Philippine Land Mammals (Part 1 of 2) by Filipiniana Online

PHILIPPINE LAND MAMMALS ,/'

EDWARD H. TAYLOR /

MANILA BUREAU OF PRINTING 1934 259463

R~ if r~L_

111l !,JI

THE GOVERNMENT OF THE PHILIPPINE ISLANDS DEPARTMENT OF AGRlCULTURE AND COMMERCE BUREAU OF SCIENCE MANILA

MONOGRAPHS OF THE BUREAU OF SCIENCE WILLIAM H. BROWN, Editor A. S. ARGUELLES, Associate Editor R. C. MCGREGOR, Associate Editor

"'I

Monograph 30 PHILIPPINE LAND MAMMALS By EDWARD H. TAYLOR

(Actual date of publication June 30, 1934) 2

-I

.:21.:2

PREFACE This study was undertaken at the suggestion of Dr. Elmer D. Merrill, then director of the Bureau of Science, Manila, who pointed out the need of a monographic catalogue of the mammals such as had been prepared for other vertebrate groups. A splendid checklist was prepared by N. Hollister * in 1912, but this was of no direct aid in the study and identification of Philippine forms. The literature on the group was scattered and difficult of access. It was believed that such a pUblication would be a great stimulus to the study of this least-known group of Philippine vertebrates, as well as an immediate aid in the identification of Philippine species. It was hoped that it would likewise be a guide to collectors and sportsmen. The work was begun in 1918, but due to my absenc-e in western Siberia during the winter of 1918-1919 and my subsequent return to the United States during the winter of 1919-1920 very little was accomplished until the spring of 1920, when an effort was begun to bring together a representative mammal collection. From March, 1920, to November, 1923, a large part of my time was spent in the field. Numerous islands were visited, and a collection of more than one thousand specimens was brought together. This collection, made chiefly in Tablas, Luzon, Mindanao, and Palawan, with a few important specimens from the other islands, forms the basis of this monograph. The collection of the Bureau of Science was only partly available as the bulk of it was on deposit in the United States National Museum. However, a number of important specimens of this collection together with certain other specimens in this museum were made available to me. The older collections in the Santo Tomas Museum, Manila, and the Ateneo de Manila were of meager use for study, since the pelage of the bulk of the specimens, long exposed to light, was changed beyond recognition, and the skulls, forming a part of the mounts, were not available for accurate identification. Then, too, locality data in numerous cases were missing altogether and in others obviously wrong. Many of the speci,.

* Philip.

Journ. Sci. ยง D 7 (1912) 1-64. 3

PREFACE

mens in these collections were obtained from the Mehan Botanical Garden in Manila, in which place no accurate data regarding acquisitions are kept. The specimens kept there on display are from numerous localities, and when turned over to the museum, dead, might be given the locality "Manila." A small collection in the College of Agriculture, University of the Philippines, was placed at my disposal by the authorities of this institution, and several important specimens were presented to me by the authorities of the School of Forestry at Los Banos, Laguna. Specimens of Philippine mammals in foreign scientific institutions have been ably reported on by various European mammalogists, and those in the United States National Museum have been reported on by N. Hollister, Gerrit S. Miller, Jr., J. A. Allen, Knud Andersen, D. G. Elliot, and Edgar A. Mearns. I am under deep obligation to the authorities of the United States National Museum for the loan of valuable specimens for comparison and description. Just as this paper is going to press I have learned of a small collection of Ph~lippine mammals hI the Museum of the University of Minnesota most of which are unidentified. My own collection, representing something more than one hundred species and subspecies, has produced twenty-one new forms, representing approximately 20 per cent of the species in the collection. As originally planned, the work Was to comprise only the smaller mammals. However, on advice, I have included the larger forms, although I feel the inadequacy of the treatment here given. This applies particularly to the ungulates. In bringing together, housing, and transporting my collection, I have placed myself under great obligation to many of my friends, and I take this opportunity of expressing my sincerest thanks to Dr. Elmer D. Merrill, former director of the Bureau of Science, for placing at my disposal the facilities of that in,. stitution; to Dr. Albert W. C. T. Herre, for ample laboratory space, innumerable courtesies, and specimens; to Gregorio Lopez, for much assistance and many specimens; to Capt. Francis Link, of the Philippine Constabulary, for assistance and specimens; to Mr. E. H. Pagenhart, director of the Coast and Geodetic Survey in the Philippines, for the courtesy of transportation on the steamship Pathfinder; to Capt. F. G. Engle, of the Pathfinder, for innumerable courtesies, assistance in collecting, and specimens; to Mr. Charles ,McIlvain, manager of the Celebes

PREFACE

Plantation Co., Saub, Cotabato, for extraordinary assistance, hospitality, and specimens; to John Dutton Wright, for assistance in financing my expedition to Palawan and extended hospitality at Baguio; to the officers of the Iwahig Penal Colony, for facilitating my collecting at Iwahig; to Mr. Macasaet, of Iwahig, Palawan, for examination of specimens in his private collection; to Dr. A. H. Aberin, Iwahig, Palawan, for specimens; to Mrs. Ida M. McCrory, for specimens; to Mr. R. C. McGregor, for specimens; to my wife, Hazel Clark Taylor, for patient assistance and cooperation in making possible the undertaking of the collections and the preparation of this monograph. For the housing of my bulky collection, care of specimens, and the use of a private laboratory I am under the greatest obligation to my friend Charles D. Bunker, assistant curator in charge of the Museum of Birds and Mammals, University of Kansas. Lastly, I desire. to express my sincerest thanks to Dr. H. H. Lane, head of the Department of Zoology, University of Kansas, who has tendered me able advice, has read and criticized the manuscript, and whose faith and sympathetic assistance have made it possible that the work be carried to a conclusion. EDWARD

Lawrence, Kansas, May 31, 1926.

H. TAYLOR.

CONTENTS Page.

ILLUSTRATIONS ...... _---_ .. ---_ .... -----_ ... _--_ ... ---------------------------------------------.------_ .. _----INTRODUCTION

15 21

Genus Manis Linnreus .... - ...................................................................... Manis javanica Desmarest ........................................................... .

22 30 52 56 59 61 64 66 67 68 68 68

ORDER INSECTIVORA .......•..................................................................................

FAMILY ERINACElIDlE ...... .................. ............................................................ ... .

74 74 75 76 76 77

Historical review of collections ........................................................... . Annotated bibliography ........................................ :.............................. . Collection and preservation of specimens ....................................... . Local names for mammals .................................................................... Distribution and faunal relations ....................................................... . Species erroneously credited to the Philippines ............................... . SYSTEMATIC CONSIDERATION ......................................................................... . CLASS MAMMALIA ........................................................................................... . ORDER LEPIDOTA .......................... ............................................................ ...... .. . . FAMILY MANIDlE .......................... ......•...........................................................

Genus Podogymnura Mearns ................................. ............ .............. . Podogymnura tr'uei Mearns ....................................................... . FAMILY

SORICID.JE

......................................................................................... .

Genus Pachyura Selys-Longchamps ................................................... . Pachyura occultidens Hollister ................................................... . Pachyura palawanensis sp. nov ................................................... . Pachyura luzoniensis (Peters) ................................................... . Genus Crocidura Wagler ................................................................... . Crocidura parvacauda sp. nov ....................................................... . Crocidura halconus Miller ........................................................... . Crocidura beatus Miller ............................................................... . Crocidura grayi Dobson ............................................................... . Crocidura mindorus Miller .......................................................... . Crocidura grandis Miller ........................................................... . Crocidura palawanensis sp. nov.................................................... Crocidura edwardsiana Trouessart ............................................ .

80 82 83 85 85

86 87 88

ORDER MENOTYPHLA ....................................................................................... . FAMILY TUPAIIDlE ........................................................................................... .

90 91 91 92 92 97 97

Genus Tupaia Raffles ..................................................................... ..... . Tupaia cuyonis Miller ................................................................. . Tupaia palawanensis (Thomas) ............................................... . Tupaia mollendorffi Matschie ....................................................... .

99 101 103 105

ORDER DERMOPTERA .......................................................................... ............. .. . FAMILY GALEOPTERIDlE ............................................... .'................................. .

Genus Cynocephalus Boddaert .............. _.......................................... . Cynocephalus volans (Linnams) ............................................... .

CONTENTS

FAMILY TUPAIIDJE-Continued. Genus Urogale Mearns ........................................................................ . Urogale everetti (Thomas) ........................................................... . ORDER CHIROPTERA ....................................................................................... . SUBORDER MEGACHIROPTERA ......................................................................... . FAMILY PTEROPIDJE .. ........................................~ .............................................. . SUBFAMILY HARPYIONYCTERINJE ................................................................... . Genus Harpyionycteris Thomas ......................................................... . Harpyionycteris whiteheadi Thomas ....................................... . SUBFAMILY MACROGLOSSINJE ......................................................................... . Genus Macroglossus Cuvier ............................................................... . Macroglossus lagochilus lagochilus (Matschie) ....................... . Macroglossus fructivorus sp. nov............................................... . Genus Eonycteris Dobson ..................................................................... . Eonycteris robusta Miller ........................................................... . Eonycteris longicauda sp. nov....................................................... . SUBFAMILY PTEROPINJE ................................................................................. . Genus Acerodon Jourdan ..................................................................... . Acerodon lucifer (Elliot) ........................................................... . Acerodon jubatus (Eschscholtz) ............................................... . Acerodon jubatus jubatus (Eschscholtz) ................................. . Acerodon jubatus mindanensis Andersen ............................... . Genus Pteropus Brisson ....................................................................... . Pteropus leucopterus Temminck ............................................... . Pteropus vampyrus (Linnams) ............... _.................................. . Pteropus vampyrus lanensis (Mearns) ..................................... . Pteropus hypomelanus Temminck ............................................. . Pteropus hypomelanus cagayanus (Mearns) ........................... . Pteropus hypomelanus tomesi Peters ......................................... . Pteropus mearnsi Hollister ......................................................... . Pteropus ptimilus Miller .............................................. : ................ . Pteropus speciosus Andersen ..................................................... . Pteropus mimus Andersen ........................................................... . Pteropus balutus Hollister ........................................................... . Pteropus tablasi sp. nov............................................................... . Genus Rousettus Gray ...................................................................... . Rousettus amplexicaudatus (Geoffroy) ..................................... . Genus Dobsonia Palmer ....................................................................... . Dobsonia peronii (Geoffroy) ..................................................... . Genus Ptenochirus Peters ................................................................... . Ptenochirus jagorii (Peters) ....................................................... . Genus Cynopterus Cuvier ..................................................................... . Cynopterus archipelagus sp. nov ................................................ . Cynopterus brachyoiis Mtiller ..................................................... . Cynopterus brachyotis brachyotis (M tiller) ............................. . Genus Thoopterus Matschie ............................................................... . Thoopterus nigrescens (Gray) ...... ............................... .............. . Genus Megrerops Peters ............................................... :....................... . Megrerops wetmorei sp. nov .................................................. 路.. 路... 路 FAMILY EMBALLONURIDJE ............................................................................. .

Palte.

106 106 107 113 116 116 117 118 119 119 122 125 126 127 131 134 135 136 138 138 142 147 160153 153 158 158 160 162 163 164

166 168 169 171

172 176 17'1

178 178 181 182 184 184 188 189 190 191 194

CONTENTS

9 Page.

SUBFAMILY EMBALLONURINlE .. ______ .. _... __..... __ .. _.. _.. __... __._ .. __.__ .... _..... __ ._. _____ ._.__ Genus Emballonura Temminck ...... _.. _....... ___ ...... __.... _......... __ ._ ..... __ .__ .. _. Emballonura alecto (Eydoux and Gervais) _. __._ ..... __ . __ . ____ ... __ ... _ Emballonura alecto alecto (Eydoux and Gervais) .. _... _..... ____ ._ Emballonura alecto palawanensis subsp. nov __ ______ __. _______ .____ .____ Genus Taphozous Geoffroy __ _._. _._. __ __ ____ . ___ . ___ ._ .. __ ______ . __._. ___ _. ______ . __ ._. __ . Taphozous philippinensis Waterhouse . _____ ____________ __ ._. ___ _. __ .____ .__ Genus Taphonycteris Dobson __ .__ _____ _._ .. ___ ._. __ .. ___ _. ______ . ____ _._. __ ._ ...... _. __ .__ ._ Taphonycteris capito Hollister .________ ._. ______ ._._______________________ ________ Taphonycteris pluto (Miller) __ . _______ __ _. ______ ._. ____ ..... _____ .. ___ ___ .. _._ __ FAMILY MOOADERMIDlE __ ..___ .. __ . __ .. __ _.__ ._ .__ .. __....__ .. _.__ '_.... __ ... __ ........__ .. ___ ... _.. __ ...... Genus Megaderma Geoffroy .______ .. _.. __ ____ _.. __ . ___ .__ . ________ .____ _._. ___ __.______ ___ _ Megaderma spasma spasma (Linnreus) ... __ __ . ______ ._ .. _._. _________ .. __ FAMILY RHINOLOPHIDlE Miller .... __ ._____ __ . ___ __ __ . ___ __ .. __ . ___ .. __ _._.. _. _._. __.. _.. _.. __ ___ __ Genus Rhinolophus Lacepede . __ .__ . ___ .. __ . ___ .____ _____ ____ __ __ . _____ ___ __ ____ . ____ .__ ___ Rhinolophus philippinensis Waterhouse __ __ ... ____ .. _____ __ ._ .. ____ ._. ___ ._ Rhinolophus hirsutus An der se n .__ .... _.. __ .__ ...... ____ . __ __ . ____ . _________ _.... Rhinolophus virgo Andersen __ __ ___ ... _.. __ _____ ... ______ ..... _.. __ .. ______ .____ . __ . Rhinolophus a nderseni CabreJra . ____ __ __ ._..... __ __ __ ____ .. __ .. __ .____ ._ .. __ ... _. Rhinolophus anderseni andersen i (Cabrera) __ .__. ______ .... __ ._... __.. _ Rhinolophus anderseni requalis Allen __ .. __ .. __ ......... _..... ___ .. __. __ .. ___ .. Rhinolophus subrufus An dersen ......... ____ . ___ __ __ .. __ . __ . __ .______ .___ ... _._.. Rhin'olophus arcuat us arcuatus (Peters) ______ ._. __________ ._. __ _______ .. ___ Rhinolophus arcuatus exiguus Andersen _. __.. ______ ... __ ...... ____ _.___ Rhinolophus inops Andersen .. _.... _.... _. ___ ...... __ .. _........ _. __ ... _... ___ ... _.. Rhinolophus bunkeri sp. nov ...... __ ..................... _.... _... __ ........... _._. FAMILY HIPPOSIDERIDlE ........................ _. __... ___ .... _......... _........ _. _.... _._._ ...... _.. ___ . Genus Hipposideros Gray __ ........ __ _............... __ .... _....... _._ ....... _.... _... _._._ __ Hipposideros obscurus (Peters) ........... _.. ___ _... _.. _... _.............. _.. _. Hipposideros coronatus (Peters) ....... _........ _._ ... __ .. __ ............ ____ ... _ Hipposideros bicolor (Temminck) ..................... ____ ._._...... _. __ ....... Hipposideros wrighti sp. nov.... __ ._...................... __ ._ .. __ .... __ ..... _...... Hipposideros pygmreus (Waterhouse) .. _.. _... __ .. ______ . ___ .__ .. _____ .. Hipposideros antricola (Peters) ________ ___ ...................__ .... ______ ...... Hipposideros diadema (Geoffroy) ........ _...... __ . _____ .......... _....... ____ ... _. Hipposideros diadema griseus (Meyen) __ .... __ ...... __ ........... _........ Hipposideros diadema anderseni subsp. nov.... __ .. _.. ____ .... _..... __..... Genus Chilophylla Miller . __ ..................... ______ ...... __ ._ ...... ___.. __ ..... _... ___ ._. Chilophylla hirsuta Mille r . ___ ___ .. __ .__ ... _.. __ .__ _.. ___ .. __ .. _.. __ ._ ... _..... ___ .... FAMILY VESPERTILIONIDlE _________ .. _... _... __ ._. __ .__ .__ .... __ ..... _... _. ___ ._ .... _. __ .. _______ . __ ._._ SUBFAMILY MURININlE . _______ ._... _. __ . _______________ . ___ ... __ . __ .__ ..__ ._ .. _.__ .___ .____ .___ ......__ .__ Genus Murina Gray .____ ._ .. _______ . ____ .... _____ .... __ . __________ .. ____ . ___________ .. ____ . _____ Murina cycIotis Dobson .____ .. ___ .. _. __ ._ ... __ ._. ___ ... _. __ . ___ . ____ . _______ .______ .___ SUBFAMILY KERIVOULINlE . __ . _______ . ____________________ . ____ ... __ .. _. __ ._ .... _____ ._. __ ._ ...... ___ .__ Genus Kerivoula Gray ____________ ._._. ______ ___ .. ___ ._____ . __ ... _____ . __ . _________ _...... ______ Kerivoula whiteheadi Thomas ._. ______ ._._. ______ .____ ._ .. _._____ ... __ .. _________ Kerivoula jagorii (Peters ) ______________ ._ .. _______ . __ ._____ .. ______ .__ . ___ .______ ._.. Kerivoula hardwickii (Horsfield) ________________ ___ ._______ . ___ . ____ .___________ Kerivoula pellucida (Water hou se) ___ .. ___ ._ .... _. ___... ______ ... ___ . __ ._ .. __ .

195 195 196 196 197 200 200 204 204 205 206 207 208 210 211 214 215 216 219 219 220 221 224 227 228 228 229 230 232 234 236 237 239 241 241 242 246 247 248 249 251 251 252 253 253 256 257 258 261

CONTENTS Page.

SUBFAMILY MINIOPTERINlE ........................................................................... .

Genus Miniopterus Bonaparte ...................................................... ' .... . Miniopterus pusillus Dobson ....................................................... . Miniopterus australis Tomes ......................................................... . Miniopterus paululus Hollister .................................. ............. . Miniopterus tibialis Tomes ... ~ ..................................................... . Miniopterus eschscholtzii (Waterhouse) ................................. . Miniopterus tristis (Waterhouse) ............................................ .

264 264 266 268 269 272

272 275

SUBFAMILY VESPERTILIONINlE ..................................................................... .

277

Genus Myotis Kaup ............................................................................... . Myotis rufopictus (Waterhouse) • ............................................... Myotis macrotarsus (Waterhouse) ... ;....................................... . Myotis jeannei sp. nov................................................................... . Myotis patricire sp. nov................................................................. . Myotis browni sp. nov..................................................................... . Myotis herrei sp. nov................................................................... . Genus Glischropus Dobson ................................................................... . Glischropus tylopus (Dobson) ................................................... . Genus Pipistrellus Kaup ............... :.................................................. . Pipistrellus irretitus (Cantor) ................................................... . Pipistrellus tenuis (Temminck) ................................................... . Pipistrellus abramus (Temminck) ............. ~ ............................. . Pipistrellus imbricatus (Horsfield) ........................................... . Genus Pterygistes Kaup ....................................................................... . Pterygisies stenopterus (Dobson) ............................................ . Genus Tylonycteris Peters ................................................................... . Tylonycteris pachypus (Temminck) ......................................... . Genus Scotophilus Leach .........................................................:........... . ScotophiIus temminckii (Horsfield) ........................................... .

277 279 281 284 286 288 290

FAMILY MOLOSSIDlE ........................•...............................................................

Genus Chrerephon Dobson .................................................................. . Chrerephon luzon us Hollister ....................................................... . Genus Philippinopterus gen. nov ....................................................... . Philippinopterus lanei sp. nov................................................... . Genus Cheiromeles Horsfield ............................................................... . Cheiromeles torquatus Horsfield ............................................... . ORDER PRIMATES ............................................................................................. .

SUBORDER LEMUROIDEA ....................................•............................................. .. FAMILY T ARSIIDlE ........................................................................................... .

Genus Tarsius Storr ............................................................................. . Tarsius phiIippensis Meyer ........................................_.................. . Tarsius fraterculus Miller ........................................................... . Tarsius carbonarius Heude ......................................................... . FAMILY NYCTICEBIDJE ..................................................................................... .

G€nus Nycticebus Geoffroy ................................................................. . Nycticebus menagensis (Lydekker) ......................................... .

292

293 294 296

297 298 300 301 302 303 304 305 306 309 310 311

314 314 320 321 323 324 325 325 326 327 331· 332 332 333

CONTENTS

11 Page.

SUBORDER ANTHROPOIDEA .............................................................................. FAMILY CERCOPITHECIDJE ................................................................................

Genus Pithecus Geoffroy........................................................................ Pithecus mindorus Hollister ........................................................ Pithecus philippinensis (Geoffroy) ..........................................., Pithecus mindanensis (Mearns) ................................................ Pithecus suluensis (Mearns) ...................................................... Pithecus cagayanus (Mearns) .............:..................................... ORDER

CARNIVORA

..........................................................................................

FAMILY FELIDJE ..............................................................................................

Genus Felis Linnreus................................................................................ Felis catus Linnreus ........................................................................ Felis minuta Temminck ................................................... ............. FAMILY CANIDJE ...................... ........... ..... ............................. ............. ..............

Genus Canis Linnreus.............................................................................. Canis familiaris Linnreus ..... ... ............................ ....... ....... .......... FAMILY

MUSTELIDJE

SUBFAMILY

......................................................................................

MELINlE

......................................................................................

Genus Mydaus Cuviel' ............................................................................ Mydaus schadenbergii J entink ................................ _.................. . Mydaus marchei Huet .................................................................. SUBFAMILY

LUTRINJE

........â&#x20AC;˘...........................................................................

Genus Aonyx Lesson .......................................................................... .. Aonyx cinerea (IlUger) ...................... .......................................... FAMILY VIVERRIDlE

........................................................................................

SUBFAMILY VIVERRINlE .............. -'.... .........., ................................................. .'..

Genus Arctictis Temminck .................................................................. Arctictis whitei Allen .................................................................... Genus Viverra Linnreus ........................................................................ Viverra tangalunga Gray ............................................................ Genus Paradoxurus Cuvier ................................................................ Paradoxurus philippinensis Jourdan ........................................ Parado:x;urus torvus Thomas ........................................................ Paradoxurus minax Thomas ........................................................ SUBFAMILY HERPESTINJE ........................... ........ .... .. ,. .....................................

Genus Mungos Geoffroy and Cuvier ................................................ Mungos palawanus Allen ............................................................ Mungos parvus (Jentink) ............................................................ ORDER RODENTIA ...................................................... ......... ......... ................ .. .... FAMILY SCIURIDlE .................................................;............. .. ........................... SUBFAMILY NANNOSCIURINlE ............................................... .... ..... ........... .....

Genus Nannosciurus Trouessart .......... ............................... ............... Nannosciurus concinnus (Thomas) .............................................. Nannosciurus surrutilus Hollister ............................................ Nannosciurus samaricus Thomas ................................................ Nannosciurus luncefordi sp. nov................................................

335 336 336 337 338 340 342 342 343 345 345 345 346 348 348 348 348 349 350 350 351 352 352 353 353 354 354 355 357 358 360 360 361 362 364 364 365 365 366 367 368 368 369 370 372 373

CONTENTS Page.

SUBFAMILY SCIURINlE

Genus Sciurus Linnreus -.-----.... -.. -------.----------------.---.----.---.-.------.-- .. ---.--Sciurus steerii Gi.lnther ______ .__ .. ___ .. , ___ ._____ .__ ..... __ .____ .. __ .__ .... ____ . __ .. __ _ Sciurus juvencus Thomas __________ .______ ..... ____ ... __ . ____ .______ .__ .____ . ____ .__ . Sciurus albicauda Matschie ___ :__ .__ .......... ______ .__ .__ .... ____ . __ .____________ _ S'ciurus mollendorffi Ma tschie __ .. __ .____ .____ .__ .. __________ .____ . __ . __ .____ .__ __ Sciurus mindanensis Steere ____ .____ . ______ . __ . ____ .____ .______________ .. ________ .. Sciutus philippinensis Waterhouse ________ .______ .______ ... __ .________ .____ .__ Sciurus samarensis Steere ________ .____ .________ .____ .. __ . __ ... __ ..... ____________ .. __ FAMILY PETAURISTIDlE

__ .... ________ . __ .. ____ ... ___ .________ .. __ . ______ ... __ . __ . ____ . ___________ . ______ _

Genus Sciuropterus Cuvier __ ... ____ .__ ._._ .. ____ ._. __ ... _._. ____ .___ ._._._ .. ___ .____ .. _______ _ Sciuropterus nigripes Thomas __ . _____ . ___________________ . __ .. __________ .____ .__ _ Sciuropterus nigripes nigripes (Thomas) __ ... ______ .__ . __ . ______________ __ Sciuropterus nigripes elassodontus Osgood. ____ .. __ .__ . ________________ . Sciuropterus crinitus Hollister . ____ . __ .____ ... _______________ .____ .. ____ .________ . FAMILY MURIDlE ___ . __ . _________ . _____ . _____ .. _. __ . __ . ____ ... ____ ._. ___ . __ . _______________ .. _._ ... _________ . __ SUBFAMILY PHL<EOMYINlE ______________ .. ______ .. __ . ___ .. __ ........ __ ... __ . __ ... ____ ...... _______ . ___ __

Genus Phlceomys W aterhouse _______________ .____ .__ .. __ ._________________ .. ________________ . Phlreomys cumingi Waterhouse _______ .__ .__________ .... ________ . ___ .______ .______ _ Phlreomys pallidus (Nehring) _________________ .____ .. _________ .____ . ______________ _ Phlreomys elegans (CabJlera) _____________________________________ . __ ..... __ .. __ .. __ . Genus Crateromys Thomas _______ ._ .. _. _______ .___ ._____ .__ .____ .. ___ .__________ ... __________ _ Crateromys schadenbergi (Meyer) ____________ ... ________ .__ .__ .____________ __ SUBFAMILY MURINlE ____________________ .. _____ . _____ ._._. ___ . __________ . _____ . __ . __ ._ .. ____ ._. _____ . ___ . __ _

Genus Celrenomys Thomas _________________ . ________ .______ .________ .. ____ . ----- ___ .--. __ . --.. Celrenomys siIaceus (Thomas) _______________ .. __ .__________ . __________________ ... Genus Rhynchomys Thomas _______________ .______ .. _______ . __ .________ . ___________ .________ ._ Rhynchomys soricoides Thomas _.______________ .. __________ .___________________ __ Genus Chrotomys Thomas _______ . __ . ___________________ .__ .__________ .__ .____ .______________ _ Chrotomys whiteheadi Thomas __________ .______ .________ . ______ ._____ .______ .____ _ Genus Crunomys Thomas __________ .. __________________ .... ___________ .__ ... _____ .. _________ .__ Crunomys fallax Thomas __ .______________________ . ________ ... ______________ .. _____ __ Crunomys melanius Thomas _______ .. ____ . ______ .____ .. __ .____ .________ . __________ __ Genus Bullimus Mearns __ .____ . __ . ______________ .__ .. _____________________ .... ___________ .____ _ Bullimus luzonicus (Thomas) ___________ . ________ . _______ . __ . __ . _______________ __ Bullimus bagobus 'Mearns _________ . ____________ . __ . __ .______ .. ______________ . ______ __ Genus Rattus Fischer ___________________ .________ . ___________________ ... ____________ . __ .. ____ .... __ Pala wanensis group _______________ .____________ .. ________ . __ . ____ . ____ .____ .. ____ . __ .. ___ __ Rattus palawanensis sp. nov _______ . __ .. ________________ . __ .. __ .____ .______ _ Everetti group .__________________________ ._____ ._______ .... ____ ._._ .... _______ .__ ... ___ .... ___ Rattus everetti (Gunther) ____ ._______________ .________ ._____________ .. ____ . Rattus gala (Miller) _________ .__ . ________ . __ .__________ . _________ .______ .. ____ __ Rattus tagulayensis (Mearns) ________________________________________ __ Rattus albigularis (Mearns) . ___________ ._________ .. __ .__ .. __________ .__ .. N orvegicus group _____________ .. ____ .___ .___________________________________ .. __ .__________ __ Rattus norvegicus (Erxleben) .______ .____ .____ .. _______________ ... __ . __ .. Rattus tyrannus (Miller) ________ .______ . __ .___________ ... ________ .. __ .____ ..

375 375 376 379 379 380 381 382 382 383 383 384 384 388 389 390 391 391 392 394 396 397 398 400 402 402 404 405 406 406 407 408 409 411 411

413 414 416 416 419 420 421 422 423 425 425 427

CONTENTS MURINlE-Continued. Genus Rattus Fischer-Continued. Rattus group .................................................................. ................. . Rattus magnirostris (Mearns) ........................................... . Rattus rattus (Linn::eus) ..................................................... . Rattus mindorensis (Thomas) ........................................... . Rattus coloratus (Hollister) ............................................. . Rattus robiginosus (Hollister) ........................................... . Rattus mindanensis rnindanensis (Mearns) .......... ............ Rattus mindanensis tablasi subsp. nov............................. . Alexandrinus group ..................................................................... . Rattus kelleri (Mearns) .................................................... .. Rattus zarnboang::e (Mearns) ............................................. . Dat::e group ..................................................................................... . Rattus benguetensis (Hollister) ....................................... . Rattus dat::e (Meyer) ........................................................... . Concolor-ephippium group ........................................................... . Rattus negrinus (Thomas) ................................................. . Rattus todayensis (Mearns ) ................................ .............. .. Rattus vigoratus (Hollister) ............................................. . Rattus querceti (Hollister) ................................................. . Rattus luteiventris (Allen) ................................................. . Rattus pantarensis (Mearns) ........................................... . Rattus vulcani (Mearns) ................................................... . Rattus basil anus (Hollister) ................................................ .. Rattus leucoph::eatus (Hollister) ......................................... . Rattus rnayonicus. (Hollister) ........................................... . Rattus ornatulus (Hollister) ............................................. . Rattus calcis (Hollister) ..................................................... . Genus Apomys Mearns ....................................................................... . Apomys hylocretes Mearns ........................................................... . Apomys insignis Mearns .. ............................................................ . Apomys insignis insignis (Mearns) ............................................ Apomys insignis bardus (Miller) ................................................. . Apomys petr::eus Mearns ............................................................. . Apomys major Miller ................................. ............. .................. ... . Apomys musculus Mearns ........................................................... . Apomys micro don Hollister ....................................................... . Genus Insul::emus gen. nov.......... ......................................................... . Insul::emus calamianâ&#x201A;Źnsis sp. nov ............................................... . Genus Carpomys Thomas .................................... :....... ......................... . Carpomys melanurus Thomas ........................................ ............ Carpomys ph::eurus Thomas ....................................................... . Genus Tryphomys Miller ..................................................................... . Tryphomys adustus Miller ........................................................... . Genus Mus Linn::eus ........................................................................... . Mus castaneus Waterhouse ....................................................... . Mus commissarius Mearns ....................................... _.................. .

SUBFAMILY

Page.

429 430 431 433 434 435 437 439 440 440 442 443 443 444 445 446 448 449 450 452 453 454 455 456 457 457 458 459 461 462 462 463 465 466 468 469 469 470 471

472 474 476 476 477 478 479

CONTENTS

SUBFAMILY MURINlE-Continued. Genus Batomys Thomas --------------------------------------:--------------------------------Batomys grantii Thomas -------------------------------------------------------------Batomys dentatus Miller -------------------------------------------------------------Genus Limnomys Mearns -------------------------------------------------------------------Limnomys sibuanus Mearns -------------------------------------------------------Limnomys picinus Hollister --_____________________________________________________ _ Limnomys mearnsi Hollister -------------------------------------------------------Genus Tarsomys Mearns -------------------- __________________________________ ----:--------Tarsomys apoensis Mearns ______________________________________________________ ---FAMILY HYSTRICIDlE -______________________________ --------------------------------- --------- ----- -------SUBFAMILY HYSTRICINlE ______________________________________________________ -------------------------Genus Thecurus Lyon ______________________________________________________ ---------------------Thecurus pumilus (Gunther) _______________________________________________________ _ ORDER ARTIODACTYLA ______________________________________________________ ------------------------------- FAMILY SUIDlE ______________________________________________________ -------------------------------------------Genus Sus Linnreus ______________________________________________________ ------ _________________ _ Sus philippensis Nehring ___________________________________________________________ _ Sus calamianensis Heude _____________________________________________________________ _ Sus minutus Meude _______________________________________________________________________ _ Sus cebifrons Heude _____________________________________________________________________ _ Sus inconstans Heude ______ "' ___________________________________________________________ _ Sus mindanensis Major _______________________________________________________________ _ Sus ahoenobarbus Huet _______________________________________________________________ _ Sus barbatus balabacensis Major _____________________________________________ _ F AMILY TRAGULIDlE - -- ------------ - --,.. -_. --.---."---".--_. --- -.. ---" --_. _.". --_. _.. -_ .. .. ". ---_. _.. -_. Genus Tragulus Brisson _________ . ________________ ,.-___________________________________________ _ Tragulus nigricans Thomas ______ . ________________________________________________ _ FAMILY CERVIDlE ___________________________________________________________________________________________ _ Genus Rusa H. Smith _____________________________________________________________________________ _ Rusa philippinus (Smith) ___________________________________________________________ _ Rusa alfredi (Sclater) _______________________________________________________________ _ Rusa barandanus (Heude) _________________________________________________________ _ Rusa francianus (Heude) ___________________________________________________________ _ Rusa basilanensis (Heude) _________________________________________________________ _ Rusa nigellus Hollister _________________________________________________________________ _ Rusa calamianensis (Heude) _______________________________________________________ _ Rusa nigricans (Brooke) _____________________________________________________________ _ FAMILY BOVIDlE _____________________________________________________________________________________________ _ Genus Bubalus Smith ______________________________________________________ ---------------------Bubalus mainitensis Heude _________________________________________________________ _ Bubalus moellendorffi Nehring ___________________________________________________ _ Bubalus mindorensis Heude _______________________________________________________ _ -~

Bubalus bubalus (Linnreus) -------------------------------------------------------ADDENDUM ______________________________________________________ -------------------------------------------------INDEX ______________________________________________________ ------------------------------------------------------------

Page_

480 482 483 484 485 486 487 487 488 489 490 490 490 493494 494496 498 499 500 501 501 50Q'

504505 506 50S 507 508 513" 51!), 516 517"

51S518

519

522 520-

526 527" 527"

528 529

530 5H

ILLUSTRATIONS PLATE 1

FIGs. 1 and 2. Tupaia palawanensis (Thomas) ; No. 19, Sir J. Brooke Point, Palawan; enlarged. 3 and 4. Tupaia palawanensis (Thomas); No. 20, Sir J. Brooke Point, Palawan; enlarged. 5 and 6. Tupaia cuyonis Miller; Bureau of Science No. 24, Cuyo Island; para type. PLATE

FIG. 1. Cynopterus brachyotis bra,chyotis Muller; E. H. T. collection No. 214, Polillo Island; a, skull, dorsal view; b, skull, ventral view; c, skull, lateral view; d, lower jaw, dorsal view; e, lower jaw, lateral view. (Drawings by Dolores Hood.) Enlarged. 2. Urogale eve?'etti (Thomas), t ype, British Museum No. 79.5.3.11, Zamboanga, Mindanao; a, skull, lateral view; b, lower jaw, lateral view; c, skull, dorsal view; d, skull, ventral view; e, lower tooth row, I, upper tooth I'OW; X 1. (From Lyon.) PLATE 3 FIG. 1. Acet'odon jubatus jubatus (Eschscholtz), female; S. Negros, British Museum No. 77.12.10.11; a, skull, dorsal view; b, skull, lateral view; c, skull, ventral view; d, lower jaw, dorsal view; e, lower jaw, lateral view; X 1. (From Andersen.) 2. Thoopterus nigrescens (Gray), female; type, British Museum No. 62.10.21.6 Morotai; a, skull, dorsal view; b, skull and lower jaw, lateral view; c, lower jaw, dorsal view; d, skull, ventral view; e, skull and jaw, front view; X 1. (From Andersen.) PLATE 4

FIG. 1. Ptenochirus jagorii (Peters) ; E. H. T. collection No. 215, Odiongan, Tablas; a, skull, dorsal view; b, skull, ventral view; c, skull, lateral view; d, lower jaw, dorsal view; e, lower jaw, lateral view; actual length of skull, 39 millimeters. (Drawings by Dolores Hood.) 2. Pteropus tablasi sp. nov.; type, E. H. T. collection No. 312; a, head, dorsal view; b, head, ventral view; c, head, lateral view; d, lower jaw, lateral view; e, lower jaw, dorsal view; total length, 46 millimeters. PLATE 5

FIG. 1. Pteropus hypomelanus tom.e si Peters, female; British Museum No. 94.7.14.2, Darvel Bay, northeastern Borneo; a, skull, dorsal view; b, skull, ventral view; c, skull, lateral view; d, lower jaw, ventral view; e, lower jaw, lateral view; X 1. (From Andersen.) 15

ILLUSTRATIONS

FIG. 2. Pte1'opus leucopte?'us Temminck (type of Pteropus chinensis) ; British Museum; a, skull, dorsal view; b, skull, ventral view; c, skull, lateral view; d, lower jaw, dorsal view; e, lower jaw, lateral view; X 1. (From Andersen.) PLATE 6

FIGS. 1 to 5. Eonycteris robusta Miller; 1, adult female, No. 442; 2, adult male, No. 445; 3, adult male, No. 436; 4, No. 562; 5, young male, No. 449. 6 to 8. Eonycteris longicauda sp. nov.; 6, No. 438; 7, No. 4'35; 8, adult male, No. 439. PLATE 7

FIG. 1. Harpyionycteris whiteheadi Thom~s; type, British Museum No. 97.5.2.7, Mindoro; a, upper jaw, ventral view, enlarged; b, upper jaw, lateral view, enlarged; c, lower jaw, lateral view, enlarged; d, lower jaw, dorsal view, enlarged; e, skull, dorsal view; X 1. (From Thomas.) 2. Ptenochir;us jagori Peters; male, type, Berlin Museum No. 2523; a, skull, dorsal view, Xl; b, skull, ventral view, Xl; c, skull, lateral view, Xl; d, lower jaw, lateral view, Xl; e, lower jaw, dorsal view, Xl; I, palate ridges, Xl; g, skull and jaw, front view, X 2. 3. Macroglossus lagochilus lagochi~us (Matschie); female, United States National Museum No. 123440, Philippine Islands; a, skull, dorsal view; b, skull, ventral view; c, skull, lateral view; d, lower jaw, lateral view; X H. (From Miller.) PLATE 8

FIG. 1. Megaderma spasma spasma (Linnreus) ; a, face, front view; b, skull, dorsal view; c, skull, ventral view. (From Dobson.) 2. Cheirome~es torqu,atus Horsfield; a, male, front view of face; b, female, front view of face. (From Dobson.) 3. Glischropus tylopus (Dobson); front view of face. (From Dobson.) 4. Rhinolophus philippinensis Waterhouse; a, face, lateral view; b, nose leaf, much enlarged. (From Dobson.) PLATE 9

FIG. 1. Pterygistes stenopterus (Dobson); body, head, and wing, ventral view. (From Dobson.) 2. Scotophilus temmincki (Horsfield); a, skull, dorsal view; b, teeth, front view. (From Dobson.) 3. Ke1'ivoula pellucida (Waterhouse); ear, front view. (From Dobson.) 4. Kerivoula hardwickii (Horsfield); ear, front view. (Fl'om Dobson.) 5. Murina cyclotis Dobson; head, lateral view. (From- Dobson.) 6. Hipposide~'os pygmreus (Waterhouse); nose leaf, enlarged. (From Dobson.)

ILLUSTRATIONS

FIG. 7. Hipposideros diadema (Geoffroy); head, front view. (From Dobson.) 8. Hipposide?'os bicolo?' (Temminck); head, front view. (From Dobson.) PLATE 10 FIGS. 1 and 2. C?'ateromys schadenbergi (Meyer); Bureau of Science collection 'No. 139, Mount Data, Luzon; 1, skull, dorsal view; fJ, skull, ventral view. 3 and 4. Pte?'opus speciosus Andersen. Malinipa Island; 3, skull, dorsal view; 4, skull, ventral view. 5 and 6. Phlr:eomys pallidus Nehring; E. H. T. collection, No. 650, Kalinga, Luzon; 5, mandibles, dorsal view; 6, skull, ventral view. PLATE

FIGS. 1 and 2 Vive?"?"a tangalunga Gray; Polillo, No. 708. 3 and 4. Vive?"?'a tang(kl~tn!la Gray; Los Banos, Luzon, No. 705. PLATE

FIGS. I, 2, and 3. Sciuropte?"us nigripes Thomas; E. H. T. collection No. 647; Sir J. Brooke Point, Palawan; skull and jaws, about X 1. 4, 5, and 6. Sciu1'opterus nigripes Thomas; E. H. T. collection No. 648; Iwahig, Palawan, skull and jaws, about X 1. PLATE

FIG. 1. Batomys grantii Thomas; type, British Museum No. 95.8.2.16, Mount Data, Luzon; skull, dorsal, ventral, and lateral views; lower jaw, lateral view; molar teeth, greatly enlarged. (From Thomas.) 2. C(krpomys melanurus Thomas; type? British Museum No. 95.8.2.12, Mount Data, Luzon; skull, dorsal, ventral, and lateral views; lower jaw, lateral view; molar teeth, greatly enlarged. (From Thomas.) PLATE 14 FIG. 1. Bullimus luzonicus (Thomas); type, Mount Data, Lepanto, Luzon; a, skull, dorsal view; b, skull, ventral view. (From Thomas.) 2. Carpomys ph::eurus Thomas; type, Mount Data, Lepanto, Luzon; a, skull, dorsal view; b, skull, ventral view; c, skull, lateral view; d, lower jaw. (From Thomas.) 3. Rhyncho?nys sor;icoides Thomas; type, Mount Data, Lepanto, Luzon; a, skull, dorsaT view; b, skull, ventral view; c, skull, la~ral view; d, lower jaw. (From Thomas.) 4. CrateromI!Js schadenbe1'gi (Meyer); Mount Data, Lepanto, Luzon; upper and lower molar teeth. (From Thomas.) PLATE

FIG. 1. Chrotomys whilteheadi Thomas; Mount Data, Lepanto, Luzon; (I, skull, dorsal view; b, skull, ventral view; c, skull, lateral view; d, lower jaw; e, upper and lower molars, enlarged. (From Thomas.) 269463-2

ILLUSTRATIONS

FIG. 2. Celrenomys silaceus (Thomas); type, Mount Data, Lepanto, Luzon; a, skull, dorsal view; b, skull, ventral; c, skull, lateral; d, lower jaw; e, upper and lower molar teeth enlarged. (From Thomas.) 3. Cruncnnys fallax Thomas; type, Isabela, central-northern Luzon; a, skull, dorsal view, X 2; b, skull, ventral view, X 2; c, skull, lateral view, X 2; d, lower jaw, X 1; e, skull, dorsal view, X 1; f, upper and lower molars, enlarged. (From Thomas.) PLATE

FIG.!. Tryphcnnys adustus Miller; type, United States National Museum No. 151511, Haights-in-the-Oaks, Benguet, Luzon; a, skull, dorsal view; b, skull, lateral; c, skull, ventral; d, lower jaw, internal view; e, lower jaw, external view; X 1. (From Miller.) 2. Batomys dentatus Miller; type, United States National Museum No. 151506, Haights-in-the-Oaks, Benguet, Luzon; a, skull, dorsal view; b, skull, lateral; c, skull, ventral; d, lower jaw, internal view; e, lower jaw, external view; X 1. (From Miller.) 3. Apomys musculus Mearns; type, United States National Museum No. 145770, Camp John Hay, Baguio, Benguet, Luzon; a, skull, dorsal view; b, skull, lateral; c, skull, ventral; d, lower jaw, internal view; e, lower jaw, external view; X 1. (From Miller.) PLATE

FIG. 1. Rattus tyrannus (Miller); type, Philippine Bureau of Science collection No.8 (in the United States National Museum), Ticao Island; a, skull, dorsal view; b, skull, lateral view; c, skull, ventral view; d, lower jaw, internal view; e, lower jaw, external view; X 1. (From MiNer.) 2. Chilophylla hirsuta Miller; type, No. 144821, Alag River, Mindoro; face, front view; X 3. (From Miller.) PLATE

FIG. 1. Thecurus pumilus (Gunther); Palawan, No. 18; skull, X 1. 2. Thecurus pumilus? Busuanga, No. 138;' skull, X 1. 3. Thecurus pwmilus? Busuanga, No. 1000; skull, X 1. PLATE 19

FIG. 1. ChilophyUa hi1路suta Miller; type, noseleaf, greatly enlarged. (From Miller.) 2. Pteropus leucopterus Temminck; Philippine Islands; a, skull and . lower jaw, lateral view; b, lower jaw, dorsal view; c, upper jaw, ventral view; X 1. (From Dobson.) 3. Megaderma spasma Linnreus, Johore; a, skull, dorsal view; b, skull, ventral view; c, skull, lateral view; d, lower jaw; X 1. (From Miller.) PLATE 20

Pithecus mindorus Hollister; type, United States National Museum No. 144674; Alag River, Mindoro; 1, skull, lateral view; 2, lower jaw, dorsal view; 3, lower jaw, lateral view; 4, skull, dorsal view; 5, skull, ventral view.

ILLUSTRATIONS PLATE

Rusa Ilig ellus Hollister; type, United States National Museum collection No. 14-1711, Mount Malindang, Mindanao; 1, skull, dorsal view, reduced; 2, upper molar teeth; 8, lower jaw, reduced. PLATE

FIG. 1. Sus philippensis Nehring; male, Luzon; a, skull, dorsal view; b, skull, ventral; c, skull and lower jaw, lateral; d, lower jaw, dorsal; e, lower jaw, ventral; X 搂. 2. Sus philippem;is Nehring; Luzon; young female; skull and lower jaw, lateral view. PLATE 23 FIGs. 1 and 2. Caingins at Saub, Cotabato Province, Luzon. Type locality of Crocidllra par l)ica uda, Rhinolophus bunkeri, Myotis browni, Philippinopterus lanei, and Nannosciu1"US lunce/m路di. PLATE

Thecu1'US pumilus (Gunther). Specimens captured and kept alive at the Bur eau of Science. 2. Tragulus nigricans Thomas.

FIG. 1.

PLATE

Busuanga

FIG. 1. Tamarao recently killed. 2. A young tamarao in the Mehan botanical garden, Manila. TEXT FIGURES P a ge.

FIG. 1.

2. 3. 4. 5. 6. 7. 8. 9. 10. 11. 12.

The Philippine Islands, showing where collections of mammals have been made. Density of stippling suggests the relative amount of collecting.................................................... Distribution of SOl'icidre in the Philippines............................ Dist.ribution of CynocephalUS, Tupaia, and Erinaceidre in the Philippines .................. ____ .. __ __ ..... ..____ ... ____ .__ ...... __ ............. __ ... Distribution of Macroglossus, Eonycte1"is, Harpyiony ct el'is, and six species of Rattus in the Philippines .. __ .................... Distribution of Acerodon in the Philippines ... ........ ____ ...... __ ..... Distribution of Pte7'ol1'Us in the Philippines ...... __ .................... Distribution of Rousettus, Dobsoniq., and Ptenochi7"US in the Philippines ........................ ....... __ ...................... ....... __ ............ ____ .. __ Distribution of Cynopterus, Mega:?'ops, and Emballonuridre in the Philippines ________ ..... __ __ ...... __ ...... __ ....... ............ __ ................ __ Distribution of Megaderma and five species of Rhinolophus in the Philippines __ .. __ ...... __ ...... .... __ ...... ____ ........... ..................... __ .. Distribution of five species of Rhinolophus in the Philippines ........... __ .. ________ .____ .____ .. __ __ .... ____ .. __ .. __ ... ____ .... __ ....... __ ____ .......... __ . Distribution of five species of Hipposideros in the Philippines --....... ____ .. __ . ______ ... __ . ____ ....... ____ __ __ ..... __ ... ____ ..... ____ .. __ ........__ .. ...... Distribution of Hipposidel'os, Chilophylla, and Myotis in the Philippines .. __ .... __ ........ __ ......... ................... ____ ............ __ ........ __ ...... __

53 79 100 121 136 149 173 183 214 225 235 243

ILLUSTRATIONS Page.

FIG. 13. Distribution of Murina, Kerivoula, and Miniopterus in the Philippines .................................................................................... 14. Distribution of Scotophilus, Tylonycteris, Pterygistes, and Pipistrellus in the Philippines................................................ 15. Distribution of Molossidre, Felidre, and Mustelidre in the Philippines _.. _... _....... _...... _._ ............... _........ __ ................ _... _............ 16. Distribution of Viverridre in the Philippines ............. _............. _ 17. Distribution of N annosciurus and Sciuropterus in the Philippines .. __ ._ ........ __ .... _._ .... _..... _._ ... _...... ___ .. _....... _... _.. _...... _._._.............. 18. Dist.ribution of Celamo1nys, Rhynchomys, Chrotomys, Crunomys, and Sciurus in the Philippines................................ 19. Distribution of Phlmom;ys, C1'ate1~omys, and Apomys in the Philippines ........... _.................................. "\i••••••• .••••• _.................... 20. Distribution of seven species of Rattus in the Philippines.... 21. Distribution of seven members of the Rat'tus rattus group in the Philippines ........ _._ ..... _......................... .............................. 22. Distribution of four species of Rattus in the Philippines........ 23. Distribution of six species o~ Rattus in the Philippines.... 24. Distribution of Bullimus, Insul:;emus, Carpomys, and Tryphomys in the Philippines ................ _............. _......................... 25. Distribution of Mus, Batomys, Limnomys, and Tarsomys in the Philippines ............................................................................

254 295 315 356 371 377 392 417 428 441 447 473 481

PHILIPPINE LAND MAMMALS By

EDWARD

TAYLOR

INTRODUCTION As is true of many other groups of Philippine animals, the first notable collection of mammals was brought together by Hugh Cuming, a British collector, who visited the Islands during the period 1836-1840. The localities visited were numerous, but since no intensive mammal collecting was done the material is only fairly representative of the localities visited and chiefly of the smaller mammals. After Cuming's time the next most important collection was made by John Whitehead. This collection (the bulk of which was from one locality, Mount Data, Lepanto, Luzon) contained new and important genera, apparently confined to the highmountain region of Lllzon. Certai~ of these genera, as has later been discovered, have representatives also on other Philippine islands. Shortly after American occupation of the Philippines, Maj. Edgar A. Mearns, a mammalogist of long experience, began collecting there-particularly in certain of the more outlying parts-and brought together a remarkable collection containing fifty-four new forms. This contribution was the largest ever made to mammal study in the Philippines. My own collection, while approximating that of Mearns in size and number of species, contains fewer new forms. This collection, which forms the basis of this monograph, contains the following new species and subspecies: Pachyu?"a palawanensis, Taytay, Palawan. Crocidura Tjarvacauda, Saub, Cotabato Province, Mindanao. CrOCidU1"a palawanensis, Sir J. Brooke Point, Palawan. Mac?'oglossus !ructivo't路us, Tatayan, Cotabato Province, Mindanao. Pteropus tablasi, Tablas. Cynopterus a't'chipelagus, Polillo. Meg:erops wetmO?'ei, Tatayan, Cotabato Province, Mindanao. Eonycte?'is longicauda, Montalban caves, Rizal Province, Luzon. Emballonura alecto palawanensis, Iwahig, Palawan. 21

PHILIPPINE LAND MAMMALS

Rhinolophus bunlce?~i, Saub, Cot.abato Province, Mindanao. Hipposide'ros wright-i, Baguio, Benguet Subprovince, Luzon. Hipposide?'os diadema anderseni, Montalban, Rizal Province, Luzon. Myotis jeannei, Caldera, Zamboanga Province, Mindanao. Myotis patricire, Agusan Provinoo, Mindanao. Myotis browni, Saub, Cotabato Province, Mindanao. Myotis he?'rei, Montalban, Rizal Province, Luzon. Philippinopterus lanei, Saub, Cotabato Province, Mindanao. Nannosciurus luncefo?'di, Saub, Cotabato Province, Mindanao. Rattus palawanensis, Sir J. Brooke Point, Palawan. Rattus mindanensis tablasi, Tablas. Insulremus calamianensis, Busuanga.

Aside from the four collections mentioned many smaller collections have been made and their content is discussed together with the four mentioned collections in' the following pages. HISTORICAL REVIEW OF COLLECTIONS

References to Philippine mammals occur in records of practically all of the early voyages to the Philippines. ,For the most part the larger mammal forms, such as the deer, w:ild boar, and monkeys are mentioned; references also occur to the bats, which form a very conspicuous part of the fauna. Among the earlier records of mammals is that of William Dampier, * who visited Mindanao in 1686. He remarks: "The Hogs are ugly Creatures; they have all great knobs growing over their Eyes, and there are multitudes of them in the Woods. They are commonly very poor, yet sweet." Probably one of the earliest collections made in the Philippines was that of a Jesuit missionary, George Joseph Camel (or as frequently spelled, Kamel or Camelle). There is some doubt as to where his specimens were taken, but certain forms, notably a form of Tarsius reported as being from Luzon, certainly is not from that island. A paper of Camel's published by Jacobo Petiver t contained Camel's list of Philippine animals. These, for the most part, cannot be specifically identified. Apparently no specimens were sent to Europe but merely papers which consisted of notes and drawings. The native names listed, while giving a certain

* ANew Voyage Round the W orId. London

(1729). Quadrupedibus Philippensibus Tractat. a Reverendo Georg. Jos. Camel. transmissus Jacobo Petiver, Pharmacop. et Societ. Regire. Soc. Londini. Philos. Trans. 25 (1706-1707-1708) 2200-2201.

t De

INTRODUCTION

amount of evidence as to localities and species, are not at all conclusive evidence of origin of the specimens. * Friederich Eschscholtz obtained some specimens in the Philippines about 1825 (1823-1826) among which was the type of Acerodon jubatus. This species, described in 1831, is probably the first Philippine specimen of mammal to receive a specific "Linnrean" designation. Fortune Eydoux and Paul Gervais (1830-1832) obtained a few specimens of mammals, among which were the types of Rh~no足 lophus luctus rufa (= Rhinolophus rufus) and Vespertilio (Nycticeus) alecto (= Emballonura alecto). Hugh Cuming collected wi'dely in the Philippines during the period from 1836 to 1840. While primarily interested in other classes of animals he collected some mammals. These specimens were sent to the Zoological Society of London and most of the new forms were described by G. R. Waterhouse in the proceedings of that society. A few specimens remained for others to describe. The collection contained the following: Crocidura grayi Dobson. Galeopithecus philippinensis Waterhouse [= Cynocephalu8 volans (Linnreus) ] . Cynopterus marginatus var. philippensis Gray [= Cynopterus b1'achyotis brachyotis (Muller)]. Cynopte7"US marginatus var. c'U.lTlting-U Gray [ = Cynopterus brachyotis brachyotis (Muller)]. Eleutherura philippinensis Gray [= Roussettus amplexicaudatu8 (Geoffroy) ] . Pte7'opus chinensis Gray (= Pteropus leucopterus Temminck). Taphozous philip'[linensis Waterhouse. Megader?na philippinensis Waterhouse [ = Megaderma spasma spasma (Linnreus)]. Pteropus jubatus Eschscholtz [= Acerodon jubatus juhatus (Eschscholtz)] . Rhinolophus philippinensis Waterhouse. Rhinolophus pygmaeus Waterhouse = Hipposideros pygmaeus (Waterhouse) .

* Cabrera, Journ. Mamm. 4 (1923) 89, has tried to show that the Linnrean species Simia syrichta is really a species of tarsier and not a monkey as many authors conclude. He has succeeded apparently in proving that the type is a tarsier but its specific identity, whethel' Tarsius carbonarius Heude, Tarsius fraterculus Miller, Tarsius philippensis Meyer, or an unnamed form actually inhabiting Luzon, there is no known method of determining.

PHILIPPINE LAND MAMMALS

Vespe?'tilio ?-v.fopictus Waterhouse [= Myotis rufopictus (Waterhouse)] , VespeTtilio mam'otarsus Waterhouse [ Myotis macro tarsus (Waterhouse)] . Vespe1,tilio meyeni Waterhouse [= Pipistrellus irTetitus (Cantor)]. Vespertilio eschscholtzii Waterhouse [= Miniopterus eschscholtzii (Waterhouse) ]. VespeTtilio tristis Waterhouse [ Miniopterus tristis (Wa1:.€rhouse»). Vespe?,tilio pellucidus Waterhouse [= Kerivoula pellucida (Waterhouse)] . Miniopte?'Us pusillus Dobson. Sciu1'Lt.'1 1Jhilippinensis Waterhouse. Mus (Phlmomys) cumingi Waterhouse [= Phlmomys cumingi (Waterhouse) ]. lIItls castaneus Waterhouse. Macacus cynomolgus val', cmningii Gray [= Pithecus philippinensis (1. Geoffroy) ] ,

Aside from the specimens that were made the types of new species, others were identified as known species. ,In several cases these have received specific definition. Thus, Dysopes tenuis (so identified by Waterhouse) may very probably be the earliest known specimen of ChfErephon luzonus Hollister. The Cuming collection contained the first European specimen of PhZreomys pallid'us Nehring, although it was not separated from the cumingi form. Several other known species were also collected by Cuming, in many cases the first Philippine specimens of the species to reach Europe. F. Jagor was in the Philippine Islands from 1859 to 1861 and obtained'a large number of animals among which were numerous mammals. Peters studied the collections and described the following as new: Vespe?·tilio (Kerivoula) jago?'ii Peters [= Kerivoula jagorii (Peters)] . EmballonU7:a discolor Peters [= Emballonwra alecto (Eydoux and Gervais) ]. Rhinolophus 1-ufus Peters (= Rhinolophus sub rufus Andersen), Rhinolophus arcuatus a1'cuatus (Peter,,), Phyllorhina co?~onata Peters [= Hipposider'os cOl'onatus (Peters)]. Phyllo?'hina ant?'icola Peters [= Hipposideros antricola (Peters)]. Phyllo')"'hina obsoum Peters [ = Hipposideros obscura (Peters)]. Pacltysoma (Ptenochi?'us) jagO?'ii Peters [= Ptenochirus jagoTii (Peters)], Pachysoma luzoniense Peters [= Cynopterus brachyotis brachyotis (Muller) ]. C?'ocidu?'a (P.) luzoniensis Peters [= Pachyura luzoniensis (Petel's)] .

INTRODUCTION

A. H. Everett, a noted British naturalist and collector, spent considerable time in the Philippines. While his chief interest was directed towards birds he collected numerous mammals, among which were the types of Rattus everetti (Gunther) and Thecu'fus pumilus Gunther. Several other known forms were also taken by Everett, including the following species: Felis minuta Temminck. Sci~拢?"us philippi.n ensis Waterhouse. Pithecus philippinensis (Geoifl'oy) (probably wrongly identified). Cynocephalus volans (Linnreus). Pteropus hypomelanus Temminck (probably P. h. cagayanus). Ace?路odon. jubatus (Eschscholtz) (probably A. j. j'L~batus). Hipposide?'os obscu?"t(s (Peters), Pachyu1"a l'Lt-zoniensis (Peters). Pa?'adoxu1'US philippiJnensis Jourdan.

Gii.nther published on this collection (see Bibliography). Prof. J. B. Steere, who collected in various parts of the Philippine Archipelago as early as 1887, obtained the types of the following: Sciu?'uS steerei Gunther. Sciurus mindanensis Sreere. Sciurus sama?'ensis Steere. Miniopterus paululus Hollister. Hipposideros bicolor (Temminck). Rhinolophus anderseni aequalis G. M. Allen. Probubalus mindorensis Steere (= Bubalus mindO?'ensis Heude). Hipposideros diadema griseus (Meyen). Pte?'opus vampyrus lanensis (Mearns). Taphozous philippinensis Waterhouse. Myotis 1'ufo-piotus (Waterhouse). Pipistrellus imb?'icatus (Horsfield).

Besides these a number of other species were collected, some twenty-five in all. The collection was badly scattered in various museums, and I have been unable to trace all his specimens. It is probable that a greater number of species was taken than is recorded here. The collections made by Steere were published on chiefly by A. Gunther, D. G. Elliot, N. Hollister, G. M. Allen, and by Steere himself. The collection made by John Whitehead was particularly important in that it included representatives of several new endemic genera. The collection contained the following new species, each representing a new genus: Celamomys silaceus (Thomas). Chrotomys whiteheadi Thomas.

PHILIPPINE LAND MAMMALS

Rhynchomys soricoides Thomas. Batomys granti Thomas. Carpomys melanurus Thomas. Carpomys phreurus Thomas. Crunomys fallaxThomas. Harpyionycteris whiteheadi Thomas.

Aside from the species listed above several new species belonging to known genera were found, as follows: Kerivoula whiteheadi Thomas. N annosciurus samaricu8 Thomas. Mus luzonicus Thomas [= Rattus luzonicus (Thomas)]. Mus mindorensis Thomas [ Rattus mindorensis (Thomas)]. Mus ephippium negrinus Thomas ( = Rattu8 negrinu8 Thomas).

Besides the new species, Whitehead obtained twenty-three other species, one of which, described by Meyer as "Phlreomys (?)" was made a new genus, Crateromys, by Thomas. This collection or at least a large part of it became the property of the Zoological Society of London and later of the British Museum of Natural History. Some of the forms are, I .believe, known only from the type series. A small collection was made by the Menage Expedition to the Philippines. I have been unable to trace this material, but some of it is at present in the museum of the University of Minnesota. The collection made by Maj. Edgar A. Mearns, Medical Corps, United States Army', shortl~ after American occupation of the Islands, and now in the United States National Museum, is undoubtedly the largest single collection. The collection was made possible largely through the interest and cooperation of Maj.-Gen. Leonard Wood. The collection totaled something over one thousand specimens, containing ninety-three species and subspecies, of which fifty-four were made types of new species and subspecies and eight species were made types of genera. Mearns's collections were from a very large number of localities and islands, among which perhaps the most important were Davao Province and Mount Apo; Zamboanga; Mount Bliss, Mount Malindang, northern Mindanao; Cagayan Sulu; Basilan; Mount Ha1con and Bulalacao, Mindoro; and Manila, Benguet, Ilocos Norte, Isabela, and Cagayan, on Luzon. Numerous other localities are represented. The new species from this large collection were treated chiefly in three papers. Mearns* described six new genera (Urogale,

* Proc.

U. S. Nat. Mus. 28 (1905) 425-460.

INTRODUCTION

Podogymnura, Bullimus, Tars O'YYI!!JS , Limnomys, and Apomys)

and the following new species and SUbspecies: Cynomolgus cagayanus Mearns [= Pithecus cagayanus (Mearns)]. Cynomolgus mindanensis Mearns [ = Pithecus mindanensis mindanensis (Mearns)]. Cynomolgus mindanensis apoensis Mearns [= Pithecus mindanensis apoensis (Mearns)]. Cynomolgus suluensis Mearns [ = Pithecus suluensis (Mearns)]. Pteropus lanensis Mearns [ = Pteropus vampyrus lanensis (Mearns) J. Pteropus cagayanus Mearns [= Pteropus hypomelanus cagayanus (Mearns)]. Urogale cylindrura Mearns. Podogymnura truei Mearns. Mus tagulayensis Mearns [= Rattus tagulayensis (Mearns)]. Mus albigularis Mearns [= Rattus albigularis (Mearns)]. Mus magnirostris Mearns [ = Rattus magnirostris (Mearns)]. Mus mindanensis Mearns [= Rattus mindanensis (Mearns) J. Mus zamboang:E Mearns [= Rattus zamboang:E (Mearns)]. Mus kelleri Mearns [ = Rattus kelleri (Mearns)]. Mus todayensis Mearns [ = Rattus todayensis (Mearns)]. Mus vulcani Mearns [= Rattus vulcani (Mearns)]. Mus vulcani apicis Mearns [= Rattus vulcani (Mearns)]. MU8 pant~rensis Mearns [ = Rattus pantarensis (Mearns)]. Mus commissariu8 Mearns. Bull;imus bagobU8' Mearns. Limnomys sibuanus Mearns. Tarsomys apoensis Mearns. Apomys hyloccetes Mearns. ApomJy8 petr83us Mearns. ApomY8 insignis Mearns.

The second paper was published by Gerrit S. Miller in 1910.* Two genera, ChilophyZla and Tryphomys, are described. The following species are described from the Mearns collection: Crocidura halconu8 Miller. Crocidura beatus Miller. Crocidura mindorus Miller. Crocidura grandis Miller. Pteropus pumilus Miller. Chilophylla hirsuta Miller. Taphozous pluto Miller. Epimys gala Miller [= Rattus gala (Miller)]. Tryphomus adustus Miller. Batomys dentatus Miller. Apomys bardu8 Miller. Apomys major Miller. Apomys musculus Miller.

* Proc.

U. S. Nat. Mus. 38 (1910) 391-404.

PHILIPPINE LAND MAMMALS

The follo wing species in the same paper are from the Bureau of Science collection, collected by R. C. McGregor and A. Celestino: Tupaia cuyon is Miller. Epimys tymnnus Miller [= Rattus tyrannus (Miller)]. T al'sius fTatercu lu s Miller.

Thâ&#x201A;Ź third paper published on the Mearns collection appeared in 1912 * by N. Hollister and included the following species: Chael'ephon luzom ls Hollister. Nannosciu1'u s surrutilus Hollister. Epimys coloratus Hollister [= Rattus coloratus (Hollister)]. Epimys mayom'cus Holliste r [= Rattus mayoni c'U.s (Hollister)] . Epimys vigoratus Hollister [= Rattus vigor atu8 (Hollister)]. Epimys basilanus Holliste r [= Rattus basilanus (Hollister)]. Epimys benguetensis Hollister [ = Rattu s benguetensis (Hollister)]. Limnomys rnearnsi Hollister. Limnomys picinus Hollister. Pithecus 1nindorus Hollister. R~(sa nigellus Hollister.

In t he same paper there also appeared descriptions of Taphonycteris capito Hollister, Epimys leucophmtus Hollister [ = Rattus leucophmtus (Hollister)], and Apomys micro don Hollister, collected by D. B. Mackie; Pachyura occultidens Hollister, collected by Dr. Carrol Fox; Miniopterus paululus Hollister, collected by J. B. Steere, and Epimys robiginosus Hollister [ = Rattus robiginosus (Hollister)] and Epimys ornatulus Hollister [= Rattus ornatulus (Hollister)], collected by R. C. McGr egor and A. Celestino. Hollister published descriptions of two new rats from the Mearns collection in 1911.t These were Epimys calcis Hollister [ = Rattus calcis (Hollister)] and Epimys querceti Hollister [ = Rattus querceti (Ifollister)]. In the same journal,t he likewise described t wo new bats from the collection. These were Pteropus balutus Hollister and Pteropus mearnsi Hollister. One bat, Rhinolophus in ops Andersen, from the collection was described by Knud Andersen. Several other smaller collections have reached the United States National Museum as follows: The George C. Lewis collection contained twenty-eight specimens and seven species. They were collected in central Luzon,

* Proc. t hoc.

U. S. Nat. Mus. 46 (1912) 299-341. BioI. Soc. Washington 24 (1911) 89-90. :j: hoc. BioI. Soc. Washington 26 (1913) 111-112.

INTRODUCTION

from Manila north to San Fernando, La Union. No new species were present. The Dr. Carrol Fox collection numbered twenty-six speCimenB with nine species represented. One new species, Pachyura occultidens Hollister, has been described. The collection was made in Panay and about Manila in 1912. The D. B. Mackie collection of twenty-six specimens made on Catanduanes in 1909, contained three new species, Taphonyctens capito Hollister, Ra.ttns leucopluetus Hollister, and Apomys mierodon Hollister. Six species are represented. The L. M. McCormick collection made in southern Luzon, chiefly in Camarines, contained twenty specimens and six species. One new species, Rhinolophu,.s V iTgO Andersen, was described in 1905 by Knud Andersen.* Dr. Paul Bartsch made a collection of bats from the caves at Montalban, near Manila. Fifty-seven specimens were taken, representing seven species. One new species, Eonyeteris robustus Miller, was described. Dr. Robert Grubb's collection contained thirty-five specimens representing four species. The collection is from Iligan, Mindanao. Wm. D. Carpenter collected ten bats, representing two species, at Malabon, near Manila. Dr. J. C. Le Hardy collected ten specimens in Min'd oro; among them were specimens of the Mindoro deer and the timarau. A few other persons have collected specimens now in the United States National Museum, in no case, however, more than four. No new species is represented in these. They are from widespread localities. The Bureau of Science collection deposited in the United States National Museum t consisted originally of ninety-eight specimens, representing twenty-six species. The collections were made for the most part by Richard C. McGregor and Andres Celestino. Types of Rattus robiginosus (Hollister), Rattus ornatuLus (Hollister), Tupaia cuyonis Miller, R.attus tyrannus (Miller), Seiuropterus erinitus Hollister, and Tarsius frateremus Miller are in this collection. It was made during 19031910, in numerous localities. '" Proc. Zool. Soc. London (1905).

t A few . of these specimens were returned Manila in 1922.

to the Bureau of Science in

PHILIPPINE LAND MAMMALS ANNOTATED BIBLIOGRAPHY

In preparing this bibliography I have found the bibliographical list published by Hollister * especially helpful. Several of the papers listed there are omitted from this list since they are not pertinent, and a number of others, not included, or published since this work appeared, are here included. Certain of the papers 'I have not been able to examine, but most of the rarer works have been made available through the courtesy of The John Crerar Library, of Chicago. ALLEN, GROVER M. Bats from Palawan, Philippine Islands. Occasional Papers Mus. Zool. Univ. Michigan No. 110 (February 25, 1922) 1-5. This paper is based on a part of the Beal-Steere Expedition collection preserved in the University of Michigan Museum of Zoology. The following known species are listed: Pteropus vampyrus lanensis Mearns, Taphozous philippinensis Waterhouse, Hipposideros bicolor (Temminck), Hipposideros diadema griseus (Meyen), Myotis rufopictU8 (Waterhouse), Pipistrellus imbricatus (Horsfield). A single form is described as a new subspecies, Rhinolophus ande?'seni aequalis Allen. ALLEN, J. A. Mammals from Palawan Island. Bull. Am. Mus. Nat. Hist. 28 (1910) 13-17. (January 29.) Contains type descriptions of Mus luteivent1'is ( Rattus luteivent1-1.S) , Ar,ctitis (sic) whitei (= Arctictis whitei) , and Mungos palawanus. The following species are likewise recorded: Manis sp. (= Manis javanica) , Sus ahrenobarbus Huet, Muntiacus nigricans (= Tragulus nigricans Thomas), Sciurus stee1'i Gunther, Sciuropterus nigripes Thomas, Felis minuta Temminck, Viverra t'lCngalunga (sic) Gray ( = Viverr,a tangalunga Gray), Paradoxurus philippinensis Jourdan, Aonyx cinerea (Illiger), Tupaia ferruginea palawanensis Thomas ( Tupaia palawanensis Thomas). This list represents the collection of Col. John R. White, Philippine Constabulary. The collection contained thirtythree specimens, all collected at Iwahig, Palawan, except a single specimen of Tragulus that came from Balabac Island.

ANDERSEN, KNUD. On the bats of the Rhinolophus philippinensis group with descriptions of five new species. Ann. & Mag. Nat. Hist. VII 16 (1905) 243-257. A synopsis of the Rhinolophus phillippinensis group. ANDERSEN, KNUD. On the bats of the Rhinolophus arcuatus group, with descriptions of five new forms. Ann. & Mag. Nat. Hist. VII 16 (1905) 281-288. The type descriptions of Rhinolophus inops, Rhinolophus subrufus, and Rhinolophus arcuatus exiguus appear. ANDERSEN, KNUD. On the bats of the Rhinolophus macrotis group, with descriptions of two new forms. Ann. & Mag. Nat. Hist. VII 16 (1905) 289-292. The type description of Rhinolophus hirsutus appears.

* Philip.

Journ. Sci. ยง D 7 (1912) 52-64.

INTRODUCTION

ANDERSEN, KNUD. On some bats of the genus Rhinolophus, with remarks on their mutual affinities, and descriptions of twenty-six new forms. Proc. Zool. Soc. London (1905) 75-145, pIs. 3, 4. (October, 1905.) The original description of Rhinolophus virgo appears. ANDERSEN, KNUD. On Ripposideros diadema and its closest allies. Ann. & Mag. Nat. Rist. VII 16 (1905) 497-507. Meyen's name RhiJnorLophus g'riseus is revived as a subspecies, Hipposideros diadema griseus. ANDERSEN, KNUD. A list of the species and subspecies of the genus Rhinolophus with some notes on their geographical distribution. Ann. & Mag. Nat. Rist. VII 16 (1905) 648-662. A review of the genus Rhinolophus is given. Several Philippine forms are mentioned. ANDERSEN, KNUD. On Pterocyon, Rousettus, and Myonycteris. Ann. & Mag. Nat. Rist. VII 19 (1907) 501-515. Revision of the genus Roussettus. The Philippine form is recorded. ANDERSEN, KNUD. Twenty new forms of Pteropus. Ann. & Mag. Nat. Rist. VIn 2 (1908) 361-370. The original description of Pte?'oP~tS specwsus is given. ANDERSEN, KNUD. Notes on the genus Acerodon, with a synopsis of its species and subspecies, and descriptions of four new forms. Ann. & Mag. Nat. Rist. VIn 3 (1909) 21-29. (January.) Type description of Acerodon jttbatu8 mindanensis. ANDERSEN, KNUD. On the characters and affinities of "Desmalopex" and Pteralopex. Ann. & Mag. Nat. Rist. VIII 3 (1909) 213-222. (February.) Desmalopex Miller is regarded a synonym of Pteropus. Presumes that the type of Gray's Pte1'opus chinensis came from Luzon. ANDERSEN, KNUD. On the fruit-bats of the genus Dobsonia. Nat. Rist. VIn 4 (1909) 528-533. (December.) A review of the genus Dobsonia.

Ann. & Mag.

ANDERSEN, KNUD. Six new fruit bats of the genera Macroglossus and Syconycteris. Ann. & Mag. Nat. Rist. VIII 7 (1911) 641-643. (June.) A review of the genus Macroglossus. ANDERSEN, KNUD. Catalogue of the Chiroptera in the Collection of the British Museum 2d ed. London 1 (1912) I-CI, 1-854, numerous text figures. This is an extraordinary monograph on the Megachiroptera. It includes all known forms of this group to that date and gives detailed descriptions. Pteropus hypomelanus tomesi is first recorded from the Philippine Islands. Two specimens collected by A. Everett, 1893 are listed from Sibutu and one from Lamboyan, off southern Palawan, collected by Douglas Cator, August or September, 1894. ANDERSEN, KNUD, and R. C. WROUGHTON. On the bats of the family Megadermatidre. Ann. & Mag. Nat. Rist. VII 19 (1907) 129-145. A monographic treatment of the genus Megaderma and other bats of the same family.

PH.ILIPPINE LAND MAMMALS

BALFOUR, EDWARD. Cyclopredia of India and of Eastern and Southern Asia, Commercial, Industrial, and Scientific. Madras, 2d ed. (18711873) 1-5. Some notes on Philippine mammals. Of no special systematic importance. BOURNS, FRANK S., and DEAN C. WORCESTER. Preliminary Notes on the Birds and Mammals Collected by the Menage Scientific Expedition to the Philippine Islands. Occasional Papers Minn. Acad. Nat. Sci. 1 (1894) 1-64. Pages 61 to 64 contain a list of mammals which is, for the most part, very untrustworthy since numerous forms are identified as extra-Philippine species. Macacus philippinensis Geoffroy (= PithecUS species) ("Occurs on every island visited by us"); Nycticebu8 tardigmdus Linnreus [=Nycticebus menagensis (Leydekker)] Bongao, Tawitawi; Tarsius spect1路um Pallas (=Tarsius philippinensis Meyer) Samar; T~~paia javanica Horsf. = (Tupwia species); Galeopithecus philippinensis W aier house [= Cynocephalus volans (Linnreus)] Basilan, Mindanao, Samar; Mydaus marchei Ruet, Palawan and Calamianes (the latter record probably Mydaus schadenbergii Jentink); Arctitis binturong Raffles [=Arctictus whitei (Allen)] Palawan; ParadoxU1'US philippinensis Jourdan (= Paradoxu?'us species) Mindoro, Panay, Negros, Mindanao, Basilan, Palawan; Viverra tangalunga Gray, Siquijor, Panay, Mindoro, Mindan3l0, Palawan, Calamianes; Felis bengalensis Kerr (= Felis minuta Temminck) Panay, Negros, Cebu; Hystrix pumila Gunther [= Thecurus pumUus (Gunther)] Palawan, Calamianes; Sciuropterus nigripes Thomas, Palawan; Sciurus steerii Gunther, Palawan; Sciurus samarensis Steere, Samar; Sciurus rrvindanensis Steere, Mindanao and Basilan; Sciurus coccinus Thomas (= ? Sciurus p1dlippinensis Waterhouse), Mindanao, Basilan; Bubalus kerabaw ferus Nehring (= Bubalus species) Mindoro, Calamianes, Masbate, Luzon, Negros, Mindanao; Bubalus mindorensis Reude, Mindoro; Tragulus javanicus Gmelin (= Tragulus nigricans Thomas) Balabac; Manis javanica Desmarest, Palawan, Calamianes; Sus ahamobarbus Ruet, Palawan, Calamianes. Other species were collected and remain unidentified. These consist of deer and bats and a few other mammals, both skeletal and alcoholic preparations. BROOKE, SIR VICTOR. On the deer of the Philippine Islands, with the description of a new species. Proc. Zool. Soc. London (1877) 51-60, pIs. 8-10. Original description of Cervus nigricans (= Rusa nigricans), Cervus alfredi [=Rusa alfredi (Sclater)], Cervus philippinus (=Rusa phili'P'Pinus (Smith)], and Cervus mariannus [= Rusa philippinus (Smith)] are also listed. CABRERA, ANGEL. Bol. Real Soc. Espanola Rist. Nat. (1901) 372, fig. 2. (December.) Describes Capromys elegans ( =PhlCl3omys elegans) from an old specimen in the Madrid Museum, presumably from Cuba or a near-by island.

INTRODUCTION

CABRERA, ANGEL. Trab. Mus. Cien. Nat. No.3 (1912) 20. Establishes the fact that the type locality of Capromys elegans is the Philippine Islands and places it in the genus Phlmomys. CABRERA, ANGEL. Sobre los Loris, y en especial sobre la forma Filipina. Bol. Real Soc. Espanola Hist. Nat. 8 (1908) 135-139. Cabrera proposes a name Nycticebus philippinus for the Philippine slow lemur named by Lydekker Lemur ? menagensis. CABRERA, ANGEL. Un nuevo "Rhinolophus" filipino. Bol. Real Soc. Espanola Hist. Nat. 9 (1909) 304-306. Gives the type description of Rhinolophus anderseni. CABRERA, ANGEL. On the identification of Simia syrichta Linnreus. â&#x20AC;˘J ourn. Mammalogy 4 (1923) 89-91. Cabrera believes Simia syrichta to be Tarsius philippensis Meyer. He has apparently proved that the genus is Tarsius, but it may be any one of the three species known from the Islands. CAMEL, GEORGE JOSEPH. De Quadrupedibus Philippensibus. Philosoph. Trans. London 21 (1699) 2197-2204. This work contains a list of Philippine mammals with short descriptions in some cases. Some t hirty-four forms are given, including the elephant, which he states is abundant in Jolo. Much of the paper is devoted to Bezoar stones. In the list are 3 bats, 3 monkeys, 1 deer, 3 wild boars. Camel (Kamel) was a German Jesuit who died in Manila 1702. CANTOR, THEODORE. General features of Chusan, with remarks on the flora and fauna of that island. Animals observed at Chusan. Ann. & Mag. Nat. Hist. 9 (1842) 481-493. Gives original description of Pipistrellus irTetitus from Chusan Island, China. CUVIER, F. Des .Dents de Mammiferes, Considerees comme Characteres Zoologiques 1-55 (1825) 1-259. Diagnosis of Sciuropterus. DELGADO, JUAN J. Historia general sacro-profano, politico y natural de las Islas del poniente llamadas Filipinas. Manila (1892). This is a work only partly prepared by Delgado. His writing was done between 1711and 1744, but the exact date is probably not known. It was not published until 1892. He is known to have been in Samar, Cebu, Mindanao, Leyte, Bohol, and Luzon. He lived in the Islands about thirty-four years and died in Mexico. It is generally believed that the parts dealing with natural history is largely the work of two Jesuits, P. Clain and G. Camel. The book contains descriptions of several Philippine mammals. DESMAREST, A. G. Mammalogie ou Description des Especes de Mammiferes. Seconde partie. Paris (1822). Original descriptions of Manis javanica and Mustela nudipes. 269453-8

PHILIPPINE LAND MAMMALS

DOBSON, G. E. Notes on some bats c.olIected by Captain W. G. Murray in the northwestern Himalaya, with description of new species. Proc. Asiatic Soc. Bengal (1872) 208-210. Original description of Murina cyclotis from Darjeeling, northeastern Bengal. DOBSON, G. E. Descriptions of new or little-known species of bats of the genus Vesperugo. Proc. Zool. Soc. London (1875) 470-474. Original description of Vesperugo tylopus (= Glisehropus tylo'/YUs). DOBSON, G. E. Monograph of the Asiatic Chiroptera and Catalogue of the Species of Bats in the Collection of the Indian Mus,e um, Calcutta. London, Taylor and Francis (1876). The f ollowing species are attributed to the Philippines: Pteropu8 ju batus Eschscholtz [= Acerodon jubatus (Eschscholtz)]; Cynopierus 17w1'ginatus Geoffroy ( = Cynopterus braehyotis braehyotis (S. Muller) ; Cymopte1'us (Ptenoehirus) jagori Peters [= PtenocJvir1,~s jago1'i (Pet er s ) J; Cy on y eteris amplexicaudata Geoffroy [= Rousettus amIJlexicaudatus (Geoffroy)]; Rhinolophus luetus Temminck (= ? Rhinoloph us philippinensis Water.house) ; Rhinolophus ruf~[s Peters (= RhinolOPM~ s~~b 1'ufus Anaersen); Phyl101-hina diadema Geoffroy [= Hipposideros diadema griseus (Meyen)]; Phyllorhina pygmaea Waterhouse [= Hipposideros 'PYg'frW3US (Waterhouse)]; Phyllorhina bicolor Temminck [= Hipposid&os bicolorr (Temminck)]; Phyllorhina eo?'onata Peter s [= Hipposiderros ooronatfb (Peters)]; P11l!Jllorrhina obsm~ra Peters (= Hipposideros obso:urus Peters); Megaderma spasma Linnreus (= Megaderma spasma spasma Linnams,); Vesperugo (Tylonycteris) paehypus Temminck [= Tylonyot'erys pachypus (Temminck)]; Seotophilus temminokii (Horsfield); Vespertilio capacei'Yllii Bonaparte (= ?); Vesperrtilio maerotarsu$ Waterhouse [= Myotis macro tarsus (Waterhouse)]; KerivouZa peUucida (Waterhouse); Kerivoula jagorii (Peters); Miniop't erus sehreiberrsii Naterrer [= Miniop,te1"U8 eschseholtzii (Waterhouse)]; Miniopterus tristis (Waterhouse); Emballonura montioola Temminck (= Emballonura aleeto Eydoux); Tarphozous m elanopogon Temminck (=?); and Taphozous pJvilippinensis Waterhouse. DOBSON, G. E. Description of new species of Crocidura .. Ann. & Mag. Nat. Hist. VI 6 (1890) 4'94-497. Croeidura grayi is described from a specimen in the Cuming collection. The exact type locality is not given. ELERA, CASTO DE. CataJogo Sist.ematico de Toda la Fauna de Filipinas Conocido . hasta el Presente, y a la vez el de la Coleccion Zoo16gica del Museo de P. P. Dominicos del Colegio-Universidad de Santo Tomas de Manila, Escrito con Motivo de la Exposicion Regional Filipina. Manila 1 (1895). Vertebrados, 1-51 and 623, Mammiferos. A list of species of animals known or reported from the Philippines. A few new names, (nomina nuda) occur. Many extra-Philippine forms ar-e listed as being from the Islands. Most of the forms listed purport to be present in the Universidad de Santo Tomas. At present the specimens in many cases are no longeT extant.

