Roosting behavior.

Demog aphy and Na u al His o y
o he Common F ui Ba ,
A ibeus jamaicensis,
on Ba o Colo ado Island, Panama
CHARLES O. HANDLEY, JR.,
DON E. WILSON,
and
ALFRED L. GARDNER
EDITORS
SMITHSONIAN CONTRIBUTIONS
TO
ZOOLOGY
•
NUMBER
511
SERIES PUBLICATIONS OF THE SMITHSONIAN INSTITUTION
Emphasis upon publica ion as a means o "di using knowledge" was exp essed by he i s
Sec e a y o he Smi hsonian. In his o mal plan o he Ins i u ion, Joseph Hen y ou lined a
p og am ha included he ollowing s a emen : "I is p oposed o publish a se ies o epo s,
gi ing an accoun o he new disco e ies in science, and o he changes made om yea o yea
in all b anches o knowledge." This heme o basic esea ch has been adhe ed o h ough he
yea s by housands o i les issued in se ies publica ions unde he Smi hsonian imp in ,
commencing wi h Smi hsonian Con ibu ions o Knowledge in 1848 and con inuing wi h he
ollowing ac i e se ies:
Smi hsonian Con ibu ions o An h opology
Smi hsonian Con ibu ions o As ophysics
Smi hsonian Con ibu ions o Bo any
Smi hsonian Con ibu ions o he Ea h Sciences
Smi hsonian Con ibu ions o he Ma ine Sciences
Smi hsonian Con ibu ions o Paleobiology
Smi hsonian Con ibu ions o Zoology
Smi hsonian
Folkli e
S udies
Smi hsonian S udies in Ai and Space
Smi hsonian S udies in His o y and Technology
In hese se ies, he Ins i u ion publishes small pape s and ull-scale monog aphs ha epo
he esea ch and collec ions o i s a ious museums and bu eaux o o p o essional colleagues
in he wo ld o science and schola ship. The publica ions a e dis ibu ed by mailing lis s o
lib a ies, uni e si ies, and simila ins i u ions h oughou he wo ld.
Pape s o monog aphs submi ed o se ies publica ion a e ecei ed by he Smi hsonian
Ins i u ion P ess, subjec o i s own e iew o o ma and s yle, only h ough depa men s o he
a ious Smi hsonian museums o bu eaux, whe e he manusc ip s a e gi en subs an i e e iew.
P ess equi emen s o manusc ip and a p epa a ion a e ou lined on he inside back co e .
Robe McC. Adams
Sec e a y
Smi hsonian Ins i u ion
SMITHSONIAN CONTRIBUTIONS
TO
ZOOLOGY
•
NUMBER
511
Demog aphy and Na u al His o y
o he Common F ui Ba ,
A ibeus jamaicensis,
on Ba o Colo ado Island, Panama
Cha les O. Handley, J .,
Don E. Wilson,
and
Al ed L. Ga dne
EDITORS
SMITHSONIAN INSTITUTION PRESS
Washing on, D.C.
1991
ABSTRACT
Handley, Cha les O., J ., Don E. Wilson, and Al ed L. Ga dne , edi o s. Demog aphy and
Na u al His o y o he Common F ui Ba , A ibeus jamaicensis, on Ba o Colo ado Island,
Panama. Smi hsonian Con ibu ions o
Zoology,
numbe
511,
173 pages, 69 igu es, 62 ables,
1991.—Ba s we e ma ked and moni o ed on Ba o Colo ado Island, Panama, o s udy seasonal
and annual a ia ion in dis ibu ion, abundance, and na u al his o y om 1975 h ough 1980.
Da a ga he ed ad ances ou knowledge abou locking; abundance; eeding s a egies; social
beha io ; species
ichness;
popula ion s uc u e and s abili y; age and sex
a ios;
li e expec ancy
and longe i y; nigh ly, seasonal, and annual mo emen s; synch ony wi hin and be ween species
in ep oduc i e ac i i y; iming o ep oduc i e cycles; su i al and dispe sal o ec ui s;
in a-and in e -speci ic ela ionships; and day and nigh oos selec ion.
Ba o Colo ado Island (BCI) ha bo s la ge popula ions o ba s ha eed on he ui o canopy
ees,
especially igs. These ees a e abundan , and he indi idual asynch ony o hei ui ing
hy hms esul s in a ai ly uni o m abundance o ui . When igs a e sca ce, a a ie y o o he
ui s is a ailable o eplace hem. This ela i ely dependable ood supply a ac s
a
ema kably
ich guild o ba s.
Al hough we ma ked all ba s caugh , we ied o maximize he numbe o A ibeus
jamaicensis ne ed, because i is abundan
Q-h
o
he
o al ca ch o ba s on BCI), easily cap u ed
by con en ional means (mis ne s se a g ound le el), and esponds well o handling and
ma king.
An a e age A ibeus jamaicensis is
a 45
g ugi o e ha ea s oughly i s weigh in ui e e y
nigh . These ba s p e e igs and o en seek hem ou e en when o he ypes o ui hey migh
ea a e a mo e abundan . They commu e se e al hund ed me e s o eeding ees on he
a e age, eeding on ui om one o ou ees each
nigh ,
and e u ning o a single ui ing ee
an a e age o ou nigh s in succession. The ba s end o ly a he when ewe ig ees a e
bea ing ipe ui , and hey eed om ewe ees, on he a e age, when he moon is nea ly ull.
These
ba s,
like hei congene s, do no eed in he ui ing ee
i sel .
Ins ead, hey selec a ui
and ca y i o a eeding oos ypically abou 100 m away be o e ea ing i . We u ilized adio
eleme y o assess eeding a es om he numbe o " eeding passes"— ansi s be ween ui
ee and eeding oos . Ba s a e o en ne ed while ca ying ui , e ealing hei die . Feces also
e eal die a y in o ma ion.
Adul emale
A.
jamaicensis li e
in
ha ems o h ee o 30 indi iduals wi h
a
single adul male.
On BCI he ha em g oups oos du ing he day in hollow ees. The e is p esumably a la ge
popula ion o su plus males ha oos oge he wi h nonadul s o bo h sexes in oliage. Females
commu e an a e age o 600 m om hei day oos s o eeding si es, and ha em males a el less
han
300 m.
Twice
a
yea mos emales gi e bi h o
a
single young, once in Ma ch o Ap il, and
again in July o Augus ; ac i e ges a ion a e ages abou 19 weeks. Ju eniles a e i s ne ed
when hey a e abou en weeks old, and emales usually i s bea young in Ma ch o Ap il
ollowing hei yea o bi h.
OFFICIAL
PUBLICATION
DATE
is hands amped in a limi ed numbe o ini ial copies and is
eco ded in he Ins i u ion's annual epo ,
Smi hsonian
Yea .
SERIES
COVER
DESIGN:
The co al
Mon as ea ca e nosa (Linnaeus).
Lib a y o Cong ess Caialoging-in-Publicalion Da a
Handley, Cha les O. (Cha les O e oil), 1924-
Demog aphy and na u al his o y o he common ui ba , A ibeus jamaicensis, on Ba o Colo ado Island, Panama /
Cha les O. Handley, J ., Don E. Wilson, and Al ed L. Ga dne .
p.
cm. — (Smi hsonian con ibu ions o zoology, no. 511)
Includes bibliog aphical e e ences (p. ).
1.
A ibeus jamaicensis—Panama—Ba o Colo ado Island.
I. Wilson, Don E. II. Ga dne , Al ed L. III. Tide. IV. Se ies.
QL1.S54 no. 511 [QL737.C57] 591
s-dc20
[599.4] 91-13449
@ The pape used in his publica ion mee s he minimum equi emen s o he Ame ican
Na ional S anda d o Pe manence o Pape o P in ed Lib a y Ma e ials Z39.48—1984.
Con en s
Page
1.
INTRODUCTION
by
Cha les
O.
Handley,
J ., Don E.
Wilson,
and
Al ed
L.
Ga dne
1
2.
PHYSIOLOGY
by
Eugene
H.
S udie
and Don E.
Wilson
9
3.
REPRODUCTION IN A CAPTIVE COLONY
by Lucinda Keas Ta and Cha les
O. Handley,
J 19
4.
REPRODUCTION
ON
BARRO COLORADO ISLAND
by Don E.
Wilson,
Cha les
O.
Handley,
J ., and
Al ed
L.
Ga dne
43
5.
SURVIVAL AND RELATIVE ABUNDANCE
by Al ed L. Ga dne , Cha les O.
Handley,
J ., and Don E.
Wilson
53
6.
POPULATION ESTIMATES
by
Egbe
G.
Leigh,
J ., and
Cha les
O.
Handley,
J .. . 77
7.
MOVEMENTS
by
Cha les
O.
Handley, J ., Al ed
L.
Ga dne ,
and
Don
E.
Wilson
89
8.
ROOSTING BEHAVIOR
by
Douglas
W.
Mo ison
and
Cha les
O.
Handley,
J . . . 131
9.
FORAGING BEHAVIOR
by
Cha les
O.
Handley, J .,
and
Douglas W. Mo ison
. . 137
10.
FOOD HABITS
by
Cha les
O.
Handley,
J .,
Al ed
L.
Ga dne ,
and
Don
E.
Wilson
141
11. DIET AND FOOD SUPPLY
by Cha les O. Handley, J ., and Egbe G. Leigh, J .. 147
12.
APPENDIX: METHODS
OF
CAPTURING
AND
MARKING TROPICAL BATS
151
LITERATURE CITED
167

FRONTISPIECE.—A ibeus jamaicensis. Pencil ske ch by Nancy Mo an, Ba o Colo ado Island, Panama^ Oc obe ,
1976.
Demog aphy and Na u al His o y
o he Common F ui Ba ,
A ibeus jamaicensis,
on Ba o Colo ado Island, Panama
1.
In oduc ion
Cha les
O.
Handley, J ., Don E. Wilson,
and Al ed
L.
Ga dne
BCI Ba P ojec
In 1974, Cha les Handley was in i ed o de elop a p ojec o
moni o ba s as pa o he Smi hsonian T opical Resea ch
Ins i u e's (STRI) long- e m en i onmen al moni o ing p o-
g am on Ba o Colo ado Island (BCI). The STRI moni o ing
p og am, launched in 1970 and suppo ed by he Smi hsonian
En i onmen al Sciences P og am (ESP), sough o moni o a
wide a ay o bio ic and physical en i onmen al componen s o
he island con inuously o e a long pe iod o ime.
The BCI Ba P ojec was bo n unde he adminis a i e i le:
"Biomass and ene ge ics o selec ed popula ions in Panama:
Ba s."
We wan ed o moni o demog aphic pa ame e s and
na u al his o y o all he ba s egula ly ound on BCI. Based on
ou ea lie expe iences, we hough he auna migh o al 40
species o ba s. The leng h o he p ojec was designed o
con inue h ough a gene a ion o
ba s,
howe e long ha migh
be.
The only clue o possible du a ion was he epo (Wilson
and Tyson, 1970) o a se en-yea -old A ibeus jamaicensis on
BCI.
A he ou se i was e iden ha ESP unds we e sp ead o e
Cha les O. Handley, J ., and Don E. Wilson, Na ional Museum o
Na u al His o y, Smi hsonian Ins i u ion, Washing on, D.C. 20560.
Al ed L. Ga dne , NERC, US. Fish and Wildli e Se ice, Na ional
Museum o Na u al His o y, Washing on, D.C. 20560.
Re iew Chai man: W. Ronald Heye , Smi hsonian Ins i u ion. Th ee
anonymous e iewe s a e g a e ully acknowledged.
oo many p ojec s o be able o suppo a eally meaning ul
moni o ing p ojec o ba s. Clyde Jones, hen Di ec o o he
Na ional Fish and Wildli e Labo a o y, U.S. Fish and Wildli e
Se ice, o e ed o p o ide bo h inancial and pe sonnel
suppo , and he Ba P ojec became a join en u e o he
Smi hsonian Ins i u ion and he U.S. Fish and Wildli e Se ice,
wi h Michael A. Bogan, Al ed L. Ga dne , and Don E. Wilson
joining Handley as ield c ew leade s.
Handley made se e al ips o BCI in 1975 and 1976 o
become amilia wi h he island and i s ba s, as well as o
de e mine wha was easible and how o o ganize he p ojec .
On 2 July 1977 a yea - ound cap u e and ma king p og am
began. Wi h he help o collabo a o s and dozens o olun ee s,
we ook u ns manning he ield su ey on BCI un il No embe
1980 when his phase o he P ojec was comple ed. The ea e ,
Handley con inued wo k on BCI on a pe iodic basis— he all
o 1981, he all o 1982, and he 12 mon hs om Sep embe
1984 h ough Augus 1985—wi h he suppo o he Smi h-
sonian's ESP, STRI, and Na ional Museum o Na u al His o y
(NMNH). The pu pose o he con inuing s udy was o main ain
he pool o ma ked ba s, e ine he demog aphic da a, and gain
u he in o ma ion on he biology o he ba s, pa icula ly hei
esponses o ood sou ces.
Du ing he econnoi e ing phase a he beginning o he
p ojec we ocused much o ou a en ion on de eloping a
eliable, long-las ing, ha mless ma king sys em. We es ab-
lished colonies o ba s a he Na ional Zoological Pa k (NZP)
ha we used in ma king expe imen s and in es ablishing
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
s anda ds o desc ibing age and ep oduc i e s a e. As a esul ,
we disca ded o ea m banding, a ooing, and hea b anding and
se led
on
necklacing wi h s ainless s eel ball chain. Band-
bea ing necklaces we e i s a ached o ee-li ing ba s on BCI
on 18 Oc obe 1976. By he all
o
1980 we had placed 18,953
necklaces on ba s.
The p ojec e ol ed apidly.
We
soon ealized ha
ou
esou ces we e
no
su icien
o
moni o simul aneously
all
species
o
ba s
on
BCI
in an
e ec i e manne . We se led
on
s udying
he ba
ha
had
p o ed
o be ou
p inciple
ca ch—A ibeus jamaicensis Leach,
he
common ui -ea ing
ba .
A i s we es ablished ne ing s a ions
a
places ha , based
on expe ience elsewhe e, "looked good
o
ba s," such
as
s eam alleys, ails h ough open o es , and gaps along idges
* The use o p oduc b and names in his publica ion is no in ended as
an endo semen o he p oduc s by he Smi hsonian Ins i u ion.
whe e unde b ush and o he ege a ion did no in e e e wi h
he ne s o obs uc ly
ways.
Howe e , i became appa en ha
abundance o ba s coincided wi h nea by ui sou ces and ha
wha
a
i s appea ed o be complica ed dis ibu ional pa e ns
o he island's ba s p o ed
o be
no hing mo e ha
a
di ec
co ela ion be ween o aging ac i i y and he une en dis ibu-
ion and a ailabili y
o
ui . We imp o ed ou ne ing success
by making sys ema ic su eys along he ails on BCI o loca e
pelle s
o
ui pulp d opped
by
eeding ba s and hen se ing
ou ne s nea by whe e hey we e mos likely o ca ch ba s.
We ealized ea ly
in he
p ojec ha squeaking ba s o en
a ac ed o he s,
and in
1978
we
began
o use he
wooden
Audubon Bi d Call as
a
subs i u e
o
a ac ba s in o he ne s.
A
a
good si e,
an
Audubon Bi d Call*,
an
occasional
squeaking ba , and one o wo ne s could p oduce enough ba s
o keep e e yone busy o hou s.
In he beginning we some imes caugh only wo o h ee ba s
in
a
nigh , and we we e sa is ied wi h ca ches
o
30 ba s. La e ,
TABLE
1-1.—Ba s
eco ded
on
Ba o
Colo ado
Island,
Ga un
Lake,
Panama.
Family
EMBALLONURIDAE
Rhynchonyc e is naso (Wied)
Saccop e yx bllineala (Temminck)
Saccop e yx lep u a (Sch ebe )
Co mu a b e i os is (Wagne )
Cen onyc e is maximiliani (Fische )
Family
NOCTOJONIDAE
Noc ilio albi en is Desma es
Noc ilio lepo inus Linnaeus
Family
MORMOOPIDAB
P e ono us gymnono us Wagne
P e ono us pa nellii (G ay)
Family
PHYLLOSTOMIDAE
Sub amily
PHYLLOSTOMINAE
Mic onyc e is b achyo is (Dobson)
Mk onyc e s hi su a (Pe e s)
Mic onyc e is megalolis (G ay)
Mic onyc e is nice o i Sanbom
Mic onyc e is schmid o um Sanbom
Mac ophyllum mac opkyllum (Schinz)
Tona ia bidens (Spix)
Tona ia sil icola D'O bigny
Mimon c enulalum (E. Geo oy)
Pkyllos omus discolo Wagne
Pkyllos omus haslalus(PMas)
Pkyllode ma s enops Pe e s
T achops ci hosus (Spix)
Ch olop e us au i us (Pe e s)
Vampy um
spec um (Linnaeus)
Sub amily
GLOSSOPHAOINAE
Glossophaga commissa is i Ga dne
Glossophaga so icina (Pallas)
Lonchophylla obus a Mille
Sub amily
CAROLLINAE
Ca ollia b e icauda (Schinz)
Ca ollia cas anea H. Allen
Ca ollia pe spicilla a (Linnaeus)
Sub amily STURNIRINAE
S u ni a luisi Da is
Sub amily
STENODERMATTNAE
U ode ma biloba um Pe e s
U ode ma magni os um Da is
Pla y hinus helle i (Pe e s)
Vampy odes ca accioli (Thomas)
Vampy essa nymphaea Thomas
Vampy essa pus ilia (Wagne )
Chi ode ma illosum Pe e s
Mesophylla macconnelli Thomas
A ibeus ha ii Thomas
A ibeus jamaicensis Leach
A ibeus li u a us (Ol e s)
A ibeus phaeo is (Mille )
A ibeus wa soni Thomas
Ame ida cen wio G ay
Cen u io senex G ay
Sub amily
DESMODONTINAE
Desmodus o undus (E. Geo oy)
Family
THYROPTERTOAE
Thy op e a disci/e a (lich ens ein and Pe e s)
Tky op e a icolo Spix
Family
VESPERTIUONIDAE
Myo is albescens (E. Geo oy
Myo is nig icans (Schinz)
Rhogeessa umida H. Allen
Family
MOLOSSIDAE
Tada ida la icauda a (E. Geo oy)
Molossus bondae J.A. Allen
Molossus coibensis J.A. Allen
Molossus molossus (Pallas)
NUMBER 511
we we e disappoin ed wi h less han 100 ba s pe nigh , and we
logged many nigh s wi h ca ches exceeding 200. Ou bes ca ch
came on 25 Oc obe 1979 a a gian Ficus dugandii wi h ipe
ui whe e we ne ed 282 ba s in abou ou hou s.
As o 1985, we had ound 56 species o ba s on BCI (Table
1-1). Bonacco so (1979) ca ego ized he ba s o he island in o
nine " eeding guilds." The dis ibu ion o he 56 species among
Bonacco so's guilds is: canopy ugi o es (14), g ounds o y
ugi o es
(4),
sca enging ugi o es (2), omni o es (4),
sanguini o es (1), gleaning ca ni o es (12), slow- lying hawk-
ing insec i o es (14), as - lying hawking insec i o es (4), and
pisci o es (1). A. jamaicensis, he majo subjec o his epo ,
is a canopy ugi o e.
By e e y measu e A. jamaicensis is he mos widesp ead and
abundan ba on BCI (Table 1-2). On a yea ly basis, i a e aged
60%
o he o al ca ch o ba s, and we caugh i almos e e y
nigh ha ne s we e se . Al oge he , in he pe iod 1975-1980
(including ba s cap u ed be o e ma king wi h necklaces began),
we eco ded 17,820 cap u es o
A.
jamaicensis.
By he end o 1980, we had lea ned enough abou A.
jamaicensis including i s popula ions, ep oduc ion, mo e-
men s, o aging, and physiology o jus i y a pause o
TABLE 1-2.—Measu es o abundance o ba s cap u ed on BCI du ing 1979. Ba s we e ne ed on 157 nigh s, and
cap u es (including bo h ma ks and ecap u es) o alled 9118 ba s. Tabula ions a e by equency o cap u e
(numbe
and
pe cen age o nigh s caugh ), numbe caugh ( o al), ca ch pe species nigh ( o al o
a
species caugh
di ided by numbe o nigh s i was caugh ), and ca ch pe ne ing nigh ( o al o a species caugh di ided by o al
nigh s o ne ing).
Species
A ibeus jamaicensis
U ode ma biloba um
A ibeus li u a us
Chi ode ma illosum
Ca ollia pe spicilla a
Vampy odes ca accioli
Phyllos omus discolo
A ibeus phaeo is
Ca ollia cas anea
Mic onyc e is hi su a
Vampy essa pusilla
Vampy essa nymphaea
P e ono us pa nellii
Tona ia sil icola
Tona ia bidens
A ibeus walsoni
Mic onyc e is megalo is
Glossophaga so icina
Phyllos omus has a us
Mimon c enula um
Rhogeessa umida
T achops ci hosus
Co mu a b e i os is
Pla y hinus helle i
Mic onyc e is b achyo is
Mac ophyllum mac ophyllum
Mic onyc e is nice o i
Saccop e yx bilinea a
Desmodus o undus
Glossophaga commissa isi
Myo is nig icans
Mic onyc e is schmid o um
Phyllode ma s enops
Cen u io senex
Vampy um spec um
U ode ma magni os um
Lonchophylla obus a
Ca ollia b e icauda
A ibeus ha ii
Mean ca ch
pe species
nigh
36.3
9.2
6.0
4.4
4.0
3.9
3.3
2.2
2.0
1.8
1.8
1.8
1.7
1.6
1.6
1.6
1.6
1.6
1.6
1.5
1.5
1.4
1.4
1.3
1.2
1.2
1.1
1.1
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
1.0
Rank
1
2
3
4
5
6
7
8
9
10
10
10
13
14
14
14
14
14
14
20
20
22
22
24
25
25
27
27
29
29
29
29
29
29
29
29
29
29
29
To al
5484
551
717
172
428
325
130
235
154
91
87
51
86
113
77
67
35
28
26
50
3
70
7
31
13
11
21
8
12
8
7
5
5
3
2
2
1
1
1
Mean ca ch
pe ne ing
nigh
34.9
3.5
4.6
1.1
2.7
2.1
0.8
1.5
1.0
0.6
0.6
0.3
0.6
0.7
0.5
0.4
0.2
0.2
0.2
0.3
0.02
0.4
0.05
0.2
0.08
0.07
0.1
0.05
0.08
0.05
0.05
0.03
0.03
0.02
0.01
0.01
0.01
0.01
0.01
Nigh s caugh
N
151
60
120
39
107
83
40
108
78
52
49
28
51
71
49
43
22
17
16
33
2
49
5
23
11
9
19
7
12
8
7
5
5
3
2
2
1
1
1
%
96
38
76
25
68
53
25
69
50
33
31
18
32
45
31
27
14
11
10
21
1
31
3
15
7
6
12
4
8
5
4
3
3
2
1
1
1
1
1
10SMITHSONIAN CONTRIBUTIONS
TO
ZOOLOGY
in a T, ange o 8.0°-33.2°C. The eg ession coe icien o his
ela ionship
(0.933;
SE = 0.202) does no di e signi ican ly
om 1.0 and we conclude ha in A. jamaicensis, Tb pa allels
Ta. Indi idual ba s main ain a cons an ange o Tb 6.6°-8.3°C
highe han T, h oughou he es ed T, ange.
P e iously, Mo ison and McNab (1967) epo ed li le daily
luc ua ion in Tb cycles in cap i e A. jamaicensis a an
es ima ed T, o
27°C.
Du ing dayligh , some o hei da a poin s
(13/128 = 10.1%) we e signi ican ly lowe han a e age. They
demons a ed ha A. jamaicensis exhibi ed he he mo egula-
o y pa e n o a homeo he mic endo he m h oughou a T,
exposu e ange o 5°-39°C. McNab (1969) ob ained simila
esul s in cap i e A. jamaicensis held o up o wo weeks
be o e es ing; no ba s had educed Tb ela i e o hei mean
empe a u es.
We ein es iga ed hese con adic o y esul s and concluded
ha al hough di e ences in he mo egula o y pa e ns migh
ha e been ela ed o gene ic di e ences in he popula ions
s udied, mo e likely he a ia ion e lec ed di e en me hodo-
logical app oaches (S udie and Wilson, 1979). The p ima y
signi icance o ou 1979 s udy was he demons a ion o a
ansi ion om a he e o he mic pa e n o Tb egula ion on he
day o cap u e o a homeo he mic pa e n a e h ee days o
cap i i y. Such a "cap i i y e ec " o
A.
jamaicensis may help
o esol e appa en ly con lic ing he mo egula o y pa e ns in
o he mammals, o example, Myo is luci ugus (S ones and
Wiebe s, 1967; S udie and O'Fa ell, 1972) and Pe omyscus
leucopus (Gae ne e al., 1973; Hill, 1977).
Al hough he cap i i y e ec explains di e gen da a on
he mo egula ion, i does no show which da a ep esen he
na u al he mo egula o y pa e n. S udie and O'Fa ell (1972)
ound ha he Tb o Myo is luci ugus and M. hysanodes in hei
na u al oos ing si es was highly a iable and simila o he da a
o newly caugh , labo a o y- es ed indi iduals. Appa en ly,
da a om ba s es ed soon a e cap u e be e e lec na u al
he mo egula o y pe o mance, whe eas da a om cap i e ba s
held longe e lec hei g ea es homeo he mic capabili ies.
We did no a emp o de e mine which componen s o he
cap i i y e ec a e esponsible, ei he singly o in combina ion,
o he changeo e in he mo egula o y pe o mance, bu
se e al possibili ies exis . The inc eased homeo he mic e-
sponse p obably does no esul om he mal acclima ion. We
did no y o hold cap i e ba s a cons an T,. Ins ead, T, o
cap i e ba s luc ua ed in sligh ly mu ed ashion wi h ha o
BCI's na u al en i onmen (S udie and Wilson, 1970).
Fu he mo e, he cap i i y e ec does no seem o be a gene al
s ess esponse because s ess would be g ea es du ing he
ini ial hou s o cap i i y; hus ba s would be expec ed o exhibi
he mos igid homeo he my in day ze o es ing. The cap i i y
e ec may esul om indi idual o combined ac ions o
educed ac i i y while caged o he con inual p esence o
excess ood, which he ba s can ea a will.
In i s na u al en i onmen , A. jamaicensis is p obably a
homeo he mic endo he m du ing pe iods o eeding and ligh
ac i i y, bu loosens Tb con ol (becoming a nonhomeo he mic
endo he m) du ing oos ing (non eeding and non lying epi-
sodes).
The sligh educ ion in Tb a such imes would conse e
la ge amoun s o ene gy (S udie , 1981). He e o he my in A.
jamaicensis conse es 38.7% and 67.4% o he ene gy ha
would be equi ed o homeo he mic indi iduals a ambien
empe a u es o 30°C and 25°C, espec i ely (S udie and
Wilson, 1970). This would amoun o a majo ene gy cos
educ ion du ing a oos ing pe iod. We assume ha he sligh
Tb educ ion in A. jamaicensis (Tb o 35.2°C a Ta o 30°C and
Tb o 32.5°C a Ta o 25°C) would no educe esponsi eness o
en i onmen al s imuli du ing oos ing no would i p eclude
ini ia ion o ligh .
The mo egula ion p obably is dependen on nu i ional s a e
in cap i e ba s. In he wild, A. jamaicensis eeds on ui ha
a ies seasonally om sca ce o plen i ul. Indi idual ba s may
unde go a na u al pe iod o diel o po , whe eas cap i e animals
wi h unlimi ed ood may ne e become o pid as long as hei
ood supply is cons an and plen i ul. Cap i e ba s ou inely
weigh mo e han wild-caugh indi iduals (see Sec ion 3,
Rep oduc ion in a Cap i e Colony).
The signi ican ques ions ega ding possible e ec o
nu i ion on he a iabili y o Tb in A. jamaicensis a e: Does he
he e o he mic pa e n o ba s es ed immedia ely a e cap u e
e lec unde nou ished indi iduals? O , is he homeo he mic
pa e n o ba s kep in cap i i y a esponse o o e nou ishmen
and inac i i y? Unde nou ished o no , poo Tb egula ion in
newly caugh A. jamaicensis is no ela ed o physiological
compe ence, bu e lec s educed me abolic hea p oduc ion
and a e o deple ion o ene gy s o es. Manakins, which a e
small ugi o ous bi ds, also exhibi he e o he my on BCI
(Ba holomew e al., 1983).
McNab (1969) epo ed a es ing me abolic a e o 1.70
cc/g/h o
A.
jamaicensis wi hin i s he mal neu al zone (TNZ)
and a he mal conduc ance o 0.17 cc/g/h /°C (below i s TNZ).
As wi h o he ugi o ous ba s, his mass-speci ic s anda d
me abolic a e is sligh ly highe han would be p edic ed by
Kleibe 's (1932) classic ela ion be ween me abolism and mass
in mammals. The ene ge ic cos o high Tb homeo he my in A.
jamaicensis is, he e o e, highe han expec ed o
a
mammal o
i s size.
McNab (1983) p esen ed ex ensi e a gumen s conce ning
ene ge ics, body sizes, and he limi s o endo he my ha can be
exp essed by a minimum bounda y cu e o high Tb
homeo he my. This cu e es ima es he smalles mass a which
con inuous endo he my can occu o a gi en me abolic a e.
When his minimum bounda y cu e is d awn by ela ing he Tb
o Ta di e en ial as a unc ion o body mass, da a o A.
jamaicensis alls almos exac ly on he cu e (McNab, 1982).
A.
jamaicensis, he e o e, would be p edic ed o be ma ginally
able o main ain i s epo ed high homeo he mic Tb o Ta
di e en ial. I is, he e o e, no su p ising ha unde nou ished
(o no mally nou ished) ba s es ed immedia ely a e cap u e
main ain a ma kedly lowe Tb and, consequen ly, lowe Tb o T

