Ba
Biology
and
Conse a ion
Edi ed,
by
Thomas
H.
Kunz
and
Paul
A.
Racey
SMITHSONIAN
INSTITUTION
PRESS
Washing on
and
London
24
Geog aphic
Pa e ns,
Ecological
G adien s,
and
he
Main enance o
T opical
F ui
Ba
Di e si y
The
Philippine
Model
RUTH
C.
B.
UTZURRUM
Like
mos
opical
a eas
wo ldwide,
Philippine
ain o es s
ha e
been
los
o
exploi a ion ( esou ce
ex ac ion
and
con-
e sion),
mono ypic
e o es a ion,
and
eplacemen
o
na-
i e
species
wi h
exo ics.
The
a chipelago's
o es s,
es ima ed
o
co e
80%
o
i s
o al
land
a ea
in
he
1800s,
ha e
been
educed
o
12%
wi h
mos
o
he
decline
occu ing
wi hin
he
las
six
decades
(Mye s
1988;
Kumme
1990).
Gi en
he
a chipelagic
na u e
o
he
Philippines,
an h opogenic
deg a-
da ion
and
agmen a ion
o
he
o es
habi a
could
e y
well
exace ba e
e ec s
o
isola ion
on
popula ions
o
plan s
and
animals.
By
he
la e
1980s,
2
o
he
26
species
o
Philip-
pine
ui
ba s
(Family
P e opodidae)
we e
epo ed
o
be
ex inc
(Ace odon
luci e
and
Dobsonia
chapmani;
Heaney
and
Heideman
1987).
A
ecen
edisco e y
o
a
popula ion
o
A.
luci e
on
Bo acay
Island
(o
Panay)
awai s
con i ma ion
(E.
E.
Ma o,
pe sonal
communica ion;
bu
see
Heaney
e
al.
[n.d.]
ques ioning
he
species
s a us
o
Ace odon
luci e ).
I
is
ecognized
ha
a
leas
5
o
he
24
ex an
species
(including
Ace odon
juba us,
A.
leuco is,
Eonyc e is
obus a,
Nyc imene
abo i,
and
P e opus
leucop e us)
a e
se iously
h ea ened
by
habi a
des uc ion
and
hun ing
(Heaney
and
U zu um
1991;
Micklebu gh
e
al.
1992;
U zu um
1992).
In
his
chap e ,
I
summa ize
esul s
o
ele a ional
an-
sec
in en o ies
conduc ed
in
ecen
yea s
and
use
hese
da a
o
assess
he
e ec s
o
habi a
agmen a ion
on
he
main enance
o
Philippine
ui
ba
di e si y.
Th ee
ques-
ions a e
examined:
(1)
Wha
a e
he
pa e ns
o
species
di e si y
in
un agmen ed
(i.e.,
local
g adien s)
and
na u-
ally
agmen ed
a eas
(i.e.,
biogeog aphic)
landscapes?
(2)
Wha
impac
will
habi a
agmen a ion
ha e
on
he
main enance
o
di e si y?
and
(3)
Do
mac o- and
mic o-
geog aphic
pa e ns
o
communi y
associa ions
p o ide
p ac ical
insigh s
in o
conse a ion?
The
conse a ion
im-
plica ions
o
hese
ques ions
a e
(1)
o
examine
p ope ies
o
popula ions
and
communi y
assemblages
ha
could
de-
ine
hei
esponse
o
e ec s
o
habi a
agmen a ion;
(2)
o
iden i y
measu es
ha
may
mi iga e
e ec s
o
agmen-
a ion
gi en
limi ed
in o ma ion;
and
(3)
o
assess
esea ch
needs
o
he
de elopmen
o
sound
conse a ion
s a e-
gies.
P ima y
emphasis
is
gi en
o
he
pa e ns
o
local
g adien
( a he
han
biogeog aphic
pa e ns)
because
hese
bea
he
mos
ele ance
o
he
discussion
o
habi a
agmen a ion
and
i s
impac
on
he
main enance
o
spe-
cies
di e si y.
342
Main enance
o
F ui
Ba
Di e si y
343
Da a
Sou ces
and
Me hods
o
Analysis
S udy
Si es
The
Philippine
A chipelago,
lying
be ween
4°40'
N
o
2i°50'
N
la i ude
and
116°50'
E
o
136°35'
E
longi ude,
is
opical
h oughou
i s
ange.
Habi a
di e si y
is
mo e
ma ked
along
al i udinal
han
ac oss
la i udinal
o
longi u-
dinal
g adien s,
al hough
a iabili y
in
he
local
lo a
occu s
in
associa ion
wi h
clima ic
sub egions
(Alcala
1976).
Local
g adien s
encompass
h ee
p ima y
ege a ion
zones:
low-
land
dip e oca p
o es ,
mon ane
o es ,
and
mossy o es
(dip e oca p,
lowe
mon ane,
and
uppe
mon ane
ain
o -
es ,
in
Whi mo e
1984).
The
speci ic
ele a ions
a
which
hese
o es
ypes
occu
a y
among
moun ain
si es,
la gely
as
a
esul
o
di e ences
in
maximum
ele a ion
and
he
amoun
and
dis ibu ion
o
local
ain all
(Whi mo e
1984).
Moun ains
ha
a e
a
leas
1,500
m
in
ele a ion
suppo
well-de eloped
p ima y
ege a ional
ypes
o e
wide
al i-
udinal
anges,
and
ansi ion
zones
be ween
majo
o -
es
ypes
exis
as
dis inc
bands
(e.g.,
M .
Gui ingui ing,
Sibuyan
[Goodman
and
Ingle
1993];
M .
Guisayawan,
Ne-
g os
[Heaney
e
al.
1989];
and
M .
Isa og,
Luzon
[Ricka
e
al.
1991]).
Small
moun ains
exhibi
comp ession
o
ege a-
ional
zones,
which
hus
occu
a
ela i ely
lowe
ele a ions
in
wha
is
known
as
he
"Massene hebung
e ec
(e.g.,
M .
Pangasugan,
Ley e
[Heaney
e
al.
1989;
Ricka
e
al.
1993],
and
M .
Konduko,
Bili an
[Ricka
e
al.
1993])
(G ubb
and
Whi mo e
1966;
F ahm
and
G ads ein
1991).
M .
Talinis,
cen e ed
app oxima ely
a
9°16'
N,
123
12
E
on
Neg os
Island,
ex ends
upwa d
o
1850
m
and
exhibi s
well-de ined
lowland,
mon ane,
ansi ional
mon ane-mossy,
and
mossy
o es s
(see
ollowing
sec ion).
The
deg ee
o
an h opogenic
dis u bance
o
na u al
habi a s
a ies
g ea ly
among
hese
si es.
In
mos
loca ions,
p ima y
lowland
o es
is
absen
below
500
m,
al hough
limi ed
na u al
o
eplan ed
seconda y
o es
may
exis
be-
low
his
ele a ion.
Mosaic
pa ches
o
dis u bance
wi hin
p ima y
o es s
esul ing
om
small-scale
imbe
ex ac ion
o
shi ing
ag icul u e
a e,
likewise,
a
common
ea u e
o
he
o es s,
e en
in
a eas
designa ed
as
na ional
pa ks.
Field
Me hods
and
Da a
Sou ces
The
p incipal
in o ma ion
on
biogeog aphic
pa e ns
o
dis-
ibu ion
is
ha
epo ed
by
Heaney
(1986,
1991a)
and
Heaney
and
Ricka
(1990).
Addi ional
de ails
a e
de i ed
om
Heaney
and
Rabo
(1982),
Heaney
e
al.
(1991),
U zu -
um
(1992),
Goodman
and
Ingle
(1993),
and
Vincgue a
and
M ille
(1993).
The
summa y
p esen ed
he e
does
no
ake
in o
accoun
he
p oposed
change
in
he
species
s a us
o
Ace odon
luci e
(Heaney
e
al.
n.d.).
Pa e ns
o
ele a ional
g adien s
a e
summa ized
om
a
su ey
I
conduc ed
on
M .
Talinis
(Neg os
Island)
and
om
published
s udies
on
Bili an
(M .
Konduko:
Ricka
e
al.
1993),
Ley e
(M .
Pangasugan:
Heaney
e
al.
1989;
Ricka
e
al.
1993),
and
Neg os
(M .
Guinsayawan:
Heaney
e
al.
1981,
1989;
Heideman
and
Heaney
1989)
(Figu e
24.1).
Whe e
a ailable
and ele an ,
unpublished
da a
om
ecen s udies
a e
p o ided.
Da a
om
he
1990
ele a ional
ansec
s udy
on
M .
Talinis,
Neg os
Island,
p o ided
he
ocal
poin
o
analysis
on
ele a ional
ends.
Al hough
p e ious
s udies
on
a
neigh-
bo ing
moun ain
sys em,
M .
Guisayawan
(Heaney
e
al.
1989;
Heideman
and
Heaney
1989),
we e
mo e
ex ensi e
in
scope
and
e o ,
a
g ea e
p opo ion
o
he
ne ing
was
done
a
si es
ep esen ing
a
single
habi a
ype,
and
he
ull
ange
o
ele a ional
sampling
was
no
unde aken
wi hin
he
same
yea .
