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Zoology 

NEW  SERIES,  NO.  106 


The  Mammals  and  Birds  of  Camiguin  Island, 
Philippines,  a  Distinctive  Center  of  Biodiversity 

Lawrence  R.  Heaney,  Editor 


April  5,  2006 
Publication  1537 


PUBLISHED  BY  FIELD  MUSEUM  OF  NATURAL  HISTORY 


UNIVERSITY  I 
LIBR/ 

AT  URBANA-CHA 
BIOL: 


-S4*^|| 


Frontispiece 

Two  species  of  mammals  and  one  species  of  bird  live  only  on  Camiguin  Island:  Apomys  sp.  (described 
herein;  center  left),  Bullimus  gamay  (described  by  Rickart  et  al.  in  2002;  lower  right),  and  Loriculus  sp. 
(described  herein;  upper  center).  The  distinctive  volcanic  peaks  of  Camiguin  form  the  background. 


8'OLOGY  UBRAfly 

OCT  2  7  2006 


FIELDIANA 


Zoology 

NEW  SERIES,  NO.  106 


The  Mammals  and  Birds  of  Camiguin  Island, 
Philippines,  a  Distinctive  Center  of  Biodiversity 

Lawrence  R.  Heaney,  Editor 

Division  of  Mammals,  Department  of  Zoology 
Field  Museum  of  Natural  History 
1400  South  Lake  Shore  Drive 
Chicago,  Illinois  60605-2496 


Accepted  16  April  2004 
Published  April  5,  2006 
Publication  1537 


PUBLISHED  BY  FIELD  MUSEUM  OF  NATURAL  HISTORY 


©  2006  Field  Museum  of  Natural  History 

ISSN  0015-0754 

PRINTED  IN  THE  UNITED  STATES  OF  AMERICA 


Contents 

1.  Mammal  and  Land  Bird  Studies  on  Camiguin  Island,  Philippines:  Background  and 

Conservation  Priorities 1 

Laurence  R.  Heaney  and  Bias  R.  Tabaranza,  Jr. 

2.  A  New  Species  of  Forest  Mouse,  Genus  Apomys  (Mammalia:  Rodentia:  Muridae),  from 

Camiguin  Island,  Philippines 14 

Lawrence  R.  Heaney  and  Bias  R.  Tabaranza,  Jr. 

3.  Synopsis  and  Biogeography  of  the  Mammals  of  Camiguin  Island,  Philippines 28 

Lawrence  R.  Heaney,  Bias  R.  Tabaranza,  Jr.,  Natalie  Rigertas,  and  Danilo  S.  Balete 

4.  A  New  Species  of  Hanging-Parrot  (Aves:  Psittacidae:  Loriculus)  from  Camiguin  Island, 

Philippines 49 

Jose  G.  Tello,  Jacob  F.  Degner,  John  M.  Bates,  and  David  E.  Willard 

5.  An  Annotated  Checklist  of  the  Land  Birds  of  Camiguin  Island,  Philippines 58 

Danilo  S.  Balete,  Bias  R.  Tabaranza,  Jr.,  and  Lawrence  R.  Heaney 

List  of  Figures  (abbreviated  titles) 

Frontispiece.  Color  drawing  of  three  endemic  species  from  Camiguin  Island  (one  bird, 

two  rodents) i 

Chapter  1. 

1.  Photo  of  Camiguin  Island 2 

2.  Map  of  Camiguin  Island,  showing  extent  of  forest  and  park  boundaries 3 

3.  Map  of  Camiguin  island,  showing  elevational  isoclines  and  site  locations 4 

Chapter  2. 

1 .  Phylogeny  of  Apomys 17 

2.  Photo  of  adult  Apomys  camiguinensis 18 

3.  Ventral  surface  of  hind  feet  of  Apomys 20 

4.  Photos  of  crania  of  four  species 21 

5.  SEMs  of  anterior  portions  of  skulls 22 

6.  SEMs  of  posterior  portions  of  skulls 23 

7.  SEMs  of  maxillary  toothrows 24 

8.  SEMs  of  mandibular  toothrows 24 

Chapter  3. 

1.  Elevational  ranges  of  non-volant  small  mammals 31 

2.  Elevational  ranges  of  fruit  bats 34 

3.  Species  accumulation  curve 44 

Chapter  4. 

1 .  Map  of  the  central  and  southern  Philippines 50 

2.  Results  of  the  Principal  Components  Analysis  of  morphological  data 55 

Chapter  5. 
No  figures 


List  of  Tables  (abbreviated  titles) 

Chapter  1. 
None 

Chapter  2. 

1.  Cranial  and  external  measurements  of  Apomys 16 

Chapter  3. 

1.  Numbers  of  nonvolant  small  mammals,  by  site 32 

2.  External  and  cranial  measurements  of  shrews 33 

3.  Numbers  of  fruit  bats,  by  site 34 

4.  External  and  cranial  measurements  of  fruit  bats 35 

5.  External  and  cranial  measurements  of  micro  bats 38 

6.  External  and  cranial  measurements  of  murid  rodents 41 

Chapter  4. 

1.  Results  of  analysis  of  variance,  by  sex 53 

2.  Results  of  analysis  of  variance,  by  taxon,  for  males 54 

3.  Results  of  the  principal  components  analysis 54 

Chapter  5. 
None 


Preface 


The  Philippine  Islands  have  been  recognized 
for  over  a  century  as  having  a  fauna  that  is 
characterized  by  large  numbers  of  endemic  and 
highly  distinctive  species.  For  most  of  this 
period,  the  mammals  and  birds  were  thought  to 
be  well  known,  based  on  studies  conducted  in  the 
period  1880-1950.  However,  the  description  of 
new  species  during  the  1980s  implied  that 
diversity  was  higher  than  had  been  believed 
and  suggested  that  there  might  be  additional 
localized  centers  of  endemism  yet  to  be  found.  In 
addition,  it  was  during  the  1980s  that  the  extent 
of  rain-forest  habitat  destruction  became  appar- 
ent and  increased  the  need  for  extensive  and 
detailed  documentation  of  biological  diversity  to 
guide  development  and  conservation  for  the 
benefit  of  both  wildlife  and  the  human  popula- 
tion of  the  Philippines. 

With  this  in  mind,  the  authors  of  this  volume 
and  their  colleagues  began  investigations  of 
places  likely  to  be  of  special  interest  and  initiated 
a  series  of  publications  in  Fieldiana:  Zoology  to 
document  the  findings.  The  field  studies  have 
been  conducted  in  a  standardized  fashion  to 
allow  direct  and  meaningful  comparison  between 
study  areas  so  that  geographic  patterns  of  species 
richness  between  islands  and  mountain  ranges 
and  along  elevational  gradients  could  be  docu- 
mented. However,  older  museum  collections, 
which  often  were  unreported  in  the  published 
literature,  have  also  been  utilized,  increasing  the 
rate  of  our  progress  in  understanding  the  re- 
markable mammal  and  bird  faunas  of  the 
Philippines.  The  first  publications  in  Fieldiana 
concerned  the  mammals  of  Leyte  and  nearby 
islands  north  of  Mindanao,  the  birds  and 
mammals  of  Mt.  Isarog  National  Park  in 
southern  Luzon,  and  the  birds  of  Sibuyan  Island. 
Plans  are  in  place  for  similar  publications  on  the 
mammals  and  birds  of  the  Kitanglad  Range  of 
northern  Mindanao  and  the  mammals  of  Sibu- 
yan Island,  and  others  are  likely  to  follow. 


This  volume  contains  the  results  of  our  studies 
of  Camiguin  Island,  a  small  island  that  lies  just 
north  of  central  Mindanao.  It  is  easily  over- 
looked on  a  map  of  the  Philippines,  yet,  as 
shown  here,  the  island  supports  an  endemic 
species  of  small  parrot  and  two  endemic  species 
of  small  mammals,  all  discovered  in  the  course  of 
our  studies.  Announcements  in  the  Philippine 
press  of  the  discovery  of  the  mammals  in  1994 
and  1995  played  an  important  role  in  encourag- 
ing the  declaration  of  the  remaining  rain  forest 
on  Camiguin  as  a  national  park,  a  movement 
that  is  continuing  to  gain  support.  It  is  our  hope 
that  this  publication  will  help  guide  the  planning 
for  this  protected  area  and  its  management  when 
it  has  been  formally  initiated.  The  data  contained 
here  constitute  a  baseline  from  which  changes 
may  be  measured  in  the  future  and  also  make 
clear  how  much  additional  study  is  needed. 
Although  Camiguin  is  a  small  island,  it  repre- 
sents an  ideal  natural  laboratory  in  which  to 
investigate  many  aspects  of  the  evolution  and 
conservation  of  biological  diversity. 

The  assistance  of  many  individuals  is  acknowl- 
edged in  each  of  the  chapters  that  follow,  but  we 
must  give  special  recognition  here  to  the 
Philippine  Department  of  Environment  and 
Natural  Resources  for  providing  both  permits 
and  encouragement;  we  especially  thank  Dr. 
Angel  C.  Alcala,  Dr.  Corazon  Catibog-Sinha, 
Mr.  Carlo  C.  Custodio,  Atty.  Wilfredo  Pollisco, 
and  Dr.  Mundita  Lim  for  their  steadfast  support. 
This  project  has  benefited  greatly  from  the 
financial  support  provided  by  the  John  D.  and 
Catherine  T.  MacArthur  Foundation,  from  the 
Ellen  Thorne  Smith  and  Marshall  Field  Funds  of 
the  Field  Museum,  and  especially  the  Barbara 
Brown  Fund  for  Mammal  Research  of  the  Field 
Museum. 

L.  R.  Heaney 

November  2005 
Chicago,  Illinois 


FIELDIANA:  ZOOLOGY,  N.S.,  NO.  106,  APRIL  5,  2006,  P.  vii 


Mammal  and  Land  Bird  Studies  on  Camiguin  Island, 
Philippines:  Background  and  Conservation  Priorities 

Lawrence  R.  Heaney    and  Bias  R.  Tabaranza,  Jr. 


Abstract 

Camiguin  Island,  with  an  area  of  ca.  265  km'  and  maximum  elevation  of  ca.  1620  m,  lies 
about  10  km  north  of  Mindanao  but  is  isolated  from  Mindanao  by  a  deep  (385  m)  channel.  It 
originated  from  volcanic  activity  as  a  dryland  island  not  earlier  than  1  million  years  ago,  but 
most  growth  of  the  island  has  occurred  within  the  past  340,000  years.  Current  landforms  are 
dominated  by  large,  scenic  volcanic  peaks,  several  of  which  are  active.  Lowland  rain  forest 
originally  occurred  up  to  about  1100  m  elevation,  with  montane  rain  forest  from  1100  m  to 
about  1350  m  and  mossy  forest  from  1350  m  to  the  peaks.  By  the  mid-1990s,  deforestation 
had  removed  most  natural  vegetation  below  about  600  m,  with  degree  of  disturbance  to  forest 
decreasing  with  elevation  and  ending  at  about  1250  m.  The  climate  is  tropical,  with  rainfall  of 
2-3  m  per  year  in  the  lowlands  and  probably  about  7.5  m  near  the  peaks.  Mammal  and/or 
bird  specimens  are  available  from  18  sites  from  the  1960s  and  1990s;  these  sites  are  here 
located  and  described  to  the  extent  possible.  Given  the  presence  of  two  endemic  species  of 
mammals  (one  described  in  this  volume),  one  endemic  bird  (described  in  this  volume),  and 
previously  described  endemic  plants  and  a  frog,  Camiguin  is  one  of  the  smallest  but  most 
distinctive  centers  of  biodiversity  in  the  Philippines  and  should  be  a  priority  site  for 
conservation.  The  remaining  forest  on  Camiguin  is  essential  habitat  for  these  unique  species, 
but  it  is  also  essential  for  watershed  protection  and  control  of  floods  and  landslides,  and  it 
contributes  significantly  to  the  tourism  trade  that  provides  substantial  income  on  the  island. 
Deforestation  for  logging  and  agriculture  and  overhunting  are  current  threats.  A  protected 
area  on  the  island  should  include  the  full  range  of  original  habitat  diversity,  which  would 
encompass  both  the  existing  high-quality  forest  at  upper  elevations  and  also  significant  tracts 
of  disturbed  but  natural  lowland  forest,  especially  along  rivers  and  streams,  that  should  be 
allowed  to  regenerate  in  the  future. 


Introduction 

The  mammal  and  bird  faunas  of  the  Philippine 
Islands  are  remarkable  for  the  high  total  di- 
versity   of  the    fauna    and    especially    for    the 


1  Field  Museum  of  Natural  History,  1400  South 
Lake  Shore  Drive,  Chicago,  IL  60605-2496,  U.S.A. 

2  Department  of  Biology,  Iligan  Institute  of  Tech- 
nology, Mindanao  State  University,  Iligan  City, 
Lanao  del  Norte,  Philippines. 


remarkably  large  number  of  endemic  species 
(e.g.,  Dickinson  et  al.,  1991;  Mittermeier  et  al., 
1997,  1999;  Heaney  &  Regalado,  1998;  Heaney  et 
al.,  1998;  Stattersfield  et  al.,  1998).  For  example, 
at  least  512  of  the  898  species  of  breeding 
terrestrial  vertebrates  (57%)  are  endemic  to  the 
Philippines,  an  unusually  high  value  (Heaney  & 
Regalado,  1998).  Most  of  the  species  endemic  to 
the  country  occur  on  the  large  islands  of  Luzon, 
Mindanao,  Mindoro,  Negros,  and  Palawan  (e.g., 
Hauge  et  al.,  1986;  Heaney,  1986;  Heaney  et  al., 
1998;  Collar  et  al.,  1999;  Peterson  et  al.,  2000), 


FIELDIANA:  ZOOLOGY,  N.S.,  NO.  106,  APRIL  5,  2006,  PP.  1-13 


Fig.  1.     Photograph  of  Camiguin  Island  from  south-southeast,  taken  on  27  March  1995.  All  the  volcanic  peaks 
visible  on  the  island  are  less  than  350.000  years  old:  Mt.  Ginsiliban,  in  the  left  foreground,  is  the  oldest  (see  text). 


but  significant  numbers  of  endemic  mammals  are 
restricted  to  some  of  the  smaller  islands  as  well 
(e.g..  Heaney.  1986:  Goodman  &  Ingle,  1993: 
Heaney  &  Tabaranza.  1997;  Musser  et  al.,  1998; 
Heaney  &  Mallari,  2002:  Rickart  et  al..  2002). 
Evidence  has  suggested  that  those  small  islands 
with  endemic  species  are  nearly  always  islands 
surrounded  by  deep  water  (more  than  125  m).  so 
that  they  were  not  connected  to  adjacent  larger 
islands  during  periods  of  low  sea  level  during  the 
"ice  ages"  of  the  middle  and  late  Pleistocene, 
when  sea  level  dropped  to  no  more  than  120  m 
below  present  level  (Heaney.  1986.  1991a.b: 
Heaney  &  Regalado.  1998:  Hanebuth  et  al., 
2000;  Siddall  e^l.,  2003).  Clearly,  if  we  are  to 
understand  the  overall  patterns  of  the  evolution 
and  ecology  of  diversity  in  the  Philippines,  we 
must  understand  the  biodiversity  patterns  of 
these  smaller  centers  of  endemism  as  well  as  the 
larger  ones. 

One  of  the  smaller  Philippine  islands  sur- 
rounded by  deep  water  and  still  retaining 
moderate  rain-forest  cover  is  Camiguin,  located 
about  10  km  off  the  north-central  shore  of 
Mindanao  Island,  with  minimum  intervening 
water  depth  of  385  m  (the  island  of  Camiguin 
Norte,  which  lies  north  of  Luzon,  is  often 
confused  with  Camiguin).  It  is  a  steeply  moun- 
tainous island  (Fig.  1)  of  about  265  km2  with 
several  active  volcanic  cones  that  reach  to 
a  maximum  elevation  of  about  1620  m.  A  series 
of  biological  surveys  on  Camiguin  in  the  late 
1960s  (described  below)  that  focused  on  birds 
also  yielded  some  mammal  specimens.  An  earlier 


report  on  the  mammals  (Heaney,  1984)  conclud- 
ed that  the  island  had  no  endemic  mammal 
species  and  that  it  was  depauperate  relative  to  its 
area.  Subsequent  studies  on  other  islands  made 
us  suspect  that  those  earlier  mammal  surveys 
were  incomplete  because  so  few  mammal  species 
had  been  obtained  and  because  the  number  of 
nonvolant  mammal  voucher  specimens  was  small 
(thus  indicating  limited  sampling  effort).  To 
investigate  the  hypothesis  that  the  previously 
measured  species  richness  of  mammals  was  low 
because  of  incomplete  surveys,  we  returned  to 
Camiguin  briefly  in  1992  and  more  extensively  in 
1994  and  1995  to  conduct  additional  mammal 
inventories  in  all  the  major  habitats  along  the 
elevational  gradient,  especially  by  trapping  small 
mammals  at  higher  elevations  (Heaney  &  Taba- 
ranza. 1997),  and  we  obtained  some  new  data  on 
the  birds  as  well.  In  this  volume,  we  summarize 
results  from  both  the  1960s  and  the  1990s 
surveys  and  present  evidence  that  both  endemic 
mammal  and  bird  species  are  present  on  the 
island  (Heaney  et  al.,  2006;  Tello  et  al.,  2006). 

There  is  a  second  reason  that  we  conducted 
surveys  on  Camiguin  in  the  mid-1990s.  While  the 
Philippines  is  increasingly  being  recognized  as 
a  global  center  for  biodiversity,  with  unusually 
high  levels  of  endemism.  it  has  simultaneously 
vaulted  into  public  view  as  one  of  the  most 
severely  deforested  of  the  tropical  countries  and 
home  to  what  may  be  the  greatest  concentration 
of  endangered  species  of  mammals  and  birds 
(Collar  et  al.,  1994,  1999;  Heaney  &  Regalado, 
1998:  Stattersfield  et  al.,  1998;  Mittermeier  et  al.. 


FIELDIANA:  ZOOLOGY 


■  9°13 


-9°11 


124°  38' 


9°  15' 


124°  40' 


124°  42' 


124°  44' 


124°  46' 


124°  48' 


White 
Island 


9°09 


9  07' 


9°05' 


Mantigue 

Island 

o 


5  kilometers 


Fig.  2.  Map  of  Camiguin  Island  showing  the  locations  of  mountain  peaks  referred  to  in  the  text,  boundaries  of 
municipalities  in  the  mid-1990s,  the  approximate  boundaries  of  the  proposed  Timpoong-Hibok-hibok  Natural 
Monument  (dotted  line),  and  extent  of  forest  cover  in  1987  (gray  area  bounded  by  dashed  line)  from  National 
Mapping  and  Resource  Information  Authority  (1988).  The  names  of  the  municipalities  are  adjacent  to  the  primary 
population  centers  (=  poblacion)  of  each  municipality,  shown  as  solid  squares. 


1999;  Environmental  Science  for  Social  Change, 
2000;  Ong  et  al.,  2002).  Satellite  maps  of  forest 
cover  from  1986  (National  Mapping  and  Re- 
source Information  Authority,  1988)  showed 
a  substantial  area  of  forest  cover  in  the  center  of 
Camiguin  (Fig.  2);  such  forest  cover  is  now  a  rarity 
in  the  Visayas  (the  islands  of  the  central  Philip- 
pines) and  adjacent  regions  (Heaney  &  Regalado, 
1998;  Environmental  Science  for  Social  Change, 


2000),  a  portion  of  the  country  where  the 
concentration  of  endangered  mammals  and  birds 
is  especially  high  (Wildlife  Conservation  Society 
of  the  Philippines,  1997;  Collar  et  al.,  1999; 
Heaney  &  Mallari,  2002).  Because  deforestation 
has  generally  proceeded  rapidly  all  over  the 
country  in  recent  decades,  we  felt  a  sense  of 
urgency  to  obtain  current  information.  Simulta- 
neously,   the    prospect    of    finding    species    of 


HEANEY  AND  TABARANZA:  MAMMAL  AND  LAND  BIRD  STUDIES 


-  9M5' 


ST13' 


124°42' 


Site  9 
ir 

Site  1 1 

Site  10 


-  gro9' 


-gro7' 


I I I I I I 

5  kilometers 


Fig.  3.  Map  of  Camiguin  Island  showing  200-m  elevational  contours  and  locations  of  collecting  sites  in  the 
1990s  (solid  dots)  and  1960s  (dashed  circles);  as  noted  in  the  text,  the  locations  of  the  latter  are  approximate.  The 
municipal  population  centers  are  shown  also  (solid  squares). 


mammals  or  birds  in  a  place  where  enough  forest 
remained  to  support  stable  populations  of  any 
endemic  or  endangered  species  was  quite  exciting, 
posing  the  potential  for  positive  action  and  the 
development  of  successful  conservation  pro- 
grams. We  therefore  proceeded  to  Camiguin  with 
a  sense  of  cautious  optimism  and  found  condi- 
tions that  confirmed  all  our  best  hopes  but  also 
some  of  our  fears,  as  we  shall  describe  here. 


Geology,  Vegetation,  and  Climate 

Camiguin  is  composed  almost  entirely  of 
Quaternary  volcanic  material  from  currently 
active  volcanoes,  and  eruptions  have  occurred 


within  historic  time  (Hamilton.  1979;  Mitchell  et 
al.,  1986;  Hall,  1998,  2002),  with  the  most  recent 
serious  eruptions  in  1871  and  1951  (Agoo,  1995). 
Five  major  and  several  secondary  volcanic  peaks 
(Figs.  1-3)  dominate  the  island's  landscape  and 
serve  as  a  tourist  attraction;  hikers  often  reach 
the  crater  lake  in  Mt.  Hibok-hibok  and  visit  the 
steam  vents  along  its  sides. 

Camiguin  is  the  northernmost,  somewhat 
isolated  portion  of  the  Central  Mindanao  Arc 
of  volcanic  activity,  which  includes  areas  from 
Mt.  Apo  in  the  south  to  Mts.  Kalatungan  and 
Kitanglad  in  the  center  and  Camiguin  in  the 
north  (Sajona  et  al.,  1997).  Volcanic  activity  in 
this  arc  originated  about  2.5  million  years  ago 
(Ma),  though  most  has  taken  place  since  about 


FIELDIANA:  ZOOLOGY 


1.25  Ma.  Camiguin  is  probably  derived  from 
a  single  source  of  magma  that  first  produced 
undersea  lavas  in  the  late  Pliocene  or  early 
Pleistocene  (ca.  2  Ma),  with  dryland  appearing 
not  earlier  than  1  Ma.  Potassium-argon  dating 
shows  the  oldest  dated  magmatic  materials  on 
the  island  to  be  those  of  the  Mt.  Ginsiliban  and 
Mt.  Butay  volcanoes  in  the  southeastern  corner 
of  the  island  (Fig.  1),  which  date  to  0.34  Ma.  The 
peaks  and  volcanic  flows  associated  with  Mt. 
Mambajao  that  form  much  of  the  center  of  the 
island  are  much  younger,  and  Mt.  Hibok-hibok 
in  the  northwestern  portion  of  the  island  is  the 
youngest  and  only  currently  active  vent  (Pu- 
nongbayan  &  Solidum,  1985;  Sajona  et  al.,  1997; 
Castillo  et  al.,  1999).  We  interpret  this  to  indicate 
that  Camiguin  originated  as  a  dryland  island  not 
earlier  than  1  Ma  and  had  high  peaks  and 
substantial  area  by  at  least  300,000  years  ago, 
though  possibly  somewhat  earlier  (Heaney  1986, 
1991a,  b).  The  island  is  surrounded  by  deep 
water,  with  a  minimum  depth  to  Mindanao  of 
385  m,  which  far  exceeds  the  lowest  documented 
late  or  middle  Pleistocene  sea  lowering  of  ca. 
120  m  (Siddall  et  al.,  2003);  thus,  there  is  no 
evidence  of  a  dryland  connection  to  Mindanao  at 
any  time  in  the  past. 

Camiguin  Island  lies  within  a  climate  zone 
characterized  by  annual  rainfall  of  2-3  m  or 
more  in  the  lowlands,  with  seasonal  variation 
that  includes  a  moderately  dry  period  from 
March  to  May  (2.0-5.5  inches/mo;  ca.  5-14  cm/ 
mo),  especially  wet  from  October  to  January 
(10-15  inches/mo;  ca.  25-38  cm/mo),  and  moist 
during  the  rest  of  the  year;  mean  annual  rainfall 
for  a  24-year  period  at  Mambajao  was 
99.4  inches  (ca.  252.5  cm;  Manalo,  1956).  Tem- 
perature declines  and  rainfall  increases  with  an 
increase  in  elevation;  based  on  patterns  elsewhere 
in  the  Philippines  (Heaney  &  Regalado,  1998), 
we  estimate  that  rainfall  at  1500  m  is  roughly 
triple  that  at  sea  level  (i.e.,  about  7.5  m/year),  but 
no  details  from  Camiguin  are  available. 

The  original  vegetation  of  Camiguin  was 
lowland  tropical  rain  forest  from  near  the  sea 
to  about  800  m  elevation,  with  montane  and 
mossy  forest  to  the  peaks,  but  by  the  mid-1990s, 
virtually  no  original  vegetation  remained  below 
300  m  and  little  from  300  to  about  800  m  (Agoo, 
1995).  Beach  and  mangrove  forest  once  occurred 
along  the  coast,  but  little  remains.  Agoo  (1995, 
p.  4)  described  the  upper  elevations  of  Mt. 
Hibok-hibok  as  a  submontane  plant  community 
"dominated  by  shrubby  plants  of  Radermachera, 


Rhododendron,  Medinilla,  and  Vaccinium.  Gym- 
nosperms  are  conspicuously  absent.  .  .  .  The 
vegetation  at  the  peak  is  devoid  of  tall  trees 
and  shrubs.  It  is  composed  mostly  of  turfs  of 
grasses."  Descriptions  of  vegetation  at  our 
sampling  areas  during  the  1990s  are  below. 

Agoo  (1995)  noted  the  presence  of  the 
following  plants  endemic  to  Camiguin:  Miquelia 
reticulata  (Icacinaceae),  Medinilla  multiflora, 
Memecylon  subcaudatum  (Melastomataceae),  Sy- 
zigium  camiguense  (Myrtaceae),  Coelogyne  con- 
fusa,  and  Goodyera  ramosii  (Orchidaceae). 

A  small  frog  (Oreophryne  nana)  was  the  only 
endemic  vertebrate  known  from  Camiguin  when 
we  began  our  work  (Alcala  &  Brown,  1998). 


Methods 

Prior  Reports 

Aside  from  a  very  small  number  of  scattered 
records  (e.g.,  Gray  1843,  who  reported  Paradox- 
urus  hermaphroditus)  and  a  few  specimens  of  birds 
from  the  1930s  deposited  at  the  Museum  of 
Comparative  Zoology,  the  first  significant  surveys 
of  birds  and  mammals  on  Camiguin  were 
conducted  by  field  teams  from  Silliman  Univer- 
sity at  10  sites  in  1967,  1968,  and  1969  and  on  the 
small  offshore  island  of  Mantigue  in  1969.  The 
field  team  in  1967  was  led  by  D.  P.  Empesso  and 
R.  B.  Gonzales;  we  know  of  only  bats  collected  by 
this  team,  all  of  which  were  deposited  in  the  Royal 
Ontario  Museum.  Labels  on  specimens  indicate 
that  collecting  was  conducted  at  three  sites,  listed 
below  as  Sites  1 6—1 8.  No  field  catalogs  or  notes 
were  kept,  and  no  information  is  available  on 
their  field  procedures,  but  we  suspect  that  they 
followed  roughly  the  same  procedure  as  was  used 
in  subsequent  years  by  Dioscoro  S.  Rabor,  as 
follows.  Field  teams  in  1968  and  1969,  which 
produced  all  the  bird  specimens  and  many  of  the 
mammals  cited  here,  were  headed  by  Rabor;  his 
teams  worked  at  seven  sites,  according  to  the 
labels  on  specimens  in  the  Field  Museum  of 
Natural  History  (fmnh)  and  Delaware  Museum 
of  Natural  History  (dmnh).  B.R.T.  Jr.  worked  on 
several  of  Rabor's  teams  during  the  1970s  and 
observed  that  Rabor  followed  a  standardized 
approach  to  fieldwork.  The  field  crew  simulta- 
neously occupied  a  base  camp,  where  Rabor 
remained  most  of  the  time,  and  several  satellite 
camps.  The  satellite  teams,  made  up  of  two  or 


HEANEY  AND  TABARANZA:  MAMMAL  AND  LAND  BIRD  STUDIES 


three  persons,  would  go  to  areas  chosen  by  Rabor 
and  operate  for  several  days  to  several  weeks. 
Members  in  each  satellite  camp  would  set  nets 
each  day.  sometimes  would  run  a  small  number  of 
large  snap  traps,  and  would  hunt  using  shotguns 
over  an  area  that  could  readily  be  covered  on  foot. 
Once  each  day.  one  member  would  hike  to  the 
base  camp  with  the  specimens  obtained  since  the 
last  such  trip,  then  return  to  the  satellite  camp.  A 
similar  team  of  collectors  would  operate  in  the 
area  of  the  base  camp  as  well.  At  the  time  of 
collection,  specimens  were  labeled  simply,  usually 
with  the  name  of  the  site  (often  the  name  of 
a  barangay  or  sitio.  which  could  cover  many 
square  kilometers)  and  rough  estimate  of  elevation 
based  on  topographic  maps.  Rabor  would  work 
with  a  team  to  prepare  specimen  labels  (including 
measurements,  date,  and  locality  data)  and  to  skin 
and  stuff  specimens.  No  field  catalog  or  notes  were 
usually  kept:  data  were  recorded  only  on  the 
specimen  skin  tags  (see  also  Rabor.  1966). 

We  have  reconstructed  the  activities  at  Ra- 
bor's  field  sites  by  listing  all  the  locality  names 
that  were  noted  and  compiling  the  dates  on 
which  specimens  were  obtained.  Because  field 
teams  operated  simultaneously,  the  dates  over- 
lap. We  noted  the  elevation,  when  present,  for 
each  locality  name,  providing  us  with  a  range  of 
elevations  for  that  site:  many  labels  lack  eleva- 
tion, but  we  assume  that  the  elevations  noted 
apply  to  all  specimens  from  a  given  site.  We 
believe,  however,  that  many  of  the  elevations  as 
indicated  on  the  labels  of  the  specimens  collected 
during  this  period  were  overestimated  and  that 
they  should  be  used  with  caution.  Additional 
comments  on  elevational  records  can  be  found  in 
specific  site  descriptions. 

Recent  Data 

Field  studies  that  focused  on  mammals  but 
also  obtained  some  data  on  birds  were  conducted 
during  three  periods  in  1992.  1994.  and  1995  by 
representatives  from  the  fmnh  and  Mindanao 
State  University-Iligan  Institute  of  Technology. 
The  first  reconnaissance  trip  extended  from  26  to 
30  May  1992:  bats  were  netted  at  a  single 
agricultural  site  by  a  field  team  led  by  A.  T. 
Peterson.  The  second  period  extended  from  4 
May  1994  to  4  June  1994:  specimens  were 
collected  at  one  agricultural  site  and  three  forest 
sites  by  a  team  led  by  B.R.T.  Jr.  The  third  period 
extended  from  12  to  26  March  1995:  specimens 


were  collected  at  two  forest  sites  by  a  team  led  by 
L.R.H.  and  B.R.T.  Jr.  Our  methods  for  sampling 
mammals  and  birds  are  presented  in  the  relevant 
chapters. 

Study  Sites 

Camiguin  is  dominated  by  a  series  of  volcanic 
peaks,  and  most  localities  refer  to  those  peaks. 
We  note  that  the  three  adjacent  high  peaks  near 
the  center  of  the  island  (Fig.  3)  are  often 
collectively  referred  to  as  Mt.  Mambajao.  though 
we  follow  local  people  in  referring  to  the 
northeasternmost  of  the  three  major  peaks  as 
Mt.  Timpoong  and  the  southwesternmost  as  Mt. 
Mambajao. 

Sites  from  1990s 

We  surveyed  mammals  at  one  site  in  1992. 
four  in  1994.  and  two  in  1995:  these  are  referred 
to  in  the  species  accounts  by  number,  and  their 
locations  are  shown  in  Figure  3.  Additional 
minor  sites  are  described  in  the  species  accounts. 
The  dates  include  only  those  days  when  speci- 
mens were  obtained.  Distances  are  given 
from  the  municipal  town  centers,  as  shown  in 
Figure  2. 

Site  1 — Barangay  Balbagon.  1  km  S.  2  km  E 
of  Mambajao.  10  m.  9°14.5'N,  12444'E  (28-29 
May  1992).  This  site  was  situated  in  a  highly 
disturbed  lowland  agricultural  area.  The  area 
originally  was  lowland  rain  forest  but  in  1992 
was  a  mosaic  of  agricultural  fields,  pastures,  and 
small  bits  of  secondary  lowland  forest.  Several 
patches  of  shrubbery  with  heights  of  2-4  m  were 
present,  as  were  several  large,  thick  patches  of 
erect  bamboo.  Six  nets  were  set  for  a  single  night. 
Bats  and  some  birds  were  collected. 

Site  2 — Barangay  Balbagon.  7  km  S.  2  km  E 
of  Mambajao.  1000  m.  9r10.5'N.  124  44'E  (28- 
29  May  1992).  A  few  specimens  of  birds  were 
netted  in  extensive,  regenerating  secondary  forest 
at  this  site. 

Site  3 — Barangay  Manuyog.  Sagay  Munici- 
pality. 7  km  S.  3  km  W  of  Mahinog.'  0-300  m. 
9°5.5'N,  124r45.5'E  (5-11  May  1994).  Sampling 
was  conducted  in  heavily  disturbed  lowland 
agricultural  land,  with  very  few  scattered  patches 
of  degraded  lowland  forest  along  steep  slopes 
beside  rivers.  Trapping  and  netting  were  con- 
ducted from  close  to  the  shoreline  (ca.  50  m  from 
the  beach)  up  to  300  m  along  the  southwestern 
slopes  of  Ginsiliban  Peak,  extending  partially 


FIELDIANA:  ZOOLOGY 


into  Ginsiliban  Municipality,  for  a  total  of  three 
net-nights  and  148  trap-nights. 

Site  4 — Barangay  Kital-is,  Sagay  Municipali- 
ty, Vi  km  N,  6Va  km  W  of  Mahinog,  1000  m, 
9°9.5'N,  124°43.5'E  (14-24  in  May  1994).  This 
site  was  in  disturbed  transitional  lowland/lower 
montane  forest  along  the  southwestern  slopes  of 
Mt.  Mambajao  at  900-1100  m  elevation.  The 
slope  was  moderate,  and  the  ground  was  covered 
with  abundant  leaf  litter.  The  vegetation  con- 
sisted mostly  of  small  trees  with  dbh  of  15-25  cm 
and  a  few  large  trees  standing  20-30  m  high  and 
with  40-60  cm  dbh;  some  lianas  (5-10-cm  di- 
ameter), rattan  {Calamus  spp.),  and  climbing 
bamboo  {Schizostachyum  spp.)  clung  to  these 
emergents.  Ficus  spp.  were  common.  The  most 
common  epiphytes  were  orchids,  moss,  and 
ferns.  Tree  ferns  (Cyathea  spp.)  and  rattan 
(Calamus  spp.)  seedlings  were  abundant  as 
ground  cover.  Scattered  areas  of  humus  were 
up  to  30  cm  thick.  Our  total  sampling  effort  at 
this  site  consisted  of  24  net-nights  and  907  trap- 
nights. 

Site  5 — XA  km  N,  6  !/2  km  W  of  Mahinog, 
Sagay  Municipality,  1200  m,  9°9.5'N,  124°43.5'E 
(26-29  May  1994).  This  site  was  situated  in 
disturbed  lower  montane  forest  on  moderate  slope 
at  1000-1300  m  elevation,  in  the  vicinity  of  Lasak- 
lasak  (Site  12).  There  was  some  evidence  of  small- 
scale  illegal  logging  done  about  a  year  previously. 
Emergent  trees  had  dbh  of  between  40  and  60  cm 
and  heights  up  to  30  m.  Lianas  and  canopy 
epiphytes  (mainly  ferns  and  mosses)  were  present. 
Rattan  {Calamus  spp.)  and  climbing  bamboo 
{Schizostachyum  spp.)  were  also  present,  with 
pitcher  plants  {Nepenthes)  present  but  rare.  Un- 
derstory  and  ground  cover  consisted  of  rattan 
seedlings  and  ferns,  with  some  sedges.  Leaf  litter 
covered  about  80%  of  the  ground,  and  humus  was 
typically  about  30  cm  thick.  Our  total  sampling 
effort  consisted  of  339  trap-nights;  at  this  site,  nets 
were  concurrently  maintained  with  those  at  Site  6 
for  a  total  of  14  net-nights. 

Site  6 — Barangay  Kital-is,  Sagay  Municipali- 
ty, on  a  small  peak  near  Mt.  Mambajao,  1  km  N, 
7'/2km  W  of  Mahinog,  1300m,  9°10'N, 
124°43'E  (26-29  May  1994).  This  site  was 
situated  in  primary  mossy  forest  at  1200- 
1400  m.  Hanging  moss  was  abundant  on  trees, 
but  moss  cover  on  the  ground  was  light  and  the 
humus  layer  not  more  than  a  few  centimeters 
thick.  Our  total  sampling  effort  consisted  of  348 
trap-nights  for  this  site  and  14  net-nights  for 
Sites  5  and  6  combined. 


Site  7 — Mt.  Timpoong,  2  km  N,  6  Vi  km  W  of 
Mahinog,  1275  m,  9°10.5'N,  124°43.5'E  (17-25 
March  1995).  This  site  was  situated  in  primary 
montane  forest  (Fig.  3)  at  1225-1350  m  eleva- 
tion, in  the  vicinity  of  Lasak-lasak.  The  average 
slope  was  ca.  35°  and  was  often  steep.  The  forest 
had  a  fairly  low  and  relatively  open  canopy;  the 
height  of  emergent  trees  was  usually  20-25  m, 
but  a  few  reached  30  m;  dbh  was  12-30  cm,  and 
none  had  buttresses.  The  canopy  was  broken  by 
many  treefalls,  and  canopy  leaf  sizes  were  small, 
3-6  cm,  often  with  serrated  edges.  Lower  strata 
leaf  sizes  were  4-20  cm  and  usually  6-10  cm. 
Epiphytes  were  abundant,  including  ferns,  or- 
chids, and  mosses.  Arborescent  pandans  (Pan- 
clanus  sp.),  melastome  shrubs  (Melastoma  spp.), 
and  tree  ferns  (Cyathea  spp.)  were  common,  and 
climbing  rattan  (Calamus  spp.)  and  viney  pan- 
dans  (Freycinetia  spp.)  were  abundant;  Ficus  and 
Musa  were  rare  to  absent.  This  area  had  thin  to 
moderate  leaf  litter  cover  and  thin  to  moderate 
humus  (up  to  15  cm  deep);  the  soil  consisted  of 
lightly  weathered  volcanic  rock  with  many  stones 
at  the  surface  and  was  generally  very  shallow. 
There  were  no  signs  of  human  disturbance,  but 
many  trees  had  fallen  because  of  very  shallow 
soil,  probably  during  occasional  typhoons.  There 
were  eight  net-nights  and  655  trap-nights. 