NUMBER
51111
di e en ials. As s a ed p e iously, he sligh Tb educ ion ound
in A. jamaicensis sa es 38.7%-67.4% o he ene gy equi ed
o main enance o highe Tb. Such ene gy conse a ion would
seem c i ically impo an in a species such as A. jamaicensis
ha has se e ely limi ed ese es o s o ed ene gy.
BODY
FAT.—To al
body a is a di ec indica ion o o e all
nu i ional s a us in e eb a es. Annual a iabili y in body a
con en in nume ous Neo opical ba s, including A. jamaicen-
sis,
has been epo ed (McNab, 1976; 1982). Howe e ,
Neo opical ugi o es and nec a i o es demons a e less o he
seasonal a ia ion and none o he gende - ela ed di e ences
ha cha ac e ize empe a e zone insec i o es. All Neo opical
ba s exhibi low peak a ese e le els when compa ed wi h
empe a e zone ba s (Bake e al., 1968; Ewing e al., 1970;
Pagels, 1975; Webe and Findley, 1970). The ex emely low a
ese es epo ed by McNab (1976) o A. jamaicensis
emphasize i s need o educe daily ene gy expendi u es. Lack
o a ese es and high in ake o die a y ca bohyd a e sugges
ha glycogen should be examined as he no mal ene gy ese e
o A. jamaicensis.
Glycogen le els ha e no been epo ed o A. jamaicensis,
bu compa able da a a e a ailable om megachi op e ans ha ,
al hough no ela ed o A. jamaicensis, a e nu i ional equi a-
len s.
Daily a ia ion in glycogen and a le els in li e and
ligh muscle issue in Eidolon hel um, a Paleo opical
ugi o ous ba , ha e been epo ed by Okon e al. (1978).
Thei indings show ha li e glycogen le els a sun ise (90.0
mg/g) a e ex emely high in compa ison wi h le els seen in
la ge domes ica ed mammals (Wa and Me ill, 1963). Li e
glycogen hen d ops p ecipi ously in E. hel um un il sunse
when le els (35.0 mg/g) each a ange no mal o la ge
mammals. B eas muscle glycogen in his ba emains low
(6.0-8.1 mg/g) and cons an h oughou he oos ing pe iod.
The ex eme d op in li e glycogen sugges s ha glycogen (as
glucose) is he p ima y ene gy sou ce o E. hel um h oughou
i s oos ing pe iod. Fa concen a ions in li e and b eas
muscle in E. hel um show sligh inc eases a sunse , bu he
ange o all alues (5.0-10.3 mg/100 g) is nea ly wo o de s o
magni ude less han a le els in li e and muscle in o he
mammals, la ge and small (Ki kham and All ey, 1972; Wa
and Me ill, 1963). These glycogen le els e lec he high
ca bohyd a e, low lipid composi ion o he die o hese ba s.
Van de Wes huyzen's (1978) epo on ano he Paleo opi-
cal ui ba , Rouse us aegyp iacus, p o ides addi ional suppo
o he ex eme impo ance o glucose o glycogen and he
ela i e unimpo ance o a as an ene gy sou ce in opical
ugi o ous ba s. He epo ed he diu nal cycle o se e al
me aboli es including glucose, ee a y acids, lac ic acid, and
py u ic acid in cap i e ba s du ing no mal eeding cycles as
well as a e a "p olonged" as . The mos salien ea u es o
his s udy a e he cycles o blood glucose and ee a y acid
le els. He ound ha blood glucose le els ollow he expec ed
pa e n and all wi hin no mal concen a ions o mammals in
gene al. Mos o he ba s s udied died du ing 31-32 hou as s;
howe e , ba s ha su i ed showed no u he change in blood
glucose le el a 35.3 mg/100 ml. The diu nal pa e n o ee
a y acid (FFA) plasma le els ollows he expec ed gene al
in e se ela ion o blood glucose le els. Nigh ime FFA's a e
essen ially cons an a abou 0.5 milliequi alen s/li e (mEq/L),
which is qui e no mal o mammals. I ood is wi hheld o
h ee hou s a e sunse , plasma FFA concen a ions ise o 4.0
mEq/L concomi an wi h he all in blood glucose le el.
Al hough his ochemical s udies o he muscles o ba s
(A ms ong e al., 1977; Talesa a and Kuma , 1974) demon-
s a e he ela i e impo ance o a s and glucose as ene gy
subs a es, muscle enzyme p o ile s udies such as hose o
Mulle and Baldwin (1978) and, especially, hose o Yacoe e
al.
(1982), a e pa icula ly ge mane o he p esen discussion.
Yacoe e al. (1982) de e mined enzyme ac i i y le els o
ci a e syn he ase, hexokinase, 3-hyd oxyacyl-CoA dehyd oge-
nase (HOAD), and phospho ylase in wo ugi o ous species
(one o which was A ibeus li u a us) and eigh insec i o ous
species o ba s. The ou enzymes measu ed a e indi ec
indica o s (in sequence) o ci ic acid cycle capaci y, blood
glucose oxida ion capaci y, be a-oxida i e capaci y, and glyco-
genoly ic capaci y.
As expec ed, ci a e syn he ase ac i i y was ex emely high,
among he highes epo ed o mammalian skele al muscle,
and he e we e no in e speci ic di e ences. Ele a ed HOAD
ac i i y le els in all species indica ed he expec ed high
capaci y o a y acid oxida ion. Enzymes pa icipa ing in
glucose s o age, mobiliza ion, and cell en y, howe e , p o-
ided he mos in iguing pic u e. Hexokinase ac i i y in he
ugi o ous species was om wo o h ee imes highe han in
he insec i o ous species. Phospho ylase ac i i y in all species
was on he high end o he no mal mammalian ange, and,
al hough no s a is ically signi ican , phospho ylase ac i i y in
A.
li u a us was highe han in any o he species es ed.
F ugi o ous ba s, he e o e, e ain he capabili y o apidly
me abolizing a s as a uel sou ce bu also ha e unusually high
glycogenoly ic abili y. The combina ion o high ae obic and
high glycogenoly ic ac i i ies p e iously has been hough o
be mu ually exclusi e in mammalian muscle ibe (Bu leigh
and Schimke, 1969). Such a combina ion de ies easy classi ica-
ion in he slow (I) and as (IIA) and as (IIB) ca ego ies o
mammalian muscle ibe ypes (Lamb, 1984). Al hough simila
esul s ha e been epo ed o Aus alian ba s (Mulle and
Baldwin, 1978), ex emely high capaci ies o ae obic glucose
oxida ion ha e been epo ed p ima ily in ligh muscle o
insec s, which ha e a no mal die wi h high glucose densi y
(Beenakke s, 1969; Beenakke s e al., 1975; Hein ich, 1979).
Because o i s p esumed high die a y glucose densi y and
ex emely high glucose assimila ion e iciency (Mo ison,
1980b), A. jamaicensis, along wi h o he ugi o es and
nec a i o es, should ha e much mo e glucose a ailable o
oxida ion han do ba s o o he die a y p e e ences. Howe e ,
he e is a signi ican ene gy penal y o con e ing die a y
ca bohyd a e o a (Ma in and Lieb, 1979).
12SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
F om he o egoing, i may be p esumed ha A. jamaicensis
p oduces li le a om glucose and does no ake ad an age o
he educed weigh and high calo ic densi y gained by s o ing
ene gy as a . Fu he mo e, A. jamaicensis mus exhibi an
ex eme acili y in he s o age, mobiliza ion, and u no e o
glycogen. Finally, he o e all ene gy ese oi in A. jamaicen-
sis is se e ely limi ed. In hei na u al en i onmen , hese ba s
main ain no signi ican posi i e daily ene gy balance ha
would allow o s o age o su plus calo ic ene gy.
SALIVATION.—Like
many ugi o es, A. jamaicensis has
excep ionally la ge sali a y glands (Phillips e al., 1977).
Resul s o ou his ological examina ion o hese glands in A.
jamaicensis (S udie , Boyd e al., 1983) a e in ag eemen wi h
hose o Wimsa (1956). In spi e o he la ge size o he glands
he e is no ela i e inc ease in numbe o duc s, no is he e
e idence o ela i e excess p oduc ion o se ous sec e ion. The
unc ional signi icance o hese s uc u al obse a ions include
possible inc eases in sali a y amylase p oduc ion, inc eased
in ol emen in egula ion o mine al and wa e balance, and
o e all inc ease in p oduc ion o sali a, all ela ed o he
unusually la ge size o he glands.
Sali a y componen s may ac chemically o neu alize
alkaloids in igs. Mucus may hold oge he he pelle s ha a e
d opped du ing eeding (Dalques e al., 1952). We ha e
sugges ed ha he abundan sali a may s ongly bu e gas ic
sec e ions, hus p e en ing gas ic con en s om becoming
acidic, while simul aneously coa ing he gas ic epi helium
wi h an ex ensi e bu e ing ba ie (S udie , Boyd e
al.,
1983).
I gas ic luid emains abo e pH 4, sali a y amylase should
con inue o unc ion. This may be o conside able impo ance
in iew o he apid passage ime and consequen b ie
diges ion exhibi ed by A. jamaicensis (Mo ison, 1980b;
S udie , Boyd e al., 1983).
The gas ic glands in he s omach and duodenum o many
ugi o ous ba s con ain ypical pa ie al and zymogen cells
(Bhide, 1980; Fo man, 1972; Rouk and Glass, 1970). B un-
ne 's glands a e educed o absen in ugi o ous ba s,
including species o A ibeus. The unc ion o B unne 's glands
among mammals is s ill deba ed, bu o a long ime a
connec ion wi h p o ec ion o he duodenum om damage by
highly acidic chyme lea ing he s omach has been suspec ed. I
sali a o A. jamaicensis has su icien bu e ing capaci y o
p e en chyme om becoming highly acidic, educ ion o
absence o B unne 's glands in hese ugi o es would be
compensa ed. Sali a y bica bona e concen a ion has been
shown o
Be
di ec ly p opo ional o he a e o sali a o ma ion
(Bu gen and Emmelin, 1961). Because he bica bona e bu e
sys em accoun s o he bulk o he bu e ing powe o he
sali a (Izu su, 1981), inc eased o high ela i e a es o sali a
p oduc ion may indica e ele a ed sali a y bu e ing capaci y in
A.
jamaicensis. Fo u he discussion o
his
possibili y and o
de ails o gas ic ul as uc u e see Phillips e al. (1984).
Calo ic Balance
To al daily ene gy equi emen s can be es ima ed by a ious
me hods (Kunz and Nagy, 1988). Main enance ene gy (ME)
equi emen s a e mos simply calcula ed as a unc ion o body
mass (W). Fo endo he mic mammals, which main ain cons an
high Tb, ME (Kcal/day) = 106 W0-75, whe e W is in kilog ams
(Na ional Resea ch Council, 1978). Fo an A. jamaicensis
weighing 45 g, es ima ed ME equals 10.3 Kcal (43.3 Kj)/day
(Table 2-1). Daily ene gy equi emen s o an A. jamaicensis
weighing 45 g also can be es ima ed based on ime pa i ioning
(Table 2-1). The i s es ima e is based on Mo ison's (1978d)
da a, in which he pa i ioned ligh ime in o a ious ac i i ies
and ligh dis ances. Me abolic cos o ligh was based on he
wind unnel s udies o Thomas (1975) on o he ugi o ous
ba s whe e me abolic cos o ho izon al ligh o a 45 g ba is
0.30 Kj. A an a e age ligh speed o
5
m/sec, he cos o lying
100 m is 0.092 Kj. App op ia e inc eases in he ene gy cos o
ligh o anspo o igs o eeding oos s also a e included
(Mo ison, 1978d). These ligh cos s a e added o ene gy needs
o main aining he basal me abolic a e.
Using da a om McNab (1969) o well- o o e ed A.
jamaicensis (1.7 cc/g/h), we es ima ed a basal ene gy cos o
TABLE 2-1.—Daily ene gy expendi u es o a 45 g A ibeus jamaicensis. All
alues a e Kcal/days; equi alen Kj/day a e gi en in pa en heses. See ex o
u he de ails.
Pa ame e s
Based on mass*
Based on ime pa i ioning
Fligh cos s'"
sea ching
commu ing
be ween eeding passes
eeding passes
ansmi e e
sub o al
Basal me abolism (well- o o e ed)'1
To al**1
Basal me abolism (no mally- o unde ed)8
To al1"'
Based on ime pa i ioning
Fligh cos s
Basal me abolism (well- o o e ed)d
To als
Basal me abolism (no mally- o unde ed)*
To al 4«
Kcal/day
10.30
0.20
0.28
0.19
1.04
0.11
1.82
8.77
10.59
3.87
5.69
3.20
8.77
11.97
3.87
7.07
Kj/day
(43.30)
(0.84)
(1.17)
(0.79)
(4.33)
(0.50)
(7.63)
(36.70)
(44.33)
(16.20)
(23.83)
(13.40)
(36.70)
(50.10)
(16.20)
(29.60)
• Na ional Resea ch Council, 1978.
b Mo ison, 1978d.
c Addi ional 1% inc emen as cos o ca ying ansmi e in ligh .
d McNab, 1969.
e S udie and Wilson, 1979.
'Mo ison, 1980b.
NUMBER 51113
8.77 Kcal/day. Using me abolic a es o unde ed (= no mally
ed) A. jamaicensis (0.75 cc/g/h; S udie and Wilson, 1979),
educes basal ene gy cos o 3.87 Kcal/day. In ei he
ci cums ance, daily ene gy cos o basal me abolism ep esen s
he majo ac ion (67.9%-82.7%) o o al daily ene gy
equi emen s. An addi ional ime-pa i ioning ene gy budge
(Mo ison, 1980b) is based simply on a minimum es ima ed
ligh ime o 45 min/day (Table 2-1). Al hough his second
ime-pa i ioning budge ma kedly inc eases me abolic ene gy
expendi u e o ligh ( om 1.8 o 3.2 Kcal/day), basal
me abolic cos s s ill ep esen he majo ac ion (54.8%-
73.1%) o o al daily ene gy budge s. Kunz (1980) p oposed a
daily ene gy budge o ba s in gene al in which calo ic needs
(in Kcal/day) equal 0.92m0-767 (whe e m is mass in g ams).
Kunz's (1980) es ima e o daily ene gy needs, howe e ,
includes da a om lac a ing and p egnan ba s as well as om
ep oduc i ely inac i e indi iduals. The e o e, he o e es i-
ma ed calo ic needs.
The es ima es o o al daily ene gy cos s o a ep oduc i ely
inac i e, endo he mic A. jamaicensis, main aining a high Tb a e
ema kably consis en wi h a o al ange o 10.3-12.0 Kcal/day
(43.3-50.1 Kj/day) and p obably ep esen ealis ic es ima es
o a ba ha emains cons an ly homeo he mic. The minimal
daily ene gy budge o 5.7 Kcal/day (23.8 Kj/day) o
he e o he mic indi iduals is su ely an unde es ima e because
he ba s a e no con inuously he e o he mic. The ma ked
educ ion in daily ene gy demand associa ed wi h he e o h-
e my, howe e , may be in aluable o ee- lying indi iduals
ha ha e a ma ginal ene gy in ake. Fleming (1988) es ima ed
daily ene gy budge s o 41.9-47.3 Kj/day o Ca ollia
pe spicilla a, a smalle phyllos omid wi h a mo e a ied die
and di e en o aging s a egy. The simila i y be ween hese
igu es is s iking.
A.
jamaicensis ex ac s 55.5 g o juice pe 100 g o esh ui
o F. insipida (Mo ison, 1980b). Mo ison epo ed ha juice
om hese ipe igs con ained 0.315 Kcal/g; we ound 0.415
Kcal/ml (S udie , Boyd e al., 1983). The speci ic g a i y o an
a i icial ig juice solu ion (111 mg glucose pe ml) is 1.042
g/cc. Mos o he ene gy in ig juice is in dissol ed glucose and
he ene gy assimila ion e iciency o ig juice by A. jamaicensis
is 98.3% (Mo ison, 1980b). Using he maximum es ima e o
a daily ene gy budge o 12.0 Kcal/day, an A. jamaicensis
weighing 45 g would need o assimila e all he ene gy om
28.9 o 36.5 ml o ig juice. Gi en he 98.3% assimila ion
e iciency, his would equi e 29.4-37.2 ml o inges ed ui
pulp juice o 55.1-69.8 g o esh ipe ui . A 7 g pe ui , an
A. jamaicensis would equi e 8-10 whole ui s o
F.
insipida
pe day o mee i s calo ic equi emen en i ely om he
inges ion o he ui s o igs. I a e age-weigh ipe ui s
weighing 5.6 g (Mo ison, 1978a) we e inges ed, equi ed
in ake would be 9.8-12.5 igs. These numbe s o whole igs
co espond nicely o he 7 ± 2 nigh ly eeding passes obse ed
by Mo ison (1978a) when A. jamaicensis was eeding
exclusi ely on ui s o . insipida.
In summa y,
A.
jamaicensis has meage , i any, a ese es;
p obably exhibi s ex ao dina y glucose assimila ion, s o age,
mobiliza ion, and glycoly ic capaci ies; p obably has ex eme
daily luc ua ion in glycogen le els wi h li le ese e capaci y;
and is ma ginally able o main ain calo ic balance on a no mal
daily in ake o 7 ± 2 Ficus insipida ui s. Inges ion o nine
ui s p obably would allow
A.
jamaicensis o main ain a high
Tb o a 24-hou pe iod bu inges ion o ewe ui s would no
allow calo ic balance as a high Tb homeo he m. Such ba s
would conse e ene gy and hus emain in calo ic balance by a
d op in egula ed Tb, and a co esponding d op in ene gy needs.
Ni ogen Balance
Ni ogen exc e ion is ela ed o me abolic a e. Conse-
quen ly, ni ogen equi emen s a e app op ia ely ela ed o
me abolic body mass (W°-75) (B ody, 1945; Kleibe , 1975). The
daily ni ogen equi emen (mg/day) o high Tb homeo he ms
is 200 W075, whe e W is in kilog ams (Na ional Resea ch
Council, 1978). The ni ogen equi emen can be con e ed o
a minimum p o ein equi emen by mul iplying by 6.25
(He bs , 1988). Al e na i ely, die a y p o ein equi emen can
be calcula ed as a unc ion o inges ed calo ic in ake. Minimum
p o ein o main enance is 10.7 mg p o ein pe Kcal inges ed.
Fo a 45 g A. jamaicensis, he daily minimum p o ein
equi emen is 122 mg, based on body mass alone. Using
maximal es ima ed daily ene gy expendi u e (12.0 Kcal/day
om Table 2-1), calcula ed daily minimum main enance
p o ein in ake is 128 mg. The p o ein densi y o
F.
insipida ui
juice is 4.7 mg/ml (calcula ed om Mo ison, 1980b).
Assuming ig juice o be he only sou ce o die a y ni ogen o
A.
jamaicensis, cons an ly homeo he mic indi iduals main ain-
ing high Tb would equi e 26.0-27.2 ml o ig juice/day (a
100%
assimila ion) o main ain ni ogen balance. This es i-
ma ed equi ed in ake is less han he calcula ed daily olume
o ig juice needed o main ain calo ic balance in high Tb
homeo he mic indi iduals (29.4—37.2 ml/day).
Diges ibili y o p o ein in low- ibe die s in a a ie y o
mammals anges om 77% o 90% (Mayna d and Loosli,
1969).
Assuming an assimila ion o 85% o A. jamaicensis,
daily minimum in ake o ig juice would ise o 30.6-32.0
ml/day. This would mean ha he daily olume o ig juice
necessa y o mee p o ein needs is equal o o less han ha
needed o calo ic balance and ha main enance o ni ogen
balance is less o a p oblem o he ba s han main aining
calo ic economy. The calcula ed p o ein minimum, howe e , is
o he "ideal" p o ein whose amino acid composi ion exac ly
e lec s he needs o he subjec . Rasweile (1977) poin ed ou
ha animal p o eins gene ally ha e amino acid composi ions
ha co espond mo e closely o mammalian equi emen s and
may be mo e eadily diges ible han p o eins o plan o igin,
which o en a e incomple e in e ms o essen ial amino acids.
14SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
The calcula ed equi emen o ig juice olume gi en abo e
is su ely a minimum o low es ima e o ac ual juice needs. As
is he case o calo ic economy, i seems likely ha a high Tb
homeo he m such as A. jamaicensis could main ain ni ogen
balance only ma ginally on a die o F. insipida ui . Again, he
abili y o a ee-li ing
A.
jamaicensis o educe i s egula ed Tb
ac s as a sa e y al e, no only o calo ic balance, bu also by
educing die a y ni ogen equi emen s o a le el su icien o
allow ni ogen balance.
Al hough s udies o he s uc u e and unc ion o kidneys o
Neo opical ba s will be discussed in mo e de ail wi h espec o
wa e and mine al balance, some o he da a a e pe inen o
ni ogen balance. S udie , Wisniewski e al. (1983) examined
kidneys in 25 species o Neo opical ba s and sugges ed ha
enal mo phology is p ima ily a unc ion o die a y p o ein
densi y. Among he phyllos omids, membe s o he p ima ily
ugi o ous/nec a i o ous sub amilies Glossophaginae, Ca ol-
liinae, and S enode ma inae (including A. jamaicensis), mos o
which ha e low-p o ein die s, possess enal medullae ha
canno be subdi ided eadily in o inne and ou e zones.
Membe s o he phyllos omid sub amilies Phyllos ominae and
Desmodon inae, as well as all membe s hus a s udied in all
o he amilies o New Wo ld ba s, ha e enal medullae ha a e
easily subdi ided in o inne and ou e zones. The die a y
p e e ences o his second g oup o ba s a e a ied, bu all
species egula ly consume some ood o animal o igin, which
is high in p o ein.
Ou da a on u ine composi ion u he suppo he p oposed
ela ion be ween die a y p o ein densi y and enal unc ion in
Neo opical ba s (S udie and Wilson, 1983). We ound u ina y
ammonia and u ea ni ogen le els o
A.
jamaicensis (495 mg%
N),
al hough highly a iable, o be ma kedly lowe han le els
o he insec i o ous species, Myo is nig icans (1887 mg% N).
I he minimal ni ogen equi emen o a 45 g A. jamaicensis is
19.54 mg/day (= 200 x 0.045075), hen o al u ina y N is 434
mg/kg/day, which app oxima es u ina y ni ogen exc e ion
a es in many mammalian he bi o es (Al man and Di me ,
1961).
I he wo abno mally high alues o u ina y ammonia
and u ea ni ogen we ound in A. jamaicensis a e dis ega ded,
he ecalcula ed a e age le el is 370 mg% N (n = 17, S udie
and Wilson, 1983).
Wa e Balance
We now ha e su icien da a o p esen a ough wa e balance
accoun o A. jamaicensis (Table 2-2). Values in Table 2-2
ep esen es ima es o a 45 g, high Tb homeo he mic
indi idual. As p e iously discussed, daily in ake o ig juice is
29.4-37.2 ml. Swallowed juice is 90% wa e (Mo ison,
1980b). Daily in ake o wa e in ood, he e o e, is 27.6-34.9 g.
Es ima ed me abolic wa e assumes comple e ae obic oxida ion
o all glucose in he consumed juice (111 mg/ml, om S udie ,
Boyd e al., 1983). The calcula ed alue o me abolic wa e is
p obably sligh ly low because oxida ion o he small amoun s
TABLE 2-2.—Wa e economy budge o a 45 g, high Tb, homeo he mic
A ibeus jamaicensis. All alues a e g/day. EWL • e apo a i e wa e loss. See
ex o u he de ails.
Pa ame e sg/day
WATER GAINS
In ood
F om me abolism
D unk
To al gain
WATER LOSSES
EWL a es
EWL in ligh
In u ine
In eces
To al loss
UNACCOUNTED LOSSES
27.6-34.9
1.9-2.4
0
29.5-37.3
5.2
1.3
9.4-11.8
11.8-14.9
27.7-33.2
1.8-4.1
o die a y a s and p o eins will p oduce a li le addi ional
wa e . The e is no indica ion ha A. jamaicensis needs o
consume ee wa e when eeding on igs. The s udies o
Phillips e al. (1984) on gas ic ul as uc u e in A ibeus also
indica ed a lack o impo ance o ee wa e inges ion.
Assuming cons an empe a u e and humidi y, e apo a i e
wa e loss (EWL) a es is calcula ed as
log EWL = log 0.398 + 0.672 log W
whe e EWL is g ams o wa e /animal/day and W is in g ams
(S udie , 1970). We es ima ed EWL in ligh based on 45
minu es o al lying ime/day (Mo ison, 1980b) and scaling up
he EWL (957 mg/h) ound by Ca pen e (1969) o lying
Lep onyc e is sanbomi (24.4 g). Es ima es o u ina y wa e
loss and wa e los in ecal ma ix a e de i ed om Mo ison's
(1980b) da a showing ha u ina y wa e ep esen s 31.8% and
ecal wa e ep esen s 40.1% o swallowed ig juice, espec-
i ely. I all inges ed ni ogen appea ed in he u ine, A.
jamaicensis would p oduce u ine olumes o 4.0-5.3 ml/day,
assuming a minimum daily equi ed ni ogen in ake o 19.54
mg/day and u ina y ammonia and u ea ni ogen le els o
370-495 mg%. These alues a e conside ably lowe han
es ima es based on Mo ison's (1980b) da a, bu s ill ep esen
ema kably high u ine ou pu .
Conside ing he mul iple sou ces and assump ions used o
cons uc he wa e economy budge o A. jamaicensis (Table
2-2),
he alues come ema kably close o balancing and a e
p obably accu a e. Some use ul compa isons can be made wi h
alues o o he mammals. Wa e u no e a es o a 45 g A.
jamaicensis (655-829 ml/kg/day) a e ex eme in con as o
alues o 40-273 ml/kg/day epo ed o a wide a ie y o
mammals (Al man and Di me , 1961). Simila ly, p e o med
wa e consump ion in A. jamaicensis o 613-775 ml/kg/day is
ele a ed in compa ison wi h alues o o he mammals
(35-211 ml/kg/day; Al man and Di me , 1961). These
NUMBER 51115
excep ional alues ela e o he essen ially liquid die o A.
jamaicensis, coupled wi h i s poo enal wa e conse a ion,
and ela i ely small size. Compa a i e alues o o al body
u no e and p e o med wa e u no e a e 330 ml/kg/day and
150 ml/kg/day, espec i ely, o an 8.4 g Myo is hysanodes
(O'Fa ell e al., 1971), and 250 ml/kg/day and 180 ml/kg/day,
espec i ely, o M. luci ugus (O'Fa ell e al., 1971). To al
body wa e u no e in Pizonyx (= Myo is) i esi is es ima ed a
480 ml/kg/day (Ca pen e , 1968). P e o med wa e u no e o
a common ampi e, Desmodus o undus, weighing 34.2 g is
410 ml/kg/day (Wimsa , 1969), assuming bo ine blood o be
78.5%
wa e (McNab, 1973). P e o med wa e u no e also
can be es ima ed om published da a o he nec a i o ous ba ,
Lep onyc e is sanbo ni, a 465-712 ml/kg/day (Howell, 1974),
and he nec a i o ous bi d, Selaspho us lammula, a 692
ml/kg/day (Hainswo h and
Wol ,
1972).
U ina y ou pu in A. jamaicensis a ies om 209 o 262
ml/kg/day in con as o many o he mammals wi h alues ha
a y om 2.5 o 74 ml/kg/day (Al man and Di me , 1961).
These a es o u ine p oduc ion in A. jamaicensis (6.5-8.2
mic oli e s/min) a e oughly 30 imes he maximal a e o u ine
p oduc ion (0.23 mic oli c s/min) ound by Basse and
Wicbe s (1979) o he insec i o ous M. luci ugus. Ha ing
collec ed u ine samples om many species o
ba s,
we ha e no
di icul y belie ing ha a es o u ine p oduc ion in A.
jamaicensis and o he ugi o ous/nec a i o ous ba s a
exceed u ine olumes conside ed no mal in o he species.
As men ioned ea lie , A. jamaicensis and o he ugi o ous
and nec a i o ous ba s possess kidneys wi h undi ided enal
medullae. Such species ou inely p oduce na u al u ine o low
osmo ic p essu e and low u ina y ni ogen le els (S udie and
Wilson, 1983) compa ed wi h hose species wi h enal medulla
di isible in o inne and ou e zones (S udie , Wisniewski e al.,
1983).
Mean maximal u ine concen a ion (MMUQ in A.
jamaicensis is 972 mOsm/kg (S udie , Boyd e al., 1983). The
o al medulla y (M) hickness o co ical (C) hickness a io
(M/C) o A. jamaicensis (2.4), while ypical o ugi o es, is
ma kedly lowe han M/C o ba s o o he eeding p e e ences
and does no a y wi h habi a a idi y (S udie , Wisniewski e
al.,
1983).
Geluso (1980) p esen ed a highly p edic i e equa ion
ela ing MMUC o insec i o ous ba s o M/C in which MMUC
(mOsm/kg) = 702 + 387 (M/C). Based on his equa ion, A.
jamaicensis would p oduce MMUC o 1,620 mOsm/kg, a alue
nea ly double he obse ed MMUC. Simila obse a ions hold
ue o o he ugi o ous/nec a i o ous species wi h undi ided
enal medullae (Ca pen e , 1969; S udie and Wilson, 1983).
None o hese species p oduce na u al u ine concen a ions
app oaching hose p edic ed by Geluso's equa ion, sugges ing
ha his o mula does no apply o ba s wi h undi ided enal
medullae (S udie and Wilson, 1983).
We made se e al a emp s o induce MMUC in A.
jamaicensis, ha included dehyd a ion/s a a ion, loading wi h
s ongly hype osmo ic sal solu ions, and eeding dehyd a ed
ui s o igs (S udie , Boyd e al., 1983). Al hough none o
hese me hods wo ked well, inges ion o dehyd a ed igs, he
unc ional equi alen o Geluso's (1975, 1978) "wa e denied"
expe imen s wi h insec i o ous ba s, esul ed in he mos
uni o m and highes u ine concen a ions o his species. I is
o pa icula in e es ha al hough dehyd a ion and sal loading
caused an expec ed and p edic able inc ease in osmo ic
p essu e o he blood, hese ea men s we e no associa ed wi h
he expec ed inc ease in osmo ic p essu e o he u ine (S udie ,
Boyd e
al.,
1983). Dehyd a ion in Myo is luci ugus, induced by
Basse and Wiebe s (1979), howe e , esul ed in expec ed
inc eases o osmo ic p essu es in bo h blood and u ine. The e
seems o be a slow o minimal elease o an idiu e ic ho mone
(ADH) o a slow o educed enal ubula esponse o ADH in
esponse o ising osmo ic p essu e in he blood o A.
jamaicensis.
U ine samples aken om A. jamaicensis a sunse in May
we e signi ican ly mo e concen a ed and less a iable han
samples aken in No embe (S udie , Boyd e al., 1983). The
May samples we e aken a he end o he d y season and he
No embe sampling occu ed owa d he end o he we season
(Smy he, 1974). P esumably hea /dehyd a ion s ess is g ea e
in he d y season han in he we . In May, he e was a ma ked
apid dec ease in o al u ine concen a ion 0.5-1.5 hou s a e
sunse in ee- lying
A.
jamaicensis. This dec ease is associa ed
wi h apid ood passage ime in his species, inges ion o
adequa e hypo onic luid o ehyd a ion, and apid assimila-
ion and equilib a ion wi h he inges ed ig pulp juice. U ine
hen became p og essi ely mo e concen a ed h oughou he
emainde o he nigh . Osmo ic p essu es o u ine in cap i e,
ehyd a ed ( aken wo hou s a e sunse ) indi iduals we e
iden ical h oughou he nigh wi h u ine concen a ions o
ee- lying
A.
jamaicensis (S udie , Boyd e al., 1983). This
sugges s ha ehyd a ion occu s ea ly in he nigh ly eeding
pe iod and is una ec ed by subsequen eeding bou s. We
know om o he s udies ha A. jamaicensis eeds spo adically
h oughou he nigh (Mo ison 1978a; 1978b; 1978c).
Mine al Balance
How he bi o ous mammals inges adequa e amoun s o
die a y sodium (Na+) has gene a ed conside able in e es
(Blai -Wes e al., 1968; Cowan and B ink, 1949; Dalke e al.,
1965;
He be and Cowan, 1970; Jo dan e al., 1973; S ocks ad
e al., 1953; Weeks and Ki kpa ick, 1976; 1978). Sodium
le els in mos plan s and plan pa s a e ypically low (Likens
and Bo mann, 1970; Sauchelli, 1969; Weeks, 1978) and ela ed
o soil Na+ le els, which in u n a e highly a ec ed by he Na+
le els in ain all (Blai -Wes e al., 1968) and he equency o
ain all. T opical ain o es may be pa icula ly suscep ible o
loss o nu ien s by leaching due o apid decomposi ion o
li e and hea y, equen ains (Jo dan and He e a, 1981).
Al hough some da a on cha ac e is ics and composi ion o he
soils o BCI a e a ailable (Knigh , 1975), he e is no

16SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
in o ma ion on soil sodium le els.
Fig ui s con ain li le sodium (Diem, 1962; Heinz In e na-
ional Resea ch Cen e , 1964; Oa es, 1978). Sodium le els in
ipe ui s o F. insipida and
F.
yoponensis ha e been measu ed
a 0.49 and 0.48 mg/g d y weigh , espec i ely (Nagy and
Mil on, 1979). We de e mined sodium le els in igs ca ied by
A.
jamaicensis o be simila o hose ound by Nagy and Mil on
(1979),
bu po assium concen a ions we e lowe (S udie ,
Boyd e al., 1983). Sodium densi y in d ied pulp is abou 2.4
imes he sodium le el o d ied seeds and is iden ical o he
sodium le el o pulp
juices.
Thus, he consump ion o ig juice
a he han he en i e ui g ea ly inc eases die a y Na+ densi y
in A. jamaicensis and signi ican ly educes he weigh o igs
es ima ed o be equi ed o main enance o Na+ balance. On
he o he hand, he K+ le el o ig juice is less han hal he
concen a ion o ha ion in d ied pulp. This implies ha K+ is
concen a ed in speci ic o ganelles wi hin he ig pulp and is
no likely o be ex ac ed by A. jamaicensis. I is eleased in he
p ocess o homogeniza ion o he d ied pulp o labo a o y
analysis. The lowe ed die a y K+ le el ob ained om pulp
juices as opposed o whole pulp p obably also lowe s he Na+
equi emen s o A. jamaicensis as i does in o he mammals
(Meye e al., 1950; S aaland e al., 1980; Weeks and
Ki kpa ick, 1978).
Minimal sodium equi emen s ha e been es ima ed o ew
small mammals. Sodium equi ed o g ow h in labo a o y a s
and mice is es ima ed a
10
and 18 mg/animal/day, espec i ely
(Na ional Resea ch Council, 1978). Assuming he a e age
equi emen o g ow h in A jamaicensis is 14 mg/animal/day,
ba s would equi e 69 ml (72 gms) o
F.
insipida ui juice a
8.8 mEq/L sodium le el o main ain sodium balance. Such
amoun s a exceeds he equi emen s o ig juice p e iously
calcula ed o main enance o calo ic and ni ogen balance
(29.4-37.2 and 30.6-32.0 ml/day, espec i ely). Es ima ed
daily Na+ equi emen s o g ow h a e p obably highe han
equi emen s o main enance. Howe e , i would appea ha
acqui ing adequa e sodium o daily equi emen s om ig
juice is mo e likely o be a nu i ional limi ing ac o han
inges ion o su icien juice o mee calo ic and ni ogen needs.
Mammals su e ing Na+ de iciency cha ac e is ically exhibi
hype ophy and hype plasia o he zona glome ulosa o
ad enal glands and g ea e de elopmen o s ia ed and
exc e o y duc s wi hin sali a y glands (Blai -Wes e al., 1968).
These ea u es, ac ing h ough he enin-angio ensin-
aldos c one sys em, esul in enal and sali a y Na+ e en ion.
The ad enal zona glome ulosa o A. jamaicensis shows no
ob ious hype ophy o hype plasia when compa ed wi h he
condi ion in Neo opical insec i o ous ba s, al hough possible
sub le di e ences may ye be de ec ed (S udie , Boyd e al.,
1983).
We ha e no ound an inc ease in he numbe o duc s
wi hin sali a y glands, ano he ac ha a gues agains a
p obable Na+ de iciency in A. jamaicensis. The ino dina e size
o hese glands, howe e , indica es a p opo iona e inc ease in
absolu e numbe o duc s pe g am o body mass o he ba .
Ene gy and Wa e Balance du ing Lac a ion
Because he o egoing discussion has conce ned nu i ional
equi emen s and balances needed o main enance, i is use ul
o es ima e nu i ional inc emen s needed unde s ess such as
du ing lac a ion (see Sec ion 3, Rep oduc ion in a Cap i e
Colony). A 45 g
A.
jamaicensis should p oduce milk a a a e o
12.3 gm/day based on Linzel's (1972) measu e o milk
p oduc ion as a unc ion o body mass (daily milk p oduc ion in
Kg/day = 0.126075 Kg/Kg). Milk o
A.
jamaicensis con ains 2.3
Kcal/g (Jenness and S udie , 1976). This ene gy le el is
compa able o milk ene gy con en o o he ba s, bu is highe
han milk ene gy le els o many la ge mammals. The high
ene gy con en o A. jamaicensis milk is in acco d wi h Ben
Shaul's (1962) sugges ion ha mammals ha nu se hei young
on a scheduled basis p oduce milk o highe ene gy con en
han mammals ha nu se con inuously o on demand.
I one assumes ha ood ene gy can be con e ed o milk
ene gy wi h no cos , he p oduc ion o 12.3 g o milk pe day a
2.3 Kcal/g imposes a minimal addi ional calo ic equi emen
o A. jamaicensis o 28.3 Kcal/day. This would aise he o al
daily calo ic equi emen o abou 40 Kcal/day o a high Tb,
homeo he mic, lac a ing emale compa ed o 12.0 Kcal/day o
a nonlac a ing indi idual. The o al milk p o ein le el o A.
jamaicensis is 4.7 g% (1.1 g% casein and 3.6 g% whey p o ein)
(Jenness and S udie , 1976). Again assuming no ene gy cos s
o milk p o ein syn hesis, a high Tb, homeo he mic, lac a ing
emale would equi e an addi ional inc emen al daily p o ein
in ake o 578 mg o a o al daily p o ein equi emen o
700-706 mg compa ed o 122-128 mg/day o a nonlac a ing
indi idual. Milk o A. jamaicensis is abou 70% wa e ;
he e o e, p oduc ion o 12.3 g o milk would equi e an
addi ional in ake o 8.6 ml o wa e o a o al daily wa e need
o 38.1-45.9 ml in lac a ing emales compa ed o he
29.5-37.3 ml wa e equi emen s o nonlac a ing ba s.
Du ing lac a ion, he e o e, he addi ional inc emen al needs
o ex a calo ic and ni ogen in ake a e massi e in compa ison
o addi ional wa e needs. Whe eas es ima ed milk p oduc ion
may seem somewha high, ene gy e iciency o milk p oduc-
ion is ce ainly no 100%. B ody (1945) calcula ed g oss
ene ge ic e iciency o milk p oduc ion o be 28%-34% o
humans and 44%-48% o
a s.
We sugges ha du ing pe iods
o high nu i ional demands, such as g ow h and especially
p egnancy and lac a ion, A. jamaicensis would be expec ed o
lowe hei egula ed Tb o educe calo ic and ni ogen needs
and ha du ing such imes, hese ba s may supplemen hei
s aple ig die wi h ood i ems o highe calo ic and p o ein
densi y. S udies di ec ed a shi s in ood habi s du ing
p egnancy and lac a ion could es his hypo hesis.
Summa y
S udies on ene gy balance in A. jamaicensis we e ocused on
his ba 's use o he ui s o Ficus insipida, he ood ha i used
NUMBER 51117
mos on BCI. P obably hese ba s always a e in sligh ood
s ess in he wild and a e mo e likely o beha e as acul a i e
he e o he ms. Poo body empe a u e egula ion in newly
cap u ed ba s is no ela ed o physiological compe ence, bu
e lec s educed me abolic hea p oduc ion and a e o deple ion
o ene gy s o es. A. jamaicensis alls e y nea he minimum
bounda y cu e o high body empe a u e homeo he my,
sugges ing ha ene gy sa ings om he e o he my may be
c i ical o his ba in he wild.
A ibeus jamaicensis has ex emely low a ese es and a
high in ake o die a y ca bohyd a e, sugges ing ha glycogen
may be i s no mal ene gy ese e. This ba , in i s na u al
en i onmen , main ains no signi ican posi i e daily ene gy
balance ha would allow o s o age o su plus calo ic ene gy.
The ene gy cos o basal me abolism ep esen s he majo
ac ion (55%-83%) o o al daily ene gy equi emen s. Daily
ene gy budge s o 10-12 Kcal/day combined wi h he calo ic
con en o daily ood in ake equi es an in ake o 9.8-12.5 igs
pe day, a igu e no inconsis en wi h he 7 ± 2 nigh ly eeding
passes obse ed in adio eleme y s udies.
Ni ogen and p o ein equi emen s can be me by he amoun
o daily ood in ake necessa y o main ain calo ic balance,
al hough ques ions emain abou he diges ibili y o plan
p o eins. U ina y ammonia and u ea ni ogen le els a e
ma kedly lowe in A. jamaicensis han in ba s ha use animal
p o ein.
Based on ou s udies o
calo ic,
ni ogen, and wa e balances,
he e is no indica ion
ha
A.
jamaicensis needs o consume ee
wa e when eeding on igs. Wa e u no e a es o A.
jamaicensis (655-829 ml/kg/day) a e ex eme in con as o
alues epo ed o a wide a ie y o o he mammals. Mean
maximal u ine concen a ion in A. jamaicensis is 972 mOsm/
kg,
a s ikingly low alue indica ing poo u ine concen a ing
abili y. Published p edic i e equa ions o maximal u ine
concen a ion do no apply o ugi o ous ba s. Dehyd a ion in
A.
jamaicensis esul s in inc eased blood osmo ic p essu e, bu
wi hou a concomi an inc ease in u ine osmo ic p essu e.
Sodium is po en ially limi ing in opical animals es ic ed
o a ugi o ous die , bu s uc u al ea u es o he kidneys and
sali a y glands o A. jamaicensis a gue agains any p obable
ch onic sodium de iciency in hese animals. Du ing pe iods o
high nu i ional demands such as g ow h, p egnancy, and
lac a ion, A. jamaicensis p obably lowe s i s egula ed body
empe a u e o educe calo ic and ni ogen needs and may use
o he , mo e ene gy- and p o ein- ich ood esou ces.
3.
Rep oduc ion in a Cap i e Colony
Lucinda Keas
Ta
and Cha les
O.
Handley, J .
We epo on he ep oduc i e biology o A ibeus jamaicen-
sis based upon s udies o a colony held cap i e a he Na ional
Zoological Pa k (NZP). Al hough in o ma ion is a ailable on
many aspec s o he ep oduc ion o ba s, mos is based on
empe a e zone Vespe ilionidae. The mos ecen e iews o
p e-
and pos na al de elopmen among he espe ilionids a e
hose o O (1970) and Tu le and S e enson (1982). Racey
(1988) summa ized me hodology o ep oduc i e assessmen
and Wilson (1988) p o ided in o ma ion on main aining ba s
o cap i e s udies. The e is li le in o ma ion, howe e , on
ep oduc ion and on ogeny o o he ba s. Kleiman and Da is
(1979) e iewed he a ailable li e a u e on de elopmen and
ma e nal ca e in phyllos omids. The mos de ailed in o ma ion
on on ogeny is o Ca ollia pe spidlla a (Kleiman and Da is,
1979) and Desmodus o undus (Schmid and Manske, 1973).
Phyllos omid ba s show a g ea e di e si y o social
sys ems, die a y habi s, ep oduc i e s a egies, and selec ion o
oos si es han do he espe ilionids (Bake e al., 1976,1977,
1979).
The selec i e p essu es wi h which phyllos omids mus
cope a e qui e di e en om hose encoun e ed by he
be e -known espe ilionids o he empe a e zone. Compa i-
sons be ween he wo amilies should be p oduc i e.
Each yea mos emale A. jamaicensis gi e bi h o a single
young du ing wo ep oduc i e episodes. This pa e n o
bimodal polyes y is common o many Neo opical ugi o ous
and nec a i o ous ba s (Wilson, 1979). A. jamaicensis is
unusual, howe e , because de elopmen o he implan ed
blas ocys is delayed o 2.5-3 mon hs du ing one o he bi h
episodes (Fleming, 1971; Fleming e al., 1972). Delayed
de elopmen has been documen ed only in Panamanian
popula ions o A. jamaicensis, bu i is likely o p o e
widesp ead. Delayed de elopmen also occu s in a phylo-
gene ically and ecologically di e se a ay o ba s, Mac o us
(B adshaw, 1962), Miniop e us (Medway, 1971), Hipposide os
(Be na d and Mees e , 1982), and Haplonyc e is (Heideman,
1988).
Lucinda Keas Ta , Na ional Zoological Pa k, Smi hsonian Ins i u-
ion,
Washing on, D.C. 20560.
Cha les O. Handley, J ., Na ional Museum o Na u al His o y,
Smi hsonian Ins i u ion, Washing on, D.C. 20560.
Pa u i ion and neona al appea ance ha e been desc ibed
(Bha naga , 1978; Jones, 1945,1946), bu li le is known abou
ea ly de elopmen o he young. In Panama^ whe e he young
o A. jamaicensis usually a e bom in ee holes (Mo ison,
1975),
he s udy o ea ly de elopmen is di icul because many
ee hole oos s a e inaccessible and he ba s show a s ong
endency o dese a oos i dis u bed. The e o e, obse a ions
on cap i es is s ill he bes means o gaining in o ma ion on
ep oduc ion and de elopmen .
No ick (1960), who was success ul in b eeding and
long- e m main enance o
A.
jamaicensis in cap i i y, belie ed
ha eedom om handling was necessa y o success ul
ep oduc ion. Imp o ed a i icial die s (Rasweile and de
Bonilla, 1972) ha e enhanced he success ul main enance o a
numbe o Neo opical ugi o ous and nec a i o ous phyllos-
omids (G eenhall, 1976; Rasweile , 1975,1977; Rasweile and
de Bonilla, 1972; Rasweile and Ishiyama, 1973). Kleiman and
Da is (1979) ound ha cap i e Ca ollia pe spidlla a success-
ully b ed in spi e o he handling necessa y o weekly
examina ions o he adul s and young.
His o y o he Resea ch Colony a NZP
We es ablished a esea ch colony o A. jamaicensis a he
NZP in Washing on, D.C, wi h 24 ba s cap u ed in June 1978
nea Co ozal, Panama^ 35 km SE o Ba o Colo ado Island
(BCI).
We s udied ep oduc ion and de elopmen in his colony
om July 1978 h ough Augus 1981 and, consequen ly, we
can desc ibe se e al aspec s o he biology o A. jamaicensis in
conside able de ail.
Among he o iginal 24 ba s, h ee males and 12 emales we e
subadul s, bo n ea ly in 1978. Fou males and i e emales we e
adul s. The emales we e nei he lac a ing no ob iously
p egnan a cap u e. P io o he i s bi hs in cap i i y (Janua y
1979),
wo adul males and ou subadul emales died. One o
hese was a emale ha appa en ly did no adjus o he cap i e
eeding egime. On he 47 h day o cap i i y all ba s we e
wing-banded wi h plas ic bands and h ee days la e ano he
emale died, possibly o complica ions om banding-
associa ed auma. The o he ou ba s died because o
19
26SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 3-7.—Leng h
o
in e bi h in e als
in
he NZP colony
o
A ibeus
jamaicensis.
In e al
be ween
bi h g oups
i-n
Il-III
III-IV
i -
V-VI
Numbe o
indi idual
emales
7
6
14
13
13
Mean in e al
in days ( ange)
134.6(120-153)
162.2(132-184)
131.2(113-158)
190.9 (159-220)
120.2(112-137)
198.29
443.37
372.34
377.74
50.74
Compa ison*
X
Y
X
z
X
Sou ce
Be ween
Wi hin
To al
SS
40561.185
13388.740
53949.925
ANOVA (one-way)
d
MS
4 10140.296
48
52
F
36.354
P
< 0.001
* S uden -Newman-Keuls mul iple compa ison es Iden ical le e s deno e ha di e ences we e
no
signi ican
a
he 5%
le el.
In e als whe e delayed emb yonic de elopmen was p esumed
o
ha e occu ed.
P<0.01.
de elopmen occu ed
in
he
cap i e emales
a
app oxima ely
he same ime
i
occu ed
in
ee-li ing emales, hen
he
in e bi h in e als be ween BG-II
and
BG-III
and
be ween
BG-IV
and BG-V
would
be
expec ed
o be
longe han
in e bi h in e als whe e de elopmen p oceeded no mally.
These da a
and
in e bi h in e als
o
indi idual emales
be ween consecu i e bi h g oups (Table
3-7)
sugges ha
delayed emb yonic de elopmen
did
occu
in he
cap i es
bu
was
o
diminished du a ion. The a e age in e bi h in e al
o
no mal p ena al de elopmen would
be
abou 122 days
and
he
in e bi h in e al wi h delayed emb yonic de elopmen would
be abou
213
days acco ding
o
Fleming's (1971) da a
o
ee-li ing Panamanian A. jamaicensis.
Di e ences be ween in e als p esumed
o
ep esen no mal
ges a ions we e
no
signi ican ly di e en om
one
ano he
in
he
NZP
ba s
as
e ealed
by a
pos e io i es ing (Table
3-7).
The p esumed delayed in e als, howe e , we e signi ican ly
di e en om
he
no mal in e als
as
well
as
om each o he .
The o e all end
in he
colony's ep oduc i e ou pu
was
owa d
a
sho ening
o
he in e bi h in e als
and
ea lie onse s
o successi e bi h g oups han expec ed
i he
cap i es
had
emained synch onized wi h
he
wild popula ion.
O
he
wo
in e als whe e delayed de elopmen
was
expec ed, mean
leng h
o
he
second in e al (BG-IV
o
BG-V)
was
nea ly
30
days longe han ha
o
he i s , al hough
BG-V
s ill occu ed
app oxima ely h ee mon hs ahead
o he
Panamanian cycle.
Loss
o
some en i onmen al clue, such
as day
leng h, may ha e
impac ed
he
cycle
o
he
cap i e ba s.
Ju enile Mo ali y
Fo y- h ee babies died be o e weaning (Table 3-4). Fac o s
con ibu ing
o
dea hs we e
as
ollows
(by
age
g oups):
NEONATAL
PERIOD
(1s day; 14 dea hs).—Mo ali y was he
esul
o
p ema u e bi h, s ill bi h, congeni al de o ma ion,
and ma e nal neglec .
PREVOLANT PERIOD
(2-30 days o age; 17 dea hs).—This
pe iod
was
cha ac e ized
by
mo he s equen ly lea ing hei
in an s una ended,
o in he
company
o
o he young and/o
adul s. Mo he s we e
in
es us du ing
he
ea ly pa
o
his
pe iod
and
we e equen ly pu sued
by
males.
A
highe han
no mal densi y
o
males (ne e less han i e adul males)
may
ha e con ibu ed
o
inc eased le els
o
s ess
in
he colony.
The
g ea es pe cen age
o
in an mo ali y occu ed
a
his ime,
possibly
due
o
his s ess.
NEWLY VOLANT PERIOD
(31-60 days o age; 5 dea hs).—
Young we e newly olan ,
bu no ye
weaned. Ini ially, hey
emained
in he
oos , exe cised hei wings
and
pec o al
muscles,
and
accompanied hei mo he s
on
sho ligh s.
I
hey we e
no
s ong enough
o a
e u n ligh
o a
oos a ea
and could
no
ind
a
subs a e
up
which hey could c awl
o a
oos ,
o a
place high enough
o
ini ia e ano he ligh , hey
would weaken
and die
unless hey we e e ie ed
by
hei
mo he s
o
disco e ed
by
he
keepe s.
WEANING
PERIOD
(61-110 days;
7
dea hs).—Ju eniles we e
becoming mo e independen , weaning was nea ing comple ion,
and
he
mo he s we e s a ing
o
ejec hei young
by
chasing
hem om he oos . T ansi ion
o
subadul hood was beginning.
P eweaning mo ali y among in an s
o
wild-caugh emales
was 35.4% (Table 3-8). Much
o
his mo ali y was a ibu able
o
wo
emales
(F4 and F12) who
consis en ly ailed
o
ea
o sp ing (se en babies be ween hem
o BG-I o
BG-VI)
because
o
congeni al de ec s (e.g., skele al de o mi ies)
o
in es inal ncma ode in es a ions
in he
young. These emales
we e able
o
ea young success ully beyond weaning age when
hey we e emo ed om he colony
and
placed
in a
la ge ligh

