Conservation of bats on remote Indo-Pacific islands.

Ba
Biology
and
Conse a ion
Edi ed
by
Thomas
H.
Kunz
and
Paul
A.
Racey
SMITHSONIAN
INSTITUTION
PRESS
Washing on
and
London
23
Conse a ion
o
Ba s
on
Remo e
Indo-Paci ic
Islands
WILLIAM
E.
RAINEY
Oceanic
island
bio as
ha e
played
key
oles
in
bo h
he
heo y
and
p ac ice
o
conse a ion
biology.
They
p o-
ided
he
basis
o
he
heo y
o equilib ium
island
bio-
geog aphy
(MacA hu
and
Wilson
1967),
which
has
since
been
widely
applied
o
o he
discon inuous
habi a s.
They
u nish
some
o
he
bes -documen ed
examples
o
an h o-
pogenic
ex inc ions
and
in oduc ions
(Simbe lo
1995),
and
o e
si es
o
manageable
scale
whe e
expe imen al
in e en ion,
such
as
ansloca ion
o
p eda o
ex e mina-
ion,
can
be
unde aken
on
behal
o
endange ed
species
(Clou
and
C aig
1995).
While
esea ch and
managemen
e o s
on
emo e-
island
e eb a es
ha e
emphasized
endemic
land
bi ds
and
seabi ds,
he
less-di e se
ba communi ies
ha e
ecen ly
a -
ac ed
a en ion,
d i en
by
conce ns
abou
biodi e si y,
ecosys em
unc ion,
and
he
main enance
o
adi ional
human
esou ces
(Falan uw
1988;
Cox
e
al.
1991).
Th ee
ecen
compila ions
(Micklebu gh
e
al.
1992;
Wilson
and
G aham
1992;
Flanne y
1995)
co e ed
many
opics
ele an
o
ba
conse a ion
on
paleo opical
islands,
including
sys-
ema ics,
biogeog aphy,
human
u iliza ion,
and
he
s a us
o
species.
Addi ionally,
Micklebu gh
e
al.
(1992)
ha e
o -
e ed
a
p og am
o
conse a ion
ac ion
o
p e opodids.
The
ollowing
discussion
emphasizes
he
special
ci cum-
s ances
o
he
small,
la gely
endemic
ba
aunas
o
emo e
Indo-Paci ic
islands
bu
d aws
on
he
mo e-di e se
aunas
om
Sou heas
Asia
and
Aus alia
o
examine
speci ic
hemes.
Ba s
om
emo e
islands
a e
la gely
p e opodids
o
he
wide- anging
genus
P e opus,
mos
o
whose
membe s
a e
island
endemics
(Rainey
and
Pie son
1992).
E olu iona y
and
Ecological
Con ex
Among
mammals,
ba s
a e
ou lie s
in
e ms
o
maximum
li e
span
in
ela ion
o
body
size
(Aus ad
and
Fishe
1991).
Long
mass-speci ic
li e
spans
in
bi ds
and
olan
mammals
link
ligh
and
low
mo ali y
which,
in
u n,
a e
co ela ed
wi h
low
annual
ep oduc i e
ou pu
(Holmes
and
Aus ad
1995).
While
cons ain s
o
ligh
may
ha e
shaped
he
ep oduc i e
specializa ions
o
ba s
(Hayssen
and
Kunz
1996),
popula ions
a e
adap ed
o
low
p eda ion
and
low
s ochas ic
adul
mo ali y.
E olu iona y
esponses
o
e e-
b a e
popula ions
o
al e ed
selec ion
p essu es
on
islands
can
be
apid.
The
mos
nume ous
examples
in ol e
mo -
phological
change
(Smi h
e
al.
1995),
bu
Aus ad
(1993)
demons a ed
ha
an
island
popula ion
o
Vi ginia
opos-
326
Conse a ion
o
Remo e
Indo-Paci ic
Island
Ba s
327
sums
(Didelphis
i giniana),
wi h
low
p eda ion
o
abou
5,000
yea s,
had
bo h
educed
U e
size
and
educed
a es
o
senescence
ela i e
o
mainland
popula ions.
Assuming
e olu iona y
scope
o
e en
lowe
ep oduc i e
a e
in
island-dwelling
ba s,
such
adap a ions
could
heigh en
he
isk
o
ex inc ion
wi h
he
ab up
inc ease
in
p eda ion
ha
accompanies
human
coloniza ion
o
islands.
While
ends
in
body
size
e sus
island
a ea
a y
ac oss
ba
axa
(K zanowski
1967),
McNab
(1994)
demons a ed
a
gene al
end
owa d
smalle
body
size
and
lowe
mass-
speci ic
me abohc
a es
o
some
species
o
P e opus
on
small
islands.
He
no ed
ha
he
long- e m
pe sis ence
o
small
popula ions
is
highly
sensi i e
o
popula ion
size
and
ha
dec eased
indi idual
equi emen s
pe mi
la ge
popu-
la ions
o
esou ce-Hmi ed
species.
O ni hologis s,
in
pa -
icula ,
ha e
in ensi ely
explo ed
(wi hou
eaching
consen-
sus
on
cause)
an
"insula
synd ome"
o
inc eased
densi ies,
b oadened
habi a
o
die a y
niches,
and,
occasionally,
e-
duced
e i o iaU y
in
bi ds
on
islands
ela i e
o
he
same
o
ela ed
axa
on
la ge
adjacen
land
masses
(ThioUay
1993).
In e ac ions
among
di e en
species
a e
simpli ied
in
he
depaupe a e
communi ies
o
small
islands.
In
wha
a e
ypi-
cally
e olu iona ily
asymme ical
ela ionships,
small
guilds
o
gene alis s,
including
ugi o ous
and
nec a i o ous
ba s,
pol ina e
o
dispe se
seeds
o
nume ous
plan s
(Woodell
1979;
Cox
e
al.
1991).
The e
a e
no ably
ew
e eb a e
dispe se s
o
he
o en
la ge-seeded
ui s
o
island
canopy
ees.
Thus,
declines
o
ex inc ions
in
his
guild
aise
ques-
ions
abou
long- e m
impac s
on
o es
egene a ion
(Rainey
e
al.
1995),
and
abou
cascading
e ec s
on
o he
species,
including
humans,
whose
adi ional
subsis ence
sys ems
may
exploi
la ge-seeded
o es
ees
(Wiles
and
Fuji a
1992).
Noc u nal
ae ial
insec i o es
on
emo e
islands
a e
ew,
and
he
ecosys em
ole
o
insec i o ous ba s
in
hese
locales
is
li le
known.
Ex inc ions
and
Ex i pa ions
o
Ba s
on
Islands
A chaeological
Pe spec i es
on
Human
Impac
Accumula ing
a chaeological
da a
(Flanne y
1995;
S ead-
man
1995)
make
i
clea
ha
human
coloniza ion
o
smalle
oceanic
islands
in
he
Paci ic
was
accompanied
by
nume ous
ex inc ions
o
endemic
e eb a es
(la gely
land
bi ds)
and
local
ex i pa ions
o
indigenous
species.
As
human
popula-
ions
g ew
on
many
o
hese
islands,
aunas
we e
inc eas-
ingly
in luenced
by
widesp ead
in oduc ions
o
domes i-
ca ed and
commensal
e eb a es
and
by
al e a ions
in
ege a ion
s uc u e
and
composi ion,
especially
in
lowland
o es s
(S eadman
and
Role
1996).
Iden i ying
axa
ound
in
a cheological
deposi s
as
an h opogenic
ex inc ions
equi es
ca e ul
in e p e a ion.
Bones
may
be
p ehis o ical
impo s,
ag an s
(W agg
1995),
o
axa
ex i pa ed
by
en i onmen al
change
(Weisle
and
Ga ge
1993).
None heless,
models
o
biogeog aphic
p ocesses
ha
ea
human-in luenced
is-
lands
as
isola ed
excep ions
(Lack
1976;
Adle
1992)
mus
be
e iewed
ca e ully
agains
e idence
o
pe asi e
human
coloniza ion
and
ex ensi e
de auna ion
o
islands
in
he
opics
and
sub
opics.
Then,
as
now,
exploi a ion
o
island
wildli e
emphasized
subsis ence,
and
easily
ha es ed
o ms
declined
i s
(e.g.,
colonial
g ound-nes ing
seabi ds,
la ge
pigeons,
and
e es ial
o
ligh less
species).
Addi ionally,
cul u al
p e e ences
some imes
emphasized
pa icula
axa
(e.g.,
he
ed
ea he s
o
he
Rima a a
lo ikee ,
Vini
kuhlii)
and
con ibu ed
o
local
declines
(Wa ling
1995).
A cheological
e idence
o
educ ions,
ex i pa ions,
and
ex inc ions
o
ba
popula ions
on
Indo-Paci ic
islands
a e
summa ized
in
Table
23.1.
Al hough
a chaeological
su eys
ha e
e ealed
nume ous
ex inc ions
o
bi d
axa,
he e
is
only
one
eco d
o
ba
ex inc ion,
an
undesc ibed
mic ochi-
op e an
om
Maui,
Hawaii
(James
e
al.
1987).
The
eco d
does,
howe e ,
sugges
o
documen
declines
and
local
ex-
i pa ion
o
a
numbe
o
o he
axa.
Remains
o
ba s
om
he
wo
Eas
Paci ic
islands
a e
no
om
human-occupied
si es,
bu
adioca bon
da es,
s a ig aphy,
and
eco ds
o
o he
auna
a e
co ela ed
wi h
he
disappea ance
o
ex i -
pa ion
o
hese
species
wi h
human
coloniza ion
(S eadman
1986;
James
e
al.
1987,
pe sonal
communica ion).
The
ap-
pa en
pa e n
o
ex i pa ion
and
ex inc ion
may
be
biased
agains
de ec ion
o
small
e eb a es,
including
mic ochi-
op e an
ba s,
pa icula ly
a
human
occupa ion
si es.
Small
e eb a es
a e
less
likely
o
be
ha es ed,
and
hei
emains
a e
less
likely
o
su i e
ood
p ocessing,
consump ion,
and
ex ended
bu ial.
Addi ionally,
coa se
sc eens,
mo e
com-
monly
used
in
he
pas
o
sampling
a chaeological
deposi s,
ha e
selec ed
agains
eco e y
(Flanne y
e
al.
1988).
The
only
island
a chaeo auna
showing
loss
o
se e al
ba
species
is
on
'Eua,
Tonga
(Koopman
and
S eadman
1995).
