: me = bo rere aot Os o or 1 Ley, etetones fe" "O80 ehy, oe + “10 olen. te. ttete me. fa teen ape he ek Ao lrerneig haa. novener anes s = ome ": 8 we acess Pe mck ete eoaenen a Gieres cor eS Aes on A . wehcae ieee BIOLOGY LIBRARY 791 BURRILL HALL FEB 07 i996 ratormation tar Coniributars to Fieldiana Cue cab | Frtlaon is thine: a journal fot Miele (Myseum start menmbers and: esearch ‘associates. although Mans iots Tay Mins iated authors tay he Conse pel as space permits. ar the Gouttal, cursed a papceirbe Of $65.00 pan pinnwed, page on Cec Hor, thereat, Payment of ut teast 50% of page ¥ dharges Qualsihe A Paver TORR be Chee rote souna Vee rer Moes 4He publication dime.’ QComtinbutions Hon staff, research " usediates ana iwiled anlhor wih be eensidévedtor poblicn lon ai gandiéss of ability Lopay page charges, however, the ful 4, Chaise i hancdtin Jor nonufiilated anchors of vnsclicted manuscripts, “Farce complete oHpies Of (he text Cinctuding title £ faze and abstract) and of tne Ley ehe arcing should. be, subniitted (ond original copy. plus: to review copics which may be: mache comes. No manysehipts sill be considered. ror Gublication on submitted to reviewers’ Delore all materials are’ complete and jn ake hands oh tie Maen Rgitog, | Mandsctipty sci be sytmulted te Scienmtinic Tiditor Preldiana. Piela Miseun of Naural History, Chicago, Iitinois. ” S605 -PAIG: SA’, itry Pout: Marustripus: basi be typewritten double-spaced on staridard-weight, BiA- by Tisinch paper with wide margins ‘ On ‘all fade sides. LE tiped On aa TENGcompatiblé computer “using MS-DOS, sats submit) text of S¥e-inch disketic (WordPerfect 4a, 42, v0 5:0) Mile Mate Disphaywrite 2, 9 Sed) Waity PC, Santina Microsolt Word) Volksariter, | WorniStar programs or ASCILY. ; é Pox payers over 200 orahuseripl pages apthous are requested: to submita "Table of Cobtents,”a,"List of Llustrations, and a “List of Fables" inmediatélysfollowing tiie’ page. lo most cards, the text shuui¢ be’ preceded oy an ‘Abstract and” should-condlude. witiy “Acknowledgments” {if any) and “Literature Cited? aA any : Ail measneements should DEAN Eke Tettiosyeletn (perinds are mot used after abbreviated measurements). The format - and style of headings should follow thut of/recent issues of Meidiana: 1, Sete For more iddtaiied styie. information, ste The Chicago Mamialof Sole (13th ed), published by, The University off) Chicago’ Préss, and dino recent issucs ob Pieler. a oa References: ii "Literature Cited,” bookand journal titles should be given in ‘full: Where abbreviations are desirable (¢.g;, in citation OF synonyniies), authors consisently should tollow Botanico-Perioditum-Huntiantun and FL-2 Taxonomic- Diterature BY FA. States & R. §. Cowan (1976 et seq.) (botanical papers) of Sertal Sources for me Hiosis Data Base (1983) published by the BioSciences Information’ Sexice. Names of botanical authors shotitd follow the "Draft Index of Author Abbreviations, Royal Botanic Gardens; Kew,” 1984 edition, or (2-2. gt, Reéfertnces should be typed inthe following form: a Croat, TB) 1978; Fiora Of Barro Colorado Island, Stanford University Press; Stanford, Calif.) 943 pp. nea a Grubb, Fo, YR. Liovd, and T. D. Pennington, 1963, A comparison of mantane add lowland rain forest in Beuador,” L Phe forest structure, physidgnomy, and floristics. Journal of Ecology, St: 67-601. DME Bid: Langdon, Bw. M1979) Yage among the Siona: Cultural patterns in’ visions, pp. 63-80; Ja Browman, D. Land ROAL Schwarz, eds.) Spitits; Shamans, and Stats. Mouton Publishers, The Hague, Netherlands. Paes i: Murra, 1. 1946, "The historic tribes of Hedador, pp, 785-821. In Steward, J. FL, ed., Handbook of South Amexican: Indians, Volk. 2. fhe Andean Givilizations. Bolletin’ 143, Bureau of American Ethnology, Smithsonian (astitution, Washington, D.C. : Mie tae Siolve, R. G. T9S1) Ferns and fern allies of Guatemala, Part Tl, Polypodiaceae. Fieldiana: Botany, ms., 6 1-522. > s Ilustratioas: INustrations aré referred to as "figures" in the: text (nat as "plates"). Figures must be accompanied By. gome indication of scale, normally a reference bat. Statements in figure captions alone, such as "x 0.8," are not acceptable. Captions should be typed doublé-spaced and consecutively. See recent issues of Fte/diana for details of style. © aN All illustrations should be marked on the reverse with author's name, figure number(s), and "top." aes Figures as submitted. should, whenever practicable, be 84% 11 inches (22% 28 cm), and may pot exceed 11% x 16M ineves (30x 42 cm). Whistrations should be mounted: on boards in the arrangement to be obtained in the printed’ work, This® onginal set’ should be suitable for ttaiismission io the printer as,follows: Pen atid ink drawings maybe. originals (preferred), or phatustats;’shaded drawings amat be originals, but within the size limitation; and photostats must be high-quality, glossy, blackdabd-white prides. Oviginal iflustrarions willbe returned to the corresponding author upon publication unicss otherwise’ upecified. Re ete re Authors who Wish: to publish fpures ‘that require costly special paper Or eelot, reproduction ‘must make prior, atrangemants with, the, Scientific Bditon, eae D : ; i ve Page Proots: ivldiancl exmploys a two-step correction system. ‘The cortesponding alithior will normally receive a copy, Of the edited manserint on which deletions) additions, and changes can be made and quences answered. Only oneiset of page probs will be sen AU desinsd ‘cortections of type dust be made on the siygle: ser OF page roots. Changes it page! prods fas wpposed 16 corsections) are mary expensive. Anthor-generated changes trpage proots can only be made if the author apiees ta advance toopay for term. it pst ee REC FT FANS PUBLICATION IS PRINTED ON AGID-PREE PAPER | KIELLDIA: Zoology NEW SERIES, NO. 69 A Key to the Bats of the Philippine Islands Nina R. Ingle Department of Natural Resources Fernow Hall Cornell University Ithaca, New York 14853 Lawrence R. Heaney Department of Zoology Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 Accepted March 20, 1992 Published October 30, 1992 Publication 1440 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1992 Field Museum of Natural History Library; of Congress Catalog Card Number: 92-73951 ~ Pa ISSN 0015-0754 PRINTED IN THE UN ITED STATES OF AMERICA t Table of Contents POMRTRACT? oh 8c SO i a Pes rs | ENTRODUCTION) oc eo Cee a here l METHOUG 566. oon ee eee eA 3 ANATOMY AND MEASUREMENTS OF A BAT .... 3 IGE OF THE REY ite a ee ee 4 Determination of Age and Sex ........... 5 Identification to Family and Species ...... 6 Use of Measurement Tables .............. 6 Comparison with Skull Drawings ......... 6 COLLECTING AND PRESERVING VOUCHER SPEC- IMENS 9 5.S5 SO a en Eo 5 ctor We | 2 KEY TO FAMILIES OF PHILIPPINE BATS ........ 7 FAMILY PTEROPODIDAE: FRUIT BATS ......... 8 Kev to: Pleropognine: 2001s eo ses 10 FAMILY EMBALLONURIDAE: SHEATH-TAILED BATS ae chisy o cout kta cere carers 14 Key to Emballonuridae .........25524.4.3. 14 FAMILY MEGADERMATIDAE: FALSE VAMPIRE BATS 2c 2G in ee eh ee es 14 FAMILY RHINOLOPHIDAE: HORSESHOE AND LEAF INOGED BATS. Goce ces eke 15 Key to'Rhinolophidae: 6 oo. coco ee 15 FAMILY VESPERTILIONIDAE: EVENING BATS 18 Key: to Vespertilionidae 6.50645 Dee ys 20 FAMILY MOLOSSIDAE: FREE-TAILED BATS .... 22 ROY 10 MOIOSSIURE: 1G eee eee 23 ett Re Et ME AO ABR Ke ae Sore Os eee 23 PEA NOWLEDOMENTS: 2555 2 oo 8k tat 25 ESTRRATURE CITED: 2.535200 os oe ee ae 25 List of Illustrations 1. Map of the Philippines ................ 2 2. Generalized pteropodid bat ............ a 3. Ears of representative bats ............. 4 4. Bat with external measurements indicat- Dee ore Gen etek Oo ee Murs Pa ey 4 5. Bat skull with teeth and cranial mea- Beremients Indicated: 8 oe i es 5 6. Joints in digits of wing of adult and ju- WORMS OBIE me ic ei an ice ee as Sn 5 7. Tails of representatives of Philippine Ny SHINING 5 iso 25 or a os ei ewes 8 8. Heads of representatives of Philippine RE TMMNOENNOR 5 ch, on cis oc fads ne eens els 9 9. Frontal views of fruit bat skulls, show- ing incisors and canines ............... 12 10. Dorsal views of fruit bat muzzles ...... 13 11. Posterior attachment of wing membrane RENE MIE phe reine se eee ee eines 14 ill . Skull of Chaerephon plicata . Noseleaves of Rhinolophus and Hipposi- 7 Ft \ ORE LCS BD: te ORES Fa cla A ee 16 . Connecting processes of some rhinolo- ONIGS hs Sate res eee enn 18 . Ventral views of muzzles of Murina and DIVOUS. o Focin oe aor aon ee 21 . External ears (pinnae) of some vespertil- HOMOGE. So ke Siok ace eee a 21 . Hind feet of Myotis spp., showing at- tachment of wing membrane .......... Ze . Pad on wrist of Glischropus ........... 22 (EMIS Ol- 2 VIONICLONIS. one ooh 22 . Skull of Acerodon jubatus ............. 28 Skull of Cynopterus brachyotis ......... 29 . Skull of Dobsonia chapmani .......... 29 . Skull of Eonycteris robusta ............ 29 . Skull of Eonycteris spelaea ............ 30 . Skull of Haplonycteris fischeri ......... 30 . Skull of Harpyionycteris whiteheadi .... 30 . Skull of Macroglossus minimus ........ eH 2 Skull of Nyctimene rabori <5 2 a 31 . Skull of Ptenochirus jagori ............ 31 . Skull of Pteropus hypomelanus ........ 32 . Skull of Pteropus pumilus ............. a2 . Skull of Pteropus vampyrus ........... 33 . Skull of Rousettus amplexicaudatus .... 33 . Skull of Emballonura alecto ........... 33 Skull of Saccolaimus saccolaimus ...... 34 . Skull of Taphozous melanopogon ...... 34 . Skull of Megaderma spasma .......... 5 hs . Skull of Hipposideros ater ............. 35 . Skull of Hipposideros diadema ........ 36 . Skull of Hipposideros obscurus ......... 36 . Skull of Hipposideros pygmaeus ....... 37 . Skull of Rhinolophus arcuatus-l ........ 37 . Skull of Rhinolophus arcuatus-s_ ....... 38 . Skull of Rhinolophus inops ............ 38 . Skull of Rhinolophus macrotis ......... 39 . Skull of Rhinolophus philippinensis 39 . Skull of Rhinolophus virgo ............ 40 . Skull of Kerivoula hardwickii .......... 40 . Skull of Miniopterus australis ......... 40 . Skull of Miniopterus schreibersi ........ 41 Skull of Miniopterus tristis ............ 41 ew OF Mfarind CPClONIS. 4.0% 5% sake 41 . Skull of Myotis horsfieldii ............. 42 . Skull of Myotis macrotarsus ........... 42 . Skull of Myotis muricola .............. 42 . Skull of Myotis rufopictus ............. 43 . Skull of Philetor brachypterus ......... 43 . Skull of Pipistrellus javanicus .......... 43 . Skull of Pipistrellus tenuis ............. 44 . Skull of Scotophilus kuhlii ............ 44 Gas ete 44 ~ 3 . List of Tables 1. Measurements of Philippine Pteropodi- IRE ie oct re eke ated nde ee tall viele ee Rice 11 2. Measurements of Philippine Emballonur- idae and Megadermatidae .............. 15 3. Measurements of Philippine Rhinolophi- if Gm i Saree fae ere orate Aa erie tae ad 17. 4. Measurements of Philippine Vespertilion- LAG axes cts ok Mh oats Whe Soi ie Oh Oe 19 5. Measurements of Philippine Molossidae.. 24 iv A Key to the Bats of the Philippine Islands Nina R. Ingle and Lawrence R. Heaney Abstract An identification guide is presented for the six families and 70 species of bats now known from the Philippine Islands, based on a key and a set of standardized measurements. Most critical characters are illustrated, and detailed drawings are provided of the skulls of 42 species. Introduction The Philippine Islands (fig. 1) support a large and diverse fauna of mammals: over 170 species are now known, compared, for example, to the 105 species known from Madagascar, which has nearly twice the land area (Heaney et al., 1987; Jenkins, 1987). About 100 species are endemic to the Philippines, giving the country an unusually high number and percentage of unique species (Hauge et al., 1986; Heaney, 1986, 1991; Heaney et al., 1987; Koopman, 1989). One of the most diverse and, in general, poorly known mammalian orders in the Philippines is the Chiroptera. Sixty-eight species of bats were known from the Philippines when the last checklist was prepared (Heaney et al., 1987), and two more are now known (Pteropus dasymallus and Harpi- ocephalus harpia). In number of species, bats ex- ceed even rodents, of which 67 are now known (Heaney et al., 1987; Musser and Heaney, 1992). By our best estimate, 22 species of bats, about 31%, are endemic to the Philippines, again an un- usually high number (Heaney, 1991; Koopman, 1989). The high levels of species richness and ende- mism are factors of special importance currently because of the rapid rate of loss of natural habitat in the Philippines. Roughly 94% of the Philippine land area was once covered by forest; that figure had been reduced to 40% at the end of World War II, and current estimates of forest cover range from 25% to less than 20%, depending in part on the amount of degraded forest that is included (Col- lins, 1990; Hauge et al., 1986; Myers, 1988; Ut- zurrum, 1991). The ongoing forest destruction poses an especially grave problem because many species of bats, especially endemics, depend pri- marily on forest (Heaney et al., 1987; Heaney and Utzurrum, 1991; Heideman and Heaney, 1989). Two species of bats (Acerodon lucifer and Dob- sonia chapmani) are believed to have become ex- tinct in the last 100 years, and many others are threatened (Heaney and Heideman, 1987; Heaney et al., 1987). No identification guide to the 70 species of bats recorded from the Philippines currently exists. The most recent work describing Philippine bats, Tay- lor’s “Philippine Land Mammals”, was published in 1934 and does not include the many species subsequently described or recorded from the Phil- ippines, nor does it reflect the many changes in taxonomy that have taken place over nearly 60 years. Moreover, Taylor’s keys were intended for use with museum study specimens, not live ani- mals in the field. Thus, identification of Philippine bats in recent years has been primarily by com- parison with museum specimens. This requires access to a comprehensive reference collection and is not an option for researchers in many places. This key is intended to permit identification of all bat species that have been recorded from the Philippines, to the extent that current knowledge permits. It is our hope that this key will encourage more research on the Philippine bat fauna, which, with its diversity and distribution over many is- lands with differing habitats and climates, serves as an excellent subject for studies of biogeography and many aspects of ecology. We also hope that a greater knowledge about Philippine bats will con- tribute to efforts toward their conservation. FIELDIANA: ZOOLOGY, N.S., NO. 69, OCTOBER 30, 1992, PP. 1-44 | peaks 145 10 118° GREATER PALAWAN — GREATER NEGROS- PANAY Pa GREATER SULU 122° l x GREATER MINDANAO 1 LF 126° Less than 120 meters current water depth 0 200 | ae L 1 a 18 Kilometers GREATER LUZON 14° Maripipi Biliran Leyte Dinagat 104 64 126 ° l l FIELDIANA: ZOOLOGY Forearm \S 2 . ant - 1st digit: thumb Metacarpals 2nd digit 1st phalanges Terminal phalanges 3rd digit 4th digit Fic. 2. Generalized pteropodid bat, with important structures