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BIX S^NCY*"* 3 - SYSID -BL- EW SERIES Systematic Review of Southeast Asian Longtail M&cm Macaca fascicular™ (Raffles, [11 590 . 5 BIX FI N.S. 31 KJ0" Publication 1470 PUBLISHED BY F IELD i Confabs :d for puti FIELDIANA 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 Abstract 1 Introduction 1 Geographic Distribution 2 Pelage 3 General Characterization 3 Sex and Age Variation 11 Geographic Variation 11 Dorsal Pelage Color Saturation 11 Dorsal Pelage Erythrism 13 Crown Color 15 Lateral Facial Crest Pattern 18 External Measurements and Propor- tions 21 Sex and Age Variation 21 Geographic Variation 22 Head and Body Length 22 Tail Length 23 Cranial Characters 30 Sex and Age Variation 30 Geographic Variation 33 Molecular Biology and Genetics 35 Mitochondrial DNA 35 Nuclear DNA 39 Multiple Alpha-Globin Genes 39 Highly Repeated Restriction Patterns ... 41 Blood Proteins 42 Blood Groups 45 Serum Cholesterol Response 46 Karyology 47 Diseases 47 Malaria 47 Simian T-Lymphotrophic Retrovirus, Type 1 49 Susceptibility to Attenuated Polio Virus ... 49 Natural History 49 Habitats 49 Arboreality/Terrestriality 50 Swimming 51 Troop Size and Composition 51 Home Range, Day Range 51 Density 53 Diet 54 Predators 54 Intertroop Behavior 56 Interspecific Behavior 57 Reproduction 58 Fossils and Subfossils 63 Systematics 65 Subspecific Taxonomy 65 Key to Recognized Subspecies 69 Subspecies Accounts 70 Macaca fascicularis fascicularis (Raffles, [1821]) 70 Macaca fascicularis aurea I. Geoffroy, [1831] 77 Macaca fascicularis philippinensis I. Geoffroy, [1843] 79 Macaca fascicularis umbrosa Miller, 1902b 79 Macaca fascicularis fusca Miller, 1903a 81 Macaca fascicularis lasiae (Lyon, 1916) 81 Macaca fascicularis atriceps Kloss, 1919c 82 Macaca fascicularis condorensis Kloss, 1926 84 Macaca fascicularis tua Kellogg, 1944 86 Macaca fascicularis karimondjawae Sody, 1949 86 Macaca fascicularis subspecies undeter- mined 87 Evolution and Dispersal 87 Stage I, > 1 Ma 88 Stage II, ca. 160 Ka 88 Stage III, > 18 Ka 90 Stage IV, ca. 18 Ka 95 Stage V, < 18 Ka 95 Stage VI, ca. 4.5 Ka 96 Acknowledgments 97 Literature Cited 98 Appendix 1 : Specimens Examined 1 20 Appendix 2: Gazetteer of Non-Philippine Macaca fascicularis Localities 1 28 Appendix 3: Dorsal Pelage Color Satur- ation in Fringing-Island Samples of Macaca fascicularis Compared with Saturation in Core-Area Reference Samples 181 Appendix 4: Dorsal Pelage Color Erythrism in Fringing-Island Samples of Macaca fascicularis Compared with Erythrism in Core-Area Reference Samples 1 84 Appendix 5: Crown Color Pattern Frequency in Samples of Macaca fascicularis That Include Dark-Crowned Specimens 187 Appendix 6: Geographic Variation of Lateral Facial Crest Pattern in Samples of Macaca fascicularis .... 188 Appendix 7: Head and Body Length in in Fringing-Island Samples of Macaca fascicularis compared with head and Body Length in Core-Area Reference Samples Collected at Similar Latitudes 1 90 Appendix 8: Tail Length in Fringing- Island Samples of Macaca fascicu- laris Compared with Tail Length in Core-Area Reference Samples Collected at Similar Latitudes 193 Appendix 9: Relative Tail Length in Fring- ing-Island Samples of Macaca fas- cicularis Compared with Relative Tail Length in Core-Area Reference Samples Collected at Similar Latitudes 196 Appendix 10: Blood-Protein Allele Fre- quencies at Polymorphic Loci in 1 1 Geographic Samples of Macaca fascicularis 198 Appendix 1 1 : Monthly Distribution of Recorded Reproductive Events in Natural Populations of Macaca fascicularis 200 Appendix 12: Subspecific Variation of Head and Body Length, Relative Tail Length, and Greatest Length of Skull in Macaca fascicularis .... 203 Subject Index 204 Index to Scientific Names 206 List of Illustrations 1 . Known locality records of Macaca fasci- cularis 3 2. Detail maps of non-Philippine localities of Macaca fascicularis A. Northwestern quadrant 4 B. Southwestern quadrant 6 C. Southeastern quadrant 9 3. Approximate extent of dry land in South- east Asia during last glacial maximum, ca. 18 Ka 12 4. Core area, shallow-water fringing islands and deep-water fringing islands inhabited by Macaca fascicularis 13 5. External characters in Macaca fascicu- laris 14 6. Dorsal pelage color saturation extremes in Macaca fascicularis 15 7. Crown color pattern in dark-crowned Macaca fascicularis 18 8. Infrazygomatic and transzygomatic lat- eral facial crest patterns in Macaca fas- cicularis 19 9. Geographic variation of lateral facial crest pattern in samples of Macaca fascicularis collected in Indochinese Peninsula, Isth- mus of Kra, Malay Peninsula, and neigh- boring islands 21 10. Latitudinal variation of head and body length in adult core-area specimens of Macaca fascicularis 24 1 1 . Latitudinal variation of head and body length in adult shallow-water fringing-is- land specimens of Macaca fascicularis compared with that in adult core-area specimens 26 12. Latitudinal variation of head and body length in adult deep-water fringing-island specimens of Macaca fascicularis com- pared with that in adult core-area speci- mens 27 1 3. Latitudinal variation of tail length in adult core-area specimens of Macaca fascicu- laris 29 14. Latitudinal variation of relative tail length in adult core-area specimens of Macaca fascicularis 30 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 deep-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens 32 17. Latitudinal variation of relative tail length in adult shallow-water fringing-island specimens of Macaca fascicularis com- pared with that in adult core-area speci- mens 34 1 8. Latitudinal variation of relative tail length in adult deep-water fringing-island speci- mens of Macaca fascicularis compared with that in adult core-area specimens 35 1 9. Skull of adult female Macaca fascicularis, near topotype 36 20. Skull of adult male Macaca fascicularis, near topotype 37 21. Latitudinal variation of greatest length of skull in adult core-area specimens of Ma- caca fascicularis 40 22. Latitudinal variation of greatest length of skull in adult shallow-water fringing- island specimens of Macaca fascicularis compared with that in adult core-area specimens 41 23. Latitudinal variation of greatest length of skull in adult deep-water fringing-island specimens of Macaca fascicularis com- pared with that in adult core-area speci- mens 43 24. Blood-protein dendrograms for geograph- ic samples of Macaca fascicularis: Den- drogram that excludes Mauritius sample compared with dendrogram that includes Mauritius sample; consensus dendrogram 47 25. Type localities and limits of distribution of recognized subspecies of Macaca fas- cicularis and type localities of synony- mous nominal species or subspecies ... 64 26. Variation of greatest length of skull in adult Macaca fascicularis fusca (Indonesia: Pu- lau Simeulue) compared with variation in adult Sumatran M. f fascicularis 83 27. Tail length vs. head and body length in Macaca fascicularis fusca and M. f la- siae 83 28. Craniometric comparisons of adult Ma- caca fascicularis atriceps, M. f condor- ensis, and mainland Indochinese M. fas- cicularis 85 29. Latitudinal variation of adult head and body length in samples of Macaca fasci- cularis collected in the southern part of the Indochinese Peninsula and adjacent Isthmus of Kra, north of 10°N, compared with that in samples of parapatric hi. mu- latta collected in the northern part of the Indochinese Peninsula 92 30. Latitudinal variation of adult greatest length of skull in samples of Macaca fas- cicularis collected in the southern part of the Indochinese Peninsula and adjacent Isthmus of Kra, north of 10°N, compared with that in samples of parapatric M. mu- latta collected in the northern part of the Indochinese Peninsula 93 3 1 . Latitudinal variation of adult tail length in samples of Macaca fascicularis col- lected in the southern part of the Indo- Chinese Peninsula and adjacent Isthmus of Kra, north of 10°N, compared with that in samples of parapatric M. mulatta col- lected in the northern part of the Indo- Chinese Peninsula 94 List of Tables 1 . Dorsal pelage color saturation in core-area Macaca fascicularis: summary of varia- tion 16 2. Dorsal pelage color saturation in core-area Macaca fascicularis: geographic variation of mean value in 2-degree latitude-lon- gitude blocks 16 3. Dorsal pelage erythrism in core-area Macaca fascicularis: summary of varia- tion 17 4. Dorsal pelage erythrism in core-area Ma- caca fascicularis: geographic variation of mean value in 2-degree latitude-longitude blocks 17 5. Crown color pattern in Macaca fascicu- laris: summary of variation 18 6. Lateral facial crest pattern in Macaca fas- cicularis: summary of variation 20 7. External measurements and proportions in age/sex classes of wild-collected Ma- caca fascicularis 22 8 . Head and body length in core-area Macaca fascicularis: summary of variation 23 9. Summary comparison of head and body length in fringing-island and core-area samples of Macaca fascicularis 25 10. Tail length and relative tail length in core- area Macaca fascicularis: summary of variation 28 1 1 . Summary comparison of tail length and relative tail length in fringing-island and core-area samples of Macaca fascicularis 33 1 2. Cranial measurements and proportions in age/sex classes of wild-collected Macaca fascicularis 38 13. Schedule of emergence of deciduous teeth in captive Macaca fascicularis 38 14. Schedule of emergence of permanent teeth in captive Macaca fascicularis 39 1 5 . Greatest length of skull in core-area Macaca fascicularis: summary of variation 39 16. Summary comparison of greatest length of skull in fringing-island and core-area samples of Macaca fascicularis 42 17. Frequency of mtDNA types revealed by five restriction enzymes in six geographic samples of Macaca fascicularis 44 18. Frequency of alpha-globin gene haplotypes in geographic samples of Macaca fascicu- laris compared with frequencies in samples of M. mulatta and M. fuscata 45 19. Blood proteins: summary of frequencies of major alleles at polymorphic loci in 1 1 geographic samples of Macaca fascicu- laris 46 20. Summary of allele frequencies at the vita- min D binding protein locus in geographic samples of Macaca fascicular is 47 2 1 . Allele frequencies at the complement C6 locus in three imprecisely localized geo- graphic samples of Macaca fascicularis 48 22. Geographic variation of A-B-O and Lewis blood group phenotype frequencies in Macaca fascicularis 48 23. Geographic distribution of malaria spe- cies identified as natural parasites in Ma- caca fascicularis samples 49 24. Known vectors of malaria species that have been identified as natural parasites in Macaca fascicularis 49 25. Simian T-lymphotrophic retrovirus, type 1 : frequency of Macaca fascicularis in- dividuals or groups positive for anti- bodies 50 26. Recorded habitats of Macaca fascicu- laris 52 27. Recorded size of nonprovisioned troops of Macaca fascicularis 53 28. Ratio of sexually mature males to sexually mature females reported in nonprovisioned troops of Macaca fascicularis 54 29. Estimated home range and day range in troops of Macaca fascicularis 55 30. Foods reported eaten by nonprovisioned troops of Macaca fascicularis 56 3 1 . Daily waking-hour time budget estimates for Macaca fascicularis 57 32. Copulatory behavior data in two Macaca fascicularis groups 60 33. Macaque fossils or subfossils collected within geographic range of Macaca fas- cicularis 62 34. Summary comparison of external char- acters in recognized subspecies of Macaca fascicularis 66 35. Subspecies of Macaca fascicularis recog- nized as valid by Chasen (1940a) and re- cent authors 68 36. Macaca fascicularis aurea: external and cranial variation in adult specimens ex- amined 80 37. Sexual dimorphism of greatest length of skull compared in Macaca fascicularis fusca and Sumatran M. f fascicularis . . 82 38. Comparison of cranial proportions in In- dochinese Macaca fascicularis, M.f atri- ceps, and M. f condorensis 84 39. Hypothetical stages in evolution and dis- persal of Macaca fascicularis 88 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 Sumatra 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 Macaca fascicularis (Raffles, [1821]) is widely distributed in mainland and insular Southeast Asia from approximately 2 1°N to 1 0°S latitude and from 92°Eto 1 2 6°E longitude (Fig. 1). Vernacular names applied to this species include crab-eating, cynom- olgus, kra, and longtail macaque. The natural abundance of M. fascicularis is indicated by the large number of specimens in museum collections (Appendix 1), which greatly exceeds that of any other species of macaque. The existence of this species was first made known to Western science by the English seaman W. Dampier (1697, p. 220) and the German mis- sionary G. J. Camel (in Petiver, 1705, p. 2199), both of whom observed M. fascicularis in the Phil- ippines. The name "macaque" was first applied to this monkey by Buffon (in Buffon & Daubenton, 1766, p. 190), based on an imported specimen of unknown origin. The present review of Macaca fascicularis is based on examination of 2,049 museum speci- mens (Appendix 1 ), survey of relevant literature, and observation of natural populations in Thai- land and Bali. Specimens examined are preserved in the following institutions, which hereafter are cited by means of the following abbreviations: aiuz Anthropologisches Institut der Uni- versitat Zurich amnh American Museum of Natural His- tory, New York ansp Academy of Natural Sciences, Phil- adelphia bm(nh) British Museum (Natural History), London bnhs Bombay Natural History Society, Bombay ctnrc Centre for Thai National Reference Collections, Thailand Institute of Scientific and Technological Re- search, Bangkok fmnh Field Museum of Natural History, Chicago irsn Institut Royal des Sciences Natu- relles de Belgique, Brussels mcz Museum of Comparative Zoology, Harvard University, Cambridge, Massachusetts FIELDIANA: ZOOLOGY, N.S., NO. 81, NOVEMBER 30, 1995, PP. 1-206 Bell Museum of Natural History, University of Minnesota, Minne- apolis Museum National d'Histoire Na- turelle (Mammiferes), Paris Museum Zoologicum Bogoriense, Bogor Naturhistorisches Museum, Basel Naturhistorisches Museum, Bern Naturhistoriska Riksmuseet, Stockholm Natur-Museum Senckenberg, Frankfurt Philippine National Museum, Ma- nila Primate Research Institute, Kyoto University, Inuyama Royal College of Surgeons, Odon- tological Museum, London Rijksmuseum van Natuurlijke His- toric, Leiden Simian Conservation, Breeding & Research Center, Inc., Tanay, Lu- zon Sarawak Museum, Kuching Staatliches Museum fur Tierkunde, Dresden Museum of Zoology, University of Michigan, Ann Arbor University of the Philippines at Los Banos, College of Forestry, Los Banos, Luzon University of the Philippines at Los Banos, Zoology Department, Los Banos, Luzon National Museum of Natural His- tory, Washington, D.C. Zoological Laboratory of Utrecht University, Utrecht Zoologisches Museum des Hum- boldt-Universitat, Berlin Zoologisches Museum der Univ- ersitat Zurich Zoological Reference Collection, Department of Zoology, National University of Singapore Zoologisches Sammlung des Bay- erischen Staates, Munich Zoological Survey of India, Na- tional Zoological Collection, Cal- cutta Geographic names used in this review generally are officially approved names. However, the fol- lowing conventional names are retained because MMNH MNHN NHMBa NHMBe NHRM NMS PNM rcs(om) RMNH SICONBREC SMK SMTD UMMZ UPLBCF UPLBZD USNM ZLUU ZMB ZMUZ ZRC ZSBS ZSI of their greater familiarity (alternative names in parentheses): Burma (Myanmar), Java (Jawa), Lesser Sunda Islands (Nusa Tenggara), and Su- matra (Sumatera). Cited geographic names fre- quently include the following generic terms: gun- ong (Malaysian) or gunung (Indonesian) = moun- tain; ko (Thai) = island; pulau (Malaysian and Indonesian) = island; and teluk (Indonesian) = bay. This review begins with a general account of the geographic distribution of M. fascicular is. Follow- ing are discussions of (1) pelage characters, (2) external measurements and proportions, (3) cra- nial characters, (4) molecular and genetic char- acters, and (5) disease susceptibility; these discus- sions emphasize patterns of geographic variation. Subsequent sections summarize available infor- mation concerning natural history, reproduction, and paleontology. A discussion of subspecific tax- onomy, supplemented by a key to recognized sub- species, is based on material presented previously. For each recognized subspecies, a formal subspe- cies account provides the following: annotated synonymy; information concerning types, type lo- cality, and distribution; subspecific diagnosis; and summary of specimens examined (complete list in Appendix 1). The final section of this review pre- sents a hypothetical interpretation of the evolution and dispersal of M. fascicularis. Geographic Distribution Macaca fascicularis inhabits tropical Southeast Asia (Figs. 1, 2). Its natural geographic range ex- tends from southernmost Bangladesh and south- ern Burma southward and eastward through the southern part of the Indochinese Peninsula (south of 1 7°N), the Isthmus of Kra, the Malay Peninsula, Sumatra, Borneo, Java, and the Lesser Sunda Is- lands as far east as Timor, the Philippine Islands, and numerous small adjacent islands, including the southernmost three Nicobar Islands. The presence of M. fascicularis in some islands in Southeast Asia has been attributed to prehis- toric human introduction (see below, Fossils and Subfossils; Evolution and Dispersal). During his- toric times, successful introductions have occurred in Mauritius Island (I.), east of Madagascar, prob- ably in the sixteenth century (Sussman & Tatter- sall, 1986, p. 30); southwestern Sulawesi, date un- known (Miiller, [1840], p. 17; M. Weber, 1890b, p. 102; Dammerman, 1929, p. 6); Anguar Island, 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, fascicular is. For details, see Figure 2A-C. (Poirier & Smith, 1974, p. 264); Hong Kong, ca. 1945 (Marshall, 1967, p. 44; Southwick & Manry, 1 987, p. 48); and Pulau (P.) Tinjil, south of western Java, 1988-1990 (Kyes, 1993, p. 78). Physiographically, the geographic range of M. fascicularis includes mainland Southeast Asia, shallow- water islands on the Sunda Shelf, and deep- 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 Shelf and extended mainland Southeast Asia to include Sumatra, Borneo, Java, and other shallow- water islands (Heaney, 1986, p. 131; 1991a, p. 56). During the preceding glacial maximum, ca. 160 Ka, sea level may have been reduced by about 1 70 m, which exposed more of the Sunda Shelf but still left most deep-water islands isolated. To facilitate zoogeographic analysis, in subse- quent discussions of geographic variation the range of M. fascicularis is subdivided into three com- ponents: (1) a core area of distribution that com- prises mainland Southeast Asia, Sumatra, Borneo, and Java, four large landmasses that were inter- connected by dry land ca. 1 8 Ka; (2) shallow-water fringing islands— small islands within the 120-m bathymetric line that have been isolated from the core area for less than 1 8 K years; and (3) deep- water fringing islands— islands beyond the 120-m bathymetric line that have been isolated from the core area at least since the beginning of the pre- ceding interglacial, ca. 1 20 Ka (Van Couvering & Kukla, 1988, p. 461) and that may never have been joined to the core area. Pelage General Characterization (Fig. 5) Dorsal pelage color in M. fascicularis varies from buffy to yellowish gray to golden brown to reddish brown to blackish. Individual hairs generally are conspicuously marked by one or two pale subter- minal bands that vary in color from pale yellowish to golden to rufescent. The crown usually is more brightly colored than the back (pale subterminal bands more conspicuous in crown hairs), but in some specimens the crown is contrastingly darker and duller than the back (pale subterminal bands reduced or absent in crown hairs). The anterior edge of the crown is bordered by a transverse 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) 1. Bilasodia; Bimirdia; Ghorardia; Ochodia; Ruku- modia. 2. Jolir dia. Burma (Bu) 1. 2. 3. 4. 5. 6. 7. Lai char. Myengun Kyun. Arakan Division. Hainggyi Kyun. Preparis Island. Desertion Creek. Rangoon. 8. Pegu. 9. Wimpong. 10. Haungtharaw. 1 1 . Ataran River; Ye Forest. 12. Moscos Island Game Sanctuary. 13. Tavoy River. 14. Taungbyauk Chaung. 15. Kathema Kyun; Mibya Kyun. 16. Kaser Doo Wildife Sanctuary. 17. Kadan Kyun. 18. Great Tenasserim River; Mergui. 19. Tagoot; Tenasserim; Thagyet. 20. Letsok-aw Kyun. 2 1 . Lanbi Kyun. 22. Ban Sadein; Pakchan River, near Bankachon; Pak- chan River, near Maliwun. 23. Ru, Pulo. 24. Zadetkyi Kyun. Cambodia (C) 1. Siemreab. 2. Pang Roloem-Sur Sdei area. 3. Laem Ngop-Phumi Cham Yearn (Cambodia or Thailand). Laos (L) 1. Thateng, Muang. Nicobars (N) 1 . Ol-kolo-kwak vicinity. 2. Little Nicobar. 3. Kopenheat; Pulo Nyur. 4. Campbell Bay vicinity. Thailand (T) 1 . Ban Nam Lai Tai. 2. Ban Mae Na Ree. 3. Wong, Nam Mae, 53 mi (= 85 km) E of Um Pang. 4. Ping, Mae Nam; Wat Khao Noh. 5. Ban Pak Nam Pho; Wat Krieng Krai Klang. 6. Wat Khao Wong Kot. FIELDIANA: ZOOLOGY often form an irregular tuft or crest. Hairs on the side of the head are pale ochraceous gray to pale ochraceous brown and form a variably prominent lateral facial crest that usually extends from near the angle of the jaw to the crown, but may be restricted to the mandibular region. The face gen- erally is thinly haired; the facial skin is brownish to pinkish, except for the upper eyelids, which are sharply defined whitish. On the limbs, the proxi- mal part of the outer surface is approximately the same color as the adjacent surface of the trunk. More distally, the outer surface of the limbs be- comes paler, pale grayish to pale golden brown at the wrists and ankles. The dorsal surface of the tail is golden brown to blackish basally, becoming paler (pale grayish to grayish brown) distally. Pel- age on the ventral surface of the trunk, limbs, and tail is thin and pale gray to whitish. 9. 10. 11. 12. 13. 14. 15. 16. 17. 18. 19. 20. 21. 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49 Sarn Pra Karn. Wat Khao Sompoad. Lat Bua Khao. Ban Sakaerat. Aranyaprathet. Kosumphi Forest Park. Wat Koo Pra Kona. Ban Kosum. Phu Phan. Don Poo Tao; Prang Koo. Ban Wan; Wat Ban Kan Yai. Wong, Nam Mae, 40 mi (= 65 km) E of Um Pang. Kata Taek; Sap Khao; Thap Salao, Huai. Ban Tamrong Phato; Khwae Noi, Mae Nam. Ban Phu Toie. Khao Phatowee. Wat Noi Chompoo. Wat Phra Buddha Chai. Pak Chong, Sathani. Ban Huai Maenam Noi. Siam, 13°45'N, 99°25'E. Khao Suan Luang; Tham Chomphon; Wat Cha-Am Kiri; Wat Huai Takhaeng; Wat Khao Chong Phran; Wat Khao Khan Hok; Wat Khao Yod Thong; Wat Ngern Rung Sawang; Wat Ratch Singkhorn; Wat Tham Kunchhorn. Wat Tham Sala. Bangkok; Chao Phraya, Mae Nam. Khangkhao, Ko; Samuk, Khao; Si Chang, Ko; Si Racha vicinity. Wang Kaew. Laem Sing mountains. Ven-Ven. Chang, Ko. Kut, Ko. Khao Wang; Phet Buri; Samut Songkhram vicinity; Wat Ban Rai Don; Wat Bun Thawi; Wat Khao Ban- dai It; Wat Khao Takhrao; Wat Khao Tamon; Wat Kut. Pran Buri, Mae Nam.; Wat Khao Takieb. Khram Yai, Ko; Klet Kaeo, Ko. Khao Sam Roi National Park. Prachuap Khiri Khan, few miles north; Wat Tham- mikaram Varaviharn. Wat Tham Khao Phlu. Wat Tha Mai Lai. Chumphon, Khlong. Phayam, Ko. Laem Son National Park. Chiew Larn Reservoir. Thung Thong Waterfowl Reserve. Khlong Pah Yie, Ko Samui. 50. Phangan, Ko. 51. Khao Lampi-Hat Thai Muang National Park; Phangnga Bay National Park; Wat Tham Suwan Khuha. 52. Na Ka Yai, Ko; Yao Noi, Ko. 53. Rang Yai, Ko. 54. Phi Phi Don, Ko. 55. Lanta Yai, Ko. 56. Ban Phra Muang; Hat Chao Mai National Park; Kantang; Muk, Ko; Talibong, Ko. 57. Mu Ko Phetra National Park. 58. Butang, Ko. 59. Telok Wau, Ko Tarutao. 60. Ban Nong Kok; Ban Nong Put; Ban Thap Plik, 1 km NE; Hat Noppharat Thara-Mu Ko Phi Phi Na- tional Park; Tham Horn; Wat Tham Sua. 61. Tyching. 62. Khao Rang Kai. 63. Nakhon Si Thammarat. 64. BanNa. 65. Wat Khuha Sawan; Wat Suwankuha. 66. Nang Kham, Ko. 67. Khao Noi/Khao Tangkuan. 68. Nong Chik region; Pattani; Pho, Laem; Yaring, tidal creeks near; Yaring region. 69. Ban Sai Kau; Kampong Biserat; Wat Khuha Phi- muk. Vietnam (V) 1. Sontra Peak. 2. Lac Giao. 3. Xa Trang Bom. 4. Ho Chi Minh City. 5. Tay Ninh. 6. Phu Quoc Dao. 7. Airfield building vicinity, Con Son; Ben Dam vi- cinity, Hon Ba; Con Son. Absence reports a. North Andaman Island. b. Middle Andaman Island. c. Henry Lawrence Island. d. Barren Island. e. South Andaman Island. f. Little Jolly Boy Island. g. Rutland Island. h. Little Andaman Island. i. Car Nicobar Island. j. Tillanchong Island, k. Camorta Island. 1. Nancowry Island; Trinkat Island, m. Phai, Ko. FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS d Specimens examined Literature records a- n Absence reports """W-Maffn*. Java (J) 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) 1. We, Pulau. Simeulue, Pulau: 2. Dalam, Lhok; Sibaboh, Lugu; Survey Site LB; Sur- vey Site LS; Survey Site SEM. 3. Ajer Dingin; Labuhanbajau; Sinabang. 4. Lasia, Pulau. 5. Tuangku, Pulau. Nias, Pulau: 6. Lafau. 7. Soliga. 8. Hilisimaetano. 9. Samasama. 10. Siaba, Teluk. 1 1 . Mursala, Pulau. 12. Tanahmasa, Pulau. 13. Tanahbala, Pulau. Java (J), including nearby islands 1. Cilacap. 2. Kalipucang; Pangandaran. 3. Tjeringin. 4. Tasikmalaya. 5. Tilu, Gunung. 6. Malabar; Preanger district. 7. Bandung, 2000 ft; Bandung, near. 8. Ciwangi; Singkil, Gunung; Takokak Reserve. 9. Bantargebang; Pelabuhanratu, Teluk. 10. Jasinga. 11. Cihara. 12. Sarongen; Tamandjaija. 13. Camara. 14. Handeuleum, Pulau; Niur; Panaitan, Pulau; Peu- cang, Pulau; Ujungkulon, Suaka Margasatwa. 15. Danau, Rawa. 16. Gunung Halimun Reserve. 17. Bogor; Depok; Salak, Gunung. 18. Gede, Gunung; Pangrango, Gunung. 19. Cikarang forest. 20. Cikujang; Pasir Carolina. 21. Indramayu; Jatibarang. 22. Majalengka; "P. Sember." 23. Cirebon; Ciremay, Gunung; Linggajati. 24. Baturaden; Kaligoea. South China Sea (SCS) Pinang, Pulau [2]; Redang, Pulau. Acheh, Pulau. Pulau Tioman: Camp II, Juara side; Juara, Telok; Kampong Mukut; Nipah, Telok, vicinity; Sedagong; Tekek. Pemanggil, Pulau. Aur, Pulu. Tinggi, Pulau. Sentosa Island; Singapore Island: Bukit Timah Na- ture Reserve; Changi; MacRitchie Reservoir Nature Reserve; northwestern part; Pandan, Sungai; Pung- gol; Sembawang, Sungai; Singapore Botanical Gar- dens; western part. FIELDIANA: ZOOLOGY 8. Pulau Bintan: north coast; Pasir Panjang; Sungei Biru. 9. Mentigi, Pulau Mapur. 10. Tanjong Sauh, Pulau Batam. 1 1 . Galang, Pulau; Nguwal, Pulau. 12. Bakung, Pulau; Sebangka, Pulau. 13. Lingga, Pulau. 14. Selayar, Pulau. 15. Bulan, Pulau, south. 16. Durian, Pulau; Durian-kecil, Pulau; Sugi, Pulau; Ungar, Pulau. 17. Karimun-kecil, Pulau; Selatbliat, Pulau Kundur; Pulau Karimun: Mensuda Bay; Monos; Pemeral; Pongka, Kampung. Bangka, Pulau: 18. Jebus; Muntok; Pamuja, Tanjung; Rengsam, Tan- jung; Simpang. 19. Sungaiselan. 20. Batu, Tanjung, Pulau Belitung. 21. Serutu, Pulau. 22. Pai, Teluk, Pulau Karimata. 23. Pelapis Tengah, Pulau; Penebangan, Pulau. 24. Lemukutan, Pulau. 25. Pejantan, Pulau. 26. Benua, Pulau; Jela, Pulau; Selintang, Pulau; Tam- belan Besar, Pulau; Uwi, Pulau. 27. Airabu, Pulau; Piling, Pulau. 28. Jemaja, Pulau. 29. Telaga, Pulau. 30. Matak, Pulau; Mubur, Pulau; Siantan, Pulau. 31. Laut, Pulau. Natuna Besar, Pulau: 32. Ulu, Sungai. 33. Binjai, Sungai. 34. Lagong, Pulau. 35. Subi-kecil, Pulau. 36. Serasan, Pulau. Strait of Malacca (SM) 1. Burau, Pulau; Langkawi, Pulau. 2. Pulau Pinang [1]: Pantai Krachut; Penang Botanical Gardens; Penang Hill; Telok Bahang. 3. Pangkor, Pulau. 4. Pintu Gedong, Pulau. 5. Rupat, Pulau. 6. Kapos Tinggi, Pulau Bengkalis; Padang, Pulau, north coast. Sumatra (S) 1. Meulaboh vicinity. 2. Meulaboh, ca. 35 km NW. 3. Meulaboh, ca. 60 km NW. 4. Banda Aceh, ca. 40 km S and 45 km S. 5. Banda Aceh, near and ca. 20 km SSW. 6. Banda Aceh, ca. 30 km ENE. 7. Banda Aceh, ca. 85 km ESE. 8. Takengon, ca. 40 km NNW. 9. Bur ni Bebuli; Takengon, ca. 15 km NNW. 10. Lhokseumawe, 80 km ESE. 1 1 . Lesten. 12. Gunung Leuser Reserve. 1 3. Alas, Lae, between Agusan and Ketambe Research 14. 15. 16. 17. 18. 19. 20. 21, 22. 23. 24. 25. 26. 27. 28. 29. 30. 31. 32. 33. 34. 35. 36. 37. 38. 39. 40. 41 42, 43, 44 45 46 47 Station; Goenoengsetan-Meloewak; Gumpang, near; Ketambe Research Station; Ketambe, ca. 5 km S and 10 km S. Ketambe, ca. 30 km SSE. Alas, Lae, between Muara Setulen and Lae Renun, between Lae Renun and Bengkong River, and be- tween Bengkong River and Gelombang; Bukit La- wang, ca. 35 km SW; Renun, Lae. Alas, Lae, between Gelombang and mouth of river. Singkel vicinity. Tamiang. Medan, ca. 75 km NW; Pangkalansusu; Serangjaya- hilir; Sikundur. Bukitlawang; Bukitlawang, Area I and Area II; Bun- gara, Area III; Ober Langkat district; Serapit Tand- jung; Tandjung Bringin; Tandjung Butus; Unter Langkat district. Batangkuis; Belawan; Deli, Sungai; Labbuhandeli vicinity; Medan, forest near, near sea level, and vi- cinity; Medan, 20 km N; Medan-Siantar; Tanjung- morawa; Terbanjawan. Serdang district. Batu Bara district; Lauttador; Padang/Bedagei dis- trict; Paguruan; Tandjung; Tebingtinggi vicinity, ca. 25 km SSE and ca. 35 km ESE. Tebingtinggi, ca. 55 km ESE and ca. 65 km and 75 km SE. Bukitlawang, ca. 65 km S. Medan, ca. 100 km SSW. Dolok Tinggi Radja Reserve. Medan, ca. 105 km SSE. Tebingtinggi, ca. 75 km SSW. Pematangsiantar; Tebingtinggi, ca. 60 km SSW. Batang Garut. Tapanuli, Teluk. Padangsidempuan vicinity. Padangsidempuan, ca. 45 km NNW. Rantauprapat vicinity. Telukpanji. Bangko, ca. 15 km, 20 km, and 25 km ENE, ca. 20 km N, and ca. 30 km NNW. Rokan-kanan, Sungai. Bangko, ca. 80 km S. Bangko, ca. 90 km SSE; Pagansan; Pap-ka; Siak Co- patta; Sumatra, east-central. Tapung-kanan, Sungai. Gasip, Sungai, ca. 10 km and 20 km above mouth; Mandau, Sungai; Mempura; Perawang, 4 km NW; Siak, Sungai, near Pekanbaru, ca. 1 5 km, 30 km, 45 km, 60 km, and 75 km below Pekanbaru, ca. 60 km, 48 km, 36 km, 24 km, and 1 2 km above mouth, and near mouth. Lubuksikaping; Pinagar; Pinagar, ca. 5 km SW and ca. 20 km E. Bukit Cangang. Bukittinggi and vicinity and ca. 20 km S; Maninjau; Payakumbuh. Pajo. Lubukminturun area; Meru, Gunung; Padang; Pangkalan; Pasar Usang area; Sungailundang; Ta- rusan, Teluk. Kambang. Siulakderas. Sandaran Agong. FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. Telukkayubutih vicinity, ca. 15 km E, ca. 15 km NNE, ca. 1 5 km and 30 km WNW, ca. 30 km and 45 km ESE, and ca. 60 km SE. Kerinci, Gunung, northern foot; Pakan Selasa and ca. 10 km SSE. Sijunjung. Landai vicinity. Japura. Indragiri district. Kateman, Sungai. Indragiri, Sungai. Jambi, ca. 60 km NNW. Jambi, ca. 90 km ENE. Musi, Air, ca. 30 km and 1 5 km above Palembang, near Palembang, and ca. 12 km, 24 km, 36 km, 48 km, 60 km, 72 km, and 84 km below Palembang; Palembang district; Palembang, ca. 30 km, 40 km, 45 km, 50 km, 65 km, 70 km, 80 km, and 100 km N, ca. 30 km and 60 km NNE, ca. 40 km, 60 km, 65 km, and 70 km NNW, ca. 45 km and 60 km NW, and ca. 35 km SW. Harileko, Batang, ca. 12 km, 24 km, 36 km, 48 km, 60 km, 72 km, 84 km, and 96 km above mouth; Musi, Air, ca. 15 km, 30 km, 45 km, 60 km, 75 km, and 90 km above mouth of Batang Harileko; Musi, Air, ca. 50 km, 70 km, and 85 km above Palembang. Palembang, ca. 95 km and 100 km SW and ca. 90 km WSW. Lubuklinggau and vicinity and ca. 1 5 km N. Lubuklinggau, ca. 40 km NNW. Barisan Selatan. Kelabong, Bukit, vicinity. Jeleket vicinity and ca. 10 km W. Bengkulu. Bengkulu, ca. 40 km ENE. Sanggul, Bukit. Lubuklinggau, ca. 60 km SE. Baturaja, ca. 70 km WNW and ca. 75 km NW; Lahat. Baturaja, ca. 20 km and 30 km NNE. Palembang, ca. 60 km, 65 km, and 70 km SW. Muaradua. Suka Bandjar, NE and SE. Bukit Barisan Selatan, Taman Nasional. Wonosobo. Lampung, Propinsi. Kotabumi. Moro Batin and SSE; Telukbetung. Sukadana. Sungsan, ca. 50 km N; Kambas, Wai. Sungsan, ca. 20 km NNE. Kiambang. Kalianda. West Malaysia (WM) 1 . Kampong Titi Tinggi. 2. Perlis, Sungai. 3. Perangin, Bukit. 4. Pong. 5. Larut, Daerah; Ulu Ijok. 6. Hantu, Tanjong. 7. Cameron Highlands; Kinta, Daerah. 8. Telom, Sungai. 9. Berangkat, Gunong, east. 10. Terengganu, Sungai, vicinity. 1 1 . Kenyam, Sungai; Negara, Taman; Tahan, Sungai; Tembeling, Sungai. 1 2. Sungai Tekam Forestry Concession. 13. Berapit, Bukit, east; Kampong Cherok Paloh; Kam- pong Tanah Puteh; Kuantan; Pekan vicinity. 14. Kertau, Bukit. 1 5. Benom, Gunong; Benom, Gunong, NE slope; Kuala Lompat; Kuala Lompat Post, 0-1 km NW and 0- 2 km W; Lompat, Sungai, ca. 3 km and 4 km ENE of Kuala Serloh; Lompat, Sungai, ca. 3 km, 4 km, and 6 km W of Kuala Lompat Post; Patong, Bukit, ca. 1 km W; Tens, Sungai. 16. Keroh Forest Reserve; Telok Anson. 17. Morib. 18. Changkat Mentri; Cherakah, Bukit; Dusun, Sungai; Kampong Rantau Panjang; Kampong Sungai Buloh; Kuala Selangor; Kuala Selangor estuary; Lima Belas Estate; Pacific Tin; Port Swettenham vicinity; Tengi, Sungai; Tunggal, Bukit. 19. Bunga Buah; Damansara; Dusun Tua; Gaik Liew Estate; Kelang Road; Kuala Lumpur vicinity; Nan- as, Bukit; Templer Park; Ulu Gombak Forest Re- serve. 20. Pasoh Forest Reserve. 21. Lesong. 22. Endau, Sungai, vicinity; Kuala Rompin; Tanjong Panjair. 23. Telapak Burok, Gunong. 24. Menyala, Sungai; Port Dickson; Tanjong Tuan. 25. Kuala Pilah vicinity; Nuri Valley. 26. Melaka. 27. Nyalas. 28. Selai, Sungai. 29. Sekol, Sungai. Absence reports a. Berhala, Pulau. b. Jarak, Pulau. c. Sembilan Kepulauan. d. Babi, Pulau. e. Bangkaru, Pulau. f. Siberut, Pulau. g. Sipura, Pulau. h. Pagai Utara, Pulau. i. Pagai Selatan, Pulau. j. Enggano, Pulau. k. Anak Krakatau, Pulau; Rakata, Pulau; Rakata- kecil, Pulau; Sertung, Pulau. 1. Sebesi, Pulau. m. Datuk, Pulau. n. Temaju, Pulau. Absence report not represented because of map con- gestion: Sanglang-besar, Pulau (0°36'-0°38'N, 103°41'- 103°42'E). FIELDIANA: ZOOLOGY Sabah (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) 1. Bandar Seri Begawan; Brunei Bay area; Kampong Menuggol. Unmapped Brunei locality: Kuala Belalong Field Studies Centre (4°33'N, 1 15°08'E; for details, see Gaz- etteer, Appendix 2). Java (J), including nearby islands 1-24. See Figure 2B. 25. Candiroto. 26. Semarang. 27. Karimunjawa, Pulau; Kemujan, Pulau. 28. Pangonan. 29. Gedangan. 30. Ngawi. 31. Tamansari. 32. Tulungagung. 33. Kawarasan; Kediri district; Margomulio, Gunung; Manggis. 34. Wonokojo. 35. Meru Betiri Nature Park. 36. Mendit; Tengger, Pegunungan. 37. Pulau Doem area. 38. Bawean, Pulau. 39. Madura, Pulau. 40. Kangean, Pulau. 4 1 . Baluran Game Reserve. 42. Banyuwangi. 43. South Banyuwangi Nature Park. Kalimantan (K), including nearby eastern islands 1 . Cabang Panti Research Station; Gunung Palung Na- ture Reserve; Kampung Baru Study Area. 2. Ambawang, Sungai. 3. Senoeang. 4. Perbuah. 5. Roema Manoeal. 6. Semitau. 7. Sintang. 8. Riam. 9. Sekonyer Kan an River. 10. Tanjung Puting Reserve. 1 1 . Katingan, Sungai, right bank, 1 40 km above mouth. 12. Parit. 13. Katingan, Sungai, left bank, 160 km above mouth. 14. Katingan, Sungai, left bank, 200 km above mouth. 15. Kahayan, Sungai, right bank, 180 km above mouth of Sungai Rungan. 16. Liang Koeboeng. 17. Putussibau. 18. Poelau. 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 bank, 1 km below mouth of Sungai Busang and 2 km and 4 km above Muarajuloi; Julai, Sungai, right bank, 1 km above mouth of Sungai Busang, 3 km below mouth of Sungai Busang, and 6 km above Muarajuloi; Murung, Sungai, left bank, 1 km below mouth of Sungai Danau, 2 km above mouth of Sun- gai Beriwit, 2 km above mouth of Sungai Turusan, 2 km below mouth of Sungai Turusan, and 8 km above Muarajuloi; Murung, Sungai, left bank, at mouth of Sungai Beriwit, 1 km below mouth of Sungai Beriwit, 5 km above mouth of Sungai Tu- rusan, and 1 km below mouth of Sungai Turusan; TelukJolo, 12 km N. 20. Kahayan, Sungai, left and right banks, 1 20 km above mouth of Sungai Rungan. 2 1 . Kahayan, Sungai, left bank, 60 km above mouth of Sungai Rungan. 22. Rungan, Sungai, right bank, 50 km above mouth. 23. Rungan, Sungai, right bank, above mouth; Kahay- an, Sungai, left bank, above and below mouth of Sungai Rungan, and right bank, below mouth of Sungai Rungan. 24. Kapuas, Sungai, right bank, 25 km above mouth. 25. Barito, Sungai; Kambang, Pulau; Kaget, Pulau. 26. Pelaihari; Pleihari Tanah Laut Game Sanctuary. 27. Karangintan. 28. Rantau. 29. Hantakan; Telang. 30. Tanjung. 31. Buntok. 32. Kampong Hadjak; Muaratewe. 33. Purukcahu. 34. Tibang, Mt. 35. Badang. 36. Sungai Kayan-Sungai Mentarang Nature Reserve. 37. Sembakung, Sungai. 38. Long Peleben; Long Pangian. 39. Kaboerau. 40. Karangtigau, Tanjung. 41. Maratua, Pulau. 42. Berau, Sungai; Birang, Sungai. 43. Merah. 44. Karangan, Sungai; Pelawan, Sungai. 45. Kariorang. 46. Kutai Nature Reserve, northeast corner; Sengata; Sengata, Sungai, Kutai Reserve; Sengata, Sungai, at Mentoko camp and 1 km, 2 km, 3 km, 5 km, 7 km, 9 km, 1 1 km, and 1 3 km below; Sengata, Sungai, below Sengata village. 47. Juyan, Sungai, right bank, near mouth; Mentoko Research Center; Sengata, Sungai, right bank, 1 km below mouth of Sungai Nubung. 48. Kembangjanggut. 49. Goson Djerong, near; Jembayan, Sungai; Karang- mumus, Sungai; Loa Bambam; Mahakam, Sungai, left bank, 1 km and 4 km above Sebulu; Mahakam, Sungai, north bank, above Samarinda; Mahakam, Sungai, right bank, 8 km above Sebulu and 2 km below Sebulu; Tangarveng Island. 50. Sepaku, Sungai. 5 1 . Klumpang, Teluk. 52. Sebuku, Pulau. 53. Matasiri, Pulau. Lesser Sundas (LS) Bali, Pulau, including nearby islet: 1. Bali Barat National Park; Banjoe Wetan; Gilima- nuk; Pulaki, Tanjung; Sendang; Trima, Teluk. 2. Jembrana. 3. Bangli, near; Batur, Gunung; Botanical Gardens; Bratan, Danau; Bratan, Gunung; Buyan, Danau- Danau Bratan region; Catur, Gunung; Desa Poet- jang; Gitgit; Kukuh; Sangeh; Ubud. 4. Penida, Nusa. Lombok, Pulau: 5. Pengsong, Gunung; Pusuk, Gunung; Suranadi. 6. Kuta. 7. Pusuk forst; Rinjani, Gunung; Sewela. Sumbawa, Pulau, including nearby islet: 8. Ai Beta; Batudulang; Maman; Semongkat. 9. Moyo, Pulau. 10. Ampang area. 1 1 . Dompu; Oo vicinity. 12. Ntori; Raba; Rite. Flores, Pulau, including nearby islets: 13. Ginggo, Teluk, Pulau Rinca; Kode, Nusa; Mangia- tan, Pulau; Mbura, Pulau Flores; Nanga Look, Pulau Flores; Seraya Besar, Pulau. 14. Sano, Wai. 15. Rana Mese. 16. Bari. 17. Reo. 18. Pota. 19. Sika. 20. Solor, Pulau. 21. Adonora, Pulau. Sumba, Pulau: 22. Payeti-Kambaniru. 23. Mao Marroe. Timor, Pulau, including nearby islets: 24. Kambing, Pulau; Oeassa vicinity, Pulau Semau. 25. Amarassie. 26. Benu; Bokong; Kuatnana; Lelogama; Timau, Fatu. 27. Nikiniki. 28. Mutis, Gunung. 29. Fatuboi. Philippines Unnumbered locality: Bongao Peak, Bongao Island (5°01'N, 1 19°45'E; for details, see Gazetteer, Appendix 2). Sabah (Sab) 1 . Padas Bay. 2. Rayoh. 3. Papar. 4. Bongkabong; Rugading. 5. Talibang; Tuaran; Tuaran-Kampong Tenghilan Road. 6. Keningau. 7. Garau; Kampong Bundu Tuhan; Kampong Kiau; Kampong Kiau-Tenampok Pass; Kenokok; Kiau- lan; Kinabalu, Mount; Kinabalu National Park; Lumu Lumu; Ranau; Tempasuk, Sungai; Tenom- pok; Tinonkok. 10 FIELDIANA: ZOOLOGY Sex and Age Variation Geographic Variation Dorsal pelage color is generally similar in adult, subadult, and juvenile males and females, but pel- age in adult males tends to be longer and sleeker than in other age/sex classes. In newborn infants, dorsal pelage is blackish (on head, trunk, limbs, and tail), and the facial skin is bare, unpigmented, and pinkish (Dang, 1977, p. 11; Aldrich-Blake, 1980, p. 147; Crockett & Wilson, 1980, p. 170; Fittinghoff & Lindburg, 1980, p. 189; Wheatley, 1982, p. 205; Koyama, 1985, p. 106). The gradual transition from blackish neonatal pelage to grayish or brownish postneonatal pelage begins at about age 2-3 months, which is the age when deciduous canines and first molars erupt (Table 1 3) and in- fants begin to obtain some of their food indepen- dently (see below, Reproduction). The change of pelage color occurs first on the appendages, then on the trunk, and last on the crown. In specimens examined, the transition is complete well before eruption of the permanent first molars (age ca. 15 mo; Table 14). Dorsal Pelage Color Saturation— To in- vestigate geographic variation in dorsal pelage col- or saturation in M. fascicularis, specimens ex- amined have been compared with standard skins, as in a previous study of Philippine M. fascicularis (Fooden, 1991, p. 5). In the previous study, the range of standard saturation index (SI) values ex- tended from 1 .0 (pale yellowish brown) to 3.0 (dark brown). In the present study, this range extends from 0.5 (buffy; fmnh 9965 1 , Thailand: Kata Taek) to 4.0 (blackish; usnm 114168, Indonesia, P. Si- meulue: Sibaboh, Lugu) (Fig. 6). Skins of infants (specimens with deciduous teeth only) are exclud- ed from the analysis. Also excluded is the aber- rantly albinistic (pale ochraceous) skin of an adult female (amnh 107094) collected at Perbuah, northwestern Kalimantan; eight other specimens from the same locality are normally colored. In the core area of distribution of M. fascicularis (see above, Geographic Distribution), dorsal pel- age color in 640 postinfantile specimens varies 8. Balambangan, Pulau. 9. Banggi, Pulau, south, and Karakit; Maliangin Besar, Pulau. 10. Sabor, Pulau Banggi. 1 1 . Malawali, Pulau. 12. Sabatik, Pulau. 13. Ulu Segama Reserve. 14. Bukit Garam. 15. Lokan, Sungai. 16. Betotan; Lungmanis Station. 17. Labuk Road; Sandakan, 8 mi (= 13 km) W; Sibuga Besar, Sungai. 18. Sapagaya Forest Reserve. 19. Abai; Kinabatangan, Sungai; Trusan Kinabatangan. 20. Dewhurst Bay; Kretam Besar, Sungai; Kretam Kechil, Sungai. 21. Lahad Datu; Segama, Sungai. 22. Darvel Bay. 23. Tawau Hills National Park. 24. Tawau. Sarawak (Sar) 1. Samunsam Wildlife Sanctuary. 2. Penrissen, Gunung. 3. Paku Cave. 4. Bako National Park; Kuching; Kuching, 10th mile; Sarawak, Sungai, mouth. 5. Bukar, Sungai; Entawa, Tanjong. 6. Pelandok, Sungai. 7. Simunjan, Sungai. 8. Lingga. 9. Paku, Saribas. 10. Jumpit. 1 1 . Lanjak-Entimau Orang-Utan Sanctuary (proposed). 12. Belaga. 13. Dulit, Bukit. 14. Selikan, Bukit. 15. Similajau National Park. 16. Niah Caves. 17. Niah National Park. 18. Baram, Batang; Baram district; Miri; Miri, Sungai; Miri district. 19. Long Ekang. 20. Kalulong, Bukit. 2 1 . Gunong Mulu National Park; Melinau Gorge; Mulu, Gunung. 22. Punang, Sungai. Absence reports a. Manuk Manka Island, Philippines (see Fooden, 1991, p. 40). b. Masalembu Besar, Pulau. c. Sangeang, Pulau. d. Banta, Pulau; Bugis, Gili; Lawadarat, Gili; Lawa- laut, Gili; Longos, Nusa; Misa, Pulau; Papagaran- besar, Pulau; Sababi, Pulau; Sabajor Besar, Pulau; Sebolon Besar, Pulau; Siaba Besar, Pulau; Tatawa, Pulau. e. Kelapa, Pulau; Komodo, Pulau; Langkoi, Pulau; Lengah, Pulau; Mbarapu, Nusa; Motang, Gili; Pa- dar, Pulau; Padar-kecil, Pulau. f. Lomblen, Pulau. g. Pantar, Pulau. 