 99 Fac o d MS FSigni icance
Pos o bi al cons ic ion
Mean 6.80 6.66 7.03 6.83 Sex 1.620 15.01 <.001
SD —.19 .23 .16 Wi hin 46 .041
SE —.07 .05 .04
CV —2.85 3.27 2.34
n1720 20
Mas oid b ead h
Mean 19.30 18.14 19.46 18.00 Sex 123.627 145.84 <.001
SD -.28 .51 .32 Wi hin 46 .162
SE —.10 .11 .07
CV —1.54 2.62 1.78
n1720 20
B ead h o b aincase
Mean 14.00 13.59 14.04 13.63 Sex 12.080 28.22 <.001
SD -.11 .29 .30 Wi hin 46 .074
SE —.04 .07 .07
CV —.81 2.07 2.20
n1720 20
Ros al b ead h
Mean 9.50 9.09 9.65 9.14 Sex 13.099 56.09 <.001
SD —.23 .25 .24 Wi hin 46 .055
SE -.09 .06 .05
CV —2.53 2.59 2.63
n1720 20
Heigh o b aincase
Mean 16.70 16.1
1
17.1
1
16.15 Sex 110.631 48.38 <.001
SD -.27 .62 .34 Wi hin 46 .220
SE —.10 .14 .08
CV -1.68 3.62 2.1 1
n1720 20
B ead h ac oss he uppe mola s
Mean 13.00 12.86 13.56 13.22 Sex 11.997 23.00 <.001
SD -.31 .24 .27 Wi hin 46 .087
SE —.12 .05 .06
CV —2.4! 1.77 2.04
n1720 20
B ead h ac oss he uppe canines
Mean 9.30 8.67 9.75 9.01 Sex 17.590 121.03 <.001
SD -.18 .23 .21 Wi hin 46 .063
SE —.07 .05 .05
CV —2.08 2.36 2.33
n1720 20
Leng h o he maxilla y oo h ow
Mean 13.10 12.70 13.33 12.81 Sex 13.414 54.65 <.001
SD -.33 .28 .19 Wi hin 46 .063
SE —.12 .06 .04
CV —2.60 2.10 1.58
n1720 20
Leng h o he uppe mola i o m oo h ow
Mean 10.20 9.83 10.09 9.91 Sex 1.480 12.24 <.001
SD -.24 .21 .17 Wi hin 46 .039
SE -.09 .05 .04
CV -2.44 2.08 1.72
n1720 20
54 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 11. —Con inued.
Caa inga Ce ado Analysis o a iance
<3(3 9$ <3.3 99 Fac o d MS FSigni icance
Wid h o he wides mola
Mean 3.30 3.41 3.43 3.32 Sex 1.066 3.52 .067
SD —.19 .15 .10 Wi hin 46 .019
SE —.07 .03 .02
CV —7.48 4.37 3.01
n1720 20
G ea es leng h o he mandible
Mean 24.90 24.1
1
25.50 24.16 Sex 120.801 90.70 <.001
SD —.27 .55 .46 Wi hin 46 .229
SE —.10 .12 .10
CV —1.12 2.16 1.90
n1720 20
Leng h o he mandibula oo h ow
Mean 14.80 14.14 15.23 14.47 Sex 18.027 84.45 <.001
SD —.35 .30 .26 Wi hin 46 .095
SE —.13 .07 .06
CV —2.48 1.96 1.80
n1720 20
Leng h o he co onoid p ocess
Mean 10.70 9.86 11.13 10.31 Sex 19.932 68.63 <.001
SD —.26 .44 .27 Wi hin 46 .145
SE -.10 .10 .06
CV —2.64 3.95 2.62
n1720 20
Table 12.— Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o/T achops ci hosus males and emales om he Caa inga biome. Aone-way analysis o
a iance (Model I) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual a ia ion is indica ed
by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
<5<5 99 <5<3 99 Fac o d MS FSigni icance
To al leng h
Mean 100.81 98.58 Sex 143.331 .70 .409
SD 7.63 8.07 Wi hin 33 61.972
SE 1.91 1.85
CV 7.57 8.19
n16 19
Tail leng h
Mean 15.75 15.63 Sex 1.122 .03 .875
SD 2.32 2.09 Wi hin 33 4.831
SE .58 .48
CV 14.73 13.37
n16 19
Hind oo leng h
Mean 16.25 16.26 Sex 1.002 .00 .963
SD .68 .93 Wi hin 33 .687
SE .17 .21
CV 4.18 5.72
n16 19
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 55
Tabic 12. —Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 9$ <5<3 9$ Fac o d MS FSigni icance
Ea leng h
Mean 32.31 33.58 Sex 1.617 .17 .681
SD 2.02 1.77 Wi hin 33 3.578
SE .51 .41
CV 6.25 5.27
n16 19
T agus leng h
Mean 13.31 13.58 Sex 1.617 .72 .401
SD .95 .90 Wi hin 33 .851
SE .24 .21
CV 7.14 6.63
n16 19
Fo ea m leng h
Mean 60.94 61.21 Sex 1.648 .27 .604
SD 1.29 1.72 Wi hin 33 2.367
SE .32 .39
CV 2.12 2.81
n16 19
Weigh
Mean 38.88 39.16 Sex 1.695 .02 .898
SD 4.46 7.68 Wi hin 33 41.221
SE 1.12 1.76
CV 11.47 19.61
n16 19
Leng h o digi one
Mean 14.19 14.95 Sex 1.001 .00 .975
SD .77 1.03 Wi hin 33 .845
SE .19 .24
CV 5.15 6.89
n16 19
Leng h o digi h ee
Mean 115.50 115.58 Sex 1.054 .00 .951
SD 3.16 4.18 Wi hin 33 14.080
SE .79 .96
CV 2.74 3.62
n16 19
Leng h o digi ou
Mean 85.06 85.68 Sex 13.357 .44 .511
SD 2.57 2.91 Wi hin 33 7.607
SE .64 .67
CV 3.02 3.40
n16 19
Leng h o digi i e
Mean 86.13 87.00 Sex 16.650 .78 .382
SD 2.75 3.04 Wi hin 33 8.477
SE .69 .70
CV 3.19 3.49
n16 19
Tibia leng h
Mean 28.13 28.00 Sex 1.136 .05 .821
SD .81 2.06 Wi hin 33 2.599
SE .20 .47
CV 2.88 7.36
n16 19
62 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 14 .—Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion. CV; sample size, n) q/'Lonchophylla mo dax males and emales om he Caa inga biome. Aone-way analysis
o a iance (Model I) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual a ia ion is indica ed
by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
66 92 66 22 Fac o d MS FSigni icance
To al leng h
Mean 64.81 66.14 Sex 131.914 2.86 .095
SD 3.47 3.19 Wi hin 70 11.142
SE .57 .54
CV 5.35 4.82
n37 35
Tail leng h
Mean 10.05 10.17 Sex 1.248 .15 .699
SD 1.37 1.18 Wi hin 70 1.641
SE .23 .20
CV 13.63 11.60
n37 35
Hind oo leng h
Mean 9.03 9.20 Sex 1.538 1.53 .220
SD .65 .53 Wi hin 70 .351
SE .1 1.09
CV 7.20 5.76
n37 35
Ea leng h
Mean 14.60 14.77 Sex 1.562 .69 .409
SD .93 .88 Wi hin 70 .816
SE .15 .15
CV 6.37 5.96
n37 35
T agus leng h
Mean 6.00 5.97 Sex 1.015 .04 .840
SD .67 .51 Wi hin 70 .357
SE .11 .09
CV 11.17 8.54
n37 35
Fo ea m leng h
Mean 34.65 35.14 Sex 14.393 4.35 .041
SD 1.14 .85 Wi hin 70 1.010
SE .19 .14
CV 3.29 2.42
n37 35
Weigh
Mean 8.32 8.87 Sex 15.384 8.14 .006
SD .60 .99 Wi hin 70 .661
SE .10 .17
CV 7.21 11.16
n37 35
Leng h o digi one
Mean 7.68 7.54 Sex 1.317 .97 .327
SD .58 .56 Wi hin 70 .326
SE .10 .10
CV 7.55 7.43
n37 35
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 63
Table 14 .—Con inued.
Caa inga Ce ado Analysis o a iance
66 99 (3<5 99 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 66.70 67.77 Sex 120.54 5.71 .020
SD 2.12 1.63 Wi hin 70 3.60
SE .35 .28
CV 3.18 2.41
n37 35
Leng h o digi ou
Mean 49.43 50.23 Sex 111.400 6.58 .012
SD 1.24 1.40 Wi hin 70 1.732
SE .20 .24
CV 2.51 2.79
n37 35
Leng h o digi i e
Mean 47.27 47.66 Sex 12.692 1.30 .259
SD 1.41 1.48 Wi hin 70 2.074
SE .23 .25
CV 2.98 3.1
1
n37 35
Tibia leng h
Mean 15.08 15.14 Sex 1.069 .11 .739
SD .68 .88 Wi hin 70 .615
SE .11 .15
CV 4.51 5.81
n37 35
Calca leng h
Mean 5.57 5.86 Sex 11.508 4.52 .037
SD .56 .60 Wi hin 70 .334
SE .09 .10
CV 10.05 10.24
n37 35
Noselea leng h
Mean 5.05 4.89 Sex 1.510 1.84 .180
SD .52 .53 Wi hin 70 .278
SE .09 .09
CV 10.30 10.84
n37 35
G ea es leng h o skull
Mean 22.58 22.59 Sex 1.001 .00 .944
SD .43 .37 Wi hin 68 .161
SE .07 .07
CV 1.90 1.64
n37 33
Condylobasal leng h
Mean 21.33 21.36 Sex 1.012 .08 .784
SD .46 .33 Wi hin 67 .163
SE .08 .06
CV 2.16 1.54
n36 33
64 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 14 .—Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 22 <5(5 22 Fac o d MS FSigni icance
Pos o bi al cons ic ion
Mean 4.22 4.30 Sex 1.109 4.12 .046
SD .13 .19 Wi hin 69 .026
SE .02 .03
CV 3.08 4.42
n37 34
Mas oid b ead h
Mean 9.08 9.01 Sex 1.068 1.79 .185
SD .16 .23 Wi hin 67 .038
SE .03 .04
CV 1.76 2.55
n36 33
B ead h o b aincase
Mean 8.39 8.31 Sex 1.114 3.68 .059
SD .19 .16 Wi hin 69 .031
SE .03 .03
CV 2.26 1.93
n37 34
Ros al b ead h
Mean 3.62 3.53 Sex 1.150 6.88 .011
SD .16 .14 Wi hin 69 .022
SE .03 .02
CV 4.42 3.97
n37 34
Heigh o b aincase
Mean 8.73 8.75 Sex 1.002 .03 .871
SD .26 .24 Wi hin 59 .062
SE .05 .05
CV 2.98 2.74
n32 29
B ead h ac oss he uppe mola s
Mean 5.10 5.14 Sex 1.024 .97 .328
SD .16 .16 Wi hin 57 .025
SE .03 .03
CV 3.14 3.1
1
n32 27
B ead h ac oss he uppe canines
Mean 3.71 3.56 Sex 1.314 14.57 <.001
SD .15 .15 Wi hin 58 .022
SE .03 .03
CV 4.04 4.21
n32 28
Leng h o he maxilla y oo h ow
Mean 7.75 7.81 Sex 1.049 1.10 .298
SD .25 .16 Wi hin 57 .045
SE .04 .03
CV 3.23 2.05
n32 27
Leng h o he uppe mola i o m oo h ow
Mean 5.82 5.86 Sex 1.028 1.01 .318
SD .18 .15 Wi hin 57 .028
SE .03 .03
CV 3.09 2.56
n32 27
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 65
Table 14. —Con inued.
Caa inga Ce ado Analysis o a iance
<53 29 <5<3 99 Fac o d MS FSigni icance
Wid h o he wides mola
Mean .92 .96 Sex 1.022 5.38 .024
SD .07 .05 Wi hin 58 .004
SE .01 .01
CV 7.61 5.21
n32 28
G ea es leng h o he mandible
Mean 15.71 15.78 Sex 1.076 .53 .470
SD .45 .29 Wi hin 69 .145
SE .07 .05
CV 3.86 1.84
n37 34
Leng h o he mandibula oo h ow
Mean 8.12 8.07 Sex 1.033 .66 .419
SD .20 .25 Wilhin 59 .050
SE .04 .05
CV 2.46 3.10
n32 29
Leng h o he co onoid p ocess
Mean 3.87 3.79 Sex 1.110 2.38 .128
SD .21 .22 Wi hin 67 .050
SE .03 .04
CV 5.43 5.80
n37 32
Caa ingas whe e i was usually associa ed wi h se -
o e habi a s, bu i was locally abundan in open
a eas o Ce adao o in Ce ado habi a s on he
Chapada do A a ipe. In he Caa ingas popula ion,
44% o he 25 cap u ed indi iduals we e males,
hence^ he sexes occu in equal p opo ion (Binomial
Tes , P>.05). In con as , only 21% o he 270
cap u ed specimens in he Ce ado we e males; his
indica es ha males and emales occu in signi i-
can ly di e en p opo ions (Binomial Tes , P<.01)
on he Chapada do A a ipe.
Mo phome ic da a a e epo ed o B azilian
specimens o A. g. geoj oyi by Dobson (1878), Lima
(1926), and Cunha Viei a (1942). Mo e ecen ly,
Ande son (1957) was unable o de ec s a is ically
signi ican seconda y sexual a ia ion in la ge sam-
ples o males and emales om Chiapas, Mexico.
He did, howe e , ind o ea m leng h and skull leng h
o be signi ican ly di e en o specimens om Sou h
Ame ica and Chiapas.
S a is ically signi ican seconda y sexual a ia ion
in specimens om he No heas o B azil is e i-
denced by h ee ex e nal and i e c anial cha ac e s;
geog aphic a ia ion be ween Caa ingas and Ce -
ado habi a s is s a is ically signi ican o wo ex-
e nal and ou c anial cha ac e s (Table 15).
Sub amily Ca olliinae
Ca ollia pe spicilla a (Linnaeus, 1758)
This ugi o e was abundan and ubiqui ous in
all habi a s o he Caa ingas and Ce ado; i was
equen ly ound oos ing in man-made s uc u es
in associa ion wi h Glossophaga so icina. O he
h ee subspecies cu en ly ecognized, C. p. pe spici-
lla a is he designa ion applied o specimens om
he No heas o B azil (Jones and Ca e , 1976).
F ui s o Vismia composed he majo po ion o his
species’ die on he Chapada do A a ipe. O he 467
cap u ed adul specimens om Ce ado and Ce -
adao habi a s, 42% we e males; he p opo ion o
males and emales we e no s a is ically equal (Bi-
nomial Tes , Pc.01). Al hough signi ican , he
magni ude o di e ence be ween males and emales
was small; his sugges s ha ac o s o he han social
s uc u e ( o example, inc eased ca chabili y o less
maneu e able p egnan emales may in la e hei
66 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 15.— Summa y s a is ics o ex e naIand c anial cha ac e s (mean; s anda d de ia ion. SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o Anou a geo oyi males and emales om bo h Caa inga and Ce ado biomes. A wo-way
analysis o a iance (Model I)wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual o geog aphic
a ia ion is indica ed by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
66 9$ 66 99 Fac o d MS FSigni icance
To al leng h
Mean 63.73 63.57 65.45 67.85 A ea 1137.287 23.85 <.001
SD 2.57 2.88 2.44 1.84 Sex 119.198 3.34 .073
SE .78 .77 .55 .41 AXS124.901 4.33 .042
CV 4.03 4.53 3.73 2.71 E o 61 5.756
n1
1
14 20 20
Hind oo leng h
Mean 10.27 10.79 10.80 10.55 A ea 1.324 .54 .466
SD .65 .58 1.11 .51 Sex 1.264 .44 .510
SE .20 .16 .25 .11 AXS12.219 3.69 .059
CV 6.33 5.38 10.28 4.83 E o 61 .602
n1 1 14 20 20
Ea leng h
Mean 14.55 14.43 14.35 14.65 A ea 1.003 .00 .949
SD 1.04 1.02 .67 .49 Sex 1.128 .21 .649
SE .31 .27 .15 .11 AXS1.663 1.08 .302
CV 7.15 7.07 4.67 3.34 E o 61 .611
n1
1
14 20 20
T agus leng h
Mean 6.27 5.71 5.90 5.70 A ea 1.571 1.66 .203
SD .65 .47 .64 .57 Sex 12.193 6.36 .014
SE .20 .13 .14 .13 AxS1.490 1.42 .238
CV 10.37 8.23 10.85 10.00 E o 61 .345
n1
1
14 20 20
Fo ea m leng h
Mean 42.82 42.14 42.80 43.00 A ea 12.683 2.46 .122
SD 1.17 .54 1.06 1.21 Sex 1.861 .79 .378
SE .35 .14 .24 .27 AXS12.921 2.68 .107
CV 2.73 1.28 2.48 2.81 E o 61 1.091
n1
1
14 20 20
Weigh
Mean 15.50 14.36 15.13 15.55 A ea 12.550 2.07 .155
SD 1.14 1.34 1.00 1.03 Sex 11.964 1.60 .21
1
SE .34 .36 .22 .23 AXS19.370 7.61 .008
CV 7.35 9.33 6.61 6.62 E o 61 1.231
n1114 20 20
Leng h o digi one
Mean 8.00 7.93 8.45 8.15 A ea 11.719 7.27 .009
SD .45 .48 .51 .49 Sex 1.526 2.22 .141
SE .14 .13 .11 .11 AXS1.199 .84 .363
CV 5.63 6.05 6.04 6.01 E o 6
1
.237
n1
1
14 20 20
Leng h o digi h ee
Mean 84.91 84.50 85.75 85.50 A ea 112.918 1.68 .200
SD 4.23 1.99 2.63 2.37 Sex 11.656 .22 .644
SE 1.28 .53 .59 .53 AxS1.097 .01 .911
CV 4.98 2.36 3.07 2.77 E o 61 7.691
n1114 20 20
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 67
Table 15 .—Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 92 66 22 Fac o d MS FSigni icance
Leng h o digi ou
Mean 62.00 60.36 61.65 61.10 A ea 1.588 .17 .678
SD 1.84 1.74 2.01 1.71 Sex 118.330 5.44 .023
SE .56 .46 .45 .38 AXS14.553 1.35 .250
c 2.97 2.88 3.26 2.80 E o 61 3.370
n1
1
14 20 20
Leng h o digi i e
Mean 54.82 52.71 54.20 53.40 A ea 1.017 .00 .946
SD 1.54 1.94 2.44 1.57 Sex 132.144 8.43 .005
SE .46 .52 .55 .35 AXSI6.481 1.70 .197
CV 2.81 3.68 4.50 2.94 E o 61 3.811
n1
1
14 20 20
Tibia leng h
Mean 16.64 16.21 16.55 16.65 A ea 1.465 .88 .353
SD .67 .43 .89 .75 Sex 1.395 .74 .392
SE .20 .11 .20 .17 AxS11.039 1.96 .167
CV 4.03 2.65 5.38 4.50 E o 61 .531
n1
1
14 20 20
Calca leng h
Mean 3.27 3.21 3.10 2.95 A ea 1.728 2.30 .134
SD .65 .70 .45 .5! Sex 1.166 .52 .472
SE .20 .19 .10 .11 AXS1.032 .10 .752
CV 19.88 21.81 14.52 17.29 E o 61 .316
n1 1 14 20 20
Noselea leng h
Mean 3.82 3.71 4.10 3.90 A ea 1.833 3.16 .081
SD .60 .47 .55 .45 Sex 1.352 1.33 .253
SE .18 .13 .12 .10 AxS1.035 .13 .716
CV 15.71 12.67 13.41 11.54 E o 61 .264
n1114 20 20
G ea es leng h o skull
Mean 24.76 24.34 24.95 24.83 A ea 11.740 8.04 .006
SD .43 .34 .62 .38 Sex 11.165 5.38 .024
SE .13 .09 .14 .08 AxS1.350 1.62 .209
CV 1.74 1.40 2.48 1.53 E o 61 .217
n1
1
14 20 20
Condylobasal leng h
Mean 24.10 23.67 24.1 124.06 A ea 1.591 2.54 .116
SD .39 .37 .64 .40 Sex 1.891 3.84 .055
SE .12 .10 .14 .09 AXS1.532 2.29 .136
CV 1.62 1.53 2.65 1.66 E o 61 .232
n1114 20 20
Zygoma ic b ead h
Mean 10.85 10.83 10.94 10.86 A ea 1.031 .44 .513
SD .35 .23 .26 .19 Sex 1.019 .28 .602
SE .12 .10 .07 .07 AxS1.083 .12 .732
CV 3.23 2.12 2.38 1.75 E o 32 .070
n8 6 14 8
68 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 15 .—Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 $9 <5<5 99 Fac o d MS FSigni icance
Pos o bi al cons ic ion
Mean 5.13 5.04 5.19 5.17 A ea 1.130 3.08 .085
SD .16 .24 .23 .18 Sex 1.038 .89 .349
SE .05 .06 .05 .04 AxS1.018 .43 .513
CV 3.12 4.76 4.43 3.48 E o 61 .042
n1
1
14 20 20
Mas oid b ead h
Mean 10.30 10.12 10.36 10.30 A ea 1.217 2.69 .106
SD .26 .28 .28 .30 Sex 1.217 2.69 .106
SE .08 .08 .06 .07 AXS1.054 .67 .418
CV 2.52 2.77 2.70 2.91 E o 61 .081
n1
1
14 20 20
B ead h o b aincase
Mean 9.71 9.61 9.78 9.80 A ea 1.255 5.20 .026
SD .12 .24 .21 .26 Sex 1.023 .46 .500
SE .04 .06 .05 .06 AXS1.061 1.25 .268
CV 1.24 2.50 2.15 2.65 E o 61 .049
n1
1
14 20 20
Ros al b ead h
Mean 4.28 4.14 4.31 4.16 A ea 1.009 .40 .532
SD .17 .17 .13 .14 Sex 1.323 14.86 <.001
SE .05 .05 .03 .03 AXS1.000 .00 .988
CV 3.97 4.1
1
3.02 3.37 E o 61 .022
n1
1
14 20 20
Heigh o b aincase
Mean 9.92 9.66 9.89 9.94 A ea 1.214 3.91 .053
SD .14 .29 .23 .23 Sex 1.156 2.86 .096
SE .05 .08 .05 .05 AXS1.363 6.64 .013
CV 1.41 3.00 3.33 2.31 E o 61 .055
n914 20 20
B ead h ac oss he uppe mola s
Mean 6.24 6.15 6.24 6.21 A ea 1.010 .30 .587
SD .10 .1
6
.22 .19 Sex 1.053 1.53 .221
SE .03 .05 .05 .04 AXS1.019 .54 .465
CV 1.60 2.60 3.53 3.06 E o 58 .034
n913 20 20
B ead h ac oss he uppe canines
Mean 4.93 4.52 5.02 4.66 A ea 1.166 6.79 .102
SD .16 .17 .16 .14 Sex 12.033 83.24 <.001
SE .05 .05 .04 .03 AXS1.011 .43 .513
CV 3.25 3.76 3.19 3.00 E o 58 .024
n913 20 20
Leng h o he maxilla y oo h ow
Mean 9.41 9.19 9.44 9.39 A ea 1.183 2.76 .102
SD .31 .19 .26 .27 Sex 1.256 3.88 .054
SE .1 1.05 .06 .06 AXS1.114 1.73 .193
CV 3.29 2.07 2.75 2.88 E o 58 .066
n 9 13 20 20
Leng h o he uppe mola i o m oo h ow
Mean 7.87 7.68 7.87 7.87 A ea 1.121 1.95 .168
SD .27 .25 .24 .25 Sex 1.125 2.02 .161
SE .09 .07 .05 .06 AXS1.125 2.02 .161
CV 3.43 3.26 3.05 3.18 E o 58 .062
n9 13 20 20
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 69
Table 15. —Con inued.