INTRODUCTION

ELERA, CASTO DE. Contribucion a la Fauna Filipina. Manila (1915) 1-284. Numerous plates. This work, which treats only of mammals, was prepared by the author prior to his death August 29, 1903. It was edited and published by Fr. Florencio Llanos, direcior of the Museum of the University of Santo Tomas in 1915. ELLIOT, D. G. On sundry collections of mammals received by the Field Columbian Museum from different localities, with descriptions of supposed new species and sub-species. Field Columbian Museum Pub!. No. 11 Zoo!. Ser. 1 'No.3 (1896) 67-82, pIs. 6-13. A report on mammals obtained by the Menage Expedition to Borneo and the Philippines; the following are listed: Sus aluenobarbus Huet, Palawan: Cel'vus? [= Rusa calam路ianensis (Heude)], Culion: Nycticebus tardigrad'us (Linnreus [ = Nycticebus menagensis (Lydekker)], Bongao; Scotophilus temmincki Horsfield, Panay. Elliot recognizes that the deer listed is probably already described by Hende. It is illustrated by figures of skulls of both males and females. From the Steere expedition the following species are described or listed; Cervus stee?'ei [= Rusa nigricans (Brooke)] Basilan; Pterop us ccw'i-nuchalis (=Ace1'odon ju.batus j1tbatu.,s Eschscholt.z), Leyte, and Pt.e-r路opus lucifer [= Ace1'odon l~wife1' (Elliot)] are described as new. The following are listed: Trag~ilus nig1'icans Thomas, Balabac: Pte1'opus jubatus (=' Acerodon j'Urbatns jub'atus Eschscholtz), Concepcion, Panay; Pteropus edu~is Geoffroy [:::: Pteropus va?npyrus lanensis (Mearns) ], Leyte; Pte1"opus hypomelanus Temminck [= Ptel'opus hypomelanus cagayanus (Mearns)], Panay and Guimaras; Cynonycteris amplexicaudatus Geoffroy [= Rousettns a?nplexicalldatus (Geoffroy)], Negros; Phyllorliina diadema Geoffroy [ = Hipposide?'os diadema griseus (Meyen)], Guimaras. Plates are given of tne head and skulls, male and female of Rusa nigricans, skulls of Acerodon jubatus jubatus, Acerodon lucife?', and Tragul~iS nig1路icans. ELLIOT, D. G. Remarks upon two species of deer of the genus Cervus from the Philippine Archipelago. Field Columbian Museum Publ. 20 Zoo!. Ser. 1 No.7 (1897) 157-159. Ce1'vus culionensis is proposed for a species of deer described from Cui ion in the preceding paper after he has examined Heude's paper, Memoires concernant I'Histoire N aturelle de l'Empire Chinois 2 pt. 1 (1888) 1-50, pIs. 1-19. Elliot is convinced that his Cervus steerei is distinct after consideration of Rusa philippinus and R. nig?路icans. ELLIOT, D. G. A Catalog of the Collection of Mammal" in the Field Columbian Museum. Field Columbian Mus. Pub!. 115 Zool. Ser. 8 (1907) 1-694. Numerous Philippine specimens are listed, chiefly from the J. B. Steere collection, a few from the Menage Scientific Expedition to the Philippines, and one collected by Everett. He also lists as present Sus ce:ebensis and Macacus fascicularis Raffles. These probably are incorrectly identified. Many of the records of the forms listed have appeared in other papers by Elliot, Steere, and Bourns and Worcester.

PHILIPPINE LAND MAMMALS

ELLIOT, D. G. A Review of the Primates. Monograph 1 Am. Mus. Nat. Hist. 1 (1913) I-CXXVII, 1-317, I-XXXVIII, numerous plates; 2 (1913) I-XVIII, 1-382, I-XXVI, numerous plates; 3 (1913) I-XIV, 1-262, I-CLXVII, numerous plates. In volume 1 are redescribed Tarsius philippensis Meyer and Tarsius ft'ate?'culus Miller. Apparently he has overlooked Heude's species Tarsi~~s carbonarius Heude. In volume 2, four forms of Pithem~s are listed, philippinensis (Geoff;roy), philippinensis apmnsis (Mearns), cagatyanus (Mearns) and suluensis (Mearns). Elliot regards apmnsis as of doubtful validity, and makes ?nindanensis a synonym of philippinensis. He feels that cagayanus and suluensis are based on insufficient material. In volume 3 there are no Philippine forms. ERXLEBEN, J. C. P. Systema Regni Animalis. Classis I, Mammalia. Lipsire (1777). Contains the original description of Rattus norvegicus. ESCHSCHOLTZ, F. ZoOlogischer Atlas, enthaltend Abbildungen und Beschreibungen Neuer Thierarte:n, wahrend des Flottencapitans von Kotzebue Zweiter Reise urn diâ&#x201A;Ź Welt, auf der Russisch-Kaiserlichen Kriegsschaluppe Predpriaetre in den Jahren 1823-1826. Berlin (1831) 1-19. Contains the type description of Acerodon jubatus. EVERETT, A. H. Remarks on the zoogeographical relationships of the Island of Palawan and some adjacent islands. Proc. Zool. Soc. London (1889) 220~228, 1 map. Gives a list of the known species of mammals common to Borneo and Palawan. Discusses the probable geological relationship. EVERETT, A. H. A nominal list of mammals inhabiting the Bornean group of islands. Proc. Zool. Soc. London (1893) 492-496. (May 16.) Notes on the Palawan fauna. Palawan species in the check lists are preceded by two asterisks. EYDOUX, FORTUNE, and PAUL GERVAIS. Zoologie. Voyage autour du monde par les Mers de I'Inde et de Chine execute sur la corvette de l'Etat la Favorite pendant les annees 1830, 1831, et 1832, sous Ie commandement de M. Laplace Captaine de Fregate. Paris (1839). Contains original descriptions of Rhinolophus luctus val'. 'l'Ufa (= ? Rhinolophus subrufus Andersen) Vespe?,tilio (Nycticeus) alecto [ = En~ballomLTa alec to alecto (Eydoux and Gervais)]. GEOFFROY SAINT-HILAIRE, E. Description des Roussettes et des Cephalotes, deux nouveaux genres de la Famille des Chauve-souris. Ann. duo Mus. d'Hist. Nat. 15 (1810) 86-108, pIs. 4-7. Contains original descriptions of Pteropus amplexicaudatus (= Rousettus amplexicaudatus) , Timor; and Cephalotes peroni [= Dobsonia peToni (E. Geoffroy)], Timor. GEOFFROY SAINT-HILAIRE, E. Sur un Genre de Chauve-souris, sous Ie nom de Rhinolophes. Ann. du Mus. d'Hist. Nat. 20 (1813) 254-266, pIs. 5-6. Contains the type description of Rhinolophus diadema (= Hippo8?" deros diadema).

INTRODUCTION

GEOFFROY SAINT-HILAIRE, ISIDORE. Description des Mammiferes nouveaux ou imparfaitement connus de la Collection du Musee d'Histoire Naturelle, et Remarques sur la Classification et les Caracteres des Mammiferes. Premier Memoire, Famille des Singes. Archive du Mus. d'Hist. Nat. 2 1841 (1843) 485-592, pIs. 29-34. Contains the original description of Macacus philippinensis (= Pithecus philippinensis).

GEOFFROY SAINT-HILAIRE, ISIDORE. Catalogue de la Collection des Mammiferes, de la Collection des Oiseaux â&#x201A;Źt des Collections annexes. Premiere Partie. Mammiferes, Catalogue des Primates. Paris (1851). Contains the original description of Macacus palpebrosus (= Pitke-

cus philippinensis). GERVAIS, PAUL. Voyage de la Bonite, Zoologie 1. Paris (1841) 43-50, pI. 8. Discusses a form of Phlmom;ys and gives a picture of the same. GORGORZA Y GONZALEZ, JOSE. Datos para la fauna filipina. There are 35 species of mammals listed in this work. in synonymies.)

Vertebrados. (Not quoted

GRAY, J. E. On the family of Viverridre and its generic subdivisions; with an enumeration of the species of Paradoxurus and characters of 'sever~l new ones. Proc. Zool. Soc. London (1832) 63-68. Contains the original description of Viverra tangalunga. GRAY, J. E. List of Mammals in the British Museum (1843) 55. A specimen of Paradoxurus is listed under the nomen nudum ot Paradoxurus zelanica. The specimen is probably Pa?'adoxurus minax Thomas. It is from Camiguin Island, P. 1. Other Philippine forms are listed. GRAY, J. E. Catalogue of Monkeys, Lemurs, and Fruit-eating Bats in the Collection of the British Museum. London (1870). This contains the original descriptions of Eleuther'wra philippinensis [= Roussettus amplexicaudatus (Geoffroy) ], Manila; Cynopterus rrw.rginatus var. nigrescens [= Thoopte?'Us nigrescens (Gray)], Morty Island, Malay Archipelago; Macacus cynomolg'us var. cumingii (=? Pithecus phi[;ippinensis Geoffroy), Philippine Islands. Certain other forms are listed from the Philippines. GUNTHER, ALBERT. Report on some of the additions to the collection ot Mammalia in the British Museum. Proc. Zool. Soc. London (1876) 735-36. (On a collection from the Philippine Islands.) Lists 111acacus philippinensis Geoffroy [=Pithecus phil1'ppinensis (Geoffroy)], Majayjay, Luzon; Galeopithecus phibippinensis Waterhouse [= Cynocephalus volans (Linnreus)], Bohol; Pteropus jubatus Temminck, Mindanao; Tupaia javanica Horsfield, Palawan; Viverra tangalunga Gray, Panay; Sciurus exilis S. Milller (= ?); SCiU1'US mindanensis Steere, Basilan ("Also found in Mindanao"); Sciurus steere'i is described as new from Balabac; Tupaia javanica ( = T. palawanensis Thomas).

PHILIPPINE LAND MAMMALS

GUNTHER, ALBERT. List of the mammals, reptiles, and batrachians sent by Mr. Everett from the Philippine Islands. Proc. Zool. Soc. London (1879) 74-79. (Mammals 74-75.) The following species are listed: Macacus philippinensis Geoffroy [= ? Pithecus rnindanensis (Mearns)], from Butuan andSurigao; Galeopithecus philippinensis Water house [ = Cynocephalus volans (Linnreus)], Surigao and Dinagat Island; Pte1"oprus edulis Geoffroy (= ?), Dinagat and Rasol (near Dinagat); Pteropus hypomelanus Temminck [=Pteropus hypomelanus cagayanus (Mearns)], Surigao and Dinagat; Pteropus J'ubatus Eschscholtz (= Ace?"odon iubatus mindanensis Andersen), Dinagat, S. Leyte, and S. Negros; Phyllorhina obscura Peters [= Hipposide'f'os obscurus (Peters)]; Crocidura luzoniensis Peters [= Pachyura luzoniensis (Peters)], Cebu, also Luzon, Manila; Felis minuta Temminck, S. Negros; Paradoxurus philippinensis Jourdan, North Mindanao (also a record for Manila, Luzon; Macroxus philippinensis Waterhouse, Placer, northeastern Mindanao. Mus everetti [= Rattus everetti (Gunther)] is described as new. Most of the specimens were collected in extreme northeastern Mindanao and Dinagat Island. GUNTHER, ALBERT. Description of a new species of porcupine from the Philippine Islands. Ann. & Mag. Nat. Hist. V 4 (1879) 106-107. This contains the original description of Hystrix pumila (= Thecurus pumilus) from a specimen collected at Puerto Princesa, Palawan, by Mr. A. Everett. HEUDE, P. M. Gatal. des Cerfs Tachetes (1885) 1. Describes C (ervus) soloensis later described under the name Sikelaphis soloensis (vide infra). I have not seen this work. HmJDEi, P. M. Cerfs des Philippines et de l'Indo-Chine. Mem. Hist. Nat. Emp. Chinois 2 pt. 1 (1888) 1-50 pIs. 0-19. This paper contains the original descriptions of Hyelaphus calamianensis (= Rusa calamianensis); U ssa c?'assicornis; U ssa gorrichanus; USsa barandanus (= Rusa barandanus); Ussa francianus (= Russa francianus); U ssa nublanus; U ssa tuasoninus; U ssa spatharius; U ssa ralJ'l'1J.O'smnus; U ssa amlirosianus; U ssa macarianus; U ssa elorzanus,. U ssa garcianus; U ssa guidoteanus; U ssa rosarianus; U ssa maraisianus,. U ssa dailliardianus; U ssa roxasianus; U ssa microdontus; Us sa brachyceros; Ussa corteanus; Ussa rubiginosus; Ussa hipolitianus; U ssa chrysotrichos; U ssa gueva?"anus; U ssa marzaninus; U ssa longicuspis,. U ssa gonzalirnus; U ssa telesforianus; U ssa verzosanus; M elanaxis basilanensis (= Rusa basilane,n sis); M elanaxis masbatensis; Melanaxis breviceps; and Melanaxis elegans. Most of these names probably rep,r esent the individual variation of a single species and this species not improbably Rusa philipp1tnus (Smith). One copy of this work I have examined was one acquired by the Bureau of , Science at my request but the plates dealing with Philippine deer had been removed by the Fathers at the Zi Ka Wei Museum presumably (for the purpose of) suppressing them. Since no less than 24 of the presumed species (4 others in another paper making

INTRODUCTION

28 in all) are from central southern Luzon all within a radius of 150 miles of Manila, this is not surprising. HEUDE, P. M. Note sur Ie Petit BufHe Sauvage de PIle de Mindoro (Philippines). Mem. Hist. Nat. Emp. Chinois 2 (1888) 50-51. (August.) Contains the type description of BubaltÂŁs mindorensis. HEUDE, P. M. Etudes Odontologiques Part I. Chap. 1. Mem. Hist. Nat. Emp. Chinois 2 (1892) 65-82. Contains on page 67 the type description of Sus inconstans. HEUDE, P. M. Etude sur les Suilliens. Chap. 11. Mem. Hist. Nat. Emp. Chinois 2 (1892) 85-115 pIs. 19A-29C. This paper contains original descriptions of Sus miinutus, Laguna to Tarlac, Luzon ?; Sus calarnianensis, Calamianes Islands; Sus ejJren'us, Jala JaIa, Laguna de Bay, Luzon; Sus micTotis, Jala Jala, Laguna de Bay, Luzon; Sus arietinus, Manila ?; Sus frenatus, Jala Jala, Laguna de Bay, Luzon; Sus cebifrons, Cebu and Masbate. Hollister places Sus a1'ietanus, S. ejJ?'e?ius, S. microtis, and S. frenatus as synonyms of Sus philippensis Nehring. HEUDE, P. M. Catalogue Revise des Cerfs Tachetes (Sika) de la Chine Centrale. Mem. Hist. Nat. Emp. Chinois 2 pt. 3 (1894) 146-163. C (ervus) soloensis is described under the name Sikelaphus soloensis (p. 147). HEUDE, P. M, Etudes Odo'ntologiques Part I, Chap. IV. Mem. Hist. Nat. Emp. Chinois 2 (1894) 170-211. Contains the original description of Bubalus ?nainitensis. HEUDE, P. M. Premiere Revision du Genre Ussa H. et Rectification de nomenclature. Mem. Hist. Nat. Emp. Chinois 4 pt. 3 (1899) 134-141. Contains the original descriptions of Ussa baryce?'os, Laguna and Batangas, Luzon; Ussa- cinereus, Cebu Island; Ussa villime1<ia,nus, San Miguel de Murcia, Tarlac, Luzon; Ussa michrelinus, San Miguel de Murcia, Tarlac, Luzon; Ussa vidalinus, San Miguel de Murcia, Tarlac, Luzon; Us sa atheneensis, Luzon. These names are probably referable to other known species but which ones it is difficult to say. See note under Heude, 1888. HEUDE, P. M. Etudes Odontologique. Part 4 Chap. I Lemuriens, Tarsiens, Galeopitheciens. Mem. Hist. Nat. Emp. Chinois 4 (1899) 155172. Contains the type description of Tarsius carbonariu8. HOFFMAN, B. tiber Saugethiere aus dem Ostindischen Archipel. Abhand. und Berichte des Konig!. Zool. Mus. Dresden No.3 (1886-1887) 1-29. Notes on rats, bats, and buffaloes from the Philippines. HOLLISTER, N. Two new Species of Epimys from Luzon. Proc. BioI. Soc. Washington 24 (1911) 89-90. (May 15.) Describes EpimY8 calcis ( = Rattus calcis) and Epimys querceti (= Rattus que1'ceti) from Benguet Province, Luzon, collected by Dr. E. A. Mearns.

PHILJPPINE LAND MAMMALS

HOLLISTER, N. Description of a new' Philippine flying-squirrel. Proc. BioI. Soc. Washing1;on 24 (1911) 185-186. (June 23.) Contains the original description of Sciuropterus crinitus collected in Basilan by R. C. McGregor and A. Celestino. HOLLISTER, N. The generic name of the African buffalo. Proc. BioI. Soc. Washington 24 (1911) 191-194. (June 23.) Regards the generic name Bubalus as applying only to Indian and Malayan buffaloes. He discusses Bubalus 'l'YUindo7'ensis Heude and regards 'it as congeneric with other oriental buffaloes. HOLLISTER, N. A list of the mammals of the Philippine Islands exclusive of the Cetacea. Philip. Journ. Sci. ยง D 7 (1912) 1-64. This is the most important work on Philippine mammals to its date of issue. It consists of an annotated list giving all apparently valid species with type localities and short diagnostic characters of genera. It likewise includes a list of erroneously credited species and a tabulation of type localities. A relatively complete bibliography is appended. No new species are described but many new combinations appear. HOLLISTER, N. Two new Philippine fruit bats. Proc. BioI. Soc. Washington 26 (1913) 111-112. (May 3.) Pteropus balutus, from Balut Island, Sarangani Group, Mindanao, and Pteropus mearnsi, from Isabela, Basilan, are described from the Mearns collection. HOLLISTIDR, N. A review of the Philippine land mammals in the United States National Museum. Proc. U. S. Nat. Mus. 46 (1913) 299-341, pIs. 27-29. This very important paper lists 133 species and subspecies of mammals represented in the United States National Museum by 1,454 specimens. Eighteen new species are described as follows: Pachyura occultidens, Iloilo, Panay; Taphonycteris capito, Pandan, Catanduanes; Miniopter~Ls paululus, Guimaras; Chaerephon luzonus, Cagayan, Cagayan, Luzon; Nannosciurus surrutilus, Mount Bliss, Mindanao; Epimys coloratus (= Rattus coloratus) , Isabela, Basilan; Epimys robiginosus (= Rattus r,o biginosus) , Cagayancillo, Cagayan Islands; Epimys mayonic~Ls (= Rattus mayonicus) , Mayon Volcano, Albay, Luzon; Epimys leucophmatus (= Rattus leucophmatus,) , Bagamanoc, Catanduanes i Epimys vigoratus ( = Rattus vigoratus), Mount Halcon, Mindoro; Epimys basilanus (= Rattus basilanus) , Basilan; Epimys ornatulus ( = Rattus ornatulus) , Cagayancillo, Cagayan Islands; Epimys bemguetensis ( = Rattus benguetenS1.:s) , Baguio, Benguet. Luzon; Limnomys mearnsi, Grand Malindang Peak, Mindanao; Limr nomys picinus, Mount Halcon, Mindoro; Apomys mic?'odon, Biga, Catanduanes; Pithecus mindorus, Alag River, Mindoro; Rusa nigellus, Mount Malindang, Mindanao. Of the eighteen forms described E. A. Mearns collected 11 species; McGregor and Celestino, 2; D. B. Mackie, 3; Dr. C. Fox, 1; and J. B. Steere, 1. HORSFIELD, THOMAS. Zoological Researches in Java and the -Neighboring Islands. London (1824). (Pages and plates not numbered.)

INTRODUCTION

This work contains original descriptions of Vespertilio hardwickii (=Kerivoula hardwickii) ; Vespe?,tilio imb?'icatus (=Pipist?>ellus imbriaa,tus); Vespe?"tilio te'YI'll1ninckii (.=Scotophilus temm,inckii); Cheirom,eles torqua,t us. All these forms are based on extra-Philippine types. HUET, J. Note sur une Espece nouvelle de Mammiferes du Genre Mydaus Provenant de l'Ile Palaouan. Le Naturaliste II 9 路 annee No. 13 (1887) 149-151. Figs. 1-3. (September 15.) Contains the original descl'iption of Mydaus ma1'chei, from a specimen collected by Marche in Palawan. The form is compared with Mydaus meliceps Cuvier. Figures 1 and 2 give a detailed drawing of the entire animal and three views of the skull. HUET, J. Sur deux Especes Nouvelles de Sangliers. Le Naturaliste II No. 20 (1888) 6-8, figs. 1-4. Contains original descriptions of Sus ahmnobarbus and Sus marchei. The first is from "Palaouan" (Palawan), the second species from "Laguan" (presumably Laguna Province or Laguna de Bay, central Luzon, so interpreted by N. Hollister). The figures show the entire animals, with details of lower jaws, skulls, and molar teeth. JENTINK, F. A. On two mammals from the Calamianes Islands. Notes Leyden Mus. 17 No. 9 (1895) 41-48. (August.) Original descriptions of Mydaus schadenbe?'gii and Mtmgos parvU8 are given. Jentink is apparently not aware that a species of Mydaus has been described :Drom Palawan. Some later writers regard the forms as belonging to the same species. JENTINK, F. A. Revision of the genera Macroglossus and Syconycteris and description of a new genus and species, Odontonycteris meyel'i. Notes Leyden Mus. 23 No. 3 (1901) 131-142. (July 15.) Original description of Odontonycte?'is meye?'i (= 1Ilam'oglossus lagochiltls lagochilus Matschie) from Gl'eat Sangi Island (extra-Philippine). This form is based on a presumed anomalous specimen having an additional molar. JORDANA y MORERA, RAM6N. Bosquejo geographico e hist6rico natural del Archipielago Filipino. Madrid (1885). Contains much information on the mammals of the Philippines. References to this work are not quoted in synonymies. JOURDAN, M. Memoire sur quelque mammiferes 'nouveaux. Compt. Rend. Acad. Sci. 5 (1837) 521-524. Contains the very brief original description of Parado'Xurus philip-

pinensis. LINNlEUS, C. Systema Natural ed. 10 1 (1758) 14-77 Mammalia. Original descriptions Bubaltcs bubalis, Mus -rattus (= Rattus ?'attus) , Simia syrichta (= Tarsius sp.), Felis catus, Lemur' volans (= Cynocephalus volans), and VespertiUo spasma (= M egaderma spasma). The type locality of Sitmia syrichta is Luzon (doubtful) and Le'YILu1' volans is from Pampanga, Luzon, according to Thomas, Proc. Zool. Soc. London (1911) 130. Neither of these forms occurs in Luzon now, The other forms are from extra-Philippine localities.

PH;ILIPPINE LAND MAMMALS

LYDEKKER, R. Mammalia. Zool. Rec. 29. (1892) 24-25. Gives a generic combination to Nachtreib's name menagensis for the Philippine slow lemur which gives the name an authentic place in scientific nomenclature, hence the "original description" of the form. LYDEKKER, R. The Deer of All Lands. A History of the Family Cervidre, Living and Extinct. London (1898). A rather general account of various species of Philippine deer. The following forms are recognized: Cervus unicolor philippinus [=Rusa marianus (Desmarest) J; Cervus unicolo1' nigricans Brooke (= Rusa nig?'icans Brooke) ; Cervus alfredi Sclater [= Rusa alfredi (Sclater)]; and Ce?'v1GS c1~lionensis Elliot [=Rusa calamw,lItensis (Heude) J. LYDEKKER, R. Wild Oxen, Sheep, and Goats of All Lands, Living and Extinct. London (1898). Account of Philippine Bovidre. LYDElKKER, 'R. An undescribed type of rusine deer. Ann. & Mag. Nat. Hist. VII 6 (1900) 204-205. Contains the original description of Cervus tavistocki from an unknown locality, "probably Philippines." LYON, MARCUS WARD, Jr. Notes on the porcupines of the Malay Peninsula and Archipelago. Proc. U. S. Nat. Mus. 32 (1907) 575-594, pIs. 54-57. (June 29.) A general discussion of the Oriental porcupines. Hystrix pwmila Gunther is placed in the genus Thecurus. LYON, MARCUS WARD, Jr. The authority for the name Nycticebus menagensis. Proc. BioI. Soc. Washington 22 (1909) 89. (April 17.) Shows the authority for the name Nycticebus menagensis to be Lydekker (vide supra). LYON, MARCUS WARD, Jr. Remarks on the Insectivores of the genus Gymnura. Proc. U. S. Nat. Mus. 36 (1909) 449-456, pIs. 34-37. (May 27.) Discusses the type of Podogymnura, truei and figures it on plates 35, 36, and 37. LYON, MARCUS WARD, Jr. Treeshrews: an account of the mammalian family Tupaiidre. Proc. U. S. Nat. Mus. 45 (1913) 1-187. Gives detailed descriptions of Philippine species of Tupaia and Urogale. MAJOR, C. 1. FORSYTH. On Sus verrucosus Mull. & Schleg., and allies from the Eastern Archipelago. Ann. & Mag. Nat. Hist. VI 19 (1897) 521-542. Sus ve1TUCOSUS rnrinda,nensis (= S1.ÂŁS mindanensis) and Sus barbatus balabacensis listed. This is a very important paper on Philippine pigs. The literature on the group is reviewed. MARTINEZ DE ZUNIGA, JOAQUIN. Estadismo de las islas Filipinas, 0 mie viajes por este pais. Madrid (1893) 2 vols, This work, written sometime between 1786 and 1818 (the dates of the author's arrival in the Philippines and his death), was not published until 1893. Part of the second volume contains many data on the mammals.

INTRODUCTION

MATSCHIE, PAUL. tiber Saugethiere von den Philippinen. Sitz.-Ber. Geti. Nat. Freunde Berlin 5 (1898) 38-43. (May 17.) Tupaja mollendo1'ffi ( = Tupaia 1nollendorffi) , Sciul"us 11Lollendo?'fji, and Sciurus albicauda, all from Culion Island, are described as new. The following species are listed, with not.es: Taphozus philippinensis Waterhouse, Manila; lIIiniopterus tibialis Tomes, Manila; lviiniopte?"Us pusillus Tomes, Manila; Rhinolophus TUJUS Petel's, Tablas; Rhinolophus (J;r'cuatus Peters (= Rhinolophus arcuatus w'cuatu,s Peters), Manila; Hipposideros antr-icola Peters, Manila; Macr'oglossus aust?'alis Peters [= Macroglossus lagochilus lagochilus (Matschie)] Tablas; Galeopithecu,s philippinensis Waterhouse [= Cynocephalus volans (Linnreus)] Samar; Phlmomys cumingi Waterhouse, Marinduque; Sciurus stee?'ii Gunther, Mindanao; Chrotomys whiteheadi Thomas, Luzon; Cer'vu8 calamianensis Heude [=R~t.Sa caiamiane?'u,is (Heude)]; Cer'VUS philippinensis Brooke ( = Rusa ? ); Cer'vus crassicornis Heude ( = ?) , Samar. Cm'vus basilanensis Heude (=R~t.Sa basilanensis), Mindanao; Bubalus ket'abau Jet'us Nehring [=Bubalus bubalus (Linnreus)], Middle Luzon; Bubalus moellendorffi Nehring, Busuanga. MATSCHIE, PAUL. Die Fledermause des Berliner Museums fUr Naturkunde. I. Lieferung. Die Megachiroptera. Berlin (1899) 1-102, pIs. 1-14. Lists certain Philippine species. Contains the original description of Macroglossus lagochilus with type locality Buru. MEARNS, EDGAR A. Description of new genera and species of mammals from the Philippine Islands. Proc. U. S. Nat. Mus. 28 (1905) 425-460. Mearns describes the following new genera and species from collections made by himself: Urogale, Podogymnur'a, Bullimus, Limnomys, Tarsomys, and Apomys. Cynomolgus mindanensis (= Pithecus mindanensis) , Cynomolgus m~'l'Uianensis apoensis (=Pithecus mindanensis) , Cynomolgus suluensis (=Pithemt.S suluensis) , Cynol1wlgus cagayanus (=Pithecus cagayanus) , PWr'optIM lanensis (=Pteropus Vatmpyrus lanensis) , Pter'opus cagayanus (= Pter'opus hypomelanus cagayanu8) , Urogale cylindrura, Podogymnura truei, Mus tagulayensis (= Rattus tagulayensis), Mus albigularis (=Rattus albigularis) , Mus magnirostris (= Rattus magnirostris) , Mus mindatnensis (= Rattus mindanensis) , Mus zamboang;;c (=Rattus zamboang;;c) , Mus kelleri (=Rattus keller路i) , Mus todayensis (=Rattus todayensis) , Mus vulcani (=Rattus vulcani) , Mus pantarensis ( = Rattus pantarensis) , Mus commissarius, Bullimus bagobus, Liwnwmys sibttanus, Tarsomys apoensis, Apomys hylocmtes, Apomys petrt拢us, Apomys insignis. The Mearns collection formed the basis of several other pa:pers listed in the bibliography. MEYER, A. B. Proc. Zool. Soc. Lond<m (1879) 666. A note concerning the type locality of Ce?'vus alfredi. the type was from Samar or Leyte.

Presumably

MEYER, A. B. Zool. Gart. (1890) 199. Data on Phlmomys. MEYER, A. B. Description of a new squirrel from the Philippine Islands. Proc, Zool. Soc. London (1890) 579-601. Contains the type description of Sciurus cagsi (=Sciur'us mindanensis Steere).

PHILIPPINE LAND MAMMALS

MEYER, A. B. Eine neue Tarsius Art. Abhandl. und Ber. des Konigl. Zoo!. Mus. Dresden 1 (1894) 1-2 (1895). Tarsius philippensis is described from specimens from Samar, Leyte, and Mindanao. Meyer is under the impression that the Tarsius from Bohol is of the same species, and also gives Luzon as a locality, the latter perhaps on the sole authority of Camel or Petiver (see bibliography) . MEYER, A. B. Eine Neue Phlreomys-Art. Abhandl. und Ber. des Konigl. Zool. Mus. Dresden 6 (1894-95) 1-2 (1895). Describes Phlceomys (?) schadenbergii (=Crate1'omys schadenbergii) from Mount Data, Lepanto, Luzon. The skull of the type wa.:l wanting, hence its association with Phlceomys. MEYER, A. B. Der Stinkdachs del' Philippinen (Mydaus marchei Huet) , Abhand. und Bel'. des KonigI. Zool. Mus. Dresden No. 13 (1894-95) 1-4. (August 25, 1895.) Myda.us schadenbergii Jentink said to be a synonym of Mydaus marchei Huet. MEYER, A. B. Saugethiere von Celebes und Philippinen-Archipe!. 1. Abhandl. und Bel'. des Konigl. Zool. Mus. Dresden No. 6 (1896-97) (1896) . This important paper gives excellent notes on Philippine mammals with illustrations. MEYER, A. B. Saugethiere von Celebes und Philippinen-Archipel. 1. AbhandI. und Bel'. des Konigl. Zool. Mus. Dresden No.7 (1898-99) (1899). Mus datm (=Rattus datm) is described from a specimen from Mount Data, Lepanto, Luzon. The differences hetween this form and M. r;hrysocomus are pointed out. Other Philippine mammal records are given. MILLER, GERRIT S., Jr. A second Specimen of Odontonycteris meyeri Jentink. Proc. BioI. Soc. Washington 18 (1905) 253. (December 9.) A note on a specimen of Odontonycteris meyeri Jentink [=Mac1'oglossus lagochilus lagochilus (Matschie)] from Cagayan Sulu, collected by Edgar A. Mearns. MILLER, GERRIT S., Jr. The families and genera of bats. Nat. Mus. 57 (1907) I-XVII, 1-282, pIs. 1-14. This important work lists a few Philippine species.

~ull.

U. S.

MILLER, GERRIT S., Jr. Descriptions of two new genera and sixteen new species of mammals from the Philippine Islands. Proc. U. S. Nat. Mus. 38 (1910) 391-404, pIs. 18-20. (August 19.) This important paper contains the type descriptions of the following species: C?'ocidu1'a halconus, C1'ociduTa beatus, Crocidura mindorus, Crocidura gr'andis, Tupaia cuyonis, Pteropus pumilus, Chilophylla hirst~ta, Taphozous 1Jluto, Epimys tyr'annus (=Rattt~ tyr'annus) , Epimys gala (= Rattus gala), Tryphomys adustus, Batomys dentatU8, Apomys bardus, Apomys maยงor, Apomys musculus, Tarsius jraterculU8. The two new genera are Chilophylla and Tryphomys. Thirteen of the

INTRODUCTION types are from a collection made by Dr. Edgar A. Mearns. based on specimens from the Bureau of Science, Manila.

Three were

MILLER, GERRIT S., Jr. Note on the Mus commissarius of Mearns. Proc. BioI. Soc. Washington 24 (1911) 38. (February 24.) Miller reports two specimens of this species from Luzon. He further identifies 8 specimens from Japan (Tokyo and Misaki) as belonging to this species. MILLER', GERRIT S., Jr. The cranial and dental characters of Chilophylla. Proc. BioI. Soc. Washington 25 (1912) 115-118. (June 29.) The cranial and dental characters were omitted from the type specimen since the skull had been lost. These data are here presented, taken from a specimen collected by Arthur de C. Sowerby, Port Swettenham, Federated Malay States. MILLER., GERRIT S., Jr. A new pteropine bat from Luzon. Proc. BioI. Soc. Washington 26 (1913) 73-74, March 22. Eonycteris robusta, is described from an adult male specimen collected by Dr. Paul Bartsch, in a cave at Montalban, Luzon, July 5, 1908. MULLER, PHILIPP LUDWIG STATIUS. Des Ritters Carl von Linne, Koniglich Schwedishchen Leibarztes Vollstandiges N atursystem. Supplementsand Register-Band libel' aIle Sechs Theile oder Classen des Thierreichs. Nlirenburg (1776). Contains the type description of T1'ichecus dugon (=Dugong dugon). NACHTRIEB, HENRY F. A new lemur (Menagensis). Zool. Anzeiger 15 (1892) 147-148. The new lemur without a generic designation is given the name Mena,gensis. The name has, however, been rehabilitated by Lydekker. NEHRING, ALFRED. Dber Zwei Schadel des Sus longirostris Nehring von Borneo und Java. Sitz.-Ber. Ges. Nat. Freunde Berlin No.5 (1886) 80-85, figs. 1-3. Contains in a footnote the original description of Sus philippensis. Nehring gives Meyer credit for the name since he quotes Meyer's manuscript. The author must be regarded as Nehring since he was the first to publish. NEHRING, ALFRED. Dber Sus celebensis und Verwandte. AbhandI. und Ber. des Konigl. Zool. Mus. Dresden (1888-89) 1-34, pIs. 1-2. Nehring discusses several species and subspecies of wild boars in various German museums. He discusses Sus philippinensis and describes Sus ba?'ba,tu8 pala,vensis from a skin and skeleton specimen collected Nov. 3, 1887, at Puerto Princesa, Palawan, by Platen. This is an important paper on the Oriental forms of Sus. NEHRING, ALFRED. Dber Sus celebensis and Verwandte. Nat. Freunde Berlin No.1 0 (1889) 196. A short note on Sus barba,tus pa,la,vensis.

Sitz.-Ber. Ges.

NEHRING, ALFRED. Dber Einen Unterkiefer des Philippinen Wildschweins. Sitz.-Ber. Ges. Nat. Freunde Berlin No. 1 (1890) 8-11. General notes on Philippine pigs.

PHILIPPINE LAND MAMMALS

NEHRING, ALFRED. Ubel' Saugethiere ,der Philippinen, namentlich tiber Phlceomys cumingi Waterh. und Bubalis mindol'ensis Heude. Sitz.-Ber. Ges. Nat. Freunde Berlin No.6 (1890) 101-108. (June 17.) Contains the type description of Phlceomys cumingi var. pallida. ( = Phlceom ys pallida). NEHRING, ALFRED. Uber Phlceomys cumingi val'. pallida. Sitz.-Ber. Ges. Nat. Fl'eunue Ber lin ~ o. 8 (1890) 153-154. (October.) A note on the genus Phlceomys. NEHRIN G, ALFRED. Uber Siiugethiere von den Philippinen, namentlich von del' Palawan-Gruppe. Sitz.-Ber. Ges. Nat. Freunde Berlin. No. 8 (1894 ) 179-193 , fi gs. 1-3. Th is pa per contains the type descriptions of Bubalus moellendorffi, f rom Busuanga, and B u balus kerabau fents [= Bubalus bubalus (Linnreus) ]. Phlceom;ys pallidus Nehring, Tupaja ferruginea (=Tu1Jaia ?noellend01'ffi Matschie), Sus bOl'batus pcdavensis, f r om Taytay, P a lawan, and Sus ba?'batus calamianensis (Heude) are listed with notes. The specimens a r e from a small collection sent by von Moellendorff f r om Manila. NEHRING, ALFRED. tiber Sus marchei Huet und Tragulus nigl'icans Thom as. Sitz.-Ber. Ges. Nat. Frednde Berlin No.9 (1894) 219-226. (November 20.) Discussion of Philippine pigs and the chevl'otain. PETERS, W. Uber die von Hrn. F. Jago r bisher auf Malacca, Borneo, Java. und den Philippinen Gesammelten Saugethiere aus den Ordnungen der Halbaffen, Pelzf1atterer und Fleder thiere. Monatsb. Konigl. Preuss, Akad. der Wiss. zu Berlin (1861) 706-712. (July 25.) This work contains the type descriptions of Emballonura discolo,," [=Em ballon ur a alecto (Eydoux and Gervais)], Paracali, Luzon; Rhinolo phu s ?'Ufus (=Rhinolophus subrufu.s Andersen), Paracali, Luzon; Phy llorhina obscura (=Hipposide?'os obscura), Paracali, Luzon; Phyllorhina antricola (=Hipposide?'os antricola); Pachysoma luzoniensis [= Cynopterus b?'achyotis brachyotis (M tiller) ], Iriga Volcano, Luzon, and Pachysoma (Ptenochirus) jagori [=Ptenochirus jagori (Peters)]. PETERS, W. Uber Neue oder Ungentigend Bekannte Flederthiere (Vampyrops, Uroderma, Chiroderma, Ametrida, Tylostoma, Vespertilio. Vesperugo) und Nager (TylomY03, Lasiomys). Monatsb. Konigl. Preuss. Akad. der Wiss. zu Berlin (1866) 392-411. (1867). Contains the original description of Vespertilio (Kerivoula) jagori (=Kerivoula jagorii), from Samar. PETERS, W. uber Neue Arten von Spitzmausen des Konigl. Zoologischen Museums aus Ceylon, Malacca, Borneo, China, Luzon, und Ost-Afrika. Monatsb. Konigl. Preuss. Akad. der Wiss. zu Berlin (1870) 584--596. (1871.) Contains the original description of Crocidura (P.) luzoniensis ( = Pachytbra luzoniensis) , from Luzon. PETERS, W. uber die Gattungen und Arten der Hufiisennasen, Rhinolophi. Monatsb. Konig!. Preuss. Akad. del' Wiss. zu Berlin (1871) 301-332. (June, 1872.)

INTRODUCTION

Contains the original descriptions of Phyllorhina C01'onata (=Hipposideros cO~'onatus) Mainit, Mindanao; and Rhinolophus arcuatus (=Rhinolophus a1'cuatus arcuatus), Luzon. PETIVER, JACOB. Cat. Class. Gazophylacium Natur. 21 plate 13, fig. 11. Drawing of Ceroopithecus luzonicus minimus (=Tarsius sp.). SANCHES, DOMINGO. Los Mamiferos de Filipinas, Anales de la Soc. Espanola de Hist. Nat. II 27 (1898) 93-110. Notes on numerous Philippine mammals with distributional data. SANCHEZ, DOMINGO. Los Mamiferos de Espanola de Hist. Nat. II 29 (1900) A discussion of Philippine mammals eliminates forms erroneously credited revised list.

Filipinas. Anales de la Soc. 177-290. with regard to distribution. He to the Philippines and gives a

SCLATER, P. L. Report on additions to the society's menagerie. Proc. Zool. Soc. London (1870) 380-383, pI. 28. Contains the original description of Ce1'vus alJredi (=Rusa alf~'edi). SCLATER, P. L. The "tamaron" of the Philippine Islands. Nature 38 (1888) 363-364. (August 16.) Extracts from a letter from J. B. Steere, regarding <CAnoa minMj'ensis" (=Bubalus mindo'/'路ensis). SLACK, J. H. Mammalogical notices. Proc. Acad. Nat. Sci. Philadelphia 19 (1867) 34-38, pI. 1. Contains the original description of Macacus JU1' [ = Pithecus philippinensis( Geoffroy) ] . SMITH, CHARLES HAMILTON. Supplement to the order Ruminantia. Griffith's Animal Kingdom 4 (1827) 33-428. Contains the original description of Cervus philippinus (=Rusa philippinus) on page 147. STEERE, J. B. In Selater, Nature 38 (1888) 364. (August 16.) Brief description of Anoa mindo1'ensis (=Bubalus mindorensis . Heude), in extracts from a letter. STEERE, J. B. (A letter from J. B. Steere to the Zoological Society of London). Proc. Zool. Soc. London (1888) 413-415. Reports on the Mindoro buffalo, the Ta1'l1.araon (sic). A good description is given with notes on habits. STEERE, J. B. A list of the Birds and Mammals Collected by the Steere Expedition to the Philippines with Localities apd Brief Preliminary Descriptions of Supposed New Species. Ann Arbor, Michigan (1890) 1-30. (July 14.) Contain:;; the original brief descriptions of SciuT'/.('s sa'l'narensis and Sciurus mindanensis. Probubalus mindorensis (=Bubalus mindorensis Heude) is likewise listed as a new species. Several other species are listed with their localities. Macacus philippinensis [= Pithecus philippinensis (Geoffroy)], Tarsius spectrum (= ? Tarsius cwrbonarius Heude and Tarsius philippensis Meyer, since Samar is also given as a locality r路ecord), Pteropus edulis [ = Pteropus vampyrus lanensis (Mearns) ], Pteropus jubatus [= A cerodon jubatus jubatus (Esch-

P H'ILIPPINE LAND MAMMALS

scholtz)], Pte?'opus hypomelanus [ = Pteropus hypomelanus cagayanus (Mearns)], Cynonycte?'is amplexicaudata (= Rottsettus amplexicaudata Geoffroy), Phyl101'hina diadema [ = Hipposideros diadema grf,geus (M eyen) ], Galeopithec'us philippinensis [ = Cynocephalus volans (Linnmus)], Tupaia sp. (=?), Paradoxurus philippinensis Jourdan, MacroXiUs philippinensis (=perhaps Sciurus philippinensis Waterhouse although that species appears later in the list), Sus philippinensis (= ? Basilan and Mindoro), Tragulus sp. (= Tragulus nigricans Thomas), Cervus philippinus (=? Rusa nigricans), Phlreomys cumingii; Sciu~ philippinensis, Sciurus steeri, Hystrix pumila (= Thecurus pumilus Gunther), Pholidotus dahlmanni (= Manis javanica Desmarest). Steere states that a number of other smaller species await identification. Apparently some of these specimens are listed by Hollister, Proc. U. S. Nat. Mus. 46 (1913) 229-341 Dec. 31. TEMMINCK, C. J. Monographies de Mammalogie, Tome Premier, Paris (1827) I-XXXII, 1-268. Contains the original description of Felis minuta from Java. TEMMINCK, C. J. Monographies de Mammalogie ou Description de quelques genres de mammiferes, dont les especes ont eM observees dans les differents musees de l'Europe 2 (1841) 1-392, pIs. 28--70. Leiden (1835-1841) . Contains the original descriptions of Vespertilio abramus (=Pipistrellus abramus) , Tylonycteris pachypus, Pipistrellus tenuis, Rhinolophus bicolor (= Hipposideros bicolor") , and RhinolopJuus luctus Temminck (=Rhinolophus sp. ?). THOMAS, OLDFIELD. Diagnoses of four New Mammals from the Malayan region. Ann. & Mag. Nat. Hist. VI 2 (1888) 407. Contains the original description of Nannosciurus concinnrus. THOMAS, OLDFIELD. Preliminary notes on the characters and synonymy of t.he different species of otter. Proc. Zoo!. Soc. London (1889) 190-200. The specific name cinerea is revived for the Eastern otter. THOMAS, OLDFIELD. On some new Mammalia from the East Indian Archipelago. Ann. & Mag. Nat. Hist. VI 9 (1892) 250-254. Contains the type descriptions of Tr"agulus nigricans and Urogale everetti. THOMAS, OLDFIELD. Description of a new Sciuropterus from the Philippines. Ann. & Mag. Nat. Hist. VI 12 (1893) 30. Describes Sciuropterus nigripes. THOMAS, OLDFIELD. On the Palawan representative of Tupaia ferruginea. Ann. & Mag. Nat. Hist. VI 13 (1894) 367. Contains the original description of Tupaia palwwanensis as a subspecies of T. ferruginea. THOMAS, OLDFIELD. Descriptions of two new bats of the Genus Kerivoula. Ann. & Mag. Nat. Hist. VI 14 (1894) 460-462. Contains the type description of KerivO'Ula whiteheadti.

INTRODUCTION

THOMAS, OLDFIELD. Preliminary diagnoses of new mammals from northern Luzon, collected by Mr. John Whitehead. Ann. & Mag. Nat. Hist. VI 16 (1895) 160-164. Contains short preliminary descriptions of Chrotomys whitehetUU, Mus luzonicus (=Bullimus luzonicus Thomas), RhynchomJJjs soricoides, Xeromys ? silaceus (= Celamomys silaceus) , Carpomys melam~rus. Carpomys phmurus, and Batomys g?'anti. Complete descriptions of these forms are published in the Transactions of the Zoological Society of London, vide infra. THOMAS, OLDFIELD. On mammals from Celebes, Borneo, and the Philippines recently received at the British Museum. Ann. & Mag. Nat. Hist. VI 18 (1896) 241-250. Contains the original description of the genus Harpyonycteris, and of the type of the genus, Harpyonycteris whiteheadi, from Mindoro. THOMAS, OLDFIELD. Minutes of the Proc. Zool. Soc. London for June 15, 1897 (published June 19). Contains preliminary diagnoses of Nannosmurus swmaricus, CrunomJJjS fallax, and Mus mindorensis = (Rattus mindorensis). THOMAS, OLDFIELD. On the mammals obtained by Mr. John Whitehead during his recent expedition to the Philippines. Trans. Zoo1. Soc. London 14 pt. 6 (1898) 377-412, pIs. 30-36. This is one of the most important papers on Philippine mammalogy. Some 38 species are listed with copious notes, and in a large number of cases with complete detailed descriptions. The original description of the genera Celmnomys and Crunomys and of the species Mus ephippium negrinu[f (= Rattus negrinus) are given. The following genera and species are listed: M acacus cynomolgus Linnreus (=Pithecus philippinensis I. Geoffroy), Ta?'l"ius philippensis Meyer, Pterop~Ls jubatu $ [ = Acerodon jubatus jubatus (Eschscholtz)], Pte'ropus vwmpy?"Us Linnreus [= Ptet路opus vampyrus lanensis (Mearns)], Xantharpia a?nplexicaudata Geoffroy [=Roussettus atrnplexicaudatus (Geoffroy) J, Harpyionycteris, Harpyionycte?-is whiteheadi Thomas, Carponycteris australis Peters (= Miniopterus australis Thomas), Hipposideros diadema Geoffroy [ = Hipposideros diadema griseus (Meyen) J, Pipistrellus imbricatus (Horsfield), Myotis macro tarsus (Waterhouse), Kerivoula whiteheadi Thomas, Miniopte?"Us schreibe?'si pusillus Dobson (= MinioptM-;US pusillus Dobson), Galeopithecus philippinensis Waterhouse [= Cynocephalus volans (Linnreus)J ', Crocidura (Croc.) gTayi Dobson ( = Crocidura grayi Dobson), Felis minuta Temminck, Felis domestica Linnreus, (= Felis catus Linnams), Viverra tangalunga Gray, Pa?'adoxurus philippinensis Jourdan, Sciurus samarensis Steere, NannoBciurus samaricus, Celamomys, Celmnomys silaceus (Thomas), Chrotomys, Chrotomys whiteheadi Thomas, Crunomys fallax sp. nov., Rhynchomys, Rhynchomys soricoides Thomas, Phlceomys pallidus Nehring, Mus evet'etti Gunther ( = Rattus everetti Gunther), Mus luzonicus Thomas (= Bullimus luzonicus Thomas), Mus decumanus val'. Pallas [ = Rattus norvegicus (Linmeus)], Mus rattrus (Linnreus) val'. [ = Rattus rattus (Linnreus)], Mus windorensis Thomas (= Rattus mindo269463-4

PHILIPPINE LAND MAMMALS

?'ensis Thomas), Mus cht'ysocomus Hoffman [ = Rattus datre (Meyer)], Mus ephippium negrinus Thomas (= Rattus negrinus Thoma.;;), Batomys, Batomys g1'anti Thomas, Carpomys, Carpomys melanurus Thomas, Carpo?nys phreurus Thomas, Crateromys, Crater,omys schadenbergi (Meyer), Bubalus mindorensis Heude, Sus celebensis philippinensis Nehring. Nine species are figured in colored plates, while the skull and tooth details of several other forms are given. THOMAS, OLDFIELD. On mammals collected in Mindanao, Philippines, by Mr. M. ,P . Anderson for the Duke of Bedford's exploration of eastern Asia. Abstracts Proc. Zoo!. Soc. London No. 39 (1907) 5. (February 12.) Original description of Crunomys melanius. THOMAS, OLDFIELD. The Duke of Bedford's zoOlogical exploration in eastern Asia.-III. On mammals obtained by Mr. M. P. Anderson in the Philippine Islands. Proc. Zool. Soc. London (1907) 140-142. The complete description of Crunomys melanius Thomas appears here. He lists Urogale cylindru1:a Mearn.;; (but thinks it only a local form of U. everetti), Sciurus mindanensis, Mus sp. (of Rattus group), Mus bagobus (Mearns) = (Bullimus bagobus Mearns), Mus vulcani Mearns [ = Rattus vulcani (Mearns)], Sus sp. These specimens. were obtained in "North Central Mindanao" and on Mount Apo. The collections were made by Mr. M. P. Anderson who visited the region to collect topotypes of the species discovered by Mearns. Due to illness he made only very meager collections. THOMAS, OLDFIELD. The nomenclature of the flying-lemurs. Ann. & Mag. Nat. Hist. VIII 1 (1908) 252-255. Shows that Cynocephalus volans is the correct name for the Philippine species of flying lemur. The type locality as given by Petiver as Philippine Islands, later specifically given as Province Pampanga, southern Luzon. * Fixes the name of family Galeopteridre. THOMAS, OLDFIELD. The nomenclature of certain lorises. Ann. & Mag. Nat. Hist. VIII 1 (1908) 4'67-469. Disagrees with Cabrera as to Lemur tardigradus Linnreus. Believes menagensis of Nachtrieb described without a genus, is validated by Trouessart who places it in the Genus Nycticebus, and thus replaces philippinus as the specific designation for the Tawitawi form of the slow lemur. (Hollister believes, Proc. U. S. Nat. Mus. 46: 328, that Cabrera's name is available since he believes the Mindanao form to be specifically distinct from the Tawitawi form.) THOMAS, OLDFIELD. The squirrels described as Sciurus steeri from Balabac and Palawan. Ann. & Mag. Nat. Rist. VIII 2 (1908) 498. Regards the form SC'iturus steeri Giinther as belonging to Balabac, and name.;; a form occurring in Palawan Sciu?"Us juvencus.

* Pallas, Act. Ac. Petrop. 4 (1870) 208. Galeopithecus volans Camellu desc?'iptus. It was Father George Joseph Camel, who sent from the Philippines, the description published by Petiver, itself in turn the oasis of Linnreus's Lemur V olans.

INTRODUCTION

THOMAS, OLDFIELD. New species of Paradoxurus of the P. philippinensis group and a new Paguma. Ann: & Mag. Nat. Hist. VIII 3 (1909) 374-377. Contains original descriptions of Parado :kn~rus mina x and Paradoxurus torvus. THOMAS, OLDFIELD. The mammals of the tenth edition of Linnreus; an attempt to fix the types of the genera and the exact bases and localities of the species. Proc. Zoo!. Soc. London (1911) 120-158. Endeavors to show that Si'YJ1lia syrichta should be applied to the form now known as Pithecus philippinensis Geoffroy. (Cabrera shows that the name applies to a species of Tat"sius. See Bibliography under Cabrera.) TOMES, ROBERT F. A monograph of the genus Miniopterus. Pl'oc. Zoo!. Soc. London (1858) 115-128. Contains original descriptions of Miniopte1"Us wust?'alis and Minioptet"US tibialis" Tomes r€gards Vespertilis eschscholtzii Waterhouse as a synonym of Miniopten~s blepotis Temminck and gives measurements of the type specimen. Miniopterus tr"istis (Waterhouse) is discussed. TOMES, ROBERT F. On the characters of four species of bats inhabiting Europe and Asia, and the description of a new species of Vespertilio inhabiting Madagascar. Proc. Zoo!. Soc. London (1858) 78. A redescription of the type of Vespe?"tilio rufopictus Waterhouse (=Myotis ?'ufo-pictus) is given with descriptions and comparisons of related forms. TOMES, ROBEkl.T F. A monograph of the genus Kerivoula. Proc. Zool. Soc. London (1858) 322. Tomes concludes that Vespeq·tilio pellucidus Waterhouse (=Kerivoula pellucidus) is identical with Ke?'ivoula hat"dwicki Horsfield. TROUESSART, E. L. Description d'une Espece Nouvelle de Musaraigne de la Collection de Musee de Paris. Le Naturaliste No. 42 (1880) 330" (December 15.) Contains the original description of Crocidu?'a edwardsiana. TROUESSART, E. L. Catalogus Mammalium tam Viventium quam Fossilium. Nova editis (Prima Completa) Berolini Tomus 1 (1898-1899) 1-664. Primates, Prosimire, Chiroptera, Insectivora, Carnivora, Rodentia, Pinnipedia. Numerous species are listed from the Philippines. Tomus 2. Furth€r species listed from the Philippines. Quinquennale supplementum (1904) 1-929. Lists 2 Primates, 2 Prosimire, 35 Chiroptera, 1 Dermoptera, 6 Insectivora, 7 Carnivora, 28 rodents, and 15 UngUlata from the Philippines. S€veral forms are included that may be regarded as doubtful. WATERHOUSE, G. R. On the flying lemurs. Proc. Zoo\. Soc. London (1838) 119-120. Describes Galeopithecus philippinensis = Cynocephalus volans (Linnreus) and compares it with G. temminckii.

PHILIPPINE LAND MAMMALS

WATERHOUSE, G. R. On a new genus of ~odents from the Philippine Islands. Proc. Zool. Soc. London 7 (1839) 107-108. The genus Phlceomys is described as a new subgenus with the species Phlceomys cumingi. WATERHOUSE, G. R. A new species of squirrel, sent by Hugh Cuming, Esq., from one of the Philippine Islands. Proc. Zool. Soc. London 7 (1839) 117-118. SCiU1'US philippinensis is described as new. WATERHOUSE, G. R. Various species of bats from the Philippine Islands, collected by Hugh Cuming, Esq. Proc. Zool. 'Soc. London (1843) 66-69. The following species from the Cuming collection are described: Rhinolophus pygml£us (=Hipposideros pygmlEus), Rhinolophus philippinensis, Megaderma philippinensis [=Megaderma spasma spasma (Linnreus)]. Aside from the presumed new species the following fOTms are listed: Pte1'opus jubatus Eschscholtz, Pteropus sp. (perhaps new), Pachysomrlf amplexicaudatum (GeoffTOY) (=Roussettus amplexicaudatus) , Pachysoma titthlEcheilum (Temminck) (=Cynop·t eru8 brach'yotis brachyotis Muller), Pachysoma brevicaudatU'rn (Is. Geoffroy) [=? Cynopte1'us b'f'achyo·t is brachyotis (Miiller)], Macroglossus minimus (Geoffroy) (=M acroglossus lagochilus lagochilus Matschie), Rhinolophus nobilis Horsfield [=Hipposideros diadema griseus (Meyen)], Rhinolophus bicolor Temminck (=Hipposide?'os bicolo?·). WATERHOUSE, G. R. On a new species of MegadeTma. Proc. Zool. Soc. London (1843) 69. Type description of Megaderma philippinensis = Megaderma spasma spasma (Linmeus). WATERHOUSE, G. R. (Exhibition of and description of two new species of mouse.) Ann. & Mag. Nat. Hist. 12 (1843) 134. Contains description of Mus castaneus. WATERHOUSE, G. R. Description of species of bats collected in the Philippin·e Islands and presented to the society by H. Cuming. Proc. Zool. Soc. London (1845) 3-10. (January 14.) Descriptions of new bats from the Cuming collection with a key. The following are described as new: Vespertilio tristis (=Miniopterus t?'istis), Vespertilio eschscholtzii (=Miniopterus eschscholtzii), Vespertilio macro tarsus ( = Myotis macro tarsus ), Vespertilio pellucidus (= Kerivoula pellucida), Vespertilio meye'rii [=PipistrelLus irretitus (Cantor)], Vespertilio 1'ufo-pictus (=Myotis rrufopictus). Waterhouse also lists Nycticejus borbonicus (=Scotophilus temmincki Horsfield and Dysopes tenuis Horsfield (= ? ChlErephon luzonus Hollister). WATERHOUSE, G. R. (New Philippine bats.) Ann. & Mag. Nat. Hist. 16 (1845) 49-56. (July.) Contains much the same material as appears in the preceding paper, Proc. Zool. Soc. London (1845) 3-10. (April.) COLLECTION AND PRESERVATION OF SPECIMENS

The collections of mammals that have been made in the Philippine Islands are very meager when one considers the extent

INTRODUCTION

and diversity of territory involved. Careful collecting has been confined to only a few places, and even in such well known localities as Mount Data, made famous by Whitehead's collecting, and Mount Apo, by Mearns's collections, intensive efforts will doubt120'

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126

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PHILIPPINE ISLANDS

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The Phili pp in e Islands, show in g wh ere collections of m amm als have been ma de. Density of stippling suggests the r elat ive a mount of collectin g .

less yield other novelties. Many of the islands do not have even a single record for a mammal (see fig. 1). Much collecting needs to be done before the fauna can be even moderately well known. The following localities will doubtless yield the greatest percentage of new forms.

PHILIPPINE LAND MAMMALS

Luzon and neighbo1"ing islands.-The mountains along the eastern part of northern Luzon, ' probably representing island territory recently added to the Luzon mainland, will repay careful investigation. The islands about the coast, particularly the Babuyan and Batan groups to the north, and Polillo and Catanduanes to the east, are very important and should not be overlooked by the collector. Mindoro.-Mount Halcon and the central range culminating in Mount Baco with an elevation of 2,489 meters will doubtless prove to be a fertile field for collections. Such collections will prove conclusively whether or not Mindoro has received any of its fauna from Borneo via the Palawan Islands, a question at present without answer. Palawan.-The high mountains in southern Palawan are as yet wholly unknown to the scientist, and will probably offer to an intrepid collector-explorer the greatest reward of new species and new records. I visited southern Palawan with the intention of scaling these mountains, which reach an elevation of 2,087 meters, but 'found that the dangers in the form of poisoned traps and unfriendly peoples prevented my obtaining carriers and other assistants to accompany me. There, no one will cross the land of another unless the owner is willing to guide him. Difficulties such as these could be overcome if they were anticipated and time allowed to overcome them. The Calamianes.-Many of the species known from this group of islands have proved to differ from those on surrounding islands. A new genus of rodents is reported in this paper and very probably other forms new to science await di~covery. On Busuanga I recently discovered a new genus of primitive ribbed frogs of a family unknown to any other East Indian island. This may bespeak a remnant fauna of greater age than that of the surrounding islands. Penon de Coron Island, because of the remarkable character of the limestone rocks, suggests desert conditions in spite of a heavy rainfall. The absence of soil has modified to a great extent its tropical flora, and may have brought about decided variation in its mammalian fauna. This island, due to its geological formation, is a formidable and forbidding territory, for the coast for the most part is a vertical face of rock, 200 to 400 feet high, deeply undercut. There are only a few points where one can actually gain access to the island, which is practically uninhabite'd.

INTRODUCTION

Sulu Islands.-A careful study of the fauna of the numerous islands of this group will doubtless give some certain basis for judging the derivation of the faunas of the eastern part of the Philippine Islands, provided the vicarious ups and downs of the Sulu group during recent geological time have not destroyed the evidence of this migration. Minda.nao.-Probably the least known part of this island is the hig,h mountain range along the northwestern coast. This territory is an old island joined in recent times to the mainland of Mindanao. Dinagat and Siargao are northern extensions of these mountains. The localities mentioned above are certainly not the only profitable collecting localities, but in my opinion do represent the places where the greatest number of novelties await the thorough collector, who is able and willing to undergo the hardships necessary to explore these out-of-the-way places. One, however, does not need to be a trained collector to contribute to our knowledge of the mammalian fauna. Probably 'no mammalian or'd er is more poorly , known in the Philippines than that to which the pig and deer belong. Adult skulls of both sexes of these animals, as well as those of the young, are greatly desired; and the sportsman will do a great service to science if he will remove the skin and skull of specimens he kills and ship them to the Bureau of Science' in Manila, or to some other museum where such specimens are wanted. Only specimens with careful data are desired. The following directions, taken from Hollister, * give the technic of collecting these larger forms: The average traveler little realizes what he may do for zoology. It is in the power of sportsmen, especially, to enrich museum collections, at little trouble to themselves, with the many valuable specimens which come into their hands, and for which they have no personal use. The skins and skulls of all animals killed should be carefully preserved and deposited, with full data, in some institution of standing; it will surprise the average sportsman to find how gladly they are received and what scientific interest is attached to material he assumes to be worthless. The making of study skins of small mammals is really a very simple process and one of considerable interest; and the pleasures to be reali70ed from trapping add greatly to a traveler's experiences. A small pamphlet issued by the United States National Museum, (Washington, D. C.) gives instructions for preparing specimens of mammals in the most approved

* Philip.

Journ. Sci. ยง D 7 (1912) 50-52.

PHILIPPINE LAND MAMMALS

styles for scientific study. To the sport.sman, the gl路eatest interest naturally lies in the medium sized and larger mammals, the skins of which, when intended chiefly for study, are easily prepared. A primary incision should be made from the breast to the tip of the tail, and other incisions from the sole of each foot up the leg to the main longitudinal cut. In horned mammals, an additional cut from the base of the head to between the horns, branching in a Y to around each horn is necessary; the skull in these being removed through this opening. The skin is then removed entire, special care being take~ to skin out the feet, tail, ears, and lips. The skin sho'Uld then be well salted, salt rubbed in on the most fleshy parts and around the edges. It is a good plan to separate the skin of the ear from the cartilage and force salt to the tip, salting the feet well also. The skin may then be rolled up for a few hours, as over night, and afterward spread out in the shade to drain and dry. As the drying process advances, the skin may be folded, before too dry, into a convenient bundle for transportation. The skull, which should always accompany the skin, may be disjointed from the vertebrae, care being taken not to cut or otherwise injure its base. The flesh should be roughly cut off, the brain removed through the natural opening at back of skull with a folded and bent wire, and the skull dried in the sun or near a fire. The skin and skull of the same individual should always be tagged with the same number so that there will be no uncertainty as to what skull belongs with a particular skin when the specimens reach the museum. Even when it is not possible to save the skin, or when it is desired for other purposes, the skull of a medium sized or large mammal should always be saved. Large series of skulls from all localities are greatly desired for study and comparison. Where the sportsman wishes to retain the horns they may be sawed off, either alone or with a portion of the bone of the frontal region, and the specimen will still be of great service to the specialist, to whom the teeth and lower surface of the skull are more important than the horns. When this is done a photograph of the skull before the horns are removed should be taken, to accompany the specimen. Weathered skulls, particularly when the full set of teeth is present, are often of great value when their exact locality is known. Specimens should be labeled with the exact locality where the animal was killed, the date, sex, and the collector's name. The length of the animal before skinning, from nose to root of tail, the length of the tail vertebrae, and length of hind foot are valuable measurements and should be taken if the time and conveniences are available. Bats are always important and may be preserved entire in spirits, or in a weak solution of formalin; a small incision in the belly will allow the fluid to enter the cavity and aid in preservation. Alcohol and formalin must not be used at their full strength. Add to commercial alcohol (95 per cent) one-fifth its volume of water. Add to commercial formalin 15 times its volume of water. LOCAL NAMES FOR MAMMALS

The following names have been taken in part from published literature and in part obtained directly from peoples in various parts of the Archipelago. Most of the names have been given

INTRODUCTION

a generic rather than a specific identification since only occasionally, and this among the forest dwelling peoples, are specific differences recognized. Thus, for example, the name ambao and its variants may be applied to many species of Rattus and allied genera. ABABA. Sus (Camel). * ACNO. Pithecus. Visayan (Elera). AHM-MO. Pithecus. Lanao Moro (Mearns). AHMBOUGH. Rattus (generally) ; Lanao Moro (Mearns). ALALACSIN. Sciu?·us. Samar and Leyte. ALAMlD. Pa~·ado,,·urus. AMAC. Tarsius. Visayan. AMBo. Rattus. Tablas. AMBAO. Rattus. Manobo, Bagobo. AMO. PithecuB. AUGBo. Rattus. Cuyo. AYONG. Pithecus. Cagayan (Elera). BABUY. Sus (generally). BACULAO. Pithecus. Visayan (Oamel). BAKTIN. Sus (domestic). Tablas. BALAC-BALAC. Cynopterus (Elera) . BALATNA MATINIC. Thecurus (Elera). BALINTONG. Manis. BALECON. Manis. Cuyo. BAYONG. Sus (Camel). BINATANG. Sus. Coast Manobo. BISING. Sciurus. Cuyo. BO-OT. Tarsius (Camel). BOOOT. Tarsius (Elera). BOTOTOC. Pithecus. Visayan (Elera) . BUBUIT. C1'ocidura and Pachyu7·a. (Sometimes Mus.) BUCALAO. Pithecus. Mindanao (Elera) . BULILIT. C7'ocidura and Pachyurl1, (Elera) . BUTA-BUTA. Chaerephon (Elera). Bu-uc. Probably same as bu-ut. BU-UT. Cmteromys schadenbergi. Igorrote (Whitehead). BUYATAT. ?

* Names

(aper montanus) BAYONG. Sus (Camel) . CAABUIT. ? CABAG. Small and medium-sized bats. Also ca,big. CASAT-CABAT. Rhinolophtts and Hipposideros. CABUG-CABUG. Bats (Camel). CABAYO. Equus. Common corruption of the Spanish name. CABAO. Ca1·abao. Manobo. CAGO. Cynocephalus volans. CAGS!. SCiU1·US. S. Min dan a 0 (Meyer) . CAGUANG. Cynocephalus volans. Visayan. CAGNING. Rhinolophus and Hippo-

sideros. CALABIDANG. Chaerephon. CALAMBAY. ? CALAPNIT. Small and medium-sized bats. CALIBAGA. Chaerephon and other small bats (Elera). CALUMBIRONG. Small chiroptera. Tablas. CARABAO. Bubalus. CIMMARON. Wild carabao. Northern Luzon. CHONGO. Pithecus. Tagalog. COCANE. Nycticebus menagensis (Elliot) . COLAGO. Cynocephalus volans (Camel) . COGLE. Rattus 01' Crocidura (Camel). COLALAPNIT. Bats (Camel). CUAYAPNIT. Smaller Chiroptera. Tablas. CUYAYAPNIT. Smaller Chiroptera. CUFANIT. Smaller Chiroptera. CUT!. Felis. Cuyo.

in parentheses are authorities for the native names.

PHILIPPINE LAND MAMMALS

DAGA. R'lJJttus (generally). DINGALONG. ViverrlJJ (Camel). DINGUIN. Aonyx cinerelJJ. Palawan. DURIAN. Thecurus. Cuyo and Palawan. DUGUIANT. Thecurus. DUGIAN. Thecurus. Palawan. DURIANT. Thecurus. EUT-EUT. fhlmomys. Igorl'ote (Whitehead) . GAGUA. CynocephIJJlus volans. GIGUA. CynocephaLus volwns. GADIA. Elephas (Camel). GARIA. Elephas (Camel). Elephants, formerly in J 010. HALINTONG. Manis j(wanicus. IRO. Canis, used generally for dog. IYAM. Dog. Mindanao Visayan. ILo. Paradoxurus. Tablas. KAHBOOG. Pte?'opus and Acerodon. Lanao Moro. KURING. Cat. MindanaQ Visayan. KOH-LAGH-CHEWEY. Urogale everetti. Bagobo (Mearns). LANGAM. Crocidura, Pachyura (Elera) . LAMIRAN. Paradoxurus? (Camel) . LAMPOG. ParlJJdoxurus. Loo TOONG. Pithecus. Coast Manobo (Mearns). LUNIPAO. Rattus. Coast Manobo. MACHING. Pithecus. Tag a log (Elera) . MAGO. Tarsius. Samar. MAGAU. Tarsius. Leyte. MAGATILOKOK. Tarsius. Coast Manobo. MAJO. Tarsius (Camel). MALMAY. Tarsius. MANANIR. Pithecus. Visayan (Camel). MANTURON. A~'ctictis. Palawan. MILO. Felis. MIMMAL. Cynocephalus "IJolans. Samal. MIRo. A?'ctictis. MUNDO. Paradoxurus. MUNGAo. Paradoxwrus.

MUSANG. Paradoxurus (Whitehead). Also Viverra tangalunga. P AGIL. SUS (Camel). PANIQUE. Pteropus and Acerodon. PANTOG. Mydaus. Palawan. PANTOT. Mydaus. Palawan. PARRETT. Phlmomys. PASIMUNDOT. Crocidura and Pachyt~ra.

PILANDOK. T?'agulus. Balabac. POLATUCA. Sciuropterus. Palawan. QUINGUA. Cynocephalus vol a n s (Elera) . REEAH. Rattus. Lanao Moro. SALADUNG. Cervus. Cotabato Moro. (EleSAMBO. Pithecus. Visayan ra) . SANGIO. C?'ocidura and Pachyura. SINGALONG. Felis and Viverra. TAGNAN. Sciwropterus. T AGBANUAS. Crocidura and Rattus. Southern Palawan. TAHDEEAHN. Apomys and probably Ra.ttus. Bagobo. TALIBATA. Batop (Camel). TALIBATAB. Megaderma. spasma (Elera) . TALONON. Sus (wild). Tablas. T APILAC. Sciuropterus. Paia wan. T AMARAO. Bubalus mindorensis. Mindoro. T AMARAN. Bubalus mindore.n8i..'!. Mindoro. TARRA BAHBOOEY. Urogale and Podogymnura. Bagobo. TINUCAY. Tarsius (Elera). USA. Cervus. UMBAO TALIKOOBUNG. Rattus tagulayensis and albigularis. Bagobo (Mearns) . UMBAO KAHWEE-LEE. Rattus. Bagobo (Mearns). UMBAO OMBBOISAY. Rattus and Bullimus (Mearns). UMBAO TAKOBO. Rattus and allied genera. Bagobo. VILle. Shrews (Camel). VISIN. Tupaia, Sciurus (Elera).

INTRODUCTION

DISTRIBUTION AND FAUNAL RELATIONS

In a study of the distribution and faunal relations of the Philippine mammals the following assumptions and facts must be taken into consideration: That animals may be transported on natural rafts whose movements depend on water currents or prevailing winds. That animals are transported as pets or as domestic animals from island to island. That flying forms may be assisted by strong winds and typhoons to make longer flights than normal. That larger forms, such as deer, pigs, and buffaloes, may swim no inconsiderable distances from island to island. That interbreeding of differentiated species may occur when brought together by intention or chance (Cerws, Pachyura, etc.). That domestication tends to bring about abnormalities and profound changes in certain forms (deer, dogs, etc.). That a given island or territory did not receive its fauna at a single migration, but at intervals through longer or shorter periods of time. That faunas, of continuous land masses that later become broken up into archipelagoes, tend to have the original species differentiate into island species, that differ from the original forms to a greater or less degree. That the absence of a species from an island does not necessarily postulate its absence there in the past. That the evolution of mammals on island groups apparently takes place with greatei1 rapidity than on larger land masses. That some of the present land masses show strong evidence of former connections with other islands; while others have been recently broken up into smaller areas or have been below the sea level; certain larger islands have been formed by the union of smaller islands.

There are a number of problems of distribution of Philippine mammals that are no longer puzzling if one accepts the above assumptions. These may be tabulated as follows: The number of endemic genera apparently confined to Luzon. The endemic genera occurring in Mindanao. The distribution of Tarsius, Nycticebus, Cynocephalus, Nannosciurus, Podogymnura, and Urogale in the southern and southeastern parts of the Archipelago. The presence of Sciurus and Sci~~ropterus in both Mindanao and/or Basilan and Palawan. The presence in the Palawan group of the genera Mungos, Manis, Thecurus, Arctictis, Mustela, Mydaus, Aonyx, Tupaia, Trag'Ulus, and Insulremus. The presence of distinct species of Rusa and Bubalus in the Calamian Islands and their absence in Palawan.

PHILIPPINE LAND MAMMALS

It has been customary * to assume that a double migration has taken place; namely, an older migration from land to the south of the Philippines, one route from the present Borneo land mass; another route from the region now occupied by Celebes; and a second later migration from Borneo through two ancient peninsulas; one the Balabac-Palawan-Calamianes-Cuyo peninsula; the second the Tawitawi-Jolo-Basilan peninsula. The entrance of the older fauna at some early Pleistocene orpre-Pleistocene era has allowed sufficient time for the development of the endemic genera from the primitive migrant stock. The failure of newly developed species to become generally distributed may be attributed to the fact that land areas from the time of their development have been discontinuous or that the species having developed in high mountains, low lying land connections have been disregarded. Thus we have a ~eries of endemic genera apparently confined to the northern half of Luzon, and a like series in Mindanao,. chiefly high mountain forms that do not become generally distributed even when the land masses of the islands themselves increase.t That we postulate a northward migration is due to the fact that no evidence to be gleaned from a study of the faunas shows definite relationship with Formosa and the islands to the north. On the other hand there is some evidence to show that the presumed earlier migration was not necessarily earlier but that it reached the Philippines (exclusive of the Sulu, Palawan groups) through land bridges via Sumatra-Java-Celebes, excluding Borneo completely in the route. Such genera or species held in common by Borneo and the Philippines (exclusive of the Sulu, Palawan groups) were received by Borneo from the same source from which the Philippines received them. The question of the relation of the fauna of the Palawan group, however, appears simpler and is due to direct migration from the mainland of Borneo, via land bridges, at a time when Palawan and the Calamianes were a peninsula of that island. The forms held in common by Palawan group and the remainder of the Philippines were, (excepting in case of rafting or other

* See Dickerson et al., Distribution of Life in the Philippines, Bureau of Science Monograph 21 (1928). t Mindanao has in Recent or very late Pleistocene times been formed by the union of four or more smaller islands.

INTRODUCTION

3ccidental means of transportation, or transportation by man), derived from different sources, as suggested above. The distribution of Tarsius, Cynocephalus and Nannosciurus in the Southern part of the Archipelago to Mindanao, BasiIan, Samar, Leyte, and Bohol, * tends to show connections among this group by recent iand bridges. This seems all the more probable when one takes into consideration the fauna as a whole, particularly Aves, Reptilia, and Amphibia. To return to our consideration of the northern part of the Palawan group, we find that the Calamianes appear to have retained certain forms t not found on Palawan or Balabac. A -species of Cervus and one of Bubalus occur which are wanting in Palawan; also the genus Insul;;emus that has not been discovâ&#x201A;Źred in Palawan. The retention of these forms in these smaller islands, points to the fact that the land comprising the Calamianes group is older; that is, was not submerged so recently as Palawan proper. The relationship of the fauna for the most part seems to be with the Asiatic continent. However, several of the rodent genera seem to show relat ionship with New Guinea and Australia. Dicke.rson suggests that natural rafts from the latter countries, carried by the Kuro-siwo current, might account for this eastern Malasian element. One would presume that any land bridges would have admitted representatives of the Marsupials among the mammals, as well as members of the populous family Hylidre in the amphibia. Since representatives of these populous families are both wanting it would seem that continuous land bridges between New Guinea and the Philippines have never existed. SPECIES ERRONEOUSLY CREDITED TO THE PHILIPPINES

The most recent work on Philippine mammals, that of Elera,:j: contains a large number of species which do not occur in the Archipelago. They are listed in the order in which they appear in this work. Hylobates funer"tls. Jo10? Semnopithecus albipes. Jo10. Cynopithecus othiops. Jo10, Negros, and Guimaras.

* N annosciu1'US

is not known from Bohol.

t Barbourula, a new genus of discog10ssid frogs, was discovered on Busuanga of the Ca1amianes group by the author. :j: Contribuci6n a la fauna Filipina. (Manila (1915).

PHILIPPINE LAND MAMMALS

M[acacus] nemestrinus. Luzon. N[ycticebus] tardig1·adus. Bohol, Bungau, Tawitawi. T[arsius] spect'r um. Leyte, Bohol, Mindanao, Luzon? Samar. Pteropus edulis. Samar, Dinagat. Pte?'opus keraudrenii. Filipinas. Pteropus niger ·6 philippinensis (plate). Cynopte?'us marginatus. Filipinas. Cynopterus melanocephalus. Macr'oglossus minimus. Panay, Samar, Cuyo, Luzon. Rhinolophus luctus. Luzon, Paragua, L€yte, Mindanao. Rhinolophus rufus. Luzon (Manila). Phyllorhina larvata. Filipinas. Vespertilio capaccini. Filipinas. Vespertilio muricula. J 010. Miniopterus sch?·iber·sii. Luzon, Samar. Emballonura monticola. Filipinas. Taphosous melanopogon. Luzon, Mindanao. Nyctinom'Us plicatus. Luzon. Galepithecus r:ufus. Bohol, Samar, Mindanao. Galeopithecus variegatus. Leyte, Samar. TupaJia ferruginea. Culion, Paragua. Tupaya javanica. Mindanao. So'rex (Crocidura) indicus. (plate.) Crocidura caerulesceus. Manila, Ilocos, Bulacan, Rizal, Samal, Leyte, J 010, Marinduque. Crocidura murina. Luzon, Paragua. Pteromys petaurista. Paragua. Pteromys petaurista var. paUida. Paragua. Sciuropteru.s melanotis. Mindanao, Balabac. lEosciurus affinis. Calamianes, Culion. Heterosciurus lateralis. (Nomen nudum.) Samar, Paragua, Bohol, Culion, Calamianes. Nannosciurus exilis. Basilan, Mindanao. Funambu7:us vittatus. Samar. Bandicota gigantea. Luzon. Phlmomys albayensis. (Nomen nudum.) Poephagomys ater? Filipinas. Felis javanensis. Panay, Negros, Cebu, Paragua, Calamianes. Viverra zibetha. Paragua, Negros, Panay. Viverri(}ula ?ruLlacensis. Filipinas. Paradoxura nigrifrons. Luzon, Cagayan. Paradoxurus crossi. Ilocos Norte. Artictis binturong. Paragua, Balabac. [Canis pollokensis]. (Name for domestic dog in Mindanao.) Mydaus meliceps. Paragua, Calamianes, Balabac. Lutra SU?ruLtrana. Paragua, Balabac. Tragulus javani(}us. Balabac. Trwgulus fulviventer. Balabac. Tragulus pygm;eus. Balabac. Sus vittatus. Filipinas. Sus papuensis. Filipinas.

INTRODUCTION

Elera, in a previous work, Catalogo Sistematico de Toda la Fauna de Filipinas, lists also Macacus speciosus, Sulu; Pteromys inornatus, Palawan; Vespe?'Ugo noctula, Luzon; Vesperugo maurus, Luzon; and Vespertil.io chinensis, Luzon. It is not improbable that a few of these species may be found in the Philippines. However, the bulk of the names listed are errors of identification and should hold no place in the Philippine "faunal lists. * A few of the names are synonyms.

* Note list of species excluded by Sanchez, Anal. de la Soc. Espanola de Rist. Nat. 29 (1900) II 177-290; and Hollister, Philip. Journ. Sci. ยง D 7 (1912) 4'6.