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) (Tables 1 , 2); relatively few of these specimens (4.5%) are darker than medium brown (SI = 2.0). Pelage color in core-area specimens averages pal- est in the Indochinese Peninsula (mean SI = 0.99, n = 93), becomes darker southward and reaches maximum saturation in Sumatra (mean SI = 1.38, n = 178) and Borneo (mean SI = 1.47, n = 239), and then becomes somewhat paler farther south- ward in Java (mean SI = 1.26, n = 77). Within the Indochinese Peninsula, pale specimens are particularly concentrated near the northwestern margin of the species range, north of 1 4°N between 98°E and 102°E (mean SI = 0.73, n = 42). In all parts of the core area, variation in dorsal pelage saturation is relatively great, and overlap in sat- uration between samples from different parts of the core area is extensive. For example, a pale golden brown adult male collected in western Thailand (fmnh 99642, Ban Tamrong Phato) al- most perfectly matches an adult male collected in southern Sumatra (amnh 102765, Muaradua), and a pale yellowish brown adult male collected in east-central Sumatra (usnm 1131 69, Indragiri, Sungai) matches an adult male collected in western Java (usnm 156291, Bantargebang). Fringing-island specimens examined include 49 1 postinfantile skins— 234 collected in 71 shallow- water islands and 257 collected in 27 deep-water islands (Appendix 3). Although dorsal pelage sat- uration in most fringing-island specimens broadly overlaps that in core-area specimens, mean SI in fringing-island samples tends to exceed that in neighboring core-area samples (53 of 71 shallow- water fringing-island samples, 23 of 27 deep-water fringing-island samples). Strikingly dark fringing-island samples (con- spicuously darker than any other M. fascicularis samples examined) have been collected in six deep- water islands that constitute three geographic units: (1) Nicobar Islands (Katchall, Little Nicobar, Great Nicobar), northwest of Sumatra (mean SI = 3.9, n = 8); (2) P. Simeulue and P. Lasia, west of Su- 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- ripheral and disjunct distribution of these intense- ly saturate populations of M. fascicularis was not- ed previously by Kellogg (1944, p. 76). Dorsal pelage also is relatively dark in many islands of the Philippine Archipelago (deep-water); in west- ern, northern, and eastern Philippine islands, mean SI varies from 2.4 to 3.0 (cf. Fooden, 1991, p. 5). A brief, equivocal comment alluding to pelage sat- uration in M. fascicularis in deep-water P. We, north of Sumatra, has been published by Scheff- rahnetal. (1994, p. 136). The darkest shallow-water fringing-island sam- ple was collected in P. Karimunjawa, north of Java (mean SI = 2.7, n = 6; cf. Sody, 1949, p. 132). A shallow-water fringing-island sample from Ko Khram Yai, in the Inner Gulf of Thailand, also is conspicuously darker (mean SI = 1 .8, n = 10) than neighboring core-area specimens (cf. Kloss, 1919c, p. 347). Inconclusively small shallow-water island samples of dark specimens have been collected in P. Redang, east of the Malay Peninsula (mean SI = 2.5, n = 2), Ko Na Ka Yai, west of peninsular Thailand (SI = 2.5, n = 1), and P. Belitung, be- tween Sumatra and Borneo (mean SI = 2.3, n = 2). Dorsal Pelage Erythrism— Dorsal pelage er- ythrism in M. fascicularis has been investigated by comparing specimens examined with selected standard specimens, in a procedure parallel to that used in investigating saturation (see above). The range of standard erythrism index (EI) values ex- tends from 1.0 to 3.0, as follows: EI = 1.0, noner- ythristic, color of pale hair bands pale yellowish (fmnh 99651, Thailand: Kata Taek); EI = 2.0, moderately erythristic, pale hair bands golden (fmnh 33505, Vietnam: Ho Chi Minh City); and EI = 3.0, intensely erythristic, pale hair bands ru- fescent (fmnh 62913, Philippines: Palawan, Puer- to Princesa). Specimens marginal in erythrism be- tween primary standards have been assigned EI values of 1 .5 or 2.5. Infant skins are excluded from the analysis. In core-area samples, erythrism in M. fascicu- laris averages relatively high in Sumatra and in 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- amined, 24 are intensely erythristic (EI = 3.0); all of these intensely erythristic specimens originated either in Sumatra or in the adjacent Malay Pen- insula. Within Sumatra and the Malay Peninsula, intense erythrism apparently is an individual vari- able, not a local variable; in local samples (n > 1) that include intensely erythristic specimens, the frequency of such specimens varies from 0. 1 2 (Su- matra: Batangkuis; n = 8) to 0.50 (West Malaysia: Port Dickson; n = 4). Intense erythrism (EI = 3.0) occurs in relatively pale core-area specimens (zrc 4-075, West Malaysia: Port Dickson, SI = 1.0) as well as in darker specimens (zsbs C- 1 76, Sumatra: Kampong Baru, SI = 2.5). The high incidence of intense erythrism in M. fascicularis in the Malay Peninsula has been indicated previously by Kloss (1919c, p. 347) and Chasen (1940a, p. 67). Aggi- marangsee (1992, p. 119) reported on erythrism in a provisioned population of M. fascicularis at Kosumphi Forest Park, northeastern Thailand. In fringing-island samples, erythrism data are available for 234 postinfantile specimens collected in 7 1 shallow-water islands and 2 1 8 postinfantile specimens collected in 25 deep-water islands (Ap- pendix 4). In shallow-water fringing-island sam- ples, mean EI tends to exceed that in neighboring core-area samples (mean EI greater in 48 fringing- island samples, mean EI less in 1 6 fringing-island samples, mean EI equal in 7 fringing-island and core-area samples). Geographic variation of ery- thrism in shallow-water fringing-island samples generally parallels that in neighboring core-area samples: 1 3 intensely erythristic specimens (EI = 3.0) have been collected in 10 shallow- water fring- ing islands (Bintan, Bulan, Durian, Karimun, and Nguwal, in the Riau Archipelago, east of Sumatra; Redang, Siantan, and Tioman, east of the Malay Peninsula; Pinang [1] and Pintu Gedong, west of the Malay Peninsula; and Singapore); all 10 shal- low-water fringing islands from which intensely erythristic M. fascicularis specimens have been collected are adjacent to Sumatra or the Malay Peninsula, where intensely erythristic core-area specimens have been collected. In these 1 fringing islands, the frequency of intensely erythristic spec- imens varies from 0.09 (Tioman, n = 1 1) to 1.00 (Redang, n = 2). 14 FIELDIANA: ZOOLOGY Fig. 6. Dorsal pelage color saturation extremes in Macaca fascicular is— 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- atively high in most islands of the Philippine Ar- chipelago (Appendix 4) and is particularly high in Palawan, as noted previously (Fooden, 1991, p. 8). Erythrism is low in the very dark insular pop- ulations west of Sumatra (Appendix 4), and ery- thrism also is generally low in the deep-water Less- er Sunda Islands, as in nearby Java (Tables 3, 4). Crown Color— As previously indicated, the crown in M. fascicularis usually is more brightly colored than the back; however, in 63 of 1,103 postinfantile specimens examined (Table 5), the crown is darker than the back (Fig. 7) as a result of reduction or absence of pale bands in crown hairs. Darkness of the crown in these specimens varies from a dilute blackish wash to an irregular blackish streak to a clearly defined blackish patch. In dark-crowned specimens, the periphery of the crown patch usually is darker than the center. Of 648 core-area postinfantile specimens ex- amined, dark crowns are clearly defined in 1 and weakly defined in 13 (Table 5, Appendix 5). Thir- teen of these 14 dark-crowned specimens, includ- ing the one with a clearly defined dark crown, were FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 15 Table 1. Table 2). Dorsal pelage color saturation in core-area samples of Macaca fascicularis 1 : summary of variation (cf. N Frequencies (%) at SI values Mean SI Sample area 0.5 1.0 1.5 2.0 2.5 3.0 value Indochinese Peninsula 2 93 41.9 29.0 18.3 10.8 0.99 Malay Peninsula 3 53 11.3 34.0 45.3 9.4 1.26 Sumatra 178 4.5 23.6 67.4 1.7 1.7 1.1 1.38 Borneo 239 0.8 43.9 24.3 21.8 9.2 1.47 Java 77 10.4 41.5 35.1 10.4 2.6 1.26 All 640 9.8 35.0 38.5 12.2 4.2 0.3 1.33 1 Excludes infants. 2 Includes Isthmus of Kra north of 1 0°N. 3 Includes Isthmus of Kra south of 10°N. collected in the Indochinese Peninsula or adjoin- ing Isthmus of Kra north of 10°N (n = 94); one, with a weakly denned dark crown, was collected in Borneo (n = 242). The Indochinese specimen with a clearly defined dark crown (bm(nh) 1881. 6. 30. 2, adult male) was collected in southern Vietnam ("Cochin China"; cf. Elliot, 1909, p. 252; Kloss, 1926, p. 358). In fringing-island samples, most dark-crowned specimens also were collected in the vicinity of the Indochinese Peninsula and northern part of the Isthmus of Kra (Table 5, Appendix 5). Of 452 Table 2. Dorsal pelage color saturation in core-area Macaca fascicularis (n = 506) 1 : geographic variation of mean value in 2-degree latitude-longitude blocks (sample size indicated by italicized figures in parentheses) (cf. Table 1). Longitude (°E) 96- 98- 100- 102- 104- 106- 108- 110- 112- 114- 116- 118- Latitude 98 100 102 104 106 108 110 112 114 116 118 120 16-18 (°N) 1.0 (1) 0.6 (14) 1.0 (1) 1.5 (1) 14-16 0.8 (21) 0.8 (7) 1.0 (1) 12-14 1.1 (22) 1.0 (2) 1.0 (3) 0.5 U) 1.0 (1) 10-12 1.6 (6) 1.5 (12) 8-10 0.8 (8) 1.0 (1) 6-8 1.1 (6) 1.0 (3) 1.3 (20) 4-6 1.0 (1) 1.5 (6) 1.0 (1) 1.6 (8) 1.4 (16) 1.2 (29) 2-4 1.5 1.4 1.4 1.4 1.6 1.6 1.5 (1) (66) (76) (11) (12) (8) (2) 0-2 1.3 (2) 3.0 (1) 1.5 (32) 1.8 (6) 1.5 (1) 1.7 (3) 0-2 (°S) 1.4 (76) 1.0 (l) 1.5 (1) 1.2 (5) 2.1 (4) 1.7 (3) 2-4 1.0 (1) 1.2 (9) 1.8 (2) 1.4 (26) 1.4 (4) 4-6 1.1 (17) 6-8 0.5 (1) 1.1 (23) 1.3 (32) 1.3 (4) 1.0 (/) 8-10 1.0 (3) Excludes 1 34 imprecisely localized specimens that are included in Table 1 . 16 FIELDIANA: ZOOLOGY Table 3. Dorsal pelage erythrism in core-area Macacafascicularis 1 : summary of variation (cf. Table 4). N Frequencies (%) at EI values Mean EI Sample area 1.0 1.5 2.0 2.5 3.0 value Indochinese Peninsula 2 93 69.9 5.4 23.6 1.1 1.28 Malay Peninsula 3 53 37.7 17.0 30.2 5.7 9.4 1.66 Sumatra 180 23.3 8.3 56.7 1.1 10.6 1.84 Borneo 241 46.9 34.4 18.3 0.4 1.36 Java 78 57.7 23.1 19.2 1.31 All 645 44.2 20.2 30.8 1.1 3.7 1.50 Excludes infants. Includes Isthmus of Kra north of 10°N. Includes Isthmus of Kra south of 10°N. fringing-island postinfantile specimens examined, dark crowns are clearly denned in 1 7 and weakly denned in 32. Thirty-seven of the 49 dark-crowned specimens were collected in shallow-water islands adjacent to the Indochinese Peninsula or Isthmus of Kra. Of the 1 7 specimens with clearly defined dark crowns, 16 originated in three islands adja- cent to the Indochinese Peninsula: Ko Khram Yai, in the Inner Gulf of Thailand (n = 10: dark crown clearly defined, 7; weakly defined, 3); Con Son, in the South China Sea (n = 9: clearly defined, 8; weakly defined, 1); and Hon Ba, an islet adjacent to Con Son (n = 3: clearly defined, 1; weakly de- fined, 2) (cf. Kloss, 1916b, p. 32; 1919c, p. 347; Table 4. Dorsal pelage erythrism in core-area Macacafascicularis (n = 5 10) 1 : geographic variation of mean value in 2-degree latitude-longitude blocks (sample size indicated by italicized figures in parentheses) (cf. Table 3). Longitude (°E) 96- 98- 100- 102- 104- 106- 108- 110- 112- 114- 116- 118- Latitude 98 100 102 104 106 108 110 112 114 116 118 120 16-18 (°N) 1.0 V) 1.4 (14) 2.0 (1) 2.0 (1) 14-16 1.3 (21) 1.1 (7) 2.0 (1) 12-14 1.1 (22) 1.0 (2) 1.3 (3) 1.0 (1) 1.0 (/) 10-12 1.4 (6) 1.5 (12) 8-10 1.3 (8) 1.5 (D 6-8 1.4 (6) 2.0 (3) 1.1 (20) 4-6 2.0 (D 1.6 (6) 2.0 (1) 1.4 (8) 1.4 (16) 1.1 (29) 2-4 2.0 1.8 2.1 1.6 1.5 1.4 1.3 (D (68) (16) (11) (12) (8) (2) 0-2 1.8 (2) 2.0 (D 1.5 (32) 1.1 (6) 1.0 (/) 1.2 (3) 0-2 (°S) 2.1 (16) 1.3 (1) 2.0 (1) 1.4 (5) 1.4 (4) 2.0 (3) 2-4 1.5 (/) 1.6 (9) 2.5 (2) 1.5 (26) 1.4 (5) 4-6 1.5 (17) 6-8 1.0 (1) 1.2 (24) 1.2 (32) 1.1 (4) 1.5 (1) 8-10 1.0 (3) 1 Excludes 135 imprecisely localized specimens that are included in Table 3. FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 17 Table 5. Crown color pattern in Macaca fascicularis ] : summary of variation. For details, see Appendix 5. Frequencies of crown color patterns Crown with diffuse Crown with clearly Crown colored like blackish streak defined blackish N back, or brighter or wash patch 2 Sample area N % N % N % Core area 648 634 97.8 13 2.0 1 0.2 Indochinese Peninsula 3 94 81 86.1 12 12.8 1 1.1 Malay Peninsula 4 53 53 100.0 Sumatra 179 179 100.0 Borneo 242 241 99.6 1 0.4 Java 80 80 100.0 Shallow-water fringing islands 234 190 81.2 28 12.0 16 6.8 Deep-water fringing islands 221 216 97.7 4 1.8 1 0.5 Totals 1,103 1,040 94.3 45 4.1 18 1.6 1 Excludes infants. 2 Cf. Figure 7. 3 Includes Isthmus of Kra north of 1 0°N. 4 Includes Isthmus of Kra south of 10°N. 1921, p. 76; 1926, p. 358). In Ko Khram Yai specimens, the crown patch is distinctively narrow (Fig. 7) and is notably darker peripherally than centrally (see below, M.f. atriceps). The only non- Indochinese fringing-island specimen with a clear- ly defined dark crown is one of two specimens (usnm 144666) collected in Basilan I., a deep-wa- ter island in the south-central Philippines. Al- though the crown is blackish in M. fascicularis specimens collected in P. Simeulue and P. Lasia, deep-water islands west of Sumatra, the color of the crown in these specimens does not contrast with the color of the back, which also is blackish. Lateral Facial Crest Pattern (Fig. 8)— The Fig. 7. Crown color pattern in dark-crowned Macaca fascicularis— usnm 2366 1 8 (left), adult male, Thailand: Ko Khram Yai, M.f. atriceps; usnm 357241 (right), adult male, Vietnam: Hon Ba, 3.1 km W and 0.6 km S of Ben Dam, M. f condorensis. Scale bar 2 cm. FIELDIANA: ZOOLOGY 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 by the convergence of variably elongated poste- riorly directed hairs of the anterolateral facial re- gion and anteriorly directed hairs of the postero- lateral facial region. In most specimens examined, this crest sweeps upward from near the angle of the jaw to the lateral margin of the crown, passing between the eye and ear (transzygomatic crest pat- tern); less commonly, the crest is restricted to the mandibular region and terminates superiorly in a whorl low on the cheek (infrazygomatic crest pat- tern) (cf. Pocock, 1939, p. 78). In the infrazygo- matic pattern, unlike the transzygomatic pattern, hairs of the temporal region are smoothly directed posteriorly from the posterior margin of the eye to the anterior margin of the ear (anteriorly di- rected hairs are absent in this region); as a result, the infrazygomatic crest lacks the upper (preaur- icular) portion of the transzygomatic crest. Al- though the infrazygomatic pattern is relatively rare in M. fascicularis, it is common in M. mulatta, where the transzygomatic pattern occurs infre- quently (Stewart, 1933, p. 30). Transzygomatic and infrazygomatic patterns can be distinguished in the hair tracts of infants as well as in postinfants. In 819 specimens examined, the lateral facial crest pattern is transzygomatic in 728, infrazygo- matic in 83, and asymmetric (transzygomatic on FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 19 Table 6. Lateral facial crest pattern in Macacafascicularis 1 : summary of variation. For details, see Appendix 6. Frequencies of lateral facial crest patterns Asymmetric: infrazygomatic/ N Infrazygomatic Transzygomatic transzygomatic Sample area N % N % N % Core area 396 59 14.9 332 83.8 5 1.3 Indochinese Peninsula 2 98 57 58.1 37 37.8 4 4.1 Malay Peninsula 3 48 1 2.1 47 97.9 Sumatra 50 49 98.0 l 4 2.0 Borneo 163 l 5 0.6 162 99.4 Java 37 37 100.0 Shallow-water fringing islands 218 ll 6 5.0 204 93.6 V 1.4 Deep-water fringing islands 205 13 8 6.3 192 93.7 Totals 819 83 10.1 728 88.9 8 1.0 1 See Figure 8; tabulation includes infants. 2 Includes Isthmus of Kra north of 10°N. 3 Includes Isthmus of Kra south of 10°N. 4 Collected at Kotabumi (sample size = 1). 5 Collected at Mt. Tibang, Kalimantan (sample size = 1). 6 Includes 10 specimens collected in islands west of the Indochinese Peninsula and Isthmus of Kra (see Appendix 6) and one geographically anomalous specimen collected in Indonesia: P. Bintan (sample size = 8), southeast of Singapore. 7 Collected in islands west of the Isthmus of Kra and Malay Peninsula (see Appendix 6). 8 Collected in the Nicobar Islands, P. Simeulue, and P. Lasia (see Appendix 6). one side and infrazygomatic on the other) in 8 (Table 6, Appendix 6). Of the 83 specimens with the infrazygomatic pattern, 5 1 originated in a re- stricted area bordering the Bay of Bengal, at the northwestern margin of the species range (Fig. 9). No specimens with the transzygomatic pattern have been collected in this area, which includes south- ernmost Bangladesh, southern Burma, nearby shallow- water islands in the Mergui Archipelago, and a small part of west-central Thailand. The eastern boundary of the restricted area that is ho- mogeneous for the infrazygomatic pattern gener- ally coincides with the mountain ranges that form the border between Burma and Thailand. Of the remaining 32 specimens with the infra- zygomatic pattern, 1 7 originated in two areas ad- jacent to the restricted area homogeneous for the infrazygomatic pattern (Fig. 9); both of these ad- jacent areas are heterogeneous for the lateral facial crest pattern. Nine of these specimens originated at five scattered localities in the central and eastern parts of the Indochinese Peninsula, immediately east of the area homogeneous for the infrazygo- matic pattern, along the northern margin of the range of M. fascicularis; seven specimens with the transzygomatic pattern also have been collected in this area (central and eastern Indochinese Pen- insula), which includes two localities where spec- imens with both crest patterns have been collected together. Eight specimens with the infrazygomatic pattern originated at three localities in the eastern and southern parts of the Isthmus of Kra, im- mediately southeast of the area homogeneous for the infrazygomatic pattern; 1 1 specimens with the transzygomatic pattern also have been collected in this area (eastern and southern Isthmus of Kra, two adjacent west-coast islands). The remaining 1 5 specimens with the infrazy- gomatic pattern originated at localities distant from the restricted area homogeneous for the infrazy- gomatic pattern. Three of these specimens origi- nated in the Nicobar Islands, northwest of Su- matra, together with five specimens with the transzygomatic pattern. Ten specimens originated in P. Simeulue and P. Lasia, west of Sumatra, together with five specimens with the transzygo- matic pattern. One specimen originated in P. Bin- tan, southeast of Singapore, together with seven specimens with the transzygomatic pattern. The final specimen with the infrazygomatic pattern originated in north-central Borneo, at Mt. Tibang, Kalimantan. 20 FIELDIANA: ZOOLOGY V ' / 1 i 1 1 Area homogeneous for infrazygomatic pattern \ N \ _/ 20 - V^ S (' ; Areas heterogeneous for lateral facial crest pattern _ ^s» \ e # ^ N\ \ \ \ \ \ \ \ \ V* • v» N. • N /(»Ol 2 3 • <4sr!-(MO] --^ "No 2 / 0» i / O 3\ l f 6 x J v ^ [•OI\ 15 \ /_°2 \(«X>> \2 °2 • 1 •• 1 2 (CO)! 11 ° v '111 1 t— s 2°/ */' j 10 X £ \ / 9 IX 6> 3 Lateral facial crest pattern l ^i\ • Infrazygomatic 2 *\ v~_ o Transzygomatic °° \ 3 \»» 2 si c Asymmetric 80\ ° , V°2 5 l „— ..J ^-_ I — 95 100 105 Fig. 9. Geographic variation of lateral facial crest pattern in samples of Macaca fascicularis collected in the Indochinese Peninsula, Isthmus of Kra, Malay Peninsula, and neighboring islands (see Appendix 6). Numerals indicate number of individuals represented by symbols, where this number exceeds 1 ; heterogeneous samples collected at single localities are indicated by parentheses. Of the eight specimens with the asymmetric lat- eral facial crest pattern, seven originated in the two areas heterogeneous for the lateral facial crest pattern that are adjacent to the restricted area ho- mogeneous for the infrazygomatic pattern (Fig. 9). Two of these specimens originated in the central part of the Indochinese Peninsula and five origi- nated in the eastern part of the Isthmus of Kra or in islands adjacent to the southern part of the west coast of the Isthmus of Kra. The remaining spec- imen with the asymmetric lateral facial crest pat- tern originated, somewhat incongruously, in southeastern Sumatra, at Kotabumi. External Measurements and Proportions Sex and Age Variation Collectors' measurements of wild-collected specimens of M. fascicularis indicate that head and body length (HB) in adult males (mean = 465.6 mm, n = 238) averages approximately 13% more than in adult females (mean = 412.0 mm, n = 161) (Table 7). Weight in adult males (mean = 5.36 kg, n = 69) averages 49% more than in adult females (mean = 3.59 kg, n = 46). Relative length of appendages is similar in adult males and fe- males: mean relative tail length (T/HB x 100) is 1 17.0 (extremes 66.7-149.5) in 392 adults, mean relative hind foot length (HF/HB x 100) is 28.7 (extremes 23.2-37.8) in 365 adults, and mean rel- ative ear length (E/HB x 100) is 8.0 (extremes 4.8-11.1) in 301 adults. (Note: Karrer [1970, p. 171] reported that the tail is semiprehensile in captive M. fascicularis.) Relative length of the tail and of the hind foot in infants and juveniles exceed that in fetuses, sub- adults, and adults (Table 7); similarly, relative length of the ear in infants and juveniles exceeds that in subadults and adults (unknown in fetuses). This implies that, during prenatal and early post- natal life, the growth rate of the tail, hind foot, FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 21 Table 7. External measurements and proportions in age/sex classes of wild-collected Macaca fascicular is. Age/sex class 2 Head and body length (mm) Relative tail length 3 (T/HB x 100) Relative hind foot length 3 (HF/HB x 100) Relative ear length (E/HB x 100) Weight 3 (kg) Fetuses 82.5 78.8 23.7 — — 54-111 64.8-92.8 19.4-27.9 (2) (2) (2) Infants 254.8 ± 54.00 125.8 ± 24.52 33.6 ± 5.89 11.9 ± 2.51 0.76 ± 0.342 4 153-380 61.8-167.3 15.3-43.7 7.6-18.3 0.24-1.36 (52) (50) (51) (42) (13) Juveniles 361.3 ± 51.90 125.7 ± 15.83 32.0 ± 2.95 8.9 ± 1.74 2.36 ± 0.917 240-490 60.3-167.8 23.6^0.0 5.1-14.1 0.90-4.54 (208) (205) (197) (183) (75) Subadults — 120.4 ± 17.66 29.7 ± 2.24 7.9 ± 1.20 — 59.7-164.1 24.7-35.7 4.8-11.1 (106) (97) (81) Subadult females 397.6 ± 32.33 120.9 ± 17.63 29.8 ± 1.88 7.6 ± 1.40 3.41 ± 0.731 326^175 59.7-159.7 26.5-33.5 4.9-9.6 2.50-4.88 (39) (39) (36) (29) (14) Subadult males 453.8 ± 44.36 120.1 ± 17.81 29.6 ± 2.43 8.0 ± 1.06 5.15 ± 0.967 356-600 63.7-164.1 24.7-35.7 4.8-11.1 3.86-7.26 (67) (67) (67) (52) (23) Adults — 117.0 ± 14.50 28.7 ± 2.42 8.0 ± 1.34 — 66.7-149.5 23.2-37.8 4.8-11.1 (392) (365) (301) Adult females 412.0 ± 36.86 116.4 ± 15. 55 5 28.8 ± 2.43 7.8 ± 1.28 3.59 ± 0.690 6 - 7 315-545 70.4-148.4 23.5-36.2 5.0-10.5 2.35-5.44 (161) (159) (153) (131) (46) Adult males 465.6 ± 42.48 117.6 ± 13.37 8 28.7 ± 2.42 8.1 ± 1.37 5.36 ± 1.438 6 - 7 370-630 69.2-149.5 23.2-37.8 4.8-11.1 3.40-12.00 (238) (232) (212) (170) (69) 1 Mean ± SD (where n > 2), extremes, and sample size (italicized figures in parentheses). 2 Dental specifications: infants, deciduous teeth only; juveniles, some permanent teeth erupted; subadults, M3 in females or C in males incompletely erupted (cf. Spiegel, 1952, p. 129); adults, all permanent teeth completely erupted. 3 Cf. Spiegel (1985, pp. 27, 37, 53), who recorded postnatal growth of tail length, hind foot length, and body weight in captive M. fascicularis; cf. Shimizu et al. (1994, p. 175), who reported on effect of dietary restriction on growth in captive M. fascicularis. 4 In captivity, mean birth weight is 0.34 kg for females (n = 156) and 0.37 kg for males (n = 166) (Dang et al., [1993], p. 150; cf. Berkson, 1968, p. 352). 5 Excludes bobtailed specimen zrc 4-127, West Malaysia: P. Tioman, Telok Juara (HB = 409, T = 215, T/HB x 100 = 52.6). 6 Cf. J. Suzuki and Varavudhi (1989, p. Ill) and Varavudhi et al. (1989a, p. 225), who have reported higher mean weights in artificially provisioned troops studied in Thailand (adult females, 4.03-5.86 kg; adult males, 6.80-9.33 kg; cf. Aggimarangsee, 1992, p. 139). 7 Cf. Bakaretal. (1981, p. 12). 8 Excludes bobtailed specimens mzb 6483, Java: Cikujang (HB = 540, T = 250, T/HB x 100 = 46.3); and bm(nh) 1955.1511, Thailand: Ko Butang (HB = 403, T = 269, T/HB x 100 = 66.7). and ear exceeds the growth rate of the head and body and that subsequently— probably beginning in late infancy— the growth rate of the head and body exceeds that of the appendages (cf. Lumer & Schultz, 1941, p. 284; Karrer, 1970, p. 172; Yosh- idaet al., 1993, p. 438). Geographic Variation Head and Body Length— In 184 core-area adults, geographic variation of HB is generally similar in females and males (Table 8, Fig. 10). Surveying this variation from south to north, HB 22 FIELDIANA: ZOOLOGY Table 8. Head and body length in core-area Macaca fascicularis: summary of variation (cf. Fig. 10). Adult females Adult males Sample area N Mean ± SD Extremes N Mean ± SD Extremes Indochinese Peninsula 1 16 449.1 ± 50.26 354-545 3 531.0 ± 98.02 434-630 Malay Peninsula 2 11 393.6 ± 21.75 360-432 14 445.5 ± 46.62 370-519 Sumatra 5 420.6 ± 27.69 380-458 20 484.9 ± 37.04 420-551 Borneo 43 408.0 ± 27.34 354-469 42 473.1 ± 43.05 400-590 Java 11 435.1 ± 33.30 350-480 19 512.7 ± 46.29 434-610 All 86 418.0 ± 37.22 350-545 98 481.0 ± 49.24 370-630 1 Includes Isthmus of Kra north of 10°N. 2 Includes Isthmus of Kra south of 10°N. in both sexes is relatively great in Java (ca. 7°S); generally declines northward and reaches a min- imum at 0°-5°N in the Malay Peninsula, Sumatra, and Borneo; increases to about 12°N in the Isth- mus of Kra; and decreases farther northward to about 1 6°N, the latitude of the northernmost mea- sured specimens in the Indochinese Peninsula (cf. Aimi et al., 1982, p. 53). At similar latitudes in the Malay Peninsula, Sumatra, and Borneo, HB in each sex is approximately equal. The pattern of geographic variation of HB in core-area M. fas- cicularis parallels that previously reported for skull length in core-area samples of this species (Fooden & Albrecht, 1993, p. 526); for both HB and skull length, size variation in core-area M. fascicularis generally follows Bergmann's rule (cf. Mayr, 1963, p. 3 1 9), except for the aberrant northward decrease of size north of 1 2°N in the Indochinese Peninsula. One adult female collected at 7°01'S in Java ap- pears abnormally small (mcz 12757, Ban targe- bang, HB = 350 mm); a subadult female (usnm 156292, HB = 380 mm) collected at the same locality is larger. Fringing-island HB data are available for 215 adult specimens collected in 59 shallow- water is- lands and 2 1 deep-water islands (Appendix 7). In most of these islands, sample size is too small to determine conclusively whether or not insular HB differs from core-area HB at similar latitudes; however, mean HB in shallow-water fringing-is- land samples generally is less than in correspond- ing core-area samples (Table 9, Fig. 1 1). This pat- tern of HB variation in shallow-water fringing is- lands suggests a tendency toward insular dwarfism, as previously also observed for skull length vari- ation (Fooden & Albrecht, 1993, p. 525). HB is particularly small, compared with HB in core-area reference samples, in Pinang [1] (west of the Malay Peninsula) and Bali (east of Java). Among deep-water fringing-island samples, only those from the Lesser Sunda Islands show a clear tendency toward insular dwarfism (Table 9, Fig. 12, Appendix 7); in this respect, these samples resemble the sample from nearby Bali (see above), the shallow-water member of the Lesser Sunda group. In other deep-water fringing islands, HB in male samples tends to exceed that in core-area reference samples, and in Mindanao (Philippines) HB in both sexes conspicuously exceeds that in core-area reference samples. In one of four adult females collected in P. Simeulue (west of Sumatra; usnm 121513, HB = 325 mm), HB appears to be abnormally small; the skull and recorded body weight of this female are not particularly small (greatest skull length = 102.4 mm, weight = 3.74 kg). Tail Length— Collectors' measurements of tail length (T) are available for 183 core-area adults and 222 fringing-island adults (Table 10, Appen- dix 8). Although T averages less in females than in males, relative T (T/HB x 100) is approxi- mately equal in the two sexes, as indicated above (Table 7). Because geographic variation of T is not always concordant with geographic variation of HB, T variation and relative T variation are con- sidered separately in the following discussion. In core-area specimens of each sex, T tends to be negatively correlated with latitude (Fig. 1 3), in accord with Allen's rule (Mayr, 1963, p. 323). T averages greatest near the equator in the Malay Peninsula, Sumatra, and Borneo, somewhat less in Java (6°-8°S), and least at 15°-17°N in the In- dochinese Peninsula. In the Malay Peninsula, Su- matra, and Borneo, T— like HB (see above)— is approximately equal in each sex at similar lati- tudes. In Java, T is highly variable. T reduction in Indochinese M. fascicularis has been noted pre- viously (Fooden, 1964, p. 363; 1971, p. 29). FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 23 650 600 550- 500 ~ 450- E E O) c O X) T3 C CO ca CD X 400- 350 550 500 450- 400- 350 300 Males D oo $ + 01 o| -e-j O x x ..+_OJ< 4 00 o D o I o o &£ x.. O.l « CA > of FI > CA> islands CA 1 FI 2 islands CA 1 FI 2 1 . FI sample includes both sexes a. Sexes in FI sample deviate concordantly from those in CA sample b. Sexes in FI sample deviate discordantly from those in CA sample 1) Females, FI > CA; males, CA > FI 2) Females, CA > FI; males, FI > CA 2. FI sample includes only one sex a. Females b. Males Totals 13 10 1 0.5 0.5 1 0.5 0.5 5 2.5 2.5 5 2.5 2.5 7 2 5 2 1 1 33 7 26 8 5 3 59 15 44 21 10 11 1 FI sample mean exceeds CA sample mean. 2 CA sample mean exceeds FI sample mean. p. 1 00) for two core-area specimens identified by these authors as M. fascicularis; these specimens were collected in 1965 and 1967 at Sontra Peak, Vietnam (16°07'N, 108°18'E). The problematic measurements (in millimeters) are the following: adult male, total length = 730, T = 222 (HB = 508, relative T = 43.7%), and immature female, total length = 665, T = 230 (HB = 435, relative T = 52.9%). The skull of the male and the skin and skull of the female are preserved in the col- lection of the usnm (No. 356979, adult male; No. 356968, subadult female). Collectors' field notes, kept in the archives of the Division of Mammals, usnm (photocopies provided by Linda Gordon and David F. Schmidt), confirm the accuracy of measurements reported by Van Peenen et al. (1 97 1) for the male, but these notes indicate that total length of the female is 6 1 5 (HB = 385, relative T = 59.7%), not 665, as re- ported by Van Peenen et al. The collector's no- tation on the skin tag of the female likewise in- dicates that the total length is 615. Although both of these Sontra Peak specimens are identified by Van Peenen et al. (1971, p. 134; cf. 1968, p. 609) as M. fascicularis, the field notes indicate that they were originally identified by the collectors as M. mulatta. Dorsal pelage of the available skin of the subadult female (usnm 356968) matches that of M. fascicularis fmnh 33505 (Vietnam: Ho Chi Minh City) and therefore confirms the identification made by Van Peenen et al. Because the skin of the adult male (usnm 356979) is lacking, its identification cannot be in- dependently verified. This specimen may be either M. fascicularis or M. mulatta; Sontra Peak is near the border between the ranges of these parapatric species (Fooden, 1971, p. 28). At my request, P. F. D. Van Peenen recently reexamined his files, and he reports that he no longer has notes that would resolve this ambiguity (pers. comm., 27 Feb. 1992). Because of the unresolved ambiguity, data for the male (usnm 356979) are excluded from the present analysis of characters of M. fascicularis. In any event, the short tail of the subadult female (usnm 356968, relative T = 59.7%) is in accord with the pattern of tail reduction in M. fascicularis northward in the Indochinese Peninsula (see above). If the adult male (usnm 356979, relative T = 43.7%) had pelage characters of M. fascicu- laris, it would indicate that even more extreme tail reduction has occurred in this species at Sontra Peak. (Note: The tail also is relatively short in two infants and one juvenile— amnh 87266, ansp 15136, 15138— collected at nearby Muang Tha- teng, Laos.) In a core-area adult female (bm(nh) 1939.181) collected in 1 938 in West Malaysia at Bukit Nanas (formerly known as Weld's Hill; 3°09'N, 101°42'E), the tail is unusually short (T = 3 1 5 mm, relative T = 78.8%), considering the latitude of collection (Fig. 13). The tail also appears short in an adult male collected (date unknown) at the same locality (bm(nh) 1939.180, no collector's measurements; estimated relative T = ca. 75%, based on mea- FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 25 650 600 550 500- ~ 450 E E B 400 c O 350- X) C 550 n CO ■a co CD X 500-1 . Males ■ o o 400 350 300 O o o o a o <■ o o o <* o Females b ' OB ^ ■ o 10°S 10 Latitude |e o. . ■' LA j 0. 15°N ■ Shallow-water fringing-island specimens Core-area specimens Fig. 1 1 . Latitudinal variation of head and body length in adult shallow-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens (cf. Table 9 and Appendix 7). surements of dry skin). In another adult male col- lected in 1 9 1 3 at Bukit Nanas, T is normal (bm(nh) 1955.1512, T = 577 mm, relative T = 126.8%). Two additional short-tailed monkeys were ob- served at Bukit Nanas in 1966 (Bernstein, 1966, p. 1559; 1968a, p. 121); these two monkeys, which were associated with a troop of normal-tailed M. fascicularis, were inferred to be hybrids of M. fas- cicularis and M. nemestrina. In fringing islands, T data are available for small 26 FIELDIANA: ZOOLOGY 650 600 550 500 ~ 4504 E E ■g) 400 c o > o 350 X3 c 550 CO "O co CD X o s ^0 o o o oo o 0° A. ■ o ol i&o <>: ■- o o O I . -0 ■ o . 9 I A _. 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). samples of adults 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. 13, 14).' Tail length and relative tail length in core-area Macaca fascicularis: summary of variation (cf. Figs. Tail length Relative tail length 2 Sample area N Mean ± SD Extremes N Mean ± SD Extremes Adult females Indochinese Peninsula 3 16 434.4 ± 56.53 334-538 16 97.1 ± 10.95 76.1-114.7 Malay Peninsula 4 11 481.5 ± 63.44 315-556 11 122.6 ± 16.78 78.8-143.3 Sumatra 5 5 519.6 ± 12.18 508-535 5 124.0 ± 8.28 112.4-134.2 Borneo 43 507.3 ± 34.64 455-628 43 124.7 ± 9.78 110.2-144.0 Java 11 468.6 ± 52.37 345-555 11 108.5 ± 16.21 75.5-140.0 All 86 486.2 ± 52.91 315-628 86 117.2 ± 16.01 75.5-144.0 Adult males Indochinese Peninsula 3 3 513.3 ± 50.14 456-549 3 97.9 ± 11.28 84.9-105.1 Malay Peninsula 4 13 579.3 ± 41.02 515-640 13 132.6 ± 12.30 102.0-149.5 Sumatra 5 20 556.6 ± 38.59 490-623 20 115.3 ± 10.78 99.4-137.9 Borneo 6 43 576.7 ± 45.93 470-680 42 122.4 ± 11.32 103.4-147.2 Java 7 18 556.5 ± 84.86 360-715 18 110.1 ± 21.73 69.2-148.9 All 97 567.2 ± 54.33 360-715 96 119.2 ± 15.71 Both sexes 69.2-149.5 Indochinese Peninsula 3 19 97.2 ± 10.68 76.1-114.7 Malay Peninsula 4 24 128.0 ± 15.07 78.8-149.5 Sumatra 25 117.0 ± 10.76 99.4-137.9 Borneo 85 123.6 ± 10.57 103.4-147.2 Java 29 109.5 ± 19.52 69.2-148.9 All 182 118.3 ± 15.84 69.2-149.5 1 Cf. J. Suzuki and Varavudhi (1989, pp. 1 1 1-1 13), who reported tail length in artificially provisioned troops of M. fascicularis in Thailand (cf. Aggimarangsee, 1992, p. 104) 2 Relative tail length = tail length/head and body length x 100. 3 Includes Isthmus of Kra north of 10°N. 4 Includes Isthmus of Kra south of 1 0°N. 5 Cf. Bakaretal. (1981, p. 12). 6 One male specimen with tail length measurement lacks head and body measurement. 7 Excludes bobtailed specimen mzb 6483, collected at Cikujang; HB = 540, T = 250, T/HB x 100 = 46.3. ing islands, there is no general pattern of tail re- duction (Fig. 16, Table 11). In deep-water P. Si- muelue and P. Nias (both west of Sumatra) and in the deep-water Lesser Sunda Islands, T tends to be less than in core-area reference samples, but in the Nicobar Islands (northwest of Sumatra) and in many of the Philippine Islands, T tends to be greater than in core-area reference samples (Ap- pendix 8). The greatest reduction of T in fringing-island samples, relative to core-area reference samples, is evident in both sexes in P. Karimun and P. Bintan (both shallow-water islands east of Su- matra), P. Tioman (shallow- water, east of the Ma- lay Peninsula), and P. Simeulue and P. Nias (deep- water, west of Sumatra) (Figs. 15, 16; Appendix 8). T reduction may also be large in P. Pintu Ge- dong (west of the Malay Peninsula), P. Bangka (east of Sumatra), Ko Samui (east of the Isthmus of Kra), and P. Belitung (west of Borneo), shallow- water islands known from male samples only. The tail is unusually long in a male specimen collected in P. Acheh (shallow-water, east of the Malay Pen- insula, T = 698 mm; cf. Fig. 15). In P. Pinang [1] (shallow- water, west of the Malay Peninsula), one member of a troop of M. fascicularis had a de- formed tail (Lee Chin Thuan, 1964, p. 172). Relative T in shallow-water fringing-island samples is generally similar to relative T in core- area samples collected at similar latitudes (Fig. 1 7, Appendix 9); this implies that HB and T tend to be similarly reduced in shallow-water fringing-is- land populations, relative to HB and T in corre- sponding core-area populations. Conspicuous ex- ceptions occur in P. Tioman (east of the Malay Peninsula), where T is more reduced than HB, and, conversely, in Ko Khram Yai (south of the In- dochinese Peninsula), where HB is more reduced than T (cf. Figs. 11, 15). Relative T in deep-water fringing-island sam- ples frequently deviates from relative T in core- area samples collected at similar latitudes (Fig. 1 8, 28 FIELDIANA: ZOOLOGY 700 650 600 550 .".oo 450 400 c o 'c0 350 650 600 350 500 450 400- 350 ol □ | D 1 i La Males ° ° I °i $ *s x i + v \> + | J 1 ^ 00 i x X ! $ t + ? %> x |o * i + % ! O x*x |§ S{ + o °! o x t !# n ' + Ol + x X 1+ i ! o X D a a + |o° X 1 1 D 300 10°S Females D 1 ° x x ° i+ + i i 00 + X X a v x<0 D 1 r i I j D x x X i X* D 1 ! 1 x ; i ' i ' i ' j X 10 Latitude 15°N O Borneo □ Java x Mainland + Sumatra Fig. 13. Latitudinal variation of tail length in adult core-area specimens of Macaca fascicularis (cf. Table 10). Appendix 9). Relative T in P. Simeulue and P. Nias (west of Sumatra) is significantly less than at corresponding latitudes in West Malaysia, Su- matra, and Borneo, and relative T in Mindoro and Luzon (northern Philippines) is significantly great- er than at corresponding latitudes in the Indo- chinese Peninsula. In P. Simeulue and P. Nias, T is more reduced than HB, relative to HB and T in West Malaysia, Sumatra, and Borneo; in Min- doro and Luzon, both HB and T are increased— the latter more than the former— relative to HB and T in the Indochinese Peninsula, where both FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 29 150 5 Latitude 15"N 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. Figs. 10, 13). In the deep-water Lesser Sundas, as in Bali (cf. Fig. 17), HB and T tend to be iso- metrically reduced, relative to HB and T in Java. Cranial Characters Sex and Age Variation (Figs. 19, 20) In wild-collected M. fascicularis, greatest length of skull (GL) in adult males (1 18.7 mm, n = 454) averages approximately 1 8% greater than in adult females (100.4 mm, n = 439) (Table 12). Zygo- matic arches are relatively narrow in both sexes; relative zygomatic breadth (ZB/GL x 100) aver- ages 67.9 in 440 males and 65.9 in 428 females. The rostrum is relatively long and narrow, partic- ularly in males; the rostral-postrostral ratio (R/ PR x 100) averages 56.7 in 316 males and 47.6 in 24 1 females. Supramaxillary ridges and lateral maxillary concavities vary from weakly to strongly defined in both sexes. Canines in males are rela- tively large. A median sagittal crest, formed by progressive convergence of the temporal lines, is prominent in many old males. During development from infancy to adult- hood, zygomatic breadth increases slightly more rapidly than skull length, and rostral length in- creases much more rapidly than postrostral length (Table 12; cf. N. Fujiwara, 1963, p. 57; Morimoto, 1982, p. 98; Nanda et al., 1987, p. 217). Recent studies indicate that the conspicuous sexual di- morphism of R/PR in adult M. fascicularis prob- ably is a result of both a higher rate and a greater duration of allometric growth of the rostrum in males (Ravosa, 1991, p. 408; Richtsmeier et al., 1993a, p. 28); prenatal and postnatal growth pat- terns contribute differentially to development of adult cranial morphology (Richtsmeier et al., 1 993b, p. 322). Fluctuating asymmetry of the skull generally increases with age (Halgrimsson, 1993, p. 431). The sequence and timing of dental emergence 30 FIELDIANA: ZOOLOGY 700 650 600 550- 500- 450 400 350 | 1 ol | ■ ■ Males o 1 \ ■ o| CO | o ^> o _ o o.. ■ ■ M • a %% 9 & -j ■ ■ s ■o <£*> o ■ ■ » ■ §■■ ■ : f'T" t « ■ ■ |o ■ ol 7<>i 1 ■ i ■ 1 B o | 1 ■ ■ V * ■ ■■ I ■■ ■ ■ ■ !o° ■ ■ 1 ■ ■ 1 ■ o en c 9 650-i 600 J 550 500 450- 100 350- 300 Females o lo' o o ?$°l Od> ■ o d> o o <0 0" o % ■ ■ ■ ■ I 10°S 10 15°N Latitude ■ Shallow-water fringing-island specimens o Core-area specimens Fig. 15. Latitudinal variation of tail length in adult shallow-water fringing-island specimens ofMacacafascicularis compared with that in adult core-area specimens (cf. Table 1 1 and Appendix 8). in captive M. fascicularis have been investigated p. 250). Results of these investigations are in gen- by Spiegel (1952, p. 127), Berkson (1968, p. 354), eral agreement (Tables 13, 14; cf. Richtsmeier et Bowen and Koch (1970, p. 118), B. H. Smith et al., 19.93a, p. 4); however, Bowen and Koch (1970, al. (1994, pp. 214, 226), and Ostyn et al. (1995, p. 1 14) cautioned that the timing of dental emer- FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 31 700 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). gence in captivity may not be identical to that in and the last deciduous tooth (m 2 ) 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 of emergence of deciduous 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 core-area (CA) samples of Macaca fascicularis. 1 For details, see Appendixes 8 and 9. Shallow-water FI samples Deep-water FI samples Categories of FI samples Number of islands FI > CA 2 CA > FI 3 Number of islands FI > CA 2 CA > FI 3 Tail length 1 . FI sample includes both sexes a. Sexes in FI sample deviate concordantly from those in CA sample b. Sexes in FI sample deviate discordantly from those in CA sample 1) Females, FI > CA; males, CA > FI 2) Females, CA > FI; males, FI > CA 2. FI sample includes only one sex a. Females b. Males Totals All samples (including both sexes) 15 11 2 1 1 0.5 1 0.5 2 1 1 8 31 57 6 11.5 8 25 45.5 2 10 22 1 3 8 1 7 14 Relative tail length 57 25 32 21 8 13 1 Samples for tail length differ slightly from those for relative tail length because both head and body length and tail length are not available for all specimens. 