Caa inga Ce ado Analysis o a iance
66 $9 66 99 Fac o d MS FSigni icance
Wid h o he wides mola
Mean 1.19 1.17 1.14 1.19 A ea 1.004 1.05 .309
SD .06 .05 .06 .07 Sex 1.002 .62 .436
SE .02 .01 .01 .02 AXS1.015 4.01 .050
c 5.04 4.27 5.26 5.88 E o 58 .004
n913 20 20
G ea es leng h o he mandible
Mean 8.08 7.74 8.17 8.20 A ea 11.113 8.34 .005
SD .40 .32 .44 .29 Sex 1.376 2.82 .099
SE .12 .08 .10 .07 AXS1.505 3.78 .056
CV 4.95 4.13 5.39 3.54 E o 6
1
.134
n1
1
14 20 20
Leng h o he mandibula oo h ow
Mean 9.88 9.62 9.95 9.78 A ea 1.195 3.27 .076
SD .28 .19 .23 .27 Sex 1.649 10.90 .002
SE .09 .05 .05 .06 AXS1.030 .50 .483
CV 2.83 1.98 2.31 2.76 E o 58 .060
n913 20 20
Leng h o he co onoid p ocess
Mean 4.38 3.92 4.34 4.13 A ea 1.094 1.93 .170
SD .31 .18 .18 .23 Sex 11.713 35.27 <.001
SE .09 .05 .04 .05 AXS1.239 4.92 .030
CV 7.08 4.60 4.15 5.67 E o 6
1
.049
n1
1
14 20 20
sample equency, esul ing in unequal sex a ios in
la ge samples) a ec he obse ed p opo ion o he
sexes. In he Caa ingas, 47% o he 366 cap u ed
adul specimens we e males; he sexes occu ed in
s a is ically equal equency (Binomial Tes , P>
.05).
Al hough ag ea deal o in o ma ion is a ailable
on he mo phome ic cha ac e is ics o C. pe spi-
cilla a (see Swanepoel and Genoways, 1979), li le
has been published on specimens om B azil. Hahn
(1907), Cunha Viei a (1942), Pine (1972), Pi lo
(1972), and Taddei (1975Z?) ha e epo ed mea-
su emen s om B azil, bu only Taddei (
1
915b) has
pe o med igo ous s a is ical analyses. He ound
ha emales a e la ge on he a e age han males
when conside ing ex e nal cha ac e s, bu s a is i-
cally di e en o only ou o he 17cha ac e s (head-
body leng h, ea leng h, o ea m leng h, and leng h
o me aca pal II). Con e sely, on he a e age, males
a e la ge han emales when conside ing ag oup o
15c anial cha ac e s, bu only one cha ac e , mas-
oid b ead h, is s a is ically signi ican ly di e en .
Tamsi and Valdi ieso (1963) s a e ha males and
emales om Colombia do no di e in size, and
ha specimens on opposi e sides o he Andes do
no di e in any cha ac e .
Only wo ex e nal cha ac e s ( o al leng h and
weigh ) exhibi s a is ically signi ican sexual a i-
a ion in specimens om No heas B azil, and no
end is appa en wi h ega d o a e age size ela-
ionships be ween he sexes. Howe e , when con-
side ing c anial cha ac e is ics, he sample means o
males a e gene ally la ge han hose o emales and
o six cha ac e s, s a is ically signi ican di e ences
a e de ec ed (see Table 16). Caa ingas specimens
ha e la ge sample means han Ce ado specimens
o 11o he 15analyzed c anial cha ac e s; s a is-
ically signi ican di e ences a e indica ed o six
cha ac e s (Table 16). Size ends o ex e nal cha -
ac e s a e no appa en and only wo cha ac e s ( a-
gus leng h and weigh ) ha e s a is ically dis inguish-
able means.
Sub amily S enode ma inae
S u ni a lilium (E. Geo oy, 1810)
O he six en a i ely ecognized subspecies, V. /.
lilium is he designa ion p ope ly applied o speci-
mens om he No heas o B azil (Jones and Ca -
70 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 16. —Summa y s a is ics o ex e nal and c ania Icha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o Ca ollia pe spicilla a males and emlaes om bo h Caa inga and Ce ado biomes. A wo-
way analysis o a iance (Model I) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual o
geog aphic a ia ion is indica ed by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
<53 2$ 33 9$ Fac o d MS FSigni icance
To al leng h
Mean 72.80 75.65 71.95 73.45 A ea 146.513 3.49 .066
SD 3.76 4.02 2.80 3.90 Sex 194.613 7.09 .009
SE .84 .90 .63 .87 AxS19.113 .68 .411
CV 5.16 5.31 3.89 5.31 E o 76 13.338
n20 20 20 20
Tail leng h
Mean 10.70 11.05 10.75 11.30 A ea 1.450 .15 .700
SD 1.92 1.85 1.68 1.46 Sex 14.050 1.34 .250
SE .43 .41 .38 .33 AXS1.200 .07 .797
CV 17.94 16.74 15.63 12.92 E o 76 3.015
n20 20 20 20
Hind oo leng h
Mean 11.85 11.65 11.60 11.75 A ea 1.113 .15 .701
SD .88 .59 1.05 .91 Sex 1.013 .02 .898
SE .20 .13 .23 .20 AXS1.613 .81 .372
CV 7.43 5.06 9.05 7.74 E o 76 .759
n20 20 20 20
Ea leng h
Mean 19.55 19.30 19.00 19.15 A ea 12.450 2.46 .121
SD 1.10 1.03 1.03 .81 Sex 1.050 .05 .823
SE .25 .23 .23 .18 AXS1.800 .80 .373
CV 5.63 5.34 5.42 4.23 E o 76 .996
n20 20 20 20
T agus leng h
Mean 7.70 7.90 8.35 8.30 A ea I5.513 13.62 <.001
SD .66 .64 .49 .73 Sex 1.113 .28 .600
SE .15 .14 .11 .16 AXS1.313 .77 .382
CV 8.57 8.10 5.87 8.80 E o 76 .405
n20 20 20 20
Fo ea m leng h
Mean 42.70 42.95 42.25 42.70 A ea 12.450 1.11 .295
SD 1.87 1.10 1.37 1.49 Sex 12.450 1.11 .295
SE .42 .25 .31 .33 AXS1.200 .09 .764
CV 4.38 2.56 3.24 3.49 E o 76
n20 20 20 20
Weigh
Mean 18.72 20.88 18.50 18.70 A ea 128.561 6.02 .016
SD 1.35 3.30 1.33 2.1 1Sex 127.848 5.87 .018
SE .30 .74 .30 .47 AXS119.208 4.05 .048
CV 7.21 15.80 7.19 11.28 E o 76 4.742
n20 20 20 20
Leng h o digi one
Mean 10.85 10.95 11.10 11.30 A ea 11.800 4.97 .029
SD .49 .69 .64 .57 Sex 1.450 1.24 .268
SE .1 1.15 .14 .13 AXS1.050 .14 .711
CV 4.52 6.30 5.77 5.04 E o 76 .362
n20 20 20 20
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 71
Table 16. —Con inued.
Caa inga Ce ado Analysis o a iance
<36 99 66 99 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 89.60 90.80 89.65 90.50 A ea 1.313 .03 .860
SD 3.24 2.93 3.63 2.78 Sex 121.013 2.10 .151
SE .72 .66 .81 .62 AXS1.613 .06 .805
c 3.62 3.23 4.05 3.07 E o 76 9.994
n20 20 20 20
Leng h o digi ou
Mean 65.35 65.50 65.50 65.50 A ea 1.113 .03 .860
SD 1.69 1.85 2.07 1.93 Sex 1.113 .03 .860
SE .38 .41 .46 .43 AXS1.113 .03 .860
CV 2.59 2.82 3.16 2.95 E o 76 3.573
n20 20 20 20
Leng h o digi i e
Mean 64.50 64.30 64.00 64.95 A ea 1.113 .03 .871
SD 1.73 2.08 2.56 1.79 Sex 12.813 .66 .419
SE .39 .47 .57 .40 AxS16.613 1.55 .217
CV 2.68 3.23 4.00 2.76 E o 76 4.265
n20 20 20 20
Tibia leng h
Mean 19.80 19.60 19.70 19.60 A ea 1.050 .04 .851
SD .83 .82 1.38 1.54 Sex 1.450 .32 .574
SE .19 .19 .31 .34 AXS1.050 .04 .851
CV 4.19 4.18 7.01 7.86 E o 76 1.408
n20 20 20 20
Calca leng h
Mean 7.70 7.95 7.50 8.15 A ea 1.000 .000 1.000
SD .87 1.23 .89 1.04 Sex 14.050 3.91 .052
SE .19 .28 .20 .23 AxS1.800 .77 .382
CV 11.30 15.47 11.87 12.76 E o 76 1.036
n20 20 20 20
Noselea leng h
Mean 6.60 6.65 6.45 6.40 A ea 1.800 1.73 .192
SD .75 .75 .61 .60 Sex 1.000 .000 1.000
SE .17 .17 .14 .13 AXS1.050 .11 .743
CV 11.36 11.28 9.46 9.38 E o 76 .462
n20 20 20 20
G ea es leng h o skull
Mean 22.26 22.00 22.10 21.77 A ea 1.722 3.99 .049
SD .43 .28 .45 .52 Sex 11.741 9.61 .003
SE .10 .06 .10 .12 AXS1.025 .14 .714
CV 1.93 1.27 2.04 2.39 E o 76 .181
n20 20 20 20
Condylobasal leng h
Mean 20.23 20.03 20.05 19.81 A ea 1.780 4.33 .041
SD .46 .36 .35 .51 Sex 1.946 5.25 .025
SE .10 .08 .08 .11 AxS1.010 .06 .813
CV 2.27 1.80 1.75 2.57 E o 76 .180
n20 20 20 20
Pos o bi al cons ic ion
Mean 5.68 5.55 5.63 5.51 A ea 1.037 .79 .376
SD .19 .20 .18 .27 Sex 1.300 6.59 .012
SE .04 .05 .04 .06 AXS1.001 .02 .876
CV 3.35 3.60 3.20 4.90 E o 76 .046
n20 20 20 20
78 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 18 .—Summa y s a is ics o ex e nal and c ania Icha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o/'Vampy ops linea us males and emales om bo h Caa inga and Ce ado biomes. A wo-
way analysis o a iance (Model I) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual o
geog aphic a ia ion is indica ed by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
<5(3 99 <5<5 99 Fac o d MS FSigni icance
To al leng h
Mean 64.40 67.85 63.55 66.70 A ea 120.000 2.63 .109
SD 2.98 2.60 2.78 2.66 Sex 1217.800 28.61 <.001
SE .67 .58 .62 .59 AXS1.450 .06 .809
CV 4.63 3.83 4.37 3.99 E o 76 7.612
n20 20 20 20
Hind oo leng h
Mean 11.20 11.60 11.55 11.90 A ea 12.1 13 7.06 .010
SD .62 .60 .51 .45 Sex 12.813 9.40 .003
SE .14 .13 .11 .10 AXS1.013 .04 .839
CV 5.54 5.17 4.42 3.87 E o 76 .299
n20 20 20 20
Ea leng h
Mean 18.05 18.15 17.80 18.15 A ea 1.313 .45 .506
SD 1.05 .67 .89 .67 Sex 11.013 1.45 .233
SE .24 .15 .20 .15 AXS1.313 .45 .506
CV 5.82 3.69 5.00 3.69 E o 76 .701
n20 20 20 20
T agus leng h
Mean 7.15 7.35 7.30 7.10 A ea 1.050 .11 .736
SD .75 .59 .57 .72 Sex 1.000 .00 1.000
SE .17 .13 .13 .16 AXS1.800 1.84 .179
CV 10.49 8.03 7.81 10.14 E o 76 .436
n20 20 20 20
Fo ea m leng h
Mean 47.20 47.45 46.00 47.20 A ea 110.513 4.09 .047
SD 1.94 2.04 1.12 1.06 Sex 110.513 4.09 .047
SE .43 .46 .25 .24 AxS14.513 1.76 .189
CV 4.1 14.30 2.43 2.25 E o 76 2.570
n20 20 20 20
Weigh
Mean 23.93 26.44 23.10 26.48 A ea 13.121 .46 .501
SD 1.32 3.73 1.78 2.90 Sex 1173.461 25.45 <.001
SE .30 .84 .40 .65 AX S13.698 .54 .464
CV 5.52 14.1 17.71 10.95 E o 76 6.815
n20 20 20 20
Leng h o digi one
Mean 10.65 10.95 10.60 11.10 A ea 1.050 .09 .759
SD .75 .76 .75 .64 Sex 13.200 6.06 .016
SE .17 .17 .17 .14 AXS1.200 .38 .540
CV 7.04 6.94 7.08 5.77 E o 76 .528
n20 20 20 20
Leng h o digi h ee
Mean 96.80 97.10 94.00 97.35 A ea 132.513 3.37 .070
SD 3.49 3.70 2.64 2.41 Sex 166.613 6.90 .010
SE .78 .83 .59 .54 AXS146.513 4.82 .031
CV 3.61 3.81 2.81 2.48 E o 76 9.652
n20 20 20 20
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 79
Table 18 .—Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 99 <5<3 99 Fac o d MS FSigni icance
Leng h o digi ou
Mean 70.95 71.90 69.35 71.05 A ea 130.013 6.48 .013
SD 2.04 2.55 2.03 1.93 Sex 135.1 13 7.58 .007
SE .46 .57 .46 .43 AxS12.813 .61 .438
c 2.88 3.55 2.93 2.72 E o 76 4.635
n20 20 20 20
Leng h o digi i e
Mean 65.75 67.25 64.35 66.10 A ea 132.513 7.10 .009
SD 1.94 2.86 1.90 1.65 Sex 152.813 11.54 .001
SE .44 .64 .43 .37 AxS1.313 .07 .795
CV 2.95 4.25 2.95 2.50 E o 76 4.577
n20 20 20 20
Tibia leng h
Mean 18.00 18.35 17.95 18.50 A ea 1.050 .09 .766
SD .46 .88 .76 .83 Sex 14.050 7.24 .009
SE .10 .20 .17 .19 AX S1.200 .36 .552
CV 2.56 4.80 4.23 4.49 E o 76 .559
n20 20 20 20
Calca leng h
Mean 4.75 4.30 4.25 4.30 A ea 11.250 3.41 .069
SD .85 .57 .44 .47 Sex 1.800 2.18 .144
SE .19 .13 .10 .10 AXS11.250 3.41 .069
CV 17.89 13.26 10.35 10.93 E o 76 .367
n20 20 20 20
Noselea leng h
Mean 6.95 7.00 6.85 7.25 A ea 1.113 .18 .672
SD .76 .86 .88 .64 Sex 11.013 1.63 .206
SE .17 .19 .20 .14 AXS1.613 .99 .324
CV 10.94 12.29 12.85 8.83 E o 76 .623
n20 20 20 20
G ea es leng h o skull
Mean 24.32 24.40 24.38 24.41 A ea 1.018 .09 .762
SD .46 .37 .56 .35 Sex 1.061 .31 .579
SE .10 .08 .13 .08 AXS1.013 .06 .801
CV 1.89 1.52 2.30 1.43 E o 76 .195
n20 20 20 20
Condylobasal leng h
Mean 21.90 22.00 21.83 21.89 A ea 1.181 .78 .379
SD .44 .46 .61 .38 Sex 1.128 .56 .459
SE .10 .10 .14 .09 AXS1.005 .02 .889
CV 2.01 2.09 2.79 1.74 E o 76 .231
n20 20 20 20
Zygoma ic b ead h
Mean 14.32 14.37 14.25 14.17 A ea 1.365 2.61 .111
SD .34 .40 .37 .39 Sex 1.005 .03 .858
SE .08 .09 .08 .09 AXS1.098 .70 .405
CV 2.37 2.78 2.60 2.75 E o 76 .140
n20 20 20 20
Pos o bi al cons ic ion
Mean 6.26 6.29 6.23 6.22 A ea 1.045 1.07 .304
SD .19 .20 .18 .25 Sex 1.001 .03 .871
SE .04 .04 .04 .06 AXS1.010 .24 .626
CV 3.04 3.18 2.89 4.02 E o 76 .042
n20 20 20 20
80 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 18 .—Con inued.