The
ex i pa ed
species—P e opus
samoensis,
No op e is
mac-
donaldi,
and
Chae ephon
jobensis—a e
cu en ly
unknown
elsewhe e
in
he
Tongan
a chipelago,
al hough
all
a e
ex-
an
in
Fiji
(Flanne y
1995).
No op e is
macdonaldi
and
C.
jobensis
a e
p esen
in
Vanua u
and
P.
samoensis
in
Samoa.
P e opus
onganus
and
Emballonu a
semicauda a
ha e
been
eco e ed
om
p ehuman
s a a
on
'Eua,
and
bo h
species
pe sis
o
he
p esen
(Koopman
and
S eadman
1995).
The
p ehis o ic
ex i pa ion
o
N.
macdonaldi
and
C.
jobensis
may
be
a ibu ed
o
he
high
ulne abili y
o
ela i ely
la ge-
bodied,
agg ega ed,
ca e- oos ing
species
o
o e ha es ing
and
dis u bance.
Flanne y
(1995)
epo ed
ha
C.
jobensis
is
cu en ly
collec ed
o
ood
om
ca es
in
Fiji.
Al hough
in
328
W.
E.
RAINEY
Table
23.1
A chaeological
E idence
o
Reduc ions
in
Indo-Paci ic
Popula ions
o
Island
Ba s
Locali y
Species
Impac
Local
dis ibu ion
Commen s
Sou ce
Rod igues
Round
Island,
Mau i ius
Okinoe abu,
Ryukyu
Island
Ro a,
Ma ianas
Island
'Eua,
Tonga
Mangaia,
Cook
Island
Ai u aki,
Cook
Island
Ma'uke,
Cook
Island
Maui,
Hawaiian
Island
Flo eana
(=San a
Ma ia),
Galapagos
Island
P e opus
nige
P e opus
od icensis
P e opus
dasymallus
Emballonu a
semicauda a
P e opus
sa noensis
No op e is
macdonaldi
Chae ephon
jobensis
P e opus
onganus
P e opus
onganus
P e opus
onganus
Undesc ibed
espe ilionid
Lasiu us
bo ealis
P ehis o ic
ex i pa ion?
P ehis o ic
ex i pa ion?
P ehis o ic
ex i pa ion?
P ehis o ic
ex i pa ion
P ehis o ic
ex i pa ion
Declining
ha es
h ough
ime
P ehis o ic
ex i pa ion
P ehis o ic
ex i pa ion
P ehis o ic
ex inc ion
His o ic
ex i pa ion
Pe sis s
on
Mau i ius
Pe sis s
on
Rod igues
Ex an
elsewhe e
in
a chipelago
Ex an
elsewhe e
in
a chipelago
Absen
elsewhe e
in
Tonga;
p esen
in
adjacen
a chipelagoes
P esen
in
island
in e io
Ex an
elsewhe e
in
a chipelago
Ex an
elsewhe e
in
a chipelago
Remains
also
e-
co e ed
elsewhe e
in
a chipelago
Ex an
elsewhe e
in
a chipelago
Skull
in
ca e
wi h
ex inc
axa
Skull
in
bone
issu e
ill
Occupa ion
si e;
limi ed
emains
Occupa ion
si e;
limi ed
emains
Rap o
p ey
deposi s
and
occupa ion
si es;
wo
o he
ba
species
pe sis
o
p esen
Human
occupa ion
si e
Occupa ion
si e;
limi ed
emains
Occupa ion
si e
Ca e
ap;
bones
o
ex an
Lasiu us
cine eus
semo us
also
p esen
in
deposi
Ba n
owl
p ey
deposi ;
owl
ex i pa ed
mid-
1800s
Cheke
and
Dahl
1981
Cheke
and
Dahl
1981
Nishinakagawa
e
al.
1994
S eadman
1992
Koopman
and
S eadman
1995
Ki ch
e
al.
1992
S eadman
1991
Wal e
1990
(in
Koopman
and
S eadman
1995)
James
e
al.
1987;
H.
F.
James, pe sonal
communica ion
S eadman
1986
Samoa
oday
P.
samoensis
is
much
less
nume ous
han
P.
onganus
(C aig
e
al.
1994),
he
ex i pa ion
o
one
o
he
wo
la ge
(0.3-0.5
kg)
canopy- oos ing
ba s
on
'Eua
is
di icul
o
explain
wi hou
in oking
cul u al
p ey
p e e -
ence
o
se e e
p ehis o ical
educ ion
in p ima y
o es
(on
which
he
egional
endemic
P.
samoensis
appea s
mo e
de-
penden ).
Beha io al
obse a ions
(Wilson
and
Engb ing
1992;
Pie son
e
al.
1996)
o
P.
samoensis
and
P.
onganus
do
no
sugges
di e en ial
ulne abili y
o
hun ing
me hods
adi ionally
used
by
Polynesians.
His o ical
and
Recen
Ex inc ions
and
Ex i pa ions
Se e al
ba
species,
collec ed
on
Indo-Paci ic
islands
a e
he
onse
o
Eu opean
explo a ion,
a e
ei he
ex inc
o
locally
ex i pa ed
(Table
23.2).
Fo
hose
i s
collec ed
in
he
nine een h
cen u y,
he
eco d
is
equen ly
enigma ic.
The e
a e
usually
one
o
a
ew
specimens
wi h
unce ain
p o enance,
and
na u al
his o y
desc ip ions
p o ide
no
clues
on
he
cou se
and
causes
o
decline.
Simila
e en s
in
he
wen ie h
cen u y
a e
some imes
be e
documen ed,
so
ha
ecological
ai s
which
made
hem
(and
p esumably
o he
ex an
species)
ulne able
can
be iden i ied.
The
nine een h-cen u y
demise
o
P e opus
subnige
on
bo h
Mau i ius
and
Reunion
(wi h
he
la ge
P e opus
nige
ex i pa ed
on
Reunion,
bu
s ill
ex an on
Mau i ius)
is
well
documen ed
(Cheke
and
Dahl
1981).
Bo h
species
we e
ini ially
so
common
ha
plans
we e
made
o
expo
en-
de ed
ba
oil.
This
s udy
sugges ed
ha
he
ex inc ion
o
P.
subnige
was
linked
o
i s
unusual
oos ing
habi s.
While
mos
P e opus
species
o
which
oos ing
habi s
a e
known
cling
ex e nally
o
ee
b anches
(Pie son
and
Rainey
1992),
P.
subnige
oos ed
in
agg ega ions
as
many
as
400
indi idu-
als
in
ee
ca i ies
and
ock
c e ices,
whe e
i
was
highly
ulne able
o
collec ion
(Cheke
and
Dahl
1981).
Risks
o
his
species
likely
in ensi ied
as
mo e
o es
was
clea ed.
The
a e
o
P.
subnige
sugges s
we
should
pay
pa icula
a en ion
o
o he
species
wi h
simila
oos ing
habi s.
Re-
iew
o
museum
eco ds
and
ecen
ield
s udies
indica es
ha
ca i y
oos ing
may
be
mo e
common
han
was
p e-
iously ealized
among
smalle ,
li le-known
P e opus.
Fo
example,
P e opus
e ulus
on
New
Caledonia
is
one
o
se -
e al
ex an
species
ha
esembles
P.
subnige
in
ha ing
sho ,
u ed
ea s
which
ba ely
p o ude
abo e
a
dense,
woolly
pelage.
In o ma ion
on
specimen
ags
a
he
Ame ican
Mu-
seum
o
Na u al
His o y
indica es
se e al
P.
e ulus
we e
cap u ed
om
a
hollow
ee
limb
a
1,200
m
ele a ion.
Conse a ion
o
Remo e
Indo-Paci ic
Island
Ba s
329
Table
23.2
His o ical
Ex i pa ions
and
Ex inc ions
o
Indo-Paci ic
Island
Ba s
Da e
o
ex inc ion
o
Locali y
Species
ex i pa ion
P obable
cause
Commen s
Sou ce"
Mau i ius,
P e opus
subnige
ca.
1870,
1860
Hun ing;
habi a
Agg ega ed
ee
oos s
Cheke
and
Dahl
1981
Reunion
al e a ion
ulne able
Reunion
P e opus
nige
Ex i pa ed
Hun ing;
habi a
Pe sis s
on
Mau i ius
Cheke
and
Dahl
1981
be o e
1801
al e a ion
Reunion
Sco ophilus
bo bonicus
La e
19
h
Unknown;
habi a
Taxonomic
s a us
unce ain;
Cheke
and
Dahl
1981
cen u y?
al e a ion
ex ensi e
possible
species
pe sis s;
wo
o he
ex an
Mic ochi op e a
Okinawa,
Ryukyu
P e opus
ma iannus
A e
1849
Unknown;
habi a
Two
specimens;
Ande sen
1912;
Island
loochoensis
al e a ion
ex ensi e;
P.
dasymallus
pe sis s
K.
Koopman,
pe sonal
locali y
e o
communica ion;
possible
H.
Oh a,
pe sonal
communica ion
Palau
P e opus
pilosus
A e
1874
Unknown;
habi a
Two
specimens;
la ge
ba ;
Ande sen
1912;
Wiles
1997
ela i ely
in ac
P.
m.
pelewensis
pe sis s
Tobi,
Palau
P e opus
ma iannus
A e
1908
Hun ing?
Pe sis s
elsewhe e
in
Palau
Wiles
1997
pelewensis
Nendo,
Solomon
Nyc imene
ca.
1900
Unknown;
possibly
One
specimen;
no
Flanne y
1995
Island
sanc ac ucis
habi a
al e a ion
con empo a y
na u al
his o y
da a
Panay,
Philippine
Ace odon
luci e
A e
1892
Habi a
al e a ion;
Pe sis ed
in
pe iphe y
o
U zu um
1992
Island
hun ing
ag icul u al
landscapes
Addu
A oll,
P e opus
hypomelanus
A e
1922
Unknown;
possibly
a
One
specimen;
no
Hill
1958;
Holmes
1994
Maldi es
ma is
wai
o
locali y
e o
con empo a y
na u al
his o y
da a;
P.
gigan eus
a iel
pe sis s
Guam
P e opus
okudae
A e
1968
Unce ain;
habi a
Th ee
specimens
Wiles
1987a
al e a ion;
ba
hun ing
Neg os,
Dobsonia
chapmani
A e
1964
Fo aging
habi a
Mo ali y
om
guano
Heaney
and
Heideman
Philippine
clea ance;
hun ing;
mining
and
hun ing
1987
Island
oos
dis u bance
in
he
ca es
Kasho o
Island,
P e opus
dasymallus
A e
1986
Hun ing; habi a
—
H.