indicated. ) interfemoral \N XZ membrane Tail Wing membrane Hind foot Methods The descriptions and measurements in this key have been based primarily on examination of specimens in the American Museum of Natural History (AMNH), British Museum (Natural His- tory) (BMNH), Delaware Museum of Natural His- tory (DMNH), Field Museum of Natural History (FMNH), Royal Ontario Museum (ROM), Philippine National Museum (PNM), Silliman University Mu- seum of Natural History (su), University of Mich- igan, Museum of Zoology (UMMz), and United States National Museum of Natural History (USNM). Our descriptions and measurements are based on Philippine specimens except in a few cases involving species that are rare in collections; in those cases, specimens from elsewhere or char- acters from published accounts were utilized. For standard external measurements we relied preferentially on data from animals that we had collected, but we also utilized data on specimen labels and measured specimens in collections. We took cranial measurements with dial or digital cal- ipers following deBlase and Martin (1974) and Heaney and Peterson (1984); these measurements are described briefly in the following section, Anatomy and Measurements of a Bat. We consulted a variety of publications as aids in building the keys and descriptive sections, re- lying heavily on Harrison (1966), Hill (1983), Lawrence (1939), Lekagul and McNeely (1977), Medway (1969), Miller (1907), Payne et al. (1985), and Taylor (1934). These publications, the refer- ences cited in Hill (1983), and publications re- ferred to in the sections for each bat family should be consulted for descriptions of characters not treated in the key. Anatomy and Measurements of a Bat Familiarity with basic terminology on the anat- omy of a bat is necessary to use this key; Figure 2 shows the most important external structures. The external ear (pinna) has two structures that can be useful for identification. The tragus is a oe Fic. 1. islands is indicated by the shaded areas. Map of the Philippines. Recent islands are outlined by continuous lines, and the extent of late Pleistocene INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 3 Fic. 3. Left external ears (pinnae) of representative bats. A, Pteropodidae; B, Rhinolophidae; C, Vespertil- ionidae. projection from the base of the ear (fig. 3C); this structure is absent in members of the Pteropodidae (fig. 3A) and Rhinolophidae (fig. 3B) and is quite small in the Molossidae. The antitragus is the flap along the lower posterior margin of the ear (figs. 3B, 12); it is especially well developed in rhinol- ophids and most molossids. Rhinolophids and megadermatids have noseleaves, structures formed from elaborate folds of skin on their noses (figs. 8B,C, 12); bats of other families have no such structure. All bats have four types of teeth: incisors, ca- nines, premolars, and molars (fig. 5). All bats pos- sess a pair of large, sharp, conical teeth called ca- nines on the upper and lower jaws. The smaller teeth at the front of the mouth, anterior to the Total length canines, are incisors. Posterior to the canines are the premolars and molars, collectively called cheek teeth. The large teeth in this series are referred to as molariform teeth. Molariform teeth are usually low and broad with cutting or grinding surfaces; the anterior cheek teeth are tiny pegs in some spe- cies. External and cranial measurements provide substantial aid in identification; we have therefore included tables containing typical adult measure- ments. The definitions of standard external mea- surements are illustrated in Figure 4. The cranial measurements included in the tables are shown in Figure 5 and are defined as follows: condylobasal length (CBL), distance from the posterior edge of the occipital condyles to the anterior tip of the premaxillaries; condylocanine length (CCL), dis- tance from the posterior edge of the occipital con- dyles to the anterior edge of the base (alveolus) of the canines; and maxillary toothrow (C' to last M), distance from the posterior edge of the last upper molar to the anterior edge of the upper canine, taken along the bone line of the alveolus. Use of the Key Using the key most effectively requires several steps that are explained in detail in the following sections. First, the bat should be identified as to — LL Body length f0OO COC 7 att —as — pert tere peeemt Poses seman rece sence cline: foment mere Fic. 4. Bat with external measurements indicated. FIELDIANA: ZOOLOGY Incisors and Molars Fic. 5. age and sex. It should then be identified to family with the Key to Families, and then to species with the key to that family. Next, all external mea- surements should be taken and matched against those in the tables of measurements; if the mea- surements do not match, the identification is ques- tionable and the process should be begun again from the first step. Finally, if a skull is available, it should be visually compared with Figures 19- 60 to see if it matches the features of that species or genus. Determination of Age and Sex The age of bats can be estimated by the degree of ossification of the joints in the digits of the wing (Anthony, 1988). In juvenile bats, the joints have cartilaginous discs where growth takes place. When a light is shown through the wing of a live bat, the bands of cartilage at the joints appear partly trans- lucent (fig. 6B). In adult bats, the bones are fully ossified and the joints appear opaque (fig. 6A). The shape of the joints of the digits also differs between juvenile and adult bats; this is especially useful in determining the age of animals prepared as stuffed skins or fluid-preserved specimens. In juvenile bats — CC Ondylobasal length__, 1 Condylocanine length__} Bat skull with teeth and cranial measurements indicated. the joints are swollen and tapered (fig. 6B), whereas in adult bats the joints are knobby and more dis- tinct from the bone shaft (fig. 6A). Bats can be sexed by examination of the external genitalia. Males have a conspicuous penis (except A B Fic. 6. Joints in digits of wing. Stippled areas rep- resent bone and open areas represent cartilage. A, adult; B, juvenile. INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 5 subadult male Rousettus, in which the penis is retracted into the abdomen). Both sexes possess axillary nipples on the upper chest, usually near the armpit (axilla). However, the nipples of adult females are more prominent than those of males. In female rhinolophids and megadermatids, a pair of inguinal papillae that look like nipples is present just anterior to the genitals. Identification to Family and Species Following determination of age and sex, the bat should be identified to family with the Key to Families, and then to species with the key for the appropriate family. The keys consist of sets of al- ternatives arranged in order of increasing restric- tiveness. Alternatives belonging to the same set are preceded by the same number with different superscripts (e.g., 1, 1’, 1”). For each set, the user should choose the best alternative; this indicates either the next set of alternatives to be considered, or the name of the family or species. All distin- guishing characters should be checked carefully before a choice is made. This will minimize the likelihood of incorrectly identifying a bat. Re- searchers should bear in mind that this key in- cludes only those species now known from the Philippines, and that discovery of undescribed species and previously unreported species is likely. Figures illustrating