Site  8— Mt.  Timpoong,  2%  km  N,  6 '/a  km  W 
of  Mahinog,  1475  m,  9°11'N,  124°43.5'E  (22-25 
March  1995).  Sampling  at  this  site  was  con- 
ducted in  mossy  forest.  The  site  was  on  a  steep 
slope,  averaging  50°,  and  ranged  from  30-70°, 
including  steep  gullies.  The  height  of  the  canopy 
was  usually  8-10  m  but  varied  from  2  to  3  m  in 
exposed  spots  to  a  maximum  of  ca.  18  m  in  low, 
protected  places;  trees  were  generally  gnarled, 
with  dbh  of  12-20  cm  (rarely  25  cm).  Canopy 
leaves  were  small,  with  serrated  edges,  typically 
1-8  cm,  but  most  were  4-5  cm.  Oaks,  laurels, 
tree  ferns,  and  arborescent  Pandanus  were 
common.  Lower-strata  leaves  were  small,  usually 
4-8  cm.  Ficus  and  Musa  were  rare  to  absent. 
Epiphytes  were  abundant,  including  mosses, 
ferns,  orchids,  and  saplings,  and  pitcher  plants 
(Nepenthes  spp.)  were  common.  Canopy  vines 
(Freycinetia  spp.  and  Calamus  spp.)  were  abun- 
dant. Ground  plants  included  ferns,  saplings, 
and  abundant  pandans  (Freycinetia).  The  ground 
was  covered  by  abundant  and  thick  leaf  litter 
over  deep  humus  (over  a  half  meter  thick,  with 
many  tunnels  and  vacuities).  Moss  covered 
nearly  all  tree  trunks,  branches,  and  fallen  logs 
and  hung  from  branches  in  sheets  and  often 


HEANEY  AND  TABARANZA:  MAMMAL  AND  LAND  BIRD  STUDIES 


covered  the  ground.  No  human  disturbance  was 
seen,  but  there  was  evidence  of  some  large 
landslides  and  many  tree  falls.  No  netting  was 
performed  at  this  site;  there  were  386  trap- 
nights. 

Sites  from  the  1960s 

As  noted  above  (see  Methods),  field  teams 
from  Silliman  University  conducted  surveys  on 
Camiguin  at  10  sites  in  1967-1969  plus  an 
additional  site  on  the  small,  adjacent  Mantigue 
Island.  As  noted  above  (Methods),  we  have 
estimated  the  location  of  these  sites;  our  esti- 
mates are  shown  in  brackets.  The  vegetation 
types  and  condition  at  these  sites  were  not  noted 
by  the  collectors.  We  surmise,  on  the  basis  of 
specimens  collected  here  and  the  condition  of  the 
remaining  vegetation  on  Camiguin  in  the  1990s, 
that  when  the  collecting  was  done,  the  vegetation 
below  800  m  ranged  from  secondary  lowland 
forest  to  heavily  populated  agricultural  areas.  By 
the  time  of  our  surveys  in  the  1990s,  the  lowland 
forest  was  almost  totally  cleared  and  replaced 
with  coconuts,  and  hardly  any  area  below  800  m 
supported  remnant  forest  except  along  steep 
slopes.  During  the  1960s,  from  ca.  1000  m  up  to 
the  peak,  the  sites  would  have  been  covered  with 
primary  montane  and  mossy  forest,  as  they  were 
in  the  1990s. 

Site  9 — Mt.  Catarman,  Catarman  Municipal- 
ity, 2000-4950  ft  (ca.  600-1500  m)  [approx. 
5.5  km  S,  4.5  km  W  Mambajao,  9  12'N, 
124  40.5' E]  (10  29  June  1968).  This  was  the 
highest  site  surveyed  in  1968;  both  birds  and 
mammals  were  collected.  Mammals  were  collect- 
ed at  2500-4500  ft  (ca.  750-1400  m).  The  ma- 
jority of  the  birds  specimens  were  taken  on  12-21 
June,  but  a  few  others  were  collected  on  10,  25, 
and  29  June.  Bird  specimens  were  taken  from 
2000  to  4950  ft  (ca.  600-1500  m).  This  would 
suggest  that  the  team  sent  to  this  camp  concen- 
trated their  efforts  from  the  middle  to  the  upper 
slopes,  including  the  peak  of  Mt.  Catarman,  but 
several  were  taken  at  1000  ft  (ca.  300  m)  on  14 
and  16  June  1968.  However,  we  also  know  that 
collecting  at  this  site  was  done  simultaneously 
with  Site  1 1 ,  and  specimens  from  both  sites  were 
then  taken  down  to  the  base  camp  at  Site  10  for 
processing  by  D.  S.  Rabor.  We  suspect  that  in 
the  process,  there  was  some  mixing  up  of 
elevational  data  for  some  specimens  (see  also 
Site  1 1 ).  Furthermore,  the  highest  peak  of  the 
Mt.  Catarman  does  not  exceed  1400  m,  so  all  the 


specimens  cited  by  the  team  to  have  been 
collected  on  this  mountain  above  this  elevation 
were  probably  obtained  close  to  the  peak. 

Site  10 — Gidag-on,  Catarman  Municipality, 
500-1500  ft  (ca.  1 50-450  m)  [approx.  9T0.5'N, 
124°39.5'E]  (13-28  June  1968).  This  was  the 
lowest  campsite  during  the  1968  field  season; 
birds  and  mammals  were  collected.  Birds  were 
collected  mostly  from  23  to  28  June  1968,  with 
a  few  taken  on  earlier  dates  (13,  16-17,  and  19 
June).  Some  birds  collected  on  13  June  were 
noted  to  have  been  collected  at  2000  ft  (ca. 
600  m),  possibly  the  highest  point  reached  from 
this  site.  Because  it  was  the  lowest  campsite  in 
1968,  it  is  the  most  likely  location  of  the  base 
camp  that  year.  We  are  unable  to  locate  this  site, 
but  some  collecting  dates  at  this  site  overlapped 
with  Sites  9  and  1 1;  we  suspect  that  it  was  on  the 
lower  slopes  of  Mt.  Catarman.  No  information 
on  the  vegetation  when  the  specimens  were 
collected  at  this  site  is  available,  but  we  surmise 
that  it  would  have  been  partly  secondary  lowland 
forest  and  partly  agricultural. 

Site  11 — Kasangsangan,  Catarman  Munici- 
pality, 1000-2500  ft  (300-800  m)  [approx. 
9  ll'N,  12440' E]  (11-22  June  1968).  This  was 
the  middle  campsite  in  1968.  We  are  unable  to 
locate  this  site,  but  because  collecting  dates 
overlapped  with  Site  9,  we  suspect  that  it  was 
mainly  on  the  lower  to  middle  slopes  of  Mt. 
Catarman.  Several  specimens  were  collected  up 
to  4950  ft  (ca.  1500  m),  which  might  indicate 
that  the  team  assigned  to  this  site  ventured  all  the 
way  to  the  peak  of  Mt.  Catarman,  but  we  suspect 
mixing  of  localities,  as  noted  above.  No  in- 
formation on  the  vegetation  when  the  specimens 
were  collected  is  available,  but  we  surmise  that, 
similar  to  Site  9,  it  would  have  been  partly 
secondary  lowland  forest  and  partly  agricultural. 

Site  12 — Mt.  Timpoong,  Lasak-lasak,  Mahi- 
nog  Municipality,  4400-5700  ft  (ca.  1350- 
1700m)  [approx.  9  1  l'N,  12443. 5'E]  (19-28 
June  1969).  We  traveled  through  this  site  in 
1995;  it  is  located  near  the  headwaters  of  the 
Sagay  River,  from  1200  m  to  roughly  the  peak. 
This  was  the  highest  site  surveyed  in  1969;  birds 
and  mammals  were  collected.  Collecting  activi- 
ties were  conducted  from  19  to  28  June.  The 
majority  of  the  bird  specimens  were  taken  at 
4800-5700  ft  (ca.  1500-1700  m),  indicating  that 
the  team  concentrated  their  collecting  in  the 
high-elevation  habitats  on  Mt.  Timpoong.  Sev- 
eral specimens  were  taken  slightly  farther  down, 
at  4400  ft  (ca.  1350  m).  On  23  June  1969,  some 


FIELDIANA:  ZOOLOGY 


specimens  attributed  to  this  site  were  noted  as 
taken  at  800  m.  As  with  the  previous  year  at  Sites 
9-11,  collecting  at  this  site  was  done  simulta- 
neously with  Site  13,  and  specimens  were  taken 
down  to  the  base  camp  at  Site  14  for  processing. 
We  suspect  that  some  mixing  up  of  elevational 
data  for  some  specimens  from  among  the  three 
sites  occurred.  While  the  vegetation  at  this  site  was 
not  indicated  when  the  1960s  collecting  was 
undertaken,  it  would  have  been  primary  mossy 
forest,  based  on  the  condition  of  the  habitat  that 
we  saw  on  Mt.  Timpoong  in  the  1990s. 

Site  13 — Mt.  Timpoong,  Matugnao,  Mahinog 
Municipality,  3150  ft(ca.  950  m)  [approx.  9°10'N, 
124°44.5'E]  (12-26  June  1969).  This  was  the 
middle  campsite  established  in  1969;  birds  and 
mammals  were  collected.  The  majority  of  the  bird 
specimens  bear  the  elevation  3150  ft  (ca.  950  m), 
with  a  few  at  3250  ft  (ca.  1000  m),  around  the 
midslopes  of  Mt.  Timpoong.  A  few  specimens, 
however,  were  noted  as  taken  at  800  ft  (ca.  250  m) 
on  1 6  June  and  at  4800  ft  (ca.  1 500  m)  on  20  June. 
Collecting  at  this  site  was  done  simultaneously 
with  Sites  12  and  14,  and  we  suspect  that  some 
mixing  of  elevational  data  occurred. 

Site  14 — Puntod,  Mahinog  Municipality, 
800  ft  (ca.  250  m)  [approx.  2  km  N,  2  km  W 
Mahinog;  9°9.5'N,  124°46'E]  (24-29  June  1969). 
This  was  the  lowest  campsite  established  in  1969; 
birds  and  mammals  were  collected.  Bird  collect- 
ing took  place  on  24-29  June.  Elevations  were  all 
given  as  800  ft  (ca.  250  m),  with  a  single  bird 
specimen  bearing  an  elevation  of  700  ft  (ca. 
200  m).  Puntod  is  a  village  located  on  the  lower 
slopes  of  Mt.  Timpoong,  about  2  km  west  of  the 
town  center  of  Mahinog  (Fig.  3).  This  probably 
was  the  base  camp  during  the  1969  field  season. 
The  vegetation  in  the  1960s  is  unknown,  but 
given  its  situation  as  a  settled  village  during  that 
time,  the  habitat  was  probably  already  largely 
agricultural  with  remnant  lowland  forest. 

Site  15 — Mount  Timpoong  Peak,  Mahinog 
Municipality,  5700  ft  (ca.  1600  m)  [approx. 
9°11'N,  124°43.5'E].  Only  a  single  specimen  of 
Simcus  murinus  bears  this  locality.  Given  the 
restriction  to  Mahinog  Municipality,  the  site 
must  have  been  at  the  peak  on  the  eastern  side  of 
the  mountain,  near  to  or  including  Site  8.  The 
vegetation  at  this  site  would  have  been  mossy 
forest,  as  described  for  Site  8. 

Site  16 — Mt.  Mambajao,  Mahidlaw,  Catar- 
man  Municipality,  2500-3500  ft  (ca.  800- 
1000  m)  [approx.  9  9.5'N,  124°42.5'E]  (24  and 
26  May  1967).  Only  a  few  bats  were  collected  at 


this  site.  Mt.  Mambajao's  highest  peak  is  ca. 
1600  m;  this  location  in  Catarman  Municipality 
suggests  that  specimens  were  taken  from  the 
middle  slopes,  on  the  southern  to  southwestern 
flank  of  the  mountain. 

Site  17 — Mt.  Mambajao,  Sangsangan,  Catar- 
man Municipality,  1400-3300  ft  (ca.  400- 
1000  m)  [approx.  9°9'N,  124°42.5'E]  (14-31 
May,  15  June  1967).  Only  mammals  were 
collected  at  this  site.  This  location  suggests  that 
specimens  were  taken  on  the  lower  to  middle 
slopes  of  Mt.  Mambajao,  on  the  southern  to 
southwestern  flank  of  the  mountain. 

Site  18 — Tag-ibo  Cave,  Catarman  Municipal- 
ity, 400  ft  (ca.  100  m)  (31  May  1967).  A  few  bats 
were  collected  at  this  site,  which  we  have  been 
unable  to  locate.  Because  it  was  visited  on  the 
same  day  as  specimens  were  obtained  from  Site 
17,  they  are  probably  near  each  other. 

Site  19 — Mantigue  Island  [approx.  2  km  N, 
4  km  E  Mahinog;  9°10.5'N,  124°49.5'E]  (28  June 
1969).  This  site  is  a  4-ha  coralline  island,  situated 
ca.  3  km  east  of  Barangay  Hubangon,  Mahinog 
Municipality  (Figs.  2  and  3).  It  was  visited  on  28 
June  1969,  and  only  White-collared  Kingfisher 
{Halcyon  cloris),  Yellow-vented  Bulbul  {Pycno- 
notus  goiavier),  Pied  Triller  (Lalage  nigra),  and 
Olive-backed  Sunbird  {Nectar  una  jugular  is)  were 
collected.  The  vegetation  when  it  was  visited  in 
1969  was  most  likely  beach  forest,  with  some 
disturbance. 

Conservation 

As  documented  by  Balete  et  al.  (2006),  Heaney 
et  al.  (2006),  and  Tello  et  al.  (2006),  Camiguin 
supports  at  least  24  species  of  mammals  and  at 
least  54  species  of  birds,  and  additional  fieldwork 
is  likely  to  document  the  presence  of  additional 
species.  This  includes  two  species  of  mammals  (in 
the  genera  Apomys  and  Bullimus)  and  one  species 
of  bird  (genus  Loriculus)  that  are  unique  to 
Camiguin;  Camiguin  is  the  smallest  island  in  the 
Philippines  currently  known  to  support  unique 
species  of  mammals  (Heaney,  1986;  Heaney  et 
al.,  1998)  and  is  smaller  than  any  island  pre- 
viously known  to  support  an  endemic  bird 
(Peterson  et  al.,  2000).  The  endemic  species  of 
mammals  have  been  documented  to  occur  in 
lowland,  montane,  and  mossy  forest  from 
1000  m  to  near  the  peaks  but  probably  occur  at 
lower  elevations  as  well  where  we  were  unable  to 
sample;  the  endemic  bird  (Tello  et  al.,  2005)  has 
been  documented  from  ca.  300  to  1200  m,  which 


HEANEY  AND  TABARANZA:  MAMMAL  AND  LAND  BIRD  STUDIES 


encompasses  lowland  and  lower  montane  forest. 
The  presence  of  these  unique  species,  along  with 
the  frog  and  plants  that  are  restricted  to  the 
island,  have  caused  Camiguin  to  be  listed  as  a  key 
national  and  global  priority  site  for  conservation 
(Mallari  et  al.,  2001:  Heaney  &  Mallari.  2002: 
Ong  et  al..  2002)  that  was  overlooked  by 
previous  assessments  that  were  based  on  in- 
complete biological  surveys  (e.g..  Hauge  et  al.. 
1986:  Heaney.  1993:  Peterson  et  al.,  2000). 

These  diverse  mammal  and  bird  faunas 
originally  occurred  along  the  entire  elevational 
and  habitat  gradient.  The  most  fundamental 
requirement  for  their  conservation  is  the  contin- 
ued existence  of  substantial  areas  of  all  original 
types  of  habitat  in  mature,  good  condition.  The 
areas  must  be  large  enough  for  the  existence  of 
substantial  populations,  not  just  a  few  individu- 
als, since  populations  often  are  vulnerable  to 
extinction  when  they  drop  below  about  1000 
individuals  (Primack.  1998). 

Fortunately.  Camiguin  retains  enough  forest 
cover  to  make  this  possible,  though  there  are 
a  number  of  challenges  to  surmount.  The  results 
of  the  Swedish  Satellite  Survey  of  forest  cover  in 
1987  (National  Mapping  and  Resource  Informa- 
tion Authority.  1988)  showed  a  fairly  large  area 
of  forest  on  Camiguin.  mostly  at  upper  eleva- 
tions on  Mt.  Mambajao  and  Mt.  Timpoong  and 
associated  highlands,  as  shown  in  Figure  2.  Our 
surveys  in  1994  and  1995  showed  that  much  of 
that  forest  still  remained,  though  the  edges  had 
been  further  degraded.  Downslope.  the  degree  of 
disturbance  progressively  increased  so  that  little 
if  any  lowland  forest  remained  in  a  primary 
condition:  indeed,  most  had  been  replaced  with 
coconut  plantations,  agricultural  areas,  and 
grassland.  In  2001.  it  was  estimated  that  S29c 
of  the  island,  ca.  20.847  ha.  was  covered  by 
croplands  and  other  inhabited  areas:  almost 
half  the  island's  area  was  covered  with  coco- 
nut plantations  (http://agrilO.norminet.org.ph/ 
NMProfile/profile_camiguin.htm).  leaving  \S9c 
as  secondary  and  primary  forest. 

According  to  local  residents,  much  of  the 
forest  on  Camiguin  was  removed  by  commercial 
logging  operations  in  the  1980s,  with  the  logged 
areas  subsequently  being  cleared  for  agriculture. 
During  the  1990s,  small-scale  commercial  log- 
ging, operating  under  salvage  permits,  was  the 
main  agent  responsible  for  the  continuing  de- 
nudation of  the  areas  below  1000  m.  These 
salvage  permits  technically  allowed  only  dead 
trees  to  be  cut.  but  local  residents  told  us  that 


trees  were  often  killed  deliberately  by  the  loggers 
so  that  they  could  then  be  cut  down.  Slash-and- 
burn  agriculture  {kaingin)  of  the  remnant  forest 
usually  followed  quickly  (Heaney  &  Tabaranza. 
1997).  By  the  time  we  visited  the  island  in  1995. 
areas  below  300  m  had  no  native  forest,  and  only 
a  little  second  growth  remained  up  to  ca.  800  m. 
Only  the  montane  and  mossy  forest  was  in 
relatively  good  condition,  but  small-scale  logging 
and  kaingin  was  creeping  farther  up  on  the 
mountains:  we  noted  a  new  kaingin  at  1200  m  on 
the  side  of  Mt.  Timpoong  in  1995  (Heaney  & 
Tabaranza.  1997:  Mallari  et  al..  2000). 

It  is  essential  to  note  that  conservation  of  the 
forest  will  have  great  benefits  to  the  people  of 
Camiguin  as  well  as  to  the  wildlife.  The  forest 
can  serve  as  a  permanent  source  of  wood  and 
other  nontimber  forest  products  to  the  local 
inhabitants  if  this  is  carefully  managed.  Perhaps 
most  crucially,  the  forests  of  Camiguin  provide 
the  people  with  abundant  water  for  domestic, 
agricultural,  and  industrial  uses.  As  an  addition- 
al essential  ecological  service,  the  forest  helps 
prevent  soil  erosion  and  landslides.  In  early 
November  2002.  about  200  human  lives  were  lost 
and  thousands  of  houses  and  other  properties 
damaged  by  landslides  during  a  typhoon,  partic- 
ularly in  Mahinog.  This  disaster  demonstrates 
the  great  economic  and  human  cost  of  forest 
degradation.  Additionally,  the  continued  popular- 
ity of  Camiguin  as  a  tourist  destination  will  rely 
strongly  on  its  ability  to  provide  beautiful  scenery, 
attractive  areas  for  hiking,  fresh  and  clean  water, 
and  health}  coral  reefs  (which  require  low  levels  of 
siltation  from  adjacent  rivers). 

The  importance  of  protecting  the  remaining 
forest  of  Camiguin  can  be  appreciated  when  one 
considers  its  fast-growing  population.  In  1980,  the 
population  of  Camiguin  was  about  57.000;  two 
decades  later,  in  2000.  the  population  had  in- 
creased to  more  than  73.000:  in  2020.  it  is  projected 
to  increase  to  90.000  (http://www.popcom.gov. 
ph/sppr/statistics/Ieastvis_wesmin_northmin.htm). 
Consequently,  the  population  density  of  Cami- 
guin increased  from  279  persons/km2  in  1980  to 
306  persons/km"  in  2000:  these  were  well  above 
the  national  density  of  160  persons/km2  and 
251  persons/km2  in  1980  and  2000.  respective- 
ly (http://www.popcom.gov.ph/sppr/statistics/ 
lleastvisz_wesmin_northmin.htm:  http://www. 
popcorn. gov.ph/sppr/statistics/table3. htm). 

In  response  to  all  these  issues,  including  our 
discoveries  in  1994-1995  of  endemic  species  of 
mammals,  the  local  gov  ernment  and  Department 


10 


FIELDIANA:  ZOOLOGY 


of  Environment  and  Natural  Resources  (DENR) 
have  moved  steadily  in  recent  years  to  have  the 
upland  areas  where  forest  remains  declared  the 
"Timpoong-Hibok-hibok  Natural  Monument." 
At  the  time  of  this  writing,  the  process  of 
officially  designating  the  protected  area  has 
reached  the  office  of  the  secretary  of  the  DENR 
and  is  expected  to  soon  be  endorsed  to  the 
president.  If  it  is  successful  at  that  level,  as  is 
anticipated,  it  must  then  be  voted  on  by  the 
Philippine  Congress  for  final  designation,  a  pro- 
cess that  may  yet  require  several  years.  The 
approximate  proposed  boundary  of  the  natural 
monument,  with  an  area  of  2,228  ha,  plus 
a  1,423-ha  buffer  zone,  is  shown  in  Figure  2. 
Although  both  the  location  and  area  of  forest 
and  the  natural  monument  as  shown  in  Figure  2 
are  approximate,  it  is  clear  that  the  majority  of 
the  primary  forest  lies  within  the  proposed 
boundaries  of  the  park. 

We  fully  endorse  these  recommendations  and 
actions.  But  further,  as  part  of  this  program  of 
environmental  protection  and  stabilization,  we 
recommend  the  following  to  the  local  govern- 
ment of  Camiguin  and  the  Philippine  DENR  as 
essential  activities:  1)  continue  and  expand  active 
enforcement  efforts  to  protect  existing  forests, 
wildlife,  and  environment;  2)  continue  and 
expand  current  reforestation  projects,  using  only 
native  species  of  trees,  not  exotics,  because  the 
exotic  trees  do  not  provide  habitat  for  the 
biodiversity  of  Camiguin  and  are  injurious  to 
the  soil;  3)  cancel  existing  salvage  cutting  permits 
for  dead  and  fallen  trees  because  these  have 
often  been  abused;  and  4)  complete  the  process 
of  declaring  the  remaining  forest  and  key 
watershed  areas  a  national  protected  area  as 
soon  as  possible.  To  the  maximum  extent 
possible,  all  portions  of  the  protected  area 
should  be  connected  by  corridors  of  mature  (or 
regenerating)  habitat,  especially  along  rivers  and 
streams. 

In  connection  with  the  above  recommenda- 
tions, it  is  essential  to  both  wildlife  conservation 
and  watershed  protection  that  national  and  local 
governments  include  substantial  lowland  areas 
(those  below  800  m)  in  the  proposed  national 
park,  regardless  of  the  present  condition  of  the 
habitats  and  vegetation,  especially  including  all 
areas  within  200  m  of  streams  and  rivers.  We 
therefore  recommend  that  the  natural  monument 
be  expanded  to  include  all  good-quality  second- 
ary forest  at  all  elevations  and  to  include  all 
major  watercourses  down  to  at  least  600  m  of 


elevation.  This  will  enable  future  management  to 
rehabilitate/restore  lowland  forest,  which  is  one 
of  the  most  critical  habitats  on  Camiguin. 


Acknowledgments 

For  assistance  with  the  often  challenging  field 
work  on  Camiguin,  we  thank  N.  Antoque,  E. 
Batara,  N.  Batocael,  N.  Bojo,  M.  Carmona,  R. 
Fernandez,  A.  DeOcampo,  M.  Jayoma,  L. 
Mostrales,  A.  T.  Peterson,  G.  Rosell,  A.  Tabar- 
anza,  B.  Tabaranza  III,  and  D.  Tabaranza. 
Permits  were  provided  by  the  Protected  Areas 
and  Wildlife  Bureau  of  the  Department  of 
Environment  and  Natural  Resources  (DENR); 
we  offer  special  thanks  to  A.  C.  Alcala,  C. 
Custodio,  M.  Mendoza,  and  W.  Pollisco  and  to 
the  Region  10  Office  of  the  DENR.  Emmauel 
Aranas  and  Primitivo  Espinas  provided  essential 
assistance  and  logistical  support  on  Camiguin 
during  the  fieldwork.  We  thank  E.  Canete,  C. 
Custodio,  and  G.  Rosell-Ambal  for  information 
on  the  status  of  the  protected  area.  For  the  loan  of 
specimens  under  their  care,  we  are  indebted  to  M. 
Carleton  and  H.  Kafka  (usnm)  and  P.  Myers 
(ummz);  we  especially  thank  G.  Hess  (dmnh)  for 
much  assistance  and  encouragement  over  many 
years.  The  maps  and  graphs  were  prepared  by 
Lisa  Kanellos;  the  photograph  of  Camiguin  was 
digitally  prepared  by  Rebecca  Banasiak.  We 
thank  P.  Janney  for  information  on  the  geological 
history  of  Camiguin  and  J.  Bates,  N.  Collar,  and 
E.  Rickart  for  constructive  comments  on  an 
earlier  draft  of  the  manuscript.  This  research 
was  supported  by  the  World  Environment  and 
Resources  Program  of  the  John  D.  and  Catherine 
T.  MacArthur  Foundation  and  by  the  Marshall 
Field  Fund,  the  Ellen  Thorne  Smith  Fund,  and 
the  Barbara  Brown  Fund  for  Mammal  Research 
of  the  Field  Museum  of  Natural  History. 


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HEANEY  AND  TABARANZA:  MAMMAL  AND  LAND  BIRD  STUDIES 


13 


A  New  Species  of  Forest  Mouse,  Genus  Apomys 
(Mammalia:  Rodentia:  Muridae),  from  Camiguin 
Island,  Philippines 

Lawrence  R.  Heaney1  and   Bias  R.  Tabaranza,  Jr.2 


Abstract 

An  inventory  of  the  mammals  of  Camiguin  Island  conducted  in  1994  and  1995  documented 
the  presence  of  a  previously  unknown  species  of  Philippine  forest  mouse  of  the  endemic 
Philippine  genus  Apomys,  which  is  here  named  and  described.  Based  on  molecular  data,  the 
new  species  is  most  closely  related  to  two  species  {A.  hylocoetes  and  A.  insignis)  from 
Mindanao  Island  and  to  an  unnamed  species  from  Leyte,  Biliran,  and  Bohol  islands.  The  new 
species  is  diagnosed  in  comparison  to  its  three  closest  relatives  on  the  basis  of  slightly  browner 
and  less  russet  fur,  slightly  greater  size  overall,  a  moderately  long  and  broad  hind  foot  with 
small  plantar  pads,  large  tail  scales,  slightly  narrower  zygomatic  width  and  mastoid  breadth, 
deep  rostrum  of  moderate  length,  a  long  orbit  and  braincase,  narrow  palate,  large  incisive 
foramina,  short  distance  from  the  posterior  edge  of  the  incisive  foramina  to  the  anterior  edge 
of  the  first  upper  molar,  bony  palate  that  extends  well  to  the  posterior  of  the  posterior  edge  of 
the  last  upper  molar,  bullae  that  are  more  strongly  oriented  toward  the  cranial  midline  axis, 
third  upper  molar  without  a  conspicuous  anterolabial  cusp,  and  a  number  of  more  subtle 
features.  It  is  one  of  two  species  of  mammals  now  known  to  be  endemic  to  Camiguin,  the 
other  being  Bullimus  gamay  (Rickart  et  al.,  2002).  Both  are  common  in  rain  forest  on 
Camiguin  Island  at  upper  elevations.  The  presence  of  two  endemic  mammals  on  this  small 
(265  km")  island  is  remarkable;  there  are  no  smaller  islands  in  the  Philippines  known  to 
support  endemic  mammal  species. 


Introduction 

The  Philippine  Islands  are  notable  for  their 
large  number  of  unique  species  of  mammals;  of 
172  species  known  in  1998,  111  (64%)  occurred 
nowhere  else  in  the  world,  one  of  the  greatest 


1  Field  Museum  of  Natural  History,  1400  South 
Lake  Shore  Drive,  Chicago,  IL  60605-2496,  U.S.A. 

2  Department  of  Biology,  Iligan  Institute  of  Tech- 
nology, Mindanao  State  University,  Iligan  City, 
Lanao  del  Norte,  Philippines. 


concentrations  of  unique  mammalian  diversity 
worldwide  (Heaney  et  al.,  1998).  These  species 
are  usually  not  widely  distributed  within  the 
Philippines  but  rather  are  confined  to  one  or 
a  few  islands.  Recent  studies  have  shown  that  the 
geological  history  of  the  archipelago  is  largely 
responsible  for  the  pattern  of  distribution,  with 
most  species  of  mammals  found  on  only  one  of 
the  several  islands  that  formed  during  periods  of 
low  sea  level  in  the  late  and  middle  Pleistocene. 
Each  of  these  Pleistocene  islands  is  surrounded 
by    deep    water    (greater    than    120  m    current 


14 


FIELDIANA:  ZOOLOGY,  N.S.,  NO.  106,  APRIL  5,  2006,  PP.  14-27 


depth),  and  each  has  remained  as  an  isolated 
oceanic  island  throughout  its  existence.  Howev- 
er, though  they  are  isolated  by  deep-water 
channels,  the  channels  are  not  wide,  usually  not 
more  than  25  km  and  often  much  narrower 
(Heaney,  1986,  1991,  1993,  2004;  Heaney  and 
Rickart,  1990;  Heaney  &  Regalado,  1998). 

Camiguin  Island  was  noted  by  Heaney  (1984, 
1986)  as  an  apparent  exception  to  this  pattern: 
collections  made  on  Camiguin  in  the  1960s  by 
field  teams  from  Silliman  University  did  not 
include  any  endemic  mammals,  even  though  the 
island  seemed  large  enough  to  support  them 
(Heaney,  1986,  2004).  After  the  discovery  of  four 
endemic  species  on  Sibuyan  Island,  another  small 
oceanic  island  in  the  archipelago  (Goodman  & 
Ingle,  1993;  Heaney  et  al.,  1998),  we  suspected 
that  the  mammals  of  Camiguin  might  not  have 
been  fully  surveyed,  so  we  returned  in  1994  and 
1995  for  further  investigations  (Heaney  et  al., 
2004;  Heaney  &  Tabaranza,  2006).  In  the  course 
of  those  field  studies,  we  documented  the  presence 
of  two  previously  unknown  species  of  mammals 
(Heaney  &  Tabaranza,  1997),  Bullimus  gamay 
(Rickart  et  al.,  2002)  and  a  species  of  forest 
mouse,  genus  Apomys.  It  is  the  purpose  of  this 
paper  to  describe  this  new  species  of  forest  mouse. 


Materials  and  Methods 

Specimens  examined  for  this  study  are  housed 
in  the  Delaware  Museum  of  Natural  History 
(dmnh),  Field  Museum  of  Natural  History 
(fmnh),  Mindanao  State  University-Iligan  In- 
stitute of  Technology  (msu-iit),  National  Muse- 
um of  the  Philippines  (nmp),  University  of 
Michigan  Museum  of  Zoology  (ummz),  and 
United  States  National  Museum  of  Natural 
History  (usnm).  Half  the  specimens  from  Cami- 
guin now  housed  in  fmnh  will  be  sent  to  nmp. 
Material  examined  included  specimens  prepared 
as  study  skins  with  skulls  (and  some  with 
postcranial  skeletons),  complete  skeletons,  and 
formalin-fixed  specimens  stored  in  70%  ethyl 
alcohol,  many  with  skulls  subsequently  removed 
and  cleaned.  The  following  samples  were  exam- 
ined: Apomys  hylocoetes — Mindanao  Island: 
Bukidnon  Province:  Mt.  Kitanglad  Range: 
15  km  S,  12.5  km  W  Dalwangan,  elev.  2800  m, 
fmnh  148055;  16.5  km  S,  4  km  E  Camp  Phillips, 
elev.  1900  m,  fmnh  147871-872,  148123-124; 
18.5  km  S,  4  km  E  camp  Phillips,  elev.  2250  m, 


fmnh  147874-876,  147880,  147900-904,  147906, 
148125-128,  148132,  148135-138.  Apomys  in- 
signis — Mindanao  Island:  Bukidnon  Province: 
Mt.  Kitanglad  Range:  16.5  km  S,  4  km  E  Camp 
Phillips,  elev.  1900  m,  fmnh  148152;  17  km  S, 
7  km  E  Baungon,  elev.  1550  m,  fmnh  146703; 
18  km  S,  7  km  E  Baungon,  elev.  1800  m,  fmnh 
146704-710,  146712-714,  146716-718,  147088- 
089,  147091-092,  147094,  147098-099,  147102. 
Apomys  sp. — Biliran  Island:  3  Vi  km  S,  5  Vi  km 
W  Caibiran,  elev.  700  m,  ummz  160314,  160290, 
160429^130;  4'/2  km  S,  5  km  W  Caibiran,  elev., 
920  m,  ummz  160290-291,  160315-316.  Leyte 
Island:  9  km  N,  3  km  E  Baybay,  elev.  750  m, 
ummz  160318;  8 !/2  km  N,  2'/2  km  E  Baybay, 
elev.  500  m,  ummz  160317,  160441.  Apomys  n. 
sp. — Holotype  and  referred  specimens  from 
Camiguin  Island  (see  below). 

Specimens  were  assigned  to  age  categories  as 
follows.  Subadult  animals  are  those  that  have 
not  completed  cranial  growth,  especially  those 
having  unfused  basicranial  sutures;  these  young 
animals  have  pelage  that  is  usually  softer  and 
grayer  than  that  of  adults  and  are  noticeably 
lower  in  weight,  and  females  are  usually  nullip- 
arous.  Young  adults  are  older;  they  have  unworn 
adult  pelage  and  have  nearly  completed  cranial 
growth  but  have  not  yet  reached  adult  weight 
and  usually  have  not  yet  reproduced  or  are 
pregnant  for  the  first  time.  Adults  have  complet- 
ed cranial  growth  and  have  adult  pelage,  and 
usually  the  females  are  multiparous.  Terminolo- 
gy for  description  of  external  features  follows 
Brown  (1971)  and  Brown  and  Yalden  (1973). 
Terminology  for  cranial  and  dental  features 
follows  Musser  and  Heaney  (1992).  Scanning 
electron  micrographs  of  skulls  and  teeth  were 
made  with  uncoated  specimens. 

External  measurements  (total  length,  tail 
length,  hind  foot  length,  length  of  ear  from 
notch,  weight  in  grams)  were  taken  from 
collectors'  field  catalogs  or  specimen  labels. 
Fourteen  cranial  measurements  (Table  1)  were 
taken  with  digital  calipers  to  the  nearest  0.01  mm 
by  Heaney;  comparisons  made  in  the  text  refer 
only  to  specimens  also  measured  by  Heaney. 


Results 

The  endemic  Philippine  genus  Apomys  was 
described  by  Mearns  (1905)  to  contain  three 
species:  A.  hylocoetes  (as  the  type  species),  A. 


HEANEY  AND  TABARANZA:  A  NEW  SPECIES  OF  FOREST  MOUSE 


15 


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d  on'  d  ^  d  «S  dm  cci  d  °.  d  ^  d  ^  d  d  d  ^  d  ^  d  °^  r-'  2 

CnI  </^  —  —  —  tJ-  NC  —  —  nC  nC  r»  N 

+  1         +11  +1  +1         +1    :     +11     +1         +11+1    l  +11+1     ,    +1    l     +1    I 

tj-  —    oc  oc  rj  nC  r*".  -sj-    Or*".    oiN    iT;  x    O  O  Mr,  "st  v,    r*",  ^    O  t~~-    x  r,    <r4  r~J    Cr    —  —    C" 

1 1^  - :  tt  °.  tj-"  °°.  <n  "*.  —  x.  r-;  •"i  ^  -.  h  ^  on"  °.  w-;  ^  nc'  ^:  no  t^  ^  d  -  —      d      no" 

x  n    t.  w~i  —  rsi  —    cn)  —    m         nc         <->-.  —  on  nC         no         r^         ^         ^         f.         cn         r-, 

rN  —  —         —  —  <N         — 

vCOO  it,  Jn-  ^  N    C  ifl    M  v^  (N  X  it,  C  (N  —  o-,  ^O 

^t  c   no  m  r^  —  ■*   r<-.  o   rs  rs   —  NC-str-  —  rs  c.  r«-,   —  c   rs  iri   r  ^  n  -   <n  o 

d  on'  d  "1  d  vi  d  rn  d  rn  d  ^  d  ^  d  rn  d  d  d  "J  d  ^  d  ^  dc  uS  _  — '  c,        cm  r^  m 

+  1     '     +1  +1     ■  +1    J     +1  +lT    +|7    +1     :  +1  +1  +11     +1     I     +|7    +|     I     +1     I    2     !     +lZ 

—  r^  —  OONO^-r*-,  nC  —  r,  r,  O  r  ?■    -x  ^t 

3NONinONrNiONt^r-'Ni-o-^->^ooo^3-ONr~-N3    " 

^rN^.^VlTf^N^ri00-"'  ^   ^  -i  <*  ~  ° 

3crsiw~> r*~.  —    rsi  —    r< 

Cs|  _  — 


:  nc 


i/~-  d    On  ^  O  f*"'    r-  rNj    CM  —    v,  r»% 

<-:r.-    _JfM  fN- 

r-         ir,  tj- 

rN  — 


i^tj-u-,         ONrNirNi-^-ONr^-^-         on         onon   —  r~~o         —         no 

■^-r-    rsi  —   f,  r,   -Nt  t~~    -^-nC    —  •<*■    ex    t  ?-    t*~,  no    rN  — —  no 

•    -:  nT-,    _;      ■    _;      •    _•  _•    _■  oc    _•  rr-.    _;  _•    _:  _•    _:  —    _;  io    _;  in 


-X    O:    Om    CM    O  —    O 
fN  w-i  —  —  — 

+  1  _    +1     I,    +!  +1  +1  +1 


O—    OO     O.    O.    O.    nCnC    ■*  i/-,    —  Tj-    —  O    r<",  rs) 


rN     ,  ,  — 


r~-oc    ton    it,  ^    m  on    nc-^-    ocm    no  —    —  oc    oo*    't  ?-    it,  (N    f.  r%i    oc-st    r-.  tj-    >^-,  r<-,    o  —    inc. 


"*.  —  P  d  ^°. 


—  O    r~  f*~,    r-,  o    ■<*■ 


"tN    —  On    c.  rsi    o 

o  on   d  7   d :  w!   d  rN 


rr  oc    rsi  oc    —  3 


ON  OC  —  >/~, 

2    «~>     <S~,    —      X   Tf     T3-   l~- 

n—  -i^    rN  i,-    oc  ,^.    oc  rv» 


— ; :    rN  _■    o 


r*~.  tj  cnu-,  —  r-   —  r-~   rN^j-   -<3-v^   -^--^  tj- 

o  ^  d(N  do  d  ^  d  *?  o"  "~;  ^  S  t^  2  d  ^ 

—  —  NONCr-rN          —          r^, 

+  1  .  +1  A  +!  J.    +1  ^    +1    I     +1^+1^  +1 


- 


on    o  tj-    rsi 


T  —    rN  — 


—  Tj-  © 

_    VI    rNO     ooc    </-,  —    CN3N  NIT,    OC-^"    (Nf, 

Tt  ^    q_   N--    CX   ^f  ,r.-    m  ^  _•  rN 
nC         rN  —    o  —    no 


^  oc" 

+11" 


i  n3    o 


5      - 


S3 


^         22 


r^ 


rN         — 


=        —        N 


.2? 