2 FI sample mean exceeds CA sample mean. 3 CA sample mean exceeds FI sample mean. 1952, p. 129; 16 females, 22 males, not all teeth studied in all specimens). The first permanent tooth (M,) may emerge as early as age 1 .2 years, and the last permanent tooth (M 3 ) may not emerge until age 9.5 years (Table 14). The permanent canines of males emerge approximately 1 year later (C,, age 4.4 yr; C 1 , age 4.8 yr) than the much smaller permanent canines of females. Spiegel (1952, p. 128) indicated that the interval between initial emergence of a tooth and complete eruption to its final height is 1-2 months for most teeth, some- what longer for third molars, and several years for male canines. Geographic Variation In core-area specimens examined, skull length varies latitudinally (Table 15, Fig. 21), mostly in accord with Bergmann's rule, as previously re- ported by Fooden and Albrecht (1993, p. 525). In both sexes, skull length is high in Java (ca. 7°S), decreases to a minimum in Sumatra and Borneo (ca. 0°), increases to a second peak in the northern part of the Isthmus of Kra (ca. 1 2°30'N), and de- creases farther northward — contrary to Berg- mann's rule— in the Isthmus of Kra and Indo- chinese Peninsula (northernmost measured spec- imens collected ca. 1 7°N). Within each sex, skull length is approximately equal in specimens col- lected at similar latitudes in the Malay Peninsula, Sumatra, and Borneo; a similar latitudinal ho- mogeneity of skull length in these three disjunct landmasses was previously noted in M. nemestri- na (Fooden, 1975, p. 85). Four core-area skulls stand out as aberrantly large for their latitudes of collection (Fig. 21). Of these, one female skull (bm(nh) 1939.181, GL = 1 19.2 mm) and one male skull (bm(nh) 1939. 180, GL = 1 32.0 mm), both collected in West Malaysia at Bukit Nanas (3°09'N), belong to specimens that were previously cited for their aberrantly short tails (see above, External Measurements and Pro- portions); another adult male (bm(nh) 1955.1512, GL = 1 16.9 mm) collected at Bukit Nanas has a skull and tail of normal length. The other two aberrantly large skulls were collected at widely sep- arated localities in Borneo— a female (fmnh 68700, GL = 115.8 mm) in Sabah at Kretam Besar (5°32'N) and a male (usnm 521837, GL = 131.0 mm) in Kalimantan at Hantakan (2°38'S); the tail is of normal length in both of these specimens. Collectors' measurements of head and body length are available for three of the four specimens with aberrantly large skulls (all except bm(nh) 1 939. 1 80); in these three specimens, HB is large, but not aber- rantly so. In shallow-water fringing islands, geographic FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 33 50 140 I -jo 20- m i t no .c c V 100 ® 90 to DC 80 70- 60 Both sexes o ■ o . Si 0° •■■&■ ■ o. si i * * if ; ° 10°S 5 10 Latitude 15°N ■ 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 with Bergmann's rule, but skull length, like HB (see above), tends to be somewhat smaller in these islands than in core-area populations at similar latitudes (Fig. 22, Table 16); this indicates a gen- eral tendency toward insular dwarfism in shallow- water fringing islands (cf. Fooden & Albrecht, 1 993, p. 525). For four shallow-water fringing islands, samples are sufficiently large to establish that both females and males are significantly smaller than in core-area reference samples; these islands are P. Bintan (Indonesia: Riau Archipelago, 1°09'N), P. Tioman (West Malaysia: east coast, 2°48'N; in- cluding one aberrantly small male, usnm 101744, GL = 97.4 mm), Ko Kut (Thailand: southeast coast, 1 1°40'N), and Bali (Indonesia: Lesser Sunda Islands, 8°15'S). In five shallow-water fringing-is- land samples, skull length may be larger than in core-area reference samples; four of these islands are in the Java Sea, between Borneo and Java (Indonesia: P. Karimunjawa, 5°51'S; P. Kemujan, 5°51'S; P. Bawean, 5°48'S; P. Matasiri, 4°47'S), and one is north of Borneo (Sabah: P. Banggi, 7°09'N). In deep-water fringing islands, a tendency to- ward insular dwarfism is evident only in the Lesser Sunda Islands (8°32'-9°42'S) (Fig. 23, Table 16); skull length variation in deep-water Lesser Sunda Islands is similar to that in Bali, the only shallow- water Lesser Sunda Island (cf. Fig. 22). Skull length in deep-water fringing islands tends to be greater than in core-area reference samples in the Nicobar Islands (6°58'-8°00'N), northwest of Sumatra; in P. Simeulue (2°39'N) and nearby P. Lasia (2°10'N), west of Sumatra; and probably in P. Maratua (2°15'N), east of Borneo (cf. Fooden & Albrecht, 1993, p. 531). In the Philippine Islands (5°24- 1 8°3 1 'N), skull length tends to increase with lati- tude, extending the Bergmannian trajectory evi- dent in core-area populations between the equator and about 12°30'N (Fig. 23; Fooden, 1991, p. 10); variation of skull length in Philippine samples col- lected north of about 12°30'N does not follow the anti-Bergmannian trajectory evident in core-area 34 FIELDIANA: ZOOLOGY 150 140 3 130 O X 120 CO I 1 — t 110- o 100 0) 90 & 80 70- 60 O ■<£ |o 10°S Both sexes v o : o .o°x> 1 a^i * o ^t , \ M * lo = '. o- 00 5 10 Latitude 15°N ■ 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- mus of Kra and Indochinese Peninsula. Skull length is aberrantly small in one male specimen collected at Mahayahaya (13°55'N; GL = 109.3 mm), Lu- zon, Philippines. Molecular Biology and Genetics Mitochondrial DNA In a unique and valuable series of studies, Ha- rihara et al. (1986, p. 357; 1988, p. 1 18; 1991, p. 611) have employed five restriction enzymes to investigate restriction fragment length polymor- phism in the mitochondrial DNA (mtDNA) of six geographic samples of M. fascicularis (Table 1 7). One of these samples consists of five captive-born individuals— one born to a female imported from Vietnam and four born to females imported from Cambodia. Two samples consist of a total of 98 individuals livetrapped in Thailand; one of these samples originated north of the Isthmus of Kra (< 7 localities), and the other originated south of the Isthmus of Kra (2 localities). Each of the other three samples consists of 48 individuals, imported, respectively, from Malaysia, Indonesia, and the Philippines. For the last three samples, no further details are available concerning the regions or is- lands of origin of individuals in the samples or concerning possible geographic heterogeneity within the samples (Harihara et al., 1986, p. 357; 1988, p. 125); this is unfortunate, because Malay- sia, Indonesia, and the Philippines each include widely dispersed landmasses inhabited by isolated populations of M. fascicularis. In Table 1 7, the six sample areas of Harihara et al. are arranged in approximate geographic order. They are: (1) Vietnam and Cambodia, both east of (2) Thailand north of the Isthmus of Kra, which obviously is north of (3) Thailand south of the Isthmus of Kra; the latter sample area is north of (4) West Malaysia and northwest of East Malaysia 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 East Malaysia, together considered one sample area, are mostly north of (5) Indonesia; and Ma- laysia and Indonesia are southwest of (6) the Phil- ippine Islands. In the absence of information con- cerning whether any members of the Malaysian and/or Indonesian samples originated in Borneo, the geographic relationship of the Philippine sam- ple to the Malaysian and Indonesian samples re- mains ambiguous. For the four initially studied non-Thai samples, treatment of M. fascicularis mtDNA with each of the five restriction enzymes yielded three to six distinctive digestion patterns (enzyme morphs), as follows: BamHl and EcoRl, three morphs each; Hpal and Sstl, four morphs each; and Hi ndlll, six morphs (Table 17). In the 149 individuals in- cluded in these four samples, 21 different com- binations (mtDNA types) of these enzyme morphs were detected. In the 98 individuals included in the two subsequently studied Thai samples, four additional enzyme morphs and three additional 36 FIELDIANA: ZOOLOGY Fig. 20. Skull of adult 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 these two samples, all individuals were uniform in their mtDNA type (Harihara et al., 1991, p. 611). Except for the peculiar mtDNA type in the Thai- land south of the Isthmus of Kra sample (Harihara et al., 1991, p. 612), the distribution of enzyme morphs and mtDNA types in samples generally conforms to the geographic relationships of sam- ples (Table 17; cf. Melnick & Hoelzer, 1993, p. 3). Although the small Indochinese sample shares nei- ther of its two mtDNA types with other samples, it resembles the Malaysian sample in its high fre- quency ofSstl morph 2. Mitochondrial DNA types in the Thailand north of the Isthmus of Kra sample are closely related to those in the Indochinese and Malaysian samples (Harihara et al., 1 99 1 , p. 6 1 2). The Indonesian sample (10 mtDNA types), which FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 37 Table 12. Cranial measurements 1 and proportions in age/sex classes of wild-collected Macaca fascicularis. 2 Relative zygomatic Postrostral Greatest length breadth length Rostral- -post rostral Age/sex class 3 (mm) (ZB/GL x 100) (mm) ratio (R/PR x 100) Fetuses 60.8 56.7 52.0 23.7 U) (1) (/) V) Infants 75.9 ± 6.21 63.1 ± 2.48 64.2 ±4.11 25.7 ± 3.60 56.0-93.7 57.0-68.8 50.6-70.8 18.2-34.1 (80) (78) (57) (57) Juveniles 93.3 ± 9.40 64.6 ± 2.44 71.8 ± 4.08 40.6 ± 6.28 72.2-130.0 57.0-78.1 62.7-85.4 28.3-55.5 (482) (476) (287) (287) Subadult females 97.6 ± 4.54 65.6 ± 2.54 73.3 ± 3.12 44.0 ± 3.23 89.7-109.9 60.2-72.9 67.2-80.2 37.7-51.5 (70) (69) (40) (40) Subadult males 111.5 ± 6.70 66.9 ± 2.72 78.4 ± 3.52 53.0 ± 4.28 95.7-128.0 59.8-75.4 70.2-88.1 42.3-63.4 (123) (122) (92) (92) Adult females 100.4 ± 5.63 65.9 ± 2.40 74.0 ± 3.26 47.6 ± 4.21 84.0-119.2 59.4-74.1 64.3-87.2 36.9-61.0 (439) (428) (241) (241) Adult males 118.7 ± 7.29 67.9 ± 2.66 80.9 ± 4.02 56.7 ± 3.94 97.4-140.1 59.8-76.4 70.6-93.1 41.9-66.3 (454) (440) (316) (316) 1 For definition of measurements, see Fooden (1969, p. 40). 2 Mean ± SD (where n > 2), extremes, and sample size (italicized figures in parentheses). 3 Dental specifications: infants, deciduous teeth only; juveniles, some permanent teeth erupted; subadults, M3 in females or C in males incompletely erupted (cf. Spiegel, 1 952, p. 1 29); adults, all permanent teeth completlely erupted. may be geographically intermediate between the Malaysian and Philippine samples (see above), shares 2 mtDNA types with the Malaysia sample (9 mtDNA types) and shares 2 other mtDNA types with the Philippine sample (4 mtDNA types). The Philippine sample shares no mtDNA types with any sample other than that from Indonesia (cf. Lawler et al., 1995; p. 137). Known genetic dis- Table 13. Schedule of emergence of deciduous teeth in captive Macaca fascicularis 3 ; emergence age in days. Spiegel (1952, p. 131) Berkson (1968, p. 355) Bowen and Koch (1970, p. 118) Tooth Mean Extremes N Median 4 Extremes N Mean Extremes N ii 14 0-41 37 30 15-75 10 18 7-35 18 i 1 18 5-39 37 30 15-75 10 20 7^*2 18 la 26 0-58 37 30 30-75 10 36 14-49 18 i 2 40 18-77 37 45 30-75 10 47 21-63 18 c, 71 28-187 37 75 45-105 10 72 35-98 18 c 1 77 28-173 36 75 45-105 10 75 35-91 16 m, 1 m 1 J 66 35-98 37 90 90 75-120 75-120 10 10 74 72 56-91 56-98 20 17.5 s m 2 154 98-252 36 165 150-210 10 176 119-287 18 m 2 168 105-280 36 180 150-225 10 189 119-238 18 12 Designate positions of teeth. 3 Cf. B. H. Smith et al. (1994, p. 214), who cite the following mean emergence ages (days) without specifying sample sizes or extreme values: i,, 14; i\ 21; i 2 , 28; i 2 , 33; c„ 76; c 1 , 76; m„ 76; m 1 , 76; m 2 , 167; m 2 , 198. 4 Estimated to nearest half month. 5 Examined unilaterally in one subject. 38 FIELDIANA: ZOOLOGY Table 14. Schedule of emergence of permanent teeth in captive Macaca fascicularis 5 ; emergence age in years. Spiegel (1952, p. 134) 6 Bowen and Koch (1970, p. 119) Females Males Both sexes Tooth Mean Extremes N Mean Extremes N Mean Extremes N 7 M, M 1 I, I 1 I 2 I 2 M 2 M 2 P 3 P 3 P4 P 4 C, c M 3 M 3 1.38 1.54 2.38 2.38 2.62 2.85 3.38 3.54 3.62 3.92 3.38 3.85 5.54 6.23 1.23-1.69 1.31-1.92 2.08-3.23 2.31-3.00 2.38-2.92 2.38-3.15 3.00-3.69 3.23-3.92 3.23^.38 3.46-4.62 3.00-3.92 3.23-4.38 5.31-7.69 5.92-9.54 13 13 10 10 10 10 10 8 10 10 10 3 3 1.54 1.69 2.46 2.54 2.62 2.92 3.38 3.46 4.00 4.38 4.77 6.00 6.46 1.38-1.85 1.46-2.08 2.08-2.69 2.31-2.92 2.15-2.08 2.85-3.38 2.92-3.69 3.00-3.85 3.15-5.08 3.85-4.69 4.08-5.15 5.38-6.46 5.69-6.62 10 10 1.34 1.46 2.40 2.41 2.54 2.68 3.68 3.83 3.68 3.60 3.68 3.79 3.52 3.94 1.17-1.50 1.17-1.67 1.92-2.75 1.92-2.83 2.08-2.83 2.17-3.08 3.33-3.83 3.33^1.08 3.33-3.83 3.08-3.83 3.33-3.83 3.25-4.08 3.00-4.08 3.33-4.25 16 15 10 10.5 6.5 6.5 3 1.5 3 3 3 1.5 3 1.5 1-4 Designate positions of teeth. 5 Cf. B. H. Smith et al. (1994, p. 226), who cite the following mean emergence ages (yr) in females without specifying sample sizes or extreme values: M„ 1.50; M l , 1.75; I„ 2.50; I 1 , 2.50; I 2 , 2.50; I 2 , 2.50; M 2 , 3.25; M 2 , 3.25; P 3 , 3.50; P 3 , 3.25; P 4 , 3.50; P 4 , 3.25; C„ 3.00; C, 3.25; M 3 , 5.50; M 3 , 6.00. Also cf. Ostyn et al. (1995, p. 250), who provide the following emergence ages (yr; mean ± SD) for six teeth: females (n = 3)— M,, 1.33 ± 0.09; M 1 , 1.46 ± 0.09; I,, 2.54 ± 0.16; I 1 , 2.70 ± 0.08; I 2 , 2.70 ± 0.08; I 2 , 2.85 ± 0.19; males (n = 6, except I 2 , n = 4)-M„ 1.35 ± 0.11; M 1 , 1.53 ± 0.10; I„ 2.48 ± 0.21; I 1 , 2.50 ± 0.23; I 2 , 2.57 ± 0.24; I 2 , 2.70 ± 0.30. 6 In the present table, Spiegel's data are converted from "Messjahres" to calendar years [calendar years = (Messjahre x 48/52) + (Messmonate x 4/52)]; cf. Spiegel (1952, p. 127, footnote 2). 7 Decimal fractions indicate unilateral examination of subjects. tances between sample pairs also generally con- form to geographic relationships (Table 17, foot- note 1). Further interpretation of the taxonomic and phylogenetic significance of these mtDNA data is hindered by the imprecise provenance and pos- sible geographic heterogeneity of the samples from Malaysia, Indonesia, and the Philippines. Nuclear DNA Multiple Alpha-Globin Genes— Restriction- endonuclease analysis of M. fascicularis DNA (n = 168) indicates that individuals of this species have from one to four alpha-globin genes per hap- loid genome and that the frequency of alpha-glo- Table 15. Greatest length of skull in core-area Macaca fascicularis: summary of variation 1 (see Fig. 21) Adult females Adult males Sample area N Mean ± SD Extremes N Mean ± SD Extremes Indochinese Peninsula 2 22 105.0 ± 6.50 95.7-116.2 10 123.9 ± 5.12 116.4-132.3 Malay Peninsula 3 13 101.9 ± 7.33 92.6-119.2 20 115.0 ± 7.05 106.6-132.0 Sumatra 39 98.4 ± 3.42 93.1-103.9 80 116.2 ± 4.37 107.4-128.2 Borneo 71 99.1 ± 4.40 89.7-115.8 69 115.6 ± 5.36 103.2-133.5 Java 40 104.3 ± 4.26 97.0-114.0 44 126.3 ± 5.76 115.9-138.1 All 185 101.0 ± 5.40 89.7-119.2 223 118.2 ± 6.81 103.2-138.1 1 Includes specimens without precise localities that are excluded in Fooden and Albrecht (1993, p. 537). 2 Includes Isthmus of Kra north of 10°N. 3 Includes Isthmus of Kra south of 10°N. FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 39 145 ~ 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 H O 2 LU D CORE AREA no h 105 100 h 95 UJ 90 - _J < 85 - 80 B = BORNEO M = MAINLAND J = JAVA S = SUMATRA M — M~~~ M ^ \ ""M "^"-^^ M FEMALES 10°S 8°S 6°S 4°S 2°S 0° 2°N 4°N 6°N 8°N 10°N 12°N 14°N 16°N 18°N 20°N LATITUDE Fig. 2 1 . Latitudinal variation of greatest length of skull in adult core-area specimens of Macaca fascicularis (Fooden & Albrecht, 1993, p. 526). Solid lines indicate third-order polynomial regressions of skull length on latitude for females and males; dashed lines indicate 95% confidence limits. bin gene haplotypes varies geographically (Table 1 8 and below, Blood Proteins; Barnicot et al., 1966, p. 241; 1970, p. 380; D. G. Smith & Ferrell, 1980, p. 558). Judging from available evidence, the sin- gle-gene haplotype is rare, having been detected (heterozygously) in only 1 of 1 4 west-central Thai individuals and 1 of 20 Indonesian individuals. The double-gene haplotype is most common; it is very frequent (0.88-1.00) in Thailand north of the Isthmus of Kra; less frequent (0.52-0.62) in Thai- land south of the Isthmus of Kra, in Malaysia, and in Indonesia; and, somewhat incongruously, it is very frequent (0.98) in the Philippines. The triple- gene haplotype, conversely, is relatively rare (0- 0.12) in Thailand north of the Isthmus of Kra; moderately frequent (0.28-0.43) in Thailand south of the Isthmus of Kra, Malaysia, and Indonesia; and absent in the Philippines. The quadruple-gene 40 FIELDIANA: ZOOLOGY I F CD z LU D I F CD D XL CO lu 90 - _l < ^ 85 UJ LL 80 45 40 35 30 - 25 - 20 " 15 - 10 - 05 - 00 20 h 15 10 - 05 - 00 95 h SHALLOW-WATER FRINGING ISLANDS MALES REGRESSIONS AND CONFIDENCE LIMITS TAKEN FROM CORE-AREA ANALYSES FEMALES 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 of greatest 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- stricted to Malaysia, Indonesia, and the Philip- pines. In M. mulatta and M. fuscata, two other members of the fascicularis species group (Food- en, 1991, p. 2), the frequency of alpha-globin gene haplotypes apparently is similar to that in Thai- land north of the Isthmus of Kra. Highly Repeated Restriction Patterns— A study by Crovella et al. (1994, p. 66) of highly repeated nuclear DNA sequences has revealed that, for nine enzymes employed, restriction patterns are indistinguishable in Sumatran, Javan, and Philippine M. fascicularis. Eight of the nine en- 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 Macaca fascicularis. 1 For details, see Fooden and Albrecht (1993, Tables 2, 3). Shallow-water FI samples Deep-water FI samples Categories of FI samples Number Number of FI > CA > of FI > CA > islands CA' FP islands CA" FF 1 . FI sample includes both sexes a. Sexes in FI sample deviate concordantly from those in CA sample b. Sexes in FI sample deviate discordantly from those in CA sample 1) Females, FI > CA; males, CA > FI 2) Females, CA > FI; males, FI > CA 2. FI sample includes only one sex a. Females b. Males Totals 17 12 12 3 1.5 1.5 1 0.5 0.5 1 0.5 0.5 2 1.0 1.0 9 4 5 3 1 2 33 12 21 11 7 4 63 23 40 29 16.5 12.5 FI sample mean exceeds CA sample mean. CA sample mean exceeds FI sample mean. Hpal, Pstl, and Pvull— also failed to distinguish the three geographic samples of M. fascicularis from Indian and Chinese samples of M. mulatta. One enzyme— Xmn\— distinguished the M. fas- cicularis samples from the M. mulatta samples. Blood Proteins The most important body of data on blood- protein allele frequencies in natural populations of M. fascicularis is the product of a long-term research project that has been pursued since 1979 by Dr. Y. Kawamoto and associates (Kawamoto, 1982, p. 68; Kawamoto & Ischak, 1981, p. 240; Kawamoto & Suryobroto, 1985, p. 36; Kawamoto et al., 1981, p. 20; 1982b, p. 275; 1984, p. 135; 1987, p. 98; 1988, p. 172; 1989, p. 97; 1991, p. 600; Kondo et al., 1991, p. 20; 1993, p. 171; for references to other studies of blood-protein allele frequencies in this species, see Fooden & Lanyon, 1 989, p. 235). Dr. Kawamoto has generously made available an interim summary of his group's cur- rent data on M. fascicularis, partitioned into 1 1 regional or insular samples (Table 19, Appendix 1 0). Of the 2 1 polymorphic loci studied by this group, 14 exhibit relatively limited variation, with the same major allele common to all 1 1 samples; the 7 more variable loci are Alb, CA-II, Dia, HbA- II, IDH, Pi, and Tf (for key to abbreviations, see Table 19, footnote 1). As noted previously (cf. Fooden & Lanyon, 1989, p. 223; Kawamoto et al., 1991, p. 600), blood-protein polymorphism tends to be reduced in insular populations. Two dendrograms (Fig. 24a) of blood-protein relationships among these 1 1 samples have been published by Dr. Kawamoto's group (Kawamoto et al., 1991, p. 600; Kondo et al., 1993, p. 177); one of these dendrograms includes a sample of the introduced Mauritius population of M. fascicu- laris. A consensus tree (Fig. 24b) of stable clusters common to both dendrograms delineates group- ings that frequently are discordant with current and Late Pleistocene (ca. 1 8 Ka) land connections of the regions or islands where the respective sam- ples were obtained (cf. Fig. 3). The most conspic- uous of these discrepancies is the grouping of sam- ples from Thailand north of the Isthmus of Kra with samples from Bali (a shallow-water island that was connected to Java during the Late Pleis- tocene glacial maximum) and Lombok (a deep- water island that was connected to Sumbawa dur- ing the Late Pleistocene glacial maximum). Allele frequencies at two loci (HbA-II and Pi) tend to link the samples from Thailand north of the Isthmus of Kra with those from Bali and Lom- bok, and allele frequencies at one locus (Tf) tend to link the Bali and Lombok samples with each other but not with samples from Thailand north of the Isthmus of Kra (Table 1 9). None of the relevant HbA-II, Pi, or Tf alleles are exclusive to samples from Thailand north of the Isthmus of Kra, Bali, or Lombok. Similar allele frequencies 42 FIELDIANA: ZOOLOGY 145 _ 140 E E 135 ** X 130 i- n z 125 UJ —i _i 120 _i 73 * 115 CO UJ -1 110 < 5 105 100 DEEP-WATER FRINGING ISLANDS MALES REGRESSIONS AND CONFIDENCE LIMITS TAKEN FROM CORE-AREA ANALYSES 120 E \ b E 115 \ \ X __ \ \ c - ^Z Q L^ L X 110 \ \ — " x Bu O- M L H O z UJ _i _i _i 105 100 \ \ ^- \ \ ~ «" T X Bv x *kjw&-i — ** ^^^ \ 95 i ! \ E ^ X " ^ \ V> « v ^ ^"X \ UJ 90 ^ •*' \ _l *~ — — ■ < 2 UJ 85 FEMALES LL 80 ■ i i i i i i i i i i i i i i 10°S 8°S 6°S 4°S 2°S 0° 2°N 4°N 6°N 8°N 10°N 12 nit nit.fi |v iia Cambodia (98 > n of Kra Malaysia nesia pines no. Ba Ec Hi Hp Ss (n - 5 3 ) > 65") (n < 33 4 ) (n = 48) (n = 48) (n = 48) 20 1 1 1 2 2 80.0 21 1 1 4 2 2 20.0 22 5 23 5 7 ? ? ? 7 ? 7 7 ? 7 o } 100.0 246 ? ? 7 ? ? 100.0 6 1 1 3 1 3 10.4 7 1 2 1 1 2 10.4 15 1 1 1 1 4 2.1 16 3 1 1 1 2 2.1 17 1 1 3 1 2 2.1 18 1 1 5 1 2 2.1 19 1 1 1 3 2 2.1 1 1 1 1 1 2 60.4 6.3 3 1 1 1 1 3 8.3 37.5 5 1 2 2 2 1 29.2 8 1 2 2 1 1 8.3 9 1 3 1 2 1 8.3 10 1 1 4 1 3 2.1 11 1 1 1 4 3 2.1 12 1 3 6 2 1 2.1 2 1 1 1 1 1 2.1 52.1 4 1 2 1 1 1 2.1 43.8 13 1 2 1 2 1 2.1 14 2 2 1 1 1 2.1 1 Genetic distances between non-Thai sample pairs: Indochina-Malaysia, 0.00679 ± 0.001 15; Indochina-Indonesia, 0.01284 ± 0.00157; Indochina-Philippines, 0.02073 ± 0.001 10; Malaysia-Indonesia, 0.00848 ± 0.00184; Malaysia- Philippines, 0.01110 ± 0.00068; Indonesia-Philippines, 0.00270 ± 0.00058 (Harihara et al., 1988, p. 124). 2 Enzyme abbreviations: Ba = BamHl; Ec = EcoRl; Hi = Hindlll; Hp = Hpal; Ss = Sstl. 3 Vietnam, n = 1; Cambodia, n = 4. 4 Cf. Kawamoto et al. (1989, p. 95) and Harihara et al. (1991, p. 611). s Similar to mtDNA type nos. 1-21, particularly those in continental Southeast Asian samples (Harihara et al., 1991, p. 612). 6 Widely distinct from mtDNA type nos. 1-23 (Harihara et al., 1991, p. 612). Asia, Sumatra, Borneo, or Java. The Mauritius population, which apparently was introduced dur- ing the sixteenth century, probably originated from monkeys imported from Java. Computed genetic distances indicate remarkable divergence — ca. 100 times greater than expected— between allele fre- quencies in the Mauritius population and those in Asian populations of M. fascicularis; Kondo et al. (1993, p. 1 79) suggested that this may be the result of a genetic bottleneck (cf. Lawler et al., 1995, p. 138). Allele frequencies at the vitamin D binding pro- tein (DBP) locus have been studied by Tanaka et al. (1989, p. 104; 1991, p. 126) in 251 livetrapped Thai M. fascicularis and in 529 imported or cap- tive-bred Malaysian, Indonesian, and Philippine M. fascicularis (Table 20). Based on frequencies of the three most common alleles at this locus, the sample areas may be subdivided into three groups: (1) Thailand north of the Isthmus of Kra— DBP 1 very high (0.86-1.00), DBP 2 very low (0-0.01), DBP 3 very low (0-0.01); (2) Thailand south of the Isthmus of Kra, Malaysia, and Indonesia— DBP 1 moderately high (0.48-0.57), DBP 2 low (0-0.19), DBP 3 moderately high (0.30-0.40); and (3) Phil- ippines— DBP 1 moderately high (0.40), DBP 2 moderately high (0.59), DBP 3 absent. Allele frequencies at the complement C6 locus have been studied by Omoto et al. (1991, p. 603) in 150 imported or captive-bred Malaysian, In- donesian, and Philippine M. fascicularis (Table 21). The Malaysian and Indonesian samples are 44 FIELDIANA: ZOOLOGY 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). Region or island N Alpha-globin gene haplotypes Country Single Double Triple Quadruple M. fascicularis Thailand North of Isthmus of Kra East-central and southeast 30 1.00 West-central 14 0.04 0.96 Southwest 36 0.88 0.12 South of Isthmus of Kra 22 0.61 0.39 Malaysia Unknown 22 0.52 0.43 0.05 Indonesia Unknown 20 0.02 0.62 0.28 0.08 Philippines Unknown 24 M. mulatta 0.98 0.02 India Unknown 13 0.96 0.04 China Unknown 14 M . fuscata 0.93 0.07 Japan Unknown 30 0.02 0.96 0.02 similar in allele frequencies and differ from the Philippine sample (0.025 > P > 0.01), particularly in their lower frequency of alleles A and M4. The Philippine sample is notably less polymorphic than the Malaysian and Indonesian samples. In a recent study, Scheffrahn et al. ( 1 994, p. 135) investigated variation of blood-protein allele fre- quencies in M. fascicularis in Sumatra (nine groups representing four local populations) and four near- by islands (one shallow-water, three deep-water). Based on examination of five to seven polymor- phic loci, this study finds that (1) differentiation among six adjacent social groups in Sumatra is relatively great, (2) differentiation between these six adjacent groups and three other local popula- tions in Sumatra is roughly proportional to geo- graphic distance (25-350 km), (3) allele frequen- cies in a shallow-water fringing-island population in P. Tuangku (area ca. 300 km 2 ) are similar to those in the four local populations in Sumatra, (4) populations in the deep-water fringing islands P. Simeulue (ca. 2,000 km 2 ) and P. Nias (ca. 4,000 km 2 ), both west of Sumatra, cluster together, and (5), differentiation is greatest in P. We (ca. 250 km 2 ), a small deep-water fringing island north of Sumatra. Genetic drift is inferred to be the most important factor in blood-protein differentiation in insular populations. Although blood-protein al- lele frequencies in these M. fascicularis popula- tions in Sumatra and nearby islands are variably differentiated, all of these populations share one or more alleles at each of the five to seven loci. Blood Groups Limited information is available concerning geographic variation of human-type blood group frequencies in M. fascicularis (Table 22). In a study of the A-B-O system in M. fascicularis saliva and serum, type B was much less frequent in Malaysian (26.1%) and Indonesian (15.8%) samples than in a Philippine sample (76.9%); in an earlier study limited to A-B-O agglutinins in serum, type B ap- parently was similarly less frequent in Thai and Malaysian samples than in a Philippine sample. In M. mulatta, a species closely related to M. fas- cicularis and parapatric with M. fascicularis in Thailand (Fooden, 1980, p. 5), the frequency of type B is 97.0% (n = 200; Socha & Ruffie, 1983, p. 47; cf. Nakajima et al., 1970, p. 246); contrary to expectation, this is closer to the frequency in Philippine M. fascicularis than to that in Thai, Malaysian, and Indonesian M. fascicularis. Lewis group frequencies in M. fascicularis are similar in Thai, Malaysian, and Philippine samples. In a preliminary report on geographic variation of simian-type blood group H * in M. fascicularis, Honjo et al. (1984, pp. 73-74; cf. Terao, 1985, p. 5 1) indicated that the gene frequency of H 1 is mod- erately high in a Malaysian sample (n = 295), lower in an Indonesian sample (n = 284), and zero in a Philippine sample (n = 201). In simian-type blood group T^, the gene frequency of T 2 is represented (graphically) as lower in the Malaysian sample than in the Indonesian and Philippine samples. Nu- FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 45 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. Sampl e areas Thailand North of Isthmus of kra South Central and of south- South- Isthmus West Lom- Sum- Philip- Loci 1 i and east west of Kra Malaysia Sumatra Java Bali bok bawa Timor pines 2 alleles (727) (124) (33) (140) (276) (222) (136) (35) (81) (7) (142) Plasma proteins Alb A 84 20 52 94 90 100 100 100 100 93 100 B 16 80 48 6 8 7 Alp A 100 > 99 100 100 100 100 99 100 100 100 97 ChEs 1 100 100 100 100 100 99 100 100 100 100 100 Pi B 18 1 95 54 82 69 22 79 64 92 C 82 88 5 46 18 31 78 100 21 36 8 TBPA F 89 98 100 85 94 97 100 100 52 100 94 Tf C3 4 24 4 2 10 60 57 Dl 2 8 62 74 62 58 < 1 3 40 43 99 D2 32 13 2 1 G2 < 1 1 5 < 1 2 14 98 83 Erythrocyte proteins Acp A 97 99 97 96 > 99 100 100 100 100 100 100 ADA 2 100 92 100 100 100 100 100 100 100 100 99 AK 1 96 100 97 98 100 100 100 100 100 100 92 CA-I a 99 95 100 96 98 99 100 100 100 100 100 CA-II a 61 54 18 39 46 16 11 61 b 39 46 82 61 54 84 100 100 89 100 39 CellEs 1 100 100 100 99 > 99 97 100 100 80 86 95 Dia A 25 48 7 8 1 < 1 C 72 44 93 91 99 > 99 100 100 100 100 100 EsD 1 100 100 100 > 99 100 100 100 100 100 100 94 HbA-I 1 100 100 95 99 > 99 100 100 100 100 100 100 HbA-II 99 100 33 38 13 69 99 97 2 1 67 62 87 31 1 3 100 100 100 HbB 1 100 100 100 100 100 100 97 100 100 100 100 IDH 1 47 39 51 80 90 65 15 57 100 86 99 2 53 61 49 19 9 35 85 43 14 1 LDHA 1 94 97 100 100 98 100 100 100 100 100 100 PGD A 98 93 97 92 90 99 100 100 100 100 100 PHI 1 100 83 100 99 98 100 98 97 100 100 81 1 Abbreviations: Acp = acid phosphatase, ADA = adenosine deaminase, AK = adenylate kinase, Alb = albumin, Alp = alkaline phosphatase, CA = carbonic anhydrase, CellEs = esterase, ChEs = cholinesterase, Dia = NADH- diaphorase, EsD = esterase D, HbA = hemoglobin alpha-chain, HbB = hemoglobin beta-chain, IDH = isocitrate dehydrogenase, LDHA = lactate dehydrogenase-A, PGD = 6-phosphogluconate dehydrogenase, PHI = phosphohex- ose isomerase, Pi = protease inhibitor, TBPA = thyroxin-binding prealbumin, Tf = transferrin. 2 Island or islands unknown. merical values of these gene frequencies are not reported in this publication. Serum Cholesterol Response The effect on serum cholesterol level of two cho- lesterol-containing diets— one screening diet and one test diet— was studied in samples of adult male "Malayan" (?West Malaysian) and Philippine M. fascicularis (Taub & Bond, 1982, p. 339, abstract only, sample sizes not specified). In response to both diets, serum cholesterol level was signifi- cantly lower in the Malayan sample (screening diet, 235 mg/dl; test diet, 370 mg/dl) than in the Phil- ippine sample (screening diet, 353 mg/dl; test diet, 46 FIELDIANA: ZOOLOGY ■c 3 a. Dendrogram excluding Mauritius sample compared with dendrogram including Mauritius sample. Thailand, Cent. & SE 1 Thailand, SW 1 Bali Lombok Timor Sumbawa Sumatra W Malaysia Philippines Thailand, S 2 Java Mauritius c D- rC 4: Consensus dendrogram. Thailand, Cent. & SE 1 Thailand, SW 1 Bali Lombok Thailand, S 2 W Malaysia Sumatra — Philippines •• Java Sumbawa Timor - : .'-'.' -€ North of Isthmus of Kra. 2 South of Isthmus of Kra. Fig. 24. Blood-protein dendrograms for geographic samples of Macaca fascicular is: a. Dendrogram that ex- cludes Mauritius sample (Kawamoto et al., 199 1 , p. 600) compared with dendrogram that includes Mauritius sample (Kondo et al., 1993, p. 177). b. Consensus den- drogram. 487 mg/dl). In response to the screening diet, high- density lipoprotein levels were significantly higher in the Malayan sample (64 mg/dl) than in the Phil- ippine sample (5 1 mg/dl), and serum lipid levels also were consistently and significantly different; in response to the test diet, serum lipid levels were less consistently different. Karyology Hirai et al. (1991, p. 619) studied the chromo- somal G-banding pattern in 297 imported or cap- tive-born M. fascicularis individuals that origi- nated in Vietnam, Cambodia, Malaysia, Indone- sia, and the Philippines; the sample size for each country is not specified. No significant differences in banding pattern were detected among samples from these five countries, although one female from Vietnam exhibited possible interstitial hetero- chromatin in one autosome. The chromosome number of M. fascicularis was confirmed to be 42. Diseases Malaria Of the seven species of malaria {Plasmodium) that have been recorded as natural parasites in macaques, five species naturally infect hi. fasci- cularis (Fooden, 1994, p. 576). Of these, P. inui is quartan (72-hr asexual erythrocytic cycle); P. cy- nomolgi, P. fieldi, and P. coatneyi are tertian (48- hr cycle); and P. knowlesi is quotidian (24-hr cy- cle). Natural infections with all five of these species have been recorded in West Malaysia, which ap- parently is the center of malaria infection in M. fascicularis (Table 23). For P. fieldi, natural infec- tions are known only in West Malaysia. For P. coatneyi, infections also have been reported in the southern Philippine islands of Palawan (question- 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). Region or island N Alleles Country DBP 1 DBP 2 DBP 3 DBP* DBP 16 Other Thailand North of Isthmus of Kra East-central and southeast 46 1.000 West-central 57 0.860 0.140 Southwest 118 0.970 0.013 0.013 0.004 South of Isthmus of Kra 30 0.567 0.400 0.033 Malaysia Unknown 204 0.476 0.130 0.380 0.007 0.007 Indonesia Unknown 167 0.491 0.192 0.305 0.003 0.009 Philippines Unknown 158 0.405 0.592 0.003 FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 47 Table 21. Allele frequencies at the complement C6 locus in three imprecisely localized geographic samples of Macaca fascicularis (Omoto et al., 1991, p. 604). Alleles Hetero- Country N A Al A2 B Bl M2 M3 M4 Other 1 zygosity Malaysia 50 0.48 0.03 0.06 0.06 0.06 0.08 0.14 0.03 0.06 0.7298 Indonesia 50 0.43 0.05 0.01 0.08 0.02 0.08 0.18 0.05 0.10 0.7622 Philippines 50 0.66 0.10 0.09 0.14 0.01 0.5266 Total 150 0.52 0.03 0.02 0.05 0.03 0.09 0.14 0.07 0.05 - Seven minor alleles. been reported in Tioman, Java (questionable), Pa- lawan, and Cebu. For P. cynomolgi, infections have been reported from Cambodia in the north to Java in the south and from the Nicobar Islands in the west to Cebu in the east. Similarly, for P. inui, infections have been reported from Thailand in the north to Java in the south and from the Nic- obar Islands in the west to Mindanao in the east. Negative screening results suggest that malaria in- fections may be absent in M. fascicularis popu- lations that inhabit Luzon (n = 24) and Bali (n = 60). In malarious areas, the mean incidence of natural malaria infections in M. fascicularis is about 0.18 (n > 2,300); individual monkeys often are simultaneously infected with two or more species of malaria. Natural malaria infections in M. fas- cicularis and other macaques are relatively benign. The known vectors of malaria infections in M. fascicularis are five species of mosquitoes that be- long to the Leucosphyrus Group of the genus Anopheles (Table 24); geographic ranges of these five species of mosquitoes are partly overlapping (cf. Fooden, 1994, p. 581). One of these species, A. hackeri, is a proved vector of all five species of malaria that infect M. fascicularis. Individual mosquitoes sometimes harbor two or more species of malaria simultaneously. Although natural malaria infections in M. fas- cicularis are benign, untreated experimental infec- tions with one species of malaria, P. knowlesi, are Table 22. Geographic variation of A-B-O and Lewis blood group phenotype frequencies in Macaca fascicularis. Country Region or island Sample size and frequency (%) A-B-O groups (saliva, serum) 1 N B AB Malaysia Unknown Indonesia Unknown Philippines Unknown A-B-O agglutinins (serum) 2 N Anti-A Anti-B Anti-A + B Lewis groups (erythrocytes) 2 - 3 N Le"- positive O 245 45.4 26.1 26.9 1.6 165 55.1 15.8 27.9 1.2 130 2.3 76.9 19.2 1.6 None Thailand Southwest (north of Isthmus of Kra) 146 26.7 26.0 35.0 12.3 Malaysia West Malaysia 232 24.6 32.3 30.2 12.9 Philippines Mindanao 60 88.3 5.0 5.0 1.7 N Le b - positive Thailand Southwest (north of Isthmus of Kra) Malaysia West Malaysia Philippines Mindanao 149 4.0 86 4.6 231 4.8 181 10.5 60 3.3 60 8.3 1 Reference: Terao et al., 1981, p. 76 (cf. Honjo et al., 1984, p. 73). 2 References: Nakajima et al., 1970, pp. 246, 248; Omoto et al., 1970, p. 217. 3 Tested by direct agglutination with human anti-Le a serum and rabbit anti-Le b serum. 48 FIELDIANA: ZOOLOGY Table 23. Geographic distribution of malaria species identified as natural parasites in Macacafascicularis samples. For details, see Fooden (1994, p. 576). Geographic origin of M. fascicularis sample Malaria species (Plasmodium) cynomolgi fieldi coatneyi knowlesi sp. Thailand X Cambodia X West Malaysia X X Tioman X Nicobar Islands X X Sumatra Java X X Borneo ? Palawan X X Cebu X X Mindanao X almost invariably fatal in samples of M. fascicu- laris from the Indochinese Peninsula and possibly from the northern part of the Isthmus of Kra, both outside of the natural range of P. knowlesi (Food- en, 1994, p. 585); in continental Southeast Asia, P. knowlesi is not known to occur north of West Malaysia. Conversely, untreated experimental in- fections with P. knowlesi are relatively benign in samples of M. fascicularis from Cebu, which is within the natural range of this species of malaria. This suggests that populations of M. fascicularis that inhabit areas within the natural range of P. knowlesi have evolved partial resistance to the del- eterious effects of this parasite. Simian T-Lymphotrophic Retrovirus, Type 1 Samples of M. fascicularis populations in sev- eral areas have been tested for antibodies to reveal infections with simian T-lymphotrophic retrovi- rus, type 1 (STLV-1) (Table 25). Infections with STLV- 1 are relatively frequent in M. fascicularis in Indonesia, particularly in P. Sumbawa (Lesser Sunda Islands, between P. Lombok and P. Flores), and rare or absent in other natural populations sampled. STLV-1 infections also are fairly fre- quent in the introduced population of M. fasci- cularis in Mauritius. Susceptibility to Attenuated Polio Virus A retrospective analysis of the frequency of spi- nal cord lesions induced by inoculation with var- ious dilutions of two types of polio vaccines in- dicates that M. fascicularis captives imported from the Philippines were up to 1 ,000 times more sus- ceptible to induction of lesions than M. fascicularis captives imported from Indonesia, Malaysia, and the Indochinese Peninsula (Chino et al., 1992, p. 11). Natural History Habitats Although M. fascicularis is most commonly en- countered at low elevations, its known altitudinal range in the core area of its distribution extends to at least 1 200 m in West Malaysia (Gunong Be- nom; possibly to 1500 m in Cameron Highlands), to 2000 m in Sumatra (Bur ni Bebuli), to 1 800 m in Borneo (Lumu Lumu), and to 2000 m in Java (Gunung Salak); in the northern part of the core area (altitudinal data available for Bangladesh, Vi- etnam, Cambodia, and Thailand), this species has not been recorded above 400 m (Lac Giao, Viet- Table 24. Known vectors of malaria species that have been identified as natural parasites in Macaca fas- cicularis. For details, see Fooden (1994, p. 581). Malaria species (Plasmodium) Vector species cyno- coat- know- (Anopheles) inui molgi fieldi neyi lesi balabacensis ? ? ? ? dims, sensu lato x x hacked x x x x x introlatus x x leucosphyrus, sensu lato x FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 49 Table 25. Simian T-lymphotrophic retrovirus, type 1: frequency of Macaca fascicularis individuals or groups positive for antibodies. Data for individuals Data for groups Individuals Groups Refer- Origin N positive % N positive % ence Blood samples obtained from monkeys in country of origin Thailand, north of Isthmus ofKra 220 7 1 Thailand, south of Isthmus ofKra 30 2 1 Thailand 117 4 3.4 — — — 2 West Malaysia 199 — — — 2 Indonesia 245 34 13.9 23 8 34.8 3 Sumatra — — — 4 1 25.0 Java — — — 7 2 28.6 Bali — — 4 Lombok — — 3 Sumbawa — — ca. 50 5 5 100.0 Blood samples obtained from imported laboratory monkeys Malaysia 113 4 3.5 — — — 4 Indonesia 103 20 19.4 — — — 4 Philippines 83 - - - 4 Blood samples obtained from introduced population Mauritius 102 15 14.7 6 - - 5 1 Key to references: /, Ishida & Varavudhi, 1992, p. 164. 2, Ishida et al., 1985, pp. 840-842. 3, Hayami et al. 1983, p. 620. 