Ten
Cons uc ion
and
Use
by
U ode ma
biloba um
in
Coconu
Palms
(Cocos
nuci e a)
in
Cos a
Rica
ROBERT
M.
TIMM'
AND
SUSAN
E.
LEWIS2
ABSTRACT
Ten
cons uc ion
and
use,
uni o mi y
o
en s,
and
ond
selec ion
we e
s udied
in
a
popula ion
o
U ode ma
biloba um
oos ing
in
coconu
palms
(Cocos
nuci e a)
in
Guanacas e
P o ince
o
no h-
wes e n
Cos a
Rica
du ing
July
1988.
Palm
lea le s
we e
cu
a
hei
mid ibs
in
a
line
con e ging
dis-
ally
wi h
he
ond
mid ib,
and
he
lea le s
col-
lapsed
downwa d
o
o m
a
la ge
enclosed
en .
Ten
heigh ,
numbe
o
lea le s
cu ,
and
angle
be-
ween
he
line
o
cu
lea le s
and
he
mid ib
o
he
onds
we e
measu ed
o
assess
uni o mi y
o
en
cons uc ion.
To
asce ain
i
ba s
we e
selec ing
speci ic
ees
o
onds,
we
measu ed
he
angle
o
o ien a ion
o
cu
onds,
numbe
o
onds
hang-
ing
abo e
a
en ,
and
ee
heigh .
Ba
en s
we e
ound
in
palms
wi h
a
na owe
ange
o
heigh s
han
he
o e all
ee
popula ion,
and
ees
wi h
en s
we e
alle
on
a e age
han
ees
wi hou
en s.
A
single
al e ed
ond
p o ides
excellen
p o ec ion
om
ain all.
Ba s
do
no
seem
o
p e e
onds
based
on
numbe
o
o e hanging
onds
o
angle
o
o ien a ion.
The
age
o
he
modi ied
ond
may
be
an
impo an
ac o
in
oos
si e
selec ion,
as
en s
in
younge
onds
we e
mo e
likely
o
be
occupied
han
hose
in
olde
onds.
The
numbe
o
ba s
oos ing
unde
en s
anged
om
1
o
15
adul s
and
subadul s.
The
colony
was
composed
la gely
o
adul
emales
and
wo
age
classes
o
young.
INTRODUCTION
I
has
been
known
o
o e
hal
a
cen u y
ha
some
ba s
c ea e
hei
own
oos ing
si es
by
modi ying
he
shapes
o
lea es
(Ba bou ,
1932;
Chapman,
1932).
Ba s
c ea e
hese
s uc u es
by
se e ing
he
eins
and,
in
some
cases,
he
in e connec ing
issues
o
lea es
o
a ious
species.
The
sides
o
he
lea es
hen
collapse
downwa d
along
he
mid ib
o
o m
a
da k,
secluded
oos ing
si e
o
he
ba s.
Because
some
s yles
o
hese
modi ied
lea es
a e
py amidal o
" en -shaped,"
all
modi ied
lea es
a e
now
called
ba
en s
and
he
ba s
ha
modi y
hem
a e
called
en -making
ba s.
Fou een
species
o
New
Wo ld
phyllos-
omid
ba s,
all
in
he
sub amily
S enode -
ma inae,
a e
known
o
cons uc
en s
(Timm,
1987).
Addi ionally,
wo
Old
Wo ld
species
o
lying
oxes-
Cynop e us
b achyo is
and
C.
sphinx
( amily
P e opodidae)
-ha e
been
e-
po ed
o
modi y
palms
o
p oduce
diu nal
oos ing
s uc u es
(Phillips,
1924;
Goodwin,
1979).
Re iews
o
en
cons uc ion
and
use
by
ba s
we e
p o ided
by
Kunz
(1982)
and
Timm
(1987).
In
Cos a
Rica
and
elsewhe e
in
he
Neo-
opics
we
ha e
ound
ha
(1)
en
ba s
a e
o en
highly
localized
in
occu ence;
(2)
en
ba s
seem
o
be
mos
concen a ed
in
a eas
ha
ha e
an
abundan
supply
o
he
p e e ed
plan
species
used
in
en
cons uc ion;
(3)
qui e
o en
en s
a e
concen a ed
in
localized
a eas
e en
hough
he
p e e ed
plan
species
is
mo e
widely
dis ibu ed;
(4)
he
loca ion
o en s
wi hin
plan s
a ies,
as
does
he
shape
I
Cu a o
o
Mammals
and
Associa e
P o esso ,
Depa men
o
Sys ema ics
and
Ecology
and
Museum
o
Na u al
His o y,
Uni e si y
o
Kansas,
Law ence,
Kansas
66045.
2
Ph.D.
Candida e,
Depa men
o
Ecology,
E olu ion,
and
Beha io ,
Uni e si y
o
Minneso a,
Minneapolis,
Minneso a
55455.
251
BULLETIN
AMERICAN
MUSEUM
OF
NATURAL
HISTORY
o
en s
cons uc ed
by
a
gi en
species
o
ba ;
and
(5)
he e
a e
o en
many
mo e
en s
p es-
en
in
an
a ea
han
a e
occupied
by
ba s.
These
ac o s
sugges
ha
he
ba s
a e
selec ing
spe-
ci ic
lea es
o
en s
and
using
only
ce ain
en s
o
hose
a ailable
on
a
daily
basis.
