1
J.
ZooL,
Lond.
(1970)
161,
237-245
Roos -si e
selec ion
among
la -headed
ba s
(Tylonyc e is
spp.)
LORD
MEDWAY
School
o
Biological
Sciences,
Uni e si y
o
Malaya,
Kuala
Lumpu ,
Malaysia
AND
ADRIAN
G.
MARSHALL
Depa men
o
Na u al
His o y,
Ma ischal
College,
Uni e si y
o
Abe deen,
Sco land
{Accep ed
9
Decembe
1969)
(Wi h
2
pla es
and
1
igu e
in
he
ex )
Du ing
six
yea s,
1962
68,
he
biology
o
he
ba s
Tylonyc e is
pachypus
and
T.
obus ula
has
been
s udied
a
he
Uni e si y
o
Malaya
Field
S udies
Cen e,
Ulu
Gombak,
Selango ,
Malaysia.
Bo h
species
we e
o en
ound
oos ing
wi hin
in e nodes
o
he
bamboo
Gigan o-
chloa
sco echinii,
en e ing
and
lea ing
h ough
na ow
e ical
sli s
caused
by
he
bee le
Lasiochila
go yi.
In
his
pape
he
oos
si es
a e
desc ibed
and
e idence
is
p esen ed
ha
each
species
o
ba
ac i ely
selec ed
oos s
o
a
di e en
ange
o
dimensions.
The
ecological
implica ions
o
he
sepa a ion
o
he
species
by
his
ac o
a e
discussed.
Con en s
In oduc ion
Me hods
Resul s
Discussion
Summa y
Re e ences
Page
237
238
239
244
245
245
In oduc ion
The
genus
Tylonyc e is
Pe e s
(Chi op e a:
Vespe ilionidae)
comp ises
wo
species
T
pachypus
(Temmmck)
and
T.
obus ula
Thomas
(Medway,
in
p ess)
bo h
o
which
ha e
ex o si e
o e lapping
anges
m
Sou h
and
Sou h-eas
Asia
(Walke ,
1964),
In
he
S a es
Malaya,
Wes e n
Malaysia,
ecen
collec ions
ha e
shown
ha
bo h
a e
sympa ic
4(^ o
nT20
m
^Z,7 ;ni O eS namICVa l71S
anging
he
lowland'To^
4J0U
ee
(1220
m).
Al hough
dis inguishable
in
he
hand
by
a
numbe
o
cha ac e *
l^
.g ex u eandcolou
o
he
pelage,
body
measu emen s
and
weigh ,
bo h
species
a e
:
small
(Table
I)
and
simila
in
gene al
appea ance.
Bo h
exhibi
he
p incipal
cha ac e s
o
he
genus,
iz.
a
do so en ally
la ened
skull
( om
which
he
colloouial
Fnoli h
is
de i ed)
and
expanded
leshy
pads
a
he
base
i
he
pollex
oo .
These
cha ac e s
seem
o
be
adap a ions
o
oos ing
in
p o ec ed
ca i ies
wWch
can
18
237
238
LORD
MEDWAY
AND
A.
G.
MARSHALL
be
en e ed
only
h ough
na ow,
sli -shaped
ape u es.
In
Malaya
he
in e nodal
spaces
o
bamboo
p o ide
jus
such
oos
si es
when
he
culm
wall
has
been
pie ced
by
ail
open-
ing
o
app op ia e
shape
(see
also
Ha ison,
1966;
Lim,
1967).
The
Uni e si y o
Malaya
Field
S udies
Cen e
ese e
a
UluGombak
o ms
a
small
pa
o
an
ex ensi e
S a e Fo es
Rese e
co e ing
he
wes e n
slopes
o
he
Main
Range
o
Malaya
om
600
ee
(183
m)
ele a ion
upwa ds,
in
he
headwa e s
o
he
i e
Gombak
and
i s
ibu a ies,
a
3°
20'
N,
101°
45'
E.
A
majo
sec o
o
he
Field
S udies
Cen e
ese e
was
logged
comme cially
in
1956-58,
and
among
he
egene a ing
ege a ion
he
la ge
bamboo
Gigan ochloa
sco echinii
Gamble
is
abundan .
In
some
localized
a eas
his
bamboo
is
agg ega ed
o
o m
nea ly
pu e
s ands;
mo e
commonly,
isola ed
clumps
a e
sepa a ed
by
b oad-lea ed
o es .
Du ing
he
cou se
o
his
in es iga ion,
be ween
1962
TABLE
I
Selec ed
measu emen s
and
weigh s
o
adul
Tylonyc e is
pachypus
and
T.
obus ula;
aken
om
Medway
(1969)
T.
pachypus
T.
obus ula
Head
and
body
leng h
(mm)
35-40
45-50
Fo ea m
leng h
(mm)
25-28
27-32
Condylocanine
leng h
(mm)
10-4-10-6
12-3-12-7
Li e
weigh
(g)
3-5-5-0
7-0-10-0
and
1968,
a
o al
o
448
oos ing
pa ies
(i.e.
one
o
mo e
ba s
a
oos
on
one
occasion
in
a
single
in e node)
o
bo h
species
o
Tylonyc e is
we e
aken
om
in e nodes
o
G.
sco e-
chinii.