4, Hayami et al., 1984, p. 181.5, Matsubayashi et al., 1992, p. 284. nam). In shallow-water fringing islands, the alti- tudinal range extends to 900 m in P. Tioman (Se- dagong) and 1 200 m in Bali (Gunung Bratan). In deep-water fringing islands, where M. fascicularis often is the only native monkey, the altitudinal range extends to 1 700 m in Lombok (Gunung Rin- jani), 2100 m in Timor (Gunung Mutis), 1950 m in Mindanao (Mt. McKinley), 1800 m in Negros (Canlaon Volcano), 1 900 m in Mindoro (Mt. Hal- con), and 2300 m in Luzon (Mt. Data). At low elevations, the preferred habitats of M. fascicularis apparently are seashore, riverbanks, and swamp forest (Table 26). Along the seashore, mangrove forest is reported as a frequent habitat of this species from Bangladesh in the north (M. A. R. Khan & Wahab, 1983, p. 102) to Flores in the south (Auffenberg, 1 98 1 , p. 242): the only oth- er species of monkeys that regularly inhabit man- grove forest in Southeast Asia are two colobines, Trachypithecus cristatus and Nasalis larvatus. Ma- caca fascicularis also has been reported inland in lowland secondary and primary forest and, less frequently, in upland secondary and primary for- est. At nearly 2000 m in northern Sumatra, Volz (1912, p. 88) reported, "To my astonishment, up here in this cool damp high-altitude vegetation, I still observed several monkeys, namely Macacus cynomolgus [= M. fascicularis]" (translated from German). This species also inhabits heavily dis- turbed areas, including the immediate vicinity of villages and farms; following logging, the density of M. fascicularis sometimes increases (Johns, 1992, p. 438). Arboreality/Terrestriality On the seacoast and riverbanks, M. fascicularis often forages on the ground (25% of encounters in Sumatra [Crockett & Wilson, 1980, p. 165], 26.5% of encounters in Kalimantan [Fittinghoff & Lind- burg, 1980, p. 189]). In inland forests, M. fasci- cularis is predominantly arboreal (98% in West Malaysia [Bernstein, 1967, p. 204; Aldrich-Blake, 1980, p. 158], ca. 97% in Kalimantan [Wheatley, 1980, p. 216]). This species apparently favors the lower and middle forest strata; in more than 50% of forest sightings, troops were in branches less than 20 m above the forest floor (West Malaysia [Aldrich-Blake, 1980, p. 159; cf. Harrison, 1961, p. 14], Sumatra [Rijksen, 1978, p. 127], Kaliman- tan [Rodman, 1978, p. 476]). Observations in Sumatra indicate that smaller groups of M. fasci- cularis tend to occur higher in the canopy (van 50 FIELDIANA: ZOOLOGY Schaiketal., 1983b, p. 216; D. R. Vosetal., 1992, p. 388). In response to danger, troops may flee either on the ground (Burma [Tickell, 1854-1875, p. [17], Great Nicobar I. [Das & Ghosal, 1977, p. 265], Sumatra [Volz, 1912, p. 369; Ulmer in Mil- ler, 1942, p. 127], Lesser Sunda Islands [B. Rensch, 1930, p. [17]; Mertens, 1936, p. 320]), in the can- opy (Thailand [Fooden, [1975], p. 100], West Ma- laysian [Bernstein, 1967, p. 204], P. Simeulue [Su- gardjito et aL, 1989, p. 200], Sumatra [Crockett & Wilson, 1980, p. 165], Kalimantan [Kurland, 1973, p. 250; Wheatley, 1980, p. 216]), or partly on the ground and partly in the canopy (Thailand [Food- en, [1975], p. 100]). Accidental falls of this species out of trees have been observed in northern Su- matra (Karssemeijer et al., 1990, p. 286). Details of locomotor behavior are discussed by Kurland (1973, p. 250), Fleagle (1980, p. 198), Cant (1988, p. 31), and Cannon and Leighton (1994, p. 512). At night, M. fascicularis usually sleeps in tall, rel- atively bare trees near a river (Singapore [Ridley, 1906, p. 142], Sumatra [Moszkowski, 1909, p. 24], Sarawak [Hornaday, 1910, p. 358], and numerous later accounts, particularly Fittinghoff & Lind- burg, 1980, p. 190 -Kalimantan). Swimming Observers agree that M. fascicularis frequently enters bodies of water, apparently often for pleas- ure, and that it is an excellent swimmer. In Burma, a wounded male that was attempting to escape from humans swam about 50 m underwater (Tick- ell, 1854-1875, p. [18]). In a captive insular group of M. fascicularis in Japan, a young male that was defeated in a dominance fight apparently swam 100 m to a breakwater (Furuya, 1965, p. 325). However, during the course of 3 years in the same captive insular group, most of about 60 individ- uals that were driven away by dominant group members died by drowning; none apparently reached the next nearest island, which is about 250 m distant and which may not have been vis- ible to the escaping monkeys. Drownings also have been reported in natural populations in coastal Thailand (Aggimarangsee, 1992, p. 130). Troop Size and Composition Mean size of nonprovisioned troops of M. fas- cicularis generally varies from about 12 to 25 (Ta- ble 27). Mean troop size is exceptionally small (ca. 5) in a highly disturbed area in Vietnam and ex- ceptionally large in western Java (ca. 47, n = 5 troops) and Mauritius (77.5, n = 2 troops, intro- duced population). Crockett and Wilson (1980, p. 149) suggested that troop size in Sumatra may be smallest in mangrove forest, larger in inland pri- mary forest, and largest in inland secondary forest (cf. Ruiter, 1993, pp. 90-91), but the observed differences are acknowledged not to be statistically significant. In provisioned troops in Thailand, troop size (mean = 76.9,n = 33 troops) generally exceeds that in nonprovisioned troops (Aggimar- angsee, 1992, p. 150). Solitary males, not associ- ated with any troop, have been reported in Thai- land (Fooden, 1971, p. 24), Singapore (Furuya, 1965, p. 294), Sumatra (Wilson & Wilson, 1973, p. 5; Rijksen, 1978, p. Ill; Norikoshi, 1984, p. 5), Sabah (Kurland, 1973, p. 253), Java (Hoog- erwerf, 1970, p. 408), and Mindanao (Hoogstraal, 1951, p. 45). In sample areas studied, the pooled ratio of sexually mature males to sexually mature females varies from 0.22 to 0.90 (Table 28); within troops, known sex ratios vary from 0.14 to 1.67. During a 10-year period of investigation at Kua- la Lompat, West Malaysia, the size of local troops tended to remain relatively constant (Chivers & Raemaekers, 1980, p. 25 1). However, troops often temporarily subdivide into smaller units while for- aging (West Malaysia [Aldrich-Blake, 1980, p. 147], Sumatra [van Schaik et al., 1983b, p. 214], Kali- mantan [Kurland, 1973, p. 253; Rodman, 1978, p. 470; Wheatley, 1978, p. 348]). At each of two localities in Sumatra, a large troop has been ob- served to permanently subdivide into two un- equal-sized daughter troops (van Schaik et al., 1983a, p. 174; Norikoshi, 1984, p. 6). Home Range, Day Range Reported home-range area in nonprovisioned troops varies from 1 2.5 ha in P. Simeulue to more than 300 ha in West Malaysia (Table 29). Overlap between home ranges of adjacent troops generally is slight, but the home range of a daughter troop may overlap extensively with that of the troop from which it split (van Schaik & van Noordwijk, 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- jacent troops are separated by rivers (van Noord- wijk & van Schaik, 1 985, p. 850), the home ranges of a pair of adjacent troops in Borneo overlapped on both sides of a river (Fittinghoff & Lindburg, 1980, p. 207). FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 51 Table 26. Recorded habitats of Macaca fascicular is. Sample area Lowland forest Upland 1 forest _, , . ( nlti- Popu- lated Man- Other Riv- Sec- Sec- vated area; Refer- grove coastal erine ondary Primary ondary Primary area temple ence 2 Core area X 7 X X X X 2 X X X X X X X X X 3 4 X X X X X X 5 X X X X X X X X 6 X X X X X XXX X 7 Bangladesh Burma Thailand, > 10°N Vietnam Thailand, < 10°N West Malaysia Sumatra Borneo Java Mergui Archipelago Con Son Tioman Singapore Riau-Lingga Archipelago 3 Kepulauan Natuna 4 Banggi group 5 Bali Nicobar Islands 6 Simeulue Nias Lesser Sundas 7 Philippines Shallow-water fringing islands Deep-water fringing islands 10 11 12 13 14 15 16 17 18 19 20 21 22 1 Altitude > 500 m. 2 Key to references: 1, M. A. R. Khan & Wahab, 1983, p. 102. 2, Heifer, 1838, p. 858; Phayre in Blyth, 1844, p. 475; Mason, 1851, p. 220; Tickell, 1854-1875, p. [17]; Shortridge in Wroughton, 1915, p. 700, 3, Gairdner, 1914, p. 28; Kloss, 1916b, p. 32; Gyldenstolpe, 1917b, p. 6; Fooden, 1971, p. 15; Lekagul & McNeely, 1977, p. 293; Eudey, 1980, p. 75; Aggimarangsee, 1992, p. 119. 4, Van Peenen et al., 1971, p. 134; Lippold, 1977, p. 521. 5, Robinson in Bonhote, 1903, p. 4; Fooden, [1975], p. 100; Boonratana, 1988, pp. 73 fF.; Aggimarangsee, 1992, p. 137; Eudey, 1994, p. 273. 6, Kelsall, 1894b, p. 16; Rower, 1900, p. 316; Shebbeare, 1940, p. 57; Tweedie & Harrison, 1954, p. 16; Furuya, 1965, p. 287; Bernstein, 1967, p. 198; Harrison, 1969, p. 176; Medway, 1972b, p. 120; Southwick & Cadigan, 1972, p. 8; Olivers & Davies, 1979, pp. 19-20; Lim & Sasekumar, 1979, p. 106; J. R. MacKinnon & MacKinnon, 1980, pp. 176, 186; Mah & Aldrich-Blake, 1980, p. 354; J. R. Marsh & Wilson, 1981, p. 232; Johns, 1986b, p. 207. 7, Snelleman, 1887, p. 10; Hagen, 1890, p. 80; Miller, 1902a, p. 158; Schneider, 1905, p. 72; Moszkowski, 1909, p. 24; Volz, 1912, p. 88; H. C. Robinson & Kloss, 19 18, p. 6; F. A. Ulmer, Jr., in Miller, 1942, p. 129; Wilson & Wilson, 1973, p. 5; Rijksen, 1978, p. Ill; van Schaik & van Noordwijk, 1985a, p. 141; Ghiglieri, 1986, p. 108; Bismark, 1992, p. 13; Yanuar & Sugardjito, 1993, p. 34. 8, Wallace, 1869, pp. 82, 326; Hose, 1893, p. 8; J. Buttikofer in Jentink, 1897, p. 39; Hornaday, 1910, p. 358; Shelford, 1916, p. 10; Mjoberg, 1930, p. 25; Allen & Coolidge, 1940, p. 147; Burgess, 1961, p. 146; Davis, 1962, p. 57; Yoshiba, 1964, p. 25; Kawabe & Mano, 1972, p. 216; Kurland, 1973, p. 247; Banks, 1978, pp. 166 ff.; Mittermeier, 1980, p. 252; Joines, 1981, p. 9; Chivers & Burton, [1991], p. 140; Rodman, 1991, p. 364; Cannon & Leighton, 1994, p. 509, 9, J. J. Menden, 3 Jan.-8 Apr. 1933, amnh 101811, 102015, 102017-102022 (Linggajati; Ciremay); Hoogerwerf, 1970, p. 408; Bismark, 1992, p. 13. 10, Carpenter, 1887, p. 53. 11, Van Peenen et al., 1970, p. 421. 12, Medway, 1966, p. 16; D. W. Lee, 1977, p. 21. 75, Ridley, 1895, p. 24; Chasen, 1924a, p. 78 (cf. zrc 4-089); Chuang, 1973, p. 3; Lucas & Corlett, 1991, p. 203. 14, W. L Abbott, 23 Jul. 1899, usnm 101603 (P. Lingga); Abbott in Miller, 1906c, p. 281. 75, A. Everett in Thomas & Hartert, 1894, p. 654. 16, Shukor Md. Nor, pers. comm., 1 1-20 Jul. 1991. 17, Anonymous, 1931, p. 457; Angst, 1975, p. 372; Wheatley, 1988, p. 517; Wheatley & Harya Putra, 1994, p. 247. 75, W. L Abbott, 25-27 Feb., 23 Mar. 1901, usnm 1 1 1795-1 1 1797, 1 1 1799 (field catalog); Kloss, 1903a, pp. 1 14, 150; Abdulali, 1967, p. 143; Kalra, 1980, p. 50; Das & Ghosal, 1977, p. 265. 19, W. L. Abbott, 18 Nov. 1901, usnm 1 14162 (field catalog); van Schaik & van Noordwijk, 1985a, p. 140; Sugardjito et al., 1989, p. 201. 20, F. A. Ulmer, Jr., in Miller, 1942, p. 129. 21, Mertens, 1936, pp. 318-320; Pfeffer, 1959, p. 198; Auffenberg, 1981, p. 242; Kitchener et al., 1990, p. 98. 22, Cf. Fooden, 1991, p. 20. 3 Information available for P. Sugi and P. Lingga. 4 Information available for P. Serasan and P. Natuna Besar. 52 FIELDIANA: ZOOLOGY Table 27. Recorded size of nonprovisioned troops of Macaca fascicular is. Troop size Number of troops Sample area Mean Minimum Maximum Reference 1 Core area Bangladesh 16.0 9 29 6 1 Burma ca. 15 5 20 ? 2 Thailand ca. 25 < 10 ca. 100 16 3 Vietnam ca. 5 3 ca. 10 ? 4 West Malaysia ca. 25 14 ca. 70 > 8 5 Sumatra ca. 20 6 ca. 65 63 6 Borneo < 20 < 10 30 > 33 7 Java ca. 47 39 Deep-water fringing islands 58 5 8 Little Nicobar I. ca. 25 ca. 20 ca. 30 ? 9 Great Nicobar I. ca. 18 6 30 6 10 Simeulue 12.5 10 15 10 11 Nias ca. 12 ? ? ? 12 Flores 15 8 32 4 13 Philippines ca. 18 ca. 5 ca. 50 > 7 14 Introduced populations Mauritius 77.5 66 89 2 15 Angaur ca. 15 ca. 10 ca. 20 9 16 1 Key to references: 1, M. A. R. Khan & Wahab, 1983, p. 104. 2, Tickell, 1854-1875, p. [17]; Yin, 1967, p. 9. 3, Fooden, 1971, pp. 15, 25; [1975], p. 100; Lekagul & McNeely, 1977, p. 293; Eudey, 1980, pp. 75-76. 4, Van Peenen et al., 1971, p. 134; Lippold, 1977, p. 521. 5, Furuya, 1965, p. 291; Bernstein, 1967, p. 199; Aldrich-Blake, 1980, p. 149; J. R. MacKinnon & MacKinnon, 1980, p. 168; Johns, 1986a, p. 685; cf. Flower, 1900, p. 316. 6, Snelleman, 1887, p. 10; F. Kurt & W. Sinaga in Angst, 1975, p. 332; Crockett & Wilson, 1980, p. 157; van Schaik & van Noordwijk, 1986, p. 297; Ruiter, 1993, p. 90. 7, Davis, 1962, p. 58; Yoshiba, 1964, p. 25; Kurland, 1973, p. 251; Macdonald, 1982, pp. 63, 71; Chivers & Burton, [1991], p. 141; A. Suzuki, 1991, p. 53. 8, Hoogerwerf, 1970, p. 408; Angst, 1975, p. 337. W. L. Abbott, 27 Feb. 1901, usnm 111797 (field tag). 10, Das & Ghosal, 1977, p. 266. 11, van Schaik & van Noordwijk, 1985a, p. 142. 12, F. A. Ulmer, Jr., in Miller, 1942, p. 129. 13, Auffenberg, 1981, p. 242. 14, Fooden, 1991, p. 18. 15, Sussman & Tattersall, 1986, p. 36. 16, Matsubayashi et al., 1987, p. 84. Reported mean day-range length varies from 325 m to 1,900 m (Table 29). In Sumatra, day- range length is positively correlated with troop size (van Schaik et al., 1983a, p. 176). Following a severe forest fire in Kalimantan, the mean day- range length in one troop declined from about 1 ,450 m to 1,300 m (Berenstain, 1986, p. 260). Density In a broad survey of M. fascicularis in Sumatra conducted in 1971-1973, reported density varied from 0.60 troops/km 2 (11.2 individuals/km 2 ) in hill scrub-grassland to 7.71 troops/km 2 (143.4 in- dividuals/km 2 ) in Rhizophora mangrove swamp (Crockett & Wilson, 1980, p. 160); overall mean density was 2.98 troops/km 2 (55 individuals/km 2 ) in 1 1 1 .45 km 2 surveyed. Roughly similar densities have been recorded in Bangladesh (M. A. R. Khan & Wahab, 1983, p. 104), West Malaysia (Chivers & Davies, 1979, p. 17; J. R. MacKinnon & MacKinnon, 1980, p. 168; Marsh & Wilson, 1981, p. 232; cf. Southwick & Cadigan, 1972, p. 8), Su- matra (J. R. MacKinnon, 1973, p. 240; Rijksen, 1978, p. 122; van Schaik & van Noordwijk, 1985a, p. 142; Bismark, 1992, p. 14; Yanuar & Sugardjito, 1993, p. 35), Kalimantan (Kurland, 1973, p. 251; Wilson & Wilson, 1975, p. 254; Wheatley, 1982, p. 205; cf. Rodman, 1978, p. 472; A. Suzuki, 1991, p. 53), and Java (Bismark, 1992, p. 14). Investigators who have studied the density of this species in primary and secondary forest 5 Information available for P. Balambangan, P. Banggi, P. Malawali, and P. Maliangin Besar. 6 Information available for Katchall I., Little Nicobar I., and Great Nicobar I. 7 Information available for P. Lombok, P. Sumbawa, P. Rintja, and P. Flores. FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 53 Table 28. Ratio of sexually mature males to sexually mature females reported in nonprovisioned troops of Macaca fascicularis. Number of Number of troops sexually mature individuals Troop sex ratios Pooled sex ratio Sample area Minimum Maximum Reference 1 Bangladesh 6 55 0.17 0.25 0.22 7 Burma ? ? 0.20 0.25 0.22 2 West Malaysia 1 13 — — 0.62 3 Sumatra 7 116 0.69 1.67 0.90 4 Borneo 6 44 0.44 1.00 0.57 5 Java 4 99 0.62 1.00 0.83 6 Simeulue 10 85 0.14 1.00 0.55 7 1 Key to references: 1, M. A. R. Khan & Wahab, 1983, p. 104. 2, Tickell, 1854-1875, p. [17]. 3, Aldrich-Blake, 1980, p. 149. 4, van Noordwijk & van Schaik, 1985, p. 850; Ruiter, 1993, pp. 98, 99. 5, Kurland, 1973, p. 251; Macdonald, 1982, p. 63; Cannon & Leighton, 1994, p. 509. 6, Angst, 1975, p. 337. 7, Sugardjito et al., 1989, p. 202. agree that the density is greater— by a factor of 1.3-9.8— in secondary forest. In P. Simeulue, a deep-water island west of Sumatra, density (> 100 individuals/km 2 ) reportedly is about twice that in comparable habitats in Sumatra (van Schaik & vanNoordwijk, 1985a, p. 142). Density reportedly also is high (> 400 individuals/km 2 ) in P. Pe- utjang, a small shallow- water island off the western tip of Java (Angst, 1975, p. 329). In provisioned troops at Ubud, Bali, density is about 1,600 in- dividuals/km 2 (Wheatley, 1989, p. 65). Based on rough estimates of population density and remaining available habitat, K. S. MacKinnon (1986, p. Ill) calculated provisionally that the population of M. fascicularis in Indonesia at that time was 3,726,860, and J. R. MacKinnon and MacKinnon (1 987, p. 1 89) calculated that the pop- ulation in mainland Southeast Asia north of West Malaysia was 309,360. If these provisional cal- culations are reasonable, the total population of this species about 1 years ago in its entire natural range— which, in addition to Indonesia and main- land Southeast Asia north of West Malaysia, in- cludes the Nicobar Islands, Malaysia, Brunei, and the Philippines-may have been approximately 5 million. Diet Fruits, of at least 185 species (Lucas & Corlett, 1991, p. 205; 1992, p. 45; cf. Ungar, 1994, p. 217; 1995, p. 232), apparently are the main food of M. fascicularis (Table 30). Ingestion of fruit frequent- ly (ca. 57% of observations) involves use of the incisors (Ungar, 1994, p. 210). When M. fasci- cularis feeds on fruit, seeds that are more than about 4 mm wide are separated from the sur- rounding pulp and rejected, either manually or by spitting, whereas smaller seeds usually are swal- lowed and defecated intact (Corlett & Lucas, 1 990, p. 167). Macaca fascicularis also consumes leaves and other plant parts, invertebrates, and small verte- brates. Widely reported invertebrate prey includes crustaceans, bivalves, and snails, along the sea- coast and riverbanks, and insects inland. Oysters sometimes are broken open by smashing with a stone that a monkey may transport up to 75 m for this purpose (Carpenter, 1887, p. 53). The capture and consumption of an adult sparrow by a captive M. fascicularis in the Calcutta Zoo is described by Mandal (1990, p. 435). This species often raids cultivated crops (Kloss, 1 903a, p. 1 28; Miller, 1 903a, p. 438; Das & Ghos- al, 1977, p. 266; Wilson & Wilson, 1977, p. 211). In Sumba, it has learned to eat the fruit of Opuntia elatior, an introduced American cactus (Dam- merman, 1928, p. 301). Relative proportions of various components of the natural diet vary seasonally and in response to severe environmental perturbations, such as forest fires (Aldrich-Blake, 1980, p. 160; Beren- stain, 1986, p. 258). These proportions also vary according to the monkey's age and sex (van Schaik & van Noordwijk, 1986, p. 305). Feeding is esti- mated to occupy 1 3-5 5% of the daily waking hours of M. fascicularis (Table 31). Predators Known natural predators on M. fascicularis in- clude crocodiles (^2 species), the Komodo giant monitor, the python, the Philippine eagle, leop- 54 FIELDIANA: ZOOLOGY Table 29. Estimated home range and day range in troops of Macaca fascicularis. Home range Day range Number Sample area Habitat (ha) (m) of troops Reference 1 Nonprovisioned troops Bangladesh Mangrove 80 325 6 / West Malaysia Mangrove 200 — 1 2 Lowland forest 46.2 1,400 ? 3 Lowland forest 41 1,062 1 4 Lowland forest 14 100 5 5 Lowland forest > 300 — ? 5 Sumatra Mangrove >25 — ? 6 Nonmangrove 75 — ? 6 Lowland forest 50 1,500 7 7 Borneo Lowland forest 80 700 1 8 Lowland forest 125 1,900 1 9 Lowland forest 112.5 1,450 2 1 10 Lowland forest 60 — 11 11 Simeulue Lowland forest 12.5 Semiprovisioned troops ~ < 10 12 West Malaysia Mangrove 35 — 1 13 Singapore Lowland forest 33 Urban troop 1 14 West Malaysia Golf course 7.2 1,009 1 4 1 Key to references: /, M. A. R. Khan & Wahab, 1983, pp. 104, 107. 2, Furuya, 1965, p. 288. 3, J. R. MacKinnon & MacKinnon, 1980, p. 168. 4, Mah & Aldrich-Blake, 1980, pp. 355-356. 5, Caldecott, 1986a, p. 155. 6, Crockett & Wilson, 1980, p. 168. 7, van Schaik et al., 1983a, p. 176; 1983b, p. 213. 8, Kurland, 1973, p. 252. 9, Wheatley, 1980, p. 233. 10, Berenstain, 1986, pp. 257, 260. 11, A. Suzuki, 1991, p. 53. 12, van Schaik & van Noordwijk, 1985a, p. 141; Sugardjito et al., 1989, p. 201. 13, Lim & Sasekumar, 1979, p. 107. 14, Lucas & Corlett, 1991, p. 203. 2 Mean for 1 1 mo prior to a major forest fire; the mean for 4 mo after the fire is 1,300 m. ards (2 species), and the tiger. Crocodiles (Croc- odylus sp., Tomistoma schlegeli) have been ob- served preying on M. fascicularis in Thailand (Mouhot, 1864, p. 152), Sumatra (Volz, 1912, p. 369), and Borneo (Shelford, 1916, p. 10; Mjoberg, 1930, p. 25; Galdikas & Yeager, 1984, p. 50). The Komodo giant monitor (Varanus komodoensis) is known to prey on M. fascicularis in Flores and nearby P. Rintja (Hoogerwerf, 1955, p. 26; Dar- evsky & Kadarsan, 1964, p. 1358; Pfeffer, 1959, p. 231; Auffenberg, 1981, pp. 228, 243; Sumardja, 1981, p. 4). A python {Python sp.) was observed to take a juvenile M. fascicularis in northern Su- matra (van Schaik et al., 1983b, p. 220). The Phil- ippine eagle (Pithecophaga jefferi) has been ob- served preying on monkeys in Samar and Min- danao (Fooden, 1991, p. 18). In western Java, leopards (Panthera pardus) are reported to prey on M. fascicularis in Ujung Kulon National Park and at Gunung Pangrango (Bartels, 1929, p. 81; Hoogerwerf, 1970, p. 402), and in eastern Java, leopards and tigers (P. tigris) are reported to prey on this monkey in Meru-Betiri Reserve (Seiden- sticker, 1983, p. 324); clouded leopards (Neofelis nebulosa) are reported to prey on M. fascicularis in Borneo (Banks, 1931, p. 77). Domestic dogs also attack and sometimes kill M. fascicularis (Ridley, 1895, p. 25; Seidensticker, 1983, p. 325), and humans are known to hunt this species for food (Labang & Medway, 1979, p. 56; Aggimar- angsee, 1992, p. 154). Under experimental conditions, hi. fascicularis exhibited fear of a stuffed python and a snake model (van Schaik & Mitrasetia, 1990, p. 105; Vitale et al., 1991, p. 281). In nature, M. fascicularis has been observed to mob a python (van Schaik & Mitrasetia, 1990, p. 106) and to engage in antag- onistic behavior directed toward crocodiles, Ko- modo giant monitors, and domestic dogs (Mou- hot, 1864, vol. 1, p. 152; Hagen, 1890, p. 81; Auffenberg, 1981, p. 243; Wheatley, 1991, p. 172). Field studies indicate that predator detection is a major determinant of social behavior and group size in M. fascicularis (van Schaik et al., 1983b, p. 220; van Schaik & van Noordwijk, 1985a, p. 139). FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 55 Table 30. Foods reported eaten by nonprovisioned troops of Macacafascicularis; dietary proportions are indicated where available. Foods eaten Crustaceans, Other Fruit, Other Insects, mollusks, animal Refer- Sample area Leaves seeds plant parts spiders fish prey ence 1 Core area Bangladesh 20% 4% 25% 51% ; Burma X X X X 2 Thailand X X X 3 West Malaysia X X X X X X 2 4 16.1% 52.4% 8.3% 23.3% 5 24.0% 63.7% 8.8% 4.4% X 3 6 Sumatra X X X X X 7 4.4% 63.3% 18.1% 8 4 Borneo X X X X X X' 9 8% 87% 3.5% 1% 10 Shallow-water fringing islands Mergui Archipelago X X 11 Singapore X X X 12 Banggi X 13 Deep-water fringing islands Nicobar Islands X X 14 Lesser Sundas X X X X 15 Philippines X X X* 16 Introduced population Mauritus 9% 70% 14% 5% 7 17* 1 Key to references: 1, M. A. R. Khan & Wahab, 1983, p. 106. 2, Heifer, 1838, p. 858; A. P. Phayre in Blyth, 1844, p. 475; Mason, 1851, p. 220; Tickell, 1854-1875, p. [17]; Blyth, 1859, p. 275; G. C. Shortridge in Wroughton, 1915, p. 700; H. C. Smith, 21-30 Apr. 1936, bm(nh) 1936.9.10.8, 9.10.10, 9.10.14 (field tags); Pocock, 1939, p. 82. 3, H. C. Robinson in Bonhote, 1903, p. 4; Gairdner, 1914, p. 28; Gyldenstolpe, 1914, p. 3; Fooden, 1971, p. 24; [1975], p. 99; McNeely, 1977, p. 10; Lekagul & McNeely, 1977, p. 293. 4, Tweedie & Harrison, 1954, p. 16; Harrison, 1961, p. 7; Furuya, 1965, p. 289; Lim & Sasekumar, 1979, p. 109 (semiprovisioned troop); Lambert, 1990, p. 455. 5, Aldrich-Blake, 1980, p. 160. 6, J. R. MacKinnon & MacKinnon, 1980, p. 178. 7, Volz, 1912, p. 369; F. A. Ulmer, Jr., in Miller, 1942, p. 127; Rijksen, 1978, p. 112; Crockett & Wilson, 1980, p. 164; Whitten et al., 1984, p. 136; van Schaik & van Noordwijk, 1988, p. 82. 8, Ungar, 1994, p. 210. 9, Shelford, 1916, p. 10; Mjoberg, 1930, p. 25; Kern, 1964, p. 185; Roedelberger & Groschoff, 1967, p. 30; Kurland, 1973, p. 254; Rodman, 1978, p. 469; Fittinghoff & Lindburg, 1980, p. 190; Wheatley, 1980, p. 216; Joines, 1981, p. 9; Macdonald, 1982, p. 63; Leighton & Leighton, 1983, p. 185; Berenstain, 1986, p. 257; 10, Wheatley, 1978, p. 348; reported diet also includes 0.5% clay. 11, Carpenter, 1887, p. 53; H. C. Smith, 20-23 Apr. 1936, bm(nh) 1936.9.10.4-6, 1936.9.10.11. 12, Ridley, 1895, p. 24; Chasen, 1924a, p. 79; Lucas & Corlett, 1991, p. 203 (semiprovisioned troop). 13, Shukor Md. Nor, pers. comm., 11 Jul. 1991. 14, Kloss, 1903a, p. 129; Das & Ghosal, 1977, p. 266; Kalra, 1980, p. 50. 75, Dammerman, 1928, p. 301; Pfeffer, 1959, p. 198; Auffenberg, 1981, p. 242. 16, Fooden, 1991, p. 18. 17, Sussman & Tattersall, 1981, p. 200. 2 Sipunculid worms; lizards. 3 Frogs. 4 14.2% of diet is unspecified. 5 Birds' eggs or nestlings. 6 Earthworms; lizards; birds. 7 Includes all invertebrates. 8 2% of diet is unspecified. Intertroop Behavior Judging from reports of long-term observations, direct encounters between adjacent nonprovi- sioned troops are relatively rare (West Malaysia [Aldrich-Blake, 1 980, p. 1 5 1], Sumatra [van Schaik & van Noordwijk, 1988, p. 80], Kalimantan [Fit- tinghoff & Lindburg, 1980, p. 206], Java [Angst, 1973, p. 626]), which suggests mutual avoidance. In Sumatra, however, small daughter troops fre- quently encounter the large troops from which they have split (van Schaik & van Noordwijk, 1988, p. 56 FIELDIANA: ZOOLOGY Table 31. Daily waking-hour 1 time budget estimates (%) for Macaca fascicularis. 2 Activity Sample area Feeding Travel Other Reference Bangladesh Ebb tide Full tide West Malaysia Kalimantan Mauritius (introduced) 55 22 35 13 32 15 3 20 45 4 30 75 45 3 42 64 M. A. R. Khan & Wahab, 1983, p. 106 Aldrich-Blake, 1980, p. 161 Wheatley, 1980, p. 221 Sussman & Tattersall, 1981, p. 197 1 Estimates of total daily waking hours: West Malaysia, 13 hr (J. R. MacKinnon & MacKinnon, 1980, p. 185); Sumatra, 12.5 hr (van Schaik & van Noordwijk, 1988, p. 85); Kalimantan, 1 1.4 hr (Wheatley, 1980, p. 222). 2 Cf. Lim and Sasekumar (1979, p. 1 1 1), J. R. MacKinnon and MacKinnon (1980, p. 185), and Leon et al. (1993, p. 177). 3 Computed by subtraction. 80), and in P. Peutjan, an islet off western Java that is densely populated by M. fascicularis (see above), adjacent troops are often intermixed (Angst, 1 973, p. 626). Intertroop displays that may function to maintain intergroup spacing have been reported in West Malaysia and Kalimantan. In P. Simeulue, west of Sumatra, M. fascicularis has a unique rapid bark that probably functions in in- tertroop communication (van Schaik & van Noordwijk, 1985a, p. 143). During intertroop en- counters in Sumatra, copulations between mem- bers of different troops have been observed infre- quently (van Noordwijk, 1985, p. 289). In Brunei, northern Borneo, a larger troop was observed to displace a smaller troop (Macdonald, 1982, p. 71). In artificially provisioned troops, intertroop fight- ing is frequent (Furuya, 1 965, p. 288; Angst, 1975, p. 373; Norikoshi, 1984, p. 1 1; Koyama, 1984, p. 25; Wheatley, 1991, p. 171). Macaca fascicularis males have been observed to leave one troop and take up residence in another troop in northern Sumatra (Rijksen, 1978, p. Ill; van Noordwijk & van Schaik, 1985, p. 85 1), west- ern Sumatra (Norikoshi, 1 984, p. 2; Koyama, 1 984, p. 20; 1985, p. 1 19), eastern Kalimantan (Wheat- ley, 1982, p. 207; Berenstain, 1986, p. 258), and Bali (Koyama et al., 1981, p. 8). Such intertroop movement by males presumably is universal in natural populations of this species, as in other spe- cies of macaques. Males apparently leave their na- tal troop as late juveniles or subadults, before about age 7 years, and most males probably change troops several more times during their life; intertroop movement apparently precludes close inbreeding in natural troops (Ruiter et al., 1992, p. 186). In six troops studied in northern Sumatra, 40 of 44 adult and subadult (immigrant) males that were present in January 1 980 had emigrated by March 1984. In western Sumatra, the mean duration of troop residence of an adult male was estimated to be 1 .2 years. Most intertroop movements by males in northern Sumatra occurred near the beginning of the peak copulatory season. The maximum re- corded interval between a male's emigration from one troop and immigration into another troop was 2 months (semisolitary interval). Immigration of a new male often is followed by changes in a troop's dominance hierarchy. Females also occasionally move from one troop to another, but apparently at a much lower frequency than males (5 of 57 recorded migrations in northern Sumatra; 1 of 7 in western Sumatra; of 1 1 in eastern Kalimantan; of 2 in Bali; total, 6 of 77 = 7A Interspecific Behavior Interactions have been reported between M. fas- cicularis and M. nemestrina, leaf monkeys (Pres- bytis spp., Trachypithecus spp.; cf. Groves, 1993, p. 270), Nasalis larvatus, Hylobates spp., and Pon- S° pygmaeus. In a 3 -km 2 study area in eastern Kalimantan, Rodman (1973, p. 657) found that the local occurrence of M. fascicularis was nega- tively correlated with that of M. nemestrina, Pres- bytis aygula (= P. hosei), Hylobates moloch, and Pongo pygmaeus. Noncompetitive associations of M. fascicularis and M. nemestrina have been observed in West Malaysia (Bernstein, 1967, p. 201), and seven mixed troops of these species have been recorded in Sumatra (Rijksen, 1978, p. Ill, single M. fas- cicularis individuals in six M. nemestrina troops; Crockett & Wilson, 1980, p. 175, M. nemestrina individual in M . fascicularis troop). These two spe- cies, however, generally occupy different ecologi- FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 57 cal niches; M. fascicularis is rare in primary ev- ergreen rain forest, which is favored by M. ne- mestrina (Rodman, 1973, p. 656; 1978, p. 476; 1991, p. 364; J. R. MacKinnon & MacKinnon, 1978, p. 316; Crockett & Wilson, 1980, p. 162). Macaca fascicularis and Trachypithecus crista- tus frequently have been observed together in West Malaysia (Furuya, 196 1-1 962b, p. 56; Bernstein, 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 species reportedly are generally peaceful, but M. fascicularis often displaces T. cristatus. Hooger- werf (1970, p. 408) observed one violent inter- specific fight in Java, during which an immature T. cristatus ". . . fell to the ground torn to pieces." Associations of M. fascicularis and Presbytis melalophos have been observed in West Malaysia (Bernstein, 1967, p. 201; J. R. MacKinnon & MacKinnon, 1978, p. 319) and Sumatra (Snelle- man, 1887, p. 10; Koyama, 1985, p. 105). Most accounts indicate that these associations are peaceful . Koyama (1985, p. 105) reported that these two species often share the same sleeping tree, but J. R. MacKinnon and MacKinnon (1978, p. 319) indicated that M. fascicularis displaces P. mela- lophos from feeding and sleeping trees. Bernstein (1967, p. 201) reported noncompeti- tive association of M. fascicularis with Trachypi- thecus obscurus in West Malaysia, and Kurland (1973, p. 254) reported noncompetitive associa- tion with Presbytis aygula (= P. hosei) in Kali- mantan. Ridley (1895, p. 25) reported that leaf monkeys in West Malaysia fought for more than 2 hours with M. fascicularis over occupation of a feeding tree ("biting them [M. fascicularis] and throwing them out of the tree and into the river"). Banks (1978, p. 188) reported that M. fascicularis often prevented leaf monkeys in Borneo from en- tering sleeping trees. Peaceful association of M. fascicularis and Nasalis larvatus in mangrove swamps in Borneo has been observed frequently (Davis, 1962, p. 57; Kern, 1964, p. 185; Kawabe & Mano, 1972, p. 216; Kurland, 1973, p. 254; Macdonald, 1982, p. 117). Noncompetitive association of M. fascicularis and Hylobates lar in West Malaysia was reported by Bernstein (1967, p. 201). Macaca fascicularis and Pongo pygmaeus have been observed together on 32 occasions in Sumatra (Rijksen, 1 978, p. 112) and on two occasions in Kalimantan (Fittinghoff & Lindburg, 1980, p. 201); these observations in- clude peaceful interspecific sharing of feeding and sleeping trees and rudimentary interspecific threats— a few by P. pygmaeus and one by M. fascicularis. In West Malaysia, racket-tailed drongos (Dicru- rus paradiseus) have been observed following M. fascicularis through the canopy and catching in- sects stirred up by the monkeys' activity (Ridley, 1901, p. 105; cf. Fooden, 1969, p. 52). Reproduction Although copulations, pregnancies, and births have been recorded throughout the year in natural populations of M. fascicularis, long-term studies indicate that all three of these reproductive events exhibit seasonal peaks in this species (Appendix 1 1). The timing of these reproductive peaks ap- parently varies geographically and varies between years at the same locality. In a study in northern Sumatra, the annual birth peak tended to occur during or shortly after the local fruiting peak; in years when fruit production was relatively low, the birth peak tended to be delayed by 2 or 3 months (van Schaik & van Noordwijk, 1985b, p. 538). In Thailand, on the other hand, fertile matings— not births— tend to peak shortly after the fruiting peak, which occurs in April-June (Varavudhi et al., 1989a, pp. 221-222; cf. Tangpraprutigul & Var- avudhi, 1982, p. xci; Aggimarangsee, 1992, p. 129). In a 2- to 3-year study of captive M. fascicularis, annual variation of sperm concentration in semen collected by electroejaculation was investigated (Okamoto, 1994, p. 27); results were inconclusive. In natural populations of M. fascicularis, fe- males probably become reproductively active at about age 3.5 years. A pregnant juvenile female, aged about 3.5 years, was collected in western Thailand (fmnh 99646, with erupting C,, P 3 , and M 2 ; cf. Table 14; Fooden, 1 97 1 , p. 24), and females in Sumatra reportedly produce their first infants at about age 4 years (Koyama, 1985, p. 106; van Noordwijk & van Schaik, 1987, p. 586; cf. Var- avudhi et al., 1989a, p. 222). In captivity, females may copulate as early as age 2.5 years (Chance et al., 1 977a, p. 6 1 9), and one captive female is known to have given birth at age 3 years (Timmermans et al., 1981, p. 121); however, fertility in captive females usually does not begin until about age 4 years (Spiegel, 1954, p. 230; Dang, 1983, p. 38). Mean age at menarche in 43 captive females was 2.5 ± 0.7 years (Honjo et al., 1984, p. 69; cf. Dang, 1983, p. 36). Mean length of the menstrual 58 FIELDIANA: ZOOLOGY cycle in 28 captive females was 30.9 days (SD = 4.8 days, extremes = 19-43 days, mode = 28 days, n = 595 cycles) (Dukelow, 1977, p. 34; cf. Joach- imovitz, 1928, p. 462; Comer, 1932, p. 404; Spie- gel, 1954, p. 235; Fujiwara et at, 1967, p. 506; Kerber & Reese, 1 969, p. 976; Valerio et al., 1 969, p. 287; MacDonald, 1 97 1 , p. 374; Nawar & Hafez, 1972, p. 45; Dang, 1977, p. 3; Goodman et al., 1977, p. 480; Zumpe & Michael, 1 983, p. 58; Hon- jo et al., 1984, p. 63; Wallis et al., 1986, p. 87). Males in natural populations usually begin to copulate at age 5-6 years, as subadults, after they have left their natal troops (van Noordwijk, 1985, p. 288; van Noordwijk & van Schaik, 1985, p. 852; cf. Varavudhi et al., 1989a, p. 222). As pre- viously indicated (see above, Natural History), close inbreeding probably is rare or absent in nat- ural populations of M. fascicularis. In Sumatra, subadult males that were still in their natal troops accounted for only 5 of 664 observed copulations. In Bangladesh, copulation attempts by immature males were actively thwarted by adult males (M. A. R. Khan & Wahab, 1983, p. 108). In captive males, testicular descent occurs at age 2.5 years, and adult testicular size is achieved at age 4.0-4.5 years (Cho et al., 1973, p. 408; Chance et al., 1 977a, p. 6 1 9). Fertile copulations have been recorded for three captive males at ages 3.5 years (1 male; Honjo et al., 1984, p. 68) and 3.8 years (2 males; Spiegel, 1954, p. 230). At or near the onset of sexual maturity in fe- males, the bare skin between the root of the tail and the anus swells prominently and the skin around the vulva frequently reddens (Joachimov- itz, 1928, p. 464; Corner, 1932, p. 408; Spiegel, 1 954, p. 232; Nawar & Hafez, 1 972, p. 49; Fooden, [1975], p. 100; Emory et al., 1980, p. 250; van Noordwijk, 1985, p. 281; Meishvili & Chalyan, 1986, p. 14; C. M. Anderson & Bielert, 1994, p. 285). This pubertal swelling and reddening ap- parently reaches its maximum about midway through the menstrual cycle and diminishes a few days later. In postpubertal females, sexual swelling and reddening generally are less conspicuous and are highly variable in size and duration; in many fully fertile females, sexual swelling and reddening are slight or absent (cf. Pocock, 1 906, p. 558; 1 939, p. 78; Kurland, 1973, p. 258; Wilson & Wilson, 1977, p. 211; Fittinghoff & Lindburg, 1980, p. 189). However, in a carefully studied population in northern Sumatra, sexual swellings generally were larger during the peak copulation season (Jan- Jun.), and copulations and consortships of most females tended to increase with increasing size of their sexual swelling or increasing redness of their perivulval skin (van Noordwijk, 1985, pp. 281, 287). In some females, sexual swelling or redden- ing and copulations were observed during preg- nancy (cf. Deputte & Goustard, 1980, p. 96); fol- lowing parturition, swelling was not observed for several months. Variably complete and generally concordant ac- counts of copulatory behavior in M. fascicularis are available from natural populations (Tickell, 1854-1875, p. [17]; Furuya, 196 1-1 962a, p. 76; Kurland, 1973, p. 258; M. A. R. Khan & Wahab, 1983, p. 104; Koyama, 1984, p. 31; 1985, p. 107; van Noordwijk, 1985, p. 281), introduced free- ranging populations (Poirier & Smith, 1974, p. 300; Sussman & Tattersall, 1981, p. 203), colony groups (Spiegel, 1954, p. 238; Goustard, 1961, p. 313; 1963, p. 710; 1968, p. 464; Furuya, 1965, p. 313; de Benedictis, 1973, p. 1470; Angst, 1974, p. 52; Chevalier-Skolnikoff, 1975, p. 207; Emory & Harris, 1978, p. 227; Deputte & Goustard, 1980, p. 85; Emory et al., 1980, p. 251; Shively et al., 1982, p. 375), and laboratory test pairs (Kanagawa et al., 1972, p. 453; Kanagawa & Hafez, 1973, p. 234; Michael & Zumpe, 1988, p. 379; Zumpe & Michael, 1983, p. 58; 1990, p. 148). Copulations in this species have been observed throughout the day, but evidence from troops in northern Su- matra suggests that copulations may be more fre- quent in the morning (ca. 60%) than in the after- noon (van Noordwijk, 1985, p. 286; cf. de Bene- dictis, 1973, p. 1473; Sussman & Tattersall, 1981, p. 203). Copulations occur both in trees and on the ground (Kurland, 1973, p. 258; Angst, 1974, p. 53; Sussman & Tattersall, 1981, p. 203; Koy- ama, 1985, p. 117). Copulations in M. fascicularis may be initiated by either sex. A female initiates copulation by star- ing at or approaching a male and presenting her hindquarters or by stereotyped head or arm move- ments; in laboratory test pairs, female use of head or arm movements to initiate copulation peaked near the middle of the menstrual cycle (Michael & Zumpe, 1988, p. 381). A male initiates copu- lation by staring at or approaching a female and raising her tail. Both female- and male-initiated copulations usually proceed with examination by the male of the perineum of the female; this ex- amination may be visual, digital, olfactory, or, rarely, lingual. If the female is acceptable (cf. de Benedictis, 1973, pp. 1479-1480), the male mounts the female, by gripping her hips with his hands and gripping her calves with his feet, and begins 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) SME copulations (N) Duration of SME mount 1 (sec) Thrusts per SME mount 1 MME copulations (N) Duration of MME mount 1 (sec) Thrusts per MME mount 1 Duration of MME sequence 1 (min) Mounts per MME sequence 1 Reference 169 139(82.2%) 7.1 (2.6-12.0) 16.4(5-32) 30(17.8%) 6.4 (3.0-9.4) 15(6-31) 7.1 (2-35) 2.4 (2-6) Wheatley, 1991, p. 171 17 8(47.1%) 8.5 (6-14) 17(14-21) 9 (52.9%) 6(2-11) 13(4-29) 41 (2-85) 3(2-6) Shively et al., 1982, p. 376 Mean and extremes. and mounting attempts tended to increase slightly during the first half of the menstrual cycle and tended to decrease more precipitously during the second half (Zumpe & Michael, 1983, p. 66). Ejaculation in M. fascicularis may occur either as a result of a single mount (single-mount ejac- ulations [SME]) or as a result of a series of mounts separated by dismounts (multiple-mount ejacu- lation [MME]). In a provisioned natural group in Bali, 82% of observed ejaculatory copulations were SME and 1 8% were MME; in a seminatural colony group, 47% of observed ejaculatory copulations were SME and 53% were MME (Table 32; cf. Fu- ruya, 196 1-1 962a, p. 76; van Noordwijk, 1985, p. 281). In laboratory tests, the frequency of SME was higher in first copulations of test pairs than in second copulations (27% of 191 first copula- tions, 6% of 154 second copulations) (Zumpe & Michael, 1983, p. 60; cf. Kanagawa et al., 1972, p. 453). In the natural and colony groups, the du- ration of both SME and MME mounts averaged about 7 seconds, and the number of pelvic thrusts per mount averaged about 15 (cf. Furuya, 1965, p. 313; Zumpe & Michael, 1983, p. 61); similar mount durations and thrust/mount rates have been reported for natural populations of M '. fascicularis in Bangladesh (10-15 seconds, 8-10 thrusts/mount; M. A. R. Khan & Wahab, 1983, p. 108) and Ka- limantan (5-7 sec, 4-12 thrusts/mount; Kurland, 1973, p. 258) and for a free-ranging introduced population in Angaur Island ("brief," 8-10 thrusts/ mount; Poirier & Smith, 1974, p. 300). The num- ber of mounts in an MME sequence averaged about 3 in the natural and colony groups; the duration of an MME sequence averaged about 7 minutes in the natural group and about 4 1 minutes in the colony group. Copulation in M. fascicularis usually is accom- panied by a characteristic staccato female vocal- ization and sometimes by a characteristic male vocalization (Goustard, 1963, p. 710; 1968, p. 470; Furuya, 1965, p. 313; Angst, 1974, p. 61; Poirier & Smith, 1974, p. 300; Chevalier-Skolnikoff, 1975, p. 207; Deputte & Goustard, 1980, p. 83; Wheat- ley, 1 982, p. 206; 1 984, p. 391; 1991, p. 171; Zumpe & Michael, 1983, p. 60; van Noordwijk, 1985, p. 281). At or near the time of ejaculation, the male pauses, and the female usually grasps one of the male's arms or legs and turns her face toward his (cf. Kanagawa et al., 1972, p. 453). Shortly after ejaculation, the male dismounts and the male and female move apart. After copulation, a pair may remain associated in a consortship for a variable period, ranging from about 1 hour to 3 weeks (Fu- ruya, 1965, p. 314; Poirier & Smith, 1974, p. 301; Sussman & Tattersall, 1981, p. 203; van Noord- wijk, 1985, p. 283); during a consortship, most, but not all, copulations of consortship partners are with each other. In natural groups, females may copulate as often as 3-4 times per hour, through- out the day, often with many different partners (Ruiter et al., 1992, p. 177). Dominant males in M. fascicularis apparently participate in a disproportionately large share of copulations. In northern Sumatra, the highest ranking 2 of 7 males engaged in 52.7% of 659 copulations (van Noordwijk, 1985, p. 285); in western Sumatra, 2 of 7 males engaged in 60.9% of 46 ejaculatory copulations (Koyama, 1984, p. 35; cf. Norikoshi, 1984, p. 12); in eastern Kali- mantan, 1 of 6 males engaged in 50.0% of 48 cop- ulations (Wheatley, 1982, p. 207; alpha status shifted during this study); and in Bali, 1 of 4 males engaged in 50.9% of 126 copulations (Wheatley, 1991, p. 171). Similar high frequencies of copu- lation by dominant males have been reported in 60 FIELDIANA: ZOOLOGY colony groups (Furuya, 1965, p. 315; Shively et al., 1982, p. 377). In natural and colony groups, some low-ranking adult males were never ob- served to copulate. In laboratory test pairs, the copulatory performance of a subordinate male was strongly inhibited by close proximity of a sepa- rately confined dominant male (Zumpe & Mi- chael, 1990, p. 154). In West Malaysia and Kali- mantan, aggression between males reportedly in- tensified shortly before periods of increased sexual activity (Aldrich-Blake, 1980, p. 151; Wheatley, 1982, p. 210). In troops observed in northern and western Sumatra, subordinate males frequently at- tempted to interfere with the copulations of dom- inant males, who, in turn, responded aggressively (Norikoshi, 1984, pp. 9, 12; Koyama, 1985, p. 107; van Noordwijk, 1985, p. 283). The copula- tory partners of high-ranking males in Sumatra generally tended to be females who were high rank- ing, not young, and not nulliparous (van Noord- wijk, 1985, p. 289). Paternity tests, using blood-protein analysis and DNA fingerprinting, have been conducted for 45 offspring in three troops in northern Sumatra (Rui- ter et al., 1992, p. 184; cf. Ruiter, 1993, p. 99; Ruiter & van Hooff, 1993, p. 518; Ruiter et al., 1994, p. 211); these tests indicate that dominant males sire most of the offspring in natural troops, as would be expected from observed copulation frequencies. Of 42 offspring for which probable fathers could be determined, 31 probably were sired by alpha males, 7 by beta males, and only 4 by approximately 10 lower ranking males. In ar- tificial colony groups, dominant males apparently do not maintain this reproductive advantage; pa- ternity tests in one colony revealed that, despite their higher frequency of copulations, dominant males did not sire a disproportionately large num- ber of offspring (Shively & Smith, 1985, p. 131). In a large captive group of M. fascicularis, mean gestation length for 1,141 live births was 163.5 ± 5.8 (SD) days (Honjo et al., 1984, pp. 63-64); in other captive groups, reported mean gestation lengths vary from 160.0 to 168.0 days (extremes 141-225 days) (Spiegel, 1954, p. 245; T. Fujiwara & Imamichi, 1966, p. 226; Valerio et al., 1969, p. 295; MacDonald, 1971, p. 374; Dang, 1977, p. 2; 1983, p. 41; Dukelow et al., 1979, p. 44; Vara- vudhi et al., 1989a, p. 222). Mean birth weight in the large captive group was 318.2±45.2gfor563 female infants and 347.5 ± 55.2 g for 600 male infants. The ratio of male infants to female infants tends to be greater for high-ranking captive fe- males than for low-ranking females (van Schaik et al., 1989, p. 151). Most births probably occur be- tween sunset and sunrise; in a captive group main- tained in a 14-hour light/ 10-hour dark environ- ment, 90% of 1 52 observed births occurred during dark hours, 1900-0500 (M. T. Suzuki et al., 1990, p. 252; cf. Ridley, 1906, p. 142; Spiegel, 1954, p. 248; Erwin, 1977, p. 358; Banks, 1978, p. 188; Kemps & Timmermans, 1982, p. 84). In captivity, 23 infants were nursed for an av- erage of 15.1 months (SD = 3.0 mo, extremes = 9-22 mo) (Spiegel, 1954, p. 262; cf. Chance et al., 1977b, p. 31); following 9 of these 23 nursing pe- riods, the mother's lactation continued without interruption through the nursing of one or more succeeding infants. In natural populations, infants begin to obtain some of their food independently about age 3 months (Karssemeijer et al., 1990, p. 288). The mean duration of postpartum amen- orrhea in captive nursing mothers was 6.6 months (SD = 2.6 mo, extremes = 2-13 n = 28) (Spiegel, 1954, p. 262; cf. Dang, 1979, p. 377; Cho, 1981, p. 254; Honjo et al., 1984, p. 66; Varavudhi et al., 1989a, p. 221). In northern Sumatra, in six to seven nonpro- visioned troops that included 42-53 adult females, the annual birth rate (births/adult females) from 1980-1981 to 1983-1984 was 0.45, 0.82. 