The mammals of Paracou, French Guiana: a Neotropical lowland rain forest fauna. Pt. 1. Bats.

2
CONTENTS
Abs ac ....................................................................... 3
Re´sume´ ........................................................................ 4
In oduc ion .................................................................... 6
Acknowledgmen s ............................................................ 7
Si e Desc ip ion ................................................................ 8
Topog aphy, Soils, and Vege a ion .............................................. 8
Clima e and Phenology ...................................................... 13
Ba In en o y Me hods ........................................................ 14
Mis ne ing ................................................................ 15
Sea ching o Roos s ........................................................ 17
Specimen Collec ion and P ese a ion ......................................... 19
Acous ic Su ey ............................................................ 21
Scheduling and Pe sonnel .................................................... 21
Sys ema ic Accoun s .......................................................... 22
Emballonu idae ............................................................. 23
Noc ilionidae ............................................................... 42
Mo moopidae .............................................................. 45
Phyllos omidae ............................................................. 45
Desmodon inae ........................................................... 46
Glossophaginae .......................................................... 47
Phyllos ominae ........................................................... 57
Ca olliinae ............................................................... 94
S enode ma inae .......................................................... 97
Fu ip e idae ............................................................... 128
Thy op e idae ............................................................. 129
Vespe ilionidae ........................................................... 131
Molossidae ............................................................... 146
Analyses o Sampling ........................................................ 169
Sampling Resul s om Di e en Me hods .................................... 169
Sampling Resul s in Di e en Habi a s ....................................... 177
Sampling Resul s in Di e en Yea s ......................................... 181
Es ima ing Comple eness ................................................... 182
Discussion .................................................................. 185
Taxonomic Composi ion and Biogeog aphy ................................... 185
Species Richness .......................................................... 190
T ophic Guilds, Habi a Use, and O he Topics ................................ 192
Sugges ions o Fu u e Wo k .................................................. 198
Imp o ing In en o y E iciency ............................................. 199
S anda ds o Repo ing In en o y Da a ...................................... 200
Quan i a i e Me hods o Di e si y Compa isons .............................. 201
P ospec s o Rapid Di e si y Assessmen .................................... 202
Re e ences .................................................................. 202
Appendix 1: Ba s P e iously Repo ed om F ench Guiana o Su inam, bu no .....
Cap u ed a Pa acou ....................................................... 215
Appendix 2: Species Ma ix o 14 Neo opical Rain o es Ba In en o ies .......... 218
1998 3SIMMONS AND VOSS: PARACOU BATS
ABSTRACT
This epo desc ibes he esul s o ba in en-
o y ieldwo k a Pa acou, a lowland ain o es lo-
cali y in no he n F ench Guiana. Wo king wi hin
a 3-km adius o e he cou se o 168 sampling
days om 1991 o 1994, we cap u ed 3126 ba s,
o which abou 78% we e aken in g ound-le el
mis ne s, 10% in mis ne s suspended abo e
g ound le el, and 12% a oos s. We iden i ied a
o al o 78 species, including 10 emballonu ids, 2
noc ilionids, 1 mo moopid, 49 phyllos omids, 1
u ip e id, 1 hy op e id, 5 espe ilionids, and 9
molossids.
Among ou axonomic esul s, we desc ibe a
new species o Mic onyc e is (sensu s ic o) o
hono And e´ B osse , pionee ing monog aphe o
ain o es ba aunas in India, A ica, and Sou h
Ame ica. In addi ion, we epo he i s eco ds
o eigh o he species om F ench Guiana: Cen-
onyc e is maximiliani,Pe op e yx kapple i,Sac-
cop e yx gymnu a,Mic onyc e is homezi,Mic o-
nyc e is schmid o um,Molossops pa anus,Mo-
lossus sinaloae, and P omops cen alis. Mos o
hese we e p e iously known om Su inam, bu
he ange ex ensions a e signi ican o Saccop-
e yx gymnu a (ca. 900 km), Mic onyc e is hom-
ezi (2200 km), and M.schmid o um (1500 km).
Al oge he , he known ba auna o F ench Guiana
now consis s o 102 species.
The ollowing signi ican axonomic esul s a e
also epo ed he ein. (1) Compa ison o Pa acou
specimens e e able o Pe op e yx mac o is (Wag-
ne ) wi h he holo ype o P. ini a is Mille sup-
po s he conclusions o ecen in es iga o s ha
hese axa a e sepa a e species. (2) Mo phological
a ia ion among specimens o small Choe oniscus
om Pa acou, oge he wi h examina ion o ype
ma e ial and a c i ical e iew o he li e a u e,
sugges ha C.mino (Pe e s), C.in e medius (Al-
len and Chapman), and C.inca Thomas a e con-
speci ic; he oldes a ailable name o he species
is Pe e s’. (3) Glyphonyc e is Thomas (including
Ba iconyc e is Hill as a synonym), Mic onyc e is
G ay (including Xenoc enes Mille as a synonym),
and T inyc e is Sanbo n a e ediagnosed as dis-
inc gene a; Lamp onyc e is Sanbo n and Neo-
nyc e is Sanbo n, wo o he e s while subgene a
o Mic onyc e is (sensu la o), should also be ea -
ed as ull gene a. (4) Mic onyc e is homezi Pi lo ,
based on a los holo ype and p e iously consid-
e ed a nomen dubium, is edesc ibed and edi-
agnosed as a alid species. (5) Mic onyc e is
megalo is (G ay) and M.mic o is Mille a e dis-
inc species ep esen ed by sympa ic collec ions
om Pa acou and o he ma e ial simila ly in e -
p e ed by ecen in es iga o s. (6) Mimon benne -
ii (G ay) and M.cozumelae Goldman a e diag-
nosable as dis inc species by consis en ex e nal
and c anioden al cha ac e di e ences. (7) Ec o-
phylla H. Allen is ediagnosed o include Meso-
phylla Thomas in ecogni ion o he sis e -g oup
ela ionship be ween E.alba H. Allen and E.mac-
connelli (Thomas). (8) The ecen hypo hesis ha
S u ni a lilium (E. Geo oy) and S.luisi Da is
a e conspeci ic is ejec ed as implausible because
o enchan c anial cha ac e di e ences. (9) The
Venezuelan and F ench Guianan specimens e-
cen ly iden i ied in he li e a u e as Ep esicus an-
dinus J. A. Allen a e no conspeci ic wi h he ho-
lo ype o ha species; ins ead, examina ion o
ype specimens, o he compa a i e ma e ial, and
he p ima y li e a u e sugges s ha his ma e ial
is e e able o E.chi iquinus Thomas. (10) All
cu en ly accep ed synonymies o axa included
wi hin Da is’s (1966) andinus g oup o Ep esicus
a e appa en ly inco ec ; in ou iew, E.andinus
is a senio synonym o E.mon osus Thomas and
E.chi alensis An hony, whe eas E.chi iquinus is
a senio synonym o E.inca Thomas. (11) We
e iew he con en s o Cynomops Thomas, cu -
en ly anked as a subgenus o Molossops Pe e s,
and abula e diagnos ic cha ac e s o he ou
species we ega d as alid: M.ab asus (Tem-
minck), M.g eenhalli (Goodwin), M.pa anus
(Thomas), and M.plani os is (Pe e s). (12) Mo-
lossus ba nesi Thomas is a alid species eadily
dis inguishable om bo h M.molossus (Pallas)
and M.coibensis J. A. Allen.
Analyses o ou sampling esul s indica e ha
(1) dis inc se s o species a e e ec i ely sampled
by di e en cap u e me hods; (2) dis inc se s o
species inhabi di e en local habi a s; and (3) in-
c eased sampling e o wi h any me hod gene -
ally esul s in mo e species, al hough he a e o
accumula ion declines wi h sample size (numbe
o cap u es). Based on nonpa ame ic s a is ical
ex apola ions, we es ima e ha he Pa acou ba
auna p obably consis s o somewhe e be ween 85
and 95 species; he mo e conse a i e ichness
es ima o sugges s ha ou in en o y is pe haps
abou 90% comple e. Judging om he known o
in e ed beha io s o he a e axa (single ons and
double ons) in ou da a, mos o he local species
missing om his in en o y a e p obably ae ial
insec i o es, gleaning insec i o es, o nec a i-
o es.
In e ms o highe axonomic composi ion, he
ba auna a Pa acou is ypical o hose ound
h oughou he humid Neo opical lowlands. A
quan i a i e analysis o aunal simila i y a he
species le el among 14 ain o es locali ies cho-
sen as exempla s clus e s he Pa acou lis wi h
o he s p e iously epo ed om he Guiana sub-
4 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
egion o Amazonia, nex wi h lis s om else-
whe e in Amazonia, and las ly wi h Cen al
Ame ican lis s. No su p isingly, pai wise simila -
i y alues show a posi i e co ela ion be ween
aunal esemblance and geog aphic p oximi y
wi hin he Neo opical ain o es biome. Many
(47%) o he ba species in he Pa acou auna a e
essen ially pan-Neo opical in dis ibu ion and
mos o hese a e also known om habi a s o he
han ain o es . The emaining species exhibi
mo e es ic ed geog aphic dis ibu ion pa e ns,
bu ue Amazonian endemics cons i u e only a
mino ac ion o he Pa acou ba auna.
Species ichness compa isons among in en o y
si es a e complica ed by p oblems o inconsis en
me hodology, habi a ep esen a ion, and sam-
pling e o . Fo example, he appa en ly excep-
ional di e si y o emballonu ids, phyllos omines,
and molossids in he Pa acou auna is plausibly
explained by ou in ensi e use o ele a ed ne ing
and oos su eys, and by p olonged e o , all o
which ac o s ac o educe he well-known cap-
u e bias o g ound-le el mis ne s (which consis-
en ly unde sample hese axa in he sho e m).
Howe e , he low ichness o ca olliines and s e-
node ma ines a Pa acou by compa ison wi h mos
o he Amazonian (especially wes e n Amazonian)
locali ies is appa en ly eal. The only app oxi-
ma ely alid s a is ical compa ison o species
ichness ha we can make be ween si es based on
published cap u e- equency da a sugges s an in-
c ease o app oxima ely 50% in unde s o y ba s
om eas e n Cen al Ame ica o Amazonia, bu
he eal o a i ac ual na u e o his es ima ed di -
e ence emains o be e alua ed.
A ophic classi ica ion o Pa acou ba s indi-
ca es ha ae ial insec i o es a e he mos speciose
eeding guild in he local auna, ollowed by
gleaning animali o es, ugi o es, and nec a i-
o es; omini o es, sangui o es, and pisci o es a e
mino componen s. Pa e ns o di e en ial habi a
use among species wi hin some eeding guilds can
be in e ed om ou cap u e- equency da a, no-
ably o ae ial insec i o es and ugi o es. By
con as , gleaning animali o es appea o be
la gely es ic ed o p ima y o es , a puzzling
phenomenon p e iously epo ed om o he Neo-
opical ain o es locali ies.
To acili a e u u e in en o y ieldwo k we p o-
ide (1) de ailed desc ip ions o su ey and cap-
u e me hods, (2) illus a ions o mos local hab-
i a s ecognized as dis inc , (3) comple e b eak-
downs o cap u e equencies by me hod and hab-
i a o each species, (4) pho og aphs o nume ous
oos s a which ba s we e cap u ed, and (5) de-
sc ip ions and/o illus a ions o use ul cha ac e s
o iden i ying species hi he o equen ly con-
used in he ield. Finally, we make ecommen-
da ions o imp o ing ba in en o y e iciency,
sugges minimal s anda ds o epo ing in en o y
da a, u ge he adop ion o quan i a i e me hods
o in e si e di e si y compa isons, and commen
on he p ospec s o apid di e si y assessmen o
ain o es ba aunas.
RE
´SUME
´
Dans ce appo nous de´c i ons les e´sul a s de
l’in en ai e du peuplemen de chau es-sou is e -
ec ue´ su le e ain a` Pa acou, un si e de o eˆ
hyg ophile de basse al i ude en Guyane anc¸aise.
Dans un ayon de 3 km e´ udie´ pendan 168 jou s
d’e´chan illonnage de e ain de 1991 a` 1994, nous
a ons cap u e´ 3126 chau es-sou is, soi 78% dans
des ile s p e`s du sol, 10% dans des ile s e´le e´s
suspendus au-dessus du sol, e 12% a` des do oi s.
Nous a ons iden i ie´ 78 espe`ces au o al, don 10
emballonu ide´s, 2 noc ilionide´s, 1 mo moopide´,
49 phyllos omide´s, 1 u ip e ide´, 1 hy op e ide´, 5
espe ilionide´s e 9 molosside´s.
Pa mi nos e´sul a s axonomiques nous de´c i-
ons une espe`ce nou elle de Mic onyc e is (sensu
s ic o) en l’honneu d’And e´ B osse , au eu de
monog aphies inno a ices su les aunes de chau-
es-sou is des o eˆ s hyg ophiles de l’Inde, de
l’A ique e de l’Ame´ ique du Sud. De plus, nous
p e´sen ons les p emie` es donne´es conce nan la
p e´sence de hui au es espe`ces en Guyane an-
c¸aise: Cen onyc e is maximiliani,Pe op e yx
kapple i,Saccop e yx gymnu a,Mic onyc e is
homezi,Mic onyc e is schmid o um,Molossops
pa anus,Molossus sinaloae e P omops cen alis.
La plupa de ces espe`ces e´ aien de´ja` connues du
Su iname, mais les ex ensions des ai es de e´pa -
i ion son subs an ielles pou Saccop e yx gym-
nu a (p e`s de 900 km), Mic onyc e is homezi
(2200 km) e M.schmid o um (1500 km).
Dans ce a ail, nous appo ons e´galemen les
impo an s e´sul a s axonomiques sui an s. (1) La
compa ison d’exemplai es de Pa acou se appo -
an a`Pe op e yx mac o is (Wagne ) a ec
l’holo ype de P. ini a is Mille , con i me la con-
clusion d’au eu s e´cen s, que ces axons appa -
iennen a` des espe`ces di e´ en es. (2) La a ia-
bili e´ mo phologique de´celable pa mi les exem-
plai es de pe i s Choe oniscus de Pa acou, con-
join emen a ec l’e´ ude des ypes e a ec une
e´ ision c i ique de la li e´ a u e, sugge` en que C.
mino (Pe e s), C.in e medius (Allen e Chap-
man) e C.inca Thomas son conspe´ci iques; le
plus ancien nom disponible pou l’espe`ce es celui

1998 5SIMMONS AND VOSS: PARACOU BATS
de Pe e s. (3) De nou elles diagnoses son don-
ne´es pou Glyphonyc e is Thomas (qui inclu Ba -
iconyc e is Hill comme synonyme), e pou T i-
nyc e is Sanbo n; Lamp onyc e is Sanbo n e
Neonyc e is Sanbo n, deux anciens sous-gen es de
Mic onyc e is (sensu la o) de aien e´galemen
eˆ e conside´ e´s comme des gen es. (4) Mic o-
nyc e is homezi Pi lo , onde´ su un holo ype pe -
du e conside´ e´ p e´ce´demmen comme un nomen
nudum, es ede´c i e diagnos ique´a` nou eau
comme une espe`ce alable. (5) Mic onyc e is
megalo is (G ay) e M.mic o is Mille son des
espe`ces dis inc es ep e´sen e´es pa des exemplai-
es e´col e´s en sympa ie a` Pa acou e pa d’au e
ma e´ iel, ce qui es in e p e´ e´ de ac¸on similai e
pa des che cheu s e´cen s. (6) Mimon benne ii
(G ay) e M.cozumelae Goldman peu en eˆ e
diagnos ique´es comme deux espe`ces dis inc es
g aˆce a` des di e´ ences de ca ac e` es ex e nes e
c anioden ai es. (7) Le gen e Ec ophylla H. Allen
es diagnos ique´a` nou eau pou y inclu e Meso-
phylla Thomas, ce qui econnai ainsi les appo s
d’espe`ces-soeu s en e E.alba H. Allen e E.mac-
connelli (Thomas), ce e de nie` e appele´e aupa-
a an Mesophylla. (8) L’hypo he`se, e´cemmen
e´mise, que S u ni a lilium (E. Geo oy) e S.luisi
Da is son conspe´ci iques es eje e´e comme e´ an
peu plausible a` cause de di e´ ences ma que´es des
ca ac e` es c aniaux. (9) Les exemplai es du Ve´-
ne´zue´la e de Guyane anc¸aise e´cemmen de´ e -
mine´s dans la li e´ a u e comme appa enan a` Ep-
esicus andinus J. A. Allen ne son pas conspe´-
ci iques a ec l’holo ype de ce e espe`ce; en e-
anche, l’e´ ude des exemplai es ypes, de ma e´ iel
compa a i addi ionnel, e de la li e´ a u e o igi-
nale, sugge` en que ce ma e´ iel peu eˆ e appo e´
a` E.chi iquinus Thomas. (10) Tou es les syno-
nymies accep e´es a` l’heu e ac uelle pou les ax-
ons inclus au sein du g oupe d’Ep esicus appele´
g oupe andinus pa Da is (1966) semblen eˆ e
inco ec es; d’ap e`s nous, E.andinus es un syn-
onyme plus ancien d’E.mon osus Thomas e de
E.chi alensis An hony, andis que E.chi iquinus
es un synonyme plus ancien d’E.inca Thomas.
(11) Nous e´ isons la eneu de Cynomops Tho-
mas, ac uellemen classe´ comme un sous-gen e de
Molossops Pe e s, e p e´sen ons sous o me a-
bulai e les ca ac e` es diagnos iques des qua e es-
pe`ces que nous conside´ ons comme alables: M.
ab asus (Temminck), M.g eenhalli (Goodwin),
M.pa anus (Thomas), e M.plani os is (Pe e s).
(12) Molossus ba nesi (Thomas) es une espe`ce
alable, aise´men di e´ enciable a` la ois de M.
molossus (Pallas) e de M.coibensis J. A. Allen.
L’analyse de no e e´chan illonnage indique que:
(1) des assemblages d’espe`ces dis inc s son e -
ec i emen e´chan illonne´s pa di e´ en es me´ ho-
des de cap u e; (2) des assemblages d’espe`ces dis-
inc es occupen des bio opes di e´ en s au sein de
no e zone d’e´ ude; e (3) un e o acc u
d’e´chan illonnage, quelle que soi la me´ hode,
p odui en gene al da an age d’espe`ces, bien que
le aux d’accumula ion baisse a ec une augmen-
a ion de la aille de l’e´chan illon (nomb e de cap-
u es). Su la base d’ex apola ions s a is iques
non-pa ame´ iques, nous es imons que le peuple-
men de chau es-sou is de Pa acou con ien en e
85 e 95 espe`ces; l’es ima ion conse a ice de
ce e ichesse sugge` e que no e in en ai e es
peu -eˆ e a` 90% comple . Si nous nous basons su
les compo emen s, connus ou in e´ e´s, des axons
a es ( ep e´sen e´s dans no e e´chan illonnage pa
des exemplai es uniques ou des double s), la plu-
pa des espe`ces qui manquen enco e a` ce in-
en ai e son p obablemen des insec i o es
ae´ iens, des insec i o es glaneu s ou des nec a i-
o es.
En ce qui conce ne sa composi ion axono-
mique sup age´ne´ ique, la aune de chau es-sou is
de Pa acou es ypique de celles que l’on ou e
dans l’ensemble des e´gions humides ne´o opi-
cales de basse al i ude. Une analyse quan i a i e
de la essemblance aunis ique au ni eau spe´ci-
ique pa mi 14 si es choisis comme exemples de
o eˆ hyg ophile, place la lis e de Pa acou,
d’abo d pa mi d’au es men ionne´es aupa a an
de la sub e´gion guyanaise de l’Amazonie, ensui e
pa mi des lis es p o enan d’au es e´gions de
l’Amazonie, e en in pa mi des si es d’Ame´ ique
cen ale. Les aleu s des indices de simila i e´,
compa e´es deux a` deux, mon en une co e´la ion
posi i e en e la essemblance aunis ique e la
p oximi e´ge´og aphique des si es au sein du bio-
me o eˆ hyg ophile ne´o opicale. De nomb euses
espe`ces de chau es-sou is du peuplemen de Pa -
acou (47% du o al) on une e´pa i ion essen iel-
lemen pan-ne´o opicale, e la plupa d’en e elles
son connues d’habi a s au es que la o eˆ hyg o-
phile. Les au es espe`ces mon en des sche´mas
de dis ibu ion ge´og aphiquemen plus es ein s,
mais les ais ende´miques amazoniens ne cons i -
uen qu’une ac ion mineu e du peuplemen de
chau es-sou is de Pa acou.
Les compa aisons de ichesse spe´ci ique pa mi
les si es in en o ie´s p e´ce´demmen son endues
complique´es pa des p oble`mes de me´ hodologie
ine´gale, de e´p esen a ion d’habi a s e d’e o
d’e´chan illonnage. Pa exemple, l’explica ion de
la di e si e´, a` p emie` e ue excep ionnelle, des
emballonu ide´s, phyllos omine´s e molosside´s
dans la aune de Pa acou, es endue plausible pa
no e emploi in ensi de ile s e´le e´s e pa no e
e´ ude de do oi s, ainsi que pa un e o p olonge´,
me´ hodes d’e´chan illonnage qui ou es e´duisen
la endance bien connue de su cap u e lo sque les
ile s son pose´s p e`s du sol, ce qui donne, dans
6 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
le cou e me, un sous-e´chan illonnage de ces
axons. Tou e ois, la aible ichesse des ca olliine´s
e des s e node ma ine´s a` Pa acou, compa e´e a`
celle de la plupa des au es locali e´s amazonien-
nes (spe´cialemen d’Amazonie occiden ale), ne
pa aıˆ pas a e ac uelle. La seule compa aison s a-
is ique a` peu p e`s alable que nous pou ons ai e
de ichesse spe´ci ique en e si es, base´e su des
donne´es de e´quences de cap u es dans la li e´-
a u e, sugge` e que la p opo ion de chau es-sou -
is du sous-bois augmen e d’en i on 50% en e
l’Ame´ ique cen ale o ien ale e l’Amazonie, mais
la na u e e´elle ou a i ac uelle de ce e di e´ ence
es ime´e es e a`eˆ e e´ alue´e.
Une classi ica ion ophique des chau es-sou is
de Pa acou indique que les insec i o es ae´ iens
son la guilde du peuplemen local con enan le
plus d’espe`ces, sui ie pa les animali o es gla-
neu s, les ugi o es e les nec a i o es; les omni-
o es, sangui o es e pisci o es ne ep e´sen en
que des ac ions mineu es. Les pa ons
d’u ilisa ion di e´ en ielle des habi a s au sein de
ce aines guildes alimen ai es peu en eˆ e de´dui s
a` pa i de nos donne´es de e´quences de cap u es,
no ammen pou les insec i o es ae´ iens e les u-
gi o es. Pa con e, les animali o es glaneu s
semblen eˆ e la gemen limi e´s a` la o eˆ p imai-
e, un phenome`ne de´conce an , de´ja` men ionne´
aupa a an pou d’au es locali e´s de la o eˆ hy-
g ophile ne´o opicale.
Dans le bu de acili e de u u s in en ai es de
e ain: (1) nous donnons des desc ip ions de´ ail-
le´es de nos me´ hodes d’e´ ude e de cap u e, (2)
nous illus ons la plupa des habi a s locaux e-
connus comme dis inc s, (3) nous indiquons, pou
chaque espe`ce, la e´pa i ion comple` e des e´-
quences de cap u e pa me´ hode e pa habi a , (4)
nous p esen ons des pho og aphies des nomb eux
do oi s ou` nous a ons cap u e´ des chau es-sou -
is, e (5) nous de´c i ons e /ou illus ons des ca -
ac e` es u iles pou la de´ e mina ion d’espe`ces qui
on e´ e´ e´quemmen con ondues jusqu’a` p e´sen
su le e ain. Finalemen , nous aisons des ecom-
manda ions isan a` ame´lio e l’e icaci e´ des in-
en ai es de chau es-sou is, nous sugge´ ons des
ni eaux minimaux pou les appo s de donne´es
d’in en ai es, nous conseillons i emen
l’adop ion de me´ hodes quan i a i es pou des
compa aisons de di e si e´ in e -si es, e nous ai-
sons des ema ques au suje de l’e´ alua ion apide
de la di e si e´ des aunes de chau es-sou is en
o eˆ hyg ophile.
INTRODUCTION
The mos species- ich mammalian aunas
in he wo ld a e p obably hose ha inhabi
he lowland ain o es s o ce ain pa s o
Cen al and Sou h Ame ica, bu compelling
e idence o suppo his conjec u e is spa se.
Much o wha we know abou Neo opical
ain o es mammal di e si y comes om jus
a hand ul o si es whe e la ge species lis s
we e buil up o e many yea s as byp oduc s
o ield esea ch on beha io , communi y
ecology, demog aphy, and o he na u al his-
o y opics (e.g., An hony, 1921; Ende s,
1935; Glanz, 1982; Janson and Emmons,
1990; Handley e al., 1991; Timm, 1994;
Hu e e e al., 1995). Such haphaza dly col-
lec ed da a a e di icul o analyze o com-
ple eness, and es ima ing species ichness a
mos si es is he e o e a ma e o mo e-o -
less educa ed guesswo k (Voss and Emmons,
1996).
Fo bo h esea ch and conse a ion pu -
poses i would be use ul o know how much
ime and e o and wha combina ion o in-
en o y me hods a e necessa y o ob ain ea-
sonably comple e species lis s o mammals
a ain o es ed Neo opical locali ies. Un o -
una ely, he cu en li e a u e does no p o-
ide an adequa e basis o making such es-
ima es. Among o he p oblems, ew in en-
o y epo s ha e included su icien me h-
odological in o ma ion o know exac ly wha
was done, he local habi a s ac ually sampled
o mammals a e seldom iden i ied explici ly,
ele an measu es o sampling e o a e a e-
ly p o ided, and he equency da a neces-
sa y o judge in en o y comple eness a e al-
mos ne e published.
Species iden i ica ions a e ano he majo
impedimen o in e p e ing in en o y esul s.
Mos gene a o Neo opical ain o es mam-
mals ha e ne e been e ised, o we e e-
ised long ago om spa se and geog aphi-
cally sca e ed ma e ial. Much o he second-
a y li e a u e commonly consul ed o spec-
imen iden i ica ion (e.g., keys, checklis s,
and ield guides) he e o e e lec s axonomic
con en ions ha ha e no been es ed by e-
cen analyses o cha ac e da a. Fo many
gene a, eliable iden i ica ions can only be
ob ained by consul ing he p ima y li e a u e
1998 7SIMMONS AND VOSS: PARACOU BATS
and by di ec ly compa ing ouche speci-
mens wi h ypes. Un o una ely, published
species lis s om ain o es mammal in en-
o y p ojec s a e seldom accompanied by
su icien documen a ion o judge he eli-
abili y o he iden i ica ions hey con ain.
To p o ide a baseline o mo e e ec i e
esea ch on Neo opical ain o es mammal
di e si y, we looked o a s udy si e whe e
we could e u n yea a e yea o ob ain e-
sul s om sus ained sampling e o using
he ull ange o me hods equi ed o axo-
nomically comp ehensi e su eys (Voss and
Emmons, 1996). We we e a ac ed o F ench
Guiana because o i s unique combina ion o
ex ensi e p ima y ain o es in close p ox-
imi y o mode n anspo a ion and commu-
nica ion ne wo ks. A Pa acou—a esea ch
a ea wi h p imi i e bu adequa e li ing qua -
e s in he mids o a la ge ac o o es only
a ew kilome e s by excellen oads om
pos o ices, elephones, ax machines, ma -
ke s, and hospi als—we ound an ideal lo-
ca ion o ou p ojec .
This is he i s o wo monog aphs based
on ou ieldwo k a Pa acou, which began in
1991 and con inued o 1994. He ein we de-
sc ibe he esea ch a ea, explain he me hods
used o sample he ba auna, and epo ou
ba -sampling esul s. We documen all axo-
nomic iden i ica ions by e iewing he ele-
an li e a u e, by p o iding ables o ex e -
nal and c anioden al measu emen s o ouch-
e specimens, and by de ailed compa isons
wi h ypes and o he ele an ma e ial in
cases whe e we encoun e ed signi ican p ob-
lems. We summa ize ield obse a ions o
each species by cap u e me hod and habi a ,
and we ema k no ewo hy ecological di e -
ences be ween congene s o o he closely e-
la ed axa. We analyze ou sampling esul s
o assess me hodological bias, habi a di e -
ences, empo al a ia ion, and comple eness.
We selec 13 o he Neo opical ain o es ba
in en o ies as compa a i e exempla s, and
we discuss hese oge he wi h he Pa acou
auna in e ms o axonomic composi ion,
biogeog aphy, species ichness, and ophic
s uc u e. Finally, we o e sugges ions o
u u e wo k based on he ou come o ou
ield and museum esea ch wi h Pa acou
ba s.
ACKNOWLEDGMENTS
We hank he adminis a i e s a o CIRAD
in F ench Guiana, especially V. Fa ichon,
O. Hamel, J.-F. Julia, and O. La oussinie, o
hei pe mission o wo k a Pa acou, o
space o s o e ou ield equipmen in Kou ou,
and o nume ous cou esies ha made ou
ieldwo k mo e p oduc i e and enjoyable.
We a e also g a e ul o G. Dubos o his ini-
ial in i a ion o wo k a Pa acou, and o his
ad ice and encou agemen h oughou he
cou se o ou ieldwo k. P.-M. Fo ge kindly
helped us ge s a ed in 1991.
Special hanks a e due o P. Pe onelli, a
gi ed ama eu na u alis and long- ime esi-
den a Pa acou, who gene ously sha ed his
special knowledge o he o es besides e-
pai ing almos e e y hing ha b oke o mal-
unc ioned a ound camp. Wi hou his help i
would ha e aken us much longe o lea n
he ail sys em, iden i y plan s, ind ba
oos s, and gene ally use ou ime e icien ly
in he ield. Wha e e measu e o success we
achie ed a Pa acou is in la ge pa owed o
him.
Fo hei assis ance wi h ba in en o y
ieldwo k we hank Da in Lunde, who was
always he e o help om 1991 o 1993; o
Suzanne Smi h, who compe en ly bo e he
b un o he mis ne ing e o in 1993; o Ro-
land Kays, who climbed ees and assis ed in
many o he ways in 1993; o And ea Pe ley,
who did almos e e y hing almos e e y day
in 1994; and o Nancy Voss, whose s alwa
assis ance helped make ou las ield season
he mos p oduc i e e e . Amy Be ko and
Louise Emmons also helped us ne ba s in
1993 and 1994, espec i ely.
Back a he museum we we e g a e ul o
he assis ance o Eleano S e ling who i s
se up he Pa acou da abase, o Da in Lun-
de and Bu on Ba ne who comple ed and
p oo ed i , and o Tenley Conway who pa-
ien ly made nume ous e isions and p in -
ou s as ou species iden i ica ions we e e-
ined. Da in Lunde, Eleano S e ling, and
And ea Pe ley ca aloged specimens, pulled
skulls, and cu a ed he collec ions. Pa
B unnaue esou ce ully acked down doz-
ens o e e ences in he AMNH lib a y, and
Don Clyde always ound he es some-
whe e else. Pe e Goldbe g skill ully p in -
8 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
ed all o he pho og aphs we ook in he
ield, and Pa Wynne did all o he splendid
a wo k.
Fo he loan o specimens o o hei hos-
pi ali y while we isi ed hei espec i e in-
s i u ions (abb e ia ed as below), we hank P.
D. Jenkins (BMNH); D. A. Schli e and S.
B. McLa en (CM); L. R. Heaney and B. D.
Pa e son (FMNH); L. G anjon and M. T an-
ie (MNHN); V. Pacheco (MUSM); M. D.
Engs om (ROM); and M. D. Ca le on, L. K.
Go don, and C. O. Handley, J . (USNM).
C i ical eedback on he i s d a o his
epo was p o ided by L. H. Emmons, M.
D. Engs om, E. K. V. Kalko, and B. D. Pa -
e son, all o whom ook ime om hei busy
esea ch schedules o help imp o e ou ex .
F anc¸ois Vuilleumie e y kindly ansla ed
ou abs ac .
Ou ield esea ch was ini ia ed wi h unds
om he Depa men o Mammalogy and he
esea ch adminis a ion o he Ame ican Mu-
seum o Na u al His o y. Funding o he las
wo ield seasons was p o ided by g an
numbe 4991–93 om he Na ional Geo-
g aphic Socie y. We hank bo h o ganiza ions
o hei gene ous suppo .
We dedica e his epo o ou la e iend and
colleague, Ka l F. Koopman, whose pa ience,
help, and encou agemen we e essen ial a
many poin s in ou museum esea ch. The sys-
ema ic accoun s would no ha e been he
same wi hou him, and we a e saddened ha
he did no li e o see hem comple ed.
SITE DESCRIPTION
TOPOGRAPHY, SOILS, AND
VEGETATION
The collec ions and obse a ions epo ed
below we e ob ained wi hin he Domaine Ex-
pe imen al Pa acou (he ea e , Pa acou), a la ge
ac o land adminis e ed as a esea ch con-
cession by he Cen e de Coope´ a ion In e -
na ionale en Reche che Ag onomique pou le
De´ eloppemen (CIRAD). F om a ield camp
a 5
8
16
9
31
0
N, 52
8
55
9
25
0
W (app oxima ely 12
km SSE o Sinnama y and 33 km WNW o
Kou ou; igs. 1, 2), we wo ked in a oughly
ci cula a ea wi h a adius o abou 3 km.
1
The
local e ain consis s o low, densely o es ed
hills d ained by he headwa e s o ou s eams:
C ique Moges e n and C ique Ve lo (bo h ib-
u a ies o he lowe Sinnama y Ri e ), and
C ique Pa akou and C ique Malmanou y
(which low sepa a ely in o he A lan ic
Ocean). The minimum ele a ion abo e sea
le el wi hin ou s udy a ea is abou 4 m, he
maximum abou 45 m (IGN, 1991).
1
We es ima ed he coo dina es o ou camp as he
mean o ou eadings ob ained in 1994 wi h a Panasonic
KX-G5500 GPS ecei e using he WGS84 (s anda d)
map da um. A ew ba s collec ed unde old highway
b idges c ossing C ique Renne (ca. 7.2 km o he NE)
and C ique Pa acou (ca. 7.1 km N) a e epo ed among
ou ouche ma e ial, bu all o hese belong o species
ound wi hin ou 3-km sampling adius.
Local soils a e he esul o in si u wea h-
e ing om qua zi es and schis s o he Bon-
ido o se ies (Ba i eau, 1993; Du ieu de
Mad on, 1993). O e mos o ou s udy a ea,
he subs a e is an acidic b ownish o eddish
sandy clay (mixed wi h i ons one conc e ions
on some idge ops and hillsides), bu a ew
pa ches o almos pu e whi e sand also occu
(Ba hes, 1991). Al hough hea ily wea he ed
qua zi e boulde s a e occasionally exposed
in s eams, he e a e no la ge ocky ou c ops
o ca es in he egion. Local wa e cou ses
a e all small (mos ly 5 m o less in wid h),
shallow (usually 1 m o less in dep h), and
hea ily shaded, wi h anspa en , ea-colo ed
(‘‘black’’) wa e lowing slowly o e p e-
dominan ly sandy beds.
Mos o his landscape is co e ed wi h p i-
ma y ain o es
2
( ig. 3), bu he new asphal
highway om Kou ou o Sinnama y cu s
2
No mac oscopic soil cha coal was obse ed when D.
P. Lunde and R. W. Kays dug 55 pi s, each abou 40 cm
deep and 30 cm in diame e , o ins all pi all aps in 1993
(Voss e al., in p ep.). We likewise saw no cha coal in he
spoil om nume ous soil sample co es made in a 25-ha
plo (Pa cel 16) by a o es y esea ch class in 1994. Soil
cha coal has been in e p e ed as e idence o p ehis o ic ag-
icul u e a o he Neo opical ain o es locali ies (e.g., by
Lo ejoy and Bie egaa d, 1990; McDade and Ha sho n,
1994), and i s absence sugges s ha he ma u e o es a
Pa acou has no been clea ed o cen u ies.
1998 15SIMMONS AND VOSS: PARACOU BATS
The e o e, we eely changed ou equipmen
and modi ied ou p ocedu es om season o
season (some imes om day o day) as sug-
ges ed by ou own accumula ing esul s,
p omp ed by ecommenda ions om ou col-
leagues, o demanded by changing clima ic
and phenological ci cums ances in he ield.
MISTNETTING
A he begining o ou su ey (in 1991)
we used s anda d-weigh 70-denie (70 d)
mis ne s o ca ch ba s. We la e (1992)
swi ched o ligh -weigh 30- and 50-denie
(30 d and 50 d) ne s and used hem almos
exclusi ely he ea e . The ne s we used a
Pa acou we e 2.6 m high and came in 6-, 13-,
and 18-m leng hs; he size o he nylon mesh
was 36 mm.
G
ROUND
-L
EVEL
N
ETTING
: To ca ch ba s a
g ound le el, 0–3 m by ou con en ion, we
moun ed ne s on poles (usually made om
saplings cu and immed in si u) h us in o
he soil and b aced by guys o b aided nylon
co d ( ig. 7). To minimize dis u bance o he
unde s o y ege a ion ha migh ala m ba s
pa olling amilia e i o y, we usually ied
o se ne s along p eexis ing ails. E en so,
some clea ing was usually necessa y o a oid
en angling ne s in ailside ege a ion, and o
lea e enough space o people o wo k on
each side. Ligh weigh (30 d) ne s a e so
agile and ha d o clean o en angled li e
ha i is gene ally also necessa y o sweep
he g ound clean o lea es and wigs unde
each ne .
As a gene al ule, ne s mus be mo ed e -
e y nigh because ba s quickly lea n o a oid
hem. Ne ing e ec i ely on a nigh ly sched-
ule he e o e equi ed a leas wo pe sons:
one o s ay in camp and p ocess he las
nigh ’s ca ch, and he o he o su ey he o -
es o new ne loca ions, clea new ne lanes,
and se up o he nex nigh ’s wo k. We
chose ne loca ions o sample as many o es
mic ohabi a s as possible: well-d ained si es,
palm swamps, s eambeds, unusual plan as-
socia ions, e c.
We o en opened ne s sho ly be o e dusk
(when i was s ill ligh enough o ead easily)
because some ba s eme ge om hei oos s
in he la e a e noon. Once he ne s we e
opened we s ayed wi h hem cons an ly be-
cause la ge ba s chew holes in ne s unless
hey a e p omp ly emo ed, and because
small ba s o en escape om ne s unless hey
a e immedia ely seized. We egula ly kep
ne s open om dusk o dawn in 1991 and
1992, bu ne ing a e 22:00 hou s was sel-
dom p oduc i e, so we usually closed ou
ne s be o e midnigh in subsequen yea s.
In addi ion o passi ely wai ing o ba s o
ly in o ou ne s, we used he Audubon Bi d
Call ( ig. 8) o a ac hem. This simple de-
ice p oduces high-pi ched (bu no ul ason-
ic) squeaks esembling (among o he hings)
he dis ess calls o s enode ma ines, which
some imes lock o he sound in la ge num-
be s (Handley e al., 1991). O g ea e in e -
es o in en o y wo k, some phyllos omines
ha a e o he wise ha d o cap u e (e.g.,
Ch o op e us au i us,Glyphonyc e is da ie-
si,Tona ia schulzi,Vampy um spec um) a e
likewise a ac ed, pe haps because hey a e
in e es ed in dis essed ba s, singing o hop-
e ans (Tu le e al., 1985), o o he small an-
imals wi h high-pi ched ocaliza ions as
p ey. We i s used Audubon Bi d Calls spa -
ingly in 1992, bu in la e yea s we equen -
ly used hem when bo ed by he absence o
ba ac i i y a ound ou ne s.
Ba s we e emo ed om ne s as soon as
possible a e cap u e and placed in clo h
bags closed by d aws ings. When oo many
ba s we e cap u ed o place each in i s own
bag, we o en pu wo o mo e conspeci ics
oge he , bu we ne e knowingly mixed spe-
cies in bags. In gene al, we bagged all cap-
u ed ba s whe he o no hey we e equi ed
as specimens in o de o p e en ecap u es
on he same nigh ; a e he ne s we e closed,
unwan ed ba s we e checked o co ec iden-
i ica ion and sex, and hen eleased. Bags
con aining cap u ed ba s we e always sus-
pended om co ds ied ho izon ally be ween
nea by ees o keep hem away om p owl-
ing opossums.
E
LEVATED
N
ETTING
: To ca ch ba s in he
subcanopy ( ig. 9) and in he canopy i sel
( ig. 10), we moun ed ou ne s on slende
poles, he sawn-o op and bo om ends o
which we e w apped wi h duc ape ( o p e-
en spli ing) and i ed wi h small eyebol s.
Swi el-snaps we e used o a ach he poles
o a leng h o
1
⁄
8
-inch (ca. 3 mm) b aided ny-
lon line ha o med a con inuous unning