Roos
si e
selec ion
in
en
ba s
has
been
in es i-
ga ed
p e iously
o
only
a
ew
species
o
he
smalle
en -make s:
A ibeus
phaeo is
(Timm,
1987),
A ibeus
wa soni
(Choe
and
Timm,
1985),
Ec ophylla
alba
(Timm
and
Mo ime ,
1976;
B ooke,
1990),
and
Vam-
py essa
nymphaea
(B ooke,
1987).
These
s udies
concluded
ha
ba s
selec
speci ic
spe-
cies
o
plan s
o
en
cons uc ion
and
ha
hey
o en
selec
speci ic
ages
o
lea es.
Be-
cause
o
he
la ge
size
o
many
al e ed
lea es,
especially
hose
o
pinna ely
compound
plan s,
a
conside able
amoun
o
ene gy
p obably
is
expended
by
he
ba
o
ba s
in
c ea ing
hese
oos
si es.
The
ene ge ic
cos s
associa ed
wi h
he
elabo a e
na u e
o
many
s yles
o
ba
en s,
especially
hose
cons uc -
ed
by
ba s
o
he
genus
U ode ma,
sugges
ha
s ong
selec ion
p essu es
a e
in ol ed.
In e es ingly,
no
s udies
ha e
add essed
oos
si e
selec ion
and
en
use
by
Pe e s's
en -making
ba
(U ode ma
biloba um),
he
i s
species
o
Neo opical
ba
disco e ed
o
modi y
lea es
and
one
o
he
la ges
and
mos
widely
dis ibu ed
o
he
Neo opical
en -
making
ba s.
Ou
disco e y
o
a
sizable
pop-
ula ion
o
U.
biloba um
in
he
Paci ic
low-
lands
o
Cos a
Rica
oos ing
in
al e ed
onds
o
coconu
palms
p o ided
an
ideal
oppo -
uni y
o
explo e
se e al
aspec s
o
en
con-
s uc ion
and
use
by
hese
ba s.
The
goals
o
his
s udy
we e
o
(1)
de e mine
whe he
U.
biloba um
chooses
speci ic
ees
o
onds
o
en
cons uc ion,
(2)
assess
whe he
hey
cons uc
en s
o
a
uni o m
shape
o
heigh ,
(3)
de e mine
i
en s
a e
e ec i e
p o ec ion
om
ain,
and
(4)
e alua e
pa e ns
o
en
use
(including
g oup
size,
g oup
composi ion,
and
mo emen
be ween
en s).
ACKNOWLEDGMENTS
We
a e
g a e ul
o
S .
Guille mo
Canessa
Mo a
o
El
Depa amen o
de
Vida
Sil es e
and
S .
Fe nando
Co es
o
Se icio
de
Pa -
ques
Nacionales
o
Cos a
Rica
o
making
ou
wo k
a
Palo
Ve de
possible.
We
hank
he
O ganiza ion
o
T opical
S udies
o
p o-
iding
logis ic
and
inancial
suppo ,
and
William
S.
Al e son
and
Be e
A.
Loiselle
o
assis ance
wi h
logis ics.
Kimbe ly
J.
Babbi ,
Nina
R.
Ingle,
John
M.
Kasme ,
Ke in
Ken-
nedy,
John
E.
Mi le ,
Sean
O'Donnell,
and
B uce
E.
Young
p o ided
supe b
assis ance
in
he
ield.
Ba ba a
L.
Clauson,
Thomas
A.
G i i hs,
Don
E.
Wilson,
and
an
anonymous
e iewe
made
cons uc i e
commen s
on
ea lie
d a s
o
he
manusc ip .
Ka h yn
E.
S one
gene ously
p o ided
us
wi h
he
in-
o ma ion
she
ob ained
om
his
same
col-
ony
o
ba s
in
1990.
Lewis
was
suppo ed
by
inancial
assis ance
om
he
College
o
Bi-
ological
Sciences
and
he
Depa men
o
Ecology,
E olu ion,
and
Beha io ,
Uni e -
si y
o
Minneso a.
METHODS
F om
16
h ough
21
July
1988, indi iduals
and
g oups
o
U ode ma
biloba um
con ex-
um
and
hei
en s
in
coconu
palms
(Cocos
nuci e a)
we e
obse ed.
The
plan ed
coco-
nu
g o e
con ained
56
ees
ha
a ied
in
heigh
and
age,
and
was
loca ed
app oxi-
ma ely
0.5
km
eas
o
he
adminis a i e
buildings
a
Re ugio
Nacional
de
Fauna
Sil-
es e
D .
Ra ael
Lucas
Rod iguez
Caballe o
(commonly
known
as
Palo
Ve de)
in
he
Pa-
ci ic
lowlands
o
no hwes e n
Cos a
Rica.
Palo
Ve de
is
a
wildli e
e uge
loca ed
ap-
p oxima ely
2
km
sou h
and
12
km
eas
o
Bolson
in
Guanacas e
P o ince
(10°30'N,
85'20'W;
ele .
10
m).
The
a ea
lies
wi hin
he
T opical
D y
Fo es
Li e
Zone
(Hold-
idge,
1967)
and
is
domina ed
by
lowland
deciduous
and
i e ine
swamp
o es s
and
a
la ge
seasonal
ma sh.