Thus,
esul s
o
he
M .
Guisayawan
s udies
may
be
less
compa able
wi h
da a
om
mo e
s anda dized
sampling
used
in
ecen
su eys
conduc ed
elsewhe e
in
he
Philippines.
S anda dized
su eys
o
he
ui
ba
auna
along
ele a-
ional
ansec s
ypically
in ol ed
unning
a
se ies
o
unde -
s o y
mis
ne s,
du ing
h ee
o
i e
nigh s,
wi hin
a
50-
o
100-m
ele a ional
band
a
si es
co esponding
o
each
ma-
jo
habi a
ype
(Ricka
1993).
On
M .
Talinis,
I
an
an
ele a ional
ansec
be ween
May
and
July
1990
using
six
ne ing
si es.
The
si es
we e
(1)
an
a ea
o
low-in ensi y
ag icul u e
and
seconda y
g ow h
(500
m),
cha ac e ized
by
s ands
o
coconu s,
co ee,
and
plo s
o
cul i a ed
ege-
ables
and
lowe s,
in e spe sed
wi h
pa ches
o
sh ubs
(Melas oma
spp.),
sawg ass
(Impe a a
sp.),
and
sca e ed
wild
igs
(Ficus
spp.);
(2)
a
na u ally
egene a ed
seconda y
o es
(500
m)
wi h
emnan
an h opogenic
plan s
such
as
banana
and
abaca
(Musa
spp.),
a ocado
(Pe sea
ame icana),
and
jack-
ui
(A oca pus
he ephyllus);
(3)
p ima y
lowland
o es
(750
m)
punc ua ed
by
small
(<0.5-ha)
dis u bed
pa ches
in
a i-
ous
s ages
o
egene a ion;
(4)
p ima y
mon ane
o es
(1,100
m)
ela i ely
ee
o
and
dis an
om
dis u bance,
(5)
ansi ional
mon ane-mossy o es
(1,250
m)
wi h
elemen s
o
p e ious
an h opogenic
dis u bance
(e.g.,
Musa
spp.
and
Bambusa
sp.);
and
(6)
p ima y
mossy
o es
(1,625
m)
a
he
summi
o
he
idge
sys em
on
which
he
i e
o he
sam-
pling
si es
we e
loca ed.
A
each
sampling
loca ion,
en
nylon
mis
ne s
(12
m
long
X
2.6
m
high)
we e
un
om
1800
o
0600
o
i e
consecu-
i e
nigh s.
Ex e nal
measu emen s
(size
measu emen s
and
body
mass)
we e
eco ded
o
each
ui
ba
cap u ed.
All
samples
we e
iden i ied
o
species
ollowing
Heaney
e
al.
(1987)
and
Ingle
and
Heaney
(1992),
and
assessed
o
age
and
344
R.
C.
B.
UTZURRUM
Pleis ocene
islands
as
de ined
by
cu en
1
20
m
ba hyme ic
line
0
200
1 1 1
1 1
II
Kilome e s
•18°
Luzon
Faunal
Region
1
6:8:1
G ea e
Mindo o
1
1:6:1
Palawan
Faunal
Region
7:1:1
Neg os-
Panay
Faunal
Region
Figu e
24.1.
Ex en
o
Pleis ocene
land
connec ions
(shaded
a eas,
which
co espond
o
he
cu en
120-m
ba hyme ic
line)
in
compa ison
o
he
cu en
opog aphy
(solid
lines)
o
he
Philippines.
The
i s
o
he
h ee
numbe s
ollowing
each
Pleis ocene
egion
name
co esponds
o
he
o al
numbe
o
ui
ba
species
p esen ;
he
second
numbe ,
he
subse
o
his
o al
ha
a e
endemic
o
he
Philippines;
and
he
hi d,
he
numbe
o
endemic
species
ha
a e
unique
o
he
aunal
egion.
Le e s
indica e
he
p incipal
loca ions
o
s udy
si es
men ioned
in
he
ex :
C,
moun ain
on
Ca anduanes;
G,
M .
Guisayawan,
Neg os;
G ,
M .
Gui ingui ing,
Sibuyan;
I,
M .
Isa og,
sou heas e n
Luzon;
Ki,
M .
Ki anglad,
Mindanao;
K,
M .
Konduko,
Bili an
Island;
P,
M .
Pangasugan,
Ley e;
T,
M .
Talinis,
Neg os.
ep oduc i e
s a us
by
me hods
modi ied
om
Heideman
(1987).
Vouche
specimens
we e
deposi ed
in
he
Field
Mu-
seum
o
Na u al
His o y
(Chicago),
Philippine
Na ional
Mu-
seum
(Manila),
Silliman
Uni e si y
Museum
o
Na u al
His-
o y
(Dumague e
Ci y),
and
he
eaching
collec ion
o
he
Depa men
o
Biology,
Bos on
Uni e si y.
Da a
Analysis
Analyses
o
da a
se s
o he
han
om
M .
Talinis
included
calcula ions
o
species
ichness
(S,
he
o al
numbe
o
di e en
species
cap u ed
a
each
si e),
o al
abundance
(exp essed
as
numbe s
o
ba s
cap u ed
pe
ne -nigh ),
and
ela i e
abundance
(numbe
o
ba s
o
a
gi en
species
cap-
u ed
pe
ne -nigh ).
Simila
ea men s
we e
applied
o
he
M .
Talinis
da a
se .
The
Shannon-Wiene
index
o
di e -
si y
(H')
was
also
calcula ed
o
he
M .
Talinis
( his
chap e )
and
M .
Pangasugan
(Heaney
e
al.
1989;
Ricka
e
al.
1993)
ele a ional
ansec
da a.
Wi hin-moun ain
and
be ween-
moun ain
di e si y
indices
we e
compa ed
using
a
- es
(Magu an
1988).
La ge
lying- oxes
(Ace odon
and
P e opus
species)
a e
cus oma ily
excluded
om
da a
analysis
be-
Main enance
o
F ui
Ba
Di e si y
345
Table
24.1
Dis ibu ions
o
he
26
Species
o
Philippine
F ui
Ba s
(P e opodidae)
ac oss
Pleis ocene
Faunal
Regions
(sensu
Heaney
1986)
as
Upda ed
om
Heaney
(1991)
Dis ibu ion
and
endemism
Spec'
Species
widesp ead
in
Indo-Aus alia
(6/26;
23%)
Species
sha ed
wi h
nea by
a chipelagos;
o
limi ed
dis ibu ion
in
he
Philippines
(4/26;
15%)
Endemic
species
widesp ead
in
oceanic
Philippines
(6/26;
23%)
Endemic
species
on
wo
o
mo e
Pleis ocene
islands
(2/26;
8%)
Endemic
species
on
only
one
Pleis ocene
island
(8/26;
31%)
Cynop e us
b ackyo is
Eonyc e is
spelaea
Mac oglossus
minimus
P e opus
hypomelanus
P e opus
ampy us
Rouse us
amplexicauda us
Dyacop e us
spadiceus
Megae ops
we mo ei
P e opus
dasymallus3
P e opus
speciosus
Ace odon
juba us
Eonyc e is
obus a
Haplonyc e is
ische i
Ha pyionyc e is
whi eheadi
P enochi us
jago i
P e opus
pumilus
P e opus
leucop e usuz
Nyc imene
abo i4-6
Ace odon
leuco is
Ace odon
luci e
Alionyc e is
pauciden a a
Dobsonia
chapmani
Haplonyc e is
sp.4
O op e opus
ca ilagonodus
P enochi us
mino
P e opus
sp.5
"Supe sc ip
numbe s
indica e
sou ces
o
upda es:
(1)
Heaney
and
Rabo
1982;
(2)
Heaney
e
al.
1991;
(3)
U zu um
1992;
(4)
Goodman
and
Ingle
1993;
(5)
Heaney
1993,
and
unpublished;
and
(6)
Vincigue a
and
Mulle
1993.
cause
hey
a e
no
eliably
sampled
in
unde s o y
ne
se s
(Heaney
e
al.
1989;
Heideman
and
Heaney
1989;
Ricka
e
al.
1993).
Howe e ,
I
included
he
smalles
o
he
lying-
oxes,
P e opus
pumilus,
in
he
analysis
o
he
M .
Talinis
da a.
The
ne s
o
he
M .
Talinis
ansec
we e
se
on
na ow
idges
and
we e
e ec i e
in
cap u ing
lying- oxes
ha
commu e
o e
idge ops.
Resul s
P e opodids
p esen ly
known
om
he
Philippines
ange
in
size
om
16
g
(e.g.,
Alionyc e is
pauciden a a;
Mac oglossus
minimus)
o
mo e
han
1
kg
(e.g.,
Ace odon
juba us
and
P e -
opus
ampy us).
O
he
26
species
p esen ,
16
(62%)
a e
es ic ed
o
he
Philippines,
including
6
species
in
he
en-
demic
gene a:
Alionyc e is
(1),
Haplonyc e is
(2),
O op e opus
(1),
and
P enochi us
(2)
(Heaney
e
al.
1987;
Heaney
1991a;
U zu um
1992).