16 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 7. G ound-le el mis ne s ( u led in his day ime pho og aph) in swampy p ima y o es ; po -
ions o h ee ne s (a ows) suppo ed by guyed poles a e shown. We caugh 30 species o ba s in 11
nigh s o g ound-le el ne ing a his si e om 1991 o 1994, including ou only examples o Glypho-
nyc e is da iesi. The mos commonly cap u ed species he e we e Ca ollia pe spicilla a,A ibeus ja-
maicensis,A.obscu us,Phyllos omus elonga us,Rhinophylla pumilio,Tona ia sau ophila, and T achops
ci hosus.
1998 17SIMMONS AND VOSS: PARACOU BATS
Fig. 8. The Audubon Bi d Call (a ailable
om R. W. Eddy, Box 0172, Newing on CT
06131) consis s o a pew e key in a hollow
wooden cylinde ; wis ing he key p oduces high-
pi ched squeaks ha can be used o a ac ba s o
ne s. Handley e al. (1991: ig. 12-1) p o ided a
spec og aphic analysis o he sounds p oduced by
his use ul de ice, all o which a e in he audible
ange.
loop by means o which he ne could be
aised o lowe ed ( lagpole ashion) a each
end. The unning loops we e h eaded
h ough o e head suppo s (eyebol s o b ass
ings), he posi ions o which de e mined he
heigh o which ne s could be aised.
When ees on ei he side o a sui able
canopy gap o lyway we e close oge he
(wi hou enough oom be ween hei c owns
o suspend a ne ) and no oo la ge (
#
35 cm
diame e ), we climbed hem using F ench
ee-climbing spikes (Model 502, a ailable
om ETS Lacos e, 24160 Excideuil, F ance;
see Mo i, 1987: ig. I-3A) and sc ewed la ge
eyebol s di ec ly in o he unks. To ix sup-
po s ac oss la ge gaps, we used he gian
slingsho designed by Munn (1991) o shoo
lead ishing weigh s ied o hea y mono ila-
men line (on a spin-cas ing eel) in o o o e
he c owns o ees on ei he side.
4
B ass
ings ( h ough which he unning loops we e
i s h eaded) we e hen d awn up on b aided
line; ou simples ne igs used a single op-
line o hold he ings in place.
In addi ion o ha ing sui able o e head
suppo s on each side o a canopy gap, ele-
a ed ne si es mus be clea enough a
4
D. P. Lunde and R. W. Kays cons uc ed and es ed
ou line-shoo ing equipmen and helped ins all he i s
canopy ne s a Pa acou.
g ound le el ha ne s can be aised and low-
e ed wi hou obs uc ion. Un o una ely,
mos na u al gaps in he canopy a e ee alls,
which a e always li e ed wi h p os a e
unks and limbs. Roads and clea ings p o-
ide canopy gaps ha a e gene ally ee o
such obs acles ( ig. 11), bu manmade open-
ings in he o es a e o en bo de ed by
young ees ha a e no all enough o sup-
po high ne s. These cons ain s, oge he
wi h he labo equi ed o ig ele a ed ne s,
p ecluded equen si e changes in sampling
he canopy ba auna a Pa acou.
We measu ed he app oxima e heigh o el-
e a ed ne s by a aching a ape measu e o
he bo om o one o he ne poles; cap u e
heigh was hen eco ded as he 3-m in e al
om he bo om o he op o he ne (i.e.,
10–13 m o ba s caugh in a ne wi h low-
e mos pole ends 10 m abo e he g ound).
Howe e , we ound i essen ial o keep a
sha p eye on ele a ed ne s as hey we e low-
e ed o emo e en angled ba s because low-
lying species we e o en cap u ed in he p o-
cess; cap u e heigh da a om ca elessly
moni o ed ne s could he e o e o e es ima e
he e ical ac i i y ange o unde s o y spe-
cies.
SEARCHING FOR ROOSTS
Many ain o es ba s ha a e ha d o ca ch
in mis ne s can easily be collec ed o ob-
se ed a hei diu nal oos s. We ound some
oos s by chance, when ba s lew up sudden-
ly as we passed an unsuspec ed e ea in he
o es unde s o y. In such cases, we s epped
a ew me e s away and s ood quie ly un il he
animals e u ned (usually in jus a ew min-
u es). Mos oos s, howe e , we e disco e ed
by delibe a e sea ching along ails. Among
o he po en ial oos si es in p ima y and sec-
onda y o es , we sea ched benea h unde cu
s eambanks, unde he bu esses o allen
ees, inside hollow logs, inside hollow
s anding ees, unde loose ba k on s anding
ees, and in oliage (especially among he
lea es o la ge monoco s such as palms, Hel-
iconia spp., Phenakospe mum guyannense,
Philodend on spp.). Al hough we some imes
made special e o s o ind pa icula oos
ypes (e.g., lea en s o hollow ees), we
ne e igno ed o he kinds o oos s encoun-
18 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 9. Subcanopy o well-d ained p ima y o es a Pa acou, he e iewed om abou 20 m abo e
he g ound (pho og aph by R. W. Kays). The subcanopy is much less clu e ed by lea y ege a ion han
he unde g ow h, which may explain he highe cap u e a es o ae ial insec i o es like Co mu a b e-
i os is,Pe op e yx kapple i,Saccop e yx bilinea a, and S.lep u a a his le el. A leas 16 species o
phyllos omids a e p obably also ac i e in his habi a , al hough mos o ou ele a ed ne cap u es o
hem we e o e oads, no inside he o es .
e ed as we sea ched. In addi ion o looking
o na u al oos s, we in es iga ed epo s o
ba s oos ing in houses and s o age sheds,
and we sea ched e e y cul e unde oads
wi hin ou sampling adius.
Mos ba s we e loca ed isually in hei di-
u nal e ea s, and some wi h dis inc i e pos-
u e, colo a ion, o o he ex e nal cha ac e s
could be eliably iden i ied wi hou cap u e.
Likely pa ches o subcanopy oliage we e
scanned wi h binocula s, which we e also use-
ul o examining well-li oos s in he o es
unde s o y ha could no be closely ap-
p oached wi hou igh ening he inhabi an s.
Roos s unde allen ees, inside hollow logs,
and o he da k e ugia we e examined wi h a
headligh (which lea es he hands ee o push
aside obs uc ing ege a ion, wield a ne , aim
a gun, o ake no es). Sho -handled bu e ly
ne s we e some imes use ul o cap u ing
oos ing ba s, bu in many si ua ions shoo ing
(wi h numbe 12 sho in .22 calibe o .410
loads) was he only op ion.
Occasionally, oos ing ba s could be hea d
lu e ing inside ee holes, hollow logs, o
o he places whe e hey could no be seen o
eached. In such si ua ions, we plugged all
bu one en ance, a ound which we buil a
mis ne enclosu e o ca ch he animals as
hey eme ged a dusk. We used sho (6 m)
70-d mis ne s w apped a ound up igh poles
o enclose he oos opening on all sides, and
hen ha ched he enclosu e wi h palm onds
o p e en eme ging ba s om escaping up-
wa d. Such enclosu es usually cap u ed e -
e y eme ging ba .
To ob ain da a on oos occupancy in suc-
cessi e yea s, we assigned e e y oos a
unique numbe ; du able oos s ( hose ha
could be expec ed o las o mo e han a ew
mon hs, e.g., hollow logs, ee ca i ies) we e
iden i ied by nailed aluminum ags. We ied
1998 19SIMMONS AND VOSS: PARACOU BATS
Fig. 10. View o e he o es canopy om he c own o an eme gen ee, abou 45 m abo e he
g ound (pho og aph by R. W. Kays). The unobs uc ed ai space abo e he ees is p obably used mos ly
by species o Diclidu us,Ep esicus,Lasiu us, and molossids, al hough mos o ou cap u es o hese
axa we e made close o he g ound in ne s suspended ac oss a oad h ough he o es o o e oadside
puddles. Many phyllos omids a e p obably ac i e in he canopy i sel , al hough we cap u ed only Phyl-
los omus has a us,Ame ida cen u io, and Chi ode ma illosum in ou highes ne s.
o e is each du able oos e e y subsequen
ield season and eco d he species and num-
be s o ba s in esidence, bu some oos s we e
des oyed when new land was clea ed o ag-
icul u al expe imen s and o he s could no be
ound again. Some oliage oos s we e e is-
i ed se e al imes pe ield season ollowing
emo al o he o iginal occupan s o de e mine
whe he lea - en s used by one species a e e e
subsequen ly colonized by ano he .
To explo e he e ec s o obse e bias in
sea ching o oos s, and o ob ain da a on
oos densi y, we sys ema ically su eyed a
25-ha plo o p ima y o es (Pa cel 16 o
CIRAD’s o es y esea ch p og am), which
was con enien ly di ided by a squa e g id o
su eyed ails in o 1-ha quad a s ( ig. 12).
Fou sea che s wi h widely a ying p io ex-
pe ience we e assigned quad a s by lo e y in
1994. Each 1-ha quad a was hen su eyed
o oos s by one sea che who i s walked
he pe ime e ails ( o aling 400 m) and hen
he ailless in e io ; he in e io was sea ched
in ou pa allel ansec s spaced 20 m apa .
Thus, no poin wi hin any quad a was mo e
han 10 m om a sea ched pa h o ansec .
Sea che s we e ins uc ed o in es iga e all
likely oos ing si es and o lag all disco e ed
oos s so ha hey could be subsequen ly e-
isi ed and con i med by he mos expe i-
enced sea che . Because Pa cel 16 was se
aside o nondes uc i e esea ch, i was no
possible o de e mine he iden i ica ion o all
oos ing ba s ound in his su ey.
SPECIMEN COLLECTION AND
PRESERVATION
We p ese ed ouche specimens using
s anda d p ocedu es desc ibed and illus a ed
by Handley (1988). In collec ing specimens,
ou objec i e was o p ese e enough ex-
amples o each species o assess he axo-
nomic s a us o he Pa acou popula ion wi h
20 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 11. A na ow di oad ac oss which we aised ne s o ca ch high- lying ba s. The oad is
bo de ed by young seconda y g ow h and selec i ely logged o es , bu p ima y o es occu s wi hin
abou 50–100 m on ei he side.

1998 21SIMMONS AND VOSS: PARACOU BATS
Fig. 12. Map o a 25-ha plo o p ima y o es (Pa cel 16) di ided by a g id o su eyed ails in o
1-ha quad a s (le ) and he plan used o sea ch each 1-ha quad a o ba oos s ( igh ). Two ails on
he eas side p o ided access om a di oad ca. 0.5 km away. Mos o he su eyed habi a was well-
d ained o es , bu quad a s 5, 11, 12, 16, 17, 21, 22, and 25 also con ained swampy o es . The s anda d
sea ch p ocedu e o each quad a is explained in he ex .
espec o popula ions sampled a o he lo-
cali ies. Usually, we limi ed ou emo al o
abou 20 indi iduals (10 males and 10 e-
males) pe species, enough o ob ain mean-
ing ul es ima es o he ange o a ia ion and
cen al endency in each sex o cha ac e s o
axonomic in e es . Fo open popula ions o
mos ain o es ba s, des uc i e sampling on
his scale is p obably i ial by compa ison
wi h na u al demog aphic p ocesses.
5
5
Leigh and Handley (1991) es ima ed a popula ion den-
si y o abou 200 A ibeus jamaicensis pe squa e kilome e
o lowland o es in cen al Panama. Because hei s udy
si e was ich in ig ees, he p ima y ood esou ce o A.
jamaicensis, his es ima e is pe haps on he high side o
he species. Howe e , A.jamaicensis is la ge (45 g) han
mos Neo opical ain o es ba s, whose calo ic equi e-
men s can p obably be ob ained in smalle home anges.
On he assump ion ha 100 indi iduals/km
2
is a conse -
a i e a e age alue o many s enode ma ines, we could
expec abou 2800 indi iduals o each species wi hin ou
s udy a ea (ca. 28 km
2
). An annual su i o ship o 64%
( he highes igu e sugges ed by Leigh and Handley o A.
jamaicensis) would hen imply abou 1000 na u al dea hs
pe species pe yea a Pa acou. P esumably, such mo ali y
is annually compensa ed by ec ui men in all species ha
main ain s able local popula ions.
ACOUSTIC SURVEY
In he las yea o ou wo k a Pa acou
(1994), we used a cus om-made elec onic
ba de ec o bo owed om Elisabe h K. V.
Kalko (Uni e si y o Tuebingen, Ge many,
and Smi hsonian T opical Resea ch Insi u e,
Panama) o eco d he echoloca ion calls o
ba s o aging along oads and in manmade
and na u al clea ings. The signals we e am-
pli ied, ed in o a ansien eco de , and ead
ou a
1
⁄
15
educed speed on a Sony Walkman
P o essional casse e ape eco de . We made
198 eco dings o e he cou se o 10 nigh s
be ween 9 Oc obe and 30 Oc obe 1994.
Analyses o hese eco dings a e cu en ly in
p og ess, and he esul s will be p esen ed
elsewhe e.
SCHEDULING AND PERSONNEL
We wo ked a Pa acou om 1991 o 1994,
wi h he da es, pe sonnel, and me hods lis ed
below. Al hough all pe sonnel helped o
some ex en wi h he ba in en o y, some
we e p ima ily engaged in o he ac i i ies;
only hose chie ly esponsible o he lis ed
22 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
me hods a e iden i ied pa en he ically. I alics
indica e he au ho s o ieldno es (deposi ed
in he AMNH Depa men o Mammalogy
a chi es) ha should be consul ed o de-
ailed in o ma ion abou ele an me hods.
1991: 26 June–27 Augus ; Da in P.
Lunde, Nancy B. Simmons, and Robe S.
Voss; g ound-le el ne ing (Simmons) and
sea ching o oos s (Voss).
1992: 20 Oc obe –18 No embe ; Da in
P. Lunde, Nancy B. Simmons, and Robe S.
Voss; g ound-le el ne ing (Simmons, Voss)
and sea ching o oos s (Voss).
1993: 20 July–15 Sep embe ; Roland W.
Kays, Da in P. Lunde, Suzanne E. Smi h,
and Robe S. Voss; g ound-le el and ele-
a ed ne ing (Smi h, Voss) and sea ching o
oos s (Voss).
1994: 19 Sep embe –13 No embe ;
Louise H. Emmons, And ea L. Pe ley, Nan-
cy B. Simmons, Nancy A. Voss, and Robe
S. Voss; g ound-le el and ele a ed ne ing
(Pe ley, Simmons, N. A. Voss, R. S. Voss),
sea ching o oos s (Pe ley, Simmons, N. A.
Voss, R.S.Voss), and ul asonic census (Pe -
ley, Simmons).
SYSTEMATIC ACCOUNTS
The ollowing accoun s a e a anged sys-
ema ically by amily (and by sub amily
wi hin Phyllos omidae), and hen alphabe i-
cally by genus and species. Unless no ed o h-
e wise, he nomencla u e and axonomic se-
quence used he ein ollows Koopman
(1993). The o ganiza ion o mos accoun s
employs se e al s anda d subheadings.
V
OUCHER
M
ATERIAL
: This sec ion sum-
ma izes in o ma ion abou specimens col-
lec ed, which a e lis ed by sex and museum
ca alog numbe . Indi iduals measu ed o a-
bles a e indica ed wi h as e isks.
I
DENTIFICATION
: In he ield, we made ex-
ensi e use o Emmons’ (1990) gene ally ex-
cellen desc ip ions o ex e nal cha ac e s o
make p elimina y iden i ica ions. Al hough
subsequen museum s udy o ouche speci-
mens con i med mos o ou ield de e mi-
na ions, we also encoun e ed nume ous ax-
onomic p oblems ha had o be esol ed o
jus i y he nomencla u e adop ed below.
In cases whe e iden i ica ion was ela i ely
s aigh o wa d, we simply lis he echnical
e e ences we consul ed a he beginning o
each accoun and ema k any no ewo hy
poin s in which ou ma e ial di e s om
published desc ip ions. Species iden i ica-
ions in ol ing signi ican axonomic p ob-
lems a e ea ed a leng h, wi h e iews o
he ele an li e a u e and analyses o cha -
ac e a ia ion in he specimens examined.
We also discuss cu en ly ecognized subspe-
cies in his sec ion, al hough we do no use
inomial headings below.
Measu emen s o ouche specimens and
compa a i e ma e ial a e p o ided in ables
accompanying each species accoun . These
mo phome ic summa ies include only mea-
su emen s o adul s wi h closed epiphyses.
Linea measu emen s o ex e nal and c anio-
den al dimensions a e epo ed in millime e s
(mm); weigh s a e epo ed in g ams (g). The
ollowing measu emen s we e aken o all o
some o he species ea ed below.
To al leng h: Dis ance om he ip o he snou
o he ip o he las caudal e eb a.
Tail leng h: Measu ed om he poin o do sal
lexu e o he ail wi h he sac um o he ip o
he las caudal e eb a.
Hind oo leng h: F om he an e io edge o he
base o he calca o he ip o he claw o he
longes oe.
Ea leng h: F om he no ch o he leshy ip o
he pinna.
Fo ea m leng h: F om he elbow ( ip o he olec-
anon p ocess) o he w is (including he ca -
pals). This measu emen is made wi h he wing
a leas pa ially olded.
Tibia leng h: F om he p oximal end o he ibia
o he pos e io base o he calca .
Thumb leng h: F om he me aca pal-phalangeal
join o he ip o he claw.
G ea es leng h o skull: F om he pos e io mos
poin on he occipu o he an e io mos poin
on he p emaxillae (excluding he inciso s).
Condyloincisi e leng h: F om he pos e io mos
poin on he occipi al condyles o he an e i-
o mos poin on he uppe inciso s.
Condylocanine leng h: F om he pos e io mos
poin on he occipi al condyles o he an e i-
o mos poin on he uppe canines.
Lac imal b ead h: G ea es b ead h ac oss he lac-
1998 23SIMMONS AND VOSS: PARACOU BATS
imal (
5
an eo bi al) idges, when p esen and
well de ined.
Pos o bi al b ead h: Leas b ead h ac oss he
on als pos e io o he pos o bi al p ocesses o
bulges.
Zygoma ic b ead h: G ea es b ead h ac oss he
zygoma ic a ches.
B aincase b ead h: G ea es b ead h o he glob-
ula pa o he b aincase.
Mas oid b ead h: G ea es c anial b ead h ac oss
he mas oid egion.
Maxilla y oo h ow leng h: F om he an e io mos
edge o he canine c own o he pos e io mos
edge o he c own o M3.
B ead h ac oss mola s: G ea es b ead h ac oss
he ou e edges o he c owns o he uppe mo-
la s.
B ead h ac oss canines: G ea es b ead h ac oss
he ou e edges o he c owns o he uppe ca-
nines.
To con i m iden i ica ions o p oblema ic
species, we compa ed ou ouche ma e ial
o museum specimens om o he locali ies,
and we o en consul ed li e a u e desc ip ions
o ypes ha we did no pe sonally examine.
We iden i y he museums in which ouche s
and o he specimens a e p ese ed by he ol-
lowing abb e ia ions: AMNH, Ame ican
Museum o Na u al His o y (New Yo k);
BMNH, B i ish Museum o Na u al His o y
(London); CM, Ca negie Museum o Na u al
His o y (Pi sbu gh); FMNH, Field Museum
o Na u al His o y (Chicago); MNHN, Mu-
se´um Na ional d’His oi e Na u elle (Pa is);
MUSM, Museo de His o ia Na u al de la
Uni e sidad Nacional Mayo de San Ma cos
(Lima); RMNH, Rijksmuseum an Na uu -
lijke His o ie (Leiden); SMN, S aa liches
Museum u¨ Na u kunde (S u ga ); USNM,
Na ional Museum o Na u al His o y (Wash-
ing on, D.C.); ZMH, Zoologisches Museum
Hambu g (Hambu g).
F
IELD
O
BSERVATIONS
: This sec ion sum-
ma izes ou cap u e eco ds and oos obse -
a ions, and includes discussion o habi a s,
beha io , and o he na u al his o y opics. In
epo ing ou obse a ions o each species,
we i s lis cap u e equencies by in en o y
me hod, and hen p o ide a b eakdown o
cap u e equencies by habi a o each me h-
od. I e e y eco ded cap u e is known o
ep esen a unique indi idual, we s a e he
numbe o indi iduals cap u ed. Al hough we
did no ma k eleased ba s, he numbe o
indi iduals known o ha e been cap u ed is
occasionally la ge han he numbe o
ouche s we p ese ed because only one in-
di idual o each sex was eleased o some
species. I mo e han one indi idual o ei he
sex we e eleased, we s a e he numbe o
cap u es eco ded and no e ha hese ‘‘pos-
sibly’’ o ‘‘p obably’’ (a subjec i e judg-
men ) include some ecap u es. Fo each spe-
cies caugh in ele a ed ne s, we summa ize
cap u e heigh eco ds as he in e al om
he bo om o he lowes ele a ed ne in
which he species was aken o he op o he
highes ne (e.g., 10–21 m o a species aken
once in a ne suspended 10–13 m abo e he
g ound and again in a ne 18–21 m abo e he
g ound). We also no e any conspicuous di -
e ences in mis ne cap u e-habi a o cap-
u e-heigh equencies be ween congene s o
o he closely ela ed species. We nex p o-
ide in o ma ion abou oos ing habi s, es-
pecially oos ype (hollow log, ee ca i y,
lea - en , e c.), oos ing habi a , numbe s o
indi iduals in oos ing g oups obse ed, and
he age and sex o cap u ed oos ing-g oup
membe s.
In epo ing habi a da a o each species,
we use he ollowing ca ego ies: (1) well-
d ained p ima y o es , (2) swampy p ima y
o es , (3) c eekside p ima y o es , (4) ee-
all openings in p ima y o es , (5) manmade
clea ings ( oadways, ga dens, plan a ions,
e c.), (6) closed-canopy seconda y g ow h
(including selec i ely logged o es ), and (7)
oadside puddles. The e m ‘‘p ima y o es ’’
(wi hou modi ie s) includes ca ego ies 1–4,
whe eas ca ego ies 5–7 a e o en e e ed o
collec i ely as ‘‘modi ied habi a s.’’
EMBALLONURIDAE
We cap u ed 10 emballonu id species a
Pa acou, h ee o which a e he e epo ed o
he i s ime om F ench Guiana. The spe-
cies we iden i ied om mo phological
ouche ma e ial ep esen he gene a Cen-
onyc e is,Co mu a,Diclidu us,Pe op e yx
(including Pe onymus), Rhynchonyc e is,
and Saccop e yx. Published eco ds om
elsewhe e in F ench Guiana and Su inam in-
clude ano he ou emballonu id species ha
migh also occu in ou s udy a ea (appendix
1).
24 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Cen onyc e is maximiliani (Fische )
Figu e 13
V
OUCHER
M
ATERIAL
: 1 male (AMNH
267397); see able 1 o measu emen s.
I
DENTIFICATION
: P e iously ega ded as
mono ypic (e.g., by Koopman, 1993), he ge-
nus Cen onyc e is was ecen ly shown o
con ain a leas wo species by Simmons and
Handley (1998). Acco ding o hese au ho s,
he known ange o Cen onyc e is maximi-
liani includes sou he n Venezuela, he Guia-
nas, and no he n, eas e n, and sou heas e n
B azil, whe eas C.cen alis Thomas (1912b)
occu s in Cen al Ame ica, Colombia, Pe u,
and on bo h sides o he Andes in Ecuado .
Bo h species a e known om only a ew
specimens, howe e , and hei ue geog aph-
ic anges may be conside ably la ge . In ac ,
because no known biogeog aphic ba ie s
sepa a e he anges o C.maximiliani and C.
cen alis, hese ba s migh occu sympa i-
cally almos anywhe e in Amazonia. Ou
Pa acou ouche is he i s known specimen
o C.maximiliani om F ench Guiana.
Measu emen s and desc ip ions o Cen-
onyc e is maximiliani om he Guianas and
elsewhe e we e p o ided by Thomas (1913),
Husson (1962, 1978), Williams e al. (1983),
and Simmons and Handley (1998). The la e
au ho s summa ized c anioden al cha ac e s
and measu emen s ha dis inguish C.maxi-
miliani om C.cen alis. Ou Pa acou
ouche o med much o he basis o hei
desc ip ion o C.maximiliani.
F
IELD
O
BSERVATIONS
: Ou single indi id-
ual o his species was sho in he la e a e -
noon (abou 16:00 hou s) as i pe ched abou
3 m abo e he g ound on he unde side o a
la ge (ca. 20
3
30 cm) unmodi ied lea o a
small melas oma aceous ee beside a ail in
well-d ained p ima y o es .
Co mu a b e i os is (Wagne )
V
OUCHER
M
ATERIAL
: 17 emales (AMNH
*266009, *267070, 267076, *267078,
*267389, *267391, *267823, *267824,
*267825, *267826, *267828; MNHN
*1995.823, *1995.824, *1995.825, *1995.826,
*1995.833, *1995.834) and 18 males
(AMNH *265994, *266002, *266003,
*267069, *267071, *267074, *267075,
267079, *267390, *267394, *267822,
*267827; MNHN *1995.827, *1995.828,
1995.829, *1995.830, *1995.831, *1995.832);
see able 1 o measu emen s.
I
DENTIFICATION
:Co mu a b e i os is can
be dis inguished easily om o he emballon-
u ids by he unique loca ion o he an eb a-
chial wing sac (Sanbo n, 1937: ig. 37; Jones
and Hood, 1993: ig. 1). C anioden al cha -
ac e s and measu emen s o C.b e i os is
om he Guianas and o he pa s o Sou h
Ame ica we e discussed by Sanbo n (1937),
Husson (1962, 1978), B osse and Cha les-
Dominique (1990), and Jones and Hood
(1993). Husson (1962) p o ided a e ised
desc ip ion o Co mu a and a de ailed dis-
cussion o Myop e yx pullus, which he and
all subsequen au ho s ha e conside ed o be
a junio synonym o C.b e i os is. No sub-
species a e cu en ly ecognized (Jones and
Hood, 1993; Koopman, 1994).
Ou specimens om Pa acou con o m o
p e ious desc ip ions o Co mu a b e i os-
is and all wi hin he ange o size a ia ion
p e iously documen ed in Su inam and
F ench Guiana.
F
IELD
O
BSERVATIONS
: We eco ded 42 cap-
u es (possibly including some ecap u es) o
Co mu a b e i os is a Pa acou: 7 in
g ound-le el mis ne s, 12 in ele a ed mis -
ne s, and 23 a oos s. O he se en g ound-
le el mis ne cap u es, one was in well-
d ained p ima y o es , h ee we e in c eek-
side p ima y o es , wo we e in a ee all
opening in c eekside p ima y o es , and one
was in a manmade clea ing. Eigh ele a ed-
ne cap u es we e be ween 10 and 21 m
abo e a na ow di oad, and ou we e
made be ween 10 and 38 m o e a ee all
gap in well-d ained p ima y o es .
We encoun e ed 10 di e en oos ing
g oups o Co mu a b e i os is a eigh
unique oos si es (one si e was e isi ed
wice). Fi e oos si es we e on he da k un-
de sides o allen ees (e.g., ig. 14), bu one
oos ing g oup was ound clinging o an un-
modi ied lea o Phenakospe mum guyan-
nensis ( ig. 15), ano he occupied a shallow
ca i y in he base o a li ing ee suspended
by i s oo s o e a s eam, and ano he was
ound unde a conc e e b idge. In addi ion o
he b idge oos , wo na u al oos s (bo h in
p ima y o es ) we e o e small s eams, bu
1998 31SIMMONS AND VOSS: PARACOU BATS
Fig. 16. Roos successi ely occupied by h ee species o emballonu ids in well-d ained p ima y
o es a Pa acou. The da k conca e unde side o his b oken unk (a ow) shel e ed oos ing g oups o
h ee Pe op e yx mac o is in 1991, ou P.kapple i in 1992, and h ee Co mu a b e i os is in 1994;
he oos was ound acan in 1993.