NUMBER 51127
TABLE
3-8.—Co ela ion o in an and ju enile mo ali y and le el o ma e nal expe ience. Numbe s o in an
dea hs in a ious age g oups a e abula ed in an -by-c con ingency able wi h expec ed alues in pa en heses
(Choi, 1978). X1
=
20.697, = 4, P
<
0.001.
O igin and
ep oduc i e expe ience
o emales
Wild-caugh
Cap i e-bo n
mul ipa ous
p imipa ous
To al o colony
Numbe
o
bi hs
65
10
14
89
Numbe o dea hs
du ing p eweaning
<,
30 days
15(21.91)
3 (3.37)
12 (4.72)
30
31-110 days
8 (7.30)
1 (1.12)
1 (1.57)
10
Numbe su i ing
beyond weaning
110+days
42 (35.79)
6 (5.51)
1 (7.71)
49
Pe cen
p eweaning
mo ali y
35.4
40.0
92.9
44.9
Pe cen
pos weaning
su i al
64.6
60.0
7.1
55.1
TABLE 3-9.—Analysis o mo ali y among cap i e-bom ba s by sex. Numbe s o in an dea hs in a ious age
g oups a e abula ed in an -by-c con ingency able wi h expec ed alues shown in pa en heses (Choi, 1978). X1
= 5.860, d = 3, 0.25 > P > 0.10.
Sex
Males
Females
To al
To al
bi hs
45
38
83
To al
dea hs
23
11
34
<.
7 (4.34)
1 (3.66)
8
Age in days
2-30
10 (8.67)
6 (7.33)
16
31-110
6 (5.42)
4 (4.58)
10
Su i o s
110+
22 (26.57)
27 (22.43)
49
cage wi h ewe ba s. Thus, hey may ha e been in poo
nu i ional condi ion om excess s ess o social exclusion
om ood, ac o s ha could ha e a ec ed hei o sp ing's
g ow h in u e o and pos pa um de elopmen Excluding he
young bo n o emales F4 and F12 would educe mo ali y o
ju eniles o wild-caugh mo he s o abou 27%.
P eweaning mo ali y among o sp ing o mul ipa ous
(expe ienced) cap i e-bo n mo he s was 40%, a ibu able
en i ely o he dea hs o all young bo n du ing BG-VI. We ha e
no explana ion o his ca as ophe. In con as , app oxima ely
64%
o he young bo n o p imipa ous (inexpe ienced) emales
did no li e mo e han a day, and nea ly 86% did no su i e a
mon h. Six y pe cen o mo e o he young o expe ienced
mo he s (wild-caugh emales and mul ipa ous cap i e-bo n
emales) su i ed beyond weaning, bu ew (7%) o he young
o p imipa ous emales su i ed ha long.
Appa en ly, p eweaning mo ali y and pos weaning su i-
o ship a e dependen on he mo he 's le el o ep oduc i e
expe ience (Table 3-8). Dea hs o young we e no e enly
dis ibu ed be ween he sexes (Table 3-9). In ou colony, males
died a a a e g ea e han expec ed and emales died a a a e
lowe han expec ed, bu he di e ences we e no signi ican .
Lowe concep ion a es and inc eased in an mo ali y
among p imipa ous cap i e-bo n emales may be a ibu able o
a a ie y o causes. In na u al popula ions, young A. jamaicen-
sis p obably a e o ced o dispe se om hei na al oos be o e
bi h o he nex young, and hus pa en ing beha io would no
be acqui ed h ough lea ning. Despi e p oximi y o expe ienced
emales a he NZP, inexpe ience may ha e ope a ed o inc ease
mo ali y o in an s bo n o ou cap i e, p imipa ous emales.
Inc eased mo ali y due o inb eeding could also ha e been
in ol ed, because he inexpe ienced emales could ha e ma ed
wi h hei male pa en , male siblings, o o he ela ed males.
Howe e , he subsequen success ul ep oduc ion in hese
emales sugges s ha his was no an impo an ac o .
Kleiman (1980) discussed se e al physiological and beha -
io al means whe eby ep oduc ion may be supp essed in
socially subo dina e emale mammals in cap i e si ua ions.
Abo ions, s illbi hs, inadequa e ma e nal ca e, and one
appa en case o dep essed lac a ion we e he causes o dea h
o se e al o
he
i s young bo n o ou cap i e- ea ed emales.
In Panama and Mexico, g oups o 4-11 adul emale A.
jamaicensis accompanied by hei ju enile o sp ing and a
single adul male, occupied ee hole oos s, whe eas oliage
oos s (palm onds and subcanopy ees) we e occupied by
soli a y males o small g oups o subadul s (Mo ison, 1979).
T ee hole oos s p obably a e p e e ed o e oliage oos s by
ep oducing emales, as hey p o ide g ea e p o ec ion om
ain, p eda o s, and luc ua ions in empe a u e (see Sec ion 8,
Roos ing Beha io ). While o aging, emale A. jamaicensis
usually lea e hei young in he oos s (Fen on, 1969).
T une and Slobodchiko (1976) ound ha clus e ed
An ozous pallidus we e less agi a ed and had less weigh loss,
lowe oxygen consump ion, smalle body empe a u e-ambien
28SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
empe a u e di e en ials, los less hea o he en i onmen , and
conse ed mo e me abolic ene gy while inac i e, han ba s
oos ing indi idually. Any o all o hese ac o s could be
impo an o p ena al and pos na al de elopmen .
Ou cap i e-bo n emale A ibeus jamaicensis i s p oduced
young while hey we e s ill in nonclus e ing "subg oups." The
implica ion is ha hese emales we e no in he app op ia e
social en i onmen (ha em) necessa y o in an de elopmen .
Fo some ee-li ing popula ions, ee hole oos s may be
c i ical esou ces o ep oduc ion and he clus e o ba s wi hin
he oos s may be i al as a sou ce o "helpe s" o main ain he
app op ia e he mal condi ions o he young ba s. These
ac o s migh explain he ailu e o ou p imipa ous cap i e
emales o success ully ea o sp ing.
Neona al Physiognomy
Be ween 29 Janua y 1979 and 30 Ap il 1981, a leas 78
ull- e m ba s we e bo n a
NZP.
Unless o he wise no ed in he
desc ip ions ha ollow, obse a ions on neona es e e o 37 o
hese in an s on hei i s day o li e. Neona es we e i s
examined a he beginning o he da k pe iod and we e
p obably om 1 o 14 hou s old when i s seen. All neona es
e ained a segmen o d ied umbilical co d. F equen ly, bi s o
d ied amnio ic memb ane we e p esen on hei bodies as well.
By he second day, he co d and all d ied memb anes we e
gone, excep o wo in an s ha s ill e ained umbilical co ds.
All neona es had hei eyes open, and mos had ea pinnae
and noselea e ec when hey we e i s examined. A ew had
he pinnae and he noselea la ened agains he head, bu hey
became e ec wi hin six hou s. Ea openings usually we e
appa en and he neona es wi ched hei pinnae o bodies in
esponse o sounds. Six babies did no show ea openings o
espond o sounds un il as la e as ou days ollowing bi h.
Howe e , all in an s examined had he abili y o emi ul asonic
sounds in he 80 kHz ange on hei i s day. A. jamaicensis
emi hei ul asonic o ien a ion calls h ough he nose (G i in,
1958),
and, acco dingly, he mou hs o in an s we e closed
when hey p oduced ul asonic emissions. Thei nos ils and he
g oo es a he base o he noselea on he ou e edges o he
nos ils qui e ed isibly du ing such ocaliza ions.
The neona es usually we e co e ed spa sely wi h da k g ay
u on he do sum and op o he head. The ea s, muzzle, wings,
and en e we e hai less. Non u ed skin usually was pink.
Howe e , a ew neona es had da k g ay skin.
The ee o neona es we e well de eloped and disp opo ion-
a ely la ge. The wings appea ed sho , na ow, and ela i ely
less de eloped compa ed o he o he wise p ecocial physical
condi ion o he neona es. Bones o inge s and join s we e so
and lexible. The bones and skin o he wing ips we e
unpigmen ed and anslucen , and he ip o digi III o en
cu ed in owa d he body mo e han 180 deg ees.
A e age physical measu emen s o neona es we e aken on
22 indi iduals, each o which su i ed a leas h ee weeks.
Males and emales did no di e signi ican ly in mass, o ea m
leng h, wing span leng h, o wing a ea, al hough a e age size o
emale neona es exceeded ha o males (Table 3-10).
Phyllos omid neona es a e compa a i ely la ge a bi h, wi h
neona e- o-mo he mass a ios usually exceeding 0.25 and
o ea m a ios exceeding 0.41 (Kleiman and Da is, 1979;
Figu es 3-2 and 3-3; Table 3-11). A. jamaicensis compa es well
wi h hese obse a ions, wi h a neona e- o-mo he mass a io o
0.26 and a o ea m a io o 0.54 (Table 3-10).
Neona al Beha io
Newbo n young in he zoo colony ypically we e ound
hanging beside hei mo he s wi h mou h g asping one o he
mo he 's nipples. Young hus a ached kep hei wings olded
and did no appea o use hei humbs o clinging. O en, a
mo he shielded he in an , a leas pa ially, wi h a wing. I was
a e o ind a neona e wi h i s ee also a ached o i s mo he
while es ing quie ly in he oos . On he o he hand, i was no
unusual o ind newbo n in an s hanging alone in he oos .
They hung quie ly, wi h bo h ee a ached o he ceiling, hei
wings olded, and o en had hei eyes closed. They appea ed o
be sleeping.
Neona es we e able o c awl abou in a slow and wobbly
TABLE 3-10.—Body mass and measu emen s o wings o neona e A ibeus jamaicensis in he NZP colony.
Va iable
Mass (g)
Fo ea m leng h (mm)
Wing span (mm)
Wing a ea (cm2)
Wing loading (g/cm2)
Aspec a io
(wing span2/wing a ea)
Female, N = 12
mean
14.1
33.9
235.8
77.2
0.187
7.41
SD
2.100
2.749
18.195
18.865
0.025
0.869
Male.W
mean
13.7
33.0
227.5
71.6
0.198
7.49
= 10
SD
1.655
2.088
15.501
17.666
0.034
1.252
Combined,
mean
13.9
33.5
232.0
74.7
0.192
7.45
N = 22
SD
1.878
2.453
17.159
18.120
0.029
1.034
*
0.513
0.811
1.144
0.715
0.825
0.168
P
ns
ns
ns
ns
ns
ns
Sex
a io
(F/M)
1.031
1.026
1.036
1.078
0.944
0.990
Pe cen
adul
size
0.264
0.542
0.504
0.276
1.118
0.942
* S uden 's /- es o signi ican di e ences be ween he means o males and emales (one- ailed).
NUMBER 51129
1.2-1
HI
i-
<
z
o
w
z
o
o
0.4-
0.4-
D
0.6 1.2 1.8
LOG ADULT MASS (g)
FIGURE 3-2.—The ela ionship be ween neona e mass and adul mass o 18
species o ba s. Log neona e mass (g) is plo ed agains log adul mass (g). The
eg ession line o espe ilionids (solid ci cles) is y = -0.60 + 0.93 x, ( =
0.942, P < 0.001). Phyllos omid (open iangles) and molossid (open squa es)
da a poin s a e shown o compa ison. Da a a e om Table 3-11.
ashion and we e ac i e in he ea achmen p ocess upon
eunion wi h hei mo he s. They showed a s ong endency o
keep hei heads downwa d. When a mo he was held in a
head-up posi ion, he a ached neona e eadjus ed i s posi ion
wi h i s ee un il i g asped i s mo he 's head. Tu ning he
mo he head-down caused he baby o esume he no mal
head-down pos u e. Neona es placed on a e ical wi e mesh
su ace wi h hei heads up immedia ely s a ed o in e . They
also ended o c awl upwa d (achie ed by "walking" backwa d
wi h he ee ). Olde babies ended o c awl upwa d un il
s opped by an obs acle o lack o a oo hold, bu neona es
usually c awled upwa d only a ew cen ime e s and hen
s opped.
Au og ooming in A. jamaicensis consis s o licking he body
su aces and aking o sc a ching body su aces wi h one oo
while hanging by he o he . Neona es we e seen licking
hemsel es, especially hei wing su aces, bu hey ne e hung
by one oo in o de o use he o he oo o g oom.
Allog ooming is known in his species o include he g ooming
o in an s by hei mo he s. Occasionally an in an ex ended a
wing o i s mo he as i o solici g ooming, and he mo he hen
licked he baby's wing. Mo he s equen ly sni ed young o he
E
E
HI
CC
o
U_
LJ
<
Z
o
LU
o
o
1.60n
1.25-
Q90-
1.41.61.8
LOG ADULT FOREARM (mm)
FIGURE 3-3.—The ela ionship be ween neona e and adul o ea m leng hs o
19 species o ba s. Log neona e o ea m (mm) is plo ed agains log adul
o ea m (mm). The eg ession line o espe ilionids (solid ci cles) is y = -0.24
+ 0.88 x, ( = 0.879, P < 0.001). Phyllos omid (open iangles) and molossid
(open squa es) da a poin s a e shown o compa ison. Da a a e om Table 3-11.
han hei own and o en huddled wi h hem, bu hey licked
only hei own young. We ne e obse ed allog ooming
be ween adul s a he NZP, bu i mus occu occasionally in
wild adul A. jamaicensis. Tha would bes explain he chewed
necklace bands we no ed in some indi iduals ha we e
cap u ed on BCI.
When we sepa a ed a neona e om i s mo he 's nipple, i
appea ed anxious and was quick o es o e a hold as wi h i s
mou h, bu i o en was no disc imina o y abou he subs a e
i happened o g asp. As hey we e being examined he
newbo ns equen ly succeeded in bi ing ou inge s, and on
wo occasions babies emained a ached and began o suck
igo ously. By he second day pos pa um, he babies we e
mo e disc imina ing abou wha hey would g asp wi h hei
mou hs. In o de o a oid alling, hey would bi e ou Finge s
when p e en ed om clasping ou hands wi h hei humbs and
hind ee , bu none ied o suck he skin. In an s we e ne e
ound a ached o emales o he han hei own mo he s.
Dis u bed mo he s ca ied hei babies wi h hem in ligh .
30
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
£2
Si
I1*
III
PJ
"1 „ £ ^ „ o o •» g s ^l-aS^S^^"0
«
-
2.
"51~.
^
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1
-
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"-
-
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I
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Z
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Q*o^oou:ao^^SJ2^a:&.SSQaaa
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NUMBER
51131
Appa en ly he babies ins an aneously de ached hei ee om
he ceiling as he mo he sp ead he wings o ake o . When a
mo he wi h a baby a ached o one o he nipples was ne ed in
ligh , he in an usually was ound in a posi ion pa allel o i s
mo he s's body, i s ee g ipping he emu o inguinal egion.
Ra ely, an in an was ound in a c osswise pos u e such as ha
no ed by Kleiman and Da is (1979) o Ca ollia pe spicilla a.
In his case, he in an 's ee we e a ached o he opposi e
nipple egion and he baby was ca ied ac oss he mo he 's
ches jus behind he h oa .
G ow h and De elopmen o Young
GROWTH
OF
HAIR.—Follicula
ac i i y p eceding g ow h o
u caused he colo o he skin o change om he neona al
pink o g ay. Skin o he unde pa s da kened a ound se en
days o age, and hai g ow h began a 12 days. A his ime,
young also began o g oom hemsel es wi h hei ee . The
spa se, app essed u on he head and do sum was e ec and
s a ing o hicken on day
15.
The muzzle a ea da kened a ound
day 17, and acial u g ow h s a ed abou day 20. By day 22
65-i
60-
^
55-
E
E
o
z
UJ
50H
45-
UJ
§
40-
u.
35-
30J
0
25
50
AGE
(days)
75
100
FIGURE
3-4.—G ow h o o ea m in he NZP colony o
A ibeus
jamaicensis.
Daily
means,
s anda d
e o s,
and
anges
a e
shown.

32
SMITHSONIAN
CONTRIBUTIONS
TO ZOOLOGY
he en um was co e ed wi h spa se u . The gene al and
supe cilia y ib issae we e only ed spo s abou he co ne s o
he mou h and eyes un il day 25, when hey began o p o ude.
The ea e , sho , so ib issae we e also e iden . Young had
ull pelage by 30 days o age and whi ish acial s ipes we e
appa en on some indi iduals.
BODY
SIZE.—G ow h
cu es o he ou measu ed cha ac-
e s ( o ea m leng h, wing span, wing a ea, and mass) a e
p esen ed in Figu es 3-4 o 3-7. Daily means, s anda d e o s,
and anges a e shown o 22 young om BG-I and II. Da a
de i ed om de o med o ma kedly unde de eloped young
ha died wi hin a ew days o bi h we e no included in he
compu a ions.
Fo ea m leng h showed he g ea es de elopmen a bi h
ela i e o o he measu ed cha ac e s and he as es g ow h a e
(X = 0.9 mm/day du ing he maximum linea g ow h phase;
Figu e 3-4). Fo ea m g ow h s abilized a ound 50 days o age
a X = 61 mm, 1 mm less han he a e age o ea m leng h o
ba s o he o iginal adul colony (Tables 3-1 and 3-12).
Wing span inc eased by X = 5.8 mm pe day and wing a ea
by X = 3.8 cm2 pe day du ing he maximum g ow h phase, and
bo h eached adul p opo ions in X = 70 days (Figu es 3-5 and
3-6). Body mass showed he slowes a e o inc ease (-0.5 g
pe day; Figu e 3-7) and ook nea ly 80 days o s abilize a X
= 48 g, which was 5 g less han he a e age adul body mass o
500-
E 400-
E
z
Q.
o
z
300-
2OOJ
o
25
I
50
AGE
(days)
— -
75
100
FIGURE
3-5.—G ow h o wing span in he NZP colony o A ibeus
jamaicensis.
Means,
s anda d
e o s,
and
anges
a e
shown.
NUMBER 51133
TABLE 3-12.—A e age asymp o ic measu emen s by sex o A ibeus jamaicensis in he NZP colony.
Va iable
Mass (g)
Fo ea m (mm)
Wing span (mm)
Wing a ea (cm2)
mean
48.4
61.7
466.7
264.73
Females
SD
3.462
1.377
10.630
18.155
n
20
19
15
7
mean
47.6
60.7
457.1
257.77
Males
SD
2.923
1.635
91.400
8.789
n
18
18
14
6
*
0.077
2.058
2.600
0.853*
Pi
ns
<
0.025
<0.01
ns
Sex a io
(F/M)
1.017
1.016
1.021
1.027
* S uden 's /- es o signi ican di e ences be ween he means.
One- ailed.
$ The a iances o measu emen s o male and emale wing a ea we e no homoscedas ic, so a Wilcoxon
wo-sample es was also pe o med (U = 25, ns).
300-1
250-
o
AREA
O
z
200
150
100-
50 J
0[
25l
50
AGE
(days)
75100
FIGURE 3-6.—G ow h o wing a ea in he NZP colony o A ibeus jamaicensis. Means, s anda d e o s, and
anges a e shown.
34SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
he colony's o iginal membe s (Tables 3-1 and 3-11).
Sexual dimo phism was no as appa en in he asymp o ic
size o he cap i e- ea ed young as i was in he wild-caugh
ba s.
Again, emales exceeded males in all measu ed a iables,
bu he di e ences we e signi ican only o o ea m leng h and
wing span. The magni ude o he di e ences be ween he sexes
was also educed, and emales exceeded males in size by only
2%-3%
(Table 3-12). The a iable bes co ela ed wi h age
was mass ( = 0.9387, P < 0.0001; whe e is he co ela ion
coe icien and P is p obabili y). All size a iables we e highly
co ela ed wi h one ano he . The a e age asymp o ic masses o
19 cap i e-bo n young we e compa ed wi h hei neona al
masses, and a signi ican ela ionship was ound ( = 0.7097,
, = 4.1459, P <
0.001;
Figu e 3-8). La ge in an s ended o
become la ge adul s.
INFANT-MOTHER
SIZE
RATIO.—The e was a
endency
o
la ge emales o p oduce la ge in an s (Figu e 3-9), bu he
ela ionship was no signi ican (n = 12; = 0.552; 0.10 > P >
O)
50-
45-
40-
35-
30-
25-
20-
15-
1OJ
o
2550
AGE
(days)
751OO
FIGURE 3-7.—G ow h o mass in he NZP colony o A ibeus
jamaicensis.
Means, s anda d
e o s,
and anges a e
shown.
NUMBER
511
35
a
CO
5
50H
o
o
46-
a.
2
>-
<
42-
10
15
NEONATAL MASS (g)
20
FIGURE 3-8.—The ela ionship be ween he asymp o ic mass
o
cap i e- ea ed young and hei neona al mass.
The equa ion o he eg ession line is
y =
32.94
+
1.05 x, ( = 0.710, P
<
0.001).
1.25-1
>
<
5
UJ
z
o
UJ
o
o
1.15-
1.05-
1.68 1.70 1.72 1.74 1.76 1.78
LOG FEMALE NON-GRAVID MASS
(g)
FIGURE
3-9.—The ela ionship be ween neona e and mo he mass in he NZP
colony
o
A ibeus jamaicensis.
A
mean nong a id mass was de e mined
o
each adul emale. Means (solid ci cles) and anges (ba s)
o
in an mass
a e
shown. The equa ion o he eg ession line
is y =
-0.14
+
0.74 x, ( = 0.552,
0.10<P<0.05).
0.05). Howe e , ela i e in an size
(as a
pe cen age
o he
mo he 's nonp egnan mass) was co ela ed wi h he mo he 's
po en ial weigh -bea ing capaci y (wing a ea/nong a id mass).
A posi i e ela ionship be ween ela i e neona al mass and
he mo he 's wing weigh -bea ing capaci y
has no
been
eco ded
o
any o he species
o
ba . Howe e , Kunz (1974)
no ed ha emale wing size was la ge han male wing size
in
Ep esicus uscus
in
Kansas whe e he no mal li e is one.
He
specula ed ha
i
he e
is
any selec i e ad an age gained
by
la ge wing size, he deg ee
o
sexual dimo phism in
E.
uscus
should be e en g ea e in he mo e eas e n s a es whe e wo
is
he usual li e size.
Mo e a en ion should
be
paid
o
species' wing-loading
alues
and
o he a iables
o
wing shape
and
ae odynamic
abili y
and
hei ela ionships
o a
a ie y
o
de elopmen al,
beha io al, physiological,
and
ecological ac o s. Wings
a e
essen ial o he loca ion, pu sui , and anspo
o
ood. O he
po en ial unc ions
o
wings ha also could
be
wing-size
dependen a e hea dissipa ion, e apo a i e wa e loss, ans-
po o shielding o young, and beha io al displays.
The mass
o a
de eloping e us mus
be o
pa icula
signi icance
o
hose ba s ha o age while lying. Howe e ,
beyond
a
ce ain poin
in
ela i e size,
he
la ge wings
hemsel es would ep esen
a
signi ican weigh load wi h
much inc eased d ag and would p obably be qui e di icul
o

