UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BIOLOGY APR 9 1992 Zoology ystematic Review of Philippine Macaques ^imates, Cercopithecidae: lacaca fascicularis subspp.) : ooden 91 1476 'ontributors to Fieldiana i mb ■ i nd arch as pai ; ( ■• rjj ^ •:., ■ ... (1 thl ' . .. ' ll :\ which 'educes ihc publication time, Contributions f /: . of ability to pay page ;f ; e.ii i .-.•■■■.;■ hrec c i ilen ies of th< : ■ •■ IU in ■ origin ,; . | is tw ■ ■ - ■ ' to reviewers befor Editor, Fieldiana, Field Museum of Natural Histot si ■■• h ; - •' by lb nch j ising MS-DOS, also subi Lit & 4, Wane PC, Sar a • most cast erature Cited." - used after al reviate ii ma. ■■■< nual of Style (13th ed .) ■ ■ ■'.■■ i-l •dicui, Vaxonon cat paper dial Sources for tip. or*; should follow . ' v! tJnivi itanfo d, Cal 1.963. >mp : ' '• tics i iloj 51 5 3-80. In Br pp. 785-821. In Steward, J. II, ed., Handb ■ ,. I.Jui merii in the te> its musi t< author upon ,;..■; idin rwiil ■••■; and queries an< set i ■ ■ ■ lior-j'cneratcd changes in page proofs i H PAF£J FTELDIANX Zoology NEW SERIES, NO. 64 Systematic Review of Philippine Macaques (Primates, Cercopithecidae: Macaco, fascicularis subspp.) Jack Fooden Research Associate Division of Mammals Department of Zoology Field Museum of Natural History Chicago, Illinois 60605-2496 Accepted December 6, 1990 Published May 31, 1991 Publication 1426 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1991 Field Museum of Natural History Library of Congress Catalog Card Number: 91-70915 ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents Abstract 1 Introduction 1 External, Cranial, and Molecular Char- acters 3 Pelage Color 3 External Measurements 9 Cranial Characters 9 Blood Proteins 12 Natural History 13 Systematics 21 Subspecific Taxonomy 21 Subspecies Accounts 22 Macaca fascicularis fascicularis (Raffles, [1821]) 22 Synonymy 22 Type 22 Type Locality 22 Distribution 22 Diagnosis 22 Macaca fascicularis philippinensis I. Geoffrey, [1843] 22 Synonymy 22 Type 23 Type Locality 23 Distribution 23 Diagnosis 24 Taxonomic Note 24 M. f. fascicularis/ M. f philippinensis Contact Zone 24 Synonymy 24 Distribution 24 Diagnosis 24 Zoogeography and Evolution 24 Summary 28 Acknowledgments 28 Literature Cited 29 Appendix 1 : Specimens Examined 32 Appendix 2: Gazetteer 34 List of Illustrations 1 . Known locality records of Philippine M. fascicularis 2 2. Selected standard skins of subadult or adult male Philippine M. fascicularis ... 4 3. Geographic variation of dorsal pelage color saturation in samples of Philip- pine and Sabah M. fascicularis 6 4. Skull of adult female Philippine M. fas- cicularis 10 5. Skull of adult male Philippine M. fascicu- laris 11 6. Size extremes in adult male skulls of Philippine M. fascicularis 12 7. Latitudinal variation of greatest skull length in adult Philippine and Sabah M. fascicularis 13 8. Accessory entostyle variation in Philip- pine M. fascicularis, lingual view of left M 2 " 3 18 9. Known limits of distribution of Philip- pine M. f fascicularis, M. f philippinen- sis, and contact-zone populations 25 10. Map of Southeast Asia showing limits of distribution of pale to intermediate subspecies of M. fascicularis, location of deep-water insular habitats of dark sub- species of M. fascicularis, and depth of intervening channels 26 1 1 . Dark non-Philippine M. fascicularis skins compared with pale and dark Philippine M. fascicularis skins 27 List of Tables 1 . Dorsal pelage color: saturation in Phil- ippine and Sabah M. fascicularis 5 2. External measurements recorded by col- lectors for adult Philippine M. fascicu- laris 7 3. Interisland variation of external mea- surements in adult Philippine and Sa- bah M. fascicularis 8 4. Relative length of tail, hind foot, and ear in various age/sex classes of Philip- pine M. fascicularis 9 5. Cranial measurements: insular, latitudi- nal, and local variation in adult Philip- pine and Sabah M. fascicularis 14 6. Cranial measurements and ratios in var- ious age/sex classes of Philippine M. fascicularis 17 7. Accessory entostyles: frequency in Phil- ippine and Sabah M. fascicularis 17 8. Local variation of accessory entostyle frequency in M 2 and M 3 in Mindanao M. fascicularis 19 9. Blood-protein allele frequencies at poly- ni morphic loci in Philippine and non- 1 1 . Monthly distribution of births in cap- Philippine M. fascicularis 19 tive colony of Philippine M. fascicularis 10. Habitat observations of Philippine M. maintained outdoors at Tanay, near fascicularis 20 Manila 21 IV Systematic Review of Philippine Macaques (Primates, Cercopithecidae: Macaca fascicularis subspp.) Jack Fooden Abstract Based on study of 352 museum specimens and review of relevant literature, the systematics of Philippine macaques (Macaca fascicularis) is examined. Information is presented concerning pelage color, external measurements, cranial characters, blood proteins, and natural history. Two subspecies of Philippine macaques are recognized— M. f. fascicularis and M.f philippinen- sis; between the exclusive ranges of these subspecies is a subspecific contact zone. Subspecific synonymies, type data, distribution summaries, and diagnoses are provided. Zoogeographic hypotheses to account for the present distribution of Philippine macaques are investigated. All known Philippine macaque localities are listed in an annotated gazetteer. Introduction The occurrence of monkeys in the Philippine archipelago was revealed to science nearly 300 years ago in separate reports by the English seaman W. Dampier (1697, p. 220) and the German mission- ary G. J. Camel (in Petiver, 1705, p. 2199). The first known museum specimen, now lost, was sent from Luzon to London in 1837 by H. Cuming (1839, p. 93); on receipt, this specimen was iden- tified by Waterhouse (1838, p. 8) as Macacus cy- nomolgus (= Macaca fascicularis). Based on living captives, I. Geoffrey ([ 1 843], p. 568; 1851, pp. 29, 93) attempted to distinguish Philippine macaques from non-Philippine M. fascicularis. The first zo- ologist to work with reasonably adequate series of properly localized specimens from various islands of the archipelago was Mearns ( 1 905, p. 426). Lat- er, Mearns collected additional specimens that were studied by Hollister ( 1 9 1 3, p. 328). The treatment of macaques in Taylor's (1934, p. 336) classic re- view of Philippine mammals essentially follows those of Mearns and Hollister. More recently, Lawrence (1939, p. 62) reported on a small col- lection of macaques that she obtained in Luzon and Mindoro, and Sanborn (1952, p. 113) briefly reviewed the large collection obtained in six is- lands in 1 946-1 947 by H. Hoogstraal, D. S. Rabor, and associates. Additional unreported material, collected principally by D. S. Rabor and associ- ates, has accumulated subsequently. In current checklists, Philippine macaques are routinely al- located to one or two endemic subspecies of M. fascicularis (cf. Hill, 1974, p. 522; Napier, 1981, P. 12). The present review is based on study of 352 specimens of Philippine macaques (Appendix 1) augmented by a survey of relevant literature. In- formation is available concerning monkeys col- lected or observed at 1 54 localities in 29 islands of the archipelago (fig. 1, 1 54 localities represented by 1 28 locality symbols; Appendix 2). To establish a broader geographic context for this study, ex- ternal and cranial characters of Philippine speci- mens are compared with those of 132 specimens of M. fascicularis collected in neighboring Sabah, northern Borneo. Specimens examined are pre- served in the following institutions, which here- after are cited by means of the indicated abbre- viations: AIUZ AMNH Anthropologisches Institut der Uni- versitat Zurich, Zurich American Museum of Natural His- tory, New York FTELDIANA: ZOOLOGY, N.S., NO. 64, MAY 31, 1991, PP. 1-44 -T" 118 122 c Sabah 126° 300 km • Specimens examined o Literature records Fig. 1 . Known locality records of Philippine M. fascicularis. For details, see Gazetteer (Appendix 2). Key to Locality Numbers— Balabac: /, Minagas Point. Palawan: 2, Mantalingajan, Mount. 3, Brookes Point; Macagua. 4, FIELDIANA: ZOOLOGY bbm Bernice Bishop Museum, Honolulu (cited for field catalog data only; no Philippine macaque specimens in collection) bm(nh) British Museum (Natural History), London fmnh Field Museum of Natural History, Chicago irsn Institut Royal des Sciences Natu- relles de Belgique, Brussels mcz Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts mmnh Bell Museum of Natural History, University of Minnesota, Minne- apolis mnhn Museum National d'Histoire Na- turelle (Mammiferes), Paris nhrm Naturhistoriska Riksmuseet, Stock- holm pnm Philippine National Museum, Ma- nila rmnh Rijksmuseum van Natuurlijke His- toric Leiden siconbrec Simian Conservation, Breeding & Research Center, Inc., Tanay, Lu- zon, Philippines SMTD UMMZ im bo UPLBZD I'SNM ZMB Staatliches Museum fur Tierkunde, Dresden Museum of Zoology, University of Michigan, Ann Arbor University of the Philippines at Los Banos, College of Forestry, Los Ba- nos, Luzon, Philippines University of the Philippines at Los Banos, Zoology Department, Los Banos, Luzon, Philippines National Museum of Natural His- tory, Washington, D.C. Zoologisches Museum des Hum- boldt-Universitat, Berlin External, Cranial, and Molecular Characters Pelage Color Dorsal pelage color in Philippine M . fascicularis varies from pale yellowish brown agouti (oliva- ceous) through golden brown agouti to dark brown agouti (fig. 2). Erythrism is variable. Individual dorsal hairs in the interscapular region of adult Lapulapu. 5, Puerto Princesa; Puerto Princesa area; Puerto Princesa, E; Puerto Princesa, "Mt. wooded area." 6, Tarabanan. 7, Malabusog. 5, Malampaya Sound. Cuuon: 9, Makinis; San Pedro. Busuanga: 10, Dimana. 77, San Nicolas. Mindoro: 12, Bulalacao. 13, Pinamalayan. 14, Pasi. 15, Naujan Lake National Park. 16, Calapan. 17, Alag River. 18, Halcon, Mount. Tablas: 19, Tablas I. Romblon: 20, Romblon I. Sibuyan: 21, Guitinguitin, Mount. Masbate: 22, Masbate I. Marinduque: 23, Marinduque I. Luzon: 24, Albay Prov. 25, Mount Isarog National Park. 26, Lopez. 27, Quezon National Park. 28, Batangas. 29, Mahayahaya. 30, Tuy. 31, Santa Cruz. 32, Bataan National Park. 33, Maquiling, Mount. 34, Los Banos. 35, Jalajala. 36, Daraitan, Mount. 37, Manila. 38, Bulacan Prov. 39, Biak-na-bato National Park. 40, Zambales Prov. 41, Pangasinan Prov. 42, Aurora Memorial Park. 43, Data, Mount. 44, Dimalasud Barrio; San Mariano Munic. 45, Fuyot Spring National Park. 46, Cagayan Valley. 47, Barit. 48, Bessang Pass National Park. 49, Lagangilang. 50, Massisiat. 51, Laoag. 52, Nagpartian. Catanduanes: 53, Catan- duanes I. Samar: 54, Catubig River. 55, Matuguinao. 56, San Sebastian. 57, Sohoton Natural Bridge National Park. Maripipi: 58, Maripipi, 2 km N, 3 km W; Maripipi, 2 km N, 4 km W; Maripipi I. Biliran: 59, Naval, 5 km N, 10 km E; Sayao, Mount. Panay: 60. Calantas forest. Guimaras: 61, Guimaras I. Leyte: 62, Patoc Barrio. 63, Baybay, 8.5 km N, 2.5 km E; Pangasugan, Mount. 64, Balinsasayo Barrio. Bohol: 65, Sandayong. 66, Cantaub. Cebu: 67, Cebu I. Negros: 68, Canlaon National Park; Canlaon Volcano. 69, Pagyabonan. 70, Amio; Naliong. 71, Kabungahan. 72, Pamo-at. 73, Balangbang; Kandomao. 74, Kauitan. 75, Inubungan. 76, Siaton, 10 km N. 77, Bufiga Barrio. 78, Balinsasayo, Lake; Balinsasayo, Lake, [N bank]; Balinsasayo, Lake, 6 km N and 14 km W of Dumaguete City; Pamplona, 12 km S, 8 km W; Pamplona, 18 km S; Talinis. 79, Pamplona, near. 80, Mabaja. Siquuor: 57. Sequijor I. Mindanao: 82, Surigao. 83, La Union. 84, Agusan River. 85, Mainit Hot Spring National Park. 86, Cateel River and Agusan River, crest between. 87, Baganga River. 88, Cateel River, lower. 89, Caraga. 90, Manay River. 91, Kamansi. 92. Sumlog River. 93, Parak Creek; Tagulaya River. 94, McKinley, Mount, E slope, 4800 ft; McKinley, Mount, E slope, 6400 ft. 95, Apo, Mount; Mainit, Mount Apo, 3800 ft. 96, Matutungan. 97, Badiang; Pantod, Mount. 98, Kibawalan. 99, Caburan. 100, Busaw, Mount. 101, Maculi Point. 102, Burungkot. 103, Bugusan. 104, Camp Vicars-Malabang, between. 105, Lanao, Lake. 706. Pantar. 707, Tangub. 705, Catagan. 709, Masawan, Mount Malindang, 3500-4500 ft; Masawan, Mount Malindang, 4400-5000 ft. 770, Gubat; Libu; Sigayan; Situbo; Tacuta; Tampalan. 777, Canibongan. 772, Mamara. 77 J, Bucong. 114, Kabasalan River. 775, Santa Maria. 776, Banga, Port. 777. Pulunbato, Mount; Zamboanga; Zamboanga area. 775, Ayala; San Ramon. Pangapuyan: 779, Pangapuyan I. Balut: 720, Balut I. Cagayan Sulu: 727, Cagayan Sulu I. Basilan: 722, Isabela. 123, Camp No. 2-Camp No. 3, between; Camp No. 4-Camp No. 5, between. 124, Tiputipu. 725, Basilan I., E end. Jolo: 726, Crater Lake Mountain, foot. Tawitawi: 727, Tawitawi I. 725, Balimbing. FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES Fig. 2 . Selected standard skins of subadult or adult male Philippine M. fascicularis — fmnh 6 7 7 3 3 (left), Mindanao: Tacuta (saturation index value, 1.0); fmnh 66348 (center), Negros: Balinsasayo, Lake, N bank (2.0); fmnh 62913 (right), Palawan: Puerto Princesa, E (3.0). Scale bar 10 cm. (Photo by John Weinstein, Division of Photography, FMNH.) males are 3-5 cm long. Each hair is pale at the root, dark brown over most of its length, and marked subterminally by one or two yellowish to gold bands; the tip of each hair is blackish. In pale specimens, the subterminal bands are broader and paler yellowish than in dark specimens. The crown is somewhat more brightly colored than the back and usually bears an irregular tuft, crest, or cowlick at the vertex. The anterior edge of the crown is bordered by a blackish superciliary band. Hairs on the side of the head, below the ear, are pale ochraceous gray, elongated, and anteriorly directed; these anteriorly directed subauricular hairs meet the posteriorly directed cheek hairs to form a prominent pale lateral facial crest. The face is thinly haired; facial skin is brown, except for the upper eyelids, which are sharply defined whit- ish. On the proximal part of the outer surface of the limbs, pelage is approximately the same color as on the adjacent surface of the trunk; more distally, pelage on the outer surface of the limbs becomes paler— variably gold to yellowish to pale gray at the wrists and ankles. The dorsal surface of the tail is blackish proximally, becoming pale grayish distally. Pelage on the ventral surface of the trunk and tail and on the inner surface of the limbs is thin and pale gray to whitish. Dorsal pelage color is generally similar in adult males, adult females, and juveniles collected in the same area, but the pelage in adult males is longer and sleeker than in adult females and juveniles. The pelage of young infants (fmnh 33510, 3351 1; probably less than age 6 months, judging from dental eruption) is fine in texture, blackish dor- sally, and grayish ventrally. In older infants (fmnh 66345, 67740, 67741, 75600, 75601; probably age 6-12 months), pelage texture and color approach those in juveniles and adult females. Paleness or darkness (saturation) of dorsal pel- FTELDIANA: ZOOLOGY Table 1 . Dorsal pelage color: saturation in Philippine and Sabah M. fascicularis. Frequencies (% ) at saturation index values Inter- Mean Pale mediate Dark saturation Island : N 1.0 1.5 2.0 2.5 3.0 index value Philippine Islands Balabac (7) 1 100.0 3.0 Palawan 12 25.0 75.0 2.9 Palawan, S (3) 3 33.3 66.7 2.8 Palawan, N (4-6) 9 22.2 77.8 2.9 Culion (9) 6 83.3 16.7 2.6 Busuanga (70) 1 100.0 2.0 Mindoro (12-14, 16-18) 7 28.6 71.4 2.9 Luzon 24 16.7 83.3 2.9 Luzon, S (26, 28, 29. 33. 36. 38) 12 100.0 3.0 Luzon, N (43. 44. 46. 47. 49. 52) 12 33.3 66.7 2.8 Samar (55) 3 100.0 3.0 Leyte (62. 64) 5 100.0 3.0 Negros, S (70-75. 77. 78. 80) 45 6.7 35.6 33.3 24.4 2.4 Mindanao 68 47.1 13.2 19.1 13.2 7.4 1.6 Mindanao, NE (82, 84) 2 100.0 3.0 Mindanao, SE (91, 92, 94-97) 12 16.7 33.3 33.3 16.7 2.2 Maculi Pt. (101) 2 100.0 2.5 Burungkot; Bugusan (102, 103) 4 25.0 25.0 25.0 25.0 2.1 Pantar (106) 8 25.0 62.5 12.5 1.9 Catagan (108) 4 25.0 50.0 25.0 1.9 Bucong (7/ 3) 5 80.0 20.0 1.1 Katipunan Munic. (110-112) 23 78.3 17.4 4.3 1.1 Zamboanga area (777, 775) 8 100.0 1.0 Balut (720) 1 100.0 3.0 Cagayan Sulu (727) 1 100.0 2.0 Basilan (722. 72 J, 725) 3 66.7 33.3 1.7 Tawitawi (127) 2 100.0 1.0 All 179 19.0 Sabah 7.8 17.9 21.2 34.1 2.2 Borneo 58 77.6 10.3 12.1 1.2 1 Excludes infants. 2 Locality numbers in parentheses (see fig. 1). Intraisland regional data provided for Palawan, Luzon, and Mindanao. age color in Philippine macaques varies geograph- ically, as previously indicated by Mearns (1905, p. 426), Hollister (1913, p. 328), and Sanborn (1952, p. 1 13). In order to analyze this variation, I have selected five subadult or adult male skins to serve as standards of comparison. These five skins include two of the palest available Philippine specimens (fmnh 65440, 67733; pale yellowish brown), two of the darkest specimens (fmnh 629 1 3, 87718; dark brown), and one intermediate speci- men (fmnh 66348; golden brown). Specimens that match the pale standards have been assigned a saturation index value of 1 .0, those that match the intermediate standard have been assigned a value of 2.0, and those that match the dark standards have been assigned a value of 3.0 (fig. 2). Speci- mens marginal in saturation between the primary standards have been assigned values of 1 .5 or 2.5. Infant skins (from specimens that lack permanent teeth) are excluded from this analysis. Of 14 islands for which postinfant skins are available, 4 are represented exclusively by pale or intermediate specimens (saturation index values 1.0-2.0), 8 are represented exclusively by dark specimens (values 2.5-3.0), and 2 are represented by pale, intermediate, and dark specimens (values 1.0-3.0) (table 1; fig. 3). Islands represented ex- clusively by pale or intermediate specimens are Tawitawi (N = 2), Cagayan Sulu (1), Basilan (3), and Busuanga ( 1 ). Islands represented exclusively by dark specimens are Balabac ( 1 ), Palawan ( 1 2), Culion (6), Mindoro (7), Luzon (24), Samar (3), Leyte (5), and Balut (1). Islands represented by pale, intermediate, and dark specimens are Min- danao (67) and Negros (45). Within Mindanao, western samples are pale to intermediate, central FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES Fig. 3. Geographic variation of dorsal pelage color saturation in samples of Philippine and Sabah M.fascicularis. For details, see table 1 . FIELDIANA: ZOOLOGY Table 2. External measurements (mm) recorded by collectors for adult Philippine M. fascicularis. Question marks indicate measurements of doubtful validity. Island Loc. no.' Nlat. Head and body length Tail length Hind foot length Ear length Females Palawan 2 2 8°48' 420 7417 108 30 Busuanga ir- 12°04' 420 530 115 28 Luzon 50' 17°35' ?950 515 125 52* i8°3r 420 580 131 Negros 71 9°30' 424, 462 536, 538 118, 130 24,35 78 9°21' 400,410,433 460, 470, 455 126, 122, 125 26, 27, 29 Mindanao . . .5 7°30' 420 539 115 29 103 7°27' 433 564 130 118 TOO' ?370 7570 115 32 95 6 6°59' 445, 465 455, 445 128, 120 35,35 102' 6°52' 430 551 122 Tawitawi 127 5° 10' 394 Males 504 105 31 Balabac P 7°54' ?332, 450 544, 510 132, 126 40,42 Palawan 5 9°44' 467, 475 487, 550 132, 136 39,50 6 2 io°or 530 560 135 43 7 2 10°04' 410 7 510 7 120 7 31 7 Culion 9* n°5r 463 496 132 36 9> n°5r 496, ?563 536, 548 126, 134 38,36 Busuanga 10 12°03' ?381 7595 145 36 Mindoro 13 13°02' 450 580 50 14 13°07' 476 552 47 17 w 13°22' 520 580 140 Luzon 33 14°08' 500 592 36 14°37' 505 510 135 43 50' 17°35' ?1060, 71080, 71115 550, 530, 560 140, 145, 140 52* 18°31' 500 625 150 Leyte 62 11°05' 7425 490 130 27 ... ii 11°00' 7430 7700 135 64 10°40' 7410 590 135 30 Negros 71 9°30' 7686 564 140 32 72 9°30' 466 575 145 36 70 9°24' 539 578 147 32 Mindanao 106" 8°04' 500 600 137 39 94 7°06' 508 536 137 44 95 6°57' 7575 7425 137 40 9P 6°56' 7390, 490 7615, 585 7292 117" 6°54' 445 510 102' 6°52' 7443 7633 136 39 97 6°49' 490 531 140 40 Tawitawi 127 5° 10' 386 6 507 6 118 6 30* 1 See figure 1 . 