In
he
p esen
pape
he
oos
si es
a e
desc ibed
and
e idence
is
p esen ed
ha
each
species
o
ba
ac i ely
selec ed
oos s
o
a
di e en
ange
o
dimensions.
The
ecological
implica ions
o
he
sepa a ion o
he
species
by
his
ac o
a e
discussed.
Me hods
Fla -headed
ba s
a
oos
in
bamboo
we e
aken
by
wo
me hods. Fi s ly,
wi hin
a
small
a ea
o
egene a ing
o es
a
he
Field
S udies
Cen e,
78
ac i e
oos
si es
we e
indi idually
ma ked
by
se ial
numbe .
A
in e als
a ying
i egula ly
om
one
day
o
se e al
mon hs,
hese
oos s
( e e ed
o
in
he
ex
below
as
"ma ked
oos s")
we e
inspec ed
and
any
ba s
p esen
we e
caugh
on
eme ging
by
means
o
a
simple
wi e
mesh
unnel- ap
ied
o e
he
en ance
hole,
secondly,
an
addi ional
83
oos ing
g oups
we e
collec ed
in
1966-68
by
elling
he
bamboo
culm,
and
cu ing
ou
he
occupied
in e node
which
was
hen
b ough
back
o
he
Field
S udies
Cen e.
These,
e e ed
o
below
as
"collec ed
oos s",
we e
aken
om
di e en
sec o s
o
he
Ulu
Gombak
S a e
Fo es
Rese e
wi hin
a
adius
o
2
miles
om
he
a ea
o
ma ked
oos s.
Selec ed
measu emen s
we e
aken
o
he
bamboo
in e nodes
o
collec ed
oos s
a e
elling,
and
o
he
ma ked
oos s
in
si u.
The
dimensions
o
he
en ance
holes
we e
measu ed
o
he
nea es
0-1
mm
by
enginee s'
sliding
calipe s
wi h
a
e nie
scale,
he
leng h
and
wid h
eco ded
being
he
g ea es
dimensions
measu ed
a
any
poin
o
he
hole.
Culm
diame e
was
measu ed
by
calipe s
o
he
nea es
1
mm,
and
o he
dimensions
by
ule
o
lexible
ape
o
he
nea es
cm.
ROOST-SITE
SELECTION
AMONG
BATS
239
Resul s
The
s em
wall
o
he
s anding,
unb oken
bamboo
culm
may
be
pie ced
by
holes
o
many
shapes
and
sizes
a
di e en
heigh s
om
he
g ound
as
a
esul
o
he
ac i i y o
a
a ie y
o
di e en
agen s.
Acciden al
damage
o
he
young
shoo
in
an
ea ly
s age
o
g ow h
can
p oduce
longi udinal
ows
o
la ge,
mo e
o
less
coalesced
holes
in
he
wall
o
he
ma u e
culm
a
any
heigh .
An
unde e mined
numbe
o
bo ing
insec
la ae
make
holes
and
unnels
o
a ying
diame e s
cha ac e is ic
o
each
species.
And
inally
he
ac i i y
o
e eb a es,
including
woodpecke s,
he
Pencil- ailed
ee
mouse
Chi opodomys
gli oides
and
man,
will
p oduce
new
openings
in
he
bamboo
s em
wall
o
modi y
exis ing
holes.
Despi e
he
a ie y
o
holes
a ailable,
we
ound
ha
in
he
Ulu
Gombak
ese e
bo h
species
o
la -headed
ba
oos ed
only
in
in e nodes
en e ed
by
holes
o
a
limi ed
and
cha ac e is ic
size
ange
(Fig.
1).
These
holes
in
e e y
case
we e
na ow
e ical
sli s
in
he
walls
o
s anding
g een
culms
o
he
one
bamboo
species,
Gigan ochloa
sco echinii.
All
collec ed
oos s
we e
b oken
open
and
examined
in e nally.
In
mos
cases
only
one
sizeable
90
85
80
75
70
65
60
$
o
o
55
-
:
50
45
40
<J
o
J
o
i
o
o
•
o o
•
o
H
• •
A
•
A
A
o
O
A
•
35.
20
25
30
35
40
45
60
65
70
75
80
85
50
55
Leng h
(mm)
FIG.
1.
Size
o
oos
en ance
hole
in
ela ion
o
species
o
Tylonyc e is.
T.
pachypus:
O,
single
collec ion;
•,
h ee
o
mo e
appings.
T.
obus ula:
•,
single
collec ion;
•,
h ee
o
mo e
appings.
T.
pachypus
and
T.
obus ula:
A,
h ee
o
mo e
appings.
240
LORD
MEDWAY
AND
A.
G.
MARSHALL
hole
ga e
access
o
he
in e nodal
space
in
which
he
ba s
we e
ound,
and
he
noda!
sep um
a
each
end
was
in ac .