Biogeog aphic
Dis ibu ion
Pa e ns
Biogeog aphic
analysis
o
species
ichness
among
Philip-
pine
ui
ba s
shows
dis ibu ions
conco dan
wi h
land
masses
o med
du ing
lowe ing
o
sea
le els
in
he
Pleis o-
cene
(Heaney
1991a).
Six
(23%)
o
he
nonendemic
species
a e
widesp ead
wi hin
he
Philippines
and
in
he
Indo-Aus-
alian
egion;
he
o he
ou
(15%)
a e
es ic ed
wi hin
he
Philippines
and
a e
sha ed
wi h
nea by
islands
(Table
24.1;
see
also
Figu e
24.1)
(Heaney
e
al.
1987;
Heaney
1991a).
Among
he
endemic
species,
h ee
basic
pa e ns
o
geog aphic
dis ibu ion
eme ge:
(1)
species
ha
a e
wide-
sp ead
in
oceanic
Philippines
(six);
(2)
species
ha
a e
sha ed
a
leas
be ween
wo
Pleis ocene
islands
( wo);
and
(3)
species
ha
a e con ined
o
only
one
Pleis ocene
island
(eigh )
(Table
24.1;
Figu e
24.1)
(Heaney
and
Rabo
1982;
Heaney
1991a
1993;
U zu um
1992;
Goodman and
Ingle
1993).
346
R.
C.
B.
UTZURRUM
Da a
om
well-in en o ied p esen -day
islands
e eal
a
signi ican
posi i e
ela ionship
be ween
species
numbe s
and
island
size
(Heaney
1991a).
Howe e , his
ela ionship
does
no
hold
ue
o
endemic
species.
Luzon
Island
(108,171
km2)
is
he
la ges
island
o
hold
an
endemic
(i.e.,
O op e opus
ca ilagonodus).
The
islands
o
Mindanao
(99,078
km2;
Alionyc e ispauciden a a),
Neg os
(13,670
km2;
Dobsonia
chapmani),
Panay
(12,300
km2;
Ace odon
luci e ),
Palawan
(11,785
km2;
Ace odon
leuco is),
and
Mindo o
(9,735
km2;
undesc ibed
P e opus
sp.)
also
ha e
one
endemic
species
each
(Heaney
1991a,
1993;
Heaney
e
al.
n.d.).
Sibuyan
Island
(463
km2;
undesc ibed
Haplonyc e is
sp.),
an
oceanic
island
wi h
no
his o ical
connec ion
o
any
o
he
Pleis ocene
islands,
is
he
smalles
Philippine
island
cu en ly
known
o
ha e
an
endemic
ba
species
(U zu um
1992;
Goodman
and
Ingle
1993).
The
high
deg ee
o
species
o e lap
wi hin
and
among
Pleis ocene
islands
sugges s
he
impo ance
o
o e wa e
coloniza ion
in
shaping
di e si y
and
dis ibu ion
(Heaney
and
Ricka
1990;
Heaney
1991a).
Measu es
o
gene
low
con i m
hese
pa e ns
(Pe e son
and
Heaney
1993).
Pleis-
ocene
land-b idge
islands
in e media e
in
size
be ween
Ne-
g os
and
Sibuyan
ha
lack
endemic
species
(e.g.,
Dinaga
and
Ley e)
u he
suppo
he
impo ance
o
Pleis ocene
land
connec ions,
o
con e sely
he
lack
he eo ,
in
shaping
specia ion
e en s
wi hin
he
a chipelago.
Species
wi h
dis-
junc
dis ibu ions
pose
an
in e es ing
puzzle
in
ou
unde -
s anding
o
hese
aunal
pa e ns
and
he
mechanisms
ha
shaped
hem.
Pleis ocene
land
connec ions
among
islands
do
no
accoun
o
he
disjunc
dis ibu ion
o
Nyc imene
abo i
(in
Cebu,
Neg os,
and
Sibuyan
islands
only;
Heaney
e
al.
1987;
Goodman
and
Ingle
1993,
Vincigue a
and
Miille
1993),
o
o
P e opus
leucop e us
(in
Ca anduanes,
Dinaga ,
and
no he n
Luzon;
Heaney
and
Rabo
1982;
Heaney
e
al.
1987,
1991).
These
cases
sugges
p ocesses
ha
ha e
in luenced
ex inc ion
e en s
in
he
pas —o
may
simply
e lec
in o ma ion
gaps
ha
need
o
be
illed
om
mo e
ho ough
in en o ies.
Ele a ional
G adien s
in
Species
Di e si y
and
Abundance:
Gene al
T ends
Gene al
pa e ns
ha e
eme ged
om
ele a ional
ansec
su eys:
(1)
species
ichness
eaches
i s
maximum
in
p i-
ma y
lowland
o es
and
declines
wi h
ele a ion;
(2)
o al
abundance
is
highes
in
dis u bed
a eas
and
dec eases
wi h
ele a ion
in
o es
habi a s;
and
(3)
species
assemblages
in
dis u bed
a eas
end
o
be
cha ac e ized
by
he
p esence
o
geog aphically
widesp ead
species,
whe eas
endemic
spe-
cies
end
o
be
associa ed
wi h
o es
habi a s
(Heaney
e
al.
1981,
1989;
Heideman
and
Heaney
1989;
Ricka
e
al.
1993).
Endemic
species
a e
some imes
ound
in
low
num-
be s
in
mode a ely
dis u bed
habi a s
p o ided
ha
hese
a e
adjacen
(wi hin
1
km)
o
p ima y
habi a s
(Heideman
and
Heaney
1989;
Ricka
e
al.
1993).
Despi e
he
abundance
o
ui
ba s
in
u ban
o cha ds
and
ag icul u al
a eas
a
emo ed
om
o es ed
si es,
en-
demic
species
ha e
no
been
eco ded
a
hese
loca ions
(Heideman
1987;
Heaney
e
al.
1989;
Heideman
and
Heaney
1989;
Ricka
1993),
wi h
he
excep ion
o
P e opus
pumilus
(R.
C.
B.
U zu um,
unpublished
eco ds
o
cap-
u es
in
Dumague e
Ci y
[1986]
and
Sia on
[1992]
on
Ne-
g os).
In
mos
cases,
species
a
high
ele a ions
ep esen
a
subse
o
he
lowland
communi y,
indica ing
a
lack
o
high-
ele a ion
specialis s.
In
1992
and
1993,
Alionyc e ispauciden-
a a
was
cap u ed
in
conside able
numbe s
on
M .
Ki an-
glad,
Mindanao,
bu
only
a
ele a ions
abo e
1,500
m
in
mon ane
and
mossy
o es
(L.
R.
Heaney,
pe sonal
commu-
nica ion).
Ele a ional
G adien s
on
M .
Talinis,
Neg os
Island
O
he
15
species
o
ui
ba s
occu ing
in
Neg os
Island,
9
we e
ne ed
in
his
s udy
(Table
24.2).
The
cap u e
o
Cynop e us
b achyo is
a
1,250
m
is
a
new
ele a ional
eco d
o
he
species;
all
o he
species
ha e
been
eco ded
a
ele-
a ions simila
o
o
g ea e
han
in
he
p esen
s udy
(Heaney
and
Heideman,
unpublished
da a).
Missing
om
he
samples
we e
3
species
o
lying- oxes
(Ace don
juba us,
P e opus
hypomelanus,
and
P.
ampy us)
no
expec ed
o
be
cap u ed
in
unde s o y
ne s,
an
uncommon
endemic
spe-
cies,
Eonyc e is
obus a
(U zu um
1992),
and
he
epo edly
ex inc
Dobsonia
chapmani
(Heaney
and
Heideman
1987).
The
o al
numbe s
o
ui
ba s
cap u ed we e
highes
a
he
ag icul u al
si e
(3.24
ba s/ne -nigh )
and
dec eased
wi h
ele a ion
in
o es
( om
1.9
ba s/ne -nigh
in
lowland
o es
o
0.22
ba s/ne -nigh
in
mossy
o es ;
Table
24.2).
This
o e all
end
was
ue
o
all
he
nonendemic
species
as
well
as
he
endemic
P enochi us
jago i
(Table
24.2).
All
o he
endemic
species
we e
uncommon
o
absen
a
he
ag icul u al
si e.
Ins ead,
hey
we e
ound
in
highe
num-
be s
in
lowe
ele a ion
o es
(lowland
o
mon ane),
al-
hough
Nyc imene
abo i
was
ela i ely
uncommon
e en
in
o es
habi a .
These
pa e ns
o
abundance
and
dis ibu-
ion
a e
consis en
wi h
a
p e iously
obse ed
ecological
dicho omy
be ween
endemic
and
nonendemic
species
(Heaney
e
al.
1989).
As
p edic ed
by
Heaney
e
al.
(1989),
species
ichness
was
highes
in
lowland
p ima y
o es
(S
=
9)
and
lowes
in
mossy o es
(S
=
3)
(Table
24.2).
Unexpec -
edly
high
le els
o
species
ichness
and
abundance
we e
eco ded
a
he
mon ane-mossy
ansi ional
zone.