2 Measurements from field catalog, bbm. ' Measurements from field catalog, fmnh. 4 Measurements from field catalog, usnm. 5 Zamboanga del Sur Prov. 6 Measurements from Mearns ( 1 905, pp. 429, 430). 7 Abnormally small specimen; measurements excluded from summary statistics. 8 Makinis. * San Pedro. '" Measurements from Hollister (1913, p. 329). " Leyte I.; measurements recorded on skull, usnm. 12 Measurements from Martens (1876, p. 206). and southeastern samples are mixed pale-inter- mediate-dark, and a small northeastern sample is dark. In summary, available specimens of Phil- ippine M. fascicularis are pale to intermediate in the south-central islands of Tawitawi, Cagayan Sulu, and Basilan and in adjacent western Min- danao; they are mixed pale— intermediate-dark in central and eastern Mindanao and in southern Ne- gros; and they are dark elsewhere in the Philip- pines (except Busuanga, which is represented by one intermediate skin). This pattern of color vari- ation in Philippine M. fascicularis does not appear to be closely correlated with climate variation (Djambatan N.V., 1964, p. 10). Fifty-eight skins FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES Table 3. Interisland variation of external measurements (mm) in adult Philippine and Sabah M. fascicularis. For details, see table 2. Island 1 Head and body length Tail length Hind foot length Ear length Females Philippine Islands Palawan 420 108 30 Busuanga 420 530 115 28 Luzon 420 547.5 (2) 515-580 128.0(2) 125-131 Negros 425.8 ± 23.9 (5) 491.8 ± 41.6(5) 124.2 ± 4.5(5) 28.2 ± 4.2 (5) 400-462 455-538 118-130 24-35 Mindanao 438.6 ± 17.2(5) 510.8 ± 56.3(5) 121.7 ± 6.3(6) 32.8 ± 2.9 (4) 420-465 445-564 115-130 29-35 Tawitawi 394 504 105 31 All 426.9 ± 20.4(14) 510.1 ± 45.3(14) 120.9 ± 7.8(16) 30.1 ± 3.7(12) 394-465 445-580 105-131 24-35 Sabah Borneo 2 393.3 ± 14.8(18) 494.1 ± 29.7(18) 116.3 ± 4.1 (18) 24.8 ± 4.1 (16) 369-420 455-555 Males 108-123 20-35 Philippine Islands Balabac 450 527.0(2) 129.0(2) 41.0(2) 510-544 126-132 40-42 Palawan 490.7 ± 34.3 (3) 532.3 ± 39.6(3) 134.3 ± 2.1 (3) 44.0 ± 5.6 (3) 467-530 487-560 132-136 39-50 Culion 479.5 (2) 526.7 ± 27.2 (3) 130.7 ± 4.2(3) 36.7 ± 1.2(3) 463^96 496-548 126-134 36-38 Busuanga 145 36 Mindoro 482.0 ± 35.4 (3) 570.7 ± 16.2(3) 140 48.5 (2) 450-520 552-580 47-50 Luzon 501.7 ± 2.9 (3) 561.2 ± 41.8(6) 142.0 ± 5.7 (5) 43 500-505 510-625 135-150 Leyte 540.0 (2) 133.3 ± 2.9(3) 28.5 (2) 490-590 130-135 27-30 Negros 502.5 (2) 572.3 ± 7.4 (3) 144.0 ± 3.6(3) 33.3 ± 2.3 (3) 466-539 564-578 140-147 32-36 Mindanao 486.6 ± 24.4 (5) 552.4 ± 38.3(5) 137.4 ± 1.5(5) 40.4 ± 2.1 (5) 445-508 510-600 136-140 39-44 All 487.9 ± 27.1 (19) 550.7 ± 35.5 (27) 137.2 ± 6.0(26) 39.0 ± 5.9 (22) 445-539 487-625 126-150 27-50 Sabah Borneo 451.8 ± 30.8(18) 559.8 ± 48.1 (18) 132.8 ± 8.2(18) 29.4 ± 5.7(18) 400-501 470-650 115-148 22-42 1 Where insular sample size exceeds two, entries indicate mean ± SD (N) and extremes. 2 Excludes one abnormally large specimen: fmnh 68700, Kretam Besar, HB = 450 mm. of M. fascicularis collected in Sabah, northern Bor- neo, are pale to intermediate, matching in satu- ration the Philippine skins collected in Tawitawi, Cagayan Sulu, Basilan, and western Mindanao. The similarity of pale pelage color in M. fascicu- laris in northern Borneo and Mindanao was pre- viously indicated by Buzeta and Bravo (1850, p. 40). Erythrism occurs variably in specimens that are pale (usnm 114697, adult male, Mindanao), in- termediate (fmnh 66329, adult female, Negros), and dark (fmnh 62902, 62913, usnm 477840, all adult males collected in Palawan). In erythristic specimens, the entire length of each dorsal hair, including both pale and dark segments, is more intensely pigmented than in nonerythristic speci- mens. Erythrism appears to be somewhat more prevalent in Palawan than in other islands; in Pa- lawan, the color of strongly erythristic specimens tends toward dark chestnut. Sabah specimens av- FIELDIANA: ZOOLOGY Table 4. ± SD(N). Relative length of tail, hind foot, and ear in various age/sex classes of Philippine M. fascicularis: mean Relative hind Head and body length Relative tail length foot length Relative ear length Age/sex class (mm) (T/HB x 100) (HF/HB x 100) (E/HB x 100) Infants 238.7 ± 48.9 (9) 138.9 ± 21.6(9) 38.2 ± 4.9 (8) 12.9 ± 1.8(6) Juveniles 360.7 ± 40.6 (30) 126.4 ± 10.2(30) 32.2 ± 2.0 (28) 9.4 ± 1.4(27) Subadult females 396.0 ± 6.2 (4) 123.8 ± 19.7(4) 29.8 ± 1.7(4) 8.4 ± 0.7 (2) Subadult males 422.8 ±41.1(4) 124.6 ± 6.5 (4) 31.0 ± 4.0(3) 8.2 ± 1.4(2) Adult females 426.9 ± 20.4(14) 119.6 ± 12.6(13) 28.4 ± 1.8(14) 7.0 ± 0.7(11) Adult males 487.9 ± 27.1 (19) 113.3 ± 7.8 (19) 27.8 ± 1.6(14) 8.5 ± 1.3(14) erage somewhat less erythristic than Philippine specimens. Albinism is sporadic in Philippine M. fascicularis (I. Geoffrey, [1843], p. 568; Sclater, 1875, p. 349; Elera, 1915, p. 34; mnhn 373/265; siconbrec, living captive observed 4 Aug. 1989). In a series of three dark brown specimens col- lected in Samar (fmnh 87717, adult female; 877 1 8, subadult male; 877 19, adult female), hair banding is relatively inconspicuous. Tips of the dorsal hairs in all three of these specimens are truncated and frayed. The cause of this unusual truncation and fraying is unknown. External Measurements Head and body length (HB) in adult Philippine M. fascicularis varies from 394 to 465 mm in 14 females (426.9 ± 20.4 mm, mean ± SD) and from 445 to 539 mm in 19 males (487.9 ± 27.1 mm) (tables 2, 3; cf. Rabor, 1986, p. 138); this excludes measurements of two aberrantly small adult males (siconbrec 1475, Tawitawi, HB = 386 mm; usnm 477842, Palawan: Malabusog, HB = 410 mm). Relative tail length (T/HB x 100) averages 1 19.6 in 13 adult females and 1 13.3 in 19 adult males (table 4). Relative hind foot length (HF/HB x 100) averages 28.4 in 14 adult females and 27.8 in 14 adult males; relative ear length (E/HB x 100) av- erages 7.0 in 1 1 adult females and 8.5 in 14 adult males. Weights of wild-collected specimens are known for only one adult female (ca. 4.0 kg) and one adult male (ca. 7.7 kg), both collected at Nag- partian, northern Luzon. Weights of two captive adult females are 3.75 and 3.93 kg (origins— Tawi- tawi; Mindanao: Zamboanga del Sur), and weights of three captive adult males are 4.25, 5.3, and 7.54 kg (Tawitawi; Leyte; Luzon: Daraitan, Mt.). Available external measurements of adults are inadequate to establish whether or not head and body length increases with latitude in Philippine M. fascicularis; such an increase would be ex- pected in view of the latitudinal increase of greatest skull length (see below). Available measurements also are inadequate to establish whether or not tail length varies geographically in Philippine M. fas- cicularis. Mean head and body length in Philippine M. fascicularis exceeds that in Sabah M. fascicu- laris (table 3; /-test, P < 0.00 1 , females and males); however, overlap between the Philippine and Sa- bah samples is extensive. Mean relative tail length in Sabah M. fascicularis (females, 125.6; males, 1 24.4) exceeds that in Philippine M. fascicularis. Relative length of the tail, hind foot, and ear all decrease with age in Philippine M . fascicularis (ta- ble 4). This implies that postnatal growth of these extremities is slower than that of the head and body. The extreme example is postnatal growth of the ear in females, which apparently is negligible from infancy to adulthood. Cranial Characters (figs. 4, 5) Greatest skull length (GL) in Philippine M. fas- cicularis varies from 93.7 to 1 1 5.8 mm in 75 adult females (105.3 ± 4.3 mm, mean ± SD) and from 109.7 to 139.1 mm in 89 adult males (124.0 ± 5.4 mm) (table 5); this excludes one adult male (bm(nh)) 1876.10.4.9, Luzon: Mahayahaya, GL = 109.3 mm) that is abnormally small compared with other specimens collected in the same area. Relative zygomatic breadth (ZB/GL x 100) av- erages 65.2 in adult females and 66.0 in adult males (table 6). The rostrum is relatively long and nar- row, particularly in adult males; rostral-postros- tral ratio (%) averages 49.5 in adult females and 59.0 in adult males. Supramaxillary ridges and lateral maxillary concavities in adults vary from weakly to strongly defined. Canines in adult males are relatively large. A median sagittal crest, formed FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES Fig. 4. Skull of adult female Philippine M.fascicularis—FMNH 87424, Bohol: Cantaub. (Photos by John Weinstein, Division of Photography, fmnh.) by progressive ontogenetic convergence of the temporal lines, is prominent in old males. Skull length in Philippine M. fascicularis tends to increase with latitude (figs. 6, 7). Skull length in Sabah M. fascicularis averages less than in Phil- ippine M. fascicularis (table 5; Mest, P < 0.001, females and males). However, Sabah means (fe- males, 100.2 mm; males, 116.4 mm; latitude, 5°49'N) are close to values predicted for that lat- itude by the Philippine M. fascicularis regression equations (females, 101.3 mm; males, 120.4 mm) (fig. 7). Upper molars in M . fascicularis frequently bear one or more variably defined accessory entostyles that arise from the lingual cingulum between the protocone and hypocone (fig. 8). Because M 1 is often severely worn, the incidence of accessory entostyles is most conveniently studied in M 2 and M 3 (table 7); the incidence of entostyles in M 3 (0.36, N = 425 teeth) in Philippine and Sabah M. fascicularis somewhat exceeds that in M 2 (0.23, N = 575 teeth). In most islands of the Philippine archipelago, the combined incidence of well-de- fined accessory entostyles in M 2 and M 3 in M. fascicularis is less than 0.25. However, in Min- danao (N = 242 teeth) and Mindoro (N = 1 8 teeth), the incidence is 0.44; the high frequency of acces- sory entostyles in these two islands was previously noted by Hollister (1913, p. 329). The incidence may also be higher than average in Panay (0.62, 10 FIELDIANA: ZOOLOGY Fig 5. Skull of adult male Philippine M. fascicular is— fmnh 67996, Negros: Balangbang. (Photos by John Wein- stein. Division of Photography, fmnh.) N = 8 teeth), Balut (0.50, N = 4), and Basilan (0.38, N = 8), but available samples for these three islands are small. Within Mindanao, the incidence is relatively high (0.41-0.69) in all areas sampled except the Zamboanga area (0.12, N - 8 teeth), extreme southwestern Mindanao, which is repre- sented by a total of only three skulls (table 8). In Sabah M. fascicularis, the incidence of accessory entostyles is relatively high (0.39, N = 106), as in Mindanao and Mindoro. Ontogenetically, growth of the zygomatic arch from infancy to adulthood is approximately iso- metric with growth of greatest skull length (table 6), as in other macaques. Rostral length (facial skeleton), however, increases allometrically rela- tive to postrostral length (cranial skeleton), also as in other macaques. The power function for the ontogenetic relationship between rostral length (y) and postrostral length (x) determined by the meth- od of reduced major axes is log y = 3.725 log x — FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 11 Fig 6. Size extremes in adult male skulls of Philippine M. fascicularis— usnm 125325 (left), Cagayan Sulu, ca. 7°02'N; usnm 144680 (right), Luzon: Nagpartian, 18°31'N. (Photos by Division of Photography, usnm.) 5.456 (r = 0.917); 95% confidence limits for the slope are L, = 3.716 and L 2 = 3.733. Ontogenetic development of zygomatic breadth and rostral length in Sabah M. fascicularis is similar to that in Philippine M. fascicularis. Blood Proteins Thirty-two electrophoretic blood-protein loci have been studied in Philippine M. fascicularis (Fooden & Lanyon, 1989, p. 211). Of these, 22 loci are essentially fixed in Philippine and non- Philippine M . fascicularis (i.e., frequencies of com- mon major alleles are greater than 0.95 in all pop- ulations sampled). For the 10 polymorphic loci, allele frequencies in Philippine M . fascicularis and non-Philippine M. fascicularis are compared in table 9. Unfortunately, no information is available con- cerning geographic variation of allele frequencies within the Philippine archipelago, and no blood- protein data from Bornean M. fascicularis are available for comparison with Philippine data. Part of the Philippine sample is known to have origi- nated in Mindanao; the island of origin of the balance of this sample is unknown. Because blood- protein data are not available from Bornean M. fascicularis, the Philippine sample is compared with a sample of Sumatran M. fascicularis and with a composite of all other non-Philippine samples of M. fascicularis. The most striking blood-protein difference be- tween Philippine and non-Philippine M. fascicu- laris is at the Tf locus (for abbreviations, see table 9); in Philippine M. fascicularis this locus is mono- morphic for allele D, whereas in non-Philippine populations the maximum frequency of allele D is 0.66 (West Malaysia). Philippine M. fascicularis also is less variable than non-Philippine M. fas- cicularis at loci Alb (monomorphic), IDH, PGD (monomorphic), and PI. Only locus PHI is notably more variable in Philippine M. fascicularis than in non-Philippine M. fascicularis; at this locus, allele 5 is relatively frequent (0.17-0.43) in Phil- ippine M. fascicularis, whereas it is rare (ca. 0.01) 12 FIELDIANA: ZOOLOGY 140 E E 130 - g 120 CD CO CD 2 1 10 CO 100 90 1 1 ■ l ■ ■ ■ ■ ■ ■■V — ■ ■ _^- — - * ■ ■ m ■ ■ ^ ■ ■ ■ "■ m ■ 1 ■ ■ ■ ■ ■ 'l i ■ ■ ■ - * ■ 1 m □ D ■ D • D •* □ --"CD D ■ D D D □ - * D 1 h D D * D *fc D B D ^^ § ^** D B — 5 D Fema es Males ** □ 1 Phil i ipp Sa ines bah D 1 ■ 8° 12° Latitude (N) 16 20° Fig. 7. Latitudinal variation of greatest skull length in adult Philippine and Sabah M. fascicularis (cf. table 5). Regression lines indicated for Philippine data (females, slope = 0.91 ± 0.27 [95% confidence limits], y-intercept = 96.0, r = 0.649; males, slope = 0.85 ± 0.33, ^-intercept = 1 15.5, r = 0.498). in non-Philippine M. fascicularis. The tendency for blood-protein loci that are variable elsewhere to become fixed in Philippine M. fascicularis and in other insular macaques has been noted previ- ously (Goodman et al., 1965, p. 886; Ishimoto, 1973, p. 12; Darga et al., 1975, p. 809; Nozawa etal., 1977, p. 24). Natural History Macaca fascicularis is widespread in the Phil- ippine archipelago (fig. 1). Isolated islets as small as 22 km 2 are known to support viable populations (Balut; Maripipi; Romblon). Islands in the archi- pelago that possibly or probably are not inhabited by M. fascicularis are Batan, 200 km N of Luzon; Dinagat and Siargao, 25 km NE of Mindanao; and Manuk Manka, 25 km S of Tawitawi (for details, see Gazetteer, Appendix 2). Prior to about 1960, the abundance of Philippine M. fascicularis was generally reported to be high, but subsequent trap- ping activities and forest clearance have drastically reduced populations (Cabrera, 1973, p. 252; Ra- bor, 1977, p. 35). The habitats in which Philippine M. fascicularis has been observed are extremely diverse, ranging from the seashore to oak-pine montane forest at FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 13 Table 5. fascicularis. Cranial measurements (mm): insular, latitudinal, and local variation in adult Philippine and Sabah M. Loc. Zygomatic Postrostral Island no. 2 Nlat. Greatest length breadth length Rostral length Females Philippine Islands Palawan 104.9 (2) 67.2 (2) 75.9 (2) 36.5 (2) 2 8°48' 102.7 67.8 74.7 35.1 6 10°01' 107.1 66.6 77.1 37.8 Busuanga 11 12°04' 105.5 72.5 74.0 38.9 Luzon ... 111.2 ± 3.2(9) 69.0 ± 1.4(9) 79.1 ± 3.6(9) 40.4 ± 2.3 (9) 105.2-115.8 67.2-71.3 75.1-87.2 36.8^3.9 28 13°45' 110.9 69.4 76.2 40.9 33 14°08' 105.2 71.3 75.6 36.8 46 17°25' 111.2 68.2 78.7 41.2 50 17°35' 108.6 68.4 75.1 41.3 52 18°31' 112.9 ± 2.4(5) 68.7 ± 1.5(5) 81.2 ± 3.4(5) 48.8 ± 2.5 (5) 110.0-115.8 67.2-70.5 78.5-87.2 37.7^3.9 Samar 55 12°08' 107.3 (2) 66.6 (2) 76.3 (2) 39.0 (2) 104.9-109.6 65.6-67.5 76.0-76.6 35.4-^2.5 Maripipi 58 1 1°47' 110.2(2) 68.7 (2) 76.7 (2) 44.0 (2) 108.4-112.0 67.9-69.5 76.3-77.2 41.5-46.5 Leyte 63 10°46' 104.8 67.5 75.8 36.5 Bohol 106.5 ± 1.5(4) 66.3 ± 1.6(4) 75.3 ± 0.5 (4) 39.2 ± 1.6(4) 105.0-107.9 64.5-68.2 74.6-75.9 37.3-41.1 66 9°48' 106.0 ± 1.5(3) 65.6 ± 1.3(3) 75.3 ± 0.6 (3) 39.1 ± 1.9(3) 105.0-107.7 64.5-67.0 74.6-75.9 37.3-41.1 65 9°45' 107.9 68.2 75.3 39.7 Negros 105.3 ± 2.6(19) 70.7 ± 1.7(19) 74.8 ± 1.7(19) 37.4 ± 2.3(19) 100.8-110.0 68.0-73.7 71.7-79.1 34.0-42.6 71 9°30' 106.6(2) 71.2(2) 76.0 (2) 38.4 (2) 103.2-110.0 68.8-73.7 72.8-79.1 37.6-39.1 79 9°30' 104.7 72.4 76.0 35.4 70 3 9°26' 104.0 ± 2.4(4) 69.7 ± 1.7(4) 74.8 ± 0.3 (4) 36.8 ± 1.7(4) 101.6-107.4 68.0-71.7 74.4-75.0 35.1-39.1 78" 9°23' 104.4(2) 70.5 (2) 74.5 (2) 36.0 (2) 101.9-106.9 68.1-72.9 73.1-75.8 35.7-36.2 75 9°23' 106.3 ± 2.1(3) 70.3 ± 1.6(3) 75.9 ± 0.7 (3) 38.3 ± 1.7(3) 104.8-108.7 69.0-72.1 75.2-76.6 36.8^0.2 78 s 9°22' 107.7(2) 71.7(2) 73.3 (2) 41.1(2) 106.5-108.9 70.3-73.0 72.2-74.3 39.5^2.6 78 6 9°21' 104.9 ± 2.9 (4) 70.7 ± 0.9 (4) 74.1 ± 1.6(4) 37.3 ± 2.9 (4) 100.8-107.7 69.6-71.7 71.7-75.0 34.3^1.1 . . .7 9°15' 103.2 72.1 74.8 34.0 Mindanao 102.6 ± 4.4 (25) 68.4 ± 2.5 (26) 75.0 ± 2.6 (25) 35.7 ± 2.8 (26) 93.6-114.1 63.6-73.6 71.3-82.8 30.7^11.0 110 s 8°31' 108.5(2) 73.1 (2) 77.8 (2) 40.1(2) 102.8-114.1 72.5-73.6 72.8-82.8 39.2-41.0 HO 9 8°31' 104.7 68.3 75.7 37.5 HO 10 8°31' 100.0 66.0 75.3 35.5 7/0" 8°31' 100.5 ± 2.9 (3) 68.6 ± 1.2(3) 74.7 ± 1.2(3) 34.6 ± 3.0 (3) 98.2-103.7 67.5-69.8 73.5-75.9 31.6-37.5 110 12 8°31' 102.0 73.2 73.7 35.5 111 8°25' 98.3 67.8 72.3 33.1 112 8°25' 100.3 (2) 65.0 (2) 73.1(2) 33.8 (2) 98.4-102.1 63.6-66.4 72.5-73.7 32.2-35.3 109 8°20' 103.8 (2) 67.4 (2) 75.6 (2) 35.4 (2) 102.1-105.5 66.5-68.3 74.5-76.7 34.5-36.2 108 8° 10' 100.3 ± 3.1 (3) 66.7 ± 1.0(3) 74.9 ± 2.5 (3) 34.1 ± 2.7(3) 96.8-102.7 65.6-67.4 72.2-77.2 32.2-37.2 106 8°04' 107.9 (2) 68.7 (2) 76.3 (2) 40.1(2) 105.7-110.0 67.7-69.6 75.4-77.1 39.1^11.0 14 FIELDIANA: ZOOLOGY Table 5. Continued. Loc. Zygomatic Postrostral Island no.