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Zoology NEW SERIES, NO. 81

Systematic Review of Southeast Asian Longtail Macaques, Macaca fascicularis (Raffles, [1821])

Jack Fooden

Research Associate Division of Mammals Department of Zoology Field Museum of Natural History Chicago, Illinois 60605-2496

Accepted January 13, 1995 Published November 30, 1995 Publication 1470

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1995 Field Museum of Natural History Library of Congress Catalog Card Number: 95-61092 ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents Subspecies Accounts 70 Macaca fascicularis fascicularis (Raffles, [1821]) 70 Abstract 1 Macaca fascicularis aurea I. Geoffroy, Introduction 1 [1831] 77 Geographic Distribution 2 Macaca fascicularis philippinensis I. Pelage 3 Geoffroy, [1843] 79 General Characterization 3 Macaca fascicularis umbrosa Miller, Sex and Age Variation 11 1902b 79 Geographic Variation 11 Macaca fascicularis fusca Miller, Dorsal Pelage Color Saturation 11 1903a 81 Dorsal Pelage Erythrism 13 Macaca fascicularis lasiae (Lyon, Crown Color 15 1916) 81 Lateral Facial Crest Pattern 18 Macaca fascicularis atriceps Kloss, External Measurements and Propor- 1919c 82 tions 21 Macaca fascicularis condorensis Kloss, Sex and Age Variation 21 1926 84 Geographic Variation 22 Macaca fascicularis tua Kellogg, Head and Body Length 22 1944 86 Tail Length 23 Macaca fascicularis karimondjawae Cranial Characters 30 Sody, 1949 86 Sex and Age Variation 30 Macaca fascicularis subspecies undeter- Geographic Variation 33 mined 87 Molecular Biology and Genetics 35 Evolution and Dispersal 87 > Mitochondrial DNA 35 Stage I, 1 Ma 88 Nuclear DNA 39 Stage II, ca. 160 Ka 88 Multiple Alpha-Globin Genes 39 Stage III, > 18 Ka 90 Highly Repeated Restriction Patterns ... 41 Stage IV, ca. 18 Ka 95 Blood Proteins 42 Stage V, < 18 Ka 95 Blood Groups 45 Stage VI, ca. 4.5 Ka 96 Serum Cholesterol Response 46 Acknowledgments 97 Karyology 47 Literature Cited 98

Diseases 47 Appendix 1 : Specimens Examined 1 20 Malaria 47 Appendix 2: Gazetteer of Non-Philippine

Simian T-Lymphotrophic Retrovirus, Macaca fascicularis Localities 1 28 Type 1 49 Appendix 3: Dorsal Pelage Color Satur- Susceptibility to Attenuated Polio Virus ... 49 ation in Fringing-Island Samples of Natural History 49 Macaca fascicularis Compared with Habitats 49 Saturation in Core-Area Reference Arboreality/Terrestriality 50 Samples 181 Swimming 51 Appendix 4: Dorsal Pelage Color Troop Size and Composition 51 Erythrism in Fringing-Island Home Range, Day Range 51 Samples of Macaca fascicularis Density 53 Compared with Erythrism in

Diet 54 Core-Area Reference Samples 1 84 Predators 54 Appendix 5: Crown Color Pattern Intertroop Behavior 56 Frequency in Samples of Macaca Interspecific Behavior 57 fascicularis That Include Reproduction 58 Dark-Crowned Specimens 187 Fossils and Subfossils 63 Appendix 6: Geographic Variation of Systematics 65 Lateral Facial Crest Pattern in Subspecific Taxonomy 65 Samples of Macaca fascicularis .... 188 Key to Recognized Subspecies 69 Appendix 7: Head and Body Length in

in Fringing-Island Samples of Macaca 7. Crown color pattern in dark-crowned fascicularis compared with head Macaca fascicularis 18 and Body Length in Core-Area 8. Infrazygomatic and transzygomatic lat- Reference Samples Collected at eral facial crest patterns in Macaca fas-

Similar Latitudes 1 90 cicularis 19 Appendix 8: Tail Length in Fringing- 9. Geographic variation of lateral facial crest Island Samples of Macaca fascicu- pattern in samples of Macaca fascicularis laris Compared with Tail Length collected in Indochinese Peninsula, Isth- in Core-Area Reference Samples mus of Kra, Malay Peninsula, and neigh- Collected at Similar Latitudes 193 boring islands 21 Appendix 9: Relative Tail Length in Fring- 10. Latitudinal variation of head and body ing-Island Samples of Macaca fas- length in adult core-area specimens of cicularis Compared with Relative Macaca fascicularis 24

Tail Length in Core-Area Reference 1 1 . Latitudinal variation of head and body Samples Collected at Similar length in adult shallow-water fringing-is- Latitudes 196 land specimens of Macaca fascicularis Appendix 10: Blood-Protein Allele Fre- compared with that in adult core-area

quencies at Polymorphic Loci in 1 1 specimens 26 Geographic Samples of Macaca 12. Latitudinal variation of head and body fascicularis 198 length in adult deep-water fringing-island

Appendix 1 1 : Monthly Distribution of specimens of Macaca fascicularis com- Recorded Reproductive Events in pared with that in adult core-area speci- Natural Populations of Macaca mens 27

fascicularis 200 1 3. Latitudinal variation of tail length in adult Appendix 12: Subspecific Variation of core-area specimens of Macaca fascicu- Head and Body Length, Relative laris 29 Tail Length, and Greatest Length 14. Latitudinal variation of relative tail length of Skull in Macaca fascicularis .... 203 in adult core-area specimens of Macaca Subject Index 204 fascicularis 30 Index to Scientific Names 206 15. Latitudinal variation of tail length in adult shallow-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens 31 16. Latitudinal variation of tail length in adult List of Illustrations deep-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens 32 tail 1 . Known locality records of Macaca fasci- 17. Latitudinal variation of relative length cularis 3 in adult shallow-water fringing-island 2. Detail maps of non-Philippine localities specimens of Macaca fascicularis com- of Macaca fascicularis pared with that in adult core-area speci- A. Northwestern quadrant 4 mens 34 B. Southwestern quadrant 6 1 8. Latitudinal variation of relative tail length C. Southeastern quadrant 9 in adult deep-water fringing-island speci- 3. Approximate extent of dry land in South- mens ofMacacafascicularis compared with east Asia during last glacial maximum, ca. that in adult core-area specimens 35 18 Ka 12 1 9. Skull of adult female Macacafascicularis, 4. Core area, shallow-water fringing islands near topotype 36 and deep-water fringing islands inhabited 20. Skull of adult male Macaca fascicularis, by Macaca fascicularis 13 near topotype 37 5. External characters in Macaca fascicu- 21. Latitudinal variation of greatest length of laris 14 skull in adult core-area specimens of Ma- 6. Dorsal pelage color saturation extremes caca fascicularis 40 in Macaca fascicularis 15 22. Latitudinal variation of greatest length of skull in adult shallow-water fringing- List of Tables island specimens of Macaca fascicularis compared with that in adult core-area specimens 41

23. Latitudinal variation of greatest length of 1 . Dorsal pelage color saturation in core-area skull in adult deep-water fringing-island Macaca fascicularis: summary of varia- specimens of Macaca fascicularis com- tion 16 pared with that in adult core-area speci- 2. Dorsal pelage color saturation in core-area mens 43 Macacafascicularis: geographic variation 24. Blood-protein dendrograms for geograph- of mean value in 2-degree latitude-lon- ic samples of Macaca fascicularis: Den- gitude blocks 16 drogram that excludes Mauritius sample 3. Dorsal pelage erythrism in core-area compared with dendrogram that includes Macaca fascicularis: summary of varia- Mauritius sample; consensus dendrogram tion 17 47 4. Dorsal pelage erythrism in core-area Ma- 25. Type localities and limits of distribution caca fascicularis: geographic variation of of recognized subspecies of Macaca fas- mean value in 2-degree latitude-longitude cicularis and type localities of synony- blocks 17 mous nominal species or subspecies ... 64 5. Crown color pattern in Macaca fascicu- 26. Variation ofgreatest length ofskull in adult laris: summary of variation 18 Macaca fascicularisfusca (: Pu- 6. Lateral facial crest pattern in Macacafas- lau Simeulue) compared with variation in cicularis: summary of variation 20 adult Sumatran M. f fascicularis 83 7. External measurements and proportions 27. Tail length vs. head and body length in in age/sex classes of wild-collected Ma- Macaca fascicularis fusca and M. f la- caca fascicularis 22

siae 83 8 . Head and body length in core-area Macaca 28. Craniometric comparisons of adult Ma- fascicularis: summary of variation 23 caca fascicularis atriceps, M. f condor- 9. Summary comparison of head and body ensis, and mainland Indochinese M. fas- length in fringing-island and core-area cicularis 85 samples of Macaca fascicularis 25 29. Latitudinal variation of adult head and 10. Tail length and relative tail length in core- body length in samples of Macaca fasci- area Macaca fascicularis: summary of cularis collected in the southern part of variation 28

the Indochinese Peninsula and adjacent 1 1 . Summary comparison of tail length and Isthmus of Kra, north of 10°N, compared relative tail length in fringing-island and with that in samples of parapatric hi. mu- core-area samples of Macaca fascicularis latta collected in the northern part of the 33 Indochinese Peninsula 92 1 2. Cranial measurements and proportions in 30. Latitudinal variation of adult greatest age/sex classes of wild-collected Macaca length of skull in samples of Macaca fas- fascicularis 38 cicularis collected in the southern part of 13. Schedule of emergence of deciduous teeth the Indochinese Peninsula and adjacent in captive Macaca fascicularis 38 Isthmus of Kra, north of 10°N, compared 14. Schedule of emergence of permanent teeth with that in samples of parapatric M. mu- in captive Macaca fascicularis 39

latta collected in the northern part of the 1 5 . Greatest length ofskull in core-area Macaca Indochinese Peninsula 93 fascicularis: summary of variation 39

3 1 . Latitudinal variation of adult tail length 16. Summary comparison of greatest length in samples of Macaca fascicularis col- of skull in fringing-island and core-area lected in the southern part of the Indo- samples of Macaca fascicularis 42 Chinese Peninsula and adjacent Isthmus 17. Frequency of mtDNA types revealed by of Kra, north of 10°N, compared with that five restriction enzymes in six geographic in samples of parapatric M. mulatta col- samples of Macaca fascicularis 44 lected in the northern part of the Indo- 18. Frequency of alpha-globin gene haplotypes Chinese Peninsula 94 in geographic samples of Macaca fascicu- laris compared with frequencies in samples mature females reported in nonprovisioned of M. mulatta and M. fuscata 45 troops of Macaca fascicularis 54 19. Blood proteins: summary of frequencies 29. Estimated home range and day range in of major alleles at polymorphic loci in 1 1 troops of Macaca fascicularis 55 geographic samples of Macaca fascicu- 30. Foods reported eaten by nonprovisioned laris 46 troops of Macaca fascicularis 56

20. Summary of allele frequencies at the vita- 3 1 . Daily waking-hour time budget estimates min D binding protein locus in geographic for Macaca fascicularis 57 samples of Macaca fascicularis 47 32. Copulatory behavior data in two Macaca

2 1 . Allele frequencies at the complement C6 fascicularis groups 60 locus in three imprecisely localized geo- 33. Macaque fossils or subfossils collected graphic samples of Macaca fascicularis within geographic range of Macaca fas- 48 cicularis 62 22. Geographic variation ofA-B-O and Lewis 34. Summary comparison of external char- blood group phenotype frequencies in acters in recognized subspecies of Macaca Macaca fascicularis 48 fascicularis 66 23. Geographic distribution of malaria spe- 35. Subspecies of Macaca fascicularis recog- cies identified as natural parasites in Ma- nized as valid by Chasen (1940a) and re- caca fascicularis samples 49 cent authors 68 24. Known vectors of malaria species that 36. Macaca fascicularis aurea: external and have been identified as natural parasites cranial variation in adult specimens ex- in Macaca fascicularis 49 amined 80 25. Simian T-lymphotrophic retrovirus, type 37. Sexual dimorphism of greatest length of

1 : frequency of Macaca fascicularis in- skull compared in Macaca fascicularis

dividuals or groups positive for anti- fusca and Sumatran M. f fascicularis . . 82 bodies 50 38. Comparison of cranial proportions in In- 26. Recorded habitats of Macaca fascicu- dochinese Macaca fascicularis, M.f atri- laris 52 ceps, and M. f condorensis 84 27. Recorded size of nonprovisioned troops 39. Hypothetical stages in evolution and dis- of Macaca fascicularis 53 persal of Macaca fascicularis 88 28. Ratio of sexually mature males to sexually Systematic Review of Southeast Asian Longtail Macaques, Macaca fascicularis (Raffles, [1821]) Jack Fooden

Abstract

The longtail macaque, Macaca fascicularis (Raffles, [1821]), is systematically reviewed, based on examination of 2,049 museum specimens, study of relevant literature, and observation of natural populations. Macaca fascicularis inhabits tropical Southeast Asia, where its natural range extends from southernmost Bangladesh to the Philippines and from islands west of to Timor. This review of M. fascicularis includes analyses of geographic variation in pelage characters, external measurements and proportions, cranial characters, molecular and genetic characters, and disease susceptibility. Evidence concerning natural history, reproduction, and paleontology also is investigated. Ten subspecies of M. fascicularis are recognized, and a key to these subspecies is provided. Subspecies accounts present synonymies, type and type locality information, geographic distributions, and diagnoses. Six hypothetical stages in the evolution and dispersal of this species are proposed and discussed. An annotated gazetteer lists approximately 1,150 localities at which M. fascicularis has been collected or observed.

Introduction land and Bali. Specimens examined are preserved in the following institutions, which hereafter are cited of the abbreviations: Macaca fascicularis (Raffles, [1821]) is widely by means following distributed in mainland and insular Southeast Asia from approximately 2 1°N to 1 0°S latitude and from aiuz Anthropologisches Institut der Uni-

92°Eto 1 2 6°E longitude (Fig. 1). Vernacular names versitat Zurich applied to this species include crab-eating, cynom- amnh American Museum of Natural His- olgus, kra, and longtail macaque. The natural tory, New York abundance of M. fascicularis is indicated by the ansp Academy ofNatural Sciences, Phil- large number of specimens in museum collections adelphia

(Appendix 1), which greatly exceeds that of any bm(nh) British Museum (Natural History), other species of macaque. London The existence of this species was first made bnhs Bombay Natural History Society, known to Western science by the English seaman Bombay W. Dampier (1697, p. 220) and the German mis- ctnrc Centre for Thai National Reference sionary G. J. Camel (in Petiver, 1705, p. 2199), Collections, Thailand Institute of both ofwhom observed M. fascicularis in the Phil- Scientific and Technological Re- ippines. The name "macaque" was first applied to search, Bangkok this monkey by Buffon (in Buffon & Daubenton, fmnh Field Museum of Natural History, 1766, p. 190), based on an imported specimen of Chicago unknown origin. irsn Institut Royal des Sciences Natu- The present review of Macaca fascicularis is relles de Belgique, Brussels based on examination of 2,049 museum speci- mcz Museum of Comparative Zoology,

mens (Appendix 1 ), survey of relevant literature, Harvard University, Cambridge, and observation of natural populations in Thai- Massachusetts

FIELDIANA: ZOOLOGY, N.S., NO. 81, NOVEMBER 30, 1995, PP. 1-206 MMNH Bell Museum of Natural History, of their greater familiarity (alternative names in University of Minnesota, Minne- parentheses): Burma (), (Jawa), apolis Lesser Sunda Islands (Nusa Tenggara), and Su- MNHN Museum National d'Histoire Na- matra (Sumatera). Cited geographic names fre- turelle (Mammiferes), Paris quently include the following generic terms: gun- = Museum Zoologicum Bogoriense, ong (Malaysian) or gunung (Indonesian) moun- = Bogor tain; ko (Thai) island; pulau (Malaysian and = = NHMBa Naturhistorisches Museum, Basel Indonesian) island; and teluk (Indonesian) NHMBe Naturhistorisches Museum, Bern bay. NHRM Naturhistoriska Riksmuseet, This review begins with a general account of the Stockholm geographic distribution of M. fascicular is. Follow- NMS Natur-Museum Senckenberg, ing are discussions of (1) pelage characters, (2) Frankfurt external measurements and proportions, (3) cra- PNM Philippine National Museum, Ma- nial characters, (4) molecular and genetic char- nila acters, and (5) disease susceptibility; these discus- Primate Research Institute, Kyoto sions emphasize patterns of geographic variation. University, Inuyama Subsequent sections summarize available infor- rcs(om) Royal College of Surgeons, Odon- mation concerning natural history, reproduction, tological Museum, London and paleontology. A discussion of subspecific tax- RMNH Rijksmuseum van Natuurlijke His- onomy, supplemented by a key to recognized sub- toric, Leiden species, is based on material presented previously. SICONBREC Simian Conservation, Breeding & For each recognized subspecies, a formal subspe- Research Center, Inc., Tanay, Lu- cies account provides the following: annotated zon synonymy; information concerning types, type lo- SMK Museum, Kuching cality, and distribution; subspecific diagnosis; and SMTD Staatliches Museum fur Tierkunde, summary of specimens examined (complete list in

Dresden Appendix 1). The final section of this review pre- UMMZ Museum of Zoology, University of sents a hypothetical interpretation of the evolution Michigan, Ann Arbor and dispersal of M. fascicularis. UPLBCF University ofthe Philippines at Los Banos, College of Forestry, Los Banos, Luzon Geographic Distribution UPLBZD University ofthe Philippines at Los Banos, Zoology Department, Los Macaca fascicularis inhabits tropical Southeast Banos, Luzon Asia (Figs. 1, 2). Its natural geographic range ex- USNM National Museum of Natural His- tends from southernmost Bangladesh and south- tory, Washington, D.C. ern Burma southward and eastward through the ZLUU Zoological Laboratory of Utrecht southern part of the Indochinese Peninsula (south University, Utrecht of 1 7°N), the Isthmus of Kra, the Malay Peninsula, ZMB Zoologisches Museum des Hum- Sumatra, , Java, and the Lesser Sunda Is- boldt-Universitat, Berlin lands as far east as Timor, the Philippine Islands, ZMUZ Zoologisches Museum der Univ- and numerous small adjacent islands, including ersitat Zurich the southernmost three Nicobar Islands. ZRC Zoological Reference Collection, The presence of M. fascicularis in some islands Department of Zoology, National in Southeast Asia has been attributed to prehis- University of Singapore toric human introduction (see below, Fossils and ZSBS Zoologisches Sammlung des Bay- Subfossils; Evolution and Dispersal). During his- erischen Staates, Munich toric times, successful introductions have occurred ZSI Zoological Survey of , Na- in Mauritius Island (I.), east of Madagascar, prob- tional Zoological Collection, Cal- ably in the sixteenth century (Sussman & Tatter- cutta sall, 1986, p. 30); southwestern Sulawesi, date un- Geographic names used in this review generally known (Miiller, [1840], p. 17; M. Weber, 1890b, are officially approved names. However, the fol- p. 102; Dammerman, 1929, p. 6); Anguar Island, lowing conventional names are retained because north of New Guinea, early twentieth century

FIELDIANA: ZOOLOGY a Specimens examined

Literature records

10

100 110 120

Fig. 1. Known locality records of Macaco, fascicularis. For details, see Figure 2A-C.

(Poirier & Smith, 1974, p. 264); Hong Kong, ca. core area for less than 1 8 K years; and (3) deep- 1945 (Marshall, 1967, p. 44; Southwick & Manry, water fringing islands— islands beyond the 120-m 1 987, p. 48); and Pulau (P.) Tinjil, south ofwestern bathymetric line that have been isolated from the Java, 1988-1990 (Kyes, 1993, p. 78). core area at least since the beginning of the pre- Physiographically, the geographic range of M. ceding interglacial, ca. 1 20 Ka (Van Couvering & fascicularis includes mainland Southeast Asia, Kukla, 1988, p. 461) and that may never have shallow-water islands on the Sunda Shelf, and deep- been joined to the core area. water islands at or beyond the edge of the Sunda Shelf (Figs. 3, 4). During the most recent glacial maximum, ca. 18 Ka, sea level was reduced by about 1 20 m, which exposed most of the Sunda Pelage Shelf and extended mainland Southeast Asia to include Sumatra, Borneo, Java, and other shallow- General Characterization (Fig. 5) water islands (Heaney, 1986, p. 131; 1991a, p. 56). During the preceding glacial maximum, ca. 160 Dorsal pelage color in M. fascicularis varies from Ka, sea level may have been reduced by about 1 70 buffy to yellowish gray to golden brown to reddish m, which exposed more of the Sunda Shelf but brown to blackish. Individual hairs generally are still left most deep-water islands isolated. conspicuously marked by one or two pale subter- To facilitate zoogeographic analysis, in subse- minal bands that vary in color from pale yellowish quent discussions ofgeographic variation the range to golden to rufescent. The crown usually is more of M. fascicularis is subdivided into three com- brightly colored than the back (pale subterminal ponents: (1) a core area of distribution that com- bands more conspicuous in crown hairs), but in prises mainland Southeast Asia, Sumatra, Borneo, some specimens the crown is contrastingly darker and Java, four large landmasses that were inter- and duller than the back (pale subterminal bands connected by dry land ca. 1 8 Ka; (2) shallow-water reduced or absent in crown hairs). The anterior fringing islands— small islands within the 120-m edge of the crown is bordered by a transverse bathymetric line that have been isolated from the blackish supraorbital streak; hairs at the vertex

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS d Specimens examined Literature records

a-m Absence reports

240 25 ^ ,9 „/ \\.\j "I Vietnam (V)

L $r-^3 * Cambodia

Fig. 2A. Detail map of non-Philippine localities of Macaca fascicularis, northwestern quadrant. (Philippine localities are mapped in Fooden, 1991, p. 2.) For documentation, see Gazetteer (Appendix 2). Abbreviations in parentheses are those used in Gazetteer locality codes.

Bangladesh (Ba) 22. Ban Sadein; Pakchan River, near Bankachon; Pak- chan River, near Maliwun. 1. Bilasodia; Bimirdia; Ghorardia; Ochodia; Ruku- 23. Ru, Pulo. modia. 24. Kyun. 2. Jolir dia.

Cambodia (C) Burma (Bu) 1. Siemreab. 1. Lai char. 2. Pang Roloem-Sur Sdei area. 2. Myengun Kyun. 3. Laem Ngop-Phumi Cham Yearn (Cambodia or 3. Arakan Division. Thailand). 4. Hainggyi Kyun. 5. Island. Laos (L) 6. Desertion Creek. 1. Thateng, Muang. 7. Rangoon. 8. Pegu. Nicobars (N) 9. Wimpong. 10. Haungtharaw. 1 . Ol-kolo-kwak vicinity. 2. Little Nicobar. 1 1 . Ataran River; Ye Forest. 3. Pulo 12. Moscos Island Game Sanctuary. Kopenheat; Nyur. 13. Tavoy River. 4. Campbell Bay vicinity. 14. Taungbyauk Chaung. Thailand (T) 15. Kathema Kyun; Mibya Kyun.

16. Kaser Doo Wildife Sanctuary. 1 . Ban Nam Lai Tai. 17. Kadan Kyun. 2. Ban Mae Na Ree. 18. Great Tenasserim River; Mergui. 3. Wong, Nam Mae, 53 mi (= 85 km) E of Um Pang. 19. Tagoot; Tenasserim; Thagyet. 4. Ping, Mae Nam; Wat Khao Noh. 20. Letsok-aw Kyun. 5. Ban Pak Nam Pho; Wat Krieng Krai . 2 1 . Lanbi Kyun. 6. Wat Khao Wong Kot.

FIELDIANA: ZOOLOGY often form an irregular tuft or crest. Hairs on the mal part of the outer surface is approximately the side of the head are pale ochraceous gray to pale same color as the adjacent surface of the trunk. ochraceous brown and form a variably prominent More distally, the outer surface of the limbs be- lateral facial crest that usually extends from near comes paler, pale grayish to pale golden brown at the angle of the jaw to the crown, but may be the wrists and ankles. The dorsal surface of the restricted to the mandibular region. The face gen- tail is golden brown to blackish basally, becoming erally is thinly haired; the facial skin is brownish paler (pale grayish to grayish brown) distally. Pel- to pinkish, except for the upper eyelids, which are age on the ventral surface of the trunk, limbs, and sharply defined whitish. On the limbs, the proxi- tail is thin and pale gray to whitish.

Sarn Pra Karn. 50. Phangan, Ko. Wat Khao Sompoad. 51. Khao Lampi-Hat Thai Muang National Park; 9. Lat Bua Khao. Phangnga Bay National Park; Wat Tham Suwan 10. Ban Sakaerat. Khuha. 11. Aranyaprathet. 52. Na Ka Yai, Ko; Yao Noi, Ko. 12. Kosumphi Forest Park. 53. Rang Yai, Ko. 13. Wat Koo Pra Kona. 54. Phi Phi Don, Ko. 14. Ban Kosum. 55. Lanta Yai, Ko. 15. Phu Phan. 56. Ban Phra Muang; Hat Chao Mai National Park; 16. Don Poo Tao; Prang Koo. Kantang; Muk, Ko; Talibong, Ko. 17. Ban Wan; Wat Ban Kan Yai. 57. Mu Ko Phetra National Park. 18. Wong, Nam Mae, 40 mi (= 65 km) E of Um Pang. 58. Butang, Ko. 19. Kata Taek; Sap Khao; Thap Salao, Huai. 59. Telok Wau, Ko Tarutao. 20. Ban Tamrong Phato; Khwae Noi, Mae Nam. 60. Ban Nong Kok; Ban Nong Put; Ban Thap Plik, 1 21. Ban Phu Toie. km NE; Hat Noppharat Thara-Mu Ko Phi Phi Na- 22. Khao Phatowee. tional Park; Tham Horn; Wat Tham Sua. 23. Wat Noi Chompoo. 61. Tyching. 24. Wat Phra Buddha Chai. 62. Khao Rang Kai. 25. Pak Chong, Sathani. 63. Nakhon Si Thammarat. 26. Ban Huai Maenam Noi. 64. BanNa. 27. Siam, 13°45'N, 99°25'E. 65. Wat Khuha Sawan; Wat Suwankuha. 28. Khao Suan Luang; Tham Chomphon; Wat Cha-Am 66. Nang Kham, Ko. Kiri; Wat Huai Takhaeng; Wat Khao Chong Phran; 67. Khao Noi/Khao Tangkuan. Wat Khao Khan Hok; Wat Khao Yod Thong; Wat 68. Nong Chik region; Pattani; Pho, Laem; Yaring, tidal Ngern Rung Sawang; Wat Ratch Singkhorn; Wat creeks near; Yaring region. Tham Kunchhorn. 69. Ban Sai Kau; Kampong Biserat; Wat Khuha Phi- 29. Wat Tham Sala. muk. 30. Bangkok; Chao Phraya, Mae Nam. Vietnam (V) 31. Khangkhao, Ko; Samuk, Khao; Si Chang, Ko; Si Racha vicinity. 1. Sontra Peak. 32. Wang Kaew. 2. Lac Giao. 33. Laem Sing mountains. 3. Xa Trang Bom. 34. Ven-Ven. 4. Ho Chi Minh City. 35. Chang, Ko. 5. Tay Ninh. 36. Kut, Ko. 6. Phu Quoc Dao. 37. Khao Wang; Phet Buri; Samut Songkhram vicinity; 7. Airfield building vicinity, Con Son; Ben Dam vi- Wat Ban Rai Don; Wat Bun Thawi; Wat Khao Ban- cinity, Hon Ba; Con Son. dai It; Wat Khao Takhrao; Wat Khao Tamon; Wat Absence reports Kut. 38. Pran Buri, Mae Nam.; Wat Khao Takieb. a. North Andaman Island. 39. Khram Yai, Ko; Klet Kaeo, Ko. b. Middle Andaman Island. 40. Khao Sam Roi National Park. c. Henry Lawrence Island. 41. Prachuap Khiri Khan, few miles north; Wat Tham- d. Barren Island. mikaram Varaviharn. e. South Andaman Island. 42. Wat Tham Khao Phlu. f. Little Jolly Boy Island. 43. Wat Tha Mai Lai. g. Rutland Island. 44. Chumphon, Khlong. h. Little Andaman Island. 45. Phayam, Ko. i. Car Nicobar Island. 46. Laem Son National Park. j. Tillanchong Island, 47. Chiew Larn Reservoir. k. Camorta Island. 48. Thung Thong Waterfowl Reserve. 1. Nancowry Island; Trinkat Island, 49 Khlong Pah Yie, Ko Samui. m. Phai, Ko.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS d Specimens examined Literature records

Java (J) a- n Absence reports """W-Maffn*.

Fig. 2B. Detail map of non-Philippine localities of Macaca fascicularis, southwestern quadrant. (Philippine localities are mapped in Fooden, 1991, p. 2.) For documentation, see Gazetteer (Appendix 2). Abbreviations in parentheses are those used in Gazetteer locality codes.

Indian Ocean (IO) 10. Jasinga. 11. Cihara. 1. We, Pulau. 12. Sarongen; Tamandjaija. 13. Camara. Simeulue, Pulau: 14. Panaitan, Pulau; Peu- 2. Dalam, Lhok; Sibaboh, Lugu; Survey Site LB; Sur- Handeuleum, Pulau; Niur; Suaka vey Site LS; Survey Site SEM. cang, Pulau; Ujungkulon, Margasatwa. 15. Rawa. 3. Ajer Dingin; Labuhanbajau; Sinabang. Danau, 16. Gunung Halimun Reserve. 4. Pulau. Lasia, 17. Bogor; Depok; Salak, Gunung. 5. Tuangku, Pulau. 18. Gede, Gunung; Pangrango, Gunung. 19. Cikarang forest. Nias, Pulau: 20. Cikujang; Pasir Carolina. 6. Lafau. 21. Indramayu; Jatibarang. 7. Soliga. 22. "P. Sember." 8. Hilisimaetano. Majalengka; 23. Cirebon; Linggajati. 9. Samasama. Ciremay, Gunung; 24. Baturaden; Kaligoea. 10. Siaba, Teluk. South China Sea (SCS) 1 1 . Mursala, Pulau. 12. Tanahmasa, Pulau. Pinang, Pulau [2]; Redang, Pulau. 13. Tanahbala, Pulau. Acheh, Pulau. Pulau Tioman: Camp II, Juara side; Juara, Telok; Java (J), including nearby islands Kampong Mukut; Nipah, Telok, vicinity; Sedagong; 1. Cilacap. Tekek. 2. Kalipucang; . Pemanggil, Pulau. 3. Tjeringin. Aur, Pulu. 4. Tasikmalaya. Tinggi, Pulau. 5. Tilu, Gunung. Sentosa Island; Singapore Island: Bukit Timah Na- 6. Malabar; Preanger district. ture Reserve; Changi; MacRitchie Reservoir Nature 7. Bandung, 2000 ft; Bandung, near. Reserve; northwestern part; Pandan, Sungai; Pung- 8. Ciwangi; Singkil, Gunung; Takokak Reserve. gol; Sembawang, Sungai; Singapore Botanical Gar- 9. Bantargebang; Pelabuhanratu, Teluk. dens; western part.

FIELDIANA: ZOOLOGY 8. Pulau Bintan: north coast; Pasir Panjang; Sungei Station; Goenoengsetan-Meloewak; Gumpang, near; Biru. Ketambe Research Station; Ketambe, ca. 5 km S 9. Mentigi, Pulau Mapur. and 10 km S. 10. Tanjong Sauh, Pulau Batam. 14. Ketambe, ca. 30 km SSE.

1 1 . Galang, Pulau; Nguwal, Pulau. 15. Alas, Lae, between Muara Setulen and Lae Renun, 12. Bakung, Pulau; Sebangka, Pulau. between Lae Renun and Bengkong River, and be- 13. Lingga, Pulau. tween Bengkong River and Gelombang; Bukit La- 14. Selayar, Pulau. wang, ca. 35 km SW; Renun, Lae. 15. Bulan, Pulau, south. 16. Alas, Lae, between Gelombang and mouth of river. 16. Durian, Pulau; Durian-kecil, Pulau; Sugi, Pulau; 17. Singkel vicinity. Ungar, Pulau. 18. Tamiang. 17. Karimun-kecil, Pulau; Selatbliat, Pulau Kundur; 19. Medan, ca. 75 km NW; Pangkalansusu; Serangjaya- Pulau Karimun: Mensuda Bay; Monos; Pemeral; hilir; Sikundur. Pongka, Kampung. 20. Bukitlawang; Bukitlawang, Area I and Area II; Bun- gara, Area III; Ober Langkat district; Serapit Tand- Bangka, Pulau: jung; Tandjung Bringin; Tandjung Butus; Unter 18. Jebus; Muntok; Pamuja, Tanjung; Rengsam, Tan- Langkat district. jung; Simpang. 21, Batangkuis; Belawan; Deli, Sungai; Labbuhandeli 19. Sungaiselan. vicinity; Medan, forest near, near sea level, and vi- 20. Batu, Tanjung, Pulau Belitung. cinity; Medan, 20 km N; Medan-Siantar; Tanjung- 21. Serutu, Pulau. morawa; Terbanjawan. 22. Pai, Teluk, Pulau Karimata. 22. Serdang district. 23. Pelapis Tengah, Pulau; Penebangan, Pulau. 23. Batu Bara district; Lauttador; Padang/Bedagei dis- 24. Lemukutan, Pulau. trict; Paguruan; Tandjung; Tebingtinggi vicinity, ca. 25. Pejantan, Pulau. 25 km SSE and ca. 35 km ESE. 26. Benua, Pulau; Jela, Pulau; Selintang, Pulau; Tam- 24. Tebingtinggi, ca. 55 km ESE and ca. 65 km and 75 belan Besar, Pulau; Uwi, Pulau. km SE. 27. Airabu, Pulau; Piling, Pulau. 25. Bukitlawang, ca. 65 km S. 28. Jemaja, Pulau. 26. Medan, ca. 100 km SSW. 29. Telaga, Pulau. 27. Dolok Tinggi Radja Reserve. 30. Matak, Pulau; Mubur, Pulau; Siantan, Pulau. 28. Medan, ca. 105 km SSE. 31. Laut, Pulau. 29. Tebingtinggi, ca. 75 km SSW. 30. Pematangsiantar; Tebingtinggi, ca. 60 km SSW. Natuna Besar, Pulau: 31. Batang Garut. 32. Ulu, Sungai. 32. Tapanuli, Teluk. 33. Binjai, Sungai. 33. Padangsidempuan vicinity. 34. , Pulau. 34. Padangsidempuan, ca. 45 km NNW. 35. Subi-kecil, Pulau. 35. Rantauprapat vicinity. 36. Serasan, Pulau. 36. Telukpanji. 37. Bangko, ca. 15 km, 20 km, and 25 km ENE, ca. 20 Strait ofMalacca (SM) km N, and ca. 30 km NNW. 1. Burau, Pulau; Langkawi, Pulau. 38. Rokan-kanan, Sungai. 2. Pulau Pinang [1]: Krachut; Botanical 39. Bangko, ca. 80 km S. Gardens; Penang Hill; Telok Bahang. 40. Bangko, ca. 90 km SSE; Pagansan; Pap-ka; Siak Co- 3. Pangkor, Pulau. patta; Sumatra, east-central. 4. Pintu Gedong, Pulau. 41 Tapung-kanan, Sungai. 5. , Pulau. 42, Gasip, Sungai, ca. 10 km and 20 km above mouth; 6. Kapos Tinggi, Pulau ; Padang, Pulau, north Mandau, Sungai; Mempura; Perawang, 4 km NW; coast. Siak, Sungai, near , ca. 1 5 km, 30 km, 45 km, 60 km, and 75 km below Pekanbaru, ca. 60 Sumatra (S) km, 48 km, 36 km, 24 km, and 1 2 km above mouth, 1. Meulaboh vicinity. and near mouth. 2. Meulaboh, ca. 35 km NW. 43, Lubuksikaping; Pinagar; Pinagar, ca. 5 km SW and 3. Meulaboh, ca. 60 km NW. ca. 20 km E. 4. Banda , ca. 40 km S and 45 km S. 44 Bukit Cangang. 5. Banda Aceh, near and ca. 20 km SSW. 45 Bukittinggi and vicinity and ca. 20 km S; Maninjau; 6. Banda Aceh, ca. 30 km ENE. Payakumbuh. 7. Banda Aceh, ca. 85 km ESE. 46 Pajo. 8. Takengon, ca. 40 km NNW. 47 Lubukminturun area; Meru, Gunung; Padang; 9. Bur ni Bebuli; Takengon, ca. 15 km NNW. Pangkalan; Pasar Usang area; Sungailundang; Ta- 10. Lhokseumawe, 80 km ESE. rusan, Teluk.

1 1 . Lesten. Kambang. 12. Gunung Leuser Reserve. Siulakderas. 1 3. Alas, Lae, between Agusan and Ketambe Research Sandaran Agong.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 51. Telukkayubutih vicinity, ca. 15 km E, ca. 15 km 6. Hantu, Tanjong. NNE, ca. 1 5 km and 30 km WNW, ca. 30 km and 7. ; Kinta, Daerah. 45 km ESE, and ca. 60 km SE. 8. Telom, Sungai. 52. Kerinci, Gunung, northern foot; Pakan Selasa and 9. Berangkat, Gunong, east. ca. 10 km SSE. 10. , Sungai, vicinity.

53. Sijunjung. 1 1 . Kenyam, Sungai; Negara, Taman; Tahan, Sungai; 54. Landai vicinity. Tembeling, Sungai. 55. Japura. 1 2. Sungai Tekam Forestry Concession. 56. Indragiri district. 13. Berapit, Bukit, east; Kampong Cherok Paloh; Kam- 57. Kateman, Sungai. pong Tanah Puteh; ; Pekan vicinity. 58. Indragiri, Sungai. 14. Kertau, Bukit. 59. , ca. 60 km NNW. 1 5. Benom, Gunong; Benom, Gunong, NE slope; Kuala 60. Jambi, ca. 90 km ENE. Lompat; Kuala Lompat Post, 0-1 km NW and 0- 61. Musi, Air, ca. 30 km and 1 5 km above Palembang, 2 km W; Lompat, Sungai, ca. 3 km and 4 km ENE near Palembang, and ca. 12 km, 24 km, 36 km, 48 of Kuala Serloh; Lompat, Sungai, ca. 3 km, 4 km, km, 60 km, 72 km, and 84 km below Palembang; and 6 km W of Kuala Lompat Post; Patong, Bukit, Palembang district; Palembang, ca. 30 km, 40 km, ca. 1 km W; Tens, Sungai. 45 km, 50 km, 65 km, 70 km, 80 km, and 100 km 16. Keroh Forest Reserve; Telok Anson. N, ca. 30 km and 60 km NNE, ca. 40 km, 60 km, 17. . 65 km, and 70 km NNW, ca. 45 km and 60 km 18. Changkat Mentri; Cherakah, Bukit; Dusun, Sungai; NW, and ca. 35 km SW. Kampong Rantau Panjang; Kampong ; 62. Harileko, Batang, ca. 12 km, 24 km, 36 km, 48 km, Kuala ; estuary; Lima Belas 60 km, 72 km, 84 km, and 96 km above mouth; Estate; Pacific Tin; Port Swettenham vicinity; Tengi, Musi, Air, ca. 15 km, 30 km, 45 km, 60 km, 75 km, Sungai; Tunggal, Bukit. and 90 km above mouth of Batang Harileko; Musi, 19. Bunga Buah; Damansara; ; Gaik Liew Air, ca. 50 km, 70 km, and 85 km above Palembang. Estate; Kelang Road; vicinity; Nan- 63. Palembang, ca. 95 km and 100 km SW and ca. 90 as, Bukit; ; Ulu Gombak Forest Re- km WSW. serve. 64. Lubuklinggau and vicinity and ca. 1 5 km N. 20. Pasoh Forest Reserve. 65. Lubuklinggau, ca. 40 km NNW. 21. Lesong. 66. Barisan Selatan. 22. Endau, Sungai, vicinity; ; Tanjong 67. Kelabong, Bukit, vicinity. Panjair. 68. Jeleket vicinity and ca. 10 km W. 23. Telapak Burok, Gunong. 69. Bengkulu. 24. Menyala, Sungai; ; Tanjong Tuan. 70. Bengkulu, ca. 40 km ENE. 25. vicinity; Nuri Valley. 71. Sanggul, Bukit. 26. Melaka. 72. Lubuklinggau, ca. 60 km SE. 27. Nyalas. 73. Baturaja, ca. 70 km WNW and ca. 75 km NW; 28. Selai, Sungai. Lahat. 29. Sekol, Sungai. 74. Baturaja, ca. 20 km and 30 km NNE. Absence reports 75. Palembang, ca. 60 km, 65 km, and 70 km SW. 76. Muaradua. a. Berhala, Pulau. 77. Suka Bandjar, NE and SE. b. Jarak, Pulau. 78. Bukit Barisan Selatan, Taman Nasional. c. Sembilan Kepulauan. 79. Wonosobo. d. Babi, Pulau. 80. , Propinsi. e. Bangkaru, Pulau. 81. Kotabumi. f. Siberut, Pulau.

82. Moro Batin and SSE; Telukbetung. g. Sipura, Pulau. 83. Sukadana. h. Pagai Utara, Pulau. 84. Sungsan, ca. 50 km N; Kambas, Wai. i. Pagai Selatan, Pulau.

85. Sungsan, ca. 20 km NNE. j. Enggano, Pulau. 86. Kiambang. k. Anak Krakatau, Pulau; Rakata, Pulau; Rakata- 87. Kalianda. kecil, Pulau; Sertung, Pulau.

1. Sebesi, Pulau. West (WM) m. Datuk, Pulau.

1 . Kampong Titi Tinggi. n. Temaju, Pulau. 2. , Sungai. Absence report not represented because of map con- 3. Perangin, Bukit. gestion: Sanglang-besar, Pulau (0°36'-0°38'N, 103°41'- 4. Pong. 103°42'E). 5. Larut, Daerah; Ulu .

FIELDIANA: ZOOLOGY (Sab)

Fig. 2C. Detail map of non-Philippine localities of Macaca fascicularis, southeastern quadrant. (Philippine lo- calities are mapped in Fooden, 1991, p. 2.) For documentation, see Gazetteer (Appendix 2). Abbreviations in parentheses are those used in Gazetteer locality codes.

Brunei (B) 4 1 . Baluran Game Reserve. 42. Banyuwangi. 1. Bandar Seri Begawan; area; Kampong 43. South Banyuwangi Nature Park. Menuggol.

Unmapped Brunei locality: Kuala Belalong Field (K), including nearby eastern islands Studies Centre (4°33'N, 1 15°08'E; for details, see Gaz- etteer, Appendix 2). 1 . Cabang Panti Research Station; Gunung Palung Na- ture Reserve; Kampung Baru Study Area. Java (J), including nearby islands 2. Ambawang, Sungai. 1-24. See Figure 2B. 3. Senoeang. 25. Candiroto. 4. Perbuah. 26. Semarang. 5. Roema Manoeal. 27. Karimunjawa, Pulau; Kemujan, Pulau. 6. Semitau. 28. Pangonan. 7. Sintang. 29. Gedangan. 8. Riam. 30. Ngawi. 9. Sekonyer Kan an River. 31. Tamansari. 10. Reserve.

32. Tulungagung. 1 1 . Katingan, Sungai, right bank, 1 40 km above mouth. 33. Kawarasan; Kediri district; Margomulio, Gunung; 12. . Manggis. 13. Katingan, Sungai, left bank, 160 km above mouth. 34. Wonokojo. 14. Katingan, Sungai, left bank, 200 km above mouth. 35. Meru Betiri Nature Park. 15. Kahayan, Sungai, right bank, 180 km above mouth 36. Mendit; Tengger, Pegunungan. of Sungai Rungan. 37. Pulau Doem area. 16. Liang Koeboeng. 38. Bawean, Pulau. 17. Putussibau. 39. Madura, Pulau. 18. Poelau. 40. Kangean, Pulau. 19. Busang, Sungai, left bank, 6 km above mouth, and

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS right bank, 1 km above mouth; Julai, Sungai, left Lesser Sundas (LS) bank, 1 km below mouth of Sungai Busang and 2 Bali, Pulau, including nearby islet: km and 4 km above Muarajuloi; Julai, Sungai, right 1. Bali Barat National Park; Banjoe Wetan; Gilima- bank, 1 km above mouth of Sungai Busang, 3 km nuk; Pulaki, Tanjung; Sendang; Trima, Teluk. below mouth of Sungai Busang, and 6 km above 2. Jembrana. Muarajuloi; Murung, Sungai, left bank, 1 km below 3. Bangli, near; Batur, Gunung; Botanical Gardens; mouth of Sungai Danau, 2 km above mouth of Sun- Bratan, Danau; Bratan, Gunung; Buyan, Danau- gai Beriwit, 2 km above mouth of Sungai Turusan, Danau Bratan region; Catur, Gunung; Desa Poet- 2 km below mouth of Sungai Turusan, and 8 km jang; Gitgit; Kukuh; Sangeh; . above Muarajuloi; Murung, Sungai, left bank, at 4. Penida, Nusa. mouth of Sungai Beriwit, 1 km below mouth of Sungai Beriwit, 5 km above mouth of Sungai Tu- Lombok, Pulau: rusan, and 1 km below mouth of Sungai Turusan; 5. Pengsong, Gunung; Pusuk, Gunung; Suranadi. TelukJolo, 12 km N. 6. . 20. Kahayan, Sungai, left and right banks, 1 20 km above 7. Pusuk forst; Rinjani, Gunung; Sewela. mouth of Sungai Rungan. Sumbawa, Pulau, including nearby islet: 2 1 . Kahayan, Sungai, left bank, 60 km above mouth of 8. Ai Beta; Batudulang; Maman; Semongkat. Sungai Rungan. 9. Moyo, Pulau. 22. Rungan, Sungai, right bank, 50 km above mouth. 10. Ampang area. 23. Rungan, Sungai, right bank, above mouth; Kahay- 1 1 . Dompu; Oo vicinity. an, Sungai, left bank, above and below mouth of 12. Ntori; Raba; Rite. Sungai Rungan, and right bank, below mouth of Sungai Rungan. Flores, Pulau, including nearby islets: 24. Kapuas, Sungai, right bank, 25 km above mouth. 13. Ginggo, Teluk, Pulau Rinca; Kode, Nusa; Mangia- 25. Barito, Sungai; Kambang, Pulau; Kaget, Pulau. tan, Pulau; Mbura, Pulau Flores; Nanga Look, Pulau 26. Pelaihari; Pleihari Tanah Laut Game Sanctuary. Flores; Seraya Besar, Pulau. 27. Karangintan. 14. Sano, Wai. 28. Rantau. 15. Rana Mese. 29. Hantakan; Telang. 16. Bari. 30. Tanjung. 17. Reo. 31. Buntok. 18. Pota. 32. Kampong Hadjak; Muaratewe. 19. Sika. 33. Purukcahu. 20. Solor, Pulau. 34. Tibang, Mt. 21. Adonora, Pulau. 35. Badang. Sumba, Pulau: 36. Sungai Kayan-Sungai Mentarang Nature Reserve. 22. Payeti-Kambaniru. 37. Sembakung, Sungai. 23. Mao Marroe. 38. Long Peleben; Long Pangian. 39. Kaboerau. Timor, Pulau, including nearby islets: 40. Karangtigau, Tanjung. 24. Kambing, Pulau; Oeassa vicinity, Pulau Semau. 41. Maratua, Pulau. 25. Amarassie. 42. Berau, Sungai; Birang, Sungai. 26. Benu; Bokong; Kuatnana; Lelogama; Timau, Fatu. 43. Merah. 27. Nikiniki. 44. Karangan, Sungai; Pelawan, Sungai. 28. Mutis, Gunung. 45. Kariorang. 29. Fatuboi. 46. Nature Reserve, northeast corner; Sengata; Philippines Sengata, Sungai, Kutai Reserve; Sengata, Sungai, at Mentoko camp and 1 km, 2 km, 3 km, 5 km, 7 km, Unnumbered locality: Bongao Peak, Bongao Island 9 km, 1 1 km, and 1 3 km below; Sengata, Sungai, (5°01'N, 1 19°45'E; for details, see Gazetteer, Appendix below Sengata village. 2). 47. Juyan, Sungai, right bank, near mouth; Mentoko Sabah (Sab) Research Center; Sengata, Sungai, right bank, 1 km

below mouth of Sungai Nubung. 1 . Padas Bay. 48. Kembangjanggut. 2. Rayoh. 49. Goson Djerong, near; Jembayan, Sungai; Karang- 3. Papar. mumus, Sungai; Loa Bambam; Mahakam, Sungai, 4. Bongkabong; Rugading. left bank, 1 km and 4 km above Sebulu; Mahakam, 5. Talibang; Tuaran; Tuaran-Kampong Tenghilan Sungai, north bank, above ; Mahakam, Road. Sungai, right bank, 8 km above Sebulu and 2 km 6. Keningau. below Sebulu; Tangarveng Island. 7. Garau; Kampong Bundu Tuhan; Kampong Kiau; 50. Sepaku, Sungai. Kampong Kiau-Tenampok Pass; Kenokok; Kiau-

5 1 . Klumpang, Teluk. lan; Kinabalu, Mount; Kinabalu National Park; 52. Sebuku, Pulau. Lumu Lumu; Ranau; Tempasuk, Sungai; Tenom- 53. Matasiri, Pulau. pok; Tinonkok.

10 FIELDIANA: ZOOLOGY Sex and Age Variation Geographic Variation

Dorsal pelage color is generally similar in adult, Dorsal Pelage Color Saturation— To in- subadult, and juvenile males and females, but pel- vestigate geographic variation in dorsal pelage col- age in adult males tends to be longer and sleeker or saturation in M. fascicularis, specimens ex- than in other age/sex classes. In newborn infants, amined have been compared with standard skins, dorsal pelage is blackish (on head, trunk, limbs, as in a previous study of Philippine M. fascicularis and tail), and the facial skin is bare, unpigmented, (Fooden, 1991, p. 5). In the previous study, the and pinkish (Dang, 1977, p. 11; Aldrich-Blake, range of standard saturation index (SI) values ex- 1980, p. 147; Crockett & Wilson, 1980, p. 170; tended from 1 .0 (pale yellowish brown) to 3.0 (dark Fittinghoff & Lindburg, 1980, p. 189; Wheatley, brown). In the present study, this range extends

1 1982, p. 205; Koyama, 1985, p. 106). The gradual from 0.5 (buffy; fmnh 9965 , Thailand: Kata Taek) transition from blackish neonatal pelage to grayish to 4.0 (blackish; usnm 114168, Indonesia, P. Si- or brownish postneonatal pelage begins at about meulue: Sibaboh, Lugu) (Fig. 6). Skins of infants age 2-3 months, which is the age when deciduous (specimens with deciduous teeth only) are exclud- canines and first molars erupt (Table 1 3) and in- ed from the analysis. Also excluded is the aber- fants begin to obtain some of their food indepen- rantly albinistic (pale ochraceous) skin of an adult dently (see below, Reproduction). The change of female (amnh 107094) collected at Perbuah, pelage color occurs first on the appendages, then northwestern Kalimantan; eight other specimens on the trunk, and last on the crown. In specimens from the same locality are normally colored. examined, the transition is complete well before In the core area of distribution of M. fascicularis eruption of the permanent first molars (age ca. 15 (see above, Geographic Distribution), dorsal pel- mo; Table 14). age color in 640 postinfantile specimens varies

8. Balambangan, Pulau. 10. Jumpit. 9. Banggi, Pulau, south, and Karakit; Maliangin Besar, 1 1 . Lanjak-Entimau Orang-Utan Sanctuary (proposed). Pulau. 12. Belaga. 10. Sabor, Pulau Banggi. 13. Dulit, Bukit.

1 1 . Malawali, Pulau. 14. Selikan, Bukit. 12. Sabatik, Pulau. 15. Similajau National Park. 13. Ulu Segama Reserve. 16. Niah Caves. 14. Bukit Garam. 17. Niah National Park. 15. Lokan, Sungai. 18. Baram, Batang; Baram district; Miri; Miri, Sungai; 16. Betotan; Lungmanis Station. Miri district. 17. Labuk Road; Sandakan, 8 mi (= 13 km) W; Sibuga 19. Long Ekang. Besar, Sungai. 20. Kalulong, Bukit.

18. Sapagaya Forest Reserve. 2 1 . Gunong Mulu National Park; Melinau Gorge; Mulu, 19. Abai; Kinabatangan, Sungai; Trusan Kinabatangan. Gunung. 20. Dewhurst Bay; Kretam Besar, Sungai; Kretam Kechil, 22. Punang, Sungai. Sungai. Absence reports 21. ; Segama, Sungai. 22. Darvel Bay. a. Manuk Manka Island, Philippines (see Fooden, 1991, 23. Tawau Hills National Park. p. 40). 24. Tawau. b. Masalembu Besar, Pulau. c. Sangeang, Pulau. Sarawak (Sar) d. Banta, Pulau; Bugis, Gili; Lawadarat, Gili; Lawa- 1. Samunsam Wildlife Sanctuary. laut, Gili; Longos, Nusa; Misa, Pulau; Papagaran- 2. Penrissen, Gunung. besar, Pulau; Sababi, Pulau; Sabajor Besar, Pulau; 3. Paku Cave. Sebolon Besar, Pulau; Siaba Besar, Pulau; Tatawa, 4. Bako National Park; Kuching; Kuching, 10th mile; Pulau. Sarawak, Sungai, mouth. e. Kelapa, Pulau; , Pulau; Langkoi, Pulau; 5. Bukar, Sungai; Entawa, Tanjong. Lengah, Pulau; Mbarapu, Nusa; Motang, Gili; Pa- 6. Pelandok, Sungai. dar, Pulau; Padar-kecil, Pulau. 7. Simunjan, Sungai. f. Lomblen, Pulau. 8. Lingga. g. Pantar, Pulau. 9. Paku, Saribas. h. Alor, Pulau.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 11 120

Fig. 3. Approximate extent of dry land in Southeast Asia during last glacial maximum, ca. 18 Ka; estimate based on present-day 120-m bathymetric line (Heaney, 1986, p. 137; 1991b, p. 147; U.S. Defense Mapping Agency bathymetric charts Nos. 63020, 1993; 63400, 1990; 71000, 1975a; 71006, 1975b; 93036, 1985; 94004, 1986).

= = from buffy (SI 0.5) to dark brown (SI 3.0) a pale yellowish brown adult male collected in

1 (Tables , 2); relatively few of these specimens east-central Sumatra (usnm 1131 69, Indragiri, = (4.5%) are darker than medium brown (SI 2.0). Sungai) matches an adult male collected in western Pelage color in core-area specimens averages pal- Java (usnm 156291, Bantargebang). = est in the Indochinese Peninsula (mean SI 0.99, Fringing-island specimens examined include 49 1 = n 93), becomes darker southward and reaches postinfantile skins— 234 collected in 71 shallow- = maximum saturation in Sumatra (mean SI 1.38, water islands and 257 collected in 27 deep-water = = = n 178) and Borneo (mean SI 1.47, n 239), islands (Appendix 3). Although dorsal pelage sat- and then becomes somewhat paler farther south- uration in most fringing-island specimens broadly = = ward in Java (mean SI 1.26, n 77). Within overlaps that in core-area specimens, mean SI in the Indochinese Peninsula, pale specimens are fringing-island samples tends to exceed that in particularly concentrated near the northwestern neighboring core-area samples (53 of 71 shallow- margin of the species range, north of 1 4°N between water fringing-island samples, 23 of 27 deep-water = = 98°E and 102°E (mean SI 0.73, n 42). In all fringing-island samples). parts of the core area, variation in dorsal pelage Strikingly dark fringing-island samples (con- saturation is relatively great, and overlap in sat- spicuously darker than any other M. fascicularis uration between samples from different parts of samples examined) have been collected in six deep- the core area is extensive. For example, a pale water islands that constitute three geographic units: golden brown adult male collected in western (1) Nicobar Islands (Katchall, Little Nicobar, Great = Thailand (fmnh 99642, Ban Tamrong Phato) al- Nicobar), northwest of Sumatra (mean SI 3.9, = most perfectly matches an adult male collected in n 8); (2) P. Simeulue and P. Lasia, west of Su- = = southern Sumatra (amnh 102765, Muaradua), and matra (mean SI 4.0, n 15); and (3) P. Maratua,

12 FIELDIANA: ZOOLOGY Fig. 4. Core area, shallow-water (< 120 m) fringing islands, and deep-water (> 120 m) fringing islands inhabited by Macaca fascicularis (cf. Fig. 3). For names of islands, see Figure 2 and Fooden (1991, p. 2).

= = = = east of Borneo (mean SI 4.0, n 4). The pe- Thailand (SI 2.5, n 1), and P. Belitung, be- = = ripheral and disjunct distribution of these intense- tween Sumatra and Borneo (mean SI 2.3, n ly saturate populations of M. fascicularis was not- 2). ed previously by Kellogg (1944, p. 76). Dorsal Dorsal Pelage Erythrism— Dorsal pelage er- pelage also is relatively dark in many islands of ythrism in M. fascicularis has been investigated the Philippine Archipelago (deep-water); in west- by comparing specimens examined with selected ern, northern, and eastern Philippine islands, mean standard specimens, in a procedure parallel to that

SI varies from 2.4 to 3.0 (cf. Fooden, 1991, p. 5). used in investigating saturation (see above). The A brief, equivocal comment alluding to pelage sat- range of standard erythrism index (EI) values ex- = uration in M. fascicularis in deep-water P. We, tends from 1.0 to 3.0, as follows: EI 1.0, noner- north of Sumatra, has been published by Scheff- ythristic, color of pale hair bands pale yellowish = rahnetal. (1994, p. 136). (fmnh 99651, Thailand: Kata Taek); EI 2.0, The darkest shallow-water fringing-island sam- moderately erythristic, pale hair bands golden ple was collected in P. Karimunjawa, north ofJava (fmnh 33505, Vietnam: Ho Chi Minh City); and = = = (mean SI 2.7, n 6; cf. Sody, 1949, p. 132). A EI 3.0, intensely erythristic, pale hair bands ru- shallow-water fringing-island sample from Ko fescent (fmnh 62913, Philippines: Palawan, Puer- Khram Yai, in the Inner Gulf of Thailand, also is to Princesa). Specimens marginal in erythrism be- = = conspicuously darker (mean SI 1 .8, n 10) than tween primary standards have been assigned EI neighboring core-area specimens (cf. Kloss, 1919c, values of 1 .5 or 2.5. Infant skins are excluded from p. 347). Inconclusively small shallow-water island the analysis. samples of dark specimens have been collected in In core-area samples, erythrism in M. fascicu- P. Redang, east of the Malay Peninsula (mean SI laris averages relatively high in Sumatra and in = = 2.5, n 2), Ko Na Ka Yai, west of peninsular the adjacent Malay Peninsula, south of 10°N (Ta-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 13 Fig. 5. External characters in Macaca fascicularis—left, Thailand: Lop Buri, Sarn Pra Karn, adult male; right, Indonesia: P. Bali, Sangeh, adult female and adult male.

bles 3, 4). Of 645 postinfantile core-area skins ex- specimens collected in 25 deep-water islands (Ap- = amined, 24 are intensely erythristic (EI 3.0); all pendix 4). In shallow-water fringing-island sam- of these intensely erythristic specimens originated ples, mean EI tends to exceed that in neighboring either in Sumatra or in the adjacent Malay Pen- core-area samples (mean EI greater in 48 fringing- insula. Within Sumatra and the Malay Peninsula, island samples, mean EI less in 1 6 fringing-island intense erythrism apparently is an individual vari- samples, mean EI equal in 7 fringing-island and able, not a local variable; in local samples (n > 1) core-area samples). Geographic variation of ery- that include intensely erythristic specimens, the thrism in shallow-water fringing-island samples frequency of such specimens varies from 0. 1 2 (Su- generally parallels that in neighboring core-area = = matra: Batangkuis; n 8) to 0.50 (West Malaysia: samples: 1 3 intensely erythristic specimens (EI = = Port Dickson; n 4). Intense erythrism (EI 3.0) 3.0) have been collected in 10 shallow-water fring- occurs in relatively pale core-area specimens (zrc ing islands (Bintan, Bulan, Durian, Karimun, and = 4-075, West Malaysia: Port Dickson, SI 1.0) as Nguwal, in the Archipelago, east of Sumatra; well as in darker specimens (zsbs C- 1 76, Sumatra: Redang, Siantan, and Tioman, east of the Malay = Kampong Baru, SI 2.5). The high incidence of Peninsula; Pinang [1] and Pintu Gedong, west of intense erythrism in M. fascicularis in the Malay the Malay Peninsula; and Singapore); all 10 shal- Peninsula has been indicated previously by Kloss low-water fringing islands from which intensely (1919c, p. 347) and Chasen (1940a, p. 67). Aggi- erythristic M. fascicularis specimens have been marangsee (1992, p. 119) reported on erythrism collected are adjacent to Sumatra or the Malay in a provisioned population of M. fascicularis at Peninsula, where intensely erythristic core-area Kosumphi Forest Park, northeastern Thailand. specimens have been collected. In these 1 fringing In fringing-island samples, erythrism data are islands, the frequency of intensely erythristic spec- = available for 234 postinfantile specimens collected imens varies from 0.09 (Tioman, n 1 1) to 1.00 = in 7 1 shallow-water islands and 2 1 8 postinfantile (Redang, n 2).

14 FIELDIANA: ZOOLOGY Fig. 6. Dorsal pelage color saturation extremes in Macaca fascicularis— fmnh 99651 (left), adult male, Thailand: Kata Taek, buffy (saturation index value, 0.5); usnm 1 14168 (right), adult male, Indonesia: P. Simeulue, Lugu Sibaboh, blackish (SI value, 4.0). Scale bar 10 cm.

In deep-water fringing islands, erythrism is rel- crown is darker than the back (Fig. 7) as a result atively high in most islands of the Philippine Ar- of reduction or absence of pale bands in crown chipelago (Appendix 4) and is particularly high in hairs. Darkness of the crown in these specimens Palawan, as noted previously (Fooden, 1991, p. varies from a dilute blackish wash to an irregular 8). Erythrism is low in the very dark insular pop- blackish streak to a clearly defined blackish patch. ulations west of Sumatra (Appendix 4), and ery- In dark-crowned specimens, the periphery of the thrism also is generally low in the deep-water Less- crown patch usually is darker than the center. er Sunda Islands, as in nearby Java (Tables 3, 4). Of 648 core-area postinfantile specimens ex- Crown Color— As previously indicated, the amined, dark crowns are clearly defined in 1 and crown in M. fascicularis usually is more brightly weakly defined in 13 (Table 5, Appendix 5). Thir- colored than the back; however, in 63 of 1,103 teen of these 14 dark-crowned specimens, includ- postinfantile specimens examined (Table 5), the ing the one with a clearly defined dark crown, were

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 15 1 Table 1. Dorsal pelage color saturation in core-area samples of Macaca fascicularis : summary of variation (cf. Table 2). 1 Table 3. Dorsal pelage erythrism in core-area Macacafascicularis : summary of variation (cf. Table 4). ] Table 5. Crown color pattern in Macaca fascicularis : summary of variation. For details, see Appendix 5. INFRAZYGOMATIC TRANSZYGOMATIC

Gervais, 1854

Pocock, 1939

Fig. 8. Infrazygomatic and transzygomatic lateral facial crest patterns in Macaco, fascicularis, as depicted by Gervais (1854, p. 87) and Pocock (1939, PI. V). In both depictions, note posteriorly directed preauricular hairs in the infrazygomatic pattern and anteriorly directed preauricular hairs in the transzygomatic pattern.

pale lateral facial crest in M. fascicularis is formed posteriorly from the posterior margin of the eye by the convergence of variably elongated poste- to the anterior margin of the ear (anteriorly di- riorly directed hairs of the anterolateral facial re- rected hairs are absent in this region); as a result, gion and anteriorly directed hairs of the postero- the infrazygomatic crest lacks the upper (preaur- lateral facial region. In most specimens examined, icular) portion of the transzygomatic crest. Al- this crest sweeps upward from near the angle of though the infrazygomatic pattern is relatively rare the jaw to the lateral margin of the crown, passing in M. fascicularis, it is common in M. mulatta, between the eye and ear (transzygomatic crest pat- where the transzygomatic pattern occurs infre- tern); less commonly, the crest is restricted to the quently (Stewart, 1933, p. 30). Transzygomatic and mandibular region and terminates superiorly in a infrazygomatic patterns can be distinguished in whorl low on the cheek (infrazygomatic crest pat- the hair tracts of infants as well as in postinfants. tern) (cf. Pocock, 1939, p. 78). In the infrazygo- In 819 specimens examined, the lateral facial matic pattern, unlike the transzygomatic pattern, crest pattern is transzygomatic in 728, infrazygo- hairs of the temporal region are smoothly directed matic in 83, and asymmetric (transzygomatic on

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 19 1 Table 6. Lateral facial crest pattern in Macacafascicularis : summary of variation. For details, see Appendix 6.

Table 7. External measurements and proportions in age/sex classes of wild-collected Macaca fascicularis.

Head and Relative tail Relative hind Relative 3 3 3 body length length foot length ear length Weight 2 Age/sex class (mm) (T/HB x 100) (HF/HB x 100) (E/HB x 100) (kg)

Fetuses Table 8. Head and body length in core-area Macaca fascicularis: summary of variation (cf. Fig. 10). 650

600

550-

500 oo

$ + O x x ~ 450- ..+_OJ< 4 E 01 E o| 00 o O) 400- -e-j c Males

O 350 X) T3C 550 CO

ca CD X 500

D

450- D

o I o o

400- &£ x.. « O.l

350

Females

300 10°S 5 10 15°N Latitude

O Borneo Java x Mainland + Sumatra

Fig. 10. Latitudinal variation of head and body length in adult core-area specimens of Macaca fascicularis (cf. Table 8).

Relative T in core-area specimens also is neg- x 100) with latitude is exaggerated relative to that atively correlated with latitude (Fig. 1 4). Since HB ofT. re- generally is positively correlated with latitude (Fig. Problematic T measurements have been et al. 1 97 1 1 cf. 1 10), the negative correlation of relative T (T/HB ported by Van Peenen ( , p. 34; 969, 24 FIELDIANA: ZOOLOGY Table 9. Summary comparison of head and body length in fringing-island (FT) and core-area (CA) samples of Macaca fascicularis. For details, see Appendix 7.

Shallow-water FI samples Deep-water FI samples Number

Categories of FI samples 650

600

550

500- .

~ 450 E E

B 400 c Males

O 350- X)

C 550 n CO a co XCD 500-1 o o O o o a o o o o

<* o < o ' 400 |e o. . LA b ' ^ OB

350 o j 0.

Females

300 10°S 10 15°N Latitude

Shallow-water fringing-island specimens Core-area specimens of Macaca Fig. 1 1 . Latitudinal variation of head and body length in adult shallow-water fringing-island specimens fascicularis compared with that in adult core-area specimens (cf. Table 9 and Appendix 7).

surements of dry skin). In another adult male col- p. 1559; 1968a, p. 121); these two monkeys, which lected in 1 9 1 3 at Bukit Nanas, T is normal (bm(nh) were associated with a troop of normal-tailed M. = = 1955.1512, T 577 mm, relative T 126.8%). fascicularis, were inferred to be hybrids of M. fas- Two additional short-tailed monkeys were ob- cicularis and M. nemestrina. served at Bukit Nanas in 1966 (Bernstein, 1966, In fringing islands, T data are available for small 26 FIELDIANA: ZOOLOGY 650

600

o 550 s o ^0 o o o 500 oo 0° A. . -0

o ol i&o ~ 4504 o <>: - E 9 A E . I o o O I O l A <>_. g) 400 Males o 350 X3 c 550 CO "O co CD X 500

450

400-

350

300 10°S 10 15°N Latitude

Deep-water fringing-island specimens c Core-area specimens

Fig. 12. Latitudinal variation of head and body length in adult deep-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens (cf. Table 9 and Appendix 7).

of adults samples collected in 57 shallow-water lected at similar latitudes (Fig. 15, Table 1 1); this islands and 22 deep-water islands (Appendix 8). parallels the general reduction of HB in shallow- In shallow-water fringing islands, T is generally water fringing islands relative to HB in core-area reduced relative to T in core-area samples col- reference samples (see above). In deep-water fring- FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 27 Table 10. Tail length and relative tail length in core-area Macaca fascicularis: summary of variation (cf. Figs. 13, 14).' 700 ol

| 650 D 1 i La

600

550

.".oo

450

400

350 c o 650

'c0 600

350

500

450

400- 350 150

5 15"N Latitude

O Borneo Java x Mainland + Sumatra

Fig. 14. Latitudinal variation of relative tail length in adult core-area specimens of Macaca fascicularis (cf. Table 10).

of these measurements decrease latitudinally (cf. maxillary concavities vary from weakly to strongly Figs. 10, 13). In the deep-water Lesser Sundas, as defined in both sexes. Canines in males are rela-

in Bali (cf. Fig. 17), HB and T tend to be iso- tively large. A median sagittal crest, formed by metrically reduced, relative to HB and T in Java. progressive convergence of the temporal lines, is prominent in many old males. During development from infancy to adult- hood, zygomatic breadth increases slightly more Cranial Characters rapidly than skull length, and rostral length in- creases much more rapidly than postrostral length Sex and Age Variation (Table 12; cf. N. Fujiwara, 1963, p. 57; Morimoto, (Figs. 19, 20) 1982, p. 98; Nanda et al., 1987, p. 217). Recent studies indicate that the conspicuous sexual di- In wild-collected M. fascicularis, greatest length morphism of R/PR in adult M. fascicularis prob- = of skull (GL) in adult males (1 18.7 mm, n 454) ably is a result of both a higher rate and a greater averages approximately 1 8% greater than in adult duration of allometric growth of the rostrum in = females (100.4 mm, n 439) (Table 12). Zygo- males (Ravosa, 1991, p. 408; Richtsmeier et al., matic arches are relatively narrow in both sexes; 1993a, p. 28); prenatal and postnatal growth pat- relative zygomatic breadth (ZB/GL x 100) aver- terns contribute differentially to development of ages 67.9 in 440 males and 65.9 in 428 females. adult cranial morphology (Richtsmeier et al., The rostrum is relatively long and narrow, partic- 1 993b, p. 322). Fluctuating asymmetry ofthe skull ularly in males; the rostral-postrostral ratio (R/ generally increases with age (Halgrimsson, 1993, PR x 100) averages 56.7 in 316 males and 47.6 p. 431). in 24 1 females. Supramaxillary ridges and lateral The sequence and timing of dental emergence 30 FIELDIANA: ZOOLOGY 700

| 650 1

600

550-

500-

450

400

350 700

15°N Latitude

Deep-water fringing-island specimens Core-area specimens

Fig. 16. Latitudinal variation of tail length in adult deep-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens (cf. Table 1 1 and Appendix 8).

2 gence in captivity may not be identical to that in and the last deciduous tooth (m ) apparently natural populations. emerges before age 1 months (Table 1 3); no sex- In captivity, the first deciduous tooth (i,) may ual differences have been observed in timing or emerge (i.e., penetrate the gingiva) before birth, sequence ofemergence ofdeciduous teeth (Spiegel, 32 FIELDIANA: ZOOLOGY Table 11. Summary comparison of tail length and relative tail length (T/HB x 100) in fringing-island (FT) and 1 core-area (CA) samples of Macaca fascicularis. For details, see Appendixes 8 and 9.

Shallow-water FI samples Deep-water FI samples Number Number of FI > CA > of FI > CA > 2 3 2 3 Categories of FI samples islands CA FI islands CA FI

Tail length

1 . FI sample includes both sexes a. Sexes in FI sample deviate concordantly from those in CA sample 15 11 b. Sexes in FI sample deviate discordantly from those in CA sample 1) Females, FI > CA; males, CA > FI 2 2) Females, CA > FI; males, FI > CA 1

2. FI sample includes only one sex a. Females b. Males Totals

All samples (including both sexes) 50

Both sexes 140 o

* ; ° si * I -jo i if

o . Si m 20- o. i 0° t no •& .c c V 100

® 90

to

DC 80

70-

60 10°S 5 10 15°N Latitude

Shallow-water fringing-island specimens o Core-area specimens

Fig. 17. Latitudinal variation of relative tail length in adult shallow-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens (cf. Table 1 1 and Appendix 9).

variation of skull length also is in general accord and one is north of Borneo (Sabah: P. Banggi, with Bergmann's rule, but skull length, like HB 7°09'N). (see above), tends to be somewhat smaller in these In deep-water fringing islands, a tendency to- islands than in core-area populations at similar ward insular dwarfism is evident only in the Lesser latitudes (Fig. 22, Table 16); this indicates a gen- Sunda Islands (8°32'-9°42'S) (Fig. 23, Table 16); eral tendency toward insular dwarfism in shallow- skull length variation in deep-water Lesser Sunda

water fringing islands (cf. Fooden & Albrecht, 1 993, Islands is similar to that in Bali, the only shallow-

p. 525). For four shallow-water fringing islands, water Lesser Sunda Island (cf. Fig. 22). Skull length samples are sufficiently large to establish that both in deep-water fringing islands tends to be greater females and males are significantly smaller than than in core-area reference samples in the Nicobar in core-area reference samples; these islands are Islands (6°58'-8°00'N), northwest of Sumatra; in P. Bintan (Indonesia: Riau Archipelago, 1°09'N), P. Simeulue (2°39'N) and nearby P. Lasia (2°10'N), P. Tioman (West Malaysia: east coast, 2°48'N; in- west of Sumatra; and probably in P. Maratua

cluding one aberrantly small male, usnm 101744, (2°15'N), east of Borneo (cf. Fooden & Albrecht, = GL 97.4 mm), Ko Kut (Thailand: southeast 1993, p. 531). In the Philippine Islands (5°24- coast, 1 1°40'N), and Bali (Indonesia: Lesser Sunda 1 8°3 1 'N), skull length tends to increase with lati- Islands, 8°15'S). In five shallow-water fringing-is- tude, extending the Bergmannian trajectory evi- land samples, skull length may be larger than in dent in core-area populations between the equator core-area reference samples; four of these islands and about 12°30'N (Fig. 23; Fooden, 1991, p. 10); are in the Java Sea, between Borneo and Java variation of skull length in Philippine samples col- (Indonesia: P. Karimunjawa, 5°51'S; P. Kemujan, lected north of about 12°30'N does not follow the 5°51'S; P. Bawean, 5°48'S; P. Matasiri, 4°47'S), anti-Bergmannian trajectory evident in core-area 34 FIELDIANA: ZOOLOGY 150

Both sexes 140 O

130 3O vo X 120 : o a^i * ICO * 1— <£ ^t o, \ M lo t 110- .o°x> 1

|o = '. o 100 o- 00

0) 90

& 80

70-

60 10°S 5 10 15°N Latitude

Deep-water fringing-island specimens o Core-area specimens

Fig. 18. Latitudinal variation of relative tail length in adult deep-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens (cf. Table 1 1 and Appendix 9).

samples collected at similar latitudes in the Isth- individuals livetrapped in Thailand; one of these mus ofKra and Indochinese Peninsula. Skull length samples originated north of the Isthmus of Kra (< is aberrantly small in one male specimen collected 7 localities), and the other originated south of the = at Mahayahaya (13°55'N; GL 109.3 mm), Lu- Isthmus of Kra (2 localities). Each of the other zon, Philippines. three samples consists of48 individuals, imported, respectively, from Malaysia, Indonesia, and the Philippines. For the last three samples, no further details are available concerning the regions or is- Molecular Biology and Genetics lands of origin of individuals in the samples or concerning possible geographic heterogeneity

Mitochondrial DNA within the samples (Harihara et al., 1986, p. 357; 1988, p. 125); this is unfortunate, because Malay- In a unique and valuable series of studies, Ha- sia, Indonesia, and the Philippines each include rihara et al. (1986, p. 357; 1988, p. 1 18; 1991, p. widely dispersed landmasses inhabited by isolated 611) have employed five restriction enzymes to populations of M. fascicularis. investigate restriction fragment length polymor- In Table 1 7, the six sample areas of Harihara et phism in the mitochondrial DNA (mtDNA) of six al. are arranged in approximate geographic order. geographic samples of M. fascicularis (Table 1 7). They are: (1) Vietnam and Cambodia, both east One of these samples consists of five captive-born of (2) Thailand north of the Isthmus ofKra, which individuals— one born to a female imported from obviously is north of (3) Thailand south of the Vietnam and four born to females imported from Isthmus of Kra; the latter sample area is north of Cambodia. Two samples consist of a total of 98 (4) West Malaysia and northwest of FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 35 Fig. 19. Skull of adult female Macacafascicularis, near topotype— amnh 102764, Indonesia: Sumatra, Muaradua. (Photographs by James L. Balodimas, Division of Photography, fmnh.)