Oh a,
pe sonal
Taiwan
o mosus
al e a ion
communica ion
"Species
accoun s
o
all
excep
Sco ophilus
bo bonicus
a e
in
Micklebu gh
e
al.
(1992).
In
con as
o
ea lie
epo s
(Sanbo n
and
Nicholson
1950)
in
which
P e opus
o na us
was
common
and
P.
e ulus
a e,
Flanne y
(1995)
ecen ly
ound
ha
P.
e ulus
was
com-
mon
in
subcanopy
mis
ne
cap u es,
despi e
in ensi e
local
ba
hun ing.
Supposing
ha
ca i y
oos ing
is
common
o
his
species,
a
key
di e ence
om
he
Masca ene
species
may
be
ha
New
Caledonian
ba s
ha e
ecen ly
been
hun ed
by
spo ligh ing
and
shoo ing,
o en
a
nigh
om
ehicles
(A.
Baue ,
pe sonal
communica ion).
Wi h
his
me hod,
c yp ic
agg ega ed
oos s
and
s ic
noc u nali y
may
in
ac
educe
ulne abili y
o hun e s.
Flanne y
(1995)
also
epo ed
a
ee-ca i y
colony
o
abou
30
P e opus
ad-
mi ali a um
on
Malai a,
Solomon
Islands,
and
no ed
he
Moluccan
P e opus
caniceps
oos s
as
pai s
in
ee
hollows.
Flanne y
(1995)
also
sugges ed
ha
all
membe s
o
he
li le-s udied
genus
P e alopex
may
oos
in
ee
ca i ies.
He
epo ed
ha
an
unnamed
P e alopex
in lowland
New
Geo -
gia
and
Vangunu,
Solomon
Islands,
oos s communally
in
ca i ies
o
la ge
ees
and
is
easily
cap u ed
by
hand.
Flan-
ne y
also
desc ibed
how
his
species
was
obse ed
lying
away
om
a eas
o
logging
on
Kolombanga a
and
has
no
been
seen
he e
since
he
mid-1970s.
In
lowland
o es s
now
subjec
o
in ensi ying
imbe
managemen
o
o he
ac i i-
ies
ha
unca e
he
age
s uc u e
o
ees,
ba s
which
330
W.
E.
RAINEY
depend
on
la ge
hollow
ees
a e
ce ainly
mo e
ulne able
han
hose
ha
oos
in
canopy
oliage.
Ace odon
luci e ,
om
Panay,
Philippines,
was
las
col-
lec ed
in
1892
om
he
ma gins
o
ag icul u al
a eas
(U zu um
1992).
The
clea ance
o
land
on
Panay
has
now
educed
o es
co e
o
10%,
wi h
esidual
o es s
con ined
p ima ily
o
idges.
Ex ensi e
hun ing
and
habi a
loss
ha e
p esumably
led
o
he
ex inc ion
o
A.
luci e
(Micklebu gh
e
al.
1992).
Dobsonia
chapmani,
an
agg ega ed
ca e- oos ing
species,
was
s ill
common
on
sou he n
Neg os,
Philippine
Islands,
in
1964
(Heaney
and
Heideman
1987).
In
la e
su -
eys
hese
au ho s
concluded
ha
apid
clea ing
o
o ag-
ing
habi a
in
lowland
o es
nea
ca e
oos s,
along wi h
dis u bance
and
mo ali y
om
guano
mining
and
hun ing
in
he
ca es,
caused
ex inc ion
be o e
1981.
Reasons
o
he
ecen
decline
and
p esumed
ex inc ion
o
P e opus
okudae
on
Guam
a e
obscu e
(Wiles
1987a;
Mickle-
bu gh
e
al.
1992).
When
disco e ed,
his
small
species
was
e y
a e
ela i e
o
he
sympa ic
P e opus
ma iannus
ma ian-
nus.
Two
specimens
we e
collec ed
in
1930,
and
a
hi d
was
1
o
100
ba s
sho
du ing
he
hun ing
season
o
1968.
Remain-
ing
o es
habi a
on
Guam,
al hough
hea ily
al e ed,
could
p obably
s ill
suppo
many
mo e
han
he
se e al
hund ed
P.
m.
a iannus
cu en ly
on
he
island.
Rapid
popula ion
in-
c ease
o
Boiga
i egula is,
he
p eda o y
a bo eal
snake
in-
oduced
a e
Wo ld
Wa
II,
pos da es
he
disappea ance
o
P.
okudae
(Wiles
1987a;
Rodda
e
al.
1992).
This
species
is
no
known
om
he
adjacen ,
less
dis u bed
island
o
Ro a
o
elsewhe e
in
he
Ma ianas,
despi e
epea ed
su eys
(G.
Wiles
and
D.
Wo hing on,
pe sonal
communica ion).
P e opus
dasymallus
o mosus,
which
pe sis ed
un il
a
leas
1986
on
Kasho o,
an
o sho e
island
o
Taiwan,
is
appa -
en ly
he
mos
ecen
ex inc ion
o
P e opus.
Se e al
o he
subspecies
o
P.
dasymallus
in
he
Ryukyus
ba ely
su i e
(Micklebu gh
e
al.
1992;
H.
Oh a,
pe sonal
communica-
ion).
Fo es
co e
on
Kasho o
has
been
g ea ly
educed
by
ag icul u al
con e sion,
bu
he
ul ima e
cause
o
ex inc-
ion
is
likely
hun ing
o
ood.
Ku oda
(1933)
concluded
ha
his o ical
specimens
o
P.
d.
o mosus
om
locali ies
on
Tai-
wan
(Ande sen
1912)
o igina ed
elsewhe e,
bu
no
zooa -
chaeological
su eys
a e
a ailable
o
es
his
hypo hesis.
In en o y,
S a us Assessmen ,
and
Popula ion
Moni o ing
In en o y
Scien i ic
knowledge
o
many
ba
species,
pa icula ly
in
he
mo e
di e se
aunas
o
he
sou hwes
Paci ic,
has
o en
been
limi ed
o
hei
axonomic
desc ip ion
based
on
a
ew
specimens.
Ea ly
ba
collec ions
we e
equen ly
inciden al
o
bi d
sampling,
and
hus
in en o ies
we e
incomple e.
Wi h
accele a ing
habi a
al e a ion
in
ecen
decades,
he
con inued
exis ence
o
species
on
small
islands,
especially
hose
dependen
on
lowland
o es
a eas,
has
been
an
open
ques ion.
Pa ly
om
conce n
o
he
po en ial
loss
o
bio-
di e si y,
he
museum
adi ion
o
aunal
in en o ies
has
been
enewed,
especially
by
he
Aus alian
Museum,
he
Wes e n
Aus alian
Museum,
and
egional
collabo a o s.
F om
hese
su eys
Flanne y
(1995)
has
epo ed
ecen
collec ions
o
many
o
he
li le-known
P e opus
om
he
sou hwes
Paci ic
(e.g.,
P.
ch ysop oc us,
P.
junda us,
P.
maha-
ganus,
P.
melanopogon,
P.
ni endiensis,
P.
ocula is,
P.
pohlei,
P.
ayne i
cogna us,
and
P.
emmincki).
These
su eys
also
yielded
wo
new
species
(one
discussed
ea lie )
o
P e alopex
om
he
Solomon
Islands
(Flanne y
1991,
1995).
On
Gua-
dalcanal,
P e alopex
pulch a
is
unusual
in
ha
i
appea s
e-
s ic ed
o
high,
mossy,
mon ane
o es ,
whe e
i
eplaces
he
(now
declining)
lowland
species
P e alopex
a a a.
The
we ,
high-ele a ion
habi a
is
analogous
o
ha
o
he
a e
Fijian
endemic,
P e alopex
ac odon a
(Hill
and
Beckon
1978;
also
ecollec ed
by
Flanne y
1995).
In
Fiji,
howe e ,
no
lowland
P e alopex
species
is
known.
S a us
Assessmen s
and
Popula ion
Moni o ing
Se e al
su eys
ha
ha e
assessed
he
s a us
o
ba
popula-
ions
pos da e
he
megachi op e an
compila ion
by
Mickle-
bu gh
e
al.
(1992).
A
1991
su ey
o
ba s
in
Palau
ound
ha
P e opus
ma iannus
pelewensis,
o me ly
e y
hea ily
hun ed
o
comme cial
expo
o
he
Ma ianas,
was
common
o
abundan
a
40%
o
54
e ening
census
locali ies
and
ha
P e opus
pilosus
is
almos
ce ainly
ex inc
(Wiles
e
al.
1997).
G an
(1994)
obse ed
P e opus
onganus
on
he
isola ed
is-
land
o
Niue,
and
ob ained
local
es ima es
ha
1,200-5,000
indi iduals
a e
aken
annually
by
shoo ing,
sugges ing
a
much
la ge
popula ion
han
p e iously
epo ed.
In
e-
iewing
he
dis ibu ion
and
cu en
s a us
o
ba s
known
om
Tonga,
Koopman
and
S eadman
(1995)
epo ed
a
leas
2,400
P.
onganus
on
'Eua
and
se e al
housand
a
Kolo ai,
Tonga apu,
in
1988.
Holmes
(1994)
desc ibed
P e opus
gigan eus
a iel
in
he
Maldi es
as
widesp ead,
bu
no
nume ous,
and
subjec
o
le hal
con ol
e o s
in ended
o
educe
ui
dep eda ion.
Su eys
conduc ed
o e
se e al
yea s
o
P e opus
li ing-
s onii,
he
a e
o
he
wo
P e opus
species
in
he
Como es,
yielded
minimum
popula ion
es ima es
o
380
on
Anjouan
and
60
on
Moheli
(W
T ewhella,
pe sonal
communica ion).
Reason
and
T ewhella
(1994)
no ed ha
he
p ima y
h ea
o
his
species
is
habi a
loss
as
o es
agmen s
a e
con-
e ed
o
ag icul u al
use
o
suppo
a
apidly
g owing
human
popula ion.
P e opus
li ings onii
cu en ly
oos s
in
Conse a ion
o
Remo e
Indo-Paci ic
Island
Ba s
331
he
canopy
o
mon ane
o es
pa ches
on
s eep
slopes
and
elies
mo e
ex ensi ely
on
na i e
ui s
o
ood
han
he
o he
P e opus
species,
P e opus
seychellensis
como ensis.