- Nlat. Greatest length breadth length Rostral length 107 8°03' 101.1 68.7 71.8 36.0 113 7°54' 99.6 68.2 73.5 33.8 103 7°27' 108.3 68.8 77.7 36.6 118 7°00' 102.5 70.2 74.6 35.1 95 6°59' 107.4 70.7 (2) 70.0-71.4 79.1 36.6 (2) 33.5-39.6 117 6°54' 93.6 64.3 71.3 30.7 Balut 120 5°24' 99.0 63.8 71.9 33.8 Unknown 106.0 ± 2.4 (9) 66.5 ± 1.7(9) 103.0-109.6 64.4-69.3 All 105.3 ± 4.3(75) 68.6 ± 2.5 (76) 75.6 ± 2.7 (66) 37.4 ± 3.2 (67) 93.6-115.8 63.6-73.7 71.3-87.2 30.7-46.5 Sabah Borneo 1 5°49" 4 100.2 ± 3.8(28) 65.9 ± 3.6 (28) 73.2 ± 2.5 (26) 33.6 ± 2.4 (26) 93.6-107.8 59.4-72.5 69.4-77.8 29.9-39.0 Males Philippine Islands Balabac Palawan Culion Mindoro Luzon 1 Maripipi Biliran Panay / 7°54' 116.3(2) 80.6 (2) 79.5 (2) 44.7 (2) 115.9-116.7 80.4-80.8 79.0-79.9 44.6-44.7 123.4 ± 6.9(5) 81.5 ± 5.0(5) 83.3 ± 4.5(4) 51.5 ± 2.9(4) 115.3-129.9 75.8-88.5 76.7-86.4 47.4-53.6 5 9°44' 127.4(2) 82.7 (2) 85.0(2) 52.6 (2) 125.8-129.0 82.0-83.3 83.8-86.1 51.6-53.6 6 io°or 129.9 88.5 86.4 53.5 7 10°04' 115.3 77.8 76.7 47.4 ? 116.8 75.8 124.0 ± 1.6(3) 82.8 ± 2.1 (3) 82.6 ± 2.9 (3) 48.0 ± 1.7(3) 122.3-125.5 81.2-85.1 80.1-85.8 46.2^9.4 9" 1 1°5 1 ' 122.3 82.0 80.1 48.5 OI6 11°51' 124.9(2) 83.2 (2) 83.9 (2) 47.8 (2) 124.2-125.5 81.2-85.1 82.0-85.8 46.2^*9.4 128.8 ± 5.1 (4) 82.3 ± 3.5 (4) 86.7 ± 1.8(4) 51.1 ± 3.2(4) 125.4-136.4 77.2-85.1 84.8-89.2 46.7-54.2 13 13°02' 127.2 77.2 84.8 52.6 14 13°07' 126.3 83.8 86.8 50.9 17 13°22' 136.4 85.1 89.2 54.2 ? 125.4 83.0 86.1 46.7 129.5 ± 5.7(12) 81.3 ± 3.6(12) 87.7 ± 3.0(7) 53.0 ± 2.8 (7) 121.6-139.1 74.8-88.6 84.3-92.2 48.9-55.8 26 13°53' 124.3 77.8 33 14°08' 132.9 80.1 85.4 54.2 35 14°21' 122.2(2) 121.6-122.7 80.4 44 17°00' 137.0 85.9 90.1 55.8 47 17°20' 133.2 82.2 50 17°35' 128.6 ± 3.0(3) 80.2 ± 0.9 (3) 85.5 ± 1.3(3) 50.7 ± 2.6(3) 126.4-132.0 79.1-80.9 84.3-86.9 48.9-53.6 52 18°31' 132.2 ± 7.1 (3) 82.0 ± 6.9(3) 91.1 (2) 54.4 (2) 125.0-139.1 74.8-88.6 89.9-92.2 52.9-55.8 7 82.9 58 1 1°47' 125.7 ± 2.8(3) 82.4 ± 1.9(4) 83.8 ± 1.7(3) 51.3 ± 1.7(4) 122.7-128.3 79.5-83.5 81.9-84.9 49.8-53.4 59 U°33' 123.9 82.1 83.7 49.1 60 11°33' 120.4(2) 82.0 (2) 81.9(2) 47.7 (2) 119.9-120.9 81.5-82.5 81.8-81.9 46.9^48.4 FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 15 Table 5. Continued. Loc. Zygomatic Postrostral Island no. 2 Nlat. Greatest length breadth length Rostral length Leyte 128.8 ± 4.1 (4) 82.2 ± 1.6(4) 85.8 ± 2.1 (4) 51.2 ± 3.2(4) 123.6-133.6 80.6-83.8 83.0-88.2 47.2-54.1 62 11°05' 123.6 80.6 83.0 47.2 63 10°46' 128.7 83.8 85.7 50.0 64 10°40' 133.6 83.3 88.2 54.1 ? 129.4 81.2 86.3 53.4 Bohol . '.. 125.6 ± 2.4(3) 80.5 ± 2.4 (3) 83.5 ± 1.9(3) 51.1 ± 0.9(3) 123.3-128.1 79.1-83.2 82.0-85.7 50.2-51.9 66 9°48' 125.5 79.1 82.8 51.9 65 9°45' 123.3 79.1 82.0 50.2 ? 128.1 83.2 85.7 51.2 Negros 121.4 ± 3.6(15) 83.1 ± 2.3(15) 81.8 ± 1.7(15) 47.8 ± 2.2(15) 116.6-129.6 79.6-87.8 79.6-84.9 44.0-52.4 80 9°39' 129.6 84.8 84.9 52.4 71 9°30' 116.6 81.1 79.6 44.7 72 9°30' 119.2 82.9 81.2 46.3 70 3 9°26' 120.8(2) 81.8(2) 81.3(2) 48;0 (2) 120.0-121.6 79.6-84.0 81.0-81.5 46.7-49.2 7i 18 9°25' 122.9(2) 84.2 (2) 83.6 (2) 48.3 (2) 121.0-124.7 83.4-84.9 82.8-84.4 48.0-^8.5 70" 9°24' 127.2 85.7 84.3 49.9 78 4 9°23' 122.2(2) 82.6 (2) 80.9 (2) 48.3 (2) 121.6-122.8 80.8-84.4 80.7-81.1 48.2^8.4 73 20 9°23' 118.8 83.4 80.0 45.7 74 9° 18' 121.1 81.4 81.3 49.9 78 2> 9°18' 120.6 87.8 83.2 47.8 . . .7 9°15' 116.6 81.6 79.6 44.0 77 9°12' 119.9 80.0 82.0 47.3 Mindanao 123.6 ± 4.2(26) 82.3 ± 2.3 (26) 82.9 ± 2.7 (25) 48.7 ± 2.8 (25) 118.5-132.9 77.8-85.9 78.9-90.8 42.5-54.4 82 9°45' 118.8 84.8 no* 8°31' 118.6 82.6 81.0 46.8 7/0" 8°31' 132.9 84.6 90.8 51.5 709 8°20' 122.0 ± 4.0(5) 80.5 ± 1.7(5) 82.3 ± 2.7 (5) 46.6 ± 3.3 (5) 118.5-127.2 78.3-82.4 78.9-85.3 42.5-51.0 108 8° 10' 124.7 ± 3.6(4) 83.0 ± 0.7 (4) 83.0 ± 2.3 (4) 49.8 ± 1.3(4) 121.5-128.4 82.4-83.9 81.0-85.5 48.2-51.4 106 8°04' 121.9 ± 2.7(4) 81.2 ± 2.1 (4) 82.3 ± 2.0 (4) 47.1 ± 2.5(4) 120.0-125.9 78.1-82.7 79.9-84.7 45.0-50.5 107 8°03' 128.7 84.2 82.1 54.4 105 7°53' 121.1 (2) 80.5 (2) 81.2(2) 47.9 (2) 120.5-121.6 80.3-80.7 80.5-81.9 47.1-48.6 94 7°06' 124.3(2) 84.2 (2) 82.8 (2) 49.3 (2) 123.4-125.1 82.8-85.5 82.3-83.3 47.5-51.0 95 6°57' 130.1 84.9 86.5 51.4 91 6°56' 127.8(2) 85.5 (2) 85.8 (2) 50.9 (2) 125.1-130.5 85.1-85.9 85.3-86.2 48.9-52.9 102 6°52' 121.0 81.4 80.0 48.1 97 6°49' 120.5 77.8 80.9 48.3 Cagayan Sulu 121 7°01' 109.7 77.5 75.0 43.9 Basilan 22 125 6°35' 117.4 81.3 80.1 44.7 Jolo 126 5°58' 125.6 89.9 87.2 47.4 Unknown 120.5 ± 4.8(6) 114.9-128.3 76.3 ± 3.4 (6) 71.5-79.6 All ... 124.0 ± 5.4(89) 81.8 ± 3.2(90) 83.4 ± 3.2 (76) 49.2 ± 3.1 (77) 109.7-139.1 71.5-89.9 75.0-92.2 42.5-55.8 Sabah Borneo 5°49" 1 16.4 ± 4.0 (20) 80.0 ± 3.6 (20) 79.8 ± 2.7 (20) 44.2 ± 3.0 (20) 110.7-123.4 74.3-87.9 75.7-85.7 38.2^17.6 16 FIELDIANA: ZOOLOGY Table 6. _^ Sample areas' Zamboanga; San Ramon (7/7, 775) Katipunan Munic. (7 10-112) Mt. Malindang area; Bucong (107-109, 113) Pan tar; Lake Lanao (705, 706) Bugusan; Bumngkot; Maculi Pt. (101-103) Mt. Apo area; Caburan; Kamansi (91. 94, 95, 97, 99) Mat. E long. teeth frequen 7°00' 122°00' 8 0.12 8°30' 123°15' 74 0.41 8°00' 123°30' 76 0.39 8°00' 124°15' 32 0.69 6°45' 124°15' 14 0.50 7°00' 125°15' 38 0.42 Listed in west-east order. Locality numbers in parentheses (see fig. 1). ly distribution of births in the Tanay colony of M. fascicularis generally resembles that in M. fascicu- laris in West Malaysia, where births peak in June (Kavanagh & Laursen, 1984, p. 23; reference pro- vided by anonymous reviewer). The birth peak in May in the Tanay colony also is close to the mid- June birth season of the fruit bat Haplonycteris fischeri studied in northern Luzon, southern Ne- gros, and southern Bohol by Heideman (1988, p. 586); in fruit bat populations studied in Biliran and central Leyte, the birth season occurs later, in July or August. Copulations in the Tanay colony peak during the period October-March (R. G. Re- suello, siconbrec, pers. comm., 4 Aug. 1989), which accords with the 5. 5-month gestation period in M. fascicularis (Ardilo, 1976, p. 216). In 1987, the annual reproductive rate in the Tanay colony was 0.76 surviving weaned young per breeding female (Hobbs, 1989, p. 64). In nature, females enter the breeding population Table 9. Blood-protein allele frequencies at polymorphic loci in Philippine and non-Philippine M. fascicularis. For references and details, see Fooden and Lanyon (1989, pp. 216, 222). Allele frequencies (%) and sample sizes (in parentheses) IvOCUS Philip- Philip- pines: Abbrevia- pines: island Other M. tion Allele Mindanao unknown Sumatra' fascicularis' Alb A 100(58) 100(21) 89 (222) 98 (734) CA-II A 62(16) 59(133) 2 B 38 41 HbA 1 50 (59) 50(21) 58 (222) 72(1,077) 2 50 50 42 24 IDH 1 98(21) 89 (222) 58(516) 2 2 10 42 PGD A 100(78) 100(21) 89(218) 95(912) PGM-I 1 90(21) 98(221) 96(481) 4 10 3 PHI 1 57 (30) 81 (21) 99 (223) 99 (648) 2 2 5 43 17 1 < 1 PI A 2(54) 0(73) 1 (291) 1 (1,034) B 93 96 80 60 C 5 4 19 38 TBPA F 94(58) 96 (73) 90 (274) 90(108) S 6 4 10 10 Tf D 100(78) 100(73) 59 (244) 50(913) Albumin, plasma Carbonic anhydrase-II, cell Hemoglobin-alpha, cell Isocitrate dehydrogenase, cell 6-phosphogluconate dehydrogenase, cell Phosphoglucomutase-I, cell Phosphohexose isomerase, cell Protease inhibitor, plasma Thyroxine-binding prealbumin, plasma Transferring, plasma 1 Table excludes minor alleles present in 2 Includes unlegalized sample (N = 67; Lanyon (1989, table IV). non-Philippine M. fascicularis but absent in Philippine M. fascicularis. 0.75 A, 0.25 B) inadvertently omitted from data table in Fooden and FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 19 Table 10. Habitat observations of Philippine M. fascicular is. Observer Year Island Reference Seashore J. B. Steere 1874 Basilan, Luzon, Mindanao, Palawan Giinther, [1877], p. 735 P. C. Frier et al. 1906 Palawan Mearns ms. (usnm) E. A. Mearns 1906 Tawitawi Mearns ms. (usnm) Sand Ridges E. A. Mearns 1907 Luzon Swamp Forest Mearns ms. (usnm) E. A. Mearns 1903 Jolo Mearns ms. (usnm) E. A. Mearns 1904 Cagayan Sulu Riverine Forest Mearns ms. (usnm) E. A. Mearns 1906 Samar Mearns ms. (usnm) H. Hoogstraal et al. 1946 Luzon fmnh 62273-62274, notes D. S. Rabor and F. Werner 1947 Palawan Cultivated Field fmnh 62901, note E. A. Mearns 1906 Basilan Mearns ms. (usnm) H. Hoogstraal 1947 Mindanao fmnh 56995, note E. A. Rickart et al. 1987 Maripipi Rickart et al., in prep. Mr. Siloy 1987 Leyte Open Grassland L. R. Heaney (fmnh), field notes E. A. Mearns 1904 Mindanao Second-Growth Forest Mearns ms. (usnm) P. Afionuevo 1947 Culion pnm 1 196, note D. S. Rabor 1947 Culion fmnh 62908, note D. S. Rabor 1947 Palawan fmnh 62914, note H. T. Wright et al. 1947 Palawan fmnh 62905, 62913, notes E. A. Rickart et al. 1987 Primary Rain Forest, Edge Biliran Primary Rain Forest E. v. Martens 1861 Luzon, Mindanao E. A. Mearns 1903 Mindanao M. Celestino 1946 Mindanao, 3 localities H. Hoogstraal et al. 1946 Luzon D. S. Rabor 1947 Busuanga H. Hoogstraal 1947 Mindanao L. R. Heaney 1984 Leyte E. A. Rickart et al. 1987 Maripipi Montane Forest (850 m) E. A. Rickart et al. 1987 Biliran Mossy Forest (2000 m) H. Hoogstraal 1946 Mindanao Oak -pine Montane Forest (2300 J. Whitehead 1895 Luzon Rickart et al., in prep. Martens, 1876, p. 193 usnm 123450, note fmnh 56435-56437, notes fmnh 62275-62276, notes fmnh 62906, note fmnh 56495, note ummz 161311, note (pers. comm. 14 Mar. 1990) Rickart et al., in prep. Rickart et al., in prep. Hoogstraal, 1951, p. 44 m) Thomas, 1898, p. 381 before they achieve dental maturity. A wild-col- lected female (usnm 477849, Apr. 1 962, field note) with third molars unerupted (age < 4.5 years; Hurme, 1960, p. 797; Bowen & Koch, 1970, p. 122) was already pregnant, and another (usnm 144675, Nov. 1906, field note) with third molars erupting (age ca. 6 years) evidently had already nursed an infant. 20 FIELDIANA: ZOOLOGY Table 1 1 . Monthly distribution of births in captive colony of Philippine M. fascicularis maintained outdoors at Tanay, near Manila (Hobbs, 1 989, p. 67). 1985 1986 1987' Total 1 Month N % N % N % N % Jan. 2 0.6 30 5.7 56 5.8 88 5.0 Feb. 6 1.8 26 4.9 52 5.4 84 4.8 Mar. 28 8.2 54 10.3 62 6.4 144 8.2 Apr. 26 7.6 62 11.8 116 12.0 204 11.6 May 60 17.6 88 16.7 154 15.9 302 17.1 June 41 12.0 72 13.7 138 14.3 251 14.2 July 42 12.3 74 14.0 116 12.0 232 13.2 Aug. 28 8.2 36 6.8 75 7.8 139 7.9 Sep. 54 15.9 18 3.4 100 10.4 172 9.7 Oct. 32 9.4 17 3.2 97 10.0 146 8.3 Nov. 12 3.5 22 4.2 nd 2 Dec. 10 2.9 28 5.3 nd 2 Totals 341 100.0 527 100.0 966 100.0 1,762 100.0 Ten-month period. No data. Systematics Subspecific Taxonomy Geographic variation within or between Phil- ippine M. fascicularis and Sabah M . fascicularis is apparent in (1) dorsal pelage color, (2) head and body length, (3) relative tail length, (4) greatest skull length, and (5) incidence of accessory ento- styles (see above). Of these, dorsal pelage color traditionally has served as the primary basis for recognition of subspecies in M. fascicularis and, as discussed below, continues to provide the most useful criterion for subspecific allocation of Phil- ippine and Sabah populations of this species. Head and body length and greatest skull length average greater in Philippine M. fascicularis than in Sabah M. fascicularis, and, conversely, relative tail length averages greater in Sabah M. fascicularis than in Philippine M. fascicularis (tables 3, 5). However, overlap between Philippine and Sabah samples is substantial in all three of these char- acters. Greatest skull length varies clinally within Philippine M. fascicularis and probably also be- tween Philippine M. fascicularis and Sabah M. fascicularis (fig. 7). Similar clinal variation within and between Philippine and Sabah M . fascicularis also may occur in head and body length and rel- ative tail length. The incidence of accessory entostyles is geo- graphically erratic in Philippine and Sabah M. fas- cicularis (table 7). The incidence of entostyles is relatively high in Sabah, Mindanao, and Mindoro and generally is low elsewhere in the Philippines. Dorsal pelage color in M. fascicularis is distinc- tively dark in most islands of the Philippine ar- chipelago (fig. 3). The major known exceptions are the south-central islands of Cagayan Sulu, Tawi- tawi, Basilan, and western Mindanao. In these is- lands, dorsal pelage color is pale to intermediate, matching in saturation that in nearby Sabah M. fascicularis. In central and southeastern Mindanao and in southern Negros, dorsal pelage color varies from pale to intermediate to dark. Variation of dorsal pelage color in Philippine M. fascicularis is at least partly independent of variation of skull length; in southern Philippine samples, skull length is small but pelage color may be either pale or dark (figs. 3, 7). Similarly, vari- ation of pelage color also is partly independent of variation of incidence of accessory entostyles; pel- age color is pale in Sabah and dark in Mindoro, but entostyle incidence is relatively high in both of these places (table 7). Variation of pelage color probably also is at least partly independent of vari- ation of head and body length and relative tail length. Taxonomically, pale to intermediate popula- tions of M. fascicularis in south-central Philip- pines are referable to the same subspecies as those in neighboring Sabah (fig. 9); assuming that these pale populations are inseparable from those in southern Sumatra, the name that applies to this pale subspecies is M '. fascicularis fascicularis Raf- FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 21 fles, [1821] (type locality, Bengkulu vicinity, southern Sumatra). Dark populations of Philip- pine M. fascicularis that are geographically con- tiguous constitute another subspecies, for which the name M. fascicularis philippinensis I. Geoffrey, [1843] (type locality, "Manile"), is available. The area of mixed populations of pale, intermediate, and dark individuals in central and southeastern Mindanao and southern Negros is geographically intermediate between the subspecific ranges of M. f. philippinensis and M. f. fascicularis as defined above and may be regarded as a subspecific contact zone. The current taxonomic practice (cf. Napier, 1 98 1 , p. 1 3) of uniting pale and dark populations of Philippine M. fascicularis in a single taxon that excludes pale Sabah M. fascicularis is untenable. For most nonvolant mammals, the faunal re- lationship between Borneo and Palawan is closer than that between Borneo and Mindanao (Heaney, 1986, p. 141). The reverse is true for subspecies of M. fascicularis (fig. 9). A single specimen collected in Busuanga Island, in the Palawan-Mindoro chain, and another col- lected in Balut Island, south of Mindanao, are problematical (fig. 3). The Busuanga specimen is intermediate in dorsal pelage color saturation and may be assigned either to M. fascicularis or to the contact-zone area; both of these assignments are geographically incongruous. The Balut specimen is dark, which implies either that this island is part of the subspecific contact zone or that it is a dis- junct extension of the range of M. f philippinensis. Subspecies Accounts Macaca fascicularis fascicularis (Raffles, [1821], p. 246), part, Philippine Islands only Macaca suluana: Alcasid, [1970], p. 24— listed. Macaca sulvensis: Chiarelli, 1972, p. 213— species name incorrectly spelled. Cynomolgus cagayanus Mearns, 1905, p. 431— ho- lotype, usnm 125325, adult male, skin and skull, collected 25 Feb. 1904 by E. A. Mearns, Cagayan Sulu. Napier, 1981, p. 13— a synonym of Macaca fascicularis philippinensis. Macaca cagayana: Lyon and Osgood, 1909, p. 283 — holotype cataloged. Macaca irus cagayana: Hill, 1974, p. 525— external and cranial characters ex Mearns, 1905. M[acaca] p[hilippinensis] cagayanus: Rabor, 1986, p. 138— distribution, Cagayan Sulu. Pithecus cagayanus: Hollister, 1912, p. 36— listed. Type— None preserved. The name Simia fas- cicularis Raffles, [1821], is based on specimens collected between 1818 and 1820 in Sumatra by T. S. Raffles. Three syntypes formerly were pre- served in the collection of the Zoological Society of London (Waterhouse, 1838, p. 7). Type Locality— Indonesia: Sumatra, near Bengkulu (Hill, 1974, p. 477). Distribution— The known range of Philippine Macaca f fascicularis is restricted to the south- central part of the Philippine archipelago, includ- ing Cagayan Sulu, the Sulu archipelago (Tawitawi, Basilan), and western Mindanao (west of 123°30'E) (figs. 3, 9). A single specimen with golden brown agouti pelage also has been collected in Busuanga, which is disjunct from the range given above; the zoogeographic significance of this specimen is un- clear (see above, Subspecific Taxonomy). Diagnosis— Dorsal pelage color in Philippine M. f fascicularis is pale yellowish brown agouti (olivaceous) to golden brown agouti and is vari- ably erythristic. Pelage color in Philippine M. f fascicularis is similar in saturation to that in Sabah M. f fascicularis, but averages somewhat more erythristic. Macacus philippensis: Murray, 1885, p. 665 (not Rei- chenbach, 1862)— Mindanao: Mount Pulunbato. Macacus philippinensis: Steere, 1 890, p. 28 (part, not I. Geoffroy, [1843])— Basilan. Bourns and Worces- ter, 1894, p. 61 — Basilan, Jolo, Tawitawi. Cynomolgus suluensis Mearns, 1905, p. 430— holo- type, usnm 125324, adult male, skull only, collected 16 Nov. 1903 by E. A. Mearns, Jolo: foot of Crater Lake Mountain. Hill, 1974, p. 522— a synonym of Macaca irus philippinensis. Macaca suluensis: Lyon and Osgood, 1909, p. 284— holotype cataloged. M[acaca] p[hilippinensis] suluensis: Rabor, 1986, p. 138— distribution, Sulu archipelago. Pithecus sulensis: Hollister, 1912, p. 37— listed. Elliot, [1913], p. 252— specific status indeterminate. Macaca fascicularis philippinensis I. Geoffroy, [1843] Macacus cynomolgus: Waterhouse, 1838, p. 8— part, specimen collected in Luzon. Cuming, [1841], p. 33— color variation. Gray, 1849, p. 4— part, Phil- ippines included in geographic range. Thomas, 1 898, p. 381— Luzon: Bark. C[ynamolgus] cynamolgus: Elera, 1895, p. 2— part, Philippines included in geographic range. Elera, 1915, p. 32— part, dorsal pelage color. Macacus carbonarius: Waterhouse, 1839, p. 2— part, revised identification of Macacus cynomolgus: Wa- terhouse, 1838. 