0.30, and 0.75; the overall annual birth rate for these 4 years was 0.58 (van Schaik & van Noordwijk, 1985b, p. 538). In western Sumatra, in three pro- visioned troops that included 22-3 1 adult females, the annual birth rate from 1980 to 1984 was 0.73, 0.48, 0.8 1 , 0. 1 1 , and 1 .00, with an overall average of 0.63 (Koyama, 1985, p. 119). High and low annual birth rates tend to occur in alternate years in both the nonprovisioned and provisioned troops. The limited available data suggest that a biennial birth cycle may be common in these Sumatran troops and that most females may tend to produce their young during the same alternate years. In provisioned troops at Ubud, Bali, where the number of adult females varied from 27 to 46, the annual birth rate reported in 1986 and 1990-1992 was 0.59, 0.32, 0.41, and 0.33 (Wheatley & Harya Putra, 1994, p. 248). In four free-ranging provi- sioned groups of M. fascicularis introduced in P. Tinjil, south of Java, the annual birth rate is es- timated to be 0.56 (Kyes, 1 993, p. 8 1). In captivity, M. fascicularis females are capable of producing young every year (Ridley, 1895, p. 24; Chance et al., 1977a, p. 612; cf. Hadidian & Bernstein, 1979, p. 440; Timmermans et al., 1981, p. 120; Dang, 1983, p. 41; Honjo et al., 1984, p. 66). 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. Number of fossils or Estimated subfossils reported M. fasci- M. nemes- Macaca Refer- Locality Epoch age (Ka) cularis trina sp. ence 1 West Malaysia Bukit Chintamani Holocene ? 1 1 Kota Tongkat Holocene < 4 1 1 Gua Cha, Trench 79 A Holocene ? > 1 2 Gua Cha, Trench 79B Holocene < 3 > 1 2 Gua Cha, Trench 79B Holocene 6.3 > 1 2 Gua Madu Holocene Sumatra 7 > 1 3 Djamboe Cave Holocene/Pleistocene 2 ? 11 4 Lida Ajer Cave Holocene/Pleistocene 2 ? 1 6 1 4 Sibrambang Cave Holocene/Pleistocene 2 ? 5 74 10 4 Unspecified cave Holocene/Pleistocene 2 ? 1 11 4 4 Borneo: Sarawak Paku Cave ?Holocene ? 4 3 5 Unspecified cave ?Holocene ? l 3 5 Niah Cave Holocene 4 5 21 4 6 Niah Cave Pleistocene 4 15 37 6 Niah Cave Pleistocene 4 25 7 6 Niah Cave Pleistocene 4 Java 35 5 1 6 Sampung Holocene 5 5 3 7 Goea Djimbe Holocene 10 26 4 Goea Ketjil Holocene 10 1 4 Cave near Wajak Holocene 10 l 6 4 Punung Pleistocene 120 2 16 8 Ngandong Pleistocene 7 < 800 l 8 9 Sangiran (?Kabuh) Pleistocene 7 800 1 1 10 Sangiran Pleistocene 7 ?800 2 11 Glagahombo (Kabuh) Pleistocene 7 800 1 12 Kali Brangkal (Kabuh) Pleistocene 7 800 1 12 Ndangklampok (Kabuh) Pleistocene 7 800 1 12 Saradan Pleistocene 7 ?800 1 13 Trinil Pleistocene 7 1,000 7 4 Trinil Pleistocene 7 1,000 1 14 Bangle Pleistocene 7 Flores ? 1,000 2 4 Liang Toge Holocene Timor 10 3.5 9 1 15 Bui Ceri Uato Holocene < 1.5 2 16 Uai Bobo 1 Holocene 1.6 2 16 Lie Siri Holocene <4.3 l 3 16 Uai Bobo 2 Holocene 4.5 4 16 1 Key to references: 1, Matthews, 1961, pp. 18, 41; Dunn, 1975, p. 122. 2, Adi, 1985, pp. 66-67. 3, Tweedie, 1940, p. 7. 4, Hooijer, 1962b, pp. 50, 54-55, 58. 5, P. H. Napier, 1981, pp. 9, 20. 6, Hooijer, 1962a, p. 440. 7, Dammerman, 1934, p. 492. 8, Badoux, 1959, p. 88; J. de Vos, 1983, p. 421; J. de Vos et al., 1994, pp. 132, 134. 9, Aziz, 1989, p. 52. 10, Aimi, 1981, p. 409. 11, Hooijer, 1964, p. 76. 12, Aimi & Aziz, 1985, pp. 161-163. 13, Deninger, 1910, p. 1; Hooijer, 1962b, p. 54; Aimi, 1981, p. 412. 14, Stremme, 1911, p. 140; Hooijer, 1962b, p. 54. 15, Hooijer, 1967, p. 160. 16, Glover, 1986, pp. 78, 121, 158, 192. 2 See J. de Vos (1983, p. 422); J. de Vos et al. (1994, p. 132). 3 Minimum number of individuals; species identification tentative. 4 For tabulation details, see Fooden (1975, p. 61). 5 See Bell wood (1985, p. 200). 6 Cf. Brink (1982, p. 180). 62 FIELDIANA: ZOOLOGY about age 20 years (Spiegel, 1954, p. 265; Angst & Thommen, 1977, p. 212; van Noordwijk & van Schaik, 1987, p. 587), which suggests that an av- erage female in a natural population may produce about eight to nine infants during her reproductive life (age ca. 4—20 yr). The maximum known life span in captive M. fascicular is is 37 years 1 month (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 to 1983-1984 was 0.21, 0.12, 0.25, and 0.06, with an overall average of 0.14 (van Schaik & van Noordwijk, 1985b, p. 538); neonatal infant mor- tality was high in years when the birth rate was low (see above). The average infant death rate be- fore age 1 year was 0.22 in northern Sumatra (van Noordwijk & van Schaik, 1 987, p. 585); this infant death rate is comparable to that reported in a pro- visioned free-ranging group at Ubud, Bali (0.25; Wheatley & Harya Putra, 1994, p. 248), and in a large captive group of M. fascicularis (> 0. 1 5; Lu- der, 1993, p. 142). Infanticide, committed by adult males, probably is one of the important causes of infant deaths (cf. Thompson, 1967, p. 18; Wash- burn & Hamburg, 1968, p. 473; Angst & Thom- men, 1977, p. 208; Erwin, 1977, p. 359; Tim- mermans et al., 1981, p. 120; Wheatley, 1982, p. 211; Pallaud, 1984, p. 92; Koyama, 1985, p. 106; van Noordwijk & van Schaik, 1 987, p. 585; Ruiter etal., 1994, p. 218). In the Ketambe study area, northern Sumatra, a natural nonprovisioned population of M. fas- cicularis increased from 70-75 in 1972 to more than 190 in 1986 (van Schaik & van Noordwijk, 1988, p. 94), an annual rate of increase of about 7%; at this rate, which obviously is not indefinitely sustainable, the population would double every 10.5 years. The introduced nonprovisioned pop- ulation in Angaur increased in less than 75 years from a few individuals to 480-600 in 1 973 (Poirier & Smith, 1974, pp. 264, 271; cf. Kawamoto et al., 1988, p. 176; Matsubayashi et al., 1989, p. 54). In Mauritius, another introduced nonprovisioned population apparently increased in less than 500 years from a few individuals to about 30,000 (Sussman & Tattersall, 1986, pp. 30, 38; Lawler et al., 1995, p. 138). In Florida, a seminatural menagerie group increased from 6 individuals (4 females, 2 males) in 1933 to about 150 individuals in 1966 (Dumond, 1967, p. 203), an annual rate of increase of about 1 0%. Fossils and Subfossils Known fossils indicate that M. fascicularis has inhabited the Sunda Shelf since at least later Early Pleistocene, approximately 1 Ma (Table 33). All of the earliest macaque fossils in this area, dating from 0.8 to 1.0 Ma, have been collected in Java, where, at this time, M. fascicularis apparently was sympatric with M. nemestrina. In Java, M. fas- cicularis fossils of this age outnumber those of M. nemestrina, but most of these fossils consist of isolated teeth or small jaw fragments, and some may be misidentified. The long period from 800 Ka to 35 Ka has yielded no macaque fossils within the present geo- graphic range of M. fascicularis. However, fossils and subfossils of M. fascicularis and M. nemes- trina have been collected in subsequent Late Pleis- tocene and Holocene deposits in West Malaysia, Sumatra, Borneo, and Java (M. fascicularis fossils only). In these deposits in Borneo and Java, M. fascicularis fossils and subfossils outnumber those of M. nemestrina. In deposits in West Malaysia and Sumatra, by contrast, M. nemestrina fossils and subfossils outnumber those of M. fascicularis. This is anomalous, because, at present, M. fasci- cularis generally outnumbers M. nemestrina in the broad area of overlap of their respective ranges, except in the interior of primary evergreen rain forest (Fooden, 1975, p. 61; above, Natural His- tory). The explanation for the anomalous relative frequencies of fossils and subfossils of these species in Sumatra and West Malaysia is unclear. In the Lesser Sunda Islands, Holocene subfossils 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 of distribution of recognized subspecies of Macaca fascicularis and type localities of synonymous nominal species or subspecies (cf. Fig. 1; Fooden, 1982, p. 576); in the Philippine Islands, the indicated 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: M. f. aurea 1 . Pithecus vitiis Elliot, 1910; Burma, Letsok-aw Kyun. M. f. fascicularis 2. Pithecus validus Elliot, 1909; Vietnam: "Cochin China." 3. Pithecus capitalis Elliot, 1910; Thailand, Trang Province: Ban Phra Muang. 4. Macaca irus argentimembris Kloss, 1911b; Malay- sia, West Malaysia, P. Pinang [2]. 5. Pithecus agnatus Elliot, 1910; Indonesia, P. Tuang- ku. 6. Macacus phaeur a Miller, 1 903b; Indonesia, P. Nias: Teluk Siaba. 7. Pithecus mansalaris Lyon, 1916; Indonesia, P. Mur- sala. 8. Macacus irus I. Geoffroy, 1 826; Indonesia, Sumatra. 9. Macacus carbonarius I. Geoffroy, 1826; Indonesia, Sumatra (technical name also mistakenly applied to M. f. aurea and M. f. umbrosa). 10. Pithecus alacer Elliot, 1909; Indonesia, P. Kundur: Selatbliat. 11. Pithecus karimoni Elliot, 1909; Indonesia, P. Kar- imun: Monos. 12. Pithecus dollmani Elliot, 1909; Singapore, Singa- pore I.: Changi. 13. Pithecus laetus Elliot. 1909: Malaysia, West Malay- sia, P. Tinggi. 14. Pithecus bintangensis Elliot, 1909; Indonesia, P. Bintan: Sungei Biru. 15. Pithecus impudens Elliot, 1910; Indonesia, P. Sugi. 16. Pithecus lingae Elliot, 1910; Indonesia, P. Lingga. 17. Pithecus lapsus Elliot, 1910; Indonesia, P. Bangka: Tanjung Pamuja. 18. Pithecus lautensis Elliot, 1910; Indonesia, P. Laut. 1 9. Pithecus lingungensis Elliot, 1910; Indonesia, P. La- gong. 20. Pithecus sirhassensis Elliot, 1910; Indonesia, P. Ser- asan. 2 1 . Macacus pumilus Miller, 1 900; Indonesia, P. Benua. 22. Pithecus mandibularis Elliot, 1910; Indonesia, Bor- neo, Kalimantan: Sungai Ambawang, near Pontia- nak. 23. Pithecus carimatae Elliot, 1910; Indonesia, P. Kar- imata: Teluk Pai. 64 FIELDIANA: ZOOLOGY mains, have been collected in caves in Timor and Flores, both of which are east of Wallace's Line (passing between Bali and Lombok). In four care- fully excavated caves in Timor, where basal de- posits date back to ca. 13.5 Ka, the oldest M. fascicularis subfossils do not appear until ca. 4.5 Ka (Glover, 1986, p. 212). At lower levels, the only mammals included among the human food remains are murid rodents and bats, of which thousands of skeletal fragments were collected; these lower levels also yield stone tools. At ap- proximately the same level where M. fascicularis first appears, remains of palm civet and cuscus also appear, along with remains of domestic goat, pig, and dog and pottery. Glover (1986, p. 159) concluded that "... there is strong presumptive evidence that these species [M. fascicularis, palm civet, and cuscus] were introduced into Timor di- rectly by, or through the agency of man, between about 4000-5000 years ago." For M. fascicularis, a less plausible alternative hypothesis is that it was present in Timor earlier but was not hunted by the preceramic cave inhabitants, possibly because they lacked suitable projectile weapons (cf. Adi, 1985, p. 65). In Flores, a subfossil molar of M. fascicularis was collected in cave deposits, dated ca. 3.5 Ka, that also contain human artifacts and human skel- etal remains. Other mammals represented in these deposits are rat species, fruit bat, porcupine, and pig (Hooijer, 1967, p. 160). Based on available zoogeographic evidence, including Glover's ar- cheological data from Timor (see above), Musser ( 1 98 1 , p. 133) concluded that M. fascicularis, por- cupine, and pig probably were introduced into Flo- res by humans. Systematics Subspecific Taxonomy The geographic range of M. fascicularis is broad and encompasses mainland Southeast Asia and numerous large and small islands, both on and beyond the Sunda Shelf (Figs. 3, 4). It is therefore not surprising that this species exhibits great in- dividual, local, and geographic variation in a di- verse array of characters (see above); variation among populations of M. fascicularis ranges from trivial, to statistically significant, to locally dis- continuous, to totally discontinuous. Based on this variation, 50 specific or subspecific names have been proposed for populations of this species (Fig. 25). The evaluation of intraspecific variation for the purpose of defining taxonomically useful sub- species presents serious theoretical and practical difficulties, as previously discussed by Chasen (1940a, p. 66; cf. Table 35) in his classic review of M. fascicularis. In the present taxonomic analysis, subspecific status is accorded only to those geographic pop- ulations in which variation of at least one character is completely— or nearly completely— discontin- uous with variation of the same character in the nominotypical subspecies, M. f fascicularis. The reason for selecting character-state discontinuity as the critical requisite for subspecific recognition is that, without such discontinuity, unambiguous diagnosis of subspecies becomes difficult or im- possible. The requirement of character-state dis- continuity denies subspecific recognition to local populations that are distinct from adjacent pop- ulations of M. f fascicularis but that fall within 24. Macaca resima Thomas & Wroughton, 1909b; In- donesia, Java: Tasikmalaya. 25. Macaca modax Thomas & Wroughton, 1909c; In- donesia, Java: Cilacap. 26. [Simia] Aygula Linnaeus, 1758; Indonesia, Java. 27. Pithecus baweanus Elliot, 1910; Indonesia, P. Baw- ean. 28. Pithecus cupidus Elliot, 1910; Indonesia, P. Mata- siri. 29. Macaca irus submordax Sody, 1949; Indonesia, P. Bali: Desa Poetjang. 30. Macaca irus sublimitus Sody, 1932; Indonesia, P. Sumba: Payeti-Kambaniru and Mao Marroe. 31. Pithecus fascicularis limitis Schwartz, 1913; Indo- nesia, P. Timor: Lelogama. 32. Cynomolgus cagayanus Mearns, 1905; Philippines, Cagayan Sulu I. 33. Cynomolgus suluensis Meanrs, 1905; Philippines, Jolo I.: foot of Crater Lake Mountain. M. f. fascicularis/ M. f. philippinensis contact zone 34. Cynomolgus mindanensis apoensis Mearns, 1905; Philippines, Mindanao I.: Mt. Apo. 35. Cynomolgus mindanensis Mearns, 1905; Philip- pines, Mindanao I.: Pantar. M. f. philippinensis 36. Pithecus mindorus Hollister, 1913; Philippines, Mindoro I.: Alag River. 37. Macacus palpebrosus I. Geoffroy, 1851; Philippines, Luzon I.; "forets de Manille." 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 Dorsal Lateral Head and Relative pelage Crown facial crest Thigh (outer body length, 4 tail length, 4 Subspecies 2 color color pattern 3 surface), color ad. males ad. males aurea Buffy to me- Golden Infrazyg. Similar to dors. 560 ± 61 95 ± 9 dium brown pel. col. 520-630 85-104 brown N = 3 atriceps Buffy to me- Dark brown Transzyg. Similar to dors. 452 ± 17 118 ± 7 dium to blackish pel. col. 425-465 110-128 brown (narrow) 5 N = 5 condorensis Buffy to me- Dark brown Transzyg. Similar to dors. 450 ± 18 115 ± 7 dium to blackish pel. col. 435^180 109-127 brown (broad) 5 N = 5 fascicularis Buffy to dark Golden Transzyg. Similar to dors. 462 ± 43 118 ± 15 brown brown pel. col. 370-610 N= 193 69-150 N = 189 karimondjawae Dark grayish Dark brown 7 Similar to dors. 501 108 brown to diffusely blackish pel. col. N = 1 philippinensis Dark brown Dark golden Transzyg. Similar to dors. 480 ± 33 114 ± 9 brown pel. col. 410-530 N = 101-129 13 tua Blackish Yellowish brown Transzyg. 6 Brownish gray 440 N = 131 1 umbrosa Blackish Yellowish Infrazyg. or Pale brownish gray 502 ± 25 115 ± 1 brown transzyg. 7 475-525 N = 470 115-116 3 118 lasiae Blackish Blackish 8 Infrazyg. or Similar to dors. transzyg. pel. col. 9 N = 1 fusca Blackish Blackish 10 Infrazyg. or Similar to dors. 478 ± 14 97 ± 6 transzyg. pel. col." 460-495 N = 90-105 6 1 Abbreviations: ad. = adult; infrazyg. = infrazygomatic; transzyg. = transzygomatic. 2 Listed in order of increasing pelage color saturation. 3 See Figure 8. 4 Entries indicate mean ± SD, extremes, and sample size; relative tail length = tail length/head and body length 100. 5 See Figure 7. 6 Subauricular hairs pale ochraceous-buff, conspicuously elongated. 7 Subauricular hairs pale brownish, not elongated. 8 Crown hairs inconspicuously annulated. 9 Outer surface of shanks pale grayish brown. 10 Crown hairs conspicuously annulated with pale yellowish. 1 ' Outer surface of shanks blackish. the range of variation of more distant populations of this subspecies. This is not intended to depre- ciate the theoretical importance of locally deviant populations, which often have great zoogeographic or evolutionary significance (see below); appreci- ation of this significance does not require formal subspecific recognition. A total of 1 subspecies of M. fascicularis are recognized here (Fig. 25, Table 34, Appendix 12). These are M.f. fascicularis (core area, many shal- low-water fringing islands, some deep-water fring- ing islands); M. f aurea (core area northwest of M.f fascicularis, adjacent shallow-water fringing islands); M. f atriceps, M. f condorensis, and M. karimondjawae (shallow-water fringing islands); and M. f umbrosa, M. fascicularis fusca, M. f lasiae, M. f tua, and M. f philippinensis (deep- water fringing islands). Characters and distribu- tions of these subspecies are briefly reviewed be- low and are treated in greater detail in subsequent subspecies accounts. 1 . M.f. fascicularis. The type locality of M. fas- cicularis— and, hence, of M. f fascicularis— is Bengkulu, southwestern Sumatra, a locality that fortuitously lies within the previously defined core area of distribution of the species. Judging from 66 FIELDIANA: ZOOLOGY available evidence, variation of characters in Su- matra is continuous with that in the Malay Pen- insula (including the Isthmus of Kra north to ca. 1 0°N), Borneo, and Java (see above); all of these parts of the core area are therefore included within the geographic range of M. f. fascicularis, as here defined. In the southern part of the Indochinese Peninsula, character-state variation in M. fasci- cularis also is continuous with that in the Malay Peninsula, Sumatra, Borneo, and Java, with the possible exception of variation of mtDNA types (Table 17) and resistance to experimental infec- tions with Plasmodium knowlesi malaria (see above; Fooden, 1994, p. 585). Because evidence for discontinuity of variation of mtDNA types and P. knowlesi resistance is incomplete, the southern part of the Indochinese Peninsula is provisionally retained within the geographic range of M. f. fas- cicularis. Although now disjunct, the range of M. f fascicularis in the southern part of the Indo- Chinese Peninsula was connected by dry land to the range of this subspecies in the Malay Peninsula during the last glacial maximum, ca. 1 8 Ka (Fig. 3). In most shallow-water fringing-island popula- tions of M. fascicularis, variation is continuous with that in core-area M. f fascicularis (Appen- dixes 3-9; Fooden & Albrecht, 1993, p. 537); therefore, these islands are included within the geographic range of M. f fascicularis. In eight of these shallow-water fringing-islands, populations are distinct from nearby core-area populations of M.f fascicularis but fall within the range of vari- ation of more distant populations of this subspe- cies; as indicated above, these populations are not recognized here as separate subspecies, although some have been recognized as subspecies by other authors. These eight populations, sorted according to characters that distinguish the fringing-island populations from nearby core-area populations of M.f. fascicularis, inhabit the following islands: P. Mursala (west of Sumatra), P. Redang (east of the Malay Peninsula), P. Lagong (west of Borneo), and P. Uwi (west of Borneo)— distinguished by dorsal pelage erythrism (Appendix 4); P. Tioman (east of the Malay Peninsula) and P. Belitung (west of Bor- neo)— distinguished by T and/or relative T (Ap- pendixes 8, 9); and Ko Kut and Phu Quoc Dao (both south of the Indochinese Peninsula)— dis- tinguished by skull length (Fooden & Albrecht, 1993, p. 528). In many other fringing-island pop- ulations, character-state variation is statistically significantly different from, but continuous with, variation in nearby core-area populations (Ap- pendixes 3-9); some of these populations also have been recognized as separate subspecies by other authors. The deep-water fringing-island population of M. fascicularis in P. Nias (west of Sumatra) resembles the eight shallow-water fringing-island popula- tions listed above in being distinct— in tail col- oration and T— from nearby core-area popula- tions of M. f fascicularis in Sumatra but falling within the range of variation of more distant pop- ulations of this subspecies. Although the proxi- modorsal surface of the tail is more intensely blackish in P. Nias than in Sumatra, as empha- sized by Miller (1903b, p. 63) in his description of the P. Nias monkey as Macacus phaeura, tail coloration in M. fascicularis in P. Nias broadly overlaps that in specimens collected in other parts of the species range (cf. mcz 36030, usnm 121871 — P. Nias; usnm 196823— Kalimantan; mcz 37415 — Sabah; usnm 124710-P. Bangka; usnm 124969, 124970-P. Belitung). T and relative T in P. Nias similarly overlap T and relative T in other parts of the species range (Figs. 15, 17; Appendixes 8, 9). Lacking character-state discontinuity, M. fas- cicularis in P. Nias is regarded here as subspecif- ically inseparable from M. f fascicularis. Deep- water fringing-island populations of M. fascicu- laris in the Lesser Sundas and southern Philippines are retained within M.f. fascicularis because known variation in these populations is continuous with that in nearby core-area populations of M. f fas- cicularis (Appendixes 3-5, 7-9; Fooden, 1991, p. 21; Fooden & Albrecht, 1993, p. 537). 2. M. f aurea. In the western part of the In- dochinese Peninsula and western part of the Isth- mus of Kra south to about 1 0°N, variation of the lateral facial crest pattern (infrazygomatic) is al- most completely discontinuous with variation of this character (transzygomatic) in the rest of the core area (Figs. 8, 9). This warrants recognition of the subspecies M. f aurea (type locality, Pegu, southern Burma). In all samples available from shallow-water islands in the Mergui Archipelago, west of the northern part of the Isthmus of Kra, the lateral facial crest pattern also is infrazygo- matic; these islands are therefore included within the geographic range of M. f aurea. Two areas immediately east of the range of M. f aurea are heterogeneous for the lateral facial crest pattern (Fig. 9). The more southern of these two areas, in eastern and southern parts of the Isthmus of Kra and adjacent southwestern islands, may be regarded as an intersubspecific contact zone 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 (— ) indicates that subspecies is extralimital in the cited work. Author and date w. c. o. P. H. Groves and Chasen Hill Napier Scott Weitzel Fa This Subspecies (1940a) (1974) (1981) (1982) (1988)» 1989 paper argentimembris X X X X X atriceps X2 X X X X X aurea X 3 X X X X baweana X X X X — X bintangensis X X cagayana — X — — capitalis X X X X condorensis X2 X X X 3 X cupida X X X — — X fascicularis (= irus) X X X X X X X fusca X X X — — X X impudens X — karimondjawae X X X — X laeta X X X X X X lasiae X — — X X limitis — X X X — X lingungensis X X mandibularis X X mindanensis — — — X mindora — — — X mordax X X X X X X phaeura X X X — X 3 X philippinensis — X X — — X X pumila X X X X — X sirhassensis X X sublimitis — X X — X submordax X — tua — — X X umbrosa X2 X X — — X X valida — X X X 3 X Subsp. (Borneo, north) X X Subsp. (P. Banggi) X Subsp. (P. Siantan) X In Weitzel et al. (1988, pp. 5, 96). Subspecies recognized, name not applied. Subspecies recognized provisionally. aurea. The more northern of these two areas, in central and eastern parts of the Indochinese Pen- insula, along the northern margin of the species range, may be either an intersubspecific contact zone between the ranges of M. f. fascicularis and M. f. aurea or an interspecific contact zone be- tween the ranges of M. fascicularis and M. mulatta (cf. Fooden, 1971, p. 24); future study of variation in M. mulatta may contribute to resolution of this ambiguity. 3. M. f. atriceps. The distinctive narrow dark crown patch warrants recognition of this subspe- cies in shallow-water Ko Khram Yai in the Gulf of Thailand (Fig. 7, Table 5, Appendix 5). 4. M.f. condorensis. The distinctive broad dark crown patch warrants recognition of this subspe- cies in shallow-water Con Son and nearby Hon Ba, both in the South China Sea (Fig. 7, Table 5, Appendix 5). 5. M. f karimondjawae. Variation of dorsal pelage color saturation in the population of M. fascicularis in P. Karimunjawa and presumably also in nearby P. Kemujan, both shallow- water islands north of Java, overlaps minimally with that in M. f fascicularis (Appendix 3). The dark grayish brown dorsal pelage color of this popula- tion is the basis for recognition of M. f. kari- mondjawae. 6. M.f. umbrosa. The population of M. fasci- cularis in the Nicobar Islands, northwest of Su- 68 FTELDIANA: ZOOLOGY matra, is one of four deep-water fringing-island populations with distinctively blackish dorsal pel- age (Appendix 3). The crown is yellowish brown and the thighs are pale brownish gray in M. f. umbrosa. 7. M. fascicularis fusca. In the population of M. fascicularis in deep-water P. Simeulue, west of Su- matra, the back, crown, and thighs are blackish, and the tail is shorter than usual in M. fascicularis (Figs. 16, 18; Appendixes 3, 8, 9). 8. M.f lasiae. In the population in deep-water P. Lasia, near P. Simeulue, the back, crown, and thighs are blackish, as in M. fascicularis fusca, but the tail is of normal length (Figs. 16, 18, 27; Ap- pendixes 3, 8, 9). 9. M.f. tua. In the population in deep-water P. Maratua, east of Borneo, the back is blackish, the crown is yellowish brown, and the thighs are brownish gray (Appendix 3). In M. f tua, unlike M. f. umbrosa, subauricular hairs are elongated and pale ochraceous-buff. 10. M. f philippinensis. In populations of M. fascicularis in western, northern, and eastern is- lands of the Philippine Archipelago, dorsal pelage color is distinctively dark brown (Appendix 3; Fooden, 1991, p. 3). In eastern and central Min- danao, southern Negros, and perhaps in some nearby islands, mixed populations of dark and pale individuals occur; this area of mixed populations is regarded as a contact zone between M. f phi- lippinensis and M. f fascicularis. Brief casual remarks have been published con- cerning dorsal pelage coloration in M. fascicularis in P. We, a deep-water island off the northern tip of Sumatra (Scheffrahn et al., 1994, p. 136). These remarks are inadequate for determination of the subspecific status of this population. The present subspecific classification of M. fas- cicularis differs from recent classifications pro- posed by W. C. O. Hill (1 974, p. 504), R. H. Napier (1981, p. 12), Scott (1982, unpubl. M.A. thesis, p. 182), Weitzel et al. (1988, p. 96), and Fa (1989, p. 54) (Table 35). P. H. Napier, whose list of sub- species follows that of W. C. O. Hill with a single exception (nonrecognition of subspecies cagayana Mearns, 1905), explicitly acknowledges that her classification is unsatisfactory and that many of the recognized subspecies cannot be diagnosed. Scott's classification is based on a sample of 148 specimens (including 1 34 skins and 69 skulls), more than 90% of which are in two museums (mzb, zrc); this sample, which apparently is almost identical with that studied by Chasen (1940a, p. xix), pro- vides an inadequate representation of variation in M. fascicularis. Weitzel et al. indicate that their classification is largely based on Scott's research. Fa, without further explanation or documentation, lists 2 1 subspecies of M. fascicularis that he rec- ognizes as valid; most, but not all, of these sub- species are the same as those recognized by authors cited above. 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 General color of dorsal surface of trunk dark brown, 1 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 Outer surface of shanks pale grayish brown; tail length > 1 1 2% of head and body length . . . lasiae 9. Subauricular hairs pale ochraceous-buff, elongated (4-6 cm), extending to apex of lateral facial crest forming a conspicuous pale lateral facial patch; tail length > 1 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 Macaca fascicularis fascicularis (Raffles, [1821], p. 246) Jawanska Markattor: Osbeck, 1757, p. 99— external characters and behavior of monkeys collected and ob- served in Java by P. Osbeck, Jul. 1751; specimens not preserved. [Simia] Aygula Linnaeus, 1758, p. 27 — based on Ja- wanska Markattor: Osbeck, 1757, and supplementary information in unpublished letters from Osbeck to Linnaeus (P. H. Napier & Groves, 1 983, p. 1 1 7). Tho- mas & Wroughton, 1909a, p. 373 — misidentified as a species of leaf monkey. P. H. Napier & Groves, 1983, p. 118— shown to be a senior synonym of Macaca fascicularis (Raffles, [1821]); suppression requested. International Commission on Zoological Nomencla- ture, 1986, p. 229— specific name officially suppressed. [Cercopithecus] aygula: Erxleben, 1777, p. 39— external and cranial characters. Cercop[ithecus] Aigula: Schinz, 1821, p. 108— incorrect spelling of [Simia] Aygula Linnaeus, 1758. Schinz, 1825, p. 256 — said to be indistinguishable from Inuus cynamolgus [— Macaca fascicularis]. Macaca aygula: Daudin in Lacepede & Daudin, [1802], p. 148— listed as "Variete A" of Macaca cynomolgus [= M. fascicularis]. Pithecus aygula: E. Geoffroy, 1 803, p. 24— external char- acters. Cercocebus aygula: E. Geoffroy, 1812, p. 99— external characters. Macaque: Buffon in Buffon & Daubenton, 1766, p. 190, pis. 20, 22-24— based on specimen of unknown origin, not preserved; species said to inhabit "Congo & des autres parties de l'Afrique meridionale." F. Cuvier, 1819, liv. 3, p. 1, 2 pis. — illustrations of captives des- ignated as Irus by F. Cuvier, 1818; habitat, "vra- isemblablement du Senegal ou de la cote de Guinee." Aigrette: Buffon in Buffon & Daubenton, 1766, p. 190, pi. 21— based on specimen of unknown origin, not preserved; regarded as "variete" of Macaque: Buffon in Buffon & Daubenton, 1766. 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, 1876, p. 101 —name said to have been based on mis- identification. Blanford, [1888a], p. 624— name said to have been based on misidentification. Cercopithecus (Cynamolgus): Zimmermann, 1780, p. 186 (part; not Linnaeus, 1758, p. 28)— misidentification of Macaque: Buffon in Buffon & Daubenton, 1766. Cercocebus cynamolgos: Schlegel, 1876, p. 101 (not Lin- naeus, 1758)— taxonomic comparisons. [Cercopithecus] Cynomolgus: Erxleben, 1 777, p. 28 (part; not Linnaeus, 1766, p. 38)— misidentification of Ma- caque: Buffon in Buffon & Daubenton, 1766. Simia Cynomolgus: Audebert, 1798-1799, p. 5 (part; not Linnaeus, 1766, p. 38)— misidentification of Ma- caque: Buffon in Buffon & Daubenton, 1766. Simia cynomolgos: G. Cuvier, 1798, p. 98— incorrect spelling of [Simia] Cynomolgus: Linnaeus, 1766; mis- identification of Macaque: Buffon in Buffon & Dau- benton, 1766. Macaca cynomolgus: Daudin in Lacepede & Daudin, [1802], p. 148 (not Linnaeus, 1766)— misidentifica- tion of Macaque: Buffon in Buffon & Daubenton, 1766. [M]acacus cynomolgus: Desmarest, 1820, p. 65 (part; not Linnaeus, 1766)— misidentification of Macaque: Buffon in Buffon & Daubenton, 1766. Blanford, 1888b, pp. 21, 23— binomial said to be based on misidenti- fication, accepted as valid. Macacus ' cynomolgus' Auct.: Miller, 1900, p. 239 — specimens collected in P. Lingga and P. Tioman. Mil- ler, 1903a, p. 476 — specimens collected at Teluk Ta- panuli, Sumatra, and in P. Tuangku and P. Mursala. Cyn[ocephalus] cynomolgus: Latreille, 1 804, p. 292 (part; not Linnaeus, 1766, p. 38)— misidentification of Ma- caque: Buffon in Buffon & Daubenton, 1766. Cercocebus cynomolgus: E. Geoffroy, 1812, p. 99 (not Linnaeus, 1766)— external and cranial characters. [P]ithecus cynomolgus: Desmarest, 1817, p. 323 (part; not Linnaeus, 1766, p. 38)— misidentification of Ma- caque: Buffon in Buffon & Daubenton, 1766. Pithecus Cynomolgos: Blainville, [1839], pi. 7 — incorrect spelling of [Simia] Cynomolgus: Linnaeus, 1 766. Pith[ecus] (Mac[acus]) cynomolgus: Dahlbom, 1856, p. 1 18 (not Linnaeus, 1766, p. 38)— external characters. 70 FIELDIANA: ZOOLOGY I[nuus] cynomolgus: Wagner, [1839], p. 135 (not Lin- naeus, 1766, p. 38)— external characters; distribution; taxonomic comparisons. Irus: F. Cuvier, 1818, p. 120— unavailable name, not published in combination with generic name; pro- posed as substitute specific technical name for Ma- caque: Buffon in Buffon & Daubenton, 1766 (cf. I. Geoffrey, 1826, p. 588; [1831], p. 56: Gervais, 1854, p. 85; Miller, 1942, p. 127); habitat "vraisemblable- ment du Senegal ou de la cote de Guinee" (corrected to "du midi de l'Asie, et particulierement de Sumatra" by F. Cuvier, 1825, liv. 52, p. 1). Simia fasciculcris Raffles, [1821], p. 246— for details concerning holotype, see below. Cantor, 1846, p. 176 — cited as a synonym of Cercopithecus cynomolgus. H. C. Robinson, 1916, p. 63— type locality restriction. Semn[nopithecus]l fascicularis: [Vigors], 1830, p. 642— geographic distribution; type history; "doubtful whether it is a true Semnopithecus." Semnopithecus fascicularis?: Waterhouse, 1838, p. 4— geographic distribution. S[emnopithecus] fascicularis: Martin, 1838, p. 435— ex- ternal characters. Macacus fascicularis: Bonhote, 1903, p. 3— taxonomic history. Macaca fascicularis: Miller, 1906b, p. 65— specimens collected in P. Karimata. Wroughton, 1918, p. 556— taxonomic history. Miller, 1942, p. 126— taxonomic history. Hooijer, 1962b, p. 44— subspecific taxonomy. Macaca fascicularis group: Thomas, 1928, p. 832— spec- imens collected in Vietnam. [Cynomolgus] fascicularis: Trouessart, 1904, p. 16— geo- graphic distribution. Pith ecus fascicularis: Elliot, 1913, p. 233— taxonomic comparison. [Silenus] fascicularis: Stiles & Nolan, 1929, p. 529— parasites. P[ithecus] f[ascicularis] fascicularis: Schwarz, 1913, p. 297— taxonomic comparison. M[acaca] fascicularis fascicularis J. R. Napier & Napier, 1967, p. 403— geographic distribution. Macaca mulatto fascicularis: Fooden, 1964, p. 364— taxonomic comparison. Kra Buku: Raffles, [1821], p. 247— vernacular name of "smaller species" than Simia fascicularis Raffles, [1821], p. 246 [apparently juvenile Macaca fascicu- laris], specimens apparently not obtained; habitat, Su- matra and "other Malay islands." Macacus irus I. Geoffroy, 1826, p. 588— based on Ma- caque: Buffon in Buffon & Daubenton, 1 766, and liv- ing captives reported by F. Cuvier (1818, p. 112; 1819, liv. 3, p. 1); specimens not preserved, origin unknown; binomial incorrectly attributed to F. Cuvier, 1818. I. Geoffroy, [1831], p. 56— cited as a synonym of Ma- cacus cynomolgus Desmarest, 1820; taxonomic his- tory. Blanford, [1888a], p. 624— taxonomic history. Cabrera, 1910, p. 620— taxonomic history; type lo- cality, Sumatra. H. C. Robinson & Kloss, 1914, p. 394— type locality, probably Malacca (= Melaka). Miller, 1942, p. 127— cited as a synonym of Macaca fascicularis (Raffles, [1821]); binomial attributed to Blanford, [1888a]. Fooden, 1976, p. 226-cited as a synonym of Macaca fascicularis (Raffles, [1821]); bi- nomial attributed to I. Geoffroy, 1826. Macaca irus: H. C. Robinson & Kloss, 1915a, p. 130 — specimens collected in Ko Samui and Ko Phangan. [Macaca] zVus-group: Miller, 1933, pp. 5, 6— taxonomic history. [Macaca] ira: Weinman & Wiratmadja, 1969, p. 498 — trypanosomes. Pithecus irus: Elliot, 1913, p. 229— taxonomic compar- ison. [Silenus] irus: Stiles & Nolan, 1929, p. 530— parasites. Cynomolgus irus: Furuya, 1962, p. 377— dermatoglyph- ics. Macaca irus irus: H. C. Robinson & Kloss, 1918, p. 6— collected in Sumatra; taxonomic history. Chasen, 1940a, p. 66— taxonomic comparisons. Kellogg, 1945, pp. 116, 129— taxonomic comparisons. Sody, 1949, p. 130— taxonomic comparison. Carbonarius: F. Cuvier, 1825, liv. 52, p. 2, pi.— un- available name, not published in combination with generic name; based on living captive, origin un- known; specimen not preserved. Macacus carbonarius I. Geoffroy, 1826, p. 588— based on Carbonarius: F. Cuvier, 1825; external characters; taxonomic comparison. Lesson, 1827, p. 42— said to inhabit Sumatra. I. Geoffroy, [1831], p. 63— doubt- fully distinct. Eydoux & Souleyet [& Gervais], 1841, p. 6— cited as a synonym of Macacus aureus I. Geof- froy, [1831] [= Macaca fascicularis aurea]. Blyth, 1875, p. 7— cited as a synonym of Macacus cynomolgus [= Macaca fascicularis]. S[imia] carbonaria: Fischer, 1829, p. 26— habitat, Su- matra. C[ynamolgus] carbonarius: Reichenbach, 1862, p. 136 — taxonomic comparison. Semnopithecus kra Lesson, [1830], p. 20— replacement name for Simia fascicularis Raffles, [1821], p. 246. Semnopithecus Buku Martin, 1 838, p. 435 —specific name proposed provisionally for Kra Buku: Raffles, [1821], p. 247. J. Anderson, 1879, p. 74— cited as a synonym of Macacus cynomolgus [= Macaca fascicularis]. Inuus cercopithecus Mason, 1851, p. 220— unavailable name attributed to Blyth, cited in synonymy, never treated as available. Macacus cancrivorus Mason, 1 85 1 , p. 22 1 —unavailable name attributed to Blyth, cited in synonymy, never treated as available. C[ynamolgus] cynocephalus: Reichenbach, 1862, p. 133 (not Linnaeus, 1766, p. 38)— natural history. Macacus assamensis: Gray, 1 870, p. 3 1 (not McClelland in Horsfield, [1840], p. 148)— misidentification; tax- onomic comparison. Macacus Sinicus: Morice, 1875, p. 41 (not Linnaeus, 1 77 1 , p. 52 1)— misidentification of monkeys observed in Vietnam. Macacus pumilus Miller, 1900, p. 241 —holotype, usnm 101639, adult male, skin and skull, collected at Pulo Bunoa (= Pulau Benua), Kepulauan Tambelan, In- donesia, by W. L. Abbott, 6 Aug. 1899; paratypes, usnm 101638 (juvenile female, P. Benua, 5 Aug.), usnm 101666 (adult female, P. Uwi, 13 Aug.), and usnm 101711 (subadult male, P. Siantan, 8 Sep.), skins and skulls, collected in Indonesia by W. L. Abbott, 1899. Lyon & Osgood, 1909, p. 284— holotype cataloged. Weitzel et al., 1988, p. 112— topoparatypic specimens (P. Siantan) referred to M. fascicularis sirhassensis (El- liot, 1910). FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 71 [Cynomolgus] pumilus: Trouessart, 1904, p. 16— distri- bution. Pithecus pumilus: Elliot, 1 9 1 3, p. 252— external and cra- nial characters. Macaca irus pumilus: Chasen & Kloss, 1928b, p. 29 — taxonomic comparison. Macaca irus pumila: Chasen, 1940a, p. 69— geographic distribution. Poole & Schantz, 1942, p. 245 — holotype cataloged. Weitzel et al., 1988, pp. Ill, 1 13— cited as name previously applied to specimens referred to M. fascicularis lingungensis (Elliot, 1910) and M. f. sir- hassensis (Elliot, 1910). M[acaca] f[ascicularis] pumila: J. R. Napier & Napier, 1967, pp. 349, 403— geographic distribution. Macacus phaeura Miller, 1903a, p. 63 — holotype, usnm 121870 (Coll. No. 2399), adult male, skin and skull, collected at Teluk Siaba, P. Nias, Indonesia, by W. L. Abbott, 20 Mar. 1903; paratypes, usnm 121868 (Coll. No. 2393, adult male, 16 Mar.), usnm 121869 (Coll. No. 2395, adult female, 18 Mar.), and usnm 121871 (Coll. No. 2400, adult male, 20 Mar.), skins and skulls, collected at Teluk Siaba, P. Nias, Indonesia, by W. L. Abbott, 1903. Lyon & Osgood, 1909, p. 284-holo- type cataloged. Macaca phaeura: Lyon, 1906, p. 606— specimens col- lected in P. Bangka and P. Belitung. Miller, 1942, p. 128 — specimens collected in P. Nias. Pithecus phaeurus: Elliot, 1913, p. 243— external and cranial characters. Pithecus phaeura: Lyon, 1916, p. 458— geographic dis- tribution. Macaca irus phaeura: Chasen, 1940a, p. 69— geographic distribution. Poole & Schantz, 1942, p. 245— holotype cataloged. M[acaca] /[ascicularis] phaeura: J. R. Napier & Napier, 1967, pp. 349, 403— geographic distribution. Cynomolgus suluensis Mearns, 1905, p. 430— holotype, usnm 125324 (Coll. No. 5750), adult male, skull only, collected at foot of Crater Lake Mountain, Jolo I., Philippine Islands, by E. A. Mearns, 16 Nov. 1903. W. C. O. Hill, 1974, p. 522-cited as a synonym of Macaca irus philippinensis I Geoffroy, [ 1 843]. Fooden, 1991, p. 22— cited as a synonym of Macaca fascicu- laris fascicularis (Raffles, [1821]). Macaca suluensis: Lyon & Osgood, 1909, p. 284— ho- lotype cataloged. Macaca suluana: Alcasid, [1970], p. 24— incorrect spell- ing of Cynomolgus suluensis Mearns, 1905. Macaca sulvensis: Chiarelli, 1972, p. 2 1 3 — incorrect spelling of Cynomolgus suluensis Mearns, 1905. [Macacus] suluensis: Raven, 1935, p. 237— geographic distribution. Pithecus suluensis: Hollister, 1912, p. 37 — listed. Elliot, 1913, p. 252— specific status indeterminate. M[acaca] p[hilippinensis] suluensis: Rabor, 1986, p. 1 38 — geographic distribution. Cynomolgus cagayanus Mearns, 1905, p. 431— holo- type, usnm 125325 (Coll. No. 5771), adult male, skin and skull, collected in Cagayan Sulu I., Philippine Is- lands, by E. A. Mearns, 25 Feb. 1904. P. H. Napier, 1981, p. 13— cited as a synonym of Macaca fascicu- laris philippinensis I. Geoffroy, [1843]. Fooden, 1991, p. 22— cited as a synonym of Macaca fascicularis fas- cicularis (Raffles, [1821]). Macaca cagayana: Lyon & Osgood, 1909, p. 283 — ho- lotype cataloged. Poole & Schantz, 1942, p. 241 — holotype cataloged. [Macacus] cagayanus: Raven, 1935, p. 236— geographic distribution. Pithecus cagayanus: Hollister, 1912, p. 36— geographic distribution. Macaca irus cagayana: W. C. O. Hill, 1974, p. 525 — external and cranial characters. M[acaca] p[hilippinensis] cagayanus: Rabor, 1986, p. 138— geographic distribution. Cynomolgus fuscus: Moszkowski, 1909, pp. 143, 302 (not Miller, 1903a, p. 476)— misidentification of spec- imens collected in Sumatra. Pithecus validus Elliot, 1909, p. 252 — holotype, bm(nh) 1881.6.30.2, adult male, skin and skull, collected in "Cochin China," Vietnam, by M. Boucard, before 1 882. Macaca validus: Kloss, 1919c, p. 348— taxonomic com- parison. M[acaca] i[rus] validus: Kloss, 1921, p. 75— taxonomic comparison. Macaca irus valida: Kellogg, 1945, p. 1 19— geographic distribution. M[acaca] f[ascicularis] valida: J. R. Napier & Napier, 1967, pp. 349, 403— geographic distribution. P. H. Napier, 1981, pp. 13, 19— external characters; holo- type cataloged. Macaca fascicularis validus: Van Peenen et al., 1971, p. 134 (part)— female specimens collected at Sontra Peak; geographic variation. Pithecus alacer Elliot, 1909, p. 253— holotype, bm(nh) 1909.4.1.36 (Coll. No. 1454), adult male, skin and skull, collected at Bliah (= Selatbliat), P. Kundur, Ke- pulauan Riau, Indonesia, by H. C. Robinson and E. Seimund, 18 Aug. 1908; paratype (available but not explicitly cited in original description), bm(nh) 1909.4.1.37 (Coll. No. 1495, adult female, skin and skull), collected at Selatbliat, P. Kundur, Indonesia, by H. C. Robinson and E. Seimund, 20 Aug. 1908. H. C. Robinson & Kloss, 1914, pp. 393, 394-type his- tory. Chasen, 1925, p. 93— cited as a synonym of Ma- caca irus F. Cuvier (I. Geoffroy, 1 826). Chasen, 1940a, p. 66— cited as a synonym of Macaca irus irus F. Cu- vier (I. Geoffroy, 1826). P. H. Napier, 1981, pp. 13, 15— cited as a synonym of Macaca fascicularis fasci- cularis (Raffles, [1821]); holotype cataloged. [Macacus] alacer: Raven, 1935, p. 236— geographic dis- tribution. M[acaca] i[rus] alacer: Dammerman, 1926b, p. 316 — geographic distribution. Pithecus karimoni Elliot, 1909, p. 254— holotype, bm(nh) 1909.4.1.34 (Coll. No. 1662), adult male, skin and skull, collected at Monos, P. Karimun, Kepulauan Riau, Indonesia, by H. C. Robinson and E. Seimund, 30 Aug. 1908; paratype, bm(nh) 1909.4.1.35 (Coll. No. 1636, adult female, skin and skull), collected at Mon- os, P. Karimun, Indonesia, by H. C. Robinson and E. Seimund, 29 Aug. 1908. H. C. Robinson Kloss, 1914, p. 393, 394— type history; taxonomic comparison. Chasen, 1925, p. 93— cited as a synonym of Macaca irus F. Cuvier (I. Geoffroy, 1826). Chasen, 1940a, p. 66— cited as a synonym of Macaca irus irus F. Cuvier (I. Geoffroy, 1826). P. H. Napier, 1981, pp. 13, 15- cited as a synonym of Macaca fascicularis fascicularis (Raffles, [1821]); holotype cataloged. [Macacus] karimoni: Raven, 1935, p. 236— geographic distribution. 