(Sarawak and Sabah, both in Borneo); West and treatment of M. fascicularis mtDNA with each of East Malaysia, together considered one sample the five restriction enzymes yielded three to six area, are mostly north of (5) Indonesia; and Ma- distinctive digestion patterns (enzyme morphs), as laysia and Indonesia are southwest of (6) the Phil- follows: BamHl and EcoRl, three morphs each; ippine Islands. In the absence of information con- Hpal and Sstl, four morphs each; and Hindlll, cerning whether any members of the Malaysian six morphs (Table 17). In the 149 individuals in- and/or Indonesian samples originated in Borneo, cluded in these four samples, 21 different com- the geographic relationship of the Philippine sam- binations (mtDNA types) of these enzyme morphs ple to the Malaysian and Indonesian samples re- were detected. In the 98 individuals included in mains ambiguous. the two subsequently studied Thai samples, four For the four initially studied non-Thai samples, additional enzyme morphs and three additional

36 FIELDIANA: ZOOLOGY Fig. 20. Skull ofadult male Macacafascicularis, near topotype—amnh 106566, Indonesia: Sumatra, Bukit Sanggul. (Photographs by James L. Balodimas, Division of Photography, fmnh.)

mtDNA types were detected; at each locality in ples (Table 17; cf. Melnick & Hoelzer, 1993, p. 3). these two samples, all individuals were uniform Although the small Indochinese sample shares nei- in their mtDNA type (Harihara et al., 1991, p. ther of its two mtDNA types with other samples, 611). it resembles the Malaysian sample in its high fre- Except for the peculiar mtDNA type in the Thai- quency ofSstl morph 2. Mitochondrial DNA types land south ofthe Isthmus ofKra sample (Harihara in the Thailand north ofthe Isthmus ofKra sample in the Indochinese and et al., 1991, p. 612), the distribution of enzyme are closely related to those et 1 99 1 6 1 morphs and mtDNA types in samples generally Malaysian samples (Harihara al., , p. 2). conforms to the geographic relationships of sam- The Indonesian sample (10 mtDNA types), which

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 37 1 2 Table 12. Cranial measurements and proportions in age/sex classes of wild-collected Macaca fascicularis. 14. teeth in 5 in Table Schedule of emergence of permanent captive Macaca fascicularis ; emergence age years. CORE AREA 145

~ 140 E E 135

X 130 e> Z 125 111 -J 120 - _l ^ 115 (A

LU 110 " _l < S 105 h

100

_ 120 - E E 115 -

I no h H O 2 105 LU 100 h

D 95

- UJ 90 _J < 85 -

80 SHALLOW-WATER FRINGING ISLANDS 45

40

35

I 30 - F CD z 25 - LU 20 " D 15 -

10 -

- 05 MALES

00

REGRESSIONS AND CONFIDENCE LIMITS TAKEN FROM CORE-AREA ANALYSES

20 h

15

I 10 - F CD 05 -

00 D XL 95 h CO

lu 90 - _l < FEMALES ^ 85 UJ LL 80

J I I L 10°S 8°S 6°S 4°S 2°S 0°2 N4 N6 N8°N 10°N 12°N 14°N 16°N 18°N 20°N LATITUDE

Fig. 22. Latitudinal variation ofgreatest length of skull in adult shallow-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens (Fooden & Albrecht, 1993, p. 528). For key to shallow- water fringing-island symbols, see Fooden and Albrecht (1993, p. 537). Solid lines indicate third-order polynomial regressions for core-area specimens; dashed lines indicate 95% confidence limits (see Fig. 21).

haplotype is relatively rare (0.02-0.08) and is re- Highly Repeated Restriction Patterns—A stricted to Malaysia, Indonesia, and the Philip- study by Crovella et al. (1994, p. 66) of highly pines. In M. mulatta and M. fuscata, two other repeated nuclear DNA sequences has revealed that, members of the fascicularis species group (Food- for nine enzymes employed, restriction patterns en, 1991, p. 2), the frequency of alpha-globin gene are indistinguishable in Sumatran, Javan, and haplotypes apparently is similar to that in Thai- Philippine M. fascicularis. Eight of the nine en- land north of the Isthmus of Kra. zymes—Alul, BamHI, EcoRl, Hindlll, Hinfl,

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 41 Table 16. Summary comparison of greatest length of skull in fringing-island (FI) and core-area (CA) samples of

1 Macaca fascicularis. For details, see Fooden and Albrecht (1993, Tables 2, 3).

Shallow-water FI samples Deep-water FI samples Number

Categories of FI samples

Table 17. Frequency (%) of mtDNA types revealed by five restriction enzymes in six geographic samples of Macaca fascicularis (Harihara et al., 1988, p. 121; 1991, p. 611).' Table 18. Frequency of alpha-globin gene haplotypes in geographic samples of Macaca fascicularis compared with frequency in samples of M. mulatta and M.fuscata (O. Takenaka et al., 1989, p. 87; A. Takenaka et al., 1991, p. 324; A. Takenaka & Takenaka, 1991, p. 632). Table 19. Blood proteins: summary of frequencies (%) of major alleles at polymorphic loci in 1 1 geographic samples of Macaca fascicularis (sample size indicated by italicized figures in parentheses). For details, see Appendix 10. a. Dendrogram excluding Mauritius sample compared with also were consistently and significantly different; dendrogram including Mauritius sample. in response to the test diet, serum lipid levels were Thailand, Cent. & SE 1 less consistently different. Thailand, SW 1 Bali c Lombok 3 Karyology Timor c Sumbawa D- Hirai et al. (1991, p. 619) studied the chromo- Sumatra somal G-banding pattern in 297 imported or cap- tive-born M. individuals that W Malaysia fascicularis origi- nated in Vietnam, Cambodia, Malaysia, Indone- Philippines sia, and the Philippines; the sample size for each Thailand, S 2 country is not specified. No significant differences Java in banding pattern were detected among samples Mauritius from these five countries, although one female from Consensus dendrogram. Vietnam exhibited possible interstitial hetero- Thailand, Cent. & SE 1 chromatin in one autosome. The chromosome rC Thailand, SW 1 number of M. fascicularis was confirmed to be 42. Bali

4: Lombok

Thailand, S 2 Diseases W Malaysia Sumatra

: Malaria - .'-'.' — Philippines

•• Java Of the seven species of malaria {Plasmodium) Sumbawa that have been recorded as natural parasites in -€ Timor macaques, five species naturally infect hi. fasci- 2 North of Isthmus of Kra. South of Isthmus of Kra. cularis (Fooden, 1994, p. 576). Of these, P. inui is asexual P. Fig. 24. Blood-protein dendrograms for geographic quartan (72-hr erythrocytic cycle); cy- samples of Macaca fascicularis: a. Dendrogram that ex- nomolgi, P. fieldi, and P. coatneyi are tertian (48- cludes Mauritius et 199 1 sample (Kawamoto al., , p. 600) hr cycle); and P. knowlesi is quotidian (24-hr cy- compared with dendrogram that includes Mauritius cle). Natural infections with all five ofthese species sample (Kondo et al., 1993, p. 177). b. Consensus den- have been recorded in West Malaysia, which ap- drogram. parently is the center of malaria infection in M.

fascicularis (Table 23). For P. fieldi, natural infec- 487 mg/dl). In response to the screening diet, high- tions are known only in West Malaysia. For P. density lipoprotein levels were significantly higher coatneyi, infections also have been reported in the in the Malayan sample (64 mg/dl) than in the Phil- southern Philippine islands of Palawan (question- ippine sample (5 1 mg/dl), and serum lipid levels able) and Cebu. For P. knowlesi, infections have

Table 20. Summary of allele frequencies at the DBP locus in geographic samples of Macaca fascicularis (Tanaka etal., 1989, p. 106; 1991, p. 131). Table 21. Allele frequencies at the complement C6 locus in three imprecisely localized geographic samples of Macaca fascicularis (Omoto et al., 1991, p. 604). Table 23. Geographic distribution ofmalaria species identified as natural parasites in Macacafascicularis samples. For details, see Fooden (1994, p. 576).

Malaria species (Plasmodium) Geographic origin of M. fascicularis sample cynomolgi fieldi coatneyi knowlesi sp.

Thailand Table 25. Simian T-lymphotrophic retrovirus, type 1: frequency of Macaca fascicularis individuals or groups positive for antibodies. Schaiketal., 1983b, p. 216; D. R. Vosetal., 1992, 5) in a highly disturbed area in Vietnam and ex- = p. 388). In response to danger, troops may flee ceptionally large in western Java (ca. 47, n 5 = either on the ground (Burma [Tickell, 1854-1875, troops) and Mauritius (77.5, n 2 troops, intro- p. [17], Great Nicobar I. [Das & Ghosal, 1977, p. duced population). Crockett and Wilson (1980, p. 265], Sumatra [Volz, 1912, p. 369; Ulmer in Mil- 149) suggested that troop size in Sumatra may be ler, 1942, p. 127], Lesser Sunda Islands [B. Rensch, smallest in forest, larger in inland pri- 1930, p. [17]; Mertens, 1936, p. 320]), in the can- mary forest, and largest in inland secondary forest opy (Thailand [Fooden, [1975], p. 100], West Ma- (cf. Ruiter, 1993, pp. 90-91), but the observed laysian [Bernstein, 1967, p. 204], P. Simeulue [Su- differences are acknowledged not to be statistically gardjito et aL, 1989, p. 200], Sumatra [Crockett & significant. In provisioned troops in Thailand, = = Wilson, 1980, p. 165], Kalimantan [Kurland, 1973, troop size (mean 76.9,n 33 troops) generally p. 250; Wheatley, 1980, p. 216]), or partly on the exceeds that in nonprovisioned troops (Aggimar- ground and partly in the canopy (Thailand [Food- angsee, 1992, p. 150). Solitary males, not associ- en, [1975], p. 100]). Accidental falls of this species ated with any troop, have been reported in Thai- out of trees have been observed in northern Su- land (Fooden, 1971, p. 24), Singapore (Furuya, matra (Karssemeijer et al., 1990, p. 286). Details 1965, p. 294), Sumatra (Wilson & Wilson, 1973, of locomotor behavior are discussed by Kurland p. 5; Rijksen, 1978, p. Ill; Norikoshi, 1984, p. (1973, p. 250), Fleagle (1980, p. 198), Cant (1988, 5), Sabah (Kurland, 1973, p. 253), Java (Hoog- p. 31), and Cannon and Leighton (1994, p. 512). erwerf, 1970, p. 408), and Mindanao (Hoogstraal, At night, M. fascicularis usually sleeps in tall, rel- 1951, p. 45). In sample areas studied, the pooled atively bare trees near a river (Singapore [Ridley, ratio of sexually mature males to sexually mature 1906, p. 142], Sumatra [Moszkowski, 1909, p. 24], females varies from 0.22 to 0.90 (Table 28); within Sarawak [Hornaday, 1910, p. 358], and numerous troops, known sex ratios vary from 0.14 to 1.67. later accounts, particularly Fittinghoff & Lind- During a 10-year period of investigation at Kua- burg, 1980, p. 190 -Kalimantan). la Lompat, West Malaysia, the size of local troops tended to remain relatively constant (Chivers &

Raemaekers, 1980, p. 25 1). However, troops often Swimming temporarily subdivide into smaller units while for- aging (West Malaysia [Aldrich-Blake, 1980, p. 147], Kali- Observers agree that M. fascicularis frequently Sumatra [van Schaik et al., 1983b, p. 214], enters bodies of water, apparently often for pleas- mantan [Kurland, 1973, p. 253; Rodman, 1978, ure, and that it is an excellent swimmer. In Burma, p. 470; Wheatley, 1978, p. 348]). At each of two a wounded male that was attempting to escape localities in Sumatra, a large troop has been ob- from humans swam about 50 m underwater (Tick- served to permanently subdivide into two un- Schaik et ell, 1854-1875, p. [18]). In a captive insular group equal-sized daughter troops (van al., of M. fascicularis in Japan, a young male that was 1983a, p. 174; Norikoshi, 1984, p. 6). defeated in a dominance fight apparently swam 100 m to a breakwater (Furuya, 1965, p. 325). Home Range, Day Range However, during the course of 3 years in the same insular most of about 60 individ- captive group, Reported home-range area in nonprovisioned uals that were driven dominant away by group troops varies from 1 2.5 ha in P. Simeulue to more members died none by drowning; apparently than 300 ha in West Malaysia (Table 29). Overlap reached the next nearest which is about island, between home ranges of adjacent troops generally 250 m distant and which not have been vis- may is slight, but the home range of a daughter troop ible to the also have escaping monkeys. Drownings may overlap extensively with that of the troop been reported in natural populations in coastal from which it split (van Schaik & van Noordwijk, Thailand (Aggimarangsee, 1992, p. 130). 1988, p. 80), and home ranges of troops in some small islands also may overlap extensively (Angst, 1973, p. 625). Although home ranges of some ad- Troop Size and Composition jacent troops are separated by rivers (van Noord- wijk & van Schaik, 1 985, p. 850), the home ranges Mean size of nonprovisioned troops of M. fas- of a pair of adjacent troops in Borneo overlapped cicularis generally varies from about 12 to 25 (Ta- on both sides of a river (Fittinghoff & Lindburg, ble 27). Mean troop size is exceptionally small (ca. 1980, p. 207).

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 51 Table 26. Recorded habitats of Macaca fascicular is.

Sample area Table 27. Recorded size of nonprovisioned troops of Macaca fascicularis. Table 28. Ratio of sexually mature males to sexually mature females reported in nonprovisioned troops of Macaca fascicularis. Table 29. Estimated home range and day range in troops of Macaca fascicularis. Table 30. Foods reported eaten by nonprovisioned troops of Macacafascicularis; dietary proportions are indicated where available. 1 2 Table 31. Daily waking-hour time budget estimates (%) for Macaca fascicularis. cal niches; M. fascicularis is rare in primary ev- sleeping trees and rudimentary interspecific ergreen rain forest, which is favored by M. ne- threats— a few by P. pygmaeus and one by M. mestrina (Rodman, 1973, p. 656; 1978, p. 476; fascicularis. 1991, p. 364; J. R. MacKinnon & MacKinnon, In West Malaysia, racket-tailed drongos (Dicru- 1978, p. 316; Crockett & Wilson, 1980, p. 162). rus paradiseus) have been observed following M. Macaca fascicularis and Trachypithecus crista- fascicularis through the canopy and catching in- tus frequently have been observed together in West sects stirred up by the monkeys' activity (Ridley, Malaysia (Furuya, 196 1-1 962b, p. 56; Bernstein, 1901, p. 105; cf. Fooden, 1969, p. 52). 1968b, p. 8; Lim & Sasekumar, 1979, p. 108), Sumatra (Wilson & Wilson, 1973, p. 5), and Java (Hoogerwerf, 1970, p. 408, including three inter- specific mixed groups). Relations between these Reproduction species reportedly are generally peaceful, but M. fascicularis often displaces T. cristatus. Hooger- Although copulations, pregnancies, and births werf (1970, p. 408) observed one violent inter- have been recorded throughout the year in natural specific fight in Java, during which an immature populations of M. fascicularis, long-term studies

T. cristatus ". . . fell to the ground torn to pieces." indicate that all three of these reproductive events Associations of M. fascicularis and Presbytis exhibit seasonal peaks in this species (Appendix

melalophos have been observed in West Malaysia 1 1). The timing of these reproductive peaks ap- (Bernstein, 1967, p. 201; J. R. MacKinnon & parently varies geographically and varies between MacKinnon, 1978, p. 319) and Sumatra (Snelle- years at the same locality. In a study in northern man, 1887, p. 10; Koyama, 1985, p. 105). Most Sumatra, the annual birth peak tended to occur accounts indicate that these associations are during or shortly after the local fruiting peak; in

peaceful . Koyama (1985, p. 105) reported that these years when fruit production was relatively low, the two species often share the same sleeping tree, but birth peak tended to be delayed by 2 or 3 months J. R. MacKinnon and MacKinnon (1978, p. 319) (van Schaik & van Noordwijk, 1985b, p. 538). In indicated that M. fascicularis displaces P. mela- Thailand, on the other hand, fertile matings— not lophos from feeding and sleeping trees. births— tend to peak shortly after the fruiting peak,

Bernstein (1967, p. 201) reported noncompeti- which occurs in April-June (Varavudhi et al., tive association of M. fascicularis with Trachypi- 1989a, pp. 221-222; cf. Tangpraprutigul & Var- thecus obscurus in West Malaysia, and Kurland avudhi, 1982, p. xci; Aggimarangsee, 1992, p. 129). (1973, p. 254) reported noncompetitive associa- In a 2- to 3-year study of captive M. fascicularis, tion with Presbytis aygula (= P. hosei) in Kali- annual variation of sperm concentration in semen mantan. Ridley (1895, p. 25) reported that leaf collected by electroejaculation was investigated monkeys in West Malaysia fought for more than (Okamoto, 1994, p. 27); results were inconclusive. 2 hours with M. fascicularis over occupation of a In natural populations of M. fascicularis, fe- feeding tree ("biting them [M. fascicularis] and males probably become reproductively active at throwing them out of the tree and into the river"). about age 3.5 years. A pregnant juvenile female, Banks (1978, p. 188) reported that M. fascicularis aged about 3.5 years, was collected in western leaf in often prevented monkeys Borneo from en- Thailand (fmnh 99646, with erupting C,, P3 , and 2 trees. cf. 1 1 females tering sleeping Peaceful association of M. M ; Table 14; Fooden, 97 , p. 24), and fascicularis and Nasalis larvatus in mangrove in Sumatra reportedly produce their first infants swamps in Borneo has been observed frequently at about age 4 years (Koyama, 1985, p. 106; van (Davis, 1962, p. 57; Kern, 1964, p. 185; Kawabe Noordwijk & van Schaik, 1987, p. 586; cf. Var-

& Mano, 1972, p. 216; Kurland, 1973, p. 254; avudhi et al., 1989a, p. 222). In captivity, females Macdonald, 1982, p. 117). may copulate as early as age 2.5 years (Chance et Noncompetitive association of M. fascicularis al., 1 977a, p. 6 1 9), and one captive female is known and Hylobates lar in West Malaysia was reported to have given birth at age 3 years (Timmermans by Bernstein (1967, p. 201). Macaca fascicularis et al., 1981, p. 121); however, fertility in captive and Pongo pygmaeus have been observed together females usually does not begin until about age 4 on 32 occasions in Sumatra (Rijksen, 1 978, p. 112) years (Spiegel, 1954, p. 230; Dang, 1983, p. 38). and on two occasions in Kalimantan (Fittinghoff Mean age at menarche in 43 captive females & Lindburg, 1980, p. 201); these observations in- was 2.5 ± 0.7 years (Honjo et al., 1984, p. 69; cf. clude peaceful interspecific sharing of feeding and Dang, 1983, p. 36). Mean length of the menstrual

58 FIELDIANA: ZOOLOGY = cycle in 28 captive females was 30.9 days (SD their sexual swelling or increasing redness of their = = 4.8 days, extremes 19-43 days, mode 28 days, perivulval skin (van Noordwijk, 1985, pp. 281, = n 595 cycles) (Dukelow, 1977, p. 34; cf. Joach- 287). In some females, sexual swelling or redden- imovitz, 1928, p. 462; Comer, 1932, p. 404; Spie- ing and copulations were observed during preg- gel, 1954, p. 235; Fujiwara et at, 1967, p. 506; nancy (cf. Deputte & Goustard, 1980, p. 96); fol- Kerber & Reese, 1 969, p. 976; Valerio et al., 1 969, lowing parturition, swelling was not observed for

1 1 p. 287; MacDonald, 97 , p. 374; Nawar & Hafez, several months.

1972, p. 45; Dang, 1977, p. 3; Goodman et al., Variably complete and generally concordant ac- 1977, p. 480; Zumpe & Michael, 1 983, p. 58; Hon- counts of copulatory behavior in M. fascicularis jo et al., 1984, p. 63; Wallis et al., 1986, p. 87). are available from natural populations (Tickell, Males in natural populations usually begin to 1854-1875, p. [17]; Furuya, 196 1-1 962a, p. 76; copulate at age 5-6 years, as subadults, after they Kurland, 1973, p. 258; M. A. R. Khan & Wahab, have left their natal troops (van Noordwijk, 1985, 1983, p. 104; Koyama, 1984, p. 31; 1985, p. 107; p. 288; van Noordwijk & van Schaik, 1985, p. van Noordwijk, 1985, p. 281), introduced free- 852; cf. Varavudhi et al., 1989a, p. 222). As pre- ranging populations (Poirier & Smith, 1974, p. viously indicated (see above, Natural History), 300; Sussman & Tattersall, 1981, p. 203), colony close inbreeding probably is rare or absent in nat- groups (Spiegel, 1954, p. 238; Goustard, 1961, p. ural populations of M. fascicularis. In Sumatra, 313; 1963, p. 710; 1968, p. 464; Furuya, 1965, p. subadult males that were still in their natal troops 313; de Benedictis, 1973, p. 1470; Angst, 1974, p. accounted for only 5 of 664 observed copulations. 52; Chevalier-Skolnikoff, 1975, p. 207; Emory & In Bangladesh, copulation attempts by immature Harris, 1978, p. 227; Deputte & Goustard, 1980,

males were actively thwarted by adult males (M. p. 85; Emory et al., 1980, p. 251; Shively et al., A. R. Khan & Wahab, 1983, p. 108). 1982, p. 375), and laboratory test pairs (Kanagawa In captive males, testicular descent occurs at age et al., 1972, p. 453; Kanagawa & Hafez, 1973, p. 2.5 years, and adult testicular size is achieved at 234; Michael & Zumpe, 1988, p. 379; Zumpe & age 4.0-4.5 years (Cho et al., 1973, p. 408; Chance Michael, 1983, p. 58; 1990, p. 148). Copulations et al., 1 977a, p. 6 1 9). Fertile copulations have been in this species have been observed throughout the recorded for three captive males at ages 3.5 years day, but evidence from troops in northern Su- (1 male; Honjo et al., 1984, p. 68) and 3.8 years matra suggests that copulations may be more fre- (2 males; Spiegel, 1954, p. 230). quent in the morning (ca. 60%) than in the after- At or near the onset of sexual maturity in fe- noon (van Noordwijk, 1985, p. 286; cf. de Bene- males, the bare skin between the root of the tail dictis, 1973, p. 1473; Sussman & Tattersall, 1981, and the anus swells prominently and the skin p. 203). Copulations occur both in trees and on around the vulva frequently reddens (Joachimov- the ground (Kurland, 1973, p. 258; Angst, 1974,

itz, 1928, p. 464; Corner, 1932, p. 408; Spiegel, p. 53; Sussman & Tattersall, 1981, p. 203; Koy-

1 954, p. 232; Nawar & Hafez, 1 972, p. 49; Fooden, ama, 1985, p. 117). [1975], p. 100; Emory et al., 1980, p. 250; van Copulations in M. fascicularis may be initiated Noordwijk, 1985, p. 281; Meishvili & Chalyan, by either sex. A female initiates copulation by star- 1986, p. 14; C. M. Anderson & Bielert, 1994, p. ing at or approaching a male and presenting her 285). This pubertal swelling and reddening ap- hindquarters or by stereotyped head or arm move- parently reaches its maximum about midway ments; in laboratory test pairs, female use of head through the menstrual cycle and diminishes a few or arm movements to initiate copulation peaked days later. In postpubertal females, sexual swelling near the middle of the menstrual cycle (Michael and reddening generally are less conspicuous and & Zumpe, 1988, p. 381). A male initiates copu- are highly variable in size and duration; in many lation by staring at or approaching a female and fully fertile females, sexual swelling and reddening raising her tail. Both female- and male-initiated

are slight or absent (cf. Pocock, 1 906, p. 558; 1 939, copulations usually proceed with examination by p. 78; Kurland, 1973, p. 258; Wilson & Wilson, the male of the perineum of the female; this ex- 1977, p. 211; Fittinghoff & Lindburg, 1980, p. amination may be visual, digital, olfactory, or, 189). However, in a carefully studied population rarely, lingual. If the female is acceptable (cf. de in northern Sumatra, sexual swellings generally Benedictis, 1973, pp. 1479-1480), the male mounts were larger during the peak copulation season (Jan- the female, by gripping her hips with his hands Jun.), and copulations and consortships of most and gripping her calves with his feet, and begins females tended to increase with increasing size of pelvic thrusting. In laboratory test pairs, mounts

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 59 Table 32. Copulatory behavior data in two Macaca fascicularis groups.

Provisioned natural group, Bali Colony group, USA

Ejaculatory copulations observed (N) 169 17

SME copulations (N) 139(82.2%) 8(47.1%)

1 Duration of SME mount (sec) 7.1 (2.6-12.0) 8.5 (6-14) Thrusts per SME mount 1 16.4(5-32) 17(14-21)

MME copulations (N) 30(17.8%) 9 (52.9%)

1 Duration of MME mount (sec) 6.4 (3.0-9.4) 6(2-11) Thrusts per MME mount 1 15(6-31) 13(4-29) Duration of MME sequence 1 (min) 7.1 (2-35) 41 (2-85) Mounts per MME sequence 1 2.4 (2-6) 3(2-6)

Reference Wheatley, 1991, p. 171 Shively et al., 1982, p. 376

Mean and extremes. and mounting attempts tended to increase slightly panied by a characteristic staccato female vocal- during the first half of the menstrual cycle and ization and sometimes by a characteristic male tended to decrease more precipitously during the vocalization (Goustard, 1963, p. 710; 1968, p. 470; second half (Zumpe & Michael, 1983, p. 66). Furuya, 1965, p. 313; Angst, 1974, p. 61; Poirier Ejaculation in M. fascicularis may occur either & Smith, 1974, p. 300; Chevalier-Skolnikoff, 1975, as a result of a single mount (single-mount ejac- p. 207; Deputte & Goustard, 1980, p. 83; Wheat- ulations [SME]) or as a result of a series of mounts ley, 1 982, p. 206; 1 984, p. 391; 1991, p. 171; Zumpe separated by dismounts (multiple-mount ejacu- & Michael, 1983, p. 60; van Noordwijk, 1985, p. lation [MME]). In a provisioned natural group in 281). At or near the time of ejaculation, the male Bali, 82% of observed ejaculatory copulations were pauses, and the female usually grasps one of the SME and 1 8% were MME; in a seminatural colony male's arms or legs and turns her face toward his group, 47% of observed ejaculatory copulations (cf. Kanagawa et al., 1972, p. 453). Shortly after were SME and 53% were MME (Table 32; cf. Fu- ejaculation, the male dismounts and the male and ruya, 196 1-1 962a, p. 76; van Noordwijk, 1985, female move apart. After copulation, a pair may p. 281). In laboratory tests, the frequency of SME remain associated in a consortship for a variable was higher in first copulations of test pairs than period, ranging from about 1 hour to 3 weeks (Fu- in second copulations (27% of 191 first copula- ruya, 1965, p. 314; Poirier & Smith, 1974, p. 301; tions, 6% of 154 second copulations) (Zumpe & Sussman & Tattersall, 1981, p. 203; van Noord- Michael, 1983, p. 60; cf. Kanagawa et al., 1972, wijk, 1985, p. 283); during a consortship, most, p. 453). In the natural and colony groups, the du- but not all, copulations of consortship partners are ration of both SME and MME mounts averaged with each other. In natural groups, females may about 7 seconds, and the number of pelvic thrusts copulate as often as 3-4 times per hour, through- per mount averaged about 15 (cf. Furuya, 1965, out the day, often with many different partners p. 313; Zumpe & Michael, 1983, p. 61); similar (Ruiter et al., 1992, p. 177). mount durations and thrust/mount rates have been Dominant males in M. fascicularis apparently reported for natural populations of M'. fascicularis participate in a disproportionately large share of in Bangladesh (10-15 seconds, 8-10 thrusts/mount; copulations. In northern Sumatra, the highest M. A. R. Khan & Wahab, 1983, p. 108) and Ka- ranking 2 of 7 males engaged in 52.7% of 659 limantan (5-7 sec, 4-12 thrusts/mount; Kurland, copulations (van Noordwijk, 1985, p. 285); in 1973, p. 258) and for a free-ranging introduced western Sumatra, 2 of 7 males engaged in 60.9% population in Angaur Island ("brief," 8-10 thrusts/ of 46 ejaculatory copulations (Koyama, 1984, p. mount; Poirier & Smith, 1974, p. 300). The num- 35; cf. Norikoshi, 1984, p. 12); in eastern Kali- ber of mounts in an MME sequence averaged about mantan, 1 of 6 males engaged in 50.0% of 48 cop- 3 in the natural and colony groups; the duration ulations (Wheatley, 1982, p. 207; alpha status of an MME sequence averaged about 7 minutes shifted during this study); and in Bali, 1 of 4 males in the natural group and about 4 1 minutes in the engaged in 50.9% of 126 copulations (Wheatley, colony group. 1991, p. 171). Similar high frequencies of copu- Copulation in M. fascicularis usually is accom- lation by dominant males have been reported in

60 FIELDIANA: ZOOLOGY colony groups (Furuya, 1965, p. 315; Shively et al., 1989, p. 151). Most births probably occur be- al., 1982, p. 377). In natural and colony groups, tween sunset and sunrise; in a captive group main- some low-ranking adult males were never ob- tained in a 14-hour light/ 10-hour dark environ- served to copulate. In laboratory test pairs, the ment, 90% of 1 52 observed births occurred during copulatory performance ofa subordinate male was dark hours, 1900-0500 (M. T. Suzuki et al., 1990, strongly inhibited by close proximity of a sepa- p. 252; cf. Ridley, 1906, p. 142; Spiegel, 1954, p. rately confined dominant male (Zumpe & Mi- 248; Erwin, 1977, p. 358; Banks, 1978, p. 188; chael, 1990, p. 154). In West Malaysia and Kali- Kemps & Timmermans, 1982, p. 84). mantan, aggression between males reportedly in- In captivity, 23 infants were nursed for an av- = = tensified shortly before periods of increased sexual erage of 15.1 months (SD 3.0 mo, extremes activity (Aldrich-Blake, 1980, p. 151; Wheatley, 9-22 mo) (Spiegel, 1954, p. 262; cf. Chance et al., 1982, p. 210). In troops observed in northern and 1977b, p. 31); following 9 of these 23 nursing pe- western Sumatra, subordinate males frequently at- riods, the mother's lactation continued without tempted to interfere with the copulations of dom- interruption through the nursing of one or more inant males, who, in turn, responded aggressively succeeding infants. In natural populations, infants (Norikoshi, 1984, pp. 9, 12; Koyama, 1985, p. begin to obtain some of their food independently 107; van Noordwijk, 1985, p. 283). The copula- about age 3 months (Karssemeijer et al., 1990, p. tory partners of high-ranking males in Sumatra 288). The mean duration of postpartum amen- generally tended to be females who were high rank- orrhea in captive nursing mothers was 6.6 months = = = ing, not young, and not nulliparous (van Noord- (SD 2.6 mo, extremes 2-13 n 28) (Spiegel, wijk, 1985, p. 289). 1954, p. 262; cf. Dang, 1979, p. 377; Cho, 1981, Paternity tests, using blood-protein analysis and p. 254; Honjo et al., 1984, p. 66; Varavudhi et al., DNA fingerprinting, have been conducted for 45 1989a, p. 221). offspring in three troops in northern Sumatra (Rui- In northern Sumatra, in six to seven nonpro- ter et al., 1992, p. 184; cf. Ruiter, 1993, p. 99; visioned troops that included 42-53 adult females,

Ruiter & van Hooff, 1993, p. 518; Ruiter et al., the annual birth rate (births/adult females) from 1994, p. 211); these tests indicate that dominant 1980-1981 to 1983-1984 was 0.45, 0.82. 0.30, males sire most of the offspring in natural troops, and 0.75; the overall annual birth rate for these 4 as would be expected from observed copulation years was 0.58 (van Schaik & van Noordwijk, frequencies. Of 42 offspring for which probable 1985b, p. 538). In western Sumatra, in three pro- fathers could be determined, 31 probably were visioned troops that included 22-3 1 adult females, sired by alpha males, 7 by beta males, and only 4 the annual birth rate from 1980 to 1984 was 0.73,

10 lower males. In ar- 1 0. 1 1 1 with an overall by approximately ranking 0.48, 0.8 , , and .00, average tificial colony groups, dominant males apparently of 0.63 (Koyama, 1985, p. 119). High and low do not maintain this reproductive advantage; pa- annual birth rates tend to occur in alternate years ternity tests in one colony revealed that, despite in both the nonprovisioned and provisioned troops. their higher frequency of copulations, dominant The limited available data suggest that a biennial males did not sire a disproportionately large num- birth cycle may be common in these Sumatran ber of offspring (Shively & Smith, 1985, p. 131). troops and that most females may tend to produce In a large captive group ofM. fascicularis, mean their young during the same alternate years. gestation length for 1,141 live births was 163.5 ± In provisioned troops at Ubud, Bali, where the 5.8 (SD) days (Honjo et al., 1984, pp. 63-64); in number of adult females varied from 27 to 46, the other captive groups, reported mean gestation annual birth rate reported in 1986 and 1990-1992 lengths vary from 160.0 to 168.0 days (extremes was 0.59, 0.32, 0.41, and 0.33 (Wheatley & Harya 141-225 days) (Spiegel, 1954, p. 245; T. Fujiwara Putra, 1994, p. 248). In four free-ranging provi- & Imamichi, 1966, p. 226; Valerio et al., 1969, p. sioned groups of M. fascicularis introduced in P. 295; MacDonald, 1971, p. 374; Dang, 1977, p. 2; Tinjil, south of Java, the annual birth rate is es-

1983, p. 41; Dukelow et al., 1979, p. 44; Vara- timated to be 0.56 (Kyes, 1 993, p. 8 1). In captivity, vudhi et al., 1989a, p. 222). Mean birth weight in M. fascicularis females are capable of producing the large captive group was 318.2±45.2gfor563 young every year (Ridley, 1895, p. 24; Chance et female infants and 347.5 ± 55.2 g for 600 male al., 1977a, p. 612; cf. Hadidian & Bernstein, 1979, infants. The ratio of male infants to female infants p. 440; Timmermans et al., 1981, p. 120; Dang, tends to be greater for high-ranking captive fe- 1983, p. 41; Honjo et al., 1984, p. 66). males than for low-ranking females (van Schaik et Effective fertility in females probably ceases at

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 61 Table 33. Macaque fossils or subfossils collected within geographic range of Macaca fascicularis. about age 20 years (Spiegel, 1954, p. 265; Angst years from a few individuals to about 30,000 & Thommen, 1977, p. 212; van Noordwijk & van (Sussman & Tattersall, 1986, pp. 30, 38; Lawler Schaik, 1987, p. 587), which suggests that an av- et al., 1995, p. 138). In Florida, a seminatural erage female in a natural population may produce menagerie group increased from 6 individuals (4 about eight to nine infants during her reproductive females, 2 males) in 1933 to about 150 individuals life (age ca. 4—20 yr). The maximum known life in 1966 (Dumond, 1967, p. 203), an annual rate span in captive M. fascicularis is 37 years 1 month of increase of about 1 0%. (M. L. Jones, 1982, p. 1 17; cf. Dumond, 1967, p. 203; Angst, 1975, p. 350; van Noordwijk & van Schaik, 1988, p. 30). In northern Sumatra, neonatal infant mortality (deaths before age 3 mo/births) from 1980-1981 Fossils and Subfossils to 1983-1984 was 0.21, 0.12, 0.25, and 0.06, with an overall average of 0.14 (van Schaik & van Known fossils indicate that M. fascicularis has Noordwijk, 1985b, p. 538); neonatal infant mor- inhabited the Sunda Shelf since at least later Early tality was high in years when the birth rate was Pleistocene, approximately 1 Ma (Table 33). All low (see above). The average infant death rate be- of the earliest macaque fossils in this area, dating fore age 1 year was 0.22 in northern Sumatra (van from 0.8 to 1.0 Ma, have been collected in Java, Noordwijk & van Schaik, 1 987, p. 585); this infant where, at this time, M. fascicularis apparently was death rate is comparable to that reported in a pro- sympatric with M. nemestrina. In Java, M. fas- visioned free-ranging group at Ubud, Bali (0.25; cicularis fossils of this age outnumber those of M. Wheatley & Harya Putra, 1994, p. 248), and in a nemestrina, but most of these fossils consist of large captive group of M. fascicularis (> 0. 1 5; Lu- isolated teeth or small jaw fragments, and some der, 1993, p. 142). Infanticide, committed by adult may be misidentified. males, probably is one of the important causes of The long period from 800 Ka to 35 Ka has infant deaths (cf. Thompson, 1967, p. 18; Wash- yielded no macaque fossils within the present geo- burn & Hamburg, 1968, p. 473; Angst & Thom- graphic range of M. fascicularis. However, fossils men, 1977, p. 208; Erwin, 1977, p. 359; Tim- and subfossils of M. fascicularis and M. nemes- mermans et al., 1981, p. 120; Wheatley, 1982, p. trina have been collected in subsequent Late Pleis- 211; Pallaud, 1984, p. 92; Koyama, 1985, p. 106; tocene and Holocene deposits in West Malaysia, van Noordwijk & van Schaik, 1 987, p. 585; Ruiter Sumatra, Borneo, and Java (M. fascicularis fossils etal., 1994, p. 218). only). In these deposits in Borneo and Java, M. In the Ketambe study area, northern Sumatra, fascicularis fossils and subfossils outnumber those a natural nonprovisioned population of M. fas- of M. nemestrina. In deposits in West Malaysia cicularis increased from 70-75 in 1972 to more and Sumatra, by contrast, M. nemestrina fossils than 190 in 1986 (van Schaik & van Noordwijk, and subfossils outnumber those of M. fascicularis. 1988, p. 94), an annual rate of increase of about This is anomalous, because, at present, M. fasci- 7%; at this rate, which obviously is not indefinitely cularis generally outnumbers M. nemestrina in the sustainable, the population would double every broad area of overlap of their respective ranges, 10.5 years. The introduced nonprovisioned pop- except in the interior of primary evergreen rain ulation in Angaur increased in less than 75 years forest (Fooden, 1975, p. 61; above, Natural His- from a few individuals to 480-600 in 1 973 (Poirier tory). The explanation for the anomalous relative

& Smith, 1974, pp. 264, 271; cf. Kawamoto et al., frequencies offossils and subfossils ofthese species

1988, p. 176; Matsubayashi et al., 1989, p. 54). In in Sumatra and West Malaysia is unclear. Mauritius, another introduced nonprovisioned In the Lesser Sunda Islands, Holocene subfossils population apparently increased in less than 500 of M. fascicularis, consisting of human food re-

7 Age estimates from Theunissen et al. (1990, p. 51; cf. Swisher et al., 1994, p. 1 1 19; J. de Vos & Sondaar, 1994, p. 1726; Swisher, 1994, p. 1727). 8 Cf. Koenigswald (1951, p. 219) and Medway (1972c, p. 80). 9 SeeMusser(1981,p. 72). 10 Age estimate indicates age of earliest fossil collected at each locality; number of fossils indicates minimum number of individuals collected at each locality.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 63 Fig. 25. Type localities and limits ofdistribution of recognized subspecies of Macacafascicularis and type localities in the indicated of synonymous nominal species or subspecies (cf. Fig. 1; Fooden, 1982, p. 576); Philippine Islands, the intersubspecific contact zone includes an area for which the subspecies is undetermined (cf. Fooden, 1991, p. 25). Because the geographic origins of the holotypes of Macacus cristatus Gray, 1870, and Macacus cynomolgus . . . var. Cumingii Gray, 1870, are inadequately known, these two synonyms of M. fascicularis are excluded from this figure (see Fooden, 1991, pp. 23, 24). Key to type localities of synonyms of recognized subspecies:

11. Pithecus karimoni P. Kar- M. f. aurea Elliot, 1909; Indonesia, imun: Monos. 1 . Pithecus vitiis Elliot, 1910; Burma, Letsok-aw Kyun. 12. Pithecus dollmani Elliot, 1909; Singapore, Singa- pore I.: Changi. M. f. fascicularis 13. Pithecus laetus Elliot. 1909: Malaysia, West Malay- 2. Pithecus validus Elliot, 1909; Vietnam: "Cochin sia, P. Tinggi. China." 14. Pithecus bintangensis Elliot, 1909; Indonesia, P. 3. Pithecus capitalis Elliot, 1910; Thailand, Trang Bintan: Sungei Biru. Province: Ban Phra Muang. 15. Pithecus impudens Elliot, 1910; Indonesia, P. Sugi. 4. Macaca irus argentimembris Kloss, 1911b; Malay- 16. Pithecus lingae Elliot, 1910; Indonesia, P. Lingga. sia, West Malaysia, P. Pinang [2]. 17. Pithecus lapsus Elliot, 1910; Indonesia, P. Bangka: 5. Pithecus agnatus Elliot, 1910; Indonesia, P. Tuang- Tanjung Pamuja. ku. 18. Pithecus lautensis Elliot, 1910; Indonesia, P. Laut. 6. Macacus phaeura Miller, 1 903b; Indonesia, P. Nias: 1 9. Pithecus lingungensis Elliot, 1910; Indonesia, P. La- Teluk Siaba. gong. 7. Pithecus mansalaris Lyon, 1916; Indonesia, P. Mur- 20. Pithecus sirhassensis Elliot, 1910; Indonesia, P. Ser- sala. asan.

8. Macacus irus I. Geoffroy, 1 826; Indonesia, Sumatra. 2 1 . Macacus pumilus Miller, 1 900; Indonesia, P. Benua. 9. Macacus carbonarius I. Geoffroy, 1826; Indonesia, 22. Pithecus mandibularis Elliot, 1910; Indonesia, Bor- Sumatra (technical name also mistakenly applied to neo, Kalimantan: Sungai Ambawang, near Pontia- M. f. aurea and M. f. umbrosa). nak. 10. Pithecus alacer Elliot, 1909; Indonesia, P. Kundur: 23. Pithecus carimatae Elliot, 1910; Indonesia, P. Kar- Selatbliat. imata: Teluk Pai.

64 FIELDIANA: ZOOLOGY mains, have been collected in caves in Timor and Systematics Flores, both of which are east of Wallace's Line (passing between Bali and Lombok). In four care- Subspecific Taxonomy fully excavated caves in Timor, where basal de- posits date back to ca. 13.5 Ka, the oldest M. The geographic range of M. fascicularis is broad fascicularis subfossils do not appear until ca. 4.5 and encompasses mainland Southeast Asia and Ka (Glover, 1986, p. 212). At lower levels, the numerous large and small islands, both on and only mammals included among the human food beyond the Sunda Shelf (Figs. 3, 4). It is therefore remains are murid rodents and bats, of which not surprising that this species exhibits great in- thousands of skeletal fragments were collected; dividual, local, and geographic variation in a di- these lower levels also yield stone tools. At ap- verse array of characters (see above); variation proximately the same level where M. fascicularis among populations of M. fascicularis ranges from first appears, remains of palm civet and cuscus trivial, to statistically significant, to locally dis- also appear, along with remains of domestic goat, continuous, to totally discontinuous. Based on this pig, and dog and pottery. Glover (1986, p. 159) variation, 50 specific or subspecific names have concluded that "... there is strong presumptive been proposed for populations of this species (Fig. evidence that these species [M. fascicularis, palm 25). The evaluation of intraspecific variation for civet, and cuscus] were introduced into Timor di- the purpose of defining taxonomically useful sub- rectly by, or through the agency of man, between species presents serious theoretical and practical about 4000-5000 years ago." For M. fascicularis, difficulties, as previously discussed by Chasen a less plausible alternative hypothesis is that it was (1940a, p. 66; cf. Table 35) in his classic review present in Timor earlier but was not hunted by of M. fascicularis. the preceramic cave inhabitants, possibly because In the present taxonomic analysis, subspecific they lacked suitable projectile weapons (cf. Adi, status is accorded only to those geographic pop- 1985, p. 65). ulations in which variation ofat least one character In Flores, a subfossil molar of M. fascicularis is completely— or nearly completely— discontin- was collected in cave deposits, dated ca. 3.5 Ka, uous with variation of the same character in the that also contain human artifacts and human skel- nominotypical subspecies, M. f fascicularis. The etal remains. Other mammals represented in these reason for selecting character-state discontinuity deposits are rat species, fruit bat, porcupine, and as the critical requisite for subspecific recognition pig (Hooijer, 1967, p. 160). Based on available is that, without such discontinuity, unambiguous zoogeographic evidence, including Glover's ar- diagnosis of subspecies becomes difficult or im- cheological data from Timor (see above), Musser possible. The requirement of character-state dis-

1 1 concluded that M. ( 98 , p. 133) fascicularis, por- continuity denies subspecific recognition to local cupine, and pig probably were introduced into Flo- populations that are distinct from adjacent pop- res by humans. ulations of M. f fascicularis but that fall within

24. Macaca resima Thomas & Wroughton, 1909b; In- 33. Cynomolgus suluensis Meanrs, 1905; Philippines, donesia, Java: Tasikmalaya. Jolo I.: foot of Crater Lake Mountain. 25. Macaca modax Thomas & Wroughton, 1909c; In- Java: donesia, Cilacap. M. f. fascicularis/M. f. philippinensis contact zone 26. [Simia] Aygula Linnaeus, 1758; Indonesia, Java. 34. mindanensis 27. Pithecus baweanus Elliot, 1910; Indonesia, P. Baw- Cynomolgus apoensis Mearns, 1905; ean. Philippines, Mindanao I.: Mt. Apo. 35. mindanensis 28. Pithecus cupidus Elliot, 1910; Indonesia, P. Mata- Cynomolgus Mearns, 1905; Philip- I.: siri. pines, Mindanao Pantar. 29. Macaca irus submordax Sody, 1949; Indonesia, P. Bali: Desa Poetjang. M. f. philippinensis 30. Macaca irus sublimitus Sody, 1932; Indonesia, P. Sumba: Payeti-Kambaniru and Mao Marroe. 36. Pithecus mindorus Hollister, 1913; Philippines, 31. Pithecus fascicularis limitis Schwartz, 1913; Indo- Mindoro I.: Alag River. nesia, P. Timor: Lelogama. 37. Macacus palpebrosus I. Geoffroy, 1851; Philippines, 32. Cynomolgus cagayanus Mearns, 1905; Philippines, Luzon I.; "forets de Manille." Cagayan Sulu I. 38. Macacus fur Slack, 1867; Philippines, Luzon I.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 65 Table 34. Summary comparison of external characters in recognized subspecies of Macaca fascicularis. x available evidence, variation of characters in Su- pendixes 3-9); some ofthese populations also have matra is continuous with that in the Malay Pen- been recognized as separate subspecies by other insula (including the Isthmus of Kra north to ca. authors. 1 0°N), Borneo, and Java (see above); all of these The deep-water fringing-island population ofM. parts of the core area are therefore included within fascicularis in P. Nias (west of Sumatra) resembles

the geographic range of M. f. fascicularis, as here the eight shallow-water fringing-island popula- defined. In the southern part of the Indochinese tions listed above in being distinct— in tail col- Peninsula, character-state variation in M. fasci- oration and T— from nearby core-area popula- cularis also is continuous with that in the Malay tions of M. f fascicularis in Sumatra but falling Peninsula, Sumatra, Borneo, and Java, with the within the range of variation of more distant pop- possible exception of variation of mtDNA types ulations of this subspecies. Although the proxi- (Table 17) and resistance to experimental infec- modorsal surface of the tail is more intensely tions with Plasmodium knowlesi malaria (see blackish in P. Nias than in Sumatra, as empha- above; Fooden, 1994, p. 585). Because evidence sized by Miller (1903b, p. 63) in his description for discontinuity ofvariation ofmtDNA types and of the P. Nias monkey as Macacus phaeura, tail P. knowlesi resistance is incomplete, the southern coloration in M. fascicularis in P. Nias broadly part of the Indochinese Peninsula is provisionally overlaps that in specimens collected in other parts

retained within the geographic range of M. f. fas- ofthe species range (cf. mcz 36030, usnm 121871 — cicularis. Although now disjunct, the range of M. P. Nias; usnm 196823— Kalimantan; mcz 37415 — f fascicularis in the southern part of the Indo- Sabah; usnm 124710-P. Bangka; usnm 124969, Chinese Peninsula was connected by dry land to 124970-P. Belitung). T and relative T in P. Nias the range ofthis subspecies in the Malay Peninsula similarly overlap T and relative T in other parts during the last glacial maximum, ca. 1 8 Ka (Fig. of the species range (Figs. 15, 17; Appendixes 8, 3). 9). Lacking character-state discontinuity, M. fas- In most shallow-water fringing-island popula- cicularis in P. Nias is regarded here as subspecif- tions of M. fascicularis, variation is continuous ically inseparable from M. f fascicularis. Deep- with that in core-area M. f fascicularis (Appen- water fringing-island populations of M. fascicu- dixes 3-9; Fooden & Albrecht, 1993, p. 537); laris in the Lesser Sundas and southern Philippines therefore, these islands are included within the are retained within M.f. fascicularis because known geographic range of M. f fascicularis. In eight of variation in these populations is continuous with these shallow-water fringing-islands, populations that in nearby core-area populations of M. f fas- are distinct from nearby core-area populations of cicularis (Appendixes 3-5, 7-9; Fooden, 1991, p. M.f fascicularis but fall within the range of vari- 21; Fooden & Albrecht, 1993, p. 537). ation of more distant populations of this subspe- 2. M. f aurea. In the western part of the In- cies; as indicated above, these populations are not dochinese Peninsula and western part of the Isth- recognized here as separate subspecies, although mus of Kra south to about 1 0°N, variation of the some have been recognized as subspecies by other lateral facial crest pattern (infrazygomatic) is al- authors. These eight populations, sorted according most completely discontinuous with variation of to characters that distinguish the fringing-island this character (transzygomatic) in the rest of the populations from nearby core-area populations of core area (Figs. 8, 9). This warrants recognition of M.f. fascicularis, inhabit the following islands: P. the subspecies M. f aurea (type locality, Pegu, Mursala (west of Sumatra), P. Redang (east of the southern Burma). In all samples available from Malay Peninsula), P. Lagong (west of Borneo), and shallow-water islands in the , P. Uwi (west of Borneo)— distinguished by dorsal west of the northern part of the Isthmus of Kra, pelage erythrism (Appendix 4); P. Tioman (east of the lateral facial crest pattern also is infrazygo- the Malay Peninsula) and P. Belitung (west of Bor- matic; these islands are therefore included within neo)— distinguished by T and/or relative T (Ap- the geographic range of M. f aurea. pendixes 8, 9); and Ko Kut and Phu Quoc Dao Two areas immediately east of the range of M. (both south of the Indochinese Peninsula)— dis- f aurea are heterogeneous for the lateral facial tinguished by skull length (Fooden & Albrecht, crest pattern (Fig. 9). The more southern of these 1993, p. 528). In many other fringing-island pop- two areas, in eastern and southern parts of the ulations, character-state variation is statistically Isthmus of Kra and adjacent southwestern islands, significantly different from, but continuous with, may be regarded as an intersubspecific contact zone variation in nearby core-area populations (Ap- between the ranges of M. f fascicularis and M. f

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 67 Table 35. Subspecies of Macaca fascicularis recognized as valid by Chasen (1940a) and recent authors; dash is in the cited work. (— ) indicates that subspecies extralimital matra, is one of four deep-water fringing-island Brief casual remarks have been published con- populations with distinctively blackish dorsal pel- cerning dorsal pelage coloration in M. fascicularis age (Appendix 3). The crown is yellowish brown in P. We, a deep-water island off the northern tip

and the thighs are pale brownish gray in M. f. of Sumatra (Scheffrahn et al., 1994, p. 136). These umbrosa. remarks are inadequate for determination of the 7. M. fascicularisfusca. In the population ofM. subspecific status of this population. fascicularis in deep-water P. Simeulue, west of Su- The present subspecific classification of M. fas- matra, the back, crown, and thighs are blackish, cicularis differs from recent classifications pro- and the tail is shorter than usual in M. fascicularis posed by W. C. O. Hill (1 974, p. 504), R. H. Napier

(Figs. 16, 18; Appendixes 3, 8, 9). (1981, p. 12), Scott (1982, unpubl. M.A. thesis, p. 8. M.f lasiae. In the population in deep-water 182), Weitzel et al. (1988, p. 96), and Fa (1989, P. Lasia, near P. Simeulue, the back, crown, and p. 54) (Table 35). P. H. Napier, whose list of sub- thighs are blackish, as in M. fascicularisfusca, but species follows that of W. C. O. Hill with a single the tail is of normal length (Figs. 16, 18, 27; Ap- exception (nonrecognition of subspecies cagayana pendixes 3, 8, 9). Mearns, 1905), explicitly acknowledges that her 9. M.f. tua. In the population in deep-water P. classification is unsatisfactory and that many of Maratua, east of Borneo, the back is blackish, the the recognized subspecies cannot be diagnosed. crown is yellowish brown, and the thighs are Scott's classification is based on a sample of 148 brownish gray (Appendix 3). In M. f tua, unlike specimens (including 1 34 skins and 69 skulls), more M. f. umbrosa, subauricular hairs are elongated than 90% ofwhich are in two museums (mzb, zrc); and pale ochraceous-buff. this sample, which apparently is almost identical 10. M. f philippinensis. In populations of M. with that studied by Chasen (1940a, p. xix), pro- fascicularis in western, northern, and eastern is- vides an inadequate representation of variation in lands of the Philippine Archipelago, dorsal pelage M. fascicularis. Weitzel et al. indicate that their color is distinctively dark brown (Appendix 3; classification is largely based on Scott's research.

Fooden, 1991, p. 3). In eastern and central Min- Fa, without further explanation or documentation, danao, southern Negros, and perhaps in some lists 2 1 subspecies of M. fascicularis that he rec- nearby islands, mixed populations ofdark and pale ognizes as valid; most, but not all, of these sub- individuals occur; this area of mixed populations species are the same as those recognized by authors is regarded as a contact zone between M. f phi- cited above. lippinensis and M. f fascicularis.

Key to Recognized Subspecies

The 10 subspecies of Macaca fascicularis recognized here are distinguished in the following key, which is based on external characters of postinfants:

1 . General color of dorsal surface of trunk buffy to yellowish gray to medium brown to dark brown, variably erythristic 2 General color of dorsal surface of trunk blackish 7

2. Preauricular hairs directed posteriorly, partly covering ears (lateral facial crest infrazygomatic) .... aurea

Preauricular hairs directed anteriorly, forming part of lateral facial crest, not covering ears (lateral facial crest transzygomatic) 3 3. General color of dorsal surface of trunk buffy to yellowish gray to medium brown, variably erythristic 4

1 General color of dorsal surface of trunk dark brown, variably erythristic 6 4. Crown conspicuously darker than back 5 Crown not conspicuously darker than back fascicularis

1 Note: Approximately 5% of M.f. fascicularis specimens examined are dark brown and, if of unknown geographic origin, probably would be misidentified as either M. f. karimondjawae or M. f. philippinensis.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 69 5. Dark crown patch narrow, extending laterally approximately as far as middle of each eye, sharply delimited laterally and posteriorly atriceps Dark crown patch broad, extending laterally to or beyond lateral margin of each eye, not sharply delimited laterally and posteriorly condorensis 6. Pale annulations of dorsal hairs pale yellowish karimondjawae Pale annulations of dorsal hairs golden to rufescent philippinensis

1 . Outer surface of thighs blackish, approximately same color as dorsal surface of trunk 8 Outer surface of thighs brownish gray, contrastingly paler than dorsal surface of trunk 9 8. Outer surface of shanks blackish; tail length < 1 12% of head and body length fusca

1 1 Outer surface of shanks pale grayish brown; tail length > 2% of head and body length . . . lasiae 9. Subauricular hairs pale ochraceous-buff, elongated (4-6 cm), extending to apex of lateral facial crest

1 . . forming a conspicuous pale lateral facial patch; tail length > 22% of head and body length . tua Subauricular hairs brownish gray to pale gray, not elongated (2-4 cm), not extending to apex of lateral facial crest, relatively inconspicuous; tail length < 122% of head and body length umbrosa

Subspecies Accounts Simia Cynamolgos: Schreber, [1774], p. 91, pi. 13 (part; not Linnaeus, 1758, p. 28)— misidentification of Ma- caque: Buffon in Buffon & Daubenton, 1 766. Schlegel, Macaca fascicularis fascicularis 1876, p. 101 —name said to have been based on mis- p. (Raffles, [1821], 246) identification. Blanford, [1888a], p. 624— name said to have been based on misidentification.

Cercopithecus (Cynamolgus): Zimmermann, 1780, p. 186 Jawanska Markattor: Osbeck, 1757, p. 99— external (part; not Linnaeus, 1758, p. 28)— misidentification characters and behavior of monkeys collected and ob- of Macaque: Buffon in Buffon & Daubenton, 1766. served in Java by P. Osbeck, Jul. 1751; specimens not Cercocebus cynamolgos: Schlegel, 1876, p. 101 (not Lin- preserved. naeus, 1758)— taxonomic — comparisons. [Simia] Aygula Linnaeus, 1758, p. 27 based on Ja- [Cercopithecus] Cynomolgus: Erxleben, 1 777, p. 28 (part; wanska Markattor: Osbeck, 1757, and supplementary not Linnaeus, 1766, p. 38)— misidentification of Ma- information in unpublished letters from Osbeck to caque: Buffon in Buffon & Daubenton, 1766. Linnaeus H. & Groves, 1 983, p. 1 1 7). Tho- Simia 5 (P. Napier — Cynomolgus: Audebert, 1798-1799, p. (part; mas & Wroughton, 1909a, p. 373 misidentified as a not Linnaeus, 1766, p. 38)— misidentification of Ma- species of leaf monkey. P. H. Napier & Groves, 1983, caque: Buffon in Buffon & Daubenton, 1766. p. 118— shown to be a senior synonym of Macaca Simia cynomolgos: G. Cuvier, 1798, p. 98— incorrect fascicularis (Raffles, [1821]); suppression requested. spelling of [Simia] Cynomolgus: Linnaeus, 1766; mis- International Commission on Zoological Nomencla- identification of Macaque: Buffon in Buffon & Dau- ture, 1986, p. 229— specific name officially suppressed. benton, 1766. [Cercopithecus] aygula: Erxleben, 1777, p. 39— external Macaca cynomolgus: Daudin in Lacepede & Daudin, and cranial characters. [1802], p. 148 (not Linnaeus, 1766)— misidentifica- Cercop[ithecus] Aigula: Schinz, 1821, p. 108— incorrect tion of Macaque: Buffon in Buffon & Daubenton, 1766. spelling of [Simia] Aygula Linnaeus, 1758. Schinz, [M]acacus cynomolgus: Desmarest, 1820, p. 65 (part; 1825, p. 256 — said to be indistinguishable from Inuus not Linnaeus, 1766)— misidentification of Macaque: cynamolgus [— Macaca fascicularis]. Buffon in Buffon & Daubenton, 1766. Blanford, 1888b, Macaca aygula: Daudin in Lacepede & Daudin, [1802], pp. 21, 23— binomial said to be based on misidenti- p. 148— listed as "Variete A" of Macaca cynomolgus fication, accepted as valid. ' [= M. fascicularis]. Macacus cynomolgus' Auct.: Miller, 1900, p. 239 — Pithecus aygula: E. Geoffroy, 1 803, p. 24— external char- specimens collected in P. Lingga and P. Tioman. Mil- acters. ler, 1903a, p. 476 — specimens collected at Teluk Ta- Cercocebus aygula: E. Geoffroy, 1812, p. 99— external panuli, Sumatra, and in P. Tuangku and P. Mursala. characters. Cyn[ocephalus] cynomolgus: Latreille, 1 804, p. 292 (part; Macaque: Buffon in Buffon & Daubenton, 1766, p. 190, not Linnaeus, 1766, p. 38)— misidentification of Ma- pis. 20, 22-24— based on specimen of unknown origin, caque: Buffon in Buffon & Daubenton, 1766. not preserved; species said to inhabit "Congo & des Cercocebus cynomolgus: E. Geoffroy, 1812, p. 99 (not autres de meridionale." F. external and cranial characters. parties l'Afrique— Cuvier, Linnaeus, 1766)— 1819, liv. 3, p. 1, 2 pis. illustrations of captives des- [P]ithecus cynomolgus: Desmarest, 1817, p. 323 (part; ignated as Irus by F. Cuvier, 1818; habitat, "vra- not Linnaeus, 1766, p. 38)— misidentification of Ma- isemblablement du Senegal ou de la cote de Guinee." caque: Buffon in Buffon & Daubenton, 1766. — Aigrette: Buffon in Buffon & Daubenton, 1766, p. 190, Pithecus Cynomolgos: Blainville, [1839], pi. 7 incorrect pi. 21— based on specimen of unknown origin, not spelling of [Simia] Cynomolgus: Linnaeus, 1 766. preserved; regarded as "variete" of Macaque: Buffon Pith[ecus] (Mac[acus]) cynomolgus: Dahlbom, 1856, p. in Buffon & Daubenton, 1766. 1 18 (not Linnaeus, 1766, p. 38)— external characters.

70 FIELDIANA: ZOOLOGY I[nuus] cynomolgus: Wagner, [1839], p. 135 (not Lin- Macaca irus: H. C. Robinson & Kloss, 1915a, p. 130 — naeus, 1766, p. 38)— external characters; distribution; specimens collected in Ko Samui and Ko Phangan. taxonomic comparisons. [Macaca] zVus-group: Miller, 1933, pp. 5, 6— taxonomic Irus: F. Cuvier, 1818, p. 120— unavailable name, not history. published in combination with generic name; pro- [Macaca] ira: Weinman & Wiratmadja, 1969, p. 498 — posed as substitute specific technical name for Ma- trypanosomes. caque: Buffon in Buffon & Daubenton, 1766 (cf. I. Pithecus irus: Elliot, 1913, p. 229— taxonomic compar- Geoffrey, 1826, p. 588; [1831], p. 56: Gervais, 1854, ison. p. 85; Miller, 1942, p. 127); habitat "vraisemblable- [Silenus] irus: Stiles & Nolan, 1929, p. 530— parasites. ment du Senegal ou de la cote de Guinee" (corrected Cynomolgus irus: Furuya, 1962, p. 377— dermatoglyph- to "du midi de l'Asie, et particulierement de Sumatra" ics. by F. Cuvier, 1825, liv. 52, p. 1). Macaca irus irus: H. C. Robinson & Kloss, 1918, p. 6— Simia fasciculcris Raffles, [1821], p. 246— for details collected in Sumatra; taxonomic history. Chasen, concerning holotype, see below. Cantor, 1846, p. 176 — 1940a, p. 66— taxonomic comparisons. Kellogg, 1945, cited as a synonym of Cercopithecus cynomolgus. H. pp. 116, 129— taxonomic comparisons. Sody, 1949, C. Robinson, 1916, p. 63— type locality restriction. p. 130— taxonomic comparison. Semn[nopithecus]l fascicularis: [Vigors], 1830, p. 642— Carbonarius: F. Cuvier, 1825, liv. 52, p. 2, pi.— un- geographic distribution; type history; "doubtful whether available name, not published in combination with it is a true Semnopithecus." generic name; based on living captive, origin un- Semnopithecus fascicularis?: Waterhouse, 1838, p. 4— known; specimen not preserved. geographic distribution. Macacus carbonarius I. Geoffroy, 1826, p. 588— based S[emnopithecus] fascicularis: Martin, 1838, p. 435— ex- on Carbonarius: F. Cuvier, 1825; external characters; ternal characters. taxonomic comparison. Lesson, 1827, p. 42— said to Macacus fascicularis: Bonhote, 1903, p. 3— taxonomic inhabit Sumatra. I. Geoffroy, [1831], p. 63— doubt- history. fully distinct. Eydoux & Souleyet [& Gervais], 1841, Macaca fascicularis: Miller, 1906b, p. 65— specimens p. 6— cited as a synonym of Macacus aureus I. Geof- collected in P. Karimata. Wroughton, 1918, p. 556— froy, [1831] [= Macacafascicularis aurea]. Blyth, 1875, taxonomic history. Miller, 1942, p. 126— taxonomic p. 7— cited as a synonym of Macacus cynomolgus [= history. Hooijer, 1962b, p. 44— subspecific taxonomy. Macaca fascicularis]. Macacafascicularis group: Thomas, 1928, p. 832— spec- S[imia] carbonaria: Fischer, 1829, p. 26— habitat, Su- imens collected in Vietnam. matra. carbonarius: 136 — [Cynomolgus] fascicularis: Trouessart, 1904, p. 16— geo- C[ynamolgus] Reichenbach, 1862, p. graphic distribution. taxonomic comparison. kra 20— Pithecus fascicularis: Elliot, 1913, p. 233— taxonomic Semnopithecus Lesson, [1830], p. replacement for Simia 246. comparison. name fascicularis Raffles, [1821], p. Semnopithecus Buku Martin, 1 838, p. 435 —specific name [Silenus] fascicularis: Stiles & Nolan, 1929, p. 529— proposed provisionally for Kra Buku: Raffles, [1821], parasites. p. 247. J. Anderson, 1879, p. 74— cited as a synonym P[ithecus] f[ascicularis] fascicularis: Schwarz, 1913, p. of Macacus cynomolgus [= Macaca fascicularis]. 297— taxonomic comparison. Inuus cercopithecus Mason, 1851, p. 220— unavailable M[acaca] fascicularisfascicularis J. R. Napier & Napier, name attributed to Blyth, cited in synonymy, never 403— distribution. 1967, p. geographic treated as available. mulatto Macaca fascicularis: Fooden, 1964, p. 364— 1 1 1 Macacus cancrivorus Mason, 85 , p. 22 —unavailable taxonomic comparison. name attributed to Blyth, cited in synonymy, never Kra Buku: Raffles, [1821], p. 247— vernacular name of treated as available. "smaller species" than Simia fascicularis Raffles, C[ynamolgus] cynocephalus: Reichenbach, 1862, p. 133 [1821], p. 246 [apparently juvenile Macaca fascicu- (not Linnaeus, 1766, p. 38)— natural history. laris], specimens apparently not obtained; habitat, Su- Macacus assamensis: Gray, 1 870, p. 3 1 (not McClelland islands." matra and "other Malay in Horsfield, [1840], p. 148)— misidentification; tax- Macacus irus I. Geoffroy, 1826, p. 588— based on Ma- onomic comparison. caque: Buffon in Buffon & Daubenton, 1 766, and liv- Macacus Sinicus: Morice, 1875, p. 41 (not Linnaeus,

F. 1 77 1 ing captives reported by Cuvier (1818, p. 112; 1819, , p. 52 1)— misidentification of monkeys observed liv. 3, p. 1); specimens not preserved, origin unknown; in Vietnam. binomial incorrectly attributed to F. Cuvier, 1818. I. Macacus pumilus Miller, 1900, p. 241 —holotype, usnm Geoffroy, [1831], p. 56— cited as a synonym of Ma- 101639, adult male, skin and skull, collected at Pulo cacus cynomolgus Desmarest, 1820; taxonomic his- Bunoa (= Pulau Benua), Kepulauan Tambelan, In- tory. Blanford, [1888a], p. 624— taxonomic history. donesia, by W. L. Abbott, 6 Aug. 1899; paratypes, Cabrera, 1910, p. 620— taxonomic history; type lo- usnm 101638 (juvenile female, P. Benua, 5 Aug.), usnm cality, Sumatra. H. C. Robinson & Kloss, 1914, p. 101666 (adult female, P. Uwi, 13 Aug.), and usnm 394— type locality, probably (= Melaka). 101711 (subadult male, P. Siantan, 8 Sep.), skins and Miller, 1942, p. 127— cited as a synonym of Macaca skulls, collected in Indonesia by W. L. Abbott, 1899. fascicularis (Raffles, [1821]); binomial attributed to Lyon & Osgood, 1909, p. 284— holotype cataloged. Blanford, [1888a]. Fooden, 1976, p. 226-cited as a Weitzel et al., 1988, p. 112— topoparatypic specimens synonym of Macaca fascicularis (Raffles, [1821]); bi- (P. Siantan) referred to M. fascicularis sirhassensis (El- nomial attributed to I. Geoffroy, 1826. liot, 1910).

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 71 [Cynomolgus] pumilus: Trouessart, 1904, p. 16— distri- lotype cataloged. Poole & Schantz, 1942, p. 241 — bution. holotype cataloged. Pithecus pumilus: Elliot, 1 9 1 3, p. 252— external and cra- [Macacus] cagayanus: Raven, 1935, p. 236— geographic distribution. nial characters. — Macaca irus pumilus: Chasen & Kloss, 1928b, p. 29 Pithecus cagayanus: Hollister, 1912, p. 36— geographic taxonomic comparison. distribution. Macaca irus pumila: Chasen, 1940a, p. 69— geographic Macaca irus cagayana: W. C. O. Hill, 1974, p. 525 — distribution. Poole & Schantz, 1942, p. 245 — holotype external and cranial characters. cataloged. Weitzel et al., 1988, pp. Ill, 1 13— cited as M[acaca] p[hilippinensis] cagayanus: Rabor, 1986, p. name previously applied to specimens referred to M. 138— geographic distribution. fascicularis lingungensis (Elliot, 1910) and M. f. sir- Cynomolgus fuscus: Moszkowski, 1909, pp. 143, 302 hassensis (Elliot, 1910). (not Miller, 1903a, p. 476)— misidentification of spec- M[acaca] f[ascicularis] pumila: J. R. Napier & Napier, imens collected in Sumatra. 403— distribution. Pithecus validus Elliot, 1909, p. 252 — 1967, pp. 349, geographic — holotype, bm(nh) Macacus phaeura Miller, 1903a, p. 63 holotype, usnm 1881.6.30.2, adult male, skin and skull, collected in 121870 (Coll. No. 2399), adult male, skin and skull, "Cochin China," Vietnam, by M. Boucard, before 1 882. collected at Teluk Siaba, P. Nias, Indonesia, by W. L. Macaca validus: Kloss, 1919c, p. 348— taxonomic com- Abbott, 20 Mar. 1903; paratypes, usnm 121868 (Coll. parison. No. 2393, adult male, 16 Mar.), usnm 121869 (Coll. M[acaca] i[rus] validus: Kloss, 1921, p. 75— taxonomic No. 2395, adult female, 18 Mar.), and usnm 121871 comparison. (Coll. No. 2400, adult male, 20 Mar.), skins and skulls, Macaca irus valida: Kellogg, 1945, p. 1 19— geographic collected at Teluk Siaba, P. Nias, Indonesia, by W. L. distribution. Abbott, 1903. Lyon & Osgood, 1909, p. 284-holo- M[acaca] f[ascicularis] valida: J. R. Napier & Napier, type cataloged. 1967, pp. 349, 403— geographic distribution. P. H. Macaca phaeura: Lyon, 1906, p. 606— specimens col- Napier, 1981, pp. 13, 19— external characters; holo- lected in P. Bangka and P. Belitung. Miller, 1942, p. type cataloged. — 128 specimens collected in P. Nias. Macaca fascicularis validus: Van Peenen et al., 1971, p. Pithecus phaeurus: Elliot, 1913, p. 243— external and 134 (part)— female specimens collected at Sontra Peak; cranial characters. geographic variation. Pithecus phaeura: Lyon, 1916, p. 458— geographic dis- Pithecus alacer Elliot, 1909, p. 253— holotype, bm(nh) tribution. 1909.4.1.36 (Coll. No. 1454), adult male, skin and Macaca irus phaeura: Chasen, 1940a, p. 69— geographic skull, collected at Bliah (= Selatbliat), P. Kundur, Ke- distribution. Poole & Schantz, 1942, p. 245— holotype pulauan Riau, Indonesia, by H. C. Robinson and E. cataloged. Seimund, 18 Aug. 1908; paratype (available but not M[acaca] /[ascicularis] phaeura: J. R. Napier & Napier, explicitly cited in original description), bm(nh) 1967, pp. 349, 403— geographic distribution. 1909.4.1.37 (Coll. No. 1495, adult female, skin and collected at P. Cynomolgus suluensis Mearns, 1905, p. 430— holotype, skull), Selatbliat, Kundur, Indonesia, C. E. usnm 125324 (Coll. No. 5750), adult male, skull only, by H. Robinson and Seimund, 20 Aug. 1908. H. C. Robinson his- collected at foot of Crater Lake Mountain, Jolo I., & Kloss, 1914, pp. 393, 394-type Philippine Islands, by E. A. Mearns, 16 Nov. 1903. tory. Chasen, 1925, p. 93— cited as a synonym of Ma- caca irus F. Cuvier 1 W. C. O. Hill, 1974, p. 522-cited as a synonym of (I. Geoffroy, 826). Chasen, 1940a,

I 1 p. 66— cited as a of Macaca irus irus F. Cu- Macaca irus philippinensis Geoffroy, [ 843]. Fooden, synonym vier P. H. 1991, p. 22— cited as a synonym of Macaca fascicu- (I. Geoffroy, 1826). Napier, 1981, pp. 13, cited as a laris fascicularis (Raffles, [1821]). 15— synonym of Macaca fascicularis fasci- cularis Macaca suluensis: Lyon & Osgood, 1909, p. 284— ho- (Raffles, [1821]); holotype cataloged. alacer: 236— dis- lotype cataloged. [Macacus] Raven, 1935, p. geographic tribution. Macaca suluana: Alcasid, [1970], p. 24— incorrect spell- M[acaca] i[rus] alacer: Dammerman, 1926b, p. 316 — ing of Cynomolgus suluensis Mearns, 1905. geographic distribution. Macaca sulvensis: Chiarelli, 1972, p. 2 1 3 — incorrect Pithecus karimoni Elliot, 1909, p. 254— holotype, bm(nh) spelling of Cynomolgus suluensis Mearns, 1905. 1909.4.1.34 (Coll. No. 1662), adult male, skin and [Macacus] suluensis: Raven, 1935, p. 237— geographic skull, collected at Monos, P. Karimun, Kepulauan Riau, distribution. Indonesia, by H. C. Robinson and E. Seimund, 30 Pithecus suluensis: Hollister, 1912, 37 — listed. Elliot, p. Aug. 1908; paratype, bm(nh) 1909.4.1.35 (Coll. No. 1913, p. 252— specific status indeterminate. 1636, adult female, skin and skull), collected at Mon- suluensis: 1 38 — M[acaca] p[hilippinensis] Rabor, 1986, p. os, P. Karimun, Indonesia, by H. C. Robinson and E. geographic distribution. Seimund, 29 Aug. 1908. H. C. Robinson Kloss, 1914, Cynomolgus cagayanus Mearns, 1905, p. 431— holo- p. 393, 394— type history; taxonomic comparison. type, usnm 125325 (Coll. No. 5771), adult male, skin Chasen, 1925, p. 93— cited as a synonym of Macaca and skull, collected in Cagayan Sulu I., Philippine Is- irus F. Cuvier (I. Geoffroy, 1826). Chasen, 1940a, p. lands, by E. A. Mearns, 25 Feb. 1904. P. H. Napier, 66— cited as a synonym of Macaca irus irus F. Cuvier 1981, p. 13— cited as a synonym of Macaca fascicu- (I. Geoffroy, 1826). P. H. Napier, 1981, pp. 13, 15- laris philippinensis I. Geoffroy, [1843]. Fooden, 1991, cited as a synonym of Macaca fascicularis fascicularis p. 22— cited as a synonym of Macaca fascicularis fas- (Raffles, [1821]); holotype cataloged. cicularis (Raffles, [1821]). [Macacus] karimoni: Raven, 1935, p. 236— geographic Macaca cagayana: Lyon & Osgood, 1909, p. 283 — ho- distribution.

72 FIELDIANA: ZOOLOGY M[acaca] i[rus] karimoni: Dammerman, 1926b, p. 316— M[acaca] i[rus] bintangensis: Dammerman, 1926b, p. geographic distribution. 3 1 6 —geographic distribution. Pithecus laetus Elliot, 1909, p. 255— holotype, bm(nh) M[acaca] f[ascicularis] bintangensis: Groves & Weitzel 1909.4.1.21 (Coll. No. 849), subadult male, skin and in Weitzel et al., 1988, pp. 5, 96, 99-external char- skull, collected in P. Tinggi, West Malaysia, by H. C. acters; geographic distribution. Robinson, 25 Jun. 1908; paratype (available but not Macaca mordaxThomas & Wroughton, 1909c, p. 380— explicitly cited in original description), bm(nh) holotype, bm(nh) 1909.1.5.27 (Coll. No. 613), sub- 1909.4.1.27 (Coll. No. 844, juvenile female, skin and adult male, skin and skull, collected at Tjilatjap (= skull), collected in P. Tinggi, by H. C. Robinson, date Cilacap), sea level, W Java, Indonesia, by G. C. Shor- unspecified, presumably Jun. 1908. Elliot, 1913, p. tridge, 19 Oct. 1907; paratype, bm(nh) 1909.1.5.28 236— geographic distribution, P. Tinggi and P. Tio- (Coll. No. 783, juvenile female, skin and skull), col- man, West Malaysia. H. C. Robinson & Kloss, 1914, lected at Cilacap, sea level, W Java, Indonesia, by G. p. 393— type history; taxonomic comparisons. C. Shortridge, 12 Nov. 1907 (cf. Thomas & Wrough- Macaca laetus: Kloss, 191 la— taxonomic comparison. ton, 1909a, p. 373). Miller, 1933, p. 8 -possible phys- [Macacus] laetus: Raven, 1935, p. 236— geographic dis- iological differentiation. tribution. [Macacus] mordax: Raven, 1935, p. 236— geographic Macaca irus laetus: H. C. Robinson, 1919, p. 325 — distribution. zoogeography. Pithecus mordax: Elliot, 1913, p. 232— geographic dis- Macaca irus laeta: Chasen, 1940a, p. 69— geographic tribution; external and cranial characters. distribution. P[ithecus]f[ascicularis] mordax: Schwarz, 1913, p. 296— Macacafascicularis laeta: Medway, 1966, p. 16— natural taxonomic comparison. history. P. H. Napier, 1981, pp. 13, 18— external char- Macaca fascicularis mordax: Hooijer, 1962b, p. 46— acters; holotype cataloged. cranial characters; taxonomic comparisons. P. H. Na- M[acaca]f[ascicularis] laeti.J. R. Napier & Napier, 1967, pier, 1981, pp. 13, 18— external characters; holotype pp. 349, 403— geographic distribution. cataloged. Pithecus dollmani Elliot, 1909, p. 256— holotype, bm(nh) Macaca irus mordax: Dammerman, 1928, p. 300— spec- 1909.4.1.20 (Coll. No. 1065), late subadult male, skin imens collected in Sumba; taxonomic comparisons. and skull, collected at Changi, Singapore I., by H. C. Sody, 1929, p. 165— included in fauna of Java. Sody, Robinson and E. Seimund, 22 Jul. 1908 (cf. P. H. 1933, p. 93— specimens collected in Bali; taxonomic Napier, 1981, p. 15). H. C Robinson & Kloss, 1914, comparisons. Mertens, 1936, p. 315 — specimens col- p. 393— type history. Kloss, 1919c, p. 347— taxonomic lected in Bali, Lombok, Sumbawa, and Flores; taxo- comparison. Chasen, 1924b, p. 59— taxonomic com- nomic comparisons. Sody, 1949, p. 131— specimens parison. Chasen, 1940a, p. 66— cited as a synonym of collected in Java; taxonomic comparisons. — Macaca irus irus F. Cuvier (I. Geoffroy, 1826). P. H. Macaca resima Thomas & Wroughton, 1909c, p. 381 Napier, 1981, pp. 13, 15— cited as a synonym of Ma- holotype, bm(nh) 1909.1.5.31 (Coll. No. 1219), late

1 skin cacafascicularisfascicularis (Raffles, [ 82 1]); holotype subadult male, and skull, collected at Tasikmalaja cataloged. (= Tasikmalaya), 1 145 ft (= 350 m), Preanger (region), [Macacus] dollmani: Raven, 1935, p. 236— geographic W Java, Indonesia, by G. C. Shortridge, 18 Jan. 1908. distribution. Dammerman in Chasen, 1940a, pp. 68, 70— cited as [Macaca irus] dollmani: Sody, 1949, table 1— external a synonym of Macaca irus mordax Thomas & and cranial characters. Wroughton, 1909c. P. H. Napier, 1981, pp. 13, 18- Pithecus bintangensis Elliot, 1909, p. 257— holotype, cited as a synonym of Macaca fascicularis mordax bm(nh) 1909.4.1.23 (Coll. No. 812), adult male, skin Thomas & Wroughton, 1 909c; holotype cataloged. and skull, collected at Sungei Biru, P. Bintan, Kepu- [Macacus] resimus: Raven, 1935, p. 237— geographic lauan Riau, Indonesia, by H. C. Robinson, date un- distribution. specified, presumably Jun. 1908; paratypes, bm(nh) Pithecus resimus: Elliot, 1 9 1 3, p. 224— external and cra- 1909.4.1.24 (Coll. No. 780, adult female, Sungei Biru, nial characters; taxonomic comparison. P. Bintan, 1 1 Jun.), bm(nh) 1 909.4. 1 .25 (Coll. No. 784, Pithecus lapsus Elliot, 1910, p. 343— holotype, usnm subadult female, Sungei Biru, P. Bintan, 12 Jun.), 124863 (Coll. No. 3418), adult male, skin and skull, bm(nh) 1 909.4. 1 .26 (Coll. No. 746, adult female, Pasir collected at Tanjung Pamuja, P. Bangka, Indonesia, Panjang, P. Bintan, 8 Jun.), and bm(nh) 1909.4.1.29 by W. L. Abbott, 1 9 Jun. 1 904; paratypes, usnm 1 24969 (Coll. No. 870, adult female, Tanjong Sauh, P. Batam, (Coll. No. 35 19, adult male, skin and skull fragments,

in 1 9 and 1 1 10 Jun.), skins and skulls, collected Kepulauan Riau, Jul.) usnm 24970 (Coll. No. 352 , adult male, Indonesia, by H. C. Robinson and/or E. Seimund, skin and skull, 20 Jul.), collected at Tanjung Batu, P. 1908. Elliot, 1913, p. 246— type series information. Belitung, Indonesia, by W. L. Abbott, 1904. Chasen, Robinson & Kloss, 1914, pp. 393, 394— type history; 1940a, p. 66— cited as a synonym of Macaca irus irus taxonomic comparison. H. C. Robinson, 1916, p. 62— F. Cuvier, 1818 (I. Geoffroy, 1826). Poole & Schantz, cited as a synonym of Pithecus fascicularis (Raffles, 1942, p. 243 -holotype cataloged. P. H. Napier, 1981, [1821]). Chasen, 1924b, p. 59— cited as a synonym of p. 13— cited as a synonym of M. fascicularis fascicu- Macaca irus F. Cuvier (I. Geoffroy, 1826). Chasen, laris (Raffles, [1821]). 1940a, p. 66— cited as a synonym of Macaca irus irus [Macacus] lapsus: Raven, 1935, p. 236— geographic dis- F. Cuvier (I. Geoffroy, 1826). P. H. Napier, 1981, pp. tribution. 13, 15— cited as a synonym of Macaca fascicularis Macaca irus lapsus: Sody, 1937, p. 247— specimens col- fascicularis (Raffles, [1821]); type history. lected in P. Bangka; external and cranial characters; [Macacus] bintangensis: Raven, 1935, p. 236— geo- taxonomic comparison. graphic distribution. Pithecus agnatus Elliot, 1910, p. 344— holotype, usnm

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 73 1 14409 (Coll. No. 1471), adult male, skin and skull, collected at Teluk Pai, P. Karimata, Indonesia, by W. collected in P. Tuangku, Kepulauan Banyak, Indo- L. Abbott, 24 Aug. 1 904; paratype (available but not nesia, by W. L. Abbott, 26 Jan. 1 902; paratypes, usnm explicitly cited in original description), usnm 125102 1 14408 (Coll. No. 1464, late subadult male, skin and (Coll. No. 3661), adult male, skin and skull, collected skull, 24 Jan.), usnm 111410 (Coll. No. 1472, adult at Teluk Pai, P. Karimata, Indonesia, by W. L. Abbott, male, skin and skull, 26 Jan.), usnm 1 1 44 1 1 (Coll. No. 27 Aug. 1904. Lyon, 1911, p. 137-type history. Chas-

?, subadult male, skull only, 28 Jan.), and usnm 1 14643 en, 1935b, p. 2— taxonomic comparison. Chasen, (Coll. No. ?, late juvenile male, skull only, 25 Jan.), 1940a, p. 66— cited as a synonym of Macaca irus irus collected in P. Tuangku, Kepulauan Banyak, Indo- F. Cuvier, 1818 (I. Geoffroy, 1826). Poole & Schantz, nesia by W. L. Abbott, 1902. Chasen, 1940a, p. 66- 1942, p. 242— holotype cataloged. P. H. Napier, 1981, cited as a synonym of Macaca irus irus F. Cuvier, 1818 p. 13— cited as a synonym of M. fascicularis fascicu-

(I. Geoffroy, 1826). Poole & Schantz, 1942, p. 241- laris (Raffles, [1821]). holotype cataloged. P. H. Napier, 1981, p. 13— cited [Macacus] carimatae: Raven, 1935, p. 236— geographic as a synonym of M. fascicularis fascicularis (Raffles, distribution. [1821]). [Macaca irus] carimatae: Sody, 1949, Table 1 —external [Macacus] agnatus: Raven, 1935, p. 236— geographic and cranial characters. distribution. Macaca irus? carimatae: Sody, 1 949, p. 1 3 1 —specimens [Macaca irus] agnatus: Sody, 1949, Table 1— external collected in P. Pelapis Tengah and P. Serutu, Kepu- and cranial characters. lauan Karimata, Indonesia. Pithecus lingungensis Elliot, 1910, p. 344— holotype, Pithecus mandibularis Elliot, 1910, p. 347— holotype, usnm 104853 (Coll. No. 492), adult male, skin and usnm 142225 (Coll. No. 4196), late subadult male, skull, collected in Pulo Lingung (= P. Lagong), Ke- skin and skull, collected at Sungei Sama (= Sungai pulauan Natuna, Indonesia, by W. L. Abbott, 19 Jun. Ambawang), near Pontianak, Kalimantan, Indonesia, 1900. Chasen, 1940a, p. 69— cited as a synonym of by W. L.Abbott, 18 Jun. 1905. Lyon, 1911, p. 137- Macaca irus pumila Miller, 1 900. Poole & Schantz, type history. Chasen & Kloss, 1931, p. 10— said to be 1942, p. 243-holotype cataloged. P. H. Napier, 1981, based on individual variables. Poole & Schantz, 1 942, p. 14— cited as a synonym of M. fascicularis pumila p. 244— holotype cataloged. P. H. Napier, 1981, p.