In
one
ins ance
among
collec ed
oos s
and
in
a
lea-.
h ee
ins ances
among
ma ked
oos s
he e
we e
wo
o
mo e
holes
in
he
culm
wall
la ge
enough
o
allow
he
ba s
o
en e
bu
in a iably
one
could
be
dis inguished
as
he
unc ion-
al
en ance
by
a
de ini e
smoo hing
o
i s
edges,
p esumably
he
esul
o
ic ion
o
he
ba s
passing
h ough
i .
,
In
all
cases
i
appea ed
ha
u ilized
en ance
holes
we e
o iginally
made
by
he
bee le
Lasiochila
go yi
(Gue in)
(Coleop e a,
Ch ysomelidae)
on
eme gence
om
he
in e node
(see
Pla e
I).
Salien
aspec s
o
he
biology
o
his
bee le
in
he Ulu
Gombak
Fo es
Rese e
ha e
been discussed
by
Macdonald
(1960)
unde
he
name
Anisode a
go yi,
and
ou
in es iga ions
con i m
his
obse a ions.
A
leas
i e
o he
species
o
bamboo
occu
in
he
Ulu
Gombak
Fo es
Rese e
bu
no
ins ances
o
a ack
by
L.
go yi
has
been
obse ed
among
hese,
and
no
la -headed
ba s
ha e
been
collec ed
om
hem.
The
adul
bee le
appa en ly
lays
on
he
g owing
ips
o
young
bamboo
shoo s.
On
ha ching
he
la a
bo es,
h ough
he
culm
wall
in o
he
in e nodal
space,
whe e
i
eeds
on
he
in e nal
lining
o
pa enchyma ous
issue.
When
ully
g own
he
la a
exca a es
a
sho
e ical
unnel
in
he
culm
wall,
whe e
i
pupa es.
On
eme gence,
he
adul
bee le
inally
b eaks
he
hin
skin
co e ing
he
pupal
chambe .
A e
a
ime
he
esidual
dead
issue
su ounding
he
eme gence
unnel
decays,
becoming
so
and
iable
so
ha
i
b eaks
away
easily.
The
ac ion
o
woodpecke s
in
some
cases
has
clea ly
modi ied
he
ou line
o
he
ape u e,
and
he
ba s
hemsel es,
me ely
by
passing
in
and
ou ,
will
ub
o
ough
edges.
In
he
cou se
o
he
su ey,
we
ha e
obse ed
la -headed
ba s
bo h
en e ing
and
lea ing
hei
oos s.
Be o e
eme ging,
he
ba 's
muzzle
can
be
seen
h ough
he
sli
(Pla e
11(a)).
appa en ly
sensing
he
ou side
en i onmen
by
smell
o
echoloca ion,
o
bo h.
A e
a
sho
pause
he
ba
comes
ou
head
i s ,
wi h
he
body
u ned
so
ha
i s
do sal
and
abdomi
-
nal
su aces
a e
pa allel
o
he
long
axis
o
he
ape u e. The
p ocess
o
eme gence
is
ex emely
apid,
and
we
ha e
no
succeeded
in
pho og aphing
i .
As
a
as
can
be
seen,
he
w is s
eme ge
alongside
he
head
and
a e
applied
o
he
ex e nal
su ace
o
he
bamboo.
By
ex ension
o
he
elbow
join ,
he
ba
hen
hea es
i s
body
h ough
he
na ow-
sli .
We
ha e
gained
a
momen a y
imp ession
o
he
ba
s a ic
b ie ly
wi h
i s
unk
ou -
side
he
hole
and
i s
o ea ms
mo e
o
less
ully
ex ended;
an
ins an
la e
i
is
in
he
ligh
(Pla e
11(b)).
The
p ocess
o
en e ing
is
equally
apid.
Usually
he
ba
makes
mo e
han
one
app oach,
on
each
occasion
educing
ligh
speed
so
ha
i
almos
ho e s
in
on
o
he
hole—again
appa en ly
scanning
by
acous ic
o
ol ac o y
sense.
On
he
inal
app oach,
i
en e s
di ec ly
wi hou
a
isible
b eak
in
ligh .
I
goes
in
head
i s ,
again
wi h
he
body
u ned
sideways.
The
en i e
mo emen
is
no
less
apid
han
eme gence
and,
in
he
absence
o
pho og aphs,
de ails
o
he
ac ion
a e
no
clea .
Once
inside
he
culm,
he
ba s
oos
a
he
uppe
end
o
he
in e nodal
ca i y,
suppo ed
p incipally
by
he
claws
o
hei
hind
ee
a aching
o
small
i egula i ies
in
he
su ace.
The
expanded
pads
on
he
soles
and
he
ca pal
join s
appa en ly
assis
he
ba s
in
g ipping
such
a
su ace.
They
a e
no
uly
suc o ial,
bu
he
ba s
can
o
ins ance
climb
backwa ds
up
he
wall
o
a
glass
ja
o
sui able
diame e .
I
is
no ewo hy
ha
he
pollex
is
p opo -
iona ely
sho
and
i s
claw
educed.
