The distribution and ecology of mammals on Leyte, Biliran, and Maripipi Islands, Philippines


Rickart, E.A.; Heaney, L.P.; Heideman, P.D.; Utzurrum, R.C.B.

Fieldiana Zoology 72: 1-62

1993


The land mammal faunas of the Philippine islands of Leyte, Biliran, and Maripipi are described. Collectively, a total of 48 indigenous species are known to occur on the three islands: 45, 30, and 25 from Leyte, Biliran, and Maripipi, respectively. These include 1 insectivore, 1 dermopteran, 33 bats, 2 primates, 7 rodents, 2 carnivores, and 2 ungulates. All but three of these species occur on the large neighboring island of Mindanao. The exceptions include a widespread but rarely captured vespertilionid bat and two rodents that have sister-species on Mindanao. Among the remaining 45 species, 9 are restricted to the Mindanao faunal region, 10 are more widely distributed in the oceanic Philippines, and 26 have distributions that extend outside of the Philippines. One species from Biliran and two from Maripipi (all poorly known insectivorous bats) have not been recorded from Leyte. Otherwise the faunas of the two smaller islands are subsets of the Leyte fauna, which is, in turn, a subset of the Mindanao fauna. Data indicate that the present-day mammal faunas of these islands have been shaped primarily by events that occurred during the last 12,000 years. Results generally support predictions concerning elevational patterns in species richness and abundance (decreasing with elevation for small fruit bats, increasing with elevation for small non-volant species) and habitat associations of endemic versus non-endemic species (the former restricted to pristine or lightly disturbed habitats, the latter predominant in disturbed habitats). However, some of these patterns appear to break down on small land-bridge islands with relatively depauperate faunas. Most notably, commensal or widespread Asian species are abundant in undisturbed or lightly disturbed habitats when there are few endemic species present. Many of the results have direct implications for wildlife conservation in the Philippines. Patterns of deforestation render lowland species most vulnerable. Habitat associations of species indicate that disturbances to natural habitat have a disproportionately greater negative impact on the unique endemic portions of a local fauna. The depauperate nature of faunas on small land-bridge islands demonstrates that relatively large faunal reserves are required to provide long-term protection for many species. Furthermore, elevational range restrictions or special ecological requirements of some species necessitate reserves that encompass the full range of regional habitat diversity.

The
Distribution
and
Ecology
of
Mammals
on
Leyte,
Biliran,
and
Maripipi
Islands,
Philippines
Eric
A.
Rickart
Lawrence
R.
Heaney
Paul
D.
Heideman
Ruth
C.
B.
Utzurrum
Abstract
The
land
mammal
faunas
of
the
Philippine
islands
of
Leyte,
Biliran,
and
Maripipi
are
de-
scribed.
Collectively,
a
total
of
48
indigenous
species
are
known
to
occur
on
the
three
islands:
45,
30,
and
25
from
Leyte,
Biliran,
and
Maripipi,
respectively.
These
include
1
insectivore,
1
dermopteran,
33
bats,
2
primates,
7
rodents,
2
carnivores,
and
2
ungulates.
All
but
three
of
these
species
occur
on
the
large
neighboring
island
of
Mindanao.
The
exceptions
include
a
widespread
but
rarely
captured
vespertilionid
bat
and
two
rodents
that
have
sister-species
on
Mindanao.
Among
the
remaining
45
species,
9
are
restricted
to
the
Mindanao
faunal
region,
10
are
more
widely
distributed
in
the
oceanic
Philippines,
and
26
have
distributions
that
extend
outside
of
the
Philippines.
One
species
from
Biliran
and
two
from
Maripipi
(all
poorly
known
insectivorous
bats)
have
not
been
recorded
from
Leyte.
Otherwise
the
faunas
of
the
two
smaller
islands
are
subsets
of
the
Leyte
fauna,
which
is,
in
turn,
a
subset
of
the
Mindanao
fauna.
Data
indicate
that
the
present-day
mammal
faunas
of
these
islands
have
been
shaped
primarily
by
events
that
occurred
during
the
last
12,000
years.
Results
generally
support
predictions
con-
cerning
elevational
patterns
in
species
richness
and
abundance
(decreasing
with
elevation
for
small
fruit
bats,
increasing
with
elevation
for
small
non-volant
species)
and
habitat
associations
of
endemic
versus
non-endemic
species
(the
former
restricted
to
pristine
or
lightly
disturbed
habitats,
the
latter
predominant
in
disturbed
habitats).
However,
some
of
these
patterns
appear
to
break
down
on
small
land-bridge
islands
with
relatively
depauperate
faunas.
Most
notably,
commensal
or
widespread
Asian
species
are
abundant
in
undisturbed
or
lightly
disturbed
hab-
itats
when
there
are
few
endemic
species
present.
Many
of
the
results
have
direct
implications
for
wildlife
conservation
in
the
Philippines.
Patterns
of
deforestation
render
lowland
species
most
vulnerable.
Habitat
associations
of
species
indicate
that
disturbances
to
natural
habitat
have
a
disproportionately
greater
negative
impact
on
the
unique
endemic
portions
of
a
local
fauna.
The
depauperate
nature
of
faunas
on
small
land-bridge
islands
demonstrates
that
rela-
tively
large
faunal
reserves
are
required
to
provide
long-term
protection
for
many
species.
Furthermore,
elevational
range
restrictions
or
special
ecological
requirements
of
some
species
necessitate
reserves
that
encompass
the
full
range
of
regional
habitat
diversity.
Introduction
The
Philippine
Islands
support
a
diverse
mam-
mal
fauna
of
about
170
species,
many
of
which
are
endemic
to
the
archipelago
(Heaney,
1986,
1991a;
Heaney
et
al.,
1987).
Most
studies
of
Phil-
ippine
mammals
have
focused
on
the
faunas
of
the
highlands
of
the
two
largest
islands,
Luzon
and
Mindanao,
principally
because
early
work
re-
vealed
many
endemic
species
in
these
areas.
As
a
result
of
this
limited
focus,
the
mammals
of
most
of
the
smaller
Philippine
islands
remain
poorly
known.
Recent
studies
on
some
small
and
medi-
um-sized
islands
in
the
archipelago
have
yielded
a
wealth
of
information
on
patterns
of
mammalian
distribution
(Heaney,
1986;
Heaney
et
al.,
1989,
FIELDIANA:
ZOOLOGY,
N.S.,
NO.
72,
AUGUST
31,
1993,
PP.
1-62
1991;
Heaney
&
Rickart,
1990)
and
have
under-
scored
the
importance
of
such
areas
both
as
nat-
ural
laboratories
and
as
potential
faunal
reserves.
Unfortunately,
this
unique
fauna
is
now
se-
verely
threatened
by
habitat
destruction.
Primary
forest
cover
in
the
Philippines
as
a
whole
has
been
reduced
from
more
than
80%
of
the
total
land
area
near
the
turn
of
the
century
to
ca.
8%
in
1988
(Myers,
1988),
and
the
destruction
continues
at
an
alarming
rate.
The
impact
of
deforestation
on
wildlife
may
be
particularly
severe
on
small
is-
lands,
where
population
and
community
struc-
tures
may
be
less
robust.
Forest
destruction
also
poses
a
very
direct
threat
to
human
welfare.
In
addition
to
the
loss
of
invaluable
forest
resources,
the
elimination
of
natural
watershed
protection
in
upland
regions
greatly
increases
the
likelihood
of
seasonal
flooding.
On
5
November
1991,
this
last
possibility
became
a
terrible
reality
in
northern
Leyte,
where
thousands
of
Filipinos
lost
their
lives
in
catastrophic
floods
and
landslides
associated
with
a
tropical
storm.
The
scope
of
this
disaster
was
certainly
magnified
by
the
extensive
logging
that
has
virtually
eliminated
interior
forests
throughout
most
of
the
region.
In
this
paper
we
present
the
first
synoptic
lists
of
mammals
from
three
islands
in
the
central
Vi-
sayan
group:
Leyte,
the
eighth
largest
island
in
the
Philippine
archipelago,
and
Biliran
and
Maripipi,
two
small
islands
immediately
north
of
Leyte
(fig.
1).
All
three
were
part
of
the
island
of
Greater
Mindanao
during
a
period
of
lower
sea
level
in
the
late
Pleistocene,
becoming
isolated
about
10,000
years
ago
(Heaney,
1986).
In
relation
to
present-
day
Mindanao
(with
an
area
of
99,078
km
2
),
Leyte
(7,213
km
2
),
Biliran
(498
km
2
),
and
Maripipi
(22
km
2
)
represent
a
graded
series
of
sizes,
each
dif-
fering
from
the
next
by
a
similar
factor.
Each
of
these
islands
retains
some
areas
representing
the
three
major
types
of
natural
forest
vegetation:
low-
land
forest,
montane
forest,
and
mossy
forest
(or
dipterocarp,
lower
montane,
and
upper
montane
rain
forest;
sensu
Whitmore,
1984).
It
is
our
purpose
to
assess
the
taxonomic
status
and
summarize
available
ecological
information
for
all
mammal
species
known
to
occur
on
each
island.
Because
many
of
these
species
are
poorly
known,
we
provide
as
much
detailed
information
as
is
practical.
Further
analysis
of
distributional
data
is
presented
elsewhere
(e.g.,
Heaney
et
al.,
1989;
Heaney
&
Rickart,
1990;
Heaney,
1991a).
History
of
Investigations
Although
there
are
scattered
early
distributional
records
of
mammals
from
Leyte
(Hollister,
1912,
1913;
Taylor,
1934),
the
history
of
comprehensive
collecting
on
these
islands
is
limited.
In
1963,
D.
P.
Empesso
of
Silliman
University
collected
bats
from
within
and
around
the
Cathedral
Cave
com-
plex
near
Inopacan,
Hindang
Municipality,
Leyte
(specimens
in
the
Royal
Ontario
Museum).
Dur-
ing
the
same
decade,
G.
L.
Alcasid
and
M.
Celes-
tino
of
the
Philippine
National
Museum
and
D.
S.
Rabor
of
Silliman
University
made
collections
in
the
highlands
of
northern
and
central
Leyte
(specimens
in
the
American
Museum
of
Natural
History
and
the
Delaware
Museum
of
Natural
History).
Most
of
these
localities
are
mapped
and
briefly
described
by
Parkes
(1973).
Only
small
por-
tions
of
these
collections
have
been
described
in
the
literature
(Heaney,
1985a;
Musser,
1982a,b;
Peterson,
1981;
Peterson
&
Fenton,
1970).
Prior
to
our
field
studies
(1981-1987),
no
surveys
had
been
conducted
on
either
Biliran
or
Maripipi.
A
preliminary
listing
of
non-volant
mammal
species
from
all
three
islands
appeared
in
Heaney
(1986);
however,
this
did
not
include
the
results
from
our
extensive
surveys
of
1987.
We
have
attempted
to
include
information
on
all
specimens
now
in
museums.
However,
most
of
the
information
presented
here
is
based
on
our
own
field
surveys.
Our
work
on
Leyte
was
con-
ducted
along
an
elevational
transect
on
the
west-
central
portion
of
the
island
(figs.
1,
2).
A
brief
reconnaissance
trip
was
made
to
this
region
from
23
to
28
May
1984
(voucher
specimens
in
the
University
of
Michigan
Museum
of
Zoology
[uMMz]),
but
most
work
took
place
from
2
March
to
13
April
1987
(specimens
in
the
Philippine
Na-
tional
Museum
[PNA,
the
Silliman
University
Museum
[su],
the
United
States
National
Museum
of
Natural
History
[usNm],
the
Visayas
State
Col-
lege
of
Agriculture
[viscA],
and
the
Western
Aus-
--)
FIG.
l
.
Map
of
Leyte
Island,
showing
the
relative
positions
of
Biliran
and
Maripipi
islands
and
the
locations
of
Leyte
collecting
sites
(as
enumerated
in
text).
A
=
site
of
the
elevational
transect
in
the
viscA—Mt.
Pangasugan
region
(sites
L
I—L6)
shown
in
Figure
2.
Contour
lines
in
meters.
2
FIELDIANA:
ZOOLOGY
o
Maripipi
1
125°
0
Biliran
0
45
0
450
s o
.00
ca
20
19
A
1
6
-11°
17
8-1
1070
11°-
18
15
Leyte
5
21
All-
14
22
45
0
-10°
10
°
-
0
100Km
I
1
125°
1
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
3
tralian
Museum
[wAm]).
In
1984
we
initiated
work
on
Biliran
with
brief
surveys
of
two
sites
from
27
April
to
3
May
(specimens
in
ummz).
During
a
second
field
season
from
10
to
28
April
1987,
we
investigated
two
sites
intensively
and
three
others
incidentally
(specimens
in
PNM,
su,
and
usNm).
Field
work
on
Maripipi
took
place
from
4
to
7
July
1981
(specimens
in
ummz),
from
4
to
8
May
1984
(specimens
in
ummz),
and
from
15
to
27
April
1987
(specimens
in
PNM,
su,
and
usNm).
Geology
The
geological
history
of
the
Philippines
has
been
outlined
in
detail
elsewhere
(Heaney,
1985b,
1986).
Here
we
provide
only
a
general
summary
as
it
applies
to
the
study
region.
The
Philippine
Islands
have
formed
along
sub-
duction
zones
at
the
margins
of
lithospheric
plates,
primarily
as
a
result
of
anticlinal
folding
and
vol-
canic
activity.
Although
small
amounts
of
conti-
nental
crustal
material
are
present
on
some
is-
lands,
these
lie
beneath
younger
marine
deposits.
Evidence
therefore
indicates
that
the
archipelago
emerged
de
novo
as
an
oceanic
group.
The
island
of
Leyte
consists
primarily
of
Miocene
limestones
and
marine
volcanics,
indicating
that
it
probably
did
not
emerge
before
the
Pliocene.
There
are
also
several
Quaternary
volcanoes
on
Leyte,
and
much
of
the
island
interior
is
covered
by
relatively
young
volcanic
material.
Likewise,
Biliran
and
Maripipi
are
entirely
covered
by
volcanic
material
(Ham-
ilton,
1979;
011ier,
1985;
Mitchell
et
al.,
1986).
The
periodic
advance
and
recession
of
conti-
nental
glaciers
during
the
Pleistocene
epoch
were
accompanied
by
global
changes
in
sea
level.
In
the
Philippine
archipelago,
the
lower
sea
levels
asso-
ciated
with
glacial
maxima
resulted
in
larger
island
areas
and
periods
of
connection
between
many
present-day
islands.
With
the
exception
of
the
Pa-
lawan
region,
however,
the
Philippines
have
never
had
a
direct
land
connection
with
mainland
Asia.
During
the
latest
glacial
episode,
about
18,000
years
ago,
sea
level
was
approximately
120
m
lower
than
at
present
(Donn
et
al.,
1962;
Bloom,
1983).
Leyte,
Biliran,
and
Maripipi,
which
are
now
separated
from
one
another
by
relatively
shallow
channels
(<70
m
minimum
water
depth),
were
then
en-
compassed
within
the
single
large
Pleistocene
is-
land
of
Greater
Mindanao
(Heaney,
1985b,
1991a,
b).
At
that
time,
Biliran
and
Maripipi
were
rela-
tively
isolated
volcanic
highlands
in
a
vast
lowland
plain
extending
between
present-day
Leyte
and
Samar
(fig.
1
in
Heaney,
1985b).
Description
of
Study
Areas
Field
sites
visited
by
us
on
each
island
are
listed
and
described
in
detail
below.