4.
Rep oduc ion on Ba o Colo ado Island
Don E. Wilson, Cha les
O.
Handley, J .,
and Al ed
L.
Ga dne
Da a on ep oduc ion in ba s ha e accumula ed a an
accele a ing a e du ing he pas 25 yea s (Racey, 1982;
Wilson, 1979). Because A ibeus jamaicensis is common and
widesp ead i s ep oduc i e cycle is be e known han hose o
mos o he Neo opical ba s. Howe e , in-dep h s udies a a
single locali y o e long pe iods, which a e essen ial o
in e p e ing isola ed pieces o in o ma ion and o elucida ing
de ails o a ep oduc i e cycle, ha e been lacking.
We augmen ed ou unde s anding o ep oduc ion in A.
jamaicensis, gained om ou colony a he Na ional Zoological
Pa k (NZP), by ga he ing in o ma ion om popula ions on
Ba o Colo ado Island (BCI) and icini y. Ou da a on
ep oduc ion, accumula ed o e se e al annual cycles om
wild-caugh ba s, demons a e a high deg ee o ep oduc i e
synch ony in his species.
In 1971 Fleming p esen ed e idence ha emb yos o A.
jamaicensis unde go e a ded de elopmen du ing pa o he
yea , and Fleming e al. (1972) demons a ed bimodal polyes y
o be he basic ep oduc i e pa e n in his species. Ou la ge
da a se om BCI and he adjacen mainland, oge he wi h he
esul s o he s udy o ou NZP colony (see Sec ion 3,
Rep oduc ion in a Cap i e Colony) p o ide us he oppo uni y
o examine his cycle in some de ail. We base ou ou line o he
ep oduc i e cycle o A. jamaicensis on 4447 indi idual
emales cap u ed be ween 1972 and 1980 on BCI and icini y.
The Basic Pa e n
We con i m he p e iously pos ula ed pa e n o bimodal
polyes y (Fleming e al., 1972; Wilson, 1979), and Fleming's
(1971) s a emen s on delayed emb yonic de elopmen (Figu e
4-1,
Table 4-1). In gene al, emales a e palpably p egnan in
Janua y and begin o gi e bi h by la e Feb ua y o ea ly Ma ch.
They a e so synch onized ha no mally mo e han hal o he
Don E. Wibon, and Cha les O. Handley, J ., Na ional Museum o
Na u al His o y, Smi hsonian Ins i u ion, Washing on, D.C. 20560.
Al ed L. Ga dne , NERC, US. Fish and Wildli e Se ice, Na ional
Museum o Na u al His o y, Washing on, D.C. 20560.
popula ion gi es bi h in hose wo mon hs. Du ing a
pos pa um es us mos o hem become p egnan again. This
esul s in a b ie pe iod when indi iduals a e bo h p egnan and
lac a ing. Palpably p egnan , bu s ill lac a ing indi iduals a e
mos likely in Ap il, May, o June, be ween he Feb ua y-
Ma ch and July-Augus bi h peaks.
Young bo n du ing he ini ial bi h peak (Feb ua y and
Ma ch) a e weaned in Ap il and May. The second cycle o
bi hs commences in July and Augus wi h he same pa e n o
pos pa um es us and lac a ion. The young om he second
bi h peak a e hen weaned in Sep embe and Oc obe . The
di e ence be ween he wo pe iods o ep oduc i e ac i i y
occu s when he blas ocys om he second pos pa um es us
implan s, bu does no de elop a he same a e as he p eceding
one.
The pe iod o delayed emb yonic g ow h co e s app oxi-
ma ely h ee mon hs, and is ollowed by a pe iod o essen ially
no mal de elopmen du ing he emaining ou .
We we e able o ecognize indi iduals ha we e p egnan
( e minal six weeks), lac a ing, o pos lac a ing, and as hese
e en s acked one ano he o e he yea s o ou s udy, we
eco ded peak imes o each e en . Some a ia ion om yea
o yea is ob ious (Figu e 4-1), bu he amoun o cong uence
om one yea o he nex is s iking.
Ano he way we summa ized he da a was o combine he
in o ma ion om all yea s in o a single "a e age" yea (Figu e
4-2).
Thus summa ized and di ided in o 3-week in e als, hese
a e ages show he bimodal cycle and emphasize he 12-week
diapause be ween July and No embe .
Indi idual Va ia ion
In spi e o he g ea numbe o ma ked ba s in ou sample,
and e en hough we sampled almos con inuously o h ee
yea s,
we we e unable o accumula e a comple e eco d o an
indi idual emale's ep oduc i e his o y h oughou he li e-
ime.
To do so, we would ha e o cap u e he same emale a
leas wice pe yea du ing he bi h pe iods. We did ca ch some
indi iduals wo o mo e imes each yea , bu no always a
app op ia e s ages o ep oduc ion. Al hough ou da a a e no
based on comple e indi idual ep oduc i e his o ies, i seems
43
44SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE
4-1.—Rep oduc i e condi ion (by week) o adul emale A ibeus jamaicensis on BCI, 1975 h ough 1980.
Week
1975
To al
1976
To al
1977
To al
1978
9
10
11
12
10
11
12
13
41
42
43
44
45
46
47
48
27
28
29
30
31
32
33
34
35
36
37
38
39
40
41
42
43
44
45
46
47
48
49
50
51
52
1
2
4
5
6
7
8
9
10
11
12
13
14
15
16
17
18
19
20
21
22
24
26
27
No
ep oduc ion
N
12
14
8
1
35
27
3
1
2
16
38
51
9
27
35
28
37
274
0
3
3
2
0
0
1
3
10
5
15
5
1
1
8
33
13
9
7
16
2
1
10
53
21
69
291
101
24
2
4
16
6
4
1
0
5
1
3
5
5
0
0
0
0
0
2
0
1
4
1
%
75.0
87.5
34.8
33.3
51.9
50.0
2.8
5.9
84.2
97.4
89.5
64.3
96.4
92.1
100.0
100.0
0.0
9.4
6.3
15.4
0.0
0.0
14.3
27.3
43.5
45.5
65.2
63.3
25.0
100.0
28.6
51.6
40.6
58.3
53.8
80.0
100.0
50.0
100.0
84.1
55.3
94.5
99.0
70.6
100.0
80.0
66.7
40.0
6.9
4.3
0.0
8.6
1.8
13.6
9.6
17.9
0.0
0.0
0.0
0.0
0.0
40.0
0.0
33.3
18.2
20.0
P egnan
N
4
2
11
2
19
25
3
35
32
0
0
0
0
0
0
0
0
95
1
19
13
2
0
0
2
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
7
16
2
62
1
9
0
1
8
9
54
21
25
16
20
3
2
1
0
0
0
1
0
0
0
2
7
1
%
25.0
12.5
47.8
66.7
48.1
50.0
97.2
94.1
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
100.0
59.4
27.1
15.4
0.0
0.0
28.6
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
11.1
42.1
2.7
1.0
26.5
0.0
20.0
33.3
60.0
93.1
91.3
78.1
27.6
35.7
13.6
3.8
3.6
0.0
0.0
0.0
12.5
0.0
0.0
0.0
66.7
31.8
20.0
Lac a ing
N
0
0
4
0
4
0
0
0
0
2
1
6
3
0
1
0
0
13
0
5
14
4
3
1
0
1
6
2
2
0
0
0
3
1
2
0
0
0
0
0
0
1
1
1
47
0
0
0
0
0
0
0
0
7
35
30
14
31
14
10
1
3
0
0
1
0
0
4
0
%
0.0
0.0
17.4
0.0
0.0
0.0
0.0
0.0
10.5
2.6
10.5
21.4
0.0
2.6
0.0
0.0
0.0
15.6
29.2
30.8
42.9
100.0
0.0
9.1
26.1
18.2
8.7
0.0
0.0
0.0
10.7
1.6
6.3
0.0
0.0
0.0
0.0
0.0
0.0
1.6
2.6
1.4
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
21.9
60.3
53.6
63.6
59.6
50.0
55.6
50.0
30.0
0.0
0.0
20.0
0.0
0.0
18.2
0.0
Young
on ea
N
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
%
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
4.5
0.0
P egnan &
lac a ing
N
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
2
0
%
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
().()
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
9.1
0.0
Pos -
lac a ing
N
0
0
0
0
0
0
0
0
0
1
0
0
2
1
2
0
0
6
0
5
18
5
4
0
4
7
7
4
6
1
3
0
17
30
17
7
6
4
0
1
0
2
0
1
149
0
1
0
0
0
0
0
1
0
2
5
2
14
8
8
1
7
7
2
2
1
0
4
3
%
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
5.3
0.0
0.0
14.3
3.6
5.3
0.0
0.0
0.0
15.6
37.5
38.5
57.1
0.0
57.1
63.6
30.4
36.4
26.1
16.7
75.0
0.0
60.7
46.9
53.1
43.8
46.2
20.0
0.0
50.0
0.0
3.2
0.0
1.4
0.0
2.9
0.0
0.0
0.0
0.0
0.0
4.3
0.0
3.4
8.9
9.1
26.9
28.6
44.4
50.0
70.0
87.5
100.0
40.0
100.0
0.0
18.2
60.0
To al
16
16
23
3
58
52
6
36
34
19
39
57
14
28
38
28
37
388
1
32
48
13
7
1
7
11
23
11
23
6
4
1
28
64
32
16
13
20
2
2
10
63
38
73
549
102
34
2
5
24
15
58
23
32
58
56
22
52
28
18
2
10
8
2
5
1
3
22
5
NUMBER
51145
TABLE
4-1.—Con inued.
Week
28
29
30
31
32
33
43
44
45
46
47
48
49
50
51
52
To al
1979 1
2
3
4
5
7
9
10
11
12
13
14
15
16
17
18
19
20
21
22
23
24
26
29
36
37
38
39
40
41
42
43
44
45
46
47
48
49
'50
51
To al
1980 1
2
3
4
5
6
7
8
9
13
14
15
No
ep oduc ion
N
0
1
3
2
0
1
11
4
14
13
35
16
45
7
21
59
417
28
13
8
2
2
1
8
4
3
3
7
0
4
9
4
1
3
3
4
2
1
2
0
1
12
36
34
26
9
43
36
19
11
90
95
48
5
27
70
1
677
8
59
33
6
3
17
2
17
1
1
10
3
%
0.0
100.0
9.7
100.0
0.0
14.3
91.7
80.0
82.4
86.7
92.1
94.1
95.7
100.0
95.5
98.3
96.6
100.0
29.6
22.2
33.3
33.3
47.1
20.0
3.0
18.8
15.6
0.0
3.8
7.4
6.9
100.0
25.0
25.0
10.5
3.9
3.7
3.7
0.0
9.1
52.2
56.3
72.3
78.8
69.2
74.1
92.7
95.0
100.0
100.0
94.1
98.0
83.3
93.1
98.6
100.0
88.9
96.7
97.1
85.7
75.0
94.4
50.0
63.0
100.0
2.4
8.9
2.2
P egnan
TV
3
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
185
0
0
19
7
4
2
2
3
4
0
1
0
5
11
7
0
1
5
24
38
18
39
2
1
0
0
0
0
0
0
1
0
0
0
4
1
0
2
0
0
201
1
2
1
1
1
1
2
10
0
1
6
18
%
75.0
0.0
3.2
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
70.4
77.6
66.7
66.7
11.8
15.0
4.0
0.0
2.2
0.0
4.8
9.1
12.1
0.0
8.3
41.7
63.2
74.5
66.7
72.2
66.7
9.1
0.0
0.0
0.0
0.0
0.0
0.0
2.4
0.0
0.0
0.0
4.0
2.0
0.0
6.9
0.0
0.0
11.1
3.3
2.9
14.3
25.0
5.6
50.0
37.0
0.0
2.4
5.4
13.2
Lac a ing
N
1
0
16
0
3
4
0
1
0
0
0
0
0
0
0
0
175
0
0
0
0
0
0
6
10
89
12
31
2
63
71
26
0
0
2
7
2
7
13
1
9
6
6
4
3
0
1
0
0
0
0
0
0
1
0
0
0
372
0
0
0
0
0
0
0
0
0
35
82
84
%
25.0
0.0
51.6
0.0
50.0
57.1
0.0
20.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
35.3
50.0
89.9
75.0
68.9
100.0
60.0
58.7
44.8
0.0
0.0
16.7
18.4
3.9
25.9
24.1
33.3
81.8
26.1
9.4
8.5
9.1
0.0
1.7
0.0
0.0
0.0
0.0
0.0
0.0
16.7
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
83.3
73.2
61.8
Young
on
ea
N
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
%
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
1.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
P egnan
&
lac a ing
N
0
0
1
0
0
0
0
0
0
0
0
0
0
0
0
0
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
3
0
0
0
0
0
0
0
0
0
0
0
0
%
0.0
0.0
3.2
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
7.9
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Pos -
lac a ing
N
0
0
10
0
3
2
1
0
3
2
3
1
2
0
1
1
97
1
0
0
0
0
0
1
3
3
1
6
0
32
30
21
0
8
2
0
9
1
0
0
0
5
22
9
4
4
14
2
1
0
0
2
0
0
0
1
0
182
0
0
0
0
0
0
0
0
0
5
14
31
%
0.0
0.0
32.3
0.0
50.0
28.6
8.3
0.0
17.6
13.3
7.9
5.9
4.3
0.0
4.5
1.7
3.4
0.0
0.0
0.0
0.0
0.0
5.9
15.0
3.0
6.3
13.3
0.0
30.5
24.8
36.2
0.0
66.7
16.7
0.0
17.6
3.7
0.0
0.0
0.0
21.7
34.4
19.1
12.1
30.8
24.1
4.9
5.0
0.0
0.0
2.0
0.0
0.0
0.0
1.4
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
11.9
12.5
22.8
To al
4
1
31
2
6
7
12
5
17
15
38
17
47
7
22
60
878
29
13
27
9
6
3
17
20
99
16
45
2
105
121
58
1
12
12
38
51
27
54
3
11
23
64
47
33
13
58
41
20
11
90
101
49
6
29
71
1
1436
9
61
34
7
4
18
4
27
1
42
112
136
46SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE
4-1.—Con inued.
weeK
16
18
19
20
21
32
33
34
35
36
37
38
39
40
41
42
43
To al
No
ep oduc ion
N
0
0
9
2
7
8
15
9
13
5
26
36
14
33
59
38
6
440
%
0.0
0.0
32.1
8.3
38.9
61.5
27.3
13.8
23.2
14.3
40.0
64.3
66.7
64.7
73.8
88.4
100.0
P egnan
N
0
2
1
13
7
0
0
0
0
0
0
0
0
0
0
0
0
67
%
0.0
66.7
3.6
54.2
38.9
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Lac a ing
N
2
0
3
7
0
4
36
20
9
8
13
3
1
1
2
0
0
310
%
50.0
0.0
10.7
29.2
0.0
30.8
65.5
30.8
16.1
22.9
20.0
5.4
4.8
2.0
2.5
0.0
0.0
Young
on ea
N
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
%
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
P egnan &
lac a ing
~N
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
0
%
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
0.0
Pos -
lac a ing
N
2
1
15
2
4
1
4
36
34
22
26
17
6
17
19
5
0
261
%
50.0
33.3
53.6
8.3
22.2
7.7
7.3
55.4
60.7
62.9
40.0
30.4
28.6
33.3
23.8
11.6
0.0
To al
4
3
28
24
18
13
55
65
56
35
65
56
21
51
80
43
6
1078
0)
CO
#•*
*
mm
9
o
k.
o
Q.
90 -
80-
70-
60 -
50 -
40 -
30-
20-
10-
1977
n I
V
'•
:
l
I
•
"•
•
•
1978
a
1 *.
l
••
•:<-
V"?i
/
•' ' hi,
•••
1
I
A
/
/
i
5
/
1979 '
I
! K1
» Mi1
1 T !
11 1 «
i i i ii '
•'
• 1
!
/I/M
h
1980
#
n
n
n
11
i i
i i
i i
•
1SA
• I
V.
•
*
•
*»
:
•
*.
'.
10 20 30 40 50 8 18 28 38 48 6 16 26 36 46 4 14 24 34 44
Weeks
FIGURE 4-1.—Pe cen ages o p egnan (solid line), lac a ing (dashed line), and pos lac a ing (do ed line) emale
A ibeus jamaicensis on BCL Each
eco d
ep esen s
ou weeks combined.
NUMBER 51147
70-
60-
50
H
O
O)
« 40-
c
©
o
S 30H
Q.
20-
10-
Jan Feb Ma Ap May Jun Jiil Aug Sep Oc No Dec
FIGURE 4-2.—A e age pe cen age o p egnan A ibeus jamaicensis on BCI (based on da a om 1972 h ough
1980,
pooled in h ee week inc emen s).
clea ha no e e y emale akes pa in e e y b eeding pe iod.
The e a e eco ds om each week o he yea o emales
showing no e idence o ep oduc i e ac i i y (Table 4-1). A
he peak o each ep oduc i e episode, he numbe o p egnan
emales a e ages abou 70% (Figu e 4-2).
Fo emales o iginally banded as ju eniles o subadul s, we
ha e 894 cap u e eco ds in he yea ollowing hei bi h. O
hese, 547 o sligh ly mo e han 60% we e eco ded as
non ep oduc i e (nei he palpably p egnan , lac a ing, no
pos lac a ing). F om a sample o 1121 cap u e eco ds o adul
emales (indi iduals ha had expe ienced an ea lie p eg-
nancy),
449 o 40% we e non ep oduc i e. Du ing any gi en
sampling pe iod, he e a e almos always mo e ep oduc i ely
ac i e adul s han he e a e yea lings (Figu es 4-3 and 4-4). A
summa y g aph (Figu e 4-5) combining da a om 1977
h ough 1980 shows ha yea lings no only b eed less
equen ly, hey a e mo e likely o be ou o synch ony wi h he
ep oduc i ely expe ienced adul s.
The Rep oduc i e Cycle
ESTRUS.—Each
emale has wo pe iods o pos pa um es us
pe yea . The i s occu s immedia ely ollowing he bi h in
Feb ua y-Ma ch; he second immedia ely ollowing he bi h in
July-Augus . Thus, he es ous cycle is easily con olled by he
iming o pa u i ion excep in hose emales ha , o wha e e
eason, ail o gi e bi h du ing a pa icula ep oduc i e pe iod.
We do no know wha physiological and en i onmen al ac o s
cue es us a he app op ia e
ime.
Pe haps es us in emales ha
ailed o become p egnan is esynch onized by con ac wi h

48
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
0
O>
(0
*1
9
O
h.
<D
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100-
90-
80-
70-
60-
50
40 -
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1977
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ILL
i
1980
i /
' i /
/
10 20 30 40 50 8 18 28 38 48 6 16 26 36 46 4 14 24 34 44
Weeks
10 20 30 40 50 8 18 28 38 48 6
Weeks
16 26 36 46 4 14 24 34 44
NUMBER
51149
105
Jan
Feb Ma Ap May Jun Jul Aug Sep Oc No Dec
FIGURE 4-5.—A e age pe cen ages
o
p egnan yea ling (solid line)
and
olde (dashed line) emale A ibeus
jamaicensis
on BCI,
based
on
da a om 1977-1980 combined. Each eco d ep esen s
one
week.
adul emale oos ma es who a e ollowing a no mal cycle.
COPULATION.—Males
ha e enla ged es es du ing he
pe iods when emales a e unde going pos pa um es us (see
Sec ion 3, Rep oduc ion in a Cap i e Colony). The deg ee o
asynch ony in he sys em indica es ha males may be capable
o insemina ing emales o ela i ely long pe iods o e lapping
bo h bi h peaks.
EMBRYONIC
DEVELOPMENT.—Fe iliza ion
and
implan a-
ion ollow a no mal sequence du ing each o he ep oduc i e
episodes, bu subsequen de elopmen di e s be ween he wo.
The no mal ges a ion pe iod ollowing implan a ion du ing he
i s ep oduc i e episode o he yea is 3.5-4 mon hs.
FIGURE
4-3
(opposi e, op).—Pe cen ages
o
p egnan yea ling (solid line)
and
olde (dashed line) emale A ibeus jamaicensis
on
BCI. Each eco d ep esen s
ou weeks combined.
FIGURE
4-4
(opposi e, bo om).—Pe cen ages
o
non ep oduc i e yea ling
(solid line)
and
nonyea ling (dashed line) emale A ibeus jamaicensis
on BCI.
Each eco d ep esen s ou weeks combined.
Implan a ion du ing he second episode is ollowed by delayed
emb yonic de elopmen , which esul s in a ges a ion pe iod o
abou se en mon hs (Fleming, 1971). This is a unique ea u e
o he ep oduc i e cycle o A. jamaicensis. The cues a e
unknown, bu because he cycle pe sis s when animals a e
mo ed in o cap i i y unde en i onmen al condi ions unlike
he na u al ones (see Sec ion 3, Rep oduc ion in a Cap i e
Colony), i mus be unde some gene ic con ol. E en ually, he
synch oniza ion b eaks down unde cons an condi ions o
cap i i y, sugges ing ha one o mo e en i onmen al cues a e
necessa y o ese he sys em.
PARTURITION.—The
wo peaks o pa u i ion a e in Feb u-
a y-Ma ch and July-Augus . The neona es a e well-enough
de eloped o be able o hang by hemsel es wi hin a ew hou s
o bi h (see Sec ion 3, Rep oduc ion in a Cap i e Colony).
Only on wo occasions did we cap u e a emale wi h an
a ached young. Appa en ly, he young a e le behind sho ly
a e bi h while he emales o age.
LACTATION.—Lac a ion
is easily de ec ed and las s o abou
wo mon hs (see Sec ion 3, Rep oduc ion in a Cap i e Colony).
50
SMITHSONIAN
CONTRIBUTIONS
TO
ZOOLOGY
I
| | | I I I | I I I
I jl
I I I I I I I I I I
I ( I
I I I I ' I I ' I I
' | '
' ' ' ' I ' I I I
I
I
1977 1978 1979 1980
O
CO
O
Q.
50 •
25-
0 -
25
50 •
O
(0
O)
o
50
25
0
25
50
50
25
0
25
50
60
40
J FMAMJJASONDJ FMAMJJ ASON
DJ
FMAMJJ ASON
DJ
FMAMJ
J
ASOND
1977 1978 1979 1980
FIGURE
4-6.—Schema ic
ep esen a ion
o he ep oduc i e cycle o A ibeus jamaicensis on BCL showing
empo al a ia ion
in
pe cen ages
o
p egnan ,
lac a ing,
and pos lac a ing emales. Mean
ain all
is shown in he
ba
g aph
a he
bo om.
Time is in
mon hs
and yea s.
Peaks in pe cen ages o lac a ing emales ollow abou a mon h
behind he peaks o p egnan emales (Figu es 4-1 and 4-6).
Lac a ion in cap i i y las s a bi longe in emales ha a e
ca ying do man emb yos ollowing he second b eeding
episode. This phenomenon can be in e ed in ou wild
popula ion (Figu e 4-6).
POSTLACTATION.—We
we e able o glean addi ional in o -
ma ion abou he cycle om indi iduals we ecognized as
pos lac a ing. Appa en ly he e is mo e indi idual a ia ion in
he leng h o ime ha nipples a e classi ied as pos lac a ing
han in o he ep oduc i e c i e ia (Figu es 4-1 and
4-6).
Pa o
his a ia ion may be an a i ac o s ages o e inemen o ou
de ini ion o pos lac ion. Ea ly in he Ba P ojec , we p esumed
a ba o be pos lac a ing i no milk could be exp essed om
nipples al hough he nipples we e la ge, labby, and su ounded
by naked skin. La e , he ca ego y was expanded o include
emales showing new hai g owing in a ound he eg essing
nipple, ega dless o i s size.
NUMBER 51151
Seasonali y, Rain all, and Abundance o Food
P e ious hypo heses (Wilson, 1979) ha e ocused on
ene ge ics as he p ima y con ol on ep oduc ion, and his
indeed seems he mos logical explana ion o he basic pa e n
o he cycle in A. jamaicensis. O he ac o s being equal, one
would expec animals o achie e maximum ep oduc i e ou pu
wi hou sac i icing su i o ship o he young.
I we assume ha he physiological cons ain s ope a ing on
an A. jamaicensis esul in a li e size o one and a ges a ion
pe iod o 3.5-4 mon hs, hen in a wo ld o abundan ene ge ic
esou ces, emales simply should ep oduce con inually,
p oducing up o h ee young pe
yea .
Tha hey do no sugges s
ha he e mus be a ime when ene gy sou ces a e in sho
supply.
The ime o maximum s ess migh occu a any o se e al
c i ical poin s in he ep oduc i e cycle. P egnancy, as does
lac a ion, ca ies wi h i an inc ease in ene gy demands o
indi idual emales (see Sec ion 2, Physiology; S udie e al.,
1972).
Rep oduc i e success should be enhanced i hese
s ess ul pe iods coincide wi h pe iods o peak esou ce
a ailabili y (B adbu y and Veh encamp, 1977b).
The c i ical ime o mos ba s is when he young a e weaned.
No only do he g owing young ha e high ene gy demands,
hey a e o ced o leam o o age a he same ime, and
p esumably i would be easie o do so when ood is eadily
a ailable. Readily a ailable ood esou ces also a e c i ical a
he ime o weaning in a species showing synch onized
ep oduc i e cycles, because he popula ion size is ma kedly
inc eased a ha ime. The e is now a conside able body o
e idence suppo ing he posi ion ha ep oduc i e e en s in
ba s a e imed o synch onize wi h seasonally abundan ood
esou ces (Dine s ein, 1983; Hei haus e al., 1975; Racey,
1982).
Young bo n in he i s ep oduc i e episode (Feb ua y-
Ma ch) a e weaned in Ap il-May, and hose bo n in he second
pe iod (July-Augus ) a e weaned in Sep embe -Oc obe . This
pa e n is con i med by ou cap u e eco ds, as he numbe o
ju eniles and subadul s inc eases d ama ically du ing hese
pe iods. On BCI his esul s in a gene al pa e n in which he
young a e weaned du ing he ainy season (Figu e 4-6). I he
ba s concei ed du ing he second episode we e o de elop
no mally (in 3.5-4 mon hs), hey would be bo n in No embe
and weaned a he beginning o he d y season in Janua y.
Fos e (1982) has shown ha he peaks in ui a ailabili y on
BCI occu in Ap il-May and Sep embe -Oc obe , wi h a low
in Janua y a he beginning o he d y season (Figu e 4-7).
The e o e, i seems ha delayed de elopmen has e ol ed as a
mechanism o a oid weaning young du ing he d y season
when ui ing ees a e sca ce. Thus, he ep oduc i e cycle o
A.
jamaicensis seems o be an adap a ion o he seasonal cycle
o ood abundance. E idence om o he locali ies suppo s his
hypo hesis (Wilson, 1979).
O
<
u.
Aug Sep Oc No Dec Jan Feb Ma Ap May Jun Jul
FIGURE 4-7.—Schema ic ep esen a ion o seasonali y o ui ing ac i i ies o canopy ees on BCI. "Ac i i y"
ep esen s he p opo ion o species ha a e ui ing (de e mined by seed aps). Red awn om Fos e (1982, ig.
12).
s
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62SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
LU
>
UJ
i
O
z
40-
30-
20
10-
40
30
20
10
UJ
O 40
oc
UJ
Q.30
20
10
M
A=c Sp ing •= 9
A=c Fall O= 9 Fall
a
I
3I
4I
5
I
6i
8
HALF YEARS AFTER MARK
FIGURE 5-5.—Compa ison o su i o ship o seasonal coho s o A ibeus jamaicensis ma ked on BCI and
adjacen mainland om 1976 o 1980; a = adul s, b = subadul s, and c = ju eniles.
NUMBER
511
63
basis
and,
al hough p oducing impo an in o ma ion
on
mo emen s, hese ba s did no o m
a
majo pa
o
he main
popula ion we we e s udying. When he A. jamaicensis om
hese o -island si es and hei ecap u e eco ds a e emo ed
om he da a (p esen ed
in
Table 5-9), he eby es ic ing he
analysis
o
ba s ma ked
on
BCI and O chid Island, su i al
es ima es a e highe and a y om a low
o
17% o adul males
ma ked
in he
i s hal 1980
o a
high
o
71% o ju enile
emales ma ked in he sp ing
o
1978 (da a om Table 5-10).
JUVENILES.—O e all
su i o ship in o he second hal yea
o li e by ju enile A. jamaicensis was 31% (Figu e 5-6, Tables
5-3 and 5-4). Su i o ship o ju eniles anged om 25%
o
males ma ked in he alls
o
1977 and 1979 o 57% o emales
ma ked
in
he sp ing
o
1978 (Table 5-3). Su i o ship om
combined da a om
he
all
o
1977
o he
sp ing
o
1980
sugges s ha app oxima ely 68%
o
he ju enile coho is los
by he ime ju eniles en e hei second hal yea (Tables 5-3,
and 5-4). We had an icipa ed high mo ali y du ing he i s
six
mon hs
o
li e based
on
s udies
o
empe a e-zone ba s
(B enne , 1968; Da is, 1967; Fos e
e a ,
1978; Humph ey
and Cope, 1970,1976, 1977; Keen and Hi chcock, 1980; Mills
e al., 1975; Pea son
e
al., 1952; S e enson and Tu le, 1981;
Tu le and S e enson, 1982). Su i o ship om he second
o
six h hal yea a e ma king among ba s ma ked as ju eniles
a e ages lowe han ha
o
adul s and subadul s (Figu e 5-6).
Dispe sal du ing
he
i s yea
o
li e
is he
mos likely
explana ion o his highe obse ed "mo ali y" o disappea -
ance a e. Ne e heless, ac o s such
as
dea h, lea ned
ne
a oidance, and loss
o
necklace (which ende s he indi idual
un ecognizable), as well as dispe sal away om he a ea o ou
s udy, con ibu e o he disappea ance
o
hese ba s. Su i o -
ship by he end o he second hal yea o li e (second hal yea
a e ma k) a e aged 17% (Figu e 5-6, Tables 5-3 and 5-4).
The ea e , he a e
o
decline pa allels ha
o
subadul s and
adul s.
Necklace loss may be g ea e in ju eniles because we had o
i
ba s
o
his age class wi h adul -size necklaces. Howe e ,
necklace loss canno explain
he
mo e apid decline
in he
second hal yea
o
li e
by
which ime he ba s ha e a ained
nea adul size. Su i o ship by he end o he second hal yea
o li e (second hal yea a e ma k) a e aged 17% (Figu e 5-6,
Tables 5-3 and 5-4). The ea e , he a e o decline pa allels ha
o subadul s and adul s.
We examined he numbe
o
ba s caugh
in
each hal yea
e sus he numbe known o be ali e
in
ha hal yea (Figu e
5-7, Table 5-11) o unde s and he p obable causes o he mo e
apid decline om
he
i s
o
second hal yea
o
li e
in
ju eniles. The consis en ly highe pe cen age
o
males caugh
o hose known
o be
ali e
in he
i s hal yea a e being
ma ked e lec s an ini ially highe ecap u e a e among males
a
HI
DC
40 n
I- 30
a.
o
LJJ
OC
20
LU
O
CC
LJJ
Q.
10 -
M
A ibeus jamaicensis
O= Ju eniles
A= Subadul s
•
=
Adul s
O
2 3 4 5 6
HALF YEARS AFTER MARK8
FIGURE 5-6.—Su i o ship cu es
o
he h ee age classes o A ibeus jamaicensis ma ked on BCI and adjacen
mainland om 1976 o 1980.

SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
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NUMBER 51165
LU
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LU
Q.
80-
60-
40-
20
80
60-
40-
• o
o
• c
80-
60-
40
20-F S F S F
1977 1978 1979
o •
F F S F S
1976 1977 1978 1979
FIGURE 5-7.—Compa ison o numbe s o A ibeus jamaicensis ma ked on BCI and adjacen mainland and
ecap u ed in he i s and second hal yea s a e ma k. Values plo ed a e ecap u es exp esssed as pe cen ages
o he numbe s known o be ali e in he same hal yea , open ci cles = emales, closed ci cles = males. Adul s,
(a) i s hal yea and (b) second hal yea o li e; subadul s, (c) i s hal yea and (d) second hal yea o li e; and
ju eniles, (e) i s hal yea and ( ) second hal yea o li e.
66
SMITHSONIAN CONTRIBUTIONS
TO
ZOOLOGY
(mo e ecap u es wi hin a sho e ime). The same pa e n
among subadul s (Table 5-12) and adul s (Table 5-13) o he
hal yea ollowing hei being ma ked sugges s beha io al
di e ences be ween he sexes ha in u n inc eases suscep ibil-
i y o ecap u e among males. The pa e n is simila among he
h ee age classes o he second hal yea a e ma k, excep ha
he ela ionship is e e sed o ju eniles in he all o 1977 and
he sp ing o 1978, and o subadul s in he all o 1978.
The ea e , he e is no pa e n emphasizing g ea e ca chabili y
o males.
Ju eniles o ei he sex ma ked in hei i s hal yea o li e
a e known o be ali e in he second hal yea in abou equal
numbe s (Tables 5-3 and 5-4; 30% o males e sus 32% o
emales). Ne e heless, by he hi d and ou h hal yea s, he
su i o ship a e is highe in emales (21% e sus 14%, X2 =
13.383,
P
<
0.01, o hi d hal yea ; 14% e sus 9%, X2 =
9.159, P
<
0.02), sugges ing ei he g ea e mo ali y in males,
di e en ial emig a ion, o bo h, along wi h possible lea ned ne
a oidance esul ing om highe ac ual ecap u e a e du ing he
i s yea a e ma k.
Chi-squa e analysis o ba s ma ked as ju eniles showed
some signi ican di e ences in su i o ship co ela ed wi h sex
and season o bi h. In gene al, emales ha e highe su i al
a es han males. Sp ing newbo n o bo h sexes su i e be e
in o he second hal yea o li e han do all newbo n wi h he
di e ence be ween season o bi h o emales signi ican a he
0.05 le el (35% e sus 28%, espec i ely; X2 = 6.103). In
subsequen hal yea s, he su i al a e is simila ega dless o
bi h season. Sp ing-bo n young may ha e he ad an ages o
long ges a ion (se en mon hs e sus ou ), bi h in he d y
season, weaning a he beginning o he we season, and
a ainmen o subadul age (and nea adul size) by he ime o
ig sca ci y a he heigh o he ains. Fall-bo n young, he
p oduc o a sho (4-mon h) ges a ion pe iod, a e bo n in he
we season, weaned a he heigh o he ains when ipe igs a e
beginning o be sca ce, and become subadul s in he d y season.
P eweaning e en s such as leng h o ges a ion and coinci-
dence o bi h wi h we o d y seasons, may ha e li le e ec on
su i al o baby ba s. Pos weaning condi ions, howe e , when
young ba s a e on hei own o he i s ime, may be c i ical.
Fall-bo n ba s a e weaned in he season o hea ies ains when
he e is a g ea e p obabili y o ge ing we while o aging. The
likelihood o ge ing we is inc eased by he sca ci y o igs
because he e is mo e eason o ly in he ain i quali y ood is
ha de o ind. We ing and consequen chilling could be a al
o weanling A. jamaicensis because o hei es ic i e,
ela i ely in lexible ene gy budge (see Sec ion 2, Physiology).
Su i o ship o he second hal yea ( i s hal yea a e
ma k) is no signi ican ly di e en o males and emales
ma ked as ju eniles. Howe e , emales appea o su i e a a
highe a e by he hi d and ou h yea (P
<
0.01 and P < 0.02,
espec i ely; based on da a in Tables 5-3 and 5-4). Su i al in
he hi d hal yea (second hal yea a e ma k) a o ed
all-bo n emales o e all-bo n males (P
<
0.05). Reasons o
hese di e en ial su i al a es a e unclea . The highe ini ial
ecap u e a e cha ac e is ic o males and hei highe subse-
quen a enua ion a e in he popula ion a e bo h e lec ed in
hese a es. Ne e heless, we can no be su e how much o he
appa en a enua ion a e ac ually e lec s g ea e di e en ial
emig a ion by males (highe p obabili y o con inued esidency
by emales) ins ead o mo ali y.
SUBADULTS.—Specula ion
on su i al and ecap u e a es o
ba s ma ked as subadul s mus include he assump ion ha
hese ba s a e al eady in he second hal yea when i s
cap u ed. Ba s ma ked as subadul s in he all we e bo n he
p e ious sp ing, and hose ma ked in he sp ing we e bo n he
p e ious all. The e o e, he i s hal yea a e being ma ked is
he hi d hal yea a e bi h. Any compa isons o hese eco ds
wi h hose o ju eniles a e on his basis.
The e is some una oidable "slop" in aging subadul s.
Subadul s caugh la e in he all include all indi iduals bo n in
he p e ious sp ing and some bo n ea ly in he all. In he i s
weeks o sp ing he pool o subadul s includes all young o he
p e ious yea , and should be a i s la ges . Midway h ough
sp ing he pool should ha e diminished o i s lowes le el
because mos ha e become adul s ( ep oduc i e) ega dless o
age.
La e in he sp ing, ju eniles o ha sp ing begin o c oss
he h eshold o subadul hood. Howe e , numbe s o subadul s
ma ked in he all g ea ly exceed he numbe ma ked in he
sp ing (Tables 5-5, 5-6, and 5-9). This is simply because in a
yea - ound cap u e p og am, mos o he subadul s a e cap u ed
and ma ked in he all, lea ing ewe unma ked ba s o be
cap u ed in he sp ing. Ba s a e aged as ju eniles only on he
basis o open epiphyses o he me aca pals and phalanges o he
wings. T ansi ion om ju enile o subadul s a us (ossi ica ion
o he epiphyses in he wing) is ela i ely apid (see Sec ion 3,
Rep oduc ion in a Cap i e Colony).
Assuming ha subadul s al eady a e in hei second hal yea
when i s cap u ed, su i al a e is sligh ly highe han in
ju eniles (Figu e 5-6). Ne e heless, he p opo ional loss om
he popula ion o he coho ma ked as ju eniles h ough he
second hal yea a e ma king mus ha e al eady been
abso bed by he coho ma ked as subadul s when hese ba s
comple e he i s hal yea a e ha ing been ma ked. Reasons
o ha loss a e he same as hose ou lined o losses among
ba s ma ked as ju eniles, excep ha necklace loss may no be
as impo an a ac o .
Obse ed o e all su i o ship h ough he i s hal yea
a e ma k (34%) o subadul s was no signi ican ly di e en
om ha o ju eniles (32%, X2 = 3.225), whe eas ecap u e
a e o he second hal yea a e ma k was signi ican ly lowe
(17%
e sus 23%,^ = 22.300, P
<
0.001) o ju eniles (Tables
5-3 and 5-5). The highes su i al a e o ba s ma ked as
subadul s was 43% in he all o 1978 (males, 45%; emales,
39%;
Table 5-9). As we sugges ed o ou su i o ship da a on
ju eniles, he su i al a e o subadul s is undoub edly highe
han he
43%
we eco ded du ing he se en hal -yea pe iods in
which we accumula ed su i o ship da a. I we es ic ou da a
NUMBER
511
67
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74
SMITHSONIAN CONTRIBUTIONS
TO
ZOOLOGY
a ack. We ha e a ac ed he Spec acled Owl (Pulsa ix
pe spicilla a) o ne s by imi a ing ba squeaks, and ha e caugh
hese owls in ne s se o ba s. Handley wa ched a Mo led Owl
(Ciccaba i ga a) ambushing Myo is nig icans and Molossus
coibensis a hei oos a dawn when he ba s we e milling
abou be o e en e ing. The owl s a ioned i sel in a ee o o
he side and highe han he oos and made equen so ies
down h ough he g oup o milling ba s, usually success ully.
We caugh Mo led Owls ai ly equen ly in ne s and
occasionally ha e caugh he Ve micula ed Sc eech-Owl (O us
gua emalae). A leas se en species o owls a e known o occu
on BCI, and some o hem a e a he abundan .
The Ba Falcon (Falco u igula is), mainly a c epuscula
eede and known o be an e ec i e p eda o o ba s (Ridgely,
1976), is a e o absen on BCI bu nes s nea by. The iming o
he depa u e om and e u n o day oos s by A. jamaicensis
makes i suscep ible o p eda ion by he Ba Falcon. Diu nal
hawks o en ake ba s, especially when hey ha e been
dis u bed in hei oos du ing he day. Se e al A ican hawks
egula ly in e cep ba s along hei ligh pa hs (Black e al.,
1979;
Fen on, Cumming, e al., 1977).
Habi s cha ac e is ic o A. jamaicensis such as luna phobia
and picking ui om a ee and ca ying i a hund ed o mo e
me e s away o be ea en a a eeding oos , a e p esumably
de ensi e s a egies (Howe, 1979). The equen use o he
onds o he spiny- unked black palm (As oca yum s andley-
anu i) and he a ching onds o he palm Oenoca pus
panamanus as dining oos s by A. jamaicensis may be a
de ensi e beha io o p o ec i sel om p eda o s (see Sec ion
8, Roos ing Beha io ). Response o dis ess calls o ba s ha
Augus (1979) called "mobbing" may also be a de ensi e
mechanism. A any a e, he e a e enough seemingly de ensi e
beha io al ai s in A. jamaicensis o sugges ha i has
conside able exposu e o p eda o s.
ACCIDENTS.—Many
ne ed A. jamaicensis ha e damaged
wings. Ne e heless, hey appea o su i e acciden s ha ea
wing memb anes o b eak inge s. These pa s heal ema kably
apidly and ligh is no no ably impai ed, as we ha e seen in
ou NZP colony. Common sou ces o inju y a e igh ing among
indi iduals and encoun e s wi h sha p plan spines o ho ns.
Bi e wounds a e mos equen in males, pa icula y young
males ying o ge access o emales (obse ed in he NZP
colony). Judging by ne ing esul s, ba s may a oid a eas in he
o es whe e spines and ho ns a e pa icula ly p e alen ;
al hough many, i no mos , o he ea s in ligh memb anes
ha we ha e obse ed mus o igina e om his sou ce. B oken
bones may esul om s uggles o ge ee when wings ge
angled. A b oken o ea m o hume us, ende ing he ba
ligh less, is in a iably a al.
DISEASE.—A ibeus
jamaicensis is known o ha e been
in ec ed wi h abies and ypanosomiasis on he Paci ic side o
he Canal de Panama a ea, bu disease in ba s is unknown on
BCI.
Summa y
On BCI A. jamaicensis made up abou 66% o he o al
nigh ly ca ch o ba s yea - ound. Numbe s ended o be highes
when young we e i s olan and lowes when he popula ion
was composed mos ly o subadul s and adul s.
The en i e sample o A. jamaicensis ma ked be ween
Oc obe 1976 and June 1980, consis ed o 8907 ba s cap u ed
a o al o 15,728 imes. Fi y-se en pe cen (5061 indi iduals)
we e caugh only once, lea ing 3846 cap u ed wo o mo e
imes. Subsequen cap u es pe ba diminished un il we had a
single indi idual cap u ed 11 imes. The p obabili y o
ecap u ing a ba was oughly he same ega dless o he
numbe o p e ious cap u es.
O e all su i o ship in o he second hal yea by ju enile A.
jamaicensis was 31%. Su i o ship om he second o six h
hal yea a e ma king among ba s ma ked as ju eniles was
lowe han ha o adul s and subadul s. Dispe sal du ing he
i s yea is he mos likely explana ion o his high a e o
disappea ance. Mo e equen ecap u e o males in he hal
yea ollowing ma king sugges s beha io al di e ences be-
ween he sexes ha inc ease suscep ibili y o ea lie and mo e
equen ecap u e among males.
Ju eniles o ei he sex ma ked in he i s hal yea a e
known o be ali e in he second hal yea in abou equal
numbe s (30% o males e sus 32% o emales). Bu by he
hi d and ou h hal yea s, he su i o ship a e is highe in
emales, sugges ing ha males a e mo e likely o emig a e,
ha e g ea e mo ali y, and possibly lea n o a oid ne s because
o hei highe ac ual ecap u e a e du ing he i s yea a e
ma k. O e all, emales ha e highe su i al ( esidency) a es
han males.
F equency o cap u e seems o ha e less o do wi h age han
wi h sex. Appa en ly mos ba s a e easie o cap u e he i s
ime, ega dless o age: 57% caugh once, 25% caugh only
wice, 18% caugh mo e han wice, and he ea e he cap u e
cu e le els ou
In ou sample o almos 17,000 eco ds o A. jamaicensis
(Oc obe 1976 o Oc obe 1980, adul and subadul emales
ou numbe ed adul and subadul males 55 : 45, while ju enile
emales a e ou numbe ed by ju enile males 48: 52. The
ecap u e his o ies o ba s ma ked as ju eniles also suppo he
con en ion ha emales ha e highe su i al (o esidency)
po en ial han males.
Among known-age ba s, he mean age a las cap u e was
2.02 yea s in he emales (n = 110) and 1.65 y s in he males (n
= 115). The a e age o he en i e sample (n = 225) was 1.83
yea s. The oldes male was ou and he oldes emale was 6.5
yea s old. Subsequen o 1980 Handley caugh wo A.
jamaicensis ha we e nine yea s old.
Wi h po en ially g ea longe i y, and wi h mos emale A.
jamaicensis p oducing wo young each yea , he species mus
be unde se e e popula ion con ols. Dea h a e mus be high,
bu i s causes a e specula i e. Mo ali y ac o s include