32 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 17. Roos o Pe op e yx leucop e a and Fu ip e us ho ens in well-d ained p ima y o es a
Pa acou (some unde s o y ege a ion was selec i ely emo ed o pho og aphy). Two indi iduals o P.
leucop e a we e ound oos ing in a ho izon al chambe (a ow) be ween wo bu esses o his allen
ee in 1992, and one F.ho ens occupied a simila chambe on he o he side a he same ime. This
oos was acan in 1993 and was des oyed by a ee all in 1994. Deep ecesses be ween he bu esses
o allen ees like his p o ide oos s o se e al ba species besides he wo ound he e, including
Pe op e yx kapple i,Choe oniscus mino , and Lonchophylla homasi. Howe e , whe eas emballonu ids
usually occupy dimly ligh ed si es be ween la e al bu esses, glossophagine oos s a e always on he
da ke unde side (whe e hey can be seen only by c awling unde nea h such ees wi h a headligh ).
1998 33SIMMONS AND VOSS: PARACOU BATS
Fig. 18. Roos o Pe op e yx leucop e a a he ma gin o an expe imen al plo in well-d ained o es
a Pa acou. Felled by cu s h ough he bu esses abou 1 m abo e he g ound, his ee was he only
oos o P.leucop e a ha we ound unobs uc ed by dense unde s o y ege a ion. Se en o eigh ba s
we e obse ed oos ing in he ca e-like ecess (a ow) o med by he s anding base and he allen unk.
mu a b e i os is in 1992, and we once
ound Fu ip e us ho ens simul aneously
oos ing benea h he same allen ee as P.
leucop e a, bu in a di e en chambe sepa-
a ed by la ge bu esses. O he wise, his spe-
cies was no obse ed o sha e oos si es
wi h o he ba s.
On se e al occasions we saw ba s wi h
whi e wings, almos ce ainly Pe op e yx
leucop e a, lying egula bea s 1–2 m abo e
he g ound a dusk in ou camp clea ing.
Pe op e yx mac o is (Wagne )
Figu es 19, 20
V
OUCHER
M
ATERIAL
: 2 emales (AMNH
*266006, *267396) and 3 males (AMNH
*266005, *266007; MNHN *1995.843); see
able 4 o measu emen s.
I
DENTIFICATION
: B osse and Cha les-Dom-
inique (1990) ecognized wo small Pe op-
e yx species in F ench Guiana ha hey
iden i ied as P.mac o is (o iginally de-
sc ibed by Wagne [1843] based on a B a-
zilian specimen om Ma o G osso) and P.
ini a is (desc ibed by Mille [1899] based
on ou specimens om T inidad). P e ious-
ly, Sanbo n (1937) and Goodwin and G een-
hall (1961) had concluded ha ini a is was
no mo e han subspeci ically dis inc om
mac o is because specimens e e able o
hese axa a e simila and we e no hen
known om sympa y. Al hough Handley
(1976) subsequen ly epo ed ha P.mac o-
is and P. ini a is occu sympa ically in
Venezuela, he p o ided no discussion o di-
agnos ic cha ac e s. In consequence, mos e-
cen au ho s (e.g., Jones and Hood, 1993;
Koopman, 1993, 1994) ha e con inued o
ecognize ini a is as a subspecies o P.ma-
c o is.
Acco ding o B osse and Cha les-Domi-
nique (1990), Pe op e yx mac o is and P.
ini a is can be dis inguished in F ench Gui-
ana based on size (e.g., o ea m leng h 43.0–
34 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 19. Do sal (A), en al (B), and la e al (C) iews o he skull o Pe op e yx mac o is (AMNH
266005; male) om Pa acou. Scale ba
5
5 mm.
1998 35SIMMONS AND VOSS: PARACOU BATS
Fig. 20. Close-up la e al iews o he uppe igh den i ion o (A)Pe op e yx mac o is (AMNH
267396; emale) om Pa acou and (B) he holo ype o P. ini a is (AMNH 7496; emale) om T inidad.
Occlusal iews o he uppe igh den i ion (C)P.mac o is (AMNH 267396; emale) and (D) he
holo ype o P. ini a is (AMNH 7496; emale) a e shown below. No e he species di e ence in mo -
phology o he an e io uppe p emola . Scale ba s
5
5 mm.
36 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
47.0 mm, maxilla y oo h ow leng h 5.3–6.0
mm in P.mac o is, o ea m 39.5–40.4 mm,
maxilla y oo h ow 4.7–4.8 in P. ini a is),
skull shape (sho e , mo e ounded in P. in-
i a is), and ea and agus shape (bo h na -
owe in P. ini a is). They also obse ed
di e ences in sexual dimo phism ( emales
la ge han males in P.mac o is bu no in
P. ini a is) and social beha io (social
g oups o 1–4 indi iduals in P.mac o is,
much la ge [
.
100] in P. ini a is). Un o -
una ely, all o hei obse a ions conce ning
P. ini a is we e based on a single colony
om which only wo specimens (one male
and one emale) we e collec ed (B osse and
Cha les-Dominique, 1990).
Ou ouche ma e ial om Pa acou ap-
pea s o ep esen he same species ha B os-
se and Cha les-Dominique iden i ied as Pe -
op e yx mac o is despi e some disc epancies
in measu emen s. Whe eas all measu emen s
o he wo emales in ou sample ( able 4)
all wi hin he ange o a ia ion hey e-
po ed o emale P.mac o is (op. ci .), he
o ea m measu emen s o ou h ee males
(39.7, 40.1, and 42.2 mm) co espond mo e
closely o alues hey epo ed o P. ini-
a is. Examina ion o skull shape and ea
mo phology, howe e , sugges ha all o ou
specimens ep esen P.mac o is sensu B os-
se and Cha les-Dominique. The e o e, i
he e a e wo diagnosable axa o small Pe -
op e yx in F ench Guiana, i appea s ha
males canno be dis inguished on he basis o
o ea m measu emen s alone.
To es ou iden i ica ion o he Pa acou
specimens, we compa ed ou se ies wi h he
adul emale holo ype (AMNH 7496) o Pe -
op e yx ini a is. The o ea m o he holo-
ype measu es 41.9 mm, much smalle han
hose o ei he o ou Pa acou emales. Al-
hough he skull o he holo ype is badly
damaged, he os um and oo h ows emain
la gely in ac ( ig. 20). The maxilla y oo h-
ow leng h o AMNH 7496 is 5.21 mm, and
he b ead h ac oss he uppe mola s is 5.74
mm, alues ha likewise all well below
hose o ou emale Pa acou ouche s ( able
4) and o he emale specimen iden i ied as
P.mac o is by B osse and Cha les-Dominique
(1990). We also no ed one quali a i e di e -
ence be ween he holo ype o ini a is and
ou ouche ma e ial: he an e io uppe p e-
mola is peglike and lacks well-de ined an-
e io and pos e io cusps in ini a is, bu
his oo h is la ge and has a dis inc pos e io
cusp in he Pa acou specimens. Pa a ypes
(AMNH 7493, 7494, 7495) and opo ypes
(AMNH 175556, 175558, 175559) o ini-
a is esemble he holo ype in size and also
ha e iny, peglike an e io uppe p emola s.
Acco ding o Ca e and Dolan (1978), he
holo ype o Pe op e yx mac o is is an unca-
alogued adul emale p ese ed in alcohol a
he Na u his o isches Museum Wien. Un o -
una ely, he skull is missing and p esumed
los . Howe e , he o ea m measu emen o
he holo ype epo ed by Ca e and Dolan,
45.8 mm, co esponds closely wi h ha o
ou emale specimen om Pa acou (45.0
mm) and wi h hose iden i ied as P.mac o is
by B osse and Cha les Dominique (43.9–
47.0 mm). These measu emen s a e la ge
han, and do no o e lap wi h, measu emen s
p e iously epo ed o emales o P. ini-
a is (e.g., by Goodwin and G eenhall, 1961).
We he e o e ollow B osse and Cha les-
Dominique in e e ing he la ge o he wo
small Pe op e yx om F ench Guiana o P.
mac o is.
F
IELD
O
BSERVATIONS
: We cap u ed only
i e indi iduals o Pe op e yx mac o is, all
o hem a oos s. One oos ing g oup o
h ee ba s ( wo adul males and one adul e-
male) was ound benea h he b oken unk o
a allen ee can ile e ed abou 1.5 m abo e
he g ound in well-d ained p ima y o es
( ig. 16). Ano he g oup o h ee indi iduals
(o which only one adul male and one sub-
adul emale we e cap u ed) was ound in a
la ge hollow log (ca. 60 cm in inside diam-
e e ), also in well-d ained p ima y o es .
Rhynchonyc e is naso (Wied)
V
OUCHER
M
ATERIAL
: 4 emales (AMNH
*265985, *265988, *267373; MNHN
*1995.844) and 6 males (AMNH *265981,
*265986, *265987, *267372; MNHN
*1995.845, *1995.846); see able 7 o mea-
su emen s.
I
DENTIFICATION
: Sanbo n (1937) and Jones
and Hood (1993) discussed c anial and den-
al cha ac e s o Rhychonyc e is naso, and
Husson (1962, 1978) p o ided a de ailed de-
sc ip ion and compa a i e measu emen s.

1998 37SIMMONS AND VOSS: PARACOU BATS
Measu emen s o specimens p e iously col-
lec ed in F ench Guiana we e epo ed by
B osse and Cha les-Dominique (1990). No
subspecies a e cu en ly ecognized (Koop-
man, 1994).
Ou ouche specimens con o m o p e-
ious quali a i e desc ip ions o Rhynchon-
yc e is naso and measu emen s all wi hin
he ange o size a ia ion p e iously epo -
ed o he species. Howe e , ou mo pho-
me ic da a ( able 7) somewha exceed he
ange o size a ia ion known om he
Guianas, ep esen ing specimens bo h sligh -
ly smalle and sligh ly la ge han hose e-
po ed by Husson (1962, 1978) and B osse
and Cha les-Dominique (1990).
F
IELD
O
BSERVATIONS
: We cap u ed 10 in-
di iduals o Rhynchonyc e is naso, all o
which we e ound oos ing o e wa e in
la ge (3–4 m in diame e ) me al cul e s o
unde conc e e b idges. The i e oos ing
g oups we obse ed anged in size om
h ee o abou nine indi iduals. One g oup o
h ee ba s consis ed o wo adul males and
one adul emale, and one g oup o abou
nine con ained a leas wo adul males, bu
ou in o ma ion abou he age and sex com-
posi ion o he o he g oups is oo incomple e
o be in o ma i e.
Saccop e yx
Re e ences use ul o iden i ying species
o Saccop e yx include Sanbo n (1937) and
Jones and Hood (1993); bo h pape s p o ide
keys based on ex e nal and c anioden al mo -
phology. Al hough h ee size-g aded species
( om la ge o small, S.bilinea a,S.lep u a,
and S.canescens) a e commonly collec ed
sympa ically in Amazonia, he small species
a Pa acou is S.gymnu a, p e iously un-
known om he Guianas.
Saccop e yx bilinea a (Temminck)
V
OUCHER
M
ATERIAL
: 11 emales (AMNH
*265963, *265965, *267058, *267060,
*267377, *267379, *267842; MNHN
*1995.847, *1995.848, *1995.849, *1995.850)
and 11 males (AMNH *265962, 265964,
*265966, *267057, *267064, *267374,
*267378, *267840; MNHN *1995.851,
*1995.852, *1995.853); see able 8 o mea-
su emen s. One indi idual o unknown sex
(AMNH 266977) was eco e ed om he
c op o a ba alcon.
I
DENTIFICATION
: Desc ip ions and measu e-
men s o Saccop e yx bilinea a om he
Guianas and elsewhe e we e p o ided by
Thomas (1904), Sanbo n (1937), Goodwin
and G eenhall (1961), Husson (1962, 1978),
and B osse and Cha les-Dominique (1990).
The e is conside able disag eemen conce n-
ing inomial nomencla u e. Thomas (1904)
explici ly ecognized only wo subspecies, S.
b.bilinea a in no he n Sou h Ame ica (in-
cluding T inidad), and S.b.cen alis in Cen-
al Ame ica. Subsequen ly, Sanbo n (1937)
a gued ha size a ia ion in his species was
con inuously dis ibu ed and p ecluded delin-
38 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
ea ion o subspecies in he absence o o he
signi ican cha ac e s; in his e ision, he
names insignis,pe spicilli e , and cen alis
we e he e o e ega ded as s ic synonyms
o S.bilinea a.
Al hough many au ho s ha e ollowed
Sanbo n’s (1937) opinion (e.g., Husson,
1962, 1978), o he s ha e no . Cab e a
(1958), o example, ecognized S.pe spicil-
li e ( om T inidad) as a dis inc species, and
Goodwin and G eenhall (1961) la e ea ed
pe spicilli e as a alid subspecies o S.bili-
nea a. Mos ecen ly, Koopman (1994) ec-
ognized S.b.bilinea a as anging om op-
ical Mexico h oughou Cen al Ame ica and
opical Sou h Ame ica excep no he n Ven-
ezuela, T inidad, and Tobago, and S.b.pe -
spicilli e as occu ing in no he n Venezuela,
T inidad, and Tobago).
Recogni ion o pe spicilli e as a axon
dis inc om bilinea a is appa en ly based
en i ely on size, wi h specimens om no h-
e n Venezuela, T inidad ( he ype locali y),
and Tobago supposedly being la ge and
mo e obus han specimens om elsewhe e
in he Neo opics (Mille , 1899; Goodwin
and G eenhall, 1961). Howe e , his pu a i e
geog aphic pa e n is no suppo ed by a ail-
able da a. Ins ead, ou pe sonal obse a ions
and da a epo ed by Thomas (1904), San-
bo n (1937), Husson (1962, 1978), and B os-
se and Cha les-Dominique (1990) indica e
ha specimens om eas e n Ecuado , Guy-
ana, Su inam, F ench Guiana, B azil, and
Boli ia exhibi simila o ea m, skull, and
den al measu emen s as specimens om
no he n Venezuela, T inidad, and Tobago.
Thus, al hough signi ican geog aphic a ia-
ion may exis among some popula ions o
Saccop e yx bilinea a, none o he subspecies
1998 39SIMMONS AND VOSS: PARACOU BATS
Fig. 21. Roos o Saccop e yx bilinea a and Mic onyc e is hi su a in c eekside p ima y o es a
Pa acou. Small oos ing g oups o S.bilinea a occupied he hal -open basal pa o he cen al ca i y
(la ge a ow) whe e hey we e jus isible wi hou a i icial illumina ion in 1992 and 1994. A oos ing
g oup o M.hi su a occupied he comple ely enclosed da k in e io (small a ow) se e al me e s abo e
he colony o S.bilinea a in 1992. In 1993 his oos was occupied only by Ca ollia pe spicilla a.
40 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
adi ionally ecognized by au ho s appea o
be jus i ied by he e idence a hand.
Ou ouche ma e ial om Pa acou con-
o ms wi h p e ious desc ip ions o he spe-
cies, and measu emen s all wi hin he ange
o size a ia ion p e iously documen ed om
he Guianas.
F
IELD
O
BSERVATIONS
: We eco ded 34 cap-
u es (possibly including some ecap u es) o
Saccop e yx bilinea a: 14 in g ound-le el
mis ne s, 11 in ele a ed mis ne s, and 9 a
oos s; addi ionally, 1 specimen was eco -
e ed om he c op o a oad-killed Falco u-
igula is (which had also ea en a specimen o
S.lep u a). O he 14 g ound-le el mis ne
cap u es, 4 we e made in well-d ained p i-
ma y o es , 2 in swampy p ima y o es , 3
in c eekside p ima y o es , and 5 in man-
made clea ings. Six S.bilinea a we e aken
in ne s suspended 10–21 m o e a na ow
di oad, wo we e ne ed 10–13 m o e a
ee all gap in c eekside p ima y o es , and
h ee we e ne ed 7–10 m abo e he g ound
in he subcanopy o swampy p ima y o es .
Six o ou mis ne cap u es we e made be o e
da k, be ween 18:00 and 18:35 hou s.
We ound i e oos ing g oups o Saccop-
e yx bilinea a a Pa acou. All o hese oc-
cupied mo e-o -less e ical ca i ies (by con-
as wi h he essen ially ho izon al chambe s
ypically used by Co mu a b e i os is and
Pe op e yx spp.) wi h no -qui e-comple ely
da k in e io s ( he oos ing ba s we e usually
jus isible wi hou illumina ion), bu o he
oos cha ac e is ics di e ed. One oos was
in a 2-m-high chimney, open abo e and be-
low, o med by he used bu esses o a la ge
ee ( ig. 3); ano he was in he semicylind i-
cal basal opening o a cen al ca i y ha ex-
ended a up in o he unk o a much small-
e ee ( ig. 21); wo we e in he open bases
o hollow ees big enough o a g own man
o s and inside; and he i h was an open
cle , 12 m abo e he g ound a i s lowes
poin , in he unk o ano he big ee (ac-
cessed by climbing a ope). Roos ing g oups
ha we we e able o coun a ied om one
o i e indi iduals. Two indi iduals ha we
ound oos ing alone we e bo h adul males.
One g oup o i e ha we cap u ed in i s en-
i e y consis ed o wo adul males, wo adul
emales, and one ju enile.
Saccop e yx gymnu a Thomas
Figu e 22
V
OUCHER
M
ATERIAL
: 2 males (AMNH
*267843; MNHN *1995.862); see able 8 o
measu emen s.
I
DENTIFICATION
: Cha ac e s and measu e-
men s o Saccop e yx gymnu a we e dis-
cussed by Thomas (1901b) and Sanbo n
(1937). No subspecies a e cu en ecognized
(Koopman, 1994).
Ou ma e ial o Saccop e yx gymnu a
om Pa acou ep esen s a ange ex ension o
app oxima ely 900 km o his species, which
was p e iously known only om a ew lo-
cali ies along he lowe Amazon in B azil
(Koopman, 1994). Despi e his conside able
geog aphic hia us, he Pa acou specimens
con o m closely wi h published desc ip ions
o he holo ype (Thomas, 1901b; Sanbo n,
1937) and a e indis inguishable om o he
B azilian ma e ial e e able o S.gymnu a
(AMNH 93519, 93520; USNM 392995,
460080).
The only species ha migh be con used
wi h Saccop e yx gymnu a is S.canescens.
Al hough simila in size, hese iny ba s can
be dis inguished unambiguously by (1) do -
sal u colo (da k b own in gymnu a, b own
wi h s ong g ayish o yellowish os ing in
canescens), (2) do sal u ma kings (s ipes
absen o e y ain in gymnu a, a pai o
whi e s ipes always isible and o en b igh
in canescens), (3) en al u banding (uni-
colo ed da k b own in gymnu a, bicolo ed
black o da k b own wi h an ips in canes-
cens), (4) o igin o he pos e io pa o he
wing memb ane ( om he me a a sals in
gymnu a, om he ibia in canescens), and
(5) leng h o he o ea m (33.5–35.3 mm in
gymnu a, 35.8–40.8 mm in canescens). Al-
hough he e is some species o e lap in
leng h o he maxilla y oo h ow, ou speci-
mens o gymnu a ha e sho e oo h ows
(4.6 mm) han hose p e iously epo ed o
specimens o canescens (4.9–5.0 mm) om
F ench Guiana (B osse and Cha les-Domi-
nique, 1990).
F
IELD
O
BSERVATIONS
: Bo h o ou speci-
mens o Saccop e yx gymnu a we e cap u ed
in g ound-le el mis ne s, one in a small
clea ing bo de ed by well-d ained p ima y
o es ( ig. 2), he o he o e a na ow di
1998 47SIMMONS AND VOSS: PARACOU BATS
F ench Guiana by B osse and Cha les-Dom-
inique (1990).
F
IELD
O
BSERVATIONS
: All o he i e Des-
modus o undus we cap u ed a Pa acou we e
aken in g ound-le el mis ne s. Two indi id-
uals we e aken in well-d ained p ima y o -
es , wo in swampy p ima y o es , and one
in a manmade clea ing.
Diaemus youngi (Jen ink)
V
OUCHER
M
ATERIAL
: 1 emale (AMNH
*268571) and 2 males (AMNH *266347;
MNHN *1995.984); see able 11 o mea-
su emen s.
I
DENTIFICATION
: Desc ip ions and measu e-
men s o Diaemus youngi appea in Husson
(1962, 1978), Swanepoel and Genoways
(1979), Hall (1981), Koopman (1988), and
G eenhall and Schu (1996). Some au ho s
place D.youngi in he genus Desmodus (e.g.,
Handley, 1976; Emmons, 1990), bu we e-
ain Diaemus as a dis inc genus ollowing
Cab e a (1958), Husson (1962, 1978), Jones
and Ca e (1976), Hall (1981), Koopman
(1993, 1994), and G eenhall and Schu
(1996). Al hough Thomas (1928c) named D.
y.cypselinus based on ma e ial om Pe u,
Husson (1962: 198) no ed ha cypselinus
was p obably ‘‘no subspeci ically di e en
om he ypical o m.’’ Subspecies o Diae-
mus youngi ha e no been ecognized by
mos au ho s (e.g., Cab e a, 1958; Koopman,
1993, 1994; G eenhall and Schu , 1996).
Ou ma e ial o Diaemus youngi om Pa -
acou con o ms closely wi h p e ious quali-
a i e desc ip ions o he species. Al hough
o ea m and c anial dimensions o ou spec-
imens a e conside ably smalle han hose e-
po ed o specimens om Mexico, Cos a
Rica, Pe u, Venezuela, and T inidad, hey a e
only sligh ly smalle han hose p e iously
epo ed om he Guianas (Thomas, 1928b;
Goodwin and G eenhall, 1961; Husson,
1962; Hall, 1981; B osse and Cha les-Dom-
inique, 1990; G eenhall and Schu , 1996).
The signi icance o his a ia ion is no clea ,
bu we no e ha he Guyanese holo ype o
D.youngi (RNH 12088) is one o he small-
es specimens hi he o epo ed in he li e a-
u e (Husson, 1962; Ca e and Dolan, 1978).
Thus, i subspecies a e ecognized in some
u u e e isiona y s udy, ou ma e ial would
p esumably be e e ed o he nomina e o m.
F
IELD
O
BSERVATIONS
: Two o he h ee
Diaemus youngi we cap u ed a Pa acou
we e aken in g ound-le el mis ne s, one in
well-d ained p ima y o es , and he o he in
a manmade clea ing. The hi d specimen was
cap u ed in a mis ne suspended 17–20 m
abo e a na ow di oad.
GLOSSOPHAGINAE
We cap u ed i e glossophagine species a
Pa acou, bu one o wo addi ional species
migh also be expec ed in ou s udy a ea (ap-
pendix 1). G ea ca e mus be exe cised in
iden i ying glossophagines in he ield, as
e en some gene a can be ha d o dis inguish
ex e nally un il amilia i y is gained wi h he
nuances o cha ac e exp ession in his
g oup.
Anou a caudi e a (E. Geo oy)
V
OUCHER
M
ATERIAL
: 1 male (AMNH
*267290); see able 12 o measu emen s.
I
DENTIFICATION
: Keys o he species o An-
ou a we e p o ided by Tamsi and Nago sen
(1982) and Handley (1984), bu new species
ha e been desc ibed subsequen ly (Molina i,
1994) and addi ional axa s ill awai desc ip-
ion (Emmons, 1997). Handley (1984) and