72 FIELDIANA: ZOOLOGY M[acaca] i[rus] karimoni: Dammerman, 1926b, p. 316— geographic distribution. Pithecus laetus Elliot, 1909, p. 255— holotype, bm(nh) 1909.4.1.21 (Coll. No. 849), subadult male, skin and skull, collected in P. Tinggi, West Malaysia, by H. C. Robinson, 25 Jun. 1908; paratype (available but not explicitly cited in original description), bm(nh) 1909.4.1.27 (Coll. No. 844, juvenile female, skin and skull), collected in P. Tinggi, by H. C. Robinson, date unspecified, presumably Jun. 1908. Elliot, 1913, p. 236— geographic distribution, P. Tinggi and P. Tio- man, West Malaysia. H. C. Robinson & Kloss, 1914, p. 393— type history; taxonomic comparisons. Macaca laetus: Kloss, 191 la— taxonomic comparison. [Macacus] laetus: Raven, 1935, p. 236— geographic dis- tribution. Macaca irus laetus: H. C. Robinson, 1919, p. 325 — zoogeography. Macaca irus laeta: Chasen, 1940a, p. 69— geographic distribution. Macaca fascicularis laeta: Medway, 1966, p. 16— natural history. P. H. Napier, 1981, pp. 13, 18— external char- acters; holotype cataloged. M[acaca]f[ascicularis] laeti.J. R. Napier & Napier, 1967, pp. 349, 403— geographic distribution. Pithecus dollmani Elliot, 1909, p. 256— holotype, bm(nh) 1909.4.1.20 (Coll. No. 1065), late subadult male, skin and skull, collected at Changi, Singapore I., by H. C. Robinson and E. Seimund, 22 Jul. 1908 (cf. P. H. Napier, 1981, p. 15). H. C Robinson & Kloss, 1914, p. 393— type history. Kloss, 1919c, p. 347— taxonomic comparison. Chasen, 1924b, p. 59— taxonomic com- parison. Chasen, 1940a, p. 66— cited as a synonym of Macaca irus irus F. Cuvier (I. Geoffroy, 1826). P. H. Napier, 1981, pp. 13, 15— cited as a synonym of Ma- caca fascicularis fascicularis (Raffles, [ 1 82 1]); holotype cataloged. [Macacus] dollmani: Raven, 1935, p. 236— geographic distribution. [Macaca irus] dollmani: Sody, 1949, table 1— external and cranial characters. Pithecus bintangensis Elliot, 1909, p. 257— holotype, bm(nh) 1909.4.1.23 (Coll. No. 812), adult male, skin and skull, collected at Sungei Biru, P. Bintan, Kepu- lauan Riau, Indonesia, by H. C. Robinson, date un- specified, presumably Jun. 1908; paratypes, bm(nh) 1909.4.1.24 (Coll. No. 780, adult female, Sungei Biru, P. Bintan, 1 1 Jun.), bm(nh) 1 909.4. 1 .25 (Coll. No. 784, subadult female, Sungei Biru, P. Bintan, 12 Jun.), bm(nh) 1 909.4. 1 .26 (Coll. No. 746, adult female, Pasir Panjang, P. Bintan, 8 Jun.), and bm(nh) 1909.4.1.29 (Coll. No. 870, adult female, Tanjong Sauh, P. Batam, 10 Jun.), skins and skulls, collected in Kepulauan Riau, Indonesia, by H. C. Robinson and/or E. Seimund, 1908. Elliot, 1913, p. 246— type series information. Robinson & Kloss, 1914, pp. 393, 394— type history; taxonomic comparison. H. C. Robinson, 1916, p. 62— cited as a synonym of Pithecus fascicularis (Raffles, [1821]). Chasen, 1924b, p. 59— cited as a synonym of Macaca irus F. Cuvier (I. Geoffroy, 1826). Chasen, 1940a, p. 66— cited as a synonym of Macaca irus irus F. Cuvier (I. Geoffroy, 1826). P. H. Napier, 1981, pp. 13, 15— cited as a synonym of Macaca fascicularis fascicularis (Raffles, [1821]); type history. [Macacus] bintangensis: Raven, 1935, p. 236— geo- graphic distribution. M[acaca] i[rus] bintangensis: Dammerman, 1926b, p. 3 1 6 —geographic distribution. M[acaca] f[ascicularis] bintangensis: Groves & Weitzel in Weitzel et al., 1988, pp. 5, 96, 99-external char- acters; geographic distribution. Macaca mordaxThomas & Wroughton, 1909c, p. 380— holotype, bm(nh) 1909.1.5.27 (Coll. No. 613), sub- adult male, skin and skull, collected at Tjilatjap (= Cilacap), sea level, W Java, Indonesia, by G. C. Shor- tridge, 19 Oct. 1907; paratype, bm(nh) 1909.1.5.28 (Coll. No. 783, juvenile female, skin and skull), col- lected at Cilacap, sea level, W Java, Indonesia, by G. C. Shortridge, 12 Nov. 1907 (cf. Thomas & Wrough- ton, 1909a, p. 373). Miller, 1933, p. 8 -possible phys- iological differentiation. [Macacus] mordax: Raven, 1935, p. 236— geographic distribution. Pithecus mordax: Elliot, 1913, p. 232— geographic dis- tribution; external and cranial characters. P[ithecus]f[ascicularis] mordax: Schwarz, 1913, p. 296— taxonomic comparison. Macaca fascicularis mordax: Hooijer, 1962b, p. 46— cranial characters; taxonomic comparisons. P. H. Na- pier, 1981, pp. 13, 18— external characters; holotype cataloged. Macaca irus mordax: Dammerman, 1928, p. 300— spec- imens collected in Sumba; taxonomic comparisons. Sody, 1929, p. 165— included in fauna of Java. Sody, 1933, p. 93— specimens collected in Bali; taxonomic comparisons. Mertens, 1936, p. 315 — specimens col- lected in Bali, Lombok, Sumbawa, and Flores; taxo- nomic comparisons. Sody, 1949, p. 131— specimens collected in Java; taxonomic comparisons. Macaca resima Thomas & Wroughton, 1909c, p. 381 — holotype, bm(nh) 1909.1.5.31 (Coll. No. 1219), late subadult male, skin and skull, collected at Tasikmalaja (= Tasikmalaya), 1 145 ft (= 350 m), Preanger (region), W Java, Indonesia, by G. C. Shortridge, 18 Jan. 1908. Dammerman in Chasen, 1940a, pp. 68, 70— cited as a synonym of Macaca irus mordax Thomas & Wroughton, 1909c. P. H. Napier, 1981, pp. 13, 18- cited as a synonym of Macaca fascicularis mordax Thomas & Wroughton, 1 909c; holotype cataloged. [Macacus] resimus: Raven, 1935, p. 237— geographic distribution. Pithecus resimus: Elliot, 1 9 1 3, p. 224— external and cra- nial characters; taxonomic comparison. Pithecus lapsus Elliot, 1910, p. 343— holotype, usnm 124863 (Coll. No. 3418), adult male, skin and skull, collected at Tanjung Pamuja, P. Bangka, Indonesia, by W. L. Abbott, 1 9 Jun. 1 904; paratypes, usnm 1 24969 (Coll. No. 35 19, adult male, skin and skull fragments, 1 9 Jul.) and usnm 1 24970 (Coll. No. 352 1 , adult male, skin and skull, 20 Jul.), collected at Tanjung Batu, P. Belitung, Indonesia, by W. L. Abbott, 1904. Chasen, 1940a, p. 66— cited as a synonym of Macaca irus irus F. Cuvier, 1818 (I. Geoffroy, 1826). Poole & Schantz, 1942, p. 243 -holotype cataloged. P. H. Napier, 1981, p. 13— cited as a synonym of M. fascicularis fascicu- laris (Raffles, [1821]). [Macacus] lapsus: Raven, 1935, p. 236— geographic dis- tribution. Macaca irus lapsus: Sody, 1937, p. 247— specimens col- lected in P. Bangka; external and cranial characters; taxonomic comparison. 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 in P. Tuangku, Kepulauan Banyak, Indo- nesia, by W. L. Abbott, 26 Jan. 1 902; paratypes, usnm 1 14408 (Coll. No. 1464, late subadult male, skin and skull, 24 Jan.), usnm 111410 (Coll. No. 1472, adult male, skin and skull, 26 Jan.), usnm 1 1 44 1 1 (Coll. No. ?, subadult male, skull only, 28 Jan.), and usnm 1 14643 (Coll. No. ?, late juvenile male, skull only, 25 Jan.), collected in P. Tuangku, Kepulauan Banyak, Indo- nesia by W. L. Abbott, 1902. Chasen, 1940a, p. 66- cited as a synonym of Macaca irus irus F. Cuvier, 1818 (I. Geoffroy, 1826). Poole & Schantz, 1942, p. 241- holotype cataloged. P. H. Napier, 1981, p. 13— cited as a synonym of M. fascicularis fascicularis (Raffles, [1821]). [Macacus] agnatus: Raven, 1935, p. 236— geographic distribution. [Macaca irus] agnatus: Sody, 1949, Table 1— external and cranial characters. Pithecus lingungensis Elliot, 1910, p. 344— holotype, usnm 104853 (Coll. No. 492), adult male, skin and skull, collected in Pulo Lingung (= P. Lagong), Ke- pulauan Natuna, Indonesia, by W. L. Abbott, 19 Jun. 1900. Chasen, 1940a, p. 69— cited as a synonym of Macaca irus pumila Miller, 1 900. Poole & Schantz, 1942, p. 243-holotype cataloged. P. H. Napier, 1981, p. 14— cited as a synonym of M. fascicularis pumila Miller, 1900. [Macacus] lingungensis: Raven, 1935, p. 236— geo- graphic distribution. [Macaca irus] lingungensis: Sody, 1949, Table 1— ex- ternal and cranial characters. M[acaca] /[ascicularis] lingungensis: Groves & Weitzel in Weitzel et al., 1988, pp. 5, 96, 99— external char- acters; geographic distribution. Pithecus lautensis Elliot, 1910, p. 345— holotype, usnm 104854 (Coll. No. 614), adult male, skin and skull, collected in P. Laut, Kepulauan Natuna, Indonesia, by W. L. Abbott, 9 Aug. 1900. Chasen, 1940a, p. 69- cited as a synonym of Macaca irus pumila Miller, 1900. Poole & Schantz, 1942, p. 243— holotype cat- aloged. P. H. Napier, 1 98 1 , p. 1 4— cited as a synonym of M. fascicularis pumila Miller, 1900. [Macacus] lautensis: Raven, 1935, p. 236— geographic distribution. [Macaca irus] lautensis: Sody, 1949, Table 1— external and cranial characters. Pithecus sirhassensis Elliot, 1 9 1 0, p. 345 — holotype, usnm 104852 (Coll. No. 468), subadult male, skin and skel- eton, collected in Sirhassen I. (= P. Serasan), Kepu- lauan Natuna, Indonesia, by W. L. Abbott, 8 Jun. 1900. Chasen, 1940a, p. 69— cited as a synonym of Macaca irus pumila Miller, 1 900. Poole & Schantz, 1942, p. 245 -holotype cataloged. P. H. Napier, 1981, p. 14— cited as a synonym of M. fascicularis pumila Miller, 1900. [Macacus] sirhassensis: Raven, 1935, p. 237— geograph- ic distribution. [Macaca irus] sirhassensis: Sody, 1949, Table 1— exter- nal and cranial characters. M[acaca] f[ascicularis] sirhassensis: Groves & Weitzel in Weitzel et al., 1988, pp. 5, 96, 99— external char- acters; geographic distribution. Pithecus carimataeEWiot, 1910, p. 346— holotype, usnm 125101 (Coll. No. 3646), adult male, skin and skull, collected at Teluk Pai, P. Karimata, Indonesia, by W. L. Abbott, 24 Aug. 1 904; paratype (available but not explicitly cited in original description), usnm 125102 (Coll. No. 3661), adult male, skin and skull, collected at Teluk Pai, P. Karimata, Indonesia, by W. L. Abbott, 27 Aug. 1904. Lyon, 1911, p. 137-type history. Chas- en, 1935b, p. 2— taxonomic comparison. Chasen, 1940a, p. 66— cited as a synonym of Macaca irus irus F. Cuvier, 1818 (I. Geoffroy, 1826). Poole & Schantz, 1942, p. 242— holotype cataloged. P. H. Napier, 1981, p. 13— cited as a synonym of M. fascicularis fascicu- laris (Raffles, [1821]). [Macacus] carimatae: Raven, 1935, p. 236— geographic distribution. [Macaca irus] carimatae: Sody, 1949, Table 1 —external and cranial characters. Macaca irus? carimatae: Sody, 1 949, p. 1 3 1 —specimens collected in P. Pelapis Tengah and P. Serutu, Kepu- lauan Karimata, Indonesia. Pithecus mandibularis Elliot, 1910, p. 347— holotype, usnm 142225 (Coll. No. 4196), late subadult male, skin and skull, collected at Sungei Sama (= Sungai Ambawang), near Pontianak, Kalimantan, Indonesia, by W. L.Abbott, 18 Jun. 1905. Lyon, 1911, p. 137- type history. Chasen & Kloss, 1931, p. 10— said to be based on individual variables. Poole & Schantz, 1 942, p. 244— holotype cataloged. P. H. Napier, 1981, p. 13— cited as a synonym of M '. fascicularis fascicularis (Raffles, [1821]). [Macacus] mandibularis: Raven, 1935, p. 236— geo- graphic distribution. Macaca irus mandibularis: Gyldenstolpe, 1920, pp. 3, 14— specimen collected at Kaboerau, Kalimantan, In- donesia; said to be probably "not [valid] as a distinct race." Sody, 1949, p. 130— taxonomic comparison. [Macaca irus] mandibula: Sody, 1949, Table 1— incor- rect spelling of Pithecus mandibularis Elliot, 1910: ex- ternal and cranial characters. M[acaca]f [ascicularis] mandibularis: Groves & Weitzel in Weitzel et al., 1988, pp. 5, 96, 99— external char- acters; geographic distribution. Pithecus baweanus Elliot, 1910, p. 347 — holotype, usnm 151829 (Coll. No. 5565), adult male, skin and skull, collected in P. Bawean, Java Sea, Indonesia, by W. L. Abbott, 24 Nov. 1907; paratype (available but not explicitly cited in original description), usnm 151830 (Coll. No. 5566), adult female, skin and skull, collected in P. Bawean, Indonesia, by W. L. Abbott, 24 Nov. 1907. Lyon, 1911, p. 137-type history. [Macacus] baweanus: Raven, 1935, p. 236— geographic distribution. Macaca irus baweana: Chasen, 1940a, p. 70— geographic distribution. Poole & Schantz, 1 942, p. 24 1 — holotype cataloged. Macaca irus baweanus: Sody, 1949, p. 132 — specimens collected in P. Bawean; taxonomic comparison. M[acaca]f [ascicularis] baweana: J . R. Napier & Napier, 1967, pp. 349, 403— geographic distribution. Pithecus cupidus Elliot, 1910, p. 348— holotype, usnm 151831 (Coll. No. 5584), adult male, skin and skull, collected in P. Matasiri, Java Sea, Indonesia, by W. L. Abbott, 8 Dec. 1907; paratype (available but not explicitly cited in original description), usnm 154368 (Coll. No. 6388), adult male, cranium only, collected 74 FIELDIANA: ZOOLOGY in P. Matasiri, Indonesia, by W. L. Abbott, presum- ably 1907-1908. Lyon, 1911, p. 137— type history. [Macacus] cupidus: Raven, 1935, p. 236— geographic distribution. Macaca irus cupida: Chasen, 1940a, p. 70— geographic distribution. Poole & Schantz, 1942, p. 242— holotype cataloged. [Macaca irus] cupidus: Sody, 1949, Table 1— external and cranial characters. M[acaca /[ascicularis] cupida: J. R. Napier & Napier, 1967, pp. 349, 403— geographic distribution. Pithecus lingae Elliot, 1910, p. 349— holotype, usnm 101603 (Coll. No. ?), subadult male, skin and skull, collected in Linga I. (= P. Lingga), Kepulauan Lingga, Indonesia, by W. L. Abbott, 23 Jul. 1899; paratype, usnm 101602 (Coll. No. ?), late juvenile male, skin and skull, collected in P. Lingga, Indonesia, by W. L. Abbott, 23 Jul. 1899. Chasen, 1925, p. 93 -cited as a synonym of Macaca irus F. Cuvier (I. Geoffroy, 1 826). Chasen, 1940a, p. 66— cited as a synonym of Macaca irus irus F. Cuvier, 1818 (I. GeofTroy, 1826). Poole & Schantz, 1942, p. 243— holotype cataloged. P. H. Na- pier, 1981, p. 13— cited as a synonym of M. /ascicu- laris f ascicularis (Raffles, [1821]). [Macacus] lingae: Raven, 1935, p. 236— geographic dis- tribution. M[acaca] i[rus] lingae: Dammerman, 1926b, p. 316 — geographic distribution. Pithecus impudens Elliot, 1910, p. 350— holotype, usnm 115675 (Coll. No. 1956), adult male, skin and skull, collected in P. Sugi, Kepulauan Riau, Indonesia, by W. L. Abbott, 24 Aug. 1902. Chasen, 1925, p. 93- cited as a synonym of Macaca irus F. Cuvier (I. Geof- froy, 1826). Chasen, 1940a, p. 66— cited as a synonym of Macaca irus irus F. Cuvier, 1 8 1 8 (I. Geoffroy, 1826). Poole & Schantz, 1942, p. 242— holotype cataloged. P. H. Napier, 1981, p. 13— cited as a synonym of M. /ascicularis /ascicularis (Raffles, [1821]). M[acaca] impudens: Raven, 1935, p. 236— geographic distribution. M[acaca] i[rus] impudens: Dammerman, 1926b, p. 316— geographic distribution. Macaca irus? impudens: Sody, 1949, p. 129— specimens collected in P. Durian, Kepulauan Riau, Indonesia. Pithecus capitalis Elliot, 1910, p. 350— holotype, usnm 83271 (Coll. No. ? ), adult male, skin and skull, col- lected at Trong (= Ban Phra Muang, Trang Province), peninsular Thailand, by W. L. Abbott, 7 Mar. 1896; paratype cited in original description, usnm 83272 (Coll. No. ?, adult male, skin and skull), collected in Telibon I. [= Ko Telibong], Thailand, by W. L. Ab- bott, 27 Feb. 1896; paratypes available but not ex- plicitly cited in original description, usnm 83273 (Coll. No. ?, late juvenile male, skin and skull, Ban Phra Muang, 8 Mar.), usnm 83274 (Coll. No. 122, adult female, skin and skull, Tyching, 19 May), and usnm 83275 (Coll. No. ?, infant, skin and skull, Tyching, 19 May), collected in Trang Province, peninsular Thai- land, by W. L. Abbott, 1896. Kloss, 1919c, p. 347- taxonomic comparison. Weitzel et al., 1988, p. 109— cited as a synonym ofMacaca/ascicularis bintangensis (Elliot, 1909). Macaca capitalis: Gyldenstolpe, 1919, p. 131— geo- graphic distribution. Macaca irus capitalis: Chasen, 1940a, pp. 68-69— geo- graphic distribution; said to be possibly a junior syn- onym of [Pithecus] vitiis Elliot, 1 9 1 0, p. 346 (= Macaca /ascicularis aurea I. Geoffroy, [1831]). Poole & Schantz, 1942, p. 241— holotype cataloged. M[acaca] /[ascicularis] capitalis: J. R. Napier & Napier, 1967, pp. 349, 403— geographic distribution. Macaca irus argentimembris Kloss, 191 la, p. 1 16— ho- lotype, bm(nh) 1949.426 (Coll. No. 3814, SM [= Se- langor Museum] 2068/10), adult male, skin and skull, collected in P. Pinang [2], West Malaysia, by C. B. Kloss, 3 Sep. 1910; paratypes, bm(nh) 1955.1523 (Coll. No. 3491, SM 2069/10, adult female, P. Pinang [2], 3 Sep.), zrc 4-054 (Coll. No. 3488, SM 2070/10, late subadult male, P. Redang, 2 Sep.), and zrc 4-055 (Coll. No. 3833, SM 2071/10, latejuvenile male, P. Redang, 5 Sep.), skins and skulls, collected in West Malaysia by C. B. Kloss, 1910. J. E. Hill, 1960, p. 31 -type history. Weitzel et al., 1988, p. 109— cited as a syn- onym of M. /ascicularis bintangensis (Elliot, 1 909). [Macacus] irus argentimembris: Raven, 1935, p. 236— geographic distribution. Macaca /ascicularis argentimembra: Kloss, 1911a, pp. 177, 181 —external and cranial characters; taxonomic comparison; type series information. Macaca /ascicularis argentimembris: J. R. Napier & Na- pier, 1967, pp. 349, 403— geographic distribution. P. H. Napier, 1981, pp. 13, 16— external characters; ho- lotype cataloged. Macaca [nemestrinus-Gruppe] aff. adusta: Elbert, 1912, p. 101 (not Miller, 1906a, p. 559)— misidentification of specimens collected in P. Sumbawa. Schwarz, 1912, p. 304— misidentification of specimens collected in P. Sumbawa. Pithecus /ascicularis aff. limitis Schwarz, 1912, p. 304— nomen nudum. Pithecus /ascicularis limitis Schwarz, 1913, p. 296— ho- lotype, zsbs 1 9 1 1 /2 1 04 (Coll. No. 1 9), adult male, skin and skull, collected at Lelogama, P. Timor, Indonesia, by C. B. Haniel, 28 May 1911; paratypes, zsbs 1911/ 2101 (Coll. No. 15, juvenile male, skin and skull, Fatu Timau, NE, 1200 m, 20 May), zsbs 191 1/2102 (Coll. No. 16, adult female, skin and skull, Lelogama, 22 May), zsbs 191 1/2103 (Coll. No. 17, adult male, skin only, Lelogama, 27 May), zsbs 1911/2105 (Coll. No. 21, infant male, skin and skull, Lelogama, 29 May), zsbs 1 9 1 1/2 1 06 (Coll. No. 22, juvenile male, skin only, Lelogama, 30 May), zsbs 1911/2107 (Coll. No. 24, infant male, skin only, Lelogama, 3 Jun.), zsbs 1911/ 21082 (Coll. No. 63, adult male, skin and skull, Kuat- nana, 300 m, 20 Jul.), zsbs 191 1/2109 (Coll. No. 69, adult male, skull only, Bokong, 180 m, 26 Jul.), and zsbs 1911/2110 (Coll. No. 68, adult male, skin only, Bokong, 180 m, 24 Jul.) collected in P. Timor, In- donesia, by C. B. Haniel, 1911. Schwarz, 1914, p. 117— type series information. M[acaca] /[ascicularis] limitis: J. R. Napier & Napier, 1967, pp. 349, 404— geographic distribution. [Macacus] irus limitis: Raven, 1935, p. 236— geographic distribution. Macaca irus limitis: Kellogg, 1945, p. 119— geographic distribution. Macaca irus limitus: Sody, 1949, p. 133, Table 1— in- correct spelling of Pithecus /ascicularis limitis Schwarz, 1913; taxonomic comparison. Pithecus mansalaris Lyon, 1916, p. 452— holotype, usnm FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 75 114560 (Coll. No. 1639), adult male, skin and skull, collected in Pulo Mansalar (= P. Mursala), west of Sumatra, Indonesia, by W. L. Abbott, 10 Mar. 1902; paratypes, usnm 1 14559 (Coll. No. 1624, adult male, skin and skull, 7 Mar.) and usnm 1 14561 (Coll. No. ?, adult female, skull only, 9 Mar.), collected in P. Mursala, Indonesia, by W. L. Abbott, 1902. Chasen, 1940a, p. 66— cited as a synonym of Macaca irus irus F. Cuvier, 1818 (I. Geoffroy, 1826). Poole & Schantz, 1942, p. 244— holotype cataloged. P. H.Napier, 1981, p. 13— cited as a synonym of Macaca fascicularis fas- cicularis (Raffles, [1821]). [Macacus] mansalaris: Raven, 1935, p. 236— geographic distribution. [Macaca irus] mansalaris: Sody, 1949, Table 1 —external and cranial characters. Pithecus sp.: Thomas, 1928, p. 832 — misidentification of juvenile male collected at Tay Ninh, Vietnam. Macaca mulatta: Thomas, 1928, p. 832 (not Zimmer- mann, 1780, p. 195)— misidentification of specimens (Coll. Nos. 777, 778) collected at Phu Quoc (not Phu- Qui), Vietnam. Macaca irus sublimitis Sody, 1932, p. 338— based on five specimens, unseen by Sody, reported by Dam- merman (1928, p. 300) as Macaca irus mordax Tho- mas & Wroughton, 1909c; type series, mzb 865 (adult female, Kambera [= Payeti-Kambaniru], 26 Mar.), mzb 866 (adult male, Kambera, 27 Mar.), mzb 867 (juvenile male, Mao Marroe, 4 May), mzb 868 (late juvenile female, Mao Marroe, 8 May), and mzb 869 (subadult female, Mao Marroe, 10 May), collected in P. Sumba, Indonesia, by P. F. Franck and/or Denin, 1925. [Macaca irus] sumbae: Sody, 1933, p. 94— lapsus for Macaca irus sublimitis Sody, 1932. Macaca irus sublimitus: Sody, 1 949, p. 1 34, Table 1 — incorrect spelling of Macaca irus sublimitis Sody, 1 932; taxonomic comparisons, distinctiveness of subspecies "still waits for thorough confirmation"; geographic distribution; type series information. W. C. O. Hill, 1974, p. 529— recognized provisionally, said to be probably a synonym of Macaca irus mordax Thomas & Wroughton, 1909c. M[acaca] /[ascicularis] sublimitis: J. R. Napier & Na- pier, 1967, pp. 349, 404— geographic distribution. Macaca irus submordax Sody , 1949, p. 133, Table 1 — holotype, rmnh unnumbered (Coll. No. E85), adult male, skin and skull, collected at Desa Poetjang, G. Agoeng (= Gunung [Mountain] Agung), E Bali, In- donesia, by H. J. V. Sody, date unspecified, apparently 1927-1931; paratypes, rmnh (Coll. No. E34, juvenile male, skin and skull, Sendang), rmnh (Coll. No. E64, adult male, skin and skull, Sendang), rmnh (Coll. No. E74, adult male, skin and skull, Sendang), rmnh (Coll. No. El 36, adult male, skull only, Jembrana, Negara district), and rmnh (Coll. No. El 39, subadult female, skin and skull, Jembrana, Negara district), collected in Bali, Indonesia, by H. J. V. Sody, apparently 1927— 1931; mzb 2001 (Coll. No. 1), adult female, skin and skull, collected at Batu Meringgit, Bali, Indonesia, by P. F. Franck, 8 Oct. 1928; mzb 6521 (Coll. No. 17/ 92/33, juvenile male), mzb 6522 (Coll. No. 15/93/33, adult male), and mzb 6523 (Coll. No. 16/04/33, adult male), skins and skulls, collected at Banju Wetan, Bali, Indonesia, by J. J. Menden, 18 Jul. 1933; and one unlocated additional female specimen cited by Sody in Table 1. W. C. O. Hill, 1974, p. 525 -cited as a synonym of Macaca irus mordax Thomas & Wrough- ton, 1909c. P. H. Napier, 1981, p. 13 -cited as a syn- onym of Macaca fascicularis mordax Thomas & Wroughton, 1909c. Simia mauritius: W. C. O. Hill, 1974, p. 507 (not Grif- fith, 1821, p. 58; an "entirely black" monkey)— cited as a synonym ofMacaca irus irus. P. H. Napier, 198 1, p. 13— cited as a synonym of Macaca fascicularis fas- cicularis. Holotype— Not preserved. Simia fascicularis Raffles, [1821] (p. 246) apparently is based on a single specimen, probably an adult female ("The body is about twenty inches long, and the tail a little more. . . . Canines short."). This specimen was part of a collection made in Sumatra for Raf- fles by P. Diard and A. Duvaucel during the period March 1819-March 1820 (T. S. Raffles in S. Raf- fles, 1830, pp. 703, 713); the original description of Simia fascicularis is included in Raffles's pub- lished catalog of this collection. The fate of the holotype is unclear. Part of the Raffles/Diard/Duvaucel Sumatra collection, possibly accompanied by the manu- script of Raffles's catalog, was sent to England in March 1820, and another, larger part was sent to England in or before June 1 820 (T. S. Raffles in S. Raffles, 1830, pp. 440, 447, 453, 715; Watson et al. in S. Raffles, 1830, p. 716). The text of Raf- fles's catalog was read at a meeting of the Linnaean Society of London on 5 December 1820. Three specimens of M. fascicularis (sexes unspecified) sent by Raffles from Sumatra ultimately reached the museum of the Zoological Society of London (Waterhouse, 1838, p. 7; cf. Vigors, 1830, p. 642), and two additional specimens sent by Raffles from Sumatra or Java reached the museum of the East India Company, London (Horsfield, 1851, pp. iii- iv, 17). When these two museums were disband- ed—the former in 1852-1860 and the latter in 1879— their collections were transferred to the bm(nh) (Thomas, 1906, pp. 40, 63), but none of Raffles's specimens of M. fascicularis are now pre- served in the bm(nh) (cf. P. H. Napier, 1981, p. 16). Another part of the Sumatra collection was transferred by Raffles to Diard and Duvaucel and subsequently was sent to France by Duvaucel in 1820-1821 (T. S. Raffles in S. Raffles, 1830, p. 720; Lacaze, 1856, col. 536). Included in this part of the collection was an adult female specimen of M. fascicularis (mnhn 362/233), which coinciden- 76 FIELDIANA: ZOOLOGY tally is a paralectotype of M. f. aurea I. Geoffroy, [1831] (see below). This female, however, almost certainly is not the holotype of Simla fascicularis; Raffles considered the specimens that he trans- ferred to Diard and Duvaucel to be "duplicates," not the "originals," which he cataloged and sent to England. Type Locality— Sumatra, restricted to "neigh- bourhood of Bencoolen [= Bengkulu]" by H. C. Robinson (1 9 1 6, p. 63). Bengkulu, in southwestern Sumatra, was the headquarters of Raffles, Diard, and Duvaucel (cf. Steenis-Kruseman, 1 950, p. 425). Distribution (Fig. 25)— The broad geographic range of M. f. fascicularis in Southeast Asia in- cludes the following continental and insular com- ponents: the southern part of the Indochinese Pen- insula (including southern Vietnam, Cambodia, and southern Thailand) and adjacent islands (ex- cluding the Con Son group and Ko Khram Yai); 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- wai, and P. Enganno); Borneo and adjacent islands (excluding P. Maratua), Java and adjacent islands (excluding P. Karimunjawa and P. Kemujan); Lesser Sunda Islands from Bali to Timor (exclud- ing P. Komodo and many nearby islets); and the south-central Philippines (including Cagayan Sulu, Sulu Archipelago, and western Mindanao). At the margins of the subspecific range are three inter- subspecific or interspecific contact zones, inhab- ited by mixed-phenotype populations, as follows: (1) central and eastern parts of the Indochinese Peninsula— an intersubspecific contact zone be- tween ranges of M. f. fascicularis and M.f aurea, or an interspecific contact zone between ranges of M. fascicularis and M. mulatta; (2) eastern and southern parts of the Isthmus of Kra and adjacent southwestern islands— an intersubspecific contact zone between ranges of M. f. fascicularis and M. f aurea; and (3) eastern and central Mindanao, southern Negros, and possibly nearby islands— an intersubspecific contact zone between ranges of M. f. fascicularis and M. f philippinensis (see below, M. f. aurea— Distribution; M. f. philippinensis— Distribution). diagnosis— General color of dorsal surface of trunk buffy to yellowish gray to medium brown (825 of 868 postinfantile specimens examined), occasionally dark brown (43 specimens; cf. Table 1 , Appendix 3); erythrism variable, pale hair an- nulations pale yellowish to golden to rufescent (cf. Table 3, Appendix 4); crown same color as back, or brighter (847 of 873 postinfantile specimens examined), occasionally with an indistinct black- ish wash (24 specimens), rarely contrastingly darker than back (2 specimens; cf. Table 5, Appendix 5); preauricular hairs directed anteriorly, forming part of lateral facial crest (crest transzygomatic, 501 of 504 specimens examined), rarely directed poste- riorly, partly covering ears (crest infrazygomatic, 2 specimens; crest asymmetric, 1 specimen; cf. Table 6, Appendix 6); T 67-150% of HB in 311 adult specimens examined (cf. Table 10, Appendix 9). Remarks— For comments on nine quasi-dis- tinct populations that are here included within M. f fascicularis, see above (Subspecific Taxonomy). Specimens Examined— Total, 1,550: skins and skulls, 739; skins only, 285; skulls only, 526 (see Appendix 1). Macaca fascicularis aurea I. Geoffroy, [1831], p. 58 Macacus aureus I. Geoffroy, [1831], pp. 58, 76, pi. 2— for details concerning type series, see below. Eydoux & Souleyet [& Gervais], 1841, p. 6, pi. 2— taxonomic comparisons. I. Geoffroy, [1843], p. 566— taxonomic history. Gray, 1849, p. 4— cited as a synonym of Ma- cacus cynomolgus [= Macaca fascicularis]. I. Geoffroy, 1851, p. 27— taxonomic comparison; type series in- formation. Blyth, 1875, p. 7— cited as a synonym of Macacus cynomolgus [= Macaca fascicularis]; said to be based on "a casual individual variety from Pegu." J. Anderson, 1879, p. 73— cited as a synonym of Ma- cacus cynomolgus [= Macaca fascicularis]; type said to be probably a market specimen. Elliot, 1913, p. 229— cited as a synonym of Pit hecus irus [= Macaca fascicularis]. Rode, 1938, p. 223— cited as a synonym of Macaca irus (F. Cuvier, 1818; I. Geoffroy, 1826); holotype cataloged. Macacus auratus Miiller, [1840], p. 49 (not E. Geoffroy, 1812, p. 93)— incorrect spelling of Macacus aureus I. Geoffroy, [1831]. Blyth, 1863, p. 9-cited as a syn- onym of Macacus carbonarius F. Cuvier (I. Geoffroy, 1826). I[nuus] aureus: Wagner, [1839], p. 138 (part)— external characters. Pith[ecus] (Mac[acus]) aureus: Dahlbom, 1856, p. 118 — external characters. Macaca irus aurea: Pocock, 1939, p. 79— taxonomic comparison; type locality restricted to Pegu. [Macaca irus] aureus: Sody, 1 949, Table 1 —external and cranial measurements. M[acaca] f[ascicularis] aurea: J. R. Napier & Napier, 1967, pp. 349, 403— distribution. Cercopithecus Cynosurus: Heifer, 1838, p. 858 (not Sco- poli, 1786, p. 44)— misidentification; natural history. FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 77 M[acacus] cynomolgus(?): Blyth, 1844, pp. 474, 476 (not Linnaeus, 1766, p. 38)— specimens collected in Ara- kan. Macacus cynomolgus: Blyth, 1875, p. 7 (part; not Lin- naeus, 1766)— taxonomic comparisons. M\acacus] carbonarius: Blyth, 1847, p. 732 (not I. Geof- froy, 1826)— said to be common in Arakan. C\ynamolgus] mulatta: Reichenbach, 1862, p. 136 (part; not Zimmermann, 1780, p. 195)— misidentification; taxonomic history. Pithecus vitiis Elliot, 1910, p. 346— holotype, usnm 124176 (Coll. No. 3076), subadult male, skin and mandible (cranium lost Jan. 1938), collected at Domel I. (= Letsok-aw Kyun), Mergui Archipelago, Burma, by W. L. Abbott, 26 Jan. 1904; paratypes, usnm 104442 (Coll. No. 287, late juvenile male, Sullivan's I. [= Lanbi Kyun], 30 Jan.) and usnm 111898 (Coll. No. 757, adult female, St. Matthews I. [= Zadetkyi Kyun], 9 Dec), skins and skulls, collected in Mergui Archi- pelago, Burma, by W. L. Abbott, 1900. Pocock, 1939, p. 79— cited as a synonym of Macaca irus aurea. Chas- en, 1 940a, p. 68 — said to be possibly a senior synonym of Pithecus capitalis Elliot, 1910, p. 350 (= M. fasci- cularisfascicularis (Raffles, [1821 ])). P. H. Napier, 1981, p. 13— cited as a synonym of Macaca fascicular is au- rea. Macaca vitiis: Poole & Schantz, 1942, p. 246 — holotype cataloged. [Macaca irus] vitiis: Sody, 1949, Table 1— external and cranial measurements. Pithecus fascicularis: Wroughton, 1915, p. 699 (part; not Raffles, [1821], p. 246)— taxonomic history. Type Series — Macacus aureus I. Geoffroy, [1831, pp. 58, 76] is explicitly based on four spec- imens of Macaca fascicular is: (1) mnhn 362/234, Type Cat. No. 58a (lectotype, designated by Rode, 1938, p. 223), subadult or adult male, mounted skin only (skull measured by Elliot, 1913, p. 231; reported absent by Rode, 1938, p. 223), obtained in "Bengale" (now partly in eastern India and part- ly in Bangladesh) by L. T. Leschenault de la Tour, presumably in 1819-1820, acquired by the mnhn in Jul. 1822 (cf. Leschenault de la Tour, 1820, p. 359; 1822, p. 262; I. Geoffroy, 1851, p. 27; Rode, 1938, p. 223; Steenis-Kruseman, 1950, p. 321); (2) unpreserved specimen (not listed in catalog of I. Geoffroy, 1 85 1 , p. 27), sex unspecified, obtained in "Pegou" (= Pegu, Burma) by A. A. M. Reynaud, 1827-1828 (cf. de Rossel et al., 1829, p. 603; de Rossel, 1829, p. 609); (3) mnhn 362/233, Type Cat. No. 58b, adult female, mounted skin only, obtained in Sumatra (no further locality infor- mation available) by A. Duvaucel, 1 8 1 9-1 820, ac- quired by the mnhn 7 Sep. 1821 (cf. I. Geoffroy, 1851, p. 27; Rode, 1938, p. 223; de Lacaze, 1856, col. 535); and (4) unpreserved specimen (listed by I. Geoffroy, 1851, p. 29; not listed by Rode, 1938, p. 223), male, skin ("poils uses," I. Geoffroy, [ 1 843], p. 567) with or without skull, obtained in Java (no further locality information available), by P. Diard, 1821. I. Geoffroy ([1831], pp. 58, 76) indicated that dorsal pelage in all four of these specimens was conspicuously erythristic ("Dessus du corps d'un beau roux tiquete de noir") and cited this as the primary diagnostic character of Macacus au- reus, "Le Macaque Roux Dore." However, the infrazygomatic lateral facial crest (see above, Pel- age) that is now regarded as diagnostic of Macaca fascicularis aurea is clearly figured by I. Geoffroy ([1831], PI. 2) in his illustration of either the "Ben- gale" or Pegu specimen and is alluded to in his detailed description of the illustrated specimen ("longs poils blancs diriges en arriere . . . cachent en partie les oreilles," p. 77). No holotype is designated in the original de- scription of Macacus aureus I. Geoffroy, [1831]. In I. Geoffroy's (1851, p. 27) catalog of primates in the mhnh, the "Bengale" specimens ("L 'un des types de respece") and the Sumatra specimen {"L'un des types") are treated as coordinate type specimens of this taxon; the Java specimen is transferred to another taxon (p. 29), and the Pegu specimen is not mentioned. Ambiguous state- ments by J. Anderson (1879, p. 76) and Elliot (1913, p. 231) may or may not constitute desig- nations of the "Bengale" specimen as lectotype. The "Bengale" specimen is decisively designated as lectotype by Rode (1938, p. 223; cf. Interna- tional Code of Zoological Nomenclature, 1985, Article 74). Type Locality— Pegu, Burma (restricted by Pocock, 1 939, p. 79; cf. International Code of Zoo- logical Nomenclature, 1985, Article 72h). The lec- totype of Macacus aureus I. Geoffroy, [183 1], ob- tained in "Bengale," probably was a captive pur- chased in Calcutta (J. Anderson, 1879, p. 76); sim- ilar captive specimens in the Calcutta market, or nearby, have been reported by C. Belanger (in I. Geoffroy, [1831], p. 77), Eydoux and Souleyet [& Gervais] (1841, pp. xiv, 6), and J. Anderson (1879, p. 76). The original provenance of the lectotype is unknown. Based on the locality of one of the para- lectotypes (see above), Pocock (1939, p. 79) amended the type locality to Pegu. Distribution (Figs. 9, 25) — Southernmost Bangladesh, southwestern and southern Burma (including Preparis I.), adjacent islands in the Mer- gui Archipelago, and a small area in west-central Thailand; most of the geographic range is west of the mountain chains that form the border between 78 FIELDIANA: ZOOLOGY Burma and Thailand. Adjacent to the geographic range of M.f. aurea are two areas— one in central and eastern parts of the Indochinese Peninsula and the other in eastern and southern parts of the Isth- mus of Kra— where specimens with the diagnostic lateral facial crest character of M. f. aurea and specimens with the diagnostic lateral facial crest character of M. f. fascicularis have been collected, sometimes at the same locality. The area in eastern and southern parts of the Isthmus of Kra may be regarded as a M.f. aurea/ M.f. fascicularis contact zone; the area in central and eastern parts of the Indochinese Peninsula may be either a M.f. aurea/ M.f. fascicularis contact zone or a M. fascicularis/ M. mulatta contact zone (see below, Evolution and Dispersal). Diagnosis— General color of dorsal surface of trunk buffy to medium brown, pale hair annula- tions pale yellowish to golden, rarely rufescent; crown usually brighter than back, occasionally with an indistinct blackish wash; preauricular hairs di- rected posteriorly, partly covering ears (lateral fa- cial crest infrazygomatic); T 76-104% of HB in 12 adult specimens examined. For external and cra- nial measurements, see Table 36. Specimens Examined— M. f aurea, 53: skins and skulls, 42; skins only, 8; skulls only, 3. Inter- subspecific or interspecific contact zones, 45: skins and skulls, 41; skins only, 3; skull only, 1 (see Appendix 1). Macaca fascicularis philippinensis I. Geoffroy, [1843], p. 568 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 from type catalog published by Rode, 1938, p. 222). This monkey was obtained alive in Manila by A. Chenest and presented by him to the me- nagerie of the mnhn on 6 Aug. 1841; it died on 29 Aug. 1 842 (cf. I. Geoffroy, 1 85 1 , p. 29). A figure of the holotype accompanies the original descrip- tion (pi. 5). Type Locality— Philippine Islands, probably Luzon ("Chenest . . . l'a acquis a Manille, et il le croit originaire de cette ile," I. Geoffroy, [1843], p. 570). Distribution (Fig. 25)— Western, northern, and eastern islands of the Philippine Archipelago, in- cluding Balabac, Palawan, Culion, Mindoro, Lu- zon, Samar, Leyte, and probably other islands north of about 10°N (Fooden, 1991, pp. 6, 23). Based on two available skins, northeastern Mindanao formerly was also included in the range of this subspecies (Fooden, 1 99 1 , p. 23); however, in June 1991 one of the critical specimens (bm(nh) 1877.10.6.2, Butuan River [= Agusan River]) was discovered to consist of a late infant or early ju- venile skin mismatched with a subadult female skull, which weakens the evidence for retaining northeastern Mindanao in the subspecific range. In eastern and central Mindanao, southern Ne- gros, and possibly in nearby islands, between the ranges of M. f philippinensis and M.f. fascicularis, is a contact zone inhabited by mixed-phenotype populations. Diagnosis— General color of dorsal surface of trunk dark brown, pale hair annulations golden to refescent (pale golden in 1 subadult male, fmnh 87718); crown brighter than back (indistinct blackish streak in 1 adult male, bm(nh) 1877.10.6.1); preauricular hairs directed anteri- orly, forming part of lateral facial crest (crest transzygomatic); T 101-138% of HB in 14 adult specimens examined. For further details, see Fooden (1991, pp. 3 ff). Specimens Examined— M. f philippinensis, 98: skins and skulls, 53; skins only, 15; skulls only, 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- nensis/ 'M.f fascicularis contact zone, 12: skins and skulls, 2; skin only, 1 ; skulls only, 9 (see Appendix 1). Macaca fascicularis umbrosa Miller, 1902b, p. 789 Cercopithecus cynomolgus: Cantor, 1846, p. 176 (part; not Linnaeus, 1766, p. 38)— Nicobar Islands included in distribution. Macacus cynomolgus: Barbe, 1846, pp. 365 (not Lin- naeus, 1766)— sight records. Blyth, 1846, p. 367 (part) — zoogeography. M[acacus] cynomolgos: Blyth, 1863, p. 9 (part; not G. Cuvier, 1798, p. 98)— specimen collected in "Nico- bars." Cercocebus carbonarius:Ze\eboT, [ 1 869], p. 7 (not I. Geof- froy, 1826, p. 588)— cited as "Var. a" of Inuus cy- nomolgus; specimen collected in Great Nicobar I. 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. Latitude Adult females Adult males HB T/HB GL HB T/HB GL Locality (N) (mm) x 100 (mm) (mm) x 100 (mm) Haungtharaw 16°30' 125.7 Ye Forest 16°10' 108.4 Wong, 65 km E 15°55' 400 96.3 99.0 Wong, 85 km E 15°55' 460 76.1 98.5 Ban Tamrong Phato 14°54' 486 90.7 113.0 529 103.8 128.2 Taungbyauk Chaung 13°45' 116.2 Kathema Kyun 13°39' 126.3 Kadan Kyun 12°30' 520 95.2 129.8 Mergui 12°26' 128.7 Tenasserim 1 12°05' 480 490 512 95.8 94.9 94.7 104.4 114.0 109.1 630 84.9 132.3 Zadetkyi Kyun 9°57' 470 89.4 114.3 Mean 471.1 91.1 108.54 559.7 94.6 128.50 SD 35.4 7.1 6.62 61.1 9.4 2.41 N 7 7 9 3 3 6 1 Summary statistics for female sample: HB, mean = 494.0, SD = 16.3; T/HB x 100, mean = 95.1, SD = 0.6; GL, mean = 109.17, SD = 4.80. not I. Geoffrey, 1 826)— cited as "Smaller var." of Ma- cacus cynomolgus; external characters of specimen collected in Nicobar Islands. Macacus umbrosus Miller, 1902b, p. 789 — for details concerning type series, see below. Lyon & Osgood, 1909, p. 285— holotype cataloged. Macaca umbrosa: Miller, 1933, p. 8— possible physio- logical differentiation. Poole & Schantz, 1942, p. 246— holotype cataloged. [Macacus] umbrosa: Raven, 1935, p. 237— geographic distribution. [Cynomolgus] umbrosus: Trouessart, 1904, p. 16— geo- graphic distribution. Pithecus umbrosus: Elliot, 1913, p. 228— external and cranial characters. [Silenus] umbrosus: Stiles & Nolan, 1929, p. 537 — par- asites. Macaca irus umbrosa: Pocock, 1 939, p. 82— external and cranial characters. Kellogg, 1944, p. 76— taxonomic comparisons; zoogeography. M[acaca]f[ascicularis] umbrosa: J. R. Napier & Napier, 1967, pp. 349, 402— geographic distribution. Type Series— Holotype (by original designa- tion), usnm 111795 (Coll. No. 888), adult male, skin and skull, collected in Little Nicobar I., Ni- cobars, India, by W. L. Abbott, 25 Feb. 1901; paratypes, usnm 1 1 1796 (Coll. No. 889, subadult male, Little Nicobar I., 26 Feb.), usnm 111797 (Coll. No. 893, adult male, Little Nicobar I., 27 Feb.), usnm 1 1 1 792 (Coll. No. 9 1 8, subadult male, Great Nicobar I., 8 Mar.), usnm 111793 (Coll. No. 929, subadult female, Great Nicobar I., 12 Mar.), usnm 111799 (Coll. No. 939, subadult female, Great Nicobar I., 23 Mar.), usnm 111801 (Coll. No. 886, adult male, Katchall I., 21 Feb.), and usnm 111802 (Coll. No. 887, juvenile male, Katchall I., 21 Feb.), skins and skulls, collected in Nicobars, India, by W. L. Abbott, 1901. Type Locality— Little Nicobar I., Nicobars, India. Distribution (Fig. 25)— Katchall I., Little Nic- obar I., and Great Nicobar I., Nicobars, India. Kloss (1903a, p. 114) reported that monkeys are absent in the Nicobars north of Katchall I. Diagnosis— General color of dorsal surface of trunk blackish (dark brownish gray in 1 adult male, usnm 1 1 1801), pale hair annulations pale yellow- ish; crown yellowish brown, hairs conspicuously annulated with pale yellowish; preauricular hairs directed anteriorly, forming part of lateral facial crest (crest transzygomatic, 5 of 8 specimens ex- amined), or preauricular hairs directed posteri- orly, partly covering ears (lateral facial crest in- frazygomatic, 3 specimens); subauricular hairs pale brownish gray, not elongated; outer surface of thighs and shanks pale brownish gray; T 115-11 6% of HB in 3 adult specimens examined. For external and cranial measurements, see Appendixes 7-9 and 12 and Fooden and Albrecht (1993, p. 538). Specimens Examined— Total 8: skins and skulls, 8 (see Appendix 1). 