NUMBER 51175
s a a ion, p eda ion, acciden s, and disease, bu we ha e li le
di ec e idence and we ha e seen ew dead ba s.
Young o he "sp ing" bi h g oup a e weaned in a season o
abundan ood (Ap il-June) a he beginning o he ainy
season. Young o he "summe " g oup, on he o he hand, a e
weaned in a season o ood sca ci y (Augus -Oc obe ) in he
middle o he ainy season when hey could easily s a e be o e
becoming e icien o age s.
Young A. jamaicensis ace a numbe o isks. Wi h a high
me abolic a e and a low p o ein, high ca bohyd a e die one
would expec s a a ion o be ab up , pe haps in a single nigh
i ood sou ces a e no adequa e. Young A. jamaicensis and all
ages o he smalle -size species o A ibeus and Vampy essa
become unable o ly, lose con ol o body empe a u e, chill,
and die in as li le as an hou unde s ess and ood dep i a ion.
Rain mus inc ease he isk o s a a ion in young weaned in he
ainy season.
A ibeus jamaicensis elies on a ood sou ce ha is s ongly
pulsed in ime and space. I mus sea ch ou esou ces o he
u u e while ha es ing a cu en sou ce. I s beha io and die
adds isk om p eda o s ha may be a ac ed o he ood
sou ce.
Snakes and owls a e impo an p eda o s o ba s. Falcons,
opossums, coa imundis, and he la ge ca ni o ous ba Vampy-
um spec um also a e po en ial p eda o s. The e a e enough
seemingly de ensi e beha io al ai s in A. jamaicensis o
sugges ha i has conside able exposu e o p eda o s.
A ibeus jamaicensis is a s ong, obus ba ha usually
su i es acciden s ha ea he wing memb anes o b eak
inge s. These pa s heal ema kably apidly and ligh is no
no ably impai ed. Many ne ed A. jamaicensis ha e damaged
wings. Common sou ces o inju y a e igh ing among
indi iduals and encoun e s wi h sha p plan spines o ho ns.
6. Popula ion Es ima es
Egbe G. Leigh, J ., and Cha les O. Handley,
J .
Be ween
20
Oc obe 1976
and 19 May
1980,
Ba
P ojec
pa icipan s ma ked 8907 A ibeus jamaicensis, which
had
been cap u ed
a
o al
o
15,728 imes
by
20 Oc obe 1980,
he
las nigh
o
ield wo k co e ed by his sec ion. Can his eco d
ell us how long indi idual A. jamaicensis li e, and how many
he e a e on Ba o Colo ado Island (BCI)? Fo pu poses
o
he
ensuing analyses,
we
will assume ha
we a e
sampling
he
en i e popula ion
o A.
jamaicensis
on
BCI,
and
ha
i is a
closed popula ion.
A Simple Es ima e
AVERAGE
LIFETIME.—Suppose
A. jamaicensis has an expo-
nen ial "li e able,"
in
which he p obabili y
o a
ba li ing pas
age
y is emy,
whe e
e
( he base
o
Napie ian loga i hms)
is
2.71828, and m
is
he annual dea h a e pe capi a. Then,
i
he
cap u e e o , a e aged o e a pe iod du ing which
a
qua e o
he ba s a e eplaced, does no a y excessi ely om one such
pe iod o ano he , we may es ima e he a e age li e ime 1/m
o
hese ba s
by he
a e age ime elapsed be ween i s and las
cap u es
o
ba s cap u ed mo e han once. We accomplish his
as ollows:
1.
Le cd
be
he p obabili y ha
a
gi en ba , ali e be ween
ime
and
ime
+ d , is
cap u ed du ing his ime in e al.
He e,
c
is he cap u e a e and d deno es an "in ini esimal" ime
in e al. Then he p obabili y
o a
ba ha li es
L
yea s being
caugh n imes
in
i s li e
is
[(cL)n/n!]exp(-cL)
2.
I
he p obabili y ha
a
ba 's o al li e ime lies be ween
L
and L
+
dL yea s is (mdL) exp -(mL), hen he p obabili y ha
a ba will
be
caugh n imes in
i s
li e
is
P( l)
=
IT (mdL) exP (-mL)[(cL)n
/
n!] exp (-cL)
=
J
(cL)n
/
n!](mdL) exp
R n + c)L]
= (m/c)[c/(c
+
m)]n+1
Egbe
G.
Leigh,
J .,
Smi hsonian
T opical
Resea ch
Ins i u e,
Uni
0948,
APO AA
34002-0948,
o
Apa ado
Box
2072,
Balboa,
Republic
o
Panama.
Cha les
O.
Handley,
J .,
Na ional
Museum
o
Na u al
His o y,
Smi hsonian
Ins i u ion,
Washing on,
D.C.
20560.
3.
The a e age age a which
a
ba is i s cap u ed
is
J (c d ) exp
-
(m
+ c)
(cd ) exp - (m
+ c)
=
1
/
(m
+ c)
4.
This ime in e al
l/(m + c) is
also
he
a e age ime
in e al be ween cap u es,
as
long
as
cap u ing
a
ba does no
al e i s beha io . Thus, he a e age ime be ween i s and las
cap u es
is
[P(2)
+
2P(3)
+
3P(4)
+
4P(5)
(m
+
c)[P(2)
+ P(3)
=
1/m,
whe e
1/m is
he a e age li e ime.
The a e age ime elapsed be ween i s
and
las cap u e
o
ba s cap u ed mo e han once
is an
exac es ima e
o
a e age
li e ime,
i
he li e able
is
exponen ial. Al hough he ba s
we
caugh mo e han once li e longe han ou eco ds sugges , he
ba s we cap u ed mo e han once ended o be he ba s ha li e
longe han a e age in he i s place, and he e o s cancel.
We ha e calcula ed he a e age ime be ween i s and las
cap u es
by Ba
P ojec pa icipan s
o
ba s i s ma ked
by
Mo ison (1978b)
and
Bonacco so (1979) be ween 1972
and
1974.
Judging om he a e age in e al be ween i s and las
cap u e
by
Ba P ojec pe sonnel
o
he 33 emales and eigh
males o iginally ma ked by ei he Mo ison o Bonacco so and
cap u ed mo e han once du ing
he BCI Ba
P ojec (Table
6-1),
he
a e age li e ime
o
such ba s
was 1.5
yea s.
I we
conside only emales,
he
es ima e
o
a e age li e ime
is 1.6
yea s.
These ba s we e wo o mo e yea s old when i s caugh
by Ba P ojec pe sonnel, bu
i
he li e able is exponen ial, he
a e age expec a ion o u he li e does no depend on he ini ial
age o he ba .
The a e age ime be ween i s and las cap u e
o
301 ba s
caugh i e
o
mo e imes du ing
he
Ba P ojec
is 1.6
yea s.
This is
a
su p isingly low igu e o ba s caugh so many imes,
because in heo y he mo e imes
a
ba is caugh , he g ea e he
ime be ween i s and las cap u e.
EXPONENTIAL LIFE
TABLE.—Ou
basis o assuming an
exponen ial li e able
is as
ollows. The numbe s n(x)
o
he
77
78SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 6-1.—In e al be ween i s and las cap u e by he Ba P ojec o A ibeus jamaicensis i s ma ked on
BCI by Mo ison and Bonacco so.
Ca ego y
Bo h sexes
Females only
0.25*
6
5
0.50
9
6
1.00
2
2
Numbe o yea s be ween i s and las cap u e
1.50
9
8
2.00
4
2
2.50
4
3
3.00
6
6
5.00
1
1
To al
41
33
* Minimum assumed elapsed ime o ba s caugh o he i s ime and ecap u ed o he las ime in he same hal yea .
TABLE 6-2.—Obse ed and expec ed equency o cap u e o A ibeus jamaicensis ma ked on BCI by Mo ison
and Bonacco so, 1972-1974, and by he Ba P ojec , 1976-1980.
MORRISON-BONACCORSO
BATS
Obse ed (N = 146)
Expec ed (69)(0.5274)'1
BAT
PROJECT
BATS
Obse ed (N= 15728)
Expec ed (89O7)(0.4337)"1
69
69
8907
8907
>2
39
36
3846
3863
>3
22
19
1683
1675
>4
11
10
734
727
Numbe o imes caugh
>5
3
5
317
315
>6
1
3
146
137
>7
1
1.5
65
59
>8
0.8
21
26
>9
0.4
6
11
>10
0.2
2
5
>11
0.1
1
2
ba s ma ked by Mo ison and Bonacco so and caugh a leas x
imes du ing he ba p ojec o m a geome ic se ies whe e n(x)
= 69(77/146)*"1 (see Table 6-2). Mo eo e , one o he mos
s iking ea u es o he Ba P ojec 's cap u e eco ds is ha he
numbe n(x) o ba s cap u ed a leas x imes o ms a nea ly
pe ec geome ic se ies: /i(x + 1) = /i(l)A whe e n(l) = 8907
is he o al numbe o ba s caugh a leas once, and A is he
o al numbe o cap u es o ba s caugh a leas wice, di ided
by he o al numbe o cap u es, o 6821/15728 = 0.4337 (see
Table 6-2).
I he p obabili y ha a ba ali e a ime is caugh be ween
imes and + d is cd (whe e cap u e a e c is cons an and d
is he leng h o a "sho " ime in e al), hen i all ba s li e he
same li e ime (L), he p obabili y, ollowing he law o Poisson,
ha a ba may be caugh n imes in i s li e is
[(cL)n/n!]exp(-cL)
On he o he hand, i he ba s ha e an exponen ial li e able
wi h an a e age li e ime o 1/m, he chance P(x) ha a ba will
be cap u ed x imes in i s li e is
P(x) = [m/(c + m)] [c/(c + m)]
a geome ic se ies in x. This ag ees wi h ou obse a ion i
A = c/(c + m).
I hese ba s ha e an exponen ial li e able, hei age should
no a ec hei p ospec s o u he li e. Is an a e age li e ime
o 1.6 yea s easonable o emale A. jamaicensisl No ice ha
i he a e age li e ime 1/m o emales is 1.6 yea s, hen m =
1/1.6 = 0.625. Thus, su i al a e pe yea is exp (-m) =
53.5%,
and su i al a e pe hal yea is exp (-m/2) = 73.2%.
Hal o he emales ha li e long enough o ep oduce can
bea a young a hal yea a e hey a e bo n ( hey a e p oduc s
o he second ep oduc i e episode); all can bea a young e e y
hal yea he ea e . I all ha can bea a young do so, and i hal
he young a e emale, he expec ed numbe o young emales a
newbo n emale ba will bea in he li e is
(l/4)(0.732) + (1/2X0.535) [1 + 0.732 + (0.732)2 + ...]
= 0.183 + (0.2675)/[I-0.732] x 1.18.
I , as ou eco ds show, one o e e y en ma u e emale ba s
in each b eeding season ails o ep oduce (see Sec ion 3,
Rep oduc ion in a Cap i e Colony, and Sec ion 4, Rep oduc-
ion on BCI), he popula ion will be e y nea ly in balance.
The e o e, he a e age expec a ion o u he li e o hese ba s
p obably ma ches he a e age li e ime o all emale A.
jamaicensis a he closely and sugges s ha hei li e able is
indeed exponen ial.
NUMBER
OF
BATS.—Fo
he sake o analysis, we shall make
wo mo e assump ions, e en hough hey a e no comple ely
ue (see Sec ion 5, Su i al and Rela i e Abundance).
(1) Cap u ing a ba does no a ec he p obabili y o
cap u ing i again la e .
(2) Ba s a e sampled om a pool in which all indi iduals
a e equally liable o cap u e.
NUMBER
511
79
Then
we
may es ima e
he
o al numbe
o
ba s a ailable
o
cap u e du ing he Ba P ojec om he numbe
o
ba s ac ually
ma ked, di ided by he chance
a
ba in he pool will
be
ma ked,
whe e
he
la e
is
assumed equal
o he
p opo ion
o
ma ked
ba s ha
a e
ecap u ed
a
leas once.
In
o he wo ds,
he
o al
numbe a ailable du ing
he
p ojec equals
he
numbe
o
ba s
ma ked, di ided
by he
chance
o
ma king
a ba
(equals
he
numbe
o
ba s ma ked di ided
by he
numbe
o
ba s
ecap u ed).
The numbe
o
ba s a ailable
o
cap u e
a any one
ime
is
he o al numbe a ailable o e
he
du a ion
o he
p ojec ,
mul iplied
by he
a e age
ba
li e ime
1/m
(including bo h
sexes),
and
di ided by he o al du a ion
o
he sampling pe iod
(4 yea s). Ano he
way o
saying his
is: he
o al numbe
o
ba s a ailable
a a
gi en ime equals
he
o al a ailable du ing
he p ojec , imes
he
mean li e ime
o a ba ,
di ided
by he
du a ion
o
he p ojec .
I A ibeus jamaicensis
has an
exponen ial li e able,
and i
all ba s "a ailable
o
cap u e"
a e
equally likely
o be
caugh ,
hen
he
o al numbe
o
a ailable ba s du ing
he
ou yea s
o
he sampling pe iod
is
(8907)/(0.4337)
=
20535.
The
numbe
a ailable
a
any one ime
is
he o al, imes
he
li e ime
o
hese
ba s (a e aged
o
bo h sexes), di ided
by he
ou yea s'
du a ion
o
he p ojec ,
o
[(20535)(1.5)]/4
=
7701.
An equi alen way
o
calcula e popula ion size
o
hese ba s
is
o
ind
he
cap u e a e
c by
se ing
A =
0.4337 equal
o
c/(c
+
m), and o assume ha he mo ali y a e m o bo h sexes
concu en ly
is
0.667.
We
ind
c =
0.5106, which implies ha ,
on
he
a e age, (0.5106)/12
o
4.166%
o he
ba s
a e
caugh
each mon h.
As
(15,728)/48,
o 328,
ba s we e caugh
pe
mon h,
on
he a e age, he o al popula ion o
A.
jamaicensis on
BCI
is
328/(0.04166)
=
7873.
VALIDITY
OF
ASSUMPTIONS.—How
alid a e he assump-
ions behind hese calcula ions?
I
all a ailable ba s a e equally
liable
o
cap u e
and i all
ba s ha e equal p ospec s
o
u he
li e ime, ega dless o cu en age, hen he chances o ecap u e
o
all
ba s ma ked wi hin
a
gi en hal yea will
be he
same,
ega dless
o
age
o
sex.
This
is no
ue. Adul s ma ked
in he
all
o
1976
and 1977
we e
a
leas
as
likely
o be
ecap u ed
as he
ju eniles
o
subadul s ma ked
a
ha ime, al hough adul s ma ked la e
on
we e ecap u ed much less o en han we e ju eniles
o
subadul s (Table
6-3; and
Sec ion
5,
Su i al
and
Rela i e
Abundance, Table 5-9). Few
o
he adul s newly ma ked
in he
all
o 1979 and
sp ing
o 1980
we e caugh again, while
ecap u e a es we e mo e nea ly no mal
o
ju eniles
and
subadul s. Many
o
hese newly ma ked adul s we e caugh
om peninsulas
on he
mainland su ounding
BCI,
whe e
we
ne ed
a
less o en han
on he
island.
In
gene al,
he
a io
o
he p opo ion
o
newly ma ked ju eniles
and
subadul s,
o he
TABLE
6-3.—Pe cen ages,
by
sex
and age
class,
o
A ibeus
jamaicensis
ma ked
in
successi e
hal
yea s
on BCI
and
subsequen ly ecap u ed; p opo ion
o
young
o
adul ecap u es;
and
p opo ion
o
adul s ma ked among
o al
cap u es
o
adul s (ma ks
and
ecap u es) caugh
in
each hal yea .
Age
class
Pe cen
ecap u ed
Ju eniles
Subadul s
Adul s
Ra io
o
pe cen age
ecap u ed
Ju eniles
and Subadul s
/
Adul s
P opo ion
o
new ma ks among cap u ed ba s
Numbe
adul s ma ked
/
To al cap u es
o
adul s
Pe cen
ecap u ed
Ju eniles
Subadul s
Adul s
Ra io
o
pe cen age
ecap u ed
Ju eniles
and Subadul s
/
Adul s
P opo ion
o
new ma ks among cap u ed ba s
Numbe
adul s ma ked
/
To al cap u es
o
adul s
Fall
1976
0.3784
0.4185
0.5225
0.7820
1.0000
0.2955
0.3526
0.3841
0.8677
1.0000
Fall
1977
0.3814
0.4413
0.5055
0.8259
0.8044
0.4215
0.4600
0.5050
0.8754
0.7910
Sp ing
1978
0.7273
0.5404
0.3586
1.6372
0.5380
0.6154
0.5778
0.5364
1.0966
0.4314
Fall
1978
FEMALES
0.3855
0.4879
0.2524
1.8427
0.3787
MALES
0.3093
0.3459
0.3271
1.0151
0.4632
Sp ing
1979
0.5551
0.5315
0.3668
1.4918
0.5303
0.5368
0.4935
0.2934
1.7762
0.5000
Fall
1979
0.4743
0.3113
0.2194
1.8373
0.2476
0.4220
0.3163
0.2893
1.3404
0.3710
Sp ing
1980
0.3723
0.2105
0.1237
2.9450
0.3811
0.3109
0.2154
0.2055
1.4383
0.2219

80SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 6-4.—To al cap u es o A ibeus jamaicensis on BCI in successi e hal yea s by sex and age class, by
p opo ion o young o adul emales, by p opo ion o new ma ks o o al cap u es (ma ks and ecap u es) among
adul emales.
Age class
Ju eniles
Subadul s
Adul s
Ju eniles
Subadul s
Adul s
All ba s
Nonadul ba s
Ju eniles and Subadul s / Adul emales
Adul emale ma ks / Adul emale
cap u es
Fall
1976
76
251
213
95
201
173
1009
623
2.925
0.836
Fall
1977
247
495
587
285
421
534
2569
1448
2.467
0.624
To al all nonadul s /
Sp ing
1978
56
241
579
48
232
522
1678
577
0.997
0.496
Fall
1978
FEMALES
85
354
296
MALES
101
200
256
TOTALS
1292
740
Sp ing
1979
307
197
771
308
232
514
2329
1044
PROPORTIONS
2.500
0.348
To al all adul emales = ?306 ~
1.354
0.477
2.950
Fall
1979
397
535
666
506
409
658
3171
1847
2.773
0.233
Sp ing
1980
401
49
533
367
132
318
1800
949
1.781
0.364
Fall
1980
612
543
544
625
364
430
3118
2144
3.941
0.248
p opo ion o newly ma ked adul s ecap u ed subsequen ly,
was highe he g ea e he p opo ion o adul s al eady ma ked
(Table 6-3).
Ou assump ion ha all a ailable ba s a e equally liable o
ecap u e is clea ly w ong. I seems, a he , ha as mo e ba s
we e ne ed, we eached a poin whe e mos o he adul ba s on
BCI had been ma ked, so mos ba s a ailable o ma king on
he island we e ju eniles and subadul s. The ea e , many o he
ba s ma ked as adul s esided in locali ies whe e p ospec s o
ecap u e we e no g ea o hey lea ned o a oid he ne s.
No only a e ba s in ce ain places mo e liable o ecap u e,
younge ba s a e mo e liable o ecap u e han olde ones. The
a io o he o al numbe o ju eniles and subadul s o bo h
sexes o he numbe o adul emales handled (coun ing each
ins ance o cap u e) in he all o 1976 was 2.925; combining all
he all ca ches o he sampling pe iod oge he , i was 2.950
(Table 6-4). Because adul emales can ha e no mo e han wo
young a yea , adul emales mus be nea ly wice as ha d o
ca ch as ju eniles o ei he sex.
A Re ined Es ima e
Popula ion size in hese ba s can be be e es ima ed a e wo
in e media e s eps.
(1) Calcula ing mo ali y a es o bo h sexes.
(2) Calcula ing he o al numbe s o ma ked ba s o each
sex ali e in successi e hal yea s (Dowdeswell e al.,
1940;
Fishe and Fo d, 1947).
The i s s ep enables he second, because he o al numbe o
indi iduals M( ) ali e in hal -yea o he Ba P ojec ha we e
ma ked ea lie is he numbe m'{ ) ma ked in he i s hal yea
( all 1976), imes he p opo ion p( - 1) su i ing o hal -yea
, plus he numbe m'(2) ma ked in he second hal yea (sp ing
1977) imes he p opo ion p( -2) o hose su i ing o
hal -yea , and so on. In summa y:
M( ) = m'(l)p( - 1) + m'(2)p( -2) + . . . + m'( - l)p(l).
MORTALITY
ESTIMATES.—I
we assume ha he numbe s o
A.
jamaicensis on BCI do no change subs an ially om yea o
yea , he simples way o calcula e he mo ali y o adul s is o
conside he ecap u es o
a
sample o ma ked ba s as long a e
hey we e i s ma ked as possible. The ba s ma ked by
Mo ison (1978b) and Bonacco so (1979) p o ide a sui able
sample. Be ween 1972 and 1974 hey ma ked 1212 A.
jamaicensis, o which abou hal we e emale. O hese emales,
47 we e caugh by he Ba P ojec a e
1
July 1977. Because 93
o he 178 adul emale A. jamaicensis ma ked by he Ba
P ojec in he all o 1976 we e ecap u ed a e 1 July 1977, i
seems easonable o assume ha 93/178 o he emales ma ked
by Mo ison and Bonacco so, and s ill li ing in he all o 1976,
we e ecap u ed a e
1
July 1977. I
so,
hen 47(178/93), o 90,
o Mo ison and Bonacco so's emale ba s we e ali e in he all
o 1976, implying a su i al a e o (90/606)1/35, o 58%, a
yea , and an expec a ion o u he li e o 1.835 yea s.
Simila ly, 16 o hei ma ked male ba s we e caugh a e
1
July
1977.
As 58 o
151
adul males ma ked by he Ba P ojec in he
NUMBER 51181
all o 1976 we e caugh a e
1
July 1977, oughly 16(151/58),
o 42, o he ba s ma ked by Mo ison and Bonacco so we e
ali e du ing he all o 1976, implying a su i al a e o
46.7%
a yea . I he popula ion we e declining, hese es ima es would
be oo high, and ice e sa.
A second way o calcula e su i al a e is o conside he
da es o i s cap u e o he ba s ecap u ed in a gi en hal yea .
Le n( , x) be he numbe o ba s cap u ed in hal -yea ha
we e i s ma ked in hal -yea x, and le m'(x) be he o al
numbe o ba s ma ked in hal -yea x (Tables 6-5 and 6-6).
Then m'(x)p( -x) is he o al numbe o ba s ma ked in
hal -yea x su i ing o hal -yea . I a ac ion c( ) o he ba s
ali e in hal -yea a e cap u ed hen, n( , x) = c( )m'(x)p( - x).
I he ba s ha e an exponen ial li e able, hen
p( -x) = exp[-m( -x)/2],
whe e exp (-m) is he a e age su i al a e pe yea . Mo eo e ,
n(U = c( ) exp [-m( -
sugges ing ha we calcula e m as he coe icien o eg ession
o log [n( , x)/m'(x)] on x.
This me hod o es ima ing m su e s om he disad an age
ha ba s ma ked la e in he p ojec a e mo e likely o be om
in equen ly sampled si es. Thus, ba s i s ma ked in hal -yea
7 a e less likely o be ecap u ed in hal -yea 8 han al eady
ma ked ba s caugh in hal -yea 7. On he o he hand, his
es ima e o m does no depend on he s abili y o he ba
popula ion as a whole.
None heless, es ima es (Table 6-7) o su i al a es o
emale ba s based on ecap u es o hal -yea 7 and hal -yea 9,
o which he co ela ion be ween he hal -yea x o ma king
and he loga i hm o he p opo ion n( , x)/m'(x) o ba s ma ked
hen ha we e ecap u ed in hal -yea is ela i ely close, ag ee
wi h each o he , and wi h he es ima e based on ba s ma ked by
Mo ison and Bonacco so and ecap u ed by he Ba P ojec .
The a e age o he o me wo es ima es is 57.4%, compa ed
wi h 57.99% om he Mo ison and Bonacco so ba s.
On he o he hand, es ima es o su i al a es o male ba s
based on ecap u es o hal -yea s 7 and 9 a e age 37.2% a
yea , ma kedly lowe han he 46.7% es ima ed om ecap u es
o male ba s ma ked by Mo ison and Bonacco so. Do olde
males su i e be e ? The 46.7% igu e is based on a a he
small sample o ecap u es. Ano he piece o e idence is he
ecap u e in he all o 1981 o a male ma ked by Mo ison and
Bonacco so. I 37.2% o he males su i e each yea , a male
has one chance in 4000 o su i ing he 8.5 yea s o he all o
1981,
so he chances a e
1
in 7 ha one o hei 606 ba s would
s ill be li ing, and pe haps hal ha , had i been ali e, we would
ha e caugh i hen. Tha cap u e eco d sugges s, bu does no
p o e, ha a leas some olde males su i e a he be e .
Finally, i he popula ion is s able, we may calcula e adul
mo ali y a es om he p opo ions n(x, )/[m'(x) + n(x)] o
ba s ma ked in a gi en hal -yea among he o al numbe o
ba s—bo h ma ks, m x), plus ecap u es, n(x)—caugh in la e
hal yea s. I m'(x) + n(x) = c(x)N(x), whe e c(x) is he
p opo ion o he N(x) ba s ali e in hal -yea
x
ha we e caugh
du ing ha hal yea , while
n(x, ) = c(x)m'( ) exp -m(x -1)/2,
hen n(x, )/[m'(x) + n(x)] = [m'( )/N(x)] exp -m(x -1)/2. I N
is cons an , he eg ession on x o he loga i hm o his
p opo ion is m/2. I he ba popula ion is g owing by a ac o
exp ( /2) pe hal yea , hen N(x) = N( ) exp (x -1)/2, and ou
eg ession gi es (m + )/2.
Mo ali y es ima es o ba s, bo h male and emale, ma ked
in he all o 1976 and he sp ing o 1978, ag ee wi h each o he ,
and a e only sligh ly lowe han hose calcula ed om ma king
da es o ba s ecap u ed in he all o 1979 and he all o 1980.
Mo ali y es ima es o ba s ma ked in he all o 1977 a e much
lowe , bu like hose o ba s ma ked in he sp ing o 1978, he
all 1977 es ima es a e based on s ong co ela ions, illus a ing
he unce ain ies in ou calcula ions.
I he chances o cap u ing a ba depend on i s age class, hen
his es ima e equi es ha he age composi ion o ba s cap u ed
in successi e hal yea s be he same. I he chances o cap u ing
a ba depend on whe he i is al eady ma ked, hen he
p opo ion o ecap u es among he ba s handled should also be
cons an . Finally, i chances o ecap u e a y om place o
place, ne ing e o should be dis ibu ed o e he island in he
same way du ing successi e hal yea s.
A e ou igu es ue mo ali ies? Al hough ba s do occasion-
ally mo e be ween BCI and he su ounding mainland, his
exchange does no seem o be g ea (see Sec ion 7, Mo e-
men s). We do no hink we a e mis aking emig a ion o
mo ali y. Howe e , some ba s do lose hei necklaces. Adding
oge he he in e als be ween ime o i s necklacing and o
las ecap u e o each ba conce ned, ou p ojec has moni o ed
ba s ca ying Mo ison and Bonacco so wing bands o o e
i y ba yea s. Du ing his in e al, i e ba s los hei
necklaces, sugges ing ha he su i al a e o necklaces is 90%
a yea . I so, he a e age su i al a e o
A.
jamaicensis is
11
%
highe han ou eg essions sugges , pe haps be ween 62% and
65%
a yea o adul emales.
Gi en he su i al a e o ju eniles ela i e o adul s, and he
b eeding a e o adul emales, he su i al a e o adul emales
mus be nea
60%
a yea i
he
popula ion is o be in balance. To
show his, we make he ollowing assump ions and obse a-
ions.
Hal o he emales ( hose ha a e bo n in he second
ep oduc i e episode) can bea a young a hal yea a e bi h;
all can bea a young when hey a e a yea old, and all can b eed
once each hal yea he ea e ; hal hei o sp ing a e emale.
Twen y-eigh o he 74 ju enile emales caugh in he all o
1976 we e ecap u ed in successi e hal yea s, while 87 o he
178 adul emales ma ked hen we e ecap u ed la e (see
Sec ion 5, Su i al and Rela i e Abundance, Table 5-9). Thus,
a ma ked ju enile emale had 0.7242 imes he chance o being
ecap u ed as an adul as did an adul emale ma ked hen. I in
gene al, he chance o a ju enile emale su i ing h ough i s
82SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 6-5.—Numbe s o emale A libeus jamaicensis ma ked on BCI in successi e hal yea s by age class, by
numbe s n(x, y) o hese ecap u ed in la e hal -yea s y, and by numbe n(x) o ecap u es in hal -yea s x o hese
ba s ma ked in p e ious hal
yea s;
and based on hese alues, p opo ions n( , x)/m'(x) o hese ba s ecap u ed
in selec ed hal -yea s among hose ma ked in ea lie hal -yea s x, and p opo ions n(x, )/[m'(x) + n(x)] o ba s
ma ked in selec ed hal yea s among hose caugh in la e hal yea s.
Females
Ma ked
Ju eniles
Subadul s
Adul s
To al [m'(x)]
Recap u es om hal -yea y
1
3
4
5
6
7
8To al
Recap u ed/Ma ked
n(9,
x)/m'(x)
«(8,
x)/m'(x)
«(7,
x)/m'(x)
n(x, l)/[m'(x) + n(x)]
«(x. 3)/Im'(x) + n(x)]
n(x, 4)/Im'(x) + «(x)]
1
Fall
1976
74
227
178
479
16/479
23/479
33/479
3
Fall
1977
236
358
366
960
91
91
41/960
47/960
88/960
91/1051
4
Sp ing
1978
66
198
290
554
Recap u es n(x,
68
181
249
38/554
31/554
67/554
68/803
181/803
5
Fall
1978
83
289
103
475
Hal -yea x
6
Sp ing
1979
272
141
368
781
7
Fall
1979
331
257
155
743
8
Sp ing
1980
368
19
194
583
y) in hal -yea x o ba s i s caugh in hal -yea y
27
81
61
169
46/475
32/475
100/475
27/644
81/644
61/644
36
102
74
114
326
P opo ions
76/781
75/781
183/781
36/1107
102/1107
74/1107
33
88
67
100
183
471
115/743
107/743
33/1214
88/1214
62/1214
23
47
31
32
75
107
315
157/583
23/898
47/898
21/898
9
Fall
1980
500
286
135
921
16
41
38
46
76
115
157
489
16/1410
41/1410
38/1410
i s yea is 0.7242 imes he chance (p2) o an adul emale
su i ing a ull yea , and i a emale has a 1/4 chance o bea ing
a emale young when i is a yea old, and a 1/2 chance o doing
so each hal yea he ea e , hen he numbe (Ro) o emale
young a newbo n emale A. jamaicensis can expec o bea
du ing he li e is
Ro = 0.7242
= 0.1810p + 0.362/^/(1 -p).
I su i al a e p2 pe yea is
0.5625
(so p = 0.75), hen
Ro = 0.95; i
p2
= 0.6 (sop = 0.7746), hen Ro = 1.10; and
i p2
= 0.64 (sop = 0.8), hen Ro = 1.30.
In ac , no all emales b eed in each b eeding season. In six
ep oduc i e episodes in ou cap i e colony wild-caugh adul
emales we e p egnan in 66 o 72 chances (91.7%), mul ipa-
ous cap i e-bo n emales we e p egnan in 12 o 14 chances
(85.7%),
and p imipa ous cap i e-bom emales we e p egnan
in 14 o 20 chances (70%) (see Sec ion 3, Rep oduc ion in a
Cap i e Colony, Tables 3-4 and 3-5). On BCI in he mon h o
Ap il, when all emales should be ep oduc i e, we in a iably
caugh a ew non ep oduc i e adul emales and some
subadul s who ailed o ep oduce on ime (undoub edly
9-mon h-old young o he p e ious July-Augus bi h-g oup).
Fo example, in Ap il 1978,9 o 102 adul emales caugh we e
non ep oduc i e, and we caugh 22 subadul emales; in Ap il
1979,
7 o 47 newly ma ked adul emales we e non ep oduc-
i e.
We can make a minimum co ec ion o his by
mul iplying ou alues o Ro o di e en su i al a es by 0.89,
he pe cen age (133/149) o adul emales caugh in 1978 and
NUMBER 51183
TABLE 6-6.—Numbe s o male A ibeus jamaicensis ma ked on BCI in successi e hal yea s by age class, by
numbe s n(x, y) o hese ecap u ed in
la e
hal -yea s y, and by numbe n(x) o
ecap u es
in hal -yea s x o hese
ba s ma ked in p e ious hal
yea s;
and based on hese alues, p opo ions «( , x)/m'(x) o hese ba s ecap u ed
in selec ed hal -yea s among hose ma ked in ea lie hal -yea s x, and p opo ions «(x, )/[m'(x) + n(x)] o ba s
ma ked in selec ed hal yea s among hose caugh in la e hal yea s.
Males
Ma ked
Ju eniles
Subadul s
Adul s
To al [m'(x)]
Recap u es om hal -yea y
1
3
4
5
6
7
8To al
Recap u ed/Ma ked
«(9,
x)/m'(x)
«(8,
x)/m'(x)
n(7,
x)/m'(x)
«(x.
l)/[m'(x)+n(x)]
n(x,3)/[m'(x)+«(x)]
l(x,4)/[m'(x)+n(x)]
1
Fall
1976
88
173
151
412
3/412
9/412
9/412
3
Fall
1977
261
300
299
860
79
79
10/860
18/860
56/860
79/939
4
Sp ing
1978
52
135
220
407
5
Fall
1978
97
159
107
363
Hal -yea x
6
Sp ing
1979
272
154
242
668
7
Fall
1979
410
196
197
803
8
Sp ing
1980
341
65
73
479
Recap u es «(x, y) in hal -yea x o
ba s
i s caugh in hal -yea y
55
235
290
13/407
22/407
44/407
55/697
235/697
14
50
60
124
15/363
31/363
61/363
14/487
50/487
60/487
19
65
57
101
242
P opo ions
46/668
50/668
164/668
19/910
65/910
57/910
9
56
44
61
164
334
90/803
126/803
9/1137
56/1137
44/1137
9
18
22
31
50
126
256
114/479
9/735
18/735
22/735
9
Fall
1980
496
198
101
795
3
10
13
15
46
90
114
291
3/1086
10/1086
13/1086
1979 ha we e ac ually b eeding.
The e may be o he e o s in ou calcula ions. On he one
hand, sp ing-bo n ju eniles should su i e be e han all ones;
on he o he , ou igu es o ju enile mo ali y do no include
dea hs be o e he ju eniles a e old enough o ly in o ou ne s.
All in all, we belie e he su i al a e o adul emale A ibeus
jamaicensis is be ween 60% and 64% a yea .
ESTIMATE OF SURVIVING MARKED BATS IN SUCCESSIVE
HALF
YEARS.—Le
M^ ) ep esen he numbe o ma ked
adul emale ba s ali e a he beginning o hal -yea , and
M ( ) be he numbe o ma ked subadul emale ba s also
ali e hen. In addi ion, p ep esen s he su i al a e o adul
emales pe hal yea .
Eigh y- h ee o he 227 subadul ba s ma ked in he all o
1976 we e ecap u ed in succeeding hal
yea s,
while 87 o 178
adul s ma ked hen we e ecap u ed la e (Sec ion 5, Su i al
and Rela i e Abundance, Table 5-9), sugges ing ha a subadul
emale was 0.8010 imes as likely as an adul o su i e he
nex hal yea . I his is ue in any hal yea , hen a ju enile
emale is (0.7242)/(0.8010) =
0.9041
imes as likely as an adul
emale o su i e he nex hal yea . I we make he con en ion
ha a ac ion -ip o he adul s, a ac ion 0.8010 Jp o he
subadul s, and a ac ion
0.9041
<p
o he ju eniles ma ked in
a gi en hal yea su i e o he beginning o he nex hal yea ,
and ha ju eniles become subadul s, and subadul s become
adul s, a ha ime, hen we may assume he numbe Ms ( ) o
ma ked subadul emales a he beginning o a hal yea o be
0.9041
<p
m^ ( - 1), whe e mj ( - 1) is he numbe o ju enile
emales ma ked in hal -yea - 1.
The ma ked adul emales ali e a he beginning o hal -yea
90SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
4546 47 48 49 50
11ii1
0 12 3 4 5
kilome e s
FIGURE 7-1.—Dis ibu ion o locali y g oups (agg ega ions o locali ies) whe e ba s we e cap u ed on BCI and
adjacen mainland ( om Table 7-1). The locali y g oups a e as ollows: (1) Lu z; (2) Ba bou -Hood; (3)
Shannon-AMNH; (4) Shannon-Balboa; (5) Lake-Wheele ; (6) Ba bou
S eam;
(7) Mille
Ridge;
(8) Fue es; (9)
Con ad; (10) Pla eau; (11) D ay on End; (12) A mou End; (13) Ze ek 21; (14) S andQey Ridge; (15) S andley
End; (16) O chid Island; (17) G oss Poin ; (18) Chapman; (19) Ha a d; (20) Mona G i a; (21) Gigan e; (22)
F ijoles Rd.; (23) Bohio; (24) Buena Vis ; (25) Pena Blanca.