48 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Molina i (1994) a e especially use ul e e -
ences because hey abula ed measu emen
da a and p o ided de ailed mo phological
compa isons among species ha could easily
be con used. O he use ul e e ences con ain-
ing desc ip ions and measu emen s o A.cau-
di e a
6
a e Husson (1962, 1978), Tamsi and
Valdi ieso (1966), Taddei (1975b), Swane-
poel and Genoways (1979), Williams and
Genoways (1980a), Webs e and Jones
(1984), and B osse and Cha les-Dominique
(1990). No subspecies o A.caudi e a a e
cu en ly ecognized (Koopman, 1994; Mol-
ina i, 1994).
Al hough ou single specimen ag ees in
mos espec s wi h p e ious desc ip ions o
Anou a caudi e a, se e al o i s measu e-
men s all in he zone o mo phome ic o e -
lap be ween A.caudi e a and a smalle spe-
cies ecen ly desc ibed by Molina i (1994),
A.luismanueli. Like wo indi iduals o cau-
di e a epo ed om Su inam by Williams
and Genoways (1980a), he Pa acou speci-
men lacks a dis inc ail enclosed wi hin he
u opa agium, and hus o al body leng h is
educed. Fo ea m leng h o he Pa acou spec-
imen likewise alls a he bo om end o he
ange o a ia ion p e iously epo ed o A.
caudi e a (e.g., by Tamsi and Valdi ieso,
1966). Molina i (1994) sugges ed ha Tam-
si and Valdi ieso may ha e included spec-
imens o luismanueli o ano he small un-
desc ibed Anou a in hei samples, bu Mol-
ina i’s es ima e o he ange o o ea m
leng h in caudi e a di e s li le om hose
o p e ious au ho s. Ou specimen has a
o ea m sligh ly smalle han Molina i e-
po ed o ue caudi e a (33.6 mm as com-
pa ed wi h 34.6), a di e ence ha we do no
ega d as signi ican gi en he ange o a i-
a ion commonly seen wi hin o he phyllos o-
mid species. The Pa acou specimen also has
a sho e maxilla y oo h ow han do speci-
mens o caudi e a p e iously epo ed om
F ench Guiana and Venezuela, bu in his e-
spec i esembles Su inamese ma e ial mea-
su ed by Husson (1962, 1978), Williams and
Genoways (1980a), and Molina i (1994).
Compa isons o ou ouche o Molina i’s
6
We ollow Handley’s (1984) spelling o he speci ic
epi he , which Koopman (1993) and mos o he au ho s
ha e inco ec ly spelled ‘‘caudi e .’’
(1994) desc ip ions o Anou a luismanueli
and A.caudi e a a e o he wise consis en
wi h i s iden i ica ion as a small example o
caudi e a. The Pa acou specimen has a lon-
ge calca and b oade u opa agium han seen
in luismanueli, and i lacks a con inuous,
dense inge o hai s along he u opa agial
edge. Ins ead, he ee ma gin o he u opa-
agium is ligh ly u ed nea he midline bu
is naked la e ally, a ai diagnos ic o cau-
di e a (Molina i, 1994). O he aspec s o he
pelage o he Pa acou specimen ag ee wi h
Molina i’s desc ip ion o caudi e a wi h he
excep ion o he colo o he bases o he
hai s in ou specimen, which a e g ayish-
whi e a he han yellowish- o c eamy-whi e
as he desc ibed. Gi en he ange o a ia ion
in basal u colo ha we ha e obse ed in
o he species, we in e p e his as wi hin-spe-
cies a ia ion.
F
IELD
O
BSERVATIONS
: Ou single example
o Anou a caudi e a was aken in a g ound-
le el mis ne in c eekside p ima y o es .
Choe oniscus mino (Pe e s)
Figu e 24
V
OUCHER
M
ATERIAL
: 8 emales (AMNH
*266120, *266121, *266123, *266377,
*267153; MNHN *1998.668, 1998.669,
*1998.670) and 5 males (AMNH *266124,
*267946, *267947, *267948; MNHN
1998.671); see ables 13 and 14 o mea-
su emen s.
I
DENTIFICATION
: The complex axonomic
his o y o Choe oniscus, pa icula ly o he
smalle species o iginally placed in he genus
Choe onyc e is, is un o una ely ele an o
unde s anding he co ec iden i ica ion o
ou Pa acou ouche s.
Choe onyc e is mino was i s desc ibed
by Pe e s (1868) based on a specimen om
Su inam. Al hough Goodwin and G eenhall
(1961) s a ed ha he ype was des oyed in
Wo ld Wa II, Husson (1962) epo ed ha
Pe e ’s o iginal specimen was, in ac , s ill
in ac a he S u ga museum. Husson
(1962) published measu emen s and a pho-
og aph o he holo ype (SMN 441), and also
p o ided a de ailed desc ip ion. A second
small species, Choe onyc e is in e media,
was desc ibed by Allen and Chapman (1893)
based on specimens om T inidad, and Tho-
1998 49SIMMONS AND VOSS: PARACOU BATS
mas (1912a) named a hi d, Choe onyc e is
inca, based on a specimen om Pe u. Tho-
mas (1928a) subsequen ly subdi ided Cho-
e onyc e is, placing mino ,in e media, and
inca in his new genus Choe oniscus.
Conside able con usion has su ounded
he use o hese h ee speci ic epi he s (Tho-
mas, 1912a; Sanbo n, 1954; Goodwin and
G eenhall, 1961; Husson, 1962; Koopman,
1978; and Williams and Genoways, 1980a),
much o i he esul o small a ailable sam-
ple sizes and ma ked sexual dimo phism
(Genoways e al., 1973; Koopman, 1978).
The la e ac o is c ucial because he ho-
lo ypes o mino and inca a e males, while
ha o in e medius is a emale. Con lic ing
epo s conce ning he mo phology and sex
o he supposedly los holo ype o mino
only se ed o compound he eal biological
p oblem, whe he hese names eally ep e-
sen h ee o ewe species.
The e is gene al ag eemen ha Allen and
Chapman (1893) named hei Tinidadian ma-
e ial as a new species in pa because se e al
measu emen s o mino we e inco ec ly e-
po ed by Pe e s (1868). Thomas (1912a:
404) obse ed ha
The T inidad ba desc ibed by Allen and Chapman as
Ch.in e media is e y p obably he same as Ch.mi-
no , as hose au ho s we e decei ed by Pe e ’s im-
possible measu emen o 11 mm. o he calca , an
o gan d awn as abou 5 mm. long in he mo e ecen ly
published pla e o he la e species . . .
Thomas (1912a) hen desc ibed inca based
on den al p opo ions (leng h and wid h o
he p emola s and mola s) in which i sup-
posedly di e ed om mino . He also no ed
ha he skull o inca was la ge and b oade
han ha o mino , bu his compa ison was
appa en ly based on a specimen o mino
o he han he holo ype, as measu emen s o
he la e do no con i m his obse a ion ( a-
ble 13).
Mos ecen au ho s ha e ollowed Koop-
man (1978) in ega ding inca as a junio syn-
onym o mino , and in e medius as a po en-
ially dis inc species. Koopman’s (1978: 8)
conclusion ha wo species could be ecog-
nized was based on obse a ions conce ning
den al a ia ion and leng h o he os um:
S udy o a numbe o skulls o Choe oniscus om
T inidad, B i ish Guiana, B azil, Pe u, and Ecuado
[has] con inced me ha wo species can be dis in-
guished . . . by os al leng h, which is e lec ed in
maxilla y oo h ow leng h. Thomas (1912[a]) dis in-
guished inca om mino solely by he ela i e sizes
o he p emola s and mola s. On he basis o hese
skulls, I can see conside able a ia ion in his cha -
ac e , bu no clea cu dis inc ion be ween wo o ms.
The species wi h he longe os um would hen be
called C.mino (inca a synonym) and he species
wi h he sho e os um would s and as C.in e media
[sic].
Koopman (1994) ga e he ange o mino as
ex ending om coas al Ecuado o he mou h
o he Amazon, no hwa d o eas e n Vene-
zuela, and sou hwa d o no hwes e n Boli -
ia. Al hough mos specimens o small Cho-
e oniscus ecen ly collec ed in he Guianas
ha e been iden i ied as in e medius (e.g., by
B osse and Cha les-Dominique, 1990), a
long-muzzled emale om Guyana (AMNH
140471) was e e ed o mino by Williams
and Genoways (1980a).
In a emp ing o iden i y ou ma e ial om
Pa acou, we examined he holo ype o in e -
medius; Husson’s (1962) measu emen s, pho-
og aphs, and desc ip ion o he holo ype o
mino ; Thomas’s (1912a) desc ip ion o he
holo ype o inca; specimens e e ed o in-
e medius and mino in he AMNH; and
measu emen s o addi ional specimens e-
e ed o in e medius and mino by Geno-
ways e al. (1973), Genoways and Williams
(1980a), Ochoa and Fe na´ndez (1982), B os-
se and Cha les-Dominique (1990), and An-
de son (1997). Al hough we kep males and
emales sepa a e, and in es iga ed a ia ion
in size, skull shape, and den al mo phology,
we ound no consis en di e ences among
named o ms o small Choe oniscus. Fo ex-
ample, den al mo phology in ou se ies o 13
specimens om Pa acou is highly a iable,
wi h some indi iduals ha ing much sho e
o longe ee h han o he s, o he s ha ing
na owe o b oade ee h, e c. These di e -
ences o m no consis en pa e n and do no
appea o be co ela ed wi h body size o
wi h os al p opo ions. Because ou sample
includes indi iduals wi h bo h ‘‘mino - ype’’
and ‘‘inca- ype’’ ee h (sensu Thomas,
1912a), as well as o he s wi h in e media e
condi ions, we ollow Koopman (1978, 1993,
1994) in concluding ha mino and inca a e
conspeci ic.
Den al mo phology has ne e been used o
sepa a e mino (o inca) om in e medius.
50 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 24. Ven al iews o he skull o six indi iduals o Choe oniscus mino om Pa acou showing
in aspeci ic a ia ion in c anioden al mo phology. (A) AMNH 266377 ( emale); (B) MNHN 1995.1271
( emale); (C) AMNH 266123 ( emale); (D) AMNH 267948 (male); (E) AMNH 267946 (male); (F)
AMNH 267947 (male). No e a ia ion in ela i e size o he p emola s and mola s, skull leng h and
b ead h, os al p opo ions, and o m o he pala e. This se ies includes indi iduals wi h ‘‘mino - ype’’
1998 51SIMMONS AND VOSS: PARACOU BATS
←
den i ions (sho p emola s and long mola s; e.g., E), o he s wi h ‘‘inca- ype’’ ee h (p emola s and
mola s subequal in leng h; e.g., D), and den al in e media es (e.g., A). I also includes la ge indi iduals
wi h a ela i ely long (‘‘mino - ype’’) os um (e.g., C), smalle specimens wi h a sho e (‘‘in e medius-
ype’’) os um (e.g., A), and in e media es (e.g., B). Scale ba
5
10 mm.
Ins ead, measu emen s o he skull, os um,
and maxilla y oo h ow appea o be he only
cha ac e s so employed in ecen yea s
(Koopman, 1978; Williams and Genoways,
1980a). Ra he han con i ming ha mino
and in e medius a e dis inc species, ou
compa isons o measu emen da a show he
opposi e. Whe eas measu emen s o ou e-
male specimens om Pa acou b oadly o e -
lap hose o he emale holo ype o in e -
medius, measu emen s o ou males o e lap
hose o he male holo ypes o mino and
inca ( ables 13, 14). Fo bo h sexes, mea-
su emen s o ou sample co espond closely
o hose epo ed o a la ge se ies (10 males
and 26 emales) o opo ypical in e medius
om T inidad (Genoways e al., 1973). Mea-
su emen s o o he specimens om Venezue-
la and he Guianas ( epo ed by Genoways
and Williams [1980a], Ochoa and Fe na´ndez
[1982], and B osse and Cha les-Dominique
[1990]) and Boli ia (Ande son, 1997) u he
ill he mo phome ic gap ha Koopman
(1978) obse ed be ween his small se ies o
mino and in e medius. Conside ing he pos-
sibli y ha ou la ges ouche specimens
52 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
migh ep esen mino and he smalles in-
e medius, we ca e ully examined a ia ion
in ex e nal and c anioden al mo phology
wi hin ou se ies, bu ul ima ely ound no e -
idence ha he Pa acou ma e ial ep esen s
mo e han one species. In ou iew, speci-
mens iden i ied as in e medius in he li e a-
u e simply ep esen he lowe end o a no -
mal ange o wi hin-species size a ia ion,
while hose iden i ied as mino and inca ep-
esen he uppe end o his ange. We he e-
o e conclude ha hese epi he s ep esen a
single species o which he oldes a ailable
name is Choe oniscus mino (Pe e s, 1868).
Recogni ion o subspecies appea s o be un-
jus i ied based on he da a cu en ly a hand.
F
IELD
O
BSERVATIONS
: We cap u ed 13 in-
di iduals o Choe oniscus mino a Pa acou,
o which 7 we e aken in g ound-le el mis -
ne s and 6 a oos s. Th ee indi iduals we e
ne ed in swampy p ima y o es , h ee in
c eekside p ima y o es , and one in a man-
made clea ing. One oos ing g oup o six
ba s was ound on he da k unde side o a
allen ee wi h la ge bu esses (simila o
ha shown in ig. 17) in well-d ained p i-
ma y o es ; o his g oup, one adul male
and ou adul emales we e cap u ed, and
one ba escaped. Ano he oos ing ba , a sol-
i a y adul male, was ound hanging benea h
he unde cu bank o a d y s eambed in
well-d ained p ima y o es .
Glossophaga so icina (Pallas)
V
OUCHER
M
ATERIAL
: 10 emales (AMNH
*267134, *267137, *267138, *267448,
*267449, *267953, *267956, 267958; MNHN

1998 53SIMMONS AND VOSS: PARACOU BATS
*1995.1040, *1995.1041) and 25 males
(AMNH *266090, *266091, *266092,
*266093, *266094, *266095, 266099,
*267139, *267140, *267949, *267950,
*267951, 267954, *267955, 267957, *267959;
MNHN *1995.1042, *1995.1043, *1995.1044,
*1995.1045, *1995.1046, *1995.1047,
1995.1048, *1995.1049, *1995.1050); see a-
ble 15 o measu emen s.
I
DENTIFICATION
: The mos use ul e e ence
o iden i ying species o Glossophaga is
Webs e ’s (1993) e ision, which includes a
key as well as de ailed desc ip ions and com-
pa a i e measu emen s. Al a ez e al. (1991)
p o ided a good desc ip ion o G.so icina,
bu he p incipal e e ence o he species is
also Webs e ’s e ision. Fi e subspecies o
G.so icina a e cu en ly ecognized, o
which h ee occu in Sou h Ame ica: G.s.
handleyi (Mexico h oughou Cen al Ame -
ica o no he n and wes e n Colombia), G.s.
so icina (Sou h Ame ica eas o he Andes
om Colombia and Venezuela in he no h
o Pa aguay and no he n A gen ina), and G.
s. alens (d ie a eas o wes e n Ecuado and
Pe u) (Al a ez e al., 1991; Webs e , 1993;
Koopman, 1994). Measu emen s o speci-
mens aken in he Guianas we e epo ed by
Husson (1962, 1978), B osse and Cha les-
Dominique (1990), and Webs e (1993).
Ou Pa acou ma e ial con o ms in all e-
spec s wi h Webs e ’s (1993) desc ip ion o
Glossophaga so icina so icina, and hei
measu emen s ( able 15) all wi hin he ange
o a ia ion p e iously epo ed o G.so i-
cina in B azil and he Guianas (Husson,
1962, 1978; Taddei, 1975b; B osse and
Cha les-Dominique, 1990; Webs e , 1993).
F
IELD
O
BSERVATIONS
: We eco ded 56 cap-
u es (possibly including some ecap u es) o
Glossophaga so icina a Pa acou, o which
45 we e in g ound-le el mis ne s and 11
we e a oos s. Se en o he mis ne cap u es
we e in well-d ained p ima y o es , 11 we e
in swampy p ima y o es , 3 we e in c eek-
side p ima y o es , and 24 we e in manmade
clea ings. The p opo ional di e ence in cap-
u e-habi a equencies be ween his species
and Lonchophylla homasi, he only o he
common glossophagine a Pa acou (see be-
low), is no ewo hy: G.so icina was mo e
commonly ne ed in modi ied habi a s and L.
homasi in p ima y habi a s han would be
expec ed i hese sympa ic nec a i o es did
no di e in habi a use ( able 16). Al hough
his esul is consis en wi h B osse and
54 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Cha les-Dominique’s (1990) cha ac e iza ion
o G.so icina as a species o modi ied bio-
opes, we no e ha specimens a e in ac
known om ecologically p is ine locali ies in
F ench Guiana (e.g., USNM 548471,
548472, collec ed by L. H. Emmons a Sau
Pa a e´ in 1984).
O he h ee oos s o Glossophaga so i-
cina we ound a Pa acou, wo we e in ee
ca i ies ( igs. 25, 26) and one was unde un-
de a conc e e b idge. The b idge oos con-
ained many G.so icina, pe haps hund eds
(o which se en males and one emale we e
collec ed as ouche s), as well as oos ing
g oups o Rhynchonyc e is naso and Ca ollia
pe spicilla a. One ee ca i y oos occupied
by an adul male- emale pai o G.so icina
( ig. 26) was also sha ed wi h C.pe spicil-
la a; he o he ee ca i y oos was occupied
a soli a y adul male.
Lichonyc e is obscu a Thomas
V
OUCHER
M
ATERIAL
: 1 emale (AMNH
*267960); see able 15 o measu emen s.
I
DENTIFICATION
: Two colo mo phs occu
among specimens o Lichonyc e is, a da k
b own o m ha ag ees wi h Thomas’ (1895)
o iginal desc ip ion o L.obscu a, and a pale
b own o m ha is o en iden i ied as L.de-
gene Mille (1931). Howe e , he axonomic
s a us o he la e is p oblema ic due o con-
usion ega ding pa e ns o a ia ion in bo h
c anioden al and pelage cha ac e s (Ga dne ,
1976; Hill, 1985; Ochoa e al., 1993). Pend-
ing a e ision o he genus, we ollow Ga d-
ne (1976), Hill (1985), Ochoa e al. (1993),
and Koopman (1993, 1994) in ega ding de-
gene as a junio synonym o obscu a. De-
sc ip ions and compa a i e measu emen s o
L.obscu a so de ined can be ound in Hus-
son (1962, 1978), Ga dne (1976), Swane-
poel and Genoways (1979), Hill (1985),
B osse and Cha les-Dominique (1990), and
Ochoa e al. (1993).
Ou chocola e-b own specimen ag ees
closely wi h p e ious desc ip ions o Lichon-
yc e is obscu a, and alls wi hin he ange o
a ia ion in measu emen s p e iously epo -
ed in he li e a u e.
F
IELD
O
BSERVATIONS
: Ou specimen was
cap u ed in a g ound-le el mis ne in well-
d ained p ima y o es .
Lonchophylla homasi J. A. Allen
V
OUCHER
M
ATERIAL
: 15 emales (AMNH
*266100, *266107, *266108, *266109,
*267147, *267148, *267149, *267451,
*267452, *267939; MNHN *1995.1096,
*1995.1097, *1995.1098, *1995.1099,
*1995.1100) and 23 males (AMNH
*266101, *266102, *266103, *266104,
*266105, *266106, *266110, *266113,
*266114, *266116, *266117, 267150,
*267940, *267941, *267942, *267943;
MNHN *1995.1101, *1995.1102, *1995.1103,
*1995.1104, *1995.1105, 1995.1106,
*1995.1107); see able 17 o measu emen s.
I
DENTIFICATION
: The mos use ul e e ence
o iden i ying species o Lonchophylla is
Taddei e al. (1983), who p o ided a key and
discussed a ia ion. We consul ed desc ip-
ions and compa a i e measu emen s o Lon-
chophylla homasi in Husson (1962, 1978),
Hill (1964, 1980), Ga dne (1976), Taddei e
al. (1978, 1983), Swanepoel and Genoways
(1979), and B osse and Cha les-Dominique
(1990). No subspecies o L. homasi a e cu -
en ly ecognized (Taddei e al., 1983; Koop-
man, 1994).
Ou ouche ma e ial, one o he la ges
se ies a ailable om a single locali y, con-
o ms in all espec s o p e ious desc ip ions
o Lonchophylla homasi.
F
IELD
O
BSERVATIONS
: We eco ded 55 cap-
u es (possibly including some ecap u es) o
1998 55SIMMONS AND VOSS: PARACOU BATS
Fig. 25. A la ge kimbo o, P adosia cochlea ia (Sapo aceae), a common canopy eme gen in well-
d ained p ima y o es a Pa acou ha o en has o ed ca i ies shel e ing ba s and o he mammals. In
1991 he ca e nous lowe chambe o his ee (a ow) con ained a soli a y Glossophaga so icina, bu
in 1992 we ound a oos ing g oup o ou Mic onyc e is megalo is and ano he consis ing o one
Ca ollia pe spicilla a in he same space; in 1993 and 1994 his oos was occupied by small g oups o
C.pe spicilla a only.
56 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 26. Roos o Glossophaga so icina,Phyllos omus elonga us, and Ca ollia pe spicilla a in well-
d ained p ima y o es a Pa acou. This ee ca i y, abou 60–70 cm in diame e and se e al me e s
high, con ained abou 20 C.pe spicilla a ( wo o which, bo h adul emales, we e collec ed as ouche s)
and one adul male P.elonga us in 1992. In 1993 he same space con ained abou 10 C.pe spicilla a
and 2 G.so icina (an adul male- emale pai ). I is possible ha his oos was used by Vampy um
spec um in 1991, when RSV obse ed wo huge ba s ocalizing loudly and lying a ound he opening
a nigh ; howe e , he oos was acan when we climbed he ee o in es iga e wo days la e .
1998 63SIMMONS AND VOSS: PARACOU BATS
Fig. 28. Do sal (A), en al (B), and la e al (C) iews o he skull o he holo ype o Mic onyc e is
b osse i (MNHN 1995.1030; male). Scale ba
5
10 mm.

64 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
*266034, *266035, *267419), all o which
we e likewise collec ed a Pa acou. See a-
bles 22 and 23 o measu emen s.
R
EFERRED
M
ATERIAL
: In addi ion o he
ype se ies om Pa acou, we also e e nine
specimens o Mic onyc e is b osse i om
ou o he Sou h Ame ican locali ies as ol-
lows: PERU, Lo e o, Pue o Indiana, Rı´o
1998 65SIMMONS AND VOSS: PARACOU BATS
Amazonas nea mou h o Rı´o Napo (AMNH
73495, 73496, 73498, 73499); Pe u, Lo e o,
Jena o He e a (MUSM 5528); Pe u, Lo e o,
Rı´o Ya a ı´ Mi ı´m, Queb ada Espe anza
(FMNH 89100, 89101, 89102); BRAZIL,
Sa˜o Paulo, Rio Juquia´, Ba a (FMNH
92997). One o hese specimens (MUSM
5528) was p e iously iden i ied as M.
schmid o um by Asco a e al. (1991a).
D
ISTRIBUTION
: Al hough specimens o Mi-
c onyc e is b osse i a e cu en ly known only
om eas e n Pe u, F ench Guiana, and sou h-
eas e n B azil, he species could easonably
be expec ed o occu h oughou Amazonia
and he B azilian A lan ic o es .
E
TYMOLOGY
: We name his species o hon-
o And e´ B osse in ecogni ion o his many
impo an con ibu ions o knowledge o
opical ba aunas in India, A ica, and
Sou h Ame ica. In pa icula , B osse and
Cha les-Dominique (1990) p o ided an in-
dispensable guide o he ba auna o F ench
Guiana, as he a e ed co e s, c eased pages,
and anno a ed ma gins o ou well- humbed
ield copy eloquen ly a es .
D
IAGNOSIS
: A small Mic onyc e is wi h
da k b own do sal u and pale (g ay o bu )
en al u ; do sal hai s bicolo ed, 7–8 mm
long in shoulde egion, wi h whi e base
comp ising
⅓
–
½
o each hai ; u on ou side
o medial hi d o pinna sho (
#
3 mm) and
dense; pinnae la ge wi h ounded ips, con-
nec ed ac oss c own o head by low in e -
au icula band wi h shallow midline no ch;
humb small; second phalanx o wing digi
IV sho e han i s ; calca longe han oo ;
mas oid b ead h less han zygoma ic b ead h;
c own heigh o uppe inciso s no educed;
no gap p esen be ween ou e uppe inciso
and canine; P3 and P4 subequal in an e o-
pos e io leng h, heigh o P3 slighly less
han ha o P4; P4 wi h small lingual heel
and poo ly de eloped lingual cusp; M1 na -
owe han M2; no no iceable gap be ween
pos e io edge o cingulum o M2 and
an e olingual edge o M3 when oo h ow
seen in occlusal iew; lowe inciso s small
and bilobed; p2 and p4 subequal in an e o-
pos e io leng h and heigh , p3 smalle in
bo h dimensions; co onoid p ocess high, up-
pe ma gin o ascending p ocess wi h s eep
slope (25–30
8
).
O he cha ac e s lis ed abo e, ou a e
pa icula ly use ul o ield iden i ica ion o
M.b osse i: colo o he en al u (pale
g ay o bu ), leng h o he u on he leading
edge o he pinna (sho ,
#
3 mm), ela i e
leng h o he i s and second phalanges o
wing digi IV (second sho e han i s ), and
he calca (longe han oo ). Wi hin he ge-
nus Mic onyc e is, his combina ion o ai s
is unique o M.b osse i.
M
EASUREMENTS
: A summa y o measu e-
men s o Mic onyc e is b osse i, including
hose o he ype se ies and o all o he e-
e ed specimens, is p o ided in able 23.
D
ESCRIPTION AND
C
OMPARISONS
:Mic o-
nyc e is b osse i equi es close compa ison
wi h six o he like-size congene s wi h which
i migh be con used, namely M.homezi,M.
megalo is,M.mic o is,M.minu a,M.san-
bo ni, and M.schmid o um. By con as , M.
hi su a s ands ou om all o he membe s o
he genus by i s la ge size and unique lowe
inciso mo phology (see able 24 and species
accoun o hi su a below). The ollowing
pa ag aphs he e o e omi hi su a om u -
he conside a ion and ocus only on he e-
maining se en species.
Mic onyc e is b osse i is one o he small-
es membe s o he genus, wi h mean alues
o mos o i s measu emen s alling well be-
low hose o o he species ound a Pa acou
( ables 23–25). Despi e some mo phome ic
o e lap be ween la ge indi iduals o b os-
se i and a ew smalle specimens in he con-
gene ic se ies measu ed by Simmons (1996b:
able 2), measu emen s a e clea ly use ul o
dis inguishing b osse i om mos o he Mi-
c onyc e is species whe e hey occu sym-
pa ically. A Pa acou, he only species om
which b osse i is indis inguishable in uni-
a ia e mo phome ic compa isons ( ables
23–25) is mic o is.
A Pa acou, b osse i is consis en ly small-
e han megalo is in body weigh ( emales
only), ea leng h, humb leng h, g ea es
leng h o skull (males only), b aincase
b ead h ( emales only), mas oid b ead h, zy-
goma ic b ead h ( emales only), maxilla y
oo h ow leng h, and b ead h ac oss mola s
(males only). Simila ly, b osse i is smalle
han homezi (known only om one male; see
species accoun below) in body weigh , ea
leng h, o ea m leng h, ibia leng h, and in
66 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
all c anioden al measu emen s. In compa i-
son wi h minu a,b osse i is consis en ly
smalle in body weigh , ea leng h, g ea es
leng h o skull ( emales only), condyloinci-
si e leng h ( emales only), pos o bi al
b ead h, b aincase b ead h ( emales only),
mas oid b ead h, maxilla y oo h ow leng h
( emales only), and b ead h ac oss mola s
( emales only). Likewise, b osse i is smalle
han schmid o um in many measu emen s,
including body weigh , ea leng h, ibia
leng h, humb leng h, and all c anioden al
measu emen s.
The en i e do sal pelage o b osse i is
composed o bicolo ed hai s wi h whi e bas-
es and b own ips. In he uppe back egion,
he whi e bases comp ises app oxima ely
one- hi d o one-hal o he leng h o each
1998 67SIMMONS AND VOSS: PARACOU BATS
hai . This is compa able o he pa e n seen
in mic o is,megalo is,schmid o um, and
homezi, in which he whi e bases comp ise
one- ou h o one-hal o each hai on he
uppe back. In con as , he whi e bases a e
much mo e ex ensi e in minu a (one-hal o
wo- hi ds o each hai ) and sanbo ni ( wo-
hi ds o h ee- ou hs o each hai ).
The leng h o he do sal u o e he uppe
back in b osse i is app oxima ely 7–8 mm,
compa able o ha o mic o is and homezi.
In con as , minu a and sanbo ni ha e sho e
u o e he shoulde s (5–7 mm), while
megalo is and schmid o um ha e longe u
(8–11 mm).
The en al u o b osse i (including ha
co e ing he neck and h oa ) is ei he pale
g ay o pale bu , whe eas ha o megalo is
68 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
and mic o is is b own (app oxima ely he
same shade as he do sal u ). The unde pa s
o schmid o um and minu a a e also pale
(g ay o bu ), like b osse i, bu sanbo ni has
b igh whi e en al u .
All o he Pa acou specimens o b osse i
ha e a pale g ay en e , as do some speci-
mens om Pe u (e.g., MUSM 5528). How-
e e , some olde and somewha aded Pe u-
ian specimens o b osse i ha e pale bu
en e s (e.g., AMNH 73495–73499, collec -
ed in 1926). I is no clea i he bu colo
is he esul o pos mo em al e a ion o wi h-
in-species a ia ion. Bo h en al u -colo
a ian s can be obse ed among museum se-
ies o o he species (e.g., minu a and
schmid o um), so his a ia ion (wha e e i s
cause) is e iden ly no peculia o b osse i.
The u on he ou side o he medial hi d
o he ea pinna is sho (
#
3 mm) and dense
in b osse i, simila o he condi ion seen in
mic o is,minu a, and sanbo ni. By con as ,
he au icula u o megalo is,schmid o um,
and homezi is much longe (5–8 mm).
As in all o he species o Mic onyc e is,
he ea s o b osse i a e connec ed ac oss he
c own o he head by an in e au icula band
o skin. In b osse i, his band is low and
nea ly ho izon al in p o ile, and is ma ked a
he midline by a shallow no ch; a simila
condi ion is seen in megalo is and mic o is.
This mo phology con as s sha ply wi h he
in e au icula band in minu a,sanbo ni, and
homezi, which is much highe and di ided
by a deep median no ch in o wo oughly i-
angula laps (see species accoun o homezi
below). The in e au icula band o schimid-
o um is o in e media e heigh wi h a mod-
e a ely deep median no ch.
The second phalanx o wing digi IV is
sho e han he i s in b osse i, as i is also
in megalo is,mic o is,schmid o um, and
homezi. By con as , he i s and second pha-
langes o digi IV a e subequal in minu a and
sanbo ni.
The calca is dis inc ly longe han he
hind oo in b osse i, esembling he condi-
ion seen in megalo is,mic o is, and schmid-
o um. The calca is app oxima ely equal in
leng h o he hind oo in sanbo ni, howe e ,
and he calca is sho e han he oo in min-
u a and homezi.
As no ed by Simmons (1996b), c anial
mo phology is ema kably simila in all spe-
cies o Mic onyc e is. Like o he membe s o
he genus, b osse i has a ela i ely long os-
um, no an eo bi al in la ion, a la ge b ain-
case, and shallow basisphenoid pi s. Mas oid
b ead h is less han zygoma ic b ead h in
b osse i, as i is likewise in megalo is,mi-
c o is, and schmid o um; mas oid b ead h is
g ea e han zygoma ic b ead h in minu a,
sanbo ni, and homezi.
Like all o he congene ic species, b osse i
has a den al o mula o I 2/2, C 1/1, P 2/3,
M 3/3
3
2
5
34. The c own heigh o he
uppe inciso s is no educed, and no gap is
p esen be ween he ou e uppe inciso and
he canine. In his espec , b osse i esembles
mos species o Mic onyc e is; he con as -
ing mo phology is seen only in sanbo ni.
Al hough di e ences in he uppe p emo-
la den i ion a e sub le, a ia ion in his e-
gion p o ides an impo an means o dis in-
guishing species o Mic onyc e is (see Sim-
mons, 1996b). In b osse i, P3 and P4 a e
subequal in an e opos e io leng h, and he
heigh o P3 is e y slighly less han ha o
P4. This is simila o he condi ion seen in
megalo is,mic o is,schmid o um, and hom-
ezi. Some a ia ion is known in he o me
h ee species, wi h P3 and P4 subequal in
heigh in some indi iduals and P3 sligh ly
smalle han P4 in o he s. Al hough we did
no obse e such a ia ion in b osse i (o
homezi, known om only one specimen), i
would no be unexpec ed. In con as o he
condi ions desc ibed abo e, P3 is no iceably
smalle han P4 in bo h leng h and heigh in
minu a and sanbo ni.
The deg ee o de elopmen o he pos e o-
lingual heel and lingual cusp on P4 a ies
among and wi hin species o Mic onyc e is.
In b osse i, P4 has a small lingual heel and
a poo ly de eloped lingual cusp. The lingual
heel o P4 in b osse i is simila in size o
hose o megalo is,mic o is,schmid o um,
and homezi, bu he lingual heel is gene ally
smalle in minu a and la ge in sanbo ni. The
lingual cusp o P4, which is o med om he
aised edge o he oo h, is poo ly de eloped
and lacks a sha p poin in b osse i,homezi,
minu a, and sanbo ni, and i is en i ely lack-
ing in many specimens o schmid o um.By
con as , his cusp is be e de eloped and a-
pe s o a sha p poin in mos specimens o