important descriptive char- acteristics are referred to in the key and should be consulted. Fur coloration can be useful in identi- fication; it should be observed on a dry animal or study skin. Use of Measurement Tables Once a bat has been identified to species, it is good practice to check if its measurements agree with those given in the tables. Cranial measure- ments should be taken with calipers. External measurements can be taken with a ruler, although calipers provide more accurate measurements of forearm length and of small structures such as parts of the noseleaves of rhinolophids. Because forearm length is a very useful mea- surement for identifying adult bats, ranges for fore- arm length are given in the keys for each species, usually at the end of the final description for that species. Note that forearm lengths are given even when overlap in the ranges of two or more species means that the measurement cannot be used to fully distinguish between the species. When other measurements are particularly useful for separat- ing similar species, they are also given in the key. Measurements should not be used to identify ju- venile bats, although hind foot length is sometimes useful because the feet of juvenile bats approach adult size long before most other structures have ended growth. Comparison with Skull Drawings Skulls are very important in identification of bats. Thus, although we have written this key with the intention that live animals and preserved spec- imens can be identified without examination of their skulls, we have included skull drawings of common Philippine bats (figs. 19-60). The draw- ings will aid in identification of museum speci- mens with cleaned skulls and will provide a means of verifying an identification based on external characteristics. The skull drawings also serve as a guide to head shape. Even if use of the key yields an apparently unambiguous identification, it is al- ways best ifa skull can be examined from a vouch- er specimen. The skull should be compared with the figures, and cranial measurements should be taken and compared with those in the tables. Den- tition, although most easily visible on a skull, can also be observed, at least partially, on a live animal and should be examined when possible. The mouth can be gently pried and held open by a toothpick, and the lips can be lifted to view the cheek teeth. With small bats a magnifying lens is a great help. Note that if the jaws are closed, some teeth, par- ticularly the canines, may obstruct the view of others. Users of this key should not overlook the fact that variation in both quantitative and qualitative characters is present in all species. Whenever pos- sible, samples of more than one individual for each species should be examined and the most common character states used in the key. The identities of a few species of Philippine bats that are extremely poorly represented in collec- tions (sometimes only by the holotype) are uncer- tain. When possible, these species have been in- cluded in the key on the basis of whatever descriptions are available, and further comments on their identification are provided in the Notes section. We also comment briefly on two new re- cords of bats for the Philippines in this section. FIELDIANA: ZOOLOGY Collecting and Preserving Voucher Specimens In any study, several individuals of each species should be preserved as voucher specimens and deposited in a museum collection. Voucher spec- imens are necessary to verify identification, since many species can be identified definitively only after careful study of the external and cranial mor- phology. Such specimens are invaluable for adding to our current, very limited knowledge of Philip- pine bats, particularly of distribution and inter- and intraspecific variation. Specimens should be collected and prepared carefully, following all reg- ulations for the scientific study of wildlife. Bats are most easily preserved in fluid. They should be killed in a quick and painless manner, examined for reproductive condition, and weighed, and their standard external measurements taken (total length, tail length, ear length, and hind foot length). The date, habitat, collection locality, el- evation, collector, collector’s field number, and Key to Families of Philippine Bats external measurements should be recorded in In- dia ink (which is best) or pencil (which is usually acceptable), not with other kinds of ink (which are not permanent). These data should accompany the specimen as part of the permanent record. The specimen should have a permanent label attached that bears the collector’s name and field number; other data may go onto the same label or into a field catalog. The specimens should then be rinsed lightly with soapy water, injected with a 10% so- lution of formalin, and immersed in a 10% solu- tion of formalin. After about three days, they can be rinsed with clean water and transferred to 70% alcohol (methanol is best) for permanent storage. Skulls can be extracted and cleaned in the muse- um. The Philippine National Museum, University of the Philippines at Los Banos, Silliman Univer- sity, and a number of other institutions maintain research collections and accept voucher specimens for their permanent collections, and the staff may be able to help with identifications. 1. Interfemoral membrane is absent or reduced, forming narrow margin along insides of legs; tail short (up to 20% of body length) or absent, never completely enclosed by interfemoral membrane (fig. 7A); both thumb and second finger with claw (except Eonycteris and Dobsonia; fig. 2); both tragus and antitragus absent (figs. 3A, 8A); ear margin forms continuous ring ... Megachiroptera: Pteropodidae . Interfemoral membrane is a continuous expanse of skin stretching between legs (figs. 7B—F; except Coelops, which has a concave posterior margin to the membrane and no tail); tail present (except Megaderma and Coelops, which both possess noseleaves, structures not present in any pteropodids), usually comprising more than 20% of body length; the second finger does not have a claw; either tragus (fig. 3C) or antitragus (fig. 3B) or both present; ear margin does not form continuous ring .. (Microchiroptera) 2 + ROOSIR A584 070 OKe) se) 60 86.56, ah be BAe. 618 6° 616) O 6%, 0.016 [656 OF OG ele! 8 Abra! fey. 0166 S50 08s, Mo wre oe! ose We) (ete 16) @: w, “BTeL Us fay 6 lee ss INGMOIERT PROMENE COGS, SIs. hit so coh are sen OU et oh cl eRe et Lan ean een ek a ae 3 ee PACERS MOAT CHER Ee ES)) iz 0. duh, sone cheeks ete alee ees ee oe PN a a 4 3. Large ears connected at top of forehead (fig. 8B); tragus long and forked; external tail absent but interfemoral membrane well developed (fig. 7B) ..............0.0e cee c eee ceece Megadermatidae 3’. Ears not connected across top of forehead; tragus absent, but antitragus usually well developed (fig. 3B); tail present, enclosed by interfemoral membrane except at extreme tip (fig. 7D; except Coelops, which has no tail and a reduced interfemoral membrane ....................0-5 Rhinolophidae 4. Tail extends to posterior margin of interfemoral membrane (extreme tip may project 1-2 mm beyond membrane; fig. 7C); ears variable, usually not fleshy (fig. 8F); anterior edge of bony palate deeply Ly Ls genes aig ase aoad A Rly SGA OE CARE RUS ORTOP RAE BREE Ae It AA COR ORI gon ROC ae Vespertilionidae 4’. Tail emerges dorsally from interfemoral membrane but is shorter than membrane when legs and membrane are outstretched (fig. 7F); ears not noticeably thick and fleshy (fig. 8D); anterior edge of DOMy Tmlate GOCOLy CISreI Ale oe isk. ea aan ogg as aio wih oie bl Se Emballonuridae 4”. Tail projects beyond posterior margin of interfemoral membrane for over half its length (fig. 7E); ears thick and fleshy (fig. 8E); anterior edge of bony palate continuous, not emarginate 0.3; 8 1 8' O06 oe o40 he RxD 63S 1G Oi 0, B00 :0 8) C'S 6) 612 OO BO. 000 66) 6. 