22 FIELDIANA: ZOOLOGY Macacus philippinensis I. Geoffrey, [1843], p. 568, pi. 5 (animal)— name based on captive albino, pro- posed provisionally. I. Geoffrey, 1851, p. 29— type history. Gervais, 1854, p. 88— behavior of holotype in captivity. Schlegel, 1876, p. 104— a synonym of Cercocebus cynamolgus. Macacus philippensis: Anderson, 1879, p. 74— species name incorrectly spelled. Macaca philippinensis: Kellogg, 1945, p. 119— listed. Macaca philippensis: Kuntz, 1969, p. 217— species name incorrectly spelled. Macaca phylippinensis: Chiarelli, 1972, p. 213— spe- cies name incorrectly spelled. Pith[ecus] (Mac[acus]) Philippinensis: Dahlbom, 1856, p. 1 18— external characters. C\ynamolgus] philippensis: Reichenbach, 1862, p. 134— species name incorrectly spelled. C[ynamolgus] philippinensis: Elera, 1 895, p. 2— listed. Cynomolgus philippinensis: Mearns, 1905, p. 426— external and cranial characters. Zati philippensis: Reichenbach, 1862, p. 135— pre- ferred binomial. Cercocebus Philippinus: Martens, 1876, p. 207— spe- cies name incorrectly spelled. [Silenus] philippinensis: Stiles and Nolan, 1929, p. 532— new generic combination. Macacus cynamolgos . . . var. Philippinus: Martens, 1876, pp. 193, 362 — Luzon: Los Banos. Macaca philippinensis philippinensis: Sanborn, 1952, p. 1 13— taxonomic comparison. Macaca irus philippinensis: Hill, 1974, p. 522— tax- onomy. [Macaca fascicularis] philippinensis: Napier and Na- pier, 1967, p. 404 — listed. Macacus palpebrosus I. Geoffrey, 1851, pp. 88, 93 — syntypes, four captives presented by M. Dugast to menagerie of mnhn about 1851, obtained in Luzon: "forets de Manille," apparently not preserved (cf. Rode, 1938, p. 222); name proposed provisionally. I. Geoffrey, [1852], p. 543— type history. Schlegel, 1876, p. 104— a synonym of Cercocebus cynamol- gos. Martens, 1876, p. 206— a synonym of Macacus cynamolgos var. Philippinus. Mearns, 1905, p. 426— a synonym of Cynomolgus philippinensis. I[nuus] palpebrosus: Wagner, 1855, p. 54— specific status questionable. C\ynamolgus] palpebrosus: Reichenbach, 1862, p. 137— external characters ex I. Geoffrey, 1851. Macacus fur Slack, 1867, p. 36, pi. 1 (animal; skull)— holotype, formerly preserved in Academy of Nat- ural Sciences, Philadelphia (ansp), no. 1254, adult male, skin and skull, obtained "some years since" in Paris by J. H. Slack ( 1 867), said to inhabit Luzon; holotype now lost (J. E. Cadle, ansp, letter, 14 Nov. 1989; cf. Koopman, 1976, p. 7). Schlegel, 1876, p. 104— a synonym of Cercocebus cynamolgos. Hol- lister, 1912, p. 37— a synonym of Pithecus syrichta. C\ynamolgus] fur: Elera, 1895, p. 3 — listed. Macacus cynomolgus . . . var. Cumingii Gray, 1870, p. 30— holotype, bm(nh) 1859.7.9.5, juvenile fe- male, skin and skull, obtained (presumably in 1 859) from J. Verreaux, "Hob. Philippine Islands." Schle- gel, 1876, p. 104— allocated to Philippine "conspe- cies." Hollister, 1 9 1 2, p. 37— a synonym of Pithecus syrichta. C\ynamolgus\ var. cumingii: Elera, 1895, p. 3 — listed. [ Cynomolgus] fascicularis: Trouessart, 1904, p. 16— part, Philippines included in geographic range. Macacus fascicularis: Elliot, 1907, p. 567 — Panay. M[acaca] syrichta: Thomas, 1 9 1 1 , p. 129 (not Lin- naeus, 1 758)— misidentification of holotype of Sim- ia syrichta Linnaeus, 1758, p. 29 (a tarsier; cf. Ca- brera, 1 923, p. 90). Elliot, [ 1 9 1 3], p. 249-a synonym of Pithecus philippinensis. Pithecus syrichta: Hollister, 19 1 2, p. 37 (not Linnaeus, 1758)— distribution: Luzon, Leyte, Mindoro, Ne- gros, Palawan, Samar. Hollister, 1913, pp. 328, 330— invalid designation of neotype. S[ ilenus] syrichta: Stiles and Nolan, 1929, p. 537 (not Linnaeus, 1758)— nomenclature. Pithecus mindorus Hollister, 1913, p. 328, pis. 27-28 (skull)— holotype, usnm 144674, adult male, skin and skull, collected 1-8 Nov. 1906 by E. A. Mearns, Mindoro: Alag River. Hill, 1974, p. 522— a syn- onym of Macaca irus philippinensis. Macaca mindorus: Poole and Schantz, 1942, p. 244— holotype cataloged. M\acaca] p[hilippinensis] mindorus: Rabor, 1986, p. 1 3 8 — distribution , M i ndoro. Macaca mindora: Lawrence, 1939, p. 63— taxonomic comparison. [Macaca fascicularis] mindora: Napier and Napier, 1967, p. 404-listed. Macaca irus: Pocock, 1939, p. 78 — part, Philippine Islands included in distribution. Type— The holotype of Macacus philippinensis I. Geoffrey, [ 1 843], is mnhn 373/265, albino adult male, mounted skin only (omitted from type cat- alog published by Rode, 1938, p. 222). This mon- key was obtained alive in Manila by A. Chenest and presented by him to the menagerie of mnhn on 6 Aug. 1841; it died on 29 Aug. 1842 (cf. I. Geoffroy, 1851, p. 29). Type Locality— Philippine Islands: probably Luzon. Concerning the provenance of the holo- type, I. Geoffroy [1843, p. 570] reports: "M. Adolphe Chenest, qui a fait don au Museum de Singe precieux, l'a acquis a Manille, et il le croit originaire de cette tie." Distribution— The known range of M. f. phil- ippinensis includes Balabac, Palawan, Culion, Mindoro, Luzon, Samar, Leyte, and northeastern Mindanao (north of ca. 9°00'N) (figs. 3, 9). The subspecific range probably also includes other is- lands in central Philippines, north of ca. 10°00'N lat. A single specimen with dark dorsal pelage was collected in Balut, south of Mindanao; the zoo- geographic significance of this specimen is ambig- uous (see above, Subspecific Taxonomy). FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 23 Diagnosis— Dorsal pelage color in M. f philip- pinensis is dark brown agouti, variably erythristic. Taxonomic Note— Macacus cristatus Gray, 1870 (p. 30) has been treated as a synonym of Macacus philippinensis I. Geoffroy, [ 1 843], by El- liot ([191 3], p. 249) and subsequent authors. How- ever, this name may not be based on a Philippine macaque. The holotype of Macacus cristatus is bm(nh) 1858.4.28.9, an albinistic late juvenile male, skin and skull, obtained in 1858 from the collection of Th. G. van Lidth de Jeude, Utrecht (cf. Thomas, 1906, p. 43; Napier, 1981, p. 20). No information was available to Gray concerning the provenance of the specimen ("Hab. ?"). In Gray's description of cristatus, he compares ex- ternal characters of the albinistic holotype with those of the albino holotype of Macacus philip- pinensis I. Geoffroy; this may be the source of Elliot's assumption that the holotype of cristatus originated in the Philippines (cf. Anderson, 1879, p. 76). Independent retrospective inquiry into the geographic origin and subspecific identity of the holotype of cristatus is impeded by the immaturity and deficient pigmentation of the specimen. Macaca fascicularis fascicularis/ Macaca fascicularis philippinensis Contact Zone Macaca philippinensis: Steere, 1 890, p. 28 (part, not I. Geoffroy, [1843])— Mindanao, Negros. Cynomolgus mindanensis Mearns, 1905, p. 428— ho- lotype, uskm 123450, adult male, skin and skull, collected 26 Aug. 1 903 by E. A. Mearns, Mindanao: Pantar. Elliot, [1913], p. 250-a synonym of Pithe- cus philippinensis. Hill, 1974, p. 522— a synonym of Macaca irus philippinensis. Macaca mindanensis: Lyon and Osgood, 1909, p. 284— holotype cataloged. Pithecus mindanensis: Hollister, 1913, p. 329— tax- onomy. Pithecus mindanensis mindanensis: Hollister, 1912, p. 37— listed. Macaca philippinensis mindanensis: Sanborn, 1952, p. 1 13— taxonomic comparison. Rabor, 1986, p. 138— distribution, Panay, Samar, Leyte, Bohol, Cebu, Negros, Mindanao, Basilan. [Macaca fascicularis] mindanensis: Napier and Na- pier, 1967, p. 404— listed. Cynomolgus mindanensis apoensis Mearns, 1905, p. 429— holotype, usnm 125321, adult female, skin and skull, collected 25 June 1904 by E. A. Mearns, Mindanao: Mount Apo. Hollister, 1913, p. 329 — subspecies regarded as unrecognizable. Macaca mindanensis apoensis: Lyon and Osgood, 1909, p. 283-holotype cataloged. Pithecus mindanensis apoensis: Hollister, 1 9 1 2, p. 37— listed. Pithecus philippinensis apoensis: Elliot, [1913], p. 250— subspecies recognized provisionally. Distribution— Mixed populations that include individuals with pale, intermediate, and dark dor- sal pelage are documented in southern Negros (south of 9°40'N) and in central and southeastern Mindanao (south of 8°10'N, east of 1 23°40'E) (figs. 3,9). Diagnosis— Dorsal pelage color in the M.f. fas- cicularis/ M. f. philippinensis contact zone varies from pale yellowish brown agouti to golden brown agouti to dark brown agouti and is variably ery- thristic. Zoogeography and Evolution The Philippine archipelago is separated from Borneo and other islands on the Sunda Shelf by deep trenches (minimum depth 145 m, Balabac- Palawan chain; 290 m, Sulu archipelago; Heaney, 1985b, p. 54). Most Philippine nonvolant mam- mals probably dispersed to the archipelago from Borneo during periods of Pleistocene continental glaciation when worldwide sea-level regression re- duced the width of intervening water gaps (Hea- ney, 1986, p. 152;VanCouvering&Kukla, 1988b, p. 507). During the most recent glacial maximum (ca. 18,000 years B.P.), sea level was about 120 m lower than at present; during the immediately pre- ceding glacial maximum (ca. 160,000 years B.P.), sea level was about 1 60 m lower than at present (Heaney, 1985a, p. 129; Van Couvering & Kukla, 1988a, p. 462). The most conspicuous zoogeographic pattern in Philippine M. fascicularis is the restricted distri- bution of pale to intermediate M.f. fascicularis in south-central islands (Cagayan Sulu, Sulu archi- pelago, western Mindanao), the relatively broad distribution of dark M.f. philippinensis in western, northern, and eastern islands (Balabac, Palawan, Culion, Mindoro, Luzon, Samar, Leyte, north- eastern Mindanao, and presumably others), and the intervening distribution of pale to intermedi- ate to dark contact-zone populations (eastern and southern Mindanao, southern Negros) (figs. 3, 9). Pale to intermediate M.f. fascicularis also inhabits Sabah. Either of two alternative hypotheses can account for this pattern. A "one-wave" hypothesis postulates that M. fascicularis dispersed from Bor- neo to the Philippines once, becoming progres- 24 FIELDIANA: ZOOLOGY 118° 122" 126* - 14° 100 200 300 km 18* nsis ?&-■- M. f. fascicularis 9ps <&..-'' 122' 126" — 14*' 10°' Fig. 9. Known limits of distribution of Philippine M. f. fascicularis, M. f. philippinensis, and contact-zone pop- ulations (cf. fig. 3). FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES >bars>-^ o° Fig. 10. Map of Southeast Asia showing limits of distribution of pale to intermediate subspecies of M. fascicular is, location of deep-water insular habitats of dark subspecies of M. fascicularis, and depth of intervening channels. (References: Chasen, 1940, p. 68; Kellogg, 1944, p. 76; Defense Mapping Agency Hydrographic Center, 1973, 1975, 1979; Fooden, 1980, p. 5; Heaney, 1985b, p. 54; Khan, 1985, p. 29; above, figs. 3, 9.) sively darker as it spread through the archipelago, thus producing the present pattern of color vari- ation. A "two-wave" hypothesis, discussed in greater detail below, postulates that M. fascicularis dispersed to the Philippines twice, with the first wave now represented by dark M. f. philippinensis and the second wave now represented by pale to intermediate M. f fascicularis. The species-wide geographic distribution of pale to intermediate and dark populations of M. fascicularis tends to favor the two-wave hypothesis. Outside of the Philippines, pale to intermediate populations of M \ fascicularis are broadly distrib- uted, from southernmost Bangladesh, through the Indochinese and Malay peninsulas, Sumatra, Java, Borneo, and numerous adjacent smaller islands, to Timor in the Lesser Sunda Islands (fig. 10; Cha- sen, 1940, p. 68; Kellogg, 1944, p. 76). Non-Phil- ippine dark populations of M. fascicularis are re- stricted to three island groups or islands around the periphery of the specific range: (1) Nicobar Islands, NW of Sumatra; (2) Pulau Simeulue and Pulau Lasia, W of Sumatra; and (3) Pulau Mara- tua, E of Borneo (fig. 1 1). Most islands inhabited by pale to intermediate populations of M. fascicu- laris are shallow- water islands on the Sunda Shelf; the only deep-water islands inhabited by pale to intermediate M. fascicularis are the south-central Philippine Islands and the Lesser Sunda Islands. By contrast, all four island groups or islands that are inhabited by dark populations of M. fascicu- laris, including M.f. philippinensis, are deep-water islands, separated from the Sunda Shelf by deep trenches (Nicobar Islands, > 1000 m; Pulau Si- 26 FIELDIANA: ZOOLOGY Luzon Maratua Simeulue Nicobar Fig. 1 1 . Dark non-Philippine M. fascicularis skins (usnm 1 97662, Pulau Maratua; usnm 1 1 4 1 68, Pulau Simeulue; usnm 1 1 1 797, Little Nicobar I.) compared with pale and dark Philippine M. fascicularis skins (usnm 1 44666, Basilan: E end; usnm 144676, Luzon: Nagpartian). Scale bar 10 cm. meulue and Pulau Lasia, > 200 m; Pulau Maratua, > 200 m). Dorsal pelage is pale to intermediate not only in most populations of M. fascicularis but also in Macaca mulatta, a close relative of M '. fascicularis (Fooden, 1980, p. 7). This suggests that pale to intermediate pelage is primitive in M . fascicularis and that dark pelage evolved independently in the four deep-water island populations, which pre- sumably have been isolated longer than pale to intermediate shallow-water island populations. These pale to intermediate shallow-water island populations of M. fascicularis probably were geo- graphically and genetically continuous during the most recent glacial maximum (ca. 18,000 years B.P.), and they probably have been isolated only since the subsequent postglacial rise of sea level. Macaca f philippinensis and the other three dark populations may have dispersed to their deep-wa- ter island habitats during the preceding glacial maximum (ca. 160,000 years B.P.) and may have been isolated since the corresponding interglacial rise of sea level. Complete land bridges to the deep- water islands would not have been required for M. fascicularis to reach them; this species swims well and often forages along the seashore, and therefore was a prime candidate for active or pas- sive overwater dispersal when the penultimate gla- cial sea-level depression (ca. 160 m) greatly nar- rowed the water gaps that separate deep-water islands from the Sunda Shelf. To recapitulate, the first wave of M. fascicularis is assumed to have reached the Philippines from Borneo during the penultimate glacial maximum; in isolation, during the subsequent interglacial, this population differentiated into dark M. f. philip- pinensis. The second wave of M. fascicularis reached the Philippines from Borneo during the most recent glacial maximum; during its much shorter period of postglacial isolation, this popu- lation has retained the pale to intermediate pelage of M. f. fascicularis. (The dispersal of M. fascicu- laris to the Lesser Sunda Islands may also have occurred during the most recent glacial maximum; this is suggested by the pale to intermediate pelage of Lesser Sunda populations.) The dispersal route FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 27 of the first wave may have been either by way of the Balabac-Palawan chain or by way of the Sulu archipelago, or by both. The dispersal route of the second wave evidently was by way of the Sulu archipelago. The slow loris, Nycticebus coucang, with one subspecies inhabiting both Borneo and the Sulu archipelago (Groves, 1971, p. 49; Musser & Heaney, 1985, p. 29), provides additional evi- dence of a relatively recent dispersal route for pri- mates from Borneo via the Sulu archipelago. Ca- gayan Sulu, the Sulu archipelago, and western Mindanao formerly may have been inhabited by dark first-wave populations of M. f. philippinensis that were displaced by pale to intermediate sec- ond-wave populations of M. f. fascicularis. The M.f.fascicularis/M. f. philippinensis contact zone in central and southeastern Mindanao and south- ern Negros may be the result of interbreeding be- tween dark first-wave populations and pale to in- termediate second-wave populations. A dark specimen collected in Balut Island, off southern Mindanao (fig. 3), may represent a population that is a geographic relict of the first wave. After becoming isolated in the Philippines, in addition to undergoing darkening of dorsal pelage, M . fascicularis also apparently increased in size— at least cranially— as it dispersed northward; this is a frequent pattern of geographic variation in macaque species (cf. Fooden, 1988, p. 8). The in- cidence of accessory entostyles apparently has de- creased independently in several islands following isolation of populations of M \ fascicularis; the high frequency of entostyles that presumably charac- terized the ancestral population in Borneo appar- ently has been retained in Mindanao and Mindoro (possibly also in Panay, Balut, and Basilan). Blood- protein loci apparently have tended to become fixed in isolated populations of Philippine M. fas- cicularis; additional localized blood-protein data from Bornean and Philippine M. fascicularis would provide a critical test of the two-wave hypothesis. In the absence of other species of macaques, Phil- ippine M. fascicularis apparently has successfully expanded its ecological range to include primary evergreen rain forest ("ecological release")- Pro- liferation of M '. fascicularis in the Philippines may have been a factor in the origin and evolution of the endemic Philippine eagle Pithecophaga jef- feryi, a known predator on these monkeys. Transport by humans of pet monkeys is fre- quently assumed to have been a major factor in the dispersal of M. fascicularis to and within the Philippine archipelago (Blyth, 1875, p. 8; White- head in Thomas, 1898, p. 381; Taylor, 1934, p. 340; Darlington, 1957, p. 504). No data are avail- able on which to base an estimate of the impact of human introductions on the distribution of Philippine macaques. However, as indicated above, the habits of this species are conducive to natural overwater dispersal. In any event, it seems highly unlikely that the broad coherent pattern of distri- bution in the archipelago of M. f fascicularis, M. f philippinensis, and contact-zone populations is essentially an artifact produced by the random transport of pets. Summary Dorsal pelage color in Philippine M. fascicularis is pale to intermediate in south-central islands of the archipelago— as in Sabah, northern Borneo— and dark in most other islands; in central and southeastern Mindanao and in southern Negros, dorsal pelage color varies from pale to interme- diate to dark. Available external measurements are inadequate to establish a pattern of geographic variation of size; greatest skull length, however, increases latitudinally from Sabah to Luzon. Blood- protein loci tend to be less variable in Philippine M. fascicularis than in non-Philippine M. fascicu- laris. Philippine M. fascicularis frequently occurs in primary evergreen rain forest, an uncommon habitat for non-Philippine M. fascicularis. Births in Philippine M. fascicularis probably peak in May. Taxonomically, pale to intermediate populations of Philippine macaques are referable to M. f fas- cicularis, which also is distributed widely outside of the Philippines; dark populations are referable to M. f philippinensis, which is endemic to the Philippines; mixed pale-intermediate-dark pop- ulations may be regarded as M. f fascicularis/ M. f philippinensis contact-zone populations. Zoo- geographic evidence suggests that ancestors of M. f philippinensis dispersed to the Philippine archi- pelago from Borneo during the penultimate glacial maximum (ca. 160,000 years B.P.) and that an- cestors of Philippine M. f fascicularis dispersed to the archipelago from Borneo during the most recent glacial maximum (ca. 18,000 years B.P.); contact-zone populations probably are a product of interbreeding between these two successive waves of macaque dispersal from Borneo to the Philippines. Acknowledgments I thank curators and staff members of institu- tions listed in the Introduction for generous per- 28 FIELDIANA: ZOOLOGY mission to study specimens in their custody and for assistance in conducting the research. For valu- able supplementary information, discussion, and help, I thank P. L. Alviola III (uplbzd), J. E. Cadle (Academy of Natural Sciences, Philadelphia, Pennsylvania), W. G. Dee (Protected Areas and Wildlife Bureau, Quezon City, Philippines), E. Delson (amnh), J. Erwin (Diagnon Corporation, Rockville, Maryland), P. Gonzales (pnm), C. O. Handley, Jr. (usnm), J. -J. Hwang (fmnh), P. D. Jenkins (bm(nh)), K. F. Koopman (amnh), F.