Additional
Leyte
sites
visited
by
previous
collectors
were
taken
di-
rectly
from
specimen
labels
or
from
the
literature.
Older
sites
are
identified
by
the
name
of
a
local
barrio
or
principal
geographic
feature
and
the
en-
compassing
municipality.
Although
these
cannot
be
located
precisely,
we
have
included
approxi-
mate
latitude
and
longitude
coordinates
for
each.
All
collecting
localities
are
plotted
by
site
number
on
the
island
maps.
Leyte
The
island
of
Leyte
(fig.
1)
is
centered
at
about
10°50'N,
124°50'E
and
is
located
approximately
50
km
north
of
the
northern-most
point
on
Min-
danao.
It
is
separated
from
Samar
to
the
northeast
by
a
narrow
(2
km)
channel.
Minimum
mid-chan-
nel
water
depth
to
Mindanao
is
80
m,
but
only
ca.
20
m
to
Samar.
The
island
consists
of
a
low
coastal
plain
of
variable
width
and
rugged
interior
moun-
tain
ranges
with
a
maximum
elevation
of
1350
m.
Most
of
the
coastal
plain
has
been
cleared
for
ag-
riculture
(principally
rice,
maize,
and
sugar
cane),
but
small
patches
of
heavily
disturbed
secondary
forest
still
existed
in
some
coastal
areas
in
1987.
Much
of
the
forest
in
the
interior
has
been
de-
stroyed
or
disturbed
through
extensive
logging
and
upland
agriculture.
In
the
area
of
our
field
surveys
on
the
west-central
portion
of
the
island,
undis-
turbed
forest
extended
down
to
about
250
m
el-
evation
in
1987.
At
that
time
we
estimated
that
less
than
10%
of
the
island
retained
primary
or
moderately
disturbed
forest.
Mean
annual
precip-
itation
from
1971
to
1986
at
Baybay
(elev.
ca
.
5
m)
on
the
western
coastal
plain
was
205
cm,
with
minimum
and
maximum
monthly
means
of
7.5
and
24.5
cm
in
May
and
December,
respectively
(Heideman,
1988).
Elsewhere
in
the
central
Phil-
ippines,
mid-elevation
mountain
regions
receive
up
to
three
times
as
much
rainfall
as
adjacent
low-
lands
(Manalo,
1956;
Heideman
&
Erickson,
1987).
Therefore,
mean
annual
precipitation
in
the
in-
terior
highlands
on
Leyte
may
exceed
500
cm.
Precipitation
profiles
on
Biliran
and
Maripipi
are
4
FIELDIANA:
ZOOLOGY
San
Augustin
5
50
0
psngssu
gs
"
River
—10°45'
2
Mf
g
1
VISCA
0
1
2Km
Gabes
(
124
°
50'
FIG.
2.
Map
of
the
elevational
transect
in
the
viscA-Mt.
Pangasugan
region,
Leyte
(sites
L1-L6).
Contour
lines
in
meters.
presumably
similar,
with
some
differences
due
to
the
effect
of
smaller
island
area.
We
conducted
field
work
at
seven
field
sites
on
Leyte.
Most
of
our
work
was
concentrated
along
an
elevational
transect
on
Mt.
Pangasugan
(sum-
mit
elev.
1150
m)
situated
immediately
northeast
of
the
campus
of
the
Visayas
State
College
of
Ag-
riculture
(figs.
1-3).
Along
this
transect,
lowland
dipterocarp
forest
extended
up
to
approximately
500
m
(fig.
4),
followed
by
mixed
lowland
dipter-
ocarp-lower
montane
forest
to
approximately
800
m
(fig.
5),
and
finally
ridgetop
mossy
forest
along
the
mountain
crest
from
850
m
to
the
summit
(fig.
6).
We
established
four
camps
in
primary
forest
at
300,500,700,
and
950
m
elevation
that
served
as
focal
points
for
intensive
survey
work
(figs.
3-
7).
Less
extensive
collecting
was
done
at
the
base
of
Mt.
Pangasugan
in
areas
of
disturbed
forest
and
mixed
agricultural
land
between
50
and
100
m
elevation.
We
also
collected
bats
at
the
Cathedral
Cave
complex
of
limestone
caves
near
Inopacan.
Site
L
1
Visayas
State
College
of
Agriculture,
7
km
N
Baybay,
Baybay
Munic.,
Leyte
Prov.,
5-
10
m
elev.,
10°45'N.
124°47'30"E.
We
did
limited
netting
in
mixed
residential
and
agricultural
areas
on
the
college
campus,
where
there
was
a
great
variety
of
fruit
trees
and
flowering
plants.
We
also
set
a
few
traps
for
commensal
rodents
in
campus
buildings.
Site
L2
—Mt.
Pangasugan,
7
km
N,
1
1
/
2
km
E
Baybay,
Baybay
Munic.,
Leyte
Prov.,
50
m
elev.,
10°45'N,
124°48'E.
We
collected
at
the
base
of
Mt.
Pangasugan
immediately
above
the
campus
of
the
Visayas
State
College
of
Agriculture.
We
netted
bats
at
two
sites
about
400
m
apart
along
a
small
river
in
an
area
of
patchwork
agricultural
land,
second
growth,
and
disturbed
forest.
The
first
site
was
surrounded
by
agricultural
land
planted
with
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
5
A
110
11
W
.
1
-
I:
1
lipi."411re
410
FIG.
3.
Mt.
Pangasugan,
Leyte
(summit
elev.
1150
m).
View
is
northeast
from
the
VISCA
campus
(site
LI,
ca.
10
m
elev.).
Site
L6
(950
m
elev.)
was
located
at
the
summit
of
the
large
subsidiary
peak
north
(left)
of
the
main
peak.
Photograph
taken
in
March
1987
by
E.
A.
Rickart.
coconut
palms
(Cocos
nucifera)
and
abaca
(Musa
textilis),
with
scattered
remnant
forest
trees
and
second
growth.
The
second
site
was
in
dense
sec-
ond
growth
with
scattered
small
patches
of
abaca,
sweet
potato,
and
maize.
A
few
traps
were
set
in
forest
at
the
second
site
(ca.
20
trap-nights).
We
also
hunted
in
this
area.
Site
L3—Mt.
Pangasugan,
10
km
N,
2
km
E
Baybay,
Baybay Munic.,
Leyte
Prov.,
300
m
elev.,
10°46'N,
124°49'E.
This
site
was
near
the
lower
limit
of
remaining
primary
forest
on
Mt.
Panga-
sugan.
It
was
located
on
the
side
of
a
small
valley
from
ca.
280
m
along
a
small
stream
to
ca.
400
m
along
an
adjacent
ridge.
Emergent
trees
(predom-
inantly
dipterocarp
species)
had
large
buttresses,
diameters
at
breast
height
(dbh)
of
0.8-2.0
m,
and
heights
of
45-55
m.
The
relatively
unbroken
can-
opy
ranged
from
25
to
40
m
high.
Large
woody
vines
up
to
10
cm
in
basal
diameter
were
common.
Epiphytic
ferns
and
orchids
were
uncommon,
and
moss
was
scarce.
An
open
understory
consisted
of
saplings,
small
palms,
tree
ferns
(Cyathea),
erect
terrestrial
pandans
(Pandanus),
and
low
ferns.
There
was
a
thin
covering
of
leaf
litter
over
rocky
volcanic
soil.
Site
IA
—Mt.
Pangasugan,
8
1
/
2
km
N,
2
1
/
2
km
E
Baybay,
Baybay
Munic.,
Leyte
Prov.,
500
m
elev.,
10°45
'30"
N,
124°49'30"E.
This
site
was
located
on
a
ridge
system
bounded
by
steep
valleys
in
undis-
turbed
lowland
forest
(fig.
4).
The
canopy
was
25-
30
m
high
and
broken
by
numerous
tree-falls.
Emergent
trees
with
large
buttresses
had
dbh
of
0.5-1.5
m
and
heights
up
to
40
m.
Woody
vines
were
common.
Epiphytes,
particularly
ferns,
were
common
but
not
abundant.
Moss
was
limited
to
trunks
of
large
trees
and
fallen
logs.
The
understory
was
similar
to
that
at
site
L3
but
more
dense.
Scattered
patches
of
sedges
were
also
present.
There
was
a
moderate
layer
of
leaf
litter
and
humus
(up
to
20
cm
deep)
over
rocky
soil.
Site
L5—Mt.
Pangasugan,
10
1
/
2
km
N,
4
km
E
Baybay,
Baybay
Munic.,
Leyte
Prov.,
700
m
elev.,
10°46'N,
124°49'30"E.
This
site
was
located
along
a
narrow
ridge
bounded
by
cliffs
and
steep
slopes.
Vegetation
was
transitional
lowland/montane
for-
est
and
included
some
mossy
forest
elements
(fig.
6
FIELDIANA:
ZOOLOGY
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.
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t
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.
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.
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IP
ke.
all
11,
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e
,
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7
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-
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te'r4°
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7.1
art
...
Ai,
ti
0
411
11
"
7
'0*
••,
4t
4-
Jr"
Fro.
4.
Lowland
dipterocarp
forest
at
ca.
550
m
elevation
on
Mt.
Pangasugan,
Leyte
(near
site
L4).
Rattus
everetti
was
the
most
common
small
mammal
trapped
in
this
habitat,
followed
by
Apomys
littoralis
and
Bullimus
bagobus.
The
fruit
bats
Ptenochirus
minor,
P.
jagori,
and
Rousettus
amplexicaudatus
were
common,
as
was
the
microchiropteran
Rhinolophus
inops.
Photograph
taken
in
March
1987
by
P.
D.
Heideman.
5).
The
broken
canopy
was
at
15-20
m.
Emergent
trees
with
little
or
no
buttressing
had
dbh
of
up
to
1
m
and
heights
up
to
30
m.
Dipterocarps
were
uncommon,
and
an
arborescent
gymnosperm
was
present.
No
trees
had
leaves
with
serrated
edges.
Woody
vines
and
rattan
(Calamus)
were
common,
and
climbing
pandans
(Freycinetia)
were
present.
Epiphytes
were
common,
especially
moss,
orchids,
and
ferns.
Pitcher
plants
(Nepenthes)
were
rare.
Understory
and
ground
cover
were
dense,
con-
sisting
of
the
same
general
elements
as
at
lower
sites
with
the
addition
of
sedges
and
low
viny
pan-
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
7
k
"--.11IfEk•O'
f
.14
111111
1
4
MINIMBE-,
1r
4
4;
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ars
4,14
,
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4
,
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t
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for
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IL
e.'21
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:
'
tit
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it
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it.
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c
k*
y
ir
lt
i
71
f.
k
v.
Jt.
1
,4
•.4
4
.
111
Or'
FIG.
5.
Transitional
lowland-montane
forest
at
ca.
700
m
elevation
on
Mt.
Pangasugan,
Leyte
(near
site
L5).
Batomys
salomonseni
was
the
most
common
species
trapped
here,
followed
by
Rattus
everetti
and
Apomys
littoralis.
Ptenochirus
minor
was
the
most
abundant
bat,
followed
by
Harpyionycteris
whiteheadi,
Haplonycteris
ftscheri,
and
Rhinolophus
inops.
Photograph
taken
in
March
1987
by
P.
D.
Heideman.
dans.
Leaf
litter
and
humus
were
thick,
often
in
excess
of
0.5
m.
Site
L6
—Mt.
Pangasugan,
10
km
N,
4
1
/
2
km
E
Baybay,
Baybay
Munic.,
Leyte
Prov.,
950
m
elev.,
10°47'N,
124°50'E.
This
site
was
located
on
a
high,
narrow
ridge
along
the
summit
crest
of
Mt.
Pan-
gasugan
(fig.
6).
The
ridge
varied
in
width
from
5
to
20
m,
with
narrow
points
bounded
by
steep
slopes
or
sheer
cliffs.
Vegetation
was
ridgetop
mossy
forest
characterized
by
the
predominance
of
trees
in
the
families
Fagaceae
and
Ericaceae
and
the
absence
of
dipterocarps.
The
open
canopy
was
5-
8
FIELDIANA:
ZOOLOGY
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s-
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. .
fi
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•=.
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,
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.
i i
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.
r.
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.,
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-
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,
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rt,
14
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r
••
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YL
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-
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FIG.
6.
Ridgetop
mossy
forest
at
930
m
elevation
on
Mt.
Pangasugan,
Leyte
(near
site
L6).
Batomys
salomonseni,
Apomys
littoralis,
and
Crocidura
beatus
were
most
abundant
in
this
habitat.
Ptenochirus
minor
and
Rhinolophus
inops
were
the
only
bat
species
that
were
relatively
common
at
this
elevation.
Photograph
taken
in
March
1987
by
P.
D.
Heideman.
12
m
high.
The
largest
trees
had
dbh
of
10-30
cm
and
were
not
buttressed.
Climbing
pandans
were
abundant,
as
were
epiphytic
pitcher
plants
and
orchids.
Moss
was
abundant
on
nearly
all
surfaces.
The
ground
was
densely
covered
by
viny
pandans,
moss,
sedges,
and
other
herbaceous
plants.
Leaf
litter
and
humus
were
very
thick,
exceeding
0.5
m
in
many
places.
Site
L7
—Cathedral
Cave
area,
4
km
S,
I
kin
E
Inopacan,
Hindang
Munic.,
Leyte
Prov.,
60
m
elev.,
10°28'N,
124°45E.
We
surveyed
caves
in
1984
and
1987
that
lie
along
a
limestone
cliff
face
lo-
cated
about
4
km
south
of
Inopacan.
We
inves-
tigated
one
large
cave
and
a
few
of
the
many
small
caves
at
this
site.
Vegetation
uphill
from
the
caves
was
heavily
disturbed
secondary
forest.
Areas
downslope
were
predominantly
agricultural
land
with
scattered
small
patches
of
second
growth.
This
is
the
same
area
in
which
D.
Empesso
collected
in
1963.
Judging
from
the
specimens
that
he
ob-
tained
(in
the
Royal
Ontario
Museum
[ROM]),
he
collected
both
within
the
caves
and
in
surrounding
forest
and
agricultural
areas.
Undoubtedly,
local
forests
were
much
more
extensive
and
less
dis-
turbed
at
that
time.
The
largest
cave
at
this
locality,
locally
known
as
Cathedral
Cave,
had
a
main
entrance
at
the
center
of
a
large
concavity
in
the
limestone
cliff.
Several
shallow
solution
pits
were
located
along
the
face
of
the
concavity
outside
the
cave
entrance.
The
cave
was
about
100
m
deep
with
two
primary
rooms.
One
room
was
about
40
m
wide
and
30
m
long,
varied
from
10
to
30
m
high,
and
had
many
small
grottoes
on
the
sides.
The
second
room
was
larger
(ca.
50
m
by
60
m)
and
higher
(ca.
40
m),
contained
few
side
passages,
and
was
well
lit
through
a
large
opening
in
the
ceiling.
A
second
site,
located
about
100
m
downhill
from
the
entrance
to
Cathedral
Cave,
consisted
of
cavities
under
large
pieces
of
limestone
that
had
separated
from
the
cliff.
The
resultant
system
of
narrow
passageways
probably
extended
along
much
of
the
cliff
face.
In
1984
we
visited
two
additional
caves
located
about
1
/
2
km
south
of
Cathedral
Cave
along
the
same
hillside.
One
cave
was
moderately
large,
with
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
9
a
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4.?
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t
tI
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r
re
I
t
-
i
e.
I
.t
.
Fio.
7.
View
northeast
from
the
crest
of
Mt.
Pangasugan
(near
site
L6),
showing
extent
of
forest
between
ca.
