Biology of Bats of the New World Family Phyllostomatidae. Part I

SPECIAL PUBLICATIONS
THE
MUSEUM
TEXAS TECH UNIVERSITY
Biology
o
Ba s
o
he
New
Wo ld Family
Phyllos oma idae. Pa I
Edi ed
by
Robe
J.
Bake ,
J.
Knox
Jones, J .,
and
Dil o d
C.
Ca e
June
1976
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No.
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TEXAS
TECH
UNIVERSITY
G o e
E.
Mu ay, P esiden
Glenn E. Ba ne , Execu i e Vice P esiden
Regen s.-Clin
Fo mby (Chai man), J. F ed Bucy, J ., Bill E. Collins,
John
J. Hinchey,
A.
J. Kemp, J ., Robe
L.
P luge , Cha les G. Sc uggs, Judson F. Williams, and Don
R.
Wo kman.
Academic
Publica ions Policy
Commi ee.-J.
Knox Jones, J . (Chai man), Dil o d C.
Ca e (Execu i e Di ec o ), C. Leona d Ainswo h, Samuel E. Cu l, Ha old
E.
D egne,
Hugh
H.
Genoways, Ray
C.
Janeway, William
R.
Johnson,
S.
M.
Kennedy, Thomas
A.
Lang-
o d, Geo ge F. Meenaghan, Ha ley
D.
Obe helman. Robe L. Packa d, and Cha les W.
Sa gen .
The Museum
Special Publica ions No.
10
218 pp.
25
June
1976
$6.00
Special Publica ions
o
The Museum a e numbe ed sepa a ely and published
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basis unde he auspices
o
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o
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o
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lica ions, and in coope a ion wi h he In e na ional Cen e o A id and Semi-A id Land
S udies. Copies may be ob ained on
an
exchange basis om,
o
pu chased h ough, he Ex-
change Lib a ian, Texas Tech Uni e si y, Lubbock, Texas 79409.
Texas Tech P ess, Lubbock, Texas
1976
SPECIAL PUBLICATIONS
THE
MUSEUM
TEXAS
TECH
UNIVERSITY
Biology
o
Ba s
o
he
New
Wo ld Family
Phyllos oma idae. Pa I
Edi ed
by
Robe
J.
Bake ,
J.
Knox
Jones, J .,
and
Dillo d
C.
Ca e
No.
10
June
1976
TEXAS
TECH
UNIVERSITY
G o e
E.
Mu ay,
P esiden
Glenn
E. Ba ne , Execu i e Vice P esiden
Regen s.-Clin
Fo mby
(Chai man),
J.
F ed
Bucy,
J .,
Bill E. Collins,
John
J. Hinchey,
A.
J. Kemp, J .,
Robe
L. P luge ,
Cha les
G. Sc uggs,
Judson
F. Williams,
and
Don
R.
Wo kman.
Academic
Publica ions Policy
Commi ee.-J.
Knox
Jones, J .
(Chai man),
Dil o d C.
Ca e
(Execu i e Di ec o ), C.
Leona d
Ainswo h,
Samuel
E. Cu l,
Ha old
E.
D egne,
Hugh
H. Genoways, Ray
C.
Janeway,
William
R.
Johnson,
S.
M.
Kennedy,
Thomas
A. Lang-
o d,
Geo ge
F.
Meenaghan,
Ha ley
D.
Obe helman,
Robe
L.
Packa d,
and
Cha les
W.
Sa gen .
The
Museum
Special Publica ions No. 10
218 pp.
25
June
1976
$6.00
Special Publica ions
o
The
Museum
a e
numbe ed
sepa a ely
and
published
on
an
i egula
basis
unde
he
auspices
o
he
Dean
o
he
G adua e
School
and
Di ec o
o
Academic
Pub-
lica ions,
and
in
coope a ion
wi h
he
In e na ional
Cen e
o
A id
and
Semi-A id
Land
S udies. Copies
may
be
ob ained
on
an
exchange basis om,
o
pu chased
h ough,
he
Ex-
change
Lib a ian,
Texas
Tech
Uni e si y, Lubbock, Texas 79409.
Texas
Tech
P ess, Lubbock, Texas
1976
CONTENTS
INTRODUCTION
5
ANNOTATED
CHECKLIST,
WITH
KEYS
TO
SUBFAMILIES
AND
GENERA.
. . . . . . 7
J. Knox Jones, J ., and Dil o d
C.
Ca e ,
The Museum, Texas Tech Uni e si y, Lubbock, 79409.
ZOOGEOGRAPHY
39
Ka l
F.
Koopman, Depa men
o
Mammalogy,
The
Ame ican Museum
o
Na u al His o y,
Cen al Pa k Wes
a
79 h
S ., New Yo k, 10024.
CHIROPTERAN
EVOLUTION
49
James Dale Smi h, Depa men
o
Biological Sciences,
Cali o nia S a e Uni e si y, Fulle on, 92634.
COLLECTING
TECHNIQUES
71
Me lin
D.
Tu le, Ve eb a e Di ision,
Milwaukee Public Museum, Milwaukee, Wisconsin 53233.
CARE
IN
CAPTIVITY.
........................................
..
89
A hu M. G eenhall, U.S. Fish and Wildli e Se ice,
Na ional
Fish
and Wildli e Labo a o y, Na ional Museum
o
Na u al
His o y, Washing on, D.C. 20560.
ECONOMICS
AND
CONSERVATION.
. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
..
133
Clyde Jones, U.S. Fish and Wildli e Se ice, Na ional Fish and Wildli e
Labo a o y, Na ional Museum
o
Na u al His o y,
Washing on, D.C. 20560.
BRAIN
ANATOMY
147
V.
Rick McDaniel, Di ision
o
Biological Sciences,
A kansas S a e Uni e si y, Jonesbo o, 72467.
LACTATION
AND
MILK
201
Robe Jenness and Eugene
H.
S udie , Depa men
o
Biochemis y,
Uni e si y
o
Minneso a, S . Paul, 55108, and Depa men
o
Biology,
Uni e si y
o
Michigan, Flin , 48503.

INTRODUCTION
Because
o
hei adap i e di e si y and,
in
many ins ances, unique mo phologi-
cal a ibu es, ba s
o
he amily Phyllos oma idae ha e long ascina ed biologis s.
Known only om he New Wo ld, mos gene a
o
phyllos oma ids a e s ic ly
limi ed o opical en i ons, bu some ep esen a i es occu as a no h as he
sou hwes e n Uni ed S a es and o he s sou hwa d o he no he n pa s
o
A -
gen ina and Chile; some species also a e dis ibu ed in he Bahamas and
on
he
islands
o
he G ea e and Lesse An illes. Wi h he ad en in ela i ely ecen
yea s
o
imp o ed me hods
o
collec ing ba s (see Tu le, his olume), a e-
mendous weal h
o
in o ma ion on phyllos oma ids has been ga he ed and i
is
he
pu pose
o
his publica ion, which ul ima ely will con ain mo e han 20 indi idual
chap e s, o b ing hese da a oge he
in
o de o assess wha now
is
known abou
he amily and o p o ide adepa u e poin o u he s udies.
Owing o he la ge numbe
o
con ibu ions, all
o
which we e solici ed by us
om pe sons we el o be knowledgeable
o
he subjec ma e , and he ac ha
se e al con ibu ions a e necessa ily leng hy, he decision was made o g oup
chap e s in o h ee pa s. Each pa will be asepa a ely numbe ed Special Pub-
lica ion
o
The Museum a Texas Tech Uni e si y.
In
o de o es ablish awo k-
able app oach by which e e ence could be made consis en ly o axa h oughou
he se ies, he anno a ed checklis
by
Jones and
Ca e
was ci cula ed o all
au ho s. Each was asked o ollow he nomencla u e and sys ema ic a angemen
in he checklis o , al e na i ely, o documen depa u es he e om. This sys em,
i
is hoped, will allow eade s o ela e in o ma ion om one chap e o he nex
wi hou he handicap
o
con lic ing names o he same o ganism.
Manusc ip s o mos con ibu ions i s we e solici ed in 1973. Mos manu-
sc ip s had been ecei ed by he end
o
1974. As edi o ial wo k p og essed, some
au ho s p o ided up-da ed in o ma ion and all au ho s
o
chap e s
in
Pa 1had
he oppo uni y o inse limi ed ma e ials a he ime hey ecei ed galley p oo s
(in mos cases Oc obe 1975). The e o e, con en is as cu en as easonably
could be an icipa ed o ap ojec
o
his kind. O ganiza ion and edi o ial s yle
ollows ha es ablished o he Special Publica ions
o
The Museum
a
Texas
Tech Uni e si y. O he wise, au ho s we e allowed b oad la i ude conce ning ma-
e ial o be included in hei chap e s. Acco dingly, and o ob ious o he easons,
some chap e s will o e lap o he s
in
con en .
E en hough some edundacy has eSUl ed,
we
hough i bes o ha e asec ion
on he ci ed li e a u e wi h each con ibu ion. Ci a ions o manusc ip s in his
collec ed se ies a e ca ied in ex as " his olume," which does no necessa ily
indica e ha he chap e appea s in he same pa
o
he se ies as he one in
which i is ci ed.
No embe 1975
5
Robe J. Bake
J. Knox Jones, J .
Dil o d
C.
Ca e
ANNOTATED
CHECKLIST,
WITH
KEYS
TO
SUBFAMILIES
AND
GENERA
J.
KNOX JONES,
JR.,
AND
DILFORD
C.
CARTER
Lea -nosed ba s
o
he New Wo ld amily Phyllos oma idae a e p ima ily
limi ed in dis ibu ion o opical and sub opical egions. A
ew
species each
sub empe a e a eas. The anlily has aknown ossil eco d da ing back o Miocene
imes. Mos phyllos oma ids a e ui ea e s
o
nec a eede s, bu some, p ima ily
species in he sub amily Phyllos onla inae, a e ca ni o ous o insec i o ous, and
he unique desmodon ines a e sangui o ous.
The amily
is
unusually di e se om an e olu iona y poin
o
iew, comp ising
six sub amilies, 49 cu en ly ecognized Recen gene a, and 137 Recen nominal
species. Twen y- ou gene a a e mono ypic. The sub amilies con ain he ollowing
numbe s
o
gene a and species as he e ecognized: Phyllos oma inae,
11
and 32;
Glossophaginae,
13
and 32; Ca olliinae, wo and se en; S enode minae, 17 and
54; Phyllonyc e inae, h ee and se en; and Desmodon inae, h ee and h ee.
Sys ema ic inqui y in he pas decade has ended o educe he numbe
o
ecognized gene a and species, bu he disco e y
o
new axa con inues. Some
species a e a e
in
museum collec ions and hei ela ionships poo ly unde s ood.
Va ious
new
echniques applied in ecen yea s o he s udy
o
phyllogene ic ela-
ionships ha e esul ed
in
ecogni ion
o
new axonomic alignmen s-- o ex-
ample, inclusion
o
he ampi e ba s as asub amily
o
he Phyllos oma idae
(Fo man e aI., 1968) and exclusion
o
he Mo moopidae (Smi h, 1972), o me ly
ega ded
as
asub amily
o
his g oup.
As s anda d e e ences o apoin
o
depa u e in compila ion
o
his anno a ed
lis , we used Hall and Kelson (1959) o No h Ame ica and Cab e a (1958) o
Sou h Anle ica. A a ie y
o
publica ions has appea ed subsequen o hese wo
basic documen s in which he dis ibu ion and sys ema ics
o
phyllos oma ids a e
ea ed.
O
hese, e isions and e iews a e ci ed a he app op ia e places
in
he
accoun s. Faunal epo s
o
special in e es a e no ed below. Recou se o he
li e a u e we ha e ci ed will lead he in e es ed esea che o mos
o
he pub-
lished sou ces used in compiling his synopsis.
Villa-R. (1967) summa ized ma e ial on Chi op e a
o
Mexico. Publica ions
since ha ime on Chihuahua (Ande son, 1972), Jalisco (Wa kins
e
aI., 1972),
Oaxaca (Goodwin, 1969), Sinaloa (Jones
e
al., 1972), he Yuca an Peninsula
(Jones e al., 1973), and Zaca ecas (Genoways and Jones, ]968; Ma son and
Pa en, 1975) ea majo aunal uni s as awhole.
Fo
Cen al Ame ica, he
pape s
o
Jones (1966) on Gua emala, Bu and S i on (1961) on
EI
Sal ado ,
Jones e al. (1971
b)
and Bake and Jones (1975) on Nica agua, S a e and Case-
bee (1968) and Ga dne e al. (1970) on Cos a Rica, and Handley (1966) on
Panama a e use ul, as well as hose by Da is e al. (1964) and Ca e e al.
(1966)
7
8
SPECIAL
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on he egion as awhole. Choa e and Bi ney (1968), Koopman (1968) and Jones
and Phillips (1970) a e use ul ecen e e ences o ba s in he An illean egion.
Rela i ely
ew
majo con ibu ions ha e been published
on
Sou h Ame ica
since Cab e a's (op. ci .) compendiunl, hose
on
T inidad and Tobago (Goodwin
and G eenhall, 1961), Su inam (Husson, 1962), Pe u (Tu le, 1970), Colombia
(Aellen, 1970; Ma inkelle and Cadena, 1972), and U uguay (Ximenez e ai.,
1972) being especially no ewo hy. S udies
o
amo e limi ed scope, such as Hill's
(1964) epo
o
asmall collec ion om Guyana, B osse 's (1965) and Bake 's
(1974) pape s on Ecuado , and he publica ion by Villa-R. and Co nejo (1969) on
no he n A gen ina also ha e p o ed use ul (see also he appendix
o
he con i-
bu ion
on
zoogeog aphy by
K.
F. Koopman in his olume).
I
is
o
in e es ha
ew epo s on he B azilian auna ha e appea ed since Cab e a's wo k, pape s by
Handley (1967)
on
he Belem a ea and by Pine e
al.
(1970) on acollec ion onl
Ma o G osso, and Pe acchi and de Albuque que (1971) on he s a es
o
Rio
de
Janei o and
Guanaba a
being no able excep ions. [See also Ga dne 's (1976)
ecen pape
on
Pe u. ]
In a ecen
pape
on he mammalian auna
o
he An illes, Va ona (1974) in-
co po a ed anumbe
o
sys ema ic changes wi h espec o ba s ound in ha
egion.
Fo
example, he ega ded all species
o
A dops, A i eus, and Phyllops as
assignable o he subgenus A i eus
o
he genus S enode ma,
and
placed Mono-
phyllus
as
asubgenus
o
Giossophaga. Because Va ona p esen ed no e idence
suppo i e
o
hese and o he changes, we ha e no ollowed his a angemen
he e.
We a e indeb ed o anumbe
o
colleagues, p incipally Robe J. Bake , Al ed
L.
Ga dne , Hugh H. Genoways, Clyde Jones, Ka l F. Koopman, and Don E.
Wilson, o sc u inizing
an
ea ly d a
o
his manusc ip .
SUBFAMI
LY
PHYLLOSTOMATINAE
Genus
MICRONYCTERIS
G ay
Mic onyc e is megalo is (G ay, 1842)
Dis ibu ion.-Wes e n (Jalisco) and eas e n (Tamaulipas) Mexico sou h-
eas wa d h ough Middle Ame ica and much
o
no he n and cen al Sou h
Ame ica o Amazonian Pe u and Sao Paulo, B azil; also eco ded om G enada
in he Lesse An illes.
Sys ema ics.-Fou
subspecies cu en ly a e ecognized: megaio is (mos
o
Sou h Ame ican segmen
o
species dis ibu ion); homezi (no hwes e n Vene-
zuela); mexicana (Mexico sou h o wes e n Nica agua and adjacen Cos a Rica);
mic o is (eas e n Nica agua sou heas wa d o Panama
and
adjacen pa s
o
no hwes e n Sou h Ame ica).
Mic onyc e is schmid o um Sanbo n, 1935
Dis ibu ion.-
Yuca an Peninsula
o
Mexico sou heas wa d o no hwes e n
Sou h Ame ica.
Sys ema ics.-M.
schmid o um
is
amono ypic species.
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE 9
Mic onyc e is minu a (Ge ais, 1855)
Dis ibu ion.-Nica agua sou heas wa d o Sou h An}e ica (including T ini-
dad) a leas o B azil and eas e n Pe u.
Sys ema ics.-M.
minu a
is
cu en ly ega ded as amono ypic species.
Toge he wi h megalo is and schmid o um his species ep esen s he subgenus
Mic onyc e is.
Mic onyc e is hi su a (Pe e s, 1869)
Dis ibu ion.-Hondu as sou heas wa d o no he n Sou h Ame ica (Colon1-
bia, Venezuela, Guyana, T inidad, and Pe u).
Sys ema ics.-M.
hi su a
is
amono ypic species and he sole ep esen a i e
o
he subgenus Xenoc enes.
Mic onyc e is b achyo is (Dobson, 1878)
Dis ibu ion.-Oaxaca sou heas wa d h ough Cen al Ame ica o Amazonian
B azil.
Sys ema ics.-M.
b achyo is
is
amono ypic species and ep esen s he sub-
genus Lamp onyc e is.
The
speci ic name pla yceps, widely used o his ba o
se e al decades, is asynonym
o
b achyo
is.
Mic onyc e is pusilla Sanbo n, 1949
Dis ibu ion.-No he n
B azil, eas e n Colombia, p obably adjacen egions
o
Sou h Ame ica.
Sys ema ics.-M.
pusilla is amono ypic species and ep esen s he subgenus
Neonyc e is.
Mic onyc e is nice o i Sanbo n, 1949
Dis ibu ion.-Nica agua o no he n Sou h Ame ica (including T inidad and
sou h a leas o no he n B azil and no he n Amazonian Pe u).
Sys ema ics.-M.
nice/o i
is
amono ypic species and he only ep esen a i e
o
he subgenus T inyc e is.
Mic onyc e is syl es is (Thomas, 1869)
Dis ibu ion.-Wes e n
(Naya i ) and eas e n (Ve ac uz) Mexico sou heas -
wa d h ough Cen al Ame ica o Panama and in o no he n Sou h Ame ica a
leas
as
a eas
as
T inidad, no heas e n B azil, and eas e n Pe u.
Sys ema ics.-M.
syl es is is hough o be amono ypic species.
Mic onyc e is behni (Pe e s, 1865)
Dis ibu ion.-Known
only om cen al B azil and Pe u.
Sys ema ics.-
This nominal species is poo ly known. Along wi h M. syl es is,
wi h which i e iden ly
is
closely ela ed, behni cons i u es he subgenus Gly-
phonyc e is.
Lonchophylla mo dax Thomas, 1903
Dis ibu ion.-Repo ed
om Ecuado , Boli ia, and B azil.
Sys ema ics.-L.
mo dax
is
conside ed he e o be amono ypic species bu may
include conca a
as
ano he n subspecies.
Lonchophylla obus a Mille , 1
91
2
Dis ibu ion.-Repo ed
onl Nica agua, Cos a Rica, Panama, Calombia,
Venezuela, and Pe u.
Sys ema ics.-L.
obus a
is
amono ypic species.
Lonchophylla homasi
J.
A. Allen, 1904
Dis ibu ion.-Known
om Panama, Venezuela, Guyana, Su inam, B azil,
Pe u, and Boli ia.
Sys ema ics.-L.
homasi
is
amono ypic species.
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TEXAS
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Genus
ANouRA
G ay
Anou a geo oyi G ay, 1838
Dis ibu ion.-Wes e n
(Sinaloa) and eas e n (San Luis Po osi) Mexico sou h
o sou heas e n B azil and no hwes e n A gen ina.
Sys ema ics.-
Th ee nominal subspecies a e: geo oyi (Venezuela, Su inam,
T inidad, B azil, A gen ina, and Boli ia); lasiopyga (Mexico sou h o no he n
Colombia); and pe uana (Colombian Andes sou h o Pe u).
Genus
LIONYCTERIS
Thomas
Lionyc e is spu elli Thomas, 1
91
3
Dis ibu ion.-Eas e n
Panama and no he n Sou h Ame ica ( eco ded om
Guyana, no he n B azil, Venezuela, Colombia, and Amazonian Pe u).
Sys ema ics.-L.
spu elli
is
amono ypic species.
Lonchophylla conca a Goldman, 1914
Dis ibu ion.-Repo ed
om Cos a Rica, Panama, Colombia, and Pe u.
Sys ema ics.-L.
conca a
is
ecognized p o isionally
as
amono ypic species
dis inc om mo dax, wi h which
i
may be conspeci ic. Handley (1966) con-
side ed conca a o be ano he n subspecies
o
mo dax, bu ecen au ho s ha e
no ollowed ha a angemen .
Anou a caudi e
(E.
Geo oy S .-Hilai e, 1818)
Dis ibu ion.-No he n
Sou h Ame ica sou h o Pe u and B azil.
Sys ema ics.-
Two subspecies cu en ly a e ecognized: caudi e (Colombia
eas h ough Venezuela and he Guianas and sou h in eas e n B azil o Sao
Paulo); aequa o is (Ecuado and Pe u).
16

BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
Anou a
cul a a Handley, 1960
Dis ibu ion.-Known
om
Cos a
Rica,
Panama,
and
Venezuela.
Sys ema ics.-A.
cul a a is amono ypic species.
17
Anou a
we ckleae S a e , 1
969
Dis ibu ion.-Known
only om
Cos a
Rica.
Sys ema ics.-A.
we ckleae
is
amono ypic species closely ela ed o A.
cul a a.
Ano;:; a b e i os um Ca e , 1
968
Dis ibu ion.-Reco ded om
Colombia
(San ande ) and eas e n Pe u.
Sys ema ics.-As
p esen ly known, A. b e i os um
is
amono ypic species.
Genus
SCLERONYCTERIS
Thomas
Scle onyc e is ega Thomas, 1895
Dis ibu ion.-
This a e species is known only om B azil and Venezuela.
Sys ema ics.-S.
ega
is amono ypic species.
Genus
LICHONYCTERIS
Thomas
Lichonyc e is degene Mille , 1931
Dis ibu ion.-Known
only om lowe Amazon egion
o
B azil.
Sys ema ics.-L.
degene is amono ypic species.
Lichonyc e is obscu a Thomas, 1895
Dis ibu ion.-Gua emala sou heas wa d o Sou h
Ame ica
a leas as a as
Su inam
and eas -cen al Pe u.
Sys ema ics.-L.
obscu a is amono ypic species.
Genus
HYLONYCTERIS
Thomas
Hylonyc e is unde woodi Thomas, 1903
Dis ibu ion.-Wes e n
Mexico (no h o Jalisco) sou heas wa d o wes e n
Panama.
Sys ema ics.-
Two
subspecies (Phillips
and
Jones, 1971), unde woodi (Ve a-
c uz
and
no he n
Oaxaca
sou heas wa d o Panama) and mino (wes e n Mexi-
co),
a e
ecognized.
Genus
PLATALINA
Thomas
Pla alina geno ensium Thomas, 1928
Dis ibu ion.-
This
a e
ba is known only om Pe u, p incipally wes
o
he
Andes.
Sys ema ics.-P.
geno ensium
is
amono ypic species.
Choe oniscus pe iosDs Handley, 1966
Dis ibu ion.-Known
only om Paci ic Coas
o
Colombia.
Sys ema ics.-C.
pe iosus
is
adis inc i e, mono ypic species.
Choe oniscus mino (Pe e s, 1868)
Dis ibu ion.-Reco ded
om B azil, Colombia, Ecuado , Pe u,
and
Su inam.
Sys ema ics.-C.
mino
is
amono ypic species closely ela ed o
C.
inca and
C. in e medius.
Choe oniscus in e medius (J. A. Allen
and
Chapman,
1893)
Dis ibu ion.-
Though o be es ic ed o T inidad, bu epo ed also om
Pe u by Tu le (1970).
Sys ema ics.-C.
in e medius
is
amono ypic species closely ela ed o
C.
mino and
C.
inca.
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Genus
CHOERONYCTERIS
Tschudi
Choe onyc e is mexicana Tschudi, 1844
Dis ibu ion.-Ex eme
sou he n pa s
o
Cali o nia, A izona, and New
Mexico sou hwa d o Hondu as.
Sys ema ics.-C.
mexicana
is
he e ega ded as amono ypic species. How-
e e , Pi lo (1967) desc ibed asubspecies
(ponsL)
om no hwes e n Venezuela;
we a e uncon inced by Pi lo 's b ie desc ip ion ha his wo specinlens a e e-
e able o he genus Choe onyc e is.
Choe oniscus inca (Thomas, 1912)
Dis ibu ion.-Reco ded
om Guyana, Ecuado , Pe u, and Venezuela.
Sys ema ics.-C.
inca is amono ypic species closely ela ed o he wo p eced-
ing axa. Specimens
o
he genus Choe oniscus a e a e in museum collec ions.
Only abou wo dozen indi iduals
o
he mino -in e medius-inca complex
ha e been epo ed in he li e a u e, and he cha ac e is ics
o
he h ee species
ne e ha e been de ined in acompa a i e sense. Clea ly, his g oup is in need
o
sys ema ic e iew.
I
may well
b~
ha mino , in e medius, and inca ep esen
asingle species.
Genus
CHOERONISCUS
Thomas
Choe oniscus godmani (Thomas, 1903)
Dis ibu ion.-Wes e n
Mexico (Sinaloa) sou heas wa d o Colombia
and
Venezuela.
Sys ema ics.-C.
godmani
is
amono ypic species.
18
BIOLOGY OF
THE
PHYLLOSTOMATIDAE 19
Genus
MUSONYCTERIS
Schaldach and McLaughlin
Musonyc e is ha isoni Schaldach and McLaughlin, 1960
Dis ibu ion.-P esen ly known only om he s a es
o
Colima and Gue e o
in
wes e n Mexico.
Sys ema ics.-M.
ha isoni
is
amono ypic species. Al hough some ecen
au ho s ha e ega ded Musonyc e is as asynonym
o
Choe onyc e is, we ollow
Phillips (1971) in ega ding i as adis inc genus.
SUBFAMILY
CAROLLIINAE
Genus
CAROLLIA
G ay
Ca ollia cas anea H. Allen, 1890
Dis ibu ion.-Hondu as sou heas wa d h ough Colombia, Ecuado , and
Pe u o Boli ia.
Sys ema ics.-Acco ding o Pine (1972),
C.
cas anea
is
amono ypic
species.
Ca ollia sub u a (Hahn, 1905)
Dis ibu ion.-Wes e n Mexico (Jalisco) sou heas wa d, mos ly
in
he Paci ic
e san
o
Middle Ame ica, o Nica agua.
Sys ema ics.-C.
sub u a was ega ded by Pine (1972) as amono ypic species.
Ca ollia b e icauda (Schinz, 1821)
Dis ibu ion.-Eas e n Mexico (sou he n San Luis Po osi and adjacen Ve a-
c uz) sou heas wa d o no he n and wes e n Sou h Ame ica (no heas e n B azil,
Colombia, Venezuela, Ecuado , Amazonian Pe u, and Boli ia).
Sys ema ics.-As
in he case
o
he p e ious wo species
o
Ca
o
Ilia,
Pine
(1972) conside ed
C.
b e icauda o be mono ypic.
Ca ollia pe spicilla a (Linnaeus, 1758)
Dis ibu ion.-Ve ac uz and Oaxaca sou heas wa d o Sou h Ame ica, whe e
he species
is
widely dis ibu ed sou h o Boli ia, Pa aguay, and sou he n B azil;
also epo ed om T inidad, Tobago, and he An illean island
o
G enada ( e-
co ded occu ences on Jamaica and Redondo Island, in he no he n Lesse
An illes, a e ques ionable).
Sys ema ics.-
Two subspecies cu en ly a e en a i ely ecognized (Pine,
1972), pe spicilla a in much
o
he Sou h Ame ican ange
o
he species and
az eca
in
Middle Ame ica and adjacen no hwes e n Sou h Ame ica. Pine (op.
ci .) also no ed ha he name
C.
p. icolo migh apply o specimens om he
sou he n pa
o
he ange
o
he species.
Rhinophylla ale hina Handley, 1966
Dis ibu ion.-Known
only om wes e n Colombia.
Sys ema ics.-R.
ale hina
is
amono ypic species.
Rhinophylla ische ae Ca e , 1966
Dis ibu ion.-Known
only om Amazonian pa s
o
Pe u
and
B azil, and
adjacen Colombia and Ecuado .
Sys enla ics.-R.
ische ae
is
amono ypic species.
Genus
RHINOPHYLLA
Pe e s
Rhinophylla pumilio Pe e s, 1865
Dis ibu ion.-No he n
Sou h Ame ica in Guyana, Su inam, Venezuela,
Colombia, B azil, and eas e n
Ecuado
and Pe u.
Sys ema ics.-R.
pumilio
is
amono ypic species.
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20
SUBFAMILY
STENODERMINAE
Genus
STURNIRA
G ay
The sys ema ics
o
ba s
o
he genus S u ni a a e, o he mos pa , poo ly
unde s ood. Se e al new species ha e been named in he pas decade
o
so.
The
lis p esen ed he e
is
p o isional, pending publica ion
o
Luis
de
la To e's long
awai ed e ision
o
he genus.
S u ni a homasi
de
la
To e
and Schwa z, 1966
Dis ibu ion.-Known
only om Guadeloupe, Lesse An illes.
Sys ema ics.-S.
homasi is en a i ely ecognized he e as a alid, mono-
ypic species because i di e s in se e al ways om
o he
named An illean
popula ions
o
S u ni a.
I
is ela ed o lilium and ul ima ely may p o e bes
ega ded as asubspecies
o
ha species.
S u ni a Iilium
(E.
Geo oy S .-Hilai e, 1810)
Dis ibu ion.-Widely
dis ibu ed om wes e n (Sono a) and eas e n (Tamau-
lipas) Mexico sou hwa d h ough Middle Ame ica and h oughou mos
o
opi-
cal and sub opical Sou h Ame ica o U uguay, no he n A gen ina, and possibly
Chile; also in sou he n Lesse An illes and epo ed om Jamaica.
Sys ema ics.-
The
ollowing subspecies a e en a i ely ecognized: !ilium
(mos
o
Sou h Ame ica, including T inidad); angeli (Dominica in Lesse An il-
les); luciae (S . Lucia in Lesse An illes); paulsoni (S . Vincen in Lesse An il-
les); pa idens (Mexico sou heas wa d o Colombia); zygoma icus (Ma inique
in Lesse An illes).
The
axa
angeli and paulsoni, o iginally named as species,
a e he e lis ed as subspecies
o
lilium ollowing Koopman (1968) and Jones and
Phillips (1970, 1976).
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
2l
S u lli a ildae de la To e, 1959
Dis ibu ion.-O iginally named om T inidad, his species now
is
known o
be widely dis ibu ed in no he n and cen al Sou h Ame ica, sou h a leas o
Ma o G osso, B azil, and Amazonian Pe u.
Sys ema ics.-S.
ildae
is
amono ypic species.
S u ni a magna de la To e, 1966
Dis ibu ion.-Known
only om Amazon d ainage
in
Colombia, Ecuado , and
Pe u.
Sys ema ics.-S.
magna
is
amono ypic species.
S u ni a mo dax (Goodwin, 1938)
Dis ibu ion.-Reco ded
only om Cos a Rica.
Sys ema ics.-S.
mo dax
is
amono ypic species desc ibed o iginally as he
sole ep esen a i e
o
he genus S u ni ops, possibly alid
a~
asubgenus (see
Da is e al., 1964).
S u ni a bidens (Thomas, 1915)
Dis ibu ion.-Known
only om nea Ta lbo, Colombia, he ype locali y a
Baeza, Ecuado , and cloud o es s
o
eas e n Pe u ian Andes.
Sys ema ics.-S.
bidens
is
amono ypic species and o many yea s was placed
in he genus Co i a. Ga dne and O'Neill (1969) educed Co i a o subgene ic
s a us unde S u ni a.
S u ni a nana Ga dne and O'Neill,
1971
Dis ibu ion.-Known
only om he ype locali y, Huanhuachayo, Ayacucho,
Pe u.
Sys ema ics.-S.
nana
is
amono ypic species in he subgenus Co i a (Ga dne
and O'Neill, 1971).
S u ni a a a a homasi Pe e son and Tamsi , 1968
Dis ibu ion.-Sou hwes e n Colonlbia and Ecuado wes
o
Andes.
Sys ema
ics.-S.
a a a homasi
is
amono ypic species.
S u ni a ludo ici An hony, 1
924
Dis ibu ion.-Wes em
(Sinaloa) and eas e n (Tamaulipas) Mexico sou heas -
wa d h ough Cen al Ame ica a leas o Colombia, Venezuela, Ecuado , and
Pe u; limi s
o
ange poo ly unde s ood owing o con using sys ema ic pic u e
(see below).
Sys ema ics.-
Two subspecies p esen ly ecognized in he li e a u e a e
ludo ici (cen al Mexico o Sou h Ame ica) and occiden alis (wes e n Mexico).
Howe e , se e al named kinds
o
S u ni a (including hondu ensis, bogo ensis,