3  '5 


16 


FIELDIANA:  ZOOLOGY 


insignis,  and  A.  petraeus  (the  last  synonymized 
with  A.  hylocoetes  by  Musser,  1982),  all  from  Mt. 
Apo,  from  which  the  name  was  derived.  As 
related  by  Musser  (1982),  additional  species  were 
named  from  Luzon  and  Catanduanes,  and  the 
genus  was  found  throughout  much  of  the 
archipelago  (Ruedas,  1995;  Heaney  et  al., 
1998).  However,  because  the  initial  description 
was  vague  and  many  genera  of  Indo-Australian 
rodents  poorly  studied,  Apomys  was  briefly 
synonymized  with  the  genus  Rattus  at  a  time 
when  most  Indo-Australian  murids  were  placed 
in  that  genus.  Musser  (1982)  thoroughly  rede- 
scribed  Apomys,  pointing  out  its  many  distinctive 
characters,  and  redefined  the  species  then 
known.  Musser  and  Heaney  (1992)  further 
defined  and  compared  Apomys  to  other  Philip- 
pine murids,  and  they  pointed  out  its  apparent 
close  relationship  to  Chrotomys,  Celaenomys, 
and  Rhynchomys,  also  endemic  to  the  Philip- 
pines. Using  data  from  Musser  and  Heaney 
(1992),  Heaney  and  Rickart  (1990)  postulated 
that  Apomys  was  basal  to  the  clade  including 
Chrotomys,  Celaenomys,  and  Rhynchomys  and 
noted  the  diversification  of  this  clade  within  the 
Philippines  as  an  example  of  adaptive  radiation. 
Heaney  et  al.  (1998)  noted  the  presence  of  many 
undescribed  species  of  Apomys,  including  the 
species  from  Camiguin  reported  by  Heaney  and 
Tabaranza  (1997). 

Rickart  and  Heaney  (2002)  showed  that  Ap- 
omys hylocoetes,  A.  insignis,  and  the  Leyte  Apomys 
(as  well  as  most  others  from  Greater  Luzon  and 
Greater  Negros-Panay)  have  distinctive  karyo- 
types, but  the  Apomys  from  Camiguin  has  not 
been  karyotyped.  Steppan  et  al.  (2003)  used 
molecular  data  to  assess  phylogenetic  relation- 
ships and  geographic  patterns  of  diversification 
within  Apomys.  Analysis  of  variation  in  cyto- 
chrome^ in  10  species  (Fig.  1)  showed  the 
presence  of  three  major  clades:  one  containing 
Apomys  datae  and  A.  gracilirostris;  a  second  clade 
containing  A.  microdon,  A.  musculus,  and  two 
undescribed  species  from  Negros  and  Sibuyan  (the 
"Greater  Luzon,  Mindoro,  and  Negros  clade"); 
and  a  third  clade  containing  A.  hylocoetes  and  A. 
insignis  from  Mindanao,  plus  an  undescribed 
species  from  Leyte,  Biliran,  and  Bohol  and 
another  from  Camiguin  ( the  "Greater  Mindanao 
clade").  The  second  and  third  of  these  three  clades 
form  a  monophyletic  clade  that  Musser  (1982) 
described  as  being  "species  ...  of  small  or 
medium  body  size  in  which  the  canal  for  the 
internal  maxillary  artery  [also  described  as  the 


rC" 

I A 


A.  "A/C"  (Negros) 
"B"  (Sibuyan) 


r 


c 


A.  musculus 
A.  microdon 
A.  hylocoetes 
A.  insignis 

A.  "F"  (Leyte) 

A.  camiguinensis 

A.  datae 

A.  gracilirostris 

Rhynchomys  isarogensis 

Chrotomys  gonzalesi 


Fig.  1.  Hypothesis  of  phylogenetic  relationships 
within  the  genus  Apomys  based  on  parsimony  analysis 
of  cytochrome-b  molecular  data  (based  on  Steppan  et 
al.,  2003). 


infraorbital  branch  of  the  stapedial  artery]  is 
partially  open  and  part  of  the  artery  is  exposed 
on  the  ventral  surface  of  each  pterygoid  plate. 
That  feature,  combined  with  bright  tawny 
upperparts,  buffy  underparts,  brown  or  pale 
buffy  feet,  and  patterning  on  the  tail  (usually 
monocolored  or  mottled,  rarely  sharply  bi- 
colored)  suggest  the  five  species  [known  in 
1982]  may  be  more  closely  related  to  each  other 
than  to  any  others  in  Apomys."  Musser's  (1982) 
recognition  and  diagnosis  of  this  clade,  based 
on  very  few  specimens,  was  prescient  and 
remains  accurate.  In  the  analysis  by  Steppan 
et  al.  (2003),  the  species  from  Camiguin  is  basal 
to  the  others  in  the  "Greater  Mindanao  clade" 
(Fig.  1),  and  they  estimated  the  time  of 
divergence  of  the  Camiguin  mouse  from  the 
other  members  of  its  clade  at  about  2.3  million 
years  ±  about  25%. 

We  note  that  the  Apomys  from  Biliran,  Bohol, 
and  Leyte  (which  we  hereafter  refer  to  as  "the 
Leyte  Apomys''')  was  tentatively  considered  to 
represent  either  A.  microdon  (by  Musser,  1982) 
or  A.  littoralis  (Rickart  et  al.,  1993;  Heaney  et  al., 


HEANEY  AND  TABARANZA:  A  NEW  SPECIES  OF  FOREST  MOUSE 


17 


Fig.  2.     Photograph  of  an  adult  Apomys  camiguinensis,  taken  on 
March  1995. 


Mt.  Timpoong,  Camiguin  Island,  in 


1998).  but  the  combination  of  the  morphological 
data  presented  here,  karyotypic  data  (Rickart  & 
Heaney,  2002).  and  molecular  data  (Steppan  et 
al.,  2003)  have  led  us  to  conclude  that  the  Leyte 
Apomys  is  a  distinct  species;  further  details  and 
description  will  be  published  elsewhere. 

The  specimens  from  Camiguin  are  morpho- 
logically similar  to  series  of  A.  hylocoetes  and  A. 
insignis  from  Mindanao  and  the  undescribed 
species  from  Leyte  and  Biliran  but  readily 
distinguished  from  all  three  of  these  species  on 
the  basis  of  external  and  cranial  features.  Heaney 
et  al.  (1998)  and  Steppan  et  al.  (2003)  referred  to 
this  animal  informally  as  "'Apomys  sp.  D."  We 
now  name  the  species  from  Camiguin  as  Apomys 
camiguinensis,  new  species. 


Apomys  camiguinensis,  new  species 

Holotype  Adult  male,  fmnh  167878,  collect- 
ed 16  May  1994  by  B.  R.  Tabaranza,  Jr. 
Specimen  originally  fixed  in  formalin,  trans- 
ferred to  im  ethyl  alcohol  with  skull  removed 


and  cleaned.  Specimen  is  currently  on  deposit  at 
fmnh  and  is  to  be  transferred  to  nmp. 

Type  Locality — Barangay  Kital-is,  Sagay  Mu- 
nicipality, 2  km  N,  62  km  W  Mahinog,  1000  m 
elevation,  Camiguin  Province,  Camiguin  Island. 
9°9.5'N,  124°43.5'E  (see  Heaney  &  Tabaranza, 
2006,  for  further  details). 

Referred  Specimens  and  Localities — In  addi- 
tion to  the  holotype,  19  paratypes  are  known 
from  three  localities  ranging  from  1000  to 
1400  m  (fmnh  154815-154816,  154854-154860, 
167878-167882,  plus  6  specimens  at  msu-iit);  for 
localities,  see  Heaney  et  al.  (2006).  All  were 
originally  preserved  in  formalin  and  are  now 
stored  in  ethyl  alcohol,  many  with  skulls  re- 
moved and  cleaned,  or  were  prepared  as 
complete  skeletons.  Tissue  samples  are  housed 
at  fmnh.  Half  of  the  specimens  will  be  deposited 
at  nmp. 

Distribution — Known  only  from  the  upper 
elevations  on  Mt.  Timpoong,  Camiguin  Island, 
but  probably  occurring  throughout  the  montane 
and  mossy  rain  forest  on  Camiguin  Island  (see 
fig.  2  in  Heaney  et  al.,  2004)  and  possibly  at 
lower  elevations. 


IS 


FIELDIANA:  ZOOLOGY 


Measurements — Table  1. 

Etymology — The  specific  name  refers  to  the 
sole  island  on  which  the  species  is  found.  We 
suggest  the  common  English  name  "Camiguin 
forest  mouse." 

Diagnosis — A  species  of  the  genus  Apomys,  as 
defined  by  Musser  (1982)  and  Musser  and 
Heaney  (1992),  including  the  following  distinc- 
tive generic  features:  rostrum  long  and  moder- 
ately narrow;  viewed  laterally,  rostrum  with 
a  rectangular  shape,  with  premaxillaries  project- 
ing well  anterior  to  the  anterior  edge  of  the 
upper  incisors;  incisive  foramina  broad  relative 
to  length;  bony  palate  wide  and  long,  densely 
pitted  and  perforated;  upper  third  molar  re- 
duced to  a  large  round  peg;  lower  third  molar 
also  peg-like  but  retaining  an  anterior  lamina, 
without  evidence  of  the  two  cusps  that  usually 
form  this  lamina;  occlusal  surface  of  each  first 
and  second  upper  molar  consisting  of  two  simple 
chevron-shaped  laminae  followed  by  a  small  oval 
lamina,  without  evidence  of  cuspidation;  audito- 
ry bulla  separated  from  the  squamosal  and 
alisphenoid  by  a  gap  that  is  formed  by  the 
coalescence  of  the  postglenoid  foramen,  the 
postalar  fissure,  and  the  middle  lacerate  fora- 
men. 

As  described  in  more  detail  in  the  following 
section,  the  Camiguin  mouse  is  defined  by  the 
following  characters  or  unique  combination  of 
characters:  moderate  body  size  but  somewhat 
robust  build  for  the  genus  overall;  the  tail  is  long 
relative  to  body  length,  with  unusually  large 
scales,  and  more  often  with  a  sharp  transition 
from  dark  brown  dorsum  to  pale  brown  venter 
(i.e.,  sharply  bicolored)  than  in  other  species; 
moderately  long  but  unusually  broad  hind  foot 
with  small  plantar  pads,  and  slightly  shorter  ear. 
The  pelage  has  more  brown  in  the  generally 
russet-brown  dorsum  than  on  its  closest  relatives 
and  has  more  conspicuous  salt-and-pepper 
speckling  dorsally;  the  mystacial  and  genal 
vibrissae  are  long  but  moderate  for  the  genus. 
The  cranium  has  an  unusually  long  braincase 
and  orbit,  somewhat  narrow  zygomatic  and 
mastoid  width,  and  a  moderately  long  but  deep 
and  robust  rostrum.  The  palate  is  rather  narrow, 
the  posterior  edge  of  the  palate  extends  un- 
usually far  posterior  to  the  last  molar,  the 
incisive  foramina  long  and  wide,  and  the  distance 
from  the  posterior  edge  of  the  incisive  foramina 
is  unusually  short.  The  longest  axis  of  the  bullae 
is  about  35°  from  the  midline  axis  of  the  skull. 
The  toothrows  are  of  moderate  size;  the  ante- 


rolabial  cusp  of  the  third  upper  molar  is  barely 
evident  in  most  individuals. 

Description  and  Comparisons — Apomys  cami- 
guinensis  is  an  attractive  mouse  with  large  eyes 
and  ears,  long  tail,  and  soft  pelage  (see  Frontis- 
piece, this  volume,  and  Fig.  2).  As  with  other 
members  of  the  genus,  the  pelage  is  soft  and 
dense,  without  spines  or  stiff  hairs.  The  dorsal 
coloration  is  a  rich  brownish-russet  with  a  small 
amount  of  salt-and-pepper  speckling;  underfur  is 
pale  slate-gray.  The  venter  is  paler,  usually 
nearly  white  with  a  wash  of  buffy  or  pale  russet, 
but  some  individuals  have  blazes  of  pure  white 
(usually  on  the  chest)  or  are  much  darker  brown 
or  russet-brown.  There  is  a  narrow  area  of  bare 
skin  around  the  eyes;  the  ears  are  moderately 
dark  brown,  with  short  hairs  apparent  on  the 
outer  surface,  and  present  but  tiny  and  nearly 
invisible  on  the  inner  surface.  The  mystacial 
vibrissae  are  long  and  conspicuous.  The  dorsal 
surface  of  the  fore  and  hind  feet  are  mostly  buffy 
or  pale  brown  but  with  a  narrow  band  of 
scattered  darker  hairs  around  the  midline,  and 
these  decrease  in  number  and  length  toward  the 
distal  end  of  each  foot  (with  only  pale  hairs  on 
the  dorsal  surface  of  the  toes).  The  feet  are  lightly 
pigmented  or  unpigmented  on  the  ventral 
surface,  with  conspicuous  plantar  pads  on  the 
ventral  surface  (Fig.  3).  The  tail  is  long  with 
conspicuous  scales;  fine  hairs  that  are  present 
between  the  scales  are  most  visible  on  the  dorsal 
and  lateral  surfaces  and  least  visible  ventrally. 
The  tail  is  darker  on  the  dorsal  surface  than  on 
the  ventral  surface.  The  scrotum  of  adult  males  is 
fairly  small  and  projects  beyond  the  abdomen 
only  partially  on  the  posterior  portion  and  has 
black  or  dark  brown  pigment  at  the  posterior  tip, 
about  3-5  mm  in  length.  Females  have  two  pairs 
of  inguinal  mammae. 

Apomys  camiguinensis  is  easily  distinguished 
from  most  members  of  the  genus  by  its  in- 
termediate size  (only  A.  insignis  and  A.  hylo- 
coetes  are  similar)  and  from  those  two  species  by 
both  external  and  cranial  characters.  Apomys 
camiguinensis  has  total  length  (average  254— 
260  mm)  slightly  greater  than  A.  hylocoetes 
(248-252  mm)  and  A.  insignis  (251-252  mm) 
and  substantially  greater  than  the  Leyte  Apomys 
(238-245  mm;  Table  1).  The  average  tail  length 
(146-148)  is  equal  to  that  of  A.  insignis  (147— 
148  mm),  and  substantially  longer  than  that  of 
A.  hylocoetes  (141-142  mm)  and  the  Leyte 
Apomys  (140-145  mm).  The  tail  averages  57% 
of  the  total  length  in  A.  camiguinensis,  compared 


HEANEY  AND  TABARANZA:  A  NEW  SPECIES  OF  FOREST  MOUSE 


19 


Fig.  3.  Ventral  surface  of  the  right  hind  feet  of  (A)  Apomys  camiguinensis,  (B)  A.  hylocoetes,  (C)  A.  insignis, 
and  (D)  the  undescribed  Apomys  from  Leyte,  all  to  roughly  the  same  scale.  Those  of  A.  hylocoetes  and  A.  insignis 
are  redrawn  from  Musser  (1982). 


to  56.5%  in  A.  hylocoetes,  58%  in  A.  insignis,  and 
59%  in  the  Leyte  Apomys.  The  hind  foot  (about 
33  mm)  is  about  equal  in  length  to  that  of  A. 
insignis  (33  mm)  and  is  substantially  longer  than 
in  A.  hylocoetes  (31-32  mm)  and  the  Leyte 
Apomys  (30-31  mm);  the  hind  foot  of  A. 
hylocoetes  is  notably  the  broadest  (Fig.  3;  see 
also  fig.  7  in  Musser,  1982).  Ear  length  is  greatest 
in  A.  hylocoetes  (20.2  mm),  with  A.  insignis 
(19.5  mm),  A.  camiguinensis  (19.0  mm),  and  the 
Leyte  Apomys  (18-19  mm)  progressively  slightly 
smaller  (Table  1 ).  The  weight  of  A.  camiguinensis 
(38-41  g)  averages  the  greatest  of  the  four, 
followed  by  A.  hylocoetes  (36-39  g),  A.  insignis 
(37  38  g),  and  the  Leyte  Apomys  (28-31  g).  In 
other  words,  the  Camiguin  Apomys  is  relatively 
heavy  and  long  and  has  a  relatively  long  tail  (but 
slightly  less  long  proportionately  than  some  close 
relatives),  a  moderately  long  hind  foot,  and 
somewhat  short  ear. 

The  following  qualitative  external  characters 
also  distinguish  Apomys  camiguinensis  from  its 
three  closest  relatives.  The  dorsal  coloration  of 
the  Camiguin  mouse  is  less  russet  and  more 
brown  than  in  A.  hylocoetes  and  A.  insignis,  with 


more  of  the  salt-and-pepper  appearance;  the 
Leyte  mouse  is  dorsally  brighter  red  than  the 
others,  with  more  red  and  orange  in  the  russet 
than  the  other  three  and  almost  no  salt-and- 
pepper.  Ventral  coloration  is  generally  similar  in 
all  four,  though  with  the  variation  noted  above, 
but  A.  hylocoetes  tends  to  have  more  of  an 
orange  wash  than  the  others.  The  dorsal  surface 
of  the  hind  feet  usually  is  palest  in  the  Leyte 
mouse  and  darkest  in  the  Camiguin  mouse.  The 
mystacial  vibrissae  are  long  on  all  four  species 
but  longest  on  A.  insignis  (up  to  56-60  mm 
maximum),  on  which  they  reach  past  the  middle 
of  the  back,  and  intermediate  on  the  Camiguin 
mouse  (52-55  mm),  A.  hylocoetes  (51-55  mm), 
and  Leyte  mouse  (50-55  mm).  Genal  vibrissae 
reach  to  the  anterior  edge  of  the  largest  lateral 
pad  on  the  Camiguin  and  Leyte  mice  but  farther 
forward,  to  the  base  of  the  toes,  on  A.  hylocoetes 
and  A.  insignis. 

The  hind  foot  (Fig.  3)  differs  markedly  among 
the  four.  The  foot  of  A.  camiguinensis  is  about 
the  same  length  as  that  of  A.  hylocoetes  but  is 
broader  and  has  smaller  plantar  pads.  The  hind 
foot  of  the  Leyte  mouse  is  proportioned  similarly 


20 


FIELDIANA:  ZOOLOGY 


Fig.  4.  Photographs  of  dorsal,  ventral,  and  lateral  views  of  the  crania  of  Apomys  camiguinensis  (A;  fmnh 
167878,  holotype),  A.  hylocoetes  (B;  fmnh  148146),  A.  insignis  (C;  fmnh  147092),  and  the  undescribed  Apomys 
from  Leyte  (D;  ummz  160290),  all  to  the  same  scale. 


HEANEY  AND  TABARANZA:  A  NEW  SPECIES  OF  FOREST  MOUSE 


21 


Fig.  5.  Scanning  electron  micrographs  of  the  ventral  view  of  the  anterior  portion  of  the  skulls  of  Apomys 
eamiguinensis  (A;  fmnh  167878,  holotype),  A.  hylocoetes  (B;  fmnh  148146),  A.  insignis  (C;  fmnh  147092),  and  the 
undescribed  Apomys  from  Leyte  (D;  ummz  160290),  all  to  same  scale. 


to  A.  hylocoetes  but  is  smaller.  The  hind  foot  of        sturdy  and  sharply  pointed;  those  of  A.  hylo- 


A.  insignis  is  very  long  and  narrow,  with  plantar 
pads  of  moderate  size.  On  all  species,  the  claws 
on  all  five  digits  are  unpigmented  and  laterally 
compressed.  On  A.  eamiguinensis,  the  claws  are 


eoetes  are  slightly  thinner  and  more  sharply 
pointed  and  the  digits  bearing  them  slightly  more 
slender.  The  digits  and  claws  of  A.  insignis  are 
shorter  and  more  slender;  the  digits  and  claws  of 


FIELDIANA:  ZOOLOGY 


Fig.  6.  Scanning  electron  micrographs  of  the  ventral  view  of  the  posterior  portion  of  the  skulls  of  Apomys 
camiguinensis  (A;  fmnh  167878,  holotype),  A.  hylocoetes  (B;  fmnh  148146),  A.  insignis  (C;  fmnh  147092),  and  the 
undescribed  Apomys  from  Leyte  (D;  ummz  160290),  all  to  same  scale. 


the  Leyte  mouse  are  still  smaller  and  more 
slender.  The  forefeet  of  A.  camiguinensis  have 
smaller  pads  than  those  of  the  others  (but  only 
proportionately  in  the  case  of  the  generally 
smaller  Leyte  mouse),  but  the  feet  are  slightly 
broader  and  more  robust  than  in  the  other 
species.  All  four  species  have  a  flat,  unpigmented 
nail  on  the  pollex.  The  relative  size  and  thickness 
follows  the  same  pattern  as  the  hind  feet,  with  A. 


camiguinensis  being  the  most  robust,  A.  hylo- 
coetes slightly  shorter  and  thinner,  A.  insignis 
much  shorter  and  more  slender,  and  the  Leyte 
mouse  much  like  A.  insignis  but  a  bit  smaller 
overall. 

The  tail  scales  on  the  Camiguin  mouse  are 
largest,  with  12-12.5  scales/cm  near  the  base, 
compared  to  14-15  scales/cm  in  A.  hylocoetes, 
13-14  in  A.   insignis,  and   13-15  in  the  Leyte 


HEANEY  AND  TABARANZA:  A  NEW  SPECIES  OF  FOREST  MOUSE 


Fig  7  Scanning  electron  micrographs  of  the  occlusal  surface  of  the  maxillary  toothrows  of  Apomys 
camiguinensis  (A;  fmnh  167878,  holotype),  A.  hylocoetes  (B;  fmnh  148146),  A.  insignis  (C;  fmnh  147092),  and  the 
undescribed  Apomys  from  Leyte  (D;  ummz  160290),  all  to  same  scale. 


Fig.  8.  Scanning  electron  micrographs  of  the  occlusal  surface  of  the  mandibular  molariform  toothrows  of 
Apomys  camiguinensis  (A;  fmnh  167878,  holotype),  A.  hylocoetes  (B;  fmnh  148146),  A.  insignis  (C;  fmnh  147092), 
and  the  undescribed  Apomys  from  Leyte  (D;  ummz  160290),  all  to  same  scale. 


24 


FIELDIANA:  ZOOLOGY 


mouse.  The  size  of  the  scales  remains  about  the 
same  more  distally  on  a  given  species  to  about 
the  midpoint;  from  there  to  the  tip  scale  size 
decreases,  mostly  in  the  last  quarter  of  the 
length,  to  about  half  the  length  (and  one-fourth 
the  size)  of  scales  near  the  base  of  the  tail.  The 
scales  of  the  tail  are  dark  brown  dorsally  and 
nearly  white  ventrally  on  all  four  species,  but  the 
transition  is  most  often  an  abrupt  line  on  the 
Camiguin  mouse,  often  producing  a  sharply 
bicolored  tail,  and  most  often  gradual  on  the 
other  three  species.  Typically,  three  hairs  grow 
from  beneath  each  scale,  projecting  only  slightly 
laterally  from  the  tail;  on  the  basal  quarter  of  the 
tail,  one  of  the  three  hairs  is  missing  from  a  given 
scale  about  one-third  of  the  time.  On  A. 
camiguinensis,  the  hairs  are  about  one  and  one- 
third  the  length  of  a  scale  near  the  base  of  the 
tail,  have  about  the  same  length  25%  toward  the 
tip,  are  slightly  longer  at  the  midpoint,  are  about 
one  and  two-thirds  the  length  of  a  scale  75% 
toward  the  tip,  and  are  about  five  times  the 
length  of  a  scale  near  the  tip,  where  scale 
length  has  been  reduced  by  about  half  relative 
to  the  base.  On  A.  hylocoetes,  hairs  are  about 
one  and  a  half  the  length  of  a  scale  near  the  base 
of  the  tail,  are  similar  25%  toward  the  tip,  are 
about  twice  the  length  of  a  scale  near  the 
midpoint,  are  about  two  and  a  half  times 
the  length  of  a  scale  75%  toward  the  tip,  and 
are  more  than  five  times  the  length  of  a  scale 
near  the  tip.  On  A.  insignis,  hairs  are  one  and 
one-third  a  scale  length  near  the  tail's  base, 
similar  at  25%,  one  and  two-thirds  near  the 
midpoint,  about  two  and  one-half  of  a  scale 
length  near  75%,  and  about  three  and  one-half 
the  length  of  a  scale  near  the  tip.  On  the  Leyte 
mouse,  tail  hairs  are  slightly  less  than  the 
length  of  a  scale  near  the  base  of  the  tail,  similar 
at  the  midpoint  and  at  75%,  and  about  two  times 
the  length  of  a  scale  near  the  tip.  In  all  four 
species,  the  hair  becomes  slightly  greater  in 
diameter  distally  than  it  is  proximally;  combined 
with  the  trend  for  greater  length,  this  means 
that  the  tail  become  more  heavily  covered  with 
hair  distally.  None  of  the  species  shows  elongat- 
ed hairs  at  the  very  tip  of  the  tail  (i.e.,  there  is 
no  "pencilling").  Hairs  on  the  dorsal  surface 
are  more  heavily  pigmented  dorsally  than 
ventrally  in  all  four  species.  On  adults  of  all  four 
species,  the  dorsal  surface  of  the  tip  (2-5  mm)  of 
the  tail  becomes  worn,  with  most  scales  and 
many  hairs  absent,  leaving  a  smooth,  leathery 
surface. 


The  cranium  of  Apomys  camiguinensis  (Figs.  4- 
6)  has  basioccipital  length  (28.3-29.0  mm)  slight- 
ly greater  than  that  of  A.  hylocoetes  (28.3- 
28.8  mm),  clearly  more  than  A.  insignis  (27.8- 
28.0  mm)  and  substantially  more  than  the  Leyte 
Apomys  (25.3-26.3  mm);  the  interorbital  width  is 
proportioned  similarly  (Table  1).  Zygomatic 
breadth  averages  greatest  in  A.  hylocoetes 
(15.0-15.2  mm),  followed  by  A.  camiguinensis 
(14.9-15.0  mm),  A.  insignis  (14.7-14.8  mm),  and 
the  Leyte  Apomys  (13.1-13.7  mm),  with  mastoid 
breadth  following  the  same  pattern  (Table  1). 
Nasal  length  averages  substantially  greater 
in  A.  hylocoetes  (12.3-12.5  mm)  than  in  A. 
insignis  (11.6-11.9  mm),  A.  camiguinensis  (11.1- 
11.4  mm),  or  the  Leyte  Apomys  (10.3-10.7  mm), 
and  anterior  nasal  breadth  is  similarly  patterned 
(Table  1).  Rostral  depth  averages  greatest  in  A. 
camiguinensis  (6.3-6.4  mm),  with  A.  hylocoetes 
(6.2-6.3  mm)  and  A.  insignis  (6.3  mm)  slightly 
less  deep  and  very  similar  to  each  other  and  the 
Leyte  Apomys  generally  less  deep  (5.9-6.1  mm). 
Rostral  length  (Fig.  5)  is  clearly  greatest  in  A. 
hylocoetes  (13.1-13.2  mm),  with  A.  camiguinensis 
(11.6-12.0  mm)  and  A.  insignis  (11.9-12.0  mm) 
similar  to  each  other  but  much  greater  than  the 
Leyte  Apomys  (10.6-11.2  mm).  Orbital  length 
averages  greatest  in  A.  camiguinensis  (10.2- 
10.3  mm),  followed  by  A.  insignis  (10.0- 
10.2  mm)  and  A.  hylocoetes  (9.7-10.0)  and  the 
small  Leyte  Apomys  (9.2-9.6).  To  summarize,  A. 
camiguinensis  is  characterized  by  the  longest 
skull  and  deepest  rostrum,  but  A.  hylocoetes 
has  slightly  greater  zygomatic  and  mastoid  width 
as  well  as  greater  nasal  length  and  breadth  and 
rostral  length.  Although  the  rostrum  of  A. 
camiguinensis  is  moderate  in  length,  the  orbital 
region  and  braincase  are  unusually  long.  The 
cranium  of  A.  insignis  tends  to  be  smaller  but 
generally  similarly  proportioned  to  A.  camigui- 
nensis, and  the  Leyte  Apomys  is  smaller  in  all 
dimensions,  though  it  seems  to  follow  the  pattern 
of  A.  camiguinensis  and  A.  insignis. 

The  maxillary  toothrow  (Fig.  7)  of  A.  cami- 
guinensis (6.0  mm)  averages  nearly  identical  in 
length  to  that  of  A.  hylocoetes  (6.0  mm)  and  A. 
insignis  (6.0  mm),  and  all  are  much  greater  than 
that  of  the  Leyte  Apomys  (5.2-5.3  mm).  Palatal 
breadth  at  M1  (Fig.  4)  is  greatest  in  A.  insignis 
(6.5-6.6  mm),  followed  by  A.  hylocoetes 
(6.5  mm)  and  A.  camiguinensis  (6.3-6.5  mm) 
and  the  Leyte  Apomys  (5.7-5.8  mm).  Diastema 
length  (Figs.  4  and  5)  in  A.  camiguinensis 
(7.3  mm)  averages  slightly  greater  than  in  A. 


HEANEY  AND  TABARANZA:  A  NEW  SPECIES  OF  FOREST  MOUSE 


25 


hvlocoetes  (7.2  -7.3  mm),  more  than  A.  insignis 
(7.0  mm),  and  least  in  the  Leyte  Apomys  (6.5 
6.8  mm).  In  other  words,  the  maxillary  tooth- 
rows  of  the  three  larger  Apomys  are  all  similar, 
with  the  Leyte  Apomys  having  disproportionate- 
ly short  toothrow,  but  the  palate  of  A.  camigui- 
ncnsis  is  disproportionately  narrow  (as  is  that  of 
the  Leyte  Apomys),  and  that  of  A.  hvlocoetes  is 
disproportionately  wide. 

In  addition,  we  note  the  following  qualitative 
characters.  The  incisive  foramina  (Fig.  5)  are 
widest  in  A.  eamiguinensis  and  in  A.  hvlocoetes 
and  longest  in  A.  eamiguinensis  and  A.  insignis,  so 
that  the  area  of  the  foramina  is  greatest  in  A. 
eamiguinensis.  The  distance  from  the  posterior 
edge  of  the  incisive  foramina  to  a  line  between  the 
anterior  edges  of  the  first  maxillary  molar  is 
shortest  in  A.  eamiguinensis,  slightly  greater  in 
A.  insignis,  and  longest  in  A.  hvlocoetes  and 
the  Apomys  from  Leyte.  The  braincase  of  A. 
eamiguinensis  is  slightly  more  elongate  and  that  of 
the  Leyte  Apomys  proportionately  most  squarish 
among  the  four  species  (Fig.  6).  The  posterior 
edge  of  the  bony  palate  (Fig.  6)  extends  farthest 
posterior  to  the  posterior  edge  of  the  last 
maxillary  molar  in  A.  eamiguinensis,  slightly  less 
far  in  A.  hvlocoetes  and  the  Apomys  from  Leyte. 
and  least  far  in  A.  insignis.  The  long  axis  of  the 
bullae  (Fig.  6)  is  about  45  from  the  cranial 
midline  axis  in  the  Apomys  from  Leyte,  about  40 
in  A.  hvlocoetes  and  A.  insignis,  and  about  35c  in 
A.  eamiguinensis.  The  hard  palate  of  A.  hvlocoetes 
and  the  Leyte  Apomys  are  usually  most  heavily 
pitted  and  perforated  with  vacuities  and  A. 
eamiguinensis  and  A.  insignis  less  so  (Figs.  4  and 
7).  The  third  upper  molar  (Fig.  7)  has  an 
anterolabial  cusp  (probably  tl;  see  Musser  & 
Heaney,  1992,  p.  65)  that  is  well  developed  in  A. 
hvlocoetes,  less  conspicuous  in  A.  insignis,  and 
barely  evident  in  A.  eamiguinensis  and  the  Apomys 
from  Leyte.  The  first  upper  molar  of  A.  hvlocoetes 
tends  to  have  a  more  conspicuous  anterolingual 
cleft  than  do  the  other  three  species  (Fig.  7).  Both 
upper  and  lower  toothrows  of  A.  insignis  are  the 
most  massive  (Figs.  7  and  8),  with  A.  eamigui- 
nensis somewhat  less  massive,  A.  hvlocoetes  sub- 
stantially less  so,  and  the  Apomys  from  Leyte 
smallest  overall.  All  four  species  have  the  canal 
for  the  infraorbital  branch  of  the  stapedial  artery 
partially  open  and  part  of  the  artery  exposed  on 
the  ventral  surface  of  each  pterygoid  plate  (Fig.  6), 
as  noted  by  Musser  (1982). 

Ecology    Sec  Heaney  et  al.  (2006)  for  ecolog- 
ical information 


Discussion 

The  presence  of  Apomys  eamiguinensis  as  an 
endemic  species  on  a  small  island,  along  with  the 
additional  murine  rodent  Bullimus  gamay  (Rick- 
art  et  al.,  2002)  and  the  Hanging-Parrot  (Lor- 
iculus  sp.,  described  in  this  volume),  as  noted 
above,  clearly  indicates  the  importance  of 
Camiguin  Island  as  a  unique  center  of  biological 
diversity  that  is  worthy  and  in  need  of  conser- 
vation (Heaney  &  Tabaranza,  2005).  In  addition, 
the  distinctiveness  of  this  species  confirms  pre- 
dictions made  on  the  basis  of  biogeographic 
models  (Heaney,  2004)  of  the  expected  presence 
of  endemic  small  mammals  on  Camiguin.  Fur- 
ther studies  of  the  mammals,  birds,  and  other 
organisms  are  clearly  warranted  to  determine, 
for  example,  the  degree  of  genetic  difference 
from  closest  relatives  (most  of  which  occur  on 
Mindanao)  as  a  means  of  assessing  the  role  of 
colonization  and  gene  flow  in  determining 
patterns  of  species  richness  and  endemism  in 
the  Philippines.  In  other  words,  Camiguin 
represents  a  natural  experiment,  as  a  young 
oceanic,  volcanic  island  that  is  near  to  a  large, 
rich  source  of  species  (Mindanao),  in  which  we 
can  measure  the  impact  of  genetic  isolation  in 
animals  and  plants  of  varying  vagility  under 
standardized  conditions.  Such  studies  are  certain 
to  produce  new  insights  into  the  process  by 
which  biological  diversity  is  generated  in  the 
Philippines  and  in  other  oceanic  archipelagos. 


Acknowledgments 

We  thank  Rebecca  Banasiak  for  assistance  in 
photographing  skulls,  Betty  Strack  for  assistance 
with  making  scanning  electron  microscope 
images,  and  Lisa  Kanellos  for  preparation  of 
figures;  Figure  3D  was  drawn  by  Jodi  Sedlock. 
The  Protected  Areas  and  Wildlife  Bureau  of  the 
Philippine  Department  of  Environment  and 
Natural  Resources  (denr)  provided  encourage- 
ment and  permits,  and  the  Region  10  office  of 
the  denr  provided  logistical  support.  Specimens 
for  comparison  were  kindly  loaned  by  G.  Hess 
(dmnh),  P.  Myers  (ummz),  and  M.  Carleton,  L. 
Gordon,  and  H.  Kafka  (usnm).  The  manuscript 
was  much  improved  by  comments  from  D.  S. 
Balete,  G.  G.  Musser,  P.  Myers,  and  E.  A. 
Rickart.  Funding  was  provided  by  the  Marshall 
Field  Fund,  Ellen  Thorne  Smith  Fund,  and  the 


26 


FIELDIANA:  ZOOLOGY 


Barbara  Brown  Fund  for  Mammal  Research  of 
the  Field  Museum  of  Natural  History. 


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ecology  of  mammals  on  Leyte,  Biliran,  and  Maripipi 
Islands,  Philippines.  Fieldiana  Zoology,  n.s.,  72: 
1-62. 

Rickart,  E.  A.,  L.  R.  Heaney,  and  B.  R.  Tabaranza, 
Jr.  2002.  Review  of  Bullimus  (Muridae:  Murinae) 
and  description  of  a  new  species  from  Camiguin 
Island,  Philippines.  Journal  of  Mammalogy,  83: 
421^136. 

Ruedas,  L.  A.  1995.  Description  of  a  new  large-bodied 
species  of  Apomys  Mearns  1905  (Mammalia: 
Rodentia:  Muridae)  from  Mindoro  Island,  Philip- 
pines. Proceedings  of  the  Biological  Society  of 
Washington,  108:  302-318. 

Steppan,  S.,  C.  Zawadski,  and  L.  R.  Heaney.  2003.  A 
molecular  assessment  of  phylogenetic  relationships 
and  patterns  of  phylogenesis  in  the  Philippine  murid 
rodent  Apomys.  Biological  Journal  of  the  Linnean 
Society,  80:  699-715. 


HEANEY  AND  TABARANZA:  A  NEW  SPECIES  OF  FOREST  MOUSE 


27 


Synopsis  and  Biogeography  of  the  Mammals  of  Camiguin 
Island,  Philippines 

Lawrence  R.  Heaney,1  Bias  R.  Tabaranza,  Jr.,1  Danilo  S.  Balete,1,3  and 
Natalie  Rigertas1 


Abstract 

Biodiversity  surveys  in  the  1960s  and  1990s  on  Camiguin  Island,  a  geologically  young, 
volcanically  active  oceanic  island  surrounded  by  deep  water,  have  demonstrated  the  presence 
of  24  species  of  land  mammals.  Three  species  (one  insectivore  and  two  rodents)  are  not  native 
to  the  Philippines,  but  all  others  (one  insectivore,  12  bats,  one  monkey,  four  rodents,  two 
small  carnivores,  and  one  ungulate)  are  indigenous.  Among  those  captured  in  the  1990s  were 
two  previously  unknown  species  of  murid  rodents  in  the  genera  Apomys  and  Bullimus  that  are 
endemic  to  Camiguin.  The  discovery  of  two  new  species  on  such  a  small  island  (ca.  265  km  )  is 
remarkable;  Camiguin  is  currently  the  smallest  island  in  the  Philippines  known  to  have  unique 
species  of  mammals.  Total  species  richness  of  nonvolant  mammals  on  Camiguin  is  low,  but 
relative  to  islands  that  were  once  part  of  Pleistocene  Greater  Mindanao,  Camiguin  is  not 
depauperate.  However,  its  fauna  is  not  ecologically  balanced  in  the  same  way  as  the  faunas  of 
the  islands  that  were  part  of  Greater  Mindanao:  ground-living  shrews  (Crocidura)  and  rodents 
(Apomys,  Bullimus,  Crunomys,  and  Rat t us)  from  lowland  forest,  along  with  some  large 
mammals  (Macaco,  Paratloxwus,  and  Sus)  are  well  represented  on  Camiguin,  but  all  the 
arboreal  small  mammals  that  characterize  lowland  forest  on  Mindanao  (Sundasciurus, 
Exilisciurus,  Cynocephalus,  and  Tarsius),  ground-living  small  mammals  from  montane 
habitats  ( Urogale,  Podogymnura,  Batomys,  Limnomys,  and  Tarsomys),  and  one  large  mammal 
(Cervus)  are  absent.  Additionally,  at  least  two  genera  of  fruit  bats  (Haplonycteris  and 
Megaerops)  that  are  fairly  common  in  lowland  rain  forests  on  Mindanao  are  absent  on 
Camiguin.  The  presence  of  some  nonvolant  mammals  demonstrates  that  dispersal  across  the 
deep  but  narrow  intervening  channel  takes  place,  but  the  presence  of  two  species  endemic  to 
Camiguin  and  the  absence  of  other  species  that  are  present  on  nearby  Mindanao  implies  that 
dispersal  probably  is  rare.  The  Asian  house  shrew  (Suncus  murinus)  was  remarkably  abundant 
in  primary  forest  at  high  elevation;  this  species  has  also  been  found  to  be  abundant  in 
montane  primary  forest  on  Negros  Island,  which  also  has  low  total  species  richness.  Species 
richness  of  small  nonflying  mammals  was  greatest  at  fairly  high  elevation. 


1  Field  Museum  of  Natural  History,   1400  South  Introduction 
Lake  Shore  Drive.  Chicago.  IL  60605-2496.  U.S.A. 