SYSTEMATIC CONSIDERATION The arrangement and interpretation of the larger groups of the Mammalia are still in an unsettled state, and the number of orders of mammals varies with each systematist who treats of them. Older systematists were prone to place together in single orders animals of diverse structure and apparently only remotely related. . The old Edentata is a typical example, as well as the present Marsupialia. The latter group contains all the marsupial forms, which are as diverse as the orders of true placentates. By this same criterion one might place all the true placentates in an order "Placentalia." The tendency at the present time, however, is to separate groups that are not related definitely, in spite of the fact that it may be necessary to create an order for a single species. This is the logical course, provided the structure warrants; hence such orq,ers as Dermoptera, Tubulidentata, etc. The systematists are frequently just as likely to disagree on what constitutes a species. The old criterion was that one form unable to produce a fertile hybrid with other closely related forms merited specific designation; forms having the ability to produce fertile crosses were regarded as subspecies providing certain other characters could separate them. Forms occurring on islands, although closely related to those on other nearby islands which did not cross with the neighboring form because of water barriers, were usually considered as species in spite of the fact that they would cross if brought together. At the present time many zoOlogists no longer consider the above criterion important since occasionally forms of different genera, possibly also of different families, may cross. I have no doubt that the present so-called races of mankind are in many cases species as distinct as those recognized in other groups of mammals. The fact that they are fertile inter se has no real bearing on the question. The choosing of a single common name for animals occurring in a country where so many languages and dialects are spoken as in the Philippines seems a futile course, and I have not done so. Only among some of the more primitive Filipinos is any 64

SYSTEMATIC CONSIDERATION

recognition given to different forms of the rats, bats, and insectivores that occur in one locality. A generic designation such as "bat" or "rat" suffices for a large number of species having certain characters in common. However, on page 57 .is given a list, far from complete, of native names of Philippine mammals. The mammalian fauna of the Philippines, while especially diverse in certain orders of small mammals, notably Chiroptera and Rodentia, is lacking or only poorly represented in many of the orders of larger mammals. The following orders of land forms occur: Lepidota, 1 species; Insectivora, 12; Dermoptera, 1; Menotyphla (here considered as an order' distinct from the Insectivora), 4 species; Chiroptera, 77 species and sUbspecies; Primates, 9; Carnivora, 13 forms (including the dog and cat) ; Rodentia, 75 species and subspecies; Artiodactyla, 21 species (including the domestic carabaos). My failure to treat of the sea mammals in this work is largely because of the fact that practically no material from these groups is present in collections. There have been reports of various whales, porpoises, dolphins, and sirenians in Philippine waters, but for the most part the reports a1!'e from field observations and one is not certain that identifications are correct. A few forms ' certainly occur. Of the Sirenia the species Dugong dugon Muller is rather well known along the southern Mindanao coast. A mounted specimen appears in the Santo Tomas Museum, Manila. The following whales are listed by Elera:*

Bahena australis, Baltenoptera swinhoei, Baltenoptera mindor'ensis, Catodon macrocephalus, Balmna mysticetus; dolphins and porpoises, Delphinus longirostris and Steno chinensis. In Science of May 22, 1925, Albert W. Herre, then chief of the division of fisheries, Bureau of Science, Manila, reported a specimen of Balmnoptera rostrata in Manila Bay. The specimen had stranded at Bacoor, Cavite Province. This at least must be considered as an authentic identification. Specimens of whales are not infrequently stranded, but rarely is anyone available to identify them. Careful photographs of the animal as a whole, as well as the parts of the animal, such as the mouth, eyes, and fins, and careful measurements of the body, fins and their relative position from the snout, should be made together with notations as to color, etc., and these sent

* Elera, 269453-6

Cont. a la Fauna Filipina (1915) 209-219.

PHILIPPINE LAND MAMMALS

to the Bureau of Science, Manila, or to the United States National Museum, Washington D. C., for accurate identification. It is only by this means that the aquatic mammalian fauna may become known. When possible the skuIIs should be preserved and sent.

Class MAMMALIA Key to the Philippine orders 0/ land ma-rnmals. a '. Body covered with scales; teeth lacking (pangolin) ____ LEPIDOTA (p. 67). a '. Body covered with fur, lacking scales; teeth always present. b 1. Forelimb modified into a true wing, with greatly elongated digits (bats) _________________ :. __ .__ .. _.... ___________ .. _________________ .__ ._ CHIROPTERA (p. 107). b'_ Forelimb not modified into a true wing. 0'. Gliding forms, with a membrane between hind limbs involving tail; also a lateral membrane from arm to leg (caguang) DERMOPTERA (p. 91). c ' . Terrestrial, arboreal, or fossorial forms usually lacking lateral membrane (present in some Rodentia), and invariably lacking membrane between hind limbs involving tail. d 1. Small- or medium-sized forms with a single pair of路 greatly elongate chisel-like incisors in upper and lower jaws, separated from the molar-premolar series (rats, squirrels, porcupines, etc.) _____ . ______________________________________________ RODENTIA (p. 366). d ". No pair of chisel-shaped teeth alone present in anterior part of upper and lower jaws_ e '. Functional digits of fore and hind limbs reduced to two; digits with hoofs instead of claws or nails (buffalo, deer, pigs, etc.) _________________ .________________________ ._ ARTIODACTYLA (p. 493). e 2. Functional digits on limbs more than two, never hoofed, but with claws or nails. /1. Claws alone present. g'. Canine teeth not distinctly differ路entiated from the incisors. h 1. Small shrewlike forms, heavily furred except on the tails which are more or less naked; brain case l'educed in size, terrestrial or burrowing (shrews) INSECTIVORA (p. 71). h '. Somewhat larger, shrewlike or squirrel-like fOl'ms, with tails more or less heavily furred; arboreal. MENOTYPHLA (p. 97). g '. Canine teeth strongly differentiated from incisors; medium-sized burrowing, terrestrial or arboreal forms (civets, weasels, cats, dogs, etc.) ____ CARNIVORA (p. 343). / '. Nails present on digits (claws may be present on some of the digits (Ta?'sius) (monkeys, lemurs, and tars~ers), PRIMATES (p. 323).

LEPIDOTA

Order LEPIDOTA

Body covered with horny imbricated scales except on belly, throat, chin, sides of face, and on inner side of limbs; teeth wanting; tongue elongate, protractile; limbs short with five digits on each foot; no accessory articular processes on lumbar vertebrre but anterior zygapophyses largely developed and deeply concave. The scaly anteaters have long been considered as having the rank of a suborder under the order Edentata. Lane has offered what appears to be the best disposition of this group. He considers the Edentata a superorder under which he associated four orders, one of which, the Lepidota, comprises the scaly anteaters. Superorder Edentata Vicq d'Azyr. Order 1. Tamiodonta Cope. Family Conoryctidre W ortman. Family Stylinodontidre Marsh. Order 2. Xenarthra Gill. Suborder Pilosa Flower. Family Bradypodidre Bonaparte. Family Megalonychidre Zittel. Family Megatheriidre Owen. Family Myrmecophagidre Bonaparte. Family Orophodontidre Ameghino. Suborder Loricata Flower. Family Dasypodidre Bonaparte. Family Glyptodontidre Burmeister. Order 3. Lepidota Lane. Family Manidre Gray. Order 4. Tubulidentata Huxley. Family Oryctel'opodidre Bonaparte.

Lane ':' states, regarding the name Pholidota Weber: A further examination of the literature reveals the fact that the term PHOLIDOTA Weber (1904), comprising the Manidre is antedated by PHOLIDOTA Merrem (Tentamen systematis amphibiorum 1820) applied to the Reptilia. As SQUAMATA Huxley (1872), which also has been frequently used to designate the Manidce, is itself antedated by SQUAMATA Oppel 1811, applied to an order or superorder (Osborn) of Reptilia, it seems necessary to adopt some other name for this group. I therefore propose that the order to which the Manidre belong, be called the Lepidota (Gr. A~7r16oToS. Scaly) .

* Science N. S'. 31 (1910) 913-914. Also see Lane, State University of Oklahoma, Research Bulletin 2 (1909).

PHILIPPINE LAND MAMMALS

Family MANIDlE A single family, having characters of the order, is recognized; this contains the single genus Manis. Genus MANIS Linnreus

Manis LINNlEUS, Syst. Nat. 10 ed. 1 (1758) 36. Pholidotus BRISSON, Regn. Anim. 12 (1762) 18.

Characters of the order. Skull suggestive of a cone with the small end turned forwards, free from crests or ridges; no distinction between orbits and temporal fossre; zygomatic arch usually incomplete, due to absence of jugal bone; no distinct lachrymal; palate long and narrow; pterygoids extend backwards as far as tympanics but do not meet in middle line below; rami of mandible slender, straight; a sharp conical toothlike process projects upwards and outwards from near anterior extremity of upper edge; no clavicles; no third trochanter on femur; ungual phalanges bifid at their terminations; caudal vertebrre long, with strong transverse processes and numerous chevron bones; tongue vermiform, flattened toward tip; no crecum; a gall bladder; eyes and pinna of ear very small; body elongated, narrow; tail elongated, convex above, flat underneath; testes lie in inguinal canal; penis external and well developed; uterus bicornuate, vagina not divided; placenta diffused and nondeciduate. These are burrowing animals that feed almost exclusively on termites. They roll themselves into a ball on the approach of danger. One species, Manis javanica Desmarest, is known in the Philippines and probably is confined to Palawan, and possibly also the Calamianes. MANIS JAVANICA Desmarest

Manis javanica DESMAREST, Mamalogie 2 (1822) 377 (Java); BOURNS and WORCESTER, Occ. Papers Minnesota Acad. Nat. Sci. 1 (1894) 1-64 (Palawan); ALLEN, Bull. Am. Mus. Nat. Hist. 28 (1910) 13 (Palawan); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 35; Proc. U. S. Nat. Mus. 46 (1913) 338; SCHULTZE, Philip. J ourn. Sci. ยง D 9 (1914) 95, pis. 1 and 2. Pholidotus dahlma,nni (1) STEERE, List Birds and Mammals Stoore Exped. (1910) 30 (Paragua). 1 Pholidotus culionen.';Iis ELERA, Contr. a la Fauna Filipina (1915) 274 (Culion). P(holidotus) javanicus ELERA, Contr. a la Fauna Filipina (191fl) 273 (Paragua, Culion).

Description of the species.-Head conical, covered with small scales, forming a triangle; sides of head, chin, and throat without scales but covered sparsely with fine, stiff hair, thicker about

MANIS

ears; nose naked; mouth very small, inferior, upper jaw extending beyond lower; eyes small; ears close behind eyes, made visible externally by a skin fold raised and rounded; body, except on middle line of belly, throat, chin, and inside of limbs covered with large subtriangular scales varying in size; each scale finely striated at base, those low on sides with a strong keel; unsealed areas with short, sparse, fine, stiffened hairs; tail with three rows of very slightly keeled scales above, the median largest; laterally with a single row of folded scales with a strong keel forming lateral edge of tail; below with three rows, the two outer heavily, the inner lightly, keeled; larg-est body scales on posterior part of rump and at base of tail; at base of each scale on its underside two (rarely more) small stiff hairs arise on each side, visible between scales; forelegs with five digits with claws of varying size; middle finger bears the largest claw, which reaches a length of more than 30 millimeters; two contiguous digits bear claws about 22 millimeters long; claws of outer digits both less than 10 millimeters long; claws on toes shorter, median, 25; two, adjoining, 16; outer, each about 8 long. These measurements from a young specimen. Scales on hind legs very heavily keeled. Skull.-Characters of skull very similar to those given for genus. Brain cavity with two bony septa extending inward and partially dividing brain cavity; palate arched, alveolar surface of jaws with a slight keel and a few indistinct tubercles; palatal foramina at extreme tip of palate, small; nasals terminating in a point anteriorly; nasal septa extremely complex. Measurements of Manis javanica Desmarest.* mm.

Total length Tail Length of head to posterior edge of ear Snout tip to eye Width of tail at base Median claw on front foot Lateral claw

765 275 86

52 86 35 28

Remarks.-The Palawan pangolin has been regarded by most authors as being specifically identical with that occurring in Java. Elera has essayed to describe as new a specimen from

* From a large specimen in the possession of Mr. Macasaet, Iwahig, Palawan. This specimen also gives the following scale data: Scale rows around middle of body, 21; between ears, 9; on tip of snout, 3; around foreleg at base, 11; around hind leg at base, 8; keeled scales above on tail, 3; 7 rows on hind leg keeled.

PHILIPPINE LAND MAMMALS

Culion, one of the Calamianes. He also gives Palawan ("Paragua") as a locality, apparently not distinguishing it from the Palawan form. It is, of course, not improbable that the Culion form is specifically distinct, but until material is available from Culion and Palawan for comparison I shall consider the name as a synonym. of Manis javanica Desmarest. I obtained a young specimen at Iwahig Penal Colony, Palawan, in the early part of November, 1923, and took it to Manila. I carried with me a few arboreal termite nests to provide food for the specimen. It lived in captivity for some three weeks, when it was preserved. W. Schultze gives a spirited account of the general habits of the P'alawan form as follows: Upon receiving the pangolin, I offered it various species of large true ants, but it paid no attention to them and refused to eat. I then secured a fresh brood comb from a terrestrial termite nest with its included termites, and the pangolin quickly consumed all the insects. To supply the animal in this way with sufficient food presented considerable difficulties, and remembering the destroyed and partly destroyed nests of the arboreal termites that I had observed in the forests the idea occurred to me that the pangolin was probably responsible for their destruction and that these particular termite路s, to a large degree, supplied the food of the animal. I accordingly secured several of these arboreal termite nests, and placed them in the cage with the animal. The pangolin commenced to break open a nest shortly after dark. Its method of accomplishing this is very peculiar. First, it is necessary to give some idea of the characters of the nest in order better to understand how well the animal is adapted to its food supply. The termite nests are usually subglobose, sometimes being rather irregular in shape, and from 2'0 centimeters to 50 centimeters in diameter. In color they are usually dark brown or black, and externally have a rather ;flaky appearance. The outermost part of the nest is rather thin, and is brittle in texture. The entire interior of the nest is made up of a somewhat porous material that is more or less spongelike in appearance, but not at all sponge-like in texture, being very hard and somewhat brittle. In the region where the queen chamber is located, the surrounding material is still harder, and the passages or cells are slightly smaller than in the external portions. In opening the nest the pangolin at first removes the outer layer from one side by means of its powerful claws. The animal then commences working its way into the :nterior of the nest by inserting its claws into the passages or cells and using them as levers, thus breaking away, in small pieces, the very hard material that makes up the interior of the nest. While doing this work, the animal may assume any positionstanding on its hind legs or lying down on its side, on its belly, or even on its back on the top or on the sides of the nest-depending 0!l1 the location of the nest. As soon as it has worked a short distance into the interior, it reaches the regions inhabited by the termites, and, while

INSECTIVORA

busily engaged with its claws in breaking down the interior of the nest, it keeps its tongue constantly protruded, licking up the termites that are disturbed or dislodge4, in its effort.s to reach the interior of the nest. In this manner, the pangolin hollows out the nest sometimes to such a degree that only a thin crust or shell about 3 centimeters in thickness is left. If the nest be very large, it is generally more or less broken up; small nests are rarely broken, but retain their original shape after being hollowed out completely. The pangolin under observation consumed the contents of as many as four medium-sized nests in one night. Very probably during the dry season, the Palawan pangolin lives mostly on these arboreal termites. In Palawan, the terrestrial termite nests or mounds are mostly found in rather open brush or grass lands and are so hard that the animal could not burrow into them during the dry season. It probably uses them as food supply during the rainy season. The pangolin seems to have poor eyesight, at least during the daytime. Howevf'r, its sense of smell is apparently very acute. Its sense of direction is undoubtedly largely dependent on scent. The animal under observation was repeatedly liberated, and soon after being removed from its cage it raised its head and sniffed in various directions. It then invariably turned toward the nearest forest or thicket and walked away in that direction. When it had decided upon a given course, no amount of turning could deflect it, and after being turned about it always resumed its original direction. Even lifting the animal by its tail and quickly revolving it failed to confuse it. It never turned toward open places or toward the sea. There is little doubt that it depends largely upon its sense of smell in locating termite nests, especially those that are placed high in the trees. If disturbed when walking about, it quickly puts its head between its front legs, turns a somersault, and rolls up into a ball, making a hissing noise when so doing. Its powerful tail, which has a horny pad on the end, is a great aid to the animal in climbing and in hanging on branches of trees. The animal has a very peculiar odor.

Order INSECTIVORA

The species that have long been grouped in the order Insectivora, while agreeing in numerous characters, present considerable diversity in general appearance, structure, and habits, as well as in essential anatomical characters. Recently, a group of species has been placed in a new order, the Dermoptera. The remaining forms have been assigned to two suborders or orders (here treated as orders), approximately of equal rank, either of which, were we dealing with larger animals, would unquestionably be considered of ordinal rank. These two are the Lipotyphla and the Menotyphla. The former contains- the living families Erinaceidre, Soricidre, Talpidre, Potomogalidre, Solenodontid~, Centetidre, and Chrysochloridre; the latter, the families Tupaiidre and Macroscelididre. The Menotyphla differ from the Lipotyphla, or true Insectivora, in having ossified intercentra in the dOl'solumbar region

PHILIPPINE LAND MAMMALS

(unique save in the Dermoptera), a crecum (due probably to the fact that they are more herbivorous than insectiv'Orous), a long pubic symphysis, a comparatively large brain case, the orbit encircled with bone, and a well-developed zygomatic arch. In applying the law of adaptive radiation we might suppose that the factors and conditions of adaptive evolution have been at work sd long among the members of this primitive group, and have caused such wide divergenee that the earlier relationships are all but 'Obliterated and we may be forced to recognize our aerial forms in the related order Chiroptera; the volant-arboreal and saltatorial forms in the Menotyphla and Dermoptera, and the more-primitive, more-generalized terrestrial and fossorial forms in the Lipotyphla. Aside from the two living groups there are two extinct suborders, the Hyopsodonta and the Proglires, the latter being an aberrant diprotodont group having the general appearance of rodents. In the treatment given here the question of these more general relationships need concern us but little and the question of the evaluation of the suborders must be left to authors who deal with the Insectivora as a whole and not to one who deals with a few isolated species such as come within the scope of this work. The order Insectivora as here considered includes the shrews, moles, hedgehogs, and a few other, less-known groups. The forms Vary greatly in habit and habitat. Some are burrowing, some arboreal, some terrestrial, while others are semiaquatic. A few species have beoome almost as domestic as the house mouse. By far the greater number of species are nocturnal and insectivorous. The Insectivora are small mammals having plantigrade or semiplantigrade, generally pentadactylate feet; clavicles are present (one exception in Potomogale) ; there are always more than two incisors in the mandible; the enameled molars have tuberculated crowns and well-developed roots. The extremity of the muzzle usually extends far beyond the end of the mandible and may almost be regarded as a characteristic of the order. The testes are inguinal or abdominal, and are not received into a scrotum (a primitive character) ; the uterus is two-horned; the placenta discoidal and deciduate; and the smooth cerebral hemispheres do not extend backwards over the cerebellum (likewise primitive characters). The differentiation of the teeth into the various groups of incisors, canines, premolars, and molars is less pronounced than

INSECTIVORA

in most other mammals. It is sometimes difficult to differentiate the teeth into the various groups, and one must depend upon the position in the jaw and a study of the milk dentition for satisfactory interpretation of the various dental elements. The character of the palatal ridges differs somewhat in different species, but is not diagnostic. Distribution of the living families of Insectiv ora.

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This order of mammals is entirely wanting in Australia, New Guinea, New Zealand, the Pacific islands, and all of South America save the extreme northern part. One family only, the Soricidre, enters the eastern Malayan islands, and northwestern South America. The distribution within the Philippines of the two families Erinaceidre and Soricidre is of interest. Only one species of the Erinaceidre has been found. This is represented by a single specimen from Mount Apo, Mindanao. It is highly probable that it will eventually be taken on other Mindanao mountains. The Soricidre are represented by two genera; namely, Pachyura, from Luzon, Marinduque, Cebu, and Palawan; and Crocidura, from Luzon, Mindoro, Mindanao, J 010, and Palawan. Both genera are probably more widely distributed in the Archipelago. Key to the Philippine families and genera of the Insectiv 01'a. a 1. Skull without postorbital processes; zygomatic arch slender or wanting; no crecum; symphysis pubis short or wanting; auditory bullre imperfect; terrestrial forms. b'. Crown of first and second upper molars with a central fifth cusp; zygomatic arch present; wood shrews of moderate size. Family ERINACEIDA!J (p. 74). c'. Body covered with fur; without spines; tail long. Subfamily GYMNURINA!J. d 1. Small ratlike shrew, with long, soft fur; upper parts mixed with long, red-brown hair ... _ Genus Podogymnura Mearns (p. 74).

PHILIPPINE LAND MAMMALS

b 2. Crown of first and second molars without the fifth CUSp; zygomatic arch wanting ; small .................................... Family SORIClDlE (p. 76). c '. Small shrews, brownish to slate gray; four small conical teeth behind each anterior upper incisor. Genus Pachyura Selys-Longchamps (p. 76). c ' . Small mouselike shrews darker than Pachyura; three small conical teeth behind each anterior upper incisor. Genus Crocidura Wagler (p. 82).

Family ERINACEIDlE Insectivora with plantigrade feet provided with simple claws. Radius and ulna large, fibula not anchylosed below to tibia; clavicles long, slender; a narrow symphysis pubis; zygomatic arches路 with well-developed pterygoid fossre; malar bones present (rarely absent), suspended in zygomatic arches; a low ridge in front of orbit; postorbital processes wanting; nasals separate; tympanic bone ring-shaped, not forming a bulla; first and second upper molars each with five cusps; central cusp minute, united by a ridge on each side to the bases of the two internal cusps. No crecum. (After Dodson and Blanford.) One genus, Podogymnura, of this fam1ly is found in Mindanao. No representative ha'8 been found elsewhere in the Philippines. Genus PODOGYMNURA Mearns

Podogyrnnura MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 436; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 4; LYON, Proc. U. S. Nat. Mus. 36 (1909) 449.

Related to Gymnura and Hylomys. Dental formula: i. 3- 3, c. 1- 1, pm. 3- 3, m. 3-=-3 = 40 teeth. 3- 3 1- 1 3- 3 3-3 Second incisor smallest; tail robust and moderately hairy, more than one-third the length of head and body; hind foot seminaked, three-fourths the length of head; whiskers long, reaching shoulders; diameter of eye one-sixteenth the length of head; head less than one-fourth of total length; ears large, seminaked, height from meatus more than one-third the length of head; forefoot seminaked above, entirely so below, with four tubercles, one at base of second digit, one at base of fifth, a large vase-shaped one at junction of third and fourth, and a very prominent one at base of palm; hind foot 5-tuberculate, a small tubercle at base of first digit, a large one at base of second, a. vase-shaped one at j unction of third and fourth, a mediumsized one at base of fifth, and a prominent one about midway between heel and middle digits toward inner side. Pelage long, full, and soft, with fur of different lengths. (From Mearns.)

PODOGYMNURA

Only one specimen, the type of PodogymnuTa truei Mearns, is known. It was collected on Mount Apo, Mindanao. The two related genera Gymnura and Hylomys have four premolars in both jaws instead of three. The tail length of Podogymnura IS intermediate between that of the other two genera. PODOGYMNURA TRUEI Mearns

PodogymnuTa truei MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 437; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 5; Proc. U. S. Nat. Mus. 46 (1913) 302; LYON, Proc. U. S. Nat. Mus. 36 (1909) pI. 37 (photograph of the type).

Description of the species.-After the type description.

The general characters of the species are those given for the genus. Color in life.-Nose and toes flesh color; snout, lips, ears, tail, and feet purplish flesh color; pelage of upper parts slate-gray mixed with coarse reddish brown hairs; underparts hoary, slightly mixed with coarse brown hairs; claws horn color, with a reddish brown spot at middle. Meas~~rements

of the type of PodogymnuTa t?'uei Meal路ns. mm.

Zygomatic breadth, estimated; brain case smashed 18 Least interorbital breadth 9 Median length of palate 21 Length of internasal suture 13.6 Greatest length of internasal bone 15.1 Greatest breadth of nasals 2.5 Height of narial opening 4.9 Breadth of narial opening 3.5 Breadth of palate between anterior molars 5 Greatest breadth of palate outside of teeth 10.3 Diastema between first upper incisors 2 Distance between outer border of first upper incisors 3.3 Between upper canines 4.1 Between outer border of upper canines 6.1 From anterior border of first upper incisor to posterior margin of last molar 20 From anterior border of upper canine to posterior margin of last molar 15 From anterior border of first upper incisor to canine 5 United length of three upper molars 6.5 Length of first upper molar 2.5 Breadth of first upper molar 2.6 Length of first upper incisor from alveolus 2.3 Length of first upper canine from alveolus 3.1 From anterior border of first lower incisor to posteriol' border of last lower molar 18 Combined length of three lower molars 7.5 Total length 210

PHILIPPINE LAND MAMMALS

Measurements of the type of Podogymnura truei Mearns-Continued. mm.

Tail Hind foot Forefoot Head, about Ear from crown Ear from notch at base From tip of nose to eye From t.ip of nose to angle of mouth From tip of nose to ear Diameter of eye Tibial joint Forearm

36 23.5 49 13 19 26.5 17.5 32 3

33 25

The species was named for Dr. Frederick W. True, of the United States National Museum. I have seen no specimen of this species, which appears to be known only from the type. The native Bagobo name for the species is given by Mearns as tali-rah bali booey (ground pig). The type, in the United States National Museum, was collected on Mount Apo, Davao, Mindanao, at an altitude of about 1,820 ~ meters, June 25, 1904, by Edgar A. Mearns. Family SORICIDJE Terrestrial shrews covered with soft hair; head elongated, eyes very small, snout pointed and projecting considerably beyond lower lip; ear conch, when present, rounded, resembling somewhat a human ear; skull elongated without postorbital processes and zygomatic arches; tympanic bone annular, not forming a bulla; symphysis pubis wanting; tibia and fibula not united; molars with W -shaped cusps. Two genera of this family are found in the Philippines; namely, Pachyura with three species and Crocidura with eight. Key to the genera of Soricidre known

the Philippines.

Four small conical teeth behind each anterior upper incisor. Pachyura Selys-Longchamps (p. 76). a '. Three small conical teeth behind each anterior upper incisor. Crocidura Wagler (p. 82). a

Genus P ACHYURA Selys- Longchamps

Pachyura SELys-LoNGCHAMPS, Etudes de micro-mammalogie (1839) 32.

Small, long-snouted, mouselike shrews, uniformly slate gray or brownish gray; eyes very small; vibrissre on snout directed

PACHYURA

outward or forwards; an odoriferous gland on side; four small conical teeth behind the enlarg~d upper incisors. Dental for.. . . 3-3 1-1 2-2 3-3 mula for PhIlIppme forms: 1. 2-2' c. 0-0' pm. 1-1' m. 3-3 = 30. Three species of this genus are known from the Philippines. They are Pachyura luzoniensis (Peters), P. occultidens Hollister, and P. palawanensis sp. nov. The first appears to be confined to Luzon. * The second species is known from the western Visayan Islands, Panay and N egros, and also from Mindanao. The third is from Palawan. A species, Crocidura edwardsiana Trouessart, has long been listed under the genus Pachyura, but Hollister t has rightly placed it in the genus Crocidura. Hollister says: The shl'ew originally described as Crocidu'ra edwa?'dsiana 1 is placed in the genus Pachyura in the Supplement to Trouessart's Catalogue, and I followed this disposition of the species in my recent list of Philippine mammal names! I am lately informed by Doctor Trouessart that the type of the Sulu species is an undoubted Crocidu?路a.

Key to the Philippine species of Pachyu?'a Selys-Longchamps.

a'. First premolar not v,isible fr(i)m outer lateral view of jaw. P. occultidens Hollister (p. 77). a'. First premolar distinctly visible from lateral view of jaw. b t. "Canine" larger than third incisor .... P. palawanensis sp. nov. (p. 78). b". "Canine" equal to third incisor ________ P. luzoniensis (Peters) (p. 80). PACHYURA OCCULTIDENS Hollister

Pachyu1'a occultidens HOLLISTER, Proc. U. S. Nat. Mus. 40 (1913) 303.

Description of the type specim.en.-(After Hollister.) Large, in general much like the larger specimens of Pachyura luzoniensis, but skull relatively wider interorbitally; skull very similar to that of the Luzon form, but heavier inter orbitally ; teeth large, crowded, the rows shortened; first three unicuspids large, second only slightly smaller than third; fourth unicuspid greatly reduced, lying entirely posterior to a line between anterior tips of large premolars, and not visible from outer side; large premolar, from this view, in contact with third unicuspid. * A specimen has been recorded from Oebu by Gunther, Proc. Zool. Soc. London (1879) 75, but it is probable that this belongs to the species described by Hollister from Panay, P. occultidens. t Proc. U. S. Nat. Mus. 46 (1913) 303. 1 Trouessart, Le Naturaliste, No. 42, p. 330, 1880. [Hollister's note.] , Philippine Journ. Sci. sec. D, vol. 7, No.1, February, 1912. [Hollister's note.]

PHILIPPINE LAND MAMMALS

ColO1'.-Similar to PachyuT(}, luzoniensis in color; slate gray above and below; snout, ears, feet, and tail brownish. MeaSU1'e1nents of the type of Pachyura occttltidens Holliste?路. mm.

Head and body Tail vertebrre Hind foot, without claws Condylobasal skull length Mastoid breadth Least interorbital breadth Upper toot.h row, entire Mandible Lower tooth row, entire

115 70

19 30.7 13.6

5.7 13.6 17.0 12.2

Remarks.-The type was collected at Iloilo, Panay, August, 1912, by Carroll Fox; it is No. 175761, United States National 路Museum. Hollister examined four specimens. The .species is found on Panay, N egros, and Mindanao. The relatively wider and heavier skull and the size of the second unicuspid teeth differentiate this species from the Luzon species. PACHYURA PALAWANENSIS sp. nov .

. Type.-No. 89, E. H. Taylor collection; collected in Taytay, Palawan, May, 1913; collector unknown. Skin made up from alcoholic specimen, skull. Paratype (No. 90) from the same locality. Diagnosis; description of the tyve.-M uzzle not sharply pointed; anterior part of head distinctly widened due to the swelling of the upper lips, as is typical, particularly in the young of Pachyura luzoniensis; snout with numerous elongate vibrissre directed outward or backward; muzzle naked, snout on sides covered with fine, sparse, short hair; elongate body fur extends forward on snout to a point halfway between eye and tip of snout on a median line, standing as if roached; ear much less prominent than in P. luzoniensis and apparently smaller; fur on body apparently thicker and longer than on head; hands, feet, and tail covered with very fine, short, sparse hair; tail with a scattered growth of elongate hairs directed backward, 5 to 10 millimeters in length; the fine hair grows in circles on the tail giving it an annulated appearance; fingers and toes normal for the genus. Color in alcohol.-Uniform brownish, apparently having faded somewhat as a result of immersion in alcohol for several years. I presume that the color in life is grayish or ashen.

PACHYURA

Skull and teeth.-Skull and teeth similar to those of Pachyu1'a luzoniensis but differing in numerous characters that are difficult to define. Cones of' molars distinctly more declivous, third molar distinctly larger; basal parts of anterior incisors 120

118

124

122

126

Batan Is 20

PHILIPPINE ISLANDS

® Pachyura occultidens ® Pachyura luzoniensis © Pachyura palawanensis Crocidura parvacauda ® Crocidura halconus ® Crocidura beatus @ Crocidura grayi ® Crocidura mindorus CD Crocidura palawanensis Q) Crocidura edwardsiana ® Crocidura grandis

@ 18

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120

FIG. 2.

122

124'

126

Distribution of Soricidm in the Philippines.

fail to show such short curves as those of P. luzoniensis; basal cusp of first incisor with a distinctly higher elevation than the corresponding part in P. luzoniensis. In the mandible, the molar series, particularly the third molar, is distinctly more elongate in P. palawanensis than in P. luzon-

PHILIPPINE LAND MAMMALS

iensis; the first incisors are slenderer. other small differences.

There are numerous

Measurements of the type and the paratype of Pachyura palawanensis sp. nov. No. 89 No. 90 Total ' length Head and body Tail Hind foot Forefoot Ear: Greatest height Greatest length Diameter of eye Skull, condylobasal length to incisor tip Breadth of brain case Orbital constriction Maxillary tooth row to InCISOr tip Mandible, total length to incisor tip Mandibular tooth row to incisor tip Height of coronoid process

mm.

139.5 85.0 54.5 18 12

140.2 86.2 54.0 18 12

9.5 8.5 1.2 27 11.6 6 12.1 18 11.5 7.8

9.4 8.5 1.2 30 12 6 12.5 19.5 12 8

Remarks.-Very little variation can be noted between the two specimens. N either has the basal suture of the skull fully coalesced and the teeth show but little wear. It may be noted that the orbital constriction is less and the breadth of the brain case is greater than in P. luzoniensis. It is not surprising to find this genus in Palawan, since it has long been known from Borneo. PACHYURA LUZONIENSIS (Peters)

Crocidura (P.) luzoniensis PETERS, Monatsb. Akad. Wiss. Berlin (1870) 595. ? Crocidura luzoniensis GUNTHER, Proc. Zoo!. Soc. London 1879) 75 (Cebu). Pachyum luzoniensis HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 5; Proc. U. S. Nat. Mus. 46 (1913) 303.

Description of the species.-(From No. 28, E. H. Taylor collection; collected at Manila, Philippine Islands, December 8, 1920 by E. H. Taylor.) Muzzle sharply pointed; nostrils elongate, rather tubular, surrounding integument naked; snout covered with short hair with numerous elongate vibrissre directed backward and outward; mouth ventral; eyes very small, concealed in fur; ears small, rather flattened against head, slightly resembling a human ear; cartilage of ear naked; pelage soft; small space behind ear naked; tail slightly longer than half

PACHYURA

the length of head and body; hair on tail arranged in whorls, giving the tail an annulated appearance; scent glands well developed, on sides behind foreleg; tail with scattered short hair and with numerous long hairs directed backward; feet with fine short hair extending to claws; hairs about claws elongated; lips glandular, much thickened above, giving snout a flattened appearance. Color.-Above grizzled ash gray with brownish suffusion anteriorly; not perceptibly darker below; hair on feet brownish; tail brown, the hairs occasionally tipped with white. Skull and teeth.- Skull narrow, elongate, with a slight sagittal crest, and with strong occipital (lambdoid) crests; anterior lower incisor with a strong secondary cusp; second incisor rests against cingulum of first; canine not differentiated from premolars; anterior premolar very small; a large preorbital fossa. . 3- 3 1- 1 2- 2 3- 3 Dental formula: 1. 2-2' c. 0- 0' pm. 1-1' m. 3-3 = 30. MeasU7'ements of Pachyura luzoniensis (Peters). mm.

Total length Head and body Tail Longest hairs on snout Hind foot and cIa ws Snout from eye Skull, condylobasal length, with incisors Breadth of brain case Orbital constriction Mandible length to incisor tip Maxillary tooth row Mandibular tooth row

208 140 68

28 22 18 30.5 13 7 19 12.5 12

RemaTks.-The types were collected in Daraga, southern Luzon, by Jagor, and one in Manila by V. Martens. Hollister states: ... There is an immense and most puzzling variation in size in a large series of specimens from Manila Bay, but after long study I am forced to believe that it is genuine individual variation within a single species.

About half of the specimens that I have examined were distinct ashy to grizzled gray, with the tip of the snout, the feet, and the tail showing a brownish color. The other half were distinctly brownish on the back as well as on the extremities. Hollister gives the measurements of thirteen males and

* Proc. 259463---6

U . S. Nat. Mus. 46 (1913) 303.

PHILIPPINE LAND MAMMALS

six females, all fully adult. The following variation is apparent. Minimum and maximum measurements are given: MeaS1,拢1'ernents of Pachyura luzoniensis (Pete?路s). Males . mm.

CondJ>:lobasal length Zygomatic breadth Mastoid breadth Interorbital breadth Palatal length Mandible Maxillary tooth row, entire Mandibular tooth row, entire

28.5-36.0 8.7-10.2 11.9-14.5 5.3- 6.1 12.1-14.7 14.8-17.9 12.4-13.7 11.5-12.5

Females. mm.

25.7-30.7 8.4- 9.5 11.0-13.2 5.0- 5.7 11.6-13.0 14.1-16.5 11.6-13.1 10.7-12.0

This unusual cranial variation is not associated wit.h constant external chara~ters and decided variation occurs in the measurements of individuals belonging to the various color phases, which range from gray to brownish. Hollister has suggested that this may be due to an admixture of various races from shipping in the harbor. The species is domestic and is a common nuisance in towns and cities in the island. It lives about foundations of houses and under sidewalks and the shrill cries can be heard at night as it forages indoors or outdoors for its evening meal. The mammre are 3-3, located in the inguinal region. The described specimen has two teeth missing in each upper jaw. These correspond to the first premolar and the canine; . all the teeth show great wear. The common name for the species in central Luzon is bu-ot. I collected the young of a species of Pachyura on Marinduque that I believe are of this species. Genus CROCIDURA Wagler

Crocidura WAGLER, Isis von Oken (1832) 275; FLOWER and LYDEKKER, Mamm. Liv. and Ext.inct (1891) 626; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 5.

Small mouselike shrews darker in color than Pachyura. Male or female organs forming a . short cloaca with anal orifice; tail long with a mixture of long and short hairs; ears moderately large. Teeth white. 1-1 1-1 3-3 . 3-3 Dental formula: 1. 2-2' c. 0-0' pm. 1-1 m. 3-3 = 28. The known ,s pecies of the 路genus are all small and rarely have a head-body length exceeding 100 millimeters. Tail never as long as head and body.

CROCIDURA

Key to the Philippine species of CTocidu1'a Wagler. a '. Smaller species. b'. Above brownish, head and body, 61 millimeters. C. paTvacauda sp. nov. (p. 83). b'. Above dark sepia, darker posteriorly; head and body, 74 millimeters. C. halconus Miller (p. 85). b". Above brownish with black suffusion; head and body, 77 millimeters. C. beatus Miller (p. 85). b '. Above uniform red brown; head and body, 77 millimeters. C. grayi Dobson (p. 86). a'. Larger species. b '. Similar to C. halconus .. head and body, 91 millimeters. C. mindorus Miller (p. 87). b '. Fur with slaty tinge; head and body, 95 millimeters. C. grandis Miller (p. 88). b 3. Fur slaty gray with brownish suffusion; head and body, 82 millimeters ________________________ ______ __ ______ ______ C. palawanensis sp. nov. (p. 88). b '. Fur uniform black, passing to a deep ash below; head and body, 84 millimeters ____ .________ .__________________ C. edwardsiana Trouessart (p. 90).

Eight species of CTociduTa are recorded from the Philippines. Crocidura grayi Dobson, from Luzon; C. halcomts Miller, from Mount Halcon, Mindoro; and C. beatus Miller, from Mount Bliss, Mindanao, are closely related. Crocidura mindorus Miller, from Mount Halcon, Mindoro, and C. g1'andis Miller, from Mount Malindang, Mindanao, are likewise related. It is significant that C. halcomls and C. mindorus are from the same type locality; C. beatus and C. grandis are likewise from the same locality. Mearns obtained a specimen of C. halconus at Bulalacao, Mindoro, which proves that C. halconus is not confined to high mountains since Bulalacao is near sea level. Two new species are described in this paper. The type of C. palawanensis is the first record of the genus from Palawan. CROCIDURA PARVACAUDA

S}).

nov.

Type.-No. 737, immature female, E. H. Taylor collection; collected at Saub, Cotabato, April 27, 1923, by E. H. Taylor. Skin (dry) and skull. Diagnosis; descTiption of the type.-Diminutive shrew; snout produced, tip bare, divided by a deep groove that extends as deep as naked portion of snout, giving the latter a distinct Yshaped appearance; anterior part of head very slender, lips not, or but slightly, thickened; ear small, upper part of conch extending much anterior to auricular opening; snout covered with thin short hair, averaging from 0.5 to 1 millimeter in length; vibrissre numerous, the longest extending behind ear;

PHILIPPINE LAND MAMMALS

a few short vibrissre on lower jaw; body covered with dense short fur of uniform length; fur on forehead begins suddenly at a point 7 millimeters from snout tip and laterally at a point approximately 11 millimeters from snout tip; tail sparsely covered with short hair 0.5 to 0.75 millimeter long; towards the base, par~icularly on the underside, are a few, more-elongate hairs, the longest about 3 millimeters; legs covered with sparse hair not longer than that on tail except hair at distal end of phalanges which is longer than claws. ColO1'.-Almost uniform grayish to light seal brown, both above and below as well as on extremities; abdomen a slightly grayer shade; unhaired tip of snout dark blackish brown; individual hairs on back silver basally, gradually merging into slaty gray, which in turn changes to brownish on distal fourth of hairs. M easurements of the type of Crocidura parvacatuda sp. nov. mm.

Total length Head and body Tail Snout to ear Height of ear, moistened Widt.h of ~ar, moistened Hind foot to claw tip Skull, condylobasal length Width of brain case, approximate Depth of brain case, approximate Maxillary tooth row Molars and premolars, upper Width of premaxillaries Width between third molars, upper Length of mandible to incisor tip Height of coronoid process Length of lower tooth row Length of lower molar row

96 61 35 16 6

5 14.2

20 9

5.5 9.1 5.1 2.5 2.5 15 5.8 9

4.5

Remarks.-Crocidura beat~ts Mearns and C. grandis Mearns have been ' described from Mindanao. Both of these differ from the described specimen in having much longer tails. In C. beatus the tail equals 71 per cent of the total length of the head and body; in C. g1"andis the proportion is 70 per cent, while in the present form the tail is equal to only 57 per cent of the head and body length. The specimen is obviously immature and the brain case is badly broken. The condylobasallength exceeds that of Crocidura beatus. The growth of fur on the head is strongly delimited from the short hair that covers the snout.

CROCIDURA

It does not merge gradually as is the case in C. palawanensis, but the thick fur begins suddenly at a point 7 millimeters from the tip of the snout medially, and laterally at 11 millimeters from the snout on a level with the front of the eye. The single specimen was obtained at Saub, Cotabato, after a heavy flood; it was caught and killed by a native as the skull will testify. CROCIDURA HALCONUS Miller

Croeidu1'a, hCLleonus MILLER, Proe. U. S. Nat. Mus. 38 (1910) 391; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 6; Proe. U. S. Nat. Mus. 46 (1913) 392.

Description of the species.-This shrew is related to CTocidura grayi, but the unicuspid teeth are more contrasted in size. Color.-Entire animal dark sepia, blackening on posterior half of back; underparts tinged with broccoli brown; feet and tail blackish, long hairs on tail a slightly contrasted brown. Skull and teeth.-The skull does not differ appreciably from that of CrGcidura grayi, though the interorbital region seems inclined to be narrower and the brain case less deepened; there is a slight increase in the size of the first unicuspid tooth and a reduction in the size of the second, the third remaining unchanged, while the large premolar and first molar exceed in size the corresponding teeth in C. gmyi. (After Miller.) Meas'U1'e?nents of the type of C?'oeidura hale onus Mille?'. mm,

Head and body Tail Hind foot, dry Skull, condylobasal length Breadth of brain case Dept.h of brain case at middle Mandible Maxillal'Y tooth row, entire Mandibular tooth row, entire

74 54 13 19 9 5 10.2

8.8 8.2

Remarks.-The type and paratypes from an altitude of 1,910 meters on Mount Halcon, Mindoro, were collected by Edgar A. Mearns, November, 1906. Four specimens were taken. They are in the United States National Museum. The type is No. 144652. CROCIDURA BEATUS Miller

Croeidura beatus MILLER, Proc. U. S. Nat. Mus. 38 (1910) 392; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 5; Proc. U. S. Nat. Mus. 46 (1913) 303.

PHILIPPINE LA-ND MAMMALS

Description of the species.-Similar to Crocidu1+a g'rayi and C. halconus, but the color is darker and the skull slightly less robust. Color.-The color is similar to that of C. halconus but the blackish suffusion on the back is more extensive, involving nearly the entire. dorsal surface. Skull and teeth.-The skull, save that it is less robust, resembles that of other Philippine species of Crocidura. The teeth are like those of C. halconus. (After Miller.) Measurements of the type of C?'ocidiura beatus Mille?". mm.

Head and body Tail Hind foot, dry Skull, condylobasal l~mgth Breadth of brain case Depth of brain case Mandible Maxillary tooth row, entire Mandibular tooth row

77 55 14.4 19.2 9

5 10.4 8.8 8.2

Remarks.-Only the type is known; it was collected on Mount Bliss, Mindanao, at an elevation of 1,461 meters by Edgar A. Mearns, May 28, 1906. The type is No. 144647, in the United States Nationa:r Museum. CROCIDURA GRAYI Dobson

Crocidura grayi DOBSON, Ann. & Mag. Nat. Hist. VI 6 (1890) 484; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 5; Bull. U. S. Nat. Mus. 46 (1913) 303 (Haights, Benguet).

Description of the species.-Like C. horsfieldi but considerably larger, although the tail is not longer than in that species, and is similarly nearly naked; the fur (so far as can be ascertained from an inspection of specimens preserved in alcohol) is very similar in length and in color to that of that species; the lateral gland is as well developed as in the males of C. horsfieldi, perhaps somewhat larger; bn the single female available for examination (which is, however, not quite fullgrown) there is no trace of one. Color.-Fur above reddish brown, beneath grayish brown, the basal three-fourths of the hairs somewhat slaty. Skull and teeth.-Skull larger than that of C. h01'sfieldi; teeth * like those of that species, the last upper incisor nearly

* See

Dobson, Monograph of the Insectivora pt. 3, fasc. 1, pI. 28, fig. 11.

CROCIDURA

equal to the anterior maxillary tooth in vertical extent, but less than it in cross section. (After Dob~on.) Measu?'ements of the type of Crocidum gmyi Dobson. mm.

Length, adult male, head and body Tail Ear Elbow to end of middle digit, without claw Hand, without claw Foot, without claw

68 51 10 18 8

13.5

Rema1路ks.-The type, now in the British Museum, was collected by H. Cuming in the Philippines; the exact locality is unknown. Miller reports it from Luzon, and Hollister records one from Haights-in-the-Oaks, Benguet, collected by E. A. Mearns. CROCIDURA MINDORUS Miller

C'rocidu?'a ?nindo?'us MILLER, Proc. U. S. Nat. Mus. 38 (1910) 392; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 5; Proc. U. S. Nat. Mus. 46 (1913) 392.

Description of the species.-Color and general appearance as in Crocidura grayi and allied forms, but size noticeably greater. Color.-So .similar to C. halconus as to need no special description; the blackish suffusion on the back is, however, apparently a little less evident than in C. halcom('s. Skull and teeth.-At once distinguished by its conspicuously larger size, the skull shows no pecularities of form as compared with C. g'rayi; teeth similar to those of smaller species but second and third unicuspids more nearly equal in size. (After Miller.) Measurements of the type of C?'ocidum mind01'us Miller. mm.

Head and body Tail Hind foot, dry Skull, condylobasal length Depth of brain case Breadth of brain case Mandible Maxillary tooth row, entire Mandibular tooth row

91 72 16.4 21.6

15.8 10.0 12

9.8 9.0

Rema1'ks.-The type, from Mount Halcon at an elevation of 1,938 meters, was collected November 19, 1906, by Edgar A. Mearns. The type, No. 144654, United States National Museum, is the only specimen known.

PHILIPPINE LAND MAMMALS CROCIDURA GRANDIS Miller

Crocidtwa gmndis MILLER, Proc. U. S. Nat. Mus. 38 (1911) 393; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 5; Proc. U. S. Nat. Mus. 46 (1913) 392. .

Description of the species.-Like Crocidura mindorus but larger. Color.-Color essentially that of Crocidu1"a mindorus; it differs from that of the other Philippine members of the genus in the presence of a slight but evident slaty tinge throughout the pelage; the exact shade is intermediate between hair brown and mouse gray but darker than either; feet and tail an indefinite dark brown obviously different from the blackish color of the tail in other species. Skull and teeth.-Skull slightly larger than that of C. mindorus. The second unicuspid distinctly smaller than the third; teeth otherwise as in C. mindorus. (After Miller.) Measurements of the type of Crocidura grandis Mille?'. mm.

Head and body Tail Hind foot Skull, condylobasal length . Breadth of brain case Depth of brain case Mandible Maxillary tooth row, entire Mandibular tooth row, entire

67 18.6 23 10.2 6 12.8 10.2 9.4

Remarks.-The type, United States National Museum No. 144648, is the only specimen known. It was collected on Grand lVIalindang Mountain, Mindanao, by Edgar A. Mearns. CROCIDURA PALAWANENSIS sp. nov.

Type.-No. 55, E. H. Taylor collection; collected at Sir J. Brooke Point, Palawan, September 29, 1923, by E. H. Taylor. Skin made up from alcoholic specimen; skull. Diagnosis; desc?'iption of the type.-Typical shrewlike habitus; snout pointed (the extreme tip injured), covered with a thick growth of short hair gradually merging into the longer fur of the face; vibrissre numerous, the longest reaching at least 10 millimeters behind ears; a few elongate vibrissalike hairs on underside of lower jaw; both sides of ear covered with fine short hairs; tail with a sparse growth of fine hairs rarely ex-

CROCIDURA

ceeding 1 millimeter in length; a few elongate hairs (2 to 4 millimeters) can be discer ned with difficulty near base; hair at tip of tail about 2 millimeters long; hair on feet and toes sparse, similar to that on tail; ear moderate, upper part of conch reaching considerably anterior to auricular opening. Color.-Fur slaty to ashy gray except at tip, which is slaty brown, so that viewed in certain lights the surface color seems slightly brownish; short hair on snout, ear, and tail blackish brown; area on throat and thorax and extending some distance on belly strong reddish brown to broccoli brown; posterior abdomen grayish; hair on upper part of hand and foot with a brown cast. Skull and teeth.-Teeth not or but slightly worn . . 3- 3 1- 1 1-1 3-3 Dental formula: 1. 2-1 c. 0- 0' pm. 1-1' m. 3-3' = 28. First pair of upper incisors strongly curved, never ~n contact as in Sorex; third incisor distinctly smaller than canine; cusps of second and third incisors more compressed than in the corresponding teeth in PachyU't'a luzoniensis; inferior cusp of first incisor strongly developed; W-pattern strongly pronounced in first and second molars. ,1jI!eastl1'ernents of the type of C1"ocidw'a pala'wanensis sp. nov . 111m.

Total length, in life Body and head Tail Hind foot to claw tip Ear height Greatest length Skull, condylobasal length, to incisor tips VVidth of brain case Greatest width of upper jaw Interorbital width Premaxillary width Length, upper tooth row Length of upper molars VVidth between last molars Length, mandible to incisor tip Height of coronoid process Length, mandible bone Length of lower molar row

154 82 72

16 6.5 9

23.5 10,

7.5 5 3 10

4.3 2.5 15.3 7.1 11.2 4.8

Remarks.-This small shrew was trapped at night in deep forest near the sea at Sir J. Brooke Point, Palawan. The speci-

PHILIPPINE LAND MAMMALS

men is an adult male. The teeth, however, show practically no wear but the basal suture is obliterated. At night I frequently heard shrill cries which I attributed to this species. Only a single specimen was collected. The gland on the side of the body, somewhat behind insertion of foreleg~ is present and well developed. The odor secreted was not so offensive as that of species of PachyuTa occurring in the Philippines, at least not at the time: it was collected. The natives, Tagbanuas, did not differentiate between this species and young rats. CROCIDURA EDW ARDSIANA Trouessart

Crocidut"a edwardsiana TROUESSART, Le Naturaliste 330; HOLLISTER, Proc. D. S. Nat. Mus. 46 (1913) Pachyu1"a cae1"ulea edWa1"dsiana TROUESSART, Cat. (1904-5) 139. Pachyura edwardsiana HOLLISTER, Philip. Journ. Sci.

No. 42 (1880) 303 and 334. Mamm. Suppl. ยง D 7 (1912) 5.

Description of the species.-Ear bent back, exterior and interior covered with very short brown hairs, a little longer on edge of valvules, bare at its base and behind, and well separated from pelage; sides 0:(, snout puffed out and naked up to near eye, covered with long vibri.ssalike hairs; pelage of head elongate on median line up to snout, forming a straight band which is extended not more than 1 or Z millimeters; tail cylindrical, finely annulated, thirty-six annuli to the centimeter, much thickened at base, narrowing gradually to tip; feet long and robust, some black hairs between the scales below digits; an odoriferous gland present on sides indicated by short hair. ColoT.-Color similar to that of Rattus rattus; uniform black above, passing to deep ash below; hair short, of nearly the same color throughout its length; vibrissre on snout and tail white to their tips; claws yellowish; all bare parts of nose, tail, and feet deep brown. Skull and teet h.-Posterior lobe of first upper incisor .shorter than anterior lobe, its summit scarcely reaching the neck of second incisor; third incisor and canine subequal, but the latter a little heavier and its tip reaching that of third incisor; cones of anterior premolars well developed but reaching the level of tip of canine; first lower incisor somewhat wavy on the upper cutting surface; second lower incisor twice as long as high; tip of anterior upper molar not well developed; total number of teeth 28. (After Trouessart.)

GALEOPTERIDJE

Measul'e?ne'nts of C?'ocidw'a

edwa?'ds!~ana

TTouessart, mm.

Length of body and tail Tail Head Ear: Height Width Tip of nose to eye li:ye to ear opening Skull, length

130 46 35 8 6 15 9 27

Remarks.-The species was founded on two young females, which do not appear to be fully developed and are apparently not the mothers of certain very young specimens that accompany them. The species is said to be related to Crocidura foetida and C. doriB'3, of Borneo, but it differs in having much larger feet than these species. From the known Philippine forms it may be readily distinguished by the black pelage. The types were collected in Jol0, Sulu Islands.

Order DERMQPTERA

The Dermoptera have long been regarded as a suborder under the Insectivora. More recent authors have recognized differences between the true insectivores and this strange aberrant form which seem to warrant the rank of an order for the single family and genus to which it belongs. Upper and lower incisors compressed, multicuspidate, the lower deeply pectinate, or comblike; fore and hind legs connected by a broad integumentary expansion forming a parachute; orbit nearly surrounded by bone; zygomatic arches well developed; tympanic forming bulla; ulna united with radius; tibia and fibula distinct; pubic symphysis long; penis pendent; testes received into inguinal pouches; mam:rnre axillary; uterus bicornuate; crecum large. Family GALEOPTERIDJE Characters the same as those of the order. This family name has been proposed * to replace GaZeopithecidB'3 (auct.) and Colugiclm Miller since the generic names Galeopithecus and Colugo are invalidated. The genera Galeopterus

* Thomas,

Ann. & Mag. Nat. Hist. VIII 1 (1908) 254,

PHILIPPINE LAND MAMMALS

Thomas and Cynocephalus Boddaert are recognized; only the latter occurs in the Philippines. Genus CYNOCEPHALUS Boddaert

Cynocephalus BODDAERT, Dierk. Meng. 2 (1768) (footnote); THOl';IAS, Ann. & Mag. Nat. Hist. VIII 1 (1908) 252-255; HOLLISTER, Philip. Journ .. Sci. 搂 D 7 (1912) 7. Galeopit hecus PALLAS, Acta. Acad. Sci. Imp. Petropolis IV 1 (1780) 208, pI. 7,8; CUVIER, Tableau Elem. (1788) 106; SHAW, Gen. ZooI. 1 Mamm. (1800) 115-121, pI. 38. Galeo'JYUS' RAFINESQUE, Analyse de la Nature (1815) 54. De?'nwpte?'us BURNETT, Quart. Journ. Sci. Lit. Art. 27 (1829) 268. Pleu?'opterus BURNETT, Quart. Journ. Sci. Lit. Art. 27 (1829) 268. Colugo GRAY, Cat. Monk. Lem. Fruit-eat. Bats Brit. Mus. (1870) 98.

The characters of the genus are those given for the order. Dental formula:

.2123 1.

3' c. T pm. 2' m. "3 = 34.

Only one species is recognized * from the Islands. Great variation does occur, especially in color and markings, and to a lesser extent in other characteristics, but this maximum variation may be evident in a group of specimens from a single locality and is due to age, sex, and individual variation and cannot be correlated with geographic distribution. In the Philippines the species occurs in southern Mindanao (apparently absent at least in certain parts of northwestern Mindanao), Basilan, Samar, Leyte, and Bohol. Elera gives Cebu as a locality, but if it was once present in that island, it appears to be extinct. CYNOCEPHALUS VOLANS (Linnreus)

Lemur volans LINN.tEUS,t Syst. Nat. 10 ed 1 (1758) 30. Cynocephalus volans BODDAERT, Dierkundid Mengehvert 2 (1768) 8, footnote 1. Galeopithecus volans PALLAS, Act. Acad. Petropolis 4 (1780) 208.

* Elera, Contr. a la Fauna Filipina, lists Galeopithecus rufus Geoff. (no locality); Galepithecus variegatus Geoff., Cebu, Leyte, Samar, Java, Borneo, Sumatra; Geleopithecus philippinensis Wat., no locality; G. volans Shaw, Bohol, Cebu, y toda la Malasia; G. marmoratus Tern., Basilan, Samar, Banka. t Pre-Linnrean names: Cato-Simius volans camellii PETIVER, Gaz. Nat. Artis. t. 9 (1702) f. 8, and Act. AngI. 277 n. 1065 [= Phil. Trans. Royal Soc. pt. 277 in Vol. 23 (1704) J. Vespe1路tilio admirabilis BONTIUS, Java (1658) 68. Felis volans ternatea SEBA, Mus. 1 (1735) 93, pI. 58, figs. 2 and 3.

CYNOCEPHALUS

Galeopithecus (Colugo) philippinensis GRAY, Cat. Monk. Fruit-eat. Bats Brit. Mus. (1870) 98. Galeopithemts philippinensis W ATERHOUSID, Proc. Zool. Soc. London (1838) 119-120; STEERE, List Birds Mammals Steere Exped. (1890) 28 (Basilan, Bohol, Mindanao); THOMAS, Trans. Zool. Soc. London 14 (1898) 386 (Bohol, Samar, Leyte); GUNTHER, Proc. Zool. Soc. London (1879) 74 (Dinagat); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 7; Proc. U. S. Nat. Mus. 46 (1913) 304. Colugo philippinensis MILLER, Proc. BioI. Soc. Washington 19 (1906) 41.

Description of the species.-(From No. 112, Bureau of Science collection; collected January 22, 1907, Basilan Island, by Andres Celestino.) Adult female. Head, except nose, covered with short fur, extremely short at base of nose and bordering lips, growing gradually longer on forehead and between ears; ears small, narroW, on posterior side furred to the summit; within, ear covered with sparse minute hairs; body and greater part of membranes covered with soft 'fine wavy fur; antebrachial membranes attach on inner digit of foot and on sides of neck behind and much below ear; lateral membranes most heavily furred near body, fur growing scantier and very much shorter along outer edge; lateral membranes attach at base of claws; interfemoral membranes strongly furred along posterior part of body and along limb, but growing nearly naked on posterior portion; above tail a line of fur extending to tip; throat, median part of belly, antebrachial membrane, and lateral membranes covered with somewhat scant fur; ,sides of belly, especially about mammre, almost entirely naked; interfemoral membrane attaches to claws of outer toes and almost entirely naked except a scant line of hair which follows tail; the latter entirely inclosed in the membrane; limbs elongate, subequal; digits of forelimb armed with strong claws which are nearly as broad at base as they are long; digits webbed, the membrane covered with a fine growth of hair; pads under toes and sole of foot naked; underside of femur scantily furred; digits of fore limbs longer and stronger than those of hind' limbs; mammre four, paired and axillary. Color.-Head generally grayish resulting from buff and gray hairs; two small buff spots on face between eyes followed by an area somewhat more brownish; a small white spot on back of head; color of body above generally vinaceous buff, tips of hair dark to buff grayish; basal portion of fur purplish brown;

PHILIPPINE LAND MAMMALS

fur on wing membranes darker than body, grayish or olivegrayish portion of hair much longer, many of the hairs tipped with buff; basal purplish brown portion of hairs short; outer portion of wing membrane deep brown; hind limbs and interfemoral membrane generally uniform grayish with a buff streak above tail; above forearm darker with a distinct deep black-brown area and buff-white spots; deep brown mixed with whitish spots above toes; undersurface generally buff or orange-buff except at base of interfemoral where the fur is deep chestnut-brown growing gradually lighter toward outer edge of membrane. Skull and teeth.-Skull low, rostrum arched, a depression between orbits; upper edges of orbits raised and strongly proj ecting; postorbital process extends 6 millimeters; process from zygomatic arch 4 millimeters; distance between tips of processes 9 millimeters; two low crests run back from postorbital pro-cesses across brain case almost coming together in occipital region and then separating at posterior edge of skull; zygomatic arches strong, base of anterior attachment greatly widened forming base of eye socket; palate slightly concave, incisive foramina large, their combined width equal to their length; posterior edge of palate with two rounded indentations separated by a septum which continues in a low crest nearly as far as base of zygoma. First pair of upper incisors small, widely separated (17 millimeters), second pair high and large, caniniform, larger than canines; shaft a flattened cone with lateral ears; this tooth and the following very similar in shape, the canine of greater anteroposterior length and thicker; first upper premolar about as wide as long, molariform; third premolar and upper molars similar in character, each much broader than long, complex, each with four or five cones arranged about a deep depression, inner sides of cones finely striated; first lower incisors strongly compressed, built like a comb for the hair, with ten- "teeth," each "tooth" flattened and not or scarcely in contact with its neighbor; second lower incisors not, or but barely, in contact with first, and distinctly fleparated from third, similar to first but considerably wider, !Somewhat shovel-shaped, with twelve "teeth" in the "comb;" third lower incisor somewhat molariform with four cones; lower canine 2-rooted, much higher than premolars; first lower premolar elongate with three cones; coronoid process present; condyle transversely elongated; posterior end of mandible rounded.

CYNOCEPHALUS

Measuxements of Cynocephalus volans (Linnmus). mm.

Length, head to tail, dry skin Head Tail Height of ear Length of forefoot to tip of longest digit Length of hind foot to tip of longest digit Length of claw Width of claw Skull, greatest length Zygomatic width Diameter of orbit Height of brain case Shortest interorbital width Width between tips of postorbital processes Width of brain case Width between canines Width between last molars Palate from incisive foramina Upper tooth row complete, alveolar Lower tooth row from anterior edge of canine Length of mandiBle bone

658 78 244

19 89 84 10 9 75

51 17 27.5

20.6

34 25.5 21 17 28 37 29 49

Variation.-The most obvious vruriations in the species are in color and markings, due perhaps entirely to age and sex. No. 110, Bureau of Science collection; adult female, Sevilla, Bohol. Face and forehead dark with two whitish buff spots, one behind nose, the other between eyes; a small buffy white spot between ears; fur on back generally dirty whitish with gray tips or bands of gray tipped with grayish white; wing membranes generally lavender brown as are the bases of the individual hairs on the back; outer wing membranes lavender brown of a more distinct shade, becoming buff near their attachment to hind limb; buff above tail; forelimb dark brownish with a very indefinite wash of buff in places and scattered buff spots on digits and interdigital membranes; below generally buff, somewhat lighter on the belly than on membranes. No. 107, Bureau of Science collection; adult female, Sevilla, Bohol. General coloration and markings very similar to No. 110, but a s~ronger ,suffusion of smoky gray on back, and no white spot between the ears. No. 74, Bureau of Science collection; young female, Guindulman, Bohol. Fur for greater part of length whitish, outer portion ringed with smoky to brownish gray; on shoulders and rump hairs tipped with whitish buff forming indefinite lighter areas; on wing membranes the main color is buff, nearly con-

PHILIPPINE LAND MAMMALS

cealed by the smoky to blackish gray tips, rarely olive gray; feet with a buff wash on outer portions; interfemoral membrane dark brown edged with lighter; fur fairly well developed to tip of tail, brownish hairs tipped slightly with dark buff. No. 75, Bureau of Science collection; adult male. Head almost blackish ~ith a single large white spot between eyes; another indefinite large white spot between ears; above entire body and wing membranes dark, variegated, lavender brown with occasional groups of light hairs which on the limbs form distinct small spots; on belly brownish buff; nearly brown on membranes. Nos. 742 and 743, E. H. Taylor collection, from Saub, Cotabato, resemble the form described from Basilan but there is considerable variation in details; the chestnut brown along the body is almost identical in three specimens; No. 742 has the fur less rough and irregular than No. 743, and in fact, the fur is almost straight and smooth, and is, at least on the body, more brownish than gray; on the rump and limbs are small tufts of elongate white hairs which are conspicuous. No. 741, E. H. Taylor collection, a young female, Saub, Cotabato, has the upper J?art of the body a deep buff brown, the tips of hairs edged with buff, and on the outer edges of body hairs edged with black; small elongate wisps of cream-colored hairs push out several millimeters beyond the fur along sides of body on proximal portions of limbs and on rump. No. 774, E. H. Taylor collection, Saub, Cotabato, is an adult male, strongly resembling the males listed from Bohol. Remarks.-The species occurs in Bohol, Samar, Leyte, Mindanao, and Basilan. . Capt. Francis Link~ Philippine Constabulary, reports (in letter) the presence of this or an allied species on the islands of the Tonkil group in the Sulu Archipelago, and its absence in Jolo as well as other islands near by. The animal feeds on leaves of trees. It is nocturnal in habit, beginning to move about in the forest at dusk. My specimens were obtained when a portion of forest near the seashore, at Saub, Cotabato, was being felled, the specimens being taken by hand by the natives. The common name * of the species is cfLguan, or gaguang throughout the Visayan islands where it occurs; the name in Samal Moro is mimmfLl.

* Elera lists the following native names; Gigua (Tagalog and Pampango!); cago, calago gagua, caa,b uit, (Visayan); calamby (Cagayan) Qllingua (?)

TUPAIIDJE

Order MENOTYPHLA

Upper molars broadly multicuspidate with more or less welldefined W -shaped crowns; symphysis pubis long; radius and ulna, and tibia and fibula separate; zygomatic arch invariably present; orbit either completely or incompletely surrounded by bone; brain case large; jugal perforated. The Tupaiidre and the Macroscelididre, belong to this order; only the former occurs in the Philippines. This group is usually considered a suborder under the Insectivora. The characters by which it differs from the typical insectivores seem to warrant its separation from that group; its relation to the Lemuroidea warrants placing it near the Primates. K ey to Philippine fami lies and genera of the Menotyphla. a '. Postorbital process and crecum present; orbital ring encircled by bone; metatarsus moderate; symphYl'\is pubis long. Squirrel-like, arboreal forms ........................................................................ TUPAIIDJE (p. 97). b '. Tail flattened, with long hairs on top and sides; second incisors and lower canines moderate .................................... Tu paia Raffles (p. 99). b 2. Tail cylindrical, with short hairs ; second incisors and lower canines very large .............., ......................................... Ur og!!le Mea rns (p. 106).

Family TUP AIIDJE Arboreal Insectivora, with comparatively large brain case; orbit encircled with bone, and a well-developed zygomatic arch; malar bone perforated; tympanics form bullre; pubic symphysis long; tibia and fibula distinct, metatarsus little longer than tarsus; molars broad, with W -sha ped cusps; intestinal canal with a short crecum. In general members of this group resemble squirrels. Aside from the remarkable difference of the teeth, they differ in having the bushy hajr of the tail absent on the underside and to some extent above. The muzzle is longer and the ears are shorter, and more flattened against the head. In the Philippines the group is confined to Palawan and Mindanao. In northern Palawan specimens of Tupaia palawanensis Thomas are fairly common. They move about in the forest climbing trees and vines, and running over the ground with great rapidity. While I was trapping squirrels on Palawan a specimen of the above species was unexpectedly captured in a trap placed at the base of a tree, baited with nuts, oatmeal, and insects mixed in a paste. This happened only a few minutes after the trap had 269458-7

PHILIPPINE LAND MAMMALS

been set. The trap was again set and when visited contained what I supposed was another specimen of the same species. Upon examination, however, it proved to be a specimen of Sciurus. The general similarity in the two forms is striking. In the Philippines so far as is known, Tupaia is confined to the Palawan group of islands; Urogale is confined to Mindanao. Besides the two genera listed, there are elsewhere three other genera commonly recognized. These are Dendrogale, Tana, and Anathana. The five genera belong to the subfamily Tupaiime. The other subfamily, Ptilocercinre, has only a single genus" Ptilocercus, and is not represented in the Philippines. Distribution of the genera of TupaiidlB.

Lyon * points out that this family does not cross Wallace's Line. He states: The Tupaiidre as a whole range from India on the west to and including Mindanao of the Philippine Islands on the east, and from southern China on the north southward to and including Java and the chain of islands off the southwest coast of Sumatra. They are not found eastward of Java, nor on the 'Celebes, Formosa, Ceylon, or the Andaman Islands so far as known. I know of no specimens or records of the Tupaiidre on the island of Bali, off the east end of Java and just west of Wallace's Line. It would not be surprising to find them on Bali when the fauna of that island becomes better known. They are found on practically all the smaller islands of the Malayan Archipelago, within the limits just mentioned, and more frequently than not develop geographic races or species on them.

Another significant fact seems to be that while Ptilocercus, Dendrogale, and Tana are present in Borneo and Sumatra (in the case of Dendrogale in Indo China), they appear to be

* Proc.

U. S. Nat. Mus. 45 (1913) 15.

TUPAIA

totally wanting in Java. It suggests that Java was separated prior to the separation between Borneo and Sumatra or Borneo and the Asiatic mainland. Genus TUP AlA Raffies

Sorex DIARD, Asiat. Joum. Month. Register 10 (1820) 478 (non Linnmus 1758). Tupaia * RAFFLES, Trans. Linn. Soc. London 1 3 (1822) 256 (published in May, 1821); LYON, Proc. U. S. Nat. Mus. 45 (1913) 30. Sorex-glis GEOFFROY and CUVIER, Rist. Nat. Mamm. 3 (1821) 1 (published in December, 1821). Glisorex DESMAREST, Mammalogie (1822) 536. Cladobates CUYlER, Dents des Mammiferes (1825) 60, pI. 17. Hylogale TEMMINCK, Monogr. Mamm. (1827) 19. Erinaceus BLAINVILLE, Osteo-g. Mamm. Insect. (1840) 112, pI. 6, fig. 1. Hylogalea SCHLEGEL and MULLER, Verh. Nat. Gesch. N ederl. Overz. Bezitt. (1843) 159. Gliso8orex GIEBEL, Odontographie (1855) 18. Sciuru8 ELLIS, in Gray, Ann. & Mag. Nat. Rist. III 5 (1860) 7l. Tapaia GRAY, Ann. & Mag. Nat. RistJ III 5 (1860) 71. (Error?) Glirisorex SCUDDER, Nomenclator Zool. pt. 2 (1882) 13l. Glipora JENTINK, Cat. Syst. Mus. Rist. Nat. Pay-Bas. Mamm. 12 (1888) 118.

The species of the genus are squirrel-like tree shrews with elongated snouts. They are distinguished from other members of the family by having the ears very small; the finely reticulated naked area of the nose, not encroaching on haired area of nose; the tail well haired; the face lacking markings; a more or less distinctly defined oblique shoulder stripe; mammre in one to three pairs (two in Philippine forms). Rostrum moderately .short and heavy; distance from lachrymal notch to end of premaxilla about equal to distance from notch to the region of external auditory meatus. The teeth are more generalized than in other members of the family; the two pairs of upper incisors are usually subequaI, although the first is nearly always a little the larger; there are three pairs of lower incisors all well developed and functional, but the third pair is much smaller than the others; the second pair somewhat larger than the first pair; there are three pairs each of upper and lower premolars; the first is small and almost functionless; the others increase in size and complexity from before backward.

* From tupai,

a Malay word for these arboreal shrews and squirrels.

100

PHILIPPINE LAND MAMMALS

The type of the genus is Tupaia jeT1'u,ginea Raffles. The genus is distributed in India, Burma, Siam, China, Malay Archipelago to Java and north to the Philippines, extending only to the Palawan group. No specimens have been collected in 118 "

120'

122' Batan Is

126

PHILIPPINE ISLANDS

124

® Tupala palawanensis @ Tupaia cuyonls

CO Tupaia mollendorffi Urogale everetti ® Podosymnura truei ® Cynocephalus volans @

'" ""'.

12'

..... ~ 10

'"' ~

<>Casayan Sulu

". ®,....8asilan ,-r®,

~®®

~Jolo d'~

••

,4>

~Tawitawi Sibutu :D ~ ungau

120

FTG. 3,

122

124

126

Distribution of Cyn,cephalus, Tupa;a, and Erinaceidre in the Philippines.

other parts of the Philippines. * Some forty-four species and subspecies are recorded in 'Lyon's monograph.

* A British Museum specimen listed by Lyon (op. cit. 80) has the locality "Palawan or Basilan." The former island is doubtless the proper locality. '

101路

TUPAIA

The three species described from the Palawan group, Tupaia pala'Wanensis Thomas, T. cuyonis Miller and T. mollendortfi Matschie, constitute a small group having a generally grayish or ochraceous color without characteristic marks or bright colors, and with the mammary formula 2 + 2 = 4 . Dental formula:

. 2 I.

3' c. l' pm. 3' m. 3 = 38.

Key to the Phili1Jpine species of Tupaia Raffles. a 1. Tail distinctly different in color from lower back; teeth larger. b '. Tail more grayish than back ........ T. mollend01'ffi Matschie (p. 105). b". Tail more blackish than back ....... T. palawanensis Thomas (p. 103). a ' . Tail approximately the same color as lower back; teeth smaller. T. cnyonis Miller (p. 101). TUP AlA CUYONIS Miller

Twpaia cuyonis MILLER, Proc . .U. S. Nat. Mus. 38 (1910) 393 (August 19); LYON, Proc. U. S. Nat. Mus. 45 (1913) 82, pI. 9, fig. 1; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 82; Proc. U. S. Nat. Mus. 46 (1913) 304.

Description of the species.-(From No. 24, adult male, Bureau of Science collection; collected January 14, 1903. Cuyo Island, P. 1., by R. C. McGregor and A. Celestino. Formerly No. 239183, United States National Museum, one of the paratypes.) General habitus that of a small squirrel; snout pointed, tip bare, finely reticulated with a distinct groove on anterior part; nasal openings in the form of narrow curves; a well-defined groove separates the naked portion of the snout from the haired portion save in the median part; a .s mall naked area at tip of lower jaw; vibrissre on snout short, laterally placed, extending on upper lip to corner of mouth; a tuft of two or three elongate vibrissalike hairs above anterior part of eye; a group of similar hairs on side of head anterior to the lower edge of ear; a few elongate hairs, usually, on throat; all elongate hairs black; body covered with moderately long hair, softer and sparser on posterior part of belly; tail, dorsally and laterally, with a heavy growth of long hair (lateral hair, 30 millimeters; dorsal, 20); ventrally, hair short (abou't 5 millimeters) ; hair on tip of tail 50 millimeters: an ill-defined group of stiffened elongate hairs on outer anterior part of forearm; ears small, pressed fiat against head, naked save for scattered fine short hair; legs well developed; mammre (in females) 2 2 = 4. Color.-Upper part of body and legs, sides of head, neck, and body a grizzle of black and ochraceous, basal portion of

102

PHILIPPINE LAND MAMMALS

the hairs for nearly half their length slate gray, distal half with bands of black and various shades of buff, the latter predominating; an ill-defined diagonal stripe showing an excess of ochraceo us buff on shoulder; underside of forelegs, throat, and chin lighter, belly not especially differing from sides save for a russet tinge in certain lights; tail similar to body, darker above than below. .. Skull.-Skull elongate, tapering anteriorly; orbit entirely inclosed; zygomatic arches strong; brain case relatively large; individual bones of skull largely grown together; a distinct longitudinal sagittal crest between interparietals and a transverse crest at juncture of supraoccipital with interparietals and parietals, this crest the most posterior part of skull; auditory bullre well developed. Measu.rements of Tu.pr;tia cu.yonis Miller. mm.

Total length, field measurement to tip of tail vertebrre Head and body{ field measurement Tail length, field measurement Hind foot, field 'measurement Forefoot, dry Head, about Condylobasal length Zygomatic width VVidth of brain case Interorbital width Diameter of orbit Orbit to tip of premaxillary Length, upper tooth row Length of maxillary tooth row Length of three molars Length of palate to level of first incisors Width between third upper molars Width between upper canines Length of mandible to incisor tips Last lower molar to tips of incisors Length of three lower molars Width between third lower molars VVidth between lower canines

365 â&#x20AC;˘ 191 174

42 24 53 43.8

25 18.5 14

10 20 24 16 7.1

24 9.0 6.0 34.5

22 8!5 8.0 4.0

Remarks.-The teeth of the described specimen are well worn. The type specimen is apparently smaller than the present one Head and body measurement stated by Lyon (op. cit. p. 83) to be 145 millimeters. This appears to be an error. The dry skin measures, head and body, 179 millimeters and the field tag gives 191 millimeters. ,The dry tail measures 178 millimeters, the tag gives 174 millimeters. U

TUPAIA

103

since the measurements are only 154 and 166 millimeters, respectively, for head and body and for tail. The skull measurements of the type are all slightly smaller than those given here. Miller (loc. cit.) states that save in coloration this species is similar to Tupaia mollendorjJi Matschie. TUP AlA P ALA WANENSIS (Thomas)

Tupaia /errltginea palawanewis THOMAS, Ann. & Mag. Nat. Hist. VI 13 (1894) 367; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 6. Tupaia palawanensis LYON, Proc. U. S. Nat. Mus 46 (1913) 304; HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 304.

Description of the species.-(From No. 20, E. H. Taylor collection; collected at Sir J. Brooke Point, southern Palawan, September 20, 1923, by E. H. Taylor.) Adult female.-Similar to Tupwia cuyonis Miller but differing in the darker pelage of body and tail; vibrisS'::E on snout and head with the same disposition as in T. cuyonis .. a few elongated hairs on anterior part of forearm; hair on tail distinctly thinner and shorter; mammre 2 + 2 = 4. Color.-Head covered with nearly uniformly black and ochraceo us buff hair, black predominating; no darker area along median line as in the type, and only a faint suggestion of a darker area above eye; fine short hair on ear black; immediately behind ear ochraceous predominates; a curved, fairly distinct, buff-colored line on shoulder; body and sides almost uniformly grizzled, the darker shade predominating on the back, the ochraceous on the sides; sides of belly and underside of forelegs ochraceous-rufous, the median area with tints of slatish buff. Skull and teeth.-Teeth, of molar series in particular, large, having distinctly greater width and length than those of Tupaia cuyonis; skull appears less heavy, bone on posterior part of orbit more fragile than in T. cuyonis.. lachrymal foramen larger and more pronounced; coronoid process of mandible rises more perpendicularly; auditory bullre slightly larger than those of Tupaia cuyonis. Variation.-Of the specimens listed above, Nos. 19 and 20 are from Sir J. Brooke Point on the eastern coast of Palawan near the southern end of the island, and No. 560* was taken in the extreme northern part of the island, on the coast near Shark * The following additional measurements of No. 560 were made on the freshly killed specimen. Tip of snout to eye, 27 millimet"ers; snout to ear, 43; height of ear, 16; width, 12; forefoot with claw, 25; hair at tip of tail, 31. .

104

PHILIPPINE LAND MAMMALS

Fin Bay. The latter specimen, while agreeing with the southern specimens in skull characters and dentition, exhibits certain color variations. On the head, from snout to forehead, the black color of the grizzle is so prominent as to form an inde... finite darker median line; above the eye is a suggestion of two lighter areas separated by a darker; the hair on the ears is more brown than black, likewise the back of the hand and foot; the back and tail do not differ markedly in coloration save that the back shows more rufous. The tail above and below is lighter than the southern form. The underside of the tail shows three fairly well-defined ochraceous-buff lines on either side of the median part. It is not improbable that the northern specimens approach Tupaia mollendorfji of the Calamianes. It may be that the northern form will warrant subspecific designation when more specimens are available. M easu'rem ents of specimens of Tupaia palawanensis (Thomas). E. H. T. collection No. 19

20 , female.

----------------~~~--~~-~------------I------

mm. H ead and body_____ _ _ _ __ __ _ __ _ _ _ _ _ _ _ _ __ _ ___ _ _ _ __ _ _ _ _ __ _ _ TaiL __ ____ _ ____ _ ___ _ ____ ____ __ ____ _________ _ _____ __ __ __

362 184 178

Hind foot ___ â&#x20AC;˘â&#x20AC;˘ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ __ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _

Skull, condylobasallength ________ ______ _____ ___ __ _____ ___ Zygomatic width _ __ _ _ _ _ _ _ _ _ __ _ ___ _ _ _ _ __ ___ _ _ _ __ _ ___ _ _ __ _

46 25

Width of brain case_ ______ __ ____ _______ ________ _________ _ L ea st interorbital width_ __ _ _ _ _ _ __ _ _ _ _ ____ _ __ _ _ _ _ _ _ ___ _ __ _ Diameter of orbiL __ ____ _ __ __ _ _ _ _ _ _____ __ ___ _ ______ _ _ ____

18 13 10 . 5

Orbit to tip of premaxillary__ _________________ ____ _______ _ L ength oj upper tooth row, inner measurement_ __ _ _ _ _ __ _ _ _ _ _ Length of three upper molars _ _ __ _ _ ____ __ __ __ _ ___ __ _ _ __ ___

22.2 26 . 1 9.1

L engt h of maxillary tooth row_ __ ___ ____ ______ __________ __ L ength of palate to anterior level of first incisors_ __ _ _ _ _ __ __ _ Width between third upper molars_ __ ____ __ ______ __ ______ __ Width betwe en upper canines_____ _ _____ __ __________ ____ __ Length of mandible to incisor tip____ ____ __________ ___ __ ___ Last lower molar to tips of incisors ________ _____ ___________

18 27 9 5.1 35 . 1 24 . 1

L ength of three lower molars_______ ___ ___ ______ _________ __

mm. 365 195 170 44 46 26 18 14 10 . 5 21.2 25 . 6 9.1 17 . 5 26

9 6

35 23

9. 8

560

----711m.

344 167 177 45 47 25 5 20 . 8 14 . 1 10 . 5 22 . 5 26 9 18 . 4 26 9 . <l

5.5

37 2~

9.6

Remarks.-The typical Tupaia palawanensis is described as having a black tail. My specimens and apparently most of those studied by Lyon (op. cit. 78, 79) do not show this condition. Lyon examined eighteen sItecimens, nine of which were from near Puerto Princesa, near the eastern, middle part of the island. One was from Balabac Island and another from the Calamianes.

105

TUPAIA

The last specimen (in Berlin Museum), if correctly labeled, would seem to give us two forms occurring in the Calamianes, one of whose islands, Culion, is the type locality of Tupaia mollendorffi. The apparently greater length of the tail in the specimen from northern Palawan may be an added character since it actually (in a freshly killed specimen) exceeds the length of head and body. In none of the .specimens of T. mollendorffi or T. palawanensis examined by Lyon, did the tail reach such a length. However, the tail is longer than head and body in certain T. cuyonis (presumably field measurements). The type locality is "Pala~Tan." The type was collected by A. Everett, and is No. 94.2.1.3 in the British Museum. TUPAIA

M~LLENDORFFI

Matschie

T[upaja] mollendorffi MATSCHIE, Sitz.-Ber. Ges. Nat. Freunde, Berlin (1898) 39. Tupaia mOUendorffi HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 6; Proc. U. S. Nat. Mus. 46 (1913 ) 304; LYON, Proc. U. S. Nat. Mus. 45 (1913) 81.