and opo ophilum) ela ed o ludo ici, bu no cu en ly ecognized in li e a u e,
may, in ac , be alid species
o
subspecies.
Vampy ops do salis Thomas, 1900
Dis ibu ion.-Known
om in e media e ele a ions in Cos a Rica, Panama,
Colombia, Ecuado , Pe u, and ques ionably om Venezuela.
S u ni a e y h omos (Tschudi, 1844)
Dis ibu ion.-P esen ly eco ded only om eas e n slope
o
Andes
in
Pe u,
bu p obably widely dis ibu ed in no he n Sou h Ame ica.
Sys ema ics.-S.
e y h omos
is
cu en ly ega ded
as
amono ypic species.
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Genus VAMPYROPS Pe e s
Vampy ops in uscus Pe e s, 1881
Dis ibu ion.-Colombia
sou h o Pe u and B azil.
Sys ema ics.-
V.
in uscus
is
amono ypic species.
Vampy ops i a us (Pe e s, 1860)
Dis ibu ion.-Known
o occu a in e media e ele a ions (900 o 2600
me e s) om Cos a Rica sou h o Pe u and eas o Venezuela.
Sys ema ics.-
V.
i a us
is
amono ypic species.
Genus
URODERMA
Pe e s
U ode ma biloba um Pe e s, 1866
Dis ibu ion.--Sou he n Mexico (Ve ac uz and Oaxaca), Cen al Ame ica,
Sou h Ame ica as a sou h
as
sou he n Pe u and adjacen Boli ia, and sou h-
eas e n B azil.
Sys ema ics.-Cu en ly ecognized subspecies (Da is, 1968; Bake and Mc-
Daniel, 1972) include: biloba um (eas e n Boli ia, B azil, he Guianas, and Vene-
zuela); con exum (Paci ic e san
o
Middle Ame ica om Nica agua sou heas -
wa d o adjacen Sou h Ame ica); da isi (Paci ic e san
o
Middle Ame ica om
Chiapas o
EI
Sal ado and p obably Hondu as); mola is (Ca ibbean e san
o
Middle Ame ica om Ve ac uz o Cos a Rica); homasi (Ecuado , Pe u, no h-
wes e n Boli ia); ini a um (T inidad).
U ode ma magni os um Da is, 1968
Dis ibu ion.-Chiapas sou heas wa d in Paci ic e san
o
Middle Ame ica
o Panama, and no he n and cen al Sou h Ame ica eas
o
Andes ( epo ed
om no he n Boli ia, B azil, Colombia, eas e n Pe u, eas e n Ecuado , and
Venezuela).
Sys ema ics.-A.
magni os um
is
amono ypic species.
22
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
23
Sys ema ics.-
This species
is
in
need
o
sys ema ic e iew, bu p obably
is
poly ypic, wi h disjunc popula ions occu ing abo e 900 me e s om eas e n
Panama sou h o Pe u and eas in o Venezuela. We ollow Ga dne and Ca e
(1972) and Ca e and Rouk (1973)
in
ou ea men
o
V.
do salis.
Vampy ops au a ius Handley and Fe is, 1972
Dis ibu
ion.-Known
only om he Guiana Highlands
o
Venezuela.
Sys ema ics.-
V.
au a ius
is
ecognized p o isionally
as
a alid species, bu
may p o e o be asynonym
o
V.
do salis.
Vampy ops nigellus Ga dne and Ca e , 1972
Dis ibu ion.-Reco ded
only om Colombia and Pe u, bu p obably occu s
also
in
Ecuado .
Sys ema ics.-
V.
nigellus
is
conside ed o be amono ypic species.
Vampy ops b achycephalus Rouk and Ca e , 1972
Dis ibu ion.-Known
om
ColoITlbia,
Venezuela, Guyana, Amazonian B azil,
Ecuado , and Pe u.
Sys ema ics.-
V.
b achycephalus is he e conside ed
as
mono ypic and o in-
clude
V.
la us and
V.
l.
saccha us
o
Handley and Fe is.
Vampy ops helle i Pe e s, 1867
Dis ibu ion.--Sou he n Mexico sou h h ough Middle Ame ica and no he n
Sou h Ame ica o Pe u, Boli ia, and B azil; also ound on T inidad.
Sys ema ics.-As
poin ed ou
by
Rouk and Ca e (] 972), ce ain di e ences
exis be ween specimens
o
helle i om Mexico sou h h ough Middle Ame i-
ca and hose in Pe u, bu oo
ew
specimens a e a ailable o in e p e hese di -
e ences. The name inca um Thomas, 1912, would apply o Pe u ian specimens
and p obably o he Amazonian ma e ial should he di e ences p o e o be
o
subspeci ic impo .
V.
za hinus is conside ed o be asynonym
o
V.
helle i, and
he holo ype o ha e come om Panama.
Vampy ops linea us
(E.
Geo oy S .-Hilai e, 1810)
Dis ibu ion.-Repo ed
om Cen al (Ma o G osso) and eas e n (Bahia)
B azil sou h o U uguay, Pa aguay, Boli ia, and no he n A gen ina (Chaco).
Al hough eco ded by se e al au ho s om locali ies in wes e n Sou h Ame ica,
hese epo s e iden ly e e o o he species
o
ba s.
Sys ema ics.-
V.
linea us
is
amono ypic species
as
p esen ly unde s ood, bu
appea s closely allied o
V.
eci inus, wi h which i may be conspeci ic.
Vampy ops eci inus Thomas, 1901
Dis ibu ion.-Known
om he B azilian s a e
o
Pe nambuco and pu po ed
o occu
in
hose
o
Bahia and Sao Paulo.
Sys ema ics.-P o isionally ecognized
as
amono ypic species closely ela ed
o
V.
linea us, om which i may no be dis inc e en a he subspeci ic le el.
Vampy essa bidens (Dobson, 1878)
Dis ibu ion.-Ecuado , Pe u, Colombia, no he n B azil, and Guyana.
Vampy essa b ocki Pe e son, 1968
Dis ibu ion.-P esen ly
known only om Guyana and Colombia.
Sys ema ics.-
V.
b ocki
is
amono ypic species.
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Genus V
AMPYRESSA
Thomas
Vampy essa pusilla (Wagne , 1843)
Dis ibu ion.-Sou he n
Mexico (Ve ac uz), Cen al Ame ica, no he n and
cen al Sou h Anle ica sou h a leas o sou heas e n B azil and Pe u.
Sys ema ics.-Goodwin
(1963), who e iewed he genus Vampy essa, ecog-
nized h ee subspecies (pusilla, hyone, and enUla). La e , Pe e son (1968), in
his synopsis
o
he genus, lis ed only wo ( ega ding enUla as indis inc om
hyone). Handley (1966) did no ecognize subspecies in
V.
pusilla.
The
species
na e e i, named by Goodwin (op. ci .), is he e ega ded as asynonym
o
pusilla
ollowing Pe e son (op. ci .).
Genus V
AMPYRODES
Thomas
Vampy odes ca accioloi (Thomas, 1889)
Dis ibu ion
..
--Sou he n
Mexico (Oaxaca, Ve ac uz) sou heas wa d h ough
Cen al Ame ica o Sou h Ame ica as a sou h as no he n B azil and Ama-
zonian Pe u.
Sys ema ics.-Two
subspecies a e ecognized, ollowing Handley (1966):
ca accioloi (T inidad and Tobago, and adjacen egions
o
no heas e n Sou h
Ame ican mainland); majo (Mexico o Pe u, o na us asynonym). Some ecen
au ho s ha e ega ded majo
as
aspecies dis inc om, bu closely ela ed o,
ca accioloi.
Vampy essa nymphaea Thomas, 1909
Dis ibu ion.-Repo ed
om Nica agua, Cos a Rica, Panama, and wes e n
Colombia.
Sys ema ics.-
V.
nymphaea is amono ypic species ep esen ing, along wi h
V.
b ocki, he subgenus Me a ampy essa.
Vampy essa melissa Thomas, 1926
Dis ibu ion.-Known
only om eas e n slope
o
Andes
in
Pe u.
Sys ema ics.-
V.
melissa is amono ypic species, which oge he wi h
V.
pusilla
cons i u es he subgenus Vampy essa.
24
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
25
Sys ema ics.-
V.
bidens
is
amono ypic species and he sole ep esen a i e
o
he subgenus Vampy iscus, which has been used in he gene ic sense
by
some
ecen au ho s.
Genus
CHIRODERMA
Pe e s
Chi ode ma do iae Thomas,
1891
Dis ibu ion.-Eas e n
B azil (Minas Ge ais).
Sys ema ics.-C.
do iae is amono ypic species.
Chi ode ma imp o isum Bake and Genoways, 1976
Dis ibu ion.-Known
only om he Lesse An illean island
o
Guadeloupe.
Sys ema ics.-C.
imp o isum is amono ypic species known only om he
holo ype.
Chi ode ma illosum Pe e s, 1860
Dis ibu ion.--Sou he n Mexico (Oaxaca and Ve ac uz) sou h o Pe u,
Boli ia, and B azil.
Sys ema ics.-
Two subspecies a e ecognized, illosum (T inidad and adjacen
Venezuela sou h o Pe u and B azil) and jesupi (Mexico sou h h ough Cen al
Ame ica o no he n Colombia).
Chi ode ma sal ini Dobson, 1878
Dis ibu ion.-Wes e n
Mexico (Chihuahua) sou h o Colombia and Ecuado .
Sys ema ics.-
Two subspecies cu en ly a e ecognized, sal ini (Puebla, Mexi-
co, sou h o no he n Sou h Ame ica) and scopaeum (wes e n Mexico om Chi-
huahua sou h o Gue e o).
Chi ode ma ini a um Goodwin, 1958
Dis ibu ion.-Panama
eas o T inidad and sou h o Pe u, Boli ia, and B azil
(Ma o G osso).
Sys ema ics.-
Two subspecies a e ecognized, ini a um (T inidad and
Ama-
zonian Sou h Ame ica sou h o Pe u and B azil) and go gasi (Da ien, Panama,
eas o Venezuela).
Genus
ECTOPHy.LLA
H. Allen
Ec ophylla alba H. Allen, 1892
Dis ibu ion.-Known
only om Nica agua, Cos a Rica, and wes e n Panama.
Sys ema ics.-E.
alba
is
amono ypic species.
Ec ophylla macconnelli Thomas, 1901
Dis ibu ion.--Sou h Ame ica ( epo ed om Anlazonian Ecuado and
Pe u, Boli ia, B azil, Colombia, Venezuela, Guyana, and T inidad) and epo ed
om Cos a Rica and Panama in No h Ame ica.
4.
Two
lowe
p emola s
.........................................
..
Phyllos omus
Th ee
lowe
p emola s
(second
some imes
c owded
o
lingual
side
o
oo h ow)
5
5.
Ros um
as
long
as
b aincase
Vampy um
Ros um
sho e
han
b aincase
............................................. 6
6.
Second
lowe
p emola
la ge,
subequal
in
size
o
i s
and
hi d
p emola s
7
Second
lowe
p emola
small
o
minu e,
much
smalle
han
i s
and
hi d
p emola s
..
· 8
Phyllos oma inae
1.
One
lowe
inciso
2
Two
lowe
inciso s
4
2.
Two
lowe
p emola s
Mimon
Th ee
lowe
p emola s
(second
small
o
minu e)
3
3.
Second
lowe
p emola
c owded
o
lingual
side
o
oo h ow,
i s
and
hi d
lowe
p e-
mola s
usually
in
con ac
Ch o op e lls
Second
lowe
p emola
no
c owded
om
oo h ow,
i s
and
hi d
lowe
p emola s
no
in
con ac
........................................................
..
Tona ia
2.
Noselea
udimen a y,
wi hou
dis inc
up igh
p ocess;
ail
p esen
Phyllonyc e inae
Noselea
usually
well
de eloped;
ail
absen
i
noselea
udimen a y
3
3.
Tongue
elonga e,
wi h
conspicuous
b is lelike
papillae
on
an e odo sal
su ace;
i s
uppe
p emola
usually
dis inc ly
sepa a ed
om
canine
and
a ely
in
con ac
wi h
second
uppe
p emola
( i s
uppe
p emola
some imes
in
con ac
wi h
canine
in
Mono-
phY//llS,
bu
dis inc ly
sepa a ed
om
second
uppe
p emola )
Glossophaginae
Tongue
no
elonga e,
lacking
conspicuous
b is lelike
papillae;
i s
uppe
p emola
in
con ac
wi h
canine
and
usually
wi h
second
uppe
p emola
4
4.
Zygoma ic
a ch
incomple e
Ca o//iinae
Zygoma ic
a ch
comple e
5
5.
Mola s
dilambdodon
(dis inc
W-shaped
pa e n
o
lophs
on
occlusal
su ace)
.
·.......................................................
..
Phyllos o 11a inae
Mola s
lacking
dilambdodon
pa e n
S enode minae
SPECIAL
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7.
Audi o y
bullae
la ge,
g ea es
diame e
much
exceeding
dis ance
be ween
hem
.....
·..............................................................
..
Mac o is
Audi o y
bullae
small,
g ea es
diame e
less
han
dis ance
be ween
hem
.
·..........................................................
..
Mic onyc e is
8.
Second
lowe
p emola
displaced
lingually
om
oo h ow;
i s
and
second
lowe
p e-
mola s
in
con ac
o
nea ly
so 9
Second
lowe
p emola
no
displaced
lingually
om
oo h ow;
i s
and
second
lowe
p emola s
usually
no
in
con ac
10
9.
G ea es
leng h
o
skull less
han
20
mm
Mac ophyllum
G ea es
leng h
o
skull
mo e
han
20
mm.
T achops
10.
Do sal
p o ile
o
os um
s ongly
con ex;
deep
dep ession
p esen
be ween
o bi s
.....
·...........................................................
..
Loncho hina
Do sal
p o ile
o
os um
no
con ex;
no
dep ession
be ween
o bi s
Phy//ode ma

BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
33
Glossophaginae
1.
Pe manen
lowe inciso s lacking 2
Two
pai s
o
pe manen
lowe
inciso s, usually
weJl
de eloped 8
2.
P emola s
3/3
Anou a
P emola s 2/3 3
3.
Mola s 2/2
Lichonyc e is
Mola s
3/3 4
4. P e ygoids highly modi ied,
expanded
a base and in la ed in
appea ance;
p e ygoid
wings long
and
in con ac ,
o
nea ly so, wi h
audi o y
bullae 5
P e ygoids
no mal,
no
expanded
a base
o
in la ed in
appea ance:
p e ygoid wings
sho
and
no in
con ac
wi h
audi o y
bullae 7
5.
Fi s and second
uppe
inciso s sepa a ed
by
dis inc gap;
uppe
p emola s
low, ba ely
exceeding heigh
o
mola s
Choe oniscus
Fi s
and
second
uppe
inciso s in con ac ,
o
nea ly so;
uppe
p emola s
dis inc ly
highe
han
mola s
6
6.
Ros um
dis inc ly
longe
han
pos os al pa
o
c anium:
uppe
mola s
essen ially
equal
in
size, all wi h adis inc me as yle
Musonyc e is
Ros um
abou equal in leng h o pos os al pa
o
c anium;
hi d
uppe
mola
some-
wha
smalle
han
i s wo and lacking adis inc me as yle
Choe onycle is
7.
Uppe
mola s
lacking mesos yle; lowe
mola s
long
and
na ow;
known
only
om
Middle
Ame ica
Hylonycle is
Mesos yle p esen
on
all
uppe
mola s;
lowe
mola s
only
mode a ely
comp essed:
known
only om B azil
and
Venezuela
Scle onyc e is
8.
Mola s 2/2
Lep onyc e is
Mola s 3/3 9
9.
Zygoma ic
a ch comple e, i s
uppe
inciso
no
ma kedly
enla ged
and
spa ula e
10
Zygoma ic
a ch incomple e, i s
uppe
inciso enla ged
and
spa ula e
11
10. E iden
gap
be ween
uppe
p emola s
and be ween
hem
and
adjacen ee h; ail ela-
i ely long
and
ex ending
beyond
pos e io
bo de
o
u opa agium
Monophyll
us
Uppe
p emola s
usually in
con ac
and illing space be ween
canine
and
i s mola :
ail sho
and
no ex ending beyond
pos e io
bo de
o
u opa agium
.....
Glossophaga
11.
Ros um
elonga e,
longe
han
pos os al pa
o
c anium;
pos canine
maxilla y ee h
educed in size
and
wi h e iden gaps be ween
hem
Pla alina
Ros um
no elonga e,
no
longe
han
pos os al
pa
o
c anium;
pos canine
maxilla y
ee h
o
no mal size, las
p emola
and
mola s
in
con ac
(o
nea ly so)
12
12. Fi s
uppe
p emola
smalle
han
second and la e ally comp essed
Lonchophylla
Fi s
uppe
p emola
essen ially
same
size as second, no la e ally
comp essed
( iangula
in ou line)
Lionyc e is
Ca olliinae
1.
Tail p esen ;
uppe
p emola s
essen ially equal in size Ca ollia
Tail absen ; i s
uppe
p emola
much
smalle
han
second . .
..
Rhinophylla
]0. Second
uppe
mola
no iceably
la ge
han
i s ;
uppe
p emola s
sepa a ed
om each
o he
and
om adjacen ee h by e iden gaps
Ec ophylla
(pa )
Second uppe
mola
equal in size o,
o
smalle
han, i s ; no gaps be ween
an e io
uppe
cheek ee h
11
II.
Inciso s 2/l
o
2/2; heigh
o
i s inciso
g ea e
han
heigh
o
i s
p emola ;
g ea es
leng h
o
skull less
han
22
Vampy essa
(pa )
Inciso s 2/2; heigh
o
i s inciso
much
less
han
heigh
o
i s
p emola ;
g ea es
leng h
o
skull
mo e
han
24
Vampy odes
12.
Uppe
den al
a cade
expanded la e ally
o
o m
semici cula
a c
13
Uppe
den al
a cade
no expanded la e ally, U-shaped in occlusal iew
14
13. O bi al space wide
han
long; in e o bi al
cons ic ion
less
han
5
Ame ida
O bi al space
longe
han
wide; in e o bi al
cons ic ion
mo e
han
5 .
·
Sphae onyc e is
14.
Pala e
sho ,
pos e io
pala al
ema gina ion
eaching le el
o
i s
uppe
mola
.....
15
Pala e
o
medium
leng h
o
long,
pos e io
bo de
a iously
ema gina e
bu ne e
o
le el
o
oo h ow
......................................................
..
17
S enode minae
]. Mola s 2/2 2
Mola s 2/3
o
3/3 7
2.
Uppe
den al a cade semici cula ,
os um
less
han
hal
as long as b aincase .
·..............................................................
Cen u io
Uppe
den al
a cade
no semici cula ,
os um
mo e
han
hal
as long as b aincase
...
3
3.
Ros um in la ed, nea ly cuboid in o m ,
Pygode ma
Ros um
no in la ed
o
cuboid in o m 4
4.
Pos e io
ma gin
o
ex e nal na es wi h
ma ked,
ly e-shaped
ema gina ion
.
·............................................................
..
CII
i ode l11a
Pos e io
ma gin
o
ex e nal na es lacking ly e-shaped
ema gina ion
5
5.
Second
uppe
mol
a
ma kedly
la ge
han
i s ;
uppe
p emola s
sepa a ed om each
o he
and adjacen ee h by e iden gaps . . . . . . . . . . . . . . . . . . . . . .
..
Ec ophylla
(pa )
Second
uppe
mola
essen ially equal in size o,
o
smalle
han,
i s ; no gaps be ween
an e io
uppe cheek ee h 6
6.
Pos e io
ma gin
o
ex e nal na es
mo e
o
less s aigh ; second
uppe
mola
much
smalle
han
i s and di e ing in o m
A ihells
(pa )
Pos e io
ma gin
o
ex e nal na es b oaQly V-shaped; second
uppe
mola
esembling
i s in size and o m
..
...................................
..
Vampy essa
(pa )
7.
Mola s
2/3
8
Mola s 3/3
12
8.
Pala e
sho ,
pos e io
bo de
ha ing deep U-shaped
ema gina ion
ha
eaches le el
o
i s
mola
A i eus
Pala e
long,
pos e io
bo de
ha ing shallow
ema gina ion
ha alls a sho
o
le el
o
oo h ow
..............................................................9
9.
Fi s
uppe
inciso
ma kedly
bi id, less
han
wice size
o
second inciso .
·
A ibells
(pa )
Fi s
uppe
inciso no bi id
o
only weakly so,
mo e
han
wice size
o
second
inciso .
.
.......................................................................
]0
34
SPECIAL
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PHYLLOSTOMATIDAE
35
15.
Pala al
ema gina ion
b oadly V-shaped
Phyl/ops
Pala al
ema gina ion
deeply U-shaped
16
J
6.
Well-de eloped V-shaped idge om sagi al c es o
an e io
ma gin
o
o bi s, o ming
deep os al dep ession . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . .
..
S enode ma
V-shaped idge om sagi al c es o
an e io
ma gin
o
o bi s lacking,
os um
no mal
A d()ps
17.
Uppe
mola s dis inc ly g oo ed longi udinally,
he
i s wo
subquad a e
in
ou line
and
lacking well-de eloped cusps; i s
uppe
inciso
app oxima ely
hal
as high as
canine
S u ni a
Uppe
mola s lacking longi udinal g oo e, he i s wo no
subquad a e
in
ou line
and
possessing well-de eloped cusps; i s
uppe
inciso much less
han
hal
as high as
canine
18
18. Fi s uppe inciso less
han
wice size
o
second and esembling i in shape:
uppe
in-
ciso s in
con ac
and
ill ing space be ween canines
19
Fi s
uppe
inciso
mo e
han
wice size
o
second and di e ing om i in shape; e iden
gaps p esen be ween
uppe
inciso s .........................................20
19.
Fi s
uppe
inciso deeply bi id; m3, i p esen ,
minu e
and
peglike·
A ihells
(pa )
Fi s
uppe
inciso no bi id; m3 ela i ely la ge and well de eloped
Enchis henes
20.
C owns
o
i s
uppe
inciso s pa allel, deeply bi id; lowe inciso s in
con ac
.
·.............................................................
..
U ()de I11a
C owns
o
i s
uppe
inciso s con e ge dis ally, no deeply bi id; lowe inciso s sepa-
a ed by dis inc
gaps.
..........................................
..
Vampy ops
Phyllonyc e inae
1.
Tail no ex ending beyond edge
o
u opa agium
B achyphylla
Tail ex endin_g beyond edge
o
u opa agium
2
2.
Zygoma ic a ch comple e; second
and
hi d
lowe
mola s dis inc ly cuspida e .
·
E ophylla
Zygoma ic a ch incomple e; second and
hi d
lowe mola s no dis inc ly cuspida e
....
·..........................................................
..
Phyllonyc e is
Desmodon inae
1.
Fi s lowe inciso s in con ac ; in e emo al
memb ane
wi h dis inc inge
o
mode a ely
long hai s
Diphylla
Fi s
lowe inciso s no in con ac ; in e emo al
memb ane
wi hou inge
o
hai
2
2.
Lowe
inciso s no bi id; wing
whi e
om middle
o
p oximal
phalanx
o
ip .
·..............................................................
..
Diaelnlls
Lowe inciso s bi id; wing usually pigme ed
o
ip
(i
whi e- ipped, whi e
does
no
ex end p oximally
o
i s phalanx) .................................
..
Desl110d
us
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Philande
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36
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TEXAS
TECH
UNIVERSITY
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
37
GOODWIN,
G.
G.
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1.
HALL,
E. R.,
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K.
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The
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No h
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P ess,
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Yo k,
]:xxx+
1-546+
79.
HANDLEY,
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0.,
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Desc ip ions
o
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o
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]967. Ba s
o
he
canopy
o
an
Amazonian
o es .
A as
do
simposio
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a
bio a
Amazonica,
5:2 1
1-2
15.
HILL,
J. E. ]964.
No es
on
ba s
om
B i ish
Guiana,
wi h
he
desc ip ion
o
anew
genus
and
species
o
Phyllos omidae.
Mammalia,
28:553-572.
HUSSON,
A.
M.
]962.
The
ba s
o
Su iname.
Zoo!. Ve hand., 58: 1-282.
IRWIN,
D. W.,
AND
R.
J.
BAKER.
]967.
Addi ional
eco ds
o
ba s
om
A izona
and
Sinaloa.
Sou hwes e n
Na .,
12: 195.
JONES,
J. K.,
JR.
1966. Ba s
om
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Uni .
Kansas
Publ., Mus.
Na .
His .,
16:439-472.
JONES,
J. K., JR.,
AND
C. J.
PHILLIPS.
1970.
Commen s
on
sys ema ics and zoogeog aphy
o
ba s
in
he
Lesse An illes.
S udies
on
he
Fauna
o
Cu acao
and
o he
Ca ib-
bean
Islands, 32:131-145.
1976. Ba s
o
he
genus
S u ni a
in
he
Lesse An illes. Occas.
Pape s
Mus.,
Texas
Tech
Uni .,
40:]-16.
JONES,
J. K., JR.,
AND
A.
SCHWARTZ.
1967.
B edin-A chbold-Smi hsonian
Biological
Su -
ey
o
Dominica.
6.
Synopsis
o
ba s
o
he
An illean
genus
A dops.
P oc.
U.S.
Na .
Mus., 124(3634):1-13.
JONES,
J. K., JR., H.
H.
GENOWAYS,
AND
R.
H.
BAKER.
1971a.
Mo phological
a ia ion
in
S enade ma
u um. J.
Mamm.,
52:244-247.
JONES,
J.
K., JR., J. D.
SMITH,
AND
R. W.
TuRNER.
1971
b.
No ewo hy
eco ds
o
ba s
om
Nica agua,
wi h
achecklis
o
he
chi op e an
auna
o
he
coun y.
Occas.
Pape s
Mus.
Na .
His ., Uni . Kansas, 2: 1-35.
JONES,
J. K., JR., J.
R.
CHOATE,
AND
A.
CADENA.
1972.
Mammals
om
he
Mexican
s a e
o
Sinaloa.
II.
Chi op e a.
Occas.
Pape s
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Na .
His ., Uni .
Kansas,
6:
1-29.
JONES,
J. K., JR.,
J.
D.
SMITH,
AND
H.
H.
GENOWAYS.
1973.
Anno a ed
checklis
o
mam-
mals
o
he
Yuca an
Peninsula,
Mexico.
I.
Chi op e a.
Occas.
Pape s
Mus.,
Texas
Tech
Uni ., 13:1-31.
KOOPMAN,
K. F. 1958.
Does
pygode ma
occu
in
No h
Ame ica?
J.
Mamm.,
39:
584-585.
1968.
Taxonomic
and
dis ibu ional
no es
on
Lesse
An illean
ba s.
Ame .
Mus.
No i ., 2333:1-13.
MARINKELLE,
C.
J.,
AND
A.
CADENA.
1972.
No es
on
ba s
new
o
he
auna
o
Colombia.
Mammalia,
36:50-58.
MATSON,
J.
0.,
AND
D.
R.
PATTEN.
1975.
No es
on
some
ba s
om
he
s a e
o
Zaca ecas,
Mexico.
Con ib.
Sci., Los
Angeles
Co.
Mus.
Na .
His ., 263:1-12.
OJASTl, J.,
AND
O. J.
LINARES.
1971.
Adiciones
de
la
auna
de
mu cieJagos
de
Venezuela
con
no as
sob e
las especies del
gene o
Diclidu us
(Chi op e a).
Ac a
BioI.
Venezolana,
7:421-441.
PARADISO,
J. L. 1967. A e iew
o
he
w inkle- aced
ba s
(Cen u ia
senex
G ay),
wi h
desc ip ion
o
a
new
subspecies.
Mammalia,
31
:595-604.

PERACTHI,
A. L.,
AND
S.
T.
DE
ALBUQUERQU
E.
1971.
Lis a
p o iso ia
dos
qui op e os
dos
es
ados
do
Rio
de
Janei o
e
Guanaba a,
B asil
(Mammalia,
Chi op e a).
Re .
B asil. BioI.,
31
:405-413.
PETERSON,
R. L. 1965. A
e iew
o
he
ba s
o
he
genus
Ame ida,
amily
Phyllos omi-
dae.
Con ib.
Li e Sci.,
Royal
On a io
Mus.,
65:1-13.
1968. A
new
ba
o
he
genus
Vampy essa
om
Guyana,
Sou h
Ame ica.
Con-
ib.
Li e
Sci., Royal
On a io
Mus., 73:1-17.
PHILLIPS,
C.
J.
1971.
The
den i ion
o
glossophaginae
ba s:
de elopmen ,
mo phological
cha ac e is ics,
a ia ion,
pa hology,
and
e olu ion.
Misc. PubJ. Mus.
Na .
His .,
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54:1-138.
PHILLIPS,
C.
J.,
AND
J.
K.
JONES,
JR. 1971. A
new
subspecies
o
he
long-nosed
ba ,
Hylonyc e is
lInde l 'oodi,
om
Mexico.
J.
Mamm.,
52:77-80.
PINE,
R. H. 1972.
The
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o
he
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Ca o//ia.
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Ag ic.
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PINE,
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I.
R.
BISHOP,
AND
R. L.
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1970.
P elimina y
lis
o
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o
he
Xa an ina/Cachimbo
expedi ion
(cen al
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Royal
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T op.
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Hygiene,
64:668-670.
PIRLOT,
P.
1967.
Nou elle
ecol e
de
chi op e es
dans
oues
du
Venezuela.
Mammalia,
31
:260-274.
POWER,
D.
M.,
AND
J.
R.
TAMSITT.
1973.
Va ia ion
in
Phyllos omus
discolo
(Chi op e a:
Phyllos oma idae).
Canadian
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Zool.,
5]
:461-468.
ROUK,
C. S.,
AND
D.
C.
CARTER.
1972. A
new
species
o
Vampy ops
(Chi op e a:
PhyJ-
los oma idae)
om
Sou h
Ame ica.
Occas.
Pape s
Mus.,
Texas
Tech
Uni .,
1:
1-7.
SCHWARTZ,
A.,
AND
J.
K.
JONES,
JR. 1967.
B edin-A chbold-Smi hsonian
Biological
Su ey
o
Dominica.
7.
Re iew
o
ba s
o
he
endemic
An illean
genus
Mono-
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P oc.
U.S.
Na .
Mus.,
124(3635):1-20.
SILVA
TABOADA,
G.,
AND
R. H.
PINE.
1969.
Mo phological
and
beha io al
e idence
o
he
ela ionship
be ween
he
ba
genus
B achyphylla
and
he
Phyllonyc e inae.
Bio opica,
1:10-19.
SMITH,
J.
D. 1972.
Sys ema ics
o
he
chi op e an
amily
Mo moopidae.
Misc.
Pub
I.
Mus.
Na .
His ., Uni .
Kansas,
56: 1-132.
STARRETT,
A.,
AND
R.
S.
CASEBEER.
1968.
Reco ds
o
ba s
om
Cos a
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Con ib.
Sci.,
Los
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Co.
Mus.,
]48:1-21.
TUTTLE,
M.
D. 1970.
Dis ibu ion
and
zoogeog aphy
o
Pe u ian
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wi h
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on
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Kansas
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49:45-86.
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L.
S.
1974.
Ca alogo
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i ien es
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ex inguidos
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ill
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iii +1-139 pp.
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Nac.
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AND
M.
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CORNEJO.
1969.
Algunos
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del
no e
de
A gen ina.
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Na .
His .,
Uni .
Kansas,
51
:407-428.
WATKINS,
L.
C.,
J.
K.
JONES,
JR.,
AND
H. H.
GENOWAYS.
1972.
Ba s
o
Jalisco,
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Texas
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Uni .,
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XIMENEZ,
A.,
A.
LANGGUTH,
AND
R.
PRADERI.
1972.
Lis a
sis ema ica
de
los
mami e os
del
U uguay.
An. Mus.
Nac.
His .
Na .
Mon e ideo,
se .
2,7(5):1-49.
38
SPECIAL
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UNIVERSITY
ZOOGEOGRAPHY
KARL
F.
KOOPMAN
O
he nine amilies
o
ba s
in
he Wes e n hemisphe e, h ee (Emballonu idae,
Vespe ilionidae, Molossidae) a e ound also in he Old Wo ld.
O
he six
endemic New Wo ld (and chie ly Neo opical) amilies, he Phyllos oma idae
is
by
a he la ges g ouping. The Noc ilionidae, Fu ip e idae, and Thy op e idae
ha e only wo species each, he Na alidae p obably only six species, and he
Mo moopidae eigh species (see Smi h, 1972). The Phyllos oma idae, howe e ,
ha e 136 species as ecognized in he classi ica ion in his olume.
Excep o a
ew
a eas (Wes Indies, sou heas e n B azil, no he n Mexico)
e en he Pleis ocene ossil eco d
o
ba s o he Neo opical egion
is
poo .
To
my
knowledge he only p e-Pleis ocene eco d
o
ba s in he Neo opical egion
is
o
No onyc e is (a phyllos oma id) om he Miocene
o
Colombia. This shows
ha he Phyllos oma idae we e in Sou h Ame ica by a leas ha ime. In he
Miocene, Sou h Ame ica was s ill
an
island con inen sepa a ed om o he con i-
nen s by ocean ba ie s. Judging by p esen di e si y in Sou h Ame ica,
i
is
likely ha he Phyllos oma idae was he i s chi op e an amily o each Sou h
Ame ica du ing i s long pe iod
o
isola ion, and may e en ha e o igina ed as a
amily on ha con inen . The o he i e amilies mo e
o
less con ined o he
Neo opical egion ha e oo
ew
species o any hing e y de ini e o be said
abou hei a ea
o
o igin (see Koopman, 1970). While i
is
hus p obable ha
Sou h Ame ica was he p ima y cen e
o
phyllos oma id e olu ion, i is clea
ha bo h Middle Ame ica and he Wes Indies ha e been impo an seconda y
cen e s.
In
he ollowing sec ions, a e asho discussion
o
dis ibu ion
o
he sub-
amilies, he a ious special egions
o
Sou h Ame ica, Middle Ame ica, and he
Wes Indies will be aken up
in
u n wi h adiscussion
o
he phyllos oma ids
ound wi hin each
o
hem I should be emphasized ha oceanic s ai s
as
well
as
high moun ains and cold empe a e lowland a eas (such as Pa agonia) cons i u e
o midable ba ie s o he Phyllos oma idae.
DISTRIBUTION
OF
MAJOR
PHYLLOSTOMATID
GROUPS
O
he six sub amilies
in
o which he Phyllos oma idae a e cu en ly di ided,
all bu he Phyllonyc e inae a e widely dis ibu ed
in
Sou h and Middle Ame ica.
The phyllonyc e ines a e endemic o he Wes Indies, which also ha e phyl-
los onla ines, glossophagines, and s enode mines. Vampi es a e known in he
ossil eco d
o
Cuba, bu he e a e no ce ain eco ds
o
ca olliines in he Wes
Indies as de ined he e. Se e al gene a
o
glossophagines and s enode mines a e
con ined o he An illes.
On he mainland, many gene a and e en species a e dis ibu ed o e ala ge
pa
o
he o al ange
o
he amily. O he s, pa icula ly
in
he Glossophaginae
(and o alesse ex en he S enode minae), ha e es ic ed anges ( o example,
39
Pla alina).
I
should also be men ioned. ha he S u ni ini (all now included in he
genus S u ni a) a e s ongly concen a ed in he no he n Andean egion wi h
ela i ely ew species a away om
i .
Fo
u he in o ma ion
on
dis ibu ion
o
indi idual species, he eade
is
e e ed o he p eceding a icle in his olume
and he appendix
o
his pape .
REGIONAL
BAT
FAUNAS
Sou h Ame ica
Pa agonian
sub egion.-
This
is
ha
po ion
o
Sou h Ame ica sou h
o
he opical o es s. I has ne e been p ecisely de ined bu would ce ainly in-
clude, o
ou
pu poses, all
o
Chile and U uguay, also all
o
A gen ina excep
small po ions
o
he no heas and no hwes .
The
high Andes
and
al iplano
o
wes e n Boli ia and sou he n Pe u also would be included as well as d ye a eas
o
wes e n Pa aguay, sou he n Boli ia, and ex eme sou heas e n B azil. Al-
hough he e a e se e al species
o
ba s ha a e mos ly con ined o he Pa agonian
egion, all a e espe ilionids. A
ew
species
o
phyllos oma ids, he main ange
o
which lies a he no h, do each he Pa agonian sub egion o alimi ed deg ee,
bu only six species eally pene a e he sub egion.
O
hese, only one, Desmodus
o undus, eaches any dis ance
sou hwa d- o
cen al A gen ina and e en
cen al Chile. While i
is
possible ha his may in
pa
e lec he man-made a ail-
abili y
o
ca le as ood, his
is
by no means ce ain. S u ni a /ilium also eaches
cen al A gen ina, bu an old eco d om Chile is appa en ly e oneous.
O
he o he ou species, Ch o op e us au i us, Glossophaga so icina, and A ibeus
li u a us ange no a he han no he n A gen ina, whe eas Vampy ops
linea us has ecen ly been eco ded om U uguay.
Eas e n B azilian highlands and
coas .-
The
d y chaco zone
o
no he n
A gen ina, wes e n Pa aguay, and sou heas e n Boli ia con inues in modi ied
o m as he caa inga, abel
o
sc ub o es ( eally asa anna), which eaches he
A lan ic coas ali le o he wes
o
eas e nmos B azil. This isola es he moun-
ain and coas al o es s
o
eas e n B azil, eas e n Pa aguay, and no heas e n
A gen ina om hose
o
he Amazon basin. As a esul , he e a e anumbe
o
mammals, pa icula ly p ima es and oden s,
ha
a e con ined o his eas enl
a ea. Howe e , ou
o
36 species
o
phyllos oma ids known om he egion, only
ou (Tona ia b asiliense, Vampy ops eci inus, Chi ode ma do iae, and possibly
Lonchophylla mo dax) a e, as a p esen ly ecognized, endemic o i . In iew
o
he ac ha ba s a e able o
ly
ac oss sho s e ches
o
un a o able habi a ,
howe e , his
is
no su p ising. The e a e, on he
o he
hand, some
42
species
o
phyllos oma ids
in
he Amazon basin ha a e no known om he eas e n B azili-
an highlands and coas al a ea, al hough some may e en ually be ound he e.
Anlazon
Basin.-This
ep esen s pe haps he eaJ hea land
o
he Neo opi-
cal egion.
I
includes he en i e Amazon d ainage
o
B azil and also includes
no heas e n Boli ia (wi h an ex ension along he eas e n ace
o
he Andes in o
no hwes e n A gen ina), he eas e n lowlands
o
Pe u and Ecuado , he Amazon
and O inoco d ainages
o
Colombia and Venezuela, and also he Guianas. Some
40
SPECIAL
PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
BIOLOGY OF
THE
PHYLLOSTOMATIDAE
41
74 species
o
phyllos oma ids a e known om his a ea, 14
o
which appa en ly
a e endemic. Howe e , he lis
o
endemics (Mic onyc e is behni,
M.
da iesi,
Loncho hina o inocensis, Tona ia ca ike i, Phyllos o nus la i olius, Scle onyc-
e is ega, Choe oniscus inca, Lichonyc e is degene , Rhinophylla ische ae,
Vampy ops in uscus,
V.
au a ius, Vampy essa b ocki,
V.
bidens, and A ibeus
concolo ) includes se e al species ha a e poo ly known
o
o
dubious alidi y.
Fou
o he species
(Mimon
benne i, Phyllos omus elonga us, Rhinophylla
pumilio, and Pygode ma bilabia um) a e sha ed only wi h he eas e n B azilian
a ea. Mo e axonomic wo k undoub edly will change conside ably he igu es
bo h o o al numbe and numbe
o
endemics.
Eas e n slopes
o
he no he n
Andes.-
The
uppe o es ed slopes on he
eas e n side
o
he Andes om Colombia o Boli ia, while ecologically con inuous
wi h he Amazonian lowlands o he eas , a e en i onmen ally dis inc i e o he
ex en ha many lowland species ex end only o alimi ed ex en up hese slopes,
whe eas o he species a e con ined o highe ele a ions. Un o una ely he al i-
udinal dis ibu ions
o
phyllos oma ids a e no well known
in
mos
o
his
Andean bel and, a p esen , i
is
only abou Pe u ha much can be w i en. I
ha e chosen o igno e species ha do no occu abo e abou 1000 me e s bu in-
clude as endemics all species ha a e con ined o ele a ions abo e 500 me e s.
Using hese c i e ia, some 25 species a e known om he uppe slopes
o
he
eas e n Andes, and h ee
o
hese
(Mimon
koepkeae, S u ni a nana, and
Vampy essa melissa) a e endemic. The e would be mo e endemic species (such
as
S u ni a e y h omos and
S.
bidens) i highe ele a ions
o
he in e nal
Andean alleys
o
Colombia we e included.
No he n coas and
islands.-I
would de ine his a ea on he mainland as ex-
ending om he no he n end
o
he Co dille a Occiden al
Uus
eas
o
he
Gul
o
U aba) eas along he coas o he Pa ia Peninsula in no heas e n Venezuela.
In Venezuela, i would include only a a he na ow coas al s ip including he
moun ains di ec ly o he sou h, bu in Colombia would ex end up he i e
alleys be ween he co dille as, bu no wes
o
he Co dille a Occiden al no eas
o
he Co dille a O ien al.
The
bounda ies a e mos di icul o d aw in
Colombia. He e he highe ele a ions in he in e nal Andean alleys a e pe haps
be e placed wi h eas e n slope highlands.
The
lowlands
o
he
Cauca
Valley,
on he o he hand, a e almos equally well placed wi h lowlands
o
he Paci ic
coas . Howe e , in he absence
o
ag ea deal mo e de ailed dis ibu ional in o -
ma ion, Iha e been unable o d aw abe e bounda y. Anumbe
o
islands a e
also included, chie ly A uba,
Cu a~ao,
Bonai e, Ma ga i a, T inidad, Tobago,
and G enada. Iha e published p e iously on he ba aunas
o
hese islands
(Koopman, 1958), bu anumbe
o
species ha e been eco ded since. Ip e iously
(Koopman, 1959) ea ed G enada and he G enadines as pa
o
he Wes
Indies, bu , as explained in he Lesse An illean sec ion, Ibelie e hese islands
a e be e placed he e. Pe haps he g ea es signi icance
o
his no he n coas and
island a ea
is
ha anumbe
o
he species occu ing he e ha e a ini ies wi h
Cen al Ame ica (and some imes he Wes Indies)
a he
han wi h he Amazon
Basin. Some 64 species a e known om his a ea. Al hough only Lep onyc e is
CHIROPTERAN
EVOLUTION
JAMES
DALE
SMITH
One
o
he mos in iguing p oblems in e eb a e e olu ion
is
he e olu ion
and di e si ica ion
o
he mammalian o de Chi op e a. Among e eb a es, ba s
sha e wi h bi ds and possibly he ep ilian p e osau s ( he la e may ha e been
~dap ed
o gliding a he han ue ligh ) he unique abili y
o
sus ained ligh .
Whe eas he pel ic appendages
o
bi ds ha e emained ela i ely unchanged o
e es ial locomo ion and only he pec o al appendages modi ied o ligh , ba s
ha e become o ally commi ed,
in
an
ana omical sense, o as a egy o ligh .
The di e ence in he mode
o
adap a ion o ligh by bi ds and ba s no doub e-
lec s wo qui e di e en selec i e egimes in ol ing bipedal and quad upedal
ances y, espec i ely. Ba s appa en ly became adap ed o an ae ial exis ence
in
o de o exploi
an
ae ial insec i o ous ood sou ce.
On
he o he hand, bi ds
ini ially may ha e de eloped ligh o pu sue ap edacious mode
o
li e, o escape
p eda o s, o dispe sal,
o
acombina ion o hese (Os om, 1974). Admi edly,
ae ial insec i o y has been impo an in he adap a ion and di e si ica ion
o
bi ds, bu his pa icula eeding s a egy has no been he cen al ocus
in
hei
specia ion.
Al hough ba s a e a ema kably success ul g oup and comp ise he second
la ges mammalian o de , hey emain one
o
he leas known g oups
in
e ms
o
a ossil eco d. The delicacy
o
he chi op e an skele on and he ca e and o es -
dwelling habi
o
ba s ha e appa en ly con ibu ed o he pauci y
o
ossils. The
an iqui y
o
he Chi op e a
is
con i med
by
Ica onyc e is om he ea ly Eocene
o
Wyoming and F ance (Russell e al., 1973; includes desc ip ion and compa i-
sons
o
Ica onyc e is? menui); Palaeochi op e yx and A chaeonyc e is om he
ea ly Eocene
o
Aus ia; Cecilionyc e is om he middle Eocene
o
Ge many;
Ageina om he ea ly Eocene
o
F ance; and he ex an genus Hipposide os om
he middle Eocene
o
Eu ope. By he Oligocene and Miocene, six chi op e an
amilies (P e opodidae, Rhinolophidae, Emballonu idae, Phyllos oma idae,
Vespe ilionidae, and Molossidae) a e ep esen ed
in
he geologic eco d. Un-
o una ely, mos
o
he ossilized emains o ba s a e ex emely agmen a y
wi h he excep ion
o
Ica onyc e is index, which
is
beau i ully p ese ed.
Ma in (1972) compiled asynopsis
o
la e Pliocene and Pleis ocene ba s (in-
cluding phyllos oma ids) om No h Ame ica and he An illes. Fo he mos
pa , his lis includes ex an species
o
ex inc species ha a e clea ly ela ed o
li ing axa. Paula Cou o (1938) epo ed nume ous Pleis ocene ossil ba s om
B azil bu hese, oo, we e ex an species
o
ela ed he e o. Two espe ilionids,
Miomyo is /o idanus and Suap enos whi ei, we e desc ibed
by
Law ence (1943)
om he ea ly Miocene
o
Flo ida, and, mo e ecen ly, Su on and Genoways
(1974) desc ibed Ancenyc e is asmusseni om la e Miocene deposi s
in
Gal-
la in Coun y, Mon ana. Galb ea h (1962) desc ibed Oligomyo is casemen i om
Middle Oligocene deposi s
in
Logan Coun y, Colo ado. In addi ion o hese os-
49