600
and
1100
m
elevation.
Photograph
taken
in
March
1987
by
P.
D.
Heideman.
a
length
of
25-30
m,
a
width
of
10-20
m,
and
an
average
height
of
5-6
m;
the
ceiling
was
irregular,
with
bats
clustering
in
deep
solution
holes.
The
other
cave
was
about
100
m
away
and
was
25
m
long,
5
m
wide,
and
1-2.5
m
high.
ADDITIONAL
LEYTE
COLLECTING
LOCALITIES—
Site
L8
—Barrio
Buri,
Mt.
Lobi
Range,
Burauen
Munic.,
Leyte
Prov.,
600-850
m
elev.,
ca.
11°00'N,
124°50'E.
Site
L9
—Barrio
Tambis,
Mt.
Lobi
Range,
Bu-
rauen
Munic.,
Leyte
Prov.,
150-700
m
elev.,
ca.
11°00'N,
124°50'E.
The
areas
around
Barrio Buri
and
Barrio
Tambis
in
the
Mt.
Lobi
region
(sites
L8
and
L9)
were
visited
by
D.
S.
Rabor
in
1964
(specimens
in
the
Delaware
Museum
of
Natural
History
[DMNH]).
At
that
time,
areas
below
500
m
elevation
were
a
mixture
of
cultivated
fields,
cleared
hills,
parang
(mixed
second
growth
and
grassland),
and
secondary
forest.
Areas
of
primary
diptero-
carp
forest
were
still
extant
above
500
m
elevation,
with
transitional
montane
forest
at
the
highest
el-
evations
(Parkes,
1973).
Site
L10
—Barrio
Patok,
Mt.
Lobi
Range,
Da-
gami
Munic.,
Leyte
Prov.,
ca.
11°01'N,
124°50'E.
This
was
the
primary
collection
area
of
G.
L.
Al-
casid
and
M.
Celestino
in
1961
(specimens
in
the
American
Museum
of
Natural
History
[AMNH]).
At
that
time,
the
vicinity
of
Patok
was
agricultural
(abaca
and
coconuts).
Forests
persisted
above
300
m
elevation
but
were
broken
in
many
places
up
to
600
m
elevation
by
recently
logged
clearings
(Parkes,
1973).
Site
L11
—Bulog
Pk.,
Mt.
Kabalanti-an,
Ma-
haplag
Munic.,
Leyte
Prov.,
1500-2000
ft
(450-
600
m)
elev.,
ca.
10°36'N,
125°00'E.
Site
L12—Barrio
Paniniklan,
Mt.
Kabalanti-an,
Mahaplag
Munic.,
Leyte
Prov.,
800-1000
ft
(250-
300
m)
elev.,
ca.
10°36'N,
125°00'E.
Site
L13
—Barrio
Santa
Cruz,
Mahaplag
Mu-
nic.,
Leyte
Prov.,
ca.
10°36'N,
125°00'E.
Localities
in
the
vicinity
of
Mt.
Kabalanti-an
in
the
Munic-
ipality
of
Mahaplag
(sites
L11—L13)
were
visited
by
D.
S.
Rabor
in
1964
(specimens
in
DMNH
and
the
University
of
the
Philippines
[uPLB]).
At
that
time,
areas
at
lower
elevations
were
predomi-
nantly
croplands
(corn
and
rice).
Higher
elevations
were
covered
with
mixtures
of
primary
and
sec-
ondary
forest
(Parkes,
1973).
10
FIELDIANA:
ZOOLOGY
100
500
100
900
3
0
5
500
2
1
100
-11°30'
100
0
10Km
124°30'
FIG.
8.
Map
of
Biliran
Island,
showing
locations
of
collecting
sites
(as
enumerated
in
text).
Contour
lines
in
meters.
Site
L14
—Paril
Cave,
Mahaplag
Munic.,
Leyte
Prov.,
ca.
10°36'N,
125°00'E.
An
undescribed
lo-
cality
visited
by
Alcasid
and
Celestino
in
1961
(specimens
in
AMNH).
Site
L15
—Barrio
Balinsasayao,
AhuyogMunic.,
Leyte
Prov.,
10°40'N,
124°57'E.
An
undescribed
locality
visited
by
Alcasid
and
Celestino
in
1961
(specimens
in
AMNH).
Site
L16
—Palo,
Leyte
Prov.,
11°10'N,
125°00E.
An
undescribed
locality,
probably
urban,
for
series
of
Scotophilus
kuhlii
and
Rattus
rattus
(specimens
in
the
Field
Museum
of
Natural
History
[FMNH]).
Site
L17
—Palompon,
Leyte
Prov.,
11°03'N,
124°23'E.
An
undescribed
site,
probably
urban,
for
a
series
of
Scotophilus
(specimens
in
the
ROM:
not
seen).
Site
L18
Vicinity
of
Ormoc,
Leyte
Prov.,
11°01'N,
124°37'E.
Undescribed
localities
in
this
vicinity
are
cited
for
Cynopterus
brachyotis
(spec-
imens
in
RoM)
and
Cervus
mariannus
(specimen
at
vIscA).
Site
L19—
Vicinity
of
Tacloban,
Leyte
Prov.,
11°14'N,
125190/E.
An
undescribed
locality
for
a
specimen
of
Cynocephalus
volans
(specimen
in
USNM).
Site
L20
—Small
island
in
channel
between
Ley-
te
and
Samar,
ca.
11°25'N,
124°55'E.
An
unde-
scribed
locality,
presumably
an
island
roost,
where
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
11
-4,
.4
4
4
4
,
„C
.
,
•••
P
e''‘
1•
.41
fArr.
r.
4
`.;
.Z77
4
`:41
u
Val
ito
,
•••
SOL
_
-
m{,
/
k••'4-
F1G.
9.
View
of
the
southern
slope
of
Mt.
Konduko,
Biliran
(summit
elev.
1045
m)
and
the
adjacent
valley
near
site
B
1
(located
at
the
extreme
left
center
of
the
photograph).
Site
B3
was
located
in
a
saddle
on
the
north
slope
of
Mt.
Konduko
below
the
small
peak
just
visible
behind
the
western
(left)
flank
of
the
mountain.
Lower
elevations
support
a
mosaic
of
agricultural
land,
second
growth,
forest
plantation,
and
small
patches
of
remnant
forest.
Large
forest
tracts
are
restricted
to
slopes
above
800
m
elevation.
Photograph
taken
in
April
1987
by
L.
R.
Heaney.
Steere
(1890)
collected
a
series
ofAcerodonjubatus
(specimens
in
FMNH
and
usNm).
Site
L21
—Consolacion,
Southern
Leyte
Prov.,
10°23'N,
125°00E.
An
undescribed
locality
for
Tarsius
cited
by
Taylor
(1934).
Site
L22—
Vicinity
of
Hinunangan,
Southern
Leyte
Prov.,
ca.
10°24'N,
125°11'E.
An
unde-
scribed
locality
for
Cervus
mariannus
(specimen
at
viscA).
Biliran
This
volcanic
island
(fig.
8),
centered
at
11°35'N,
124°30'E,
is
separated
from
northwestern
Leyte
by
a
sea
channel
1
km
wide.
A
broad
coastal
plain
around
much
of
the
island
perimeter
grades
into
low
hills
and
an
interior
that
is
dominated
by
five
small
ridge
systems.
The
highest
interior
peaks
range
in
elevation
from
1000
to
1340
m.
As
of
1987,
most
of
the
interior
land
below
500
m
el-
evation
had
been
cleared
for
agriculture
(primarily
the
production
of
coconut
and
abaca
hemp),
with
small
remnant
patches
of
heavily
disturbed
forest
restricted
to
the
steepest
ridges.
Between
500
and
800
m,
gentler
slopes
had
been
cleared
and
the
remaining
forest
had
been
extensively
logged
(figs.
9,
10).
Disturbance
due
to
logging
was
less
exten-
sive
above
800
m,
but
large
trees
had
been
selec-
tively
cut
from
all
but
the
steepest
slopes
and
least
accessible
areas;
the
small
areas
of
montane
forest,
where
trees
are
naturally
small,
were
undisturbed.
We
estimated
that
less
than
50
km
2
(less
than
10%
of
the
total
island
area)
remained
covered
by
pri-
mary
or
lightly
disturbed
forest.
Most
of
the
timber
removed
in
recent
years
had
been
cut
illegally
by
loggers
using
handsaws,
with
shaped
timbers
dragged
out
to
roads
by
water
buffalos
(fig.
11).
Most
logging
had
occurred
after
1980,
when
a
bridge
between
Biliran
and
Leyte
was
completed.
Logging
continued
at
a
rapid
rate
during
our
1984
and
1987
field
seasons.
Local
residents
told
us
that
12
FIELDIANA:
ZOOLOGY
40
4
as
-
Mr
F
.•
FIG.
10.
Eastern
slope
of
Mt.
Konduko,
Biliran,
at
ca.
700
m
elevation.
View
is
toward
the
summit
from
a
point
along
the
cleared
ridge
seen
at
the
right
edge
of
Figure
9.
The
foreground
and
middleground
were
recently
cleared
and
burned
and
are
dominated
by
cogon
grass
(I
mperata)
and
scrubby
vegetation
with
scattered
young
coconut
palms.
Higher
slopes
(above
800
m)
remain
forested.
Photograph
taken
in
April
1987
by
L.
R.
Heaney.
the
principal
market
was
a
single
business
in
Na-
val,
which
provided
trucks
for
transporting
lum-
ber
from
roadsides.
Site
B1
-21
km
N,
11
km
E
Naval
town,
Cai-
biran
Munic.,
450
in
elev.,
11°35'N,
124°30'E.
In
1987
we
netted
(8
net-nights)
and
trapped
(61
trap-
nights)
in a
Bureau
of
Forest
Development
refor-
estation
station
at
400-450
m.
A
20-ha
forest
plan-
tation
and
an
adjacent
20-ha
patch
of
heavily
logged
remnant
primary
forest
formed
an
island
of
trees
separated
from
primary
forest
by
about
0.5
km
of
agricultural
land
and
sparse
second
growth.
The
plantation
was
dominated
by
ipil-ipil
(Leucaena)
and
other
fast-growing
trees
with
a
volunteer
un-
derstory
of
native
trees
and
herbs.
Adjacent
ag-
ricultural
land
included
rice
fields,
coconut
groves,
and
fruit
tree
orchards.
The
patch
of
remnant
for-
est
covered
a
hillside
from
400
to
500
m
elevation.
The
canopy
was
discontinuous
and
the
understory
dense
with
small
trees,
shrubs,
and
vines.
The
few
remaining
large
trees
(dbh
>
15
cm;
height
to
30
m)
were
hollow
or
otherwise
damaged.
Site
B2
-3
1
/
kin
S,
5
1
/
2
kin
W
Caibiran
town,
Caibiran
Munic.,
700
m
elev.,
11°32'N,
124°32'E.
In
1984
we
netted
(ca.
36
net-nights)
and
trapped
(ca.
212
trap-nights)
in
mixed
partially
logged
and
primary
lowland
forest.
Although
the
forest
was
in
the
process
of
being
logged,
trees
of
up
to
50
cm
dbh
were
still
common
(fig.
11).
The
canopy
was
at
15-20
m
and
broken
by
abundant
small
clearings
where
large
trees
had
been
felled.
A
brief
inspection
in
1987
showed
the
area
to
be
covered
by
abaca
and
dense
second
growth
beneath
a
high-
ly
discontinuous
canopy
of
trees
up
to
20
cm
dbh.
Site
B3
—Mt.
Konduko,
5
kin
N,
10
km
E
Naval
town,
Naval
Munic.,
850
in
elev.,
11°36'30"N,
124°29'E.
In
1987
we
netted
(74
net-nights)
and
trapped
(451
trap-nights)
in
montane
forest
cen-
tered
on
a
saddle
along
a
ridge
at
800-950
m
on
the
north
side
of
Mt.
Konduko
(summit
elev.
1045
m).
The
forest
on
the
saddle
received
moisture-
laden
air,
producing
some
characteristics
of
higher
elevation
mossy
forest.
The
canopy
was
12-15
m
in
height
with
numerous
gaps
due
to
tree-falls.
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
13
44'
A
.74
C.
,
,
4
e,
,
-••
f14
1
.
1
111
4t.
'
'
4
1,
Jr
,
irdir
84.
o••
1^,,•.
:=
:
"
Vit-irbr
ir6.
-
.444
,
1.
_
AV:r5,
7
:41:i
14
a v'
4.
40
••'.
t
ip
:40K
FIG.
11.
Area
of
recent
illegal
logging
in
lowland
forest
at
700
m
elevation
on
Biliran
(near
site
B2).
Apomys
littoralis,
Batomys
salomonseni,
Rattus
everetti,
R.
exulans,
and
R.
rattus
were
trapped
in
this
area.
Haplonycteris
fischeri,
Ptenochirus
minor,
P.
jagori,
and
Macroglossus
minimus
were
relatively
common.
Photograph
taken
in
April
1984
by
P.
D.
Heideman.
Most
large
trees
were
stunted
and
broken
due
to
wind
damage.
Ferns
and
orchids
were
common
epiphytes,
and
moss
was
common
on
the
trees.
There
was
a
moderately
dense
understory
of
tree
ferns,
saplings,
small
palms,
erect
terrestrial
pan-
dans,
viny
pandans,
and
wild
banana.
Soil
was
rocky,
with
scattered
deep
pockets
of
humus
and
some
leaf
litter.
Trees
below
the
saddle
were
larger
(20
m
canopy,
emergents
to
25
m,
dbh
up
to
50
cm)
and
supported
similar
but
denser
epiphyte
loads.
Canopy
height
declined
to
3
m
on
nearby
higher
ridges;
epiphytes
and
moss
were
uncom-
mon
in
such
places.
Trees
were
absent
from
the
crests
of
the
highest
knolls
where
viny
pandans,
sedges,
and
low-growing
shrubs
predominated.
Site
B4
4
1
/
2
km
S,
5
km
W
Caibiran
town,
Caibiran
Munic.,
920
m
elev.,
11°32'N,
124°32'E.
In
1984
we
conducted
limited
trapping
(91
trap-
nights)
and
netting
(4-8
net-nights)
in
disturbed
ridgetop
montane
forest
above
site
B2.
Site
B5
—Naval
town,
5
m
elev.,
11°34'N,
124°24'E.
In
1984
and
1987
we
made
incidental
collections
of
bats
in
and
around
residential
build-
ings
in
the
town
of
Naval.
BILIRAN
CAVE
LOCALITIES
—The
caves
that
we
found
were
all
in
igneous
rock,
either
cracks
in
cliff
faces
or
gaps
between
large
boulders
in
breakdown
piles
below
cliffs.
All
were
small
with
narrow
and
mostly
shallow
passages.
We
saw
no
limestone
on
the
island
and
thus
doubt
that
there
are
large
caves.
Site
B6
4
km
N,
9
1
/
2
km
E
Naval
town,
Naval
Munic.,
500
m
elev.,
11°36'N,
124°29'E.
In
1987
we
collected
a
series
of
Emballonura
alecto
from
a
small
rockfall
cave
located
in
a
boulder
field
at
the
base
of
a
steep
granite
cliff
on
the
southwest
flank
of
Mt.
Konduko.
The
cave
had
a
single
large
chamber
measuring
ca.
1
m
by
3
m.
Surrounding
vegetation
was
brushy
second
growth.
Site
B7
—Caves
on
Tincansan
Island,
Kawayan
Munic.,
11°40'30"N,
124°19'30"E.