Caa inga Ce ado Analysis o a iance
<56 92 66 22 Fac o d MS FSigni icance
Mas oid b ead h
Mean 12.12 12.22 12.21 12.16 A ea 1.005 .05 .822
SD .26 .31 .24 .32 Sex 1.013 .14 .707
SE .06 .07 .07 .07 AXS1.113 1.28 .262
CV 2.15 2.54 1.97 2.63 E o 76 .088
n20 20 20 20
B ead h o b aincase
Mean 10.51 10.56 10.56 10.58 A ea 1.021 .37 .544
SD .29 .26 .23 .16 Sex 1.021 .37 .544
SE .06 .06 .05 .04 AXS1.003 .06 .815
CV 2.76 2.46 2.18 1.51 E o 76 .057
n20 20 20 20
Ros al b ead h
Mean 7.46 7.42 7.03 6.86 A ea 14.950 57.83 <.001
SD .26 .31 .36 .21 Sex 1.210 2.45 .121
SE .06 .07 .08 .05 AXS1.078 .91 .342
CV 3.49 4.18 5.12 3.06 E o 76 .086
n20 20 20 20
Heigh o b aincase
Mean 12.48 12.42 12.44 12.36 A ea 1.045 .63 .429
SD .23 .31 .29 .24 Sex 1.105 1.48 .228
SE .05 .07 .07 .05 AXS 1.001 .02 .900
CV 1.84 2.50 2.33 1.94 E o 76 .071
n20 20 20 20
B ead h ac oss he uppe mola s
Mean 10.19 10.40 10.17 10.13 A ea 1.406 5.92 .017
SD .23 .31 .25 .25 Sex 1.153 2.23 .139
SE .05 .07 .06 .06 AXS1.325 4.74 .033
CV 2.26 2.98 2.46 2.47 E o 76 .069
n20 20 20 20
B ead h ac oss he uppe canines
Mean 6.05 6.15 6.09 6.02 A ea 1.045 1.08 .302
SD .22 .23 .15 .21 Sex 1.006 .15 .703
SE .05 .05 .03 .05 AXS1.153 3.67 .059
CV 3.64 3.74 2.46 3.49 E o 76 .042
n20 20 20 20
Leng h o he maxilla y oo h ow
Mean 8.59 8.75 8.76 8.62 A ea 1.010 .11 .738
SD .33 .32 .28 .27 Sex 1.001 .01 .911
SE .07 .07 .06 .06 AXS1.435 4.83 .031
CV 3.84 3.66 3.20 3.13 E o 76 .090
n 20 20 20 20
Leng h o he uppe mola i o m oo h ow
Mean 7.09 7.16 7.05 7.98 A ea 1.231 3.95 .050
SD .26 .28 .21 .20 Sex 1.000 .00 .963
SE .06 .06 .05 .05 AXS1.105 1.80 .184
CV 3.67 3.91 2.98 2.51 E o 76 .059
n20 20 20 20
Wid h o he wides mola
Mean 2.51 2.53 2.42 2.45 A ea 1.145 8.03 .006
SD .1 1.13 .14 .16 Sex 1.018 1.00 .320
SE .02 .03 .03 .04 AXS1.001 .03 .868
CV 4.38 5.14 5.79 6.53 E o 76 .018
n20 20 20 20
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 81
Table 18.—Con inued.
Caa inga Ce ado Analysis o a iance
66 22 66 22 Fac o d MS FSigni icance
G ea es leng h o he mandible
Mean 16.62 16.74 16.49 16.49 A ea 1.722 4.73 .033
SD .48 .28 .45 .32 Sex 1.072 .47 .494
SE .11 .06 .10 .07 AxS1.085 .55 .459
CV 2.89 1.67 2.73 1.94 E o 76 .153
n20 20 20 20
Leng h o he mandibula oo h ow
Mean 9.53 9.60 9.54 9.46 A ea 1.098 1.25 .267
SD .26 .33 .25 .26 Sex 1.001 .01 .937
SE .06 .08 .06 .06 AXS1.113 1.44 .235
CV 2.73 3.44 2.62 2.75 E o 76 .078
n20 20 20 20
Leng h o he co onoid p ocess
Mean 5.69 5.87 5.76 5.74 A ea 1.018 .28 .595
SD .25 .25 .29 .21 Sex 1.128 2.03 .159
SE .06 .06 .07 .05 AxS1.181 2.86 .095
CV 4.39 4.26 5.03 3.66 E o 76 .063
n20 20 20 20
Table 19 .—Summa y s a is ics o ex e naIand c anial cha ac e s (mean; s anda d de ia ion, SD: s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o A ibeus concolo males and emales om he Ce ado biome. Aone-way analysis o
a iance (Model I) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual a ia ion is indica ed
by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
66 22 66 22 Fac o d MS FSigni icance
To al leng h
Mean 58.33 61.50 Sex 130.083 7.02 .024
SD 2.07 2.07 Wi hin 10 4.283
SE .84 .85
CV 3.55 3.37
n66
Hind oo leng h
Mean 10.33 10.33 Sex 1.000 .000 1.000
SD .52 .52 Wi hin 10 .267
SE .21 .21
CV 5.03 5.03
n66
Ea leng h
Mean 17.00 17.00 Sex 1.000 .000 1.000
SD .63 .63 Wi hin 10 .400
SE .26 .26
CV 3.71 3.71
n6 6
T agus leng h
Mean 6.50 7.50 Sex 13.000 6.00 .034
SD .84 .55 Wi hin 10 .500
SE .34 .22
CV 12.92 7.33
n66
82 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 19 —Con inued.
Caa inga Ce ado Analysis o a iance
36 92 33 92 Fac o d MS FSigni icance
Fo ea m leng h
Mean 47.00 47.33 Sex 1.333 .09 .771
SD 2.19 1.63 Wi hin 10 3.733
SE .89 .67
CV 4.66 3.44
n66
Weigh
Mean 18.42 21.42 Sex 127.000 16.45 .002
SD .86 1.59 Wi hin 10 1.642
SE .35 .65
CV 4.67 7.42
n 6 6
Leng h o digi one
Mean 10.67 10.17 Sex 1.750 1.22 .296
SD .52 .98 Wi hin 10 .617
SE .21 .41
CV 4.87 9.64
n6 6
Leng h o digi h ee
Mean 96.33 99.50 Sex 130.083 2.02 .186
SD 4.68 2.81 Wi hin 10 14.883
SE 1.91 1.15
CV 4.86 2.82
n66
Leng h o digi ou
Mean 73.17 73.50 Sex 1.333 .04 .850
SD 3.49 2.35 Wi hin 10 8.833
SE 1.42 .96
CV 4.77 3.20
n66
Leng h o digi i e
Mean 69.17 68.67 Sex 1.750 .07 .794
SD 4.12 1.97 Wi hin 10 10.417
SE 1.68 .80
CV 5.96 2.87
n66
Tibia leng h
Mean 18.50 18.33 Sex 1.083 .08 .787
SD 1.05 1.03 Wi hin 10 1.083
SE .43 .42
CV 5.68 5.62
n66
Calca leng h
Mean 5.67 6.67 Sex 13.000 6.43 .030
SD .82 .52 Wi hin 10 .467
SE .33 .21
CV 14.46 7.80
n66
Noselea leng h
Mean 6.67 7.33 Sex 11.333 5.00 .049
SD .52 .52 Wi hin 10 .267
SE .21 .21
CV 7.80 7.09
n66
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 83
Table 19 .—Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 99 <5.5 99 Fac o d MS FSigni icance
G ea es leng h o skull
Mean 21.22 21.23 Sex 1.001 .00 .957
SD .40 .40 Wi hin 9.160
SE .18 .17
CV 1.89 1.88
n56
Condylobasal leng h
Mean 18.84 19.18 Sex 1.322 3.21 .107
SD .35 .29 Wi hin 9.100
SE .16 .12
CV 1.86 1.51
n56
Zygoma ic b ead h
Mean 13.00 13.17 Sex 1.076 1.19 .304
SD .19 .29 Wi hin 9.064
SE .08 .12
CV 1.46 2.20
n56
Pos o bi al cons ic ion
Mean 5.44 5.47 Sex 1.002 .20 .662
SD .09 .10 Wi hin 9.010
SE .04 .04
CV 1.65 1.83
n56
Mas oid b ead h
Mean 11.12 11.53 Sex 1.466 11.00 .009
SD .13 .25 Wi hin 9.042
SE .06 .10
CV 1.17 2.17
n56
B ead h o b aincase
Mean 9.90 9.80 Sex 1.027 .35 .568
SD .31 .25 Wi hin 9.078
SE .14 .10
CV 3.13 2.55
n56
Ros al b ead h
Mean 6.96 7.43 Sex 1.61 114.27 .004
SD .20 .22 Wi hin 9.043
SE .09 .09
CV 2.87 2.96
n56
Heigh o b aincase
Mean 11.60 11.42 Sex 1.092 1.02 .339
SD .27 .32 Wi hin 9.090
SE .12 .13
CV 2.33 2.80
n56
B ead h ac oss he uppe mola s
Mean 9.14 9.48 Sex 1.322 6.29 .034
SD .20 .25 Wi hin 9.051
SE .09 .10
CV 2.19 2.64
n56
84 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 19 .—Con inued.
Caa inga Ce ado Analysis o a iance
66 99 66 99 Fac o d MS FSigni icance
B ead h ac oss he uppe canines
Mean 5.78 5.87 Sex 1.021 .24 .639
SD .21 .35 Wi hin 9.087
SE .09 .14
CV 3.63 5.96
n56
Leng h o he ma
?
dlla y oo h ow
Mean 6.82 7.02 Sex 1.106 2.28 .165
SD .26 .17 Wi hin 9.046
SE .12 .07
CV 3.81 2.42
n56
Leng h o he uppe mola i o m oo h ow
Mean 5.68 6.02 Sex 1.309 5.00 .052
SD .18 .29 Wi hin 9.062
SE .08 .12
CV 3.17 4.82
n56
Wid h o he wides mola
Mean 2.38 2.45 Sex 1.013 1.91 .200
SD .08 .08 Wi hin 9.007
SE .04 .03
CV 3.36 3.26
n56
G ea es leng h o he mandible
Mean 13.84 13.98 Sex 1.056 .55 .478
SD .37 .27 Wi hin 9.102
SE .16 .11
CV 2.67 1.93
n56
Leng h o he mandibula oo h ow
Mean 7.52 7.63 Sex 1.035 .45 .519
SD .37 .18 Wi hin 9.078
SE .17 .07
CV 4.92 2.36
n56
Leng h o he co onoid p ocess
Mean 5.56 5.65 Sex 1.022 .57 .468
SD .11 .24 Wi hin 9.039
SE .05 .10
CV 1.98 4.25
n56
p obable sys ema ic ela ions), hence i seems p e-
ma u e o assign asubspeci ic designa ion o No h-
eas B azilian popula ions. This species was abun-
dan and ubiqui ous in bo h Ce ado and Ce adao
habi a s on he Chapada do A a ipe; Vismia was
he p edominan ui in i s die he e. I was p esen
in he Caa ingas and eached i s highes densi y on
o nea se o es. The p opo ion o males was no
equal o he p opo ion o emales wi hin ei he Caa-
ingas o Ce ado biomes (Binomial Tes s, P<.01).
Twen y-eigh pe cen o he 204 cap u ed adul
specimens om he Caa ingas we e males, whe eas
40% o he 270 cap u ed adul specimens om he
Ce ado we e males.
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 85
Aple ho a o e e ences including mo phome ic
da a on A. jamaicensis a e ci ed by Swanepoel and
Genoways (1979), bu only Rehn (1900, 1902), G.
M. Allen (1908), Ande sen (1908), and Cunha Viei a
(1942) epo measu emen s om B azilian speci-
mens. Da is (1970) examined indi idual, seconda y
sexual, and geog aphic a ia ion in specimens o A.
jamaicensis om Middle Ame ica, bu no one has
examined a ia ion in as a is ically igo ous man-
ne o Sou h Ame ican specimens.
Table 20 de ails he esul s o s a is ical analyses
o indi idual, seconda y sexual and geog aphic a i-
a ion in B azilian specimens o A. jamaicensis om
Caa ingas and Ce ado biomes. S a is ically signi -
ican geog aphic a ia ion was de ec ed o se en
ex e nal and en c anial cha ac e s; in all o hese
cases, he sample means o Caa ingas specimens
a e la ge han he sample means o Ce ado spec-
imens. These wo dis inc ye geog aphically p ox-
ima e popula ions a e clea ly dimo phic. Seconda y
sexual a ia ion is no appa en o A. jamaicensis
when conside ing ex e nal mo phome ic a iables
(only weigh is s a is ically signi ican ); howe e ,
males ha e la ge sample means han emales o
ele en c anial cha ac e s and he a ia ion is s a is-
ically signi ican o i e o hem. Females, on he
o he hand, we e ne e la ge han males when con-
side ing sample means o c anial cha ac e s. The e
is hen, app eciable seconda y sexual a ia ion ex-
hibi ed by c anial cha ac e s and s ong e idence
o sexual dimo phism o size wi h males being
la ge han emales. This is unlike he esul s ob-
ained by Da is (1970) who ound no signi ican
seconda y sexual a ia ion in samples om nu-
me ous locali ies in Cen al Ame ica. This ailu e
o de ec signi ican a ia ion was mos likely caused
by he la ge quan i y o a ia ion a ibu able o
geog aphic he e ogenei y in Da is’ samples which
was no s a is ically emo ed om his analysis o
seconda y sexual a ia ion.
A ibeus li u a us (Ol e s, 1818)
In aspeci ic ela ionships o his species a e
poo ly unde s ood a he p esen (see o example
Koopman, 1976); as such, i is bes o a oid using
ap o isional subspeci ic appella ion o popula ions
om he B azilian No heas . This ugi o e was
widesp ead and abundan in bo h Ce ado and Ce -
adao habi a s; Vismia was he p edominan ui
in i s die on he Chapada do A a ipe. I was wide-
sp ead bu uncommon in he Caa ingas. Wi hin bo h
Caa ingas and Ce ado biomes, he sexes occu ed
in equal equency (Binomial Tes , P>.05). O he
44 cap u ed adul specimens om he Caa ingas,
45% we e males. Simila ly, 58% o he 366 cap u ed
adul s om he Ce ado we e males.
Mo phome ic da a on A. li u a us appea s in he
li e a u e many imes (see Swanepoel and Geno-
ways, 1979), bu only G. M. Allen (1908), Ande sen
(1908), Cunha Viei a (1942), and Pi lo (1972) con-
side B azilian specimens. Seconda y sexual a ia-
ion in A. li u a us was examined by Ande son (1960)
and Tamsi and Valdi ieso (1963), bu only he
o me pe o med s a is ical analyses, and he e was
no signi ican seconda y sexual a ia ion de ec ed
in ha s udy. Geog aphic a ia ion in mensu al
cha ac e s o A. li u a us was examined by Dalques
(1950) and Tamsi and Valdi ieso (1963) bu no
s a is ical esul s a e epo ed. Fu he , indi idual
a ia ion in mo phome ic cha ac e s has no been
in es iga ed in his species.
The esul s o as a is ical analysis o indi idual,
seconda y sexual, and geog aphic a ia ion o pop-
ula ions o A. li u a us om Ce ado and Caa ingas
biomes a e summa ized in Table 21. Fou ex e nal
cha ac e s exhibi s a is ically signi ican seconda y
sexual a ia ion, whe eas none o he c anial cha -
ac e s exhibi s a is ical di e ences be ween males
and emales. Ob ious ends in mean alues o
males and emales a e no obse ed; ends in mean
alues o Caa ingas and Ce ado popula ions a e
also no e iden . Howe e , ag oup o i e c anial
cha ac e s exhibi s s a is ically signi ican seconda y
sexual a ia ion in which ou male mean alues
a e la ge han he compa able emale mean alues.
Only asingle ex e nal cha ac e shows s a is ically
signi ican sexual a ia ion. Seconda y sexual a i-
a ion occu s in popula ions o A. li u a us bu is no
ex ensi e.
Sub amily Desmodon inae
Desmodus o undus (E. Geo oy, 1810)
O he wo ex an subspecies, D. . o undus is he
designa ion p ope ly applied o No heas B azilian
popula ions o his sanguini o e (Jones and Ca e ,
1976). I was common in he Caa ingas whe e i was
locally abundan on se o es. O he 365 cap u ed
adul specimens om he Caa ingas, 45% we e males;
as such he sexes occu ed in s a is ically indis in-
guishable p opo ions (Binomial Tes , P>.05). I
was a e in Ce ado and Ce adao habi a s o he
Chapada do A a ipe, pe haps due o he pauci y o
li es ock and o he la ge wild mammals.
86 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 20. —Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean. SE;
coe icien o a ia ion. CV; sample size, n) o A ibeus jamaicensis males and emales om bo h Caa inga and Ce ado biomes. A wo-
way analysis o a iance (Model I) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual o
geog aphic a ia ion is indica ed by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
<5<5 92 <5(5 99 Fac o d MS FSigni icance
To al leng h
Mean 74.85 75.85 73.20 73.75 A ea 170.313 7.36 .008
SD 2.98 3.08 3.12 3.18 Sex 112.013 1.26 .266
SE .67 .69 .70 .71 AxS11.013 .11 .746
CV 3.98 4.06 4.26 4.31 E o 76 9.553
n20 20 20 20
Hind oo leng h
Mean 13.25 13.65 13.30 13.35 A ea 1.313 .43 .513
SD .79 1.09 .66 .81 Sex 11.013 1.40 .241
SE .18 .24 .15 .18 AxS1.613 .85 .361
CV 5.96 7.99 4.96 6.07 E o 76 .724
n20 20 20 20
Ea leng h
Mean 20.55 20.55 20.40 20.35 A ea 1.613 .65 .421
SD .95 .95 .88 1.09 Sex 1.013 .01 .908
SE .21 .21 .20 .24 AxS1.013 .01 .908
CV 4.62 4.62 4.31 5.36 E o 76 .938
n20 20 20 20
T agus leng h
Mean 8.15 8.20 8.10 8.10 A ea 1.113 .34 .563
SD .59 .62 .55 .55 Sex 1.013 .04 .847
SE .13 .14 .12 .12 AXS1.013 .04 .847
CV 7.24 7.56 6.70 6.79 E o 76 .334
n20 20 20 20
Fo ea m leng h
Mean 58.95 59.45 57.40 57.20 A ea 172.200 16.43 <.001
SD 2.16 1.47 2.14 2.48 Sex 1.450 .10 .750
SE .48 .33 .48 .56 AXS12.450 .56 .458
CV 3.66 2.47 3.73 4.34 E o 76 4.393
n20 20 20 20
Weigh
Mean 40.98 46.38 39.40 41.03 A ea 1239.778 14.13 <.001
SD 3.48 5.84 2.08 4.17 Sex 1246.753 14.54 <.001
SE .78 1.31 .46 .93 AXS171.253 4.20 .044
CV 8.49 12.59 5.28 10.16 E o 76 16.970
n20 20 20 20
Leng h o digi one
Mean 11.65 11.70 11.75 11.45 A ea 1.113 .19 .666
SD '.75 .87 .72 .76 Sex 1.313 .52 .472
SE .17 .19 .16 .17 AXS1.613 1.02 .315
CV 6.44 7.44 6.13 6.64 E o 76 .598
n20 20 20 20
Leng h o digi h ee
Mean 113.30 114.45 1 1 1.65 111.30 A ea 1115.200 5.89 .018
SD 4.23 4.81 3.58 4.94 Sex 13.200 .16 .687
SE .95 1.08 .80 1.11 AXS111.250 .58 .451
CV 3.73 4.20 3.21 4.44 E o 76 19.551
n20 20 20 20
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 87
Table 20. —Con inued.