Main enance
o
F ui
Ba
Di e si y
347
Table
24.2
Summa y
o
Cap u es
o
F ui
Ba s
(P e opodidae)
on
M .
Talinis,
Neg os
O ien al,
May-July
1990
Si e
(and
ele a ion)
A
B
c
D
E
F
Species
(500
m)
(500
m)
(725
m)
(1,100
m)
(1,250
m)
(1,625
m)
Cynop e us
b achyo is
0.74
0.68
0.54
0.08
0.10
0.02
1.04
0.10
0
0
0.16
0.04
0.02
0
0
0
0
Eonyc e is
spelaea
0.38
0.14
0.02
0
0.10
0.02
1.04
0.10
0
0
0.16
0.04
0.02
0
0
0
0
*
Haplonyc e is
ische i
0.04
0.10
0.26
0.42
0.10
0.02
1.04
0.10
0
0
0.16
0.04
0.02
0
0
0
0
*Ha pyionyc e is
whi eheadi
0 0
0.04
0
0.10
0.02
1.04
0.10
0
0
0.16
0.04
0.02
0
0
0
0
Mac oglossus
minimus
0.94
0.20
0.54
0.10
0.36
0
0
0.16
0.04
0.02
0
0
0
0
*Nyc imene
abo i
0 0
0.06
0.06
0.02
0
0.04
0
0
0
0.16
0.04
0.02
0
0
0
0
'P enochi us
jago i
0.36
0.04
0.16
0
0.02
0
0.04
0
0
0
0.16
0.04
0.02
0
0
0
0
*P e opus
pumilus
0.02
0
0.22
0
0.02
0
0.04
0
0
0
0.16
0.04
0.02
0
0
0
0
Rouse us
amplexicauda us
0.76
0.08
0.06
0
0.02
0
0.04
0
0
0
0.16
0.04
0.02
0
0
0
0
All
nonendemics
2.82
1.10
1.16
0.18
0.48
0.02
All
endemics
(*)
0.42
0.14
0.74
0.48
1.20
0.20
AH
species
3.24
1.24
1-90
0.66
1.68
0.22
(S
=
7)
(S
=
6)
(S
=
9)
(S
=
4)
(S
=
7)
(S
-
3)
No es:
Da a
a e
gi en
as
numbe
o
ba s
cap u ed
pe
ne -nigh
Each
si e
had
a
o al
ne ing
e o
o
50
ne -mgjus.
Si e
designa ions:
A,
mixed
ag icul u e/seconda y
g ow h;
B,
seconda y
lowland
o es ;
C,
p ima y
lowland
o -
es -
D
mon ane
o es ;
E,
ansi ional
mon ane-mossy;
F,
mossy
o es .
An
as e isk
deno es
endemic
species.
Discussion
Biogeog aphic
In o ma ion
and
he
Design
o
a
Sys em
o
P o ec ed
A eas
Pa e ns
o
biogeog aphic
dis ibu ion
o
species
a e
ele-
an
o
ui
ba
conse a ion
a
wo
le els.
Fi s ,
hey
p o-
ide
a
biological
basis
o
he
selec ion
o
impo an
si es
o
p o ec ion.
Second,
island
size
and
species
di e si y
e-
la ionships
e ealed
om
biogeog aphic
analysis
p o ide
es ima es
o
a eal
equi emen s
o
he
main enance
o
species
di e si y
based
on
es ima ed
a es
o
coloniza ion
and
ex inc ion
(Heaney
1986).
Gi en
he
high
deg ee
o
simila i y
in
he
composi ion
o
ui
ba
assemblages
be-
ween
Pleis ocene
aunal
egions
and
among
islands
o
a
egion,
a
minimum
o
eigh
p o ec ed
a eas,
o
be
loca ed
in
he
cu en
islands
o
Luzon
(one
in
he
no he n
ip
and
a
second
in
he
sou heas e n
peninsula),
Mindanao,
Ne-
g os,
Panay,
Palawan,
Mindo o,
and
Sibuyan, may
heo e i-
cally
p o ec
all
26
species
(Table
24.3).
In e es ingly,
he
pa e ns
o
species
ichness
and
le els
o
endemism
seen
in
ui
ba s
ela i e
o
Pleis ocene
aunal
egions
a e
conco -
dan wi h
hose
o
o he
e eb a e
g oups,
including
non-
olan
mammals
(Heaney
1986,
1993;
Heaney
and
Ricka
1990),
bi ds
(Dicke son
1928;
Dickinson
e
al.
1991),
and
amphibians
and
ep iles
(B own
and
Alcala
1970;
Hague
e
al.
1986).
These
zoogeog aphic
pa e ns
also
o e lap
well
wi h
phy ogeog aphic
pa e ns
o
di e si y
and
endemism
(D.
A.
Madulid,
pe sonal
communica ion;
Heaney
1993).
Thus,
a
pa ks
sys em
modeled
on
he
biogeog aphy
o
hese
mo e
speciose
e eb a es
and
plan s
would
subsume
p o ec ion
o
he
ui
ba s.
In
he
ecen
In eg a ed
P o ec ed
A eas
Sys ems
(IPAS)
ini ia i e
owa d
he
ede elopmen
o
he
Philippine
pa ks
sys em
(IUCN
1991),
ecommended
p io i y
a eas
based
on
mammalian
and
lo is ic
di e si y
pa e ns
ha e
led
o
he
inclusion
o
si es
in
Luzon,
Neg os,
and
Mindanao
(Heaney
1993).
This
plan will
inco po a e
habi a s
o
85%
(22
o
26)
o
all
ui
ba s,
including
75%
(12
o
16)
o
he
endemic
species
(see
Table
24.3).
I
should
be
emphasized
ha
a eal
size
equi emen s
based
on
es ima ed
a es
o
coloniza ion
and
ex inc ion
should
be
ea ed
as
conse a i e guidelines,
gi en
ha
hese
and
o he
e olu iona y
p ocesses
ha e
oc-
cu ed
in
a
con ex
o
habi a s
(i.e.,
con inuous
expanse
o
o es s)
la gely
di e en
om
p esen -day
condi ions
(i.e.,
discon inuous
pa ches
o
o es s).
Thus,
whene e
possible,
he
la ges
con inuous
a ea
o
sui able
habi a
a ailable
a
each
p io i y
si e
should
be
chosen.
The e
a e
pe cei ed
poli ical
ba ie s
agains
he
designa-
ion
o
small
islands
o
p o ec ion
e en
when
s ong
bio-
logical
easons
exis
(see
U zu um
1991
o
discussion).
348
R.
C.
B.
UTZURRUM
Table
24.3
Theo e ical
Pe cen ages
o
Species
Tha
Would
Be
P o ec ed
by
a
Designa ed
Pa k
in
he
Philippine
Islands
All
species
Endemic
species
( o al,
26)
( o al,
16)
Island
Cumula i e
Cumula i e
(and
a ea)
No.
%
No.
%
Mindanao
(99,078
km2)
17
65%
8
50%
Luzon
(108,171
km2)
+3
77%
+2
62%
Neg os
(13,670
km2)
+2
85%
+2
75%
Panay
(12,300
km2)
+
1
88%
+
1
81%
Mindo o
(9,735
km2)
+
1
92%
+
1
88%
Palawan
(11,785
km2)
+
1
96%
+
1
94%
Sibuyan
(463
km2)
+
1
100%
+
1
100%
No es:
Mindanao
is
anked
i s
because
i
has
he
mos
species.
Luzon
and
Neg os
ollow,
based
on
he
numbe
o
species
ha
hese
islands
will
add
o
he
heo e ical
p o ec ed
pool.
The
las
ou
islands
a e
anked
on
he
basis
o
conse a ion
p io -
i y
o
he
addi ional
species
unique
o
each
heo e ical
ese e
(see
Heaney
1993;
Heaney
and
U zu um
1991;
U zu um
1992).
Fou
o
six
o es
pa ks
designa ed
by
he
In eg a ed
P o ec ed
A eas
Sys em
(IPAS)
will
be loca ed
in
Mindanao
(M .
Ki anglad),
Luzon
(Palanan
Wilde ness
A ea
and
Subic
Bay),
and
Neg os
(M .
Kanlaon).
Such
may
be
he
case
o
Sibuyan
Island.
A
leas
i e
unde-
sc ibed
species
o
mammals
we e
ecen ly
(i.e.,
since
1990)
disco e ed
on
his
island
(one
ui
ba ,
Haplonyc e is
sp.,
and
ou
mu ids,
Apomys
(2),
Ch o o nys
(1),
and
Ta somys
(1);
Goodman
and
Ingle
1993).
I s
deg ee
o
isola ion
and
ela-
i e
s a e
o
"unde de elopmen ,"
howe e ,
elemen s
inhe -
en ly
a o ing
species
pe sis ence,
could
wo k
agains
i s
selec ion
o
p o ec ion
unde
he
na ional
pa ks
scheme.
Ongoing
poli ical
machina ions
exis
ha
exploi
he
ich
imbe
esou ces
on
M .
Gui ingui ing
(N.
Ingle,
pe sonal
communica ion),
bu
he
island's
ela i e
isola ion
om
ade
and
communica ion
p ecludes
he
na ional
isibili y
ha
could
highligh
a
need
o
include
i
in
a
sho
lis
o
p io i y
pa ks.