In
1981
we
col-
lected
bats
at
two
sea
caves
on
Tincansan
Island,
a
small
(ca.
1
/
8
km
2
)
islet
located
about
300
m
off
the
northwest
coast
of
Biliran.
One
consisted
of
a
single
concavity
about
3
m
wide,
4
1
/
2
m
high,
and
14
FIELDIANA:
ZOOLOGY
12
100
A
5
400
A
11
•7
•13
700
—11°
A
6
48'
A4
0
700
14
3
A2
1
400
100
0
2Km
124°20'
10
A
FIG.
12.
Map
of
Maripipi
Island,
showing
locations
of
collecting
sites
(as
enumerated
in
text).
Contour
lines
in
meters.
5
m
deep.
The
second
cave
was
larger,
consisting
Maripipi
of
a
single
passageway
6
m
wide,
4
m
high,
and
ca.
20
m
deep.
This
second
cave
was
exposed
to
This
small,
roughly
circular
volcanic
island
(fig.
surf
and
the
floor
was
covered
with
deep
water.
12)
is
centered
at
1
1°47'N,
124°20'E
immediately
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
15
FiG.
13.
The
island
of
Maripipi
(maximum
elev.
924
m),
view
from
the
southeast.
Photograph
taken
in
April
1987
by
P.
D.
Heideman.
north
of
Biliran.
It
is
separated
from
Biliran
by
a
channel
about
9
km
wide
with
a
minimum
mid-
channel
depth
of
60
m.
The
island
rises
steeply
from
the
sea
to
form
a
single
major
mountain
with
two
high
peaks
(fig.
13).
The
larger
peak
is
formed
by
three
interconnected
ridges
that
rise
to
924
m;
the
subsidiary
peak
is
806
m
high.
The
coastal
bench
is
only
about
1
km
wide
and
has
been
com-
pletely
cleared
and
planted,
primarily
with
coco-
nut
palms.
In
1987
the
lowest
primary
forest
was
at
200
m
on
the
steepest
slopes.
Most
of
the
gentler
slopes
had
been
cleared
to
an
elevation
of
about
500
m.
We
estimated
that
an
area
of
3-4
km
2
(15-
20%
of
the
total
island
area)
was
covered
by
pri-
mary
or
lightly
disturbed
forest
in
1987.
Site
M1-1
km
N,
1
1
/
2
km
W
Maripipi
town,
300-400
m
elev.,
11°47'30"N,
124°20'E.
In
1981
we
netted
(10
net-nights)
and
trapped
(100
trap-
nights)
in
a
steep
valley
along
forest
edge
and
in
disturbed
lowland
forest
(fig.
14).
The
extant
forest
was
restricted
to
steep
slopes,
where
only
the
larg-
est
trees
had
been
removed.
Eighty-eight
trap-
nights
were
in
this
forest,
most
along
the
banks
of
a
dry,
rocky
streambed.
Soil
was
thin
and
rocky.
Twelve
trap-nights
were
in
a
nearby
camote
patch.
Nets
were
placed
in
the
forest
and
on
the
forest
edge.
Site
M2-
1
/
2
km
N,
2
1
/
2
km
W
Maripipi
town,
600-700
m
elev.,
11°47'N,
124°19'E.
In
1984
we
netted
(15
net-nights),
trapped
(137
trap-nights),
and
hunted
in
secondary
forest
and
old
second
growth
on
the
upper
slopes
of
the
800
m
peak.
The
forest
had
been
burned
more
than
30
years
prior
to
our
collecting.
Some
flat
land
had
been
cultivated
at
about
that
time.
The
low
canopy
(5-
10
m)
included
tree
ferns
and
was
broken
by
fre-
quent
gaps.
Trees
had
dbh
of
less
than
20
cm.
The
understory
was
rich
with
ferns
and
herbaceous
plants.
Vines
were
abundant,
but
moss
and
other
epiphytes
were
rare.
Site
M3-
1
/
2
km
N,
2
1
/
2
kin
W
Maripipi
town,
800
m
elev.,
11°47'N,
124°19'E.
In
1984
we
did
limited
collecting
(127
trap-nights,
6
net-nights)
in
a
small
area
of
ridgetop
mossy
forest
above
site
M2
(fig.
15).
The
habitat
was
similar
to
that
of
site
M4.
16
FIELDIANA:
ZOOLOGY
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digtiotio.
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7
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-apt_
...i..--Irr
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a
'
''
''''''
.5-
-
'
'
-.
•s
i
e
.
1
.0
°,
'
NI
''it
..r.
r
.
....;•--
-
i.....
7
..
v.,
,,,,
,-
.1-a.:
.
..,41,..,
'
..
j :
(
e
t
,
•'
P
. -
1,,t
6 1.'
t.-
.
-
7
%
LC-
J.
14.
5
.0
•••••
.5,
-
V
,
rf•
-
it"
,
"-.1t
4
.
V.
,
"'"
••
„,,
to
.
Ask
FIG.
14.
Clearing
in
disturbed
lowland
forest
at
ca.
300
m
on
Maripipi
(site
M1).
Crocidura
beatus
and
Rattus
rattus
were
trapped
in
this
area.
Cynopterus
brachyotis
was
the
most
common
fruit
bat.
Photograph
taken
in
July
1981
by
P.
D.
Heideman.
Site
M4-2
km
N,
3
kin
W
Afaripipi
town,
600-
800
in
elev.,
11°48'N,
124°19'E.
In
1987
we
netted
(49
net-nights)
and
trapped
(611
trap-nights)
in
primary
and
lightly
disturbed
ridgetop
montane
forest
from
600
to
800
m
on
the
ridge
system
of
the
highest
peak
on
the
island.
The
forest
had
a
broken
canopy
of
6-8
m
on
the
ridge
crests
and
a
few
meters
higher
on
ridge
sides.
Trees
of
the
fam-
ilies
Lauraceae
and
Ericaceae
(dbh
up
to
50
cm)
were
predominant.
An
arborescent
gymnosperm
was
also
present,
whereas
dipterocarps
were
ab-
sent.
Most
trees
had
moss
on
their
trunks,
some
of
which
was
up
to
8
cm
thick.
Orchids
and
other
epiphytes
were
abundant,
as
were
rattans,
viny
pandans
(Freycinetia),
and
climbing
bamboo
(Schizostachyum).
Ground
cover
was
dense
with
abundant
ferns,
sedges,
and
pandans.
Soil
was
thin
and
rocky.
In
some
flat,
protected
areas,
emergent
trees
up
to
25
m
with
dbh
of
100
cm
were
present
(although
the
canopy
was
generally
no
higher).
In
these
places
ground
cover
was
slightly
less
dense
and
epiphytes
were
generally
confined
to
emergent
trees.
At
600
m
elevation,
a
small
partial
clearing
created
by
removal
of
understory
growth
was
re-
planted
with
cacao
(Theobroma)
seedlings.
Site
MS
Vicinity
of
Viga,
sea
level
to
200
in
elev.,
11°48'30"N,
124°18'E.
In
1987
we
conduct-
ed
a
small
amount
of
netting
(5
net-nights)
in
and
around
the
town
of
Viga
in
agricultural
areas
ad-
jacent
to
second
growth
and
remnant
forest.
We
also
set
a
few
traps
for
commensal
species
in
build-
ings
(11
trap-nights).
Site
M6—
Ca.
2
kin
N,
3
1
/
2
kin
W
Maripipi
town,
500
m
elev.,
11°48'N,
124°19'E.
In
1987
we
ob-
tained
a
commensal
mouse
(Mus)
that
had
been
hand-caught
at
this
site
(a
farm
building
in
a
clear-
ing
at
the
edge
of
primary
forest).
Site
M7—
Ca.
2
kin
N,
4
kin
W
Maripipi
town,
100-200
in
elev.,
11°48'N,
124°18'E.
In
1987
we
received
some
specimens
that
had
been
caught
in
this
area
of
disturbed
lowland
forest
and
coconut
plantation.
Site
M8-3
1
/
2
km
N,
3
1
/
2
km
W
Maripipi
town,
50
in
elev.,
11°49'N,
124°19'E.
In
1987
we
ob-
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
17
al.
+4",,141V
s
t
-
4
N
l
tel
.
.
11
"..
,
..
AP
0.,
i
••••
''4
1
"
rs
,
:•••••
1
-
e
4
.
TZ
4
ti
;.
Tr
!Le
f
4
)
104
1*
••
,.
t.
ta,
-
V
A
F1G.
15.
Ridgetop
mossy
forest
at
800 m
elevation
on
Maripipi
(site
M3).
Crocidura
beatus
and
Rattus
everetti
were
trapped
at
this
site,
and
the
fruit
bats
Harpyionycteris
whiteheadi,
Pteropus
pumilus,
and
Macroglossus
minimus
were
netted
here.
Photograph
taken
in
April
1984
by
P.
D.
Heideman.
served
and
collected
Pteropus
hypomelanus
at
roosts
in
two
35
m
remnant
forest
trees
in
a
grove
of
15-20
m
coconut
palms
near
the
coast.
The
roost
trees
were
separated
by
300
m
and
were
at
least
500
m
from
the
nearest
disturbed
forest.
Site
M9-1
1
/
2
km
N,
1
1
/
2
km
W
Maripipi
town,
200-400
m
elev.,
11°47'30"N,
124°20'E.
In
1984
and
1987
we
observed
and
collected
Acerodon
ju-
batus
in
an
area
of
mixed
remnant
forest
and
co-
conut
plantation
between
200
and
400
m
eleva-
tion.
Several
separate
roost
trees
occurred
in
an
area
of
ca.
30
ha:
a
35
m
remnant
emergent
sur-
rounded
by
15
m
coconut
palms,
a
20
m
Ficus
located
among
smaller
trees
at
the
edge
of
a
cliff
ca.
500
m
from
the
first
roost,
and
a
1
ha
cluster
of
about
10
emergent
trees
on
the
edge
of
the
same
cliff
300
m
from
the
Ficus.
Site
M10—
Vicinity
of
Maripipi
town,
near
sea
level,
11°47'N,
124°21'E.
In
1981
we
obtained
a
few
specimens
from
buildings
in
the
town
of
Mari-
pipi
and
from
nearby
forest.
MARIPIPI
CAVE
LOCALMES—All
of
the
caves
that
we
found
were
in
igneous
rock.
These
consisted
of
cracks
in
large
rocks
or
cliff
faces,
or
of
gaps
beneath
large
rocks
in
the
breakdown
piles
at
the
bases
of
cliffs.
We
found
nine
such
caves
from
the
edge
of
the
sea
to
800
m
elevation
but
found
bats
only
in
caves
between
sea
level
and
200
m.
All
these
caves
were
small,
with
narrow
passageways,
and
often
with
multiple
entrances.
None
had
ne-
gotiable
passages
deeper
than
10
m.
A
complete
circuit
of
the
island's
coast
revealed
no
limestone;
therefore,
we
doubt
there
are
any
large
caves.
Site
Mll
Ca.
3
km
N,
5
km
W
Maripipi
town,
50
m
elev.,
11°48'N,
124°18'E.
In
1987
we
pur-
chased
some
Emballonura
alecto
that
had
been
collected
by
local
boys
from
a
small
rockfall
cave
at
this
approximate
locality.
Site
M12
4
km
N,
3
1
/
2
km
W
Maripipi
town,
10
m
elev.,
11°49'N,
124°19'E.
In
1987
we
col-
lected
Emballonura
alecto,
Taphozous
melanopo-
gon,
and
Hipposideros
ater
from
a
rockfall
cave
complex
in
an
agricultural
area
on
the
northern
coast
of
the
island.
The
cave
was
located
among
boulders
at
the
base
of
a
30
m
cliff
near
sea
level.
Multiple
openings
led
to
several
large
chambers
up
to
10
m
deep
that
were
connected
by
narrow,
twisting
passageways.
Site
M13
3
km
N,
4
km
W
Maripipi
town,
100-200
m
elev.,
11°48'N,
124°18'30"E.
In
1987
18
FIELDIANA:
ZOOLOGY
we
obtained
a
series
of
Emballonura
alecto
from
a
small
rockfall
cave
at
this
site.
The
cave
was
among
boulders
at
the
base
of
a
cliff
in
an
area
of
mixed
agricultural
land,
second
growth,
and
rem-
nant
forest.
Site
M14
Ca.
1
km
N,
1
/
2
km
W
Maripipi
town,
ca.
200
m
elev.,
11°47'30"N,
124°20'30"E.
In
1981
we
collected
a
series
of
Emballonura
alecto
at
this
site,
located
in
primary
forest
approximately
1
km
east
of
site
M
1.
The
cave
consisted
of
a
single
chamber
approximately
5
m
long
and
3
m
high,
with
one
main
entrance
and
several
smaller
open-
ings.
Site
M15
—Ca.
1
/
2
km
N,
1
1
/
2
km
W
Maripipi
town,
200
m
elev.,
11°47'N,
124°20'E.
In
1984
we
obtained
a
small
series
of
Emballonura
alecto
that
had
been
collected
from
an
undescribed
rockfall
cave
at
this
approximate
location.
Methods
Our
field
studies
were
conducted
using
proce-
dures
that
evolved
over
time.
The
primary
pur-
pose
of
surveys
in
1981
and
1984
was
to
ascertain
species
diversity
on
the
separate
islands.
The
1987
surveys
had
the
additional
aim
of
documenting
elevational
patterns
in
species
richness
and
abun-
dance.
These
later
surveys
involved
greater
stan-
dardization
of
field
methods,
particularly
with
re-
spect
to
baits
used
in
small
mammal
trapping,
placement
of
traps
and
mist
nets,
number
of
nights
that
traps
and
nets
were
run,
and
total
effort
(num-
ber
of
trap-
and
net-nights)
at
a
given
locality
(Hea-
ney
et
al.,
1989).
Murid
rodents
and
shrews
were
caught
in
trap
lines
containing
a
mixture
of
Victor
rat
traps
(ca.
65%),
National
live
traps
(25%),
and
Sherman
live
traps
(10%)
spaced
5-15
m
apart
with
one
trap
per
station.
Most
traps
were
set
on
the
ground
(90%),
most
often
along
runways,
near
holes,
or
within
root
tangles.
Other
traps
were
set
aboveground
on
fallen
trees,
horizontal
branches,
or
large
vines.
Separate
trap
lines
of
20-50
traps
typically
were
operated
for
4-6
days;
a
few
were
run
longer
to
increase
captures
of
trap-shy
species.
Traps
were
baited
in
the
late
afternoon
with
fresh-fried
co-
conut
and
peanut
butter
(occasionally
with
addi-
tion
of
small
amounts
of
other
ingredients
such
as
essential
oils
and
banana)
and
checked
and
re-
baited
near
dawn.
Occasionally
traps
were
also
checked
in
the
early
afternoon
and
evening.
Al-
ternative
trapping
methods
(e.g.,
snares,
drift
fenc-
es,
pitfalls)
that
may
be
more
effective
in
capturing
certain
species
were
used
on
a
few
occasions.
We
did
not
use
live
earthworms
as
bait,
although
they
subsequently
have
proven
to
be
very
effective
in
capturing
some
species
that
are
marginally
attract-
ed
to
standard
baits
(Rickart
et
al.,
1991).
Bats
were
caught
in
mist
nets
set
on
ridgetops,
across
established
trails
and
streams,
or
at
the
edge
of
clearings
in
forests,
and
among
fruit
trees
in
agricultural
areas.
Most
nets
were
2.6
m
high
and
12
m
long
and
had
36
mm
mesh
size,
but
smaller
nets
were
used
occasionally.