Tha
his
digi
is
unimpo an
in
main aining
hold
a
oos
is
indica ed
by
a
specimen
o
T.
obus ula,
collec ed
as
an
adul
a
oos
wi h
o he s,
which
lacked
a
claw
on
bo h
pollices
by
congeni al
de o mi y,
appa en ly
wi hou
su e ing
ill
e ec .
J.
Zoo/.,
Loud.
(1970)
161
PLATE
I.
(a)
A
s and
o
he
bamboo
Gigan och/oa
sco echinii
(c oss
poles
used
o
mo e
om
one
oos ing
si e
o
ano he ).
(b)
Adul
Lasiochila
go yi
bu owing
in
bamboo.
I
will
eme ge
om
he
bamboo
o
he
le
a
he
op
o
he
bu ow;
he
walls
do sal
and
en al
o
he
bee le
will
hen
collapse
o ming
he
oos
en ance
hole.
(c)
Tylonyc e is
pachypus
by
a
ypical
oos
en ance
hole.
[To
ace
page
240
J.
Zool.
Loud,
(1970)
161
PLATE
II.
Tylonyc e is
obus ula
eme ging
om
i s
oos
in
he
bamboo
Gigan ochloa
sco echinii.
(a)
A
momen
be o e
eme ging,
he
muzzle
o
he
ba
is
seen
in
he
uppe
pa
o
he
oos
en ance;
(b)
he
ba
sho ly
a e
eme ging.
(J.
D.
B oomhall.)
ROOST-SITE
SELECTION
AMONG
BATS
241
Du ing
he
cou se
o
his
su ey,
he
wo
species
o
Tylonyc e is
we e
only
once
ound
a
oos
oge he .
The
composi ion
o
402
oos ing
pa ies
was
accu a ely
de e mined
(Medway
&
Ma shall,
in
p ep.).
O
hese,
193
comp ised
only
T.
obus ula, 208
only
T.
pachypus,
and
one
bo h
species.
Fo y-six
oos s
we e
classed
as
incomple ely
collec ed,
ei he
because
one
o
mo e
ba s
escaped
be o e
being
examined,
o
because
one
o
mo e
ailed
o
eme ge
and
be
apped;
mixed
species
we e
no
ound
among he
emaining
ba s
in
hese
46
oos ing
pa ies.
Ye
epea ed
apping
a
he
ma ked
oos s
showed
ha
a
p opo ion
we e
occupied
by
bo h
species
on
di e en
occasions.
O
he
78
ma ked
oos s,
T.
pachypus
only
was
aken
om
34
(44%),
T.
obus ula
only
om
22
(28%)
and
bo h
species
om
22
(28%).
In
an
a emp
o
de e mine
he
ac o s
go e ning
he
selec ion
o
oos
si es
by
he
wo
species
o
ba ,
he
ollowing
measu emen s
ha e
been
analysed:
(i)
heigh
abo e
g ound
o
oos
en ance,
(ii)
leng h
o
occupied
in e node,
(iii)
ex e nal
diame e
o
culm
(i )
he
wid h
o
he
en ance
hole,
( )
e ical
leng h
o
he
en ance
hole,
( i)
posi ion
o
he
en ance
hole
in
ela ion
o
he
nodal
sep a.
Heigh
abo e
g ound
o
oos
en ance
The
heigh
abo e
g ound
o
he
occupied
in e nodes
o
collec ed
oos s
was
no
measu ed
exac ly
bu
appa en ly
a ied
om
abou
1
m
o
a
leas
10
m.
Among
he
ma ked
oos s,
he
heigh
o
he
base
o
he
en ance
hole
abo e
g ound
a ied
om
250-830
cm.
The
uppe
limi in
his
case
was
de e mined
by
he
g ea es
heigh
o
which
we
o
ou
assis an s
we e
able
o
climb,
and
is
acco dingly
biased
o
we
saw
appa en ly
sui able
holes
beyond
ou
each.
I
he e
is
an
uppe
limi ,
i
is
likely
o
be
imposed
by
he
p og essi e
educ ion
o
he
diame e
o
he
bamboo
culm
owa ds
i s
ip
(below).
Bo h
he
highes
and
he
lowes
ma ked
oos
yielded
only
T.
obus ula
on
apping,
and
he e
is
no
e idence
o
di e en
p e e ences
by
he
wo
species.
Leng h
o
in e node
The
ex e nal
in e nodal
leng h
and
ex e nal
diame e
o
he culm
we e
measu ed
o
all
collec ed
oos s
and
many
ma ked
oos s.
The in e nodal
leng h
o
36
collec ed
oos s
occupied
by
T.
obus ula
a ied
om
32-63
cm,
mean
48
cm,
and
o
45
collec ed
oos s
occupied
by
T.
pachypus
a ied
om
38-66
cm,
mean
also
48
cm.
In
his
species
o
bamboo,
he
ange
o
na u al
a ia ion
in
in e nodal
leng h
p obably
does
no
all
a
ou side
he
limi s
o
his
dimension
eco ded
among
occupied
oos s.
The e
is
he e o e
no
e idence
o
ac i e
selec ion
o
his
cha ac e
by
he
ba s.