2  Department  of  Biology,  Iligan  Institute  of  Tech-  Tne  Philippine  Islands  present  a  remarkable 
nology,  Mindanao  State  University,  Iligan  City,  theater  for  the  study  of  the  ecology  and  evolution 
Lanao  del  Norte.  Philippines.  of  mammalian  diversity.  Its  islands  range  in  size 

1  Laksambuhay   Conservation.    Inc.,    10241    Mt.  from  less  than  one  hectare  to  over  100,000  km2, 

Bulusan  Street,  U-2.  Los  Banos,  Laguna.  Philippines.  with  geological  age  varying  from  under  1  million 

28  FIELDIANA:  ZOOLOGY,  N.S.,  NO.  106,  APRIL  5,  2006,  PP.  28-48 


years  to  over  40  million  years.  These  islands 
represent  many  sets  of  historically  distinct  geo- 
logical units  of  remarkably  varied  origins;  some 
had  land-bridge  connections  to  the  Asian  main- 
land in  the  past  (those  of  the  Palawan  group), 
but  most  are  purely  oceanic  in  origin  (Heaney, 
1986,  1991b,  2000;  Heaney  &  Rickart,  1990; 
Hall,  1998,  2002).  The  mammals  that  have 
evolved  in  this  diverse  archipelago  include  at 
least  1 1 1  species  endemic  to  the  archipelago  out 
of  172  native  terrestrial  species;  with  endemism 
at  64%,  the  Philippine  fauna  is  one  of  the  most 
distinctive  in  the  world  (Mittermeier  et  al.,  1997, 
1999;  Heaney  &  Regalado,  1998).  While  most  of 
the  endemic  species  occur  on  the  large  islands  of 
Luzon,  Mindanao,  Mindoro,  Negros,  and  Pala- 
wan (e.g.,  Heaney,  1986,  1993,  2000;  Heaney  et 
al.,  1998;  Rickart  et  al.,  1998),  significant 
numbers  occur  on  the  smaller  islands  as  well, 
especially  those  surrounded  by  deep  water  (e.g., 
Heaney,  1986,  2004;  Goodman  &  Ingle,  1993; 
Heaney  &  Tabaranza,  1997;  Musser  et  al.,  1998). 
As  noted  by  Heaney  and  Tabaranza  (2006a), 
Camiguin,  an  island  of  265  km2  located  about 
10  km  north  of  Mindanao  in  the  Bohol  Sea,  is 
one  such  deep-water  island,  with  a  minimum 
depth  to  Mindanao  of  385  m.  It  is  steeply 
mountainous,  with  several  active  volcanic  cones 
that  reach  to  a  maximum  elevation  of  about 
1600  m.  A  series  of  biological  surveys  on 
Camiguin  in  the  late  1960s  that  focused  on  birds 
(see  Balete  et  al.,  2006)  also  yielded  some 
mammal  specimens,  and  an  earlier  report  on 
those  surveys  (Heaney,  1984)  concluded  that  the 
island  had  no  endemic  mammal  species  and  was 
depauperate.  Subsequent  studies  on  other  islands 
made  us  suspect  that  those  earlier  surveys  were 
incomplete  because  so  few  mammal  species  had 
been  obtained  and  because  the  number  of 
nonvolant  mammal  specimens  was  small  (thus 
indicating  limited  sampling  effort).  Further,  on 
the  basis  of  biogeographic  patterns  elsewhere  in 
the  Philippines,  we  predicted  the  presence  of 
several  endemic  small  mammals  on  Camiguin 
(Heaney,  2004).  To  investigate  the  hypotheses 
that  the  previously  measured  species  richness  was 
low  because  of  incomplete  surveys  and  that 
about  two  endemic  species  should  be  present,  we 
returned  to  Camiguin  briefly  in  1992  and  more 
extensively  in  1994  and  1995  to  conduct  addi- 
tional mammal  inventories  in  all  the  major 
habitats  along  the  elevational  gradient,  especially 
by  trapping  small  mammals  at  higher  elevations 
where  there  were  few  records  from  the  1 960s.  As 


indicated  in  a  brief  preliminary  report  (Heaney  & 
Tabaranza,  1997),  we  found  eight  additional 
species  on  the  island  that  are  widespread  in  the 
Philippines,  plus  two  previously  unknown  en- 
demic species  of  rodents.  The  purpose  of  this 
paper  is  to  summarize  the  results  of  the  1994  and 
1995  mammal  surveys  and  integrate  those  data 
with  information  from  the  1960s,  including 
information  on  habitat  associations,  relative 
abundance,  and  ecology  of  the  species. 


Methods 
Prior  Reports 

The  first  report  of  mammals  from  Camiguin 
Island  was  that  of  Gray  (1843),  who  reported 
Paradoxurus  hermaphroditus.  Three  field  teams 
from  Silliman  University  led  by  Dioscoro  S. 
Rabor  collected  mammals  on  Camiguin  in  1967, 
1968,  and  1969;  specimens  were  deposited  at  the 
Delaware  Museum  of  Natural  History  (dmnh) 
and  Royal  Ontario  Museum  (rom);  for  details, 
see  Heaney  and  Tabaranza  (2006a).  Several 
specimens  were  reported  by  Peterson  and  Fenton 
(1970);  all  known  specimens  from  the  1960s  were 
examined  and  summarized  by  Heaney  (1984).  All 
data  included  in  this  paper  from  the  1960s 
specimens  are  based  on  data  in  Heaney  (1984), 
except  as  noted  below. 

Recent  Data 

Field  studies  were  conducted  during  three 
periods  in  1992,  1994,  and  1995;  general  methods 
and  site  descriptions  are  given  in  Heaney  and 
Tabaranza  (2006a).  Sampling  during  1995  fol- 
lowed methods  used  on  Leyte,  Luzon,  Negros, 
and  other  islands  (Heaney  et  al.,  1989,  1999; 
Rickart  et  al.,  1991,  1993)  to  facilitate  quantita- 
tive comparisons.  Nonvolant  small  mammals 
were  caught  in  traps;  during  1995,  all  traps  were 
Victor  rat  snap  traps.  Most  were  baited  with 
fresh  fried  coconut  coated  with  peanut  butter, 
but  a  few  were  baited  with  live  earthworms. 
During  1994,  several  National  live  traps  were 
used,  in  addition  to  Victor  rat  traps,  and  were 
baited  with  coconut  bait.  Bats  were  captured  in 
12-m  mist  nets.  Voucher  specimens  were  pre- 
pared in  fluid  or  as  skeletons  and  have  been 
deposited  at  The  Field  Museum  of  Natural 
History  (fmnh),  National  Museum  of  the  Phi- 


HEANEY  ET  AL.:  THE  MAMMALS  OF  CAMIGUIN  ISLAND 


29 


lippines  (nmp),  and  Mindanao  State  University- 
lligan  Institute  ol'  Technology  (msu-iit).  Most 
specimens  were  autopsied  for  reproductive  in- 
formation. The  si/e  of  embryos  was  measured  as 
crown  to  rump  length  (CRL).  Subadult  animals 
are  defined  here  as  those  that  have  not  completed 
cranial  growth,  especially  those  having  unfused 
basicranial  sutures;  these  young  animals  have 
pelage  that  is  usually  softer  and  grayer  than  that 
of  adults  and  are  noticeably  lower  in  weight  and 
females  are  usually  nulliparous.  Young  adults 
are  older;  they  have  nearly  completed  cranial 
growth  but  have  not  yet  reached  adult  weight 
and,  usually,  have  not  yet  reproduced  or  are 
pregnant  for  the  first  time.  Adults  have  complet- 
ed cranial  growth  and  adult  pelage,  and  usually 
the  females  are  multiparous.  Comments  on 
distribution  and  use  of  scientific  names  are  based 
on  Heaney  et  al.  (1998)  unless  additional  sources 
are  mentioned.  Records  of  specimens  examined 
are  summarized  at  the  end  of  each  account;  such 
summaries  include  site  number  and  the  number 
of  specimens  (in  parentheses). 

External  measurements  and  weights  reported 
here  were  taken  in  the  field  by  members  of 
the  field  team  on  fresh  animals.  Cranial  mea- 
surements were  taken  by  Heaney  with  digital 
calipers  graduated  to  0.01  mm.  Comparisons  of 
cranial  measurements  are  to  published  records  of 
specimens  measured  in  the  same  manner  by 
Heaney. 


Accounts  of  Species 

Order  Insectivora 

Family  Soricidae — Shrews 

Crocidura  beatus  Miller,  1910 

The  Mindanao  shrew  is  widespread  on  islands 
in  the  Mindanao  Faunal  Region  (Heaney  & 
Ruedi,  1994;  Heaney  et  al.,  1998);  these  are  the 
first  records  from  an  island  that  was  not  part  of 
the  late  Pleistocene  island  of  Greater  Mindanao 
(Heaney.  1986).  This  shrew  has  most  often  been 
found  in  primary  forest,  especially  at  higher 
elevations;  is  usually  uncommon  in  secondary 
forest;  and  is  absent  outside  of  forest  (Heaney  et 
al..  1989;  Rickart  et  al.,  1993). 

Crocidura  beatus  was  trapped  on  Camiguin  at 
three  forest  sites  in  May  1994  and  March  1995 
(Fig.  1.  Table  1).  It  was  uncommon  in  secondary 
lowland  forest  at  1000  m  elevation  (Site  4),  in 
disturbed    lower    montane    forest    at     1200  m 


elevation  (Site  5),  and  in  primary  montane  forest 
at  1275  m  (Site  7).  It  was  most  often  trapped 
under  tree  roots  and  live  vegetation.  None  were 
taken  in  agricultural  areas  at  150  m  (Site  3),  or  in 
mossy  forest  at  1475  m  (Site  8,  where  there  was 
limited  sampling;  Table  1 ).  This  use  of  habitat  is 
consistent  with  data  from  islands  on  Greater 
Mindanao  (Rickart  et  al.,  1993;  Heaney  et  al., 
unpubl.  data). 

In  1994  and  1995,  three  adult  females  were 
trapped;  one  was  pregnant  with  a  single  embryo 
(CRL  =  10  mm).  A  multiparous,  nonpregnant 
female  weighed  13  g;  an  adult  male  weighed  7  g. 
Both  cranial  and  external  measurements 
(Table  2)  are  within  stated  ranges  for  Mindanao 
(Heaney  &  Ruedi,  1994)  but  are  slightly  smaller 
than  those  of  series  taken  on  Biliran,  Leyte,  and 
Maripipi  (Rickart  et  al.,  1993). 

Specimens  Examined — Total  5.  Site  4  (2  fmnh, 
1  msu-iit);  Site  5  (1  fmnh);  Site  7  (1  fmnh). 

Suncus  murium  (Linnaeus,  1 766) 

The  Asian  house-shrew  occurs  widely  in  Asia 
and  Indo-Australia;  it  now  occurs  throughout 
the  Philippines,  though  it  is  not  native  to  the 
country.  It  is  abundant  in  urban  and  agricultural 
areas;  on  islands  with  low  mammal  species 
richness  such  as  Negros,  it  is  sometimes  abun- 
dant in  disturbed  forest  and  occasionally  in 
primary  forest  (Heideman  et  al.,  1987;  Heaney  et 
al.,  1989,  1991),  but  on  islands  of  average  species 
richness,  it  is  usually  rare  or  absent  from  forest 
(Heaney  et  al.,  1989,  1998.  unpubl.  data;  Rickart 
et  al.,  1993). 

A  single  subadult  specimen  from  Mt.  Tim- 
poong  Peak  was  available  previously  (Heaney 
1984).  In  1994-1995,  we  captured  this  species 
from  150  to  1475  m,  and  it  was  the  most 
common  species  at  the  three  highest  sites,  all  in 
primary  forest  (Table  1,  Fig.  1).  It  was  especially 
abundant  in  montane  forest  at  1275  m  (Site  7).  It 
was  moderately  abundant  in  primary  mossy 
forest  (Site  6;  elev.  1300  m)  and  in  lower  mossy 
forest  at  1475  m  elevation  (Site  8)  but  was  much 
less  common  in  heavily  disturbed  lowland 
agricultural  land  at  150  m  (Site  3).  This  pattern 
of  abundance  is  quite  different  from  that  on  the 
species-rich  islands  of  Biliran,  Leyte,  Luzon, 
Maripipi,  and  Mindanao,  where  specimens  were 
never  caught  in  primary  forest  (Heaney  et  al., 
1989,  1999,  unpubl.  data;  Rickart  et  al.,  1993) 
but  similar  to  the  species-poor  island  of  Negros, 
where  S.  murinus  was  abundant  in  transitional 
mossy/montane  forest  and  in  mossy  forest  at 


30 


FIELDIANA:  ZOOLOGY 


1800  i- 
1600  =" 
1400  — 


~  1200 

O   1000   — 

re  _ 

|     800 


600  — 
400  — 
200 

0 


*=1960s 
■=1990s 


11 


•a  5) 


o 


maximum  elevation 


J 1 1 1. ■ 1 1 

L  " 


o  .c 
c  £ 


t  c 

O    03 
Q..C 

.5) 


OQ 


W    CO 

II 

5  c 
o  cc 

II 


0) 


</>   c/> 
5  ^ 


mossy 


montane 


lowland 


primary 
disturbed 


no  forest 


II 

c 

"3 


Fig.  1.  Elevational  range  of  nonvolant  small  mammals  (Insectivora  and  Rodentia)  on  Camiguin  Island, 
Philippines.  Records  from  the  1960s  are  indicated  with  stars  and  from  the  1990s  by  solid  squares.  The  approximate 
original  boundaries  of  primary  lowland,  montane,  and  mossy  rain  forest  along  the  elevational  gradient  are 
indicated.  The  condition  of  forest  along  our  transect  in  the  middle  1990s  is  indicated  as  nearly  absent  (below 
600  m),  disturbed  by  logging  and  agriculture  but  present  as  second  growth  (about  600-1250  m),  and  primary  or 
lightly  disturbed  by  human  activities  and  landslides  (above  about  1250  m).  Elevations  from  the  1960s  were  rough 
estimates  (see  text). 


1280  m  (Heaney  et  al.,  1989),  which  is  similar  to 
Sites  6  and  7. 

Suncus  murinus  was  most  often  trapped  in 
runways  or  clear  areas  beneath  fallen  and  rotting 
logs,  under  roots  of  trees,  or  under  horizontal 
trunks  of  live  trees  as  well  as  in  runways  near 
large  boulders.  Many  were  caught  during  day- 
light hours. 

Five  adult  females  with  a  mean  weight  of  32  ± 
4.5  g  (range  =  27-39  g)  were  pregnant;  litter 
sizes  for  four  of  these  were  one,  two,  three,  and 
three.  Thirteen  nonpregnant  parous  females 
(those  with  large  mammae)  weighed  an  average 
of  28.8  ±  4.1  g  (range  =  22-35  g),  and  nullipa- 
rous  females  (those  with  small  mammae) 
weighed  20.6  ±  2.2  g  (range  =  17.5-23  g,  N  = 
10).  Adult  males  (defined  as  those  with  large 
testes)  had  a  mean  weight  of  36  ±  5.8  g  (range 
24-48  g,  N  =  27).  Males  are  conspicuously 
larger  than  females  in  this  species  (Table  2). 


Specimens  Examined — Total  78.  Site  3  (2  msu- 
iit);  Site  6(10  msu-iit);  Site  7  (56  fmnh);  Site  8  (9 
fmnh);  Site  15  (1  dmnh). 

Order  Chiroptera 

Family  Pteropodidae — Fruit  Bats 

Cynopterus  brachyotis  (Muller,  1838) 

The  common  short-nosed  fruit  bat  is  wide- 
spread in  Southeast  Asia  and  is  common 
throughout  the  Philippines.  It  ranges  from  sea 
level  to  at  least  1250  m  and  is  typically  found  in 
agricultural  areas;  it  is  also  common  in  second- 
ary lowland  forest  but  usually  rare  in  primary 
forest  (Heaney  et  al.,  1998). 

Our  limited  netting  on  Camiguin  during  1994- 
1995  (Table  3)  indicated  that  C.  brachyotis  was 
abundant  in  a  highly  disturbed  lowland  agricul- 
tural area  at  10  m  elevation  (Site  1)  and  was 
common  in  a  heavily  disturbed  lowland  agricul- 
tural area  at  100  m  (Site  2;  Table  3).  It  was  less 


HEANEY  ET  AL.:  THE  MAMMALS  OF  CAMIGUIN  ISLAND 


31 


Tabu  1.  Numbers  of  nonvolant  small  mammals  captured  in  traps  in  heavily  disturbed  lowland  agricultural 
area  (Site  3)  secondary  lowland  forest  (Site  4).  disturbed  lower  montane  forest  (Site  5),  primary  mossy  forest  (Site 
6)  primary  montane  forest  (Site  7),  and  lower  mossy  forest  (Site  8)  on  Camiguin  Island.  The  numbers  of  captures 
per  KM)  trap-nights  are  given  in  parentheses.  See  Heaney  and  Tabaranza  (2005a)  for  full  site  descriptions.  Asterisks 
mark  species  presumed  to  be  present;  see  Methods. 


Site  3, 

Site  4, 

Site  5, 

Site  6, 

Site  7, 

Site  8, 

Scientific  name 

150  m 

1000  m 

1200  m 

1300  m 

1275  m 

1475  m 

Total 

Crocidura  bcutits 

0 

3  (0.3) 

1  (0.3) 

0* 

1  (0.2) 

0 

5 

Sanctis  marinas 

2(1.4) 

0* 

0* 

10(2.9) 

56(8.5) 

9  (2.3) 

77 

Apomys  camiguinensis 

0 

14(1.5) 

0* 

2  (0.6) 

9(1.4) 

0 

25 

Bullimus  gatnay 

0 

4  (0.4) 

2  (0.6) 

3  (0.9) 

10(1.5) 

1  (0.3) 

20 

C  'runomys  melanius 

0 

2  (0.2) 

1  (0.3) 

0* 

2  (0.3) 

0 

5 

Rattas  everetti 

0 

7(0.8) 

2  (0.6) 

3  (0.9) 

2  (0.3) 

2  (0.5) 

16 

Rat t us  exulans 

2(1.4) 

0* 

0* 

0* 

5  (0.8) 

0 

7 

Rattas  tanczumi 

21  (14.2) 

8  (0.9) 

4(1.2) 

1  (0.3) 

0 

0 

34 

Total  small  mammals 

24 

38 

10 

19 

85 

12 

188 

Total  trap-nights 

148 

907 

339 

348 

655 

386 

2783 

Number/ 100  trap-nights 

16.2 

4.2 

2.9 

5.5 

9.9 

3.1 

6.8 

Total  small  mammal  species 

3 

6(+l) 

5  (+3) 

5  (+3) 

7 

3 

8 

Native  small  mammal  species 

0 

5 

4(+l) 

3  (+2) 

5 

2 

5 

common  in  disturbed  lowland  forest  at  1000  m 
(Site  4)  and  uncommon  in  disturbed  lower 
montane  forest  at  1000-1300  m  and  mossy  forest 
between  1200  and  1400  m  elevation  (Sites  5  and 
6).  Limited  netting  did  not  detect  this  species  in 
montane  primary  forest  at  1275  m  elevation  (Site 
7).  The  occurrence  of  C.  brachyotis  in  these 
habitats  is  consistent  with  records  from  other 
Philippine  islands;  for  example,  on  Catanduanes, 
Leyte,  Luzon,  and  Negros,  this  species  was 
most  abundant  in  agricultural  land  and  second- 
ary forest  (Heaney  et  al.,  1989,  1991,  1999; 
Heideman  &  Heaney,  1989;  Ingle,  1992;  Rickart 
et  al.,  1993),  similar  to  Sites  1  and  3  to  5  on 
Camiguin.  Records  from  the  1960s  indicate  that 
the  species  occurs  along  the  entire  elevational 
gradient,  from  sea  level  to  near  the  peaks, 
although  most  specimens  are  from  below 
1000  m  (i.e.,  below  the  transition  to  montane 
forest;  Fig.  2). 

Eight  adult  females  (mean  =  29.5  ±  4.72  g) 
taken  in  May  1992  and  1994  were  pregnant  with 
single  embryos  (CRL  =  5-26  mm).  Two  of  the 
pregnant  females  (22  and  35  g)  and  three  non- 
pregnant ones  (32.5-39  g)  were  lactating.  Adult 
females  with  enlarged  mammae  but  not  pregnant 
or  lactating  had  mean  weight  of  29.2  ±  2.3  g 
(range  =  26  33.2  g,  N  =  13).  Five  nulliparous 
females  had  a  mean  weight  of  16.5  g  ±  2.0  g 
(range  =  14-19  g).  Males  with  abdominal  testes 
had  a  mean  weight  of  27.4  ±  2.6  g  (range  =  22- 
32.5  g,  N  =  15).  On  Negros  Island,  this  species 
probably  produces  two  young  per  year,  one  in 


March/April  and  another  in  August/September 
(Heideman,  1995). 

Females  are  slightly  larger  than  males  in  most 
external  and  cranial  dimensions  (Table  4),  a  trend 
quite  similar  to  specimens  from  Mt.  Kitanglad, 
Mindanao  (Heaney  et  al.,  unpubl.  data).  Speci- 
mens from  Biliran,  Leyte  and  Maripipi,  in 
contrast,  showed  the  opposite  trend  (Rickart  et 
al.,  1993). 

Specimens  Examined — Total  91.  Site  1  (27 
fmnh);  Site  3  (5  msu-iit);  Site  4  (4  fmnh,  14  msu- 
iit);  Site  6  (1  fmnh,  3  msu-iit);  Site  11(12  dmnh); 
Site  12  (3  dmnh);  Site  13  (6  dmnh);  Site  16  (2 
rom);  Site  17  (14  rom). 

Harpyionycteris  whiteheadi  Thomas,  1 896 

The  harpy  fruit  bat  has  been  reported  pre- 
viously from  Camiguin  (Peterson  &  Fenton, 
1970),  as  well  as  from  Marinduque,  Masbate, 
Mindoro,  Negros,  southern  Luzon,  and  through- 
out Greater  Mindanao  (Heaney  et  al.,  1998, 
1999).  It  is  generally  restricted  to  primary  or 
lightly  disturbed  forest;  it  is  usually  rare  in 
lowland  forest  but  often  is  moderately  common 
in  montane  forest  from  roughly  800  m  to  at 
least  1800  m.  It  apparently  feeds  heavily  on 
the  fruits  of  viney  pandans  (Freycinetia  spp.)  and 
figs  (Ficus  spp.);  (Heaney  et  al.,  1989,  1999; 
Heideman  &  Heaney,  1989;  Rickart  et  al., 
1993). 

In  May  1992,  we  netted  one  pregnant  female 
weighing  123  g  with  a  single  embryo  (CRL  = 
28  mm)    at     10  m    elevation,     in    a    lowland 


32 


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anao (Kitanglad  Range),  pregnant  females 
were  similarly  recorded  in  May,  with  further 
records  of  pregnancy  in  April,  August,  and 
October;  lactation  was  noted  in  March,  April, 
and  August  (Heaney  et  al.,  unpubl.  data).  In 
March  1995,  at  Site  7  (1275  m  in  primary 
montane  forest),  where  Freycinetia  spp.  were 
abundant,  we  often  heard  the  distinctive  Har- 
pyionycteris  whistles  (Rickart  et  al.,  1993;  Hea- 
ney et  al.,  1999),  but  none  were  captured. 
Combined  with  records  from  the  1960s  (Fig.  2), 
these  data  indicate  that  it  occurs  on  this  island 
from  sea  level  to  1500  m  in  lowland,  montane, 
and  mossy  forest. 

External  and  cranial  measurements  of  H. 
whiteheads  on  Camiguin  are  comparable  to  those 
from  Mindanao  (Mt.  Kitanglad)  but  are  slightly 
larger  than  those  found  on  Leyte  and  Luzon 
(Mt.  Isarog)  (Heaney,  1984;  Rickart  et  al.,  1993; 
Heaney  et  al.,  1999,  unpubl.  data). 

Specimens  Examined — Total  4.  Site  1  (1 
fmnh);  Site  11  (1  dmnh);  Site  12  (1  dmnh);  Site 
16  (1  ROM). 

Macroglossus  minimus  (E.  Geoffroy,  1810) 

The  dagger-toothed  flower  bat  occurs  from 
Thailand  to  Australia  and  is  found  throughout 
the  Philippines  (Heaney  et  al.,  1998).  Within  the 
Philippines,  it  occurs  in  virtually  every  habitat  in 
the  country,  from  sea  level  to  at  least  2250  m.  It 
is  often  abundant  in  agricultural  and  heavily 
disturbed  areas,  is  common  in  secondary  forest, 
and  usually  is  uncommon  in  primary  forest.  It  is 
most  often  associated  with  domestic  or  wild 
banana  (Musa  spp.;  Heaney  et  al.,  1989,  1999; 
Heideman  &  Heaney,  1989;  Rickart  et  al., 
1993). 

On  Camiguin,  our  limited  netting  showed  M. 
minimus  to  be  abundant  in  a  highly  disturbed 
lowland  agricultural  area  at  ca.  10  m  elevation 
(Site  1),  present  in  a  heavily  disturbed  lowland 
agricultural  area  at  100  m  (Site  3),  uncommon  in 
disturbed  lowland  forest  at  1000  m  (Site  4), 
common  in  disturbed  lower  montane  forest  at 
1000-1300  m  (Site  5),  and  present  in  mossy 
forest  between  1200  and  1400  m  elevation  (Site 
6;  Table  3).  Limited  netting  did  not  detect  this 
species  in  primary  montane  forest  at  1275  m 
elevation  (Site  7).  Combined  with  prior  records, 
it  is  apparent  that  this  species  occurs  on 
Camiguin  from  sea  level  to  at  least  1200  m,  in 
lowland  and  montane  forest,  in  both  disturbed 
and  undisturbed  forest  (Fig.  2). 


HEANEY  ET  AL.:  THE  MAMMALS  OF  CAMIGUIN  ISLAND 


33 


Tabu:  3.  Numbers  of  fruit  bats  captured  in  mist  nets  in  a  lowland  agricultural  area  (Site  1),  heavily  disturbed 
lowland  agricultural  area  (Site  3),  secondary  lowland  forest  (Site  4),  disturbed  lower  montane  forest  and  primary 
mossy  forest  (Sites  5  and  6  concurrently),  and  primary  montane  forest  (Site  7)  on  Camiguin  Island  during  1992, 
1994,  and  1995.  The  number  of  captures  per  net-night  are  given  in  parentheses.  See  Heaney  and  Tabaranza  (2006a) 
for  full  site  descriptions.  Asterisks  indicated  species  observed  but  not  netted  (see  text). 


Scientific  name 


Site  I, 
10  m 


Site  3, 
150  m 


Site  4, 
1000  in 


Sites  5  and  6, 
1200-1400  m 


Site  7, 
1275  m 


( 'ynopterus  brach  vol  is 

27  (4.5) 

5(1.67) 

18(0.75) 

4  (0.29) 

0 

liar  prion  vcteris  whitehead! 

1  (0.17) 

0 

0 

0 

0* 

Macroglossus  minimus 

15  (2.5) 

1  (0.33) 

4(0.17) 

9  (0.64) 

0 

Picnot  hints  jagori 

8(1.33) 

0 

16(0.67) 

1  (0.07) 

0 

Total  fruit  bats 

51 

5 

35 

14 

0 

Total  net-nights 

6 

3 

24 

14 

8 

Fruit  bats  per  net-night 

8.5 

1.67 

1.46 

1.00 

0 

Six  adult  females  netted  in  May  1992  and 
1994.  with  a  mean  weight  of  15.2  ±  3.2  g  (range 
=  12.5-20.3  g),  were  pregnant  with  single 
embryos.  Four  parous  females  with  large  mam- 
mae but  neither  pregnant  nor  lactating  had 
a  mean  weight  of  18.5  ±  1.78  g  (range  =  16- 
20  g).  Three  adult  males  weighed  between  16  and 
19  g.  A  juvenile  male  and  a  juvenile  female  each 
weighed  5  g.  On  Mindanao  (Kitanglad  Range), 
pregnant    females   were    recorded    in    April    to 


August  and  October,  while  lactating  females 
were  recorded  in  May,  September,  and  October 
(Heaney  et  al.,  unpubl.  data).  Heideman  (1995) 
documented  that  this  species  undergoes  aseaso- 
nal  breeding  and  postpartum  estrus  on  Negros 
Island  and  has  several  young  per  year. 

Comparison  with  specimens  of  M.  minimus 
from  Biliran,  Dinagat,  Leyte,  Luzon  (Mt. 
Isarog),  Maripipi,  and  Mindanao  (Kitanglad 
Range)  shows  that  while  the  overall  variation 


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HEANEY  ET  AL.:  THE  MAMMALS  OF  CAMIGUIN  ISLAND 


35 


in  cranial  and  external  measurements  is  slight, 
Camiguin  specimens  tend  to  cluster  consistently 
within  the  upper  ranges  of  most  features 
measured  (Table  4;  Heaney  and  Rabor,  1982; 
Heaney  et  al.,  1991,  1999,  unpubl.  data;  Rickart 
et  al.,  1993). 

Specimens  Examined — Total  56.  Site  1  (15 
fmnh);  Site  3  ( 1  msu-iit);  Site  4  (4  msu-iit);  Site  5 
(8  msu-iit);  Site  6  (1  msu-iit);  Site  11(12  dmnh); 
Site  13  (5  dmnii);  Site  14  (2  dmnii);  Site  17  (8 
dmnii). 

Ptenochirus  jagori  ( Peters,  1861) 

The  musky  fruit  bat  is  a  common  Philippine 
endemic,  occurring  throughout  the  archipelago 
with  the  exception  of  the  Batanes/Babuyan  and 
Palawan  faunal  regions  from  sea  level  to  at  least 
1800  m  (Heaney  et  al.,  1998).  Our  limited  netting 
on  Camiguin  showed  Ptenochirus  jagori  to  be 
common  in  a  lowland  agricultural  area  at  10  m 
elevation  (Site  1),  less  common  in  disturbed 
lowland  forest  at  1000  m  (Site  4),  and  scarce  in 
primary  mossy  forest  at  1300  m  (Site  6;  Table  3). 
Combined  with  records  from  the  1960s,  these 
data  indicate  that  the  species  is  widespread  in 
lowland  and  montane  rain  forest  from  sea  level 
to  at  least  1200  m  (Fig.  2),  though  probably  its 
abundance  declines  with  increasing  elevation  and 
with  increasing  levels  of  disturbance  (Table  3),  as 
noted  elsewhere  (Heaney  et  al.,  1989,  1999, 
unpubl.  data;  Heideman  &  Heaney,  1989;  Rick- 
art et  al.,  1993;  Lepiten,  1997). 

Three  adult  females,  netted  in  May  1992  and 
1994,  weighing  an  average  of  70  g  (range  =  68- 
75  g),  were  pregnant  with  a  single  embryo  each 
(CRL  =  5  10  mm).  Three  nonpregnant  females 
with  large  mammae  had  an  average  weight  of 
73.5  g  (range  =  72-74  g),  and  two  nulliparous 
females  weighed  67  and  68.5  g.  Eleven  adult 
males  had  a  mean  weight  of  73.2  ±  4.7  g  (range 
=  64-78  g,  N  =  11).  Pregnant  females  of  P. 
jagori  have  been  recorded  also  in  May  on  Luzon 
(Mt.  Isarog)  (Heaney  et  al.,  1999).  On  Mindanao 
(Kitanglad  Range),  pregnant  females  were  re- 
corded in  March,  May,  July,  and  August  and 
lactating  females  in  May  to  June  and  August  to 
December  (Heaney  et  al..  unpubl.  data).  Heide- 
man and  Powell  (1998)  found  that  on  Negros 
Island,  P.  jagori  gives  birth  to  a  single  young 
twice  each  year:  the  first  in  late  March  or  early 
April  and  the  second  in  August.  It  was  further 
discovered  that  this  species  undergoes  delayed 
implantation  and  early  development  that  lasts 
for   five    months,    shorter   than    in    two   other 


endemic  species  of  Philippine  pygmy  fruit  bats, 
Haplonycteris  fischeri  and  Otopteropus  eartilago- 
nodus,  where  the  phenomenon  was  first  detected 
(Heideman,  1989;  Heideman  et  al.,  1993;  Heide- 
man &  Powell,  1998).  Additionally,  this  condi- 
tion was  apparently  exhibited  by  primiparous 
young  adult  females  only,  allowing  them  to  give 
birth  only  once  in  their  first  year,  which  had  the 
effect  of  enabling  them  to  synchronize  breeding 
with  the  adult  females  the  following  year  (Heide- 
man &  Powell,  1998). 

Males  are  somewhat  larger  than  females  in 
most  cranial  and  external  dimensions,  as  on 
Biliran,  Leyte,  and  Maripipi  (Table  4;  Rickart  et 
al.,  1993).  Cranial  and  external  measurements 
(Table  4)  are  noticeably  larger  than  those  for 
series  from  Catanduanes  and  southern  Luzon 
and  were  similar  to  those  from  Biliran,  Leyte, 
and  Maripipi  (Heaney,  1984;  Heaney  et  al.,  1991, 
1999;  Rickart  et  al.,  1993). 

Specimens  Examined — Total  46.  Site  1  (8 
fmnh);  Site  4(16  msu-iit);  Site  6  (1  msu-iit);  Site 
11  (6  dmnh);  Site  13  (6  dmnh);  Site  17  (9  rom). 

Pteropus  hypomelanus  Mearns,  1905 

The  common  island  flying  fox  occurs  from 
Thailand  to  Australia  and  is  found  throughout 
the  Philippines  with  the  exception  of  the  Palawan 
faunal  region.  It  is  often  common  in  agricultural 
areas  from  sea  level  to  ca.  900  m  and  is  absent  in 
primary  forest  (Heideman  &  Heaney,  1989; 
Heaney  et  al.,  1991,  1998;  Utzurrum,  1992; 
Rickart  et  al.,  1993).  Records  from  the  1960s 
document  it  from  Camiguin  at  elevations  from 
about  250  to  1500  m  (Fig.  2),  but  we  netted  none 
in  the  1990s;  because  this  species  typically  flies 
above  the  canopy  and  our  nets  were  set  not  more 
than  about  4  m  above  the  ground,  our  failure  to 
catch  any  does  not  necessarily  indicate  any 
change  in  their  abundance. 

External  and  cranial  measurements  show  only 
slight  variations  with  those  of  specimens  from 
Dinagat,  Panay,  Leyte,  and  Maripipi  (Heaney  & 
Rabor,  1982;  Rickart  et  al.,  1993). 

Specimens  Examined — Total  8.  Site  10  (2 
dmnh);  Site  12  (1  dmnh);  Site  13  (2  dmnh);  Site 

17  (3  ROM). 

Pteropus  pwnilus  Miller,  1910 

The  little  golden-mantled  flying  fox  is  endemic 
to  the  Philippines  (aside  from  a  single  population 
on  Miangas  Island.  Indonesia,  adjacent  to 
Mindanao)  and  occurs  throughout  the  archipel- 
ago, with  the  exception  of  the  Batanes/Babuyan 


36 


FIELDIANA:  ZOOLOGY 


and  Palawan  regions  (Heaney  et  al.,  1998). 
Previously  reported  from  Camiguin  as  P.  tablasi, 
the  species  was  revised  to  include  P.  tablasi  and 
P.  balutus  as  synonyms  under  Pteropus  pumilus 
(Klingener  &  Creighton,  1984).  It  is  associated 
with  primary  and  good  secondary  lowland  forest 
from  sea  level  to  about  1100  m,  and  it  is 
uncommon  outside  of  forest.  Additionally,  it  is 
most  common  on  small  islands  and  is  uncommon 
to  rare  on  larger  islands.  Pteropus  pumilus  often 
is  netted  in  clearings  or  on  ridgetops  (Heideman 
&  Heaney,  1989;  Utzurrum,  1992;  Rickart  et  al., 
1993). 

We  did  not  encounter  P.  pumilus  during  the 
1990s,  but  the  1967-1969  surveys  obtained  42 
individuals  at  four  sites  from  ca.  250  to  nearly 
1000  m  elevation  (Fig.  2).  As  with  Pteropus 
hypomelanus,  this  species  usually  flies  above  the 
canopy,  so  our  failure  to  catch  any  in  the  1990s 
does  not  necessarily  indicate  a  change  in  their 
status  on  the  island. 

Specimens  Examined — Total  42.  Site  11  (19 
dmnh);  Site  13  (2  dmnh);  Site  14  (1  dmnh);  Site 

17  (20  ROM). 

Family  Emballonuridae — Sheath-tailed  Bats 

Emballanura  alecto  (Eydoux  and  Gervais,  1836) 
The  Philippine  sheath-tailed  bat  is  a  common 
cave-dwelling  species  that  occurs  throughout 
most  of  the  Philippines  and  is  also  known  from 
Borneo,  the  Moluccas,  and  Sulawesi  (Koopman, 
1989).  It  has  been  recorded  only  in  lowland  areas 
(450  m  and  below)  in  disturbed  forest  and 
agricultural  areas  with  scattered  remnant  forest, 
with  most  captures  recorded  from  caves,  under 
large  boulders,  or  in  man-made  tunnels  (Heaney 
et  al.,  1991,  1999;  Rickart  et  al.,  1993).  We  did 
not  record  this  species  during  our  fieldwork  in 
1994  and  1995,  but  in  1967  specimens  were  taken 
from  Tag-ibo  Cave  at  400  ft  (ca.  120  m)  and  at 
1400-3300  ft  (ca.  400-1000  m)  on  Mt.  Mamba- 
jao  (see  also  Heaney,  1984). 

Comparison  of  external  and  cranial  measure- 
ments with  series  from  Leyte  and  Biliran  shows 
little  variation  (Table  4;  Rickart  et  al.,  1993). 

Specimens  Examined — Total  4.  Site  17  (2 
rom);  Site  18  (2  rom). 

Family  Rhinolophidae — Horseshoe-nosed  Bats 

Rhinolophus  arcuatus  Peters,  1871 

The  arcuate  horseshoe  bat  is  widespread  from 
Sumatra  to  New  Guinea  and  throughout  the 
Philippines  (Heaney  et  al.,  1998),  including  the 
Palawan  faunal  region  (Esselstyn  et  al.,  2004).  It 


is  most  often  encountered  from  lowlands  to  at 
least  1350  m  in  agricultural  lands  to  primary 
lowland  and  montane  forest  and  occasionally 
roosts  in  caves  (Heaney  et  al.,  1991,  1999; 
Rickart  et  al.,  1993).  Two  groups  that  differ  in 
size  and  habitat  are  recognized  within  this 
species:  a  smaller  morphotype,  designated  R. 
arcuatus — s,  that  occurs  in  the  lowlands  or 
disturbed  habitats,  and  a  larger  one,  designated 
R.  arcuatus — 1,  found  in  forest  at  higher  eleva- 
tions (Ingle  &  Heaney,  1992). 

In  May  1994,  we  netted  this  species  at  1000  m 
elevation  in  disturbed  lowland  forest  (Site  4).  Of 
two  adult  females  netted,  one  (15  g)  was 
pregnant  with  one  large  embryo  (CRL  = 
28  mm),  and  the  other  was  lactating  (12.5  g). 
Pregnancies  in  this  species  were  recorded  also  in 
March  on  Luzon  (Mt.  Isarog)  and  in  April  on 
Biliran,  Leyte,  and  Maripipi  (Rickart  et  al.,  1993; 
Heaney  et  al.,  1999). 

Cranial  and  external  measurements  (Table  5) 
of  the  Camiguin  specimens  are  consistently 
larger  than  those  in  series  from  Biliran,  Leyte, 
Luzon  (Mt.  Makiling),  Maripipi,  and  Mindanao 
(Mt.  Kitanglad),  which  all  fall  within  the 
dimensions  of  the  smaller  morphotype,  R. 
arcuatus — s  (Ingle  &  Heaney,  1992;  Rickart  et 
al.,  1993;  Heaney  et  al.,  1999,  unpubl.  data). 
Instead,  their  external  and  cranial  dimensions  are 
comparable  to  the  larger  morphotype,  R.  arcua- 
tus— 1  (Ingle  &  Heaney,  1992).  Systematics  in  this 
"species"  are  badly  in  need  of  detailed  study. 

Specimens  Examined — Total  10.  Site  4  (4  msu- 
iit);  Site  13  (6  dmnh). 