Rain all
is
ex emely
seasonal,
wi h
mos
o
he
mean
annual
p e-
cipi a ion
o
1700+
mm
alling
be ween
Ap il
and
Oc obe .
De ails
o
he
ege a ion,
hab-
i a
ypes,
and
clima e
o
Palo
Ve de
ha e
been
desc ibed
by
Slud
(1980)
and
Ha sho n
(1983).
All
ba
en s
we e
su eyed
daily,
and
he
numbe
o
oos ing
ba s
p esen
in
each
en
was
coun ed.
Nu sing,
non olan
ju enile
o sp ing
( o ea m
=
32.5
mm
[N
=
1])
we e
clea ly
isible
clinging
o
23
adul
emales
( o ea m
x
=
42.5
mm
[N
=
2]),
enabling
us
o
easily
dis inguish
hem
om
o he
ba s.
252
NO.
206
TIMM,
LEWIS:
URODERMA
TENT
CONSTRUCTION
Fig.
1.
Do sal
iew
o
a
coconu
ond
showing
he
lea le s
cu
by
U ode ma
biloba um
o
o m
a
en .
Twen y- h ee
o
he
41
adul
and
subadul
ba s
p esen
we e
cap u ed
in
mis
ne s,
and
21
o
hese
we e
ma ked
o
u u e
iden i i-
ca ion.
Volan
ba s
we e
judged
o
be
adul s
i
he
phalangeal
epiphyses
we e
used,
and
subadul s
i
he
epiphyses
we e
no
ye
used.
To
es
whe he
U ode ma
selec s
speci ic
ees
o
onds
o
en
cons uc ion,
he
ol-
lowing
measu emen s
we e
aken.
We
mea-
su ed
he
dis ance
om
he
g ound
o
he
poin
whe e
he
lowes
g een
ond
a ached
o
he
ee
( o
he
nea es
0.1
m).
This
p o-
ided
an
es ima e
o
ee
heigh ,
because
co-
conu
ees
shed
he
lowes
lea es
as
hey
253
1991
BULLETIN
AMERICAN
MUSEUM
OF
NATURAL
HISTORY
g ow
alle
and
lea es
a e
concen a ed
a
he
e y
op
o
he
ee.
To
assess
whe he
ba s
we e
selec ing
onds
wi h
espec
o
he
di-
ec ion
he
onds
hung,
we
measu ed
he
compass
di ec ion
o
each
cu
ond
o
he
nea es
deg ee.
The
numbe
o
onds
di ec ly
abo e
a
en
was
coun ed
o
de e mine
i
ba s
we e
cons uc ing
en s
in
onds
ha
had
onds
abo e,
possibly
p o iding
addi ional
shade
o
p o ec ion
om
ain.
Two
aspec s
o
he
coconu
ees
and
al-
e ed
onds
we e
measu ed
o
assess
he
de-
g ee
o
uni o mi y
among
en s
in
his
a ea.
Fi s ,
lea le s
hanging
pe pendicula ly
o
he
plane
o
he
ond
up
o
he
poin
whe e
he
line
o
cu s
con e ged
wi h
he
mid ib
o
he
ond
we e
coun ed
as
cu
lea le s
ha
con-
ibu e
o
he
en ;
any
cu
lea le s
dis al
o
he
poin
o
con e gence
wi h
he
mid ib
we e
no
coun ed
( ig.
1).
Second,
he
angle
be-
ween
he
mid ib
and
one
o
he
wo
ows
o
cu
lea le s
was
es ima ed
(wi h
a
p o ac o )
o
he
nea es
deg ee.
We
also
assessed
p o ec ion
om
ain all
unde
en s
by
measu ing
wa e
collec ed
o e
a
4-day
pe iod
in
a
250-ml
beake
placed
di ec ly
unde
he
cen e
o
a
ypical
en ,
in
he
posi ion
ba s
no mally
hang.
Du ing
he
same
pe iod,
wo
addi ional
beake s
we e
placed
in
he
open
o
measu e
o al
ain all.
To
e alua e
pa e ns
o
en
use,
we
su -
eyed
g oup
size
and
composi ion
unde
in-
di idual
en s
h oughou
he
s udy.
Two
ac-
o s
-age
o
onds
and
en
heigh
-we e
examined
o
asce ain
i
ba s
occupied
spe-
ci ic
en s
o
he
o al
en s
a ailable.
The
age
o
he
cu
ond
(young
o
old)
was
es ima ed
by
he
ond's
posi ion
on
he
ee
in
ela ion
o
o he
onds.
Ten
heigh
was
measu ed
( o
he
nea es
0.1
m)
as
he
dis ance
om
he
g ound
o
he
poin
a
which
he
line
o
cu
lea le s
con e ged
wi h
he
ond
mid ib.
A
e e ence
specimen
o
U ode ma
bilo-
ba um
om
his
popula ion
was
deposi ed
in
he
Museo
Nacional
de
Cos a
Rica
in
San
Jose;
he
museum's
mammals
ecen ly
we e
ans e ed
o
INBio
( he
new
Na ional
Bio-
di e si y
Ins i u e,
San o
Domingo,
Cos a
Rica).
RESULTS
A
Palo
Ve de,
U ode ma
biloba um
se e s
he
mid ib
o
lea le s
on
he
la ge,
pinna ely
compound-lea ed
coconu
palm,
Cocos
nu-
ci e a,
o
o m
da k,
secluded
diu nal
oos s
and
ma e ni y
si es.