Description of the species.-(After Lyon.) Upper parts finely grizzled, ochraceous and blackish; the tail coarsely black and buff, 2 = 4. more gray in color than upper parts; mammre 2 Color.-Upper parts and sides of head, neck, and body a fine grizzle of ochraceous to ochraceous-buff and blackish, the lighter color being in excess, especially along sides and rump; underparts, including throat and inner side of legs, usually dirty buff or ochraceous-buff; in the type the chin and throat cream-buff, distinctly lighter than rest of underparts; outer side of legs essentially like upper parts of body; feet similar but the grizzle very fine and the dull ochraceous-buff predominating; shoulder stripe fairly well defined, buffy; tail a coarse grizzle of buff and blackish both above and below, the blackish color slightly in excess above, and the buff below; tail from above distinctly grayer than rest of upper parts, and noticeably different in color. Skull and teeth.-The skull is somewhat smaller than in the Palawan species; the teeth show little variation.

Measurements of the type of T'ttpaia mollendorffi Matschie. mm.

Head and body, dry skin Tail Hind foot Zygomatic width VVidth of brain case Maxillary tooth "row

200 160 43 24 18 16.5

106

PHILIPPINE LAND MAMMALS

Remarks.-The type specimen was collected by von Mollendorff on Culion Island, Calamian group. It is No. 9858, in the Royal ZoOlogical Museum, Berlin. Genus UROGALE Mearns

Urog'ale MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 435; HOLLISTER, Phiiip. Journ. Sci. ยง D 7 (1912) 6; LYON, Proc. U. S. Nat. Mus. 45 (1913) 154, text figs. 12 and 13.

Large; feet and claws relatively large; tail short-haired and cylindrical. Brain case narrow and elongate; zygomatic vacuities obsolete; rostrum produced, enlarged and greatly swollen for the accommodation of enormous second incisors; second upper incisor caniniform; third lower incisor minute and erect; lower canines enormous. (From Mearns.) The type of the genus is Urogale cylindrura Mearns which is regarded as synonymous with Tupaia everetti Thomas. It is not improbable that Mearns's species represents a color variety, but Lyon is convinced that the species are identical. UROGALE EVERE1lTI (Thomas)

Tupaia everetti THOMAS, Ann. & Mag. Nat. Hist. VI 9 (1892) 250. Urogale cylind!rura MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 435; HOLLISTER, Philip. Journ. Sci. ,ยง D 7 (1912) 7.

Urogale everetti HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 7; Proc. U. S. Nat. Mus. 46 (1913) 304; LYON, Proc. U. S. Nat. Mus. 45 (1913) 157, text figs. 12 and 13; pI. 11, fig. 6.

Description of the species.-General characters the same as those of the genus. Color.-General effects of upper parts of head, neck, body, tail, and outer side of legs a dark mummy to seal brown, produced by a fine grizzling of blackish and tawny, the blackish brown in excess; about nose and sides of head, the tawny replaced by raw sienna and more of it; feet similar to legs but darker, and with almost none of the lighter color; underparts including inner side of legs, varying from ochraceous to ochraceous-rufous, brightest in the region of the chest, the slaty bases of the hairs showing through in places to a considerable extent; underside of tail similar to upper side but the light and dark color elements about equally mixed; shoulder stripe rather poorly defined ochraceous. Skull and teeth.-Sknlllarge, angular, with a relatively heavy rostrum rising abruptly from rest of skull, enlarged just back of extremity to accommodate roots of the large second pair of in-

107

CHmOPTERA

cisors; temporal ridges short but prominent; sagittal crest rather long and high; bony palate well ossified usually with vacuities; postorbital constriction less than preorbital constriction; second pair of upper incisors enormousily developed functioning as canines; third lower incisors small functionless and sometimes deciduous; last upper and lower premolars wen developed especially the protoCiones on upper tooth; lower canines conspicuously developed. (After Lyon.) Measurements of Urogale eve7'etti (Thomas)" mm .

Head and body, dried skin (approximately) Tail Hind foot Skull, condylobasal length Zygomatic width Width of braincase Maxillary tooth row a

23 3 170

51 58 29 21

Largest specimen listed by Lyon, Proc. U. S. Nat. Mus. 45 (1913) 168.

Remarks.-The type of Urogale everetti was collected at Zamboanga, Mindanao. It is No. 79.5.3.11 in the British ,Museum. Mearns's specimens, including the type of UTogale cylindrura, were collected on Mount Apo. Lyon mentions a young shrew collected by Mearns on Mount Malindang. The skull is missing, but it appears to represent a different species, which cannot be determined without the skull and the adult pelage. Order CHIROPTERA Chiroptera

BLUMENBACH,

Handbuch der Naturgeschichte (1779) 74.

Mammals with the front limbs modified for flight; digits of front limbs greatly elongated (the length of the third usually exceeding or equalling the length of head and body), joined together by a naked membrane which attaches to the side of body (sometimes near middle of back, Hypodermis) and along side of hind limb. Shoulder girdle more strongly developed than pelvis; sternum is usually keeled; knee directed backwards owing to rotation of leg to support wing membrane. Body covered with fur above and below, rarely approaching nudity (Cheiromeles) . Colors usually nearly uniform. Bats are found in all parts of the world to the northern limits of tree growth. The Galapagos Islands and the Hawaiian Islands have received their bats from the Americas, while New Zealand, Samoa, and the Carolines have received theirs from Asia.

108

PHILIPPINE LAND MAMMALS

Key to the suborders of Chiroptera,. a.' Second finger maintaining an evident degree of independence, its ungual phalanx present; humerus with trochiter and trochin small, the former never articulating with the scapula; mandible with angular process broad and low and practically absent; margin of ear forming a complete ring ............................................................ MEGACHIROPTERA (p. 113). a,! Second finger scarcely if at all independent from third, its ungual phalanx" absent; humerus with trochiter and trochin large, the former usually articUlating with scapula; mandible with angular process well developed, long and narrow; margin of ear not forming a complete ring ................................................ MICROCHIROPTERA (p. 194). Distribution of the fa,milies of Chi1·optem. -----

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- --" Pteropida> __ . __ . _______ __ _____ __________ __ _ ~ __ . x x x x x x ______ __ Rhinopomidal _______ __ ________ ___________ __ ~_ __ x x __ __ x __ __ X ______ __ ---Emballonuridal _____ ____________ __ __ ____ ______ __ ____ X X X X X ____ X X Noctilionidal __ ______ _______ • ~ _ ___ __ ____ ____ ____ __ __ __ __ __ __ __ __ __ __ __ __ ____ x X Nycteridal ____ _________ _____ , _________ ____ __ x ____ __ __ __ __ x ___ ___ __ __ __

---

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______

---- x --- - ---- -- -00--

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-- -- -- --

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~i~~o;~<~ - : --: ~c :- :- ::-}-~-: -}-~I~:-[~ :~:~-~I ---:~: -~~. vespertilionida? __ ____ ______ _____ __ ______ ____ X Mystacopida? __ __ __ __ __ __ __ _ __ __ __ __ __ __ __ _ x Molossid:l' _____________ __ ______ __ _____ _________

X X x x x x X x X __________ __ !____ __ __ -- --:- -- - -- -- -- -X , x x I x x ix x x x

---

Key to the higher g?'oups of Philippine Chiroptera. a.' Second finger more or less free from third, the ungual phalanx present; margin of ear forming a complete ring; no nose leaves; tragus never present; mandibular incisors never more than 2-2. Suborder MEGACHlROPTERA (p. 113). b.' Characters of suborder ................................ Family PTEROPIDJE (p. 116). c.' Premaxillaries separate or barely united anteriorly. do' Tongue not highly extensible; molars and incisors not specially reduced in size; brain case never strongly deflected; upper surface of mandibular symphysis forming conspicuous angle with alveolar line .... Subfamily PTEROPINJE (p. 134). d." Tongue highly extensible; molars and incisors considerably reduced in size; brain case strongly deflected; upper surface of mandibular symphysis parallel with alveolar line. Subfamily MACROGLOSSINJE (p. 119). 0

...........................

CHIROPTERA

c." Premaxillaries broadly united;

109

molars with five sharp cusps each .................................... Subfamily HARPYIONYCTERINlE (p. 116). ct.' Second finger scarcely if at all independent from third, its ungual phalanx absent; margin of the ear not forming a complete ring. Suborder MICROCHIROPTERA (p. 194). b'. Premaxillaries absent ............................ Family MEGADERMIDlE (p. 206). b." Premaxillaries present. co' Premaxillaries usually free, always incomplete, their boundaries never obliterated; no nose leaf. do' Premaxillaries represented by a nasal branch only, or with a very incomplete palatal branch, second finger without phalanges; postorbital processes present. Family EMBALLONURIDlE (p. 194). d. 2 Same characters ................ Subfamily EMBALLONURINlE (p. 195). d.' Premaxillaries represented by palatal branch only; nose leaves present; no tragus. e.' Toes with two phalanges each; pelvis with postacetabular foramen .................................... Family HIPPOSIDERIDlE (p. 229). e." Toes except hallux with three phalanges each; pelvis without postacetabular foramen; premaxillaries partly cartilaginous, free from each other and from maxillaries. Family RHINOLOPHIDlE (p. 210). c.' Premaxillaries always fused with surrounding parts, complete or incomplete, their boundaries very early obliterated. d.] Fibula very robust, its diameter usually about half that of tibia contributing largely to strength of short stout leg; ears connected with band; tragus very small. Family MOLOSSIDlE (p. 309). d.' Fibula very slender or rudimentary not contributing essentially to strength of long, slender leg; tragus large, no nose leaf; ears not connected with a band .... Family VESPERTILIONIDlE (p. 249). e.' Anterior upper premolar large not differing conspicuously from posterior premolar in size or form; nostrils tubularly elongate ............................................ Subfamily MURININlE (p. 251). e.' Anterior upper premolars greatly reduced in size or absent; when present, conspicuously different from posterior premolar in both size and form; nostrils not tubularly elongated. t.' Sternum short and broad, its length in middle line not twice as great as breadth of presternum; ear slightly but evidently funnel-formed ........ Subfamily KERIVOULINlE (p. 253). t.' Sternum slender; ear not funnel-formed. g.' Presternum with median lobe larger than body of bone; scapula with coracoid straight directed inward; second phalanx of third finger nearly three times as long as first .............................. Subfamily MINIOPTERINlE (p. 264). g.' Presternum with median lobe much smaller than body of bone; scapula with coracoid curved outward; second phalanx of third finger not specially elongated. Subfamily VESPERTILIONINlE (p. 277).

110

PHILIPPINE LAND MAMMALS

Distribution of the families arid the subfamilies of Oriental bats.

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Pteropid", __________________ __ X X X X X X X X X X X X Pteropin", __________ ______ X - - - - X X X X X X - - -- X X X X MacrogloBsin", ________ __ __ X - - -- X X X X -- -- X - - -- - - -- X X Nyctimenin", ______ _______ -- -- -- -- X X X X - --- - - -- - --- - - -- - - -- - - -- - Harpyionycterin"' ___ ______________________________________ X _______________ _

Rhinopomid"' ___________________________________________________________________ _ Emballonurid"' _______________ X ________ X --- - ____________________ X X Emballonurin"' ___________ X ________ X ________________ X ____ X X Nycteridre _____ __ ________________________________________________ __ ______ X X Megadermidre ________________________ X -- -- - -- - - -- - - -- X - - -- - - -- X - - -- - --Rhino]ophidre _______________________ _ X X X - - -- - - -- X X X X X

Hipposiderid:E___ __ _ ___ __ ____ _ X X --- - - - -- - -' - - -- - --- - --- - - -Vespertilionidre__ __ ___ ___ __ _ __ X X X X X X X X X X Vespertilioninre ______ - ---- X X X X X X X X X X Miniopterinre _____________________________________________________

X X X X

MurininaL _______________ ---- --- - ---- ---- -- -- ---- ---- ---- ---- ---- X Kerivoulinm __________ _____ • ___________ X X X i X X ____ ---- X

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Pteropidm___ __ __ __ __ __ __ __ ___ X Pteropin m___ ____ ___ _ __ __ _ X MacrogloBSin:E ____________ X

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Hipposiderid"' ________________ X X Xx xX Xx XX Xx Xx ---- ---VespertilionidllL ______________ X X X X X Vespertilioninre _____ ______ X X X X X X X X X X X Miniopterinre _____________ X X X X X X X ---- ---- ---Murininm ______ __ ________ X X X X X ________ - -- - X ---- X Kerivoulinm _________ .. ____ X X X X X X ---- ---- ---- ---- ---NyctophilinaL ______________1._____________________________________ ___ __

---- --X-X X X ---- ------- ----

X X X Nyctimeninm __ ___ ____________ ---Harpyionycterin:P _____ ________ ---Rhinopomid lll ___ _____________ X X EmballonuridllL ______________ X X Emballonurinm ___________ X X

X X X X X X X X X X X X X ---X ---- ---- ---- ---- ------- ---- ---- ---- ---X X X X -- -X X X X X X X X X X

-- --

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I ___

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-- --

-- - - ---

---- ---X X

CHIROPTERA

111

Key to the genera of Oriental bats. a 1. Megachiroptera. Margin of ear making a complete ring. b 1, Pteropidre. Tragus never present; no nose leaf. e '. Pteropinre. Tongue slightly extensile: premaxillaries separate or barely united; molars not reduced. d ' . Teeth 30. e 1. Small bats with short tails, terminal half free from interfemoral membrane; claw on second finger . Cyn opter us Cuvier (p. 181) . e '. Tail shorter, concealed in fur; cheek teeth greatly enlarged. Thoopt erus Matschie (p. 188). W2. Teeth 28. e'. Small bats; outer upper incisors shorter than middle ones; fou r upper and two lower incisors. Ptenoehirus Peters (p. 178). e '. Medium-sized bats; no claw on second finger; two upper and two lower incisors ___ __.__ .. __ ___ ___ __________ Do路bsonia Palmer (p. 176). dJ路. Teeth 34. e'. Medium-sized bats; muzzle long, second finger with claw; tail 16 millimeters _______ ____ __________ , ________ __ ___ _ Rousettu s Gray (p. 171). e '. Large fruit bats; t.ail absent; interfemoral membrane narrow; claw on second finger _______ ___________ __ Pteropus Brisson (p. 147). e". Large bats; like Pteropus but with smaller canines and larger molars; middle incisors smaller than outer. Acerodon Jourdan (p. 135). e'. Macroglossinre. Tongue strongly extensible; molars and incisors reduced in size. d'. Teeth 34; small reddish brown fruit bats; body entirely furred; claw on second finger; no external tail; five upper molar-premolars teeth ___ ____ ____ _____ ____ ___ _______ ___ ___ Maeroglossus Cuvier (p. 119). d '. Teeth 34; similar to Maeroglossus but occipital portion of skull slightly de,tlected, the alveolar line when projected backward passing through the root of zygoma. Eonycteris Dobson (p. 126). e 3. Harpyionycterinre. Premaxillaries broadly united; to-ngue moderate; teeth 30; medium size, claw present on second finger; very short legs ___________ ____ __ _______ __ ________ Harpyionycteris Thomas (p. 117). a'. Microchiroptera. Margin of ear not making complete ring. b 1. Fmballonuridre. Premaxillaries represented by nasal branches, no nose leaf; tragus present; eyes large; tail perforates the interfemoral membrane and appears above, distinctly back from the edge. e" Teeth 34; no depression between eyes. Emballonura Temminck (p. 195). e '. Teeth 30; deep hollow between eyes; last 10 millimeters of tail free. Taphozous Geoffroy (p. 200). e". Similar to Taphozous but there is a total lack of emargination on the inner side of the audital bullre, and the upper molars are larger ____ ___ ____ _________ _: ___._____ ___ ___ ___ __ . ___ __ Taphonyeteris Dobson (p. 204).

112

PHILIPPINE LAND MAMMALS

b', Megadermidre. Premaxillaries absent; nose leaf present; small bats with long erect nose leaf, very short tails and large ears with bifid tragus; no upper incisor; teeth 28. Megaderma Geoffroy (p, 207), b 3. Rhinolophidre. Premaxillaries present; nose leaf present; toes with three phalanges except hallux. c '. Teeth 32; small bats with conspicuous nose leaves; ears large without tragus; tail well developed contained entirely within the interfemoral membrane........ Rhinolophus Lacepede (p. 211). c'. Teeth 30; small bats with large funnel-shaped ears; nose leaf small, inconspicuous except two lappets projecting over upper lip; tail short ...................................................... Chilophylla Miller (p. 247). b'. Molossidre. Premaxillary present, fused with surrounding parts; band connecting ears; fibula robust. c '. Teeth 28 to 30; bats with ears connected by a low band across head. d ' . Body heavily furred; tragus very small; tail long, free from membrane for half its length ...... Cha3rephon Dobson (p. 310). d '. Similar to Chawephon but with only 28 teeth. Philippinopterus gen. nov. (p. 314). c~. Teeth 26; earS not connected; body heavy and nearly naked; tail short; a neck gland present .......... Cheiromeles Horsfield (p. 320), b 5. Vespertilionidre. Premaxillary present; fibula slender; ears not connected. c '. Vespertilioninre. Slender sternum; ear not funnel-shaped. d 1. Teeth 38, tail long, ear well developed with slender tragus; interfemoral membrane large, its surface furred at extreme base below .......................................................... Myotis Kaup (p. 277). d 2. Teeth 34. e \ Similar to Myotis; ear shorter and broader, tragus less acutely pointed ...................................... Pipistrellus Kaup (p. 294). e 2. Similar to Pipistrellus but callosities on ball of foot and thumb more developed; upper incisor forced outward. Glischropus Dobson (p. 292). e 3 . Similar to Pipistrellus; fifth finger much shortened, scarcely longer than metacarpal of third and fourth; callosities on foot and thumb not well developed. Pterygistes Kaup (p. 301). d 3. Teeth 32; small bats with flattened head; ears normal with small tragus ................................ Tylonycteris Peters (p. 303). d ' . Teeth 30, ears small with long sharply pointed tragus. Scotophilus Leach (p. 305). c '. Kerivoulinre. Sternum short and broad; teeth 38; small, delicately formed bats; ears extending beyond nostrils when brought forward, slightly funnel-shaped, with long slender tragi. Kerivoula Gray (p. 253). c 3. Murininre. Nostrils tubularly elongate; ears nearly circular; tragus long, tapering to a fine point, wing membrane attached along

MEGACHIROPTERA

0'.

113

the whole length of outer toe to base of claw; tip of tail free; teeth 34 __ ______ ________ ___ _____ ___ ______ _______ ___ ___ ___ ___ ________ Murina Gray (p. 251). Miniopterinre. Ears small; tail as long as head and body, wholly contained within the interfemoral membrane; tragus high, slender, curved at tip; teeth 36 ________ Miniopterus Bonaparte (p. 264).

Suborder MEGACHIROPTERA F1-uctiVO?'/B GRAY, London Medical Repository 15 (1821) 299. Frugivora GILL, Arrangement of Families of Mammals (1872) 18. Megachiroptera DOBSON, Ann. & Mag. Nat. Rist. IV 16 (1875) 346; Cat. Chirop. Brit. Mus. (1878) 2; MATSCHIE, Flâ&#x201A;Ź'derm . des Berliner Mus. fur Naturk. (1899) 1; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 44.

Tragus never present; skull varying in rostral length but rostrum never strongly specialized ; premaxillaries strongly developed, usually free, always without palatal branch; postorbital processes well developed; teeth modified for fruit eating, those of upper and lower jaws resembling each other; never more than four incisors in lower jaw; claw present or absent on second finger of forelimb; second finger maintaining an evident degree of independence, the ungual phalanx always present; the humerus with trochiter and trochin reduced, the former never articulating with scapula; mandible with angular process broad and low or practically absent; margin of ear forming a complete ring. Approximately 230 forms of fruit bats are recognized. These belong to the family Pteropidre Gray. The Megachiroptera occur only in Asia, Africa, Malay Archipelago, Pacific Islands, and Australia. The distribution is not uniform, an'd in large areas no species of the group is known. The entire northern part of Africa, save Egypt, has no representative of the family. None occurs in Europe. Very few species extend north of the Himalayas. They are absent from the Hawaiian Islands, New Zealand, and Tasmania. The greatest number of genera and species occur in the Malay and Philippine Archipelagoes, particularly the large islands Sumatra, Java, Borneo, Celebes, Mindanao, and Luzon. The Philippines alone have no less than 23 forms, 21 of which are distinct species. Borneo has 11 forms, Java 9, Celebes 16, and Sumatra 11. The Philippines have 11 genera, Java 6, Celebes 11, Borneo 9, and Sumatra 7. 269468-8

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116

PHILIPPINE LAND MAMMALS

Several genera represented in neighboring areas have no species in the Philippines, but some of these are to be expected. Genera represented in neighboring areas Borneo.

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not in the PhJilippines.

Celebes.

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The occurrence of M egmrops in the Philippines is recorded for the first time in this work. The records of Dobsonia and Thoopterus for the Philippines are not recognized by Andersen. * Family PTEROPIDJE Pteropidm

GRAY, London Med. Repository 1 5 (1821) 299; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 45. (Most other authors have used this name or a variation of it.)

The characters of the family are those of the suborder Megachiroptera. The family is divided into four subfamilies, three of which occur in the Philippines. Key to the Philippine subfamilies of the Pte?'opidre. a 1. Premaxillaries broadly united anteriorly.... HARPYIONYCTERINJE (p. 116). Premaxillaries separate or barely united anteriorly. b'. Tongue highly extensible; molars and incisors considerably reduced in size; brain case strongly deflected ........ MACROGLOSSINJE (p. 119). b 2 . Tongue not highly extensible; molars and incisors not specially reduced in size, brain case never strongly deflected. PTEROPINJE (p. 134).

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Subfamily HARPYIONYCTERIN.JE:

Harpyionycterinre MILLER, Bull. U. S. Nat. Mus. 57 (1907) 77; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 799.

Premaxillaries broadly and solidly fused anteriorly (length of median suture considerably less than distance between canines), all their boundaries completely lost in adults; bony palate narrowing rapidly behind tooth rows; canines crossing each other at nearly right angles when jaws are closed, the lower canines

* Catalogue of the Chiroptera of the British Museum 2d ed. 1 (1912) 1-854.

RARPYIONYCTERIS

117

tricuspidate, almost in contact with each other, and lower incisors probably absent (a single one present in type); cheek teeth very unusually cuspidate, each molar with five or six distinct sharply pointed cusps. Only the genus Harpyionycteris Thomas is known. Genus HARPYIONYCTERIS Thomas

Ha?'pyionyctel'is THOMAS, Ann. & Mag. Nat. Rist. VI 18 (1896) 24'3; Trans. Zool. Soc. London 14 (1898) 384; MATSCHIE, Flederm. des Berliner Mus. fur Naturk. (1898) 70; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 77; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 11; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 799.

Upper incisor very large, filling all space between canines; canines with an obliquely projecting hooked cusp and a broad posterior heel, the two cusps in contact at middle, then diverging outward; lower incisor a minute spicule probably not normally present (in type present on one side only) ; canines slanting noticeably forward, very broad, with well-developed cingulum and large posterior secondary cusps, this in the upper canine almost equal in diameter to main cusp, though much shorter; in the lower canine the cingUlum forms a second supplemental cusp in front of the main shaft of the tooth; small upper premolar well developed though not' usually large, with distinct anterior cusp and narrow posterior heel; the corresponding tooth in the lower jaw, the second premolar, much larger, its size when viewed from the side nearly equaling that of the first molar; in form it somewhat resembles the canines though the cingulum and secondary cusp are less developed; middle upper premolar large, almost equal to canine; witb well-developed inner and posterior secondary cusps; it is closely resembled by the last lower premolar, the main cusp of which is, however, slender; last premolar, both above and below, with a distinct inner and posterior secondary cusp and a bifid main cusp; first lower molar of the same type but shorter; second and third molars, both above and below, much shorter than the other teeth when viewed from the side, each with three outer and three inner cusps between which extends the fairly well-defined median groove; skull not unlike that of Pte't opus in general form but occipital region apparently not tubular; deflection of occiput so slight that the alveolar line when proj ected backwards passes through base of zygoma; externally characterized by unusually short legs, absence of tail, presence of claw on index finger and attachment of wings low on sides of body.

118

PHILIPPINE LAND MAMMALS

The only species, Harpyionyoteris whiteheadi Thomas and H. celebensis Hollister and Miller, are confined to Mindoro and Celebes, respectively. HARPYIONYCTERIS WHITEHEADI Thomas

H a?'pyionycte?"is whiteheadi THOMAS, Ann. & Mag. Nat. Hist. VI 18 (18~6) 244; Trans. Zool. Soc. London 14 (1898) 384 pI. 30, fig. 1 (animal, and pI. 35 figs. 1-4 (skull and teeth); TROUESSART, Cat. Mamm. 1 (1897) 85; Cat. Mamm.,. SuppI. (1904) 61; MATSCHIE, Megachiroptera (1899) 70; SANCHEZ, An. Soc. Espanola Nat. Hist. 29 (1900) 276; MILLER, Bull. U. S. Nat. Mus. 58 (1907) 78; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 805 (also page LI) fig. 78, p. 800 and fig. 79 p. 802; HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 11; Proc. U. S. 'Nat. Mus. 46 (1913) 335.

Description of the species.- Characters of subfamily and genus. Size about that of Rousettus amplexicaudatus .. fur soft and wooly especially posteriorly; ears of medium length, rounded at tips; fur of back extending thinly on forearms, covering hind limbs densely to roots of claws; interfemoral membrane barely 2.8 millimeters wide) wholly buried in fur. Color.-General color above and below uniform chocolate brown, a little darker on face and a little lighter on nape and shoulders; wing membranes dark, with a few scattered whitish spots. Measu?'ements of the type of Harpyionycteris whiteheadi Thomas. mm.

Forearm Head and body Ear Index finger and claw Third finger metacarpal First phalanx Second phalanx Skull: Basal length Greatest breadth Interorbital breadth, tip to tip of postorbital processes Front of canine to back of second upper molar

84 140 17 60 59 44 54 37.5 23.8 6.9 17

Remarks.-The type is No. 97.5.2.7 in the British Museum; it was collected in Mindoro at an elevation of 1,660 meters, December 1895, by John Whitehead. Whitehead says:* This interesting new Frllit-Bat was shot by me in the Highlands of Mindoro at an altitude of 5000 feet. It was ~fIying around some high trees

* Trans.

Zoo!. Soc. London 14 (1898) 385.

MACROGLOSSUS

119

at dusk, at which time I generally sat out near my camp on the look-out for nocturnal birds. The specimen, when shot, fell into some tangled undergrowth, and it was only after a careful search with a lamp that my servant found it.

A related species has recently been discovered in Celebes by H. C. Raven. Subfamily MACROGLOSSINLE

Mam'oglossus DOBSON, Ann. & Mag. Nat. Rist. IV 16 (1875) 346; Cat. Chirop. Brit. Mus. (as part of Pteropidre). Carponycte?'in:e FLOWER and LYDEKKER, Mamm. Liv. and Extinct (1891) 654. Kiodotin:e PALMER, Proc. BioI. Soc. Washington 12 (1898) 111; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 68. Pteropid:e MATSCHIE, Flederm. des Berliner Mus. Naturk. (1899) 1 (part). Macroglossin:e ANDERSON, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 723.

Premaxillaries at first separate, but uniting and losing their distinctness later in life; bony palate narrowing gradually behind tooth-rows; mandibular symphysis elongated, its upper surface parallel with alveolar line; tongue highly extensible; teeth, except canines, much reduced in size. The subfamily has eight genera distributed from West Africa, to India, Malay region, Australia, and even extending in the Pacific to the Fij i Islands and the Carolines. Eonycteris and Macroglossus are known in the Philippines. Eonycteris is undoubtedly less modified and more nearly related to the Rousettus section of the Pteropidil:e than is Macroglossus. The chief differences between Eonycteris and Rousettus are the shorter tongue and the presence of a claw on the second digit. A third genus formerly recognized, Odontonycteris, has been shown by Andersen * to have been founded on an anomalous specimen having upper cheek teeth 6-6 instead of 5-5 as in M acroglossus. Key to Philippine genera of Mac?路oglossin:e. a '. Claw present on second finger; smaller .... Macroglossus Cuvier (p. 119). a'. Claw wanting on second finger; larger ........ Eonycte?'is Dobson (p. 126). Genus MACROGLOSSUS Cuvier

Macroglossus F. CUYlER, Dents. Mamm. (1824) 248; SCHINZ, Naturgesch. undo Abbild. Saugeth. (1824) 71; DOBSON, Cat. Chirop. Brit. Mus. (1878) 95; MATSCHIE, Flederm. des Berliner Mus. fiir

* Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 754.

120

PHILIPPINE LAND MAMMALS N aturk. (1899) 95; ANDE~sEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 746. Macroglossa LESSON, Man. de Mamm. (1827) 115. Kiodotus BLYTH, Cuvier's Anim. King. (1840) 69; PALMER, Proc. BioI. Soc. Washington 12 (1898) 117; MILLER, Bun. u. S. Nat. Mus. 57 (1907) 70. Rhynchocyon GISTEL, Naturgs. Thier. fur hahere Schulen (1848) 9 (non . Peters). Carpyonycteris LYDEKKER, in Flower and Lydekker Mamm. Liv. and Extinct (1891) 654. Odontonycteris JENTINK, Notes Leyden Mus. 23 (1902) 140. MILLER, Proc. BioI. Soc. Washington 18 (1905) 253; Bull. U. S. Nat. Mus. 57 (1907) 71.

Upper incisors subequal, minute, simple, distinctly projecting forward, the diameter of the crowns much less than that of posterior molars, the teeth of each pair separated by a space about equal to diameter of tooth at afveolus, the pairs separated from each other by a slightly wider space and from canines by diastema twice as wide; lower incisors similar to upper in form and position but outer tooth appreciably larger than inner, and median space much wider than that between outer incisor and canine, the latter interval about equal to that between the incisors of each pair; canines strong, well developed, without secondary cusp or prominent cingula, maxillary canines with deep longitudinal groove on anterior surface; cheek teeth small without distinct contrast of size and form, two anterior premolars in each jaw with crowns compressed and elevated into distinct though blunt cusps; all other cheek teeth with narrow nearly flat crowns, bounded by indistinct ridges, of which that on outer side is less developed; anterior premolar both above and below close to canine, but separated from second premolar by a diastema twice as great as width of its crown; in both jaws the anterior premolar is smaller than other premolars. Skull with a weak rostrum an'd a large brain case, occipital region not tubular but so strongly deflected that alveolar line projected backward passes between root of the zygomata and middle of brain case; postorbital processes short and little curved; development of audital bullre about as in Cynopterus,. mandible very weak, and ramus distinctly concave beneath; symphysis long, its greatest diameter equal to about twice the distance between outer sides of canines; externally characterized by presence of a welldeveloped claw on second , or index finger, the attachment of the

121

MACROGLOSSUS

wing membrane to base of fourth toe and low on sides of body; no external tail. Two species occur in the Philippines. r---~1I~8r-----~12o,O~-----'1"22~-------n12~------·12~6;r------.

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122

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126

Distribution of Macroglo8S11.8. Eonycteris. HOITPyionycteris, and six species of Rattus in the Philippines.

Key to the Philippine species of Macroglossus Cuvier. a '. Upper incisors small, subequal, terete; orbit extends to a point above the second upper molar or behind .... M. lagochilus Matschie (p. 122). a 2. Upper incisors large, subequal with broadened cutting edges; the orbit extends forward to a point above first molar. M. fructivorus sp. nov. (p.125).

122

PHILIPPINE. LAND MAMMALS MACROGLOSSUS LAGOCHILUS Matschie

Andersen recognizes four named forms of this species, separated by small average , differences in measurements. Only the typical form occurs in the Philippines. It is rather widespread, reaching from Luzon to Arnboina. The forms do not overlap in territory. MACROGLOSSUS LAGOCHILUS LAGOCHILUS (Matschie)

Mcw?'oglos sus minimus WATERHOUSE, Proc. Zool. Soc. London (1893) 67 (Philippines); DOBSON, Cat. Chirop. Brit. Mus. (1878) 96 (Philippines) ; ELERA, Cat. Sisto Fauna de Filipinas 1 (1895) 7 (Panay, Samar, Cuyo); Contrib. a la Fauna Filipina (1915) 79 (Luzon). Mam'oglossus minimus var. GRAY, Cat. Monk. Lemurs and Fruit-eat. Bats Brit. Mus. (1870) 115 (Philippines). Macroglossus aust?'alis (nec Peters) THOMAS, Proc. Zoo I. Soc. London (1888) 476 (part.) (Philippines.) Ca'rponycteris australis THOMAS, Trans. Zool. Soc. London 14 (1898) 385 (Canlaon Volcano, Negros) ; MATSCHIE, Sitzb. Ges. Nat. Freunde (1898) 39 (Tablas). Macroglossus lagochilus MATSC~IE, Megachirop. Berlin Mus. (1899) 97; HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912). Kiodotus lagochilus MILLER, Bull. U. S. Nat. Mus. 57 (1907) 71, fig. 10 (skull) (Philippines). Odontonycteris meyeri JENTINK, Notes Leyden Mus. 23 (1902) 140 (Tabukan, Great Sanghir); MILLER, Proc. BioI. Soc. Washington 18 (1905) 253; BulL U. S. Nat. Mus. 57 (1907) 72; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 11 (Cagayan Sulu). Macroglossus lagochilus lagochilus ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 763 (Negros); HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 306 (Cagayan Sulu).

Description of the species.-From No. 332, E. H. Taylor collection; collected at Culasi, Panay, May, 1918, by Richard C. McGregor. Adult male.-Muzzle elongate doglike, nostrils slightly projecting, openings large, directed forward and outward, separated by a groove extending from posterior extension of "tubes" to the lower lip; two short obsolescent grooves on either side of naked granular area; lower lip with two naked granular areas separated by a groove; either side of lower lip with wide elongate glandular area with a tubercle at posterior epds probably caused by the upper canines which push into lower lip in pockets which terminate directly opposite the rounded tubercles; eyes relatively large with an elongate tubercle above each, and a tubercle near angle of mouth; ears moderately large, tips somewhat truncate, anterior edge somewhat thickened and curving to top; a

MACROGLOSSUS

123

slight emargination posterior to tip; antitragal lobe distinct, triangular; ten transverse ridges on ear becoming shorter near middle of ear; tongue very long; wing membranes ample, antebrachial membrane attaching on proximal end of first phalanx of first digit; wing membranes attach on sides of body and to the metatarsal of fourth toe; calcar short; tail reduced to a slight vestige; interfemoral membrane narrow; foot rather small, toes elongate, the outer shortest; penis small; muzzle covere'd with very short fine hair; fur on forehead not or scarcely reaching to anterior line of eyes; back with moderately long fur; forearm furred on proximal half with short fur; tibia and interfemoral membrane covered with sparse long fur. Color in alcohol.-Above, pelage cream-buff, tips of the hairs somewhat cinnamon-buff; below on throat and belly creamy buff; on sides below wings with a cinnamon tinge on the hair !tips; wing membranes orange to cinnamon-buff. The colors are probably faded somewhat by immersion in alcohol. Skull and teeth.-Brain case strongly deflected, the alveolar line continued backward passes through brain case some distance above lambdoid crest; rostrum elongate, slender; anterior edge of orbit usually behind last molar; postorbital foramen present; temporal ridges low on sides of cranium never developing a sagittal crest; mandible narrow, fragile. Teeth small; upper incisors very small, subequal, with simple cutting edge; a wide space between first two incisors; upper canines with a deep vertical groove on anterior surface; premolars and molars very narrow, linear; last premolar and first upper molar with slightly concave crushing surfaces, with scarcely any trace of the cusplike elevation anteriorly; lower incisors small, subequal, or inner pair smaller; inner separated by a space more than double that between first and second incisors which is equal to distance between canine and second incisor; lower canine curving outward, edges rather sharp and angular, a faint shallow groove on anterior outer surface; first and second lower premolars separated by a wide diastema more than double the distance between the first lower premolar and canine; following teeth separated by diastemata growing smaller posteriorly. Remarks.-The maximum and minimum measurements are taken from Andersen. It will be noted that the measurements for my specimen approach the maximum measurements throughout. This form is rather widely distributed, and is known from Borneo, the Philippines, Celebes, and Amboina.

124

PHILIPPINE J.,AND MAMMALS

Measurements of the Philippine species of Macroglossus Cuvier.

--M.

lagochilu~

lagochilus.

M. frurtivarus.

E.H.T. collection. No. 332. male.

Total length _____ _ Head ______________________________________________ _ Tail _________________________________ . ______________ __

Maximumminimum.

mm. mm. 76 -----------27 .8 ------- ---- 0-1 1 5

E.H.T. collection, No. 710, male.

mm. 51 24 .2 . 75

Ear: Height from meatus ______________________________ _ Width ___________________________________________ _ Forearm ____________ . ____________ " ___________________ _ First finger and claw_______________

_ ______ . _________ _

Second finger: Metacarpal ______ __ _______________ __ __ __ _________ _

13-14 9. 5-11 38-44.5 15-18

16 9. 6 42 5 19

13. 2 8.9 35 .2 15

24 5 4

19 . 5-23 .5 4-5 4 . 5-6

19 4 5

Second phalanx ________________________________ _

33 24 28

27-33 19 . 5-23.5 24-28

25 18 . 5 22

Fourth finger: MetacarpaL _ ___ ___ __ __ __ _ ____ ____________ . _____ _ First phalanx- _ __ __ __ ____ _ _ __ ___ _ ___ _ ______ _ Second phalanx __ ____________________________ . ___ _

33 17 18

27-33 14-17 . 5 15 . 5-19

25 13 .2 14

33 15 17 17 .5

27-33 . 5 11.5-15 13.5-17 . 5 15.5-17

25 . 3 11 . 1 132 14 . 5

24 8-27 .2 7. 8- 9 .2 12.8-15 . 8 9. 8-11 6 . 5- 7. 3

22 .2 6.2 11.8 10 . 2 6 4.4 7 4 5 8

First phalanx- ___________________________________ _ Second phalanx __________ . _______ . __ ___________ _ Third finger:

Fifth finger : Metacarpal ______________________________________ _ First phalanx ___ ____ _____________________________ _ Second phalanx ____ ______ . ______________________ _ Tihia ________________ __ _____________________________ _

Skull: I Totallength ______________________________________________ _ Rostrum orbit to nares _____ __________________ . ___________ . __ Zygomatic width ______ _________ _________________ . ________ _ Width of brain case at zygomata _____ ________________________ _ Lachrymal breadth _________________________ . _ ___ __ _ __ __ __ _

Width across crowns of canines externally _____ . ___ ___ ___ _ ___ __ Postorbital breadth ________________________________ : ----. -Interorbital breadth _____________________________ . ' __________ 1 Orbital diameter __ _____ --- -- - - ----- -- - -- ---- ---- -

--1- ----- ----

upper tooth row ___ --- __ -- __ ----- --- ___ --- -- -- --

'1_ ----- ----

Lower tooth row ___________________________ - . _ _ __ _ Mandible_____________________ ___________________

9 8

19 . 8

4 5- 5.3 1 6 8- 7. 7 4.6- 5.1 I 5. 6- 6 .2 8- 9 2 8 8- 10 .5 18 2-20 7

Odontonycteris meyeri J entink is regarded by Andersen as undoubtedly a synonym of this form. The dental formula with six cheek teeth above and below is presumably an anomaly. He lists several specimens of Macroglossus having either more or less than the normal formula. One of the two known forms referred to Odontonycteris was taken in the Sanghir Islands,

MACROGLOSSUS

125

the other in Cagayan Sulu. It is not improbable that these apparently anomalous characters have become constant in these localities. The separation of the subspecies of Macroglossus lagochilus is based for the most part on small average measurement differences of which the typical form is somewhat the larger. The species is not particularly common in the Philippines or its habits are such that few specimens are taken. MACROGLOSSUS FRUCTIVORUS sp. nov.

Type.-No. 710, E. H. Taylor collection; collected at Tatayan, Cotabato, March 5, 1923, by E. H. Taylor. Alcoholic specimen with skull separate. Diagnosis; description of the type.-Y oung male; resembling in general habitus Macroglossus lagochilus lagochilus but with the nostril directed slightly more outward; small vibrissre-bearing tubercles on snout arranged in longitudinal rows on side of muzzle; ears smaller, narrower, distinctly more narrowed at tip, the emargination behind tip greater; anti tragal lobe angular, not thickened, prominent; wing membranes similar, attaching likewise above fourth toe metacarpal; interfemoral membranes narrower and the calcar shorter; other external characters similar to those of M. lagochilus lagochil~lS. Color.-General color of head and back cinnamon to russet, basal half of the hairs light wood brown approaching buff-ecru to drab; below nearly uniform buff-ecru approaching cinnamon high on sides under wings; membranes darker brown than fur; ears with edges blackish; feet dark blackish brown. Skull and teeth.-Skull even more strongly deflected than in M. Zag 0 chilus ; no temporal crests evident, probably due to its youth; supraorbital foramen present, small; postorbital processes small; rostrum short, the distance from orbit to nares contained in total skull length morâ&#x201A;Ź than 3.5 times; the orbit consequently is farther forward, its anterior edge reaching anteriorly to a point above the first upper molar; palate strongly concave; tooth rows gradually widening from canines backward; a median ridge on palate with two grooves on each side; upper incisors large, suggesting those of SyconycÂˇt eris; the four teeth separated by nearly equal distances, all with widened cutting edges, the two innâ&#x201A;Źr teeth directed inwards; second incisor separated from canine by a wide diastema; canines large proportionally, with a strongly defined deep groove from near tip to base on anterior surface; lateral surface with a single faint

126

PHILIPPINE LAND MAMMALS

broad depression; a strong heel behind; first premolar large, placed close behind canine; diastemata between three premolars subequal; lower incisors arranged in pairs separated medially by a distance equal to or more than double diameter of outer incisor; first and second lower incisors separated from each other by a diastema less than one-third the diameter of the first incisor; second incisor separated from canine by a like or slightly greater distance; canines large, curved outward or backward; a distinct broad shallow groove on outer surface, and a similar one on inner surface; a deeper more-pronounced groove on posterior surface; anterior edge sharp; first lower premolar close behind canine but separated from following premolar by a considerable diastema, equal to 1.5 times first premolar; last molars both above and below just beginning to appear. Meas~~rements.-See table under Macro'glossus lagochilus Zagochil.us. Remarks.-Only the type has been examined. The similarity to Macroglossus lagochilus externally and the fact that the skull and teeth seem to be midway between Syconycteris and Macroglossus have made this form difficult to place. In the cliaracter of the upper incisors, the more-forward position of the orbit, and the shorter rostrum it agrees with Syconycteris. In the characters of the lower incisors and of the first upper premolar it is more like Macroglossus. It is much smaller than recognized forms of Syconycteris. The type is the only specimen known. Genus EONYCTERIS Dobson

Eonycteris DOBSON, J.ourn. Asiat. Soc. Bengal 42 (1873) 204; Cat. Chirop. Brit. Mus. (1878) 94; MATSCHlE, Flederm. des Berliner Mus. fur Naturk. (1899) 89; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 69.

Externally like Rousettus, muzzle more elongate and index finger without claw; tail and calcar well developed; tongue long, sharply pointed, highly extensible; skull slenderer than in Rousettus, and the occipital region more deflected, alveolar line when continued backward passing through or slightly above root of zygoma; mandible not as heavy as in Rousettus generally and coronoid not rising as abruptly, otherwise differing only in the character of the symphysis peculiar to the subfamily; upper incisors of equal size, simple, shorter than in Rousettus but not distinctly different in form or position; lower incisors apparently similar to upper, directed distinctly forward, separated from each other and from canines by subequal spaces about as wide

EONYCTERIS

127

as diameter of incisors at alveolus; canines slender and not very large, without secondary cusps, and with barely indicated cingulum; front surface of upper canines marked by a deep longitudinal groove; cheek teeth essentially as in Ro'USettus, but lateral ridges and median grooves less distinct. . 2-2 1-1 3-3 2-2 Dental formula: 1. 2- 2' c. 1-1' pm. 3-3' m. 3-3 = 34. Five species belong to the genus, two of which-Eonycteris robusta Miller and E. longic(}'ud(}' sp. nov.-are known from the Philippines. The type locality for both forms is Montalban Caves, near Manila, Luzon. Key to the Philippine species of Eonycte?'is Dobson. a 1. Forearm, 71 to 76 millimeters; scrotal pouches furred with distinct gland present, ear rounded; tail, 15 to 18 millimeters. E ,. robusta Miller (p. 127). a t. Forearm, 75 to 81 millimeters; scrotal pouches naked without gland; ear rather pointed, longer; ; tail, 26 to 33 millimeters. E. longicauda sp. nov. (p. 131). EONYCTERIS ROBUSTA Miller

Eonycteris 1'obusta MILLER, Proc. tJ. S. Nat. Mus. 24 (1913) 73 (March 22); HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 305.

Description of the species.-(From No. 436, E. H. Taylor collection; collected in Montalban caves, Montalban, Rizal Province, Luzon, by W. Schultze, Albert T. Herre, and E. H. Taylor.) Muzzle moderately thin; nostrils not, or only slightly tubular, separated by a deep groove which continues to mouth, dividing two naked areas on upper lip; tip of lower lip naked; muzzle with very fine, short hair with a few longer vibrissalike hairs; eyes large, a small distinct tubercle above eye; ears short, rounded at tip without noticeable emargination anterior or posterior to tip; a small antitragal lobule posterior to basal notch; anterior edge of ear somewhat thickened; head covered with short velvety fur not exten'd ing to anterior edge of eyes; ear naked posteriorly, the anterior face with minute scattered hairs; back of neck and between shoulders sparsely furred; shoulders and back with short velvety fur growing sparse toward tail; tibia and interfemoral membranes naked save for minute scattered hairs; forearm with short rather dense hair on proximal third both above and below; wing membranes with a noticeable growth of long hair in region of elbow continuing to body but much shorter near body; attachment of wings high on back separated by a distance of 12 millimeters; on limb the attachment is between first and

436,

male.

44fi,

male.

449,

young male.

437,

female.

I fern44 444, female.

447, female.

Second phalanx---- --------------------- --- -------

Firat phalanx _____________________________ - - - - - - --

~et8carpal -- ----- -"---- --- --- --------- ------- - - - -

Fifth finger:

First phalanx ______ _____ _______ ______ __ _- __ - - - - . -Second phalanx- _____ __ ___________ __ _________ - __ - - '

~etacarpal-- ------ ------ ------ ---------- ---------

Second phalanx - _ ___ ___ __ __ - - - - - - -- - - --- - - -- - - --Fourth finger:

First phalanx _______________________________ - _- - --

~etacarpal ----------------------------- ---- ------

Third finger:

~::::::.::;:;;:::::::::-::::::::::::::-:--::::::-I

Second finger:

First finger and claw _________ _________ ____ ____ ________

Fore:;~_t~._ ~~~~~~~~~ ~ ~~ ~ ~ ~~ ~~~~ ~ ~~ ~ ~ ~~ ~~ ~ ~ ~ ~ ~ ~ ~ ~~ ~~ ~~ I

46 22 20

43 22 19

42 21 19

45 26 25

47 27 24

47 28 25

45 32 39

17.6 12 . 1 72 22 .5

119 104 38_ 15

17 11 . 5 7223

136 119 39 17

47 33 39

48 34 43

35 11 7

13.5 76 25

Total length __________________________________________ 140 Head and body ______ _________ ______ _______ ! ___________ 122 Head ___________ ______ ____________ ____ ___ __________ __ 40 Tail __________________________________ 18 ------------Ear: Height [rom notch _________________________________ I 17.5

45 20 111

4 2

2 2

47 26 25

43 21 19

47 26 25

47 26 24 44 21 19

49 31 42

7 47 32 42

4 3 3

18.2 12 73 23

19 12 74 23

47 5 31 41

32 11 8

129 114 40 15

129 112 38 17

7 2

3 1

12 11

11 . 5

132 U5 40 17

--------------------------------------------------------,-------, -------,-------,-------,------- - - - - - -

I 562,

female.

563,

young female .

43 21 20

47 27 26

50 32 45

23 . 5

18 12 . 5

133 116 40 17

42 21 18

47 25 24

48 32 41

32 9 7

19 13 73 23

------- .-

117 40

--.------

42 20 16

46 25 22

47 31 38

31 10 7

16.5 12 70 21. 5

121 105 39 16

- - - - - - - --

453,

female.

E. H. Taylor collection No. and sex.

[Measurements in millimeteI'B.]

Measurements of Eonyoteris 1'obusta Miller.

43 21 ]:9

47 27 24

48 32 42

32 11

18 12 72 23

126 110 40 16

---

565,

female.

I:'

t;'1

....

'"d '"d

....

~ .... t"

"1:1

Jo-'

t<>

......'"

Height of coronoid _________________________________

u~' Mandible _________________________________________

. . ."--------- ------------

Skull: Total length _______________________________ - - - - --Orbit to nares _________________________________ - - -. Zygomatic width ________________________________ . Width of brain cas(! ________________________________ Width across last molars- __________________________ Interorbital breadth ____________________________ --Width between canines _____________________________ Orbital diameter_--- _______________________________ Upper teeth _______________________________________

;~:i:~~d~~~~~::::::::::::::::::::::::::::::::::::::::1

35.1 11.1 24 16 8.9 7.4 4.7 8.5 13.3 14.3 27.2 9.8

35 17 35 11.5 22 15 9 6.8 4. 6 8.4 12 12.9 26.6 10

34 11.8 19 158.7 6.2 4.2 8.1 12.8 14. 4 27.5 8.2

31. 6 16.5

32 16.5

20 15.5 8.6 6.8 4.5 8.2 12.8 i3.9 27 . 0 9.2

32 16 32 16

31 17

32 18

8.5 12.4 14 . 1 27.5 10

7 6

36 12 21.1 15.2 9.8

31 18

30 18

12.11]------: 13.3 ______ _ 26 _______ _ 8 . 5 ___ ____ _

33 .2 10.9 19 16 8 .2 6.9 4.2 7.9

29 18

t".l

l\:) ~

~ ....

>-3

ISO

PHILIPPINE! LAND MAMMALS

second toes; calcar short, about 6 millimeters long; tail short, flattened, free for about two-thirds of its length; testes very large (16 by 10 millimeters) pushed back in two scrotal pouches at the tip of which on either side of the anus is an irregular kidney-sh~ped gland (6 by 3 millimeters) ; scrotal pouches covered with short fur save immediately surrounding the glands; penis very large with an anterior roughened basal area 4 by 4 millimeters; nuchal ruff prominent, the fur 6 to 10 millimeters long; fur on belly and breast very short, 2 to 2.5 millimeters long; feet moderately large, no claw on second digit. Color in life.-Pelage above, dark Mars brown; on head distinctly darker; on belly the fur tipped with silver, the basal twothirds light brown; wing membranes dark blackish, b.rown; the ruff on underside of neck light gray, brown at tip, ,growing darker toward base. Skull and teeth.-Skull distinctly 'd eflected, alveolar line projected passing just above attachment of zygoma; temporal ridges separated throughout their length but c0ming Â˘ldse together posteriorly; a slight occipital crest on each side, obsolete medially; postorbital processes strong. Upper incisors small, terete, separated by distinct spaces; a very strong anterior and a posterior .groove on canines; a broad shallow internal groove; outer surface of tooth flattened; a small cingulum visible posteriorly; first upper premolar twice the diameter and from three to four times the bulk of incisor, equidistant from canine and second premolar; lower incisors small, somewhat bilobed, arranged in pairs, the teeth in each pair very close to each other but separated from canines by a considerable space; lower canines .c urving outward and backward; an obsolescent groove on anterior face, slightly depressed laterally; cingulum visible posteriorly. Variation.-Three males, one a young specimen, and eight females, were obtained from the Montaiban caves. The adult males agree with the type in general characters; the young specimen does not have the testes extruded from the body cavity; the glands are smaller in the young specimen. Females differ from males in being of smaller size; the temporal ridges are usually more widely separated, and the neck ruff is wanting; in females 'the underside of neck is vert thinly furred; there is very slight difference in the color of the two sexes. Certain of the females contain embryos.

j - - - - --- - .-- - - - ._-

_ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _-=.,J .....

a~...

'-'

131

EONYCTERIS

R emarks.-This species was taken in Montalban caves together with numerous other species of cave-haunting bats. They were grouped together at the highest point in the roof of the cavern and there were several hundred specimens in the group. In the same cave several specimens of another species of Eonycteris were taken. However, only males of this species were obtained. The two forms are very similar in general appearance, and I think it surprising to find the two forms in such close proximity. The differences between the two forms are listed in the following table: Eonyc/eria robua/a.

mm. 71- 76 15- 18 Tibia ____ ______ ___ __ __ ______ ____________ ___ ______ ___ ___ _ 29-35 16 .. 19 132- 140 33-35 Height of coronoid procegs __ ____ _______________ __I___ _____ 8- 10 Furred. Present. R ounded, less emarginate.

______

mm. 75- 81 26-33 86- 42 20- 21 136- 153 35-38 10- 13

~:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~-~-~~~~~~~ ~~~~~~~

L -____________________________________

EOnyc: : ! lonoicau da .

Naked. Absent. More pointed, distinctly em a rginate an t e rio r ly a nd posteriorly.

____

Eonycteris robusta is obviously related to E. spelaea and E . majo'f and seems to be intermediate in size between the two forms. It differs in color and markings from E. sp ela ea. Measurements of E. major (known only from the type) for tail and ear as well as description of these same parts are wanting but it is presumed they are similar to E. spelaea. The tibia of E. 'i"obusta is distinctly larger than either of the two ~ther forms. EONYCTERIS LONGICAUDA

"p.

nov.

Type.-N o. 561, E. H. Taylor 'collection; collected in Montalban caves, Montalban, Rizal Province, Luzon, by W. Schulb:e, Albert Herre, and E. H. Taylor. Alcoholic skin and skull. Diagnosis .. description of the species.-This species is closelY related to Eonycteris 'fobusta Miller, but the following differ-

132

PHILIPPINE LAND MAMMALS

ences are evident. Larger and somewhat less robust, narrower through chest with more elongate head, body, and tail; distribution of fur on head similar; ears more elongate with a distinct emargination both anterior to and posterior to tip which is regularly rounded; tubercles somewhat more prominent above anterior part of eye; fur on back of head distinctly longer, on shoulders and back only slightly longer; lower third of forearm likewise furred; fur on breast, on belly, and in anal region distinctly longer; scrotal pouches when testes are extruded rounded and entirely naked, surrounded by sparse hair 5 to 7 millimeters long; when testes are withdrawn the pouch shows as a very strongly wrinkled area of skin; the gland at sides of anus apparently wanting; testes very much smaller; penis smaller with the roughened area near base when extruded apparently wanting; tail less flattened, nearly twice as long, ending in a small rounded tip; feet slightly longer and somewhat slenderer, membranes attaching between the first and second toes; antebrachial membr~ne below with a noticeable growth of elongate hairs, as are the wing membranes about the elbow and in region between elbow and femur; forearm longer and second phalanx of third digit very noticeably longer; also femur and tibia longer than in Eonycteris robusta. , Color.-Practically identical with preceding species. Skull and te(3!th.~Skull slightly less deflected than in E. robusta; alveolar line p'a ssing through base of zygoma when extended; rostrum somewhat longer and distance between anterior and posterior base of zygoma somewhat greater; audital bullre distinctly larger, middle incisors nearly double the size of same teeth in E. robusta; canines somewhat larger with posterior and anterior grooves; there is no groove on the inner side; molars and premolars distinctly narrower and somewhat longer, not visible when viewed from above, as is usually the case in E. robusta; lower incisors heavier, bilobed, arranged in pairs, members of each pair in contact, the two pairs separated by a space 2.5 times as great as their distance from the canines; lower molar-premolar series with teeth narrower, slightly more elongate with wider diastemata between the premolars; grooves on lower canines almost wholly obsolete; coronoid process more strongly developed than in Eonycteris robusta, whole mandible slighter and more fragile.

133

EONYCTERIS

M ea~ur,.ements of males of Eonycteris longicauda sp. nov .

[Meas urements in millimeters.]

E. H. T",,, '"

561

554

4391441

~ll",;o" No. 445

45.

438

- - - - - - --- --- --- ---- - - - --Totallength __________ 147 Head and body _____ __ 120 Head ___ ____ _________ 43 TaiL ________ __ ___ _ - - 27 Ear: Height from notch 20 Width flattened ___ 13 Forearm _________ - - - - 77 . First finger and claw __ 28 Second finger: MetacarpaL __ ___ _ 34 First phalanx ____ _ 11 Second phalanx __ _ 6 Third finger: MetacarpaL ___ ___ 52 First phalanx _____ 36 Second phalanx ___ 49 Fourth finger: MetacarpaL _____ _ 50 First phalanx ___ __ 28 Second phalanx __ _ 32 Fifth finger: MetacarpaL __ __ __ 47 First phalanx __ ___ 25 Second phalanx __ _ 25 Tibia _____ ___ _______ _ 38 Foot and claws _ _____ _ 21 Skull: Totallength ____ __ 36 Orbit to nares __ __ 12 Zygomatic width __ 22 Width of brain

case _________ . _ 16

146

136

122 41 27

120 43 26

115 41 -21

139 117 42 -21

138

133

116 42 -22

120 43 33

147 118 43 30

21 14 81. 5 26

21 14 80 27

21 14 76 27

21 14 77 26

20 12 . 6 77 26

20 14 79 26

20 14 78 26

20.5 14 . 1 76 24

20 13 . 8 75 26

37 11 7

37 10 8

35 10

33 10 7

35 11 8

31 10 8

35 10 7

34 10 6

54 37 52

54 35 49

50 34 46

50 34 47

52 34 48

52 34 50

52 36 47

51 34 47

49 36 47

52 30 32

52 28 30

49 28 31

49 27 31

51 28 32

51 29 30

51 29 31

49 26 30

50 29 31

51 25 26 42 22

51 23 24 39 22

46 19 24 36 21

47 22 23 38 20

48 22 24 38 22

50 24 23 39 21

50 25 24 40 21

47 23 24 36 20

47 23 23 37 22

-- - --- ----- - - - --- - - - .... ------ ------

35 12 22 . 5

38 12 . 5 23 . 8

36 12 22

36 12 . 2 22 . 5

15 . 1

------ - - - - -- ------ ----- -

15 . 2

16 . 5

15.6

------1------Âˇ_----- -----------1------ ----- ------

9.2

9. 5

9.3

9 .2

7.3

5 9.4 14 . 4 15 . 7 29 12

4.3 9 13 . 2 14 . 2 28 . 1 12

36 ---- -- - - - --12 . 2 --- --- -- ---23 ----- - --- - --

Width across last molars _________

9 6

Interorbital breadth ____ ____

7 .9

Width between canines __ ____ __

4.6 Orbital diameter. _ 9 Upper teeth ______ 13 . 6 Lower teeth ______ 14 . 6 I ~ible _ _______ 28 Height of coronoid 13

149

- --

4.9 ------ - -- --- - - ---.-8 . 8 ------ ------ ------ -----13 . 7 ------ - - - - -- ---- -- - -- - -14 . 8 -- - - -- ------ ------ .----29 - -- - -- ------ - -- -- ... - - -J2 ~ - - - --- - - -.- -._--- ------1

- --

â&#x20AC;˘ Tail injured.

5 8.8 13 . 7 15 . 1 28 . 7 11 . 6

141 115 42 26

144 115 40 29

4 .4 9 13 . 7 15 . 1 28 10 . 7

134

PHILIPPINE LAND MAMMALS

Variations.-The variation in measurements is relatively slight; the tip of the tail is injured in several specimens; the temporal ridges vary, some of them forming a sagittal crest, others remaining separate. In certain specimens the first and second incisors are not fully in contact and the median diastema is less th~m in the type. Remarks.-Not a single female of this form was obtained, }vhile more than a dozen males were taken; all the specimens are from the Montalban caves, near Montalban, Rizal Province, I ~uzon.

Subfamily PTEROPIN.IE

Pteropodidw DOBSON, Cat. Chirop. Brit. Mus. (1878) 3; MATSCHIE, Flederm. des Berliner Mus. fur Naturk. (1899) 1 (part.). PteropodinlE FLOWER and LYDEKKER, Mammals Living and Extinct (1891) 650. PteropinlE MILLER, Bull. U. S. Nat. Mus. 57 (1907) 45.

Premaxillaries separate, though usually in contact anteriorly, retaining their distinctness throughout life; bony palate narrowing gradually behind tooth rows, the width of interpterygo,i d fossa, including hamuli, distinctly less than distance between posterior molars; canines parallel, when jaws are closed; cheek teeth well developed without unusual development of cusps; tongue not especially elongated. The subfamily has about twenty-seven genera, eight of which Occur in the Philippines. Key to the genera of the Pte1'opinlE. p, '. Wing membranes attach on sides of body; calcar present; a claw on second finger. b '. Tail wanting; teeth 34; claw present on index finger. c '. First and second lower molars with broad, ,fiat, posterointernal heel in addition to the two longitudinal ridges. Acerodon Jourdan (p. 135). c 2. First and second lower 'molars without posterointernal heel. Pteropus Brisson (p. 147). , b 2. Tail present. c '. Teeth 34; claw on index finger .................... Rousettus Gray (p. 171). c ': Teeth 28. d '. Wings attach on middle of back; no claw on index finger. Dobsonia Palmer (p. 176). d 2. Wings attach on sides of body; claw present on index finger. Ptenochirus Peters (p. 178). c s. Teeth 30. d '. Tail short, terminal half free; teeth without special modifications .. ., ................................................ Cynopteru8 Cuvier (p. 181).

135

ACERODON d '. Tail reduced; cheek teeth enlarged, subquadrate.

Thoopterus Matschie (p. 188). b 3. Tail absent or extremely short; teeth 28.

Meg:ÂŁrops Peters (p. 190). Genus ACERODON Jourdan

Acerodon JOURDAN, L'Echo du Monde Savant et L'Rermes 4 (1837) 156; CUVIER, Ann. Sci. Nat. Paris 2, 8 Zool. (1837) 309; MATSCHIE, Megachirop. (1899) 9; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 59; ANDERSEN, Ann. & Mag. Nat. Rist. VIII 3 (1909) 20; Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 412. Pteropus DOBSON, Cat. Asia. Chirop. (1878) 15 (part.). ,

Like Pteropus but with smaller canines and larger more complex molariform teeth; upper incisors as in PteroP'l-f,s but somewhat more elongate and less bluntly pointed; lower incisors differing only slightly from those of Pteropus; canines mllch shortened as compared with most species of Pteropus, the mandibular canine exceeding the height of the third lower 'premolar; though reduced in length the canines retain their thickness, and the cingulum is even better developed than in mBst species of the related genus; small premolars as in Pteropus .. upper molariform teeth differing from those of Pteropus in relatively greater size, greater breadth of crowns, greater distinctness of cusps and a peculiar trenchant character of the edges and ridges; the first large premolar essentially as in Pteropus .. the three succeeding teeth differ from those of all other Pteropidre in the presence of a broad, flat posterointernal heel or shelf, sharply differentiated from the two ridges and extending considerably beyond them, especially behind. Cranial and external characters as in Pteropus. The type species is AceTodon fubatus Eschscholtz. . 2-2 1-1 3-3 2-2 Dental formula: 1. 2-2' c. 1-1' pm. 3-3' m. 3-3' = 34. Six species with nine forms have been recognized in this genus. They are confined as far as is known to the eastern Malayan Region (comprising Timor, Flores, Alor, Sumba, Celebes, Salayer, and Talant Islands) and the Philippines. Three forms occur i.n the Philippines-two subspecies of Acerodon iubatus and A. lucifer. Key to the species of Acerodon Jourdan. a. \ Forearm, 165 millimeters ................................ A. lucifer Elliot (p. 136). a.'. Forearm, 182 to 205 millimeters ............ A. jttbatus Eschscholtz (p. 138).

136

PHILIPPINE LAND MAMMALS ACERODON LUCIFER (Elliott)

Pteropus lucifer ELLIOT, Field Col. Mus. Publ., Zool. 1 (1896) 78, pI. 13 (skull) (Concepcion, Panay); Cat. Mamm. Field Col. Mus. (1907) 492; SANCHEZ, An. Soc. Espanola Hist. Nat. 29 (1900-1901) 276. 122 Batan I.

120

118

126

tfJ

PHILIPPINE ISLANDS

® Acepodon jubatus Jubatus Acepodon j . mtndanensis ~ Acepodon lucifer>

"<:)

lubans '<:;,o:'.

I;i.

DCatenduones

o Qlue

MINDORO

Busuan8a~ Culion~.'

abl.fJSifJya~~'ic·o ' . ~A~'~. .

/ANAY @®

••

; 8ur;.~ ~ol1)blon

Cuyo

Dumapan

.~ .

~r% ~~ ®

EBU

0.,0,0']' ,o"R05 PAL AWAN

C:)

<>t"geyan SuI" •

'DQ . :. Basdan

~jolO

<f~

•

~'ra~itawi

Sibutu:' \ ungau 120

FIG. 5.

122

124

126

Distribution of Acerodon in the Philippines.

Acerodon lucifer ANDERSEN, Ann. & Mag. Nat. Hist. VIII 3 (1909) 24; Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 432; HOLLISTER, Philip. Journ. Sci. § D 7 (1912) 10; Proc. U. S. Nat. Mus. 46 (1913) 306. Pteropus (Acerodon) lucifer TROUESSART, Cat. Mamm. Suppl. (1904) 48.

ACERODON

137

Description of the species.-Large fruit bat; ears broad, narrowing towards the tip, which is rounded; interfemoral membrane very narrow and entirely covered with hair in the middle; hair extending a short distance on the wing membrane above, along the humerus, and for nearly two-thirds the length of the forearm; tibia covered with hair for half the length above, naked below. Color.-Hair on top of head and occiput tipped with bright yellow; broad band on nape orange-ochraceous passing into reddish brown on upper back; sides of head beneath ears dark chestnut; rest of back to rump blackish brown, with a few yellowish hairs on middle of back; throat and entire underparts black, with numerous grayish hairs scattered over the body, and a slight reddish tinge below the throat; hair on wing membrane above, along humerus, blackish brown; below, hair reddish brown. Skull and teeth.-Sagittal and occipital crests present; orbits incompletely surrounded by bone; first premolar deciduous; upper incisors pointed, outer broader than inner; lower incisors very small but filling space between canines, the outer about twice the width of the inner; upper premolar, first and second upper molars, and second lower premolar with two anterointernal basal cusps, the anterior one very small; first lower premolar slightly larger than last lower molar and both with round crowns; last upper molar triangular, with one anteroexternal cusp. (After Elliot.) Measurements.-See under Acerodon jubatus jubatus, page 140. Remarks.-The type was collected at Concepcion, Panay, by J. B. Steere, and is in the Field Columbian Museum with one topotype. Steere (Elliot op. cit.) writes: The black bats of Panay were found occupying a roost in some tall clumps of spiny bamboos, among the rice fields and almost over some of the native houses. They were thickly clustered in the tops of the bamboos, hundreds of them together. As we fired among them they rose and Sew about overhead, squealing as they :flew, but soon settled again. There seemed to be at least three species, inhabiting the roost together, but each species in trees by itself. All the species of Pteropus from Panay sent you, were from this roost. Elliot also states, "The sagittal and occipital crests vary apparently among individuals, possibly on account of age, for while the type has low crests, the other specimen possesses crests of considerable height."

138

PHILIPPINE LAND MAMMALS

ACERODON

JUB~TUS

(Eschscholtz)

This form, originally described from the Philippines by Eschscholtz, has recently been split into two subspecies, Acerodon jubatus jubatus Eschscholtz, and A. j. mindanensis Andersen. They differ only in size. There is considerable variation between i-ndividuals of the two forms in color and markings, but the same variations occur in members of both subspecies. The typical form appears to be present in most of the larger islands north of Mindanao; the second fOil'!IIl is known only from Mindanao. Key to the subspecies of Acerodon juba,tus Eschscholtz. a '. Averaging smaller; forearm, 182 to 198 millimeters.

A. j. jubatus Eschscholtz (p. 138). a '. Averaging larger; forearm, 205 millimeters. A. j. mindanensis Andersen (p. 142). ACERODON JIIBATUS JUBATUS (Eschscholtz) Pteropus juba,tus ESCHSCHOLTZ, Zool. Atl. pt. 4 (1831) 1, pI. 16 (Manila): LESSON, Hist. Nat. Mamm. 5 (1836) 59; TEMMINCK, Mon. Mamm. 2 (1837) 59 (Luzon) ; WATERHOUSE, Cat. Mamm. Mus. Zoo!. Soc. 2d ed. (1838) 13, (Luzon); WAGNER, Schreber's Saug., Suppl. 1 (1839) 343 (Luzon); BLAINVILLE, Ost. Mamm., CMir. (1840) 100, pIs. 1, 2, and 9-13 (Luzon); WATERHOUSE, Proc. Zool. Soc. London (1843) 67 (Philippines); SCHINZ, Syst. Verz. Saug. 1 (1844) 120; DESMAREST, Dict. Univ. d'Hist. Nat. 11 (1848) 250; GRAY, Zool. 'Samarang,' Vert. (1849) 11 (Philippines); WAGNER, Schreber's Saug., Suppl. 5 (1853-55) 595 (Luzon); GERVAIS, Rist. Nat. Mamm. 1 (1854) 187; GEIBEL, Saug. (1855) 995; FITZINGER, Sitzb. Akad. Wien 42 (1860) 389 (Manila); PETERS, Monatsb. Akad. Berlin (1861) 707 (San Fernando, Albay, Luzon); (1867) 332 (Luzon); ZELEBOR, Reise 'Novara' Saug. (1869) 12 (Manila); GRAY, Cat. Monk. Lemurs and Fruit-eat. Bats (1870) 105 (Philippines); FITZINGER, Sitzungsb. Akad. Wien 60, Abth. 1 (1870) 403; MARCHI, Atti Soc. Ital. Sci. Nat. 15 (1873) 515 (hair structure); DOBSON, Mon. Asiat. Chirop. (1876) 28 and 188; Cat. Chirop. Brit. Mus. (1878) 68 p!. 4. fig. 6 (Luzon, Dinagat) .; TROUESSART, Rev. and Mag. Zoo!. III 6 (1879) 203; GUNTHER, Proc. Zoo!. Soc. (1879) 74 (Dinagat, S. Leyte, S. Negros); STEERE, List Bds. & Mamm. Steere Exp. Philip. (1890) 28 (Leyte, Panay) ; ELERA, Cat. Sisto Faun. de Filip. 1 (1895) 6 (part.) (Luzon, Dinagat); Contr. a la Fauna Filipina (1915) 71 (part.) (Luzon, Dinagat) ; ELLIOT, Field Col. Mus. Publ., Zoo!. 1 (1896) 76 (Panay); TROUESSART, Cat. Mamm. 1 (1897) 83; THOMAS, Trans. Zool. Soc. 14 (1898) 383 (Barit, Abra Dist., N. Luzon); SANCHEZ, Ann. Soc. Espanola Hist. Nat. 29 (1900-1901) 242, 275, 288; ELLIOT, Cat. Mamm. Field Col. Mus. (1907) 4'92 (Panay). Ace?'odon juba,tus LESSON, N. Tabl. R. An., Mamm. (1842) 14; HEtJDE, ~em. Hist. Nat. Emp. Chin. 3 (1896) pI. 5, figs. 11, 12; MILLER, Bull. U. S. Nat. - Mm;. 57 (1907) 59.

ACERODON

139\

Pteropus pyrrhocephalus MEYEN, Nova. Acta Acad. Caes. Leop.-Carol. 16 2 (1833) 604 pIs. 45, 46; SCHINZ, Sysrt. Verz. Saug. 2 (1845) 14. Pteropus aurinuchalis ELLIOT, Field Col. Mus. Publ., Zoo!. 1 (1896) 77, pI. 12 (skull, Leyte); SANCHEZ, An. Soc. Espanola Hist. Nat. 29 (1900-1901) 276; ELLIOT, Cat. Mamm. Field Col. Mus. (1907) 492 (Leyte). Pteropus (Acerodon) aU'rinuchalis TROUESSART, Cat. Mamm., Suppl. (1904) 48. Acerodon jubatus jubatus ANDERSEN, Ann. & Mag. Nat. Hist. VIII 3 (1909) 24, 26-29; Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 427; HOLLISTER, Phil. Journ. Sci. ยง D 7 (1912) 10; Proc. U. S. Nat. Mus. 46 (1913) 306 (Luzon, Mindoro, Leyte).

Description of the species.-Large; ears about as long as muzzle, obtusely pointed; interfemoral membrane narrow in the center, covered thinly with fur; fur on throat thin; body fur, except on back below shoulders, short and smooth; little hair on wing membranes above; on lower side humerus completely covered with blackish-brown fur; this extends over membrane to two-thirds length of forearm. Color.-Sides of face, forehead, a line over eyes, and throat black; crown of head dusky, hairs tipped with yellow; nape golden yellow, a narrow line of orange on back of neck; sides of neck and upper part of back at base of neck maroon, grading into brownish black, and succeeded on rest of back by reddish brown; fur on lower side of body black, a maroon tinge on front of neck; many yellow hairs interspersed among black hairs on body below chest; thighs covered with blackish brown fur; membranes pale brown. Skull and teeth.-First upper premolar deciduous; premolars and molars without very noticeable posterior basal projections; upper incisors with a slight space between the middle and outer, the middle the larger, widely separated from canines; lower incisors grooved on top and behind, middle ones distinctiy smaller than outer, separated from each other and from canines by equal spaces and filling all the space between canines; a wide diastema between upper canine and second premolar; canine large, heavy, with an anterior groove, a slight internal groove and a posterior groove; cingulum prominent, forming a slight cusp; molars and premolars longer than broad in both jaws, inner cusps well developed; last lower molar with a round crown; last upper molar longer than first lower premolar, which is about equal to last lower molar; teeth of both jaws, with the

140

PHILIPPINE LAND MAMMALS

exception of last two lower molars, sharp and pointed; lower canine small with a very slight cingulum cusp. A moderate sagittal and occipital crest; the process from zygomatic arch and postorbital process fails to encircle orbit by a space about 3.5 millimeters wide; a deep frontal groove; nasals much widened at anterior end; supraorbital foramen present. Measu?'ements of Acerodon jubatus jubatus and A. lucifer, from Andersen. [Measurements in millimeters.] Acerodon jUba,tU8 Acerodo1t jUba,tus. ~ucifer.

Height of ear from orifice Width of ear flattened Forearm First finger and claw Second finger: Metacarpal First phalanx Second and third phalanges Third finger: Metacarpal First phalanx Second phalanx Fourth finger: Metacarpal First phalanx Second phalanx Fifth finger: Metacarpal First phalanx Second phalanx Eye to tip of muzzle Depth of interfemoral at center Tibia Foot with claws Calc&r Skull: Total length Orbit to tip of nasals Zygomatic width Width of brain case Width across outer canines Interorbital constricti~m Or bital diameter Mandible length Coronoid height Upper tooth row, without incisors Lower tooth row, without incisors

30.5- 33 19 - 21.5 182 -198 77 - 86.5

165 72

85 - 99 18 - 25 19 - 24

82 21 20.5

120 -132.5 92 - 99.5 127 -143.5

115 88.5 116

117 -130 74.5- 82 70 - 82

114 70.5 64

122 -136 54 - 59.5 50 - 59 32 - 35 6 - 10 86 - 94 56 - 60 21.5- 29

120 52.5 49

77.225 41.326.715 10.514.862 31.532 34.2-

84 29 47 27.5 17.2 13.2 16 68 36.8 36 40

79 ?50

71.8 23.3 38.5 26.2 15.2 11.8 15 58 28.3 30.8 32.8

ACERODQN

141

Variation.-There is distinct variation in color. The color on the back is generally seal brown but occasionally approaching burnt umber. The breast, belly, and flanks vary from dull brown to maroon; there is some variation in the number of buff-colored hairs scattered through the pelage; the anterior part of neck in most specimens is chocolate or chestnut, but often blackish like the chin and throat; the neck spot varies from cream-buff to yellowish, usually tinged with ochraceousbuff. The variations do not depend upon age, sex, or locality. Remarks.-It is probable that the Cabag vel Panic qui of Camel, * from the Philippines, is this species. Meyen's Pteropus pyrrhocephalus and Steere's Pter-opus aurinuchalis have been examined by Andersen and both are regarded as synonyms of this species. Steere's specimens were obtained in Leite by himself. He writes (Elliot, loco cit.) : The two islands (Samar and Leyte) are practically one in fauna, and are only separated by a narrow river-like arm of the sea. In this channel, at a small low island, about half a mile from either shore, I procured the bats sent you, labelled Leite. Here there seemed to be several species inhabiting th~ same roost and they were hanging from the branches of tall, bare bamboos. The fruit bats of the Philippines prefer small islands ,f or their roosts, but will take up with other isolated localities. r found one roost on Negros occupying one immense hardwood tree standing by itself far from the foresll on the plains of the western side of the island. Where they are not hunted io.r food by the natives, they sometimes roost near the native houses or villages for pretection. Their roosts seem to be permanently occupied. We feund the specimens we procured had been feeding upon the palm juice which the natives were collecting for saquir (toddy). The bats visit the trees at night and drink the juice from the cups hUng on the trees.

J. Whitehead (in Thomas, loco cit.) writes: This large Fruit-Bat was in immense numbers in the Province of Abra, N. Luzon, where it had taken possession of a long, low range of hills, well covered with forest. Just at sunset th~se Bats issued from their roosting-place in thousands toward all points of the compass. Numbers of those that passed the Abra river dipped to drink in the stream, but seemed afraid, making often several attempts before they dared to come low enough to touch the water. On 'the sea-coast also the large Fruit-Bats often dip to drink in the sea on calm everungs. This Bat has a peculiar, though not disagreeable, odour. The wings are quite sticky to the touch.

Through the kindness of Mr. C. D. Bunker, curator of Birds and Mammals of the Dyche Museum, University of Kansas, I was enabled to examine three skeletons of this species from

* Phil.

Trans. 25 (1708) ?198.

142

PHILIPPINE LAND MAMMALS

Manila; the body bones are largely disarticulated. The skulls are complete and are well within the measurements here given for the subspecies. They were presented to the museum by Mr. Handel T. Martin, curator of Vertebrate Paleontology at the University of Kansas; the skins apparently were not preserved. ACERODON JUBATUS MINDANENSIS Andersen

Pte?"oprus jubatus GUNTHER, Proc. Zool. SOC. London (1876) 735 (Mindanao); DOBSON, Cat. Chirop. Brit. Mus. (1878) 68; TROUESSART, Rev. & Mag. Zool. III 6 (1879) 203 (part.); Cat. Mamm. 1 (1897) 83 (part.); ELERA, Cat. Sisto Fauna de Filip. 1 (1895) 6 (part.); Contrib. a la Fauna Filipina (1915) 71 (part.). Pteropus (Acerodon) jubatus, MATSOHIE, Megachirop. (1899) 9; TROUESSART, Cat. Mamm. Suppl. (1904) 48. Acerodon jubatus mtindanensis ANDERSEN, Ann. & Mag. Nat. Hist. VIII 3 (1909) 24-29 (Mindanao); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 10; Proc. U . S. Nat. Mus. 46 (1913) 306 (Mindanao); ANDERSEN, Cat. Chirop. &rit. Mus. 2d ed. 1 (1912) 429.

Descriptio.n of the species.-(From No. 641, E. H. Taylor collection; collected April 8, 1923, at Kraan Point, Cotabato coast, Mindanao-.) , Very large, similar to A cerodon jubatus jubatus but rather larger in average measurements; nostrils somewhat tubular, separated by a deep groove which continues to mouth giving a harelip appearance; anterior part of muzzle apart from rhinarium clothed in very fine short hair, with a fringe of longer hair about lip (10 to 12 millimeters) ; fur on forehead and sides of head rather short (7 to 8 millimeters) ; fur on upper neck somewhat wavy and longer and continuing thick to a line between anterior attachment of wing membranes (8 to 10 millimeters) ; on back fur scant and shorter, straight (about 5 millimeters) ; on rump longer and wavy; interfemoral membrane proximally heavily furred; outer edge naked; upper part of tibia covered with fur on inner side; foreneck and lower part of body covered with elongate wavy fur; lower part of forearm with scant elongate fur; wing membranes attach to base of first phalanx of second toe; interfemoral membrane about 13 to 14 millimeters deep medially; penis very large, pendent; ear large, curving very gradually anteriorly and posteriorly to a bluntly pointed or semirounded top; a ruff, on each side of neck, composed of extremely dense (one cann0t push them aside to expose skin) stiffened hairs intermingled with hairs considerably longer; shorter hairs 30 millimeters; hairs composing ruff 15 to 20 millimeters. Wing spread 1.550 millimeters.

ACERODON

143

Colo'f .-Anterior part of head, chin and sides of head dark blackish, the hairs mixed with numerous gray hairs; on upper part of head from a point between eyes and to middle of upper neck a large spot, anteriorly cream-buff intermingled with black hairs, varying to gray-buff with only occasionally a dark hair; posterior part of spot yellowish to orange-buff where it merges into the chestnut-red of mantle on sides and chocolate-brown of posterior part; lower parts of ruff darker chestnut; foreneck brownish to chocolate; back and rump seal to blackish brown. intermingled with few gray to yellowish buff hairs; belly darker chocolate to blackish with numerous gray to silvery hairs; wing membranes and feet dark blackish; ears brownish. Skull and teeth.- In general the teeth are the same as those in the northern subspecies. Upper incisors equally spaced from each other, the two inner distinctly longer than outer, and extending much farther, as the first and second pairs do not rise from the same level; a slight bench on inner side of teeth; canines very large, distinctly curved backward, a long narrow groove on anterior face, and a less conspicuous groove on posteroexterior face; a shallow groove on the antero-internal face and a deeper more promiNent one on the postero-internal part of tQoth; cingulum prominent, rising to form a slight heel cusp; first premolar deciduous (present in some specimens as a small terete tooth about one-fifth the bulk of incisors placed E짜luidistant between canines and second premolar) ; second premolar large molariform, bearing a small anterointernal cusp which is more developed on third premolar and on first molar; teeth in profile show a distinct anterior notch; lower canines without distinct posterior or anterior grooves; cingUlum pronounced but not forming a cusp; lower incisors unequal, first much shorter and much less than half the bulk of second; latter close to canines; all lower incisors somewhat bifid; first lower premolar somewhat larger than last lower molar; lower mOlar-premolar series with a strong longitudinal groove which divides the tooth into two large cusps, the inner longer and more blunt than outer save in case of second premolar; posterior heel present on second premolar, more prominent exteriorly; on following teeth more prominent internally; last molar small, resembling first premolar.*

* The teeth of. No. 641 are somewhat worn, and details were supplied from skull No. 636.

~etacarpal

~etacarpaL _____________ First phalanx ____________ Second phalanx. _________ Fifth finger: ______________ First phalanx- ___________ Second phalanx __________

Fourth finger:

First phalanx----- . ______ Second phalanx __________

~etacarpal--------------

First phalanx- ___________ Se('ond finger: ~etacarpaL _____________ First phalanx- ___________ Second and third phalanges. Third finger:

~etacarpal--------------

First finger: Length with claw _________

Height _________________ . Width ___________________ Forearm _____________________

Ear:

637. male.

641. male.

644. male.