-W. Lange (siconbrec), J. R. Leuterio (uplbcf), P. Lowther (fmnh), J. B. Nazareno (siconbrec), B. D. Patterson (fmnh), D. S. Rabor (uplbcf), R. G. Resuello (siconbrec), R. W. Thorington, Jr. (usnm), and W. C. Wozencraft (usnm). Special thanks are due L. R. Heaney (fmnh) for numerous helpful suggestions and for generously sharing his extensive knowledge of the Philippines and its mammalian fauna. Literature Cited Alcala,A. C 1976. Philippine Land Vertebrates: Field Biology. New Day Publishers, Quezon City, Philip- pines, 167 pp. Alcasid, G. L. [1970]. Checklist of Philippine Mam- mals. National Museum, Manila, 51 pp. [For date of publication, see Heaney and Rabor, 1 982.] Anderson, J. 1879. Anatomical and Zoological Re- searches: Comprising an Account of the Zoological Results of the Two Expeditions to Western Yunnan in 1868 and 1875, vol. 1. Bernard Quaritch, London, 985 pp. [For date of publication, see Corrigenda, be- tween pp. xii-xiii.] Andrews, R. C 1911. Around the world for the Mu- seum. American Museum Journal, 11: 21-24. Ardito, G. 1976. Check-list of the data on the ges- tation length of primates. Journal of Human Evolu- tion, 5: 213-222. Barbour, T. 1939. Collections from the Philippine Islands: Introduction. Bulletin of the Museum of Com- parative Zoology at Harvard College, 86: 5-27. Blasius, W. 1888. Die Vogel von Palawan. Ornis, 4: 300-320. Blyth, E. 1875. Catalogue of mammals and birds of Burma. Journal of the Asiatic Society of Bengal, 44(2), extra number: 1-167. Bourns, F. S., and D. C. Worcester. 1894. Prelim- inary notes on the birds and mammals collected by the Menage Scientific Expedition to the Philippine Is- lands. Minnesota Academy of Natural Sciences. Oc- casional Papers, 1(1): 1-64. Bowen, W. H, andG. Koch. 1970. Determination of age in monkeys (Macaca irus) on the basis of dental development. Laboratory Animals, 4: 1 13-123. Buzeta, M., and F. Bravo. 1850. Diccionario Geo- grafico, Estadistico, Historico de las Islas Filipinas. D. Jose C de la Pena, Madrid, 567 pp. Cabrera, A. 1923. On the identification of Simia sy- richta Linnaeus. Journal of Mammalogy, 4: 89-91. Cabrera, B. D. 1973. A review of some wild land mammals of Palawan, Republic of the Philippines. Experimental Animals (Jikken Dobutsu), Tokyo, 22(suppl.): 247-252. Census of the Philippines: 1 960; Population and Hous- ing, Report by Province. 1962-. Bureau of the Cen- sus and Statistics, Manila. Chasen, F. N. 1940. A handlist of Malaysian mam- mals. Bulletin of the Raffles Museum, 15: 1-209. Chiarelli, A. B. 1972. Taxonomic Atlas of Living Primates. Academic Press, London, 362 pp. Clemens, J. 1907. Notes from the Philippines. Condor, 9: 92-93. Cuming, H. 1 839. Letter from. Proceedings of the Zoo- logical Society of London, 1839: 93. . [1841]. Letter, relating to specimens sent to the Society. Proceedings of the Zoological Society of Lon- don, 1840: 33. [For date of publication, see Proceed- ings of the Zoological Society of London, 107(ser. A): 79(1937).] Dahlbom, A. G. 1856. Studia Zoologica . . . , vol. 1, fasc. 1. Berligianis, Lund, 240 pp. Dampier, W. 1697. A New Voyage Round the World. Argonaut Press, London (reprint, 1927), 376 pp. Darga, L. L., M. Goodman, M. L. Weiss, G. W. Moore, W. Prychodko, H. Dene, R. Tashian, and A. Koen. 1975. Molecular systematics and clinal variation in macaques, pp. 797-812. In Markert, C. L., ed.. Iso- zymes IV: Genetics and Evolution. Academic Press, New York. Darlington, P. J., Jr. 1 957. Zoogeography: The Geo- graphical Distribution of Animals. John Wiley and Sons. New York, 675 pp. Defense Mapping Agency Hydrographic Center. 1973. Sheet No. 72120, Borneo- East Coast: Pulau Maratua to Tandjung Mangkalihat, 2nd ed., revised. Washington, D.C. . 1975. SheetNo. 71006, Sumatera— Northwest Part: Tanjung Jambuair to Sinkil, 2nd ed. Washington, D.C. . 1979. Sheet No. 72035, Eastern Portion of Jawa, Including Madura, Bali and Lombok, 3rd ed. Washington, D.C. Djambatan N. V. 1 964. Atlas of South-East Asia. Mac- millan, London, [92] pp. Elera, Casto de. 1 895. Catalogo Sistematico de Toda la Fauna de Filipinas . . . , vol. 1. Colegio de Santo Tomas, Manila, 701 pp. . 1915. Contribucion a la Fauna Filipina. Cole- gio de Santo Tomas, Manila, 284 + xi pp. Elliot, D. G. 1 907. A catalogue of the collection of mammals in the Field Columbian Museum. Field Co- lumbian Museum Publication No. 1 1 5, Zoological Se- ries, 8: 1-694. . [1913]. A Review of the Primates, vol. 2. American Museum of Natural History, New York, 382 + xxvi pp. [For date of publication, see Zoological Record: Mammals, 1913, p. 9.] Famatiga, E. G 1973. Leptospirosis in Philippine monkeys. Southeast Asian Journal of Tropical Med- icine and Public Health, 4: 316-318. FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 29 Fooden,J. 1969. Taxonomy and evolution of the mon- keys of Celebes (Primates: Cercopithecidae). Biblio- theca Primatologica, 10: 1-148. . 1980. Classification and distribution of living macaques (Macaca Lacepede, 1 799), pp. 1-9. In Lind- burg, D. G., ed., The Macaques: Studies in Ecology, Behavior and Evolution. Van Nostrand Reinhold, New York. -. 1982. Ecogeographic segregation of macaque species. Primates, 23: 574-579. -. 1988. Taxonomy and evolution of the sinica group of macaques. 6. Interspecific comparisons and synthesis. Fieldiana: Zoology, n.s., 45: 1-44. Fooden, J., and S. M. Lanyon. 1989. Blood-protein allele frequencies and phylogenetic relationships in Macaca: A review. American Journal of Primatology, 17:209-241. Geoffroy Saint-Hilaire, I. [1843]. Description des mammiferes nouveaux ou imparfaitement con- nus. . . . Premier memoire. Famille des singes. Ar- chives du Museum d'Histoire Naturelle, Paris, 2: 485- 592. [For date of publication, see I. Geoffroy, 1851, P- 29.] . 1851. Catalogue Methodique de la Collection des Mammiferes. . . . Gide et Baudry, Paris, 96 pp. . [1852]. Description des mammiferes nouveaux ou imparfaitement connus. . . . Troisieme memoire. Famille des singes; supplement. Archives du Museum d'Histoire Naturelle, Paris, 5 529-584. [For date of publication, see Royal Society of London, Catalogue of Scientific Papers ( 1 800-1 863), vol. 2, p. 836 ( 1 868).] Gervais, P. 1854. Histoire Naturelle des Mammi- feres. . . . L. Curmer, Paris, [420] pp. Gogorza y Gonzalez, J. 1888. Datos para la fauna Filipina. Anales de la Sociedad Espanola de Historia Natural, 17: 247-303. Gonzales, R. B. 1 968. A study of the breeding biology and ecology of the monkey-eating eagle. Silliman Jour- nal, 15:461^191. Goodman, M., A. Kulkarni, E. Poulik, and E. Reklys. 1965. Species and geographic differences in the trans- ferrin polymorphism of macaques. Science, 147: 884- 886. Gray, J. E. 1849. Vertebrata, pp. 1-43. In Adams, A., ed., The Zoology of the Voyage of H.M.S. Sama- rang. . . . Reeve, Benham, and Reeve, London. . 1870. Catalogue of Monkeys, Lemurs, and Fruit-Eating Bats in the Collection of the British Mu- seum. British Museum, London, 137 pp. Groves, C. 1971. Systematics of the genus Nycticebus, pp. 44-53. In Biegert, J., and W. Leutenegger, eds., Proceedings of the Third International Congress of Primatology, Zurich 1970. Vol. 1, Taxonomy, Anat- omy, Reproduction. S. Karger, Basel. Gunther, A. [1877]. Report on some of the additions to the collection of mammals in the British Museum: 1. On a collection from the Philippine Islands. Pro- ceedings of the Zoological Society of London, 1876: 735-736. [For date of publication, see Proceedings of the Zoological Society of London, 107(ser. A): 73 (1937).] . 1879. List of the mammals, reptiles, and ba- trachians sent by Mr. Everett from the Philippine Is- lands. Proceedings of the Zoological Society of Lon- don, 1879: 74-79. Heaney, L. R. 1985a. Zoogeographic evidence for middle and late Pleistocene land bridges to the Phil- ippine Islands. Modern Quaternary Research in Southeast Asia, 9: 127-143. . 1985b. Systematics of Oriental pygmy squirrels of the genera Exilisciurus and Nannosciurus (Mam- malia: Sciuridae). Miscellaneous Publications, Muse- um of Zoology, University of Michigan, No. 170: 1- 58. . 1986. Biogeography of mammals in SE Asia: Estimates of rates of colonization, extinction and spe- ciation. Biological Journal of the Linnean Society, 28: 127-165. Heaney, L. R., P. C. Gonzales, R. C. B. Utzurrum, and E. A. Rickart. In preparation. The mammals of Catanduanes Island, Philippines. Heaney, L. R., P. D. Heideman, and K. M. Mudar. 1981. Ecological notes on mammals in the Lake Ba- linsasayao region, Negros Oriental, Philippines. Sil- liman Journal, 28: 122-131. Heaney, L. R., and D. S. Rabor. 1982. Mammals of Dinagat and Siargao islands, Philippines. Occasional Papers of the Museum of Zoology, University of Mich- igan, No. 699: 1-30. Heideman, P. D. 1988. The timing of reproduction in the fruit bat Haplonycteris fischeri (Pteropodidae): Geographic variation and delayed development. Jour- nal of Zoology, London, 215: 577-595. Hill, W. C. O. 1 974. Primates: Comparative Anatomy and Taxonomy. VII. Cynopithecinae: Cercocebus, Macaca, Cynopithecus. John Wiley and Sons, New York, 934 pp. Hobbs, K. R. 1989. The SICONBREC project-Con- ditioning and breeding facilities for the cynomolgus monkey (Macaca fascicularis) in the Philippines: A review after the first five years. Animal Technology, 40: 55-68. Hollister, N. 1912. A list of the mammals of the Philippine Islands, exclusive of the Cetacea. Philip- pine Journal of Science, ser. D, 7: 1-64. . 1913. A review of the Philippine land mam- mals in the United States National Museum. Pro- ceedings of the United States National Museum, 46: 299-341. Hoogstraal, H. 1951. Philippine Zoological Expe- dition 1946-1947: Narrative and itinerary. Fieldiana: Zoology, 33: 1-86. Hurme, V. O. 1960. Estimation of monkey age by dental formula. Annals of the New York Academy of Sciences, 85: 795-799. Ishimoto, G. 1973. Blood protein variations in Asian macaques: III. Characteristics of the macaque blood protein polymorphism. Journal of the Anthropologi- cal Society of Nippon, 81: 1-13. [In Japanese, English summary.] IUCN. 1971. United Nations List of National Parks and Equivalent Reserves, 2nd ed. Hayez, Brussels, 60 1 pp. Kavanagh, M., and E. Laursen. 1984. Breeding sea- 30 FIELDIANA: ZOOLOGY sonality among long-tailed macaques, Macacafascicu- laris, in peninsular Malaysia. International Journal of Primatology, 5: 1 7-29. Kellogg, R. 1944. A new macaque from an island off the east coast of Borneo. Proceedings of the Biological Society of Washington, 57: 75-76. . 1945. Macaques, pp. 113-134. In Aberle, S. D., ed., Primate Malaria. National Research Council, Division of Medical Science, Office of Medical Infor- mation, Washington, D.C. Kennedy, R. S. 1977. Notes on the biology and pop- ulation status of the monkey-eating eagle of the Phil- ippines. Wilson Bulletin, 89: 1-20. . 1981. Saving the Philippine eagle. National Geographic Magazine, 159: 847-856. Khan, R. 1985. Mammals of Bangladesh. Nazma Reza, Dhaka, 92 pp. Koopman, K. 1 976. Catalog of type specimens of re- cent mammals in the Academy of Natural Sciences at Philadelphia. Proceedings of the Academy of Natural Sciences of Philadelphia, 128: 1-24. Kuntz, R. E. 1969. Vertebrates taken for parasitolog- ical studies by U.S. Naval Medical Research Unit No. 2 on Si 1 1 1 man University-Bishop Museum Expedition to Palawan, Republic of the Philippines. Quarterly Journal of the Taiwan Museum, 22: 207-220. La Gironiere, P. P. de. [1853?]. Twenty Years in the Philippines. James Vizetelly and Henry Vizetelly, London, [372] pp. Lawrence, B. 1939. Collections from the Philippine Islands: Mammals. Bulletin of the Museum of Com- parative Zoology at Harvard College, 86: 28-73. Lewis, R. E. 1988. Mt. Apo and other national parks in the Philippines. Oryx, 22: 100-109. LrNNAEUS, C. 1758. Systema Naturae, 10th ed., vol. 1. Impensis Direct. Laurentii Salvii, Holmiae, [824] pp. Lyon, M. W., Jr., and W. H. Osgood. 1909. Catalog of the type-specimens of mammals in the United States National Museum, including the Biological Survey collection. United States National Museum Bulletin, 62: 1-325. Map of Negros Province, Showing Road System . 1975. Office of the Provincial Engineer, [Dumaguete City?], Philippines. Marche, A. 1970. Luzon and Palawan. Filipiniana Book Guild, Manila, 296 pp. [Translation by Ojeda, C, and J. Castro of Lucon et Palaouan; six annees de voyages aux Philippines (1887).] Martens, E. v. 1 876. Die Preussische Expedition nach Ost-Asien. Nach Amtlichen Quellen. Zoologischer Theil, vol. 1. Koniglichen Geheimen Ober-Hofbuch- druckerei, Berlin, 412 pp. Mearns, E. A. 1905. Descriptions of new genera and species of mammals from the Philippine Islands. Pro- ceedings of the United States National Museum, 28: 425^60. Meyer, A. B. 1896. Saugethiere vom Celebes- und Philippinen-Archipel, I. Abhandlungen und Berichte des Koniglichen Zoologischen und Anthropologisch- Ethnographischen Museums zu Dresden, 1896/97(6): 1-36. Murray, J. 1885. Report on the Scientific Results of the Voyage of H.M.S. Challenger . . . Narrative, vol. I. Second Part. Her Majesty's Stationery Office, Lon- don, 51 1-1 110 pp. Musser, G. G., and L. R. Heaney. 1985. Philippine Rattus: A new species from the Sulu Archipelago. American Museum Novitates, No. 2818: 1-32. Napier, J. R., and P. H. Napier. 1967. A Handbook of Living Primates. Academic Press, London, 456 pp. Napier, P. H. 1 98 1 . Catalogue of Primates in the Brit- ish Museum (Natural History) and Elsewhere in the British Isles. Part II: Family Cercopithecidae, Subfam- ily Cercopithecinae. British Museum (Natural Histo- ry), London, 203 pp. Nozawa, K, T. Shotake, Y. Ohkura, and Y. Tanabe. 1977. Genetic variations within and between species of Asian macaques. Japanese Journal of Genetics, 52: 1 5-30. Ogilvie Grant, W. R. 1897. On the birds of the Phil- ippine Islands. — Part IX. The islands of Samar and Leite. Ibis, 7th ser., 3: 209-250. Petiver, J. 1705. De Quadrupedibus Philippensibus tractat. a Reverendo Georg. Jos. Camel. Philosophical Transactions, Royal Society, London, 25: 2197-2204. PococK, R. I. 1939. The Fauna of British India, in- cluding Ceylon and Burma. Mammalia. Vol. 1, Pri- mates and Camivora (in part). Taylor and Francis, London, 463 pp. Poole, A. J., and V. S. Schantz. 1942. Catalog of the type specimens of mammals in the United States Na- tional Museum, including the Biological Surveys col- lection. United States National Museum, Bulletin, 178: 1-705. Rabor, D. S. 1955. Notes on mammals and birds of the central northern Luzon highlands, Philippines. Part I: Notes on mammals. Silliman Journal, 2: 193-218. . 1977. Philippine Birds and Mammals. Uni- versity of the Philippines, Science Education Center, Quezon City, Philippines, 284 pp. 1986. Guide to Philippine Flora and Fauna: Birds, Mammals. Natural Resources Management Center, Ministry of Natural Resources, and University of the Philippines, Quezon City, Philippines, 211 pp. Raffles, T. S. [1821]. Descriptive catalogue of a zoo- logical collection, made on account of the Honourable East India Company, in Sumatra and its vicinity. . . . Transactions of the Linnean Society of London, 13: 239-274. [For date of publication, see Horsfield, T. (1821-1824), Zoological Researches in Java, and the Neighbouring Islands: Tapirus Malayanus. p. 2.] Rai, N.K. 1990. Living in a lean-to: Philippine Negrito foragers in transition. Anthropological Papers, Mu- seum of Anthropology, University of Michigan, No. 80: 1-184. Rand, A. L., and D. S. Rabor. 