Miller, 1900. 13— cited as a synonym of M'. fascicularis fascicularis [Macacus] lingungensis: Raven, 1935, p. 236— geo- (Raffles, [1821]). graphic distribution. [Macacus] mandibularis: Raven, 1935, p. 236— geo- [Macaca irus] lingungensis: Sody, 1949, Table 1— ex- graphic distribution. ternal and cranial characters. Macaca irus mandibularis: Gyldenstolpe, 1920, pp. 3, M[acaca] /[ascicularis] lingungensis: Groves & Weitzel 14— specimen collected at Kaboerau, Kalimantan, In- in Weitzel et al., 1988, pp. 5, 96, 99— external char- donesia; said to be probably "not [valid] as a distinct acters; geographic distribution. race." Sody, 1949, p. 130— taxonomic comparison. Pithecus lautensis 345— usnm Elliot, 1910, p. holotype, [Macaca irus] mandibula: Sody, 1949, Table 1— incor- 104854 No. adult skin and (Coll. 614), male, skull, rect spelling of Pithecus mandibularis Elliot, 1910: ex- collected in P. Laut, Kepulauan Natuna, Indonesia, ternal and cranial characters. W. L. Abbott, 9 Aug. 1900. Chasen, 1940a, p. 69- by M[acaca]f[ascicularis] mandibularis: Groves & Weitzel cited as a synonym of Macaca irus pumila Miller, in Weitzel et al., 1988, pp. 5, 96, 99— external char- 1900. Poole & Schantz, 1942, p. 243— holotype cat- acters; geographic distribution. P. 1 98 1 1 cited as a aloged. H. Napier, , p. 4— synonym Pithecus baweanus Elliot, 1910, p. 347 — holotype, usnm of M. fascicularis pumila Miller, 1900. 151829 (Coll. No. 5565), adult male, skin and skull, [Macacus] lautensis: Raven, 1935, p. 236— geographic collected in P. Bawean, Java Sea, Indonesia, by W. L. distribution. Abbott, 24 Nov. 1907; paratype (available but not [Macaca irus] lautensis: Sody, 1949, Table 1— external explicitly cited in original description), usnm 151830 and cranial characters. — (Coll. No. 5566), adult female, skin and skull, collected Pithecus sirhassensis Elliot, 1 9 1 0, p. 345 holotype, usnm in P. Bawean, Indonesia, by W. L. Abbott, 24 Nov. 104852 (Coll. No. 468), subadult male, skin and skel- 1907. Lyon, 1911, p. 137-type history. eton, collected in Sirhassen I. (= P. Serasan), Kepu- [Macacus] baweanus: Raven, 1935, p. 236— geographic lauan Natuna, Indonesia, by W. L. Abbott, 8 Jun. distribution. 1900. Chasen, 1940a, p. 69— cited as a synonym of Macaca irus baweana: Chasen, 1940a, p. 70— geographic Macaca irus pumila Miller, 1 900. Poole & Schantz, — distribution. Poole & Schantz, 1 942, p. 24 1 holotype 1942, p. 245 -holotype cataloged. P. H. Napier, 1981, cataloged. p. 14— cited as a synonym of M. fascicularis pumila irus 132 — Miller, 1900. Macaca baweanus: Sody, 1949, p. specimens collected in P. taxonomic [Macacus] sirhassensis: Raven, 1935, p. 237— geograph- Bawean; comparison. ic distribution. M[acaca]f[ascicularis] baweana: J . R. Napier & Napier, [Macaca irus] sirhassensis: Sody, 1949, Table 1— exter- 1967, pp. 349, 403— geographic distribution. nal and cranial characters. Pithecus cupidus Elliot, 1910, p. 348— holotype, usnm M[acaca] f[ascicularis] sirhassensis: Groves & Weitzel 151831 (Coll. No. 5584), adult male, skin and skull, in Weitzel et al., 1988, pp. 5, 96, 99— external char- collected in P. Matasiri, Java Sea, Indonesia, by W. acters; geographic distribution. L. Abbott, 8 Dec. 1907; paratype (available but not Pithecus carimataeEWiot, 1910, p. 346— holotype, usnm explicitly cited in original description), usnm 154368 125101 (Coll. No. 3646), adult male, skin and skull, (Coll. No. 6388), adult male, cranium only, collected

74 FIELDIANA: ZOOLOGY in P. Matasiri, Indonesia, by W. L. Abbott, presum- graphic distribution; said to be possibly a junior syn- ably 1907-1908. Lyon, 1911, p. 137— type history. onym of [Pithecus] vitiis Elliot, 1 9 1 0, p. 346 (= Macaca [Macacus] cupidus: Raven, 1935, p. 236— geographic /ascicularis aurea I. Geoffroy, [1831]). Poole & Schantz, distribution. 1942, p. 241— holotype cataloged. Macaca irus cupida: Chasen, 1940a, p. 70— geographic M[acaca] /[ascicularis] capitalis: J. R. Napier & Napier, distribution. Poole & Schantz, 1942, p. 242— holotype 1967, pp. 349, 403— geographic distribution. cataloged. Macaca irus argentimembris Kloss, 191 la, p. 1 16— ho- [Macaca irus] cupidus: Sody, 1949, Table 1— external lotype, bm(nh) 1949.426 (Coll. No. 3814, SM [= Se- and cranial characters. langor Museum] 2068/10), adult male, skin and skull, M[acaca /[ascicularis] cupida: J. R. Napier & Napier, collected in P. Pinang [2], West Malaysia, by C. B. 1967, pp. 349, 403— geographic distribution. Kloss, 3 Sep. 1910; paratypes, bm(nh) 1955.1523 (Coll. Pithecus lingae Elliot, 1910, p. 349— holotype, usnm No. 3491, SM 2069/10, adult female, P. Pinang [2], 101603 (Coll. No. ?), subadult male, skin and skull, 3 Sep.), zrc 4-054 (Coll. No. 3488, SM 2070/10, late collected in Linga I. (= P. Lingga), Kepulauan Lingga, subadult male, P. Redang, 2 Sep.), and zrc 4-055 (Coll. Indonesia, by W. L. Abbott, 23 Jul. 1899; paratype, No. 3833, SM 2071/10, latejuvenile male, P. Redang, usnm 101602 (Coll. No. ?), late juvenile male, skin 5 Sep.), skins and skulls, collected in West Malaysia and skull, collected in P. Lingga, Indonesia, by W. L. by C. B. Kloss, 1910. J. E. Hill, 1960, p. 31 -type Abbott, 23 Jul. 1899. Chasen, 1925, p. 93 -cited as a history. Weitzel et al., 1988, p. 109— cited as a syn- synonym of Macaca irus F. Cuvier (I. Geoffroy, 1 826). onym of M. /ascicularis bintangensis (Elliot, 1 909). Chasen, 1940a, p. 66— cited as a synonym of Macaca [Macacus] irus argentimembris: Raven, 1935, p. 236—

irus irus F. Cuvier, 1818 (I. GeofTroy, 1826). Poole & geographic distribution. Schantz, 1942, p. 243— holotype cataloged. P. H. Na- Macaca /ascicularis argentimembra: Kloss, 1911a, pp. pier, 1981, p. 13— cited as a synonym of M. /ascicu- 177, 181 —external and cranial characters; taxonomic laris fascicularis (Raffles, [1821]). comparison; type series information. [Macacus] lingae: Raven, 1935, p. 236— geographic dis- Macaca/ascicularis argentimembris: J. R. Napier & Na- tribution. 403— distribution. P. — pier, 1967, pp. 349, geographic M[acaca] i[rus] lingae: Dammerman, 1926b, p. 316 H. Napier, 1981, pp. 13, 16— external characters; ho- geographic distribution. lotype cataloged. Pithecus impudens Elliot, 1910, p. 350— holotype, usnm Macaca [nemestrinus-Gruppe] aff. adusta: Elbert, 1912, 115675 (Coll. No. 1956), adult male, skin and skull, p. 101 (not Miller, 1906a, p. 559)— misidentification collected in P. Sugi, Kepulauan Riau, Indonesia, by of specimens collected in P. Sumbawa. Schwarz, 1912, W. L. Abbott, 24 Aug. 1902. Chasen, 1925, p. 93- p. 304— misidentification of specimens collected in P. cited as a synonym of Macaca irus F. Cuvier (I. Geof- Sumbawa. froy, 1826). Chasen, 1940a, p. 66— cited as a synonym Pithecus /ascicularis aff. limitis Schwarz, 1912, p. 304—

of Macaca irus irus F. Cuvier, 1 8 1 8 (I. Geoffroy, 1826). nomen nudum. Poole & Schantz, 1942, p. 242— holotype cataloged. Pithecus /ascicularis limitis Schwarz, 1913, p. 296— ho- P. H. Napier, 1981, p. 13— cited as a synonym of M. lotype, zsbs 1 9 1 1 /2 1 04 (Coll. No. 1 9), adult male, skin /ascicularis /ascicularis (Raffles, [1821]). and skull, collected at Lelogama, P. Timor, Indonesia, M[acaca] impudens: Raven, 1935, p. 236— geographic by C. B. Haniel, 28 May 1911; paratypes, zsbs 1911/ distribution. 2101 (Coll. No. 15, juvenile male, skin and skull, Fatu M[acaca] i[rus] impudens: Dammerman, 1926b, p. 316— Timau, NE, 1200 m, 20 May), zsbs 191 1/2102 (Coll. geographic distribution. No. 16, adult female, skin and skull, Lelogama, 22 Macaca irus? impudens: Sody, 1949, p. 129— specimens May), zsbs 191 1/2103 (Coll. No. 17, adult male, skin collected in P. Durian, Kepulauan Riau, Indonesia. only, Lelogama, 27 May), zsbs 1911/2105 (Coll. No. Pithecus capitalis Elliot, 1910, p. 350— holotype, usnm 21, infant male, skin and skull, Lelogama, 29 May),

83271 (Coll. No. ? ), adult male, skin and skull, col- zsbs 1 9 1 1/2 1 06 (Coll. No. 22, juvenile male, skin only, lected at Trong (= Ban Phra Muang, Trang Province), Lelogama, 30 May), zsbs 1911/2107 (Coll. No. 24, peninsular Thailand, by W. L. Abbott, 7 Mar. 1896; infant male, skin only, Lelogama, 3 Jun.), zsbs 1911/ paratype cited in original description, usnm 83272 21082 (Coll. No. 63, adult male, skin and skull, Kuat- (Coll. No. ?, adult male, skin and skull), collected in nana, 300 m, 20 Jul.), zsbs 191 1/2109 (Coll. No. 69, Telibon I. [= Ko Telibong], Thailand, by W. L. Ab- adult male, skull only, Bokong, 180 m, 26 Jul.), and bott, 27 Feb. 1896; paratypes available but not ex- zsbs 1911/2110 (Coll. No. 68, adult male, skin only, plicitly cited in original description, usnm 83273 (Coll. Bokong, 180 m, 24 Jul.) collected in P. Timor, In- No. ?, late juvenile male, skin and skull, Ban Phra donesia, by C. B. Haniel, 1911. Schwarz, 1914, p. Muang, 8 Mar.), usnm 83274 (Coll. No. 122, adult 117— type series information. female, skin and skull, Tyching, 19 May), and usnm M[acaca] /[ascicularis] limitis: J. R. Napier & Napier, 83275 (Coll. No. ?, infant, skin and skull, Tyching, 19 1967, pp. 349, 404— geographic distribution. May), collected in Trang Province, peninsular Thai- [Macacus] irus limitis: Raven, 1935, p. 236— geographic land, by W. L. Abbott, 1896. Kloss, 1919c, p. 347- distribution. taxonomic comparison. Weitzel et al., 1988, p. 109— Macaca irus limitis: Kellogg, 1945, p. 119— geographic cited as a synonym ofMacaca/ascicularis bintangensis distribution. (Elliot, 1909). Macaca irus limitus: Sody, 1949, p. 133, Table 1— in- Macaca capitalis: Gyldenstolpe, 1919, p. 131— geo- correct spelling of Pithecus/ascicularis limitis Schwarz, graphic distribution. 1913; taxonomic comparison. Macaca irus capitalis: Chasen, 1940a, pp. 68-69— geo- Pithecus mansalaris Lyon, 1916, p. 452— holotype, usnm

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 75 114560 (Coll. No. 1639), adult male, skin and skull, unlocated additional female specimen cited by Sody collected in Pulo Mansalar (= P. Mursala), west of in Table 1. W. C. O. Hill, 1974, p. 525 -cited as a Sumatra, Indonesia, by W. L. Abbott, 10 Mar. 1902; synonym of Macaca irus mordax Thomas & Wrough- paratypes, usnm 1 14559 (Coll. No. 1624, adult male, ton, 1909c. P. H. Napier, 1981, p. 13 -cited as a syn- skin and skull, 7 Mar.) and usnm 1 14561 (Coll. No. onym of Macaca fascicularis mordax Thomas &

?, adult female, skull only, 9 Mar.), collected in P. Wroughton, 1909c. Mursala, Indonesia, by W. L. Abbott, 1902. Chasen, Simia mauritius: W. C. O. Hill, 1974, p. 507 (not Grif- 1940a, p. 66— cited as a synonym of Macaca irus irus fith, 1821, p. 58; an "entirely black" monkey)— cited

F. Cuvier, 1818 (I. Geoffroy, 1826). Poole & Schantz, as a synonym ofMacaca irus irus. P. H. Napier, 198 1, 1942, p. 244— holotype cataloged. P. H.Napier, 1981, p. 13— cited as a synonym of Macaca fascicularis fas- p. 13— cited as a synonym of Macaca fascicularis fas- cicularis. cicularis (Raffles, [1821]). [Macacus] mansalaris: Raven, 1935, p. 236— geographic distribution. Holotype— Not preserved. Simia fascicularis [Macaca irus] mansalaris: Sody, 1949, Table 1 —external and cranial characters. Raffles, [1821] (p. 246) apparently is based on a — Pithecus sp.: Thomas, 1928, p. 832 misidentification single specimen, probably an adult female ("The at Vietnam. of juvenile male collected Tay Ninh, body is about twenty inches long, and the tail a Macaca mulatta: Thomas, 1928, p. 832 (not Zimmer- little more. . . . Canines short."). This specimen mann, 1780, p. 195)— misidentification of specimens was part of a collection made in Sumatra for Raf- (Coll. Nos. 777, 778) collected at Phu Quoc (not Phu- fles P. Diard and A. Duvaucel the Qui), Vietnam. by during period Macaca irus sublimitis Sody, 1932, p. 338— based on March 1819-March 1820 (T. S. Raffles in S. Raf- five specimens, unseen by Sody, reported by Dam- fles, 1830, pp. 703, 713); the original description merman (1928, p. 300) as Macaca irus mordax Tho- of Simia fascicularis is included in Raffles's pub- mas & Wroughton, 1909c; type series, mzb 865 (adult lished catalog of this collection. The fate of the female, Kambera [= Payeti-Kambaniru], 26 Mar.), mzb is unclear. 866 (adult male, Kambera, 27 Mar.), mzb 867 (juvenile holotype male, Mao Marroe, 4 May), mzb 868 (late juvenile Part of the Raffles/Diard/Duvaucel Sumatra Mao 8 and mzb 869 female, Marroe, May), (subadult collection, possibly accompanied by the manu- female, Mao Marroe, 10 May), collected in P. Sumba, script of Raffles's catalog, was sent to England in Indonesia, by P. F. Franck and/or Denin, 1925. March 1820, and another, larger part was sent to [Macaca irus] sumbae: Sody, 1933, p. 94— lapsus for in 1 Macaca irus sublimitis Sody, 1932. England or before June 820 (T. S. Raffles in Macaca irus sublimitus: Sody, 1 949, p. 1 34, Table 1 — S. Raffles, 1830, pp. 440, 447, 453, 715; Watson

incorrect of irus sublimitis 1 spelling Macaca Sody, 932; et al. in S. Raffles, 1830, p. 716). The text of Raf- taxonomic comparisons, distinctiveness of subspecies fles's catalog was read at a meeting of the Linnaean "still waits for thorough confirmation"; geographic Society of London on 5 December 1820. Three distribution; type series information. W. C. O. Hill, of M. 1974, p. 529— recognized provisionally, said to be specimens fascicularis (sexes unspecified) probably a synonym of Macaca irus mordax Thomas sent by Raffles from Sumatra ultimately reached & 1909c. Wroughton, the museum of the Zoological Society of London M[acaca] /[ascicularis] sublimitis: J. R. Napier & Na- (Waterhouse, 1838, p. 7; cf. Vigors, 1830, p. 642), pier, 1967, pp. 349, 404— geographic distribution. and two additional specimens sent by Raffles from irus 1 — Macaca submordax Sody , 1949, p. 133, Table Sumatra or Java reached the museum of the East holotype, rmnh unnumbered (Coll. No. E85), adult male, skin and skull, collected at Desa Poetjang, G. India Company, London (Horsfield, 1851, pp. iii- Agoeng (= Gunung [Mountain] Agung), E Bali, In- iv, 17). When these two museums were disband- H. V. date donesia, by J. Sody, unspecified, apparently ed—the former in 1852-1860 and the latter in 1927-1931; paratypes, rmnh (Coll. No. E34, juvenile 1879— their collections were transferred to the male, skin and skull, Sendang), rmnh (Coll. No. E64, but none of adult male, skin and skull, Sendang), rmnh (Coll. No. bm(nh) (Thomas, 1906, pp. 40, 63), E74, adult male, skin and skull, Sendang), rmnh (Coll. Raffles's specimens of M. fascicularis are now pre- No. El 36, adult male, skull Jembrana, only, Negara served in the bm(nh) (cf. P. H. Napier, 1981, p. district), and rmnh (Coll. No. El 39, subadult female, 16). skin and skull, Jembrana, Negara district), collected Another part of the Sumatra collection was in Bali, Indonesia, by H. J. V. Sody, apparently 1927—

1931; mzb 2001 (Coll. No. 1), adult female, skin and transferred by Raffles to Diard and Duvaucel and skull, collected at Batu Meringgit, Bali, Indonesia, by subsequently was sent to France by Duvaucel in P. F. Franck, 8 Oct. 1928; mzb 6521 (Coll. No. 17/ 1820-1821 (T. S. Raffles in S. Raffles, 1830, p. 92/33, juvenile male), mzb 6522 (Coll. No. 15/93/33, 720; Lacaze, 1856, col. 536). Included in this part adult male), and mzb 6523 (Coll. No. 16/04/33, adult of the collection was an adult female of male), skins and skulls, collected at Banju Wetan, Bali, specimen Indonesia, by J. J. Menden, 18 Jul. 1933; and one M. fascicularis (mnhn 362/233), which coinciden-

76 FIELDIANA: ZOOLOGY tally is a paralectotype of M. f. aurea I. Geoffroy, Table 3, Appendix 4); crown same color as back, [1831] (see below). This female, however, almost or brighter (847 of 873 postinfantile specimens certainly is not the holotype of Simla fascicularis; examined), occasionally with an indistinct black- Raffles considered the specimens that he trans- ish wash (24 specimens), rarely contrastingly darker

ferred to Diard and Duvaucel to be "duplicates," than back (2 specimens; cf. Table 5, Appendix 5); not the "originals," which he cataloged and sent preauricular hairs directed anteriorly, forming part to England. of lateral facial crest (crest transzygomatic, 501 of Type Locality— Sumatra, restricted to "neigh- 504 specimens examined), rarely directed poste- bourhood of Bencoolen [= Bengkulu]" by H. C. riorly, partly covering ears (crest infrazygomatic, Robinson (1 9 1 6, p. 63). Bengkulu, in southwestern 2 specimens; crest asymmetric, 1 specimen; cf. Sumatra, was the headquarters of Raffles, Diard, Table 6, Appendix 6); T 67-150% of HB in 311

and Duvaucel (cf. Steenis-Kruseman, 1 950, p. 425). adult specimens examined (cf. Table 10, Appendix Distribution (Fig. 25)— The broad geographic 9).

range of M. f. fascicularis in Southeast Asia in- Remarks— For comments on nine quasi-dis- cludes the following continental and insular com- tinct populations that are here included within M. ponents: the southern part of the Indochinese Pen- f fascicularis, see above (Subspecific Taxonomy). insula (including southern Vietnam, Cambodia, Specimens Examined— Total, 1,550: skins and and southern Thailand) and adjacent islands (ex- skulls, 739; skins only, 285; skulls only, 526 (see

cluding the Con Son group and Ko Khram Yai); Appendix 1). the northeastern part of the Isthmus of Kra; the Malay Peninsula and adjacent islands; Sumatra and adjacent islands (excluding the Nicobar Is- lands, the P. Simeulue group, Kepulauan Menta- Macaca fascicularis aurea wai, and P. Enganno); Borneo and adjacent islands I. Geoffroy, [1831], p. 58 (excluding P. Maratua), Java and adjacent islands (excluding P. Karimunjawa and P. Kemujan); Macacus aureus I. Geoffroy, [1831], pp. 58, 76, pi. 2— for details see below. Lesser Sunda Islands from Bali to Timor (exclud- concerning type series, Eydoux & Souleyet [& Gervais], 1841, p. 6, pi. 2— taxonomic ing P. Komodo and many nearby islets); and the comparisons. I. Geoffroy, [1843], p. 566— taxonomic south-central Philippines Sulu, (including Cagayan history. Gray, 1849, p. 4— cited as a synonym of Ma- Sulu Archipelago, and western Mindanao). At the cacus cynomolgus [= Macacafascicularis]. I. Geoffroy, margins of the subspecific range are three inter- 1851, p. 27— taxonomic comparison; type series in- formation. Blyth, 1875, p. 7— cited as a of subspecific or interspecific contact zones, inhab- synonym Macacus cynomolgus [= Macaca fascicularis]; said to ited by mixed-phenotype populations, as follows: be based on "a casual individual variety from Pegu." central and eastern of the Indochinese (1) parts J. Anderson, 1879, p. 73— cited as a synonym of Ma- Peninsula— an intersubspecific contact zone be- cacus cynomolgus [= Macaca fascicularis]; type said to be a market specimen. Elliot, 1913, p. tween ranges of M. f. fascicularis and M.f aurea, probably 229— cited as a synonym of Pithecus irus [= Macaca or an interspecific contact zone between ranges of fascicularis]. Rode, 1938, p. 223— cited as a synonym M. and M. mulatta; (2) eastern and fascicularis of Macaca irus (F. Cuvier, 1818; I. Geoffroy, 1826); southern parts of the Isthmus of Kra and adjacent holotype cataloged. southwestern islands— an intersubspecific contact Macacus auratus Miiller, [1840], p. 49 (not E. Geoffroy, 1812, p. 93)— incorrect spelling of Macacus aureus I. zone between ranges of M. f. fascicularis and M. Geoffroy, [1831]. Blyth, 1863, p. 9-cited as a syn- f aurea; and (3) eastern and central Mindanao, onym of Macacus carbonarius F. Cuvier (I. Geoffroy, southern Negros, and possibly nearby islands— an 1826). intersubspecific contact zone between ranges ofM. I[nuus] aureus: Wagner, [1839], p. 138 (part)— external characters. f. fascicularis and M. f philippinensis (see below, Pith[ecus] (Mac[acus]) aureus: Dahlbom, 1856, p. 118 — M. f. aurea— Distribution; M. f. philippinensis— external characters.

Distribution). Macaca irus aurea: Pocock, 1939, p. 79— taxonomic diagnosis— General color of dorsal surface of comparison; type locality restricted to Pegu. trunk buffy to yellowish gray to medium brown [Macaca irus] aureus: Sody, 1 949, Table 1 —external and cranial measurements. (825 of 868 postinfantile specimens examined), M[acaca] f[ascicularis] aurea: J. R. Napier & Napier, occasionally dark brown (43 specimens; cf. Table 1967, pp. 349, 403— distribution. 1 hair an- , Appendix 3); erythrism variable, pale Cercopithecus Cynosurus: Heifer, 1838, p. 858 (not Sco- nulations pale yellowish to golden to rufescent (cf. poli, 1786, p. 44)— misidentification; natural history.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 77 476 I. 1 M[acacus] cynomolgus(?): Blyth, 1844, pp. 474, (not p. 223), male, skin ("poils uses," Geoffroy, [ 843], Linnaeus, 1766, p. 38)— specimens collected in Ara- p. 567) with or without skull, obtained in Java (no kan. further locality information available), by P. Diard, Macacus cynomolgus: Blyth, 1875, p. 7 (part; not Lin- 1821. I. indicated naeus, 1766)— taxonomic comparisons. Geoffroy ([1831], pp. 58, 76) M\acacus] carbonarius: Blyth, 1847, p. 732 (not I. Geof- that dorsal pelage in all four of these specimens said to be in Arakan. froy, 1826)— common was conspicuously erythristic ("Dessus du corps C\ynamolgus] mulatta: Reichenbach, 1862, p. 136 (part; d'un beau roux tiquete de noir") and cited this as not Zimmermann, 1780, p. 195)— misidentification; the primary diagnostic character of Macacus au- taxonomic history. Pithecus vitiis Elliot, 1910, p. 346— holotype, usnm reus, "Le Macaque Roux Dore." However, the 124176 (Coll. No. 3076), subadult male, skin and infrazygomatic lateral facial crest (see above, Pel- mandible lost Jan. collected at Domel (cranium 1938), age) that is now regarded as diagnostic of Macaca I. (= Letsok-aw Kyun), Mergui Archipelago, Burma, fascicularis aurea is clearly figured by I. Geoffroy by W. L. Abbott, 26 Jan. 1904; paratypes, usnm 104442 PI. in his illustration of either the "Ben- (Coll. No. 287, late juvenile male, Sullivan's I. [= ([1831], 2) Lanbi Kyun], 30 Jan.) and usnm 111898 (Coll. No. gale" or Pegu specimen and is alluded to in his 757, adult female, St. Matthews I. [= Zadetkyi Kyun], detailed description of the illustrated specimen 9 Dec), skins and skulls, collected in Mergui Archi- ("longs poils blancs diriges en arriere . . . cachent pelago, Burma, by W. L. Abbott, 1900. Pocock, 1939, en partie les oreilles," p. 77). p. 79— cited as a synonym of Macaca irus aurea. Chas- en, 1 940a, p. 68 — said to be possibly a senior synonym No holotype is designated in the original de- of Pithecus capitalis Elliot, 1910, p. 350 (= M. fasci- scription of Macacus aureus I. Geoffroy, [1831]. cularisfascicularis (Raffles, [1821 ])). P. H. Napier, 1981, In I. Geoffroy's (1851, p. 27) catalog of primates p. 13— cited as a synonym of Macaca fascicularis au- in the the 'un des rea. mhnh, "Bengale" specimens ("L de and the Sumatra Macaca vitiis: Poole & Schantz, 1942, p. 246 — holotype types respece") specimen cataloged. {"L'un des types") are treated as coordinate type external and [Macaca irus] vitiis: Sody, 1949, Table 1— specimens of this taxon; the Java specimen is cranial measurements. transferred to another taxon (p. 29), and the Pegu Pithecus fascicularis: Wroughton, 1915, p. 699 (part; not specimen is not mentioned. Ambiguous state- Raffles, [1821], p. 246)— taxonomic history. ments by J. Anderson (1879, p. 76) and Elliot (1913, p. 231) may or may not constitute desig- Type Series — Macacus aureus I. Geoffroy, nations of the "Bengale" specimen as lectotype. [1831, pp. 58, 76] is explicitly based on four spec- The "Bengale" specimen is decisively designated imens of Macaca fascicularis: (1) mnhn 362/234, as lectotype by Rode (1938, p. 223; cf. Interna- Type Cat. No. 58a (lectotype, designated by Rode, tional Code of Zoological Nomenclature, 1985, 1938, p. 223), subadult or adult male, mounted Article 74). skin only (skull measured by Elliot, 1913, p. 231; Type Locality— Pegu, Burma (restricted by reported absent by Rode, 1938, p. 223), obtained Pocock, 1 939, p. 79; cf. International Code ofZoo- in "Bengale" (now partly in eastern India and part- logical Nomenclature, 1985, Article 72h). The lec- ly in Bangladesh) by L. T. Leschenault de la Tour, totype of Macacus aureus I. Geoffroy, [183 1], ob- presumably in 1819-1820, acquired by the mnhn tained in "Bengale," probably was a captive pur- in Jul. 1822 (cf. Leschenault de la Tour, 1820, p. chased in Calcutta (J. Anderson, 1879, p. 76); sim- 359; 1822, p. 262; I. Geoffroy, 1851, p. 27; Rode, ilar captive specimens in the Calcutta market, or 1938, p. 223; Steenis-Kruseman, 1950, p. 321); nearby, have been reported by C. Belanger (in I. (2) unpreserved specimen (not listed in catalog of Geoffroy, [1831], p. 77), Eydoux and Souleyet [&

I. 1 1 Geoffroy, 85 , p. 27), sex unspecified, obtained Gervais] (1841, pp. xiv, 6), and J. Anderson (1879, in "Pegou" (= Pegu, Burma) by A. A. M. Reynaud, p. 76). The original provenance of the lectotype is

1827-1828 (cf. de Rossel et al., 1829, p. 603; de unknown. Based on the locality of one of the para- Rossel, 1829, p. 609); (3) mnhn 362/233, Type lectotypes (see above), Pocock (1939, p. 79) Cat. No. 58b, adult female, mounted skin only, amended the type locality to Pegu. obtained in Sumatra (no further locality infor- Distribution (Figs. 9, 25) — Southernmost mation available) by A. Duvaucel, 1 8 1 9-1 820, ac- Bangladesh, southwestern and southern Burma

quired by the mnhn 7 Sep. 1821 (cf. I. Geoffroy, (including Preparis I.), adjacent islands in the Mer- 1851, p. 27; Rode, 1938, p. 223; de Lacaze, 1856, gui Archipelago, and a small area in west-central col. 535); and (4) unpreserved specimen (listed by Thailand; most of the geographic range is west of I. Geoffroy, 1851, p. 29; not listed by Rode, 1938, the mountain chains that form the border between

78 FIELDIANA: ZOOLOGY Burma and Thailand. Adjacent to the geographic cluding Balabac, Palawan, Culion, Mindoro, Lu- range of M.f. aurea are two areas— one in central zon, Samar, Leyte, and probably other islands north and eastern parts ofthe Indochinese Peninsula and of about 10°N (Fooden, 1991, pp. 6, 23). Based the other in eastern and southern parts of the Isth- on two available skins, northeastern Mindanao mus of Kra— where specimens with the diagnostic formerly was also included in the range of this

1 1 lateral facial crest character of M. f. aurea and subspecies (Fooden, 99 , p. 23); however, in June specimens with the diagnostic lateral facial crest 1991 one of the critical specimens (bm(nh) character of M. f. fascicularis have been collected, 1877.10.6.2, Butuan River [= Agusan River]) was sometimes at the same locality. The area in eastern discovered to consist of a late infant or early ju- and southern parts of the Isthmus of Kra may be venile skin mismatched with a subadult female regarded as a M.f. aurea/M.f. fascicularis contact skull, which weakens the evidence for retaining zone; the area in central and eastern parts of the northeastern Mindanao in the subspecific range. Indochinese Peninsula may be either a M.f. aurea/ In eastern and central Mindanao, southern Ne- M.f. fascicularis contact zone or a M. fascicularis/ gros, and possibly in nearby islands, between the M. mulatta contact zone (see below, Evolution and ranges of M. f philippinensis and M.f. fascicularis, Dispersal). is a contact zone inhabited by mixed-phenotype Diagnosis— General color of dorsal surface of populations. trunk buffy to medium brown, pale hair annula- Diagnosis— General color of dorsal surface of tions pale yellowish to golden, rarely rufescent; trunk dark brown, pale hair annulations golden to crown usually brighter than back, occasionally with refescent (pale golden in 1 subadult male, fmnh an indistinct blackish wash; preauricular hairs di- 87718); crown brighter than back (indistinct rected posteriorly, partly covering ears (lateral fa- blackish streak in 1 adult male, bm(nh) cial crest infrazygomatic); T 76-104% of HB in 12 1877.10.6.1); preauricular hairs directed anteri- adult specimens examined. For external and cra- orly, forming part of lateral facial crest (crest nial measurements, see Table 36. transzygomatic); T 101-138% of HB in 14 adult Specimens Examined—M. f aurea, 53: skins specimens examined. For further details, see and skulls, 42; skins only, 8; skulls only, 3. Inter- Fooden (1991, pp. 3 ff). subspecific or interspecific contact zones, 45: skins Specimens Examined—M. f philippinensis, 98: and skulls, 41; skins only, 3; skull only, 1 (see skins and skulls, 53; skins only, 15; skulls only,

Appendix 1). 30. Probably M. f philippinensis, 15: skin only, 1; skulls only, 14. M.f. philippinensis/M. f fascicu- laris contact zone, 110: skins and skulls, 82; skins only, 5; skulls only, 23. Probably M. f philippi- Macaca fascicularis philippinensis nensis/'M.ffascicularis contact zone, 12: skins and

I. skin 1 skulls Geoffroy, [1843], p. 568 skulls, 2; only, ; only, 9 (see Appendix

1). For synonymy, see Fooden (1991, pp. 22, 24). Holotype— Macacus philippinesis I. Geoffroy, [1843, p. 568], is based solely on mnhn 373 (265), albino adult male, mounted skin only (omitted Macaca fascicularis umbrosa from type catalog published by Rode, 1938, p. Miller, 1902b, p. 789 222). This monkey was obtained alive in Manila A. Chenest and him to the me- by presented by Cercopithecus cynomolgus: Cantor, 1846, p. 176 (part; nagerie of the mnhn on 6 Aug. 1841; it died on not Linnaeus, 1766, p. 38)— Nicobar Islands included

1 I. 1 1 in distribution. 29 Aug. 842 (cf. Geoffroy, 85 , p. 29). A figure Macacus cynomolgus: Barbe, 1846, pp. 365 (not Lin- of the holotype accompanies the original descrip- naeus, 1766)— sight records. Blyth, 1846, p. 367 tion (pi. 5). (part) — zoogeography. Type Locality— Philippine Islands, probably M[acacus] cynomolgos: Blyth, 1863, p. 9 (part; not G. collected in "Nico- Luzon ("Chenest . . . l'a acquis a Manille, et il le Cuvier, 1798, p. 98)— specimen bars." croit originaire de cette ile," I. Geoffroy, [1843], Cercocebus carbonarius:Ze\eboT, [ 1 869], p. 7 (not I. Geof- p. 570). froy, 1826, p. 588)— cited as "Var. a" of Inuus cy- Distribution and (Fig. 25)— Western, northern, nomolgus; specimen collected in Great Nicobar I. eastern islands of the Philippine Archipelago, in- M[acacus] carbonarius: J. Anderson, 1881, p. 65 (part;

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 79 Table 36. Macaca fascicularis aurea: external and cranial variation in adult specimens examined. Macaca fascicularis fusca specimens); subauricular hairs pale gray, not elon- 476 Miller, 1903a, p. gated; outer surface of thighs and shanks blackish; T 90-108% of HB in 10 adult specimens exam- — Macacus fuscus Miller, 1903a, p. 476 for details con- ined. For external and cranial measurements, see see below. & cerning type series, Lyon Osgood, 1909, Table 37, Figures 26 and 27, and Appendix 12. p. 283— holotype cataloged. Remarks— For comments on T and crown hair Macaca fusca: Thomas, 1923, p. 591— specimen col- lected in P. Simeulue. annulation in M. fascicularis fusca, see below (M. [Cynomolgus] fuscus: Trouessart, 1904, p. 16— geo- f. lasiae— Remarks). Behavior in M. fascicularis graphic distribution. fusca has been compared with that in Sumatran Pit hecus fuscus: Elliot, 1913, p. 228— external and cra- M. f. fascicularis by van Schaik and van Noord- nial characters. wijk (1985a, p. 141) and Sugardjito et al. (1989, Pithecus fuscus fuscus: Lyon, 1916, p. 457— geographic distribution. p. 197). These authors suggest that sexual dimor- Macaca irus fusca: Chasen, 1940a, p. 69— geographic phism of size may be less in M. fascicularis fusca distribution. Poole & Schantz, 1 942, p. 242— holotype than in Sumatran M. f. fascicularis; however, re- cataloged. Kellogg, 1944, p. 76— taxonomic compar- duced sexual dimorphism is not apparent in avail- isons; zoogeography. able cranial measurements of M. M[acaca] f[ascicularis] fusca: J. R. Napier & Napier, fascicularisfusca 1967, pp. 349, 403— geographic distribution. (Table 37, Fig. 26). Specimens Examined— Total, 19: skins and

Type Series— Holotype (by original designa- skulls, 12; skins only, 7 (see Appendix 1). tion), usnm 1 14164 (Coll. No. 1348), adult male, skin and skull, collected at Telok Dalam (= Lhok Macaca fascicularis lasiae Dalam), east coast of Simalur I. (= P. Simeulue), (Lyon, 1916, p. 453) Indonesia, by W. L. Abbott, 20 Nov. 1901 (for collecting locality details, see collector's field cat- Macacusfuscus: Miller, 1903a, p. 476 (part)— specimens alog, usnm); paratypes, usnm 114162 (Coll. No. collected in P. Lasia; taxonomic comparisons. 1346, adult female, P. Simeulue, 18 Nov. 1901), Pithecus fuscus lasiae Lyon, 1916, p. 453— for details usnm 1 14163 (Coll. No. 1347, adult male, P. Si- concerning type series, see below. W. C. O. Hill, 1 974, 5 1 1 —cited as a of Macaca irus Mil- meulue, 18 Nov. 1901), usnm 114165 (Coll. No. p. synonym fusca ler, 1903a. P. H. Napier, 1981, p. 13-cited as a syn- 1349, adult female, P. Simeulue, 20 Nov. 1901), onym of Macaca fascicularis fusca Miller, 1903a. usnm 114166 (Coll. No. 1350, adult female, P. Macaca irus lasiae: Chasen, 1940a, p. 69— geographic Simeulue, 20 Nov. 1901), usnm 114167 (Coll. No. distribution. Poole & Schantz, 1942, p. 242— holotype 1354, adult male, P. Simeulue, 26 Nov. 1901), cataloged. Kellogg, 1944, p. 76— taxonomic compar- isons; zoogeography. usnm 114168 (Coll. No. 1387, adult male, Siba- boh, P. Simeulue, 17 Dec. 1901), usnm 114169 Type Series— Holotype (by original designa- (Coll. No. 1396, adult male, Badjan Bay tion), usnm 1 14248 (Coll. No. 1398), adult male, [= Labuhanbajau], P. Simeulue, 1 Jan. 1902), usnm skin and skull, collected in P. Lasia, southeast of 114247 (Coll. No. 1397, late juvenile male, P. P. Simeulue, Indonesia, by W. L. Abbott, 5 Jan. Lasia, 4 Jan. 1902), and usnm 1 14248 (Coll. No. 1902; paratype, usnm 114247 (Coll. No. 1397), 1398, adult male, P. Lasia, 5 Jan. 1902), skins and late juvenile male, skin and skull, collected in P. skulls, collected in Indonesia by W. L. Abbott. Lasia, Indonesia, by W. L. Abbott, 4 Jan. 1902. Type Locality— Telok Dalam (= Lhok Dal- These two specimens also are paratypes of M. fas- am), east coast of Simalur Island (= P. Simeulue), cicularis fusca Miller, 1903a, p. 476 (see above; west of northern Sumatra, Indonesia. cf. International Code of Zoological Nomencla- Distribution (Fig. 25)— P. Simeulue, west of ture, 1985, Article 72d). northern Sumatra, Indonesia. Type Locality— Palau Lasia, southeast of P. Diagnosis— General color of dorsal surface of Simeulue, west of northern Sumatra, Indonesia. trunk blackish, pale hair annulations pale yellow- Distribution (Fig. 25)— Pulau Lasia, west of ish; crown blackish, hairs conspicuously annulated northern Sumatra, Indonesia. with pale yellowish; preauricular hairs usually di- Diagnosis— General color of dorsal surface of rected posteriorly, partly covering ears (lateral fa- trunk blackish, pale hair annulations pale yellow- cial crest infrazygomatic, 9 of 13 specimens ex- ish; crown blackish, hair annulations inconspic- amined), occasionally directed anteriorly, forming uous; preauricular hairs directed posteriorly, part- part of lateral facial crest (crest transzygomatic, 4 ly covering ears (lateral facial crest infrazygomatic,

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 81 Table 3 7 . Sexual dimorphism ofgreatest length of skull (mm) compared in Macacafascicularisfusca and Sumatran Macaca fascicularis fascicularis (cf. Fig. 26).

Table 38. Comparison of cranial proportions in Indochinese (I-C) Macaca fascicularis, M. f. atriceps, and M. J condorensis (cf. Fig. 28; Kloss, 1919c, p. 348; 1921, p. 76; 1926, p. 358).

Diagnosis— General color of dorsal surface of gated; outer surface of thighs brownish gray, be- trunk buffy to medium brown (with blackish mid- coming pale brownish gray on shanks; T 1 26-1 3 1 % dorsal streak in 2 juveniles, zrc 4-017, 4-018), of HB in 2 adult specimens examined. For external pale hair annulations pale yellowish to golden; and cranial measurements, see Appendix 12. crown with a broad dark brown to blackish patch Specimens Examined— Total, 4: skins and skulls,

(Fig. 7) that extends laterally as far as lateral mar- 4 (see Appendix 1). gin of each eye and posteriorly as far as vertex or occiput, margin of crown patch not sharply delim- ited; preauricular hairs directed anteriorly, form- ing part of lateral facial crest (crest transzygo- Macaca fascicularis karimondjawae matic); T 109-127% of HB in 6 adult specimens Sody, 1949, p. 132 examined. For external and cranial measure- ments, see Table 38, Figure 28, and Appendixes Macacus cynomolgus: Willink, 1905, p. 175 (part, not Linnaeus, 1766, p. 38)— "P. Karimon-Djawa (P. Ka- 7-9 and 12 and Fooden and Albrecht (1993, p. moedian)" included in geographic distribution. 537). Macaca irus karimondjawae Sody, 1949, p. 132, Table — — Re- Remarks See above (M. / atriceps 1 —for details concerning type series, see below. W. C. marks). O. Hill, 1974, p. 528— external and cranial characters. Macaca P. H. Specimens Examined— Total, 16: skins and fascicularis karimondjawae: Napier, 1981, p. 13— type locality information. 1 1 3 skulls, ; skins only, 2; skulls only, (see Ap- pendix 1). Type Series— Holotype (by original designa- tion), rmnh 10608 (mzb 2719), adult male, skin and collected in Karimon P. Kar- Macaca fascicularis tua skull, Djawa (= imunjawa), Java Sea, Indonesia, by W. Rom- Kellogg, 1944, p. 75 swinckel, 28 Nov. 1930 (incorrectly given as "28.VI.1930" in original description); paratypes Macaca irus tua Kellogg, 1944, p. 75 — for details con- cerning type series, see below. W. C. O. Hill, 1974, p. (cf. Sody, 1949, p. 132, Table 1), mzb 1454 (adult 507— cited as a synonym of Macaca irus irus F. Cu- male, cranium only, 7 May), mzb 1455 (adult fe- vier, 1818 1826). P. H. Napier, 1981, p. (I. Geoffroy, male, skin and skull, 9 May), mzb 1456 (juvenile 13— cited as a synonym of Macaca fascicularis fasci- female, skin and skull, 10 May), mzb 1457 (adult cularis (Raffles, [1821]). skin and 11 and mzb 1458 M[acaca]f[ascicularis] tua: J. R. Napier & Napier, 1967, female, skull, May), pp. 349, 404— geographic distribution. (infant male, skin and skull, 14 May), collected in P. Karimunjawa, Indonesia, by K. W. Dammer- Type Series— Holotype (by original designa- man, P. F. Franck, and/or Denin, 1926; mzb 2717 tion), usnm 197663 (Coll. No. 626), adult male, (juvenile male) and mzb 2718 (late juvenile fe- skin and skull, collected in Pulo Muara Tua (= P. male), skins and skulls, collected in P. Karimun- Maratua), east of Borneo, Kalimantan, Indonesia, jawa, Indonesia, by W. Romswinckel, 26 Nov. by H. C. Raven, 21 May 1913; paratypes, usnm 1930; and mzb 2720 (adult female, cranium only) 197660 (Coll. No. 623, juvenile female), usnm collected in P. Kemujan, < 1 km northeast of P. 197661 (Coll. No. 624, adult female), and usnm Karimunjawa, Indonesia, by W. Romswinckel, 25 197662 (Coll. No. 625, subadult male), skins and Nov. 1930. skulls, collected in P. Maratua, Kalimantan, In- Type Locality— Karimon Djawa (= P. Kari- donesia, by H. C. Raven, 21 May 1913. munjawa), Java Sea, Indonesia. Type Locality— Pulau Maratua, Kalimantan, Distribution (Fig. 25)— Pulau Karimunjawa Indonesia. and, presumably, nearby P. Kemujan (known from Distribution (Fig. 25)— Pulau Maratua, east of one cranium only), Java Sea, 60 km north of cen- northeastern Kalimantan, Indonesia. tral Java, Indonesia. Diagnosis— General color of dorsal surface of Diagnosis— General color of dorsal surface of trunk blackish, pale hair annulations pale yellow- trunk dark grayish brown, pale hair annulations ish; crown pale yellowish brown, hairs conspicu- pale yellowish; crown frequently with a blackish ously annulated with pale yellowish; preauricular wash; preauricular hair direction not studied; T hairs directed anteriorly, forming part of lateral 101-1 12% of HB in 3 adult specimens examined. facial crest (crest transzygomatic); subauricular For external and cranial measurements, see Ap- hairs pale ochraceous-buff, conspicuously elon- pendix 12 and Fooden and Albrecht (1993, p. 538).

86 FIELDIANA: ZOOLOGY Specimens Examined— Total 9: skins and skulls, the penultimate glacial maximum, ca. 1 60 Ka, sea

6; skin only, 1; skulls only, 2 (see Appendix 1). level was about 160 m lower than at present, there- by adding some deep-water fringing islands to the temporarily consolidated large Sunda Shelf land- mass. Similar cyclic sea-level changes, of impre- Macaca fascicularis Subspecies Undetermined cisely known magnitude, have occurred with a pe- riodicity of ca. 100 Ka since the beginning of the Macacus cristatus Gray, 1870, p. 30— holotype, bm(nh) Pleistocene. 1858.4.28.9, late juvenile male, skin (albinistic) and skull, acquired in 1858 from collection of Th. G. van Tectonic forces also have affected the distribu- Lidth de Jeude, Utrecht, provenance unknown. Schle- tion of land and sea within the range of M. fas- 101— cited as a of Cercocebus gel, 1876, p. synonym cicularis. Prominent examples of such effects are cynamolgos: Schlegel (= Macaca fascicularis (Raffles, the approximately 500-m uplift of the north coast [1821])). Elliot, 1913, p. 249-cited as a synonym of of P. Sumba et al., the Pithecus philippinensis (I. Geoffroy, [1843]). W. C. O. (Pirazzoli 1991, p. 1835), Hill, 1974, pp. 477, 522— cited as a synonym of Ma- approximately 500-m subsidence and subsequent caca irus I. P. H. Na- philippinensis Geoffroy, [1843]. reelevation of P. Maratua (Kuenen, 1947, p. 8), pier, 1981, pp. 13, 20— cited as a synonym of Macaca and the massive volcanic explosions of Rakata (= I. Food- fascicularis philippinenesis Geoffroy, [1843]. and Toba 1 1 Krakatau) (Dammerman, 1948, p. 4; en, 99 , p. 24— subspecies not determined; taxonom- Chesner et 1 1 ic history. al., 99 , p. 200; Rampino & Self, 1993, Macacus albus: P. H. Napier, 1981, p. 20— unavailable p. 269). However, the zoogeographic implications author written on of manuscript name, unknown, tag of Pleistocene tectonic changes in this area are holotype of Macacus cristatus Gray, 1870. more localized than those of Pleistocene eustatic changes. 2. Swimming. The swimming distance limit of M. fascicularis probably is about 1 00 m (see above, Evolution and Dispersal Natural History). 3. Natural rafting. Monkeys may be transport- Available information concerning the distri- ed across water gaps on floating trees or larger bution, variation, natural history, and paleontol- entwined masses of vegetation (cf. Wallace, 1895, ogy of M. fascicularis provides a basis for hypo- p. 74). The tendency for M. fascicularis to exploit thetical interpretation of the evolution and dis- coastal habitats (see above, Natural History) makes persal of this species. Because most of the geo- this species a prime candidate for such passive graphic range of M. fascicularis is insular, a large dispersal on natural rafts. A specific instance is part of this interpretation concerns water gaps and reported by Hickson (1889, p. 190): "Some days their effect on dispersal. Four major factors are after the eruption of in 1883 a female pertinent to dispersal in the insular part of the green monkey [M. fascicularis] was found floating range of M. fascicularis. on some drifting timber in the Sunda Straits. She 1. Eustatic changes in sea level. Since the be- was terribly scorched, but completely recovered, ginning of the Pleistocene, ca. 1.75 Ma, worldwide and is, I believe, still alive." sea level has been strongly controlled by, and in- The average surface current speed in the Sunda versely correlated with, the cyclic growth of con- Shelf area is about 1 km/hr (E. G. W. Smith, 1974, tinental glaciers (Van Couvering & Kukla, 1988, Fig. 2.3-1; Couper, 1989, p. 50). Arbitrarily as- p. 459; Heaney, 1991a, p. 56; Cande & Kent, 1 992, suming that some members of a small group of pp. 13,936, 13,938). Sea level has been approxi- M. fascicularis could survive on a natural raft for mately as at present since ca. 5 Ka (Clark et al., up to about 3 days, such a group potentially could 1978, p. 283; Fairbanks, 1989, p. 639). It rose colonize a new island about 75 km distant from sigmoidally to this level from about 1 20 m lower its original habitat; during monsoons and other during the last glacial maximum, ca. 1 8 Ka, when storms, the current would be stronger and the dis- the Sunda Shelf was exposed and the Indochinese tance covered would be greater. The reproductive Peninsula, the Malay Peninsula, Sumatra, Borneo, capability of a small group of M. fascicularis in- Java, and shallow-water fringing islands were unit- dividuals to multiply and populate islands not pre- ed to form a single large landmass (Fig. 3). During viously inhabited by this species is demonstrated the preceding interglacial, ca. 120 Ka, sea level by the success of introduced populations in Mau- was high, and components of this large landmass ritius and Angaur (Poirer & Smith, 1 974, p. 264; were isolated, approximately as at present. During Sussman & Tattersall, 1986, p. 30).

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 87 Table 39. in evolution and dis- Hypothetical stages Available evidence indicates a minimum of six persal of Macaca fascicularis. major stages in the evolution and dispersal of M. These are > fascicularis (Table 39). discussed below. Stage I, 1 Ma Dispersal of M. fascicularis into Sunda Shelf area.

Stage II, ca. 160 Ka > Stage I, 1 Ma Dispersal and isolation of progenitors of strongly dif- ferentiated deep-water fringing-island popula-

tions. The earliest record of M'. fascicularis is at Trinil, M. umbrosa: Nicobar Islands f. east-, ca. 1 Ma (Table 33); the Trinil M. f. fusca, M. f. lasiae: Simeulue and Lasia fossils are part of the Dubois collection that also M. f. tua: Maratua includes Homo erectus and two species of leaf M. f. philippinensis: western, northern and eastern Philippines monkeys, Presbytis comata and Trachypithecus

auratus (Hooijer, 1962b, p. 5; Theunissen et al., Stage III, > 18 Ka Differentiation of progenitors of populations in the 1990, p. 41; Groves, 1993, p. 273). Assuming that and northern of the Indochinese Peninsula part M. fascicularis, H. erectus, and leaf monkeys dis- Isthmus of Kra. persed to Java from mainland Southeast Asia, they M. f. aurea: Indochinese Peninsula, Isthmus of probably reached Java by dry land during one of Kra, Mergui Archipelago the Pleistocene of advance M. f. fascicularis: Indochinese Peninsula, Isthmus Early periods glacial of Kra and low sea level (cf. J. de Vosetal., 1994, p. 131). earliest is in Stage IV, ca. 18 Ka The record of M. nemestrina also Dispersal and isolation of progenitors of weakly dif- east-central Java but is somewhat later (ca. 800 ferentiated popula- deep-water fringing-island Ka) than that of M. fascicularis (Table 33). Despite tions. this later date, zoogeographic evidence indicates M. f. fascicularis: Nias that M. nemestrina M. fasci- M. f fascicularis: southern Philippines probably preceded cularis in the Sunda Shelf area (Fooden, 1975, p. Stage V, < 18 Ka 70). Macaca is absent in Sulawesi and Isolation of progenitors of shallow-water fringing-is- fascicularis land populations and populations in Penida and Kepulauan Mentawai, faunistically distinctive is- Lombok (deep-water). lands off the Sunda Shelf that are inhabited by M. M. karimondjawae: Karimunjawa, Kemujan f nemestrina or its close relatives. Since M. fasci- M. f. atriceps: Khram Yai cularis is otherwise much more widely distributed M. f. condorensis: Con Son, Hon Ba thanM nemestrina and its relatives M. f fascicularis: other shallow-water fringing is- (Fooden, 1980, lands, excluding Bali p. 4), it appears that the intrinsic ability of M. M. Lombok f fascicularis: Bali, Penida, fascicularis to disperse is not inferior to that of M. Stage VI, ca. 4.5 Ka nemestrina. This suggests that the M. nemestrina and isolation of of Dispersal progenitors populations stock dispersed from the Sunda Shelf to Sulawesi in eastern Lesser Sunda Islands (deep-water). and Kepulauan Mentawai at some time prior to M. f fascicularis: Sumbawa-Timor 1 Ma, before M. fascicularis had arrived on the Sunda Shelf. During Late Pleistocene or Holocene, M. fas- 4. Transport by humans. Interisland transport cicularis may have been less abundant than M. of captive M. fascicularis by humans is reported nemestrina in the Malay Peninsula and Sumatra by Wallace (1869, p. 294) and presumably has a (see above, Fossils and Subfossils). If so, this would long, if undocumented, history in Southeast Asia. require explanation, because M. fascicularis is now Homo erectus reached Java by 1-2 Ma, and H. generally more abundant than M. nemestrina in sapiens reached Borneo and Australia by ca. 40 the Malay Peninsula, Sumatra, and elsewhere Ka (Bellwood, 1985, pp. 29, 89, 98; cf. Swisher et (Fooden, 1975, p. 61). al., 1994, p. 1119). Although much of the dispersal of hominids in the Sunda Shelf area probably oc- curred over dry land during glacial periods of low Stage II, ca. 160 Ka sea level, the dispersal of H. sapiens to Australia must have been over water, which implies that The most strongly differentiated populations of some means of over-water transport was available M. fascicularis inhabit four deep-water islands or

to humans in this area at least 40 Ka (cf. Sondaar island groups: (1) Nicobar Islands, northwest of etal., 1994, p. 1261). Sumatra; (2) P. Simeulue and P. Lasia, west of

FIELDIANA: ZOOLOGY Sumatra; (3) P. Maratua, east of Borneo; and (4) p. 802; Chaturvedi, 1980, p. 133; Musser & Carle- Philippines, excluding southern islands (Fig. 25). ton, 1993, pp. 651, 660); no information is avail- All of these populations are very dark (Appendix able concerning the date of first arrival of humans 3), and their deep-water island habitats are sym- in the Nicobars (Fuchs, 1973, p. 285; Gratton, metrically arrayed adjacent to the northwestern 1 992, p. 209). The poverty of the fauna, together and northeastern margins of the Sunda Shelf (cf. with the deep ocean trenches that surround this Kellogg, 1 944, p. 76). Assuming that dark dorsal group ofislands, strongly implies that these islands pelage is derived in M. fascicularis, the similar have never had a land connection to Sumatra or pelage saturation and geographic relationships of any other source of the rich mammalian fauna of these populations suggest that they may have had Southeast Asia. This in turn implies that M. fas- a similar evolutionary history. These populations cicularis reached the Nicobars by over-water dis- appear to have been isolated longer than popu- persal; the absence of this species in the Andaman lations of M. fascicularis inhabiting shallow-water Islands indicates that it dispersed to the Nicobars fringing islands, which were isolated by rising sea from Sumatra, the Malay Peninsula, or the Isth- level subsequent to the last glacial maximum, ca. mus of Kra— not from Burma. As noted above, 1 8 Ka. A working hypothesis is that the progen- the degree of differentiation of M. fascicularis in itors of these strongly differentiated populations the Nicobars suggests that this species may have reached their deep-water habitats during the pen- reached these islands during the penultimate gla- ultimate glacial maximum, ca. 1 60 Ka, when sea cial maximum, ca. 1 60 Ka; at this early date, dis- level was about 1 60 m lower than at present, and persal by natural rafting would be more likely than that these populations have been isolated at least dispersal by human introduction. since the subsequent interglacial, ca. 1 20 Ka. Fu- Following the arrival and isolation of M. fas- ture new evidence concerning the paleontology, cicularis in the Nicobars, dorsal pelage in the im- morphology, or molecular biology of these pop- migrant population apparently darkened, and skull ulations may provide a test of this hypothesis. size increased, resulting in differentiation of the Nicobar Islands— This group includes 19 is- subspecies M. f umbrosa. The present morpho- 2 lands, of which 1 1 are larger than about 2 km . logical uniformity of populations of this subspe-

M. fascicularis inhabits only Great Nicobar I., Lit- cies in the three islands that it inhabits implies tle Nicobar I., and Katchall I. (Kloss, 1903a, p. relatively recent genetic exchange among these 1 14)— the southernmost 3 of the 1 1 larger islands. populations; this exchange presumably occurred Forest cover in the Nicobars is restricted to these during the last glacial maximum, ca. 1 8 Ka, when three islands and Tillanchong I., one of the more Great Nicobar was joined to Little Nicobar and northern islands (Kloss, 1903a, p. 109). Monkeys, the gap between Little Nicobar and Katchall was including M. fascicularis, are absent from the An- reduced to narrow channels (< 10 km wide). daman Islands, a forested group of deep-water is- The lateral facial crest pattern is infrazygomatic lands that lies between the Nicobar Islands and in three of eight M. f umbrosa specimens exam- Burma (Kloss, [1928], p. 802; Chaturvedi, 1980, ined (one of three Great Nicobar specimens, two p. 134). of three Little Nicobar specimens, zero of two

Great Nicobar is separated from Sumatra by an Katchall specimens) (see Appendix 6). Three al- ocean trench that is 1 50 km wide and more than ternative interpretations would account for the rel- 200 m deep; for half of its width, this trench is atively high frequency of the infrazygomatic pat- more than 1,000 m deep. The channel between tern in M. f. umbrosa: (1) it may indicate that the Great Nicobar and Little Nicobar is 6 km wide progenitors of M. f umbrosa dispersed to the Ni- and less than 100 m deep, which indicates that cobars from the Malay Peninsula or Isthmus of these two islands were united during the last glacial Kra, in or near the geographic range of M.f aurea maximum. Narrow channels (< 10 km wide) be- (Figs. 9, 25); (2) it may indicate that the range of tween Little Nicobar and Katchall are more than M. f aurea extended to Sumatra during the pen- 1 20 m deep, which probably indicates that these ultimate glacial maximum, when progenitors of two islands were not united during the last glacial M. f umbrosa are hypothesized to have dispersed maximum. to the Nicobars; or (3) it may be the result of The nonvolant mammal fauna of the Nicobars independent local mutation. is limited to one species each of shrew, tree shrew, P. Simeulue and P. Lasia— These two islands and pig, four to six species of rats, Mfascicularis, are approximately 1 20 km west of northern Su- and humans (Miller, 1902b, p. 792; Kloss, [1928], matra (Fig. 25). Pulau Simeulue is relatively large

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 89 (ca. 100 x 25 km) and is about 20 km northwest central Borneo (Fig. 25; Kuenen, 1947, p. 5). It is of the much smaller P. Lasia (ca. 6x3 km). Al- separated from the Bornean shelf by a deep trench though M. fascicularis inhabits both of these is- (> 180 m) that is 17 km wide; the water gap is lands, it is absent in P. Babi (ca. 8x7 km), which narrowed by the presence of a stepping-stone is- 2 is only 3 km south of P. Lasia (Abbott in Miller, land, P. (area ca. 5 km ), which inter- 1903a, p. 479). venes in the middle of the trench. Deep trenches (> 200 m) separate P. Simeulue The only mammal collector known to have vis- and P. Lasia from Sumatra and from each other. ited P. Maratua is H. C. Raven, who worked there The deep trench between P. Simeulue and Sumatra in August 1912 and May 1913 (Deignan, [1960], is about 1 5 km wide, and that between P. Simeulue p. 267). In P. Maratua, Raven collected M. fas- and P. Lasia is about 4 km wide. The narrow cicularis, two species of rats, possibly a few other channel between P. Lasia and P. Babi probably is murine rodents, and one species of bat (Kellogg,

less than 1 00 m deep. 1944, p. 75; Musser & Carina, 1982, p. 6; M. D. The nonvolant mammal fauna of the Simeulue Carleton, usnm, letter 16 Dec. 1993). Judging from group, as reported by Kloss ([1928], p. 802; cf. the degree of differentiation of M. fascicularis in Sugardjito et al., 1989, p. 197; Musser & Carleton, P. Maratua, it may have dispersed to this island 1993, pp. 652, 659), is limited to one species each from Borneo during the penultimate glacial max- ofcivet and pig, two species of rats, M. fascicularis, imum, ca. 1 60 Ka; the dispersal probably was over and humans. This fauna, like that of the Nicobars water— presumably by natural rafting. In isola- (see above), presumably dispersed over water. tion, dorsal pelage in the founder population dark- Judging from the degree of differentiation of M. ened, resulting in differentiation of the subspecies fascicularis in P. Simeulue and P. Lasia, this spe- M.f. tua. In all four M.f. tua specimens examined, cies may have dispersed to these islands— presum- the lateral facial crest pattern is transzygomatic, ably by natural rafting— during the penultimate as in Bornean M. fascicularis. glacial maximum. Following isolation, the pelage Philippines, Excluding Southern Islands—

Genetic 1 1 the of the immigrant population darkened. As previously discussed (Fooden, 99 , p. 24), exchange between the populations in P. Simeulue progenitors of current populations of M. fascicu- and P. Lasia probably occurred during the last laris in western, northern, and eastern islands of glacial maximum. During subsequent postglacial the Philippine Archipelago may have dispersed to reisolation, the tail apparently shortened in the P. these islands during the penultimate glacial max- Simeulue population (Fig. 27; cf. above, M.f las- imum, ca. 1 60 Ka. Following isolation, dorsal pel- iae— Remarks), resulting in differentiation of M. age in the founder population became darker and fascicularisfusca and M.f. lasiae. Because P. Lasia also tended to become erythristic, resulting in dif- probably was joined to P. Babi during the last ferentiation of M. f philippinensis. glacial maximum, a dark population of M. fasci- cularis probably also inhabited P. Babi at that time but subsequently became locally extinct. Stage III, > 18 Ka The lateral facial crest pattern is infrazygomatic in 9 of 1 3 M. fascicularis fusca specimens exam- Indochinese Peninsula and Northern Part ined and in 1 of 2 M.f. lasiae specimens examined of Isthmus of Kra— Although zoogeographic ev- (Appendix 6). The geographic location of P. Si- idence, cited below, indicates that M. f aurea ex- meulue and P. Lasia strongly indicates that the isted as a differentiated subspecies in the western progenitors of M. fascicularisfusca and M.f. lasiae part of the Indochinese Peninsula and northern dispersed to these islands from Sumatra, not from part of the Isthmus of Kra prior to the last glacial

the Malay Peninsula or Isthmus of Kra (i.e., in or maximum, ca. 18 Ka, the evolutionary history of near the present range ofM.f. aurea; Fig. 25). This this subspecies and that of neighboring popula- implies either that the range of M. f aurea for- tions ofM.f. fascicularis remain enigmatic. Equiv- merly extended to Sumatra or that there has been ocal evidence concerning variation in M. fasci- independent mutation in these islands from the cularis in the Indochinese Peninsula and Isthmus transzygomatic crest pattern to the infrazygomatic of Kra is presented in the following paragraphs. crest pattern. The relationship between variation in M. fasci- P. Maratua— This U-shaped island (area ca. cularis and variation in M. mulatta—a. closely re- 2 25 km ) is a raised coral atoll 50 km east of north- lated species that replaces M. fascicularis in the

90 FIELDIANA: ZOOLOGY northern part of the Indochinese Peninsula (Food- pattern, which is infrazygomatic and which con- en, 1982, p. 576)— also is indicated. trasts with the transzygomatic pattern that pre- 1. Dorsal pelage color averages paler and less dominates in M. f fascicularis. Adjacent to the erythristic in the Indochinese Peninsula and geographic range of M. f aurea are two areas— northern part of the Isthmus of Kra than in the one in central and eastern parts of the Indochinese Malay Peninsula and Sumatra (Tables 1, 3). The Peninsula and the other in eastern and southern pattern of pelage color resemblances is indepen- parts of the Isthmus of Kra— inhabited by hetero- dent of local land connections; the Indochinese geneous populations that include some individuals Peninsula and Isthmus of Kra are continuous with with the infrazygomatic pattern, some with the the Malay Peninsula, which is separated by a water transzygomatic pattern, and some with the asym- gap from Sumatra. The latitude of pelage color metric pattern; occasionally, heterogeneous pop- transition is about 10°N, near the southern end of ulations have been sampled at a single locality. the Isthmus of Kra. The pale dorsal pelage color The southern part of the Indochinese Peninsula is in M. fascicularis north of 10°N generally ap- inhabited by a now-disjunct population of M. f proaches that in neighboring allopatric popula- fascicularis (homogeneous for the transzygomatic tions of M. mulatta; however, the posterior part pattern). of the dorsal surface is strongly erythristic in M. The lateral facial crest pattern in shallow-water mulatta— and not in Indochinese M. fascicularis. fringing-island populations of M. fascicularis 2. In the Indochinese Peninsula north of about matches that in adjacent mainland populations in 1 3°N, which is at the northern end of the Isthmus Southeast Asia; the crest is infrazygomatic in the of Kra, head and body length and skull length Mergui Archipelago— shallow-water islands west begin to decline northward in M. fascicularis (Fig. of the Isthmus of Kra, and the crest is transzy- 10; Fooden & Albrecht, 1993, p. 532). This de- gomatic in shallow-water islands east of the Isth- cline, which is anti-Bergmannian, is retrograde to mus of Kra and adjacent to the Malay Peninsula the general relationship between size and latitude (Fig. 9). This implies that M. f aurea had already in core-area M. fascicularis and also is retrograde differentiated, and was approximately restricted to to the relationship between size and latitude in M. its present geographic range, prior to the last glacial fascicularis in the Philippines, at approximately maximum (ca. 1 8 Ka), when M. fascicularis pre- the same latitude as the Indochinese Peninsula. sumably dispersed from mainland Southeast Asia The northward decline of size in Indochinese M. to adjacent shallow-water islands. fascicularis tends to bring the size of this species Pelage color, head and body length, tail length, near to that of neighboring populations of M. mu- and skull length in populations of M. f aurea are latta (Figs. 29, 30). generally similar to those in populations of M. f 3. Tail length declines rapidly in the Indo- fascicularis that inhabit similar latitudes in the

Chinese Peninsula north of about 1 3°N (Fig. 1 3). Indochinese Peninsula and Isthmus of Kra. The This is an acceleration of the general decline of lateral facial crest pattern in M. f aurea is similar tail length in core-area M. fascicularis north of the to that in M. mulatta, where the infrazygomatic equator (Allen's rule); there is no comparable ac- pattern is predominant (Stewart, 1933, p. 30). celerated northward decline of tail length in Phil- 5. The frequencies of mtDNA types, blood- ippine M. fascicularis. Parallel to the trend noted protein alleles, and resistance to Plasmodium above for head and body length and skull length, knowlesi malaria in populations of M. fascicularis the accelerated northward decline of tail length in north of the Isthmus of Kra differ markedly from Indochinese M. fascicularis tends to bring the tail those in populations south of the Isthmus of Kra length of this species near to that of neighboring (Tables 17, 19; Fooden, 1994, p. 585). The fre- populations of M. mulatta (Fig. 31; cf. Fooden, quency off. knowlesi resistance in populations of 1971, p. 29). M. fascicularis north of the Isthmus of Kra is sim- 4. The subspecies M. f aurea inhabits a re- ilar to that in M. mulatta. stricted area bordering the , mostly Based on this evidence, the following conclu- west of the mountain ranges that form the border sions may be drawn. Although suggestive, these between Burma and Thailand, along the western conclusions are not adequate to explain the evo- side of the Indochinese Peninsula and Isthmus of lutionary history of M. fascicularis in the Indo- Kra, south to about 10°N (Figs. 9, 25). This sub- Chinese Peninsula and northern part of the Isth- species is distinguished by its lateral facial crest mus of Kra.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 91 650 -r-

600

550

500

4-50

Males

L00 J

£ 550 -r

IV V V 500 jV -7 IV V

4-50-

4-00 * v •* ; v Females

550 ! 10° 12 135

130 —

125-

120-

115

110-

o 550-

500-

450-

400-

550-

300-

250-

?oo- 3. Populations of M. f. aurea and M. f. fasci- ulations that include pale, intermediate, and dark cularis in the northern part of the Isthmus of Kra individuals may be the result of interbreeding be- and Indochinese Peninsula tend to be transitional tween early-arriving M. f philippinensis and late- between M. fascicularis in the Sunda Shelf area arriving M. f fascicularis. and M. mulatta, which replaces M. fascicularis in the northern part of the Indochinese Peninsula.

Stage V, < 18 Ka

Stage IV, ca. 18 Ka Shallow-Water Fringing Islands, Ex- cluding P. Bali— During the last glacial maxi- P. Nias— Although P. Nias is a deep-water is- mum, ca. 1 8 Ka, sea level was approximately 1 20 land, its population of M. fascicularis is not strong- m lower than at present, and shallow-water fring- ly differentiated (see above, Subspecific Taxono- ing islands— together with Sumatra, Borneo, and my), which suggests relatively recent isolation of Java— were part of a single large landmass that this population. The strait between P. Nias (area extended from the Indochinese Peninsula to P. 2 ca. 3,500 km ) and the west coast of northern Su- Bali (Fig. 3; Heaney, 1991a, p. 55). This permitted matra is about 90 km wide, but the deep channel dispersal, or redispersal, of M. fascicularis over (120-180 m) in this strait is only about 20 km dry land to what are now shallow-water islands; wide. The nonvolant mammal fauna reported for as a result, populations of M. fascicularis in all of P. Nias by Kloss ([1928], p. 802; cf. Musser & these present-day islands were genetically contin- Califia, 1982, pp. 11, 18; Musser &Carleton, 1993, uous. Subsequently, between 1 8 Ka and 5 Ka, sea pp. 614, 633, 652) is limited to M. fascicularis, level rose to its present level, and present-day shal- one species each oftree shrew, binturong, pig, trag- low-water insular populations of M. fascicularis ulid, and pangolin, two species of deer, and five became isolated. species of murine rodents. Following isolation, populations of M. fascicu- Macaca fascicularis may have dispersed to P. laris in many shallow-water fringing islands ap- Nias from Sumatra during the last glacial maxi- parently have tended to become slightly darker, mum, ca. 1 8 Ka, when the intervening water gap more erythristic, and/or smaller than their ances- was reduced to about 20 km. During subsequent tral stock, assuming that this stock is represented isolation, the proximodorsal surface of the tail in by core-area populations (Table 9; Appendixes 3, the P. Nias population apparently became darker 4; Fooden & Albrecht, 1993, p. 533). The darkest and the tail apparently became shorter. In all 12 shallow-water fringing-island population, M. f specimens examined of M. fascicularis from P. karimondjawae in P. Karimunjawa and presum- Nias, the lateral facial crest pattern is transzygo- ably also in P. Kemujan, may have become iso- matic, as in Sumatran M. fascicularis. lated somewhat earlier than most other shallow- Philippines, Southern Islands— As previous- water fringing-island populations (Appendix 3). ly noted (Fooden, 1991, p. 6), dorsal pelage color The distinctive dark crown patches of M. f atri- in M. fascicularis in southern islands of the Phil- ceps in Ko Khram Yai and M. f condorensis in ippine Archipelago is paler than in M. f philip- Con Son and Hon Ba presumably also developed pinensis, which inhabits western, northern, and in isolation following elevation of sea level since eastern islands in this archipelago. The pale south- the last glacial maximum. During the same inter- ern populations, which are allocated to the nom- val, dorsal pelage erythrism in M.f fascicularis in inotypical subspecies M. f fascicularis, are indis- Borneo apparently differentiated somewhat from tinguishable in pelage color from populations in that in the same subspecies in Sumatra and the Borneo. This strongly suggests that progenitors of Malay Peninsula (Tables 3, 4). the southern populations dispersed from Borneo Despite postglacial isolation, many shallow-wa- to the Philippines more recently— possibly during ter island populations of M. fascicularis appear to the last glacial maximum— than progenitors ofM. have retained ancestral characters. Size is similar f philippinensis (see above). Present distributions at similar latitudes in currently isolated popula- indicate that the relatively recent dispersal of M. tions in the Malay Peninsula, Sumatra, and Bor- f fascicularis to the southern Philippines occurred neo (Fig. 10; Fooden & Albrecht, 1993, p. 533). via the Sulu Archipelago (Fooden, 1991, p. 28). Dorsal pelage erythrism and tail length are more In Negros and Mindanao, a zone of mixed pop- similar in populations in the Malay Peninsula and

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 95 Sumatra, which are now isolated, than in popu- The characters that distinguish M. fascicularis lations in the Malay Peninsula and the Indo- and T. auratus in P. Bali from their respective chinese Peninsula, which are connected by dry land conspecifics in Java link populations of these spe- (Tables 3, 10). The infrazygomatic lateral facial cies in P. Bali to those in P. Lombok. This strongly crest pattern apparently has persisted in popula- suggests that, subsequent to their differentiation tions in the Mergui Archipelago despite the post- in P. Bali, populations of both species dispersed— glacial isolation of these islands from the Isthmus over water— from P. Bali to P. Lombok, as pre- ofKra(Fig. 9). viously proposed by Kawamoto and Suryobroto Lesser Sunda Islands: P. Bali, Nusa Penida, (1985, p. 39); the dispersal of M. fascicularis to and P. Lombok— Pulau Bali is the only shallow- Nusa Penida, located between P. Bali and P. Lom- water fringing island in the Lesser Sunda group bok, presumably occurred at the same time.