1998 69SIMMONS AND VOSS: PARACOU BATS
mic o is and megalo is; howe e , some in-
di iduals o bo h species ha e a poo ly de-
eloped cusp, while his cusp may be lacking
en i ely in o he s.
The i s uppe mola is no iceably na ow-
e han M2 in b osse i as i is in mos o he
species o Mic onyc e is. Uniquely, he lin-
gual po ion o M1 is expanded so ha M1
and M2 a e subequal in wid h in sanbo ni,
which also di e s om all o he species in
ha ing a la ge gap isible be ween he pos-
e io edge o he cingulum o M2 and he
an e olingual edge o M3 when he oo h ow
is seen in occlusal iew.
The lowe den i ion o b osse i closely e-
sembles ha o mos o he species, wi h mi-
no di e ences appa en only in he p emo-
la s. In b osse i and schmid o um, p2 and p4
a e subequal in an e opos e io leng h and in
heigh , while p3 is sligh ly smalle in bo h
dimensions. In minu a,sanbo ni, and home-
zi, p3 is u he educed ela i e o p2 and
p4, which a e la ge and subequal. By con-
as , p3 is a la ge oo h (app oxima ely he
same size as p4) in mic o is and megalo is;
in hese axa, p2 is o en sligh ly la ge han
ei he p3 o p4, al hough he e is some wi h-
in-species a ia ion in his ai and all h ee
lowe p emola s a e subequal in some indi-
iduals o bo h species.
The co onoid p ocess in b osse i is high,
and he uppe ma gin o he ascending p o-
cess has a s eep slope (25–30
8
), simila o he
mandibula mo phology o megalo is,mic o-
is, and schmid o um. In con as , he co o-
noid p ocess is compa a i ely low and he
uppe ma gin o he ascending amus has a
shallow slope (16–18
8
)inminu a,sanbo ni,
and homezi.
F
IELD
O
BSERVATIONS
: O he eigh speci-
mens o Mic onyc e is b osse i ha we col-
lec ed a Pa acou, one was aken in a g ound-
le el mis ne and he emaining se en om
a single oos in a hollow ee; bo h cap u e
si es we e in well-d ained p ima y o es .
The oos ee, abou 1 m in diame e a
b eas heigh , had a single en ance hole
(abou 20 cm in diame e ) 1 m abo e he
g ound. A mis ne enclosu e a ound he base
o he ee cap u ed se en o he eigh ba s
ha eme ged (be ween 18:45 and 19:15
hou s) on 19 Augus 1991; he cap u ed an-
imals consis ed o ou adul males and h ee
adul emales. In addi ion, he mis ne enclo-
su e cap u ed wo Phyllos omus elonga us on
he ou side as hey came swooping owa d
he oos opening, ob iously in ending o
ambush he much smalle eme ging ba s. We
we e unable o e isi his oos in subse-
quen yea s because he si e was clea ed o
ag icul u al esea ch in 1992.
Mic onyc e is hi su a (Pe e s)
V
OUCHER
M
ATERIAL
: 3 emales (AMNH
*267857, *267858; MNHN *1995.800) and
6 males (AMNH *267093, *267894,
*267096, 267860; MNHN *1995.801,
1995.802); see able 24 o measu emen s.
I
DENTIFICATION
:Mic onyc e is hi su a is
easy o iden i y owing o i s la ge size (see
Simmons [1996b] and ables 23–25) and
uniquely high-c owned lowe inciso s ( ig-
u ed by Genoways and Williams, 1986). De-
sc ip ions and compa a i e measu emen s o
M.hi su a om he Guianas and elsewhe e
we e p o ided by Sanbo n (1949), Goodwin
and G eenhall (1961), Hill (1964), Da is
(1976a), Genoways and Williams (1986),
B osse and Cha les-Dominique (1990), and
Simmons (1996b). No subspecies a e cu -
en ly ecognized (Simmons, 1996b).
Ou specimens o Mic onyc e is hi su a
con o m in all espec s o p e ious desc ip-
ions o he species.
F
IELD
O
BSERVATIONS
: We cap u ed nine in-
di iduals o Mic onyc e is hi su a a Pa a-
cou, h ee o which we e aken in g ound-
le el mis ne s ( wo in well-d ained p ima y
o es and one in swampy p ima y o es ),
and he o he six a oos s. The wo oos s
we ound we e simila : bo h we e hollow
ees nea s eams in p ima y o es , bo h
ees had sloping unks, and he cen al ca -
i y o each ee had a lowe en ance 1–2 m
abo e he g ound and ano he smalle ape -
u e se e al me e s highe up he unk. F om
one ee ( ig. 21) we collec ed an adul male
and an adul emale (p obably no he en i e
oos ing g oup); om he o he ( ig. 29) we
collec ed wo adul males, one adul emale,
and one subadul male (possibly he en i e
oos ing g oup). The loo o he la e oos
was li e ed wi h he wings o la ge o hop-
e ans, p esumably he emains o p ey ca -
70 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 29. Roos o Mic onyc e is hi su a in swampy p ima y o es a Pa acou. Like he oos illus-
a ed in igu e 21, his is an inclined ee (sloping away om he came a in his iew) wi h a hollow
cen al ca i y accessed by a la ge lowe ape u e (a ow) and a second smalle opening much highe up
he unk (no isible). The p esence o ba s in his oos was in e ed om a pile o la ge o hop e an
wings ound a he base o he lowe ape u e. In he o eg ound a e cu palm onds ha we used o
ha ch a mis ne enclosu e in which ou eme ging ba s we e cap u ed.
1998 71SIMMONS AND VOSS: PARACOU BATS
ied back o he oos and ea en by he in-
habi an s.
Mic onyc e is homezi Pi lo
Figu es 30–32
V
OUCHER
M
ATERIAL
: 1 male (AMNH
*267414); see able 25 o measu emen s.
I
DENTIFICATION
:Mic onyc e is homezi was
o iginally desc ibed by Pi lo (1967), who
conside ed i a subspecies o M.megalo is.
P io o he p esen s udy, he only specimens
e e ed o homezi we e hose o Pi lo ’s ype
se ies om he Ma acaibo Basin o no h-
wes e n Venezuela (ca. 2200 km WNW o
Pa acou). Simmons (1996b: 4) discussed he
b ie his o y o his axon:
Mic onyc e is megalo is homezi Pi lo , 1967, was
named based on h ee specimens collec ed in he
wes e n Venezuelan s a e o Zulia. These specimens
(including he holo ype) we e des oyed along wi h
he es o Pi lo ’s collec ion some ime du ing he
1970s (Pi lo , phone con e sa ion wi h T. G i i hs).
The published desc ip ion o homezi is incomple e,
bu se e al ea u es desc ibed by Pi lo (e.g., pale
en e , deep no ch in ea memb ane) sugges ha
homezi is no ela ed o megalo is. I is no clea om
he ype desc ip ion i homezi ep esen s a dis inc
species o is synonymous wi h minu a,schmid o um,
o sanbo ni.
Simmons (1996b) ound Pi lo ’s desc ip ion
o homezi o be inadequa e o se e al ea-
sons, including (1) no desc ip ion o he den-
i ion, (2) no measu emen s o he han leng h
o he o ea m, and (3) no men ion o ela i e
size o he calca . The only uly dis inc i e
cha ac e o homezi desc ibed by Pi lo was
a ‘‘c eux cu ane´ glab e’’ (smoo h cu aneous
ossa) on he op o he head be ween he ea s
behind he in e au icula band. Pi lo (1967:
265) desc ibed his peculia s uc u e as ol-
lows:
C’es une pe i e plage subo ale, pa iellemen glab e,
limi e´e pa un bou ele e`s ne . Les deux e san s
an e´ ieu e pos e´ ieu de ce e plage se e minen
dans le sillon ans e sal qui a d’une o eille a` l’au e.
Su le e san an e´ ieu , une ange´e de poils ela i e-
men longs es implan e´e.
An accompanying d awing o he do sum o
he head wi h he ea s and in e au icula band
olded o wa d (op. ci .: ig. 1) shows li le
mo e han he loca ion o he s uc u e in
ques ion. Al hough Pi lo did no explici ly
discuss sexual dimo phism, he appa en ly
had bo h male and emale examples o hom-
ezi in hand (op. ci .: 262), sugges ing ha he
cu aneous ossa is p esen in bo h sexes.
Ha ing ne e seen a s uc u e esembling
Pi lo ’s cu aneous ossa, Simmons (1996b)
in e ed ha he was jus desc ibing he naked
pa ch o skin ha lies unde nea h he in e -
au icula band and he a ached hai u in
all Mic onyc e is species. The in e au icula
band is no mally olded back agains he op
o he head so ha he pos e io ( en al) su -
ace o he band is closely app essed o he
skin o he head. The c anial skin co e ed by
he band appea s naked (al hough a spa se
coa o ine hai s can be seen unde magni-
ica ion), and i is ully exposed only when
he ea s a e d awn o wa d. This naked pa ch
a ies in size among species o Mic onyc e -
is, being la ges in species wi h a high in e -
au icua band (e.g., minu a,sanbo ni) and
smalles in hose wi h a low in e au icula
band (e.g., megalo is,mic o is). Howe e ,
his in e p e a ion o Pi lo ’s ‘‘c eux cu ane´
glab e’’ now appea s inco ec .
One o he specimens ha we collec ed a
Pa acou—AMNH 267414, an adul male—
has a well-de ined, hai less ossa on he op
o he head be ween he ea s and behind he
in e au icula band ( ig. 30). Smoo h and gla-
b ous inside, his pi is su ounded by a im
o skin wi h coa se p ojec ing hai s, exac ly
as Pi lo desc ibed. Al hough no his ological
wo k has ye been done, his s uc u e ap-
pea s o be a glandula pouch o a pouch in
which glandula sec e ions migh be s o ed.
To he bes o ou knowledge, his s uc u e
is unique among ba s ( o a summa y o
known chi op e an glandula s uc u es, see
Quay, 1970).
Se e al o he cha ac e s o ou specimen
co espond wi h Pi lo ’s (1967) desc ip ion
o homezi, including (1) small size ( o ea m
leng h 36.5 mm in Pa acou specimen, 31.4–
37.4 mm in Pi lo ’s se ies), (2) a pale en e
(much ligh e han he do sal u ), and (3) a
high in e au icula band di ided in o oughly
iangula laps by a deep median no ch. Be-
cause ou ma e ial ag ees closely wi h Pi -
lo ’s desc ip ion, and because o i s clea dis-
inc ness om any o he named congene ic
o m, we conclude ha AMNH 267414 ep-
esen s a alid species ha should be called
Mic onyc e is homezi.
72 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 30. Li e iews o Mic onyc e is homezi (AMNH 267414; male). No e he pale en al u , he
long hai s on he leading edge o each pinna, and he e y deep, V-shaped no ch in he in e au icula
band; he cu aneous ossa is isible h ough he no ch in he in e au icula band. The hai s co e ing he
ossa in his iew a e oo ed in he skin su ounding he ossa, and on he unde side (pos e io su ace)
o he in e au icula band; he ossa i sel is hai less.
1998 79SIMMONS AND VOSS: PARACOU BATS
ly eiden i ied as ei he M.sanbo ni (Sim-
mons, 1996b) o M.b osse i (see abo e).
Simmons (1996b) p o ided a summa y o
measu emen s o Mic onyc e is schmid o um
sensu s ic o om h oughou i s known geo-
g aphic ange. No subspecies a e cu en ly
ecognized (Simmons, 1996b).
Al hough ou ouche ma e ial gene ally
ag ees wi h p e ious desc ip ions o Mic o-
nyc e is schmid o um, Pa acou specimens
ha e pale g ay en al u a he han he pale
bu en e s seen in specimens om o he
locali ies. Because we did no ind any o he
consis en di e ences, and because o he
congene s wi h pale en al u (e.g., M.
b osse i and M.minu a) exhibi simila ch o-
ma ic a iabili y, we conclude ha his ep-
esen s no mal in aspeci ic a ia ion.
In addi ion o cha ac e s discussed p e i-
ously, we ound ha ibia leng h was help ul
o dis inguishing Mic onyc e is schmid o-
um om sympa ic species in he hand, a
leas a Pa acou. Whe eas bo h M.schmid-
o um and M.homezi ha e long ibias (15.3–
15.8 mm), he lowe leg is consis en ly sho -
e (12.8–14.6 mm) in M.b osse i,M.mega-
lo is,M.mic o is, and M.minu a.
F
IELD
O
BSERVATIONS
: Ou wo examples o
Mic onyc e is schmid o um we e bo h aken
in g ound-le el mis ne s in well-d ained p i-
ma y o es .
Mimon benne ii (G ay)
Figu es 34–37
V
OUCHER
M
ATERIAL
: 1 emale (AMNH
*267109) and 1 male (MNHN *1995.988);
see able 26 o measu emen s.
I
DENTIFICATION
: Al hough Koopman
(1993, 1994) conside ed Mimon benne ii
and M.cozumelae o be conspeci ic, many
o he au ho s ha e ecognized hem as dis-
inc species (Dalques , 1957; Handley, 1960;
Ca e e al., 1966; Ga dne and Pa on,
1972; Jones and Ca e , 1976; Swanepoel and
Genoways, 1979; McCa hy, 1987; McCa -
hy e al., 1993). Whe eas he es ic ed ype
locali y o M.benne ii is Ipanema in he
B azilian s a e o Sa˜o Paulo (He shko i z,
1951), he ype locali y o M.cozumelae is
Cozumel Island in he Mexican s a e o
Quin ana Roo (Goldman, 1914). Cu en ly,
he known ange o cozumelae ex ends om
sou he n Mexico o no hwes e n Colombia,
and ha o benne ii ex ends om he Guia-
nas o sou heas e n B azil (Koopman, 1994).
Dalques (1957) epo ed addi ional spec-
imens o Mimon cozumelae collec ed in he
decades ollowing Goldman’s (1914) o iginal
desc ip ion, and discussed di e ences be-
ween cozumelae and benne ii, which he
ea ed as dis inc species. Howe e , as
Handley (1960) subsequen ly no ed, Dal-
ques ’s only compa a i e example o benne -
ii was a ju enile. Schaldach (1964) subse-
quen ly summa ized cha ac e s supposedly
dis inguishing hese axa, including size
(smalle in benne ii), do sal pelage colo
(da ke in benne ii, b igh e in cozumelae),
and leng h and woolliness o he middo sal
hai s (long and no woolly in benne ii, sho
and woolly in cozumelae). Like Dalques ’s
(1957) accoun , howe e , Schaldach’s de-
sc ip ion o he pelage o benne ii seems o
ha e been based on ju enile o subadul
specimens. Despi e he mo phological di e -
ences he no ed, Schaldach concluded ha
benne ii and cozumelae a e conspeci ic, a
decision based in la ge pa on he geog aph-
ic gap be ween hei known geog aphic ang-
es. Hall (1981) and Koopman (1993, 1994)
ollowed Schaldach (1964) wi hou addi ion-
al discussion.
Con a Schaldach (1964), Hall (1981), and
Koopman (1993, 1994), we conside Mimon
benne ii and M.cozumelae o ep esen dis-
inc species based on appa en ly consis en
di e ences in he ollowing cha ac e s: (1)
do sal pelage colo (mo e eddish in adul
benne ii, less eddish in cozumelae), (2) col-
o o he wing ip (da k in benne ii, whi e in
cozumelae), (3) shape o he middle uppe
inciso s ( ape ing o poin s in benne ii, mo e
spa ula e in cozumelae), (4) o m o he low-
e inciso s (na owe in benne ii), (5) mo -
phology o m3 ( alonid la ge and be e de-
eloped in benne ii), and (6) mo phology o
he pos e io pala al ma gin (b oade wi h U-
shaped mesop e ygoid no ch in benne ii,
na owe wi h V-shaped no ch in cozume-
lae).
8
Al hough some o e lap exis s, benne -
8
This lis o cha ac e s is based on bo h li e a u e ac-
coun s and ou examina ion o ouche specimens. Spec-
imens examined in addi ion o hose om Pa acou: Mi-
mon benne ii (B azil: USNM 123393, 391027); Mimon
cozumelae (Mexico: AMNH 144508, 185862–185872).

80 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 34. Do sal (A) and en al (B) iews o he skull o Mimon benne ii (MNHN 1995.988; male)
om Pa acou. Do sal (C) and en al (D) iews o he skull o M.cozumelae (AMNH 265107; male)
om Gua emala. No e he species di e ences in inciso mo phology and in he shape o he mesop e y-
goid no ch o he pos e io pala e. Scale ba s
5
10 mm.
1998 81SIMMONS AND VOSS: PARACOU BATS
Fig. 35. La e al iews o he skull and lowe jaw o (A)Mimon benne ii (MNHN 1995.988; male)
and (B)M.cozumelae (AMNH 265107; male). Scale ba s
5
10 mm.
ii is gene ally smalle han cozumelae (e.g.,
o ea m leng h o 50.0–56.6 mm in benne ii,
54.6–60.7 mm in cozumelae; Swanepoel and
Genoways, 1979; Hall, 1981; B osse and
Cha les-Dominique, 1990). Al hough hese
axa ha e no been collec ed in sympa y, i
is coun e p oduc i e o ega d hem as sub-
species gi en he magni ude o hei mo -
82 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 36. An e io iews o he uppe and low-
e den i ion o (A)Mimon benne ii (MNHN
1995.988; male) and (B)M.cozumelae (AMNH
265107; male). No e ha he middle uppe inci-
so s o benne ii ape o poin s, whe eas hese
ee h a e mo e spa ula e in cozumelae. Scale ba s
5
5 mm. Fig. 37. Occlusal iews o he c own o he
igh m3 in (A)Mimon benne ii (MNHN
1995.988; male) and (B)M.cozumelae (AMNH
265107; male). No e ha he alonid is la ge and
be e de eloped in benne ii han i is in cozu-
melae. Scale ba s
5
1 mm.
phological di e gence; un il p o en alse, he
app op ia e null hypo hesis is ha hey ep-
esen dis inc species.
Desc ip ions and measu emen s o Mimon
benne ii sensu s ic o we e p o ided by G ay
(1838), Dalques (1957), Husson (1962,
1978), Hill (1964), Swanepoel and Geno-
ways (1979), and B osse and Cha les-Dom-
inique (1990). Ou specimens closely esem-
ble o he s p e iously epo ed om Guianas,
pa icula ly he b igh o ange- usse adul s
desc ibed by B osse and Cha les-Dominique
(1990). Based on ou limi ed compa isons,
ecogni ion o subspecies does no seem wa -
an ed.
F
IELD
O
BSERVATIONS
: One o ou wo spec-
imens o Mimon benne ii was cap u ed in a
g ound-le el mis ne and he o he was aken
a a oos ; bo h cap u es we e in c eekside
p ima y o es . The oos si e was a la ge ee
(ca. 1.5 m in diame e a b eas heigh ) wi h
a cen al ca i y ex ending om g ound le el
o nea he c own. We su ounded he only
1998 83SIMMONS AND VOSS: PARACOU BATS
oos opening a g ound le el, a small hole
ca. 15 cm in diame e , wi h a mis ne enclo-
su e ha caugh one adul male Mimon ben-
ne ii and 27 adul male Ca ollia pe spicil-
la a be ween 18:10 and 18:35 hou s on 17
No embe 1992 ( he Mimon benne ii
eme ged a 18:25 hou s); no eme ging ba s
escaped. We did no e isi his oos in sub-
sequen yea s.
Mimon c enula um (E. Geo oy)
V
OUCHER
M
ATERIAL
: 7 emales (AMNH
*267114, *267115, *267437, *267887,
*267889; MNHN *1995.1032, 1995.1033)
and 10 males (AMNH *267111, 267113,
*267880, 267884, *267885, *267886,
267888; MNHN *1995.1034, *1995.1035,
*1995.1036); see able 26 o measu emen s.
I
DENTIFICATION
: Use ul desc ip ions and
measu men s o Mimon c enula um we e
p o ided by Handley (1960), Goodwin and
G eenhall (1961), Husson (1962, 1978), Hill
(1964), Genoways and Williams (1979),
Swanepoel and Genoways (1979), Ga dne
and Pa on (1972), Hall (1981), B osse and
Cha les-Dominique (1990), and Ped o e al.
(1994). O he i e subspecies ecognized by
Koopman (1994), M.c.c enula um occu s
om T inidad and eas e n Venezuela
h oughou he Guianas o sou he n Amazo-
nian B azil. Howe e , we ollow Ga dne and
Pa on (1972) in ega ding M.koepckeae
( om he highlands o cen al Pe u) as a dis-
inc species con a Koopman (1978, 1993,
1994).
Ou ouche ma e ial con o ms wi h p e-
ious desc ip ions o Mimon c enula um
c enula um. Measu emen s o Pa acou spec-
imens gene ally all wi hin he ange o a i-
a ion p e iously documen ed o Guianan
popula ions (Husson, 1962, 1978; Hill, 1964;
Genoways and Williams, 1979; B osse and
Cha les-Dominique, 1990), al hough some o
ou males a e sligh ly smalle in some di-
mensions han specimens p e iously epo ed
om he egion.
F
IELD
O
BSERVATIONS
: All o he 17 Mimon
c enula um we cap u ed a Pa acou we e
aken in g ound-le el mis ne s: 9 in well-
d ained p ima y o es , 4 in swampy p ima y
o es , and 4 in c eekside p ima y o es . An
unusually la ge p opo ion (88%) o cap u es
we e in he e y ea ly e ening, be o e 19:00
hou s.
84 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Phyllode ma s enops Pe e s
V
OUCHER
M
ATERIAL
: 3 emales (AMNH
*266078, *267441; MNHN *1995.1075) and
5 males (AMNH *266077, *267440,
*267890, *267891; MNHN *1995.1076);
see able 27 o measu emen s.
I
DENTIFICATION
: Desc ip ions and measu e-
men s o Phyllode ma s enops om he
Guianas and elsewhe e we e p o ided by
Husson (1962, 1978), Hill (1964), Ga dne
(1976), Genoways and Williams (1979),
Swanepoel and Genoways (1979), Williams
and Genoways (1980a), B osse and Cha les-
Dominique (1990), and Ande son (1997).
Th ee subspecies a e cu en ly ecognized: P.
s.sep en ionalis (sou he n Mexico o Cos a
Rica), P.s.s enops (Panama o sou he n B a-
zil), and P.s.boli iensis (cen al Boli ia)
(Ba quez and Ojeda, 1979; Koopman, 1994;
Ande son, 1997).
Ou ouche ma e ial con o ms in all e-
spec s o p e ious desc ip ions o Phyllode -
ma s enops s enops, and measu emen s all
wi hin he ange o a ia ion epo ed p e i-
ously o his axon.
F
IELD
O
BSERVATIONS
: All eigh o he
Phyllode ma s enops we cap u ed a Pa acou
we e aken in g ound-le el mis ne s: wo in
well-d ained p ima y o es , h ee in swampy
p ima y o es , wo in c eekside p ima y o -
es , and one in closed-canopy seconda y
g ow h.
Phyllos omus discolo Wagne
V
OUCHER
M
ATERIAL
: 10 emales (AMNH
*267116, *267118, *267120, *267121,
*267123, *267124; MNHN *1995.1077,
*1995.1078, *1995.1079, *1995.1080) and 5
males (AMNH *267117, *267119, *267984,
*267986; MNHN *1995.1081); see able 28
o measu emen s.
I
DENTIFICATION
: Use ul desc ip ions and
measu emen s o Phyllos omus discolo may
be ound in Goodwin and G eenhall (1961),
Husson (1962, 1978), Powe and Tamsi
(1973), Taddei (1975a), Swanepoel and Gen-
oways (1979), B osse and Cha les-Dominique
(1990), and Ande son (1997). Two subspe-
cies a e o en ecognized: P.d. e ucosus
(Mexico o no hwes e n Pe u) and P.d.dis-
colo (Sou h Ame ica eas o he Andes o
no hwes e n A gen ina; Ma ga i a Island;
T inidad) (Koopman, 1994). Powe and Tam-
si (1973), howe e , ques ioned he ecog-
ni ion o hese axa based on compa isons o
specimens om nea he supposed con ac
zone in Colombia.
Ou Pa acou specimens con o m closely
wi h p e ious desc ip ions o Phyllos omus
discolo , and measu emen s all wi hin he