Ne -
ing
e o s
o
collec
P.
li ings onii
o
an
ex
si u
cap i e
b eeding
p og am
inciden ally
cap u ed
mo e
han
150
Rouse us
obli iosus,
a
hi d
li le-known
p e opodid
(Reason
e
al.
1994).
In
mo e
a luen
Ame ican
Samoa,
human
popula ion
g ow h
is
compa ably
apid,
de o es a ion
o
limi ed
low-
lands
is
well
ad anced,
and
clea ing
o
s eepe
slopes
is
ongoing.
Howe e ,
he
majo
ac o
a ec ing
ecen
popu-
la ion
ends
o
P.
onganus
and
P.
samoensis
has
been
mo -
ali y,
la gely
om
hun ing,
a he
han
habi a
limi a ion.
C aig
e
al.
(1994)
has
summa ized
he
esul s
o
oos
coun s
and
diu nal
su eys,
which
began
in
1987
on
Tu uila
a e
local
legisla ion
was
enac ed
o
hal
comme cial
ex-
po
o
ba s
o
he
Ma ianas
(see
ollowing).
These
au ho s
showed
ha
ba
coun s
d opped
p ecipi ously
ollowing
a
cyclone
in
1990.
While
some
ba s
died
o
s a ed,
inc eased
oppo unis ic
hun ing
was
he
p ima y
cause
o
pos cy-
clone
mo ali y.
Hun e s
epo ed
double
he
ypical
annual
ha es
in
he
yea
ollowing
ha
s o m.
A
second
se e e
cyclone
less
han
2
yea s
la e
compounded
e ec s
on
ba
popula ions.
C aig
e
al.
(1994)
es ima ed
an
80%-90%
popula ion
decline
o
bo h
species
o e
5
yea s
and
en a i ely con-
cluded
ha
200-400
P.
samoensis
and
1,500-2,500
P.
onganus
we e
on
he
island
in
la e
1992.
Diu nal
oos
coun s
o
P.
onganus
on
Tu uila
ha e
shown
a
s eady
inc ease
om
1,700
in
1991
o
5,700
in
1996.
Radio acking
indica es
ha
diu nal
ligh
obse a ions
unde es ima e
P.
samoensis
num-
be s, which
may
be
close
o
1,000
(A.
B ooke,
pe sonal
communica ion).
Pe haps
pa ly
as
a
consequence
o
con-
inued
illegal
hun ing,
oos s
o
bo h
species
a e
concen-
a ed
on
s eep
o es ed
slopes
in
unde eloped
a eas.
A
subs an ial
a ea
o
o es
is
con ained
wi hin
he
ecen ly
es ablished
Na ional
Pa k
o
Ame ican
Samoa,
bu
his
may
no
be adequa e
o
main ain
iable
popula ions
o
ba s
and
o he
wildli e
i
o es
clea ance
in
o he
a eas
con inues
a
he
cu en
a e.
A
s udy
conduc ed
in
Wes e n
and
Ame ican
Samoa
ound
di e ences
be ween
he
wo
P e opus
species
in
e-
sponse
o
a
pos cyclone
educ ion
in
ood
a ailabili y
(Pie -
son
e
al.
1996).
The
less
abundan ,
egionally
endemic
P.
samoensis
emained
in
he
o es ,
ini ially
eeding
la gely
on
lea es,
while
he
mo e
widesp ead
P.
onganus
o aged
ex-
ensi ely
on
esidual
o
allen
ui
in
inhabi ed
a eas
and
expe ienced
highe
mo ali y
om
human
hun e s
and
p e-
da ion
by
domes ic
animals.
This
s udy
ound
ha
o es
ese es
on
Sa ai'i,
Wes e n
Samoa,
appea ed
o
p o ec
small
numbe s
o
P.
samoensis
because
hei
limi ed
diu nal
o aging
mo emen s
emained
wi hin
he
ese e
whe e
hun ing
was
banned.
P e opus
onganus
equen ly
lew
ou -
side
he
ese e
o
eed
and
was
likely
subjec
o
hea ie
hun ing
p essu e.
The
emnan
P.
ma iannus
ma iannus
popula ion
on
Guam,
usually
ound
oos ing
as
a
single
colony
on
a
U.S.
mili a y
base
(a
de
ac o
ese e),
has
been
moni o ed
since
1981
(Wiles
e
al.
1995).
These
au ho s
epo ed
seasonal
luc ua ions
in
popula ion
size,
wi h
200-400
animals p es-
en
om
June
o
Sep embe
and
400-750
om
No embe
o
Feb ua y.
Wiles
and
Glass
(1990)
epo ed
obse a ions
o
o sho e
ba
ligh s
among
he
Ma ianas
and
used
changes
in
colony
size
o
in e
in e island
g oup
mo emen s,
including
se e al
(60
km
o e
he
ocean)
be ween
Guam
and
Ro a
in
he
Commonweal h
o
he
No he n
Ma ianas
(CNMI).
Wiles
e
al.
(1995)
sugges ed
ha
he
seasonal
luc ua ions
in
ba
numbe s
on
Guam
e lec
annual
mo emen
be ween
he
wo
islands.
P eda ion
on
non olan
ju enile
ba s
by
he
in oduced
snake
Boiga
i egula is
appa endy
p e en s
local
ec ui men
(Wiles
1987b),
so
ha
he
pe sis ence
o
he
Guam
popula ion
could
depend
on
mo emen
om
Ro a.
His o ically,
ba s
we e
hea ily
hun ed
on
Guam,
bu
poach-
ing
has
no
been
a
h ea
o
he
esiden
ba
colony
in
ecen
yea s
(G.
Wiles,
pe sonal
communica ion).
Guam
is
an
un-
usually
a luen ,
egional
anspo a ion
hub
wi h
apidly
expanding
ou is
acili ies
and
ela ed
in as uc u e.
In
1993,
he
U.S.
go e nmen
c ea ed
a
wildli e
e uge
om
ela i ely
in ac
o es
on
mili a y lands.
While
his
has
me
wi h
some
local
suppo ,
he e
has
also
been
conside able
esis ance,
based
pa ly
on
p e-Wo ld
Wa
II
p i a e
land
claims
o
hese
a eas
(Wiles
1994).
In
addi ion
o
de el-
opmen
p essu es,
emaining
o es
a eas
a e
subjec
o
on-
going
deg ada ion
by
in oduced
ungula es
(Wiles
e
al.
1995).
Ro a
ha bo s
he
only
subs an ial
popula ion
o
P.
ma i-
annus
in
he
sou he n
islands
o
he
CNMI
(S inson
e
al.
1992).
Es ima es
o
ba
numbe s
on
Ro a
om
1986
un il
a
majo
cyclone
in
Janua y
1988
we e
2,000-2,500.
Sub-
sequen
coun s
d opped
o
app oxima ely
1,000,
owing
o
pos cyclone
hun ing
and
possibly
emig a ion.
The
popula-
ion
has
emained
a
his
lowe
le el
h ough
1995,
p obably
because
o
ch onic
illegal
hun ing
(Wo hing on
and
Taisa-
can
1996).
Wiles
e
al.
(1989)
su eyed
he
less-de eloped
islands
no h
o
Saipan
and
es ima ed
ha
7,450
ba s
we e
p esen
in
1983.
Illegal
ma ke
hun ing
has
since
inc eased.
Also,
a
su ey
o
Ana ahan
iden i ied
apidly
g owing
e al
goa
popula ions
as
a
se ious
new
h ea
o
o es
habi a
(Ma shall
e
al.
1995).
The
CNMI
has
a
egula o y
ame-
wo k
o
achie ing
sus ainable
ba
ha es ,
bu
public
and
ins i u ional
esis ance
o
es ic ions
on
ba
hun ing
and
ade
a e
s ong.
Habi a
conse a ion
a eas
ha e
been
ou -
332
W.
E.
RAINEY
lined,
bu
en o cemen
o
local
wildli e
laws
emains
mini-
mal.
Wiles
and
Glass
(1990)
sugges ed,
as
a
way
o
a ional-
ize
he
a ied
axonomic
and
egula o y
s a us
o
popula-
ions
on
di e en
islands,
ha
all
P.
ma iannus
in
he
sou he n
Ma ianas
should
be
managed
as
a
single
uni .
Recen
epo s
o
Emballonu a
semicauda a,
he
only
mi-
c ochi op e an
ba
in
much
o
Mic onesia
and
Polynesia,
sugges
ha
a
pa chy,
angewide
decline
is
in
p og ess.
A
he
eas e n
limi
o
i s
cu en
ange
in
Polynesia,
G an
e
al.
(1994)
epo ed
a
apid
educ ion
o
E.
semicauda a
o
only
a
ew
indi iduals
in
Ame ican
Samoa.
They
no ed ha
simila
declines,
om
hund eds
o
pe haps
housands
o
ba s
o
isola ed
indi iduals,
seem
o
ha e
occu ed
con em-
po aneously
o
colonies
on
Upolu,
Wes e n
Samoa.
These
declines
lag
by
decades
simila
changes
o
popula ions
o
E.
semicauda a
and
ca e
swi le s
(Collocalia
aniko ensis)
in
Guam
and
adjacen
islands
in
he
CNMI
(Lemke
1986;
S eadman
1992).
In
Samoa,
howe e ,
swi le
popula ions
ha
sha e
ca es
wi h
E.
semicauda a
ha e
emained
s able
o
inc eased.
G an
e
al.
(1994)
discussed
he
ole
o
a
se e e
cyclone
in
he
educ ion
o
E.
semicauda a
popula ions
in
Ame ican
Samoa,
bu
did
no
link
he
long- e m
declines
o
he
ex-
plana ions
o e ed
o
he
Ma ianas—mili a y
des uc ion
o
ca es,
ex ensi e
pes icide
use
o
ec o
con ol,
and
guano
mining.
Recen
su eys
in
he
Ma ianas
ha e
gen-
e ally
con i med
a
pa e n
epo ed
by
Lemke
(1986)
sug-
ges ing
ha
E.
semicauda a
is
ex inc
on
Guam
and
Ro a
and
absen
on
all
o he
islands,
excep
Aguiguan
(no
hu-
man
inhabi an s)
and
pe haps
Saipan
(Wiles
e
al.
1995;
Wo hing on
and
Taisacan
1996).
Island-by-island
pa e ns
o
pe sis ence
o
ba
and
swi le
popula ions
in
he
Ma i-
anas
a e
also
no
cong uen
(D.
Wo hing on,
pe sonal
communica ion).