(Fig. 3; Kitchener et al., 1990, p. 1 1 1). The strait Whether the dispersal of M. fascicularis and T. between Java and P. Bali is approximately 3 km auratus from P. Bali to P. Lombok was by natural

wide and has a maximum depth of about 65 m. rafting or by human transport is uncertain (cf. Ev-

The strait between P. Bali and Nusa Penida is erett in Hartert, 1896, p. 593; Kitchener et al., about 10 km wide and has a depth of more than 1990, p. 112). 1 20 m for about half of its width. The strait be- tween Nusa Penida and P. Lombok is approxi- mately 23 km wide and has a depth of more than Stage VI, ca. 4.5 Ka 1 20 m for almost all of its width. All three of these islands are inhabited by M. Lesser Sunda Islands: P. Sumbawa-P. Tim- fascicularis (Fig. 2C). The only other nonhuman or—Macaca fascicularis is known to inhabit 13 primate known to inhabit the Lesser Sundas is one of the deep-water Lesser Sunda Islands east of P. species of leaf monkey, Trachypithecus auratus, Lombok (listed in west-east order): P. Sumbawa, the range ofwhich includes P. Bali and P. Lombok. P. Moyo, P. Sumba, P. Rinca, Nusa Kode, P. Man- In these two islands, T. auratus is represented by giatan, P. Seraya Besar, P. Flores, P. Adonara, P. an endemic subspecies, T. a. kohlbruggei (P. H. Solor, P. Semau, P. Kambing, and P. Timor (Fig. Napier, 1985, p. 56; Weitzel & Groves, 1985, p. 2C). This species reportedly is absent in P. Ko- 402; Groves, 1993, p. 273). modo and P. Padar (between P. Sumbawa and P. During the last glacial maximum, ca. 1 8 Ka, P. Rinca, north of P. Sumba), and in many nearby Bali was joined to Java, but Nusa Penida and P. islets, and also is absent in P. Lomblen, P. Pantar,

Lombok were separate islands (Kitchener et al., and P. Alor (east of P. Adonara and P. Solor, north 1990, p. lll;Heaney, 1991a, p. 55). At that time, of P. Timor) (for documentation, see Gazetteer, populations of M. fascicularis and T. auratus in Appendix 2). No other nonhuman primate inhab- P. Bali presumably were genetically continuous its any of the Lesser Sunda Islands east of P. Lom- with those in Java. It is unclear whether or not the bok. last glacial maximum was the time of initial dis- None of these islands were connected to P. Bali persal of M. fascicularis and T. auratus from Java during the last glacial maximum, although some to P. Bali; these species may have dispersed from of these islands were interconnected at that time Java during a previous glacial cycle, and their in- to form four larger islands (Lombok-Sumbawa- sular populations may merely have been reunited Moyo, Sumba, Komodo-Padar-Rinca-Kode- during the last glacial maximum. Mangiatan-Seraya Besar-Flores-Alor-Solor, and In either event, sometime before 5 Ka, rising Semau-Kambing-Timor) (Heaney, 1991a, p. 55). sea level separated P. Bali from Java and, conse- The isolation of these deep-water islands is reflect- quently, separated populations of M. fascicularis ed in their relatively poor nonvolant mammal fau- T. and auratus in P. Bali from those in Java. Fol- nas (Laurie & Hill, 1954, pp. 13 ff.). In P. Flores 2 lowing this separation, M. fascicularis in P. Bali (area ca. 20,000 km ), for example, Musser (1981, apparently underwent reduction in body size (Figs. p. 134) listed 16 species of nonvolant mammals

1 5, 22; Appendix 7) and also underwent a marked (shrews, 3; M. fascicularis; civet, 1; pigs, 2; deer, shift in allele frequencies at blood-protein loci 1; porcupine, 1; rats, 5; mice, 2), of which some HbA-II, Pi, and Tf (Table 19). Concurrently, T. probably were introduced by humans. This con- auratus in P. Bali apparently became subspecifi- trasts with 38 species of nonvolant mammals re- 2 cally distinct from T. auratus in Java. corded for P. Bangka (area ca. 1 2,000 km ), a shal-

96 FIELDIANA: ZOOLOGY low-water island on the Sunda Shelf (Heaney, 1 984, Auffenberg (1981, p. 242) suggests that the ab- P. 12). sence of M. fascicularis in P. Komodo (area 393 2 2 Four items of partly contradictory evidence seem km ) and P. Padar (13 km ) may be attributable most relevant to attempting an interpretation of to an insufficient year-round supply of fruit in these the history of M. fascicularis in deep-water Lesser islands. This is puzzling, because small islands im- Sunda Islands east of P. Lombok: (1) Dorsal pelage mediately east ofP. Komodo and P. Padar support 2 color in these islands is essentially similar to dorsal populations of M. fascicularis (P. Rinca, 278 km ; in P. P. Nusa P. 9.6 2 P. pelage color Java, Bali, and Lombok (Ap- Kode [= Oewada Sami], km ; Ser- size in Besar P. 2 pendixes 3, 4). (2) External and cranial these aya [= Seraja], 3.6 km ; P. Mangiatan [= 2 islands is similar to size in P. Bali and P. Lombok P. Mengjatan], 1 km ). The insular distribution of (Figs. 15, 16, 22, 23); size is greater in Java. (3) M. fascicularis in this area may be affected in some Blood-protein allele frequencies at loci HbA-II, Pi, way by the distribution of the Komodo giant mon- and Tf in these islands are similar to allele fre- itor, a known predator ofthese monkeys (see above, quencies in Java (Table 1 9); frequencies of these Natural History) that is restricted to the following alleles are divergent in P. Bali and P. Lombok. (4) seven islands (listed in west-east order): P. Ko- in 2 Subfossils of M. fascicularis P. Timor cave de- modo, Nusa Mbarapu (0.6 km , occasionally vis- posits appear relatively late (ca. 4.5 Ka), concur- ited by giant monitors), P. Padar, P. Rinca, Nusa rently with the appearance of remains ofdomestic Kode, Gili Motang (= Gili Mota), and western P. animals and pottery (see above, Fossils and Flores (Auffenberg, 1981, p. 40). M. fascicularis Subfossils). coexists with the giant monitor only in three of The evidence of dorsal pelage color and Timor the more easterly of these islands (P. Rinca, Nusa cave deposits suggests relatively recent dispersal Kode, and P. Flores). of M. fascicularis to deep-water Lesser Sunda Is- The relatively recent dispersal ofM. fascicularis lands east of P. Lombok. The blood-protein evi- to deep-water Lesser Sunda Islands, indicated by dence indicates that the differentiated populations evidence cited above, is somewhat surprising. This of M. fascicularis in P. Bali and P. Lombok prob- species has been present in east-central Java since ably were not directly involved in dispersal of this ca. 1 Ma (see above), and, during the maximum species to islands east of P. Lombok. Assuming Pleistocene glaciation, the widest water gap be- that the differentiated populations in P. Bali and tween Java and Sumbawa may have been only 400 P. Lombok inhabited those islands before M. fas- m or less (Kitchener et al., 1 990, p. 1 1 0). However, cicularis dispersed to islands east of P. Lombok as indicated by Heaney (1991a, p. 57), during gla- (see above), the implication is that populations of cial maxima, rainfall in the Lesser Sundas was M. fascicularis east of P. Lombok were derived much lower than at present. Perhaps forest cover from a source in Java, or elsewhere west of P. Bali. was inadequate to sustain M. fascicularis in these This geographic incongruity, together with the ar- islands during glacial maxima, when low sea level cheological evidence of the Timor cave deposits, and narrow water gaps might otherwise have fa- strongly supports the hypothesis that the popu- cilitated the eastward dispersal of this species. lations of M. fascicularis in the Lesser Sundas east of P. Lombok are the result of human introduc- tion, beginning ca. 4.5 Ka; this hypothesis has been previously proposed by Musser (1981, p. 133), Acknowledgments Kawamoto and Suryobroto (1985, p. 39), and Glover (1986, p. 159). The generous cooperation of curators and staff Multiple introductions would be required to es- members of institutions in which specimens ex- tablish populations of M. fascicularis in all of the amined are preserved (see Introduction) has been islands east of P. Lombok that it now inhabits. If indispensable to the conduct of this research and Java was the source of these introductions, exter- is gratefully acknowledged. For valuable supple- nal and cranial size became convergently reduced mentary advice and assistance, I also thank the in these islands. This is inconvenient to the hy- following: H. Abdulali (bnhs), N. Aggimarangsee pothesis, but not impossible, considering the fre- (Chiang Mai University, Thailand), M. Aimi (pri), quency of insular dwarfing in M. fascicularis (see G. H. Albrecht (University ofSouthern California, above, External Measurements and Proportions; Los Angeles), W. Y. Brockelman (Mahidol Uni- Cranial Characters). versity, Bangkok), M. D. Carleton (usnm), S.

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FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 119 Appendix 1: Specimens Examined Sintang— ansp 14037, 14042, 14043; Tangarveng (Total 2,049) I. -usnm 1 96804; Telang— usnm 521838; Tibang, Mt.— mcz 22277; no locality— ansp 14041, bm(nh) Macaca fascicularis fascicularis, 1856.9.3.6, zsbs Coll. Nos. 272, 738, 788, 1227, Total 1,550 1354, 1373, 1527;5w/a«(3),south-usNM 144419; no locality— zrc 4-098, 4-099; Durian (2), no lo- Skins and Skulls, 739 cality— mzb 241, 242; Flores (7), Mbura— mzb 2384-2387; Rana Mese-NMS 1 6630; Sano, Wai-

CAMBODIA, 2. Mainland'(2), Siemreab— mnhn mzb 2388; no locality— bm(nh) 1864.4.12.1; Gal- 1929.460; no locality— mnhn 1961.61 1. ang (4), no locality— zrc 4-100-4-103; Java (84), INDONESIA, 456. Bali (21), Banjoe Wetan- Bandung, near— usnm 521839; Bantargebang— mzb 6521-6523; Batoe-meringgit— mzb 2001; mcz 12757, usnm 156291, 156292, 156295; Ban- Bratan, Gunung, amnh 107555-107557; Desa yuwangi— rmnh 959; Batoeraden— rmnh Coll. No. Poetjang— rmnh Coll. No. E85 (external measure- 89C; Bogor— mzb 1884; Camara— mzb 6482; Can- ments from Sody, 1933, p. 94); Gilimanuk— amnh diroto— rmnh Coll. No. 107B; Cihara— usnm 107561-107568; Gitgit-NMS 16632; Jembrana- 156456, 156458; Cikujang-MZB 6483, 6485; Cil- rmnh Coll. No. El 39 (external measurements from acap-BM(NH) 1909,1.5.27, 1909.1.5.28; Cire- Sody, 1933, p. 94); Sendang— rmnh Coll. Nos. bon—amnh 101891, 102016; Ciremay, Gunung- E34, E64, E74 (external measurements for all three amnh 102015, 102017-102022; Ciwangi-BM(NH) from Sody, 1933, p. 94); Bangka (4), Pamuja, Tan- 1909.1.5.29, 1909.1.5.30; Depok-Mcz 12755 (ex- jung— usnm 124863; Rengsam, Tanjung— usnm ternal measurements from usnm field cat.), 1 2756; 1247 10; no locality-RMNH Coll. Nos. Bk52, Bk53; Gedangan— rmnh Coll. Nos. 74, V9, V10; Indra- Batam (4) "Tanjong Sauh"-BM(NH) 1909.4.1.29, mayu— mzb 2961, 2962; Jasinga— mzb 2052, 3189; bnhs 5080; Tanjong Turut-BM(NH) 1909.4.1.27, Java, West— rmnh 2b, 3c, 4h; Kaligoea— mzb 1909.4.1.28; Bawean (8), no locality-MZB 1841- 574, 575; Kalipucang-BM(NH) 1909.1.5.19, 1846, usnm 151829, 151830; Belitung (2), Batu, 1909.1.5.32, 1909.1.5.33; Linggajati-AMNH Tanjung-USNM 124969, 124970; Bengkalis (1), 101811; Majalengka-AMNH 101808-101810; Kapos Tinggi— usnm 143582; Benua (2), no lo- Pangandaran— bm(nh) 1909.1.5.24; Pangonan— cality-usNM 101638, 101639; Bintan (8), north rmnh Coll. No. M40; Pasir Carolina— mzb 6484; coast— usnm 115676; Pasir Panjang— bm(nh) Pelabuhanratu, Teluk— usnm 156457, zrc 4-134; 1909.4.1.26, usnm 115677; Sungei Biru-BM(NH) Salak, Gunung— rmnh Coll. No. A 147; Saron- 1 909.4. 1 .23-1 909.4. 1 .25; no locality-ZRC 4-093, gen— mcz 12759; Singkil, Gunung— rmnh 5919o, 4-094; Borneo: Kalimantan (93), Ambawang, Sun- 5919p, 5919q; Tamadjaija— mcz 12758; Taman- gai — usnm 142225; Badang— amnh 103730, sari— bm(nh) 1954.51, zrc 4-133; Tasikmalaya— 103731, 106024-106026, mzb 6513; Berau, Sun- bm(nh) 1909.1.5.31, mzb 1604-1609; Tiloe, gai— usnm 196817-196822; Birang, Sungei— usnm Goenoeng— rmnh 5919n; Tjeringin— rmnh Coll. 196825 (includes skinned head in fluid); Buntok— Nos. 13f, 25f, 26f; Wonokojo— rmnh Coll. Nos. bm(nh) 1910.4.5.23; Goson Djerong — usnm Won3, Won4, Won5; no locality— nms 16628, 196814; Hantakan — usnm 521837; Karangin- rmnh 2505a, 2505b, 2527, 3094, 3095, 3097, zmb tan— bm(nh) 1910.4.5.22; Karangmumus, Sun- 113, zmuz 11627 (skin)/11630 (skeleton); Kan- — gai— usnm 196816; Karangtigau, Tanjung— usnm gean (4), no locality — bm(nh) 1 9 10.4.6. 1 196824; Kembangjanggut— mzb 8130; Liang Koe- 1910.4.6.4; Karimata (2), Pai, Teluk — usnm boeng— rmnh 160, 178; Loa Bambam— usnm 125101, 125102; Karimun (7), Mensuda Bay- 196815; Long Peleben-AMNH 106027, mzb 6514; usnm 122849; Monos-BM(NH) 1909.4.1.34, Mahakam, Sungai — usnm 196813; Merah — 1909.4.1.35; Pemeral-BM(NH) 1909.4.1.30- — mzb 1164; Muaratewe bm(nh) 1910.4.5.19, 1909.4.1.33; Kundur (2), Selatbliat-BM(NH) 1910.4.5.20; Parit-AMNH 103644-103658, 1909.4.1.36, 1909.4.1.37; Lagong (1), no locali- 103660-103663, mzb 6515-6517, zmb 48005, ty— usnm 104853; Laut (1), no locality— usnm 48006, 92191; Pelawan, Sungai-USNM 198301; 104854; Lingga (2), no locality— usnm 101602, Perbuah-AMNH 107091-107096, 107098, 101603 (external measurements from Miller,

107099, mzb 6519, 6520; Putussibau-ANSP 14038, 1906c, p. 276); Lombok (3), Pussuk forest— mzb 14039, 14044; Riam-amnh 106283-106285, mzb 6528; Sewela-NMS 16634, zmb 92306; Mapur(l), 6518; Roema Manoeal— rmnh 23, 24; Sembak- Mentigi— zrc 4-097; Matasiri (1), east coast— ung, Sungai— mzb 576, 6512; Semitau— rmnh 6; usnm 151831; Mursala (2), no locality— usnm

120 FIELDIANA: ZOOLOGY 114559, 114560; Natuna Besar (2), Binjai, Sun- 114408-114410; Uwi (1), no locality-uSNM gai—zrc 4-139, 4-140; Nguwal (6), no locality— 101666. zrc 4-104-4-109; Nias (1 1), Hilisimaetano— ansp MALA YSIA, 1 63. MALA YSIA: SABAH, Bang- 20404-20406; Lafau-USNM 141372; Samasa- gi (5), Banggi, Pulau, south— zrc 4-116-4-118; ma—usnm 141371; Siaba, Teluk— usnm 121868- Karakit-FMNH 140939; Sabor-FMNH 140938; 121871; Soliga-Mcz 36029, 36030; Pelapis Ten- Borneo (74), Abai-Mcz 35600, 35606, 35611- gah (1), no locality— mzb 2914; Penida (3), no lo- 35613, 35629, 35641, 35642, 35693, 35694, cality— amnh 107558-107560; Serasan (2), no lo- 35700, 35701, 35722, 35724, 35726, 35727, cality— usnm 104852, zrc 4-144; Serutu (1), no 35729, 35731, 35732, 35744, 35761; Betotan,ZRC locality— mzb 29 1 5; Siantan (3), no locality —usnm 4-115; Bongkabong-Mcz 37409, 37411, 37412; 101711, zrc 4-136, 4-137; Subi-kecil (3), no lo- Darvel Bay—zmb A86.10; Garau—mcz 37349, cality—zrc 4-141-4-143; Sugi (1), no locality— 37351, 37414, 37418; Kampong Bundu Tuhan- usnm 115675; Sumatra (123), Batangkuis— zsbs usnm 292555-292558; Kampong Kiau — mcz 4-1 4-121 Coll. No. 9; Bungur-Buikt-ZMB A25.09; Deli, 37348, 37352, zrc 20, ; Keningau-AMNH Sungai— ansp 20217, 20219; Goenoengsetan- 188343; Kenokok zrc 4-122; Kiaulan— mcz Meloewak— mzb 6481; Indragiri, Sungai— usnm 37350, 37353; Kinabatangan, Sungai — usnm 113169; Kalianda- amnh 1 02904-1 02906; Kam- 19193; Kretam Besar, Sungai— fmnh 68700; Kre- bang— zmb A25.09; "Kg. Baru"— zsbs Coll. Nos. tam Kechil, Sungai— fmnh 68699; Lahad Datu— 156, 176, 180; Kotabumi-ZRC 4-124; Lesten- zmb A89.04; Papar— usnm 317190; Ranau— mcz mzb 6480; Lubuklinggau— amnh 102211; Me- 37346, usnm 3 1719 1-3 17 195; Rayoh-ZRC 4-1 19; dan-ANSP 20221, zsbs Coll. Nos. 1/10, 5, 6, 15, Rugading— mcz 37357; Sapagaya Forest Re- 16, 21, 22, 26, 30-32, 35, 36, 42, 45-48, 51-53, serve— fmnh 68701, 68702; Sibuga Besar, Sun- 68 (Bruegel), 68 (Widnmann), 76, 79, 82, 89, 91, gai— fmnh 33545; Talibang— mcz 37354, 37406, 92, 101, 105, 134, W2,Wll;Medan, forest near- 37413; Tawau—amnh 31288; Tempasuk, Sun- zsbs Coll. No. 11; Muaradua— amnh 102763- gai— mcz 37417; Tenompok— mcz 37415, 37416; 102770; Padang-RMNH 518, 981, 1051; Padang Tinonkok— mcz 37347; Tuaran-Kampong Tengh- highlands— rmnh e2; Pajo— bm(nh) 1879.6.28.4; ilan Road-Mcz 37355, 37356, 37405, 37408; no Pematangsiantar— mcz 35937, 35938; Sandaran locality-uSNM 344989-344992; Sebatik (2), no Agong— zrc 4-110; Sanggul, Bukit— amnh locality— usnm 175896 (skin and skull)/amnh 106565-106569, mzb 6477-6479; Serdang-ZSBS 30620 (postcranial skeleton), usnm 175897. MA- Coll. No. 134;Siulakderas-BM(NH) 1919.11.12.8; LAYSIA: SARAWAK, Borneo (15), Baram, Ba- Sukadana—amnh 102907-102910; Sumatra, [east- tang-BM(NH) 1894.6.12.2, USNM 83945; Belaga central]— zmb A 14.08/37; Sumatra, east coast— or Kalulong, Bukit-BM(NH) 1951.66, 1951.68; zsbs Coll. Nos. 105, 135, 156, 166-168, 174; Su- Bukar, Sungai— zrc 4-113; Entawa, Tanjong— matra, west coast— bm(nh) 1882.7.24.1; Tanjung- bm(nh) 1955.710; Kuching-SMK 092/3, 092/8; morawa— zsbs Coll. Nos. 7 1 [b], 72, 1 37; Tapanuli, Kuching, probably— zmb 4939; Long Ekang— Teluk— usnm 114505, 114506; Tapung-kanan, fmnh 88582; Paku, Saribas-BM(NH) 1955.711, Sungai— zmb A14.08; Tarusan, Teluk— usnm 1955.712; Pelandok, Sungai-ZRC 4-112; Punang, 141145; Telukbetung— fmnh 14805; Telukpan- Sungai— zrc 4-114; Selikan, Bukit— usnm 83944. ji— zrc 4-111; no locality— NHMBa 3347, 3348, MALA YSIA: WEST MALA YSIA, mainland (29), rmnh 1479, usnm 271018, zsbs Coll. Nos. 28, 35, Benom, Gunong— zrc 4-065, 4-066; Changkat 36, 38, 58, 64, 67, 93, 107, 112, 135, 157, 184, Mentri— bm(nh) 1955.1510, zrc 4-064; Cherak- 1 85, 203-205, 2 1 2, 232; Sumba (8), Langgaliroe- ah, Bukit-BM(NH) 1955.1516, zrc 4-069; Dusun mzb 6526, 6527, zmb 92075; Mao Marroe— mzb Tua— zrc 4-068; Gaik Liew Estate — bm(nh) 867-869; Payeti-Kambaniru— mzb 865, 866; 1955.1515; Hantu, Tanjong -zrc 4-063; Kelang — Sumbawa (8), Batudulang— mzb 6529, 6530, zmb Road-BM(NH) 1971.2611; Melaka bm(nh) 92307, 92308; Dompu-NMS 1025; Kambing, Pu- 1875.3.29.1, rmnh 41g; Nanas, Bukit— bm(nh) lau— mzb 6531; Ntori— zmb 92309; Oo vicinity— 1939.180, 1939.181, 1955.1512; Nyalas-ZRC nms 1026; Tanahbala (2), no locality— usnm 4-078 (external measurements from Weitzel et al.,

121802, 121803; Timor (9), Kuatnana-ZSBS 1911/ 1988, p. 107); Pong-BM(NH) 1934.7.18.1; Port 2108; Lelogama-ZSBS 1911/2102, 1911/2104, Dickson -bm(nh) 1939.229 (skin)/RCS(OM) A85.4 1911/2105; Mutis, Gunung— mzb 6524, zmb (skull), zrc 4-073-4-075; Tanjong Panjair— bm(nh)

1 3 1 1 92 , 92 32; Nikiniki-MZB 6525; Timau, Fatu- 1955.1514, zrc 4-067; Tanjong Tuan, Keramat— zsbs 1911/2101; Tuangku (3), north coast— usnm zrc 4-072; Telom, Sungai-BM(NH) 1934.7.18.4,

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 121 1934.7.18.5; Ulu Ijok-BM(NH) 1934.7.18.2, Kut (8), no locality-BM(NH) 1915.11.4.8- 1934.7. 18.3; no locality-usNM 301 758; Acheh(l), 1 9 1 5. 1 1 .4. 1 0, 1 976. 1 826 (skin)/zRC 4-040 (skull), no locality — bm(nh) 1955. 1 5 IS; A ur (2), no local- usnm 201552, 201553 (external measurements ity- bm(nh) 1955.1520, zrc 4-080; Burau (1), no from Kloss, 1916a, p. 67), 254741, zrc 4-041; locality— bm(nh) 1955.1519; Langkawi (1), no lo- Phangan (2), southwest— zrc 4-044, 4-045; Phi cality— bm(nh) 1 909. 11.1.1; Pemanggil (2), no lo- Phi Don (4), no locality —zrc 4-046-4-049; Samui cality-BM(NH) 1955.1508, usnm 1 12500; Pinang (2),west-BM(NH) 1955. 1522, zrc 4-050; Talibong flj(8), Pantai Krachut-ZRC 4-062; Telok Bahang (2), no locality— usnm 83272, zrc 4-051; Tarutao bm(nh) 1955.1521, zrc 4-056-zrc 4-061; Pinang (6), Sungei Adang-BM(NH) 1909.11.1.2,

[2] (2), no locality -bm(nh) 1949.426, 1955.1523; 1909.1 1.1.3; Telok Wau-BM(NH) 1955. 1509, zrc Pintu Gedong (3), no locality— bm(nh) 1955.1517, 4-053; no locality-usNM 123990, 123991. zrc 4-070, 4-071; Redang (2), no locality— zrc VIETNAM, 8. Mainland (6), Cochin China- 4-054, 4-055; Tinggi (5), no locality— bm(nh) bm(nh) 1881.6.30.2, mnhn 1882.2; Ho Chi Minh 1909.4.1.21, 1909.4.1.22, 1955.1527, zrc 4-131, City— fmnh 33505; Ho Chi Minh City, Zoological 4-132; Tioman (11), Juara, Telok — bm(nh) Garden-MNHN 1929.459; Sontra Peak, 3.9 km 1908.1.25.1, 1908.2.25.1, 1955.1524, 1955.1525, W, 0.3 km S-usnm 356968; Xa Trang Bom-ZRC zrc 4- 1 25-4- 1 27; Sedagong, zrc 4- 1 29, 4- 1 30; no 4-020 (external measurements from Weitzel et al., locality-BM(NH) 1955.1526 (skin)/zRC 4-128 1988, p. \ 14); Phu Quoc(2)-bm(nh) 1928.7.1.10, (skull), usnm 101744. mnhn 1929.455.

PHILIPPINES, 43. Basilan (3), Basilan I., east end— usnm 144666; Camp No. 4 and Camp No. Skins only, 285 5, between— usnm 144665; Isabela— usnm 125326; - Cagayan Sulu (1), no locality— usnm 125325; CAMBODIA, 1. Mainland (1), no locality Mindanao (39), Bucong— fmnh 65440-65444; RMNH 38. Canibongan— fmnh 67737-67739; Gubat— fmnh CAMBODIA OR VIETNAM, 2. Mainland (2),

67725, 67729-6773 1; Libu-FMNH 67727, 67728; "Camboja or Cochinchine"— bm(nh) 1 878.6. 1 7.4, Mamara— fmnh 67717, 67718; Pulunbato, Mt.— 1878.6.17.5. bm(nh) 1880.1 1.24.1; San Ramon-FMNH 33507- INDONESIA, 241. Bali (1), Bratan, Danau- 33511; Sigayan-FMNH 67722, 67732, 67740- bm(nh) 1913.3.6.7; Bangka (1), Simpang— zsbs 67742; Situbo-FMNH 67719-67721, 67723, Coll. No. 25; Borneo: Kalimantan (82), Borneo, 67724; Tacuta — fmnh 67733-67735; Tampa- southeast — rmnh 35v (skull inside); Borneo, Ian— fmnh 67736; Zamboanga— usnm 144667, southwest— rmnh 32s, 33t, 34u (skulls inside); 144668, 144698, Kampong Hadjak— irsn 9796; Karangtigau, Tan- — SINGAPORE, 1. Singapore (7), Botanical Gar- jung— usnm 196823; Muaratewe bm(nh) den-ZRC 4-082, 4-084; Changi-BM(NH) 1910.4.5.21; Parit-AMNH 103659 (skull not lo- 1909.4.1.20; Punggol-BM(NH) 1955.1513, zrc cated, cranial measurement from G. H. Albrecht, 4-086; no locality— zmb 5444, zrc 4-090. pers. comm., Oct. 1991), zmb 92190; Pelaihari— THAILAND, 60. Mainland (32), Aranyaprath- rmnh 36w (skull inside); Perbuah— amnh 107097;

et— mcz 35931; Ban Huai Maenam Noi— fmnh Poelau— rmnh 229 (cf. Hooijer, 1962b, p. 46, no.

99649, 99650; Ban Nong Kok-ZRC 4-034; Ban 48); Rantau— rmnh 4617 (cf. Hooijer, 1962b, p. Nong Put— ctnrc (catalog number unknown); Ban 44, no. 11); Riam— amnh 106282; Senoeang— Pak Nam Pho-Mcz 23812, 23813, zrc 4-031; amnh 1 07 1 00 (male skin mismatched with female Ban Phra Muang-USNM 83271, 83273; Ban Phu skull); Sintang— ansp 14036, 14040; no locality— Toie-FMNH 99639, 99640; Ban Sai Kau-BM(NH) zmb (unnumbered, 24 Apr. 1902), zsbs Coll. Nos. 1903.2.6.3; Ban ThapPlik, 1 kmE-FMNH 105654; 3PS,4PS,7PS, 10-12, 1 5 Guvenile male), 15 (adult Ban Thap Plik, 1 km NE-fmnh 105653; Kam- male), 16, 21, 36(450), 48, 157, 201, 205, 276, pong Biserat-BM(NH) 1903.2.6.1, 1903.2.6.4; 277, 297, 298, 306, 372, 373, 445(316), 587, 616- Kantang— zrc 4-032; Kata Taek— fmnh 99651- 618, 694, 701, 709, 710, 730, 740, 741, 743, 749, 99656; Khao Rang Kai— ctnrc (catalog number 764-766, 771, 780, 1013, 1014, 1228, 1358, 1359, unknown), fmnh 105689; Laem Sing mountains— 1372, 1378, 1386, 1387, 1414, 1437, 1438, 1452, usnm 251662; Lat Bua Khao, Sathani— usnm 1464, 1465, 1477, 1478, 1488, 1505, 1510, 1534, 236631, zrc 4-033; Tham Hom-FMNH 105649; unnumbered (Breugel), unnumbered (Breugel, Tyching-USNM 83274, 83275; Butang(l), no lo- "von Zengen"); Java (25), Bantargebang— mcz cality— bm(nh) 1955.1511; Chang (3), no locali- 12760; Gedangan— rmnh (unnumbered); Indra- ty-BM(NH) 1915.1 1.4.7, usnm 201551, zrc 4-039; mayu— mzb 2939, 2960; Kawarasan— rmnh 2067;

122 FIELDIANA: ZOOLOGY Pangandaran— bm(nh) 1909.1.5.26; Pangrango, side); Kuching—smk 092/5 (skull inside); Ku- Gunung— rmnh 2067, 2067/II (skulls inside); Se- ching, 10th mile— smk 092/12 (skull inside); Ling- marang— zsbs (unnumbered); Tasikmalaya— mzb ga-SMK 092/2, 092/7 (skulls inside); Miri-Mcz 1610; Java, West— rmnh 1, 5e, 6, 7g (skulls in- 6662 (skull inside); Miri district— NHMBe 817; side), 8; no locality— NHMBa 3131, 3161, 3352, Penrissen, Gunong-SMK 092/4, 092/1 1/A3.312 rmnh 9i, lOj, Ilk, 121, 13m, 14 (skulls inside); (skulls inside); Punang, Sungai— smk 092/6, 092/ smtd (unnumbered); Kambing (1), no locality— 1 0; Segobang, Sungai -smk 092/ 1 1/47 1 3 (skull in- rmnh 30p (skull inside); Laut (1), no locality— side); Dindding(l), no locality— bm(nh) 1905.3. 1.3. bm(nh) 1939.1054 (skull inside); Natuna Besar MALAYSIA: WEST MALAYSIA, mainland (4), (6), Sinubing-BM(NH) 1939.1053 (skull inside); Melaka-MNHN 1848.401, 1848.402 (skulls in- Ulu, Sungai— zrc 4-138; no locality— bm(nh) side), rmnh 40, 42 (skull inside); Aur (1), no lo- 1939.1055, 1964.428 (skulls inside), mcz 6660, cality—zrc 4-079. 6661 (skulls inside); Mas (1), no locality— zrc PHILIPPINES, 3. Mindanao (1), Zamboanga 4-135 (skull not located); Pejantan (1), no locali- del Sur Prov.— siconbrec 1586 (living captive); ty— zsbs I; Sumatra (114), Batangkuis— zsbs Coll. Tawitawi (2), no locality— siconbrec 1225, 1475 Nos. 3, 1 3, 1 5, 38, 47, 88, 96, 97, 1 97, 200; "Bran- (living captives). dan?"-ZMB 38542; Deli, Sungai— ansp 20218, SINGAPORE, 5. Singapore (5), Botanical Gar- 20220 (fetuses in fluid); "I. Lendung"— zsbs Coll. den— zrc 4-081 (skull inside), 4-083; Punggol— Nos. 32, 74, unnumbered juvenile; "Kg. Baru"— zrc 4-087, 4-088; Sembawang, Sungai— zrc 4-089. zsbs Coll. Nos. 63, 77, 79, 81, 113; Medan-ZSBS THAILAND, 3. Mainland (3), Bangkok— mnhn Coll. Nos. 1, 6/II, 7, 7/II, 8/II, 9/II, 14, 24, 28 1219 (skull inside); Ban Thap Plik, 1 km E—fmnh (juvenile), 29, 42/111, 43, 47/11, 52, 58, 60, 62, 67, 105655 (infant in fluid); Kampong Biserat— bm(nh) 67/11, 71, 74, 75, 77, 80, 81, 81/11, 82, 90, 93, 94, 1903.2.6.2.

98, 99/11, 100/11, 102, 103, 104, 106, 111, 115, VIETNAM, 5. Mainland (5), Ho Chi Minh 1 16/VIII, 119, 124/III; Menam-ZMB 40820; Pa- City-NMS 5786, 5787; Ho Chi Minh City, Bo- dang—rmnh 44x, 45y, 46z (skulls inside), 47a, tanical Gardens— bm(nh) 1928.7.1.9, mnhn 48b, 49c, 50d (skulls inside), zmuz 1 1625, 1 1626; 1929.457; Tay Ninh-BM(NH) 1928.7.1.7. Palembang, Kotamady— zsbs Coll. Nos. 22, 23; Pangkalansusu— usnm 502457 (infant in alcohol); Skulls only, 526 1 1 1 1 Sumatra, east coast— zsbs Coll. Nos. 58, 70, 7 ,

189; Sumatra, south-ZMB 38540, 38541; Tan- INDONESIA, 434. Bali (2), Bratan, Danau- jungmorawa— zsbs Coll. Nos. 60, 73[b], 75[b]; no bm(nh) 1913.3.6.5 (possibly skull of bm(nh) locality— mnhn 362/233, zmb Coll. Nos. 9, 25, 1913.3.6.7, skin only); Jembrana—rmnh Coll. No. unnumbered juvenile, unnumbered male, zmuz E136 (external measurements from Sody, 1933, p. 11619 (skull inside), zsbs Coll. Nos. 14, 27, 52, 94); Bangka (173), Jebus-RMNH Coll. No. Bk7; 61, 68, 70, 78, 78/1905, 95, 111, 115, 117, 123, no locality— rmnh Coll. No. Bk74, zluu 1, 2, 4- 141, 155, 198, 199, 201, 202, 216, 235, 236, 242; 16, 18, 20, 22, 25-30, 32-37, 39^13, 45-62, 64- Sumbawa (1), Batudulang— nms 16631 (cranial 67, 73-75, 77-105, 106 (juvenile cranium only), measurements from Mertens, 1936, p. 319, and 106 (adult mandible only), 107-112, 113, (juve- G. H. Albrecht, pers. coram., Oct. 1991); Timor nile cranium only), 113 (adult mandible only), 1 1 4- (6), Amarassie— rmnh 29o (skull inside); Bo- 116, 117 (cranium and mandible), 1 17 (mandible kong— zsbs 1911/2110; Lelogama— zsbs 1911/ only), 118, 119, 120 (cranium only), 120 (man- 2103, 1911/2106, 1911/2107; Nikiniki-ZMB dible only), 121-134, 136, 137, 140-165, 167- 92133; island unknown (1), Sunda Islands— zmb 171, 173-178, 179 (mandible only), 181-184 111. (mandibles only), 185, 187-189 (mandibles only), INDONESIA OR MALAYSIA, 1. Borneo (1), zsbs 1964/232; Bintan (2), Pasir Panjang— zrc Sendung—zmb (unnumbered). 4-092, Sungei Biru-ZRC 4-091; Borneo: Kali- MALAYSIA, 24. MALAYSIA: SABAH, Borneo mantan (24), Borneo, southeast— rmnh p, zmb (2), Abai-Mcz 35695; Tuaran-Mcz 37407 (male 38543; Gosong Djerong— usnm 196827; Jembay- skin mismatched with female skull); Sebatik (1), an, Sungai-USNM 199181, 199183, 199184, Kar- no locality- amnh 30619. MALAYSIA: SARA- angan, Sungai— usnm 198300; Kariorang—zmb WAK, Borneo (15), Baram, Batang—nms 4689, 48494, unnumbered; Loa Bambam—usnm 196826; smk 092/1 3 (skull inside); Belaga or Kalulong, Bu- Long Peleben— mnhn 1974; 144; Purukcahu— kit— bm(nh) 1 95 1 .67 (female skin mismatched with bm(nh) 1910.4.5.18; Tanjung—NHMBa 3747; no male skull); Dulit, Bukit— smk 092/9 (skull in- locality— amnh 107100 (female skull mismatched

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 123 1 1 1 with male skin), rcs(om) G 66.3 , zsbs 944/3009, Sumatra, east coast— zsbs Coll. Nos. 157, 165, 169, Coll. Nos. 12-4-12-6, 20, 363/368, 748, 848, 864; 175; Tamiang— rmnh Coll. No. F143; Tanjung- Java (123), Bandung— rmnh 3438, zluu (unnum- morawa— rmnh q, z, unnumbered skeleton, zsbs bered); Cirebon — nms 6974, rcs(om) A85.6, Coll. Nos. 71 (adult male), 73 (subadult male), 75 A85.61-A85.65; Gedangan-RMNH Coll. VI, un- (adult female); Wonosobo— rmnh Coll. No. 60; numbered adult males (2), unnumbered infant (1); no locality-NMS 1045, 1046, 1101, 16635/IE.2.a, Java, West— rmnh cat. ost. b-cat. ost. e (4); Jock- rmnh 1839, 1872a (female), 1872a (male), 3322, aboemi—zmb A34.09; Malabar-ZLUU (unnum- 4658, zmb Coll. Nos. 6 Guvenile), 6 (adult male), bered); Manggis-ZSBS 1909/4062, 1909/4601, 15, unnumbered female, unnumbered male, zsbs unnumbered specimens (7), Margomulio, Gun- Coll. Nos. 33, ?5 1 (adult male), 62 (subadult male), ung— zsbs (unnumbered); Ngawi— rmnh 4660; 93, 105, 109, 112, 115-117, 120, 202; Sumbawa Pangandaran bm(nh) 1909.1.5.25 (with mounted (4), Dompu—nms 1027, 16633; Semongka— pri skin, not examined); Pangrango, Gunung— nhrm 1399; no locality— NHMBa 5341; Tanahmasa (1), Z2923-Z2925, Z3146-Z3150, Z3152-Z3154, no locality— usnm 121836 (external measure- Z3165, Z3166, Z3292, Z3293, Z3295, Z3314, ments recorded on skull tag); Timor (2), Bokong— Z3316, Z3320-Z3322, Z3324 (adult female), zsbs 1911/2109; no locality — NHMBa 2703;

Z3324 (subadult female), Z3325, Z3328-Z3330; Tuangku (2), north coast— usnm 1 1441 1, 1 14643 Salak, Gunung— rmnh Coll. Nos. A 137, A221; (external measurements recorded on skull tag); is- Tjeringin— rmnh Coll. No. 12f; Tulungagung— land unknown (1), Sunda Islands— NHMBa 2981. rmnh 4650-4655; no locality— ansp 12, NHMBa MALAYSIA, 82. MALAYSIA: SABAH, Borneo (unnumbered), nhrm Z2698-Z2700, Z2702, (65), Abai-Mcz 25699, 25711, 35569, 35571, Z2709-Z2711, Z2718-Z2720, Z2726-Z2729, 35576, 35578, 35587, 35608, 35619, 35622,

1 1 1 Z2730 (mandible only), Z273 , Z2739, Z3 5 , nms 35623, 35626, 35633, 35634, 35643, 35651, 1044, 1100, rmnh 2615i, 2651b-2651h, 2683, 35652, 35655, 35656, 35658, 35661, 35673, 3087-3089, 3099, 3106, 3107, 3109, 31 10, 31 13- 35677, 35681, 35725, 35730, 35734-35736, 3115, 3177-3179, 3188, 3229, 4644, 6684, cat. 35738, 35739, 35741, 35742, 35746, 35748- ost. a, cat. ost. i-cat. ost. 1 (4), cat. ost. n, zmb 35755, 35758 (cranium embedded in plaster), 43867; Karimun (1), Pongka, Kampung— usnm 35759, 35764-35767, 35768/491, 37663, 37664 122848; Kundur(2), no locality-ZRC 4-095, 4-096; (external measurements for 50 of these 5 1 speci-

Lombok (2), Suranadi— pri 1400, no locality— mens, all except mcz 35622, are listed on expe- bm(nh) 1 920. 1 .26.3; Matasiri(l), east coast— usnm dition record cards); Kinabalu, Mt.— mcz 37781— 154368;Mwrsfl/tf(l)-ushJMll4561;Mas(5),Sia- 37786, 37787-37789 (three mandibles only); Kin- ba, Teluk-usNM 121872-121874; no locality - abatangan, Sungai— usnm 19192; Sandakan, 8 mi zmb 48417, 48419; Padang{\), north coast— usnm W— fmnh 33547; Segama, Sungai— zmb A85.10; 143583; Sumatra (87), Balbalan-ZMB 34005; Ba- Talibang— mcz 57836; Tuaran— mcz 37407 (fe- tangkuis— zsbs Coll. Nos. 110, 114; Bengkulu— male skull mismatched with male skin). MALA Y- rmnh 4648; Bukittinggi— zmb 34241; Indragiri SIA: SARAWAK, Borneo (12), Baram, Batang- district— NHMBa 3774-3776, 3778-3781, zmuz ansp 6149, bm(nh) 1894.6.12.13; Belaga or Ka- 11668; Kateman, Sungai— usnm 123147; "Kg. lulong, Bukit— bm(nh) 1951.67 (male skull mis- Baru"— zsbs Coll. Nos. 62 (adult male), 113; La- matched with female skin), 1 95 1 .69, 1 95 1 .70; Du- hat— NHMBa zsbs 1805, 4829; Lampung, Propinsi— lit, Bukit— zmb (unnumbered); Jumpit— zmb rmnh (unnumbered); Lubuksikaping— 4659; Me- A 1870; Kuching-SMK Coll. Nos. 3-315, 3-320; dan— mcz 41 zsbs Coll. Nos. 28 167; (adult male), Melinau Gorge— bm(nh), unnumbered mandibu- zmb A 14.08 67, 151; Mempura— (subadult male), 1 1 2. 1 lar fragment; Mulu, Gunong— bm(nh) 894.6. ; A 14.08/02 1 (infant); Padang-ZMUz 1652; Pagan- Puram-ANSP 6113. MALAYSIA: WEST MA- san-ZMB A 14.08 (subadult female), A 14.08/03- LAYSIA, mainland (5), Benom, Gunong, north- A 14.08/05; Pangkalan-RMNH 4645-4647; Pang- east slope-BM(NH) 1979.2869; Nyalas-ZRC 4-076 kalansusu— usnm 536025; Pap-ka—zmb A 14.08; (external measurements from skull tag), 4-077 (ex- Payakumbuh— rmnh 4649; Rawas—zmb 34025; ternal measurements from Weitzel et 1988, p. Rokan-kanan, Sungai-ZMB A 14.08/08, A 14.08/ al., Telok Anson-NHRM 3260. 1/07, A 14.08/11/01; Serangjaya-hilir-RMNH Coll. 107); 3259, Nos. F144, F145; Siak Copatta-SMB A14.08/53; PHILIPPINES, 2. Jolo (1), Crater Lake Moun- Sijunjung— rmnh w; Soekaranda— zmb A22.05; tain— usnm 1 25324; Mindanao (I), Zamboanga— Sumatra, [east-central]— zmb A 14.08 (infant), zmb A2920 (external measurements from Mar- A 14.08 (juvenile female), A 14.08 (subadult male); tens, 1876, p. 206).

124 FIELDIANA: ZOOLOGY SINGAPORE, 1. Singapore (1), Botanical Gar- 65 km E of Um Pang—amnh 54677, bm(nh) den— zrc 4-085. 1924.9.2.8; Wong, Nam Mae, 85 km E of Um THAILAND, 4. Mainland (3), Ban Sakaerat- Pang—amnh 54679. nhrm Coll. No. 1/3; Kampong Biserat— smtd B4346; Pattani— smtd B4348; Tarutao (1), no lo- Skins Only, 8 cality— usnm 123992. VIETNAM, 3. Mainland (3), Cochin China- BURMA, 6. Mainland (6), Arakan Div.— zsi mnhn 1869.297; Ho Chi Minh City, Botanical 1 1990; Haungtharaw-zsi 4389, 4390, 4392 (skulls Gardens—mnhn 1962.1445; Ho Chi Minh City, inside all three); Mergui, bnhs 5078, zsi 5238. Zoological Garden— mnhn 1962.1443. COUNTRY UNKNOWN, 2. "Bengale"-MNHN 362/234 (lectotype), 368/247. Supplementary information has been derived from 1 2 unexamined specimens of M. f. fascicu- laris collected at the following localities: INDO- Skulls Only, 3. NESIA, Borneo: Kalimantan, Kaboerau (Gyld- BURMA, 3. Mainland (2), Mergui— bm(nh) enstolpe, 1920, pp. 14-15); Loa Bambam (Coll. 1856.5.6.16; Tenasserim-BM(NH) 1972.1312; No. 10, H. C. Raven field catalog, usnm); Long Lanbi (1), no locality— usnm 1241 12. Pangian (Gyldenstolpe, 1920, pp. 13-14); Pelai- hari (Kohlbrugge, 1 896a, p. 185); Java, Kediri dis- Supplementary information has been derived trict— zsbs 191 1/2363, 191 1/2364 (specimens not from a published photograph of M. f. aurea ob- seen, cranial measurements from G. H. Albrecht, served at BANGLADESH, mainland, Whykeong

pers. comm., Oct. 1991); Tengger, Pegunungan Union Council (M. A. R. Khan, 1985, fig. 15). (Kohlbrugge, 1896b, p. 280); Sumatra, Labbu- handeli vicinity (Hagen, 1890, p. 82); Sumbawa, Dompu—nms 16629 (not seen, cranial measure- ments from Mertens, 1936, p. 319, and G. H. Macaca fascicularis aurea/ Albrecht, pers. comm., Oct. 1991). MALAYSIA: Macaca fascicularis fascicularis SABAH, Borneo, Abai—mcz 35768/492 (not seen, Contact Zone or external measurements from expedition record Macaca fascicularis/Macaca mulatta cards). PHILIPPINES, Mindanao, Zamboanga- Contact Zone, Total 45 2 specimens (Martens, 1876, p. 206). Skins and Skulls, 41

BURMA, 11. Mainland (10), Pakchan R., near Maliwun—amnh 54972; Pakchan R., near Ban- Macaco fascicularis aurea, Total 55 kachon-BM(NH) 1914.12.8.11, 1914.12.8.14; Thagyet-BM(NH) 1914.12.8.12, 1914.12.8.13, Skins and Skulls, 44 bnhs 5071, 5072, 5074; fmnh 82805, zrc 4-024; BURMA, 33. Mainland (23), Ban Sadein— amnh Ru (1), no locality— bmnh 5077. 54968; Haungtharaw-zsi 1 1989; Tagoot, bnhs 5073, LAOS, 3. Mainland (3), Thateng, Muang—amnh 5075; Taungbyauk Chaung-BM(NH) 1936.9.10.8, 87266, ansp 15136, 15138 (external measure- 1936.9.10.9, 1936.9.10.12, 1936.9.10.13; Tavoy ments from amnh catalog). R.-bm(nh) 1936.9.10.10, 1936.9.10.14; Tenas- THAILAND, 26. Mainland (22), Ban Mae Na serim-BM(NH) 1914.12.8.15-1914.12.8.18, bnhs Ree-FMNH 99657-99659; Ban Na-535154; Ban 5070, 5076, 5079, fmnh 82804, zrc 4-02 1-4-023; Nam Lai Tai— ctnrf (catalog number unknown), Wimpong— bm(nh) 1885.8.1.15; Ye Forest— fmnh 99660-99663, 99666, 99667; Chumphon, bm(nh) 1910.12.24.1; Kadan (1), no locality— Khlong-ZRC 4-029, 4-030; Nakhon Si Tham- bm(nh) 1925.7.2.1; Kathema (4), no locality— marat— usnm 251661, zrc 4-035-4-037; Pak bm(nh) 1936.9.10.4-1936.9.10.7; Lanbi (1), no Chong, Sathani—zrc 4-025, 4-026; Phu Phan— locality— usnm 104442; Letsok-aw (1), no locali- usnm 3077 14 (skin)/307732 (skull); Pran Buri, Mae

ty— usnm 124176 (cranium missing); Mibya (2), Nam-ZRC 4-027, 4-028; Naka Yai (1), no local- no locality-BM(NH) 1936.9.10.3, 1910.9.10.11; ity— zrc 4-043; Phayam (1), no locality— zrc Zadetkyi (1), no locality— usnm 1 1 1898. 4-038; Rang Yai (1), no locality— zrc 4-042; Yao

THAILAND, 11. Mainland (1 1), Ban Tamrong Noi (1), no locality— zrc 4-052. Phato— ctnrc (catalog number unknown), fmnh VIETNAM, 1. Mainland (I), Lac Giao-FMNH 99641, 99642, 99644-99648; Wong, Nam Mae, 46523.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 125 Skins Only, 3 merly known only from its skin, is now known from both skin and skull. LAOS, 2. Mainland(2), Thateng, Muang— amnh 4. bm(nh) 1872.9.20.1, skin and skull, Negros I., 87270,87271. S (M.f. philippinensis/M.f fascicularis Contact THAILAND, 1. Mainland (1), Ban Nam Lai Zone), not included in previous lists, was ex- Tai— fmnh 99664 (infant in fluid). amined after those lists were prepared. 5. fmnh 65453, skin only, formerly listed as of Skull Only, 1 unknown origin and undetermined subspecies, THAILAND, 1. Mainland (1), Nakhon Si is now known to have been collected at Negros: Thammarat— usnm 251660. Amio riverbank (M.f. philippinensis/M.f. fas- cicularis Contact Zone).

Supplementary information has been derived M. / philippinensis, from two unexamined specimens of M. f philip- Total 98 (Skins and Skulls, 53; pinensis collected at the following localities: Skins Only, 15; Skulls Only, 30) PHILIPPINES, Luzon, Nagpartian — usnm 144677 (transferred to Bureau of Science, Manila; Probably M. /. external measurements from Hollister, 1913, p. philippinensis, Total 15 330); Palawan, Brookes Point— fmnh 62916 (in- (Skin Only, 1; Skulls Only, 14) fant in fluid, not seen; external measurements from field catalog). M. / philippinensis/ M. f. fascicularis Contact Total 110 Zone, Macaco fascicularis umbrosa, Total 8 (Skins and Skulls, 82; Skins Only, 5; Skulls Only, 23) Skins and Skulls, 8

INDIA, 8. Great Nicobar(3), Kopenheat— usnm Probably M. /. 111799; no locality-usNM 111792, 111793; philippinensis/M. f. fascicularis Katchall (2), no locality-usNM 1 1 1801, 1 1 1802; Contact Zone, Total 12 Little Nicobar (3), no locality— usnm 1 1 1795 (ho- (Skins and Skulls, 2; Skin Only, 1; lotype), 111796, 111797. Skulls Only, 9)

For detailed lists of M. f philippinensis and M. Macaca fascicularis fusca, Total 19 f. philippinensis/M. f. fascicularis contact zone specimens examined, see Fooden (1991, p. 32). Skins and 1 2 Five revisions have been made to the previously Skulls,

lists, as follows: published INDONESIA, 1 2. Simeulue (12), Ajer Dingin- Lhok-usNM 1 14162, 1. bm(nh) 1877.10.6.2, Mindanao: Agusan River, bm(nh) 1923.10.7.1; Dalam, 1 14164 114165-114167; La- is now recognized as a composite consisting of 114163, (holotype), — usnm the skin of an infant or juvenile mismatched buhanbajau 114169; Sibaboh, Lugu— 121511-121513. with the skull of a subadult female (see above, usnm 114168, M. f philippinensis account, Distribution). This Skins 7 skin and skull, formerly assigned to M. f phi- Only, lippinensis, are now tentatively reassigned, as INDONESIA, 7. Simeulue (7), Ajer Dingin- two to separate specimens, "Probably M.f phi- bm(nh) 1923.10.7.2; Sinabang— rmnh Coll. Nos. lippinensis/M. Contact Zone." f fascicularis 20, 140, 1160, 1161;nolocality-RMNH574, 1330. 2. 1 1 1 bm(nh) 877. 0.6. , skin and skull, Mindanao: Surigao, collected near the Agusan River (see above) and formerly assigned to M. f philip- Macaca fascicularis lasiae, Total 2 pinensis, is now also tentatively reassigned to "Probably M.f. philippinensis/M.f fascicularis Skins and Skulls, 2 Contact Zone." 2. no -usnm 3. bm(nh) 1872.8.20.5, Negros I., S (M.fphilip- INDONESIA, Lasia (2), locality pinensis/M. f fascicularis Contact Zone), for- 114247, 114248 (holotype).

126 FIELDIANA: ZOOLOGY Macaco fascicularis atriceps, Total 1 1 imen was subsequently located and examined by G. H. Albrecht (pers. comm., Oct. 1991), who Skins and Skulls, 1 1 kindly provided the measurement of its greatest - length, excluding incisors. THAILAND, 11. Khram Yai(l 1), no locality bm(nh) 1939.891, 1939.892, usnm 236618- Skulls Only, 2 236621, 236622 (holotype), zrc 4-012, 4-013 (skull)/4-733 (skin), 4-015, 4-016. For external INDONESIA, 2. Karimunjawa (1), no locali- measurements of zrc specimens, cf. Kloss (1921, ty—mzb 1454; Kemujan(l), no locality— mzb 2720. p. 77) and Weitzel et al. (1988, p. 105).

Macaca fascicularis, Subspecies Undetermined, Total 95 Macaca fascicularis condorensis. Total 16

Skins and Skulls, 1 5 Skins and Skulls, 1 1

MAURITIUS, 3. Mauritius (3), Plaine Sophie - VIETNAM, 11. Ba (3), Ben Dam, 1.2 km W, 1950. no 0.1 km S-usnm 357242; Ben Dam, 1.7 km W- bm(nh) 1458; locality -bm(nh) 1861.6.1.6, 1909.3.12.1. usnm 357239; Ben Dam, 3.1 km W, 0.6 km S— PHILIPPINES, 10. See Fooden (1991, p. 33). usnm 357241; Con Son (8), Airfield building, 2.5 THAILAND, 2. Mainland (2), locality un- km E, 0.7 km N—usnm 357014; Airfield building, known-BM(NH) 1860.4.20.2, zsbs Coll. No. 95. 3.3 km E, 0.6 km N— usnm 357349; no locality— bm(nh) 1939.893, 1939.894, 1947.1498 (holo- Skins Only, 20 type), 1955.1528, zrc 4-018, 4-019.

1BURMA, 3. Mainland (3), "Rangoon"— rmnh Skins 2 Only, 51/j, 52/k; no locality— bnhs Coll. No. 7. MAURITIUS, 2. Mauritius (2), Kanaka - VIETNAM, 2. Con Son (2), no locality— mnhn bm(nh) unnumbered female; no locality— rmnh 1882.5 (skull inside), zrc 4-017. L-2 (skin inside).

9. See Fooden 99 1 and Skulls Only, 3 PHILIPPINES, (1 , p. 33) above, M. f. philippinensis/M. f. fascicularis Con- VIETNAM, 3. Ba (1), Ben Dam, 1.7 km W- tact Zone, fmnh 65453. usnm 357240. Con Son Airfield 0.5 (2), building, THAILAND, 5. Mainland (5), Phet Buh-mnhn km 4 km E— usnm no mnhn N, 357401; locality— 1 220 (lateral facial crest not examined); no local- 1882.1. ity— amnh 37208, mnhn 462, 767, zsbs Coll. No. 89. COUNTRY UNKNOWN, 1. No locality-ZRC Macaca fascicularis tua, Total 4 4-123.

Skins and Skulls, 4 Skulls Only, 60

INDONESIA, 4. Maratua no usnm (4), locality— 1BURMA, 2. Mainland (2), "Bengalen"— nms 197660-197662, 197663 (holotype). 1043, zmb 116.

MAURITIUS, 1. Mauritius (1), no locality - bm(nh) 1955.12.26.43. Macaca Total 9 fascicularis karimondjawae, PHILIPPINES, 52. See Fooden (1991, p. 34).

THAILAND, 5. Mainland {5), Ping, Nam Mae- Skins and 6 Skulls, amnh 5468 1 (external measurements from amnh catalog); "13°45'N, 99°25'E"-bm(nh) 1914.8.22.3, INDONESIA, 6. Karimunjawa (6), no locali- no Coll. Nos. 123. ty-MZB 1455-1458, 2717, 2718. 1914.8.33.4; locality-zsBS 27,

Skin Only, 1

INDONESIA, 1. Karimunjawa (1), no locali- ty— rmnh 10608 (holotype). The skull ofthis spec-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 127 Appendix 2: Gazetteer of 1 2. Locality code as indicated in distribution maps Non-Philippine (Fig. 2A-C). Macaca fascicularis Localities Abai; Borneo, MALAYSIA: SABAH; 5°42'N, 118°23'S; collected 2 Jun.-3 Aug. 1937 by S. as noted Introduction), Except previously (see L. Washburn and A. H. Schultz (see Coolidge, names listed as primary entries in this gaz- locality 1940, p. 124); mcz, 73 (including 1 skin only, etteer are official names approved in preferentially 51 skulls only). FAS; C:Sab-19. the U.S. Board on Geo- gazetteers published by Acheh, Pulau, MALAYSIA: WEST MALAYSIA; Bur- graphic Names (USBGN; Bangladesh, 1976; 2°39'-2°41'N, 103°46'-103°47'E; collected 18 1971a; India, 1952; In- ma, 1966a; Cambodia, Jul. 1915 by H. C. Robinson; bm(nh), 1. FAS; donesia, 1982; Laos, 1973; Malaysia, Singapore, B:SCS-2. and Brunei, 1970; Thailand, 1966b; Vietnam, Adenare. See Adonara, Pulau. in addition to lists ofplace- 1964,1971 b); providing Adonara, Pulau, Lesser Sunda Islands, INDO- names and USBGN also coordinates, gazetteers NESIA; 8°14'-8°25'S, 123°00'-123°20'E; re- contain useful glossaries ofgeneric geographic terms ported present before 1 937 by R. Mertens (1936, for each country. Macaque localities that are not p. 319). FAS;C:LS-21. listed in USBGN are spelled here as in gazetteers Agung, Gunung, See Desa Poetjang, Gunung the sources. with cross- original Secondary entries, Agung. references to entries, in- corresponding primary Ai Beta; Pulau Sumbawa, Lesser Sunda Islands, dicate variant spellings or alternate locality names INDONESIA; ca. 8°33'S, 1 17°25'E; blood sam- that on tags, in published litera- appear specimen ples taken Jul. 1981-Jan. 1982 by Y. Kawa- or in manuscripts on macaques. ture, unpublished moto, Tb. M. Ischak, and J. Supriatna (1982a, No secondary entries are supplied for variant p. 58). FAS; CLS-8. of Indonesian that contain spellings place-names Airabu, Pulau, INDONESIA; 2°43'-2°48'N, "oe" in of "u." For a of place gazetteer Philippine 106°12'-106°15'E; observed 18 Aug. 1899 by localities, see Fooden (1991, p. 34). macaque W. L. Abbott (in Miller, 1900, p. 244; cf. Kloss, The sequence of information presented in pri- 1903b, p. 68). FAS; B:SCS-27. entries is as follows: mary Airfield building, 0.5 km N, 4 km E, 80 m; Con 5 1 . Locality name. Son, VIETNAM; 8°44'N, 106°40'E; collected 2. Altitude, if reported by collector or observer. Mar. 1968 by M. L. Cunningham (see Duncan 3. Name of island (for noncontinental localities), et al., 1970, p. 180); usnm, 1 (skull only). CON; italicized. A:V-7. 4. Name of country, in capital letters. Airfield building, 2.5 km E, 0.7 km N, [0-590 m]; 5. Coordinates of locality (principal sources— Con Son, VIETNAM; 8°44'N, 106°37'E; col- USBGN gazetteers; published or unpublished lected 28 Feb. 1968 by J. F. Duncan (see Van field notes of collectors or observers). Peenenetal., 1969, pp. 100,285; 1970, p. 421); 6. Date of collection or observation. usnm, 1. CON; A:V-7. 7. Name of collector or observer. Airfield building, 3.3 km E, 0.6 km N, ca. 160 m; 8. Bibliographic reference (in parentheses) to Con Son, VIETNAM; 8°44'N, 106°37'E; col- published or unpublished field notes, if any. lected 12 Mar. 1968 by M. L. Cunningham (see

9. Abbreviated name of museum (see Introduc- Van Peenen et al., 1969, p. 100, 285; 1970, p. tion) where specimens are preserved. 421); usnm, 1. CON; A:V-7. 10. Number of specimens available (with indi- Ajer Dingin; Pulau Simeulue, INDONESIA; cation of part preserved, if skin and skull are 2°29'N, 96°23'E; collected 10 Mar. 1913 by E. not both present). Jacobson (1917, p. 276); bm(nh), 2 (including 1

1 1 . Abbreviated subspecific identification: ATR skin only). FUS; B:IO-3. = = M.f. atriceps; AUR M. f. aurea; AUR/ Akyab harbor. See Myengun Kyun. = FAS/MUL M. f aurea/M. f. fascicularis Alas, Lae, between Agusan and Ketambe Research contact zone or M. fascicularis/M. mulatta Station; Sumatra, INDONESIA; ca. 3°48'N, contact zone; CON = M.f. condorensis; FAS 97°33'E; observed Oct. 1984 by M. P. Ghiglieri = = M.f. fascicularis; FUS M.ffusca; KAR (1986, p. 106). FAS;B:S-13. = = M. f karimondjawae; LAS M. f lasiae; Alas, Lae, between Bengkong River and Gelom- = = TUA M.f. tua; UMB M. f umbrosa. bang; Sumatra, INDONESIA; ca. 2°53'N,

128 FIELDIANA: ZOOLOGY 97°53'E; observed Oct. 1984 by M. P. Ghiglieri Allen & Coolidge, 1940, p. 147); mcz, 1. FAS; (1986, p. 108). FAS;B:S-15. A.T-11. Alas, Lae, between Gelombang and mouth of riv- Arranya. See Aranyaprathet. er; Sumatra, INDONESIA; 2°32'N, 97°49'E; Ataran. See Ye Forest, Ataran district, Moulmein observed Oct. 1984 by M. P. Ghiglieri (1986, region. p. 108). FAS; B:S-16. Ataran River, between Podowk and Kya-eng; Alas, Lae, between Lae Renun and Bengkong Riv- BURMA; ca. 16°02'N, 98°00'E; observed 3 Feb. er; Sumatra, INDONESIA; ca. 3°02'N, 97°55'E; 1 859 by S. R. Tickell (1 859, p. 426). AUR; A:Bu- observed Oct. 1984 by M. P. Ghiglieri (1986, 11. p. 108). FAS;B:S-15. Atjeh. See Lesten, Daerah Istimewa Aceh. Alas, Lae, between Muara Setulen and Lae Renun; Aur, Pulau, MALAYSIA: WEST MALAYSIA; Sumatra, INDONESIA; ca. 3°10'N, 97°55'E; 2°26'-2°28'N, 104°30'-104°33'E; collected 14 observed Oct. 1984 by M. P. Ghiglieri (1986, Jun. 1912 by H. C. Robinson; bm(nh), 1. Col- lected 1-8 1927 2 p. 106). FAS; B:S-15. May by N. Smedley; zrc, (1 Alas Kedaton. See Kukuh. skin only). FAS; B:SCS-5. Babi, 96°37'- Alas River. See Alas, Lae. Pulau, INDONESIA; 2°03'-2°07'N, 96°42'E; primates reported absent 7-14 Jan. Alor, Pulau, Lesser Sunda Islands, INDONESIA; 1902 by W. L. Abbott (see Miller, 1903a, p. 8°08'-8°28'S, 124°20'-125°08'E; M.fascicularis 479). B:d. reported absent before 1937 by J. J. M. F. Sy- Badang, 400-600 m; Borneo: Kalimantan, IN- mons (Mertens, 1936, p. 319). C:h. DONESIA; 2°45'N, 1 15°43'E; collected 21 May- Amarassie; Pulau Timor, Lesser Sunda Islands, 1 1 Jun. 1935 by V. von Plessen (see Stresemann, INDONESIA; 10°20'S, 1 2 3°40'E; collected Jun. 1938, p. 109); amnh, 5; mzb, 1. FAS; C:K-35. 1829 by S. Muller and H. C. Macklot; rmnh, 1 , Sungai. See Badang. (skin only, skull inside). FAS; GLS-25. Bako National Park; Borneo, MALAYSIA: SA- Ambawang, Sungai, near Pontianak; Borneo: Ka- RAWAK; ca. 1°40'N, 110°26'E; reported pres- limantan, INDONESIA; 0°02'S, 109°42'E; col- ent in 1971 by G. Rothchild (1971, p. 169). lected 1 8 Jun. 1 905 W. L. Abbott by (field catalog; Observed Sep.-Dec. 1980 by R. E. Salter and in Lyon, 1907, p. 548); usnm, 1. FAS; CK-2. K. M. Aken (1983, p. 8). FAS; C:Sar-4. Pulau Lesser Sunda Is- Ampang area; Sumbawa, Bakong. See Bakung, Pulau. lands, INDONESIA; ca. 8°47'S, 1 18°00'E; blood Bakung, Pulau, INDONESIA; 0°0r-0°10'N, taken Jan. 1979-Dec. 1981 Y. Ka- samples by 104°23'-104°30'E; reported present 15-22 Jul. wamoto GLS-10. (1982, p. 67). FAS; 1903 by W. L. Abbott (see Miller, 1906c, p. Amphoe Fhang. See Ban Na. 283). FAS; B:SCS-12. Anak Krakatau, Pulau, INDONESIA; 6°06'S, Balambangan, Pulau, MALAYSIA: SABAH; 105°25'E; primates reported absent 1928-1934 7°10'-7°21'N, 1 16°51'-1 17°01'E; observed Feb.- by K. W. Dammerman (1948, p. 61); island Jun. 1991 by Shukor Md. Nor (pers. comm., 20 emerged in 1928. B:k. Jul. 1991). FAS; C:Sab-8. Anamba. See Siantan, Pulau. Balbalan; Sumatra, INDONESIA; not located, Angaur Island, MICRONESIA; 6°53'-6°55'N, 5°38'N-5°57'S, 95°12'-106°05'E; collected 4 Jul. 134°08'-134°09'E; introduced population, ob- 1904 by W. Volz; zmb, 1 (skull only). FAS; not served Jun.-Aug. 1973 by F. E. Poirier and E. mapped. O. Smith (1974, p. 258). Blood samples taken Bali, Pulau, Lesser Sunda Islands, INDONESIA; Nov.-Dec. 1986 by Y. Kawamoto, K. Nozawa, 8°04'-8°48'S, 114°26'-115°42'E; reported pres- K. Matsubayashi, and S. Gotoh (1988, p. 170). ent ca. May 1845 by H. Zollinger (1845, p. 46). Subspecies uncertain; not mapped. FAS; GLS-1 through LS-3. Arakan Division; BURMA; 17°22'-21°33'N, Bali Barat National Park; Pulau Bali, Lesser Sunda 92°10'-94°54'E; collected in 1871 by museum Islands, INDONESIA; ca. 8°08'S, 1 14°30'E; re- collector; zsi, 1 (skin only, skull inside). Two ported in 1982 by A. H. Robinson and Y. Rus- skulls (2 ID, 2 IE) presented in 1843 by A. P. tandi (1982, p. 14). Observed May-Jun. 1990 Phayre (see J. Anderson, 1881, p. 63) are not by A. Fuentes (in Wheatley et al., 1993, p. 1). now in the zsi collection. AUR; A:Bu-3. FAS; GLS-1. Aranyaprathet; THAILAND; 13°41'N, 102°30'E; Balipor. See Pajo, Danau Singkarak, Balipor dis- collected 6 May 1937 by J. A. Griswold, Jr. (see trict.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 129 Baluran Game Reserve; Java, INDONESIA; ca. V. Sody (1937, p. 248); rmnh, 3 (including 1 7°51'S, 11 4°22'E; observed Sep. 1969-Jan. 1971 skull only). Reported present Nov. 1971-Jan.

by W. Angst (1975, p. 326). FAS; CJ-41. 1 973 by C. L. Darsono (Crockett & Wilson, 1 980, Banda Aceh, ca. 20 km SSW; Sumatra, INDO- p. 156). FAS; B:SCS-18, SCS-19. NESIA; ca. 5°27'N, 95°16'E; observed Nov. Bangkaru, Pulau, Kepulauan Banyak, INDO- 1971-Jan. 1973 by C. M. Crockett and W. L. NESIA; 2°00'-2°07'N, 97°04'-97°09'E; mon- Wilson (1980, p. 156). FAS; B:S-5. keys reported absent 16-21 Jan. 1902 by W. L. Banda Aceh, ca. 30 km ENE; Sumatra, INDO- Abbott (see Miller, 1903a, p. 479). B:e. NESIA; ca. 5°36'N, 95°35'E; observed Nov. Bangko, ca. 1 5 km ENE; Sumatra, INDONESIA; 1971-Jan. 1973 by C. M. Crockett and W. L. ca. 1°49'N, 100°55'E; observed Nov. 1971-Jan. Wilson (1980, p. 156). FAS; B:S-6. 1 973 by C. M. Crockett and W. L. Wilson (1980, Banda Aceh, ca. 40 km S; Sumatra, INDONESIA; p. 156). FAS; B:S-37. ca. 5°10'N, 95°19'E; observed Nov. 1971-Jan. Bangko, ca. 20 km ENE; Sumatra, INDONESIA; 1 973 by C. M. Crockett and W. L. Wilson (1 980, ca. 1°49'N, 100°58'E; observed Nov. 1971-Jan. p. 156). FAS;B:S-4. 1973 by CM. Crockett and W.L.Wilson (1980, Banda Aceh, ca. 45 km S; Sumatra, INDONESIA; p. 156). FAS; B:S-37. ca. 5°07'N, 95°18'E; observed Nov. 1971-Jan. Bangko, ca. 20 km N; Sumatra, INDONESIA; ca. 1 973 by C. M. Crockett and W. L. Wilson (1980, 1°55'N, 100°51'E; observed Nov. 1971-Jan.

p. 156). FAS;B:S-4. 1 973 by C. M. Crockett and W. L. Wilson (1 980, Banda Aceh, ca. 85 km ESE; Sumatra, INDO- p. 156). FAS; B:S-37. NESIA; ca. 5°17'N, 96°02'E; observed Nov. Bangko, ca. 25 km ENE; Sumatra, INDONESIA; 1971-Jan. 1973 by C. M. Crockett and W. L. ca. 1°41'N, 101°02'E; observed Nov. 1971-Jan.

Wilson (1980, p. 156). FAS; B:S-7. 1 973 by C. M. Crockett and W. L. Wilson ( 1 980, Banda Aceh, near; Sumatra, INDONESIA; ca. p. 156). FAS;B:S-37. 5°34'N, 95°20'E; blood samples taken Nov.-Dec. Bangko, ca. 30 km NNW; Sumatra, INDONESIA; 1986 by J. R. de Ruiter (1993, p. 91). FAS; ca. 2°03'N, 100°46'E; observed Nov. 1971-Jan.

B:S-5. 1 973 by C. M. Crockett and W. L. Wilson ( 1 980, Bandar Seri Begawan, near; Borneo, BRUNEI; ca. p. 156). FAS; B:S-37. 4°53'N, 114°56'E; observed ca. 1980 by R. A. Bangko, ca. 80 km S; Sumatra, INDONESIA; ca. Mittermeier (1980, p. 252). FAS; C:B-1. 1°00'N, 100°49'E; reported Nov. 1971-Jan. 1973 Bandjar. See Tjeringin, near Banjar, Preanger re- by local residents (Crockett & Wilson, 1980, p. gion, east. 156). FAS;B:S-39. Bandung; Java, INDONESIA; 6°54'S, 107°36'E Bangko, ca. 90 km SSE; Sumatra, INDONESIA; collected 29 Apr. 1938 by N. Gosselaar; rmnh ca. POO'N, 101°05'E; reported Nov. 1971-Jan. 1 (skull only). FAS; B:J-7. 1973 by local residents (Crockett & Wilson, Bandung, 2000 ft (= 600 m); Java, INDONESIA 1980, p. 156). FAS;B:S-40. ca. 6°54'S, 107°36'E; collected in 1936 by D. P Bangkok; THAILAND; 13°45'N, 100°31'E; col- Bosscha Erdbrink; zluu, 1 (skull only); FAS lected in 1869 by M. Boucourt; mnhn, 1 (skin B:J-7. only, skull inside). FAS; A:T-30. Bandung, near, 700 m; Java, INDONESIA Bangli, near; Pulau Bali, Lesser Sunda Islands, "6°54'S, 107°37'E"; collected 7 Oct. 1971 by INDONESIA; ca. 8°27'S, 1 15°21'E; observed ca. NAMRU Djakarta Detachment; usnm, 1. FAS 1993 by B. P. Wheatley, A. Fuentes, and D. K.

B:J-7. Harya Putra (1993, p. 1). FAS; QLS-3. Banggi, Pulau, south, MALAYSIA: SABAH; ca. Banguey Island. See Banggi, Pulau, south. 7°06'N, 1 17°04'E; collected 4-5 Sep. 1927 by F. Ban Huai Maenam Noi, ca. 15 km SE, ca. 75 m; N. Chasen and C. B. Kloss (1931, p. 1); zrc, 3. THAILAND; 14°25'N, 98°51'E; collected 16-

C:Sab-9. 1 7 1 1 1 1 FAS; Feb. 967 by J. Fooden ( 97 , p. 6); fmnh, Bangka, Pulau, INDONESIA; r30'-3°07'S, 2. FAS; A:T-26. 105°06'-106°51'E; reported present before 1852 Banjak Islands. See Tuangku, Pulau and Bang- by P. Bleeker (1851, p. 527). Collected in 1898 karu, Pulau, Kepulauan Banyak. by Blonk; zsbs, 1 (skull only). Collected in 1 900 Banjar. See Tjeringin, near Banjar, Preanger re- by A. A. W. Hubrecht (see Kohlbrugge, 1902, gion, east. p. 322); zluu, 170 (including 158 skulls only, Banjoewangi. See Banyuwangi. = 12 mandibles only). Collected ca. 1936 by H. J. Banjoe Wetan (? Banyuwedang), sea level; Pulau

130 FIELDIANA: ZOOLOGY Bali, Lesser Sunda Islands, INDONESIA; Banta, Pulau, Lesser Sunda Islands, INDONE- ?8°08'S, 114°36'E; collected 18 Jul. 1933 by J. SIA; 8°24'-8°29'S, 119°16'-119°18'E; monkeys J. Menden; mzb, 3. FAS; OLS-1. reported absent 1969-1973 by W. Auffenberg Banka. See Bangka, Pulau. (1981, p. 40). C:d. Bankachon. See Pakchan River, near Bankachon. Bantam region. See Camara; Cihara; Cikujang; Pe- Ban Kosum; THAILAND; 15°50'N, 104°00'E; re- labuhanratu, Teluk; and Tamandjaija. ported present before 1978 by B. Lekagul and Ban Tamrong Phato, ca. 100 m; THAILAND; J. A. McNeely (1977, pp. 293, 725). Subspecies 14°54'N, 98°31'E; collected 10-1 1 Feb. 1967 by

uncertain; A:T-14. J. Fooden (1971, p. 15); ctnrc, 1; fmnh, 7. Ban Mae Na Ree, ca. 200 m; THAILAND; AUR; A:T-20. 16°25'N, 99°23'E; collected 13 Mar. 1967 by J. Bantargebang; Java, INDONESIA; 7°01'S,

1 1 1 3. collected Oct. Fooden ( 97 , p. 8); fmnh, AUR/FAS/MUL; 106°40'E; 21-23 1909 by O. Bry- A:T-2. ant and W. Palmer; mcz, 2 (including 1 skin Ban Me Thuot. See Lac Giao. only); usnm, 3. FAS; B:J-9. Ban Na; THAILAND; ca. 8°10'N, 100°10'E; col- area. See Danau, Rawa. lected 11 Feb. 1954 by B. Lekagul; usnm, 1. Ban Thap Plik. See Ban Nong Put; Tham Horn, AUR/FAS/MUL; A:T-64. 4 km W of Ban Thap Plik. Ban Nam Lai Tai, ca. 5 km W, ca. 300 m; THAI- Ban Thap Plik, 1 km NE, ca. 75 m; THAILAND; LAND; 16°10'N, 99°20'E; collected 4-5 Apr. 8°1 l'N, 98°53'E; collected 4-5 Jun. 1973 by J.

1 1 1 1 1 1 967 by J. Fooden ( 97 , p. 9); ctnrc, ; fmnh, Fooden ([1975], p. 98); fmnh, 3 (including 7 (including 1 infant in fluid). AUR/FAS/MUL; infant in fluid). FAS; A:T-60. A:T-1. Ban Wan; THAILAND; 15°23'N, 104°H'E; ob- Ban Nong Kok; THAILAND; 8°06'N, 98°52'E; served 12-14 Jul. 1989 by N. Aggimarangsee collected 1 1 Jan, 1918 by local collectors em- (1992, pp. 109, 120; pers. coram., Oct. 1993). ployed by H. C. Robinson and C. B. Kloss (1919, Subspecies uncertain; A:T-17. p. 8 7); zrc, 1. FAS; A:T-60. Ban Wang Kalang. See Khwae Noi, Mae Nam. Ban Nong Put, ca. 75 m; THAILAND; 8°11'N, Ban Wang Phato. See Ban Tamrong Phato. 98°53'E; collected 5 Jun. 1973 by J. Fooden Banyuwangi; Java, INDONESIA; 8°1 2'S, 1 14°2 1 'E; ([1975], p. 98); ctnrc, 1. FAS; A:T-60. collected 16 Dec. 1919 by unknown collector; Ban Pak Nam. See Chumphon, Khlong, mouth. rmnh, 1. FAS; CJ-42. Ban Pak Nam Pho, ? 1 50 m; THAILAND; 1 5°43'N, Baram, Batang; Borneo, MALAYSIA: SARA- 100°09'E; collected 2 Apr. 1924 by J. H. Cham- WAK; 4°35'N, 114°00'E; collected 1884-1898 brai [and K. G. Gairdner] (see Kloss, 1919a, p. by C. Hose and E. Hose (see Medway, 1 977, pp. 49; 1930, p. 61); zrc, 1. Collected 29-30 Aug. 3-4); ansp, 1 (skull only); bm(nh), 2 (including 1926 by C. J. Aagaard; mcz, 2. FAS; A:T-5. 1 skull only); nms, 1 (skin only); smk, 1 (skin Ban Palian. See Khao Rang Kai. only, skull inside); usnm, 1. Collected Jul. 1896 Ban Phra Muang; THAILAND; 7°18'N, 99°28'E; by W. H. Furness III (1896, p. 310) and H. M. collected 7 Mar. 1896 by W. L. Abbott (see Hiller; ansp, 1 (skull only). FAS; C:Sar-18. Riley, 1938, p. 12); usnm, 2. FAS; A:T-56. Baram district; Borneo, MALAYSIA: SARA- BanPhuToie, 100-200 m; THAILAND; 14°42'N, WAK; ca. 4°35'N, 1 13°58'E; collected date un- 99°07'E; collected 31 Jan. 1967 by J. Fooden known by unknown collector; University Mu- (1971, p. 14); fmnh, 2. FAS; A:T-21. seum of Zoology, Cambridge, 1 (skeleton only; Ban Sadein; BURMA; 10°20'N, 98°32'E; collected not seen, data from P. H. Napier, 1981, p. 14). 18 Feb. 1928 by Faunthorpe-Vernay Expedi- FAS; C:Sar-18. tion; amnh, 1. AUR; A:Bu-22. Baramfluss. See Baram, Batang. Ban Sai Kau, Nong Chik region; THAILAND; Bari; Pulau Flores, Lesser Sunda Islands, INDO- 6°38'N, 101°08'E;captivesobtained 17 Jun. 1902 NESIA; 8°21'S, 120°H'E; collected 21 Nov. by N. Annandale and H. C. Robinson (1903, p. 1888-9 Jan. 1889 by M. Weber (1890a, p. viii; xxxvii; cf. Bonhote, 1903, p. 3); bm(nh), 1. FAS; 1890b, p. 102); museum unknown, 1 (skull only, A:T-69. not seen). FAS; C:LS-16. Ban Sakaerat; THAILAND; 14°30'N, 101°56'E; Barisan Selatan; Sumatra, INDONESIA; ca. collected 8 Jan. 1 9 1 2 by N. Gyldenstolpe (1913, 3°00'S, 102°15'E; observed in 1983 by K S. p. 3; 1914, p. 1; cf. Thonglongya, 1967, p. 187); MacKinnon (1986, p. 1 1 2). Observed Jun. 1988 nhrm, 1 (skull only). FAS; A:T-10. by M. Bismark (1992, p. 11). FAS; B:S-66.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 131 Barito, Sungai. See Purukcahu, Sungai Barito. SIA; ca. 4°00'S, 104°17'E; observed Nov. 1971- Barito, Sungai, right bank, above ; Jan. 1973 by C. M. Crockett and W. L. Wilson Borneo: Kalimantan, INDONESIA; ca. 3°13'S, (1980, p. 156). FAS; B:S-74. 1 14°32'E; observed Jul. 1984-May 1986 by K. Baturaja, ca. 30 km NNE; Sumatra, INDONE- M. Burton (see Chivers & Burton, [1991], p. SIA; ca. 3°56'S, 104°20'E; observed Nov. 1971- 140). FAS; CK-25. Jan. 1973 by C. M. Crockett and W. L. Wilson Barren Island, , INDIA; 12°15'- (1980, p. 156). FAS; B:S-74. 12°17'N, 93°50'-93°52'E; primates reported ab- Baturaja, ca. 70 km WNW; Sumatra, INDONE- sent before 1903 by G. S. Miller, Jr. (1902, p. SIA; ca. 3°47'S, 103°39'E; observed Nov. 1971- 792; cf. Kloss, [1928], p. 802; Chaturvedi, 1980, Jan. 1973 by C. M. Crockett and W. L. Wilson p. 134). A:d. (1980, p.156). FAS;B:S-73. Batam, Pulau, Kepulauan Riau, INDONESIA; Baturaja, ca. 75 km NW; Sumatra, INDONESIA; o 58'-l°12'N, 103°54'-104°10'E; reported pres- ca. 3°42'S, 103°42'E; observed Nov. 1971-Jan. ent 15-17 Sep. 1 905 by W. L. Abbott (see Miller, 1973 by C. M. Crockett and W. L. Wilson (1980, 1906c, p. 282). FAS; B:SCS-10. p. 156). FAS; B:S-73. Batang Garut; Sumatra, INDONESIA; 2°23'N, Bawean, Pulau, INDONESIA; 5°43'-5°53'S, 99°38'E; observed Nov. 1971-Jan. 1973 by C. 112°33'-112°44'E; collected 24 Nov. 1907 by

M. Crockett and W. L. Wilson (1980, p. 156). W. L. Abbott (in Lyon, 191 1, p. 61); usnm, 3. FAS;B:S-31. Collected 6-12 May 1928 by Denin and P. F. Batangkuis; Sumatra, INDONESIA; 3°40'N, Franck; mzb, 6. FAS; C:J-38. 98°44'E; collected 1 906-1 9 10 by C. Bruegel; zsbs, Belaga; Borneo, MALAYSIA: SARAWAK; 1 3 (including 1 skins only, 2 skulls only). FAS; 2°42'N, 1 13°37'E; collected Oct.-Nov. 1932 by B:S-21. Oxford University Expedition to Borneo (see Batoe-meringgit; Pulau Bali, Lesser Sunda Is- Harrisson, 1933, p. 402; P. H. Napier, 1981, p. lands, INDONESIA; not located, 8°04'-8°55'S, 1 4); bm(nh), undetermined portion of 6 speci- 114°26'-115 43'E; collected 8 Oct. 1928 by P. mens (including 1 skin only and 3 skulls only)

F. Franck; mzb, 1 . FAS; not mapped. collected at Belaga and/or Kalulong (C:Sar-20). Battam. See Batam, Pulau. FAS;C:Sar-12. Batu, Tanjung; Pulau Belitung, INDONESIA; ca. Belawan; Sumatra, INDONESIA; 3°47'N, 98°41'E; 2°56'S, 107°32'E; collected 19-20 Jul. 1904 by blood samples taken Jan. 1979-Dec. 1981 by W. L. Abbott (in Lyon, 1906, p. 608); usnm, 2. Y. Kawamoto (1982, p. 67). FAS; B:S-21. FAS; B.SCS-20. Belitung, Pulau, INDONESIA; 2°32'-3°16'S, Batu Apoi Forest Reserve. See Kuala Belalong Field 107°32'-108°17'E; reported present before 1906 Studies Centre. by T. Willink (1905, p. 175). FAS; B:SCS-20. Batu Bara district; Sumatra, INDONESIA; ca. Ben Dam, 1.2 km W, 0.1 km S; Hon Ba, VIET- 3°00'N, 99°15'E; collected 1897-1899 by G. NAM; 8°39'N, 106°33'E; collected 2 Mar. 1968

Schneider (1905, p. 72); museum and number by J. F. Duncan (in Van Peenen et al., 1970, p. of specimens unknown (not seen). FAS; B:S-23. 421); usnm, 1. CON; A:V-7. Batudulang, 800-900 m; Pulau Sumbawa, Lesser Ben Dam, 1.7 km W, ca. 20 m; Hon Ba, VIET- Sunda Islands, INDONESIA; 8°36'S, 1 17°19'E; NAM; 8°39'N, 106°33'E; collected 2 Mar. 1968 collected 5-8 May 1927 by Sunda-Expedition by P. F. D. Van Peenen (in Van Peenen et al., Rensch (see B. Rensch, 1930, p. 79; Mertens, 1970, p. 421); usnm, 2 (including 1 skull only). 1930, p. 133; I. Rensch, 1934, p. 226); mzb, 2; CON; A:V-7. nms, 1 (skin only); zmb, 2. FAS; C:LS-8. Ben Dam, 3.1 km W, 0.6 km S, 100 m; Hon Ba, Batu Islands. See Tanahbala, Pulau; Tanahmasa, VIETNAM; 8°39'N, 106°33'E; collected 2 Mar. Pulau. 1968 by M. L. Cunningham (see Van Peenen et Batur, Gunung, inside cone; Pulau Bali, Lesser Sun- al., 1970, p. 421); usnm, 1. CON; A:V-7. da Islands, INDONESIA; 8°14'S, 115°23'E; un- "Bengale"; country unknown; acquired by L. T. confirmed report ca. 1993 by unspecified infor- Leschenault de la Tour (1820, p. 359; 1822, p.

mant (Wheatley et al., 1993, p. 1). FAS; CLS-3. 262), probably in Calcutta, Sep. 1 8 1 9-Jan. 1 820; Baturaden, Gunung Slamet, 700 m; Java, IN- mnhn, 1 (skin only). Captive acquired 5-27 Apr. DONESIA; ca. 7°19'S, 109°14'E; collected 9 Oct. 1837 at Diamond's Harbor, near Calcutta, by 1929 by H. J. V. Sody; rmnh, 1. FAS; B:J-24. J. F. T. Eydoux and L. Souleyet (1841, p. xiv, Baturaja, ca. 20 km NNE; Sumatra, INDONE- 6); mnhn, 1 (skin only). AUR; not mapped.