1998 85SIMMONS AND VOSS: PARACOU BATS
ange o a ia ion p e iously epo ed om
he Guiana egion. Ou compa isons o pub-
lished measu emen s om h oughou he
known geog aphic ange o P.discolo sug-
ges ha Powe and Tamsi (1973) we e co -
ec and ha he e is no cu en jus i ica ion
o ecognizing subspecies.
F
IELD
O
BSERVATIONS
: We cap u ed 17
Phyllos omus discolo a Pa acou: 14 in
g ound-le el mis ne s and 3 in ele a ed ne s.
Fou o he g ound-le el cap u es we e in
well-d ained p ima y o es , eigh we e in
swampy p ima y o es , and wo we e in
c eekside p ima y o es . One o he ele a ed
mis ne cap u es was made a 18–21 m o e
a na ow di oad and wo we e made a 10–
13 m abo e a ee all gap in c eekside p i-
ma y o es . Se e al indi iduals ne ed in he
d y season we e co e ed wi h pollen. Mos
ba s we e caugh singly, bu wo adul males
and wo adul emales (possibly membe s o
a o aging lock) we e aken wi hin a 10-min-
u e in e al a one mis ne ing si e in swampy
p ima y o es on 24 Oc obe 1992, and an-
86 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
o he adul emale was caugh a ew hou s
la e a he same place.
Phyllos omus elonga us (E. Geo oy)
V
OUCHER
M
ATERIAL
: 14 emales (AMNH
*266051, *266055, *266058, *266062,
*266063, *266064, *266067, *266068,
*267152, *267897; MNHN *1995.1082,
*1995.1083, *1995.1084, *1995.1085) and
12 males (AMNH *266050, *266052,
*266053, *266054, *266065, *266066,
*266069, *267127; MNHN *1995.1086,
*1995.1087, *1995.1088, *1995.1089); see
able 28 o measu emen s.
I
DENTIFICATION
: Desc ip ions and measu e-
men s o Phyllos omus elonga us om he
Guianas and elsewhe e we e p o ided by
Husson (1962, 1978), Hill (1964), Swane-
poel and Genoways (1979), B osse and
Cha les-Dominique (1990), and Ande son
(1997). No subspecies a e cu en ly ecog-
nized (Koopman, 1994).
Ou ma e ial om Pa acou ag ees in all
espec s wi h p e ious desc ip ions o Phyl-
los omus elonga us. Like o he conspeci ic
ma e ial om he Guiana egion, ou speci-
mens all nea he uppe end o he known
ange o size a ia ion o P.elonga us.
F
IELD
O
BSERVATIONS
: We eco ded 162
cap u es (p obably including some ecap-
u es) o Phyllos omus elonga us a Pa acou.
One hund ed i y-nine cap u es we e in
g ound-le el mis ne s,
9
one was in a mis ne
suspended 13–16 m abo e a na ow di
oad, and wo we e a oos s. Se en y- ou o
he g ound-le el mis ne cap u es we e in
well-d ained p ima y o es , 45 we e in
swampy p ima y o es , 27 we e in c eekside
p ima y o es , 1 was in a na u al ee all
opening in p ima y o es , 7 we e in man-
made clea ings, and 5 we e in closed-canopy
seconda y g ow h. The wo ba s aken a
oos s we e bo h soli a y males; one was col-
lec ed om a ee ca i y wi h a single la ge
opening abou 5 m abo e he g ound ( ig.
26), he o he om he cen al sha o a hol-
9
This igu e includes wo ba s caugh on he ou side
o he mis ne enclosu e desc ibed abo e in he accoun
o Mic onyc e is b osse i. Howe e , because hose cap-
u es we e inciden al o oos sampling, hey a e exclud-
ed om he quan i a i e analysis o mis ne ing esul s
below.
low ee wi h a la ge lowe en ance a
g ound le el and second smalle ape u e ca.
15–20 m a he up he unk ( his oos was
sha ed wi h Saccop e yx bilinea a and Ca -
ollia pe spicilla a). Bo h oos s we e in well-
d ained p ima y o es .
By con as wi h he mul iple ele a ed-
mis ne cap u es we eco ded o Phyllos o-
mus discolo and P.has a us (see abo e and
below), he almos comple e absence o P.
elonga us in ne s suspended mo e han a ew
me e s abo e he g ound is no ewo hy.
Phyllos omus has a us (Pallas)
V
OUCHER
M
ATERIAL
: 11 emales (AMNH
*266071, *266072, *266073, *266075,
*267433, *267901, *267902, *267903;
MNHN *1995.1090, *1995.1091, *1995.1092)
and 8 males (AMNH *266070, *266074,
*267434, *267904, *267907; MNHN
*1995.1093, *1995.1094, *1995.1095); see
able 28 o measu emen s.
I
DENTIFICATION
: Desc ip ions and measu e-
men s o Phyllos omus has a us ha e ap-
pea ed in many publica ions; we consul ed
hose in Goodwin and G eenhall (1961),
Husson (1962, 1978), Taddei (1975a),
Swanepoel and Genoways (1979), B osse
and Cha les-Dominique (1990), and Ande -
son (1997). Two subspecies o a e cu en ly
ecognized: P.h.panamensis (Hondu as o
Pe u and eas o Venezuela) and P.h.has-
a us (eas e n Venezuela sou h o no he n
Boli ia and sou heas e n B azil) (Koopman,
1994). These a e dis inguished p incipally on
he basis o size, wi h P.h.has a us being
he smalle o m.
Ou ouche ma e ial con o ms closely
wi h p e ious desc ip ions o Phyllos omus
has a us has a us, wi h measu emen s gen-
e ally alling wi hin he ange o a ia ion
p e iously epo ed om he Guianas. One
excep ion is a pa icula ly small adul male
(AMNH 267907), some measu emen s o
which a e less han any p e iously epo ed
o he species (e.g., o ea m leng h o 77.5
mm, compa ed wi h 79.0 mm o he smalles
male epo ed by Taddei [1975a]). Compa -
isons o his diminu i e indi idual wi h o he
specimens in ou sample, howe e , ailed o
e eal any mo phological di e ences o he
han size.
1998 87SIMMONS AND VOSS: PARACOU BATS
F
IELD
O
BSERVATIONS
: We eco ded 56 cap-
u es (possibly including some ecap u es) o
Phyllos omus has a us, o which 34 we e in
g ound-le el mis ne s and 22 we e in ele a -
ed mis ne s. The 34 g ound-le el cap u es in-
cluded 9 in well-d ained p ima y o es , 11
in swampy p ima y o es , 6 in c eekside p i-
ma y o es , and 8 in manmade clea ings (7
we e aken in a banana/cacao plan a ion on
a single nigh ). O he 22 ele a ed mis ne
cap u es, 17 we e in ne s suspended 10–23
m abo e a na ow di oad, 4 we e made a
20–38 m abo e a ee all gap in well-d ained
p ima y o es , and 1 was a 7–10 m abo e
a ee all in c eekside p ima y o es .
Tona ia
Al hough keys o he species o Tona ia
p o ided by Genoways and Williams (1984)
and Medellı´n and A i a (1989) a e use ul
ools o iden i ica ion, nei he is comple e;
he o me co e s only he species known
om Su inam, and he la e was published
be o e T.bidens and T.sau ophila we e ec-
ognized as dis inc species (see below). All
o he i e species o Tona ia known om
he Guiana sub egion o Amazonia a e sym-
pa ic (and appa en ly syn opic) a Pa acou.
Tona ia b asiliense (Pe e s)
V
OUCHER
M
ATERIAL
: 3 emales (AMNH
*267103, *267104; MNHN *1995.1052) and
6 males (AMNH *267101, *267102,
*267916, *267917; MNHN *1995.1053,
*1995.1054); see able 29 o measu emen s.
I
DENTIFICATION
: Desc ip ions and measu e-
men s o Tona ia b asiliense om he Guia-
nas and elsewhe e can be ound in Goodwin
(1942), Swanepoel and Genoways (1979),
Genoways and Williams (1984), and B osse
and Cha les-Dominique (1990). No subspe-
cies a e cu en ly ecognized (Genoways and
Williams, 1984; Koopman, 1994).
As he smalles membe o i s genus, Ton-
a ia b asiliense is mos likely o be con used
wi h species o Mic onyc e is, which a e also
small g ay-b own phyllos omines wi h la ge,
ounded ea s. Genoways and Williams
(1984) men ioned se e al c anioden al cha -
ac e s ha can be used o dis inguish hese
axa, bu ailed o no e one ob ious ex e nal
cha ac e : whe eas all species o Mic onyc-
e is ha e a pai o la ge de mal pads o m-
ing a ‘‘V’’ on he ip o he chin, he ip o
he chin in T.b asiliense (and o he conge-
ne s) has a U-shaped ow o iny de mal pa-
pillae (Emmons, 1990, 1997). We also ob-
se ed ha T.b asiliense olds i s ea s back
agains he c own o he head when ouched,
a beha io seen in se e al o he species o
Tona ia (see below) bu no in Mic onyc e is.
Ou ouche ma e ial con o ms closely
wi h p e ious desc ip ions o Tona ia b asi-
liense, pa icula ly hose based on ma e ial
om he Guianas. P obably because ou se-
ies con ains mo e indi iduals han do ea lie
collec ions om he egion, i exhibi s some-
wha g ea e size a iabili y han p e iously
epo ed. Howe e , measu emen s o ou
specimens all wi hin he ange o a ia ion
p e iously documen ed o he species as a
whole.
As ema ked by Genoways and Williams
(1984), Tona ia b asiliense as cu en ly ec-
ognized may be composi e. Should his
p o e o be he case, compa isons wi h pub-
lished measu emen s (e.g., hose in Goodwin
[1942] and Swanepoel and Genoways
[1979]) sugges ha ou specimens, along
wi h o he s om he Guianas, would be e-
e ed o T.b asiliense a he han o o he
axa cu en ly ea ed as synonyms (e.g., min-
u a,nica aguae, and enezuelae; Handley
[1966], Ga dne [1976], Jones and Ca e
[1976], Genoways and Williams [1984], Me-
dellı´n and A i a [1989], Koopman [1993,
1994]).
F
IELD
O
BSERVATIONS
: All o he nine spec-
imens o Tona ia b asiliense ha we caugh
a Pa acou we e aken in g ound-le el mis -
ne s: six in swampy p ima y o es and h ee
in c eekside p ima y o es .
Tona ia ca ike i (J. A. Allen)
V
OUCHER
M
ATERIAL
: 1 emale (AMNH
*267918); see able 29 o measu emen s.
I
DENTIFICATION
:Tona ia ca ike i has been
collec ed only a ely, so desc ip ions and
measu emen s o mos known specimens
ha e been epo ed in he li e a u e, o ex-
ample, by Goodwin (1942), Husson (1962,
1978), Ga dne (1976), Swanepoel and Gen-
oways (1979), McCa hy e al. (1983, 1992),
Genoways and Williams (1984), and McCa -
88 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
hy and Handley (1987). No subspecies a e
cu en ly ecognized (Koopman, 1994).
Ou single emale specimen o Tona ia
ca ike i ag ees wi h p e ious desc ip ions o
he species in all espec s. Like se e al o he
congene s, T.ca ike i olds i s ea s back
o e he c own o he head when hey a e
ouched.
F
IELD
O
BSERVATIONS
: The single example
o Tona ia ca ike i we cap u ed a Pa acou
was aken in a g ound-le el mis ne in
swampy p ima y o es domina ed by he
palm Eu e pe ole acea.
Tona ia sau ophila
Koopman and Williams
V
OUCHER
M
ATERIAL
: 10 emales (AMNH
*266044, *266045, *266046, *266047,
*267429, *267908, *267914; MNHN
*1995.1060, *1995.1061, *1995.1062) and
11 males (AMNH *266049, *267099,
1998 95SIMMONS AND VOSS: PARACOU BATS
clea .’’ Al hough he use o inomial nomen-
cla u e o Ca ollia pe spicilla a is no cu -
en ly jus i ied, u u e mo phological o mo-
lecula s udies may ye show ha some sub-
speci ic dis inc ions a e wa an ed.
Ou specimens o Ca ollia pe spicilla a
con o m in all espec s o p e ious desc ip-
ions o he species.
F
IELD
O
BSERVATIONS
: We eco ded 1142
cap u es (including ecap u es) o Ca ollia
pe spicilla a a Pa acou, o which 1048 we e
in g ound-le el mis ne s, 18 we e in ele a ed
mis ne s, 75 we e a oos s, and 1 was in a
ha p ap. O he 1049 g ound-le el mis ne
and ha p- ap cap u es, 252 we e in well-
d ained p ima y o es , 279 we e in swampy
p ima y o es , 110 we e in c eekside p i-
ma y o es , 1 was in a ee all opening in
p ima y o es , 325 we e in manmade clea -
ings, 9 we e in closed-canopy seconda y
g ow h, and 73 we e o e oadside puddles.
The 18 cap u es in ele a ed mis ne s we e
made be ween 5 and 13 m abo e he g ound:
7 o e a na ow di oad, 7 o e ee alls,
and 4 in he subcanopy o swampy p ima y
o es .
O he 11 oos s a which we cap u ed o
obse ed Ca ollia pe spicilla a a Pa acou, 4
we e in cul e s unde oads, 1 was unde a
b idge, and 6 we e in ee ca i ies (e.g., igs.
21, 25, 26, 33). We ne e ound C.pe spi-
cilla a unde allen ees, inside hollow logs,
o in oliage o any kind. O he ba s obse ed
oos ing wi h C.pe spicilla a included Rhyn-
chonyc e is naso,Saccop e yx bilinea a,Mi-
c onyc e is megalo is,M.mic o is,Mimon
benne ii,Phyllos omus elonga us,T achops
ci hosus, and Glossophaga so icina.Ca -
ollia pe spicilla a was he only species ha
we ound oos ing in la ge bachelo g oups;
o example, 27 males we e cap u ed in a
mis ne enclosu e as hey eme ged om an
opening in he base o a la ge hollow ee (a
oos sha ed wi h Mimon benne ii, see
abo e). Ano he ee-ca i y oos ing g oup
ha we cap u ed in i s en i e y consis ed o
wo adul males, wo lac a ing adul emales,
and wo ju eniles.
Rhinophylla pumilio Pe e s
V
OUCHER
M
ATERIAL
: 25 emales (AMNH
*266168, *266171, *266178, *266184,
*266186, *266188, *266189, *266193,
*266196, *266198, *267159, *267456,
*267457, *267458, *267459, *267971;
MNHN *1998.623, *1998.624, *1998.625,
*1998.626, *1998.627, *1998.628, *1998.629,
*1998.630, *1998.631) and 24 males
(AMNH *266174, *266175, *266179,
*266180, *266181, *266182, *266183,
*266185, *266187, *266190, *266191,
*266192, *266194, *266197, *267158;
MNHN *1998.632, *1998.633, *1998.634,
*1998.635, *1998.636, *1998.637, *1998.638,
*1998.639, *1998.640); see able 35 o
measu emen s.
I
DENTIFICATION
: Desc ip ions and compa -
a i e measu emen s o Rhinophylla pumilio
ha we consul ed o iden i y ou ma e ial in-
cluded hose in Husson (1962, 1978), Hill
(1964), Ca e (1966), Swanepoel and Gen-
oways (1979), Williams and Genoways
(1980a), and B osse and Cha les-Dominique
(1990). No subspecies a e cu en ly ecog-
nized (Koopman, 1994).
Ou Pa acou specimens con o m closely
wi h p e ious quali a i e desc ip ions o
Rhinophylla pumilio in he li e a u e ci ed
abo e. Likewise, measu emen s o ou se ies
gene ally all wi hin he known ange o a i-
a ion o he species, al hough a ew a e
sligh ly smalle han any p e iously epo ed.
Because he small indi iduals in ou collec-
ion a e simila in all o he epec s o la ge
examples, we a ibu e his mino disc ep-
ancy o wi hin-popula ion mo phome ic
a ia ion. Like o he collec ions o R.pumi-
lio om he Guianas, ou Pa acou ouche s
all a he lowe end o he known ange o
size a ia ion o he species.
F
IELD
O
BSERVATIONS
: We eco ded 128
cap u es (p obably including some ecap-
u es) o Rhinophylla pumilio a Pa acou, o
which 106 we e in g ound-le el mis ne s, 2
we e in ele a ed mis ne s, 19 we e a oos s,
and 1 was in a ha p ap nea g ound le el.
O he 107 g ound-le el mis ne and ha p-
ap cap u es, 22 we e in well-d ained p i-
ma y o es , 44 we e in swampy p ima y o -
es , 14 we e in c eekside p ima y o es , 4
we e in ee all openings in p ima y o es ,
19 we e in manmade clea ings, and 4 we e
in closed-canopy seconda y g ow h. The wo
ba s cap u ed in ele a ed mis ne s we e aken

96 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
5–10 m abo e a ee all opening in c eekside
p ima y o es .
We ound eigh oos ing g oups o Rhin-
ophylla pumilio a six unique oos si es (one
oos was e isi ed wice). All o he oos s
we ound we e in oliage be ween 1.5 and 5
m abo e he g ound. Six oos ing g oups o
R.pumilio occupied ‘‘bi id’’ en s (sensu
Kunz e al., 1994) made om he e minal
lea le s o onds o imma u e unde s o y
palms ha we p o isionally iden i ied as As-
oca yum sciophilum.
11
In all eco ded con-
s uc ion de ails, hese en s exac ly esem-
bled hose in which we also ound A ibeus
cine eus (see igs. 43–45) and Ec ophylla
macconnelli (see ig. 47). Al hough we do
no know which (i any) o hese ba s was
ac ually esponsible o making such en s,
we once collec ed (on 6 Augus 1993) a
g oup o h ee R.pumilio om a en p e i-
ously occupied (on 30 July 1993) by se en
E.macconnelli. By con as , we ne e ob-
11
We base his iden i ica ion on ege a i e cha ac e s
desc ibed by Hende son e al. (1995) and de G an ille
(1997): he plica e and subco iaceous lea le s, egula ly
a anged and sp eading in one plane, ha e nonspinous
ma gins, and he long spines on he achis lack basal
wings (see igs. 44, 45). Howe e , none o he plan s in
which we ound ba en s a Pa acou we e e ile, so
diagnos ic ep oduc i e cha ac e s could no be de e -
mined.
se ed o he species o ba s in en s p e i-
ously ound occupied by R.pumilio. Fo his
eason, and because he delica e an e io den-
i ion o Rhinophylla seems inadequa e o he
ask o chewing h ough he ough la e al
eins o As oca yum lea es, we a o he
hypo hesis sugges ed by Cha les-Dominique
(1993) ha R.pumilio is a oos pa asi e ha
uses en s made by o he ba s.
In addi ion o inding oos s o Rhinophyl-
la pumilio in bi id As oca yum lea - en s, we
dis u bed one g oup o h ee indi iduals om
an uniden i ied loca ion in a clump o Phen-
akospe mum guyannense (S eli ziaceae) ha
con ained an ‘‘apical’’ en (Kunz e al.,
1994) simila in cons uc ion o hose in
which we ound U ode ma biloba um a o h-
e si es (see ig. 51). On ano he occasion we
ound an adul male R.pumilio oos ing
alone benea h an unmodi ied lea o P.guy-
annense.
O he ou As oca yum lea - en s in
which we ound Rhinophylla pumilio, wo
we e in well-d ained p ima y o es , one was
in closed-canopy seconda y g ow h, and one
was in p ima y o es a he edge o a
swampy a ea. The wo Phenakospe mum
oos s we e bo h in swampy p ima y o es .
None o he oos ing g oups we ound a Pa -
acou ( able 36) con ained mo e han one
adul male.
1998 97SIMMONS AND VOSS: PARACOU BATS
STENODERMATINAE
Al hough o he Amazonian si es a e
known o ha e mo e han 20 sympa ic spe-
cies o S enode ma inae, we cap u ed only 15
species o his phyllos omid sub amily a
Pa acou. Following he axonomic usage ec-
ommended by Pe ley e al. (
MS
), he Pa acou
species ep esen eigh gene a: Ame ida,A -
ibeus (including De manu a and Koopman-
ia), Chi ode ma,Ec ophylla (including Me-
sophylla), Pla y hinus ( o me ly Vampy-
ops), S u ni a,U ode ma, and Vampy essa.
Collec ions om o he locali ies in F ench
Guiana and Su inam include an addi ional
se en s enode ma ine species (appendix 1),
bu o easons explained below (see Anal-
yses o Sampling: Es ima ing Comple eness)
we doub ha any o hese no mally occu
wi hin ou s udy a ea.
Ame ida cen u io G ay
V
OUCHER
M
ATERIAL
: 7 emales (AMNH
*267973, *267274, *267275, *267276,
*267278; MNHN *1995.1037, *1995.1038)
and 3 males (AMNH *267279, *267976;
MNHN *1995.1039); see able 37 o mea-
su emen s.
I
DENTIFICATION
: Pe e son’s (1965a) e i-
98 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
sion emains he p incipal e e ence on Ame-
ida cen u io. Measu emen s o specimens
p e iously collec ed in he Guianas can also
be ound in Husson (1962, 1978), Swanepoel
and Genoways (1979), and B osse and
Cha les-Dominique (1990). No subspecies
a e cu en ly ecognized (Pe e son, 1965a;
Jones and Ca e , 1976; Koopman, 1994).
Pa acou specimens o Ame ida cen u io
con o m in all espec s o Pe e son’s (1965a)
accoun o his species, wi h measu emen s
alling wi hin he ange o a ia ion p e i-
ously epo ed in he li e a u e. In pa icula ,
mo phome ic da a om ou se ies ( able 37)
p o ide compelling con i ma ion o he e-
ma kable sexual size dimo phism o A.cen-
u io, in which emales a e subs an ially la g-
e han males. Measu emen s o Pa acou
males and emales a e comple ely nono e -
lapping in 11 o 13 dimensions, wi h sligh
o e lap occu ing only in hind oo leng h and
leng h o ea .
F
IELD
O
BSERVATIONS
: O he 10 indi iduals
o Ame ida cen u io cap u ed a Pa acou, 6
we e aken in g ound-le el mis ne s and 4 in
ele a ed mis ne s. O he six g ound-le el
cap u es, wo we e in swampy p ima y o -
es , one was in c eekside p ima y o es , and
h ee we e in manmade clea ings. Two indi-
iduals we e cap u ed be ween 5 and 20 m
o e a na ow di oad, and wo we e cap-
u ed 34–37 m abo e a ee all clea ing in
well-d ained p ima y o es .
A ibeus
Reliable species iden i ica ions wi hin he
genus A ibeus as ecognized he ein (includ-
ing De manu a and Koopmania) equi es
e e ence o Handley (1987) and Ma ques-
Aguia (1994). Al hough Lim and Wilson’s
(1993) concep o A ibeus jamaicensis di -
e s om ou s, hei keys a e also use ul o
dis inguishing he la ge species o A ibeus
(subgenus A ibeus) in no he n Sou h Ame -
ica.
A ibeus (A ibeus)jamaicensis Leach
Figu es 38, 39
V
OUCHER
M
ATERIAL
: 23 emales (AMNH
*266322, *266331, *266333, *266334,
*266335, *266336, *266337, *266338,
*266344, *267998, *268503, *268505,
*268508, *268528, *268529; MNHN
*1995.1139, *1995.1140, *1995.1141,
*1995.1142, *1995.1143, *1995.1144,
*1995.1145, *1995.1146) and 12 males
(AMNH *266321, *266332, *266341,
*266345, *267202, *267999, *268502,
*268504; MNHN *1995.1147, *1995.1148,
*1995.1149, *1995.1150); see able 38 o
measu emen s.
I
DENTIFICATION
: We ollow Handley (1987)
and Ma ques-Aguia (1994) in ega ding A -
ibeus jamaicensis as he senio synomym o
A.plani os is (con a Lim and Wilson
[1993] and Koopman [1993, 1994]). De-
sc ip ions and compa a i e measu emen s
can be ound in Goodwin and G eenhall
(1961), Swanepoel and Genoways (1979),
Koepcke and K a (1984), Handley (1987),
B osse and Cha les-Dominique (1990), Lim
and Wilson (1993), and Ma ques-Aguia
(1994). Al hough Husson (1962, 1978) p o-
ided a de ailed accoun o ‘‘A ibeus li u -
a us allax’’ (
5
A.jamaicensis) om Su i-
nam, his sample may ha e included speci-
mens o bo h A.jamaicensis and ue A.li-
u a us.
Be ween 10 and 13 subspecies o A ibeus
jamaicensis (including plani os is) a e cu -
en ly ecognized (see Hall, 1981; Da is,
1984; Handley, 1987; Ma ques-Aguia , 1994;
Koopman, 1994), o which 5 occu in Sou h
Ame ica: A.j.aequa o ialis (Paci ic slope o
he Andes om sou he n Colombia o no h-
e n Pe u), A.j.he cules (eas e n Ecuado and
Pe u), A.j. ini a is (cen al Colombia and
Venezuela no h o he O inoco), A.j. allax
(Venezuela sou h o he O inoco h oughou
he Guianas, hence sou hwa d h ough cen-
al B azil o Boli ia), and A.j.plani os is
(eas e n B azil and Pa aguay).
Ou Pa acou ma e ial con o ms closely o
published quali a i e desc ip ions o A ibeus
jamaicensis allax, al hough we obse ed
somewha mo e a ia ion in pelage colo
han p e iously desc ibed o Guianan spec-
imens (see below). Measu emen s o ou ma-
e ial all wi hin he ange o a ia ion p e-
iously epo ed by mos au ho s o A.j. al-
lax, including specimens e e ed o A.plan-
i os is by Lim and Wilson (1993).
Despi e ou bes e o s o co ec ly iden-
i y la ge A ibeus in he ield, subsequen
museum s udy showed ha we ini ially mis-
1998 99SIMMONS AND VOSS: PARACOU BATS
Fig. 38. Do sal (A) and en al (B) iews o he skull o A ibeus jamaicensis (AMNH 266321;
male) om Pa acou. Do sal (C) and en al (D) iews o he skull o A.li u a us (AMNH 267492;
male) om Pa acou. No e ha jamaicensis has a b oad in e o bi al egion, small pos o bi al p ocesses,
and h ee mola s, whe eas li u a us has a na owe in e o bi , be e de eloped pos o bi al p ocesses,
and lacks M3. Scale ba s
5
10 mm.
100 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 39. La e al iews o he skull and lowe jaw o (A)A ibeus jamaicensis (AMNH 266321;
male) and (B)A.li u a us (AMNH 267492; male). Scale ba s
5
10 mm.

1998 101SIMMONS AND VOSS: PARACOU BATS
den i ied 6 specimens (6%) ou o 94 adul s
p ese ed as ouche s. While we ne e con-
used specimens o li u a us and obscu us,
we ini ially misiden i ied one specimen o ja-
maicensis as li u a us, wo specimens o ja-
maicensis as obscu us, and h ee specimens
o obscu us as jamaicensis. All o ou iden-
i ica ion e o s in ol ed emales. Mos o
ou iden i ica ion mis akes in ol ed indi id-
uals somewha la ge o smalle han he spe-
cies no m, o hose wi h ambiguous pelage
colo s and acial ma kings. Fo example,
mos specimens o li u a us ha e much
b igh e acial s ipes han hose ypical o
jamaicensis, bu we ound some jamaicensis
wi h b igh s ipes and some li u a us wi h
pale s ipes. We also ound in aspeci ic a i-
a ion in do sal pelage colo in all h ee spe-
cies, wi h some indi iduals ha ing blacke o
b owne u han ha epo ed as ypical in
he li e a u e.
Unambiguous iden i ica ions o ou ouch-
e ma e ial we e ul ima ely based on he
quali a i e c anioden al cha ac e s desc ibed
by Handley (1987), B osse and Cha les-
Dominique (1990), and Ma ques-Aguia
(1994). In pa icula , Pa acou specimens o
A ibeus jamaicensis a e uni o mly dis in-
guished by a b oad pos o bi al egion wi h
poo ly de eloped pos o bi al p ocesses ( ig.
38), and M3 is always p esen . By con as ,
he pos o bi al egion is always na owe ( a-
ble 38) and he pos o bi al p ocesses a e well
de eloped in ou specimens o A.li u a us,
which uni o mly lack M3. Subsequen ex-
amina ion o ou measu emen da a ( able
38) showed ha whe eas mean alues o ex-
e nal dimensions o jamaicensis a e smalle
han hose o li u a us, hese species ha e
o e lapping anges o mo phome ic a ia-
ion a Pa acou.
Ma ques-Aguia (1994) ci ed wo pelage
cha ac e s as pa icula ly use ul o dis in-
guishing A ibeus jamaicensis om A.li u -
a us: (1) he do sal su ace o he base o he
o ea m is e y spa sely u ed (almos na-
ked) in jamaicensis e sus densely u ed in
li u a us; and (2) he en al u is os ed
(wi h whi e o pale g ay) in adul jamaicen-
sis, whe eas li u a us has comple ely da k
(un os ed) en al u . Examina ion o ou
ouche s con i ms ha hese ai s a e un-
ambiguously diagnos ic o jamaicensis and
li u a us a Pa acou. Un o una ely, we we e
no awa e o hese help ul iden i ica ion c i-
e ia in he ield.
Nume ous c anial cha ac e s ci ed by
Handley (1989) consis en ly sepa a e A i-
beus jamaicensis and A.obscu us, bu hese
a e ob iously no use ul in he ield. Ou
mo phome ic da a, howe e , indica e ha ja-
maicensis and obscu us can be sepa a ed un-
ambiguously (a leas in F ench Guiana) on
he basis o body weigh , o al leng h, and
o ea m leng h, in all o which dimensions
obscu us is subs an ially smalle . As no ed
by Handley (1989), jamaicensis has sho e
u han obscu us, bu a Pa acou his di e -
ence is sligh (7 mm e sus 8–9 mm) and
equi es ca e ul measu emen o be use ul o
iden i ica ion. Bo h axa, in ou judgmen ,
ha e equally ‘‘so ’’ u , con a Handley’s ob-
se a ions. Do sal u colo is ypically much
da ke in obscu us han in jamaicensis, bu
(as p e iously no ed) we ound enough o e -
lapping a ia ion in his cha ac e o comp o-
mise i s use ulness in he ield. Handley
(1989: 450) also no ed ha obscu us has
‘‘ ewe and smalle o namen al wa s on
[ he] chin,’’ bu we did no ind his o be
consis en ly ue in ou ma e ial. Mos indi-
iduals o bo h species ha e he same num-
be o chin papillae (nine small papillae a -
anged in a ‘‘U’’ a ound a la ge cen al pa-
pilla), wi h conside able a ia ion in papil-
la y size.
As no ed abo e, we suspec ha Husson’s
(1962, 1978) sample o ‘‘A ibeus li u a us
allax’’ was a composi e o indi iduals p op-
e ly e e ed o jamaicensis and li u a us.
This conclusion is based on wo obse a-
ions. Fi s , Husson (1962, 1978) epo ed
ha M3 was p esen in 26 o his 34 speci-
mens, and absen in 7 specimens. By con-
as , we ound M3 o be uni o mly p esen
in jamaicensis and uni o mly absen in li u -
a us, a pa e n ha was also obse ed by
B osse and Cha les-Dominique (1990). Sec-
ond, Husson (1962: able XX) p o ided mea-
su emen s o 10 indi iduals, 9 o which all
wi hin he ange o a ia ion ha we ob-
se ed o jamaicensis, bu 1 o which does
no . The la e indi idual (a emale om he
S u ga museum, SNM 686.1) was epo ed
o ha e a pos o bi al b ead h o 6.3 mm, a
alue alling well below he ange o a ia-
102 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
ion in ou se ies o jamaicensis, bu ag eeing
pe ec ly wi h ou measu emen da a o li-
u a us.
F
IELD
O
BSERVATIONS
: We eco ded 73 cap-
u es (possibly including some ecap u es) o
A ibeus jamaicensis a Pa acou, o which 71
we e in g ound-le el mis ne s and 2 we e in
ele a ed mis ne s. The 71 g ound-le el cap-
u es included 18 in well-d ained p ima y
o es , 38 in swampy p ima y o es , and 15
in c eekside p ima y o es . The wo ele a ed
ne cap u es we e made 5–10 m abo e he
g ound in he subcanopy o swampy p ima y
o es .
A ibeus (A ibeus)li u a us (Ol e s)
Figu es 38, 39
V
OUCHER
M
ATERIAL
: 15 emales (AMNH
*267204, *267495, *267496, *267497,
*268506, *268507, *268509, *268510,
*268512, *268513; MNHN *1995.1151,
*1995.1152, *1995.1153, *1995.1154,
*1995.1155) and 10 males (AMNH
*266346, *267206, *267492, *268511,
*268514, *268515; MNHN *1995.1156,
*1995.1157, *1995.1158, *1995.1159); see
able 38 o measu emen s.
I
DENTIFICATION
: Cha ac e s use ul o sep-
a a ing A ibeus li u a us om A.jamaicen-
sis a Pa acou a e discussed in he p eceding
species accoun . Desc ip ions and compa a-
i e measu emen s o A.li u a us can also be
ound in Goodwin and G eenhall (1961), Hill
(1964), Tamsi and Valdi ieso (1966),
Swanepoel and Genoways (1979), Da is
(1984), Koepcke and K a (1984), Handley
(1987), B osse and Cha les-Dominique
(1990), Lim and Wilson (1993), and Ma -
ques-Aguia (1994). As no ed ea lie , Hus-
son’s (1962, 1978) accoun o ‘‘A ibeus li-
u a us allax’’ was appa en ly based on a
composi e se ies o A.li u a us and A.ja-
maicensis.
Two subspecies o A ibeus li u a us a e
gene ally ecognized: A.l.palma um (Cen-
al Ame ica o no hwes e n Pe u, no he n
Colombia, no he n Venezuela, T inidad o
S . Vincen ) and A.l.li u a us ( h oughou
Amazonia o no he n A gen ina) (Da is,
1984; Koopman, 1994; Ma ques-Aguia ,
1994). A ibeus in e medius, conside ed a
dis inc species by many au ho s (e.g., Da is,
1984; Koopman, 1993, 1994), may ep esen
a subspecies o A.li u a us endemic o Mex-
ico and Cen al Ame ica (Ma ques-Aguia ,
1994).
Ou ouche ma e ial con o ms closely
wi h mos p e ious quali a i e desc ip ions
o A ibeus li u a us (excep Husson’s) in he
li e a u e ci ed abo e. In pa icula , measu e-
men s o ou specimens all wi hin he ange
o a ia ion p e iously epo ed o A.l.li-
u a us in he Guianas and elsewhe e in
no he n Sou h Ame ica.
F
IELD
O
BSERVATIONS
: We eco ded 53 cap-
u es (possibly including some ecap u es) o
A ibeus li u a us a Pa acou, o which 40
we e in g ound-le el mis ne s and 12 we e
in ele a ed mis ne s; in addi ion, 1 indi idual
was sho a nigh . O he 40 g ound-le el
mis ne cap u es, 17 we e in well-d ained p i-
ma y o es , 8 we e in swampy p ima y o -
es , 3 we e in c eekside p ima y o es , 3
we e in manmade clea ings, and 9 we e o e
oadside puddles. O he 12 indi iduals cap-
u ed in ele a ed ne s, 11 we e aken be ween
10 and 23 m abo e a na ow di oad, and
1 was aken a 5–8 m in he subcanopy o
swampy p ima y o es . The sho ba was
ound hanging om a palm ond abou 10
m abo e he g ound in well-d ained p ima y
o es .
A ibeus (A ibeus)obscu us (Schinz)
V
OUCHER
M
ATERIAL
: 27 emales (AMNH
*266271, *266273, *266279, *266281,
*266287, *267997, *268000, *268501,
*268516, *268518, *268520, 268522,
*268523, *268524, *268525, *268526,
*268527; MNHN *1995.1160, *1995.1161,
*1995.1162, *1995.1163, *1995.1164,
*1995.1165, *1995.1166, *1995.1167,
*1995.1168, *1995.1169) and 10 males
(AMNH *266272, *266286, *266288,
*267208, *267210, *268517, 268519;
MNHN *1995.1170, *1995.1171, *1995.1172);
see able 38 o measu emen s.
I
DENTIFICATION
: We ollow Handley (1989)
in using he name A ibeus obscu us ins ead
o A. uliginosus o he smalles and da kes
membe o he subgenus A ibeus ound
h oughou he we Sou h Ame ican lowlands
eas o he Andes. A ibeus obscu us is bes
iden i ied by e e ence o Handley (1987,
1998 103SIMMONS AND VOSS: PARACOU BATS
1989), Lim and Wilson (1993), and Ma ques-
Aguia (1994). Addi ional desc ip ions and
compa a i e measu emen s can also be ound
(unde he name A. uliginosus) in Swanepoel
and Genoways (1979), Koepcke and K a
(1984), and B osse and Cha les-Dominique
(1990). No subspecies a e cu en ly ecog-
nized (Handley, 1987, 1989; Ma ques-
Aguia , 1994).
Al hough ou ouche ma e ial gene ally
ag ees wi h p e ious quali a i e desc ip ions
o A ibeus obscu us, h ee o ou specimens
lack M3 comple ely, a polymo phism no ed
by Handley (1989) and Ma ques-Aguia
(1994), bu no by Handley (1987) o Lim
and Wilson (1993). Measu emen s o he
Pa acou se ies likewise all wi hin he known
ange o a ia ion o his species wi h he
excep ion o ou la ges specimens, which
ha e sligh ly longe o ea ms han p e iously
epo ed.
F
IELD
O
BSERVATIONS
: We eco ded 117
cap u es (p obably including some ecap-
u es) o A ibeus obscu us a Pa acou, o
which 104 we e in g ound-le el mis ne s, 7
we e in ele a ed mis ne s, and 6 we e a
104 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
oos s. O he 104 g ound-le el mis ne cap-
u es, 42 we e in well-d ained p ima y o es ,
37 we e in swampy p ima y o es , 19 we e
in c eekside p ima y o es , 4 we e in man-
made clea ings, 1 was in closed-canopy sec-
onda y g ow h, and 1 was o e a oadside
puddle. O he se en ele a ed mis ne cap-
u es, i e we e made be ween 7 and 20 m
abo e a na ow di oad, one was 10 m
abo e a ee all in well-d ained p ima y o -
es , and one was 5–8 m abo e he g ound in
he subcanopy o swampy p ima y o es .
We ound h ee oos ing g oups o A i-
beus obscu us unde ex olia ing pieces o
ba k 6–7 m abo e he g ound on he unks
o g ignon ees, Oco ea ub a (Lau aceae),
in well-d ained p ima y o es ( ig. 40). One
o hese g oups consis ed o an adul emale
wi h a nu sing ju enile; ano he consis ed o
an adul emale, a nu sing ju enile, and an
escaped adul o unknown sex; and he hi d
was a soli a y nea - e m p egnan emale. We
also ound a soli a y adul male A ibeus ob-
scu us oos ing benea h an unmodi ied lea
o Phenakospe mum guyannense (S eli zia-
ceae) abou 4 m abo e he g ound a he edge
o a clea ing in seconda y ege a ion.
A ibeus (De manu a)cine eus (Ge ais)
Figu es 41, 42
V
OUCHER
M
ATERIAL
: 9 emales (AMNH
*266259, *266265, *266266, *266270,
*267499, *267991; MNHN *1995.1108,
*1995.1109, *1995.1110) and 16 males
(AMNH *266260, *266261, *266290,
*266291, *266302, *266306, *266307,
*266320, *267196, *267978, *267980;
MNHN *1995.1111, *1995.1112, *1995.1113,
*1995.1114, *1995.1115); see able 39 o
measu emen s.
I
DENTIFICATION
: The mos ecen e ision
o he smalle species o A ibeus (subgenus
De manu a) is Handley’s (1987), whose me-
iculous species compa isons and key we e
ou p ima y esou ce o iden i ica ion. Ad-
di ional compa a i e measu emen s o A.ci-
ne eus and A.gnomus om F ench Guiana
p o ided by B osse and Cha les-Dominique
(1990) we e also help ul. Al hough bo h spe-
cies p obably occu in Su inam and may
ha e been mixed in ea lie collec ions, Hus-
son’s (1962, 1978) accoun o A.cine eus
seems o ha e been based en i ely upon spec-
imens p ope ly e e ed o ha species.
Koopman (1994) lis ed se en subspecies o
A.cine eus, howe e all bu wo o hese ap-
pea o ep esen o he species (see Handley,
1987, and below).
Ou ouche ma e ial, one o he la ges
se ies o A ibeus cine eus epo ed om a
single locali y, con o ms in all espec s o
Handley’s (1987) desc ip ion o he species.
Al hough ou specimens gene ally all wi hin
he ange o mo phome ic a ia ion p e i-
ously epo ed in he li e a u e, a ew indi-
iduals o bo h sexes a e sligh ly smalle in
some dimensions han hose documen ed
om he Guianas by Husson (1962, 1978)
and B osse and Cha les-Dominique (1990).
As wi h he la ge species o A ibeus
(subgenus A ibeus), we ound ha we could
no always dis inguish species o De manu a
in he ield. Mos indi iduals o cine eus a e
la ge han mos specimens o gnomus ( able
39), and mos cine eus ha e c eam-colo ed
ea ma gins and agus while hose s uc u es
a e b igh yellow in mos gnomus. Howe e ,
we cap u ed some indi iduals ha we ini-
ially misiden i ied using hese cha ac e s. O
67 collec ed specimens o De manu a,65
(97%) we e co ec ly iden i ied in he ield;
wo indi iduals (3%) we e ini ially iden i ied
inco ec ly, bo h cases in ol ing specimens
o A.cine eus (one male and one emale)
misiden i ied as A.gnomus. We had o clean
he skulls o hese and se e al o he speci-
mens ha ell in he zone o size o e lap be-
ween A.cine eus and A.gnomus (see able
39) in o de o de e mine hei co ec iden-
i ica ions.
We a e unawa e o any ex e nal cha ac e s
ha allow unequi ocal iden i ica ion o A -
ibeus cine eus and A.gnomus whe e hese
axa occu in sympa y, bu he numbe o
lowe mola s appea s o be eliable in ou
Pa acou sample: all indi iduals wi h wo
lowe mola s p esen on bo h sides p o ed o
be A.cine eus upon subsequen examina ion,
and all indi iduals wi h h ee lowe mola s
on bo h sides we e A.gnomus (one indi id-
ual o cine eus had wo lowe mola s on one
side and h ee on he o he ). Al hough he
numbe o lowe mola s is pe haps he bes
single cha ac e o dis inguishing hese axa
in he ield, i is no e y use ul o nonde-
1998 111SIMMONS AND VOSS: PARACOU BATS
Fig. 45. De ail o he lea - en oos o A ibeus cine eus shown in igu e 43. This is he lowe
su ace o he lea , which is conspicuously whi ish by con as wi h he da k g een uppe su ace ( acing
page). The long black spines, g ouped in clus e s all along he mid ib, had been chewed o sho s ubs
o he dis almos 15 cm o so, nea he apex o he en whe e he ba s we e hanging (a ow).