In
all
years
our
mist-
netting
efforts
included
data
on
the
number
of
in-
dividuals
of
each
species
caught
per
net
each
night.
In
1987
netting
was
more
standardized
with
re-
spect
to
the
number
ofnet-nights
per
site
(generally
more
than
20)
and
number
of
nights
each
net
was
operated
(three
to
six
nights).
On
most
nights,
mist
nets
were
tended
continuously
during
the
primary
activity
peak
in
the
early
evening
(ca.
1800-2000
hours),
although
some
were
tended
longer
or
checked
only
twice
per
evening.
Thereafter,
nets
remained
open
all
night
and
were
checked
at
dawn.
On
each
island
we
used
additional
means
to
census
large
mammals
and
species
that
are
oth-
erwise
difficult
to
trap
or
net.
These
efforts
in-
volved
direct
observation,
hunting
(air
rifle,
.22
caliber
rifle,
or
.410
caliber
shotgun),
limited
pur-
chase
of
old
trophy
specimens
from
local
residents,
and
interviews
with
local
residents.
Trapping
and
netting
sites
were
well
separated
to
minimize
de-
pletion
of
local
populations
and
to
ensure
that
sampling
at
one
site
did
not
influence
sampling
at
another.
Some
animals
were
released
after
positive
identification;
others
were
preserved
as
voucher
specimens
for
taxonomic
and
anatomical
studies.
Each
of
our
field
crews
consisted
of
at
least
one
of
the
authors
and
one
previously
trained
field
tech-
nician
(usually
two
of
each),
two
or
more
local
assistants,
and
often
collaborators
from
other
in-
stitutions.
On
Leyte,
two
crews
usually
worked
different
sites
simultaneously.
To
provide
comparative
data
on
geographic
variation
and
as
an
aid
to
identification,
we
have
attempted
to
examine
and
measure
as
many
spec-
imens
as
possible.
These
are
enumerated
by
lo-
cality
and
institution
under
Specimens
Examined
in
individual
species
accounts.
Cranial
measure-
ments
were
made
by
Heaney
to
the
nearest
0.1
mm
using
dial
or
digital
calipers.
Limits
of
cranial
measurements
are
defined
by
DeBlase
and
Martin
(1974).
External
measurements
were
taken
from
specimen
labels
or
collectors'
field
notes.
Speci-
mens
referred
to
are
housed
in
the
American
Mu-
seum
of
Natural
History
(AMNH),
the
Delaware
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
19
TABLE
1.
The
number
of
insectivores
and
rodents
trapped
at
principal
sites
on
Leyte,
Biliran,
and
Maripipi
islands.
The
number
of
captures
per
100
trap-nights
are
given
in
parentheses.
Species
Leyte
Biliran
Maripipi
L3
(300
m)
IA
(500
m)
L5
(700
m)
L6
(950
m)
BI
(450
m)
B2
B3
(700
m)
(850
m)
B4
(920
m)
MI
M2
M4
(400
m)
(700
m)
(740
m)
Crocidura
beatus
0
2
0*
6
0
0
1
0
1
(0.24)
(0.41)
(0.22)
(1)
Apomys
littoralis
2
7
6
16
0
4
4
8
0 0
0
(0.22)
(0.85)
(0.53)
(1.10)
(1.89) (0.89)
(8.80)
Batomys
salomonseni
0
1
14
21
0
2
1
0
0
0
0
(0.12)
(1.24)
(1.45)
(0.94)
(0.22)
Bullimus
bagobus
6
2
0
0
0 0
0 0
0
0
5
(0.65)
(0.24)
(0.82)
Rattus
everetti
9
11
8
4
0
4
11
1
0
1
8
(0.97)
(1.34)
(0.71)
(0.28)
(1.89)
(2.44)
(1.10)
(0.73)
(1.31)
Rattus
exulanst
0 0
0
0 0
2
0 0
0
0
0
(0.94)
Rattus
rattust
0
0
0
0
4
3
4
3
2
5 5
(6.56)
(1.42)
(0.89)
(3.30)
(2)
(3.65)
(0.82)
Total
captures
17
23
28
47
4
15
21 12
3
6
18
Total
trap-nights
930
819
1125
1451
61
212
451
91
100
137
611
Total
mammals
per
1.83
2.81
2.49
3.24
6.56
7.08
4.66
13.19
3
4.38
2.94
100
trap-nights
Number
of
species
3
5
3
(+
1)*
4
1
5
5
3
2 2
3
*
Inferred
from
presence
at
lower
and
higher
elevations.
t
Introduced
commensal
species.
Museum
of
Natural
History
(DMNH),
the
Field
Museum
of
Natural
History
(11vINH),
Philippine
National
Museum
(PNM),
the
Royal
Ontario
Mu-
seum
(Rom),
the
Silliman
University
Museum
(su),
the
United
States
National
Museum
of
Natural
History
(usNM),
the
University
of
Michigan,
Mu-
seum
of
Zoology
(uMMz),
the
University
of
the
Philippines
at
Los
Banos
(UPLB),
Visayas
State
College
of
Agriculture
(viscA),
and
the
Western
Australian
Museum
(wAM).
Voucher
specimens
collected
by
us
are
deposited
at
PNM,
UMMZ,
USNM,
VISCA,
and
WAM.
Taxa
are
presented
alphabetically
within
fami-
lies.
Unless
noted
otherwise,
species
names
follow
Honacki
et
al.
(1982).
Subspecies
names
are
given
where
they
have
been
used
consistently
and
un-
ambiguously
in
recent
taxonomic
literature.
For
each
species,
we
summarize
the
known
distribu-
tional
limits
(based
on
Heaney
et
al.,
1987,
unless
stated
otherwise)
and
report
known
records
of
oc-
currence
on
the
three
islands.
Specific
collecting
localities
are
referred
to
by
site
number.
We
pro-
vide
tables
of
external
and
cranial
measurements
and
discuss
discernable
patterns
of
geographic
TABLE
2.
Means
SD)
and
ranges
of
selected
measurements
of
adult
Crocidura
beatus
from
Leyte,
Biliran,
and
Maripipi
islands.
Locality
Sex
N
Condylo-
basal
length
Braincase
width
Inter-
orbital
width
Rostral
length
Rostral
width
Post-
palatal
depth
Post-
palatal
length
Occipital
condyle
to
glenoid
fossa
Leyte
d
2
21.6
9.6
4.7
8.6
2.6
3.9
9.6
8.6
(21.6-21.7)
(9.3-9.8)
(4.6-4.8)
(2.5-2.7)
(9.4-9.8) (8.5-8.6)
2
4
21.1±0.38
9.5±0.19
4.7
±
0.31
8.4±0.20
2.8
±0.24
3.9
±
0.17
9.4
±0.13
8.3
±
0.13
(20.6-21.5)
(9.4-9.8)
(4.3-5.0)
(8.3-8.7)
(2.5-3.0)
(3.7-4.1)
(9.3-9.6)
(8.1-8.4)
Biliran
2
1
9.5
3.7
9.8
8.7
Maripipi
9
2
21.0
9.4
4.8
8.5
2.6
3.7
9.4
8.2
(20.6-21.5)
(9.2-9.5)
(4.8-4.9)
(2.6-2.7)
(9.3-9.7)
(8.0-8.5)
Note:
Measurements
other
than
weight
are
in
millimeters.
20
FIELDIANA:
ZOOLOGY
variation
within
the
study
region.
Finally,
we
sum-
marize
available
information
on
ecology
and
be-
havior
of
species,
relying
primarily
on
our
field
notes.
Accounts
of
Species
Order
Insectivora
Family
Soricidae
-Shrews
Crocidura
beatus
Miller,
1910
Prior
to
our
work,
this
species
of
white-toothed
shrew
was
reported
from
Mindanao
only
(Heaney
et
al.,
1987;
Heaney
&
Ruedi,
in
press).
On
Mt.
Pangasugan,
Leyte,
shrews
were
present
in
pri-
mary
lowland
forest
at
500
m
(site
L4)
and
com-
mon
in
mossy
forest
at
950
m
elevation
(site
L6).
A
single
specimen
from
Biliran
was
trapped
in
primary
montane
forest
at
850
m
elevation
(site
B3).
On
Maripipi,
one
specimen
was
caught
near
a
dry
streambed
in
disturbed
lowland
forest
at
385
m
(site
M
1)
and
another
was
taken
in
ridgetop
mossy
forest
at
800
m
elevation
(site
M3).
The
species
has
a
broad
elevational
range
but
is
most
common
in
high-elevation
forest
(table
1).
All
of
the
specimens
we
collected
were
trapped
on
the
ground
within
leaf
litter,
in
runways,
or
along
steep,
moss-covered
embankments.
Three
specimens
were
captured
during
full
daylight.
A
female
cap-
tured
on
20
April
on
Biliran
was
carrying
a
single
embryo
(crown-rump
length
=
3
mm).
Cranial
and
external
measurements
of
adult
specimens
from
the
three
islands
are
listed
in
Table
2.
SPECIMENS
EXAMINED-Total
13.
LEYTE:
Leyte
Prov.:
site
L4
(2
usNm);
site
L6
(6
usNm);
site
L9
(2
DMNI-I).
BILIRAN:
site
B3
(1
usNm).
MARI-
PIPI:
site
M1
(1
ummz);
site
M4
(1
usNm).
Suncus
murinus
occultidens
(Hollister,
1913)
The
widespread
Oriental
house
shrew
occurs
throughout
the
Philippines,
primarily
in
the
low-
lands
near
human
habitations.
The
two
known
specimens
from
Leyte
were
hand-caught
soon
after
sunrise
near
buildings
on
the
campus
of
the
Vi-
sayas
State
College
of
Agriculture
(site
LI).
None
were
taken
on
Biliran
or
Maripipi,
but
we
did
not
make
extensive
efforts
to
obtain
commensal
spe-
cies
on
either
island.
SPECIMENS
EXAMINED-Total
2.
LEYTE:
Leyte
Prov.:
site
LI
(1
UMMZ,
1
usNm).
Order
Dermoptera
Family
Cynocephalidae
-"Flying
Lemurs"
Cynocephalus
volans
(Linnaeus,
1758)
The
kagwang,
or
Philippine
"flying
lemur,"
is
restricted
to
the
Mindanao
faunal
region;
previous
records
are
from
Basilan,
Bohol,
Dinagat,
Leyte,
Mindanao,
Samar,
and
Siargao.
Thomas
(1911)
reported
the
species
to
be
common
on
Leyte
but
did
not
list
specific
localities.
In
the
Mt.
Panga-
sugan
region,
our
field
party
obtained
two
speci-
mens
from
local
hunters.
Both
were
captured
in
an
area
of
secondary
forest
adjacent
to
a
small
coconut
plantation
at
about
50
m
elevation
(site
L2).
In
the
same
area,
we
sighted
individual
ani-
mals
on
consecutive
evenings
shortly
after
dusk
as
they
glided
between
coconut
palms.
Another
specimen
was
taken
on
Mt.
Pangasugan
in
primary
forest
at
500
m
elevation
(site
L4).
Several
other
individuals
were
sighted
at
sites
below
500
m
in
TABLE
2.
Extended.
Fourth
First
incisor
premolar
Labial
width
Palatal
width
to
third
to
third
at
second
at
third
Total
molar
molar
molars
molars
length
Tail
length
Hindfoot
Weight
(g)
9.5
5.4
6.4
2.4
145
59
16
11.2
(9.4-9.6)
(5.3-5.4)
(2.3-2.4)
(53-65)
(11.0-11.5)
9.2
±
0.24
5.2
±
0.12
6.4
±0.05
2.6±0.10
151±13.6
62±11.1
15
±
1.2
11±1.4
(9.0-9.5)
(5.1-5.4)
(6.3-6.4)
(2.5-2.7) (135-163)
(52-75)
(14-16)
(10-12)
5.3
6.3
2.3
144
56
16
11
9.3
5.4
6.5
2.7
150
68
16
9.0
(9.1-9.5)
(5.2-5.5)
(6.4-6.6)
(2.6-2.8) (146-153)
(68-69)
(15-17)
(8.5-9.5)
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
21
TABLE
3.
Selected
measurements
of
adult
Cynocephalus
volans
from
Leyte
and
Biliran
islands.
Condylo-
Inter-
Post-
Post-
basal
Zygomatic
orbital
orbital
Mastoid
Rostral
Orbital
orbital
Sex
N
length
breadth
width
width
breadth
length
length
length
Leyte*
9
Bilirant
1
1
1
66.0
41.
-
8
16.
-
7
15.0
30.2
25.3
24.2
25.7
Note:
Measurements
other
than
weight
are
in
millimeters.
*
No
cranial
measurements.
t
No
external
measurements.
both
primary
and
secondary
forest.
A
specimen
from
the
vicinity
of
Tacloban
(site
L19)
was
ob-
tained
from
Bureau
of
Forest
Development
per-
sonnel.
This
animal,
an
unweaned
juvenile
male,
was
caught
with
an
adult
female
on
18
March.
On
Biliran,
an
adult
male
(scrotal
testes
17
x
14
mm)
was
shot
at
450
m
elevation
in
an
isolated
20
ha
patch
of
remnant
lowland
primary
forest
approx-
imately
1
/
2
km
from
larger
areas
of
forest
(site
B1).
No
kagwang
were
taken
on
Maripipi,
but
local
residents
reported
that
they
were
common.
Shortly
after
dawn
on
7
July
1981,
Heideman
saw
two
kagwang
gliding
between
trees
in
partially
cleared
lowland
forest
at
300
m
elevation
(site
M1).
Ac-
cording
to
local
farmers
and
hunters
on
Leyte,
the
species
is
common
in
both
disturbed
and
primary
forest
habitats
in
lowland
areas
below
500
m
el-
evation,
is
active
both
night
and
day,
and
typically
feeds
on
young
leaves
and
fruit
of
a
variety
of
tree
species.
Specimens
from
Leyte
and
Biliran
(table
3)
fall
within
the
range
of
size
variation
seen
in
series
from
Dinagat,
Siargao,
and
Mindanao
(Hea-
ney
&
Rabor,
1982).
SPECIMENS
EXAMINED-Total
10.
LEYTE:
Leyte
TABLE
4.
Means
SD)
and
ranges
of
selected
measurements
of
adult
fruit
bats
(Acerodon
and
Cynopterus)
from
Leyte,
Biliran,
and
Maripipi
islands.