Caa inga Ce ado Analysis o a iance
66 92 66 99 Fac o d MS FSigni icance
Leng h o digi ou
Mean 86.45 87.85 84.95 85.25 A ea 118.050 7.81 .007
SD 2.74 3.27 2.52 4.30 Sex 114.450 1.34 .250
SE .61 .73 .56 .96 AxS16.050 .56 .456
CV 3.17 3.72 2.97 5.04 E o 76 10.766
n20 20 20 20
Leng h o digi i e
Mean 79.80 81.45 77.70 77.65 A ea 1174.050 22.77 <.001
SD 2.44 1.64 2.03 4.22 Sex 112.800 1.67 .200
SE .55 .37 .45 .94 AXS114.450 1.89 .173
CV 3.06 2.01 2.61 5.43 E o 76 7.643
n20 20 20 20
Tibia leng h
Mean 23.60 23.40 23.05 22.80 A ea 16.613 3.93 .051
SD 1.50 .60 1.23 1.61 Sex 11.013 .60 .440
SE .34 .13 .28 .36 AXS1.013 .01 .932
CV 6.36 2.56 5.34 7.06 E o 76 1.681
n20 20 20 20
Calca leng h
Mean 6.45 6.50 6.00 6.25 A ea 12.450 2.92 .091
SD .83 1.05 .92 .85 Sex 1.450 .54 .466
SE .19 .24 .21 .19 AXS1.200 .24 .627
CV 12.87 16.15 15.33 13.60 E o 76 .838
n20 20 20 20
Noselea leng h
Mean 6.80 6.45 6.50 6.45 A ea 1.450 1.22 .273
SD .62 .51 .51 .76 Sex 1.800 2.16 .145
SE .14 .11 .12 .17 AxS1.450 1.22 .273
CV 9.12 7.91 7.85 11.78 E o 76 .370
n20 20 20 20
G ea es leng h o skull
Mean 27.68 27.61 27.30 26.96 A ea 15.305 17.47 .001
SD .53 .43 .51 .70 Sex 1.841 2.77 .100
SE .12 .10 .11 .16 AxS1.338 1.11 .295
CV 1.91 1.56 1.87 2.60 E o 76 .304
n20 20 20 20
Condylobasal leng h
Mean 24.41 24.37 24.17 23.91 A ea 12.485 9.49 .003
SD .59 .38 .41 .62 Sex 1.465 1.78 .187
SE .13 .08 .09 .14 AXS1.253 .97 .329
CV 2.42 1.56 1.70 2.59 E o 76 .262
n20 20 20 20
Zygoma ic b ead h
Mean 17.20 17.07 16.87 16.62 A ea 13.121 17.05 <.001
SD .38 .30 .42 .58 Sex 1.761 4.16 .045
SE .08 .07 .09 .13 AXS1.061 .33 .567
CV 2.21 1.76 2.49 3.49 E o 76 .183
n20 20 20 20
Pos o bi al cons ic ion
Mean 6.99 6.92 6.99 6.82 A ea 1.050 1.03 .312
SD .24 .15 .28 .20 Sex 1.265 5.47 .022
SE .05 .03 .06 .04 AxS1.050 1.03 .312
CV 2.43 2.17 4.01 2.93 E o 76 .048
n20 20 20 20
94 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 22. —Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 29 88 22 Fac o d MS FSigni icance
Leng h o digi ou
Mean 75.80 80.50 76.40 81.71 A ea 17.433 .85 .361
SD 2.63 3.00 2.70 3.82 Sex 1226.452 25.95 <.001
SE .59 .67 1.21 1.44 AXS1.852 .10 .756
CV 3.47 3.73 3.53 4.68 E o 48 8.726
n20 20 57
Leng h o digi i e
Mean 73.65 78.85 75.20 79.29 A ea 18.904 .95 .334
SD 2.16 2.91 3.42 5.06 Sex 1194.701 20.80 <.001
SE .48 .65 1.53 1.91 AXS12.804 .30 .567
CV 2.93 3.69 4.55 6.38 E o 48 9.361
n20 20 57
Tibia leng h
Mean 27.90 29.35 29.00 30.71 A ea 113.713 8.92 .004
SD 1.29 1.09 1.00 1.60 Sex 122.609 14.71 <.001
SE .29 .24 .45 .61 AXS1.158 .10 .750
CV 4.62 3.71 3.45 5.21 E o 48 1.537
n20 20 57
G ea es leng h o skull
Mean 23.89 24.47 23.80 24.50 A ea 1.005 .02 .901
SD .49 .65 .89 .28 Sex 13.51 110.31 .002
SE .11 .15 .40 .11 AXS1.031 .09 .765
CV 2.05 2.66 3.74 1.14 E o 47 .341
n20 20 56
Condylobasal leng h
Mean 21.50 21.99 21.66 22.20 A ea 1.293 1.81 .185
SD .43 .42 .37 .23 Sex 12.273 14.04 <.001
SE .10 .09 .16 .09 AXS1.005 .03 .856
CV 2.00 1.91 1.71 1.04 E o 47 .162
n20 20 56
Zygoma ic b ead h
Mean 12.36 12.63 12.62 12.65 A ea 1.174 2.04 .160
SD .28 .31 .19 .35 Sex 1.187 2.19 .146
SE .06 .07 .09 .14 AXS1.118 1.39 .245
CV 2.27 2.45 1.51 2.77 E o 47 .085
n20 20 56
Pos o bi al cons ic ion
Mean 5.37 5.41 5.34 5.57 A ea 1.037 .72 .401
SD .29 .28 .15 .25 Sex 1.158 3.06 .087
SE .06 .06 .07 .10 AXS1.071 1.37 .248
CV 5.40 5.18 2.81 4.49 E o 47 .052
n20 20 56
Mas oid b ead h
Mean 12.73 12.84 12.76 12.90 A ea 1.017 .24 .627
SD .29 .23 .15 .25 Sex 1.134 1.85 .180
SE .06 .06 .07 .10 AXS1.002 .03 .871
CV 2.28 2.18 1.18 1.94 E o 47 .072
n20 20 56
B ead h o b aincase
Mean 12.44 12.49 12.60 12.55 A ea 1.109 2.15 .149
SD .20 .27 .14 .12 Sex 1.000 .00 .974
SE .05 .06 .06 .05 AXS1.019 .38 .539
CV 1.61 2.16 1.11 .96 E o 47 .050
n20 20 56
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 95
Table 22. —Con inued.
Caa inga Ce ado Analysis o a iance
<3(5 2$ <5.3 99 Fac o d MS FSigni icance
Ros al b ead h
Mean 7.83 7.97 7.62 7.73 A ea 1.418 4.65 .036
SD .25 .34 .35 .29 Sex 1.132 1.47 .232
SE .06 .08 .16 .12 AXS1.001 .01 .916
CV 3.19 4.27 4.59 3.75 E o 47 .090
n20 20 56
Heigh o b aincase
Mean 14.43 14.45 14.54 14.68 A ea 1.268 2.16 .149
SD .37 .31 .11 .51 Sex 1.057 .46 .501
SE .08 .07 .05 .21 AXS1.033 .26 .611
CV 2.56 2.14 .76 3.47 E o 47 .124
n20 20 56
B ead h ac oss he uppe mola s
Mean 6.74 6.93 6.76 6.67 A ea 1.122 1.47 .232
SD .26 .31 .17 .37 Sex 1.022 .27 .608
SE .06 .07 .08 .15 AxS1.178 2.15 .149
CV 3.86 4.47 2.51 5.55 E o 47 .083
n20 20 56
B ead h ac oss he uppe canines
Mean 6.13 6.36 6.22 6.20 A ea 1.008 .21 .646
SD .14 .20 .11 .32 Sex 1.095 2.62 .112
SE .03 .05 .05 .13 AxS1.134 3.71 .060
CV 2.28 3.14 1.77 5.16 E o 47 .036
n20 20 56
Leng h o he maxilla y oo h ow
Mean 3.42 3.32 3.42 3.43 A ea 1.030 1.81 .185
SD .09 .18 .08 .05 Sex 1.014 .86 .358
SE .02 .04 .04 .02 AXS1.025 1.52 .224
CV 2.63 5.42 2.34 1.46 E o 47 .017
n20 20 56
Leng h o he uppe mola i o m oo h ow
Mean 1.33 1.31 1.36 1.40 A ea 1.033 3.37 .073
SD .09 .11 .09 .11 Sex 1.001 .13 .715
SE .02 .02 .04 .05 AXS1.007 .65 .424
CV 6.77 8.40 6.62 7.86 E o 47 .010
n20 20 56
Wid h o he wides mola
Mean .99 .92 .92 .98 A ea 1.000 .01 .926
SD .11 .09 .11 .12 Sex 1.000 .01 .926
SE .03 .02 .05 .05 AxS1.038 3.54 .066
CV 11.11 9.78 11.96 12.24 E o 47 .01
1
n20 20 56
G ea es leng h o he mandible
Mean 15.06 15.49 15.50 15.67 A ea 1.802 5.56 .023
SD .40 .38 .32 .35 Sex 1.750 5.20 .027
SE .09 .08 .15 .14 AXS1.143 .99 .324
CV 2.66 2.45 2.06 2.23 E o 47 .144
n20 20 56
Leng h o he mandibula oo h ow
Mean 4.53 4.55 4.52 4.60 A ea 1.004 .12 .734
SD .13 .22 .30 .13 Sex 1.019 .52 .473
SE .03 .05 .14 .05 AXS1.009 .24 .623
CV 2.87 4.84 6.64 2.83 E o 47 .037
n20 20 56
96 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 22. —Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 22 66 22 Fac o d MS FSigni icance
Leng h o he co onoid p ocess
Mean 5.72 6.15 5.80 6.00 A ea i.01
1
.08 .777
SD .34 .41 .27 .28 Sex i.851 6.59 .014
SE .08 .09 .12 .12 AXSi.113 .88 .353
CV 5.94 6.67 4.66 4.67 E o 47 .129
n20 20 56
g aphic a ia ion in popula ions o D. o undus om
he No heas o B azil. In all bu one case, emale
mean alues a e la ge han male mean alues and
s a is ically signi ican seconda y sexual a ia ion is
e ealed o se en ex e nal and ou c anial cha -
ac e s. The obse a ions o sexual dimo phism by
Husson (1962) and He shko i z (1949) a e s a is-
ically co obo a ed. Ce ado sample means a e
la ge han Caa ingas sample means o 19 cha ac-
e s, h ee o which exhibi s a is ically signi ican
a ia ion. Caa ingas specimens ha e la ge sample
means han Ce ado specimens o only one cha -
ac e and he di e ence is no s a is ically signi ican .
The ailu e o achie e s a is ical signi icance o geo-
g aphic a ia ion is p obably due o he small sam-
ple sizes om he Ce ado. Howe e , he consis en
di ec ion o he di e ence be ween sample means
om he Caa ingas and Ce ado does sugges he
s ong possibili y o geog aphic a ia ion wi hin he
No heas o B azil.
Diphylla ecauda a Spix, 1823
Ojas i and Lina es (1971) ecognize wo subspe-
cies o his sanguini o e; D. e. ecauda a is he des-
igna ion app op ia e o specimens om he No h-
eas o B azil. This species was a e in he Caa ingas
and appa en ly absen om he Chapada do A a-
ipe.
Among he au ho s ci ed by Swanepoel and Gen-
oways(1979) o con ibu ing knowledge o D. ecau-
da a'% mo phome ic biology, only Dobson (1878),
Lima (1926), and Cunha Viei a (1942) epo ed
measu emen s om B azilian specimens. Two adul
males and wo adul emales we e collec ed in he
Caa ingas; selec ed mo phological measu emen s
(a e Swanepoel and Genoways, 1979) a e he ein
epo ed (To al leng h: 70, 66, 74, 69; ail leng h:
—,—,—,—;hind oo leng h: 15, 13, 15, 12; ea
leng h: 15, 14, 15, 14; g ea es leng h o skull: 21.2,
20.8, 21.2, 21.1; condylobasal leng h: 18.9, 18.0,
19.0, 18.7; zygoma ic b ead h: 12.2, 11.6, 12.1, 11.7;
pos o bi al cons ic ion: 6.8, 6.2, 6.5, 6.5; leng h o
he maxilla y oo h ow: 3.0, 2.9, 3.0, 2.9; and b ead h
ac oss he uppe mola s: 5.2, 4.9, 5.0, 5.0). Caa ingas
specimens a e on he a e age smalle han he g oup
o specimens om No h and Cen al Ame ica whose
measu emen s a e epo ed by Swanepoel and Gen-
oways (1979).
Family Na alidae
Na alus s amineus G ay, 1838
This ae ial insec i o e was appa en ly absen om
he Caa ingas and a e on he Chapada do A a ipe.
Because only wo adul male specimens we e cap-
u ed, only selec measu emen s (a e Swanepoel
and Genoways, 1979) a e epo ed he e ( o al leng h:
99, 104; ail leng h: 53, 57; hind oo leng h: 9, 8;
ea leng h: 15, 12; g ea es leng h o skull: —,16.0;
condylobasal leng h: —,14.9; zygoma ic b ead h:
—,8.7; pos o bi al cons ic ion: —,3.2; b ead h o
b aincase: —,—;leng h o he maxilla y oo h ow:
—,6.7; b ead h ac oss he uppe mola s: —,4.0).
Specimens om he No heas should be included
in he subspecies N. s. na alensis because hey we e
cap u ed wi hin he de ined ange o ha subspecies
(Goodwin, 1959) and because mo phome ic com-
pa ison wi h he ype specimen indica es co e-
spondence.
Family Fu ip e idae
Fu ip e us ho ens (F. Cu ie , 1828)
This poo ly known ae ial insec i o e was a e in
he Caa ingas whe e i was only cap u ed on se o es.
I was appa en ly absen om all habi a s on he
Chapada do A a ipe. Only one adul male and one
adul emale specimen we e cap u ed, hence selec ed
measu emen s (a e Swanepoel and Genoways,
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 97
1979) only will be epo ed he e ( o al leng h: 84,
80; ail leng h: 45, 44; hind oo leng h: 6, 6; ea
leng h: 10, 9; g ea es leng h o skull: 12.0, 12.0;
condylobasal leng h: 11.0, 11.0; zygoma ic b ead h:
7.2, 7.2; pos o bi al cons ic ion: 2.9, 2.8; b ead h
o b aincase: 5.6, 5.8; leng h o he maxilla y oo h-
ow: 4.3, 4.6; b ead h ac oss he uppe mola s: 4.5,
4.6). Uieda e al. (1980) summa ize and p esen new
da a on aspec s o he biology o F. ho ens] un o -
una ely, li le can be added om his s udy.
Family Vespe ilionidae
Myo is nig icans (Schinz, 1821)
This ae ial insec i o e was a e o common in
bo h Caa ingas and Ce ado biomes. O he h ee
subspecies cu en ly ecognized, popula ions om
he No heas should en a i ely be assigned o he
subspecies M. n. nig icans (LaVal, 1973); howe e ,
u he sys ema ic analyses a e equi ed o subs an-
ia e mo phome ic homogenei y wi hin he sub-
species. The sexes occu ed in s a is ically indis in-
guishable p opo ions in bo h Caa ingas and Ce ado
biomes (Binomial Tes P>.05). Only 39% o he
41 cap u ed adul specimens om he Chapada do
A a ipe we e males while 44% o he 260 cap u ed
specimens om he Ce ado we e males. M. n. ni-
g icans we e equen ly ound oos ing unde he
oo ing iles o abandoned buildings h oughou he
No heas ; howe e , indi iduals we e no ound in
agg ega ions o any size.
Mye s (1978) de ec ed s a is ical di e ences in
o ea m leng h be ween specimens o male and e-
male M. nig icans om Pa aguay. Williams and
Findley (1979) ound male sample means consis-
en ly smalle han emale sample means o six
mensu al cha ac e s; howe e , only head and body
leng h exhibi ed s a is ically signi ican a ia ion.
Table 23 summa izes he esul s o s a is ical anal-
yses o indi idual, seconda y sexual and geog aphic
a ia ion in popula ions o M. nig icans om he
No heas o B azil. S a is ically signi ican second-
a y sexual a ia ion occu s only o asingle a iable,
weigh ; u he , apa e n in he ela ionship be ween
male and emale mean alues is no appa en . M.
nig icans popula ions om he No heas o B azil
clea ly do no exhibi sexual dimo phism in ex e nal
and c anial cha ac e s.
Popula ions o M. nig icans om Caa ingas and
Ce ado biomes do no exhibi app eciable geo-
g aphic a ia ion ei he . Only wo cha ac e s, calca
leng h and leng h om he canine o he las mola
in he mandible, exhibi s a is ically signi ican a i-
a ion be ween Caa ingas and Ce ado si es; and
ends in mean alue ela ionships a e no appa en .
The Caa ingas and Ce ado popula ions a e mo -
phome ically homogeneous.
Ep esicus u inalis D’O bigny and Ge ais, 1847
This ae ial insec i o e was appa en ly absen om
he Caa ingas and a e on he Chapada do A a ipe.
Williams (1978) sugges ed ha Caa ingas specimens
o E. u inalis a e mos closely allied wi h he sub-
species E. chapmani;howe e , he specimens he
u ilized we e ac ually asubsample o he specimens
caugh in Ce ado habi a s o he Chapada do A a-
ipe.
Samples a e small bu su icien o analyzing in-
di idual and seconda y sexual a ia ion by s a is-
ical me hods; he esul s a e summa ized in Table
24. Females ha e la ge sample means han males
o en ex e nal and 14 c anial cha ac e s. Males
ha e la ge sample means han emales o i e a i-
ables. Th ee o he cha ac e s ( o ea m leng h, weigh ,
and g ea es leng h o he maxilla y) exhibi ed s a-
is ically signi ican seconda y sexual a ia ion; in
all h ee cases, he male mean alue was la ge han
he emale mean alue. Fu he , Williams (1978)
sugges ed ha sexual dimo phism exis ed in he
Chapada popula ion o E. u inalis based upon a
mul i a ia e analysis o a ia ion.
Lasiu us bo ealis (Mulle , 1776)
This ae ial insec i o e was a e on he Chapada
do A a ipe and appa en ly absen om he Caa in-
gas. I s ange ex ends om no he n No h Ame ica
o sou he n Sou h Ame ica.
Samples o L. bo ealis a e small, bu o su icien
size o a emp s a is ical analyses o indi idual and
seconda y sexual a ia ion (Table 25). Females ha e
la ge sample means han males o all cha ac e s
excep pos o bi al cons ic ion. S a is ically signi i-
can seconda y sexual a ia ion was e idenced by
ou ex e nal and six c anial cha ac e s. Sexual di-
mo phism o size wi h emales la ge han males
is s ongly indica ed by he da a. This co obo a es
he indings o Williams and Findley (1979) which
also indica ed sexual dimo phism.