EVOLUTIONARY
IMPETUS
OF
THE
CHIROPTERAN
GRADE
Because
o
he meage ossil eco d o ba s, s uden s
o
chi op e an e olu ion
ha e been o ced o ex apola e he pas his o y
o
he o de based
on
ea u es
o
li ing species.
Fo
he mos pa , ana omical ea u es
o
he ligh mechanism
and den al mo phology ha e been u ilized in his endea o , whe eas he ecological
ole, in e ms
o
eeding s a egy and niche di e si y, mos ly has been o e -
looked. The p ima y emphasis
o
chi op e an biology has been desc ip i e and
i
is
only ecen ly ha he e has been ashi o syn hesizing his in o ma ion in
e ms
o
aunal and ecological complexi y. The p oblem
is
u he agg a a ed by
he pauci y
o
in o ma ion ela ing o wo ld ecosys ems in he la e Mesozoic and
ea ly Cenozoic. Howe e , o a i e a a easonable in e p e a ion
o
he e olu ion
o
he Phyllos oma idae
as
well
as
ha
o
he Chi op e a
as
awhole, one mus a
leas be awa e ha hei adap i e adia ion p og essed as an in eg al pa
o
de-
eloping global ecological complexi y.
Based on known ossils, he chi op e an g ade was ully es ablished in he
ea ly Eocene. Reasonable conjec u e migh p ojec he o igin
o
he g oup back
as
a as he ea ly Paleocene
o
pe haps e en in o he la e C e aceous.
A
ha
poin
in
geologic ime, he angiospe m adia ion was in i s ini ial s ages (Axel od,
sils, which clea ly a e assignable o he Chi op e a, he e a e anumbe
o
ag-
men ed insec i o e enlains ha a e sugges i e
o
achi op e an g ade, bu ha
can only be ca ego ized as "ince ae sedis" (Simpson, 1945; Russell and Sige,
1970).
The de elopmen
o
ligh
by
ba s (which has in ol ed nea ly all majo o gan
sys ems) was p ima ily conce ned wi h p o iding adeli e y sys em o he eed-
ing appa a us. Based on den al mo phology
o
ex an species
as
compa ed wi h
ha o ea ly ossils, i is gene ally assunled ha ae ial insec i o y was he ini ial
impe us o chi op e an e olu ion. Subsequen di e si ica ion has been associa ed
wi h he u he pa i ioning and specializa ion
o
his gene alized eeding s a egy
in o ca ni o y, pisci o y, oliage gleaning, ugi o y, nec a i o y, and sangui o y.
[Fo con enience, Iha e selec ed he ophic ca ego ies desc ibed by Wilson
(1973) in his discussion; I ealize ha hese, in hemsel es, ep esen gene alized
s a egies ha could be u he pa i ioned. ]
I
is
he goal
o
his chap e o conside he e olu ion
o
he Phyllos oma idae.
This amily has conlmanded he in e es
o
s uden s
o
chi op e an e olu ion
because i ep esen s one
o
he mos ,
i
no he mos , di e se amilies in e ms
o
eeding
s a egy-all
ca ego ies excep pisci o y being ep esen ed wi hin he
con ex
o
he amily. Fu he mo e, al hough anking hi d in numbe
o
species
(136, as compa ed o 285 o espe ilionids and 150 o p e opodids), he phyl-
los oma ids exceed all o he amilies
o
ba s in nunlbe
o
gene a (49, Koopman
and Jones, 1970; and Jones and Ca e , his olume). Las ly, he adap i e
adia ion
o
phyllos oma ids appa en ly has been con ined o he opical egions
o
he New Wo ld. Be o e p oceeding u he wi h adiscussion
o
he phyllos o-
ma ids, Ibelie e i
is
ele an and impo an o conside some
o
he o e all as-
pec s
o
chi op e an e olu ion.
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51
1952, 1970). Also,
by
he close
o
he C e aceous, he an hophilous insec o de s
Coleop e a, Dip e a, and qui e p obably he Lepidop e a (p incipal ood e-
sou ces
o
insec i o ous ba s) we e well es ablished
in
an e olu iona y and eco-
logical sense (Leppik, 1957, 1960; Bake and Hu d, 1968). In addi ion, eu he ian
and me a he ian mammals we e di e en ia ed and bo h we e expanding in o
nume ous e es ial niches.
I
oo h mo phology is any indica ion, mos
o
he e es ial niches, open o
mammals, we e gea ed
o
insec i o y
o
some o m
o
ca ni o y, al hough
gene alized he bi o y would ce ainly ha e been wi hin he unc ional po en ial
o
hese small e eb a es.
I
is
no di icul o isualize he chi op e an ances y
as
ha ing aken he o m
o
small a bo eal insec i o es ha nlay ha e possessed
gliding memb anes.
The a gumen o an a bo eal ances o
as
opposed o as ic ly e es ial an-
ces o seems ob ious
in
ligh
o
he ac ha
all
olan mammals no mally
launch om ees
o
heigh s abo e he g ound. The ansi ion om gliding o a
mo able wing could ha e p og essed by way
o
elonga ion .o he digi s and
in e connec ed memb anes o inc ease he su ace a ea o he pa agium. The
pa agial a angemen possessed by li ing De mop e a may esemble an ea ly
s age h ough which he chi op e an ances o passed.
I
seems easonable o
suspec ha digi al elonga ion would ha e eached apoin
o
diminishing e-
u ns in ha u he p og ession would ha e p oduced an ungainly and clunlsy
s uc u e ha necessi a ed mo emen as awing a he han use as a ixed gliding
de ice.
[I
should be no ed he e ha bi ds and he ecen ly desc ibed gian
p e osau (Lawson, 1975a, 1975b) appa en ly achie ed g ea ly elonga ed wings
by
usion and elonga ion
o
nondigi al elemen s o elonga ion o asingle digi
wi h b oad-based a icula ions, espec i ely.] Ha ing success ully a e sed
his c i ical poin in wing de elopmen , appa en ly by simply u ilizing he exis -
ing do sal and en al ho acic muscula u e o d i e he wing (Vaughan, 1970a,
1970b, 1970c), ba s we e well on hei way o occupying an ae ial insec i o ous
niche. Fu he e inemen s
o
he wing p obably ela ed o such pa ame e s
as
maneu e abili y and speed.
The a gumen s o suppo he conjec u ed insec i o y o he chi op e an an-
ces o may be ega ded
as
open o ques ion. As no ed abo e, his gene al assump-
ion
is
based
on
he mo phology
o
he den al a cade
o
known ossil ba s. The
p imi i e ibosphenic den i ion
o
eu he ian-me a he ian mammals was modi-
ied
in
he ea lies ba s o adilambdodon condi ion wi h ama ked W-shaped
ec oloph. This con igu a ion, which allows an inc ease in he numbe
o
shea ing
ace s on he pos canine den i ion,
is
gene ally associa ed wi h insec i o y (o
ca ni o y) in li ing ba s
as
well
as
in
li ing insec i o es such as sh ews and
moles. The poin o be made he e is ha whe eas an a bo eal habi seems equi-
si e o he de elopmen
o
chi op e an ligh , insec o o y,
in
and
o
i sel , does
no . Su ely, niche pa i ioning would ha e played as in eg al apa in he a ious
mammalian adap i e adia ions in he la e C e aceous
o
ea ly Paleocene
as
i has
in con empo a y ecosys ems.
Wi h his in mind we migh ask he ques ion, why do animals occupy an
a bo eal niche o begin wi h?
I
li ing o ms a e any indica ion, we migh con-
side spa ial seg ega ion
o
such pa ame e s as nes ing and oos ing si es, escape
om e es ial (nona bo eal) p eda o s, and he u iliza ion
o
such ood e-
sou ces as insec s
o
o he small o ganisms gleaned om b anches and oliage,
seeds, ui s, lowe s, and he like. All
o
hese a iables would ha e been impo -
an
o he chi op e an ances o , and, ce ainly, he u iliza ion
o
a ious ood
i ems (omni o y) would ha e been well wi hin he po en ial
o
hei ibosphenic
den i ion.
The o egoing discussion poin s ou he e olu iona y impe us o
he
chi op-
e an g ade.
I
is
ele an o
ou
conside a ion
o
he e olu ion
o
he Phyl-
los oma idae (a g oup
ha
exploi s many eeding s a egies) because i es ablish-
es he a ionale and po en ial
o
he a bo eal niche wi h ega d o chi op e an
e olu ion. Al hough insec i o y, in he o m
o
oilage gleaning o pe haps ae ial
insec i o y, may ha e been impo an o he chi op e an ances o , ce ainly op-
po unis ic ca ni o y
o
e en ugi o y (u iliza ion
o
ui s, seeds, and lowe s)
would ha e been possible.
The
la e
is
especially impo an conside ing he
deg ee
o
den al specializa ion and o he ana omical depa u es om he chi op-
e an no m seen in li ing p e opodids (appa en ly exclusi e ugi o es and he
only ba s o he han phyllos oma ids o u ilize his ood esou ce).
On
his basis
alone and wi hou ag ea deal
o
conjec u e i would be possible o pos ual e
diphyly,
o
a leas an ea ly dicho omy, o ba s wi h espec o he wo dis inc
lineages-Megachi op e a
(P e opodidae) and Mic ochi op e a (all
o he
li ing
amilies
o
ba s).
PHYLOGENETIC
RELATIONSHIPS
OF
THE
CHIROPTERA
Judgemen s as o he phylogeny and e olu ion
o
majo g oups
o
ba s ha e
been mos ly in ui i e and based on ea u es exhibi ed by li ing species. Hill
(1974), in his desc ip ion
o
he new ba amily C aseonyc e idae, wa ned
o
he
inhe en di icul ies and dange s
o
his p ac ice. Wi h espec o ba s, he
p oblem his o ically has in ol ed he assessmen
o
he deg ee
o
specializa ion
o
he ligh mechanism and he den al a cade; o he sys ems mos ce ainly could
be added in his conside a ion, bu , o da e, ew ha e been examined.
Whe he igh ly
o
w ongly,
i
we a e o p oceed wi h an in e p e a ion
o
chi op enin phylogeny based on li ing
axa
and he meage sample
o
ossil ep-
esen a i es, we mus es ablish an in e ence as o he
na u e
o
he p o o ype.
The
ecen desc ip ion
o
[ca onyc e is index om he ea ly Eocene
o
Wyoming
(Jepsen, 1966, 1970; Russell and Sige, 1970; Segall, 1971) has p o ided
chi op ologis s wi h a an alizing insigh in o he p ospec i e chi op e an p o o-
ype. [ca onyc e is along wi h Paleochi op e yx, A chaeonyc e is, Cecilionyc-
e is, and Ageina es ablishes he na u e
o
awo ld-wide paleochi op e an g ade
in ea ly o middle Eocene imes.
C anially, he Paleochi op e a ese lbled upaiids
in
gene al shape, al hough
he b aincase may no ha e been as in la ed.
The
acial po ion
o
he c anium
was long and ape ed dis ally. The p emaxilla ies
o
!ca onyc e is
appea
o ha e
been used, al hough Jepsen (1970) claimed hey we e no uni ed a he midline.
The
zygoma ic pla e was b oad and he zygoma we e well de eloped.
The e
is
no
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indica ion
o
apos o bi al
ba
on any
o
he paleochi op e an ossils.
The
den al
o mula was i2-3?/3 c
1/1
P3/3 m3/3. The inciso s we e well de eloped and
he p emola s exhibi ed a educ ion in size om pos e io o an e io . The mola s
we e ypically ibosphenic and dilambdodon , wi h a W-shaped ec oloph and
small hypocone on he
uppe
ee h. The lowe mola s had well-de eloped
alonids. All mola cusps and associa ed conlmissu es we e high and con ibu ed
o he complex
o
shea ing ace s so cha ac e is ic
o
insec i o e den i ions.
The
wing
o
paleochi op e ans was ully de eloped, bu p imi i e
in
mos
aspec s. The globula head
o
he hume us was he mos p ominen ea u e
o
he p oximal po ion
o
his bone and he g ea e ube osi y was no pa icula ly
enla ged and did no ex end p oximal
o
he head
o
he hume us. Con a y o
Jepsen (1966), Idoub ha he e was a
~'seconda y
a icula ion" es ablished be-
ween he g ea e ube osi y and be sup aglenoid egion
o
he p imi i e, ye
chi op e an, scapula (Smi h, 1972). Dis ally, he hume us was a he p imi i e,
wi h asho medial p ocess and gene alized ochlea and capi ula su aces.
The
adius appea s o ha e been ypically chi op e an, and,
ju~ging
om he dis-
al a icula ion
o
he hume us, he e we e no special locking ace s p esen
in
he
elbow egion; disjoin ing s esses de eloped du ing ligh
in
his egion we e
p obably p e en ed by muscula and ligamen ous binding. The ulna
o
Ica onyc e is was no used wi h he p oximal po ion
o
he adius and, albei
educed in size,
i
appea s o ha e been nea ly comple e, ese nbling he condi ion
ound in megachi op e ans (P e opodidae).
The
i s digi ( humb)
o
Ica onyc e is was la ge and appa en ly ee
o
he pa agium, and he second digi
e mina ed wi h aclaw, again esembling he Megachi op e a. Al hough some
phalangial elemen s a e missing om
all
a ailable ossils, he wing appa en ly
was b oad,
o
low aspec , and wi hou any special ip modi ica ions (Findley
e al., 1972). The deg ee
o
sac al usion o he pel ic egion
in
paleochi op e ans
was somewha less han ha which occu s in ei he he Megachi op e a
o
Mic o-
chi op e a.
The
head
o
he emu was la ge and globula and se be ween he
langelike g ea e and lesse ochan e s. Paleochi op e yx had awelJ-de eloped
calca , whe eas he emaining paleochi op e ans appa en ly did no , al hough
his absence may be an a i ac
o
p ese a ion.
The
ail was long and slende
in
Ica onyc e is and Paleochi op e yx. Isuspec ha
A chaeonycle is
also possessed
along ail.
The
paleochi op e an g ade, as exempli ied by he Eocene ossils, was p imi-
i e and gene alized in mos espec s.
The e
is
li le doub ha hese ba s we e
insec i o ous, bu hei capaci y o acous ic o ien a ion emains ques ionable
(Segall, 1971). Wi h u he e inemen s in he wing, o speed and maneu e -
abili y, and specializa ion
o
he cochlea egion, o acous ic o ien a ion, he
Mic ochi op e a a e easily de i ed om he paleochi op e an g ade as cha -
ac e ized abo e. Isuspec ha such di e gence occu ed in he Paleocene.
The
ques ion conce ning he ela ionship
o
he Megachi op e a and he pale-
ochi op e an g ade is no so easily esol ed. Meschinelli (1903) desc ibed
A chaeop e opus
ansiens om he ea ly Oligocene
o
I aly. Meschinelli, along
wi h Ande sen (1912), Re illiod (1922), and Dal Paiz (1937), ega ded
A ch-
aeop e opus
as
ep esen ing he oldes membe
o
he Megachi op e a. This as-
signmen
is
based p ima ily upon simila i ies
o
wing mo phology, because Re il-
Hod
(1922), Russell and Sige (1970), and Slaugh e (1970) ha e poin ed ou ha
he den i ion
o
A chaeop e opus, which
is
badly agmen ed, mo e closely e-
sembles ha
o
he Mic ochi op e a in appea ance. Isuspec ha A chaeop e o-
pus ep esen s a u he di e en ia ion
o
he paleochi op e an g ade and pe -
haps
is
no a all ela ed o he Megachi op e a. The dis inc ness and ma ked de-
pa u e
o
megachi op e an den i ion om ha
o
he Mic ochi op e a, as well
as
om known Te ia y ossils, sugges s o me ha his g oup
o
ba s had hei
o igin much ea lie in he paleochi op e an g ade
o
pe haps,
as
no ed abo e,
sepa a ely om an insec i o ous ances al s ock.
I
is
impo an o poin ou he a ionale o weigh ing den al mo phology, in
de e ence o wing mo phology, a his le el
o
in e p e a ion
o
chi op e an e o-
lu ion. Since he Oligocene, he e appea s o ha e been ela i ely li le a ia-
ion in he den al mo phology
o
he Mic ochi op e a; he g ea es depa u e
om he basic dilambdodon condi ion
is
seen in he phyllonyc e ine and des-
modon ine phyllos oma ids. The educ ion and modi ica ion
o
he den i ion in
hese wo sub amilies seems o be he p edic able consequence
o
ahighly
specialized eeding s a egy and
in
bo h cases he den al pa e n is aceable o he
"p imi i e" dilambdodon condi ion (Slaugh e , 1970).
The
deg ee and con-
sis ency
o
di e ence
o
he megachi op e an den i ion as well as he appa en
o al absence in li ing
o
ossil axa
o
any den i ion enlo ely simila o he
dilambdodon condi ion, u he sugges s a a he leng hy sepa a ion om he
paleochi op e an ances o . [Slaugh e 's (1970:77, ig.
1)
a gumen o asingle
di e gence
o
he Mic ochi op e a and Megachi op e a om apaleochi op e an
p o o ype based
on
supposed simila i y
o
he den i ions
o
A chaeop e opus--
see commen
abo e--and
Ha pyionyc e is
is
weak.]
On
he o he hand, ma ked
di e ences in wing mo phology migh no be expec ed. The e en ion
o
a"p imi-
i e" wing by megachi op e ans may simply e lec he adequacy
o
his s uc u e
o he habi
o
hese ba s; whe eas, he wing
o
mic ochi op e ans has been modi-
ied o p o ide g ea e maneu e abili y
o
speed, he eby acili a ing u he
pa i ioning
o
he insec i o ous niche. The e o e, wi h ega d o he Mic o-
chi op e a, he depa u e om he paleochi op e an p o o ype had o do mo e
wi h e ining he wing o maneu e abili y and speed and wi h less emphasis on
modi ying he den al mo phology. Among he Mic ochi op e a, he phyllos o-
ma ids illus a e he g ea es di e si y in den al modi ica ion and his appea s o
ha e aken place since he Oligocene
as
will be discussed beyond.
The Mic ochi op e a, no doub , ha e hei o igin wi hin he paleochi op e an
g ade. Twen y-one gene a (age and geog aphic loca ion in pa en hesis) occu as
Te ia y ossils and a e assignable o he ollowing six li ing amilies (Re illiod,
1922; S i on, 1931; Law ence, 1943; Simpson, 1945; Galb ea h, 1962; Russell
and Sige, 1970; SJaugh e , 1970; Smi h, 1972; Su on and Genoways, 1974):
Emballonu idae-
Vespe ilia us (Eocene-Oligocene, Eu ope);
RhinoJophidae-
Palaeophyllopho a (Eocene-Oligocene, Eu ope), Pa aphyllopho a (Eocene-
Miocene, Eu ope), Palaeonyc e is (Oligocene, Eu ope), Rhinolophus (Eocene-
Recen , Eu ope), Pseudo hinolophus (Eocene-Oligocene, Eu ope), Hipposide os
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(Eocene-Recen , Eu ope); Megade nla idae-Nec oman is (Eocene, Eu ope),
Miomegade ma (Miocene, Eu ope), P o ampy us (Eocene-Oligocene, A ica);
Phyllos oma idae-No onyc e is
(Miocene, Sou h Ame ica);
Vespe ilionidae-
S ehlinia (Eocene-Oligocene, Eu ope), Nyc e obius
(=
Re illiodia) (Eocene-
Oligocene, Eu ope), Samonyc e is (Miocene? Pliocene, wes e n Asia), Suap enos
(Miocene, No h Ame ica), Miomyo is (Miocene, No h Ame ica), Ancenyc e is
(Miocene, No h Ame ica), Oligomyo is (Oligocene, No h Ame ica), Myo is
(Oligocene-Recen , Eu ope), Simonyc e is (Pliocene, No h Ame ica); Molos-
sidae-
Tada ida
(=
Nyc inomus) (Oligocene-Recen , Eu ope). In addi ion, he e
a e se e al gene a
o
such agmen a y emains ha amilial assignmen is no
possible a his ime. F om his, i
is
e iden ha mos
o
he majo amilies
o
Mic ochi op e a we e well es ablished a leas by middle Oligocene
o
Miocene.
I seems o be gene ally ag eed upon ha he majo i y
o
he Mic ochi op e a
o igina ed in he Old Wo ld. This hesis is suppo ed by he middle Te ia y oc-
cu ence
o
he Emballonu idae, Rhinolophidae, Megade ma idae, Vespe ilioni-
dae, and Molossidae in Eu opean deposi s. Wi h he excep ion
o
he Megade -
ma idae, which appa en ly has no li ing ep esen a i e in he Palea c ic ( empe -
a e Eu asia), all
o
he abo e-lis ed amilies ha e di e en ia ed in, and p esen ly
occu in, all majo zoogeog aphic egions
o
he Old Wo ld.
I
is no ewo hy ha
he di e en ia ion
o
hese i e amilies, as well as he emaining i e in he Old
Wo ld (Rhinopoma idae, C aseonyc e idae, Nyc e idae, Myzopodidae,
and
Mys acinidae), has p oceeded along he heme
o
insec i o y o , in se e al cases,
ca ni o y
o
pisci o y ( o example, Megade ma, Mac ode ma, and, pe haps,
Ca diode ma).
O
hese 10 Old Wo ld amilies, only h ee (Emballonu idae,
Vespe ilionidae, and Molossidae) also ha e adap i ely adia ed in he New
Wo ld, he emballonu ids being con ined he e o he Neo opics.
Al hough i has no been p ecisely s a ed, an emballonu oidlike ances y gen-
e ally has been accep ed as he base o he Mic ochi op e a (Fig. 1A). Seeming-
ly, his hypo hesis
is
based mo e on he appa en an iqui y
o
he g oup a he
han any pa icula se
o
p imi i e cha ac e is ics. Ce ainly, he long and slende
ee ail
o
Rhinopoma
is
eminiscen
o
acondi ion no ed in se e al paleo-
chi op e ans. Howe e , he end owa d acial sho ening by educ ion in size
and numbe
o
p emola s, os al in la ion, un used and unique o m
o
he p e-
max
ill
a ies, end owa d complica ed osseous p ocesses on and pos e io o he
do sonasal pla e, complica ed basisphenoidal pi s, and specializa ions
o
he
hume us (bo h p oximally and dis ally) sugges asomewha mo e specialized
s a e o hese ba s han would be expec ed o an ances al g oup. A u he
co olla y
o
he hypo hesis
is
ha he New Wo ld noc ilionids, mo moopids, and
phyllos oma ids e ol ed om an
Old
Wo ld emballonu oid mig an . The Neo-
opically endemic na alids, hy op e ids, and u ip e ids appa en ly we e de-
i ed somewha la e om a espe ilionoid s ock, which in u n had e ol ed
om he emballonu oid complex.
Un il he disco e y
o
lca onyc e is index (Jepsen, 1966), which e i ied he
exis ence
o
awo ld-wide paleochi op e an g ade, he o egoing hypo hesis
would no ha e been o ally un enable. Wi h !ca onyc e is, mo e ligh
is
shed,
albei dim, on he ques ion
o
ea ly chi op e an phylogeny, and i now becomes
B
P e opodidae
FIG.
J
.-A,
cladog am
o
he
gene ally accep ed iew
o
chi op e an
phylogeny wi h he
M
ic ochi op e a
de i ed om a
common
emballonu oidlike
ances y.
B,
an
al e na i e p o-
posal o
chi op e an
e olu ion
wi h
se e al
mic ochi op e an
lineages being de i ed, in-
dependen ly, om awo ld-wide
paleochi op e an
g ade
and
he
Megachi op e a
(P e opod-
idae) de i ed
ei he
sepa a ely
om an insec i o ous s ock
o
ea ly
om
he
paleochi op-
e an
g ade.
a,
Emballonu oidea;
b,
Rhinolophoidea;
c,
Phyllos oma oidea;
d, Vespe ilionoi-
dea.
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Rhinopoma idae
}a
C aseonyc e idae
Emballonu idae
Megade ma idae
}b
Nyc e idae
Rhinolophidae
Phyllos oma idae
}c
Mo moopidae
Noc ilionidae
Molossidae
Mys acinidae
Na alidae
Thy op e idae d
Fu ip e idae
Vespe ilionidae
Myzapodidae
A
P e opod
idae
Rhinopoma idae
}a
C aseonyc e idae
Emballonu idae
~b
Megade ma idae
}b
~
Nyc e idae
~o
~
:" . Rhinolophidae
~
':.--.;;
~
Phyllos oma idae
}c
§Mo moopidae
<a
Noc ilionidae
0
~
Molossidae
Mys acinidae
1
Na alidae
'-......1
Thy op e idae d
Fu ip e idae
Vespe ilionidae
Myzapodidae
56
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
57
possible o sugges an al e na i e o an exclusi ely emballonu oid ances y
o
he Mic ochi op e a, a leas
in
he New Wo ld. This al e na i e hypo hesis would
be o ecognize apaleochi op e an g ade as asou ce om which se e al lineages
o
Mic ochi op e a adap i ely adia ed (Fig. IB).
In he conside a ion
o
he phylogeny
o
he New Wo ld ba s, especially
noc ilionids, mo moopids, and phyllos oma ids, he abo e p oposi ion
is
o
pa a-
moun impo ance. The disco e y
o
No onyc e is magdalenesis (Sa age, 1951,
om he la e Miocene
o
Colombia) indica es ha phyllos oma ids we e
weJ]
es ablished
in
an e olu iona y sense
in
he la e Te ia y. The oldes embal-
lonu oids om he Que cy auna (Eocene-Oligocene
o
Eu ope) we e no ma k-
edly dissimila om mode n species o ha amily. The amoun
o
ime in ol ed
o accoun o he magni ude
o
ana omical di e en ia ion
o
he phyllos oma ids
om
an
emballonu oidlike ances o , amig an om he Old Wo ld, seenlS o
shi unduly he whole e olu iona y sequence
o
he Mic ochi op e a o amuch
ea lie and as ye undocumen ed age.
I
is
equally plausible o sugges asepa a e
and independen adia ion
o
hese h ee unique New Wo ld amilies om he
paleochi op e an g ade p esen
in
he ea ly o middle Te ia y
o
he New Wo ld.
In u he suppo
o
he al e na i e hypo hesis, aconside a ion
o
he o e aH
ana omical adap a ion o apa icula eeding s a egy
is
ele an . Wi h ega d
o he Mic ochi op e a, some a ied modes
o
insec i o y (o ca ni o y) ap-
pa en ly we e he ini ial impe us o he di e en ia ion
o
he a ious amilies.
I hink i
is
impo an o no e ha i
is
only in he New Wo ld opics ha al e -
na i e mic ochi op e an eeding s a egies such as ugi o y, nec a i o y, and
sangui o y de eloped.
I
could be a gued ha ,
in
he Old Wo ld, he a ious ugi o ous, nec a i-
o ous, and o he simila niches we e al eady occupied
by
he Megachi op e a.
This,
o
cou se,
is
en i ely possible, bu
i
is
no consis en
in
an e olu iona y
sense o in oke an adap i e po en ial o an emballonu oidlike ances o
in
he
New Wo ld (whe e al e na i e eeding niches appa en ly we e a ailable) and
no o conside he same po en ial as likely
in
he Old Wo ld opics. The e o e,
i seems easonable o expec ha al e na i e eeding s a egies would ha e been
exp essed, e en
in
amino way,
in
he adap i e adia ion
o
Old Wo ld mic o-
chi op e ans. Ye , he oldes mic ochi op e an ossils om he Old Wo ld now
a ailable o in e p e a ion as well as he en i e Old Wo ld mic ochi op e an
complex, a e specialized o insec i o y
o
a
elC;l ed
eeding s a egy.
The a gumen o ecological compe i i e exclusion
o
he Mic ochi op e a
by
he Megachi op e a o ugi o ous and nec a i o ous eeding niches in he Old
Wo ld also seems weak. I beha io o li ing ep esen a i es
o
bo h g oups and
he mode
by
which ood esou ces a e pa i ioned is any indica ion
o
pas his o y,
hen, indeed, he a ailabili y
o
al e na i e eeding niches o Mic ochi op e a in
he Old Wo ld opics
is
o be expec ed. This hesis
is
p oposed on he basis ha
he nonacous icalJy o ien ing megachi op e ans, acul a i ely, u ilize he ood
esou ces ( ui s, lowe s, nec a , and pollen) du ing he wiligh (c epuscula )
pe iod, he eby lea ing hese esou ces a ailable du ing he noc u nal hou s o
acous ically o ien ing mic ochi op e ans.
Smi h (1972), in conside ing he phylogene ic ela ionships
o
he Mo moopi-
dae, sugges ed ha he Phyllos oma idae, Mo moopidae, and Noc ilionidae we e
in ima ely ela ed o he ex en ha hey migh ha e been de i ed om common
ances y. This ela ionship
is
suppo ed on he basis
o
simila c anial and pos -
c anial skele al mo phology as well
as
on simila i ies
o
he so ana omy. The
ana omical simila i ies
o
he phyllos oma ids wi h hese wo amilies is especially
p onounced
i
one conside s he sub amily Phyllos oma inae. Wi h ega d
o he h ee amilies, he phyllos oma ids a e he mos di e gen , wi h he mo -
moopids being somewha in e media e be ween noc ilionids and phyllos oma ids
in his ega d (Fig. 1). This di e gence simply may be a e lec ion
o
he di e si y
in eeding s a egies u ilized by he la e .
RELATIONSHIPS
WITHIN
THE
PHYLLOSTOMATIDAE
The adap i e adia ion
o
he Phyllos oma idae appa en ly was a esponse
o exploi he a ious ugi o ous niches in he New Wo ld opics. One sub-
amily, he Desmodon inae, de eloped he unique eeding s a egy
o
sangui o y.
In addi ion o hese specialized s a egies, some membe s
o
he amily, pa -
icula ly he phyllos oma ines, pu sue he mo e ypical chi op e an eeding
s a egy
o
insec i o y and, in se e al ins ances, ca ni o y and omni o y. The
amily as cu en ly unde s ood
is
di ided in o six (pe haps se en) sub amilies:
Phyllos oma inae, Glossophaginae, Ca olliinae, Phyllonyc e inae, S enode -
minae (he e including he S u ni inae), and Desmodon inae. This classi ica ion
is
adi ionally based mos ly on den al mo phology.
The phylogene ic ela ionships wi hin he amily a e complex and a e no well
unde s ood a his ime.
Pa
o
he con usion may be due o simila , bu un-
ela ed, adap a ions o simila eeding s a egies. The phyllos oma ines a e gen-
e ally conside ed o ep esen he mos p imi i e
o
phyllos oma id sub amilies.
The den al a cade
o
hese ba s shows he leas amoun
o
modi ica ion when
compa ed o o he sub amilies. Slaugh e (1970) sugges ed he p o o ypic den i-
ion
o
he phyllos oma ids would ha e had a o mula
o
i2/2, c1/1, P2/3, m
3/3, which
is
ound in
mo~
li ing membe s
o
he sub amily. The uppe mola s
had well-p onounced and W-shaped ec olophs, acha ac e is ic
o
insec i o ous,
pisci o ous, and ca ni o ous ba s. As
in
noc ilionids and mo moopids, he
W-
shaped ec oloph on he uppe mola s
o
phyllos oma ines ex ends a leas hal he
wid h
o
he oo h and he p o ocone-hypoconal shel
is
no pa icula ly
b oadened. In addi ion, P3 and p3 p obably we e no much educed
in
size
in
he p o o yp ic den i ion.
In compa ing No onyc e is magdalenensis wi h o he phyllos oma ines, Sa age
(1951) ecognized se e al g oups
o
gene a wi hin he sub amily. He dis-
inguished hese p ima ily
on
he con igu a ion
o
he cusps on he cheek ee h. In
one g oup, he included No onyc e is, Ch o op e us, and Vampy um, which he
ound o di e s ikingly om Phyllos omus. Wal on and Wal on (1968) and
Smi h (1972) also no ed se e a] g oups
o
gene a wi hin he phyllos oma ine
ba s based on a ious pos c anial cha ac e is ics. In hei Mac o us ype, Wal on
and Wal on (1968) included Sa age's (1951) No onyc e is-Ch o op e us-
SPECIAL
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58
BIOLOGY OF
THE
PHYLLOSTOMATIDAE 65
Chi ode ma sal ini
Dobson
VENEZUELA
(A agua):
Cue a
de
Queb ada
Honda
(SR), Lina es, 1968.
A ibeus
jamaicensis Leach
MEXICO
(Yucauin):
La a's
Ca e
(SR),
Has
Ca e
(SR),
Lol un
Ca e
(SR),
Coyok
Ca e
(SR), Spikul
Ca e
(SR), Chacaljas
Ca e
(SR),
Ha
e ai., 1953.
CUBA:
Daiqui i
Ca e
(LP),
An hony, 1919;
Camaguey
Ca e
(LP),
Koopman
and
RUibal, 1955.
HAITI:
Ca e
nea
S . Michel (SR),
Diquini
(SR), Mille , 1929.
PUERTO
RIco:
Cue a
Mon e
G ande
(LP),
An hony,
1918;
Cue a
de
Cla a
(LP),
Cue a
del
Pe o
(LP),
Choa e
and
Bi ney,
1968;
Reynolds
e
al., 1953.
VENEZUELA
(A agua):
Cue a
de
Queb ada
Honda
(SR), Lina es, 1968.
A ibeus
cine eus Mille
MEXICO
(Yuca an):
Coyok
Ca e
(SR),
Ha
e ai., 1953.
VENEZUELA
(A agua):
Cue a
de
Queb ada
Honda
(SR), Lina es, 1968.
Enchis henes ha i (Thomas)
VENEZUELA
(A agua):
Cue a
de
Queb ada
Honda
(SR), Lina es, 1968.
Sphae onyc e is
oxophyllum
Pe e s
VENEZUELA
(A agua):
Cue a
de
Queb ada
Honda
(SR), Lina es, 1968.
Phyllops alca us (G ay)
CUBA:
Daiqui i
Ca e
(LP),
An hony,
1919;
Camaguey
Ca e
(LP),
Koopman
and
Ruibal, 1955.
Phyllops hai iensis (J. A. Allen)
HAl
TI:
Ca e
nea
S . Michel (SR),
Ca e
nea
A alaye (SR),
Diquini
(SR),
Mille ,
1929;
Ca e
nea
EnCa e
(SR), Mille , 1930.
Phyllops e us
An hony
CUBA:
Daiqui i
Ca e
(LP),
An hony, 1919.
A i eus
la escens (G ay)
JAMAICA:
Dai y
Ca e
(LP),
Williams, 1952.
S enode ma
u um
an honyi
Choa e
and
Bi ney
PUERTO
RICO:
Cue a
de
Cla a
(LP),
Cue a
del
Pe o
(LP),
Choa e
and
Bi ney,
1968.
Sub amily
Desmodon inae
Desmodus
O l ndus (Wagne )
UNITED
STATES
(Texas):
Ca e
in
Te lingua
dis ic
(LP),
Cocke ell, 1930.
MEXICO
(Yuca an): Lol un
Ca e
(SR),
Ha
e al., 1953.
CUBA:
Cue a Lamas
(LP),
Koopman,
1958.
VENEZUELA
(A agua):
Cue a
de
Queb ada
Honda
(SR), Lina es, 1968.
Desmodus
o undus pun ajudensis Woloszyn
and
Mayo
CUBA:
Cen ena io
de
Lenin, Lo na del Medio,
Pun a
Judas,
NE
coas
o
Las
Villas (SR), Woloszyn
and
Mayo,
1974
( hese
au ho s
we e
unce ain
abou
as-
signing
Koopman's,
1958, specimen om
Cue a
Lamas
o his axon).