Flanne y
(1995)
ela ed
a
long- e m
esiden 's
epo
o
simila
declines
o
E.
semicauda a
on
Vi i
Le u,
Fiji,
om
abundance
in
he
1950s
o
i ual
absence
in
he
1990s.
The
in o man
a ibu ed
declines
o
oos
dis u bance
and
he
bu ning
o o es s
nea
ca es.
Flanne y
(1995)
also
epo ed
a
small
colony
o
E.
semicauda a
esiding
in
an
inaccessible
ca e
oos
on
Ta euni,
Fiji,
p esumably
in
he
1990s.
This
species
may
ha e
also
declined
on
he
isola ed
island
o
Ro uma,
Fiji.
Clunie
(1985)
epo ed
he
ba s
as
p esen
in
mul iple
housands
in
ca es
and
obse ed
o age s
"in
a
la ge
numbe s
han
is
usual
in
Fiji."
In
1993-1994,
Cox
(pe sonal
communica ion)
isi ed
Ro uman
ca es
se e al
imes
and
no ed
a
ma ked
educ ion
in
numbe s,
consis en
wi h
epo s
o e ed
by
local
esiden s.
In
su eys
o
mo e
han
50
ca es
on
'Eua,
Tonga,
in
1988-1989,
only
a
single
colony
o
25
E.
semicauda a
was
obse ed
(Koopman
and
S eadman
1995).
In
se e al
o he
a eas
o
Mic onesia,
E.
semicauda a
has
been
ecen ly
p esen .
Wiles
and
Con y
(1990)
epo ed
a
ca e
colony
o
200
in
Palau.
Obse a ions
a
mul iple
si es
in
Palau
in
1991
showed
he
species
o
be
widesp ead
and
common,
wi h
o aging
mo emen s
by
se e al
housand
ba s
obse ed
(G.
Wiles,
pe sonal
communica ion).
In
1989,
ens
o
o aging
E.
semicauda a
and
swi le s
we e
also
seen
a
dusk
o e
he
s ee s
on
Moen
Island,
Chuuk,
bu ,
as
obse ed
ea lie
by
B une
and
P a
(1979),
his
species
is
conside ably
less
common
on
Pohnpei
(Rainey,
unpub-
lished
obse a ions).
A
common
cause
o
hese
seemingly
pa allel
declines
is
no
e iden .
Bo h
he
sou he n
Ma ianas
(wi h
ew
ba s)
and
Palau
(wi h
many)
sha e
a
his o y
o
in ense
mili a y
ac i i y
du ing
Wo ld
Wa
II
and
o me
use
o
DDT
o
insec
con ol
(Bake
1946),
bu
he
Ma ianas
ha e
ew
ba s
and
Palau
has
many.
Unlike
he
Ma ianas,
Palau
has
nume -
ous
small
ou lying
ka s ic
islands
ha
likely
o e
mo e
ex ensi e
ba
e uges.
Ch onic
oos
dis u bance
is
a
well-
documen ed
cause
o
declines
in
ca e-dwelling
ba s,
espe-
cially
a
si es
close
o
concen a ions
o
humans
and
domes-
ic
o
e al
animals.
Colonies
o
E.
semicauda a
pe sis ,
howe e ,
close
o
habi a ions
on
densely
popula ed,
hea ily
al e ed
Moen,
Chuuk,
and
a e
declining
on
much
la ge ,
less
densely
popula ed
islands
such
as
Vi i
Le u,
Fiji,
and
Upolu,
Wes e n
Samoa.
In oduced
pa hogens,
which
a e
appa en ly
esponsible
o
declines
o
na i e
Hawaiian
bi ds
in
ela i ely
in ac
habi a
(A kinson
e
al.
1995),
o e
an-
o he
possible
mechanism.
Resou ces
o
wildli e
conse a-
ion
o en
ocus
on
cha isma ic,
endemic
" lagship"
species,
bu
he
egional
decline
o
his
small
insec i o ous
ba
de-
se es
ca e ul
sc u iny.
Sys ema ics
and
Molecula
Gene ics
In
he
1980s,
use
o
p o ein
elec opho esis
in
su eys
o
Aus alian
Mic ochi op e a
e ealed
o
con i med
nume -
ous
well-di e en ia ed
sibling
species
and
some imes
con-
side able
geog aphic
di e en ia ion
wi hin
species
(Adams
e
al.
1982;
Ba e s ock
e
al.
1987).
Mo e
ecen
popula ion-
and
species-le el
su eys
o
Paci ic
p e opodids,
using
se -
e al
molecula
echniques,
o e ed
con as ing
esul s
wi h
bo h
e olu iona y
and
managemen
implica ions.
Fo
ex-
ample,
p o ein
elec opho esis
o
Cynop e us
popula ions
in
Indonesia
e ealed
ela i ely
small
gene ic
dis ances
among
species
and
li le
e idence
o
geog aphic
di e en ia ion
wi hin
islands,
sugges ing
he
ocean
is
he
p ima y
ba ie
o
gene
exchange
(Schmi
e
al.
1995).
Pe e son
and
Heaney
(1993)
compa ed
Haplonyc e is
ische i,
a
Philippine
endemic
es ic ed
o
p ima y
o es ,
and popula ions
o
he
wide- anging
Cynop e us
b achyo is
in
he
Philippines
Conse a ion
o
Remo e
Indo-Paci ic
Island
Ba s
333
and
no ed
highe
le els
o
gene ic
di e en ia ion
among
popula ions
o
H.
ische i.
In
con as ,
Ki chene
e
al.
(1993)
used
elec opho e ic
analysis
o
examine
a ia ion,
on
se -
e al Indonesian
islands,
in
Ae halops
alec o,
a
small
p e opo-
did
es ic ed
o
mon ane
o es .
They
concluded
ha
popu-
la ions
we e
nei he
well
di e en ia ed
no
gene ically
isola ed.
An
ex ensi e
s udy
o
he
highly
mig a o y
Aus alian
lying- ox,
P e opus
scapula us,
using
bo h
allozymes
and
an-
dom
ampli ied
polymo phic
DNA,
showed
low
le els
o
popula ion
s uc u e
and
indica ed
ha
his
species,
now
independen ly
managed
by se e al
ju isdic ions,
was
e ec-
i ely
panmic ic
(Sinclai
e
al.
1996).
Webb
and
Tidemann
(1996)
epo ed
simila
esul s o
P e opus
alec o
and
P e opus
poliocephalus
and
sugges ed
hey
should
be
managed
as
mi-
g a o y
species.
Thei
analysis
o
pu a i e
hyb ids
om
a -
eas
o
sympa y
e ealed
only
one
o
wo
ixed
alleles
in
23
loci
di e en ia ing
P.
alec o
om
P.
poliocephalus
o
P e opus
conspicilla us,
which
sugges s
ela i ely
ecen
di e gence
among
hese
species
(Webb
and
Tidemann
1995).
The
low
le els
o
in aspeci ic
di e en ia ion
in
P e opus
and
Cynop e us
sugges
ha
popula ions
o
P e opus
(o
o he
la ge-
o
mode a e-sized
unspecialized
p e opodids)
on
small
oceanic
islands
can
gene ally
be
iewed
as
single
managemen
uni s.
The
in e ence
ha
P.
ma iannus
egu-
la ly
c osses
as
much
as
60
km
o
ocean
(Wiles
and
Glass
1990;
Wiles
e
al.
1995)
highligh s
he
need
o
use
molecula
app oaches
o
iden i y
app op ia e
managemen
uni s
in
species
ha
lack
ob ious
mo phological
discon inui ies
and
whose
anges
encompass
mul iple
islands
sepa a ed by
signi ican
wa e
gaps.
While
he
desc ip ion
o
new
species
and
subspecies
o
smalle
p e opodid
and
mic ochi op e an
ba s
om
he
mo e
di e se
communi ies
o
he
wes e n
Paci ic
has
con-
inued
o
he
p esen
(Ki chene
e
al.
1994),
highe -le el
megachi op e an
sys ema ics
has
emained
essen ially
ha
o
Ande sen
(1912),
which
is
inexplici
by
mode n
s an-
da ds.
Micklebu gh
e
al.
(1992)
p esen ed
a
conse a ion
anking
scheme
o
geog aphic
a eas
ha
inco po a ed
weigh s
o
bo h
" axonomic
dis inc i eness"
and
species
ichness
using
a
mo phologically
based
cladog am
o
megachi op e an
gene a
(by
J.
E.
Hill;
bu
see
also
Heaney
1991).
Mo e
ecen ly,
Colgan
and
Flanne y
(1995),
Ki sch
e
al.
(1995),
and
Sp inge
e
al.
(1995)
p esen ed
independen
molecula
da a
se s
on
megachi op e an
sys ema ics.
While
hese
ine i ably
di e ,
hey
a e
consis en
wi h
some
ea lie
mo phological
and
biochemical
s udies
(Hood
1989;
Haiduk
1983)
in
one
majo
inding:
he
mo phologically
specialized
nec a - eeding
gene a,
his o ically
g ouped
in
he
sub amily
Mac oglossinae,
a e
no
a
clade.
Wi h
new
ools
and
enewed
in e es
in
his
opic,
we
can
expec
a
g adual
consensus
on
phylogene ic
ela ionships
ha
will
p o ide
new
c i e ia
o
conse a ion
anking
schemes.
Th ea s
o
Ba
Popula ions
Among
declining
e eb a e
species,
one
can
make
a
heu-
is ic
di ision
be ween
hose
ha
a e
los
wi h
hei
habi a
and
hose
ha
anish
long
be o e
hei
habi a .
The
la e
a e
usually
animals
ha
a e
ei he
la ge
(e.g.,
si enians)
o
a e
selec i ely
hun ed
o
hei
pe cei ed
high
alue
o
hu-
mans
(e.g.,
musk
dee ,
bi ds
wi h
unusual
plumes).
Al-
hough
ba s
ha e
ela i ely
small
body
mass
and
migh
be
expec ed
o
closely
ack
habi a
loss,
low
ep oduc i e
ou -
pu
and,
o
some
species,
agg ega ed
oos ing
habi s
a
adi ional
si es
make
hem
ulne able
o
selec i e
hun ing.
Loss
o
deg ada ion
o
o es
habi a
is
clea ly
an
impo an
h ea
o
he
long- e m
pe sis ence
o
ba
popula ions
on
islands.
The
ela i e
impo ance
o
habi a
loss,
exploi a-
ion,
and
o he
ac o s
in
de e mining
popula ion
ends
is
howe e
qui e
a ied
among
islands
and
gene ally
is
poo ly
known.