Lasiu us ega (Ge ais, 1856)
This ae ial insec i o e was p esen bu a e in
bo h Caa ingas and Ce ado biomes. Acco ding o
he subspecies anges sugges ed by Handley (1960),
specimens om he No heas o B azil should be
98 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 23.— Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD; s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o/'Myo is nig icans males and emales om bo h Caa inga and Ce ado biomes. A wo-way
analysis o a iance (Model I) wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual o geog aphic
a ia ion is indica ed by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
66 29 66 22 Fac o d MS FSigni icance
To al leng h
Mean 80.65 81.90 81.44 81.75 A ea 11.912 .19 .661
SD 2.61 3.95 2.61 3.09 Sex 111.489 1.17 .283
SE .58 .88 .65 .69 AXS14.136 .42 .519
CV 3.24 4.82 3.20 3.78 E o 72 9.834
n20 20 16 20
Tail leng h
Mean 35.95 35.45 35.44 35.30 A ea 12.065 .22 .641
SD 2.50 2.74 2.76 3.98 Sex 11.913 .20 .653
SE .56 .61 .69 .89 AXS1.618 .07 .798
CV 6.95 7.73 7.79 11.27 E o 72 9.389
n20 20 16 20
Hind oo leng h
Mean 5.95 6.00 5.94 5.90 A ea 1.060 .56 .458
SD .39 0.25 .45 Sex 1.001 .01 .934
SE .09 0.06 .10 AXS1.036 .34 .563
CV 6.55 04.21 7.63 E o 72 .107
n20 20 16 20
Ea leng h
Mean 12.35 12.20 12.31 12.25 A ea 1.001 .00 .973
SD .75 .77 .79 .85 Sex 1.213 .34 .561
SE .17 .17 .20 .19 AXS1.036 .06 .811
CV 6.07 6.31 6.42 6.94 E o 72 .624
n20 20 16 20
T agus leng h
Mean 7.65 7.55 7.81 7.80 A ea 1.801 2.29 .134
SD .59 .51 .54 .70 Sex 1.060 .17 .681
SE .13 .11 .14 .16 AXS1.036 .10 .749
CV 7.71 6.75 6.91 8.97 E o 72 .349
n20 20 16 20
Fo ea m leng h
Mean 34.40 33.85 34.00 33.80 A ea 1.953 .77 .382
SD 1.14 .88 1.32 1.11 Sex 12.647 2.15 .147
SE .26 .20 .33 .25 AXS1.577 .47 .456
CV 3.31 2.60 3.88 3.28 E o 72 1.230
n20 20 16 20
Weigh
Mean 4.50 4.95 4.34 4.88 A ea 1.252 .85 .359
SD .46 .63 .44 .60 Sex 14.531 15.35 <.001
SE .10 .14 .11 .14 AxS1.031 .11 .747
CV 10.22 12.73 10.14 12.30 E o 72 .295
n20 20 16 20
Leng h o digi one
Mean 4.95 4.70 4.63 4.70 A ea 1.497 1.70 .197
SD .51 .47 .62 .57 Sex 1.144 .49 .485
SE .11 .11 .16 .13 AxS1.497 1.70 .197
CV 10.30 10.00 13.39 12.13 E o 72 .293
n20 20 16 20
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 99
Table 23. —Con inued.
Caa inga Ce ado Analysis o a iance
<36 99 <33 99 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 56.50 56.65 56.06 56.60 A ea 11.118 .16 .694
SD 2.91 2.23 2.59 2.91 Sex 12.224 .31 .580
SE .65 .50 .65 .65 AxS1.707 .10 .755
c 5.15 3.94 4.62 5.14 E o 72 7.185
n20 20 16 20
Leng h o digi ou
Mean 47.45 47.10 45.88 47.30 A ea 18.897 1.51 .223
SD 2.37 2.17 2.47 2.68 Sex 15.438 .92 .340
SE .53 .49 .62 .60 AxS114.827 2.51 .117
CV 4.99 4.61 5.38 5.67 E o 72 5.899
n20 20 16 20
Leng h o digi i e
Mean 44.45 44.15 43.88 43.85 A ea 13.603 .78 .381
SD 1.54 2.01 2.16 2.74 Sex 1.497 .11 .744
SE .34 .45 .54 .6
1
AXS1.356 .08 .783
CV 3.46 4.55 4.92 6.25 E o 72 .636
n20 20 16 20
Tibia leng h
Mean 15.45 15.00 15.19 15.20 A ea 1.018 .03 .869
SD .95 .80 .75 .77 Sex 1.901 1.33 .252
SE .21 .18 .19 .17 AxS11.007 1.49 .226
CV 6.15 5.33 4.94 5.07 E o 72 .675
n20 20 16 20
Calca leng h
Mean 12.60 12.50 12.44 11.50 A ea 16.360 3.99 .050
SD 1.27 1.47 1.26 1.00 Sex 15.065 3.18 .079
SE .29 .33 .32 .22 AXS13.301 2.07 .154
CV 10.08 11.76 10.13 8.70 E o 72 1.594
n20 20 1
6
20
G ea es leng h o skull
Mean 13.46 13.50 13.49 13.51 A ea 1.006 .06 .803
SD .35 .29 .24 .30 Sex 1.013 .14 .705
SE .08 .07 .06 .07 AXS1.001 .01 .915
CV 2.60 2.15 1.78 2.22 E o 69 .092
n20 20 14 19
Condylobasal leng h
Mean 12.81 12.75 12.79 12.71 A ea 1.010 .11 .736
SD .34 .24 .28 .29 Sex 1.090 1.07 .305
SE .08 .06 .08 .07 AxS1.002 .03 .872
CV 2.65 1.88 2.19 2.28 E o 69 .085
n20 20 14 19
Zygoma ic b ead h
Mean 8.26 8.30 8.36 8.32 A ea 1.054 1.67 .202
SD .17 .16 .24 .18 Sex 1.000 .00 .952
SE .04 .04 .08 .05 AXS1.024 .73 .397
CV 2.06 1.93 2.87 2.16 E o 55 .032
n20 19 812
Pos o bi al cons ic ion
Mean 3.29 3.25 3.29 3.25 A ea 1.000 .02 .878
SD .09 .14 .12 .11 Sex 1.028 2.1
1
.151
SE .02 .03 .03 .02 AXS1.000 .02 .878
CV 2.74 4.31 3.65 3.38 E o 71 .013
n20 20 15 20
100 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 23.— Con inued.
Caa inga Ce ado Analysis o a iance
<53 2$ <5<5 22 Fac o d MS FSigni icance
Mas oid b ead h
Mean 6.94 6.93 6.93 6.92 A ea 1.002 .07 .794
SD .12 .15 .10 .18 Sex 1.002 .08 .777
SE .03 .03 .03 .04 AxS1.000 .02 .887
CV 1.73 2.16 1.44 2.60 E o 68 .022
n20 20 13 19
B ead h o b aincase
Mean 6.48 6.40 6.38 6.42 A ea 1.029 1.09 .300
SD .21 .13 .11 .17 Sex 1.006 .24 .628
SE .05 .03 .03 .04 AXS1.067 2.53 .116
CV 3.24 2.03 1.72 2.65 E o 69 .027
n20 20 14 19
Ros al b ead h
Mean 3.50 3.49 3.42 3.50 A ea 1.020 1.00 .322
SD .11 .13 .13 .18 Sex 1.020 1.00 .322
SE .02 .03 .03 .04 AXS1.042 2.13 .149
CV 3.14 3.72 3.80 5.14 E o 71 .020
n20 20 15 20
Heigh o b aincase
Mean 6.67 6.70 6.59 6.58 A ea 1.168 3.61 .062
SD .20 .20 .17 .26 Sex 1.003 .06 .812
SE .05 .05 .05 .06 AXS1.003 .06 .806
CV 3.00 2.99 2.58 3.95 E o 68 .046
n20 20 13 19
B ead h ac oss he uppe mola s
Mean 5.41 5.41 5.35 5.34 A ea 1.076 2.89 .094
SD .18 .15 .17 .15 Sex 1.001 .03 .854
SE .04 .03 .05 .03 AXS1.000 .00 .956
CV 3.33 2.77 3.18 2.81 E o 68 .026
n20 20 13 19
B ead h ac oss he uppe canines
Mean 3.42 3.48 3.46 3.48 A ea 1.008 .41 .526
SD .15 .15 .10 .13 Sex 1.024 1.30 .259
SE .03 .03 .03 .03 AXS1.005 .27 .606
CV 4.39 4.31 2.89 3.74 E o 67 .019
n20 20 12 19
Leng h o he maxilla y oo h ow
Mean 5.09 5.07 5.11 5.07 A ea 1.005 .28 .599
SD .19 .11 .14 .10 Sex 1.019 .98 .325
SE .04 .02 .04 .02 AXS1.003 .14 .709
CV 3.73 2.17 2.74 1.97 E o 70 .019
n20 20 14 20
Leng h o he uppe mola i o m oo h ow
Mean 4.22 4.21 4.16 4.22 A ea 1.010 .43 .512
SD .20 .1 1.18 .12 Sex 1.015 .63 .429
SE .04 .02 .05 .03 AXS1.021 .87 .354
CV 4.74 2.61 4.33 2.84 E o 70 .024
n20 20 14 20
Wid h o he wides mola
Mean 1.58 1.56 1.58 1.57 A ea 1.001 .12 .726
SD .06 .08 .11 .08 Sex 1.003 .37 .543
SE .01 .02 .03 .02 AXS1.000 .03 .868
CV 3.80 5.13 6.96 5.10 E o 70 .007
n20 20 14 20
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 101
Table 23 .—Con inued.
Caa inga Ce ado Analysis o a iance
33 99 66 99 Fac o d MS FSigni icance
G ea es leng h o he mandible
Mean 9.80 9.82 9.84 9.86 A ea 1.033 .44 .510
SD .30 .26 .27 .27 Sex 1.007 .10 .757
SE .07 .06 .07 .06 AXSI.000 .00 .998
CV 3.06 2.65 2.74 2.74 E o 68 .075
n20 20 14 19
Leng h o he mandibula oo h ow
Mean 5.55 5.55 5.43 5.43 A ea 1.279 7.75 .007
SD .21 .15 .17 .22 Sex 1.000 .00 .961
SE .05 .03 .05 .05 AXS1.000 .00 .961
CV 3.78 2.70 3.13 4.05 E o 68 .036
n20 20 13 19
Leng h o he co ono d p ocess
Mean 3.00 3.02 2.91 3.01 A ea 1.045 3.03 .086
SD .14 .10 .10 .13 Sex 1.055 3.73 .058
SE .03 .02 .03 .03 AxS1.030 2.01 .161
CV 4.67 3.31 3.44 4.32 E o 67 .015
n20 20 13 18
Table 24. —Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion, SD: s anda d e o o he
coe icien o a ia ion, CV; sample size, n) o/Ep esicus u inalis males and emales om he Ce ado biome. Aone-way
a iance (Mode! I)wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual a ia ion
by P alues less han o equal o .050.
mean, SE;
analysis o
is indica ed
Caa inga Ce ado Analysis o a iance
<53 99 33 99 Fac o d MS FSigni icance
To al leng h
Mean 96.71 97.40 Sex 12.244 .18 .678
SD 3.25 3.68 Wi hin 20 12.651
SE 1.23 .95
CV 3.36 3.78
n715
Tail leng h
Mean 39.14 38.33 Sex 13.127 .38 .544
SD 1.35 3.31 Wi hin 20 8.210
SE .51 .85
CV 3.45 8.64
n715
Hind oo leng h
Mean 7.00 6.33 Sex 12.121 2.77 .112
SD .82 .90 Wi hin 20 .767
SE .31 .23
CV 11.71 14.22
n715
Ea leng h
Mean 13.57 13.67 Sex 1.043 .07 .799
SD .98 .72 Wi hin 20 .652
SE .37 .19
CV 7.22 5.27
n715
102 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 24.— Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 $2 66 22 Fac o d MS FSigni icance
T agus leng h
Mean 8.14 8.53 Sex 1.728 1.00 .330
SD .90 .83 Wi hin 20 .730
SE .34 .22
CV 11.06 8.73
n715
Fo ea m leng h
Mean 38.86 40.13 Sex 17.773 5.44 .030
SD 1.35 1.13 Wi hin 20 1.430
SE .51 .29
CV 3.47 2.82
n715
Weigh
Mean 8.36 9.53 Sex 16.603 5.37 .031
SD .69 1.25 Wi hin 20 1.230
SE .26 .32
CV 8.25 13.12
n7 15
Leng h o digi one
Mean 4.86 5.40 Sex 11.407 3.33 .083
SD .69 .63 Wi hin 20 .423
SE .26 .16
CV 14.20 11.67
n715
Leng h o digi h ee
Mean 67.14 68.80 Sex 113.107 2.59 .123
SD 2.34 2.21 Wi hin 20 5.063
SE .88 .57
CV 3.48 3.21
n715
Leng h o digi ou
Mean 55.86 57.07 Sex 16.982 2.50 .129
SD 1.68 1.67 Wi hin 20 2.790
SE .63 .43
CV 3.01 2.93
n7 15
Leng h o digi i e
Mean 49.29 49.87 Sex 11.61
1
.75 .398
SD 1.38 1.51 Wi hin 20 2.158
SE .52 .39
CV 2.80 3.03
n7 15
Tibia leng h
Mean 16.00 16.27 Sex 1.339 .30 .592
SD 01.28 Wi hin 20 1.147
SE 0.33
CV 07.87
n715
Calca leng h
Mean 12.57 12.87 Sex 1.416 .28 .601
SD 1.27 1.19 Wi hin 20 1.472
SE .48 .30
CV 10.10 9.25
n715
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 103
Table 24.— Con inued.
Caa inga Ce ado Analysis o a iance
<5(5 92 (5.5 22 Fac o d MS FSigni icance
G ea es leng h o skull
Mean 14.88 14.84 Sex 1.010 ,ii .746
SD .33 .28 Wi hin 18 .088
SE .13 .08
CV 2.22 1.89
n6 14
Cond lobasal leng h
Mean 14.38 14.39 Sex 1.000 .00 .993
SD .28 .30 Wi hin 17 .086
SE .11 .08
CV 1.95 2.08
n6 13
Zygoma ic b ead h
Mean 10.52 10.56 Sex 1.005 .08 .787
SD .20 .30 Wi hin 13 .072
SE .08 .10
CV 1.90 2.84
n69
Pos o bi al cons ic ion
Mean 3.59 3.64 Sex 1.014 .56 .464
SD .17 .1
6
Wi hin 20 .025
SE .06 .04
CV 4.74 4.40
n715
Mas oid b ead h
Mean 8.25 8.35 Sex 1.040 1.01 .327
SD .15 .21 Wi hin 19 .040
SE .06 .06
CV 1.82 2.5!
n615
B ead h o b aincase
Mean 7.12 7.25 Sex 1.080 3.57 .074
SD .15 .15 Wi hin 19 .022
SE .06 .04
CV 2.1
1
2.07
n615
Ros al b ead h
Mean 4.69 4.83 Sex 1.104 2.31 .145
SD .25 .19 Wi hin 20 .045
SE .10 .05
CV 5.33 3.93
n715
Heigh o b aincase
Mean 7.42 7.47 Sex 1.013 .18 .674
SD .3! .24 Wi hin 18 .069
SE .13 .07
CV 4.18 3.21
n614
B ead h ac oss he uppe mola s
Mean 6.46 6.51 Sex 1.015 .46 .505
SD .10 .21 Wi hin 20 .033
SE .04 .05
CV 1.55 3.23
n715
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
110
Table 26. —Con inued.
Caa inga Ce ado Analysis o a iance
<5<5 $2 36 22 Fac o d MS FSigni icance
G ea es leng h o skull
Mean 15.00 15.13 14.90 15.50 Sex 1.114 1.26 .313
SD .57 .11 ——Wi hin 5.091
SE .40 .07 ——
CV 3.80 .73 ——
n2311
Condylobasal leng h
Mean 14.50 14.80 14.50 15.00 Sex 1.210 4.57 .086
SD .42 .10 ——Wi hin 5.046
SE .30 .06 — —
CV 2.90 .68 ——
n231 1
Zygoma ic b ead h
Mean 10.55 10.83 10.60 10.80 Sex 1.114 1.62 .260
SD .07 .42 — — Wi hin 5.071
SE .05 .24 ——
CV .66 3.88 ——
n2311
Pos o bi al cons ic ion
Mean 4.75 4.58 4.60 4.30 Sex 1.061 .47 .517
SD .21 .47 — — Wi hin 6.128
SE .15 .23 ——
CV 4.42 10.26 ——
n2411
Mas oid b ead h
Mean 8.75 8.90 8.70 8.60 Sex 1.014 .41 .549
SD .21 .17 ——Wi hin 5.035
SE .15 .10 ——
CV 2.42 1.91 ——
n2411
B ead h o b aincase
Mean 8.20 8.23 8.40 8.10 Sex 1.008 .09 .772
SD .42 .31 ——Wi hin 5.081
SE .30 .18 ——
CV 5.12 3.77 ——
n2411
Ros al b ead h
Mean 5.85 5.30 5.90 6.00 Sex 1.263 1.14 .335
SD .07 .62 ——Wi hin 5.231
SE .05 .36 ——
CV 1.20 11.70 ——
n2311
Heigh o b aincase
Mean 8.95 8.90 9.20 8.80 Sex 1.043 1.23 .317
SD .21 .20 ——Wi hin 5.035
SE .15 .12 ——
CV 2.35 2.25 ——
n231 1
B ead h ac oss he uppe mola s
Mean 7.10 6.88 6.90 7.30 Sex 1.010 .15 .710
SD .14 .26 — — Wi hin 6.066
SE .10 .13 ——
CV 1.97 3.78 ——
n2411
1983 WILLIG- BRAZILIAN BAT COMMUNITIES
Table 26. —Con inued.
Caa inga Ce ado Analysis o a iance
33 22 S3 22 Fac o d MS FSigni icance
B ead h ac oss he uppe canines
Mean 5.90 5.83 6.10 5.90 Sex 1.023 .54 .496
SD .14 .29 ——Wi hin 5.043
SE .10 .17 ——
CV 2.37 4.97 ——
n2311
Leng h o he maxilla y oo h ow
Mean 5.35 5.37 5.30 5.50 Sex 1.008 .57 .484
SD .07 .15 ——Wi hin 5.013
SE .05 .09 ——
CV 1.31 2.79 ——
n231 1
Leng h o he uppe mola i o m oo h ow
Mean 4.15 4.27 4.20 4.50 Sex 1.043 3.97 .103
SD .07 .06 ——Wi hin 5.011
SE .05 .03 ——
CV 1.69 1.41 ——
n231 1
Wid h o he wides mola
Mean 2.00 2.08 2.00 2.30 Sex 1.027 2.38 .174
SD 0.10 ——Wi hin 6.01
1
SE 0.05 ——
CV 04.81 ——
n2411
G ea es leng h o he mandible
Mean 11.10 11.63 11.30 10.60 Sex 1.120 .77 .413
SD .14 .13 ——Wi hin 6.156
SE .10 .06 — —
CV 1.26 1.11 ——
n2411
Leng h o he mandibula oo h ow
Mean 6.15 6.20 6.20 5.80 Sex 1.008 .14 .721
SD .07 .27 ——Wi hin 5.053
SE .05 .15 ——
CV 1.13 4.35 ——
n2311
Leng h o he co onoid p ocess
Mean 4.00 3.90 4.00 4.60 Sex 1.003 .03 .865
SD 0.25 ——Wi hin 6.095
SE 0.12 ——
CV 06.4! ——
n2411
e nal cha ac e is ics, bu he ela ionship be ween
male and emale sample means was no consis en
enough o sugges he exis ence o sexual dimo -
phism in his species.
Family Molossidae
Molossops plani os is (Pe e s, 1865)
This as - lying ae ial insec i o e was a e in he
Caa ingas and appa en ly absen om he Chapada
do A a ipe. The subspeci ic s a us o popula ions
om he No heas is unce ain. The h ee cap u ed
specimens we e ob ained om ahollow in acu ee
om Caa inga Al a.
Vizo o and Taddei (1976) s a is ically analyzed
indi idual and seconda y sexual a ia ion in M.
plani os is. They showed conclusi ely ha males
a e la ge han emales on he a e age and ha s a-
is ically signi ican seconda y sexual a ia ion oc-
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
112
cu s o mos ex e nal and all c anial cha ac e s ana-
lyzed.