Rhinolophus  inops  K.  Anderson,  1905 

The  Philippine  forest  horseshoe  bat  is  common 
to  abundant  in  primary  lowland  and  montane 
forest  from  sea  level  to  2250  m  and  is  usually 
rare  in  secondary  forest  and  mossy  forest 
(Heaney  et  al.,  1991,  1998,  1999;  Rickart  et  al., 
1993).  Improvements  in  our  understanding  of 
Philippine  Rhinolophus  lead  us  to  reidentify  the 
single  specimen  from  Site  13,  reported  as  R. 
subrufus  by  Heaney  (1984),  as  R.  inops.  We 
netted  two  additional  males  in  disturbed  lowland 
forest  at  1000  m  (Site  4).  Cranial  and  external 
measurements  (Table  5)  are  slightly  larger  in 
most  dimensions  than  those  of  the  series  from 
Biliran,  Leyte  (Rickart  et  al.,  1993),  and  Mind- 
anao (Mt.  Kitanglad;  Heaney  et  al.,  unpubl. 
data);  we  suspect  that  regional  morphs  corre- 
sponding to  the  Pleistocene  islands  (Heaney, 
1986)  are  present. 


HEANEY  ET  AL.:  THE  MAMMALS  OF  CAMIGUIN  ISLAND 


37 


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Specimens  Examined — Total  3.  Site  4  (2  msu- 
lii);  Site  13  (1  dmnh). 

Rhinolophus  virgo  K.  Anderson,  1905 

The  yellow-faced  horseshoe  bat  is  an  endemic 
species  widespread  within  the  Philippines.  It  is 
usually  encountered  in  primary  lowland 
forest  from  250  to  1100  m;  several  records  are 
from  caves  (Heaney  et  al,  1991;  Rickart  et  al., 
1993).  We  obtained  one  adult  female  from 
the  edge  of  a  small  tree-fall  gap  in  primary 
montane  forest  at  1275  m  on  22  March  1995 
(Site  7).  It  has  small  mammae  and  showed  no 
indication  of  reproductive  activity.  Cranial 
dimensions  (Table  5)  fall  within  the  range  of 
those  from  Leyte  and  Maripipi  but  are  slightly 
smaller  than  those  from  Mindanao  (Mt.  Kitan- 
glad;  Rickart  et  al,  1993;  Heaney  et  al.,  unpubl. 
data). 

Specimens  Examined — Total  1.  Site  7  (1 
fmnh). 

Family  Vespertilionidae — Common  Bats 

Murina  cyclotis  Dobson,  1872 

The  round-eared  tube-nosed  bat  is  widespread 
in  southern  Asia  and  was  previously  known  in 
the  Philippines  from  a  few  specimens  from  the 
central  and  southern  portion  of  Greater  Luzon 
as  well  as  Greater  Mindanao,  Sibuyan,  and 
Siquijor,  with  records  from  disturbed  and 
primary  lowland  and  montane  forest  from  250 
to  1500  m  (Heaney  et  al.,  1991,  1998,  1999, 
unpubl.  data;  Rickart  et  al.,  1993;  Ruedas,  1994; 
Lepiten,  1997).  We  netted  one  adult  female  in 
May  1994  in  secondary  lowland  forest  at  1000  m 
(Site  4). 

Specimens  Examined — Total  1.  Site  4  (1  msu- 
iit). 

Pipistrellus  javanicus  (Gray,  1838) 

The  Javan  pipistrelle  is  found  from  Korea 
to  Java  and  throughout  the  Philippines.  It  is 
generally  common  in  primary  montane  forest 
and  uncommon  in  primary  lowland  and  mossy 
forest,  though  it  has  been  found  from  sea 
level  to  2250  m  (Ingle,  1992;  Heaney  et  al., 
1998,  1999,  unpubl.  data).  A  single  skin  without 
a  skull,  tentatively  identified  as  this  species,  was 
obtained  from  Camiguin  by  the  1960s  surveys.  In 
March  1995,  two  were  netted  in  primary 
montane  forest  at  1275  m  (Site  7).  Cranial  and 
external  measurements  (Table  5)  are  comparable 
with  those  of  a  series  from  Mindanao  (Mt. 
Kitanglad)  in  most  dimensions  (Heaney  et  al., 
unpubl.  data). 


38 


FIELDIANA:  ZOOLOGY 


Specimens  Examined — Total  3.  Site  7  (2 
fmnh);  Site  13  (1  dmnh). 

Order  Primates 

Family  Cercopithecidae — Monkeys 

Macaca  fascicularis  (Raffles,  1821) 

The  long-tailed  macaque  occurs  from  Burma 
to  Timor  and  is  found  throughout  the  Philip- 
pines (Fooden,  1995;  Heaney  et  al.,  1998).  It  is 
found  in  agricultural  areas  near  forest,  in  second- 
growth  and  secondary  forest,  as  well  as  primary 
forest  from  sea  level  to  at  least  1800  m  in 
lowland  and  montane  forest  (Goodman  &  Ingle, 
1993;  Rickart  et  al.,  1993;  Heaney  et  al.,  1999). 
We  did  not  collect  any  specimens,  but  in  March 
1995  we  obtained  the  skull  of  an  adult  individual 
from  a  hunter  who  had  killed  the  macaque  in 
a  forested  area  above  Mahinog  a  few  years 
earlier.  Also  in  March  1995,  a  local  hunter  who 
was  a  member  of  our  team  saw  two  macaques 
a  short  distance  from  our  camp  at  1100m 
elevation  (Site  7)  in  primary  montane  forest. 
He  reported  that  the  macaques  were  fairly 
common  in  what  remained  of  lowland  primary 
forest  and  good  secondary  forest  at  that  time  and 
were  actively  hunted. 

Specimens  Examined — Total  1.  Above  Mahi- 
nog (1  fmnh). 

Order  Rodentia 

Family  Muridae — Rats  and  Mice 

Apomys  camiguinensis  Heaney  and  Tabaranza, 
2006 

A  previously  unknown  species  of  Apomys 
(Musser,  1982)  was  trapped  on  Camiguin  in 
May  1994  and  March  1995  (Heaney  &  Tabar- 
anza, 1997)  and  described  in  this  volume 
(Heaney  &  Tabaranza,  2006b).  The  Camiguin 
forest  mouse  is  moderately  large  for  the  genus, 
averaging  about  40  g.  It  is  marked  by  a  number 
of  subtle  morphological  characters,  and  molec- 
ular data  show  it  to  be  most  closely  related  to 
Apomys  hylocoetes  and  A.  insignis  from  Mind- 
anao and  to  an  unnamed  species  from  Leyte  and 
Biliran  (Steppan  et  al.,  2003;  Heaney  &  Tabar- 
anza, 2005b).  This  species  was  common  (Table  1, 
Fig.  1)  in  disturbed  lowland  forest  at  1000  m 
(Site  4)  and  in  primary  montane  forest  at  1275  m 
(Site  7)  and  relatively  uncommon  in  primary 
mossy  forest  at  1200-1400  m  (Site  6).  It  was 
not  found  in  heavily  disturbed  lowland  agri- 
cultural land  (Site  3,  elev.  150  m),  lower 
montane  forest  (Site  4,  elev.  1000-1300  m),  or 
lower  mossy  forest  (Site  8,  elev.  1475  m),  but 


there  was  limited  sampling  at  all  of  these 
(Table  1).  Most  specimens  were  taken  from  traps 
placed  in  primary  montane  forest  underneath 
root  tangles  and  beneath  fallen  and  rotten  logs 
on  a  moderate  slope.  All  individuals  were 
captured  at  night. 

A  pregnant  female  taken  on  29  May  1994 
weighed  40.5  g  and  had  a  single  embryo  (CRL  = 
30  mm).  Adult  females  with  large  mammae  but 
not  pregnant  weighed  39  ±  2.1  g  (range  =  37.5- 
42  g,  N  =  5);  two  of  these,  taken  23  May  1994 
and  17  March  1995,  each  had  two  placental 
scars.  Nulliparous  females  (with  small  mammae) 
were  lighter  and  had  an  average  weight  of  35  g 
(range  =  34-36).  Females  have  two  pair  of 
inguinal  mammae.  These  data  indicate  a  litter 
size  between  one  and  two.  Males  with  scrotal 
testes  had  a  mean  weight  of  41.3  ±  4.7  g  (range 
=  36.5^48  g,  N  =  11);  testis  size  ranged  from  15 
X  16  mm  to  8  X  12  mm.  Males  with  abdominal 
testes  weighed  33.0  ±  5.0  g  (range  =  28-34  g,  N 
=  5)  and  had  testis  size  of  5  X  8  mm. 

Specimens  Examined — Total  25.  Site  4  (14 
msu-iit),  Site  6  (2  msu-iit);  Site  7  (9  fmnh). 

Bullimus  gamay  Rickart,  Heaney  and  Tabaranza, 
2002 

A  previously  unknown  species  of  Bullimus, 
recently  described  as  B.  gamay  (Rickart  et  al., 
2002),  was  captured  on  Camiguin  in  May  1994 
and  March  1995.  The  Camiguin  forest  rat  was 
incorrectly  assigned  to  Tarsomys  by  Heaney  and 
Tabaranza  (1997)  and  was  designated  "Bullimus 
sp.  A"  by  Heaney  et  al.  (1998).  It  was  trapped 
from  900  to  1475  m  (Fig.  1,  Table  1)  but  not  in 
heavily  disturbed  lowland  agricultural  land  from 
sea  level  to  300  m  (Site  3).  Most  specimens  were 
taken  from  traps  placed  in  runways  beneath  root 
tangles  and  rotten  logs  or  near  large  rocks.  This 
species  was  most  common  in  primary  montane 
forest  at  1275  m  (Site  7).  It  was  less  common  in 
disturbed  lower  montane  forest  at  1000-1300  m 
(Site  5)  and  in  primary  mossy  forest  at  1200- 
1400  m  (Site  6).  It  was  rare  in  secondary  lowland 
forest  at  1000  m  (Site  4)  and  in  lower  mossy 
forest  at  1475  m  (Site  8).  Bullimus  bagobus 
occurs  in  similar  habitat  on  Mindanao  but  also 
occurs  at  lower  elevation  where  good  forest 
persists  (Heaney,  2001;  Heaney  et  al.,  unpubl. 
data);  this  suggest  that  B.  gamay  should  also  be 
sought  at  lower  elevations  on  Camiguin. 

Parous  females  with  large  mammae  weighed 
an  average  of  370  g  (range  =  360-390  g,  N  =  3), 
while  nulliparous  young  adult  females  with  small 


HEANEY  ET  AL.:  THE  MAMMALS  OF  CAMIGUIN  ISLAND 


39 


mammae  were  lighter,  having  a  mean  weight  of 
276  ±  53.7  g  (range  =  210-345  g,  N  =  5).  Adult 
males  with  scrotal  testes  had  an  average  weight 
of  402  g  (range  =  240-500  g;  N  -  3)  and  had 
testis  size  ranging  from  21  x  30  mm  to  27  X 
35  mm).  Among  the  nine  individuals  taken  in 
May  1994,  the  smallest  was  a  nulliparous  young 
female  weighing  210  g,  while  among  the  11 
individuals  captured  in  March  1995,  two  were 
juveniles,  a  male  weighing  120  g  and  a  female 
weighing  125  g.  No  pregnant  females  were  taken 
in  May  1994  and  March  1995,  but  a  lactating 
female  (360  g)  was  taken  on  18  March  1995,  and 
a  female  taken  on  29  May  1994  had  three 
placental  scars.  This  suggests  that  females  give 
birth  in  February  and  March.  Three  females  had 
three  pairs  of  mammae  and  one  had  four  pairs, 
indicating  that  litter  size  most  likely  does  not 
exceed  three. 

Specimens  Examined — Total  20.  Site  4  (4  msu- 
iit);  Site  5  (2  msu-iit);  Site  6  (3  msu-iit);  Site  7  (10 
emnh);  Site  8  (1  fmnh). 

Crunomys  melanius  Thomas,  1907 

The  Mindanao  shrew-mouse  was  previously 
known  only  from  Leyte  and  Mindanao  from 
near  sea  level  to  1550  m,  apparently  in  primary 
lowland  rain  forest;  it  is  rare  in  collections 
(Musser  &  Heaney,  1992;  Heaney  et  al.,  1998; 
unpubl.  data;  Rickart  et  al.,  1998).  On  Camiguin, 
we  found  it  to  be  uncommon  in  heavily  disturbed 
lowland  agricultural  land  at  1000  m  (Site  4),  in 
disturbed  lower  montane  forest  at  1200  m  (Site 
5),  and  in  primary  montane  forest  at  1275  m 
(Site  7;  Table  1,  Fig.  1).  It  was  trapped  beneath 
rotten  logs  and  wood  tangles  in  an  area  with  few 
dead  leaves  and  little  moss.  It  occurs  down  to  sea 
level  in  good  forest  on  Mindanao  (Heaney  et  al., 
1998;  Heaney,  2001)  and  should  also  be  sought 
at  lower  elevations  on  Camiguin. 

Two  of  the  five  specimens  we  captured  were 
females.  An  adult  female  taken  on  17  March 
1995,  with  enlarged  mammae  and  weighing  71  g, 
had  recently  given  birth;  it  had  three  placental 
scars  (one  left,  two  right),  and  the  uterus  was 
swollen.  The  other  female  (62.5  g),  probably 
nulliparous,  had  small  mammae.  Two  adult 
males  (60  g  and  72  g)  had  scrotal  testes  (15  X 
8  mm);  the  third  male,  taken  on  24  May  1994, 
was  a  small  juvenile  (28  g)  with  abdominal  testes. 
These  scanty  data  imply  that  March,  April,  and 
May  are  months  of  reproductive  activity.  Fe- 
males have  two  pairs  of  mammae,  indicating  that 
litter  size  is  small. 


Cranial  and  external  measurements  (Table  6) 
are  similar  to  but  slightly  larger  than  those  from 
Leyte  and  Mindanao  (Kitanglad  Range  and  Mt. 
Apo;  Rickart  et  al.,  1993,  1998;  Heaney  et  al., 
unpubl.  data). 

Specimens  Examined — Total  5.  Site  4  (2  msu- 
iit);  Site  5  (1  msu-iit);  Site  7  (2  fmnh). 

Rattus  everetti  (Gunther,  1879) 

The  common  Philippine  forest  rat  is  a  wide- 
spread endemic  species  (excluding  the  Babuyan/ 
Batanes,  Palawan,  and  Sulu  groups)  found  in 
primary  and  disturbed  lowland,  montane,  and 
mossy  forest  from  sea  level  to  2400  m  (Heaney  et 
al.,  1991,  1998,  1999;  Musser  &  Heaney,  1992; 
Rickart  et  al.,  1993). 

During  1994-1995,  we  found  this  250-350  g 
rodent  to  be  fairly  common  at  all  sampling  sites 
from  disturbed  lowland  forest  at  1000  m, 
through  montane  forest  to  mossy  forest  at 
1475  m;  specimens  from  the  1960s  were  taken 
at  500  m  (Table  1,  Fig.  1);  on  Mindanao,  it 
occurs  down  to  sea  level  (Heaney,  2001;  Heaney 
et  al.,  unpubl.  data).  It  was  trapped  under  root 
tangles  and  beneath  fallen  logs,  in  hollow  spaces 
beneath  live  trees,  and  near  mounds  of  soil  and 
rotting  leaves. 

Five  parous  adult  females  from  May  1994  and 
March  1995  had  a  mean  weight  of  271  ±  28.8 
(range  =  240-300  g,  N  =  5).  During  the  same 
months,  five  nulliparous  females  (with  adult 
pelage)  weighed  188  ±  21.4  g  (range  =  160- 
215  g),  and  four  females  with  juvenile  pelage 
weighed  116  ±  31.0  g  (range  =  77-150  g). 
Females  have  four  pairs  of  mammae,  indicating 
moderate  litter  size.  None  were  pregnant,  and  we 
were  unable  to  count  placental  scars;  previous 
studies  also  produced  little  data  on  litter  size 
(Heaney  et  al.,  1999,  unpubl.  data).  Males  with 
scrotal  testes,  recorded  in  May  1994  and  March 
1995,  had  a  mean  weight  of  288  ±  69.8  g  (range 
=  210  375  g,  N  =  4),  and  the  largest  males  had 
testes  size  of  33  X  52  mm  and  30  X  50  mm. 
Three  males  with  abdominal  testes,  during  the 
same  months,  weighed  an  average  of  207  g 
(range  =  160-295  g). 

Cranial  and  external  measurements  (Table  6) 
are  comparable  to  those  from  Biliran,  Leyte,  and 
Maripipi  but  slightly  smaller  than  series  from 
Dinagat,  Mindanao  (Kitanglad  Range),  and 
Siargao  (Heaney  &  Rabor,  1982;  Heaney  et  al., 
1991,  1999,  unpubl.;  Rickart  et  al.,  1993). 

Specimens  Examined— Total  28.  Site  4  (6  msu- 
iit);  Site  5  (2  msu-iit);  Site  6  (3  msu-iit);  Site  7  (8 


40 


FIELDIANA:  ZOOLOGY 


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HEANEY  ET  AL.:  THE  MAMMALS  OF  CAMIGUIN  ISLAND 


41 


fmnh);  Site  8  (2  fmnh);  Site  1 1  (4  dmnii);  Site  12 
(1  dmnh);  Site  13  (2  dmnii). 

Rutins  exulans  (Peale,  1848) 

Known  as  the  Polynesian  rat  or  the  spiny 
rice-field  rat,  R.  exulans  is  not  native  to  the 
Philippines;  through  mostly  accidental  dispersal 
by  humans,  it  occurs  from  Bangladesh  to  Easter 
Island  and  throughout  the  Philippines,  especially 
as  a  pest  in  agricultural  areas.  On  Camiguin,  we 
found  this  small  (60-75  g)  rat  to  be  moderately 
common  in  heavily  disturbed  lowland  agricul- 
tural land  at  ca.  150  m  (Site  3)  and  in  primary 
montane  forest  at  1275  m  (Site  7)  but  not  at 
other  sites.  Records  from  the  1960s  also  show  it 
to  be  present  in  mossy  forest  at  ca.  1500  m.  This 
distribution  is  similar  to  that  found  on  Negros, 
where  it  occurs  in  high-density  agricultural  areas 
and  in  mossy  forest  at  an  elevation  of  1500- 
1650  m  (Heideman  et  al.,  1987;  Heaney  et  al., 
1989).  Both  Camiguin  and  Negros  have  very  few 
native  rodents,  and  this  may  influence  the  ability 
of  nonnative  species  to  invade  the  forest  since  the 
nonnatives  are  absent  or  very  rare  in  mature 
forest  on  species-rich  islands  such  as  Leyte, 
Luzon,  and  Mindanao  (Heaney  et  al.,  1989, 
1999.  unpubl.  data;  Rickart  et  al.,  1993). 

In  March  1995,  one  nulliparous  female  weigh- 
ing 41  g  was  trapped.  Two  adult  males  (71-76  g) 
with  scrotal  testes  and  two  young  adults  (49- 
61  g)  with  scrotal  testes  were  trapped  at  the  same 
time.  They  typically  become  reproductive  at 
a  young  age  and  have  many  large  litters  each 
year  (Barbehenn  et  al.,  1973).  Cranial  and 
external  measurements  (Table  6)  are  comparable 
to  those  of  Dinagat,  Leyte,  and  Mindanao  (Mt. 
Kitanglad),  but  slight  variations  in  most  dimen- 
sions are  discernible  (Heaney  &  Rabor,  1982; 
Rickart  et  al.,  1993;  Heaney  et  al.,  unpubl.  data). 

Specimens  Examined — Total  15.  Site  3  (2  msu- 
iit);  Site  7  (5  fmnh);  Site  12  (1  dmnh);  Site  13  (7 

DMNH). 

Rattus  tanezwni  Temminck,  1844 

Previously  known  in  the  Philippines  as  Rattus 
rattus  and  Rattus  mindanensis,  the  Oriental  house 
rat  is  a  widespread  nonnative  rodent  in  the 
Philippines;  it  occurs  from  Afghanistan  to 
Indomalaya.  New  Guinea,  and  Micronesia 
(Musser  &  Carleton,  1993).  This  140  200-g  rat 
is  most  often  found  as  a  pest  in  urban  and 
agricultural  areas  but  can  be  common  in 
disturbed  forest  up  to  1800  m  (Heideman  et  al., 
1987;  Heaney  et  al.,  1989,  1991,  1998,  unpubl. 


data;  Rickart  et  al.,  1993).  On  Camiguin,  we 
found  R.  tanezwni  to  be  very  abundant  in  heavily 
disturbed  agricultural  land  at  150  m  (Site  3; 
Table  1).  It  was  less  common  in  secondary 
lowland  forest  at  1000  m  and  in  disturbed  lower 
montane  forest  at  1000  m  (Site  4)  and  1200  m 
(Site  5),  and  a  single  individual  was  trapped  in 
primary  mossy  forest  at  1400  m  (Site  6).  Four 
nestling  juveniles  were  found  in  a  new  slash-and- 
burn  clearing  near  Site  7,  but  no  R.  tanezwni 
were  caught  in  adjacent  mature  forest.  Taken 
together  with  records  from  the  1960s,  these  data 
indicate  that  R.  tanezwni  occurs  from  sea  level  to 
the  highest  peaks  on  Camiguin,  although  it  tends 
to  be  most  common  at  lower  elevations  and  in 
disturbed  habitats. 

Pregnant  females  had  a  mean  weight  of  126  ± 

57.6  g  (range  =  63-200  g,  N  =  4).  Females  with 
large  mammae  but  not  pregnant  had  a  mean 
weight  of  151  ±  16.7  g  (range  =  130-169  g,  N  = 
9);  three  of  these  had  placental  scars  (three,  four, 
and  eight  scars).  Nulliparous  females  had  a  mean 
weight  of  93  ±  39.8  g  (range  =  67-160  g).  Males 
with  scrotal  testes  had  a  mean  weight  of  169.4  ± 

22.7  g  (range  =  141-199  g,  N  =  8);  a  subadult 
with  scrotal  testes  weighed  105  g.  These  data 
indicate  large  litter  size  (up  to  eight)  and  very 
early  reproduction,  as  is  typical  for  the  species 
(Barbehenn  et  al.,  1973).  External  and  cranial 
measurements  (Table  6)  are  comparable  to  those 
of  Biliran,  Leyte,  Maripipi,  and  Mindanao,  with 
only  slight  variation. 

Specimens  Examined — Total  69.  Site  3  (21 
msu-iit);  Site  5  (4  msu-iit);  Site  6  (1  msu-iit);  Site 
7  (4  fmnh);  Site  9  (2  dmnh);  Site  10  (6  dmnh); 
Site  11  (3  dmnh);  Site  12  (6  dmnh);  Site  13  (22 
dmnh). 

Order  Carnivora 

Family  Viverridae — Civets 

Paradoxurus  hermaphroditus  Jourdan,  1837 

The  common  palm  civet  occurs  from  Sri 
Lanka  to  Hainan  and  the  Lesser  Sunda  Islands 
and  is  widespread  in  the  Philippines  (Heaney  et 
al.,  1998).  It  has  been  recorded  in  agricultural 
and  forested  areas  from  sea  level  up  to  at  least 
2400  m  (Heaney  et  al.,  1991,  1999,  unpubl.  data). 
We  did  not  capture  any  palm  civets,  but  in 
March  1995  we  were  given  a  mandible  from 
a  specimen  taken  by  local  hunters  in  the 
mountains  above  Mahinog,  thus  confirming  the 
earlier  report  by  Gray  (1843).  A  local  hunter 
identified  P.  hermaphroditus  from  pictures  as 
being  common  on  Camiguin. 


42 


FIELDIANA:  ZOOLOGY 


Specimens  Examined — Total  1.(1  fmnh). 

Viverra  tangalunga  Gray,  1832 

The  Malay  civet  (or  tangalung)  is  found  from 
peninsular  Malaysia  to  Sulawesi  and  is  wide- 
spread in  the  Philippines  (Heaney  et  al.,  1998).  It 
has  been  found  in  primary  and  secondary 
lowland,  montane,  and  mossy  forest  from  sea 
level  to  at  least  1700  m  (Rickart  et  al.,  1993; 
Heaney  et  al.,  1999).  We  did  not  obtain  any  on 
Camiguin,  but  a  local  hunter  identified  Viverra 
tangalunga  from  photographs  as  being  present 
on  Camiguin;  we  tentatively  accept  this  as  a  valid 
record. 

Order  Artiodactyla 
Family  Suidae — Pigs 

Sus  philippensis  Nehring,  1 886 

The  Philippine  warty  pig  is  a  Philippine 
endemic  that  occurs  in  the  Greater  Luzon, 
Greater  Mindanao,  and  Mindoro  faunal  regions; 
its  numbers  are  declining  (Oliver,  1992,  1999).  It 
formerly  was  abundant  from  sea  level  to  at  least 
2800  m  in  all  habitats;  now  it  is  common  only  in 
remote  forests  (Heaney  et  al.,  1991,  1999, 
unpubl.  data;  Oliver,  1992,  1999;  Garcia  & 
Deocampo,  1997). 

On  Camiguin  in  March  1995,  we  observed 
hoof  marks  of  wild  pigs  from  disturbed  lowland 
forest  at  1 000  m  up  to  primary  montane  forest  at 
1275  m  elevation  (Site  7).  Near  the  sampling  site 
at  1475  m  (Site  8),  we  saw  an  active  pig  nest,  and 
scattered  pig  trails  were  in  clear  evidence.  A  local 
hunter  said  that  they  were  commonly  hunted  and 
often  sold  in  a  small  local  market  at  Owakan, 
Mahinog  Municipality,  but  not  in  coastal  cities. 
We  were  given  two  adult  mandibles  by  hunters 
from  pigs  captured  in  forest  in  the  early  1 990s  in 
the  mountains  above  Mahinog. 

Specimens  Examined — Total  2.  Above  Mahi- 
nog (2  fmnh). 


Analysis  and  Discussion 

Adequacy  of  Sampling:  What  Is  Present  and  What 
Is  Absent? 

Before  interpreting  field  data,  it  is  necessary  to 
evaluate  the  extent  to  which  they  are  complete 
and  reliable.  In  doing  so  here,  we  follow  the 
procedures  used  by  Heaney  et  al.  (1989,  1991, 
1999)  and  Rickart  et  al.  (1993). 


Fruit  Bats — Because  pteropodid  bats  lack 
sonar  systems,  they  are  easily  captured  in  mist 
nets.  The  1960s  surveys,  which  focused  on  birds, 
yielded  many  fruit  bat  specimens  (Heaney,  1984), 
most  of  them  almost  certainly  from  mist  nets. 
Those  efforts  obtained  six  species  of  fruit  bats 
(Fig.  2).  Our  netting  in  1992-1994  yielded  no 
additional  species  but  was  not  extensive.  We 
consider  it  quite  possible  that  some  additional 
species  may  be  present,  especially  Eonycteris 
spelaea  and  Rousettus  amplexicaudatus,  since 
both  occur  very  widely  in  the  Philippines 
(Heaney  et  al.,  1998),  and  possibly  the  large 
Acerodon  jubatus  and  Pteropus  vampyrus,  which 
fly  high  above  the  canopy  and  so  are  difficult  to 
capture.  However,  our  sampling  was  sufficient 
that  we  consider  it  quite  unlikely  that  the  species 
that  are  abundant  and  easily  captured  on  Mind- 
anao and  associated  islands  (Heaney  et  al.,  1989, 
unpubl.  data;  Rickart  et  al.,  1993)  are  present  on 
Camiguin,  especially  Haplonycteris  fischeri  and 
Ptenochirus  minor  and  probably  Megaerops 
ecaudatus.  Suitable  habitat  for  the  high-elevation 
specialist  Alionycteris paucidentata  is  very  limited 
in  area  on  Camiguin,  if  present  at  all;  although 
we  did  limited  netting  in  that  habitat,  we  think  it 
very  unlikely  that  this  species  is  present. 

Sampling  along  the  elevation  gradient  proba- 
bly has  produced  a  partial  picture  of  variation  in 
species  richness  (Fig.  2)  but  as  noted  may  be 
incomplete  because  our  netting  in  1992-1995  was 
not  extensive  (Table  3). 

Insectivorous  Bats — Insectivorous  bats  use 
sophisticated  sonar  systems  to  navigate.  They  are 
thus  difficult  to  capture  in  mist  nets,  which  were 
our  only  means  of  capturing  them;  further,  our 
mist-netting  efforts  in  1992-1995  were  limited  in 
extent  (see  Heaney  &  Tabaranza,  2006a;  Table  3). 
For  this  reason,  we  believe  that  our  sample  of  six 
species  is  likely  to  be  quite  incomplete  and  is  not 
usable  for  estimates  of  species  richness,  as  is 
often  the  case  (Heaney  et  al.,  2002).  Our  data 
contribute,  however,  to  general  knowledge  of  the 
natural  history  of  these  poorly  known  animals. 

Nonvolant  Small  Mammals — Shrews,  ro- 
dents, and  other  small  mammals  that  can  be 
captured  in  traps  were  poorly  sampled  by  the 
1 960s  field  teams,  which  focused  their  efforts  on 
birds  (and  secondarily  on  fruit  bats).  During 
1994  and  1995,  we  accumulated  2783  trap-nights 
at  six  sites  that  included  all  major  habitat  types 
on  the  island,  yielding  188  captures  (Table  1).  A 
species-accumulation  curve  for  these  data  (Fig.  3) 
shows  that  the  number  of  species  recorded  for 


HEANEY  ET  AL.:  THE  MAMMALS  OF  CAMIGUIN  ISLAND 


No.  Species 


200   400  600   800  1000  1200  1400  1600  1800  2000  2200  2400  2600  2800 

No.  Trap-Nights 

Fig.  3.     Species  accumulation  curve  for  nonvolant  small  mammals  on  Camiguin  Island.  The  points  shown  are 
for  Sites  1  and  3  to  6.  as  described  in  the  text. 


the  island  reached  a  peak  of  eight  species 
(including  five  nonnative  species)  by  the  end  of 
sampling  at  the  second  site  (1055  trap-nights) 
and  showed  no  further  gain  as  we  approached 
the  total  of  nearly  2800  trap-nights.  Local 
hunters  either  were  unfamiliar  with  squirrels  or 
knew  them  from  Mindanao;  given  their  conspic- 
uousness  (Heaney  et  al.,  unpubl.  data),  they  are 
very  unlikely  to  be  present.  We  conclude  that  the 
island  is  unlikely  to  support  other  species  of 
small  mammals  that  are  present  on  Mindanao, 
including  gymnures  (Podogymnura),  tree  shrews 
(Urogale),  squirrels,  or  murid  rodents  (e.g., 
Batomys,  Limnomys,  and  Tarsomys).  Further, 
the  sampling  along  the  elevational  gradients  was 
sufficiently  intense  that  we  are  likely  to  have  an 
accurate  assessment  of  the  pattern  of  species 
richness  above  about  800  m,  although  it  must  be 
noted  that  extensive  habitat  destruction  at  lower 
elevations  may  have  had  an  extensive  impact  on 
the  current  patterns  (discussed  below). 

Medium  and  Large  Mammals — We  collected 
information  only  on  the  presence  and  absence  of 
large  mammals  and  a  mixed  set  of  medium-sized 
species  (none  of  which  are  readily  captured  in 
traps  such  as  the  ones  we  used),  not  on  detailed 
habitat  use.  We  observed  carefully  during  all  our 
time  in  the  field  for  any  evidence  of  additional 
species,  and  we  interviewed  many  residents  of  the 
islands  and  showed  them  photos  of  potential 
species;  some  of  these  residents  are  active 
hunters.  Neither  we  nor  our  informants  had 
any  evidence  for  tarsiers,  flying  lemurs,  or  deer 
on  the  island,  though  many  people  knew  of  deer 
and  tarsiers  on  Mindanao,  and  a  few  knew  of 
flying  squirrels  and  flying  lemurs.  We  consider  it 
unlikely  that  species  other  than  those  listed  here 
are   present   given   the   voucher   specimens   we 


obtained  and  the  concordance  between  our 
specimens  and  the  information  received  from 
local  hunters.  Thus,  we  conclude  that  deer  are 
absent,  along  with  the  smaller  but  conspicuous 
species  such  as  tarsiers,  squirrels,  and  flying 
lemurs. 

Elevational  Patterns 

Though  our  limited  netting  prevents  us  from 
commenting  on  the  patterns  of  diversity  and 
abundance  of  bats  along  the  elevational  gradient 
on  Camiguin,  our  trapping  data  (Table  1)  pro- 
vide a  reasonably  complete  picture  of  the  general 
patterns  among  nonvolant  species.  The  highest 
density  we  encountered  was  at  Site  3,  in 
agricultural  land  at  ca.  150  m.  However,  these 
were  all  nonnative  rats  and  a  shrew,  representing 
only  three  species,  a  pattern  that  we  have  often 
seen  (e.g.,  Heaney  et  al.,  1989;  Rickart  et  al., 
1991).  The  five  native  small  mammal  species 
were  all  present  (or  suspect  to  be  present)  from 
1000  to  1300  m  and  declined  in  species  richness 
to  two  species  near  the  peak  of  Mt.  Timpoong  at 
Site  8  (Table  1).  Density  (measured  as  the 
number  captured  per  100  trap-nights)  more  than 
doubled  from  1000  to  1275  m,  then  fell  to  a  much 
lower  level  near  the  peak  at  1475  m.  However, 
much  of  the  change  in  density  was  driven  by  the 
remarkably  large  numbers  of  Suncus  murium  at 
the  upper  elevations;  they  were  not  taken  at  1000 
or  1200  m  (though  they  were  taken  at  150  m), 
were  fairly  common  at  1300  and  1475  m,  and 
were  exceptionally  abundant  at  1275  m,  where 
we  often  saw  and  heard  them  during  the  day. 
These  patterns  for  species  richness,  though 
limited  in  extent,  are  similar  to  those  from 
elsewhere    in    the    Philippines,    where    species 


44 


FIELDIANA:  ZOOLOGY 


richness  is  generally  low  at  low  elevations, 
increases  to  about  the  area  of  transition  from 
montane  to  mossy  forest,  and  declines  with 
increasing  elevation  in  mossy  forest  (Rickart  et 
al.,  1991;  Heaney,  2001).  The  pattern  of  overall 
abundance  is  also  similar  to  that  documented 
elsewhere,  except  that  Suncus  murinus  is  re- 
markably abundant  in  the  high-elevation  mon- 
tane and  mossy  forest,  where  usually  it  is  absent 
(Heaney,  2001).  We  have  previously  noted  the 
abundance  of  S.  murinus  in  similar  habitat  only 
once,  on  Negros  Island  (Heaney  et  al.,  1989), 
which  is  notably  depauperate  of  small  mammals 
(Heaney,  1986).  This  raises  a  question:  Is 
Camiguin  depauperate,  relative  to  its  area,  in 
comparison  to  other  islands  in  the  Philippines? 
This  question  is  addressed  in  the  next  section. 

Biogeographic  Implications 

Bats — Because  our  focus  in  conducting  field 
studies  concentrated  primarily  on  nonvolant 
mammals,  we  note  only  that  we  are  confident 
that  the  fruit  bat  community  on  Camiguin  is  at 
least  somewhat  depauperate  relative  to  similar 
elevations  on  Mindanao  and  associated  islands. 
At  least  two  species  usually  common  at  lower 
and  middle  elevations,  Haplonycteris  fischeri  and 
Ptenochirus  minor,  are  almost  certainly  absent. 
Both  of  these  are  primarily  associated  with  old- 
growth  rain  forest  and  generally  do  not  travel 
long  distances  out  from  under  the  canopy  of 
forest  (Heaney  et  al.,  1989,  1998;  Heideman  & 
Heaney,  1989;  Rickart  et  al.,  1993).  Some 
additional  species  probably  also  are  absent 
(Alionycteris  paucidentata  and  Megaerops  wet- 
morei);  they  also  are  associated  with  old-growth 
rain  forest,  though  A.  paucidentata  in  a  distinctive 
high-elevation  habitat  (M.  wetmorei  is  poorly 
known).  This  level  of  reduction  in  species 
richness  is  striking  in  view  of  the  tight  correlation 
between  species  richness  and  island  area  pre- 
viously found  (Heaney,  1991a).  However,  the 
uncertainty  regarding  the  potential  absence  of 
Eonycteris  and  Rousettus  leads  us  to  be  cautious 
in  defining  the  extent  to  which  species  richness  is 
low. 

Nonvolant  Mammals — The  data  are  more 
certain  regarding  both  small  mammals  (shrews 
and  murid  rodents)  and  the  medium  and  larger 
mammals  (deer,  squirrels,  and  so  on).  It  seems 
certain  that  many  genera  present  in  the  lowlands 
and  in  montane  habitats  on  Mindanao  and 
associated  islands  are  absent:  tree  shrews  (Ur- 


ogale),  flying  lemurs  (Cynocephalus),  tarsiers 
(Tarsius),  tree  squirrels  (Sundasciurus),  pygmy 
squirrels  (Exilisciurus),  flying  squirrels  (Peti- 
nomys),  a  murid  (Tarsomys  echinatus),  and  deer 
(Cervus).  Among  the  murid  rodents,  Batomys  is 
also  absent  though  often  found  in  montane 
forest  on  Mindanao  and  associated  islands 
(Rickart  et  al.,  1993).  Limnomys  bryophilus,  L. 
sibuanus,  and  Tarsomys  apoensis  also  occur  on 
Mindanao  but  only  above  about  1900  m  eleva- 
tion (Musser  &  Heaney,  1992;  Heaney  et  al., 
1998;  Rickart  et  al.,  2003). 

Those  species  of  nonvolant  mammals  that  are 
present  on  Camiguin  form  a  group  that  is  quite 
consistent  in  one  respect.  Not  all  lowland  species 
from  Mindanao  are  present,  but  all  those  that 
are  present  on  Camiguin  are  among  the  relatively 
few  species  that  are  common  in  the  lowlands  of 
Mindanao — or  were  present  before  human  de- 
struction of  lowland  rain  forests.  As  shown  by 
Musser  and  Heaney  (1992),  Heaney  (2001),  and 
Heaney  et  al.  (unpubl.  data),  Crocidura,  Apomys, 
Bullimus,  Crunomys,  Rattus  everetti,  Paradox- 
urus,  Viverra,  and  Sus  are  among  the  few 
mammals  that  occur  (or  occurred  until  recently) 
on  Mindanao  in  lowland  forest.  With  only  one 
exception  (Tarsomys  echinatus),  all  those  low- 
land species  of  Mindanao  noted  above  as  being 
absent  from  Camiguin  are  arboreal  species.  In 
other  words,  the  nonvolant  mammal  fauna  of 
Camiguin  is  composed  only  of  nonarboreal  small 
mammals  from  Mindanao  (or  are  their  sister 
taxa)  and  all  the  lowland  larger  mammals  (all  of 
which  are  also  not  arboreal).  We  conclude  that 
the  mammalian  fauna  of  Camiguin  is  highly 
biased;  that  is,  it  is  composed  entirely  of  species 
shared  with  and/or  derived  from  Mindanao,  but 
it  is  not  a  random  sample  of  the  Mindanao 
fauna;  rather,  it  is  comprised  of  species  that 
occur  (or  occurred  before  deforestation)  on  the 
lowland  forest  floor  of  Mindanao,  not  in  the 
forest  canopy  and  not  in  the  montane  or  mossy 
forest. 

Does  the  absence  of  arboreal  mammals  mean 
that  Camiguin  has  a  species-poor  nonvolant 
mammal  fauna  relative  to  other  islands  in  the 
Philippines?  Perhaps  surprisingly,  the  answer  is 
clearly  no.  With  nine  native  nonvolant  mam- 
mals, Camiguin  falls  almost  precisely  on  the 
same  species-area  curve  as  the  islands  that  made 
up  Greater  Mindanao  (Mindanao,  Leyte,  Bohol, 
Biliran,  and  Maripipi).  While  the  fauna  is  biased 
toward  lowland,  ground-living  murid  rodents, 
and  small  omnivorous  carnivores,  the  species 


HEANEY  ET  AL.:  THE  MAMMALS  OF  CAMIGUIN  ISLAND 


45 


richness  is  not  reduced  relative  to  islands  that 
were  recently  connected  to  Mindanao  itself 
(Heaney.  1986;  Rickart  et  al.,  1993).  However, 
the  nonvolant  small  mammal  community  is 
apparently  sufficiently  species-poor  to  allow 
a  nonnative  shrew  (Suncus  minimis)  to  invade 
primary  montane  forest.  The  presence  of  this 
shrew  in  similar  habitat  has  been  noted  on 
Negros  Island,  which  is  also  species  poor 
(Heaney  et  al.,  1989),  but  not  on  other  islands 
in  the  Philippines  (e.g.,  Rickart  et  al..  1991. 
1993).  We  predict  that  S.  minimis  will  be  found 
in  primary  montane  forest  on  other  islands  in  the 
Philippines  with  five  or  fewer  native  shrews  and 
rodents. 