Sex
N
Condylo-
basal
length
Zygomatic
breadth
Inter-
orbital
width
Post-
orbital
width
Mastoid
breadth
Rostral
length
Orbital
length
C
to
last
M
Acerodon
jubatus
Leyte
3
80.7
43.8
12.5
9.1
24.4
29.1
30.6
32.3
(79.9-82.2)
(42.9-45.1)
(12.0-13.1)
(8.9-9.3)
(23.9-25.0)
(27.9-30.2)
(29.8-31.6)
(31.7-33.0)
9
8
80.7±
1.41
42.3
±0.58
12.4
±
0.77
9.3
±
0.44
24.4
±
0.95
29.1±
1.40
30.7±0.50
32.6
±
0.77
(79.1-83.1)
(41.2-43.7)
(11.6-13.6)
(8.8-10.1)
(23.2-25.7)
(27.5-31.2)
(30.0-31.6)
(30.9-33.3)
Maripipi
3
81.0
44.8
11.9
8.8
25.0
28.8
30.5
33.2
(78.3-85.9)
(41.3-48.3)
(11.1-13.1)
(8.3-9.4)
(24.0-26.3)
(26.9-31.7)
(29.3-32.3)
(31.8-35.7)
9
4
78.1±1.70
42.1±1.37
11.8±0.19
9.0±0.64
24.1±0.57
27.9
±
0.85
29.8
±
0.86
31.3±0.81
(76.7-80.3)
(40.1-43.2)
(11.7-12.1)
(8.4-9.9)
(23.3-24.5)
(27.2-29.1)
(29.1-30.9)
(30.2-31.9)
Cynopterus
brachyotis
Leyte
d
6
28.2±0.54
18.6±0.50
6.0±0.38
6.2±0.53
12.0±0.22
9.0±0.26
11.8±0.26
9.4±0.35
(27.4-28.8)
(17.9-19.2)
(5.4-6.5)
(5.5-6.8)
(11.6-12.2)
(8.6-9.3)
(11.4-12.1)
(9.0-9.9)
9
7
27.8±0.51
18.4±0.44
6.0±0.31
6.3±0.29
11.8±0.49
8.5
±
0.35
12.1±0.39
9.4±0.15
(27.1-28.4)
(18.0-19.1)
(5.5-6.4)
(5.9-6.6)
(11.0-12.6)
(7.9-8.9)
(11.6-12.7)
(9.2-9.6)
Biliran
d
4
28.4±0.34
19.1±0.38
6.1±0.06
6.3±0.32
12.0±0.21
9.1±0.30
12.1±0.49
9.3±0.18
(28.1-28.8)
(18.7-19.4)
(6.0-6.1)
(6.0-6.7)
(11.7-12.2)
(8.8-9.5)
(11.5-12.7)
(9.1-9.5)
9
10
27.3±0.59
18.3±0.43
6.1
±0.50
6.3±0.48
11.7±0.37
8.7±0.44
11.7±0.37
9.1±0.26
(26.5-28.4)
(17.6-19.1)
(5.5-7.0)
(5.7-7.4)
(11.0-12.1)
(8.0-9.5)
(11.2-12.3)
(8.7-9.7)
Maripipi
d
7
27.4±0.82
18.1±0.35
6.1
±0.25
6.5
±
0.54
11.9±0.31
8.7±0.24
11.8±0.46
9.3±0.21
(26.2-28.2)
(17.6-18.7)
(5.6-6.3)
(5.3-7.0)
(11.5-12.3)
(8.4-9.1)
(11.0-12.4)
(9.0-9.6)
9
8
27.2±0.79
18.4±0.36
6.1
±0.23
6.3±0.29
11.9±0.31
8.7
±
0.42
11.8±0.33
9.1
±0.22
(26.4-28.5)
(17.7-18.9)
(5.6-6.4)
(5.9-6.8)
(11.4-12.4)
(8.1-9.2)
(11.1-12.3)
(8.7-9.4)
Note:
Measurements
other
than
weight
are
in
millimeters.
*
Measured
from
dry
skins.
22
FIELDIANA:
ZOOLOGY
Palatal
C
to
Molariform
breadth
Palatal
Total
Tail
last
M
toothrow
at
last
NI
length
length
length
Hindfoot
Ear
Weight
(g)
275
81
29
1,360
645
260
86
29
1,160
31.6
20.7
14.2
32.1
-
_
Prov.:
site
L2
(2
usNm);
site
L4
(1
ummz);
site
L10
(5
AMNH);
site
L19
(1
usNm).
BILIRAN:
site
B1
(1
us/gm).
Order
Chiroptera
Family
Pteropodidae
-Fruit
Bats
Acerodon
jubatus
(Eschscholtz,
1831)
The
golden-capped
flying
fox
is
a
Philippine
en-
demic
that
is
found
throughout
the
country
with
the
exception
of
the
Palawan
region.
Gunther
(1879)
reported
this
species
from
southern
Leyte.
Specimens
taken
by
Steere
(1890)
on
a
small
island
midway
in
the
channel
between
Leyte
and
Samar
(site
L20)
were
named
Pteropus
aurinuchalis
(El-
liot,
1896),
but
they
are
not
now
recognized
as
distinct.
We
did
not
encounter
this
species
during
our
field
work
on
Leyte,
nor
did
we
obtain
any
specimens
on
Biliran.
We
received
reports
of
flying
fox
roosts
on
Biliran
but
did
not
locate
any.
On
several
occasions,
however,
we
saw
giant
fruit
bats
flying
at
dusk
over
our
campsites
on
Biliran
or
TABLE
4.
Extended.
TABLE
3.
Extended.
Molariform
toothrow
Palatal
breadth
at
last
NI
Palatal
length
Total
length
Tail
length
Hindfoot
Ear
Forearm
Weight
(g)
23.4
(22.9-23.8)
23.6
±0.68
(22.6-24.7)
16.4
(16.1-16.9)
16.4
±0.33
(15.8-16.8)
47.0
(46.6-47.4)
45.6±
1.59
(43.2-47.5)
-
-
0
0
65*
66±2.9*
(65-70)
28•
(28-29)
27±2.5*
(23-30)
203*
(198-208)
192±5.8*
(184-202)
-
-
23.7
16.2
46.2
298
0
66
39
193
954
(22.6-24.8)
(14.4-17.9)
(44.0-49.9)
(280-319)
(60-73) (37-41)
(187-203)
(750-1,140)
22.8
±0.70
16.6
±
0.85
44.5±
1.11
286±
10.1
0
65±5.2
36±0.6
192±5.1
948±
170
(22.1-23.8)
(15.9-17.7)
(43.1-45.8)
(273-296)
(60-70)
(36-37)
(186-196)
(720-1,100)
6.6
±0.23
6.2
±0.29
14.1±
0.35
101
±8.1
9±3.3
15±0.8
18±0.4
64±1.5
30±2.3
(6.4-6.9)
(5.7-6.5)
(13.6-14.5)
(96-117)
(6-15)
(14-16)
(18-19)
(62-66)
(26-32)
6.6±0.08
6.2±0.28
14.0±0.46
101±3.3
8±2.3
15±1.0
18±0.8
64±2.1
30±1.7
(6.5-6.7)
(5.8-6.6)
(13.4-14.6)
(96-105)
(4-10)
(14-16)
(17-19)
(61-67)
(28-32)
6.5±0.14
6.3±0.14
14.5±0.50
100±5.1
8±1.2
16±1.3
18±0.6
64±2.1
32±2.1
(6.4-6.7)
(6.1-6.4)
(14.1-15.2)
(93-104)
(7-9)
(15-18)
(18-19)
(61-66)
(30-34)
6.5±0.23
6.1
±
0.22
13.8±0.38
99
±5.1
8±0.6
16±1.7
18±0.8
64±2.1
33±2.7
(6.1-6.9)
(5.7-6.3)
(13.4-14.6)
(88-105)
(7-9)
(13-18)
(17-19)
(61-66)
(28-38)
6.5±0.21
6.0±0.34
14.0±0.48
97±2.9
8±1.9
14±2.0
17±1.6
62±2.2
31±1.8
(6.3-6.9)
(5.5-6.5)
(13.2-14.6)
(93-101)
(5-10)
(10-16)
(14-19)
(59-65)
(28-33)
6.5±0.15
6.2±0.27
13.9±0.45
97±3.8
6±2.3
14±0.6
17±0.5
62±
1.1
32±2.8
(6.2-6.7)
(5.8-6.6)
(13.2-14.6)
(91-103)
(3-10)
(13-15)
(17-18)
(61-64)
(29-38)
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
23
TABLE
5.
The
number
of
small
fruit
bats
(Pteropodidae)
netted
at
principal
sites
on
Leyte,
Biliran,
and
Maripipi
islands.
The
number
of
captures
per
net-night
are
given
in
parentheses.
Species
Leyte
Biliran
Maripipi
BI
(450
m)
B2
B3
(700
m)
(850
m)
B4
(920
m)
L2
(50
m)
L3
(300
m)
LA
L5
L6
(500
m)
(700
m)
(950
m)
MI
M2
M4
(400
m)
(700
m)
(740
m)
Cynopterus
39
2
4
3
0
17
6
6
0
14
11
32
brachyotis
(1.56)
(0.03)
(0.05) (0.02)
(2.13)
(0.24)
(0.08)
(1.40)
(0.73)
(0.65)
Eonycteris
0
0
0 0 0
0
1
0 0 0
1
0
robusta
(0.04)
(0.07)
Eonycteris
28
0*
2
0 0
2
1
0 0
3
0
0
spelaea
(1.12)
(0.03)
(0.25)
(0.04)
(0.30)
Haplonycteris
0
2
1
11
3
1
26
43
3
0
0
0
fzscheri
(0.03)
(0.01)
(0.09)
(0.09)
(0.13)
(1.04)
(0.58)
Harpyionycteris
0
1
6
12
0
0 0
Ot
1
Ot
Ot
2
whiteheadi
(0.01)
(0.08)
(0.10)
(0.04)
Macroglossus
20
4
3
4
1
5
10
9
4
4
1
15
minimus
(0.80)
(0.05)
(0.04) (0.03)
(0.03)
(0.63)
(0.40)
(0.12)
(0.40)
(0.07)
(0.31)
Ptenochirus
40
14
18
2
0
16
27
2
3
10
6
12
jagori
(1.60)
(0.18)
(0.23)
(0.02)
(2.00)
(1.08)
(0.03)
(1.00)
(0.40)
(0.24)
Ptenochirus
12
23
21
84
11
0
24
51
10
0
0
0
minor
(0.48)
(0.30)
(0.27)
(0.68)
(0.33)
(0.96)
(0.69)
Rousettus
13
0*
10
3
0
63
0 0
0
1
11
2
amplexicaudatus
(0.52)
(0.13)
(0.02)
(7.80)
(0.10)
(0.73)
(0.04)
Total
captures
152
46
65
119
15
104
95
111
21
32
30
63
Total
net-nights
25
77
79
123
33
8
25
74
4-8
10
15
49
Bats/net-night
6.08
0.60
0.82
0.97
0.45
13.0
3.80
1.50
3.20
2.00
1.29
Number
of
species
6 6
(+
2)*
8
7
3
6
8
6
5
6
6
5
*
Inferred
from
presence
at
lower
and
higher
elevations.
t
Known
to
be
present
(from
vocalizations)
but not
captured.
flying
toward
Biliran
from
Maripipi.
Given
the
proximity
of
large
colonies
on
Maripipi,
we
be-
lieve
these
bats
were
Acerodon
jubatus.
On
Maripi-
pi,
we
collected
10
individuals
of
this
species
from
roost
trees
in
an
area
of
mixed
remnant
forest
and
coconut
plantation
(site
M9).
In
1987
each
of
about
12
roost
trees
at
this
site
had
groups
of
50-150
bats
(apparently
all
A.
jubatus);
we
estimated
a
total
colony
size
of
ca.
1,000.
Individuals
appeared
to
be
evenly
spaced
within
the
tree
crowns.
When
one
bat
approached
another,
the
usual
result
was
an
aggressive
display
involving
vocal
threats
and
"boxing"
with
closed
wings
(striking
with
the
wrists).
Within
a
short
time,
one
bat
usually
re-
treated
or
flew
from
the
tree,
often
returning
to
another
position
within
the
same
tree.
Many
bats
hung
with
their
wings
partly
spread,
occasionally
flapping
them
gently,
and
sometimes
licking
the
wing
membranes
and
panting.
This
behavior
probably
prevented
overheating.
Two
adult
fe-
males
collected
in
May
1984
and
another
taken
on
26
April
1987
held
large
embryos
(crown-rump
lengths
=
75-106
mm;
99-148
g).
In
April
1987
the
abdomens
of
most
females
were
swollen
with
near-term
embryos,
but
no
females
were
carrying
nursing
young.
A
young
female
taken
on
this
date
(believed
to
be
a
yearling
on
the
basis
of
external
and
cranial
measurements)
was
not
pregnant.
These
observations
suggest
that
A.
jubatus
on
Maripipi
produce
their
single
yearly
young
in
late
May
and
June,
and
that
females
may
not
reproduce
until
2
years
of
age.
Leyte
and
Maripipi
specimens
exhibit
slight
sexual
dimorphism,
with
males
averaging
slightly
larger
in
most
cranial
and
external
mea-
surements
(table
4).
SPECIMENS
EXAMINED-Total
21.
LEYTE:
Leyte
Prov.:
site
L20
(2
FMNH,
9
usNm).
MARIPIPI:
site
M9
(6
ummz,
4
usNm).
Cynopterus
brachyotis
luzoniensis
Peters,
1862
The
short-nosed
fruit
bat
is
common
through-
out
the
Philippines
and
is
widely
distributed
in
24
FIELDIANA:
ZOOLOGY
Southeast
Asia
(Koopman,
1989).
It
typically
is
associated
with
forest
clearings
and
other
dis-
turbed
habitats
(Heaney
et
al.,
1989;
Heideman
&
Heaney,
1989).
In
the
Mt.
Pangasugan
region
on
Leyte,
this
was
the
most
common
pteropodid
spe-
cies
taken
in
disturbed
secondary
forest
at
50
m
(site
L2)
and
in
mixed
agricultural
and
residential
areas
below
50
m
elevation
(site
L1).
It
was
un-
common
in
primary
lowland
and
montane
forest
between
300
and
700
m
elevation
(sites
L3—L5)
and
absent
from
ridgetop
mossy
forest
(site
L6;
table
5).
Similarly,
on
Biliran
it
was
abundant
in
disturbed
lowland
forest
and
second
growth
re-
forested
areas
(site
B1)
but
rarely
encountered
in
primary
forest
(site
B3;
table
5).
On
Maripipi,
in
contrast
to
the
situation
on
all
other
islands
where
we
have
worked
(Heaney
et
al.,
1989,
1991),
C.
brachyotis
was
abundant
in
all
habitats
and
was
the
most
frequently
captured
bat
species
in
pri-
mary
mossy
forest
at
750
m
(site
M4;
table
5).
On
Maripipi,
we
found
one
individual
roosting
in
a
3
m
shrub
amid
moderately
dense
undergrowth
in
primary
forest
at
800
m
elevation.
Adult
fe-
males
taken
from
late
March
through
early
July
contained
single
embryos.
Cranial
and
external
measurements
(table
4)
fall
within
ranges
reported
for
specimens
from
various
locations
throughout
the
Philippines
(Heaney
&
Rabor,
1982;
Heaney,
1984;
Heaney
et
al.,
1991).
Males
are
slightly
larg-
er
than
females
in
some
cranial
dimensions.
Leyte
specimens
have
a
karyotype
of
2n
=
34,
FN
=
58,
that
is
indistinguishable
from
those
of
specimens
from
elsewhere
in
the
species'
geographic
range
(Rickart
et
al.,
1989a).
Kitchener
and
Mahara-
datunkamsi
(1991)
have
suggested
that
C.
b.
lu-
zoniensis
be
recognized
as
a
separate
species,
but
we
recommend
further
investigation
ofgeographic
variation
before
this
is
accepted.
SPECIMENS
EXAMINED—Total
134.
LEYTE:
Leyte
Prov.:
site
LI
(10
usNm);
site
L2
(2
PNM,
33
usNm);
site
L3
(2
usNm);
site
L4
(2
ummz,
4
usNM);
site
L5
(3
usNm).
BILIRAN:
site
B1
(2
PNM,
15
usNm);
site
B2
(8
ummz);
site
B3
(6
usNm).
MARI-
PIPI:
site
M1
(14
ummz);
site
M2
(4
ummz);
site
M4
(2
PNM,
27
USNM).
OTHER
RECORDS—LEYTE:
site
L7
(RoM);
site
L18
(RoM).