132 FIELDIANA: ZOOLOGY "Bengalen"; country unknown; acquired in 1852 Jul. 1912 by H. C. Raven (see Deignan, [1960], by E. Riippell; nms, 1 (skull only). Collected date p. 267); usnm, 6 (external measurements re- unknown by Frank; zmb, 1 (skull only). Sub- corded in field catalog). FAS; GK-42. species uncertain; not mapped. Berhala, Pulau, INDONESIA; 3°46'N, 99°31'E; Bengkalis, Pulau, INDONESIA; l 15'-l°37'N, primates reported absent in 1953 by J. L. Har- 101°59'-102°31'E; reported present 24 Feb.-3 rison and J. R. Hendrickson (1963, p. 548). B:a. Apr. 1906 by W. L. Abbott (in Lyon, 1908, p. Betotan, ca. 22 mi (= 35 km) SW of Sandakan, 31 624). FAS; B:SM-6. m; Borneo, MALAYSIA: SABAH; 5°48'N, Bengkulu; Sumatra, INDONESIA; 3°48'S, 1 1 7°50'E; collected 1 3 Aug. 1927 by F. N. Chas- 102°16'E; colelcted 1887-1941 by E. Dubois; en and C. B. Kloss (1931, p. 1; cf. Davis, 1962, rmnh, 1 (skull only). FAS; B:S-69. p. 126;Weitzeletal., 1988, p. 133); zrc, l.FAS; Bengkulu, ca. 40 km ENE; Sumatra, INDONE- C:Sab-16. SIA; ca. 3°38'S, 102°35'E; observed Nov. 1971- Bettotan. See Betotan. Jan. 1973 by C. M. Crockett and W. L. Wilson Big Kretam River. See Kretam Besar, Sungai. (1980, p. 156). FAS; B:S-70. Big Tambelan Island. See Tambelan Besar, Pulau. ca. Bengkulu, Propinsi (province); Sumatra, INDO- Bilasodia, Naf River; BANGLADESH; 2 observed 1982-Feb. NESIA; 2°23'-5°00'S, 101°06'-103°50'E; blood 1°05'N,92°12'E; Sep. 1983 M. A. R. Khan and M. A. Wahab samples taken Jan.-Nov. 1 979 by Y. Kawamoto by (1983, p. cf. M. A. R. and Tb. M. Ischak (1981, p. 238). FAS; not 104; Khan, 1981, p. 13; 1985, p. mapped. 30). AUR; A:Ba-l. Billiton. See Pulau and Benom, Gunong; MALAYSIA: WEST MALAY- Belitung, Batu, Tanjung; Pulau SIA; ca. 3°50'N, 102°06'E; collected 8 Nov. 1913 Belitung. Bima. See Kambing, Pulau. by museum collector (see H. C. Robinson & Bimirdia, Naf River; ca. Kloss, 1915b, p. 174); zrc, 2. FAS; B:WM-15. BANGLADESH; 21°05'N,92°12'E; observed Sep. 1982-Feb. 1983 Benom, Gunong, 4000 ft (= 1220 m); MALAY- by M. A. R. Khan and M. A. Wahab (1983, p. SIA: WEST MALAYSIA; ca. 3°50'N, 102°06'E; 104; cf. M. A. R. Khan, 1981, p. 13; 1985, p. observed 1968-1972 by unspecified informant 30). AUR; A:Ba-l. (Medway, 1972b, p. 120). FAS; B:WM-15. Binjai, Sungai; Pulau Natuna Besar, INDONE- Benom, Gunong, NE slope, 700 ft (= 2 10 m); MA- SIA; 3°47'N, 108°14'E; collected 27 Aug. and 5 LAYSIA: WEST MALAYSIA; 3°52'N, Sep. 1928 by F. N. Chasen (1935a, p. 5); zrc, 102°1 l'E; collected 1 Feb.-15 Apr. 1967 by Lord 2. FAS; B:SCS-33. Medway (1972a, p. 5; 1972b, p. 120; cf. Whit- Bintan, Pulau, Kepulauan Riau, INDONESIA; more, 1972, p. 12); bm(nh), 1 (skeleton only). 0°48'-l°13'N, 104°13'-104°34'E; collected 1 1 Jun. FAS;B:WM-15. 1930 by G. Nunong; zrc, 2. FAS; B:SCS-8. Benu, Pulau Timor, Lesser Sunda Islands, IN- Bintan, Pulau, north coast, Kepulauan Riau, IN- observed 24 DONESIA; 9°56'S, 123°59'E; Apr. DONESIA; ca. TIO'N, 104°30'E; collected 12 1930 by Jhr. W. C. van Heurn (1932, p. 65). Aug. 1902 by W. L. Abbott (field catalog; cf. FAS; C:LS-26. Miller, 1906c, p. 282); usnm, 1. FAS; B:SCS-8. Benua, Pulau, INDONESIA; 0°55'-0°58'N, Bintang, Pulau. See Bintan, Pulau. 107°25'-107°28'E; collected 6 Aug. 1899 by W. Birang, Sungai; Borneo: Kalimantan, INDONE- L. Abbott cf. (in Miller, 1900, p. 243; Kloss, SIA; 2°1 l'N, 1 17°27'E; collected 3 Oct. 1912 by 1903b, p. 61; Oberholser, 1919, p. 129); usnm, H. C. Raven (see Deignan, [1960], p. 268); usnm, 2. B:SCS-26. FAS; 1 (skin, skinned head in fluid; external mea- Berangkat, Gunong, east; MALAYSIA: WEST surements recorded in field catalog). FAS; GK- MALAYSIA; ca. 5°10'N, 102°06'E; reported 42. present in 1968 by D. Chivers (1971, p. 80). Biserat. See Kampong Biserat. FAS; B:WM-9. Blambangan-Purwo Nature Park. See South Ban- Berapit, Bukit, east; MALAYSIA: WEST MA- yuwangi Nature Park. LAYSIA; ca. 4°05'N, 103°20'E; reported present Bliah. See Selatbliat. in 1968 by D. Chivers (1971, p. 80). FAS; B:WM- Bodo, Gili. See Sababi, Pulau. 13. Boegies. See Bugis, Gili. Berau, Sungai, north bank; Borneo: Kalimantan, Boekit Sanggul (Benkoelan). See Sanggul, Bukit, INDONESIA; 2°09'N, 117°29'E; collected 29 Propinsi Bengkulu.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 133 Boeloengan. See Badang; Long Peleben; Sembak- Botanical Gardens, near Danau Buyan and Danau ung, Sungai. Bratan; Pulau Bali, Lesser Sunda Islands, IN- Bogor; Java, INDONESIA; 6°35'S, 106°47'E; col- DONESIA; ca. 8°15'S, 115°09'E; observed ca. lected 23 Sep. 1928 by G. Schiffer; mzb, 1. FAS; 1993 by B. P. Wheatley, A. Fuentes, and D. K.

B:J-17. Harya Putra (1993, p. 1). FAS; C:LS-3. Bohorok district. See Bungara; Bukitlawang. "Brandan?"; Sumatra, INDONESIA; not located, Bokong, 180 m; Pulau Timor, Lesser Sunda Is- 5°38'N-5°57'S, 95°12'-106°05'E; collected date lands, INDONESIA; 9°58'S, 124°04'E; collected unknown by W. Volz; zmb, 1 (skin only). FAS; 24-26 Jul. 191 1 by C. B. Haniel (see Hellmayr, not mapped. 1914, p. 5); zsbs, 2 (including 1 skin only, 1 skull Bratan, Danau, ca. 2500 ft (= 750 m); Pulau Bali, only). FAS; CLS-26. Lesser Sunda Islands, INDONESIA; 8°16'S, Bolongs, Outer. See Myengun Kyun. 1 15°1 l'E; collected Jan. 191 1 by E. Stresemann; Bolovens, Plateau des. See Thateng, Muang, Pla- bm(nh), 1 (skin only). Collected date unknown teau des Bolovens. by unknown collector; bm(nh), 1 (skull only; Bongao Peak, 70-300 m; Bongao Island, Tawitawi possibly skull of preceding). FAS; C:LS-3. Prov., PHILIPPINES; 5°01'N, 119°45'E; 4 Bratan, [Gunung], 1200 m; Pulau Bali; Lesser troops observed 24-26 Jan. 1990 by R. P. Ru- Sunda Islands, INDONESIA; 8°15'S, 1 15°12'E;

bio, P. L. Alviola III, and M. R. Felizardo ( 1 990, collected 3-8 Feb. 1 938 by V. von Plessen; amnh, p. 29). FAS; C: not numbered (new record, not 3. FAS; GLS-3. in Fooden, 1991, p. 2). Brunei Bay area; Borneo, BRUNEI; ca. 4°50'N, Bongkabong. See Rugading, near Bongkabong. 115°05'E; observed Nov. 1962-Mar. 1963 by J. Bongkabong, near sea level; Borneo, MALAYSIA: A. Kern (1964, p. 185). FAS; C:B-1. SABAH;ca. 5°59'N, 1 16°04'E; collected 21 Aug. Btg. Kwis. See Batangkuis. 1937 by J. A. Griswold, Jr. (see Coolidge, 1940, Bugis, Gili, Lesser Sunda Islands, INDONESIA; p. 123); mcz, 3. FAS; C:Sab-4. 8°30'S, 119°35'E; monkeys reported absent

1 1 1 Borneo: Kalimantan, INDONESIA; 4°25'N- 969-1 973 by W. Auffenberg ( 98 , p. 40). C:d. 4°10'S, 108°50'-119°00'E; collected in 1851 by Buitenzorg. See Bogor. J. H. Croockewit (see Medway, 1977, p. 3); Bukar, Sungai; Borneo, MALAYSIA: SARA- rmnh, 1 (skin only, skull inside). Collected be- WAK; ca. 1°16'N, 110°28'E; collected 2 Nov. fore 1857 by Verreaux; bm(nh), 1. Collected ca. 1919 by C. Ulok; zrc, 1. FAS; C:Sar-5. 1 897 by A. Harrison, Jr., and H. M. Hiller; ansp, Bukit Barisan Selatan, Taman Nasional, Lam-

1 . Collected 24 Apr. 1 902 by Dr. Pagel; zmb, 1 pung; Sumatra, INDONESIA; ca. 5°15'S, (skin only). Collected 1907-1910 by C. Bruegel; 104°15'E; observed Jun. 1988 by M. Bismark zsbs, 67 (including 60 skins only). Collected date (1992, pp. 11, 13). FAS; B:S-78. unknown by C. Bruegel; zsbs, 13 (including 4 Bukit Cangang; Sumatra, INDONESIA; ca. 0°10'S, skins only, 9 skulls only). Collected before 1948 100°05'E; observed 8 Sep. and 20-23 Oct. 1980 by unknown collector; rcs(om), 1 (skull only). by N. Koyama, A. Asuan, and N. Natsir (1981, Collected 8 Nov. 1961-6 Apr. 1962 by R. E. p. 1). Blood samples taken Jul.-Oct. 1980 by Y. Kuntz; usnm, 4. Collected date unknown by un- Kawamoto, K. Nozawa, and Tb. M. Ischak known collector; amnh, 1 (skull only; female (1981, p. 16). FAS; B:S-44. skull mismatched with male skin). FAS; not Bukit Cheraka. See Cherakah, Bukit. mapped. Bukit Garam; Borneo, MALAYSIA: SABAH; ca. Borneo: Kalimantan, southeast, INDONESIA; 5°28'N, 1 17°52'E; observed 25 May-23 Jul. 1963 4°25'N-4°10'S, 108°50'-119°00'E; collected in by K. Yoshiba (1964, p. 25). FAS; C:Sab-14. 1851 by J. H. Croockewit (see Medway, 1977, Bukitlawang; Sumatra, INDONESIA; ca. 3°30'N, p. 3); rmnh, 1 (skull only). Collected date un- 98°06'E; observed Nov. 1971-Jan. 1973 by C. known by unknown collector; zmb, 1 (skull only). M. Crockett and W. L. Wilson (1980, p. 156). FAS; not mapped. FAS; B:S-20.

Borneo: Kalimantan, southwest, INDONESIA; Bukitlawang, Area I, Bohorok district; Sumatra, o 4°25'N^l 10'S, 108°50'-119°00'E; collected in INDONESIA; ca. 3°34'N, 98°07'E; 2 troops ob- 1836 by S. Muller (see Medway, 1977, p. 3); served 11 Nov. 1980-18 Apr. 1982 by K. J. rmnh, 3 (skins only, skulls inside). FAS; not Gurmaya (1989, p. 66). FAS; B:S-20. mapped. Bukitlawang, Area II, Bohorok district; Sumatra,

134 FIELDIANA: ZOOLOGY INDONESIA; ca. 3°33'N, 98°07'E; observed 1 1 ed 2 Dec. 1908 by [H.] Schoede; zmb, 1. FAS; Nov. 1980-18 Apr. 1982 by K. J. Gurmaya not mapped. (1989, p. 66). FAS; B:S-20. Bunoa, Pulo. See Benua, Pulau. Bukitlawang, ca. 35 km SW; Sumatra, INDO- Buntok, Sungai Barito, 20 ft (= 6 m); Borneo: Ka- NESIA; ca. 3°15'N, 97°55'E; reported before limantan, INDONESIA; 1°42°S, 1 1 4°48°E; col- 1981 by M. Borner (Crockett & Wilson, 1980, lected 6 Oct. 1909 by G. C. Shortridge; bm(nh), p. 156). FAS;B:S-15. l.FAS;C:K-31. Bukitlawang, ca. 65 km S; Sumatra, INDONE- Burau, Pulau, MALAYSIA: WEST MALAYSIA; SIA; ca. 2°55'N, 98°07'E; reported Nov. 1971- 6°21'N, 99°41'E; collected 14 Dec. 1916 by H. Jan. 1973 by local residents (Crockett & Wilson, C. Robinson (see H. C. Robinson & Kloss, 1910, 1980, p. 156). FAS;B:S-25. p. 664); bm(nh), 1. FAS; B:SM-1. Bukit Timah Nature Reserve; Singapore Island; Buraw. See Burau, Pulau. ca. observed SINGAPORE; 1°21'N, 103°48'E; [7BURMA]; ?10°00'-21°00'N, 92°00'-98°30'E; before 1901 S. S. Flower Re- by (1900, p. 316). collected date unknown by unknown collector; in 1941 ported present (Burkhill, 1961, p. 162). bnhs, 1 (skin only). Subspecies uncertain; not Observed 25 Jul. 1986-6 Nov. 1987 by P. W. mapped. Lucas (Lucas & Corlett, 1991, p. 202). FAS; Bur ni Bebuli, north of Danau Laut Tawar, ca. B:SCS-7. 2000 m; Sumatra, INDONESIA; ca. 4°40'N, Bukittinggi; Sumatra, INDONESIA; 0°19'S, 96°52'E; observed 2 Nov. 1905 by W. Volz 100°22'E; collected date unknown by unknown (1912, pp. 88, 369). FAS; B:S-9. collector; zmb, 1 (skull only). FAS; B:S-45. Busang, Sungai, left bank, 6 km above mouth; Bukittinggi, ca. 20 km S; Sumatra, INDONESIA; Borneo: Kalimantan, INDONESIA; ca. 0°06'S, ca. 0°23'S, 100°25'E; observed Nov. 1971-Jan. 1 1 3°57'E; observed 2-4 Sep. 1 986 by D. J. Oliv- 1 973 by C. M. Crockett and W. L. Wilson (1980, ers and K. M. Burton ([1991], p. 143). FAS; p. 156). FAS; B:S-45. CK-19. Bukittinggi vicinity; Sumatra, INDONESIA; ca. Busang, Sungai, right bank, 1 km above mouth; 0°19'S, 100°22'E; observed Nov. 1971-Jan. 1973 Borneo: Kalimantan, INDONESIA; ca. 0°08'S, by C. M. Crockett and W. L. Wilson (1980, p. 1 1 3°58'E; observed 2-4 Sep. 1 986 by D. J. Oliv- 156). FAS; B:S-45. ers and K. M. Burton ([1991], p. 143). FAS; Bulan, Pulau, Kepulauan Riau, INDONESIA; CK-19. 0°55'-l°02'N, 103°51'-103°58'E; collected 5-10 Butang, Ko, THAILAND; 6°32'N, 99°12°E; col- Apr. 1924 by F. N. Chasen (1924b, p. 59); zrc, lected 4 Apr. 1911 by H. C. Robinson; bm(nh), 2. FAS; B:SCS-15. 1. FAS; A:T-58. Bulan, Pulau, [south], Kepulauan Riau; INDO- -Danau Bratan Pulau NESIA; ca. 0°57'N, 103°56'E; collected 22 Mar. Buyan, Danau, region; Bali, Lesser Sunda Islands, INDONESIA; ca. 8°15'S, 1907 by W. L. Abbott (field catalog; cf. Lyon, 1 15°09'E; observed ca. 1993 B. P. 1909, p. 479); usnm, 1. FAS; B:SCS-15. by Wheatley, A. and D. K. Putra Bulang, Pulau. See Bulan, Pulau. Fuentes, Harya (1993, p. 1). CLS-3. Boloh. See Kampong Sungai Buloh. FAS; Bundu Tuhan. See Kampong Bundu Tuhan. Cabang Panti Research Station, Gunung Palung Bunga Buah, 1000 m; MALAYSIA: WEST MA- National Park; Borneo: Kalimantan, INDO- LAYSIA; 3°25'N, 101°45'E; reported as prob- NESIA; 1°13'S, 110°08'E; observed Jan.-Jun. 1988 C. H. Cannon and M. ably present Jul. 1 978-Jun. 1 98 1 by C. W. Marsh by Leighton (1994, C:K-1. and W. L. Wilson (1981, p. 232). FAS; B:WM- p. 509). FAS; 19. Cagar Alam Lembah Anai; Sumatra, INDONE- not observed Feb. 1990 M. Bungara, Area III, Bohorok district; Sumatra, IN- SIA; located; by DONESIA; ca. 3°32'N, 98°12'E; 3 troops ob- Bismark (1992, p. 11). FAS; not mapped. served 10 Aug. 1983-18 Apr. 1984 by K. J. Camara, Bantam region; Java, INDONESIA; collected 27 Jul. 1932 P. Gurmaya (1989, p. 66). FAS; B:S-20. 6°36'S, 105°37'E; by F. 1. Bunguran Island. See Binjai, Sungai; Sinubing; and Franck; mzb, FAS; B:J-13. Ulu, Sungai. CAMBODIA; 10°24'-14°41'N, 102°20'-107°38'E; "Bungur-Buikt"; Sumatra, INDONESIA; not lo- collected [1858-1 86 1] by H. Mouhot (1 864, map cated, 5°38'N-5°57'S, 95°12'-106°05'E; collect- at end of vol. 2; cf. Schlegel, 1876, p. 104); rmnh,

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 135 1 (skin only). Collected 1 Dec. 1960 by unknown LAND; ca. 13°40'N, 100°33'E; observed Apr.- collector; mnhn, 1. FAS; A:C-1 through C-3. May 1912 and 1914-1915 by N. Gyldenstolpe "Camboja" or "Cochinchine"; CAMBODIA or (1914, p. 3; 1917b, p. 6). FAS; A:T-30. VIETNAM; 8°33'-14°41'N, 102°21'-107°39'E; Cheraka Klang, Bukit. See Cherakah, Bukit. collected ca. 1 878 by M. Pierre (cf. P. H. Napier, Cherakah, Bukit; MALAYSIA: WEST MALAY- 1981, p. 19; undocumented restriction of local- SIA; 3°14'N, 101°23'E; collected 16-17 Nov. ity to "Tay Ninh"); bm(nh), 2 (skins only). FAS; 1910 by museum collector; bm(nh), 1; zrc, 1 not mapped. FAS; B:WM-18. Cameron Highlands, 1 500 m; MALAYSIA: WEST Cheribon. See Cirebon. MALAYSIA; 4°28'N, 101°22'E; reported as Cherok Paloh. See Kampong Cherok Paloh. probably present Jul. 1978-Jun. 1981 by C. W. Chiang Mai, northwest; THAILAND; ca. 19°15'N, Marsh and W. L. Wilson (1981, p. 232). FAS; 98°45'E; unconfirmed report, source unspeci- B:WM-7. fied, cited by Varavudhi et al. (1992, p. 338; cf. '- Camorta Island, Nicobar Islands, INDIA; 8°0 1 Fooden, 1971, p. 28). Not mapped. 8°15'N, 93°28'-93°34'E; primates reported ab- Chiew Lam Reservoir; THAILAND; ca. 9°05'N, sent before 1903 by G. S. Miller, Jr. (1902b, p. 98°40'E; rescued from flooding 1986-1987 by 792; cf. Kloss, 1903a, p. 1 14). A:k. Royal Forest Deparment (Nakhasathien, 1989, Camp II, Juara side; Pulau Tioman, MALAYSIA: p. 150). AUR/FAS/MUL; A:T-47. WEST MALAYSIA; ca. 2°48'N, 104°12'E; ob- Chittagong Hill tracts; BANGLADESH; ca. served Mar.-Apr. 1 962 by Lord Medway (1966, 22°30'N, 92°20'E; improbable report (M. A. R. p. 16). FAS; B:SCS-3. Khan, 1981, p. 13; 1985, p. 30). Not mapped. Campbell Bay vicinity; , IN- Christmas Island, AUSTRALIA; ca. 10°23'- DIA; ca. 6°59'N, 93°55'E; observed 27 Feb.-8 10°34'S, 105°34'-105°46'E; primates reported Mar. 1986 by R. Grubh (see Abdulali, 1967, p. absent 1897-1898 by C. W. Andrews (1900, p. 143). UMB; A:N-4. 22). Not mapped. Candiroto, 600 m; Java, INDONESIA; 7°10'S, Chumphon, Khlong, mouth; THAILAND; 110°03'E; collected 20 Jun. 1929 by H. J. V. 10°22'N, 99°10'E; collected 7 Jul. 1917 by W. Sody; rmnh, 1. FAS; GJ-25. J. F. Williamson and M. A. Smith (see Kloss, Cape Patani. See Pho, Laem. 1917, p. 289); zrc, 2. AUR/FAS/MUL; A:T- Cape Rachado Forest Reserve. See Tanjong Tuan, 44. Keramat. Cihara, Bantam region; Java, INDONESIA; 6°52'S, Car Nicobar Island, INDIA; 9°07'-9°16°N, 92°44'- 106°06'E; collected 1 1 Nov. 1909 by O. Bryant 92°51'E; primates reported absent before 1903 (field catalog); usnm, 2. FAS; B:J-1 1. by G. S. Miller, Jr. (1902b, p. 792; cf. Kloss, Cikarang forest; Java, INDONESIA; 6°15'S, 1903a, p. 114). A:i. 107°09'E; collected 9 May 1858 by J. Zelebor Catur, Gunung; Pulau Bali, Lesser Sunda Islands, ([1869], p. 7); museum unknown, 1 (not seen). INDONESIA; ca. 8°14'S, 1 15°1 l'E; observed ca. FAS; B:J-19. 1993 by B. P. Wheatley, A. Fuentes, and D. K. Cikujang, Bantam region; Java, INDONESIA; Harya Putra (1993, p. 1). FAS; CLS-3. 6°41'S, 107°03'E; collected 8 Aug. 1932 by P. Celebes. See Sulawesi, Pulau. F. Franck; mzb, 2. FAS; B:J-20. Chang, Ko, THAILAND; 1 1°57'-12°09'N, Cilacap, sea level; Java, INDONESIA; 7°44'S, 102°16'-102°28'E; collected 9-11 Dec. 1914 by 109°00'E; collected 19 Oct. 1907 by G. C. Shor- C. B. Kloss (1916b, p. 28); bm(nh), 1; usnm, 1; tridge (see Thomas & Wroughton, 1 909a, p. 373); zrc, 1. FAS; A:T-35. bm(nh), 2. FAS; B:J-1. Changi; Singapore Island, SINGAPORE; 1°23'N, Cirebon; Java, INDONESIA; 6°44'S, 108°34'E; 103°59'E; collected 22 Jul. 1908 by H. C. Rob- collected 18 Dec. 1927 by K. Fritsche; nms, 1 inson (in Thomas & Wroughton, 1909b, p. 102) (skull only). Collected before 1948, possibly by and E. Seimund; bm(nh), 1. FAS; B:SCS-7. J. J. Menden; rcs(om), 6 (skulls only). FAS; B:J- Changkat Budiman. See Keroh Forest Reserve. 23. Changkat Mentri; MALAYSIA: WEST MALAY- Cirebon, 600 m; Java, INDONESIA; ca. 6°44'S, SIA; 3°44'N, 101°15'E; collected 19-23 Sep. 1918 108°34'E; collected 18 Feb. 1933 by J. J. Men- by C. B. Kloss (see P. H. Napier, 1981, p. 14); den; amnh, 2. FAS; B:J-23.

bm(nh), 2. FAS; B:WM-18. Ciremay, Gunung, 600 m (1 specimen) and 800 Chao Phraya, Mae Nam, below Bangkok; THAI- m (6 specimens); Java, INDONESIA; 6°54'S,

136 FIELDIANA: ZOOLOGY 108°24'E; collected 11 Mar.-8 Apr. 1933 by J. Deli. See Medan. o J. Menden; amnh, 7. FAS; B:J-23. Deli, Pulau, INDONESIA; 7°00'-7 01'S, 105°31'- . See Kalipucang. 105°34'E; introduced in 1987 by C. V. Primates Ciwangi, 4000 ft (= 1200 m); Java, INDONESIA; (Indonesia) (advertising leaflet, ca. 1993). Sub- 7°04'S, 107°02'E; collected 25 Sep.-l Oct. 1907 species uncertain; not mapped. by G. C. Shortridge (see Thomas & Wroughton, Deli, Sungai, between Belawan and Labuhandeli, 1909a, p. 373); bm(nh), 2. 1 FAS; B:J-8. sea level; Sumatra, INDONESIA; 3°46'N, Cochin China [region]; VIETNAM; 8°33'-12°17'N, 98°41'E; collected 17 Mar. 1939 by F. Ulmer 104°30'-107°39'E; collected before 1882 by M. (in Miller, 1 942, p. 1 27; cf. Schauensee & Rip- Boucard; bm(nh), 1. Collected before 1883 by ley, 1940a, p. 311); ansp, 4 (including 2 fetuses

M. Germain; mnhn, 1 . FAS; A:V-3 through V-5. in alcohol) FAS; D:S-21. Cocos Islands. See Tikus, Pulu. DeLisle Island. See Phayam, Ko. Condor, P. See Con Son. Deli Terbanjawan. See Terbanjawan. Condore, P. See Con Son. Depok; Java, INDONESIA; 6°24'S, 106°50'E; col- Con Son, VIETNAM; 8°40'-8°47'N, 106°32'- lected 17 Jun. and 6 Jul. 1909 by O. Bryant and 106°39'E; observed 20-28 Jan. 1780 by J. King W. Palmer (see field catalog, usnm); mcz, 2. FAS; (1784, p. 462). Observed 22-23 Aug. 1822 by B:J-17.

J. Crawfurd (1 828, p. 199). Collected before 1 893 Depok, small forest near; Java, INDONESIA; ca. by M. Germain, mnhn, 2 (including 1 skin only, 6°24'S, 106°50'E; observed 1932-1957 by A. 1 skull only). Collected 19-23 Sep. 1919 by M. Hoogerwerf (1970, p. 408). FAS; B:J-17. A. Smith; bm(nh), 3. Collected 6-18 Nov. 1920 Desa Poetjang, Gunung Agung; Pulau Bali, Lesser by W. J. F. Williamson and C. Hose (see KJoss, Sunda Islands, INDONESIA; ca. 8°21'S, 1926, p. 357; Weitzel et al., 1988, p. 110); 115°30'E; collected ca. 1930 by H. J. V. Sody

bm(nh), 1; zrc, 3 (including 1 skin only). CON; (1933, p. 93); rmnh, 2. FAS; CLS-3. A:V-7. Dewhurst Bay area; Borneo, MALAYSIA: SA- Con Son, west shore, VIETNAM; ca. 8°43'N, BAH; ca. 5°35'N, 1 18°35'E; observed May-Jun. 106°34'E; observed 20-25 Mar. 1969 by P. F. 1950 by D. D. Davis (1962, p. 57). FAS; C:Sab- D. Van Peenen, M. L. Cunningham, and J. F. 20.

Duncan (1970, p. 421). CON; A:V-7. Dindding I.; Borneo, MALAYSIA: SARAWAK; Cox's Bazar Forest Division. See Bilasodia, Bi- not located, 0°43'-4°58'N, 109°34'-115°38'E; mirdia, Ghorardia, Ochodia, and Rukumodia. collected before 1906 by H. C. Robinson (see Dalam, Lhok; Pulau Simeulue, INDONESIA; P. H. Napier, 1981, p. 14); bm(nh), 1 (skin only). 2°40'N, 96°08'E; collected 18-26 Nov. 1901 by FAS; not mapped. W. L. Abbott (see Miller, 1903a, p. 437); usnm, Dindings. See Hantu, Tanjong, Dindings. 6. FUS; B:IO-2. Dirk de Vries Bay. See Pangandaran, Teluk Parigi. Damansara; MALAYSIA: WEST MALAYSIA; Desertion Creek, Elephant Point, bank of Irra- 3°08'N, 101°38'E; laboratory animals obtained waddy River, near Rangoon; BURMA; ca. before 1982 by unknown collector (Nordin, 16°29'N, 96°20'E; collected before 27 Jan. 1876 1981, p. 164). FAS; B:WM-19. by J. Armstrong (see J. Anderson, 1881, p. 63; Dampit. See Wonokojo (?= Wonokerto), Dampit Khajuria,[1955],p. 109).zsi, 1 (not seen). AUR; district, southern region. A:Bu-6. Danau, Rawa; Java, INDONESIA; 6°09'S, Djambajan, Sungai. See Jembayan, Sungai. 105°59'E; blood samples taken in 1979 by Y. Djapura. See Japura. Kawamoto and Tb. M. Ischak (1981, p. 238). Djasinga. See Jasinga. FAS;B:J-15. Djeboes. See Jebus. Darvel Bay; Borneo, MALAYSIA: SABAH; Djembrana. See Jembrana. 4°50'N, 1 18°30'E; collected 21 Apr. 1909 by Dr. Dolok Tinggi Radja Reserve; Sumatra, INDO- Pagel; zmb, 1. FAS; C:Sab-22. NESIA; ca. 3°07'N, 98°45'E; reported present Dasun Tua. See Dusun Tua. before 1972 by IUCN (1971, p. 276). FAS; B:S- Datu, Pulau. See Datuk, Pulau. 27. Datuk, Pulau, INDONESIA; 0°09'-0°10'N, Domel Island. See Letsok-aw Kyun. 108°38'-108°39'E; monkeys reported absent 2- Dompu, 66 m; Pulau Sumbawa, Lesser Sunda Is- 4 May 1907 by W. L. Abbott (in Lyon, 1911, lands, INDONESIA; 8°32'S, 1 18°28'E; collected p. 59). B:m. 22 Dec. 1909 and in 1910 by J. Elbert (1912,

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 137 p. 98); nms, 2 (including 1 skull only). Collected 102°05'-102°24'E; primates reported absent be- 28 May 1927 by Sunda-Expedition Rensch (see fore 1928 by C. B. Kloss ([1928], p. 802; cf. B. Rensch, 1930, p. 93; Mertens, 1930, p. 135; Lyon, 1916, p. 460). Reported present Nov. I. Rensch, 1934, p. 226); nms, 2 (skulls only; 1971-Jan. 1973 by unspecified informants

including 1 not examined, measurements from (Crockett & Wilson, 1980, p. 156). Pending fur- Mertens, 1 936, p. 3 1 9, and G. H. Albrecht, pers. ther information, Kloss's report is accepted as comm., Oct. 1991). FAS; CLS-11. valid. B:j. Don Poo Tao; THAILAND; 15°45'N, 104°22'E; Entawa, Tanjong, Sungai Samarahan; Borneo, observed 16-17 Jul. 1989 by N. Aggimarangsee MALAYSIA: SARAWAK; 1°17'N, 110°29°E; (1992, pp. 109, 120; pers. comm., Oct. 1993). collected 21 Nov. 1919 by H. C. Robinson; Subspecies uncertain; A:T-16. bm(nh), 1. FAS; C:Sar-5. Dulit, Bukit, 4000 ft (= 1200 m); Borneo, MA- E. Siam. See "Siam." LAYSIA: SARAWAK; 3°21'N, 114°11'E; col- False Durian. See Sanglang-besar, Pulau. lected Sep. 1891 and Mar. 1894 by C. Hose Fatuboi; Pulau Timor, Lesser Sunda Islands, IN-

(1893, p. 8; altitude cited as 5000 ft); smk, 1 DONESIA; ca. 8°50'S, 126°20'E; observed 28 (skin only, skull inside); zmb, 1 (skull only). FAS; Apr.-3 May 1883 by H. O. Forbes (1885, p. C:Sar-13. 471). FAS;C:LS-29. Durai. See Durian, Pulau. Flores, Pulau, Lesser Sunda Islands, INDONE- Durian, False. See Sanglang-besar, Pulau. SIA; 8°14'-8°58'S, 119°48'-123°02'E; collected Durian, Pulau, Kepulauan Riau, INDONESIA; 1854-1862 by A. R. Wallace; bm(nh), 1. FAS; 0°42'-0°45'N, 103°42'-103°45'E; reported pres- C:LS- 13 through LS- 19. ent Jun.-Aug. 1903 by W. L. Abbott (see Miller, Fort de Kock. See Bukittinggi. 1906c, p. 279). Collected 18 Jun. 1923 by P. F. Gaik Liew Estate, Damansara; MALAYSIA: Franck(see Dammerman, 1926b, pp. 282, 302); WEST MALAYSIA; 3°07'N, 101°37'E; collect- mzb, 2. FAS; B:SCS-16. ed 13 Feb. 1921 by "T. H. S."; bm(nh), 1. FAS; Durian-kecil, Pulau, Kepulauan Riau, INDO- B:WM-19. NESIA; 0°43'-0°45'N, 103°39'-103°42'E; re- Galang, Pulau, Kepulauan Riau, INDONESIA; ported present 6-9 Jul. 1903 by W. L. Abbott 0°42'-0°48'N, 1 04° 1 0'-l 04° 1 8 'E; collected 1 Jan. (see Miller, 1906c, p. 280). FAS; B:SCS-16. 1925 by F. N. Chasen; zrc, 4. FAS; B:SCS-1 1. Dusun, Sungai, 10 m; MALAYSIA: WEST MA- Galang-baru, Pulau. See Nguwal, Pulau. LAYSIA; 3°40'N, 101°20'E; observed Jul. 1978- Gantang. See Kantang. Jun. 1981 by C. W. Marsh and W. L. Wilson Garau, Kampong Kiau region, Mount Kinabalu, (1981, p. 232). FAS; B:WM-18. ca. 3000 ft (= 900 m); Borneo, MALAYSIA: Dusun Tua; MALAYSIA: WEST MALAYSIA; SABAH; ca. 6°02'N, 116°31'E; collected 18-19 3°08'N, 101°50'E; collected 3 Oct. 1906 by E. Aug. 1937 by J. A. Griswold, Jr. (1939a, p. 410); Seimund (see Thomas & Wroughton, 1 909b, p. mcz, 4. FAS; C:Sab-7. 101); zrc, 1. FAS; B:WM-19. Gasip, Sungai, ca. 1 km above mouth; Sumatra, Eagle's Nest Trail, Kowloon Reservoir Area; INDONESIA; ca. 0°32'N, 101°44'E; observed UNITED KINGDOM: HONG KONG; ca. Nov. 1971-Jan. 1973 by C. M. Crockett and W. 22°21'N, 114°09'E; introduced population, ob- L. Wilson (1980, p. 156). FAS; B:S-42. served Jan.-Feb. 1987 by C. H. Southwick and Gasip, Sungai, ca. 20 km above mouth; Sumatra, D. Manry (1987, p. 48). Subspecies uncertain; INDONESIA; ca. 0°37'N, 101°43'E; observed not mapped. Nov. 1971-Jan. 1973 by C. M. Crockett and W. Elephant Point. See Desertion Creek, Elephant L. Wilson (1980, p. 156). FAS; B:S-42. Point. Gedangan, Semarang district, 65 m; Java, IN- Empang area. See Ampang area. DONESIA; 7°11'S, 110°41'E; collected Jan. Endau, Sungai, vicinity; MALAYSIA: WEST 1929-Oct. 1931 by H. J. V. Sody; rmnh, 7 (in- MALAYSIA; ca. 2°40'N, 103°38'E; observed 16 cluding 4 skulls only). Collected 6 Oct. 1931 by Oct.-2 Nov. 1892 by H. W. Lake and H. J. F. A. T. H. Verbeek; rmnh, 1 (skin only). FAS; Kelsall (see Kelsall, 1894b, p. 16). FAS; B:WM- CJ-29. 22. Gede, Gunung, Pangrango; Java, INDONESIA; Engano Id. See Enggano, Pulau. 6°47'S, 106°59'E; observed Jan. 1991 byM.Bis- Enggano, Pulau, INDONESIA; 5°17'-5°32'S, mark(1992, pp. 11, 13). FAS; B:J-18.

138 FIELDIANA: ZOOLOGY Ghirbi. See Ban Nong Kok. SIA: SARAWAK; ca. 4°00'N, 114°55'E; re- Ghorardia, Naf River; BANGLADESH; ca. ported as food of local residents Jan. 1978 by 21°05'N,92°12'E; observed Sep. 1982-Feb. 1983 D. Labang and Lord Medway (1979, p. 56). FAS; by M. A. R. Khan and M. A. Wahab (1983, p. C:Sar-21. 104; cf. M. A. R. Khan, 1981, p. 13; 1985, p. Gunung Halimun Reserve; Java, INDONESIA; 30). AUR; A:Ba-l. ca. 6°15'S, 106°30'E; observed Jul. 1989 by K. Gili Bodo. See Sababi, Pulau. M. Kool (1992, p. 32). FAS; B:J-16. Gili Lawa Darat. See Lawadarat, Gili. Gunung Leuser Reserve; Sumatra, INDONESIA; Gili Lawa Laut. See Lawalaut, Gili. ca. 3°45'N, 97°H'E; observed Apr.-Aug. 1970 Pulau Lesser Sunda IN- Gilimanuk; Bali, Islands, by F. Kurt (1973, p. 64). FAS; B:S-12. collected 8-10 DONESIA; 8°10'S, 114°26'E; Gunung Palung Nature Reserve; Borneo: Kali- Apr. 1938 by V. von Plessen; amnh, 8. FAS; mantan, INDONESIA; ca. 1°13'S, 1 10°08'E; re- CLS-1. ported present before 1 983 by G. Davies ([ 1 983], Gili Mota. See Gili. Motang, p. 148). FAS;C:K-1. Gilla, Pulo. See Jela, Pulau. Hainggyi Kyun, BURMA; 1 5°98'-l 6°0 1 'N, 94°1 T- Ginggo, Teluk; Pulau Rinca, Lesser Sunda Islands, 94°22'E; reported present before 1880 by J. An- INDONESIA; ca. 8°4 1 'S, 1 1 9°39'E; observed 26 derson (1879, p. 76). AUR; A:Bu-4. May-6 Jul. 1953 by A. Hoogerwerf (see Auffen- Halimun, Gunung. See Gunung Halimun Reserve. berg, 1981, p. 242). FAS; QLS-13. Handeuleum, Pulau, INDONESIA; 6°45'S, Gitgit, 530 m; Pulau Bali, Lesser Sunda Islands, 105°25'E; 3 individuals captured before 1974 INDONESIA; 8°1 l'S, 1 15°08'E; collected 1 Aug. by W. Angst (1973, p. 627). FAS; B:J-14. 1927 by Sunda-Expedition Rensch (see B. 3 km N of 50 Borneo: Ka- Rensch, 1930, p. 210; Mertens, 1930, p. 145; I. Hantakan, Pagat, m; limantan, col- Rensch, 1934, p. 227); nms, 1. FAS; CLS-3. INDONESIA; 2°38'S, 115°27'E; lected 2 2 Gn. Telapa Burok. See Telapak Burok, Gunong. Feb. 1971 by NAMRU Djakarta De- tachment et Goenoengsetan-Meloewak, 325-520 m; Sumatra, (see Van Peenen al., 1974, p. 391); INDONESIA; 3°45'N, 97°40'E; collected 21 Jan. usnm, 1. FAS;C:K-29. 1937 by A. Hoogerwerf (1941, p. 5; cf. Chasen, Hantu, Tanjong, Dindings; MALAYSIA: WEST 1940b, p. 485); mzb, 1. FAS; B:S-13. MALAYSIA; 4°19'N, 100°34'E; collected 19 Jul. Goson Djerong, near, Sungai Makaham, south 1918 by unknown collector; zrc, 1. FAS; bank; Borneo: Kalimantan, INDONESIA; ca. B:WM-6. 0°29'S, 117°02'E; collected 15 Jun. 1912 by H. Harileko, Batang, ca. 12 km above mouth; Su- C. Raven (see Deignan, [1960], p. 267); usnm, matra, INDONESIA; ca. 2°52'S, 104°01'E; ob- 2 (including 1 skull only). FAS; CK-49. served Nov. 1971-Jan. 1973 by C. M. Crockett Grabi. See Ban Nong Kok. and W. L. Wilson (1980, p. 156). FAS; B:S-62. Great Natuna Island. See Natuna Besar, Pulau. Harileko, Batang, ca. 24 km above mouth; Su- Great Nicobar Island, INDIA; 6°45'-7°14'N, matra, INDONESIA; ca. 2°48'S, 103°57'E; ob- 93°38'-93°57'E; captive obtained Mar. 1858 by served Nov. 1971-Jan. 1973 by C. M. Crockett J. Zelebor ([1869], p. 7; cf. Fitzinger, 1861, p. and W. L. Wilson (1980, p. 156). FAS; B:S-62. 389). Collected 8-12 Mar. 1901 by W. L. Ab- Harileko, Batang, ca. 36 km above mouth; Su- bott; usnm, 2. Observed Apr.-May 1975 by P. matra, INDONESIA; ca. 2°45'S, 103°52'E; ob- K. Das and D. K. Ghosal (1977, p. 265). UMB; served Nov. 1971-Jan. 1973 by C. M. Crockett A:N-3, N-4. and W. L. Wilson (1980, p. 156). FAS; B:S-62. Great Redang Island. See Redang, Pulau. Harileko, Batang, ca. 48 km above mouth; Su- Great Tenasserim River, mouth; BURMA; ca. matra, INDONESIA; ca. 2°42'S, 103°47'E; ob- 12°24'N, 98°37'E; observed before 1852 by F. served Nov. 1971-Jan. 1973 by C. M. Crockett Mason (1851, p. 220). AUR; A:Bu-18. and W. L. Wilson (1980, p. 156). FAS; B:S-62. Gulf of Siam, coast. See Si Racha vicinity. Harileko, Batang, ca. 60 km above mouth; Su- Gumpang, near; Sumatra, INDONESIA; ca. matra, INDONESIA; ca. 2°38'S, 103°42'E; ob- 3°5 1 'N, 97°33'E; blood samples taken Nov.-Dec. served Nov. 1971-Jan. 1973 by C. M. Crockett 1986 by J. R. de Ruiter (1993, p. 91). FAS; B:S- and W. L. Wilson (1980, p. 156). FAS; B:S-62. 13. Harileko, Batang, ca. 72 km above mouth; Su- Gunong Mulu National Park; Borneo, MALAY- matra, INDONESIA; ca. 2°34'S, 103°37'E; ob-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 139 served Nov. 1971-Jan. 1973 by C. M. Crockett Hutan Lindung Lintau Buo; Sumatra, INDO- and W. L. Wilson (1980, p. 156). FAS; B:S-62. NESIA; not located; observed Aug. 1990 by M. Harileko, Batang, ca. 84 km above mouth; Su- Bismark (1992, p. 11). FAS; not mapped. matra, INDONESIA; ca. 2°31'S, 103°32'E; ob- "I. Lendung"; Sumatra, INDONESIA; not locat- served Nov. 1971-Jan. 1973 by C. M. Crockett ed, 5°38'N-5°57'S, 95°12'-106°05'E; collected and W. L. Wilson (1980, p. 156). FAS; B:S-62. data unknown by C. Bruegel; zsbs, 3 (skins only). Harileko, Batang, ca. 96 km above mouth; Su- FAS; not mapped. ca. ob- matra, INDONESIA; 2°28'S, 103°27'E; Indau River. See Endau, Sungai, vicinity. served Nov. 1971-Jan. 1973 C. M. Crockett by Indragiri, Sungai; Sumatra, INDONESIA; ca. and W. L. Wilson B:S-62. (1980, p. 156). FAS; 0°22'S, 103°26'E; collected 21 Sep. 1901 by W. Hat Chao Mai National Park; ca. THAILAND; L. Abbott (see Miller, 1902a, p. 143); usnm, 1. 7°20'N, 99°25'E; reported present 17-21 Dec. FAS; B:S-58. 1987 by park employees (Boonratana, 1988, p. Indragiri district; Sumatra, INDONESIA; 0°10'- FAS; A:T-56. 76). 0°55'S, 101°50'-103°30'E; collected 6 Dec. 1898 Hat Noppharat Thara-Mu Ko Phi Phi National by G. Schneider (1905, p. 72); zmuz, 1 (skull Park; THAILAND; ca. 8°02'N, 98°47'E; re- only). Collected in 1 905 by H. Kummer; NHMBa, ported present 9-12 Dec. 1 987 by local residents 7 (skulls only). FAS; B:S-56. (Boonratana, 1988, p. 76). FAS; A:T-60. Indragiri (Djapura). See Japura. Haungdarau. See Haungtharaw. Indramayu;7ava, INDONESIA; 6°20'S, 108°19'E; Haungtharaw; BURMA; 16°30'N, 98°13'E; col- collected 30 Jul. 1930 and 29 Mar. 1931 by J. lected 10 Dec. 1880 by Mr. Limborg and J. J. Menden; mzb, 4 (including 2 skins only). FAS; Anderson; zsi, 4 (including 3 skins only, skulls B:J-21. inside). AUR; A:Bu-10. Indramojoe. See Haut Padang. See Padang highlands. Indramayu. northwest of Man- Henry Larence Island, Andaman Islands, INDIA; Irrawaddy River, right bank, 12°05'-12°13'N, 93°03'-93°07'E; primates re- dalay; BURMA; ca. 22°05'N, 96°00'E; captive obtained before 1880 ported absent before 1903 by G. S. Miller, Jr. (presumably introduced) Dr. Marfels J. Anderson, 1879, p. 74). (1902b, p. 792; cf. Kloss, [1928], p. 802; Cha- by (see uncertain; not turvedi, 1980, p. 134). A:c. Subspecies mapped. Hilisimaetano; Pulau Nias, INDONESIA; 0°38'N, Isle de France. See Mauritius Island. 97°44'E; collected 5-6 Jun. 1939 by F. Ulmer Jalor. See Kampong Biserat. (in Miller, 1942, p. 129; cf. Schauensee & Rip- Jambi, ca. 60 km NNW; Sumatra, INDONESIA; ley, 1940b, p. 399); ansp, 3. FAS; B:IO-8. ca. 1°05'S, 103°31'E; reported before 1981 by Ho Chi Minn City; VIETNAM; 10°45'N, 106°40'E; M. Borner (Crockett & Wilson, 1980, p. 156). collected in 1929 by T. Roosevelt (see Osgood, FAS; B:S-59. 1932, p. 208); fmnh, 1. Collected date unknown Jambi, ca. 90 km ENE; Sumatra, INDONESIA; by H. Zeltmann; nms, 2 (skins only). FAS; ca. 1°16'S, 104°21'E; reported before 1981 by A:V-4. M. Borner (Crockett & Wilson, 1980, p. 156). Ho Chi Minh City, Botanical Gardens; VIET- FAS; B:S-60. NAM; 10°45'N, 106°40'E; collected Oct. 1926- Jambu, tidal creeks near. See Yaring, tidal creeks Sep. 1927 by J. Delacour and W. P. Lowe (see near. Thomas, 1928, p. 832); bm(nh), 1 (skin only); Japura; Sumatra, INDONESIA; ca. 0°19'S, mnhn, 2 (including 1 skin only, 1 skull only). 102°21'E; collected 1897-1899 by G. Schneider FAS; A:V-4. (1905, p. 72); museum and number of speci- Ho Chi Minh City, Zoological Garden; VIET- mens unknown (possibly zmuz 11668, skull NAM; 10°45'N, 106°40'E; collected 26 Jul. 1926 only). FAS; B:S-55. and 1 1 Oct. 1926 by unknown collectors; mnhn, Jarak, Pulau, MALAYSIA: WEST MALAYSIA; 2 (including 1 skull only). FAS; A:V-4. 3°59'N, 100°06'E; primates reported absent Hong Kong. See Eagle's Nest Trail, Kam Shan 1950-1960 by J. L. Harrison and J. R. Hen- Entrance, Kowloon Reservoir Area, and Lower drickson (1963, p. 548). B:b. Taipo Road. Jasinga, Bogor district; Java, INDONESIA; 6°29'S, Huai Ong Sit. See Ban Phu Toie. 106°27'E; collected 10 Apr. 1929 and 4 Sep. Huaytakaeng. See Wat Huai Takhaeng. 1931 by P. F. Franck; mzb, 2. FAS; B:J-10. Huay Takang. See Wat Huai Takhaeng. Jatibarang; Java, INDONESIA; 6°28'S, 108°17'E;

140 FIELDIANA: ZOOLOGY clinical examination in 1980 by K. Matsubay- 3°27'S, 102°15'E; observed Nov. 1971-Jan. 1973 ashi and D. Sajuthi (1981, p. 48). FAS; B:J-21. by C. M. Crockett and W. L. Wilson (1980, p. Java, INDONESIA; 5°53'-8°47'S, 105°07'- 156). FAS; B:S-68. 114°37'E; collected in 1827 by van Swindern; Jeleket vicinity; Sumatra, INDONESIA; ca. 3°27'S,

nms, 1 . Collected in 1 832 by unknown collector; 102°20'E; observed Nov. 1971-Jan. 1973 by C. NHMBa, 1 (skull only). Collected in 1 836 by Ouv- M. Crockett and W. L. Wilson (1980, p. 156). ermer-Fischer; nms, 2 (skulls only). Two cap- FAS; B:S-68. tives obtained in 1858 by J. Zelebor or von Jemaja, Pulau, INDONESIA; ca. 2°49'-3°03'N, Frauenfeld (see Fitzinger, 1861, pp. 385, 388). 105°41'-105°51'E; observed 16-28 Sep. 1899 by Acquired before 1 859 by C. J. Temminck; rmnh, W. L. Abbott (in Miller, 1900, p. 244; cf. Kloss, 2 (skulls only). Collected 10 Jan. 1866 by Kok; 1903b, p. 74). FAS; B:SCS-28. rmnh, 1 (skin only, skull inside). Collected in Jembayan, Sungai; Borneo: Kalimantan, INDO- 1898 by Bartels; nhrm, 17 (including 16 skulls NESIA; 0°35'S, 117°00'E; collected 6-14 May only, 1 mandible only). Collected date unknown 1914 by H. C. Raven (see Deignan, [1960], p. by Bartels; nhrm, 1 (skull only). Collected be- 269); usnm, 3 (skulls only). FAS; C:K-49. fore 1 1 1627/ 1924 by Tucker; zmuz, (skin, No. Jembrana, Negara district; Pulau Bali, Lesser Sun- skeleton, No. 1 1630). Collected before 1925 by da Islands, INDONESIA; 8°22'S, 1 14°39'E; col- 3 Col- unknown collector; NHMBa, (skins only). lected ca. 1930 by H. J. V. Sody (1933, p. 93); lected 14 L. 1 Jan. 1932 by Heinrath; zmb, (skull rmnh, 2 (including 1 skin not examined). FAS; only). Collected before 1935 by unknown col- C:LS-2. 2 Collected in lector; rmnh, (skulls only). 1936 Jering region. See Yaring region. and 1937 C. 30 24 by Blazer; rmnh, (including Jessleton (= Kota Kinabalu). See Papar; Talibang; skulls only). Collected before 1942 by E. Du- Tuaran. bois; rmnh, 1 (skull Collected date un- only). Jimaja. See Jemaja, Pulau. known by van Aken; rmnh, 1 (skull only). Col- Jockaboemi; Java, INDONESIA; not located, lected date unknown by Hecht; zmb, 1 . Collected 5°93'-8°47'S, 105°12'-1 14°36'E; collected 7 Jun. date unknown by T. B. Wilson; ansp, 1 (skull 1909 by Dr. Biedermann; zmb, 1 (skull only). only). Collected date unknown by unknown col- FAS; not mapped. lector; rmnh, 9 (including 6 skins only [skulls Johore Archipelago. See Pemanggil, Pulau. inside], 3 skulls only); smtd, 1 (skin only). FAS; Jolir dia, Naf River; BANGLADESH; ca. 20°50'N, not mapped. 92°18'E; reported present 1982-1983 by M. A. Java, west, INDONESIA; 5°52'-7°48'S, 105°12'- R. Khan and M. A. Wahab (1983, p. 104; cf. 108°52'E; collected 1818-1826 by C. L. Blume; M. A. R. Khan, 1981, p. 13; 1985, p. 30). AUR; rmnh, 2 (including 1 skin only [skull inside], 1 A:Ba-2. skeleton only). Collected 1 820-1 82 1 by H. Kuhl Joloi River. See Julai, Sungai. and J. C. van Hasselt; rmnh, 1 (skeleton only). Telok; Pulau Tioman, MALAYSIA: WEST Collected 1826-1837 by S. Muller and H. C. Juara, MALAYSIA; 2°48'N, 1 04°1 3'E; collected 1 Jun. Macklot; rmnh 2 (skins only, skulls inside). Col- 1906 by H. C. Robinson; bm(nh), 1. Collected lected in 1863 by P. Diard; rmnh, 5 (including 9-10 Sep. 1907 by museum collector; bm(nh), 1 skin only, 1 skeleton only). Collected date un- 2. Collected 4-22 Jun. and 1-2 Jul. 1915 by H. known by P. Diard; rmnh, 1 (skeleton only). C. Robinson; bm(nh), 2; zrc, 2. FAS; Ba:SCS-3. Collected in 1 870 by unknown collector; rmnh, left 1 km below mouth of 1 (skin only, skull inside). Examined for para- Julai, Sungai, bank, Borneo: INDO- sites before 1970 by D. Weinman and N. S. Sungai Busang; Kalimantan, ca. 1 observed 2-A Wiratmadja (1969, p. 499). FAS; not mapped. NESIA; 0°09'S, 13°59'E; Sep. 1986 D. J. Chivers and K. M. Burton Jebus; Pulau Bangka, INDONESIA; 1°44'S, by ([1991], QK-19. 105°29'E; collected ca. 1935 by H. J. V. Sody p. 143). FAS; (1937, p. 248); rmnh, 1 (skull only). FAS; B:SCS- Julai, Sungai, left bank, 2 km above Muarajuloi; 18. Borneo: Kalimantan, INDONESIA; ca. 0°1 l'S, Jela, Pulau, INDONESIA; 0°57'-0°58'N, 107°29'- 1 14°03'E; observed 30 Aug.-8 Sep. 1986 by D. 107°30'E; observed 3-7 Aug. 1899 by W. L. J. Chivers and K. M. Burton ([1991], p. 143). Abbott (in Miller, 1 900, p. 243; cf. Kloss, 1 903b, FAS; CK-19. p. 60). FAS; B:SCS-26. Julai, Sungai, left bank, 4 km above Muarajuloi; Jeleket, ca. 10 km W; Sumatra, INDONESIA; ca. Borneo: Kalimantan, INDONESIA; ca. 0°10'S,

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 141 1 14°03'E; observed 30 Aug.-8 Sep. 1986 by D. gai Rungan; Borneo: Kalimantan, INDONE- J. Chivers and K. M. Burton ([1991], p. 143). SIA; ca. 2°08'S, 1 1 3°56'E; 2 troops observed Jul. FAS;C:K-19. 1984-May 1986 by K. M. Burton (see Chivers Julai, Sungai, right bank, 1 km above mouth of & Burton, [1991], p. 140). FAS; GK-23. Sungai Busang; Borneo: Kalimantan, INDO- Kahayan, Sungai, left bank, below mouth of Sun- NESIA; ca. 0°1 l'S, 1 13°58'E; observed 1-2 Sep. gai Rungan; Borneo: Kalimantan, INDONE-

1 986 by D. J. Chivers and K. M. Burton ([1 99 1], SIA; ca. 2°15'S, 114°00'E; observed Jul. 1984- p. 143). FAS; C:K-19. May 1986 by K M. Burton (see Chivers & Bur- Julai, Sungai, right bank, 3 km below mouth of ton, [1991], p. 140). FAS; CK-23. Sungai Busang; Borneo: Kalimantan, INDO- Kahayan, Sungai, right bank, 1 20 km above mouth NESIA; ca. 0°08'S, 1 14°00'E; observed 30 Aug.- of Sungai Rungan; Borneo: Kalimantan, IN- 1 Sep. 1986 by D. J. Chivers and K. M. Burton DONESIA; ca. 1°08'S, 113°52'E; observed Jul. ([1991], p. 143). FAS; GK-19. 1984-May 1986 by K. M. Burton (see Chivers Julai, Sungai, right bank, 6 km above Muarajuloi; & Burton, [1991], p. 140). FAS; C:K-20. Borneo: Kalimantan, INDONESIA; ca. 0°09'S, Kahayan, Sungai, right bank, 1 80 km above mouth 1 14°02'E; observed 30 Aug.-8 Sep. 1986 by D. of Sungai Rungan; Borneo: Kalimantan, IN- J. Chivers and K. M. Burton ([1991], p. 143). DONESIA; ca. 0°54'S, 1 13°12'E; observed Jul. FAS; C:K-19. 1984-May 1986 by K. M. Burton (see Chivers Jumpit [?= Jumbit]; Borneo, MALAYSIA: SA- & Burton, [1991], p. 140). FAS; C:K-15. RAWAK; ?1°07'N, 1 1 P26'E; collected date un- Kahayan, Sungai, right bank, below mouth ofSun- known by Rupert; zmb, 1 (skull only). FAS; gai Rungan; Borneo: Kalimantan, INDONE- C:Sar-10. SIA; ca. 2°15'S, 114°00'E; observed Jul. 1984- Juyan, Sungai, right bank, near mouth, Kutai Re- May 1986 by K. M. Burton (see Chivers & Bur- serve; Borneo: Kalimantan, INDONESIA; ca. ton, [1991], p. 140). FAS; CK-23. 0°27'N, 1 17°12'E; observed 13 Oct. 1985 by A. Kajan, Sungai. See Long Peleben. Suzuki (1986, p. 16). FAS; C:K-47. Kalianda, 100 m; Sumatra, INDONESIA; 5°45'S, Kaban, Pulau. See Acheh, Pulau. 105°38'E; collected 5 Aug. 1 934 by J. J. Menden; Kaboerau [?= Keburau]; Borneo: Kalimantan, IN- amnh, 3. FAS; B:S-87. DONESIA; ca. 2°4 1 'N, 1 1 7°07'E; collected 9 Jan. Kaligoea, Gunung Slamet, 1350 m; Java, IN- 1914 by C. Lumholtz (see Gyldenstolpe, 1920, DONESIA; ca. 7°14'S, 109°12'E; collected 18 p. 3); Zoological Museum, Christiana, 1 (not Jan. 1917 by Denin; mzb, 2. FAS; B:J-24. seen). FAS; CK-39. Kalipoetjang. See Kalipucang. Kadan, Kyun, 500 ft (= 150 m), BURMA; 12°18'- Kalipucang, Ci Tanduy, sea level; Java, INDO- 12°42'N, 98°18'-98°29'E; collected 3 Oct. 1923 NESIA; 7°39'S, 108°44'E; collected 10 Mar. 1908 by C. Primrose (see Lindsay, 1926, p. 42); by G. C. Shortridge (see Thomas & Wroughton, bm(nh), 1. AUR; A:Bu-17. 1909a, p. 373); bm(nh), 3. FAS; B:J-2. Kaddamayan River. See Tempasuk, Sungai. Kalulong, [Bukit]; Borneo, MALAYSIA: SARA- Kaget, Pulau; Borneo: Kalimantan, INDONESIA; WAK; 3°14'N, 114°39'E; collected Oct.-Nov. ca. 3°25'S, 1 14°30'E; observed in 1983 by K. S. 1932 by Oxford University Expedition to Bor- MacKinnon (1986, p. 112). Observed 1988- neo (see Harrisson, 1933, p. 402; P. H. Napier, 1991 by C. H. Southwick and B. Rosenbaum 1981, p. 14); bm(nh), undetermined portion of (1992, p. 88). FAS; CK-25. 6 specimens (including 1 skin only and 3 skulls Kahayan, Sungai, left bank, 60 km above mouth only) collected at Kalulong or Belaga (C:Sar- 1 2). of Sungai Rungan; Borneo: Kalimantan, IN- FAS; C:Sar-20. DONESIA; ca. 1°38'S, 113°56'E; 3 troops ob- Kambang; Sumatra, INDONESIA; 1°42'S, served Jul. 1984-May 1986 by K. M. Burton 100°42'E; collected 19 Nov. 1908 by [H.] (see Chivers & Burton, [1991], p. 140). FAS; Schoede; zmb, 1. FAS; B:S-48. C:K-21. Kambang, Poulo. See Kambing, Pulau. Kahayan, Sungai, left bank, 1 20 km above mouth Kambang, Pulau; Borneo: Kalimantan, INDO- of Sungai Rungan; Borneo: Kalimantan, IN- NESIA; 3°19'S, 1 14°32'E; photographed before DONESIA; ca. 1°08'S, 1 13°54'E; observed Jul. 1968 by Le Roux (see Roedelberger & Gros- 1984-May 1986 by K M. Burton (see Chivers choff, 1967, p. 30). FAS; C:K-25. & Burton, [1991], p. 140). FAS; CK-20. Kambaniru. See Payeti-Kambaniru. Kahayan, Sungai, left bank, above mouth of Sun- Kambas, Wai; Sumatra, INDONESIA; ca. 5°02'S,

142 FIELDIANA: ZOOLOGY 105°52'E; observed in 1983 by K. S. Mac- 114°59'E; observed in 1972 by D. Macdonald Kinnon (1986, p. 112). Observed Jun. 1988 by (1982, pp. 62, 71). FAS; C:B-1.

M. Bismark (1992, p. 1 1). Observed Jan.-Mar. Kampong Mukut, hillsides above; Pulau Tioman, 1989 by A. Yanuar and J. Sugardjito (1993, p. MALAYSIA: WEST MALAYSIA; ca. 2°43'N, 34). FAS; B:S-84. 104°11'E; observed Mar.-Apr. 1962 by Lord Kambera. See Payeti-Kambaniru. Medway (1966, p. 16). FAS; B:SCS-3. Kambing, Pulau; Pulau Sumbawa, Lesser Sunda Kampong Punkah. See Pongka, Kampung. Islands, INDONESIA; 8°27'S, 118°42'E; col- Kampong Rantau Panjang; MALAYSIA: WEST lected 28 Jul. 1 927 by Sunda-Expedition Rensch MALAYSIA; ca. 2°53'N, 101°29'E; examined

(see Mertens, 1930, p. 144; M. fascicularis re- for malaria before 1 95 1 by E. P. Hodgkin ( 1 950, portedly introduced in Pulau Kambing from p. 326). Observed Jul.-Sep. 1960 by Y. Furuya Bima by Sultan of Bima); mzb, 1. FAS; C:LS- (1965, p. 287). FAS; B:WM-18. 12. Kampong Sungai Buloh, 1 50-300 ft (= ca. 70 m); Kambing, Pulau, Lesser Sunda Islands, INDO- MALAYSIA; WEST MALAYSIA; 3°15'N, NESIA; 10°14'S, 123°26'E; collected Jun. 1829 101°18'E; observed 1947-1957 by J. L. Harri- by S. Muller and H. C. Macklot; rmnh, 1 (skin son (1969, p. 176). FAS; B:WM-18. only). FAS; CLS-24. Kampong Tanah Puteh, ; MALAY- Kamoedian, Pulau. See Kemujan, Pulau. SIA: WEST MALAYSIA; 3°35'N, 103°22'E; ex- Kamorta Island. See Camorta Island. amined for malaria ca. 1961-1962 by R. H. Kampong Biserat; THAILAND; 6°32'N, 101°14'E; Wharton, D. E. Eyles, McW. Warren, and W. collected 16 May 1901 by N. Annandale and H. H. Cheong (1964, p. 58). FAS; B:WM-18. C. Robinson (1903, p. xxv; cf. Bonhote, 1903, Kampong Tenghilan. See Tuaran-Kampong p. 3); bm(nh), 3 (including 1 skin only); smtd, Tenghilan Road, new bridge. 1 (skull only). FAS; A:T-69. Kampong Titi Tinggi; MALAYSIA: WEST MA- Kampong Bundu Tuhan, Mount Kinabalu; Bor- LAYSIA; 6°38'N, 100°1 5'E; captive obtained in neo, MALAYSIA: SABAH; 5°59'N, 116°32'E; 1964 by local trappers (Collins et al., 1970, p. collected 30 Jun.-l 8 Jul. 1 95 1 by D. H. Johnson 509). FAS;B:WM-1. (see Coolidge, 1940, p. 124; Davis, 1962, p. 126); Kampon Kadjak. See Kampong Hadjak. usnm, 4. FAS; C:Sab-7. Kampung Baru Study Area; Borneo: Kalimantan, Kampong Cherok Paloh; MALAYSIA: WEST INDONESIA; 1°06'S, 110°10'E; reported pres- MALAYSIA; 3°37'N, 103°23'E; laboratory an- ent 2 Apr.-9 May 1986 by Y. Ruhiyat (1986, imals obtained 1954-1959 by unknown collec- p. 60). FAS; CK-1. tor (Price, 1959, p. 499). FAS; B:WM-13. Kam Shan Entrance, Kowloon Reservoir Area; Kampong Hadjak, Muaratewe district; Borneo: UNITED KINGDOM: HONG KONG; ca. Kalimantan, INDONESIA; ca. 0°57'S, 1 14°53'E; 22°22'N, 114°09'E; introduced population, ob- collected 5 May 1932 by S. A. R. le Prince Le- served Jan.-Feb. 1987 by C. H. Southwick and opold (see Frechkop, 1934, p. 25); irsn, 1 (skin D. Manry (1987, p. 48). Subspecies uncertain; only). FAS; CK-32. not mapped. Kampong Kiau. See Garau; Kiaulan; Tempasuk, Kanaka; Mauritius Island, MAURITIUS; 19°59'- Sungai; Tinonkok. 20°31'S, 57°18'-57°48'E; introduced popula- Kampong Kiau, 3000 ft (= 900 m); Borneo, MA- tion, collected 29 May 1975 by R. L. Ciochan; LAYSIA: SABAH; ca. 6°02'N, 116°31'E; col- bm(nh), 1 (skull not examined). Subspecies un- lected 24-29 Apr. 1929 by F. N. Chasen (1931, certain; not mapped. p. 6); zrc, 2. FAS; C:Sab-7. Kangean, Pulau, 4 ft (= 1 m), INDONESIA; 6°49'- Kampong Kiau, ca. 2200 ft (= 670 m); Borneo, 7°00'S, 1 1 5°1 2'-l 1 5°34'E; collected 1 5-23 Nov. MALAYSIA: SABAH; ca. 6°02'N, 1 1 6°3 1 'E; col- 1909 by G. C. Shortridge; bm(nh), 4. FAS; C:J- lected 13-23 Aug. 1937 by J. A. Griswold, Jr. 40. (1939b, p. 514; mcz Asiatic Primate Expedition Kantan, See Kantang.

specimen list); mcz, 2. FAS; C:Sab-7. Kantang; THAILAND; 7°25'N, 99°31'E; collected Kampong Kiau-Tenampok Pass, trail between, [ca. 1 1 Jan. 1 9 1 8 by C. B. Kloss (see H. C. Robinson 1100 m]; Borneo, MALAYSIA: SABAH; ca. & Kloss, 1910, p. 668; Weitzel et al., 1988, p. 6°00'N, 116°30'E; observed 3 Jun. 1932 by J. 104; W. L. Abbott, 1897, unpubl. map in usnm A. Griswold, Jr. (1939a, p. 410). FAS; C:Sab-7. archives); zrc, 1. FAS; A:T-56. Kampong Menuggol; Borneo, BRUNEI; 4°53'N, Kapos Tinggi; Pulau Bengkalis, INDONESIA; ca.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 143 1°27'N, 102°18'E; collected 22 Mar. 1906 by W. 1994 (Anonymous, 1994, p. 12). Subspecies un- L. Abbott (field catalog; in Lyon, 1908, p. 623); certain; A:Bu-16. usnm, 1. FAS; B:SM-6. Kata Taek, ca. 200 m; THAILAND; ca. 15°28'N, Kapuas, Sungai. See Semitau, Sungai Kapuas. 99°23'E; collected 2-3 Mar. 1967 by J. Fooden Kapuas, Sungai, right bank, 25 km above mouth; (1971, p. 17); fmnh, 6. FAS; A:T-19. Borneo: Kalimantan, INDONESIA; ca. 2°48'S, Katchal Island. See Ol-kolo-kwak vicinity; Katch-

1 14°17'E; observed Jul. 1984-May 1986 by K. all Island. M. Burton (see Chivers & Burton, [1991], p. Katchall Island, Nicobar Islands, INDIA; 7°52'- 140). FAS; C:K-24. 8°02'N, 93°18'-93°27'E; observed before 1847 Karakit, Pulau Banggi, MALAYSIA: SABAH; by P. Barbe (1846, p. 365). UMB; A:N-1. 7°07'N, 117°05'E; collected 22 Jun. 1991 by Kateman, Sungai, < 2 ft (= < 1 m); Sumatra, Shukor Md. Nor (pers. comm., 17 Jul. 1991); INDONESIA; 0°12'N, 103°20'E; collected 8 Sep. fmnh, 1. FAS; C:Sab-9. 1903 by W. L. Abbott (in Lyon, 1908, p. 625); Karangan, Sungai; Borneo: Kalimantan, INDO- usnm, 1 (skull only). FAS; B:S-57. NESIA; 1°19'N, 117°55'E; collected 1 Nov. 1913 Kathema Kyun, BURMA; 13°39'N, 98°12'E; col- by H. C. Raven (see Deignan, [1960], p. 269); lected 20-22 Apr. 1 936 by H. C. Smith; bm(nh), usnm, 1 (skull only). FAS; CK-44. 4. AUR; A:Bu-15. Karangintan, hilly country, Sungai Martapura, 30 Katingan, Sungai, left bank, 1 60 km above mouth; ft (= 1 m); Borneo: Kalimantan, INDONESIA; Borneo: Kalimantan, INDONESIA; ca. 1°47'S, 3°26'S, 1 14°55'E; collected 19 Aug. 1909 by G. 1 13°19'E; observed Jul. 1984-May 1986 by K C. Shortridge; bm(nh), 1. FAS; CK-27. M. Burton (see Chivers & Burton, [1991], p. Karanginton. See Karangintan. 140). FAS;C:K-13. Karangmumus, Sungai, near Samarinda; Borneo: Katingan, Sungai, left bank, 200 km above mouth; Kalimantan, INDONESIA; 0°30'S, 117°09'E; Borneo: Kalimantan, INDONESIA; ca. 1°30'S, collected 3 Jul. 1912 by H. C. Raven (field cat- 1 13°10'E; observed Jul. 1984-May 1986 by K. alog); usnm, 1. FAS; GK-49. M. Burton (see Chivers & Burton, [1991], p. Karang Tigan. See Karangtigau, Tanjung. 140). FAS;C:K-14. Karangtigau, Tanjung; Borneo: Kalimantan, IN- Katingan, Sungai, right bank, 1 40 km above mouth; DONESIA; 2°26'N, 1 18°00'E; collected 4 Aug. Borneo: Kalimantan, INDONESIA; ca. 2°04'S, 1 9 1 2 by H. C. Raven (field catalog; cf. Deignan, 1 13°25'E; observed Jul. 1984-May 1986 by K

1 1 1 1 [1960], p. 267); usnm, 2 (including skin only). M. Burton (see Chivers & Burton [ 99 ], p. 40). FAS; C:K-40. FAS;C:K-11. Karawassen. See Kawarasan. Kawarasan; Java, INDONESIA; 7°49'S, 1 12°07'E; Karimata. See Pai, Teluk; Pulau Karimata. collected 20 Apr. 1895 by unknown collector; Karimon Anak. See Karimun-kecil, Pulau. rmnh, 1 (skin only). FAS; CJ-33. Karimon Djawa. See Karimunjawa, Pulau. Kayan, Sungai. See Long Peleben. Karimon-Djawa, Pulau, (P. Kamoedian). See Ke- Kebun Percobaan Haurbentes, Areal; Java, IN- mujan, Pulau. DONESIA; not located; obseved Oct. 1989 and Karimunjawa, Pulau, INDONESIA; 5°50'-5°53'S, Jan. 1991 by M. Bismark (1992, p. 11). FAS; not 1 10°25'-1 10°29'E; collected 7-14 May 1926 by mapped. K. W. Dammerman, Denin, and P. F. Franck; Kediri. See Manggis; Margomulio, Gunung. mzb, 5 (including 1 skull only). Collected 26-28 Kediri district; Java, INDONESIA; ca. 7°48'S, 1 12° Nov. 1930 by W. Romswinckel; mzb, 2; rmnh, 15 'E; collected 13 Nov. 1910 and 25 Nov. 2 1 (skull not examined). KAR: C:J-27. 1911 by V. Arnim; zsbs, (skulls only; not seen, Karimun-kecil, Pulau, Kepulauan Riau, INDO- data from G. H. Albrecht, pers. comm., Oct. NESIA; 1°07'-P10'N, 103°22'-103°25'E; re- 1991). FAS; GJ-33. ported present Jun.-Aug. 1903 by W. L. Abbott Kelabit uplands, 4000 ft. See Dulit, Bukit, 4000 (see Miller, 1906c, p. 277). FAS; B:SCS-17. ft. Kariorang; Borneo: Kalimantan, INDONESIA; Kelabong, Bukit, vicinity; Sumatra, INDONE- 0°50'N, 117°52'E; collected in 1903 by M. SIA; ca. 3°13'S, 102°26'E; reported prsent Nov. Schmidt; zmb, 2 (skulls only). FAS; C:K-45. 1971-Jan. 1973 by local residents (Crockett & Kaser Doo Wildlife Sanctuary; BURMA; 13° 15'- Wilson, 1980, p. 156). FAS; B:S-67. 13°24'N, 98°45'-99°00'E; reported present ca. Kelang Road, 7.25 km; MALAYSIA: WEST MA-

144 FIELDIANA: ZOOLOGY LAYSIA; ca. 3°05'N, 101°40'E; collected 2 Dec. 1973 by C. M. Crockett and W. L. Wilson (1980, 1953 by Scrub Typhus Research Unit; bm(nh), p. 156). FAS;B:S-13. 1. FAS; B:WM-19. Ketambe, ca. 30 km SSE; Sumatra, INDONESIA; Kelapa, Pulau, Lesser Sunda Islands, INDONE- ca. 3°27'N, 97°50'E; reported present before 1 98 1 SIA; 8°39'-8°42'S, 119°13'-119°14'E; monkeys by M. Borner (Crockett & Wilson, 1980, p. 1 56). reported absent 1969-1973 by W. Auffenberg FAS; B:S-14. (1981, p. 40). C:e. Ketambe Research Station, Gunung Leuser Na- Kembangjanggut; Borneo: Kalimantan, INDO- tional Park, ca. 350 m; Sumatra, INDONESIA; NESIA; 0°08'N, 116°22'E; collected 28 Nov. 3°40'N,97°40'E; observed Jun. 1971-Aug. 1974 1956 by A. M. R. Wegner and Saan; mzb, 1. by H. D. Rijksen (1978, p. 1 1 1). Hormonal study FAS; CK-48. Oct.-Nov. 1987 by C. P. van Schaik, M. A. van Kemujan, Pulau, INDONESIA; 5°47'-5°51'S, Noordwijk, T. van Bragt, and M. A. Blanken- stein B:S-13. 1 10°27'-1 10°30'E; reported present before 1906 (1991, p. 347). FAS; not located by T. Willink (1905, p. 175). Collected 25 Nov. "Kg. Baru"; Sumatra, INDONESIA; Gazetteer of 1930 by W. Romswinckel; mzb, 1 (skull only). (USBGN Indonesia, 1982, p. 483, KAR; C:J-27. lists 2 populated places named "Kampungbaru" in Kenepai, Gunung. See Roema Manoeal, south foot Sumatra), 5°38'N-5°57'S, 95°12'-106°05E; collected in 1906 and 1910 C. of Gunung Kenepai. by Bruegel; zsbs, 10 (including 5 skins only, 2 skulls only). FAS; Keningau, 800 ft (= 250 m); Borneo, MALAYSIA: not mapped. SABAH; 5°20'N, 1 16°10'E; collected Sep. 1960 Khangkhao, Ko, THAILAND; 13°06'-13°07'N, by R. E. Kuntz (1969, p. 193); amnh, 1. FAS; 15 Nov. C:Sab-6. 100°48'-100°49'E; reported present 1990 by local boatman (Aggimarangsee, 1992, Kenokok, 3300 ft (= 1000 m); Borneo, MALAY- pp. Ill, 130; pers. comm., Oct. 1993). Subspe- SIA: SABAH; ca. 6°04'N, 116°28'E; collected cies uncertain; A:T-3 1 . 25 Apr. 1929 by F. N. Chasen (1931, p. 6); zrc, Khao Khieo Wildlife Sanctuary; THAILAND; 1. FAS;C:Sab-7. 13°13'N, 101°04'E; introduced population, re- Kenyam, Sungai, 100 m; MALAYSIA: WEST leased before Jan. 1977, observed Jun. 1977- MALAYSIA; 4°31'N, 102°28'E; observed Jul. Jun. 1978 by P. J. Storer (1979, pp. 28, 46). 1978-Jun. 1981 by C. W. Marsh and W. L. Subspecies uncertain; not mapped. Wilson (1981, p. 232). FAS; B:WM-1 1. Khao Lampi-Hat Thai Muang National Park; Keramat. See Tanjong Tuan, Keramat. THAILAND; ca. 8°23'N, 98°20'E; reported Kerinci. See Sandaran Kerinci Siu- Agong, region; present 21-25 Nov. 1987 by local residents lakderas, Kerinci region. (Boonratana, 1988, p. 76). AUR/FAS/MUL; Kerinci, Gunung, northern foot; Sumatra, IN- A:T-51. DONESIA; 1°40'S, 101°20'E; observed Jun. Khao Naw. See Wat Khao Noh. T. Oi 1985-Mar. 1986 by (1986, p. 73). FAS; Khao Ngu. See Wat Ratch Singkhorn. B:S-52. Khao Noh. See Wat Khao Noh. Keroh Forest Reserve; MALAYSIA: WEST MA- Khao Noi/Khao Tangkuan; THAILAND; 7°1 3'N, LAYSIA; ca. 4°13'N, 101°07'E; observed 2 Jun. 100°36'E; blood samples taken Aug.-Sep. 1988 1935 by R. C. Morris (1936, p. 440). FAS; by P. Varavudhi, J. Suzuki, U. Yodyingyuad, B:WM-16. T. Nootprapand, V. Yodyingyuad, Y. Chaiseha, Kertau, Bukit; MALAYSIA: WEST MALAYSIA; K. Suwanprasert, and W. Settheetham (1989b,

ca. 3°27'N, 102°37'E; examined for malaria p. 77; cf. Kawamoto et al., 1989, p. 95). Ob- 1965-1969 by McW. Warren, W. H. Cheong, served 9-14 Apr. 1989 by N. Aggimarangsee H. K. Fredericks, and G. R. Coatney (1970, p. (1992, pp. 109, 139; pers. comm., Oct. 1993). 386). FAS; B:WM-14. FAS; A:T-67. Ketambe, ca. 5 km S; Sumatra, INDONESIA; ca. Khao Noi-Khuo Tanguan. See Khao Noi/Khao 3°38'N, 97°40'E; observed Nov. 1971-Jan. 1973 Tangkuan. by C. M. Crockett and W. L. Wilson (1980, p. Khao Paskowee. See Khao Phatowee. 156). FAS;B:S-13. Khao Phatowee; THAILAND; ca. 15°28'N, Ketambe, ca. 10 km S; Sumatra, INDONESIA; 99°45'E; observed 25 Feb. 1967 by J. Fooden ca. 3°36'N, 97°40'E; observed Nov. 1971-Jan. (1971, p. 16). Observed 1973-1974 by A. A.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 145 Eudey (1979, p. 90; 1994, p. 274). FAS; A:T- Kinabatangan, Sungai; Borneo, MALAYSIA: SA- 22. BAH; ca. 5°42'N, 1 18°23'E; collected 19-23 Dec.