The h ee indi iduals cap u ed om he Caa in-
gas do no cons i u e asu icien ly la ge sample o
s a is ical analyses so only selec ed measu emen s
(a e Swanepoel and Genoways, 1979) a e p e-
sen ed he e o an adul male and wo adul emale
specimens ( o al leng h: 96, 93, 92; ail leng h: 22,
24, 26; hind oo leng h: 5, 6, 6; ea leng h: 16, 17,
14; g ea es leng h o skull: 16.1, 15.1, 15.4; con-
dylobasal leng h: 15.8, 15.0, 14.9; zygoma ic b ead h:
11.2, 10.8, 11.0; pos o bi al cons ic ion: 4.3, 4.1,
4.2; b ead h o b aincase: 7.8, 7.5, 7.8; leng h o he
maxilla y oo h ow: 6.5, 6.3, 6.0; b ead h ac oss he
uppe mola s: 7.7, 7.7, 7.7).
Molossops emminckii (Bu meis e , 1854)
This molossid ae ial insec i o e was a e in he
Caa ingas and on he Chapada do A a ipe. Based
upon colo a ion cha ac e is ics and he dis ibu-
ional in o ma ion summa ized by Vizo o and Tad-
dei (1976), specimens om no heas e n B azil
should be conside ed M. . emminckii.
Vizo o and Taddei (1976) a e he only au ho s
o analyze mo phome ic ela ions in M. emminck-
ii ia s a is ical echniques. None o he 17 ex e nal
cha ac e s hey examine exhibi s a is ically sig-
ni ican seconda y sexual a ia ion, and consis en
mean alue ela ions be ween males and emales a e
no e iden . On he o he hand, males a e consis-
en ly la ge han emales o c anial cha ac e s, bu
only wo cha ac e s exhibi s a is ically signi ican
di e ences.
In o de o pe o m s a is ical analyses o indi-
idual and seconda y sexual a ia ion o specimens
om he No heas o B azil, samples a e combined
om Caa ingas and Ce ado biomes; he esul s o
he analyses a e summa ized in Table 27. None o
he ex e nal o c anial cha ac e s exhibi s a is ically
signi ican seconda y sexual a ia ion, howe e , male
sample means a e consis en ly la ge han hose o
emales. Al hough da a om he No heas as well
as ha o Vizo o and Taddei (1976) ail o de ec
signi ican di e ences be ween males and emales,
bo h analyses s ongly sugges sexual dimo phism
wi h males la ge han emales, a leas o c anial
cha ac e s. La ge sample sizes o amul i a ia e
analysis o a iance would acili a e he de ec ion
o sexual dimo phism in his species.
Tada ida la icauda a (E. Geo oy, 1805)
This as - lying ae ial insec i o e was a e in he
Caa ingas and on he Chapada do A a ipe. Samples
om he No heas o B azil a e small, so selec ed
measu emen s (a e Swanepoel and Genoways,
1979) o an adul male and emale om he Caa-
ingas and an adul male om he Chapada do A a-
ipe a e he ein epo ed ( o al leng h: 96, 96, 107;
ail leng h: 37, 39, 36; hind oo leng h: 8, 8, 8; ea
leng h: 18, 18, 20; g ea es leng h o skull: 15.7, 16.2,
—;condylobasal leng h: 14.9, 15.2, —;zygoma ic
b ead h: 9.3, 9.3, —;pos o bi al cons ic ion: 3.3,
3.2, —;b ead h o b aincase: 8.0, 7.7, —;leng h o
he maxilla y oo h ow: 6.1, 6.2, —;and b ead h
ac oss he uppe mola s: 6.6, 6.6, —).
Neopla ymops ma og ossensis (Cunha Viei a, 1942)
This small molossid ae ial insec i o e was com-
mon in ocky habi a s and on se o es in he Caa-
ingas whe e i oos ed in low-lying ock c e ices.
App oxima ely 46% o he 48 cap u ed adul spec-
imens om he Caa ingas we e males; as such, he
sexes occu ed in s a is ically indis inguishable p o-
po ions (Binomial Tes , P>.05). N. ma og os-
sensis is absen om he Chapada do A a ipe.
The biology o his species is poo ly known. O he
han he o iginal desc ip ion (Cunha Viei a, 1942)
and he e ision o la -headed ba s by Pe e son
(1965), only isola ed dis ibu ional eco ds appea
in he li e a u e (Handley, 1976; Sazima and Tad-
dei, 1976). In amul i a ia e analysis o he amily
Molossidae, F eeman (1981) ecen ly sugges ed al-
loca ing specimens in his species o he genus Mo-
lossops;howe e aconsensus on his axonomic
ques ion is no ye appa en . Bo h Pe e son (1965)
and Sazima and Taddei (1 976) epo mensu al da a
o N. ma og ossensis bu no one has s a is ically
examined indi idual o seconda y sexual a ia ion
in his species. Willig (unpublished ms.) has de ec ed
s a is ically signi ican geog aphic a ia ion wi hin
and among popula ions o Neopla ymops om he
No heas o B azil, sou he n Venezuela, and sou h-
wes e n B i ish Guiana.
Table 28 summa izes he esul s o as a is ical
analysis o indi idual and seconda y sexual a ia-
ion in his species. Like o he molossids, Neopla-
ymops exhibi s app eciable sexual dimo phism.
Males ha e la ge sample means han emales o
12 o he 13 ex e nal cha ac e s. Males ha e la ge
sample means han emales o all 16 c anial cha -
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 113
Table 21 .—Summa y s a is ics o ex e naland c ania! cha ac e s (mean; s anda d de ia ion, SD: s anda d e o o he mean, SE;
coe icien o a ia ion, CV; sample size, n) o/ Molossops emminckii males and emales om bo h Caa inga and Ce ado biomes. Due
o small sample sizes, da a om bo h biomes we e combined o de e mine he exis ence o signi ican (P <.050) seconda y sexual
a ia ion ia aone-way analysis o a iance (Model I) wi h eplica ion.
Caa inga Ce ado Analysis o a iance
<3<3 9$ 66 99 Fac o d MS FSigni icance
To al leng h
Mean 78.00 82.67 79.25 81.33 Sex 124.55 2.83 .127
SD —3.22 2.50 4.16 Wiihin 98.67
SE —1.86 1.25 2.40
CV —3.90 3.15 5.1 1
n1343
Tail leng h
Mean 21.00 24.00 26.00 29.00 Sex 16.136 .43 .510
SD —3.46 3.74 1.72 Wi hin 914.389
SE —2.00 1.87 1.00
CV —14.42 14.38 5.97
n1343
Hind oo leng h
Mean 6.00 5.00 5.25 5.00 Sex 1.436 .55 .479
SD —1.00 .96 1.00 Wi hin 9.800
SE —.48 .48 .58
CV —20.00 18.29 20.00
n1343
Ea leng h
Mean 12.00 12.67 13.00 12.33 Sex 1.246 .51 .492
SD —.58 .82 .58 Wi hin 9.478
SE —.33 .41 .33
CV —4.58 6.31 4.70
n1343
T agus leng h
Mean 3.00 2.67 2.25 3.33 Sex 1.982 1.23 .297
SD -.58 .50 1.53 Wi hin 9.800
SE —.33 .25 .88
CV —21.72 22.22 45.94
n!343
Fo ea m leng h
Mean 33.00 31.67 31.50 31.00 Sex 1.594 .87 .375
SD -.58 .58 1.00 Wi hin 9.682
SE —.33 .29 .58
CV -1.83 1.84 3.23
n1343
Weigh
Mean 6.50 7.00 6.88 6.33 Sex 1.049 .08 .787
SD —.50 1.03 .76 Wi hin 9.626
SE —.29 .52 .44
CV —7.14 14.97 12.01
n1343
Leng h o digi one
Mean 5.00 4.33 4.75 4.67 Sex 1.246 .96 .353
SD —.58 .50 .58 Wi hin 9.256
SE —.33 .25 .33
CV —13.39 10.53 12.42
n1343
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
114
Table 21 .—Con inued.
Caa inga Ce ado Analysis o a iance
66 9$ 66 99 Fac o d MS FSigni icance
Leng h o digi h ee
Mean 62.00 61.33 60.50 61.00 Sex 1.367 .02 .880
SD —1.53 6.56 1.00 Wi hin 915.293
SE —.88 3.28 .58
CV —2.49 10.84 1.64
n1343
Leng h o digi ou
Mean 52.00 51.00 51.50 51.00 Sex 1.982 .32 .524
SD —1.73 2.52 1.00 Wi hin 93.022
SE —1.00 1.26 .58
CV —3.39 4.89 1.96
n1343
Leng h o digi i e
Mean 38.00 35.67 36.50 35.67 Sex 13.503 1.42 .263
SD —1.53 1.92 1.53 Wi hin 92.459
SE —.88 .96 .80
CV —4.29 5.26 4.29
n1343
Tibia leng h
Mean 11.00 10.00 10.50 10.33 Sex 1.512 2.27 .166
SD —0.58 .58 Wi hin 9.226
SE —0.29 .33
CV —05.52 5.61
n1343
Calca leng h
Mean 10.00 10.33 9.25 8.34 Sex 1.012 .01 .941
SD —1.16 1.26 1.53 Wi hin 92.059
SE —.67 .63 .88
CV —11.23 13.62 18.35
n1343
G ea es leng h o skull
Mean 13.20 13.23 13.43 13.20 Sex 1.060 .54 .485
SD —.31 .45 .36 Wi hin 8.112
SE —.18 .26 .21
CV —2.34 3.35 2.73
n1333
Condylobasal leng h
Mean 13.10 13.10 13.10 12.83 Sex 1.043 .24 .634
SD —.27 .66 .38 Wi hin 8.174
SE —.15 .38 .22
CV —2.06 5.04 2.96
n13 3 3
Zygoma ic b ead h
Mean 9.00 9.27 9.40 9.20 Sex 1.008 .08 .784
SD —.06 .53 0Wi hin 7.099
SE —.03 .31 0
CV —.65 5.64 0
n1332
Pos o bi al cons ic ion
Mean 3.70 3.67 3.70 3.70 Sex 1.001 .06 .812
SD —.06 .10 .17 Wi hin 8.01
1
SE —.03 .06 .10
CV —1.63 2.70 4.59
n13 3 3
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 115
Tabic 21 .—Con inued.
Caa inga Ce ado Analysis o a iance
66 99 66 99 Fac o d MS FSigni icance
Mas oid b ead h
Mean 8.80 8.67 8.90 8.40 Sex 1.280 2.20 .177
SD —.15 .62 .20 Wi hin 8.128
SE —.09 .36 .11
CV —1.73 6.97 2.38
n133 3
B ead h o b aincase
Mean 6.90 7.13 7.17 7.17 Sex 1.006 .12 .736
SD —.15 .32 .21 Wi hin 8.049
SE —.09 .19 .12
CV —2.10 4.46 2.93
n1333
Ros al b ead h
Mean 5.10 5.20 5.03 4.90 Sex 1.000 .00 1.000
SD -0.21 .17 Wi hin 8.036
SE —0.12 .10
CV —04.17 3.47
n1333
Heigh o b aincase
Mean 6.50 6.83 7.17 6.67 Sex 1.150 .68 .432
SD -.31 .61 .47 Wi hin 8.219
SE —.18 .35 .27
CV —4.54 8.51 7.05
n1333
B ead h ac oss he uppe mola s
Mean 6.60 6.47 6.73 6.70 Sex 1.033 .44 .524
SD -.15 .46 .10 Wi hin 8.074
SE —.09 .27 .06
CV —2.32 6.84 1.49
n1333
B ead h ac oss he uppe canines
Mean 3.70 3.83 4.00 3.73 Sex 1.048 1.63 .237
SD —.06 .17 .21 Wi hin 8.030
SE -.03 .10 .12
CV -1.57 4.25 5.63
n1333
Leng h o he maxilla y oo h ow
Mean 5.50 5.13 5.17 5.20 Sex 1.017 .73 .419
SD —.15 .15 0Wi hin 8.030
SE —.09 .09 0
CV —2.92 2.90 0
n1333
Leng h o he uppe mola i o m oo h ow
Mean 4.50 4.40 4.33 4.33 Sex 1.000 .02 .886
SD -.10 .06 .06 Wi hin 8.008
SE -.06 .03 .03
CV -2.27 1.39 1.39
n1333
Wid h o he wides mola
Mean 2.00 1.93 1.80 1.80 Sex 1.001 .05 .826
SD —.06 .10 .10 Wi hin 8.013
SE -.03 .06 .06
CV —3.11 5.56 5.56
n1333
116 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 21 .—Con inued.
Caa inga Ce ado Analysis o a iance
66 29 <3<5 22 Fac o d MS FSigni icance
G ea es leng h o he mandible
Mean 10.50 10.40 10.37 10.30 Sex 1.006 .09 .776
SD —.36 .32 .17 Wi hin 8.069
SE —.21 .19 .10
CV —3.46 3.09 1.65
n133 3
Leng h o he mandibula oo h ow
Mean 5.90 5.80 5.93 5.80 Sex 1.038 1.12 .321
SD —0.25 .27 Wi hin 8.033
SE —0.15 .15
CV —04.22 4.66
n13 3 3
Leng h o he co onoid p ocess
Mean 3.40 3.20 3.20 3.20 Sex 1.006 .15 .713
SD —.10 .36 .20 Wi hin 8.041
SE —.06 .21 .06
CV —3.13 11.25 3.13
n133 3
Table 28.— Summa y s a is ics o ex e nal and c anial cha ac e s (mean; s anda d de ia ion. SD; s anda d e o o he mean. SE;
coe icien o a ia ion. CV; sample size, n) o/'Neopla ymops ma og ossensis males and emales om he Caa inga biome. Aone-way
analysis o a iance (Model I)wi h eplica ion is p esen ed o each cha ac e . The exis ence o signi ican seconda y sexual a ia ion is
indica ed by P alues less han o equal o .050.
Caa inga Ce ado Analysis o a iance
66 22 66 22 Fac o d MS FSigni icance
To al leng h
Mean 79.14 76.81 Sex 164.621 4.19 .046
SD 3.12 4.49 Wi hin 46 15.405
SE .67 .88
CV 3.94 5.85
n22 26
Tail leng h
Mean 25.50 25.65 Sex 1.282 .04 .843
SD 2.84 2.50 Wi hin 46 7,074
SE .61 .49
CV 11.14 9.75
n22 26
Hind oo leng h
Mean 5.27 4.96 Sex 11.154 4.70 .040
SD .55 .45 Wi hin 46 .246
SE .12 .09
CV 10.44 9.07
n22 26
Ea leng h
Mean 13.36 13.12 Sex 1.734 1.71 .197
SD .58 .71 Wi hin 46 .429
SE .12 .14
CV 4.34 4.51
n22 26
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 117
Table 28. —Con inued.
Caa inga Ce ado Analysis o a iance
66 22 66 22 Fac o d MS FSigni icance
T agus leng h
Mean 3.14 3.04 Sex i.114 .21 .650
SD .89 .60 Wi hin 46 .556
SE .19 .12
CV 28.34 19.74
n22 26
Fo ea m leng h
Mean 30.05 30.23 Sex 1.409 .43 .514
SD .90 1.03 Wi hin 46 .947
SE .19 .20
CV 3.00 3.41
n22 26
Weigh
Mean 6.1
1
5.39 Sex 16.333 8.42 .006
SD .64 1.02 Wi hin 46 .753
SE .14 .20
CV 10.47 18.92
n22 26
Leng h o digi one
Mean 5.54 5.16 Sex 11.739 3.15 .083
SD .60 .85 Wi hin 45 .551
SE .13 .17
CV 10.83 16.47
n22 25
Leng h o digi h ee
Mean 55.22 54.16 Sex 113.330 6.30 .016
SD 1.11 1.70 Wi hin 45 2.1 16
SE .24 .34
CV 2.01 3.14
n22 25
Leng h o digi ou
Mean 46.05 44.04 Sex 147.064 4.00 .052
SD 3.36 3.49 Wi hin 45 11.776
SE .72 .70
CV 7.30 7.92
n22 25
Leng h o digi i e
Mean 33.64 32.96 Sex 15.353 1.91 .968
SD 1.59 1.74 Wi hin 45 2.801
SE .34 .35
CV 4.73 5.28
n22 25
Tibia leng h
Mean 10.27 10.04 Sex 1.634 1.34 .254
SD .63 .74 Wi hin 45 .474
SE .14 .15
CV 6.13 7.37
n22 25
Calca leng h
Mean 9.09 8.60 Sex 12.820 2.77 .103
SD 1.15 .87 Wi hin 45 1.018
SE .25 .17
CV 12.65 10.12
n22 25
BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
Table 28. —Con inued.
Caa inga Ce ado Analysis o a iance
<55 22 66 2$ Fac o d MS FSigni icance
G ea es leng h o skull
Mean 14.1 113.47 Sex 14.485 38.05 <.001
SD .35 .34 Wi hin 41 .118
SE .08 .07
CV 2.48 2.52
n21 22
Condylobasal leng h
Mean 13.91 13.25 Sex 14.738 49.40 <.001
SD .31 .31 Wi hin 41 .096
SE .07 .07
CV 2.23 2.34
n21 22
Zygoma ic b ead h
Mean 9.76 9.31 Sex 11.290 20.31 <.001
SD .25 .25 Wi hin 24 .064
SE .07 .08
CV 2.56 2.69
n15 11
Pos o bi al cons ic ion
Mean 3.36 3.29 Sex 1.059 7.01 .011
SD .10 .08 Wi hin 45 .008
SE .02 .02
CV 2.98 2.43
n22 25
Mas oid b ead h
Mean 9.60 9.06 Sex 12.143 19.92 <.001
SD .40 .22 Wi hin 28 .108
SE .10 .06
CV 4.17 2.43
n 16 14
B ead h o b aincase
Mean 7.08 6.81 Sex 1.768 14.58 <.001
SD .28 .18 Wi hin 41 .053
SE .06 .04
CV 3.95 2.64
n21 22
Ros al b ead h
Mean 4.33 3.88 Sex 12.431 49.88 <.001
SD .22 .22 Wi hin 45 .049
SE .05 .04
CV 5.08 5.67
n22 25
Heigh o b aincase
Mean 6.08 5.94 Sex 1.203 3.19 .082
SD .27 .24 Wi hin 44 .064
SE .06 .05
CV 4.44 4.04
n20 22
B ead h ac oss he uppe mola s
Mean 6.54 6.40 Sex 1.246 12.29 .001
SD .13 .15 Wi hin 44 .020
SE .03 .03
CV 1.99 2.34
n21 25
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 119
Table 28. —Con inued.
Caa inga Ce ado Analysis o a iance
66 99 66 99 Fac o d MS FSigni icance
B ead h ac oss he uppe canines
Mean 3.92 3.56 Sex 11.457 65.91 <.001
SD .15 .15 Wi hin 43 .022
SE .03 .03
CV 3.83 4.21
n21 24
Leng h o he maxilla y oo h ow
Mean 5.57 5.34 Sex 1.607 28.30 <.001
SD .16 .13 Wi hin 44 .022
SE .04 .03
CV 2.87 2.43
n21 25
Leng h o he uppe mola i o m oo h ow
Mean 4.27 4.20 Sex 1.058 2.84 .099
SD .12 .16 Wi hin 44 .021
SE .03 .03
CV 2.81 3.81
n2! 25
Wid h o he wides mola
Mean 2.00 1.96 Sex 1.014 2.31 .135
SD .08 .08 Wi hin 44 .006
SE .02 .02
CV 4.00 4.08
n21 25
G ea es leng h o he mandible
Mean 10.56 10.25 Sex 11.020 10.28 .003
SD .33 .30 Wi hin 42 .099
SE .07 .06
CV 3.13 2.93
n22 22
Leng h o he maxilla y oo h ow
Mean 5.91 5.69 Sex 1.965 69.28 <.001
SD .10 .13 Wi hin 43 .014
SE .02 .03
CV 1.69 2.28
n21 24
Leng h o he co onoid p ocess
Mean 3.76 3.46 Sex 1.950 25.87 <.001
SD .22 .15 Wi hin 41 .037
SE .05 .03
CV 5.85 4.34
n22 21
ac e s and s a is ically signi ican di e ences we e
de ec ed o all bu h ee o hese cha ac e s. N.
ma og ossensis is clea ly dimo phic wi h males
being la ge han emales.