Eonycteris
robusta
Miller,
1913
This
nectar
bat
is
a
widespread
Philippine
en-
demic
with
records
from
Catanduanes,
Luzon,
Mindanao,
Negros,
and
Siargao
Islands
(Heaney
&
Rabor,
1982;
Heideman
&
Heaney,
1989;
Hea-
ney
et
al.,
1991).
Some
authors
have
considered
it
to
be
a
subspecies
of
E.
major
(Honacki
et
al.,
1982;
Koopman,
1989).
Eonycteris
robusta
is
not
common
and
may
rely
on
primary
forest
habitat.
We
captured
none
during
the
course
of
our
field
work
on
Leyte,
but
specimens
were
taken
at
Tam-
bis
(site
L9),
Paniniklan
(site
L12),
and
Santa
Cruz
(L13)
in
Leyte
Province
during
the
1960s.
On
Bi-
liran
in
April
1984,
we
netted
a
68
g
adult
female
carrying
a
13
g
neonate
in
partially
logged
primary
forest
at
700
m
(site
B2).
On
Maripipi,
an
adult
male
was
netted
at
600
m
on
a
cleared
ridgetop
above
old
second
growth
forest
(site
M2).
Mea-
surements
of
specimens
from
all
three
islands
are
larger
than
those
of
series
from
Luzon
and
Catan-
duanes
(Heaney
&
Rabor,
1982;
Heaney
et
al.,
1991).
In
the
large
series
from
Leyte,
males
are
substantially
larger
than
females
(table
6).
SPECIMENS
EXAMINED—Total
20.
LEYTE:
Leyte
Prov.:
L9
(1
DMNH);
L
12
(1
DMNH);
L13(15
DMNH).
BILIRAN:
site
B2
(2
ummz).
MARIPIPI:
site
M2
(1
UMMZ).
Eonycteris
spelaea
glandifera
Lawrence,
1939
The
cave
nectar
bat
occurs
from
India
to
Timor
and
is
common
throughout
the
Philippines.
On
Leyte
in
1987,
we
found
large
numbers
of
this
species
within
solution
cavities
in
the
ceiling
of
Cathedral
Cave
(site
L7).
Several
separate
groups
contained
between
200
and
500
individuals,
with
a
total
population
probably
in
excess
of
2,000
in-
dividuals.
The
species
has
been
heavily
persecuted
at
this
site:
a
large
pile
of
dried
skins
and
charred
bones
representing
several
hundred
individuals
was
found
at
the
cave
entrance,
and
we
found
evidence
of
torches
and
fires
within
the
cave.
Nectar
bats
were
common
in
mixed
second
growth
and
agri-
cultural
areas
at
the
base
of
Mt.
Pangasugan
but
rare
in
primary
forest
at
500
m
on
the
mountain
(sites
L2
and
L4;
table
5).
On
Biliran,
two
speci-
mens
were
netted
in
mixed
agricultural
land
and
replanted
forest
at
400
m
(site
B1),
and
one
was
taken
in
partially
logged
primary
forest
at
700
m
(site
B2).
On
Maripipi,
three
specimens
were
cap-
tured
at
the
edge
of
disturbed
forest
at
300
m
(site
M
1).
On
Negros,
Eonycteris
spelaea
is
associated
with
forest
clearings
and
other
disturbed
habitats
(Heideman
&
Heaney,
1989).
Based
on
our
rela-
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
25
TABLE
6.
Means
SD)
and
ranges
of
selected
measurements
of
adult
fruit
bats
(Eonycteris
and
Haplonycteris)
from
Leyte,
Biliran,
and
Maripipi
islands.
Sex
N
Condylo-
basal
length
Zygomatic
breadth
Inter-
orbital
width
Post-
orbital
width
Mastoid
breadth
Rostral
length
Orbital
length
C
to
last
M
Eonycteris
rohusta
Leyte
d
9
6
11
36.2±2.11
(32.7-37.9)
33.2±
1.04
(32.1-35.0)
23.1±0.58
(22.5-23.8)
20.0±0.74
(19.1-21.0)
7.4±0.59
(6.6-8.1)
7.1
±0.27
(6.6-7.6)
7.7±0.61
(7.4-8.6)
8.2
±0.51
(7.1-8.9)
14.7
±
0.66
(13.6-15.3)
13.5±0.36
(13.0-14.1)
13.8
±
1.23
(11.9-14.7)
12.6±0.95
(11.6-14.1)
12.8
±0.62
(12.1-13.6)
12.4
±
0.61
(11.7-13.9)
13.1±0.96
(11.8-14.0)
12.1±0.40
(11.6-12.9)
Biliran
9
1
35.9
20.8
7.1
7.9
14.2
13.9
13.1
13.4
Maripipi
d
1
35.4
22.0
6.9
7.2
14.1
13.3
12.7
12.8
Eonycteris
spelaea
Leyte
d
6
34.3±0.76
21.7±
1.11
6.9
±
0.18
7.7±0.37
14.4
±0.61
12.9±0.35
12.5±0.35
12.1
±0.56
(33.2-35.2)
(19.7-23.0)
(6.6-7.1)
(7.2-8.3)
(13.6-15.3)
(12.3-13.4)
(12.1-13.1)
(11.6-13.2)
9
8
33.3±0.75
20.0
±0.43
6.7±0.12
7.5
±
0.42
13.4±0.36
12.2±0.54
12.3±0.29
12.1
±0.56
(32.1-34.3) (19.3-20.7)
(6.5-6.9)
(7.0-8.2)
(12.8-14.0)
(11.2-12.7)
(11.9-12.7)
(11.3-13.3)
Biliran
d
1
33.7
22.6
7.0
7.9
14.9
12.6
12.8
11.9
Maripipi
9
2
33.2
19.4
6.6
7.5
13.0
12.4
12.6
12.2
Haplonycteris
fischeri
(33.0-33.3)
(19.1-19.8)
(7.4-7.6)
(12.8-13.1)
(11.6-13.3)
(12.5-12.6)
(12.0-12.3)
Leyte
d
8
23.9
±0.64
16.5
±0.40
6.3±0.29
6.0
±
0.28
11.0
±
0.26
7.8
±0.55
10.1
±
0.22
8.6±0.29
(23.1-24.8)
(15.8-17.1)
(5.8-6.8)
(5.7-6.6)
(10.7-11.4)
(7.1-8.5)
(9.8-10.4)
(8.2-8.9)
9
8
23.9
±0.60
16.5
±0.42
6.0±0.42
5.9±0.14
10.8±0.18
7.7±0.34
10.1
±
0.33
8.4±0.33
(22.8-24.8)
(15.9-17.3)
(5.3-6.7)
(5.7-6.1)
(10.6-11.0)
(7.2-8.1)
(9.8-10.6)
(7.7-8.6)
Biliran
d
8
24.3
±0.58
16.4±0.51
6.3±0.14
6.2
±0.29
10.9±0.40
7.8±0.26
10.3±0.48
8.5±0.31
(23.3-24.8)
(15.7-17.3)
(6.0-6.4)
(5.5-6.4)
(10.4-11.6)
(7.5-8.2)
(9.7-10.9)
(8.1-8.9)
9
10
24.3±0.73
16.4±0.42
6.2±0.31
6.1±0.33
10.8
±0.31
7.7±0.43
10.3±0.39
8.4±0.31
(22.9-24.8)
(15.7-17.3)
(5.4-6.4)
(5.6-6.7)
(10.4-11.5)
(6.8-8.4)
(9.8-11.0)
(7.8-9.1)
Note:
Measurements
other
than
weight
are
in
millimeters.
tive
netting
success,
the
species
is
most
common
in
lowland
agricultural
areas
and
is
rare
or
absent
in
primary
forest
(table
5;
Heaney
et
al.,
1989).
It
is
most
abundant
in
the
vicinity
oflimestone
caves
that
serve
as
primary
roost
sites.
Pregnant
females
containing
single
embryos
(crown-rump
lengths
=
4-33
mm)
were
taken
during
April.
Lactating
fe-
males
with
dependent
young
were
taken
in
March
at
Cathedral
Cave
on
Leyte.
Cranial
measure-
ments
of
males
are
slightly
larger
than
those
of
females
(table
6).
Measurements
fall
within
the
ranges
of
specimens
from
Catanduanes,
Luzon,
and
Siargao
(Heaney
&
Rabor,
1982;
Heaney
et
al.,
1991).
Karyotypes
of
specimens
from
Leyte
are
2n
=
36,
FN
=
66,
and
are
indistinguishable
from
those
reported
for
specimens
taken
elsewhere
in
the
species'
distribution
(Rickart
et
al.,
1989a).
SPECIMENS
EXAMINED-Total
80.
LEYTE:
Leyte
Prov.:
site
L2
(1
su,
6
usNM);
site
L4
(2
usNm);
site
L7
(37
DMNH,
2
PNM,
17
UMMZ,
9
usNm).
BI-
LIRAN:
site
B1
(2
PNM);
site
B2
(1
ummz).
MARI-
PIPI:
site
M1
(3
ummz).
Haplonycteris
fischeri
Lawrence,
1939
Fischer's
pygmy
fruit
bat
is
a
Philippine
endem-
ic
that
occurs
throughout
the
archipelago
with
the
probable
exception
of
the
Palawan
faunal
region.
It
is
found
almost
exclusively
in
primary
forest
and
lightly
disturbed
forest
and
is
typically
most
abundant
at
middle
elevations
(Heaney
et
al.,
1989;
Heideman
&
Heaney,
1989).
On
Leyte,
we
netted
specimens
between
300
m
and
950
m
on
Mt.
Pan-
gasugan.
It
was
uncommon
in
lowland
forest
at
300
and
500
m
(sites
L3
and
L4)
and
common
in
lower
montane
forest
and
ridgetop
mossy
forest
at
700
and
950
m
elevation
(sites
L5
and
L6;
table
5).
On
Biliran,
Haplonycteris
was
abundant
in
pri-
mary
montane
forest
at
850
m
(site
B3),
less
corn-
26
FIELDIANA:
ZOOLOGY
TABLE
6.
Extended.
Palatal
Molariform
breadth
Palatal
Total
Tail
toothrow•
at
last
M
length
length
length
Hindfoot
Ear
Forearm
Weight
(g)
9.1
±0.70
(8.1-9.9)
8.6±0.30
(8.1-9.1)
7.8±0.29
(7.5-8.3)
7.4
±
0.20
(7.1-7.7)
19.7±0.57
(19.2-20.5)
18.3
±
0.58
(17.4-19.2)
-
-
-
-
-
-
-
-
9.3
7.9
19.5
143
18
23
24
74
73
9.0
7.6
19.0
151
27
21
22
82
68
8.6±0.38
7.5±0.23
18.7±0.44
135±8.4
14
±
2.1
20±
1.2
21±
1.7
74
±2.8
69±12.8
(8.2-9.2)
(7.1-7.7)
(17.9-19.1)
(124-145)
(13-18)
(18-21)
(19-23)
(71-79)
(57-90)
8.5±0.46
7.6
±
0.30
18.3
±
0.71
128
±5.0
15
±
2.1
20
±0.9
21±
1.1
74
±2.6
58
±8.9
(7.8-9.3)
(7.2-8.2)
(16.8-18.9)
(121-137)
(12-18)
(19-21)
(19-22)
(70-77)
(43-70)
8.8
8.6
7.7
7.4
18.1
18.3
137
125
16
14
19
18
21
22
73
73
72
51
(8.5-8.7)
(7.3-7.4)
(18.1-18.5)
(122-128)
(13-16)
(18-19)
(21-22)
6.0
±
0.18
5.1
±0.22
11.6±
0.51
73
±
3.9
0
12±
1.0
14
±0.6
48±0.8
18±1.0
(5.7-6.2)
(4.9-5.6)
(10.7-12.2)
(70-80)
(11-13)
(13-14)
(47-49)
(17-19)
5.9±0.18
5.1
±0.18
11.5
±0.33
70±
1.5
0
11±0.5 14±0.5
49±0.8
18
±1.3
(5.5-6.0)
(4.9-5.3)
(11.0-12.0)
(68-71)
(11-12)
(13-14)
(48-50)
(16-19)
5.9
±0.24
5.2
±0.34
11.8±0.44
73
±3.3
0
13±
1.0
14±
1.0
49±1.6
18±1.4
(5.5-6.2)
(4.8-5.8)
(11.0-12.3)
(68-77)
(12-14)
(12-15)
(47-52)
(17-21)
5.8
±0.22
5.4
±
0.33
12.0±0.49
74
±
2.9
0
13±1.2
14
±0.7
50±2.0
19±1.6
(5.4-6.3)
(4.5-5.8)
(10.9-12.6)
(68-79)
(11-15)
(13-15)
(44-53)
(16-21)
mon
in
lightly
disturbed
lowland
forest
at
700
m
(site
B2),
and
rare
in
mixed
forest
plantation
and
agricultural
land
at
400
m
(site
Bl;
table
5).
De-
spite
considerable
netting
effort
in
forest
habitat
that
was
apparently
excellent
for
them
(65
net-
nights;
sites
M1,
M3,
and
M4),
we
took
no
Hap-
lonycteris
on
Maripipi
and
doubt
that
they
occur
there.
Females
collected
from
mid-March
to
late
April
contained
single
embryos
(crown-rump
lengths
=
3-4
mm)
in
arrested
early
development
(Heideman,
1988,
1989).
Large
series
of
speci-
mens
from
Leyte
and
Biliran
show
no
indication
of
sexual
size
dimorphism
(table
6).
Leyte
speci-
mens
are
similar
in
body
size
to
those
from
Dina-
gat,
Catanduanes,
and
Luzon
(Heaney
&
Rabor,
1982;
Heaney
et
al.,
1991),
whereas
those
from
Biliran
are
slightly
larger.
Specimens
from
Leyte
and
Biliran
have
karyotypes
of
2n
=
58,
FN
'
•=1
66,
possessing
the
highest
known
diploid
number
for
the
suborder
Megachiroptera
(Rickart
et
al.,
1989a).
SPECIMENS
EXAMINED-Total
94.
LEYTE:
Leyte
Prov.:
site
L3
(2
usNm);
site
L4
(3
ummz,
1
USNM,
1
VISCA);
site
L5
(1
PNM,
2
UMMZ,
10
usNm);
site
L6
(3
usNm);
site
L12
(7
DMNH).
BILIRAN:
site
B
1
(2
usNm);
site
B2
(26
UMMZ);
site
B3
(2
PNM,
1
su,
19
USNM,
1
VISCA,
4
w
Am);
site
B4
(9
UMMZ).
Harpyionycteris
whiteheadi
whiteheadi
Thomas,
1896
The
harpy
fruit
bat
is
endemic
to
the
Philip-
pines,
with
prior
records
from
Camiguin,
Min-
danao,
Mindoro,
and
Negros
islands
(Peterson
&
Fenton,
1970;
see
also
Koopman,
1989,
who
con-
sidered
H.
celebensis
of
Sulawesi
to
be
conspecific).
On
Negros,
it
probably
is
restricted
to
primary
forest
and
very
lightly
disturbed
forest
and
is
most
common
at
middle
elevations
(Heaney
et
al.,
1989;
Heideman
&
Heaney,
1989).
On
Mt.
Pangasugan,
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
27
TABLE
7.
Means
SD)
and
ranges
of
selected
measurements
of
adult
fruit
bats
(Harpyionycteris
and
Macroglossus)
from
Leyte,
Biliran,
and
Maripipi
islands.