Ins ead,
p o ec ion
o
a eas
such
as
Sibuyan,
and
o he
si es
suppo ing
single
island
endemics,
may
de-
pend
on
he
de elopmen
o
a
ne wo k
o
seconda y
e-
gionally
managed
conse a ion
pa ks
ha
complemen s
he
na ional
pa ks
sys em
(U zu um
1991).
The e
a e
many
po en ially
species- ich
and
biogeo-
g aphically
in e es ing
a eas
ha
emain
ela i ely
un-
known
in
he
Philippines.
Recen
e o s
o
sys ema ically
in en o y
local
mammalian
auna
ha e
been
concen a ed
mos ly
on
a
la i udinal
band
ex ending
om
sou he n
Luzon
(in
he
no h)
o
no he n
Mindanao
(in
he
sou h)
(see
Figu e
24.1).
Al hough
Palawan
has
been
e y
a ac-
i e
o
esea che s
o
bi ds
(Dickinson
e
al.
1991)
and,
in
pa ,
o
mammals
(Heaney
1986),
i s
ba
auna
is
poo ly
s udied
(Heaney
1991a).
The
no he nmos
egions
o
Luzon
(especially
he
no heas e n
bo de )
s ill
suppo
ex-
ensi e
o es s,
ye
mos
o
he
ecen
su eys
in
he
a ea
ocused
p ima ily
on
bi ds
(Malla i
and
Jensen
1993).
These
esea ch
ends
e lec
in
pa
he
oppo unis ic
na u e
o
esea ch
wo k
in
he
Philippines,
in
e ms
o
unding,
po-
li ical
s abili y,
and
he
expe ise
a ailable.
Habi a
A ini y
and
I s
Implica ions
o
Conse a ion
I
is
widely
ecognized
ha
o es s
a e
essen ial
o
he
conse a ion
o
biodi e si y
in
opical
egions.
Mo e
im-
po an ly,
he
design
and
managemen
o
p o ec ed
a eas
should
inco po a e
p o isions
o
di e en
a ini ies
among
species
o
g adien s
in
ege a ional
s uc u e
and
composi-
ion
and
abio ic
condi ions
h oughou
he
local
ange
o
a
o es
habi a .
Thus,
a
basic
unde s anding
o
local
pa e ns
o
species
dis ibu ion
and
species-habi a
associa ions
is
o
u mos impo ance.
Lowland
o es s
a e
essen ial
o
he
main enance
o
maximum
local
and,
he e o e,
o e all
di e si y
o
Philip-
pine
ui
ba s
(see
Resul s).
This
equi emen
necessi a es
he
inclusion
o
lowland
o es
in o
pa ks
o
ese es.
Fo -
es s
a
low
ele a ions
may
also
be
c i ical
o
he
pe sis ence
o
highe -ele a ion
o es s.
These
uppe -ele a ion
habi a s,
in
u n,
may
be
in eg al
o
he
main enance
o
lowland
o es
di e si y.
Mon ane
o es
habi a s
on
Philippine
moun-
ains
ha e
expe ienced
episodes
o
expansion
and
e ac ion
associa ed
wi h
clima e
changes
in
he
Pleis ocene
(Heaney
1991b).
Al hough
he
ele ance
o
hese his o ical
changes
in
ege a ional
co e
o
specia ion
e en s
is
mo e
appa en
o
oden s
(Heaney
and
Ricka
1990),
hey
may
ha e
in-
luenced
specia ion
e en s
o
pa e ns
o
habi a
specializa-
ion
in
ui
ba s
as
well.
The
dis ibu ional
pa e n
o
Alio-
nyc e is
pauciden a a
on
M .
Ki anglad
indica es
ha
o he
uppe -ele a ion
habi a
specialis s
among
he
ui
ba s
a e
likely
o
occu in
associa ion
wi h
b oadly
dis ibu ed
and
well-de eloped
mon ane-mossy
o es s.
As
on
M .
Ki anglad,
hese
condi ions
may
occu in
moun ains
exceeding
1,500
m
in
ele a ion.
Howe e ,
ew
Philippine
moun ains
bea ing
ele a ions
close
o
2,000
m
o
g ea e
ha e
been
su eyed.
Fu he mo e,
we
ha e
a
e y
limi ed
unde s anding
o
he
na u e
and
dynamics
o
mic ogeog aphic
p e e ences
among
and
wi hin
ui
ba
species.
While
su eys
indica e
lowland
o es s
as
he
local
cen e s
o
species
ichness,
hey
do
no
necessa ily
e eal
o
wha
ex en
adjoining
ac s
o
o es s
con ibu e
o
he
main enance
o
his
di e si y.
Fo
example,
p elimina y
analysis
o
wi hin-species
di e ences
in
he
ele a ional
dis ibu ion
o
O op e opus
ca ilagonodus
on
M .
Isa og
and
Zambales
e ealed
ele a ional
seg ega-
Main enance
o
F ui
Ba
Di e si y
349
ion
be ween
males
and
emales
o
he
species,
a
mecha-
nism
ha
may
educe
in e sexual
compe i ion
(Ruedas
e
al.
1994).
The
impo ance
o
mon ane
and
mossy
o es
habi a s
o
he
main enance
o
biodi e si y
is
e en
mo e
c ucial
o
non olan
mammals.
Philippine
mu id
oden s
exhibi
peaks
in
species
ichness
and
endemism
in
mon ane
and
mossy
o es s
(Heaney
and
Ricka
1990;
Ricka
e
al.
1991).
In
many
o
he
moun ains
su eyed,
pa e ns
o
a ian
di e -
si y
mi o
ha
o
ui
ba s,
wi h
lowland
o es
as
he
locus
o
maximum
species
ichness.
Howe e ,
endemism
i sel
may
be
cen e ed
in
he
uppe -ele a ion
o es s,
as
was
docu-
men ed
in
he
no he n
Sie a
Mad e
(Malla i
and
Jensen
1993).
Toge he ,
hese
indings
indica e
he
need
o
a
ull
ele a ional
complemen
o
o es
habi a s
in
any
designa ed
conse a ion
si e
i
maximum
species
di e si y
is
o
be
p o ec ed.
Impac s
o
Fo es
F agmen a ion
on
F ui
Ba
Di e si y
Conse a ion
o
biodi e si y
mus
add ess
whe he
and
how
con empo a y
deg ada ion
and
agmen a ion
o
habi-
a s
wi hin
islands
will
in luence
ecological
p ocesses
and,
hus,
local
pa e ns
o
species
assemblage
and
dis ibu ion.
I
s ong
p e e ence
o
o es
habi a s
inhibi s
ui
ba
mo emen s
o e
agmen ed
landscapes,
despi e
hei
in-
he en
agili y,
hen
mechanisms
o
species
main enance
and
geog aphic s uc u ing
may
be
a ec ed.
Indeed,
gene ic
a ia ion
in
Cynop e us
b achyo is
(a
nonendemic)
and
Haplo-
nyc e is
ische i
(an
endemic)
sugges s
ha
he
educed
gene
low
seen
in
he
la e
species
ela es,
in
pa ,
o
i s
g ea e
a ini y
o
speci ic
habi a s
(Pe e son
and
Heaney
1993).
The
occu ence
o
an
endemic
species
o
Haplonyc e is
on
Sibuyan
Island
demons a es
e olu iona y
s abili y
in
small
isola ed
popula ions
(Pe e son
and
Heaney
1993).
Thus,
i
is
di icul
o
p edic
whe he
inc eased
agmen a ion
o
once
con inuous
popula ions
(wi hin
islands)
in
he
ecen
pas
will
shi
e olu iona y
p ocesses
owa d
he
nega i e
ajec-
o ies
associa ed
wi h
inc easingly
smalle
popula ion
sizes
(see
Lande
1988).
Ele a ional
g adien s
in
ui
ba
di e si y
and
abundance
on
Neg os
and
Ley e
islands
illus a e
how
habi a
agmen-
a ion
o
deg ada ion
a ec s
local
communi y
s uc u e.
Neg os
and
Ley e
we e
pa s
o
wo
di e en
Pleis ocene
islands
bu
exhibi
a
mode a ely
high
deg ee
o
simila i y
in
hei
ui
ba
auna
(Heaney
1991a).
Bo h
islands
suppo
13
ex an
ui
ba
species,
wi h
12
species
sha ed
by
bo h
a eas
(Heaney
e
al.
1989).
Whe eas
Neg os
Island
has
Nyc imene
abo i,
Ley e
has
P enochi us
mino ,
bo h
o
which
a e
en-
demic and
sha e
an
a ini y
o
o es
habi a s
(Heaney
e
al.
1989;
Ricka
1993).
Thus,
hey
may
be
conside ed
ecologi-
cal
equi alen s
o
he
pu pose
o
compa ing
gene al
pa -
e ns
be ween
he
wo
islands.
On
M .
Talinis
(Neg os),
p ima y
o es
was
absen
om
below
he
500-m
ele a ion
bu
ex ended
o e
a
g ea e
dis ance
o
a
summi
o
1,850
m.