Khao Rang Kai, 4 km E. of Ban Palian, 75 m; 1887 by C. F. Adams (see Medway, 1977, p. 4); THAILAND; 7°19'N, 99°48'E; collected 6 Jul. usnm, 2 (including 1 skull only). FAS; C:Sab- 1973 by J. Fooden ([1975], p. 98); ctnrc, 1; 19. fmnh, 1. FAS; A:T-62. King's Island. See Kadan, Kyun. Khao Sam Muk. See Sammuk, Khao. Kinta, Daerah (district); MALAYSIA: WEST Khao Sam Roi National Park; THAILAND; MALAYSIA; ca. 4°30'N, 101°12'E; collected 12°05'-12°17'N, 99°53'-100°02'E; reported before 1 90 1 by unknown collector (Rower, 1 900, present before 1994 by J. W. K. Parr, N. Ma- p. 316); specimens reportedly in Taiping Mu- hannop, and V. Charoensiri (1993, p. 245). seum (not seen). FAS; B:WM-7. AUR/FAS/MUL; A:T-40. Klaeng. See Wang Kaew. Khao Suan Luang; THAILAND; 1 3°35'N, 99°45'E; Klang Road, 4V2 miles. See Kelang Road, 7.25 km. observed 24 Dec. 1990 by N. Aggimarangsee Klang Straits. See Pintu Gedong, Pulau. (1992, pp. Ill, 128; pers. comm., Oct. 1993). Klet Kaeo, Ko, THAILAND; 12°46'N, 100°51'E; FAS; A:T-28. trapped spring 1967 by G. Berkson (1970, p. Khao Wang; THAILAND; 13°06'N, 99°56'E; ob- 286). Subspecies uncertain; A:T-39. served 12-13 Aug. 1989 and 9-10 Feb. 1991 by Kloet, Goenoeng. See Manggis, Gunung Kelud; N. Aggimarangsee (1992, pp. 109, 112, 130); Margomulio, Gunung. pers. comm., Oct. 1993). Subspecies uncertain; Klong Pah Yie; Ko Samui, northwest, THAI- A:T-37. LAND; ca. 9°32'N, 99°57'E; collected 7 May Khram Yai, Ko, THAILAND; 12°40'-12°43'N, 1913 by H. C. Robinson (1915, p. 129) and E. 100°45'-100°48'E; collectd 30 Oct. 1916 by C. Seimund; zrc, 1. FAS; A:T-49. B. Kloss (1919c, p. 335); bm(nh), 2; usnm, 5; Klumpang, Teluk; Borneo: Kalimantan, INDO- zrc, 2. Collected 1 Jun. 1917 by W. J. F. Wil- NESIA; ca. 3°00'S, 1 16°12'E; observed 8 Jan.- liamson; zrc, 2. ATR; A:T-39. 13 Mar. 1908 or 18-19 Apr. 1909 by W. L.

Khulna disrict, Sundarbans; BANGLADESH; Abbott (in Lyon, 191 1, p. 58). FAS; GK-51. 22°20'N, 89°27'E; improbable report (Anony- Kode, Nusa, Lesser Sunda Islands, INDONESIA; mous, 1977, p. 14; cf. Sarker & Sarker, 1984, 8°47'-8°49'S, 1 19°37'-1 19°40'E; reported pres- p. 9; M. A. R. Kahn, 1985, p. 30). Not mapped. ent 1969-1973 by W. Auffenberg (1981, p. 40). Khwae Noi, Mae Nam, right bank, ca. 10 km be- FAS;C:LS-13. low Ban Wang Kalang; THAILAND; ca. Koh Alang Yai. See Rang Yai, Ko. 15°03'N, 98°30'E; observed 3 Feb. 1967 by J. Koh Chang Island. See Chang, Ko. Fooden (1971, p. 15). AUR; A:T-20. Kohhang. See Pran Buri, Mae Nam, mouth. Kiambang; Sumatra, INDONESIA; 5°27'S, Koh Kram Island. See Khram Yai, Ko. 105°44'E; observed Nov. 1971-Jan. 1973 by C. Koh Kut Island. See Kut, Ko. M. Crockett and W. L. Wilson (1980, p. 156). Koh Lak. See Prachuap Khiri Khan. FAS; B:S-86. Koh Lang. See Rang Yai, Ko. Kiau. See Garau; Kampong Kiau; Kampong Kiau- Koh Naka Yai. See Naka Yai, Ko. Tenampok Pass, trail between; Kiaulan; Tem- Koh Nam Kam. See Nang Kham, Ko. pasuk, Sungai; and Tinonkok. Koh Pennan. See Phangan, Ko. Kiaulan, Kampong Kiau region, ca. 2200 ft (= 670 Koh Piam. See Phayam, Ko. m); Borneo, MALAYSIA: SABAH; ca. 6°02'N, Koh Pipidon. See Phi Phi Don, Ko. 116°31'E; collected 17 Aug. 1937 by J. A. Gris- Koh Samui. See Samui, Ko. wold, Jr. (1939b, p. 514); mcz, 2. FAS; C:Sab-7. Koh Si Chang. See Si Chang, Ko. Kinabalu, Mount; Borneo, MALAYSIA: SABAH; Ko Khanghkao. See Khangkhao, Ko. ca. 6°05'N, 1 16°33'E; collected 31 May-28 Aug. Ko Khangkao. See Khangkhao, Ko.

1937 by J. A. Griswold, Jr. (1 939a, p. 402; 1 939b, Ko Klet Kaeo. See Klet Kaeo, Ko. p. 504); mcz, 9 (including 6 skulls only, 3 man- Komodo, Pulau, Lesser Sunda Islands, INDO- dibles only). FAS; C:Sab-7. NESIA; 8°26'-8°46'S, 119°22'-119°34'E; mon- Kinabalu National Park; Borneo, MALAYSIA: keys reported absent Jul. 1969-Jun. 1970 by W. SABAH; ca. 6°05'N, 1 16°35'E; reported present Auffenberg (1981, p. 242). C:e. before 1983 by G. Davies([1983],p. 148). FAS; Kopenheat; Great Nicobar Island, INDIA; 6°58'N, C:Sab-7. 93°43'E; collected 23 Mar. 1901 by W. L. Ab-

146 FIELDIANA: ZOOLOGY bott (field catalog; cf. Miller, 1902b, p. 751; LAYSIA; 3°43'N, 102°17'E; observed Jul. 1978- Kloss, 1 903a, map facing p. 8); usnm, 1 UMB; Jun. 1981 by C. W. Marsh and W. L. Wilson A:N-3. (1981, p. 232). Observed Mar. 1984-Oct. 1986 Korinchi. See Sandaran Agong. by F. Lambert (1990, p. 455). FAS; B:WM-15. Ko Sichang. See Si Chang, Ko. Kuala Lompat Post, 0-1 km NW; MALAYSIA: Kosumpee. See Kosumphi Forest Park. WEST MALAYSIA; ca. 3°43°N, 102°17'E; ob- Kosumphi Forest Park; THAILAND; 16°15'N, served Jul.-Dec. 1977 by D. Chivers and G. 103°05'E; blood samples taken Aug.-Sep. 1988 Davies (1979, p. 19). Observed Jul. 1974-Jan. by Y. Kawamoto, T. Ishida, J. Suzuki, O. Tak- 1976 by F. P. G. Aldrich-Blake (1980, p. 147). enaka, and P. Varavudhi (1989, p. 95). Ob- FAS; B:WM-15. served 21 Jul. 1989 and 12 Jan. 1991 by N. Kuala Lompat Post, 0-2 km W; MALAYSIA: Aggimarangsee (1992, pp. 109, 111, 119;pers. WEST MALAYSIA; ca. 3°42'N, 102°17'E; ob- comm., Oct. 1993). Subspecies uncertain; A:T- served Jul.-Dec. 1977 by D. Chivers and G. 12. Davies (1979, p. 19). FAS; B:WM-15. Kosumphisai. See Kosumphi Forest Park. Kuala Lumpur. See Nanas, Bukit, Kuala Lumpur. Kotabumi, Propinsi Lampung, 23 m; Sumatra, Kuala Lumpur vicinity; MALAYSIA: WEST MA- INDONESIA; 4°50'S, 104°54'E; collected 6 May LAYSIA; ca. 3°10'N, 101°42'E; observed 1901- 1940 by Vogelpol; zrc, 1. FAS; B:S-81. 1 902 by N. Annandale and H. C. Robinson (see Kota Kinabalu. See Papar; Talibang; Tuaran. Bonhote, 1903, p. 4). Observed Aug.-Dec. 1970

Kotapinang. See Telukpanji, Kotapinang region. by C. H. Southwick and F. C. Cadigan, Jr. ( 1 972, Kotawaringin. See Riam, Kotawaringin district. p. 13). Observed Jun. 1976-Jul. 1977 by Y. L. Kowloon Reservoir Area; UNITED KINGDOM: Mah and F. P. G. Aldrich-Blake (1980, p. 354). HONG KONG; ca. 22°21'N, 114°09'E; intro- Laboratory animals obtained 1 979-1 980 by un- duced population, reported present ca. 1967 by known collectors (Burke et al., 1981, p. 928). P. Marshall (1967, p. 44). Subspecies uncertain; FAS; B:WM-19. not mapped. Kuala Pilah vicinity; MALAYSIA: WEST MA- Krakatau, Pulau. See Rakata, Pulau. LAYSIA; ca. 2°44'N, 102°15'E; examined for Krakatau Ketjil, Pulau. See Rakata-kecil, Pulau. malaria before 1994 by A. N. Rain, J. W. Mak, Krau Game Reserve. See Kuala Lompat. and R. Zamri (1993, p. 386). FAS; B:WM-25. Krau River. See Benom, Gunong. Kuala Rompin, 5 m; MALAYSIA: WEST MA- Kretam Besar, Sungai, Kinabatangan district, 70 LAYSIA; 2°50'N, 103°26'E; observed Jul. 1978- m; Borneo, MALAYSIA: SABAH; 5°32'N, Jun. 1981 by C. W. Marsh and W. L. Wilson 1 1 8°32'E; collected 25 May 1 950 by D. D. Davis (1981, p. 232). FAS; B:WM-22.

1 1 1 1 . ( 962, p. ; field catalog); fmnh, FAS; C:Sab- Kuala Selangor, 0-40 m; MALAYSIA: WEST 20. MALAYSIA; 3°21'N, 101°15'E; observed Feb. Kretam Kechil, Sungai, Kinabatangan district, 70 1965-Jun. 1966 by I. S. Bernstein (1968b, p. 8). m; Borneo, MALAYSIA: SABAH; 5°31'N, Observed Aug.-Dec. 1970 by C. H. Southwick 1 1 8°33'E; collected 1 3 May 1 950 by D. D. Davis and F. C. Cadigan, Jr. (1972, p. 10). FAS; B:WM-

1 1 1 field 1 . 18. ( 962, p. ; catalog); fmnh, FAS; C:Sab- 20. Kuala Selangor, sea level; MALAYSIA: WEST Reserve. See Keroh Forest Reserve. MALAYSIA; 3°20'N, 101°17'E; reported pres- Kuala Belalong Field Studies Centre; Borneo; ent before 1973 by B. E. Weber (1972, p. 468). BRUNEI; ca. 4°33'N, 115°08'E; reported pres- Observed Jul. 1978-Jun. 1981 by C. W. Marsh ent 1991-1992 by Earl of Cranbrook (1993, p. and W. L. Wilson (1981, p. 232). FAS; B:WM- 274). FAS; not mapped (record discovered after 18. Fig. 2C was prepared). Kuala Selangor estuary, south bank; MALAYSIA: Kuala Binjai. See Binjai, Sungai. WEST MALAYSIA; ca. 3°21'N, 101°15'E; ob- Kuala Indau. See Endau, Sungai, vicinity. served Mar.-Apr. 1977 by Lim Boon Hock and Kuala Lompat; MALAYSIA: WEST MALAY- A. Sasekumar (1979, p. 106). FAS; B.WM-18. SIA; ca. 3°42'N, 102°17'E; observed 1969-1970 Kuantan; MALAYSIA: WEST MALAYSIA; by D. Chivers (1971, p. 80). Observed 1969- 3°48'N, 103°20'E; captive obtained ca. 1963 by 1970 by Lord Medway and D. R. Wells (1971, unspecified collectors (Garnham, 1963, p. 156). p. 246). FAS;B:WM-15. FAS; B:WM-13. Kuala, Lompat, 50 m; MALAYSIA: WEST MA- Kuatnana, 300 m; Pulau Timor, Lesser Sunda Is-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 147 lands, INDONESIA; ca. 9°50'S, 124°10°E; col- Abbott (see Miller, 1903a, p. 437); usnm, 1. lected 20 Jul. 1911 by C. B. Haniel (see Hell- FUS; B:IO-3. mayr, 1914, p. 5); zsbs, 1. FAS; QLS-26. Labuhandeli. See Deli, Sungai, between Belawan Kuching; Borneo, MALAYSIA: SARAWAK; and Labuhandeli. 1°33'N, 1 10°20'E; collected 23 Sep. 1893 by un- Labuhandeli vicinity; Sumatra, INDONESIA; known collector; smk, 1. Collected Mar. 1898 3°45'N, 98°4 1 'E; collected 1881-1883 by B. Ha- by unknown collector; smk, 1. Collected 25 Jul. gen (1890, p. 82); museum unknown, 1 (not 1 898 by C. Ulok; smk, 1 (skin only, skull inside). seen). FAS; B:S-21. Collected date unknown by unknown collector; Labuk Road, Sepilok Forest Reserve; Borneo, smk, 2 (skulls only). FAS; C:Sar-4. MALAYSIA: SABAH; ca. 5°53'N, 118°00'E; Kuching, 10th mile; Borneo, MALAYSIA: SA- observed 1968-1969 by M. Kawabe and T. RAWAK; ca. 1°33'N, 1 10°20'E; collected 16 Jan. Mano (1972, p. 216). FAS; C:Sab-17. 1895 by unknown collector; smk, 1 (skin only, Lac Giao, 400 m; VIETNAM; 12°40'N, 108°03'E; skull inside). FAS; C:Sar-4. collected 18 Mar. 1937 by W. H. Osgood (1941, [Kuching, probably]; Borneo, MALAYSIA: SA- p. 1; field catalog); fmnh, 1. AUR/FAS/MUL; RAWAK; 1°33'N, 110°20'E; collected 1865- A:V-2. 1866 by G. Doria (see Beccari, 1904, p. 30); Lacon. See Nakhon Si Thammarat. zmb, 1. FAS; C:Sar-4. Lacon Stritamarat. See Nakhon Si Thammarat. Kuda. See Poelau (?= Kuda), Sungai Sibau. La Datu. See Lahad Datu. Kukuh; Pulau Bali, Lesser Sunda Islands, IN- Laem Ngop-Phumi Cham Yeam, river between; DONESIA; ca. 8°31'S, 1 15°12'E; blood samples CAMBODIA or THAILAND; ca. 12°00'N, taken in 1980 by Y. Kawamoto, K. Nozawa, 102°47'E; observed in 1914 by C. B. Kloss - andTb. M. Ischak(1981,p. 16). Observed Jul. (1916a, p. 32). FAS; A:C-3. Sep. 1993 by M. F. Small (1994, p. 10). FAS; Laem Sing mountains; THAILAND; 12°29'N, QLS-3. 102°04'E; collected 7 Jun. 1926 by H. M. Smith Kundur, Pulau, Kepulauan Riau, INDONESIA; (see Riley, 1938, p. 9); usnm, 1. FAS; A:T-33. 0°38'-0°53'N, 103°21'-103°30'E; reported pres- Laem Son National Park; THAILAND; ca. 9°17'N, ent Jun.-Aug. 1 903 by W. L. Abbott (see Miller, 98°31'E; observed 25-28 Nov. 1987 by R. 1906c, p. 279). Collected 19 Aug. 1908 by E. Boonratana (1988, p. 76). AUR/FAS/MUL; Seimund; zrc, 2 (skulls only). FAS; B:SCS-17. A:T-46. Kuprakona. See Wat Koo Pra Kona. Lafau; Pulau Nias, INDONESIA; 1°23'N, 97°13'E;

Ku Prakonna. See Wat Koo Pra Kona. collected 24 Mar. 1905 by W. L. Abbott (cf. Kut, Ko, THAILAND; 1 1°33'-1 1°46'N, 102°32'- Lyon, 1916, p. 458); usnm, 1. FAS; B:IO-6. 102°37'E; collected 23-29 Dec. 1914 by C. B. Lagong, Pulau, INDONESIA; 3°34'-3°38'N, Kloss (1916b, p. 28); bm(nh), 3; bm(nh) (skull)/ 108°04'-108°08'E; collected 19 Jun. 1900 by W. zrc (skin), 1; usnm, 2; zrc, 1. Collected 1 Jun. L. Abbott (see Miller, 1901, p. Ill); usnm, 1. 1917 by H. M. Smith (see Riley, 1938, p. 2); FAS; B.SCS-34. usnm, 1. FAS; A:T-36. Lahad Datu; Borneo, MALAYSIA: SABAH; Kuta; Pulau Lombok, Lesser Sunda Islands, IN- 5°02'N, 118°19'E; collected 10 Nov. 1903 by DONESIA; 8°55'S, 116°17'E; blood samples Dr. Pagel; zmb, 1. FAS; C:Sab-21. taken Jan. 1979-Dec. 1981 by Y. Kawamoto Lahat; Sumatra, INDONESIA; 3°48'S, 103°32'E; (1982, p. 66). FAS; QLS-6. collected in 1875 by unknown collector; NHMBa, Kutai Nature Reserve, northeast corner; Borneo: 2 (skulls only). FAS; B:S-73. Kalimantan, INDONESIA; ca. 0°30'N, Lai char; BURMA; ca. 21°05'N, 92°12'E; ob- 117°30'E; observed 1 May 1970-31 Jul. 1971 served Sep. 1982-Feb. 1983 by M. A. R. Khan by P. Rodman (1973, p. 655). Observed Oct. and M. A. Wahab (1983, p. 104; cf. M. A. R. 1974-Jun. 1976 by B. P. Wheatley (1978, p. Khan, 1985, p. 30). AUR; A:Bu-l. 347). FAS; CK-46. Lampung. See Wonosobo, Propinsi Lampung. Kute. See Kuta. Lampung, Propinsi (province); Sumatra, INDO- Kya-eng. See Ataran River. NESIA; ca. 5°10'S, 104°45'E; collected in 1908 Labuan Badjan Bay. See Labuhanbajau. by J. Elbert; zsbs, 1 (skull only). Blood samples Labuhanbajau; Pulau Simeulue, INDONESIA; taken Jan.-Nov. 1 979 by Y. Kawamoto and Tb. 2°24'N, 96°28'E; collected 1 Jan. 1902 by W. L. M. Ischak (1981, p. 238). FAS; B:S-80.

148 FIELDIANA: ZOOLOGY Lampungs. See Lampung, Propinsi (province). Lasia, Pulau, INDONESIA; 2°08'-2°12'N, 96°37'- Lamukotan, Pulau. See Lemukutan, Pulau. 96°40'E; collected 4-5 Jan. 1902 by W. L. Ab- Lanbi Kyun, BURMA; 10°42'-10°59'N, 98°02'- bott (see Miller, 1903a, p. 438); usnm, 2. LAS; 98°18'E; collected 30 Jan. 1900 and 6 Jan. 1904 B:IO-4. by W. L. Abbott; usnm, 2 (including 1 skull Lat Bua Kao. See Lat Bua Khao, Sathani. only). AUR; A:Bu-21. Lat Bua Khao, Sathani; THAILAND; 14°52'N, Lancang Kuning; Pulau Bintan, Kepulauan Riau, 101°36'E; collected 10 Oct 1916 by C. B. Kloss INDONESIA; not precisely located, 0°48'- (1919c, p. 335); usnm, 1; zrc, 1. FAS; A:T-9. 1°13'N, 104°13'-104°34'E; observed 18 Jan.-2 Laut, Pulau, INDONESIA; 4°40'-4°46'N, 107°56'- Feb. 1992 by A. Yanuar (1994, p. 2). FAS; 108°02'E; collected Jul. 1 894 by C. Hose; bm(nh), B:SCS-8. 1 (skin only). Collected 9 Aug. 1900 by W. L.

Landai vicinity; Sumatra, INDONESIA; ca. Abbott (see Miller, 1901, p. 11 1); usnm, 1 . FAS; 0°01'S, 100°37'E; reported Nov. 1971-Jan. 1973 B:SCS-31. by local residents (Crockett & Wilson, 1980, p. Lauttador; Sumatra, INDONESIA; 3°18'N, 156). FAS;B:S-54. 99°15'E; collected 1897-1899 by G. Schneider Landak. See Perbuah, Sungai Landak. (1905, p. 72); museum and number of speci- Langgaliroe; Pulau Sumba, Lesser Sunda Islands, mens unknown (not seen). FAS; B:S-23. INDONESIA; not precisely located, 9° 17'- Laut Tawar, Danau. See Bur ni Bebuli. 10°19'S, 1 18°57'-120°50'E; collected 24-29 May Lawadarat, Gili, Lesser Sunda Islands, INDO- 1932 by G. Stein; mzb, 2; zmb, 1. FAS; C:LS- NESIA; 8°28'-8°29'S, 119°33'-119°34'E; mon- 22, LS-23. keys reported absent 1969-1973 by W. Auffen- Lang Island. See Rakata-kecil, Pulau. berg(1981,p. 40). C:d. Langkawi, Pulau, MALAYSIA: WEST MALAY- Lawalaut, Gili, Lesser Sunda Islands, INDONE- SIA; 6°15'-6°28'N, 99°38'-99°55'E; collected 10 SIA; 8°27'-8°28'S, 119°33'-119 35'E; monkeys Feb. 1909 by museum collector (see H. C. Rob- reported absent 1969-1973 by W. Auffenberg inson & Kloss, 1910, p. 664; H. C. Robinson, (1981, p. 40). C:d. 1917, p. 130); bm(nh), 1. FAS; B:SM-1. Lelogama, 845 m; Pulau Timor, Lesser Sunda Is- Langkawi, Pulau, 50 m, MALAYSIA: WEST MA- lands, INDONESIA; 9°44'S, 123°57'E; collected LAYSIA; 6°15'-6°28'N, 99°38'-99°55'E; ob- 22 May-3 Jun. 191 1 by C. B. Haniel (see Hell- served Jul. 1978-Jun. 1981 by C. W. Marsh and mayr, 1 9 1 4, p. 5); zsbs, 6 (includes 3 skins only); W. L. Wilson (1981, p. 232). FAS; B:SM-1. FAS; CLS-26. Langkoe. See Langkoi, Pulau. Lem Sing Mts. See Laem Sing mountains. Langkoi, Pulau, Lesser Sunda Islands, INDO- Lemukutan, Pulau, INDONESIA; 0°43'-0°47'N, NESIA; 8°44'S, 1 1 9°22'E; monkeys reported ab- 108°42'-108°44'E; reported present 7-10 May sent 1969-1973 by W. Auffenberg(1981, p. 40). 1907 by W. L. Abbott (in Lyon, 191 1, p. 59). C:e. FAS; B:SCS-24. Lanjak-Entimau Orang-Utan Sanctuary (pro- Lengah, Pulau, Lesser Sunda Islands, INDONE- posed); Borneo, MALAYSIA: SARAWAK; ca. SIA; 8°4 l'S, 1 1 9°28'E; monkeys reported absent 1°30'N, 11 2°05'E; observed Aug.-Nov. 1981 by 1969-1973 by W. Auffenberg (1981, p. 40). C:e. Sarawak Forest Department— World Wildlife Lesong, 100 m; MALAYSIA: WEST MALAY- Fund team (Kavanagh, 1982, p. 320). FAS; SIA; 2°44'N, 103°09'E; reported as probably C:Sar-ll. present Jul. 1978-Jun. 1981 by C. W. Marsh Lankawi, P. See Langkawi, Pulau. and W. L. Wilson (1981, p. 232). FAS; B:WM- Lanta Yai, Ko, THAILAND; 7°28'-7°4rN, 21. 99°02'-99°08'E; collected 9-12 Jan. 1917 by H. Lesten, Daerah Istimewa Aceh, 700 m; Sumatra, C. Robinson and E. Seimund (H. C. Robinson, INDONESIA; 4°10'N, 97°40'E; collected 19 1917, p. 135); museum unknown, 1 (not seen). Mar. 1937 by A. Hoogerwerf (1941, p. 7; cf. FAS; A:T-55. Chasen, 1940b, p. 485); mzb, 1. FAS; B:S-11. Larut, Daerah (district); MALAYSIA: WEST Letsok-aw Kyun, BURMA; 1 1°27'-1 1°48'N, MALAYSIA; ca. 4°55'N, 100°47'E; collected 98°10'-98°20'E; collected 26 Jan. 1904 by W. before 1901 by unknown collector (Flower, 1900, L. Abbott; usnm, 1 (skin and mandible only). p. 316); specimens reportedly in Taiping Mu- AUR; A:Bu-20. seum (not seen). FAS; B:WM-5. Leuser, Gunung. See Gunung Leuser Reserve.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 149 Lhokseumawe, ca. 80 km ESE; Sumatra, INDO- Loa Bambam; Borneo: Kalimantan, INDONE- NESIA; ca. 4°55'N, 97°48'E; reported present SIA; 0°29'S, 1 17°02'E; collected 1-15 Jun. 1912 Nov. 1971-Jan. 1973 by local residents (Crock- by H. C. Raven (see Deignan, [1960], p. 267); ett & Wilson, 1980, p. 156). FAS; B:S-10. usnm, 2 (including 1 skull only; third specimen Liang Koeboeng; Borneo: Kalimantan, INDO- listed in field catalog). FAS; CK-49. NESIA; 0°37'N, 1 13°08'E; collected 10-18 Apr. Lo Bon Bon. See Loa Bambam. 1894 by J. Buttikofer (1897, p. 16); rmnh, 2 (see Loeboe; Sumatra, central, INDONESIA; not lo- Jentink, 1897, p. 39, who also lists a third spec- cated, 5°38'N-5°57'S, 95°12'-106°05'E; ob- imen). FAS; C:K-16. served 1877-1879 by J. F. Snelleman (1887, p. Lima Belas Estate, ca. 50-120 m; MALAYSIA: 10). FAS; not mapped. WEST MALAYSIA; ca. 3°46'N, 101°21'E; ob- Loeboek Linggan. See Lubuklinggau. served Jul. 1965-Jun. 1966 by I. S. Bernstein Loeboek Sikaping. See Lubuksikaping. (1967, p. 199). Observed Jan. 1980-May 1981 Logo. See Lengah, Pulau. by J. O. Caldecott (1986b, p. 21). FAS; B:WM- Loho Gringgo. See Ginggo, Teluk. 18. Loka; Pulau Sulawesi, INDONESIA; ca. 5°26'S, Lima Bias Estate. See Lima Belas Estate. 1 19°54'E; introduced, reported present 21 Oct.- Linga Island. See Lingga, Pulau. 15 Nov. 1888 by Malay hunters employed by Lingartjati. See Linggajati. M. Weber (1890a, p. vii; 1890b, p. 102). Sub- Lingga; Borneo, MALAYSIA: SARAWAK; species uncertain; not mapped. 1°21'N, 111°10'E; collected Nov. 1899 by mu- Lokan, Sungai; Borneo, MALAYSIA: SABAH; ca. seum collectors; SMK, 2 (skins only, skulls in- 5°32'N, 1 17°33'E; observed 1969-1970 by P. S. side). FAS; C:Sar-8, Rodman (1991, p. 359). FAS; C:Sab-15. Lingga, Pulau, INDONESIA; 0°02'N-0°22'S, Lomblen, Pulau, Lesser Sunda Islands, INDO- 104°26'-105°00'E; collected 23 Jul. 1899 by W. NESIA; 8°H'-8°35'S, 123°13'-123°55'E; M. L. Abbott (see Miller, 1900, p. 242; 1906c, p. fascicularis reported absent before 1937 by J. J. 284; Kloss, 1903b, p. 54); usnm, 2. FAS; B:SCS- M. F. Symons (Mertens, 1936, p. 319). C:f. 13. Lombok, Pulau, Lesser Sunda Islands, INDO- Linggajati, Cirebon district, 600 m; Java, IN- NESIA; 8°13'-8°58'S, 115°50'-116°43'E; re- DONESIA; 6°52'S, 108°28'E; collected 3 Jan. ported present before 1 848 by H. Zollinger (1847, 1933 by J. J. Menden; amnh, 1. FAS; B:J-23. p. 203). Reported present May-Jul. 1896 by A. Linglung, Pulo. See Lagong, Pulau. Everett (in Hartert, 1896, p. 593). Purchased ca. Lingung, Pulo. See Lagong, Pulau. 1920 by W. G. Wallace; bm(nh), 1 (skull only). Little Andaman Island, INDIA; 10°31'-10°54'N, FAS; CLS-5 through LS-7. 92°22'-92°36'E; primates reported absent be- Lompat, Sungai, ca. 3 km ENE of Kuala Serloh; fore 1903 by G. S. Miller, Jr. (1902b, p. 792; cf. MALAYSIA: WEST MALAYSIA; ca. 3°41'N,

1 1 1 Kloss, [ 928], p. 802; Chaturvedi, 980, p. 34). 102°1 l'E; observed Jul.-Dec. 1977 by D. Oliv- A:h. ers and G. Davies (1979, p. 20). FAS; B:WM- Little Condor Island. See Ben Dam. 15. Little Durian. See Durian-kecil, Pulau. Lompat, Sungai, ca. 3 km W of Kuala Lompat Little Jolly Boy Island, Andaman Islands, INDIA; Post; MALAYSIA: WEST MALAYSIA; ca. 11°31'-11 32'N, 92°36'-92°37'E; primates re- 3°42'N, 102°16'E; observed Jul.-Dec. 1977 by ported absent before 1903 by G. S. Miller, Jr. D. Chivers and G. Davies (1979, p. 20). FAS; (1902b, p. 792; cf. Kloss, [1928], p. 802; Cha- B:WM-15. turvedi, 1980, p. 134). A:f. Lompat, Sungai, ca. 4 km ENE of Kuala Serloh; Little Karimon. See Karimun-kecil, Pulau. MALAYSIA: WEST MALAYSIA; ca. 3°41'N, Little Kretam River. See Kretam Kechil, Sungai. 102°12'E; observed Jul.-Dec. 1977 by D. Chiv- Little Nicobar Island, INDIA; 7°13'-7°25'N, ers and G. Davies (1979, p. 20). FAS; B:WM- 93°37'-93°45'E; observed before 1847 by P. 15. Barbe (1 846, p. 365). Collected 25-27 Feb. 1 90 1 Lompat, Sungai, ca. 4 km W of Kuala Lompat by W. L. Abbott (see Miller, 1902b, p. 751; Post; MALAYSIA: WEST MALAYSIA; ca. Kloss, 1903a, pp. 122, 128); usnm, 3. UMB; 3°42'N, 102°15'E; observed Jul.-Dec. 1977 by A:N-2. D. Chivers and G. Davies (1979, p. 20). FAS; Little Tenasserim River. See Thagyet, Little Ten- B:WM-15. asserim River. Lompat, Sungai, ca. 6 km W of Kuala Lompat

150 FIELDIANA: ZOOLOGY Post; MALAYSIA: WEST MALAYSIA; ca. by C. M. Crockett and W. L. Wilson (1980, p. 3°42'N, 102°14'E; observed Jul.-Dec. 1977 by 156). FAS;B:S-64. D. Chivers and G. Davies (1979, p. 20). FAS; Lubukminturun area; Sumatra, INDONESIA; ca. B:WM-15. 0°52'S, 100°23'E; blood samples taken Jan. Long Ekang; Borneo, MALAYSIA: SARAWAK; 1979-Dec. 1981 by Y. Kawamoto (1982, p. 67). 3°55'N, 1 14°27'E; collected 26 Feb. 1956 by T. FAS; B:S-47. A. Chavasse (see Arnold, 1959, p. 204); fmnh, Lubuksikaping; Sumatra, INDONESIA; OWN, l.FAS;C:Sar-19. 100°10'E; collected 1 Aug. 1930 by E. Jacobson; Long Ikang. See Long Ekang. rmnh, 1 (skull only). FAS; B:S-43. Longo, Pulau. See Longos, Nusa. Lumu Lumu, 6000 ft (= 1830 m); Borneo, MA- Longos, Nusa, Lesser Sunda Islands, INDONE- LAYSIA: SABAH; ca. 6°02'N, 116°35'E; cap- SIA; 8°20'-8°21'S, 120°07'-120°09'E; monkeys tives acquired Jun. 1932 by J. A. Griswold, Jr. reported absent 1969-1973 by W. Auffenberg (1939b, p. 506). FAS; C:Sab-7. (1981, p. 40). C:d. Lungmanis Station, Sungai Segaliud; Borneo, MA- Long Pangian (?= Long Pangean); Borneo: Kali- LAYSIA: SABAH; ca. 5°40'N, 117°45'E; ob- mantan, INDONESIA; 2°42'N, 116°42'E; col- served in 1958 by K. Stott, Jr. (1964, p. 13). lected 24 Apr. 1914 by C. Lumholtz (see Gyld- FAS; C:Sab-16. enstolpe, 1920, pp. 3, 14-15); Zoological Mu- Macarah [?= Moearah] Doewa. See Muaradua. 1 C:K- seum, Christiana, (not examined). FAS; MacRitchie Reservior Nature Reserve; Singapore 38. Island; SINGAPORE; ca. 1°21'N, 103°50'E; re- Long Pelban. See Long Peleben. ported present before 1974 by S. H. Chuang Long Peleben; Borneo: Kalimantan, INDONE- (1973, p. 3). FAS; B:SCS-7. SIA; 2°47'N, 116°35'E; collected 30 Jun. 1935 Madgalengka. See Majalengka. V. von Plessen cf. by (1936, p. 100; Stresemann, Madura, Pulau, INDONESIA; 6°52'-7°15'S, 1 1; mzb, 1. Collected 1938, p. 10); amnh, Aug. 1 1 2°4 1 '-1 1 4°07'E; reported present before 1 90 1 1956 P. Pfeffer; 1 (skull by mnhn, only). FAS; by A. G. Vorderman (1900, p. 143). FAS; C:J- CK-38. 39. Pulau. See Lanta Ko. Lontar, Yai, Maenam Chao Phaya. See Chao Phraya, Mae Nam, Lopburi. See Sarn Pra Kara, Lopburi. below Bangkok. Lower Taipo Road, Kowloon Reservoir Area; Mae Nam Khwae Noi. See Khwae Noi, Mae Nam. ca. UNITED KINGDOM: HONG KONG; Mahakam, Sungai, left bank, 1 km above Sebulu; ob- 22°21'N, 114°09'E; introduced population, Borneo: Kalimantan, INDONESIA; ca. 0°16'S, 1980 198 1 C. served Jul.-Aug and Jul.-Aug. by 1 1 7°00'E; observed 29-30 Aug. 1983 by S. Azu- K. L. Southwick H. Southwick and (1983, p. ma, A. Suzuki, and Y. Ruhiyat (1984, p. 48). 19). Subspecies uncertain; not mapped. FAS; C:K-49. Luang Ko. See Rang Yai, Ko. Mahakam, Sungai, left bank, 4 km above Sebulu; Lubuklinggau, ca. 15 km N; Sumatra, INDO- Borneo: Kalimantan, INDONESIA; ca. 0°17'S, NESIA; ca. 3°10'S, 102°50'E; observed Nov. 1 16°58'E; observed June 1972 by C. L. Darsono 1971-Jan. 1973 by C. M. Crockett and W. L. (see Wilson & Wilson, 1975, p. 257). Observed Wilson (1980, p. 156). FAS; B:S-64. 29-30 Aug. 1983 by S. Azuma, A. Suzuki, and Lubuklinggau, ca. 40 km NNW; Sumatra, IN- Y. Ruhiyat (1984, p. 48). FAS; QK-49. DONESIA; ca. 2°58'S, 102°43'E; observed Nov. Mahakam, Sungai, north bank, above Samarinda; 1971-Jan. 1973 by C. M. Crockett and W. L. Borneo: Kalimantan, INDONESIA; ca. 0°30'S, Wilson (1980, p. 156). FAS; B:S-65. 1 17°05'E; collected 31 May 1912 by H. C. Ra- Lubuklinggau, ca. 60 km SE; Sumatra, INDO- ven (field catalog); usnm, 1. FAS; QK-49. NESIA; ca. 3°40'S, 103°15'E; observed Nov. Mahakam, Sungai, right bank, 2 km below Sebulu; 1971-Jan. 1973 by C. M. Crockett and W. L. Borneo: Kalimantan, INDONESIA; ca. 0°17'S, Wilson (1980, p. 156). FAS; B:S-72. 1 1 7°00'E; observed 29-30 Aug. 1 983 by S. Azu- Lubuklinggau, sea level; Sumatra, INDONESIA; ma, A. Suzuki, and Y. Ruhiyat (1984, p. 48). 3°18'S, 102°52'E; collected 21 Nov. 1933 by J. FAS; QK-49. J. Menden; amnh, 1. FAS; B:S-64. Mahakam, Sungai, right bank, 8 km above Sebulu; Lubuklinggau vicinity; Sumatra, INDONESIA; ca. Borneo: Kalimantan, INDONESIA; ca. 0°18'S, 3°18'S, 102°52'E; observed Nov. 1971-Jan. 1973 1 16°57'E; observed 29-30 Aug. 1983 by S. Azu-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 151 ma, A. Suzuki, and Y. Ruhiyat (1984, p. 48). lected 4-10 May 1925 by P. F. Franck and Den- FAS; CK-49. in (see Dammerman, 1926a, p. 91); mzb, 3. FAS; Mahasarakham. See Kosumphi Forest Park. GLS-23. Majalengka, Cirebon district, 600 m; Java, IN- Mapor, Pulau. See Mentigi. DONESIA; 6°50'S, 108°13'E; collected 22 Dec. Marah. See Merah. 1932 by J. J. Menden; amnh, 3. FAS; B:J-22. Maratua, Pulau, INDONESIA; 2°10'-2°19'N, Malabar, Pengalengan district, 6000 ft (= 1 800 m); 1 1 8°33'-l 1 8°39'E; collected 2 1 May 1 9 1 3 by H. Java, INDONESIA; 7°07'S, 107°36'E; collected C. Raven (in Riley, 1930, p. 2); usnm, 4. TUA; in 1937 by D. P. Bosscha Erdbrink; zluu, 1 CK-41. (skull only). FAS; B:J-6. Margo-Molio, Gunung. See Margomulio, Gun- Malacca. See Melaka. ung. Malakka. See Melaka. Margomulio, Gunung, Gunung Kelud group, Ke- Malang. See Wonokojo (?= Wonokerto), Dampit diri district; Java, INDONESIA; 7°55'S, district, southern Malang region. 1 1 2°1 5'E; collected Aug. 1 909 by V. Arnim; zsbs, Malawali, Pulau, MALAYSIA: SABAH; 7°01'- 1 (skull only). FAS; C:J-33. 7°05'N, 117°16'-117°22'E; observed Feb.-Jun. Masalembu Besar, Pulau, INDONESIA; 5°32'- 1991 by Shukor Md. Nor (pers. comm., 20 Jul. 5°36'S, 114°24'-114°27'E; monkeys reported 1991). FAS; C:Sab-ll. absent 4-6 Dec. 1 907 by W. L. Abbott (in Lyon, Malaya. See MALAYSIA: WEST MALAYSIA. 1911, p. 61). C:b. MALAYSIA: WEST MALAYSIA; 1°16'-6°43'N, Mata, Pulo. See Matak, Pulau. 100°07'-104°18'E; collected date unknown by Matak, Pulau, INDONESIA; 3°13'-3°23'N,

unknown collector; usnm, 1 . FAS; not mapped. 106°14'-106°1 8'E; observed 24 Aug.-5 Sep. 1 899 Maliangin Besar, Pulau, MALAYSIA: SABAH; by W. L. Abbott (in Miller, 1900, p. 245; cf. 7°04'-7°05'N, 1 17°02'-1 17°03'E; observed Feb.- Kloss, 1903b, p. 72). FAS; B:SCS-30. Jun. 1991 by Shukor Md. Nor (pers. comm., 20 Matasiri, Pulau, east coast; INDONESIA; ca. Jul. 1991). FAS; C:Sab-9. 4°47'S, 115°50'E; collected 8 Dec. 1907 and Maliwun. See Pakchan River, near Maliwun. 1907-1908 by W. L. Abbott (field catalog; in Maman; Pulau Sumbawa, Lesser Sunda Islands, Lyon, 1911, p. 61); usnm, 2 (including 1 skull INDONESIA; ca. 8°37'S, 1 17°29'E; blood sam- only). FAS; C:K-53. ples taken Jul. 1981-Jan. 1982 by Y. Kawa- Maung district. See Phu Phan, Maung district. o o moto, Tb. M. Ischak, and J. Supriatna (1982a, Mauritius Island, MAURITIUS; 19 59'-20 31'S, p. 58). FAS; C:LS-8. 57°18'-57°48'E; introduced population, ob- Mandalay. See Irrawaddy River, right bank, served 19 Apr. 1753 by M. l'Abbe de la Caille northwest of Mandalay. (1763, p. 230). Collected before 1 862 by E. New- Mandau, Sungai; Sumatra, INDONESIA; ca. ton; bm(nh), 1. Collected 20 Jul. 1875 by J. Au- 0°48'N, 101°47'E; observed ca. 1907 by M. debert; rmnh, 1 (skin only, skull inside). Col- Moszkowski (1909, p. 143, map following p. lected before 1 9 1 by Franco-British Commis- 192). FAS;B:S-42. sion; bm(nh), 1. Collected before 1956 by un- . See Bantargebang. known collector; bm(nh), 1 (skull only). Observed Manggis, Gunung Kelud, Kediri district; Java, IN- 1977-1984 by R. W. Sussman and I. Tattersall DONESIA; 7°49'S, 1 12°14'E; collected in 1909 (1986, p. 34). Blood samples taken Oct. 1989 by V. Arnim; zsbs, 9 (skulls only). FAS; C:J-33. by M. Kondo, Y. Kawamoto, K. Nozawa, K. Mangiatan, Pulau, Lesser Sunda Islands, INDO- Matsubayashi, T. Watanabe, O. Griffiths, and NESIA; 8°33'S, 1 1 9°4 1 'E; reported present 1 969- M.-A. Stanley (1993, p. 168). Subspecies un-

1 1 973 by W. Auffenberg (1 98 , p. 40). FAS; C:LS- certain; not mapped. 13. Mbarapu, Nusa, Komodo area, Lesser Sunda Is- Maninjau; Sumatra, INDONESIA; 0°18'S, ands, INDONESIA; 8°37'S, 1 19°32'E; monkeys 100°14'E; collected Mar.-Sep. 1888 by M. We- reported absent 1969-1973 by W. Auffenberg

ber (1890a, p. iii; 1890b, p. 102); museum un- (1981, p. 40). C:e. known, 1 (skeleton only; not seen). FAS; B:S- Mbura; Pulau Flores, Lesser Sunda Islands, IN- 45. DONESIA; 8°34'S, 1 19°52'E; collected 17 Oct. Mansalar, Pulo. See Mursala, Pulau. 1929 by J. K. de Jong; mzb, 4. FAS; GLS-13. Mao Marroe, 450 m; Pulau Sumba, Lesser Sunda Medan, ca. 20 km N; Sumatra, INDONESIA; ca. Islands, INDONESIA; 9°58'S, 120°30'E; col- 3°47'N, 98°42'E; observed Nov. 1971-Jan. 1973

152 FIELDIANA: ZOOLOGY by C. M. Crockett and W. L. Wilson (1980, p. 5°38'N-5°57'S, 95°12'-106°05'E; collected 25 156). FAS;B:S-21. Jul. 1928 by W. Comberg; zmb, 1 (skin only). Medan, ca. 75 km NW; Sumatra, INDONESIA; FAS; not mapped. ca. 4°04'N, 98°09'E; observed Nov. 1971-Jan. Mendawai, Sungai. See Katingan, Sungai. 1973 by C. M. Crockett and W. L. Wilson (1980, Mendit; Java, INDONESIA; ca. 8°00'S, 1 12°50'E; p. 156). FAS;B:S-19. clinical examination in 1980 by K. Matsubay- Medan, ca. 100 km SSW; Sumatra, INDONESIA; ashi and D Sajuthi (1981, p. 48). FAS; C:J-36. ca. 2°45'N, 98°1 8'E; reported present before 1 98 1 Mengjatan, Pulau. See Mangiatan, Pulau. by H. D. Rijksen (Crockett & Wilson, 1980, p. Mensuda Bay; Pulau Karimun, northeast, Kepu- 156). FAS;B:S-26. lauan Riau, INDONESIA; 1°07'N, 103°23'E; Medan, ca. 105 km SSE; Sumatra, INDONESIA; collected 29 May 1903 by W. L. Abbott (field ca. 2°40'N, 98°42'E; reported present Nov. 197 1- catalog; cf. Miller, 1906c, p. 277); usnm, 1. FAS; Jan. 1 973 by local residents (Crockett & Wilson, B:SCS-17. 1980, p. 156). FAS; B:S-28. Mentarang, Sungai. See Sungai Kayan-Sungai Medan, forest near; Sumatra, INDONESIA; ca. Mentarang Nature Reserve. 3°35'N, 98°40'E; collected in 1906 by C. Brue- Mentigi; Pulau Mapur, west, Kepulauan Riau; IN- gekzsBS, 1. FAS; B:S-21. DONESIA; ca. 1°00'N, 104°48'E; collected 6 Jun. Medan, near sea level; Sumatra, INDONESIA; 1915 by H. C. Robinson (1916, p. 62); zrc, 1. 3°35'N, 98°40'E; collected in 1905 by C. Brue- FAS; B:SCS-9.

gel, zsbs, 1 (skin only). Collected 1906-1910 by Mentoko Research Center, Kutai National Park, C. Widnmann; zsbs, 78 (including 41 skins only, ca. 50 m; Borneo: Kalimantan, INDONESIA; 3 skulls only). Collected 21 May 1939 by F. 0°24'N, 11 7°06'E; observed Jun. 1982-Sep. 1983 Ulmer (in Miller, 1942, p. 127; cf. Schauensee by L. Berenstain (1986, p. 258). FAS; C:K-47.

& Ripley, 1 940a, p. 3 1 6); ansp, 1 . Collected date Menyala, Sungai; MALAYSIA: WEST MALAY- unknown by Mrs. P. W. Cremer-Jansen; mcz, 1 SIA; ca. 2°29'N, 101°55'E; collected in 1972 by (skull only). FAS; B:S-21. unknown collector; Department of Anatomy, Medan-Siantar; Sumatra, INDONESIA; ca. Cambridge, 1 (in fluid; not seen, data from P. 3°35'N, 98°40'E; observed Mar.-May 1939 by H. Napier, 1981, p. 14). FAS; B:WM-24. F. A. Ulmer, Jr. (in Miller, 1942, p. 127). FAS; Me Ping River. See Ping, Mae Nam. B:S-21. Merah; Borneo: Kalimantan, INDONESIA; Medan vicinity; Sumatra, INDONESIA; ca. 0°50'N, 116°48'E; collected 25 Nov. 1925 by 3°35'N, 98°40'E; reported present Nov. 1971- Madzoed; mzb, 1. FAS; C:K-43. Jan. 1 973 by local residents (Crockett & Wilson, Mergui; BURMA; 12°26'N, 98°36'E; collected be- 1980, p. 156). FAS; B:S-21. fore 1857 by Prof. Oldham; bm(nh), 1 (skull Melaka; MALAYSIA: WEST MALAYSIA; only). Collected 25 Dec. 1881 by J. Anderson, 2°12'N, 102°15'E; collected 1818-1863 by P. zsi, 1 (skin only). Collected 1 Mar. 1914 by G. Diard; rmnh, 1 (skin only). Collected before 1 849 C. Shortridge (in Wroughton, 1915, p. 696); by M. de Montigny; mnhm, 2 (skins only, skulls bnhs, 1. AUR; A:Bu-18. inside). Reported present before 1852 (Anony- Meru, Gunung; Sumatra, INDONESIA; ca. 1°06'S, mous, 1851, p. 444). Collected before 1876 by 100°27'E; 2 troops observed 21 Jun.-28 Oct. Gerrard; bm(nh), 1. Collected in 1875 by 1980 by N. Koyama, A. Asuan, and N. Natsir

Schneider; rmnh, 2 (including 1 skin only, skull (1981, p. 1). Observed 27 Aug. 1984-5 Jan. 1985 inside). FAS; B:WM-26. by N. Koyama (1985, p. 106). FAS; B:S-47. Melinau Gorge, rock shelter near RGS Camp 5, Meru Betiri Nature Park; Java, INDONESIA; ca. Mulu National Park, ca. 200 m; Borneo, MA- 8°28'S, 1 1 3°48'E; observed 24 Aug.-1 4 Sep. 1 97 1 LAYSIA: SARAWAK; 4°08'N, 114°54'E; col- by A. Hoogerwerf (1974, p. 13). Observed in lected 24 Aug. 1977 by Lord Medway; bm(nh), 1976 by J. Seidensticker (1983, p. 324). FAS; 1 (mandibular fragment only). FAS; C:Sar-21. CJ-35. Meloewak. See Goenoengsetan-Meloewak. Meulaboh, ca. 35 km NW; Sumatra, INDONE- Mempura; Sumatra, INDONESIA; 0°43'N, SIA; ca. 4°21'N, 95°54'E; observed Nov. 1971- 101°59'E; collected ca. 1907 by M. Moszkowski Jan. 1973 by C. M. Crockett and W. L. Wilson (1909, p. 151); zmb, 2 (skulls only). FAS; B:S- (1980, p. 156). FAS; B:S-2. 42. Meulaboh, ca. 60 km NW; Sumatra, INDONE- Menam; Sumatra, INDONESIA; not located, SIA; ca. 4°30'N, 95°44'E; observed Nov. 1971-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 153 Jan. 1973 by C. M. Crockett and W. L. Wilson Moro Batin; Sumatra, INDONESIA; 5°26'S, (1980, p. 156). FAS; B:S-3. 105°06'E; observed Nov. 1971-Jan. 1973 by C. Meulaboh vicinity; Sumatra, INDONESIA; ca. M. Crockett and W. L. Wilson (1980, p. 156). 4°09'N, 96°08'E; reported present Nov. 1971- FAS; B:S-82. Jan. 1 973 by local residents (Crockett & Wilson, Moro Batin, south-southeast; Sumatra, INDO- 1980, p. 156). FAS;B:S-1. NESIA; ca. 5°35'S, 105°10'E; observed Nov. Mewong River. See Wong, Nam Mae. 1971-Jan. 1973 by C. M. Crockett and W. L. Mibya Kyun, mouth of Tavoy River, BURMA; Wilson (1980, p. 156). FAS; B:S-82. 13°36'N, 98°12'E; collected 23 Apr. 1936 by H. Moro Besar. See Durian, Pulau. C. Smith; bm(nh), 2. AUR; A:Bu-15. Moro Kechil. See Durian-kecil, Pulau. Middle Andaman Island, INDIA; 12°05'-12°54'N, Moscos Island Game Sanctuary; , 92°42'-93°00'E; primates reported absent be- BURMA; 13°47'-14°27'N, 97°47'-97°56'E; re- fore 1928 by C. B. Kloss ([1928], p. 802; cf. ported present 1938-1939 by H. A. Maxwell Chaturvedi, 1980, p. 134). A:b. (see Yin, 1954, p. 281). AUR; A:Bu-12. Miri; Borneo, MALAYSIA: SARAWAK; 4°23'N, Mota, Gili. See Motang, Gili. 113°59°E; collected Jun. 1890 by E. Hose and Motang, Gili, Lesser Sunda Islands, INDONESIA; C. Hose; mcz, 1 (skin only, skull inside). Col- 8°48'-8°49'S, 119°46'-119°48'E; monkeys re- lected date unknown by unknown collector; ported absent 1969-1973 by W. Auffenberg Duckworth Laboratory of Physical Anthropol- (1981, p. 40). C:e. ogy, Cambridge, 1 (skull and limb bones only; Moulmein. See Ye Forest, Ataran district, Moul- not seen, data from P. H. Napier, 1981, p. 14). mein region. FAS; C:Sar-18. Moyo, Pulau, Lesser Sunda Islands, INDONESIA; Miri, Sungai; Borneo, MALAYSIA: SARAWAK; 8°09'-8°23'S, 1 17°29'-1 17°42'E; blood samples ca. 4°23'N, 113°58'E; collected date unknown taken Jan. 1979-Dec. 1981 by Y. Kawamoto by unknown collector; Duckworth Laboratory (1982, p. 67). FAS; CLS-9. of Physical Anthropology, Cambridge, 1 (skel- Mrengoo Island. See Myengun Kyun. eton only; not seen, data from P. H. Napier, Mt. Setan-Meloewak. See Goenoengsetan-Mel- 1981, p. 14). FAS; C:Sar-18. oewak. Miri district, Batang Baram; Borneo, MALAY- Mt. Sontra. See Sontra Peak. SIA: SARAWAK; ca. 4°23'N, 113°59'E; col- Muaradua, Palembang district, 100 m; Sumatra, lected Jan. 1895 by C. Hose; NHMBe, 1 (skin INDONESIA; ca. 4°32'S, 1 04°0 5 'E; collected 10- only). FAS;C:Sar-18. 17 Jun. 1934 by J. J. Menden; amnh, 8. FAS; Misa, Pulau, Lesser Sunda Islands, INDONESIA; B:S-76. 8°31'S, 119°45'E; monkeys reported absent Muaratewe, Sungai Barito, 65 ft (= 20 m); Borneo: 1969-1973 by W. Auffenberg(1981, p. 40). C:d. Kalimantan, INDONESIA; 0°57'S, 114°53'E; Mobur, Pulo. See Mubur, Pulau. collected 21 Sep. 1909 by G. C. Shortridge; Moeara Tewech. See Kampong Hadjak, Muara- bm(nh), 3 (including 1 skin only). FAS; C:K-32. tewe district. Muaratewe vicinity; Borneo: Kalimantan, IN- Moera Tewe. See Kampong Hadjak, Muaratewe DONESIA; ca. 0°57'S, 1 14°53'E; observed 1979- district. 1986 by J. Marshall, D. J. Chivers, and K. M. Moeria, Mt. See Pangonan, Gunung Muria. Burton (see Chivers & Burton, [1991], p. 141). Moeriah. See Pangonan, Gunung Muria. FAS; C.K-32. Mokut. See Kampong Mukut. Muara Teweh. See Muaratewe. Monos; Pulau Karimun, Kepulauan Riau; IN- Muara Tua, Pulo. See Maratua, Pulau. DONESIA; 1°08'N, 103°23'E; collected 29-30 Mubur, Pulau, INDONESIA; 3°18'-3°23'N, Aug. 1908 by H. C. Robinson (in Thomas & 106°10'-106°14'E; observed 24 Aug.-5 Sep. 1899 Wroughton, 1909b, pp. 105, 107; cf. H. C. Rob- by W. L. Abbott (in Miller, 1900, p. 245; cf. inson & Kloss, 1914, p. 394) and E. Seimund; Kloss, 1903b, p. 72). FAS; B:SCS-30. bm(nh), 2. FAS; B:SCS-17. Mugitriman, Hak Pegusahaan Hutan (HPH), Jam- Morib; MALAYSIA: WEST MALAYSIA; ca. bi; not located; Sumatra, INDONESIA; ob- 3°50'N, 100°50'E; examined for malaria before served Mar. 1989 by M. Bismark (1992, p. 11). 1994 by A. N. Rain, J. W. Mak, and R. Zamri FAS; not mapped. (1993, p. 386). FAS; B:WM-17. Muk, Ko, THAILAND; 7°22'N, 99° 1 8'E; observed

154 FIELDIANA: ZOOLOGY 4-8 Jan. 1917 by H. C. Robinson and E. Sei- Murung, Sungai, right bank, 1 km below mouth mund (H. C. Robinson, 1917, p. 134). FAS; of Sungai Beriwit; Borneo: Kalimantan, IN- A:T-56. DONESIA; ca. 0°09'S, 1 14°15'E; observed 4-7 Mu Ko Phetra National Park; THAILAND; ca. Sep. 1986 by D. J. Chivers and K M. Burton 6°57'N, 99°42'E; observed 21-23 Dec. 1987 by ([1991], p. 143). FAS; CK-19. R. Boonratana also observed (1988, pp. 76, 81); Murung, Sungai, right bank, 1 km below mouth on unspecified adjacent islands. FAS; A:T-57. of Sungai Turusan; Borneo: Kalimantan, IN- See Mulu National Park. Mulu, Gunong. Gunong DONESIA; ca. 0°1 l'S, 1 14°09'E; observed 7-8 2000 ft 600 Mulu, Gunung, (= m); Borneo, MA- Sep. 1986 by D. J. Chivers and K M. Burton col- LAYSIA: SARAWAK; 4°03'N, 114°56'E; ([1991], p. 143). FAS; CK-19. lected Oct. 1893 by C. Hose; bm(nh), 1 (skull Murung, Sungai, right bank, 5 km above mouth FAS; C:Sar-21. only). of Sungai Turusan; Borneo: Kalimantan, IN- Muntia, Pulau. See Muk, Ko. DONESIA; ca. 0°09'S, 114°12'E; observed 4-8 Muntok; Pulau Bangka, INDONESIA; 2°04'S, Sep. 1986 by D. J. Chivers and K M. Burton 105°10'E; collected in 1900 by A. A. W. Hu- ([1991], p. 143). FAS; CK-19. brecht([1895], pp. 33, 88; cf. Kohlbrugge, 1902, Musi, Air, ca. 1 2 km below Palembang; Sumatra, p. 322; Zuckerman, [1933], p. 1062); zluu, un- INDONESIA; ca. 2°58'S, 104°51'E; observed specified portion of Pulau Bangka collection (see Nov. 1971-Jan. 1973 by C M. Crockett and W. Bangka, Pulau). FAS; B:SCS-18. L. Wilson (1980, p. 156). FAS; B:S-61. Mursala, Pulau, INDONESIA; 1°37'-1°42'N, Musi, Air, ca. 15 km above mouth of Batang Har- 98°27'-98°36'E; collected 7-10 Mar. 1 902 by W. ileko; Sumatra, INDONESIA; ca. 2°54'S, L. Abbott (see Miller, 1903a, p. 438); usnm, 3 103°58'E; observed Nov. 1971-Jan. 1973 by C (including 1 skull only). FAS; B:IO-ll. M. Crockett and W. L. Wilson (1980, p. 156). Murung, Sungai, left bank, 1 km below mouth of FAS; B:S-62. Sungai Danau Tolong; Borneo: Kalimantan, IN- Musi, Air, ca. 1 5 km above Sumatra, DONESIA; ca. 0°10'S, 114°13'E; observed 4-7 Palembang; INDONESIA; ca. 3°02'S, 104°37'E; observed Sep. 1986 by D. J. Chivers and K. M. Burton Nov. 1971-Jan. 1973 by C M. Crockett and W. ([1991], p. 143). FAS; CK-19. L. Wilson (1980, p. 156). FAS; B:S-61. Murung, Sungai, left bank, 2 km above mouth of ca. 24 below Sungai Beriwit; Borneo: Kalimantan, INDO- Musi, Air, km Palembang; Sumatra, ca. observed NESIA; ca. 0°09'S, 1 14°17'E; observed 4-7 Sep. INDONESIA; 2°53'S, 104°54'E; Nov. 1971-Jan. 1973 M. Crockett and W. 1 986 by D. J. Chivers and K M. Burton ([1991], by C L. Wilson B:S-61. p. 143). FAS; CK-19. (1980, p. 156). FAS; Murung, Sungai, left bank, 2 km above mouth of Musi, Air, ca. 30 km above mouth of Batang Har- Sungai Turusan; Borneo: Kalimantan, INDO- ileko; Sumatra, INDONESIA; ca. 2°53'S, NESIA; ca. 0°1 l'S, 1 14°1 l'E; observed 4-8 Sep. 103°51'E; observed Nov. 1971-Jan. 1973 by C 1986 by D.J. Chivers and KM. Burton ([1991], M. Crockett and W. L. Wilson (1980, p. 156). p. 143). FAS; QK-19. FAS; B:S-62. Murung, Sungai, left bank, 2 km below mouth of Musi, Air, ca. 30 km above Palembang; Sumatra, Sungai Turusan; Borneo: Kalimantan, INDO- INDONESIA; ca. 3°04'S, 104°29'E; observed NESIA; ca. 0°12'S, 1 14°08'E; observed 7-8 Sep. Nov. 1971-Jan. 1973 by C M. Crockett and W. 1986 by D.J. Chivers and KM. Burton ([1991], L. Wilson (1980, p. 156). FAS; B:S-61. p. 143). FAS; CK-19. Musi, Air, ca. 36 km below Palembang; Sumatra, Murung, Sungai, left bank, 8 km above Muara- INDONESIA; ca. 2°47'S, 104°56'E; observed juloi; Borneo: Kalimantan, INDONESIA; ca. Nov. 1 97 1-Jan. 1 973 by C M. Crockett and W. 0°10'S, 1 14°08'E; observed 7-8 Sep. 1986 by D. L. Wilson (1980, p. 156). FAS; B:S-61. J. Chivers and K M. Burton ([1991], p. 143). Musi, Air, ca. 45 km above mouth of Batang Har- FAS; CK-19. ileko; Sumatra, INDONESIA; ca. 2°49'S, Murung, Sungai, left bank, at mouth of Sungai 103°44'E; observed Nov. 1971-Jan. 1973 by C Beriwit; Borneo: Kalimantan, INDONESIA; ca. M. Crockett and W. L. Wilson (1980, p. 156). 0°10'S, 1 14°15'E; observed 4-7 Sep. 1986 by D. FAS; B:S-62. J. Chivers and K M. Burton ([1991], p. 143). Musi, Air, ca. 48 km below Palembang; Sumatra, FAS; CK-19. INDONESIA; ca. 2°41'S, 104°56'E; observed

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 155 Nov. 1971-Jan. 1973 by C. M. Crockett and W. 98°29'E; collected 4 Feb. 1 9 1 8 by local collectors L. Wilson (1980, p. 156). FAS; B:S-61. employed by H. C. Robinson and C. B. Kloss Musi, Air, ca. 50 km above Palembang; Sumatra, (1919, p. 87);zrc, 1. AUR/FAS/MUL; A:T-52. INDONESIA; ca. 2°59'S, 104°19'E; observed Nakhon Si Thammarat; THAILAND; 8°26'N, Nov. 1971-Jan. 1973 by C. M. Crockett and W. 99°58'E; collected 27 Mar. 1924 and 19-29 Sep. L. Wilson (1980, p. 156). FAS; B:S-62. 1926 by H. M. Smith (see Riley, 1938, p. 10); Musi, Air, ca. 60 km above mouth of Batang Har- usnm, 2 (including 1 skull only); zrc, 3. AUR/ ileko; Sumatra, INDONESIA; ca. 2°45'S, FAS/MUL; A:T-63. 103°39'E; observed Nov. 1971-Jan. 1973 by C. Nakhorn Sawan. See Wat Khao Noh. M. Crockett and W. L. Wilson (1980, p. 156). Nakon Sritammarat. See Nakhon Si Thammarat. FAS; B:S-62. Nam Kam, Koh. See Nang Kham, Ko. Musi, Air, ca. 60 km below Palembang; Sumatra, Nanas, Bukit, Kuala Lumpur; MALAYSIA: WEST INDONESIA; ca. 2°35'S, 104°56'E; observed MALAYSIA; 3°09'N, 101°42'E; collected 11 Nov. 1971-Jan. 1973 by C. M. Crockett and W. Nov. 1913, 6 Jun. 1938, and date unknown by L. Wilson (1980, p. 156). FAS; B:S-61. museum collectors; bm(nh), 3. Observed Mar.- Musi, Air, ca. 70 km above Palembang; Sumatra, Jun. 1966 by I. S. Bernstein (1966, p. 1559; INDONESIA; ca. 3°03'S, 104°10'E; observed 1968a, p. 121). Observed Aug.-Dec. 1970 by C. Nov. 1971-Jan. 1973 by C. M. Crockett and W. H. Southwick and F. C. Cadigan, Jr. (1972, p. B:S-62. L. Wilson (1980, p. 156). FAS; 11). FAS;B:WM-19. ca. 72 km below Musi, Air, Palembang; Sumatra, Nancowry Island, Nicobar Islands, INDIA; 7°55'- ca. observed INDONESIA; 2°28'S, 104°56'E; 8°02'N, 93°30'-93°34'E; primates reported ab- Nov. 1 97 1-Jan. 1 973 C. M. Crockett and W. by sent before 1903 by G. S. Miller, Jr. (1902b, p. L. Wilson B:S-61. (1980, p. 156). FAS; 792; cf. Kloss, 1903a, p. 1 14). A:l. ca. 75 km above mouth of Har- Musi, Air, Batang Nang Kham, Ko; THAILAND; ca. 7°20'N, ileko; Sumatra, ca. 2°44'S, INDONESIA; 100°22'E; collected 12 Apr. 1899 by R. Evans 103°32'E; observed Nov. 1971-Jan. 1973 by C. and F. F. Laidlaw (see P. H. Napier, 1981, p. M. Crockett and W. L. Wilson (1980, p. 156). 17; Bonhote, [1901], p. 870); University Mu- FAS; B:S-62. seum of Zoology, Cambridge, 1 (skull only, not Musi, Air, ca. 84 km below Palembang; Sumatra, seen). FAS; A:T-66. INDONESIA; ca. 2°22'S, 104°54'E; observed Nanga Look, near Nggoer; Pulau Flores, Lesser Nov. 1 97 1-Jan. 1973 by C. M. Crockett and W. Sunda Islands, INDONESIA; ca. 8°42'S, L. Wilson (1980, p. 156). FAS; B:S-61. 119°48'E; observed 1969-1973 by W. Auffen- Musi, Air, ca. 85 km above Palembang; Sumatra, berg (1981, p. 242). FAS; GLS-13. INDONESIA; ca. 2°57'S, 104°05'E; observed Nankauri Island. See Nancowry Island. Nov. 1971-Jan. 1973 by C. M. Crockett and W. NatunaBesar, Pulau, INDONESIA; 3°37'^°13'N, L. Wilson (1980, p. 156). FAS; B:S-62. 107°59'-108°25'E; collected 17 Oct. 1893 by A. Musi, Air, ca. 90 km above mouth of Batang Har- Everett (in Thomas & Hartert, 1894, p. 654; cf. ileko; Sumatra, INDONESIA; ca. 2°43'S, W. Rothchild, 1894, p. 468); bm(nh), 1 (skin 103°25'E; observed Nov. 1971-Jan. 1973 by C. only, skull inside). Collected Aug. 1 894 and Jan. M. Crockett and W. L. Wilson (1980, p. 156). 1896 by C. Hose and E. Hose; bm(nh), 1 (skin FAS; B:S-62. only, skull inside); mcz, 2 (skins only, skulls in- Musi, Air, near Palembang; Sumatra, INDONE- side). FAS; B:SCS-32, SCS-33. SIA; ca. 2°59'S, 104°45'E; observed Nov. 1971- Naunchik. See Chik Jan. 1973 by C. M. Crockett and W. L. Wilson Nong region. State. See Chik region. (1980, p. 156). FAS;B:S-61. Nawngchik, Nong Mutis, Gunung, 2100 m; Pulau Timor, Lesser Negara, Taman; MALAYSIA: WEST MALAY- ca. Sunda Islands, INDONESIA; 9°34'S, 124°14'E; SIA; ca. 4°42'N, 102°28'E; reported present K. B. Khan collected 28 Feb.-22 Mar. 1932 by G. Stein; 1986 by M. M. (1988, p. 267). FAS; B:WM-11. mzb, 1; zmb, 2. FAS; C.LS-28. MyengunKyun, BURMA; 19°50'-20°05'N, 92°55'- Negris Island. See Hainggyi Kyun. 93°02'E; collected in 1 847 by S. R. Tickell (1 854- Ngarai Sianok, Bukit Tinggi; Sumatra, INDO- 1875, p. [18]); museum unknown (not seen). NESIA; not located; observed Feb. 1990 by M. AUR; A:Bu-2. Bismark (1992, p. 11). FAS: not mapped. Naka Yai, Ko, THAILAND; 8°03'-8°04'N, 98°28'- Ngawi; Java, INDONESIA; 7°24'S, 1 1 1°26'E; col-

156 FIELDIANA: ZOOLOGY lected 1887-1941 by E. Dubois; rmnh, 1 (skull 101°12'E; collected 17 Sep. 1901 by N. Annan- only). FAS; OJ-30. dale and H. C. Robinson (1903, p. xxxiii; cf. Nggoer. See Nanga Look, near Nggoer. Bonhote, 1903, p. 3); museum unknown, 1 (not Nguwal, Pulau, Kepulauan Riau, INDONESIA; seen). FAS; A:T-68. 0°38'-0°40'N, 104°13'-104°15'E; collected 3 Jan. Nongkok. See Ban Nong Kok. 1925 by F. N. Chasen (1925, p. 93); zrc, 6. FAS; North Andaman Island, INDIA; 12°53'-13°35'N, B:SCS-11. 92°48'-93°05'E; primates reported absent be- Niah Caves; Borneo, MALAYSIA: SARAWAK; fore 1928 by C. B. Kloss ([1928], p. 802; cf. 3°49'N, 1 13°47'E; reported present before 1959 Chaturvedi, 1980, p. 134). A:a. by Lord Medway (1958, p. 634). FAS; C:Sar- North Borneo. See Sabah. 16. North Natuna Island. See Laut, Pulau. Niah National Park; Borneo, MALAYSIA: SA- North Pagi Island. See Pagai Utara, Pulau. RAWAK; ca. 4°00'N, 114°00'E; reported as Ntori, Sape district, 420 m; Pulau Sumbawa, Less- probably present before 1983 by G. Davies er Sunda Islands, INDONESIA; 8°33'S, ([1983], p. 148). FAS; C:Sar-17. 118°52'E; collected 2 Jun. 1927 by Sunda-Ex- Nias, Pulau, INDONESIA; 0°33'-l°32'N, 97°03'- pedition Rensch (see Mertens, 1930, p. 136); 97°56'E; observed before 1864 by J. T. Nieu- zmb, 1. FAS;C:LS-12. wenhuisen and H. C. B. von Rosenberg (1863, Nu Pau. See Nang Kham, Ko. p. 19). Collected before 1890 by E. Modigliani Nuri Valley; MALAYSIA: WEST MALAYSIA;

( 1 889, p. 239); museum unknown, 1 8 specimens ca. 2°34'N, 102°15'E; captive obtained ca. 1953 (not seen). Collected Nov. 1 904 by W. L. Abbott by J. F. B. Edeson and D. G. Davey (1953, p.