1960. Birds of the Philippine Islands: Siquijor, Mount Malindang, Bo- hol, and Samar. Fieldiana: Zoology, 35: 225-441. Reichenbach, L. 1862. Die Vollstandigste Naturge- schichte der Affen. Expedition der Vollstandigste Na- turgeschichte, Dresden/Leipsig, 204 pp. + 501 figs. Rickart, E. A., L. R. Heaney, P. D. Heideman, and R. B. Utzurrum. In preparation. The mammals of Leyte, Biliran, and Maripipi islands. Philippines. FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 31 Rode, P. 1 938. Catalogue des types de mammiferes du Museum Natinal d'Histoire Naturelle. I. Ordre des Primates. A.— Sous-ordre des simiens. Bulletin du Museum National d'Histoire Naturelle, Paris (ser. 2), 10:202-251. Sanborn, C. C. 1952. Philippine Zoological Expedi- tion 1946-1947: Mammals. Fieldiana: Zoology, 33: 89-158. . 1953. Mammals from Mindanao, Philippine Islands, collected by the Danish Philippine Expedition 1951-1952. Videnskabelige Meddelelser fra Dansk Naturhistorisk Forening, 115: 283-288. SAnchez y Sanchez, D. 1900. Los mamiferos de Fi- lipinas (continuation). Anales de la Sociedad Espanola de Historia Natural, 29: 177-290. Schlegel, H. 1876. Monographic 40: Simiae. Revue Methodique, Museum d'Histoire Naturelle des Pays- Bas, 7: 1-356. Sclater, P. L. 1875. Report on additions to the So- ciety's Menagerie in April 1875. Proceedings of the Zoological Society of London, 1875: 348-349. Sharpe, R. B. 1876. Prof. Steere's expedition to the Philippines. Nature, 14: 297-298. Slack, J. H. 1867. Mammalogical notices. Proceedings of the Academy of Natural Sciences of Philadelphia, 19: 34-38. Steere, J. B. 1888a. A month in Palawan. American Naturalist, 22: 142-145. . 1888b. Six weeks in southern Mindanao. American Naturalist, 22: 289-294. . 1890. A List of the Birds and Mammals Col- lected by the Steere Expedition to the Philippines. . . . Courier Office (Printers), Ann Arbor, 30 pp. Stiles, C. W., and M. O. Nolan. 1929. Key catalog of primates for which parasites are reported. U.S. Hy- gienic Laboratory Bulletin, No. 152: 409-601. Swire, H. 1938. The Voyage of the Challenger: A Per- sonal Narrative of the Historic Circumnavigation of the Globe in the Years 1872-1876, vol. 2. Golden Cockerel Press, London, 169 pp. Taylor, E. H. 1934. Philippine Land Mammals. Bu- reau of Science, Manila, 548 pp. Thomas, O. 1898. On the mammals obtained by Mr. John Whitehead during his recent expedition to the Philippines. Transactions of the Zoological Society of London, 14: 377-^12. . 1906. 1. Mammals, pp. 3-66. In The History of the Collections Contained in the Natural History Departments of the British Museum. Vol. II, Separate Historical Accounts of the Several Collections Includ- ed in the Department of Zoology. British Museum, London. . 1911. The mammals of the tenth edition of Linnaeus; an attempt to fix the types of the genera and the exact bases and localities of the species. Proceed- ings of the Zoological Society of London, 1911: 120- 158. Timm, R. M., and E. C. Birney. 1 980. Mammals col- lected by the Menage Expedition to the Philippine Islands and Borneo, 1890-1893. Journal of Mam- malogy, 61: 566-571. Trouessart, E.-L. 1904. Catalogus Mammalium Tarn Viventium Quam Fossilium, suppl. 5. R. Friedlander and Sohn, Berlin, 929 pp. Tweeddale, A. 1880. On the birds collected in the Philippine Islands, pp. 5-25. In Thomson, C. W., ed., Report on the Scientific Results of the Voyage of H.M.S. Challenger during the Years 1 873-76 . . . Zoology, vol. II. Her Majesty's Stationery Office, London. Ulack, R., and G. Pauer. 1989. Atlas of Southeast Asia. Macmillan, New York, 1 7 1 pp. U.S. Board on Geographic Names. 1953. Prelimi- nary Gazetteer: Philippine Islands, 2 vols. Washing- ton, D.C., 1303 pp. Van Couvering, J. A., and G. Kukla. 1988a. Pleis- tocene, pp. 459-464. In Tattersall, I., E. Delson, and J. A. Van Couvering, eds., Encyclopedia of Human Evolution and Prehistory. Garland, New York. . 1988b. Sea-level change, pp. 505-510. In Tat- tersall, I., E. Delson, and J. A. Van Couvering, eds., Encyclopedia of Human Evolution and Prehistory. Garland, New York. Wagner, J. A. 1855. Schreber's Die Saugthiere, suppl. vol. 5. T. O. Weigel, Leipzig, 810 pp. Walden, A. 1875. A list of the birds known to inhabit the Philippine archipelago. Transactions of the Zoo- logical Society of London, 9: 125-252. Waterhouse, G. R. 1838. Catalogue of the Mammalia Preserved in the Museum of the Zoological Society of London, 2nd ed. Richard and John E. Taylor, London, 68 pp. . 1839. Supplement to the Catalogue of the Mammalia Preserved in the Museum of the Zoological Society of London. Richard and John E. Taylor, Lon- don, 13 pp. Whitehead, J. 1899. Field-notes on birds collected in the Philippine Islands in 1893-6. Ibis, ser. 7, 5: 81- 111. Worcester, D. C. 1898. The Philippine Islands and Their People. Macmillan, New York, 529 pp. Worcester, D. C, and F. S. Bourns. 1898. Contri- butions to Philippine ornithology. Part I.— A list of the birds known to inhabit the Philippine and Palawan islands, showing their distribution within the limits of the two groups. Proceedings of the United States Na- tional Museum, 20: 549-625, fold-out map. . [1905]. Letters from the Menage Scientific Ex- pedition to the Philippine Islands. Bulletin of the Min- nesota Academy of Natural Sciences, 4: 1 3 1-1 72. [For date of publication, see Ruch, T. C, 1941, Bibliogra- phica Primatologica, Charles C Thomas, Springfield, Illinois, p. 196.] Appendix 1: Specimens Examined Total, 484 (Philippines, 352; Sabah, 132). Lo- cality numbers (italicized) in parentheses; for de- tails, see Gazetteer (Appendix 2) and figure 1 . 32 FIELDIANA: ZOOLOGY Philippines Skins and Skulls, 1 89 M.f.fascicularis, 43 — Mindanao: Gubat (1 10), fmnh 67725, 67729-67731; Libu (110), fmnh 67727-67728; Sigayan (/ 10), fmnh 67722, 67732, 67740-67742; Situbo (110), fmnh 67719-67721, 67723-67724; Tacuta (770), fmnh 67733-67735; Tampalan (770), fmnh 67736; Canibongan (777), fmnh 67737-67739; Mamara (7 72), fmnh 677 1 7- 677 1 8; Bucong (7 13), fmnh 65440-65444; Pulun- bato, Mount (777), bm(nh) 1880.11.24.1; Zam- boanga(777),uSNM 144667-144668, 144698; San Ramon (118), fmnh 33507-33511. Cagayan Sulu: Cagayan Sulu I. (727), usnm 125325. Basi- lan: I salvia (722), usnm 125326; Camp No. 4- Camp No. 5, between (123), usnm 144665; Ba- silan I., E end (725), usnm 144666. M. f. philippinensis, 55 — Balabac: Minagas Point (7), usnm 477841. Palawan: Brookes Point (3), fmnh 62914, pnm 1 197; Macagua (3), usnm 477849; Lapulapu (4), fmnh 62901; Puerto Prin- cesa, E (5), fmnh 62902, 62905, 62913, fmnh 62903 (skull)/PNM 1191 (skin); Puerto Princesa, "Mt. wooded area" (5), usnm 333197; Tarabanan (6), usnm 477840, 477848. Culion: Makinis (9), pnm 1196; San Pedro (9), fmnh 62907-62908, pnm 1 193-1995. Mindoro: Pinamalayan (13), mcz 35059-35060, 35294; Pasi (14), mcz 35295; Alag River (77), usnm 144674; Halcon, Mount (18), usnm 144675. Luzon: Lopez (26), bm(nh) 1899.1.8.1; Batangas (28), usnm 1 14140-1 14142; Mahayahaya (29), bm(nh) 1876.10.4.9; Maqui- ling. Mount (33), amnh 1 75450, uplbzd (unnum- bered); Jalajala (35), mnhn 83/56; Bulacan Prov. (38), zmb 5443; Data, Mount (43), bm(nh) 1 897.5.2. 1 ; Dimalasud Barrio (44), amnh 1 872 1 8; San Mariano Munic. (44), amnh 1 872 1 5-1 872 1 7; Cagayan Valley (46), usnm 25607 1-256072; Barit (47), bm(nh) 1895.8.2.1; Lagarigilang (49), mcz 35057-35058; Nagpartian (52), usnm 144676, 144678. Samar: Matuguinao (55), fmnh 87717- 87719. Leyte: Patoc Barrio (62), amnh 187939- 187940; Balinsasayo Barrio (64), amnh 187936- 187938. Mindanao: Surigao (82), bm(nh) 1877.10.6.1; Agusan River (84), bm(nh) 1877.10.6.2. M. f. fascicularis/M. f. philippinensis Contact Zone, 80 — Negros: Amio (70), fmnh 65452; Na- liong (70), fmnh 67988-67993; Kabungahan (77), fmnh 65449-65451; Pamo-at (72), fmnh 65445- 65448; Balangbang (73), fmnh 67996-67997; Kandomao (73), fmnh 67994-67995; Kauitan (74), fmnh 67998; Inubungan (75), fmnh 66327-66333; Bunga Barrio (77), ummz 161308; Balinsasayo, Lake, [N bank] (78), fmnh 66339-66348; Pam- plona, 12 km S, 8 km W (78), ummz 157015; Pamplona, 18 km S (78), ummz 130418; Talinis (78), fmnh 75598-75603; Mabaja (80), fmnh 66334-66337. Mindanao: Kamansi (97), fmnh 56493; Sumlog River (92), usnm 1 25323; McKin- ley, Mount, E slope, 4800 ft (94), fmnh 56161- 56 1 62, pnm 1 1 84; Apo, Mount (95), usnm 125319- 125322; Mainit, Mount Apo, 3800 ft (95), fmnh 61026; Matutungan (96), pnm 1 185; Badiang (97), fmnh 56435; Pantod, Mount (97), pnm 1 186; Bu- rungkot (702), fmnh 56490, 56492, fmnh 56491 (skull)/PNM 1 188 (skin); Bugusan (70J), pnm 1 187; Pantar (706), usnm 123448-123450, 123452- 123456; Catagan (108), usnm 144671-144672, 144692, 144694, 144696-144697. Probably M. f. fascicularis/M.. f. philippinensis Contact Zone, 1 —Mindanao: Tangub (707), usnm 144669 (skin incomplete). Subspecies undetermined, 10 — Balut: Balut I. (720), usnm 144664. Island Unknown (not mapped): ?Manila, rmnh 80 1 ; no locality, bm(nh) 1859.7.9.5, rmnh 1714, usnm 29680, 308725, 332896-332897, 344993-344994. Skins Only, 34 M.f.fascicularis, 3 — Mindanao: Zamboanga del Sur Prov. (not mapped), siconbrec 1586 (liv- ing captive). Tawitawi: Tawitawi I. (727), si- conbrec 1225, 1475 (living captives). M. f. philippinensis, 15 — Palawan: Puerto Princesa (5), rmnh q, zmb A4644; Puerto Princesa, E (5), pnm 1 192. Mindoro: Bulalacao (72), usnm 151717; Calapan (76), amnh 30595. Luzon: Ma- quiling, Mount (33), uplbcf (unnumbered, 5 spec- imens); Jalajala (35), mnhn 80/1684; Daraitan, Mount (36), siconbrec 100 (living captive); Ma- nila (37), irsn 9323, mnhn 373/265; no locality, rmnh 55. Probably M. f. philippinensis, 1— Maripipi: Maripipi I. (58), ummz 156931 (in fluid). M. f. fascicularis/M. f. philippinensis Contact Zone, 5 — Negros: Negros I., S (not mapped), bm(nh) 1872.8.20.5. Mindanao: Busaw, Mount (700), pnm 1190; Maculi Point (707), amnh 242092-242093; Pantar (706), usnm 123451. Subspecies Undetermined, 10 — Busuanga: Di- mana (70), fmnh 62906. Island Unknown: amnh 185134, bm(nh) 1939.622, fmnh 65453, mnhn 1880/634, rmnh 1447, ummz 102455, uplbzd, (unnumbered), usnm 308724, 399509. FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 33 Skulls Only, 1 29 M. f. fascicularis, 2 — Mindanao: Zamboanga (J17), zmb A2920. Jolo: Crater Lake Mountain, foot (126), usnm 125324. M. f. Philippine nsis, 30 — Balabac: Minagas Point (/), usnm 477844-477846. Palawan: Man- talingajan, Mount (2), usnm 477843; Malabusog (7), usnm 477842; Palawan I. (not mapped), mnhn 83/1270. Mindoro: no locality, mmnh 4738. Luzon: Batangas (28), usnm 114139; Jalajala (35), mnhn 80/2460; Manila (37), nhrm M769; Mas- sisiat (50), fmnh 62273-62276; Nagpartian (52), usnm 144679-144681, 144682 (fetus)- 144687; Luzon I. (not mapped), zmb 5442, zmb (unnum- bered). Leyte: Baybay, 8.5 km N, 2.5 km E (63), ummz 151311; Pangasugan, Mount (63), usnm 458725-458726; Leyte I. (not mapped), usnm 282628. Leyte or Samar: no locality, usnm 277638. Probably M. f. philippinensis, 14 — Maripipi: Maripipi, 2 km N, 4 km W (58), usnm 458727; Maripipi I. (58), ummz 161312-161315, 161316 (mandible only), usnm 458728-458730, 458731 (mandible only), 458732^58734. Biliran: Sayao, Mount (59), usnm 458724. M. f. fascicularis/ M. f. philippinensis Contact Zone, 23— Negros: Pagyabonan (69), fmnh 66338; Naliong (70), pnm (coll. nos. 525, 549, 597, 601); Balangbang (73), pnm (coll. no. 626); Kandomao (73), pnm (coll. no. 535); Balinsasayo, Lake (78), ummz 158623; Balinsasayo, Lake, 6 km N and 14 km W of Dumaguete City (78), ummz 161309; Pamplona, near (79), ummz 161310; Negros I., S (not mapped), smtd B437, ummz 161317-161318, zmb 1448. Mindanao: Kamansi (97), pnm (coll. no. 1 760); McKinley, Mount, E slope, 4800 ft (94), fmnh 56160; Caburan (99), pnm (coll. no. 1809); Maculi Point (101), amnh 242094; Lanao, Lake (105), usnm 151661-151662; Catagan (108), usnm 144673, 144693, 144695. Probably M. f. fascicularis/ M. f philippinensis Contact Zone, 8— Mindanao: Tangub (107), usnm 144670; Masawan, Mount Malindang, 3500-4500 ft (109), fmnh 87428-87434. Subspecies Undetermined, 52— Busuanga: San Nicolas (77), usnm 477847. Sibuyan: Guitingui- tin, Mount (27), fmnh 135714. Panay: Calantas forest (60), fmnh 1 172 (an unlabeled dark brown skin in mmnh, apparently the "Panay" skin listed by Timm & Birney, 1 980, p. 568, may belong with this skull), mmnh 4740. Bohol: Sandayong (65), fmnh 87419-87420, 87427; Cantaub (66), fmnh 87422-87426; Sandayong or Cantaub (65 or 66), fmnh 87421. Cebu: Cebu I. (67), smtd 1069. Island Unknown: "Manila" (not mapped), aiuz 18-19, 28^11, 341-343, 347-348, 350-352, 354, AS344-AS346, AS349, AS353; no locality, pnm (unnumbered, 2 specimens), usnm 144688- 144689, 257991, 258587, 279563, 308723. Sabah (Localities Not Mapped) Skins and Skulls, 66 M.f fascicularis— Borneo: Abai, mcz, 21 spec- imens; Betotan, Sungai, zrc, 1 ; Bongkabong, mcz, 3; Bundu Tuhan, usnm, 2; Garau, mcz, 4; Kada- maian, Sungai, mcz, 1 ; Keningau, amnh, 1 ; Keno- kok, zrc, 1; Kiau, mcz, 2, zrc, 2; Kiaulan, mcz, 2; Kinabatangan, Sungai, usnm, 1 ; Kretam Besar, Sungai, fmnh, 1; Kretam Kechil, Sungai, fmnh, 1; Ranau, mcz, 1, usnm, 5; Rayoh, Sungai, Na- tional Reference Collection, National University of Singapore, Singapore, 1; Rugading, mcz, 1; Sa- pagaya Forest Reserve, fmnh, 2; Sibuga Besar, Sungai, fmnh, 1; Talibong, mcz, 3; Tawao, amnh, 1 ; Tenompok, Kampong, mcz, 2; Tinonkok, mcz, 1; Tuaran, mcz, 5. Skins Only, 2 M. f fascicularis— Borneo: Papar, usnm, 1 specimen; Talibong, mcz, 1. Skulls Only, 64 M.f. fascicularis— Borneo: Abai, mcz, 52 spec- imens; Bundu Tuhan, usnm, 2; Darvel Bay, zmb, 1 ; Kinabalu, Mount, mcz, 6; Lahad Datu, zmb, 1 ; Sandakan, W, fmnh, 1 ; Segama, Sungai, zmb, 1 . Appendix 2: Gazetteer Locality names listed as primary entries in this gazetteer preferentially are official names ap- proved by the U.S. Board on Geographic Names (USBGN Gazetteer: Philippine Islands, 1953). For Philippine macaque localities that are not included in the USBGN gazetteer, locality names are spelled here as in the original source. Secondary entries, with cross references to corresponding primary en- tries, indicate variant spellings or alternate locality names that appear on specimen tags, in published literature, or in unpublished manuscripts on Phil- ippine macaques. The sequence of information presented in pri- mary entries is as follows: 34 FIELDIANA: ZOOLOGY 1. Locality name; 2. Altitude, if reported by collector or observer; 3. Name of island and province; 4. Coordinates of locality (principal sources— USBGN Gazetteer; Census of the Philippines: 1960; Map of Negros Province, 1975; infor- mation in field notes of collector or observer); 5. Date of collection or observation; 6. Name of collector or observer; 7. Bibliographic reference (in parentheses) to published or unpublished locality notes, if any; 8. Abbreviated name of museum (see Introduc- tion) where specimens are preserved; 9. Number of specimens available (with indi- cation of part preserved, if skin and skull are not both present); 10. Abbreviated subspecific identification (Mff= M. f. fascicularis; Mfp = M.f philippinensis; MfflMfp = M. f. fascicularis/ M. f. philippi- nensis contact zone; subsp. indet. = subspe- cies undetermined); 1 1. Italicized locality number as shown in distri- bution map (fig. 1). Abra Prov.; Luzon I.; 17°10'-18°00'N, 120°25'- 121°05'E; reported by M. Buzeta and F. Bravo (1850, p. 265). Mfp; see 47-50. Abuyog. See Balinsasayo Barrio. Agusan River, mouth; Mindanao I., Agusan del Norte Prov.; 9°00'N, 125°31'E; collected May 1877 by A. Everett (in Giinther, 1879, p. 74); bm(nh), 1. Mfp; 84. Alag River, ca. 350 ft (= ca. 100 m); Mindoro I., Oriental Mindoro Prov.; ca. 13°22'N, 121°04'E; collected Nov. 1906 by E. A. Mearns (unpubl. ms., usnm); usnm, 1. Mfp; 17. Albay Prov.; Luzon I.; 13°00'-13°30'N, 123°15'- 123°55'E; reported by Casto de Elera (1895, p. 3); specimen(s) in Museo de Santa Tomas, Ma- nila (not seen). Mfp; 24. Amio, river bank, [ < 1 000 ft (= < 300 m)]; Negros I., Negros Oriental Prov.; ca. 9°24'N, 122°58'E; collected 13 May 1948 by D. S. Rabor; fmnh, 1 (second specimen — skin only— listed in fmnh catalog, not located). Altitude from A. L. Rand, unpublished sketch map and notes, fmnh. Mffl Mfp; 70. Apo, Mount, 6000 ft (= 1800 m); Mindanao I., Davao del Sur Prov.; ca. 6°59'N, 125°16'E; col- lected 25-29 June 1904 by E. A. Mearns (1905, p. 429); usnm, 4. Reported by R. E. Lewis ( 1 988, p. 102). MfflMfp; 95. Arubal River, South Coast Range; Mindanao I., province unknown; not located; observed 30 Sep.-5 Oct. 1903 by E. A. Mearns (unpubl. ms., usnm). Subsp. indet.; not mapped. Aurora Memorial Park; Luzon I., Aurora Prov.; ca. 15°55'N, 1 2 1°30'E; reported by IUCN( 1971, p. 390). Mfp; 42. Ayala; Mindanao I., Zamboanga del Sur; 6°57'N, 121°57'E; reported by J. B. Steere (1888b, p. 292). Mff 118. Badiang, Tagabuli region, Santa Cruz Munic; Mindanao I., Davao del Sur Prov.; ca. 6°49'N, 1 25°20'E; collected 9 Dec. 1946 by M. Celestino (see Hoogstraal, 1951, pp. 16, 24, 37); fmnh, 1. MfflMfp; 97. Baganga River, upper; Mindanao I., Davao Ori- ental Prov.; ca. 7°30'N, 126°30'E; observed 30 Apr. 1904 by E. A. Mearns (unpubl. ms., usnm). Subsp. indet.; 87. Balabac I.; Palawan Prov.; 7°50'-8°05'N, 1 16°55'- 117°05'E; reported by J. B. Steere (1888a, p. 144), possibly observed in Aug.-Sep. 1874 (see Sharpe, 1876, p. 297). Mfp; see /. Balambing. See Balimbing. Balangbang, Tolong(= Bayawan Munic), [ca. 300 m]; Negros I., Negros Oriental Prov.; ca. 9°25'N, 122°52'E; collected 9-12 May 1950 by D. S. Rabor; fmnh, 2; pnm, 1 (skull only). Coordi- nates from J. R. Leuterio, uplbcf (pers. comm., 31 July 1989); altitude from A. L. Rand, un- published sketch map and notes, fmnh. Mffl Mfp; 73. Balimbing; Tawitawi L, Tawitawi Prov.; 5°05'N, 119°57'E; unsuccessfully pursued 6 Jan. 1906 by L. Wood, Jr. and E. A. Mearns (unpubl. ms., usnm). Mff; 128. Balinsasayo Barrio, Abuyog Munic; Leyte I., Ley- teProv.; 10°40'N, 1 24°5 7' E; collected 9-16 July 1961 by G. Alcasid and M. Celestino; amnh, 3. Mfp; 64. Balinsasayo, Lake; Negros I., Negros Oriental Prov.; ca. 9°21'N, 123°10'E; reported in June 1 98 1 by local residents (Heaney et al., 1981, pp. 123, 127). Collected 11 Aug. 1982 by P. D. Heideman; ummz, 1 (skull only). MfflMfp; 78. Balinsasayo, Lake, [N bank], 3500-4000 ft (= 1000-1200 m); Negros I., Negros Oriental Prov.; ca. 9°23'N, 123°04'E; collected 21 May-2 June 1949 by D. S. Rabor; fmnh, 10. Locality details from J. R. Leuterio, uplbcf (pers. comm., 31 July 1989). MfflMfp; 78. Balinsasayo, Lake, 6 km N and 14 km W of Du- maguete City; Negros I., Negros Oriental Prov.; 9°22'N, 123°1 1'E; collected 2 Feb. 1984 by L. R. Heaney; ummz, 1 (skull only). MfflMfp; 78. Balit. See Barit. FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 35 Balut I., Sarangani Group; Davao del Sur Prov.; 5°20'-5°25'N, 125°20'-125°25'E; collected 23 Jan. 1906 by E. A. Mearns and Lt. Johnston; usnm, 1 . Observed 6 Oct. 1 906 by E. A. Mearns (unpubl. ms., usnm). Subsp. indet.; 120. Banga I. See Banga, Port. Banga, Port; Mindanao I., Zamboanga del Sur Prov.; 7°30'N, 122°26'E; observed 1 Feb. 1904 by E. A. Mearns (unpubl. ms., usnm) and L. Wood, Jr. Mff; 116. Banganan River, South Coast Range; Mindanao I., province unknown; not located; observed 30 Sep.