The
cu
lea le s
old
downwa d,
pe pendicula
o
he
g ound,
c e-
a ing
la ge,
angula
en s
( ig.
2).
Ba s
cu
he
mid ib
o
lea le s
bu
do
no
appea
o
se e
su ounding
issue,
no
do
hey
se e
he
mid ib
o
he
ond.
The
lea le s
closes
o
he
ee
unk
a e
se e ed
a
he
g ea es
dis-
ance
om
he
mid ib
o
he
ond.
This
dis-
ance
dec eases
as
he
cu s
p oceed
dis ally
o
a
poin
a
which
hey
con e ge
wi h
he
ond
mid ib
( ig.
1).
A
a iable
numbe
o
lea le s
beyond
he
poin
o
con e gence
may
also
be
cu .
Ten
shape
is
a
combina ion
o
he
numbe
o
lea le s
cu
and
he
angle
a
which
he
line
o
cu s
is
made.
O
he
56
ees
in
he
coconu
g o e,
23
con ained
en s
cons uc ed
by
U.
biloba um.
Fo y- ou
en s
we e
loca ed
on
ou
i s
su -
ey.
Th ee
addi ional
en s
we e
cons uc ed
o e
he
nex
i e
nigh s.
O
hose
ees
ha
had
en s,
he
mean
numbe
o
en s
pe
ee
was
2.0
( ange
1-5,
SD
=
1.2).
The
oad
and
ails
wi hin
a
1.5-km
adius
o
he
s udy
si e
we e
sea ched
o
addi ional
en s
o
la ge-
lea ed
ees
app op ia e
o
en s,
bu
none
we e
ound.
O
all
he
ees
in
he
g o e,
he
ange
o
heigh s
o
ees
wi h
en s
encompassed
he
alle
ees,
bu
excluded
he
e y
alles .
The
a e age
heigh
o
he
lowes
g een
ond
(ou
es ima e
o
ee
heigh )
o
all
ees
was
1.8
m
( ange
0.5-5.4
m,
SD
=
0.96).
Ba
en s
we e
ound
in
ees
ha
a e aged
2.3
m
in
heigh
o
he
lowes
ond
( ange
1.6-3.0
m,
SD
=
0.42).
The
a e age
heigh
o
ees
wi h
en s
(2.3
m,
N
=
23)
was
highe
han
he
a e age
o
ees
wi hou
en s
(1.5
m,
N
=
33,
ange
0.5-5.4
m,
SD
=
1.08;
Wilcoxon
ank
sum,
P
<
0.001).
The
o ien a ion
o
he
ond
and
he
num-
be
o
onds
abo e
a
en
did
no
a ec
which
onds
he
ba s
selec ed
o en
cons uc ion.
The e
was
no
pa e n
o
he
o ien a ion
o
cu
onds;
he
compass
di ec ion
anged
om
0
o
3520
(x
=
1910,
SD
=
1010).
The
numbe
o
onds
abo e
a
gi en
en
anged
om
0
o
3
(Fx
=
0.7,
SD
=
0.85).
Al hough
he e
was
a
de ini e
in e ed
V-shaped
pa e n
o
he
gene al
o m
o
en s
cons uc ed,
ac ual
en
shape
was
a iable.
O
26
en s
we
we e
able
o
measu e
and
coun ,
17
had
lea le s
cu
on
bo h
sides
and
254
NO.
206
TIMM,
LEWIS:
URODERMA
TENT
CONSTRUCTION
Fig.
2.
Ten
o
U ode ma
biloba um
in
he
pinna ely
lea ed
coconu
palm
Cocos
nuci e a.
9
had
lea le s
cu
on
only
one
side.
The
mean
numbe
o
lea le s
cu
pe
side
was
12.8
( ange
0-36,
SD
=
7.9).
The
angle
o med
by
he
ond
mid ib
and
he
line
o
cu
lea le s
a -
e aged
32.30
( ange
20-46',
SD
=
7.70).
Two
beake s
placed
in
he
open
each
col-
lec ed
app oxima ely
90
ml
o
ain all
du ing
a
4-day
pe iod.
The
beake
placed
unde
a
ba
en
du ing
he
same
pe iod
collec ed
only
2
ml
o
ain.
Al hough
44
47
en s
(3
en s
we e
con-
s uc ed
du ing
ou
s udy)
we e
a ailable
o
he
oos ing
ba s,
only
9-11
we e
occupied
on
any
gi en
day
du ing
he
s udy.
Six y-nine
pe cen
o
he
occupied
en s
we e
in
use
4
o
mo e
days.
Thi y- ou
en s
we e
ne e
occupied
du ing
he
s udy.
Occupied
en s
appea ed
o
be
spa ially
clumped
wi hin
he
coconu
g o e.
Occupied
and
unoccupied
en s
di e ed
in
ond
age
and
heigh .
Occupied
en s
ended
o
be
in
younge
onds
( onds
highe
in
he
ee)
han
did
unoccupied
en s
(X2
=
10.9,
P
<
0.05).
Six y-se en
pe cen
o
occupied
en s
1991
255
BULLETIN
AMERICAN
MUSEUM
OF
NATURAL
HISTORY
we e
ound
in
young
onds,
whe eas
84
pe -
cen
o
unoccupied
en s
we e
in
old
onds.