6S OL 0 6 a8 0. 6) 014) 06) * ©0018, O18 6 0.6.60. 2 0.0 6 UF © 0.4 4 6' 0 0.0)8 08 0 6% 6 6 6 INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 7 AY a od yes ar . Fic. 7. Tails of representatives of Philippine bat families. A, Pteropodidae (Rousettus); B, Megadermatidae (Megaderma); C, Vespertilionidae (Miniopterus); D, Rhinolophidae (Rhinolophus); E, Molossidae (Chaerephon); F, Emballonuridae (Taphozous). Family Pteropodidae: than those of most other bats; the large eyes are Fruit Bats associated with their dependence on vision for ori- entation and their lack of echolocation (sonar) sys- All fruit bats are characterized by dog-like heads tems (except for the very simple and limited one (fig. 8A), with eyes that are proportionately larger used by Rousettus). They do not have the elaborate — Fic. 8. Heads of representatives of Philippine bat families (not to same scale). A. Pteropodidae (Rousettus). Note that the ear margin is continuous, neither tragus nor noseleaf is present, the eyes are large, and the face is dog-like. B. Megadermatidae (Megaderma). Note that a noseleaf is present, the tragus is forked, and the large ears are connected across the top of the forehead. C. Rhinolophidae (Rhinolophus). Note the elaborate noseleaf, the well-developed 8 FIELDIANA: ZOOLOGY ) ao antitragus, and the absence of a tragus. D. Emballonuridae (Taphozous). Note that there is no noseleaf and that a tragus is present. E. Molossidae (Chaerephon). Note that there is no noseleaf and that the ears are thick and fleshy. F. Vespertilionidae (Miniopterus). Note that there is no noseleaf and that a tragus is present. INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 9 folds of skin in the nasal region (also associated with the use of echolocation) that are present in two other families of bats in the Philippines (Megadermatidae and Rhinolophidae). The exter- nal portion of the ear is simple and of moderate size, and the margin of the ear forms a continuous ring. Fruit bats do not have a tragus or antitragus (figs. 3A, 8A). All but three species (Dobsonia chapmani, Eonycteris robusta, and E. spelaea) have a claw on the second digit (fig. 2), along the leading edge of the wing; bats belonging to other families lack this claw. Many species have no tail, and in none does the tail comprise more than 20% of the body length. Their size varies greatly, with forearm lengths ranging from 40 to 215 mm, and weight from about 16 g (Alionycteris paucidentata) to at least 1,200 g (Acerodon jubatus; table 1). Most species show substantial sexual dimor- phism; males are larger than females and often have broader, more heavily muscled heads. Males of many species possess a ruff of fur (often colored yellow or rusty red) around the neck and on the shoulders that is lacking or poorly developed in females. Subadults of both sexes resemble adult females. Parous adult females of all species have a pair of axillary nipples that are large and prom- inent; males and subadult females have very small but visible nipples. The skulls of all pteropodids are characterized by the presence of prominent postorbital process- Key to Pteropodidae es, and all but a few species have postorbital fo- ramina (figs. 19-32). The dental formula is vari- able. Most species have broad, blunt, rounded molariform teeth, but the nectarivorous species have greatly reduced premolars and molars. All species have long, prominent canines. In many species, males have longer canines than do females and often have more strongly developed cranial crests. As the English name for the family implies, most fruit bats feed on fruit, but three species (Eonyc- teris robusta, E. spelaea, and Macroglossus mini- mus) feed primarily on nectar and pollen. Fruit bats often are abundant, being uncommon only in upper montane and mossy forest (Heaney and Rickart, 1990; Heaney et al., 1989). In forested areas, small to medium-sized species roost in hol- low trees and in foliage, either alone or in groups; the large flying foxes roost in exposed treetops. Several species roost in caves, often forming col- onies in the hundreds or thousands. Andersen (1912) provided descriptions of most species of pteropodids found in the Philippines. Additional descriptions may be found in Berg- mans (1975, 1978), Francis (1989), Heaney and Peterson (1984), Klingener and Creighton (1984), Kock (1969a,b,c), Musser et al. (1982), Peterson and Fenton (1970), Rookmaaker and Bergmans (1981), and Yoshiyuki (1979). + Claw On. thunib: Dut nor On SECONGANOIE Avcce7, lee nara epee erg amaied stele ts pete eri ea re ae 2 i Claws: on: both thunmib and second Gisit. (igi 2): ce seins coals. cea Mee ee eared 4 2. Wings attach to body along midline of back; two upper incisors and two minute lower incisors; forearm: 123-13 F mim; skull a5 in Figure 2 ee. i 2 a A ek Dobsonia chapmani 2’. Wings attach along sides of body; four upper and four lower incisors, all very small; forearm 67— PAE 61 5) OARS ete a AI RPOe EEIL MOR ATO SERCO VOL ariel TSI PRE RLE NY PR COA nts W RCP Eo ROE (Eonycteris) 3 3. Pair of prominent 2-6-mm-long kidney-shaped glands lateral to anus; tail 12-20 mm; forearm 67-— SOiromi: skull-as it Pisute 2934505 esse Pain yc ole ee Uae Oe eS Eonycteris spelaea 3’. No glands near anus; tail 20-28 mm; forearm 67-82 mm; skull as in Figure 22 ............... Bes eee hteetihe Gam sacs ago Stig eaters eenipast eh oe tin SAO tadantes at hier esh pS AUL ami eet Eonycteris robusta 4. Forearm 94-215 mm; tail absent; four upper and four lower incisors ....................0-. 5 4’. Forearm 41-92 mm; tail present or absent; number of incisors varies between species ....... 13 5. Wings with prominent pale blotches, particularly along thumb and anterior edge of wing adjacent to first and second digit, especially at wing tips; forearm 135-141 mm..... Pteropus leucopterus 5’. Wings dark brown without prominent pale blotches; forearm 94-215 mm .................. 6 6. -Pelage on dorsal surface of lower. back pale brown: Or gray = 664 cee ooo es BS rr 6’. Pelage on dorsal surface of lower back dark brown or black, sometimes with yellow flecks .... 