Dis u bed
pa ches
wi hin
he
p ima y
o es
we e
no
un-
common.
The
p ima y
o es
in
M .
Pangasugan
(Ley e)
was
comp essed
o e
a
na owe
ele a ional
ange
because
he
summi
was
lowe
(1,150
m).
Dis u bances
wi hin
o -
es s
we e
minimal
and
con ined
wi hin
he
lowe
300-m
ele a ion;
lowland
o es
g aded
in o
ag icul u al
a eas
be-
low
200
m.
M .
Talinis
has
sha pe
opog aphical
ea u es
consis ing
o
deep,
s eep
gulleys
bisec ing
sha p
na ow
idges.
Thus,
he
in e aces
be ween
low-
and
high-ele a ion
zones,
and
be ween
o es
and
ag icul u al
o
seconda y
g ow h
a eas,
we e
g ea e
he e
han
on
M .
Pangasugan.
On
Neg os,
endemic
species,
such
as
Haplonyc e is
ische i
and
P enochi us
jago i,
we e
absen
a
ele a ions
below
500
m,
wi h
he
excep ion
o
P e opuspumilus
(Figu e
24.2).
This
lack
coincides
wi h
he
absence
o
o es
habi a
below
his
ele a ion.
Con e sely,
he
pe sis ence
o
endemic
species
nea
sea
le el
on
Ley e
coincides
wi h
he
lowe
ex en
o
o es
on
M .
Pangasugan.
Bo h
a eas
exhibi ed
maximum
species
ichness
in
p i-
ma y
lowland
o es
(Table
24.4).
The
ansi ion
om
low-
land
o
mon ane
o es
was
he
uppe
limi
o
a
signi ican
shi
in
species
ichness.
Di e si y
indices
o
si es
abo e
lowland
o es
do
no
di e
signi ican ly,
al hough
alues
show
a
gene al
downwa d
end
wi h
inc easing
ele a ion
(Magu an
1988:
- es
on
H'
alues
and
a iances
o
H',p
>
0.05)
(Table
24.4).
Thus,
lowland
o es
zones
may
de ine
he
uppe
ele a ional
bounda y
o
maximum
ui
ba
di-
e si y.
Species
di e si y
le els
(H')
do
no
di e
s a is ically
(Magu an
1988:
- es
on
H'
and
a iance
o
H',
p
>
0.05)
be ween
pai s
o
equi alen
habi a s
e en
when
hese
zones
occu ed
a
di e en
ele a ions
on
each
moun ain
(Table
24.4).
A enua ion
in
species
numbe s
occu ed
a
he
an-
si ion
om
mon ane
o
mossy
o es
on
M .
Pangasugan,
bu
no
on
M .
Talinis.
On
M .
Talinis,
species
ichness
was
highe
in
he
mon ane-mossy
ansi ion
o es
han
ex-
pec ed.
I
a ibu e
his
o
he
a ypical
occu ence
o
he
nonendemic
species
Cynop e us
b achyo is
and
Eonyc e is
spelaea
a
he
si e
(see
Table
24.2).
On
M .
Guinsayawan
(no heas
o
Talinis),
nei he
o
hese
species
occu ed
abo e
uppe
mon ane
o es
(Heaney
e
al.
1989),
and
unpublished
da a
om
M .
Isa og
(in
Luzon;
L.
R.
Heaney,
pe sonal
communica ion)
e ealed
he
same
pa e n
o
ele a ional
dis ibu ion
o
hese
species.
I
in e -
p e
his
unexpec edly
high
le el
o
species
ichness
a
he
ansi ion
zone
o
mon ane
and
mossy
o es s
on
M .
Talinis
as
an
upwa d
ange
ex ension
o
species
ypical
o
lowland
350
R.
C.
B.
UTZURRUM
(A)
Neg os
c
o
•*—
03
>
0)
LU
2000
(B)
Ley e
OJ
c75
3
%
£
CC
>•
O
0<W3
^
J.
OUJSO^ za-
ll
O
UJ
2
£
Q.
CL
Figu e
24.2.
Compa a i e
ele a ional
dis ibu ion
o
ui
ba s
on
(A)
Neg os
Island
(a
composi e
om
M .
Guinsayawan
and
M .
Talinis
s udies)
and
(B)
Ley e
(M .
Pangasugan).
Species
a e
iden i ied
by
he
i s
wo
le e s
o
he
genus
and
species
names
(see
Table
24.1
o
lis ).
The
ho izon al
lines
deno e
lowe
ele a ional
limi s
o
he
a ious
ypes
o
o es s:
solid
line,
p ima y
o es ;
dashed
line,
mon ane
o es
(app oxi-
ma e);
do ed
line,
ansi ion
in o
mossy
o es ;
dashed-and-do ed
line,
well-de eloped
mossy
o es .
P ima y
o es
on
Neg os
begins
a
an
ele a ion
o
500
m, whe eas
on
Ley e
i
is
s ill
p esen
a
50
m.
Maximum
ele a ions
a
he
s udy
si es
we e
1,800
m
(Neg os)
and
1,150
m
(Ley e).
o
dis u bed
habi a
in
esponse
o
habi a
dis u bance
a
his
si e
and
i s
inc eased
p oximi y
o
cul i a ed
ields
on
adjoining
slopes.
This
esponse
should
be
di e en ia ed
om
inc eased
di e si y
ha
may
occu
a
zones
whe e
communi ies
o m
eco ones
(Rickle s
1979).
In
his
pa icu-
la
case,
he
la e
phenomenon
does
no
uly
apply
be-
cause
he
ui
ba
species
ound
in
he
adjoining
mon ane
and
mossy
si es
we e
no
dis inc
om
each
o he .
I
d aw
wo
poin s
o
ele ance
om
he
p eceding com-
pa isons
o
he
main enance
o
species
di e si y.
Fi s ,
changes
in
he
quali y
and
quan i y
o
o es
habi a s
al e
he
na u e
o
species
assemblages
wi hin
o es
ypes
by
a ec ing
bo h
he
ela i e
numbe s
among
species
and
he
ypes
o
species
p esen .
As
a
co olla y,
hese
esul s
sugges
ha
(1)
ligh
o
mode a e
le els
o
habi a
dis u bance,
whe e
he
p ima y
o es
s uc u e
and
composi ion
and
clima ic
condi ions
a e
e ained,
esul
in
highe
species
ichness
han
would
be
expec ed
in
undis u bed
o es
o
Table
24.4
To al
Abundance
o
F ui
Ba s
and
Measu es
o
Thei
Species
Richness,
Di e si y,
and
E enness
along
Two
Moun ains
Si e
Ele a ion
(m)
Abundance
(ba s/ne -nigh )
No.
o
ba
species
Di e si y
(H'y
E ennes
M .
Pangasugan,
Ley e
Island''
Ag icul u e
and
dis u bed
lowland
o es
50
6.08
6
1.690*
0.943
Lowland o es ,
2
si es:
(1)
P ima y
o es ,
dis u bed
300
0.60
6
1.277*
0.712
(2)
P ima y
o es
500
0.82
8
1.829*
0.880
P ima y
mon ane
o es
700
0.97
7
1.065
0.547
P ima y
mossy
o es
950
0.45
3
0.730 0.660
M .
Talinis,
Neg os
Island
Ag icul u e
and lowland
seconda y
g ow h
500
3.24
7
1.618*
0.831
Lowland
o es ,
2
si es:
(1)
Seconda y
o es
500
1.24
6
1.360*
0.759
(2)
P ima y
o es ,
dis u bed
750
1.90
9
1.792*
0.816
P ima y
mon ane
o es
1,100
0.66
4
1.047
0.755
T ansi ional
mon ane-mossy
o es
1,250
1.68
7
1.157
0.595
P ima y
mossy
o es
1,625
0.22
3
0.760
0.691
"An
H'
alue
( he
Shannon-Weine
index)
ma ked
by
an
as e isk
(*)
di e s
signi ican ly
om
he
H'
alue
o
he
nex
highe
ele a ion
on
he
same
moun ain.
'M .
Pangasugan
da a
a e
om
Heaney
e
al.
(1989)
and
Ricka
e
al.
(1993).
No e
ha ,
unlike
M .
Talinis,
M .
Pangasugan
had
no
ansi ional
mon ane-mossy
o es .
Main enance
o
F ui
Ba
Di e si y
351
simila
ype
and
ele a ion;
(2)
he
changes
in
he
commu-
ni y
s uc u e
ela ing
o
mode a e
habi a
dis u bance
e-
sul
p ima ily
om
local
ange
ex ensions
o
nonendemic
species
ha
a e
ypically
associa ed
wi h
dis u bed
habi a s
and
a e
a e
o
absen
in
p ima y
o es
(especially
abo e
lowland
o es );
(3)
la ge-scale
habi a
dis u bances
esul -
ing
in
deg ada ion
o
he
p incipal
o es
s uc u e,
he
al e a ion
o
associa ed
clima e
condi ions,
and
(o )
ma ked
agmen a ion
o
o me ly
con inuous
ac s
o
o es s
may
esul
in
lowe
le els
o
species
ichness
han
would
be
expec ed;
and
(4)
he
dec ease
in
species
ichness
in
hea ily
dis u bed
o
agmen ed
o es
habi a s
is
associa ed
p i-
ma ily
wi h
he
disappea ance
o
endemic
species
o
wi h
hei
inc easing
a i y.