Acco dingly,
en
heigh
di e ed
signi ican ly
be ween
occupied
(x
=
4.6
m,
SD
=
0.8)
and
unoccupied
en s
(x
=
3.8
m,
SD
=
0.7
m;
Wilcoxon
ank
sum
=
2.3,
P
<
0.05).
Fou een
ees
had
wo
o
mo e
en s,
bu
in
only
one
ins ance
was
mo e
han
one
en
occupied
in
a
single
ee.
Tha
ee
had
i e
en s,
one
o
which
was
occupied
on
5
o
he
6
days
by
a
la ge
clus e
o
ba s.
On
one
o
hose
days,
an
addi ional
en
was
occupied
by
a
single
ba .
The
o he
h ee
en s
in
his
ee
we e
ne e
occupied.
On
h ee
occasions
we
obse ed
U.
bilo-
ba um
oos ing
unde
coconu
onds
ha
clea ly
had
no
been
al e ed
o
o m
a
en .
The
lea le s
o
each
o
hese
onds
d ooped
na u ally
pe pendicula
o
he
g ound,
simi-
la
o
he
pa e n
seen
in
cu
lea le s.
Two
o
hese
onds
we e
occupied
by
single
ba s
o
1
day
only.
The
hi d
was
used
by
h ee,
wo,
and
ou
ba s
on
consecu i e
days.
O
he
13
di e en
en s
used
by
ba s
(and
3
unal e ed
onds),
8
we e
occupied
by
single
ba s
and
8
by
g oups
o
wo
o
mo e
ba s.
The
a e age
g oup
size
was
5.2
( ange
1-15,
SD
=
3.3).
Twen y- h ee
o
he
34
g oups
(68%)
we e
composed
p ima ily
o
emales
wi h
nu sing
o sp ing.
Fo y-one
adul
o
olan
subadul
U.
bi-
loba um
we e
obse ed
in
he
coconu
g o e
on
he
i s
day,
and
o
hese
23
we e
adul
emales
ha
had
nu i ionally
dependen ,
ju-
enile
o sp ing.
We
cap u ed
16
adul
e-
males,
1
adul
male,
5
subadul
males,
and
1
subadul
emale.
Fou een
o
he
adul
e-
males
we e
lac a ing,
one
was
isibly
p eg-
nan ,
and
one
was
pos lac a ing.
Six
emales
we e
cap u ed
while
ca ying
hei
o sp ing
wi hin
he
i s
30
minu es
a e
sunse .
Fe-
males
ha
we e
lac a ing
bu
no
ca ying
hei
o sp ing
we e
cap u ed
la e
in
he
e ening.
The
single
adul
male
obse ed
had
ully
sc o al
es es.
No
o he
species
o
ba s
we e
ound
oos ing
unde
he
coconu
onds.
O e
he
6-day
pe iod
when
ba s
we e
ma ked,
he
lac a ing
emales
wi h
nu sing
young
oos ed
in
six
o
he
en s
( able
1).
No
all
o
hese
six
en s
we e
con inuously
occupied
o e
he
6-day
pe iod.
O
ou
28
obse a ions
o
en s
occupied
by
lac a ing
TABLE
1
Dis ibu ion
and
Numbe s
o
U ode ma
biloba um
Roos ing
in
13
Ten s
and
3
Unal e ed
F onds
(in-
dica ed
by
*)
in
Coconu
Palms
(Cocos
nuci e a)
a
Palo
Ve de
in
1988a
Ten
16
17
18
19
20
21
no.
July
July
July
July July
July
1
15/6
13/10
9/6
3/3
0
1/1
2
6/5
4/4
8/8
7/6
10/7
9/6
3
6/5
6/5 6/5
5/4 5/4
5/4
4
7/7
4/4
0
3/3
1/1
4
5
1
2/2
2/1
2/1
2/1
2/1
6
0
2
3
4
3
4
7
1
2
1
1
1
1
8
1
1
1
1
1
1
9
1
1
1
1
0
0
10
1
1
0
0
1
1
11
1
0
0
0
0
0
12
1
0
0
0
0
0
13
0
0
0
0
1
1
14*
0
0
1
3
3/1
1/1
15*
0
2
0
0
0
0
16*
0
0
0
0
0
1
a
The
nume a o
(o
single
numbe
in
a
column)
is
he
numbe
o
adul s
and/o
olan
subadul s;
he
denom-
ina o
is
he
numbe
o
ju eniles
(non olan )
obse ed.
0
=
no
ba s
p esen
in
en
ha
day.
emales,
on
10
occasions
he
g oups
included
only
he
mo he /o sp ing
pai (s)
and
on
18
occasions
included
mo he /o sp ing
pai (s)
and
1-9
(x
=
1.9)
o he
nonlac a ing
ba s.
On
h ee
occasions
we
obse ed
single,
lac a ing
emales
oos ing
alone
wi h
hei
o sp ing.
Each
o
hese
emales
oos ed
in
a
di e en
en ,
and
each
mo he /o sp ing
pai
was
ound
alone
only
o
1
day.
Th ee
o
he
ma ked
subadul s
oos ed
singly,
one
oos ed
wi h
an
adul
emale
and
he
o sp ing,
one
oos ed
wi h
one
o
h ee
o he
ba s,
and
one
oos ed
in
a
la ge
clus e
o
ba s
ha
included
lac a ing
adul
emales
and
hei
nu sing
o -
sp ing.