(cf. Lyon, 1916, p. 458); zrc, 1 (skull not ex- 259). FAS; B:WM-25. amined). Reported present before 1906 by T. Nusa Mbarapu. See Mbarapu, Nusa. Willink (1905, p. 175). Collected date unknown Nusa Pinda. See Pinda, Nusa. by H. Raap and R. Knuth; zmb, 2 (skulls only). Nusa Rondong. See Rondong, Nusa. Reported present Nov. 1971-Jan. 1973 by local Nusa Vadju. See Vadju, Nusa. residents (Crockett & Wilson, 1980, p. 156). Nyalas; MALAYSIA: WEST MALAYSIA; 2°26'N, Blood samples taken from captives 1984-1989 102°28'E; collected 23 Oct. 1910 by museum by W. Scheffrahn, J. R. de Ruiter, and J. A. R. collector; zrc, 3 (including 2 skulls only; exter- A. M. van Hooff (1994, p. 135). FAS; B:IO-6 nal measurements recorded on skull tag and in

through IO- 10. Weitzel et al., 1988, p. 107). FAS; B:WM-27. Nicobar Islands, INDIA; 6°45'-8°02'N, 93°20'- Ober Langkat district; Sumatra, INDONESIA; ca. 93°57'E; collected ca. 1846 by H. Lewis (see 3°45'N, 98°20'E; collected 1897-1899 by G. Blyth, 1846, p. 366; J. Anderson, 1881, p. 65; Schneider (1905, p. 72); museum and number Khajuria, [1955], p. 109); formerly preserved in of specimens unknown (not seen). FAS; B:S-20. zsi (not seen). Examined for malaria parasites Ochodia, NafRiver; BANGLADESH; ca. 2 1°05'N, in 1979 by L. Kalra (1980, p. 50). Examined for 92°12'E; observed Sep. 1982-Feb. 1983 by M. malaria parasites ca. 1984 by S. Sinha and A. A. R. Khan and M. A. Wahab (1983, p. 104; Gajanana (1984, p. 567). UMB; A:N-1 through cf. M. A. R. Khan, 1981, p. 13; 1985, p. 30). N-4. AUR; A:Ba-l. Nikiniki, [720 m]; Pulau Timor, Lesser Sunda Is- Oeassa (?= Oebuah) vicinity; Pulau Semau, Lesser lands, INDONESIA; 9°49'S, 124°28'E; collected Sunda Islands, INDONESIA; ca. 10°07'S, 28 Mar. 1932 by G. Stein (see Hellmayr, 1914, 123°27'E; observed ca. May 1 859 by A. R. Wal-

p. 5); mzb, 1; zmb, 1 (skin only). FAS; CLS-27. lace (1869, p. 294). FAS; GLS-24. Nipah, Telok, vicinity; Pulau Tioman, MALAY- Oebuah. See Oeassa. SIA: WEST MALAYSIA; ca. 2°44'N, 104°08'E; Oewada Sami, Pulau. See Kode, Nusa. observed Mar.-Apr. 1962 by Lord Medway Ok Yam. See Laem Ngop-Phumi Cham Yearn. (1966, p. 16). FAS; B:SCS-3. Ol-kolo-kwak vicinity; Katchall Island, Nicobar Niur, Suaka Margasatwa Ujungkulon; Java, IN- Islands, INDIA; 7°52'-8°02'N, 93°18'-93°27'E; DONESIA; 6°40'S, 105°21'E; observed 20 Aug. collected 21 Feb. 1901 by W. L. Abbott (see 1941 by A. Hoogerwerf (1970, p. 408). FAS; Miller, 1902b, p. 751; Kloss, 1903a, p. 114); B:J-14. usnm, 2. UMB; A:N-1. Noesa Penida. See Penida, Nusa. [Oo vicinity], Pulau Sumbawa, Lesser Sunda Is- Nong Chik region; THAILAND; ca. 6°48'N, lands, INDONESIA; ca. 8°25'S, 118°35'E; col-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 157 lected 27 Nov. 1909 by J. Elbert (1912, p. 58); Pagai Selatan, Pulau, INDONESIA; 2°47'-3°21'S, nms, 1. FAS;C:LS-11. 100°10'-100°28'E; M. fascicularis reported ab- Outer Bolongs. See Myengun Kyun. sent before 1 928 by C. B. Kloss ([ 1928], p. 802). Pacific Tin; MALAYSIA: WEST MALAYSIA; ca. B:i. 3°25'N, 101°25'E; examined for malaria ca. Pagai Utara, Pulau, INDONESIA; 2°30'-2°52'S, 1 96 1-1 962 by R. H. Wharton, D. E. Eyles, McW. 99°58'-100°13'E; M. fascicularis reported ab-

Warren, and W. H. Cheong (1964, p. 58). FAS; sent before 1 928 by C. B. Kloss ([ 1 928], p. 802). B:WM-18. B:h. Padang; Sumatra, INDONESIA; 0°57'S, 100°21'E; Pagansan; Sumatra, INDONESIA; ca. 1°00'N, collected in 1836 by S. Miiller; rmnh, 7 (skins lOTOO'E; collected ca. 1907 by M. Moszkowski only, skulls inside). Collected in 1890 by H. (1909, maps following pp. 192, 328); zmb, 4 Meyer; zmuz, 2 (including 1 skin only). Col- (skeletons only). FAS; B:S-40. lected date unknown by unknown collector; Pagat. See Hantakan, 3 km N of Pagat. rmnh, 3; zmuz, 1 (skull only). Clinical exami- Pagurawan. See Paguruan, Sungai. nation Aug.-Nov. 1980 by K. Matsubayashi and Paguruan, Sungai; Sumatra, INDONESIA; ca. D. Sajuthi (1981, p. 48). FAS; B:S-47. 3°26'N, 99°20'E; collected 1897-1899 by G. Padang, Pulau, north coast, INDONESIA; ca. Schneider (1905, p. 72); museum and number 1°22'N, 102°20'E; collected 27 Mar. 1906 by W. of specimens unknown (not seen). FAS; B:S-23. L. Abbott (field catalog; cf. Lyon, 1908, p. 624); Pai, Teluk; Pulau Karimata, INDONESIA; 1°32'- usnm, 1 (skull only; external measurements re- 1°40'S, 108°48'-108°59'E; collected 24-27 Aug. corded on skull tag). FAS; B:SM-6. 1904 by W. L. Abbott (see Miller, 1906b, p. 55); Padang/Bedagei district; Sumatra, INDONESIA; usnm, 2. FAS; B:SCS-22. ca. 3°20'N, 99°10'E; collected 1897-1899 by G. Paillin. See Pang Roloem-Sur Sdei area. Schneider (1905, p. 72); museum and number Pajakombo. See Payakumbuh. of specimens unknown (not seen). FAS; B:S-23. Pajeti. See Payeti-Kambaniru. Padang highlands; Sumatra, INDONESIA; ca. Pajo, Danau Singkarak, Balipor district; Sumatra, 0°57'S, 100°21'E; collected in 1878 by J. F. INDONESIA; 0°37'S, 100°30'E; collected be- Snelleman(1887, p. 10); rmnh, 1. FAS; B:S-47. fore 1880 by C. Bock (see P. H. Napier, 1981, Padangsidempuan, ca. 45 km NNW; Sumatra, p. 16); bm(nh), 1. FAS; B:S-46. INDONESIA; ca. 1°44'N, 99°07'E; reported Pakan Selasa; Sumatra, INDONESIA; 1°34'S, present Nov. 1971-Jan. 1973 by local residents 101°08'E; observed Nov. 1971-Jan. 1973 by C. (Crockett & Wilson, 1980, p. 156). FAS; B:S- M. Crockett and W. L. Wilson (1980, p. 156). 34. FAS; B:S-52. Padangsidempuan vicinity; Sumatra, INDONE- Pakan Selasa, ca. 10 km SSE; Sumatra, INDO- SIA; ca. 1°22'N, 99°16'E; reported present Nov. NESIA; ca. 1°39'S, lOTlO'E; observed Nov. 1971-Jan. 1973 by local residents (Crockett & 1971-Jan. 1973 by C. M. Crockett and W. L. Wilson, 1980, p. 156). FAS; B:S-33. Wilson (1980, p. 156). FAS; B:S-52. Padar, Pulau, Lesser Sunda Islands, INDONE- Pakchan River, near Bankachon; BURMA; ca. SIA; 8°38'-8°42'S, 119°32'-119°37'E; monkeys 10°09'N, 98°36'E; collected 29 Jan. 1914 and reported absent Jul. 1 969-Jun. 1 970 by W. Auf- Dec. 1913-Apr. 1914 by G. C. Shortridge (in fenberg(1981,p. 242). C:e. Wroughton, 1915, p. 696); bm(nh), 2. AUR/ Padar-kechil, Pulau. See Padar-kecil, Pulau. FAS/MUL; A:Bu-22. Padar-kecil, Pulau, Lesser Sunda Islands, IN- Pakchan River, near Maliwun; BURMA; 10°14'N, DONESIA; 8°4 l'S, 1 1 9°32'E; monkeys reported 98°37'E; collected 28 Feb. 1928 by Faunthorpe-

absent 1969-1973 by W. Auffenberg (1981, p. Vernay Expedition; amnh, 1 . AUR/FAS/MUL; 40). C:e. A:Bu-22. Padas Bay; Borneo, MALAYSIA: SABAH; ca. Pak Chong, Sathani; THAILAND; 14°42'N, 5°13'N, 115°34'E; observed Jul.-Oct. 1968 by 101°25'E; collected 29 Feb. 1924 by H. M. Smith T. Mano (see Kawabe & Mano, 1972, p. 215). (see Kloss, 1916a, p. 2; Riley, 1938, p. l;Weitzel FAS;C:Sab-l. et al., 1988, p. 106); zrc, 1. Collected 21 May

Padeco, Hak Pengusahaan Hutan (HPH); Suma- 1 93 1 by C. J. Aagaard; zrc, 1 . AUR/FAS/MUL; tra, INDONESIA; not located; observed Dec. A:T-25. 1988 by M. Bismark (1992, p. 11). FAS; not Pak Jong. See Pak Chong, Sathani. mapped. Pak Klong Pran. See Pran Buri, Mae Nam, mouth.

158 FIELDIANA: ZOOLOGY Pak Nam Chumphon. See Chumphon, Khlong, Palembang, ca. 60 km NW; Sumatra, INDO- mouth. NESIA; ca. 2°37'S, 104°21'E; observed Nov. Paknampoh. See Ban Pak Nam Pho. 1971-Jan. 1973 by C. M. Crockett and W. L. Paku. See Pelandok, Sungai, Paku, Saribas. Wilson (1980, p. 156). FAS; B:S-61. Paku, Saribas; Borneo, MALAYSIA: SARAWAK; Palembang, ca. 60 km SW; Sumatra, INDONE- 1917 C. 1°27'N, 1H°27'E; collected Mar. by SIA; ca. 3°20'S, 104°21'E; observed Nov. 1971- Chunggat and H. C. Robinson (see P. H. Napier, Jan. 1973 by C. M. Crockett and W. L. Wilson 2. C:Sar-9. 1981, p. 14); bm(nh), FAS; (1980, p. 156). FAS; B:S-75. Paku Cave; Borneo, MALAYSIA: SARAWAK; Palembang, ca. 65 km N; Sumatra, INDONESIA; 1°25'N, 1 10°1 l'E; collected 1870 and 1879, prob- ca. 2°24'S, 104°41'E; observed Nov. 1971-Jan. ably by A. H. Everett; bm(nh), Subdepartment 1 973 by C. M. Crockett and W. L. Wilson (1980, of Anthropology, Department of Palaeontology, p. 156). FAS;B:S-61. 2 (1 skeleton only, 1 skull only; not seen, data Palembang, ca. 65 km NNW; Sumatra, INDO- from P. H. Napier, 1981, p. 20). FAS; C:Sar-3. NESIA; ca. 2°27'S, 104°32'E; observed Nov. Palembang, ca. 30 km N; Sumatra, INDONESIA; 1971-Jan. 1973 by C. M. Crockett and W. L. ca. 2°43'S, 104°46'E; observed Nov. 1971-Jan. Wilson (1980, p. 156). FAS; B:S-61. 1 973 by C. M. Crockett and W. L. Wilson (1980, Palembang, ca. 65 km SW; Sumatra, INDONE- p. 156). FAS;B:S-61. SIA; ca. 3°20'S, 104°16'E; observed Nov. 1971- Palembang, ca. 30 km NNE; Sumatra, INDO- Jan. 1973 by C. M. Crockett and W. L. Wilson NESIA; ca. 2°44'S, 104°51'E; observed Nov. (1980, p. 156). FAS;B:S-75. 1971-Jan. 1973 by C. M. Crockett and W. L. Palembang, ca. 70 km N; Sumatra, INDONESIA; Wilson (1980, p. 156). FAS; B:S-61. ca. observed Nov. 1971-Jan. Palembang, ca. 35 km SW; Sumatra, INDONE- 2°20'S, 104°42'E; 1 973 by C. M. Crockett and W. L. Wilson (1 980, SIA; ca. 3°13'S, 104°33'E; observed Nov. 1971- p. 156). FAS;B:S-61. Jan. 1973 by C. M. Crockett and W. L. Wilson ca. 70 km Sumatra, INDO- (1980, p. 156). FAS;B:S-61. Palembang, NNW; ca. observed Nov. Palembang, ca. 40 km N; Sumatra, INDONESIA; NESIA; 2°29'S, 104°23'E; 1971-Jan. 1973 C. M. Crockett and W. L. ca. 2°38'S, 104°48'E; observed Nov. 1971-Jan. by Wilson B:S-61. 1973 by C. M. Crockett and W. L. Wilson (1980, (1980, p. 156). FAS; p. 156). FAS;B:S-61. Palembang, ca. 70 km SW; Sumatra, INDONE- Palembang, ca. 40 km NNW; Sumatra, INDO- SIA; ca. 3°23'S, 104°16'E; observed Nov. 1971- NESIA; ca. 2°39'S, 104°34'E; observed Nov. Jan. 1973 by C. M. Crockett and W. L. Wilson 1971-Jan. 1973 by C. M. Crockett and W. L. (1980, p. 156). FAS;B:S-75. Wilson (1980, p. 156). FAS; B:S-61. Palembang, ca. 80 km N; Sumatra, INDONESIA; Palembang, ca. 45 km N; Sumatra, INDONESIA; ca. 2°15'S, 104°48'E; observed Nov. 1971-Jan. ca. 2°36'S, 104°43'E; observed Nov. 1971-Jan. 1 973 by C. M. Crockett and W. L. Wilson (1980, 1973 by C. M. Crockett and W. L. Wilson (1980, p. 156). FAS;B:S-61. p. 156). FAS;B:S-61. Palembang, ca. 90 km WSW; Sumatra, INDO- Palembang, ca. 45 km NW; Sumatra, INDO- NESIA; ca. 3°19'S, 104°01'E; observed Nov. NESIA; ca. 2°42'S, 104°28'E; observed Nov. 1971-Jan. 1973 by C. M. Crockett and W. L. 1971-Jan. 1973 by C. M. Crockett and W. L. Wilson (1980, p. 156). FAS; B:S-63. Wilson (1980, p. 156). FAS; B:S-61. Palembang, ca. 95 km SW; Sumatra, INDONE- Palembang, ca. 50 km N; Sumatra, INDONESIA; SIA; ca. 3°28'S, 104°05'E; observed Nov. 1971- ca. 2°32'S, 104°48'E; observed Nov. 1971-Jan. Jan. 1973 by C. M. Crockett and W. L. Wilson 1973 by C. M. Crockett and W. L. Wilson (1 980, (1980, p. 156). FAS;B:S-63. p. 156). FAS;B:S-61. Palembang, ca. 100 km N; Sumatra, INDONE- Palembang, ca. 60 km NNE; Sumatra, INDO- SIA; ca. 2°05'S, 104°47'E; observed Nov. 1971- NESIA; ca. 2°27'S, 104°51'E; observed Nov. Jan. 1973 by C. M. Crockett and W. L. Wilson 1971-Jan. 1973 by C. M. Crockett and W. L. (1980, p. 156). FAS;B:S-61. Wilson (1980, p. 156). FAS; B:S-61. Palembang, ca. 100 km SW; Sumatra, INDO- Palembang, ca. 60 km NNW; Sumatra, INDO- NESIA; ca. 3°26'S, 103°58'E; observed Nov. NESIA; ca. 2°32'S, 104°26'E; observed Nov. 1971-Jan. 1973 by C. M. Crockett and W. L. 1971-Jan. 1973 by C. M. Crockett and W. L. Wilson (1980, p. 156). FAS; B:S-63. Wilson (1980, p. 156). FAS; B:S-61. Palembang district; Sumatra, INDONESIA; ca.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 159 2°55'S, 104°45'E; collected in 1905 by B. Hagen; Pantai Krachut; Pulau Pinang [1], MALAYSIA: zsbs, 2 (skins only). FAS; B:S-61. WEST MALAYSIA; 5°27'N, 100°1 l'E; collect- Palung, Gunung. See Gunung Palung Nature Re- ed 24 Mar. 1911 by E. Seimund (see Weitzel et serve. al., 1988, p. 1 14); zrc, 1. FAS; B:SM-2. Pamuja, Tanjung; Pulau Bangka, INDONESIA; Pantar, Pulau, Lesser Sunda Islands, INDONE- ca. 1°40'S, 105°20'E; collected 19 Jun. 1904 by SIA; 8°11'-8°33'S, 123°55'-124°19'E; M.fasci- W. L. Abbott (in Lyon, 1906, p. 607); usnm, 1. cularis reported absent before 1937 by J. J. M. FAS;B:SCS-18. F. Symons (Mertens, 1936, p. 319). C:g. Panaitan, Pulau, INDONESIA; 6°32'-6°46'S, Papagaran. See Papagaran Besar, Pulau. 105°07'-105°16'E; observed 1932-1957 by A. Papagaran Besar, Pulau, Lesser Sunda Islands, Hoogerwerf (1970, p. 408). FAS; B:J-14. INDONESIA; 8°34'-8°35'S, 1 19°43'-1 19°44'E; Pandan, Sungai; Singapore Island; SINGAPORE; monkeys reported absent 1969-1973 by W. Auf- ca. 1°19'N, 103°45'E; observed before 1901 by fenberg(1981,p. 40). C:d. S. S. Flower (1900, p. 316). FAS; B:SCS-7. Papar, 20 mi (= 32 km) SW of Kota Kinabalu; Pandang, Sungei. See Pandan, Sungai. Borneo, MALAYSIA: SABAH; 5°44'N, Panebangan, Pulau. See Penebangan, Pulau. 1 1 5°56'E; collected 1 2 Sep. 1 960 by R. E. Kuntz Pangadaran. See Pangandaran. (1969, p. 193); usnm, 1. FAS; C:Sab-3. Pangandaran, Teluk Parigi; Java, INDONESIA; Pap-ka; Sumatra, INDONESIA; ca. TOO'N, 7°41'S, 108°39'E; collected 4-11 Apr. 1908 by 101WE; collected Apr. 1908 by M. Mosz- G. C. Shortridge (see Thomas & Wroughton, kowski (1909, maps following pp. 192, 328); 1909a, p. 373); bm(nh), 3 (including 1 skin only, zmb, 1 (skull only). FAS; B:S-40. 1 skull only [with mounted skin, not examined]). Parepare; Pulau Sulawesi, INDONESIA; 4°01'S, Clinical examination in 1980 by K. Matsubay- 1 19°38'E; captive (presumably introduced) ob- ashi and D. Sajuthi (1981, p. 48). FAS; B:J-2. served 6-15 Oct. 1888 by M. Weber (1890a, p. Pangerango, Mt. See Pangrango, Gunung. v; 1890b, p. 102). Subspecies uncertain; not Pangkalan; Padang highlands; Sumatra, INDO- mapped. NESIA; ca. 0°57'S, 100°21'E; collected ca. 1889 Parit, primary swamp forest, sea level; Borneo: by E. Dubois; rmnh, 3 (skulls only). FAS; B:S- Kalimantan, INDONESIA; 2°10'S, 113°00'E; 47. collected 15 Jun.-16 Jul. 1935 by J. J. Menden; Pangkalansusu, Sumatra, INDONESIA; 4°07'N, amnh, 20 (incuding 1 skull not examined, mea- 98°13'E; collected 15 Jun. 1972 by P. F. D. Van surements from G. H. Albrecht, pers. comm., Peenen; usnm, 1 (infant in fluid). Collected be- Oct. 1991); mzb, 3; zmb, 4 (including 1 skin fore 1979 by NAMRU 2 Djakarta Detachment; only). FAS; C:K-12. usnm, 1 (skull only). FAS; B:S-19. Pasar Besar, Hak Pengusahaan Hutan (HPH); Su- Pangkor, Pulau, MALAYSIA: WEST MALAY- matra, INDONESIA; not located; observed Jul. SIA; 4°12'-4°16'N, 100°32'-100°35'E; reported 1989 by M. Bismark (1992, p. 11). FAS; not present ca. 1963 by J. L. Harrison and J. R. mapped. Hendrickson (1963, p. 548). FAS; B:SM-3. Pasar Usang area; Sumatra, INDONESIA; ca. Pangonan, Gunung Muria; Java, INDONESIA; ca. 1°00'S, 100°25'E; blood samples taken Jan.

6°36'S, 1 10°53'E; collected 19 Dec. 1928 by H. 1 979-Dec. 1 98 1 by Y. Kawamoto ( 1 982, p. 67). J. V. Sody; rmnh, 1. FAS; CJ-28. FAS; B:S-47. Pangrango, Gunung; Java, INDONESIA; 6°46'S, Pasir Carolina, Purwakarta district; Java, IN- 106°57'E; collected in 1899 by Bartels; nhrm, DONESIA; ca. 6°34'S, 107°26'E; collected 17 27 (skulls only). Collected 4 Jul. 1902 and Feb. Aug. 1932 by Kloster; mzb, 1. FAS; B:J-20. 1 903 by unknown collectors; rmnh, 2 (skins only, Pasir Panjang; Pulau Bintan, Kepulauan Riau, IN- skulls inside). FAS; B:J-18. DONESIA; ca. 1°10'N, 104°30'E; collected 7 Pangrolim. See Pang Roloem-Sur Sdei area. Aug. 1902 by W. L. Abbott (field catalog; cf. Pang Roloem-Sur Sdei area; CAMBODIA; ca. Miller, 1906c, p. 282); usnm, 1. Collected 8 Jun. 12°50'N, 102°45'E; observed Oct. 1962 by D. 1 908 by H. C. Robinson (in Thomas & Wrough- E. Eyles, R. H. Wharton, W. H. Cheong, and ton, 1909b, p. 104); bm(nh), 1; zrc, 1 (skull McW. Warren (1964, p. 9). FAS; A:C-2. only). FAS; B:SCS-8. Panjaer, Tanjong. See Tanjong Panjair, Sungai Pasoh Forest Reserve, 50 m; MALAYSIA: WEST Rompin. MALAYSIA; ca. 2°57'N, 102°22'E; observed Jul. Panjang, Pulau. See Mentigi. 1978-Jun. 1981 by C. W. Marsh and W. L.

160 FIELDIANA: ZOOLOGY Wilson (1981, p. 232). Observed Jan. 1979-Jul. SIA: SARAWAK; 1°32'N, 1 10°44'E; collected 15

1980 by J. O. Caldecott (1986b, p. 21). FAS; Aug. 1916 by H. C. Robinson (see Weitzel et al., B:WM-20. 1988, p. 142); zrc, 1. FAS; C:Sar-6. Patani. See Pattani. Pelapis Tengah, Pulau, INDONESIA; 1°17'- o Patani, Cape. See Pho, Laem. 1°20'S, 109°08'-109 10'E; reported present 29 Patong, Bukit, ca. 1 km west; MALAYSIA: WEST May-1 June 1907 by W. L. Abbott (in Lyon,

ca. observed 1 1 1 1 1 1 L. MALAYSIA; 3°43'N, 102°02'E; 9 , p. 60). Collected 2 Mar. 93 by Coo- Jul.-Dec. 1977 by D. Chivers and G. Davies mans de Ruiter and Madzoed (see Chasen,

(1979, p. 22). FAS; B:WM-15. 1935b, p. 2); mzb, 1. FAS; B:SCS-23. collected Pattani; THAILAND; 6°52'N, 101°16'E; Pelawan, Sungai; Borneo: Kalimantan, INDO- 2 Jun. 1901 H. C. Robinson Annandale by (see NESIA; 1°11'N, 117°55'E; collected 18 Dec. & Robinson, 1903, p. xxi; Bonhote, 1903, p. 3); 1913 by H. C. Raven (field catalog; cf. Deignan, 1 A:T-68. smtd, (skull only). FAS; [1960], p. 269); usnm, 1. FAS; C:K-44. IN- Payakumbuh, Padang highlands; Sumatra, Pelehari. See Pelaihari. DONESIA; ca. 0°14'S, 100°38'E; collected ca. Peling, Pulo. See Piling, Pulau. 1889 by E. Dubois; rmnh, 1 (skull only). FAS; Pemanggil, Pulau, MALAYSIA: WEST MALAY- B:S-45. SIA; 2°34'-2°36'N, 104°18'-104°21'E; collected Payeti-Kambaniru; Pulau Sumba, Lesser Sunda 12 Jun. 1901 by W. L. Abbott (see Riley, 1938, Islands, INDONESIA; 9°39'S, 120°18'E; col- p. 14); usnm, 1. Collected 12 Jul. 1915 by H. lected 26-27 Mar. 1925 by P. F. Franck (see C. Robinson; bm(nh), 1. FAS; B:SCS-4. Dammerman, 1926a, p. 38); mzb, 2. FAS; C:LS- Island. See Pulau. 22. Pemangil Pemanggil, Sumatra, Pegou. See Pegu. Pematangsiantar, INDONESIA; 2°57'N, 99°03'E; collected Jul. 1937 B. Lawrence; Pegu; BURMA; 17°20'N, 96°29'E; obtained in by mcz, 2. FAS; B:S-30. 1827-1828 by A. A. M. Reynaud (see de Rossel Pulau Karimun, IN- et al., 1829, p. 603; de Rossel, 1829, p. 609; I. Pemeral; Kepulauan Riau, collected 1 1 Geoffroy, 1831, pp. 58, 77); museum unknown, DONESIA; 1°00'N, 103°25'E; Aug. 1 908 H. C. Robinson Thomas & 1 (not seen). Observed before 1889 by W. T. by (in Wrough- and E. Blanford (1888b, p. 22). AUR; A:Bu-8. ton, 1909b, pp. 105, 107) Seimund; 4. B:SCS-17. Pejantan, Pulau, INDONESIA; 0°07'-0°09'N, bm(nh), FAS; 107°12'-107°15'E; observed 1-2 Aug. 1899 by Penang Botanical Gardens; Pulau PinangflJ, MA- W. L. Abbott (in Miller, 1900, p. 243; cf. Kloss, LAYSIA: WEST MALAYSIA; ca. 5°25'N, C. 1903b, p. 59). Collected 15 Mar. 1907 by C. 100°20'E; observed 1940-1961 by K. Cheang Bruegel; zsbs, 1 (skin only). FAS; B:SCS-25. (1962, p. 73). Observed Jul.-Sep. 1960 by Y. Pekan vicinity; MALAYSIA: WEST MALAY- Furuya (196 1-1 962a, p. 76). Observed in 1974 SIA; ca. 3°30'N, 103°25'E; observed 25-28 Jun. by C. Spencer (1975, p. 83). FAS; B:SM-2. 1891 by H. N. Ridley, W. Davison, and H. J. Penang Hill; Pulau Pinang [1], MALAYSIA: Kelsall (see Kelsall, 1894a, p. 34). FAS; B:WM- WEST MALAYSIA; ca. 5°26'N, 100°16'E; re- 13. ported present before 1 976 by C. Spencer (1975, Pelabuhanratu, Teluk, 100 ft (= 30 m); Java, IN- p. 85). FAS; B:SM-2. DONESIA; 6°58'S, 106°32'E; collected 18 Mar. Penang Island. See Pinang, Pulau [1]. Waterfall 1920 by C. B. Kloss (see Weitzel et al., 1988, p. Penang Gardens. See Penang Botanical 142); zrc, l.FAS;B:J-9. Gardens. Pelabuhanratu, Teluk, Bantam region; Java, IN- Penebangan, Pulau, INDONESIA; l°10'-ri3'S, DONESIA; ca. 6°58'S, 106°32'E; collected 10 109°13'-109°16'E; observed 16 May-21 Sep. Nov. 1909 by O. Bryant (see field catalog of W. 1907 by W. L. Abbott (in Lyon, 1911, p. 60). Palmer); usnm, 1. FAS; B:J-9. FAS; B:SCS-23. Pelaihari; Borneo: Kalimantan, INDONESIA; Pengiki, Pulo. See Pejantan, Pulau. 3°48'S, 1 14°45'E; collected 20 Aug. 1866 by J. Pengsong, Gunung; Pulau Lombok, Lesser Sunda Semmelink; rmnh, 1 (skin only, skull inside). Islands, INDONESIA; ca. 8°36'S, 116°06'E; Collected before 1897 by J. H. F. Kohlbrugge blood samples taken Jan. 1979-Dec. 1981 by (1896a, p. 184); museum unknown, 1 (not ex- Y. Kawamoto (1982, p. 66). FAS; C:LS-5. amined). FAS; C:K-26. Penida, Nusa, 300 m, Lesser Sunda Islands, IN- Pelandok, Sungai, Paku, Saribas; Borneo, MALAY- DONESIA; 8°43'-8°48'S, 115°30'-115°36'E;

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 161 collected 26 Feb. 1938 by V. von Plessen; amnh, Phayam, Ko, THAILAND; 9°42'-9°46'N, 98°23'- 3. FAS; C:LS-4. 98°26'E; collected 20 Feb. 1919 by H. C. Rob-

Penrissen, Gunung, 4000 ft (= 1 200 m); Borneo, inson and C. B. Kloss; zrc, 1 . AUR/FAS/MUL; MALAYSIA: SARAWAK; 1°07'N, 110°13'E; A:T-45. collected May 1899 by E. A. W. Cox and R. W. Phet Buri; THAILAND; 13°06'N, 99°57'E; col- L. Shelford (see Shelford, 1 9 1 6, p. 2 1 7; cf. Med- lected in 1862 by M. Boucourt; mnhn, 1 (skin

1 7 1 altitude as way, 977, p. , who cited 4500 ft); only). Subspecies uncertain; A:T-37. smk, 2 (skins only, skulls inside). FAS; C:Sar-2. Phi Phi Don, Ko, THAILAND; 7°43'-7°47'N, Penuba. See Selayar, Pulau. 98°46'-98°48'E; collected 3-5 Feb. 1919 by H. Pepidon, Ko. See Phi Phi Don, Ko. C. Robinson (1921, p. 5) and C. B. Kloss; zrc, Perak, Sungai, upper. See Pong, upper Sungai Per- 4. FAS; A.T-54. ak. Pho, Laem; THAILAND; ca. 6°57'N, 101°16'E; Perangin, Bukit, 75-360 m; MALAYSIA: WEST reported present Jun. 1901 by N. Annandale

MALAYSIA; 6°22'N, 100°31'E; reported as and H. C. Robinson (1903, p. xl). FAS; A:T-68. probably present Jul. 1978-Jun. 1981 by C. W. Phu Phan, Maung district; THAILAND; 1 6°42'N, Marsh and W. L. Wilson (1981, p. 232). FAS; 104°25'E; collected date unknown by R. E. El- B:WM-3. bel; usnm, 1 (skin No. 307714, skull No. Perawang, 4 km NW; Sumatra, INDONESIA; ca. 307732). AUR/FAS/MUL; A:T-15. 0°40'N, 10 1°35'E; observed Sep.-Nov. 1981 and Phu-Qui. See Phu Quoc, Dao. Apr. 1983-Sep. 1984 by E.N. Megantara(1989, Phu Quoc, Dao, VIETNAM; 10°00'-10°27'N, p. 183). FAS;B:S-42. 103°50'-104°05'E; observed 1873-1874 by A. Perboewah, Kp. See Perbuah, Sungai Landak. Morice (1876, p. 31). Collected 30 Dec. 1927 Perbuah, Sungai Landak, 900 m; Borneo: Kali- by J. Delacour, W. P. Lowe, [and P. Jabouille]

mantan, INDONESIA; 0°53'N, 110°10'E; col- (in Delacour, 1928, p. 265); bm(nh), 1; mnhn, lected 27 Jul.-10 Aug. 1937 by J. J. Menden; l.FAS; A:V-6. amnh, 9 (including 1 skin only); mzb, 2. FAS; Phu-Quoi. See Phu Quoc, Dao. C:K-4. Piling, Pulau, INDONESIA; 2°44'-2°46'N, Perlis, Sungai, sea level; MALAYSIA: WEST MA- 106°10'-106°12'E; observed 17 Aug. 1899 by LAYSIA; 6°24'N, 1 00° 10'E; observed Jul. 1978- W. L. Abbott and C. B. Kloss (in Miller, 1900, Jun. 1981 by C. W. Marsh and W. L. Wilson p. 244; cf. Kloss, 1903b, p. 68). FAS; B:SCS-27. (1981, p. 232). FAS; B:WM-2. Pinagar; Sumatra, INDONESIA; OWN, 99°55'E; Petchaburi district. See "Siam." observed Nov. 1971-Jan. 1 973 by C. M. Crock- Peucang, Pulau, INDONESIA; 6°43'-6°45'S, ett and W. L. Wilson (1980, p. 156). FAS; B:S- 105°14'-105°16'E; observed 1932-1957 by A. 43. Hoogerwerf (1970, p. 408). Observed Sep. 1969- Pinagar, ca. 5 km SW; Sumatra, INDONESIA; ca. Jan. 1971 by W. Angst (1973, p. 626; 1975, p. 0°07'N, 99°52'E; observed Nov. 1971-Jan. 1973 326). FAS; B:J-14. by C. M. Crockett and W. L. Wilson (1980, p. Peutjang, Pulau. See Peucang, Pulau. 156). FAS;B:S-43. Pexabury. See Phet Buri. Pinagar, ca. 20 km E; Sumatra, INDONESIA; ca. Phai, Ko, THAILAND; 12°55'-12°57'N, 100°40'- OWN, 100°04'E; observed Nov. 1971-Jan. 100°41'E; primates reported as apparently ab- 1973 by C. M. Crockett and W. L. Wilson (1980, sent Jan. 1915 by collectors employed by C. B. p. 156). FAS;B:S-43. Kloss (1915a, p. 157; 1915b, p. 221). A:m. Pinang, Pulau [1], up to 2000 ft (= 600 m) MA- Phangan, Ko, southwest, THAILAND; ca. 9°40'N, LAYSIA: WEST MALAYSIA; 5°15'-5°29'N, 100°00'E; collected 27 May 1913 by H. C. Rob- 100°11'-100°20'E; reported present before 1847

inson (1915, p. 129) and E. Seimund; zrc, 2. by T. Cantor ( 1 846, p. 1 76). Observed before 1 90 1 FAS; A:T-50. by S. S. Flower (1900, p. 316). FAS; B:SM-2. Phangnga Bay National Park; THAILAND; ca. Pinang, Pulau [2], MALAYSIA: WEST MALAY- o 8°25'N, 98°37'E; observed 17-21 Nov. 1987 by SIA; 5°44'-5°45'N, 103°00'-103 01'E; collected R. Boonratana (1988, p. 76); also seen on un- 3 Sep. 1910 by C. B. Kloss (1911a, p. 177); specified adjacent islands. Subspecies uncertain; bm(nh), 2. FAS;B:SCS-1. A:T-51. Pinda, Nusa, Komodo area, Lesser Sunda Islands,

Phattalung. See Wat Suwankuha. INDONESIA; not located, ca. 8°30'S, 1 19°30'E;

162 FIELDIANA: ZOOLOGY monkeys reported absent 1 969-1 973 by W. Auf- Pota; Pulau Flores, Lesser Sunda Islands, IN- fenberg (1981, p. 40). Not mapped. DONESIA; 8°20'S, 120°46'E; observed Apr.- Ping, Mae Nam; THAILAND; ca. 16°00'N, May 1930 by Jhr. W. C. van Heurn (1932, p. 99°50'E; collected 7 Mar. 1924 by A. S. Vernay 65). FAS; GLS-18. (see Lowe, 1932, p. 202; 1933, p. 260); amnh, Poulo Condor. See Con Son. 1 (skull only; external measurements recorded Prachuap Khiri Khan, few miles north; THAI- A:T-4. in amnh catalog). Subspecies uncertain; LAND; ca. 1 1°50'N, 99°48'E; observed Nov. Pintu Gedong, Pulau, MALAYSIA: WEST MA- 1914-Feb. 1915 by N. Gyldenstolpe (1916, p. col- LAYSIA; 2°53'-2°56'N, 101°14'-101°16'E; 10; 1917b, p. 6). AUR/FAS/MUL; A:T-41. lected 2-3 Nov. 1912 by museum collector; Prahmon. See Ban Phra Muang. 2. B:SM-4. bm(nh), 1; zrc, FAS; Pran Buri, Mae Nam, mouth; THAILAND; P. See Pulau. Kemoedjan. Kemujan, 12°24'N, 100°00'E; collected 28 Jun. 1917 by Plaine Mauritius Island, MAURITIUS; Sophie; W. J. F. Williamson and M. A. Smith (see Kloss, 19°59'-20°31'S, 57°18'-57°48'E; introduced 1917, p. 289); zrc, 2. AUR/FAS/MUL; A:T- population, collected 25 Apr. 1949 by R. d'Un- 38. ienville; bm(nh), 1. Subspecies uncertain; not Prang Koo; THAILAND; 15°40'N, 104°23'E; ob- mapped. served 17-18 Jul. 1989 by N. Aggimarangsee P. Lankawi. See Langkawi, Pulau. (1992, pp. 109, 120; pers. comm., Oct. 1993). Plateau des Bolovens. See Thateng, Muang, Pla- Subspecies uncertain; A:T-16. teau des Bolovens. Preanger. See Ciwangi; Tasikmalaya, Preanger re- Pleihari Tanah Laut Game Sanctuary; Borneo: Ka- gion; Tjeringin, near Banjar, Preanger region. limantan, INDONESIA; ca. 3°48'S, 114°45'E; Preanger district; Java, INDONESIA; 6°36'- reported present before 1983 by G. Davies 7°48'S, 106°22'-108°46'E; collected 12 Nov. ([1983], p. 148). FAS; GK-26. 1890-8 Feb. 1891 by A. A. W. Hubrecht (see Pleyharie. See Pelaihari. Zuckerman, [1933], p. 1062); museum un- Podowk. See Ataran River. known (not seen). FAS; B:J-6. Poelau (?= Kuda), Sungai Sibau; Borneo: Kali- Preparis Island, BURMA; 14°51'-14°53'N, 93°40'- mantan, INDONESIA; ca. 1°02'N, 112°59'E; 93°42'E; reported present before 1929 by C. B. collected 19 Jun. 1894 by J. Buttikofer (1897, Kloss ([1928], p. 806). AUR; A:Bu-5. p. 20); rmnh, 1 (skin only). FAS; GK-18. "P. Sember"; Java, INDONESIA; ca. 6°50'S, Pong, upper Sungai Perak, 900 ft (= 275 m); MA- 108°15'E; clinical examination in 1980 by K. LAYSIA: WEST MALAYSIA; 5°36'N, 1 Matsubayashi and D. Sajuthi (1 98 , p. 48). FAS; 101°02'E; collected 16 Jan. 1932 by A. B. Hol- B:J-22. loway and A. S. Vernay (see P. H. Napier, 1981, Pulaki, Pulau Bali, Lesser Sunda Islands, p. 14); bm(nh), 1. FAS; B:WM-4. Tanjung; observed Ponggol. See Punggol. INDONESIA; 8°07'S, 114°35'E; Sep. 1969-Jan. 1971 W. Pongka, Kampung; Pulau Karimun, Kepulauan by Angst (1975, p. 326). Blood taken in 1980 Y. Riau, INDONESIA; 1°06'N, 103°24'E; collect- samples by Kawamoto, K. Nozawa, and Tb. M. Ischak (1981, p. 16). ed 25 May 1903 by W. L. Abbott (field catalog; C:LS-1. cf. Miller, 1906c, p. 277); usnm, 1 (skull only). FAS; FAS;B:SCS-17. Pulau Doem area; Java, INDONESIA; ca. 7°35'S, taken 1979-1981 Pontianak. See Ambawang, Sungai, near Pontia- 112°30'E; blood samples by nak. Y. Kawamoto (1982, p. 67). FAS; GJ-37. Port Dickson; MALAYSIA: WEST MALAYSIA; Pulau Kaban. See Acheh, Pulau. Pulau Lontar. See Lanta Ko. 2°31'N, 101°48'E; collected 6 Jan.-2 May 1937 Yai, Misa. See Pulau. by E. Seimund; zrc, 3. Collected before 1940 Pulau Misa, Pulau Muntia. See Ko. by F. Colyer; bm(nh) (skin)/RCS(OM) (skull), 1. Muk, FAS; B:WM-24. Pulau Panjang. See Yao Noi, Ko. Pulau Anak. See Yao Ko. Port Swettenham vicinity; MALAYSIA: WEST Panjang Noi, MALAYSIA; ca. 3°00'N, 101°24'E; reported Pulau Sulawesi, southern, INDONESIA; ca. 5°00'S, present Aug.-Dec. 1970 by unspecified inform- 120°00'E; introduced, reported present before ants (Southwick & Cadigan, 1972, p. 10). FAS; 1883 by H. von Rosenberg (1882, p. 1 14). Re- B:WM-18. ported present before 1930 by K. W. Dammer-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 163 man (1929, p. 6). Subspecies uncertain; not 1 1 Apr. 1927 by Sunda-Expedition Rensch (see mapped. B. Rensch, 1930, p. 46; Mertens, 1930, p. 129; Pulau Telebong. See Talibong, Ko. I. Rensch, 1934, p. 227; Elbert, 1911, map 2); Pulau Telibun. See Talibong, Ko. mzb, 1. FAS; GLS-7. Pulo Beng Kalis. See Kapos Tinggi. Putussibau; Borneo: Kalimantan, INDONESIA; Pulo-Condore. See Con Son. 0°50'N, 112°56'E; collected 19 Nov.-17 Dec.

Pulo Gilla. See Jela, Pulau. 1 897 by A. Harrison, Jr., and H. M. Hiller; ansp, Pulo Mata. See Matak, Pulau. 3. FAS; CK-17. Pulo Mobur. See Mubur, Pulau. Raba; Pulau Sumbawa, Lesser Sunda Islands, IN- Pulo Nyur; Great Nicobar Island, west coast, IN- DONESIA; 8°27'S, 118°46'E; blood samples DIA; ca. 6°55'N, 93°40'E; observed 2 1 Mar. 1 90 1 taken Jul. 1981-Jan. 1 982 by Y.Kawamoto, Tb. by C. B. Kloss (1903a, p. 150). UMB; A:N-3. M. Ischak, and J. Supriatna (1982a, p. 58; cf. Pulo Peling. See Piling, Pulau. Kawamoto, 1982, p. 66; USBGN Gazetteer of Pulo Pengiki. See Pejantan, Pulau. Indonesia, 1982, p. 1036). FAS; C:LS-12. Pulo Piling. See Piling, Pulau. Rachaburi. See Tham Chomphon, Wat Huai Tak- Pulo Riabu. See Airabu, Pulau. haeng, and Wat Ratch Singkhorn. Pulo Selindang. See Selintang, Pulau. Rakata, Pulau, INDONESIA; 6°08'-6°10'S, Pulo Terutau. See Tarutao, Ko. 105°25'-105°28'E; primates reported absent Pulo Tioman. See Tioman, Pulau. 1883-1934 by K. W. Dammerman (1948, p. Pulu Tikus. See Tikus, Pulu. 315); volcanic explosion in 1883. B:k. Punang, Sungai; Borneo, MALAYSIA: SARA- Rakata-kecil, Pulau, INDONESIA; 6°05'-6°06'S, WAK; 4°54'N, 1 15°21'E; collected 16 Apr. and 105°26'-105°27'E; primates reported absent 14-15 May 1939 by G. Nunong; smk, 2 (skins 1883-1934 by K. W. Dammerman (1948, p. only); zrc, 2. FAS; C:Sar-22. 55); volcanic explosion in 1883. B:k. Punggol; Singapore Island, SINGAPORE; 1°25'N, Rana Mese, 1250-1400 m; Pulau Flores, Lesser 103°54'E; collected 8-11 Nov. 1910 by A. H. Sunda Islands, INDONESIA; 8°38'S, 120°34'E;

Wong; bm(nh), 1; zrc, 4-086, 1. Collected 15- collected 19 Jun. 1927 by Sunda-Expedition 1 6 Dec. 1 9 1 by unknown collector; zrc, 2 (skins Rensch (see B. Rensch, 1930, p. 148; Mertens, only). FAS; B:SCS-7. 1930, p. 139; I. Rensch, 1934, p. 227); nms, 1. Punoebo. See Selayar, Pulau. FAS; GLS-15. Puram; Borneo, MALAYSIA: SARAWAK; not Ranau, ca. 1500 ft (= 450 m); Borneo, MALAY- located, 0°43'^°58'N, 109°34'-115°38'E; col- SIA: SABAH; 5°57'N, 116°41'E; collected 23 lected Jul. 1896 by W. H. Furness III (1896, p. July 1937 by J. A. Griswold, Jr. (see Coolidge, 310) and H. M. Hiller (1896, p. 321); ansp, 1 1940, p. 123);mcz, 1. Collected 19-23 Sep. 1960 (skull only). FAS; not mapped. by R. E. Kuntz (1969, p. 193); usnm, 5. FAS; Pura Uluwatu, Bukit Peninsula; Pulau Bali, south- C:Sab-7. western tip, Lesser Sunda Islands, INDONE- "Rangoon"; BURMA; 16°47'N, 96°10'E; captive SIA; not located, 8°04'-8°48'S, 114°26'- acquired 16 Dec. 1870 by unknown collector; 115°42'E; observed Jul.-Sep. 1993 by M. F. rmnh, 1 (skin only). Captive acquired 8 May Small (1994, p. 8). FAS; not mapped. 1 875 by unknown collector; rmnh, 1 (skin only). Purukcahu, Sungai Barito; Borneo: Kalimantan, Subspecies uncertain; A:Bu-7. INDONESIA; 0°35'S, 114°35'E; collected be- Rangoon, near. See Desertion Creek, Elephant fore 191 1 by G. C. Shortridge; bm(nh), 1 (skull Point. only). Observed 1979-1986 by J. Marshall, D. Rang Yai, Ko, THAILAND; 7°57'N, 98°26'E; col- J. Chivers, and K. M. Burton (see Chivers & lected 8 Feb. 1 9 1 8 by local collectors employed Burton, [1991], p. 141). FAS; QK-33. by H. C. Robinson and C. B. Kloss (1919, p. Pussuk forest. See Pusuk forest. 87); zrc, 1. AUR/FAS/MUL; A:T-53. Pusuk, Gunung; Pulau Lombok, Lesser Sunda Is- Rantau; Borneo: Kalimantan, INDONESIA; lands, INDONESIA; ca. 8°27'S, 1 16°07'E; blood 2°56'S, 1 15°09'E; collected 10 Mar. 1916 by F. samples taken Jan. 1979-Dec. 1981 by Y. Ka- C. E. van der Putten (see Hooijer, 1962b, p. 44); wamoto (1982, p. 66). FAS; CLS-5. rmnh, 1 (skin only). FAS; CK-28. Pusuk forest; Pulau Lombok, Lesser Sunda Is- Rantau Panjang. See Kampong Rantau Panjang. lands, INDONESIA; 8°25'S, 1 1 6°3 1 'E; collected Rantauprapat vicinity; Sumatra, INDONESIA;

164 FIELDIANA: ZOOLOGY ca. 2°06'N, 99°50'E; reported present Nov. 1 97 1- wamoto, Tb. M. Ischak, and J. Supriatna (1982a, Jan. 1 973 by local residents (Crockett & Wilson, p. 58). FAS;C:LS-12. 1980, p. 156). FAS;B:S-35. Roema Manoeal, south foot of Gunung Kenepai; Ranun River. See Renun, Lae. Borneo: Kalimantan, INDONESIA; 0°40'N, Ratburi district. See "Siam." 111°42'E; collected 27 Dec. 1893 by J. Biitti- Rawa Danau. See Danau, Rawa. kofer (1897, p. 12); rmnh, 2. FAS; C:K-5. Rawas; Sumatra, INDONESIA; not located, 5°38'- Roi-ed. See Wat Koo Pra Kona. 5°57'S, 95°12'-106°05'E; collected in 1906 by Roi-et. See Wat Koo Pra Kona. W. Volz; zmb, 1 (skull only). FAS; not mapped. Rokan-kanan, Sungai; Sumatra, INDONESIA; Rayoh; Borneo, MALAYSIA: SABAH; 5°15'N, 1°23'N, 100°56'E; collected ca. 1907 by M. 1 15°50'E; collected 24 Jun. 1928 by G. Nunong Moszkowski (1909, map following p. 328); zmb, (see Davis, 1962, pp. 10, 127; Weitzel et al., 3. FAS; B:S-38. 1988, p. 143); zrc, 1. FAS; C:Sab-2. Rokankunan. See Rokan-kanan, Sungai. Rayong. See Wang Kaew. Rompin, Sungai. See Tanjong Panjair, Sungai Redang, Pulau, MALAYSIA: WEST MALAY- Rompin. SIA; 5°45'-5°49'N, 102°59'-103°03'E; collected Rondong, Nusa, Komodo area, Lesser Sunda Is- 2-5 Sep. 1910 by C. B. Kloss (1911a, p. 177); lands, INDONESIA; not located, ca. 8°30'S, zrc, 2. FAS; B:SCS-1. 1 19°30'E; monkeys reported absent 1969-1973 Renau. See Ranau. by W. Auffenberg (1981, p. 40). Not mapped. Rengsam, Tanjung; Pulau Bangka, INDONESIA; Ru, Pulo, BURMA; 9°56'-9°57'N, 98°32'-98° 34'E; 2°07'S, 105°35'E; collected 21 May 1904 by W. collected 13 Nov. 1913 by G. C. Shortridge (in

L. Abbott (in Lyon, 1 907, p. 607); usnm, 1 . FAS; Wroughton, 1915, p. 696); bnhs, 1. AUR/FAS/ B:SCS-18. MUL; A:Bu-23. Renun, Lae; Sumatra, INDONESIA; ca. 3°05'N, Rugading, near Bongkabong, near sea level; Bor- 97°55'E; observed May-Dec. 1971 by J. R. neo, MALAYSIA: SABAH; ca. 5°59'N, MacKinnon (1973, p. 240). FAS; B:S-15. 1 16°04'E; collected 22 Aug. 1937 by J. A. Gris- Reo; Pulau Flores, Lesser Sunda Islands, INDO- wold, Jr. (see Coolidge, 1940, p. 123); mcz, 1. NESIA; 8°19'S, 120°30'E; observed 21 Nov. FAS; C:Sab-4. 1888-9 Jan. 1889 by M. Weber (1890a, p. viii; Rukumodia, Naf River; BANGLADESH; ca. 1890b, p. 102). Observed Apr.-May 1930 by 21°05'N,92°12'E; observed Sep. 1982-Feb. 1983 Jhr. W. C. van Heurn (1932, p. 65). FAS; C:LS- by M. A. R. Khan and M. A. Wahab (1983, p. 17. 104; cf. M. A. R. Khan, 1981, p. 13; 1985, p. Riabu, Pulo. See Airabu, Pulau. 30). AUR; A:Ba-l. Riam, Kotawaringin district, 300 m; Borneo: Ka- Rungan, Sungai, right bank, 50 km above mouth; limantan, INDONESIA; 1°50'S, 111°54'E; col- Borneo: Kalimantan, INDONESIA; ca. 1°47'S, lected 29 Oct-3 Nov. 1 935 by J. J. Menden; amnh, 1 13°42'E; observed Jul. 1984-May 1986 by K. 4 (including 1 skin only). Collected 31 [Oct.] 1935 M. Burton (see Chivers & Burton, [1991], p. by J. J. Menden; mzb, 1. FAS; C:K-8. 140). FAS; CK-22. Rinca, Pulau, Lesser Sunda Islands, INDONE- Rungan, Sungai, right bank, above mouth; Borneo: SIA; 8°37'-8°48'S, 119°37'-119°47'E; observed Kalimantan, INDONESIA; ca. 2°09'S, 1 13°53'E; 26 May-6 Jul. 1953 by A. Hoogerwerf (1954, 2 troops observed Jul. 1984-May 1986 by K. p. 225; 1955, p. 26). Observed 18 Apr.-14 Jul. M. Burton (see Chivers & Burton, [1991], p. 1956 by P. Pfeffer (1959, p. 231). FAS; C:LS- 140). FAS; CK-23. 13. Rupat, Pulau, INDONESIA; 1°41'-2°08'N, Rinjani, Gunung, 1 700 m; Pulau Lombok, Lesser 101°23'-101°47'E; reported present 24 Feb.-3 Sunda Islands; INDONESIA; ca. 8°24'S, Apr. 1906 by W. L. Abbott (in Lyon, 1908, p. 1 16°28'E; observed 23 Sep.-31 Oct. 1987 by D. 624). FAS; B:SM-5. Kitchener, Boeadi, L. Charlton, and Mahara- Rupat Tinggi. See Kapos Tinggi. datunkamsi (1990, p. 98). FAS; CLS-7. Rutland Island, Andaman Islands, INDIA; 11*21'- Rintja, Pulau. See Rinca, Pulau. 1 1°31'N, 92°35'-92°42'E; primates reported ab- Rite; Pulau Sumbawa, Lesser Sunda Islands, IN- sent before 1903 by G. S. Miller, Jr. (1902b, p.

DONESIA; ca. 8°33'S, 1 1 8°53'E (cf. Raba); blood 792; cf. Kloss, [1928], p. 802; Chaturvedi, 1980, samples taken Jul. 1981-Jan. 1982 by Y. Ka- p. 134). A:g.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 165 Sababi, Pulau, Lesser Sunda Islands, INDONE- Samut Song Kram. See Samut Songkhram. o SIA; 8 21'-8 22'S, 120°01'-120°02'E; monkeys Sandakan, 8 mi (= 13 km) W; Borneo, MALAY- reported absent 1969-1973 by W. Auffenberg SIA: SABAH; ca. 5°50'N, 118°00'E; collected (1981, p. 40). C:d. 29 Aug. 1 929 by F. C. Wonder (see Davis, 1 962, Sabah; Borneo, MALAYSIA: SABAH; 4°06'- p. 10); fmnh, 1 (skull only). FAS; C:Sab-17. 7°02'N, 1 15°26'-1 19°17'E; collected 24 Aug.-4 Sandaran Agong, Kerinci region, 2450 ft (= 750 Oct. 1960 by R. E. Kuntz (1969, p. 193); usnm, m); Sumatra, INDONESIA; 2°07'S, 101°32'E; 4. FAS; not mapped. collected 6 Jun. 1914 by H. C. Robinson and Sabajor. See Sabajor Besar, Pulau. C. B. Kloss (1918, p. 6; in Pendlebury, 1936, p. Sabajor Besar, Pulau, Lesser Sunda Islands, IN- 27);zrc, l.FAS;B:S-50. DONESIA; 8°29'-8 31'S, 119°43'-119 44'E; Sangeang, Pulau, Lesser Sunda Islands, monkeys reported absent 1 969-1 973 by W. Auf- INDONESflA; 8°08'-8°16'S, 1 19°00'-1 19°07'E; fenberg (1981, p. 40). C:d. monkeys reported absent 1 969-1 973 by W. Auf- Sabalan. See Sebolon Besar, Pulau. fenberg (1981, p. 40). C:c. Sabor; Pulau Banggi, MALAYSIA: SABAH; Sangeh; Pulau Bali, Lesser Sunda Islands, IN- 7°14'N, 117°14'E; collected 19 Jun. 1991 by DONESIA; 8°28'S, 115°14'E; observed Sep. Shukor Md. Nor (pers. comm., 17 Jul. 1991); 1969-Jan. 1971 by W. Angst (1975, p. 326). fmnh, 1. FAS; C:Sab-10. Observed 21 Aug. 1973 by J. Fooden. Observed Saigon. See Ho Chi Minh City. in 1980 by A. Bakar, M. Amir, and Marshal Saint Matthew's Island. See Zadetkyi Kyun. (1981, p. 11). Observed Jul.-Sep. 1993 by M. Sakerat. See Ban Sakaerat. F. Small (1994, p. 8). FAS; CLS-3. Salak, Gunung, 2000 m; Java, INDONESIA; Sanggul, Bukit, Propinsi Bengkulu, 500 m; Su- 6°42'S, 106°44'E; collected Nov. 1931 and date matra, INDONESIA; 3°50'S, 102°37'N; col- unknown by H. J. V. Sody; rmnh, 3 (including lected 15-19 Aug. 1936 by J. J. Menden; amnh, 2 skulls only). FAS; B:J- 17. 5;mzb, 3. FAS;B:S-71. Salikan, Mt. See Selikan, Bukit. Sanglang- besar, Pulau, Kepulauan Riau, INDO- Sama, Sungei. See Ambawang, Sungai, near Pon- NESIA; 0°36'-0°38'N, 103°41'-103°42'E; mon- tianak. keys reported absent 10-1 1 Jul. 1903 by W. L. Samarinda. See Karangmumus, Sungai, near Sa- Abbott (see Miller, 1906c, p. 280). Not mapped. marinda, and Mahakam, Sungai, north bank, Sanglar. See Sanglang-besar, Pulau. above Samarinda. Sano, Wai, 650 m; Pulau Flores, Lesser Sunda Samasama; Pulau Nias, INDONESIA; 0°58'N, Islands, INDONESIA; 8°42'S, 120°00'E; col- 97°54'E; collected 2 1 Feb. 1 905 by W. L. Abbott lected 18 Nov. 1929 by Denin; mzb, 1. FAS; (field catalog; cf. Lyon, 1916, p. 458); usnm, 1. QLS-14. FAS; B:IO-9. Sapagaya Forest Reserve, near sea level; Borneo, Samui, Ko, center of island, 300 ft (= 90 m), MALAYSIA: SABAH; 5°37'N, 118°04'E; col- THAILAND; ca. 9°30'N, 100°00'E; collected 15 lected 9 Aug. 1950 by D. D. Davis (1962, p. May 1913 by H. C. Robinson (1915, p. 129; cf. 1 1); fmnh, 2. FAS; C:Sab-18. P. H. Napier, 1981, p. 17) and E. Seimund; Sapeh. See Ntori, Sape district. bm(nh), l.FAS; A:T-49. Sap Khao, 280 m; THAILAND; 15°35'N, 99°18'E; Samuk, Khao; THAILAND; 13°19'N, 100°54'E; observed 29 Nov. 1973 and 25 Jan. 1975 by A. reported present in 1981 by W. Y. Brockelman A. Eudey (1979, pp. 90, 198). Subspecies un- (1981, p. 13). Observed 10-16 Nov. 1990 by N. certain; A:T-19. Aggimarangsee(1992, pp. 1 10, 129; pers. comm., Sarawak; Borneo, MALAYSIA: SARAWAK; Oct. 1993). FAS; A:T-31. 0°52'-4°58'N, 109°33'-115°38'E; collected in Samunsam Wildlife Sanctuary; Borneo, MALAY- 1879, probably by A. H. Everett; bm(nh), Sub- SIA: SARAWAK; ca. 2°00'N, 109°38'E; re- department of Anthropology, Department of ported present in 1980 by S. Joines (1981, p. 9). Palaeontology, 3 (1 maxillary fragment, 2 iso- Observed before 1987 by G. Hohmann and W. lated molars; not seen, data from P. H. Napier, P. Peter (1986, pp. 89, 92). FAS; C:Sar-l. 1981, p. 20). FAS; not mapped. Samut Songkhram vicinity; THAILAND; ca. Sarawak, Sungai, mouth; Borneo, MALAYSIA: 13°24'N, 100°00'E; observed Mar. 1909 by K. SARAWAK; ca. 1°38'N, 110°27'E; observed

G. Gairdner ( 1 9 1 4, p. 28). Subspecies uncertain; 1897-1912 by R. W. C. Shelford (1916, p. 10). A:T-37. FAS; C:Sar-4.

166 FIELDIANA: ZOOLOGY Saribas. See Pelandok, Sungai, Paku, Saribas. LAYSIA; 1°54'N, 103°39'E; observed Jul. 1978- Sarn Pra Karn, Lopburi; THAILAND; 14°48'N, Jun. 1981 by C. W. Marsh and W. L. Wilson 100°37'E; observed 24 Feb. and 18 Apr. 1967 (1981, p. 232). FAS; B:WM-29. by J. Fooden (1971, p. 16). Observed before Sekonyer Kanan River (?= Sikunir, Sungai), Tan- 1976 by F. C. Cadigan and Lim Boo Liat (1975, jung Puting National Park; Borneo: Kaliman- p. 86). Observed 22 Jun.-9 Jul. 1989 by N. Ag- tan, INDONESIA; ca. 2°50'S, 111°47'E; ob- gimarangsee (1992, pp. 109, 124; pers. comm., served 1977-1983 by B. M. F. Galdikas and C. Oct. 1993). Subspecies uncertain; A:T-7. P. Yeager (1984, p. 50). FAS; CK-9. Sarongen; Java, INDONESIA; ca. 6°27'S, Selai, Sungai; MALAYSIA: WEST MALAYSIA; 106°00'E; collected 13 Nov. 1909 by O. Bryant 2°16'N, 103°25'E; observed Jul. 1978-Jun. 1981 (see field catalog in usnm); mcz, 1. FAS; B:J-12. by C. W. Marsh and W. L. Wilson (1981, p. Sarongge. See Sarongen. 232). FAS; B:WM-28. Sebang. See Sebangka, Pulau. Selama. See Ulu Ijok. Sebangka, Pulau, INDONESIA; 0°01'-0°15'N, Selatbliat; Pulau Kundur, Kepulauan Riau, IN- 104°29'-104°43'E; reported present 26-31 Jul. DONESIA; ca. 0°53'N, 103°22'E; collected 18- 1903 by W. L. Abbott (see Miller, 1906c, p. 20 Aug. 1908 by H. C. Robinson (in Thomas 284). FAS; B:SCS-12. & Wroughton, 1909b, pp. 105, 107) and E. Sei- Sebatik, Pulau; MALAYSIA: SABAH; 4° 10'- mund; bm(nh), 2. FAS; B:SCS-17. 4°17'N, 117°38'-117°53'E; collected 4 Jan. 1910 Selayar, Pulau, INDONESIA; 0°17'-0°20'S, by R. C. Andrews (1911, p. 22; 1943, p. 69); 104°23'-104°29'E; reported present 2-6 Aug. amnh, 1 (skin only); usnm, 2. FAS; C:Sab-12. 1903 by W. L. Abbott (see Miller, 1906c, p. Sebattick Island. See Sebatik, Pulau. 280). FAS; B:SCS-14. Sebattik Island. See Sebatik, Pulau. Selikan, Bukit; Borneo, MALAYSIA: SARA- Sebesi, Pulau, INDONESIA; 5°55'-5°58'S, WAK; 3°31'N, 1 14°04'E; collected July 1895 by 105°28'-105°32'E; primates reported absent Sep. E. Hose and C. Hose; usnm, 1. FAS; C:Sar-14. 1920-Jan. 1922) by K. W. Dammerman (1948, Selindang, Pulo. See Selintang, Pulau. p. 68). B:l. Selintang, Pulau, INDONESIA; 0°57'-0°58'N, Sebesy Island. See Sebesi, Pulau. 107°28'-107°29'E; observed 3-7 Aug. 1899 by Sebolon Besar, Pulau, Lesser Sunda Isllands, IN- W. L. Abbott (in Miller, 1900, p. 243; cf. Kloss, DONESIA; 8°24'S, 1 19°49'E; monkeys reported 1903b, p. 60). FAS; B:SCS-26. absent 1969-1973 by W. Auffenberg (1981, p. Semao Island. See Oeassa. 40). C:d. Semarang; Java, INDONESIA; 6°58'S, 1 10°25'E; Sebuku, Pulau, INDONESIA; 3°22'-3°38'S, collected 1907-1908 by C. Bruegel and/or U. 116 19'-116°27'E; reported present 31 Dec. Schaule; zsbs, 1 (skin only). FAS; C:J-26. 1907-5 Jan. 1908 by W. L. Abbott (in Lyon, Sembakung, Sungai; Borneo: Kalimantan, IN- 1911, p. 62). FAS;C:K-52. DONESIA; ca. 3°50'N, 1 17°00'E; collected Jul. Sechawa. See Bakung, Pulau. 1912 by Mohari (see Sody, 1949, p. 180); mzb, Sedagong, 900 m; Pulau Tioman, MALAYSIA: 2. FAS; QK-37. WEST MALAYSIA; ca. 2°48'N, 104°1 l'E; col- Sembaloen. See Pusuk forest. lected 14-20 May 1927 by N. Smedley: zrc, 2. Sembawang, Sungai; Singapore Island, SINGA- FAS; B:SCS-3. PORE; 1°27'N, 103°50'E; collected 21 May 1922 Segaliud River. See Lungmanis Station, Sungai Se- by F. N. Chasen; zrc, 1 (skin only). FAS; galiud. B:SCS-7. Segama, Sungai; Borneo, MALAYSIA: SABAH; Sember, P. See "P. Sember." 4°43'-5°25'N, 1 17°38'-1 18°47'E; collected 22 Sembilan Kepulauan, MALAYSIA: WEST MA- Aug. 1907 by Dr. Pagel; zmb, 1 (skull only); FAS; LAYSIA; ca. 4°02'N, 100°33'E; primates re- C:Sab-21. ported absent in 1953 by J. L. Harrison and J. Segobang, Sungai; Borneo, MALAYSIA: SARA- R. Hendrickson (1963, p. 548). B:c. WAK; 2 rivers by this name in Sarawak (mouth Sembrong River. See Endau, Sungai, vicinity. of one at 1°23'N, 110°E; mouth of other at Semitau, Sungai Kapuas; Borneo: Kalimantan, 4°47'N, 1 14°58'E); collected 18 Sep. 1892 by E. INDONESIA; 0°33'N, 111°58'E; collected 15 Bartlett; smk, 1 (skin only, skull inside). FAS; Dec. 1893 by J. Biittikofer (1897, p. 11); rmnh, not mapped. 1. FAS; C:K-6. Sekol, Sungai, 100 m; MALAYSIA: WEST MA- Semongkat; Pulau Sumbawa, Lesser Sunda Is-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 167 lands, INDONESIA; 8°35'S, 1 17°20'E; collected Kalimantan, INDONESIA; ca. 0°27'N, 31 Oct. 1981 by M. Aimi (pers. comm., 3 Aug. 1 17°35'E; 12 troops observed 14 Apr. 1973 by 1990; cf. Kawamoto et al., 1984, p. 132); pri, 1 C C Wilson and W. L. Wilson (1975, p. 257). (skeleton only). Blood samples taken Jul. 1 98 1- FAS; CK-46. Jan. 1982 by Y. Kawamoto, Tb. M. Ischak, and Sengata, Sungai, Kutai Reserve; Borneo: Kali- J. Supriatna (1982, p. 58). FAS; C:LS-8. mantan, INDONESIA; 0°32'N, 117°28'E; ob- Sendang; Pulau Bali, Lesser Sunda Islands, IN- served 2-6 Apr. 1973 by C C Wilson and W.

DONESIA; 8°08'S, 1 14°38'E; collected ca. 1930 L. Wilson ( 1 975, p. 257). Observed 26 Jan. 1 973- by H. J. V. Sody (1933, p. 93); rmnh, 3. FAS; 31 Jan. 1974 by N. A. Fittinghoff, Jr., and D. C:LS-1. G. Lindburg (1980, p. 185). FAS; CK-46. Sendung; Borneo, country unknown; not located, Sengata, Sungai, right bank, 1 km below mouth 7°02'N-4°11'S, 108°51'-119°17'E; collected 24 of Sungai Nubung, Kutai Reserve; Borneo: Ka- Apr. 1902 by Dr. Pagel; zmb, 1 (skin only). FAS; limantan, INDONESIA; ca. 0°32'N, 117°07'E; not mapped. observed 13 Oct. 1985 by A. Suzuki (1986, p. Sengata; Borneo: Kalimantan, INDONESIA; 16). FAS; CK-47. 0°28'N, 1 17°33'E; observed summer 1971 by J. Sengatta River. See Sengata, Sungai. A. Kurland (1973, p. 250). FAS; CK-46. Senimba Bay; Pulau Batam, Kepulauan Riau, IN- Sengata, Sungai, 1 km below Mentoko camp; Bor- DONESIA; 1°05'N, 103°56'E; observed 22 Mar. neo: Kalimantan, INDONESIA; ca. 0°31'N, 1906 by C. B. Kloss (1908, p. 65). FAS; B:SCS- 117°22'E; observed 1983-1989 by A. Suzuki 10. (1989, p. 32). FAS; CK-46. Senoeang, 500 m; Borneo: Kalimantan, northeast, Sengata, Sungai, 2 km below Mentoko camp; Bor- INDONESIA; ca. TOO'N, 11 OWE; collected neo: Kalimantan, INDONESIA; ca. 0°31'N, 29 Aug. 1937 by J. J. Menden; amnh, 1 (skin 117°23'E; observed 1983-1989 by A. Suzuki only; male skin mismatched with female skull); (1989, p. 32). FAS; CK-46. FAS, CK-3. Sengata, Sungai, 3 km below Mentoko camp; Bor- Sentosa Island, SINGAPORE; 1°14'-1°16'N, neo: Kalimantan, INDONESIA; ca. 0°32'N, 103°48'-103°51'E; observed before 1901 by S. 117°24'E; observed 1983-1989 by A. Suzuki S. Rower (1900, p. 316). FAS; B:SCS-7. (1989, p. 32). FAS; C:K-46. Sepaku, Sungai, ca. 350 m NE of camp, 15-50 m; Sengata, Sungai, 5 km below Mentoko camp; Bor- Borneo: Kalimantan, INDONESIA; ca. 0°53'S, neo: Kalimantan, INDONESIA; ca. 0°32'N, 116°43'E; observed 11-13 Mar. 1973 by C C 117°25'E; 2 troops observed 1983-1989 by A. Wilson and W. L. Wilson (1975, pp. 248, 254). Suzuki (1989, p. 32). FAS; CK-46. FAS; CK-50. Sengata, Sungai, 7 km below Mentoko camp; Bor- Sepilok Forest Reserve. See Labuk Road, Sepilok neo: Kalimantan, INDONESIA; ca. 0°33'N, Forest Reserve. 117°26'E; observed 1983-1989 by A. Suzuki Seraja, Pulau. See Seraya Besar, Pulau. (1989, p. 32). FAS; CK-46. Serang Daja. See Serangjaya-hilir. Sengata, Sungai, 9 km below Mentoko camp; Bor- Serang Djaja. See Serangjaya-hilir. neo: Kalimantan, INDONESIA; ca. 0°33'N, Serangjaya-hilir; Sumatra, east coast, INDONE- 1 17°27'E; 2 troops observed 1983-1989 by A. SIA; 4°15'N, 98°12'E; collected Oct.-Nov. 1931 Suzuki (1989, p. 32). FAS; CK-46. by H. J. V. Sody; rmnh, 2 (skulls only). FAS; Sengata, Sungai, 1 1 km below Mentoko camp; B:S-19. Borneo: Kalimantan, INDONESIA; ca. 0°30'N, Serapit Tandjung; Sumatra, INDONESIA; ca. 117°28'E; observed 1983-1989 by A. Suzuki 3°33'N, 98°20'E; collected 1897-1899 by G. (1989, p. 32). FAS; CK-46. Schneider (1905, p. 72); museum and number Sengata, Sungai, 13 km below Mentoko camp; of specimens unknown (not seen). FAS; B:S-20. Borneo: Kalimantan, INDONESIA; ca. 0°30'N, Serasan, Pulau, INDONESIA; 2°28'-2°33'N, 117°29'E; observed 1983-1989 by A. Suzuki 108°58'-109°07'E; observed Sep. 1893 by A. (1989, p. 32). FAS; CK-46. Everett (in Thomas & Hartert, 1894, p. 654). Sengata, Sungai, at Mentoko camp; Borneo: Ka- Collected 8 Jun. 1 900 by W. L. Abbott (see Mil- limantan, INDONESIA; ca. 0°30'N, 117°22'E; ler, 1901, p. Ill); usnm, 1. Collected 27 Aug. observed 1983-1989 by A. Suzuki (1989, p. 32). 1931 by P. M. de Fontaine (see Chasen, 1935a, FAS; CK-46. p. 5); zrc, 1. FAS; B:SCS-36. Sengata, Sungai, below Sengata village; Borneo: Seraya Besar, Pulau, Lesser Sunda Islands, IN-

168 FIELDIANA: ZOOLOGY DONESIA; 8°22'-8°24'S, 1 19°50'-1 19°52'E; re- INDONESIA; ca. 1°01'N, 102°07'E; observed ported present 1969-1973 by W. Auffenberg Nov. 1 97 1-Jan. 1 973 by C. M. Crockett and W. (1981, p. 40). FAS; QLS-13. L. Wilson (1980, p. 156). FAS; B:S-42. Serdang district; Sumatra, INDONESIA; ca. Siak, Sungai, ca. 45 km below Pekanbaru; Su- 3°30'N, 99°00'E; collected 2 Feb. 1905 by H. matra, INDONESIA; ca 0°47'N, 101°44'E; ob-

Diirk (see Schneider, 1 905, map 1); zsbs, 1 . FAS; served Nov. 1971-Jan. 1973 by C. M. Crockett B:S-22. and W. L. Wilson (1980, p. 156). FAS; B:S-42. Sertung, Pulau, INDONESIA; 6°03'-6°07'S, Siak, Sungai, ca. 48 km above mouth; Sumatra, 105°22'-105°24'E; primates reported absent INDONESIA; ca. 0°55'N, 102°05'E; observed 1883-1934 by K. W. Dammerman (1948, p. Nov. 197 1-Jan. 1973 by C. M. Crockett and W. 50); volcanic explosion in 1883. B:k. L. Wilson (1980, p. 156). FAS; B:S-42. Serutu, Pulau, INDONESIA; 1°42'-1°44'S, Siak, Sungai, ca. 60 km above mouth; Sumatra, 108°41'-108°48'E; collected 25 Mar. 1931 by L. INDONESIA; ca. 0°48'N, 102°05'E; observed Coomans de Ruiter and Madzoed (see Chasen, Nov. 197 1-Jan. 1973 by C. M. Crockett and W. 1935b, p. 2); mzb, 1. FAS; B:SCS-21. L. Wilson (1980, p. 156). FAS; B:S-42. Sewela; Pulau Lombok, Lesser Sunda Islands, IN- Siak, Sungai, ca. 60 km below Pekanbaru; Su- DONESIA; 8°32'S, 116°35'E; collected 22-26 matra, INDONESIA; ca 0°45'N, 101°51'E; ob- Mar. 1927 by Sunda-Expedition Rensch (see B. served Nov. 1971-Jan. 1973 by C. M. Crockett Rensch, 1930, p. 12; Mertens, 1930, p. 127; I. and W. L. Wilson (1980, p. 156). FAS; B:S-42. 2. C:LS-7. Rensch, 1934, p. 228); nms, FAS; Siak, Sungai, ca. 75 km below Pekanbaru; Su- S'gobang. See Segobang, Sungai. matra, INDONESIA; ca. 0°48'N, 101°58'E; ob- Shanghai; CHINA; 31°14'N, 121°28'E; captive served Nov. 1971-Jan. 1973 by C. M. Crockett obtained Jul. 1 858 J. Zelebor or (introduced) by and W. L. Wilson (1980, p. 156). FAS; B:S-42. von Frauenfeld (Zelebor, [1869], p. 8; cf. Fit- Siak, Sungai, near mouth; Sumatra, INDONE- zinger, 1861, p. 389). Subspecies uncertain; not SIA; ca. 1°20'N, 1 02° 10'E; observed Nov. 1971- mapped. Jan. 1973 by C. M. Crockett and W. L. Wilson Siaba. See Siaba Besar, Pulau. (1980, p. 156). FAS;B:S-42. Siaba, Teluk; Pulau Nias, INDONESIA; 1°31'N, Siak, Sungai, near Pekanbaru; Sumatra, INDO- 97°24'E; collected 16-20 Mar. 1903 by W. L. NESIA; ca. 0°32'N, 101°27'E; observed Nov. Abbott (see Lyon, 1916, p. 458); usnm, 7 (in- 1971-Jan. 1973 by C. M. Crockett and W. L. cluding 3 skulls only, 2 with external measure- Wilson (1980, p. 156). FAS; B:S-42. ments recorded on skull tags); FAS; B:IO-10. Siak Copatta; Sumatra, INDONESIA; ca. 1°00'N, Siaba Besar, Pulau, Lesser Sunda Islands, IN- 101°00'E; collected ca. 1907 by M. Moszkowski DONESIA; 8°32'-8°33'S, 119°41'-119 42'E; (1909, map following p. 192); zmb, 1 (skull only). monkeys reported absent 1 969-1 973 by W. Auf- FAS; B:S-40. fenberg (1981, p. 40). C:d. Siam. See THAILAND. Siaboh. See Sibaboh, Lugu. "Siam"; "13°45'N, 99°25'E"; col- Siak, Sungai, ca. 12 km above mouth; Sumatra, THAILAND; lected 18 Jun. 1913 and date unknown K. INDONESIA; ca. 1°14'N, 102°10'E; observed by G. Gairdner cf. P. H. Nov. 197 1-Jan. 1973 by C. M. Crockett and W. (1914, pp. 34-36; Napier, 1 1 98 , p. 20, who cited this locality as "E. Siam"); L. Wilson (1980, p. 156). FAS; B:S-42. bm(nh), 2 (skulls Subspecies uncertain; Siak, Sungai, ca. 15 km below Pekanbaru; Su- only). A:T-27. matra, INDONESIA; ca. 0°35'N, 101°34'E; ob- served Nov. 1971-Jan. 1973 by C. M. Crockett Siantan, Pulau, INDONESIA; 3°04'-3°13'N, collected 8 and W. L. Wilson (1980, p. 156). FAS; B:S-42. 106°12'-106°17'E; Sep. 1899 by W. L. cf. Siak, Sungai, ca. 24 km above mouth; Sumatra, Abbott (in Miller, 1900, p. 244; Kloss, INDONESIA; ca. 1°07'N, 102°10'E; observed 1903b, p. 69; Oberholser, 1917, p. 2); usnm, 1. Collected 14 3 Oct. 1925 F. N. Nov. 1 97 1-Jan. 1 973 by C. M. Crockett and W. Sep. and by L. Wilson (1980, p. 156). FAS; B:S-42. Chasen (see Chasen & Kloss, 1 928a, p. 43); zrc, Siak, Sungai, ca. 30 km below Pekanbaru; Su- 2. FAS; B:SCS-30. matra, INDONESIA; ca. 0°42'N, 101°39'E; ob- Sibabo. See Sibaboh, Lugu. served Nov. 1971-Jan. 1973 by C. M. Crockett Sibaboh, Lugu; Pulau Simeulue, INDONESIA; and W. L. Wilson (1980, p. 156). FAS; B:S-42. 2°43'N, 96°06'E; collected 17 Dec. 1901 and 26 Siak, Sungai, ca. 36 km above mouth; Sumatra, Oct. 1902 by W. L. Abbott (field catalog; cf.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 169 Miller, 1903a, p. 437; Lyon, 1916, p. 457);usnm, 105°26'E; collected in 1905 by B. Hagen; zsbs, 4. FUS, B:IO-2. 1 (skin only). FAS; CSCS-18. Sibau, Sungai. See Poelau (?= Kuda), Sungai Si- Simujan River. See Simunjan, Sungai. bau. Simunjan, Sungai; Borneo, MALAYSIA: SARA- Siberut, Pulau, INDONESIA; 0°55'-l°48'S, WAK; ca. 1°17'N, 110°49'E; observed Aug. 1855 98°35'-99°18'E; M. fascicularis reported absent by A. R. Wallace (1869, pp. 69, 82). Collected before 1928 by C. B. Kloss ([1928], p. 802). B:f. 1876-1878 by W. T. Hornaday (1910, p. 358); Sibuga Besar, Sungai; Borneo, MALAYSIA: SA- museum unknown, 2 (not seen). FAS; C:Sar-7. BAH; ca. 5°56'N, 118°03'E; collected 21 Aug. Simunjon River. See Simunjan, Sungai. 1929 by F. C. Wonder (field catalog; cf. Davis, Sinabang; Pulau Simeulue, INDONESIA; 2°29'N, 1962, p. 10); fmnh, 1. FAS; C:Sab-17. 96°23'E; collected 3-14 Feb. 1913 by E. Jacob- Sibugal River. See Sibuga Besar, Sungai. son (1913, p. 356; 1917, p. 276) and W. C. V. Si Chang, Ko, THAILAND; 13°07'-13°1 l'N, Heurn; rmnh, 2 (skins only). Collected Jul. 1913 100°48'-100°50'E; primates reported as appar- by E. Jacobson (1913, p. 356; 1917, p. 276); ently absent Jan. 1915 by C. B. Kloss (1915a, rmnh, 2 (skins only). FUS; B:IO-3. p. 157; 1915b, p. 221). M. fascicularis reported Singapore Botanical Gardens; Singapore Island, present 15 Nov. 1990 by local residents (Aggi- SINGAPORE; 1°18'N, 103°49'E; collected 2 marangsee, 1992, pp. 1 10, 130). Pending further Apr. 1893 by unknown collector; zrc, 1 (skin information, Aggimarangsee's record is accept- only, skull inside). Observed before 1 90 1 by S.

ed as valid. Subspecies uncertain; A:T-3 1 . S. Flower ( 1 900, p. 3 1 6). Collected 26 Jul. 1912 Sidjoendjoeng. See Sijunjung, Padang highlands. by unknown collector; zrc, 1 (skin only). Col- Siemreab, 1 6 km S, 1 50 ft (= 45 m); CAMBODIA; lected 23 Mar. 1925 by museum collector; zrc, 13°22'N, 103°51'E; collected 16 Dec. 1927 by 1. Collected Apr. 1925 by unknown collector; J. Delacour (1929, p. 195) and W. P. Lowe; zrc, 1 (skull only). Collected 11 Feb. 1944 by mnhn, 1. FAS; A:C-1. Oshita; zrc, 1. Observed Jul.-Sep. 1960 by Y. Siem-Reap. See Siemreab. Furuya (1961-1962a, p. 76; 1965, p. 289). Ob- Sijunjung, Padang highlands; Sumatra, INDO- served 29 Jul. 1973 by J. Fooden. FAS; B:SCS-7. NESIA; 0°42'S, 100°58'E; collected 21 Jul. 1877 Singapore Island, northwestern part, , by J. F. Snelleman (1887, p. 10; cf. Hooijer, SINGAPORE; ca. 1°25'N, 103°44'E; reported 1962b, p. 44); rmnh, 1 (skull only). FAS; B:S- present before 1974 by S. H. Chuang (1973, p. 53. 3). FAS; B:SCS-7. Sika; Pulau Flores, Lesser Sunda Islands, INDO- Singapore Island, western part, mangroves, SIN- NESIA; 8°45'S, 122°12'E; observed 21 Nov. GAPORE; ca. 1°20'N, 103°38'E; reported pres- 1888-9 Jan. 1889 by M. Weber (1890a, p. ix; ent before 1974 by S. H. Chuang (1973, p. 3). 1890b, p. 102). FAS; GLS-19. FAS; B:SCS-7. Sikka. See Sika. Singapore Island, SINGAPORE; 1°16'-1°28'N, Sikundur; Sumatra, INDONESIA; ca. 4°00'N, 103°38'-104°00'E; captive acquired Apr. 1858 98°00'E; observed May-Dec. 1971 by J. R. by J. Zelabor ([1869], p. 8; cf. Fitzinger, 1861,

1 1 MacKinnon (1973, p. 240). FAS; B:S-19. p. 389) or von Frauenfeld. Collected [ 870-1 87 ] Simalur Island. See Simeulue, Pulau. by A. B. Meyer (see Steenis-Kruseman, 1950, Simeulue, Pulau, INDONESIA; 2°19'-2°55'N, p. 358), zmb, 1. Collected 22 Mar. 1903 by un- 95°41'-96°30'E; collected 31 Jul. 1924 and 23 known collector; zrc, 1. FAS; B:SCS-7. Dec. 1 925 by unknown collectors; rmnh, 2 (skins Singkarak, Lake. See Pajo, Danau Singkarak, Bal- only). Reported present before 1981 by J. T. ipor district. Marshall (Crockett & Wilson, 1980, p. 156). Singkel vicinity; Sumatra, INDONESIA; ca. Blood samples taken from captives 1984-1989 2°1 7'N, 97°49'E; blood samples taken Nov.-Dec. by W. Scheffrahn, J. R. de Ruiter, and J. A. R. 1986 by J. R. de Ruiter (1993, p. 91). FAS; B:S- A. M van Hooff(1994, p. 135). FUS; B:IO-2, 17. IO-3. Singkil. See Singkel vicinity. Similajau National Park; Borneo, MALAYSIA: Singkil, Gunung; Java, west, INDONESIA; not SARAWAK; ca. 3°38'N, 1 13°20'E; reported as located, 5°52'-7°48'S, 105°12'-108°52'E; col- •probably present before 1983 by G. Davies lected 19-20 Jul. 1909 by H. W. van der Weele; ([1983], p. 148). FAS; C:Sar-15. rmnh, 3. FAS; B:J-8. Simpang; Pulau Bangka, INDONESIA; 1°54'S, Sintang; Borneo: Kalimantan, INDONESIA;