142 65 62

134 62 59

135 89 87 140 63 60

145 61i 63

140 60 56

140 60 60

135 58 62

137 111 166

137 87 88

140 110 163

137 103 158

98 24 25

90 20 70

133 86 79

102 24 23

98 27 24

28 18.2 201

136 85 86

90 20 70

88 18 70

36 23

---

--- --29 18 192

631. female.

630. female.

629. female.

133 83 84

137 103 157

138 104 150

135 107 163

135 99 146

133 87 87

102 24 25

98 23 25

102 27 25

96 24 17

133 84 85

87 18 69

31 19 193

86 20 66

28 20 192

84 18 66

34 20 206

88 21 67

33 21 202

--- --- --- ---

632. male.

634. female.

636. female.

144 67 64

147 63 62

144 112 168

142 108 160

140 92 88

104 27 25

111 23 22

143 87 87

95 20 75

34 22 207

92 18 74

33 22 210

642. female.

135 86 90 142 65 65

150 62 65

85 136 58 61 134 58 55

144 92 90

140 65 53

136 88 79

143 111 156 136 98 150 128 100 152

132

142 110 156

138 108 153

102 23 24 94 23 24 95 23 19

130 87 80

97 24 020

102 28 25

84 19 65

28 18 195 86 18 68

29 20 209 90 19

31 21 195

91 20

35 20 197

--- --- --- ---

636. female.

Bureau of Science No. 147. 643. female. female.

84 18 66

33 20 200

--- --- ---

633. female.

E. H. Taylol" collection No. and sex.

[Measurements in millimeters.]

Measurements of Acerodon jubatus mindanensis Andersen.

00.

> t"'

a:: a::>

t:'

~ Z

t"' ..... '"0 '"0 .....

'"0

::r:: .....

~ ~

'"'"

""h ".---------

Lower leg ___________ - _____ - - - - -- ___ - -- - - - --Foot with claw__ . __ ____ ___ ___ 60 62 Calcar __ ___ ________ ___ ____ __ 23 22 Skull: Totallength _____________ 79 . 5 84 Pala tion to incisive fora41. 6 44 men. Front oC orbit to tip of 24 28 nasals. W~dth oC brain case ______ _ 27 26 Zygomatic width ___ __ ____ 43 . 2 45.3 25 . 8 25 6 Width across first upper molars. Width across canines ______ 16 16 Interorbital constriction ___ 13 12 Postorbital constriction_ .. __ 10 . 2 10 Orbital diameter __ ____ ___ _ 16 15 . 4 Mandible length _____ __ ___ 68 65 Coronoid process height ___ 36 33 85 34 Lower tooth row _________ 37 40 Lower incisors combined __ 7. 6? 7. 6

27 48 . 3 27

18 13 8.9 16 69 35 35 39 7. 8

26 . 8 47 25

16 12 . 5 9 16.8 66 36 34 37 7. 3

86 43

92 61 32

82 43

96 61 26

16 13.5 9 15 67 34 85 38 8

27 . 3 44 24 . 6

28 . 1

83 43

102 66 29

â&#x20AC;˘ Claw missing.

16 13 9 16 67 35 35 37 7. 5

27 45 28

26 . 5

83 42 . 6

88 68 26

16 . 2 13 . 5 8. 5 16 5 68 34 86 37.7 8

28 44 28

86 . 2 4.6

97 62 28

-- - -----

66 28

~ -------

28 44 26

82 43

99 61 28

16 16. -- ----- 12 13.5 13 9 8 9 15.6 16 15 . 5 67 66 66 36 32 -- -- ._-36 34 84 . 6 37 38 37 7.5? -- -- --- 8

28.5 26 46 44 27 -- - -----

82 43

96 60 31

8 17 68 34 34 87 8 9

16 70 33 36 38 7

- - -----

17 14 10 16 67 35 34 37

17 14

17 13

16 66 34

17 13

28 45 26

29 47 25

29 45 27 . 2

28 28.5 46 26.2

83 42

98 69 22

86 44

97 63 24

86 45

94 66 28

86 45 . 6

66 27

- -- ... ----

I-l

t:;)

ed0

;I> C'::l

146

PHILIPPINE LAND MAMMALS

Remarks.-Most of my specimens were collected near Kraan Point, Cotabato coast, Mindanao. The habitat covered 8 to 10 hectares of a small swampy forest near the mouth of Kraan River. The trees were of moderate height, and each tree held as many bats as could conveniently get a foothold on the branches. They hung by their legs, head down, continually moving and scolding, impinging on one another's roosting place, scratching themselves with their wings, or biting at some parasite in their fur. Acerodon jubatus mindanensis and Pteropus vampyrus lanensis were found in this colony. I do not know whether the species occupied separate trees or were mixed together, since the bats took flight before I was close enough to distinguish the species. After they settled down both species could be discerned on the same tree-even on the same branch. The chance collecting yielded twelve specimens of Acerodon jubatus mindanensis and five of Pteropu.c; vampyrus lanensis which probably represent the respective proportions of the two species in the colony. Many of the females carried young, which held on to the fur of the mother by means of their <Claws. The females that carried young kept one wing in a constant fanning motion while hanging, as if to help the young keep cooll, since the rather thin foliage on the trees offered only partial protection from the sun. One would expect to find a considerable accumulation of guano on the ground under the roosts, but the ground under the trees was packed hard by the tread of wild boars. These animals apparently visit the place each night and dispose of the day's accumulation of guano. The bats apparently never molest the cultivated fruits grown in the small coastal clearings; such as, pineapples, chicos, mangos, and bananas. . At sundown most of the members of the colony take flight and wing their way to the mountains 10 to 15 kilometers from the coast, where they feed on wild fruits. They apparently r.eturn before daylight as I have seen none in flight after daybreak. A few specimens remain in the colony, probably females about to give birth to young or old or wounded specimens. A conservative estimate of the bats in this colony based on the area of the forest inhabited, and on counts of specimens in several trees from various parts of the area, was about 150,000. Over two hundred were counted in several individual trees. The Acerodon species is distinctly the larger form; it has a wing spread of 1,550 millimeters (approximately 5 feet 2 inches)

PTEROPUS

147

and is the largest form occurring in the Philippines. The wing spread of the Pteropus species in no case exceeded 1,400 millimeters (approximately 4 feet 8 inches). However, one of the Malayan forms of Pteropus vampyrus is, I believe, the largest bat known. These bat colonies are not particularly rare. More frequently they exist on small islands off shore. There is a large colony on a small island off Port Banga, Zamboanga Province, where Edgar A. Mearns collected specimens of the two species. I have passed the island but circumstances did not permit of my landing. Dr. Albert T. Herre writes me of finding a large colony in the interior of Cotabato. This colony certainly contains Pteropus vampyrus lanensis, but whether other species are present I cannot say since only Pteropus was collected. There is a considerable colony on a small island off l the coast of Palawan somewhat to the north of Puerto Princesa. The island is covered with mangrove and the bats fly to the mainland of Palawam to feed. I failed to obtain specimens so I am uncertain as to what species the colony contains. The type of the subspecies was collected by J. B. Steere in Mindanao. It is No. 76.10.4.1 in the British Museum of Natural History. Genus PTEROPUS Brisson

Pteropus BRISSON, Regn. Anim., in Class. 2d ed. (1762) 13; DOBSON, Cat. Chirop. Brit. Mus. (1878) 15 (part.); MATSCHIE, Flederm. des Berliner Mus. fur Naturk. Megachir. (1899) (part.); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 56; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1913) 61. Spectrum LACE:PEDE, Tab1. des Div. Sousdiv. Ord. et. Gen. des Mamm. (1799) 15; MATSCHIE, Flederm. Berliner Mus. fur Naturk. (1899) 19 (as subgenus). Eunycte1'is GRAY, Proc. Zool. Soc. London (1866) 64; MATSCHIE, Flederm. des Berliner Mus. fur Naturk. Megachir. (1899) 30 (subgenus). Pselaphon GRAY, Cata1. Monk. Lem. Fruit-eat. Bats (1870) 110. Sericonycteris MATSCHIE, Flederm. des Berliner Mus. fur Naturk. Megachirop. (1899) 30. Desmalopex MILLER, Bull. U. S. Nat. Mus. 57 (1907) 60.

Incisors terete with slightly differentiated crowns, the lower bluntly rounded, the upper with a fairly defined cutting edge.; upper incisors in contact or nearly in contact with each other, but separated from canine by a wide diastema; they form a slightly convex row, the outer tooth of each pair distinctly shorter than the inner; lower incisors separated from each other and from canines by minute spaces, the row slightly convex and

148

PHILIPPINE LAND MAMMALS

inner tooth of each pair smaller than the outer; canines simple without secondary cusps, the cingulum rather prominently developed on posterior and inner sides and the shaft of particularly the upper teeth marked by conspicuous longitudinal furrows; anterior premolar above and below small, that in upper jaw much smaller than incisors and usually deciduous, its crown bluntly pointed; that in lower jaw slightly larger than outer incisor, permanent, its crown rounded in outline, the surface. concave, the outer edge higher than inner; posterior molars, above and below, essentially like the anterior lower premolar in size and form; but posterior upper molar usually with a more prominent inner edge and often with a distinct outer anterior CUSD or tubercle; other cheek teeth, molars and premolars rather closely resemble each other, all formed on the same general plan of a subquadrate or somewhat elongated crown with a distinct median furrow in the long axis of jaw, and a well-developed outer and inner ridge, each ridge fending to rise to a cusp in front of middle; outer ridge and cusp always larger than inner, and in the region between cusps the furrow often obliterated partly. Skull moderately elongate, length of rostrum considerably greater than lachrymal breadth; premaxillaries well developed, in contact anteriorly but not fused, their breadth along nasal suture at least equal to that at alveolus of outer incisor; occipital region strongly deflected and tubular, floor of brain case forming so great an angle with palate that alveolar line projected backward passes through tympanic region or base of zygoma; audital bulla reduced to a mere ring; taB absent; a narrow interfemoral membrane; well-developed calcar present; large claw on index finger. (After Miller.) . 2-2 1-1 3-3 2-2 Dental formula: 1. 2-2' c. 1-1' pm. 3-3' m. 3-3 = 34.

Remarks.-This genus comprises the bulk of the large fruiteating bats. The genus as here understood includes Miller's Desmalopex which Andersen has shown is not worthy of retention. This genus was made to include Pteropus leuGopterus Temminck, a species that occurs in the Philippines. Key to the Philippine species of Pteropus Brisson.

a 1, Canines heavy with broad cingulum; cheek teeth much shortened; ears short, usually hidden in fur; orbit encircled. P. Zeucopterus Temminck (p. 150). a 2. Canines moderate, with moderately broad cingulum; orbit incomplete. Sometimes complete in ________________ P. hypomelanus cagayanus (Mearns).

149

PTEROPUS

b'. Head and body, more than 250 millimeters; tibia, 90; skull, greatest length, 76 ............................ P. vampyrus lanensis (Mearns) (p. 153). b". Head and body, less than 250 millimeters. C'. 'Mantle varying from dark maroon-chestnut to cinnamon-rufous; prevailing color of back seal brown; forearm, 128 to 142.5 millimeters ...................... ..... P. hyp01nelanus t01nesi Peters (p. 160). 18

120'

122

Ic4

126

Balan Is

PHILIPPINE ISLANDS

® Pteropus ® Pteropus © Pteropus © Pteropus

. ® Pteropus ® Ptel'opus

@ pteropus

® pteropus

<D pteropus Q) pteropus

h. cagayanus h. tomesi leucopterus mearnsi speciosus v. lanensis tablasi balutus mimus pumilis

< o·

122

FIG. 6.

124'

126

Distribution of PteT(Yj)UB in the Philippines.

0'. Mantle averaging brighter, rich golden ochraceous-buff; back more tinged with Prout's bx:own; forearm, 135 to 141.5 millimeters. P. hypomelanus cagayanu,s (Mearns) (p. 158). c '. Forearm, 123 miHimetel1s; skull, 53.4; mandible, 46.5; foot, 41. P. mearnsi Hollister (p. 162).

150

PHILIPPINE LAND MAMMALS

c '. F orearm , 109 millimeters ; skull, 49.8; mandible, 39.2. P. purnilus Miller (p. 163). 0". F orearm, 120 millimeters; skull, 57; similar to P. pumilu s but skull larger; feet and claws larger; color blackish sprinkled w ith silver on back ______________ _____ _________________ P. sp eciosus Andersen (p. 164). c·. Similar to P. speciosus but having back varying shades of brown; forearm, 127.5 millimeters ____ __ ______ ____ P. mimus Andersen (p. 166). c '. Lengt h of head and body, 190 millimeters; forearm, 117; foot, 35; lighter in color than P. speciosus, larger than P . pumilus. P. balutus Hollister (p. 168). b a. Length of head and body, less than 150 millimeters; forearm, 92. P. tablasi sp. nov. (p. 169).

Andersen has divided the genus Pteropus into eighteen groups, four of which are represented in the Philippines. Andersen does not recognize the genus Desmalopex of Miller, but places the type species under the pselaphon group; nor does he recognize certain subgenera of Matschie. Andersen has undoubtedly had a greater amount of material of Pteropus than any other reviewer of the group, and his treatment seems to be the last word in a study of this sort. The four groups represented in the Philippines are as followf! : The hypomelanus group, containing P. mimus Andersen, P. 'pumilus Miller, P. speciosus Andersen, P. mearnsi Hollister, P. balutus Hollister, P. hypomelanus cagayanus (Mearns), and P. hypomelanus tomesi Peters. The pselaphon group, containing P. leucopterus Temminck. The temmincki group, containing P. tablasi sp. nov. The vampyrus group, containing P. vampyrus lttnensis (Mearns). It is probable that other species are represented in Luzon. I examined many skulls which have been collected from houses of the Igorot peoples of Bontoc and Ifugao where they serve as ornaments. In the lot were certain skulls which seem to belong to as yet undescribed species or at least to species not recorded from the Philippines. It is futile to record the characters of these skulls without data on the external character of the animals from which they came. PTEROPUS LEUCOPTERUS Temminck Pteropus leucopterus TEMMINCK, Esquiss. Zool. Sur cote de Guine (1853) 60 (Philippines); WAGNER, Schreber's Saug., Suppl. 5 (1853-1855) 599; PETERS, Monatsh. Akad. B€rlin (1867) 323; FITZINGER, Sitzungsb. Akad. Wien 60 Abth. 1 (1870) 433 (Philippines); DOBSON, Cat. Chirop. Brit. Mus. (1878) 32, pI. 4, fig. 1 (skull and teeth) (Philippines); TROUESSART, Rev. & Mag. Zool. III 6 (1879) 203; Cat. Mamm. 1 (1897) 78; JENTINK, Cat. Ost. Mamm. (1887) 253 (skull); Cat. Syst. Mamm. (1888) 140; ELERA,

PTEROPUS

151

Cat. Syst. Faun. de Filip. 1 (1895) 5 (Cagayan, Luzon); Contrib. a la Fauna Filipina (1915) 60 pI. (adjoining, unnumbered) ; SANCHEZ, An. Soc. Espanola Hist. Nat. 29 (1900-1901) 241,275,288; ANDERSEN, Ann. & Mag. Nat. Hist. VIn 3 (1909) 213; Catal. Chirop. Brit. Mus. 2d ed. 1 (1912) 311 fig. 16; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 9; Proe. U. S. Nat. Mus. 46 (1913) 3~5. Spectr1~m let~copterum GRAY, Cat. Monk. Lemurs and Fruit-eat. Bats (1870) 102 (Philippines). Pteropus chinensis GRAY, Cat. Monk. Lemurs and Fruit-eat. Bats (1870) 111 (China?) (Very probably Luzon.) Pteropus (Spectrum) leucopte't路us MATSCHIE, Megachirop. (1899) 26; TROUESSART, Cat. Mamm. Suppl. (1904) 53. Desmalopex leucopterus MILLER, Bull. U. S. Nat. Mus. 57 (1907) 60.

Description of the species.-Snout broadened, eyes slightly upturned; body heavily furred; interfemoral membrane narrow; ears short, subtriangular, half exposed; fur long, dense, semierect on back; tibia thickly furred above nearly to ankle; longish hairs on back, 16 to 19 millimeters; long mantle hairs, Z3 to 26; hairs on belly, 17 to 20. Skull and teeth.-Rostrum sholZt and stout; width of rostrum externally across alveolar borders of first upper premolars subequal to diameter of orbits; front of orbit vertically above back of last premolar or the interspace following; postorbital processes of frontalIS long, fusing with zygomatic process to completely encircle orbit; coronoid process sloping rather than vertical; angular process well pronounced; condyles distinctly above alveolar line. Upper incisors subequal, distinctly larger than in other Philippine species of Pteropus; crown in cross section nearly onethird that of canine; cingulum produced into a shelf posteriorly; upper canine without secondary cusp, moderately large; lower canine normal, curved; first upper premolar less reduced than usual, equal to or slightly larger than lower first incisor, the crown differentiated from shaft; second upper premolar with well-developed posterior basal ledge as is true of third premolar and first molar; anterointernal base of last premolar forming a projecting ledge; similar ledges, but narrower, developed in preceding premolar and following molar; lower incisors very unequal, the second five to six times the bulk of first; ledges on last two lower premolars and first lower molar; a shallow transverse depression in margin of inner longitUdinal ridge of first and second lower molars, subdividing the ridge into two incompletely differentiated rounded tubercles; a similar but less developed depression faintly indicated in outer ridge of first and

152

PHILIPPINE LAND MAMMALS

second lower molars; last lower molar rather larger than usual, subequal to second lower incisor; first premolar not deciduous. Colo?', faded.-Back buffy drab, underparts dull drab; head and mantle pale drab. Measurements of Pte'ropus T..eucopterus Temnninck. Adult female in British Museum. mm.

Forearm First finger: Total length Metacarpal First phalanx Second finger: Metacarpal First pllalanx Second and third phalanx Third finger: Metacarpal First phalanx Second phalanx Fourth finger: Metacarpal First phalanx Second phalanx Fifth finger: Metacarpal First phalanx Second phalanx Tibia Foot and claws Calcar Skull: Total length Orbit to tip of nasals Zygomatic width Width of brain case Width across first upper molars Postorbital constriction Interorbital constriction Orbital diameter Mandible length Coronoid height Upper teeth, including canine Lower teeth, including canine

142.5

Type of P. chinensis. from Luzon. Mm.

?I36

61 15 28.5

63 16.5 30.5

67.5 16.5 17

17 17

97 79

96 103

57.5 61

97 42 45 47 18

21 38 25.8

19.5 9

9.8 14.2 52' 24.5 24 27

102.5 42 48 ?66

21.8 37 24.5 18.5

9.2 10.8 14.2

51.5 23.8 26.7

Remarks.-The measurements here giv-en are taken from Andersen. * The first set are from an adult female in the British

* Cat.

Chirop. 314.

PTEROPUS

153 I

Museum. Those in the second column are those of Gray's Pteropus chinensis, which, according to Andersen, came from Luzon. He states: The type, an unregistered skin and skull, came to the Museum from Robert Fortune, who, in 1843-45, travelled in the northern provinces of China as a collector for the Horticultural Society of London; hence it was, very naturally, believed by Gray to be from "China." But the fact was overlooked that Fortune also made an excursion to Luzon (January to early in March, 184'5; see his 'Three Years' Wanderings in the N orthern Provinces of China,' pp. 332-345, 1847); when to this is added that no species of Pteropus is known to occur in China, and that the type of Pt. chinensis differs in no noteworthy character from that of Pt. leucopterus, there can be no reasonable doubt that it was obtained by Fortune during his stay in Luzon.

The type is in the Leyden Museum. PTEROPUS VAMPYRUS (Linnll!DS)

Vespertilio vampyrus LINNJEus, Syst. Nat. 10 ed. 1 (1758) 31 (part.).

This species, according to Andersen's Catalog; has several known races which differ only in size and color. The single Philippine form is one of the two smaller forms of the group. The Javanese species reaches a forearm measurement of 220 millimeters. The average measurement of Philippine forms is usually less than 200 millimeters. The wing spread of fully adult specimens averages about 1,400 millimeters. PTEROPUS VAMPYRUS LANENSIS (Mearns)

Pteropus edulis (non Geoffroy) FITZINGER, Sitzb. Akad. Wi en 42 (1860) 389 (Manila); PETERS, Monatsb. Akad. Berlin (1861) 707 (Samar); (1867) 324; ZELEBOR, Reise Novara, Zool. 1 Saug. (1869) 10 (Manila); DOBSON, Cat. Chirop. Brit. Mus. (1878) 50 (Samar and Dinagat); GUNTHER, Proc. Zool. Soc. London (1879) 74 (Dinagat, and Rasol Island, near Surigao) ; JENTINK, Cat. Ost. Mamm. (1887) 259 (part.) (Manila); Cat. Syst. Mamm. (1888) 145; STEERE, List Birds & Mamm. Steere Exp. Philip. (1890) 28 (Leyte, Panay, Negros) ; ELERA, Cat. Sisto Faun. de Filip. 1 (1895) 5 (part.) (Samar, Dinagat, Leyte, Cebu); ELLIOT, Field. Columbo Mus. Publ. 2, Zool. ser. 1 (1896) 76 (Leyte); TROUESSART, Cat. Mamm. 1 (1897) 80 (Samar); SANCHEZ, An. Soc. Espanola Hist. Nat. 29 (1900-1901) 240, 275, 288 (Luzon, Mindanao, Samar). Pteropus pluto (non Temminck) PETERS, Monatsb. Akad. Berlin (1861) 707 (Samar). Pteropus jubatus (non Eschscholtz) GERRARD, Cat. Bones Mamm. Brit. Mus. (1862) 56. Pteropus /unereus (non Temminck) MARCHI, Atti. Soc. Ital. Sci. Nat. 15 (1872-73) 515.

Pteropus vampyrus THOMAS, Trans. Zool. Soc. London 1 4 (1898) 383 (Catanduanes, southern Luzon, Samar).

154

PHILIPPINE LAND MAMMALS

P teropus celamo (non Hermann) MATSCHIEl, Megachir . (1889 ) 15 (Luzon , Samar, Mindanao); TROUElSSART, Cat. Mamm., Supp1. (1904) 50; ELLIOT, Cat. Mamm. Field. Col. Mus. (1907) 490 ( Leyte) . Pte1'opus lanensis MEARNS, Proc. U. S. Nat. Mu s. 28 (1905 ) 432 (Pan t.ar, Mindanao); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 58 . .. P teropus vamp yrus lanensis ANDERSEN, Ann. & Mag. Nat. Hist. VIII 2 (1908) 368; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 10; Proc. U. S. Nat. Mus. 46 (1913) 305, (Batac, Ilocos Norte, and S an F ernando de Union, Luzon; Panay; Leyte; Palawan; Mindanao ; Cabo Island, Mindanao).

Description of the species.-(From No. 646, E. H. Taylor collect ion; collected at Kraan Point, Cotabato coast, April 8, 1923, by E dward H. Taylor.) Large; ears long, rather pointed anter ior ly, curving to a narrow, somewhat blunt point, posterior to which a slight emargination, and then curving to notch at base; neck with heavy growth of elongate woolly fur above (about 15 millimeter'S long) continuing to shoulders; fur about 20 millimeters on sides of neck and on foreneck, longest hairs 23 to 25 millimeters; pelage of back short, stiff, rarely exceeding 5 millimeters in length; posterior rump with woolly fur, about 10 to 12 millimeters long ; fur on belly 10 millimeter.s, on sides under wings 10 to 13 millimeters in length; forearm above nearly naked (only a scattering of fine hairs) ; membranes naked above save in the interfemoral region, the proximal part of which is distinctly furred, and a fringe on the wing edge from attachment on foot to tip of phalanges of fifth finger; below, a heavy gr owth of fur on the antebrachial membrane; a heavy growth on the sides of-forearm continuing four-fifths of distance to distal end; interfemoral membrane practically naked save at base; wing membranes attach about 28 to 30 millimeters apart on back; glandular neck tufts of males only slightly differentiated, hair somewhat thicker but scarcely stiffer than on females; thumb short compared with that of typical P. v. vampyrus. Skull and teeth.-Postorbital processes reach scarcely more than halfway to arch of zygoma; a slight projection rises from zygoma; postorbital constriction distinctly greater than interorbital constriction; upper incisors unequal, the inner extending beyond and slightly larger than the outer, all (usually) slightly separated; a strong anterior groove in upper canine; groove on inner face obsolete; on posterior face a shallow narrow depression; canines with a prominent heel but no secondary cusp; premolars and molars without any peculiar modifications; lower incisors nearly equally spaced between canines, much smaller

PTEROPUS

155

than upper incisors; outer lower incisor nearly twice the bulk of inner; first lower premolar larger than last molar; lower canines small without grooves; first upper premolar usually deciduous and present usually only in young specimens. Color.-General color blackish brown; wing membranes distinctly blackish; fur on crown black with occasional white hairs; mantle generally black with an admixture of blackish brown to deep seal brown or slightly washed with same color; shoulders and rump deep burnt umber with occasional white hairs; foreneck, sides of neck, and breast, black to black-brown; belly similar, but with scattered light hairs on underside of forearm nearly chestnut. Variation.-The variation of specimens is considerable. The subspecies is separated from Pteropus vampyrus natunce from which it differs only in the coloration of the mantle. Andersen lists several specimens, as follows: Catanduanes. Malâ&#x201A;Ź adult. Back and rump blackish, sprinkled with gray. Underparts blackish; mantle pale Vandyck-brown approaching russet, heavily mixed with blackish, like foreneck; crown and sides of head similar to back; temporal region slightly washed with Vandyck-brown. Cebu. Male. Similar to above. (One specimen.) A second specimen from same locality, also adult male. The back suffused with pale Vandyckbrown approaching russet; breast and belly slightly tinged with dark Vandyck-brown. Mantle blackish and darker than back, sides of neck and foreneck similar; head blackish, somewhat tinged with Vandyck-brown on sides. Panay. Female. Very similar to second Cebu specimen. Leyte. Female. Back and rump almost uniform Mars brown; center of breast blackish shading to dark chocolate on sides of breast, and Vandyck-brown on belly. Mantle walnut brown, shading to dark Vandyckbrown on sides of neck, to dark chocolate on foreneck and throat; :!rown similar to mantle but darker. Catanduanes. Male. Back seal brown, thinly sprinkled with light grayish. Underparts almost uniform blackish seal brown from chin to inierfemoral with faint tinge of dark chocolate on foreneck. Mantle rich orange ochraceous buff shading to ochraceous-rufous on sides of neck. Crown Vandyck-brown.

Most of the specimens collected by me are of the melanistic form similar to the first listed above. However, in two specimens from Kraan Point and in one from the interior of Cotabato the mantle and body are similar to the second Catanduanes specimen. The color of the mantle in the three specimens differs somewhat but the orange-ochraceous-buff color ceases abruptly on the shoulder at a.point about one centimeter back of anterior wing attachment.

W Forearm First fi To M Fir Second M Fir, Se Third fi Me Fir Se Fourth Me' Fir Se Fifth fi Me Fir Se Depth

Ear:

II -

134 126 135 130 137 . 5 62 58 66 55 63 64 60 58 58 64 g7.{i --------c- --.---.--- --""---." •• ------ ••

:!4

14.2 64 67

-- ... --

...... ---

126 65 57

134 82 79

120 75 75

127 86 84.5

121 74 74

127 60 60 2{i

122 78.2 79 135 86 82 120 79.5 76 124 77 73 123 82 78 121 78.5 75

97 151

18 55

122 55 60

123 90 145 139 102 151 123 94 138 125 96 131 130 98 150

123.2 95 148

125 92 148

131. 6 106.5 149

124.1> 91. b 134

126 54 66 -,. -- ...- ... - . -

95 23 24 106 28 25 95 25 23 94 27 22

96 27 23

92 23 21

86 19 67 77 19 58

74 17 57

J02 26 22

93 22 18

76 17 59

82 20 62

102.5 28.6 22.5

92 20 18

79 18 62

75 19 56

----------

----~-----

83 , 5 19.5 • 45. 6

33 22 190

38 23.5 201

638. female.

37 20 192

101, male.

37 19 186

100, male.

74.5 17 • 40

37 22 190

646, female.

201

191

----------

645. female.

35 ......21 2'05

640, ,female.

Bureau of Science, No. 148, female.

-------------------

35 21

639, female.

E. H. Taylor collection No. and sex.

36.5 22 18'5

Minimum. Maximum.

From Andersen.

[Measurements in millimeters.]

Measurements of Pteropus vampyrus lanensis (M.earns).

r.n.

~ ~

t::I

t"'

Z t<.l

'"d '"d

...... t"' ......

'"d

;:q

O':l

......

28 . 8 32 . 6 6 .2

Pper teeth, canin,e s to molars ____ ______ _ Lower tooth row _ ____ _________ ________ _

Width across upper incisors __

59 . 5 27 .2

16 . 5 62.8 29.7 30 . 8 34 . 2

21. 5 9.5 11

27 . 5 46

27 .2

78 . 3 39

61 30

66 29

73 . 8 36 . 5 25 25 . 8 39 20 . 5 7.5 10 15

Mandible length __ _ Mandible, coronoid height. _____________ _

Palate length __ _ Front o. orbit to nasal tip __ ____ _______ _ _ Greatest width of brain casl' __ __ __ ______ _ Zygomatic width __ ~ idth across first molars p.xternally ______ _ Postorbital constriction __ _ Interorbital constriction ______ __________ _ Orbital diameter ____________ ___ ________ _

Skull: Total length _____ _________ ____ ________ _

Tibia __ Foot and claws ______ _ Calcar __ _________ ___ ____ _

â&#x20AC;˘ Without claw.

87 56 21

58 28

11. 5

10 15 . 5 61 28 30 34 6.8

10 . 5 Hi 67 26 29 32 6.2

10 , 3 15.5 60 31 30 34 7

26 29 33

9 .2 15 .2

15 . 8 61

9 11

24.6 27 . 5 43 . 5 20.5

58 26 . 6

~~~~~~~~~llJ - :~.5 I _ ____ _ 30 ________ __ 6. 2

74 38 . 5 24 . 5 26 41 19.1 8.5 9. 3 14 . 9 58 26

58 25

55 28

54 30

74 38 . 8 24 . 6

1~2

42 19 . 6

51 25

80 39 24 25 41 22 . 3

74 37 23.2 25 40 20

76 39 25 26 . 6 42 . 2 20

92 53 23

-:]

.......

c::

158

PHILIPPINE LAND MAMMALS

The specimens listed all come within the maximum-minimum limits of the species as here recorded from Andersen. The specimens measured are from the following localities: Kraan Point, Cotabato: 638, 639, 640, 645, 646, numbers from my own collection; Port Lebak, Zamboanga Province, Mindanao, No. 148 Bureau of Science P. 1. collection, collected by E. A. Mearns; Central Cotabato 100 and 10l. Remarks.-The type of this subspecies (given specific status by Mearns) was collected at Pantar, near Lake Lanao, Lanao Province, Mindanao, September 7, 1903, by Edgar A. Mearns. He states: "Colonies of these large bats surrounded Lake Lanao, Mindanao, which occupies a basin 2,000 feet above the sea at water level of the lake, whence many of them regularly passed over our camp at Pantar, 5 miles distant, on the Agus River, which drains Lake Lanao." The specimens in my collection came from Kraan Point, Cotabato, where they were found living with Acerodon iubatus mindanensis (see remarks under that species). PTEROPUS BYPOMELANUS Temminck

Pte1"OpUS hypomela,ruus TEMMINCK, IEsq. Zool. 1853 61 (part.) .

(Ternate)

This species has been divided into eleven SUbspecies, two of which occur in Philippine territory. The first, Pteropus hypomelanus tomesi Peters, has been reported from Sibutu Island only, by Andersen; the second is widely distributed in the Philippines. There is practically no difference in size; they can be separated only on color markings. K ey to t he Philippine subspecies of

PterOp~t8

hypomelanu8 Temminck.

at. Mantle varying from dark maroon-chestnut to cinnamon-rufous; prevailing color of back seal brown; forearm, 128 to 142.5 millimeters. P. h. tomesi Peters (p. 160). a ~ . Mantle averaging brighter, rich golden ochraceous-buff; back more tinged with Prout's brown; forearm, 135 to 141.5 millimeters. P. h. cagayanus Mearns (p. 158). PTEROPUS HYPOMELANUS CAGAYANUS (Mearns)

Pteropus (sp.) WATERHOUSE, Pl'OC. Zool. SOC. (1843) 67 (Philippines). Pteropus hypomelanus DOBSON, Cat. Chil'op. Brit. Mus. (1878) 58 (Dinagat); TROUESSART, Rev. & Mag. Zool. III 6 (1879) 202 (S. E. Philippines); Cat. Mamm. 1 (1897) 82 (Dinagat.); GUNTHER, Proc. Zool. Soc. London (1879) 74 (Dinagat and Surigao); STEERE, List. Birds and Mammals Steere Expedition (1890) 28 (Guimaras, Panay, Leyte) ; ELERA, Cat. Sisto Fauna de Filip. 1 (1895) 5 (part.) (Luzon, Dinagat, Mindanao); ELL!oT, Field. Co1. Mus. Publ. 2, Zool. Ser. 1 (1896) 76 (Panay, Guimaras); Cat. Mamm. Field Col.

PTEROPUS

159

Mus. (1907) 491 (Panay); SANCHEZ, An. Soc. Espanola Hist. Nat. 29 (1900-1901) 241, 275, 288 (Luzon, Leyte, Panay, Guimaras). Pteropus (Spectrum) hypomela,nus part. MATSCHIE, Megachirop. (1899) 23, 24 (Cuyo, Mindanao); TROUESSART, Cat. Mamm. Suppl. (1904) 52. Ptet·opus cagayanus MEARNS, Proc. U. S. Nat. Mus. 28 (1905) 433 (Cagayan Sulu); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 58; HOLLISTER, Philip. Journ. Sci. § D 7 (1912) 9. Pteropus hypomelanus hypomelanus HOLLISTER, Philip. J ourn. Sci. § D 7 (1912) 9. Pteropus hYP()1nelanus cagayanus ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912') 121-125 (Luzon, Dinagat, Surigao, Cagayan S'ulu) ; HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 305 (Luzon, Ilocos Norte, Cuyo, Panay, Cagayan Sulu).

Description of the species.-(From No. 386, E. H. Taylor collection; collected near Culasi, Panay, by R. C. McGregor.) Young adult. Of medium size; head especially doglike; ears short, the anterior edge first strongly curved, then continuing to summit, very slightly curved; posterior to summit the edge forms a straight line for nearly half length of ear, then curves suddenly; hair on ear long, scattered on front side, continuing to summit; posteriorly, the upper half of ear practically naked save near posrt erior edge where there are numerous hairs near border; hair on forehead short (5 to 7 millimeters), dense; hair of mantle medially longer (14 to 16 millimeters), on sides of neck even longer (16 to 19 millimeters) ; hair of mantle generally woolly or wavy; hair on back short (6 to 8 millimeters), straight and stiffened somewhat; on rump soft and slightly wavy; fur on foreneck 16 to 20 millimeters, on breast and abdomen 9 to 12 millimeters. Wing membranes fasten close to median part of back (in described specimen, dry skin apparently torn apart) naked above save on lower part of forearm, and a slight growth between tip of fifth finger and attachment on toe; interfemoral membrane heavily haired save in posterior region near calcar; below interfemoral membrane nearly naked; antebrachial membrane with a sparse growth of woolly hair; a similar growth about elbow and on membrane outside of forearm; scattered hairs on feet; calcar moderately developed; wings attach on base of :first phalanx of second toe. Color.-Head dark brown on sides, growing almost buff on the crown, but slightly darker chestnut in front and between ears; mantle above mixed buff, tawny, cinnamon-rufous, and chestnut, the shades changing in different lights; back dark seal brown to black-brown intermingled with numerous white hairs and chestnut hairs; foreneck and sides of neck dark black-brown

160

PHILIPPINE LAND MAMMALS

growing lighter posteriorly; breast and abdomen with a light golden area gradually becoming chestnut buff on edges of area; sides under wings blackish brown like foreneck with some white hairs intermingled; a lighter indistinct line from light area on breast over shoulder to light area on mantle. The col.or of the type is described as follows: "Head blackish brown all round but slightly paler on crown, with a few whitish hairs intermixed above; upper side of neck and shoulders wood brown, becoming tinged with tawny posteriorly, where it ends abruptly in a straight transverse line; remainder of upperparts grizzled blackish. Underside of body russet brown, becoming blackish brown on the sides; feet and wing membranes blackish." (Mearns.) Skull and t~eth.-(Type specimens.) Older specimens have the postorbital and jugal processes united, completing the osseous orbit of the eye; save for this difference and the possession of less robust teeth, the skull resembles pteropus hypomelanus. Remarks.-My specimens are somewhat immature as judged by the joints of the digits. The skulls are wanting. * The type and two paratypes, now in the United States National Museum, were collected on Cagayan Sulu Island near the west side of the Sulu Sea, north of Borneo, by Edgar A. Mearns, February 25, 1904. Mearns t remarks: "The skull and teeth of P. h. cagayanus are decidedly smaller than in Steere's specimen from Panay, which he called 'Pteropus hypomelas'." Hollister t lists twenty-one specimens from the following locali ties: Luzon, Ilocos Norte, 7 specimens (Mearns); Cuyo, 3; Panay, 8 (Steere); Cagayan Sulu, 3 including the type (Mearns) . PTEROPUS HYPOMELANUS TOMESI Peters

Pteropus hypomelanus tomesi PETERS, Monatb. Akad. Berl. (1867) 626; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1913) 119 (Sibutu) .

Description of the species.-This form differs from the preceding SUbspecies only in color. The measurements are practically identical. Practically no differences obtain in the character of the teeth.

* The skulls of my two specimens were loaned to a student instructor in the University of the Philippines and I do not have them at hand at the present t~me. t Proc. U. S. Nat. Mus. 46 (1913) 305. :j: Loc. cit.

161

PTEROPUS Measurements of Pteropus hypomelanus cagayanus (Mea?·ns). [Measurements in millimeters.]

United E. H. Taylor collection. States Na· tional Mu. I- - ----,seum, type, No. 385, No. 386, male. female. female. Head and body................. ......................... 230 Ear: 21 Height from meatus ...... ..................... " ' ' ' ' 17 . 5 Height from crown .......... ....................... . 16 Width ........... .. ...... .......................... . 138 Forearm...................... ......................... . First finger: 53 With claw ...•........... ...................•....... MetacarpaL •................................................. Second finger: Metacarpal .............. .................................... . First phalanx .......... .. . ................................. .. Second phalanx and claw...... . Third finger:

212 -------- -23 18 17 125

20 17 16 136

54 13

56 14

62 16 15

70 17 15

84 58 78

95 65 88

78 49 45

84 . 2 53 52.5

84 35 35 58 15 42

95 40 . 5 40 58 13 41

Fourth finger:

Second phalanx .. .•........•......•..•.................... Fifth finger: MetacarpaL ..........................•....................... First phalanx ..... ........................ """ ............•. Second phalanx ... ...............•.. __ ................. " " " " Tibia............... ... ... .............................. 6;1 15 Calcar ..........................•..................... 45 !~:;I~Drl claws . ........ ~ ............................. '"

Greatest length. .. ......... ......... . ............ . Basal length .......................... .. Basilarlength. . . ............................. ...... Median palatal length. ................ ............... Breadth of palate between front molars .............. . Zygomatj~hreadth ............................... Least interorbital breadth ...................... .. Breadth between tips of postorbital processes ........ .. Greatest breadth 01 brain case above zygnmata .......... , Greatest depth or brain case ....................... . Mandible....................................... .. Maxillary tooth row ............................... . Mandibular tooth row...... .............. . ..... ..

63 61. 5 59 . 5 37 11. 3 35 8 .2 26 . 5 22 . 5 18 52 24 27.5