A
ew
o
he
mo e
disc e e
h ea s
o
island
ba
popula ions
a e
discussed
nex .
In e na ional
T ade
Pa ly
as
a
consequence
o
he
la ge
body
size
o
some
Megachi op e a
(and
he
limi ed
a ay
o
animals
a ailable
on
isola ed
oceanic
islands),
human
consump ion
o
ba s
is
a
signi ican
ac o
a ec ing
ba
popula ions
on
Indo-Paci ic
islands
and
in
adjacen
a eas
o
Asia.
A i udes
owa d
ba s
as
ood
a y
ac oss
eligions,
cul u es,
and
geog aphy
(Ki ch
and
Yen
1982;
Fuji a
and
Tu le
1991).
In
Mic onesia,
o
example,
esiden s
o
bo h
he
Ma ianas
and
Yap
ha e
a
long
adi ion
o
consuming
ba s.
In
he
Ma ianas,
hey
a e
highly
a o ed
(Sheeline
1991),
while
in
Yap
hey
a e
"no
es eemed"
(Falan uw
and
Manmaw
1992).
To
he
eas
in
Chuuk
and
Pohnpei,
ba s
a e
iewed
as
somewha
epellen
and
a e
gene ally
no
pa
o
he
local
die
(Rainey,
unpub-
lished
obse a ions).
The
majo
ocus
o
in e na ional
ade
in
ba s
(p ima ily
P e opus
spp.)
has
been
Guam
and
he
adjacen
CNMI.
Wiles
(1992)
summa ized
ade
his o y
and
abula ed
legal
impo s,
showing
ha
subsequen
o
local
deple ion
ba s
we e
ini ially
acqui ed
om
nea by
islands.
In
he
1980s
he
adius
o
ade
expanded,
eaching
as
a
as
Papua
New
Guinea
and
he
Philippines.
The
mean
numbe
o
ba s
eco ded
as
impo ed
o
Guam
in
1981-1989
was
oughly
13,000
annually
(Wiles
1992).
The
mean
numbe
o
ba s
impo ed
annually
in o
he
CNMI
in
1986-1989
was
3,300
(S inson
e
al.
1992).
In
some
ins ances,
small
islands
such
as
Yap
and
Ame ican
Samoa
no ed
apid
declines
in
P e opus

334
W.
E.
RAINEY
popula ions
and
imposed
local
es ic ions
on
expo
hun -
ing
(C aig
and
Sy on
1992;
Falan uw
and
Manmaw
1992).
In
1989,
se en
cen al
and
wes
Paci ic
small
island
P e opus
species
we e
added
o
Appendix
I
o
he
Con en-
ion
on
T ade
in
Endange ed
Species
o
Wild
Flo a
and
Fauna
(CITES),
while
he
emaining
species
o
P e opus
and
all
species
o
he
allied
genus
Ace odon
we e
placed
on
Ap-
pendix
II
(B aii igam
and
Elmq is
1990).
This
lis ing
obliged
coun ies
who
we e
pa ies
o
he
ea y
(including
he
Uni ed
S a es)
o
cease
in e na ional
ade
in
Appendix
I
species
and
moni o
ade
in
Appendix
II
axa.
A e
en-
o cemen
o
CITES
p o isions
on
Guam
in
1990,
legal
impo s
om
1990-1993
came
om
Palau,
which
was
s ill
unde
U.S.
ju isdic ion
(annual
mean:
o
Guam,
7,688
ba s;
o
CNMI,
5,755)
(Wiles
e
al.
1997).
This
p ac ice
con in-
ued
un il
la e
1994
when
Palauan
independence
ended
legal
ade
in
P e opus
ma iannus
pelewensis.
Also
in
1994,
Ace odon
juba us
and
Ace odon
luci e
( he
la e
p obably
ex inc )
we e
ans e ed
o
Appendix
I,
based
on
a
pe i ion
om
he
Philippines,
which
emphasized
ongoing
illegal
ade
in o
Guam
and
CNMI.
Subsequen
o
closu e
o
he
legal
ade
om
Palau,
documen ed
impo s
ha e
essen ially
ceased
and
p ices
wi hin
CNMI
ha e
isen
o
mo e
han
US$50
pe
ba
(Wo hing on
and
Taisacan
1996).
I
is
gene ally
p esumed
ha ,
as
a
esponse
o
ma ke
o ces,
illegal
hun ing
has
inc eased
in
he
no he n
Ma ianas,
and
ha
some
in e na-
ional
smuggling
also
occu s
(Wiles
1994;
Wo hing on
and
Taisacan
1996).
No
legal
ba ie
exis s
o
comme cial
im-
po s
o
Appendix
II
o
unlis ed
species
om
a
numbe
o
coun ies.
The e
a e,
howe e ,
p ac ical
ba ie s,
such
as
ma ke
eluc ance
o
accep
un amilia
species,
complica-
ions
ega ding
ade
a angemen s,
and
he
isk
o
ligh
delays
spoiling
highly pe ishable
ca go.
In
he
pas ,
local
p e e ence
o
la ge,
s ongly
scen ed
P e opus
wi h
ew
pa asi es
led
o
p e e en ial
ma ke
hun ing
and
p icing.
Fo
example,
buye s
isi ing
Samoa
p e e ed
he
less
common
P.
samoensis
o
P.
onganus.
Smalle
P e opus
om
Palau
and
he
Fede a ed
S a es o
Mic onesia
(FSM)
we e
ini ially
o
less
in e es .
Accep ance
o
smalle
ca e-dwelling
p e opo-
dids
wi h
ob ious
ails
(e.g.,
Rouse us)
was
also
poo
(G.
Phocas,
pe sonal
communica ion;
Sheeline
1991;
D.
Wo h-
ing on,
pe sonal
communica ion).
Reeme ging
p oposals
o
impo
Rouse us
om
main-
land
Asia
o
P e opus
om
Aus alia
(G.
Wiles,
D.
Wo h-
ing on,
pe sonal
communica ion)
inc ease
he
isk
ha
peo-
ple
who
p epa e
ozen
ba s
would
be
exposed
o
le hal
pa hogens,
such
as
lyssa i uses,
which
a e
p esen
in
a
leas
some
species
o
Aus alian
P e opus
(F ase
e
al.
1996;
Young
e
al.
1996).
Al hough
hese
isks
may
no
al e
he
beha io
o
indi idual
consume s,
public
heal h
au ho i ies
will
be
obliged
o
ein e p e
impo
egula ions
o
ec o s
o
communicable
diseases
o
allow
in e na ional
ade
o
expand
along
hese
lines.
The
only
ecu ing
in e na ional
ade
ou side
o
he
Ma ianas
o
ba s
lis ed
on
CITES
in ol es
a
ew
hund ed
P e opus
expo ed
annually
om
Vanua u
o
New
Caledonia
(J.
Caldwell,
pe sonal
commu-
nica ion).
In oduced
P eda o s
Al hough
in oduced
p eda o s
ha e
had
a
se ious
impac
on
island
bio as
(A kinson
1989),
hey
ha e
no
gene ally
been
implica ed
as
he
p ima y
agen s
in
he
long- e m
declines
o
ex inc ion
o
island
ba s.
A
no able
excep ion
is
he
a e
o
wo
species
endemic
o
New
Zealand,
Mys acina
obus a
and
Mys acina
ube cula a.
The
las
known
popula-
ion
o
M.
obus a
wen
ex inc
du ing
an
i up ion
o
Ra us
a us
in
he
mid-1960s
(Daniel
1990).
Also,
a
key
ole
can
be
in e ed
o
a s
and
o he
in oduced
p eda o s
in
he
ex-
ensi e
ange
con ac ion
o
bo h
species
h oughou
New
Zealand
(Daniel
and
William
1984;
Daniel
1990).
The
en-
dency
o
Mys acina
species
o
oos
and
o age
close
o
he
g ound
may
accoun
o
hei
di e en ial
ulne abili y
(Daniel
and
William
1984).
E en
hough
a s
(especially
R.
a us)
and
e al
ca s
com-
monly
climb
ees
and
ock
su aces,
ba s
oos ing
in
ee
canopies,
in
bole
ca i ies,
and
on
he
ceilings
o
ca es
ha e
managed
o
coexis
wi h
hese
p eda o s
in
mos
se ings.
Thei
abili y,
lacking
in
bi ds,
o
mo e
h ea ened
non olan
young
has
likely
helped.
On
Ch is mas
Island
in
he
eas e n
Indian
Ocean,
Tidemann
e
al.
(1994)
ound
ha
he
canopy-
oos ing
endemic
P e opus
melano us
na alis
was
ulne able
o
e al
ca
p eda ion
when
indi iduals
o aged
nea
he
g ound
in
ui ing
sh ubs.
Limi ed
obse a ions
o
a
second
endemic,
Pipis ellus
mu ayi
(included
in
Pipis ellus
enuis
by
Koopman
1993),
showed
ha
i
o ms
small
g oups
in
ees
a he
han
ca es
(Tidemann
1985).
Despi e
hese
in-
oduced
p eda o s
and
conside able
hun ing
o
lying-
oxes
by
humans,
bo h
ba
species
we e
p esen
in
subs an-
ial
numbe s
in
he
1980s
(Tidemann
1985;
Tidemann
e
al.
1994).
Howe e ,
as
discussed
o
Samoa,
single
opical
cy-
clones
can
sha ply
inc ease
he
sho - e m
ulne abili y
o
canopy
ugi o es
and
nec a i o es
o
in oduced
e es ial
p eda o s.
The
bes -documen ed
example
o
an
in oduced
non-
human
p eda o
educing
island
ba
popula ions
is
Boiga
i egula is,
an
a bo eal
Aus alasian
snake
on
Guam.
P ob-
ably
a i ing
in
pos -Wo ld
Wa
II
mili a y
shipmen s
om
he
Admi al y
Islands,
his
snake
slowly
inc eased
o
high
densi y,
elimina ing
mos
o
he
island's
a i auna
and
appa -
en ly
p e en ing
local
ec ui men
in
P e opus
ma iannus
Conse a ion
o
Remo e
Indo-Paci ic
Island
Ba s
333
(Sa idge
1987;
Wiles
1987b;
Rodda
e
al.
1992;
Wiles
e
al.
1995).
Ex ensi e
in es iga ions,
including
con ol
and
con-
ainmen
me hods,
sugges ed
ha
his
snake
will
pe sis
on
Guam
inde ini ely
(McCoid
1991),
lea ing
long- e m
p os-
pec s
o
he
local
ba
popula ion
in
doub .