112 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
*1995.1137, *1995.1138); see able 39 o
measu emen s.
I
DENTIFICATION
: The o iginal desc ip ion o
A ibeus gnomus by Handley (1987) is s ill
he basic e e ence o his species, al hough
he measu emen s abula ed by B osse and
Cha les-Dominique (1990) o hei F ench
Guianan ma e ial a e also help ul. The ew
p oblems we encoun e ed in dis inguishing
A.gnomus om A.cine eus we e discussed
in he p eceeding species accoun . No sub-
species o A ibeus gnomus a e ecognized
(Handley, 1987).
Ou ouche ma e ial co esponds closely
wi h Handley’s (1987) desc ip ion o A i-
beus gnomus, and measu emen s o ou spec-
imens gene ally all wi hin he ange o a i-
a ion p e iously epo ed o he species. One
excep ion is a pa icula ly la ge male
(AMNH 267987), whose measu emen s in
se e al dimensions exceed any p e iously e-
po ed in he li e a u e (e.g., leng h o max-
illa y oo h ow, 6.08 mm; b ead h ac oss mo-
la s, 8.30 mm; zygoma ic b ead h, 11.47
mm). Excep in size, howe e , his specimen
ag ees mo phologically wi h he emainde
o ou specimens, and we conclude ha i
simply ep esen s an unusually la ge exam-
ple. Unpublished measu emen da a om
o he locali ies (in Venezuela, Guyana, Pe u,
and B azil; Handley, pe sonal commun.) in-
dica e ha AMNH 267987 is no unique in
his espec , and ha he no mal ange o size
a ia ion in A.gnomus is somewha g ea e
han ha o iginally epo ed by Handley
(1987).
F
IELD
O
BSERVATIONS
: We eco ded 59 cap-
u es (possibly including some ecap u es) o
A ibeus gnomus a Pa acou: 51 in g ound-
le el mis ne s, 4 in ele a ed mis ne s, and an-
o he 4 a oos s. O he 51 g ound-le el
mis ne cap u es, 6 we e in well-d ained p i-
ma y o es , 17 we e in swampy p ima y o -
es , 5 we e in c eekside p ima y o es , 22
we e in manmade clea ings, and 1 was in
closed-canopy seconda y g ow h. I is no e-
wo hy ha o he 22 g ound-le el mis ne
cap u es in manmade clea ings, 18 we e
aken in ne s e ec ed a ound small ui ing
ees, Vismia sp. (Clusiaceae), g owing along
a na ow oad h ough well-d ained p ima y
o es ; none we e made in la ge clea ings a
emo ed om p ima y o es . O he ele a -
ed mis ne cap u es, wo we e made be ween
4 and 21 m abo e a na ow di oad and wo
we e made a 7–10 m in he subcanopy o
swampy p ima y o es .
We ound ou oos ing g oups o A ibeus
gnomus, all o which occupied lea - en s.
Two la ge oos ing g oups (one wi h 4–8 in-
di iduals, he o he wi h pe haps 6–10) each
inhabi ed a single modi ied lea o he gian
he b Phenakospe mum guyannense (S eli -
ziaceae); bo h oos s we e ha d o see, abou
3 m abo e he g ound in he densely clu -
e ed unde s o y o swampy p ima y o es
( ig. 46). We cap u ed (and p ese ed as
ouche s) only a single adul emale om
each o hese oos ing g oups, he age and
sex composi ion o which a e o he wise un-
known.
The o he wo oos s, inhabi ed by soli a y
adul males, we e ‘‘apical’’ en s (sensu Kunz
e al., 1994) made om he spade-shaped
lea es o epiphy ic Philodend on sp. (A a-
ceae). One o hese oos s was abou 5 m
abo e he g ound in well-d ained dis u bed
o es , he o he abou 10 m abo e he
g ound in well-d ained p ima y o es . We
ound many epiphy ic Philodend on plan s
wi h simila ly modi ied lea es h oughou
ou s udy a ea, bu only hese wo examples
we e occupied by ba s (wasp nes s we e
ound unde he modi ied lea es o a ew
plan s).
Al hough A ibeus gnomus and A.cine eus
a e mo phologically simila , ou da a sugges
hey a e ecologically di e gen . Consis en
wi h B osse and Cha les-Dominique’s
(1990) gene aliza ions conce ning habi a use
in hese species, an analysis o g ound-le el
cap u e equency da a om Pa acou ( able
40) sugges s ha A.gnomus is mo e abun-
dan in p ima y o es han A.cine eus,
which appa en ly a o s he ea ly-succes-
sional ege a ion o modi ied habi a s.
Whe he o no A.cine eus consis en ly in-
habi s bi id palm-lea en s whe eas A.gno-
mus uses di e en ly designed en s made
om he lea es o la ge he bs is unknown,
bu his hypo he ical di e ence could easily
be es ed by ollowing he mo emen s o
posi i ely iden i ied ba s i ed wi h adio
ansmi e s. Because A.gnomus was only e-
cen ly ecognized as axonomically dis inc
om A.cine eus, and because hey a e ha d
1998 113SIMMONS AND VOSS: PARACOU BATS
Fig. 46. Roos o A ibeus gnomus in swampy p ima y o es a Pa acou. As many as eigh indi iduals
may ha e inhabi ed he da k apex o his lea - en (a ow), bu i was impossible o ob ain a clea iew o
make an accu a e coun . Phenakospe mum guyannense (S eli ziaceae) is mode a ely common in swampy
p ima y o es and seconda y ege a ion a Pa acou, whe e en - oos s made om i s la ge lea es a e also
inhabi ed by U ode ma biloba um ( ig. 51). This en was made by cu s in he la e al eins and in e s i ial
issue along bo h sides o he mid ib causing he sides o he lea o d oop (as in he ‘‘boa ’’ en s desc ibed
and illus a ed by Kunz e al., 1994), and by a deep cu h ough he mid ib, causing he apex o he lea o
hang s aigh down (a cha ac e is ic o ‘‘apical’’ en s desc ibed by he same au ho s). A simila oos
inhabi ed by ano he la ge g oup o A.gnomus was ound se e al kilome e s om his si e.
114 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
(bu no impossible) o dis inguish in he
ield, un ouche ed ecological obse a ions
epo ed in he li e a u e o hese species
should be ega ded wi h cau ion.
A ibeus (Koopmania)concolo Pe e s
V
OUCHER
M
ATERIAL
: 17 emales (AMNH
*266267, *266269, *267192, *267194,
*267476, *267478, *267479, *267487,
*267488, *267981, *267982, *267983;
MNHN *1995.1116, *1995.1117, *1995.1118,
*1995.1119, *1995.1120) and 7 males
(AMNH *267193, *267195, *267477,
*267502; MNHN *1995.1121, *1995.1122,
*1995.1123); see able 41 o measu emen s.
I
DENTIFICATION
: Fea u es use ul o iden i-
ying A ibeus concolo we e summa ized by
Handley (1987) and Acos a and Owen
(1993); addi ional desc ip ions and measu e-
men s can be ound in Husson (1962, 1978),
Hill (1964), Ba iga-Bonilla (1965), Lina es
(1969), Genoways and Williams (1979),
Swanepoel and Genoways (1979), and B os-
se and Cha les-Dominique (1990). In con-
as o he si ua ion wi h mos o he species
o A ibeus, no axonomic p oblems a e ap-
pa en ly associa ed wi h A.concolo ,so
iden i ica ion is ela i ely s aigh o wa d.
No subspecies a e cu en ly ecognized
(Handley, 1987; Acos a and Owen, 1993;
Koopman, 1994).
Ou ouche ma e ial, appa en ly he la g-
es se ies a ailable om any single locali y,
con o ms closely wi h p e ious quali a i e
and mo phome ic desc ip ions o A ibeus
concolo . In pa icula , measu emen s o ou
specimens con i m B osse and Cha les-
Dominique’s (1990) epo o conside able
size a ia ion wi hin F ench Guianan popu-
la ions o his species. The size ange among
ou specimens is e en g ea e han hey e-
po ed, ye we ound no e idence ha he
Pa acou sample includes mo e han one ax-
on.
1998 115SIMMONS AND VOSS: PARACOU BATS
F
IELD
O
BSERVATIONS
: We caugh 24 A i-
beus concolo a Pa acou, o which 22 we e
aken in g ound-le el mis ne s and 2 in ele-
a ed mis ne s. O he 22 g ound-le el cap-
u es, 3 we e in well-d ained p ima y o es ,
7 we e in swampy p ima y o es , 3 we e in
c eekside p ima y o es , 7 we e in manmade
clea ings, and 2 we e o e oadside puddles.
The wo ele a ed mis ne cap u es we e
made be ween 10 and 21 m o e a na ow
di oad.
Chi ode ma ini a um Goodwin
V
OUCHER
M
ATERIAL
: 8 emales (AMNH
*266255, *267189, *267473, *268531,
*269117; MNHN *1995.1191, *1995.1192,
*1995.1193) and 4 males (AMNH *266256,
*268532, *269118; MNHN *1995.1194);
see able 42 o measu emen s.
I
DENTIFICATION
: Desc ip ions and measu e-
men s o Chi ode ma ini a um om he
Guianas and elsewhe e can be ound in
Goodwin (1958), Goodwin and G eenhall
(1961), Ojas i and Lina es (1971), Ga dne
(1976), Genoways and Williams (1979),
Swanepoel and Genoways (1979), and B os-
se and Cha les-Dominique (1990). Al hough
subspecies o C. ini a um ha e been ec-
ognized by some au ho s (e.g., Jones and
Ca e , 1976), inc eased sampling h oughou
he ange o his species has demons a ed
mo e wi hin-popula ion a ia ion han p e-
iously suspec ed (see discussion in Williams
and Genoways, 1980a). Pending a ho ough
sys ema ic e iew, no inomial nomencla-
u e seems wa an ed (Koopman, 1994).
Ou Pa acou specimens con o m in all e-
spec s o p e ious quali a i e and mo pho-
me ic desc ip ions o Chi ode ma ini a-
um.
F
IELD
O
BSERVATIONS
: We caugh 13 Chi-
ode ma ini a um a Pa acou, o which 11
we e aken in g ound-le el mis ne s and 2 in
ele a ed mis ne s. Fou o he g ound-le el
cap u es we e made in swampy p ima y o -
es and he o he se en in manmade clea -
ings. The ele a ed mis ne cap u es we e
made a 17–20 m abo e a na ow di oad.
Chi ode ma illosum Pe e s
V
OUCHER
M
ATERIAL
: 3 emales (AMNH
*267191, *267474, *268536) and 5 males
(AMNH *267190, *267475, *268535;
MNHN *1995.1195, *1995.1196); see able
42 o measu emen s.
I
DENTIFICATION
: Desc ip ions and compa -
a i e measu emen s o Chi ode ma illosum
can be ound in Goodwin and G eenhall
116 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
(1961), Husson (1962, 1978), Hill (1964),
Genoways and Williams (1979), Swanepoel
and Genoways (1979), Hall (1981), B osse
and Cha les-Dominique (1990), and Ande -
son (1997). Two subspecies a e cu en ly ec-
ognized: C. .jesupi (Mexico o no he n
Colombia) and C. . illosum ( opical Sou h
Ame ica eas o he Andes om Colombia o
sou heas e n B azil, including T inidad and
Tobago) (Koopman, 1994).
Al hough ou Pa acou se ies o Chi ode -
ma illosum ag ees in quali a i e and quan-
i a i e cha ac e s wi h p e ious desc ip ions
o he species as a whole, c anioden al mea-
su emen compa isons indica e ha ou spec-
imens a e subs an ially smalle han mos o
hose p e iously epo ed om he Guianas.
Fo example, he obse ed ange in leng h o
he maxilla y oo h ow is 7.96–8.66 mm a
Pa acou e sus 8.6–10.2 mm a o he Guia-
nan locali ies om which measu ed speci-
mens a e epo ed in he li e a u e ci ed
abo e. An appa en ly indi idual anomaly is
ep esen ed by one o ou male specimens,
AMNH 268535, which has only one pai o
uppe and lowe inciso s; all o ou emain-
ing specimens ha e wo pai s in bo h jaws,
he no mal o mula o Chi ode ma (see
Koopman, 1994).
F
IELD
O
BSERVATIONS
: We caugh eigh
Chi ode ma illosum a Pa acou, o which
ou we e aken in g ound-le el mis ne s and
ou in ele a ed ne s. Two g ound-le el cap-
u es we e made in well-d ained p ima y o -
es , one in swampy p ima y o es , and one
in c eekside p ima y o es . Two specimens
we e cap u ed 34–37 m abo e a ee all
opening in well-d ained p ima y o es and
wo o he s we e ne ed 17–20 m abo e a na -
ow di oad.
Ec ophylla
Phylogene ic ela ionships be ween Ec o-
phylla,Mesophylla, and Vampy essa ha e
been he subjec o much deba e in he li -
e a u e. Mos wo ke s ha e e ained all h ee
as dis inc gene a (e.g., Hall, 1981; Koop-
man, 1993, 1994), bu some au ho s ha e
conside ed Mesophylla o be a junio syno-
mym o Vampy essa (e.g., Owen, 1987) o
o Ec ophylla (e.g., Goodwin and G eenhall,
1962; Handley, 1976). Recen ly, Pe ley e
al. (
MS
) ound s ong suppo o a sis e -
g oup ela ionship be ween Ec ophylla and
Mesophylla, bo h o which a e mono ypic,
and a gued ha Ec ophylla alba H. Allen
(1892) and Mesophylla macconnelli Thomas
(1901a) should be placed in a single genus
o e lec his ela ionship. We he e o e use
he olde gene ic name o bo h species, and
p o ide a o mal diagnosis o Ec ophylla as
so de ined.
E
MENDED
D
IAGNOSIS OF
E
CTOPHYLLA:
Size
small (weigh less han 11 g and o ea m
leng h less han 35 mm); do sal and en al
u pale bu , g ayish whi e, o whi e; no
whi e acial s ipes o middo sal s ipe; skin
o noselea , ea s, and humb b igh yellow;
en al bo de o na ial ho seshoe de ined by
a ee lap o skin; u opa agium sho , naked,
anslucen ; leng h o calca less han one-
hal leng h o hind oo ; den al o mula I 2/2,
C 2/2, P 2/2, M 2/2–3
3
2
5
28–30; os um
app oxima ely h ee- ou hs he leng h o he
b aincase; os um no in la ed and wi hou a
deep dep ession o long nasal ema gina ion;
in e p e ygoid space no ex ended by a deep
pala al ema gina ion; inne uppe inciso s
elonga e, unwo n c own heigh mo e han
wice ha o ou e inciso s; inne uppe in-
ciso s no deeply bi id; m1 wi hou pos e o-
lingual cusp (c own esembles ha o las
p emola ); lingual cusps o m2 es igial o
absen .
Ec ophylla macconnelli (Thomas)
V
OUCHER
M
ATERIAL
: 9 emales (AMNH
*267281, *267537, *267538, *267556,
*267558, *267559; MNHN *1995.1181,
*1995.1182, *1995.1183) and 4 males
(AMNH *267557, *267562, *268539;
MNHN *1995.1184); see able 43 o mea-
su emen s.
I
DENTIFICATION
: We consul ed desc ip ions
and measu emen s o Ec ophylla macconnelli
p o ided by Goodwin and G eenhall (1962),
Swanepoel and Genoways (1979), Williams
and Genoways (1980a), B osse and Cha les-
Dominique (1990), and Kunz and Pena
(1992). Two subspecies a e cu en ly ecog-
nized, o which he nomina e o m occu s
h oughou mos o he humid Neo opical
lowlands, including he Guianas (Koopman,
1994).

1998 117SIMMONS AND VOSS: PARACOU BATS
Ou Pa acou se ies con o ms in all e-
spec s o p e ious desc ip ions o Ec ophylla
macconnelli excep ha o B osse and
Cha les-Dominique (1990), who epo ed un-
usually high alues o leng h o he maxil-
la y oo h ow (6.7–7.4 mm) in hei F ench
Guianan ma e ial. By con as , he obse ed
ange o his measu emen in ou se ies
(5.61–6.09 mm) alls wi hin he ange p e-
iously epo ed by o he au ho s (5.5–6.6
mm). Because he o he ex e nal and c anio-
den al measu emen s epo ed by B osse and
Cha les-Dominique a e no abe an , we p e-
sume ha hei maxilla y oo h ow measu e-
men s we e e oneously epo ed.
F
IELD
O
BSERVATIONS
: We caugh 13 Ec o-
phylla macconnelli a Pa acou, o which only
1 was aken in a mis ne ; he emaining 12
we e caugh a oos s. Ou single mis ne
cap u e was a g ound le el in swampy p i-
ma y o es .
We ound h ee oos ing g oups o Ec o-
phylla macconnelli, all o which inhabi ed
lea - en s ( ig. 47) made om he bi id e -
minal lea le s o onds o young unde s o y
palms p o isionally iden i ied as As oca -
yum sciophilum. In all cons uc ion de ails
ha we no ed, hese en s exac ly esembled
hose desc ibed abo e in he accoun s o
Rhinophylla pumilio and A ibeus cine eus.
All h ee oos s we e in well-d ained p ima y
o es . One oos ing g oup o se en ba s, col-
lec ed in i s en i e y, consis ed o wo adul
males and i e adul emales. Ano he en i e
g oup o h ee consis ed o one adul male
and wo adul emales.
Ou oos obse a ions, oge he wi h
hose epo ed by Fos e (1992) and Cha les-
Dominique (1993), sugges ha Ec ophylla
macconnelli egula ly inhabi s en s manu-
ac u ed om he lea es o As oca yum
palms.
12
Because he unde su aces o As o-
ca yum lea es a e cha ac e is ically whi ish
(Hende son e al., 1995), we ag ee wi h
Hings on (1932) ha he unsually pale u o
12
Emmons (1990) was appa en ly he i s o epo
ha Ec ophylla macconnelli inhabi s palm-lea en s, bu
she did no iden i y he hos plan . Roos s ha e been
epo ed in he oliage o o he palms (and a oids), bu
mul iple independen obse a ions o occupied bi id
en s in As oca yum spp. om opposi e ends o Ama-
zonia sugges ha his genus is a o ed by Ec ophylla
macconnelli. Emmons’ (1990, 1997) sugges ion ha E.
macconnelli some imes inhabi s hollow ees was based
on Handley’s (1976: 30) epo o one indi idual ‘‘ ound
oos ing in a ee.’’ Howe e , he o iginal ield eco d
o he specimen in ques ion (USNM 405185) no es
‘‘sho in ee a e igh ened up‘‘— he ba ha ing p e-
sumably been dislodged om an unobse ed oos in he
unde g ow h. All o he published accoun s (including
Beebe [1925], Hings on [1932], Cha les-Dominique
[1993], and Kunz e al. [1994] in addi ion o e e ences
ci ed by Fos e [1992]) explici ly iden i y oliage as he
oos ing subs a e o E.macconnelli.
118 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 47. Roos o Ec ophylla macconnelli in well-d ained p ima y o es a Pa acou. Made om he
bi id e minal lea le o a young palm (As oca yum c . sciophilum), his shel e con ained h ee E.
macconnelli (one adul male and wo adul emales) hanging in a igh clus e om he mid ib abou
1.5 m abo e he g ound (a ow). No e he much spa se unde g ow h a his p ima y- o es si e han
ha su ounding an o he wise simila oos in seconda y ege a ion ( igu e 43).
1998 119SIMMONS AND VOSS: PARACOU BATS
E.macconnelli is co ela ed wi h he back-
g ound colo o i s ypical oos s, pe haps
making he ba s less conspicuous o diu nal
p eda o s.
Pla y hinus helle i (Pe e s)
V
OUCHER
M
ATERIAL
: 9 emales (AMNH
*266254, *267182, *267551, *267554,
*267555, *268540; MNHN *1995.1187,
*1995.1188, *1995.1189) and 5 males
(AMNH *267179, *267550, *268541,
*268542; MNHN *1995.1190); see able 44
o measu emen s.
I
DENTIFICATION
: A key o he species o
Pla y hinus was p o ided by Fe ell and
Wilson (1991), and Ande son (1996) help-
ully e iewed cha ac e s dis inguishing P.
helle i om P.b achycephalus (see also
Rouk and Ca e , 1972). O he use ul de-
sc ip ions and measu emen s o P.helle i can
be ound in Sanbo n (1955), Goodwin and
G eenhall (1961), Husson (1962, 1978),
Ga dne and Ca e (1972), Rouk and Ca e
(1972), Swanepoel and Genoways (1979),
B osse and Cha les-Dominique (1990), and
Ande son (1997). Two subspecies o P.hel-
le i a e cu en ly ecognized, o which P.h.
inca um anges h oughou Amazonia, in-
cluding he Guianas (Koopman, 1994).
Ou Pa acou specimens con o m closely
wi h p e ious desc ip ions o Pla y hinus
helle i. Al hough some popula ions (e.g., in
Boli ia; Ande son, 1996) exhibi a ia ion in
he numbe o accesso y cusps p esen on he
an e io ma gin o he second lowe p emo-
la , all o ou ouche s ha e only a single
accesso y cusp in his loca ion. Measu e-
men s o he Pa acou se ies ( able 44) all
wi hin he ange o a ia ion p e iously e-
po ed o P.h.inca um.
F
IELD
O
BSERVATIONS
: We cap u ed 14 in-
di iduals o Pla y hinus helle i a Pa acou,
o which 11 we e aken in g ound-le el mis -
ne s and 3 in ele a ed mis ne s. Fou g ound-
le el cap u es we e in well-d ained p ima y
o es , one in swampy p ima y o es , wo in
c eekside p ima y o es , wo in manmade
clea ings, and wo o e oadside puddles.
The h ee ele a ed ne cap u es we e made
be ween 6 and 21 m abo e a na ow di
oad.
S u ni a lilium (E. Geo oy)
Figu es 48–50
V
OUCHER
M
ATERIAL
: 19 emales (AMNH
*266210, *266226, *266231, *266234,
*266235, *266236, *267170, *268543,
*268546, *268549, *268552, *268553;
MNHN *1995.1197, *1995.1198, *1995.1199,
*1995.1200, *1998.600, *1998.601, *1998-
.602) and 35 males (AMNH *266199,
*266200, *266201, *266203, *266205,
120 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 48. Do sal (A) and en al (B) iews o he skull o S u ni a lilium (AMNH 266232; male)
om Pa acou. Do sal (C) and en al (D) iews o he skull o S. ildae (AMNH 267461; male) om
Pa acou. No e he species di e ences in inciso mo phology, size and shape o he mola s, and ela i e
b ead h o he mesop e ygoid ossa and mas oid egion. Scale ba s
5
10 mm.
1998 127SIMMONS AND VOSS: PARACOU BATS
Fig. 51. Lea - en oos o he ype inhabi ed by U ode ma biloba um, in a we opening in swampy
p ima y o es a Pa acou. This simple conical shel e , made by cu ing he mid ib o a Phenakospe mum
guyannense ond (a ow), was unoccupied when we ound i , bu an iden ical en nea by con ained
h ee ba s. Because i is di icul o app oach such oos s wi hou ala ming he inhabi an s, we ound
many emp y en s ha migh ha e con ained ba s only momen s be o e.