Conservation  and  Management 

As  noted  by  Heaney  and  Tabaranza  (1997, 
2006a),  the  native  mammal  fauna  of  Camiguin  is 
dependent  on  the  continued  survival  of  good- 
quality  forest  at  all  elevations.  The  two  mammal 
species  unique  to  Camiguin,  in  particular, 
apparently  depend  on  forest  with  little  or  no 
disturbance,  most  of  which  currently  occurs 
in  steep  areas  above  800  m  elevation.  These 
forested  areas  are  also  crucial  to  the  well-being 
and  stability  of  the  human  population,  for 
they  are  the  source  of  water  for  the  island  and 
protect  the  lowlands  from  potentially  devastat- 
ing floods  and  landslides.  Additionally,  our 
observations  indicate  that  the  medium  and  large 
mammals  have  been  depleted  by  overhunting; 
these  require  protection  if  they  are  to  survive. 
The  people  of  Camiguin  benefit  both  personally 
and  economically  from  the  beautiful  landscape 
and  seashores  of  the  island,  both  through  the 
tourism  they  make  possible  and  from  the 
environmental  stability  they  engender.  Protect- 
ing the  forests  will  benefit  the  people  and  wildlife 
equally. 


Acknowledgments 

Our  thanks  go  to  the  Protected  Areas  and 
Wildlife  Bureau  of  the  Philippine  Department  of 
Environment  and  Natural  Resources,  which 
provided  encouragement  and  permits,  and  the 
Region  10  office  of  the  DENR  for  logistical 
support.  Specimens  for  comparison  were  kindly 
loaned  by  G.  Hess  (dmnh)  and  P.  Myers  (ummz) 
and  M.  Carleton,  L.  Gordon,  and  H.   Kafka 


(lsnm).  The  manuscript  was  much  improved  by 
comments  from  P.  D.  Heideman  and  E.  A. 
Rickart.  We  thank  Lisa  Kanellos  for  preparation 
of  the  figures.  Funding  was  provided  by  the 
Marshall  Field  Fund.  Ellen  Thorne  Smith  Fund, 
and  the  Barbara  Brown  Fund  for  Mammal 
Research  of  the  Field  Museum  of  Natural 
History. 


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48 


FIELDIANA:  ZOOLOGY 


A  New  Species  of  Hanging-Parrot  (Aves:  Psittacidae: 
Loriculus)  from  Camiguin  Island,  Philippines 

Jose  G.  Tello,1'2'3  Jacob  F.  Degner,1  John  M.  Bates,1  and  David  E.  Willard1 


Abstract 

A  new  species  of  Hanging-Parrot  or  Colasisi,  Loriculus,  is  described  from  a  series  of  23 
specimens  (19  males,  4  females)  collected  in  the  1960s  on  Camiguin  Island,  Camiguin 
Province,  Philippines,  at  elevations  between  300  and  1350  m.  The  new  species  lacks  sexual 
dimorphism  in  plumage  coloration,  which  distinguishes  it  from  all  other  members  of  the  L. 
philippensis  group  and  all  other  Loriculus.  The  overall  color  pattern  of  the  new  species  appears 
most  like  females  of  L.  p.  worcesteri  and  L.  p.  apicalis  but  differs  in  plumage  characteristics 
(the  width  and  extension  of  the  orange-scarlet  crown  patch,  the  amount  and  intensity  of  blue 
in  the  face  and  thighs,  and  the  intensity  of  the  blue  in  the  tail  above  inner  edges  and  the  tail 
below).  In  addition,  males  of  the  new  species  are  larger  than  males  of  nearby  populations  of  L. 
philippensis,  having  significantly  longer  tails  and  wing  chords.  Nothing  is  known  about  the 
habits  of  the  new  species;  however,  the  small  size  of  the  island  of  Camiguin,  coupled  with 
extensive  deforestation,  makes  the  status  of  the  new  species  a  significant  conservation 
concern. 


Introduction 

The  Philippine  Hanging-Parrot  or  Colasisi 
{Loriculus  philippensis)  has  ten  described  sub- 
species distributed  throughout  the  islands  of  the 
Philippines  (Dickinson  et  al.,  1991;  Collar,  1997; 
Juniper  &  Parr,  1998;  Kennedy  et  al.,  2000).  The 
subspecies  L.  p.  apicalis  has  been  reported  to 
occur  on  the  islands  of  Bazol,  Balut,  Camiguin, 
Dinagat,  Mindanao,  and  Siargao  (Fig.  1).  How- 
ever, Austin   Rand,   a  former  Field   Museum 


1  Field  Museum  of  Natural  History,  1400  South 
Lake  Shore  Drive,  Chicago,  IL  60605-2496,  U.S.A. 

2  Biological  Sciences,  University  of  Illinois  at 
Chicago,  Chicago,  IL  60607,  U.S.A. 

3  Current  address:  Division  of  Vertebrate  Zoology- 
Ornithology,  American  Museum  of  Natural  History, 
Central  Park  West  at  79th  Street,  New  York,  NY 
10024-5192,  U.S.A. 


curator  and  Philippine  expert  (e.g.,  Rand  & 
Rabor  1960,  1969),  had  penciled  the  notation 
"subsp.  nov"  on  the  tag  of  a  specimen  from 
Camiguin  in  the  fmnh  collection.  He  never 
published  a  description.  Here,  we  quantitatively 
evaluate  the  external  morphology  and  compare 
the  plumage  color  of  specimens  referred  to  L.  p. 
apicalis  from  Camiguin  with  L.  p.  apicalis  from 
Mindanao  and  specimens  of  other  subspecies  of 
L  philippensis.  Our  results  demonstrate  that  the 
Camiguin  population  of  L.  p.  apicalis  is  separa- 
ble from  all  other  populations  of  L.  philippensis 
in  plumage.  It  is  further  separable  from  all 
neighboring  populations  in  body  size.  We  argue 
that  these  differences  warrant  designating  this 
population  as  a  distinct  species  and  not  a  sub- 
species of  L.  philippensis.  Similar  arguments  have 
been  made  in  separating  L.  bonapartei  of  the 
Sulu  Archipelago  from  the  L.  philippensis  com- 
plex (Juniper  &  Parr  1998).  We  present  a  formal 


FIELDIANA:  ZOOLOGY,  N.S.,  NO.  106,  APRIL  5,  2006,  PP.  49-57 


49 


Fig.  1.     Map  of  the  central  and  southern  Philippines  showing  the  locations  of  islands  referred  to  in  the  text. 


50 


FIELDIANA:  ZOOLOGY 


description  of  the  new  taxon  based  on  plumage 
and  morphology  of  adult  males. 


Methods 

All  qualitative  color  comparisons  were  made 
under  natural  light.  Color  names  follow  Smithe 
(1975),  and  each  color  name  (capitalized)  is 
followed  by  its  number  in  parentheses.  J.F.D. 
measured  wing  chord,  tarsus  length,  tail  length 
(from  point  of  insertion  of  central  rectrices  to  tip 
of  longest  rectrix),  culmen  length,  bill  length 
(from  anterior  edge  of  nostril),  bill  height  (at 
anterior  edge  of  nostril),  bill  width  (at  anterior 
edge  of  nostril),  and  gape  width  with  calipers  to 
the  nearest  0.05  mm.  Specimens  were  measured 
randomly  to  avoid  any  investigator  bias. 

All  statistical  analyses  were  carried  out  using 
the  program  Statistica  (Statsoft  Inc.  1995). 
Mensural  data  were  tested  for  normality  using 
Kolmogorov-Smirnov  tests  and  Lilliefors  prob- 
abilities prior  to  all  the  analyses.  Mensural 
differences  between  males  and  females  within 
each  study  population  were  evaluated  using  one- 
way analyses  of  variance  (ANOVAs).  We  used 
both  univariate  (one-way  ANOVAs  and  Bonfer- 
roni  post  hoc  tests)  and  multivariate  analysis 
(principal  components  and  discriminant  func- 
tion) on  log-transformed  data  to  test  for 
mensural  differences  between  specimens  from 
Camiguin  and  those  from  neighboring  popula- 
tions of  L.  philippensis.  Multivariate  analyses 
were  used  to  reduce  the  dimensionality  of  data 
and  facilitate  the  analysis  of  morphology  in  two 
or  three  dimensions  (Pimentel,  1979);  we  used  the 
varimax  raw  method  to  rotate  the  three  compo- 
nents that  are  reported  in  the  principal  compo- 
nents analysis  in  order  to  improve  interpretabil- 
ity  of  the  resulting  patterns.  Collecting  localities 
are  described  by  Heaney  and  Tabaranza  (2006a). 

Loriculus  camiguinensis,  new  species 

Camiguin  Hanging-Parrot 

Holotype — Field  Museum  of  Natural  History 
No.  284389,  adult  male  from  Kasangsangan, 
Municipality  of  Catarman,  Camiguin  Province, 
Camiguin  Island,  Philippines,  elevation  between 
1000  and  2000  ft  (300-600  m;  approx.  9°11'N, 
124°40'E;  see  Heaney  &  Tabaranza,  2006a,  for 
more  explanation  of  this  and  other  collecting 


localities  on  Camiguin  Island),  18  June  1968, 
collected  by  D.  S.  Rabor  and  W.  Sanguila. 

Diagnosis — A  Loriculus  hanging-parrot  with 
characteristics  of  the  philippensis  group  (see 
Front  Plate).  In  contrast  to  other  members  of 
this  group,  L.  camiguinensis  is  characterized  by 
a  lack  of  sexual  dimorphism  in  plumage  color- 
ation. The  overall  color  pattern  of  L.  camigui- 
nensis is  most  like  females  of  L.  p.  worcesteri 
from  Bohol,  Leyte,  and  Samar  and  L.  p.  apicalis 
from  Mindanao  but  differs  as  follows:  (1)  the 
scarlet  of  the  crown  of  L.  camiguinensis  does  not 
extend  as  far  onto  the  bright  olive  green  nape  as 
it  does  in  both  males  and  females  of  L.  p.  apicalis 
and  L.  p.  worcesteri;  this  pattern  differs  from  that 
of  chrysonotus,  siquijorensis,  regulus,  bournsi, 
mindorensis,  and  philippensis,  in  which  the  scarlet 
crown  is  highly  reduced.  (2)  The  width  of  the 
scarlet  crown  in  L.  camiguinensis  narrows  at  the 
rear  edge  instead  of  being  rounded  as  in  all  other 
populations.  (3)  The  scarlet  (sometimes  orange) 
throat  patch  that  is  typical  of  males  in  L. 
philippensis  is  lacking  in  L.  camiguinensis;  five 
of  the  Camiguin  males  had  data  on  gonadal 
development,  reporting  enlarged  or  slightly 
enlarged  testicles,  which  gives  an  indication  of 
their  adult  condition.  (4)  The  face  of  L. 
camiguinensis  is  extensively  turquoise  blue  and 
differs  from  that  of  females  of  L.  philippensis 
subspecies  in  that  the  blue  of  the  face  is  darker 
and  more  extensive,  extending  over  the  eye  and 
onto  the  throat.  (5)  The  turquoise  blue  in  the 
thighs  of  L.  camiguinensis  is  darker  than  that 
of  females  of  L.  philippensis  populations.  (6) 
The  blue  in  the  inner  edges  of  the  rectrices  above 
and  throughout  below  is  darker  in  L.  camigui- 
nensis. (7)  Mean  wing  chord  and  tail  length  of 
males  and  tail  length  of  females  of  L.  camigui- 
nensis are  significantly  longer  than  those  of 
nearby  L.  philippensis  subspecies  (Tables  1  and 
2).  (8)  The  overall  green  plumage  is  a  darker 
shade  with  less  of  a  yellowish  tinge,  especially  on 
the  back. 

Description  of  Holotype — General  plumage 
Parrot  Green  (160)  with  slightly  orange  tinge  in 
the  upperparts,  more  yellowish  tinge  on  under- 
pays; forehead  and  forepart  of  crown  Scarlet 
(14)  fading  to  orange  at  rear-edge;  thin  Orange 
Yellow  (18)  band  on  nape;  lores,  chin,  cheek,  and 
throat  closest  to  Turquoise  Blue  (65);  rump  and 
upper  tail-coverts  scarlet;  Turquoise  Green  (64) 
markings  on  the  sides  of  the  rump;  thighs  slightly 
paler  Turquoise  Blue;  rectrices  Emerald  Dark 
Green  (262)  above  with  dark  Cerulean  Blue  (67) 


TELLO  ET  AL.:  A  NEW  SPECIES  OF  HANGING-PARROT 


51 


tinges  on  inner  edges  of  all  but  central  reetrices; 
rectrices  dark  Cerulean  Blue  below;  flight  feath- 
ers black  above  with  Emerald  Dark  Green  outer 
edges,  flight  feathers  black  below  with  inner 
edges  with  extensive  Cerulean  Blue;  greater 
underwing-coverts  Cerulean  Blue  and  lesser  un- 
derwing-coverts  closest  to  Spectrum  Green  (62). 
Soft  part  colors  of  dried  specimen:  upper 
mandible  closest  to  Spectrum  Orange  (17)  at 
base  grading  to  yellow  with  Gray  Horn  (91)  at 
tip  and  along  tomia;  lower  mandible  with  similar 
pattern,  but  orange  at  the  base;  cere  Grayish- 
Horn;  feet  and  tarsi,  yellow-horn. 

Measurements  of  Holotype  (mm) — Wing 
(99.8),  tarsus  (11.7),  tail  (49.5),  culmen  (18.0), 
bill  length  (15.0),  bill  height  (11.1),  bill  width 
(6.4),  gape  width  (8.2). 

Distribution — Loriculus  camiguinensis  is 
known  only  from  the  forests  of  Camiguin  Island. 
Specimens  have  been  collected  between  1000  and 
4500  ft  (300-1350  m)  in  the  municipalities  of 
Catarman  and  Mahinog  (Balete  et  al.,  2006; 
Heaney  &  Tabaranza,  2006a). 

Etymology — We  name  this  species  after  the 
Philippine  Island  of  Camiguin,  to  which  this 
species  appears  to  be  endemic. 

Specimens  Examined — We  examined  the  fol- 
lowing specimens  from  Field  Museum  of  Natural 
History  (fmnh)  and  Delaware  Museum  of 
Natural  History  (dmnh): 

L.  camiguinensis  (Camiguin  Island)  (19  males, 
4  females,  all  known  specimens  of  this  new 
taxon):  Camiguin  Province:  Catarman  Munici- 
pality; Kasangsangan  (males:  fmnh  holotype, 
fmnh  284391,  284392,  284393;  dmnh  19950, 
19958,  19960,  19961,  19962:  females  fmnh 
284390;  dmnh  19959);  Camiguin  Province:  Cat- 
arman Municipality;  Catarman  Mountain  (male: 
dmnh  19949);  Camiguin  Province:  Mahinog 
Municipality;  Matugnao,  Mt.  Timpoong  (males: 
fmnh  286742,  286743,  286744,  286745;  dmnh 
19951,  19952,  19953,  19954,  19965;  females: 
fmnh  286746,  286747). 

L.  p.  apicalis  (Mindanao  Island)  (19  males,  5 
females):  North  Cotabato  Province:  Mt.  Apo, 
Todaya  (male:  fmnh  184090);  North  Cotabato 
Province:  Mt.  Apo,  Galog  (male:  dmnh  36227); 
Agusan  del  Norte:  Mt.  Hilong-Hilong,  Lewed 
(male:  fmnh  275003);  Misamis  Occidental:  Zam- 
boanga  Peninsula,  Mt.  Malindang,  Gandwan 
(males:  fmnh  227136,  227138,  227139;  female: 
fmnh  227137);  Misamis  Occidental:  Zamboanga 
Peninsula.  Mt.  Malindang,  Masawan  (males: 
fmnh  227134,  227135);  Davao  Oriental  Province; 


Mati:  Mt.  Mayo,  Limot  (male:  fmnh  277864); 
Misamis  Oriental  Province:  Manticao:  Tuod, 
Camp  Dundue  (male:  fmnh  283788;  female 
fmnh  283787);  Misamis  Oriental  Province:  Opol: 
Malubato  (male:  fmnh  283785);  South  Cotabato: 
Tupi:  Mt.  Matutum  (male:  fmnh  279330); 
Bukidnon  Province:  Malaybalay,  Mt.  Katanglad 
(male:  fmnh  262475,  262476;  female:  fmnh 
262474);  Bukidnon  Province:  Lantapan:  Ko- 
toon,  Mt.  Katanglad  SE  slope  (male:  dmnh 
2983);  Lanao  Norte  Province:  Iligan  City, 
Mainit,  Mahayahay  (female:  fmnh  283786); 
Surigao  del  Sur  Province:  Car-Can-Mad-Lan 
area  (female:  fmnh  275002);  Zamboanga  del  Sur 
Province:  Zamboanga  (male:  dmnh  36993); 
Davao  Oriental  Province:  Sigaboy  (males:  dmnh 
36224,  36226);  Davao  del  Sur  Province:  Padada 
(male:  dmnh  36233). 

L.  p.  worcesteri  (11  males,  8  females):  Bohol 
Island:  Bohol  Province:  Sierra  Bullones  (males: 
fmnh  223025,  223026,  223029,  223030,  223034, 
223036,  223037;  females:  fmnh  223027,  223327, 
223028,  223033,  223035,  223039);  Leyte  Island: 
Leyte  Province:  Burauen,  Buri,  Ma-Alngon 
(male:  fmnh  276302;  female:  fmnh  276300); 
Leyte  Province:  Burauen,  Buri,  Mt.  Lobi  range, 
Tambis  (male:  fmnh  276299;  female:  fmnh 
276298);  Samar  Province:  Mt.  Capotoan  (male: 
fmnh  247411);  Western  Samar  Province:  Matu- 
guinao  (male:  fmnh  247410). 

L.  p.  regulus  (Negros  Island)  (5  males,  4 
females):  Negros  Oriental  Province:  Bayawan, 
Basay  (male:  fmnh  257121);  Negros  Oriental 
Province:  Santa  Catalina,  Inubungan  (male: 
fmnh  219314;  male:  fmnh  188579);  Negros 
Oriental  Province:  Sicopon  River  (male:  fmnh 
185483);  Negros  Oriental  Province:  Amio  (males: 
fmnh  188545,  188548;  females:  fmnh  188544, 
188553);  Negros  Oriental  Province:  Pamo-at 
(male:  fmnh  188550). 

L.  p.  chrysonotus  (captive  specimen,  presum- 
ably from  Cebu  Island)  (1  male):  Cebu  Province: 
Exact  locality  unknown  (fmnh  252666). 

L.  p.  siquijorensis  (Siquijor  Island)  (1  male): 
Siquijor  Province:  San  Juan:  Tag-ibo  (fmnh 
222741). 

L.  p.  bournsi  (Sibuyan  Island)  (1  male,  1 
female):  Romblon  Province:  Goangan,  3  km  SE 
Magdiwang  (male:  fmnh  358288);  Romblon 
Province:  Exact  locality  unknown  (female:  fmnh 
11081). 

L.  p.  mindorensis  (Mindoro  Island)  (1  male, 
1  female):  Oriental  Mindoro  Province: 
Calapan  (male:  fmnh  19927);  Occidental  Mind- 


52 


FIELDIANA:  ZOOLOGY 


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TELLO  ET  AL.:  A  NEW  SPECIES  OF  HANGING-PARROT 


53 


Table  2.  Univariate  statistical  comparisons  be- 
tween /..  camiguinensis  and  adjacent  populations  ol'  L. 
philippensis.  Populations  ( 1  =  L.  camiguinensis:  2  =  L. 
p.  apicalis;  3  =  L.  p.  worcesteri;  and  4  =  L  p.  regains) 
are  ordered  based  on  their  mean  variation,  from 
smallest  to  largest  (left  to  right).  *  F  values  signifi- 
cantly different  at  P  <  0.05  (one-way  analysis  oi' 
variance  and  Bonferroni  post-hoc  test).  For  each 
morphometric  variable,  populations  united  by  the 
underlines  showed  non-significant  differences. 


Morphometric 
variables 


Populations 


Wing  chord 

30.5* 

Tail  length 

23.1* 

Tarsus  length 

14.8* 

Gape  width 

9.8* 

Bill  length 

3.7* 

Total  culmen 

3.5* 

Bill  height 

4.0* 

Bill  width 

5  2* 

2 

3 

4 

2 

4 

3 

2 

3 

4 

2 

3 

4 

2 

4 

3 

2 

3 

4 

4 

2 

3 

2 

4 

3 

oro  Province:  Abra  de  Hog  (female:  fmnh 
210845). 

L.  p.  philippensis  (Luzon  Island)  (1  male.  1 
female):  Bataan  Province:  Mariveles  (male:  fmnh 
73966);  Cagayan  Province:  Sierra  Madre,  Mt. 
Cagua  (female:  fmnh  258827). 

Morphometric  Differences — One-way  ANO- 
VAs  on  all  known  L.  camiguinensis  specimens 
and  specimens  in  the  fmnh  collection  from 
neighboring  islands  showed  that  certain  variables 
differed  between  sexes  in  some  of  the  study 
populations,  but  no  consistent  pattern  was  found 
(Table  1).  Because  of  the  small  sample  of  females 
available  for  this  study,  we  include  only  males  in 
the  analyses. 

Results  of  the  one-way  ANOVAs  (effect  = 
population)  and  Bonferroni  post  hoc  tests 
showed  that  males  of  L.  camiguinensis  are 
morphologically  distinct  from  males  of  other  L. 
philippensis  populations.  Wing  chord,  tail  length, 
and  tarsus  length  were  significantly  longer  in  L. 
camiguinensis  males  than  in  the  other  popula- 
tions (Table  2). 

Principal  components  analysis  for  males  from 
all  populations  resulted  in  three  rotated  (varimax 
raw)  factors  with  Eigenvalues  close  to  or  greater 
than  1.0,  which  together  explained  73.2%  of  the 
total  variance  (Table  3).  Axes  defined  by  the  first 
and  third  component  (particularly  the  third) 
demonstrate  separation  of  L.  camiguinensis  from 
the  other  populations  (Fig.  2).  The  first  compo- 
nent explained  45.2c/c  of  the  total  variance  and 


Table  3.  Fraction  of  total  variance  explained  by 
each  of  the  first  three  components  of  the  principal 
components  analyses.  The  three  components  were 
rotated  (see  Methods). 


Morphometric 
variables 


PCI 


PCII       PCIII 


Wine  chord  0.25  0.29  0.84 

Tarsus  length  0.29  0.31  0.40 

Tail  length  0.16  0.04  0.94 

Total  culmen  0.14  0.88  0.18 

Bill  length  0.05  0.92  0.09 

Bill  height  0.77  0.15  0.17 

Bill  width  0.80  -0.04  0.21 

Gap  width  0.71  0.33  0.35 

Eigenvalue  3.62  1.34  0.90 

Explained  variance  (%)      45.21  16.79  11.21 


had  high  positive  correlations  with  bill  width,  bill 
height  and  gape  width  (0.80,  0.77,  and  0.71. 
respectively).  The  second  component  explained 
16.8%  of  the  total  variance  and  had  high  positive 
correlation  with  bill  length  and  culmen  length 
(0.92  and  0.88,  respectively).  The  third  compo- 
nent explained  a  further  1 1 .2%  of  the  variance 
and  had  the  highest  positive  correlation  values 
for  tail  length  and  wing  chord  (0.94  and  0.84. 
respectively).  It  was  this  latter  axis  that  differ- 
entiates L.  camiguinensis  from  the  other  popula- 
tions. 

A  discriminant  function  analysis  using  these 
morphometric  variables  was  also  significant  (P 
<  0.001)  and  correctly  classified  88%  of  the  cases 
(plot  not  shown).  The  standardized  coefficients 
of  the  discriminant  function  separating  males  of 
L.  camiguinensis  from  those  of  other  L.  philip- 
pensis populations  weighted  tarsus  length  and 
wing  chord  heavily  (0.80  and  0.80,  respectively), 
followed  by  tail  length  (0.46)  and  bill  length 
(0.40),  with  all  the  other  variables  having 
coefficients  under  0.32. 

A  Question  of  Correctly  Sexed  Specimens — 
Unpublished  concern  has  been  expressed  re- 
garding the  ability  of  Rabor  and  his  field 
assistants  to  accurately  determine  the  sex  of 
specimens.  This  possibility  presents  a  serious 
issue  with  respect  to  interpreting  the  data  at 
hand.  In  an  attempt  to  verify  sexing.  we  tried 
unsuccessfully  to  amplify  DNA  from  toe  pads  of 
some  specimens  using  commercially  available 
primers  for  sex-linked  DNA.  However,  for  the 
following  reasons,  we  remain  convinced  that 
there  is  good  reason  to  believe  that  the  issue 
of  sexing  does  not  overshadow  the  validity  of 
this  taxon.  It  is  our  experience  that  mis-sexing 


54 


FIELDIANA:  ZOOLOGY 


o 


L  camiguinensis 
L  p.  apicalis 
L  p.  worcesteh 
L  p.  regulus 

Fig.  2.     Results  of  the  Principal  Components  Analysis  of  morphological  data  from  populations  of  Philippine 
Loriculus,  on  the  first  three  axes;  see  text  for  details. 


O 

o 

A 


most  commonly  involves  undeveloped  gonads, 
and  there  are  clearly  labeled  tags  indicating 
that  the  gonads  of  some  specimens  were  de- 
veloped. If  mistakes  were  made,  the  plumage 
characters  of  the  specimens  described  above 
would  logically  argue  that  all  males  were 
misidentified  as  females.  Again,  this  seems  highly 
unlikely  given  the  number  of  males  (19). 
Furthermore,  it  seems  highly  unlikely  that 
a  series  of  23  randomly  collected  parrots  would 
all  have  been  females.  Certainly,  there  is  nothing 
like  this  in  other  series  of  Loriculus  collected  by 
Rabor  in  other  parts  of  the  Philippines.  Thus, 
while  we  cannot  say  that  all  specimens  are 
unequivocally  identified  to  sex  correctly,  we  feel 
that  adults  of  both  males  and  females  are 
included  in  these  series. 


Discussion 

Our  results  demonstrate  that  L.  camiguinensis 
is  diagnosable  from  populations  of  L.  philippen- 
sis  in  plumage.  It  also  differs  in  size  from  all 


neighboring  populations.  We  were  able  to  make 
direct  comparisons  with  specimens  of  all  sub- 
species except  L.  p.  dohertyi  (Basilan).  It  is 
possible  that  L.  camiguinensis  is  more  closely 
related  to  some  parts  of  the  L.  philippensis  group 
than  others,  which  would  make  L.  philippensis 
paraphyletic  (Funk  &  Omland  2003).  Despite 
this  possible  relationship  to  the  widespread  L. 
philippensis  group,  we  believe  L.  camiguinensis 
sufficiently  differentiated  to  be  beyond  concerns 
expressed  about  recognizing  new  species  based 
on  minor  morphologic  differences  (e.g.,  Collar  et 
al.,  1999).  Based  on  geographic  distance  and  the 
overall  pattern  of  plumage  coloration,  L.  cami- 
guinensis most  closely  resembles  populations  of 
L.  p.  apicalis  and  L.  p.  worcesteri,  but  no 
phylogenetic  analyses  exist  yet  for  these  taxa. 
The  comparatively  dull  plumage  of  the  male 
of  L.  camiguinensis  is  consistent  with  the 
documented  tendency  for  some  insular  bird 
populations  to  lose  bright  plumage,  leading  to 
a  lack  of  sexual  dichromatism  (see  references  in 
Peterson,  1996);  L.  camiguinensis  is  the  only 
member  of  the  genus  without  sexual  dichroma- 
tism in  plumage. 


TELLO  ET  AL.:  A  NEW  SPECIES  OF  HANGING-PARROT 


The  recognition  of  this  distinctive  taxon 
coincides  with  recent  surveys  of  the  small 
mammal  fauna  of  Camiguin  Island  that  have 
discovered  two  new  species  of  rodents  (Rickart  et 
al.,  2002;  Heaney  et  al.,  2006).  Camiguin  is 
believed  to  be  the  smallest  Philippine  island  to 
harbor  endemic  species  of  birds  and  mammals 
(Heaney  &  Tabaranza,  1997,  2006a).  The  island 
has  been  continuously  isolated  from  its  large 
southern  neighbor,  Mindanao,  even  during 
periods  of  low  sea  level  during  the  "ice  ages" 
of  the  Pleistocene,  when  sea  levels  dropped  to 
120  m  below  present  levels  (Heaney,  1986, 
1991a,  1991b;  Fairbanks,  1989;  Heaney  and 
Tabaranza,  1997,  2005b;  Heaney  and  Regalado, 
1998;  Hanebuth  et  al.,  2000),  and  this  may  have 
played  a  role  in  the  differentiation  of  Camiguin's 
fauna  from  that  of  Mindanao  (Steppan  et  al., 
2003). 

The  value  of  museum  collections  is  well 
illustrated  with  this  description.  These  collec- 
tions were  essential  in  the  recognition  and 
documentation  of  L.  camiguinensis.  Had  there 
not  been  a  series  of  specimens  available  for 
study,  we  would  have  likely  dismissed  differences 
in  the  new  taxon  as  possibly  aberrant  or 
immature  plumage  or  an  error  in  sexing  of 
a  specimen  (a  female  incorrectly  identified  as 
a  male;  see  above).  However,  the  presence  of 
a  series  of  specimens  from  different  localities 
(with  data  on  gonadal  development)  has  allowed 
us  to  compile  meaningful  data  sets  on  morpho- 
logical variation  and  assess  within-population 
variation  in  color. 

This  new  species  also  illustrates  the  need  for 
additional  taxonomic  and  systematic  research  on 
the  Loriculus  hanging-parrots  to  understand  the 
evolutionary  patterns  in  the  group  and  to 
evaluate  the  possibility  that  some  of  the  other 
allopatric  forms  of  L  philippensis  may  also 
deserve  species  status.  The  issue  of  assessing  the 
taxonomic  status  of  allopatric  populations  in  the 
Philippines  has  long  been  recognized  as  a  chal- 
lenge for  conservationists  (Collar  et  al.,  1999; 
Peterson  et  al.,  2000).  To  date,  little  attention  has 
been  given  to  the  conservation  plight  of  Loriculus 
parrots.  For  instance,  none  was  included  by 
Collar  et  al.  (1999)  in  their  list  of  threatened 
Philippine  bird  species.  This  lack  of  attention  can 
be  directly  correlated  with  the  designation  of  L. 
philippensis  as  a  polytypic  species.  The  plight  of 
these  populations  is  cause  for  concern,  as 
Loriculus  p.  chrysonotus  from  the  island  of  Cebu 
is  believed  to  be  extinct  (Forshaw,  1989;  Mallari 


et  al..  2001)  and  another  form,  L.  p.  siquijorensis, 
may  be  extinct  as  well  (Forshaw,  1989;  Kennedy 
et  al.,  2000).  The  combined  populations  of 
Mindoro,  Sibuyan,  Negros,  Panay,  Tablas, 
Romblon,  Masbate,  Ticao,  Guimaras,  and  Basi- 
lan  (including  mindorensis,  bournsi,  regulus,  and 
dohertyi)  may  total  no  more  than  5000  individ- 
uals (Juniper  &  Parr,  1998).  The  current  popu- 
lation size  of  L.  camiguinensis  is  not  known  (but 
see  Heaney  and  Tabaranza,  2006a,  for  an 
assessment  of  remaining  habitat  on  the  island). 
Without  field  data  on  its  status,  we  defer  from 
suggesting  how  this  species  should  be  character- 
ized under  international  threat  criteria  (IUCN 
Species  Survival  Commission,  1994).  However, 
because  Camiguin  is  a  small  island  that  has 
experienced  extensive  deforestation,  the  conser- 
vation status  of  this  newly  described  species 
clearly  requires  assessment.  Field  study  is  needed 
to  establish  the  population  size  and  requirements 
as  a  prerequisite  for  conservation  planning  and 
action. 


Acknowledgments 

We  are  grateful  to  G.  Hess  of  the  Delaware 
Museum  of  Natural  History  for  loan  of  speci- 
mens under  his  care  and  to  M.  Skakuj  for 
painting  the  plate.  Bruce  Patterson  and  P. 
Wagner  provided  help  with  data  analyses,  and 
S.  Bober  helped  in  preparing  Figure  1.  The 
manuscript  was  greatly  improved  by  comments 
from  D.  S.  Balete,  A.  Carnaval,  L.  R.  Heaney,  S. 
Ware,  P.  C.  Rasmussen,  and  N.  J.  Collar, 
although  this  does  not  imply  that  all  agree  with 
the  status  we  convey  on  this  taxon. 


Literature  Cited 

Balete,  D.  S.,  B.  R.  Tabaranza,  Jr.,  and  L.  R. 
Heaney.  2006.  An  annotated  checklist  of  the 
land  birds  of  Camiguin  Island,  Philippines,  pp. 
58-72.  In  Heaney,  L.  R.,  ed.,  The  Mammals  and 
Birds  of  Camiguin  Island,  Philippines,  a  Distinctive 
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TELLO  ET  AL.:  A  NEW  SPECIES  OF  HANGING-PARROT 


57 


An  Annotated  Checklist  of  the  Birds  of  Camiguin 
Island,  Philippines 

Danilo  S.  Balete,1'3  Bias  R.  Tabaranza,  Jr.,2  and  Lawrence  R.  Heaney3 


Abstract 

Fifty-five  species  of  resident  breeding  and  two  species  of  migratory  land  birds  have  been 
documented  on  Camiguin  Island,  including  one  bittern,  one  eagle,  one  junglefowl,  two  rails, 
eight  doves  and  pigeons,  one  parrot,  three  cuckoos,  one  owl,  three  swiftlets,  one  dollarbird, 
two  kingfishers,  one  bee-eater,  one  hornbill,  one  pitta,  one  triller,  two  bulbuls,  one  crow,  four 
thrushes,  three  warblers,  six  flycatchers,  one  pipit,  one  wood  swallow,  two  starlings,  three 
sunbirds,  two  flowerpeckers,  two  white-eyes,  and  two  munias.  At  least  seven  species  reported 
here  are  first  records  for  Camiguin.  Ten  species  are  widespread  Philippine  endemics,  two  are 
near-endemics,  and  one  (Loriculus  camiguinensis,  described  in  this  volume)  is  endemic  to 
Camiguin.  Additionally,  four  endemic  subspecies  are  recognized  from  Camiguin,  Ixos  everetti 
catarmanensis,  Hypothymis  azurea  catarmanensis,  Diceum  trigonostigma  isidroi,  and  Zosterops 
nigrorum  catarmanensis.  While  this  list  is  not  comprehensive,  the  presence  of  57  species 
demonstrates  that  many  species  were  able  to  cross  a  narrow  but  permanent  sea  channel,  and 
the  presence  of  four  endemic  subspecies  and  one  endemic  species  indicates  that  some  genetic 
differentiation  has  resulted. 


Introduction 

Although  the  avifaunas  of  many  of  the 
Philippine  islands  have  been  reported  in  com- 
prehensive fashion  (summarized  by  Dickinson  et 
al.,  1991),  no  such  report  has  been  made  for 
Camiguin  Island,  with  the  result  that  Camiguin 
has  not  been  included  in  most  analyses  of  avian 
biogeography  and  conservation  priorities  (e.g., 
Hauge  et  al.,  1986;  Dickinson  et  al.,  1991;  Collar 


1  Laksambuhay  Conservation  Inc.,  10241  Mt.  Bu- 
lusan  Street,  U-2.  Los  Banos,  Laguna,  Philippines. 

2  Department  of  Biology,  Uigan  Institute  of  Tech- 
nology, Mindanao  State  University,  Iligan  City, 
Lanao  del  Norte,  Philippines. 

3  Field  Museum  of  Natural  History,  1400  South 
Lake  Shore  Drive,  Chicago.  IL  60605-2496,  U.S.A. 


et  al.,  1999;  Peterson  et  al.,  2000;  but  see  Mallari 
et  al.,  2001,  Ong  et  al.,  2002).  We  note  that 
Camiguin  Norte,  which  lies  north  of  Luzon,  is 
often  confused  with  Camiguin.  As  documented 
in  the  following  list,  the  land  birds  of  Camiguin 
Island  include  a  diverse  assemblage  of  at  least  57 
species.  These  records  are  based  on  voucher 
specimens  that  were  obtained  principally  during 
the  1960s  and  1990s  and  are  deposited  at  the 
Delaware  Museum  of  Natural  History  (dmnh), 
the  Field  Museum  of  Natural  History  (fmnh), 
and  Mindanao  State  University-Iligan  Institute 
of  Technology  (msu-iit),  as  detailed  below  and  in 
Chapter  1  (Heaney  &  Tabaranza,  2006a).  Addi- 
tional records  based  on  specimens  at  the 
Museum  of  Comparative  Zoology  (mcz),  Cam- 
bridge, as  cited  in  Dickinson  et  al.  (1991),  are 
included  as  well. 


58 


FIELDIANA:  ZOOLOGY,  N.S.,  NO.  106,  APRIL  5,  2006,  PP.  58-72 


The  presence  of  55  species  of  breeding  land 
birds  on  an  island  the  size  of  Camiguin  in  the 
Philippines  is  not  remarkable;  indeed,  as  docu- 
mented below,  surveys  of  birds  on  Camiguin 
were  not  exhaustive,  and  the  current  total  is  well 
below  the  total  number  likely  to  be  present. 
However,  given  that  Camiguin  is  an  oceanic 
island  surrounded  by  deep  water  but  only  about 
10  km  from  the  shore  of  Mindanao,  these  data 
are  a  useful  indicator  of  the  ability  of  these  55 
species  to  readily  cross  a  sea  channel  of  this 
breadth,  given  an  extensive  period  of  time. 
Further,  as  noted  below,  the  avifauna  of 
Camiguin  is  now  considered  to  include  four 
endemic  subspecies  of  birds  (Ixos  everetti  catar- 
manensis,  Hypothymis  azurea  catarmanensis, 
Diceum  trigono stigma  isidroi,  and  Zosterops 
nigrorum  catarmanensis)  and  an  endemic  species 
of  parrot  {Loriculus  camiguinensis;  Tello  et  al., 
2006).  This  observation  indicates  that  genetic 
differentiation  has  taken  place  in  at  least  some  of 
these  species,  leading  in  one  case  to  significant 
change.  It  is  our  hope  that  this  paper  will 
encourage  more  extensive  and  focused  analysis 
of  the  avifauna  of  Camiguin,  on  both  ecology 
and  evolution. 

For  notes  on  methods  of  collecting  and  for 
a  list  and  description  of  the  collecting  sites,  see 
Heaney  and  Tabaranza  (2006a).  As  noted  in  that 
chapter,  elevations  from  the  1960s  should  be 
viewed  as  rough  estimates.  Species  whose  names 
are  listed  in  brackets  are  of  uncertain  documen- 
tation and  are  not  included  in  our  tally  of  species 
numbers.  Unverified  sight  records  are  listed  at 
the  end  of  this  paper.  We  define  "first  records" 
as  those  not  reported  by  Dickinson  et  al.  (1991) 
and/or  Kennedy  et  al.  (2000). 


Accounts  of  Species 

Family  Ardeidae — Bitterns,  Egrets,  and  Herons 

Ixobrychus  cinnamomeus — Cinnamon  Bittern 

The  Cinnamon  Bittern  ranges  from  India  to 
China,  Ryukyus,  Taiwan,  and  Southeast  Asia;  in 
the  Philippines  it  occurs  on  most  islands,  except 
the  Babuyan  and  Batanes  group,  in  marshes,  rice 
fields,  and  wetlands  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  A  single  specimen  was 
taken  in  a  highly  disturbed  agricultural  area  in 
Balbagon,  Mambajao,  on  28  May  1992,  at  10  m, 
the  first  record  of  this  species  from  the  island 
(Site  1). 