Second,
di e ences
in
he
opog aphical
ea u es
o
moun ain
o es s
in luence
he
esponses
o
bo h
plan
and
ui
ba
communi y
s uc u e.
S eepness,
uggedness,
and
i egula i y
in
opog aphy
de e mine
he
deg ee
o
in e -
ace
be ween
habi a
bands
as
well
as
he
dep h
and
expanse
o
a
gi en
band.
Dis u bance
o
simila
scales
may
ha e
di e en
e ec s
on
wo
landscapes
o
dissimila
opog aphi-
cal
ea u es.
Hence,
e o s
should
be
made
owa d he
analysis
o
landscape
ea u es
as
hey
may
in luence
local
bio
ic
communi ies.
In
gene al,
I
p edic
ha
he
uppe
mon ane
zone
will
cons i u e
he
uppe
ele a ional
limi
o
maximum
ui
ba
di e si y.
As
o es s
a
lowe
ele a ions
disappea
and
he
lowe
edges
o
o es
p og essi ely
sh ink
upwa d
in
ele a-
ion,
we
may
see
shi s
in
he
egion
o
highes
di e si y
om
lowe
o
highe
o es
egions.
Howe e ,
his
"ele a-
ional
e ea "
may
each
i s
limi
when
condi ions
( ood,
oos ing,
clima ic)
o
he
en i onmen
necessa y
o
sup-
po ing
iable
popula ions
in
hemsel es
become
limi ing,
as
may
be
he
case
in
mossy
o es s.
While
his
p edic ion
esul s
om
s udies
associa ed
wi h
an h opogenic
des uc-
ion
o
o es s,
habi a
dis u bances
esul ing
om
na u al
ca as ophes
(e.g.,
hu icanes)
may
gene a e
simila
esul s.
One
di e ence
be ween
hese
wo
ypes
o
dis u bances
is
his:
Al hough
na u al
ca as ophes
do
no
ypically
gene -
a e
sus ained
des uc i e
s esses,
an h opogenic
p ocesses
ypically
do
so.
Fu u e
Conse a ion
Resea ch
Needs
Fu he
in en o ies
o
con inuous
habi a
g adien s
in
he
opics
a e
expec ed
o
demons a e
he
close
associa ion
be ween
o es
habi a
and
endemic
species.
In
his
con ex ,
compa isons
o
ele a ional
dis ibu ions
agains
a
backd op
o
changing
habi a
a e
use ul
as
p elimina y
indices
o
he
na u e
o
di e si y
pa e ns
and
he
ecological
p ocesses
ha
may
a ec
local
communi y
s uc u e.
A guably,
a
mo e
ho ough
analysis
o
ela ionships
be ween
habi a
agmen a ion
and
changes
in
species
di e si y
equi es
quan i a i e
measu emen s
o
habi a
dis u bance,
such
as
ela i e
a eal co e age,
spa ial
geome y,
and
ex en
o
edge
habi a s,
and
he
impac
on
local
ui
ba
assemblages.
Addi ionally,
he e
is
a
need
o
examine
he
ac ual
p o-
cesses
and
mechanisms
ha
unde pin
communi y
s uc u -
ing
and
co ela ions
among
species
di e si y,
popula ion
s uc u e,
and
habi a
quali y.
These
include:
(1)
iden i ica-
ion
and
quan i ica ion
o
hose
elemen s
o
he
habi a
o
di ec
impo ance
o
ui
ba s,
speci ically
ood
esou ces
and
oos s;
(2)
de e mina ion
o
c i ical
anges
o
en i on-
men al
condi ions
( empe a u e
and
humidi y)
ha
a e
physiologically
compa ible
wi h
he
pe sis ence
o
species
a
gi en
habi a s;
and
(3)
o e laying
spa ial
analyses
on
analyses
a
empo al
scales
o
assess
how
annual
o
seasonal
dynamics
o
ui
ba
ac i i ies
(e.g.,
ep oduc ion)
will
in luence
hei
esponses
o
habi a
changes
on
a
spa ial
scale.
Conclusions
The
po en ial
e ec s
o
habi a
agmen a ion
on
he
main-
enance
o
biodi e si y
a e
a ied
(see
Lande
1988
and
Te -
bo gh
1992
o
ecen
e iews).
Species
ex inc ions
a e
pos-
sible
end
esul s
o
hese
e ec s.
I
is
widely
belie ed
ha
such
ex inc ions
esul
om
demog aphic
s ochas ici y
a he
han
accumula ions
o
dele e ious
gene ic
changes
(Lande
1988).
In
he
inal
analysis,
bo h
demog aphic
and
gene ic
s ochas ici y
o en
become
ele an
only
a e
popu-
la ions
ha e
been
decima ed
o
in iable
numbe s.
Hence,
he
pi o al
issue
o
he
conse a ion
o
biodi e si y
is
he
p e en ion
o
declines
in
popula ion
sizes.
The
s udies
sum-
ma ized
in
his
chap e
s ongly
indica e
ha
changes
in
ui
ba
communi ies
can
occu
wi hin
he
b ie
ime
scale
in
which
o es
habi a s
a e
being
des oyed.
In
his
ega d,
i
is
appa en
ha
he
p e en ion
o
u he
o es
des uc-
ion
is
mos
c ucial
o
he
long- e m
main enance
o
ui
ba
di e si y.
Acknowledgmen s
Financial
suppo
o
he
1990
ield
esea ch
on
M .
Talinis
was
p o ided
by
he
Ame ican
Socie y
o
Mammalogis s
(g an s-in-aid
o
esea ch),
Ba
Conse a ion
In e na ional,
Inc.,
Chicago
Zoo-
logical
Socie y,
Ma shall
Field
and
Ellen
Tho ne
Smi h
Funds
o
he
Field
Museum
o
Na u al
His o y,
he
Lubee
Founda ion,
Inc.,
and
Bos on
Uni e si y.
I
hank
F.
Ca albas,
M.
Fu acan,
E.
Ma o,
and
L.
Tag-a
o
hei
aluable
assis ance
in
he
ield.
Many
o he
s udies
on
Philippine
ui
ba s
ci ed,
in
which
I
ha e
had
he
oppo uni y
o
collabo a e,
we e
suppo ed
by
he
U.S.
Na ional
352
R.
C.
B.
UTZURRUM
Science
Founda ion
(BSR-8514223),
he
John
D.
and
Ca he ine
T.
MacA hu
Founda ion,
and
he
Field
Museum
o
Na u al
His o y
(wi h
L.
R.
Heaney
as
RI.).
L.
R.
Heaney,
T.
H.
Kunz,
E.
A.
Ricka ,
and
J.
O.
Seamon
p o ided
aluable
commen s
on
his
manusc ip .
I
am
especially
hank ul
o
L.
R.
Heaney,
P.
D.
Heideman,
and
E.
A.
Ricka
o
p o iding
he
in ellec ual
a mosphe e
and
e-
sea ch
oppo uni ies
ha
ha e
shaped
my
in e es
in
Philippine
ba
esea ch.
Li e a u e
Ci ed
Alcala,
A.
C.
1976.
Philippine
Land
Ve eb a es:
Field
Biology.
New
Day
Publishe s,
Quezon
Ci y.
B own,
W
C.,
and
A.
C.
Alcala.
1970.
The
zoogeog aphy
o
he
he pe o auna
o
he
Philippine
islands,
a
inging
a chipelago.
P oceedings
o
he
Cali o nia
Academy
o
Science
38:105-130.
Dicke son,
R.
E.
1928.
Dis ibu ion
o
li e
in
he
Philippines.
Bu-
eau
o
Science
Monog aph
(Manila)
21:1-322.
Dickinson,
E.
C.,
R.
S.
Kennedy,
and
K.
C.
Pa kes.
1991.
The
bi ds
o
he
Philippines:
An
anno a ed
checklis .
B i ish
O ni holo-
gis s
Union
Checklis
12:1-507.
F ahm,
J.-R,
and
S.
R.
G ads ein.
1991.
An
al i udinal
zona ion
o
opical
o es s
using
b yophy es.
Jou nal
o
Biogeog aphy
18:
669-678.
Goodman,
S.
M.,
and
N.
R.
Ingle.
1993.
Sibuyan
Island
in
he
Philippines:
Th ea ened
and
in
need
o
conse a ion.
O yx
27:
174-180.
G ubb,
P.
J.,
and
T.
Whi mo e.
1966.
A
compa ison
o
mon ane
and
lowland
ain
o es
in
Ecuado .
II.
The
clima e
and
i s
e ec s
on
he
dis ibu ion
and
physiognomy
o
he
o es .
Jou nal
o
Ecology
54:303-333.
Hague,
P.,
J.
Te bo gh,
P.
Win e ,
and
J.
Pa kinson.
1986.
Conse -
a ion
p io i ies
in
he
Philippine
a chipelago.
Fo k ail
2:83-91.
Heaney,
L.
R.
1986.