80 FIELDIANA: ZOOLOGY Macaca fascicularis fusca Miller, 1903a, p. 476 Macacus fuscus Miller, 1903a, p. 476 — for details con- cerning type series, see below. Lyon & Osgood, 1909, p. 283— holotype cataloged. Macaca fusca: Thomas, 1923, p. 591— specimen col- lected in P. Simeulue. [Cynomolgus] fuscus: Trouessart, 1904, p. 16— geo- graphic distribution. Pit hecus fuscus: Elliot, 1913, p. 228— external and cra- nial characters. Pithecus fuscus fuscus: Lyon, 1916, p. 457— geographic distribution. Macaca irus fusca: Chasen, 1940a, p. 69— geographic distribution. Poole & Schantz, 1 942, p. 242— holotype cataloged. Kellogg, 1944, p. 76— taxonomic compar- isons; zoogeography. M[acaca] f[ascicularis] fusca: J. R. Napier & Napier, 1967, pp. 349, 403— geographic distribution. Type Series— Holotype (by original designa- tion), usnm 1 14164 (Coll. No. 1348), adult male, skin and skull, collected at Telok Dalam (= Lhok Dalam), east coast of Simalur I. (= P. Simeulue), Indonesia, by W. L. Abbott, 20 Nov. 1901 (for collecting locality details, see collector's field cat- alog, usnm); paratypes, usnm 114162 (Coll. No. 1346, adult female, P. Simeulue, 18 Nov. 1901), usnm 1 14163 (Coll. No. 1347, adult male, P. Si- meulue, 18 Nov. 1901), usnm 114165 (Coll. No. 1349, adult female, P. Simeulue, 20 Nov. 1901), usnm 114166 (Coll. No. 1350, adult female, P. Simeulue, 20 Nov. 1901), usnm 114167 (Coll. No. 1354, adult male, P. Simeulue, 26 Nov. 1901), usnm 114168 (Coll. No. 1387, adult male, Siba- boh, P. Simeulue, 17 Dec. 1901), usnm 114169 (Coll. No. 1396, adult male, Labuan Badjan Bay [= Labuhanbajau], P. Simeulue, 1 Jan. 1902), usnm 114247 (Coll. No. 1397, late juvenile male, P. Lasia, 4 Jan. 1902), and usnm 1 14248 (Coll. No. 1398, adult male, P. Lasia, 5 Jan. 1902), skins and skulls, collected in Indonesia by W. L. Abbott. Type Locality— Telok Dalam (= Lhok Dal- am), east coast of Simalur Island (= P. Simeulue), west of northern Sumatra, Indonesia. Distribution (Fig. 25)— P. Simeulue, west of northern Sumatra, Indonesia. Diagnosis— General color of dorsal surface of trunk blackish, pale hair annulations pale yellow- ish; crown blackish, hairs conspicuously annulated with pale yellowish; preauricular hairs usually di- rected posteriorly, partly covering ears (lateral fa- cial crest infrazygomatic, 9 of 13 specimens ex- amined), occasionally directed anteriorly, forming part of lateral facial crest (crest transzygomatic, 4 specimens); subauricular hairs pale gray, not elon- gated; outer surface of thighs and shanks blackish; T 90-108% of HB in 10 adult specimens exam- ined. For external and cranial measurements, see Table 37, Figures 26 and 27, and Appendix 12. Remarks— For comments on T and crown hair annulation in M. fascicularis fusca, see below (M. f. lasiae— Remarks). Behavior in M. fascicularis fusca has been compared with that in Sumatran M. f. fascicularis by van Schaik and van Noord- wijk (1985a, p. 141) and Sugardjito et al. (1989, p. 197). These authors suggest that sexual dimor- phism of size may be less in M. fascicularis fusca than in Sumatran M. f. fascicularis; however, re- duced sexual dimorphism is not apparent in avail- able cranial measurements of M. fascicularis fusca (Table 37, Fig. 26). Specimens Examined— Total, 19: skins and skulls, 12; skins only, 7 (see Appendix 1). Macaca fascicularis lasiae (Lyon, 1916, p. 453) Macacus fuscus: Miller, 1903a, p. 476 (part)— specimens collected in P. Lasia; taxonomic comparisons. Pithecus fuscus lasiae Lyon, 1916, p. 453— for details concerning type series, see below. W. C. O. Hill, 1 974, p. 5 1 1 —cited as a synonym of Macaca irus fusca Mil- ler, 1903a. P. H. Napier, 1981, p. 13-cited as a syn- onym of Macaca fascicularis fusca Miller, 1903a. Macaca irus lasiae: Chasen, 1940a, p. 69— geographic distribution. Poole & Schantz, 1942, p. 242— holotype cataloged. Kellogg, 1944, p. 76— taxonomic compar- isons; zoogeography. Type Series— Holotype (by original designa- tion), usnm 1 14248 (Coll. No. 1398), adult male, skin and skull, collected in P. Lasia, southeast of P. Simeulue, Indonesia, by W. L. Abbott, 5 Jan. 1902; paratype, usnm 114247 (Coll. No. 1397), late juvenile male, skin and skull, collected in P. Lasia, Indonesia, by W. L. Abbott, 4 Jan. 1902. These two specimens also are paratypes of M. fas- cicularis fusca Miller, 1903a, p. 476 (see above; cf. International Code of Zoological Nomencla- ture, 1985, Article 72d). Type Locality— Palau Lasia, southeast of P. Simeulue, west of northern Sumatra, Indonesia. Distribution (Fig. 25)— Pulau Lasia, west of northern Sumatra, Indonesia. Diagnosis— General color of dorsal surface of trunk blackish, pale hair annulations pale yellow- ish; crown blackish, hair annulations inconspic- uous; preauricular hairs directed posteriorly, part- ly covering ears (lateral facial crest infrazygomatic, FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 81 Table 3 7 . Sexual dimorphism of greatest length of skull (mm) compared in Macacafascicularisfusca and Sumatran Macaca fascicularis fascicularis (cf. Fig. 26). Adult females Adult males Latitude Mean SD N Mean SD N Ratio 1 M. fascicularis J usca 2°00'-2°59'N 102.22 2.81 4 121.77 1.88 7 1.19 Sumatran M. f. fascicularis 5°00'-5°59'S — — 120.15 4.58 4 — 4°00'-4°59'S 103.10 — 1 120.65 4.96 6 1.17 3°00'-3°59'S 98.35 2.90 2 117.64 3.61 7 1.20 2°00'-2°59'S 103.40 — 1 — — — 1°00'-1°59'S 94.90 — 1 114.90 — 1 1.21 0°00'-O°59'S 100.25 4.04 4 111.16 1.24 7 1.11 0°00'-0°59'N — — 119.00 — 1 — 1°00'-1°59'N 95.03 1.82 3 116.27 3.99 3 1.22 2°00'-2°59'N 95.30 0.71 2 115.40 — 1 1.21 3°00'-3°59'N 97.64 2.16 10 115.86 3.42 21 1.19 4°00'-4°59'N 103.40 — 1 120.63 4.92 3 1.17 Total 98.18 3.31 25 116.65 4.39 54 1.19 Male mean divided by female mean. 1 of 2 specimens examined), or directed anteriorly, forming part of lateral facial crest (crest transzy- gomatic, 1 specimen); subauricular hairs pale gray- ish brown, not elongated; outer surface of thighs blackish, becoming pale grayish brown on outer surface of shanks; T 1 1 8% of HB in 1 adult spec- imen examined. For external and cranial mea- surements, see Figure 27 and Appendix 12. Remarks— Although M.f. lasiae, endemic to P. Lasia, is known from only 2 specimens (1 late juvenile male, 1 adult male), T in these 2 speci- mens is clearly longer— both relatively and abso- lutely—than in 13 specimens of M. fascicularis fusca (Fig. 27), which inhabits nearby P. Simeulue and which M. f. lasiae otherwise strongly resem- bles; the difference in T was noted previously by Miller (1903a, p. 477) and Lyon (1916, p. 453). Judging from the general pattern of T variation in M. fascicularis, the longer tail in M. f. lasiae prob- ably is primitive, and the shorter tail in M. fas- cicularis fusca probably is derived (Figs. 16, 18; Appendix 12). Crown hair annulations are less conspicuous in the 2 specimens examined of M. f lasiae than in 1 3 specimens examined of M. fascicularis fusca. The blackish dorsal pelage of M. f lasiae in deep-water P. Lasia (and M. fascicularis fusca in deep-water P. Simeulue) contrasts strikingly with the yellowish brown to golden brown dorsal pelage of M. f fascicularis in shallow- water P. Tuangku (usnm 1 14408-1 14410), an island that is only 50 km east of P. Lasia but that is on the Sumatran shelf (Figs. 3, 4). The blackish dorsal pelage of M. f lasiae and M. fascicularis fusca also contrasts strikingly with the brownish dorsal pelage of M. f. fascicularis in deep-water P. Nias, which is about 95 km southeast of P. Lasia (Appendix 3). Specimens Examined— Total, 2: skins and skulls, 2 (see Appendix 1). Macaca fascicularis atriceps Kloss, 1919c, p. 347 Macaca irus atriceps Kloss, 1919c, p. 347— for details concerning type series, see below. Kloss, 1921, p. 76 — external and cranial characters; taxonomic compari- sons. Kloss, 1926, p. 358— external and cranial char- acters; taxonomic comparisons. Poole & Schantz, 1 942, p. 243— holotype cataloged. Weitzel et al., 1988, p. 105— cited as a synonym of Macaca fascicularis bin- tangensis (Elliot, 1 909) (= M. f fascicularis). M[acaca] f[ascicularis] atriceps: J. R. Napier & Napier, 1967, pp. 349, 403— geographic distribution. Type Series— Holotype (by original designa- tion), usnm 236622 (Coll. No. 2283), adult male, skin and skull, collected in Koh Khram I. (= Ko Khram Yai), near Cape Liant (= Laem Samae San), southeastern Thailand, by C. B. Kloss, 30 Oct. 1916; paratypes, bm(nh) 1939.891 (Coll. No. 2284, adult male), bm(nh) 1939.892 (Coll. No. 2287, adult female), usnm 236618 (Coll. No. 2282, 82 FIELDIANA: ZOOLOGY ■ 125- ■ ■ ■ ■ ■ A 1 ■ ■ A 120- 3 M VI 1 ■ ■ ■ ■ ■ ■ A ■ f I ■ s -• — ■ ■ I ■ length of © ■ ■ ■ * ■ Greatest 8 D a H A D 100 a A % 95- D 00 1 a a 90 1 600 6°S 4 2 2 4°N Latitude o ■ Sumatra: females, males A A Simeulue: females, males Fig. 26. Variation of greatest length of skull in adult Macacafascicularisfusca (Indonesia: P. Simeulue) com- pared with variation in adult Sumatran M.f.fascicularis (cf. Table 37). adult male), usnm 236619 (Coll. No. 2286, adult female), usnm 236620 (Coll. No. 2289, subadult female), usnm 236621 (Coll. No. 2290, juvenile male), zrc 4-012 (Coll. No. 2288, adult female), zrc 4-013, skull, and zrc 4-733, skin (Coll. No. 2285, adult male), skins and skulls, collected in Ko Khram Yai, Thailand, by C. B. Kloss, 30 Oct. 1916. Type Locality— Ko Khram Yai, near Laem Samae San, southeastern Thailand. Distribution (Fig. 25)— Ko Khram Yai, Thai- land. Diagnosis— General color of dorsal surface of trunk buffy to medium brown, pale hair annula- tions pale yellowish; crown with a narrow, sharply defined dark brown to blackish patch (Fig. 7) that extends laterally as far as middle of each eye and posteriorly as far as vertex or (rarely) occiput; preauricular hairs directed anteriorly, forming part of lateral facial crest (crest transzygomatic); T 1 09- 500 ^450 ♦ juv.* A A » ' A subad. ?age A ^ * A A A A 1 i 1 ■ ■ 400 350 300 350 400 450 500 550 Head and body length (mm) ♦ Lasia: males A A Simeulue: females, males Fig. 27. Tail length vs. head and body length in Ma- cacafascicularisfusca (Indonesia: P. Simeuleu) and M. f lasiae (Indonesia: P. Lasia); specimens are adult, except as otherwise indicated (three specimens). 1 28% of HB in 9 adult specimens examined. For external and cranial measurements, see Table 38, Figure 28, and Appendix 12. Remarks- Kloss (1919c, p. 348; 1921, p. 76; 1926, p. 358) indicated that M. f. atriceps differs cranially from M. f. condorensis and Indochinese M . fascicularis in size of the rostrum, supraorbital ridges, orbits, zygomatic arches, mandible, and molars and in shape of the tooth rows and palate. As evidence, Kloss cited measurements of some of these structures in 9 specimens of M.f. atriceps (4 adult females, including 1 young adult regarded by Kloss as a subadult, and 5 adult males) and in 3 specimens of M. f. condorensis ( 1 adult female and 2 adult males); no measurements are cited for specimens of Indochinese M. fascicularis. New data, derived from 5 additional adult spec- imens of M.f. condorensis (1 female, 4 males) and 32 adult specimens of Indochinese M. fascicularis (22 females, 10 males), permit reevaluation of the six size characters specified by Kloss (Table 38, Fig. 28). For each of these six cranial charac- ters, size variation in M. f. atriceps overlaps that in M. f. condorensis and Indochinese M. fascicu- laris. Relative sizes of the rostrum, supraorbital FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 83 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). Adult females Adult males Population Mean SD Extremes N Mean SD Extremes N Rostral -postrostral ratio (R/PR x 100) I-C M. fascicularis 49.5 4.6 41.1-56.4 20 60.8 3.1 55.5-65.1 7 M. f. atriceps 51.0 2.8 48.1-54.5 4 58.4 3.6 54.0-63.9 5 M. f condorensis 50.4 _ 46.6-54.1 2 Relative biorbital breadth (BB/GL 1 59.8 x 100) 3.4 55.3-63.3 5 I-C M. fascicularis 51.9 2.7 45.3-55.9 18 50.0 2.1 47.2-54.0 10 M. f atriceps 51.0 2.1 48.5-53.1 4 50.3 2.2 48.4-53.9 5 M. f condorensis 48.9 _ 48.0-49.7 2 Relative orbital height (OH/GL x 51.2 100) 4.4 46.5-56.8 6 I-C M. fascicularis 20.6 1.7 17.5-24.6 19 18.4 1.1 16.9-20.9 10 M. f atriceps 19.4 1.1 18.3-20.7 4 17.2 1.3 15.8-19.2 5 M. f condorensis 21.5 - 20.4-22.5 2 Relative zygomatic breadth (ZB/GL 19.1 x 100) 1.2 17.8-20.6 6 I-C M. fascicularis 67.8 2.4 63.2-71.3 22 68.9 1.7 66.4-71.2 10 M. f atriceps 66.8 2.4 63.1-68.4 4 68.4 2.3 66.8-71.8 4 M. f condorensis 65.7 - 64.9-66.6 2 Relative mandibular length (ML/GL 67.8 x 100) 2.6 65.1-71.3 6 I-C M. fascicularis 73.0 2.1 70.5-77.6 20 75.8 3.1 71.5-81.3 9 M. f. atriceps 72.4 0.5 71.8-72.9 4 75.6 1.6 73.4_77.9 5 M. f condorensis 73.2 - 72.4-74.0 2 74.9 1.6 72.6-76.4 6 Relative length of maxillary molar row (Ml M3/GL x 100) I-C M. fascicularis 19.9 1.1 18.2-22.6 22 17.4 0.8 16.2-18.6 10 M. f atriceps 21.6 0.7 20.9-22.2 4 19.8 1.1 18.9-21.3 5 M. f condorensis 20.3 - 19.8-20.8 2 18.3 1.4 15.7-19.8 6 1 Greatest length of skull, excluding incisors. ridges, zygomatic arches, and mandible are essen- tially similar in these three populations; relative size of the orbits tends to average smaller in M. f. atriceps than in the other two populations, as sug- gested by Kloss, and relative size of the molars tends to average slightly greater in M. f. atriceps than in the other two populations, as also sug- gested by Kloss. Specimens Examined— Total 11: skins and skulls, 11 (see Appendix 1). Macaca fascicularis condorensis Kloss, 1926, p. 357 monkies: King, 1784, p. 462 — said to be abundant in Jan. 1780. Macaca irus Cuv., subsp.: Kloss, 1 92 1 , p. 75 —specimens collected in Pulo Condore (= Con Son); taxonomic comparisons. Macaca irus condorensis Kloss, 1 926, p. 357 — for details concerning type series, see below. Gibson-Hill, 1949, p. 172— type history. J. E. Hill, 1960, p. 31— typ history. Macaca fascicularis condorensis: Van Peenen et al., 1970 p. 420— specimens collected in Con Son and Hon Ba external and cranial characters. P. H. Napier, 1981 pp. 13, 17— external characters; holotype cataloged. Type Series— Holotype (by original designa tion), bm(nh) 1947.1498 (CBK [C. B. Kloss] No 269 1), adult male, skin and skull, collected in Pule Condore (= Con Son), offCochin-China (= south ern Vietnam), by "Dr. Malcolm Smith's collec tor," 20 Sep. 1919; paratypes (implicitly cited ir original description; explicitly cited by Kloss, 1 92 1 p. 75), bm(nh) 1939.893 (CBK No. 2692, adul male, 23 Sep.) and bm(nh) 1939.894 (CBK No 2693, adult female, 19 Sep.), skins and skulls, col lected in Con Son, Vietnam, by "Dr. Malcolrr Smith's collector," 1919. Type Locality— Con Son, Vietnam. Distribution (Fig. 25)— Con Son and nearby Hon Ba, two islands in the South China Sea, south east of southern Vietnam. 84 FIELDIANA: ZOOLOGY I ■ I - v- • D n_ fD^ ■ nO s D A^pB ru 0° 90 100 110 120 130 Greatest length of skull (mm) 60 a ~ 50 a fl 45- | 40 (n ^ 35 30 65 - A ■ A ♦ Q« ■ ■ ■ ■ D * ♦ A S3 D □ □ A 0. A D D ♦ D D o £ a ♦ a a n f D A D □ d a ■ ■ ■ ■ ■ □1 D A 90 Afl r? A Euo D D°A n D a 70 75 80 85 90 Postrostral length (mm) 90- a a 5 so- ts ed 0) ■O I ■ ■ A ■ ■ ■ A a □ J D a n a a n cfl '0 £ o bO N 60- a a 0d a n * Ai , ■ O O D ♦ D 18- ■ , 1 , 1 1 1 1 — 1 100 110 120 130 Greatest length of skull (mm) Females Males O ♦ M. L atri.ceps A A M. f. can dor crisis □ ■ Indochinese M. fascicularis Fig. 28. Craniometric comparisons of adult Macaca fascicularis atriceps (Thailand: Ko Khram Yai), M. f. con- dorensis (Vietnam: Con Son), and mainland Indochinese M. fascicularis (cf. Table 38). FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 85 Diagnosis— General color of dorsal surface of trunk buffy to medium brown (with blackish mid- dorsal streak in 2 juveniles, zrc 4-017, 4-018), pale hair annulations pale yellowish to golden; crown with a broad dark brown to blackish patch (Fig. 7) that extends laterally as far as lateral mar- 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- matic); T 109-127% of HB in 6 adult specimens examined. For external and cranial measure- ments, see Table 38, Figure 28, and Appendixes 7-9 and 12 and Fooden and Albrecht (1993, p. 537). Remarks — See above (M. / atriceps — Re- marks). Specimens Examined— Total, 16: skins and skulls, 1 1 ; skins only, 2; skulls only, 3 (see Ap- pendix 1). Macaca fascicularis tua Kellogg, 1944, p. 75 Macaca irus tua Kellogg, 1944, p. 75 — for details con- cerning type series, see below. W. C. O. Hill, 1974, p. 507— cited as a synonym of Macaca irus irus F. Cu- vier, 1818 (I. Geoffroy, 1826). P. H. Napier, 1981, p. 13— cited as a synonym of Macaca fascicularis fasci- cularis (Raffles, [1821]). M[acaca]f[ascicularis] tua: J. R. Napier & Napier, 1967, pp. 349, 404— geographic distribution. Type Series— Holotype (by original designa- tion), usnm 197663 (Coll. No. 626), adult male, skin and skull, collected in Pulo Muara Tua (= P. Maratua), east of Borneo, Kalimantan, Indonesia, by H. C. Raven, 21 May 1913; paratypes, usnm 197660 (Coll. No. 623, juvenile female), usnm 197661 (Coll. No. 624, adult female), and usnm 197662 (Coll. No. 625, subadult male), skins and skulls, collected in P. Maratua, Kalimantan, In- donesia, by H. C. Raven, 21 May 1913. Type Locality— Pulau Maratua, Kalimantan, Indonesia. Distribution (Fig. 25)— Pulau Maratua, east of northeastern Kalimantan, Indonesia. Diagnosis— General color of dorsal surface of trunk blackish, pale hair annulations pale yellow- ish; crown pale yellowish brown, hairs conspicu- ously annulated with pale yellowish; preauricular hairs directed anteriorly, forming part of lateral facial crest (crest transzygomatic); subauricular hairs pale ochraceous-buff, conspicuously elon- gated; outer surface of thighs brownish gray, be- coming pale brownish gray on shanks; T 1 26-1 3 1 % of HB in 2 adult specimens examined. For external and cranial measurements, see Appendix 12. Specimens Examined— Total, 4: skins and skulls, 4 (see Appendix 1). Macaca fascicularis karimondjawae Sody, 1949, p. 132 Macacus cynomolgus: Willink, 1905, p. 175 (part, not Linnaeus, 1766, p. 38)— "P. Karimon-Djawa (P. Ka- moedian)" included in geographic distribution. Macaca irus karimondjawae Sody, 1949, p. 132, Table 1 —for details concerning type series, see below. W. C. O. Hill, 1974, p. 528— external and cranial characters. Macaca fascicularis karimondjawae: P. H. Napier, 1981, p. 13— type locality information. Type Series— Holotype (by original designa- tion), rmnh 10608 (mzb 2719), adult male, skin and skull, collected in Karimon Djawa (= P. Kar- imunjawa), Java Sea, Indonesia, by W. Rom- swinckel, 28 Nov. 1930 (incorrectly given as "28.VI.1930" in original description); paratypes (cf. Sody, 1949, p. 132, Table 1), mzb 1454 (adult male, cranium only, 7 May), mzb 1455 (adult fe- male, skin and skull, 9 May), mzb 1456 (juvenile female, skin and skull, 10 May), mzb 1457 (adult female, skin and skull, 11 May), and mzb 1458 (infant male, skin and skull, 14 May), collected in P. Karimunjawa, Indonesia, by K. W. Dammer- man, P. F. Franck, and/or Denin, 1926; mzb 2717 (juvenile male) and mzb 2718 (late juvenile fe- male), skins and skulls, collected in P. Karimun- jawa, Indonesia, by W. Romswinckel, 26 Nov. 1930; and mzb 2720 (adult female, cranium only) collected in P. Kemujan, < 1 km northeast of P. Karimunjawa, Indonesia, by W. Romswinckel, 25 Nov. 1930. Type Locality— Karimon Djawa (= P. Kari- munjawa), Java Sea, Indonesia. Distribution (Fig. 25)— Pulau Karimunjawa and, presumably, nearby P. Kemujan (known from one cranium only), Java Sea, 60 km north of cen- tral Java, Indonesia. Diagnosis— General color of dorsal surface of trunk dark grayish brown, pale hair annulations pale yellowish; crown frequently with a blackish wash; preauricular hair direction not studied; T 101-1 12% of HB in 3 adult specimens examined. For external and cranial measurements, see Ap- pendix 12 and Fooden and Albrecht (1993, p. 538). 86 FIELDIANA: ZOOLOGY Specimens Examined— Total 9: skins and skulls, 6; skin only, 1; skulls only, 2 (see Appendix 1). Macaca fascicularis Subspecies Undetermined Macacus cristatus Gray, 1870, p. 30— holotype, bm(nh) 1858.4.28.9, late juvenile male, skin (albinistic) and skull, acquired in 1858 from collection of Th. G. van Lidth de Jeude, Utrecht, provenance unknown. Schle- gel, 1876, p. 101— cited as a synonym of Cercocebus cynamolgos: Schlegel (= Macaca fascicularis (Raffles, [1821])). Elliot, 1913, p. 249-cited as a synonym of Pithecus philippinensis (I. Geoffroy, [1843]). W. C. O. Hill, 1974, pp. 477, 522— cited as a synonym of Ma- caca irus philippinensis I. Geoffroy, [1843]. P. H. Na- pier, 1981, pp. 13, 20— cited as a synonym of Macaca fascicularis philippinenesis I. Geoffroy, [1843]. Food- en, 1 99 1 , p. 24— subspecies not determined; taxonom- ic history. Macacus albus: P. H. Napier, 1981, p. 20— unavailable manuscript name, author unknown, written on tag of holotype of Macacus cristatus Gray, 1870. Evolution and Dispersal Available information concerning the distri- bution, variation, natural history, and paleontol- ogy of M. fascicularis provides a basis for hypo- thetical interpretation of the evolution and dis- persal of this species. Because most of the geo- graphic range of M. fascicularis is insular, a large part of this interpretation concerns water gaps and their effect on dispersal. Four major factors are pertinent to dispersal in the insular part of the range of M. fascicularis. 1. Eustatic changes in sea level. Since the be- ginning of the Pleistocene, ca. 1.75 Ma, worldwide sea level has been strongly controlled by, and in- versely correlated with, the cyclic growth of con- tinental glaciers (Van Couvering & Kukla, 1988, p. 459; Heaney, 1991a, p. 56; Cande & Kent, 1 992, pp. 13,936, 13,938). Sea level has been approxi- mately as at present since ca. 5 Ka (Clark et al., 1978, p. 283; Fairbanks, 1989, p. 639). It rose sigmoidally to this level from about 1 20 m lower during the last glacial maximum, ca. 1 8 Ka, when the Sunda Shelf was exposed and the Indochinese Peninsula, the Malay Peninsula, Sumatra, Borneo, Java, and shallow-water fringing islands were unit- ed to form a single large landmass (Fig. 3). During the preceding interglacial, ca. 120 Ka, sea level was high, and components of this large landmass were isolated, approximately as at present. During the penultimate glacial maximum, ca. 1 60 Ka, sea 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- cisely known magnitude, have occurred with a pe- riodicity of ca. 100 Ka since the beginning of the Pleistocene. Tectonic forces also have affected the distribu- tion of land and sea within the range of M. fas- cicularis. Prominent examples of such effects are the approximately 500-m uplift of the north coast of P. Sumba (Pirazzoli et al., 1991, p. 1835), the approximately 500-m subsidence and subsequent reelevation of P. Maratua (Kuenen, 1947, p. 8), and the massive volcanic explosions of Rakata (= Krakatau) and Toba (Dammerman, 1948, p. 4; Chesner et al., 1 99 1 , p. 200; Rampino & Self, 1993, p. 269). However, the zoogeographic implications of Pleistocene tectonic changes in this area are 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, Natural History). 3. Natural rafting. Monkeys may be transport- ed across water gaps on floating trees or larger entwined masses of vegetation (cf. Wallace, 1895, p. 74). The tendency for M. fascicularis to exploit coastal habitats (see above, Natural History) makes this species a prime candidate for such passive dispersal on natural rafts. A specific instance is reported by Hickson (1889, p. 190): "Some days after the eruption of Krakatoa in 1883 a female green monkey [M. fascicularis] was found floating on some drifting timber in the Sunda Straits. She was terribly scorched, but completely recovered, and is, I believe, still alive." The average surface current speed in the Sunda Shelf area is about 1 km/hr (E. G. W. Smith, 1974, Fig. 2.3-1; Couper, 1989, p. 50). Arbitrarily as- suming that some members of a small group of M. fascicularis could survive on a natural raft for up to about 3 days, such a group potentially could colonize a new island about 75 km distant from its original habitat; during monsoons and other storms, the current would be stronger and the dis- tance covered would be greater. The reproductive capability of a small group of M. fascicularis in- dividuals to multiply and populate islands not pre- viously inhabited by this species is demonstrated by the success of introduced populations in Mau- ritius and Angaur (Poirer & Smith, 1 974, p. 264; Sussman & Tattersall, 1986, p. 30). FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 87 Table 39. Hypothetical stages in evolution and dis- persal of Macaca fascicularis. Stage I, > 1 Ma Dispersal of M. fascicularis into Sunda Shelf area. Stage II, ca. 160 Ka Dispersal and isolation of progenitors of strongly dif- ferentiated deep-water fringing-island popula- tions. M. f. umbrosa: Nicobar Islands M. f. fusca, M. f. lasiae: Simeulue and Lasia M. f. tua: Maratua M. f. philippinensis: western, northern and eastern Philippines Stage III, > 18 Ka Differentiation of progenitors of populations in the Indochinese Peninsula and northern part of the Isthmus of Kra. M. f. aurea: Indochinese Peninsula, Isthmus of Kra, Mergui Archipelago M. f. fascicularis: Indochinese Peninsula, Isthmus of Kra Stage IV, ca. 18 Ka Dispersal and isolation of progenitors of weakly dif- ferentiated deep-water fringing-island popula- tions. M. f. fascicularis: Nias M. f fascicularis: southern Philippines Stage V, < 18 Ka Isolation of progenitors of shallow-water fringing-is- land populations and populations in Penida and Lombok (deep-water). M. f karimondjawae: Karimunjawa, Kemujan M. f. atriceps: Khram Yai M. f. condorensis: Con Son, Hon Ba M. f fascicularis: other shallow-water fringing is- lands, excluding Bali M. f fascicularis: Bali, Penida, Lombok Stage VI, ca. 4.5 Ka Dispersal and isolation of progenitors of populations in eastern Lesser Sunda Islands (deep-water). M. f fascicularis: Sumbawa-Timor 4. Transport by humans. Interisland transport of captive M. fascicularis by humans is reported by Wallace (1869, p. 294) and presumably has a long, if undocumented, history in Southeast Asia. Homo erectus reached Java by 1-2 Ma, and H. sapiens reached Borneo and Australia by ca. 40 Ka (Bellwood, 1985, pp. 29, 89, 98; cf. Swisher et 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 sea level, the dispersal of H. sapiens to Australia must have been over water, which implies that some means of over- water transport was available to humans in this area at least 40 Ka (cf. Sondaar etal., 1994, p. 1261). Available evidence indicates a minimum of six major stages in the evolution and dispersal of M. fascicularis (Table 39). These are discussed below. Stage I, > 1 Ma The earliest record of M '. fascicularis is at Trinil, east-central Java, ca. 1 Ma (Table 33); the Trinil fossils are part of the Dubois collection that also includes Homo erectus and two species of leaf monkeys, Presbytis comata and Trachypithecus auratus (Hooijer, 1962b, p. 5; Theunissen et al., 1990, p. 41; Groves, 1993, p. 273). Assuming that M. fascicularis, H. erectus, and leaf monkeys dis- persed to Java from mainland Southeast Asia, they probably reached Java by dry land during one of the Early Pleistocene periods of glacial advance and low sea level (cf. J. de Vosetal., 1994, p. 131). The earliest record of M. nemestrina also is in east-central Java but is somewhat later (ca. 800 Ka) than that of M. fascicularis (Table 33). Despite this later date, zoogeographic evidence indicates that M. nemestrina probably preceded M. fasci- cularis in the Sunda Shelf area (Fooden, 1975, p. 70). Macaca fascicularis is absent in Sulawesi and Kepulauan Mentawai, faunistically distinctive is- lands off the Sunda Shelf that are inhabited by M. nemestrina or its close relatives. Since M. fasci- cularis is otherwise much more widely distributed thanM nemestrina and its relatives (Fooden, 1980, p. 4), it appears that the intrinsic ability of M. fascicularis to disperse is not inferior to that of M. nemestrina. This suggests that the M. nemestrina stock dispersed from the Sunda Shelf to Sulawesi and Kepulauan Mentawai at some time prior to 1 Ma, before M. fascicularis had arrived on the Sunda Shelf. During Late Pleistocene or Holocene, M. fas- cicularis may have been less abundant than M. nemestrina in the Malay Peninsula and Sumatra (see above, Fossils and Subfossils). If so, this would require explanation, because M . fascicularis is now generally more abundant than M. nemestrina in the Malay Peninsula, Sumatra, and elsewhere (Fooden, 1975, p. 61). Stage II, ca. 160 Ka The most strongly differentiated populations of M. fascicularis inhabit four deep-water islands or island groups: (1) Nicobar Islands, northwest of Sumatra; (2) P. Simeulue and P. Lasia, west of FIELDIANA: ZOOLOGY Sumatra; (3) P. Maratua, east of Borneo; and (4) Philippines, excluding southern islands (Fig. 25). All of these populations are very dark (Appendix 3), and their deep-water island habitats are sym- metrically arrayed adjacent to the northwestern and northeastern margins of the Sunda Shelf (cf. Kellogg, 1 944, p. 76). Assuming that dark dorsal pelage is derived in M. fascicularis, the similar pelage saturation and geographic relationships of these populations suggest that they may have had a similar evolutionary history. These populations appear to have been isolated longer than popu- lations of M. fascicularis inhabiting shallow- water fringing islands, which were isolated by rising sea level subsequent to the last glacial maximum, ca. 1 8 Ka. A working hypothesis is that the progen- itors of these strongly differentiated populations reached their deep-water habitats during the pen- ultimate glacial maximum, ca. 1 60 Ka, when sea level was about 1 60 m lower than at present, and that these populations have been isolated at least since the subsequent interglacial, ca. 1 20 Ka. Fu- ture new evidence concerning the paleontology, morphology, or molecular biology of these pop- ulations may provide a test of this hypothesis. Nicobar Islands— This group includes 19 is- lands, of which 1 1 are larger than about 2 km 2 . M. fascicularis inhabits only Great Nicobar I., Lit- tle Nicobar I., and Katchall I. (Kloss, 1903a, p. 1 14)— the southernmost 3 of the 1 1 larger islands. Forest cover in the Nicobars is restricted to these three islands and Tillanchong I., one of the more northern islands (Kloss, 1903a, p. 109). Monkeys, including M. fascicularis, are absent from the An- daman Islands, a forested group of deep-water is- lands that lies between the Nicobar Islands and Burma (Kloss, [1928], p. 802; Chaturvedi, 1980, p. 134). Great Nicobar is separated from Sumatra by an ocean trench that is 1 50 km wide and more than 200 m deep; for half of its width, this trench is more than 1,000 m deep. The channel between Great Nicobar and Little Nicobar is 6 km wide and less than 100 m deep, which indicates that these two islands were united during the last glacial maximum. Narrow channels (< 10 km wide) be- tween Little Nicobar and Katchall are more than 1 20 m deep, which probably indicates that these two islands were not united during the last glacial maximum. The nonvolant mammal fauna of the Nicobars is limited to one species each of shrew, tree shrew, and pig, four to six species of rats, M fascicularis, and humans (Miller, 1902b, p. 792; Kloss, [1928], p. 802; Chaturvedi, 1980, p. 133; Musser & Carle- ton, 1993, pp. 651, 660); no information is avail- able concerning the date of first arrival of humans in the Nicobars (Fuchs, 1973, p. 285; Gratton, 1 992, p. 209). The poverty of the fauna, together with the deep ocean trenches that surround this group of islands, strongly implies that these islands have never had a land connection to Sumatra or any other source of the rich mammalian fauna of Southeast Asia. This in turn implies that M. fas- cicularis reached the Nicobars by over-water dis- persal; the absence of this species in the Andaman Islands indicates that it dispersed to the Nicobars from Sumatra, the Malay Peninsula, or the Isth- mus of Kra— not from Burma. As noted above, the degree of differentiation of M. fascicularis in the Nicobars suggests that this species may have reached these islands during the penultimate gla- cial maximum, ca. 1 60 Ka; at this early date, dis- persal by natural rafting would be more likely than dispersal by human introduction. Following the arrival and isolation of M. fas- cicularis in the Nicobars, dorsal pelage in the im- migrant population apparently darkened, and skull size increased, resulting in differentiation of the subspecies M. f umbrosa. The present morpho- logical uniformity of populations of this subspe- cies in the three islands that it inhabits implies relatively recent genetic exchange among these populations; this exchange presumably occurred during the last glacial maximum, ca. 1 8 Ka, when Great Nicobar was joined to Little Nicobar and the gap between Little Nicobar and Katchall was reduced to narrow channels (< 10 km wide). The lateral facial crest pattern is infrazygomatic in three of eight M. f umbrosa specimens exam- ined (one of three Great Nicobar specimens, two of three Little Nicobar specimens, zero of two Katchall specimens) (see Appendix 6). Three al- ternative interpretations would account for the rel- atively high frequency of the infrazygomatic pat- tern in M. f. umbrosa: (1) it may indicate that the progenitors of M. f umbrosa dispersed to the Ni- cobars from the Malay Peninsula or Isthmus of Kra, in or near the geographic range of M.f aurea (Figs. 9, 25); (2) it may indicate that the range of M. f aurea extended to Sumatra during the pen- ultimate glacial maximum, when progenitors of M. f umbrosa are hypothesized to have dispersed to the Nicobars; or (3) it may be the result of independent local mutation. P. Simeulue and P. Lasia— These two islands are approximately 1 20 km west of northern Su- 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 of the much smaller P. Lasia (ca. 6x3 km). Al- though M. fascicularis inhabits both of these is- lands, it is absent in P. Babi (ca. 8x7 km), which is only 3 km south of P. Lasia (Abbott in Miller, 1903a, p. 479). Deep trenches (> 200 m) separate P. Simeulue and P. Lasia from Sumatra and from each other. The deep trench between P. Simeulue and Sumatra is about 1 5 km wide, and that between P. Simeulue and P. Lasia is about 4 km wide. The narrow channel between P. Lasia and P. Babi probably is less than 1 00 m deep. The nonvolant mammal fauna of the Simeulue group, as reported by Kloss ([1928], p. 802; cf. Sugardjito et al., 1989, p. 197; Musser & Carleton, 1993, pp. 652, 659), is limited to one species each of civet and pig, two species of rats, M. fascicularis, and humans. This fauna, like that of the Nicobars (see above), presumably dispersed over water. Judging from the degree of differentiation of M. fascicularis in P. Simeulue and P. Lasia, this spe- cies may have dispersed to these islands— presum- ably by natural rafting— during the penultimate glacial maximum. Following isolation, the pelage of the immigrant population darkened. Genetic exchange between the populations in P. Simeulue and P. Lasia probably occurred during the last glacial maximum. During subsequent postglacial reisolation, the tail apparently shortened in the P. Simeulue population (Fig. 27; cf. above, M.f las- iae— Remarks), resulting in differentiation of M. fascicularis fusca and M.f. lasiae. Because P. Lasia probably was joined to P. Babi during the last glacial maximum, a dark population of M. fasci- cularis probably also inhabited P. Babi at that time but subsequently became locally extinct. The lateral facial crest pattern is infrazygomatic in 9 of 1 3 M. fascicularis fusca specimens exam- ined and in 1 of 2 M.f. lasiae specimens examined (Appendix 6). The geographic location of P. Si- meulue and P. Lasia strongly indicates that the progenitors of M. fascicularis fusca and M.f. lasiae dispersed to these islands from Sumatra, not from the Malay Peninsula or Isthmus of Kra (i.e., in or near the present range of M.f. aurea; Fig. 25). This implies either that the range of M. f aurea for- merly extended to Sumatra or that there has been independent mutation in these islands from the transzygomatic crest pattern to the infrazygomatic crest pattern. P. Maratua— This U-shaped island (area ca. 25 km 2 ) is a raised coral atoll 50 km east of north- central Borneo (Fig. 25; Kuenen, 1947, p. 5). It is separated from the Bornean shelf by a deep trench (> 180 m) that is 17 km wide; the water gap is narrowed by the presence of a stepping-stone is- land, P. Kakaban (area ca. 5 km 2 ), which inter- venes in the middle of the trench. The only mammal collector known to have vis- ited P. Maratua is H. C. Raven, who worked there in August 1912 and May 1913 (Deignan, [1960], p. 267). In P. Maratua, Raven collected M. fas- cicularis, two species of rats, possibly a few other murine rodents, and one species of bat (Kellogg, 1944, p. 75; Musser & Carina, 1982, p. 6; M. D. Carleton, usnm, letter 16 Dec. 1993). Judging from the degree of differentiation of M. fascicularis in P. Maratua, it may have dispersed to this island from Borneo during the penultimate glacial max- imum, ca. 1 60 Ka; the dispersal probably was over water— presumably by natural rafting. In isola- tion, dorsal pelage in the founder population dark- ened, resulting in differentiation of the subspecies M.f. tua. In all four M.f. tua specimens examined, the lateral facial crest pattern is transzygomatic, as in Bornean M. fascicularis. Philippines, Excluding Southern Islands— As previously discussed (Fooden, 1 99 1 , p. 24), the progenitors of current populations of M. fascicu- laris in western, northern, and eastern islands of the Philippine Archipelago may have dispersed to these islands during the penultimate glacial max- imum, ca. 1 60 Ka. Following isolation, dorsal pel- age in the founder population became darker and also tended to become erythristic, resulting in dif- ferentiation of M. f philippinensis. Stage III, > 18 Ka Indochinese Peninsula and Northern Part of Isthmus of Kra— Although zoogeographic ev- idence, cited below, indicates that M. f aurea ex- isted as a differentiated subspecies in the western part of the Indochinese Peninsula and northern part of the Isthmus of Kra prior to the last glacial maximum, ca. 18 Ka, the evolutionary history of this subspecies and that of neighboring popula- tions of M.f. fascicularis remain enigmatic. Equiv- ocal evidence concerning variation in M. fasci- cularis in the Indochinese Peninsula and Isthmus of Kra is presented in the following paragraphs. The relationship between variation in M. fasci- cularis and variation in M. mulatta—a. closely re- lated species that replaces M. fascicularis in the 90 FIELDIANA: ZOOLOGY northern part of the Indochinese Peninsula (Food- en, 1982, p. 576)— also is indicated. 1. Dorsal pelage color averages paler and less erythristic in the Indochinese Peninsula and northern part of the Isthmus of Kra than in the Malay Peninsula and Sumatra (Tables 1, 3). The pattern of pelage color resemblances is indepen- dent of local land connections; the Indochinese Peninsula and Isthmus of Kra are continuous with the Malay Peninsula, which is separated by a water gap from Sumatra. The latitude of pelage color transition is about 10°N, near the southern end of the Isthmus of Kra. The pale dorsal pelage color in M. fascicularis north of 10°N generally ap- proaches that in neighboring allopatric popula- tions of M. mulatta; however, the posterior part of the dorsal surface is strongly erythristic in M. mulatta— and not in Indochinese M. fascicularis. 2. In the Indochinese Peninsula north of about 1 3°N, which is at the northern end of the Isthmus of Kra, head and body length and skull length begin to decline northward in M. fascicularis (Fig. 10; Fooden & Albrecht, 1993, p. 532). This de- cline, which is anti-Bergmannian, is retrograde to the general relationship between size and latitude in core-area M. fascicularis and also is retrograde to the relationship between size and latitude in M. fascicularis in the Philippines, at approximately the same latitude as the Indochinese Peninsula. The northward decline of size in Indochinese M. fascicularis tends to bring the size of this species near to that of neighboring populations of M. mu- latta (Figs. 29, 30). 3. Tail length declines rapidly in the Indo- Chinese Peninsula north of about 1 3°N (Fig. 1 3). This is an acceleration of the general decline of tail length in core-area M. fascicularis north of the equator (Allen's rule); there is no comparable ac- celerated northward decline of tail length in Phil- ippine M. fascicularis. Parallel to the trend noted above for head and body length and skull length, the accelerated northward decline of tail length in Indochinese M. fascicularis tends to bring the tail length of this species near to that of neighboring populations of M. mulatta (Fig. 31; cf. Fooden, 1971, p. 29). 4. The subspecies M. f aurea inhabits a re- stricted area bordering the Bay of Bengal, mostly west of the mountain ranges that form the border between Burma and Thailand, along the western side of the Indochinese Peninsula and Isthmus of Kra, south to about 10°N (Figs. 9, 25). This sub- species is distinguished by its lateral facial crest pattern, which is infrazygomatic and which con- trasts with the transzygomatic pattern that pre- dominates in M. f fascicularis. Adjacent to the geographic range of M. f aurea are two areas— one in central and eastern parts of the Indochinese Peninsula and the other in eastern and southern parts of the Isthmus of Kra— inhabited by hetero- geneous populations that include some individuals with the infrazygomatic pattern, some with the transzygomatic pattern, and some with the asym- metric pattern; occasionally, heterogeneous pop- ulations have been sampled at a single locality. The southern part of the Indochinese Peninsula is inhabited by a now-disjunct population of M. f fascicularis (homogeneous for the transzygomatic pattern). The lateral facial crest pattern in shallow-water fringing-island populations of M. fascicularis matches that in adjacent mainland populations in Southeast Asia; the crest is infrazygomatic in the Mergui Archipelago— shallow- water islands west of the Isthmus of Kra, and the crest is transzy- gomatic in shallow- water islands east of the Isth- mus of Kra and adjacent to the Malay Peninsula (Fig. 9). This implies that M. f aurea had already differentiated, and was approximately restricted to its present geographic range, prior to the last glacial maximum (ca. 1 8 Ka), when M. fascicularis pre- sumably dispersed from mainland Southeast Asia to adjacent shallow-water islands. Pelage color, head and body length, tail length, and skull length in populations of M. f aurea are generally similar to those in populations of M. f fascicularis that inhabit similar latitudes in the Indochinese Peninsula and Isthmus of Kra. The lateral facial crest pattern in M. f aurea is similar to that in M. mulatta, where the infrazygomatic pattern is predominant (Stewart, 1933, p. 30). 5. The frequencies of mtDNA types, blood- protein alleles, and resistance to Plasmodium knowlesi malaria in populations of M. fascicularis north of the Isthmus of Kra differ markedly from those in populations south of the Isthmus of Kra (Tables 17, 19; Fooden, 1994, p. 585). The fre- quency off. knowlesi resistance in populations of M. fascicularis north of the Isthmus of Kra is sim- ilar to that in M. mulatta. Based on this evidence, the following conclu- sions may be drawn. Although suggestive, these conclusions are not adequate to explain the evo- lutionary history of M. fascicularis in the Indo- Chinese Peninsula and northern part of the Isth- mus of Kra. FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 91 650 -r- 600 550 500 4-50 Males L00 J £ 550 -r 500 4-50- 4-00 550 ■•* * ; v v Females IV V V jV -7 IV V 10° ! 12 ! 14- i 16 I ' ! 18 20 Latitude [N] 22 24- 26° x M. fascicularis 7 M. mulatta Fig. 29. Latitudinal variation of adult head and body length in samples of Macaco, fascicularis collected in the southern part of the Indochinese Peninsula and adjacent Isthmus of Kra, north of 10°N (cf. Fig. 10), compared with that in samples of parapatric M. mulatta collected in the northern part of the Indochinese Peninsula (Burma, Thailand, Laos, and Vietnam; specimens in amnh, bm(nh), bnhs, fmnh, mnhn, usnm, zrc, and zsi). 