DISCUSSION
The
species
o
ba s
ha
use
modi ied
lea es
as
diu nal
oos
si es
ace
bo h
ad an ages
and
disad an ages
due
o
his
oos ing
s a -
egy.
Po en ial
ad an ages
o
oos ing
in
en s
include
he
abili y
o
change
oos
si es
as
ood
a ailabili y
o
wea he
condi ions
change
256
NO.
206
TIMM,
LEWIS:
URODERMA
TENT
CONSTRUCTION
o
as
ec opa asi e
in es a ions
inc ease,
as
well
as
p o ec ion
om
p eda o s
ha
specialize
on
mo e
adi ional
ba
oos s
such
as
ca es
o
hollow
ees.
Disad an ages
include
he
ene ge ic
cos s
o
modi ying
he
lea ;
he
epheme ali y
o
he
lea ,
causing
he
need
o
cons uc ion
o
new
oos s
e e y
ew
mon hs;
ulne abili y
o
ba s
oos ing
unde
lea es
o
ain,
hea ,
and
wind;
and
ulne abili y
o
ce -
ain
a bo eal
p eda o s.
Depending
on
he
habi a ,
sui able
oos
si es
may
o
may
no
be
eadily
a ailable.
These
and
o he
cos s
and
bene i s
a e
likely
o
a ec
oos
si e
se-
lec ion
by
en -making
ba s.
A
Palo
Ve de,
U ode ma
biloba um
con-
s uc s
en s
in
coconu
palms
by
pa ially
cu ing
lea le s
along
a
line
ape ing
dis ally
owa d
he
mid ib
( ig.
1).
The
pa e n
o
lea
modi ica ion
is
simila
o
ha
in
en s
cu
by
U.
biloba um
in
he
palm
Scheelea
os a a,
as
well
as
in
a
a ie y
o
o he
la ge-lea ed
plan s
(Timm,
1987).
Bo h
he
angula
cu
and
he
selec ion
o
la ge-lea ed
plan s
seem
cha ac e is ic
o
his
species.
Al hough
he
in e ed
V-shaped
en
s yle
is
cha ac e is ic
and
easily
ecognizable,
we
obse ed
consid-
e able
a iabili y
among
indi idual
en s.
Some
en s
we e
cu
only
on
one
side,
a ying
numbe s
o
lea le s
we e
cu ,
and
he
angle
a
which
he
line
o
cu
lea le s
con e ged
o-
wa d
he
mid ib
a ied.
The
mos
impo an
ac o
we
s udied
e-
ga ding
ond
selec ion
o
en
cons uc ion
was
he
heigh
o
he
ee.
Ba s
cons uc
en s
in
ees
in
only
a
na ow
ange
o
he
o al
a ailable
ee
heigh s.
T ees
alle
han
a -
e age,
bu
no
he
alles ,
a e
used.
Talle
ees
may
p o ide
inc eased
p o ec ion
om
climbing
p eda o s,
bu
e y
all
ees
may
be
mo e
exposed
o
high- eloci y
wind.
Ba s
did
no
selec
onds
on
he
basis
o
hei
angle
o
o ien a ion
o
on
whe he
o
no
he
ond
was
o e hung
by
one
o
mo e
o he
onds.
A
i s
glance,
pinna ely
compound
palm
lea es,
wi h
hei
gaps
be ween
lea le s,
migh
no
seem
o
p o ide
adequa e
p o ec ion
om
he
o en ial
ain
showe s
o
he
lowland
opics.
Howe e ,
he
esul s
o
ou
simple
es ,
measu ing
ain all
unde
a
en
e sus
in
he
open,
s ongly
sugges
ha
en s
p o-
ide
oos ing
ba s
wi h
excellen
p o ec ion
om
ain.
Du ing
ou
s udy,
90
ml
o
ain
ell
in
his
a ea,
bu
he
beake
placed
unde
a
ba
en
in
he
posi ion
whe e
ba s
oos
collec ed
only
2
ml
o
wa e .
Age
o
he
ond
may
also
be
an
impo an
ac o
in
he
selec ion
o
a
speci ic
en
as
a
oos
si e.
Ten s
in
younge
onds
a e
mo e
likely
o
be
occupied
han
hose
in
olde
onds.
I
is
likely
ha
en s
in
mo e
ma u e
onds
ep esen
olde ,
p e iously
occupied
en s
cons uc ed
when
he
lea
was
younge .
Young
onds
a e
mo e
ende
and
p obably
easie
o
modi y
han
olde
onds.
Ten s
con-
s uc ed
in
young
onds
will
be
a ailable
as
oos ing
si es
o
a
longe
pe iod
o
ime
han
hose
in
olde
onds.
Younge
onds
a e
a -
he
om
he
g ound
and
may
be
mo e
di -
icul
o
p eda o s
o
each.
These
young
onds
also
ha e
mo e
pliable
achises,
which
may
p o ide
a
mo e
"sensi i e"
ea ly
wa n-
ing
o
he
app oach
o
a
climbing
p eda o .
Because
onds
a e
p esen
on
coconu
ees
o
2.5-3
yea s
(Vande mee ,
1983),
en s
cons uc ed
in
hese
onds
a e
po en ially
a ailable
o
ba s
o
many
mon hs.
As
en s
age,
hey
a e
inc easingly
damaged
by
wind
and
o he
en i onmen al
ac o s
and
become
less
sui able
as
oos
si es.