Ex e nal
diame e
o
culm
The
ex e nal
diame e s
o
88
oos
si es
wi h
a
a ie y
o
di e en
his o ies
a e
gi en
in
Table
II.
The
di e ence
be ween
he
mean
culm
diame e
o
{a)
collec ed
oos s
o
T.
pachypus
(70
mm)
and
o
(g)
collec ed
oos s
o
T.
obus ula
(78
mm)
is
signi ican
(P
<0-001).
The
mean
ex e nal
diame e s
o
all
ca ego ies
o
ma ked
oos s
occupied
a
leas
once
by
T.
obus ula
a e
close
o
he
igu e
o
collec ed
oos s
o
his
species,
al hough
he
samples
a e
oo
small
o
he
di e ence
be ween
means,
e.g.
(b)l( ),
o
(d)j( ),
o
be
signi ican .
I
is
no ewo hy
ha
o
ma ked
oos s
in
ca ego ies
(c),
(d)
and
(e),
i.e.
hose
om
which
bo h
species
we e
apped,
he
minimum
diame e
is
wi hin
he
limi s
o
242
LORD
MEDWAY
AND
A.
G.
MARSHALL
oos s
om
which
only
T.
obus ula
was
apped,
and
he
maximum
diame e
is
wi hin
he
limi s
o
oos s
om
which
only
T.
pachypus
was
apped.
The
lowe
pa s
o
he
culm
o
Gigan ochloa
sco echinii
wi hin
he
ange
o
heigh s
o
ou
ma ked
oos s
a e
mo e
o
less
pa allel-sided,
so
ha
he e
is
li le
a ia ion
in
diame e
be ween
successi e
in e nodes.
The
ange
o
a ia ion
be ween
di e en
culms,
howe e ,
is
ela i ely
g ea
co esponding
closely
o
he
ange
o
a ia ion
o
occupied
oos
si es.
A
Gombak
ew
culms
ha e
been
obse ed
hicke
han
he
g ea es
measu ed
diame e
o
any
occupied
oos ,
i.e.
102
mm
o
T.
obus ula,
al hough
Hol um
(1958)
eco ded
12
cm
as
he
maximum
o
he
species.
Bu
in
each
clump
he e
a e
usually
one
o
wo
abno mally
slende
culms.
The
leas
diame e
measu ed
a
Gombak
is
38
mm,
well
below
he
lowe
limi s
eco ded
o
oos s
o
T.
pachypus.
The
indica ions
a e
ha
bo h
ba s
occup
oos
si es
abo e
a
ce ain
minimum
culm
diame e
which
is
species
speci ic.
TABLE
II
Roos
si es
o
Tylonyc e is
in
Ulu
Gombak:
ex e nal
diame e
o
bamboo
culms
a
oos
si e,
and
dimensions
o
en ance
ape u e
(mm)
F equency
o
No.
o
En ance
ape u e
Reco d
occu ence
oos s
Ex e nal
diame e
Ve ical
leng h
Ho izon al
wid h
ype
pachypus
obus ula
measu ed
Range
Mean
S.D.
Range
Mean
S.D.
Range
Mean
S.D.
a
Collec ed
1
0
36
48-90
70
10-8
23-64
33
9
3-9-6-5
4-9
0-6
b
Ma ked
>3
0
5
52-97
68
17-5
26-55
42
13
4-3-5-4
5-0
0-5
c
Ma ked
>1
1
6
68-87
78
7-2
29-46
37
8
4-6-60
5-2
0-5
d
Ma ked
>1
>1
4
66-96
81
12-4
44-56
49
5
6-0-6-6
6-3
0-3
e
Ma ked
1
>1
1
94
— —
54
— —
6-4
—
—
/
Ma ked
0
>3
8
62-96
79
12-5
37-81
57
18
6
0-7-4
6-8
0-4
g
Collec ed
0
1
28
56-102
78
11-9
31-63
44
8
5-5-8-7
6-5
0-8
This
same
ac o
p esumably
also
imposes
an
uppe
limi
o
he
heigh
o
he
oos
abo e
g ound,
e en
hough
his
was
no
obse ed.
Al hough
successi e
in e nodes
a y
li le
in
diame e
wi hin
he
heigh
ange
o
ou
ma ked
oos s
he e
is
a
dis inc
ape
a
he
uppe
end
o
he
bamboo
culm,
which
may
a ain
a
leng h
o
20
m
(Hol um,
1958).
In
he
e minal
egion
p og essi e
educ ion
in
diame e
will
p esumably
exclude
i s
T.
obus ula,
hen
T.
pachypus,
and
will
inally
impose
an
uppe
limi
o
he
heigh
abo e
g ound
o
accep able
oos
si es
o
bo h
species.
Ho izon al
wid h
o
en ance
hole
Table
II
also
gi es
he
dimensions
o
he
en ance
ape u e
o
he
88
oos
si es
examined,
and
om
Fig.
1
i
is
clea
ha
bo h
species
selec
holes
alling
in
a
limi ed
ange
o
size.