-5 Oct. 1903 by E. A. Mearns (unpubl. ms., usnm). Subsp. indet.; not mapped. Bangued. See Lagangilang. Barit, 1000 ft (= 300 m); Luzon I., Abra Prov.; ca. 17°20'N, 120°42'E; collected 11 Nov. 1894 by J. Whitehead (1899, p. 84; also see Thomas, 1898, p. 381); bm(nh), 1. Mfp; 47. Basilan I.; Basilan Prov.; 6°25'-6°45'N, 121°45'- 122°20'E; collected Oct.-Nov. 1874 by J. B. Steere (see Sharpe, 1 876, p. 297; Gunther, [1877], p. 735); museum unknown. Collected Nov. 1887 by J. B. Steere (1890, pp. 5, 28); museum un- known. Reported as present 1890-1893 by F. S. Bourns and D. C. Worcester (1894, pp. 6, 61). Mff; see 122-125. Basilan I., E end; Basilan Prov.; ca. 6°35'N, 122°15'E; collected 4 Feb. 1906 by E. A. Mearns (unpubl. ms., usnm), usnm, 1. Mff, 125. Baslut Island. See Balut I. Bataan National Park; Luzon I., Bataan Prov.; ca. 14°30'N, 120°25'E; reported by IUCN (1971, p. 387). Mfp; 32. Bataan Prov.; Luzon I.; 14°25'-14°55'N, 120°15'- 120°35'E; laboratory animals obtained 1960- 1969 by Virus Laboratory of Institute of Public Health, University of Philippines, Manila (Fa- matiga, 1973, p. 3 1 6); apparently not preserved. Mfp; see 32. Batan I.; Batanes Prov.; 20°20'-20°30'N, 121°55'- 1 22°05'E; monkeys absent 28 June-1 2 July 1 989, apparently never present on this island (P. L. Alviola III, pers. comm., 4 Aug. 1989). Not mapped. Batangas; Luzon I., Batangas Prov.; ca. 13°45'N, 121°03'E; collected in 1902 for U.S. Govern- ment Board Pan-American Exposition, Buffalo; usnm, 4 (1 skull only). Mfp; 28. Baybay, 8.5 km N, 2.5 km E, 500 m; Leyte I., Leyte Prov.; 10°46'N, 1 24°49'E; collected 26 May 1984 by L. R. Heaney; ummz, 1 (skull only). Mfp; 63. Bessang Pass National Park; Luzon I., Abra Prov.; ca. 1 7°20'N, 1 20°35'E; reported by IUCN (1971, p. 393). Mfp; 48. Biak-na-bato National Park; Luzon I., Bulacan Prov.;ca. 15°06'N, 1 2 1°04'E; reported by IUCN (1971, p. 391). Mfp; 39. Biliran I.; Leyte Prov.; 1 1°30'-1 1°45'N, 124°20'- 124°35'E; reported by L. R. Heaney (1986, p. 1 32), based on specimen(s) in ummz (not located by me in Oct. 1989). Probably Mfp; see 59. Bilukan. See Bulacan. Bohol I. See Cantaub and Sandayong. Brookes Point; Palawan I., Palawan Prov.; 8°47'N, 117°50'E; collected 10-11 May 1947 by D. S. Rabor (see Hoogstraal, 1951, p. 76); fmnh, 1; pnm, 1. Mfp; 3. Bucong (? = Bukon), Pagadian region; Mindanao I., Zamboanga del Sur Prov.; ca. 7°54'N, 123°30'E; collected 15-18 June 1948 by D. S. Rabor; fmnh, 5. Mff; 113. Bugusan, 50 ft (= 15 m), Parang Munic; Min- danao I., Maguindanao Prov.; 7°27'N, 124°14'E; collected 9 Dec. 1946 by G. L. Alcasid and P. Anonuevo (see Hoogstraal, 1951, p. 62); pnm, \. Mff I Mfp; 103. Bukon. See Bucong. Bulacan Prov.; Luzon I.; 14°40'-15°15'N, 120°40'- 121°15'E; collected Jan.-Mar. 1872 by A. B. Meyer (1896, p. 4; also see Walden, 1875, p. 125); zmb, I. Mfp; 38. Bulalacao; Mindoro I., Oriental Mindoro Prov.; 12°20'N, 121°20'E; collected 17 Oct. 1906 by Dr. Porter; usnm, 1 (skin only). Mfp; 12. Bulalacca. See Bulalacao. Bunga Barrio, Dauin Munic, W; Negros I., Negros Oriental Prov.; 9°12'N, 123°16'E; collected Apr. 1984 by L. R. Heaney; ummz, 1. Mff/ Mfp; 77. Burungkot, Upi Munic; Mindanao I., Maguin- danao Prov.; collected 7-8 Jan. 1947 by F. Wer- ner, A. Gabriel, and P. Anonuevo (see Hoog- straal, 1951, pp. 62, 65); fmnh, 2; fmnh (skull) and pnm (skin), 1. Mff/ Mfp; 102. Busaw, Mount, ca. 500 ft (= 150 m); Mindanao I., Davao del Sur Prov.; ca. 5°54'N, 125°36'E; collected 26 Jan. 1947 by H. Hoogstraal (1951, pp. 37, 62); pnm, 1 (skin only). Mff/Mfp; 100. Busuanga. See Dimana and San Nicolas. Busuanga I.; Palawan Prov.; 1 1°55'-12°20'N, 119°50'-120°20'E; reported as present 1890- 1 893 by F. S. Bourns and D. C. Worcester (1894, pp. 6, 61). Subsp. indet.; see 10-11. Butuan River. See Agusan River. Cabasaran River. See Kabasalan River. Caburan, near sea level; Mindanao I., Davao del Sur Prov.; ca. 5°55'N, 125°39'E; collected 13 36 FIELDIANA: ZOOLOGY Jan. 1947 by H. Hoogstraal (1951, p. 60); pnm, 1 (skull only). Mff/ Mfp; 99. Cagayan Sulu I.; Tawitawi Prov.; 7°00'-7°05'N, 1 18°25'-1 18°35'E; collected 25 Feb. 1904 by E. A. Mearns (1905, p. 431; unpubl. ms., usnm); usnm, 1. Mff; 121. Cagayan Valley; Luzon I., Cagayan Prov. or Isa- bela Prov.; 16°30'-18°20'N, 121°35'-121°55'E; captured 14 Jan. 1931 for National Zoological Park; usnm, 2. Mfp; 46. Calamianes. See Culion, Dimana, and San Nico- las. Calantas forest; Panay I., Capiz Prov.; ca. 1 1°30'N, 122°25'E; collected 11-21 Nov. 1890 by D. C. Worcester and F. S. Bourns (Worcester, 1 898, p. 243; Worcester & Bourns, 1898, map; [1905], pp. 135, 137); fmnh, 1 (skull only; an unlabeled dark brown skin in mmnh, apparently the "Pa- nay" skin listed by Timm & Birney, 1980, p. 568, may belong with this skull); mmnh, 1 (skel- eton only). Subsp. indet.; 60. Calapan; Mindoro I., Oriental Mindoro Prov.; 13°25'N, 121°10'E; collected 19 Oct. 1909 by R. C. Andrews ( 1 9 1 1 , p. 2 1 ); amnh, 1 (skin only). Mfp; 16. Camansi. See Kamansi. Camp No. 2. See Alag River. Camp No. 2-Camp No. 3, between; Basilan I. (NW), Basilan Prov.; ca. 6°40'N, 121°50'E; ob- served 29 Jan. 1906 by E. A. Mearns (unpubl. ms., usnm). Mff; 123. Camp No. 4— Camp No. 5, between; Basilan I. (NW), Basilan Prov.; ca. 6°40'N, 121°50'E; col- lected 31 Jan. 1906 by E. A. Mearns (unpubl. ms., usnm); usnm, 1. Mff, 123. Camp No. 6. See Halcon, Mount. Camp No. 6-Camp No. 7, between. See Tiputipu, near. Camp Pantar. See Pantar. Camp Vicars-Malabang, between; Mindanao I., Lanao del Sur Prov.; ca. 7°40'N, 124°05'E; ob- served 1 1 Apr. 1904 by E. A. Mearns (unpubl. MS., usnm). Mff/ Mfp; 104. Candomao. See Kandomao. Canibongan, Katipunan Munic; Mindanao I., Zamboanga del Norte Prov.; 8°25'N, 123°13'E; collected 5 June 1950 by D. S. Rabor; fmnh, 3. Mff'; 111. Canlaon National Park; Negros I., Negros Occi- dental Prov.; ca. 10°25'N, 123°08'E; reported by IUCN (1971, p. 387). Subsp. indet.; 68. Canlaon Volcano, 6000 ft (= 1800 m); Negros I., Negros Occidental Prov.; ca. 10°25'N, 123°08'E; observed Jan.-Aug. 1 896 by J. Whitehead ( 1 899, p. 85; in Thomas, 1898, p. 381). Subsp. indet.; 68. Canloon Volcano. See Canlaon Volcano. Cantaub, 700-750 m, Sierra-Bullones Munic; Bohol I., Bohol Prov.;ca. 9°48'N, 124°18'E; col- lected 14-29 Apr. 1955 by D. S. Rabor (pers. comm., 4 Aug. 1989; see also Rand & Rabor, 1960, p. 312), fmnh, 5 (skulls only). Subsp. in- det.; 66. Caraga [Munic.]; Mindanao I., Davao Oriental Prov.; ca. 7°20'N, 126°34'E; reported by M. Buzeta and F. Bravo (1850, p. 505). Subsp. in- det.; 89. Catagan [vicinity], 1 100-5750 ft (= 335-1750 m); Mindanao I., Misamis Occidental Prov.; ca. 8°10'N, 1 2 3°40'E; collected 10-25 May 1906 by E. A. Mearns (unpubl. ms., usnm) and R. A. Schroder (in Rand & Rabor, 1 960, p. 262); usnm, 9 (3 skulls only). Mff/ Mfp; 108. Catanduanes I.; Catanduanes Prov.; 13°30'- 14°05'N, 124°00'-124°25'E; observed 1968- 1971 by P. C. Gonzales (see Heaney et al., in prep.). Reported as present in 1988 by local hunters (Heaney et al., in prep.). Mfp; 53. Cateel River and Agusan River, crest between; Mindanao I., Davao Prov.; 7°40'N, 126°10'E; observed 3 May 1904 by E. A. Mearns (unpubl. ms., usnm). Subsp. indet.; 86. Cateel River, lower; Mindanao I., Davao Oriental Prov.; ca. 7°48'N, 1 26°27'E; collected 5 Oct. 1 906 by L. Wood, Jr. (see E. A. Mearns, unpubl. ms., usnm); specimen not located (not in usnm cat- alog). Subsp. indet.; 88. Catubig River, between Laguan (= ?Laoang) and Catubig; Samar I., Northern Samar Prov.; ca. 12°30'N, 125°02'E; observed 1 Oct. 1906 by E. A. Mearns (unpubl. ms., usnm). Mfp; 54. Cavite. See Santa Cruz. Cavite Prov.; Luzon I.; 14°05'-14°30'N, 120°35'- 121°05'E; laboratory animals obtained 1960- 1969 by Virus Laboratory of Institute of Public Health, University of Philippines, Manila (Fa- matiga, 1 973, p. 3 1 6); apparently not preserved. Mfp; see 31. Cawitan. See Kauitan. Cebu I.; Cebu Prov.; 9°25'-l 1°15'N, 123°20'- 124°05'E; reported as present 1890-1893 by F. S. Bourns and D. C. Worcester ( 1 894, pp. 6,61). Collected before 1 896 by unknown collector (see Meyer, 1 896, p. 4); smtd, 1 (skull only). Subsp. indet.; 67. Concepcion. See Tarabanan. Cotabato. See Bugasan and Burungkot. Culion I.; Palawan Prov.; 1 1°35'-12°00'N, 1 19°50'- FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 37 120°05'E; reported as present 1890-1893 by F. S. Bourns and D. C. Worcester (1894, pp. 6, 61). Mfp; see 9. Crater Lake Mountain, foot; Jolo I., Sulu Prov.; ca. 5°58'N, 121°10'E; collected 16 Nov. 1903 by E. A. Mearns (1905, p. 430; unpubl. MS., usnm); usnm, 1 (skull only). Mff, 126. Dagami Munic. See Patoc Barrio. Dalawan Bay. See Minagas Point. Daraitan, Mount, Tanay Munic; Luzon I., Rizal Prov.; 14°37'N, 121°24'E; captive obtained 28 Feb. 1984; siconbrec, 1 (captive examined 4 Aug. 1989). Mfp; 36. Data, Mount, 7500 ft (= 2300 m); Luzon I., Ben- guet Prov.; ca. 16°51'N, 120°52'E; collected Feb. 1895 by J. Whitehead (1899, p. 84; in Thomas, 1898, p. 379; cf. Hoogstraal, 1951, p. 30; Rabor, 1955, p. 195); bm(nh), 1. Mfp; 43. Datu Anib's place. See Catagan [vicinity]. Davao Gulf. See Sumlog River. Davao Prov., Davao del Sur Prov., or Davao Ori- ental Prov.; Mindanao I.; 5°35'-8°00'N, 125°05'- 126°35'E; laboratory animals obtained 1960- 1969 by Virus Laboratory of Institute of Public Health, University of Philippines, Manila (Fa- matiga, 1 973, p. 3 1 6); apparently not preserved. Subsp. indet.; see 86-100. Dimalasud Barrio, San Mariano Munic, Sierra Madre; Luzon I., Isabela Prov.; ca. 17°00'N, 122°00'E; collected 19 May 1961 by American- Philippine Expedition; amnh, 1. For locality data, see amnh catalog, nos. 187121, 187125, 187179, 187180. Mfp; 44. Dimana; Busuanga I., Palawan Prov.; ca. 12°03'N, 1 20°08'E; collected 20 Mar. 1 947 by D. S. Rabor (pers. comm., 4 Aug. 1989; also see Hoogstraal, 1951, p. 81); fmnh, 1 (skin only). Subsp. indet.; 10. Dimaniang. See Dimana. Dinagat I.; Surigao del Norte Prov.; 9°50'-10°30'N, 125°30'-125°40'E; no monkeys seen 21 Apr. 1904 (E. A. Mearns, unpubl. ms., usnm). Mon- keys not included in mammalian faunal list pub- lished by L. R. Heaney and D. S. Rabor (1982, p. 1 6). Not mapped. Disabungan River. See San Mariano Munic. Dulungan, Monte. See Halcon, Mount. Fuyot Spring National Park; Luzon I., Isabela Prov.;ca. 17°10'N, 1 22°10'E; reported by IUCN (1971, p. 392). Mfp; 45. Gubat, Katipunan Munic; Mindanao I., Zam- boanga del Norte Prov.; not precisely located, ca. 8°30'N, 123°15'E; collected 28 May-1 June 1950 by D. S. Rabor; fmnh, 4. Mff; 110. Guimaras I.; 10°25'-10°45'N, 122°25'-122°45'E; Iloilo Prov.; reported as present 1890-1893 by F. S. Bourns and D. C. Worcester (1894, pp. 6, 61). Subsp. indet.; 61. Guitinguitin, Mount; Sibuyan I., Romblon Prov.; 1 2°25'N, 1 22°34'E; captive obtained 3 June 1 989 by S. M. Goodman (mother of captive shot by local resident at least 5 years previously; captive then a nursling); fmnh, 1 (skeleton only). Subsp. indet.; 21. Halcon, Mount, 6000 ft (= 1800 m); Mindoro I., Oriental Mindoro Prov.; ca. 13°16'N, 121°00'E; observed Oct. 1895-Jan. 1896 by J. Whitehead (1899, p. 85; in Thomas, 1898, p. 381). Mfp; 18. Halcon, Mount, spur of main ridge, 6300 ft (= 1900 m); Mindoro I., Oriental Mindoro Prov.; ca. 13°16'N, 121°00'E; collected 21 Nov. 1906 by Sefior Estrellus and E. A. Mearns (unpubl. ms., usnm); usnm, 1. Mfp; 18. Ifugao. See Data, Mount. Inobongan. See Inubungan. Inubungan, Santa Catalina Barrio Sitio; Negros I., Negros Oriental Prov.; ca. 9°23'N, 122°57'E; collected 20-28 Dec. 1948 by D. S. Rabor; fmnh, 7. Mff/Mfp; 75. Isabela, near; Basilan I., Basilan Prov.; ca. 6°42'N, 1 2 1°58'E; collected 26 Jan. 1 904 by E. A. Mearns; usnm, 1. Mff; 122. Isabella. See Isabela, near. Iwahig. See Lapulapu. Jalajala; Luzon I., Laguna Prov. or Rizal Prov.; ca. 14°21'N, 1 2 1°19'E; shot as agricultural pest 1819-1839 by P. P. de La Gironiere ([1853?], p. 193). Collected Feb. 1880 (and possibly other dates) by A. Marche (1970 translation, p. 59); mnhn, 3 ( 1 skull only, 1 skull in skin). Mfp; 35. Jolo I.; Sulu Prov.; 5°50'-6°05'N, 120°50'- 121°25'E; reported as present 1890-1893 by F. S. Bourns and D. C. Worcester ( 1 894, pp. 6, 6 1 ). Mff, see 126. Kabasalan River; Mindanao I., Zamboanga del Sur Prov.; ca. 7°46'N, 122°46'E; observed 30- 31 Jan. 1904 by E. A. Mearns (unpubl. ms., usnm). Mff, 114. Kabongakan. See Kabungahan. Kabungahan, Amio region; Negros I., Negros Ori- ental Prov.; ca. 9°30'N, 122°46'E; collected 11 May 1 948 by D. S. Rabor; fmnh, 3. Coordinates from J. R. Leuterio, uplbcf (pers. comm., 31 July 1989). Mff/Mfp; 71. Kamansi, Mati region; Mindanao I., Davao Ori- ental Prov.; 6°56'N, 126°1 l'E; collected 29 Dec. 1 946 by P. Convocar (see Hoogstraal, 1951, pp. 38 FIELDIANA: ZOOLOGY 37, 59); fmnh, 1; pnm, 1 (skull only). Mff/Mfp; 91. Kandomao, Tolong (= Bayawan Munic); Negros I., Negros Oriental Prov.; ca. 9°23'N, 122°52'E; collected 14 Apr. 1950 by D. S. Rabor; fmnh, 2; pnm, 1 (skull only). Coordinates from J. R. Leuterio, uplbcf (pers. comm., 31 July 1989). Mff/Mfp; 73. Katipunan. See Canibongan, Gubat, Libu, Ma- mara, Sigayan, Situbo, Tacuta, and Tampalan. Kauitan, Santa Catalina Barrio Sitio; Negros I., Negros Oriental Prov.; 9°18'N, 122°53'E; col- lected 7 July 1950 by D. S. Rabor and T. Za- mora; fmnh, 1. Mff/Mfp; 74. Kibawalan, Malalag Munic, 1 200-2200 ft (= 365- 670 m); Mindanao I., Davao del Sur Prov.; ca. 6°20'N, 125°20'E; observed 29 May 1964 by R. B. Gonzales (1968, p. 483). Mff/Mfp; 98. Kraan Point. See Maculi Point. Lagangilang; Luzon I., Abra Prov.; 17°37'N, 1 20°44'E; collected 30 Jan. 1 937 by B. Lawrence (see Barbour, 1939, p. 26); mcz, 2. Mfp; 49. Laguan. See Catubig River. Laguna Prov.; Luzon I.; 14°00'-14°35'N, 121°00'- 121°35'E; reported by Casto de Elera (1895, p. 3); specimen(s) in Museo de Santa Tomas, Ma- nila (not seen). Mfp; see 33-35. Lanao, Lake; Mindanao I., Lanao del Sur Prov.; ca. 7°53'N, 124°15'E; collected before 1913 by G. C. Lewis; usnm, 2 (skulls only). Mff/Mfp; 105. Laoag, N; Luzon I., Ilocos Norte Prov.; ca. 1 8°1 2'N, 1 20°36'E; observed 7 Feb. 1 907 by E. A. Mearns (unpubl. ms., usnm). Mfp; 51. Laoang. See Catubig River. Laoog. See Laoag, N. Lapulapu; Palawan I., Palawan Prov.; ca. 9°45'N, 1 18°39'E; collected 2 Mar. 1947 by D. S. Rabor and F. Werner (see Hoogstraal, 1951, p. 75); fmnh, 1. Mfp; 4. La Union, Cabadbaran Munic; Mindanao I., Agu- san del Norte Prov.; 9°05'N, 1 25°32'E; collected 1951-1952 by F. Salomonsen (see Sanborn, 1 953, p. 285); Zoologisk Museum, Copenhagen, 1 (not seen). Mfp; 83. Lepanto. See Data, Mount. Leyte I.; province unknown; 10°00'-1 1°35'N, 124°15'-125°15'E; collected Apr. 1888 by J. B. Steere ( 1 890, pp. 5, 28); museum unknown. Col- lected 1944-1945 by C. O. Mohr; usnm, 1 (skull only). Mfp; see 62-64. Libu, Katipunan Munic; Mindanao I., Zamboan- ga del Norte Prov.; not precisely located, ca. 8°30'N, 123°15'E; collected 30 May 1950 by D. S. Rabor; fmnh, 2. Mff; 110. Lopez; Luzon I., Quezon Prov.; 13°53'N, 122°15'E; collected before 1899 by A. Everett; bm(nh), 1. Mfp; 26. Los Banos; Luzon I., Laguna Prov.; 14°13'N, 121°1 l'E; observed May 1861 by E. von Mar- tens (1876, pp. 193, 200). Mfp; 34. Lucon. See Luzon I. Luzon I.; province unknown; 12°30'-18°40'N, 1 19°45'-124°15'E; collected in 1865 by [J.] Ver- reaux; rmnh, 1 (skin only). Obtained in Paris, "some years" before 1 867, by J. H. Slack ( 1 867, p. 36); skin and skull formerly in Academy of Natural Sciences, Philadelphia, now lost (J. E. Cadle, letter, 14 Nov. 1989). Collected Jan.