While
his
p o-
ides
a
pa ial
explana ion
o
he
obse a-
ion
ha
he e
we e
many
mo e
en s
in
he
a ea
han
we e
in
use
a
any
one
ime,
ex a
en s
may
also
p o ide
an
al e na e
oos
si e
o
ba s
ha
ha e
been
dis u bed
in
hei
en s.
We
obse ed
single
ba s
lying
om
one
en
o
ano he
du ing
ou
census
on
h ee
occa-
sions;
hese
we e
ba s
ha
we
acciden ally
dis u bed.
Ten s
in
gene al,
as
well
as
occupied
en s,
appea ed
o
be
clumped
wi hin
he
coconu
g o e.
This
clumped
dis ibu ion
could
be
due
o
he
social
o ganiza ion
o
he
ba s,
bu
mo e
likely
is
due
o
he
clumped
dis ibu ion
o
p e e ed
ees.
Talle
ees
ended
o
be
clumped
in
his
a ea.
The
ela i e
impo ance
o
hese
ac o s
could
bes
be
s udied
in
an
a ea
whe e
ee
heigh s
a e
andomly
dis ib-
u ed
o
whe e
palms
a e
all
o
equal
heigh .
The
ange
o
U ode ma
biloba um
ex ends
h oughou
Cen al
Ame ica
and
no he n
Sou h
Ame ica,
and
he
ba s
a e
ound
in
a
a ie y
o
habi a s.
In
opical
d y
o es s
such
as
a
Palo
Ve de,
ba s
seem
o
be
limi ed
in
257
1991
BULLETIN
AMERICAN
MUSEUM
OF
NATURAL
HISTORY
a ailable
oos
si es
because
he
la ge-lea ed
plan s
hey
p e e
a e
uncommon
o
absen .
A
sea ch
o
he
ails
a
he
wildli e
e uge
p oduced
no
plan s
ha
seemed
sui able
o
en
cons uc ion
by
his
species,
o he
han
he
plan ed
coconu
palms.
This
sugges s
ha
he
ba s
may
be
locally
limi ed
by
he
numbe
o
a ailable
oos
si es.
Howe e ,
he
numbe
o
ba s
oos ing
in
he
coconu
g o e
dimin-
ished
h oughou
he
6
days
o
ou
s udy,
pe haps
because
hey
we e
sensi i e
o
ou
dis u bance,
so
al e na i e
oos
si es
may
be
a ailable
in
he
a ea.
Ba s
oos ed
alone
o
in
g oups
o
up
o
15
adul s
o
subadul s
in
he
coconu
g o e.
Fe-
males
wi h
nu sing
young
we e
mos
likely
o
be
ound
wi h
a
leas
one
o he
ba
(o he
han
hei
o sp ing),
and
mos
g oups
we e
composed
o
lac a ing
emales.
Ma e ni y
oos
o ma ion
is
a
common
phenomenon
in
many
species
o
ba s.
I
seems
likely
ha
du ing
he
pe iod
o
pa u i ion
and
lac a ion,
he
en s
in
his
a ea
a e
used
by
U ode ma
biloba um
as
ma e ni y
oos s.
The
dynamics
o
ma e ni y
oos
o ma ion
and
g oup
com-
posi ion
will
be
add essed
in
a
u u e
pape .
Ou
obse a ions
o
emales
wi h
nu i ion-
ally
dependen
ju eniles
and
olan
sub-
adul s
ha
appea ed
o
be
o
uni o m
age
suppo
he
classi ica ion
o
U.
biloba um
as
being
bimodally
polyes ous,
as
sugges ed
by
p e ious
s udies
o
his
species
(Fleming
e
al.,
1972;
Wilson,
1979;
Bake ,
1981;
Bake
and
Cla k,
1987).
Ou
obse a ions,
and
hose
o
p e ious
wo ke s
(summa ized
by
Timm,
1987),
show
ha
g oups
o
U ode ma
biloba um
oos
un-
de
lea es
ha
ha e
been
al e ed
by
he
ba s
o
o m
en s.
U ode ma
selec s
la ge
lea es
ha
may
be
in
a
a ie y
o
shapes;
howe e ,
he
en s
cons uc ed
a e
cha ac e is ically
c ea ed
wi h
a
la ge,
in e ed
V-shaped
pa -
e n
in
he
cu
lea
o
lea le s.
Conside able
ene gy
is
expended
by
he
ba s
o
c ea e
hese
oos s,
as
e idenced
by
he
la ge
size
o
he
cu
a ea
made
by
U ode ma
and
by
ou
ob-
se a ions
ha
a
pe iod
o
se e al
nigh s
is
equi ed
o
he
c ea ion
o
a
new
en .
Lac-
a ing
emales
a
Palo
Ve de
usually
oos ed
in
g oups
con aining
mo e
han
one
ba .
Vo-
lan ,
non ep oduc i e
subadul s
may
oos
singly
o
wi h
g oups
o
adul
emales.
Ten
oos s
p o ide
he
ba s
wi h
excellen
p o-
ec ion
om
ain.
Al hough
en s
p obably
equi e
conside able
e o
o
cons uc ,
in
co-
conu
palms
hey
emain
a ailable
o
he ba s
o
a
pe iod
o
a
leas
se e al
mon hs,
p o-
iding
he
ba s
wi h
p o ec ion
om
he
el-
emen s
and
undoub edly
om
p eda o s
as
well.