The
mos
c i ical
dimension
is
ho izon al
wid h.
The e
is
no
o e lap
be ween
he
wid h
o
en ance
holes
o
ma ked
oos s
used
h ee
o
mo e
imes
by
T.
pachypus
only
(ca ego y
b)
and
hose
wi h
a
simila his o y
o
occupa ion
by
T.
obus ula
(J).
Among
collec ed
oos s,
he
p e ious his o y
o
occupa ion
o
which
is
unknown,
he
ange
o
a ia ion
is
g ea e
ROOST-SITE
SELECTION
AMONG
BATS
243
o
bo h
species,
wi h
an
o e lap
o
1
mm.
Howe e ,
he
di e ence
be ween
he
mean
wid h
o
collec ed
oos s
occupied
a
he
ime
o
collec ion
by
T.
pachypus
(ca ego y
a)
and
hose
occupied
by
T.
obus ula
(g)
is
signi ican
(P
=
<0-001).
The
wid h
o
he
ape u e
is
p esumably
de e mined
by
he
gi h
o
he
bee le
la a
as
i
bo es
in o
he
culm
wall
sho ly
be o e
pupa ion.
Measu emen s
o
he
ex e nal
diame e
o
he
culm
and
wid h
o
he
en ance
hole
show
no
signi ican
co ela ion
(P>0T),
and
he
wo
cha ac e s
a e
e iden ly
independen .
Ve ical
leng h
o
en ance
hole
In
he
e ical
leng h
o
he
en ance
hole
a
p opo ionally
g ea e
ange
o
a ia ion
is
obse ed
(Table
II),
and
he e
is
b oad
o e lap
among
all
ca ego ies
o
oos .
The
di e ence
be ween
he
means
o
collec ed
oos s
o
T.
pachypus
(a)
and
hose
o
T.
obus ula
(g)
app oaches
signi icance
(P
=
0-1—0-05).
The
a ia ion
in
e ical
leng h
o
all
holes
appa en ly
made
by
he
bee le
Lasiochila
go yi
is
e iden ly
g ea e
han
he
a ia ion
o
u ilized
oos
en ances
o
bo h
species
o
ba s,
pa icula ly
a
he
lowe
limi .
The
e ical
leng h
o
he
hole
in
he
bamboo
culm
wall
depends
on
he
angle
and
leng h
o
he
pupal
chambe
which
he
bee le
exca a es.
We
ha e
equen ly
obse ed
holes
less
han
20
mm
long,
down
o
a
minimum
(14
mm)
which
is
appa en ly
li le
g ea e
han
he
leng h
o
he
pupa
i sel .
I
is
he e o e
clea
ha
bo h
species
o
Tylonyc e is
u ilize
oos
en ances
o
a
limi ed
ange
o
e ical
leng h.
Posi ion
o
en ance
hole
in
in e node
The
posi ion
o
he
oos
en ance
in
he
in e node
was
assessed
by
measu ing
he
dis ance
om
he
base
o
he
hole
o
he
node
below.
Among
all
collec ed
oos s
o
bo h
species,
he
en ance
hole
was
loca ed
in
he
lowe
hal
o
he
in e node,
wi h
a
single
excep ion
(T
obus ula).
In
his case,
he
in e nodal
leng h
was
38-5
cm,
he
en ance
hole
5-0
cm
long,
and
i s
base
30
cm
abo e
he
node
below.
The e
was
also
a
second
small
hole
lowe
in
he
s em,
only
5
cm
abo e
he
node.
In
h ee
o he
collec ed
oos s
o
T.
obus ula,
and
in
one
o
T.
pachypus,
a
subsidia y hole,
no
used
by
he
ba s,
was
no ed
low
e
on
he
s em
han
he
unc ional oos
en ance.
I
hese
i e
oos s
a e
excluded,
he
pe cen age
o
he
leng h
o
he
in e node
below
he
base
o
he
en ance
hole
a ied
om
2-39
/
among
T.
obus ula,
and
2-44%
among
T.
pachypus.
Inspec ion
o
o he
bamboos
has
shown
an
appa en ly
andom
dis ibu ion
o
holes
o
app op ia e
size
h oughou
he
leng h
o
he
in e node.
The
es ic ion
o
u ilized
holes
o
he
lowe
hal
o
he
in e node
hus
once
mo e
indica es
ac i e
selec ion
by
he
ba s.
As
a
likely
explana ion
we
no e
ha
excep
in
pe iods
o
p olonged
d ough ,
Tylonyc-
e is
oos s
in a iably
con ain
a
ce ain
amoun
o
liquid.
This
luid
may
o igina e
pa ly
as
conduc ed
sap
(Macdonald,
1960),
and
pa ly
om
he
u ine
and
aeces
o
he
ba s.
Mos
o
i ,
howe e ,
is
p obably
de i ed
om
ain
wa e
which
ickles
down
he
culm
and
en e s
h ough
any
ape u e,
illing
he
in e nodal
ca i y
o
he
le el
o
he
bo om
o
he
lowes
hole.