Desmodus s ocki Jones
UNITED
STATES
(Cali o nia): Po e C eek Ca e (LP), Hu chinson, 1967;
(Flo ida): Reddick (LP), Gu , 1959, and Olsen, 1960; A edondo (LP), Ma in,
1972.
MEXICO
(Nue o
Leon): San Joseci o Ca e (LP), Jones, 1958; (Mexico): Tlap-
acoya (LP), Al a ez, 1972.
Desmodus sp.
VENEZUELA
(Monagas): Cue a del Guacha o (LP), (Clay on Ray and
Oma
Lina es, pe sonal communica ion)
[Paula Cou o (1938) epo ed Schizos oma
(=
Mic onyc e is),
Lophos oma
(=
Tona ia), Vampy us
(=
Ch o op e us, Tona ia,
o
Vampy um),Phyllos oma
( = Phyllos omus), Tylos oma
(=
Mimon
c enula um), Ca ollia, Lonchoglossa
( = Anou a), Glossophaga, Chi ode ma, S u ni a, Vampy ops, A ibeus,
Desmodus, "e c.," om Pleis ocene ca e deposi s
o
B azil. Iha e no included
hese in he abo e lis ing because he did no designa e species and hei de e -
mina ion would be di icul om he gene ic lis ha he p esen ed. No locali y
in o ma ion o he han B azil was gi en. In addi ion, he e
is
a ague e e ence o
phyllos oma id gene a ci ed
by
Pe e Lund and He lu Winge om B azilian
Pleis ocene ca es.]
SPECIAL
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LITERATURE
CITED
ALLEN,
G.
M.
1939. Ba s.
Ha a d
Uni . P ess, Camb idge, x+369 pp.
ALLEN,
H. 1898. On
he
Glossophaginae. T ans. Ame . Philos. Soc., 237-266.
ALVAREZ,
T. ]972.
Nue o
egis o
pa a
el
ampi o
del Pleis oceno.
Desmodus
s oki
[sic] de Tlapacoya, Mexico. An. Esc. Nac. Cienc. BioI., Mexico,
19:
163-165.
ANDERSEN.
K.
]912.
Ca alogue
o
he
Chi op e a
in
he
collec ion
o
he B i ish Museum.
B i ish Mus. (Na . HisL), London,
2nd
ed
.•
]:ci +1-854 +30.
ANTHONY,
H. E. ]917.
Two
new ossil ba s om
Pue o
Rico. Bull.
Ame .
Mus.
Na .
His ., 37:565-568.
1918.
The
indigenous land
mammals
o
Pue o
Rico, li ing
and
ex inc . Mem.
Ame . Mus.
Na .
His .. 2:33]-435.
1919.
Mammals
collec ed in
eas e n
Cuba
in 1917. wi h desc ip ions
o
wo
new aces. Bull. Ame . Mus.
Na .
His .,
41
:625-643.
?Phyllos oma idae ince ae sedis
Uni . Cali o nia Mus. Paleo. No. 54572, an . wo- hi ds
o
igh p2 om
UCMP
loco
V-5847
in
Big Ca Qua y, Cuyanla Valley Badlands, San a Ba -
ba a Co., Cali o nia.
Age.--ea ly
CI
a endonian. James (1963) no ed ha his
oo h was in he size ange
o
Phyllos omus, Ch o op e us, Vampy um, and
No onyc e is, bu admi ed ha gene ic assignmen was impossible and amilial
assignmen was conjec u al.
UCMP
No. 82144, le lowe canine;
UCMP
No. 80324, eden ulous and incom-
ple e igh den a y, om
UCMP
loco
V-6761 B anch Canyon Fo ma ion, San a
Ba ba a Co., Cali o nia.
Age.-Heming o dian.
Hu chison and Lindsay
(1974) no ed ha hese specimens esembled P e ono us ( amily Mo moopi-
dae), Loncho hina, and Mac ophyllum, bu de e ed gene ic assignmen .
66
AXELROD,
D.
I.
1952. A heo y
o
angiospe m e olu ion. E olu ion, 6:29-60.
1970. Mesozoic paleogeog aphy and ea ly angiospe m his o y. Bo . Re .,
36:277-319.
BAKER,
R. J. 1967. Ka yo ypes
o
ba s
o
he amily phy]]os oma idae
and
hei
axonomic
implica ions. Sou hwes e n Na ., 12:407-428.
1970.
The
ole
o
ka yo ypes in phylogene ic s udies
o
ba s. Pp. 303-312, in
Abou
ba s (B. H. Slaugh e and D. W. Wal on, eds.),
Sou he n
Me hodis Uni .
P ess, Dallas, ii +339 pp.
1973.
Compa a i e
cy ogene ics
o
he
new wo ld lea -nosed ba s (Phyllos oma -
idae).
Pe iodicum
BioI., 75:37-45.
BAKER,
H. G.,
AND
P. D.
HURD,
JR.
1968. In a loTal ecology. Ann. Re . En omol.,
13:
385-413.
COCKERELL,
T.
D.
A. 1930.
An
appa en ly
ex inc Euglandina om Texas. P oc. Colo-
ado
Mus. Na . His ., 9:52-53.
CHOATE,
J. R.,
AND
E.
C.
BIRNEY.
1968. Sub- ecen Insec i o a and
Chi op e a
om
Pue o
Rico, wi h
he
desc ip ion
o
anew ba
o
he genus S enode mll. J.
Mamm.,
49:400-412.
DAL
PiAZ,
G. 1937. I.
Mammi e i
deI Oligocene
ene o. A chaeop e opus ansiens.
Mem. Ins i . Geol., Uni . Pado a,
11:
1-8.
DE
LA
TORRE,
L. 1961.
The
e olu ion, a ia ion, and sys ema ics
o
he
neo opical ba s
o
he
genus S u ni a. Unpublished Ph.D. disse a ion, Uni . Illinois.
DoBSON,
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Ca alogue
o
he
Chi op e a
in
he
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o
he
B i ish Muse-
um. B i ish Mus. (Na . His .), London. xlii+567 pp.
FINDLEY,
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1972. Mo phologic p ope ies
o
ba
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FORMAN,
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BAKER,
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GERBER.
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Commen s
on
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sys ema ic s a us
o
ampi e
ba s ( amily Desmodon idae). Sys . Zool., 17:417-425.
GALBREATH,
E. C. 1962. Anew myo id ba om
he
middle Oligocene
o
no heas e n
Colo ado. T ans. Kansas Acad. Sci., 65:448-451.
GERBER,
J. D. 1968. Elec opho e ic
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immunologic compa isons
o
he
se um
p o-
eins
o
ba s. Unpublished Ph.D. disse a ion. Uni . Kansas.
GRA
Y,
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E.
1866. Re ision
o
he
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o
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1975.
E olu iona y
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o
he
ka yo ypes
o
he
s enode mine
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GRIFFIN,
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1955. Acous ic
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HATT,
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FISHER,
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AND
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1953.
Faunal
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HILL,
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o
ba
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Bull. B i ish Mus. (Na . His .), Zoology, 27:304-336.
HUTCHINSON,
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ba
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1-16.
HUTCHINSON,
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AND
E. H.
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1974.
The
Heming o d
mammal
auna
o
he
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locali y, B anch
Canyon
Fo ma ion,
San a
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1:
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JAMES,
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I.
Geology, aunal in e p e a ions
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JEPSEN,
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Ea ly
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BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
67
1970.
Ba
o igins
and
e olu ion. Pp. 1-64, in Biology
o
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P ess,
New
Yo k.
1:
xii+
406
pp.
JONES,
J. K.,
JR.
1958. Pleis ocene
ba s
om
San
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KOOPMAN,
K. F. 1951. Fossil
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he
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1958. A ossil
ampi e
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om
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KOOPMAN,
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AND
E.
E.
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1951. Fossil
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KOOPMAN,
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AND
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1955. Ca e- ossil
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om
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KOOPMAN,
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M.
K.
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AND
E.
LEDECKy-JANECEK.
1957.
No es
on
he
mammals
o
he
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wi h special e e ences
o
he
ba s. J.
Mamm.,
38:164-174.
KOOPMAN,
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AND
J. K.
JONES,
JR.
1970.
Classi ica ion
o
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Abou
ba s
(B. H.
Slaugh e
and
D. W.
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eds.),
Sou he n
Me hodis
Uni . P ess,
Dallas, ii+339 pp.
LAWSON,
D. A. 1975a.
P e osau
om
he
la es
C e aceous
o
wes Texas: disco e y
o
he
la ges lying
c ea u e.
Science, 187:947-948.
1975h.
Could
p e osau s
ly? Science, 188:676-678.
LAWRENCE,
B.
1943.
Miocene
ba
emains
om
Flo ida,
wi h
no es
on
he
gene ic
cha ac e s
o
he
hume us
o
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24:356-369.
LEPPIK,
E.
E.
1957.
E olu iona y
ela ionship
be ween
en omophilous
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and
an ho-
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E olu ion,
11
:466-481.
1960.
Ea ly
e olu ion
o
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LINARES,
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Qui op e os
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encon ados
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enezolanas,
Pa e
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MACHADO-ALLISON,
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The
sys ema ic
posi ion
o
he
ba s
Desmodus
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Chilonyc e is,
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on
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o
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Pliocene
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Pleis ocene
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o
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Un
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Addi ional
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o
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A chaeop e yx
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The
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o
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68
SPECIAL
PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
69
SAVAGE,
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Audi o y
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o
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wi h
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o
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Mo moops)
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o
Mo moops
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1969.
Mo phological
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beha io al
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o
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he
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G. G. ]945.
The
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o
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o
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Bull.
Ame .
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B.
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E olu iona y
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o
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Sou he n
Me hodis
Uni .
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Dallas,
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D. 1972.
Sys ema ics
o
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ST
ARRETT,
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AND
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l:xii
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ed.),
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New
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l:xii
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D.
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AND
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1968.
Compa a i e
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o
he
pel ic
and
pec o al
gi dles
o
he
Phyllos oma idae
(Chi op e a;
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Res.
Cen e ,
Sou he n
Me hodis
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..
37:]-35.
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R. L., V.
J.
TIPTON,
AND
A.
KIEWLICZ.
1966.
The
s eblid
ba lies
o
Panama
(Dip e a:
Calyp e ae:
S eblidae). Pp. 405-575, in
Ec opa asi es
o
Panama
(R. L.
Wenzel
and
V.
J.
Tip on,
eds.),
Field
Mus.
Na .
His .,
Chicago,
xii+
861 pp.
WILLIAMS,
E. E. 1952.
Addi ional
no es
on
ossil
and
sub ossil
ba s
om
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J.
Mamm.,
33: 171-] 79.
WILSON,
D. E. 1973. Ba aunas: a
ophic
compa ison.
Sys .
Zool.,
22: 14-29.
WOLOSZYN,
B.
W.,
AND
N. A.
MAYO.
1974. Pos glacial
emains
o
a
ampi e
ba
(Chi op-
e a:
Desmodlls)
om
Cuba.
Ac a
Zool.
C aco iensia.
19:253-265.
COLLECTING
TECHNIQUES
MERLIN
D.
TUTTLE
Phyllos oma ids exhibi an unusual di e si y
in
oos ing and o aging beha io .
Hence, while se e al collec ing me hods (such as mis -ne ing and apping) a e
excep ionally e sa ile, e en hese all a sho
o
cap u ing all species unde
alJ
ci cums ances. Because each echnique esul s
in
selec i e cap u e
o
ce ain
species while p ac ically excluding o he s, aunal analyses should be based upon
he wides possible a ie y
.o
collec ing me hods.
By
con as , ecological and be-
ha io al s udies
o
one
o
a
ew
species should employ only hose echniques bes
adap ed o ob aining desi ed da a while, a he same ime, minimizing dis u bance
o he popula ion.
This chap e p o ides in o ma ion on means
o
loca ing phyllos oma ids and
hei oos si es, and discusses hose collec ing echniques ha ha e p o en o be
e ec i e.
Fo
asumma y
o
o he me hods no men ioned he e see G eenhall and
Pa adiso (1968:8-15).
MATERIALS
Many ma e ials o collec ing ba s may be placed in one
o
wo
ca ego ies-
hose employed a oos s and hose used along lyways
o
a places whe e ba s
o age. Howe e , o he ma e ials a e use ul
in
bo h kinds
o
si ua ions and will be
discussed i s ; equipmen used p ima ily o specialized collec ing will be deal
wi h la e .
An
elec ic headligh is essen ial o mos ypes
o
collec ing. The bes ligh
Iha e been able o ind
is
he Jus i e Headligh (ob ainable om Jus i e Manu-
ac u ing Co., 2061 NSou hpo A enue, Chicago, Illinois 60614). This ligh
has an adjus able beam and space o s o e spa e bulbs behind he headligh e-
lec o . Agood powe sou ce is he alkaline E e eady ba e y, no. 520. Acan as
ba e y holde can be ca ied on an a my pis ol bel .
Se e al kinds
o
holding cages ha e been desc ibed (G eenhall and Pa adiso,
1968:20-21), bu Iha e ound i mo e con enien o hold cap u ed ba s in bags
made
o
nylon a my mosqui o ne ing wi h ie s ings nea he op. Muslin bags
may also be used. Howe e , muslin
is
bulky and much hea ie o ca y, o s easily
in opical en i onmen s, and ba s canno be seen wi hou opening he op
o
he
bag.
Mis ne s a e he mos e sa ile de ices o collec ing ba s. They can be pu -
chased om he ollowing supplie s: Blei z Wildli e Founda ion, 5334 Holly-
wood Boule a d, Hollywood, Cali o nia 90027; Eas e n Bi d Banding Associa-
ion, Biology Depa men , Indiana Uni e si y
o
Pennsyl ania, Indiana, Penn-
syl ania 15701; No heas e n Bi d Banding Associa ion, 37 Old B ook Road,
Wes Ha o d, Connec icu 06117; and
W.
B.
Da is,
P.
O. Box 3522, B yan,
Texas 77801. They a e a ailable in wid hs
o
6, 9, 12, and 18 me e s by 1.2
o
71

HANDLING
OF
NETS
AND
TRAPS
Mis -ne ing and apping a e he wo mos e ec i e me hods known o col-
lec ing a a ie y
o
ba s. Much
o
hei success, howe e ,
is
dependen upon
knowledge
o
how and whe e o use hem. Ope a ional de ails a e deal wi h he e,
whe eas ac o s in luencing whe e and when o use ne s will be discussed la e .
SPECIAL
PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
72
2.4 me e s high, and a e cons uc ed
o
30 o 70-denie h ead in 25 o 36 milli-
me e s
o
longe mesh. Va ious colo s a e a ailable, bu black seems o be
mos e icien o nigh use.
The
mos e sa ile ne s o ca ching ba s ha e ou
shel es, a e 6
o
12
me e s wide and 2.4 me e s high, and a e cons uc ed
o
50
o
70-denie h ead wi h 36-millime e mesh (Handley, 1968:15-16). One should
always sample one
o
a ew ne s om agi en supplie be o e o de ing mo e.
I
is
wise o check each one o he ollowing possible de ec s:
1)
imp ope ly
h eaded shel s ings, which cause une en dis ibu ion
o
ne ing; 2) shel s ings
ha ha e become un ied and mus be e h eaded;
3)
shel s ings
o
unequal
leng h; 4) inadequa e amoun
o
ne ing be ween shel s ings; and 5) ne ing ha
is
no so and pliable. Also, he loops a he ends
o
each shel s ing should be
made
o
co on because nylon loops end o become un ied easily.
A ecen ly de eloped double- amed ap has been used unde awide ange
o
condi ions and has p o en success ul in cap u ing many empe a e and opical
ba s (Tu le, 1974a).
The
ba s collide wi h ine e ical wi es and all unha med
in o ala ge can as ecep acle om which hey a e unable o escape. This ap
is
easily ca ied by one man, can be assembled
o
b oken down in 45 minu es, and
is
pa icula ly use ul
in
s udies ha equi e apid handling
o
la ge samples. A
p esen hese aps a e no p oduced comme cially, bu acomple e desc ip ion
wi h speci ica ions o cons uc ion has been published (Tu le, 1974c). Se e al
ea lie and less e sa ile aps also ha e been desc ibed (Cons an ine, 1958,
1962, 1969).
O he
impo an i ems a e amache e and glo es. In opical a eas, amache e
wi h a
15
o 18-inch blade (and abel shea h)
is
an e icien ool o clea ing
ne ing and apping si es, o p epa ing poles o ne s, and o chopping in o
oos s in small holes. Apai
o
lea he glo es should be used when ca ching oos -
ing ba s by hand and o handling cap u ed specimens.
Mis
Ne s
P epa a o y o se ing amis ne , app op ia e poles mus be ob ained.
In
opi-
cal ain o es s, one a ely expe iences di icul y in inding adequa e saplings
ha can be cu in o leng hs
o
wo and ahal o h ee me e s. These should be
s aigh , s i , and abou i e cen ime e s in diame e a he base. All wigs should
be emo ed. Amache e may be used o sha pen he la ge end
o
each pole.
Wo k
in
ela i ely d y a eas may equi e ca ying asupply
o
poles; elescoping
aluminum poles and adjus able me al pole clamps a e con enien .
When asui able ne ing si e has been ound, amache e
is
used o emo e
su icien ege a ion and deb is so ha he ne will no become angled.
A
he
same ime, i
is
impo an no o emo e oo much ege a ion, hus lea ing he
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE 73
ne exposed and conspicuous. The goal
is
o allow ba ely enough space o he
collec o o pass eely along bo h sides wi hou endange ing he ne , bu no mo e.
A windy si es, ex a space should be allowed o billowing
o
he ne .
In mos opical a eas, poles can be d i en in o he g ound
by
epea ed jabbing
and wis ing. When se e al ha e been cu ,
i
is
bes o selec he hea ies pole wi h
he ha des , sha pes poin o make holes o he es . Depending on he igidi y
o
he poles, holes should be angled
so
ha e ec ed poles lean sligh ly ou wa d.
This allows o bending om he inwa d pull
o
he ne . When apole
is
limbe
o
inadequa ely ancho ed
in
he g ound, guy lines can be used o hold
i
in place.
In ocky i e beds,
i
may
be
necessa y o p o ide addi ional suppo
by
piling
la ge ocks agains he bases
o
he poles.
Be o e he i s pole
is
inally secu ed, he loops
o
one end
o
he ne a e
placed
in
p ope sequence o e he uppe end. When anew ne
is
i s unpacked,
he main loops, which i o e he poles, usually will be ga he ed
in
he cen e
in
wo bunches in he co ec o de .
I
is
impo an ha hese be sepa a ed ca e-
ully o a oid ini ial con usion. Nex , an ou side loop
is
ound and he es
o
he
loops on ha end a e a anged
in
app op ia e sequence o e a inge , om which
all a e slipped o e he end
o
he pole. The pole
is
i mly secu ed, and he ne
is
un olded un il
i
is
pulled igh o ma k he spo whe e he second pole is o
be
placed. A his poin , one pe son may hold he ne o he g ound while ano he
p epa es he hole.
I
alone, e old he ne be o e making he hole, un olding i
only when eady o secu e he second pole. Be o e slipping he loops o e he
pole, i
is
necessa y o check he op shel s ing o be su e ha he ne
is
no
wis ed, wi h he loops in e e se sequence. When possible, ne s should be se and
adjus ed be o e da k, bu hey should no
be
opened on he poles un il i
is
ime o
use hem.
In he e ening, when collec o s end hei ne s, hey should be equipped wi h
aheadligh , agood ba e y, spa e bulbs, holding bags, apai
o
glo es, insec
epellen , and spa e s ing o epai b oken shel s ings and o guy leaning
poles. In some a eas hey may also wish o ca y agun; poisonous snakes,
caiman, c ocodiles, and la ge ca s may be a ac ed o he squeals
o
apped ba s.
Ne s should no be le una ended o long, and mus be gua ded almos con-
s an ly when se o e ails and a ound illages whe e domes ic animals and
humans a e likely o pass. E en
in
he absence
o
o he p oblems, ba s hemsel es
will soon des oy an una ended ne . Such la ge species as Ch o op e us au i us,
Phyllos omus has a us,
o
Vampy um spec um can comple ely uin ane and
escape
in
as
li le ime
as
aminu e. Also, he longe aba s uggles
in
he ne
he mo e di icul
i
is
o emo e. As a esul , ane e should be ca e ul no o
se oo many ne s, as mo e ba s may be caugh han can be emo ed, leading o
loss
o
bo h ba s and ne s.
As soon as possible a e aba s ikes he ne , he collec o should g asp
i
wi h aglo ed hand and de e mine om which side i en e ed.
I
should be held
i mly
in
he glo ed le hand ( o a igh -handed pe son) while he unglo ed
igh hand
is
used o ex ica e he ba om he open side
o
he pocke , s a ing
wi h he head. Iusually y o emo e ne ing om he ba 's mou h i s o p e-
74
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en u he damage o he ne , and hen wo k back, eeing he wings and inally
he ee . Howe e , he e e se
is
some imes mo e con enien .
Each ne mus be eadjus ed a egula in e als so ha he bag does no mo e
along he
sh~l
s ands, lea ing igh places whe e ba s bounce o
o
bunched
places ha hey mo e easily de ec .
I
ain
o
og causes wa e d ople s o collec ,
hese should be shaken om he ne as soon as possible. Also, lea es, wigs, and
insec s should be emo ed quickly be o e hey angle ane . Badly angled s icks
can be emo ed mos easily
i
i s hey a e b oken in o small pieces; la ge bee les
should be disa icula ed.
E en when one is ca e ul, an occasional la ge ba will succeed in chewing
h ough ashel s ing and, as a esul , ala ge sec ion
o
ne ceases o unc ion.
I
possible, b oken shel s ands should be e h eaded in dayligh , bu wo expe i-
enced ne e s can accomplish he job in a ew minu es a nigh . The ne loops im-
media ely abo e and below he b oken s ing should be sp ead as a apa
as
possible and aised un il he ack
o
he b oken s ing is oughly a eye le el. A
piece
o
wo en nylon ishline
o
abou he same diame e as he o iginal s ing
and ame e long should be ied o he end
o
he longes s ing so ha his leade
can be h eaded easily wi hou ension. The shel s ing hen is ca e ully h eaded
h ough abou e e y ou h mesh along he o iginal ack. When he ends mee ,
one pe son holds he wo s ings o elease ension (o an end loop
is
emo ed
om i s pole) while he o he ies he splice
in
place (including asmall piece
o
leade ) so ha he epai ed shel s ing is he same leng h as be o e. Holes in
he mesh a e no easily epai ed, and a e enough accumula e he ne should be
disca ded
o
cu in o smalle pieces.
Ane should no be closed un il all insec s, wigs and o he deb is ha e
been emo ed, a e which all
o
he loops a e pushed oge he nea he uppe
ends
o
he poles. In a eas whe e human in e e ence is no ap oblem, ne s may
be le "closed" on he poles un il he ollowing nigh , bu when hey a e e-
mo ed i is impo an o keep he loops in o de . Iusually ollow Handley's
(1968:17) me hod
o
ying apiece
o
whi e s ing abou
40
cen ime e s long o he
op loop
o
each end. Be o e emo ing ane om i s poles, he s ing a each end
is h eaded h ough he es
o
he loops and ied. Nex he loops a e emo ed
om one pole, and he ne
is
olded
by
eaching ou abou ame e a a ime o
g asp he ne , olding i back o he i s hand again and again
as
one walks owa d
he o he pole. Finally, he loops a e emo ed om he o he pole, ied, and he
ne
is
olded and s o ed
in
asmall bag.
T aps
T aps gene ally a e no b oken down be ween se ings, bu i his
is
neces-
sa y,
as
o shipping, hey a e easse lbled
in
he ollowing manne . Each ame
is assembled sepa a ely and bol ed o he o he using ou h eaded ods. The
legs a e bol ed
in
place and he ap igh ed be o e he angled ods and wi es a e
bol ed o he op
o
each ame and ca e ully un olled.
The
h eaded ods o ad-
jus ing wi e ension a e ex ended as a as possible, and he bo on angled ods
a e bol ed o he bo om on each ame. Finally he h eaded ods a he op a e
igh ened o adjus wi e ension.
BIOLOGY OF
THE
PHYLLOSTOMATIDAE 75
P ope adjus men
o
ap ames and wi es
is
essen ial. Much depends on he
speed and angle
o
app oach
by
ba s. The ap should be e ical and pe pendicu-
la o he ligh pa h. T aps a e gene ally mos e ec i e when adjus ed
so
ha
he wo pa allel se s
o
e ical wi es a e oughly se en and ahal cen ime e s
apa , al hough his dis ance may need o be a ied o di e en condi ions and
kinds
o
ba s. Du ing ini ial es ing
o
c ude ap designs, Isucceeded
in
cap u ing
an
imp essi e numbe
o
phyllos oma ids (Tu le, 1974a). Howe e , all subse-
quen ap modi ica ions we e designed o inc ease espe ilionid cap u es, wi h-
ou conside a ion
o
phyllos oma ids, and he wi e spacing was inc eased om
wo cen ime e s o wo and ahal cen ime e s. Aspacing o wo cen ime e s
o
less, combined wi h inc eased elas ici y
o
he sp ings, migh p o e ad an a-
geous o phyllos oma ids. Rega dless
o
spacing, he au ness
o
he wi es should
be p opo ional o he speed
o
he ba s. No mally, wi es should be adjus ed
so
ha hey a e ba ely igh . When ba s escape by bouncing o he ap, wi es
should
be
loosened; when ba s pass comple ely h ough he ap, bo h ames
should be igh ened. Cap u ed ba s a e easily emo ed wi h glo ed hands, and
should be so ed in o sepa a e bags o a oid placing ca ni o ous species wi h
o he ba s. Se e al housand can be handled
in
an hou . E en hough la ge
numbe s
o
ba s do no damage aps, one mus be cons an ly igilan les ba s
apidly accumula e and su oca e be o e emo al. Whene e a ap begins o ca ch
mo e ba s han can be emo ed con enien ly, i can be u ned sideways, ca ied
ou
o
he ligh pa h,
o
co e ed on one side wi h asmall can as.
CAPTURE
TECHNIQUES
Roos s
Li le
is
known abou he ecological equi emen s
o
phyllos oma id ba s, and
he e is apauci y
o
in o ma ion a ailable on oos ing beha io . Iselec ed 28
sou ces om which in o ma ion pe aining o oos ing habi s was aken (Table
1).
Pine (] 972) was used
as
he sole sou ce
o
ma e ial on Ca ollia due o p io con-
usion in iden i ica ion. Walke (1964) is ci ed only when o iginal obse a ions
could no be ound. Sou ces a e numbe ed (see pa en he ical numbe s in Li e a-
u e Ci ed), and numbe s o e e ences ci ed appea in he app op ia e places
in Table
1.
Species o which I
was
unable o ind in o ma ion on oos ing habi s
a e no included.
Ea ly li e a u e emphasized disco e y
o
new species and seldom men ioned
how
o
whe e ba s we e collec ed. Recen ly, he use
o
mis ne s has enhanced
knowledge
o
o e all dis ibu ion and p o ided much ecological da a. Ne e he-
less, ne ing has been so con enien ha
ew
esea che s ha e been o ced o look
o oos ing ba s. Sea ches o oos s ha e been limi ed o a
ew
ob ious ypes o
places.
As
a esul , oos s in ca es, houses, hollow ees,
o
cul e s a e o en
epo ed whe eas hose in oliage and o he less e iden places a e no , lea ing he
oos ing habi s
o
e en some common species unknown. Wi h his bias in mind,
Iwill p o ide sugges ions o inding he ypes o oos ing si es ha ha e
p o en mos p oduc i e.
Ca es.-Ca es
may p o ide oos ing si es o mo e di e en species
o
phyl-
los oma ids han any o he kind
o
shel e . Mos ca es a e loca ed in limes one,
How o Collec
a
Roos s
In addi ion o he mo e gene al equipmen al eady discussed, abee smoke ,
hand ne , and .22-calibe pis ol a e essen ial o collec ing a many oos s.
In
he
discussion ha ollows, i
is
assumed ha a leas wo pe sons will be wo king
oge he . In all collec ing a oos s, i
is
i al o a oid ala ming ba s wi h un-
necessa y noise, ib a ion,
o
ligh .
Hand
ne ing.-A
hand ne ideally should ha e as u dy hoop abou
40
cen i-
me e s in diame e a ached o a1.2-me e aluminum handle.
I
needed, addi-
ional sec ions
o
elescoping aluminunl can be pu chased o ex ending he
handle o i e me e s. The bag should be made
o
nylon a my mosqui o ne ing,
a leas
75
cen ime e s deep, ounded a he bo om, and sewn a he op o
hea y clo h i ing o e he hoop. Apiece
o
hea y plas ic
18
cen ime e s wide
should hang eely a ound he inside
o
he hoop, p e en ing climbing ba s om
escaping.
Hand ne s a e mos equen ly used a oos s
in
hollow ees, animal bu ows,
ock c e ices,
o
ca es. A ahollow ee, ca e ul inspec ion should be made o
de e mine he numbe and size
o
openings onl which ba s could escape. Each
po en ial exi hen should be co e ed wi h ane
o
somehow blocked. Many ba s
can be igh ened in o a emp ing o lea e
by
pounding on he ee unk wi h a
ock. I ha ails, alimbe s ick
o
app op ia e leng h may
be
Gu ,
lea ing small
b anches and oliage in ac a one end. This can be ca e ully inse ed and wi led
nea he ba s. I ba s emain s ubbo n in hei e usal o come ou , abee smoke
can
be
used. Aleng h
o
lexible ubing may be a ached, i needed, o di ec he
smoke o aspeci ic place. Eme ging ba s a e caugh
in
he hand ne s, om which
hey a e ans e ed o holding bags.
Hand-ne ing in ca es
is
much mo e di icul . Especially nea he en ance,
each dep ession
o
c e ice should be app oached cau iously, wi h he headligh
no aimed beyond he each
o
he ne . F equen ly,
an
ex a sec ion
o
handle
is
equi ed so ha ba s can be eached quickly be o e hey become ala med. When
acolony
is
hea d, he oos ing ba s should be app oached by sound a he han by
sigh , wi h he headligh aimed a he loo jus ahead. Speed
is
c ucial inasmuch
as
many ba s will
ly
as soon as possible a e sigh ing an app oaching ligh . Mos
collec o s ind i easie o wai un il e ening when eme ging ba s can be apped
o
ne ed a he ca e en ance.
Shoo
ing.-Ba s
can be sho a oos s wi h a.22-calibe pis ol and long i le
dus sho . Al hough such pis ols equen ly a e bo ed smoo h o imp o e he
sho pa e n, Iha e ne e ound ha o be necessa y. The accep able collec ing
ange
is
oughly ou o nine me e s.
Fo
g ea e ange, one can use a.32
o
.410-calibe auxilIa y ba el and dus sho in a16-gauge sho gun.
The
mos e-
quen p oblem
is
ha
o
shoo ing a oo sho a ange, hus damaging specimens.
Shoo ing
is
bes employed when collec ing a si es ha a e easy o see om a
dis ance and di icul o app oach wi hou ala ming ba s. Examples
o
such
places a e oliage oos s, ca i ies in cli aces, o e hanging oo s, la ge ca es,
cul e s, and b idges. Apis ol also can be used in la ge hollow ees and
in
small ca es, bu he e
is
dange
o
damaging he ea s
o
he collec o . Whene e
82
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83
o he op ions a e a ailable, shoo ing should be a oided, as some specimens
may be damaged while o he s, which escape, may be needlessly inju ed.
Ne ing.-Ne s
can be e ec i e
in
cap u ing ba s eme ging om oos s no
easily co e ed
by
ahand ne , and a e especially inlpo an a b idges whe e small
colonies may be di icul o app oach. A acul e , o example, one pe son
ends he ne , which
is
se o block one end, while ano he igh ens he ba s
om he opposi e end. Damaged ne s can be cu o make one o mo e small ne s,
wo o h ee me e s long, which a e handy a cul e s. When no in use, sho
ne s a e easily olled on o an aluminum pole.
Mis ne s also may be employed in on
o
such oos ing si es as buildings
and ca es du ing e ening eme gence. F equen ly, ba s occupying hese places
canno be eached
o
o ced o exi be o e hei na u al depa u e. When colonies
a e small, mis ne s can be qui e con enien ; when la ge numbe s a e in ol ed,
howe e , ne s o en en angle hund eds
o
ba s a a ime and a e uined long be o e
ba s can be ex ica ed.
T apping.-
T aps a e pa icula ly use ul a en ances o ca es con aining
la ge colonies. They may be se anywhe e along he ligh pa h
o
eme ging
ba s, bu he bes place is o en some dis ance om a oos
en ance- o
ex-
ample, whe e ba s no mally en e oliage ha can assis
in
obscu ing he ap
om de ec ion. P io obse a ion
o
eme gence pa e ns pe mi s op imal ap
placemen . F equen ly, howe e , aps simply can be se di ec ly in on
o
a
poin
o
eme gence
o
in aca e en ance; he a ea a ound he ap may be pa -
ially blocked wi h b ush o ne ing.
Fo aging Si es
A leas a
ew
o aging ba s can be ound almos anywhe e a nigh
in
he
opics; howe e , some places a e a mo e p oduc i e han o he s. The ew
examples p esen ed he e may be in ui i ely ob ious; many addi ional possibili-
ies become appa en only wi h expe ience. O en he bes places a e disco e ed
only by ca e ul obse a ion a wiligh
o
a nigh while sea ching wi h ahead-
ligh . Sea ches a nigh should include equen pauses wi h he ligh u ned o ,
lis ening o he sounds
o
alling ui , lying ba s, and he squabbling ha occu s
a majo eeding si es.
T ails.-Mos
o es ails a leas ame e in wid h a e likely o be used by
ba s, pa icula ly when he su ounding ege a ion
is
bo h all and dense. In
he opics, he bes ails a e hose ha lead om illages o ga dens
o
plan a-
ions. While hese ails a e especially p oduc i e places o collec ing ugi o ous
species, o he s connec ing pas u es
o
leading o li es ock sheds a e mo e likely o
be used
by
ampi es. The wides ails, especially sho sec ions be ween clea -
ings, a e be e o o aging insec i o es.
Fo es
edges.-Ba s
o age and
ly
along he edges
o
mos o es ed a eas, bu
edges
o
small clea ings wi hin o es s a e bes o collec ing phyllos oma ids,
excep when he e a e eeding
o
wa e ing places in la ge open a eas nea by.
S eams.--S eams p o ide na u al lyways, especially whe e su ounding
o es
is
dense. Slow- lowing s eams, h ee o 10 me e s
in
wid h, seem o ha e
How o
CollecT
a Fo aging Si es
Shoo ing.-
Though shoo ing a dusk
o
la e wi h asho gun and numbe 12
sho is an excellen me hod o collec ing many emballonu ids, noc ilionids,
espe ilionids, and nlolossids, his me hod seldom wo ks well o ob aining
he mos a ic. Swi moun ain s eams and la ge i e s equen ly a e less p o-
duc i e
o
phyllos oma ids and, in he la e case, pose di icul ies in collec ing.
Ponds.-Isola ed
ponds in a eas whe e he e a e no o he sou ces
o
a ailable
wa e o en a ac ba s
in
spec acula numbe s and a ie y in he d y season.
O he wa e ing places, some imes only a ew cen ime e s
in
diame e , appea o
be highly a ac i e o some s enode mines, e en
in
he ainy season and
in
places whe e o he wa e
is
abundan (Tu le, 1974b). These si es a e o en well
known o local na i e hun e s who isi hem in o de o hun api s (Tapi us
e es is), which also a e a ac ed in unusual numbe s.
Feeding si e
s.-Many
phyllos oma ids a e bes collec ed
in
p oximi y o hei
eeding places. Glossophagines isi many lowe ing ees and sh ubs, he bes
o
which may be ound by wa ching hummingbi ds; ce ain plan s ha a ac hese
bi ds du ing he day a e equally a ac i e o ba s a nigh . Flowe ing banana and
cashew ees a e well wo h checking.
Ba s
o
he sub amilies Ca olliinae and S enode minae a e mos commonly
collec ed nea ui ing ees
o
sh ubs. Wild igs a ac a a ie y
o
hese ba s in
la ge numbe s, and ga dens con aining ui ing bananas, gua as, papayas,
o
mangos also a e excellen a ac ions. Especially in i gin o es , ui -ea ing
bi ds and monkeys o en p o ide clues o addi ional ood sou ces.
Vampi es equen ly a e nume ous a ound he bo de s
o
illages when
chickens, dogs,
o
pigs a e p esen . Desmodus
is
encoun e ed mos equen ly
nea ca le o ho ses, whe eas Diaemus and Diphylla a e mo e likely o be ound
nea poul y.
The
p esence
o
ampi es
is
easily con i med
by
he p esence
o
d ied blood on he head and shoulde egions
o
li es ock,
o
on pe ches whe e
poul y oos a nigh .
Feeding a eas
o
phyllos oma ines seem o be mo e gene alized and unp e-
dic able, bu Iha e nea ly always succeeded
in
collec ing Phyllos omus discolo ,
P.
elonga us, and
P.
has a us on small banana plan a ions in o es ed a eas.
Goodwin and G eenhall (1961 :240) epo ed
P.
has a us lying in g oups
o
up o
100 o eed on he seeds
o
spacaia nu ees (Lecy his zabucajo),and ha
Mic onyc e is megalo is was a ac ed o ui ing gua a ees (op. ci ., 228). I
ha e nl0s equen ly collec ed o he phyllos oma ines such as Ch o op e us,
Mimon, Phyllode ma, Tona ia, and T achops in na u al clea ings benea h adense
canopy
o
i gin lowland o es , whe e hey appea ed o be o aging. Such a eas
a e ound whe e unde g ow h has been elimina ed
by
we -season looding.
Highland
passes.-A
su p ising numbe and a ie y
o
phyllos oma ids can
be collec ed as hey
ly
h ough low places along idges. These a e mos easily
ound along oads ha ollow idges. A ele a ions be ween 1400 and 2800
me e s, Iha e commonly collec ed such in e es ing gene a as Chi ode ma,
Enchis henes, and Vampy ops.
84
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85
phyllos oma ids. Fo shoo ing o
be
p ac ical, ba s mus
ly
high enough o
be
seen agains he ho izon while he e
is
s ilJ adequa e ligh , and he e should
be
ela i ely ba e g ound o wa e below so ha downed specimens can be ound.
The o aging habi s
o
only a
ew
phyllos oma ids i hese equi emen s. Some
o
he la ge species, such as Phyllos omus has a us, can be sho a dusk om
asmall boa
as
hey a emp o c oss i e s en ou e o hei eeding si es. They
loa and can be scooped om he i e
in
ahand ne . La ge ugi o ous species
some imes can
be
spo ed wi h aheadligh and sho while eeding.
Ne ing
and
apping.-Many
phyllos oma ids ha e been collec ed e icien ly
only in mis ne s and aps. Whe eas ne s ha e been used almos exclusi ely since
he la e 1950's, he po en ial
o
aps has ecei ed widesp ead a en ion only
ecen ly. Ei he ne s
o
aps can
be
se a almos any place whe e ba s a e ex-
pec ed o ly, al hough hey a e no equally p ac ical unde
alJ
ci cums ances.
T aps a e especially con enien whene e la ge numbe s
o
ba s mus be handled
apidly. They a e no easily damaged
by
ba s
o
o he animals and do no equi e
equen a en ion unless excep ionally la ge numbe s a e being caugh . Ne s
can be aised in o he o es canopy, bu he p ocedu e
is
di icul and cos ly
(Humph ey e aI., 1968). T aps, howe e , can
be
se easily
in
dense oliage on
he g ound
o
hois ed in o he canopy wi hou dange
o
becoming angled. They
a e una ec ed
by
wind, whe eas ane mus be se exac ly pe pendicula o e en
aligh b eeze
o
he ne ing quickly blows o one end, making he ne i ually
inope able. The main disad an ages
o
aps a e ha hey a e much hea ie han
ne s, and co e asmalle a ea.
Ei he ne s
o
aps may be se a any
o
he p e iously discussed kinds
o
places, bu much
o
he success
in
using hese de ices depends on he collec o 's
abili y o camou lage hem. Many eeding si es in ol e lowe ing
o
ui ing ees
whe e ne s o aps a e se
as
close
in
on
o
a ee
as
possible, o immedia ely
benea h he lowes b anches. Some imes, howe e , nea by openings
o
ails used
by
app oaching ba s p o ide easie collec ing si es. Along ails and s eams, ne s
and aps should be se in he na owes places, p e e ably whe e he e a e
na u al obs uc ions, such
as
allen ees, ha block
all
bu asmall space. T aps
a e pa icula ly e ec i e a such si es. Hanging ines, o e hanging limbs, and
sha p u ns p o ide addi ional concealmen . A ponds and small clea ings, whe e
la ge a eas mus
be
co e ed, ne s a e mo e easily used and should be se
a ound he edges pa allel o he ege a ion. A ound na i e ga dens and o he
simila si es, I equen ly ha e s ung
as
many
as
10 12-me e ne s end o end,
al e na ing he loops om wo ne s on each pole, bu such an a ay
o
ne s mus
be manned by se e al people.
I
aps a e o be used a hese places, hey mus be
se ei he whe e ege a ion o ces na u al unneling
o
he ba s o whe e a i icial
blocking a he sides can achie e he same end. O en he sides can be blocked
by
ying as ong line o he op
o
he ap on each side, unning he lines o
nea by ees. Lea y ege a ion is hen cu and hung om he lines. This
is
es-
pecially e ec i e a he app oaches o ponds
o
whe e aps a e se o e s eams.
A low passes along moun ain idges, ne s a e p e e able, and se e al may be
se end- o-end jus below he c es whe e hey blend wi h he s eep hillside.
ACKNOWLEDGMENTS
Iwish o hank D . Robe
S.
Ho mann and Ms. Diane E. S e enson o c i i-
cally eading he manusc ip and o he wise assis ing in i s p epa a ion. D .
Cha les O. Handley, J ., gene ously p o ided assis ance du ing my ea ly col-
lec ing expe ience, which was in
pa
suppo ed
by
U.S. A my con ac DA-49-
193-MD-2788.
LITERATURE
CITED
ALLEN,
G.
M.
1939. Ba s.
Ha a d
Uni . P ess,
Camb idge,
Massachuse s, x+368 pp.
ALLEN,
J.
A. 1911.
Mammals
om
Venezuela
collec ed by M . M. A.
Ca ike ,
J .,
1909-
]
9]
I. Bull.
Ame .
Mus.
Na .
His ., 30:239-273.
(l)
ANTHONY,
H.
F.
19]8.
The
indigenous
land
mammals
o
Po o
Rico, li ing
and
ex inc .
Mem.
Ame .
Mus.
Na .
His ., n.s., 2:333-435. (2)
BOND,
R.
M.,
AND
G. A.
SEAMAN.
]958.
No es
on
a
colony
o
B achyphylla
ca e na ul11.
J.
Mamm.,
39: 150-151. (3)
BURT,
W. H.,
AND
R. A.
STiRTON.
196
J.
The
mammals
o
EI
Sal ado . Misc. PubL Mus.
Zoo)., Uni . Michigan.
I]
7:
1-69. (4)
CONSTANTINE,
D.
G.
1958.
An
au oma ic
ba -collec ing de ice.
J.
Wildli e Mg .,
22: 17-22.
1961. Locali y
eco ds
and
no es
on
wes e n ba s.
J.
Mamm.,
42:404-
405. (30)
SPECIAL
PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
E ec i eness a all places will be inc eased by equen changes
o
posi ion o
coun e lea ning beha io
o
ba s.
When choices a e a ailable, ne s and aps should be se
in
he da kes places
and a imes when he e
is
li le
o
no
n100n.
Usually, he i s and las wo hou s
o
he nigh a e mos p oduc i e, bu a ew species a e mo e likely o be caugh
a o he imes.
Techniques o lu ing
ba s.-
The e a e wo me hods ha ha e conside able
po en ial o a ac ing ba s o collec ing si es. Many ba s can be a ac ed o ne s
o
aps when o he ba s a e induced o call in dis ess. In gene al he smalles
species make he bes "calle s." O en, he dis ess c ies
o
asmall ba will a ac
la ge species in addi ion o o he s
o
i s own size, whe eas ala ge species, such
as
Phylloslomus has a us, calls i s own kind while igh ening mos o he s away.
On se e al occasions Iha e achie ed excellen esul s by hanging abag con ain-
ing 20 o 30 qua eling ba s
o
se e al species
on
he side
o
aba ap
in
aplace
whe e no ba s had been caugh p e iously. Ip edic ha aped eco dings may
someday p o e in aluable o lu ing ba s in o ne s and aps.
The less es ed
o
he wo echniques
is
he use
o
bai . A one locali y
in
Pe u,
Ca ollia was so pe sis en in sea ching ou
my
a aps bai ed wi h banana
ha e en hose se benea h dense ege a ion
o
allen logs caugh hem. Al hough
his was unusual, Ca ollia o en was a ac ed o ipening s alks
o
bananas
in
na i e hu s. Ialso ha e aken Rhinophylla pumilio
in
abanana-bai ed a ap.
F ui , caged animals,
o
e en caged insec s could be hung behind aps se
in
places whe e ba s would be o ced o app oach om he opposi e side.
I
is
qui e
possible ha some glossophagines could be a ac ed o hummingbi d eede s.
Many phyllos oma ids p obably could be lu ed in o bai ed ba aps.
86
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
87
1962.
Me hodos
de
lucha
con a
los
ampi os
ansmiso es
de
la
abia.
BoI.
O ic.
Sani a ia
Paname icana,
53:7-12.
1969.
T ampa
po a il
pa a
ampi os
usada
en
p og amas
de
campana
an i-
abica.
BoI. O ic.
Sani a ia
Paname icana,
67:39-42.
DAVIS,
R.,
AND
E. L.
COCKRUM.
1963.
B idges
u ilized
as
day- oos s
by ba s.
J.
Mamm.,
44:428-430.
DAVIS,
W. B.,
AND
R.
J.
RUSSELL.
1954.
Mammals
o
he
Mexican
s a e
o
Mo elos.
J.
Mamm.,
35:63-80. (5)
DAVIS,
W. B., D. C.
CARTER,
AND
R.
H.
PINE.
1964.
No ewo hy
eco ds
o
Mexican
and
Cen al
Ame ican
ba s.
J.
Mamm.,
45:375-387. (6)
DAVIS,
W. B.,
AND
D.
C.
CARTER.
1964. A
new
species
o
ui -ea ing
ba
(genus A iheus)
om
Cen al
Ame ica.
P oc. Biol. Soc.
Washing on,
77: 119-122. (7)
GOODWIN,
G.
G.,
AND
A.
M.
GREENHALL.
1961. A e iew
o
he
ba s
o
T inidad
and
Tobago.
Desc ip ions,
abies
in ec ion,
and
ecology. Bull.
Ame .
Mus.
Na .
His ., 122:187-302 (8)
GOODWIN,
R. E. 1970.
The
ecology
o
Jamaican
ba s. J.
Mamm.,
51:571-579. (9)
GREENHALL,
A. M.,
AND
J.
L.
PARADISO.
1968. Ba s
and
ba
banding.
Resou ce
PubI.
Bu .
Spo
Fishe ies
and
Wildli e, 72:i +1-48.
HALL,
E.
R.,
AND
W. W.
DALQUEST.
1963.
The
mammals
o
Ve ac uz.
Uni .
Kansas
Publ., Mus.
Na .
His ., 14:165-362. (10)
HALL,
E.
R.,
AND
W.
B.
JACKSON.
1953.
Se en een
species
o
ba s
eco ded
om
Ba o
Colo ado
Island,
Panama
Canal
Zone.
Uni .
Kansas
PubI., Mus.
Na .
His .,
5:641-646.
(11)
HANDLEY,
C.
0.,
JR. 1966.
Checklis
o
he
mammals
o
Panama.
Pp. 753-795, in
Ec opa asi es
o
Panama
(R. L. Wenzel
and
V. J.
Tip on,
eds.),
Field
Mus.
Na .
His .,
Chicago,
xii +861 pp. (12)
1968.
Cap u ing
ba s
wi h
mis ne s. Pp. 15-19, in Ba s
and
ba
band
ing, Re-
sou ce
Publ., Bu .
Spo
Fishe ies
and
Wildli e, 72:i +1-48.
HUSSON,
A. M. 1954.
On
Vampy odes
ca acciolae
(Thomas)
and
some
o he
ba s
om
he
Island
o
Tobago
(B i ish
Wes
Indies). ZooI.
Medad.
Mus.
Leiden,
33:63-
67. (13)
1962.
The
ba s
o
Su iname.
Zool.
Ve hand.
Rijksmus.
Na .
His .
Leiden,
58:1-282,
30
pIs. (14)
INGLES,
L.
G.
1953.
Obse a ions
on
Ba o
Colo ado
Island
mammals.
J.
Mamm.,
34:266-268. (15)
JONES,
J.
K
.•
JR. 1966. Ba s
om
Gua emala.
Uni .
Kansas
PubI., Mus.
Na .
His ..
16:439-472. (16)
JONES,
J.
K
.•
JR.,
AND
A.
SCHWARTZ.
1967.
Synopsis
o
ba s
o
he
An illean
genus
A dops.
P oc. U.S.
Na .
Mus., 124(3634):1-13.
(l7)
JONES.
J. K
.•
JR.,
J.
R.
CHOATE,
AND
A.
CADENA.
1972.
Mammals
om
he
Mexican
S a e
o
Sinaloa.
II.
Chi op e a.
Occas.
Pape s,
Mus.
Na .
His ., Uni . Kansas, 6:
1-
29. (18)
MITCHELL,
H. A. 1963.
Ammonia
ole ance
o
he
Cali o nia
lea -nosed ba .
J.
Mamm.,
44:543-551. (31)
PHILLIPS,
C.
J.,
AND
J.
K.
JONES,
JR. 1971. A
new
subspecies
o
he
long-nosed
ba ,
Hylonyc e is
unde woodi,
om
Mexico. J.
Mamm.,
52:77-80. (19)
PINE,
R.
H. 1972.
The
ba s
o
he
genus
Ca ollia.
Tech.
Monog .,
Ag . Exp. S a.,
Texas
A&M
Uni .,
8:1-125. (20)
SANBORN,
C.
C. 1936.
Reco ds
and
measu emen s
o
neo opical
ba s.
Publ.
Field
Mus.
Na .
His ., Zool. Se ., 20:93-106. (21)
1951.
Mammals
om
Ma capa a,
Sou heas e n
Pe u. PubI. Mus.
His .
Na .
"Ja ie
P ado,"
Se .
A,
Zool.,
6:
1-26. (22)
1954. Ba s
om
Chiman a-Tepui,
Venezuela
wi h
ema ks
on
Choe oniscus.
Fieldiana
Zool., 34:289-293. (23)