Sex
N
Condylo-
basal
length
Zygomatic
breadth
Inter-
orbital
width
Post-
orbital
width
Mastoid
breadth
Rostra!
length
Orbital
length
C
to
last
M
Ilarpyionycteris
whiteheadi
Leyte
d
7
41.0±0.56
23.3±0.46
6.4±0.45
5.7±0.56
15.4±0.41
11.4±0.23
17.4±0.29
15.9±0.28
(40.3-41.7)
(22.9-24.1)
(5.6-6.9)
(5.2-6.7)
(14.8-15.8)
(11.0-11.7)
(17.1-17.9)
(15.6-16.3)
9
3
40.6
23.3
6.5
5.6
15.3
11.1
17.3
15.6
(39.7-41.5) (22.5-24.2)
(6.1-6.9)
(5.6-5.7)
(15.0-15.6)
(11.0-11.1)
(16.1-18.1)
(15.5-15.6)
Macroglossus
minimus
Leyte
6
3
25.0
15.3
5.0
7.1
10.2
9.7
8.2
8.7
(24.5-25.8)
(14.5-16.2)
(4.8-5.3)
(6.8-7.4)
(9.9-10.8)
(9.3-10.4)
(7.9-8.4)
(8.2-9.4)
9
7
25.0±0.63
14.0±0.21
4.9±0.27
7.2±0.17
10.0±0.22
9.5±0.35
8.0±0.17
8.6±0.38
(23.8-25.7)
(13.8-14.4)
(4.4-5.2)
(7.0-7.4)
(9.6-10.2)
(9.1-10.0)
(7.8-8.3)
(8.1-9.0)
Biiran
6
7
25.1
±0.38
15.1
±0.34
4.9
±0.24
6.9±0.35
10.1
±0.42
9.8
±
0.29
8.1
±0.17
8.9
±0.36
(24.6-25.7)
(14.7-15.6)
(4.4-5.1)
(6.4-7.3)
(9.5-10.7)
(9.5-10.2)
(7.9-8.4) (8.5-9.6)
9
5
25.0±0.72
13.9±0.50
4.6±0.30
7.3±0.33
10.0±0.25
9.6±0.38
8.0±0.34
8.7±0.33
(23.9-25.7)
(13.4-14.6)
(4.3-4.9)
(7.0-7.8)
(9.7-10.4)
(9.0-10.0)
(7.7-8.6) (8.3-9.4)
Maripipi
d
4
24.9
±
0.78
15.2
±
0.68
4.8
±0.25
7.0±0.25
10.5
±
0.25
9.5
±
0.34
8.1
±0.22
8.6
±
0.33
(24.1-25.6)
(14.4-15.9)
(4.5-5.1)
(6.7-7.3)
(10.1-10.7)
(9.0-9.8)
(7.8-8.3)
(8.2-9.0)
9
3
25.1
14.4
4.7
7.0
10.1
9.4
8.1
8.3
(24.7-25.4)
(4.6-4.8)
(6.5-7.5)
(9.6-10.4)
(9.4-9.5)
(7.8-8.3)
(7.5-8.7)
Note:
Measurements
other
than
weight
are
in
millimeters.
Leyte,
it
was
confined
to
undisturbed
primary
for-
est,
where
it
was
rare
at
300
m
(site
L3)
and
mod-
erately
common
at
500
m
and
700
m
elevation
(sites
L4
and
L5;
table
5).
On
Biliran,
we
netted
one
specimen
in
partially
logged
primary
montane
forest
at
920
m
elevation
(site
B4).
We
also
heard
the
distinctive
whistling
flight
calls
of
this
species
in
primary
forest
at
elevations
from
600
to
950
m
elevation
(site
B3).
On
Maripipi,
we
captured
two
specimens
in
primary
mossy
forest
at
800
m
ele-
vation
(site
M3).
Pregnant
females
captured
be-
tween
20
March
and
8
May
contained
single
em-
bryos
(crown-rump
lengths
=
8-50
mm).
Four
lactating
females
taken
between
18
and
30
March
had
recently
given
birth
to
single
offspring.
Three
of
these,
weighing
110, 110,
and
125
g,
were
car-
rying
nursing
young
weighing
26,
21,
and
21
g,
respectively.
Cranial
and
external
measurements
of
males
average
slightly
larger
than
those
of
fe-
males
(table
7).
This
species
has
a
distinctive
karyotype
of
2n
=
36,
FN
=
58
(Rickart
et
al.,
1989a).
SPECIMENS
EXAMINED-Total
29.
LEYTE:
Leyte
Prov.:
site
L3
(1
usNm);
site
L4
(4
ummz,
6
USNM,
1
vISCA);
site
L5
(1
PNM,
11
USNM);
site
L
11
(1
UPLB);
site
L12
(1
UPLB).
BILIRAN:
site
B4
(1
ummz).
MARIPIPI:
site
M3
(2
ummz).
Macroglossus
minimus
lagochilus
(Matschie,
1899)
The
dagger-toothed
flower
bat
occurs
from
Thailand
to
Australia
and
is
found
throughout
the
Philippines.
On
Leyte,
we
netted
this
species
in
primary
lowland,
montane,
and
mossy
forest
at
elevations
ranging
from
300
to
950
m
and
in
dis-
turbed
lowland
forest
between
50
and
100
m
el-
evation
(table
5).
It
was
found
at
all
forest
and
second
growth
sites
on
Biliran
and
Maripipi
(table
5).
Although
we
encountered
it
at
virtually
all
of
our
netting
sites,
M.
minimus
is
most
common
in
disturbed
habitats
(Heaney
et
al.,
1989;
Heideman
&
Heaney,
1989).
Most
specimens
were
netted
within
25
m
of
wild
or
domestic
bananas
and
ab-
aca
(Musa
spp.),
whose
flowers
represent
an
im-
portant
food
source
for
this
nectarivorous
species.
We
collected
females
pregnant
with
single
embry-
os
from
late
March
to
early
July.
On
average,
males
in
our
series
are
slightly
larger
than
females
in
most
cranial
and
external
dimensions
(table
7).
Size
ranges
are
comparable
to
those
seen
for
series
from
several
other
Philippine
islands
(Heaney
&
Rabor,
1982;
Heaney,
1984;
Heaney
et
al.,
1991).
The
species
has
a
standard
karyotype
of
2n
=
34,
FN
=
62,
that
is
identical
to
those
from
elsewhere
in
28
FIELDIANA:
ZOOLOGY
TABLE
7.
Extended.
Molariform
toothrow
Palatal
breadth
at
last
M
Palatal
length
total
length
"tail
length
I
lindfoot
Ear
Forearm
Weight
(g)
12.0±0.29
8.0
±
0.11
20.9
±0.29
139±5.3
0
23
±
1.5
22±1.2
84
±
2.7
100±5.6
(11.6-12.3)
(7.4-8.2)
(20.6-21.3)
(130-145)
(21-25)
(20-24)
(80-87)
(88-104)
11.8
7.6
20.6
139
0
23
21
82
110
(11.8-11.9)
(7.4-7.8)
(20.4-20.8)
(133-143)
(22-23)
(20-22)
(81-83)
5.3
5.2
13.4
74
0
12
16
44
18
(5.1-5.7)
(5.2-5.3)
(13.2-13.7)
(70-78)
(11-13)
(16-17)
(42-45)
(16-20)
5.3±0.22
5.1
±0.14
13.7±0.35
73±
1.8
0
12
±
0.8
16±
1.2
42±
1.4
17±
1.7
(5.0-5.6)
(5.0-5.4)
(13.1-14.1)
(70-76)
(11-13)
(14-17)
(40-44)
(16-20)
5.4
±
0.29
5.0±0.22
13.8
±
0.28
71
±2.6
0
13±
1.0
16
±0.6
42±0.5
17±
1.0
(5.0-5.7)
(4.8-5.4)
(13.4-14.1)
(67-73)
(12-14)
(15-16)
(42-43) (16-18)
5.3±0.23
5.1
±0.24
13.5
±
0.48
68±2.1
0
12
16
±1.2
41
±
1.2
16
±0.6
(5.1-5.6)
(4.8-5.4)
(13.0-14.1)
(66-70)
(15-17)
(40-42)
(15-16)
5.1±0.19
4.9
±0.13
13.0
±
0.43
68
±
1.9
0
12±0.8
16
±1.0
42±0.6
18±2.0
(5.0-5.4)
(4.8-5.1)
(12.5-13.5)
(67-71)
(11-13)
(15-17)
(41-42)
(16-20)
5.1 5.1
13.4
70
0
13
17
42
18
(5.0-5.1) (5.0-5.3)
(13.4-13.5)
(66-74)
(12-14)
(15-18)
(41-44)
(17-19)
the
species'
distributional
range
(Rickart
et
al.,
1989a).
SPECIMENS
EXAMINED-
Total
87.
LEYTE:
Leyte
Prov.:
site
L2
(3
PNM,
17
USNM);
site
L3
(4
USNM);
site
L4
(1
uMMz,
3
USNM,
1
viscA);
site
L5
(4
uMMz,
4
usNm);
site
L6
(1
USNM).
BILIRAN:
site
B1
(2
PNM,
3
USNM);
site
B2
(10
uMMz);
site
B3
(1
su,
7
USNM,
1
VISCA);
site
B4
(4
uMMz).
MARIPIPI:
site
M
I
(4
ummz);
site
M2
(I
ummz);
site
M3
(I
uMMz);
site
M4
(15
USNM).
OTHER
RECORDS-
LEYTE:
site
L
II
(Rom);
site
L
1
5
(AMNH).
Ptenochirus
jagori
(Peters,
1
86
1)
This
species
is
a
Philippine
endemic,
occurring
throughout
the
archipelago
with
the
exception
of
the
Palawan
region.
It
was
abundant
on
Mt.
Pan-
gasugan
in
disturbed
secondary
forest
at
50
m
el-
evation
(site
L2)
and
in
primary
lowland
forest
at
300
m
and
500
m
(sites
L3
and
L4).
It
was
rare
in
montane
forest
at
700
m
(site
L5)
and
absent
in
mossy
forest
at
950
m
(site
L6;
table
5;
Heaney
et
al.,
1989).
It
was
also
present
on
the
campus
of
the
Visayas
State
College
of
Agriculture
near
areas
of
secondary
forest
(site
L
I).
We
observed
this
species
roosting
in
small
solution
pits
(ca.
20-30
cm
across
and
30-70
cm
deep)
along
the
cliff
face
at
the
entrance
to
Cathedral
Cave
(site
L7).
Bats
were
well
shaded
within
these
cavities
but
were
clearly
visible
from
below.
We
found
eight
groups
in
such
pits,
each
containing
from
I
to
10
indi-
viduals.
Below
each
roost
was
a
pile
of
chewed
and
spat-out
remains
of
fruit,
primarily
figs
(Fi-
cus).
These
piles
of
ejecta
were
large
(1.2-1.6
m
in
diameter
and
up
to
25
cm
thick),
indicating
long-term
use
of
the
roost
sites.
We
also
found
two
small
groups
of
P.
jagori
inside
two
of
the
caves,
both
in
solution
pits
in
well-lit
places
near
the
entrances.
On
Biliran,
this
species
was
abundant
in
partially
logged
lowland
primary
forest
at
700
m
(site
B2)
and
in
the
mixed
agricultural/replanted
forest
area
at
450
m
elevation
(site
B1)
but
un-
common
in
primary
montane
and
mossy
forest
at
850
m
and
920
m
(sites
B3
and
B4;
table
5).
Pte-
nochirus
jagori
was
found
in
all
habitats
sampled
on
Maripipi
(table
5).
We
collected
females
preg-
nant
with
single
embryos
from
early
March
to
ear-
ly
July.
In
our
series,
males
are
consistently
larger
than
females
in
most
cranial
dimensions
(table
8).
There
is
also
substantial
geographic
variation
in
body
size:
specimens
from
Leyte
and
Biliran
are
RICKART
ET
AL.:
MAMMALS
OF
LEYTE,
BILIRAN,
AND
MARIPIPI
ISLANDS
29
TABLE
8.
Means
SD)
and
ranges
of
selected
measurements
of
adult
fruit
bats
(Pie
nochirus)
from
Leyte,
Biliran,
and
Maripipi
islands.
Sex
N
Condylo-
basal
length
Zygomatic
breadth
Inter-
orbital
width
Post-
orbital
width
Mastoid
breadth
Rostra!
length
Orbital
length
C
to
last
M
Ptenochirus
jagori
Leyte
d
11
35.5
±
0.85
24.9
±
0.75
7.2
±
0.46
6.3±0.16
14.5±0.35
10.8
±
0.46
15.9
±
0.62
12.5±0.46
(34.2-36.6)
(23.5-25.8)
(6.5-8.0)
(6.1-6.5)
(14.0-15.1)
(10.2-11.8)
(14.6-16.7)
(12.0-13.2)
9
13
34.7
±
0.80
24.1
±
0.74
7.0
±0.36
6.3±0.36
14.3
±
0.40
10.3
±
0.39
15.8
±
0.56
12.1±0.31
(33.3-36.1)
(23.2-25.8)
(6.5-7.8)
(5.8-7.0)
(13.7-15.1)
(9.6-11.0)
(14.9-16.7)
(11.3-12.4)
Biliran
d
10
35.0
±
0.64
24.3
±
0.76
7.1
±
0.42
6.4
±0.41
14.5
±
0.38
10.6
±
0.35
15.7
±0.43
12.3
±
0.32
(34.1-35.7)
(23.0-25.7)
(6.4-7.8)
(5.8-7.0)
(13.9-15.2)
(10.2-11.2)
(14.9-16.3)
(11.8-12.8)
9
9
34.1
±0.52
24.0
±
0.58
7.0
±
0.28
6.5±0.29
14.4
±
0.37
10.4
±
0.44
15.4
±0.30
11.8
±0.24
(33.2-34.6)
(23.0-24.8)
(6.6-7.5)
(6.0-6.9)
(13.8-14.8)
(9.5-11.0)
(15.0-15.7)
(11.4-12.1)
Maripipi
d
4
36.4
±
0.88
25.3
±
0.83
7.9
±
0.51
6.0±0.49
14.9
±
0.39
11.7
±
0.41
16.0
±0.38
12.4
±0.60
(35.4-37.1)
(24.7-26.5)
(7.1-8.2)
(5.5-6.6)
(14.6-15.4)
(11.3-12.1)
(15.5-16.4)
(12.0-13.3)
9
6
35.2
±
1.06
24.2
±
0.65
7.2±0.32
6.1±0.20
14.5
±
0.18
10.7
±
0.50
15.9
±
0.40
12.2
±
0.21
Ptenochirus
minor
(33.9-36.7)
(23.8-25.4)
(6.9-7.6)
(5.8-6.4)
(14.2-14.7)
(9.9-11.3)
(15.3-16.5)
(12.0-12.5)
Leyte
d
15
29.0
±
0.82
19.6
±
0.84
6.1
±
0.30
6.3±0.38
12.5
±
0.31
9.0
±
0.39
12.5
±
0.31
10.1
±
0.25
(28.1-30.6)
(18.2-20.3)
(5.6-6.7) (5.8-7.2)
(12.0-12.8)
(8.5-9.8)
(11.8-13.3)
(9.5-10.5)
9
17
28.7
±0.83
19.4
±
0.55
6.1±0.32
6.1±0.33
12.5
±
0.29
8.9
±
0.41
12.5
±
0.45
9.9
±0.22
(27.2-30.4)
(18.3-20.4)
(5.3-6.6) (5.5-6.6)
(11.9-12.9)
(8.1-9.6)
(11.5-13.2) (9.5-10.4)
Biliran
d
12
29.2±0.58
19.6
±
0.81
6.2
±
0.34
6.2±0.28
12.3
±0.23
9.1
±0.32
12.6
±0.34