8 7. Forearm 132-165 mm; condylobasal length 65-70 mm; three cusps on second and third upper molariform teeth, including a well-developed anterolingual cusp ............. Acerodon leucotis 7’. Forearm 94-113 mm; condylobasal length 46-52 mm; two cusps on second and third upper mo- lariform teeth (no anterolingual cusps); skull as in Figure 30 ................. Pteropus pumilus FIELDIANA: ZOOLOGY 901-69 76-08 p7-81 v7-€1 pS I-8ZI py vI-VZ1 8°8£-O're snjppnva1ixajduy snyjasnoy OI8-SZZL vO7-6LI Se-SE 0 OZE-OLZ ¢ 1€-0'°97 0°6L-8°89 snsdduiva sndosad = €€I-8I1 O€-vZ 0 ¢17-061 6°77-€'07 C19-6'€S snsoizads sndosaig 007-S¢I €1lI-+6 87-07 0 O8I-SSI ¢ 61-91 ¢1S-0'9r snjiuind sndosaig Ore IpI-SE€l 87-97 0 OvZ-S07 077-91 0°€9-¢'°9¢ snaajdoonay sndosaig OSt-SZP 6rI-9E1 ZE-87Z 0 ObZ-O1Z 9'b7—b IZ $°$9-0'9¢ snupjauoddy sndosaig 06b-08¢ (€) ZSI-€ET (€) $Z (1) 0 O€7-07Z 9°b7-€°7Z 7 ¥9-9'LS snjyjoudsop sndosaid TO-IE 81-09 titi ZI STI-86 O'1I-S$°6 VIEL AOUIUL SNATYIOUAI L6-79 06-91 S7-81 81-9 SrI-07I CCI-O'1l YE Say Yo uosvl snayIoUuald LI-SI OS-€b pI-ZI 0 9L-O0L 0°8-0'L 0°€7-8'07 snpouospjij4v2 sndo1ajdoiQ bL-19 6L-IL v7-LI O¢-07 6vI-I€I VZI-ZT1 €SEP'EE MOQDA JUIUATIIAN O7-LI 7S-Sb ¢I-ZI 9-0 9L-0L LL-UL 6°77-L IZ lasoutjam Sdosavsay 07-S I St-Ip SI-ZI 0 18-9 9°6-9'L ¢°97-L'bT SNUIUIU SNSSOjSOAID 771-66 16-08 $7-61 0 9SI-O€1 99I-F'SI €€b-L'6E Ippayanym siajaduoidiv yy 17-91 €S-9r 9I-11 0 08-s9 T6-9'L b'S7-0°7Z May Isyf Sluajaduojdv H] 06-8r 08-19 €7-81 07-7 SpI-IZI LE1-8:01 9°SE-E1E vapjads sisajaAuoq 08-SL 78-L9 €7-61 87-07 SSI-LZI OrI-9' II 6 LE-6' EE DISNGOA S14dJIAUOT = 76 (1) 7 (1) 67 (1) IZ (1) €°SI-v'vl (Z) 7 T7P-€E Ip (Z) snaoipods sniajdoovdq = I€I-€ZI LE-Se 97-€Z bezee lc 0° 17-061 S'€S-S'8b jupudoy2 piuosqoqd 1-97 89-8¢ 07-41 cI-€ OLI-88 0'OI-€°8 ¢°67-0'97Z snodyoviq snsajdoudD 8I-F1 Os-sb (S) ZI-II (S$) 0 (¢) OL-+9 (¢) ti-39 077-0 IZ pipjuapionod sisajatuoypy = OLI-L91 = 0 = 0'1€-0°67Z SEL-O'IL 4afin] uoposaoy = S9I-ZEI 7£-67 0 0S7-07Z 0°87-S$ "SZ 0°0L-0°S9 S1j0Ma] UoposaIV 0S7Z‘ 1-006 ¢17-S9I It-l€ 0 OIE-SSZ 0°9€-0°6Z C.cS-e7/ snjoqn{ uoposaop IIIA W18910 ivy LAB wsuay] [e)0L W 3S®[-.9 149 sapedg 11 ‘suoumloods ouiddiytyg WoOdj Udy¥e) 19M ‘1X9] UI PouYyop se ‘s}UIWOINsRI ‘sosoyiuosed Ul UdAIS JIB SazIs ajduIes UdYM 1d9dx9 ‘s[eNpIAIpUI Q] ISBI] Iv JUISIIdo1 sosULY ‘oLpIPOdoJolg UIddI[IYg 1[Npe JO sosueI JUSWIOINSeaW | «ATAV INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS A bo) Od Ok — eas eee Fic. 9. Frontal views of fruit bat skulls, showing incisors and canines (not to same scale). A, Otopteropus carti- lagonodus (Alionycteris paucidentata and Haplonycteris fischeri have an equal number of incisors); B, Cynopterus brachyotis; C, Nyctimene rabori; D, Ptenochirus minor (Ptenochirus jagori incisors are very similar). 8’. 10. 10’. iH, 11”, | i2.. 13: 13’. 14. 14’. £5; 1% 16. 12 Forearm 165-215 mm; condylobasal length 68-86 mm; fur on upper back sometimes completely BON eg SoS cates cate atna ada sa es NTE oe RS RSE Ne Fe ERR OS REE CN RE REE EO: 9 Forearm 118-152 mm; condylobasal length 54-66 mm; fur on upper back never completely black, BSA y BONEN Sc. ie salah aa Stee Loaele Gig erate Soden ae ANS Reet La EO Ie i Dorsal pelage may be completely dark brown or black; if upper back is golden, the posterior margin of the golden area forms a sharply defined transverse line with the dark brown lower back; tips of ears nearly pointed; two cusps on second and third upper molariform teeth (no anterolingual cusps); condylobasal length 68—79 mm; forearm 179-204 mm; skull as in Figure 31 ....Pteropus vampyrus . Dorsal pelage never completely dark brown or black; a golden patch on top of head, extending anterior to line between ears, is always present; the golden dorsal pelage never forms a transverse line along the edge of the dark brown of the lower back; tips of ears bluntly rounded; three cusps on second and third upper molariform teeth, including a well-developed anterolingual cusp; forearm hGd=2 Fs Ini; Condylopasal length 7 I—oo MI 246 4 eee oie er eA ret Wee ho aie cdi 10 Forearm 167-170 mm; condylobasal length 71-74 mm (occurs only on Panay Island; see Notes) yen et CORT ad noni aeons PERRIN tee re tM enn Ie Adie ee aire eC et oer er Acerodon lucifer Forearm 165-215 mm; condylobasal length 72-86 mm; skull as in Figure 19 ... Acerodon jubatus Forearm 118-133 mm; condylobasal length 53-62 mm (occurs only from Zamboanga to Sulu) .. IR Feige ee ira os LR SaaS Ro Os er MA Pe: Waele a A OOS AU eleva eee Res Vv gm Lyrae US Reade ANN Pteropus speciosus Forearm: 133-152 mimzcondylobasal lensth 36-60 oo soe rice cee a aaa 12 Lower legs heavily furred nearly to ankle; forearm 133-152 mm (see Notes) .................- Sper nes Encik Netnacatats aN os eA SPR rene Boil CRUST ITER Sie Mie aR RT ae ME BE RY Pteropus dasymallus Lower legs nearly naked; forearm 136-149 mm; skull as in Figure 29 ... Pteropus hypomelanus SPRUE CLEISOUAG: 3 cnt ee eecee Os eA tu Rete iy Sk Sek Se BAA Soe oN tend nie petty noe 14 ‘Lail-present (ig: 7 Ac it may be smiall.so- look carerully): .4255:5.0cs f6k oe eee ta hanes 19 Two small lumps of soft white tissue on each ear, at anterior and posterior margins; forearm 43- 108 7710 0 Rear Re OP SURUER EC Ne Sin DRO TNR OAL WA Eee maT ren NE hn One Pere Pt een Otopteropus cartilagonodus No lumps of soft white tissue at margins of ears; forearm 41-91 mm ...................45- 15 Forearm 80-91 mm; well-developed secondary cusps on canines (one on upper canines, two on lower canines); six cusps on last two upper and lower molariform teeth; dorsal surface of hind feet thickly firved: skull-ae in Pisure 25 32. cc Sos wees ae ete Harpyionycteris whiteheadi Forearm 41-53 mm; canines do not have well-developed secondary cusps; two or three cusps on molariform teeth; dorsal surface of hind feet may or may not be thickly furred ............. 16 Muzzle long and slender (fig. 10A); nostrils are not tubular and do not project beyond rest of muzzle; wing membrane attaches on top of foot, above gap between third and fourth toes from outside (fig. FIELDIANA: ZOOLOGY 7 Fic. 10. Dorsal views of fruit bat muzzles (not to same scale). A, Macroglossus; B, Haplonycteris; C, Nyctimene; D, Rousettus, E, Cynopterus. 11A); teeth, except canines, greatly reduced; forearm 41-45 mm; skull as in Figure 26 ......... AERA ty Ae ERO OF Wy BAIRE EB aN RON We ese CPD PULU US PEE anak MOP Be OSPR Boner ON Macroglossus minimus 16’. Muzzle short and broad; nostrils tubular (as in fig. 10B); wing membrane attaches on side of foot (on or above outermost toe; fig. 11C); teeth robust; forearm 43-53 mm .................... Ly 17. Four upper and two lower incisors (fig. 9D); forearm 45-52 mm .......... Megaerops wetmorei 17’. Two upper and two lower incisors (as in fig. 9A); forearm 43-53 mm ..................2-. 18 18. Band of pale fur along dorsal surface of forearm; interfemoral membrane present; forearm 46-53 MERON RU RO IR PTO Pe i ca ees cee oars week edie Whey ae We AeA ato Haplonycteris fischeri 18’. No band of pale fur along dorsal surface of forearm; no interfemoral membrane; forearm 45-50 BENIN Rs a ie ies rats Ne fa eee SE nae NE LANE Cain Wa MG att eR eke Alionycteris paucidentata 19. Ears and skin on dorsal surface of bones of wings with prominent pale yellow spots; dark stripe along most of dorsal midline; two upper and no lower incisors (fig. 9C); nostrils elongated into tubes about 2-3 mm long (fig. 10C); forearm 71-79 mm; skull as in Figure 27 ...... Nyctimene rabori 19’. Ears and wings without yellow spots; no stripe along dorsal midline; four upper incisors and at least two lower incisors; nostrils not elongated into tubes (although when viewed from above they may appear slightly tubular); forearm 45-92 mi) oA iis eis Naina BT oe Sabie 20 20. Muzzle moderately long and tapered (fig. 10D); anterior surface of upper canines with vertical groove (not always prominent); wing membrane terminates above gap between outermost toe and second toe from outside (fig. 11B); forearm 80-92 mm; skull as in Figure 32 .................. MENS ER VISR atk Pisce ake Css Fea See ie RESALE Wd RORY CR TS aA Rousettus amplexicaudatus 20’. Muzzle short and broad (fig. 10E); anterior surface of upper canines smooth, not grooved; wing membrane terminates either on outermost toe (fig. 11C) or above gap between outermost toe and second toe from outside (fig. 11B); forearm 45-92 mm ............ 0... cece eee eee eens zt 21. Four upper and two lower incisors (outer pair of upper incisors much smaller than inner pair; fig. DE BARS arb rcs RR ae TRS NS as FS Oe PEO eae AES CEU AWS Mee Tie 22 ay. Four upper and four lower incisors (fig.:9B). 