Guam
is
a
egional
cen e
o
mili a y
anspo ,
and
indi iduals
o
B.
i egula is
ha e
been
eco e ed
on
Saipan,
Kwajalein,
Oahu,
and
Diego
Ga cia
(McCoid
1991).
Because
i
is
likely
ha
dispe sal
will
con inue
and
ha
igilance
on
islands
ecei -
ing
shipmen s
may
no
always
be
su icien , esea ch
on
eme gency
e adica ion
me hods
is
impo an
(McCoid
1991).
In es iga ions
o
ep oduc ion
in
Aus alian
B.
i egu-
la is
indica ed
ha
spe m
s o age
may
enhance
i s
success
as
an
in ade
(Whi ie
and
Limpus
1996).
G eene's
(1989)
obse a ions
on
he
die
and
habi s
o
o he
Boiga
spp.
sugges ed
ha
hey
oo
pose
se ious
in asion
isks.
Di e -
ences
ha
pe mi
coexis ence
be ween
Boiga
and
P e opus
in
po ions
o
hei
na u al
ange
a e
unknown,
bu
i
is
likely
ha
snake
densi ies
a e
lowe .
A
noc u nal,
commensal
snake
om
sou heas
Asia,
Ly-
codon
aulicus,
which
was
i s
de ec ed
on
Ch is mas
Island
in
1987,
now
appea s
es ablished
(F i s
1993).
I s
maximum
epo ed
size
(84
cm
o al
leng h
[TL];
F i s
1993)
and
limi ed
gape
(H.
G eene,
pe sonal
communica ion)
make
p eda ion
on
adul
P.
m.
na alis
highly
imp obable.
I s
b oad
die a y
habi s
and
a bo eali y
sugges
ha
i
may,
howe e ,
be
a
majo
h ea
o
he
su i al
o
he
much
smalle
Pipi-
s ellus
mu ayi.
A
scena io,
simila
o
ha
obse ed
in
Guam,
o
high
snake
densi ies
causing
he
ex inc ion
o
indigenous
e eb a es
o
lowe
ecundi y,
including
ba s,
is
likely
unless
his
snake
can
be
quickly
elimina ed.
Ano he
possible
ins ance
o
an
in oduced
p eda o
ha -
ing
an
impac
on
an
island
ba
popula ion
is
he
decline
o
Coleu a
seychellensis.
Racey
and
Nicoll
(1984)
ou lined
a
his-
o y
o
epo s,
indica ing
ha
he
species
was
" e y
com-
mon"
in
1868
and
educed
o
16
indi iduals
in
hei
ecen
su eys.
Sugges ed
causes
o
decline
a e
loss
o
o es
habi-
a ,
human
dis u bance
o
oos s,
and
occupancy
o
oos
ca es
by
ba n
owls
(Ty o
alba).
Ba n
owls
we e
in oduced
o
he
Seychelles
in
1949
(Cheke
and
Dahl
1981),
and
ha e
been
shown
elsewhe e
o
be signi ican
p eda o s
on
insec-
i o ous
ba s
(Speakman
1991).
Mass
Mo ali y
om
Disease
Anecdo es
o
in oduced
diseases
decima ing
island
e e-
b a es
a e
no
uncommon
(Simbe lo
1995),
bu
only
e-
cen ly
has
he e
been
ecogni ion
ha
mass
mo ali y
om
pandemic
disease
is
a
a e
bu
widesp ead
phenomenon
among
Paci ic
island
lying- ox
popula ions
in
he
pos -
Eu opean-con ac
e a
(Flanne y
1989,
1995;
Pie son
and
Rainey
1992)
(Table
23.3).
Pe haps
because
such
e en s
migh
go
unde ec ed
(o
no
occu )
in
less-colonial
species,
epo s
all
conce ned
P e opus,
which
o m
la ge
agg ega-
ions
a
adi ional
oos s.
Desc ip ions
a e
d ama ic,
wi h
incapaci a ed
ba s
alling
om
he
sky
and
ca casses
o
bones
accumula ing
in
piles
a
oos
si es
(Coul as
1931;
Degene
1949;
Flanne y
1989).
Few
o
no
ba s
o
he
a -
ec ed
species
we e
seen
a e
epidemics,
and,
consis en
wi h
he
popula ion
biology
o
P e opus,
eco e y
is
e-
po ed
as
slow
(S ai
1887;
Flanne y
1989,
1995).
The
wo
oldes
epo s
link
hese
e en s
o
simul aneous
epidemics
in
humans
o
domes ic
animals
(S ai
1887;
Coul as
1931).
As
Flanne y
(1989)
poin ed
ou ,
high
mo ali y
a o s
he
hypo hesis
o
a
human-in oduced
pa hogen
o
which
ba s
had
no
p io
exposu e.
Again
by
analogy
o
Hawaii,
be-
cause
ew
ba s
ha
come
in o
con ac
wi h
humans
a e
la e
eleased,
ans e
o
pa hogens
o
o es -dwelling
ba s
en-
Table
23.3
Mass
Mo ali ies
o
P e opus
om
Disease
on
Indo-Paci ic
Islands
Locali y
Samoa
Kos ae
(FSM)
Vanua
Le u,
Fiji
New
Caledonia
Manus,
Papua
New
Guinea
Bougain ille
and
Buka,
Solomon
Island
Species
Da e
P.
onganus
and/o
P.
samoensis
P.
ma iannus
ualanus
P.
onganus
P.
o na us
P.
neohibe nicus
hilli
P.
ayne i
g andis
Commen s
1839
P io
epidemic
among
esiden s;
g adual
ba
popula ion
eco e y;
species
no
iden i ied
1926-27
Measles
and
dysen e y
epidemic
in
esiden s;
housands
o
ba s
died;
ew
su i o s
Be o e
1949
Hund eds
dead
unde
oos s
Ea ly
1960s
Fo me ly
common;
no
subsequen
popula ion
eco e y
unde
hun ing
p essu e
1985
Masses
dead
in
oos
a eas;
slow
eco e y;
sympa ic
P.
admi ali a um
una ec ed;
una ec ed
P.
neohibe nicus
on
adjacen
islands
1987
Una ec ed
popula ions
on
adjacen
islands
Sou ce
S ai
1887
Coul as
1931
Degene
1949
Flanne y
1995
Flanne y
1989
Flanne y
1989
336
W.
E.
RAINEY
a i ely
sugges s
an
a h opod
o
o he
animal
ec o .
The
lack
o
e idence
o
ecu en
mass
mo ali ies
om
disease
in
be e -s udied
empe a e
zone
ba
popula ions
sugges s
ha
hese
e en s
a e
no
pa
o
he
p ehuman
e olu iona y
his o y
o
island
ba s.
Howe e ,
bi ds
anspo
pa hogens
and
a h opod
ec o s
wi h
b oad
hos
anges
o
islands
(Olsen
e
al.
1993).
In e island
anspo a ion
by
humans
con inues
o
imp o e,
inc easing
he
p ospec
o
dispe sing
mic obes,
a h opod
ec o s,
and
al e na e
hos s.
Any
new
ba
epidemic
ha
is
de ec ed
dese es
ca e ul
s udy,
gi en
he
isks
o
s ochas ic
mo ali y
o
he
long- e m
pe sis-
ence
o
small
popula ions.
Global
Clima e
Change
Cu en
models
o
global
wa ming
sugges
ha
oceanically
bu e ed
clima es
in
opical
egions
a e
expec ed
o
show
small
inc eases
in
empe a u e
(Ligh hill
e
al.
1994),
he
consequences
o
which
a e
unknown
o
island
e eb a es.
Two
o he
p ojec ed
changes,
ising
sea
le els
and
al e ed
cyclone
egimes,
would
howe e
almos
ce ainly
ha e
ma-
jo
impac s
on
island
bio as.
In
he
absence
o
local
ec onic
upli ,
sea
le el
ise
h ea ens
cu en
e es ial
communi-
ies
on
low- elie
a olls,
which
accoun
o
many
o
he
wo ld's
emo e
oceanic
islands.
On
such
islands,
ela i ely
la ge
a eas
o
land
would
be
los
wi h
only
a
small
ise
in
sea
le el
(17-26
cm
by
2030
and
inc easing
he ea e ).
The
dec easing
lens
o
esh
g oundwa e
and
inc eased
e ec s
om
sal
sp ay
would
also
a ec
e eb a e
consume s
be-
cause
dec easing
plan
di e si y
will
educe
ood
esou ces
(Roy
and
Connel
1991).
Roy
and
Connel
(1991)
discussed
p ospec s
o
humans
who
inhabi
na ions
composed
la gely
o
a olls
(e.g.,
Ki iba i
o
Maldi es),
concluding
ha
he
cu en
pa e n
o
economic
emig a ion
will
likely
be
ans o med
du ing
he
nex
se e al
decades
in o
an
exodus
o
en i onmen al
e u-
gees.
Fo
coun ies
(e.g.,
FSM,
Cook
Islands)
ha
include
bo h
sca e ed
low
a olls
and
upli ed
ca bona e
o
eme -
gen
olcanic
islands
( ypically
wi h
g ea e
popula ion
and
in as uc u e),
his
mig a ion
may
be
pa ly
in e nal
bu
will
inc ease
human
densi y
and
demand
o
na u al
e-
sou ces
on
al eady
c owded
islands.
A
ew
ba
species
and
popula ions
endemic
o
low- elie
a olls
would
be
di ec ly
h ea ened
by
ising
sea
le els
(e.g.,
P e opus
hox ensis
on
On ong
Ja a;
P.
ma iannus
ul hiensis
on
Uli hi;
he
P.
insula is
popula ion
on
Namonui o
A oll,
FSM
(Rainey
and
Pie son
1992);
P.
phaeocephalus
in
he
Mo locks,
FSM;
P.
gigan eus
a iel
in
he
Maldi es).
Se e al
o he
species
(e.g.,
P.
onganus
in
Fiji
and
Tonga,
P.
insula is
on
Chuuk,
P.
ma iannus
subspp.
in
Yap
and
Palau)
occu
bo h
on
high-
and
low-lying
islands
and
hus
a e
less
acu ely
h ea ened.
How-
e e ,
a
a
scale
o
a
ew
kilome e s,
los
o
al e ed
ege a ion
will
elimina e
o aging
habi a
and
e uges
om
human
hun ing.
A
a
scale
o
many
kilome e s,
he
dis ibu ion
o
e en
small
numbe s
o
ba s
and
pa ches
o
o aging
habi a
o e
se e al
islands
o e s
escape
in
space
and
ime
om
he
ecu ing
bu
ela i ely
na ow
pa hs
o
opical
cyclones.