Mexico, Ins .
(28)
Johns
Hopk
ins P ess.
(29)
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PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
1955.
Rema ks
on
he
ba s
o
he
genus
Vllmpy ops.
Fieldiana
Zool.,
37:
403-413. (24)
SCHWARTZ,
A.,
AND
J. K.
JONES,
JR. ]967.
Re iew
o
ba s
o
he
endemic
An illean
genus
MO 1ophylllls. P oc. U.S.
Na .
Mus., 124(3635):1-20. (25)
STARRETT,
A.,
AND
L.
DE
LA
TORRE.
1964.
No es
on
acoJ)ec ion
o
ba s
om
Cen al
Ame ica,
wi h
he
hi d
eco d
o
Cy((a ops
alec o
Thomas.
Zoologia,49:53-63.
TABOADA,
G.
S.,
AND
R.
H.
PINE.
]969.
Mo phological
and
beha io al
e idence
o
he
ela ionship
be ween
he
ba
genus
B achyphyl/a
and
he
Phyllonyc e inae.
Bio opica,
1:10-]9.
(26)
TUTTLE,
M. D. 1970.
Dis ibu ion
and
zoogeog aphy
o
Pe u ian
ba s,
wi h
commen s
on
na u al
his o y.
Uni .
Kansas
Sci. Bull., 49:45-86. (27)
1974a. Ba
apping:
esul s
and
sugges ions. Ba
Banding
News,
]5:4-7.
1974/7.
Unusual
d inking
beha io
in
some
s enode mine
ba s.
Mammalia.
38:
141
-I
44.
1974('.
An
imp o ed
ap
o
ba s.
J.
Mamm.,
55:475-477.
VILLA-R., B. ]966.
Los
mu cielagos
de
Mexico. Uni .
Nac.
Au onoma
BioI., x i +491 pp.
WALKER,
E.
P.,
AND
OTHERS.
1964.
Mammals
o
he
wo ld.
Bal imo e,
1:xl iii + ]-644.
88
CARE
IN
CAPTIVITY
ARTHUR
M.
GREENHALL
In he pas
50
yea s only abou adozen a icles ha e appea ed ha deal solely
wi h ca e
o
ba s in cap i i y. Wi h he excep ion
o
he spec acula lying oxes
and he ampi e ba s, main enance
o
ba s ei he as expe imen al animals
o
as
zoological
pa k
exhibi s has been neglec ed. In e es in ba s as labo a o y animals
inc eased a e 1953, mos ly s imula ed by disco e y in he Uni ed S a es
o
abies in insec i o ous ba s (Cons an ine, 1970). As
i
became e iden ha public
heal h ela ionships exis ed be ween ba s and man, a emp s inc eased o s udy
li e ba s unde con ol1ed condi ions.
Valuable sou ces
o
he in o ma ion included
in
his chap e ha e been unpub-
lished manusc ip s
o
pape s in p ess. Iha e also included some pe sonal obse -
a ions whe e app op ia e. Iha e e iewed mos
o
he a ailable li e a u e, bu
despi e ac i e esea ch in e es , published in o ma ion
is
woe ully lacking o
cap i e Chi op e a, pa icula ly he Phyllos oma idae. Li le
o
no hing is known
abou he ca e
o
insec i o ous phyllos oma ids. Howe e , because anumbe
o
espe ilionids and molossids ha e been aised success ully on a i icial die s, I
ha e desc ibed die s, cage sys ems, and echniques o hei husband y, as apo-
en ial guide o he ca e
o
insec -ea ing phyllos oma ids.
Phyllos oma id ba s p esen a
nUITlbe
o
unusual main enance p oblems. Wha
hese p oblems a e, and how hey ha e been o may be sol ed,
is
he opic
o
his
chap e . This in o ma ion
is
in ended p ima ily o hose who main ain ba s
in
cap i i y o esea ch
o
educa ional pu poses and no o someone simply in-
e es ed
in
keeping ba s as pe s.
TRANSPORTATION
T anspo a ion
o
ba s o he labo a o y
o
zoo should be ca e ully planned.
Because phyllos oma ids a e mainly opical, anspo ime should be minimal,
and he shippe mus be awa e ha wea he condi ions may change apidly om
he ho lowlands o he cool uplands
o
om opical o empe a e la i udes.
I
is
o en possible o a ange o comme cial ca ie s o ake special in ansi p e-
cau ions wi h he animals being shipped. Iha e had excellen coope a ion om
ai lines and shipping companies in keeping my ba s away om ex eme hea o
cold
o
o he po en ially s ess ul si ua ions-.
Ba s may be anspo ed
in
me al cans, wi e cages, ligh -weigh wooden boxes,
o
in ca dboa d
o
plas ic ca ons. Mos ba s, unlike many o he mammals, will
no a emp o gnaw ou
o
con aine s, bu hey can squeeze h ough inc edibly
small holes and c acks. Ba s gene ally a el be e indi idually han in g oups,
wi h each animal placed
in
asingle compa men
o
in aligh clo h bag wi hin a
igid con aine . Vampi e ba s and ca ni o ous species should no be g ouped
wi h ba s
o
o he species.
Ca e
mus be aken o a oid exposing ba s o he sun,
o
p o ide p ope en ila ion, and o con ol empe a u e and humidi y.
Food
89
and wa e mus be p o ided o long ips. All pe sons wishing o anspo ba s
should be awa e
o
he ules and egula ions go e ning he na ional and in e -
na ional shipmen
o
li e animals.
On a i al, he shipping con aine s
in
which he animals a e ecei ed should
be ei he incine a ed
o
ho oughly cleaned and s e ilized o minimize con amina-
ion
by
disease o ganisms
o
pa asi es. I pe manen li ing qua e s a e una ail-
able, cleaned empo a y cages, such as hose used o shipping, could su ice.
The
physical condi ion
o
e e y ba should be assessed upon a i al.
THE
LABORATORY
ENVIRONMENT
Tempe a u e and Rela i e Humidi y
Among he mo e impo an ac o s in luencing he success ul main enance
o
ba s a e empe a u e and ela i e humidi y.
I
possible, hese should be con-
olled au oma ically. Unde na u al condi ions ba s a e exposed o daily luc ua-
ions
o
empe a u e and ela i e humidi y; howe e , li le is 'known abou he
op imum condi ions o cap i e opical ba s. Ne e heless, a empe a u e
o
20
o
25°C
and a ela i e humidi y
o
70
o 75 pe cen seems sa is ac o y o many
species. Low humidi ies can be inju ious o he wing memb anes (Racey, 1972).
Uwe Schmid (pe sonal communica ion) main ains his animal oom a acon-
s an empe a u e
o
27°C
and a ela i e humidi y be ween
65
and 75 pe
cen . This
is
sa is ac o y o Phyllos omus discolo , Ca ollia pe spicilla a,
A ibeus li u a us, and Desmodus o undus. Rasweile and de Bonilla (1972:659)
and Rasweile and Ishiyama (1973:56-57) main ained hei labo a o ies a
enlpe a u es be ween
21
and
28°C
and a ela i e humidi y be ween 55 and 92
pe cen , which p o ed sa is ac o y o Glossophaga so icina, Anou a caudi e ,
Phyllos omus discolo , Ca ollia pe spicilla a, A ibeus li u a us, and S u ni a
lilium.
My
ba labo a o y in T inidad was no ai -condi ioned. The daily
empe a u e anged be ween 21.1 and 29.4°C, and he ela i e humidi y be-
ween 55 and 95 pe cen , condi ions undoub edly sui able o Glossophaga
so icina, Phyllos omus discolo ,
P.
has a us, Vampy um spec um, Ca ollia
pe spicilla a, Vampy ops helle i, A ibeus jamaicensis,
A.
li u a us, and
Desmodus o undus
in
ha many
o
hese ba s li ed in he labo a o y o se e al
yea s.
The Desmodon inae do well unde a a ie y
o
labo a o y condi ions. Wimsa
and Gue ie e (1961 :450) main ained Desmodus o undus
in
an
ai -condi ioned
labo a o y a empe a u es be ween 20 and 25 °C and a ela i e humidi y be ween
30 and
65
pe cen , which app oxima ed empe a u es and humidi ies p e ious-
ly eco ded in Mexico. They also obse ed ha ampi es in he labo a o y ole -
a ed highe ela i e humidi ies and sho exposu es o lowe empe a u e, bu
ha hei ole ance
o
empe a u e abo e
25°C
was poo .
In
T inidad, G eenhall
(1965b:442) kep Desmodus and Diaemus in alabo a o y a empe a u es
be ween
21
and
29°C
and an a e age daily ela i e humidi y
o
75
pe
cen . The
colony
o
Desmodus and Diaemus s udied by Dickson and G een (1970:38)
in
London was kep
in
qua e s held a acons an empe a u e
o
240Cdu ing he
SPECIAL
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90
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
91
day bu which cooled na u ally o
21
°du ing he nigh . The ela i e humidi y
was ai ly cons an a abou 50 pe cen .
A
an ele a ion
o
2545 me e s, whe e
ampi es do no no mally occu
in
Mexico, Schmid and G eenhall (1972:243)
a emp ed o main ain alabo a o y empe a u e
o
be ween 25 °and
30°C
and
a ela i e humidi y abo e 55 pe cen o Desmodus.
A
his ele a ion he e we e
p oblems in con olling an accep able empe a u e and ela i e humidi y. Powe
ailu es, agg a a ed by he lack
o
as andby gene a o o eme gency use, com-
plica ed ma e s. We no mally used he nl0s a ically con olled elec ic hea e s
and in a ed hea lamps o main ain he empe a u e and cool mis elec ic
hunlidi ie s o con ol ela i e humidi y. Uwe Schmid (pe sonal communica ion)
epo ed Desmodus h i ed in
his
animal oom in Ge many, which was main-
ained a acons an empe a u e
o
27°C
and a ela i e humidi y
o
be ween 65
and
75
pe cen .
Daily luc ua ion in empe a u e and ela i e humidi y can be moni o ed wi h
ahyg o he mog aph. The he mog aph eadings should be calib a ed wi h an
accu a e nlinimum-maxinlum he monle e ; he hyd og aph wi h awe and d y-
bulb hyg ome e .
Ven ila ion.-
The e
is
p ac ically no in o ma ion on he impo ance
o
en ila ion and ci cula ion o ai
in
labo a o ies housing ba s. Pye (I967) cau ions
ha many ba species a e sensi i e o d augh s and o e en ila ion should be
a oided. In
my
expe ience, ba s
in
poo ly en ila ed labo a o ies appea es less.
Ligh
Ligh appea s o be an impo an ac o in egula ing he daily ac i i ies o
ba s (DeCou sey and DeCou sey, 1964). Uwe Schmid (pe sonal communica-
ion) claimed ha acolleague in es iga ing daily ba ac i i y ound ha Phyl-
los omus discolo , Ca ollia pe spicilla a, and A ibeus li u a us died when kep
con inuously
in
o al da kness o
10
days. Illumina ion
is
au oma ically con-
olled in many labo a o ies and
13
hou s
o
ligh and
II
hou s
o
da kness has
been ound o be sa is ac o y o Phyllos omus discolo , S u ni a lilium, and
A ibeus li u a us (Rasweile and Ishiyama, 1973), Glossophaga so icina, Anou a
caudi e , and Ca ollia pe spicilla a (Rasweile and de Bonilla, 1972), Desmodus
o undus and Diaemus youngii (Dickson and G een, 1970), and Desmodus
o undus (Schmid and G eenhall, 1972). Many
o
he same phyllos oma ids,
excep Diaemus, ha e been displayed success ully in ala ge simula ed Sou h
Ame ican ca e and opical ain o es exhibi a he New Yo k Zoological Pa k
whe e,
by
a ying he in ensi ies
o
whi e, blue, g een, and occasionally ed ligh ,
he ac i i y pa e ns
o
he ba s we e e e sed. Vampi e ba s kep by Wimsa
and Gue ie e (1961) we e subjec ed o low-in ensi y illumina ion om ligh
en e ing h ough wo glass-b ick windows and glass panel in adoo . The ba
cage i sel u he educed he ligh because only he on was made
o
ans-
pa en ma e ial. Elec ic ligh s we e u ned
on
in he ba oom only b ie ly when
he cages we e cleaned o he animals we e being a ended. No e o was made
by
he in es iga o s o con ol he ligh egime because wild ampi es a e ound
in oos s ecei ing a ying in ensi ies
o
ligh .
Insec i o es
disin ec an wi h a esidual a ailable iodine
o
a leas
1:
10,000,
o
1%concen a-
ion
o
soapy wa e
o
de e gen can be used. The solu ion should be au ocla ed
and disca ded a e each use. Ho soapy wa e
o
de e gen can be used o swab-
bing loo s and ables.
"Glasswa e, plas icwa e and ins umen s
...
should be disca ded in o plas ic
o
glass ecep acles con aining one
o
he disin ec an s men ioned abo e. They
should be au ocla ed.
"Ca casses and animal issue
...
a e bes disposed
o
in
plas ic bags and
incine a ed."
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In discussing he eeding habi s
o
he Phyllos oma idae in ano he chap e ,
Ga dne no ed ha he amily's only ue insec i o e may be Mac ophyllum
mac ophyllum. He ci ed examples
o
insec emains ha ing been ound
in
he
s omachs
o
ecal emains
o
all sub amilies, sugges ing ha , ega dless
o
he
basic ood p e e ences
o
each g oup, insec s a e p obably an impo an ood
componen
o
he die . Howe e ,
in
he case
o
he Desmodon inae, insec s
p obably we e inges ed acciden ally when p epa ing abi e si e
on
some animal
selec ed
as
p ey,
o
du ing g ooming ac i i y. Racey (1972:297-299),
in
his
e iew
o
he ca e and managemen
o
ba s, lis ed 33 gene a and
54
species
o
insec i o ous ba s ha ha e been kep
in
cap i i y, bu he only phyllos oma id
men ioned was Mac o us. Mic onyc e is mega/o is was kep
by
Ruschi (1953a),
who indica ed ha he ba s li ed and ep oduced, bu he did no p o ide u he
in o ma ion. Wi h he excep ion
o
Mac o us and Mic onyc e is, species
o
which
also ea ui , i
is
no su p ising ha he li e a u e
is
wan ing on he ca e
o
DIET
O
all ba s, he Phyllos oma idae p obably ha e he mos a ied ood
p e e ences. Thei na u al eeding habi s a e discussed by Ga dne ( his olume),
and his chap e on ood habi s will be indispensable o anyone who mus p epa e
adie o any phyllos oma id no ye success ully kep in cap i i y. Excep o a
ew
species, he li e a u e
is
meage abou he die s used
by
labo a o ies o
zoological pa ks when keeping New Wo ld lea -nosed ba s in cap i i y.
Fo his discussion Iha e g ouped he phyllos oma ids
as
insec i o ous,
nec a i o ous, ugi o ous, omni o ous, ca ni o ous, and sangui o ous. No ba
in cap i i y can ea exac ly
as
i
would unde na u al condi ions and speci ic
na u al ood i ems may be impossible o supply, making subs i u ions essen ial.
Adaily in ake
o
p o ein appea s necessa y, bu i is impossible o s a e wha he
equi ed amoun s should be in o mula ing abalanced die . Hope ully, howe e ,
wi h i amins, mine als, and o he ood addi i es, he nu i ional equi emen s
o
acap i e ba may be esol ed. Va ious i amin p epa a ions a e a ailable and
many in es iga o s ha e hei own p e e ences. One, S ua Fo mula Liquid,
seems almos o be apanacea o die de iciencies, no only o ba s bu o o he
cap i e small mammals (see
Appendix]
8 o sou ces
o
p oduc s men ioned
in
his chap e ).
98

insec i o ous phyllos oma id ba s. Ibelie e, he e o e, ha
i
will be
o
alue o
desc ibe die s used success ully o main ain insec i o ous ba s.
Insec
die s.-Vespe ilionid
and molossid ba s ha e been ed a a ie y
o
in-
sec s such as g eenbo le lies (bo h adul s and maggo s), house lies, ins a s
o
g asshoppe s, locus s, and c icke s in addi ion o bees,
June
bee les, e mi es,
waxwo ms, and waxmo hs (Ga es,
1936:270~
Ramage, 1947:61). Mos cap i e
insec i o ous ba s h i e on adie
o
mealwo ms (Teneb io
moli o )~
al hough
he la ae a e p e e ed, he pupae also a e ea en.
Pye (1967) cau ioned ha die a y de iciencies may occu i ba s a e ed meal-
wo ms ha ha e had apu ely a inaceous die . This de iciency may be o e come
by
he addi ion
o
agood quali y comme cial animal eed o he mealwo m's die .
Also, mealwo ms may be dus ed wi h i amin and mine al mixes (Rasweile ,
1975) o coa ed wi h i amin d ops such as hose used o child en (Ga dne ,
pe sonal communica ion). Conce ning adie
o
mealwo ms, Racey (] 972) and
Ladische e
al.
(1967) ad ised ha he e may be some oxic quinones
in
some
mealwo m imagoes.
Ga es (1938b) added pieces
o
honey bees o he die s
o
Ep esicus, Myo is,
Lasiu us, Pleco us, and Tada ida. The bees seemed o imp o e he consis ency
o
he eces and added o he pala abili y
o
he ood. He ied o he insec s, such
as g asshoppe s and
June
bee les, bu he ba s p e e ed bees. Ramage
(1
947:
61) had no success
in
pe suading a ious species
o
Myo is and Ep esicus o ea
he oods sugges ed
by
Ga es (1936) un il she p o ided he la ae, pupae, and
adul s
o
g eenbo le lies, "which can be easily ea ed
in
eno mous quan i ies."
AMyo is cali o nicus she kep e used o ea lies bu was aised success ully on
e mi es. Ramage (1947:61) commen ed, howe e : "Te mi es ha e he dual
disad an age ha hey canno be cul u ed apidly enough o eed he ba s and
mus be chlo o o med
o
killed o keep henl om c awling away be o e he
ba s ha e achance o ea hem."
Racey (1972:302), in his discussion on insec s as ood o insec i o ous ba s,
men ioned ha he la ge ba species also will ake ea ly ins a s
o
many
O hop e a,
" he
mos commonly cul u ed
o
which a e locus s and cock oaches."
O
(1954:168) mos ly ed mealwo ms o
An ozous
pallidus du ing he ea ly
phases
o
his s udy, bu la e he used ap epa ed die ecommended o him by
E nes
P.
Walke . Howe e , he o e ed (p. 234) alis ing
o
o he kinds
o
animal
oods such as a a ie y
o
lies, mo hs, and e en snails. Elsewhe e in his s udy,
O
(op. ci ., 232-233) ci ed, " eco ds
o
cap i e pallid ba s which we e obse ed
o ea wes e n skinks (Eumeces skil onianus), aSono an dese gecko (Coleonyx
a iega us), and we e suspec ed
o
ea ing he head and neck
o
aMexican ee-
ailed ba (Tada ida mexicana).
I
seems likely ha s a a ion was esponsible o
such de ia ion om
an
insec i o ous die , al hough
...
i
is
possible ha small
nigh liza ds may be p eyed upon locally by pallid ba s."
A i icial
die s.-l
ha e used insec aps o ca ch insec s o ba ood. A
imes, howe e , insec s may be ei he sca ce
o
no a ailable. Consequen ly, in-
es iga o s ha e had o de ise subs i u e die s. These a e mashes
o
mix u es com-
p ised
o
anumbe
o
i ems ha a e eadily aken
by
he ba s and usually in-
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
99
100
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clude such hings
as
banana, co age cheese, ha d-boiled egg, and i amins. Such
mix u es a e commonly called
Hgl
op
"by ba biologis s. Ga es (1936, 1938b)
ound ha a ious species
o
Myo is, Pipis ellus, Ep esicus, and Pleco us do well
in cap i i y
on
a i icial die s ha may be comple ely di e en om hei no mal
die s. He (1938b: 157) no ed:
~'Unde
cap i e condi ions hey ha e been known
o ea p ac ically e e y hing, unless
i
is
oo highly seasoned. This includes all
ce eals, b eads, c acke s, cakes, mea s, eggs, ege ables
o
all kinds, bo h esh
and cooked, le uce, cele y, and all
o
he no oo acid ui s, apples, pea s,
peaches, p unes, pineapples, and igs. All milk p oduc s, bo h esh and sou
milk, bu e milk, cheese, and e en bu e a e accep able.
In
ac , he au ho has
ha dly ound any ood which hey will no ea . They
o
cou se ha e hei p e -
e ences, appa en ly p e e ing he milde cheeses o any hing else. Howe e ,
b ead c umbs mois ened wi h bu e milk a e also g ea ly enjoyed." Ga es (1936:
270) i s sugges ed ha chi in was essen ial o he
p ope
o ma ion
o
ecal
pelle s and he p e en ion
o
in es inal obs uc ion in insec -ea ing ba s. The
ease
o
ob aining bananas and co age cheese emp ed Racey (1970) and o he s
o eed amash lacking insec s. The ba s did poo ly, howe e , and hei pelage
de e io a ed. Empi ically,
i
was disco e ed ha mealwo ms
o
o he insec s
added o he mix u e co ec ed he condi ion.
The e a e anumbe
o
ecipes o glop. Walke (1966: 138) de eloped a ood
mix u e elished by many small mammals a he Na ional Zoological Pa k (Ap-
pendix 9). Da is and Luckens (1966) used banana, c eam cheese, canned dog
ood, and mul i i amins o eed
Ep e
sicus (Appendix 10). Mohos (1961) used
he Walke (1966) o mula o eed Myo is, Pipis ellus, and Ep esicus. How-
e e (p. 371), Hoccasionally equal pa s
o
bee and bee li e we e subs i u ed
o he co age cheese, since
i
was ound ha , a e an ini ial adap a ion pe iod,
he ba s a e equally well on his die ." J. F ede ick Bell (pe sonal communica-
ion) ed Myo is luci ugus homogenized whole baby mice, which we e eadily
a ailable in his labo a o y. K u zsch and Sulkin (1958:262-264) ied a
numbe
o
eeding echniques and ood combina ions o induce hei cap i e
Tada ida b asiliensis o eed. Li e mealwo m la ae we e unsa is ac o y, and a
nu ien luid con aining amino acids, simple suga s, and i amins caused he
ba s o de elop dysen e y wi h a al esul s. They we e inally success ul
in
main-
aining Tada ida on glop.
Food
s o age.-Die
p epa a ion may be simpli ied in ha he a ious ood
ing edien s, including insec s, can be mixed in an elec ic blende , p eweighed
in wax pape
o
plas ic bags, and hen s o ed ozen a
4°C
un il used (Mohos,
1961
:37]
;Da is and Luckens, 1966:226; Ba bou and Da is, 1969:246; Ras-
weile , 1975).
The
size
o
he ood packe s should depend on he numbe
o
ba s
o be ed a
anyone
ime. Be o e eeding he ba s, ozen ood should be e-
mo ed om he eeze and allowed o haw. Once hawed, ood will las abou a
week unde o dina y e ige a ion.
I
is
impo an no o se e we mashes and
o he liquid die s oo ea ly in he day because hey may begin o spoil be o e
all ba s ha e ed. This can lead o dia hea and malnu i ion.
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
Nec a i o es
101
Un il ecen ly he s anda d die o such cap i e nec a - eeding ba s as G/os-
sophaga and
Anou a
has been suga
o
honey
in
wa e , ui juice, succulen
ui s, and i amins (Ruschi, 1953c, 1953d, 1953e; Goodwin and G eenhall,
1961 ;McNab, 1969). Rasweile (1975) claimed ha
~'labo a o y
die s o
ugi o ous
o
nec a i o ous species based solely upon ui
pUlp
andlo ui
juices may be g ossly inadequa e om anu i ional s andpoin ." He belie ed
ha addi ion
o
insec s and pollen could signi ican ly inc ease he p o ein, a ,
mine al, and i amin le els in die s o ui -ea ing ba s, as well as supply essen ial
amino acids ha a e inadequa ely ep esen ed in he ui componen
o
he
die . Rasweile and de Bonilla (1972) and Rasweile (1973) ha e o mula ed
die s (Appendix 12) ha ha e been success ul o he long- e m main enance
o
la ge numbe s
o
G/ossophaga so icina,
Anou a
geo oyi, A. caudi e , and
Ca ol/ia pe spicil/ala.
The New Yo k Zoological Pa k p epa es an a i icial nec a (Appendix 11)
dispensed om la ge wa e ing bo les hidden among he plas ic plan s
o
he
exhibi . In hese exhibi s, he nec a - eeding ba s also ha e access o he solid
die (Appendix 15) o e ed o he ugi o ous phyllos oma ids (House and
Dohe y, 1975). The ba s housed in his exhibi ha e included G/ossophaga
so icina, Anou a geo oyi, Phyllos omus discolo , and Ca ol/ia pe spicil/a a.
Donna J. Howell (pe sonal communica ion) success ully aised nec a i o ous
ba s
on
adi e en die (Appendix 13). She also main ained Phyllos omus dis-
colo and Ca ol/ia pe spicil/a a, bu ea ed hem
as
ui ea e s. Howell w o e:
HI' e kep Lep onycle is, Choe onycle is, Glossophaga, Anou a, and Hylonyc-
le is on he nec a die o pe iods exceeding ayea . All he die ing edien s
seemed necessa y o duplica e he e y nu i ious con en s
o
'ba -adap ed' pollen
and nec a om chi op e ophilous plan s. One mus be ca e ul
o
gi e he ba s
enough p o ein, ye no o e p o ein-load
o
o e suga -load hei kidneys. Con-
cen a ing abili y
o
he glossophagines is e y poo . P o ein should s ay abou
9 o 11 pe cen , suga
14
o 20 pe cen ."
F ugi o es
The
die
o
nec a i o ous and ugi o ous ba s
is
in luenced mos ly by he
seasonal abundance
o
lowe s and ui s (G eenhall, 1956, 1957; Goodwin and
G eenhall, 1961; Fleming el ai., 1972).
The
die s
o
cap i e ugi o es will be
de e mined by he a ailabili y
o
ui s. In empe a e egions, opical ui s, wi h
he excep ion
o
bananas, may no always be a ailable.
Cap i e ui -ea ing ba s gene ally p e e swee ui s such
as
bananas, man-
goes, peaches, plums, melons, g apes, and papayas (Pye, 1967; Racey, 1972).
Al hough ci us ui s a e no p e e ed, swee o anges and g ape ui occasional-
ly
may be accep ed (G eenhall, 1966). Pye (1967) men ioned ha apples can
be subs i u ed i exo ic ui s a e in sho supply.
The
banana-based die o mu-
la ed
by
Rasweile (1975) and Rasweile and de Bonilla (1972) is gi en
as
Ap-
pendix 14.
102
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When o e ing ood, i
is
ad isable o use small pieces, abou I-cen ime e
cubes, well mixed. This will p e en dominan indi iduals om selec ing all he
choice ui s
o
ob aining mo e ood han hei cage ma es. Small pieces can be
easily managed by ba s while ea ing
o
lying. The smalJe ui -ea ing ba s can
handle whole ui s only when hey a e o e ipe
o
so enough ha asmall hole
can be bi en
in
he skin h ough which he ui pulp and juice can be ex ac ed.
Some biologis s ha e ed phyllos oma ids some unusual die a y i ems.
Fo
example, Ruschi (1953j) ed A ibeus li u a us blood
as
well
as
ui and insec s,
because
he
belie ed ha his species eeds upon blood
in
he wild.
Goodwin and G eenhall (1961) s a ed ha Phyllos omus discolo will no ea
lesh
in
cap i i y and p e e s ui such as bananas, mangoes, and papayas.
McNab (] 969) epo ed, howe e , ha cap i e
P.
discolo equi e asmall, bu
egula , in ake
o
mea . Uwe Schmid (pe sonal communica ion) men ioned
ha ing success ulJy ea ed his species on sliced bananas, mealwo ms, and
neona al labo a o y mice.
McNab (1969) epo ed ha he kep anumbe
o
ugi o ous phyllos oma ids
heal hy o ex ended pe iods
o
ime, bu , un o una ely, p o ided no in o ma ion
on die s o he han he ac ha Rhinophylla pumilio, U ode ma biloba um,
A ibeus cine eus, and
A.
concolo we e ed ui . His Phyllos omus elonga us
and Vampy essa nymphaea would no ea .
Polyphagous Phyllos oma ids
Agood example
o
an omni o ous phyllos oma id
is
Phyllos omus has a us,
which eadily adjus s o cap i i y.
Dunn
(J 933) ed
P.
has a us mice, ba s, bi ds,
de ib ina ed blood, ui , and aw mea , including li e . Ruschi (1953b) and
McNab (] 969) p o ided simila a e, he o me adding cock oaches when a ail-
able. Goodwin and G eenhall (J
96])
ound ha his species, in addi ion
o
ac-
cep ing awide a ie y
o
ui , h i ed on mice and young bi ds.
I
did no hesi a e
o kill and ea o he ba s placed in i s cage al hough
i
appea ed
o
be uneasy
in
he p esence
o
Desmodus.
The
die used by he New Yo k Zoological Pa k o
hei colony
o
P.
has a us
is
gi en in Appendix 16.
Donna
J. Howell (pe sonal
communica ion) ed bo h Phyllos omus has a us and
P.
discolo aspecial die
o ugi o ous ba s (Appendix 17) and commen ed ha hese ba s also go a
dish
o
mealwo ms a each eeding.
Ca ni o es
The
bes known ca ni o ous phyllos oma id is Vampy um spec um.
I
has
been aised success ully
in
cap i i y on aw mea as
welJ
as dead whole chicks
and pigeons (Di ma s, 1935, J
936;
Goodwin and G eenhall, 1961; C andall,
1964; B adbu y, 1970). G eenhall (1968) desc ibed he ca e
o
Vampy um,
which success ully aised young du ing he i e yea s ba s we e main ained
in
cap i i y. They we e ed pigeons, chicks, wild bi ds, and dead labo a o y a s
and mice as
weI)
as aw mea cu in o 2-cen ime e chunks. Al hough hese
Vampy um ne e ough o e ood, he e was always he isk
o
inju y o hei
wings when hey s alked li e p ey because he cage was oo small. The e o e, all
BIOLOGY OF
THE
PHYLLOSTOMATIDAE 103
ood was killed and usually p esen ed o he ba s by o ceps
o
placed on he cage
loo . Food was ho oughly mas ica ed
by
he ba s; ea he s and oden ails
usually we e disca ded.
O he ca ni o ous phyllos oma ids kep in cap i i y include T achops ci -
hosus and Phyllode ma s enops, which a e liza ds (Pye, 1967), Ch o op e us
au i us, which was kep on adie
o
whi e mice
by
Villa-R. and Villa-C. (1969),
and mea and ba s
by
McNab (1969).
Vampi e Ba s
The h ee ampi e ba s, Desmodus o undus, Diaemus youngii, and Diphylla
ecauda a ha e been main ained in cap i i y wi h a ying deg ees
o
success.
Desmodus adap s easily o cap i e condi ions, whe eas Diphylla
is
mos di icul
o main ain. Diaemus does ai ly
well
once i s basic equi emen s a e ecognized
and me . P esen ly he e a e b eeding colonies
o
Desmodus
in
many labo a o ies
and zoological pa ks.
Di ma s and G eenhall (1935) we e he i s o desc ibe keeping ampi e ba s
in cap i i y. T apido (1946) demons a ed ha Desmodus easily adap ed o
labo a o y condi ions and epo ed alonge i y eco d
o
12
yea s. Wimsa and
Gue ie e (1961) de ailed he ca e
o
Desmodus
in
empe a e zone labo a o ies,
whe eas G eenhall (1965b) desc ibed i s main enance in he opics.
Wild Desmodus may consume amoun s
o
blood equalling
o
exceeding hei
body weigh (Wimsa , 1969). The a e age weigh
o
Desmodus
is
abou 30
g ams, and cap i e ba s daily may d ink up o 50 millili e s
o
blood, al hough
15
o
20
is
usually su icien (Pye, 1967). Wimsa and Gue ie e (1962) ci ed
he unusual capaci y
o
one cap i e 28-g am nonp egnan emale hey main ained
in
isola ion o
17
days, which consumed blood
in
excess
o
he body weigh on
13
days.
On
wo days he amoun s inges ed we e 47 and 52 millili e s, espec-
i ely.
Dickson and G een (1970), in o de o educe he ime and labo equi ed in
he main enance
o
ampi es, u ilized isodium ci a e
(1
millili e 3.8 pe cen
pe 10 millili e s
o
blood) o p e en blood coagula ion and dispensed he blood
meal in plas ic hoppe s. The use
o
ci a ed blood ins ead
o
de ib ina ed blood
educed he p epa a ion ime and dispensing meals in plas ic hoppe s p e en ed
spla e ing and spillage he eby educing he ime o cage cleaning. Acco ding
o Dickson and G een (1970), de ib ina ion esul s
in
he loss
o
blood olume
and he emo al
o
ac o s essen ial o he ampi e's wel a e. Blood can be
s o ed ozen a -20°C o up o six weeks wi hou becoming unpala able o
ampi es. Dickson and G een (1970) p esen ed he daily a ion wa med o
37°C
a 1630 hou s. Each ood con aine was illed wi h 100 millili e s
o
blood-
su icien o i e ba s. These hoppe s o glass- ubed d inking bo les p e-
en ed he was age and con amina ion
o
he blood
by
u ine and eces ha
usually occu s when ampi es a e ed om open dishes. Radiological examina-
ion e ealed no e idence
o
bone decalci ica ion ollowing
15
mon hs o ci a e
in he die .