Molossus a e (E. Geo oy, 1805)
This molossid ae ial insec i o e was a e in he
Caa ingas and was appa en ly absen om he Cha-
pada do A a ipe. Only one specimen, aju enile,
was cap u ed om he Caa ingas in Decembe .
Molossus molossus (Pallas, 1766)
This as - lying ae ial insec i o e was common in
bo h Caa ingas and Ce ado biomes. In bo h egions
o he No heas , M. molossus eached highes den-
si ies nea abandoned man-made s uc u es ha i
126 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
i ion, i is no clea why g ea e indi idual a ia-
ion, wi hou sex associa ed di e ences, could no
accomplish he same esul s. All o he ci cum-
s ances being equal, he e is no ap io i eason o
expec males o be la ge han emales o emales
o be la ge han males as a esul o he di e en ial
niche hypo hesis. A bes , once o he ac o s ini ially
selec o sex- ela ed size di e ences, hese di e -
ences could be accen ua ed by he bene i s o e-
duced esou ce compe i ion by niche pa i ioning
among he sexes.
The obse a ion ha emales a e equen ly la ge
han males has e ealed he gene al inadequacy o
bo h he sexual selec ion and niche pa i ioning hy-
po heses. Ralls (1976) ad ances he “Big Mo he ”
hypo hesis o accoun o cases o sexual size di-
mo phism in which emales a e la ge han males.
Fo a a ie y o easons, ala ge emale may be a
mo e i mo he ( ha is, p oduce ala ge numbe
o success ul p ogeny). La ge mo he s may p oduce
o sp ing wi h g ea e weigh s a bi h; p o ide mo e
o highe quali y milk; mo e e icien ly anspo o
de end young; o main ain homeo he mic equilib-
ium conduci e o emb yonic de elopmen . Mye s
(1978) examined size a ia ion in espe ilionid ba s
and only ound dimo phism in which emales we e
signi ican ly la ge han males. His analyses u he
showed ha he wings o emales we e p opo ion-
a ely la ge han hose o males, wi h he deg ee o
dimo phism co ela ed wi h li e size. Because o
hese obse a ions, Mye s (1978) sugges ed ha he
demands o mo he hood we e g ea e on species
p oducing mul iple young. Williams and Findley
(1979) con end ha he da a conside ed by Mye s
(1978) we e biased because monomo phic species
wi h mul iple young and highly dimo phic species
wi h single young we e excluded om he analyses.
In hei wo k (Williams and Findley, 1979), co e-
la ions be ween li e size and he deg ee o dimo -
phism we e no e iden . In acco d wi h he wo k o
Bogan (1975) and Findley and T au (1970), hey
conclude ha dimo phism in espe ilionids is
p obably ela ed o di e en ial he mo egula ion
a he han wing loading.
Wi hin he Chi op e a, sexual dimo phism is p i-
ma ily es ic ed o size, al hough he possession o
dimo phic glands in he Emballonu idae and Mo-
lossidae is qui e common (B adbu y, 1977). Six een
species o ba s om he No heas o B azil exhib-
i ed signi ican seconda y sexual a ia ion o eigh
o mo e mo phome ic cha ac e s. In app oxima e-
ly one- hi d o hose cases, he sample means o
emales we e la ge han he sample means o he
males (P. mac o is, Table 3; G. so icina, Table 13;
L. mo dax, Table 14; V. linea us, Table 18; A. con-
colo , Table 19; D. o undus, Table 22; L. bo ealis,
Table 25). Male sample means we e la ge han hose
o emales o he o he species in which s a is ically
signi ican dimo phism was de ec ed (N. lepo inus.
Table 4; T. si/ icola, Table 8; P. discolo , Table 10;
P. has a us, Table 11; A. geo oyi, Table 15; S’.
/ilium, Table 17; A. jamaicensis, Table 20; N. ma -
og ossensis, Table 28; M. molossus. Table 29).
Clea ly, dimo phism o size is common among he
Chi op e a, and unlike he si ua ion in many e -
eb a e o de s, emales a e equen ly signi ican ly
la ge han males. Ralls (1976) summa izes ou
knowledge o sexual dimo phism in ba s in which
he emales ha e been implica ed o be la ge han
males while Mye s (1978) and Williams and Findley
(1979) p esen addi ional in o ma ion on ba s in he
amily Vespe ilionidae. S a is ically signi ican di-
mo phism has only been subs an ia ed o 12 o he
45 po en ial cases ha Ralls epo s. Th ee o he
species implica ed as ha ing la ge emales han
males ei he show li le size dimo phism (A. li u a-
us) o exhibi s a is ically signi ican ly dimo phism
in which he males a e ac ually la ge han he e-
males (A. geo oyi and A. jamaicensis) in popula-
ions om B azil. The da a o D. o undus and L.
bo ealis subs an ia es Ralls’ (1976) claim ha e-
males a e la ge han males in hese species. Fu he ,
his s udy adds i e di e en species o Ralls’ lis o
ba s in which emales a e la ge han males (P. ma-
c o is, G. so icina, L. mo dax, V. linea us, A. con-
colo ).
Mos au ho s ag ee ha bo h selec ion p essu es
consis en wi h he hypo hesis o sexual selec ion
and he “Big Mo he ” hypo hesis a ec o g ea e
o lesse ex en s he obse ed size ela ions be ween
he sexes. The da a om his s udy indica es ha
selec i e p essu es a o ing la ge emales a e mo e
common han gene ally pe cei ed o he Chi op-
e a. Knowledge o ba beha iou and na u al his-
o y is somewha limi ed, bu i does no sugges a
e e sal o he heo e ical condi ions in which males
compe e among hemsel es o emales (see B ad-
bu y, 1977). Indeed, male ba s in es e y li le in
hei o sp ing o he han he ime and ene gy in-
ol ed in copula ion (see Kleiman and Malcolm,
1981). In hose cases whe e beha iou pa e ns ha e
been elucida ed, i appea s ha males compe e
among hemsel es o access o ep oduc i e e-
males (B adbu y, 1977). By implica ion, sexual se-
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 127
lec ion is p obably no he dominan ac o p o-
ducing la ge emales. Howe e , i may well be he
ac o ha educes he obse ed di e ences be ween
he sexes in hose cases whe e emales a e s a is i-
cally la ge han males. Al hough Mye s (1978) and
Williams and Findley (1979) disag ee in de ail abou
he pa icula ac o s ha a ec dimo phism in es-
pe ilionids, bo h a ibu e he obse ed di e ences
o biological condi ions consis en wi h he “Big
Mo he ” hypo hesis. This s udy canno dis inguish
be ween he he mo egula o y hypo hesis a o ed
by Williams and Findley (1979) and he wing-load-
ing hypo hesis o Mye s (1978); howe e , i can
quali y some o he con en ions hey p esen ed.
Mye s (1978) conjec u es ha small molossids
should show educed sexual dimo phism (males
la ge han emales) compa ed o la ge species in
he amily because hey sha e ce ain ligh and eed-
ing cha ac e is ics wi h he Vespe ilionidae. In he
wo species o molossids wi h su icien ly la ge sam-
ples o pe mi s a is ical analyses (M. molossus and
N. ma og ossensis), he males a e consis en ly la g-
e and s a is ically di e en han emales o bo h
c anial and ex e nal cha ac e s. Fu he , N. ma o-
g ossensis is one o he smalles membe s o he
Molossidae and i clea ly does no exhibi he e-
duced dimo phism p edic ed by Mye s (1978). Sim-
ila ly, Williams and Findley (1979) ejec Mye s’
con en ion by showing ha lasiu ine ba s in he am-
ily Ve spe ilionidae do no exhibi educed dimo -
phism ( emales la ge han males) al hough hey
would be p edic ed o do so by Mye s’ easoning.
The mo phome ic analyses o P. mac o is om he
Caa ingas a e in acco d wi h he obse a ions o
Mye s (1978) and o he s (see B adbu y, 1977) o
Emballonu ids; his small ba is dimo phic wi h e-
males la ge han males. O cou se, he p oponen s
o he he mo egula o y hypo hesis could also claim
ha he da a suppo s hei posi ion since he de-
mands o he mo egula ion a e accen ua ed o small
species. Ob iously, mo e da a need o be collec ed
om espe ilionid popula ions occu ing along la -
i udinal o al i udinal g adien s o see i species wi h
eu y he mal dis ibu ions exhibi deg ees o dimo -
phism which a e enhanced as empe a u e de-
c eases.
Al hough no nea ly as pe asi e as in he Ves-
pe ilionidae, he incidence o sexual dimo phism
wi h emales la ge han males is also common in
he Phyllos omidae (in con as , see he commen s
o B adbu y, 1977). Females a e la ge han males
in i e o he ele en species exhibi ing s a is ically
signi ican seconda y sexual dimo phism. Wi hin he
a ious phyllos omid sub amilies, i appea s ha
smalle species end o be dimo phic wi h emales
la ge han males, whe eas la ge species end o
exhibi dimo phism wi h la ge males. Al hough
Mye s (1978) sugges s a ela ion be ween eeding
ecology and dimo phism, li le e idence is a ailable
o suppo ha posi ion wi hin he Phyllos omidae.
The occu ence o dimo phism wi hin asub amily
does no seem o be ela ed o he eeding s a egies
o he cons i uen species.
ACKNOWLEDGMENTS
Ig a e ully acknowledge he ini ial inspi a ion, in aluable ad-
ice, cons an encou agemen , assis ance and iendship o D .
Michael A. Ma es, my majo ad iso . Ialso wish o since ely
hank D s. S e en J. Gauhn, Hugh H. Genoways, Richa d T.
Ha man, William Kod ich, and Robe J. Raikow, o hei
cons uc i e c i icisms and sugges ions. Nancy A. Bi a was an
island o s abili y du ing u bulen imes; he sel less aid can ne e
be epaid. Sand a Wigh and Cleon G os a e pa icula ly emem-
be ed o hei kindness, coope a ion, and skill in p oducing he
inal manusc ip . My iends and ellow esea che s in B azil,
D s. Thomas E. Lache , J ., Ka l E. S eilein, and Lau ie J. Vi ,
ole a ed much and con ibu ed in innume able ways— hei
com adeship, conce n, and assis ance we e in aluable. D . A is-
ides P. Leao, D . Paulo E. Vanzolini and he s a o he Aca-
demia B asilei a de Ciencias con ibu ed in coun less ways o he
success o he inal p ojec ; hei un i ing e o s in suppo o he
esea ch a e deeply app ecia ed. The Ins i u o B asilei o de Des-
en ol imen o Flo es al p o ided accommoda ions a he Flo-
es a Nacional A a ipe-Apodi. The manusc ip was signi ican ly
imp o ed by he c i ical e iews o D s. K. F. Koopman and D.
E. Wilson.
Many people assis ed wi h ield wo k. Joao Luna de Ca alho,
my ield assis an , iend and ’compad e’ aided in all aspec s o
he esea ch; much o he success o his esea ch can be a ibu ed
o his ai h ul and skill ul help. Ka l E. S eilein, Raimundo Lopes
da Sil a, Thomas E. Lache , J ., An onio Lemos Sil a, ‘Tico,’
and Lau ie E. Vi assis ed in he ield a imes also. Nume ous
B asilei os helped make my s ay in he No heas ewa ding bo h
p o essionally and pe sonally. In addi ion, many made hei a-
zendas a ailable o esea ch. An onio Zilclesio Pin o Sa ai a.
Isma Sa, Chico Ven u a, ‘Chame Anna,’ Rejane, Ricolice, So-
aya, Ma ia das Ne es, and he Teixei a amily a e especially
hanked o hei wa m h and conside a ion. Nancy A. Bi a ,
James D. Willig and Ma y R. Willig aided in da a compila ion
128 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
while he o me also p o ided edi o ial guidance h ough he
mo ass o o hog aphy. D . Hugh Genoways and Suzanne
McLa en kindly coope a ed in museum- ela ed aspec s o he
esea ch, while he o me also p o ided some o he species
iden i ica ions.
Field wo k was gene ously suppo ed by ag an o Michael A.
Ma es om he Academia B asilei a de Ciencias [p ojec numbe
85 (Ecology, e olu ion and zoogeog aphy o mammals), as apa
o he la ge p og am, ‘Ecological S udies o he Semi-a id Region
o No heas e n B azil’]. Addi ional unds we e p o ided h ough
D . C aig C. Black and D . Hugh H. Genoways h ough he M.
G aham Ne ing Resea ch Fund es ablished by agi om he
Co delia S. May Cha i able T us , Ca negie Museum o Na u al
His o y. Suppo was also p o ided by he Depa men o Bio-
logical Sciences, Loyola Uni e si y and aMellon Fellowship om
he Uni e si y o Pi sbu gh.
LITERATURE CITED
Ab’Sabe , A. N. 1970. P o incias geologicas edominios mo -
oclima icos no B asil. Geomo ologia Uni . Sao Paulo, Ins ,
de Geog a ia, 20:1-26.
Allen, G. M. 1908. No es on Chi op e a. Bull. Mus. Comp.
Zook, 52:25-62.
Al im, Paulo de T. 1949. Obse acoes ecologicas sob e lo a
da Regiao semi-a ida do No des e. Re is a Ce es Vicosa, 8:
105-1 11.
Ande sen, K. 1906. On ba s o he gene a Mic onyc e is and
Glyphonyc e is. Ann. Mag. Na . His ., se . 7, 18:50-65.
.1908. Amonog aph o he chi op e an gene a U o-
de ma, Enchis henes and A ibeus. P oc. Zool. Soc. London,
pp. 204-319.
Ande son, S. 1957. New eco ds o he ba , Anou a geo oyi
lasiopyga. Chicago Acad. Sci. Na . His . Misc., 159:1-3.
.1960. Neo opical ba s om Wes e n Mexico. LIni .
Kansas Publ., Mus. Na . His ., 14:1-8.
And ade, G. O. de, and R. C. Lins. 1964. In oducao ao es udo
dos “b ejos” pe nambucanos. A q. Ins . Cien. Te a, Reci e,
2:105-1 11.
Bogan. M. A. 1975. Geog aphic a ia ion in Myo is cali o -
nicus in he Sou hwes e n Uni ed S a es and Mexico. U.S.
Fish Wildl. Se ., Wildl. Res. Rep ., 3:1-31.
B adbu y, J. W. 1977. Social o ganiza ion and communica-
ion. Pp. 1-72, in Biology o ba s (W. A. Wimsa , ed.),
Academic P ess, New Yo k, 3:x i +651.
Bu meis e . H. 1854. Sys ema ische ube sich de hie e b as-
iliens welche mah end eine Reise du ch die P o inzen on
Rio de Janei o und Minas ge aes ...S. auge hie e (Mam-
malia). Geo g. Reime , Be lin, l:x +1-341.
Cab e a, A. 1917. Mami e os del Viaje al Paci ico. T ab. Mus.
Nac. Cien. Na . Zool., Mad id, 31:1-62.
.1957. Ca alogo de los Mami e os de Ame ica del Su .
I(Me a he ia-Unguicula a-Ca m o a). Re . Mus. A gen ino
Cienc. Na . "Be na dino Ri ada ia,” Zool., 4:1-307.
Cunha Viei a, C. O. da. 1942. Ensaio monog a ico sob e os
Qui op e os do B asil. A q. Zool., Es . Sao Paulo, 3:21 9—
471.
Cu ie , F. 1828. Desc ip ion d’un nou eau Gen e de Chau e-
sou is sous le nom de Fu ie. Memoi es du Museum d’His-
oi e Na u elle (Pa is), 16: 149-1 55.
Dalques , W. W. 1950. Reco ds o mammals om he Mex-
ican s a e o San Luis Po osi. Occas. Pape s Mus. Zool.,
Louisiana S a e Uni ., 23:1-15.
Daly, M., and M. Wilson. 1978. Sex, e olu ion and beha io .
Wadswo h Publishing Co., Inc., Belmon , Cali o nia.
Da win, C. 1859. On he o igin o species by means o na u al
selec ion, o he p ese a ion o a o ed aces in he s uggle
o li e. Mu ay, London.
.1871a. The descen o man, and selec ion in ela ion
o sex. Vol. I. Appel on, New Yo k.
.18716. The descen o man. and selec ion in ela ion
o sex. Vol. II. Appel on, New Yo k.
Da is, W. B. 1968. Re iew o he genus U ode ma (Chi op-
e a). J. Mamm., 49:676-698.
.1970. The la ge ui ba s (genus A ibeus) o Middle
Ame ica, wi h a e iew o he A ibeus jamaicensis complex.
J. Mamm., 51:105-122.
.1973. Geog aphic a ia ion in he ishing ba , Noc ilio
lepo inus. J. Mamm., 54:862-874.
Dias, J. 1960. Es udos sob e ageologia da Chapada do A a ipe,
no Municipio de Exu, Es ado de Pe nambuco. A q. Ins .
Pesq. Ag o., 5:33-40.
Die z, R. S., and J. C. Holden. 1970. Recons uc ion o Pan-
gaea: b eak up and dispe sion o con inen s, Pe mian o
p esen . J. Geophys. Res., 75:4939-4956.
Dobson, G. E. 1878. Ca alogue o he Chi op e a in he col-
lec ion o he B i ish Museum. B i ish Mus., London, xiii 4-
567 pp.
D'O bigny, A., and P. Ge ais. 1847. Mammi e es. In Voyage
dans 1’Ame ique Me idionale .. . annees 1826 . . .1833. (A.
D'O bigny), Pa is and S asbou g, 4(2): 1-32.
Ea ha , C. M., and N. K. Johnson. 1970. Size dimo phism
and ood habi s o No h Ame ican owls. Condo , 72:25 1—
264.
Eh lich, P. R., and P. H. Ra en. 1969. Di e en ia ion o pop-
ula ions. Science, 165:1228-1232.
Falcone , D. S. 1972. In oduc ion o quan i a i e gene ics.
The Roland P ess Co., New Yo k.
Feduccia, A., and B. H. Slaugh e . 1974. Sexual dimo phism
in ska es (Rajidae) and i s possible ole in di e en ial niche
u iliza ion. E olu ion, 28:164-168.
Fel en, H. 1956. Qui op e os (Mammalia: Chi op e a) en El
Sal ado . Comm. Ins . T op. In es . Cien ., Uni . El Sal-
ado , 5:153-170.
Findley, J. S., and G. L. T au . 1970. Geog aphic a ia ion
in Pipis eilus hespe us. J. Mamm., 50:362-367.
Fleming, T. H., E. T. Hoope , and D. E. Wilson. 1972. Th ee
Cen al Ame ican ba communi ies: s uc u e, ep oduc i e
cycles, and mo emen pa e ns. Ecology. 53:555-569.
F eise, F. W. 1938. The d ough egion o no heas e n B azil.
Geog . Re ., 28:363-378.
F o a Pessoa, O., A. B. Cou inho, D. de A. Lima, A. F. Fu -
ado, M. J. de A. Lima, S. M. Pe ei a, and E. A. Mansu .
1983 WILLIG- BRAZILIAN BAT COMMUNITIES 129
1971. Biologia No des e I. Ecologia e axionomia, Uni e -
sidade Fede al de Pe nambucco, Reci e.
Ga dne , A. L. 1976. The dis ibu ional s a us o some Pe-
u ian mammals. Occas. Pape s Mus. Zool., Louisiana S a e
Uni ., 48:1-18.
Geo oy-Sain -Hilai e, E. 1805. No e su une pe i e amille
de chau e-sou is d’Ame ique, designee sous le nom gene-
ique de Molossus. Bulle in des Sciences, pa la Socie e Phi-
loma ique de Pa is, 3(96):278—279 (=378-379).