The
a ailable
oos ing
space
abo e
wa e
le el
in
an
in e node
is
he e o e
de e mined
by
he
heigh
o
he
lowes
hole
abo e
he
nodal
sep um.
This
is
usually
he
en ance
i sel ,
and
unless
he
en ance
is
low,
he
oos
space
is
limi ed.
The
ins ance
o
a
collec ed
oos
o
T.
obus ula
men ioned
abo e
demons a es
ha
an
in e node
wi h
an
en ance
in
he
uppe
hal
will
be
selec ed
i
d ainage
is
p o ided
by
a
lowe
hole.
244
LORD
MEDWAY
AND
A.
G.
MARSHALL
Discussion
I
should
be
s essed
ha
bamboo
culms
pie ced
by
he
eme gence
holes
o
Lasiochila
go yi
do
no
p o ide
he
only
accep able oos ing
si es
o
la -headed
ba s.
E en
in
he
Ulu
Gombak
o es
ese e,
a
single
T.
pachypus
has
been
ound
oos ing
in
a
na ow-
ca i y
be ween
boulde s
(R.
L.
Rudd,
pe s.
comm.).
Nea
Kuching,
Sa awak,
Medway
has
collec ed
oos ing
g oups
o
T.
pachypus
in
he
in e nodal
spaces
o
uniden i ied
bamboos
en e ed
by
la ge ,
mo e
i egula
holes
han
hose
p oduced
by
L.
go yi
(Table
111).
Ye
in
he
Ulu
Gombak
o es
he
la ge
numbe
o
sui able
oos
si es
esul ing
om
a acks
o
he
bee le
on
he
common
bamboo
is
undoub edly
he
majo
ac o
con ibu ing
o
he
local
abundance
o
la -headed
ba s.
TABLE
III
Roos
si es
o
Tylonyc e is
pachypus
in
bamboo
a
Kampong
Segu,
nea
Kuching
(1963)
En ance
hole
dimensions
(mm)
No.
No.
Culm
diame e
Ve ical
Ho izon al
33
$$
(cm)
leng h
b ead h
Na u e
o
en ance
1
1
13-0
90
10
I egula
ea
1
12
15-5
60
13 I egula
gash
1
6
12-5
38
8
I egula
ea
Bo h
species
o
Tylonyc e is
appea
equally
able
o
ake
ad an age
o
his
si ua ion.
As
shown
abo e,
bo h
occupy
he
same
oos
si e
on
di e en
occasions.
Apa
om
sligh
di e ences
in
he
size
and
composi ion
o
oos ing
g oups,
we
ha e
been
unable
o
de ec
any
impo an
di e ences
in
hei
ecology
(Medway
&
Ma shall,
in
p ep.).
Bo h
species
ha e
been
obse ed
eeding
oge he
on
e mi e
swa ms
a
dusk,
al hough
we
ha e
li le
knowledge
o
hei
p ecise
ood
p e e ences.
The
en i e
popula ions
o
bo h
a
Gombak
appa en ly
gi e
bi h
du ing
he
same
es ic ed
pe iod
each
yea
(de ails
in
p epa a ion).
Such
ex ensi e
o e lap
o
ecological
niche
among
closely
ela ed
sympa ic
species
is
no
unique
in
he
opical
en i onmen
(c .
Klop e ,
1961,
o
bi ds),
and
i
is
o
in e es
o
examine
he
isola ing
mechanisms
which
es ic
in e speci ic
compe i ion
and
p e en
he
local
ex inc ion
o
one
species
("Gause's
hypo hesis").
Among
he
sample
o
78
si es
ep esen ed
by
he
ma ked
oos s,
28
%
we e
occupied
by
bo h
species.
Ye
o
448
oos ing
pa ies
only
one
(0-023%) was
de ini ely
known
o
ha e
included
bo h.
This
di e ence
indica es
ha
he
wo
species
a e
no mally
exclusi e
and
do no
olun a ily
associa e
a
oos .
I
he
supply
o
mu ually
accep able
oos s
was
limi ed,
he
wo
species
would
be
compe ing
o
a
common esou ce.
A
Gombak.
howe e ,
he e
is
appa en ly
a
g ea
excess
o
sui able oos ing
si es
a ailable;
on
any
one
day
only
a
e y
small
pe cen age
o
some imes
none
o
he
ma ked
oos ing
si es
examined
would
be
ound
o
con ain
ba s.
This
may
o
some
ex en
be
an
"unna u al"
si ua ion
(c .
Lack,
1968;
6-7),
in
which
he
common
esou ce
is
no
limi ed
and
Gause's
hypo hesis
is
no
applicable.
None
he
less,
ou
esul s
clea ly
indica e
ha
bo h
la -headed
ba s
ac i ely
selec
oos
si es
which
di e
wi h
espec
o
ce ain
me ical
cha ac e s.
The
ROOST-SITE
SELECTION
AMONG
BATS
245
cha ac e s
o
which
s a is ically
signi ican
di e ences
in
mean
dimensions
ha e
been
demons a ed
be ween
oos s
o
he
wo
species
a e
culm
diame e ,
and
he
leng h
and
he
wid h
o
he
en ance
hole.