NUMBER 51191
46
45
I
• •• •! .... I
2 3
kilome e s
FIGURE 7-2.—Pe cen age o o he -cap u es in he o al eco ds ( o al indi iduals plus o he -cap u es o hem) in
cap u e sui es o A ibeus jamaicensis a locali y g oups on BCI and he adjacen mainland, 1976-1980. Da a a e
om Table 7-2. Isolines a bi a ily enclose locali y g oups wi h simila pe cen ages o o he -cap u es. Ma ginal
numbe s iden i y kilome e squa es (see Appendix, Me hods o Cap u ing and Ma king T opical Ba s).
92SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
I
a. oo
H
B-
•*! §
5n
- i .s
<-
•? oo
eg « T!
< - n
H S £
so
§
-C
1
V
O he
_
3
CO
<s
cs1
o
oo
VO
o
—
CO
js
o
O
oo
c—
n
CO
s
cap u es
1
1
.
NO
12
CO
o
O
s
o
-
2
CO
CO
O
oo
o
§
oo
230
°
s
42 106
I
5309
6
.
Lu z
"™
CO
<s
o
o
o
o
o
o
CS
o
-.
CS
—
—
2
cs
-
°
19 17
640 1
]
.
B
a bou
CO
-
o
o
o
o
o
-
s
,3.
—
CO
n
O
-
cs
24 19
!
730 1
<
.
Shannon
Vn
NO
o
o
o
o
o
o
o
-
o
cs
o
CO
—
o
NO
o
cs
o
i
165
i
•a
pa
6
|
T
NO
o
o
o
o
o
o
o
-
o
CO
CO
o
o
°
cs
CO
°
156
le
J
Lake-W
-
o
o
o
o
o
o
o
-
o
,^
o
o
°
cs
«
(S
°
cs
1
234
1
,
B
a bou
ON
00
-
-
oo
o
o
o
CO
-
00
n
VO
ON
o
-
169
CO
NO
NO
CS
29 18
j
2499
2
u
idg
,
Mille R
•—<
o
o
o
o
o
o
n
-
cs
—
cs
CO
NO
cs
_
NO
139
cs
-
NO
| 1189
n
CO
cs
o
CO
o
o
o
o
o
°
s
NO
^-
cs
°
CO
_
oo
-
-
o
I
578
.
Con ad
^.
CS
-
o
o
o
o
o
oo
—«
cs
n
oo
CO
CO
8!
CO
c—
CO
50 13
[
1377 1
Pla eau
CO
CN
o
o
o
o
o
o
o
°
o
Q
o
cs
o
s
o
o
-
138
-o
W
D ay on
—
o
o
in
o
o
-
o
o
_
cs
2
2
o
o
CO
CS
456
I
A mou -
cs
.
CO
o
o
o
o
o
o
o
CO
TI-
CS
°
cs
NO
o
o
o
146
Ze ek
21
o
OO
o
•>
o
o
o
o
o
-
cs
_
n
CO
-
°
—
NO
ON
n
-
CO
cs
cs
786d«e
_u
-
s
o
o
o
o
o
CO
~
CO
o
°
cs
^^
o
CO
o
cs
649
-a
§
Slandley
n
NO
NO
o
n
cs
o
o
o
o
n
n
-
_
s
cs
°
o
^H
oo
-
o
TT
;
249
_,
J
O chid I
16.
oo
O
O
o
o
o
o
o
o
a
o
o
-
°
o
CS
o
-
o
145
.|
G oss
P(
-
o
o
cs
o
o
o
NO
—
o
o
-
CO
>n
cs
-
o
-
cs
388
Chapmai
18.
o
o
-
o
o
o
o
TT
o
o
o
o
o
°
o
o
CO
o
o
o
cs
125
Ha a d
19.
o
o
o
o
o
cs
o
o
o
©
o
o
-
°
o
o
°
o
o
o
o
•1
Mona Gi20.
CO
,—«
o
o
o
o
o
o
o
o
o
o
o
o
o
o
o
°
o
o
o
o
140
Gigan e
21.
s
o
o
o
cs
o
o
o
o
o
o
o
o
o
o
o
o
°
o
o
o
°
o
156
•3
J
F iioles ]
22.
NO
CS
ON
NO
ON
CS
o
o
o
o
o
cs
-
n
—
o
n
o
CO
^.
o
o
©
o
746
Bohio23.
ON
CO
O
CO
n
o
o
o
-
o
o
o
o
o
o
o
«
o
o
o
°
o
143
is i
Buena
V
24.
CO
n
O
o
o
o
o
o
o
o
o
o
o
o
-
o
o
o
138
5
Pe ia
Bla
25.
oo
oo
CO
_
CS
s
CO
CO
.01
~
••*
o
>n
s
-
co
s
cs
-
906
n
s
oo
cs
24
216
17,322*
16:
Ib al
NUMBER 51193
TABLE
7-2.—Tabula ion o o al eco ds, o al o he -cap u es, and pe cen age o o he -cap u es (see ex o
explana ion) o he o al numbe o A ibeus jamaicensis in each locali y g oup on BCI and adjacen mainland,
1975-1980. See Figu e 7-1 o loca ions o locali y g oups.
Locali ies
1.
Lu z
2.
Ba bou -Hood
3.
Shannon-AMNH
4.
Shannon-Balboa
5.
Lake-Wheele
6. Ba bou S eam
7.
Mille Ridge
8. Fue es
9. Con ad
10.
Pla eau
11.
D ay on End
12.
A mou End
13.
Ze ek 21
14.
S andley Ridge
15.
S andley End
16.
O chid Island
17.
G oss Poin
18.
Chapman
19.
Ha a d
20.
Mona G i a Poin
21.
Gigan e
22.
F ijoles Road
23.
Bohio
24.
Buena Vis a
25.
Pena Blanca
(A)
To al
indi iduals
5309
640
730
165
156
234
2499
1189
578
1377
138
456
146
786
649
249
145
388
125
40
140
156
746
143
138
(B)
To al
o he -cap u es
1567
233
305
65
67
57
899
421
239
524
23
92
37
248
142
66
48
111
30
4
13
2
126
39
33
(B/A)
Pe cen
o he -cap u es
30
36
42
39
43
24
36
35
41
38
17
20
25
32
22
26
33
29
24
10
9
1
17
27
24
Mona G i a Poin , 10%; Gigan e, 9%; and F ijoles Road, 1%).
These alues show ha mo e o he popula ion was ma ked and
a la ge po ion o he a ea equen ed was sampled a he
cen al locali ies. Ba s mo ing in any di ec ion om he cen al
locali ies we e likely o encoun e o he cap u e s a ions. Ba s
using he mo e isola ed lakesho e locali ies p obably o aged
ex ensi ely on he mainland; hence, we sampled smalle
ac ions o hei popula ions and he a eas ha hey equen ed.
The e ec s o sampling small ac ions o popula ions and hei
o aging a eas a e bes seen on he mainland whe e s a ions
we e ew and (excep o Bohio) cap u e e o was low. We
o en ne ed a Bohio and i anked six h in numbe s o A.
jamaicensis caugh . Howe e , Bohio was nine een h in he
numbe o o he -cap u es (17%, Table 7-2) and 83% o i s A.
jamaicensis we e caugh only once.
The lowes o he -cap u e a es (Table 7-2) on BCI we e a
D ay on End (17%) and A mou End (20%). Among he mos
isola ed BCI s a ions, and loca ed in high, old o es con aining
ew ui ees, each o hese wo si es is o e a kilome e om
he nex nea es s a ion. Ba s a he sou he n pe ime e o BCI
may o age on he adjacen mainland whe e he o es is
younge and con ains many ig ees. Among he ew ecap u es
a Gigan e and Mona G i a Poin on he mainland we e A mou
End and D ay on End ba s.
A he ou se o he Ba P ojec we g idded BCI and planned
o andomly ne ha g id. This plan was based on he
pe cep ion ha good ( ep esen a i e) ca ches nigh a e nigh
would esul i we mo ed a andom h ough ou g id, using
well-si ed and well-se ne s a choice ne ing s a ions. Ou
p esump ion was nai e; on some nigh s we caugh almos
no hing, whe eas on o he nigh s we we e o e whelmed wi h
ba s.
I became ob ious ha he nigh ly dis ibu ion o ui ba s
was no andom, bu coincided wi h he p esence o ipe ui ,
which ends o be une enly dis ibu ed in ime and space. Ne s
se a ce ain ui ing ees caugh ba s, ne s placed elsewhe e
o en did no , and a locali y migh ha e an abundance o
ba s
on
one da e and ew on ano he . When he e was only one
p e e ed ee wi h ipe ui on BCI, clea ly ha was he place
o be o ba s. The e a e many ac o s ha in luence he
dis ibu ion and mo emen s o ui ba s on BCI: dis ibu ion
and abundance o p e e ed oods; ain, wind, moon phase, and
cloud co e ; he size, opog aphy, and o he limi a ions o
insula i y. These ac o s ended o o e whelm ou e o s o se
up meaning ul measu es o cap u e e o in ol ing ime and
ne ing condi ions.
94SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 7-3.—Pe cen ages o o he -cap u es o A ibeus jamaicensis o selec ed locali y g oups on BCI weigh ed
by he numbe o o al cap u es o A. jamaicensis a each o he locali y g oups whe e he o he -cap u es we e
eco ded. Adjus ed pe cen ages we e de i ed by di iding he numbe o o he -cap u es (da a om Table 7-1) by
he numbe o o al cap u es o ha locali y g oup and educing ha alue o a pe cen age o i s column o al. The
name o each locali y g oup analyzed is ollowed by he o al numbe o o he -cap u es in pa en heses.
Locali y
g oup
1.
Lu z
2.
Ba bou -Hood
3.
Shannon-AMNH
4.
Shannon-Balboa
5.
Lake-Wheele
6. B a bou S eam
7.
Mille Ridge
8. Fue es
9. Con ad
10.
Pla eau
11.
D ay on-End
12.
A mou -End
13.
Ze ek 21
14.
S andley Ridge
15.
S andley End
16.
O chid Island
17.
G oss Poin
18.
Chapman
19.
Ha a d
20.
Mona G i a PL
21.
Gigan e
22.
F ijoles Road
23.
Bohio
24.
Buena Vis a
25.
Pena Blanca
To al
To al
cap u es
5309
640
730
165
156
234
2499
1189
578
1377
138
456
146
786
649
249
145
388
125
40
140
156
746
143
138
17,322*
Shannon-
AMNH (305)
%
6.35
8.06
10.04
11.51
6.95
1.17
2.71
1.60
2.82
10.26
9.82
1.79
1.87
1.38
0.84
3.28
1.87
7.00
8.68
1.98
Chapman
(110)
%
3.51
18.86
8.28
3.34
2.36
0.44
2.30
2.85
1.59
3.95
1.12
0.71
0.82
2.19
19.79
26.32
1.48
O he -cap u es
Lu z
(1567)
%
7.09
9.71
8.60
6.75
7.52
6.77
5.39
3.50
4.87
6.09
2.55
1.03
4.02
2.46
1.17
2.36
4.45
4.99
4.69
1.47
0.84
0.79
1.64
1.27
Mille
Ridge (899)
%
6.05
2.74
3.35
3.55
10.02
1.25
6.60
11.50
14.04
7.02
0.70
3.00
4.01
6.34
2.70
3.13
7.41
0.51
2.34
2.35
0.68
0.70
S andley
End (248)
%
1.25
1.49
1.29
9.20
3.45
4.46
1.68
1.72
9.82
16.48
28.80
1.92
1.25
7.04
6.71
3.45
* 15,736 band numbe s.
Also,
we had o adjus (p o a e) cap u e a es because he
cap u e e o di e ed among locali ies. We ied " ecap u es
pe ne -nigh " and " ecap u es pe ne -hou ," bu hese ga e
anomalous esul s, p obably because ne ing e o commonly
was in e sely ela ed o cap u e success. In o he wo ds, he
poo e ou ca ch, he mo e ne s we main ained and he longe
we wo ked hem.
A e ealizing ha cap u e e o measu ed in uni s o ime
was un eliable, we disco e ed ha he numbe s o A.
jamaicensis caugh pe locali y p o ided us he simples and
mos use ul measu e o cap u e e o . A. jamaicensis egula ly
made up abou wo- hi ds o ou ca ch and was usually he
commones ba anywhe e we ne ed. Ou accumula ed 17,322
ma k and ecap u e eco ds o his species a ied be ween
locali ies om a maximum o 5309 ba s in he Lu z Wa e shed
o a minimum o 40 a Mona G i a Poin (Table 7-3). Using
cap u e da a om A. jamaicensis (Table 7-1), we de i ed
weigh ed pe cen ages o o he -cap u es by di iding he numbe
o o he -cap u es a a locali y g oup by he o al numbe o A.
jamaicensis eco ded a ha same locali y g oup. Each alue
was hen con e ed o a pe cen age o i s column o al o
p oduce he A. jamaicensis-'weigh ed pe cen ages plo ed in
Figu es 7-3 h ough 7-8.
The e icacy o hese me hods o illus a ing dispe sion o
o he -cap u e eco ds o
A.
jamaicensis a ound a sample cen e
can be seen by compa ing esul ing pe cen ages wi h he
dis ance be ween each cap u e locali y and he cen e (Table
7-4). When g aphed (Figu e 7-9), he da a poin s app oxima e
a line declining wi h inc easing dis ance o ze o indica ing ha
he equency o o he -cap u es was p opo ional o dis ance
om he sample cen e .
Ano he way we examined he co espondence o equency
o o he -cap u es wi h dis ance om he sample cen e was by
plo ing he adjus ed alues on a map con aining concen ic
ci cles wi h adii o 1, 2, and 3 km om he sample cen e
(Figu e 7-4). The Shannon-AMNH da a i a he well
NUMBER 51195
45 46 47 48 49 50 51 52 53 54 55
i.••11..,,
12 3 4 5
kilome e s
FIGURE 7-3.—O he -cap u es o indi idual A ibeus jamaicensis eco ded a leas once in he Shannon-AMNH
locali y g oup on BCI, 1976-1980 (exp essed as a pe cen age o o al o he -cap u es o Shannon-AMNH ba s and
weigh ed by
A.
jamaicensis cap u e means; da a a e om Table
7-3;
Shannon-AMNH is ou lined wi h a hexagon).
Isolines a bi a ily enclose locali y g oups wi h simila pe cen ages o o he -cap u es. Inne mos line encloses
co e a ea. Ma ginal numbe s iden i y kilome e squa es (see Appendix, Me hods o Cap u ing and Ma king
T opical Ba s).

96SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
47
46
4545 46 47 48 49 50 51 52 53 54
• * • •
i
12 3 4 5
kilome e s
FIGURE
7-4.—O he -cap u es o indi idual A libeus jamaicensis eco ded a leas once in he Shannon-AMNH
locali y g oup on BCI, 1976-1980 (exp essed as a pe cen age o o al o he -cap u es o Shannon-AMNH ba s and
weigh ed by
A.
jamaicensis cap u e means). Ci cles a one- and wo-kilome e in e als encompass 29% and 85%,
espec i ely, o all o he -cap u es. Shannon-AMNH is ou lined wi h a hexagon.
NUMBER 51191
47
46
45
1 .,,
2 3
kilome e s
FIGURE 7-5.—O he -cap u es o indi idual A ibeus jamaicensis eco ded a leas once in he Chapman locali y
g oup on BCI, 1976-1980 (exp essed as a pe cen age o o al o he -cap u es o Chapman ba s, and weigh ed by
A.
jamaicensis cap u e means). Chapman is ou lined wi h a hexagon. Isolines a bi a ily enclose locali y g oups
wi h simila pe cen ages o o he -cap u es. Inne mos line encloses co e a ea. Ma ginal numbe s iden i y
kilome e squa es (see Appendix, Me hods o Cap u ing and Ma king T opical Ba s).
98SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
55
54
»••••'
2 3
kilome e s
FIGURE 7-6.—O he -cap u es o indi idual A ibeus jamalcensis eco ded a leas once in he Lu z locali y g oup
on BCI, 1976-1980 (exp essed as a pe cen age o o al o he -cap u es o Lu z ba s, and weigh ed by A.
jamaicensis cap u e means). Lu z is ou lined wi h a hexagon. Isolines a bi a ily enclose locali y g oups wi h
simila pe cen ages o o he -cap u es. Inne mos line encloses co e a ea. Ma ginal numbe s iden i y kilome e
squa es (see Appendix, Me hods o Cap u ing and Ma king T opical Ba s).
99
45
1....
2 3
kilome e s
FIGURE 7-7.—O he -cap u es o indi idual A ibeus jamalcensis eco ded a leas once in he Mille Ridge
locali y g oup on BCI, 1976-1980 (exp essed as a pe cen age o o al o he -cap u es o Mille Ridge ba s, and
weigh ed by A jamaicensis cap u e means). Mille Ridge is ou lined wi h a hexagon. Isolines a bi a ily enclose
locali y g oups wi h simila pe cen ages o o he -cap u es. Inne mos line encloses co e a ea. Ma ginal numbe s
iden i y kilome e squa es (see Appendix, Me hods o Cap u ing and Ma king T opical Ba s).
106SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
49 50 51
4546 47
»••"'
0 1 2 3 4 5
kilome e s
FIGURE 7-13.—O e lap o co e a eas in Wes -cen al and pe iphe al po ions o BCI. A = A mou End; BO =
Bohio; B V = Buena Vis a; CH = Chapman; C = Con ad; D = D ay on End; F = Fue es; MR = Mille Ridge; PB
= Pena Blanca.

NUMBER 511107
TABLE
7-5.—Co e a eas on BCI and icini y in which he cen al locali y had
a lowe pe cen age o o he -cap u es (see ex o explana ion) o A ibeus
jamaicensis (cap u ed a leas once he e) han some o he locali ies in he co e.
Each en y includes he cen al locali y (CAPITALIZED), pe cen age o
o he -cap u es* he e, ank among all locali ies in e ms o numbe o
indi idual A. jamaicensis caugh , and pe cen ages o o he -cap u es* a o he
locali ies wi hin he co e. * = A. jamaicensis- /eighled.
Locali y
LUTZ
Ba bou -Hood
Shannon-AMNH
Lake-Wheele
Ba bou S eam
Shannon-Balboa
MILLER
RIDGE
Fue es
Lake-Wheele
Con ad
G oss Poin
S andley Ridge
Pla eau
PLATEAU
Shannon-Balboa
Shannon-AMNH
Con ad
Lake-Wheele
Mille Ridge
A mou -End
FUERTES
Mille Ridge
Con ad
BOHIO
Pena Blanca
Buena Vis a
SHANNON-AMNH
Shannon-Balboa
Pla eau
D ay on-End
BARBOUR-HOOD
Chapman
Ha a d
Shannon-AMNH
Ba bou S eam
CONRAD
Pla eau
Mille Ridge
Fue es
S andley Ridge
ARMOUR-END
Mona G i a Poin
Shannon-Balboa
CHAPMAN
Ha a d
Ba bou -Hood
ORCHID
ISLAND
G oss Poin
Buena Vis a
Shannon -Balboa
Ze ek21
SHANNON-BALBOA
O chid Island
GROSS
POINT
O chid Island
To al
cap u es
5309
2499
1377
1189
746
730
640
578
456
388
249
165
145
Pe cen
o he -cap u es
7
10
9
8
7
7
7
11
10
14
7
6
7
6
14
12
11
8
7
6
13
14
11
16
27
18
10
11
10
10
11
24
14
11
10
10
15
13
12
11
13
15
12
20
26
19
9
44
15
10
9
14
19
70
Rank
1
2
3
4
6
7
9
10
11
12
13
15
19
TABLE 7-6.—Co e a eas on BCI and nea by mainland in which he cen al
locali y had a highe pe cen age o o he -cap u es (see ex o explan ion) o
A ibeus jamaicensis (cap u ed a leas once he e) han o he locali ies in he
co e.
Each en y includes he cen al locali y
(CAPITALIZED),
pe cen age o
o he -cap u es* he e, ank among all locali ies in e ms o numbe o
indi idual A. jamaicensis caugh , and pe cen ages o o he -cap u es* a o he
locali ies wi hin he co e. * = A. jamaicensis-weigh ed.
Locali y
STANDLEY
RIDGE
S andley-End
Con ad
STANDLEY-END
S andley Ridge
Ze ek 21
BARBOUR
STREAM
Lake-Wheele
LAKE-WHEELER
Ba bou S eam
Mille Ridge
G oss Poin
ZETEK
21
BUENA
VISTA
DRAYTON-END
PENA
BLANCA
HARVARD
Chapman
To al
cap u es
786
649
234
156
146
143
138
138
125
Pe cen
o he -cap u es
23
14
10
29
16
10
36
18
26
12
10
10
42
82
31
84
49
24
Rank
5
8
14
16
18
20
22
23
24
No o he -cap u es eco ded.
locali y wi hin each co e (Tables 7-5 and 7-6). Pe cen ages o
o he -cap u es a he cen al locali ies anged om 6% o 84%.
The lowe pe cen ages a Lu z, Mille Ridge, and Pla eau mus
e lec he numbe s o cap u es, which we e highes a hese
h ee locali ies. I o he -cap u es a a cen al locali y a e
p edominan ly local esiden s, hei ep esen a ion ou o all
ba s caugh a ha locali y will be in e sely ela ed o he o al
numbe caugh . The g ea e he numbe o indi idual ba s
in ol ed, he g ea e he likelihood ha o he -cap u es will be
widely dispe sed, hus also educing he p opo ion o
o he -cap u es a a cen al locali y.
The high pe cen ages o o he -cap u es a Buena Vis a and
Pe ia Blanca simply demons a e ideli y o hose si es, which
a e day oos s in channel ma ke s. Whene e we sampled hese
oos s, mos o he ba s caugh we e ones aken he e be o e.
The low pe cen ages o hese ba s ne ed away om hei day
oos s sugges ha hey o aged whe e we did no ne , p obably
on he mainland. A compa ison o he Bohio co e a ea (Figu e
7-13) and he S andley End a ea o equen use (Figu e 7-8)
shows ha while hese a e places whe e Buena Vis a and Pe ia
Blanca ba s o aged, hey a e p obably no hei main eeding
a eas.
The e a e ex ensi e a eas o he mainland nea hese
oos s whe e we ha e ne e ne ed (Figu e 7-13).
The e is a dicho omy o co e a eas in o hose whose cen al
locali ies ha e bo h lowe pe cen ages and ewe o he -cap u es
han o he locali ies in he co e (Table 7-5) and hose whose
cen al locali ies ha e highe pe cen ages and mo e o he -
cap u es han do neighbo ing locali ies (Table 7-6). This
di ision may be co ela ed wi h he numbe o cap u es.
108SMITHSONIAN CONTRIBUTIONS
TO
ZOOLOGY
60
55
50
45
40
I-
Z 35
LJLJ
O
30
cc
m
25
20
15
10
5
Numbe
o
indi iduals
in
each cap u e ca ego y
1
5061
57
2
2163
24
3
949
11
4
417
5
5
171
2
6
81
1
7
44
0.5
8
15
0.2
Numbe
o
eco ds
in
each
1
5061
32
2
4326
28
3
2847
18
4
1668
11
5
855
5
6
486
3
7
308
2
8
120
1
9
4
0.04
10
1
0.01
cap u e
9
36
0.2
10
10
0.01
11
1
0.01
To al
8907
ca ego y
11
11
0.01
15728
5061
.
8907
5061
15728
I
0.57
I 0.32
Indi iduals
3 4 5 6 7 8
CAPTURE CATEGORY
10 11
FIGURE 7-14.—F equency
o
cap u e
o
A ibeus jamaicensis
on
BCI. Numbe
o
indi iduals caugh
1, 2, 3, 4,
and mo e imes, and numbe
o
eco ds
in
sui es
o
indi idual ba s
in
each cap u e ca ego y.
S andley Ridge and S andley End (Table 7-6) a e excep ions
in
ha compa a i ely high numbe s o
A.
jamaicensis we e caugh
a each ( hey anked i h
and
eigh h, espec i ely),
ye
hei
numbe s
o
o he -cap u es
a e
also ela i ely high
(see
discussion
o
Fideli y, below,
in
his sec ion).
The dis ibu ion
o ig
ees p obably exe s
an
in luence
on
he ela i e numbe s
o
o he -cap u es
a
any locali y. The i s
se en locali ies
in
Table
7-5
(Lu z h ough Ba bou -Hood)
ha e
an
abundance
o ig
ees. Combined, Lu z
and
Mille
Ridge ha e o e
200 ig
ees, mos ly Ficus insipida,
he
a o ed ui
o
A. jamaicensis. Chapman, S andley Ridge, and
S andley
End
ha e
a
ai numbe
o ig
ees,
bu no
many
F.
insipida. The o he locali ies in Tables 7-5 and
7-6
ha e ew
ig
ees o he han s anglc igs (mainly
F.
ob usi olia) whose
ui s
a c no
p e e ed
by
A. jamaicensis.
I
he
numbe
o A.
jamaicensis ha ems
is
limi ed
by he
a ailabili y
o
sui able ee holes,
and i
ees wi h sui able
holes a e e enly dis ibu ed o e BCI hen he e mus be many
ha em oos s ha
a e no
nea pa ches
o ig
ees.
A
hese
places
we
should ca ch
he
ba s
o
esiden ha ems
and ew
o he s, hus accoun ing
o
high pe cen ages
o
o he -cap u es
cha ac e izing hose si es.
The
ba s
o day
oos s
no
loca ed
nea
ig
ees mus ou inely a el o o he pa s
o
he island
o
eed. Howe e , ba s wi h day oos s nea
ig
ees also commu e
o o he locali ies
o
o age whene e he ees nea hei oos s
lack ipe ui . Ne e heless, hese ba s need
o
a el less han
ba s esiding
in
ee holes
in
ig-poo places. Tu ne (1975)
showed ha Desmodus o undus ou inely changed oos si es
o emain close o
a
spacially shi ing esou ce. We assume ha
ei he oos ing si es
a e
limi ed
on BCI o
ha
A.
jamaicensis
ha e high ideli y o indi idual day oos s because we ound no
e idence
o
ou ine shi s e en among bachelo males.
Bachelo males oos
in
oliage
and no in
ee holes.
Al hough hey migh
be
expec ed
o
concen a e
in
ig- ich
a eas
and o
shi concen a ions
o
ollow ui a ailabili y,
hey appa en ly
do
no , judging
by he
unusually high a io
o
males consis en ly ound
a
S andley
End.
NUMBER
511
50
#
40-
Q
a 30
H
109
o
CO
<
CD
20-
10
0-
19
12
6%
20-
29
29
15%
30-
39
36
19%
40-
49
4
2%
50-
59
80
42%
60-
69
9
5%
70-
79
3
2%
80-
99
0
100
17
9%
I
I I 1 I I I I
0-19 20-29 30-39 40-49 50-59 60-69 70-79 80-99
CAPTURE CATEGORIES
(%)
100
FIGURE 7-15.—Fideli y o A ibeus jamaicensis o he Chapman locali y g oup on BCI. F equency
o
cap u e o
indi idual ba s wi h eco ds
o
mul iple cap u es ha we e caugh
a
leas once
a
Chapman a e g aphed
as
pe cen ages
o
eco ds
in
a ious ac ional cap u e ca ego ies
('/6
o
less
o
eco ds = 0%-19%, Vs-V*
= 20%-29%, V3 = 3O%-39%, 2/s = 40%-49%,
lh
= 50%-59%, Vs-Vi
=
60%-69%, 3A = 70%-79%.
4/s-
5/6 = 80%-99%, all = 100%). Row
A in
he able lis s ac ional cap u e ca ego ies (as pe cen ages), ow
B
lis s
numbe o indi idual ba s whose eco ds a Chapman all in o each ca ego y (n = 190), and in ow C he numbe s
a e con e ed o pe cen ages. See also Table 7-7.
Fideli y
We examined ideli y
o a
pa icula locali y g oup
by
analyzing sui es o mul iple cap u e eco ds o indi idual ba s.
Fo each locali y g oup
we
compiled eco ds
o
all
A.
jamaicensis ha ing mul iple cap u es ha we e aken
a
leas
once in ha pa icula locali y g oup be ween Oc obe 1976 and
Oc obe 1980. Fo each ba , we abula ed o al cap u es,
he
numbe
o
cap u es
in
ha pa icula locali y g oup,
and
pe cen age
o
i s o al eco ds in ha locali y g oup. F om ha
abula ion we g ouped hose ha ing mul iple cap u es acco d-
ing o he pe cen age
o
hei occu ence
a
ha locali y g oup
among o al eco ds (e.g., Chapman, Figu e 7-4).
The dis ibu ion
o
eco ds among ac ional ca ego ies
is
in luenced by he numbe o eco ds pe indi idual sui e. In all,
3846
A.
jamaicensis we e ecap u ed (Figu e 7-14). Mos
(57%) we e ecap u ed only once (i.e., had wo cap u es), 24%
we e ecap u ed wice ( h ee cap u es), 11% ecap u ed h ee
imes ( ou cap u es), and 8% ecap u ed ou
o
mo e imes.
Thus,
peaks would be expec ed (Figu e 7-15) a 50% and 100%
o wo- eco d sui es; 33%, 66%, and 100% o h ee eco ds;
and 25%, 50%, 75%, and 100% o ou eco ds, and so o h.
Because mos ecap u ed ba s we e caugh again only once,
50%
and 100% peaks pe locali y should occu mos o en.
These da a indica e ideli y
o
a
pa icula locali y g oup.
High ideli y means
a
high equency
o
eco ds
in
he
60%-100% in e al, which ep esen s 3/s, 2/3,3A, 4/s, o mo e
cap u es
in
he
same a ea.
The
hiqhe
he
equency
o
same-si e cap u es,
he
highe
he
ideli y.
Low
ideli y
is
e ealed
by
a
highe equency
o
eco ds
in he
0%-40%
in e al (2/s,
l ,
!/4, Vs, and !/6
o
cap u es
o
less)
o
ha
locali y g oup.
A
simpli ied summa y (Table 7-7)
o
locali y-
g oup ideli y, like ha
in
Figu e 7-15, shows h ee basic
pa e ns among ou da a.
Pa e n
I
(e.g., A mou End; Figu e 7-16, Table 7-7) co e s
mos
o
he locali y g oups. The e is
a
s ong simila i y among
110SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
© — O O
n l^ oo d
-^ co —
oo >n so so -
~ s oo >/•> oo
« N /I -
-^~* SO —* CO ^^ CO ^^— — co - co
I
g
NUMBER
511
60-
50
111
o
cc
0.
<
o
CO
<
00
40-
30-
20-
10-
0
J
"O = Chapman
••
• Con ad
A
-
Fue es
•A
'
O chid Island
0-19 20-29 30-39 40-49 50-59 60-69 70-79 80-99
CAPTURE CATEGORIES
(%)
FIGURE 7-16.—Fideli y
o
A ibeus jamaicensis
o
Pa e n
I
locali y g oups
on
and nea BCI: Chapman, Con ad,
Fue es, and O chid Island. See Figu e 7-15
o
u he explana ion.
100
he cap u e-ca ego y abula ions
o
locali y g oups wi h his
pa e n
in
spi e
o
hei di e se loca ions
and he
dispa i y
in
numbe s
in
each sui e
o
mul iple cap u e eco ds ( om 51
o
1321).
These locali y g oups sha e
a
low
deg ee
o
ideli y
(<19%
in
he
60%-100% column)
and
a
high equency
o
single cap u es (82%-99%
in he
0%-40%
and
0%-50%
columns).
Pa e n
II
(Lu z, Bohio,
and
S andley
End;
Figu e
7-17,
Table 7-7) comp ises h ee seemingly un ela ed locali y g oups
whose eco ds
o
ideli y
a e
ema kably simila .
All
h ee
show highe ideli y, compa ed wi h
he
Pa e n
I
locali y
g oups (30%-32%
in he
60%-l00% column),
and
ewe
single eco ds (68%-70%
in he
0%-40%
and
0%-50%
columns). Thei eco ds
may be
simila
o
di e en easons.
We sampled li le
o
he p obable o aging ange
o
he Bohio
ba s,
so
hey had li le isk
o
cap u e excep
a
Bohio, he eby
appea ing
o
ha e high ideli y
o
ha loca ion.
In
con as ,
he
ba s
o
S andley
End may
ac ually
be
seden a y because hey
we e ecap u ed mos ly
a
S andley End
o
nea by (see Figu e
7-8 and discussion
o
Co e A eas ea lie
in
his sec ion). O ,
i
we sampled only pa
o
hei o aging a eas,
he
si ua ion
a
S andley End may
be
simila o ha
a
Bohio.
We a e con iden ha we ha e sampled he ull ex en
o
he
o aging anges
o
Lu z ba s. High ideli y
o
Lu z may e lec
a la ge local esiden popula ion,
o
may esul om ca ching
he same indi iduals epea edly because
o
he a ac ion
o
he
ig g o e
in
Lu z Ra ine. These
ig
ees
may
adequa ely
suppo local popula ions
o A.
jamaicensis
and
pe iodically
a ac ba s om a a when ipe igs a e sca ce elsewhe e. I his
is ue hen some
o
he ba s showing high ideli y a e ac ually
oppo unis ic ansien o age s a he han Lu z esiden s.
Pa e n
III
(Buena Vis a and Pe ia Blanca; Figu e 7-18, Table
7-7) shows
he
highes ideli y.
The
ob ious explana ion
o
his pa e n
is
ha
we
sampled only
a
ac ion
o
he o aging
anges
o
hese ba s,
and
because
we
cap u ed hem mos ly
a
hei day oos s. We ne ed ba s only a nigh a all o he locali y
g oups and, e iden ly, we
did
no
ne
whe e he ba s
o
Buena
Vis a and Pe ia Blanca mos o en o aged.
The deg ee
o
ideli y wi hin pa e ns appea s o be ela ed o
ne ing e o . On his basis we so ed Pa e n
I
locali y g oups

112SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
50-|
40-
Q
c 30
-
O
CD
20-
10-
0J
O
=
B oh
i o
•
=
S andley
End
•
=
Lu z
I
|
••••••
0-19 20-29 30-39 40-49 50-59 60-69 70-79 80-99
CAPTURE CATEGORIES
(%)
FIGURE 7-17.—Fideli y o A ibeus jamaicensis o Pa e n II locali y g oups on and nea BCI: Bohio, Lu z, and
S andley End. See Figu e 7-15 o u he explana ion.
100
in o
wo
subpa e ns (Table 7-8).
One
wi h
19 o
mo e
ne ing-nigh s pe locali y (subpa e n la); he o he wi h ewe
(subpa e n Ib). Wi h ew excep ions (low ideli y
a
Con ad
and Lake-Wheele ; unusually high ideli y
a
D ay on End),
ideli y and ne ing e o wi hin pa e ns
a e
di ec ly
and
posi i ely co ela ed, sugges ing ha he mo e o en
a
locali y
g oup is ne ed he mo e likely he ba s using ha locali y g oup
will be ecap u ed.
Ne e heless, al hough ne ing e o di e ed g ea ly
be-
ween subpa e ns
la
and Ib, ideli y was simila . The e o e,
a ia ion
in
ideli y be ween pa e ns is independen o ne ing
e o . Subpa e n la locali ies we e ne ed om 20 o 52 nigh s
and ideli y anged om 7%
o
18% (Table 7-8). Subuni
Ib
locali ies we e ne ed only
6
o 17 nigh s apiece, ye he ideli y
ange was simila (8%-16%).
Fideli y was su p isingly simila (30%-32%) among pa e n
II locali ies e en hough ne ing e o anged om 14 nigh s a
Bohio
o
197
a
Lu z (Table 7-7). Howe e , when compa ing
ideli y on he basis o simila ne ing e o be ween pa e ns
I
and
II
he
esul s
a e
ma kedly di e en .
Fo
example,
14
ne ing nigh s demons a ed 6% and 14% ideli y in Pa e n
I,
and 30% ideli y in Pa e n II. Locali y g oups wi h 24 and 26
ne ing nigh s
in
Pa e n
I
showed
9% and
13% ideli y;
whe eas ideli y
was
mo e han double (31%) a e simila
e o a S andley End (25 ne ing nigh s) in Pa e n II.
I all ba s ha ing mul iple cap u es ha we e ecap u ed only
once
in
a
pa icula locali y g oup
a e
excluded,
we
should
elimina e mos ha we e p obably based elsewhe e. Following
ha logic, we conside ed A. jamaicensis cap u ed wo o mo e
imes in
a
locali y g oup mo e likely o be local esiden s. The
p opo ion o ba s wi h wo o mo e cap u es in a locali y g oup
o
he
o al mul iple-cap u e coho om ha locali y g oup
a ied om 0.9% o 71.9%; al hough hey s ill so ed in o he
same pa e ns I, II, and III (Table 7-7). Few indi iduals caugh
wo o mo e imes con i ms low ideli y a locali y g oups such
as Con ad (11%) and Chapman (18%). The wo-plus cap u e
ac ion a Con ad may con ain ew local
ba s,
judging om he
lowe p opo ion (54%)
o
hose ba s
in
he
60%-100%
column. Howe e ,
he
high pe cen age (78%)
in
he
60%-
100%
ac ional cap u e ca ego y a Chapman indica es ha he
wo-plus ac ion, al hough small, may ep esen mos ly local
ba s.
The high p opo ion (78%)
o
ba s
a
Pe la Blanca ha
we e caugh he e a leas wice, and he high pe cen age (92%)
o hose in he 60%-100% b acke , sugges ha his exe cise is
a alid means o es ima ing ideli y
o a
locali y because high
ideli y is o be expec ed a
a
ha em day oos .
NUMBER 511
50
w
40-
Q
LU
cc
30
Q_
O 20
H
113
CO
CD10-
0J
Buena is a Ligh
Pena Blanca Ligh
I
1 1 1 " 1 1 •—i 1
0-19 20-29 30-39 40-49 50-59 60-69 70-79 80-99
CAPTURE CATEGORIES
(%)
FIGURE 7-18.—Fideli y o A ibeus jamaicensis o Pa e n III locali y g oups nea BCI: Buena Vis a and Pena
Blanca. See Figu e 7-15 o u he explana ion.
100
We ound ha 102 (37%) o he 275 A. jamaicensis caugh a
S andley End had been caugh he e a leas wice (Table 7-7).
O hese, 83% all in o he 60%-100% ecap u e ca ego y o
he si e. This is good e idence ha a high p opo ion o he
S andley End ba s we e local esiden s,
a
conclusion u he
suppo ed when we examine hei eco ds in g ea e de ail.
Only wo (7%) o he wo-plus emales we e caugh as many as
wo imes elsewhe e (a Bohio and S andley Ridge). Also, only
six (8%) o he males caugh wo o mo e imes a S andley End
we e caugh as many as wo imes a o he si es, all nea by
(Fue es, 1; Mille Ridge, 8; and S andley Ridge, 2).
The sex a io o he S andley End A. jamaicensis (102) is
skewed owa d males (73 : 29; Table 7-9). A Lu z, ano he
Pa e n II locali y wi h simila ideli y s a is ics, he a io
is
53 :47. Du ing h ee o he ne ing episodes a S andley End
when wo-plus ba s we e caugh , males ou numbe ed emales
30 o 1. O males caugh a leas wice,
31%
we e also caugh
h ee o mo e imes while only 14% o he emales we e caugh
ha o en. In con as , only 21 (29%) males we e caugh o e a
ela i ely long pe iod ( wo o ou yea s), whe eas 14 (48%)
emales we e ecap u ed
a
S andley End du ing he same
pe iod. Close o h ee- ou hs (71%) o he males we e ound
he e o a yea o less (Table 7-9). Age dis ibu ion among ba s
cap u ed wo o mo e imes was simila in males and emales
(Table 7-10); howe e , a la ge p opo ion o he young males
han young emales s ayed in he a ea o be caugh la e as
adul s.
We belie e ha he males a S andley End we e p edomi-
nan ly bachelo s, and ha mos o all o he esiden emales
we e membe s o ha ems. O he 20 emales caugh as adul s
du ing seasons o ep oduc ion, 18 we e p egnan , lac a ing, o
pos lac a ing
a
one o mo e
o
he cap u es. The g ea es
numbe o emales caugh as adul s a S andley End in any hal
yea was 11. P esuming ha we caugh mos o he esiden
emales, hey p obably ep esen om one o h ee ha ems. In
he ou ha ems o
A.
jamaicensis ha Mo ison (1979) s udied
on BCI, he ound om ou o 11 adul emales (X= 6.5).
We s ill do no know why ideli y was so high in he h ee
Pa e n
II
locali y g oups (Lu z, Bohio, and S andley End).
Incomple e co e age
o
o aging anges may be
a
ac o a
Bohio and S andley End, bu no a Lu z, which is su ounded
by well-ne ed locali ies ha ha e ypical low Pa e n I ideli y
pe cen ages. O he pe iphe al locali y g oups such as Chapman,
S andley Ridge, and A mou End show low ideli y, sugges ing
ha pe iphe al loca ion is no he eason.
114SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE
7-8.—Fideli y o A ibeus jamaicensis o a pa icula locali y on BCI,
anked by numbe o ne ing nigh s. See Table 7-7 and ex o explana ion o
ca ego ies.
Locali y
PATTERN
la
(mo e han 19 ne ing nigh s)
Mille Ridge
Pla eau
Fue es
Shannon-AMNH
Ba bou -Hood
PATTERNIb
(less han 19 ne ing nigh s)
Chapman
S andley Ridge
Ba bou S eam
Con ad
Lake-Wheele
Ha a d
A mou End
D ay on End
Ze ek21
G oss Poin
Shannon-Balboa
O chid Island
PATTERN
II
Lu z
S andley End
Bohio
Rank
1
2
3
4
5
1
2
3
4
5
6
7
8
9
10
11
12
1
2
3
Ne ing
nigh s
52
46
26
24
20
17
16
14
14
12
11
11
10
6
5
4
4
197
25
14
60-100%
18
12
13
9
7
16
15
14
6
4
12
8
14
8
1
3
3
32
31
30
We sugges ha he high cap u e a e a Lu z is co ela ed
wi h he unusual local abundance o ig ees. Tha also could
be ue o Bohio because ou ne ing he e gene ally coincided
wi h he a ailabili y o ipe igs; howe e , ig ees we e sca ce
a S andley End. The abundance o ig ees a Mille Ridge
i als ha o Lu z, ye ideli y was low, as was ypical o o he
Pa e n I locali y g oups. Geog aphically, he Pa e n II locali y
g oups a e dissimila : Bohio is on he mainland, S andley End
is pe iphe al on BCI, and Lu z is mo e cen al. In e ms o size,
Lu z encompasses he la ges a ea, and because o i s p oximi y
o he Labo a o y Clea ing, i ecei ed he g ea es sampling
e o . Lacking a be e explana ion, one could a gue ha he
simila i ies in cap u e equency among he locali y g oups in
Pa e n II a e me ely coinciden al.
Mo emen s o Indi idual Ba s
Thus a we ha e discussed mo emen s o ba s associa ed
wi h a pa icula locali y g oup based on pooled da a. We also
examined he mo emen s o indi iduals cap u ed a wo o
mo e locali ies away om he cen al locali y unde conside a-
ion. To illus a e his we ha e mapped he mo emen s o i e
emale A. jamaicensis om Pe ia Blanca (Figu e 7-19). Each
emale was caugh a wo o mo e locali ies away om he ligh
owe , and polygons ou lining he loca ions whe e each was
cap u ed app oxima e hei known home anges. We hen
o e laid his map (Figu e 7-19) wi h a 0.5 km g id and coun ed
he numbe o imes indi idual polygons ouched each squa e.
The esul ing map (Figu e 7-20) shows he equency ( om one
o i e) o occu ence o indi idual ba s in each o he squa es.
We also mapped he same kind o in o ma ion om 11 ba s o
he day oos in he Buena Vis a ligh owe (Figu e 7-21).
This me hod p o ed e ec i e o illus a ing mo emen s
TABLE
7-10.—Age dis ibu ion o A ibeus jamaicensis cap u ed wo o mo e
imes a S andley-End, BCI.
Sex
Female
Male
Caugh only
when young
Caugh i s as young
la e as adul
Caugh only
as adul
14%
6%
38%
49%45%
Sex
FEMALE
numbe
pe cen
MALE
numbe
pe cen
TABLE
7-9.—Sexual a ia ion in age,
o mo e imes a S andley-End, BCI.
N J-S
29
73 1
1
Age du ing
cap u e span
J-A SAD
3 4
10 14
11 4
15 6
S-A
8
28
24
33
cap u e span, and numbe o cap u es o A ibeus jamaicensis caugh wo
AD
14
48
33
45
1
15
52
52
71
Cap u e span
(yea s)
2 3
7 4
24 14
48%
13 7
18 10
29%
4
3
10
1
1
Numbe o cap u es
a S andley-End
2
25
86
50
68
3 4 5+
3 1
10 4
14%
14 8 1
19 11 1
31%
2
11
38
31
42
To al
cap u es
3
9
31
20
27
4
8
28
10
14
5
1
3
8
11
6+
4
5
NUMBER 511115
45
i.••
•
i....
2 3
kilome e s
FIGURE 7-19.—Mo emen s o i e emale A ibeus jamaicensis om he day oos in he Pena Blanca ligh owe
o BCI and o he nea by a eas.
122SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
FIGURE 7-24.—Mean dis ance in kilome e s be ween cap u es o subadul emale A ibeus jamaicensis on and
nea BCI. Isolines enclose locali y g oups wi h simila mean dis ances and e lec dis ances be ween enclosed
cap u e s a ions and all o he cap u e s a ions. See ex o u he explana ion.