Biogeog aphy
o
mammals in
SE
Asia:
Es i-
ma es
o
a es
o
coloniza ion,
ex inc ion,
and
specia ion.
In
Island
Biogeog aphy
o
Mammals,
L.
R.
Heaney
and
B.
D.
Pa e son,
eds.,
pp.
127-165.
Academic
P ess,
London.
Heaney,
L.
R.
1991a.
An
analysis
o
pa e ns
o
dis ibu ion
and
species
ichness
among
Philippine
ui
ba s
(P e opodidae).
Bulle in
o
he
Ame ican
Museum
o
Na u al
His o y
206:
145-167.
Heaney,
L.
R.
1991b.
A
synopsis
o
clima ic
and
ege a ional
change
in
Sou heas
Asia.
Clima ic
Change
19:53-61.
Heaney,
L.
R.
1993.
Biodi e si y
pa e ns
and
conse a ion
o
mammals
in
he
Philippines.
Asia
Li e
Sciences
2:261-270.
Heaney,
L.
R.,
and
P.
D.
Heideman.
1987.
Philippine
ui
ba s:
Endange ed
and
ex inc .
Ba s
5(1):
3-5.
Heaney,
L.
R.,
and
D.
S.
Rabo .
1982.
Mammals
o
Dinaga
and
Sia gao
islands,
Philippines.
Occasional
Pape s
o
he
Museum
o
Zoology
o
he
Uni e si y
o
Michigan
699:1-30.
Heaney,
L.
R.,
and
E.
A.
Ricka .
1990.
Co ela ions
o
clades
and
clines:
Geog aphic,
ele a ional,
and
phylogene ic
dis ibu ion
pa e ns
among
Philippine
mammals.
In
Ve eb a es
in
he
T opics,
G.
Pe e s
and
R.
Hu e e ,
eds.,
pp.
321-332.
Museum
Alexande
Koenig,
Bonn.
Heaney,
L.
R.,
and
R.
C.
B.
U zu um.
1991.
A
e iew
o
he
conse -
a ion
s a us
o
Philippine
land
mammals.
Associa ion
o
Sys-
ema ic
Biologis s
o
he
Philippines
Communica ions
3:1-13.
Heaney,
L.
R.,
P.
C.
Gonzales,
and
A.
C.
Alcala.
1987. An
anno a ed
checklis
o
he
axonomic
and
conse a ion
s a us
o
land
mammals
in
he
Philippines.
Silliman
Jou nal
34:32-66.
Heaney,
L.
R.,
P.
D.
Heideman,
and
K.
M.
Muda .
1981.
Ecological
no es
on
mammals
in
he
Lake
Balinsasayao
egion,
Neg os
O ien al,
Philippines.
Silliman
Jou nal
28:122-131.
Heaney,
L.
R.,
P.
C.
Gonzales,
R.
C.
B.
U zu um,
and
E.
A.
Ricka .
1991.
The
mammals
o
Ca anduanes
Island,
Philippines.
P o-
ceedings
o
he
Biological
Socie y
o
Washing on
104:399-415.
Heaney,
L.
R.,
P.
D.
Heideman,
E.
A.
Ricka ,
R.
B.
U zu um,
and
J.
S.
H.
Klompen.
1989.
Ele a ional
zona ion
o
mammals
in
he
cen al
Philippines.
Jou nal
o
T opical
Ecology
5:259-280.
Heaney,
L.
R.,
D.
S.
Bale e,
L.
Dola ,
A.
C.
Alcala,
A.
Dans,
P.
C.
Gonzales,
N.
Ingle,
M.
Lepi en,
W
Oli e ,
E.
A.
Ricka ,
B.
R.
Taba anza,
J .,
and
R.
C.
B.
U zu um.
1998.
A
synopsis
o
he
mammalian
auna
o
he
Philippine
islands. Fieldiana
Zoology
(new
se ies)
88:1-61.
Heideman,
P.
D.
1987.
The
ep oduc i e
ecology
o
a
communi y
o
Philippine
ui
ba s
(P e opodidae,
Megachi op e a).
Ph.D.
disse a ion,
Uni e si y
o
Michigan,
Ann
A bo .
Heideman,
P.
D.,
and
L.
R.
Heaney.
1989.
Popula ion
biology
and
es ima es
o
abundance
o
ui
ba s
(P e opodidae)
in
Philippine
submon ane
o es .
Jou nal
o
Zoology
(London)
218:565-586.
Ingle,
N.
R.,
and
L.
R.
Heaney.
1992.
A
key
o
he
ba s
o
he
Philippine
islands.
Fieldiana
Zoology
(new
se ies)
69:1-44.
IUCN
(In e na ional
Union
o
Conse a ion
o Na u e
and
Na u-
al
Resou ces).
1991.
P o ec ed
A eas
o
he
Wo ld:
A
Re iew
o
Na ional
Sys ems.
Vol.
1:
Indomalaya,
Oceania,
Aus alia,
and
An a c ica.
Island
P ess,
Washing on,
D.C.
Kumme ,
D.
M.
1990.
De o es a ion
in
he
pos -wa
Philippines.
Ph.D.
disse a ion,
Bos on
Uni e si y,
Bos on.
Lande,
R.
1988.
Gene ics
and
demog aphy
in
biological conse a-
ion.
Science
241:1455-1460.
Magu an,
A.
E.
1988.
Ecological
Di e si y
and
I s
Measu emen .
P ince on
Uni e si y
P ess,
P ince on.
Malla i,
N.
A.
D.,
and
A.
Jensen.
1993.
Biological
di e si y
in
no h-
e n
Sie a
Mad e,
Philippines:
Implica ion
o
conse a ion
and
managemen .
Asia
Li e
Sciences
2:101-102.
Micklebu gh,
S.
P.,
P.
A.
Racey,
and
A.
M.
Hu son,
eds.
1992.
Old
Wo ld
F ui
Ba s:
An
Ac ion
Plan
o
he
Family
P e opodidae.
IUCN
P ess,
Gland,
Swi ze land.
Mye s,
N.
1988.
En i onmen al
deg ada ion
and
some
economic
consequences
in
he
Philippines.
En i onmen al
Conse a ion
15:205-214.
Pe e son,
A.
T.,
and
L.
R.
Heaney.
1993.
Gene ic
di e en ia ion
in
Philippine
ba s
o
he
gene a
Cynop e us
and
Haplonyc e is.
Bio-
logical
Jou nal
o
he
Linnean
Socie y
49:203-218.
Ricka ,
E.
R.
1993.
Di e si y
pa e ns
o
mammals
along
ele a-
ional
and
di e si y
g adien s
in
he
Philippines:
Implica ions
o
conse a ion.
Asia
Li e
Sciences
2:251-260.
Ricka ,
E.
A.,
L.
R.
Heaney,
and
R.
C.
B.
U zu um.
1991.
Dis i-
bu ion
and
ecology
o
small
mammals
along
an
ele a ional
Main enance
o
F ui
Ba
Di e si y
353
ansec
in
sou heas e n
Luzon,
Philippines.
Jou nal
o
Mam-
malogy
72:458-469.
Ricka ,
E.
A.,
L.
R.
Heaney,
P.
D.
Heideman,
and
R.
C.
B.
U zu -
um.
1993.
The
dis ibu ion
and
ecology
o
mammals
on
Ley e,
Bili an,
and
Ma ipipi
islands, Philippines.
Fieldiana
Zoology
(new
se ies)
72:1-62.
Rickle s,
R.
E.
1979.
Ecology,
2nd
Ed.
Chi on
P ess,
New
Yo k.
Ruedas,
L.
A.,
J.
R.
Demboski,
and
R.
V
Sison.
1994.
Mo phologi-
cal
and
ecological
a ia ion
in
O op e opus
ca ilagonodus
Kock,
1969
(Mammalia:
Chi op e a:
P e opodidae)
om
Luzon,
Phil-
ippines.
P oceedings
o
he
Biological
Socie y
o
Washing on
107:1-16.
Te bo gh,
J.
1992.
Main enance
o
di e si y
in
opical
o es s.
Bio opica
24:283-292.
U zu um,
R.
C.
B.
1991.
Philippine
island
biogeog aphic
pa e ns:
P ac ical
applica ions
o
esou ce
conse a ion
and
manage-
men .
Associa ion
o
Sys ema ic
Biologis s
o
he
Philippines
Communica ions
3:19-32.
U zu um,
R.
C.
B.
1992.
Conse a ion
s a us
o
Philippine
ui
ba s
(P e opodidae).
Silliman
Jou nal
36:27-46.
Vincigue a,
L.
B„
and
R.
A.
Miille .
1993.
Neue
E kenn nisse
iibe
die
Ve b ei ung
des
Roh ennasen-Flughunds
Nyc imene
abon
Heaney
&
Pe e son,
1984
au
den
Philippinen
(Mammalia:
P e opodidae).
Jah buch
des
Na u his o isches
Museum
de
S ad
Be n
11:125-129.
Whi mo e,
T.
C.
1984.
T opical
Rain
Fo es s
o
he
Fa
Eas ,
2nd
Ed.
Ox o d
Uni e si y
P ess,
Ox o d.