~~~ :~~:~: ~. ~::~: ::~ ~: I

................... .1

· · . · . -1-·····]. ·

~~~~~~~~~~ ~:~~::~~:: I· .... · .... ,...... · .. ·

Skull lost.

Color.-Ordinary phase: Breast a pale shade of hazel or cinnamon-rufous, generally washed with orange-buff; back a,.nd rump varying from blackish seal brown to pale seal brown, slightly 259463--11

162

PHILIPPINE LAND MAMMALS

or moderately, but never heavily, sprinkled with silvery whitish hairs; breast or at least center of breast pale hazel or cinnamonrufous, as a rule more or less conspicuously washed with orangebuff; on sides of breast, flanks, and belly the color gradually, sometimes rather abruptly, darkens to seal brown or blackish seal brown with or without a slight admixture of grayish hairs; mantle varying from dark maroon-chestnut to chestnut-cinnamon-rufous or chestnut-hazel, becoming darker on sides of neck, and dark chestnut or seal brown on foreneck; crown similar to mantle or rather brighter or somewhat clouded with dark brown, indistinctly sprinkled with grayish or buffy hairs, passing through a darker shade on sides of head, into blackish seal brown on throat. Some specimens show a preponderance of silvery, grayish white hairs washed with buffy, and so change the general effect of the color to buffy grayish white, sprinkled with blackish; occasionally a specimen 路 has the whole back suffused with russet, darkest on front of back, gradually merging into a paler shade of golden ochraceous on rump, and the color of the breast and belly is rather brighter, more washed with ochraceous than usual. Remarks.-Known from small islands off the northeast coast of Borneo and the Bornean coast in the same region. The only Philippine record is Sibutu, based on two specimens collected by A. Everett and reported by Andersen. l'TEROPUS MEARNS! Hollister

Pteropus mearnsi HOLLISTER, Proc. BioI. Soc. Washington 112; Proc. U. S. Nat. Mus. 46 (1913) 305.

(1913)

Description of the species.-A member of the Pteropus hypomelanus group; coloration similar to P. hypomelanus cagayanus, but the size is considerably less and the skull is smaller. Color.-Head and muzzle mummy brown, lighter on crown; mantle chestnut; back and rump brownish black with wash of rusty; breast Hessian brown, mixed with blackish; flanks and lower belly dark chestnut-brown with much blackish; center of belly lighter, golden ochraceous. Young with lighter mantle. Skull and teeth.-Skull intermediate in size between skulls of Pteropus speciosus .a nd P. h. cagayanus; in general the skull is like that of cagaya'YIIUS but considerably smaller with a relatively much larger brain case. Teeth essentially the same as in cagayanus.

163

PTEROPUS Measurements of the type of Pteropus mearnsi Holliste?'. rom.

Forearm :Pollex, total length Second finger, total length Third finger: Metacarpal First phalanx Foot Skull: Condylobasal length Zygomatic breadth Breadth braincase at zygomata Front of orbit to end of nasal Orbital diameter Least breadth of rostrum Interorbital breadth Upper tooth row, entire Upper molar-premolar row Mandible Lower tooth row, entire Lower molar-premolar row

123 55 89.5 82.5 61 41 53.4" 34 22.4 19.5 12 10.2 8.5 28 15.4 46.5 27.8 21.5

Remarks.-Except for its smaller size Pteropus mea,'rnsi resembles P. h. ca,ga,ya,nus but is conspicuously different from all other Philippine species; it approximates P. mimus in size, but differs greatly in color. The type and five paratypes, in the United States National Museum, were collected at Isabela, Basilan, and at Zamboanga, Mindanao, January, 1906, by Edgar A. Mearns. PTEROPUS PUMILUS Miller

Pteropus pU1nilus MILLER, Proc. U. S. Nat. Mus. 38 (1911) 394; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 9; Proc. U. S. Nat. Mus. 46 (1912) 305; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 816, 832.

Description of the species.-This fruit bat is similar to Pteropus speciosus, but has smaller skull, feet, and claws. Color.-Body hair brown or broccoli brown, the back irregularly suffused with dull, light buff, the underparts tinged with " ochraceous; mantle and head dull buff, the middle of neck both above and below strongly suffused with light ochraceous; face, chin, and interramia inconspicuously sprinkled with dark brown hairs. SkuU and teeth.-Save for their smaller size the skull and teeth are essentially the same as those of P. speciosus.

164

PHILIPPINE LAND MAMMALS

M easurements.-See under Pteropus mimus. Remar ks.-The type and one paratype, the only specimens known, were collected on Palmas Island, southeast of Mindanao, January 21, 1906, by Edgar A. Mearns. It is a disputed question as to whether Palmas Island is actually Philippine territory; and likewise questionable as to whether the species should be considered a Philippine mammal. PTEROPUS SPECIOSUS Andersen

Pteropus speciosus ANDERSEN, Ann. & Mag. Nat. Hist. VIII 2 (1908) 364 (Malanipa, Sibutu); Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 132; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 9; Proc. U. S. Nat. Mus. 46 (1913) (Zamboanga, Malanipa).

Description of the species.-(From No. 610, adult male, E. H. Taylor collection; collected April, 1923, on Malanipa Island by Taylor and Eickelberg.) One of the smaller members of the P. hypomelanus group; ears relatively long and prominent due to short hair on head, naked save at base; hair on head and about base of ears about 5 to 8 millimeters long; hair of mantle woolly, on back of neck about 10 millimeters long, but on sides the longest hairs reach 20 millimeters; hair very short on foreneck, and very sparse; the ruffs are only slightly developed, the fur being denser but not stiffened or waxy; hair on back between wing membranes straight (10 millimeters), growing woolly on rump and longer (10 to 14 millimeters) ; median abdominal fur 8 to 11 millimeters on sides, below membranes 15 to 20 millimeters long; a well-developed growth of sparse hair on antebrachial membrane below, as well as along the membrane external to forearm and in the region between knee and elbow up to the body; interfemoral membranes with a growth of hair only on proximal portion; the two membranes not continuous; a slight 2'rowth of hair on the forearm near elbow, and the upper surface of the interfemoral membrane sparsely covering nearly half its surface. Color.-Head a very indefinite buffy color mixed with blackish and brownish hairs; darker around the eyes, darker brown low on sides of head, lightest on crown; mantle reddish chestnut to hazel growing darker brown which suddenly changes to ochraceous posteriorly on shoulders; throat blackish; foreneck with a median lighter hazel-buff area passing back, widening considerably and changing into orange-tawny; on neck on either side of light area two well-defined darker areas that join to ruffs which are blackish seal brown; behind these somewhat lighter,

PTEROPUS

165

and then continues dark seal brown to groin along side of abdomen; back and rump very indefinitely blackish with a changeable sheen appearing brownish olive to silvery brown in certain lights; wing membrane blackish. Skull and teeth. *-Upper incisors moderately large, widened at their extremities, the inner of the series somewhat closer to each other than to other teeth; upper canine elongate, very slightly curved, with a more or less definite shallow groove; first upper premolar deciduous; second premolars with a high outer and lower inner cusp divided by a valley; a slight heel notch on outer posterior surface; first two molars similar to the premolar but lower, wider and longer; lower incisors unequal, the two median of the series farther from each other than each is from outer, the median diastema equal to or only slightly less than that between second incisor and canine; inner incisors about one-half bulk of outer; lower canines much less than upper; first lower premolar well developed, the surface showing an elevation on outer side, the tooth equal to last molar; second premolars and first and second molars somewhat similar to corresponding upper teeth. Skull rather slender, postorbital processes very long, narrowly separated from the short process arising from the arch; postorbital distance distinctly less than interorbital distance; anterior edge of orbit directly above front of second molar. Va't 'iation.-The specimens I have ' examined (1 from Zamboanga, 2 from Sibutu, and 18 from Malanipa), agree very well in general measurements, color, and tooth and skull characters. The teeth, particularly the canines, are larger in some specimens; there is distinct variation in the relation of the upper incisors, some of the skulls showing the median pair closely in contact and rather widely separated from the second pair; the groove in the front face of upper canine is obsolescent in several skulls, and distinctly present in others; the depth of the heel notch varies, it is scarcely noticeable in some teeth, distinctly present in others in the second premolar and not infrequently present in the first molar. A young specimen (No. 608, Malanipa) shows as milk dentition a functional curved incisor, a canine, and two small teeth which appear to lie at the surface of the gums and not embedded in bone. The permanent teeth can be seen; the first premolar * From No. 617, same locality and date. state.

The skull of No. 610 is in poor

166

PHILIPPINE LAND MAMMALS

is developed on one side only, on the other (left) side apparently no tooth is developed. This tooth, if normally present, is probably lost at the same time as the milk teeth since I do not find it persistent in the young specimens whose teeth show no wear; in the lower jaw a single pair of milk teeth persist in the specimen, probably the milk canines since they follow immediately behind the permanent canines. Andersen mentions a black phase; none of the specimens I examined show this. Some, however, vary in the general effect of the back, some having many silvery hairs present. The shade of the mantle varies from seal to tawny brown, and the area of ochraceous-buff on abdomen varies. The type has practically the same color as the specimens here recorded from the type locality (Malanipa Island). Andersen gives the following description of a female specimen from Sibutu: Back and rump seal-brown somewhat varied with light gray, particularly . along sides of back along membranes. Underparts seal-brown, washed with dull russet on breast. Mantle nearly Vandyck-brown becoming darker on sides of neck, gradually passing into the dark seal-brown ofl foreneck. Crown and sides of head grizzled light gray and blackish seal-brown, producing a total impression of a dark hair-brown; throat blackish.

The lighter phase from Malanipa is similar to P. h. tomesi though the color of the mantle is more like P. h. cagayanus; the dark phase is similar to P. h. geminorum. PTEROPUS MIMUS

Andersen

Pte?'opus hypomelanus (nec Temm.) DOBSON, Cat. Chirop. Brit. Mus. (1878) 57; MEYER, Abh. Mus. Dresden 7 (1899) 6 (part.). Pteropus mimus ANDERSEN, Ann. & Mag. Nat. Hist. VIII 2 (1908) 364 (South Celebes); Cat Chirop. Brit. Mus. 2d ed. 1 (1912) 133 (Luzon); HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 335.

Description of the species.-Closely related to Pteropus speciosus but differs in having the back a Vandyke brown; the rump similar, but washed with Mars brown; breast and belly pale golden ochraceous tinged with orange, heavily clouded with Mars brown on breast and crissum; purer in tinge on belly; flanks dark Prout's brown, many hairs with tawny tips; mantle rich ochraceous-buff strongly tinged with orange or a color between cinnamon and russet; sides of neck and foreneck nearly tawny; crown . similar to mantle; forehead and sides of head brownish mixed with buffy; throat seal brown.

167

PTEROPUS

Skull and teeth.-Similar in all respects to Pteropus speciosus. Measu?'ements of Pter opus mi'1'l1lU8 And.ersen and Pteropus pum1.lus Miller.

[ --

[Measurements in millimeters.]

Pteropus mimlts. Female. type, 째

Female. paratype. O

Pteropus pltmilus. Male. type.

F emale. para type.'

Ear: HeighL __ _____ - - -- - -- ---- - - -- -- - - - - - - - - - - - - - - ---- Width ____________ __-- ___ -- - -- --- - ---- - - - - - --- - -- - Forearm __ ___ ____ _. _ - - -- - - - - --- - - - -- - - - -- -- - - - - - - - - - -- First finger: With claw _____ ______________________ . --55 Metacarpal ____ __ _______________________ _ 13 First phalanx ___ __ ____ .. --- - - ----- - -- --- -Second finger: Metacarpal ______ ________________ -------

25 17 . 5 127.5

109

44 11

22 . 5

29 62 15 13

First phalanL ________ ______ - - - - - -- -- - ---Second phalanlL __ -"_____ _______ ----------

12 . 5 12

Thir~!::::;paL _________________________ ~~

89 61. 5 94

First phalanx __ ___ _____________________ - ~ _I '. Second phalanx ____ __ ___ , ___________ - __ ---

58 . 5

Fourth fi.nger: ,'.' II Metacarpal ____ _________________ - _- _____ . 80 First phalanx_ ___ __ ___ _ __ __ __ __ __________ _ 48.5 Second phalanx __ ________ ___________ __________ __ ___ _ Fifth finger: Metacarpal __ ________________________ ____ 85 First phalanx_____ __ __ ______ ___ ___________ 36 Second phalanx ___ __ ____ ____________ ____ _____ ___ ___ _ Interf(>moral width. median ________________________ ___ __ _ Tibia ________ .. . ___ _____ ____________ ______________ ______ Foot and claws__ __ ___ __ ___________ __ __ ____ __ _ 38 . 5

10 70 . 5 52 . I) 73

85 50 . 5 53

70 . 5

42.5 41. 5

91 36 40

75 30

58 36

SkuU:

Totallength ______ ___ __ _____________ _____ _ ___ ____ __ _ Orbit to na9aI8______ ___________________ __ 19 Zygomatic width _________ ___ ___ ______ ____ _____ ___ ___ Width of brain case _______ _____ ___ ____ ________ _____ _ Interorbital constriction _ _ _____ ________ _ __ _ 7.8 Orbital diameter_ __________ __________ ____ _ 12 .2 Width between canines ___ _________ ________ 5.8 Mandible length_______ ____ ____ ______ __ __ _ 45 oronOid heighL____ ________ __ __ __ _______ 20 Upper tooth row with canines_ __ ___ __ ____ __ 20.8 Lower tooth row with canines_ ___ ____ ______ 23 . 7

59 . 8 20 . 7

32 . 2 22 8 .8 12.5 6.8

29 . 6 19 . 6 ___ __ ___ _ _ 7 . 8 ___ __ _____

45 . 2

39 . 2

37 . 5

22 22.7 25

18.2

18 . 2

19 . 8

10 . 7

From Andersen.

Remarks.-Apparently only a single specimen has been reported from the Philippines. This is a specimen collected by A. B. Meyer. There is a possibility that this is a Celebean specimen collected by Meyer with incorrect label.

168

PHILIPPINE LAND MAMMALS PTEROPUS BALUTUS Hollister

Pteropus balutus

HOLLISTER, Proc. BioI. Soc. Washington 26 (1913) 111 (Balut Island, Sarangani group) Proc. U. S. Nat. Mus. 46 (1913) 305.

Description of the species.-Somewhat intermediate in characters between Pteropus speciosus and P. pumilus; coloration of upper parts much as in P. pumilus, but slightly paler; size approaching that of P. speciOstlS but slightly smaller, with noticeably smaller pollex. Color.-Crown, and anterior and posterior edges of mantle pale orange-yellow; center of mantle (nape and shoulde~s) darker, more ochraceous-tawny; general color of back and rump wood brown, a mixture of hairs of auburn and avellaneous; brighter and more reddish on hips; throat buffy brown; breast, flanks, and anal region ochraceous-tawny; middle of belly buffy. Skull and teeth.-The size and general character of the skull seem essentially as in P. speciosus with the rostrum slenderer and the interorbital breadth less. The worn teeth appear to be as in P. speciosus. Measurements of the type of Pteropus balutus HolUster. mm.

Length Alar expanse Forearm Pollex: Total Metacarpal First phalanx Second finger: Metacarpal First phalanx Second and 3rd phalanges Third finger, metacarpal Foot Front of orbit to tip of nasals Least breadth of rostrum Least interorbital breadth

190 855 117.5

47.5 11.3 25 59 13

12 78.5 35 18 9 7

Remarks.-The type, in the United States National Museum, was collected on Balut Island, Sarangani group, south of the eastern part of Mindanao, by Edgar A. Mearns, January 23,1906. Only a single specimen was found; it was taken in a forest clinging to a deciduous tree. It is related to P. speGiostlS, of Malanipa Island, but separated by its lighter color; from P. pumiltlS from Palmas Island it differs in being decidedly larger.

PTEROPUS

169

PTEROPUS TABLASI ap. nov.

Tvpe.-No. 312 E. H. Taylor collection; collected near Odiongan, Tablas Island, January, 1923, by E. H. Taylor. Skin, made up from alcoholic specimen, and skull. Diagnosis; description of the type.-A small species, probably belonging to the Pte.ropus temmincki group, but well differentiated from other members of the group; ears short, rounding anteriorly to the summit, behind which is a distinct emargination leaving ear rather pointed; following emargination ear rounded to base; lower part of ear, both inner and outer, with heavy fur, the latter continuing as scattered long hairs half way to summit; wing membranes attaching on back, at least 20 millimeters apart, and to distal end of metacarpal of the second toe; calcar strong; interfemoral membranes narrow, connected medially where membrane is 4 to 5 millimeters deep; forearm haired from elbow nearly two-thirds of its length, hair growing scant and snort distally; tibia haired completely, hair growing sparse distally and appears only as scattered hairs on foot and toes; interfemoral heavily furred with elongate hair save in region about calcar; alar membrane from point of attachment on foot to tip of fifth finger with a distinct fringe of hair; fur on head, 5 to 6 millimeters; on neck, above, 10 to 12; on shoulders, 10; on back, 8 to 10; on foreneck, 15 to 18; on belly, 12 to 15. Color.-Forehead covered with wavy silvery gray hairs with an admixture of reddish brown hairs which are nearly blackbrown at their bases, the two together giving the effect of gray with a brownish cast; back of head, neck, and shoulders with outer third of fur silvery to buff, basal part of fur blackish brown with an admixture of silver hairs not bicolored; on the back the tip of fur is dull dirty buff, basal fur same as anteriorly; foreneck and breast anteriorly grayish, posteriorly brownish buff, basal half of hairs dark black-brown strongly contrasting; in anal region the basal part slightly lighter, somewhat reddish brown; all fur wavy. Skull and teeth.-Skull slight, delicate, without sagittal crest, the excessively dim temporal crests, scarcely visible, continued posteriorly, about 6 millimeters apart; the postorbital measurement distinctly greater than interorbital measurement; the length of rostrum, orbit to nasal tip, less than one-third length of skull ; posterior part of skull strongly deflected, the alveolar line passing through middle of the supraoccipital; rostrum shows no lateral

170

PHILIPPINE LAND MAMMALS

compression; anterior edge of orbit above anterior portion of first molar. Upper incisors relatively large, the two inner in contact, the outer separated by a very short distance; canines slender, curving back slightly with a deep vertical groove anteriorly; two slight vertical grooves on outer face and two grooves on the posteroint'ernal face; the cingulum forming a heel behind; first premolar a minute terete spicule (on right side only) about 0.17 millimeter wide and 0.7 high, spaced equally from canine and second premolar; latter with dim posterior basal ledge, as is present on following tooth, but more pronounced on inner side; lower incisors unequal, the first: separated from its fellow by a distinct diastema much greater than diameter of the tooth, is only about one-third or one-fourth the bulk of second incisor; lower incisor weak without noticeable grooves, with posterior basal heel; first lower premolar larger than second incisor, about equal to last premolar; other premolars with a distinct posterior, basal ledge, less noticeable in first molar. Measure#"ents of Pte~'opus tablasi sp. nov. mm.

Total length Head Ear: Height Width Forearm First finger: Total length Metacarpal Phalanx and claw Second finger: Metacarpal First phalanx Seoond and third phalanges Third finger: Metacarpal First phalanx Second phalanx Fourth finger: Metacarpal First phalanx Second phalanx Fifth finger: Metacarpal First phalanx Second phalanx

130 48 19 12 91 43 11

33 46.5

13 12 61.5 4'5 57

61 37 35 66

28 27

171

ROUSETTUS Measurements of Pte?'opus tablasi sp. nov.-Continued. mm.

Tibia Foot and claws

40 30

C~u

Skull: Total length Orbit to nasal tip Zygomatic width Orbital diameter Width of brain case Interorbital width Postorbital width . From outer edges of first molars Posterior palate to incisors Upper tooth row, including canines Lower tooth row, including canines Mandible length

Remarks.-The four known species of the may be recognized by the follow ing key: a 1. Head striped. b t. Forearm, 100 to 116 millimeters. b 2. Forearm, 86 to 96 millimeters.

46 14 24 10.1 19.5 6.5 9 13 23.5 18 19.1 35.2 ten~mincki

group

(Bismarck Archipelago.) P. capistratus. (Gilolo and Celebes.) P. pe1路sonatus.

a'. Head not striped. b '. Forearm, 94 to 101 milHmeters; skull, 49 to 51; rostl'um, 15 to 17; mandible, 37 to 39. (Ceram, Amboina, and Timor,) P. temrwincki. b '. Forearm, 92 millimeters; skull, 46; rostrum, 14; mandible, 35. (TabIas Island, Philippines.) ........................................ P. tablasi sp. nov.

It is interesting to note that the closer relationship of the new species obtains with the species most distant in geographical habitat, rather than with the closest. The specimen was obtained from a very small cave near Odiongan, Tablas Island, It was the only specimen in the cave. Genus ROUSETTUS Gray

Rousettus GRAY, London Med. Repos. 15 (1821) 299; MILLER, Bull. U. S, Nat. Mus. 57 (1907) 54, fig. 7; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 16. Cercopterus BURNETT, Quart. Journ. Sci. Lit. and Art 27 (1829) 269. Xantharpyia GRAY, List, Mamm. Brit. Mus. (1843) 37; MATSCHIE, Flederm. des Berliner Mus. fiir Naturk. (1899) 65; BLANFORD, Fauna India Mamm. (1888-91) 261. Eleutherura GRAY, Voyage of the Sulphur, Mamm. 1 (1844) 29.

172

PHILIPPINE LAND MAMMALS

Cynonycteris PETERS, Reise nach Mossambique, Zoo I. I Saugt. (1852) 25; Monatsb. Akad. Wiss. Berlin (1867) 865; DOBSON, Catal. Chirop. Brit. Mus. (1878) 70 (part.). S enonycte?-is GRAY, Cat. Monk. Lemurs and Fruit-€ating Bats (1870) 115. M ynonycteris MATSCHIE, Flederm. des Berliner Mus. fiir Naturk. (1899) 63.

Teeth essentially as in Cynopterus but cheek teeth with cusp ridges, and grooves less strongly developed; upper incisors low, widely spaced, the central pair not higher than the outer teeth, the crown bluntly rounded but distinctly differentiated from shafts; lower incisors with front .surface marked by a groove, which in some species is so deep as to produce distinct emargination of the cutting edge; the skull differs from Cynopterus in the · slightly greater deflection of the occipital region (the alveolar line proj ected backward, passing through middle of occipital condyle) and in the longer rostrum (distance from orbit to nares greater than lachrymal breadth). External characters essentially as in Cynopterus but the muzzle less shortened; second finger with well-developed claw; tail and calcar present. (After Miller.) . 2- 2 1-1 3-3 2- 2 Dental formula: 1. 2- 2' c. 1- 1' pm. 3-3' m. 3-3 = 34. The genus, according to Andersen, contains eleven species, which are found from Africa and southern Asia to Bismarck Archipelago and the Solomon Islands. Two of the forms are referred to subgenera; these are R. (Stenonycteris) lanosus Thomas and R. (Lissonycteris) angolensis Bocage. Only a single species is known from the Philippines. Gray's Rousettus philippinensis is regarded as a synonym of R. amplexicaudatus Geoffroy by Andersen. ROUSETTUS AMPLEXICAUDATUS

(Geoffroy)

Pteropus amplexicaudatus E. GEOFFROY, Ann. Mus. d'Hist. Nat. 15 (1810) 96, pI. 4 (Timor).

Pachysoma amplexioaudatum WATERHOUSE, Proc. Zool. Soc. London (1843) 67 (Philippines).

Pte-ropus (Cynonycteris) amplexicaudatus PETERS, Monatsb. Akad. Berlin (1861) 707 (Luzon, Samar).

Xantharpia amplexicaudatus THOMAS, Trans. Zool. Soc. London 14 (1898) 383 (Luzon); MATSCHIE, Megachirop. (1899) 67.

Cynonycteris amplexicaudatus PETERS, Monatsb. Akad. Berlin (1867) 865 (Philippines); DOBSON, Mon. Asia. Chirop. (1876) 29 and 190 (Philippines); J. ANDERSON, Cat. Mamm. Ind. Mus. (1881) 103 (Philippines); ELLIOT, Field Col. Mus. Publ. Zool. 1 (1896) (Negros); ELERA, Contrib. a la Faun. Filipina (1915) 73 (Jolo).

173

ROUSETTUS

R01A,settus umplexicuuda.tu s ANDERSEN, Ann. & Mag. Nat. Rist. VII 19 (1907) 508; Cat. Chirop. Brit. Mus. 2d ed. (1912) 40 (Manila, Luzon; Benguet, N. Luzon); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 54 (part.); HOLLISTER, Philip. Journ. Sci. § D 7 (1912) 9 (Luzon, Negros, Samar, Guimaras); Proc. U. S. Nat. Mus. 4,6 (1913) 325 (Luzon, Lubang, Negros); STEERE, Birds and Mammals Collected by Stoore Exped. (1890) 20 (Guimaras). liS

122

120

126

Batan Is

PHIUPPINE ISLANDS

® Rousettus amplexiclludatus @ Dobsonia pel"onii

LubenSh W)

0 ...

Busuanga

Bur;.~

'" ~

Siffdya~~'c"o I)Rombion

•

Obl.f/

Cul;on~ •

Calanduanes

~gUqu.

\g :

MINDORO

~A¥iATN~~.

oPANAY Cuyo Oumapan

~A~N

IZ'

. '~ . 0 " ® J() ra

,.,.,"... ,NEGROS

~ .fo2 ~6

<>Cagayan Sulu

120'

FIG. 7.

122

124'

126

Distdbut ion of ROUsettus, Dobscmia., a nd Ptenoch irlts in t h e Philippines.

Eleutheru?'u philippinensis GRAY, Cat. Monk. Lemurs and Fruit-eat. Bats (1870) 119 (Manila). Pteropus philippinensis ELERA, Cat. Sisto Fauna de Filipinas 1 (1895) 6 (Luzon, Samar, Mindanao, Palawan). Rousettus philippinensis MILLER, Bull. U. S. Nat. Mus. 57 (1907) 54.

174

PHILIPPINE LAND MAMMALS

Description of the species.-(From No. 346, E. H. Taylor collection; collected on Polillo Island, July, 1920, by Edward H. Taylor.) Adult female, skin and skull. Muzzle elongate~ doglike; nostrils separated by a deep groove; inner margins moderately proj ecting; inner edges of lips with small papillre; a groove divides naked area on upper lip at tip; lower lip with two triangular_ naked areas separated by a groove; sides of snout with four ill-defined glandular ridges bearing vibrissalike hairs; a distinct tubercle above eye; hair on head extends to a point just anterior to eyes; ear relatively short, the tip rounded, without emargination, nine or ten well-defined ridges on posterior half of ear; a well-defined rounded antitragal lobe; ears naked externally save at base; inner face with scattered, somewhat elongate hairs; attachments of wing membranes on the back separated by about 10 millimeters; attachment on foot to base of digits between first and second toes; interfemoral membrane scant; calcar slender with a small postcalcaneal membrane; tail involved in interfemoral membrane about half its length, fur on head and back moderately dense, somewhat longer in neck region; wing membranes naked above; forearm with growth of fine short bair more than a third of its length; below, membranes with well-developed growth of sparse hair on antebrachial area, and behind elbow on alar membrane, and in region between elbow and femur; claw on second digit well developed; no anal glands. Color.-Head, back, and rump varying from dark olive to sepia; nape of neck wood brown; sides of neck and foreneck wood brown or tawny-olive; lower parts grayish drab to wood brown. Skull and teeth.-Skull somewhat deflected, alveolar line proj ected, passing through base of zyg<?ma; zygomatic arches rather fragile; postorbital processes rather slender; nasals do not reach anteriorly as far as tip of premaxillaries; temporal ridges distinct, save in posterior part of skull. Upper incisors with triangular summits, inner angle of triangle highest; teeth in a semicircular series, first and second on each side closer to each other than the two 1p.edian; median diastema about one-half distance of second incisor from canine; upper canines moderate; cingulum evident on posterior basal edge, and a distinct groove present both anteriorly and posteriorly on tooth; first upper premolar approaching incisors in size; lower incisors much larger than upper, bifid, equally spaced in semicircle between canines; first lower premolar strong, well developed, larger than third molar; cheek teeth strong, some-

ROUSETTUS

175

what squarish. The details of the skull are taken from No. 590, from Cebu. The skull of the described specimen is in a poor state and the teeth are badly broken and worn. M easurem ents of

RO~lsettus

am pl exicaudatus Geoffroy.

[Measurements in millimeters.]

- - - - - - - - ; - - - -- ------;------ -- . , E. H . Taylor collection No .

Totallength ____ __________ __ _________________ _ Head and body ______________________________ _ HeadL ______________________________________ _ Tail ____________ _______ _____________________ _

From Andersen .

346, female.

590, female.

Minimum .

135 121 40 14

133 115 40 18

---- -- -- --

Maximum .

-- - --- - --- - - ------ - ------- -- - - - - - --------14

19 . 7

Ear: Height from notcb ______ ________________ _ width, flattened _______ _________________ _ F~rearm ___ __ ________________ ___________ " ___ , First finger and claw ______ __ _________________ _ Second finger:

---/1

MetacarpaL - - - - -- - - - - -- - -- - - - - -- - -First phalanx ____ - - - - -- -- - - y - _ -- __ - - - Second phalanx and claw __ "'.___________ _

---I}

17 11. 5 89 29

18 .5 10 . 2 87 30

18 . 5 12 . 2 77 25

19 . 7 13 89 . 2 30

36 9 9

37 8 9

31 6 7 5

35 . 2 8. 2 9

46 . 7 30 . 2 39 2

52 35 7 47 . 2

Third finger : MetacarpaL __ -- -- -- ___ -- _-- ------ - --- _--I '; First phalan x ____________________________ _ Second phalanx ____________________ -- ____ _

58 36 46

Fourth fin ger: M etacarpaL _ ____ ____________ ___________ _ First phalanx ____________________________ _ Second phalanx _________ .. _______________ _

54 27 30

53 27 28

44. 7 23 25 . 8

51. 2 26 32

Fifth finger: MetacarpaL ____________________________ _ First phalanlL ___________________________ _ Second phalanx __________________________ _ Tihia ____________ ___________________________ _ Foot and claws ___ ___________________________ _

52 24 27 37 22

52 26 30 37 21

43 21.2 23 . 8 34 . 7 20

51 24 29 .5 38 .8 22 .8

36 . 3

35 11.7 21 14 10 . 2 7.2

39 . 7 13 . 2 24 . 5 16 . 2 11 . 8 8.6

12 . 8 14 27

14 . 2 15 . 8 31

37,

Skull: Totallength ________ _____________________ _ Orbit to nares _______ ________ ____________ _ Zygomatic width _____________ ____________ _

38 11 . 2 20 . 1 Width of brain casB ____________ __________ _ 14 . 6 Width across last molars, upper__ __ __ _ _ ___ __ _ _ _ _ _ ___ _ _ Interorbital breadth____ ____ _ ___ _ _ ___ _ ___ _ _ 8 .2 Width between upper canines __ ___ ____ __ __ _ 4 Orbital diameter ______________ ___ ________ _ 9

Pper teeth, canine and molars _____ _____ __ _ Lower teeth, canine and molars ___ ___ ___ __ _ _ Mandible ________ ____ _____ ___ ____ ___ ___ _ _

12 . 8 14 . 4 29

11.5 22 15 . 5 10 7.3 4 8.5 12 . 8 14 . 2 29

Variation.-The characters are very constant for the most part. Minimum and maximum measurements, taken from Andersen,

176

PHILIPPINE LAND MAMMALS

are included in the table. Most of the Philippine specimens fall within these limits. Rernarks.-I have never found this species in caves. It probably occurs in most of the islands. Other members of the genus, however, are cave-haunting species and feed on fruits and insects. Rousettus leschenaulti is reported as living on marine mollusks. In general appearance R. arnplexicaud:atus resembles species of Eonycteris. The claw of the second digit easily differentiates the genera. Genus DOBSONIA Palmer

Cephalotes GEOFFROY, Ann. Mus. d'Rist. Nat. Paris 15 (1810) 104 (part.); MATSCHIE, Flederm. des Berliner Mus. fiir Naturk. (1899) 85. Hypoderma I. GEOFFROY, DIct. Class, d'Rist. Nat. 14 (1828) 706 (non Latreille) . Hypodermis BLYTH, Cuvier's Anim. King. (1840) 69. Dobsonia PALMER, Proc. BioI. Soc. Washington 12 (1898) 114; THOMAS, Proc. BioI. Soc. Washington 15 (1902) 198; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 63; ANDERSEN, Ann. & Mag. Nat. Rist. VIn 4 (1909) 529 (revision of genus); Cat. Chirop. Brit. Mus. Nat. Rist. 2d ed. 1 (1912) 448.

Differs from Pteropus in having the wing membranes attaching in middle of back instead of on the sides of the body; tail well developed; no claw the first finger. Upper incisors short but well developed, in contact or nearly so, crowns about as wide as long and with distinct cutting edge; lower incisors almost structureless spicules with barely indicated blunt crowns; canines rather short, not peculiar in form, without secondary cusps, those of lower jaw almost in contact; no small upper premolar; small premolar present in lower jaw relatively larger than in Pteropus and not separated from canine or next premolar by a diastema; the other premolars above and below differ from those of Pteropus in greater development of the cusps and more trenchant ridges; small posterior molars as in Pteropus; anterior molars resembling the large premolars in their general characteristics as compared with the corresponding teeth in Pteropus, crown of each with a distinct median longitudinal ridge at middle of furrow; the ridge continuous nearly from end to end in second molar, in first molar it is confined to the region behind the cusps; in some specimens it is obsolete in first molar; skull shorter an-d more heavily built than in Pteropus, but essentially of the same type; distance from orbit to nares about equal to lachrymal width; posterior portion of brain case slightly tubular and occipital region so

177

DOBSONIA

deflected that the alveolar line, projected backward, passes just below the root of the zygoma; premaxillaries very narrow, width scarcely more than that of crown of incisors; inner extremities barely come in contact; audital bullre somewhat wider than in Pteropus but less developed than in Cynopterus . . 1- 1 1- 1 2- 2 2-2 Dental formula: 1. 1- 1' c. 1- 1' pm. 3- 3' m. 3- 3 = 28. Several species of the genus are recognized, of which Dobsonia peronii has been reported from Samar. I

DOBSONIA PERONII

(Geoffroy)

Cephalotes per onii E. GEOFFROY, Ann. Mus. d'Hist. Nat. 15 (1810) 99; Cour's d'Hist. Nat. Mamm. (1828) 29; TROUESSART, Cat. Mamm. 1 (1898-99) 87 (Samar). Pte'ropus paliatus E. GEOFFROY, Ann. Mus. d'Hist. Nat. 15 (1810) 99. Cepha~otes peroni TEMMINCK, Monog. Mamm. 1 (1825) 169. Dobsonia paliata JENTINK, Notes Leyden Mus. 28 (1906-7) 166 (part.) . Hypodermis peronii , HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 11. Dobsonia pe1'onii ANDERSEN, Ann. & Mag. Nat. Hist. YIII 4 (1909) 529; Cat. Chirop. Brit. Mus. Nat. Hist. 2d ed. 1 (1912) 467.

Description of the species.-The characters of the genus. Medium-sized fruit bats; no claw on index finger; tail well developed; two upper and two lower incisors; head and shoulders well clothed with fur; posterior part of body nearly naked. Color.-Fur of head and shoulders blackish. Skull and teeth.-The anterointernal corner of first lower molar marked off as a distinct cusp or ledge; no well-marked anterointernal ledge in first upper molar. Measurements of Dobsonia peronii Geoffroy. Total length, head and body, about Skull, total length Maxillary tooth row, crowns FOJ7earm

mm.

165 4"8.8 20 108.5

51.7 22 117

Remarks.-The species is included on the strength of a record from Samar by Trouessart. * This record, I believe, should be regarded as doubtful. Andersen has broken the species, as formerly understood, into a number of species. The Celebean form is termed Dobsonia exoleta. It is very probable that the Samar record is either an error or refers to D. exoleta or to an unknown species. None of the charts I have examined * Cat. Mamm. tam 'Yiv. quam Fossil. 969463-12

1 (1898-99) 87.

178

PHILIPPINE LAND MAMMALS

shows any island called "Samar" except in the southeastern Philippines. Genus PTENOCHIRUS Peters

Ptenochirus PETERS, Monatsb. Akad. Wiss. Berlin (1861) 707; DOB-' SON, Cat. Chirop. Brit. Mus. (1878) 81 (part.); MATSCHIE, Flederm des Berliner Mus. fur Naturk. (1899) 77; MILLER, Bull. U. S. Nat. Mus: 57 (1907) 51; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 643.

Upper incisors smaller and more pointed tha~ those of Cynopterus, distinctly though narrowly separated from each other; outer tooth of each pair three-fourths to one-half as long as inner; lower incisor smaller than corresponding tooth in Cynopterus, nearly terete (not widened anteroposteriorly) ; canines as in Cynopterus but without secondary cusps; cheek teeth differing from those of Cynopterus in a general widening and flattening of crowns, the outline of which becomes subquadrate in last路 premolar above and below and first lower molar, and the less distinctness of median furrow encroached upon the thickened inner ridge; a claw on second finger. (After Miller.) . . 2-2 1-1 3-3 1-1 Dental formula: 1. 1-1' c.. 1-1' pm. 3-3' m. 2-2 = 28. This genus may be differentiated from Cynopterus by the absence of the inner pair of lower incisors, the reduction of the size of the outer upper incisors, and in the broader, flatter molariform teeth. A single species is known, occurring in Borneo and the Philippines. PTENOCHIRUS JAGORII (Peters) Pachysoma (Ptenochirus) jagorii PE11'ERS, Monatsb. Akad. Berlin (1861) 707 (Daraga, Albay, LNzon, type locality). Ptenochirus jagorii PETERS, Monatsb. Akad. Berlin (1867) 867; ELERA, Gat. Sisto Fauna de Filipinas 1 (1895) 7; Contrib. a la Fauna Filipina (1915) 75 (South Mindanao; Pangasinan, Pampanga, and Nueva Ecija, Luzon); MATSCHIE, Megachir. (1899) 79 (Luzon; Mindoro); TROUESSART, Cat. Mamm., Suppl. (1894) 63; MILLER, Bull. U. 8. Nat. Mus. 57 (1907) 51; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. (1912) 645; fig. 51 (details of skull); HOLLISTER, Philip. Journ. Sci'. 搂 n 7 (1912) 8; Proc. U: S. Nat. Mus. 46 (1913) 334. (Mindanao and Mindoro). Cynopterrus (Ptenochirus) jagorii DOBSON, Monog. As. Chirop. (1876) 28; Cat. Chirop. Brit. Mus. (1878) 87; TROUESSART, Rev. & Mag. ZooI. III 6 (1879) 207; ROBIN, Ann. Sci. Nat. VI ZooI. 12 (1881) 4, pI. 5, figs. 31, 32; TROUESSART, Cat. Mamm. 1 (1897) 86;' MAT~ SCHIE, Megachir. (1899) pI. 12 (animal, skull, and palate ridges). Cynopterus jagori HOFFMANN, Abh. Mus. Dresden, No.3 (1886-87) 24 (S. Mindanao); SANCHEZ, An. Soc. Espanola Hist. Nat. 29 (1900) 276 and 288 (Luzon, Mindanao).

PTENOCHIRUS

179

Description of the species.-(From No. 215, E. H. Taylor collection; collected on Tablas Island, January 18, 1923, by E. H. Taylor.) Adult male. Head doglike; nostrils tubular, separated by a deep groove, tubes joined on median line posteriorly, widely separated anteriorly, as in Cynopterus; upper lip with a naked area extending to nostrils with an indistinct median groove; posterior to nostrils on sides of head a thickened glandular area naked except for very minute hairs and a few scattered vibrissre; eyes large; ears rather small, tip slightly pointed rather than distinctly rounded, with only a very slight emargination . in upper outer margin; antitragal lobe not well developed; wing membranes very ample attaching to sides of body; membranes above with a narrow inner margin covered with fine hair; a small hair growth on lower part of forearm and posterior part of antebrachial membrane; interfemoral membranes furred on inner portion; underside of antebrachial membrane covered with sparse elongate hairs; undersurface of forearm and an area of adjacent membrane with sparse elongate hairs; lower part of interfemoral membranes less hairy than above; calcar strongly developed extending slightly beyond edge of membrane; postca1caneal part of membrane small; wing membrane attaches aiong inner side of outer toe at a point beyond end of metatarsals; tail extends about three-fourths of its length beyond membrane which is reduced to a mere margin in region of tail; feet large, toes with a sparse growth of elongate hair; a sparsely furred area on neck; fur very long and dense on sides of neck, forming a ruff; fur on posterior part of back and on interfemoral membrane somewhat wavy. Color.-Head dark smoky brown, darker than general color of back; fur next to body much lighter than at tip; fur on posterior part of back and interfemoral membrane with a distinct brownish tinge; fur on sides of neck brownish amber, basal part of fur going from amber to orange; fur on belly grizzled, smoky, showing brownish tinges on sides of breast. Skull and teeth.-Skull similar to that of Cynopterus; sagittal crest distinctly developed; width of brain case little more than half zygomatic width; facial portion of skull short; alveolar line projected, passing at least 4 millimeters bElow audital bulla; postorbital processes strongly developed; supraorbital areas somewhat inflated. Outer upper incisors three-fourths the length but less than half the bulk of inner incisors and distinctly separated from them; inner incisors in contact near their summits; both pairs

180

PHILIPPINE LAND MAMMALS

of incisors much slenderer ' basally than distally; upper canines strong, curved; a slight distinct groove near outer part of tooth posteroexternally; a small but strongly defined secondary cusp posterointernally; first upper premolars very small, located in outer edge of tooth row, not crowded; second premolar and molars alike in shape, gradually decreasing in size; lower incisors only slightly smaller than inner upper incisors, in contact, narrowly separated from lower canines; latter moderate, secondary M easu?'em ents of Ptenochirus jagorii (Peters). [Measurements in millimeters.J

--=l

- -- ~~r~~i~:-- ~-. tion, No . 215.

':: 15

MInImum.- MaxImum.

: : : :: : :::::::: I ---------- -- - ------1

Ear : From meatus _______________ .. ______________________ _ F rom crown ________________________________ , ______ _

~~:B:a~~~~;-~~~-~;:~~~~~~~~~~-~~~-~~~-~~_~~j~~~~~~~~~~~

19 16 88 37

First and second phalanges ___ . ______________________ _

seco~e:an!:~:al ______________ . ________________________ c-I

Third finger: M etacarpal _____________________________________ ___ _ First phalanx _______________ ______________________ _ Secondphalanx _____________________________________ 1 ~

Fourth finger: M etacarpal ________________________________________ _ First phalanx ______________________________________ . Secondphalanx _______ ___________________________ __ ~

Fifth finger: Metacarpal _______ __________________ ______ . _______ _ _ First phalanx. _ . ___ ____________________ _________ ____ Second phalanx _ _____________ ~-------------,---- - - - Tibia _________________ ______________________________ ___ _ Foot __ ___________ __ ________________________________ ___ _ Skull : Condylobasallength ______ _________ ________________ _ _ Total length ___ _. ___ ___ ____________ ____ __ __________ __ Zygomatic width _______ _______ ______________ __ _____ _ Interorbital diameter ___ ____ ____ ____ _____ ________ ____ _ Orbit to tip of nasals ___ __________ __ __ _____ __ _____ ___ _ Diameter of orbit ___ _________ _____ _____ ____ ____ _____ _ Upper tooth row __ _______ ___ _____ ______ ____________ _ Lower tooth row ___ ___ __ ____ _____ ___ ____ __ ___ __ ____ _ Mandible __ ________ ___ ________ ___ _____ _. _____ __ ____'_,

18 . 5

------ ---- ---- ---- -80 33

86 . 5 34

40 21

34 20 . 5

36 22 . 5

59 39 55

50 . 5 34 . 4 47

56 38 51

53 32 33

48 . 5 26 . 0 29.5

52 29 . 5 32 . 5

56 28 30 33 19

60 . 5 24 . 5 26 30 20

53 . 5 26 . 5 30 32 21

37 . 5 39 28 8 9

35 37 24 6.8 8. 5 9.5 13 14 . 2 28 . 5

35 37 24 . 8 7.2 8.5 9.8 13 14 . 5 28 . 8

13 14 29

â&#x20AC;˘ Minimum and maximum measure,ments are taken from Andersen. loco cit. three spedmens, the type, and specimens from North Luzon and Mindoro.

CYNOPTERUS

181

cusp absent but a slight shelf marks its position; first lower ' premolar directly in tooth row, approximately the size of third lower molar; second lower premolar larger than molars . . Variations.-My specimen differs from Andersen's careful description of the species in that there is no anteromedial groove, but two anterolateral grooves, one on each side of the upper canines; the second upper incisors are three-fourths instead of one-half as long as the first; in other details, however, the skulls and teeth agree. The measurements of this specimen exceed somewhat other recorded measurements and are strikingly larger than the measurements recorded by Peters for the type. However, the specimen is not noticeably larger than one of Andersen's from "Caminguin North Luzon," which is noticeably larger than another, recorded from Mindoro. It is not improbable that these differences are sexual; however, the type is a male as is the largest specimen recorded by Andersen. Remarks.-My specimen was obtained from a shallow cliff cave formed by an overhanging rock. Only the single specimen was in the cave. I presume that the species is solitary. Judging by the numerous records, the species occurs throughout the Archipelago, except Palawan Province. The type, in the Berlin Museum, was collected at Daraga, Albay, southern Luzon, by F. Jagor. Genus CYNOPTERUS Cuvier

Cynopterus F. CUVIER, Des Dents des Marnm. (1825) 248; DOBSON, Cat. Chirop. Brit. Mus. (1878) 80; MATSCHIE, Flederm. des Berliner Mus. fur Naturk. (1899) 74; PmERS, Monatsb. Wiss. Ak. Berlin (1867) 867; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 47; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 613. Pa,chysoma GEOFFROY, Dict. Class. Rist. Nat. 14 (1828) 703. Naidius MILLER, Proc. BioI. Soc. Washington 19 (1906) 83.

Teeth without special modifications; upper incisors nearly terete, in contact wjth each other, separated from canines by a wide diastema, their crowns slightly flatten~d ~.nteroposteriorly, inner teeth of each pair slightly longer than outer; lower incisors slightly flattened laterally, outer of each pair slightly the larger; cutting edge blunt, nearly flat, divided by a notch into two distinct lobes; canines above and below with a secondary cusp on inner side, formed by a slight prolongation of cingulum and somewhat variable in various species; anterior upper premolar permanent, very small and short, scarcely penetrating gUmE, crown flat with a faint anteroexternal cusp; corresponding lower tooth considerably larger, the disk-shaped crown somewhat con-

182

PHILIPPINE LAND MAMMALS

cave except for minute anteroexternal cusp; three succeeding cheek teeth almost exactly resemble each other above and below; third premolar above and below higher and shorter than the others, inner ridge tending to ,f orm a blunt cusp opposite base of main cusp; crowns of molars longer than broad; second lower molar closely resembling anterior lower premolar, except that it lacks small anterointernal cusp. Skull primitive in form, ventral profile nearly straight, dorsal profile showing no peculiarities; floo-r of brain case scarcely deflected, though placed at a slight angle with palate ; occipital region without trace of tubular elongation; alveolar line projected backwards barely touching audital bulla and condyle; rostrum short, least distance from orbit less than lachrymal width; its dorsal profile concave; premaxillaries in contact anteriorly, not fused, slender throughout, tapering above to a recurved point distinctly below upper rim of nares; palate normal; audital bullre well developed, without tube or lip at meatus; mandible with broad low angular process. E xternal form short and heavy-; nostrils very prominent, almost tubular; upper lip divided by a, deep narrow ver't ical groove; second finger with well-developed claw; wings from sides of back; tail distinct, its terminal half free from interfemoral membrane; calcar weak, its length about equal to foot. (After Miller.) . 2-2 1-1 3-3 1- 1 Dental formula: 1. 2- 2' c. 1-1' pm. 3- 3' m. 2-2 = 30. Ander,sen has reduced the species to six with sixteen recognizable forms. The genus ' is represented in the Philippines by two species. K ey to th e Philippine species of Cynopt M'US Cuvie?". a ' . Alveolar line extended passing considera bly below condyle; teeth smaller, " squarish" ____________ ____________ ___ ___ ______ C. archipelagu s sp. nov. (p. 182). a 2. Alveolar line extended passing lower tip of condyle,' teeth more elongate. C. brachyotis Milller (p. 184). CYNOPTERUS ARCHIPELAGUS sp_ nov.

Type.-No. 345, E. H. Taylor collection; collected July, 1920, Polillo, Polillo 1., P. 1. by E. H. Taylor, Immature female; skin and skull. Diagnosis,. description of the type.-Head short, doglike; ears moderately long, lower anter:or margin thickened, convexly curv~d, then continuing to summit without, ~margination; summit rounded; posterior to summit an elongate shallow emargina-

183

CYNOPTERUS

tion; near base a slight angular lobule; nostrils distinctly tubular, separated by a deep groove; tail short, extending only a short distance beyond interfemoral membrane; calcar slender, elongate; feet large; pelage similar to Cynopterus brachyotis 120'

PHILIPPINE

ISL~NOS

124"

122

126

Batan Is

® Cynopterus b. brachyotis Cynopterus archipelagus © Mega-erops wetmorel @ Emb,allonuf'<1 aledo elledo ® Emballonura a. palawanensis ® Taphozous philippJnensis @ Taphonycteris pluto ® Taphonycteris capito

Il'

122

FIG. 8.

Distribution of

Oynop t~8, M eg reroPB ,

124 '

126

a nd Emballonur idm in t h e Philippi n es.

but shorter, without especially elongated fur in region of neck; fur on abdomen and over greater part of body less dense; forearm with short hairs on basal half; tibia and interfemoral membrane, except about calcar, with hairs of various lengths, longest near body; a distinct growth of fine hairs near outer upper edge

184

PHILIPPINE LAND MAMMALS

of posterior part of wing membrane extending halfway from foot to fifth finger; growth on lower edge less distinct; other characters as in Cynopterus brachyotis, but animal smaller and with a somewhat differently shaped, shorter head; temporal ridges obsolescent, widely separated. Skull a'YIJd teeth.-Skull distinctly smaller than that of Cynopterus brachyotis, rostrum shorter, supraorbital areas much less inflated, orbital diameter very distinctly less; alveolar line extended passing considerably below bulla; palate shorter than in C. brachyotis; postorbital processes shorter. Teeth very distinctly smaller than in C. brachyotis; upper incisors slenderer, canines less than half the length and bulk; first premolar about one-half the diameter, and upper cheek teeth less than one-third the size of corresponding teeth in C. brachyotis; teeth of lower jaw smaller than in C. brachyotis; two outer lower incisors almost in contact with canines and space between incisors equally divided. . Measurements.-(See table for Cynopterus brachyotis.) Remarks.-The specimen is immature. The teeth show no wear and the joints of the phalanges are not completely ossified. I have compared the specimen with an immature Cynopterus brachyotis. The difference between the two immature skulls is very striking. The nasal bones of C. archipelagus are very differently shaped, being shorter and much narrower, and show a strong constriction in the maddle. The characters of the molars, which are more squarish than those of Cynopterus brachyotis, suggest that this form is closely related to species in the "Nadius" section of Andersen, which includes C. horsfieldi, C. harpax, and C. princeps. The central cusp of the last lower premolar and the first lower molar seems to be entirely wanting. The single specimen, obtained in Polillo Island, was shot while it was suspended from the underside of a palm leaf. A second specimen was flushed from the same tree. CYNOPTERUS BRACHYOTIS Muller

This species has eight forms, only one of which is known from the Philippines. None of the others is known to occur even as near as Celebes and Borneo. CYNOPTERU:S BRACHYOTIS BRACHYOTIS (Miiller)

Pachysoma brachyotis MULLER, Tijd. Nat. Gesch. 5 (1838) 146 (Dewei River, Borneo, type locality); KREFFT, Cat. Mam. Aust. Mus. (1864) 5 (Borneo).

CYNOPTERUS

185

? Pachysoma b?'cvicaudatum WATERHOUSE, Proc. Zool. Soc. London (1843) 67 (Philippines). ? Pachysoma titth;ÂŁcheilum~ (non Temminck) WATERHOUSE, Proc. Zool. Soc. London (1843) 67 (Philippines). Cynopterus 1narginatus DOBSON, Cat. Chirop. Brit. Mus. 81 (Philippines); ELERA, Cat. Sisto Fauna de Filipinas 1 (1895) 1 (Abl'a, Luzon; Palawan); Contrib. a la Fauna Filipina (1915) 75. Pachysoma luzoniense PETERS, Monatsb. Akad. Berlin (1861) 708 (Iriga, S. Camarines, Luzon). Cynopterus sphinx luzoniensis MATSCHIE, Megachil'. Berl. Mus. (1899) 76; TROUESSART, Cat. Mamm., Suppl. (1904) 62. Cynopte?"us luzoniensis HOLLISTER, Philip. Journ. Sci. Â§ D 7 (1912) 8. Cynopte?'us ?na?'ginatus cumingii GRAY, Cat. Monk., Lemurs, and Fruit-eat. Bats (1870) 123 (Philippines). Cynoptents ma?'ginatus philippensis GRAY, Cat. Monk., Lemurs, and Fruit-eat. Bats (1870) 123 (Philippines). Cynopte1'us brachyotis b1'achyotis ANDERSEN, Cat.. Chirop. Brit. Mus. 2d ed. 1 (1912) 614 (Luzon; Mindanao, Zamboanga) (refer to this work for more complete synonymy); HQLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 334.

Description of the species.-(From adult male, in alcohol, No. 214, E. H. Taylor collection; collected on Polillo Island, July, 1920, by E. H. Taylor.) Head short, shaped like that of a dog; ears rather small, rounded at tip, with eight transverse ridges which end on posterior border, naked save about base on outer side; no antitragal lobe; a deep rounded notch in lower outer border with an angular projection posteriorly and a rounded lobule in front; anterior margin thickened, slightly folded back; nostrils prominent, projecting, tubular, and between them is a broad, furrow connecting with a slight vertical groove on upper lip, this groove bordered by elongate, granular, naked areas reaching to edge of lip; lower lip with a naked rounded area; second finger with claw much smaller than that on thumb; metacarpal bone of second finger much longer than entire first finger; metacarpal bone of third finger equals length of second; wing membranes attach to sides of body and to a point not far from base of claw of first toe; interfemoral membrane not extensive; a calcar and postcalcaneal membrane present; tail small, penetrating interfemoral membrane and extending beyond for more than half its length; wing membranes practically naked except near attachments to body; anterior part of snout (except naked area) with scant short hair and occasional longer vibrissalike hairs; fur short and dense on forehead and between eyes, growing longer on back and on neck both above and below; breast and abdomen with dense short fur; interfemoral membrane with fur

186

PHILIPPINE LAND MAMMALS

except in region anterior to calcar; antebrachial membrane with short fur on posterior part; forearm with short hair for half its length; fur on outer wing membrane near leg. Color.-Anterior part of head and body above mouse gray to brownish gray, outer tips of hair somewhat brown; fur on sides of neck, below ears, and on throat yellowish brown, rather strongly contrasted with other fur; fur on belly grizzled, showing no brownish tinge; hair on forearm dark brown; wing membrane deep blackish brown; anterior margin of ear whitish; phalanges whitish, contrasting with darker membranes. Skull and teeth.-Sagittal crest of skull very low, scarcely more than a slight ridge; postorbital processes wen developed; supraorbital area strongly inflated, the areas separated by a deep groove; facial portion of skull short, nasal bones terminating in a projecting point; alveolar line extended passes just outside audital bulla. Four upper incisors in a slightly curved group, separated from canines by a distinct space; inner teeth curved outward, in contact at middle but not at tips; outer teeth curved inward, in contact with adjoining teeth at tips; outer teeth slightly larger than inner; canines large, strongly curved backward, sharply pointed, with a well-defined anterointernal cusp; first premolar very small, nearer canine than second premolar; other cheek teeth without modification; lower incisors crowded together but none in contact or touching canines; crowns of inner teeth triangular, with a shallow triangular depression on anterior face of teeth; outer incisors much the larger; lower canines much smaller than upper; posterolateral cusp well-defined; first lower premolar about 2.5 times diameter of upper tooth; posterior lower molar with about one-fifth of crown surface of posterior upper molar. Variation.-Of this SUbspecies I have one specimen from Polillo (No. 214), two from Palawan (Nos. 313 and 591), and one (No. 329) from Mindanao. The Mindanao specimen (very immature), shows distinctly shorter measurements for the metacarpals and phalanges of the fingers; the foot is larger and the skull measurements are about equal to those of specimens from other localities. The 'dentition is too immature to show characters of value; the width of the brain case is slightly greater than that of the other skulls. The two specimens from Palawan agree very well in general measurements. There are slight differences in the characters of the teeth. The upper and lower incisors of No. 313 are slenderer than those of No. 519 (both

187

CYNOPTERUS

from near the same locality). From the described specimen from Polillo they show considerable differences both in skull and dental characters. The supraorbital areas of the Palawan forms are but slightly inflated and the longitudinal groove between them is very shallow; the facial portion of the skull is shorter while the brain case is distinctly broader than in the described Polillo specimen, and the zygomatic arches are more elongate and more expanded. MeaSU1'ements of Philippine species of Cynoptents

, -

Ct~vie?路.

[Measurements in millimetern.] Cynopterus bruehyotis bruehl/oti8.

Palawan, Ne.Polillo. No. 214.

313

fi91

ClinopterU8 arehipelagua from Mindanao, Polillo, No. 329. No . 345 .

Length, head and body (head stretch ed forward).

~:~~-~~:::::::::::::~~~:~~~~~:~=,

30 10.5

28 9

29 . 5 10 . 5

30 10

29 8

From meatus __________ ____ ____ From crown ________ __ ______ ___ I Forearm _________________ _____ _____ First finger, with claw __________ ___ __

19 16 67 27

20 15 70 . 8 27

18 . 6 15 69 27 . 5

19 15 68 28

16 14 60 . 8 27

First and second phalanges ______ Third finger: Metacarpal ________________ __ __ First phalanx- ____________ ___ __ Second phalanx __ _________ ____ _

32 15

28 18

28 15

27 17

27 16

45 31 38 . 5

42.5 29 38

42 . 3 29 38

36 23 30

40 28 36

Fourth finger: Metacarpal ________________ ____ First phalanx _____________ ___ __ Second phalanx ______________ __

43 23 25

40 22 26

40 21 25

36 20 20

38 21 23

44 20 23 25 15

43 21 22 26 17

42 . 0 20 22 25.8 16

41 17 18 24 18

38 19 20 21. 6 14

27 30 16 . 8 6 7

28 30 .2 22 6 .2

8 9.8 10. 7 22.3

28 . 2 --- ---- --29 30 . 2 19 20 7 6.2 7 6 7.8 8 10 . 1 -------.-11 ---------22 21

Ear:

Second finger: Metaearpal _____________ ___ ___ _

Filth finger: Metacarpal ________________ ____ First phalanx _____________ _____ Second phalanx ____________ ___ _ Tibia _________________________ ___ _ Foot _________________________ _____ Skull: CondylobaBallength _____ ______ _ Totallength ______________ ___ __ Zygomatic width ___________ ____ Interorbital diameter ____________ Orbit to tip of naBals _______ __ ___ Diameter of orbit____________ ___ Upper tooth row (alveolar) __ __ .__ Lower tooth row _______________ Mandible ______________________

10 . 5 11 . 2 22

I! I',

I I I

26 27 17 5. 5 6

188

PHILIPPINE LAND MAMMALS

The upper incisors of the Palawan specimens are distinctly smaller and slenderer than those of the described form and, instead of being closely crowded together, the two outer are separated from the inner pair by a di,s tinct space; the upper incisor is' much shorter and of little more than half the bulk, the first premolar is about three-fourths the size, and the molars are slightly larger than the corresponding teeth in the described form. The mandibular teeth have much the same character but are smaller in each case, and the molar-premolar series are separated by slightly wider distances, than in the described form. More material may show the necessity of separating the two forms. Remarks.-This species is not, apparently, gregarious in habit. I have occasionally startled single individuals from their diurnal sleep in clumps of dense-leaved palms. The s'p ecimens shot were hanging from the undersides of palm leaves. The species has been reported from Luzon and Mindanao. It is pl'obably a rare form in the Philippines. Mearns failed to obtain specimens. Andersen gives a very elaborate treatment of the forms of this species, listing eight. The typical form here considered is known from the Philippines, Borneo, and Celebes. These islands, except for the species herein described (Cynopterus archip,elagus, Polillo), have, so far as known, no other forms of the genus. [Genus THOOPTERUS Matschie] '"

Thoopterus MATSCHIEl, Megachirop. Berlin Mus. (1899) 72, 77; ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 662. Cynopterus DOBSON, Cat. Chirop. Brit. Mus. (1878) 80 (part.).

Related to Cynopterus; tail rudimentary, lateral membranes attach to second toe; skull in general as in Cynopterus; rostrum without special peculiarities; postorbital width less than interorbital width; postorbital foramin'a lacking; premaxillre slender, narrowing, ' barely in contact anteriorly; anterior edge of orbit directly above back of last premolar; zygomatic arches fragile; a low sagittal crest. Upper and lower incisors subterete, second upper incisor shorter than first; upper canine with a vertical groove on anterior surface, reaching from base to near top; no cingulum cusp; on lower canine a small cusp at inner base of tooth ending at level ,

'" Of doubtful occurrence in the Philippines.

189

THOOPTERUS

of incisors; first upper premolar small but heavier than an incisor; upper and lower molars and premolars short and broad; part of teeth distinctly quadrate; no inner cusp on second upper premolar; inner ridges of last upper premolar and first upper molar very low, those of corresponding teeth below perfectly obsolete, the inner edge being lower than crushing surface; last lower premolar and first lower molar with a large centrally placed surface tubercle or ridge. , . 2- 2 1- 1 3- 3 1- 1 Dental formula: 1. 2- 2' c. 1- 1' pm路 3- 3' m. 2- 2 = 30. Elera reports the species from Laguna and Batangas Provinces, Luzon; these records may be regarded as doubtful, but the species is very likely to occur in more-southern islands. [THOOPTERUS NIGRESCENS (Gray) 1

Cynopte?'us ma?'gin atus nigrescens GRAY, Cat. Monk., Lemurs, and Fruit-eat. Bats (1870) 123 (Moratai). Cynopterus (Thoopterus) nig1'eSCens MATSCHIE, Megachirop. Berlin Mus. (1899) 77; TROUESSART, Ca t. Mamm. Suppl. (1904) 62. Thoopterus nigTescens MILLElR, Bull. U. S. Nat. Mus. 57 (1907) 50; ANDERSEN, Cat. Chir op. Brit. Mus. 2d ed. 1 (1912) 665; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912 ) 8. Cynopterus latidens DOBSON, Cat. Chirop. Brit. Mus .' (1878) 86 pI. 5, fig. 3; ELERA, Cat. Sisto Fauna de Filipinas 1 (1895) 7 (Laguna and Batangas, Luzon).

Desc'ription of the specie-s.-Fur on body rather long (on back 9 millimeters, on belly 8 millimeters); tibi.re and metatarsi densely furred above; tail very short, can scarcely be seen; calcar normal; alar membranes attach to second toe; thumb smaller, but all other fingers, except fifth, longer than in Cynopterus. Color.-Back Prout's brown, underfur pale drab gray; head similar, but slightly darker, nape of neck paler; underparts, from chin to interfemoral, hair brown, slightly shaded with warmer brown on flanks and sides of breast and belly. Measurements of the type of Thoopterus nigrescens (Gray). mm.

Forearm Pollex: Total length with claw Metacarpal First phalanx S~cond finger: Metacarpal First phalanx Second and third phalanges with claws

73.5 26.5 8.5 14.5 35 8 9

190

PHILIPPINE LAND MAMMALS

Measurements of the type of Thoopterus nigrescens (Gray).-Continued, mm.

Third finger: Metacarpal First phalanx Second phalanx Fourth finger: l)ietacarpal First phalanx Fifth finger: Metacarpal First phalanx Second phalanx Tibia Foot with claws Skull: Rostrum, orbit to nares Interorbital width Postorbital width Orbital diameter Coronoid height Mandible, from condyle Upper tooth row from canine Lower tooth row from canine

51 37.5 43 48.5 29 50 22.5 22.5 26.6 20 9.1 8.7 7.7 8.5 14.5 27.8 13 14.5

Remarks.-The retention of this species is on very doubtful authority. Elera's record is wanting in the volume, Contribucion a la Fauna Filipina, a work edited and published many years after his death. There is, however, a great probability that the species occurs in Mindanao or in some of the Visayan islands. The type is a female, collected by A. R. Wallace, in Morotai, Gilolo group. Genus MEGlEROPS Peters

Megera TEMMINCK, Mon. Mamm. 2 (1841) 274 (non Wagler). MegtErops PETERS, Monatsb. Akad. Berlin (1865) 256; DOBSON, Mon. As. Chirop. (1876) 24', 29; Cat. Chirop. Brit. Mus. (1878) 81-87 (subgenus Cynopterus); MATSCHIE, Megachirop. (1899) 79 (subgenus of Ptenochirus); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 51, (genus); ANDERSEN, Cat. Chirop. Brit. Mus. 2d ed. 1 (1912) 646 (genus).

Closely related to Cynopterus, but differs from it in having usually a much deeper rostrum anteriorly and only a single pair of lower incisors, outer upper incisors somewhat reduced in length, and no tail or only a vestige of a tail; upper tooth rows slightly curved; no cusp, or only a slight secondary cusp on

MEGJEROPS

191

canine, no trace of vertical groove; molars and premolars rather short and broad, last upper molars sometimes deciduous; nostrils subtubular; odontoid papillre on inner side of lips; antebrachial membrane not extending beyond basal third of first phalanx of pollex; lateral membrane inserted on first toe near distal ex~remity of first phalanx; tibire with or without fur above; color silvery gray to brownish above . . 2+2 1-1 3-3 1-1 Dental formula: 1. 1 + l' c. 1-1' pm. 3-3' m. 2-2 = 28. The dental formula is identical with that of Ptenochirus. Magmrops differs from Ptenochirus in the absence of a welldeveloped tail, in the smaller size, and in the different type of incisors, lacking the well-defined secondary cusps. Two spedes are recognized, but only one is known from the Philippines. Key to the species of Meg;erops Pet'e rs. rL :.

Tail absent; forearm, 54 to 58 millimeters.

rL '.

M. eCrLudrLtus Temminck. Tail present, 1.5 millimeters; forearm, 49.5. M. wetmorei sp. nov. (p. 191).

The species here recorded is the first record of the genus for the Philippines. MEG.lEROPS WETMOREI aÂť. nov.

Type.-N o. 770, E. H. Taylor collection; collected near Tatayan, Cotabato, March, 1923, by E. H. Taylor. Skin and skull. Diagnosis; description of the type.-Adult female. Head somewhat mouselike, nostrils semitubular, separated by a deep groove or depression; upper lip with three grooves dividing naked area into two equal parts; low rounded tubercles, somewhat separated by short grooves, on either side bordering lip; a series of conical papillre on inner edge of lip above and below, pronounced near corners of mouth; tip of lower lip with two naked areas separated by a shallow groove; muzzle short with fine short hairs and a few vibrissre; eyes large; ears rather small, truncately rounded on tips, without or with a very slight emargination; antitragal lobe somewhat obsolete; a small triangular lobe anterior to notch, a few scattered hairs on anterior surface; posterior surface furred more than half the length from base of ear; five or six transverse grooves in ear; fur on forehead and sides of head extending anteriorly to eye; wing membranes ample, attaching on sides and to second phalanx of first

192

PHILIPPINE LAND MAMMALS

toe; limbs short; toes short, claws proportionally large; calcar short, curving backward; interfemoral membranes scant, tail scarcely visible; second digit with claw; length of second digit equal to metacarpal of third (claw not included) ; pelage long on back (7 to 10 millimeters), shorter on belly (5 to 8 millimeters) ; area on underside of neck with scant pelage; proximal third of forearm above and below with fine short fur; interfemoral membrane above with a prominent growth of elongate hairs, underside with shorter and much less dense hairs; tibia with very few, short, scattered hairs above, practically naked b~足 low; a .noticeable growth of long scattered hair on lower side of antebrachial membrane and on wing membranes about elbows and along sides of body. Color.-Head silvery to ashen gray, a slightly darker shade than on back of neck and shoulders; back ashen gray with outer tips of hair pale brown (in certain lights the color appears brown), below silvery ashen gray decidedly lighter than head and face; ears without yellow edge. Skull and teeth.-Skull like that of a diminutive Ptenochirus except that temporal ridges do not unite but continue back in subparallel lines separated by a distance of 4 millimeters; postorbital processes short, foramen distinct; supraorbital ridges somewhat inflated; occipital crest not prominent; lower jaw relatively heavy, coronoid process strongly developed. Teeth proportionally the same in general size and shape as in Meg;erops ecaudatus Temminck; upper incisors slender, styliform, separated by subequal spaces, space between second and canine greatest; second incisors shorter but slightly thicker than first; canines rather short, slightly recurved, cingulum practically wanting; a secondary cusp on anterointernal side probably well developed (in the type apparently greatly worn and not especially prominent); first upper premolars distinctly larger than incisors; lower incisors separated from each other and from canines by nearly equal spaces, slender, terete (impossible to state whether bilobed as teeth are worn) ; lower canines with a shelflike projection anterointernally which may show as a secondary cusp when unworn; anterior lower premolar about four times diameter of lower incisors; no second lower molars in lower jaw, but alveoli show that teeth are normally present, so that dental formula is probably, normally, identical with that of Meg33rops ecaudatus.

193

MEGJEROPS

Measurer/tents of the type of Mega31'ops wetmo1'ei sp. nov. mm.

Total length Head Tail Ear from meatus Ear from crown Forearm First finger with claw Second finger: Metacarpal First and second phalanx Third finger: Metacarpal First phalanx Second phalanx Fourth finger: Metacarpal First phalanx Second phalanx Fifth finger: Metacarpal First phalanx Second phalanx Skull: Total length Zygomat.ic width Interorbital diameter Orbit to tip of nasals Diameter of orbit Upper tooth row Lower tooth row, estimated Mandible

80 27 1.5

12 9

49.5

18 23 10 33 23.1 27

31 17 18 33.1 15 16.5

23.5 15.2

5.2 5.1 6 8

8.2 17

Remarks.-Closely related to M egmrops eca~(,clatus Temminck, Malay Peninsula to Borneo, the only other known species of the genus. The new species is distinguished as follows: Rostrum instead of being level from a point above last upper premolar to tip of nasals, slopes gradually, and the height at these two points is different (identical in M. ecaudatus) ; rostrum decidedly less broad and heavy than in M. ecal~datus; upper incisors more evenly spaced and separated by a greater distance from canines; lower incisors with a more-pronounced anterointernal hip or shelf than in M. ecaudat'us. Megmrops 'Wetmorei has a shorter forearm (49.5 millimeters; in M. ecaudatus, 54 to 57), and a visible external tail, 1.5 millimeters in length. Named for Dr. Alexander Wetmore, first assistant secretary of the Smithsonian Institution. 259453-13

194

PHILIPPINE LAND MAMMALS

Suborder MICROCHIROPTERA Mierochi1'optera DOBSON, Ann. & Mag. Nat. Rist. IV 16: 346.

(For

characters see key, p. 108; for families included, see p. 109.)

Family EMBALLONURIDJE V espertilionidre GRAY, London Med. Repos. 15 (1821) 299 (part.); Mag. Zoo!. and Bot. 2 (1838) 4913.

Rhinolotpha LESSON, Man. de Mamm. (1827) 81. V espertiliones part. BONAPARTE, Saggio di una Distrib. Metod. degli Anim. Vert. (1831) 15.

B1'achyura PETERS, Monatsb. Akad. Wiss. Berlin part. (1865) 257. Noctilionidm JElRDON, Mamm. India (1867) 30; GILL, Arrang. Fam. Mamm. (1872') 17.

Emballonuridm DOBSON, Ann. & Mag. Nat. Rist. IV 16 (1875) 349 (part.); Cat. Chirop. Brit. Mus. (1878) 353 (part.); GILL, Stand. Nat. Rist. (1886) 169 (part.); FLOWER and LYDEKKER, Mamm. Liv. and Extinct (1891) 666 (part.); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 82.

Humerus with. trochiter well developed though not as large as trochin, not articulating with scapula, neither tubercle rising above head; epitrochlea not especially developed, but with distinct spiny processes, capitellum nearly in line with shaft; second digit of forelimb with fully developed metacarpal, but no phalanges; third finger with two phalanges, proximal one flexed on dorsal surface of metacarpal when at rest; shoulder girdle normal; fibula complete, threadlike; tibia flattened or grooved posteriorly; pelvis norma,}, but boundaries of sacral vertebrre nearly or quite obliterated; skull with well-developed postorbital . processes (occasionally obscured by wide supraorbital ridges) ; premaxillaries represented by nasal branches only, never fused with each other or with maxillaries; palate ending in plane of last molar or produced somewhat behind it; if produced, very abruptly narrowed back of tooth rows; auditory bullre emarginate on inner side; teeth normal; tragus simple; muzzle without cutaneous outgrowths; tail perforating interfemoral membrane and appearing on upper surfaee distinctly back from edge. The family is represented in the tropical portions of both hemispheres. It occurs in the Pacific Ocean to Samoa, but is apparently absent from Australia and New Zealand. Two subfamilies are recognized, the Diclidurinre, confined to tropical America; and the Emballonurinre, found in both continents.

EMBALLONURA

195

Subfamily EMBALLONURIN..-E

Emballonurina GRAY, Ann. & Mag. Nat. Hist. III 17 (1866) 92. Emballonurinm DOBSON, Cat. Chirop. Brit. Mus. (1878) 353; GILL, Standard Nat. Hist. (1886) 169; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 85.

Characters those of the family; postorbital processes long and curved, in no way obscured by supraorbital ridges, their length at least four times their width; clavicle normal, its greatest width about one-sixth its length; tibia either subterete or with outer side flattened. The distribution of this subfamily is identical with that of the family. At least twelve genera are known; Taphozous, Taphonycteris, and Emballonura occur in the Philippines. Key to the Philippine genera of EmbalZonurVnm. a '. No radiometacarpal pouch; upper incisors, 4-; total number 34 ___ ___ __ _________ __ ______ ____ .__ ____ ____________ ___ __ ___ __ . Emballonura Temminck a'. Upper incisors, 2; total number of teeth, 30. b '. Radiometacarpal pouch present.... __ ____ ____ Tapho zous Geoffroy b 2. No radiometacarpal pouch ____ ___ ______ ___ Taphonycteris Dobson

of teeth, (p. 195). (p. 200). (p. 204).

Genus EMBALLONURA Ternrninck

Emballonura TEMMINCK, van del' Hoven's Tijdschr. Nat. Gesch. en Physiol. 5 (1838) 22; DOBSON, Cat. Chirop. Brit. Mus. (1878 ) 359; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 86; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 12; Proc. U. S. Nat. Mus. 46 (1913) 306.

Medium-sized bats with short muzzles and a deep depression between eyes; color variable; form slender; upper incisors simple, small, but well developed and permanent, subequal, inner tooth of each pair slightly the larger; lower incisors small, subequal, trifid, apparently often deciduous in old age; first and second in contact with each other, third slightly smaller than the others, and separated from them by a small space, and from canines by a much larger space; canines small, with welldeveloped cingulum, but no posterior prolongation; above and below, cingulum of canines develops an anterior and posterior cusp; small upper premolar very minute, without definite surface structure; two lower premolars almost exactly resembling canine but shorter and less slender; upper molars broad, first and second with distinct hypocones, third with metacone nearly as large as in first and second but lacking metastyle and fourth commissure; third lower molar smaller than first or second,

196

PHILIPPINE LAND MAMMALS

but with all elements complete; skull very delicate and lightly built, rostrum about as long as half the brain case, broad and flat with a distinct lateral inflation and a broad median longitudinal groove; anterior lower border of orbit expanded so tooth rows are not visible from above; postorbital processes elongated, reaching more than half the distance to zygomata; basisphenoid pits broad and shallow, median division barely indicated by a low ridge; audital bullre smaller than combined area of pits, distinctly emarginate anterointernally; mandible with angular process bent conspicuously outward; tibia scarcely flattened posteriorly. (After Miller.) . 2-2 1-1 2-2 3-3 Dental formula: 1. 3- 3' c. 1-1' pm. 2- 2' m. 3- 3 = 34. EMBALLONURA ALECTO (Eydoux and Gervais)

The specimens of this form from Pala'Yan show differences in the dentition that warrant separation from the typical species. Key to the subspecies of Emballonwra aledo (Eydoux and Gervais). Four of six median upper incisors in contact; firs't premolar .E. a. alecto Eydoux and Gervais a". Only two median upper incisors in contact. Lower canines molars reduced ........... ......... E. a. palawan ensis subsp. nov. Cb t .

larger . (p. 196). and pre(p. 197).

EMBALLONURA ALECTO ALECTO (Eydoux and Gervais)

V espertilio (Nycticeus) alecto EYDOUX and GERVAIS, Voy. autour du monde 5 pt. 2 (1839) 7 (Manila, Luzon).

Emballonura dis color PETERS, Monatsb. Konigl. Preuss. Akad. (1861) 711 (Paracale, Luzon).

Emballonura monticola HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 12; ELERA, Contrib. a la Fauna Filipina (1915) 115.

Emballonura alecto HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 306 (Guimaras) .

Description of the species.-(From No. 209, E. H. Taylor collection; collected near Caldera Bay, near Zamboanga, Mindanao, April 8, 1923, by E. H. Taylor.) Adult male. Very small; "nose" rather prominent, nostrils circular in outline, slightly tubular, opening directly forward, separated by a distinct depression; median part of upper lip with a naked, vertical ridge; lower lip with two naked areas separated by a broad shallow groove; a series of three distinct tubercles on each side of "nose," each bearing a long hair; ear rather bluntly pointed at summit, anterior margin very slightly concave below summit, then continuing in a straight line or very slightly convex to

EMBALLONURA

197

point of attachment above eye; posterior margin convex, attaching at a point below tragus; latter elongate, as wide at base as at summit, rounded slightly at summit with a tubercle near base on posterior margin; 12 transverse ridges on ear, continuing almost to summit; wing membranes attach rather high on back and to distal tip of tarsus; calcar large; interfemoral membrane ample; tail short, penetrating upper surface of membrane nearer body than distal edge; first finger with a slight callosity at distal part of metacarpal; body furred above and below with moderately long, not especially dense, fur; rostrum covered with sparse, short, dark hair and a few elongate vibrissre; wing membranes practically naked; interfemoral membrane with numerous scattered hairs above and below~ Color.-Fur bicolored, the outer three- to four-fifths of hair dark brown, shading gradually into pure white at extreme base; wing membranes dark blackish brown; muzzle dark blackish brown. Skull and teeth.-Anterior portion of skull flattened, interorbital distance much greater than narrowest constriction of skull; upper incisors small, arranged in two pairs widely separated from each other, and separated by a greater distance from canines; outer tooth of each pair larger than inner with which they are in contact, each tooth with a very small secondary cusp; canines with an anterior internal and a posterior internal cusp; first premolars -smaller than upper incisors, not rising as high as posterior cusp of canine and perceptibly separated from canine; widely separated from second premolar; second premolar with a low anterior cusp; six lower incisors forming a strongly convex row extending considerably anterior to canines, the outer widely separated from canines and by a much shorter distance from other incisor; four inner teeth in contact, innermost pair largest; lower canines and two premolars each with '3 small anterior and a posterior cusp; lower premolar,s caninelike, first smaller than second. Remarks.-Specimens of this species were taken in a dark pocket formed by a large overhanging rock near Caldera Bay, Zamboanga, Mindanao. The colony consisted of, at most, a dozen specimens. The type is from "Manila." EMBALLONURA ALECTO PALAWANENSIS subsp. nov.

Type.-N o. 72, E. H. Taylor collection; collected along I wahig River at base of Thumb Peak, Palawan, October 28, 1923, by E. H. Taylor. Alcoholic specimen and skull.

From crown ___ ___ ____ ________ ____________

Fourth finger: MetacarpaL ____ _______ ____ _______________ First phalanx _________________________ ___ _ Second phalanx _____ _____________________ J

Third finger: Metacarpal ____ __ _____ _________________ ___ First phalanx- _____________ ___ ____________ Secondphalanx ____________ ___ ____________

From anterior attaehmenL _____ ____________ Forearm__ _______ __ ____ _______________________ First finger _____ ____ _____ _____ __ ______________ Secondfinger __________________ ___ ____________

Ear:

Tragus: Height at anterior edge ____ ________ __ _______ Greatest width at summiL ______ ___________

Totallength _____ ________ ___ ___ ____ ___________ Head and body ___ ______ _______ ___ ____ _________ Tail. __ __ ______ ________ ____ ______ ____ ___ ______

34 9 8

41 14 20

12 14 45 8.5 38

3. 5 1.8

35 9. 5 8

41. 5

11 15 46 9 38 . 5

3. 8 1.8

42 12

female.

female .

54 43

72,

68,

41 14 17

15 .2 43 8.4 37 .2

3.5 1.8

03 42

female.

70,

EmbaZlonura aZecto pala Ifa nemis.

32 . 6 10 . 5 7

39 . 5 13 20

12 . 5 16 47 8 36

32 9

40 12 18

13 16 45 8 36

3.9 1.9

59 47 12

38 11 . 5 18

13 15 . 6 44 7. 0 36

3.8 1. 9

56 45

3.8 1.8

male.

207,

- - - - --

male.

20.6,

56 45

- --

male.

205, male.

209,

32 9 6

39 12 . 5 18

12 . 7 15 4路5. 2 8 36

3. 6 1.8

58 46 12

33 10 7

39 12 18

12 16 46 8.3 36

3 1.5

58 46 12

- - - - --

male.

208,

32 9 7

39 12 18

13 15 44 8 36

3.5 1.7

57 46

211,

39 11 . 2 19

13 16 45 8 36

3.5 1.7

58 45 13

34 10 8

41 13 21

13 16 48 8.3 38

3. 6 2

59 46 13

I 212, I 213, I female. female. female.

EmbaZZonu ra aZecto a/ectlo.

E. H. Taylor collection N o. a nd sex.

[Measurements in millimet ers.]

Measurement8 of Emballonura alec to palawanensis and E. a. alecto.

z l::'

t".l

"tl

~ .... t"' .... "tl "tl ....

Lower tooth row - - - ---- - -- - -- -- -- - - - - -- - -Mandible______________________.___________

Upper tooth row (to base of canine) _________

Skull: Total length to incisors _____________________ Condylobasallength _______________________ Width across molars (widest) _______________ ZYgoma tic width ______________ ____ __ ______ Interorbital width _________________________ Greatest skull constriction _______ ___________

Fifth finger: MetacarpaL ______________________________ First phalanx ___________ ______ ____________ Second phalanx __ ___ _____________ _________ Tibia ________________________________________ Foot ___ ____ _____________ _____________________ Calcar __ ____ _____________ _________________ ___

15 13. 9 6.2 8 4. 4 2.9 5. 3 5. 6 11

31 11 5 5. 5 16.5 8 12 11

5 17 8 13

5. 5 18 8.2 12 14 . 9 14 6 8 .1 4.7 3 5. 6 5. 8 11

-------- -------.-

--- ----,-

-_._--- -- --------

-------- -.=;.------

-.------- ------...,-

28 10 5 16 8 12

-------- --- --- --- ------ --------- ------ --------------- --------

29 10 5 17 . 5 9 13 . 15 . 1 13 . 8 6 9 4 .6 3 5.7 5. 9 11 . 2

18 9 13

80 10 . 2 17 11.5 18

11 6

17 8 12

30 11

29 10 5 16 8

15. 3 14.2 13.8 13 .2 5.5 6 8.7 9 4 . 1 -------- -------8 2.6 5.5 5 .6 5.8 5.8 11 11.1

29 10 5 17 7.9

15 . 1 18.8 6 82 4 2.8 5. 6 5.9

17 8 18

29 12

15 . 3 14 . 1 6 8.2 4. 5 3 5. 8 5. 9 10 . 8

81 12 6 18 8. 5 13

t;j

~ ~

t"' 0

ttl

is:

200

PHILIPPINE LAND MAMMALS

Diagnosis .. description of th,e type.-Differs from typica1 E. alec to in lower jaw being wid~r at tip; lower incisors form a broader, less-convex row; teeth spread apart, with an almost equal distance betwe.en teeth; only the two median in contact; first lower premolar about one-half as large as corresponding tooth in E. alecto; teeth slenderer, especially ,those of lower row; thil? character particularJty eviden~ in canines. In orte ~ of the specimens the first upper premolars are wanting, probably deciduous in old age; the brain case is smaller, not in actual length but in the amount of inflation. There are no pertinen.t differences in relative measurements as shown py the table of measurements under E. alecto alecto. Color.-Practically the same as for typical form. M easurements.-See under typical form. Remarks.-Five specimens were taken under' an overhanging rock along Iwahig River, near the base of Thumb Peak. Genus TAPHOZOUS Geoffroy

Taphozou8 GEOFFlROY, Descr. de l'Egypte 2 (1813) 113; DOBSON, Cat. Chirop. Brit. Mus . . (1878) 378. Saooolaimus LESSON, Nouv. Tabl. Regn. Anim. Mamm. (1842) 19.

Upper incisor in main axis of premaxillary very minute, often absent in old individuals, its tip never reaching level of cingulum cusp of canine; lower incisors large, imbricated, trifid, forming a continuous and strongly convex row between canines; canines strong, with well-developed posterior extension, especially in maxillary teeth; cingulum of upper: canine develops two small though distinct cusps, one anteriorly, one posteriorly; these cusps barely indicated in lower canines; W-pattern of molars shallow; audital bullre large. Externally characterized by a short conical muzzle; a deep hollow between eyes; long lower lip terminating anteriorly in two naked areas not or 'barely separated by a groove; foot slender; tail simple, perforating interfemoral membrane above; radiometacarpal pouch distinct . . 1-1 1-1 2-2 3-3 ' Dental formula: 1. 2-2' c. 1-1' pm. 2-2' m. 3-3 = 30. One species in the Philippines. ' TAPHOZOUS PHILIPPINENSIS

Waterhouse

TaphozOU8 philippinensis WATERHOUSE, Proc. ZOO!. SOC. London (1845) 9 (Philippine Islands) ; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 12; Bull. U. S. Nat. Mus. 46 (1913) 307 (Malabon and Montalbanj Luzon). Taphozous melanopogon (part.) ELERA, Contrib. a la Fauna Filipin'ci (1915) 117.

TAPHOZOUS

201

Description of the species.-(From No. 333, E. H. Taylor collection; collected at Tatayan, Cotabato, Mindanao, March 15, 1923, by E. H. Taylor.) Adult male. Head, from above, triangular; muzzle rather acute; nos'e slightly prominent; nostrils terminal, somewhat tubular, only slightly separated; upper lip terminating in a point, covered with a trian:g ular naked area; lower lip somewhat swollen at tip, with two triangular naked areas separated by a shallow groove and a large transverse swelling below chin; ears rounded at summit, anterior edge nearly straight to near head, then running forward on to forehead, and attaching above eye on edge of. frontal fossa, separated from its fellow by a space of about 4.2 millimeters; space between occupied by a deep frontal fossa; posterior edge of ear convex as far as ill-defined antitragal lobe which runs forward ljnd attaches below and a short distance behind angle of mouth; '.Interior ear margin with a series of very small tubercles; lateral outer margin very slightly emarginate and on inner side numerous small transverse folds or ridges; tragus, from inner base, about 6 millimeters high, about 4.8 millimeters wide at summit; summit rounded; base about 3 millimeters wide, anterior edge more strongly concave than posterior; tragus with short hair over middle portion; tail for about one-half its length involved in interfemoral membrane; latter about 23 millimeters in anteroposterior extent, naked beneath, slightly hairy above to base of free portion of tail; latter with a few long scattered hairs; feet almost naked, having only a few scattered hairs; wing mem~ branes attach on sides of back and to tarsus 4 millimeters from distal end; end of first finger metacarpal with a rounded callosity below; eyes rather large, prominent; face and jaws with very short fine hair. Color in life.-Pelage same above and below; outer tips of hairs brownish, lower parts white to creamy white; lower fur slightly lighter brown on distal portion; wing membranes brown; interfemoral membrane paler below than above; short unequal hairs on snout dark brown. Teeth and skull.-Upper incisors minute, near inner anterior edge of canines, often concealed by gums; canines large, interior edge of cingulum forming two small distinct cusps; first premolar small with two distinct cusps; lower premolars caninelike, the first largest, the second highest; lower incisors strongly trifid, tips deeply emarginate, forming a strongly convex row between canines, the two middle slightly smaller and on a lower level th!'tn Quter; skull .s imilar to description given for genus.

333,

I female. 366, I 391, male. 394,

male. I fe

--I

42 12 7

44 12 8

:!:;;:~~;:~~:~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~~I

Secondphalanx---------------------------------------- __ Fourth finger:

54 19 21

22 16 5 6.5 62 10.2 50

21 12

94 73

56 20 22

20.5 15 4.8 6 62 10 52

99 75

~etacarpal---------------------------------------------_I First phalanx- ___________________________________________

Third finger:

From lower attachment ... -- -- --- ____ -- -- _________ -- _--From crown _____________________________________________ Tragus, width .. __________________________________________ Tragus, height ___________________________________________ Forearm . . ___________________________________________________ First finger ____________________________________ ---------Secondfinger ________________________________________________

Ear:

~;::~~;:~::~~~~:~~~~~~~~~~:~~~~~~~~~~~:~:~~~~~~::~::::~:1

Tail:

Total length ______ --- -- _-- __ ---- __ ---- ____ --- ___ --- ________ Head and body length ________________________________________

--I

44 13 9

56 20.5 22

22 15 5 6 65

25 13

I- --~~---I

44 14

55 20 22.5

22 路 15.5 4.6 6 63 9.8 52

24 16

100 76

----------------------I--- --- -~-I---

male.

407,

male.

44.8 13 8

55 20 22

21 15 5.1 6.8 62 9.5 52

24 13

99 75

43 12 8

55 19 22

20.4 15 5 6.5 65 10.5 52

25 15

99 74

---- - - -

403,

male.

E. H. Taylor coIl\ection No. and sex.

[Measurements in millimeters.]

Measurements of Taphozous philippinensis Waterhouse.

45 12 8

56 21 22

22.5 14.6 5 7.1 66 10 52

23 14

98 75

---

408.

male.

44 18 8

55 20 22

22 14 4.9 7 64 10 52

23 15

98 75

---

409.

male.

44 12.5 9

58 21 23

21.5 14 5 7 66 10.5 54

26 14-

103 77

No. 144861, female.

~useum

United States National

t\:)

r.n

~ ~ t"

t::I

.....

'"d I'd

:z:.....

I'd

t\:)

Interorbital width _______________ •... _.•.....•••.... _.• __ . Upper tooth row including canines ..... _.... __ ._._ ..... __ • __ Lower tooth row including canines .......................... Man dible __ . __ .• __ . _. _. __ . _...•... _•. __ ..• __ • __ ... _. _. _.•

Skull: Total length to canine tip _________________________________ Condylobasal length ______________________________________ Zygomatic width _________________________________________ Brain case, width over bulhe. ______________________________

Calcar ______________________________________________________

~bia

Fifth finger: MetacarpaL _____________________________________________ First phalanx ____________________________________________ Secondphalanx __________________________________________ FeDlur ______________________________________________________ _______________________________________________________ Foot ________________________________________________________

21 20.5 12 11 6 9.2 10 17

34 12.5 8 23 23 13.4 17.2

5.6 9 9.8

20.5

32 11 8 24 20 12 17 21 20.5 11.6 11.8 6 9 10.1 16.5

34 13 9 22 23 13 17 21 20.5 11 11.8 6 9.2 10 17

34 12.5 7.2 22 23 13 18 21 20.5 12 12 6.1 9 10.1 17

33 13 8 22 23 13 17 21 20.5 12.5 11.4 6.1 9.5 10 16.6

34 12 8 22 23 13 17

12 11.3 6.2 9 10 16.3

~~.4

34 13 8 22.5 21 13 16

___

12 12 6.2 9 10 17

~~_..1

34 13 8 23 23 13 18 20.9 20 . 5 12.6 11.6 6.3 9 10 16

33 12 8 23 23 13 18 21 20.5 13.3 12 6.5 9.6 10.3 17

34 13 8 27 25 14 20.5

C<:>

~ c:::

> "1;j

204

PHILIPPINE LAND MAMMALS

Variation.-The specimens s~udied are from Malabon and Manila, Luzon (4 specimens) ; an~ Saub, Tatayan, and Sarangani Bay, Cotabato, Mindanao (18 specimens). The specimens from Cotabato vary imperceptibly frpm those from Luzon (Note Nos. 333, 391, 394, 399, 403, 407, 408, and 409, from Mindanao). Hollister * lists a series of Luzon specimens collected near Manila. The measurements I give , here are sHghtly greater than those recorded by Hollister for the series of twenty-six specimens. In color there are varying shades of brown and an occasional "reddish" phase which has the underfur somewhat buff. R emarks.-The Cotabato sPElcimens were obtained from shallow caves and waveworn cliffs along the seashore. The colonies were small, at most, only two or three dozen individuals clinging to the roof. They too~ flight immediately on hearing the least noise of my approach. I am indebted to Mrs. Ida M~ McCrory, of Manila, for a specimen (No. 366) collected in her home in Manila; the specimen flew into the house through an open window. The type of the species, now in the British Museum of Natural History, was collected by H. Cuming in the Philippine Islands. The exact l~cality is ,no longer known. Genus TAPHONYCTERIS Dobson

Taphonycteris HOLLISTER,

DOBSON, Cat. Akiat. Chirop. (1876) 167; (subgenu~); Proc. U. S. Nat. Mus. 46 (1913) 308 (genus).

Closely related to Taphozous in general characters but differing in having no radiometacarpal pouch and in having :the lower lip divided in the center of its upper surface and in front by a deep narrow groove; no emargination on inner side of audital bulla; small upper premolar much larger proportionally than in Taphozous. . . 1- 1 1-1 2-2 ' 3-3 = 30 Dental formula. 1. 2-2' c. 1- 1' p.m. 2-2' m. 3-3 . Key to the Philippine species of Taphonycte?'is Dobson. a '. Skull moderate; pollex and metacarpals normal.

, T. pluto (Miller) (p. 205). a '. Skull larger; with smaller pollex, metacarpals, and teeth smaller. T. capito Hollister (p. 204). TAPHONYCTERIS CAPITO Hollister

Taphonycteris capito 308.

HOLUSTEjR,

Proc. U. S. Nat. Mus. 46 (1913~

. * Proc. U. S. Nat. iMus. 46 (1913) 307.

205

TAPHONYCTERIS

Description of the species.-This species is related to Ta,phonycteris pluto but is smaller, with shorter pollex, uniformly shorter metacarpals, and smaller claws. Color.-Upper parts rich brownish black, sparingly spotted with white on head, back, anq rump; underparts sepia, marbled with whitish; wings and interfemoral membrane (in dry skin) intense black. Skull a,nd teeth.-Skull larger and, especially, broader than in T. pluto, rostral portion more massive with greater interorbital breadth; teeth actually smaller than in T. plvto. Measurements of the type of Taphonycteris capito

Holt~ster.

mm.

Forearm Third finger Metacarpal Fourth finger, metacarpal Fifth finger Metacarpal Skull: Condylobasal length Zygomatic breadth Mastoid breadth. Breadth of brain case Postorbital constriction Interorbital constriction Maxillary tooth row Mandible Mandibular tooth row

119 71.5 51.1 67 36 21.6 16.1 13.9 11.0 5.1 8.7 10.2 18.7 11.9

Remarks.-The type of the species was collected at Pandan, Catanduanes, in 1909, by D. B. Mackie. The type, No. 155149, and one paratype are in the United States National Museum. Hollister records an immature specimen that is almost pure black above with no white markings. TAPHONYCTERIS PLUTO

(Miller)

Taphozous pluto MILLER, Proc. U. S. Nat. Mus. 38 (1911) 396; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 12. Taphonycte1'is pluto HOLLISTER, Proc. U. S. Nat. Mus. 16 (1913) 307.

Description of the species.-Similar to Ta,phozous sa,ccola,imus Temminck of Java, but smaller; adult female, forearm 69 to 72 millimeters instead of 74 to 76 millimeters; condylobasal length of skull about 21 millimeters instead of about 23 millimeters. (From Miller.)

206

PHILIPPINE LAND MAMMALS

Measurements of the type of Taphonycteris pluto (Miller). mm.

Forearm 71.4 Third finger 116.0 Fifth finger 67.0 Tibia 25.5 Foot 17.3 Skull: Condylobasal lengt.h to front of alveolus of canine 21.2 Zygomatic breadth 15.8 Mastoid breadth 13.2 Breadth of brain case 11.0 Postorbital constriction 5.0 Interorbital constriction 8.0 Mandible 18.6 Maxillary tooth row 10.6 Mandibular tooth row 12.0