NUMBER 511123
kilome e s
FIGURE 7-25.—Mean dis ance in kilome e s be ween cap u es o subadul male A ibeus jamaicensis on and nea
BCI. Isolines enclose locali y g oups wi h simila mean dis ances and e lec dis ances be ween enclosed cap u e
s a ions and all o he cap u e s a ions. See ex o u he explana ion.
124SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
45 46 47 48 49 50 51 52 53 54 55
i...ii....
0 12 3 4 5
kilome e s
FIGURE 7-26.—Mean dis ance in kilome e s be ween cap u es o ju enile male A ibeus jamaicensis on and nea
BCI. Isolines enclose locali y g oups wi h simila mean dis ances and e lec dis ances be ween enclosed cap u e
s a ions and all o he cap u e s a ions. See ex o u he explana ion.
NUMBER 511125
47
46
4555
i.
••
•i...
0 12 3 4 5
kilome e s
FIGURE 7-27.—Mean dis ance in kilome e s be ween cap u es o ju enile emale A ibeus jamaicensis on and
nea BCI. Isolines enclose locali y g oups wi h simila mean dis ances and e lec dis ances be ween enclosed
cap u e s a ions and all o he cap u e s a ions. See ex o u he explana ion.
126
SMITHSONIAN CONTRIBUTIONS
TO
ZOOLOGY
TABLE
7-12.—Ca ches
o
A ibeus jamaicensis
and
o he ba s du ing episodes
o
ne ing
a wo o
mo e s a ions
on he
same nigh
on BCI.
Da e
29 No 78
HJun79
7 No 79
8 No 79
9 No 79
11
No 79
12 No 79
13
No 79
14
No 79
15
No 79
16 No 79
18
No 79
12
Oc 80
13
nigh s
Locali y
Lu z C eek
Snyde -Molino
0-2
Ba bou -La h op
16
Snyde -Molino
0-2
Mille
9
A mou -Ze ek
Je .
A mou -Ze ek
Je .
A mou -Con ad
Je .
Mille
9
Dona o
3-4
Mille
1-2
Ba bou 3
Dona o
3-4
Ba bou
3
Snyde -Molino-Lu z
S andley
15
S andley
21
Mille
4-5
S andley
21
Lu z C eek
Dona o
3-4
S andley
16
Dona o
3-4
S andley
16
S andley
16
Dona o
3-4
Mille
13-14
Mille
15
Mille
8-10
Chapman
6-9
30
si es
A ibeus
jamaicensis
26
56
109
6
77*
83*
2
27
11
47
19
24
42
27
5
10
6
2
10
1
94
103
14
68
23
42
56
30
101
49
1170
O he
ba s
15
27
33
4
47
16
3
27
18
54
25
14
47
22
3
13
10
4
26
10
136
82
27
100
54
45
40
32
12
12
958
* One
double
cap u e.
60 o 128 nigh s; see Table 7-13), only 2% ( ange, 0%-6%) o
indi idual A. jamaicensis we e cap u ed mo e han once a he
same ee on successi e nigh s. These ecap u ed ba s we e 5%
( ange, 0%-13%) o all cap u es o A. jamaicensis, which
ep esen s a pool o 7427 indi iduals amassed o e 128 nigh s.
In all, only 181 we e caugh a he same ee on wo successi e
nigh s, and se en we e cap u ed on h ee successi e nigh s.
Indi idual A. jamaicensis we e caugh mo e han once in 35 o
46 mul inigh ne ing episodes a single ipe ees.
The equency o epea s (ba s caugh mo e han once in a
nigh a he same s a ion) is simila o he equency o
ecap u es on successi e nigh s. A leas some A. jamaicensis
een e ed a ne wi hin minu es o elease, p obably because o
diso ien a ion esul ing om he handling p ocess and because
o p oximi y o he p ocessing s a ion o ne s. Ne e heless, he
imp ession was ha , once cap u ed, hese ba s a oided he ne s.
Du ing sampling pe iods o aling 133 nigh s in 1979 and 1980
we had 158 epea s o
A.
jamaicensis on 59 nigh s; an a e age
o 2.7 epea s on he nigh s wi h epea s. On he nigh s wi h
epea s, we caugh 4022 A. jamaicensis, 3.9% o which
ep esen ed epea ed cap u es. Repea ed cap u es ep esen ed
2.7%
o all A. jamaicensis caugh on all nigh s epea ed (133
nigh s, 5946 ba s).
I we could es ima e he p opo ion o ma ked and unma ked
ba s in a o aging agg ega ion we migh be able o es ima e he
numbe o
A.
jamaicensis ha come o a ui ing ee. We know
ha he ecap u e a e is low (<2%), bu we do no know wha
ou success a e is in cap u ing unma ked ba s. To judge om
he nigh ly a io o unma ked o ecap u ed ba s (Table 7-13),
hese a iables mus be changing om nigh o nigh .
I we assume ha many o he same ba s e u n nigh a e
nigh o a choice ee as long as ui emains, and ha we
ecap u e only a small ac ion o ma ked ba s, hen a ally o
TABLE
7-13.—Resul s
o
ne ing episodes in ol ing consecu i e nigh s
a
single locali ies
on BCI and he
adjacen
mainland.
The
columns
o
cap u e
eco ds,
mul iple
cap u es, and
mean cap u es
pe
nigh pe ain
o
A ibeus jamaicensis
alone. The
las
column,
o al cap u es
all species,
includes
A.
jamaicensis
and all
o he
ba s.
Da e
Jan 78
Ap 78
Jun78
No 78
May 79
No 79
Dec 79
Ap 80
Oc 80
To al
Feb78
Scp79
No 79
To al
Nigh s
N
3
4
3
2
2
2
4
4
3
27
3
2
4
9
N
291
97
85
106
79
94
253
145
319
1469
99
83
109
291
Cap u e eco ds
Ma ks
179
37
72
75
48
27
76
38
120
672
102
58
87
247
Recaps
117
63
16
32
32
67
182
109
208
826
201
141
196
538
To al
Mul iple
cap u es*
Indi iduals
N%
Reco ds
N «
Mille Ridge
(9
mul inigh episodes)
296
100
88
107
80
94
258
147
328
1498
5
3
3
1
1
0
5
2
9
29
2
3
4
1
1
0
2
1
3
2
10
6
6
2
2
0
10
4
18
58
S andley Ridge
(3
mul inigh episodes)
12
4
4
20
25
8
8
41
12
6
4
8
188
137
192
517
3
6
7
2
3
0
4
3
6
4
6
3
2
4
Mean
cap u es
pe
nigh
99
25
29
54
40
47
65
37
109
56
67
71
49
60
To al
cap u es
all
all
species
529
148
132
150
140
179
321
273
402
2274
274
205
448
927
NUMBER 511127
TABLE
7-13.—Con inued.
Da e
Dec 78
Ap 79
Sep80
Oc 80
To al
Jan 80
Aug80
Oc 80
To al
Sep79
Oc 79
No 79
Aug80
Aug80
To al
Ma 78
Ma 78
No -Dec 78
Ma 79
Oc 79
Oc 79
No 79
No 79
No 79
May 80
Aug80
Sep-Oc 80
Oc 80
To al
Aug80
Dec 78
Sep79
No 78
Sep80
Sep80
Ap 79
Ap 79
Ap 80
To al
To al o able
Nigh s
N
2
2
2
3
9
2
4
2
8
2
2
2
3
2
11
2
2
3
3
3
2
2
2
2
13
2
3
2
41
2
2
4
3
2
3
2
2
3
23
128
N
76
149
101
121
447
50
270
285
605
138
105
82
107
98
530
67
63
120
145
332
108
89
54
108
767
147
169
104
2273
147
99
326
86
199
283
165
151
130
1586
7427
Cap u e eco ds
Ma ks
64
106
88
56
314
11
176
142
329
89
65
19
75
64
312
16
19
88
54
183
45
34
15
40
637
96
93
49
1369
93
52
225
70
100
203
163
147
113
1166
4453
Recaps
13
46
18
69
146
39
102
146
287
51
41
63
35
34
224
52
44
35
99
161
64
55
39
68
157
51
80
55
960
56
52
119
19
105
86
2
6
27
472
3162
To al
Mul iple cap u es*
Indi iduals
N %
Reco ds
N
Bohio (4 mul inigh episodes)
77
152
106
125
460
1
3
5
3
12
1
2
5
3
3
2
6
10
7
25
Fue es (3 mul inigh episodes)
50
278
288
616
0
8
3
11
0
3
1
2
0
16
6
22
Pla eau (5 mul inigh episodes)
140
106
82
110
98
536
2
1
0
3
0
6
1
1
0
3
0
1
4
2
0
6
0
12
Lu z (13 mul inigh episodes)
68
63
123
153
344
109
89
54
108
794
147
173
104
2329
1
0
3
8
12
1
0
0
0
27
0
4
0
56
2
0
3
6
4
1
0
0
0
4
0
2
0
3
2
0
6
16
24
2
0
0
0
54
0
8
0
112
Miscellaneous (9 mul inigh episodes)
149
104
344
89
205
289
165
153
140
1638
7615
Ba bou -Hood
2
Chapman
5
1
5
RCS-AMNH
155
Lake-WMW
3
Con ad
6
4
3
A mou End
6 2
O chid Island
0 0
F ijoles Road
2
Gigan e
8
47
181
1
6
3
2
4
10
33
6
12
12
0
4
18
99
369
%
3
4
9
6
5
0
6
2
4
3
2
0
6
0
2
3
0
5
11
7
2
0
0
0
7
0
5
0
5
3
10
10
7
6
4
0
3
13
6
5
Mean
cap u es
nigh
39
76
53
42
51
25
70
144
77
70
53
41
37
49
49
34
32
41
51
115
55
45
27
54
61
74
58
52
57
75
52
86
30
103
96
83
72
47
71
60
To al
cap u es
all
species
160
177
156
176
669
119
325
342
786
181
393
101
131
112
918
159
143
178
260
419
149
189
90
271
969
171
230
172
3400
170
134
419$
240
234
351
1%
220
224*
2188*
11162
* Indi idual A. jamaicensis cap u ed wo o mo e imes du ing a mul inigh
ne ing episode.
One caugh h ee imes.
X
Th ee caugh h ee imes.
* Two caugh h ee imes.
• Se en caugh h ee imes.

128SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
TABLE 7-14.—La ge concen a ions o A ibeus jamaicensis a single ne ing
s a ions on BCI on successi e nigh s.
Locali y
Mille Ridge
Shannon-AMNH
Lu z
Mille Ridge
Lu z
Fue es
A mou End
Mille Ridge
Fue es
Da e
Jan 78
Sep79
Oc 79
Dec
79
May 80
Aug80
Sep80
Oc 80
Oc 80
Nigh s
N
3
4
3
4
13
4
3
3
2
A ibeus
jamaicensis
N
291
326
332
253
767
270
283
319
285
A e age
numbe pe
nigh
99
86
115
65
61
70
96
109
144
he numbe o indi iduals cap u ed du ing a mul inigh ne ing
sequence should sugges he numbe o ba s ha lock o a ee
(Table 7-14). We belie e ha , on occasion, hese numbe s mus
be la ge, and e en i we a e ca ching only a hi d o he ba s, he
o al a a ui ing ee may exceed a housand. Ou usual
imp ession a he ne s du ing a highly success ul nigh is ha
while he ai seems ull o ba s, ela i ely ew a e ge ing in o
he ne s.
G oup Mo emen s
The numbe o ba s a a ne ing s a ion, especially i i is a an
ac i e eeding oos nea a ui ing ee, a ies du ing he nigh ,
usually ab up ly. This is e idence ha he ba s a e mo ing
abou in g oups. Whe he hese g oups a e associa ions o
indi iduals ha o age oge he nigh a e nigh (and could be
conside ed ue locks), o a e me ely agg ega ions o ba s ha
o m a a eeding oos and shi oge he om one eeding si e
o ano he du ing he nigh and hen disband o inde ini e
pe iods o ime is no known.
The e is no doub abou he exis ence o agg ega ions.
Anyone who has much expe ience wo king wi h mis ne s in
he Ame ican opics has o be imp essed wi h ab up and
d ama ic shi s in numbe s o ba s du ing he nigh , signaling
he a i al and depa u e o g oups. Hei haus e al. (1974)
commen ed ha la ge ba s appea ed o a i e in g oups o eed
and ha isi s by g oups o ood sou ces was pulsed. We e e
o hese pulses as su ges, and we de ine a su ge as an ab up
inc ease in numbe s o ba s cap u ed, ollowed by an ab up
dec ease. When and whe he he e a e su ges depend on such
in luences as ain, p esence o ipe ui , he amoun o ligh ,
and he phase o he moon. Ou da a on su ges we e compiled
o h ee-mon h pe iods a he heigh o he ainy season
(Augus -No embe ) in 1979 and 1980 (n = 133 nigh s; means
summa ized in Table 7-15).
Admi edly hese da a a e biased and incomple e. On a nigh
wi h ew ba s we migh s op ne ing a e a ew hou s, hus
eco ding a nigh wi hou a su ge. On simila nigh s we
pe sis ed, pe haps wai ing o he moon o se and s ill did no
eco d a su ge; o , had he i s su ge commence as la e as
2300 h. We undoub edly missed some second su ges by
e mina ing oo soon a e he i s . Du ing he weeks o a ull
and new moon, moon ise and moonse o en coincided wi h he
beginning and end o su ges.
Ba s a e leas ac i e in he week ollowing he i s qua e o
he moon ( he week wi h mos ligh in ea ly e ening). Cap u es
we e ew (a e age 35.4 pe nigh ), su ges in equen and sho
(2.5 h), and only hal o he nigh s had any su ge a all (Table
7-15). Only 4% (one in 28 nigh s) had a second su ge. The i s
su ge was ea ly, beginning a app oxima ely 1915 h and ending
a abou 2045 h. P esence o ui and absence o ain seemed o
be o ela i ely li le consequence du ing nigh s wi h b igh
moonligh .
In spi e o inc easing ligh in ea ly e ening, A. jamaicensis
was mos ac i e in he week ollowing he new moon. Cap u e
a e was high (a e aging 53.8 pe nigh ) and su ges occu ed on
80%
o he nigh s. P obably because o he ea ly e ening ligh ,
second su ges occu ed on 20% o he nigh s. The i s su ges
we e o long du a ion, las ing abou 3.5 h, and hey began abou
1945 h and ended nea 2215 h. Mo ison (1978a) used he e m
luna phobia o desc ibe hese co ela ions o ac i i y and moon
phase.
Summa y
The ecap u e a e in ou 8907 A. jamaicensis a e aged 1.8
pe ba , indica ing ha less han 50% we e ecap u ed a all, and
less han 20% we e ecap u ed h ee o mo e imes. The low
ecap u e a e, long dis ances be ween ecap u e si es, and
po en ial home anges ha include he en i e island and pa s o
he adjacen mainland complica e mo emen analyses. We
adjus ed o some o hese di icul ies by pooling he da a o
each locali y.
We abula ed cap u es by species and by hal yea s on a
ma ix o locali y g oups and hen summa ized he en i e
ma king in e al (Oc obe 1976 h ough Oc obe 1980).
Al hough nei he ac o comple ely explains he a ia ion,
p oximi y o ne ing locali ies and di e ences in ne ing e o
led o ecap u e a es a ying om
1 %
o
43%.
Locali ies in he
cen e o he island consis en ly yielded be e ecap u e a es
han did ma ginal locali ies, sugges ing ha ba s li ing nea he
edge o he island also o age on he su ounding mainland. I
p o ed imp ac ical o use ne ing si es andomly on a g id on
BCI. Cap u e success was in luenced by ain, wind, cloud
co e , moon phase, opog aphy, and he dis ibu ion and
abundance o p e e ed oods. Thus, we had o y o
s anda dize cap u e e o by he use o such measu es as
ecap u es pe ne -nigh , o ne -hou . We se led on using he
numbe o
A.
jamaicensis caugh pe locali y as he mos use ul
and e ec i e measu e o cap u e e o .
The popula ion o A. jamaicensis on BCI is no an
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130SMITHSONIAN CONTRIBUTIONS
TO
ZOOLOGY
amo phous uns uc u ed mass o mobile ba s lying andomly
h oughou he island. Indi iduals a e mo e likely o be
ecap u ed close o whe e hey we e ma ked. Ba s ne ed a a
pa icula locali y include hose ha ha e day oos s in he a ea
as well as hose coming o o age a ui ing ees, and hose
caugh as hey a elled be ween day oos s and eeding si es
elsewhe e. We belie e ha he dis ibu ion o ig ees is a
majo in luence on he ela i e numbe o ecap u es a any
locali y.
We examined ideli y o a pa icula a ea (locali y g oup) by
analyzing sui es o mul iple cap u e eco ds o he 3846 ba s
ecap u ed and ound h ee pa e ns. Highes ideli y occu ed
whe e we sampled day oos s, a he han o aging si es.
Fideli y is a leas pa ially dependen upon ne ing e o . In
locali y g oups wi h unusually high ideli y, he sex a io was
skewed owa d males. In one such g oup, mos o he males
we e bachelo s, and mos o he emales we e membe s o
ha ems.
Da a on mo emen s o indi idual ba s was es ic ed by he
ela i ely small numbe o mul iple ecap u es. The a e age ba
lies on abou 700 nigh s du ing i s li e ime, ye we cap u ed a
ba only an a e age o 1.8 imes; a e y na ow window
h ough which o examine i s mo emen s. Radio acking and
cap u e o ba s wi h known day oos s showed a e age
mo emen s o 1-4 km be ween day oos and eeding si e.
Mean dis ances be ween cap u es show no co ela ion wi h sex
o age. P opo iona ely, eco ds o mo emen s by ju eniles
we e much mo e equen han expec ed and eco ds o adul s
a e abou he same o a li le less han expec ed, pe haps
e lec ing he ela i e di icul y in ecap u ing olde
adul s.
The
longes dis ance we eco ded be ween cap u es was abou 6 km
o h ee indi iduals. Howe e , hese we e no single-nigh
mo emen s; he e o e, hese dis ances may ep esen dispe sion
ins ead o ac ual o aging dis ances.
A ibeus jamaicensis o en isi mo e han one ee in an
e ening. We ecap u ed one indi idual a a ui ing ee 1.2 km
away om ano he ee whe e she had been cap u ed ea lie he
same e ening. Recap u e a es we e only 2% o ba s cap u ed
mo e han once a he same ee on successi e nigh s, and
simila ly low o ba s ecap u ed he same nigh a he same
s a ion.
The numbe o ba s a a ne ing s a ion shi ed du ing he
nigh , usually ab up ly, sugges ing ha he ba s a e mo ing and
o aging in g oups. These su ges o ac i i y seemed in luenced
by ain, p esence o ipe ui , and phase o he moon. Clea ly,
ba s we e leas ac i e in he week ollowing he i s qua e o
he moon (mos ligh ea ly in he e ening), and mos ac i e in
he week ollowing he new moon.
8. Roos ing Beha io
Douglas
W.
Mo ison and Cha les
O.
Handley, J .
In his sec ion we p esen a syn hesis o ou ma k- ecap u e,
adio- acking, and nigh - iewing-scope obse a ions pe inen
o day oos s o A ibeus jamaicensis on Ba o Colo ado Island
(BCI).
Fo compa ison, we ha e included da a on o he species
o ba s whe e app op ia e. We p esen a simila syn hesis o
obse a ions on nigh oos s as pa o he discussion in Sec ion
9, Fo aging Beha io .
Ma k- ecap u e and adio- acking echniques ha e inhe en
s eng hs and weaknesses. Wi h adio- agging, he beha io o
indi iduals can be in ensely moni o ed o days o weeks.
Howe e , sample sizes end o be small and he e is no
independen way o measu e how much he beha io o an
indi idual has been al e ed by he ansmi e . In con as ,
long- e m ne ing s udies can gene a e huge samples, bu he
da a a e some imes di icul o in e p e .
Taken oge he , he wo echniques complemen each o he
in impo an ways. Ques ions p o oked by adio- acking
obse a ions can p o ide a amewo k a ound which o analyze
ma k- ecap u e da a and ne ing da a p o ide independen
con i ma ion ha he beha io o adio- agged indi iduals is
ypical o he popula ion as a whole. Using he wo echniques
in conce has inc eased ou con idence in he accu acy o he
pic u e ha has eme ged.
Ba s show di e se oos ing beha io s, o en using di e en
kinds o oos s o di e en pe iods o hei daily and annual
cycles (Kunz, 1982). Day oos s o e p o ec ion om p eda o s
and he elemen s and a e gene ally used o ex ended pe iods.
In con as , nigh oos s may be less p o ec ed, empo a y si es
chosen o hei p oximi y o ood sou ces. Indi idual ba s also
may change oos s on a seasonal basis, wi h ce ain ypes o
oos s a o ed o ma ing, ea ing young, o o he ac i i ies.
Day- oos ing Si es
A.
jamaicensis appa en ly is oppo unis ic in i s selec ion o
day- oos ing si es. Whe e ca es a e a ailable, i oos s in la ge
g oups (Dalques , 1953; Kunz, 1982; Tu le, 1968). In o es ed
Douglas W. Mo ison, Depa men o Zoology and Physiology,
Ru ge s Uni e si y, 195 Uni e si y A e., Newa k, NJ. 07102.
Cha les O. Handley, J ., Na ional Museum o Na u al His o y,
Smi hsonian Ins i u ion, Washing on, D.C. 20560.
habi a s, he la ges g oups oos by day in ee hollows, while
smalle g oups and indi iduals a e ound in oliage (Mo ison,
1979).
In he mois opical o es o BCI, we loca ed 25 day
oos s used by 18 adio- agged A. jamaicensis. Six een we e in
oliage and nine in ee hollows.
DAY
ROOSTS
IN
FOLIAGE.—Foliage
oos s we e used
p ima ily by males. Se en adio- agged males made ansien
use o a a ie y o si es in he oliage, ypically occupying a si e
o 3-5 days ( ange 1-13 days) be o e mo ing o ano he
oliage si e 100 m o a he away. In con as , emales
no mally oos ed in ee hollows and a ely used oliage si es.
O 11 adio- agged emales, wo oos ed in oliage bu only
a e being cap u ed and adio- agged as hey eme ged om
hei day oos s in ee hollows.
The kinds o oliage oos s used by A. jamaicensis (a 45 g
ba ) a e ypical o hose used by canopy- o aging ui ba s
(Goodwin and G eenhall, 1961) and we e indis inguishable
om hose used by A. li u a us (70 g) and Vampy odes
ca accioli (36 g) on BCI (Mo ison, 1979, 1980a). Foliage
oos s used by adio- agged A. jamaicensis on BCI included
shel e s unde a long, a ching ond o a palm, Oenoca pus
panamanus; unde an "umb ella" o med by a single, wil ed
lea o a b oadlea epiphy e; in he c own o
a
spiny black palm,
As oca yum s andleyanum; and in a shallow shel e o med by
he o es canopy.
Radio- agged V. ca accioli (nine sigh ings) in a iably
oos ed in g oups o h ee o ou adul s, 7-12 m abo e he
g ound, unde he umb ella-like c owns o unde s o y ees
(12-20 cm Diame e a B eas Heigh (DBH)). The day oos s
o adio- agged A. li u a us (26 sigh ings) we e mo e a iable,
om 2.7-28 m abo e he g ound in a a ie y o si ua ions, such
as unde b oken o c ossed onds o Oenoca pus panamanus,
in ine- angled c owns o subcanopy ees, in ca elike ecesses
on he unde side o he c owns o canopy-heigh ees, and in
b anches o e hanging he wa e along he lakesho e.
As a iable as hese oos s migh seem, all had wo impo an
ea u es in common: hey we e di icul o see om he g ound
and hey had uns able suppo ing s uc u es. Despi e hei open
appea ance, oliage oos s may be almos as e ec i e as ee
hollows o educing he exposu e o adul ba s o p eda ion and
ain. The dense lea co e o ming he oo o hese ecesses is
131
138SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
de ec o eco dings o ac i i y (Fen on, Boyle e al., 1977).
Ecke (1982) a gued pe suasi ely ha all such e ec s a e
adap a ions o he a oidance o isually o ien ing p eda o s.
Longe - e m Fo aging Pa e ns
Pa e ns o ui ee use we e mos appa en on b igh
moonli nigh s because lying was limi ed and eeding passes
seemed especially egimen ed. Occasionally on such nigh s, a
adio- agged A. jamaicensis made wha appea ed o be a
eeding pass in he w ong di ec ion. In some cases, hese ligh s
we e simply eeding passes o ano he ui ing ee, demons a -
ing ha ba s some imes use a single nigh oos while eeding
on mo e han one ee. O , i hese we e non eeding passes hey
may ha e been econnaissance ligh s o gain in o ma ion
abou u u e eeding si es.
Al hough ba s e u ned o eed on he same ui ee o as
many as eigh consecu i e nigh s, hey discon inued eeding
isi s when he densi y o he igs declined o less han one o
wo pe squa e me e (n = 6), unless a ig ee was less han 150
m om he day oos . In swi ching ui ees, a ba ypically
made no mo e han he i s eeding pass o he nigh o he
p e iously used ee and hen immedia ely, wi hou sea ch
lying, swi ched i s eeding oos o eeding passes o a new
ee.
In 11 ins ances his shi was o an a ea ha he ba had
isi ed om 1-8 (4.2 ± 2.2) nigh s p e iously. In i e o hese
11 cases, he shi was o a ig ee ha had almos ce ainly
been he objec o ea lie econnaissance ligh s.
Flying by adio- agged A. jamaicensis du ing he b igh hal
o he luna mon h was educed o he minimum leng h o ime
equi ed o ge o a eeding oos (usually om one o i e
minu es). E en on moonless nigh s, he ba s o en made only
sho ligh s and spen mos o hei ime hanging a he eeding
oos s. Occasionally, howe e , bou s o sus ained lying ha
las ed 10-45 minu es we e eco ded. Six o 40 all-nigh
acking sessions con ained a leas one such pe iod o
p olonged lying. All bu one o hose longe ligh s occu ed
du ing he da k hal o he luna mon h. The one excep ion
occu ed in he e ening be o e a la e (2345 hou s) moon ise.
P olonged ligh s began ei he om nigh eeding oos s (n =
3) o upon eme gence a dusk om day oos s
(jn =
3). The a ea
co e ed by hese ligh s could only be app oxima ed, as i was
no possible o iangula e he ba s' cons an ly changing
posi ion. Usually he lying was con ined o he wa e shed in
which i began (n = 4), bu once i shi ed in o an adjacen
wa e shed and ano he ime in o a nonadjacen wa e shed.
These long sea ch ligh s undoub edly inc ease he chances o
encoun e ing newly ui ing ees and migh ha e been ini ia ed
o ha pu pose.
Feeding Roos s
T iangula ion by adio o o aging A. jamaicensis sugges s
ha eeding oos s usually a e in one o a ew a o ed ees,
wi hin a 0.25 o 0.5 ha a ea, 25 o 200 m om a ui ing ee.
Mo ison (1975) was able o pinpoin and con i m, by on-si e
inspec ion, wo eeding oos s used by his adio- agged A.
jamaicensis. One was in he c own o a 13-m- all, spiny black
palm (As oca yum s andleyanum). The o he was abou 3 m
abo e he g ound unde he long, a ching ond o an
Oenoca pus panamanus. Bo h si es could ha e been selec ed
o hei educed accessibili y o e es ial p eda o s (see
Sec ion 8, Roos ing Beha io ).
A e Al ed Ga dne disco e ed ha ba s d op pelle s o
chewed pulp when eeding on igs, Ba P ojec pe sonnel
ou inely used accumula ions o pelle s o ecognize si es used
as eeding oos s. Depending on he heigh om which hey a e
d opped, he pelle s may be in small piles, dis inc clus e s, o
b oadly dispe sed. The a ea co e ed can be qui e ex ensi e,
some imes exceeding 100 m2. These eeding oos s a e o en
loca ed along idges abo e ui ing ees, bu may be ound
elsewhe e, especially whe e young palms o subcanopy ees
a e nume ous. The single common ea u e is an open
unde s o y ha does no es ic ligh . This may explain why
we ha e ound so many eeding oos s along o adjacen o
ails,
which also unc ion as open ligh pa hs o ba s.
Radio- agged A. jamaicensis we e ela i ely ai h ul o
eeding oos s. In a iably he same eeding oos a ea was used
nigh a e nigh o as long as a ba ed om he same ui ee
(n = 14 ui ing ees). Fou adio- agged ba s made o aging
passes om he same eeding oos o wo simul aneously
ui ing ees 35-220 m apa , and wo ba s used he same
eeding oos a eas when hey e u ned a e h ee and eigh
nigh s, espec i ely, o ge ui om ano he ee in he
neighbo hood.
The e a e se e al possible, bu un es ed, explana ions o his
appa en ideli y o eeding oos a eas. I migh simply be he
esul o he pa chy dis ibu ion o ege a ion p e e ed o
eeding oos s. O i migh be ha A. jamaicensis is a c ea u e
o habi , p e e ing o use amilia si es along well-known ligh
pa hs.
The la e hypo hesis is suppo ed by ou obse a ion
ha adio- agged A. jamaicensis equen ly did no use (o
ailed o ind) a ig ee p oducing ipe ui close o hei day
oos s. Ins ead hey commu ed wo o h ee imes a he o
ano he ee. This sugges s ha many ui ing ees a e ound
because hey s and along amilia o aging ou es.
Feeding oos s may ha e o he unc ions no ela ed o
eeding. Fo example, i hey a e close o a ui ing ee han
he day oos , hey may se e as con enien ha ens du ing
b igh moonligh . This a leas seems o be he case wi h A.
li u a us and
V.
ca accioli, wo ui ba s ha ha e day oos s in
oliage (Mo ison, 1980a). Feeding oos s also may be si es o
social in e ac ion. Adul bachelo males may y o copula e
wi h emales a eeding oos s away om he day oos and i s
de ending ha em male because hey p obably lack o he access
o emales (Mo ison and Mo ison, 1981). Howe e , emales
may be less ecep i e o males encoun e ed ou side he
day- oos hollow i compe i ion o hollows selec s o males