Specimens  Examined — Total  1.  Site  1  (1 
fmnh). 

Family  Accipitridae — Hawks  and  Eagles 

Spilornis  cheela — Crested  Serpent-Eagle 

The  Crested  Serpent-Eagle  occurs  from  India 
to  China,  Taiwan,  Ryukyus,  and  Southeast  Asia; 
it  is  found  throughout  the  Philippines,  except  the 
Babuyan  and  Batanes  groups  of  islands,  in  forest 
and  forest  edge  (Dickinson  et  al.,  1991;  Kennedy 
et  al.,  2000).  Two  male  specimens  were  taken  on 
13  June  1968  on  Mt.  Catarman,  one  at  2000  ft 
(ca.  600  m;  Site  9),  the  other  at  an  unknown 
elevation.  We  often  saw  them  at  nearly  all 
elevations  during  fieldwork  in  1994  and  1995, 
especially  near  forest. 

Specimens  Examined — Total  2.  Site  9  (1  dmnh; 
1  fmnh). 

Family  Megapodidae — Megapodes  and  Scrubfowl 

[Megapodius  cumingii — Tabon] 

The  Tabon  ranges  from  Sulawesi  and  Borneo 
to  the  Philippines;  once  widespread  throughout 
the  country,  it  is  now  found  mainly  in  coastal 
scrub  but  also  occurs  in  lowland  to  montane 
forest  (Dickinson  et  al.,  1991;  Kennedy  et  al., 
2000).  In  an  earlier  enumeration  of  Philippine 
birds,  Camiguin  was  not  identified  as  part  of  the 
Tabon's  range  (Dickinson  et  al.,  1991).  However, 
in  the  more  recent  compilation,  Kennedy  et  al. 
(2000)  listed  Camiguin  as  part  of  the  range  of 
this  species  without  further  reference  to  the 
source  of  their  data.  Thus,  we  have  not  included 
M.  cumingii  in  our  tally  of  birds  from  Camiguin 
but  note  its  potential  presence. 

Family  Phasianidae — Partridges,  Pheasants, 
and  Quail 

Gallus  gallus — Red  Junglefowl 

The  Red  Junglefowl  occurs  from  India  to 
southern  China  and  Southeast  Asia,  including 
virtually  all  of  the  Philippines,  except  the 
Batanes  group  of  islands,  in  forest  and  forest 
edge  up  to  2000  m  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  A  male  specimen  was 
taken  on  Mt.  Timpoong  at  3150  ft  (ca.  950  m) 
on  14  June  1969  (Site  13).  In  May  1994,  a  male 
specimen  was  taken  in  a  Victor  trap  in  Kital-is, 
Sagay  at  1200-1400  m  (Site  6).  We  also  heard 
them  in  March  1995  crowing  close  to  our 
campsite  on  Mt.  Timpoong  at  1275  m  (Site  7). 

Specimens  Examined — Total  2.  Site  6  (1  msu- 
iit);  Site  13  (1  dmnh). 


BALETE  ET  AL.:  ANNOTATED  CHECKLIST  OF  THE  BIRDS  OF  CAMIGUIN  ISLAND     59 


Family  Rallidae — Coots,  Crakes,  Rails, 

and  Waterhens 

Porzana  fusca-    Ruddy-breasted  Crake 

The  Ruddy-breasted  Crake  ranges  from  India 
to  Japan,  Ryukyus,  and  Southeast  Asia;  in  the 
Philippines  it  has  been  recorded  only  on  the 
islands  of  Bohol,  Cagayancillo,  Leyte,  Luzon, 
Mindanao,  Mindoro,  Negros,  Panay,  Samar, 
and  Sibuyan,  usually  in  marshes  and  rice  fields 
but  also  along  forest  paths  up  to  1500  m 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
A  male  specimen  was  taken  in  Kasangsangan  in 
the  vicinity  of  Mt.  Catarman  at  1000  ft  (ca. 
300  m)  on  22  June  1968  (Site  11).  This  is  the  first 
record  of  this  species  on  Camiguin. 

Specimens    Examined — Total    1.    Site    11    (1 

FMNH). 

Rallina  eurizonoides — Slaty-legged  Crake 

The  Slaty-legged  Crake  occurs  from  India  and 
Sri  Lanka  to  Taiwan,  Ryukyus,  and  Southeast 
Asia.  In  the  Philippines  it  is  rather  uncommon  on 
most  islands  but  absent  on  the  Palawan  and  Sulu 
groups  of  islands,  in  scrub  and  primary  to 
secondary  forest  up  to  900  m  (Dickinson  et  al., 
1991;  Kennedy  et  al.,  2000).  The  record  on 
Camiguin,  a  first  for  this  island,  consists  of  two 
specimens,  both  taken  in  Victor  traps,  from 
Kital-is,  Sagay,  at  900-1100  m  on  19  May  1994 
(Site  4)  and  at  1200-1400  m  on  26  May  1994 
(Site  6).  The  following  year,  another  was  taken 
on  Mt.  Timpoong  in  primary  montane  forest  at 
1275  m  on  23  March  1995  (Site  7). 

Specimens  Examined — Total  3.  Site  4  (1  msu- 
iit);  Site  6  (1  msu-iit);  Site  7  (1  fmnh). 

Family  Columbidae — Doves  and  Pigeons 
Treron  vernans — Pink-necked  Green-Pigeon 

The  Pink-necked  Green-Pigeon  is  widespread 
in  Southeast  Asia;  it  has  been  recorded  through 
most  of  the  Philippines,  except  the  Babuyan  and 
Batanes  groups  of  islands,  principally  in  lowland 
second-growth  forest  below  300  m  (Dickinson, 
1991;  Kennedy  et  al.,  2000).  An  adult  male 
specimen  was  taken  on  29  June  1969  in  Puntod, 
Mahinog,  at  800  ft  (ca.  250  m). 

Specimens  Examined — Total  1.  Site  14  (1 
dmnii). 

Phapitreron  leucotis — White-eared  Brown-Dove 
The  White-eared  Brown-Dove  is  endemic  to 
the  Philippines;  it  occurs  throughout  the  country 
with  the  exception  of  the  Babuyan,  Batanes,  and 
Palawan    groups    of  islands,    in    primary    and 


secondary  forest  up  to  about  1600  m  (Dickinson 
et  al.,  1991;  Kennedy  et  al.,  2000).  It  was 
recorded  from  Mt.  Catarman  and  the  nearby 
area  of  Kasangsangan,  at  1 000-4500  ft  (ca.  300- 
1400  m)  in  June  1968  (Sites  9  and  11).  Further 
records  were  obtained  from  Mt.  Timpoong  at 
3150-4000  ft  (ca.  950-1200  m)  and  the  nearby 
area  of  Puntod  at  800  ft  (ca.  250  m)  in  June  1969 
(Sites  13  and  14,  respectively).  We  also  recorded 
it  in  May  1994  in  Kital-is,  Sagay,  at  900-1100  m 
(Site  4). 

Specimens  Examined — Total  12.  Site  4  (1  msu- 
iit);  Site  9  (3  dmnh,  1  fmnh);  Site  1 1  (3  fmnh); 
Site  13  (2  fmnh);  Site  14  (2  fmnh). 

Ptilinopus  occipitalis — Yellow-breasted  Fruit- 
Dove 

The  Yellow-breasted  Fruit-Dove  is  endemic  to 
the  Philippines;  it  is  fairly  widespread  through- 
out the  country,  except  the  Babuyan,  Batanes, 
Palawan,  and  Sulu  groups  of  islands,  in  forest  up 
to  1800  m  (Dickinson  et  al,  1991;  Kennedy  et 
al.,  2000).  It  was  recorded  on  Mt.  Catarman  and 
the  nearby  areas  of  Gidag-on  and  Kasangsangan 
at  1000^1500  ft  (ca.  300-1400  m)  in  June  1968 
(Sites  9-11).  Additional  records  came  from  Mt. 
Timpoong  at  3150^4800  ft  (ca.  950-1450  m)  in 
June  1969  (Sites  12  and  13). 

Specimens  Examined — Total  18.  Site  9  (3 
dmnh,  3  fmnh);  Site  10  (1  dmnh,  1  fmnh);  Site 
1 1  (1  fmnh);  Site  12  (1  fmnh);  Site  13  (4  dmnh,  4 
fmnh). 

Columba  vitiensis — Metallic  Pigeon 

The  Metallic  Pigeon  occurs  on  Borneo, 
Sulawesi,  Moluccas,  Lesser  Sunda  Islands,  New 
Guinea,  and  Samoa;  it  is  an  uncommon  resident 
of  most  islands  throughout  the  Philippines,  in 
lowland  to  mossy  forest  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  Records  from  Camiguin 
were  all  taken  in  1968.  A  female  specimen  was 
taken  from  Mt.  Catarman  at  2000  ft  (ca.  600  m) 
on  14  June  (Site  9)  and  three  adult  males  and  an 
adult  female  from  Kasangsangan  at  1000- 
2000  ft  (ca.  300-600  m)  in  the  vicinity  of  Mt. 
Catarman  on  11-18  June  (Site  11). 

Specimens  Examined — Total  5.  Site  9  (1 
fmnh);  Site  11  (3  dmnh,  1  fmnh). 

Macropygia  phasianella — Reddish  Cuckoo-Dove 
The  Reddish  Cuckoo-Dove  ranges  from  Bor- 
neo, the  Moluccas,  Sulawesi,  and  the  Lesser 
Sunda  islands  to  New  Guinea  and  Samoa;  it 
occurs  throughout  the  Philippines,  in  lowland  to 
mossy  forest  above  2000  m  (Dickinson  et  al., 


60 


FIELDIANA:  ZOOLOGY 


1991;  Kennedy  et  al.,  2000).  It  was  recorded  on 
Mt.  Catarman  and  the  nearby  area  of  Kasang- 
sangan  at  1000-2500  ft  (ca.  300-750  m)  in  June 
1968  (Sites  9  and  11).  Additional  records  came 
from  Mt.  Timpoong  at  3150-5700  ft  (ca.  950- 
1700  m)  and  the  nearby  area  of  Puntod  at  800  ft 
(ca.  250)  in  June  1969  (Sites  12-14).  In  May 
1994,  we  recorded  it  in  Kital-is,  Sagay,  at  1000- 
1400  m  (Sites  5  and  6). 

Specimens  Examined — Total  30.  Site  5  (1  msu- 
iit);  Site  6  (2  msu-iit);  Site  9  (1  fmnh);  Site  1 1  (3 
dmnh,  3  fmnh);  Site  12  (9  dmnh,  7  fmnh);  Site  13 
(1  dmnh,  2  fmnh);  Site  14  (1  fmnh). 

Streptopelia  bitorquata — Island  Collared-Dove 

The  Island  Collared-Dove  is  found  from 
Borneo  and  the  Lesser  Sunda  islands  to  Java; 
in  the  Philippines  it  is  a  fairly  uncommon 
resident  of  most  islands,  except  the  Batanes 
group  of  islands,  mainly  in  relatively  open  fields 
in  the  lowlands  and  sometimes  in  mangrove 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
Two  male  specimens  were  collected  from  Pun- 
tod, Mahinog,  at  700-800  ft  (ca.  200-250  m), 
one  of  which  was  a  juvenile  taken  at  800  ft  (ca. 
250  m)  on  28  June  1968  (Site  14). 

Specimens  Examined — Total  2.  Site  14  (1 
dmnh,  1  fmnh). 

Geopelia  striata — Zebra  Dove 

The  Zebra  Dove  ranges  from  Southeast  Asia 
to  Australia;  it  occurs  throughout  the  Philip- 
pines, except  the  Babuyan  and  Batanes  groups  of 
islands,  in  open  country  and  cultivated  areas  in 
the  lowlands  (Dickinson  et  al.,  1991;  Kennedy  et 
al.,  2000).  One  of  us  (B.R.T.)  netted  one  some 
50  m  away  from  the  beach  in  Manuyog,  Sagay, 
on  10  May  1994  (Site  3),  but  it  was  released.  No 
other  specimen  was  recorded  of  this  species  in 
1994.  It  was  not  recorded  during  our  fieldwork  in 
1992  and  1995. 

Specimens  Examined — None. 

Chalcophaps  indica — Common  Emerald-Dove 

The  Common  Emerald-Dove  is  found  from 
India  and  Sri  Lanka  to  China,  Taiwan,  Ryukyus 
and  Southeast  Asia,  New  Guinea,  and  Australia; 
it  occurs  throughout  the  Philippines,  often  in 
second-growth  forest  in  the  lowlands  up  to 
1000  m  (Dickinson  et  al.,  1991;  Kennedy  et  al., 
2000).  Records  on  Camiguin  consist  of  speci- 
mens taken  in  Kasangsangan,  near  Mt.  Catar- 
man, at  1000-2500  ft  (ca.  300-750  m)  in  June 
1968  (Site  11)  and  from  Puntod  at  800  ft  (ca. 
250  m)  in  May  1969  (Site  14). 


Specimens  Examined — Total  5.  Site  11  (2 
dmnh,  2  fmnh);  Site  14  (1  fmnh). 

Family  Psittacidae — Lorikeets,  Cockatoos,  Parrots, 
and  Racquet-tails 

Loriculus  camiguinensis — Camiguin  Hanging- 
Parrot 

Described  as  a  new  species  in  this  volume 
(Tello  et  al.,  2006),  this  is  the  only  species  of  bird 
currently  recognized  as  endemic  to  Camiguin.  Its 
elevational  range,  based  on  specimens  collected 
in  the  1960s,  is  300  to  1350,  thus  including 
lowland  and  montane  rain  forest.  Most  of  this 
habitat  has  been  destroyed  on  Camiguin,  leading 
to  concern  for  its  conservation  status. 

Specimens  Examined — Total  23.  Site  9  (2 
dmnh);  Site  11  (5  dmnh,  5  fmnh);  Site  13  (5 
dmnh,  6  fmnh). 

Family  Cuculidae — Cuckoos,  Malkohas,  and  Coucals 

Cacomantis  variolosus — Brush  Cuckoo 

The  Brush  Cuckoo  occurs  from  Southeast 
Asia  to  Australia  and  the  Southwest  Pacific;  it  is 
found  throughout  the  Philippines,  except  the 
Babuyan  and  Batanes  groups  of  islands,  from 
mangrove  to  mossy  forest  at  2000  m  (Dickinson 
et  al.,  1991;  Kennedy  et  al.,  2000).  The  records 
from  Camiguin  were  taken  from  Mt.  Catarman 
at  2500  ft  (ca.  750  m)  in  June  1968  (Site  9)  and 
from  Mt.  Timpoong  at  3150^1800  ft  (ca.  950- 
1450  m)  in  June  1969  (Site  13).  We  also  recorded 
it  in  Kital-is,  Sagay,  at  900-1100  (Site  4)  and 
1200-1400  m  in  May  1994  (Site  6). 

Specimens  Examined — Total  7.  Site  6  (1  msu- 
iit);  Site  9  (1  fmnh);  Site  13  (3  dmnh,  2  fmnh). 

Eudynamys  scolopacea — Common  Koel 

The  Common  Koel  occurs  from  India  through 
Southeast  Asia  to  northern  Australia;  it  is  found 
throughout  the  Philippines,  in  primary  and 
secondary  lowland  forest  (Dickinson  et  al., 
1991;  Kennedy  et  al.,  2000).  The  single  specimen 
from  Camiguin  was  taken  on  Mt.  Timpoong  at 
3150  ft  (ca.  950  m)  on  17  June  1969  (Site  13). 

Specimens  Examined — Total  1.  Site  13  (1 
fmnh). 

Centropus  viridis — Philippine  Coucal 

The  Philippine  Coucal  is  endemic  to  the 
Philippines,  where  it  occurs  throughout  the 
country,  except  the  Palawan  group  of  islands, 
from  grassland  to  forest  at  2000  m  (Dickinson  et 
al.,  1991;  Kennedy  et  al.,  2000).  The  records  on 
Camiguin    consist    of   specimens    taken    from 


BALETE  ET  AL.:  ANNOTATED  CHECKLIST  OF  THE  BIRDS  OF  CAMIGUIN  ISLAND     61 


Kasangsangan  in  the  vicinity  of  Mt.  Catarman  at 
1000  1500  it  (ca.  300  450  m)  in  June  1968  (Site 
11)  and  from  Mt.  Timpoong  at  3150  ft  (ca. 
950  m)  in  June  1969  (Site  13). 

Specimens  Examined— Total  6.  Site  11  (1 
dmnii.  2  fmnh);  Site  13(1  dmnh,  2  fmnh). 

Family  Strigidae — Owls 

Ninox  philippensis     Philippine  Hawk-Owl 

The  Philippine  Hawk-Owl  is  endemic  to  the 
Philippines,  occurring  throughout  most  of  the 
islands,  except  the  Babuyan.  Batanes,  and 
Palawan  groups,  in  the  lowlands  up  to  1 800  m 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
The  records  from  Camiguin  consist  of  a  female 
taken  from  Mt.  Catarman  at  1500  ft  (ca.  450  m) 
on  17  June  1968  (Site  9)  and  another  female  from 
Mt.  Timpoong  on  13  June  1969  at  an  unknown 
elevation.  Specimens  of  other  species  from  Mt. 
Timpoong  on  the  same  date  were  taken  at 
3150  ft  (ca.  950  m). 

Specimens  Examined — Total  2.  Site  9  (1 
fmnh);  Site  13(1  fmnh). 

Family  Apodidae — Swifts 

Collocalia  mearnsi — Philippine  Swiftlet 

The  Philippine  Swiftlet  is  endemic  to  the 
Philippines,  where  it  is  recorded  mainly  on  the 
islands  of  Bohol,  Cebu,  Luzon,  Mindanao, 
Mindoro,  Negros,  Palawan,  and  Panay,  usually 
in  forest  and  forest  clearings  above  900  m;  it  is 
absent  from  the  Babuyan,  Batanes,  and  Sulu 
groups  of  islands  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  Specimens  from  Camiguin 
were  taken  from  Mt.  Catarman  and  in  the 
nearby  Kasangsangan  at  1000-2000  ft  (ca.  300- 
600  m)  on  16  June  1968  (Sites  9  and  11)  and 
from  Mt.  Timpoong  on  23  June  1969  from  an 
unknown  elevation.  Specimens  of  other  species 
from  this  site  were  taken  at  3100-3350  ft  (ca. 
950-1000  m),  with  several  at  4000  ft  (ca. 
1200  m). 

Specimens  Examined — Total  4.  Site  9  (1 
fmnh);  Site  11  (1  fmnh);  Site  13  (2  fmnh). 

Collocalia  esculenta — Glossy  Swiftlet 

The  Glossy  Swiftlet  occurs  from  the  Anda- 
mans,  Nicobars.  and  Malay  Peninsula  to  New 
Guinea  and  the  southwest  Pacific;  it  is  found 
throughout  the  Philippines,  from  sea  level  to 
mountaintops  (Dickinson  et  al.,  1991;  Kennedy 
et  al.,  2000).  It  was  commonly  recorded  on  Mt. 
Catarman  and  in  the  nearby  area  of  Kasangsan- 


gan at  1000-2000  ft  (ca.  300-600  m)  in  June 
1968  (Sites  9  and  11)  and  on  Mt.  Timpoong  at 
3150-4800  ft  (ca.  950-1450  m)  in  June  1969 
(Sites  12  and  13). 

Specimens  Examined — Total  31.  Site  9  (1 
fmnh);  Site  11  (2  dmnh,  7  fmnh);  Site  12  (2 
dmnh,  1  fmnh);  Site  13  (9  dmnh,  9  fmnh). 

Collocalia  troglodytes— Pygmy  Swiftlet 

The  Pygmy  Swiftlet  is  endemic  to  the  Philip- 
pines, where  it  occurs  throughout  most  of  the 
country,  except  the  Babuyan,  Batanes,  and  Sulu 
groups  of  islands,  usually  at  low  to  middle 
elevations  in  forested  areas  (Dickinson  et  al., 
1991;  Kennedy  et  al.,  2000).  The  records  from 
Camiguin  were  taken  on  13  and  21  June  1968  in 
Kasansangan  at  1500-2000  ft  (ca.  450-600  m) 
and  on  Mt.  Timpoong  at  3150  ft  (ca.  950  m)  on 
25  June  1969  (Site  13). 

Specimens  Examined — Total  4.  Site  11  (3 
dmnh);  Site  13  (1  fmnh). 

Family  Coraciidae — Rollers 

Eurystomus  orientalis — Dollarbird 

The  Dollarbird  occurs  from  India  to  New 
Guinea  and  the  southwest  Pacific,  including  all 
of  the  Philippines,  except  the  Batanes  group  of 
islands,  in  forest  edge  and  clearings  in  the 
lowlands  up  to  1200  m  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  Dickinson  et  al.  (1991) 
noted  a  single  specimen  deposited  in  the  Museum 
of  Comparative  Zoology,  Harvard  University, 
taken  from  Mambajao  on  19  August  1921;  no 
elevation  was  indicated. 

Specimens  Examined — None. 

Family  Alcedinidae — Kingfishers 

Ceyx  lepidus — Variable  Dwarf-Kingfisher 

The  Variable  Dwarf-Kingfisher  occurs  from 
the  Moluccas  to  New  Guinea  and  the  southwest 
Pacific  islands;  in  the  Philippines  it  is  recorded 
on  the  islands  in  the  Central  Philippines  as  well 
as  on  the  Mindanao  and  Sulu  groups  of  islands, 
in  primary  and  secondary  lowland  forest  (Dick- 
inson et  al,  1991;  Kennedy  et  al.,  2000).  It  was 
recorded  from  Mt.  Timpoong  at  3150  ft  (ca. 
950  m)  on  17  and  18  June  1969  (Site  13).  In  May 
1994,  two  individuals  were  recorded  in  Kital-is, 
Sagay,  at  900-1 100  m  (Site  4),  including  one  that 
was  taken  in  a  Victor  trap  on  17  May. 

Specimens  Examined — Total  4.  Site  4  (2  msu- 
iit);  Site  13  (1  dmnh,  1  fmnh). 


62 


FIELDIANA:  ZOOLOGY 


Halcyon  Moris — White-collared  Kingfisher 

The  White-collared  Kingfisher  occurs  widely 
from  northeast  Africa  to  southern  China,  Ryu- 
kyus,  Southeast  Asia  to  New  Guinea,  Australia, 
and  the  southwest  Pacific;  it  is  found  nearly  all 
over  the  Philippines,  from  exposed  reefs  to  open 
country  and  forest  edge  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  It  was  recorded  in  Gidag- 
on  and  Kasangsangan,  both  in  the  vicinity  of  Mt. 
Catarman,  at  500-2500  ft  (ca.  150-600  m;  Sites 
10  and  1 1)  in  June  1968  and  on  Mt.  Timpoong  at 
3150  ft  (ca.  950  m;  Site  13),  in  Puntod  at  800  ft 
(ca.  250  m;  Site  14),  and  Mantigue  Island  (Site 
19)  in  June  1969.  We  recorded  it  further  in 
Balbagon,  Mambajao,  at  10  m  in  May  1992  (Site 
1).  On  10  May  1994,  we  also  recorded  it  in 
Manuyog,  Sagay,  at  80  m  (Site  3). 

Specimens  Examined — Total  33.  Site  1  (1 
fmnh);  Site  10  (6  dmnh,  4  fmnh);  Site  11  (1 
dmnh,  1  fmnh);  Site  13  (5  dmnh,  5  fmnh);  Site  14 
(4  dmnh,  5  fmnh);  Site  19  (1  dmnh). 

Family  Meropidae — Bee-Eaters 

Merops  viridis — Blue-throated  Bee-eater 

The  Blue-throated  Bee-eater  occurs  in  South- 
east Asia  and  the  Philippines,  where  it  is  known 
from  many  islands,  except  the  Babuyan,  Batanes, 
Palawan,  and  Sulu  groups;  it  often  found  in  open 
country,  scrubs,  and  forest  clearings  (Dickinson 
et  al.,  1991;  Kennedy  et  al.,  2000).  On  Camiguin, 
it  was  recorded  from  Mt.  Catarman  and  in 
nearby  Kasangsangan  at  1000-4500  m  (ca.  300- 
1400  m)  in  June  1968  (Sites  9  and  1 1)  and  on  Mt. 
Timpoong  at  3350  ft  (ca.  1000  m)  in  June  1969 
(Site  13). 

Specimens  Examined — Total  9.  Site  9  (1  dmnh, 
2  fmnh);  Site  11  (2  dmnh,  2  fmnh);  Site  13  (1 
dmnh,  1  fmnh). 

Family  Bucerotidae — Hornbills 

Aceros  leucocephalus — Writhed  Hornbill 

The  Writhed  Hornbill  is  endemic  to  Mind- 
anao, Dinagat,  and  Camiguin,  in  forest  up  to 
1200  m  (Dickinson  et  al.,  1991;  Kennedy  et  al., 
2000).  The  records  from  Camiguin  in  1968 
consist  of  a  male  taken  from  Kasangsangan  in 
the  vicinity  of  Mt.  Catarman  at  1000  ft  (ca. 
300  m)  on  June  16  (Site  11)  and  another  male 
from  Mt.  Catarman  at  2000-4000  ft  (ca.  600- 
1200  m)  on  June  17  (Site  9).  A  female  was  also 
obtained  from  Mt.  Timpoong  at  3150  ft  (ca. 
950  m)  on  16  June  1969  (Site  13).  We  did  not 
record  it  during  our  fieldwork  in  the  1990s. 
Mallari  et  al.  (2001)  mention,  without  further 


reference  to  the  source,  a  juvenile  captured  in 
Ginsiliban  in  1993.  The  most  recent  evaluation 
of  this  species  found  it  to  be  fairly  common 
despite  "losing  ground  clearly  to  habitat  clear- 
ance, hunting  and  trapping  for  trade"  and 
assigned  it  a  Near-Threatened  status  (Collar  et 
al.,  1999). 

Specimens  Examined — Total  3.  Site  9  (1 
dmnh);  Site  11  (1  fmnh);  Site  13  (1  fmnh). 

Family  Pittidae — Pittas 

Pitta  erythrogaster — Red-bellied  Pitta 

The  Red-bellied  Pitta  occurs  from  Sulawesi  to 
the  Moluccas  and  New  Guinea  and  northeast 
Australia;  it  is  found  throughout  the  Philippines, 
except  the  Batanes  group  of  islands,  in  primary 
and  secondary  forest  usually  below  1000  m 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
Three  individuals,  a  male  and  two  females,  were 
taken  from  Mt.  Timpoong  at  3150-5700  ft  (ca. 
950-1700  m)  on  14-25  June  1969  (Sites  12  and 
13).  We  recorded  it  further  on  the  same 
mountain  at  1475  m  on  22  March  1995  (Site  8). 
In  May  1994,  it  was  the  most  common  bird  taken 
in  Victor  traps  in  Kital-is,  Sagay,  at  900-1 100  m 
to  1200-1400  m  (Sites  4-6). 

Specimens  Examined — Total  9.  Site  4  (2  msu- 
iit);  Site  5  (1  msu-iit);  Site  6  (2  msu-iit);  Site  8  (1 
fmnh);  Site  12  (1  dmnh);  Site  13  (2  fmnh). 

Family  Campephagidae — Cuckoo-Shrikes,  Minivets, 
and  Trillers 

Lalage  nigra — Pied  Triller 

The  Pied  Triller  ranges  from  the  Nicobars  to 
Southeast  Asia;  it  occurs  throughout  the  Philip- 
pines, except  the  Batanes  and  Babuyan  groups  of 
islands,  usually  in  open  areas  in  the  lowlands  up 
to  1400  m  (Dickinson  et  al.,  1991;  Kennedy  et 
al.,  2000).  It  was  recorded  from  Gidag-on  and 
Kasangsangan  in  the  vicinity  of  Mt.  Catarman  at 
500-2000  ft  (ca.  150-600  m)  in  June  1968  (Sites 
10  and  1 1).  In  June  of  the  following  year,  records 
came  mainly  from  Mt.  Timpoong  at  3150  ft  (ca. 
950  m;  Site  13),  Puntod,  at  800  ft  (ca.  150  m;  Site 
14),  and  Mantigue  Island  (Site  19). 

Specimens  Examined — Total  30.  Site  10  (9 
dmnh,  4  fmnh);  Site  11  (2  fmnh);  Site  13  (1 
dmnh);  Site  14  (4  dmnh,  5  fmnh);  Site  19  (5 
dmnh). 

Family  Pycnonotidae — Bulbuls 

Pycnonotus  goiavier — Yellow-vented  Bulbul 

The  Yellow-vented  Bulbul  occurs  widely  in 
Southeast  Asia  and  throughout  the  Philippines, 


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except  the  Babuyan  and  Batanes  groups  of 
islands,  in  scrub  and  second-growth  forest 
(Dickinson  et  al.,  1991:  Kennedy  et  al.,  2000). 
It  was  recorded  from  Mt.  Catarman  and  its 
nearby  areas  of  Gidag-on  and  Kasangsangan  at 
500-4000  ft  (ca.  150  1200  m)  in  June  1968  (Sites 
9-11).  In  June  1969,  additional  records  were 
obtained  from  Mt.  Timpoong  at  3150-4800  ft 
(ca.  950-1450  m;  Sites  12  and  13)  and  the  nearby 
area  of  Puntod  at  800  ft  (ca.  250  m;  Site  14)  as 
well  as  from  Mantigue  Island  on  28  June  (Site 
19).  We  recorded  it  further  in  Balbagon, 
Mambajao.  at  10  m  on  28  May  1992  (Site  1). 
Dickinson  et  al.  (1991)  noted  the  presence  of 
another  specimen  from  an  unspecified  locality  on 
Camiguin,  deposited  at  the  Museum  of  Com- 
parative Zoology,  Harvard  University. 

Specimens  Examined — Total  30.  Site  1  (1 
fmnh);  Site  9  (1  dmnh,  1  fmnh);  Site  10  (5  dmnh, 
5  fmnh);  Site  11  (2  dmnh,  3  fmnh);  Site  12  (1 
dmnh);  Site  13  (5  fmnh);  Site  14  (2  dmnh,  3 
fmnh);  Site  19  (1  dmnh). 

Ixos  everetti — Yellowish  Bulbul 

The  Yellowish  Bulbul  is  endemic  to  the  islands 
of  Biliran,  Bucas  Grande,  Dinagat,  Leyte,  Mind- 
anao, Panaon,  Samar,  and  Siargao,  the  Sulu 
group,  and  Camiguin,  usually  in  primary  and 
secondary  lowland  forest  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  Three  subspecies  are 
recognized,  including  /.  e.  catarmanensis,  which 
is  restricted  to  Camiguin  (Rand  &  Rabor,  1969). 
Our  comparison  of  specimens  at  fmnh  with  those 
of  other  named  subspecies  supports  the  distinc- 
tiveness of  the  Camiguin  population;  it  is  sub- 
stantially the  largest  and  darkest  among  the  three 
races  that  are  currently  recognized  (Dickinson  et 
al..  1991;  Kennedy  et  al.,  2000).  The  call  of  this 
species  is  equally  distinct  (Kennedy  et  al.,  2000). 
The  geographic  variation  observed  in  this  species 
as  currently  defined  is  unusually  great  and  is 
suggestive  of  a  potential  species  group.  Further 
taxonomic  studies  are  warranted. 

This  species  was  recorded  in  June  1968  from 
Mt.  Catarman  and  the  nearby  areas  of  Gidag-on 
and  Kasangsangan  at  500-4500  ft  (ca.  ISO- 
MOO  m;  Sites  9-11).  Additional  records  were 
obtained  in  June  1969  from  Mt.  Timpoong  at 
3150-5700  ft  (ca.  950-1700  m;  Sites  12  and  13) 
and  the  nearby  area  of  Puntod  at  800  ft  (ca. 
250  m;  Site  14).  In  May  1994,  it  was  one  of  the 
most  common  birds  mist  netted  in  Kital-is, 
Sagay,  at  900-1 100  m  to  1200-1400  m  (Sites  4- 
6). 


Specimens  Examined— Total  94.  Site  4  (8  msu- 
iit);  Site  5  (2  msu-iit);  Site  6  (1  msu-iit);  Site  9  (6 
dmnh,  15  fmnh);  Site  10  (6  dmnh);  Site  11  (9 
dmnh,  7  fmnh);  Site  12  (2  dmnh,  2  fmnh);  Site  13 
(20  dmnh,  13  fmnh);  Site  14  (2  dmnh,  1  fmnh). 

Family  Corvidae — Crows 

Corvus  /w/oY>/7nvn/?o.v— Large-Billed  Crow 

The  Large-Billed  Crow  occurs  from  Iran  to 
Northeast  Asia,  China,  Taiwan,  Ryukyus,  and 
Southeast  Asia;  it  is  found  nearly  throughout  the 
Philippines,  in  forest  edge  to  plantations  and  in 
towns  (Dickinson  et  al.,  1991;  Kennedy  et  al., 
2000).  A  male  specimen  was  taken  from  Gidag- 
on  in  the  vicinity  of  Mt.  Catarman  at  1000  ft  (ca. 
300  m)  on  24  June  1968  (Site  10). 

Specimens  Examined — Total  1.  Site  10  (1 
fmnh). 

Family  Turdidae — Robins,  Shamas,  and  Thrushes 
Copsychus  saularis — Oriental  Magpie-Robin 

The  Oriental  Magpie-Robin  occurs  from 
Pakistan  and  India  to  southern  China  and 
Southeast  Asia;  in  the  Philippines,  it  occurs 
throughout  the  country,  except  the  Palawan, 
Babuyan,  and  Batanes  groups  of  islands,  usually 
in  second-growth  and  scrubby  forest  (Dickinson 
et  al.,  1991;  Kennedy  et  al.,  2000).  It  was 
recorded  from  Mt.  Catarman  and  in  the  nearby 
area  of  Gidag-on  at  500  3000  ft  (ca.  150-900  m) 
in  June  1968  (Sites  9  and  10).  Additional  records 
were  obtained  in  June  1969  from  Mt.  Timpoong 
at  4000  ft  (1200  m;  Site  13)  and  the  nearby  area 
of  Puntod  at  800  ft  (ca.  250  m;  Site  14).  We 
recorded  it  further  in  May  1994  in  Kital-is, 
Sagay,  at  900-1 100  m  (Site  4)  and  1200-1400  m 
(Site  6). 

Specimens  Examined — Total  16.  Site  4  (3  msu- 
iit);  Site  6  (1  msu-iit);  Site  9  (1  fmnh);  Site  10  (4 
dmnh,  3  fmnh);  Site  13  (3  fmnh);  Site  14  (1 
fmnh). 

Saxicola  caprata — Pied  Bushchat 

The  Pied  Bushchat  occurs  from  Iran  to 
southwest  China,  Southeast  Asia,  and  New 
Guinea;  it  is  a  common  resident  in  the  Philip- 
pines, except  the  Sulu,  Palawan,  Babuyan,  and 
Batanes  groups  of  islands,  usually  in  fairly  dry, 
open  country  (Dickinson  et  al.,  1991;  Kennedy  et 
al.,  2000).  A  female  specimen  was  taken  from 
Mt.  Catarman  at  2000  ft  (ca.  600  m)  on  19  June 
1968  (Site  9). 

Specimens  Examined — Total  1.  Site  9  (1 
fmnh). 


64 


FIELDIANA:  ZOOLOGY 


Zoothera  andromedae — Sunda  Ground-Thrush 

The  Sunda  Ground-Thrush  occurs  from  Java 
and  Sumatra  to  the  Lesser  Sunda  Islands;  it  is  an 
uncommon  resident  in  the  Philippines,  where  it 
has  been  recorded  mainly  on  mountains  in  the 
northern  and  central  Luzon,  Mindanao,  Mind- 
oro,  Negros,  and  Panay,  in  the  understory  of 
forest  above  1000  m  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  We  recorded  a  single 
specimen,  taken  in  a  Victor  rat  trap,  from  Mt. 
Timpoong  at  1275  m  on  23  March  1995  (Site  7); 
this  is  the  first  record  for  Camiguin. 

Specimens  Examined — Total  1.  Site  7  (1 
fmnh). 

Zoothera  dauma — Scaly  Ground-Thrush 

The  Scaly  Ground-Thrush  is  an  uncommon 
winter  visitor  to  the  Philippines  from  Siberia, 
India,  China,  Japan,  Korea,  and  Taiwan.  It  has 
been  recorded  previously  only  on  the  islands  of 
Batan,  Catanduanes,  Fuga,  Luzon,  Marinduque, 
Mindoro,  Palawan,  and  Sibuyan,  where  it 
forages  on  the  ground  and  forest  understory  at 
all  elevations  (Dickinson  et  al.,  1991;  Kennedy  et 
al.,  2000).  Our  record  on  Camiguin  consists  of 
a  single  specimen  taken  in  a  Victor  rat  trap  on 
Mt.  Timpoong  at  1275  m  on  19  March  1995  (Site 
7).  This  is  the  first  record  of  this  species  for  the 
island. 

Specimens  Examined — Total  1.  Site  7  (1 
fmnh). 

Family  Sylviidae — Old  World  Warblers 

Phylloscopus   trivirgatus — Mountain    Leaf-War- 
bler 

The  Mountain  Leaf- Warbler  occurs  from  the 
Malay  Peninsula,  Borneo,  Sumatra,  Java,  and 
the  Lesser  Sunda  Islands  to  New  Guinea;  in  the 
Philippines  it  is  found  mainly  on  the  islands  of 
Luzon,  Mindanao,  Negros,  Palawan,  and  Panay, 
in  montane  and  mossy  forest  above  800  m 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
An  endemic  subspecies,  P.  t.  diuatae,  is  recog- 
nized on  Camiguin  and  Mt.  Hilong-hilong 
(eastern  Mindanao;  Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000),  though  Dickinson  (in 
litt.)  considered  this  in  need  of  reassessment. 
Records  on  Camiguin  were  obtained  from  Mt. 
Catarman  at  2000^950  ft  (ca.  600-1500  m)  in 
June  1968  (Site  9)  and  from  Mt.  Timpoong  at 
4800-5700  ft  (ca.  1450-1700  m)  in  June  1969 
(Sites  12  and  13).  It  was  also  recorded  in  Kital-is, 
Sagay,  at  900-1100  m  in  May  1994  (Site  4). 


Specimens  Examined— Total  31.  Site  4  (1  msu- 
iit);  Site  9  (3  dmnh,  5  fmnh);  Site  12  (10  dmnh, 
10  fmnh);  Site  13  (2  dmnh). 

Megalurus  timoriensis — Tawny  Grassbird 

The  Tawny  Grassbird  occurs  from  the  Lesser 
Sunda  Islands,  Moluccas,  and  Sulawesi  to  New 
Guinea  and  Australia;  it  is  found  throughout 
most  of  the  Philippines,  excluding  the  Babuyan, 
Batanes,  Palawan,  and  Sulu  groups  of  islands,  in 
grasslands  and  disturbed  forest  up  to  2000  m 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000).  It 
was  recorded  in  1968  from  Kasangsangan  in  the 
vicinity  of  Mt.  Catarman  at  1000  ft  (ca.  300  m) 
on  June  20  (Site  11).  Additional  records  were 
obtained  from  Mt.  Timpoong  at  3250  ft  (ca. 
980  m)  in  June  1969  (Site  13).  We  also  recorded 
it  in  May  1994  in  Kital-is,  Sagay,  at  1000- 
1300  m  (Site  5). 

Specimens  Examined — Total  10.  Site  5  (1  msu- 
iit);  Site  11  (2  fmnh);  Site  13  (3  dmnh,  4  fmnh). 