104
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F ozen blood, whe he de ib ina ed
o
ci a ed, has a endency o spoil apid-
ly
a e hawing (Wimsa and Gue ie e, 1961; G eenhall, 1965
b).
The e o e,
he blood meal should
be
o e ed when he ba s no mally commence eeding.
In he ield, whe e esh blood
is
usually di icul o ob ain, an ingenious
eme gency supply was de ised
by
Wimsa and Gue ie e (1961). This "ins an
blood"
is
shell- ozen and lyophilized de ib ina ed blood and
is
econs i u ed
wi h wa e ,
as
needed.
Diaemus has been kep success ully in he labo a o y on ci a ed bo ine blood
- he
same die ed o Desmodus (Dickson and G een, 1970). Aweekly supple-
men
o
chicken blood was also p o ided by pe mi ing he ampi es o eed on
he oes
o
ali e chicken placed on he op
o
hei cage. None heless, Goodwin
and G eenhall (1961) epo ed ha Diaemus e used o d ink de ib ina ed ca le
blood e en when mixed wi h chicken blood.
Thei
ba s, howe e , would accep
chicken blood.
A
ala e da e, G eenhall (1970) epo ed obse ing Diaemus
eeding on ca le. Ibelie e ha Diaemus should be ed ci a ed whole blood
as
indi iduals appa en ly e use o d ink de ib ina ed blood.
Diphylla has no been success ully kep in cap i i y o any leng h
o
ime.
Ruschi (195
1)
s a ed ha he main ained Diphylla and
Desmodus
on bo h
ci a ed and de ib ina ed blood, and ha he la e was p e e ed. Howe e , Villa-
R. (1967) epo ed ha Diphylla would no accep ca le blood, and al hough
de ib ina ed chicken blood was consumed, he ba s died wi hin 48 hou s. Pe haps
cap i e Diphylla would accep ci a ed a he han de ib ina ed blood.
I ecommend ha liquid mul i i amins be added o all blood meals. House and
Dohe y (1975) used 2.4 cubic cen ime e s
o
"Pe
D ops"
pe pin
o
blood
(Appendix 18).
An unusual obse a ion abou ampi e ba die was epo ed by Kumm
(1932), who no ed ha Desmodus ed eadily
in
cap i i y on bananas and li e
animals. Ican ind no con i ma ion ha ampi es eed
on
ui .
Vampi e
as ing.~chmid
e a . (1971) obse ed ha wild Desmodus may
no o age e e y nigh . I ue, one mus assume ha such as s a e no dele e i-
ous. Wimsa and Gue ie e (1961) ou inely skipped eeding hei
Desmodus
on Sundays.
On
he
ew
occasions ha hei ba s we e neglec ed acciden ally o
as long as h ee days, he e ec was ma ked emacia ion bu no dea h. G eenhall
(l965b:443)
ound ha
Desmodus
could go wi hou ood o a leas 62 and a
hal hou s wi hou appa en ill e ec s.
Vampi e
cannibalism.-
The only e e ence o cannibalism among Desmodus
in cap i i y was desc ibed by Win1sa (1959). Wimsa 's colony consis ed
o
ou cages, each con aining
15
o 20 ba s. In one cage he daily a ion
o
blood
was always consumed, whe eas
in
he
o he
h ee he e was always some blood
le o e . A e se e al weeks, he ba s
in
he i s cage de eloped hai less pa ches
on he shoulde s, in e scapula egion, and back
o
he head. Associa ed lace a-
ions we e ob iously caused by bi es. This did no occu wi h he occupan s
o
he o he h ee cages, sugges ing ha his si ua ion e lec ed unusual beha io .
This abno mal beha io ceased and he inju ed animals subsequen ly eco e ed
when blood was supplied
in
adequa e amoun s.
The
eeding on cage ma es was
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
105
in e p e ed o
be
an a emp o secu e blood
in
any way possible whene e he
amoun
o
blood a ailable was de icien . G eenhall (1965b:442) obse ed ha
cannibalism migh occu anl0ng newly cap u ed ampi es, bu could easily be
p e en ed by supplying he ba s wi h
.a
g ea e quan i y
o
blood han could
be
consumed in one nigh .
P epa a ion
o
de ib ina ed and ci a ed
blood.-A
majo obs acle o keeping
a ampi e ba colony in he labo a o y
is
he unce ain y
o
acon inuous esh
supply
o
blood. De ib ina ed ca le blood has been used success ully o many
yea s, bu ci a ed blood may be p e e able inasmuch as no hing
is
emo ed.
Blood o de ib ina ing should i s be collec ed in aclean essel as
i
1l0ws
om he slaugh e ed animal. The blood mus hen be whipped o agi a ed wi h
ei he wooden applica o s, cocoa bea e s, swizzle s icks, la ge wi e bea e s,
whisks,
o
oughened glass beads. A e se e al minu es
o
agi a ion, he ib in
will adhe e o he bea ing ins umen
o
glass beads and hen may be disca ded.
The emaining blood may hen be pou ed in o ano he clean essel o s o age.
De ib ina ed blood will emain
as
aliquid and will keep unde e ige a ion o
a ew weeks. The colo , usually b igh ed, migh slowly u n pu plish. The blood
will be accep ed by he ba s i no spoiled. G eenhall (1965b) oze de ib ina ed
blood in ice cube ays and o he con aine s. F ozen de ib ina ed blood may be
main ained sa ely o se e al weeks. Inasmuch
as
i may spoil apidly, only single
a ions
o
blood should be hawed each day, and his should be used as quickly
as
possible. Iknow
o
some labo a o ies and zoos ha ha e used ou da ed human
blood and plasma om blood banks.
The
consensus
is
ha ampi es appa en ly
do no do well on ei he a e.
Dickson and G een (1970:40) desc ibed hei me hod
o
ci a ing blood
as
ollows: HBo ine blood
is
collec ed om anea by slaugh e house in 5li e con-
aine s, each con aining
500
ml
o
3.8 pe cen isodium ci a e o p e en
coagula ion. This blood
is
s ained h ough muslin in o
500
ml
poly hene bo les;
i can be s o ed ozen a -200C o up o 6weeks wi hou becoming unpala able
o he ba s."
DRINKING
WATER
In hei e iews
o
he ca e and managemen
o
ba s, Pye (1967:498) and
Racey (1972:303) s essed ha all cap i e ba s should ha e aplen i ul supply
o
d inking wa e a ailable. Mos in o ma ion abou wa e equi emen s pe ains
o espe ilionid and molossid ba s. Pe haps only a ew phyllos oma ids, such
as
Desmodus, a e able o li e o ex ended pe iods wi hou d inking wa e . Mos
ba s will lea n o use d inking ubes, nozzles
o
in e ed bo les (Racey, 1972:
303),
o
plas ic hoppe s
o
he ypes used o caged bi ds. Pye (1967), howe e ,
asse ed ha ba s do no eadily use d inking ubes and ha wa e
is
bes sup-
plied in shallow dishes. These wa e dishes should be cleaned daily. Acco ding o
Pye (1967:498), he e
is
adange ha sick animals may d own
in
wa e dishes
and he dep h
o
he wa e , o smalle ba s, should no exceed 3 o 5millime e s.
Some ba s a e eluc an o c awl on he loo so ha ood and wa e dishes should
be placed
on
ashel , as ened o he side,
o
suspended om he op
o
he cage.
GENERAL
CARE
OF
CAPTIVE
BATS
Acclima ion
The e
is
li le in o ma ion abou he acclima ion
o
ba s o cap i e condi-
ions. Rasweile (1973) ini ia ed acclima ion
o
his phyllos oma ids immedia ely
SPECIAL
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106
Racey (1972) obse ed ha espe ilionids will lap wa e om sa u a ed co on
and his
is
ause ul way
o
p o iding wa e du ing anspo . Rasweile (1975)
daily p o ided wa e o his cap i e Phyllos omus discolo ,
P.
has a us,
A ibeus
li u a us, and S u ni a lilium.
The e a e di e gen opinions
as
o whe he cap i e
Desmodus
o undus equi e
wa e o d ink. King and Saphi
(1
937) and T apido
(1
946) p o ided wa e
ad
libi um o hese ampi es. Wimsa and Gue ie e (1961) exp essed su p ise ha
hei
Desmodus
d ank li le i any wa e ; bowls
o
esh wa e we e kep in he
cages a all imes, bu we e no used e en when he ba s missed aday's eeding.
They inally discon inued he p ac ice and, a e nea ly wo yea s wi hou wa e ,
he ba s showed no
ill
e ec s. G eenhall
(l965b)
obse ed ha his ampi e
colony equi ed acons an supply
o
wa e o he ba s' well-being. In addi ion o
wa e supplied du ing he day, ano he 150 millili e s was gi en wi h he daily
meal. Du ing a wo-mon h check pe iod (May and June), 113 millili e s was he
g ea es amoun
o
wa e consumed
in
one nigh , whe eas 10 nlillili e s was
he leas . Wa e e apo a ion was no a ac o owing o he cons an high humidi y.
A high ele a ions
in
Mexico, G eenhall e al. (1971) no ed ha he daily blood
in ake
o
Desmodus
kep in glass ja s inc eased when asupply
o
d inking wa e
was added. Howe e , when la ge numbe s
o
ampi es (abou 200) we e kep a
one ime, wa e was no p o ided
in
o de o educe he main enance equi ed o
clean up o 100 ex a wa e hoppe s. O e ape iod
o
mon hs he ba s su e ed
no
ill
e ec s
as
a esul
o
wi hholding wa e . Wa e was always p o ided o he
small g oup
o
ampi es
in
he ligh cage. Uwe Schmid (pe sonal communica-
ion) p o ided his
Desmodus
wi h wa e
ad
libi um day and nigh , and blood
only du ing he nigh . He w o e:
HI
don' know whe he wa e
is
essen ial o
keeping ampi es, bu my eeling
is
hey need i o hei well-being, especially
i
hey ha e ea en spoiled blood." While main aining acolony
o
Desmodus
in
a ligh cage, Rex o d
D.
Lo d (pe sonal communica ion) obse ed:
"The
o he day Isaw a ampi e come down and d ink wa e e en be o e d inking blood
which was also a ailable. He d ank abou 5millili e s
o
wa e , hen lew up in o
he oos box." Lo d desc ibed a ood p e e ence es whe e wa e was o e ed o
]9 ba s. Each d ank an a e age
o
23.4 millili e s
o
blood and 2.5 millili e s
o
wa e nigh ly, o e ape iod
o
nine nigh s. On one nigh , no wa e was con-
sumed bu du ing ano he nigh he ba s d ank a o al
o
I32 millili e s. Lo d
conduc ed ano he es whe e ampi es we e kep indi idually
in
oil can cages
placed
in
an ou doo enclosu e. Wa e consump ion a ied wi h he ambien
empe a u e.
On
ex emely ho days, he ba s consumed 20 o 30 millili e s
o
wa e , d inking e en du ing he day.
On
cool days he ba s did no d ink. Lo d
concluded:
"I
hink wa e
is
necessa y
in
ho clima es
o
when he e
is
no con ol
o
cage enlpe a u e."
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
107
ollowing cap u e. In he ield, ood
is
placed
in
he anspo con aine s. Upon
a i al a his labo a o y,
all
ba s a e hand- ed o p o ide some nou ishmen .
This quick in oduc ion o he
new
die p obably assis s he ba s o cope wi h he
auma associa ed wi h cap u e, ansi , and he ini ial pe iod
o
adjus men o
cap i i y. Recalci an , weakened,
o
o pid Glossophaga so icina, o ex-
ample, usually can be induced o consume some
o
hei new die , i a i s a
small amoun
is
placed o e he ba 's nos ils (Rasweile , 1975). Asecond me hod
o
eeding ba s ha a e eluc an o accep hand-o e ed ood
is
o o ce ca e ully
an
eyed oppe con aining he liquid die in o he ba 's mou h (Rasweile and
Ishiyama, 1973:57). Upon as ing asmall sample
o
ood, many animals will
eadily consume mo e. The ba s a e hen ed om dishes. The same egimen was
applied o Ca ollia pe spicilla a, Phyllos omus dicolo , A ibeus li u a us, and
S u ni a lilium. Rasweile and Ishiyama (1973:58) s a ed:
"The
hand- eedings
we e gi en
in
he hope ha hey would acili a e he ansi ion
o
he animals
o alabo a o y exis ence."
T aining Ba s o Ea Mealwo ms
In he wild, mos insec -ea ing ba s ca ch hei p ey
in
ligh , whe eas a
ew
may glean hei p ey om oliage
o
o he subs a es. In cap i i y, whe e ood
is
placed
in
con aine s, many insec i o ous ba s equi e aining be o e hey
wilJ
eed hemsel es. Racey (1972), who has had much expe ience wi h cap i e in-
sec i o ous ba s, emphasized ha his aining
is
he nlos c ucial and ime-
consuming s age in he acclima ion
o
hese ba s o cap i i y. This
is
also ue
o
weanling ba s bo n
in
cap i i y. No ick (1963:51) s a ed ha cap i e Mac o us
mus be hand- ed mealwo ms be o e hey adap o aking hei own ood om a
dish. Al hough anumbe
o
o he phyllos oma id ba s eed on insec s
in
he wild,
he e
is
no
in o ma ion
as
o how hese ba s we e ained o eed on insec s
in
he
labo a o y.
Lacking his in o ma ion on insec -ea ing phyllos oma ids, i may be aluable
o desc ibe how some in es iga o s ha e ained espe ilionid and molossid
ba s o eed. The success
o
aining ba s o do wha he in es iga o wan s hem
o do depends in la ge measu e on he pa ience and skill
o
he aine , bu also
on he inclina ion
o
he ba o lea n. Some ba s equi e less aining o ea meal-
wo ms han o he s. The e a e se e al me hods, anging om holding aba
in
aglo ed hand and o e ing ood o simply allowing ba s o lea n o eed hem-
sel es om apile
o
nlealwo ms in adish.
Cons an ine (1952:397) placed mealwo ms on he cage loo hoping ha
specimens
o
Tada ida b asiliensis would ecognize hei u u e die a e hey
a e hei i s
ew
hand-o e ed wo ms. Racey (1970: 178) held he ba
in
his hand,
decapi a ed he mealwo m, and hen applied he isce a o he ba 's lips. The
ba 's jaws closed and he ba usually would chew he insec .
I
he ba did no
chew
in
hal aminu e, hen ad op
o
wa e was placed
in
he co ne
o
he ba 's
mou h. I he e was s ill no esponse, he mealwo nJ was squeezed
so
ha i s
con en s en e ed he ba 's mou h. A e he i s mealwo m was swallowed, he
ba 's nose was b ough in con ac wi h he insec s mo ing
in
adish
o
on he cage
Food Poisoning
Uwe Schmid (pe sonal communica ion) desc ibed some dea hs in his
Desmodus colony ha he a ibu ed o ood poisoning. The condi ion may
appea suddenly, e en a e ba s ha e been in cap i i y o se e a] yea s.
Schmid obse ed ha "ba s hung wi h s e ched legs om he cage op and e-
go ged blood (s omach was always comple ely illed) while u ina ing bloody
u ine." The cause, Schmid belie ed, was inges ed spoiled blood. Spoiled blood
does no appea o be dis as e ul o ampi es. Some ba s died; o he s eco e ed.
The animals ha su i ed d ank g ea amoun s
o
wa e .
The
dead ba s es ed
nega i e o abies.
So e Limbs
Swelling
o
he w is join s was obse ed
by
O
(1958), who hough ha
he high p o ein con en
o
ood, a e ayea
o
wo, may ha e caused his condi-
ion. Some ba s will de elop so es on he bo om
o
hei ee om es ing on a
ho izon al su ace such as he bo om
o
abox
o
bo le. To co ec his condi-
ion Ba bou and Da is (1969:246) sugges ed ixing asc een o pe mi he
ba s o hang head downwa d. Racey (1970) belie ed ha swelling
o
join s
in
noc ules migh
be
associa ed wi h he lack
o
exe cise. Some p e opodids and
possibly o he la ge ba s such
as
Vampy um, which no mally hang pendan and
ee, may su e om so e ee i no supplied wi h p ope oos ing su aces
such
as
ee b anches. Excessi ely cu ed claws may esul om con inued hang-
ing on wi e (Pye, 1967).
An excessi ely d y a mosphe e
in
alabo a o y may esul
in
d y and b i le
wing memb anes. Raising he ela i e humidi y and he applica ion
o
amois en-
ing skin lo ion
o
baby oil may emedy he ailmen .
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chi op e a and Mic ochi op e a. This condi ion
is
unsigh ly, bu he e do no
seem o be any o he ad e se e ec s. A e some ime
in
cap i i y on unsupple-
men ed die s, many ba s a e p one o weakening
o
he jaws and limbs, and
gene al lis lessness. S ua Fo nlula Liquid e ec s acomple e cu e
in
some
cases." O he s ha e no ed ha ba s ed on a i icial die s some imes lose hei
hai and ha mul iple i amins added o he die co ec ed he condi ion (Mohos,
1961; Da is and Luckens, 1966; Ba bou and Da is, 1969).
Rasweile (1975) men ioned co espondence om J. F ede ick Bell, who
ound ha ood con aine s mus be kep clean o p e en he loss
o
hai on ba s'
bellies. Rasweile (1975) also obse ed ha indi iduals
o
A ibeus li u a us,
unable o
ly
in small cages, we e o ced o c awl and ub agains he cage wi e,
which esul ed
in
he loss
o
en al hai . Uwe Schmid (pe sonal communica ion)
disco e ed ha la ge numbe s
o
icks
on
aba will cause lose
o
hai h ough
cons an sc a ching
by
he ba in
an
a emp o emo e he pa asi es. Tempo a y
elie nlay be p o ided
by
washing he a ec ed a ea
o
he ba wi h soap and luke-
wa m wa e .
114

MISCELLANEOUS
LABORATORY
TECHNIQUES
Anes hesia and Eu hanasia
The use
o
anes he ics and eu hanasics
in
labo a o y animals
is
o en necessa y
o humane and echnical easons. Muscle elaxan s
o
pa aly ics a e no anes-
he ics and should no be used alone o su gical es ain . They may be used
o su ge y in conjunc ion wi h d ugs known o p oduce adequa e analgesia.
E he
is
an anes he ic used
by
many biologis s. Mohos (1961) adminis e ed
e he om a egula d ipping bo le on o asmall, gauze-lined, wi e-mesh
baske co e ing he ba 's nose and mou h. P ecau ions mus be aken o a oid
hea ily soaking he gauze inasmuch
as
he ba may swallow he e he and die.
The dep h
o
anes hesia can be egula ed and eco e y
is
usually as and une en -
ul. Excessi e e he may be used o eu hanasia because ba s ha e ana ow ol-
e ance ange o his anes he ic.
Mohos (1961) ound ha nembu al, al hough easie o apply han e he , ga e
less sa is ac o y esul s. Pye (1967) p e e ed pen oba bi one sodium (nembu al)
ins ead
o
e he and used one olume
o
comme cial solu ion o nine olumes
o
10 pe cen e hyl alcohol injec ed in ape i oneally o induce su gical anes hesia,
Exe cise and ObesiTY
The e a e a ious opinions on he exe cise equi emen s
o
cap i e ba s.
O
(1958) ound ha An ozous, like many cap i e animals, ends o o e ea once
accus omed o cap i i y.
I
mo e han one ba
is
in acage,
i
is
unwise o limi
he amoun
o
ood o e ed because agg essi e ba s may consume mo e han hei
sha e. O e ea ing will lead o obesi y and one solu ion
is
egula exe cise. This
may be accomplished by pe mi ing he ba s o ly
in
a oom. Mohos (J96
J)
p o-
ided ala ge oom, 5
by
9me e s, o ba s o
ly
unhinde ed on he assump ion
ha exe cise would keep hem in good condi ion. The exe cise ligh was sa is-
ac o y o small numbe s
o
ba s. The p ac ice was la e discon inued due o
acciden s and high mo ali y when hund eds
o
ba s we e exe cised a once.
Howe e , when he exe cise was s opped, single and mul iple pyogenic in ec-
ions de eloped
in
many
o
he animals. Racey (1970) belie ed ha lying
is
no ap ac ical exe cise o la ge-scale ba husband y, bu did men ion condi-
ions he hough o be caused by he lack
o
exe cise. His se o ines o en de-
eloped so e w is join s and occasionally a al w is join in ec ions o Pseu-
domonas ae uginosa. The w is join s
o
his noc ulus also became s i and
in ec ed.
Wimsa and Gue ie e (1961) obse ed ha , al hough exe cise o ampi e
ba s migh be desi able o physiological easons, hei ba s appea ed heal hy and
igo ous wi hou exe cise. Glossophaga and Ca ollia a e able o exe cise wi hin
small cages because hey can ho e . Aldo
M.
Vou e (pe sonal communica ion)
epo ed ha he bes way he ound o keep cap i e ba s
in
good condi ion was
o place hei cage in ala ge oom and allow hem o lea e he cage and ly abou
in sea ch
o
ood as hey wished. Food was loca ed a aspeci ic spo on a
she1 ~
he ba s lea ned o ind he ood and always e u ned o hei cage o oos .
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE 1] 5
Cesa ian Sec ion
Adams and Bae (1966) desc ibed cesa ian sec ions used on Tada ida b asili-
ensis.
E he
was used. The hai
o
he abdomen was clipped and he a ea
dampened wi h one
pe
cen bensalkonium chlo ide. Amidline incision was
made h ough
he
skin and abdominal muscula u e wi h ascalpel, and he
igh
hom
o
he u e us wi hd awn.
The
u e ine wall was ca e ully incised and
he in an ba quickly wi hd awn; luids immedia ely we e sucked ou
o
i s mou h
a 30 o
50
nlillig ams
pe
kilog am
o
body weigh .
The e
is
conside able in-
di idual a ia ion in esponse and he ba should be ully a oused be o e in-
jec ion, and acons an body empe a u e
o
37° o
40°C
mus
be
main ained
a e wa ds o·.ensu e success ul anes hesia and eco e y. O e dosage wi h nem-
bu al may
be
used o kill ba s humanely (Pye, 1967).
To
anes he ize sa ely ampi e ba s, Dickson and
G een
(1970:43) in oduced
amix u e
o
oxygen (2 li e s
pe
minu e), ni ous oxide
(1
li e
pe
minu e), and
luo hane (halo hane, 2.5 uni s Fluo ec scale) in o he cages.
The
ba s we e
anes he ized in ou o i e nlinu es and eco e ed comple ely in h ee o ou
minu es. No ba s died
as
a esul
o
his me hod. Racey (1972) p e e ed he
halo hane/oxygen me hod
o
anes hesia o espe ilionid ba s.
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116
Ma king Ba s
Va ious me hods a e employed o ma k ba s o indi idual ecognI Ion.
Numbe ed me al bands a ached o o ea ms ha e been widely used. Ano he
me hod u ilizing di e en ly colo ed plas ic bands, such
as
used o cage bi ds,
is
sa is ac o y o asmall numbe
o
ba s. Some phyllos oma ids, such as Glos-
sophaga so icina, ha e apeculia s uc u e
o
he an eb achial memb ane ha
makes a achmen
o
bands o he o ea m di icul . Neona es, he o ea ms
o
which a e oo small o ake bands, may be numbe ed on he wing memb ane wi h
a a ooing o ceps (Racey, 1970). This me hod is use ul only o sho - e m ma k-
ing, because he small holes ou lining he nu nbe s heal o e and become ob-
li e a ed
in
a ew days. Howe e , he holes can be eopened wi h asui able
needle wi hou dange
o
in ec ion. When sca issue o ms, i is o en unpig-
men ed and he numbe can
be
ecognized o se e al weeks.
Rasweile (1975) ma ked ba s by means
o
spo s bleached on o hei u .
"The
bleaching solu ion consis s
o
six pe cen H202,Lady Clai ol P o ina o and
Lady Clai ol C emogenized
Hai
Ligh ene [Appendix 18]
in
he p opo ions
o
10:2:5 espec i ely.
The
H202and p o ina o a e mixed igo ously o abou
i e seconds in asmall ial.
The
ligh ene
is
hen added, and he mix u e agi-
a ed o ano he 20 seconds. The wo king solu ion e ains i s ac i i y o a
leas an hou a e p epa a ion. Immedia ely a e he applica ion
o
small
amoun s
o
he bleaching solu ion wi h ab ush, he ba s may be eleased back
in o hei cages." Up o
30
di e en numbe s a e possible by a ying he combi-
na ion and posi ion
o
he spo s on he back
and
head
o
aba . Se e al hund ed
Glossophaga and Ca ol/ia ha e been so ma ked, wi h only a ew mino cases
o
hai loss
in
adul s and skin damage
in
ju enile Ca ol/ia.
BIOLOGY OF
THE
PHYLLOSTOMATIDAE
1]7
wi h apipe e
o
wiped ou wi h aco on swab. The umbilical co d was clamped
wi h ahemos a , liga u es ied on ei he side
o
he hemos a , and he co d
cu . I was necessa y o pe o m he en i e p ocedu e apidly because any delay
mean inc eased mo ali y in he young.
AJ]
young, a e d ying, we e placed
in
an
incuba o (37°C) on asligh ly mois ened
owe1.
No men ion was made o whe he
any emales su i ed he ope a ion.
Ex ac ion
o
Ba
Milk
To de e mine some
o
he chemical and physical p ope ies
o
ba milk,
Huib eg se
(l966:551)
ela ed a echnique o emo ing milk om lac a ing
Lep onyc e is
sanbo ni and Tada ida b asiliensis as
oHows:
"The
animals we e
ligh ly anes he ized wi h sodium pen oba bi al (60
mg
20
mI)
....
Adose
o
0.03 o 0.10
ml
o
he anes he ic was ound o be sa is ac o y. This was admin-
is e ed p io o an injec ion
o
oxy ocic ho mone (Pi ochin, Pa ke-Da is)
o
less
han 0.1
m1.
The mammae we e ba hed wi h wa m wa e , and he milk exp essed
manually wi h humb and index inge . The ex uded d ople s we e collec ed
in a
smalJ
pipe e on a ubbe ube (a hemocy ome e pipe e se ed well)." A
small d op
o
10 pe cen o malin was added o p ese e e ige a ed samples.
Bleeding Ba s
Basic ex ac ion echniques ha e been de ised o ob ain blood om ba s, and
wo a e gene ally
used--one
om he hea , he o he om he eye. Ca diac
punc u e
is
he echnique mos widely used. Disposable needles and sy inges a e
ecomnlended. The app op ia e needle size depends on he size
o
he ba .
Fo
ba s
o
a e age size, Sudia e al. (1970) ecommended a25-guage, 3/8 o 5/8-
inch needle. A2-millili e sy inge
is
sa is ac o y. Asui able supply
o
needles a e
hepa inized by d awing aone pe cen solu ion
o
hepa in h ough hem and hen
allowing he needles o ai -d y. The punc u e si e should
be
cleaned and disin-
ec ed be o e inse ing he needle in o he hea .
Some in es iga o s p e e o ake blood om he o bi al sinus. Sudia e al.
(1970) ha e desc ibed his me hod. The anes he ized ba
is
held i mly
in
he
le hand, he hunlb exe ing su icien p essu e jus behind he eye o cause i o
bulge sligh ly. Amic osampling pipe e (hema oc i )
o
ei he a50
o
100 mi-
c oli e s
is
inse ed in o he pos e io o bi
o
he eye
(ca e u11y
pushing he eye-
ball o one side o a oid damaging
i )
and gen ly o a ed
so
ha he capilla ies
a e up u ed agains he bone and hus ini ia e he
low
o
blood. Once he
low
is
s a ed
i
is
necessa y o d aw he ube back sligh ly and incline i downwa d.
Usually he blood lows eely in o he ube, a imes so p o usely ha wo ubes
can be illed easily; some imes, howe e , i
is
necessa y o epea he o a ion a
ew
imes. The pipe e is hen discha ged in o a ube con aining ameasu ed
olume
o
diluen . The hema oc i ube
o
mic oli e ube may be hepa inized.
A50-mic oli e pipe e
wil1
ake up o 0.1
ml.
o
blood and a100-mic oli e
pipe e will ake up o 0.2 ml. blood ( he amoun usually aken om each ba ).
La ge amoun s
o
blood may be ob ained by holding he ba be ween he humb
and i s and second inge s o apply p essu e o he ho acic a ea. Appa en ly,
his aises he blood p essu e and inc eases he yield
o
blood om he o bi al
sinus. Ba s bled in his manne do no
appea
o
be
ha med se iously and such
bleeding may be con inued daily o up o 10 o
15
days
i
necessa y.
Sali a Collec ion
Dickson and G een (1970:41-43) equi ed he collec ion
o
sali a om
Desmodus and Diaemus. These biologis s de eloped an ingenious plas ic box o
he sa e es ain
o
hese ba s. The ampi es i s a e anes he ized as desc ibed
unde "Anes hesia." They hen a e placed in he sali a ion uni s so ha pilo-
ca pine can be adminis e ed o he buccal mucosa and he sali a collec ed as
desc ibed in de ail by he in es iga o s.
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U ine Collec ion
Bladde ca he e iza ion has been he echnique used o ob ain u ine samples
om small animals.
The
usual echniques equi e ei he con inuous anes hesia,
es ain ,
o
ex ensi e su ge y. Kan ho (1965) de ised ame hod
o
epea ed and
accu a e u ine collec ion om un es ained Myo is luci ugus (body weigh abou
7g ams).
He
w o e (p. 326):
"We
de eloped aca he e which could be passed
h ough hei ex emely well-de eloped u e h al spinc e wi h aminimum
o
subsequen i i a ion. Mo eo e , i p e en ed leakage and e en ion
o
u ine
in
he bladde , while ensu ing ee passage o pe mi collec ion
o
se ial samples
o e ex ended pe iods. Females we e animals
o
choice
as
he use
o
hese
ca he e s in males in ol ed ex ensi e su ge y."
The
ba s we e anes he ized wi h
e he . Mo e han
50
ba s ole a ed he p ocedu e and none ied o emo e he
ca he e .
The
ca he e s pe mi ed eedom
o
mo emen unde all condi ions and
emained unc ional o up o h ee days.
Ope an Condi ioning
o
Expe imen s
Ba s, unlike mos mammals, canno ope a e he con en ional gadge y
no mally used in expe imen al si ua ions wi h ei he
o
hei specialized limbs.
Beeche (1971) demons a ed ha Phyllos omus has a us could ope a e acon-
en ional pigeon key by ei he nosing
o
licking i . Schmid and G eenhall (1972)
ound ha Desmodus o undus would espond quickly o aining. Thei ba s
we e condi ioned o eed a an obse a ion able om I100 o 1300 hou s. A e
adish
o
blood was placed
on
he able, he cage was opened and acondi ioning
noise was made by sc aping wo o ceps oge he . Condi ioning could ake
up
o aweek, bu , when ba s we e ained, hey usually lew o he expe imen al
able wi hin i e minu es a e he noise in an icipa ion
o
ameal.
A e
he ba s
we e ained, a a ie y
o
small animals we e in oduced in o he cage so ha he
in es iga o s could obse e and pho og aph he ampi es as hey s alked, a acked
and ed on hei p ey. Thus, Schmid and G eenhall had o wai app oxima ely
i e minu es be o e hey we e able o obse e he eeding beha io ha o he -
wise migh ha e equi ed se e al hou s.
118
Pe sonnel P ecau ions
All pe sons handling cap i e ba s
o
in
con ac wi h hem should be awa e
o
he possible heal h isks in hei ou ine wo k. Simple hygiene p ecau ions should
be obse ed, such
as
washing he hands a e handling animals
o
immedia ely
on lea ing he animal acili ies.
In
my
labo a o ies, Iha e used wall-moun ed
dispense s con aining ei he inc u e
o
g een soap, ubbing alcohol,
o
o he
gene al an isep ics. Labo a o y coa s should be wo n o p o ec clo hing. Han-
dling ba s wi h ba e hands should be a oided
i
possible and lea he glo es should
be wo n as ap o ec ion agains bi es. Rubbe
o
disposable plas ic glo es should
be used while pe o ming ope a ions and au opsies. All cu s and ab asions
119
BIOLOGY OF
THE
PHYLLOSTOMATIDAE
Ba B ain
Remo al
Ba s ha ha e bi en people should be killed and sen o adiagnos ic labo a o y
o examina ion.
Fo
he pu poses
o
accu a e axonomic iden i ica ion, he
head p obably
is
he mos impo an pa
o
he animal o he mamnlalogis ,
whe eas b ain and o he body issues a e
o
pa amoun impo ance o he
epidemiologis . The con en ional labo a o y echniques used o b ain issue
emo al o abies diagnosis equen ly mu ila es he skin and skull, no only
making accu a e iden i ica ion di icul bu o en comple ely uining he specimen
o museum pu poses. To sol e his p oblem, G eenhall
(1965a)
de ised me hods
o issue emo al, wi h li le
o
no head and body damage, ha p o ed sa is ac-
o y o bo h mammalogis and epidemiologis . In he case
o
ex emely small ba s,
o
hose wi h specialized a achmen s be ween he ea s o unusual glandula
s uc u es on he head, i
is
unnecessa y o open he b ain case because su i-
cien b ain issue may be hypode mically wi hd awn h ough aneedle inse ed
in o he o amen magnum wi hou damage o he skull.
HUMAN
HEALTH
PROBLEMS
The Wo ld Heal h O ganiza ion (1973) epo ed ha inc easing numbe s and
kinds
o
animals a e now used
in
biomedical s udies. Zoologis s should be awa e
ha ba s may ca y diseases and, he e o e, may be ahaza d o human and animal
heal h. Jones in his chap e on economics and conse a ion ( his olume) has
discussed he diseases occu ing in wild phyllos oma ids ha may be ansmi ed
o humans and li es ock. Acomp ehensi e e iew
o
zoonoses and ba s was p e-
sen ed by Cons an ine (1970). The e
is
always he isk ha biologis s and ech-
nicians wo king wi h cap i e ba s may con ac
an
in ec ion om he pa asi es,
u ine, eces, skin, blood, and o he issues om hei own labo a o y animals.
Yunke (1964) has e iewed some
o
he common a h opod associa es
o
labo a-
o y animals ha a e haza dous o man. The public heal h impo ance o Neo-
opical ba s has been discussed in de ail by G eenhall (1964), Acha (1967),
Chalme s and Sco (1969), and Tamsi and Valdi ieso (1970).
Because li le
is
known abou he heal h
o
cap i e ba s, any in o ma ion
on he causes
o
dea h in he labo a o y
is
o
alue and au opsies should be pe -
o med when possible.