Remarks.-The type, an adult female, was collected at Mercedes, near Zamboanga, Mindanao, March 28, 1906, by Edgar A. Mearns. It appears that the species is based wholly on measurement characters. Family MEGADERMIDlE V espet路tilionid:E GRAY, London Medical Repository 15 (1821) 299 (part.); BONAPARTE, Nuovo Ann. del. Scienze Nat. Bologna Anno 1 Torno 2 (1838) 112 (part.); GRAY, Mag. Zoo!. & Bot. 2 (1838) 486. Rhinolophina LESSON, Man. de. Mamm . . (1827) 81 (part.). Rhinolophidre GERVAIS, Rist. Nat. des Mamm. (1854) 200 (part.); GRAY, Proc. Zool. Soc. London (1866) 83; WINGE, Jordfund. og nuL Flagerm. fra Lagoa Santa, Minas Geraes, Brasilien (1892) 24. Megadermatidre ALLEN, Monogr. Bats N. Am. (1864) 1 (part.); ANDERSEN and WROUGHTON, Ann. & Mag. Nat. Rist. VII 19 (1907) 129. Megadermata PETERS, Monatsb. Akad. Wiss. Berlin (1865) 256 (part.) . Megadermidre GILL, Arrangements Fam. Mamm. (1872) 17; Standard Nat. Rist. (1886) 164 (part.); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 101. Nycteridre DOBSON, Ann. & ' Mag. Nat. Hist. IV 16 (1875) 348; Cat. Chirop. Brit. Mus. (1878) 154 (subfamily Megaderminre); FLOWER and LYDEKKER, Mammals Living and Extinct (1891) 658 (part.); ELERA, Contrib. a la Fauna FiIipina (1915) 92.

Humerus with trochiter small, about equal to trochin, not articulatiI).g witn scapula, two tubercles rising slightly above head; epitrochlea large, with styloid process, capitellum carried beyond line of shaft; second finger with one phalanx, third with

MEGADERMA

207

two phalanges; shoulder girdle abnormal in the great widening of anterior portion of presternum (greatest width nearly four times length) and in fusion of this widened portion with first rib, first dorsal and seventh cervical vertebra into a solid ring of bone; keel of sternum only moderately developed; fibula threadlike, less than half as long as tibia; pelvis normal, ischia free posteriorly; skull with postorbital processes short or practically absent, usually obscured by wide supraorbital ridges; premaxillaries absent; upper canines proj ecting noticeably forward, shaft with a large secondary cusp; tragus unequally bifid; muzzle with conspicuous erect leaflike cutaneous outgrowths; upper incisors wanting; tail very short; ears very large. The family is distributed in tropical Africa, southern Asia, Malay region, and Australia. Of the five recognized genera only M egaderma is represented in the Philippines. Genus MEGADERMA Geoffroy

Megaderma GEOFFROY, Ann. Mus. Hist. Nat. Paris 15 (1810) 197; DOBSON, Cat; Ohirop. Brit. Mus. (1878) 154 (part subgenus); ANDERSEN and WROUGHTON, Ann. & Mag. Nat. Hist. VII 19 (1907) 130; MILLE!R, Bull. U. S. Nat. Mus. 46 (1907) 101; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 13; Proc. U. S. Nat. Mus. 46 (1913) 308.

Spasma GRAY, Proc. Zoo!. Soc. London (1866) 83.

No upper incisors or premaxillaries; lower incisors large, slightly trifid, forming a continuous, slightly convex row between . canines, inner tooth smaller than outer; canines large, with welldeveloped cingula, the lower without secondary cusps, upper with a minute anterior cingulum cusp and a large posterior cusp at base of shaft; anterior upper premolar minute, on inner side of tooth row in angle between canine and large premolar; first and second upper molars with three main cusps rather close together, particularly in first upper molar, and mesostyle so reduced that the W -pattern is considerably distorted by partial obliteration of median commissures; posterointernal heel well developed but without trace of hypocone; third upper molar with four cusps and two commissures; lower molars with cusps near together and those on inner side reduced in size; particularly the entoconid, which is practically absent from posterior lower molar; skull with rostrum slightly less than half as long as brain case, slender, slightly concave above, without supraorbital ridges or special widening in interorbital or lachrymal regions; postorbital processes reduced to' merest trace; basi-

208

PHILIPPINE LAND MAMMALS

sphenoid pits shallow but distinct, partly overhung by concavespatulate hamulars; audital bullre small, their greatest diameter less than distance between them; sagittal crest low and indistinct, bifurcating anteriorly into two lines terminating in rudimentary postorbital processes. (After Miller.) . 0- 0 1- 1 2- 2 3- 3 Dent~l formula: 1. 2- 2' c. 1-1' pm. 2- 2' m. 3- 3 = 28. The genus ranges from India and Ceylon throughout the Malay region. Four species are recognized; one is known in tre Philippines. MEGADERMA SPASMA SPASMA

(Linnreus)

V espertilio spasma LINN)EUS, Syst. Nat. 10 ed. (1758) 32. Megaderma spas'rna GEOFFROY, Ann. Mus. d'Hist. Nat. 15 (1810) 195 ..

Megaderma philippinensis WATERHOUSE, Proc. Zool. Soc. London (1843) 69 (Philippine Islands).

Mega.derma spasma spasma ANDERSEN and WROUGHTON, Ann. & Mag. Nat. Hist. VII 19 (1907) 132; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 13; Proc. U. S. Nat. Mus. 46 (1913) 308 (Catagan and Iligan, Mindanao); ELERA, Contrib. a la Fauna Filipina (1915) 93.

Description of the species.-(From No. 310, E. H. Taylor collection; collected at Saub, Cotabato, Mindanao, April 27, 1917, by J. P. Torres.) Nose leaf prominent, posterior part elliptical, much longer than anterior horseshoe, with a median fold which, seen from the front, appears as a ridge or keel; at its base the fold expanding into a broad heart-shaped leaf, its width equal to width of horseshoe, and wider than base of posterior leaf; forward extension covering nostril but not reaching edge of horseshoe; a small triangular area at tip of upper lip naked; a larger naked triangular area on lower jaw, divided by a broad distinct groove; two rounded swollen areas, on either side of lower jaw, bordering naked areas and separated by a median groove; a strong tubercle in front of each eye, bearing t wo stiff bristles; a similar tubercle in front of ear, bearing several bristlelike hairs; snout on each side of horseshoe swollen; ears extraordinarily large, reaching beyond tip of snout when laid forward, upper edge broadly rounded, posterior edge without notches and nearly straight; a number of short transverse ridges near outer part not extending to tip; ears joined together medially by a membrane, thus forming, when viewed from behind, a shallow pocket; tragus bifid, posterior part high, sharply

209

MEGADERMA

pointed at tip; anterior part low, bluntly pointed, anterior margin slightly serrate; eye moderate; wing membranes ample; that anterior to forearm attaches at outer tip of metacarpal of thumb; metacarpal of index finger distinctly longer than that of third, in turn shorter than fourth; fifth longest; only two phalanges in third finger; wing membranes attach low on sides, and to foot near dorsal base of outer toe; calcar somewhat shorter than foot and toes; no postcalcaneal lobe; foot small, toes slender and elongate; tail vestigial; (mammre two pairs, * anterior, abdominal, widely separated; posterior, inguinal, approximately one-third as far apart as anterior). Teeth and skull.-Practically identical with description given for genus. Color (preserved in alcohol) .-Pelage above rather ash gray, lighter at base, not extending on inner margin of web; proximal third of forearm with rather heavy brownish fur; interfemoral membrane naked; below, membranes naked except for minute hairs on posterior borders; belly fur lighter gray, whitish at tips; fur at base of ear buff; hairs on snout and nose leaf dirty buff; fur on forehead and about eyes rather grayish; feet with numerous hairs above; wing membranes brownish, that part near thumb buff; on metacarpals and phalanges buff; ear membranes light, somewhat darker on outer part; tragus with hai,:s; skin of lips and chin whitish buff. Remarks.-I obtained three specimens of this species from an old stump at Saub, Cotabato (Nos. 308, 309, and 310) ; Nos. 311 and 603 are from Los Banos, Laguna, Luzon. A larse female in the collection of the College of Agriculture, Los Banos, Luzon, contained a single large embryo. The feet are extraordinarily large, equalling the length of the feet of the mother without the claws. The type of the species was collected, it is presumed, in Ternate. Andersen, t who has examined the type of Megaderma philippinensis Waterhouse, states, "They are in every respect indistinguishable from Celebes individuals." Andersen has separated the typical form of M. spasma from that found in Java, which he records under the subspecific name trifolium. Apparently, the two forms are closely related, differing chiefly in the length of the lower leg.

* From

No. 311.

t Ann. & Mag. Nat. Rist. VII 259458-a

(1907) 132.

210

PHILIPPINE LAND MAMMALS

Measurements of Megaderma spa sma spasma (Linn:ÂŁus). [Measurements in millimeters.} E. H. Taylor collection No. and

308,

309,

310,

603,

311,

male.

female.

- -- - - - - - -- - - - - - - - - - - - - - - - - - - Total length, head and body. ________________ Head _____________________________________ Height of ear, outer side.. ___________________ Width of ear (greatest) .. ____________________ Tragus length. _____________________________ ____________________ _________ Tragus~dth

Width of horseshoe _________________________ Width of median leaL ______________________ Width of posterior leaf (not expanded) ________ Combined length of leaves. __________________ Forearm .. _________________________________ First finger and claw .. _____________________! Secondfinger ______________________________ Third finger: MetacarpaL ________________._________ __ First phalanx _________________________,_

Second phalanx - - - - - - - - - - - - - - - - - - - - - - Fourth finger: Metacarpal. ___________________________ First phalanx __________________________ Secondphalanx ________________________ Fifth finger: Metacarpal ____________________________

Firat phalanx __________________________ Interfemoral membrane, length .. _____________ Femur ____________________________________ Tibia _____________________________________ Calcar ____________________________________ Foot. _____________________________________ Skull: Totallength ___________________________ Condylobaaallength ____________________ Zygomatic ~dth _______________________

Upper tooth row

~th

canines, length _____

"0"' 'ow w;tb ~;o'-------------

Interorbital width ______________________ Mandible length .. ______________________

s~ -------'-

7.2 6.3 13.8 57 18 46

27 37 25 20 8 7 7.2 7 13 55 18 46

72 28 36 25 21 8 8 7.6 7 13.5 58 18 46

37 20.5 42

38 22 40

37 20 38

37 19 38

42 13 17

42 13 18

42 13 17

37 12.5 17

43 12.1 17

47 13.5 40 31 30 13 18

47 14 40 30 30 18

48 14 39 28 30 12 18

45 14 48 31 30 13 17

48 14 44 29 29 13 16.5

26 23 14.5 10

25 22 14.5 9.8

26 22 14 10

26 22.5 14 10

26 23 14 10

4.5 18.5

4.5 18.4

4.4 18.5

4.5 18.2

29 39 25 21 8 8 8 7 13.2 56 18 44

68 27 39 25 20 8 8 7.8 7 14 54 18 45

72 29 38 25 20 8.2

37 20 36

2J 4.5 18

Family RHINOLOPHIDJE Miller Rhinolophina LESSON, Man. de Mamm. (1827) 81 (part.). Rhinolophides GERVAIS, Rist. Nat. des Mamm. (1854') 200 (part.). Rhinolophi PETERS, Monatsb. Akad. Wiss. Berlin (1865) 256 (part.). Rhinolophid;;e GRAY, Proc. Zool. Soc. London (1866) 81 (part.); GILL, Arrang. of Fam. Mamm. (1872) 17. (part.); DOBSON, Ann. & Mag. Nat. Hist. IV 16 (1875) 348 (part.); GILL, Stand. Nat.

RHINOLOPHUS

211

Hist. (1886) 164 (part.); FLOWER and LYDEKKER, Mamm. Liv. and Extinct (1891) 656 (part.); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 106 (as nere restricted~.

Ears large, without tragus; nose leaf conspicuous; toes with three phalanges; humerus with trochiter about as large as trochin and definitely articulating with scapula by an articular surface half as large as glenoid fossa; both tubercles rising slightly above level of head; epitrochlea large and with welldeveloped styloid process; capitellum nearly in line with shaft; second finger consisting of a well-developed metacarpal; third finger with two phalanges; shoulder girdle highly abnormal, seventh cervical vertebra and first dorsal so completely fused that their boundaries cannot be detected; these fused with first rib; first rib fused with presternum and ventral half of second rib, the region between completely filled with bone, presternum appearing to be enlarged to form a broadly crescentric plate; foot normal, hallux with two phalanges, the other toes with three phalanges; fibula complete, threadlike; pelvis abnormal, ischium and pubis so reduced that dorsal and ventral profiles of innominate bone are nearly pa:r;a;llel, their width so increased that thyroid foramen is scarcely more than twice as large as acetabulum; lumbar vertebrre showing no tendency to become fused, centrum of fifth or sixth, sometimes both, with distinct bifid or double hypophysis; skull without postorbital processes, with premaxillaries represented by lingulate palatal branches only, the two bones, partly cartilaginous, fused, neither with each other nor with maxillaries; palate so deeply emarginate both anteriorly and posteriorly that its median length is less than least distance between tooth rows; teeth normal; ears large, without tragus; muzzle with conspicuous leaflike cutaneous outgrowths consisting of a horizontal horseshoe, a perpendicular median sella, and a posterior erect lancet. (After Miller.) The family contains only the genus Rhinolophus Lacepede. Genus RBINOLOPBUS Lacepede

Rhinolophus LACEPEDE, Tab. des Div. Sousdiv. Ord. et Gen. des Mamm. (1799) 15; DOBSON, Cat. Chirop. Brit. Mus. (1878) 100; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 108; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 13. Aquias GRAY, Proc. Zool. Soc. London (1847) 15. Phyllotis GRAY, Proc. Zool. Soc. London (1866) 81 (non Waterhouse). CtBlophyllus GRAY, Proc. Zool. Soc. London (1866) 427. Euryalu8 MATSCHIE, Sit~b. Ges. Natur. Freunde Berlin (1901) 225.

212

PHILIPPINE LAND MAMMALS

Ears large, without tragus; antitragus present; nose leaf conspicuous; tail well developed; upper jncisor small, but usually well formed, with distinct -rounded crown with slight cusp on inner side or distinctly bifid; lower incisors trifid with outer usually larger than inner, the four teeth forming a continuous row between canines; upper canines heavy, simple in form, without ~.econdary cusp or conspicuous cingula; lower canines rather weak; first upper premolar and second lower premolar small, functionless, usually crowded out of tooth row; other teeth without special peculiarities; first and second upper molars without hypocone, third with five cusps and three commissures; in most species a rudimentary fourth; lower molars with cusps all well developed and normal in position, Skull with distinct sagittal crest, small audital bullre, and large cochlea; basisphenoid pits absent. Other characters as in the family. . 1- 1 1- 1 2- 2 3- 3 Dental formula: 1. 2- 2' c' _I - I' pm. 3- 3' m. 3- 3 = 32. K ey to the P hilippine speoies of Rhinolophu s L acepede. a 1 . Base of sella with inter nasal lobules forming a cuplike expansion. R. philip pinensis WateThouse (p. 214). a 2 . Base of sella normal. b ' . EaTS longeT than half the forearm; sella heavily furred on anterior face ........ _._ ... __ ...... .. ____ .... __ ........... -............ R. Mi"sutus Andersen (p. 215). b 2 . Ears less than half the forearm; sella without or with only exceedingly sparse hair. c ' . Sella with sides slightly concave or parallel. d t. Length of forearm, about 38 millimeters; fiTSt phalanx of fouTth finger greater than one-eighth of fourth metacarpal. R. virgo Andersen (p. 216). d ' . Length of forearm, about 45 millimeters; first phalanx of fourth finger about one-eighth of fourth metacarpal. e '. Metacarpals graduated in length. R. ande?'senianderseni (Cabrera) (p.219). e 2 . Metacarpals about equal, skull larger. R. anderseni mqualis Allen (p. 220). 0 2 â&#x20AC;˘ Sella ovulate with convex borders; connecting process from top of sella. d ' . Apex of sella normal. e '. Length of forearm, 53 to 57 millimeters; widt.h of horseshoe, 11 to 12 millimetel's; oolla with long hair on front face. R. subrufus Andersen (p. 221). e 2 . Length of forearm, 44 to 47 millimeters; width of horseshoe, 9 to 9.8 mi~limeters; nasal protuberances of cranium more than 5.5 millimeters, R. a7'ouatus a?'ouahts (Peters) (p. 224).

RHINOLOPHUS

213

e 8. Width of horseshoe, 7 to 8 millimeters; nasal protuberances less than 5.5 millimeters. R. arcuatus exiguus Andersen (p. 227). d ' . Apex 'of sella forming a triangular pocket. R. inops Andersen (p. 228). c". Sella ovulate with concave borders; no longer hair on front face of sella; connecting process rises below summit of sella; forearm, 56 millimeters ............................ R. bunkeri sp. nov. (p. 228) .

Andersen ' has divided the genus into six major groups, four of which occur in the Philippines. These are the simplex group, to which R. virgo and R. anderseni belong; the philippinensis group, to which R. philippinensis belongs; the macro tis group, which includes R. hirsutus; and the arcuatus group, to which belong R. arcuatus exig'uus, R. arcuatus arcuatus, R. inops, R. subrufus, and R. bunkeri. Eydoux and Gervais * mentioned Rhinolophus luctus rufa, a bat from Manila which is said to be like R. luctus Temminck (the leaf exactly the same), the color differing however in that it is "roussattre," hence the name "rufa." The description is such that it cannot be applied to any known bat. Andersen t has shown that no bat of the R . luctus type has before or since (1839) been found in the Philippines and suggests that it may be an earlier name for R. philippinensis which Waterhouse described in 1843. It might even more probably apply to R. ?"'U fus of Peters, which certainly does occur at Manila and, as far as the color goes, would agree with the "roussattre" color of the bat found by Eydoux and Gervais. If such can be pr oved by an examination of the type then the bat known as R. subrufus Andersen (R. rufus Peters), will be R. 1'ufus Eydoux and Gervais. The numerous records of Elera:t of R. luctus for the Philippines (Luzon, Paragua, Leyte, and Mindoro) may be disregarded. Hollister has retained the name R. luctus, as well as the varietal name, in his list, presumably on the authority of Elera. There is not and, probably, never has been a bat of this species in the collection of the University of Santo Tomas, Manila, which collection purports to form the basis of Elera's works.

* Voyage

autour du monde la Favorite 5 pt. 2 (1839) 9. & Mag. Nat. Hist.路 VII 16 (1905) 252. :j: Contribucion a la Fauna Filipina (1915) 82.

t Ann.

214

PHILIPPINE LAND MAMMALS RHINOLOPHUS PHILIPPINENSIS

Waterhouse

Rhinolophus philippinensis WATERHOUSE, Proc. Zool. SOC. London (1843) 68 (Luzon); PETERS, Monatsb. Acad. Wiss. Berlin (1871) 305 (Luzon); DOBSON, Cat. Asiat. Chirop. (1876) 43; Cat. Chirop. 118

120

122

124

126

Batan Is

PHILIPPINE ISLANDS

20 "

~Il

® Megadepma spasma spasma @

Rhinolophus philippinensis

® Rhinolophus a. anderseni ® Rhinolophus a. <!!9ualis

"<:

~ ~ "'1;

.... ~ 10

o·

<>Cagayan Sulu

122

FIG. 9.

124·

126

Distribution of Meg'ade1'ma and five species of Rhimolophu8 in the Philippines.

Brit. Mus. (1878) 107; ELERA, Contribuci6n a la Fauna Filipina (1915) 80; ANDERSEN, Ann. & Mag. Nat. Hist. VII 16 (1905) 243, figs. 1 and 2; 24'5,651; HOLLISTER, Philip. Journ. Sci. § D 7 (1912) 13; Proc. U. S. Nat. Mus. 46 (1913) 335 (Luzon, Mindanao). Phyllotis philippensis GRAY, Proc. Zool. Soc. London (1866) 81.

215

RHINOLOPHUS

Description of the species.-Ears large, subacute, upper third of outer margin slightly concave; antitragus large and deep; horseshoe-shaped nose leaf broad, concealing muzzle; base of central leaf expanded, with turned-up edges, forming a deep cup between and above nostrils; sella broad, not diminishing in width upwards, truncate above; posterior vertical connecting process convex on upper margin, not so high as summit of sella; lancet long, narrow, and acute; lower lip with three grooves; wing membranes attach at ankles; interfemoral membrane square behind, extreme tip of tail projecting. Color.-Fur above and beneath dark brown, slightly paler beneath. Skull and teeth.-Sagittal crest low in front, sloping gently toward postnasal depression; openings of audital bullre large; brain case width greater than zygomatic width; mastoid and zygomatic width nearly equal; nasal swellings large; palatal bridge more than one-third as long as maxillary tooth row, the median anterior part opposite front of last premolar; small upper premolar invariably in tooth row. Measurements of the t'!jpe of Rhinolophus phUippinemis Waterhouse. mm.

Length, head and body (in alcohol) Tail Ear Antitragus Forearm Thumb Second finger Fourth finger Tibia Foot and claws

49.5 28 25.4 12.2 by 7.6 47 8.1 68.5 53.3 21.6 8.9

Remarks.-The type, in the British Museum of Natural History, was collected in the Philippine Islands by Hugh Cuming. The species has been reported from Luzon and Mindanao, but the exact type locality is unknown. I have been unable to examine a specimen. RHINOLOPHUS HIRSUTUS

Andersen

Rhinolophus hirsutu8 ANDERSEN, Ann. & Mag. Nalt. Hist. VII 16 (1905) 289; 651 (Guimaras); HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 14; Proc. U. S. Nat. Mus. 46 (1913) 308.

Description of the species.-Belongs to Andersen's macrotis Nose leaves large, horseshoe broad; sella tongue-shaped,

group.

216

PHILIPPINE LAND MAMMALS

parallel margined, front surface covered with long dense hair; connecting process rounded, projecting slightly, beginning at a point below sella; lancet cuneate; lower lip with three mental grooves; ears large and broad; tail one-third longer than tibia. Skull and teeth.-Skull shaped as in R. macro tis, larger; middle lower premolar half in row; first upper premolar in row, with a distinct pointed cusp. Measurements of the type of Rhinolophus hirsutus Andersen. rnm.

Forearm Ear length Ear width Tail Femul'

44.7 24 18 24

Remarks.-The type, in the United States National Museum, was collected on Guimaras Island by J. B. Steere. No other specimen is known. RHINOLOPHUS VIRGO Andel'sen

Rhinolophus virgo ANDERSEN, Proc. Zoo!. Soc. London (1905) 88 (Pasacao, South Camarines, Luzon); Ann. & Mag. Nat. Hist. VII 16 (1905) 654; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 13; Proc. U. S. Nat. Mus. 46 (1913) 308.

Description of the species.-(From No. 351, E. H. Taylor collection; collected at Saub, Cotabato, Mindanao, April 20, 1923, by E. H. Taylor.) Adult female. Small rhinolophid belonging to the simplex group of the genus; ears large, tips reaching beyond tip of snout, anterior edge slightly rounding, tips directed outward, strongly concave behind; antitragus separated by a deep notch from rest of ear; nose leaf covering snout, horseshoe divided anteriorly; an indication of a second leaf below outer edge of horseshoe; sella with sides nearly parallel, top of anterior face rounded, lower part of anterior face widest; at base of sella anteriorly a small rounded platform, the edges turned up, connecting anteriorly by a narrow bridge with horseshoe; lancet elongate, broadly triangular, rising higher than sella, forming three small pockets on each side of anterior face and a slight indentation near its base posteriorly; lower lips with three grooves; fur long, not especially thick; horseshoe and sella naked, lancet with numerous hairs; upper lips with short hair; wing membranes above almost entirely naked; below inner membranes of wings and interfemoral membrane with sparse white hair; wing membranes attach low on sides of body, and to foot at a distance of about 1 millimeter from end of tibia;

RHINOLOPHUS

217

calcar well developed, supporting a very ample interfemoral membrane; tail long, penetrating edge of membrane and proj ecting beyond less than 1 millimeter; feet slender and delicate; antebrachial membrane attaches at small pad on thumb; a second membrane anterior to second finger attaches at same place; second finger distinctly free from third, and separated by a membrane at least 4 millimeters wide; head and body approximately 44 millimeters long. Skull and teeth.-Skull with a low sag~ttal crest running forward and hranching, the branches about 1 millimeter long and ceasing at base of rounded basal swelling; interorbital constriction narrow; audital bul1re small on large snail-like cochlea; palate deeply emarginate anteriorly and posteriorly so that its median length is greatly shortened; premaxillary narrow, greatly elongated, bearing two upper incisors on lateral edges, anterior to canines and at some distance from tip of premaxillaries; second upper premolar small with cusp pointing inward, and distinctly separated from following tooth; small lower premolar not crowded but standing out of tooth row touching only anterior premolar; lower canines with prominent anterior cingulum; anterior lower incisors trilobed, forming a semicircle, two anterior in contact, each separated from second incisors; second pair larger than first, separated from canines . . 1- 1 1- 1 2- 2 3- 3 Dental formula: 1. 2- 2' c. 1- 1' pm. 3- 3' m. 3- 3 = 32.

Variation.-The significant differences between my specimens, from Mindanao and J 010, and the types, from Camarines, are the longer metacarpals and phalanges and the shorter nose leaf. These differences are scarcely greater than might be expected in a series larger than was available to Andersen. There is some difference in the position of the middle lower premolar; sometimes it is external to the tooth row, sometimes partly included. In No. 375, a female from Jolo, the l.J.pper incisors are strongly bilobate; in the Mindanao specimen the tooth shows only a single cusp. The canines are distinctly farther apart and have a thicker base than in the Mindanao specimen described. The color is distinct reddish to brownish orange, similar to Andersen's third phase of R. borneensis. This single specimen was collected on Bud Daho, a low volcanic crater in the eastern part of Jolo. It was shot in daylight, having been startled from a hiding place in a mass of leaves in a net of vines.

218

PHILIPPINE LAND MAMMALS

Measurements of Rhinolophus virgo Andersen. [Measurements in millimeters.]

- - - - - - - - - - - , - - - - - - - -.---- Collection of E. H. Taylor, No. and sex.

375, female.

From Andersen . \ type and cotype, South Camarinea, Luzon.

351 , female.

Minimum.

1--------------------1------ Total len gth .. __ _ ___ _ _ ____ __ _ __ ___ _ _ _ __ _ __ _ _ __ __ _ _ _ __ _ Ear lengt h ___ ___ - - - _ ___ __ _ _ _ __ _ _ _ _ _ __ _ _ _ _ _ _ __ _ _ __ _ _ _ _ Nose lea ves, total length . __ ____ __ ___ _____ __ _ .__ _ _ ___ _ _ _ Breadth of horseshoe_ _ __ ___ _ __ _ _ __ _ _ _ __ __ ___ _ ___ _ __ _ __ Forearm_ ___ _ _ __ _ _ _ _ __ _ _ __ ____ _ _ _ __ __ ___ _ _ _ ___ _____ __

65 14.8 8.5 7 37 . 2

66 15.3 10 7.2 39

Third finger: MetacarpaL__ _ _ _ __ _ __ _ _ _ ___ _ _ _ _ __ ___ _ _ ___ _ _____ __ First phala nx. _ _ _ _ __ _ _ _ _ __ __ ___ _ __ _ _ __ _ _ _ _ __ __ _ _ __ S ~o nd phalanx . ___ _ ___ _ _ _ _ _ __ __ _ _ __ _ _ __ __ __ _ __ _ _ _

27 11 16

Maximum.

- - - --

------- - '-------14.7 10.7 7.2 37.5

16 . 2 11 . 2 7.2 38.8

28.6 11 17

27.2 10 . 2 15 . 2

28.2 10 . 2 15.2

28 8 9

30 8.5 11 . 5

28 7.3 9

28.6 8.2 9

27 8 .7 9 20 15

29 .5 9 11 20.2 15.2 7.8

27 8.1 8.2 17.9 14.2 7.2

28.2 8.8 . 8.3 20.2 15.2 8

16.2

16 . 9

Fourth finger:

L ower leg__ _ _ _ __ __ ___ __ _ _ ___ _ _ ___ __ _ __ _ _ _ __ _ _ _ _ __ _ _ _ _ Foot_. _________________________________ ~_____________

Skull: T ot al length_ ____________ __ ____ ____ _____ __ _______ 17.5 Interorbital constriction __ _____ ___ _______ ________ ___ __ ____ _ 8 M astoid widtb . __ _ _ _ _ __ __ _ _ _ _ _ ___ _ _ _ _ _ _ _ ___ _ _ _ __ _ _

Width of brain case _____________ _____ __ ______ ___ __ Zygomatic width . __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ Supraorbital length _ __ _____ __ _ ____ __ __ _____ _ ______

9 8.8 4.8

Width of nasal swellings _____________ ______________ Mandible, length . _________ ______________ ____ . _ __ _ _

4.1 11.2

~ idth

between canin es _ - - - - - - - -- -- - - - - --- - - - - - - - - - - - - - ---U pper t eeth _ _ _ _ _ _ _ _ _ _ __ _ _ _ _ _ _ __ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ _ 6. 6 6.8 L ower t eeth _. ___ _______ ______ ___ . _ __ __ ___ _ _ __ __ _ _

16.8 3 8 8 8.1 4.6 4.3 11.8 2 6.4 7

- ----- -- - ------8 7.1 8.1 4.7 4 _3 10.8

8.2 7 .7 8.2 6 4.3 11.6

--------~ 6.1 6 .2 6.5

6.8

Remarks.-The types, in the United States National Museum, were collected in Pasacao, Camarines Sur, Luzon, by L. M. McCormick. The relationship of this species is with Rhinolophus borneensis Peters, from which it differs chiefly in size. Both of these forms belong to Andersen's simplex group of the genus, and this is the smallest known form. The described specimen, No. 351, was obtained from a dark cavity formed by

219

RHINOLOPHUS

the buttress of a tree in a dense forest near the seacoast, at Saub, Cotabato, Mindanao. RHINOLOPHUS ANDERSENI

Cabrera

This species has two recognized subspecies, the typical form of which is from an unknown locality in the Philippines. The other, R. a. tJ3qualis, is from Palawan. RHINOLOPHUS ANDERSENI ANDERSENI

(Cabrera)

Rhinolophus anderseni CABRERA, Bol. ,de la Real Soc. Espanola de Hist. Nat. (1909) 305; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 13; Proc. U. S. Nat. Mus. 46 (1913) 335.

Description of the species.-TaiI elongate, approximately equal to tibia; nasal appendages large, horseshoe very wide, covering snout completely when seen from above; borders of sella practically parallel, projections high and slightly convex (as in all species of the simplex group), drawn to a point somewhat higher than ordinary length almost to apex of sella; second phalanx of third digit more than one and one-half times length of first phalanx; first phalanx of fourth digit about onefourth metacarpal of same digit; wing membrane attached about 1 millimeter above ankle. . Color.-Dirty reddish brown above, shading slightly to a somewhat darker reddish between shoulders; paler below. Skull and teeth.-Brain case similar to that of Rhinolophus nereis, but facial outline region entirely distinct; nasal bones flattened; second premolar very small; third lower premolar outside tooth row; the other premolars in contact with each other inside. Measurements of the types of Rhinolophus anderseni ande?'seni (Cab?"em) cmd R. a. ;;equalis Allen. Rhfntolophus <lh\derseni anderseni.

mm.

Head and body Ear: Length Width Width of horseshoe of nose leaf Forearm

Rhinolophus IIInderseni

requalis. mm.

18 14.5 10 45

8 45

220

PHILIPPINE LAND MAMMALS

Measurements of the types of Rhinolophus anderseni anderseni (Cabrera) and R. a. requaZis Allen-Continued. Rhinolophus andeTseni anderseni. mm.

Third finger: Metacarpal l"irst phalanx Second phalanx Fourth finger: Met.acarpal First phalanx Second phalanx Fifth finger: Metacarpal First phalanx Second phalanx Tail Tibia Foot Skull: Total length Mastoid width VVidth of brain case Zygomatic width VVidth of nasal swellings Upper tooth row with canine Mandible; condyle to front of incisors Lower toOith row with canine

Rhinolophu8 a11.der8eni requalis.

30.5 12 21

34 13 17

32'

34.5 9

8 13

32.8 9.5 12 21 20 10

34.510 10.5 21.5 20.5 9

20 9 8.3 9.2 5.5 8 12.3 8.5

21 10 10.5 8.2 10

Remarks.-The type, an adult male in the Museo de Madrid, was received as a gift from Sr. Uhagon. The exact locality and collector are no longer known. It is, .however, from the Philippines, probably Luzon. I have seen no specimen of the species. RHINOLOPHUS ANDERSENI lEQUALIS Allen

Rhinolophus andM"Seni 33qualis ALLEN, Occ. Papers Mus. Zoology Univ. Michigan 110 (1922) 1-5 (February 25).

Description of the species-(After the type description.) Similar to Rhinolophus anderseni anderseni, but with a slightly larger skull and metacarpals slightly longer, nearly of equal length instead of slightly graduated. Color.-No color description is given. Measurements.-See under measurements of Rhinolophus anderseni anderseni (Cabrera).

RHINOLOPHUS

221

Remarks.-This subspecies was founded on No. 53112, Museum of Zoology, University of Michigan, from Puerto Prusse * (sic), Palawan, collected by J. B. Steere. RHINOLOPHUS SUBRUFUS Andersen Rhinoloph~ts

?OUf'US PETERS, Monatsb. Akad. Wiss. Berlin (1861) 710 (non Eydoux and Gervais); ELERA, Contrib. a la Fauna Filipina (1915) 83; DOBSON, Monog'. Asiatic Chirop. (1876) 44. Rhinolophus subrufus ANDERSEN, Ann. & Mag'. Nat. Hist. VII 16 (1905) 283; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 14; Proc. U. S. Nat. Mus. 46 (1913) 309.

Description of the species.-Adult male; belongs to the Rhinolophus arcuatus group; distinctly larger than R. arcuatus; horseshoe covering muzzle, of normal shape; secondary leaflet crenulate with well-defined glands on lower side, each bearing a fine vibrissalike hair, two small glands near posterior ends of horseshoe; sella ovate, sides tapering very slightly (at base, 3.4 millimeters; near 路top, 2.9); internasal leaflets with broadly triangular wings; connecting process rises from summit of sella and is gently rounded, not reaching higher than sella; sella with scattered brownish hairs, the longest 1.5 to 1.7 millimeters long; lancet tapering regularly to summit, which rises 5 millimeters above base of connecting process; three pockets on each side of lancet; three pockets formed on posterior (basal) side of sella, the median largest; ears large, anterior margin rounded to near summit, then slightly flattened toward tip; latter subacute; margin distinctly concave for a short distance, and continued convexly to antitragal notch; antitragus large, 11 millimeters wide at top; a blind pocket at base of antitragus opening from outside, forming a noticeable fold on inner side.of base of tragus; lower lip with three grooves forming four unequal calloused areas; tail somewhat shorter than tibia, tip projecting 3.5 millimeters; wing membranes attach at least 2 millimeters above distal end of tibia; sides of muzzle heavily furred; top of connecting process, area between process and lancet, and sides of lancet with long, thinly distributed hairs; body above and below densely furred; a short, noticeable growth of hair on interfemoral membrane, more prominent on outer edge near tail; hair on feet and toes short. Color in life.-Fur .on head reddish, on back cinnamon-rufous, the base of hairs orange-rufous; below similar to back, slightly more brownish red; wing membranes blackish brown.

* Probably

an incorrect spelling of Puerto Princesa.

222

PHILIPPINE LAND MAMMALS

Skull and teeth.-Nasal swellings prominent, rounded, followed by a distinct depression; supraorbital crest high anteriorly, falling gently to supraorbital ridges; ridges low and divided, a distinct depression between them; brain case slightly narrower than zygomatic width; cochlea large, bul1re small; palatal length 4 millimeters. MeaSU1"ements of Rhmolophus sub?"Ufus Andersen and R. bunkeri sp. nov. [Measurements in millimeters.] E. H. Taylor collection No. and sex. Rhinolophus bunkeri.

Rhinolophus 8ubrufUSl.

321, male.

Total length _________________ \ _________ \ __ ~ Head and body ... ________________ ___ _________ Tail.. _________ . _____________________________

334, male.

359, female.

382, female.

--- --- --- ---

367, temale. ~--

83 62 24

84 62 22

85 63 22

83.8 60 23.8

90 67 25

23 19 12 11 .5 53 10 42

23 20.5 12.1 11.2 55 9 41

23 20 12 10.5 56 10 41

23 19 11.8 9.9 55 8.9 41

25.5 19.5 12.5 9.8 56 10.2 42

37.5 16 26

37.2 15.5 26

38.5 16 26

38.5 16 28

38.2 17 20

41 11 16

39.6 11

39 10.6 15

15.5

39 12 16

40.5 12 10 25 13.5

40 12 12 27 13.2

42 13 12 27 13.5

40.5 12.5 12.8 26 12.2

41 13 15 27 15.2

25 .2 22 11.4 12.5 5 17.2 10.6 10.5

26 22 11.2 11.5 5 17.5 10 10.5

24 23 11.2 11.6 5 18 10 11

25.3 23 12 13.8 5 19 10.2 11

Ear:

Length __________________________________ Width_ .. __________________________ ______ Breadth of horseshoe. _________________________ Height of lancet. ___________________________ J Forearm路. ______________________________ ____ r

First finger with claw _________________________ Secondfinger ________________________________ Third finger: ~etacarpal------------------------------

First phalanx- ____________ _______________ Second phalanx __________________________ Fourth finger:

__________________________ First phalanx- ___________________________ Second phalanx _____._____________________ ~etacarpal----

Fifth finger: ~etacarpal------------------------------

First phalanx ____________________________ Second phalanx __________________________ Tibia _______________________________________ Foot and claws .. _____________________________

Skull: Total length _____________________________ Condylobasal length ___________________ .___ Brain case, width through bullm ____________ Zygomatic width _________ .________________ Nasal swellings, width .. ___________________ ~andible, length _________________________ Upper tooth row with canine. ______________ Lower tooth row with canine. ______________

---------------------12 5 17.6 10.1 10 . 8

RHINOLOPHUS

223

Upper canines moderately heavy; first upper premolar small, with well-developed cone, the tooth directly in tooth row; upper incisors showing only a single cusp, separated by a distance of 1.5 millimeters; lower incisors strongly trifid, outer pair larger and much thickened; second premolar entirely outside tooth row, reaching to top of cinguI:re of first and third premolars; two latter teeth in contact. Variation.-The four specimens in my collection referred to this species show considerable variation. In dentition No. 334, male, Manila, has the second lower premolar nearly half in the tooth row and the adjoining teeth separated by an appreciable distance, the first upper premolar is decidedly larger, nearly twice the size of the tooth in the described specimen; the middle lower premolar is also much larger than in the described specimen. The fur on muzzle is more prominent, and the hair on posterior part of nose leaves is longer; the tail is somewhat shorter and the color more rufous-orange (perhaps faded somewhat in alcohol) ; the tip of the lancet is less acute and slightly shorter. The skull of No. 359, female, Los Banos, Luzon, has the first upper premolar, and the second lower, nearly the same size as in No. 334; the first and third lower premolars in contact. The pelage of this form is more distinctly reddish than the described specimen. In No. 382, female, Baguio, Benguet, Luzon, the first upper premolar is about the size of that in No. 321 (Los Banos, Luzon), but the upper incisors are much smaller and nearer tip of the premaxillary; the middle lower premolar is identical in shape and position with that in No. 334. The specimen is dark brown, showing no trace of the reddish or orange color of the other three specimens; the sella is noticeably narrower and the tip of the lancet blunter than in the other specimens. For variations in size, refer to table of measurements. Remarks.-The type of Peters's Rhinolophus rufus is from Paracale, Luzon; the type of Andersen's subrufus is from Manila. Andersen has shown that the name "rufus" is preoccupied by R. luctus rufa Eydoux and Gervais. It is not at all improbable that the latter species is identical with the species here described; if such is proved the name "rufus" must stand. My specimens were captured while they were flying at night in houses.

224

PHILIPPINE LAND MAMMALS

RHINOLOPHUS ARCUATUS ARCUATUS

(Peters)

Rhinc;lophus a1'Cuatus PETERS, Monatsb. Konig!. Preuss. Akad. (1871) 305; ANDERSEN, Ann. & Mag. Nat. Hist. VII 16 (1905) 281. Rhinolophns aJ'cuatus typica ANDERSEN, Ann. & Mag. Nat. Hist. VII 16 (1905) 651. Rhinolophus a?'cuatus arcuatus HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 14; Proc. U. S. Nat. Mus. 46 (1913) 309.

Description of the species.-(From No. 155, Bureau of Science collection, collected at Irisan, Benguet.) Adult male. Small, horseshoe leaf on nose broader than upper jaw, divided anteriorly by a deep groove which is bordered by two slight, elongate, calloused ridges; a groove following outline of horseshoe, about 1 millimeter from edge; internasal lobules at base of and anterior to sella strongly angular, with a rounded point at tip; sella broadly ovate, widest at base; connecting process attached to summit of sella arcuate, upper edge forming a semicircle; posterior lancet erect with two pockets on each side of anterior surface formed by septa, third pocket on each side at base not fully surrQunded; a single thickened crenate-edged 'supplementary leaf below edges of anterior horseshoe; on posterior surface three shallow pockets formed by two lateral septa; eyes at lower extremities of two outer pockets; no glandular tuberc.1es behind lancet; ears large, anterior portion rounded at base, continuing to near summit with a straight edge; summit with an ovate tip, posterior to which the edge of ear continues in a straight or slightly curved line to the deep notch which separates the antitragal lobe; lower lip with a naked calloused area with three grooves; wing membranes attaching 'low on sides, and to tibia at a distance of 1 millimeter from lower end; tail short, projecting free for about 1.3 millimeters; interfemoral membrane scant; body densely furred above and below; upper anterior nose leaf and anterior part of sella naked; connecting proCiess and lancet covered with sparse elongate hair; ears nearly naked; wing membranes with sparse fur near body. Color in alcohol.-Above slightly reddish brown, growing lighter toward base; sparse fur on upper lip nearly white; wing membranes brown; belly buff to gray brown. Teeth and skull.-Molars and premolars normal for the group; first upper premolar very small, wholly in tooth row; upper canines some distance back from tips of premaxillaries, crowns with two cusps; cingulum represented on anterior part of upper canine by a slight projection; no posterior cusp; second lower premolar small, 路 crowded entirely out of tooth row but rising above cingUla of other premolars, slightly tricuspid (in a second

225

RHINOLOPHUS

specimen, No. 156, Bureau of Science collection, the tooth is much smaller and does not rise above cingulum) ; lower incisors trifid, forming a continuous row, not overlapping and not in contact with canines; two outer teeth distinctly larger than liB

120'

122 Balan Is

1<'4

126

PHILIPPINE ISLANDS

® Rhino lophus a. exiguus

-.;)

4J ~

.....

[.(W)

c>Cagayan Sulu .'

' . cQ-Ba9ilan

~jolo

d'~

•. lJ:)

~Tawitawi

Sibutu :) '/, "ngau 120

FIG. 10.

122

124 0

126

Distribution of five species of Rhi'nolophU8 in the Philippines.

inner; median anterior nasal swellings large, semiglobular, and abruptly projecting; palatal bridge very short, one-fourth to one-third as long as maxillary tooth row; basioccipital narrow; temporal fossa narrow, zygomatic and mastoid breadths nearly equal; sagittal crest almost obsolete; supraorbital ridges separat259453-15

226

PHILIPPINE LAND MAMMALS

ed by a groove; brain case equal to zygomatic width or greater; cochlere large, close together . . }- 1 1-1 2-2 3-3 Dental formula: 1. 2-2' c. 1- 1' pm. 3-3' m. 3- 3 = 32. Measurements of Rhinolophus arcuatus arcuatus (Pete?路s). Bureau of Science No. l'55, No. 156, male. female. mm. mm.

Total length Head and body Tail Ear: Length Width Breadth of horseshoe Height of lancet Forearm First finger with claw Second finger Third finger: Metacarpal First phalanx Second phalanx Fourth finger: Metacarpal First phalanx Second phalanx Fifth finger: Metacarpal First phalanx Second phalanx Tibia Foot Skull: Total length Condylobasal length Brain case, width Zygomatic width Width of nasal swellings Mandible length Upper teeth with canine Lower teeth with canine

61.5 46 15.5

62 45 17

18.8 15 9.8 8 45 8 32.5

19 14+ 9.4 7.5 44 7.8 32

29 11.8 18

29 12 18

30 8.6 11

29 8 11

31 9 9 21 9.5

30 9.2 10 20 9.5

23 20 10.2 10 44.8 14.3 8.5 8.6

22.5 19.8 10 9.7 4.5 13.8 8.2 8.3

Remarks.-The species was described by Peters from a Luzon specimen (the exact locality not stated). Hollister reports specimens from Irisan, Benguet; Montalban, Rizal; and Pasacao, South Camarines, in Luzon; and Mount Halcon, Mindanao (error for Mindoro). The form inhabiting the southern islands

227

RHINOLOPHUS

has been referred to the subspecies Rhinolophu.s arcuatus exi-

guus. RHINOLOPHUS ARCUATUS EXIGUUS

Ander~n

Rhinolophus arcuatus exiguu s ANDERSEN, Ann. & Mag. Nat. Hist. VII 16 (1905) 283, 651; HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 14; Proc. U. S. Nat. Mus. 46 (1913) 309.

Description of the subspecies.-Similar to typical Rhinolophus arcuatus, from Luzon, but horseshoe narrower, nasal swellings smaller, fifth metacarpal a trifle longer than fourth, and wing membrane inserted 1 to 2.5 millimeters above ankle. Color.-Dark brown, as in the dark form of the typical Luzon species. Skull and teeth.-Skull slightly shorter, width of nasal swellings less; third premolar external to tooth row, or wanting on one or both sides; anterior and posterior premolars strongly in contact, their cingula overlapping each other; otherwise as .in the typical form. Minimum a,nd maximum m easurements of exiguus Andersen.

Rhino~ophus

Minimum. mm.

Ear: Length Greatest breadth Nose leaves: Length Breadth of horseshoe Forearm Tail Lower leg Skull: Total length Mastoid width Zygomatic width Width of nasal swellings Mandibles Upper teeth Lower teeth

ar cuatus Maximum. mm .

17.2 13

19.7 14

13.8 7.7 44.8 15 20

14 8.2 46.2 16.2 21

20 9.5 9.5

20.8 9.8 9.9 5.2 14.2 8 8.7

5 13.4 7~ 9

8.5

Remarks.-The type of the subspecies, in the British Museum, was collected in Zamboanga by A. Everett. Andersen gives Guimaras as part of the range. Hollister * lists five specimens from Guimaras in the United States National Museum. I have collected no specimens of this subspecies. 'T he typical form, R. arcuatus arcuatus, appears to be confined to Mindoro and Luzon. * Proc.

U. S. Nat. Mus. 46 (1913) 309.

228

PHILIPpINE LAND MAMMALS RHINOLOPHUS INOPS Andersen

Rhinolophus inops ANDERSEN, Ann. & Mag. Nat. Hist. VII 16 (1905) 284, 651; HOLLISTER, Philip. J ourn. Sci. ยง D 7 (1912) 15; Proc. U. S. Nat. Mus. 46 (1913) 309.

Description of the species.-Intermediate in size between Rhinolophus arcuatus and R. subrufus; differs from both in that summit of sella forms a triangular pouch turning the opening downward'; horseshoe, connecting process, and lancet as in R. subrufus; greatest depth of pouch 1.5 millimeters; wing membrane inserted 1 millimeter above ankle. Color.-Entire body dark brown. Skull and teeth.-Skull of the arcuatus type but with a slightly wider temporal fossa; sagittal crest somewhat higher in front and a little more abruptly descending towards postnasal depression; middle lower premolar external to tooth row; first and third premolars in contact. Dental formula as in arcuatus. Measurements of the type of Rhinolophus inops Andersen. mm.

Forearm Ear: Length Breadth Nose leaves, length Horseshoe, breadth Tail Lower leg Skull: Total length Mastoid width Zygomatic width Width of nasal swellings Mandibles Upper teeth Lower teeth

53.8 23.3 18.3 ?18.8 12.7 16 24.7 24.2 11 11.5 6.5 17.5 10 10.9

Remarks.-The type, in the United States National Museum, was collected at Todaya, Mount Apo, Davao, Mindanao, at an elevation of 1,325 meters by Edgar A. ,Mearns. He gives the Bagobo name kohs' -set. The form may be easily differentiated by the sella, which is unique for the whole genus. I have collected no specimen of the species. RHINOLOPHUS BUNKERI sp. nov.

Type.-No. 367, E. H. Taylor collection; collected April 26, 1923, at Saub, Cotabato, Mindanao, by Edward H. Taylor. Alcoholic specimen with skull separate.

HIPPOSIDERIDlE

229

Diagnosis; description of the type.-Adult female. A member of the Rhinolophus arcuatus group, related to R. subrufus Andersen. It differs from that species in the body length being greater, wider across shoulders; the head distinctly larger and wi.der. Nose leaf of the same general pattern but with the following differences: Horseshoe somewhat wider, scarcely as long; sella without elongate hairs on front face, narrower, sides strongly concave, connecting process rising from a point below summit of sella; internasal lobules slightly less developed; lancet suddenly narrowed at a short distance from rather acute summit; secondary outer leaflet much thickened, glands bearing vibrissre on outer edge rather than on undersurface; naked area of upper lip of greater extent, ear higher and slightly narrower; forearm and tibia no gr-eater than in subrufus; foot distinctly larger and less prominently haired; second phalanx of third finger very strikingly reduced; second phalanx of fifth finger distinctly elongated. Color.-Dark drab to Prout's brown, somewhat similar to the dark phase of Rhinolophus subrufus. Skull and teeth.-Skull more m짜sive, zygoma wider, mastoid width greater, anterior part of brain case more inflated, posterior part less inflated than in R. sub rufus ; molars, above and below, much larger, first upper premolar very much smaller than in any specimen of R. subrufus I have examined; cochJere larger and farther apart; second lower premolar entirely out of tooth row, does not reach summit of cingula of adjoining teeth; the latter strongly overlapping (this tooth absent on right side). Measurements.-See table under Rhinolophus subrufus. Remarks.-The type was collected from a cavity in a buttress of a forest tree near the seashore at Saub, Cotabato. The species is named for Mr. C. D. Bunker, curator of birds and mammals, Dyche Museum, University of Kansas, who has shown me innumerable courtesies, has housed my collection, and greatly facilitated the work on this monograph. Family HIPPOSIDERIDlE Hipposideridre MILLER, Bull. U. S. Nat. Mus. 57 (1907) 109.

(Many recent authors have regarded this as a part of, or a subfamily of, the Rhinolophidre.) Like Rhinolophidre, but pectoral and pelvic girdles more highly modified, toes with two phalanges each, lumbar vertebrre showing a marked tendency to become fused into a solid rod; in pectoral girdle first and second ribs fused, involving the entire

230

PHILIPPINE LAND MAMMALS

bone to and including the corresponding dorsal vertebrre; there is thus produced a solid ring of bone consisting of seventh cervical vertebra, first and second dorsals, first and second ribs, and entire presternum, the elements of the ring indicated by a slitlike vacuity above, between ribs, and one or two small roundish vacuities below; pelvic girdle like that of the Rhinolophidre posteriorly, but anteriorly with a supplemental bridge of bone connecting acicular process in front of ilium and producing a preacetabular foramen slightly larger than thyroid foramen; lumbar vertebrre in many species so completely fused that the 'boundaries between the centra are quite obliterated; no hypophyses. (After Miller.) The family includes nine genera, of which Hipposideros and Chilophylla are Philippine. Members of the Rhinolophidre and Hipposiderid:re may be distinguished from each other by the difference in the number of phalanges in , the digits of feet; the former have three, the latter two phalanges. Key to the Philippine genera of Hipposidet路idm. a '.Ears angular at top; horseshoe normal, with terminal leaf low and rounded .......................................................... Hipposideros Gray (p. 230).

a'. Ears abnormally large, rounded; horseshoe scarcely differentiated, posterior part lancetlike .................................. Chilophylla Miller (p. 247). Genus H!lPPOSIDEROS Gray

Hipposideros GRAY, Zool. Misc. (1831) 37; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 110; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 15.

Phyllorhina BONAPARTli:, Icon. della Fauna Ital. fasc. 21 (1837); DOBSON, Cat;. Chirop. Brit. Mus. (1878) 127.

Macronycteris GRAY, Proc. Zool~ Soc. London (1866) 82. Gloionycteris GRAY, Proc. Zool. Soc. London (1866) 82. Speorifera GRAY, Proc. Zool. Soc. London (1866) 82. Rhinophylla GRAY, Proc. Zool. Soc. London (1866) 82, (non Peters). Chrysonycte1'is GRAY, Proc. Zool. Soc. London (1866) 82. Doryrhina PETERS, Monatsb. Akad. Wiss. Berlin (1871) 314 (subgenus of Phyllorhina). Sideroderma PETERS, Monatsb. Akad. Wiss. Berlin (1871) 324 (subgenus) .

Ptychorhina PETERS, Monatsb. Akad. Wiss. Berlin (1871) 325 (subgenus).

Cyclorina PETERS, Monatsb. Akad. Wiss. Berlin (1871) 325 (subgenus).

Thyreorhina PETERS, Monatsb. Akad. Wiss. Berlin (1871) 327 (subgenus) .

Syndesmotis PETERS, Monatsb. Akad. Wiss. Berlin (1871) 329. Hipposiderus BLANFORD, Proc. Zool. Soc. London (1887) 637.

HIPPOSIDEROS

231

Upper incisors very small, but well formed and distinct, rounded crowns with slight cusp on inner side; lower incisors trifid, the outer larger than inner, the four teeth forming a continuous row between canines or two inner placed anterior to outer; upper canines heavy, simple in form, without conspicuous cingulum though frequently with a secondary cusp near posterior base of shaft; lower .canines rather weak; first upper premolar small, functionless, closely crowded between cingula of canine and large premolar or forced outward completely from tooth row, occasionally absent; other teeth showing no special peculiarities; first and second upper molars with main cusps normal and hypocone absent, third usually with five cusps and three commissures, mesostyle and metacone always closely approximated, the latter sometimes lacking, leaving only two commissures; lower molars with cusps well developed and normal in position; skull with distinct though low sagittal crest; bullre small with moderately large cochlere; greatest depth of brain case (bullre not included) at least equal to depth of rostrum including molars; zygOma abruptly: expanded posteriorly, height of expanded portion much less th~n distance from last molar to glenoid fossa; lumba:r; vertebrre not fused; ears free or joined ' with a low band; tail well developed, longer than femur; caudal vertebrre 6 to 8. (After Miller.) The bulk of the species of the family are contained in this genus of which about fifty-five forms are known. The genus resembles Rhinolophus, but the character of the sagittal crest, smaller cochlere, and the absence of the small lower premolar, differentiate the genera. Several species are found in the Philippines. All have conspicuous nose leaves and large ears . . 1-1 1-1 2-2 3-3 / Dental formula: 1. 2-2' c. I-I' pm. 2-2' m. 3-3 = 30. Key to the Philippine species of Hipposideros Gray. a '. Frontal gland present; no secondary nose leaves. b 1. Terminal nose leaf not divided by septa. c '. Wings attached to metatarsus; forearm, 46 millimeters; tail, 25; tibia, 20 ........................ Hipposideros obscurus (Peters) (p. 232). c'. Wings attached to tarsus; forearm, 47 millimeters; tail, 34; tibia, 18 ................................ Hipposideros coronatus (Peters) (p. 234). b 2. Terminal nose leaf with four cells divided by septa. c 1. No dermal leaf in front of frontal gland; wingg attached to tarsus; forearm, 40.6 millimeters; tail, 30; tibia, 17.8. Hipposideros bicolor (Temminck) (p. 236). c 2. A dermal leaf in front of frontal gland; wings attached to end of tibia; forearm, 39 millimeters; tail, 27; tibia, 17. Hipposideros wrighti sp. nov. (p. 237)

232

PHILIPPINE LAND MAMMALS

a', No frontal gland; secondary nose leaves present external to horseshoe; terminal nose leaf with four cells, divided by septa. b 1. Two secondary nose leaves; wing membranes attached to tarsus; forearm, 36 millimeters; tail, 24; tibia, 12.7. Hipposide?'os pygmaJus (Waterhouse) (p. 239). b 2. Two secondary nose leaves; wing membranes attached to tarsus; forearm, 39 millimeters; tail, 30; tibia, 16.5. Hipposideros a.ntricola (Peters) (p. 241). b·. Three secondary nose leaves; wing membranes attached 2 millimeters from distal end of tibia. e 1. Forearm, 83 to 88 millimeters; tail, 43 to 54; head and body length, 94 to 104; ear, 27 to 30. Hipposideres diadema griseus (Meyen) (p. 242). 2 0 • Forearm, 76 to 77.5 millimeters; tail, 35 to 35.8; head and body length, 84; ear, 24. Hipposideros diadema anderseni subsp. nov. (p. 246). HIPPOSIDEROS OBSCURUS (Peters)

Phyllorhina obscura PETERS, Monatsb. Akad. Wiss. Berlin (1861) 707; DOBSON, Cat. Asiat. Chirop. (1876) 73; ELERA, Contrib. Fauna Filipina (1915) 88. Hipposiderus obsoura TROUESSART, Cat. Mamm. Suppl. (1904) 72; HOILLISTER; Philip. Journ. Sci. § D 7 (1912) 16; Proc. U. S. Nat. Mus. 46 (1913) 335.

Description of the species.-{From No. 337, E. H. Taylor collection; collected April 29, 1923, at Saub, Cotabato, Mindanao, by E. H. Taylor.) Adult male. Horseshoe small, not entirely covering muzzle, nostrils at base of a deep depression; internasal septum greatly reduced, circumnarial flaps small; middle leaf with a median vertical ridge, rather indistinct, with two indistinct tubercles, one on each side near summit of leaf, each bearing a vibrissa; posterior leaf not divided into cells by septa; two prominent tubercles behind this leaf, each bearing two elongate vibriss're; a large, exsertile frontal gland; ears moderately large, anterior edge strongly convex, semicircular in lower half, curving to summit; posterior to summit slightly concave for a short distance, then gently curving to notch which limits antitragal lobe; lower lip with two small tubercles separated by a groove; tip of fifth digit somewhat bifid; tail penetrates edge of interfemoral membrane; wing membranes attach to metatarsus; no secondary leaflets outside horseshoe but a distinct raised glandular ridge directly under edge of horseshoe. Color.-Pelage slightly russet brown, median portion of hairs (about one-third of the length) dull, dirty, brownish buff; below, fur grayish brown, base of hair darker; wing membranes blackish brown.

233

HIPPOSIDEROS

Skull and teeth.-Nasal swellings broad, not greatly inflated, upper surface nearly level; sagittal crest highest anteriorly, ending gradually at point of narrowest interorbital constriction where it joins the two supraorbital ridges; latter end in two slightly projecting prominences; posterior part of zygoma flattened and elevated. M easur ements of Hipposide?"os obscurus (Peters) and H. 'JY!jgm33us ( Waterhouse) . [Measurements in millimeters.] E. H. Taylor collection No. and sex. Hippo8idero8 pig-

Hipposidero8 obacurua.

m ,zuB .

Total length ______ ___ __ __~ \ __ _~ _____~ _______ Head and body ___ ___ ____ _____ ____________ ___ TaU . .. ____ ______ ______ __ _____ _______________

336, male.

337, male. ~~-

338, female.

- - -

373, female. ~~-

354, female.

---

70 51 19 5.8 6.5

67 50 17 5.8 6.2

73 51 22 5,6 6.2

75 55 20 5.4 6

63 87 26 6 6.6

Height _________________ __ ___ ____ ________ Width ... ____ _________ _________ __ ________ Forearm .. ___ __ ____ _____ ______ ____ ___ ________ First finger and claw .. __ ___ ___ ____ ____________ Second finger ____ __ ______ __________ _____ _____

17.5 16 41 7 35

17.8 16 42 7 35

17,5 16 41.5 7 36

18 . 2 15 44 7,5 37

14 13.5 39 , 5 6 31.5

Tbird finger: Metacarpal . ___ ___ _____ ________ __ ________ First phalanx ____ __ ____ __ __ ____ ____ ______ Second phalanx __ ___ ___ __ ___ __________ ___

33 13 18

32 . 5 13 17 . 5

33 . 2 13 . 8 17

33 . 5 13 16 . 5

28 12 . 5 16

Fourth finger: Metacarpal. ____ ___ __ _______ ___ _____ _____ First phalanx __ ____ ____ __ _______ ________ _ Second phalanx ______ __ ____ __ _________ ___

33 10 . 5 8

32,5 10 8

32 11 8,5

32 10 9

27 9 7

31 12 10 18 . 5 9 10

3l.2 11.5 9 17 9 9.8

31.8 12 9 19 10 10 .2

31 12 10 .5 18 10 10

25 10 8 14,2 8 7

19 17 11 5.5 7.5 7.8 14

18.8 17 10 . 8 5.3 7.3 7.8 13

5. 6 7. 2 7.8 14

19 16.3 11 5. 3 7.1 7.8 14

13.5 11 . 2 7 4.8 4.9 5. 2 9 .2

Width of horseshoe _______ ___________________ _ Width of posterior leaL _______ __ ________ -- --Ear:

Fifth finger: Metacarpal. ________ _________ __ _________ _ First phalanx _____ ________ ____ ___________ Second phalanx ____________ ___ _____ ___ ___ Tibia _____ _______ ___ ___ __ _____ ____ _____ ___ __ Calcar ___ ______________ ______ _____ ____ ______ Foot . ___ ___________________ _______ __ ________ Skull: Totallength ___ __ ________ ___ _______ ______ Condylobasal length __ _________ ___ _____ ___ Zygomatic width ___ ________________ __ __ __ Width of nasal swellings .. _____ _____ ______ _ Upper tooth series, including canine._. ___ ___ Lower tooth series, including canine... __ __ __ Mandible . . _____ ______________________ ___

234

PHILIPPINE LAND MAMMALS

Upper incisors with secondary cusp scarcely distinguishable; each canine with a broad shallow groove on anterior face, and a slight shelf at its base formed by remnant of cingulum; first upper premolar almost in tooth row, not reaching cingulum of second premolar; canine and second premolar separated, separation very slight on inside of tooth row; lower incisors trifid, the two inner smaller and somewhat in front of outer pair; a small cusp onp'osterior part of cingulum of canines. Variation.-In the females (two skulls) the upper incisors are smaller than in the two male specimens. There is also slight variation in the position of the first upper premolar; sometimes almost in alignment, and sometimes distinCltly out of alignment. There is a tendency to develop a second cusp on the first lower premolar. In the females the lower incisors are more nearly in a continuous semicircular row. R emarks.-The specimen :here described shows some marked differences in size from the type. The head and body measurements are from 5 to 10 millimeters less, and the tail from 5 to 8 millimeters less in length. The forearm is shorter, from 2 to '5 millimeters, in all my specimens. Four of the specimens (Nos. 335, 336, 337, and 338) were taken from a cavity in the buttress of a forest tree near the seashore at Saub, Cotabato; No. 373 was captured in a mine shaft at Baguio, Benguet, Luzon; the last is the largest one of the lot. The males, captured alive, kept extruding and invaginating the frontal gland. It is capable of at least a centimeter's extension. The bottom is covered with elongate hairs. I was unable to detect any peculiar odor emanating from the gland. The type of the species, in the Berlin Museum, was collected by F. Jagor at Paracale, Camarines, Luzon. The species is also known from Dinagat Island and has been reported from Mindanao. HIPPOSIDEROS CORONATUS (Peters)

Phyllorhina cO?'onata PETERS, Monatsb. Akad. Wiss. Berlin (1871) 327; DOBSON, Cat. Asiat. Chirop. (1876) 72; ELERA, Contrib. Fauna Filipina (1915) 89. Hipposidero8 coronata TROUESSART, Cat. Mamm. Suppl. (1904) 72; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912') 15.

Description of the species.-Nose leaf as in Hipposideros diadema griseus, but the concave front surface of terminal leaf

235

HIPPOSIDEROS

not divided into cells by vertical ridges; no secondary leaves on sides of muzzle external to horseshoe; a distinct frontal pore behind nose leaf; ears moderate, rounded above, upper third of Jl8

126

122

120

Batan Is

PHILIPPINE ISLANDS

® Hipposidel'os obscurus @ Hipposidel'os bicolol'

® Hipposidel"os pygmceus

Lubaog~

DCatandu.nes

~Qu~u.

MINDORO

Busuanga ~

Culion~.

: 1••

C">

8uri.~

rillROmbl!l'J~ ~

b"'V s.'i:2yan

ficao

""'

g~o

r i AA

®..

IsJ

IZ'

cu~:ANAY~'~' ~

OUnlBo8n

Cagayan Is J'NEG ROS

~A~N

s' "'Cagayan Sulu 6'

120'

FIG. 11.

122

124'

126

Distribution of five species of HiP1Iosid&rOB in tbe Philippines.

()uter margin slightly flattened as in H. bicolor; wing membranes attach at tarsus; extreme tip of tail free. Color.-Brown above, paler at base of hairs; yellowish brown below.

236

PHILIPPINE LAND MAMMALS

MeasU'fe1nents of the type of Hipposideros eoronatus (Peters). mm.

Head and body Head Tail Ear Nose leaf Forearm First" finger Second finger Third finger Fourth finger Fifth finger Tibia Foot and claws

66 22 34

15 8 47 10 35 70 56 61.7 18 9.8

Remarks.-The type of the species, in the Berlin Museum, was collected in Mainit, Surigao, northeastern Mindanao, by C. Semper. The species is known only from type. Elera's records for Mindanao, Paragua, and Samar may be disregarded. HIPPOSIDEROS BICOLOR (Temminck)

Rhinolophus bieolor TEMMINCK, Monog. Mamm. 2 (1841) 18. Phyllorhina bieolor PETERS, Monatsb. Akad. Wiss. Be~lin (1861) 709; DOBSON, Cat. Asiat. Chirop. (1876) 70; ELERA" Contrib. Fauna Filipina (1915) 89. Hipposiderrus bieolor TROUESSA,RT, Cat. Mamm. Suppl. (1904) 72; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912). 15.

Description of the specie'S.-Ears as long as head, broad, lower half of inner margin very convex; summit of ear couch rounded off broadly as far as a point on outer side where a slight but distinct flattening occurs and indicates position of tip; a distinct fold, delimiting an antitragal lobe; horseshoe small, square, scarcely as wide as transverse terminal leaf; the concave front surface of horseshoe js divided into four cells by three distinct vertical ridges; no secondary leaflets external to horseshoe; frontal sac distinct in males, rudimentary in females, practically absent in young females; wing membranes attach to ankle or tarsus; interfemoral membrane slightly angular behind; extremity of tail pierces membrane. Color.-Pelage reddish chestnut above, base of hairs for threefourths their length pale reddish white; paler below; some specimens with bases of hairs almost pure white, extremities dark reddish brown.

237

HIPPOSIDEROS I,

Measurements of HipposidfWOS bieolor (Temminek). mm.

Length, head and body Tail Head Ear length Forearm, length Third finger Fifth finger . Tibia Foot and claw

48.2 30.5 17.8 17.8 40.6 62 50.8 17.8 8

Remarks.-The type locality is, "Java, Amboina, and Timor." Peters records a specimen from Luzon. The records of Elera for Palawan and Calamianes may be disregarded. HIPPOSlDEROS WRIGHTI sp. nov.

Type.-No. 381, E". H. Taylor collection, collected July 26, 1923, in deserted mine tunnel, Baguio, Benguet (near Headwaters gold mine), by E. H. Taylor and John Suarez Wright. Alcoholic specimen with skull separate. Diagnosis; description of the type.-Adult male. Head slender, anterior part of nose leaf rather squarish, internasal part folded and inflated, bulbous posteriorly, separated from lateral wings by deep grooves; posterior leaf with four septa on anterior face making four shallow cells; a distinct small frontal pore with a transverse erect triangular leaf preceding it, situated just back of two pyramidal g'lands which are situated just back of posterior leaf; ears large, convex anteriorly, rounded broadly above to beyond greatest elevation, where a slight but distinct flattening occurs, and then rounded to point of attachment; tail with seven vertebrre, the last two free; a few scattered vibrissalike hairs on head and nose leaves; latter covered with fine short hairs; body covered with very long fur, that on belly shorter than above; a considerable growth of fur on anterior part of ear, separated on inner side by a naked space from a narrow strip of hair which goes to summit; whole surface with a few scattered hairs; outer surface of ear haired on lower half; wing membranes attach low on sides, and to lower tip of tibia; interfemoral membrane somewhat angular behind, not rounded; membranes naked except where they attach to body; tip of lower lip with two naked calloused areas scarcely separated by an obsolete groove; tail pierces interfemoral membrane.

238

PHILIPPINE LAND MAMMALS

Color.-Fur brownish black at tip, passing through gray to dirty white on basal half; ears and wing membranes dark blackish brown. Skull and teeth.-Skull normal, nasal swellings only moderately inflated; posterior part of skull rounded; sagittal crest only slightly developed, terminating anteriorly at flattened triangular facial area, which partly roofs nasal swellings; palatal bridge abcmt equal to length of molar series; canines large t narrow cingulum forming anteriorly a minute cusp, a small cusp formed at posterior point of cingulum; second upper premolar in COD tact with canines, crowding the first, which is extremely small, entirely out of tooth row; molars normal; upper incisors small, separated, summits 路pointed and curved inward and backward; not or only slightly bifid; lower canines with a distinct posterior cusp; lower incisors trifid, forming a close row, outer pair same size as inner, slightly in contact with eanines; inner pair slightly overlap outer; first lower premolar large, slightly out of alignment. . Measurement of Hipposideros wrighti sp. nov. mm.

Total length Head and body Tail Ear: Length Width Forearm First finger Second finger Third finger: Metacarpal First phalanx Second phalanx Fourth finger: Metacarpal First phalanx Second phalanx Fifth finger: Metacarpal First phalanx Second and third phalanges Nose leaf: Horizontal Posterior Tibia Foot and claws Calcar

68 41 27 17.5. 17

39 6 31.5 28.5.

16 15 28 10.5 8

27.6 12 10 4 5.3 17 7 10

239

HIPPOSIDEROS

Measurement of Hipposideros wrighti sp. nov.-Continued. mm.

Skull: Total length Condylobasal length Zygomatic width Width of nasal swellings Upper tooth series, including canine Lower tooth row, including canine Mandible

14.6 13.3 7 4 5.3

5.5 10

Remarks.-This species is obviously related to Hipposideros bicolor but differs in color, and has a short body and a shorter tail. The character of the nose leaf and the inflated internasal septum seem to distinguish the forms. The small triangular leaflet in front of the small, nearly obsolete frontal pore is likewise a distinguishing character. I doubt that these characters are present on the specimen identified as H. bicolor, reported from Luzon by Peters.* The species is named for John Suarez Wright, who helped collect the type specimen. HIPPOSIDEROS PYGMlEUS (Waterhouse)

Rhinolophus pygrnmu,s WATERHOUSE, Proc. ZooL SOC. London (1843) 67 (Philippine Islands). Phyllorhina pygmma GRAY, Proc. Zoo!. Soc. London (1866) 82; PETERS, Monatsb. Akad. Wiss. Berlin (1871) 317; DOBSON, Monog. Asiat. Chirop. (1876) 63; Cat. Chirop. Brit. Mus. (1878) 139. Hipposiderus pygmma TROUESSART, Cat. Mamm. Suppl. (1904) 71; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 16; Proc. U. S. Nat. Mus. 46 (1913) 309 (Luzon, Montalban and Pasacao, South Camarines). Rhyllorhina pygmaea ELERA, Contrib. Fauna Filipina (1915) 85 (error typ.).

Description 0/ the species.-(From No. 354, E. H. Taylor collection; collected in San Mateo Cave, Montalban, Rizal Province, Luzon, January 5, 1923 by E. H. Taylor.) Young adult female. Nose leaf relatively very large, practically covering muzzle; nasal openings not especially deeply sunken; internasal septum greatly reduced, scarcely more than indicated postel'iorly; circumnarial flaps elongate, rising above level of horseshoe, forming two small pockets anterior to nostrils; median nose leaf erect, strongly haired on upper part; posterior leaf rising well above median leaf, its upper edge semicircular, the front face divided into four cells by three distinct septa; two well-developed secondary leaves external to horseshoe; two small

* M<>natsb.

Akad. Wiss. Berlin (1861) 709.

240

PHILIPPINE LAND MAMMALS

conical glands posterior to nose leaves; no trace of frontal gland; ears broad, with short subacute tips, outer margin strongly concave in upper fourth; lower part of outer edge strongly folded in, outlining the anti tragal lobe, the summit of fold designated by a rounded calloused tip; forearm long in proportion to length of body; wing membranes attach to metatarsals; calcar elongate; tail almost twice la s long as tibia; interfemoral membranes drawn to a point behind; extremity of tail free from membrane; feet small; body covered with elongate fur above and below; fur heavy on sides of head behind eye, and posterior to nose leaves; outer third (posterior) of ears naked; wing membranes practically naked except near body; a few elongate hairs on feet; lateral leaves extend some distance beyond edge of main horseshoe. Color.-Fur blackish brown on distal part of hairs and cream white on basal portion; membranes blackish ,b rown. Skull and teeth.-N asal swellings greatly enlarged for size of skull, being nearly as large as in much larger species; sagittal crest low, terminating gently; posterior part of brain case somewhat inflated. Upper incisors at tip of premaxillaries relatively large, second cusp scarcely distinguishable; canines with a very slight cusp anteriorly and a small well-defined posterior cusp; first premolar rather large, only slightly out of alignment, separating canine from second premolar; latter with a distinct anterior cusp; lower incisors strongly trifid, about equal in size, the two inner anterior to and slightly overlapping outer; lower canine rather weak, the cingulum anteriorly and posteriorly forming a cusp. Measurements.-For measurements of the specimen see page 233, under Hipposideros obscurus. Remarks.-I obtained one specimen from San Mateo Cave, Montalban, near Manila. It was clinging to the low roof of the cavern several hundred feet from the mouth. The specimen agrees very well with the description of the type. Most of the measurements are a trifle greater. The type locality is "Philippine Islands." The type, collected by Cuming, is reported by Dobson as being in the ZoOlogical Society collection (now in the British Museum). The species has been reported from Pasacao, Camarines Sur, Luzon. Paul Bartch obtained three specimens, now in the United States National Museum, from San Mateo Cave. __ This species is the smallest of the genus known from the Philippines.

241

HIPPOSIDEROS

HIPPOSIDEROS ANTRICOLA

(Peters)

Phyllorhina antricola PETERS, Monatsb. Akad. Wiss. Berlin (1861) 709; (1871) 326 (Paracale, Luzon). Phyllorhina bicolor (part.) DOBSON, Cat. Asiat. Chirop. (1876) 70; Cat. Chirop. Brit. Mus. (1878) 148. Hipposideros antricola MATSCHIE, Sitzb. Ges. Nat. Freunde Berlin (1898) 39; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 15; Proc. U. S. Nat. Mus. 46 (1913) 309 (Pasacao, Camarines, Luzon; Cotabato, Mindanao).

Description of the species.-Ear somewhat longer than broad, tip right-angled and outer edge not c~t out; nose leaf with four cells; tail almost twice as long as tibia; wing membrane attached to tarsus. Color.-Fur brown, base white; belly fur also bicolored, but paler; wing membrane brown. Measurements of the t ype of Hipposide?'os antriMla (Pete'rs). mm.

80 50 17 30

Total length Head and body Head Tail Ear: Length Width Forearm Third finger Fifth finger Tibia Alar expanse

17 15 39 59 49 16.5 260

Remarks.-The type, in the Berlin Museum, was collected at Paracale, Luzon, by J. Jagor. HolUster reports specimens from southern Luzon and southern Mindanao. HIPPOSIDEROS DIADEMA

(Geoffroy)

Andersen * has divided this species into five subspecies, one of which occurs in the Philippines. He mentions a specimen from Zamboanga, Mindanao, collected by A. Everett, which is markedly smaller than other known forms, and expresses the opinion that it probably merits subspecific distinction. A specimen in my collection from N ovaliches, Rizal, Luzon, shows the same characteristics as the Zamboanga specimen mentioned by Andersen. I propose to differentiate this form as Hipposideros diadema anderseni subsp. nov.

* Ann. 26946:r-16

& Mag. Nat. Hist. VII 16 (1905) 497.

242

PHILIPPINE LAND MAMMALS

Key to the su bspecies of Hipposideros diadema in the Philippines. a 1. Forearm, 83 to 88 millimeters; tail, 43 to 54; body and 路 head length, 94 to 104; ear, 27 to 30 ........ H. diadem a griseus (Meyen) (p. 242). a ". Forearm, 76 to 77.5 millimeters; tail, 35 to 35.8; body and head length, 84; ea r , 24 ............................ H. diadema anderseni subsp. nov. (p. 246). HIPPOSIDEROS DIADEM A GRISEUS (Meyen)

Rhinolbphus griseus MEYEN, Nov. Act. Ac. Cres. Leop.-Car. 16 pt. 2 (1833) 608 pI. 46. Hipposiderus diadema g?'iseus ANDERSEN, Ann. & Mag. Nat. Hist. VII 16 (1905) 497; HOLLISTER, Philip. J ourn, Sci. 搂 D 7 (1912) 15; Proc. U. S. Nat, Mus. 46 (1913) 309 (Luzon, Catanduanes, Guimaras, and Mindanao).

Description of the sUbspecies.-(From No. 421, E. H. Taylor collection; collected near Zamboanga, Mindanao, April, 1923, by E. H. Taylor.) Adult female with embryO'. Large; horseshoe not covering entire muzzle; internasal septum low, circumnarial flaps reach slightly above level of horseshoe; middle leaf with a distinct vertical ridge, broadly triangular viewed in profile; two indistinct glandular prominences on either side of ridge; posterior leaf divided on anterior face into four cells by three septa, two outer septa very indistinct, with upper edge broken mesially by a slight fold or groove; two small glandular prominences behind posterior leaf; three supplementary leaflets external to horseshoe; ears large, lower anterior portion convex, then ascending to summit in a straight line; posterior edge almost a straight line, very slightly convex posterior to summit; a fold in lower part of ear designates the antitragal lobe; wing membranes attach IO'W on sides and to a point 2 millimeters from distal end of tibia; calcar large, longer than foot; tail long, tip extending some distance beyond interfemoral membrane; nose leaf covered with minute hairs, median leaf with longer hairs on summit; upper half of posterior side of ear nearly naked; body covered with long fur; wing membranes practically naked. Tips of fourth and fifth fingers are slightly bifid. Color.-Pelage brown, hairs for the middle two-thirds grayish white, basal part brownish; hair on sides Qf back above attachment of membranes and entire outer two-thirds is creamy white, forming an elongate irregular light spot; a similar white spot on shoulder; hair on sides of head grayish brown; abdomen more or less uniform drab-gray without noticeable spots except

243

HIPPOSIDEROS

on shoulder; basal part of hair darker; wing membranes dark brown. Skull and teeth.-Strong sagittal crest present, its highest elevation above narrowest interorbital constriction, then ab118

120

IZ4

122

126

Batan I.

PHILIPPINE ISUNDS

® Hipposideros antricola @ Hipposideros d. grise us

® Myotis puftopidus ® Myotis macpotapsus

@ Myotis jeannei

® Myotis patfliciae CD Myotis bflowni Myotis neflflei

.....

~ ("..

10·

:>-

'<:

<>Cagayan Sulu

6·

liB

FIG. 12.

122

124·

126

Distribution of Hipposidero8. Chi/,ophylla. and Myotis in the Philippines.

ruptly descending; upper surface of nasal swellings flattened; zygomatic arch flattened, the greatest vertical elevation of bone, 2.7 millimeters; brain case much widened above bullre, a slight transverse occipital crest; a distinct prominence marks end of low supraorbital ridges.

61 21 15

62 29 28

28 24 84 12 65

28 24 84 12 63 57 27 29

146 99 47 10 12.. 1

421, female.

423, mall'.

424, male.

H ipposideTos diadema gTiseus.

425, female.

E. H. Taylor collection No. and sex. ,

---

HippoBide rOB diadema anderBeni.

58 21 15

60 27 28

28 25 83 12 64

59 20 15

60 27 27

29 24 82 12 62

62 21 16

65 30 28

29 24 86 12 67

59 20.5 14

61 28 29

28 24 86 12 67

58 20 14

60 27 28

28 25 83 11 63

60 20 13.5

62 28.5 28

28 24.5 84 13 65

60 21 15

62 28 29

28 24 84 13 64

52.5 18.5 13

52.5 23 26

24.5 22 76

429, 431, 330, female. male. male. --- --- --- --- --- --- --143 140 149 144 135 144 119 94 I 96 99 98 100 95 96 84 44 45 44 48 40 50 35 4411 10.5 11 10.5 11 11 11 9.5 13.2 13 12.8 12.1 12 12 13 10.4

380J 1413, female. female.

152 104 48 11.5 12.2

~:::n~h;~:~:~~~~~~~~~~~~~:~~~~:~~~~~~~~~~~~~~~~~~~~~~~~~~~I ~~

Fourth finger: Metacarpal _______ _

Third finger: MetacarpaL _______ _ First phalanx ___ _ Secondphalanx _________________________________ _

Secondfinger~

Width of horseshoe ______ _ Width of posterior leal. ______ _ Ear: Height ___ : _____ _ Width_ .. _______ _ Forearm. _________ _ First finger and claw .. ___________ _ _________________________________ _

Total length_ Head and body ... __ Tail. _. _____________________ _

320, male.

[Measurements in millimeters.]

Measurements of the subspecies of Hip'poside?-os diaderna.

E= 00

~ ....

:x: ....

~ ~ ~

____ ___________ _

52 21 17 34 17 17

~andible

Lower tooth series with canine _________________________ _ .. __________________________________________ _

- - - - - - - _.

14 22

-----------------._--Width of nasal swellings. _ - -- -- -- -- - --. Width of brain case .. _____ _ -- --- -- -----. Pper tooth series with canine _________________________ _ -- -- - - ------.

Calcar __ Foot. Skull: Total length __________ _ - -Condylobasallength ____ . ______________ - -- .----- ------ - ---Zygomatic width ______ _ - - - --

First phalanx- __ _ Second phalanx ___ _ Tibia _____ ___ _________________ . _________ - --- __ -- --- - - -- --

~etacarpal

Fifth finger:

31 27 18 9.2 15 12 .8. 13 . 4 22

55 22 17 35 17 17 30 . 2 27 18 9.2 15 13 13 . 9 23

54 20 17 35 19 17 32 27 18 9 15 13 13 .8 22

51 20 17 38 21 17 33 27 18.2 10 15 13.5 14 23

56 21 17 35 20 17 81 27 17.6 9 16 . 5 18 14 22

58 20 17 36 16 16.6 31 27 17.6 10 15 13 14 23

56 20 17 36 20 17

----

31. 5 28 17.6 9 14 13 14 22

62 20 17 85 20 16

14 22.6

32 27 18.2 9.6 15

62 21 17 36 19 17

29 25 16.2 8.6 14 12 13 20

48 19 16 31 18 17

t\:)

"r:n0

l':l

.... t::l

"'d

::x: .... "'d

246

PHILIPPINE LAND MAMMALS

Upper incisors well develoPyd, unequally bifid, tips close together; canines large with a slight shelf formed by cingulum on lower anterior face; first premolar small, slightly out of alignment, not reaching cingulum of second premolar, which is widely separated from canine; anterior notch of palatal .bridge posterior to second premolar; lower incisors greatly thickened, as broad as wide, the outer indistinctly trifid, the interior bifid. VariatiOn.-Apparently little variation occurs other than is recorded on the table of measurements. Two young specimens, Nos. 418 and 428, have the body a uniform dark brown above (no white on underfur), but with two pure white spots on back just over armpits and a white stripe along each side where wings attach; belly and throat gray, tips of fur white, and a large white area on shoulder; the bulk of the specimens follow the described markings and color except that in some the light spotS' are almost obsolete. In No. 423 (adult) the fur above is almost without any white on the hairs, and the median portion is a grayish brown. The tips of the hair vary slightly in shade. A single red phase was found at Zamboanga in a colony of normally colored bats. The tips and base of hairs are russet, the median part of hair reddish buff, lighter on belly. Remarks.-There are twenty specimens * in my collection from Zamboanga, Mindanao; one from Baguio, Benguet, Luzon; one from Los Banos, Luzon; and four from localities near Manila. The Zamboanga specimens were shot in a cavity formed by a large slab of rock leaning against the bank of a river; this entire colony probably contained less than fifty individuals. Several of the specimens carry large embryos; two of the specimens are very young. Meyen's type is from San Mateo Cave, Montalban, near Manila. Specimens have also been reported from Guimaras and Catanduanes Islands. HIPPOSIDEROS DIADEMA ANDERSENI Bubsp. nov.

Type.-No. 330, E. H. Taylor collection; collected near Novaliches, Rizal Province, Luzon, March, 1922, by E. H. Taylor. Alcoholic specimen, with skull separate. Diagnosis; description of the type.-Adult male. Differs from HiJpposideros diadema griseus chieft.y in size; nose leaves narrower; forearm considerably shorter; head and body measurement considerably less; tail shorter; fingers practically the same length as in griseus.

* In the table No. 320 is from Baguio, Benguet, Luzon; No. 380 is from Los Banos, Luzon; Nos. 413 to 431 are from Zamboanga,. Mindanao.

CHILOPHYLLA

247

Color.-Same as in the preceding subspecies. Skull and teeth.-Skull and molar teeth smaller than in gri.seus; lower inciso~s distinctly smaller, upper incisors somewhat larger than corresponding teeth in griseus. Color practically the same. Measurements are given in the table on page 244. The subspecies is named for Knud Andersen, author of Chi, roptera in the British Museum, second edition. Genus CHILOPHYLLA Miller Chilophylla, MILLER, Proc. U. S. Nat. Mus. 38 (1911) 395; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 16; MILLER, Proc. BioI. Soc. Washington 2路5 (1912) 117.

A small bat having the nose leaf consisting of an ill-differentiated horseshoe and lancet, the former produced anteriorly as a pair of conspicuous lappets separated by a deep median sinus and projecting noticeably beyond, each lobe formed largely by an elongated supplemental leaflet which continues forward the outline of the obliquely truncated horseshoe proper; upper lip very low, much less noticeable than lappets, with small upright median process and two ill-defined cross flutings; nostril at inner border. of pit, the raised edges of which form a somewhat diamond-shaped figure; a freely projecting process at outer corner of diamond; a projecting wart within pit directly over opening of nostril; space between nostrils occupied by a freely projecting, somewhat bilobate, process, rather broader than high; below this process another process somewhat smaller and deeply bilobate, each lobe connected by a ridge with inner edge of corresponding lappet of horseshoe; a low slightly curved ridge above nostril at a distance about equal to vertical diameter of diamond; outer extremity of this ridge approaching, but not reaching, a wartlike excrescence near margin of horseshoe; ear very large, inferior lobe greatly developed and producing a funnel-like general form, the substance of ear everywhere thin and pellucid with no traces of ridges in region of meatus; thumb with metacarpal about one-fourth as long as that of third finger, phalanges and claw short, the combined length only about onethird that of metacarpal; leg and foot slender; calcar reduced to a mere tubercle, wing membrane inserted at ankle; antebrachial membrane extending outward as a broad fold to include entire metacarpal of thumb, its width so great as to form a conspicuous pouch at bases of metacarpals; interfemoral membranes narrow, barely extending to rudimentary calcar; tail very short, . its length about half that of femur.

248

PHILIPPINE LAND MAMMALS CHILOPHYLLA HIRSUTA MUler

Chilophylla hirsuta MILLER, Proc. U. S. Nat. Mus. 38 (1911) 395, pI. 18; Proc. BioI. Soc. Wash. 25 (1912) 117; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 16; Proc. U. S. Nat. Mus. 46 (1913) 309.

Description of the species.-A small delicately formed bat with large- funnel-shaped ears and unusually long soft fur; nose leaf inconspicuous except for the two lappets projecting on upper lip; its median portion essentially bare, its margins densely haired, especially at sides; general color light brown; fur long and glossy, hairs at middle of back about 11 millimeters long, those on belly about 7 millimeters; fur on wing membranes extends to a line joining middle of humerus with basal third of femur; ear thickly sprinkled on both surfaces with loosely spreading hairs, those along inner border about 4 millimeters long; hairs on edge of nose leaf and on neighboring portion of muzzle radiating stiffly outward, the longest near middle of nose leaf, about 5 millimeters long. Color.-Entire animal uniform light brown, between woodbrown and fawn color (Ridgway), the middle of belly faintly tinged with light buff at tips of hairs; undercolor very dark brown with a slaty tinge; ears translucent light brown; membranes an indefinite dark brown. Skull and teeth.-(The skull of the type was lost. Miller * obtained a specimen of what appears to be this species from Port Swettenham, Federated Malay States. The following diagnosis is from that specimen.) Skull like that of a small Hipposideros in geheral features but nasal swellings very small; maxillaries and premaxillaries conspicuously produced anteriorly, length of premaxillary fully twice median length of palate; distance from level of front of canines to upper margin of nares about one and one-half times that from upper margin of nares to narrowest portion of interorbital constriction; upper canine strongly projecting forward, its shaft with large anterior and posterior secondary cusps, the points of which lie in same horizontal plane as points of paracones of molars; large premolar more reduced thah in related genera; posterior lower premolar with shaft compressed laterally, the form of the cusp without evident resemblance to that of prot~conid of first molar. (After Miller.) . 1-1 1-1 2-2 3-3 _ Dental formula: 1. 2-2' c. 1-1 pm. 2-2' m. 3-3 - 30.

* Proc.

BioI. Soc. Washington J6 (1912) 117.

249

VESPERTILIONIDJE

Measurements of the type of ChilophyUQ, hirsuta Miller. mm.

Head and body Tail Femur Tibia Foot Forearm First finger Metacarpal Second finger Third finger: . Metacarpal First phalanx Second phalanx Fourth finger, metacarpal Fifth finger, metacarpal Ear: From meatus From crown Greatest width Nose leaf, greatest length Horseshoe, greatest width Height of lancet above muzzle Skull (from Federated Malay States specimen) : Condylobasal length Zygomatic breadth Breadth across nasal swellings Interorbital constriction Breadth of brain case Mandible Maxillary tooth row Mandibular tooth row

33 7 14.2 13.6 6.4 33.8 8.2 6.2 32 24.2 6.2 22 25.4 27.4 12 10 11.6 8.2 52 2.2 13 6.6 3.6 2 6.4 8.2 4.8 5

Remarks.-The type was collected on Alag River, Mindoro, December 2, 1906, by Edgar A. Mearns. It is No. 14482, United States National Museum, and consists of a skin. The skull and body were preserved in alcohol but were lost. Family VESPERTILIONIDlE Vespertilionida3 GRAY, London Med. Repository 15 (1821) 299. (Most other authors have used this name or a variation of it. Dobson excluded Natalus and Thyroptera.)

Humerus with trochiter very noticeably larger than trochin and projecting distinctly beyond head, its surface of articulation on scapula decidedly more than half as large as glenoid

250

PHILIPPINE

LAND~

MAMMALS

fossa, distinctly concave and sharply outlined; epitrochlea obsolete, capitellum scarcely out of line with shaft; ulna usually fused with radius at its head, shaft reduced to a scarcely ossified fibrous strand; second finger with fully developed metacarpal and one small bony phalanx; third finger with 'three phalanges, distal one cartilaginous throughout except at extreme base, where a distinct joint is formed with middle phalanx; shoulder girdle strictly normal in general structure, seventh cervical vertebra free except in Tomopeatinre; presternum small, with forward-projecting variously developed median lobe; mesosternum flat and scarcely keeled, usually slender; foot normal; fibula threadlike, complete, or with upper extremity cartilaginous, extending to head of tibia; pelvis normal, boundaries of sacral' vertebrre distinct, ischia widely separated posteriorly, a symphysis pubis in males; lumbar vertebrre free; skull without postorbital processes; premaxillaries without palatal branches, palate widely emarginate anteriorly, abruptly narrowed behind tooth rows, the sides of its posterior extension parallel or nearly so; teeth usually normal, though in certain genera show reduction of cusps; ears usually but not invariably separate, anterior border with distinct basal lobe (except in Tomopeatinre) ; tragus usually well developed; simple muzzle without leaflike outgrowths; tail well developed, extending to edge of wide interfemoral membrane. Members of the family may be reco路g nized externally by their simple muzzle and lips; usually separate ears with well-developed, straight, or slightly curved tragi; long tails extending to edge of wide interfemoral membrane, but never much beyond; by the presence of only two bony phalanges in third finger, and complete absence of true sucking disks on sole of foot and on thumb. (After Miller.) Miller recognizes six subfamilies, four of wlhich are represented in the Philippines. Key to the PhiliPPine subfamilies of Vespertilionidre. a '. Anterior upper premolar large, not differing conspicuously from posterior premolar in size or form; nostrils tubularly elongate. MURININJE (p. 251). a '. Anterior upper premolars greatly reduced in size or absent; when present conspicuously different from posterior in form and size; nostrils not tubularly elongated. b '. Sternum short and broad, its length in middle line not twice as great as breadth of presternum; four or five ribs connected with sternum; ear slightly but evidentI'Y funnel-shaped. KERIVOULINJE (p. 253).

MURINA

251

b 2. Sternum slender, its length in middle line more than twice as great as breadth of presternum; six or seven ribs connected with sternum; ear not funnel-shaped. c 1. Presternum with median lobe larger than body of bone; scapula with coracoid straight, directed inward; second phalanx of third finger nearly three times as long as first. MINIOPTERINlE (p. 264). c'. Presternum with median lobe much smaller than body of bone; scapula with coracoid curved outward; second phalanx of third finger not especially elongated .............. VESPERTILIONINlE (p. 277). Subfamily MURININ.JE Murininre MILLER, Bull. U. S. Nat. Mus. 57 (1907) 229.

Anterior upper premolar very slightly reduced, scarcely smaller than posterior premolar and essentially like it in form; molars essentially normal or considerably modified, the metaconid noticeably the largest cusp in the first and second upper molars; nostrils conspicuously tubular, projecting. The two genera, Murina and Harpiocephalus, differ greatly in the character of the molars. Only Murina is represented in the Philippines. Genus MURIN.oj\. Gray Murina GRAY, Ann. & Mag. Nat. Rist. 10 (1842) 258; DOBSON, Cat.

Chirop. Brit. Mus. (1878) 277 (part.); MILLER, Bull. U. S. Nat. Mus. 57 (1907) 229. Ocypetes LESSON, Nouv. Tab!. Regne Anim. (1842) 20 (non Wagler). Harpiocephalu8 DOBSON, Cat. Chirop. Brit. Mus. (1878) 276 (part.).

Small bats; ears separate, with well-developed tragus; tail long, involved in the long interfemoral membrane for its entire length; muzzle simple, nostrils very elongate, tubular; upper incisors well developed, without distinct secondary cusps, the outer larger than inner and closely crowded against canine, the four teeth almost exactly in line with each other; lower incisors with crowns about equal in length, forming a continuous strongly convex row between canines; canines well developed, not peculiar in form; second upper premolar unusually large but not peculiar in form, the cingulum resembling a cusp; anterior premolar not as large but of essentially the same structure; first and second upper molars with length along outer edge equal to transverse diameter, the cusps normal in position, the W-shaped pattern present but somewhat distorted by the reduced condition of the parastyle; hypocone absent; third upper molar not unusually reduced, cusps of normal size; mandibular teeth normal for the family; skull essentially as in medium-sized species of Myotis. (After Miller.)

252

PHILIPPINE LAND MAMMALS

. 2-2 1-1 2-2 3-3 3-3' c. 1-1' pm. 2-2' m. 3-3 =34. The genus, of some eight species, ranges from central and eastern Asia to Japan and the Malay region. Murina cycwtis Dobson has been reported from the Philippines. Dental formula:

MURINA CYCLOTIS Dobson

Murina cyclotis DOBSON, Proc. Asiat. Soc. Bengal (1872) 210; Jour. Asiat. Soc. Bengal 2 (1873) 206 pI. 14; HOFFMAN, Abhand. und Berichte des Konig!. Zoolog. Mus. Dresden (1886-87) (1887) 1-29 (Mindanao); HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 19; Proc. U. S. Nat. Mus. 46 (1913) 336. Harpiocephalus cyclotis DOBSON, Monog. ksiat. Chirop. (1876) 158, figs. a, b, c; Cat. Chirop. Brit. Mus. (1876) 282.

Description of the specie-s.-Muzzle somewhat shortened, obtusely conical, the end projecting beyond the lip and consisting of diverging tubular nostrils opening laterally with a slight depression between; ears nearly as wide as long and almost circular; the only interruption to regular convexity of external margin being a faint flattening of upper and outer portion, and a slight convexity opposite base of tragus; tragus long and narrow, somewhat angularly rounded on anterior edge, posterior edge nearly straight; tip pointed; two small indentations on posterior edge near base; a few scattered hairs on inner surface of ear conch; wing membranes attach on outer toe to base of claw; feet small, toes of nearly equal length; extreme tip of tail not involved in interfemoral membrane; upper surface of interfemoral membrane covered with hair w;hich is densest at root of tail, along tibia, and along calcar; hair also extends across tibia on wing membrane; back of feet thickly covered with hair which is longer than toes. . Color.-Fur bicolored above, basal portion dark brown, ti~s ferruginous; below paler brown throughout. Skull and teeth.-Skull similar to that of Myotis; upper incisors long and slender; outer incisor shorter than inner and almost touching canine at base but widely separated from it at extremity; at outer side of its base a small cusp projects outward; first premolar scarcely equal to half the vertical extent of canine, but about as high as second molar; first and second molars nearly equal but third narrow, forming a simple bony lamina as wide as preceding molar, but less than half its anteroposterior diameter; in lower jaw the last molar is smaller than the preceding one but has the same number of cusps. (After Dobson.*)

* Cat.

Chirop. Brit. Mus.

253

KERIVOULA

Measurements of Murina cyclotis Dobson. mm.

Head and body Tail Head Ear Tragus Forearm Thumb: Metacarpal Phalanx Third finger: Metacarpals First phalanx Second phalanx Fifth finger: Metacarpals First phalanx Second phalanx Tibia Calcaneum Foot

43.1 38.1 17.7 15.27 8.9 33.02 2.5 5 30.4 13.9 17.7 31.7 10.2 8.8 15.2 12.7 7.6

Remarks.-The type specimen, from Darjeeling, northeastern Bengal, is reported by Dobson as being in the Indian Museum, Calcutta. The species is included on Hoffman's record of a specimen from Mindanao, which appears to be the only report for the Philippines. Subfamily KERIVQULIN.A3:

Kerivoulina3 MILLER, Bull. U. S. Nat. Mus. 57 (1907) 232.