Cisticola  exilis — Bright-capped  Cisticola 

The  Bright-capped  Cisticola  ranges  from  India 
to  southern  China,  Taiwan,  Southeast  Asia 
(excluding  the  Malay  Peninsula)  to  New  Guinea, 
Australia,  and  the  southwest  Pacific;  it  occurs  on 
most  islands  of  the  Philippines,  excluding  the 
Babuyan,  Batanes,  Palawan,  and  Sulu  groups  of 
islands,  in  grassy  habitats  and  rice  fields  (Dick- 
inson et  al.,  1991;  Kennedy  et  al.,  2000).  Two 
adult  males  were  taken  in  1968,  one  from  Mt. 
Catarman  at  2000  ft  (ca.  600  m)  on  19  June  (Site 
9)  and  the  other  from  Kasangsangan  in  the 
vicinity  of  Mt.  Catarman  at  1500  ft  (ca.  450  m) 
on  21  June  (Site  11). 

Specimens  Examined — Total  2.  Site  9  (1 
dmnh);  Site  11  (1  fmnh). 

Family  Muscicapidae — Flycatchers 

Ficedula  hyperythra — Snowy-browed  Flycatcher 
The  Snowy-browed  Flycatcher  ranges  from 
the  Himalayas  to  southern  China,  Taiwan,  and 
Southeast  Asia;  it  occurs  on  most  of  the  larger 
islands  throughout  the  Philippines,  in  dense 
forest  understory  usually  above  1000  m,  but  is 
absent  on  Sibuyan  and  the  Batanes  and  Sulu 
groups  of  islands  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  On  Camiguin,  we 
recorded  this  species  for  the  first  time  in  Kital- 
is,  Sagay,  at  900-1100  m  in  May  1994  (Site  4) 
and  on  Mt.  Timpoong  at  1275  m  on  25  March 
1995  (Site  7). 


BALETE  ET  AL.:  ANNOTATED  CHECKLIST  OF  THE  BIRDS  OF  CAMIGUIN  ISLAND     65 


Specimens  Examined— Total  5.  Site  4  (4  nisi 
in);  Site  7  (1  fmnh). 

Ficedula  westermanni    Little  Pied  Flycatcher 

The  Little  Pied  Flycatcher  occurs  from  India 
to  southern  China  and  Southeast  Asia;  in  the 
Philippines,  it  has  been  recorded  only  from 
Luzon,  Mindanao,  Mindoro,  Negros,  southern 
Palawan,  and  Panay,  in  forest  and  forest  edge 
above  800  m.  usually  in  middle  story  and  forest 
canopy  (Dickinson  et  al.,  1991;  Kennedy  et  al., 
2000).  The  records  from  Camiguin  were  taken 
from  Mt.  Catarman  at  2000^950  ft  (ca.  600- 
1500  m)  in  June  1968  (Site  9)  and  on  Mt. 
Timpoong  at  3150-5700  ft  (ca.  950-1700  m)  in 
June  1969  (Sites  12  and  13).  On  29  May  1992,  we 
recorded  it  on  Mt.  Mambajao  at  1000  m  (Site  2) 
and  in  May  1994  in  Kital-is,  Sagay,  at  900- 
1300  m  (Sites  4  and  5). 

Specimens  Examined — Total  42.  Site  2  (1 
fmnh);  Site  4  (3  msu-iit);  Site  5  (1  msu-iit);  Site 
9(12  dmnh,  10  fmnh);  Site  12  (9  dmnh,  4  fmnh); 
Site  13  (2  fmnh). 

Cyornis  rufigastra — Mangrove  Blue  Flycatcher 

The  Mangrove  Blue  Flycatcher  ranges  from 
the  Malay  Peninsula,  Java,  Sumatra,  and  Sula- 
wesi to  Borneo  and  the  Philippines;  it  occurs 
throughout  most  of  the  islands  of  the  Philip- 
pines, except  the  Batanes  group,  usually  in 
disturbed  forest  habitat  (Dickinson  et  al.,  1991; 
Kennedy  et  al.,  2000).  It  was  recorded  from  Mt. 
Catarman  and  the  nearby  areas  of  Gidag-on  and 
Kasangsangan  at  500^1950  ft  (ca.  150-1500  m) 
in  June  1968  (Sites  9-11).  Further  records  were 
obtained  from  Mt.  Timpoong  at  3150-4800  ft 
(ca.  950-1450  m;  Sites  12  and  13)  and  the  nearby 
area  of  Puntod  at  800  ft  (ca.  250  m;  Site  14)  in 
June  1969.  We  also  recorded  it  in  Kital-is,  Sagay, 
at  900-1  100  m  in  May  1994  (Site  4). 

Specimens  Examined — Total  70.  Site  4  (4  msu- 
iit); Site  9  (8  dmnh,  9  fmnh);  Site  10  (7  dmnh,  4 
fmnh);  Site  11  (6  dmnh,  6  fmnh);  Site  12  (2 
dmnh,  4  fmnh);  Site  13  (7  dmnh,  5  fmnh);  Site  14 
(4  dmnh,  4  fmnh). 

Rhipidura  javanica — Pied  Fantail 

The  Pied  Fantail  ranges  from  the  Malay 
Peninsula,  Sumatra,  and  Java  to  Borneo  and 
the  Philippines,  where  it  occurs  throughout  most 
of  the  islands,  except  the  Babuyan  and  Batanes 
groups,  usually  in  lowland  second  growth, 
residential  areas,  bamboo  thickets,  and  man- 
groves (Dickinson  et  al.,  1991;  Kennedy  et  al., 


2000).  It  was  recorded  mainly  from  the  lower 
slopes  of  Mt.  Catarman  and  the  nearby  areas  of 
Gidag-on  and  Kasangsangan  in  June  1968  at 
500-2000  ft  (ca.  150-600  m;  Sites  9  11).  Addi- 
tional records  in  June  1969  were  taken  from  Mt. 
Timpoong,  at  3150  ft  (ca.  950  m;  Site  13)  and  the 
nearby  area  of  Puntod  at  800  ft  (ca.  250  m;  Site 
14).  We  also  recorded  it  in  May  1994  in  Kital-is, 
Sagay,  at  1000-1400  m  (Sites  5  and  6). 

Specimens  Examined — Total  32.  Site  5  (2  msu- 
iit); Site  6  (1  msuiit);  Site  9  (2  fmnh);  Site  10  (7 
dmnh,  7  fmnh);  Site  1 1  (3  dmnh,  3  fmnh);  Site  13 
(1  dmnh,  2  fmnh);  Site  14  (2  dmnh,  2  fmnh). 

Terpsiphone  cinnamomea — Rufous  Paradise-Fly- 
catcher 

The  Rufous  Paradise-Flycatcher  occurs 
throughout  the  Philippines,  excluding  the  Babu- 
yan, Batanes,  and  Palawan  groups  of  islands,  in 
understory  of  forest  up  to  1200  m;  it  also  occurs 
on  several  of  the  Talaud  Islands,  Indonesia 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
Three  specimens,  two  females  and  a  male,  were 
taken  on  16-23  June  1969  from  Mt.  Timpoong  at 
1500-3150  ft  (ca.  450-950  m;  Site  13);  another 
female  was  recorded  from  Puntod  at  800  ft  (ca. 
250  m)  on  27  June  1969  (Site  14).  In  May  1994,  it 
was  recorded  in  Kital-is,  Sagay,  at  900-1100  m 
(Site  4). 

Specimens  Examined — Total  5.  Site  4  (1  msu- 
iit); Site  13  (1  dmnh,  2  fmnh);  Site  14  (1  dmnh). 

Hypothymis  azurea — Black-naped  Monarch 

The  Black-naped  Monarch  ranges  from  India 
to  southern  China  and  Taiwan  to  Southeast 
Asia;  it  occurs  nearly  throughout  the  Philippines, 
except  the  Batanes  group  of  islands,  usually  in 
disturbed  forest  and  forest  edge  below  1500  m 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
The  population  on  Camiguin  is  recognized  as 
a  distinct  endemic  subspecies,  H.  a.  catarmanen- 
sis  (Rand  &  Rabor,  1969).  Records  from 
Camiguin  in  June  1968  came  from  Mt.  Catarman 
and  the  nearby  areas  of  Gidag-on  and  Kasang- 
sangan at  1000^1950  ft  (ca.  300-1450  m).  Fur- 
ther records  in  June  1969  came  from  both  Mt. 
Catarman  at  2000-4950  ft  (ca.  600-1450  m;  Sites 
9-11)  and  Mt.  Timpoong  at  3150  ft  (ca.  950  m; 
Site  13).  In  May  1994,  we  recorded  it  in  Kital-is, 
Sagay,  at  900-1 100  m  to  1200-1400  m  (Sites  4- 
6). 

Specimens  Examined — Total  33.  Site  4  (2  msu- 
iit). Site  5  (1  msu-iit);  Site  6  (2  msu-iit);  Site  9  (3 


66 


FIELDIANA:  ZOOLOGY 


dmnh,  4  fmnh);  Site  10  (1  dmnh,  1  fmnh);  Site  1 1 
(2  dmnh,  3  fmnh);  Site  13  (12  dmnh,  2  fmnh). 

Family  Motacillidae — Pipits,  Wagtails 

Anthus  hodgsoni — Olive  Tree-Pipit 

The  Olive  Tree-Pipit  is  an  uncommon  migrant 
from  eastern  Asia  to  India,  southern  China, 
Taiwan,  Ryukyus,  and  Southeast  Asia;  in  the 
Philippines  it  has  been  recorded  on  many  of  the 
larger  islands,  except  the  Batanes  and  Sulu 
groups,  usually  in  pine  and  oak  forest  above 
300  m  (Dickinson  et  al.,  1991;  Kennedy  et  al., 
2000).  On  Camiguin,  we  recorded  this  species  in 
primary  montane  forest  on  Mt.  Timpoong  at 
1275  m  on  24  March  1995  (Site  7);  it  is  the  first 
record  from  Camiguin. 

Specimens  Examined — Total  1.  Site  7  (1 
fmnh). 

Family  Artamidae — Wood  Swallows 

Artamus  leucorynchus — White-breasted  Wood- 
Swallow 

The  White-breasted  Wood-Swallow  occurs 
from  Borneo,  Sulawesi,  to  New  Guinea,  Aus- 
tralia, and  the  southwest  Pacific;  it  occurs 
throughout  the  Philippines,  except  the  Babuyan 
and  Batanes  groups  of  islands,  usually  at  forest 
edge  or  in  disturbed  forest  up  to  1800  m 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
Records  from  Camiguin  in  June  1968  came  from 
Mt.  Catarman  and  the  nearby  areas  of  Gidag-on 
and  Kasangsangan  at  500-2000  ft  (ca.  ISO- 
GOO  m;  Sites  9  and  10).  Additional  records  in 
June  1969  were  from  Mt.  Timpoong  at  3150  ft 
(ca.  950  m;  Site  13)  and  Puntod  at  800  ft  (ca. 
250  m;  Site  14). 

Specimens  Examined — Total  19.  Site  9  (1 
dmnh,  1  fmnh);  Site  10  (5  dmnh;  4  fmnh);  Site 
11  (1  dmnh);  Site  13  (2  dmnh;  4  fmnh);  Site  14  (1 
dmnh). 

Family  Sturnidae — Mynas  and  Starlings 

Aplonis  panayensis — Asian  Glossy  Starling 

The  Asian  Glossy  Starling  is  found  from 
eastern  India  to  Southeast  Asia;  it  occurs 
throughout  the  Philippines,  in  agricultural  and 
residential  areas  and  forest  edge  in  the  lowlands 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
Records  from  Camiguin  in  June  1968  came  from 
the  lower  slopes  of  Mt.  Catarman  and  nearby 
areas  of  Gidag-on  and  Kasangsangan  at  500- 
2000  ft  (ca.  150-600  m;  Sites  10  and  11).  Two 
other  specimens  were  obtained  in  Puntod  at 
800  ft.  (ca.  250  m)  in  June  1969  (Site  14).  We 


recorded  it  further  in  Balbagon,  Mambajao,  on 
28  May  1992  at  10  m  (Site  1). 

Specimens  Examined — Total  40.  Site  1  (1 
fmnh);  Site  10  (15  dmnh,  21  fmnh);  Site  11  (1 
dmnh);  Site  14  (2  dmnh). 

Sarcops  calvus — Coleto 

The  Coleto  occurs  almost  exclusively  in  the 
Philippines,  in  agricultural  plantations,  second 
growth,  and  forest  edge  in  the  lowlands;  its  only 
record  outside  the  country  is  on  Banggi  Island, 
off  Borneo  (Dickinson  et  al.,  1991;  Kennedy  et 
al.,  2000).  Within  the  Philippines  itself,  however, 
it  is  absent  from  the  Babuyan,  Batanes,  and 
Palawan  groups  of  islands  (Dickinson  et  al., 
1991;  Kennedy  et  al.,  2000).  In  June  1968,  two 
specimens  each  were  taken  from  Gidag-on  at 
500-1500  ft  (ca.  1 50^450  m;  Site  10)  and  Ka- 
sangsangan at  1000  ft  (ca.  300  m;  Site  11),  both 
in  the  vicinity  of  Mt.  Catarman. 

Specimens  Examined — Total  4.  Site  10  (2 
dmnh);  Site  1 1  (2  fmnh). 

Family  Nectariniidae — Spiderhunters  and  Sunbirds 

Anthreptes  malacensis — Plain-throated  Sunbird 

The  Plain-throated  Sunbird  occurs  throughout 
most  of  Southeast  Asia;  in  the  Philippines,  it  is 
found  on  almost  all  islands,  except  the  Babuyan 
and  Batanes  groups,  in  coconut  groves,  man- 
groves, and  second-growth  forest  in  the  lowlands 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
Records  on  Camiguin  came  mainly  from  Ka- 
sangsangan in  the  vicinity  of  Catarman  Moun- 
tain at  1500-2000  ft  (ca.  450-600  m)  in  June 
1968  (Site  1 1)  and  Puntod  at  800  ft  (ca.  250  m)  in 
June  1969  (Site  14).  A  single  specimen  was  taken 
on  Mt.  Timpoong  at  an  unknown  elevation  on 
16  June  1969.  Specimens  of  other  species  from 
the  same  site  and  date  were  taken  at  1500- 
3150  ft  (ca.  450-950  m). 

Specimens  Examined — Total  12.  Site  11  (3 
dmnh,  3  fmnh);  Site  13  (1  dmnh);  Site  14  (5 
dmnh). 

Nectarinia  jugularis — Olive-backed  Sunbird 

The  Olive-backed  Sunbird  occurs  from  South- 
east Asia  to  New  Guinea,  Australia,  and  the 
southwest  Pacific;  in  the  Philippines,  it  occurs  on 
almost  all  islands,  except  the  Babuyan  and 
Batanes  groups,  usually  in  heavily  disturbed 
habitats,  including  towns  and  cities,  below 
1000  m  (Dickinson  et  al.,  1991;  Kennedy  et  al., 
2000).  Records  from  Camiguin  were  obtained 
from  Mt.  Catarman  and  the  nearby  areas  of 


BALETE  ET  AL.:  ANNOTATED  CHECKLIST  OF  THE  BIRDS  OF  CAMIGUIN  ISLAND     67 


Gidag-on  and  Kasangsangan  at  500  3000  ft  (ca. 
150-900  m)  in  June  1968  (Sites  10  and  11).  In 
June  1969,  additional  records  came  from  Mt. 
Timpoong  at  1000-5000  ft  (ca.  300-1500  m; 
Sites  12  and  13)  and  the  nearby  area  of  Puntod 
at  800  ft  (ca.  250  m;  Site  14)  as  well  as  from  the 
island  of  Mantigue  (Site  19).  We  recorded  it 
further  in  Balbagon,  Mambajao,  on  28  May 
1992  at  10  m  (Site  1)  and  in  Kital-is,  Sagay,  at 
1000-1300  m  in  May  1994  (Site  5). 

Specimens  Examined — Total  94.  Site  1  (2 
fmnh);  Site  5  (1  msu-iit);  Site  10  (24  dmnh,  20 
fmnh);  Site  11  (4  dmnh,  6  fmnh);  Site  12  (5 
dmnh,  3  fmnh);  Site  13  (1  fmnh);  Site  14  (14 
dmnh,  6  fmnh);  Site  19  (8  dmnh). 

Nectarinia  sperata — Purple-throated  Sunbird 

The  Purple-throated  Sunbird  occurs  from 
India  and  Southeast  Asia;  it  is  found  throughout 
the  Philippines,  except  the  Batanes  group  of 
islands,  in  cultivated  areas,  mangroves,  and 
secondary  forest  in  the  lowlands  (Dickinson  et 
al.,  1991;  Kennedy  et  al.,  2000).  Records  from 
Camiguin  in  June  1968  were  obtained  from  Mt. 
Catarman  and  the  nearby  areas  of  Gidag-on  and 
Kasangsangan  at  500-3000  ft  (ca.  150-900  m; 
Sites  9-11).  In  June  the  following  year,  records 
were  obtained  from  Mt.  Timpoong  at  3150  ft 
(ca.  950  m;  Site  13)  and  in  the  nearby  area  of 
Puntod  at  800  ft  (ca.  250  m;  Site  14).  Addition- 
ally, Dickinson  et  al.  (1991)  noted  two  specimens 
from  an  unspecified  locality  and  elevation  on 
Camiguin  that  are  deposited  at  the  Museum  of 
Comparative  Zoology,  Harvard  University. 

Specimens  Examined — Total  39.  Site  9  (3 
dmnh,  3  fmnh);  Site  10  (6  dmnh,  6  fmnh);  Site 
11  (3  dmnh,  7  fmnh);  Site  13  (7  fmnh);  Site  14  (1 
dmnh,  3  fmnh). 

Family  Dicaeidae — Flowerpeckers 
Dicaeum  trigonostigma — Orange-bellied  Flower- 
pecker 

The  Orange-bellied  Flowerpecker  ranges  east- 
ern India  to  Southeast  Asia;  it  occurs  throughout 
the  Philippines,  except  on  the  Babuyan  and 
Batanes  groups  of  islands,  in  forest  edge,  second 
growth,  and  cultivated  areas  below  1500  m 
(Dickinson  et  al.,  1991;  Kennedy  et  al.,  2000). 
On  Camiguin,  this  species  is  represented  by  an 
endemic  subspecies,  D.  t.  isidroi  (Rand  &  Rabor, 
1969).  Records  from  Camiguin  were  obtained 
from  Mt.  Catarman  and  the  nearby  areas  of 
Gidag-on  and  Kasangsangan  at  500-4500  ft  (ca. 
150-1400  m)  in  June   1968  (Sites  9-11).  Addi- 


tional records  in  June  1969  were  obtained  from 
Mt.  Timpoong  at  800-3150  ft  (ca.  250-950  m: 
Site  13)  and  the  nearby  area  of  Puntod  at  800  it 
(ca.  250  m;  Site  14).  We  obtained  a  further 
record  of  it  in  Balbagon,  Mambajao,  at  10  m  on 
28  May  1992  (Site  1)  and  in  Kital-is,  Sagay,  at 
900-1 100  m  and  1200-1400  m  in  May  1994 
(Sites  4  and  6,  respectively). 

Specimens  Examined — Total  63.  Site  1  (1 
fmnh);  Site  4  (6  msu-iit);  Site  6  (5  msu-iit);  Site 
9  (5  dmnh,  4  fmnh);  Site  10  (3  dmnh,  4  fmnh); 
Site  11  (5  dmnh,  11  fmnh);  Site  13  (8  dmnh,  3 
fmnh);  Site  14  (6  dmnh,  2  fmnh). 

Dicaeum  pygmaeum — Pygmy  Flowerpecker 

The  Pygmy  Flowerpecker  is  common  through- 
out the  Philippines,  except  Mindoro  and  the 
Babuyan,  Batanes,  and  Sulu  groups  of  islands,  in 
forest,  forest  edge,  and  second  growth  mainly 
below  1000  m  (Dickinson  et  al.,  1991;  Kennedy 
et  al.,  2000).  A  single  record  exists  from 
Camiguin,  taken  on  13  June  1968  in  Kasangsan- 
gan, in  the  vicinity  of  Mt.  Catarman  at  an 
unknown  elevation.  A  specimen  of  Columba 
vitiensis  was  obtained  on  the  same  date  and 
from  the  same  site  at  1000  ft  (ca.  300  m). 

Specimens  Examined — Total  1.  Site  11  (1 
dmnh). 

Family  Zosteropidae — White-eyes 

Zosterops  everetti — Everett's  White-eye 

Everett's  White-eye  ranges  from  the  Malay 
Peninsula,  Borneo,  and  Talaud  to  the  Philip- 
pines; it  occurs  only  on  the  islands  southeast  of 
Luzon,  from  Samar  to  Mindanao  and  Cebu  to 
the  Sulu  and  Tawi-tawi  groups  of  islands,  in 
scrub  and  forest  up  to  1000  m  (Dickinson  et  al., 
1991;  Kennedy  et  al.,  2000).  Records  from 
Camiguin  were  obtained  from  Mt.  Catarman 
and  the  nearby  areas  of  Gidag-on  and  Kasang- 
sangan at  500^1950  ft  (ca.  150-1500  m)  in  June 
1968  (Sites  9  11).  Further  records  were  obtained 
in  June  1969  from  Mt.  Timpoong  at  3150  ft  (ca. 
950  m;  Site  13)  and  the  nearby  area  of  Puntod  at 
800  ft  (ca.  250  m;  Site  14). 

Specimens  Examined — Total  52.  Site  9  (4 
dmnh,  1  fmnh);  Site  10  (11  dmnh,  9  fmnh);  Site 
11  (4  dmnh,  9  fmnh);  Site  13  (4  dmnh,  8  fmnh); 
Site  14  (2  fmnh). 

Zosterops  nigrorum — Yellowish  White-eye 

The  Yellowish  White-eye  is  endemic  to  the 
Philippines,  where  it  occurs  on  Luzon,  Mindoro. 
Negros,    and    Panay    and    the    adjacent    small 


FIELDIANA:  ZOOLOGY 


islands,  in  forest  and  forest  edge  below  1000  m;  it 
is  absent  from  the  Babuyan,  Mindanao,  Pala- 
wan, and  Sulu  groups  of  islands  (Dickinson  et 
al.,  1991;  Kennedy  et  al.,  2000).  The  population 
on  Camiguin  is  the  southernmost  extension  of 
this  species'  range  and  is  treated  as  an  endemic 
subspecies,  Z.  n.  catarmanensis  (Rand  &  Rabor, 
1969).  Records  from  Camiguin  in  June  1968  were 
all  obtained  from  Mt.  Catarman  at  2000^1950  ft 
(ca.  600-1500  m;  Site  9).  Additional  records  in 
June  1969  were  all  obtained  from  Mt.  Timpoong 
at  3150-5400  ft  (ca.  950-1600  m;  Sites  12  and 
13).  We  recorded  it  further  on  Mt.  Mambajao  at 
1000  m  on  29  May  1992  (Site  2)  and  in  Kital-is, 
Sagay,  at  900-1 100  m  and  1200-1400  m  in  May 
1994  (Sites  4  and  6,  respectively). 

Specimens  Examined — Total  181.  Site  2  (2 
fmnh);  Site  4  (4  msu-iit);  Site  6  (3  msu-iit);  Site  9 
(53  dmnh,  39  fmnh);  Site  12  (36  dmnh,  15  fmnh); 
Site  13  (20  dmnh,  9  fmnh). 

Family  Estrildidae — Avadavat,  Parrotfinches, 
and  Munias 

Lonchura  leucogastra — White-Bellied  Munia 

The  White-Bellied  Munia  ranges  from  the 
Malay  Peninsula,  Sumatra,  Borneo,  and 
throughout  the  Philippines,  in  forest,  forest  edge, 
grassland,  and  in  rice  farms  (Dickinson  et  al., 
1991;  Kennedy  et  al.,  2000).  Records  of  this 
species  on  Camiguin  were  all  obtained  in  Gidag- 
on  and  Kasangsangan,  near  Mt.  Catarman,  at 
500-1000  ft  (ca.  150-300  m)  in  June  1968  (Sites 
10  and  11). 

Specimens  Examined — Total  6.  Site  10  (1 
dmnh,  3  fmnh);  Site  11  (1  dmnh,  1  fmnh). 

Lonchura  malacca — Chestnut  Munia 

The  Chestnut  Munia  is  recorded  from  India 
and  Nepal  to  southwestern  China,  Taiwan,  and 
Southeast  Asia  and  is  widespread  throughout  the 
Philippines,  associated  mainly  with  rice  fields, 
grasslands,  and  open  country.  The  only  records 
from  Camiguin  were  from  Kasangsangan  near 
Mt.  Catarman  at  1500  ft.  (ca.  450  m)  on  21-22 
June  1968  (Site  11). 

Specimens  Examined — Total  3.  Site  11  (1 
dmnh,  2  fmnh). 

Sight  Records 

Following  the  recommendation  appearing  in 
the  Rules  for  New  Records  (Kennedy  et  al., 
2000),  we  have  deliberately  omitted  from  the 
Species  Accounts  all  sight  records  that  were  not 


verified  or  supported  by  two  other  observers 
after  the  two-year  rule  for  publication.  The 
following  is  the  list  of  such  species,  including 
species  identified  by  calls  only,  which  when 
verified  would  constitute  additional  records  for 
Camiguin  and  is  indicative  of  the  potentially  far 
richer  avifauna  of  Camiguin  than  the  current 
Species  Accounts  would  suggest.  We  hope  that 
this  listing  will  encourage  other  ornithologists 
and  bird-watchers  to  conduct  more  studies  of 
Camiguin  birds. 

Family  Accipitridae 

Accipiter  sp. 

Observed  by  one  of  us  (B.R.T.)  along  a  trail 
leading  to  Site  4  on  12  March  1994  between  6  am 
and  3:30  pm. 

Family  Columbidae 

Ducula  poliocephala — Pink-necked  Pigeon 

Observed  by  one  of  us  (B.R.T.)  along  the  same 
trail  where  an  Accipiter  was  sighted  (see  above), 
on  the  same  date  and  time. 

Family  Cuculidae 

Cacomantis  merulinus — Plaintive  Cuckoo 

The  call  of  this  cuckoo  was  noted  by  B.R.T.  at 
Site  4  on  13  May  1994  as  one  of  the  most 
commonly  heard  birdcalls  at  this  site.  The  local 
name  of  the  bird  is  pitokai. 

Family  Strigidae 

Otus  sp. 

The  call  of  an  unidentified  scops-owl  was 
noted  by  B.R.T.  at  Site  4  on  14  May  1994  at  ca. 
5:40  am. 

Family  Oriolidae 

Oriolus  sp. 

The  call  of  an  unidentified  oriole  was  noted  by 
B.R.T.  at  Site  4  on  the  same  date  and  time  that 
the  scops-owl  was  heard  (see  above). 

Family  Pachycephalidae 

Pachycephala  sp. 

The  call  of  an  unidentified  whistler  was  noted 
by  B.R.T.  at  Site  4  on  the  same  date  and  time 
that  the  scops-owl  was  heard  (see  above). 


Discussion 

The   bird    fauna   of  Camiguin    is   currently 
represented    by    57    species,    consisting   of  55 


BALETE  ET  AL.:  ANNOTATED  CHECKLIST  OF  THE  BIRDS  OF  CAMIGUIN  ISLAND     69 


resident  breeders  and  two  migrants  in  26  bird 
families.  Seven  species  are  reported  here  for  the 
first  time:  Ixobrychus  cinnamomeus,  Porzana 
fusca,  Rallina  eurizonoides,  Zoothera  androme- 
dae,  Zoothera  dauma,  Ficedula  hyperythra,  and 
Anthus  hodgsoni.  Doves  and  pigeons  (Columbi- 
dae)  are  the  most  diverse  family  on  Camiguin, 
with  eight  species  recorded  on  the  island, 
followed  by  flycatchers  (Musicapidae)  with  six 
species,  and  thrushes  (Turdidae)  with  four 
species.  The  remaining  22  bird  families  are 
represented  by  one  to  three  species  only.  Because 
Camiguin  is  a  volcanic  island  of  recent  geological 
origin  and  is  separated  from  mainland  Mind- 
anao by  a  sea  channel  that  is  only  10  km  wide 
but  385  m  deep,  it  would  have  not  been 
connected  to  Mindanao  during  the  many  periods 
of  low  sea  level  in  the  Pleistocene  (Heaney,  1986; 
Heaney  &  Tabaranza,  2006a).  Thus,  the  current 
species  assemblage  reached  Camiguin  Island  by 
colonization,  and  their  presence  on  Camiguin  is 
documentation  of  the  capacity  of  these  species 
for  overwater  dispersal. 

A  brief  comparison  of  the  birds  on  Camiguin 
with  those  on  the  closest  islands,  and  sources  of 
potential  colonizers,  including  Mindanao  and 
the  central  Philippine  islands  of  Negros,  Panay, 
and  Cebu,  is  revealing.  Except  for  the  endemic 
Camiguin  Colasisi,  or  Hanging-Parrot,  Loriculus 
camiguinensis,  53  of  the  remaining  species  of 
resident  birds  are  also  found  on  Mindanao; 
Zosterops  nigrorum  does  not  occur  on  Mind- 
anao. With  Negros,  Panay,  and  Cebu,  Camiguin 
shares  50,  47,  and  42  of  its  resident  bird  species, 
respectively  (Dickinson  et  al.,  1991;  Kennedy  et 
al.,  2000).  Luzon,  which  is  farthest  away  from 
Camiguin  in  terms  of  potential  source  of 
colonizers,  also  shares  with  it  at  least  50  resident 
bird  species  (Dickinson  et  al.,  1991;  Kennedy  et 
al.,  2000).  These  data  indicate  a  remarkably  high 
faunal  similarity  with  Mindanao,  though  it  does 
not  provide  clear  evidence  of  biogeographic 
affinity,  given  an  equally  high  degree  of  similar- 
ity with  Luzon  and  Panay,  for  instance.  Thus,  it 
appears  that  Camiguin  has  been  colonized  by 
widespread  species. 

The  presence  of  at  least  one  species  endemic  to 
Camiguin  {Loriculus  camiguinensis,  described  in 
this  volume)  and  four  endemic  subspecies  of 
birds  clearly  indicates  that  colonization  rates 
have  been  so  low,  even  with  its  proximity  to 
Mindanao,  that  substantial  genetic  differentia- 
tion has  occurred.  In  this  regard,  it  is  noteworthy 
that    none    of   the    other    islands    adjacent    to 


Mindanao,  such  as  Basilan,  Bohol,  Dinagat. 
Leyte,  Samar,  or  Siargao,  regardless  of  area  and 
distance  from  it,  has  a  single-island  endemic 
(Dickinson  et  al.,  1991;  Kennedy  et  al..  2000; 
Peterson  et  al.,  2000).  All  these  islands  formed 
part  of  the  Pleistocene  island  of  Greater  Mind- 
anao and  thus  were  repeatedly  connected  by 
dryland  areas  during  much  of  the  Pleistocene 
(Heaney,  1986,  2000;  Heaney  &  Regalado,  1998; 
Steppan  et  al.,  2003),  unlike  Camiguin,  which 
remained  isolated  (Heaney  &  Tabaranza,  2006a). 

The  current  listing  of  birds  on  Camiguin  is  far 
from  comprehensive,  pending  more  systematic 
field  surveys,  but  based  on  the  quite  extensive 
collection  effort  during  the  late  1960s,  it  is 
worthwhile  to  note  here  the  apparent  absence 
or  depauperateness  of  several  families  that  are 
otherwise  well  represented  on  other  oceanic 
islands  in  the  Philippines  such  as  Sibuyan 
(currently,  the  most  comprehensively  studied 
small  oceanic  island  in  the  Philippines  for  birds; 
see  Goodman  et  al.,  1995).  Sibuyan  also  provides 
a  good  contrast  with  Camiguin  in  terms  of  the 
number  of  species  that  have  successfully  colo- 
nized oceanic  islands  in  the  Philippines.  Good- 
man et  al.  (1995)  identified  at  least  102  resident 
species  on  Sibuyan;  Camiguin's  current  list  pales 
in  comparison.  Among  raptors,  for  instance, 
only  one  eagle,  S.  cheela  (Accipitridae),  and  one 
owl,  N.  philippensis  (Strigidae),  have  been 
recorded  on  Camiguin.  Sibuyan,  in  contrast, 
has  at  least  six  species  of  eagles,  three  species  of 
owls,  and  one  species  of  falcon  (Falconidae) 
present.  A  similar  trend  is  apparent  in  rails 
(Rallidae),  swiftlets  (Apodidae),  and  kingfishers 
(Alcedinidae).  Also  noteworthy  is  the  poor 
representation  of  the  larger  doves  (Columbidae) 
on  Camiguin  that  occur  on  Sibuyan,  such  as 
Ducula  poliocephala,  D.  carola,  D.  aena,  and 
larger  parrots  (Psittacidae),  such  as  Priorniturus 
discurus  and  Tanygnatlnts  lucionensis  (Goodman 
et  al.,  1995). 

On  the  other  hand,  Camiguin  and  Sibuyan 
appear  to  share  the  absence  of  species  that  are 
generally  associated  with  wetlands  and  freshwa- 
ter swamps  (Anhingidae,  Threskiornithidae,  and 
Jacanidae)  as  well  as  species  in  high-elevation 
forest  habitats,  such  as  Orthotomus  cucullatus, 
Bradypterus  caudatus,  Rhynomyias  goodfellowi, 
Eumyias  panayensis,  Ficedula  crypta,  Culicicapa 
helianthea,  Serinus  estherae,  and  Pyrrhula  leuco- 
genis,  which  occur  on  some  of  the  larger  islands, 
including  Mindanao  (Dickinson  et  al.,  1991; 
Goodman  et  al.,   1995;  Kennedy  et  al.,  2000). 


70 


FIELDIANA:  ZOOLOGY 


Both  islands  also  lack  some  general  forest  and 
forest-edge  resident  birds  in  the  following 
families:  Podargidae,  Trogonidae,  Capitonidae, 
Chloropseidae,  Paridae,  and  Sittidae  and  Lanii- 
dae,  Dicruridae,  Oriolidae,  Rhabdornithidae, 
and  Timaliidae.  Camiguin  also  currently  lacks 
records  of  Turnicidae,  Rostratulidae,  Scolopaci- 
dae,  Tytonidae,  Caprimulgidae,  Hemiprocnidae, 
Picidae,  Eurylaimidae,  Hirundinidae,  Alaudidae, 
Oriolidae,  and  Pachycephalidae. 

It  is  further  interesting  to  note  that  on 
Camiguin,  in  contrast  to  Sibuyan  (Goodman  & 
Gonzales,  1990;  Goodman  et  al.,  1995)  and  other 
larger  and  more  speciose  islands,  several  wide- 
spread species  associated  with  open  country, 
cultivated  areas,  and  disturbed  habitats  in  the 
lowlands,  such  as  Pycnonotus  goiavier,  Rhipidura 
javanica,  and  Nectarinia  jugularis,  are  commonly 
found  in  relatively  intact  montane  and  mossy 
forest  up  to  the  peaks  of  Mt.  Timpoong.  A 
similar  pattern  in  mammals  was  earlier  observed 
on  Negros,  which  is  a  relatively  depauperate 
island  in  terms  of  mammal  diversity,  where 
several  commensal  species  {Suncus  murinus, 
Rattus  exulans,  and  R.  tanezumi)  are  present 
and  often  abundant  at  all  elevations  and  in  all 
habitat  types  (Heideman  et  al.,  1987;  Heaney  et 
al.,  1989),  and  on  Camiguin,  where  Suncus  mur- 
inus is  abundant  in  montane  forest  (Heaney  & 
Tabaranza,  1997;  Heaney  et  al.,  2006).  The  docu- 
mentation of  the  above  pattern  of  distribution 
along  elevational  gradients  has  led  to  the  hypo- 
thesis that  the  number  of  species  in  the  native 
community  of  small  mammals  in  mature  forest 
determines  the  success  of  non-native  small  mam- 
mals on  oceanic  islands  (Heaney  et  al.,  1999). 
Might  this  apply  to  birds  on  Camiguin  as  well?  Or 
might  habitat  disturbance  be  an  equally  important 
factor  contributing  to  this  trend  in  bird  diversity 
and  distribution  on  Camiguin?  Camiguin  is 
a  volcanically  active  island,  with  recorded  erup- 
tions of  at  least  two  of  its  volcanos  (Hibok-hibok 
and  Vulcan)  within  recent  times.  Mt.  Hibok-hibok 
erupted  in  1827,  1862,  1871-1875,  1897,  1902, 
and  1948-1958  and  Vulcan  in  1871  and  1874 
(http://www.volcano.si.edu/gvp/world/volcano. 
cfm?vnum= 070 1-08;  http://hannover.park.org/ 
Philippines/pinatubo/page9.html).  This  suggests 
that  the  island  has  had  bouts  of  habitat 
disturbance  brought  about  by  these  eruptions, 
aside  from  the  ongoing  human-induced  habitat 
alteration  and  fragmentation  (Heaney  &  Tabar- 
anza, 2005a).  Further  surveys  of  the  avifauna 
of  Camiguin  will  enable  a  more  comprehensive 


analysis  of  the  factors  that  influence  diversity 
and  distribution  patterns  of  birds  among 
oceanic  islands  in  the  Philippines. 

Finally,  we  note  that  Camiguin  is  the  home 
not  only  to  a  newly  described  parrot  (Loriculus 
camiguinensis;  Tello  et  al.,  2006)  and  four 
endemic  races  of  birds  (Ixos  everetti  catarma- 
nensis,  Hypothymis  azurea  catarmanensis,  Di- 
ceum  trigonostigma  isidroi,  and  Zosterops  ni- 
grorum  catarmanensis)  but  also  to  at  least  two 
species  of  mammals  (Heaney  &  Tabaranza, 
2006b),  one  amphibian,  and  seven  plants  (Hea- 
ney et  al.,  2006).  It  clearly  warrants  protection  as 
a  unique  portion  of  the  natural  heritage  of 
the  Philippines.  We  strongly  recommend  active 
protection  of  the  remaining  forest.  This  will 
benefit  the  wildlife  of  the  island  but  will  also  be 
essential  to  the  economic  and  social  stability  of 
Camiguin  because  the  mountain  forests  are  the 
source  of  the  island's  essential  and  precious 
water.  This  and  other  conservation  issues  are 
discussed  in  more  detail  in  Heaney  and  Tabar- 
anza (2006a). 


Acknowledgments 

For  assistance  with  fieldwork  on  Camiguin, 
we  thank  N.  Antoque,  E.  Batara,  N.  Batocael, 
N.  Bojo,  M.  Carmona,  A.  DeOcampo,  R. 
Fernandez,  M.  Jayoma,  L.  Mostrales,  A.  T. 
Peterson,  G.  Rosell,  A.  Tabaranza,  B.  Tabaranza 
III,  and  D.  Tabaranza.  Permits  were  provided  by 
the  Protected  Areas  and  Wildlife  Bureau  (Phi- 
lippine Department  of  Environment  and  Natural 
Resources);  we  especially  thank  A.  C.  Alcala,  C. 
Custodio,  M.  Mendoza,  and  W.  Pollisco.  We  are 
indebted  to  D.  Willard  for  access  to  specimens  at 
the  Field  Museum  and  especially  to  G.  Hess  for 
loans  of  specimens  and  information  on  collec- 
tions under  his  care  at  the  Delaware  Museum. 
We  wish  to  thank  Sean  Bober  and  Sarah  Lansing 
for  assistance  with  preparing  the  manuscript.  We 
are  grateful  to  D.  Willard  and  J.  Bates  for  helpful 
comments  on  earlier  drafts  and  to  N.  J.  Collar 
and  E.  C.  Dickinson  for  insightful  reviews  of  the 
manuscript.  Fieldwork  was  supported  by  the 
World  Environment  and  Resources  Program  of 
the  John  D.  and  Catherine  T.  MacArthur 
Foundation.  Additional  support  was  provided 
by  the  Marshall  Field  Fund,  Ellen  Thorne  Smith 
Fund,  and  Barbara  Brown  Fund  for  Mammal 
Research  of  the  Field  Museum. 


\LETE  ET  AL.:  ANNOTATED  CHECKLIST  OF  THE  BIRDS  OF  CAMIGUIN  ISLAND     71 


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72 


FIELDIANA:  ZOOLOGY 


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