170 FIELDIANA: ZOOLOGY OWN, 1 1 1°30'E; collected 14 Oct.^l Nov. 1897 Kloss, [ 1 928], p. 802; Chaturvedi, 1 980, p. 1 34). by A. Harrison, Jr., and H. M. Hiller; ansp, 5 A:e. (including 2 skins only). FAS; C:K-7. South Banyuwangi Nature Park; Java, INDO- Sinubing; Pulau Natuna Besar, INDONESIA; not NESIA; ca. 8°37'S, 114°28'E; observed 17-28 precisely located, 3°37'-4°13'N, 107°58'- Sep. 1971 by A. Hoogerwerf (1974, p. 24). FAS; 108°25'E; collected Jul. 1 894 by C. Hose; bm(nh), C:J-43. 1 (skin only, skull inside). FAS; B:SCS-32, SCS- South Island. See Pelapis Tengah, Pulau. 33. South Pagi Island. See Pagai Selatan, Pulau. Siolak Daras. See Siulakderas, Kerinci region. Province. See Sumatera Selatan, Sipora Island. See Sipura, Pulau. Propinsi. Sipura, Pulau, INDONESIA; 2°02'-2°24'S, 99°32'- Sriracha. See Si Racha vicinity. 99°52'E; M. fascicularis reported absent before St. Barbe Island. See Pejantan, Pulau. 1928 by C. B. Kloss ([1928], p. 802). B:g. Straits of Malacca. See Langkawi, Pulau. Si Racha vicinity; THAILAND; ca. 13°10'N, Suaka Margasatwa Padang Sugihan; Sumatra, IN- 1 00°56'E; observed Apr.-May 1 9 1 2 by N. Gyld- DONESIA; not located; observed Sep. 1988 by enstolpe (1914, p. 3). FAS; A:T-31. M. Bismark (1992, p. 1 1). FAS; not mapped. Sirhassen Island. See Serasan, Pulau. Subi-kecil, Pulau, INDONESIA; 3°01'-3°03'N, Sittwe harbor. See Myengun Kyun. 108°51'-108°53'E; collected 5 Aug. 1931 by P. Siulakderas, Kerinci region, 3000 ft (= 900 m), M. de Fontaine (see Chasen, 1935a, p. 5); zrc, Sumatra, INDONESIA; 1°55'S, 101°18'E; col- 3. FAS; B:SCS-35. lected 27 Mar. 1914 by H. C. Robinson and C. Sugi, Pulau, Kepulauan Riau, INDONESIA; 0°45'- collected 24 B. Kloss (1918, p. 6; in Pendlebury, 1936, p. 9); 0°53'N, 103°43'-103°5rE; Aug. bm(nh), 1. FAS; B:S-49. 1902 by W. L. Abbott (see Miller, 1906c, p. 1. B:SCS-16. Slamet, Mt. See Baturaden; Kaligoea. 281); usnm, FAS; Suka Smitau. See Semitau, Sungai Kapuas. Bandjar, northeast; Sumatra, INDONESIA; Soekadono. See Sukadana. ca. 4°55'S, 104°02'E; observed Nov. 1971-Jan. 1 973 by C. M. Crockett and W. L. Wilson (1 980, Soekaranda; Sumatra, INDONESIA; not located, 5°38'N-5°57'S, 95°12'-106°05'E; collected Aug. p. 156). FAS;B:S-77. Suka Bandjar, southeast; Sumatra, INDONESIA; 1894 by Ude; zmb, 1 (skull only). FAS; not ca. observed Nov. 1971-Jan. mapped. 5°00'S, 103°57'E; 1 973 by C. M. Crockett and W. L. Wilson (1980, Soengei Slan. See Sungaiselan. p. 156). FAS;B:S-77. Soengi Han. See Sungaiselan. Sukadana, 100 m; Sumatra, INDONESIA; 5°05'S, Soliga; Pulau Nias, INDONESIA; ca. 1°02'N, 105°33'E; 29-30 Jul. 1934; collected by J. J. 97°33'E; collected 25 Jul. 1937 by B. Lawrence Menden; amnh, 4. FAS; B:S-83. (see Schauensee & Ripley, 1 940a, map following Sulawesi, Pulau, INDONESIA; 1°45'N-5°43'S, p. 368); mcz, 2. FAS; B:IO-7. 1 18°45'-125°15'E; introduced, reported present Solombo, Pulau. See Masalembu Besar, Pulau. before 1841 by S. Miiller ([1840], p. 17). Sub- Solor, Pulau, Lesser Sunda Islands, INDONESIA; species uncertain; not mapped. 8°26'-8°37'S, 122°53'-123°11'E; reported pres- Sullivan's Island. See Lanbi Kyun. ent before 1937 by R. Mertens (1936, p. 319). Sumatera Selatan, Propinsi (province); Sumatra, FAS; CLS-20. INDONESIA; 1°39'-4°53'S, 101°57'-106°1 l'E; Songkhla. See Khao Noi/Khao Tangkuan. blood samples taken Jan.-Nov. 1 979 by Y. Ka- Sontra 0-300 ca. Peak, m; VIETNAM; 16°07'N, wamoto and Tb. M. Ischak(1981,p. 238). FAS; 1974 L. 108°18'E; observed Jun.-Aug. by K. not mapped. Lippold p. 521). A:V-1. (1977, FAS; Sumatra, [east-central], INDONESIA; ca. 1°00'N, Sontra Peak, 3.9 km W, 0.3 km S, 150-600 m; 101°00'E; collected ca. 1907 by M. Moszkowski VIETNAM; ca. 16°07'N, 108°18'E; collected 14 (1909, maps following pp. 192, 328); zmb, 4 P. F. Sep. 1967 by D. Van Peenen (see Van (including 3 skulls only). FAS; B:S-40. Peenenetal., 1969, pp. 100,290; 1971, p. 134); Sumatra, east coast, INDONESIA; 5°38'N-5°57'S, usnm, 1. FAS; A:V-1. 95°12'-106°05'E; collected date unknown by H. South Andaman Island, INDIA; 1 1°29'-12°15'N, Diirk; zsbs, 1 5 (including 4 skins only, 4 skulls 92°31'-92°48'E; primates reported absent be- only). FAS; not mapped. fore 1903 by G. S. Miller, Jr. (1902b, p. 792; cf. Sumatra, INDONESIA; 5°38'N-5°57'S, 95°12'-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 171 106°05'E; collected 7 Sep. 1 82 1 by A. Duvaucel; specified portion ofPulau Bangka collection (see mnhn, 1 (skin only). Collected in 1889 by Dr. Bangka, Pulau). FAS; B:SCS-19. Moesch; zmuz, 1 (skin only, skull inside). Col- Sungai Tekam Forestry Concession, 250-350 m; lected 1896-1897 by unknown collector; nms, 2 MALAYSIA: WEST MALAYSIA; 4°10'N, (skulls only). Collected 10 Feb. and 5 Jun. 1897 102°40'E; observed 1979-1981 by A. D. Johns by Petersen; zmb, 2 (skulls only). Collected date (1981, p. 222). FAS; B:WM-12. unknown by Petersen; zmb, 7 (including 4 skins Sungei Adang; Ko Tarutao, THAILAND; not pre- only, 3 skulls only). Collected 1905-1910 by C. cisely located, 6°3 1 '-6°44'N, 99°36'-99°42'E; Bruegel; zsbs, 52 (including 22 skins only, 1 1 collected 10-11 Mar. 1909 by museum collec- skulls only). Collected date unknown by C. tors (see H. C. Robinson & Kloss, 1910, p. 666); Bruegel; zsbs, 1 (skin only). Collected in 1924 bm(nh), 2. FAS; A:T-59. by P. Wirz; NHMBa, 2. Collected 10 Apr. 1926- Sungei Bernam. See Changkat Mentri. 18 Aug. 1930 by C. Blazer; rmnh, 4 (including Sungei Biru; Pulau Bintan, INDONESIA; ca. 3 skulls only). Collected 3 Feb. 1938 by W. 1°12'N, 104°33'E; collected Jun.-Sep. 1908 by Groenvelt, rmnh, 1 (skull only). Collected be- H. C. Robinson (in Thomas & Wroughton, fore 1942 by L. Coomans de Ruiter; rmnh, 1 1909b, pp. 100, 104); bm(nh), 3; zrc, 1 (skull (skull only). Collected 3 May 1943 by unknown only). FAS; B:SCS-8.

collector; usnm, 1 . Collected date unknown by Sungei Buloh. See Kampong Sungai Buloh. Dr. Porter; zsbs, 1 (skull only). Collected date Sungei Hdang. See Sungei Adang. unknown by unknown collector; nms, 2 (skulls Sungei Pandang. See Pandan, Sungai. only). FAS; not mapped. Sungsan, ca. 20 km NNE; Sumatra, INDONESIA; Sumatra, south, INDONESIA; 5°38'N-5°57'S, ca. 5°15'S, 105°50'E; observed Nov. 1971-Jan. 95°12'-106°05'E; collected date unknown by W. 1973 by C. M. Crockett and W. L. Wilson (1980, Volz; zmb, 2 (skins only). FAS; not mapped. p. 156). FAS;B:S-85. Sungsan, ca. 50 km N; Sumatra, INDONESIA; Sumatra, west coast, INDONESIA; 5°38'N-5°57'S, ca. observed Nov. 1971-Jan. 95°12'-106°05'E; collected before 1883 by F. 4°58'S, 105°48'E; 1973 by C. M. Crockett and W. L.Wilson (1980, von Faber; bm(nh), 1 . FAS; not mapped. p. 156). FAS; B:S-84. Sumbawa, Pulau, Lesser Sunda Islands, INDO- Suranadi; Pulau Lombok, Lesser Sunda Islands, NESIA; 8°05'-9°07'S, 116°42'-119 12'E; re- INDONESIA; 8°33'S, 116°15'E; collected 10 ported present before 1 906 by T. Willink ( 1 905, Dec. 1981 by M. Aimi (pers. comm., 3 Aug. p. 175). Collected in 1914 by Dr. Pannekoek; 1990; cf. Kawamoto et al., 1984, p. 132); pri, 1 NHMBa, 1 (skull only). FAS; C:LS-8, LS-10 (skeleton only). Morphometric study Dec. 1981 through LS- 12. by M. Aimi, A. Bakar, and J. Supriatna (1982, Sumbawang, Sungei. See Sembawang, Sungai. p. 52). Blood samples taken Jul. 1981-Jan. 1982 Sunda Islands; INDONESIA; 5°38'N-10°23'S, by Y. Kawamoto, Tb. M. Ischak, and J. Su- 95°1 2'-127°l 8'E; collected in 1 899 by P. Sarasin priatna (1982, p. 58). FAS; GLS-5. and F. Sarasin; NHMBa, 1 Collected (skull only). Sur Sdei. See Pang Roloem-Sur Sdei area. date unknown by von Altenstein; zmb, 1 (skin Survey Site LB, 5-50 m; Pulau Simeulue, IN- only). FAS; not mapped. DONESIA; ca. 2°35'N, 96°10'E; 2 troops ob- Sundarbans. See Khulna district, Sundarbans. served Jan. 1982 and 1 1 Mar.-3 Apr. 1984 by Nature Reserve; Sungai Kayan-Sungai Mentarang J. Sugardjito, C. P. van Schaik, N. A. van Borneo: Kalimantan, INDONESIA; ca. 3°00'N, Noordwijk, and T. Mitrasetia (1989, p. 198). 115°30'E; reported present before 1983 by G. FUS; B:IO-2. Davies GK-36. ([1983], p. 148). FAS; Survey Site LS, 10-120 m; Pulau Simeulue, IN- See Abai. Sungai Kinabatangau. DONESIA; ca. 2°44'N, 95°54'E; 2 troops ob- Sungailundang; Sumatra, INDONESIA; 1°05'S, served Jan. 1982 and 11 Mar.-3 Apr. 1984 by 100°29'E; morphological study Jun.-Nov. 1980 J. Sugardjito, C. P. van Schaik, N. A. van by A. Bakar, M. Amir, and Marshal (1981, p. Noordwijk, and T. Mitrasetia (1989, p. 198). 11). FAS; B:S-47. FUS; B:IO-2. Sungoxseldin; Pulau Bangka, INDONESIA; 2°24'S, Survey Site SEM, 210 m; Pulau Simeulue, IN- 105°59'E; collected in 1900 by A. A. W. Hu- DONESIA; ca. 2°45'N, 96°02'E; 5 troops ob- brecht([1895], pp. 33, 88; cf Kohlbrugge, 1902, served Jan. 1982 and 1 1 Mar.-3 Apr. 1984 by p. 322; Zuckerman, [1933], p. 1062); zluu, un- J. Sugardjito, C. P. van Schaik, N. A. van

172 FIELDIANA: ZOOLOGY Noordwijk, and T. Mitrasetia (1989, p. 198). Tamiang; Sumatra, northeast, INDONESIA; ca. FUS; B:IO-2. 4°25'N, 98°16'E; collected Nov. 1931 by H. J. Swela. See Sewela. V. Sody (see Hooijer, 1962b, p. 45); rmnh, 1 Syonan. See Singapore Island, SINGAPORE. (skull only). FAS; B:S- 18. Tabanan. See Kukuh. Tana Bala. See Tanahbala, Pulau. Tagoot (?= Tagu), Great Tenasserim River; BUR- Tanahbala, Pulau, INDONESIA; 0°15'-0°37'S, MA; 12°15'N, 99°03'E; collected 18 Apr. 1914 98°17'-98°30'E; collected 4 Feb. 1903 by W. L. by G. C. Shortridge (in Wroughton, 1915, p. Abbott (see Lyon, 1916, p. 459); usnm, 2. Re- 697; cf. Moore & Tate, 1965, p. 332); bnhs, 2. ported Nov. 1971-Jan. 1973 by local residents AUR; A:Bu-19. (Crockett & Wilson, 1980, p. 156). FAS; B:IO- Tahan, Sungai; MALAYSIA: WEST MALAY- 13. SIA; ca. 4°35'N, 102°18'E; observed Jul.-Sep. Tanah Laut. See Pleihari Tanah Laut Game Sanc- 1960 by Y. Furuya (1965, p. 287). FAS; B:WM- tuary. 11. Tanahmasa, Pulau, INDONESIA; 0°01'N-0°23'S, Tahang River. See Tahan, Sungai. 98°17'-98°34'E; collected 19 Feb. 1903 by W. Takengon, ca. 15 km NNW; Sumatra, INDO- L. Abbott; usnm, 1 (skull only). FAS; B:IO-12. NESIA; ca. 4°45'N, 96°47'E; reported present Tanah Puteh. See Kampong Tanah Puteh. Nov. 1971-Jan. 1973 by local residents (Crock- Tandjong. See Tanjung. et & Wilson, 1980, p. 156). FAS; B:S-9. Tandjung; Sumatra, INDONESIA; ca. 3°17'N, Takengon, ca. 40 km NNW; Sumatra, INDO- 99°19'E; collected 1897-1899 by G. Schneider ca. NESIA; 4°58'N, 96°42'E; reported present (1905, p. 72); museum and number of speci- 1973 local residents Nov. 1971-Jan. by (Crock- mens unknown (not seen). FAS; B:S-23. ett & Wilson, 1980, p. B:S-8. 156). FAS; Tandjung Bringin; Sumatra, INDONESIA; ca. Takokak ca. Reserve; Java, INDONESIA; 7°03'S, 3°47'N, 98°24'E; collected 1897-1899 by G. 106°59'E; present before 1972 un- reported by Schneider (1905, p. 72); museum and number specified informant 1971, p. 276). (iucn, FAS; of specimens unknown (not seen). FAS; B:S-20. B:J-8. Tandjung Butus; Sumatra, INDONESIA; ca. Tale Sap. See Nang Kham, Ko. 3°47'N, 98°24'E; collected 1897-1899 by G. Talibang, near Kota Kinabalu, near sea level; Bor- Schneider (1905, p. 72); museum and number neo, MALAYSIA: SABAH; ca. 6°H'N, of specimens unknown (not seen). FAS; B:S-20. 116°14'E; collected 15-24 Aug. 1937 by J. A. Tandjung Laut. See Tandjung. Griswold, Jr. (see Coolidge, 1940, p. 123); mcz, Tangarveng Island, Sungai Mahakam; Borneo: 4 (including 1 skeleton only). FAS; C:Sab-5. Kalimantan, INDONESIA; ca. 0°24'S, 1 16°58'E; Talibong. See Talibang. collected 2 1 Jun. 1 9 1 2 by H. C. Raven (see Deig- Talibong, Ko, northwest, THAILAND; ca. 7°17'N, nan, [1960], p. 267); usnm, 1. FAS; CK-49. 99°23'E; Collected 3 Jan. 1917 by H. C. Rob- Tanjang Sau. See Tanjong Sauh. inson (1917, p. 132); zrc, 1. FAS; A:T-56. Tanjang Turut. See Tanjong Turut. Talibong, Ko, THAILAND; 7°12'-7 17'N, 99°22'- Tanjong Batu. See Batu, Tanjung. 99°27'E; collected 27 Feb. 1 896 by W. L. Abbott Tanjong Pamuju. See Pamuja, Tanjung. (see Riley, 1938, p. 12); usnm, 1. FAS; A:T-56. MALAYSIA: Tamandjaija, Bantam region; Java, INDONESIA; Tanjong Panjair, Sungai Rompin; ca. col- 6°52'-7°00'S, 105°12'-106°33'E; collected 15 WEST MALAYSIA; 2°49'N, 103°29'E; lected 2 1 9 1 9 museum Jan. 1910 by O. Bryant; mcz, 1. FAS; B:J-12. Sep. by collector; bm(nh), Taman Negara. See Negara, Taman. 1;zrc, 1. FAS; B:WM-22. Taman Rimba Templer. See Templer Park. Tanjong Rengsam. See Rengsam, Tanjung. Tamansari, 1600 ft (= 490 m); Java, INDONE- Tanjong Sauh; Pulau Batam, east coast, Kepu- SIA; 7°57'S, 1 11°31'E; collected 20 Jan. 1920 lauan Riau, INDONESIA; ca. 1°07'N, 104°09°E; collected C. by C. B. Kloss (see Weitzel et al., 1988, p. 144); 10-11 Jul. 1908 by H. Robinson

bm(nh), 1; zrc, 1. FAS; C:J-31. (in Thomas & Wroughton, 1909b, p. 104) and Tambelan Besar, Pulau, INDONESIA; 0°57'- E. Seimund; bm(nh), 1; bnhs, 1. FAS; B:SCS- 1°02'N, 107°32'-107°36'E; observed 8-15 Aug. 10. 1899 by W. L. Abbott (in Miller, 1900, p. 244; Tanjong Tuan, Keramat; MALAYSIA: WEST cf. Kloss, 1903b, p. 62). FAS; B:SCS-26. MALAYSIA; 2°24'N, 1 1°52'E; collected 6 Apr. Tambelan Islands. See Benua, Pulau; Uwi, Pulau. 1920 by unknown collector; zrc, 1. Observed

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 173 Aug.-Dec. 1970 by C. H. Southwick and F. C. Tatawa, Pulau, Lesser Sunda Islands, INDONE- Cadigan, Jr. (1972, p. 13). FAS; B:WM-24. SIA; 8°30'-8°31'S, 119°38'-119 39'E; monkeys Tanjong Turut; Pulau Batam, east coast, Kepu- reported absent 1969-1973 by W. Auffenberg lauan Riau, INDONESIA; ca. 1°07'N, 104°09°E; (1981, p. 40). C:d. collected 12 Jul. 1908 by H. C. Robinson (in Taungbyauk Chaung, Tavoy River, Tavoy dis- Thomas & Wroughton, 1909b, p. 104) and E. trict; BURMA; ca. 13°45'N, 98°26'E; collected Seimund; bm(nh), 2. FAS; B:SCS-10. 29-30 Apr. 1936 by H. C. Smith; bm(nh), 4. Tanjung; Borneo: Kalimantan, southeast, IN- AUR; A:Bu-14. DONESIA; 2°1 l'S, 1 15°23'E; collected in 1905 Tavoy River, mouth, Tavoy district; BURMA; by A. Buxtorf; NHMBa, 1 (skull only). FAS; C:K- 14°02'N, 98°12'E; collected 21 Apr. 1936 by H. 30. C. Smith; bm(nh), 2. AUR; A:Bu-13. Tanjungmorawa, Kabupaten Deli Serdang; Su- Tawao. See Tawau. matra, INDONESIA; 3°31'N, 98°49'E; collect- Tawar-See. See Bur ni Bebuli. ed in 1882 and 1883 by B. Hagen (1890, p. 81); Tawau; Borneo, MALAYSIA: SABAH; 4°15'N, rmnh, 3 (including 2 skulls only, 1 skeleton only). 117°54'E; collected 2 Jan. 1910 by R. C. An- Collected in 1 906 and date unknown by C. Brue- drews (1943, p. 69); amnh, 1. FAS; C:Sab-24. gel; zsbs, 8 (including 3 skins only, 3 skulls only). Tawau Hills National Park; Borneo, MALAYSIA: Collected 9 Jan. 1908 by L. Weigand; zsbs, 1. SABAH; ca. 4°20'N, 1 18°00'E; reported present FAS; B:S-21. before 1983 by G. Davies ([1983], p. 148). FAS; Tanjung Puting National Park; Borneo: Kaliman- C:Sab-23. tan, INDONESIA; 2°45'-2°48'S, 1 1 1°57- TayNinh, 100 m; VIETNAM; 1 1°18'N, 106°06'E; 112°01'E; reported present Dec. 1974-Jul. 1975 collected 19 Jan. 1928 by J. Delacour (1928, p. by J. Supriatna, B. O. Manullang, and E. Soe- 268; 1929, p. 196) and W. P. Lowe; bm(nh), 1 kara (1986, p. 186). Reported present before (skin only). FAS; A:V-5. 1983 by G. Davies ([1983], p. 148). Observed Tebingtinggi, ca. 25 km SSE; Sumatra, INDO- in 1983 by K. S. MacKinnon (1986, p. 112). NESIA; ca. 3°07'N, 99°15'E; observed Nov. FAS; :K-10. 1971-Jan. 1973 by C. M. Crockett and W. L. Tapanuli, Teluk; Sumatra, west, INDONESIA; Wilson (1980, p. 156). FAS; B:S-23. 1°38'N, 98°45'E; collected 21 Feb. and 19 Mar. Tebingtinggi, ca. 35 km ESE; Sumatra, INDO- 1902 by W. L. Abbott (see Miller, 1903a, p. NESIA; ca. 3°10'N, 99°25'E; observed Nov. 438); usnm, 2. FAS; B:S-32. 1971-Jan. 1973 by C. M. Crockett and W. L. Tapanuli Bay. See Tapanuli, Teluk. Wilson (1980, p. 156). FAS; B:S-23. Tapung-kanan, Sungai; Sumatra, INDONESIA; Tebingtinggi, ca. 55 km ESE; Sumatra, INDO- 0°46'N, 101°06'E; collected ca. 1907 by M. NESIA; ca. 3°04'N, 99°36'E; observed Nov. Moszkowski (1909, p. 149); zmb, 1. FAS B:S- 1971-Jan. 1973 by C. M. Crockett and W. L. 41. Wilson (1980, p. 156). FAS; B:S-24. Tarusan, Teluk; Sumatra, INDONESIA; 1°13'S, Tebingtinggi, ca. 60 km SSW; Sumatra, INDO- 100°25'E; collected 27 Dec. 1904 by W. L. Ab- NESIA; ca. 2°54'N, 99°00'E; reported present

bott; usnm, 1 . FAS; B:S-47. Nov. 1971-Jan. 1973 by local residents (Crock- Tarusan Bay. See Tarusan, Teluk. ett & Wilson, 1980, p. 156). FAS; B:S-30. Tarutao, Ko, THAILAND; 6°3 1 '-6°44'N; 99°36'- Tebingtinggi, ca. 65 km SE; Sumatra, INDONE- 99°42'E; collected 19 Nov. 1903 and 10 Apr. SIA; ca. 2°58'N, 99°38'E; observed Nov. 1971- 1904 by W. L. Abbott (see H. C. Robinson & Jan. 1973 by C. M. Crockett and W. L. Wilson Kloss, 1910, p. 666; Riley, 1938, p. 15); usnm, (1980, p. 156). FAS;B:S-24. 3 (including 1 skull only). FAS; A:T-59. Tebingtinggi, ca. 75 km SE; Sumatra, INDONE- Tasikmalaja. Tasikmalaya. SIA; ca. 2°54'N, 99°42'E; observed Nov. 1971- Tasikmalaya; Java, INDONESIA; 7°20'S, 108°12'E; Jan. 1973 by C. M. Crockett and W. L. Wilson collected Apr -Aug. 1 926 by Dr. Kopstein; mzb, (1980, p. 156). FAS;B:S-24. 7 (including 1 skin only). FAS; B:J-4. Tebingtinggi, ca. 75 km SSW; Sumatra, INDO- Tasikmalaya, Preanger region, 1 145 ft (= 350 m); NESIA; ca. 2°43'N, 98°53'E; reported present Java, INDONESIA; ca. 7°20'S, 108°12'E; col- Nov. 1971-Jan. 1973 by local residents (Crock- lected 18 Jan. 1908 by G. C. Shortridge (see ett & Wilson, 1980, p. 156). FAS; B:S-29. Thomas & Wroughton, 1909a, p. 373); bm(nh), Tebingtinggi vicinity; Sumatra, INDONESIA; ca. 1. FAS; B:J-4. 3°20'N, 99°09'E; observed Nov. 1971-Jan. 1973

174 FIELDIANA: ZOOLOGY by C. M. Crockett and W. L. Wilson (1980, p. Telukkayubutih, ca. 15 km E; Sumatra, INDO- 156). FAS;B:S-23. NESIA; ca. 1°09'S, 102°06'E; observed Nov. Tekek; Pulau Tioman, MALAYSIA: WEST MA- 1971-Jan. 1973 by C M. Crockett and W. L. LAYSIA; ca. 2°49'N, 104°10'E; observed Mar.- Wilson (1980, p. 156). FAS; B:S-51. Apr. 1962 by Lord Medway (1966, p. 16). FAS; Telukkayubutih, ca. 15 km NNE; Sumatra, IN- B:SCS-3. DONESIA; ca. 1°06'S, 102°02'E; observed Nov. o o Telaga, Pulau, INDONESIA; 3 01'-3 05'N, 1971-Jan. 1973 by C. M. Crockett and W. L. 105°58'-105°59'E; observed 13-1 6 Sep. 1899 by Wilson (1980, p. 156). FAS; B:S-51. W. L. Abbott (in Miller, 1900, p. 245; cf. Kloss, Telukkayubutih, ca. 1 5 km WNW; Sumatra, IN- 1903b, p. 73). FAS; B:SCS-29. DONESIA; ca. 1°06'S, 101°52'E; observed Nov. Telang, 20 m; Borneo: Kalimantan, INDONESIA; 1971-Jan. 1973 by C. M. Crockett and W. L. 2°30'S, 115°24'E; collected 15 Feb. 1971 by Wilson (1980, p. 156). FAS; B:S-51. NAMRU 2 Djakarta Detachment (see Van Pee- Telukkayubutih, ca. 30 km ESE; Sumatra, IN- nen et al., 1974, p. 392); usnm, 1. FAS; C:K- DONESIA; ca. 1°12'S, 102°13'E; observed Nov. 29. 1971-Jan. 1973 by C. M. Crockett and W. L. Telapa Burok, Gunong. See Telapak Burok, Gun- Wilson (1980, p. 156). FAS; B:S-51. ong. Telukkayubutih, ca. 30 km WNW; Sumatra, IN- Telapak Burok, Gunong, 605-1060 m; MALAY- DONESIA; ca. 1°06'S, 101°44'E; observed Nov. SIA: WEST MALAYSIA; 2°49'N, 102°04'E; re- 1971-Jan. 1973 by C M. Crockett and W. L. ported as probably present Jul. 1978-Jun. 198 1 Wilson (1980, p. 156). FAS; B:S-51. by C. W. Marsh and W. L. Wilson (1981, p. Telukkayubutih, ca. 45 km ESE; Sumatra, IN- 232). FAS; B:WM-23. DONESIA; ca. 1°18'S, 102°18'E; observed Nov. Telibon Island. See Talibong, Ko, THAILAND. 1971-Jan. 1973 by C. M. Crockett and W. L. Telok Anson, near sea level; MALAYSIA: WEST Wilson (1980, p. 156). FAS; B:S-51. MALAYSIA; 4°02'N, 101°01'E; collected 27 Telukkayubutih, ca. 60 km SE; Sumatra, IN- Apr. 1915 by N. Gyldenstolpe (1917a, p. 5); DONESIA; ca. 1°23'S, 102°21'E; observed Nov. nhrm, 2 (skulls only; skins reported by Gyld- 1971-Jan. 1973 by C. M. Crockett and W. L. enstolpe, 1917a, p. 25). FAS; B:WM-16. Wilson (1980, p. 156). FAS; B:S-51. Telok Bahang; Pulau Pinang [1], MALAYSIA: Telukkayubutih vicinity; Sumatra, INDONESIA; WEST MALAYSIA; 5°28'N, 100°13'E; collect- 1°11'S, 101°59'E; observed Nov. 1971-Jan. 1973 ed 12 Mar -3 Apr. 1911 by E. Seimund (see by C. M. Crockett and W. L. Wilson (1980, p.

Weitzel et al., 1988, p. 102); bm(nh), 1; zrc, 6. 156). FAS;B:S-51. FAS; B:SM-2. Telukpanji, Kotapinang region; Sumatra, IN- Telok Betong. See Telukbetung. DONESIA; 2°02'N, 100°14'E; collected 6 Jun. Telok Dalam. See Dalam, Lhok. 1937 by M. Boogaarts; zrc, 1. FAS; B:S-36. Telok Paanji. See Telukpanji, Kotapinang region. Teluk Terima. See Trima, Teluk. Telok Wau; Ko Tarutao, west coast, THAILAND; Temaju, Pulau, INDONESIA; 0°28'-0°31'N, o ca. 6°35'N, 99°35'E; collected 25-28 Dec. 1916 108 51'-108°52'E; monkeys reported absent 5- 6 by H. C. Robinson (1917, p. 132); bm(nh), 1; May 1907 by W. L. Abbott (in Lyon, 1911, zrc, 1. FAS; A:T-59. p. 59). B:n. Telom, Sungai, 400 ft (= 120 m); MALAYSIA: Tembeling, Sungai; MALAYSIA: WEST MA- WEST MALAYSIA; ca. 4°18'N, 101°48'E; col- LAYSIA; ca. 4°30'N, 102°15'E; observed Jul.- lected 4 Mar. 1932 by F. A. B. Holloway and Sep. 1960 by Y. Furuya (1965, p. 287). FAS; A. S. Vernay; bm(nh), 2. FAS; B:WM-8. B:WM-11. Teluk Anson. See Telok Anson. Tempasuk, Sungai, Kampong Kiau region, Mount Telukbetung; Sumatra, INDONESIA; 5°27'S, Kinabalu, ca. 2000 ft (= 600 m); Borneo, MA- 105°16'E; collected before 1907 by W. Denna LAYSIA: SABAH; ca. 6°05'N, 116°27'E; col- (see Elliot, 1906, p. 49); fmnh, 1. Clinical ex- lected 9 Aug. 1937 by J. A. Griswold, Jr. (1939b, amination Aug.-Nov. 1 980 by K. Matsubayashi p. 514); mcz, 1. FAS; C:Sab-7. and D. Sajuthi (1981, p. 48). FAS; B:S-82. Templer Park, west section, near waterfall; MA- Teluk Jolo, 12 km N; Borneo: Kalimantan, IN- LAYSIA: WEST MALAYSIA; ca. 3°18'N, DONESIA; ca. 0°18'S, 1 14°03'E; observed 27- 101°36'E; observed before 1972 by M. Nadcha- 28 Aug. 1 986 by D. J. Chivers and K. M. Burton tram (1971, p. 147). FAS; B:WM-19. ([1991], p. 143). FAS; CK-19. Tenasserim; BURMA; 12°05'N, 99°01'E; collect-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 175 ed 4 Mar.-9 Apr. 1914 by G. C. Shortridge (in Tham Chomphon; THAILAND; 13°37'N, Wroughton, 1915, p. 696); bm(nh), 5 (including 99°36'E; blood samples taken Aug.-Sep. 1988 1 skull only); bnhs, 3; fmnh, 1; zrc, 3. AUR; by Y. Kawamoto, T. Ishida, J. Suzuki, O. Tak- A:Bu-19. enaka, and P. Varavudhi (1989, p. 95). Ob- Tenasserim River. See Great Tenasserim River served 20 Dec. 1 990 by N. Aggimarangsee (1992, and Tagoot (?= Tagu), Great Tenasserim River. pp. Ill, 127, pers. comm, Oct. 1993). FAS; Tengger, Pegunungan, 4000 ft (= 1200 m); Java, A:T-28. INDONESIA; 7°55'S, 112°55'E; collected be- Tham Chompol. See Tham Chomphon. fore 1897 by J. H. F. Kohlbrugge (1896b, p. Tham Horn, 4 km W of Ban Thap Plik, ca. 75 m; 280); museum unknown, 1 (not examined). FAS; THAILAND; ca. 8°11'N, 98°53'E; collected 3 GJ-36. Jun. 1973 by J. Fooden ([1975], p. 98); fmnh, Tenghilan Road. See Tuaran-Kampong Tenghilan 1. FAS; A:T-60. Road. Thap Salao, Huai, 200 m; THAILAND; 15°38'N, Tengi, Sungai, 5 m; MALAYSIA: WEST MA- 99°18'E; observed 15 Jul. 1977 by A. A. Eudey LAYSIA; 3°3 1 'N, 101°17'E; observed Jul. 1978- (1979, pp. 90, 198). Subspecies uncertain; A:T- Jun. 1981 by C. W. Marsh and W. L. Wilson 19. (1981, p. 232). FAS; B:WM-18. Thateng, Muang, Plateau des Bolovens; LAOS; Tenompok, 4700 ft (= 1430 m) and 4900 ft (= 15°26'N, 106°23'E; collected 29 Jan.-13 Feb. 1490 m); Borneo, MALAYSIA: SABAH; ca. 1932 by T. D. Carter (see Legendre, 1932, p. 6°00'N, 116°32'E; collected 6-29 Jun. 1937 by 495; 1936, p. 251); amnh, 3 (including 2 skins J. A. Griswold, Jr. (1939a, p. 410); mcz, 2. FAS; only); ansp, 2 (external measurements recorded CSab-7. in amnh catalog). AUR/FAS/MUL; A:L-1. Tenompok Pass. See Kampong Kiau-Tenampok Thung Thong Waterfowl Reserve; THAILAND; Pass, trail between. 8°50'N, 99°1 5'E; observed 22 Jul. 1 976-1 1 Mar. Terbanjawan; Sumatra, INDONESIA; ca. 3°32'N, 1977 by P. J. Storer (1978, p. 113). AUR/FAS/ 98°38'E; collected 1897-1899 by G. Schneider MUL; A:T-48. (1905, p. 72); museum and number of speci- Tibang, Mt.; Borneo: Kalimantan, INDONESIA; mens unknown (not seen). FAS; B:S-21. 1°39'N, 114°34'E; collected Nov. 1925 by E. Terengganu, Sungai, vicinity; MALAYSIA: WEST Mjoberg (1929, p. 117); mcz, 1. FAS; C:K-34. MALAYSIA; ca. 5°00'N, 103°00'E; reported Tikus, Pulu, Cocos Islands, AUSTRALIA; 12°06'S, present in 1968 by D. Chivers (1971, p. 80). 96°54'E; introduced Jun. 1905-Sep. 1906, spe- FAS; B-.WM-10. cies identification tentative (F. W. Jones, 1910, Terima, Teluk. See Trima, Teluk. p. 298). Subspecies uncertain; not mapped. Teris, Sungai, ca. 3 km S of Bukit ; MA- Tillanchong Island, Nicobar Islands, INDIA; LAYSIA: WEST MALAYSIA; ca. 3°35'N, 8°26'-8°35'N, 93°37'-93°39'E; primates report- 102°1 l'E; observed Jul.-Dec. 1977 by D. Chiv- ed absent before 1903 by G. S. Miller, Jr. (1902b, ers and G. Davies (1979, p. 20). FAS; B:WM- p. 792; cf. Kloss, 1903a, p. 114). A:j. 15. Tilu, Gunung; Java, west, INDONESIA; 7°33'S, Terutau, Pulo. See Tarutao, Ko. 107°57'E; collected 6 May 1910 by H. W. van Thaget. See Thagyet, Little Tenasserim River. der Weele; rmnh, 1. FAS; B:J-5. Thagyet, Little Tenasserim River; BURMA; Timau, Fatu, northeast, 1200 m; Pulau Timor, 12°06'N, 99°07'E; collected 25 Mar. 1914 by G. Lesser Sunda Islands, INDONESIA; ca. 9°35'S, C. Shortridge (in Wroughton, 1915, p. 697); 1 23°55'E; collected 20 May 1 9 1 1 by C. B. Haniel

bm(nh), 2; bnhs, 3; fmnh, 1; zrc, 1. AUR/FAS/ (see Hellmayr, 1914, p. 5); zsbs, 1. FAS; C:LS- MUL; A:Bu-19. 26. THAILAND; 5°37'-17°48'N, 97°21'-105°39'E; Timor, Pulau, Lesser Sunda Islands, INDONE- collected 1858-1861 by H. Mouhot (1864, map SIA; 8°19'-10°23'S, 123°27'-27°18'E; obtained at end of vol. 2); bm(nh), 1. Collected in 1862 ca. 1846 by W. H. Benson (Blyth, 1846, p. 367; and 1869 by M. Boucourt; mnhn, 2 (skins only, J. Anderson, 1881, p. 65); skin and skull for- skulls inside). Collected in 1 906 by C. Bruegel; merly preserved in zsi (not seen). Reported pres- zsbs, 4 (including 1 skin only, 2 skulls only). ent ca. 1856 by A. R. Wallace (1869, p. 326). Collected date unknown by Dr. Bulkley; amnh, Collected in 1 890 by B. Hagen; NHMBa, 1 (skull 1 (skin only). Subspecies uncertain; not mapped. only). FAS; GLS-25 through LS-29.

176 FIELDIANA: ZOOLOGY Tinggi, Pulau, MALAYSIA: WEST MALAYSIA Trusan Kinabatangan; Borneo, MALAYSIA: SA- 2°17'-2°19'N, 104°06'-104°08'E; collected 24- BAH; ca. 5°49'N, 1 18°20'E; observed May-Jun. 25 Jun. 1908 by H. C. Robinson (in Thomas & 1950 by D.D.Davis (1962, pp. 57, 127, pi. 21). Wroughton, 1909b, p. 103; cf. H. C. Robinson, FAS;C:Sab-19. 1919, p. 325) and E. Seimund; bm(nh), 3; zrc, Tuangku, Pulau, INDONESIA; 2°02'-2°14'N, 1. Collected 17 Jun. 1915 by V. Knight; zrc, 1. 97°07'-97°22'E; blood samples taken Nov.-Dec. FAS; B:SCS-6. 1986 by J. R. de Ruiter (1993, p. 91). FAS; Tingilan Road. See Tuaran-Kampong Tenghilan B:IO-5. Road. Tuangku, Pulau, north coast; INDONESIA; ca. Tinjil, Pulau, INDONESIA; 6°57'-6°58'S, 105°46'- 2°13'N, 97°14'E; collected 24-28 Jan. 1902 by 105°49'E; introduced Feb. 1988-Jun. 1990(Kyes W. L. Abbott (field catalog; cf. Miller, 1903a, p.

et al., 1991, p. 114; 1993, p. 78). Subspecies 438); usnm, 5 (including 2 skulls only; external uncertain; not mapped. measurement recorded on tag of one of these Tinonkok, Kampong Kiau region, Mount Kina- skulls). FAS; B:IO-5. balu, ca. 2200 ft (= 670 m); Borneo, MALAY- Tuan Keramat, Tanjong. See Tanjong Tuan, Ker- SIA: SABAH; ca. 6°02'N, 116°31'E; collected amat. 23 Aug. 1937 by J. A. Griswold, Jr. (1939b, p. Tuanku, Pulau. See Tuangku, Pulau. 514); mcz, 1. FAS; C:Sab-7. Tuanku, Pulo. See Tuangku, Pulau. Tioman, Pulau, MALAYSIA: WEST MALAY- Tuaran, near Kota Kinabalu, near sea level; Bor- SIA; 2°43'-2°53'N, 104°07'-104°13'E; collected neo, MALAYSIA: SABAH; 6°11'N, 116°14'E; 4 Oct. 1899 by W. L. Abbott (see Miller, 1900, collected 24 Jul. 1937 by J. A. Griswold, Jr. (see pp. 203, 246; Riley, 1938, p. 14); usnm, 1. Col- Coolidge, 1 940, p. 1 23); mcz, 2 (male skin mis- lected 20 Jun. 1 9 1 6 by C. B. Kloss; bm(nh) (skin)/ matched with female skull). FAS; C:Sab-5. zrc (skull). Examined for malaria Sep. 1 96 1 and Tuaran-Kampong Tenghilan Road, new bridge, Apr. 1962 by McW. Warren (1966, p. 156). FAS; near sea level; Borneo, MALAYSIA: SABAH; B:SCS-3. ca. 6°08'N, 116°15'E; collected 29 Jul.-8 Aug. Tjarmara. See Camara. 1937 by J. A. Griswold, Jr. (see Coolidge, 1940, Tjandiroto. See Candiroto. p. 123); mcz, 4. FAS; C:Sab-5. Tjempaga. See Parit. Tulungagung; Java, INDONESIA; 8°04'S, Tjerimai. See Ciremay, Gunung. 1 1 1°54'E; collected ca. 1 895 by E. Dubois; rmnh, Tjeringin, near Banjar, Preanger region, east; Java, 6 (skulls only). FAS; C:J-32. INDONESIA; ca. 7°22'S, 108°32'E; collected Turn Chompol. See Tham Chomphon. date unknown by H. J. V. Sody; rmnh, 4 (in- Tunggal, Bukit, 10 m; MALAYSIA: WEST MA- cluding 1 skull only). FAS; B:J-3. LAYSIA; 3°32'N, 10 1°27'E; observed Jul. 1978- Tjihara. See Cihara; Pelabuhanratu, Teluk, Ban- Jun. 1981 by C. W. Marsh and W. L. Wilson tam region. (1981, p. 232). FAS; B:WM-18. Tjikaran, Walde. See Cikarang forest. Tyching; THAILAND; 7°33'N, 99°35'E; collected Tjikoedjang. See Cikujang. 19 May 1896 by W. L. Abbott (see Riley, 1938, Tji-Tandoei River. See Kalipucang, Ci Tanduy. p. 12); usnm, 2. FAS; A:T-61. Tji Wangie. See Ciwangi. Ubud; Pulau Bali, Lesser Sunda Islands, INDO- Tk. Nipah. See Nipah, Telok, vicinity. NESIA; 8°30'S, 115°16'E; observed 21 Sep.-3 Toeroek Tjahoe. See Purukcahu, Sungai Barito. Oct. 1980 by N. Koyama, A. Asuan, and N.

Trang Bom. See Xa Trang Bom. Natsir (1981, p. 1). Observed in 1986, 1990, Trima, Teluk; Pulau Bali, Lesser Sunda Islands, 1991, and 1992 by B. P. Wheatley and D. K. INDONESIA; 8°08'S, 1 14°32'E; blood samples Harya Putra (1994, p. 246). FAS; CLS-3. taken in 1980 by Y. Kawamoto, K. Nozawa, Udon Thani, S; THAILAND; ca. 17°10'N, and Tb. M. Ischak (1981, p. 16). FAS; GLS-1. 102°50'E; unconfirmed report, source unspeci- Trinkat Island, Nicobar Islands, INDIA; 8°02'- fied, cited by Varavudhi et al. (1992, p. 338; cf. 8°08'N, 93°34'-93°37'E; primates reported ab- Fooden, 1971, p. 28; 1982, p. 576; Aggimar- sent before 1903 by G. S. Miller, Jr. (1902b, p. angsee, 1992, p. 119). Not mapped. 792; cf. Kloss, 1903a, p. 114). A:l. Udjung Kulon National Park. See Ujungkulon, Trinkut Island. See Trinkat Island. Suaka Margasatwa. Trong. See Ban Phra Muang; Tyching. Ujungkulon, Suaka Margasatwa; Java, INDO-

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 177 NESIA; 6°39'-6°53'S, 105°12'-105°30'E; ob- blood samples taken Aug.-Sep. 1988 by Y. Ka- served Sep. 1940-Jul. 1955 by A. Hoogerwerf wamoto, T. Ishida, J. Suzuki, O. Takenaka, and (1970, p. 408). Reported present ca. 1961 by R. P. Varavudhi (1989, p. 95). FAS; A:T-32. K. P. Satmoko (1961, p. 117). Observed Sep. Wat (= Temple) Ban Kan Yai; THAILAND; 1969-Jan. 1971) by W. Angst (1973, p. 626; 15°21'N, 1 04° 13'E; observed 14 Jul. 1989 by N. 1975, p. 326). Observed summer 1982 by G. Aggimarangsee(1992,pp. 109, 122;pers. comm., Hohmann and W. P. Peter (1983, p. 44). FAS; Oct. 1993). Subspecies uncertain; A:T-17. B:J-14. Wat Ban Muang Khan. See Wat Ban Kan Yai. Ulu, Sungai; Pulau Natuna Besar, INDONESIA; Wat Ban Rai Don; THAILAND; 1 3°07'N, 99°50'E; 3°53'N, 108°24'E; collected 25 Apr. 1909 by V. reported present 20 Aug. 1 989 by local nun (Ag- Knight (see Chasen, 1935a, p. 5); zrc, 1 (skull gimarangsee, 1992, pp. 110, 132; pers. comm., not examined). FAS; B:SCS-32. Oct. 1993). Subspecies uncertain; A:T-37. Forest Ulu Gombak Reserve; MALAYSIA: WEST Wat Bun Thawi; THAILAND; 13°08'N, 99°56'E; ca. observed MALAYSIA; 3°18'N, 101°47'E; observed 18 Aug. 1989 and 1 1 Feb. 1991 by N. 1962 R. H. D. E. McW. Aug. by Wharton, Eyles, Aggimarangsee (1992, pp. 109, 112, 131; pers. H. Warren, and W. Cheong (1964, p. 59). FAS; comm., Oct. 1993). Subspecies uncertain; A:T- B:WM-19. 37. Ulu Ijok, 400 ft (= 120 m); MALAYSIA: WEST Wat Cha-Am Kiri; THAILAND; 13°48'N, 5°08'N, collected 27- MALAYSIA; 100°48'E; 99°53'E; reported by local residents as exter- 30 Dec. 1930 by A. B. Holloway and A. S. Ver- minated ca. 1 980 (Aggimarangsee, 1992, pp. 110, (see P. H. Napier, 1981, p. 14); bm(nh), 2. nay 134; pers. comm., Oct. 1993). FAS; A:T-28. FAS; B:WM-5. Wat Huai Takhaeng; THAILAND; 13°35'N, Ulu Sebol. See Sekol, Sungai. 99°46'E; blood samples taken Aug.-Sep. 1988 Ulu Segama Reserve; Borneo, MALAYSIA: SA- by Y. Kawamoto, T. Ishida, J. Suzuki, O. Tak- BAH; ca. 5°10'N, 1 17°54'E; observed Jun.-Oct. enaka, and P. Varavudhi (1989, p. 95). Ob- 1968 and Oct. 1969-Oct. 1970 by J. R. served 20 Dec. 1 990 by N. Aggimarangsee (1 992, MacKinnon (1971, p. 153). FAS; C:Sab-13. pp. Ill, 128; pers. comm., Oct. 1993). FAS; Um See Nam Mae. Pang. Wong, A:T-28. Ungar, Pulau, Kepulauan Riau, INDONESIA; Wat Khao Bandai It; THAILAND; 13°06'N, 0°37'-0°42'N, 103°28'-103°32'E; reported pres- 99°56'E; observed 10-1 1 Aug. 1989 and 1 1 Feb. ent Jun.-Aug. 1 903 by W. L. Abbott (see Miller, 1991 byN. Aggimarangsee (1992, pp. 109, 112, 1906c, p. 279). FAS; B:SCS-16. 131; pers. comm., Oct. 1993). Subspecies un- Unter Langkat district; Sumatra, INDONESIA; certain; A:T-37. ca. 3°55'N, 98°20'E; collected 1897-1899 by G. Wat Khao Chong Phran; THAILAND; 13°43'N, Schneider (1905, p. 72); museum and number 99°46'E; observed 20 Dec. 1990 N. Aggi- of specimens unknown (not seen). FAS; B:S-20. by Uwi, Pulau, INDONESIA; 1°04'-1°07'N, 107°22'- marangsee (1992, pp. Ill, 127; pers. comm., Oct. A:T-28. 107°25'E; collected 13 Aug. 1899 by W. L. Ab- 1993). FAS; Wat Khao Khang. See Wat bott (in Miller, 1900, p. 244; cf. Kloss, 1903b, Ngern Rung Sawang. Wat Khao Khan p. 65; Oberholser, 1919, p. 129); usnm, 1. FAS; Hok; THAILAND; 13°50'N, B:SCS-26. 99°38'E; observed 22 Dec. 1990 by N. Aggi- Vadju, Nusa, Komodo area, Lesser Sunda Islands, marangsee (1992, pp. Ill, 126; pers. comm., Oct. A:T-28. INDONESIA; not located, ca. 8°30'S, 1 19°30'E; 1993). FAS; monkeys reported absent 1 969-1973 by W. Auf- Wat Khao Noh; THAILAND; 15°57'N, 99°53'E; fenberg (1981, p. 40). Not mapped. observed 10 Mar. 1967 by J. Fooden (1971, p. Ven-Ven (?= Mae Nam Welu); THAILAND; ca. 17). Blood samples taken Aug.-Sep. 1988 by Y. 12°20'N, 102°20'E; observed 1-4 Mar. 1859 by Kawamoto, T. Ishida, J. Suzuki, O. Takenaka,

H. Mouhot(1864, vol. 1, p. 152). FAS; A:T-34. and P. Varavudhi (1989, p. 95). Reported pres- Verlaten Island. See Sertung, Pulau. ent 22 Feb. 1991 by N. Aggimarangsee (1992, Victoria Island. See Ru, Pulo. pp. 112, 123; pers. comm., Oct. 1993). Subspe- Wai, Pulo. See Uwi, Pulau. cies uncertain; A:T-4. Wai Sano. See Sano, Wai. Wat Khao Phlu. See Wat Tham Khao Phlu. Walde Tjikaran. See Cikarang forest. Wat Khao Sompoad (?= Som Phot); THAILAND; Wang Kaew; THAILAND; ca. 12°47'N, 101°39'E; ca. 15°10'N, 101°17'E; observed 11 Jan. 1991

178 FIELDIANA: ZOOLOGY by N. Aggimarangsee ( 1 992, pp. Ill, 1 23). Sub- 100°09'E; reported present 7 Jul. 1989 by local species uncertain; A:T-8. monk (Aggimarangsee, 1992, pp. 109, 125; pers. Wat Khao Takhrao; THAILAND; 13°13'N, comm., Oct. 1993). Subspecies uncertain; A:T- 99°56'E; observed 21 Aug. 1989 and 11 Feb. 23. 1991 by N. Aggimarangsee (1992, pp. 1 10, 1 12, Wat Phra Buddha Chai; THAILAND; 14°27'N, 132; pers. comm., Oct. 1993). Subspecies un- 100°57'E; observed 11 Jan. 1991 by N. Aggi- certain; A:T-37. marangsee (1992, pp. Ill, 125; pers. comm., Wat Khao Takieb; THAILAND; 1 2°3 1 'N, 99°59'E; Oct. 1993). Subspecies uncertain; A:T-24. observed 16-17 Sep. 1 990 by N. Aggimarangsee Wat Ratch Singkhorn; THAILAND; 13°34'N, (1992, pp. 110, 134; pers. comm., Oct. 1993). 99°47'E; blood samples taken Aug.-Sep. 1988 Subspecies uncertain; A:T-38. by Y. Kawamoto, T. Ishida, J. Suzuki, O. Tak- Wat Khao Tamon; THAILAND; 13°07'N, enaka, and P. Varavudhi (1989, p. 95). Ob- 99°56'E; reported present 4 Feb. 1991 by N. served 24 Dec. 1 990 by N. Aggimarangsee (1992, Aggimarangsee (1992, pp. 112, 132; pers. comm., pp. Ill, 129; pers. comm., Oct. 1993). FAS; Oct. 1993). Subspecies uncertain; A:T-37. A:T-28. Wat Khao Wong Kot; THAILAND; 15°01'N, Wat Suwankuha (cf. Wat Khuha Sawan); THAI- 100°33'E; reported present 24 Jun. 1989 by un- LAND; ca. 7°37'N, 100°05'E; blood samples specified informants (Aggimarangsee, 1992, pp. taken Aug.-Sep. 1988 by Y. Kawamoto, T. Ish- 109, 115, 124; pers. comm., Oct. 1993). Sub- ida, J. Suzuki, O. Takenaka, and P. Varavudhi species uncertain; A:T-6. (1989, p. 95). FAS; A:T-65. Wat Khao Yod Thong; THAILAND; 13°28'N, Wat Tha Mai Lai; THAILAND; 10°30'N, 98°58'E; 99°43'E; reported present 25 Dec. 1990 by N. observed 18 Sep. 1990 by N. Aggimarangsee Aggimarangsee (1992, pp. Ill, 127; pers. comm., (1992, pp. 110, 137; pers. comm., Oct. 1993). Oct. 1993). FAS; A:T-28. AUR/FAS/MUL; A:T-43. Wat Khuha Phimuk; THAILAND; ca. 6°31'N, Wat Tham Khao Phlu; THAILAND; 10°44'N, 101°14'E; observed 21-22 Sep. 1990 by N. Ag- 99°19'E; reported present 1-2 Dec. 1990 by lo- gimarangsee (1992, pp. 110, 139; pers. comm., cal residents (Aggimarangsee, 1 992, pp. Ill, 137; Oct. 1993). FAS; A:T-69. pers. comm., Oct. 1993). AUR/FAS/MUL; A:T- Wat Khuha Sawan; THAILAND; 7°37'N, 42. 100°05'E; observed 20-21 Sep. 1990 and 26-28 Wat Tham Kunchhorn; THAILAND; 13°29'N, Feb. 1991 by N. Aggimarangsee (1992, pp. 1 10, 99°42'E; observed 19 Dec. 1990 by N. Aggi- 112, 138; pers. comm., Oct. 1993). FAS; A:T- marangsee (1992, pp. Ill, 126; pers. comm., 65. Oct. 1993). FAS; A:T-28. Wat Koo Pra Kona; THAILAND; 15°33'N, Wat Thammikaram Varaviharn; THAILAND; 103°49'E; blood samples taken Aug.-Sep. 1988 1 1°49'N, 99°48'E; observed 17 Sep. 1990 by N. by Y. Kawamoto, T. Ishida, J. Suzuki, O. Tak- Aggimarangsee (1992, pp. 1 10, 134; pers. comm., enaka, and P. Varavudhi (1989, p. 95). Ob- Oct. 1993). AUR/FAS/MUL; A:T-41. served 14-16 Jul. 1989 and 17-18 Jan. 1991 by Wat Tham Sala; THAILAND; 13°49'N, 100°07'E; N. Aggimarangsee (1992, pp. 109, 112, 120; pers. observed 12 May 1989 and 15-19 Dec. 1990 comm., Oct. 1993). Subspecies uncertain; A:T- by N. Aggimarangsee (1992, pp. 109, 1 1 1, 125; 13. pers. comm., Oct. 1993). FAS; A:T-29. Wat Krieng Krai Klang; THAILAND; 15°44'N, Wat Tham Sua; THAILAND; 8°07'N, 98°55'E; 100°11'E; observed 22 Feb. 1991 by N. Aggi- observed 1 9-20 Sep. 1 990 by N. Aggimarangsee marangsee (1992, pp. 112, 123; pers. comm., (1992, pp. 110, 138; pers. comm., Oct. 1993). Oct. 1993). Subspecies uncertain; A:T-5. FAS; A:T-60. Wat Kut; THAILAND; 13°09'N, 99°57'E; ob- Wat Tham Suwan Khuha; THAILAND; 8°26'N, served 1 9 Aug. 1 989 by N. Aggimarangsee (1992, 98°28'E; observed 17-21 Nov. 1987 by R. pp. 110, 131; pers. comm., Oct. 1993). Subspe- Boonratana (1988, p. 75). Reported present 19 cies uncertain; A:T-37. Sep. 1990 by local residents (Aggimarangsee, Wat Ngern Rung Sawang; THAILAND; 13°42'N, 1992, pp. 110, 137; pers. comm., Oct. 1993). 99°46'E; observed 20 Dec. 1990 by N. Aggi- Subspecies uncertain; A:T-51. marangsee (1992, pp. Ill, 128; pers. comm., Waw, Telok. See Telok Wau. Oct. 1993). FAS; A:T-28. Wawo. See Ntori, Sape district. Wat Noi Chompoo; THAILAND; 14°36'N, Way Kambas. See Kambas, Wai.

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 179 We, Pulau, INDONESIA; 5°47'-5°53'N, 95°13'- Wonosobo, Propinsi Lampung; Sumatra, IN- 95°23'E; reported present before 1981 by J. T. DONESIA; 5°30'S, 104°30'E; collected 28 Dec.

Marshall (Crockett & Wilson, 1980, p. 156). 1924 by H. J. V. Sody (see Hooijer, 1962b, p. Blood samples taken Nov.-Dec. 1986 by J. R. 44); rmnh, 1 (skull only). FAS; B:S-79. de Ruiter (1993, p. 91). Subspecies uncertain Wynkoops Bay. See Pelabuhanratu, Teluk, Ban- (see Scheffrahn et al., 1994, p. 136); B:IO-l. tam region. Weh, Pulau. See We, Pulau. Xa Trang Bom; VIETNAM; 10°57'N, 107°01'E; Weld's Hill. See Nanas, Bukit, Kuala Lumpur. collected 1 Jun. 1918 by C. B. Kloss (1919b, p. . See Bali Barat National 401; cf. Weitzel et al., 1988, p. 146); zrc, 1. Park. FAS; A:V-3. West Bali Wildlife Reserve. See Bali Barat Na- YaoNoi, Ko, THAILAND; 8°05'-8°l l'N, 98°34'- tional Park. 98°38'E; collected 30 Jan. 1918 by local collec- Whykeong Union Council. See Bilasodia; Bimir- tors employed by H. C. Robinson and C. B. dia; Ghorardia; Ochodia; Rukumodia. Kloss (1919, p. 87); zrc, 1. AUR/FAS/MUL; Wimpong, limestone rocks; BURMA; ca. 16°53'N, A:T-52. 97°28'E; collected ca. 1877-1878 by W. Davison Yaring, tidal creeks near; THAILAND; ca. 6°52'N, (see Hume & Davison, 1878, p. 524; Thomas, 101°22'E; observed Jun. 1901 by N. Annandale 1886, pp. 65, 66); bm(nh), 1. AUR; A:Bu-9. and H. C. Robinson (1903, p. xxxix; in Bonhote, Wong, Nam Mae, 40 mi (= 65 km) E of Um Pang, 1903, p. 4). FAS; A:T-68. 1000 ft (= 300 m); THAILAND; ca. 15°55'N, Yaring region; THAILAND; ca. 6°52'N, 101°22'E; 99°10'E; collected 9 Feb. 1924 by A. S. Vernay collected 19 Jun. 1899 by R. Evans and F. F. (see Lowe, 1932, p. 197; 1933, p. 260); amnh, Laidlaw (see P. H. Napier, 1981, p. 17; cf. Bon- 1;bm(nh), 1. AUR; A:T-18. hote, [1901], p. 870); University Museum of Wong, Nam Mae, 53 mi (= 85 km) E of Um Pang, Zoology, Cambridge, 1 (skull only, not seen). 800 ft (245 m); THAILAND; ca. 15°55'N, FAS; A:T-68. 99°25'E; collected 18 Feb. 1924 by A. S. Vernay Ye Forest, Ataran district, Moulmein region; (see Lowe, 1932, p. 197; 1933, p. 260); amnh, BURMA; ca. 16°10'N, 98°00'E; collected Nov. 1. AUR; A:T-3. 1 9 1 by G. F. B. R. Thurling (see Pocock, 1939, Wonokerto. See Wonokojo (?= Wonokerto), p. 81); bm(nh), 1. AUR; A:Bu-l 1. Dampit district, southern Malang region. Zadetkyi Kyun, BURMA; 9°50'-10°03'N, 98°07'- Wonokojo (?= Wonokerto), Dampit district, 98°18'E; collected 9 Dec. 1900 by W. L. Abbott; southern Malang region; Java, INDONESIA; ca. usnm, 1. AUR; A:Bu-24. 8°13'S, 112°35'E; collected Apr. 1930 by H. J. V. Sody; rmnh, 3. FAS; C:J-34.

180 FIELDIANA: ZOOLOGY Appendix 3: Dorsal Pelage Color Saturation in Fringing-Island Samples of Macaco, fascicularis Compared with Saturation in Core-Area Reference Samples

Core-area Fringing-island samples reference Frequencies at SI values samples FI> CA> Island 2 N 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 Mean Mean N CA3 FI4 P*

Shallow-water islands West of Sumatra Appendix 3: Continued

Core-area Fringing-island samples reference Frequencies at SI values samples FI > CA > Island 2 N 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 Mean Mean N CA3 FI 4

South of Indochinese Peninsula Khram Yai Appendix 3: Continued

Core-area Fringing-island samples reference Frequencies at SI values samples FI> CA> Island 2 N 0.5 1.0 1.5 2.0 2.5 3.0 3.5 4.0 Mean Mean N CA3 FI4

Luzon Appendix 4: Dorsal Pelage Color Erythrism in Fringing-Island Samples of Macaca fascicularis Compared with Erythrism in Core-Area Reference Samples Appendix 4: Continued Appendix 4: Continued

Fringing-island samples Core-area Frequencies at erythrism reference index values samples FI > CA > Island 2 N 1.0 1.5 2.0 2.5 3.0 Mean Mean N CA3 FI4 Mindanao Appendix 5: Crown Color Pattern Frequency in Samples of Macaco, fascicularis That 1 Include Dark-Crowned Specimens (cf. Table 5) Appendix 6: Geographic Variation of Lateral Facial Crest Pattern in Samples of Macaco, fascicularis (Including Infants) Collected or Observed in the Indochinese Peninsula, Isthmus of Kra, Malay Peninsula, Neighboring Shallow-Water Fringing Islands, Nic- obar Islands, P. Simeulue, and P. Lasia (cf. Table 6; Figs. 8, 9)

A. Samples Homogeneous for Lateral Facial Crest Pattern

1 . Indochinese Peninsula, Isthmus of Kra, Malay Peninsula = a. Area homogeneous for the infrazygomatic pattern, n 41. BANGLADESH, Whykeong (21°05'N,

90°12'E), 1 (cf. M. A. R. Khan, 1985, p. 29). BURMA, ArakanDiv.(19°27'N, 93°31'E), l;Wimpong (16°53'N, 97°28'E), 1; Haungtharaw (16°30'N, 98°13'E), 4; Ye Forest (16°10'N, 98°00'E), 1; Tavoy River (14°02'N, 98°12'E), 2; Taungbyauk (13°45'N, 98°26'E), 4; Mergui (12°26'N, 98°36'E), 2; Tagoot (12°15'N, 99°03'E), 2; Tenasserim (12°05'N, 99°01'E), 11; Ban Sadein (10°20'N, 98°32'E), 1. THAILAND, Wong, Nam Mae, 65 km E of Urn Pang (15°55'N, 99°10'E), 2; Wong, Nam Mae,

85 km E of Urn Pang(15°55'N, 99°25'E), 1; BanTamrongPhato(14°54'N, 98°31'E), 7; COUNTRY UNCERTAIN, "Bengale" (coordinates unknown), 1. = b. Area homogeneous for the transzygomatic pattern, n 66 (excludes 2 specimens with transzy- gomatic crests that were collected at unspecified localities in Thailand). THAILAND, Pak Nam Pho (15°43'N, 100°12'E), 3; Kata Taek (15°28'N, 99°23'E), 6; Lat Bua Khao (14°52'N, 101°36'E), 2; Ban Phu Toie (14°42'N, 99°07'E), 2; Ban Huai Maenam Noi (14°25'N, 98°5 l'E), 2; Aranyaprathet

(13°41'N, 102°30'E), 1; Laem Sing Mountains (12°29'N, 102°06'E), 1; Ban Thap Plik (8°11'N,

1 1 1 98°53'E), 2; Tham Horn (8°1 'N, 98°53'E), ; Ban Nong Kok (8°06'N, 98°52'E), ; Tyching (7°33'N,

99°35'E), 2; Kantang (7°25'N, 99°30'E), 1; Khao Rang Kai (7°19'N, 99°48'E), 1; Ban Phra Muang (7°21'N, 99°28'E), 2; Ban Sai Kau (6°38'N, 101°08'E), 1; Kampong Biserat (6°32'N, 101°14'E), 3. VIETNAM, Sontra Peak (16°07'N, 108°21'E), 1; Tay Ninh (1 1°18'N, 106°06'E), 1; Xa Trang Bom

(10°57'N, 107°00'E), 1; Ho Chi Minh City (10°45'N, 106°42'E), 2; Cochin China (coordinates uncertain), 1. VIETNAM OR CAMBODIA, no locality, 2. WEST MALAYSIA, 16 localities, 27. 2. Neighboring shallow-water fringing islands = a. Island group homogeneous for the infrazygomatic pattern, n 10. BURMA, Mergui Archipelago, Kathema(13°39'N, 98°12'E), 4; Mibya(13°36'N, 98°12'E), 2; Kadan(12°30'N, 98°22'E), 1; Letsok-

aw (11°37'N, 98°15'E), 1; Lanbi (10°50'N, 98°15'E), 1; Zadetkyi (9°58'N, 98°13'E), 1. = b. Islands homogeneous for the transzygomatic pattern, n 102. THAILAND, Khram Yai (12°42'N,

100°47'E), 1 1; Chang (12°00'N, 102°23'E), 3; Kut(l 1°40'N, 102°35'E), 8; Phangan(9°45'N, 100°00'E), 2; Samui (9°30'N, 100°00'E), 2; Phi Phi Don (7°45'N, 98°47'E), 4; Talibong (7°15'N, 99°29'E), 2; Tarutao (6°35'N, 99°40'E), 6; Butang (6°32'N, 99°12'E), 1. VIETNAM, Phu Quoc (10°13'N,

103°57'E), 1; Con Son (8°43'N, 106°35'E), 9; Ba (8°39'N, 106°33'E), 3. WEST MALAYSIA, 11 islands, 39. SINGAPORE (1°23'N, 103°50'E), 11.

FIELDIANA: ZOOLOGY B. Samples Heterogeneous for Lateral Facial Crest Pattern Appendix 7: Head and Body Length (mm) in Fringing-Island Samples of Macacafas- cicularis Compared with Head and Body Length in Core-Area Reference Samples 1 Collected at Similar Latitudes (cf. Table 9; Figs. 11, 12)

Fringing-island Appendix 7: Continued

Fringing-island samples Appendix 7: Continued Appendix 8: Tail Length (mm) in I ringing- Island Samples of Macaca fascicularis Compared with Tail Length in Core-Area Reference Samples Collected at Similar 1 Latitudes (cf. Table 11; Figs. 15, 16)

Fringing-islands samples Appendix 8: Continued Appendix 8: Continued

Fringing- Appendix 9: Relative Tail Length (T/HB x 100) in Fringing-Island Samples of Macaca fascicularis Compared with Relative Tail Length in Core-Area Reference Samples Col- 1 lected at Similar Latitudes ; Samples Include Both Sexes (cf. Table 11; Figs. 17, 18) Appendix 9: Continued Appendix 10: Blood-Protein Allele Frequencies at Polymorphic Loci in 11 Geographic Samples of Macaca fascicularis Appendix 10: Continued t* r- —

+

J8 + I + II +

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1 Arrows indicate that data were reported for a multimonth period. 2 Key to references: 1, M. A. R. Khan & Wahab, 1983, p. 104. 2, Fooden, 1971, p. 26. 3, Fooden, [1975], p. 99. 4, Kavanagh & Laursen, 1984, pp. 24-25. 5, Aldrich-Blake, 1980, p. 151. 6, F. N. Chasen, zrc 4-089, specimen tag. 7, Crockett & Wilson, 1980, pp. 172-173. 8, van Noordwijk, 1985, pp. 280-281. 9, van Schaik & van Noordwijk, 1985b, p. 539. 10, Karssemeijer et al., 1990, p. 276. 11, Moszkowski, 1909, p. 6. 12, Koyama et al., 1981, pp. 2, 3, 8. 13, Norikoshi, 1984, p. 13. 14, Koyama, 1985, p. 113. 15, Koyama, 1984, p. 35. 16, Zuckerman, [1933], p. 1063. 17, Davis, 1962, p. 58. 75, Kurland, 1973, p. 258. 19, Wheatley, 1982, pp. 210-211. 20, Berenstain, 1986, p. 258. 21, O. Bryant, Coll. No. 1049, usnm field catalog. 22, Hoogerwerf, 1970, p. 408. 23, Wheatley, 1991, p. 171. 24, Fooden, 1991, pp. 18, 20. 3 Multimonth reporting period extends to February 1983. 4 Figures indicate mean daily copulation vocalizations per month. 5 Copulations study period January-March 1981. 6 Copulations study period March-August 1982. 7 Figures indicate consortships per month. 8 Copulations study period January-February 1983. 9 Includes one birth recorded in January 1984. 10 Provisioned troops. " Multimonth reporting period extends to 5 January 1984.

1 2 Possibly includes data on a few specimens that originated in Java. 13 Twenty-nine percent observed in April 1976. 14 In a brief preliminary report, Angst (1975, p. 328) suggested that consortships occur throughout the year in Java and Bali, that copulations peak during the summer, and that births may peak in December-May.

202 FIELDIANA: ZOOLOGY Appendix 12: Subspecific Variation of Head and Body Length, Relative Tail Length, and Greatest Length of Skull in Macaca fascicularis (cf. Tables 8, 10, 15; Appendixes 7, 9; Fooden & Albrecht, 1993, p. 537) Subject Index

abbreviations, institutional names 1 fertility 61 paternity tests 6 1 abundance 1 fossils 63 pelage 3 Allen's rule 23 fruits, ingestion of 54 crown color pattern 1 5, 83, 86, 95 allometric growth 30 G-banding pattern 47 erythrism:

drift 45 variation 1 1 alpha-globin genes 39 genetic geographic 3, 90, 9 , altitudinal range 49 geographic names 2, 128 95 arboreal ity 50 gestation length 6 1 standards 13 Bergmann's rule 23, 33 glacial maxima 3, 87, 97 hair banding 3 birth rate, annual 6 1 habitats 49 infantile 1 1 birth weight 22, 61 hair banding 3 ontogenetic changes 1 1 blood groups 45 head and body length. See under ex- saturation:

ternal 1 1 blood proteins 42, 91, 96, 97 measurements geographic variation , 68, 69, cholesterol response 46 history of scientific knowledge 1 90, 91, 95, 97 chromosomes 47 home range 5 1 standards 1 1 classifications, recent 69 human-type blood groups 45 sexual dimorphism 1 1 collections studied 1 inbreeding avoidance 57 tail coloration 5, 67, 95 consortship 60 Indochinese faunal subregion 94 permanent teeth, emergence 33 constancy of troop size 5 1 Indomalayan faunal subregion 94 Pleistocene 3, 63, 87 contact zones infanticide 63 polio virus, susceptibility to 49 interspecific 68, 77, 79, 92 infantile pelage 1 1 population intersubspecific 67, 68, 69, 77, 79, infant mortality 63 density 53 92,95 infant sex ratio 6 1 estimates 54 copulatory behavior 59 insular dwarfism 23, 34 growth rate 63, 88 core area of distribution 3 intergradation 92 predators 54 cranial measurements 30 interspecific behavior 57 predators, fear of 55 geographic variation 33, 91, 96, interspecific contact zone 68, 77, 79, prey 54 97 92 rafting 87, 96 ontogeny 30 intersubspecific contact zone 67, 68, range sexual dimorphism 30, 8 1 69, 77, 79, 92, 95 day 53 crest. See lateral facial crest intertroop behavior 56 home 51 crop-raiding 54 intertroop movement 57 reproduction 58 crown color pattern 15, 83, 86, 95 introduction by humans 2, 65, 88, reproductive success 6 1 current speed, ocean 87 96,97 resistance to infection 48, 9 1 day range 53 introgression 93, 95 restriction patterns, DNA 35, 39 deciduous teeth, emergence 32 invertebrate prey 54 retrovirus, simian T-lymphotrophic deep-water fringing islands 3 karyology 47 49 dental emergence 32 key to subspecies 69 saturation, pelage color. See under diet lateral facial crest 1 1 54 8, 67, 89, 90, 9 , pelage dimorphism, sexual 11, 21, 30, 81 95,96 sea-level variation 3, 87 diseases 47 malaria 47 seasonal reproductive peaks 58 dispersal hypotheses 88 mammal fauna seeds, ingestion 54 dispersal, overwater 87 Flores 96 serum cholesterol response 46 distribution, geographic 2, 64 Maratua 90 sex ratio, in troops 5 1 dominance, relationship to repro- Nicobars 89 sexual dimorphism 11, 21, 30, 81 ductive success 60 Nias 95 sexual maturation drownings 5 1 Simeulue-Lasia group 90 females 58 dwarfism, insular 23, 34 mangrove forest 50 males 59 ejaculation menarche 58 sexual swelling 59 multiple-mount 60 menstrual cycle 59 shallow-water fringing islands 3 single-mount 60 mitochondrial DNA 35, 91 sidewhiskers. See lateral facial crest erythrism. See under pelage mobbing of predators 55 simian T-lymphotrophic retrovirus evolutionary hypotheses 88 molecular biology 35 49 external measurements 21 mortality, infant 63 simian-type blood groups 45 appendages 2 1 mosquito vectors 48 single-mount ejaculation 60 geographic variation 22 multiple-mount ejaculation 60 sleeping trees 5 1 head and body length, 22, 91, museums 1 subspecific recognition 65 96,97 natural history 49 subspecific taxonomy 65 relative tail length 24, 67, 82 nuclear DNA 39 Sunda Shelf 3 tail length 23, 67, 82, 91, 95 nursing of infants 6 1 suppressed name 70 ontogeny 2 1 overlap, home range 5 1 surface current speed 87 sexual dimorphism 2 1 parasites, malaria 47 swimming 51, 87

204 FIELDIANA: ZOOLOGY tail coloration 5, 67, 95 terrestriality 50 water gaps, effects on distribution tail length. See under external mea- time budgets 57 87 surements troop size and composition 5 1 weight taxonomy 65 vectors, malaria infection 48 adult 21 tectonic forces, effects on distribu- vernacular names 1 neonatal 22, 6 1 tion 87 vertebrate prey 54

FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 205 Index to Scientific Names

For macaque taxa, primary alphabetization is according to subspecific or specific name; for non- macaque taxa, primary alphabetization is according to generic name.name adusta, Macaca [nemestrinus- Cynosurus, Cercopithecus 77 Opuntia elatior 54 Gruppe] aff. 75 Dicrurus paradiseus 58 Panthera pardus 55 aqnatus, Pithecus 73 dollmani, Pithecus 73 ft'gra 55 Aigula, Cercop[ithecus] 70 fascicularis, Macacafascicularis 66, phaeura, Macacus 67, 72 alacer, Pithecus 72 69, 70, 79, 81, 82, 91, 95 phaeurus, Pithecus 72 Anopheles balabacensis 49 fascicularis, Pithecus 78 philippinensis, Macaca fascicularis dims, sensu lato 49 fascicularis, Simia 71 69, 70, 77, 79, 90, 95 hackeri 49 /«5ca, Macaca fascicularis 69, 70, Pithecophaga jefferi 55 introlatus 49 81,82,90 Plasmodium coatneyi 47 leucosphyrus, sensu lato 49 fuscata, Macaca 45 cynomolgi 47 argentimembra, Macacafascicular- fuscus, Cynomolgus 72 jfcf&fi 47 is 75 fuscus, Macacus 81 /'««/ 47 argentimembris, Macaca irus 75 Homo erectus 88 knowlesi 47, 91 assamensis, Macacus 71 sapiens 88 Pongo pygmaeus 58 atriceps, Macacafascicularis 68, 70, Hylobates lar 58 Presbytis aygula 58 82,95 moloch 57 comata 88 auratus, Macacus 78 impudens, Pithecus 75 /lose/ 58 aurea, Macaca fascicularis 67, 69, z'ra, [Macaca] 71 melalophos 58 77, 78, 90, 91 7n« 71 See also Trachypithecus aureus, Macacus 77 /rus, Macaca 68, 71, 84 pumila, Macaca irus 72 Aygula, [Simia] 70 /rus, Macacus 7 1 pumilus, Macacus 7 1 baweana, Macaca irus 74 karimondjawae, Macaca fascicu- Python sp. 55 baweanus, Pithecus 74 laris 68, 70, 86, 95 resima, Macaca 73 bintangensis, Pithecus 73 karimoni, Pithecus 72 resimus, [Macacus] 73 Buku, Semnopithecus 7 1 fcra, Semnopithecus 7 1 Sinicus, Macacus 7 1 cagayana, Macaca 72 /aeta, Macaca irus 73 sirhassensis, Pithecus 74 cagayanus, Cynomolgus 72 /a?//, M[acaca] f[ascicularis] 73 sp., Pithecus 76 cancrivorus, Macacus 71 laetus, Pithecus 73 sublimitis, Macaca irus 76 capitalis, Pithecus 75 lapsus, Pithecus 73 sublimitus, Macaca irus 76 carbonaria, S[imia] 71 lasiae, Macaca fascicularis 69, 70, submordax, Macaca irus 76 Carbonarius 7 1 81,90 suluana, Macaca 72 carbonarius, Cercocebus 79 lautensis, Pithecus 74 suluensis, Cynomolgus 72 carbonarius, M[acacus] 78 Leucosphyrus Group 48 sulvensis, Macaca 72 carbonarius, Macacus 7 1 limitis, Pithecus fascicularis 75 sumbae, [Macaca irus] 76 carimatae, Pithecus 74 limitis, Pithecus fascicularis, aff. 75 Tomistoma schlegeli 55 cercopithecus, Inuus 71 limitis, Macaca irus 75 Trachypithecus auratus 88, 96 condorensis, Macaca fascicularis 68, lingae, Pithecus 75 auratus kohlbruggei 96 70, 83, 84, 95 lingungensis, Pithecus 74 cristatus 50, 58 Crocodylus sp. 55 mandibula, [Macaca irus] 74 obscurus 58 cupida, Macaca irus 75 mandibulars, Pithecus 74 /«a, Macaca fascicularis 69, 70, 86, cupidus, Pithecus 74 mansalaris, Pithecus 75 90 Cynamolgos, Simia 70 mauritius, Simia 76 umbrosa, Macaca fascicularis 68, (Cynomolgus), Cercopithecus 70 mordax, Macaca 73 70, 80, 89 cynocephalus, C[ynamolgus] 71 mulatto, C[ynamolgus] 78 umbrosus, Macacus 80 cynomolgos, M[acacus] 79 mulatto, Macaca 19, 25, 45, 68, 76, valida, Macaca irus 72 cynomolgos, Simia 70 77, 79, 91 validus, Pithecus 72 Cynomolgus, [Cercopithecus] 70 Nasalis larvatus 50, 58 Varanus komodoensis 55 cynomolgus, Cercopithecus 80 nemestrina, Macaca 26, 33, 57, 63, v/f/Vs, Pithecus 78 cynomolgus, M[acacus] 78 88 cynomolgus, Macacus 86 Neofelis nebulosa 55

206 FIELDIANA: ZOOLOGY xien. Fieh

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