O
hese,
he
las
is
appa en ly
he
mos
c i ical,
al hough
in
all
he e
is
a
b oad
zone
o
o e lap
eco ded.
Fo
bo h
species
he e
is
appa en ly
a
p e e ed
uppe
limi
o
he
size
o
he
en ance,
which
es ic s
he
ype
o
hole
chosen
o
hose
made
by
L.
go yi.
The
equally
c i ical
lowe
limi
is
p esumably
dic a ed
by
body
size,
being
e ec i ely
he
smalles
sli
h ough
which
he
skull
and
unk
o
he
ba
can
pass.
The
smalle
species
T.
pachypus
hus
has
a ailable
o
i
a
ange
o
oos s
wi h
na owe
en ances,
om
which
T.
obus ula
is
excluded
by
i s
physical
size.
Ecological
sepa a ion
o
he
wo
species
may
be
based
only
on
his
small
di e ence.
Summa y
Tylonyc e is
pachypus
and
T.
obus ula
a e
locally
abundan
in
Ulu
Gombak,
Selango ,
Malaysia
whe e
appa en ly
sui able
oos ing
si es,
associa ed
wi h
he
bamboo
Gigan o-
chloa
sco echinii
and
he
bee le
Lasiochila
go yi,
a e
a ailable
in
g ea
excess.
Al hough
he
wo
species
appea
o
occupy
e y
simila
ecological
niches,
hey
do
no
olun a ily
associa e
a
oos ;
al hough
28%
o
ma ked
oos ing
si es
we e
seen
o
be
occupied
by
bo h
species
on
di e en
occasions,
only
0-023
%
o
oos ing
pa ies
examined
we e
known
o
ha e
included
bo h
species
o
ba
oge he .
Measu emen
o
a ious
dimensions
o
he
oos s
indica ed
ha
s a is ically
signi ican di e ences
be ween
oos s
o
he
wo
species
exis ed
be ween
he
culm
diame e ,
and
he
leng h
and
he
wid h
o
he
en ance
hole;
he
las
was
appa en ly
he
mos
c i ical.
The
lowe
limi
o
hole
wid h
is
dic a ed
by
he
body
size
o
he
ba ,
and
he
small
T.
pachypus
has
a ailable
o
i
a
ange
o
oos s
wi h
na owe
en ances
om
which
he
la ge
T.
obus ula is
excluded.
Ecological
sepa a ion
o
hese
wo
sympa ic
species
may
be
based
only
on
his
small
di e ence.
We
a e
bo h
much
indeb ed
o
all
hose
mo e
agile
han
ou sel es
who
assis ed
in he
ield
collec ion
o
la -headed
ba s,
no ably
Inche
Bah
Te a
and
Inche
Bah
Chong
o
he
Uni e si y
o
Malaya
Field
S udies
Cen e,
Ulu
Gombak.
D
J.
A.
Bullock
kindly
sugges ed
app op ia e
s a is ical
es s.
Me al
bands
o
use
in
his
p ojec
we e
p o ided
by
he
Aus alian
ba -banding
scheme,
C.S.I.R.O.,
Wildli e
Di ision.
Ma shall
is
mos
g a e ul
o
he
Ca negie
T us
o
he
Uni e si ies
o
Sco land
o
a
g an
which
made
possible
his
wo
yea
secondmen
om
he
Uni e si y
o
Abe deen
o
he
Uni e si y
o
Malaya.
REFERENCES
Ha ison,
J.
(1966).
An
in oduc ion
o
mammals
o
Singapo e
and
Malaya.
Singapo e
B anch,
Malayan
Na u e
Socie y.
Hol um,
R.
E.
(1958).
The
bamboos
o
he
Malay
Peninsula.
Ga dens
Bull.,
Singapo e
16:
1-135.
Klop e ,
P.
H.
(1961).
On he
causes
o
opical
species
di e si y;
niche
o e lap.
Am.
Na .
95:
223-226.
Lack,
D.
(1968).
Ecological
adap a ions
o
b eeding
among
bi ds.
London:
Me huen
&
Co.
Lim,
B.
L.
(1967).
Abundance
and
dis ibu ion
o
Malaysian
ba s
in
di e en
ecological
habi a s.
Fedn
Mus.
J.
(N.S.)
11
(1966):
61-76.
Macdonald,
C.
W.
(1960).
Sys ema ics
and
ecology
o
Leices e ia.
S ud.
Ins .
Med.
Res.
Malaya
29:
110-153.
Medway,
Lo d,
(1969).
The
wild
mammals
o
Malaya.
Kuala
Lumpu :
Ox o d
Uni e si y
P ess.
Medway,
Lo d
(in
p ess).
The
s a us
o
Tylonyc e is
malayana
Chasen.
Bull.
na n.
Mus.,
Singapo e.
Medway,
Lo d
&
Ma shall,
A.
G.
(in
p epa a ion).
Roos ing
associa ion
among
la -headed
ba s
(Tylonyc e is
spp.).
Walke ,
E.
P.
(1964).
Mammals
o
he
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