- Mar. 1872 by A. B. Meyer (1896, p. 4; also see Walden, 1875, p. 1 25); zmb, 2 (skulls only). Col- lected July 1888 by J. B. Steere (1890, pp. 5, 28); museum unknown. Reported as present 1 890-1 893 by F. S. Bourns and D. C. Worcester (1894, pp. 6, 61). Reported by Casto de Elera (1895, p. 3); specimen(s) in Museo de Santa To- mas, Manila (not seen). Mfp; see 24-52. Mabaha. See Mabaja. Mabaja, Bais City; Negros I., Negros Oriental Prov.; ca. 9°39'N, 1 23°0 1 'E; collected 2 1-26 Apr. 1949 by D. S. Rabor; fmnh, 4. Mff/Mfp; 80. Macagua, 250 ft (= 75 m), Brookes Point Barrio; Palawan I., Palawan Prov.; ca. 8°47'N, 1 1 7°49'E; collected 19 Apr. 1962 by D. S. Rabor; usnm, 1 . Coordinates from J. R. Leuterio, uplbcf (pers. comm., 31 July 1989). Mfp; 3. Maculi Point, Cotabato coast; Mindanao I., Sultan Kudarat Prov.; 6°07'N, 124°19'E; collected 1 920- 1923 by E.H.Taylor (1934, p. 34 1);amnh, 3 (2 skins only, 1 skull only). Mff/Mfp; 101. Mahayahaya, mountains of; Luzon I., Batangas Prov.; ca. 13°55'N, 120°52'E; collected May- July 1874 by J. B. Steere (see Sharpe, 1876, p. 297; Gunther, [1877], p. 735); bm(nh), 1. Mfp; 29. Mahayhay. See Mahayahaya. Mainit Hot Spring National Park; Mindanao I., Agusan del Sur Prov.; ca. 8°20'N, 125°35'E; re- ported by IUCN (1971, p. 391). Subsp. indet.; 85. Mainit, Mount Apo, 3800 ft (= 1150 m); Min- danao I., Davao del Sur Prov.; collected 1 7 Nov. 1946 by H. Hoogstraal (1951, p. 56); fmnh, 1. Mff/Mfp; 95. Makiling, Mount. See Maquiling, Mount. Makinis; Culion I., Palawan Prov.; ca. 11°51'N, 120°01'E; collected 29 Mar. 1947 by P. Afionu- FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 39 evo (see Hoogstraal, 1951, p. 83); pnm, 1. Mfp; 9. Malabang. See Camp Vicars-Malabang. Malabusog, Tinitian Barrio; Palawan I., Palawan Prov.; ca. 10°04'N, 1 19°07'E; collected 29 Apr. 1 962 by M. C. Thompson (see bbm field catalog); usnm, 1 (skull only). Coordinates from J. R. Leuterio, uplbcf (pers. comm., 31 July 1989). Mfp; 7. Malampaya Sound, rocky shore; Palawan I., Pa- lawan Prov.; ca. 10°51'N, 119°20'E; observed 12 Sep. 1906 by P. C. Frier et al. (see E. A. Mearns, unpubl. ms., usnm). Mfp; 8. Malindang, Mount. See Catagan and Masawan. Mamara, Katipunan Munic, 2500 ft (= 750 m); Mindanao I., Zamboanga del Norte Prov.; col- lected 21 May 1950 by D. S. Rabor; fmnh, 2. Mff 112. Manay River, above Manay; Mindanao I., Davao Oriental Prov.; ca. 7°13'N, 126°32'E; collected 6 Oct. 1906 by L. Wood, Jr. (see E. A. Mearns, unpubl. MS., usnm); museum unknown, 2 (skulls only; not seen; not listed in usnm catalog). Subsp. indet.; 90. Manila; Luzon I., National Capital Region; ca. 14°35'N, 121°00'E; collected in 1841 byKapten Nisser; nhrm, 1 (skull only). Living captive pre- sented 6 Aug. 1841 to Menagerie, mnhn, by A. Chenest (see I. Geoffroy, [1843], p. 570); mnhn, 1 (skin only). Collected in 1845 by Dechange; irsn, 1 (skin only). Mfp; 37. "Manila"; island and province unknown; living captive received 8 Jan. 1916; rmnh, 1 . Received (date unknown) from C. W. Stiles; aiuz, 30 (skulls and skeletons only). Subsp. indet.; not mapped. Manille. See Manila. Mantalingajan, Mount; Palawan I., Palawan Prov.; 8°48'N, 1 17°40'E; collected 1 1 Apr. 1962 by M. C. Thompson (see Kuntz, 1969, p. 208); usnm, 1 (skull only). Mfp; 2. Manuk Manka I.; Tawitawi Prov.; 4°48'^°50'N, 1 19°48'-1 19°52'E; monkeys absent 6 Jan. 1906 (E. A. Mearns, unpubl. ms., usnm). Not mapped. Maquiling, Mount, Los Banos Munic; Luzon I., Laguna Prov.; 14°08'N, 121°12'E; collected 16 Feb. 1921 by J. T. Zimmer; amnh, 1. Collected 29 Nov. 1947 by R. M. Sojetado; uplbzd, 1. Collected 1977-1980 by J. Leuterio (pers. comm., 31 July 1989); uplbcf, 5 (skins only). Mfp; 33. Marinduque I.; Marinduque Prov.; 13°10'- 13°35'N, 121°50'-122°10'E; reported by D. San- chez y Sanchez (1900, p. 288). Probably Mfp; 23. Maripipi, 2 km N, 3 km W, 650 m; Maripipi I., Leyte Prov.; 1 1°47'N, 124°19'E; troop heard in 1987 by E. A. Rickart et al. (in prep.). Probably Mfp; 58. Maripipi, 2 km N, 4 km W, 200 m; Maripipi I., Leyte Prov.; 11 °47'N, 1 24° 18'E; collected 20 Apr. 1987 by P. D. Heideman; usnm, 1 (skull only). Probably Mfp; 58. Maripipi I.; Leyte Prov.; 1 1°45'-1 1°50'N, 124°15'- 124°20'E; collected 4-7 July 1981 by L. R. Hea- ney; ummz, 1 (in fluid). Collected 8 May 1984 by P. D. Heideman; ummz, 5 (4 skulls only, 1 mandible only). Collected 22-25 Apr. 1987 by P. D. Heideman; usnm, 7 (6 skulls only, 1 man- dible only). Probably Mfp; 58. Masawan, Mount Malindang, 3500—4500 ft (= 1000-1400 m); Mindanao I., Misamis Occiden- tal Prov.; ca. 8°20'N, 123°36'E; collected 2-9 Apr. 1956 by D. S. Rabor (see Rand & Rabor, 1960, pp. 269, 273; pers. comm., 4 Aug. 1989); 7 (skulls only). Probably Mff I Mfp; 109. Masawan, Mount Malindang, 4400-5000 ft (= 1 340-1 520 m); Mindanao I., Misamis Occiden- tal Prov.; ca. 8°20'N, 123°36'E; collected 25 Dec. 1962 by D. S. Rabor (see Rand & Rabor, 1960, pp. 269, 273); museum unknown, 1 (not seen, listed in bbm field catalog). Probably MfflMfp; 109. Masbate I.; Masbate Prov.; 1 1°45'-12 35'N, 123°10'-124°05'E; reported by D. Sanchez y Sanchez (1900, p. 288). Subsp. indet.; 22. Massisiat, 3500 ft (= 1000 m); Luzon I., Abra Prov.; ca. 17°35'N, 120°55'E; collected 14-19 May 1946 by D. S. Rabor (1955, p. 197; also see Hoogstraal, 1951, p. 32), M. Celestino, A. Castro, H. R. Rabanal, and G. Alcasid; 4 (skel- etons only). Mfp; 50. Matchin. See Jalajala. Mati. See Kamansi. Matuguinao; Samar I., Western Samar Prov.; 12°08'N, 124°53'E; collected 11-21 Apr. 1957 by D. S. Rabor; fmnh, 3. Mfp; 55. Matutungan, Tagabuli region, Santa Cruz Munic, 2500 ft (= 760 m); Mindanao I., Davao del Sur Prov.; ca. 6°51'N, 125°18'E; collected 10 Dec. 1 946 by M. Celestino (see Hoogstraal, 1951, pp. 24, 37); pnm, 1. Mff I Mfp; 97. McKinley, Mount, E slope, 4800 ft (= 1460 m), 5700 ft (= 1740 m), 5800 ft (= 1770 m); Min- danao I., Davao del Sur Prov.; ca. 7°06'N, 125°20'E; collected 20 Aug.-4 Sep. 1946 by H. Hoogstraal (1951, pp. 37, 45, 49); fmnh, 3 (1 skeleton only); pnm, 1 . MfflMfp; 94. McKinley, Mount, E slope, 6400 ft (= 1950 m); 40 FIELDIANA: ZOOLOGY Mindanao I., Davao del Sur Prov.; ca. 7°06'N, 125°20'E; observed 1-12 Sep. 1946 by H. Hoogstraal (1951, pp. 37, 45, 49). MfflMfp; 94. Minagas Point, Dalawan Bay; Balabac I., Palawan Prov.; 7°54'N, 117°05'E; collected 22-25 Apr. 1962 by M. C. Thompson and R. Gonzales (see bbm field catalog); usnm, 4 (3 skulls only). Mfp; 1. Mindanao I.; province unknown; 5°35'-9°50'N, 121°55'-126°40'E; collected Oct.-Dec. 1887 by J. B. Steere ( 1 890, pp. 5, 28); museum unknown. Reported as present 1890-1893 by F. S. Bourns and D. C. Worcester (1894, pp. 6, 61). Subsp. indet.; see 82-1 18. Mindoro I.; province unknown; 12°10'-13°30'N, 120°15'-1 2 1°35'E; collected 1890-1893 by F. S. Bourns and D. C. Worcester (1894, pp. 6, 61); mmnh, 1 (skeleton only). Mfp; see 12-18. Monte Data. See Data, Mount. Monte Pulumbato. See Pulunbato, Mount. Mount Isarog National Park; Luzon I., Camarines Sur Prov.; ca. 13°39'N, 123°23'E; reported by IUCN (1971, p. 388). Mfp; 25. Nagpartian, ca. 400 ft (= 1 20 m); Luzon I., Ilocos Norte Prov.; 18°31'N, 120°40'E; collected 24 Feb.-2 Mar. 1907 by E. A. Mearns; usnm, 1 1 (9 skulls only). One additional specimen (usnm 144677, skin and skull) from this series was transferred in 1936 to the Bureau of Science, Manila (usnm catalog); this specimen probably was destroyed during World War II (see Hoog- straal, 1951, p. 9). Mfp; 52. Naliong, Tolong (= Bayawan Munic), [500-800 m]; Negros I., Negros Oriental Prov.; ca. 9°26'N, 122°52'E; collected 21 Apr.-l May 1950 by D. S. Rabor; fmnh, 6; pnm, 4 (skulls only). Altitude and coordinates from A. L. Rand, unpublished sketch map and notes, fmnh. MfflMfp; 70. Naujan Lake National Park; Mindoro I., Oriental Mindoro Prov.; ca. 13°10'N, 1 2 1°21'E; reported by IUCN (1971, p. 390). Mfp; 15. Naval, 5 km N, 10 km E, 800 m, 850 m; Biliran I., Leyte Prov.; 1 1°36'N, 124°29'E; observed 18 Apr. 1987 and another unknown date in 1987 by E. A. Rickart et al. (in prep.; L. R. Heaney, fmnh, field notes). Probably Mfp; 59. Negros I.; province unknown; 9°05'-l 1°00'N, 122°20'-123°35'E; collected Feb. 1888 by J. B. Steere (1890, pp. 5, 28); museum unknown. Re- ported as present 1890-1893 by F. S. Bourns and D. C. Worcester (1894, pp. 6, 61). Subsp. indet.; see 68-80. Negros I., S; province unknown; coordinates un- known; collected about 1872 by A. B. Meyer (1896, p. 4); bm(nh), 1 (skull not seen); smtd, 1 (skull only); zmb, 1 (skull only). Collected June 1984 by P. D. Heideman; ummz, 2 (skulls only). MfflMfp; see 69-80. Nituna River, South Coast Range; Mindanao I., province unknown; not located; observed 30 Sep.-5 Oct. 1903 by E. A. Mearns (unpubl. MS., usnm). Subsp. indet.; not mapped. Pagadian. See Bucong (? = Bukon). Pagyabonan, Bais City; Negros I., Negros Oriental Prov.; ca. 9°41'N, 122°59'E; collected 14 May 1949 by D. S. Rabor; fmnh, 1 (skull only). Mffl Mfp; 69. Palawan I.; Palawan Prov.; 8°20'-l 1°25'N, 117°10'-119°45'E; collected in 1883 by A. Marche ( 1 970 translation, p. 203); mnhn, 1 (skull only). Collected Aug.-Sep. 1887 by J. B. Steere (1888a, p. 143; 1890, pp. 5, 28); museum un- known. Reported as present 1890-1893 by F. S. Bourns and D. C. Worcester ( 1 894, pp. 6, 6 1 ). Reported by B. D. Cabrera (1973, p. 25 1). Mfp; see 2-8. Pamo-at, Amio region; Negros I., Negros Oriental Prov.; ca. 9°30'N, 122°41'E; collected 28 Apr.- 6 May 1 948 by D. S. Rabor; fmnh, 4. Coordi- nates from J. R. Leuterio, uplbcf (pers. comm., 31 July 1989). MfflMfp; 72. Pamplona, 12 km S, 8 km W, 840 m; Negros I., Negros Oriental Prov.; ca. 9°22'N, 123°03'E; collected June 1981 by P. D. Heideman; ummz, I. MfflMfp; 78. Pamplona, 18 km S; Negros I., Negros Oriental Prov.; ca. 9°18'N, 123°07'E; collected 8 June 1979 by K. M. Mudar; ummz, 1. MfflMfp; 78. Pamplona, near; Negros I., Negros Oriental Prov.; ca. 9°30'N, 123°07'E; collected 22 Apr. 1984 by L. R. Heaney; ummz, 1 (skull only). MfflMfp; 79. Panay. See Calantas forest. Panay I.; province unknown; 10°25'-1 1°55'N, 1 2 1°50'-1 23°10'E; reported origin of zoo animal observed in Manila in 1923 by E. H. Taylor (1934, p. 336). Subsp. indet.; see 60. Pangapuyan I.; Zamboanga del Sur Prov.; 6°54'- 6°55'N, 122°10'-122°1 l'E; observed 2 Feb. 1904 by E. A. Mearns (unpubl. MS., usnm). Mff; 1 19. Pangasinan Prov.; Luzon I.; 1 5°35'-16°l 5'N, 119°45'-120°55'E; albino(s) reported by R. P. Casto de Elera (1895, p. 2). Mfp; 41. Pangasugan, Mount; Leyte I., Leyte Prov.; 10°46'N, 124°49'E; collected Apr. 1987 by L. R. Heaney; usnm, 2 (skulls only). Mfp; 63. Pangasugan, Mount, 300 m, 500 m, 700 m; Leyte I., Leyte Prov.; 10°46'N, 124°49'E; observed 2 FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 41 Mar.-13 Apr. 1987 by E. A. Rickart et al. (in prep.). Mfp; 63. Panguil Bay. See Tangub. Pantar, 1900 ft (= 580 m); Mindanao I., Lanao del Norte Prov.; 8°04'N, 124°14'E; collected 13 Aug.-20 Sep. 1903 by E. A. Mearns (1905, pp. 428, 433); usnm, 9 (1 skin only). Mff/Mfp; 106. Pantod, Mount, Tagabuli region, Santa Cruz Mu- nic, 2500 ft (= 760 m); Mindanao I., Davao del Sur Prov.; ca. 6°49'N, 125°20'E; collected 12 Dec. 1 946 by M. Celestino (see Hoogstraal, 1951, pp. 20, 24, 37); pnm, 1. Mff/Mfp; 97. Panyabunan. See Pagyabonan. Paragua. See Palawan I. Parak Creek, Tudaya, 1 km N, 700-1200 m; Min- danao I., Davao del Sur Prov.; ca. 6°58'N, 1 25°22'E; observed as prey of eagle 1 6 Feb. 1 973 by R. S. Kennedy (1977, p. 5). Mff/Mfp; 93. Parang. See Bugasan. Pasananca. See Pulunbato, Mount. Pasi, Pola Munic; Mindoro I., Oriental Mindoro Prov.; 13°07'N, 1 2 1°19'E; collected 15 Mar. 1937 by F. S. Rivera (see Barbour, 1939, p. 26); mcz, I. Mfp; 14. Patoc Barrio, Dagami Munic; Leyte I., Leyte Prov.; 11°05'N, 124°52'E; collected 25-29 July 1961 by G. Alcasid and M. Celestino; amnh, 2. Mfp; 62. Patok, Bo. See Patoc Barrio. Pinamalayan; Mindoro I., Oriental Mindoro Prov.; 13°02'N, 121°29'E; collected 9 Mar. 1937 by F. S. Rivera (see Barbour, 1939, p. 26); mcz, 3. Mfp; 13. Port Banga. See Banga, Port. Port Princesa. See Puerto Princesa. Puerto Princesa; Palawan I., Palawan Prov.; 9°44'N, 118°44'E; collected 22 ?Feb. 1887, 23 July 1887 by C. C. Platen (see Blasius, 1888, p. 302); rmnh, 1 (skin only); zmb, 1 (skin only). Mfp; 5. Puerto Princesa area; Palawan I., Palawan Prov.; ca. 9°44'N, 1 18°44'E; collected July-Aug. 1874 by J. B. Steere (see Sharpe, 1876, p. 297; Giin- ther, [1877], p. 735); museum unknown. Mfp; 5. Puerto Princesa, E, sea level; Palawan I., Palawan Prov.; ca. 9°44'N, 118°44'E; collected 9 Mar.- 14 Apr. 1947 by H. T. Wright, C. F. Wally, and J. A. McCorkle (see Hoogstraal, 1951, p. 73); fmnh, 3; pnm, 1 (skin only; some postcranial bones in fmnh); fmnh (skull), pnm (skin, post- cranial skeleton), 1. Mfp; 5. Puerto Princesa, "Mt. wooded area"; Palawan I., Palawan Prov.; 9°44'N, 118°44'E; collected 21 May 1965 by R. E. Kuntz; usnm, 1. Mfp; 5. Pulunbato, Mount, near; Mindanao I., Zamboan- ga del Sur Prov.; ca. 6°58'N, 122°05'E; collected 2-A Feb. 1875 by H.M.S. Challenger Expedition (see Murray in Tweeddale, 1880, p. 7; Murray, 1885, p. 665; Swire, 1938, pp. 107, 1 10); bm(nh), \.Mff;117. Quezon National Park; Luzon I., Quezon Prov.; ca. 14°00'N, 121°45'E; reported by IUCN(1971, p. 392). Mfp; 27. Rio Grande. See Pulunbato, Mount. Romblon I.; Romblon Prov.; 12°30'-12°35'N, 122°15'-122°20'E; reported as present 1890- 1 893 by F. S. Bourns and D. C. Worcester ( 1 894, pp. 6, 61). Subsp. indet.; 20. Samar I.; province unknown; 1 1°05'-12°40'N, 124°15'-125°45'E; captive albino presented 23 Apr. 1875 to Menagerie of Zoological Society of London by J. Ross (see Sclater, 1875, p. 349). Collected Apr. 1888 by J. B. Steere (1890, pp. 5, 28); museum unknown. Reported as present 1 890-1 893 by F. S. Bourns and D. C. Worcester (1894, pp. 6, 61; Worcester, 1898, p. 334). Re- ported as present 1894-1897 by J. Whitehead (in Thomas, 1898, p. 381). Mfp; see 54-57. Samboangan. See Pulunbato, Mount. Sandayong, 300-350 m, Sierra-Bullones Munic; Bohol I., Bohol Prov.; ca. 9°45'N, 124°20'E; col- lected Apr-May 1955 by D. S. Rabor (pers. comm., 4 Aug. 1989; also see Rand & Rabor, 1960, p. 311), fmnh, 3 (skulls only). Subsp. in- det.; 65. San Mariano Munic, Sierra Madre; Luzon I., Isabela Prov.; ca. 17°00'N, 122°00'E; collected in 1961 by American-Philippine Expedition; amnh, 3. (For locality, see amnh catalog, nos. 187083-187200.) Reported as prey of local Ne- gritos, 1979-1980, by Rai (1990, pp. 28, 47, 182). Mfp; 44. San Nicolas, 6 km NE; Busuanga I., Palawan Prov.; ca. 12°04'N, 120°12'E; collected 23 May 1962 by M. Thompson (see bbm field catalog); usnm, 1 (skull only). Subsp. indet.; 11. San Pedro; Culion I., Palawan Prov.; 11°51'N, 120°01'E; collected 25-26 Mar. 1947 by D. S. Rabor (see Hoogstraal, 1951, p. 82); fmnh, 2; pnm, 3. Mfp; 9. San Ramon; Mindanao I., Zamboanga del Sur Prov.; 7°00'N, 121°55'E; collected 29 July 1929 by F. C. Wonder; fmnh, 5. Mff; 118. San Sebastian; Samar I., Western Samar Prov.; 1 1°43'N, 125°01'E; collected before 1888 by D. 42 FIELDIANA: ZOOLOGY Sanchez y Sanchez (see Gogorza y Gonzalez, 1 888, p. 255); specimen(s) in Museo de Ciencias naturales de Madrid (not seen). Mfp; 56. Santa Cruz; Luzon I., Cavite Prov.; ca. 14°15'N, 120°50'E; albino captive observed by R. P. Cas- to de Elera (1915, p. 34). Mfp; 31. Santa Cruz; Mindanao I. See Badiang, Matutun- gan, and Pantod, Mount. Santa Maria; Mindanao I., Zamboanga del Norte Prov.; 7°45'N, 122°07'E; observed 17 Feb. 1904 by E. A. Mearns (unpubl. ms., usnm). Mff; 115. Sara n gam Group. See Balut I. Sayao, Mount; Biliran I., Leyte Prov.; 1 1°33'N, 124°29'E; collected 27 Apr. 1987 by E. A. Rick- art; usnm, 1 (skull only). Probably Mfp; 59. Siargao I. monkeys not included in mammalian faunal list published by L. R. Heaney and D. S. Rabor(1982, p. 16). Not mapped. Siaton, 10 km N; Negros I., Negros Oriental Prov.; ca. 9°09'N, 123°02'E; collected 26 July 71962 by ?D. S. Rabor; museum unknown (data from bbm field catalog). Mff/ Mfp 76. Sibutu town; Sibutu I., Tawitawi Prov.; 4°50'N, 1 19°27'E; no monkeys seen 7 Jan. 1906 by E. A. Mearns (unpubl. ms., usnm). Not mapped. Sibuyan I.; Romblon Prov.; 12°15'-12°30'N, 122°25'-122°45'E; reported as present 1890- 1 893 by F. S. Bourns and D. C. Worcester ( 1 894, pp. 6, 61). Subsp. indet.; see 21. Sierra Bullones. See Cantaub and Sandayong. Sierra Madre. See Dimalasud Barrio and San Ma- riano Munic. Sigayan, Katipunan Munic; Mindanao I., Zam- boanga del Norte Prov.; not precisely located, ca. 8°30'N, 123°15'E; collected 23 May-5 June 1950 by D. S. Rabor; fmnh, 5. Mff; 110. Siquijor I.; Siquijor Prov.; 9°05'-9°20'N, 123°25'- 123°40'E; reported as present 1890-1893 by F. S. Bourns and D. C. Worcester ( 1 894, pp. 6,61). Probably Mff/ Mfp; 81. Situbo, Katipunan Munic; Mindanao I., Zam- boanga del Norte Prov.; not precisely located, ca. 8°30'N, 1 23°1 5'E; collected 22-26 May 1 950 by D. S. Rabor; fmnh, 5. Mff, 110. Sohoton Natural Bridge National Park; Samar I., Western Samar Prov.; ca. 1 1°30'N, 125°08'E; reported by IUCN (1971, p. 389). Mfp; 57. South Coast Range. See Arubal River, Banganan River, and Nituna River. Sulu. See Crater Lake Mountain and Jolo. Sulug. See Sumlog River. Sumlog River, mouth; Mindanao I., Davao Ori- ental Prov.; 6°53'N, 126°02'E; collected 12 June 1904 by E. A. Mearns; usnm, 1. Mff/ Mfp; 92. Surigao, immediate vicinity; Mindanao I., Surigao del Norte Prov.; ca. 9°45'N, 125°30'E; collected May 1877 by A. Everett (in Giinther, 1879, p. 74); bm(nh), 1 . No monkeys seen Apr. 1 904 by E. A. Mearns (unpubl. ms., usnm). Mfp; 82. Sutug River. See Sumlog River. Tablas I.; Romblon Prov.; 12°05'-12°40'N, 121°55'-122°10'E; reported as present 1890- 1 893 by F S. Bourns and D. C. Worcester ( 1 894, pp. 6, 61). Subsp. indet.; 19. Tacuta, Katipunan Munic; Mindanao I., Zam- boanga del Norte Prov.; not precisely located, ca. 8°30'N, 123°15'E; collected 4 June 1950 by D. S. Rabor; fmnh, 3. Mff 110. Tagabli. See Pantod, Mount. Tagabuli. See Badiang, Matutungan, and Pantod, Mount. Tagulaya River, left bank, 650 ft (= 200 m); Min- danao I., Davao Oriental Prov.; ca. 6°55'N, 125°27'E; observed 15 June 1904 by E. A. Mearns (unpubl. MS., usnm). Mff/Mfp; 93. Talinis, 3500 ft (= 1000 m); Negros I., Negros Oriental Prov.; ca. 9°21'N, 123°03'E; collected 5 Dec. 1953 by A. L. Rand (unpubl. sketch map and notes, fmnh); fmnh, 6. Mff/Mfp; 78. Tampalan, Katipunan Munic; Mindanao I., Zam- boanga del Norte Prov.; not precisely located, ca. 8°30'N, 123°15'E; collected 4 June 1950 by D. S. Rabor; fmnh, 1. Mff; 110. Tangob. See Tangub. Tangub, near, Panguil Bay, 20 ft (= 6 m); Min- danao I., Misamis Occidental Prov.; 8°03'N, 1 23°44'E; collected 9 May 1 906 by E. A. Mearns (unpubl. ms., usnm; also see Rand & Rabor, 1960, p. 262); usnm, 2 (1 skull only). Probably Mff/Mfp; 107. Tarabanan, Concepcion Barrio, Puerto Princesa region; Palawan I., Palawan Prov.; ca. 10°01'N, 119°02'E; collected 14-15 May 1962 by D. S. Rabor (see Kuntz, 1969, p. 208); usnm, 2. Mfp; 6. Tawitawi I.; Tawitawi Prov.; 5°05'-5°20'N, 119°45'-120°15'E; reported as present 1890- 1 893 by F S. Bourns and D. C. Worcester ( 1 894, pp. 6, 61). Living captives obtained 16 Feb. 1987 and 20 Mar. 1987; siconbrec, 2 (captives ex- amined 4 Aug. 1989). Mff; 127. Terabanon. See Tarabanan. Tiputipu. near; Basilan I., Basilan Prov.; ca. 6°30'N, 1 22°04'E; observed 2 Feb. 1 906 by E. A. Mearns (unpubl. ms., usnm). Mff, 124. FOODEN: SYSTEMATIC REVIEW OF PHILIPPINE MACAQUES 43 Tolong. See Balangbang, Kandomao, and Naliong. Tuy; Luzon I., Batangas Prov.; 14°01'N, 120°44'E; collected before 1888 by D. Sanchez y Sanchez (see Gogorza y Gonzalez, 1888, p. 255); speci- men^) in Museo de Ciencias naturales de Ma- drid (not seen). Mfp; 30. Upi. See Burungkot. Viritar. See Laoag, N. Zambales Prov.; Luzon L; 14°45'-15°50'N, 1 19°50'-120°30'E; laboratory animals obtained 1960-1969 by Virus Laboratory of Institute of Public Health, University of Philippines, Ma- nila (Famatiga, 1973, p. 316); apparently not preserved. Mfp; 40. Zamboanga; Mindanao I., Zamboanga del Sur Prov.;ca. 6°54'N, 122°04'E; collected 13-19 June 1861 by E. von Martens (1876, pp. 193, 206, 362) and Herr Pieschel; zmb, 1 (skull only; series originally also included 3 skins). Collected 23 Mar.-l 1 Apr. 1906 by L. Wood, Jr. and E. A. Mearns; usnm, 3. Mff; 117. Zamboanga area; Mindanao I., Zamboanga del Sur Prov.; ca. 6°54'N, 122°04'E; collected Sep.-Nov. 1874 by J. B. Steere (see Sharpe, 1876, p. 297; Gunther, [1877], p. 735); museum unknown. Mff; 117. Zamboanga del Norte Prov. or Zamboanga del Sur Prov.; Mindanao I.; 6°55'-8°45'N, 121°55'- 123°40'E; laboratory animals obtained 1960- 1969 by Virus Laboratory of Institute of Public Health, University of Philippines, Manila (Fa- matiga, 1 973, p. 3 1 6); apparently not preserved. Mff; see 110-119. Zamboanga del Sur Prov.; Mindanao I.; 6°55'- 8°20'N, 121°55'-123°40'E; captive acquired 30 Mar. 1987; siconbrec, 1 (captive examined 4 Aug. 1989). Mff; see 113, 114, 116-119. 44 FIELDIANA: ZOOLOGY itural Histor> ■ Drivi UlUlUiN Ut'U 05-2496 I 922-9410