The
isk
o
ex inc ion
inc eases
as
he
o al
geog aphic
ange
o
a
species
declines
owa d
an
a ea
po en ially
swep
by
cyclones
in
a
high- equency
yea .
Mang o e
o es s
a e
ecognized
as
an
impo an
coas al
habi a
o
ba s,
especially
P e opus.
They
p o ide
ela i ely
p o ec ed
si es
o
agg ega ed
oos ing
(e.g.,
P.
ampy us
on
Timo
[Goodwin
1979],
P.
molossinus
on
Pohnpei,
and
P.
ma iannus
ualanus
on
Kos ae
[Rainey
1990]),
wi h
some
common
ees,
no ably
Sonne a ia,
seasonally
o e ing
an
impo an
nec a
esou ce
(S a
and
Ma shall
1976).
Inpu s
o
sedimen
om
majo
i e s
may
allow
mang o e
o es s
on
con inen al
coas lines
o
keep
pace
wi h
p ojec ed
sea
le el
ise
(Jelge sma
e
al.
1993).
On
small
islands
whe e
e es ial
inpu s
o
coas al
sedimen s
a e
much
lowe , sea
le el
ise,
in
he
absence
o
ec onic
upli ,
will
exceed
soil
acc e ion
in
mang o e
o es s,
making
o es
dea h
a
likely
scena io
(Ellison
and
S odda
1991).
An
al e ed
opical
cyclone
egime
could
be
a
majo
con-
sequence
o
global
wa ming.
These
cyclic
dis u bances
play
a
key
ole
in
shaping
o es
s uc u e
and
species
composi-
ion,
and,
o
low-lying
islands,
hey
can
al e
coas al
geo-
mo phology
(Whi mo e
1974;
Shaw
1983;
Tanne
e
al.
1991;
S odda
and
Walsh
1992;
Elmq is
e
al.
1994;
Fos e
and
Boose
1995).
Ma ked
educ ions
in
ba
popula ions,
ol-
lowed
by
de e ed
ep oduc ion,
accompany
se e e
cy-
clones
(Cheke
and
Dahl
1981;
Wiles
1987a;
Pie son
and
Rainey
1992;
S inson
e
al.
1992;
C aig
e
al.
1994;
Gannon
and
Willig
1994;
G an
e
al.
1994;
Pie son
e
al.
1996).
In-
c eased
cyclone
equency
o
in ensi y
would
bode
ill
o
he
long- e m
pe sis ence
o
low- ecundi y
e eb a e
popu-
la ions
on
small
islands.
Assuming
highe
su ace
empe a u es
o
opical
seas,
as
p edic ed
by
global
clima e
models,
se e al
s udies
ha e
p ojec ed
inc eases
in
he
maximum
in ensi y,
mean
in en-
si y,
equency,
and
la i udinal
ange
o
opical
cyclones
and
modeled
hei
biological
o
economic
consequences
(Emanuel
1987;
Gable
and
Aub ey
1990;
O'B ien
e
al.
1992).
This
is
a
con o e sial
opic.
A
ecen
e iew
sug-
ges ed
ha
he
la i udinal
ange
o
cyclones
will
no
in-
c ease
and
ha
cu en
clima e
models
canno
e alua e
wha
will
happen
o
equency
o
a e age
in ensi y
o
cy-
clones
(Ligh hill
e
al.
1994).
Some
in es iga o s,
howe e ,
ha e
a gued
s ongly
ha
maximum
possible
in ensi y
will
inc ease
(Emanuel
1995).
Reanalysis
o
ela i ely
de ailed
cyclone
da a
o
he
No h
A lan ic,
Ca ibbean,
and
Gul
o
Conse a ion
o
Remo e
Indo-Paci ic
Island
Ba s
337
Mexico
shows
a
signi ican
decline
in
maximum
in ensi y
du ing
he
pas
50
yea s
(Landsea
e
al.
1996).
Conclusions
The
zooa chaeological
eco d
o
emo e
Indo-Paci ic
is-
lands,
which
is
d awn
la gely
om
limi ed
samples
in
he
less-di e se
aunas
o
Polynesia,
shows
ha
ba s
o
he
ge-
nus
P e opus
gene ally
su i ed
ex ensi e
p ehis o ical
de-
auna ion.
Thei
su i al
ela i e
o
g ound-nes ing
seabi ds
o
nume ous
ex inc
ligh less
endemic
land
bi ds
is
no
su p ising.
Howe e ,
su i al
o
ba s
ela i e
o
smalle ,
canopy-dwelling,
nec a i o ous
o
ugi o ous
bi ds
likely
e lec s
widesp ead
cul u al
p e e ences
o
hun ing
bi ds,
combined
wi h
he
high
ulne abili y
o
a ian
eggs
and
young
o
p eda o s.
Ex inc ions
and
ex i pa ions
o
island
ba s,
om
he
p ehis o ic
o
he
p esen ,
include
many
ha
a e
oo
poo ly
documen ed
o
in e p e .
Se e al
losses
and
declines, howe e ,
unde line
he
ulne abili y
o
ba s
o
o e hun ing
(e en
wi h
limi ed
echnologies),
especially
hose
species
ha
oos
colonially
in
ca es
o
ee
ca i ies.
Endemic
axa
ha
ely
on
lowland
o
lowe
mon ane
p ima y
o es
ha e
unde gone
subs an ial
popula ion
e-
duc ions
wi h
o es
clea ance.
Ye ,
in
a eas
whe e
ba
hun -
ing
is
no
in ense,
se e al
species
ha e
pe sis ed,
albei
a
low
numbe s,
on
islands
wi h
high
human
densi ies.
Se e al
such
species,
now
con ined
o
dwindling
o es
agmen s,
a e
being
aided
by
conse a ion
in e en ion.
Especially
on
a olls,
adi ional
ag icul u al
p ac ices
on
islands,
which
emphasize
ee
c ops,
may
inc ease
he
abundance
and
p e-
dic abili y
o
ui
esou ces
o
ba s
and
hus
enhance
he
ca ying
capaci y
o
hose
P e opus
ha
ha e
ca holic
die s.
F ugi o ous
ba s
ha
oos
in
seconda y
o es
can
o en
coexis
success ully
wi h
man
so
long
as
hei
eal
o
pe -
cei ed
impac
on
ui
c ops
does
no
igge
ex e mina ion
e o s.
Recen
s a us
assessmen s
sugges
ha
p e opodid
popula ions
a e
highly
ulne able
o
o e hun ing,
and
eady
a ailabili y
o
guns
is
a
signi ican
isk
ac o .
Cul u al
pe spec i es
on
ba s
can
play
a
key
ole
in
de e mining
p ospec s
o
su i al.
Du ing
he
1970s
and
1980s,
comme cial
ha es
o
ba s
o
supply
he
adi ional
luxu y
ood
ma ke s
in
Guam
and
he
No he n
Ma ianas
had
a
pe asi e
in luence
on
he
P e opus
popula ions
on
a
numbe
o
islands
in
Polynesia
and
Mic onesia.
A
pa ial
excep ion
o
his
pa e n
was
ha
o
ba
popula ions
in
Palau,
which
declined
du ing
one
pe iod
o
comme cial
hun ing
bu
eco e ed
and
appea ed
o
emain
ela i ely
common
du ing
a
second, pe haps
less-
in ense
pe iod
(Wiles
e
al.
1997).
While
demand
o
ba s
is
s ill
high
in
he
ela i ely
a luen
Ma ianas,
li le
legal
in e -
na ional
ade
now
akes
place.
Limi ed
e idence
sugges s
ha
demand
is
pa ially
me
wi h
ba s
aken
illegally
wi hin
CNMI
unde
a
egime
o
minimal
en o cemen .
Also,
o -
es
deg ada ion
by
expanding
popula ions
o
in oduced
he bi o es
h ea ens
ba
habi a
on
some
islands.
Accumula ing
da a
on
molecula
a ia ion
in
ba s
om
Aus alia
and
he
Paci ic
Islands
sugges
ha ,
o
la ge
habi a
gene alis s,
p ac ical
managemen
uni s
based
on
cu en
wa e
gaps
among
islands
will
app oxima e
uni s
ha
would
be
delinea ed
by
gene ic
su eys.
Howe e ,
some
mon ane
o es
endemics
ha e
shown
mo e
wi hin-
popula ion
gene ic
a ia ion.
Al hough
simila
ba s
in
Aus alasia
and
elsewhe e
coex-
is
wi h
a
wide
ange
o
p eda o s,
he
in oduced
a bo eal
snake
Boiga
i egula is
appa en ly
p e en s
local
ec ui men
in
he
P e opus
popula ion
on
Guam. Guam
is
a
anspo a-
ion
hub,
and
anspo
o
snakes
o
o he
islands
has
al-
eady occu ed.
In oduc ions
o
p eda o s
pose
a
se e e
h ea
o
he
nea - e m
su i al
o
endemic
island
e e-
b a es,
including
ba s,
e en
whe e
habi a
is
s ill
ela i ely
in ac .
A
second
h ea
o
unce ain
impo ance,
bu
which
can
also
dispe se
h ough
in ac
habi a ,
is
disease.
Ou -
b eaks
ha e
been
documen ed
o
cause
se e e
educ ions
o
P e opus
popula ion
a
se e al
locali ies.
While
p ojec ed
an h opogenically
induced
clima e
change
is
apid
ela i e
o
he
Pleis ocene
eco d
o
na u al
clima e
change,
i s
e ec s,
excep
o
inunda ion
o
low
a olls,
on
emo e
island
ba
popula ions
will
likely
be
unde-
ec able
agains
he
g ea e
magni ude
and
pace
o
change
induced
by
human
popula ion
g ow h,
esou ce
consump-
ion,
and
habi a
al e a ion.
Acknowledgmen s
I
hank
A.
B ooke,
E.
Pie son,
D.
S eadman,
E.
Towle,
G.
Wiles,
and
D.
Wo hing on
o
discussions
o
island
biology,
li e a u e
sugges ions,
o
unpublished
epo s.
I
also
hank
K.
Koopman,
Ame ican
Museum
o
Na u al
His o y,
and
J.
E.
Hill
and
P.
D.
Jennings,
B i ish
Museum
(Na u al
His o y),
o
discussions
and
access
o
collec ions;
and
John
Caldwell,
Wo ld
Conse a ion
Moni o ing
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o
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A.
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