ADDENDUM
Lewis
e u ned
o
Palo
Ve de
du ing
June
and
July
o
1989
and
su eyed
he
numbe s
o
U ode ma
biloba um
oos ing
unde
he
onds
o
his
same
g o e
o
coconu
palms
as
pa
o
a
s udy
on
he
popula ion
dynamics
o
hese
ba s.
She
ound
54
en s
in
26
o
he
54
coconu
ees
in
he
a ea.
In
ea ly
June,
om
6
o
10
U.
biloba um
we e
using
he
palm
en s.
As
he
s udy
p og essed,
he
g ea es
numbe
o
ba s
obse ed
oos ing
in
he
palm
g o e
on
one
day
was
35.
Bo h
male
and
emale
ba s
we e
obse ed
oos ing
un-
de
en s.
Du ing
ea ly
Feb ua y
1990,
Ka h yn
E.
S one
isi ed
Palo
Ve de
and
su eyed
he
numbe s
o
U.
biloba um
o
us
a
his
same
g o e.
O
he
42
alles
ees
in
he
coconu
g o e,
21
con ained
one
o
mo e
en s.
The
ba
en s
we e
checked
o
occupancy
du ing
ou
consecu i e
days.
On
2
Feb ua y,
h ee
ba s
we e
obse ed
in
his
g o e,
each
oos -
ing
singly
unde
en s
in
sepa a e
ees.
On
3
Feb ua y,
wo
ba s
we e
obse ed
oos ing
oge he
in
a
ou h
en
in
ano he
ee
( ig.
3);
no
o he
ba s
we e
obse ed.
On
4
Feb-
ua y,
he e
we e
no
ba s
oos ing
unde
any
o
he
en s.
On
5
Feb ua y,
only
one
ba
was
ound;
i
was
oos ing
unde
he
same
en
ha
con ained
wo
ba s
on
3
Feb ua y.
On
2
Feb ua y
1991,
Timm
e u ned
o
his
si e
o
su ey
he
numbe s
o
U.
bilo-
ba um
in
his
coconu
g o e.
He
ound
a
o al
o
41
en s
in
29
o
he
54
ees.
Only
h ee
U.
biloba um
we e
ound,
each
oos ing
singly
unde
onds
ha
had
been
al e ed
o
o m
en s.
These
addi ional
obse a ions
emphasize
he
lexibili y
o
en -making
ba s
ega ding
oos
si es.
U.
biloba um
appea s
o
use
his
g o e
seasonally.
I
is
possible
ha
some
ba s
mo e
in o
o
ou
o
he
a ea
o
ack
changing
ood
esou ces
(Kunz,
1982);
mo emen
back
o
he
g o e
migh
e ol e
a ound
colonial
NO.
206
258
TIMM,
LEWIS:
URODERMA
TENT
CONSTRUCTION
Fig.
3.
Two
U ode ma
biloba um
oos ing
in
a
coconu
palm
en .
Pho og aph
cou esy
o
Ka h yn
E.
S one .
ma e ni y
oos ing
and/o
ma ing.
Addi ion-
al
esea ch,
especially
ollowing
ma ked
ba s
h oughou
he
yea ,
would
be
ex emely
aluable
in
de e mining
he
ac o s
a ec ing
mo emen s
and
choice
o
oos s
by
hese
ba s.
REFERENCES
Bake ,
R.
J.
1981.
Ch omosomal
low
be ween
ch omo-
somally
cha ac e ized
axa
o
a
olan
mammal,
U ode ma
biloba um
(Chi-
op e a:
Phyllos oma idae).
E olu ion
35(2):
296-305.
Bake ,
R.
J.,
and
C.
L.
Cla k
1987.
U ode ma
biloba um.
Mammal.
Spe-
cies
279:
1-4.
Ba bou ,
T.
1932.
A
peculia oos inghabi
o
ba s.
Q.
Re .
Biol.
7(3):
307-312.
B ooke,
A.
P.
1987.
Ten
cons uc ion
and
social
o ganiza-
ion
in
Vampy essa
nymphaea
(Chi op-
e a:
Phyllos omidae)
in
Cos a
Rica.
J.
T op.
Ecol.
3(2):
171-175.
1990.
Ten
selec ion,
oos ing
ecology
and
so-
cial
o ganiza ion
o
he
en -making
ba ,
Ec ophylla
alba,
in
Cos a
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J.
Zool.
(London)
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Chapman,
F.
M.
1932.
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ba .
Na .
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32(6):
555-556.
Choe,
J.
C.,
and
R.
M.
Timm
1985.
Roos ing
si e
selec ion
by
A ibeus
wa -
soni
(Chi op e a:
Phyllos omidae)
on
An hu ium
a enii
(A aceae)
in
Cos a
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J.
T op.
Ecol.
1(3):
241-247.
Fleming,
T.
H.,
E.
T.
Hoope ,
and
D.
E.
Wilson
1972.
Th ee
Cen al
Ame ican
ba
commu-
ni ies:
s uc u e,
ep oduc i e
cycles,
and
mo emen
pa e ns.
Ecology
53(4):
555-
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Goodwin,
R.
E.
1979.
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o
Timo :
sys ema ics
and
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Bull.
Am.
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Ha sho n,
G.
S.
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In
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(ed.),
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1991