NUMBER 511139
who si e s onge , mo e agg essi e, o o he wise i e young.
Also,
day oos s a e p obably sa e si es o copula ing ba s
han a e nigh oos s.
G oup Fo aging
Feeding oos s migh be "in o ma ion cen e s" o ood
inding (sensu Wa d and Zaha i, 1973). In heo y, A.
jamaicensis is a likely candida e o g oup o aging in o ma ion
sha ing. A p e equisi e o he e olu ion o in o ma ion cen e s
is ha ood be ound in sho -li ed, locally supe abundan
pa ches. This so o ood dis ibu ion means ha use s mus
con inually ind new ood pa ches. Howe e , because he
pa ches con ain so much ood (and o so sho a ime), no hing
is gained by ying o conceal o de end a ood pa ch once i is
ound. Fig ees i his desc ip ion. A single ig ee on BCI
may bea 40,000 igs, o which a single A. jamaicensis would
likely ca y away no mo e han 100 (Mo ison, 1978d) be o e
he ui s we e gone o spoiled.
A second cha ac e is ic ha makes A. jamaicensis a po en ial
use o in o ma ion cen e s is ha o aging ba s agg ega e a
nigh ( eeding) oos s. A nigh oos may be occupied by
dozens, i no hund eds, o ba s, and hey may be making
eeding passes o se e al di e en ui ing ees.
A simple mechanism o in o ma ion exchange is plausible.
I an
A.
jamaicensis, which has been success ul in inding a ig
ee,
e u ns o i s eeding oos wi h a ig o he odo o igs,
any oos ma es who had been unsuccess ul in inding ood
could ollow he success ul ba ou on i s nex eeding ligh .
The in o ma ion sha ing would no need o be in en ional, bu
he in o ma ion would g ea ly bene i he unsuccess ul ba and
would no signi ican ly deple e he ood a he ee o he
success ul ba . In a g oup ha emained oge he o e a pe iod
o ime, i is possible ha onigh 's success ul ba could be
omo ow nigh 's unsuccess ul ba , and ice e sa, so all would
gain o e he long e m. No e also in his con ex he p obabili y
o ood inding econnaissance ligh s ini ia ed in ad ance o
need.
Despi e he heo e ical po en ial, i is s ill unknown whe he
A. jamaicensis on BCI o ages in cohesi e g oups. Ce ainly
his ba does o age in g oups, as he nigh ly ebb and low o
ba s a a ui ee demons a es (see Sec ion 7, Mo emen s).
Howe e , whe he he locks ha swi l abou a ui ee s ay
oge he o hou s, all nigh , se e al nigh s, o o long pe iods
o ime is unknown.
Mo ison and Mo ison (1981) acked a g oup o h ee and
ano he o ou adio- agged emale A. jamaicensis om wo
di e en ha ems o eigh comple e nigh s and ound no
e idence o cohesi eness o ha em membe s away om he day
oos . These emales le and een e ed hei day oos hole
indi idually. Because eme gences we e ypically mo e han a
minu e apa , each emale eme ged a e he p e ious one had
disappea ed om he day oos a ea. On h ee o he eigh
nigh s, h ee o ou emales did isi he same ipe- ui -
bea ing Ficus insipida in he cou se o he nigh , bu hey
mo ed be ween ui ees independen ly and did no oos
anywhe e nea each o he while eeding. This adio- acking
s udy sugges s ha ha em emales do no o age in g oups, a
leas no du ing he ci cums ances ope an when ou obse a-
ions we e made.
Ne e heless, i is s ill possible ha o he sex and age classes
o age in g oups, o ha A. jamaicensis o age oge he in
habi a s whe e ui ing ees a e no as abundan and easy o
ind as hey a e on BCI. In Mexico and Cos a Rica, o
example, clumped mis -ne ing cap u e imes ha e been
in e p e ed o mean ha he A. jamaicensis he e o age in
g oups (Dalques , 1953; Hei haus e al., 1975). G oup o aging
has been epo ed in lowe - eeding ba s in B azil (Sazima and
Sazima, 1977) and A izona (Howell, 1979).
I should be possible o de ec o aging-g oup associa ions
by analysis o he occu ence o "double ecap u e pai s" in he
ma k- ecap u e da a. A "double ecap u e pai " consis s o wo
ba s ha we e cap u ed a he same ne ing si e on he same da e
and we e subsequen ly ecap u ed oge he a ano he si e some
days,
mon hs, o yea s la e . Mo ison (1975) ound 22 such
pai s among he 259 ecap u es o 1472
A.
jamaicensis he and
Bonacco so ma ked o e a 14-mon h pe iod in 1972 and 1973.
Wha is he p obabili y ha his numbe o double ecap u e
pai s migh occu simply as a esul o chance associa ions in a
popula ion o independen ly o aging ba s? Gi en ha wo o
mo e ecap u es a e made on nigh n, le Pn s and o he
condi ional p obabili y ha any wo o hese ecap u es we e
cap u ed oge he p e iously (Mo ison, 1975). I (n - 1) = he
numbe o p e ious cap u e nigh s, { = he numbe o ba s
cap u ed on p e ious cap u e nigh i, whe e l<i<n - 1, and y =
he o al numbe o ba s banded o nigh n, hen
P = y 1)
n S
S y (y-D
The numbe o double ecap u e pai s Cj ha could be expec ed
o occu by chance on any nigh n is simply Pn imes he o al
numbe o pai combina ions o ecap u es on nigh n, o Pa
imes c2 , whe e zn is he numbe o ecap u es on nigh n. The
o al nume o ecap u es expec ed o occu by chance du ing
he en i e 14-mon h ne ing p og am is he sum o he P n. c/
alues o all 60 o he 131 sampling nigh s in which wo o
mo e ecap u es we e made. This sum was calcula ed and ound
o equal 14.2, he numbe o double ecap u e pai s expec ed o
occu simply by chance. The obse ed alue o 22 was no
signi ican ly g ea e han ha expec ed om andom asso -
men as de e mined by a one- ailed binomial es , P = 0.195 ( he
a p io i expec a ion o highe han andom asso men ,
jus i ied he use o a one- ailed es ). Thus, Mo ison concluded
ha A. jamaicensis does no o age in g oups, a leas no in
g oups wi h membe ships su icien ly cons an o e he long
e m o be de ec ed by his me hod.
Howe e , his conclusion was based on he un ealis ic
140SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
assump ion ha he e is no mo ali y o ma ked ba s. The
chance ha a ma ked ba will die educes he expec ed numbe
o double ecap u e pai s and so inc eases he signi icance le el
o he obse ed alue. To e lec his eali y we used he
equa ion
a~
He e i is easonable o make s equal o 0.57, he annual
su i o ship o adul A. jamaicensis (see Sec ion 6, Popula ion
Es ima es), and o £ we use 0.32 yea s, he a e age in e al
be ween ecap u es o Mo ison's (1975, able 1) da a. The
expec ed alue now d ops om 14.2 o 9.9 [14.2 x (0.57)0-64].
The obse ed alue o 22 double ecap u e pai s is signi ican ly
g ea e han his e ised expec ed alue (one- ailed binomial
es , P < 0.05).
Un o una ely, e en i hese e ised calcula ions e eal an
incidence o double ecap u e pai s signi ican ly g ea e han
expec ed, we s ill canno be su e ha his di e ence is due o
he exis ence o o aging g oups. O he ac o s could p oduce
he same esul . Fo example, he p obabili y o obse ing a
double ecap u e pai would be inc eased i independen ly
o aging ba s equen ed he same eeding a ea. Two ba s migh
equen he same a ea because i is especially a ac i e (e.g.,
he g ea Lu z ig pa ch) o because i is nea he day oos s o
bo h ba s. The calcula ions o si e ideli y (see Sec ion 7,
Mo emen s) indica e ha he cap u e si es we e no in equally
a ac i e a eas and ha indi idual ba s we e no equally likely
o be cap u ed a all si es. Fu he mo e, we suspec ha a e he
i s cap u e, ba s end o a oid mis ne s because hey a e
seldom ecap u ed (see Sec ion 6, Popula ion Es ima es).
Recap u e pai s migh be oge he a ui ing ees many imes
be o e one o bo h a e ecap u ed again.
These inequali ies end o inc ease he expec ed numbe o
double ecap u e pai s. Wi hou some es ima e o he magni-
ude o hese biases, any es o o aging g oups based on he
incidence o double ecap u e pai s will be inconclusi e.
Thus,
al hough clus e ed cap u e imes and double ecap u e
pai s sugges g oup o aging, we lack su icien e idence o
show ha hese associa ions a e any hing mo e han coinciden-
al.
A powe ul es o g oup o aging could be based on di ec
obse a ion o he o aging mo emen s o a la ge numbe o
adio- agged ba s cap u ed oge he a he day oos o cap u ed
in "clus e s" while o aging. This emains o be done.
Summa y
The usual pa e n o an A. jamaicensis is o lea e he day
oos hal an hou a e sunse , ly o a ui ee, ake a ui o
a nea by eeding oos , consume i (d opping d y pelle s o he
g ound benea h), make se e al mo e eeding passes o he ui
ee,
hen e u n o he day oos hal an hou be o e sun ise. On
da k moon nigh s, a ba may isi as many as i e eeding a eas.
On nigh s wi h b igh moonligh , ba s isi only one o wo
eeding a eas, and se e al hou s may be spen back in he day
oos du ing he b igh es hou s. Bo h luna phobia and he use
o eeding oos s sepa a e om he ui ing ee may educe
exposu e o p eda ion.
Radio- acking obse a ions sugges ha ba s some imes use
he same eeding oos o make eeding passes o mo e han one
ui ing ee. The ba s also appea o make b ie econnaissance
ligh s o assess he condi ion o p e iously loca ed ui ees.
The ba s spend mos o hei ime hanging in he eeding oos
and seem o minimize eeding ime e en on da k moon nigh s.
P olonged ligh s o 10-45 minu es (sea ching o new ees?)
we e a e and occu ed only in he absence o b igh moonligh .
Feeding oos s end o be g oups o a o ed ees 25-200 m
om a ui sou ce, and mos a e loca ed in a eas o open
unde s o y. Ba s a e ai h ul o indi idual eeding oos s as long
as he nea by ui ing ee emains p oduc i e. This ideli y may
esul om pa chy dis ibu ion o p e e ed oos ing ees, o
may simply e lec he habi ual use o a amilia a ea.
The ba s clea ly cong ega e a ood sou ces, bu whe he
cohesi e, long- e m g oup o aging occu s is unknown.
Clus e ed cap u e imes and double ecap u e pai s sugges
g oup o aging, bu di ec obse a ion o he phenomenon is
lacking. Radio- acking da a on emales om he same ha em
show no e idence o such locking.
Whe he o no A. jamaicensis o ms o aging g oups, he
po en ial exis s o an exchange o ood loca ion in o ma ion
among indi iduals a eeding oos s. Fo aging heo y sugges s
ha eeding oos s could be in o ma ion cen e s o inding
ees wi h ipe ui (sho -li ed, locally supe abundan ood
pa ches), especially i he ba s using a nigh oos a e making
eeding passes o di e en ui ing ees.
10.
Food Habi s
Cha les
O.
Handley, J ., Al ed
L.
Ga dne ,
and Don E. Wilson
Ini ially,
he Ba
P ojec ocused en i ely
on
cap u ing
and
ma king ba s,
and
li le a en ion
was
gi en
o
ood habi s
and
o he aspec s
o
na u al his o y. G adually
i
became appa en
ha cap u e a e usually
was
linked
o he
o aging beha io o
he ba s
and o he
loca ion
o
ne s
in
ela ion
o
ood sou ces
and eeding oos s. Consequen ly,
Ba
P ojec pe sonnel ook
inc easing in e es
in he
con en
o
he ba s' eces,
in
e idence
and loca ion
o
eeding oos s,
in
ood i ems ca ied by ba s in o
ne s,
and in he
loca ion
and
phenology
o
ui ing ees.
As a
esul , cap u e e o became mo e p oduc i e
in
numbe o ba s
ma ked
and
ecap u ed. Al hough
ou
da a
on
ood habi s lack
he p ecision
o
he smalle -scale s udies
o
Bonacco so (1979)
and Hei haus
e al.
(1975), hey
do
o e insigh s
on
ood
p e e ences, seasonali y
o
ood a ailabili y,
and he
o aging
habi s o A ibeus jamaicensis on Ba o Colo ado Island (BCI).
P ojec pe sonnel e en ually es ablished
a
p o ocol ha
included sea ching
o
e idence
o
ba ac i i y be o e selec ing
si es
as
cap u e s a ions. Along
he
ails
we
wa ched
o
pelle s
("ba chop") ha s enode ma ine ba s spi ou a e chewing
and
p essing he juice om he pulp
o
ui s such as Ficus insipida,
F. yoponensis,
and
Spondias adlko e i.
The
pelle s appea
in
disc e e piles
o
loose clus e s on
he
g ound when he ba s d op
hem om oos s
low in he
subcanopy,
o
hey
may be
sca e ed,
as
hough b oadcas , when hey a e d opped om
he
canopy. Because
he
ba s e u n again
and
again du ing
he
nigh
o he
same pe ch, piles
o
pelle s
may
ep esen many
ui s.
We also wa ched
o
agmen s
o
pa ly ea en ui s; la ge
seeds such
as
hose
o
Spondias mombin,
S.
adlko e i,
and
Dip e yx panamensis disca ded
by
ba s a e hey
had
sc aped
o
he
pulp; cas
o
skins
o
ui such
as
Qua a ibea
as e olepis;
and
ees d opping ipe ui
o
kinds known
o be
a o ed by ba s. Cong ega ions
o
noisy diu nal ugi o es such
as monkeys (Aloua a pallia a
and
Cebus capucinus), guans
Cha les
O.
Handley,
J ., and Don E.
Wilson, Na ional Museum
o
Na u al His o y, Smi hsonian Ins i u ion, Washing on,
D.C.
20560.
Al ed
L.
Ga dne , NERC,
US.
Fish
and
Wildli e Se ice, Na ional
Museum o Na u al His o y, Washing on,
D.C.
20560.
(Penelope pu pu ascens),
and
pa o s (Amazona spp.) o en
led
o
he
disco e y
o
ees wi h ipening ui .
We le
he
ails
o
sea ch unde ce ain ees (e.g.,
Oenoca pus panamanus Bailey
and
Gus a ia supe ba
(Hum-
bold , Bonpland,
and
Kun h) (Be g) pa icula ly a o ed
by
ba s
as
eeding oos s.
We
ound ha pa ches
o
hese ees
associa ed wi h se e al ui ees we e used epea edly
by A.
jamaicensis
as
eeding oos s. Thus,
in
he cou se
o
a yea ,
he
ba s
may use he
same eeding oos many imes. Taking his
in o accoun ,
we
ou inely inspec ed known eeding oos s
in
addi ion
o
walking
he
ails
in
sea ch
o
new si es.
F ui s Used
as
Food
Ou obse a ions
on
ui s ea en
by A.
jamaicensis, mos ly
la e
in he
ainy seasons (Augus
o
No embe )
o 1979 and
1980,
a e
summa ized he e (Figu e 10-1
and
Table 10-1).
Ficus insipida Willdenow: Bonacco so (1979), Fleming
(1971),
Mo ison (1978d),
and ou
own obse a ions ag ee ha
in cen al Panam£
F.
insipida
is he
a o i e ood
o A.
jamaicensis (Figu e 10-1),
A.
li u a us,
and
pe haps Vampy-
odes ca accioli,
and is
ea en
in
lesse amoun s
by
se e al o he
ba s.
I s
ui p oduc ion
is
aseasonal,
bu he
c op
is
limi ed
in
Oc obe
and
om Janua y h ough Ma ch (Table 10-1).
Few
ui s each ma u i y
in
Augus
and
Sep embe ,
wo
mon hs
when
F.
insipida does
no
seem o
be a
signi ican ood
o
ba s.
La ge concen a ions
o
ba s, mos ly la ge s enode ma ines,
we e ound nea F. insipida bea ing ipe ui
in
Ma ch, Ap il
( ou si es),
May,
June, Oc obe
( wo
si es), No embe
( wo
si es)
and
Decembe . No mally he ba s pick so , ag an , ully
ma u e ui ,
bu
occasionally
(as was
no ed
on 11
Janua y,
8
Feb ua y, 21 Oc obe ,
and 7-9
No embe when
ew
sui able F.
insipida we e a ailable)
A.
jamaicensis ca ied pa ly ea en
small, ha d,
and
la ex-laden un ipe ui s in o
he
ne s.
Ficus yoponensis Des aux: Simila
o F.
insipida
in
abundance
and
seasonal a ailabili y
on BCI
(Table 10-1),
he
small- ui ed F. yoponensis
is a
a o i e
o
smalle ugi o es
such
as
U ode ma biloba um, Vampy essa pusilla,
V. nym-
phaea,
and
Vampy ops helle i
and is
consumed
in
g ea
141
142
Ficus inslpida
Ficus yoponensis
Ficus
sp.
Ficus lgona a
Ficus ob usi olia
Ficus cos a icana
Ficus popenoel
Dlp e yx panamensis
Qua a lbea as e olepis
Spondias mombin
Spondias adlko e i
Poulsenla a ma a
Anaca dium excelsum
Calophyllum longi olium
Cec opla
sp.
Uniden i ied ui pulp
Uniden i ied seed
Pollen
on u
JAN
FEB MAR
SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
APR
MAY AUG SEP OCT NOV DEC
3
A
FIGURE 10-1.—Seasonal dis ibu ion
o
oods
o
A ibeus jamaicensis on BCI obse ed du ing he pe iod
1977-1980. Squa es ep esen seedless eces, ci cles ep esen eces wi h seeds, and iangles ep esen ui
ca ied in o ne s. Numbe s ep esen obse a ions pe mon h.
quan i ies by la ge s enode ma ines when
F.
insipida is sca ce.
Al hough we caugh la ge numbe s o
A.
jamaicensis a ui ing
F. yoponensis, we o en had he expe ience o ne ing unde a
ee d opping ipe igs and inding he a ea de oid o
ba s,
o o
ca ching A. jamaicensis ca ying F. insipida om some mo e
dis an ee. Ba s wi h eces con aining seeds o
F.
yoponensis
we e ou numbe ed by abou wo
o
one by ba s wi h eces
con aining F. insipida. Howe e , la ge concen a ions o ba s
we e ound a ipe F. yoponensis in May, Oc obe ( wo si es),
and No embe ( h ee si es).
Ficus dugandii S andley: On he b eezy nigh s o 25 and
26 Oc obe 1979,
a
gian F. dugandii, wi h a c own eme ging
abo e he canopy nea he highes poin o he island, a ac ed
(pe haps
by
wind-wa ed odo ) la ge locks
o
unma ked
s enode ma ines, p esumably om he mainland. In mis ne s
benea h he ee, which was d opping many ipe ui s, we
cap u ed only 106 A. jamaicensis,
7
A.
li u a us, and
2
V.
ca accioli, while he ca ch o smalle s enode ma ines o alled
an as onishing 265, including 138 U ode ma biloba um,
90
Chi ode ma illosum, 15 Vampy essa nymphaea,
6
V.
pusilla,
1 Vampy ops helle i,! A ibeus phaeo is,
1
A. wa soni, and he
only A. ha ii e e aken on BCI.
Ficus igona a Linnaeus: The ed-spo ed, small-seeded
ui o Ficus igona a was an impo an ood o
A.
jamaicensis
and C. illosum a he heigh o he ainy season in Sep embe
when ui o
F.
insipida and F. yoponensis was sca ce (Table
10-1).
Many ui s o
F.
igona a we e ca ied in o he ne s and
i s seeds we e common in eces (Figu e 10-1).
F.
igona a also
occasionally was ound
in
he eces
o
Ca ollia pe spicilla a
and Phyllos omus has a us. Al hough commonly seen
in
Sep embe , he lean mon h, he ui o
F.
igona a seemed no
o domina e he die o any ba in ha mon h sa e possibly
Chi ode ma illosum. Spondias mombin
and
Qua a ibea
as e olepis we e ea en mo e commonly. One F. igona a was
NUMBER
511
143
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2
3
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-3
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oo
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e
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be s
i
z
ON
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•i
:cies
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o
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eg
he e
i
0000^0000 00000000000 000000
00000—0—0 00000000000 000000
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— O
OOO S
—
—
—i S
— S — <S S —< — S
•-•CS
—
00000 — 000 000000000 — 0 000000
000000000 00000000000 000000
(SOO —
0000 — 0000 0000000000 0000000
000000 — 00 00000000 — 0 0000000
—
en — —
csen
—
O Sencs-^aocScscn
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— en N O O >• O O >.
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5 i 1

144SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
ca ied in o a ne by an A. phaeo is in Ma ch.
Ficus ob usi olia Humbold , Bonpland, and Kun h: The
la ge el e -skinned ui s o F. ob usi olia we e ca ied in o
ne s occasionally by A. jamaicensis, A. li u a us, and C.
illosum. The ui s seemed o be mo e a ac i e o he smalle
ugi o es such as C. illosum. We ound ba s concen a ed a
an
F.
ob usi olia wi h ipe ui in Ma ch.
Ficus popenoei S andley: The el e y, oblong ui s o F.
popenoei a e uncommon on BCI, and we ound only a ew ees
wi h ui . We ne ed nea one ha was d opping g ea
quan i ies o ipe ui , and al hough we caugh a numbe o
small s enode ma ines o a ious species, he ee was igno ed
by A. jamaicensis and he o he la ge ui ea e s.
Spondias mombin Linnaeus: An impo an ood o ba s
(Ga dne , 1977), S. mombin, ipens in Augus and Sep embe
when ig p oduc ion is low. I is ound in sca e ed pa ches in
he young o es on he lowe -lying a eas o he island.
Appa en ly A. jamaicensis inges s he pulp o his ui wi hou
ying o ex ac he juice and, hus, does no make pelle s.
Al hough S. mombin ui seldom was ca ied in o he ne s,
hose ne s se nea es o ees wi h ipe ui in a iably caugh
he mos A. jamaicensis and C. pe spicilla a. A la ge
concen a ion o ba s, mos ly A. jamaicensis, was ound in a
pa ch o S. mombin be ween 12 and 17 Sep embe .
Spondias adlko e i Donnell Smi h: Widesp ead on BCI,
bu nei he o ming pa ches no p oducing such g ea quan i ies
o ui as S. mombin and Q. as e olepis, S. adlko e i is an
impo an sou ce o ood o ba s in he season o ig sca ci y a
he heigh o he ains. F ui o his ee is ipe om Sep embe
o No embe and is much sough by A. jamaicensis, A.
li u a us, A. phaeo is, A. wa soni, and V. ca accioli. In 1979,
ipe ui began o all on 19 Sep embe . Many ui s we e
ca ied in o he ne s on 26 Sep embe , and by 30 Sep embe
mos A. jamaicensis and A. li u a us ha we cap u ed we e
ea ing i . A la ge concen a ion o ba s was associa ed wi h a S.
adlko e i wi h ipe ui on 7 Oc obe . The las ime we no ed
eces con aining ui pulp o his species was on 7 No embe .
Seeds o S. adlko e i wi h much o all o he pulp sc aped o
by ba s we e common unde eeding oos s h oughou
Oc obe .
Qua a ibea as e olepis Pi ie : The beau i ul Q. as e ole-
pis g ows in pa ches, which equen ly con ain many indi idu-
als,
mos ly in he old o es on he ele a ed in e io o he
island. We ound i a lowe ele a ions a Snyde -Molino 2,
Shannon 1, mou h o Ba bou C eek, S andley 16, and Wheele
26.5.
When i s ui ipens in Augus , Sep embe , and Oc obe
i is he mos abundan ee on he Pla eau wi h ui ea en by A.
jamaicensis. Indi idual ees d op la ge quan i ies o ipe ui
o e a pe iod o wo o h ee weeks o mo e. We ound
concen a ions o ba s a Q. as e olepis wi h ipe ui on h ee
occasions in Sep embe , and A. jamaicensis and C. pe spicil-
la a equen ly ca ied he ui in o ou ne s. The ib ous ui
shell emains o weeks on he g ound unde eeding oos s,
pe sis ing long a e he ui ing season has passed. Q.
as e olepis and S. mombin, oge he wi h he much less
abundan F. igona a, a e he ui s o choice o A.
jamaicensis in Sep embe , and possibly in Augus as well,
when ui s o F. insipida and F. yoponensis a e sca ce o
absen . I is o in e es ha
Q.
as e olepis and S. mombin, which
ui simul aneously and a e isi ed by he same ba s, ha e
complemen a y, nea ly nono e lapping dis ibu ions on BCI.
Bo h occu in pa ches and p oduce much mo e ui han he
e eb a e ugi o es can consume. Nea ui ing Qua a ibea
as olepis, A. jamaicensis usually was caugh in small g oups
a he han in such la ge concen a ions as o en appea ed a
a o ed ig ees a o he seasons. Some Q. as e olepis, e en
when d opping quan i ies o ui , almos seemed o be igno ed
by ba s on some nigh s when we ne ed nea hem.
Dip e yxpanamensis (Pi ie ) Reco d and Mell: Du ing he
d y season when ig p oduc i i y is low, he ui o D.
panamensis is an impo an ood o A. jamaicensis. On BCI
Dip e yx is a widesp ead and abundan ee,'some imes o ming
la ge pa ches (as a Ze ek 22 and eas o S andley 19). I s ui
is ipe om Decembe o Ma ch and occasionally a o he
seasons o he yea ( ui was ca ied in o a ne in mid-
No embe ). We ound la ge concen a ions o ba s, p incipally
A.
jamaicensis, a ipe D. panamensis in Janua y 1979 and
Feb ua y 1978, when nume ous ui s we e ca ied in o he
ne s.
B osimum alicas um Swa z: Robin Fos e (pe s. comm.)
ega ds ba s as he p incipal dispe se s o B. alicas um seeds,
and C oa (1978) belie ed ha B. alicas um is second only o
igs in impo ance as a ood o o es animals. We ound a
la ge concen a ion o ba s, including almos 100 A. jamaicen-
sis,
a a B osimum bea ing ipe ui in May. Howe e , a la ge
ee aining ipe ui in Oc obe , nea he end o he ui ing
season, was a ac i e o kinkajous (Po os la us) and pecca ies
(Tayassu ajacu), bu appa en ly was o li le o no in e es o
ba s.
Anaca dium excelsum Be e o and Balbis: Abundan and
widesp ead, and ui ing om Ma ch o May, A. excelsum is an
impo an ood o C. pe spicilla a. We o en caugh his ba ,
and A. phaeo is on one occasion, ca ying he ui o A.
excelsum. Excep o one ui ca ied in o a ne in Ma ch, we
ha e no e idence ha A. jamaicensis ea s he ui o his
species.
Poulsenia a ma a Miguel: The spiny ui s o
P.
a ma a
occasionally we e ca ied in o he ne s by A. phaeo is (Janua y,
Sep embe , and Oc obe ), A. wa soni (Oc obe ), and once by a
A.
jamaicensis (No embe ).
Calophyllum longi olium Willdenow: In Sep embe and
Oc obe we o en ound he la ge seeds o
C.
longi olium unde
dining oos s o A. jamaicensis, and occasionally we ound
emains o pulp and skin in he eces o his ba . The pelle s
om he ui o C. longi olium d opped by A. jamaicensis and
A.
li u a us seem ex emely esis an o decay and pe sis on he
g ound o se e al weeks. Al hough he la ge ound seeds a e
conspicuous, he pelle s a e much smalle and da ke han hose
NUMBER 511145
d opped by A. jamaicensis when eeding on igs, and
consequen ly we o en o e looked hem. F ui s o C. longi o-
lium ha we e ca ied in o ne s by ba s we e o e ed o A.
jamaicensis, A. li u a us, and
V.
ca accioli empo a ily held in
cap i i y on BCI. Bo h species o A ibeus a e he ui , bu V.
ca accioli did no , al hough i eadily a e igs when hose we e
o e ed.
Cec opia spp.: Occasionally we ound seeds o uniden i-
ied species o Cec opia in eces and unde eeding oos s o A.
jamaicensis.
Gue a da oliacia S andley: Pa ly ea en ui s o G.
oliacia we e ound benea h eeding oos s, p obably o A.
jamaicensis, in Sep embe 1979 and Oc obe 1978.
Hu a c epi ans Linnaeus: Nume ous leshy s amina e
lowe s ems o Hu a c epi ans we e ca ied in o ne s by
Phyllos omus discolo , A. jamaicensis, and A. li u a us in
No embe .
Uniden i ied Pollen: In No embe and Decembe , nume -
ous ba s o he species Phyllos omus has a us, Glossophaga
commissa isi, U ode ma biloba um, U. magni os um, A.
jamaicensis, and A. li u a us we e s ained yellow wi h
uniden i ied pollen (p obably mainly om lowe s o he
Bombacaceae).
T ee Selec ion
We know ha
A.
jamaicensis will ea a a ie y o ui s, and
some imes o he oods, bu mos o i s meals a e igs. On BCI
i p e e s F. insipida o e all o he igs, and i mus be a a e
nigh on he island when he e a e no ipe ui s o his species
a ailable. We ha e no idea why A. jamaicensis occasionally
chooses he ui s o B osimum, Calophyllum, Dip e yx,
Qua a ibea, o Spondias o e Ficus, bu we ha e lea ned much
abou why and when hey choose a pa icula ig ee.
The no mal massi e c op o igs o each ee goes h ough
se e al s ages o ha es as i ipens. Fi s come he howle
monkeys (Aloua a pallia a), which begin o ea he ui long
be o e i is ipe, while he pulp is ha d and he skin is s ill ull
o la ex. The monkeys a e ollowed by he local ba s om
nea by oos s. They also begin o ha es a c op be o e i is
ully ipe, p obably selec ing sca e ed ipe ui , and some-
imes picking un ipe ui . We ha e caugh A. jamaicensis
ca ying un ipe igs, bu we do no know whe he hey ac ually
ea such ui , and i hey do, whe he hey ea all o only pa
o i . Pe haps hey a e sampling c ops o de e mine he s age o
ipeness. Pelle s d opped by local ba s may be conspicuous
du ing se e al nigh s be o e enough o he ui c op is ipe o
a ac la ge g oups o ba s.
When he big c op inally is ully ipened, ba s a he ee
may numbe in he hund eds o e en housands. They come in
su ges, and some may s ay in he icini y all nigh (see Sec ion
7,
Mo emen s and Sec ion 9, Fo aging Beha io ). G oups will
e u n o a ee ha has a big c op o h ee o ou nigh s. In he
inal s age o ha es , monkeys, o he diu nal ugi o es, and
local ba s s ill e u n o he ee a e i s c op has been oo
deple ed o con inue o a ac la ge locks o ba s. Al oge he ,
local ba s may ha es igs om a ee o e a pe iod o eigh o
nine nigh s.
Some ees p oduce la ge c ops ha ipen a ew igs a a ime
du ing a p olonged pe iod (up o 10 o 15 days) a he han
ipening ab up ly as igs no mally do. These ees a e a ac i e
o local ba s and nonchi op e an ugi o es, bu no o g oups o
ba s,
excep o b ie passes.
The emnan s o ig c ops abo ed because o lack o
pollina ion by ig wasps, o because o in es a ion wi h la ae
o bee les and lies, as well as he c ops p oduced by small,
young ees, migh no a ac many ba s. Size o he ui c op
has an impo an bea ing on i s use by ba s. Small c ops may be
ha es ed by local ba s, bu usually hey a e no a ac i e o
g oups. Because monkeys {Aloua a pallia a) begin o ha es
a c op be o e ba s do, hey may s ip a small c op be o e i is
ipe enough o a ac ba s. Local ba s may help inish
ha es ing a c op ha is oo small o a ac a g oup. O , a g oup
may make one o wo eeding passes and hen lea e.
Radio- acking e ealed ha a ba may isi h ee o i e ees
in a nigh (Mo ison, 1978a).
The ui s o igs a e subjec o des uc i e p ocesses such as
in ec ion wi h ungi and in es a ion by insec s ha make hem
una ac i e o ba s. Du ing he ainy season we o en saw F.
insipida in ec ed wi h a ungus mani es ed in ully de eloped
ipe ui ha ha e a bea ded appea ance while s ill on he ee.
As alling ui accumula es, he g ound benea h he ee
becomes ca pe ed wi h he uzzy whi e ui . No hing ea s i , in
he ee o on he g ound, and e en ually i o s away. Fungi, as
well as in e eb a es o he li e , accele a e he b eakdown o
he pelle s o chewed ig pulp d opped o he g ound as ba s
eed. Only he mos disce ning eye will spo aces o hese
pelle s ou o i e days a e hey ha e been d opped.
The ui o bo h F. insipida and F. yoponensis can be
in es ed by he la ae o lies and bee les. I he in es a ion is
hea y, he ee abo s pa o all o i s c op a abou he six h
week (abou h ee- ou hs o he way h ough he de elopmen-
al cycle). These ha d, la ex-laden, and insec - iddled ui s a e
eage ly ea en by monkeys (bo h Aloua a pallia a and Cebus
capucinus),
ugi o ous bi ds, and e es ial ugi o es such as
pacas (Agou i paca), pecca ies (Tayassu ajacu), and api s
(Tapi us bai dii). Howe e , al hough hey may be la ge and
appea o be ipe, hey a e no swee , and hey a e igno ed by
ba s.
Un il we lea ned he na u e o his kind o ui -c op
abo ion, we some imes was ed e o by ne ing a hese ees.
We we e misled by he abundance o alling ui and he
e e ish ac i i y o many ugi o es in o belie ing ha ba s
also would lock o he ee.
Summa y
Because cap u e a e is clea ly linked o o aging beha io ,
we ga he ed da a on ecal con en s, eeding oos s, ood ca ied
146SMITHSONIAN CONTRIBUTIONS TO ZOOLOGY
in o ne s, and on he loca ion and phenology o ui ing ees.
The ba s' habi o d opping small pelle s o chewed ui pulp
unde a o i e eeding oos s equen ly dic a ed ou choice o
ne ing si es. Cong ega ions o diu nal ugi o es led us o ees
wi h ipening ui ha would be isi ed a nigh by ba s.
Ce ain species o ees a o ed as eeding oos s a e used
epea edly, especially in he icini y o ui ing ees.
F ui o he ig Ficus insipida is he a o i e ood o A.
jamaicensis as well as se e al o he ba s on BCI. F ui s o o he
species o igs, including F. yoponensis, F. dugandii, F.
igona a, and F. ob usi olia, a e some imes ea en by A.
jamaicensis, bu he uncommon F. popenoei is igno ed.
Spondias mombin, S. adlko e i, and Qua a ibea as e olepis
a e impo an ood sou ces la e in he ainy season when igs
a e sca ce. Dip e yx panamensis is used in he d y season as an
al e na e ood sou ce. The ui s o Anaca dium excelsum and
Poulsenia a ma a and he lowe s o Hu a c epi ans occasion-
ally a e ea en by A. jamaicensis.
The no mal massi e c op o a F. insipida is used by local
ba s o se e al days be o e he en i e c op is ully ipe, a which
ime hund eds o e en housands o ba s may isi he ee o
h ee o ou nigh s un il he c op is deple ed. Then local ba s
con inue o use he ee o se e al nigh s un il he e is no mo e
ui . T ees wi h small c ops and ees p oducing c ops ha
ipen asynch onously o e a pe iod o a week o mo e a e
isi ed by local ba s bu a e la gely igno ed by g oups.
11.
Die and Food Supply
Cha les
O.
Handley, J ., and Egbe G. Leigh,
J .
Bonacco so (1979) a gued ha ood supply limi s popula-
ions
o
ba s
on
Ba o Colo ado Island (BCI). La e
in he
ainy
season when ui is sca ce, he ound mo e ui ba s wi h emp y
s omachs han
he did a
o he imes
o
yea . Abou 83%
o he
ugi o es
he
ne ed
in
Oc obe
and
No embe
had
emp y
s omachs,
in
con as o
71%
o
hose ne ed in Ma ch and Ap il
when ui was
a
mo e abundan . Howe e , hese da a mus
be
in e p e ed wi h cau ion. Bonacco so kep
his
da a ee
o
bias
by ne ing
he
same locali ies each mon h, wi hou ega d
o
p esence
o
absence
o
ui nea
he
ne ing s a ions.
The
ba s
he caugh wi h emp y s omachs mus ha e been
on
hei way
o
someplace else whe e he e
was
ui They could
no
endu e
emp y s omachs
o
mo e han
a ew
hou s wi hou s a ing
o
dea h.
Mo eo e , Bonacco so (1979) obse ed ha bi h
(and
b eeding)
o
ba s
is
imed
o
coincide wi h seasons
o
ui
abundance, wi h
one
bi h peak
in
Ma ch
and
Ap il coinciding
wi h
he
ui ing peak
a he
onse
o he
ainy season
and
ano he
in
July
and
Augus coinciding wi h
he
ui ing peak
o
Augus
and
Sep embe (Figu es
4-6 and
4-7).
Few
ugi o ous
ba s gi e bi h be ween No embe and mid-Ma ch when ui
is
leas abundan .
Bonacco so (1979) also ound ha
he
die s
o
a ious
species
o
ba s ha o age
o
ui
in
he canopy di e ed
o he
ex en one would expec
i
hese animals we e ood limi ed.
He
ound ha la ge ba s
a e
la ge igs such
as
Ficus insipida
and
F. ob usi olia, whe eas smalle ba s concen a ed
on
smalle
igs such as
F.
popenoei and
F.
yoponensis.
Fo he
h ee la ge
ugi o es (A ibeus li u a us, A.jamaicensis,
and
Vampy odes
ca accioli),
he
ound ha he eg ession
o
he mass Y
o
a
ui
ca ied
by a ba on he
mass
X o i s
ca ie
was Y =
0.23X
- 3.92 g ( 2 = 0.46, n = 27).
P esumably, smalle ba s
ca ied ui in o
he
ne s
so
a ely ha
he
could
no
ex end
he
eg ession.
Reading
he
mass
o
hese ui s om Bonacco so's g aph
and calcula ing
he
mean
and
s anda d de ia ion
o he
loga i hms
o he
mass
o he
ui s hese ba s ca ied,
we
Cha les
O.
Handley,
J .,
Na ional Museum
o
Na u al His o y,
Smi hsonian Ins i u ion, Washing on,
D.C.
20560.
Egbe
G.
Leigh,
J .,
Smi hsonian T opical Resea ch Ins i u e, Uni
0948,
APO AA
34002-0948
o
Apa ado
Box
2072, Balboa, Republic
o Panama.
de i ed alues
o 2.30 ± 0.24 (n = 6) o A.
li u a us,
2.04 ±
0.33
(n = 17) o A.
jamaicensis,
and 1.54 ±
0.31
(n = 4) o
V.
ca accioli.
The
s anda d de ia ions
a e
oughly equal
o he
di e ences be ween neighbo ing means.
May and
MacA hu
(1972) showed ha
in an
idealized compe i i e communi y,
species could coexis secu ely
i
he sizes
o
oods ea en
by he
a ious species di e ed
o
his ex en . Excep ing A ibeus
phaeo is, which ea s
ew
igs,
he
a io
o he
mass
o
each
species
o
canopy ugi o e
o
ha
o i s
nex smalle
compe i o
was
oughly
he
same
as in
hese h ee species,
abou
1.4 : 1
(Bonacco so, 1979).
I is emp ing
o
assume ha he ela ion be ween
he
sizes
o
he smalle s enode ma ines and he sizes
o
ui hey
ea
is
he
one Bonacco so in e ed om
he ew
da a
on his
la ges
ugi o es. Fu he mo e,
we
could conclude om
he
elegan
heo y de eloped
by May
(1974)
and May and
MacA hu
(1972)
on
niche o e lap, ha
he
canopy ugi o es coexis
by
i ue
o he
di e ences Bonacco so obse ed
in he
sizes
o
ui hese ba s
ea
Ideally,
i
all sizes
o
igs we e a ailable
a
all imes and we e
uni o mly dis ibu ed,
he
smalle ba s would usually ake
he
smalle igs
and he
la ge ba s would usually ake
he
la ge
igs. Howe e , small- ui ed
and
la ge- ui ed species
o
igs
a e
no
uni o mly dis ibu ed ei he
in
ime
o
space,
and he
ba s
a e
adap able enough
o
ake wha
is
a ailable.
O
cou se,
in s anda dizing hei die s,
i is
also necessa y
o
ake in o
accoun
he
o aging beha io
o
hese ba s.
The
ene gy
expended
in
commu ing om dining oos
o
ui ing ee
makes
i
ene ge ically imp uden
o he
la ge ba s
o
ou inely
eed
on
small igs om each
o
which hey would ex ac
a
compa a i ely iny amoun
o
nu ien s
(see
Sec ion
2,
Physiology).
Fig P oduc ion
When
we
became conce ned abou he ela ion be ween ood
a ailabili y
and he
ba s' ene gy equi emen s
(see
Sec ion
2,
Physiology), we began
o
ga he in o ma ion on es ima es
o ig
p oduc ion
on BCI.
Each indi idual
ig
ee bea s ui
o i s
own hy hm (Mo ison, 1978d)
so
ha
a
any season some ees
a e bea ing ipe igs (Table 10-1). Fig p oduc ion peaks ea ly
in
147