Externally as in the Vespertilioninre. Internally, the sternum very short, its length in a median line not twice as great as breadth of presternum; median lobe of presternum small but distinct, upright; keel of mesosternum low; only four or five ribs articulate with sternum; coracoid curved outward as in the Vespertilioninre. Kerivoula is the only Philippine genus of this subfamily. Genus KERIVOULA Gray KerivCYUla GRAY, Ann. & Mag. Nat. Hist. 10 (1842) 258; DOBSON, Cat. Chirop. Brilt. Mus. (1878) 330; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 232. KerivCYUla GERVAIS, Dict. Univ. d'Hist. Nat. 13 (1849) 213. Nyctophilax FITZINGER, Sitzb. Math. Nat. Cl. Akad. Wiss. Wien 42 (1860) 390. Cerivoula BLANFORD, Fauna Brit. Ind. (1891) 338.

254

PHILIPPINE LAND MAMMALS

Small bats with strongly concave skull; ears laid forward extending slightly beyond nostril, separate, slightly funnel-shaped, outer border arises slightly in front of inner and its lower portion is unusually widened; tragus long, very slender; muzzle 118

120

PHILIPPINE ISLANDS

124

122 Batan I.

126

® Murina cyclotis @ Kerivoula whiteheadi

® Kel"ivoula peliucida ® Minioptel"us pusillus

@ Minioptel"us australis

® Minioptel"us paululus CD Minioptel"us tibialis . Q) Miniopterus eschscholtzii ® Miniopterus tristis

LUZON

~ C":l

" ~ 10

r."

:s:.

<: s' <>Cagayan Sulu

120

FIG. 13.

122

124'

126

Distribution of Mwrina. Kerivoula. and Miniopterus in the Philippines.

simple; third and fourth metacarpals about equal, the fourth possibly slightly shorter. Upper incisors well developed, the outer about half the height of inner, but of nearly the same diameter as inner and crowded between canine and inner incisor; latter may develop a sec-

KERIVOULA

255

ondary cusp; main axis of each tooth nearly perpendicular; lower incisors with three or four cusps slightly or not imbricated; canines normal in form, rather small, the point of upper tooth extending slightly beyond cingUlum of lower when jaws are closed; lower canine with small anterointernal cingulum cusp; premolars well developed, but not peculiar in form, those of mandible subequal, their crowns subquadrate in cross section and without distinct cutting edges; molars normal, first and second upper without hypocone, third with metacone and three commissures well developed, area ~f crown about half that of first or second; lower molar with all cusps present. Skull slender, lightly built, brain case large, smoothly rounded and abruptly elevated anteriorly; rostrum slender and weak; there is no indication of sagittal crest or of supraorbital or lachrymal ridges; nares very narrow, width of aperture as viewed from above scarcely or not half its length; anterior pa~ latal emargination about as wide as deep; posterior extension of palate narrowed gradually backward; hamuli very short ; audital bullre very small, covering much less than half the surface of large cochl~ . . 2- 2 1- 1 3-3 3- 3 Dental formula: 1. 3- 3' c. I - I' pm. 3- 3' m. 3- 3 =38. This genus and Myotis of the Vespertilioninre are the only Philippine genera having the maximum number of teeth of the order Chiroptera; namely, 38. More than eighteen species of the genus are recognized. These are distributed from South Africa through India and the Malayan region to New Guinea. Key to the Philippine species of K eri'Vou la Gr ay. a 1. Inner upper incisor with a secondary cusp; interfemoral membrane rather hairy above ____ ___ _________ K eri'Voula whiteheadi Thomas (p. 256). It'. No secondary cusp on upper incisor; sometimes an enlargement on cingulum_ . b 1. Two notches or concavities on outer edge of ear; wing membranes attach at base of toes; forearm, 39 millimeters. K eri'Voula jagorii (Peters) (p. 257). b 2. A single notch on outer edge of ear. c'. Very few scattered hairs on interfemoral membrane; a small toothlike projection at widest part of tragus; forearm, 32 to 33 millimeters ______ ________ ____ __ K eri'Voula hardwickii (Horsfield) (p. 258). c'. Hairs numerous on tail, hind limbs and above interfemoral membrane; two or three small toothlike prominences on tragus at widest part near base; forearm, 29 to 33 miUimeters. Keri'Voula pellucida (Waterhouse) (p. 261).

256

PHILIPPINE LAND MAMMALS KERIVOULA WHITEHEADI Thomas

Kerivoula whiteheadi THOMAS, Ann. & Mag. Nat. Hist. VI 14 (1894) 460; Trans. Zool. Soc. London 14 (1898) 386; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 20; Proc. U. S. Nat. Mus. 46 (1913) 336.

Description of the species.-Size and proportions about as in Kerivo~~la hardwickii, but ears slightly longer; lower legs shorter; upper surface of wing membranes to a line drawn from elbow to foot, whole of interfemoral membrane except terminal half-inch, and surface of legs to feet thinly but distinctly clothed with long orange-colored hairs, these parts in K. hardwickii practically naked; hinder edge of interfemoral membrane with a few short hairs along it, scarcely forming a fringe. Color.-Rufous-orange above, slaty bases of hairs showing through; dark slaty below, the lighter tips scarcely affecting the general dark tone. Skull and teeth.-Upper incisors slender with a distinct posterior secondary cusp to the tip of which the unicuspid outer incisor just reaches; other teeth as in K. hardwickii. Measurements of the type of Kerivoula whiteheadi Thomas. mm.

Forearm Head and body Tail Head Ear from notch Tip to tip of ears aCross head Length of index finger Third finger, exclusive of cartilaginous tip Fifth finger Lower leg Hind foot without claws

32 39 39 16 13 [j 28.5 31.5 61

47 16.2 8

Remarks.-The type, in the British Museum, was collected by J. Whitehead at Molino, Isabela, Luzon, May, 1894. The species is .closely related to K. hardwickii but may be distinguished by the hairy interfemoral and the character of the incisors. In my collection is a specimen which appears to belong to this species. It was collected in Panay by R. C. McGregor. The specimen, preserved originally in alcohol about 1916, is in poor condition and most of the fur is wanting. The identification has been made on body and limb measurements, and the character of the teeth. The brain case is distinctly very declivous anteriorly and where it meets the rostrum a very distinct depression occurs, which in profile appears curved. The narial

257

KERIVOULA

notch is narrowed behind but anteriorly is wider with sides nearly parallel; anterior palatal notch rounded; posterior portion of palate narrowed gradually backward, and then slightly widened; median terminal point a small rounded knob; bullre covering less than one-third of cochlere which shows no snaillike coils; inner incisor with a distinct accessory cone at a point on a level with tip of outer incisor; outer incisor more than half as long as inner; other teeth without distinctive characi~r颅 istics. Lower incisors increasing in size from inner to Guter; inner with four cusps, outer with three. This skull is from a young animal and, as a whole, appears much smaller than skulls of other species. The interfemoral membrane attaches at the extreme base of toe. Measurements of an immatu?'e specimen of Ke?'ivoula whiteheadi Thomas. mm.

Skull: Length, incisors to occiput Anterior zygomatic width Width of brain case Upper tooth row Palate Mandibular tooth row Forearm Head and body Tail Ear from notch Tip to tip of ears Second digit Third digit, exclusive of cartilaginous tip Fifth finger Hind leg Hind foot without claws

13.6 6.3 7.0 6.0 6.0 6.2 3.2 38.0 :{S.O 13.7 80.0 :U.O 61.0 46.5 15.5 7.6

KERIVOULA JAGORII (Peters)

Vespertilio (Kerivoula) jagol-ii PETERS, Monatsb. Konig路I. Preuss. Akad. Wiss. Berlin (1866) 399. Vespe?路tilio jagorii DOBSON, Monogr. Asiat. Chirop. (1876) 185. Kerivoula jagorii DOBSON, Cat. Chirop. Brit. Mus. (1878) 338; HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 19; Proc. U. S. Nat. Mus. 46 (1913) 336; ELElRA, Contrib. Fauna Filipina (1915) 108.

Description of the species.-E"ars as in K. harchuickii but with a second slight concavity about the middle of outer margin; inner margin very convex, projecting beyond eye; tragus rather broad below, tapering to an acute point and curved outward; immediately above base, a deep but narrow notch opposite base of inner margin; nostrils close together, extremity of muzzle 259453-17

258

PHILIPPINE LAND MAMMALS

projecting; tip of tail projecting slightly; margin of interfemoral membrane not fringed with hairs; wing membranes attach at base of toes. Color.-The same as that of Kerivoula hardwickii. Skull and teeth.-Skull similar to that of other species; upper inner incisor long and unicuspidate; outer incisor very short, not half the length of inner incisor; upper premolars nearly equal in vertical extent. Measurements of Kerivoula jag(Yl'ii (Peters). mm.

Length of head and body Tail Head Ear Tragus Forearm Thumb Second finger Fourth finger Tibia Foot and claws

49.6 48.3 19 2'1.2 10.2 39.4' 9.4 81.3 57.2 16.5 9

Remarks.-The type, in the Berlin Museum, was collected by J agor in Samar. The skull measurements are unknown. This appears to be the largest species of the genus known from the Philippines. KERIVOULA HARDWICKII

(Horsfield)

Vespertilio ha?'dwickii HORSFIELD, Zoo!. Res. in Java (1825) 28; Cat. Mamm. E. Ind. Co. Mus. (1851) 39; WAGNER, Suppl. Sehreb. Saugeth. 5 (1855) 736; TEMMINCK, Monog. Mamm. 2 (1841) 222; BLASIUS, Wiegmann's Archiv. 6 (1840) 2. Kerivoula hardwickii GRAY, Ann. & Mag. Nat. Rist. I 10 (1842) 208; Cat. Mamm. Brit. Mus. (1843) 27; GRAY, Zoo!. Voyage H. M. S. Samarang (1850) 7; DOBSON, Cat. Asiat. Chirop. (1876) 148 figs. a, b, c; Cat. Chirop. Brit. Mus. (1878) 335; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 19; Proc. U. S. Nat. Mus. 46 (1913) 336. Vespertilio (Kerivoula) hardwickii PETERS, Monatsb. Akad. Wiss. Berlin (1861) 711 (Nipa-Nipa, Samar). Ke?'ivoula fusca DOBSON, Proc. Asiat. Soc. Bengal (1871) 215.

Description of the species.-(From No. 355, E. H. Taylor collection; collected near Saub, Cotabato Province, Mindanao, April 15, 1923, by F. G. Engle.) Adult male. Muzzle relatively short and broad, anterior portion with scattering hairs of unequal length; "nose" rather prominent, divided medially by a

KERIVOULA

259

slight depression, nostrils rounded, directed outward and downward; inner margins separated by a distance of 2 millimeters; only lower portion naked; lower side of mouth naked; "lip" wider anteriorly; ears large, extending a very short distance beyond tip of muzzle; inner margin forms an almost regular arc of a circle from base to tip which is somewhat pointed; a deep concavity a short distance below tip, occupying upper fourth of outer margin; below this the outer side of conch becomes expanded and projects forward; outer margin curving inwards and backwards towards base of tragus; tragus long, extending to edge of emargination below tip on outer side of ear; inner margin forming a very slight continuously curved line; outer margin with a single distinct small tooth projecting; opposite base of inner margin at a point where tragus is widest; whole tragus slightly sickle-shaped, curving backwards, and becoming attenuated at tip; inner conch of ear covered with glandular papillre with small hairs arising from them; thumb large with strong claw; wing membranes attach to base of toes; posterior margin of interfemoral membrane with numerous small conical projections j body moderately furred; wing membranes totally naked above, except for a few scattering hairs near end of forearm; and naked below generally except on portion near body; interfemoral membrane above with only a few scattered hairs; a few hairs along tail and on posterior margin of membrane; a few short scattered hairs below; tail completely involved in interfemoral membrane except for a meager portion of tip; calcar prominent; membranes transparent as in Kerivoula pellucida; second finger shorter than metacarpal of third, which is longer than fourth; the latter longer than fifth; penis pendent about 3 millimeters long; toes distinctly longer than metacarpals; pelage dense above; hair on shoulders 9 millimeters long; below less dense, hairs on breast about 7.5 millimeters. Color.-Color above between cinnamon-brown and Prout's brown. Individual hairs tricolored, outer brown fading rapidly to creamy buff, which near base changes abruptly to dark Vandyke brown in strong contrast with the two other shades; tips of hairs somewhat lighter below than above, buff area shorter and basal parts of hairs a trifle darker; membranes similar to body fur but somewhat darker; ears nearly buff color. Skull and teeth.-Skull heavier and broader than in other Philippine species of the genus; rostrum short, saddlelike, with-

260

PHILIPPINE LAND MAMMALS

out a definite depression; narial notch wider than in other forms, widest point just beyond base of incisor roots; profile more curving than in other forms, especially over rostrum; anterior palatine notch nearly circular; palate far less convex than in K. whiteheadi and posterior lateral projections of palate divergent, and median point lacks the knob of K. whiteheadi; cochlere large. Inner incisors moderate, lacking a noticeable secondary cusp; outer incisor greatly reduced, not reaching as high as cingulum of canine; lower incisors somewhat imbricate, inner teeth with four cones, other two with three cones; middle incisor largest; first upper premolar equal to third in height but much smaller in cross section; middle premolar about one-third smaller, close between first and third, leaving no interval; lower premolars equal in size. Measurements of Kerivoula hanvwickii (Horsfield). mm.

Head and body Head Tail Ear Tragus Forearm First finger Second finger Third finger Fourth finger Fifth finger Tibia Foot and claws Wing expanse Skull: Incisors to occiput Zygomatic width Width of brain case Height of skull Length of palate Length of upper tooth row Length of lower tooth row Mandible

40 16.5 45 13.5 8.4

32 8

32.5 69 4'8

49 16.2

7.3 250 14.4 9.5 7.5 7 5.3 6 6.6 9.5

Remarks.-I am of the opinion that the eastern form of this species, particularly Philippine specimens, is separable from more-western specimens on the basis of the smaller size. The measurements given by Dobson show hardwickii to be an animal

261

KERIVOULA

somewhat larger, its forearm 5 to 6 millimeters longer, the wing spread 3 centimeters greater. However, I lack extra-Philippine material and cannot judge whether the transition in size is abrupt or gradual toward the east. The tragus appears to be of a somewhat different shape. My specimen agrees in general characters with that of Peters from Nipa Nipa, Samar. Thomas * mentions a Mindanao specimen. Java is the type locality. MeasUTements of K e?'iv oula hardwickii (Horsfield). [Meas u rem en ts in millimeters.] --

- - -,- - - - - ,

Sa ub, Cotabat o, Minda- Nipa Nipa, nao. No. Samar. 355, E. H. P eters. Taylor collection.

Dobson.

1------ -1---

Length, head and body, in alcohol. . __ ~ __ .. _

40 16. 5 45

40 44

59 19 43 15

12 10

Forearm .. - - - - - - - - - - - - - - - - - - - - + - - 1- -- -------. -------. Third digit __ _____ ____ _. ______ . ____ _________________ ___ _ Fifth digit. ________ ______ _____ __ . _____ . __ ___ ___________ _ Tibia _ ___ ____ .. . ____ ______ _______ ____ ____ ____________ __

I ~i::aen~:~::s~~~~~~~ ~~~~~ ~ ~~~ ~ ~ ~~ ~~~~ ~ ~ ~~~~:~ ~~~~~ ~~ ~~~

32 69

33 69 . 8

16.2 49

16.5 17 1 48H2 7 _________ _

39 70 57

7.3 260

1__________ 1

280

KERIVOULA PELLUClDA (Waterhouse)

VespeTtilio pellucidus WATERHOUSE, Proc. Zool. SOC. London (1845) 3; WAGNER, Suppl. Schreb. Saugeth. 5 (1855) 742. Kerivoula pellucida DOOSON, Monog. Asiat. Chirop. (1876) 149; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 19; Proc. D. S. Nat. Mus. 46 (1913) 336; ELERA, Contrib. Fauna FiIipina (1915) 107 (Samar) .

Description of the species.-(From No. 376 E. H. Taylor collection; collected at Bud Daho, Jolo, November, 1917, by E. H. Taylor and Francis Link.) Adult female. Muzzle and sides of head in front of ear sparsely covered with hair of unequal lengths; "nose" prominent, projecting slightly, nostrils rather widely separated, with a slight median depression; ears large,

* Trans.

Zool. Soc. 14 (1898) 386 (in Kerivoula whiteheadi).

262

PHILIPPINE LAND MAMMALS

transparent, inner edge rounding to tip which is rather more !rounded than pointed; outer edge of ear with a small notch some distance below tip; below notch a very slight convexity; \tragus long, attenuated, failing to reach edge of notch on ear, about 1.5 millimeters wide near base and 8 millimeters long; near tip less than 0.5 millimeter wide; at a point on outer edge at widest part of tragus two or three small toothlike projections; -inner side of tragus straight or nearly so, outer with a distinct elongate concavity; wing membranes ample, attaching to middle of outer toe; toes longer than metatarsals, four inner toes of nearly equal length, outer much shorter; tail, except extreme tip, involved in interfemoral membrane; calcar strongly developed; a fleshy enlargement about fourth vertebra of tail, probably glandular; fur above moderately dense, longest hairs about 8 millimeters on shoulders; on breast the longest hairs about 6 millimeters; a few scattered hairs on forearm; wing membranes generally naked above, except near body and a few scattered hairs on posterior edge of membrane between last finger and attachment to foot; hind limb to tip of toes with rather prominent growth of hair; hairs on tail short but relatively dense above gland; scattered elongate hairs on interfemoral membrane; a distinct growth of hair. on underside of membrane near posterior border; that under interfemoral membrane usually short. Color.-Fur above light Sudan brown (Ridgway), fading to a much lighter color near base, similar to color below; hairs about neck above slightly more orange; wing membranes transparent, light brown. Skull and teeth.-Skull rather slender, declivous anteriorly; rostrum with a small depression near brain case; sides of narial notch of rostrum nearly parallel; zygoma moderately expanded; anterior palatal notch not rounded behind, posterior extension with sid~s nearly parallel, median portion ending in a slight prominence. Inner incisors moderately large, caniniform, without secondary cusp (except on cingulum), outer incisor short, not reaching higher than cingulum of canine; third premolar much larger than other two but no higher than first; second of equal diameter but somewhat shorter than first; lower incisors somewhat imbricate, the inner smallest, second and third of nearly equal size, the latter somewhat thicker; inner lower incisor with four cusps, outer two with three cusps; lower canines with a rather prominent inner cingulum cusp.

263

KERIVOULA

Measurements of Kerivoula pellucioo (Waterhouse). [Measurements in millimeters.]

E. H. Toy'" W~'''ti,. NO;==:! 376, Jolo. Total length .. _ _____ ______ ____ ______ __ __ ______ Head and body_________________________ ______ TaiL_ _____ ______ __ __ _____ __ __ ______ __ _____ __ Ear from base_ _ ____ __ ____ __ __ ____ _____ _____ __ Tragus ... _______________________________ .____ Wing spread .. ________________________________

90 40 50 16 8.5 254

Forearm_____________________________________ First digit .. __ ____ __ ________ ____________ _____ _ Second metacarpaL_ ______ ________ __ __ ____ ____ Third digit. _ _____ ___ __ ___ _____ _______ ___ _____ Third metacarpaL __ __ ___ ________ _________ ____ Fourth digit. _____ ________ ___ _____ __ ___ _ ____ Fourth metacarpaL__ ___ ___ ____ ___ ____ __ ____ __

33 7 35 68.8 33.8 51 33

Fifth di~t.---------------------------------Fifth metacarpal. _______ ________ ______________ Tibia. _____________________ â&#x20AC;˘ _____ __ _____ ___ __

52 32 18

Skull: Incisors to occiput________________________ Zygomatic width .. ________________________ Width of braincase. _ __ ______ ______ __ ____ __ Palate from notch .. _______________________

15.9 7.7 7 7

_~pper tooth row. - -- - - - - - - - ---- --- -- -- -- - Lower tooth row. __________ "___ ______ _____ Mandible_ _______ __ __ ___ ______ ___ ______ __

6.5 6.8 9.2

66, Palawan.

67, Palawan.

85 36 49 16 8.6

84 36 48 15 8.5

Locality unknown.

31 7 34 68 33 50 32 60 31 17.2

30 6.6 33 64 38 50 31 46 30 16.8

83 38 45 16 9 230 29.6 6 33 62 33 49 30.6 46 29 17

16 8 7.7 6.8 6.7 6.6 10

14.6 8.2 7.6 7

14.5 8.4 7.9 6.8

6.6 10.6

6.7 9.8

---------- ---- ------

Remarks.- The unnumbered specimen is from either Baguio or Cotabato, probably the latter (the tag was lost in shipment). The J 010 form here described has a longer, slenderer skull than the other forms, the second incisors are much smaller and lower, and the groove on outer side of upper canine is more noticeable than in the other specimens. The unnumbered specimen has a short broad skull, but the outer incisor is more than half as high as inner; the ear is somewhat more pointed and the notch a little more prominent. The feet of this specimen are shorter and the digits are very strongly curved; the third outer lower incisor is distinctly larger than others. Whether the material really represents three varieties or merely individual variation I cannot say. The Jolo specimen was shot in the forest on Bud Daho. The specimen was flushed from a few dead leaves in a small shrub. The specimen flew slowly and alighted near-by, where it was shot. The Palawan specimens were in a group of seven that were found flying together in daylight. They would

264

PHILIPPINE LAND MAMMALS

alight in a group of dead leaves in shrubbery, then would take flight, the group flying slowly up and down, back and forth, within a radius of a meter. Again they would move off a few meters and perform gyrations in a narrow radius. Then they would return to the bunch of dead leaves. I was unable to determine whether or not they were breeding. The type specimen of Kerivoula pellucida Waterhouse was collected - by Hugh Cuming, in the Philippines, between 1838 and 1842. The exact locality is unknown. Subfamily MINIOPTERIN.JE

Miniopteri DOBSON, Ann. & Mag. Nat. Hist. (part.); Cat. Chirop. Brit. Mus. (1878) 170 Miniopter·t£ TROUESSART, Cat. Mamm. (1898) Miniopte?-int£ MILLER, Bull. U. S. Nat. Mus. 57

IV 16 (1875) (part.).

349

134. (1907) 227.

Vespertilionidre with median lobe of presternum enormously developed, and forming the greater part of the bone; scapula :with the coracoid straight, directed conspicuously inward; the second phalanx of third finger much elongated. The family as hete understood includes only the genus Mi-' ~iopterus Bonaparte. Genus MINIOPTERUS Bonaparte

Miniopf)erus BONAPARTE, Iconag. della Faun. Italica 1 fasc. 22' (1837); DOBSON, Cat. Chirop. Brit. Mus. (1878) 347; MILLER, Bull. U. S. Nat. Mus. 57 (190'7) 227; HOLLISTER, Philip. Journ. Sci. §, D 7 (1912) 18.

Miniopteris GRAY, Ann. & Mag. Nat. Hist. III 17 (1866) 91. Minyopterus WINGE, Jordf. og nulevende Flagermus (Chiroptera) fra Lagoa Santa, Minas Geraes, Brasilien (1892) 36. Minneopte?"us LAMPE, Jahrb. Nassau Ver. Naturk Cat. Saugeth-Samml. 53 (1900) 12.

Small bats with simple muzzle and lips; tail well developed, completely involved in the large interfemoral membranes; ears small; tragus moderately high, slender, slightly curved forward at tip; second and third metacarpals about equal, second phalanx of third finger nearly three times as long as first. Upper incisors low .and robust, subequal, the outer lower than inner but with slightly greater cross section, each tooth simple; apex of inner rather slender and directed strongly inward, that of outer broadly pointed, and nearly erect, teeth of each pair broadly in contact, but outer incisor separated from canine by distinct space; lower incisors forming a rather strongly eonvex row, their crowns trifid, the lobes of first and second subequal,

MINIOPTERUS

265

middle lobe of third much the largest, the outer almost obsolete; canines slender, not peculiar in form, the upper with a very slight cingulum, the lower with cingulum well developed and forming an indistinct anterior basal cusp; premolars showing no special peculiarities except that the anterior upper tooth is unusually large, its shaft more than one-third as high as that of canine, to base of which its crown area is nearly equal; lower premolars high and slender, with subterete though slightly four-sided shafts; molars normal, though outer section of first and second upper teeth somewhat narrowed; hypocone lacking or barely indicated; third upper molar with metacone and three commissures well developed, its crown area more than half that of first and second upper molars; lower molars with all cusps present, rather high and slender. Skull with large, high brain case, the anterior portion of which rises distinctly above occiput; rostrum slender and small, not specifically reduced or peculiar in form, its dorsal surface frequently with a slight median longitudinal concavity between orbits; sagittal crest low, usually. rather distinct, not sending forward any branches to supraorbital regions, which, with anterior border of orbits, is smoothly rounded; palate slightly concave, both longitudinally and transversely, a noticeable foramen near its border close to inner extremity of each posterior molar; audital bullre small, covering more than half the surface of cochlere; floor of brain case forming no noticeable angle with plane of palate; basisphenoid pits indicated. (After Miller.) . 2-2 1-1 2-2 3- 3 Dental formula : 1. 3-3' c. 1-1' pm. 3-3' m. 3-3 =36. Some twelve species are recognized. These are distributed from northern Africa and southern Europe to Asia, the Malay Archipelago, and the Philippines. The following additional characters mark the genus: Top of head much elevated; face short, concave in .jts longitudinal direction; muzzle obtuse, not much depressed; nostrils near together with their upper and inner margins slightly proj ecting, the space between these projecting parts being slightly emargin ate ; the notch does not pass down through upper lip which is entire and prominent; nostrils crescent-shaped; from outer side of each a vertical notch or groove passing through lip but leaving its central portion entire and slightly projecting; lateral parts of lips thick and overhanging; lower lip with a clearly defined reflex central portion; ears angular, round, short; tragus

266

PHILIPPINE LAND MAMMALS

short; tail as long as head and body; calcar short; middle phalanx of second and third finger very short; fur thick and soft. Key to the Philippine species of Miniopterus Bonapa?路te. a'. Smaller; forearm, 35 to 41 millimeters; wing membranes attach near end of tibia. b 1. Head short.; fur on inter femoral membrane extends as far back as end of third caudal vertebra; fur black throughout, the hair tips sometimes grayish; forearm, 40 millimeters; tail, 46.2. Miniopterus pusillus Dobsen (p. 266). b '. Muzzle longer; fur on interfemoral extends to a line drawn between the t.wo ankles; tibial covered with fine hairs; a small patch of short fur on wing between elbow and forearm, color dark reddish brown, darker at base of hairs, grayish at tips; forearm, 40 millimeters; tail, 42.5 .. ________________ Miniopterus australis Tomes (p. 268). b 3 . Much smaller; inter femoral haired as in M. pusillus; dark blackish brown without light underfur; forearm, 35 to 36 millimeters; foot, 7 _______________________________________ _______ MinioptelJ"us paululus Hollister (p. 269). b '. About as in Miniopterus aust1"alis, except thalt wing membranes attach one-third length of tibia from end; foot, 7.2 millimeters; forearm, 39.5 ______________ . _____ , _________ Miniopterus tibialis Tomes (p. 272). a 2. Larger; forearm, 44 to 53 millimeters. b'. Head moderate; rostrum broad; foot, 7.2 to 8.4 millimeters; forearm, 40 to 44.3 ____ Miniopterus eschscholtzii (Waterhouse) (p. 272). b 2. Head comparatively large; rostrum broader and more obtuse; foot, 10 to 11 millimeters; forearm, 51.9. M~niopterus tristis (Waterhouse) (p. 275). MINIOPTERUS PUSILL us

Dobson

Miniopterus pusillus DOBSON, Monog. Asiat. Chirop.

(1876) 1,62; MATSCHIE, Sitz.-Ber. Ges. Nat. Freunde Berlin (1898) 39 (Manila); HOLLISTER, Philip. Journ, Sci. 搂 D 7 (1912) 18; Proc. U. S. Nat. Mus. 46 (1913) 336. Mimiopterus australis TO~ES, Proc. Zool. Soc. London (1858) 125 (part.) .

Description of the species.-(From No. 370, E. H. Taylor collection; collected at Baguio, Benguet, July 20, 1923, by E. H. Taylor.) Rostrum rather broad, nostrils only moderately prominent, separated by a narrow depres'sion; upper lips glandular with a raised glandular area running back towards eyes, separated medially by a depression; a distinct groove below eye; hair on forehead grows to within 1.5 millimeters of extreme tip of snout; on underside of upper lip on each side a small serrate compressed papilla; a papilla on inside of lower lip larger and less compressed than upper; lower lip with a naked area at tip, the parts adjoining glandular, compressed.

MINIOPTERUS

267

making lip sharp and somewhat serrate; glandular area of upper lip covered with scattered hair and longer vibrissre; ear much shorter than head, truncate, somewhat funnel-shaped, inner edge attaches almost directly above eye and curves strongly to highest point which is broadly truncate, and almost straight across top; below top is a slight emargination from which ear curves forward to a notch, beyond which is a free lobule turned in directly in front of tragus; tragus not. tapering, rounded at tip about same width its entire length, inner surface concave, outer convex; no free lobule at base but a distinct thickening, the anterior face of which is smooth; short scattered hairs present on upper and under surface of antebrachial membrane; wing membranes naked except near body; tibia with extremely minute scattered hairs; interfemoral membrane furred at base, with elongate hair in mid-region along tail vertebra for more than two-thirds length of tail; underside of interfemoral with very fine hair over basal two-thirds; feet slender, elongate, metacarpals as long as phalanges with claws, second toe with a shorter, less-compressed claw than other digits; penis slender, elongate; wing membranes attach at least 2 millimeters from lower end of tibia; calcar attached to tibia for about 3 millimeters; tail entirely involved in interfemoral membrane; fur on shoulders, 8 millimeters; on breast, 5.5. Color.~Deep blackish brown (No.3 Ridgway with more bla.ck); hairs varying but little in shade at base, tips very slightly more brownish; fur on head with a very slight brown suffusion; below on throat and breast black-brown of a slightly more brownish shade than on back; membranes dark brown, underpart of inter femoral lighter than elsewhere; feet and tibia brown, claws white; tragus dark grayish near summit, creamwhite below. Skull and teeth.-Brain case rather broadened, and somewhat flattened with a transverse depression across posterior part, declivous anteriorly; rostrum broad with a median shallow longitudinal groove; narial notch rather narrowed posteriorly; anterior palatal notch more rounded, a little wider than deep. Inner incisor slanting inward, with a cingulum cusp; second incisor rises from a lower point on the bone and reaches an elevation equal to that of cingulum cusp; canines elongate, not or scarcely curved; anterior surface with a well-defined groove; cingulum developed but no accessory cusps; first premolar relatively large, not in contact with "adjoining teeth, placed internally to tooth row; molars without specific peculiarities; lower

268

PHILIPPINE LAND MAMMALS

incisors crowded, inner pair tricuspid; second pair much thicker, less distinctly tricuspid; third pair as broad as long, rather molariform, in contact with canines; six incisors form an angular series; lower canine smaller than upper, cingulum forming a cusp on anterior inner edge of tooth; two anterior premolars of same size, shape, and elevation, much smaller but of same general sh,~pe as third; second premolar not in contact with anterior or posterior; molars without specific peculiarities. M easurements.-See under Miniopterus pa.u~ulus. Remarks.-I have tentatively placed this specimen under Miniopterus pusillus. The description of M. p~tsillus omits certain important characters. The specimen listed by Dobson * differs in having shorter tibia and shorter fingers. The black (blackbrown) coloration is the same; the small haired area near the junction of the fifth finger and forearm is small and the hairs very few and short. The first Philippine specimens were collected by H. Cuming (three .specimens) . I am not certain whether these were the' types or not. I have examined only the single specimen. It was collected at Baguio on the veranda of the Country Club house, at night. MINIOPTERUS AUSTRALIS Tomes

Miniopte'l"us aust?'alis TOMES, Proc. Zool. Soc, London (1858) 125; PETERS, Monatsb. Akad. Wiss. Berlin (1861) 7'11 (Sant.ik, Camarines Sur, Luzon); DOBSON, Cat. Chirop. Brit. Mus. (1878) 351;. HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 18; Proc. U. S. Nat. Mus. 46 (1913) 335.

Description of the species.-Related to Miniopterus blepotis and M. eschscho~tzi but differs in having the face more' hairy, the ears smaller, the thumb distinctly smaller, and in being actually smaller itself; fur of underparts encroaches somewhat on membranes, while in M. blepotis they are free of hair; muzzle is distinctly longer than in M. pusillus; interfemora~ with hair as far back as a line from one heel to the other; tibia covered with fine hairs and a small area of short hairs on wing membrane between elbow and forearm. Color.-Fur with decided rufous tinge which is given by tips of hairs, being paler and redder than color of base. This is absent in some specimens and they have the dark color of M. blepotis.

* Catalogue

of the Chiroptera.

269

MINIOPTERUS

M easurements of type and cotype specimens of Miniopterus australis Tomes. [Measurements in millimeters.]

1=--1- -1 Male.

H ead and body .. . _______ ____ - - _---Tail. .. __ _______ ______ ___ __ - - -- ---. Head ___ ______________ __ _________ _ Ears _____ ____ __________ ___ _______ _

Tragus . ______________________ _____ Forearm . . ____ __________ __________ _ Longest finger ... ____ ____ __________ _ Fourth finger _______ ______ ____ ___ __ Tibia ___ _____ ____________________ _ \

F oot and c1aws .. ________________ __ _ Expanse of wing . . _____ ____ ______ ___

Male.

F emal e.

47.5 44.3 44. 3 42.2 15 . 8 16 6. 3 --- - ---- -5 . 7 - - - - -----39 . 7 39 I 76.2 --- ------46 .4 _____ _____

42 . 2 42.2 15.7 7 4.2 36 77 . 6 42 . 2

i----------

14 . 7 7.9

14.7 9 . 9 __________ 296 . 2

279.4 - - - - - - - - - -

Male.

42.2 38 15 .2 7 4. 4 38 77.6 42 . 2 14 . 7 7.6 277 .4

40 36 15. 2 6 .3 4. 2 30

79 .6 42 . 2

I I

'ill 8.4

~79.4

Remarks.-The type locality; is Loyalty Islands. The types were collected by Sir G. Grey and are in the British Museum of Natural History. The species is listed on the strength of a record by Peters, from Camarines, Luzon. MIN:IOPTERUS PAULULUS Hollister

Miniopterus paululus 311.

HOLLISTER,

Proc. U. S. Nat. Mus.

(1913)

Description of the species.-(From No. 356, E. H. Taylor collection; collected in Montalban Caves, ,M ontalban, January 5, 1923, by Albert W. Herre, W. Schultze, and E. H. Taylor.) Rostrum relatively broad; nostrils moderately prominent, openjng forward and outward, narrowly separated by a depression; area below nostrils naked; rostrum glandular on each side of "nose," the areas separated from nose by a groove; a deep groove below eye running forward; lips compressed outwardly, edges sharp and serrate above and below; median portion of rostrum depressed with a slight median elevation; fur of head r eaches to within 2.3 millimeters of extreme tip of rostrum medially ; glandular areas on rostrum with hair of varying length; on underside of upper lip a small toothlike papilla and a similar one on lower lip directly in front of a small pocket in which upper canine rests; lower lip with a narrow naked area anteriorly; ear much shorter than head, inner edge rounding to near summit, in front of which is a slight emargination; tip blunt, below which, the margin runs nearly straight to basal notch, in front of which is a well-defined anti tragal lobe; tragus widest at. ex-

270

PHILIPPINE LAND MAMMALS

treme base, the width above varying but slightly to tip, which is slightly attenuated and rounded; inner edge slightly concave, outer convex; lower edge folds over, forming a very slight, marginal, free flap; inner part of ear membrane with scattered elongate hairs except in central portion; outer surface furred to near summit; tragus with a few hairs at base; fur on top of head, 4.5 millimeters long; on shoulders, 6.5; on breast, 3.9 to 4.2; antebrachial membrane with an area of rather elongate hair near joint; below, a few exceedingly minute scattered hairs; a few scattered hairs on wing membranes as far as a line between elbow and ankle; on membranes below, hairs confined to area along body; interfemoral membrane with a definite growth of hair as far as antepenultimate vertebra; hair sparse and short below, but continues to near tip of tail; entire tail involved in membrane; feet rather small, metacarpals slightly longer than toes; .second outer toe with a rounder, shorter, less-compressed claw than other toes; a few hairs at base of claw on first finger; wing membranes attach at least 1 millimeter above end of tibia; calcar attached to tibia for about 3 millimeters; testes in postanal scrotal pouch. Color.-General color above between Vandyke and bone brown; below, between wood brown and buffy brown, slightly darker on throat and neck; wing membranes darker than pelage; tragus whitish cream without any brown; basal' fur on belly deep Vandyke brown; feet dark brown above, claws olive-horn color; toes nearly black; below, flesh color; ear membranes somewhat lighter than fur. Skull and teeth.-Skull somewhat flattened on brain case with a transverse depression posteriorly; a rather prominent crest from base of rostrum on brain case; a median longitUdinal depression on rostrum, narial notch narrowing gradually posteriorly; palatal notch as broad as deep or broader; a posterior median extension on palate. Upper inner incisor with a small secondary cusp; outer incisor not rising higher than cusp; canine with an anterior groove and a 's hallow posterior groove; no cingulum cusp; first premolar pushed out of tooth row, in contact with canine and second premolar; inner lower incisors trifid, compressed; second pair trifid, thickened, and distinctly larger than first pair; third pair molariform, cusps indistinct, more than twice as large as second incisor; canine with a very slight anterior internal cingulum cusp; second premolar slightly larger than first, in contact with first and third premolars.

271

MINIOPTERUS MeasU'~'ements

of Miniopterus pauhtlus Holliste?' and M.

pusill~ts

Dobson.

[Measurements in millimeters.]

Miniopter1l8 paulul1ts.

Miniopterus pusilZ"s. E. H. Taylor collection. United E. H. Tay!_ _ _-,-_ _---.---_ _ _ !States Na- lor collectional tion, No. 357, 356 , 371, Museum. 370, female. male. male. Type. male. ________________ - - - - - - ----1----1----

Head and body .. _________________ ____ ___ _ Tail ... _________________________________ _ Height of eaL ________________________ __ _ Forearm .. __________________________ - ___ _ First finger and claw.. ___________________ _ Second finger, metacarpaL _______________ _ Third finger: Metacarpal _________________________ _ First phalanx _______________________ _ Second and third phalanges .. ________ y Fourth finger: " MetacarpaL ________ ________________ _ First phaIanx ___________________ ____ _ Second phalanx _________ ____________ _

38 35

39 38 9.9 38 5.1 30

29 7.8 28

30 8 .1 29

30 8 27

29 6.7 14

29 6.2 13

27 6 6.5 14.1 10 4 7.2 5.3

26.5 6.2 6 15 11 4 7.2 5.4

13.5 7 4.6 6.8 7.4 5. 5 6 .1 6 .4 9 .6

13 7.1 4.3 6.8 7.1 5.4 5.9 5.8

35 5

13.5

Fifth finger: Metacarpal _________________________ _

26 First phalanx_ . _____________________ _ 6 Second phalanx _____________________ _ 6 Tibia _________________________________ _ 13 Calcar _________________________________ _ 11 Metatarsals .. ___________________________ _ 3.3 Foot and elaws.. ________________________ _ 6.3 Penis ___________________________________ ... _____ _ Skull: Greatest length _____________________________ _ Zygomatic width ______________________ . _____ _ Breadth of rostrum at third premolar __________ _ Height of brain case _________________________ _ Width of brain case.. __________________ . _____ _ Palate, median length. ______________ . ________ _ Upper tooth row, entire ______________________ _ Lower tooth row, entire __________ . ___________ _ Mandible .. ________________________ .. _______ _

42 37 10.5 37

-------30

---------------------------35

-------------- ----31

------------------29.5

------------------27

44 48 10.5 43.8 6 38 39 10 36 36 9 16 34

9 ---------10 ---------19 ---------13 -------- ----------- ---------7

---------13.4 7

------------------6.6

---------6.1 6.5 9.4

9.1 6

13.9 7.8 I)

7 8 5.3 6.3 6.3 9.8

---

Remarks.-The three sI?ecimens listed appear to agree very well among themselves in external characters except that in No. 371 the ear has the tip more rounded, and the emargination is more marked before summit is reached. The three specimens, Nos. 357, 371, and 356, are from the caves in Montalban Gorge, Rizal Province. The type and paratypes of the species were collected by J. B. Steere on Guimaras Island, December, 1887. The specimens, five in all, are in the United States National Museum (No. 105493 type). This is the smallest Philippine species of the genus.

272

PHILIPPINE LAND MAMMALS MINIOPTERUS TIBIALIS Tomes

Miniopte?'"US tibialis TOMES, Proc. Zool. Soc. London (1858) 126; MATSCHIE, Sitz-Ber. Ges. Nat. Freunde zu Berlin (1898) 39 (Manila); HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 18; Proc. U. S. Nat. Mus. 46 (1913) 336. Miniopterus sch~'eibetrsii DOBSON, Cat. Chirop. Brit. Mus. (1878) 348 (part.) .

Desc1'iption of the species.-Similar to Miniopte1'us blepotis and M. eschscholtzi but smaller; differs from other Philippine species in having wing membranes attach to tibia a third of its leng~h from distal end, calcar attached to tibia for length of free portion. Color.-Dark brown, sometimes with tips of hair paler; fur below bicolored, dark brown at base and a distinctly lighter shade at tip; in pubic region hair lighter entire length. Fur color apparently variable. Measurements of Miniopterus tibialis Tomes. mm.

Head and body Head Ear Tragus Forearm Third finger Fourth finger Tibia Foot and claws Alar expanse

63.4 14.7 7.35 4.2 40.1 73.9 42.2 14.7 7.35 292

RemOffks.-The species is included here on the strength of a single record of Matschie from Luzon. The type locality is Amboyna. . MINIOPTERUS ESCHSCHOLTZII (Waterhouse) V eS1Je1,路 tilio eschscholtz'ii W ATEnHOUSE, Proc. Zool. SOC. London (1845) 121 (Philippine Islands; Luzon). Miniopterus blepotis TOMES, Proc. Zool. Hoc. London (1858) 117 (part.); PETERS, Monatsb. Akad. Wiss. Berlin (1861) 711 (Santik and Paracale, Luzon). Miniopterus schreibersii DOBSON, Monog. Asiat. Chirop. (1876) 160; Cat. Chirop. Brit. Mus. (1878) 348; HOLLISTER, Philip. J ourn. Sci. 搂 D 7 (1912) 18; ELERA, Contrib. Fauna Filipina (1915) 109. Miniopterus elwhscholtzi路i HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 311 (Irisan, Benguet, and Malabon, Rizal, Luzon Island; Guimaras).

Description of the species.-(From No. 584, E. H. T. collection; collected March, 1923, near Caldera, Zamboanga, Mindanao,

MINIOPTERUS

273

by E. H. Taylor.) Forehead strongly arched; rostrum glandular; nostrils moderately prominent, separated by a distinct groove; fur of head covering all of rostrum except 2 millimeters of tip; a naked area below nostrils and a narrow naked area on lower lip; on inner side of upper lip a compressed serrate tubercle near angle of mouth; a rounded tubercle in front of pocket formed for tip of canine; margin of eye glandular, protruding;' a deep horizontal groove below eye; ears shorter than head; anterior attachment above eye, inner margin convex forwards, then curving back horizontally towards tip, the position of which is shown by a slight flattening of upper third of outer margin; outer margin rather straight with a deep notch in front of base of tragus and terminatjng abruptly in a lobe turned inward near angle of mouth; tragus with nearly same width throughout its length, rounded terminally, inner margin concave, the outer convex; at base the edge of tragus extending forward as a small free lobe; two slight folds across face of ear; tip of muzzle with sparse scattered hair of varying length, and a few longer vibrissre rising from glandular prominences; a small area of hair on antebrachial membrane; lower edge of forearin with short hair continuing to base of thumb; membrane between second and third fingers haired to tjp of first phalanx on third; an area of hair along base of metacarpal of fifth finger and on membranes adjoining; upper surface of interfemoral haired, except posterior fourth or fifth; tibia and wing membrane adjoining, haired to heel; below, interfemoral haired more or less about half its area basally; metatarsals longer than toes; second toe with a shorter, lesscompressed claw; tail involved in membrane its entire length. Color.-Above, fur between Vandyke brown and bone brown (Ridgway) ; on belly the basal part of fur bone brown, distal part lighter brown, which in pubic region continues to base of hairs; wing membranes blackish; ear membranes lighter with darker edges; tragus dark on upper part; skin on face brownish; tibia brownish; toes blackish above; underside of interfemoral lighter than upper side. Skull and teeth.-Skull similar to those of other Philippine species; posterior part of brain case distinctly lower than anterior part, with a transverse depression on posterior half; brain case sloping v~ry abruptly to base of rostrum; a slight longitudinal groove on rostrum; narial notch ' about as wide as deep; rostrum broad; palate arched, less so than in smaller species; a median posterior extension on palate. 259453-18

274

PHILIPPINE LAND MAMMALS

Measurements of Mimiop-terus eschscholtzii (Wa.tM路house). [Measurements in millimeters.] E. H. Taylor collection No. and sex.

584, male.

585, famale.

586, male.

383, 378, fefamale. male.

358, female.

118, male.

216, male.

- - - - - - -- - - - - - - - - - - - - - - - - - - - - Head and body __________ "- ________ 51 Tail _____________________________ 47

50 46

Height of ear. ____________________ 11 Forearm_ _ _ _ _ _ _ _ __ _ _ _ _ _ _ __ __ _ _ __ _ 43.2

First finger and claw______________ 7.2 Second finger, metacarpal. ___ . _____ 38.5 Third fin ger : Metacarpal.. _ ____ _ _ _ _ __ __ __ _ _ 39 First phalanx. ________________ 10.2 Second and third phalanges ____ 37.6 Fourth finger: Metacarpal.. _ _ _ _ _ __ _ _ _ ____ _ __ 37 First phalanx.________________ 8 Second phalanx. ______________ 16 Fifth finger: Metacarpal.. _ __ _ _ _ _ __ _ _ _ _ _ _ _ 34 First phalanx.________________ 8.5 Second pbalanx. __ _ _ __ ___ __ _ __ 8 Tibia. _______________________ ~__ _ 18 Calcar. _ _ _ __ _ _ _ _ _ _ ___ _ _ _ __ _ ___ _ _ _ 16 MetatarsaIa_ __ __ _ ____ ___ _ _ _ __ __ __ 6.2 Foot and claws_ __ _ _ _ _ ___ __ _ ____ _ _ 8.4 Penis. _ _ __ _ _ _ _ _ __ _ __ ___ _ _ _ _ ___ _ _ _ 6.2

50 46 10.4 43.4 7 38.1

50 48 43.2 7.2 38

51 46 10.7 45 6.8 38

52 47 11 44.2

39.2 10 35

38.2 9 36

38 8.5 16

36.2 8.2 16

34 8.5 8.2 18 13.8 4.9 8.9

33 8.7 8.8 18.2 14 4.5

7 39 40 10

50 45 10 43.8 5.1 38

53 48 10 44 7.6 39

39 9.2 36.2

40 10 37

39 9.9 35.2

39 10 84.9

36 7.8 16

37 8 16

37 8.2 17

37 8 16

37.5 8.2 16

32.3 8.3

33 8.1 8.4 18

33 8.2

33 8 7.6 18 13

33.6 8.6

8 17.3 14 4.9 9

7 Skull: Greatest length_ _ _ __ _ _ _ _ _ _ _ ___ 15.1 16.2 Zygomatic width .. ___________ _ 8.9 9 Breadth of rostrum at first pre6 6 molar. Height of brain case. ___ . _____ _ 8.6 8.2 Width of brain case __________ _ 8.5 8.6 Palate, median length ________ _ 6.6 6.6 Pper tooth row, entire .. ______ 7.2 ___________ _ 7.4 Lower tooth row, entire .. -- -- __ 1__ - - -7.3 Mandible_ _ _ _ _ __ _ _ __ _ _ _ ____ __ _ ___ __ 11

14 4.8 9

8 17.2 16

8.8

7.9 7

8 17 16 6 9 8

16 9 6 8.6 8.6 7 ___________________ _

-----ld--------

7.4 7.81 -----___________________ _ 12 11.1 _____________ _

Inner incisor slanting inward with a rather indistinct posterior cusp or heel; second incisor reaching somewhat above summit of heel, widely separated from canine; canine with an anterior longitudinal groove, posteroexternal groove slight or lacking; first premolar large, pushed in somewhat out of line of tooth row; molars without specific peculiarities; inner lower incisors smallest, trifid; second pair same width but noticeably thickened; third pair as broad as long across top; cusps on latter tooth rather indistinct; canine with a slight anterointernal cin-

MINIOPTERUS

275

gulum cusp; first two premolars very nearly of same size; other teeth without specific peculiarities. Remarks.-Specimens 358, 378, and 383 differ in the color of the fur which in 383 is auburn brown on head and fore part of shoulders and somewhat darker brown posteriorly; belly fur dark Vandyke brown basally, buffy brown distally. This and the two other specimens were preserved in al~ohol and the color may be faded; the other two specimens are similar but distinctly of darker shades. Six specimens from Zamboanga, Nos. 584 to 589, are similar to the described specimen; there is a larger seminaked area on the lower jaw and the median rounded glandular tubercle is more pronounced; otherwise the variation is small. Specimens 116 to 204 were obtained in a cave near Odiongan, Tablas. These specimen's seem to have the wing membranes blacker; the upper edge of the ear and the toes above are especially darker. That these specimens are correctly referred to Miniopter'lls eschscholtzii (Waterhouse) I am not absolutely certain, since a character mentioned by Waterhouse "on each side of the head is a naked groove, which runs over the eye," is lacking. Certain other characters present in these specimens are either lacking or are not mentioned by Waterhouse. The type of this species, in the British Museum of Natural History, was obtained by H. Cuming in the Philippines; the exact locality is no longer known. Hollister * gives the following measurements for Philippine specimens: Forearm, 43.9, 40.9, 44.1, 44.3; third digit, metacarpal, 38, 39.5, 35.5, 38.2, 38.2; foot with claws, 7.6, 8.4, 7.5, 7,5,7.2; skull, condylobasallength, 14.2, 14.2; zygomatic breadth, 8.4, 8.4; breadth of brain case, 7.6, 7.6; mandible, 10.8, 10.5; maxillary tooth row, 7,7; mandibular tooth row, 7.5, 7.5. These specimens are from Irisan, Benguet, 2; l\'IalaQon, Luzon, 1; Guimaras, 2. Measurements in the order given. Skull measurements are for Benguet specimens only. MINIOPTERUS TRISTIS (Waterhouse)

Vespe?'tilio t?'istis WATERHOUSE, Proc. Zool. SOC. London (1845) 3 ; GRAY, Zool. Voyage Samarang (1850) 31; WAGNER, Suppl. Schreb. Saugeth. 5 (1885) 743. Miniopterus tristis TOMES, Proc. Zool. Soc. London (1858) 124; PETERS, Monatsb. Akad. Wiss. Berlin (1861) 711; (Santik, Luzon) ;

* Proc.

U. S. Nat. Mus. 46 (1913) 311.

276

PHILIPPINE LAND MAMMALS ELERA, Cat. Sisto Fauna Filipinas 1 (1895) 14 (Luzon, Cebu, Samar); HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 19; Proc. U. S. Nat. Mus. 46 (1913) 311 (Lubang).

Description of the species.-Forehead much arched; muzzle short and obtusely rounded, very broad and hairy; lower lip with a narrow transverse naked area at tip; nostrils sublateral, moderately separated, with a slight depression between them; ears moderate, rounded, with anterior margin nea~ly straight; tragus curved and obtusely rounded at point; about 5.2 millimeters long, 3.5 wide; wings rather narrow, with black membranes; membranes attach laterally, reaching heel of foot, which has metatarsus narrow and long; distance from heel to base of toes exceeding length of toes; toes rather short, equal, claws short and but little curved; calcar short, bent back; legs rather long, interfemoral membrane ample, naked above and below excepting at base; tail involved in membrane its entire length; fur very dense, short, extending far on muzzle leaving only tip covered with very short hair. Color.-Almost uniform sooty black, hairs uniformly colored to root; tips of hairs of belly slightly lighter. Measurements

of Miniopterus t?'istis (Waterhouse). mm.

Length, from apex of nose to base of tail Tail Head Ears Tragus Forearm Longest finger Fourth finger Thumb Tibia Foot and claws Expanse of wings 路n

Original measurements are given in inches and lines.

60.9 66.04 21.6 16.5 7.62 53.3 104.1 73.6 8.38 19.05 11.4 400 After Dobson.

Remarks.-The type, in the British Museum, was collected in the Philippines by H. Cuming; the exact locality is unknown. Tomes, remarking upon the type specimen, states that the muzzle of this species is relatively broader and more obtuse than in any other species of the genus; and this peculiarity, together with its superior size, is sufficient at once to distinguish it from M. blepotis, which it otherwise resembles. Tomes also mentions the presence of glands on the lips which differ in degree of development in different specimens, thus pro-

MYOTIS

277

ducing a slight difference in the position and direction of the nostrils. He further states that the color is deep brown, appearing sooty black in spirit. Hollister records the measurements of a specimen from Lubang as follows: Forearm, 51.9 millimeters; third digit metacarpal, 47; foot with claw, 10.2; skull, condylobasal length, 18.5; zygomatic breadth, 10.4; breadth of brain case, 8.9; mandible, 14.8; maxillary tooth row, 9.1; mandibular tooth row, 10. Subfamily VESPERTILIONIN...!'E

Vespertiliones DOBSON, Cat. Chirop. Brit. Mus. (1 878) 168. Vespertilioninre and Plecotinre MILLER, N. Am. Fauna (1897) 46 and 54.

Sternum slender, its entire length considerably more than twice greatest width of presternum ; median lobe much smaller than body of presternum; six ribs connected with sternum; seventh cervical vertebra not fused with first dorsal; scapula with coracoid curved outward; nost rils simple, sometimes tubularly elongated but never margined by special outgrowths; lower incisors 3-3. The distribution is world wide. The genus contains about thirty-three genera, six of which are represented in the Philippines. Ke1! to the Philippine genera, of the Vespertilioninre. a, '. Incisors, four upper, six lower; small bats. b ' . Teeth 38; no callosities on thumb or foot ........ Myotis Kaup (P. 277). b ' . Teeth 34; skull not gTeatly flattened. c ' . Callosities on thumb and fooL ..... ..... Glischrop1ts Dobson (p. 292). c 2. No callosities on thumb or foot. d '. Fifth finger normally long, much 10ngeJ.' than metacarpals of third or fourth .................................. Pipistl"ellus Ka up (p. 294 ). d'. Fifth finger greatly shortened, scarcely longer than metacarpal of third or fourth finger.. .............. Pte?"ygistes Ka up ( p. 301). c". Teeth 32; callosities on thumb and foot; skull gr eatly flatten ed. Tylon ycteTis Peter s (p. 303). (( ' . Incisors, two upper, six lower; teeth 30; no callosities on thumb or foot; medium-sized bats .................... S cotophilus Horsfi eld (p. 305). Genus MYOTIS Kaup

Myotis KAup, Skizzirte Entw.-Gesch. u. Natiir. Syst. d Europ. Thierw. 1 (1829) 106; MILLER, Ann. & Mag. Nat. Hist. VI 20 (1897) 382; N. Am. Fauna (1897) 55; Bull. U. S. Nat. Mus. 57 (1907) 200; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 16. Nysta,ctes KAUP, Skizzirte Entw.-Gesch. u. Natiir. Syst. d Europ. Thierw. 1 (1829) 108. Leuconoe BOIS, Isis (1830) 256.

278

PHILIPPINE LAND MAMMALS

Vespertilio KEYSERLING and BLASIUS, Wiegm. Archiv fUr Naturg. 1 (1839) 306 (non Linnreus 1758); DOBSON, Cat. Chirop. Brit. Mus. (1878) 284 (non Linnreus 1758); H. ALLEN, Monog. Bats N. Am. (1894) '70. Selysius BONAPARTE, Iconog. della Fauna Italica, Intr(}. (1841) 3. Capaccinrius BONAPARTE, Iconog. della Fauna ltalica (1841) 3. Trilatitus GRAY, Ann. & Mag. Nat. Hist. 10 (1842) 258; GERVAIS, Dict. Univ. d'Hist. Nat. 13 (1849) 213. Brach",yotis KOLANATI, Allgem. deutsch. Natur. Zeit. Dresden neue folge 2 (1856) 131 (non Gould). Isotus KOLANATI, Allgem. deutsch. Natur. Zeit. Dresden neue folge 2 (1856) 131. Pternopterus PETERS, Monatsb. Wiss. Akad. Berlin (1867) 906 (subgenus of Vespertilio). Exochurus FITZINGER, Sitz. Kais. Akad. Wiss. Wien. Math-Natur. 62 (1870) 75. Aeorestus FITZINGER, Sitz. Kais. Akad. Wiss. Wien. Math-Natur. 62 (1870) 427. CO'l'/Ul,stes FITZINGER, Sitz. Kais. Akad. Wiss. Wien. Math-Natur. 62 (1870) 565.

Small bats withQut nQse leaf; upper incisQrs well develQped, subequal, clQsely crQwded, the crQwns higher than IQng, subterete; inner with a distinct PQsteriQr secQndary cusp, Quter with a well-develQped CQncave surface directed toward canine, frQm which it is separated by a space not quite equal to diameter Qf bQth incisQrs tQgether; IQwer incisQrs with crQwns abO'ut equal in length, fO'rming a cO'ntinuQus, strQngly convex rO'w between canines, first and secO'nd with rather narrQW, trilQbed cutting edge, third much wider, subterete, with three Qr fO'ur tubercles; canines well develQped, simple, with distinct thQugh rather small cingulum and nO' secQndary cusps; cheek teeth bQth abQve and belQw strictly nQrmal, first and secQnd upper mQlars rather narrQW Qn inner edge and with hypQcQne absent Qr very imperfectly develQped, third upper mQlar with three cQmmissures and fQur Qr five cusps; skull slender and lightly built withO'ut special peculiarities Qf fQrm, rQstrum nearly as IQng as brain case; depth O'f brain case, including audital bullre, abQut equal to' mastQid breadth; sagittal crest IQW but usually distinct, palate deeply emarginate in frQnt, abruptly cQntracted behind, sides Qf its PQsteriQr extensiQn parallel; audital bullre well develQped and cQvering mQre than half surface Qf cQchlere, but simple in fQrm and nQt very large, their diameter abQut equal to' distance between them; ear well develQped, slender, QC-

279

MYOTIS

casionally rather large; tragus slender and nearly or quite straight; foot variable in size, but never peculiar in form; tail about as long as outstretched hind leg; interfemoral membrane large, its surface furred at extreme base above. (After Miller.) . 2-2 1-1 3-3 3-3 Dental formula: 1. 3-3' c. 1-1' pm. 3=3' m. 3- 3 = 38.

Remarks.-About eighty-five species * are known and these are world wide in distribution. They are of very primitive form. Miller states: "It not only retains the maximum number of teeth known in the order, but it also has the slightest possible degree of special modification in external form." Key to the Philitppine species of Myotis Kaup. a t. Large; head and body, 55 to 63 millimeters. b'. Forearm, 49 millimeters; tail, 52; membranes bicolored; feet moderately large, 10 to 12 millimeters; wing membranes attach to base of toes .............................. M. rufopictus (Waterhouse) (p. 279). b 2. Forearm, 46 millimeters; tail, 45; membranes unicolored; foot very large, 13 to 15 millimeters; wing membrane attached near end of tibia ................................ M.. maC?'ota?'sus (Waterhouse) (p. 281). a', Small; head and body, 33 to 45 millimeters; wing membranes attach on side of foot. b 1. Feet large; last two caudal vertebrre almost wholly without tail membrane; forearm, 36.5 millimeters; third finger, 66; fifth finger, 48; tibia, 16.5; foot, 10; wing membranes attach near heel. M. jeannei sp. nov. (p. 284). b'. Feet small; only extreme tip of tail free. c 1. Forearm, 27 millimeters; third finger, 41; membranes attach on side of foot near base of outer toe. M. patriciEE sp. nov. (p. 286). c 2. Forearm, 30 millimeters; third finger, 56; fifth finger, 40; tibia, 12; foot, 6.1; membranes attach near base of outer toe; inner edge of tragus curved """""""""""" "" M. b?"owni sp. nov. (p. 288). c". Forearm, 31.3 millimeters; third finger, 55; fifth finger, 39; tibia, 14; foot, 7; wing membranes attach to a point on side of foot halfway between base of toe and heel; inner edge of tragus straight ............................................... M. herrrei sp. nov. (p. 290). MYOTIS RUFOPICTUS

(Waterhouse)

Vespertilio rufopictus WATERHOUSE, Proc. Zool. SOC. London (1845) 3 and 8 (Philippine Islands); TOMES, Proc. Zool. Soc. London (1858) 85.

* Miller and Allen, Bull. U. S. Nat. Mus. 144 (1928) 34-37, list 58 species and subspecies for North and South America.

280

PHILIPPINE LAND MAMMALS Ke~'ivoula nG/opicta GRAY, Zoo1. Voy. Samar. No. 5 (1849). Vespe?路tilio formosus DOBSON, Cat. Chirop. Brit. Mus. (1878) 311; TOMES, Proc. Zoo!. Soc. London (1858) 82, pI. 60. Myotis formosus HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 16 (non Hodgson). Myotis rufopictus HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 309 (Malolos, Bulacan, Luzon).

Desc'ription of the species.-Fur of moderate length; forehead slightly arched; face rather long and somewhat obtuse, but not much broader laterally than thick in a vertical direction; muzzle with tip naked or with small scattered hairs; nostrils small, near together, space between them slightly depressed; glands do not approach close to upper lip but pass backwards over eyes almost to front margin of ears, and leave a central longitudinal depression along face, up the middle of which is a narrow raised ridge producing two longitudinal grooves; lower lip with a semicircular naked space at tip, with a tubercle between this point and throat; ears long, narrow, pointed, distinctly emarginated behind; tragus 10.5 millimeters long, narrow, attenuated, pointed, a distinct lobe at its outer margin close to base above which is a considerable indentation succeeded by an obtuse angle from which it passes in a straight line to tip; inner margin nearly straight; wings large, extending to base of toes; thumb long; feet large, toes fully two-thirds entire length of foot; calcar length equivalent to two-thirds distance from foot to tail; tail wholly inclosed in membrane which is furred at base, with a few scattered hairs on upper surface of membrane; outer toe with short bristly hairs on whole of its upper surface, the others hairy only at base. Color.-Fur above light dusky gray at base, tips yellowish buff; below, uniform buffy ash with a strong tinge of yellow about axilla; wing membranes black except in region of bones of fingers and large area at base, at which parts they are bright red; a straight line drawn across wing from thumb to heel would mark boundary of basal red portion; small strip of membrane above arm red, clouded with black; limbs and interfemoral membrane red; naked tip of muzzle and ears pale flesh color; hinder toes dusky at tips. Skull 'a.nd teeth.-First and third lower premolars of each side in close contact; second premolars entirely crowded out of rows and not visible from outer sides. (After the type description.)

281

MYOTIS

MeasUTements of the type of Myotis rufopictus (Waterhouse). rom.

63.4 52.9 21.1 12.6 10.5 7.8 49.8 80.4 69.7 10.5 25.4 12.6 24 334.4

Head and body Tail Head Ears Breadth of ears Length of tragus Forearm Longest finger Fourth finger Thumb Tibia Foot and claws Calcar Expanse of wings

Remar~s.-The type, in the British Museum, was collected by H. Cuming in the Philippine Islands; the exact locality is un2-2 known. Tomes states that the premolars are 2-2' Hollister,

who examined a specimen from Luzon, states that the second premolar is absent on one side and on the other very minute, not visible from without. It is probable that Tomes failed to make careful examination of the skull. There is a single specimen in the United States National Museum from Malolos, Bulacan, Luzon, collected by George C. Lewis. MYOTIS MACROTARSUS

(Waterhouse)

Vespe1路tilio macrota1'SUS WATERHOUSE, Proc. Zool. SOC. London (1845) 3-5 (Philippine Islands); Ann. & Mag. Nat. Hist. 16 (1845) 51; DOBSON, Monog. Asiat. Chirop. (1876) 130; Cat. Chirop. Brit. Mus. (1878) 290.

Myotis mac1'otarsus THOMAS, Trans. Zool. Soc. London 14 (1898) 385 (Manila); HOLLISTER, Philip. Journ. Sci. 搂 D 7 (1912) 17; Proc. U. S. Nat. Mus. 46 (1913) 310 (Iligan and Zamboanga, Mindanao).

Description of the species.-(From No. 339, E. H. Taylor collection; collected in Manila, April, 1920, by E. H. Taylor.) Adult male. Head rather flattened, a slight depression between snout and forehead; eye very small, set back near ear; nostrils small, opening near edge of lip; two narrow naked ridges separated by a shallow groove run back from nostril on snout; a shallow depressed area between nostrils; posterior to nostrils on side of snout a glandular area with several tubercles; a small tubercle somewhat in front 9f eye and one above anterior

282

PHILIPPINE LAND MAMMALS

part of eye; tip of IO'wer lip nearly or quite naked; a glandular area bO'rders sides O'f mouth; an indistinct raised glandular area O'n chin and twO' indistinct lateral tubercles and a rounded median tubercle; ears large, tip narrO'w but distinctly rounded, anterior margin strongly curved; a small triangular lobe at base of anterior margin; posterior margin of ear nearly straight to' near base where a distinct notch O'ccurs, fO'llowed by a thickened rounded lobe at extreme base; pO'sterior margin folds strongly forward and edge slightly folded back; tragus long, terminating in a mO'derately acute but rO'unded tip, its anterior margin almost or quite straight; it widens gradually, the posterior margin uneven; just below widest point a distinct notch, followed by a small, more or less angular IO'be; ear laid forward barely reaches nostril; pelage thick on head, forehead, and posterior part of muzzle; balance O'f muzzle naked except for scattered short hair; area about eyes nearly naked; chin sparsely covered with shorter and longer hairs; body and bead above heavily furred; wing and interfemoral membranes, except basal portion, and near tip of tail naked; forearm and tibia naked; a few long hairs at base of claws; underside of wing membranes naked, except near attachment to' foot; interfemoral membrane covered with extremely numerous whitish minute tubercles each with a very short hair; extreme basal portion and base of tail with longer body fur; wing membranes transparent, terminal phalanx O'f second and third fingers fold back on digit; claw O'n thumb smaller than claws on toes; feet very large; metacarpals moderate, digits very long; O'uter toe shortest; tail, except extreme tip, inclosed in interfemoral membrane; ten vertebrre in tail; calcar well developed, much elongated, reaching about fO'ur-fifths distance to tail; no pO'stcalcaneal lobe. Color.-Brownish to' ashy gray above, dusky gray near skin; below distinctly lighter, the hair gray, tipped with whitish (in alcohol) . Skull and teeth.-Skull large, sagittal crest well defined on posterior portion of brain case; a very slight longitudinal depression on rostrum; upper narial notch extending backward a shO'rt distance farther than palatal notch and narrowed posteriorly; palatal notch rounded; palate with a m~dian posterior proj ection; bullre large, covering three-fourths or more of cochlere. Upper incisors small, subequal, separated from canines by a distinct diastema; the inner with two cusps, the anterior highest; inner with largest cone bifid, and a smaller accessory cone;

283

MYOTIS

upper canines especially elongate. without trace of secondary CUSp; cingulum low, rather indistinct; anterior premolar twice as large as middle premolar; both first and second out of line of tooth row, both visible from without, first not in contact with third; vertical extension of posterior upper premolar greater than molars, very much greater than two anterior premolars; lower incisors short, closely juxtaposed, forming a median angle; median pair wedge-shaped, widened, bifid; middle incisors trifid; posterior much the largest, molariform, with four cusps; lower anterior premolars nearly triple the bulk of middle; third about double that of first, all in tooth row. Measurements of Myotis macro tarsus (Wate?'house). E. H. Taylor coIlection. No. 339.

mm.

Head and body Head Tail Forearm Thumb and claw Second digit: Metacarpal First phalanx Third digit: Metacarpal First phalanx Second phalanx Fourth digit: Metacarpal First phalanx Second phalanx F'ifth digit: Metacarpal First phalanx Second phalanx Alar expanse Tibia Foot and claws Calcar Skull: Incisors to occiput Zygomatic width Palate from notch Width of brain case Height of brain case Upper tooth row Lower tooth row Mandible

55 21 45 46 9.3

42.5 5 43.5 18 21 42.5 12

11.5 39

10 10 312 19.5 15.5 18.6 18.5 11.4 7.5 9

8.7 8.2 8.2 12.8

284

PHILIPPINE LAND MAMMALS

Remarks.-The skull measurements are from a second specimen, collected on a small island off the coast of Tawitawi by Gregorio Lopez. It agrees very well with the Manila specimen. From the few specimens known i1his species appears to be well distributed through the Islands. The very remarkable feet and the size of the animal make it easily distinguishable from the smaller species here described. The wing membranes are unicolored, and thus it is easily separated from Myotis rufopictus. The type was collected in the Philippines but the type locality is unknown. ' Thomas * remarks on a specimen collected by Whitehead in Manila, as follows: This Bat was originally discovered by Cuming, and no other specimen has been received by the British Museum until now; I fail to see, either in the fresh specimen or the type, that the wing-membrane is attached t.o the body much nearer the spine than is usual, a character on which Dobson lays some stress. The black claws of the type, also specially mentioned by him, may have been caused by some .f luid in which the specimen had been put, for Mr. Whitehead's fresh specimen unquestionably identical specifically, has the claws of the normal pale coI~r.

Hollister t reports two specimens from Iligan, Mindanao (Grubbs) and one from Zamboa:nga (E. A. Mearns). MYOTIS JEANNEI sp. nov.

Type.-No. 340, E. H. Taylor collection; collected near Caldera, Zamboanga, April,. 1923, by E. H. Taylor. Alcoholic specimen with skull separate. Diagnosis; description of the type.-Adult female. Muzzle broad, "nose" moderately prominent, nostrils opening distinctly outward, a slight depression between nostrils or none at all; large gland on face borders nose and reaches lip; small gland above greatly reduced; a small triangular area naked on lower lip; rounded gland on chin much reduced; ear large, broad, anterior edge with a somewhat angular flap free at base; inner edge of ear curving to rounded summit; a prominent elongate concavity on outer border of ear below which the edge curves strongly to base where there is a thickened free flap; tragus rather long, inner edge straight to near tip, where it curves outward; outer edge attaches much below inner; at base of tragus a rounded flap above which is a notch that constricts tragus; outer line of notch straight to a point of greatest width, then gradually tapering to tip; a slight concavity below tip;

* Trans. Zool. Soc. London t Proc. U. S. Nat. Mus. 46

14 (1898) 385. (1903) 310.

285

MYOTIS

fur moderately dense, that on shoulders about 6 millimeters long; on sides and tip of muzzle scattered unequal hairs; outer half and inner third of ear naked; tragus naked except for a few hairs at inner point of attachment; wing membranes above and below, except near body, naked save for a meager scattering of fine short hairs, visible only with magnification; interfemoral membrane generally naked except for a few elongate hairs on membrane near tip of tail; about 2.5 millimeters of tail free from interfemoral membrane; wing membranes attach as near to heel as to base of toe; feet proportionally large, a few elongate (hairs at base of claws, with a few short hairs above digits; calcar well developed. Color.-Black-brown above, darker than mummy brown; fur at its base dull brownish black; on belly fur tips dirty buff, the lower two-thirds deep black-brown in very strong contrast; wing membranes same shade as body above; ears slightly more olivebrown; hairs at base of toes reddish orange-brown. Skull and teeth.-Skull rather flattened over brain case; rostrum with a broad shallow depression; narial notch rather rounded, narrowed behind; palatal notch distinctly rounded; palate with a posterior median extension; no median palatal ridge; bullre large, nearly covering cochlere; no sagittal crest. The inner incisor with a large anterior cusp and a smaller posterior one not bifid; outer incisor with two small cusps, the tooth with a broad shallow posterior groove; incisor separated from canine by small diastema; first premolar much larger than .second, the latter pushed out of tooth row but still separating anterior from posterior premolars. iVIeasw'ements of Myotis jeannei sp. nov. E. H. Taylor collection. No. 340.

female.

mm.

Total length Head and body Tail Forearm Ear Tragus Thumb and claw Second digit Third digit Third metacarpal Fourth digit Fourth metacarpal

77 45 32 36.5

15 5 8.5 36 66

34 50 32.5

286

PHILIPPINE LAND MAMMALS

Measurements of Myotis jeannei sp. nov.-Continued. E.H. Taylor collection, Na.34.0, felllal".

mm.

Fifth digit Fifth metacarpal Alar expanse Tibia Foot and claw Calcar Skull: Incisors to occiput Zygomatic width Palate from notch Width of brain case Depth of brain case Upper tooth row Lower tooth row Mandible

48 33

260 16.5

10 11 14.8

8.8 5.9 7.5 7 6.4

6.6 11

RemaTks.-The type specimen was collected along a small stream emptying into the sea near Caldera, Zamboanga, Mindanao, a few kilometers from the coast. The specimen was clinging to a large rock which overhung the side of the stream bed; it contains a single large embryo. The species belongs to the large-footed group of the genus represented in the Philippines by Myotis macro tarsus Waterhouse. It is named for my daughter Jeanne Louise Taylor (deceased). MYOTIS PATRICllE sp. nov.

Type.-No. 353, E. H. Taylor collection; collected in Agusan Province, Mindanao; collector and date unknown. Adult male. Alcoholic specimen with skull separate. Diagnosis,' description of the typ,e .-Very small; muzzle rather flattened; "nose" moderately projecting, with a median !\-shaped depression; nostrils directed downward, and not or only slightly outward; an enlarged irregular gland on each side of nose, reaching edge of lip and extending backward to near edge; a small triangular naked area on lower lip; a prominent rounded tubercular gland on chin; ears moderate; on inner anterior edge a free flap at base; outer anterior edge of ear curving rather irregularly to tip with a slig;h t concavity near top or none at all; tip narrowed, rounded; a distinct concavity on outer edge some distance below tip; lower half regularly rounded, ending in a small free flap which bends in and is slightly thickened;

MYOTIS

287

tragus with inner edge perfectly straight to inner point of attachment; outer edge attaches much lower; near point of attachment an angular proj ection, above which, is a notch, which ends near point of attachment of inner edge; above notch the edge forms a regular arc of a circle; muzzle with scattered hairs, only portion about nostrils actually naked; a rounded tubercle in front and slightly above eye; tragus with elongate hairs near base on anterior side, very short hairs on posterior side; ear except outer third with scattered hairs; body fur short, fur on shoulders 4 or 5 millimeters long; wing membranes above and below entirely naked, ,e xcept for an occasional very short hair; basal third of interfemoral membrane with long hairs; a few scattering hairs also on outer edge; lower surface with scattered short hairs; foot on outer half covered with long heavy hairs above; wing membrane attaches at base of outer toe; outer toe as long as inner; tail, except for extreme tip, involved in interfemoral membrane; two rounded glandular ar~as at edge of anus. Colo1..-Above, cinnamon to cinnamon-rufous, bases of hairs darker; bt:1ow, extreme tips of h~irs buff-cinnamon, the rest dark Mars brown; ears nearly the same color as upper fur; wing membranes darker brown; interfemoral membrane as light as ears; hair on feet reddish orange. Skull and teeth.-Skull sloping forward gently to rostrum; a longitudinal depression on rostrum; anterior narial notch rounded, either slightly pointed behind or not; palatal notch nearly angular; palate with a slight median ridge, nearly straight on posterior edge or with a slight median proj ection; bullre very large, all but hiding cochlere; sagittal crest wanting. Inner incisors with an anterior cusp pointing inward, and a bifid posterior cusp which is much lower; a fourth small cusp near base; outer incisor with two cusps, and a prominent groove running the length of tooth posteriorly; canines large, well developed, a slight posterior cingulum cusp; fkst premolar small, very much larger than second which is crowded partially out of tooth row but separates first and third premolars; incisors separated from canines by a distance nearly equal to diameter of one incisor; lower incisors slightly imbricate, first and second trifid, third larger, molariform, with four cusps; lower canine with an anterior and posterior cingulum cusp; first lower premolar very much larger than second, which is pushed back slightly out of line with other teeth but separating first from third premolar; the latter tooth shaped somewhat like canlne.

288

PHILIPPINE LAND MAMMALS

Measurements of Myotis pat'Y'icire sp. nov. E. H. Taylor collection. No. 353. male.

mm.

Total length Head and body Tail Forearm

Ear

12'

58.5 33 25.5

TTagus, lower outer edge Second fingeT, entire Third fingeT, entire Third metacarpal Fourth fing.er Fourth metacarpal Fifth finger Fifth metacarpal Alar expanse Tibia Foot and c1aw Calcar Skull: Incisors to occiput Zygomatic width Palate, from notch Width of brain case Depth of brain case Upper tooth row Lower tooth row Mandible

5.2 20

41 22 35 21 33

22.5 176 11.5

6 9

12.2

6.6 5 6.6 5.8 5.2

5.6 9.2

Remarks.-This is the smallest Philippine species, if not the smallest of the entire genus. The specimen appears to Le a young adult. The teeth are scarcely worn. The type is the only specimen I have seen. The species is named for my daughter Patricia Anne Taylor. MY OTIS BROWNI sp. nov.

Type.-No. 341, E. H. Taylor collection; collected near Saub, Cotabato, Mindanao, April 15, 1923, by E. H. Taylor. Alcoholic specimen with skull separate. Diagnosis; description of the type.-Small; apparently related to Myotis herrei but differing in numerous characters; "nose" somewhat more prominent, openings distinctly closer together and rounded, the depression between them less, and with a few minute hairs over surface of nose; glandular development weak, without any noticeable groove between glands and "nose;" no

MYOTIS

289

prominent rounded tubercle; longitudinal depressions nearly obsolete; tubercle in front of eye small or lacking; hair on lips and anterior rostrum scantier than in M. herrei; basal anterior lobe ,s imilar but anterior edge curves without emargination to summit; summit rounded; line of outer edge broadly emarginate to near middle where margin curves outward and again is emarginate above basal lobe which turns inward; tragus shorter, conca ve on its inner edge instead of straight; a similar basal lobe above which is a similar notch but smaller than in M. herrei; tragus, above notch, convex to a point more than two-thirds of distance to summit; tip less attenuated than in M. herrei; lower lip with a distinct small naked area; median tubercle on lower jaw small, rather indistinct; distribution of hair on membranes practically the same as in M. her<rei; calcar distinctly shorter, reaching scarcely past the halfway point between foot and tip of tail; wing membranes attach at base of outer toe. In other body characters the two forms are very similar. Color.-Color very much lighter than in M. herrei. Warm sepia above, slightly lighter on head and neck above where basal portion of fur is scarcely darker than outer portion; on back and rump basal portion of fur distinctly darker brown, about the color of membranes; ear membranes abou~ color of fur on top of head; tragus same color as ear on outer portion, lighter on lower half. Skull and teeth.-SimiIar to Myotis herrei but skull ,somewhat shorter, brain case slightly more expanded, interorbital distance greater, rostrum distinctly shorter, with a consequent shortening of tooth row; anterior palatal notch broader than deep; groove on rostrum shallower than in M. herrei. Upper incisors very similar in shape and position but actually smaller and separated from canine by a shorter distance; canine with a posterior sharp edge ending basally with a distinct cingulum cusp; first premolar nearly double the diameter of second; second premolar almost directly in tooth row, distinctly visible from outside; lower incisors distinctly smaller than in M. he1'rei but of the same general type; canine with a low posterior cingulum cusp; a cingulum cusp on posterior edge of first premolar, and a small anterior one on third lower premolar. Measwrements.-See under Myotis herrei. Remarks.-The single specimen was taken in a clump of bananas near the seacoast at Saub, Cotabato. The specimen is a female containing a young embryo. The species is named for 259453-19

290

PHILIPPINE LAND MAMMALS

my friend Dr. William H. Brown, director of the Bureau of Science, Manila. MY OTIS HERREI sp. nov.

Type.-No. 369, E. H. Taylor collection; collected January 22, 192.3, in cave at Montalban, Rizal Province, by W. Schultze, Albert W. Herre, and E. H. Taylor. Adult male. Alcoholic specimen with skull separate. Diagnusis; ,description of the type.-Small; snout moderately broadened; nostrils rather prominent, somewhat tubular, opening downward and slightly outward; a very slight triangular depression between nostrils; prominent glandular area on each side of "nose" separated from nostril by an ill-defined groove; a rounded glandular tubercle somewhat to side of nostril, behind which are two shallow longitudinal depressions; a naked area on lower lip, and a rounded glandular tubercle behind it; a small tubercle in front of eye; glandular areas rather sparsely haired; a few scattered vibrissa:! ; ear large, reaching to tip of snout when laid forward; inner edge with a large free flap at base; inner edge of ear rounding to near tip where there is a distinct emargination; tip rounded, somewhat truncate; from summit 'e ar slopes gradually to a prominent angular wrinkled notch, below which is a small rounded portion; below this is a slight emargination; at base a small free lobe turning inward; tragus with inner edge straight or very nearly so; outer edge with a rounded basal lobe above which is a rounded notch nearly as large as lobe; tragus widens above this point, the edge curving to a point nearly halfway to summit and then continues to tip in a straight line; tip rather acutely pointed; inner surface of ear with scattered elongate hairs to near tip; tragus with hair only at base; posterior side of ear naked for more than half iits length; fur on head and body above moderately dense, the hairs approximately 5 millimeters long; above wing membranes (except close to body), the antebrachial membranes and interfemoral membranes, except on basal third, naked; a few extremely minute hairs near distal end of tibia, and very heavy elongate hairs on feet; lower sides of membranes naked except that there are a few scattered hairs near distal portion of forearm, and the underside of interfemoral has scattered elongate hairs to near tip of tail; tail elongate, included in interfemoral membrane except for extreme tip; feet moderate, outer toe slightly the shorter; calcar strong, extending nearly two-thirds of distance to tip of tail; membrane with a small angle at tip

291

MYOTIS

of calcar; penis moderately large; a naked area about scrotal pouch. Color.-Blackish brown above, hairs nearly black at base, tips nearly bone brown; hair on face several shades lighter; below, Natal brown, bases of hairs nearly black; in region of penis and scrotal pouches the hairs nearly silvery gray; membranes generally same shades, perhaps darker than pelage; tragus and ears lighter than wings, tragus with a distinctly darker edge. Skull and teeth.-Brain case sloping gently forward to the nearly level extension of rostrum; a shallow longitudinal depression on rostrum; narial notch deeper than wide; anterior palatal notch wider than deep, with a slight median notch posteriorly; palate strongly concave, posterior part with a small rounded median projection; bullre large, nearly covering cochlere. Inner incisors short, anterior cusp pointed slightly forward and inwa.rd; posterior cusp rather flattened (perhaps bifid if unworn) ; outer incisor somewhat bifid, leaning forward, posterior surface curving in with a shallow longitudinal groove; canines slightly curved with posterior sharp edge; first premolar at least two and one-half times diameter of second premolar; latter pushed out of tooth row, only the tip visible from outside of tooth row; third premolar high; lower incisors increasing in size from first; first and second compressed, with three cusps; third rather molariform, with four cusps; canine with a slight anterointernal cingulum cusp; second premolar about half diameter of first, not or only slightly pushed out of line of tooth row; third lower premolar caninelike. Measurements of Myotis herrei and M . b'l'路o.wni; from the types.

Total length Head and body Tail Forearm Pollex Second finger, entire Third finger: Metacarpal First phalanx First and third phalanges Fourth finger: Metacarpal First phalanx Second phalanx

Myotis herrei,

Myotis brown.,

No. 369, male.

No. 341, female.

mm.

73 40 33 31.3 6.2 28 28 11

18 27 7 9

36 30 30 5.7 27 28 10 18 27.5 8

292

PHILIPPINE LAND MAMMALS

Measurements of Myotis herrei and M. browni; from the types-Continued. Myotis herrei. No. 369. male.

Fifth finger: Metacarpal First phalanx Second phalanx Ear: - From notch Greatest width flattened Foot with claws Calcar Tragus Tibia Skull: Total length Zygomatic width Width of brain case Interorbital width Depth of brain case Length of palate, median Upper tooth row Lower tooth row Mandible

Myotis browni, No. 341, female.

mm.

27 6 6

27 7 6

11.7 9 7 14 5.4 14

13 8.4 6.6 3 6 5.2 5.7 6

12 8 6.4 3.6 6 5.2 5.1 5.5

8.9 6.1 11 5 12

Remarks.-The species appear.s to be related to Myotis browni h erein described. The measurements are somewhat similar but the character of the tragi, the upper incisors, the second premolars, and the attachment of the alar membrane suffice to distinguish them. The upper tooth row of M. browni is distinctly shorter, and the rostrum of the skull narrower than in the species here described; from M. jeannei it is distinguished by the much ,s maller feet; from M. patricim, by the longer forearm and digits and the different character of the incisors. The single specimen of this form was collected far in the depth of the upper Montalban Cave, clinging to the low roof. The species is named fO'r my friend Dr. Albert W. Herre, ichthyologist, formerly chief of the division of fisheries, Bureau of Science, Manila, who assisted in collecting the type specimen. Genus GLISCHROPUS Dobson Glischro~~s

DOBSON, Proc. Zoot Soc. London (1875) 472; MlLLIilR, Bull. U. S. Nat. Mus. 57 (1907) 205. Vespe?'U[}o DOBSON, Cat. Chirop. Brit. Mus. (1878) 183.

The genus Glischropus was erected by Dobson as a subgenus of Vesperugo Dobson to contain forms having the upper incisO"r

GLISCHROPUS

293

forced out of the tooth row, and callosities on the sole and the ball of thumb much developed. Miller recognizes it as of generic value. Like Pipistrellus except that the upper incisor is forced outward from the tooth row and so turned that its convexity is directed outward instead of backward; ears triangular, the outer margin terminating in a distinct well-defined lobe midway between base of tragus and angle of mouth; lobule at base of outer margin of tragus small or absent; postcalcaneal lobe distinct; wings from base of toes. 2- 2 1- 1 2- 2 3-3 Dental formula: i. 3- 3' c. 1- 1' pm. 2- 2' m. 3- 3 = 34. The genus is known from Palawan, Borneo, and Batchian. Three species are recognized, of which G. tylopus Dobson has been found in the Philippines. The peculiar structure of the sole of the foot and of the inferior surface of the thumb is known only in one other genus, Tylonycteris. It is probable that the pads serve to enable the bats to cling to the undersurface of large leaves or fruits. The pads are less developed than in Thyropte1'a, a genus which has strong sucking disks. GLiSCHROPUS TYLOPUS (Dobson)

Vespe1'ugo (Glischropus) tylOp~LS DOBSON, Proc. Zoo!. Soc. London (1875) 473 (Borneo); M(mog. Asiat. Chirop. (1876) 114; ELERA, Cat. Sisto Fauna Filipinas 1 (1895) 12 (Palawan). VeS'[.ie1'ugo tylopus DOBSON, Cat. Chirop. Brit.. Mus. (1878) 236. Glisc!t1'OPZLS tylopus MILLER, Bull. U. S. Nat. Mus. 57 (1907) 205; HOLLISTER, Proc. U. S. Nat. Mus. 46 (1913) 310 (Palawan).

Description of the species.-Small; muzzle broad, evenly rounded in front; nostrils opening on a level with glandular prominences on each side, and without intervening emargination ; glands of upper lip greatly developed, forming smooth, almost naked, prominences, causing a furrow along center of face behind nostrils; ears triangular, narrower above, with rounded tips, ascending part of inner margin faintly convex, nearly straight, succeeded by a considerable convexity, causing upper third to appear concave, then distinctly angularly emarginate opposite base of tragus, and terminating in a well-defined lobe midway between base of tragus and angle of mouth, but on a lower level than mouth; tragus faintly concave along inner margin; ':outer margin slightly convex, tip obtusely rounded off; a minute, very acutely pointed projection slightly above base of outer margin; thumb long, the whole lower surface of basal half oc-

294

PHILIPPINE LAND MAMMALS

cupied by a naked rounded pale yellow callosity with transverse wrinkles; sole of foot similarly formed, but surface flat or slightly concave; light yellow lof callosities contrasts remarkably with other body color; postcalcaneal lobe distinct; extreme tip of tail projecting. Skull and teeth.-Skull much as in Pipistrelhu;, inner upper incisor long, bifid, the smaller cusp placed posteriorly and externally .;near the extremity; outer incisors very short, but in transverse diameter equal to inner ones, placed in a plane slightly anterior, the single cusp sloping inward and lying against cingulum of inner incisors; lower incisors trifid, not crowded; first upper premolar internal but visible from without. Color.-Generally dark; adhesive pads yellow. Measurements of the type of Glischropus tyloptUs (Dobson). mm.

, .-.

40 38 13.2 13 5 30 6 61 40 13.2 6

Head and body Tail Head Ear Tragus Foreann First finger Third finger Fifth finger Tibia Foot and claws

Remarks.-The type locality is "north Borneo" and the type is in the British Museum. A single specimen in the United States National Museum was collected by J. B. Steere in Palawan. Hollister states that the specimen from Palawan is smaller than any in a series taken in Borneo. Genus PIPISTRELLUS Kaup

Pipistrellus KAUP, Skizzirte Entwick.-Gesch. u. N aturl. Syst. d. Europ. Thierw. (1829) 98; BONAPARTE, Iconog. della Fauna Ital. 1 (fasc. 20) (1837); DOBSON, Proc. Asiat. Soc. Bengal (1871) 213; MILLER, Ann. & Mag. Nat. Hist. VI 20 (1897) 384; North Am. Fauna (1897) 87; MILLER, Bull. U. S. Nat. Mus. 57 (1907) 204. Romicia GRAY, Mag. Zool. & Bot. 2 (1838) 495. Romicius BLYTH, Cuvier's Anim. King. (1840) 75. Vesperugo KEYSERLING and BLASIUS, Wiegmann's Arch. fur Natlirg. 1 (1835) 312 (part.); DOBSON, Cat. Chirop. Brit. Mus. (1878) 183 (part.); H. ALLEN, Monog. Bats N. Am. (1893-1894') 121. Hypsugo KOLIDNATI, Algem. deutsch. Naturh. Zeit. Dresden neue folge 2 (1856) 131.

295

PIPISTRELLUS

Nannugo KOLENATI, Allgem. deutsch. Naturh. Zeit. Dresden neue folge 2 (1856) 13l. Scotophillus ALLEN, Monogr. Bats N. Am. (1893-1894) 27 (part.).

Small bats, averaging from 35 to 45 millimeters in length of head and body; ear usually shorter and broader, tragus less 118

120

122

1i'6

I 4

Batan Is

PHIUPPINE ISLANDS

® 5cotophilus temminckii ® Tylonyctel'is pachypus © Ptel'ygistes stenoptel'U3 © Pipistl'ellu8 imbl'icatus ® Pipistl"el\us abroamus ® Pi pistroellus il'l"etitus

@ Pi pistl'ellus tenuis

I.j:

QCagayan Sulu

6·

0' 122

FIG. 14.

124'

126

Distribution of ScotophU:U8, Tylowycteris, Pterygistes, and Pipistrellus in the Philippines.

acutely pointed than in Myotis; teeth strictly normal for the family and not differing in any essential feature from Myoti.<; except in the absence of one upper and one lower premolar; outer upper incisor not as large as inner but extending con-

296

PHILIPPINE LAND MAMMALS

siderably beyond its cingulum; inner upper incisor ~imple or more often with a well-developed secondary cusp; canines usually rather stout, posterior cutting edge of upper tooth often, though not invariably, with an incipient secondary cusp; anterior upper premolar barely or not in tooth row; other teeth with no special peculiarities; skull essentially as in Myotis though with a tendency to a greater breadth; external characters not essentially different from those of Myotis, except as mentioned. The members of the genus are recognizable by their dental formula, large outer upper incisor, unmodified skull and ears, and the normally long fifth finger; the species are all small, some of them being the smallest known bats. (After Miller.) Four species have been reported from the Philippines. Key to the Philippine species of Pipistrellus Kaup.

a 1. Tragus not expanded above, reaching its greatest width about middle of inner margin, which is concave; generally crescent shape; upper outer incisors half length of inner and less than half their transverse diameter; small postcalcaneal lobe; forearm, 33 millimeters. P. irretitus (Cantor) (p. 296). a". Tragus widest slightly above base of its inner margin; postcalcaneal lobe well developed., b t. Outer upper incisor not equaling the outer cusp of inner incisor in height; outer margin of ear conch straight or faintly concave above; forearm, 28 millimeters __ ___ ___ ... _.___ P. temLis (Temminck) (p. 297). b 2. Outer upper incisor exceeding outer cusp of inner incisor in height. c '. Outer margin of ear conch straight; forearm, 33 millimeters. P. abramus (Temminck) (p. 298). c". Similar to abramus, but tragus narrower above, slightly smaller, and lobule near its base on outer margin less distinct; head broader, thumb and feet larger; post calcaneal lobe rather indistinct ______ .__ . ________________ .____ .__ __ P. imbricatus (Horsfield) (p. 300). PIPISTRELLUS IRRETITUS

(Cantor)

Vespertilio i1'r'etitus CANTOR, Ann. & Mag. Nat. Hist. I 9 (1842) 481.

Vespertilio meyeni WATERHOUSE, Proc. Zoo!. Soc. London (1845) 3 and 7.

Pipistrellus irretitus HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 17; Proc. U. S. Nat. Mus. 46 (1913) 335.

Description of the species.-Forehead moderately arched; muzzle broad, sparingly covered with hairs, swollen at sides; two longitudinal grooves on muzzle, which are shallow at first, become deeper behind where they curve outward towards eyes; small fleshy tubercle over eye; area between grooves almost naked, but a few minute scattered hairs at tip in front and

297

PIPISTRELLUS

along edge of upper lip; lower lip with small triangular naked patch at tip; nostrils rather widely separated, opening obliquely outwards; ears short, subtriangular, shorter than head, extreme tip rounded, posterior border slightly emarginated; on inner side three or four transverse ridges; tragus narrow, somewhat pointed, with a notch in the middle of external margin; wings rather narrow, membrane extending to base of toes; thumb small; legs short; metatarsus short, about equaling length of toes, which are very nearly equal, outer slightly shorter than others; interfemoral membrane not ample, with scattered hairs below, especially near tail; above, this membrane appears to be naked, except at base; calcar moderate, with a small postcalcaneal lobe. about 4.2 millimeters long and 2.1 wide; extreme tip of tail free. Color.-Fur on body brownish gray, lighter nearest skin; body below ashy gray; muzzle blackish; wing membranes dark brown; limbs dusky, purplish red; interfemoral membrane grown above, pale beneath. MeaSUTementS of PilJistrellus in'etitus (Canto?"). [From the type of P. meyeni Waterhouse.] tnm.

Head and body Tail Forearm Ear Spread of wings

40 23.2 27.5 6.6 165.1

Remarks.-The type was collected in Chusan, China. The first Philippine specimens were collected by H. Cuming. These were made the type of Vespertilio meyeni. Cantor states that these small bats are frequently arrested in the strong webs of two large spiders, Epeira bilineata and E. heraldica, and this has given rise to the erroneous belief that spiders feed on bats. PIPISTRELLUS TENUIS (Temminck)

Vespertilio tenuis TEMMINCK, MOllogr. Mamm. 2 (1835-41) 229, pI. 57, figs. 5-7; HOFFMAN, Abhand und Berichte des KonigI. Zoolog. Mus. Dresden 3 (1886-7) 1-29; HOLLISTER, Philip. Journ. Sci. ยง D 7 (1912) 17. (PipistTellus.) Vesperugo tenuis WAGNER, SuppI. Schreb. Saugeth. V (1855) 740; DOBSON, Monog. Asiat. Chirop. (1876) 98; Cat. Chirop. Brit. Mus. (1878) 226.

Description of the species.-Ears smaller than those of Pipist'i"ellus pipistrellu8; tragus rather broad and short, its greatest