2.6 on VR IR bb cee tas 23 #2. Forearm 76-90 mm; skull as in‘ Figure 28... Sn Seti eitee de eves Ptenochirus jagori INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 13 22’. Forearm 60-78 mm (Greater Mindanao only) 22. POrearm 45-92: aon ose kee Snes He aS eRe SUNN PEt on ay eta aie Ptenochirus minor TERE RTE ee er Megaerops wetmorei 23. Anterior edges of ears pale; wing membrane terminates on side of foot, on outermost toe (fig. 11C); cheek teeth moderate in size; forearm 58-68 mm; skull as in Figure 20; (see Notes) ........:... Nigh Gh lyin ee Ce OER eee Cynopterus brachyotis 23’. No pale rims to ears; wing membrane terminates above gap between first and second toe from outside (fig. 11B); cheek teeth greatly enlarged, squarish in outline with large cusps and ridges; forearm: about 92-mms.. 3. eee eee ee Family Emballonuridae: Sheath-Tailed Bats Sheath-tailed bats are small to moderate-sized bats (forearm 44-71 mm; table 2). They are the only bats in the Philippines in which the tail per- forates the dorsal surface of the interfemoral mem- brane (fig. 7F), so that the basal portion (usually about half) of the tail is enclosed within the mem- brane and the distal portion is free, lying on the dorsal surface of the membrane. The eyes are small to moderate in size, and the ears are simple and of moderate size (fig. 8D). A tragus is always pres- ent. The muzzle is rather pointed, with the nostrils at the tip; it does not have a noseleaf. Key to Emballonuridae Ree CaN AA The ORE eT Dyacopterus spadiceus The skulls of emballonurids have postorbital processes that are very well developed (figs. 33- 35); these processes are absent in vespertilionids, which are superficially the most similar to em- ballonurids. Emballonurid premaxillaries are small and delicate; they do not fuse to each other at the anterior midline and are attached to the maxil- laries by flexible connective tissue. The anterior edge of the palate has a deep emargination. All of the sheath-tailed bats in the Philippines feed on insects. Common roosting sites are caves, hollow trees, and attics of buildings; they usually aggregate in moderate to large groups (ten to sev- eral hundred). 1. Two distinct pairs of upper incisors; forearm 44-49 mm; skull as in Figure 33 ................ BSS nO ISe OUTS) 6. 8 I. 6.49578 "9 AH) LOO ONG 16, Ow wy Ole) Ow ewes a 1h 16) 40m. METS DMS LE hoe Pda ae Per Emballonura alecto I. SONS pair OF Upper incisors. 1orearm Gl—7 1 Min 274 G i654 eis ole he ASR Le a 2 Forearm 61-65 mm; dorsal pelage pale brown to sandy at tips, very pale at bases; skull as in Figure BO Ns Pires er Ree MN Ce POLI SSD Mecca Nite Sern SR Lar Glia Taphozous melanopogon Forearm 66-71 mm; dorsal pelage very dark brown, usually with flecks of white; dark hairs do not have pale bases; skull as in Figure 34 ....... Family Megadermatidae: False Vampire Bats This is the least diverse of bat families in the Philippines, with only a single species present, Megaderma spasma (fig. 8B). M. spasma is a me- dium-sized bat (forearm 57—63 mm; table 2) easily recognized by its large ears that meet across the top of the forehead, erect, simple noseleaf, and lack of tail (although the interfemoral membrane is well developed). The skull of Megaderma spasma is of moderate size and its stoutly constructed (fig. 36). The pre- maxillaries are absent, so there are no upper in- 14 Ric ah ohn on, or eSet ated or Mn a: Saccolaimus saccolaimus cisors, and the anterior edge of the palate has a deep emargination. The postorbital processes are very poorly developed. The upper canines have a large secondary cusp. A C Fic. 11. Posterior attachment of wing membrane on left foot. A, Macroglossus; B, Rousettus; C, Ptenochirus/ Haplonycteris. FIELDIANA: ZOOLOGY TABLE 2. Measurement ranges of at least 10 individuals of adult Philippine Emballonuridae and Megadermatidae. Measurements, as defined in text, were taken from Philippine specimens. C'-last Total Hind Fore- Species CBL CCL Length Tail Foot Ear arm Weight Emballonura alecto 13.3-14.2 12.5-13.4 5.0-5.8 56-69 9-12 7-10 12-16 4449 46.5 Saccolaimus saccolaimus 20.9-23.3 — 8.8-9.9 103-117 20-28 15-19 16-20 66-71 28-36 Taphozous melanopogon 19.6-20.6 _ 8.1-8.8 100-111 20-25 11-15 20-24 61-65 20-29 Megaderma spasma _ 21.2-23.6 8.7-9.4 70-89 0 18-22 36-43 57-63 21-27 Megaderma spasma feeds primarily on large in- sects (especially cicadas and katydids) and occa- sionally on small vertebrates such as lizards, frogs, and small birds (D. Balete, pers. comm.). It has been found roosting in caves and in hollow trees, singly and in small groups. Because only a single species is present, no key to the family is necessary. Family Rhinolophidae: Horseshoe and Leaf-Nosed Bats The family Rhinolophidae is composed of about ten genera and over 100 species; it is represented in the Philippines by three genera (Coelops, Hip- posideros, and Rhinolophus) and at least 17 species (see Notes). All rhinolophids have a noseleaf, a structure consisting of elaborate folds of skin in the nasal region. The shape of the noseleaf varies among species and is an important character for identification (fig. 12). In members of the genus Rhinolophus, the posterior noseleaf is long and pointed (fig. 12A) and the anterior noseleaf is horseshoe-shaped. Between the anterior and pos- terior noseleaves lies the sella, an anterior-facing structure that is connected to the posterior noseleaf by the connecting process. In some species, sup- plementary leaflets are present lateral and ventral to the posterior noseleaf (fig. 12A). In Coelops and Hipposideros, the posterior noseleaf is low and rounded (fig. 12B) and may be divided into pock- ets by vertical septa. The intermediate noseleaf is a cushion-like structure. Supplementary leaflets are sometimes present lateral and ventral to the an- Key to Rhinolophidae terior noseleaf (not shown in fig. 12B, but see fig. 12A). Between the nostrils is the internarial sep- tum. Rhinolophids lack a tragus, but many spe- cies, especially those belonging to the genus Rhi- nolophus, possess a well-developed antitragus, a flap of skin on the lower posterior margin of the ear (fig. 12). Most rhinolophids are small (forearm 34-60 mm), but a few are medium-sized (forearm up to 90 mm; table 3). The skulls of most rhinolophids are delicate and slender, although some, especially those of the larger Hipposideros, are fairly stout. Postorbital processes are absent. Premaxillary bones are pres- ent but are small and connected to the skull by cartilaginous articulations, and so are movable in live or freshly killed animals. Incisors are present but are moderate to very small. The lower incisors are trilobed, although this is often inconspicuous. Many species have a strongly expanded nasal re- gion (which supports the noseleaf structures), giv- ing the skull a strongly sinuous dorsal profile in lateral view (figs. 37-46). Rhinolophids are insectivorous. Some species roost in caves, in numbers ranging from a few to several hundred. Other species roost in hollow trees, hollow fallen logs, and other sites in forest. Although rhinolophids usually comprise a small proportion of captures in mist nets, at times a single species may be the most common bat spe- cies netted at a given site. For further descriptions of members of the ge- nus Hipposideros, see Hill (1963a) and Jenkins and Hill (1981). For Rhinolophus, see Andersen (1905a,b,c,d). Posterior noseleaf pointed (fig. 12A); six pairs of lower cheek teeth ............ (Rhinolophus) 2 1’. Posterior noseleaf low and flattened (fig. 12B); five pairs of lower cheek teeth . Forearm 68-73 mm 2’. Forearm 38-57 mm Oere bE 20.6 Ob Bis -0°5-4 6 846.010 8) & 0 O16 ye OP Oye 6B. SC CR KOH & O98 00 3. Dorsal tip of connecting process sharply pointed (fig. 13A); forearm 47-49 mm ............... SEN Ue A TSR va ee BCR Oe bo Ce Se MO ant ae San So Se et AY ee ee Se SER Pre ae ee rey, Rhinolophus acuminatus INGLE & HEANEY: BATS OF THE PHILIPPINE ISLANDS 15 16 . Dorsal tip of connecting process not pointed (figs. 13B,C); forearm 38-57 mm .............. Posterior noseleaf Connecting process Sella Anterior noseleaf Supplementary leaflet (| \ Vertical septa Posterior noseleaf Intermediate noseleaf Internarial septum Anterior. noseleaf Fic. 12. Noseleaves of Rhinolophus and Hipposideros. A, Rhinolophus; B, Hipposideros. oS Antitragus nearly triangular (as in fig. 12A); cup at base of sella (immediately above nostrils) 8-10 mm wide, almost as wide as anterior noseleaf; forearm 51-57 mm; skull as in Figure 45 ....... Sale sk nie AGE Be ABLES te abd “hi Lae at gee ade Ss ce eta aa et ONO a rE Ter nee IDS Rhinolophus philippinensis . Antitragus most nearly rectangular; cup at base of sella 2-4 mm wide, no more than half as wide as anterior noseleal forearm as5G FT 60, sieht ais ie ei na se Fly ie fe Sea re 5 Connecting process:.attaches below tip of sella:(fig.. ISB) ..4.5..5.2.0..05 000s c00 000 Seedeiewsvess 6 > Sconnecting process attaches at tip of sélla (ig: T3C): nore ae ae tas cre anon Se IIe onlin Diesen Hipposideros pygmaeus 11’. Both pairs of lateral leaflets do not meet under anterior noseleaf; forearm 44-50 mm .......... Rees RREL Pe Dycrest ous omen Stare ry Te RU ae Percaee Mie" sic asked Mackie eee ieeds feat e neers eRe er ay ae neeeaans SEN Hipposideros cervinus 12. No vertical septa in posterior noseleaf and tail >30 mm; forearm about 47 mm (see Notes) .... eA ore eo hey Ee ee TRL BORON oe POA OR Mae RPt re 8 ORR ps eke Ae TAA at rr ORS Hipposideros coronatus 12’. Vertical septa may or may not be present in posterior noseleaf; if they are absent, then tail 18-24 MIN: AOLearin 3648 MIN: ow eee HAL IEE De Doon oe a oe ee ee eae ned ee 13 13. Anterior noseleaf 5.5-7.0 mm wide, intermediate noseleaf 5.0-6.0 mm wide, posterior noseleaf 6.0-8.0 mm wide; hind foot length 10-12 mm; forearm 42-48 mm; skull as in Figure 39 ....... SNe A Fe Oe MUN csr MC rt eerie MRA der Se Re Samat Mae nE Tene cal LON Deck Hipposideros obscurus 13’. Anterior noseleaf 4.0-5.0 mm wide, intermediate noseleaf 3.5—4.0 mm wide, posterior noseleaf 4.5-5.5 mm wide; hind foot length 7-10 mm; forearm 38-43 mm .....................0-. 14 14. Internarial septum not swollen at base; forearm about 42 mm ............ Hipposideros bicolor 14’. Internarial septum swollen at base; forearm 38-43 mm; skull as in Figure 37 ... Hipposideros ater Family Vespertilionidae: Evening Bats This is an exceptionally large and diverse family, treme tip; fig. 7C), a simple face without a noseleaf (among species in the Philippines; fig. 8F), and a well-developed tragus. Most vespertilionids are small to medium-sized (forearm 22—55 mm; table with about 40 genera and 275 species worldwide and about 11 genera and 22 species in the Phil- ippines. Vespertilionids are identified as having a long tail that is completely enclosed by the inter- femoral membrane (except occasionally the ex- Fic. 13. Lateral views of connecting processes of some rhinolophids. A, R. acuminatus; B, R. macrotis (that of R. virgo is very similar); C, R. subrufus/R. arcuatus. 18 4) and have small eyes. The skulls of vespertilionids are generally small, with a great variety of shapes and degrees of ro- bustness (figs. 47-59). The braincase is high and domed in some genera (e.g., Miniopterus, Myotis; figs. 48-50, 52-55) and exceptionally low and flat- tened in others (e.g., Tylonycteris). They all lack postorbital processes. All species have a deep emargination at the anterior end of the palate, so that the incisors on either premaxillary are widely separated. The molariform teeth are generally typ- ical of those of insectivorous bats, with broad crowns and sharp, shearing crests. All of the vespertilionids in the Philippines are insectivorous. Roosting sites include caves, build- ings, foliage, hollow trees, unfurled banana leaves, FIELDIANA: ZOOLOGY “eAReY £ Trea + “OoUIOg , = L7v7 (€) 6-8 (Z) = L£°@) SL-1L.. ). -Te-0¥ () CUO) = TI-S TI () pynisngoas si4ajoauoja [ fo | Cote $647) = $77 (Z) 09-09.) SET E L°6-1'6 €°OI-L'6 snddyovd sisajaAuoja J TC-LI €S-Lp 9I-ZI €I-0l 7S-OP I€I-Oll 69-19 8 LI-V'91 L8I-v'LI ujyny snjiydojoog v-€ ZTE-OE (bo) §=— ZI-T1 (9) L-9 (bp) ~=1€-97 (9) 91-89 (9) TZ t¥-6'€ (9) OTI-9'01 (9) =L I-71 (9) sinua] snjjasisidig 9I-ZI Or-8E (p) €1 (1) 11 (1) 9¢€ (1) 76 (1) 192s (9) €SI-6v1 (9) 6SI-I'ST (p) $-SMsaidouajs snjjasisidig = ge (1) = = ae LS (Z) OrI(Z) 8 bI-9 I (Z) isdajad snjjaaisidigd an 9€-ZE €I-0l OI-L L€-67 06-SL 0'S-S'b 371-3 11 9€I-9'TZI snoiuvanl snjjaaisidig = 6€ (1) LI (1) 6-8 (Z) Zb-0r (Z) t6-68 (PY. FESO) = pst (1) 4-4/l4030f snosiuoyd CIl-tid) 6e-pe() sI-er() 11-6 @ 6¢€-TE() 66-06 (4) O0'S-t'r (L) LEI-S71(L) = 9 vI-O'vI (L) snsajddyovag 40)9]14 EVO) 5£5-25:(0) 0z (1) vi (1) Sr (1) CEG) SC SEL) 6LI-V'9I (hb) = 76 I-T'LI CH) snjaidofna suody v-£ ve-O€ vI-Il 8-9 8E-0E 6L-69 0'S-S'b L1I-601 cis 11 pjoaianus sod py SI-0l 6b-bb 61-91 LI-S1 6b-6€ v1I-86 by Ea) 0'9I-6'F1 py LI-€°91 Snsavjosq0Ul SOA ¢9-¢ SE-SE 91-ZI ZI-Ol Ip-0€ S6-€8 Lots CCI-9 71 TvI-SEl upjayssoy suodpy 11-6 (pb) 6€-9€(b) LI-bI (hy) ZI-6 (%) 7e-BElh) I10I-06 (hy) €°9-6'S (pb) S9I-8SSI (hy) SLI-9'9I (pr) sijojodo DULNy 77-91 ¢s-I¢ LI- oology Tithes Available he Bats-of rans Systematics. Tastribunon) Ecology, By Amhony i. DeBiase, Aredia las Zoolgsy: 0.5... tno. 4, 19805424 popes, 168 ilps. 22 tales, ; . Publication 2307, S314) tudics in Néowrdpical Maraniatozy- Essays in ona of Philip Hershkoyitz, Edited ty: Brice Dxraticrson Find Robert M.Fimin ekiana,.Avology Ws. bo: 32. 198 8 SOG pages sah wi Paoblication £382, 845,00 ariability and, Sigameance of Parietal and Vernal: Scales in the Marine Saakes of the Genus apes » (Serperites: Hydrophiided) with Comments on the Occurrence of Sprty Scales in. the Genps, By Pant Grits ae Harold KV sris) Fieldigna. | Zeolagy,. 1.3), te, 56. 1990, 14 pages. 8 ilies; Wabi: Pablication L410) $7.00 eport ona’ Collection of Aniphibrans. and, Reptites from! Sichvan, Chink. ‘By, Robert PF. tage, Eni “Zhao. H. Bradley Shafter, and Guanfe Wu Hieldiana, Zcolog}, 0.5. NG. 38, VO90s 24. pages, 1 illus, ys tables... e Publication 1413, 510.00 @ Birds of Mt Isarog) National, Park, Southerh Lizon, ‘Philippines, withy Particular Reference to Altitudinal Distribution: By Sievéen Mi Goodraadiand Pedro C. Gonzales: Preliany. Zoology, 1.8., =no.. 60. 1990; 32 BABES: 6 ilius:,\5 tables, Publicatioa 1T4i5, $12.00 Binviogeiictic” Systeratics of the AMMOlODS Cyan of Ranid Frogs im Soniheastenn Asia and the Greater nai nis cable By Parton EARS Fieldiana: ee THs), 20,163, 199%. 42pages, 30 Vlus., 2 tables: Belch oaban 1A23, $14.00, tematic ‘Ronen ée Paitiveioe Macaques (Printed Cepeunihcetine, Maca Tacos subspp.). BY, ae Fooden. Fieldiana: ooltey, n. Ba no, 64/1991. 44 pages.) 11. pe 1] tables. cere a pts ee . Fie hs aber Japa Sis00 : aie) MUSEUM OF NATURAL HISIORY Sank > Library Publicalions Divisiqa”’ Roosevelt Road atLake Shore, Dive htt eau eatltpa th hari Ae ee ves. a Ce _— _ a i, oie a ee m ‘sete Ce a ee ey git gh a Te ee ' URBANA $CHGO ILLINOIS- SERIES C001 -93 § UNIVERSITY OF 2 590.5FIN FIELDIANA : ZOOLOGY. $ NEW ~75 199 66 — * Se ee, eter es, ~ SOP ees a on Sa aman . 9) von oY DAs oy eae