.1810. Su les Phyllos omes e les especes de Mega-
de mes, deux. Gen es de la amille des Chau e-sou is. An-
nales du Museum d’His oi e Na u elle (Pa is), 15:157-198.
Ge ais, P. 1856. Chei op e es sud-Ame icains. Pp. 25-88,
pis. 7-15, in F. Cas elnau, Ammaux nou eaux ou a es de
1'Ame ique du Sud. Pa is. 1855 (1856).
Goodwin, G. G. 1942. Asumma y o ecognizable species o
Tona ia wi h desc ip ions o wo new species. J. Mamm.,
23:204-209.
.1959. Ba s o he subgenus Na alus. Ame . Mus. No -
i a es, 1977:1-22.
G ay, J. E. 1838. A e ision o he gene a o ba s (Vespe il-
ionidae), and he desc ip ion o some new gene a and species.
Mag. Zool. Bo ., 2:483-505.
.1842. Desc ip ions o some new gene a and i y un-
eco ded species o Mammalia. Ann. Mag. Na . His ., se .
1. 10:255-267.
Guima Aes, L. R. 1972. Con ibuicao aepidemiologia da pes e
endemica no No des e do B asil ees ado da Bahia. Es udo
das pulgas econ adas nessa egiao. Re is a B as, de Malano-
logia eDoencas T opicais, 24:95-164.
Ha e , J. 1979. Qua e na y biogeog aphy o opical lowland
Sou h Ame ica. Pp. 107-140, in The Sou h Ame ican he -
pe o auna: i s o igin, e olu ion and dispe sal (W. E. Duell-
man, ed.), Monog . Mus. Na . His ., Uni . Kansas P ess, 7:
1-485.
Hahn, W. L. 1907. A e iew o he ba s o he genus Hemi-
de ma. P oc. U. S. Na . Mus., 32:103-1 18.
Handley, C. O., J . 1960. Desc ip ions o new ba s om Pan-
ama. P oc. U. S. Na . Mus., 112:459-479.
.1966. Checklis o he mammals o Panama. Pp. 753-
795, in Ec opa asi es o Panama (R. L. Wenzel and V. J.
Tip on, eds.). Field Mus. Na . His ., Chicago, Illinois, xii +
861 pp.
.1976. Mammals o he Smi hsonian Venezuelan p oj-
ec . B igham Young Uni . Sci. Bull., 20:1-89.
Hei haus, E. R., T. H. Fleming, and P. A. Ople . 1975. Fo -
aging pa e ns and esou ce u iliza ion in se en species o
ba s in aseasonal opical o es . Ecology, 56:841-854.
He shko i z, P. 1949. Mammals o No he n Colombia. P e-
limina y epo s No. 5: Ba s (Chi op e a). P oc. U. S. Na .
Mus., 99:429-454.
.1972. The ecen mammals o he Neo opical egion:
azoogeog aphic and ecological e iew. Pp.3 11-431, in E o-
lu ion, mammals and sou he n con inen s (A. Keas , F. E k
and B. Glass, eds.). S a e Uni . New Yo k P ess, Albany,
New Yo k, 543 pp.
Hussa , S. L. 1976. Beha io al cha ac e displacemen : e i-
dence o ood pa i ioning in insec i o ous ba s. J. Mamm.,
57:331-338.
Husson, A. M. 1962. The ba s o Su iname. Zool. Ve hand.,
Leiden, 58:1-222.
James, P. E. 1942. La in Ame ica. The Odyssey P ess, New
Yo k.
Jones, J. K., J ., and D. C. Ca e . 1976. Anno a ed checklis
wi h keys o sub amilies and gene a. Pp. 7-38, in Biology o
ba s o he New Wo ld amily Phyllos oma idae. Pa I(R.
J. Bake , J. K. Jones, J ., and D. C. Ca e , eds.), Special
Publ. Mus., Texas Tech Uni ., 10:1-218.
Jones, J. K., J ., J. D. Smi h, and R. W. Tu ne . 1971. No e-
wo hy eco ds o ba s om Nica agua, wi h achecklis o
he Chi op e an auna o he coun y. Occas. Pape s Mus.
Na . His ., Uni . Kansas, 2:1-35.
Keas , A. 1966. T ophic in e ela ionships in he ish auna o
asmall s eam. Uni . Michigan, G ea Lakes Res. Di . Publ.,
15:51-79.
Kleiman, D. G., and J. R. Malcolm. 1981. The e olu ion o
male pa en al in es men in mammals. Pp. 347-387, in Pa-
en al ca e in mammals (D. J. Gube nick and P. H. Klop e ,
eds.). Plenum P ess, New Yo k, 459 pp.
Koopman, K. F. 1978. Zoogeog aphy o Pe u ian ba s wi h
special emphasis on he ole o he Andes. Ame . Mus. No -
i a es, 265 1:1-33.
Kumme , H. 1971. P ima e socie ies: g oup echniques o eco-
logical adap a ion. Aldine-A he on, Chicago, Illinois.
Lache , T. E., J . 1981. The compa a i e social beha io o
Ke odon upes is and Galea spixii in he xenc Caa inga o
no heas e n B azil. Bull. Ca negie Mus. Na . His ., 17:1 —
71.
LaVal, R. K. 1973. A e ision o he neo opical ba s o he
genus Myo is. Na . His . Mus. Los Angeles Co., Sci. Bull.,
15:1-54.
Leach, W. E. 1821. The cha ac e s o se en gene a o ba s wi h
oliaceous appendages o he nose. T ans. Linnean Soc. Lon-
don, 13:73-82.
Lima, J. L. 1926. Os mo cegos do collec ao do Museu Paulis a.
Re . Mus. Paulis a, Sao Paulo, 14:1-87.
Linnaeus, C. 1758. Sys ema Na u ae. 10 h ed., Holimae, Vol.
1,824 pp.
MacA hu , R. H. 1965. Pa e ns o species di e si y. Biol.
Re ., 40:510-533.
McNab, B. K. 1971. The s uc u e o opical ba aunas. Ecol-
ogy, 52:352-358.
Ma es, M. A., and D. F. Williams. 1977. Expe imen al suppo
o ood pa icle size esou ce alloca ion in he e omyid o-
den s. Ecology, 58:1186-1190.
Ma es, M. A., M. R. Willig, K. E. S eilein, and T. E. Lache ,
J . 1981. The mammals o no heas e n B azil: a p elimi-
na y assessmen . Ann. Ca negie Mus., 50:81-137.
Ma es, M. A., M. R. Willig, K. E. S eilein, and T. E. Lache ,
J . The ole o he B azilian Caa ingas in Sou h Ame ican
biogeog aphy: opical mammals in ad y egion. Unpub-
lished manusc ip .
Ma es, M. A., and R. A. Ojeda. 1982. Pa e ns o di e si y
and adap a ion in Sou h Ame ican hys icogna h oden s.
Pp. 393-432, in Sou h Ame ican mammalian biology (M.
A. Ma es and H. H. Genoways, eds.). Spec. Publ. Se . Py-
ma uning Lab. Ecology, Uni . Pi sbu gh, Pi sbu gh, Penn-
syl ania, 6:xii +1-539.
Ma kham, C. G. 1972. Aspec os clima ologicos da seca no
B asil-No des e. Reci e, Sudene Assesso ia Tecnica, Di isao
de Documen aijao.
.1974. Appa en pe iodici ies in ain all a Fo aleza,
Cea a, B azil. J. Appl. Me e ology, 13:176-179.
130 BULLETIN CARNEGIE MUSEUM OF NATURAL HISTORY NO. 23
.1975. Twen y-six-yea cyclical dis ibu ion o d ough
and lood in Cea a, B azil. The P o . Geog aphe , 27:454-
456.
Ma kham, C. G., and D. R. McLain. 1977. Sea su ace em-
pe a u e ela ed o ain in Cea a, no h-eas e n B azil. Na-
u e, 265:320-323.
Ma inez, L., and B. Villa-R. 1940. Segunda con ibucion al
conocimien o de los mu cielagos Mexicanos. II. Es ado de
Gue e o. Ann. Ins . Biol., Mexico, 11:291-361.
May , E. 1964. Sys ema ics and he o igin o species. Do e
Publica ions, Inc., New Yo k.
Melo, M. L. de. 1956. Excu sion Guidebook, No. 7. No heas .
In e na ional Geog aphical Union, B azilian Na ional Com-
mi ee, Rio de Janei o. 1956, 231 pp.
Mille , G. S., J . 1898. Desc ip ions o i e new phyllos ome
ba s. P oc. Acad. Na . Sci., Philadelphia, 50:326-337.
.1913. Fi e new mammals om opical Ame ica. P oc.
Biol. Soc. Washing on, 26:31-34.
Mulle , P. 1776. Des Ri e s Ca l on Linne Volls andigen
Na u sys ems. Nu nbe g, Supplemen sband, pp. 1-384.
Mye s, P. 1978. Sexual dimo phism in size o espe iliomd
ba s. Ame . Na ., 112:701-711.
Ojas i, J., and O. J. Lina es. 1971. Adiciones de la auna de
mu cielagos de Venezuela con no as sob e las especies del
gene o Diclidu us (Chi op e a). Ac a Biol. Venezolana, 7:
421-441.
Ol e s, I. 1818. Beme kungen zu Illige 's Uebe blick de Saug-
hie e, nach ih e Ve heilung ibe die Wel heile, ucksich -
lich de S id-ame icanischen A en (Species). Pp. 192-237,
in Jou nal on B asilien (W. L. Eschwege), He 2. In Neue
Biblio hek (F. J. Be uch), Bd. 15, Weima .
Pallas, P. S. 1766. Miscellanea zoologica. Hague Comi um,
xii +224 pp.
.1767. Vespe iliones in gene e, asc. 3, 35 p., pi. 1-4.
Spicilegia zoologica, Be lin, Vol. 1.
Pa e son, B., and R. Pascual. 1968. The ossil mammal auna
o Sou h Ame ica. Qua . Re . Biol., 43:409-451.
Pe e s, W. 1865. Die zu den Vampy i geho igen Flede hie e
und na u liche S ellung de Ga ung An ozous. Mona sb.
Kon. p euss. Akad. Wiss., Be lin, pp. 503-525.
.1866. Fe ne e Mi heilungen zu Kenn ni s de Fle-
de hie e, namen lich ibe A en de Leidene und B i i-
schen Museums. Mona sb. Kon. p euss. Akad. Wiss., Be lin,
pp. 672-681.
Pe e son, R. 1965. A e ision o he la -headed ba s o he
amily Molossidae om Sou h Ame ica and A ica. Con ib.
Royal On a io Mus., 64:1-32.
Pine, R. H. 1972. The ba s o he genus Ca ollia. Tech. Monog .,
Texas Ag ic. Exp. S a., Texas A&M Uni ., 8:1-125.
Pi lo , P. 1972. Chi op e es de Moyenne Amazome. Mam-
malia, 36:7 1-85.
Powe , D. M., and J. R. Tamsi . 1973. Va ia ion in Phyllos o-
mus discolo (Chi op e a: Phyllos oma idae). Canadian J.
Zool., 51:461-468.
Ralls, K. 1976. Mammals in which emales a e la ge han
males. Qua . Re . Biol., 51:245-276.
Rand, A. L. 1952. Seconda y sexual cha ac e s and ecological
compe i ion. Fieldiana, Zool., 34:65-70.
Rehn, J. A. G. 1900. No es on Chi op e a. P oc. Acad. Na .
Sci. Philadelphia, 52:755-759.
.1902. Th ee new Ame ican ba s. P oc. Acad. Na . Sci.
Philadelphia, 54:638-641.
Reis, A. C. de S. 1976. Clima da Caa inga. Ann. Acad. B asil.
Cienc., 48:325-335.
Robinson, D. 1971. Cos a Rican mammals. Pp. 1-6, in Hand-
book o opical biology in Cos a Rica (C. E. Schnell, ed.),
O ganiza ion o T opical S udies, San Jose.
Sanbo n, C. C. 1936. Reco ds and measu emen s o neo op-
ical ba s. Field Mus. Na . His ., Zool. Se ., 20:93-106.
.1949. Ba s o he genus Mic onyc e is and i s subgene a.
Fieldiana, Zool., 31:215-233.
.1955. Rema ks on he ba s o he genus Vampy ops.
Fieldiana, Zool., 37:403-413.
Sazima, I., and V. A. Taddei. 1976. Asecond eco d o he
Sou h Ame ican la -headed ba , Neopla ymops ma og os-
sensis. J. Mamm., 57:757-758.
Schinz, H. R. 1821. Das Thie eich. S u ga and T ibingen.
Vol. 1, xxx ni +894 p.
Schnell, G. D., and R. K. Selande . 1981. En i onmen al
and mo phological co ela es o gene ic a ia ion in mam-
mals. Pp. 60-99, in Mammalian popula ion gene ics (M. H.
Smi h and J. Joule, eds.), Uni . Geo gia P ess, A hens, Geo -
gia, 380 pp.
Schoene , T. W. 1967. The ecological signi icance o sexual
dimo phism in size in he liza d Anolis conspe us. Science,
155:474-477.
.1968. The Anolis liza ds o Bimini: esou ce pa i ion-
ing in acomplex auna. Ecology, 49:704-726.
Sch ebe , J. C. D. 1774. Die Saug hie e in Abbildungen nach
de Na u mi Besch eibungen, E langen. Theil 1, He s 1-
9, pp. 1-190.
Sealande , C. 1957. On ma ing sys ems and sexual selec ion.
Ame . Na ., 99:129-141.
Selande , R. K. 1966. Sexual dimo phism and di e en ial
niche u iliza ion in bi ds. Condo , 68:115-151.
.1972. Sexual selec ion and dimo phism in bi ds. Pp. 180-
230, in Sexual selec ion and he descen o man (1871-1971)
(B. Campbell, ed.), Aldine-A he on, Chicago, Illinois.
Sick, H. 1965. A auna do Ce ado. A q. Zool., 12:71-93.
Simpson, B. B., and J. Ha e . 1978. Specia ion pa e ns in he
Amazonian o es bio a. Ann. Re . Ecol. Sys ., 9:497-518.
Simpson, G. G. 1944. Tempo and mode in e olu ion. Columbia
Uni . P ess, New Yo k.
.1964. Species densi y o No h Ame ican ecen mam-
mals. Sys . Zool., 13:57-73.
Smi h, J. D. 1972. Sys ema ics o he Chi op e an amily Mo -
moopidae. Uni . Kansas Mus. Na . His ., Misc. Publ., 56:
1-132.
Smi h, J. D., and H. H. Genoways. 1974. Ba s o Ma ga i a
Island, Venezuela, wi h zoogeog aphic commen s. Bull. S.
Cali o nia Acad. Sci., 73:64-79.
Sokal, R. R. 1977. E olu ion and sys ema ics: in oduc o y
ema ks. Pp. 35-37, in The changing scenes in he na u al
sciences, 1776-1976 (E. C. Goulden, ed.), Ful on P ess, Inc.,
Lancas e , Pennsyl ania.
Sokal, R. R., and F. J. Rohl . 1969. Biome y. W. H. F eeman
and Company, San F ancisco, Cali o nia.
Spix, J. de. 1823. Si ma um e Vespe ilionum B asiliensium
species no ae. Monachii, iii +72 pp.
S eilein, K. E. 1981. The small mammal auna o he semia id
B azilian Caa inga: adap a ions o a luc ua ing en i on-
men . Unpublished Ph.D. hesis, Uni . Pi sbu gh, 190 pp.
Swanepoel, P., and H. H. Genoways. 1979. Mo phome ies.
Pp. 13-106, in Biology o ba s o he New Wo ld amily
1983 WILLIG —BRAZILIAN BAT COMMUNITIES 131
Phyllos oma idae. Pa III (R. J. Bake . J. K. Jones, J ., and
D. C. Ca e , eds.). Spec. Publ. Mus., Texas Tech Uni ., 16:
1-441.
Taddei, V. A. 1975a. Phyllos omidae (Chi op e a) do No e-
ociden al do Es ado de Sao Paulo I—Phyllos ominae. Cien-
cia eCul u a, 27:621-632.
.19756. Phyllos omidae (Chi op e a) do No e-ociden-
al do Es ado de Sao Paulo II —Glossophaginae; Ca olliinae;
S umi inae. Ciencia eCul u a, 27:723-734.
.1979. Phyllos omadidae (Chi op e a) do No e-ociden-
al do es ado de Sao Paulo III —S enode minae. Ciencia e
Cul u a, 31:900-914.
Tamsi , J. R., and D. Valdi ieso. 1963. Rep oduc i e cycle
o he big ui -ea ing ba , A ibeus li u a us Ol e s. Na u e,
198-204.
Thomas, M. E. 1972. P elimina y s udy o he annual b eeding
pa e ns and popula ion luc ua ions o ba s in h ee ecolog-
ically dis inc habi a s in sou hwes e n Colombia. Unpub-
lished Ph.D. disse ., Tulane Uni ., New O leans, 161 pp.
Thomas, O. 1903. No es on Sou h Ame ican monkeys, ba s,
ca ni o es and oden s, wi h desc ip ions o new species.
Ann. Mag. Na . His ., se . 7, 11:376-382.
T i e s, R. L. 1972. Pa en al in es men and sexual selec ion.
Pp. 136-179, in Sexual selec ion and he descen o man
(1871-1971) (B. S. Campbell, ed.), Aldine-A he on, Chi-
cago, Illinois, 378 pp.
Tu le, M. D. 1970. Dis ibu ion and zoogeog aphy o Pe-
u ian ba s, wi h commen s on na u al his o y. Uni . Kan-
sas Sci. Bull., 49:45-86.
Uieda, W„ I. Sazima, and A. S o i Filho. 1980. Aspec os
da Biologia do mo cego Fu ip e us ho ens (Mammalia, Chi-
op e a, Fu ip e idae). Re . B asil. Biol., 40:59-66.
Valdez, R. 1970. Taxonomy and geog aphic a ia ion o ba s
o he genus Phyllos omus. Ph.D. Disse a ion, Texas A&M
Uni .
Vanzolini, P. E. 1974. Ecological and geog aphical dis ibu ion
o liza ds in Pe nambuco, no heas e n B asil (Sauna). Pa-
peis A ulsos Zool., Sao Paulo, 28:61-90.
.1976. On he liza ds o aCe ado-Caa inga con ac :
E olu iona y and zoogeog aphic implica ions (Sau ia). Pa-
peis A ulsos Zool., Sao Paulo, 29:1 11-1 19.
Vizo o, L. D., and V. A. Taddei. 1976. No as sob e Molossops
emminckii emminckii eMolossops plani osl is (Chi op-
e a-Molossidae). Na u alia, 2:47-59.
Wagne . J. A. 1843. Diagnosnen neue A en B asilische
Hand lugle . A chi . u Na u geschich e (Wiegmann, Be -
lin) 9( 1): 365—368.
Walke . E. P. 1975. Mammals o he wo ld. The Johns Hop-
kins Uni . P ess, Bal imo e, 3 d ed.
Whi ake , R. H. 1975. Communi ies and ecosys ems. Mac-
millan Publ. Co., Inc., New Yo k.
Williams, D. F. 1978. Taxonomic and ka yologic commen s
on small b own ba s, genus Ep esicus, om Sou h Ame ica.
Ann. Ca negie Mus., 47:361-383.
Williams, D. F., and J. S. Findley. 1979. Sexual size dimo -
phism in espe ilionid ba s. Ame . Midland Na ., 102: 113-
126.
Willig, M. R. The biology o Neopla ymops ma og ossensis.
(unpublished manusc ip ).
Wilson, E. O. 1975. Sociobiology. The Belnap P ess o Ha a d
Uni . P ess, Camb idge, Massachuse s.
Wilson. J. W. 1974. Analy ical zoogeog aphy o No h Ame -
ican mammals. E olu ion, 28:124-140.
Yabloko , A. V. 1974. Va iabili y o mammals. Ame ind Pub-
lishing Co., P . L d., New Delhi.