120
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should be cleansed immedia ely and agood i s aid ki should be easily accessi-
ble. Pa icula ca e should be gi en o eye p o ec ion and glasses
o
goggles
should be wo n when conduc ing pos mo em dissec ion on suspec ed abies
cases. Disposable pape ace masks se e as some p o ec ion agains he inhala-
ion
o
dus and spo es and a e ecommended o pe sons su e ing om alle gies
and espi a o y ailmen s. Whe he hese masks a e e ec i e agains in ec ious
ae osols
is
no known. S ic adhe ence o
aU
labo a o y sa e y ules should be
compulso y o s a
as
well as isi o s.
Clinical
Symp oms
o
In ec ions
Many people wo king wi h cap i e animals a e no awa e
o
he clinical signals
ha may be indica i e
o
an in ec ion acqui ed h ough exposu e o miscellane-
ous animal nla e ial. I he symp oms ail o be esol ed quickly, medical ad ice
should be ob ained. I in e
al.
(1972) lis ed he ollowing symp oms:
1)
alle gies
such as as hmalike symp oms
o
unning, i ching, and bu ning eyes, and skin
hype sensi i i y and i i a ion; 2) skin in ec ions; 3) espi a o y symp oms,
especial1y ape sis en cough, so e h oa ,
o
unning nose; 4) in luenza- ype
symp oms; 5) local in lanla ion and in ec ion, expecially
on
he hand
o
exposed
pa s
o
he body; 6) swelling
o
lymph nodes; 7) gene alized symp oms such as
e e , headache, e igo, dia hea, nausea, omi ing, and malaise.
Rabies
Rabies
is
pe haps he g ea es h ea o he chi op ologis because any ba may
con ac abies. Anumbe
o
phyl1os oma ids (no ably Phyllos omus, Glos-
sophaga, Ca ollia, A ibeus, Desmodus, Diaemus, and Diphylla) ha e been
ound o be posi i e o abies and hese ba s should be conside ed as po en-
ially dange ous labo a o y animals (Acha, )967).
Dickson and G een (1970) enlphasized ha ampi es om a eas ha a e
endemic o abies should be kep only
in
labo a o ies especially equipped o he
p o ec ion
o
he s a . The p ecau ions obse ed
in
hei labo a o y a e as ollows
(pp. 37-38):
"The
ba s a e housed
in
aqua an ine oom, which may be en e ed
only
by
pe sons immunised agains abies; gowns, masks and glo es mus be
wo n a all imes;
al1
was e ma e ials a e sealed
in
sacks be o e emo al om he
animal oom, and au ocla ed p io o disposal; he b ains
o
al1
ba s which die
a e emo ed and examined o abies i us."
Rabies in alabo a o y colony
o
ampi e
ba s.-
The
unexpec ed appea ance
o
abies in alabo a o y colony
o
ampi e ba s demons a es he impo ance
o
qua an ine measu es and he p ac ical alue
o
using cages in which he ba s
a e isible a all imes.
"The
ollowing inciden is
o
in e es because
i
was con-
a y o he usual pa e n
o
beha io
o
ampi es
in
cap i i y (Ho s and Lang-
wo hy, (1972:903).
"In
Janua y )970, 30 ampi e ba s (Desmodus o undus mu inus) we e col-
lec ed om
...
Mexico. The acclima ion
o
hese ba s o labo a o y condi ions
ollowed he pa e n desc ibed
by
Wimsa and Gue ie e (1961) and 20 ba s
su i ed his adjus men .
"On
29 Janua y 1971, 30 addi ional ampi e ba s we e ob ained in
...
Mexico
and added o he
15
emaining animals
in
he labo a o y colony. A e aweek
o
adjus men du ing which i e
o
he new animals succumbed, he colony was
s able and no unusual e en s occu ed un il he i s week
o
Ap il, wo mon hs
la e .
Up
o his ime he ba s, when dis u bed, no mally would bunch oge he ,
hiding in he da kes co ne
o
he cage. Howe e , beginning abou 1Ap il
1971, he e was in ense igh ing a he sligh es dis up i e s imulus, such as
swi ching
on
he ligh s, mo ing he cage, o sudden loud noises. These igh s
we e so in ense ha he en angled pai s would all
...
on o he loo , s ill
sc eaming and iciously bi ing each o he ." Mo ali y occu ed a a a e
o
abou
one ba pe day o abou wo weeks. Those ba s checked by he luo escen an i-
body me hod we e posi i e o abies.
Ho s and Langwo hy (1972:904) co ec ly ecommended ha "indi iduals
who main ain hese animals
in
cap i e colonies a e well ad ised o ake p ope
qua an ine p ecau ions wi h ecen ly cap u ed ampi es, les hey su e asimila
loss
o
aluable animals." Al hough Wimsa and Gue ie e (1961) desc ibed
hei ou ine ca e and main enance
o
ampi es, Icould no ind one men ion
o
he pa e n
o
acclima ion
o
ba s e e ed o by Ho s and Langwo hy
(1972:903).
Nonbi e abies in labo a o y animals and
echnicians.-Winkle
e
al.
(1972,
1973) epo ed an unusual ou b eak
o
nonbi e ansmi ed abies
in
alabo a o y
colony
o
wild ca ni o es and a a al case
o
nonbi e abies in alabo a o y wo ke
possibly caused by as ain
o
ba abies i us. Six y- ou animals died, including
39 ha had no known exposu e his o y. The human ic im had been accina ed
agains abies
13
yea s ea lie , bu had no de eloped demons able se um an i-
bodies. In es iga ion con i med ha di ec con ac ansmission did no occu and
sugges ed ha ai bo ne ba i us may ha e been esponsible. The human case
emphasizes he necessi y o biologis s and echnicians wo king wi h po en ially
abid animals o be immunized, ollowed by e i ica ion
o
demons able se um
an ibodies.
P eexposu e
immuniza ion.-The
bes p o ec ion agains abies an indi idual
can ha e
is
p eexposu e accina ion, as ecommended by he Uni ed S a es
Public Heal h Se ice (1974:16) and he Wo ld Heal h O ganiza ion (WHO)
Expe Commi ee on Rabies (1973:30): P eexposu e immuniza ion consis s
o
h ee injec ions
o
duck emb yo accine (DEY) spaced o e ape iod
o
se e al
weeks, ollowed by aboos e injec ion
o
accine one mon h la e , and las ly
by
he con i ma ion
o
an ibodies ( ha is, immuniza ion)
in
he se um
o
he ac-
cina ed indi idual. I nega i e, boos e doses should be epea ed un il an ibodies
become demons able.
Fu he
boos e injec ions should be gi en a in e als
o
one o h ee yea s as long as he pe son emains exposed. Some people ha e been
eluc an o accep he p eexposu e immunica ion because
o
he misconcep ion
ha he egimen equi ed ala ge numbe
o
daily injec ions.
Fi s aid ea men o bi e
wounds.-Acco ding
o he
WHO
Expe Commi -
ee on Rabies (1973:28) and Kaplan (1973:15-16), he
n10s
impo an i s aid
ea men o all bi e wounds and sc a ches
in
p e en ing possible abies in ec-
BIOLOGY OF
THE
PHYLLOSTOMATIDAE
121
ACKNOWLEDGMENTS
I
am
since ely g a e ul o he ollowing colleagues o p o iding ei he manu-
sc ip s ha a e in p ess
o
pe mi ing he inclusion
o
hei unpublished ob-
ion
is
he gen le washing and lushing
o
he wounds wi h soap and wa e , de e -
gen ,
o
wa e alone. Nex apply ei he 40 o 70 pe cen alcohol,
o
a 5 o 7pe
cen inc u e
o
aqueous solu ion
o
iodine,
o
0.1 pe cen qua e na y ammonium
compounds, which can kill abies i us on con ac wi hin one minu e. Alcoholic
be e ages
o
86 p oo
o
g ea e can be used in eme gencies. When soap has been
used o clean wounds, all aces
o
i should be emo ed be o e he applica ion
o
qua e na y ammonium compounds because soap neu alizes hei ac i i y. The
WHO Expe Commi ee on Rabies (1973:28) s a ed: HAl hough judicious use
o
concen a ed ni ic acid
in
punc u e wounds has i s ad oca es, he e is no
e idence ha his p oduc
is
mo e e ec i e han qua e na y ammonium com-
pounds
o
20%liquid soap solu ion."
Pos exposu e
ea men .-
The Uni ed S a es Public Heal h Se ice (1974:
1)
has ecommended he ollowing pos exposu e ea men agains abies.
"I
an
immunized pe son
is
bi en by a abid animal [ba ], he abies i us s imula es
apid p oduc ion
o
an ibodies because he indi idual has al eady been sensi-
ized
by
his p eexposu e accina ion. The e o e, an immunized pe son needs
only 1 o 6doses
o
DEV [Duck Emb yo Vaccine] e en a e being bi en by a
known abid animal, ins ead
o
he egimen
o
an ise um and up o
23
doses
o
DEY ecommended o an unimmunized pe son in he same si ua ion. Bu ,
mos impo an ly, ape son who has ecei ed p e-exposu e accina ion and e-
cei es 6doses
o
accine a e exposu e
is
conside ed signi ican ly be e p o-
ec ed han someone who ecei es only he ull pos -exposu e egimen
o
an i-
se um and accine."
The physician a ending abi e exposu e mus decide whe he an i abies ea -
men
is
indica ed and,
i
so, mus adminis e he mos e ec i e ea men a ail-
able o him.
I
se um
is
indica ed, he physician mus i s es o alle gies and
check he pa ien 's his o y o alle genic eac ion.
A
p esen he duck emb yo
accine
(DEV-Eli
Lilly &Co., Indianapolis, Indiana)
is
he mos widely used
in he Uni ed S a es. Newe and sa e accines such as he Wis a Vaccine, a e
being in es iga ed. Physicians a e s ongly u ged o check o he mos ecen
ecommenda ions a ei he
o
he ollowing wo Wo ld Heal h O ganiza ion
Rabies Re e ence Cen e s loca ed
in
he Uni ed S a es:
Uni ed S a es Public
Heal h
Se ice
Cen e
o Disease
Con ol
Bu eau
o
Epidemiology, Vi al Zoonoses Sec ion
A lan a,
Geo gia
30333
Telephone
(404) 633-3311, Ex ensions 3415
o
3683
A e 5p.m. (404) 633-2176
o
The
Wis a Ins i u e
Rabies Di ision
36 h S ee a
Sp uce
Philadelphia, Pennsyl ania 19104
Telephone
(215) EV 7-6700.
SPECIAL
PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
122
se a ions: James G. Dohe y,
H.
B ad o d House, Donna J. Howell, Rex o d
D. Lo d, John J. Rasweile , IV, Uwe Schmid , and Aldo M. Vou e. Ialso ex end
my
app ecia ion o Al ed
L.
Ga dne o his aluable c i icism. Iwish o hank
he Wo ld Heal h O ganiza ion o g an ing pe mission o ci e pa s
o
he Six h
Repo
o
he WHO Expe Commi ee on Rabies.
LITERATURE
CITED
ACHA,
P. N. 1967.
Epidemiology
o
pa aly ic
bo ine
abies
and
ba
abies.
Bol. O .
In .
Epizol.,
67:343-382.
ADAMS,
D.
B.,
AND
G.
M.
BAER.
1966.
Cesa ian
sec ion
and
a i icial
eeding
de ice
o
suckling
ba s. J.
Mamm.,
47:524-525.
BARBOUR,
R. W.,
AND
W. H.
DAVIS.
1969. Ba s
o
Ame ica.
Uni .
P ess
Ken ucky,
Lexing on,
286 pp.
BEECHER,
M. D. 1971.
Ope an
condi ioning
in
he
ba
Phyllos omus
has a us. J.
Expe .
Animal
Beha ., 16:219-223.
BRADBURY,
J.
W.
1970.
Ta ge
disc imina ion
by
he
echoloca ing
ba ,
Vampy lll11
spec um.
Exp.
Zool.,
173:23-46.
BUCKLAND-WRIGHT,
J. C.,
AND
J. D.
PYE.
1973.
Die a y
de iciency
in
ui
ba s.
In e nal.
Zoo
Yea book,
13:271-277.
BULLARD,
R. W.,
AND
S.
A.
SHUMAKE.
1973.
Food
empe a u e
p e e ence
esponse
o
Desmodus
o undus. J.
Mamm.,
54:299-302.
CHALMERS,
A.
W.,
AND
G.
R.
SCOTT.
1969.
Ecology
o
abies.
T op.
Animal
Heal h
P oduc ion,
1:33-55.
CONSTANTINE,
D.
G.
1952. A
p og am
o
main aining
he
ee ail
ba
in
cap i i y.
J.
Mamm.,33:395-397.
1970.
Ba s
in
ela ion
o
heal h,
wel a e
and
economy
o
man.
Pp. 319-449, in
Biology
o
ba s
(W.
A.
Wimsa ,
ed.),
Academic
P ess,
New
Yo k,
2:x +
1-477.
CRANDALL,
L. S. 1964.
The
ca e
and
managemen
o
wild
animals
in
cap i i y.
Uni .
Chicago
P ess,
Chicago,
x +761 pp.
DAVIS,
W.
H.,
AND
M. M.
LUCKENS,
1966.
Use
o
big
b own
ba s
(Ep esicus
[uscus) in
biomedical
esea ch.
Lab.
Animal
Ca e,
16:224-227.
DECOURSEY,
G.,
AND
P.
G.
DECOU~SEY.
1964.
Adap i e
aspec s
o
ac i i y
hy hms
in ba s.
BioI. Bull., 126:14-27.
DICKSON,
J. M.,
AND
D.
G.
GREEN.
1970.
The
ampi e
ba
(Desmodus
o undus): im-
p o ed
me hods
o
labo a o y
ca e
and
handl
ing.
Labo a o y
Animals
(London),
4:37-44.
DITMARS,
R. L. 1935.
Collec ing
ba s
in
T inidad.
Bull.
New
Yo k
Zool. Soc.,
38:213-
218.
1936. A
Vampy um
spec um
is
bo n.
Bull.
New
Yo k
Zool.
Soc., 39:162-163.
DITMARS,
R. L.,
AND
A. M.
GREENHALL.
1935.
The
ampi e
ba .
Zoologica,
19:53-76.
DUNN,
L.
H.
1933.
Obse a ions
on
he
ca ni o ous
habi s
o
he
spea -nosed
ba ,
Phyl-
los omushas a uspanamensisAlIen,
in
Panama.
J.
Mamm.,
14:188-199.
FLEMING,
T. H., E.
T.
HOOPER,
AND
D. E.
WILSON.
1972.
Th ee
Cen al
Ame ican
ba
com-
muni ies:
s uc u e,
ep oduc i e
cycles,
and
mo emen
pa e ns.
Ecology,
53:555-569.
FLORES
CRESPO,
R.,
R.
J.
BURNS,
AND
S.
B.
LINHART.
1971.
Compo amien o
del
ampi o
(Desmodus
oTundus)
du an e
su
alimen aci6n
en
ganado
bo ino
en
cau i e io.
Tecnica
Pecua ia
Mexico, 18:40-44.
GATES,
W.
H. 1936.
Keeping
ba s
in
cap i i y.
J.
Mamm.,
17:269-273.
1938a.
Raising
he
young
o
ed
ba s
on
an
a i icial die . J.
Mamm.,
19:461-
464.
1938h.
On
he
keeping
o
ba s
in
cap i i y.
P oc.
Louisiana
Acad.
Sci., 4:157-
158.
123
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
cen ime e s mul i i amin d ops (Vi-Pen a); six easpoons Ma ine P o ein Concen a e; h ee
easpoons bee ex ac . Add wa e o make
one
qua
and blend.
Then
add wo qua s
o
wa e and s i . Amoun su icien o app oxima ely 50 ba s.
16.
B onx
Zoo
Die
o
Phyllos omus has a us
House (1968: 141) used his o mula o
main ain
Phyllos omlls has a us: 50 pe cen
bananas; 30 pe cen g apes; 20 pe cen chopped mea ; i amin supplemen ;
one
easpoon
Mellin's
Food
(pe day).
The
daily amoun o e ed each ba was app oxima ely
35
g ams
ad
libi llm.
SPECIAL
PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
13.
A i icial
Die
o
Nec a i o ous
Ba s
This o mula was p o ided
by
Donna
J. Howell (pe sonal co espondence): 10 eggs; nine
ablespoons B ewe 's yeas ; nine ablespoons ins an p o ein (Alpine Ma ine);
one
able-
spoon bone meal;
18
capsules
amino
acids (Wolin's);
18
cubic cen ime e s mul i i amin
d ops (Poly-Vi e Wolin's);
18
ounces
(2
1
Acups) condensed
(no
swee ened) milk; h ee
pounds honey
o
s awbe y jam; wo small
ja s
"high-mea die " baby ood (s ained bee
o
chicken); h ee- ou hs bee ex ac (Di co Cul u e Medium);
18
ablespoons Supe
Hyd amin (Nion).
Mix he ing edien s well in blende , add six qua s wa e , and mix again. Makes one and
a
hal
gallons.
The
mix u e
is
ozen in pin con aine s and used as needed. Each
ja
mus
be shaken e y ho oughly be o e
i
is
pou ed in o ba eede s.
The
bee ex ac p o ides sal .
15.
B onx
Zoo
Die
o
F ugi o ous
Phyllos oma id
Ba s
House and Dohe y (1975) used his die o hei ugi o es: 24 bananas; one apple; one-
ou h pound g apes; 5ounces Zu/P eem P ima e Die (canned); ou ounces Zu/P eem
Feline Die (canned); six ablespoons Mellin's Food; six ablespoons Supe Hyd amin.
The
daily quan i y ed was app oxima ely
25
o
30
g ams pe ba
ad
libi um.
14.
Banana-based
Did
o
F llgi o olls
Phyllos oma id
Ba s
This die was aken om Rasweile (1976) and Rasweile and Ishiyama (1973:59): 700
g ams banana; 9.87 g ams whea ge m (pul e ized in aco ee mill); 12.90 g ams whole
milk powde ; 24.00 g ams calcium caseina e; 24.48 g ams suga ; 4.04 g ams p o ein
(Ge al);
1.71
g ams mine al supplemen ; 0.70 g ams i amin mix; 8.68 millili e s co n oil
mix u e.
The
bananas a e cu in o slices I-cen ime e hick and gen ly mixed wi h he o he in-
g edien s.
The
slices a e kep in ac so ha he ba s can hold and ca y away he pieces.
An
excess
o
ood
is
p o ided.
The
daily amoun o e ed each ba
is
app oxima ely
32
g ams
o
A ibells
li ll a lls and Phylloso mlls discolo , and abou 20 g ams o
Sw ni a
/ilium.
12.
A i icial
Die
o
Nec a i o olls
Phyllos oma id
Ba s
This o mula was gi en by Rasweile and de Bonilla (1972:661) and Rasweile (1973:
394): 700 millili e s ui base (peach nec a
o
gua a nec a ); 29.52 g ams ce eal (high
p o ein); 4.92 g ams whea ge m;
12.81
g ams milk powde ; 20.39 g ams calcium caseina e;
68.00 g ams suga ; 3.99 g ams p o ein (Ge al); 1.68 g ams mine al supplemen ; 0.69
g ams i amin mix; 9.00 millili e s co n oil mix u e; 300 millili e s wa e .
To
acili a e suspension in he liquid die , he ce eal and whea ge m a e educed
o
a
ine powde
by
passage h ough aco ee mill p io o hei mix u e wi h he
o he
d y com-
ponen s. The d y mix u e
can
be s o ed a
4°C
un il use.
The
composi ion
o
he co n oil
mix u e (polyoxye hylene so bi an mono-olea e as
an
emulsi ie and isopen ylace a e as a
la o ing agen ) can be s o ed a
4°C
un il use.
The
inal die consis ing
o
peach nec a ,
he powde ed p emix, and he co n oil mix u e
is
p epa ed eshly each day in
an
elec ic
blende and se ed o he ba s in shallow dishes.
The
amoun o e ed by Rasweile and de
Bonilla was app oxima ely 24 millili e s pe ba pe day.
130

BIOLOGY
OF
THE
PHYLLOSTOMATIOAE
131
J
7.
Die
o
F ugi o ous
and
Omni o ous
Plzyl/os oma id Ba s
Donna
J. Howell (pe sonal communica ion) ecommended his o mula o main aining
ugi o ous and omni o ous phyllos oma ids:
10
bananas ( e y ipe, chopped); h ee o
ou cups melon pieces (can aloupe
o
honeydew); eigh ounces co age cheese: one pound
hambu ge
o
one can dog ood; one- ou h ube bee ex ac ; one-hal Pe inal powde
(a ailable in pe s o es); wo eggs.
This should be ossed o mix.
o
else he i s ba a he dish will ea all he choice pieces.
Howe e , i should no be mixed in o amush
o
pas e because ba s will ge he mush on
hemsel es in hei en husiasm and compe i ion o he ood.
I
ba s a e ob iously p egnan ,
condensed milk and bone meal should be added.
The
o mula has p o ed sa is ac o y o
Phyllos omus
has a us,
P.
discolo , Ca ol/hi,
A ibeus,
and S u ni a.
18. P oduc s
Men ioned
in he Tex *
Bee
Ex ac -Di co
Labo a o ies, De oi , Michigan (Di co Cul u e Medium); Ge al-
P o ein--Lede le
Labo a o ies, Pea l Ri e , New Yo k; Ins an
P o ein--Alpine
Ma ine
Indus ies, New Bed o d, Massachuse s 02742;
Jeculin--Upjohn
Company, Kalamazoo.
Michigan 49001 (li e ex ac and i on); Lady Clai ol
P o ina o -Clai ol
Inco po a ed.
S am o d, Connec icu ; Lady Clai ol C emenogenized Hai
Ligh ene -CI
ai ol Inco po a ed,
S am o d, Connec icu ; Ma ine P o ein
Concen a e--Alpine
Ma ine Indus ies, New
Bed o d, Massachuse s 02742; Mellin's
Food-Consolida ed
Royal Chemical Company,
Chicago, Illinois 60610 (mal ose-dex in mix u e wi h added hiamin e ic glyce ophos-
pha e and po assium bica bona e);
Os e milk--Glaxo
labo a o ies, L d., G een o d,
Middlesex. England ( econs i u ed milk); Pe
D ops--Upjohn
Company. Kalamazoo,
Michigan 49001 (mul i i amins); Poly Vi e
D ops--Wolins
Company, Fa mingdale, New
Yo k (mul i i amins); S ua
Fo mula
Liquid-S ua
Company, Pasadena, Cali o nia
(mul i i amins); Supe Hyd amin
Powde -N
ion Co po a ion. Los Angeles, Cali o nia
90038; Tego
Solu ion--Hough
Hoseanson L d., Chapel S ee , Manches e J
9,
England;
The ag an--E.
R.
Squibb and Sons, New Yo k, New
Yo k
10022 (mul i i amins); Vi-
Pen a-Roche
Labo a o ies, Nu ley, New Je sey 07110; Zu/P eem
Die s--Ri iana
Food.
Inc., Hills Di ision, Topeka, Kansas 66601.
*Men ion
o
ade
names
o
p oduc s
does no imply
endo semen
by he U.S.
Go e nmen .
ECONOMICS
AND
CONSERVATION
CLYDE
JONES
His o ically, ba s ha e been conside ed
by
man
as
objec s
o
mys e y, supe s i-
ion, ea , and basically
as
indica o s
o
some unknown
o
e il signi icance. The
ole
o
ba s
in
he w i ings
o
ea ly na u alis s,
as
subjec s
o
ea ly a is s,
as
ob-
jec s
o
supe s i ion and e en wo ship, and
as
ing edien s
o
concoc ions o
a ied pu poses was summa ized in de ail by G. M. Allen (1939), who p epa ed
he i s gene al summa y on he biology
o
ba s.
In mode n imes, he e has been an inc easing awa eness
o
ba s o se e al
easons, mos ly because
o
he de elopmen
o
ools wi h which o ob ain
nume ous species o s udy, as well
as
an inc ease in knowledge
o
some diseases
o
ba s impo an o man. As a esul
o
hese and o he ac o s, he e has been a
g ea su ge
o
in e es and ac i i y
in
a ious s udies
o
ba s du ing he pas wo
decades. A
ew
o
he majo gene al wo ks on ba s wi hin his pe iod
o
ime in-
clude he summa ies p o ided by B osse (1966), Ba bou and Da is (1969),
Leen and No ick (1969), Slaugh e and Wal on (1970), and Wimsa (1970). As
an addi ional indica ion
o
cu en in e es
in
s udies
o
ba s, ou in e na ional
con e ences ha e been held since 1968, and annual con e ences ha e been held
in
he Uni ed S a es since 1970. Newsle e s abou ba s ha e appea ed du ing
ecen yea s
in
Aus alia, Eu ope, and he Uni ed S a es. Ade ailed e iew
o
he p oli e a ion
o
li e a u e on Chi op e a
in
he pas
ew
yea s was p o ided
by Ande son and Van Gelde (1970). In spi e
o
he a o emen ioned in e es s
and ac i i ies, much emains unknown wi h ega d o he biology
o
ba s. I seems
ha ce ain g oups, such as he Phyllos oma idae and some o he opical axa,
as well as special opics, such as he s a us
o
popula ions and he need o con-
se a ion' ha e been ela i ely neglec ed.
This epo a emp s o summa ize and discuss some
o
he b oad p oblems
conce ning he economic impo ance and conse a ion o he Phyllos oma idae,
and o e iew b ie ly some
o
he speci ic needs o conse a ion
o
hese ba s.
PROCEDURES
I
is
no wi hin he scope
o
his epo o p esen acomple e summa y
o
li e a-
u e con aining in o ma ion and no es on he aspec s
o
he biology
o
phyllo-
s oma ids. Re e ences a e p o ided ha con ain ei he addi ional in o ma ion
o
examples pe inen o he pu poses
o
his pape . In o ma ion was ob ained om
he li e a u e, om newsle e s and a ious epo s on ba s, and om co espond-
ence wi h esea che s who ei he wo ked on phyllos oma ids o conduc ed ield
s udies in a eas whe e hese ba s occu .
Da a
we e aken also om he iles
o
he ba -banding p og am
o
he Na ional Fish and Wildli e Labo a o y.
The scien i ic names used he ein a e
in
acco dance wi h he nomencla u e p o-
ided by Jones and Ca e ( his olume).
133
134
SPECIAL
PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
ECONOMICS
I seems impossible o a emp o de e mine meaning ul economic alues wi h
ega d o mos ba s; a leas , no p ecise de e mina ions will be made he ein. How-
e e , i
is
ele an o e iew b ie ly he ole
o
ba s wi hin ecosys ems, especial-
ly
wi h ega d o ela ionships be ween phyllos oma id ba s and ce ain hings
ha seem impo an o man.
Rela ionships wi h Plan s
Ba s
playa
ole
in
he na u al dispe sal
o
plan s. Membe s
o
he amily Phyl-
los oma idae a e he p inciple agen s
o
chi op e ocho y in he New Wo ld
opics. In gene al, mos dissemina ion
o
plan s by ba s esul s om seeds ha
a e d opped om he mou hs
o
animals ei he in ligh
o
in oos s. Rela i ely
ew
plan seeds a e passed h ough he diges i e ac s
o
ba s, al hough seeds
o
some plan s, such as Ficus, p obably a e sca e ed in his manne .
Chi op e ocho y
is
highly de eloped
in
ce ain amilies
o
plan s, especially
he Mo aceae, Palmae, Anaca diaceae, Sapo aceae, and Meliaceae. Howe e ,
his condi ion also
is
p esen in many o he amilies.
Fo
lis s
o
chi op e ocho -
ous plan s, cha ac e is ics
o
ba ui s, and discussions
o
he synd ome o dis-
pe sal
o
plan s by ba s, see he wo ks by Van de Pijl (1957, 1968, 1969, and
o he s). Sonle speci ic examples
o
synzooic ela ionships be ween plan s and
Phyllos omus discolo ,
P.
has a us, A ibeus jamaicensis,
A.
li u a us, and
Ca ollia pe spicilla a a e gi en
by
Van de Pijl (1957) and G eenhall (1956,
1965, 1966).
Phyllos oma ids a e he agen s
o
chi op e ophily
in
he Neo opical egion.
Acco ding o Van de Pijl (1969), ba lowe s occu mos ly in he gene a Musa,
Pa kia, Sonne a ia, Aga e, and Ca negiea, bu a e p esen also
in
a ious s ages
o
e olu iona y de elopmen
in
some o he axa. Flowe s pollina ed by ba s show
some specialized ai s; hey ha e an abundance
o
nec a and pollen and end o
open a nigh . The e
is
some e idence ha pollen
o
chi op e ophilous plan s ha e
mo e amino acids han do ela ed species pollina ed in o he ways (Howell,
1970). Because lowe ba s ely on lowe s o asupply
o
p o ein in he o m
o
pollen, mu ualis ic adap a ions a e exhibi ed by some pollina ing ba s.
Fo
ex-
ample, p ojec ions on he cu icula scales
o
hai s
o
some glossophagines seem-
ingly a e co ela ed wi h chi op e ophily (Howell, 1971). O he mo phological
adap a ions ela ed o eeding and ligh , as well as seasonal mo emen s
o
ba s
ha coincide wi h lowe ing
o
ce ain plan s, a e be e known han he a o e-
men ioned example.
The e
is
no doub ha phyllos oma id ba s a e impo an as agen s
o
dispe sal
o
seeds and pollen, a leas in alimi ed ange. Some bo anis s a e
o
he opinion
ha , among mammals, ba s a e he mos impo an dispe se s
o
seeds (Van de
Pijl, 1957). Recen analyses
o
ophic oles
o
ba s demons a e he ichness
o
he Neo opical egion wi h ega d o ugi o y and nec a i o y
o
he chi op-
e an auna (Wilson, 1973). The e a e simila i ies be ween he dis ibu ions
o
ce ain phyllos oma ids and chi op e ocho ous and chi op e ophilous plan s;
in ac , he anges
o
some plan s may esul om he ac ions
o
ba s on he e-
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE 135
p oduc ion
o
he plan s. Nume ous plan s wi h ypical ba lowe s and ba ui s
a e economically impo an o man ei he as sou ces
o
ood
o
o o namen al
pu poses.
Rela ionships wi h Insec s
Phyllos oma id ba s a e associa ed ecologically wi h an a ay
o
insec s.
Al-
hough usually e e ed o
as
ui ba s
o
ui -ea ing ba s, membe s
o
his amily
consume conside able quan i ies
o
insec s.
Fo
example, 33
o
143 s omachs
om a o al
o
11
axa
o
phyllos oma ids examined by A a a
e
al. (1967) con-
ained insec ma e ials. In o ma ion p o ided by Wilson (1973) e ealed he
ela i e impo ance
o
insec s
in
he die s
o
he gene a
o
phyllos oma ids.
Wilson also indica ed he need o addi ional in o ma ion
on
he ood habi s
o
ba s. Few da a a e a ailable o e alua ing he ole
o
phyllos oma ids wi h e-
ga d o he consump ion
o
insec s wi hin an ecosys em.
Fo
some limi ed in-
o ma ion on he impac
o
la ge colonies
o
insec -ea ing ba s on insec popula-
ions, see he a icles by Cock um (1969, 1970). Al hough almos impossible o
quan i y on he basis
o
cu en knowledge, i
is
ob ious ha he p eda o -p ey
ela ionships be ween phyllos oma id ba s and insec s a e impo an .
Rela ionships wi h Man
Phyllos oma id ba s a e conside ed occasionally as pes s
o
ui ees. The
possibili ies
o
hese ba s damaging ui ees we e implied in he discussion
o
he ela ionships be ween ba s and plan dispe sal and ep oduc ion. The e a e,
howe e ,
ew
epo s
o
se ious damage o ui ees by ba s in he New Wo ld
opics. Some da a on dis u bances o c op plan s by phyllos oma ids we e p e-
sen ed
by
G eenhalJ (1956, 1966). All
o
he a ailable da a indica e ha wha -
e e ha m phyllos oma ids do o he ui indus y
is
o
li le
o
no consequence,
excep o some isola ed ins ances whe e only limi ed damage is done. Phyl-
los oma ids c ea e some annoyances o man because
o
ui consump ion
in
ga dens and homes and co esponding deposi ion
o
ui pulp and o he
d opped ma e ials.
Phyllos oma ids also c ea e anuisance by oos ing
in
man-made s uc u es.
Ba s commonly occu in a ics, walls, and be ween laye s
o
ha ch
o
buildings
h oughou he Neo opical a ea. People usually objec o noises made by ei he
ocaliza ions o mo emen s
o
he animals, odo s and s ains om u ine and ecal
ma e ials, and ejec ed ood pa icles ha equen ly accumula e below oos ing
si es. In addi ion, ammonia gas may accumula e in guano deposi s and cause un-
pleasan ness o man.
Fo
an indica ion
o
he ela ionship be ween ammonia and
Mac o us cali o nicus, see he epo by Mi chell (1963). Human ood may be-
come con amina ed by d oppings om ba s ha inhabi buildings o om in-
sec s and a h opods ha li e in guano deposi s.
Guano deposi ed by ba s has been used
as
e ilize , especially in he ea ly
s ages
o
he de elopmen
o
he e ilize indus y. Excep o some igu es o
guano p oduc ion and mining in no he n Mexico and he sou hwes e n Uni ed
S a es (Hu chinson, 1950), good da a on mode n p oduc ion and use a e no
136
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TABLE
I.-Occu ence
o
bac e ial diseases in phyllos oma ids.
Bac e ia
Disease
Hos
species
Coun y
Salmonella Salmonellosis Glossophaga so icina
Panama
A ibeus
li ll a us
Colombia
S u ni a lilium
Colombia
Ba onella and Ba onellos
is
Ca ollia pe spicilla ll B azil
G ahamella
Desmodus
o undus
Pe u
a ailable. Guano
is
s ill mined, especially
in
some a eas in Mexico, and some
o
he la ge ca es appa en ly yield wo hwhile amoun s. Howe e , he impo ance
o
ba guano
as
e ilize is no wha i was p e iously because
o
he deple ion o
supplies and sou ces,
as
well
as
he de elopmen
o
o he comme cial e ilize s
in
ecen yea s. The ole
o
phyllos oma ids as p oduce s
o
guano in su icien
quan i ies o comme cial exploi a ion
is
poo ly known.
Membe s
o
he Phyllos oma idae also a e conside ed pes s because hey
ha bo ec opa asi es ha cause some conce n o humans. Howe e , no u he
discussion
is
wa an ed he ein because his opic is deal wi h in de ail by Webb
and Loomis ( his olume).
Phyllos oma ids, like o he ba s, a e o en conside ed as pes s
by
man
in
con-
nec ion wi h diseases ha may be ca ied
by
he animals and possibly ansmi ed
o man. Alis
o
he bac e ial, myco ic, and p o ozoan diseases known o occu
in
wild phyllos oma ids
is
p esen ed in Tables 1 o
3.
Known occu ences
o
i uses
in
hese ba s a e summa ized
in
Tables 4and 5. Mos
o
hese da a a e
om Cons an ine (1970), who also ga e de ailed discussions
o
each disease and
i us, including in o ma ion on how hese diseases a e mani es ed
in
humans.
Fo
addi ional in o ma ion on pa hogens
in
he Neo opical a ea, wi h special
emphasis on Pue o Rico, see he pape
by
Tamsi and Valdi ieso (1970).
The epo ed occu ences o abies i us in ec ions in wild phyllos oma ids a e
summa ized
in
Table
5.
Rabies
in
ampi e ba s
is
mo e common han
in
all o he
membe s
o
he amily. Acco ding o Cons an ine (1970), epo ed dea hs
o
humans om abies ansmi ed
by
ampi e ba s a e ela i ely insigni ican
causes
o
human mo ali y (Table 6). Howe e , abies ansmi ed
by
ampi e
ba s o li es ock
is
an impo an conce n because his disease
is
amajo cause
o
mo ali y
in
ca le
in
La in Ame ica. Rabies
in
ca le has been epo ed in all
La in Ame ican coun ies excep Chile and U uguay. Es ima es
o
annual ca le
mo ali y a y conside ably.
Fo
example, Cons an ine (] 970) p o ided asum-
ma y
o
da a on es ima ed ca le losses ha o aled hal amillion head (47.6
million dolla s) in ]966 and wo million head (100 million dolla s)
in
1969. The
magni ude
o
his p oblem
is
e lec ed in he ash
o
in es iga ions in ecen
yea s conce ned wi h he con ol
o
ei he abies
o
ampi e ba s. "Vaccina ion
has been he mos e ec i ely applied me hod
o
comba ing he p oblem in li e-
s ock" (Cons an ine, ]970). Some me hods o con olling ampi e ba s ha e
esul ed om s udies
o
ce ain biological aspec s
o
hese animals
in
p og ams
ope a ed by he Food and Ag icul u e O ganiza ion
o
he Uni ed Na ions and he

BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
TABLE
2.-0ccll ence
o
mycoTic diseases in
phyllos onw ids.
137
Fungus
His oplasma
Blas omyces
Sc pllia iopsis
C yp ococcus
Candida
To ulopsis
T ichophYTon,
Mic ospo um,
T icho~po on
Disease
His oplasmosis
Blas omycosis
Scopula iopsos
is
C yp ococcosis
Candidiasis
To ulopsosis
Supe icial
mycoses
Hos
species
Lep onyc e is
sanbo ni
Desmodus
O llndus
Phyllos omus
discolo
A ibeusjamaicensis
Ca ollia pe spicilla a
Gl ssophaga so icina
L nchophylla
hus a
Loncho hina
au i a
Mic onycle is megalo is
Tona ia bidens
Phyllos omlls has a us
An u a
geo oyi
A ibeus
li u a us
Gl ssophaga so icina
Ca o"
ia
pe spicilla a
A ibeus
lilU a us
Desmodus
o undus
Ca llia pe spicilla a
Desmodus
o undus
Leplonyc e is
sanb ni
Ca ollia pe spicilla a
Desmodus
o undus
Lep onYCle is
sanbo ni
Lep onycTe is
sanbo ni
Gl ssophaga so icina
Coun y
Uni ed S a es
Mexico, Panama,
Colombia, T inidad
El
Sal ado ,
Panama
El
Sal ado ,
Panama
Panama, Colombia,
T inidad
Panama, Colombia.
T inidad
Panama
Panama
Panama
Panama
T inidad
T inidad
Colombia
M~xico
Mexico
Mexico
Mexico
Colombia
Colombia
Uni ed S a es
Colombia
Colombia
Uni ed S a es
Uni ed S a es
Colombia
Den e Wildli e Resea ch Cen e
o
he Uni ed S a es Fish and Wildli e Se ice.
The a emp o educe abies
in
ca le by he con ol o ampi e ba s is acomplex
and some imes con o e sial issue.
Fo
addi ional in o ma ion, consul he epo s
by
Cons an ine (1970), Schmid
e
ale
(1970), Rosen hal (1972), Yos i e
al.
(1971); Mendez (1971), G eenhall (1971, 1972), Tamsi and Valdi ieso (1970),
Thompson e
al.
(1972), and Tu ne (1975).
Ba s a e ex emely use ul o man
as
objec s
o
esea ch. Some indica ions
o
he inc easing awa eness
o
his impo ance a e implici
in
he in oduc o y
ema ks wi h ega d o he p oli e a ion
o
li e a u e on hese mammals in ecen
yea s.
Ba s a e used in se e al aspec s
o
medical esea ch, especially accine de-
elopmen , epidemiological s udies, mechanisms
o
disease esis ance, aging,
and he mo egula ion. T ansillumina ion o ba wings
is
acon enien way o
make g oss and mic oscopic obse a ions o na u al and expe imen al physiologi-
cal phenomena,
as
weI)
as
o pa hological s udies and cul u ing
o
o ganisms
in
animal issues.
138
SPECIAL
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TABLE
3.-0ccll ence
o
p o ozoan
diseases in
phyllos oma ids.
P o ozoa
Disease
Hosl species
Coun y
Ba s a e impo an
in
space biology o s udies dealing wi h awide a ay
o
ole ances
o
en i onmen al ex emes and s esses. They also a e use ul
in
nume ous in es iga ions
o
ae odynamics and ela ed opics.
S udies
in
ba echoloca ion a e use ul o many pu poses, as documen ed in
some de ail
by
G i in (1958). Some in e es ing and sophis ica ed s udies
o
ul asonic o ien a ion in ba s ha e been conduc ed
in
ecen yea s.
Fo
some ex-
TABLE
4.-0ccu ence
o
i uses in
phyllos oma ids.
Hos species
Coun y
Colombia
Panama
B azil
B azil
B azil
B azil
B azil
B azil
B azil
B azil
B azil
B azil
T inidad
T inidad
B azil
B azil
B azil
B azil
B azil
B azil
B azil
B azil
Colombia
Colombia
Colombia
Colombia. Mexico
Colombia
Colombia
Colombia
Ca ollia pe spicilla a
Phyllos omus
discolo
Phyllos omus
has a us
Desmodlls
o undus
Glossophaga so icina
A ibeus
li u a us
A ibells li u a us
G oup AA bo i uses
A ibellsjamaice/lsis
Ca ol/ia pe spicil/ll a
A ibeus
li u a us
Ca ol/ia pe spicil/a({[
A ibellsjamaicensis
G oup
BA bo i uses
Rhinophyl/a
pumilio
Glossophaga so icina
Miscellaneous A bo i uses
A ibeus
li u a lls
A ibeus
li u a us
A ibeus
jamaicensi,
Ca oll
ia
pe spicil/a a
A ibeus
li u a lls
A ibeus
li u ll liS
A ihellsjll/l1llil'ensi.
A
ibe
liS
lliginosiis
A ibeus
liW a lls
A ibeus
jamaicensis
A ibells
li u a us
A ibells
jamaicensis
Phyllos omus
has ll us
Vampy ops
hel/e i
A ibellsjamaicensis
Toxoplasmosis
T ypanosomiasis
Toxoplasma
T ypanosoma
Eas e n equine
encephali is
Venezuelan equine
encephali is
Mucambo /
Vi us
I apo anga
Ca apa a
Ju ona
U inga
Taca ibe
Sain Louis
encephali is
Yellow e e
Ca u
Tacaiuma
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
TABLE
5.-0ccu ence
o
abies in phyllos oma ids.
139
Hos species
Mac o us cali o nicus
Phyllos omus discolo
Phyllos omus has a us
Glossophaga so icina
Lep onyc e is ni alis
Ca ollia pe spicilla a
U ode ma
biloba um
A ibeus
sp.
A ibeusjamaicensis
A ibeus
li u a us
Desmodlls o llndus
Diaemus
youngii
Diphylla ecallda a
Coun y
Uni ed S a es. Mexico
B i ish Hondu as. Gua emala
B azil
Mexico
Mexico
T inidad. Colombia
Panama
Mexico
Panama, T inidad, B azil
B i ish Hondu as, Gua emala, Mexico,
T inidad, B azil
B i ish Hondu as. Gua emala. Mexico.
T inidad. B azil
T inidad
B azil
amples, no e he s udies
by
Pollak e
ale
(1972), Simmons (1970), Simmons and
Howell (1971), Howell and Pylka (1972), and Simmons e al. (1972). Ba s may
se e
as
impo an bioholog aphic models in he u u e (G eguss, 1968).
In addi ion o he a o emen ioned use ulness in esea ch, ba s a e impo an
esea ch objec s o o he biologis s.
Fo
example, he e s ill
is
need o , and in-
e es in, basic s udies
o
ecology, li e his o y, dis ibu ion, mo phology, and he
like. The axonomy
o
ba s
is
s ill an in iguing opic, and conside able in e es
has de eloped
in
s udies
o
hei beha io and popula ion dynamics. This may
become amo e ac i e esea ch a ea
in
he u u e, especially wi h ega d o s udies
o
mechanisms pe mi ing ba s o su i e
in
ex emes
o
popula ion conges ion.
CONSERVATION
Needs o conse a ion
o
Ame ican ba s in gene al we e ecognized by
Man ille (1962), Da is (1967), and Cock um (1969, 1970), and p esen ed
in
hei pleas o conse a ion. Addi ional commen s on needs o ba conse a ion
we e p esen ed by Ba bou and Da is (1969), Gould (1970), G eenhall (1973),
and Findley (1973). Phyllos oma ids,
as
well as some o he ba s, ha e been gi en
TABLE
6.-Some
epo ed incidences
o
abies ansmi ed o
humans
by
ampi e ba s.
Dea hs
Yea s
Coun y
89 1925-1937
T inidad
31
1951-196] Mexico
17
1953-1961 Guyana
]1960 Boli ia
81960 B azil
51965 A gen ina
140
SPECIAL
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TABLE
7.-1njo ma ion
on
phyllos omll idsj om
su eys
oj
esea che s.
Coun y
Da es
Genus
S a us
o
popula ion
some a en ion
in
he popula and conse a ion o ien ed wo ks by Rood (1971
),
Cu y-Lindahl (1972), and No ick and Dale (1973).
Awa eness and conce n o ba conse a ion has been exp essed by nume ous
spelunke s and he Na ional Speleological Socie y.
Fo
addi ional in o ma ion
and ecommenda ions
o
h~s
o ganiza ion, see he
pape
by Moh (1972).
Some conce ns o ba popula ions in he New Wo ld ha e been exp essed by
ba esea che s.
Fo
example, esolu ions we e de eloped du ing he Thi d
Symposium on Ba Resea ch, held in San Diego, Cali o nia, on 24-25 No embe
1972 wi h some guidelines o ba conse a ion. These ac ions we e based
in pa
on
a ecen su ey ha e ealed educed popula ions
o
22 species
o
ba s
in
he Uni ed S a es, including h ee species
o
phyllos oma ids (Jones, 1971).
In
connec ion wi h he a o emen ioned su ey, limi ed da a we e accu-
mula ed on he popula ion s a us
o
phyllos oma ids a se e al places in La in
Ame ica.
O
mo e han 100 eques s o in o ma ion sen o ba esea che s, only
16
o
hose e u ned included in o ma ion on phyllos oma ids. Pe inen in o ma-
ion om hese eplies
is
summa ized in Table
7.
All epo s on phyllos oma ids
in he Uni ed S a es indica e declining popula ions; epo s o o he a eas mos ly
e eal inc easing popula ions, excep o one epo
o
dec easing popula ions
and wo epo s
o
s able popula ions. Howe e , mos esponden s s a ed ha
popula ions
o
Desmodus we e ac ually educed
o
absen
in
local a eas due o
e adica ion
o
con ol measu es.
Fo
aweal h
o
in o ma ion on popula ions
o
ampi e ba s, see he wo k
by
Tu ne
(1975).
AU
epo s included lis s
o
easons o popula ion changes. Inc eased popu-
la ions
o
ba s we e associa ed wi h inc eased cul i a ion
o
ui c ops and he
Declining
Declining
Declining
Declining
Declining
Inc easing
Inc easing
Inc easing
Inc easing
Inc easing
Inc easing
Dec easing
S able
Inc easing
S able
Inc easing
Desmodlls
Lep onyc e is,
Choe onyc e is
Lep onyc e is
Lep onyc e is,
Choe onyc e is
Mac o us
Mlic o u.
Glossophllga
Desmodlls
Desmod/l.
Desmodlls
Desmodlls, Phyllos omlls,
A ibells, VlImpy ops,
S ll ni a, U ode ma,
Glossophaga
Glossophaga, Ca ollia
Desmodlls
A ibells, Ca ol/ill,
Phyl/os omlls
Glossoplwga
1966-1967
1963-1970
1963-]
970
J965-1970
1969-1970
]953-1963
1965-1971
1967-1971
1962-1969
1960-1970
1965-]970
1961-]
965
J
96]
-1971
1968-1971
1969-197]
1957-1971
Cos a Rica
Colombia
Colombia
Colombia
A gen ina
T inidad
Uni ed S a es
Uni ed S a es
Uni ed S a es
Uni ed S a es
Uni ed S a es
Mexico
Mexico
Mexico
Mexico
Cos a Rica
Ex e nal
In e nal
Na u e
o
s udy
SPECIAL
PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
X
X
X
X
X
X
X
X
x
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
X
TABLE
I.-B ains
o
phyl!osTomaTid species sTudied.
148
Species
Phyllos oma inae
LoncllO hilla au i a
Tomes
Mac ophyl!u11l l1U1c ophyl!um (Schinz)
Mac oTus cali o nicus Bai d
Mic ollycTe is hi sl/Ta (Pe e s)
Mic onycTe is mega/oTis (G ay)
Mic onycTe is
minll a
(Ge ais)
Mic onycTe is
nice o i
Sanbo n
Mic ollVCTe is schmidTo um
Sanbo n
Mimoll'c ellll/a llm
(E.
Geo oy S .-Hilai e)
Phyllode ma
s ellops Pe e s
Phyllos om
liS
disc%
(W,agne )
Phyl!os omlls e10nga lls (E. Geo oy S .-Hilai e)
Phyl/oswml/s
has mlls (Pallas)
TOlla ia
hidens
(Spix)
TOllaTia
nica agl/lle
Goodwin
T achops ci /lOslls (Spix)
Vampy llm
spec um
(Linnaeus)
Glossophaginae
AI/ou a
geo oyi
G ay
Choe onisclIs
godmalli
(Thomas)
Choe onisclIs in e medills (Allen and
Chapman)
C/1Oc ollycTe is mexiClInli Tschudi
G/ossop/wgll a/ ico/a Da is
G/ossophaga
commissa isi
Ga dne
G/ossop/lliga so icina (Pallas)
Hy/ollyc e is IInde woodi
Thomas
LepTonyc e is . (lIlhO lli Ho meis e
Lic!1onyc e is ohscu a
Thomas
Lonchophyl!lI ohlls a Mille
MOIlOphyllll.
ed
mlllli Leach
Ca ol
Ii
inae
Ca ol!
ia
pe spicillaTa (Linnaeus)
Ca ol/ill sl/h ll a (Hahn)
Rhinophylla
pllmi/io
Pe e s
S enode minae
Amc ida
cell ll io
G ay
A Tiheus
(/~TCCUS
Ande sen
A iheus
cinc eus (Ge ais)
A Tiheus hi sl/TIIS Ande sen
A iheus
inopill({[lIS Da is and
Ca e
A Tihellsjamaiccnsis
Leach
A ihells /iTl/ mliS (Ol e s)
A iheus
p/llIeo is (Mille )
A ihi'us
To/ ecu~'
(Saussu e)
A ihells I a soni
Thomas
Cell u io
senex
G ay
Chi ode ma
sa/ ini
Dobson
Chi ode ma
1.'il/os//m Pe e s

BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
TABLE
I.-Con inued.
Ec ophy//a
l7U1cconne//i
Thomas
Enchis henes
ha ii
(Thomas)
S enode ma
Lljwn Desma es
S ll ni a
lilillm
(E.
Geo oy S .-Hilai e)
S ll ni a
Illdo ici
An hony
S ll ni a
mo dax
(Goodwin)
U ode ma
hiloha llm
Pe e s
U ode l7ll1
magni os llm
Da is
Vampy essa
nymphaea
Thomas
VlImpy essa
pllsil/a (Wagne )
Val11py odes
ca accioloi
(Thomas)
Vampy ops
do salis
Thomas
Vampy ops
he//e i
Pe e s
Val11py ops injilsclis Pe e s
Vampy ops
i1 a lls (Pe e s)
Phyllonyc e inae
B achyphy//a
ca e na llm
G ay
E ophy//a
homhi ons
(Mille )
Desmodon inae
Desl110dllS
llndliS
(E.
Geo oy S .-Hilai e)
Diaemlls
yOlillgii (Jen ink)
Diphy//a
ecallda ({ Spix
x
X
X
X
X
X
X
X
X
X
X
X
X
X
X
x
X
X
X
X
x
X
X
X
x
X
149
METHODS
AND
MATERIALS
Ba s we e mis -ne ed om na u aJ popula ions
o
we e collec ed by hand
om oos ing si es. B ains we e p epa ed by emo ing he head immedia ely
a e he specimen was killed and chipping away he pa ie al egion
o
he skull
case o expose he b ain. Fixa ion wi h 10 pe cen o malin was allowed o p o-
ceed o se e al weeks. To acili a e handling, ixed b ains we e s o ed
in
70 pe
cen e hanol. A ew species a e ep esen ed
by
specimens p ese ed o iginally
in
alcohol ha we e collec ed
by
o he wo ke s. B ains om he la e o en showed
a ying deg ees
o
in e naJ de e io a ion, and
in
some cases hei alue was
limi ed o ex e nal s udy only.
The skull and meninges we e emo ed ca e ully om each b ain. Then, he
a ailable se ies
o
b ains om each species was examined o de e mine sub-
jec i ely a" ypical" specimen o each species. These b ains we e pho og aphed
om do sal, en al, and la e al iews. Such pho og aphs se ed
as
acon-
enien basis o compa ison; howe e , inal judgmen s we e made om obse a-
ions on he specimen.
Ce ain species we e selec ed o de ailed examina ion o in e nal ana omy.
Fo
hese his ological p epa a ions, species we e selec ed ha we e:
1)
ep esen a-
i e
o
each majo ype
o
ex e nal ana omy; 2) ep esen a i e
o
each majo axo-
nomic uni ; and 3)
o
ques ionable phylogene ic posi ion (such
as
B achyphylla,
which has been assigned by pas wo ke s o one
o
se e al di e en sub amilies).
B ains o his ological s udy we e in il a ed wi h agum a abic solu ion (Huma-
son, 1967) and ozen sec ions we e made
in
he c oss-sec ional plane a 18
mic ons. E e y second sec ion ( hi d sec ion on some la ge species) was collec ed
and moun ed
in
s ep ashion on mic oscope slides. Sec ions we e s ained wi h
"LuxoJ" as blue MBSN (Ma heson Coleman and Bell) o demons a e myeli-
na ed a eas and coun e s ained wi h c esyl iole ace a e (Ma heson Coleman and
Bell) o ou line concen a ions
o
cells
in
disc e e nuclei. S aining was by amodi-
ica ion
o
echniques desc ibed
by
D u y and Walling on (1967). Speci ics o
p epa a ion
o
sec ions was as ollows:
1)
sec ions we e dehyd a ed in 95 pe cen
e hanol; 2) s ained in Luxol as blue (see McDaniel, 1973, appendix A) o
ou o eigh hou s, a 50 deg ees Cen ig ade; 3) hyd a ed h ough an e hanol
g adien o dis illed wa e ; 4) dipped in o 0.05 pe cen li hium ca bona e a
one o h ee deg ees Cen ig ade o 15 o
20
seconds (solu ion mus be eshly
p epa ed daily); 5) di e en ia ed
in
70 pe cen e hanol un il he e was aclea
dis inc ion be ween myelina ed (blue) and nonmyelina ed issues (clea ); 6) hy-
d a ed h ough
an
e hanol g adien o dis illed wa e ; 7) coun e s ained
in
c esyl
iole ace a e (see McDaniel, 1973 appendix A) o i e. o
10
minu es; 8)
coun e s ain di e en ia ed
in
95 pe cen e hanol o which a ew d ops
o
ace ic
acid had been added; 9) sec ions dehyd a ed in 100 pe cen e hanol, clea ed
in xylene, and moun ed
in
Pe moun . S eps 4and 5may
be
epea ed i
necessa y. This s aining p ocedu e yields sec ions wi h deeply blue-s ained
myelina ed a eas ( anging om la ge ac s o indi idual axons) and pu plish- ed
cell bodies embedded in apale iole ma ix.
EXTERNAL
MORPHOLOGY
OF
THE
BRAIN
Gene al Desc ip ion
The ce eb al hemisphe es
o
phyllos oma id ba s a e ela i ely smoo h and
wi hou con olu ions (Fig. 1). A
ew
dep essions, which a e usually shallow, a e
ound in mos species. In he pas , hese we e con enien ly ega ded as ue sulci,
bu mo e ecen ly, hey ha e been e med ossae
o
issu elike dep essions esul -
ing om he p esence
o
adjacen osseous and ascula ea u es (Schneide , 1957;
Henson, 1970). My compa a i e his ological examina ion
o
ase ies
o
phyl-
los oma id b ains (McDaniel, 1973) e ealed ha in some cases hese dep essions
a e no shallow, and appa en ly ep esen mo e han acon o mi y
o
b ain o skull
( o an ex eme example, see he cingula e sulcus o Phyllos omus has a us).
The e a e no co ical maps published o chi op e an b ains; he e o e, he e a e
no da a conce ning unc ional in e p e a ion
o
he sulci
o
pseudosulci, and gy i
o
pseudogy i. In he absence
o
a unc ional in e p e a ion, he ollowing e mi-
nology
is
based on mo phological simila i y a he han on ac ual,
o
e en as-
sumed, homology wi h b ains
o
highe mammals. Homology wi hin he amily
Phyllos oma idae
is
assumed, and hope ully will be subs an ia ed expe imen ally
in he u u e.
Wi hin he Phyllos oma idae, h ee sulci a e consis en ly well de eloped: he
in e hemisphe ic sulcus, which sepa a es he igh and le ce eb al hemisphe es;
he an e io hinal issu e, which sepa a es he ol ac o y bulbs om he main
mass
o
he ce eb um; and he hippocampal sulcus, which
is
en ally loca ed.
O
he emaining sulci, he mos consis en ly p esen
is
one ha di ides he ce e-
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51
b al su ace in o an e io and pos e io po ions. This issu e is simila o, bu
no homologous wi h, he cen al sulcus (Syl ian sulcus)
o
highe mammals.
In his pape i will be e e ed o
as
he pseudocen al sulcus. In some cases,
ano he issu e
is
de eloped somewha an e io o he pseudocen al sulcus.
The a ious lobes
o
he ce eb al co ex a e ela i ely unob usi e and
in
mos cases do no show a endency o bulge
in
he manne ypical
o
highe mam-
mals. All phyllos oma ids ha e well-de eloped ol ac o y bulbs on he os al
end
o
he ce eb al hemisphe es. In addi ion, caudad o he pseudocen al sulcus,
apseudo empo al lobe p ojec s en ally
o
somewha pos e io ly om he body
o
he ce eb um.
The diencephalon (Fig.
1)
is
exposed only along i s en al su ace. Apo ion
o
he base
o
he halamus
is
exposed an e io o he op ic chiasma, and he hy-
po halamus is exposed pos e io o he op ic chiasma.
The ce ebellum (Fig. 1) occu s as ado sal olia ed body pos e io o he ce e-
b al hemisphe es. The ce ebella su ace
is
ela i ely simple, and a ely has mo e
han p imi i e sulci de eloped. The ce ebella body is composed
o
amedial
e mi o m body lanked by apai
o
la e al lobes. As in o he mammals, he
ce ebellum
is
a ached o he b ain s em
by
he in e io , medial, and supe io
peduncles.
The mesencephalon
o
phyllos oma id ba s
is
a ely seen unless he hypo-
physis has been emo ed. The ce eb al peduncles a e ne e isible ex e nally, and
o
he ec al s uc u es, he enla ged in e io colliculi a e gene ally he only s uc-
u es isible (bu e en hey a e no always so). The pons is almos comple ely
co e ed en ally by he hypophysis and la e ally
by
he igeminal ne e.
The medulla oblonga a (Fig.
1)
occu s
as
a a he b oad s uc u e
in
phyl-
los oma ids. Ven ally, he an e io apezoid body, pos e io oli es, and medially
loca ed py amidal decussa ion a e isible. As
in
o he e eb a es, he medulla
g ades pos e io ly in o he spinal co d.
Aspec s
o
Va ia ion in Ex e nal Ana omy
Wi hin he Phyllos oma idae, asu p ising a ay
o
a ia ion in ea u es
o
ex-
e nal b ain ana omy
is
encoun e ed. Da a conce ning in e and in asub amilial
a ia ion a e a ailable o
65
species ep esen ing 38 gene a and six sub amilies
(McDaniel, 1973). Sub amily Phyllos oma inae
Da a a e a ailable o 10
o
he
11
gene a lis ed in his sub amily by Jones
and Ca e in he sys ema ic accoun in his olume. P onounced a ia ion
o
se e al mo phological ea u es c ea es di icul y in cha ac e izing agene alized
b ain o his sub amily.
The b ain
o
Mimon (Fig. 2) is cha ac e ized
by
he p esence
o
ex emely
sho , deep ce eb al hemisphe es, pseudo empo allobes ha p ojec en ally in a
ounded a he han angula ashion, and asligh indica ion
o
apseudocen al
sulcus. The caudal e mina ion o he ce eb al henlisphe es
is
do sally an e io o
he in e io colliculi (which a e con iguous o one ano he ), esul ing in do sal
exposu e
o
p ecollicula ec unl. Mimon
is
he only phyllos oma id ba in which
he ec um
is
b oadly exposed an e io o he in e io colliculi. The ce ebellum
o
Mimon
is
simple, ha ing shallow olia ions.
The b ain
o
Loncho hina (no igu ed) esembles ha
o
Mimon in ha ing a
sho , deep ce eb um and asimple ce ebellum.
In
Loncho hina, he pseudocen al
sulci cu mo e deeply in o he ce eb al hemisphe es han in Mimon, and he pos-
e io ma gin o he ce eb um ex ends almos o he an e io ma gin
o
he in-
e io colliculi. The b ains
o
i e species
o
Mic onyc e is (Figs. 3-7) a y only
sligh ly among species. The ce eb um
o
Mic onyc e is
is
ela i ely longe han
ha
o
Mimon.
The
ce eb um is sho es in
M.
nice o i (Fig.
3)
and
M.
minu a
(Fig. 4), longe in
M.
schmid o um (Fig. 5) and
M.
megalo is (Fig. 6), and
longes in
M.
hi su a (Fig. 7). The ce eb al pseudocen al sulci a e well de eloped
in all species excep
M.
nice o i. The e
is
shallow de elopmen
o
asulcus an-
e io o he pseudocen al sulcus. The ce ebellum
is
simple in all species. Con-
sis en ly wi hin he genus Mic onyc e is, bu
in
no o he phyllos oma ine genus,
he in e io colliculi a e exposed do sally and a e no con iguous do sally wi h
one ano he . In Mic onyc e is, he in e io colliculi a e sepa a ed by he an e io
lobe
o
he e mi o m body
o
he ce ebellum.
The b ain
o
Mac o us (Fig. 8) esembles ha
o
Mic onyc e is megalo is in
mos de ails. Howe e , he do sally exposed in e io colliculi a e con iguous.
The deg ee
o
con igui y
is
educed and app oaches he condi ion ound in
Mic onyc e is because he an e io lobe
o
he ce ebella e mi o m body
p ojec s an e io ly o co e he mos pos e io po ion
o
he in e io colliculi.
In Mac ophyllum (Fig. 9), he ce eb al hemisphe es a e elonga e ela i e o he
condi ion
in
Mimon, and a e excep ionally smoo h, wi h only ashallow in e -
hemisphe ic sulcus. The in e io colliculi a e exposed do sally and a e con iguous
wi h one ano he . The ce ebella issue
is
excep ionally nondesc ip and ligh ly
issu ed.
The b ains
o
T achops (Fig. 10) and wo species
o
Tona ia (Figs. 11-12) a e
simila
in
ex e nal ana omy.
The
b ain
o
Tona ia nica aguae (Fig. 11)
is
he
smalles and leas o namen ed
o
he h ee.
I
is
cha ac e ized by he p esence
o
deep ce eb al hemisphe es ha a e ela i ely longe han hose
o
Mac ophyl-
um.
The
pseudocen al sulci
o
he ce eb um a e well de eloped, and he e
is
shallow de elopmen
o
he sulci an e io o he pseudocen al sulci. The in e io
colliculi a e exposed do sally, and a e con iguous middo sally. As in Mac o us,
he an e io edge
o
he e mi o m body
o
he ce ebellum p o udes o wa d o
co e he pos e io po ions
o
he in e io colliculi.
The
ce ebellum
is
simple in
appea ance. The b ain
o
Tona ia bidens (Fig. 12)
is
simila o ha
o
T.
nica aguae excep ha he ce eb al hemisphe es a e ela i ely longe and he
pseudocen al sulci
o
he ce eb um a e ex emely deep. The b ain
o
T achops
(Fig. 10) di e s om ha
o
Tona ia bidens by he p esence
o
shallowe pseu-
docen al sulci, bu somewha deepe sulci an e io o he pseudocen al sulci.
The b ain
o
T achops has some seconda y olia ion a he la e al edge
o
he
e mi o m body
o
he ce ebellum.
B ains
o
he gene a Phyllos omus (Figs. 13-15) and Phyllode ma (Fig. 16)
e eal ano he subg oup wi hin he Phyllos oma inae.
The
b ains
o
bo h gene a
152
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153
a e cha ac e ized
by
massi e ce eb al hemisphe es ha a e elonga e and
an e io ly blun . Sulca ion
o
he ce eb um
is
p onounced, and he pseudocen al
sulci and sulci an e io o hem a e well de eloped. The ce eb al hemisphe es a e
well p o ided wi h small seconda y issu es adia ing om he la ge sulci. In
Phyllos omus elonga us (Fig. 13), he in e io colliculi a e exposed do sally; bu
in
Phyllos omus has a us (Fig. 14),
P.
discolo (Fig. 15), and Phyllode ma (Fig.
16), he in e io colliculi a e comple ely co e ed
by
ce eb al and ce ebella is-
sues. In all ou species, he ce ebellum has o namen a ion
in
he o m
o
secon-
da y olia ion a he la e al edges
o
he e mi o m body, which
is
i sel enla ged
o o m ap onounced medial c es o he ce ebellum.
The b ain
o
Vampy um spec um (Fig. 17)
is
ex emely la ge, bu
is
no
as
massi e in appea ance as ha
o
Phyllos omus because
o
mo e p onounced
elonga ion. The ce eb al hemisphe es a e well con olu ed and sulca ed and ha e
seconda y issu es adia ing om he majo sulci. The ce ebellum achie es i s
maximum o namen a ion in Vampy um. The e mi o m body
is
aised o o m
an
ex ao dina y medial idge, and he e
is
conside able seconda y olia ion a he
la e al edges
o
he e mi o m body.
Sub amily Glossophaginae
B ains
o
nine gene a and
12
species we e examined om he Glossophaginae.
B ains om his sub amily ha e ela i ely smoo h and shallow ce eb al hemi-
sphe es, shallow de elopmen
o
he nlajo sulci, and do sally unexposed in e io
colliculi. The ce ebellum
is
simple and wi hou seconda y o namen a ion.
The b ains
o
Choe oniscus godmani (Fig. 18) and
C.
in e medius (no igu ed)
a e p obably impossible o di e en ia e ex e nally. The b ain
o
Choe oniscus
is
cha ac e ized
by
he p esence
o
asho ce eb um ha ing asmoo h su ace and
small ol ac o y bulbs. The pseudocen al sulcus
is
ex emely shallow, and he
pseudo empo al lobes a e smoo hly ounded on he en al side. The ce ebellum
is
simple and has no seconda y'loba ion.
Hylonyc e is (Fig. 19) and Lichonyc e is (Fig. 20) ha e b ains simila o ha
o
Choe oniscus. The ce eb um
is
sho and smoo h, he ol ac o y bulbs a e
small, and he pseudocen al sulci a e shallow. The pseudo empo al lobes
o
Hylonyc e is a e en ally ounded as in Choe oniscus, bu hose
o
Lichonyc e is
a e en ally angula . The ce ebellum
is
simple in bo h gene a.
The b ains
o
Glossophaga al icola (Fig. 21),
G.
commissa isi (Fig. 22), and
G.
so icina (no igu ed) a e i ually indis inguishable ex e nally.
In
Glos-
sophaga, he ce eb al hemisphe es a e sho , smoo h, and almos lacking sulci.
The ol ac o y bulbs a e small. The pseudo empo al lobes a e somewha angula
and he ce ebella olia ions a e simple.
The b ains
o
Choe onyc e is (Fig. 23) and Monophyllus (no igu ed) a e
cha ac e ized
by
ela i ely elonga ed ce eb al hemisphe es ha a e e ically
shallow. The pseudo empo al lobes a e he shallowes wi hin he Glossophaginae
and he ce ebellum has only p ima y olia ions.
Lep onyc e is (Fig.
1)
has ab ain much like ha
o
Glossophaga al icola.
Howe e , he ce eb al sulci end o cu deepe in o he mass
o
he ce eb um
in
Lep onyc e is han
in
any
o
he species
o
Glossophaga.

Anou a (Fig. 24) has ab ain simila o ha
o
Choe onyc e is. The ce eb um
is
elonga e and smoo h, and he pseudocen al sulci a e shallow. The ol ac o y
bulbs a e ela i ely la ge, and he pseudo empo al lobes a e shallow. In Anou a,
he ce eb al hemisphe es a e somewha mo e massi e han
in
Choe onyc e is, and
hey each he g ea es ela i e leng h wi hin his sub amily.
The b ain
o
Lonchophylla (Fig. 25) has he mos massi e ce eb um wi hin
he Glossophaginae. The ce eb um
is
elonga ed and has ela i ely well-de eloped
pseudocen al sulci and la ge ol ac o y bulbs. The pseudo empo al lobes p ojec
en ally, and he ce ebellum achie es i s maximum deg ee
o
olia ion wi hin he
Glossophaginae.
Sub amily Ca olliinae
B ains
o
bo h gene a in his sub amily ha e been examined. The b ains
o
Ca ollia (Fig. 26) and Rhinophylla (Fig. 27) a e simila in almos e e y de ail.
The b ains
o
Ca ollia pe spicilla a (Fig. 26) and
C.
sub u a' (no igu ed) a e
i ually iden ical ex e nally. In Ca ollia, he ce eb um is simila o ha
o
Mic-
onyc e is in ha ing ela i ely sho and smoo h hemisphe es. The pseudocen al
sulci a e well de eloped, as a e he sulci an e io o he pseudocen al sulci. The
pseudo empo al lobes a e ounded en ally, and he in e io colliculi a e no
exposed do sally. The ce ebellum
is
simple and has only p ima y lobes.
The b ain o Rhinophylla (Fig. 27)
is
simila o ha
o
Ca ollia, bu in Rhino-
phYlla, he pseudo empo allobes p ojec en ally in an angula ashion.
Sub amily S enode minae
This la ge sub amily
is
ep esen ed by specimens om 28 species, ep esen ing
12
gene a. B ains om his sub amily no mally ha e adeep ce eb um wi h
pseudo empo allobes ha p ojec en ally in adis inc i e ashion.
The genus A ibeus (Figs. 28-35)
is
ep esen ed he ein by nine species. The
b ain
o
A.
az ecus (Fig. 29)
is
cha ac e ized by he mos shallow ce eb al hemi-
sphe es wi hin he S enode minae. The pseudo empo al lobes a e angula and
p ojec asho dis ance en ally. The pseudocen al sulci a e shallow, and he e
a e no sulci an e io o he pseudocen al sulci. The in e io colliculi a e co e ed
do sally, and he e mi o m body
o
he ce ebellum o ms alow c es .
In
A.
phaeo is (Fig. 30), he ce eb al hemisphe es a e ela i ely deepe han in A.
az ecus. The pseudo empo al lobes a e angula and p ojec en ally a he
han in
A.
az ecus. The p epseudocen al gy us is enla ged and p o udes do sally.
The pos e io po ions
o
he in e io colliculi a e exposed do sally, and he ce e-
bellum esembles ha
o
A.
az ecus. The b ain
o
A. ol ecus (Fig. 31)
is
ana o-
mically in e media e be ween he b ains
o
A.
az ecus and A. phaeo is. In A.
ol ecus, he ce eb al hemisphe es a e deep and he pseudo empo allobes p ojec
en ally in
an
angula ashion. The pseudocen al sulci a e shallow, as a e he
sulci an e io o he pseudocen al sulci. The pos e io edges
o
he ce eb al
hemisphe es co e all bu he pos e io mos edges
o
he in e io colliculi. The
ce ebellum has amedial c es and small seconda y olia ions a he la e al edges
o
he e mi o m body. The b ains
o
A.
wa soni (Fig. 32) and
A.
cine eus (Fig.
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28) a e indis inguishable excep ha
A.
cine eus has ala ge b ain han does
A. wa soni. The b ains
o
hese wo species a e cha ac e ized
by
deep and
ela i ely smoo h ce eb al hemisphe es wi h shallow sulci. In "bo h species, he
in e io colliculi a e no exposed, and he ce ebellum
is
c es ed and wi hou sec-
onda y olia ion. The b ain
o
A. inopina us (Fig. 33)
is
cha ac e ized
by
deep
ce eb al hemisphe es ha ing angula and en ally p ojec ing pseudo empo al
lobes. The pseudocen al sulci and sulci an e io o he pseudocen al sulci a e
well de eloped. The in e io colliculi a e no exposed do sally, and he ce e-
bellum
is
simple and has ahigh medial c es . A. hi su us (no igu ed),
A.
jamai-
censis (Fig. 34), and
A.
li u a us (Fig. 35) ha e b ains ha a e simila
in
mo phology. In hese species, he b ain has ela i ely well-con olu ed ce eb al
hemisphe es and well-de eloped majo sulci. The pseudo empo al lobes p ojec
en ally in an angula ashion, and he in e io colliculi a e no do sally exposed.
The ce ebellum is well c es ed and has small seconda y olia ions a he la e al
edges
o
he e mi o m body.
The b ain
o
Enchis henes (Fig. 36)
is
known o me h ough examina ion
o
one poo ly p ese ed specimen. The ela i ely smoo h ce eb um wi h deep and
angula ly p ojec ing pseudo empo al lobes esembles ha
o
A ibeus wa son;.
The in e io colliculi a e no isible om abo e, and he ce ebellum
is
simple and
media.lly c es ed.
The genus Vampy odes (Fig. 37)
is
cha ac e ized by ab ain wi h la ge and
an e io ly blun ce eb al hemisphe es ha ing poo ly de eloped sulci. The pseudo-
empo al lobes p ojec en ally
in
an angula ashion. The pos e io po ions
o
he in e io colliculi a e do sally exposed and he ce ebellum
is
simple.
The
e -
mi o m body o ms amedial c es o he ce ebellum.
The b ains
o
U ode ma biloba um (Fig. 38) and
U.
magni os um (Fig. 39)
a e simila in mos ea u es. These b ains ha e deep ce eb al hemisphe es wi h
angula pseudo empo al lobes ha p o ude en ally o alesse ex en han in
A ibeus. The pseudocen al sulci and sulci an e io o he pseudocen al sulci
a e well de eloped. The in e io colliculi a e no exposed do sally, and he e mi-
o m body
o
he ce ebellum o ms alow medial c es . The e a e seconda y olia-
ions a he la e al edges
o
he e mi o m body.
B ains
o
S u ni a [ilium (no igu ed),
S.
mo dax (Fig. 40), and
S.
ludo ici
(Fig. 41) closely esemble each o he . These b ains a e cha ac e ized by deep and
ex emely smoo h ce eb al hemisphe es. The pseudocen al sulci and sulci an-
e io o he pseudocen aJ sulci a e mo e poo ly de eloped han in o he s eno-
de mine ba s. The pseudo empo aJ lobes a e angula and p ojec en ally. The
in e io colliculi a e comple ely co e ed in
S.
lilium and
S.
ludo ici, bu in
S.
mo dax, he pos e io po ions
o
he colliculi a e do sally exposed. In all h ee
species, he ce ebellum is simple and has amedial c es .
The b ain
o
Ec ophylla macconnelli (Fig. 42)
is
simila o ha
o
A ibeus
phaeo is. In
E.
macconnelli, he ce eb al hemisphe es a e deep and ela i ely
smoo h. The majo sulci a e well de eloped and he p epseudocen al gy us p o-
udes do sally. The pseudo empo al lobes a e angula and p o ude en ally.
The in e io colliculi a e exposed do sally, and he ce ebellum is simple and
c es ed.
The b ains
o
Vampy essa nymphaea (Fig. 43) and
V.
pusilla (Fig. 44) a e no
alike. The b ain
o
V.
nymphaea
is
cha ac e ized
by
deep and somewha domed
ce eb al hemisphe es ha ing well-de eloped majo sulci. The pseudo empo al
lobes p ojec en ally in an angula ashion. The in e io colliculi a e no ex-
posed do sally. The ce ebellum
is
simple and has alow medial c es . In
V.
pusilla, he ce eb um
is
smoo h and has well-de eloped sulci, bu
i
is
no domed
as
in
V.
nymphaea. In
V.
pusilla, he in e io colliculi a e exposed
do sa11y.
The
pseudo empo al lobes and ce ebellum
o
V.
pusilla esemble hose
o
V.
nymphaea.
The b ains
o
Chi ode ma sal ini (Fig. 45) and
C.
illosum (Fig. 46) a e sim-
ila and a e cha ac e ized by massi e ce eb al hemisphe es ha a e well sulca ed,
an e io ly blun , and somewha con olu ed.
The
nlajo ce eb al sulci a e deepe
in
C.
sal ini, bu ha e small seconda y issu es adia ing om hem
in
bo h
species. The pseudo empo al lobes a e massi e and p ojec en ally
in
he ypi-
cally angula ashion. The in e io colliculi a e no exposed do sally, and he
ce ebellum
is
c es ed medially and has small seconda y olia ions along he la e al
edges
o
he e mi o m body.
B ains
o
species
o
Vampy ops (Figs. 47-49) ha e massi e ce eb al hemi-
sphe es, deep sulci and high con olu ions in mos species, and a ela i ely com-
plex pa e n
o
olia ion o he ce ebellum. Among he species exanlined,
V.
hel-
le i (Fig. 47) has he leas con olu ed ce eb um and he shallowes ce eb al sulci.
In
V.
illa us (Fig. 48) and Vampy ops in uscus (Fig. 49), he ce eb al hemi-
sphe es a e deeply sulca ed and well con olu ed. The pseudo empo al lobes a e
ela i ely la ge
in
all h ee species and p ojec en ally in an angula ashion.
In
V.
helle i, he pos e io po ions
o
he in e io colliculi a e exposed do sally,
bu in he o he wo species, he in e io colliculi a e comple ely co e ed do sally
by
he ce eb al hemisphe es. In all h ee, he ce ebellum
is
c es ed medially, and
he e a e seconda y olia ions a he la e al edges
o
he e mi o m body.
The
b ain
o
Cenlu io (Fig. 50)
is
cha ac e ized by p onounced an e opos e io
comp ession esul ing in adomed b ain. The ce eb al hemisphe es a e ela i ely
smoo h and ha e shallow sulci. The pseudo empo al lobes a e also somewha
comp essed and p ojec en ally in adi e en ashion han in o he membe s
o
he sub amily. The in e io colliculi a e exposed do sally, and he ce ebellum
is
simple and sligh ly c es ed.
Ame ida (Fig. 51) has ab ain simila o ha
o
Cenlu io.
The
ce eb um
is
comp essed and qui e smoo h, wi h almos no ace
o
he majo sulci.
The
pseudo empo al lobes a e ela i ely shallow, angula , and p ojec en ally.
The ce ebellum
is
simple and has alow c es .
The b ain
o
S enode ma (Fig. 52) also esembles ha
o
Cen u io. S enode ma
is
cha ac e ized by massi e ce eb al hemisphe es ha a e ela i ely smoo h. The
majo ce eb al sulci a e shallow, and he pseudo empo allobes a e la ge, angula ,
and en ally p ojec ing.
The
in e io colliculi a e exposed do sally.
The
ce e-
bellum
is
simple and has alow c es .
156
SPECIAL
PUBLICATIONS
MUSEUM
TEXAS
TECH
UNIVERSITY
BIOLOGY
OF
THE
PHYLLOSTOMATIDAE
157
Sub amily Phyllonyc e inae
Two species (E ophylla
bombi/ ons
and B achyphylla ca e na um) we e ex-
amined om his sub amily. Phyllonyc e is
is
' he only genus no ep esen ed.
The b ain
o
E ophylla (Fig. 53) has a ela i ely sho and smoo h ce eb um,
wi h ashallow pseudocen al sulcus and asligh ly de eloped sulcus an e io o
he pseudocen al sulcus. The pseudo empo al lobes a e ounded en ally, and
he e is asimple pa e n
o
olia ion
o
he ce ebellum. The in e io colliculi a e
do sally exposed and a e no con iguous wi h each o he . In E ophylla, he e mi-
o m body
o
he ce ebellum cons i u es abou a hi d
o
he o al do sal exp es-
sion
o
he ce ebellum.
The b ain
o
B achyphylla (Fig. 54)
is
cha ac e ized
by
ela i ely smoo h
and massi e ce eb al hemisphe es. The majo ce eb al sulci a e
wel1
de eloped,
including he sulcus an e io o he pseudocen al sulcus. The pseudo empo al
lobes a e en ally angula , bu do no p o ude en ally. The in e io colliculi
a e no isible om abo e. The e mi o m body
o
he ce ebellum
is
la e ally en-
la ged and cons i u es hal o he do sal exposu e
o
he ce ebella issues. In ad-
di ion, B achyphylla has one cha ac e is ic no ound in any o he phyllos oma id
b ain in ha he u ula po ion
o
he ce ebellum
is
g ea ly enla ged and o ms
ap ominen lobe a he pos e io edge
o
he e mi o m body along he do sal
su ace
o
he medulla.
Sub amily Desmodon inae
B ains we e examined om all h ee gene a
o
his sub amily. The b ains
o
Desmodus (Fig. 55), Diaemus (Fig. 56), and Diphylla (no igu ed) a e simila
in ha hey
al1
ha e la ge ce eb al hemisphe es ha a e deeply sulca ed and
well con olu ed. The ce ebellum
is
a iously o namen ed.
The b ain
o
Desmodus (Fig. 55)
is
cha ac e ized
by
elonga e and con olu ed
ce eb al hemisphe es ha a e deeply cu by he pseudocen al sulci and he sulci
an e io o hem. The pseudo empo al lobes p ojec en ally in an angula
ashion (as
in
he S enode minae), and he in e io colliculi a e no do sally ex-
posed. The e mi o m body
o
he ce ebellum o ms amedial c es , and he e a e
small seconda y olia ions along i s la e al edges.
In Diaemus (Fig. 56), he ce eb al hemisphe es a e less elonga e han
in
Desmodus bu a e well con olu ed and deeply sulca ed. The pseudo empo al
lobes a e angula as
in
Desmodus and he in e io colliculi a e no do sally ex-
posed. The ce ebellum has alow medial c es and seconda y olia ions along he
la e al edges
o
he e mi o m body.
Diphylla has ace eb um simila o ha
o
Diaemus. The pseudo empo al
lobes a e angula and p ojec en ally. The pos e io po ions
o
he in e io
colliculi a e do sally exposed, and he ce ebellum
is
la ge and has amedial c es .