128 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
an eadings a e cu en in he li e a u e.
Thus, Williams and Genoways (1980a) s a ed
ha V.b ocki lacks a middo sal s ipe, ci ing
Pe e son as hei au ho i y, whe eas o he au-
ho s ha e e iden ly in e p e ed Pe e son o
mean ha a s ipe is p esen in his species
(e.g., Lewis and Wilson, 1987). Pe e son
(1968: 1) himsel clea ly s a ed ha he ho-
lo ype o b ocki is ‘‘g eyish b own abo e
wi h a ain indis inc do sal whi ish s ipe.’’
A diag amma ic cha ac e summa y accom-
panying he o iginal desc ip ion (op. ci .: ig.
6), howe e , does no dis inguish be ween he
ain middo sal s ipe o b ocki and he much
b igh e ma kings o wo o he congene s
(nymphaea and bidens).
Ou Pa acou ouche s ha e a middo sal
s ipe, bu i is ex emely ain and is isible
only when he d y do sal pelage is ca e ully
b ushed. The s ipe is ha d o see unde any
condi ions because i is only sligh ly pale
han he su ounding u , and i becomes
qui e in isible when he u is we (p eclud-
ing obse a ion in alcohol-p ese ed speci-
mens).
Because he do sal s ipe is ha d o see,
Vampy essa b ocki ex e nally esembles V.
pusilla, wi h which i migh be con used in
he ield. The la e species en i ely lacks a
do sal s ipe and sha es se e al ai s wi h
b ocki, including small size, wo pai s o
lowe inciso s, and absence o m3 (Pe e son,
1968). Cha ac e s ha unambiguously dis in-
guish b ocki and pusilla include a sui e o
c anioden al ea u es (op. ci .), o which he
mos use ul include: (1) o m o he inne
uppe inciso s ( ape ing o a poin in b ocki,
bilobed in pusilla), (2) shape o he an e io
lowe p emola (low-c owned and bladelike
in b ocki, wi h highe c own and spea like
an e io cusp in pusilla), and (3) shape o he
nasal ape u e (wi h s aigh en al bo de in
b ocki, V-shaped en al bo de in pusilla).
The la e cha ac e is isible only on cleaned
skulls, bu he dis inc ion is unambiguous
and p o ides a means o unequi ocally iden-
i ying old indi iduals wi h wo n o b oken
ee h.
F
IELD
O
BSERVATIONS
: O he se en Vam-
py essa b ocki we cap u ed a Pa acou, ou
we e aken in g ound-le el mis ne s and
h ee in ele a ed ne s. One o he g ound-
le el cap u es was in well-d ained p ima y
o es , wo we e in c eekside p ima y o es ,
and one was in a manmade clea ing. The el-
e a ed ne cap u es we e made 17–21 m o e
a na ow di oad.
FURIPTERIDAE
We cap u ed a single u ip e id species a
Pa acou, he only membe o i s amily
known o occu in ain o es habi a s.
1998 129SIMMONS AND VOSS: PARACOU BATS
Fu ip e us ho ens (F. Cu ie )
V
OUCHER
M
ATERIAL
: 9 emales (AMNH
*265975, *265979, 265980, *267213,
*267214, *268572, 268573; MNHN
*1995.870, *1995.871) and 4 males (AMNH
*267212, *267507; MNHN *1995.872,
1995.873); see able 49 o measu emen s.
I
DENTIFICATION
: Husson (1962, 1978) p o-
ided a de ailed desc ip ion and measu e-
men s o Fu ip e us ho ens, and addi ional
mo phome ic da a we e published by B os-
se and Cha les-Dominique (1990). No sub-
species a e ecognized (Koopman, 1994).
Al hough ou ouche ma e ial does no
di e in any espec om Husson’s (1962,
1978) ca e ul quali a i e desc ip ion o Fu-
ip e us ho ens, measu emen s o he Pa a-
cou se ies documen a g ea e ange o size
a ia ion han ha p e iously epo ed om
he Guianas. Ou measu emen da a ( able
49) addi ionally sugges ha he species may
be sexually dimo phic ( emales a e aging
sligh ly la ge han males in mos dimen-
sions), an obse a ion ha ends o co obo-
a e Uieda e al.’s (1980) epo o sexual di-
mo phism in a no heas e n B azilian popu-
la ion. In he o he wise excellen ex e nal de-
sc ip ions o Fu ip e us ho ens, Emmons
s a ed ha he humb has ‘‘no claw’’ (1990:
82) o ‘‘almos no claw’’ (1997: 91). The
humb, small and enclosed in he wing mem-
b ane, bea s a iny bu dis inc claw in all he
specimens we examined.
F
IELD
O
BSERVATIONS
: We collec ed 13
specimens o Fu ip e us ho ens a Pa acou,
o which 12 we e aken a oos s and 1 was
sho as i lew back and o h along a egula
bea abou a me e abo e he g ound in well-
d ained p ima y o es a nigh . The nine
oos ing g oups we ound consis ed o one o
wo indi iduals, ei he soli a y adul s (o
bo h sexes) o lac a ing emales wi h nu sing
ju eniles. In no case did we ind mo e han
one adul occupying a oos . All o he oos s
we ound we e in o unde allen ees in
a ious s ages o decomposi ion. One oos
was a small ca i y in he b oken end o a
o ing log ( ig. 52), bu mos we e on he
unde sides o unks o in da k chambe s be-
ween bu esses ( ig. 17). Fou oos s we e
in well-d ained p ima y o es , one was in
swampy p ima y o es , wo we e in p ima y
o es o un eco ded cha ac e , one was in se-
lec i ely logged o es , and one was in
closed-canopy seconda y g ow h. Small,
da k, soli a y ba s ha lew away uniden i-
ied om e ugia in o unde woody deb is
on many occasions h oughou he cou se o
ou ieldwo k a Pa acou we e p obably F.
ho ens. Al hough we ne e caugh his spe-
cies in mis ne s, ou imp ession was ha
oos s o F.ho ens could be ound by ca e-
ul sea ching almos anywhe e in he o es .
THYROPTERIDAE
Al hough we caugh only one hy op e id
species a Pa acou, a second is known om
130 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 52. Roos o Fu ip e us ho ens in well-d ained p ima y o es a Pa acou. An adul emale
wi h he nu sing o sp ing inhabi ed his da k bu shallow ca i y in he b oken end o a o ing log,
abou 40 cm abo e he g ound (a ow). Such inconspicuous e ugia occu e e ywhe e in he o es and
we p obably o e looked many F.ho ens oos s e en along equen ly a eled ails h ough ou s udy
a ea.
F ench Guiana and migh also be expec ed
o occu in ou s udy a ea (appendix 1).
Thy op e a icolo Spix
V
OUCHER
M
ATERIAL
: 16 emales (AMNH
*266348, *266356, *266358, 266359,
*266364, *267215, *267216, *267217,
*267218, *268576; MNHN *1995.874,
*1995.875, *1995.876, *1995.877,
1995.878, *1995.879) and 12 males (AMNH
*266352, 266355, 266357, *266361,
*266363, 266365, *268574, 268577; MNHN
1995.880, *1995.881, 1995.882, 1995.883);
see able 50 o measu emen s.
I
DENTIFICATION
: We consul ed he desc ip-
ions and measu emen s o Thypo e a icol-
o p o ided by Husson (1962, 1978), B osse
and Cha les-Dominique (1990), and Pine
(1993) o con i m he iden i ica ion o ou
ma e ial. Al hough h ee subspecies we e
ecognized by Wilson and Findley (1977)
and Koopman (1994), he mo phological jus-
i ica ion o a inomial classi ica ion is no
clea . Fu he mo e, Pine (1993) sugges ed
ha some published obse a ions o geo-
g aphic a ia ion wi hin T. icolo may ha e
been based on ma e ial ha was no co ec ly
iden i ied o species. Pending a ho ough e-
iew o he p oblem, i cu en ly seems
poin less o employ subspeci ic nomencla-
u e.
Ou ma e ial om Pa acou con o ms wi h
published desc ip ions o Thy op e a icolo
in all espec s. Al hough Pine (1993) no ed
some a ia ion in he numbe o lappe s on
he calca in some popula ions o his spe-
cies, all o ou specimens ha e wo lappe s
on he calca . The ee po ion o he ail is
ela i ely long in all ou luid-p ese ed ma-
e ial, which con o ms o Pine’s (1993) ob-
1998 131SIMMONS AND VOSS: PARACOU BATS
se a ions. We no e, howe e , ha he ee
po ion o he ail appea s qui e sho in ou
skins, an a i ac ha esul ed om pinning
he specimens o d y wi h he u opa agium
maximally ex ended.
The size ange documen ed by ou ouch-
e specimens ( able 50) is somewha g ea e
han ha p e iously epo ed om Su ina-
mese and F ench Guianan popula ions o
Thy op e a icolo by Husson (1962, 1978)
and B osse and Cha les-Dominique (1990).
In addi ion, he Pa acou measu emen da a
sugges some sligh sexual dimo phism, wi h
emales exceeding males in a e age body
weigh and o al leng h.
F
IELD
O
BSERVATIONS
: We eco ded 40 cap-
u es (possibly including some ecap u es) o
Thy op e a icolo a Pa acou, o which 3
we e in g ound-le el mis ne s, 1 was in an
ele a ed ne , and 36 we e a oos s. One
g ound-le el mis ne cap u e was in well-
d ained p ima y o es , one was in swampy
132 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 53. Roos o Thy op e a icolo in he hal -un olled new lea o a small Heliconia g owing in
swampy p ima y o es a Pa acou. The adhesi e sucke s o he oos ing ba s a e isible as da k spo s
h ough he anslucen issue o he lea ; he ba s hemsel es (an adul male and h ee adul emales)
o m a da k mass inside hei ubula shel e (a ow).

1998 133SIMMONS AND VOSS: PARACOU BATS
Fig. 54. Roos o Thy op e a icolo in a sc olled dead lea o Phenakospe mum guyannense (a ow)
in seconda y ege a ion a Pa acou. We ound wo o he oos s o T. icolo in dead lea es like his
one, which is hanging downwa d om i s b oken pe iole. This oos con ained abou ou ba s, one o
which was collec ed as a ouche .
134 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
p ima y o es , and one was in a manmade
clea ing. Ou single ele a ed mis ne cap u e
was made a 7–8 m abo e he g ound in he
subcanopy o swampy p ima y o es .
We ound 12 oos ing g oups o Thy op-
e a icolo , all o hem in oliage ( able 51).
Mos (nine) oos s we e in he e ec , hal -
un olled new lea es o heliconias, Heliconia
sp. (Heliconiaceae) ( ig. 53), o Phenako-
spe mum guyannense (S eli ziaceae), bu
h ee oos ing g oups occupied sc olled dead
lea es hanging om la ge Phenakospe mum
plan s ( ig. 54). O he i e Heliconia oos s
we ound, one was in well-d ained p ima y
o es , wo we e in swampy p ima y o es ,
and wo we e in c eekside p ima y o es .
Fi e Phenakospe mum oos s we e in sec-
onda y ege a ion (especially along he ma -
gins o a small sa anna encla e) and wo
we e in we glades su ounded by swampy
p ima y o es . All o he lea es ha we
ound used by T. icolo as oos s we e
shaded; none was in di ec sunligh . The
numbe o ba s pe oos a ied om one o
six wi h a well-de ined mode o ou . We did
no eco d mo e han a single adul male in
any oos .
VESPERTILIONIDAE
We cap u ed i e espe ilionid species a
Pa acou belonging o he gene a Ep esicus,
Lasiu us, and Myo is. O he six o he es-
pe ilionids known om F ench Guiana and
Su inam, ou could plausibly be expec ed o
occu in ou s udy a ea also (appendix 1).
Ep esicus chi iquinus Thomas
Figu es 55, 56
V
OUCHER
M
ATERIAL
: 2 emales (AMNH
*267531, *268584) and 4 males (AMNH
*267234, *267530; MNHN *1995.961,
1995.962); see able 52 o measu emen s.
I
DENTIFICATION
: In his e ision o he
Sou h Ame ican species o Ep esicus, Da is
(1966) ecognized an andinus g oup o spe-
cies wi h long blackish u . Se e al axa we e
included in his complex: andinus Allen
(1914), chi iquinus Thomas (1920b), inca
Thomas (1920b), mon osus Thomas (1920b)
and chi alensis An hony (1926). O hese,
Da is conside ed inca and chi iquinus o be
s ic junio synonyms o E.andinus, he
la ge o he wo species he ecognized; he
smalle species, E.mon osus, included chi -
alensis as a alid subspecies.
Koopman (1978) disag eed, claiming ha
he long-hai ed o ms o Neo opical Ep es-
icus a e es ic ed o cool highland a eas,
whe eas he sho -hai ed o ms occu in he
wa m lowlands. By his in e p e a ion, he
long-hai ed axa a e local a ian s o sho -
hai ed species ha ha e adap ed o coole
condi ions a highe ele a ions. Acco dingly,
Koopman (1978, 1993, 1994) ea ed he
membe s o Da is’ (1966) andinus g oup as
subspecies o nomencla u ally olde sho -
hai ed species based on size, assigning an-
dinus o E.b asiliensis, and mon osus and
chi alensis o E. u inalis. Howe e , Koop-
man’s adap i e scena io conce ning hai
leng h is e u ed by he occu ence o long-
hai ed Ep esicus—clea ly e e able o Da is’
andinus g oup—in lowland a eas o B azil,
Panama, Venezuela, and F ench Guiana.
13
In
addi ion, he sympa ic occu ence in no h-
eas e n Venezuela o E.b asiliensis,E. u-
inalis, and a hi d species ha Ochoa e al.
(1993) iden i ied as E.andinus de ini ely in-
dica es ha Ep esicus species wi h di e en
pelage ypes can coexis a he same ele a-
ion.
Specimen da a ci ed by Da is (1966), o-
ge he wi h o he collec ions subsequen ly e-
po ed in he li e a u e (e.g., by Handley,
1976), p o ide compelling e idence ha wo
species e e able o he andinus g oup a e
sympa ic a se e al Sou h Ame ican locali-
ies. Howe e , Da is did no pe sonally ex-
amine any o he ele an holo ypes in his
complex, so his decisions abou synonymies
13
We examined da k, long- u ed specimens o Ep-
esicus, clea ly e e able o Da is’ andinus g oup, om
lowland a eas o B azil (Amazonas, Rio Madei a, San o
An onio do Guaja a´ [ca. 25 m]: AMNH 92251, 93787),
F ench Guiana (Pa acou [ca. 30 m]: AMNH 267234,
267530, 267531, 268584; MNHN 1995.961, 1995.962),
and Panama (San Blas, A mila, Queb ada Venado [sea
le el]: USNM 335411). In addi ion, unambiguous de-
sc ip ions o andinus-g oup specimens ha e been e-
po ed in he li e a u e om a lowland si e in Venezuela
(Bolı´ a , Ima aca Fo es Rese e [180 m]; Ochoa e al.,
1993) and ano he in F ench Guiana (Pis e Sain -E
´lie
[ca. 45 m]; B osse and Cha les-Dominique, 1990).
1998 135SIMMONS AND VOSS: PARACOU BATS
Fig. 55. Do sal (A) and en al (B) iews o he skull o he holo ype o Ep esicus andinus (AMNH
33807; male) om Colombia. Do sal (C) and en al (D) iews o he skull o E.chi iquinus (AMNH
267234; male) om Pa acou. E.chi iquinus has a consis en ly la ge and mo e hea ily buil skull han
does E.andinus. Also no e he la ened, iangula bony pla e a he in e sec ion o he sagi al and
nuchal c es s in andinus, a s uc u e ne e p esen in chi iquinus. Scale ba s
5
10 mm.
136 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 56. La e al iews o he skull and lowe jaw o (A) he holo ype o Ep esicus andinus (AMNH
33807; male) and (B)E.chi iquinus om Pa acou (AMNH 267234; male). No e he species di e ence
in de elopmen o he sagi al c es . Scale ba s
5
10 mm.
a e p oblema ic. Because Koopman’s (1978)
e iew o he si ua ion was ob iously no an
imp o emen , a esh app aisal o he sys-
ema ics o Neo opical Ep esicus is neces-
sa y.
To de e mine he co ec iden i ica ion o
se e al blackish, long-hai ed specimens o
Ep esicus cap u ed a Pa acou, we examined
he holo ypes o E.andinus and E.chi alen-
sis as well as 136 o he specimens o Neo-
1998 143SIMMONS AND VOSS: PARACOU BATS
Fig. 57. Do sal (A) and en al (B) iews o he skull o Myo is nig icans (AMNH 267228; emale)
om Pa acou. Do sal (C) and en al (D) iews o he skull o M. ipa ius (AMNH 268591; emale)
om Pa acou. No e di e ences in s uc u e o he an e io b aincase, pala e, and audi o y egion. Scale
ba s
5
5 mm.

144 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 58. La e al iews o he skull and lowe jaw o (A)Myo is nig icans (AMNH 267228; emale)
and (B)M. ipa ius (AMNH 268591; emale). Scale ba s
5
5 mm.
ica based on pos o bi al b ead h and b ain-
case b ead h, bo h o which a e g ea e in M.
albescens whe e hese axa occu in sympa-
y o nea -sympa y (Husson, 1962, 1978;
LaVal, 1973; Baud and Menu, 1993). Husson
(1962, 1978) no ed ha Su inamese speci-
mens o M.albescens ha e a b aincase
b ead h
.
6.8 mm and a pos o bi al b ead h
.
3.6 mm, whe eas hese dimensions a e
smalle in M.nig icans om Su inam. None
o ou specimens om Pa acou has a b ain-
case b ead h
.
6.75 mm, and only one spec-
imen has a pos o bi al b ead h
.
3.50 mm
( able 57). In ou sample o 29 indi iduals,
1998 145SIMMONS AND VOSS: PARACOU BATS
he specimen wi h he la ges pos o bi al
b ead h (MNHN 1995.943, pos o bi al
b ead h 3.65 mm) has a b aincase b ead h
iden ical o he sample mean (6.41 mm).
Based on hese da a and compa isons o ex-
e nal mo phology, we a e con iden ha
none o he Myo is collec ed a Pa acou ep-
esen s M.albescens.Myo is albescens has
ye o be epo ed in F ench Guiana despi e
i s occu ence in Su inam and no heas e n
B azil. Howe e , we no e ha se e al speci-
mens iden i ied as M.nig icans by B osse
and Cha les-Dominique we e epo ed o
ha e pos o bi al b ead h
$
3.7 mm, and i is
possible ha one o mo e o hese specimens
may ep esen M.albescens. Al hough M.al-
bescens gene ally has a dis inc i ely ‘‘ os -
ed’’ pelage, LaVal (1973) no ed ha some
specimens lack his ea u e and ex e nally e-
semble M.nig icans.
Ou ouche ma e ial o Myo is nig icans
alls wi hin he lowe ange o size a ia ion
p e iously epo ed o his species (e.g., by
LaVal, 1973; Wilson and LaVal, 1974). Wi h
he excep ion o pos o bi al b ead h (no ed
abo e), he Pa acou specimens a e simila in
all dimensions o specimens epo ed om
o he locali ies in F ench Guiana and Su i-
nam (Husson, 1962, 1978; B osse and
Cha les-Dominique, 1990). Mos o ou
specimens ha e long, silky, b ownish do sal
u . Howe e , se e al ha e u ha is b own-
ish bu woollie in ex u e, and one indi id-
ual (a male) has almos black u . The en al
u is always sligh ly ligh e in colo han he
do sal u , appea ing os ed in many speci-
mens.
F
IELD
O
BSERVATIONS
: We made 29 ouch-
e ed cap u es o Myo is nig icans a Pa acou,
25 o which we e in g ound-le el mis ne s
and 4 o which we e in ele a ed ne s. O he
25 g ound-le el cap u es, 3 we e in c eekside
p ima y o es , 9 we e in manmade clea ings,
and 13 we e o e oadside puddles. The el-
e a ed ne cap u es we e made be ween 4
and 13 m abo e a na ow di oad.
See he ollowing species accoun o hab-
i a compa isons wi h Myo is ipa ius and o
in o ma ion abou cap u es o uniden i ied
Myo is.
Myo is ipa ius Handley
Figu es 57, 58
V
OUCHER
M
ATERIAL
: 6 emales (AMNH
*267224, *267524, *268591, *268592;
MNHN *1995.946, *1995.947) and 6 males
(AMNH *266366, *266376, *267523,
*268589; MNHN *1995.948, *1995.949);
see able 57 o measu emen s.
146 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
I
DENTIFICATION
: As no ed abo e, iden i i-
ca ion o species o Neo opical Myo is e-
qui es e e ence o LaVal (1973). Desc ip-
ions and measu emen s o Myo is ipa ius
can also be ound in Handley (1960) and
B osse and Cha les-Dominique (1990). No
subspecies o M. ipa ius a e cu en ly ec-
ognized (Koopman, 1994).
Ou specimens o Myo is ipa ius con o m
o he desc ip ion p o ided by LaVal (1973).
A sagi al c es is p esen in all indi iduals,
al hough i is poo ly de eloped in wo spec-
imens; P3 is less han one- ou h he heigh
o P4 in all indi iduals wi h bo h ee h (P3
is missing in one indi idual); and P3 is shi -
ed o he inside o he oo h ow in 64% o
ou specimens. Mos o ou specimens o M.
ipa ius ha e woolly b own do sal u and
sligh ly ligh e en al u . Howe e , some
indi iduals ha e u ha is silkie in ex u e,
and one specimen (clea ly adul based on
epiphyseal usion) is g ay-b own wi h sligh -
ly da ke unde pa s. Measu emen s o ou
ma e ial esemble hose epo ed by B osse
and Cha les-Dominique (1990) o F ench
Guianan M. ipa ius, al hough ou la ge se-
ies exhibi s somewha mo e size a ia ion
( able 57).
F
IELD
O
BSERVATIONS
: We made 12 ouch-
e ed cap u es o Myo is ipa ius a Pa acou:
11 indi iduals we e aken in g ound-le el
mis ne s and 1 was sho as i lew back and
o h in a egula bea abou 5 m abo e a
na ow di oad a nigh . O he 11 g ound-
le el mis ne cap u es, 5 we e in well-d ained
p ima y o es , 4 we e in swampy p ima y
o es , 1 was in c eekside p ima y o es , and
1 was in a manmade clea ing.
The obse ed habi a di e ence in g ound-
le el mis ne cap u e equencies be ween
Myo is nig icans and M. ipa ius is no e-
wo hy despi e he a he small numbe s o
ouche ed cap u es a ailable o compa ison
( able 58). Appa en ly, M.nig icans a o s
clea ings whe eas M. ipa ius is mo e com-
monly ound benea h he p ima y o es can-
opy. To ou knowledge, ecological di e -
ences be ween sympa ic popula ions o
hese species ha e no p e iously been e-
po ed in he li e a u e, possibly due o he
di icul y o ield iden i ica ion.
In addi ion o he ouche ed cap u es o
Myo is nig icans and M. ipa ius epo ed
abo e, we eco ded i e un ouche ed cap-
u es o Myo is in g ound-le el mis ne s o e
oadside puddles. These ba s we e iden i ied
in he ield as ep esen ing ei he M.nig i-
cans o M. ipa ius, bu we e eleased wi h-
ou adequa e con i ma ion o iden i ica ion.
MOLOSSIDAE
We cap u ed nine molossid species a Pa -
acou, including membe s o he gene a Eu-
mops,Molossops,Molossus, and P omops.
Fou o hese species ha e no been epo ed
p e iously om F ench Guiana, and ano he ,
o iginally desc ibed om he depa men , has
long been los in synonymy. Ano he eigh
molossids known om elsewhe e in F ench
Guiana o Su inam migh also occu in ou
s udy a ea (appendix 1).
Eumops au ipendulus (Shaw)
V
OUCHER
M
ATERIAL
: 2 emales (AMNH
*267537, *268594) and 1 male (MNHN
*1995.950); see able 59 o measu emen s.
I
DENTIFICATION
: Ege (1977) p o ided spe-
cies diagnoses and a key in he e ision o
he genus Eumops, which has no been su-
pe ceded by any compa ably comp ehensi e
s udy. Husson (1962, 1978) ga e a de ailed
desc ip ion and compa a i e measu emen s
1998 147SIMMONS AND VOSS: PARACOU BATS
o E.au ipendulus om Su inam, and addi-
ional in o ma ion can be ound in Sanbo n
(1932) unde he accoun o E.ab asus
(now conside ed a junio synonym o au i-
pendulus; see Goodwin, 1960; Husson, 1962,
1978; Ege , 1974, 1977). Two subspecies o
E.au ipendulus a e cu en ly ecognized
wi h he ollowing con inen al dis ibu ions
(Ege , 1974, 1977; Koopman, 1994): E.a.
au ipendulus (sou he n Mexico o Amazo-
nia, including he Guianas) and E.a.majo
(no he n A gen ina, sou he n Pa aguay, and
eas e n B azil). These axa a e dis inguished
on he basis o size, wi h majo being he
la ge o he wo.
Ou ouche ma e ial con o ms in all e-
spec s o Ege ’s (1977) desc ip ion o Eu-
mops au ipendulus au ipendulus. The Pa a-
cou specimens a e also compa able in size o
hose epo ed om Su inam by Husson
(1962, 1978), al hough one o ou emales
has a sho e maxilla y oo h ow and na ow-
e skull han ei he o he emales whose
measu emen s he published (op. ci .: able
32).
F
IELD
O
BSERVATIONS
: Ou h ee specimens
o Eumops au ipendulus om Pa acou we e
cap u ed in mis ne s suspended 17--23 m
abo e a na ow di oad h ough he o es .
Eumops hansae Sanbo n
V
OUCHER
M
ATERIAL
: 1 emale (AMNH
*267538); see able 59 o measu emen s.
I
DENTIFICATION
: Diagnos ic cha ac e s and
measu emen s o Eumops hansae ha e been
discussed by Ga dne e al. (1970), Ege
(1977), and B osse and Cha les-Dominique
(1990). No subspecies a e cu en ly ecog-
nized (Ege , 1977; Koopman, 1994).
Ou Pa acou specimen con o ms o pub-
lished desc ip ions o Eumops hansae and
alls wi hin he ange o size a ia ion p e-
iously epo ed by Ga dne e al. (1970),
Ege (1977), and B osse and Cha les-Dom-
inique (1990).
F
IELD
O
BSERVATIONS
: Ou single specimen
o Eumops hansae was caugh in a mis ne
suspended 10–13 m o e a na ow di oad.
Molossops
Bo h o he species o Molossops ha we
cap u ed a Pa acou belong o he subgenus
Cynomops, he con en s o which ha e ne e
been subjec ed o mode n e isiona y ea -
men . Table 60 summa izes he diagnos ic
ai s o he ou species ha we ecognize
based on he li e a u e and ou examina ion
o ep esen a i e museum specimens. All o
148 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY

1998 149SIMMONS AND VOSS: PARACOU BATS
he subs an i e axonomic di icul ies we en-
coun e ed conce n he h ee smalle species,
an ex ended commen a y on which is p o-
ided below in he accoun o M.pa anus.
Molossops (Cynomops)ab asus
(Temminck)
V
OUCHER
M
ATERIAL
: 1 emale (AMNH
*267534); see able 62 o measu emen s.
I
DENTIFICATION
:Molossops ab asus is eas-
ily iden i ied by size and o he mo phological
ai s om o he membe s o he subgenus
Cynomops ( ables 60, 61). Fou subspecies
a e cu en ly ecognized (Williams and Gen-
oways, 1980a; Koopman, 1994): M.a.mas-
i us (Venezuela and he Guianas), M.a.b a-
chymeles (eas e n Pe u), M.a.ab asus (eas -
e n B azil), and M.a.ce as es (Pa aguay and
no he n A gen ina). These axa a e dis in-
guished p incipally on he basis o size, wi h
mas i us being he la ges (Goodwin, 1958;
Taddei e al., 1976; Williams and Genoways,
1980a, 1980b).
Ou Pa acou ouche , appa en ly he i s
eco d om F ench Guiana, con o ms o p e-
ious quali a i e desc ip ions o Molossops
ab asus and alls wi hin he ange o size
a ia ion p e iously epo ed o he species
(Goodwin, 1958; Taddei e al., 1976; Wil-
liams and Genoways, 1980a, 1980b; Koop-
man, 1994). Ou specimen, a emale, is
somewha smalle han a male iden i ied as
M.a.mas i us ha Williams and Genoways
(1980a) epo ed om Su inam, bu his di -
e ence is o he same magni ude as he sex-
ual dimo phism ha Taddei e al. (1976) doc-
umen ed in a collec ion o M.ab asus om
eas e n B azil.
F
IELD
O
BSERVATIONS
: We caugh ou only
specimen o Molossops ab asus in a mis ne
suspended 18–21 m abo e a na ow di
oad.
Molossops (Cynomops)pa anus (Thomas)
Figu es 59, 60
V
OUCHER
M
ATERIAL
: 1 male (AMNH
*267535); see able 62 o measu emen s.
I
DENTIFICATION
:Molossops pa anus, o ig-
inally desc ibed as Molossus plani os is
pa anus by Thomas (1901b), con inues o be
ecognized as a subspecies o Molossops
plani os is by au ho s (e.g, Koopman, 1978,
1993, 1994). Al hough Handley (1976) e-
po ed specimens iden i ied as Molossops
pa anus,M.plani os is, and M.g eenhalli
om he Venezuelan s a e o Bolı´ a , he did
no commen on he c i e ia he used o dis-
inguish hese axa. Pa ial diagnoses and de-
sc ip ions o he species and subspecies o
Cynomops occu h oughou he li e a u e,
bu signi ican inconsis encies exis among
published accoun s. Con lic ing desc ip ions
150 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
o en al colo a ion, p esence o absence o
whi e hai bases, and deg ee o de elopmen
o hai pa ches on he wings make unambig-
uous iden i ica ion di icul . To add ess his
p oblem, we consul ed o iginal axonomic
desc ip ions, li e a u e accoun s o addi ional
ma e ial, and examined ep esen a i e speci-
mens o all o he smalle species o de elop
wo king diagnoses o iden i ica ion.
16
16
The specimens we examined in addi ion o hose
om Pa acou a e: Molossops g eenhalli (Mexico:
USNM 511543, 523453; Panama: USNM 310264–
310268, 310270–310275, 368108, 396481, 449875;
Venezuela: USNM 387745, 517509; T inidad: AMNH
175326 [holo ype], 176285, 207071); Molossops pa -
anus (Panama: AMNH 183868, USNM 317627; Colom-
bia: ROM 41479; Venezuela: USNM 387744; Guyana:
ROM 32426, 57337–57338, 57375; B azil: AMNH
79744–79745); Molossops plani os is (Panama:
AMNH 183161, 183863; Venezuela: AMNH 17096–
17097; B azil: AMNH 37043–37049, 37050–37052,
79725, 79727, 79731, 79733, 93879–93886, 92971,
92753–92755, 94630–94653, 236221; Pa aguay:
234455–234459)
1998 151SIMMONS AND VOSS: PARACOU BATS
Fig. 59. Do sal (A) and en al (B) iews o he skull o Molossops pa anus (AMNH 267535; male)
om Pa acou. Do sal (C) and en al (D) iews o he skull o he holo ype o M.g eenhalli (AMNH
175326; male) om T inidad. No e species di e ences in size and c anial p opo ions. Scale ba s
5
10
mm.
152 NO. 237BULLETIN AMERICAN MUSEUM OF NATURAL HISTORY
Fig. 60. La e al iews o he skull and lowe jaw o (A)Molossops pa anus (AMNH 267535; male)
and (B) he holo ype M.g eenhalli (AMNH 175326; male). Scale ba s
5
10 mm.