1 . Macaca f. aurea and M. f. fascicularis be- came differentiated in the Indochinese Peninsula and northern part of the Isthmus of Kra sometime before the last glacial maximum; this differentia- tion probably occurred on opposite sides of the mountain ranges that now form the border be- tween Burma and Thailand. Each of these sub- species probably dispersed from the mainland to shallow- water islands adjacent to its respective range during the last glacial maximum (ca. 1 8 Ka), when these islands were connected to the main- land by dry land. The population heterogeneous for the lateral facial crest pattern that inhabits cen- tral and eastern parts of the Indochinese Peninsula may indicate introgression or intergradation be- tween M. f. aurea and M. f. fascicularis, or it may indicate introgression or intergradation between M. fascicularis and M. mulatta; the population heterogeneous for the lateral facial crest pattern that inhabits eastern and southern parts of the Isthmus of Kra presumably indicates introgression or intergradation between M. f. aurea and M. f. fascicularis. The disjunct population of M. f. fas- cicularis in the southern part of the Indochinese 92 FIELDIANA: ZOOLOGY 135 130 — 125- 120- 115 110- o 105 120 Q3 "5 115 o 110- 105- 100 95- Males 90 10° 12 I x X ! V X X : - X X I ; I V *x X T IV T V j ! ;V V i V " p *■ V X X j j X T i v I V xx x X X ;V ;X % | V < i V ! ; j V i j f Females ■ i ' i ■ i V . ■ j i | , j H 16 18 20 Latitude (N) 22 24 26 c M. fascicularis v M. mulatta Fig. 30. Latitudinal variation of adult greatest length of skull in samples of Macaco fascicularis collected in the southern part of the Indochinese Peninsula and adjacent Isthmus of Kra, north of 10°N (cf. Fig. 21), compared with that in samples of parapatric M. mulatta collected in the northern part of the Indochinese Peninsula (Burma, Thailand, Laos, and Vietnam; specimens in amnh, bm(nh), bnhs, fmnh, mcz, mnhn, usnm, zrc, and zsi). Peninsula was geographically continuous with the population of the same subspecies in the Malay Peninsula during the last glacial maximum (Fig. 3). 2. Within M.f. fascicularis, a strong but incom- plete barrier to gene exchange has existed in the region of the Isthmus of Kra. This barrier has had a more profound effect on differentiation of pop- ulations of this subspecies north and south of the Isthmus of Kra than the Strait of Malacca (age 5- FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 93 550- 500- 450- 400- 550- 300- 250- ?oo- x X ! V V Males v i v .7) ! v ! 1 Vj y V ' % 550 c •g 500 i— 450 400 350- 300- 250 200 150 Females xx ; •v- r 10° 12 U 16 18 20 Latitude (N) 22 24 26 c x M. fascicularis v M. mulatta Fig. 31. Latitudinal variation of adult tail length in samples of Macaco, fascicularis collected in the southern part of the Indochinese Peninsula and adjacent Isthmus of Kra, north of 10°N (cf. Fig. 13), compared with that in samples of parapatric M. mulatta collected in the northern part of the Indochinese Peninsula (Burma, Thailand, Laos, and Vietnam; specimens in amnh, bm(nh), bnhs, fmnh, mnhn, usnm, zrc, and zsi). 18 K yr) has had on populations in the Malay the Isthmus of Kra, which approximately marks Peninsula and Sumatra. Many other vertebrates— the boundary between the Indochinese and In- including amphibians, reptiles, birds, and mam- domalayan faunal subregions (Chasen, 1940a, p. mals— also are differentiated north and south of x; Corbet & Hill, 1992, p. 3). 94 FIELDIANA: ZOOLOGY 3. Populations of M. f. aurea and M. f. fasci- cularis in the northern part of the Isthmus of Kra and Indochinese Peninsula tend to be transitional between M. fascicularis in the Sunda Shelf area and M. mulatta, which replaces M. fascicularis in the northern part of the Indochinese Peninsula. Stage IV, ca. 18 Ka P. Nias— Although P. Nias is a deep-water is- land, its population of M. fascicularis is not strong- ly differentiated (see above, Subspecific Taxono- my), which suggests relatively recent isolation of this population. The strait between P. Nias (area ca. 3,500 km 2 ) and the west coast of northern Su- matra is about 90 km wide, but the deep channel (120-180 m) in this strait is only about 20 km wide. The nonvolant mammal fauna reported for P. Nias by Kloss ([1928], p. 802; cf. Musser & Califia, 1982, pp. 11, 18; Musser &Carleton, 1993, pp. 614, 633, 652) is limited to M. fascicularis, one species each of tree shrew, binturong, pig, trag- ulid, and pangolin, two species of deer, and five species of murine rodents. Macaca fascicularis may have dispersed to P. Nias from Sumatra during the last glacial maxi- mum, ca. 1 8 Ka, when the intervening water gap was reduced to about 20 km. During subsequent isolation, the proximodorsal surface of the tail in the P. Nias population apparently became darker and the tail apparently became shorter. In all 12 specimens examined of M. fascicularis from P. Nias, the lateral facial crest pattern is transzygo- matic, as in Sumatran M. fascicularis. Philippines, Southern Islands— As previous- ly noted (Fooden, 1991, p. 6), dorsal pelage color in M. fascicularis in southern islands of the Phil- ippine Archipelago is paler than in M. f philip- pinensis, which inhabits western, northern, and eastern islands in this archipelago. The pale south- ern populations, which are allocated to the nom- inotypical subspecies M. f fascicularis, are indis- tinguishable in pelage color from populations in Borneo. This strongly suggests that progenitors of the southern populations dispersed from Borneo to the Philippines more recently— possibly during the last glacial maximum— than progenitors of M. f philippinensis (see above). Present distributions indicate that the relatively recent dispersal of M. f fascicularis to the southern Philippines occurred via the Sulu Archipelago (Fooden, 1991, p. 28). In Negros and Mindanao, a zone of mixed pop- ulations that include pale, intermediate, and dark individuals may be the result of interbreeding be- tween early-arriving M. f philippinensis and late- arriving M. f fascicularis. Stage V, < 18 Ka Shallow-Water Fringing Islands, Ex- cluding P. Bali— During the last glacial maxi- mum, ca. 1 8 Ka, sea level was approximately 1 20 m lower than at present, and shallow-water fring- ing islands— together with Sumatra, Borneo, and Java— were part of a single large landmass that extended from the Indochinese Peninsula to P. Bali (Fig. 3; Heaney, 1991a, p. 55). This permitted dispersal, or redispersal, of M. fascicularis over dry land to what are now shallow-water islands; as a result, populations of M. fascicularis in all of these present-day islands were genetically contin- uous. Subsequently, between 1 8 Ka and 5 Ka, sea level rose to its present level, and present-day shal- low-water insular populations of M. fascicularis became isolated. Following isolation, populations of M. fascicu- laris in many shallow-water fringing islands ap- parently have tended to become slightly darker, more erythristic, and/or smaller than their ances- tral stock, assuming that this stock is represented by core-area populations (Table 9; Appendixes 3, 4; Fooden & Albrecht, 1993, p. 533). The darkest shallow-water fringing-island population, M. f karimondjawae in P. Karimunjawa and presum- ably also in P. Kemujan, may have become iso- lated somewhat earlier than most other shallow- water fringing-island populations (Appendix 3). The distinctive dark crown patches of M. f atri- ceps in Ko Khram Yai and M. f condorensis in Con Son and Hon Ba presumably also developed in isolation following elevation of sea level since the last glacial maximum. During the same inter- val, dorsal pelage erythrism in M.f fascicularis in Borneo apparently differentiated somewhat from that in the same subspecies in Sumatra and the Malay Peninsula (Tables 3, 4). Despite postglacial isolation, many shallow-wa- ter island populations of M. fascicularis appear to have retained ancestral characters. Size is similar at similar latitudes in currently isolated popula- tions in the Malay Peninsula, Sumatra, and Bor- neo (Fig. 10; Fooden & Albrecht, 1993, p. 533). Dorsal pelage erythrism and tail length are more similar in populations in the Malay Peninsula and FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 95 Sumatra, which are now isolated, than in popu- lations in the Malay Peninsula and the Indo- chinese Peninsula, which are connected by dry land (Tables 3, 10). The infrazygomatic lateral facial crest pattern apparently has persisted in popula- tions in the Mergui Archipelago despite the post- glacial isolation of these islands from the Isthmus ofKra(Fig. 9). Lesser Sunda Islands: P. Bali, Nusa Penida, and P. Lombok— Pulau Bali is the only shallow- water fringing island in the Lesser Sunda group (Fig. 3; Kitchener et al., 1990, p. 1 1 1). The strait between Java and P. Bali is approximately 3 km wide and has a maximum depth of about 65 m. The strait between P. Bali and Nusa Penida is about 10 km wide and has a depth of more than 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 1 20 m for almost all of its width. All three of these islands are inhabited by M. fascicularis (Fig. 2C). The only other nonhuman primate known to inhabit the Lesser Sundas is one species of leaf monkey, Trachypithecus auratus, the range of which includes P. Bali and P. Lombok. In these two islands, T. auratus is represented by an endemic subspecies, T. a. kohlbruggei (P. H. Napier, 1985, p. 56; Weitzel & Groves, 1985, p. 402; Groves, 1993, p. 273). During the last glacial maximum, ca. 1 8 Ka, P. Bali was joined to Java, but Nusa Penida and P. Lombok were separate islands (Kitchener et al., 1990, p. lll;Heaney, 1991a, p. 55). At that time, populations of M. fascicularis and T. auratus in P. Bali presumably were genetically continuous with those in Java. It is unclear whether or not the last glacial maximum was the time of initial dis- persal of M. fascicularis and T. auratus from Java to P. Bali; these species may have dispersed from Java during a previous glacial cycle, and their in- sular populations may merely have been reunited during the last glacial maximum. In either event, sometime before 5 Ka, rising sea level separated P. Bali from Java and, conse- quently, separated populations of M. fascicularis and T. auratus in P. Bali from those in Java. Fol- lowing this separation, M. fascicularis in P. Bali apparently underwent reduction in body size (Figs. 1 5, 22; Appendix 7) and also underwent a marked shift in allele frequencies at blood-protein loci HbA-II, Pi, and Tf (Table 19). Concurrently, T. auratus in P. Bali apparently became subspecifi- cally distinct from T. auratus in Java. The characters that distinguish M. fascicularis and T. auratus in P. Bali from their respective conspecifics in Java link populations of these spe- cies in P. Bali to those in P. Lombok. This strongly suggests that, subsequent to their differentiation in P. Bali, populations of both species dispersed— over water— from P. Bali to P. Lombok, as pre- viously proposed by Kawamoto and Suryobroto (1985, p. 39); the dispersal of M. fascicularis to Nusa Penida, located between P. Bali and P. Lom- bok, presumably occurred at the same time. Whether the dispersal of M. fascicularis and T. auratus from P. Bali to P. Lombok was by natural rafting or by human transport is uncertain (cf. Ev- erett in Hartert, 1896, p. 593; Kitchener et al., 1990, p. 112). Stage VI, ca. 4.5 Ka Lesser Sunda Islands: P. Sumbawa-P. Tim- or— Macaca fascicularis is known to inhabit 13 of the deep-water Lesser Sunda Islands east of P. Lombok (listed in west-east order): P. Sumbawa, P. Moyo, P. Sumba, P. Rinca, Nusa Kode, P. Man- giatan, P. Seraya Besar, P. Flores, P. Adonara, P. Solor, P. Semau, P. Kambing, and P. Timor (Fig. 2C). This species reportedly is absent in P. Ko- modo and P. Padar (between P. Sumbawa and P. Rinca, north of P. Sumba), and in many nearby islets, and also is absent in P. Lomblen, P. Pantar, and P. Alor (east of P. Adonara and P. Solor, north of P. Timor) (for documentation, see Gazetteer, Appendix 2). No other nonhuman primate inhab- its any of the Lesser Sunda Islands east of P. Lom- bok. None of these islands were connected to P. Bali during the last glacial maximum, although some of these islands were interconnected at that time to form four larger islands (Lombok-Sumbawa- Moyo, Sumba, Komodo-Padar-Rinca-Kode- Mangiatan-Seraya Besar-Flores-Alor-Solor, and Semau-Kambing-Timor) (Heaney, 1991a, p. 55). The isolation of these deep-water islands is reflect- ed in their relatively poor nonvolant mammal fau- nas (Laurie & Hill, 1954, pp. 13 ff.). In P. Flores (area ca. 20,000 km 2 ), for example, Musser (1981, p. 134) listed 16 species of nonvolant mammals (shrews, 3; M. fascicularis; civet, 1; pigs, 2; deer, 1; porcupine, 1; rats, 5; mice, 2), of which some probably were introduced by humans. This con- trasts with 38 species of nonvolant mammals re- corded for P. Bangka (area ca. 1 2,000 km 2 ), a shal- 96 FIELDIANA: ZOOLOGY low-water island on the Sunda Shelf (Heaney, 1 984, P. 12). Four items of partly contradictory evidence seem most relevant to attempting an interpretation of the history of M. fascicularis in deep-water Lesser Sunda Islands east of P. Lombok: (1) Dorsal pelage color in these islands is essentially similar to dorsal pelage color in Java, P. Bali, and P. Lombok (Ap- pendixes 3, 4). (2) External and cranial size in these islands is similar to size in P. Bali and P. Lombok (Figs. 15, 16, 22, 23); size is greater in Java. (3) Blood-protein allele frequencies at loci HbA-II, Pi, and Tf in these islands are similar to allele fre- quencies in Java (Table 1 9); frequencies of these alleles are divergent in P. Bali and P. Lombok. (4) Subfossils of M. fascicularis in P. Timor cave de- posits appear relatively late (ca. 4.5 Ka), concur- rently with the appearance of remains of domestic animals and pottery (see above, Fossils and Subfossils). The evidence of dorsal pelage color and Timor cave deposits suggests relatively recent dispersal of M. fascicularis to deep-water Lesser Sunda Is- lands east of P. Lombok. The blood-protein evi- dence indicates that the differentiated populations of M. fascicularis in P. Bali and P. Lombok prob- ably were not directly involved in dispersal of this species to islands east of P. Lombok. Assuming that the differentiated populations in P. Bali and P. Lombok inhabited those islands before M. fas- cicularis dispersed to islands east of P. Lombok (see above), the implication is that populations of M. fascicularis east of P. Lombok were derived from a source in Java, or elsewhere west of P. Bali. This geographic incongruity, together with the ar- cheological evidence of the Timor cave deposits, strongly supports the hypothesis that the popu- 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), Kawamoto and Suryobroto (1985, p. 39), and Glover (1986, p. 159). Multiple introductions would be required to es- tablish populations of M. fascicularis in all of the islands east of P. Lombok that it now inhabits. If Java was the source of these introductions, exter- nal and cranial size became convergently reduced in these islands. This is inconvenient to the hy- pothesis, but not impossible, considering the fre- quency of insular dwarfing in M. fascicularis (see above, External Measurements and Proportions; Cranial Characters). Auffenberg (1981, p. 242) suggests that the ab- sence of M. fascicularis in P. Komodo (area 393 km 2 ) and P. Padar (13 km 2 ) may be attributable to an insufficient year-round supply of fruit in these islands. This is puzzling, because small islands im- mediately east of P. Komodo and P. Padar support populations of M. fascicularis (P. Rinca, 278 km 2 ; Nusa Kode [= P. Oewada Sami], 9.6 km 2 ; P. Ser- aya Besar [= P. Seraja], 3.6 km 2 ; P. Mangiatan [= P. Mengjatan], 1 km 2 ). The insular distribution of M. fascicularis in this area may be affected in some way by the distribution of the Komodo giant mon- itor, a known predator of these monkeys (see above, Natural History) that is restricted to the following seven islands (listed in west-east order): P. Ko- modo, Nusa Mbarapu (0.6 km 2 , occasionally vis- ited by giant monitors), P. Padar, P. Rinca, Nusa Kode, Gili Motang (= Gili Mota), and western P. Flores (Auffenberg, 1981, p. 40). M. fascicularis coexists with the giant monitor only in three of the more easterly of these islands (P. Rinca, Nusa Kode, and P. Flores). The relatively recent dispersal of M. fascicularis to deep-water Lesser Sunda Islands, indicated by evidence cited above, is somewhat surprising. This species has been present in east-central Java since ca. 1 Ma (see above), and, during the maximum Pleistocene glaciation, the widest water gap be- tween Java and Sumbawa may have been only 400 m or less (Kitchener et al., 1 990, p. 1 1 0). However, as indicated by Heaney (1991a, p. 57), during gla- cial maxima, rainfall in the Lesser Sundas was much lower than at present. Perhaps forest cover was inadequate to sustain M. fascicularis in these islands during glacial maxima, when low sea level and narrow water gaps might otherwise have fa- cilitated the eastward dispersal of this species. Acknowledgments The generous cooperation of curators and staff members of institutions in which specimens ex- amined are preserved (see Introduction) has been indispensable to the conduct of this research and is gratefully acknowledged. For valuable supple- mentary advice and assistance, I also thank the following: H. Abdulali (bnhs), N. Aggimarangsee (Chiang Mai University, Thailand), M. Aimi (pri), G. H. Albrecht (University of Southern California, Los Angeles), W. Y. Brockelman (Mahidol Uni- versity, Bangkok), M. D. Carleton (usnm), S. FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 97 Chakraborty (zsi), C. Fleming (amnh), W. K.-H. Fuchs (amnh), L. Gordon (usnm), C. P. Groves (Australian National University, Canberra), L. R. Heaney (fmnh), P. D. Jenkins (bm(nh)), Y. Ka- wamoto (pri), J. C. Kerbis Peterhans (fmnh), J. W. Koeppl (fmnh), S. M. Lanyon (fmnh), G. G. Musser (amnh), Shukor Md. Nor (fmnh), B. D. Patterson (fmnh), D. P. Schmidt (usnm), S. Un- nithan (bnhs), and P. F. D. Van Peenen (U.S. Navy, retired, Chicago). I am also grateful to three re- viewers, who painstakingly scrutinized the manu- script and provided numerous helpful suggestions. Literature Cited Abdulali, H. 1967. The birds of the Nicobar Islands, with notes on some Andaman birds. Journal of the Bombay Natural History Society, 64: 139-190. Adi Haji Taha. 1985. The re-excavation of the rock- shelter of Gua Cha, Ulu Kelantan, West Malaysia. Federation Museums Journal, n.s., 30: i-xi, 1-134. Aggimarangsee, N. 1992. Survey for semi-tame col- onies of macaques in Thailand. Natural History Bul- letin of the Siam Society, 40: 103-166. Aimi, M. 1981. Fossil Macaca nemestrina (Linnaeus, 1766) from Java, Indonesia. Primates, 22: 409-413. Aimi, M., and F. Aziz. 1985. 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Further observations on the breeding of primates, with special reference to the suborders Lemuroidea and Tarsioidea. Proceedings of the Zoo- logical Society of London, 1932: 1059-1075. [For date of publication, see Proceedings of the Zoological So- ciety of London, 1933: 229.] Zumpe, D., and R. P. Michael. 1983. A comparison of the behavior of Macaca fascicularis and Macaca mulatta in relation to the menstrual cycle. American Journal of Primatology, 4: 55-72. . 1990. Effects of the presence of a second male on pair-tests of captive cynomolgus monkeys {Macaca fascicularis): Role of dominance. American Journal of Primatology, 22: 145-158. FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 119 Appendix 1: Specimens Examined (Total 2,049) Macaca fascicularis fascicularis, Total 1,550 Skins and Skulls, 739 CAMBODIA, 2. Mainland '(2), Siemreab— mnhn 1929.460; no locality— mnhn 1961.61 1. INDONESIA, 456. Bali (21), Banjoe Wetan- mzb 6521-6523; Batoe-meringgit— mzb 2001; Bratan, Gunung, amnh 107555-107557; Desa Poetjang— rmnh Coll. No. E85 (external measure- ments from Sody, 1933, p. 94); Gilimanuk— amnh 107561-107568; Gitgit-NMS 16632; Jembrana- rmnh Coll. No. El 39 (external measurements from Sody, 1933, p. 94); Sendang— rmnh Coll. Nos. E34, E64, E74 (external measurements for all three from Sody, 1933, p. 94); Bangka (4), Pamuja, Tan- jung— usnm 124863; Rengsam, Tanjung— usnm 1247 10; no locality-RMNH Coll. Nos. Bk52, Bk53; Batam (4) "Tanjong Sauh"-BM(NH) 1909.4.1.29, bnhs 5080; Tanjong Turut-BM(NH) 1909.4.1.27, 1909.4.1.28; Bawean (8), no locality-MZB 1841- 1846, usnm 151829, 151830; Belitung (2), Batu, Tanjung-USNM 124969, 124970; Bengkalis (1), Kapos Tinggi— usnm 143582; Benua (2), no lo- cality-usNM 101638, 101639; Bintan (8), north coast— usnm 115676; Pasir Panjang— bm(nh) 1909.4.1.26, usnm 115677; Sungei Biru-BM(NH) 1 909.4. 1 .23-1 909.4. 1 .25; no locality-ZRC 4-093, 4-094; Borneo: Kalimantan (93), Ambawang, Sun- gai — usnm 142225; Badang— amnh 103730, 103731, 106024-106026, mzb 6513; Berau, Sun- gai— usnm 196817-196822; Birang, Sungei— usnm 196825 (includes skinned head in fluid); Buntok— bm(nh) 1910.4.5.23; Goson Djerong — usnm 196814; Hantakan — usnm 521837; Karangin- tan— bm(nh) 1910.4.5.22; Karangmumus, Sun- gai— usnm 196816; Karangtigau, Tanjung— usnm 196824; Kembangjanggut— mzb 8130; Liang Koe- boeng— rmnh 160, 178; Loa Bambam— usnm 196815; Long Peleben-AMNH 106027, mzb 6514; Mahakam, Sungai — usnm 196813; Merah — mzb 1164; Muaratewe — bm(nh) 1910.4.5.19, 1910.4.5.20; Parit-AMNH 103644-103658, 103660-103663, mzb 6515-6517, zmb 48005, 48006, 92191; Pelawan, Sungai-USNM 198301; Perbuah-AMNH 107091-107096, 107098, 107099, mzb 6519, 6520; Putussibau-ANSP 14038, 14039, 14044; Riam- amnh 106283-106285, mzb 6518; Roema Manoeal— rmnh 23, 24; Sembak- ung, Sungai— mzb 576, 6512; Semitau— rmnh 6; Sintang— ansp 14037, 14042, 14043; Tangarveng I. -usnm 1 96804; Telang— usnm 521838; Tibang, Mt.— mcz 22277; no locality— ansp 14041, bm(nh) 1856.9.3.6, zsbs Coll. Nos. 272, 738, 788, 1227, 1354, 1373, 1527;5w/a«(3),south-usNM 144419; no locality— zrc 4-098, 4-099; Durian (2), no lo- cality— mzb 241, 242; Flores (7), Mbura— mzb 2384-2387; Rana Mese-NMS 1 6630; Sano, Wai- mzb 2388; no locality— bm(nh) 1864.4.12.1; Gal- ang (4), no locality— zrc 4-100-4-103; Java (84), Bandung, near— usnm 521839; Bantargebang— mcz 12757, usnm 156291, 156292, 156295; Ban- yuwangi— rmnh 959; Batoeraden— rmnh Coll. No. 89C; Bogor— mzb 1884; Camara— mzb 6482; Can- diroto— rmnh Coll. No. 107B; Cihara— usnm 156456, 156458; Cikujang-MZB 6483, 6485; Cil- acap-BM(NH) 1909,1.5.27, 1909.1.5.28; Cire- bon— amnh 101891, 102016; Ciremay, Gunung - amnh 102015, 102017-102022; Ciwangi-BM(NH) 1909.1.5.29, 1909.1.5.30; Depok-Mcz 12755 (ex- ternal measurements from usnm field cat.), 1 2756; Gedangan— rmnh Coll. Nos. 74, V9, V10; Indra- mayu— mzb 2961, 2962; Jasinga— mzb 2052, 3189; Java, West— rmnh 2b, 3c, 4h; Kaligoea— mzb 574, 575; Kalipucang-BM(NH) 1909.1.5.19, 1909.1.5.32, 1909.1.5.33; Linggajati-AMNH 101811; Majalengka-AMNH 101808-101810; Pangandaran— bm(nh) 1909.1.5.24; Pangonan— rmnh Coll. No. M40; Pasir Carolina— mzb 6484; Pelabuhanratu, Teluk— usnm 156457, zrc 4-134; Salak, Gunung— rmnh Coll. No. A 147; Saron- gen— mcz 12759; Singkil, Gunung— rmnh 5919o, 5919p, 5919q; Tamadjaija— mcz 12758; Taman- sari— bm(nh) 1954.51, zrc 4-133; Tasikmalaya— bm(nh) 1909.1.5.31, mzb 1604-1609; Tiloe, Goenoeng— rmnh 5919n; Tjeringin— rmnh Coll. Nos. 13f, 25f, 26f; Wonokojo— rmnh Coll. Nos. Won3, Won4, Won5; no locality— nms 16628, rmnh 2505a, 2505b, 2527, 3094, 3095, 3097, zmb 113, zmuz 11627 (skin)/11630 (skeleton); Kan- gean (4), no locality — bm(nh) 1 9 10.4.6. 1 — 1910.4.6.4; Karimata (2), Pai, Teluk — usnm 125101, 125102; Karimun (7), Mensuda Bay- usnm 122849; Monos-BM(NH) 1909.4.1.34, 1909.4.1.35; Pemeral-BM(NH) 1909.4.1.30- 1909.4.1.33; Kundur (2), Selatbliat-BM(NH) 1909.4.1.36, 1909.4.1.37; Lagong (1), no locali- ty— usnm 104853; Laut (1), no locality— usnm 104854; Lingga (2), no locality— usnm 101602, 101603 (external measurements from Miller, 1906c, p. 276); Lombok (3), Pussuk forest— mzb 6528; Sewela-NMS 16634, zmb 92306; Mapur(l), Mentigi— zrc 4-097; Matasiri (1), east coast— usnm 151831; Mursala (2), no locality— usnm 120 FIELDIANA: ZOOLOGY 114559, 114560; Natuna Besar (2), Binjai, Sun- gai— zrc 4-139, 4-140; Nguwal (6), no locality— zrc 4-104-4-109; Nias (1 1), Hilisimaetano— ansp 20404-20406; Lafau-USNM 141372; Samasa- ma— usnm 141371; Siaba, Teluk— usnm 121868- 121871; Soliga-Mcz 36029, 36030; Pelapis Ten- gah (1), no locality— mzb 2914; Penida (3), no lo- cality— amnh 107558-107560; Serasan (2), no lo- cality— usnm 104852, zrc 4-144; Serutu (1), no locality— mzb 29 1 5; Siantan (3), no locality —usnm 101711, zrc 4-136, 4-137; Subi-kecil (3), no lo- cality— zrc 4-141-4-143; Sugi (1), no locality— usnm 115675; Sumatra (123), Batangkuis— zsbs Coll. No. 9; Bungur-Buikt-ZMB A25.09; Deli, Sungai— ansp 20217, 20219; Goenoengsetan- Meloewak— mzb 6481; Indragiri, Sungai— usnm 113169; Kalianda- amnh 1 02904-1 02906; Kam- bang— zmb A25.09; "Kg. Baru"— zsbs Coll. Nos. 156, 176, 180; Kotabumi-ZRC 4-124; Lesten- mzb 6480; Lubuklinggau— amnh 102211; Me- dan-ANSP 20221, zsbs Coll. Nos. 1/10, 5, 6, 15, 16, 21, 22, 26, 30-32, 35, 36, 42, 45-48, 51-53, 68 (Bruegel), 68 (Widnmann), 76, 79, 82, 89, 91, 92, 101, 105, 134, W2,Wll;Medan, forest near- zsbs Coll. No. 11; Muaradua— amnh 102763- 102770; Padang-RMNH 518, 981, 1051; Padang highlands— rmnh e2; Pajo— bm(nh) 1879.6.28.4; Pematangsiantar— mcz 35937, 35938; Sandaran Agong— zrc 4-110; Sanggul, Bukit— amnh 106565-106569, mzb 6477-6479; Serdang-ZSBS Coll. No. 134;Siulakderas-BM(NH) 1919.11.12.8; Sukadana— amnh 102907-102910; Sumatra, [east- central]— zmb A 14.08/37; Sumatra, east coast— zsbs Coll. Nos. 105, 135, 156, 166-168, 174; Su- matra, west coast— bm(nh) 1882.7.24.1; Tanjung- morawa— zsbs Coll. Nos. 7 1 [b], 72, 1 37; Tapanuli, Teluk— usnm 114505, 114506; Tapung-kanan, Sungai— zmb A14.08; Tarusan, Teluk— usnm 141145; Telukbetung— fmnh 14805; Telukpan- ji— zrc 4-111; no locality— NHMBa 3347, 3348, rmnh 1479, usnm 271018, zsbs Coll. Nos. 28, 35, 36, 38, 58, 64, 67, 93, 107, 112, 135, 157, 184, 1 85, 203-205, 2 1 2, 232; Sumba (8), Langgaliroe- mzb 6526, 6527, zmb 92075; Mao Marroe— mzb 867-869; Payeti-Kambaniru— mzb 865, 866; Sumbawa (8), Batudulang— mzb 6529, 6530, zmb 92307, 92308; Dompu-NMS 1025; Kambing, Pu- lau— mzb 6531; Ntori— zmb 92309; Oo vicinity— nms 1026; Tanahbala (2), no locality— usnm 121802, 121803; Timor (9), Kuatnana-ZSBS 1911/ 2108; Lelogama-ZSBS 1911/2102, 1911/2104, 1911/2105; Mutis, Gunung— mzb 6524, zmb 92 1 3 1 , 92 1 32; Nikiniki-MZB 6525; Timau, Fatu- zsbs 1911/2101; Tuangku (3), north coast— usnm 114408-114410; Uwi (1), no locality-uSNM 101666. MALA YSIA, 1 63. MALA YSIA: SABAH, Bang- gi (5), Banggi, Pulau, south— zrc 4-116-4-118; Karakit-FMNH 140939; Sabor-FMNH 140938; Borneo (74), Abai-Mcz 35600, 35606, 35611- 35613, 35629, 35641, 35642, 35693, 35694, 35700, 35701, 35722, 35724, 35726, 35727, 35729, 35731, 35732, 35744, 35761; Betotan,ZRC 4-115; Bongkabong-Mcz 37409, 37411, 37412; Darvel Bay— zmb A86.10; Garau— mcz 37349, 37351, 37414, 37418; Kampong Bundu Tuhan- usnm 292555-292558; Kampong Kiau — mcz 37348, 37352, zrc 4-1 20, 4-121 ; Keningau-AMNH 188343; Kenokok zrc 4-122; Kiaulan— mcz 37350, 37353; Kinabatangan, Sungai — usnm 19193; Kretam Besar, Sungai— fmnh 68700; Kre- tam Kechil, Sungai— fmnh 68699; Lahad Datu— zmb A89.04; Papar— usnm 317190; Ranau— mcz 37346, usnm 3 1719 1-3 17 195; Rayoh-ZRC 4-1 19; Rugading— mcz 37357; Sapagaya Forest Re- serve— fmnh 68701, 68702; Sibuga Besar, Sun- gai— fmnh 33545; Talibang— mcz 37354, 37406, 37413; Tawau— amnh 31288; Tempasuk, Sun- gai— mcz 37417; Tenompok— mcz 37415, 37416; Tinonkok— mcz 37347; Tuaran-Kampong Tengh- ilan Road-Mcz 37355, 37356, 37405, 37408; no locality-uSNM 344989-344992; Sebatik (2), no locality— usnm 175896 (skin and skull)/ amnh 30620 (postcranial skeleton), usnm 175897. MA- LAYSIA: SARAWAK, Borneo (15), Baram, Ba- tang-BM(NH) 1894.6.12.2, USNM 83945; Belaga or Kalulong, Bukit-BM(NH) 1951.66, 1951.68; Bukar, Sungai— zrc 4-113; Entawa, Tanjong— bm(nh) 1955.710; Kuching-SMK 092/3, 092/8; Kuching, probably— zmb 4939; Long Ekang— fmnh 88582; Paku, Saribas-BM(NH) 1955.711, 1955.712; Pelandok, Sungai-ZRC 4-112; Punang, Sungai— zrc 4-114; Selikan, Bukit— usnm 83944. MALA YSIA: WEST MALA YSIA, mainland (29), Benom, Gunong— zrc 4-065, 4-066; Changkat Mentri— bm(nh) 1955.1510, zrc 4-064; Cherak- ah, Bukit-BM(NH) 1955.1516, zrc 4-069; Dusun Tua— zrc 4-068; Gaik Liew Estate — bm(nh) 1955.1515; Hantu, Tanjong -zrc 4-063; Kelang Road-BM(NH) 1971.2611; Melaka — bm(nh) 1875.3.29.1, rmnh 41g; Nanas, Bukit— bm(nh) 1939.180, 1939.181, 1955.1512; Nyalas-ZRC 4-078 (external measurements from Weitzel et al., 1988, p. 107); Pong-BM(NH) 1934.7.18.1; Port Dickson -bm(nh) 1939.229 (skin)/RCS(OM) A85.4 (skull), zrc 4-073-4-075; Tanjong Panjair— bm(nh) 1955.1514, zrc 4-067; Tanjong Tuan, Keramat— 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, 1934.7. 18.3; no locality-usNM 301 758; Acheh(l), no locality — bm(nh) 1955. 1 5 IS; A ur (2), no local- ity- bm(nh) 1955.1520, zrc 4-080; Burau (1), no locality— bm(nh) 1955.1519; Langkawi (1), no lo- cality— bm(nh) 1 909. 11.1.1; Pemanggil (2), no lo- cality-BM(NH) 1955.1508, usnm 1 12500; Pinang flj(8), Pantai Krachut-ZRC 4-062; Telok Bahang bm(nh) 1955.1521, zrc 4-056-zrc 4-061; Pinang [2] (2), no locality -bm(nh) 1949.426, 1955.1523; Pintu Gedong (3), no locality— bm(nh) 1955.1517, zrc 4-070, 4-071; Redang (2), no locality— zrc 4-054, 4-055; Tinggi (5), no locality— bm(nh) 1909.4.1.21, 1909.4.1.22, 1955.1527, zrc 4-131, 4-132; Tioman (11), Juara, Telok — bm(nh) 1908.1.25.1, 1908.2.25.1, 1955.1524, 1955.1525, zrc 4- 1 25-4- 1 27; Sedagong, zrc 4- 1 29, 4- 1 30; no locality-BM(NH) 1955.1526 (skin)/zRC 4-128 (skull), usnm 101744. PHILIPPINES, 43. Basilan (3), Basilan I., east end— usnm 144666; Camp No. 4 and Camp No. 5, between— usnm 144665; Isabela— usnm 125326; Cagayan Sulu (1), no locality— usnm 125325; Mindanao (39), Bucong— fmnh 65440-65444; Canibongan— fmnh 67737-67739; Gubat— fmnh 67725, 67729-6773 1; Libu-FMNH 67727, 67728; Mamara— fmnh 67717, 67718; Pulunbato, Mt.— bm(nh) 1880.1 1.24.1; San Ramon-FMNH 33507- 33511; Sigayan-FMNH 67722, 67732, 67740- 67742; Situbo-FMNH 67719-67721, 67723, 67724; Tacuta — fmnh 67733-67735; Tampa- Ian— fmnh 67736; Zamboanga— usnm 144667, 144668, 144698, SINGAPORE, 1. Singapore (7), Botanical Gar- den-ZRC 4-082, 4-084; Changi-BM(NH) 1909.4.1.20; Punggol-BM(NH) 1955.1513, zrc 4-086; no locality— zmb 5444, zrc 4-090. THAILAND, 60. Mainland (32), Aranyaprath- et— mcz 35931; Ban Huai Maenam Noi— fmnh 99649, 99650; Ban Nong Kok-ZRC 4-034; Ban Nong Put— ctnrc (catalog number unknown); Ban Pak Nam Pho-Mcz 23812, 23813, zrc 4-031; Ban Phra Muang-USNM 83271, 83273; Ban Phu Toie-FMNH 99639, 99640; Ban Sai Kau-BM(NH) 1903.2.6.3; Ban ThapPlik, 1 kmE-FMNH 105654; Ban Thap Plik, 1 km NE-fmnh 105653; Kam- pong Biserat-BM(NH) 1903.2.6.1, 1903.2.6.4; Kantang— zrc 4-032; Kata Taek— fmnh 99651- 99656; Khao Rang Kai— ctnrc (catalog number unknown), fmnh 105689; Laem Sing mountains— usnm 251662; Lat Bua Khao, Sathani— usnm 236631, zrc 4-033; Tham Hom-FMNH 105649; Tyching-USNM 83274, 83275; Butang(l), no lo- cality— bm(nh) 1955.1511; Chang (3), no locali- ty-BM(NH) 1915.1 1.4.7, usnm 201551, zrc 4-039; Kut (8), no locality-BM(NH) 1915.11.4.8- 1 9 1 5. 1 1 .4. 1 0, 1 976. 1 826 (skin)/zRC 4-040 (skull), usnm 201552, 201553 (external measurements from Kloss, 1916a, p. 67), 254741, zrc 4-041; Phangan (2), southwest— zrc 4-044, 4-045; Phi Phi Don (4), no locality —zrc 4-046-4-049; Samui (2),west-BM(NH) 1955. 1522, zrc 4-050; Talibong (2), no locality— usnm 83272, zrc 4-051; Tarutao (6), Sungei Adang-BM(NH) 1909.11.1.2, 1909.1 1.1.3; Telok Wau-BM(NH) 1955. 1509, zrc 4-053; no locality-usNM 123990, 123991. VIETNAM, 8. Mainland (6), Cochin China- bm(nh) 1881.6.30.2, mnhn 1882.2; Ho Chi Minh City— fmnh 33505; Ho Chi Minh City, Zoological Garden-MNHN 1929.459; Sontra Peak, 3.9 km W, 0.3 km S-usnm 356968; Xa Trang Bom-ZRC 4-020 (external measurements from Weitzel et al., 1988, p. \ 14); Phu Quoc(2)-bm(nh) 1928.7.1.10, mnhn 1929.455. Skins only, 285 CAMBODIA, 1. Mainland (1), no locality - RMNH 38. CAMBODIA OR VIETNAM, 2. Mainland (2), "Camboja or Cochinchine"— bm(nh) 1 878.6. 1 7.4, 1878.6.17.5. INDONESIA, 241. Bali (1), Bratan, Danau- bm(nh) 1913.3.6.7; Bangka (1), Simpang— zsbs Coll. No. 25; Borneo: Kalimantan (82), Borneo, southeast — rmnh 35v (skull inside); Borneo, southwest— rmnh 32s, 33t, 34u (skulls inside); Kampong Hadjak— irsn 9796; Karangtigau, Tan- jung— usnm 196823; Muaratewe — bm(nh) 1910.4.5.21; Parit-AMNH 103659 (skull not lo- cated, cranial measurement from G. H. Albrecht, pers. comm., Oct. 1991), zmb 92190; Pelaihari— rmnh 36w (skull inside); Perbuah— amnh 107097; Poelau— rmnh 229 (cf. Hooijer, 1962b, p. 46, no. 48); Rantau— rmnh 4617 (cf. Hooijer, 1962b, p. 44, no. 11); Riam— amnh 106282; Senoeang— amnh 1 07 1 00 (male skin mismatched with female skull); Sintang— ansp 14036, 14040; no locality— zmb (unnumbered, 24 Apr. 1902), zsbs Coll. Nos. 3PS,4PS,7PS, 10-12, 1 5 Guvenile male), 15 (adult male), 16, 21, 36(450), 48, 157, 201, 205, 276, 277, 297, 298, 306, 372, 373, 445(316), 587, 616- 618, 694, 701, 709, 710, 730, 740, 741, 743, 749, 764-766, 771, 780, 1013, 1014, 1228, 1358, 1359, 1372, 1378, 1386, 1387, 1414, 1437, 1438, 1452, 1464, 1465, 1477, 1478, 1488, 1505, 1510, 1534, unnumbered (Breugel), unnumbered (Breugel, "von Zengen"); Java (25), Bantargebang— mcz 12760; Gedangan— rmnh (unnumbered); Indra- mayu— mzb 2939, 2960; Kawarasan— rmnh 2067; 122 FIELDIANA: ZOOLOGY Pangandaran— bm(nh) 1909.1.5.26; Pangrango, Gunung— rmnh 2067, 2067/II (skulls inside); Se- marang— zsbs (unnumbered); Tasikmalaya— mzb 1610; Java, West— rmnh 1, 5e, 6, 7g (skulls in- side), 8; no locality— NHMBa 3131, 3161, 3352, rmnh 9i, lOj, Ilk, 121, 13m, 14 (skulls inside); smtd (unnumbered); Kambing (1), no locality— rmnh 30p (skull inside); Laut (1), no locality— bm(nh) 1939.1054 (skull inside); Natuna Besar (6), Sinubing-BM(NH) 1939.1053 (skull inside); Ulu, Sungai— zrc 4-138; no locality— bm(nh) 1939.1055, 1964.428 (skulls inside), mcz 6660, 6661 (skulls inside); Mas (1), no locality— zrc 4-135 (skull not located); Pejantan (1), no locali- ty— zsbs I; Sumatra (114), Batangkuis— zsbs Coll. Nos. 3, 1 3, 1 5, 38, 47, 88, 96, 97, 1 97, 200; "Bran- dan?"-ZMB 38542; Deli, Sungai— ansp 20218, 20220 (fetuses in fluid); "I. Lendung"— zsbs Coll. Nos. 32, 74, unnumbered juvenile; "Kg. Baru"— zsbs Coll. Nos. 63, 77, 79, 81, 113; Medan-ZSBS Coll. Nos. 1, 6/II, 7, 7/II, 8/II, 9/II, 14, 24, 28 (juvenile), 29, 42/111, 43, 47/11, 52, 58, 60, 62, 67, 67/11, 71, 74, 75, 77, 80, 81, 81/11, 82, 90, 93, 94, 98, 99/11, 100/11, 102, 103, 104, 106, 111, 115, 1 16/VIII, 119, 124/III; Menam-ZMB 40820; Pa- dang— rmnh 44x, 45y, 46z (skulls inside), 47a, 48b, 49c, 50d (skulls inside), zmuz 1 1625, 1 1626; Palembang, Kotamady— zsbs Coll. Nos. 22, 23; Pangkalansusu— usnm 502457 (infant in alcohol); Sumatra, east coast— zsbs Coll. Nos. 1 58, 1 70, 1 7 1 , 189; Sumatra, south-ZMB 38540, 38541; Tan- jungmorawa— zsbs Coll. Nos. 60, 73[b], 75[b]; no locality— mnhn 362/233, zmb Coll. Nos. 9, 25, unnumbered juvenile, unnumbered male, zmuz 11619 (skull inside), zsbs Coll. Nos. 14, 27, 52, 61, 68, 70, 78, 78/1905, 95, 111, 115, 117, 123, 141, 155, 198, 199, 201, 202, 216, 235, 236, 242; Sumbawa (1), Batudulang— nms 16631 (cranial measurements from Mertens, 1936, p. 319, and G. H. Albrecht, pers. coram., Oct. 1991); Timor (6), Amarassie— rmnh 29o (skull inside); Bo- kong— zsbs 1911/2110; Lelogama— zsbs 1911/ 2103, 1911/2106, 1911/2107; Nikiniki-ZMB 92133; island unknown (1), Sunda Islands— zmb 111. INDONESIA OR MALAYSIA, 1. Borneo (1), Sendung— zmb (unnumbered). MALAYSIA, 24. MALAYSIA: SABAH, Borneo (2), Abai-Mcz 35695; Tuaran-Mcz 37407 (male skin mismatched with female skull); Sebatik (1), no locality- amnh 30619. MALAYSIA: SARA- WAK, Borneo (15), Baram, Batang— nms 4689, smk 092/1 3 (skull inside); Belaga or Kalulong, Bu- kit— bm(nh) 1 95 1 .67 (female skin mismatched with male skull); Dulit, Bukit— smk 092/9 (skull in- side); Kuching— smk 092/5 (skull inside); Ku- ching, 10th mile— smk 092/12 (skull inside); Ling- ga-SMK 092/2, 092/7 (skulls inside); Miri-Mcz 6662 (skull inside); Miri district— NHMBe 817; Penrissen, Gunong-SMK 092/4, 092/1 1/A3.312 (skulls inside); Punang, Sungai— smk 092/6, 092/ 1 0; Segobang, Sungai -smk 092/ 1 1/47 1 3 (skull in- side); Dindding(l), no locality— bm(nh) 1905.3. 1.3. MALAYSIA: WEST MALAYSIA, mainland (4), Melaka-MNHN 1848.401, 1848.402 (skulls in- side), rmnh 40, 42 (skull inside); Aur (1), no lo- cality— zrc 4-079. PHILIPPINES, 3. Mindanao (1), Zamboanga del Sur Prov.— siconbrec 1586 (living captive); Tawitawi (2), no locality— siconbrec 1225, 1475 (living captives). SINGAPORE, 5. Singapore (5), Botanical Gar- den— zrc 4-081 (skull inside), 4-083; Punggol— zrc 4-087, 4-088; Sembawang, Sungai— zrc 4-089. THAILAND, 3. Mainland (3), Bangkok— mnhn 1219 (skull inside); Ban Thap Plik, 1 km E— fmnh 105655 (infant in fluid); Kampong Biserat— bm(nh) 1903.2.6.2. VIETNAM, 5. Mainland (5), Ho Chi Minh City-NMS 5786, 5787; Ho Chi Minh City, Bo- tanical Gardens— bm(nh) 1928.7.1.9, mnhn 1929.457; Tay Ninh-BM(NH) 1928.7.1.7. Skulls only, 526 INDONESIA, 434. Bali (2), Bratan, Danau- bm(nh) 1913.3.6.5 (possibly skull of bm(nh) 1913.3.6.7, skin only); Jembrana— rmnh Coll. No. E136 (external measurements from Sody, 1933, p. 94); Bangka (173), Jebus-RMNH Coll. No. Bk7; no locality— rmnh Coll. No. Bk74, zluu 1, 2, 4- 16, 18, 20, 22, 25-30, 32-37, 39^13, 45-62, 64- 67, 73-75, 77-105, 106 (juvenile cranium only), 106 (adult mandible only), 107-112, 113, (juve- nile cranium only), 113 (adult mandible only), 1 1 4- 116, 117 (cranium and mandible), 1 17 (mandible only), 118, 119, 120 (cranium only), 120 (man- dible only), 121-134, 136, 137, 140-165, 167- 171, 173-178, 179 (mandible only), 181-184 (mandibles only), 185, 187-189 (mandibles only), zsbs 1964/232; Bintan (2), Pasir Panjang— zrc 4-092, Sungei Biru-ZRC 4-091; Borneo: Kali- mantan (24), Borneo, southeast— rmnh p, zmb 38543; Gosong Djerong— usnm 196827; Jembay- an, Sungai-USNM 199181, 199183, 199184, Kar- angan, Sungai— usnm 198300; Kariorang— zmb 48494, unnumbered; Loa Bambam— usnm 196826; Long Peleben— mnhn 1974; 144; Purukcahu— bm(nh) 1910.4.5.18; Tanjung— NHMBa 3747; no locality— amnh 107100 (female skull mismatched FOODEN: SYSTEMATIC REVIEW OF MACACA FASCICULARIS 123 with male skin), rcs(om) G 1 66.3 1 , zsbs 1 944/3009, Coll. Nos. 12-4-12-6, 20, 363/368, 748, 848, 864; Java (123), Bandung— rmnh 3438, zluu (unnum- bered); Cirebon — nms 6974, rcs(om) A85.6, A85.61-A85.65; Gedangan-RMNH Coll. VI, un- numbered adult males (2), unnumbered infant (1); Java, West— rmnh cat. ost. b-cat. ost. e (4); Jock- aboemi— zmb A34.09; Malabar-ZLUU (unnum- bered); Manggis-ZSBS 1909/4062, 1909/4601, unnumbered specimens (7), Margomulio, Gun- ung— zsbs (unnumbered); Ngawi— rmnh 4660; Pangandaran bm(nh) 1909.1.5.25 (with mounted skin, not examined); Pangrango, Gunung— nhrm Z2923-Z2925, Z3146-Z3150, Z3152-Z3154, Z3165, Z3166, Z3292, Z3293, Z3295, Z3314, Z3316, Z3320-Z3322, Z3324 (adult female), Z3324 (subadult female), Z3325, Z3328-Z3330; Salak, Gunung— rmnh Coll. Nos. A 137, A221; Tjeringin— rmnh Coll. No. 12f; Tulungagung— rmnh 4650-4655; no locality— ansp 12, NHMBa (unnumbered), nhrm Z2698-Z2700, Z2702, Z2709-Z2711, Z2718-Z2720, Z2726-Z2729, Z2730 (mandible only), Z273 1 , Z2739, Z3 1 5 1 , nms 1044, 1100, rmnh 2615i, 2651b-2651h, 2683, 3087-3089, 3099, 3106, 3107, 3109, 31 10, 31 13- 3115, 3177-3179, 3188, 3229, 4644, 6684, cat. ost. a, cat. ost. i-cat. ost. 1 (4), cat. ost. n, zmb 43867; Karimun (1), Pongka, Kampung— usnm 122848; Kundur(2), no locality-ZRC 4-095, 4-096; Lombok (2), Suranadi— pri 1400, no locality— bm(nh) 1 920. 1 .26.3; Matasiri(l), east coast— usnm 154368;Mwrsfl/tf(l)-ushJMll4561;Mas(5),Sia- ba, Teluk-usNM 121872-121874; no locality - zmb 48417, 48419; Padang{\), north coast— usnm 143583; Sumatra (87), Balbalan-ZMB 34005; Ba- tangkuis— zsbs Coll. Nos. 110, 114; Bengkulu— rmnh 4648; Bukittinggi— zmb 34241; Indragiri district— NHMBa 3774-3776, 3778-3781, zmuz 11668; Kateman, Sungai— usnm 123147; "Kg. Baru"— zsbs Coll. Nos. 62 (adult male), 113; La- hat— NHMBa 1805, 4829; Lampung, Propinsi— zsbs (unnumbered); Lubuksikaping— rmnh 4659; Me- dan— mcz 41 167; zsbs Coll. Nos. 28 (adult male), 67, 151; Mempura— zmb A 14.08 (subadult male), A 14.08/02 (infant); Padang-ZMUz 1 1652; Pagan- san-ZMB A 14.08 (subadult female), A 14.08/03- A 14.08/05; Pangkalan-RMNH 4645-4647; Pang- kalansusu— usnm 536025; Pap-ka— zmb A 14.08; Payakumbuh— rmnh 4649; Rawas— zmb 34025; Rokan-kanan, Sungai-ZMB A 14.08/08, A 14.08/ 1/07, A 14.08/11/01; Serangjaya-hilir-RMNH Coll. Nos. F144, F145; Siak Copatta-SMB A14.08/53; Sijunjung— rmnh w; Soekaranda— zmb A22.05; Sumatra, [east-central]— zmb A 14.08 (infant), A 14.08 (juvenile female), A 14.08 (subadult male); Sumatra, east coast— zsbs Coll. Nos. 157, 165, 169, 175; Tamiang— rmnh Coll. No. F143; Tanjung- morawa— rmnh q, z, unnumbered skeleton, zsbs Coll. Nos. 71 (adult male), 73 (subadult male), 75 (adult female); Wonosobo— rmnh Coll. No. 60; no locality-NMS 1045, 1046, 1101, 16635/IE.2.a, rmnh 1839, 1872a (female), 1872a (male), 3322, 4658, zmb Coll. Nos. 6 Guvenile), 6 (adult male), 15, unnumbered female, unnumbered male, zsbs Coll. Nos. 33, ?5 1 (adult male), 62 (subadult male), 93, 105, 109, 112, 115-117, 120, 202; Sumbawa (4), Dompu— nms 1027, 16633; Semongka— pri 1399; no locality— NHMBa 5341; Tanahmasa (1), no locality— usnm 121836 (external measure- ments recorded on skull tag); Timor (2), Bokong— zsbs 1911/2109; no locality — NHMBa 2703; Tuangku (2), north coast— usnm 1 1441 1, 1 14643 (external measurements recorded on skull tag); is- land unknown (1), Sunda Islands— NHMBa 2981. MALAYSIA, 82. MALAYSIA: SABAH, Borneo (65), Abai-Mcz 25699, 25711, 35569, 35571, 35576, 35578, 35587, 35608, 35619, 35622, 35623, 35626, 35633, 35634, 35643, 35651, 35652, 35655, 35656, 35658, 35661, 35673, 35677, 35681, 35725, 35730, 35734-35736, 35738, 35739, 35741, 35742, 35746, 35748- 35755, 35758 (cranium embedded in plaster), 35759, 35764-35767, 35768/491, 37663, 37664 (external measurements for 50 of these 5 1 speci- mens, all except mcz 35622, are listed on expe- dition record cards); Kinabalu, Mt.— mcz 37781— 37786, 37787-37789 (three mandibles only); Kin- abatangan, Sungai— usnm 19192; Sandakan, 8 mi W— fmnh 33547; Segama, Sungai— zmb A85.10; Talibang— mcz 57836; Tuaran— mcz 37407 (fe- male skull mismatched with male skin). MALA Y- SIA: SARAWAK, Borneo (12), Baram, Batang- ansp 6149, bm(nh) 1894.6.12.13; Belaga or Ka- lulong, Bukit— bm(nh) 1951.67 (male skull mis- matched with female skin), 1 95 1 .69, 1 95 1 .70; Du- lit, Bukit— zmb (unnumbered); Jumpit— zmb A 1870; Kuching-SMK Coll. Nos. 3-315, 3-320; Melinau Gorge— bm(nh), unnumbered mandibu- lar fragment; Mulu, Gunong— bm(nh) 1 894.6. 1 2. 1 ; Puram-ANSP 6113. MALAYSIA: WEST MA- LAYSIA, mainland (5), Benom, Gunong, north- east slope-BM(NH) 1979.2869; Nyalas-ZRC 4-076 (external measurements from skull tag), 4-077 (ex- ternal measurements from Weitzel et al., 1988, p. 107); Telok Anson-NHRM 3259, 3260. PHILIPPINES, 2. Jolo (1), Crater Lake Moun- tain— usnm 1 25324; Mindanao (I), Zamboanga— zmb A2920 (external measurements from Mar- tens, 1876, p. 206). 124 FIELDIANA: ZOOLOGY SINGAPORE, 1. Singapore (1), Botanical Gar- den— zrc 4-085. THAILAND, 4. Mainland (3), Ban Sakaerat- nhrm Coll. No. 1/3; Kampong Biserat— smtd B4346; Pattani— smtd B4348; Tarutao (1), no lo- cality— usnm 123992. VIETNAM, 3. Mainland (3), Cochin China- mnhn 1869.297; Ho Chi Minh City, Botanical Gardens— mnhn 1962.1445; Ho Chi Minh City, Zoological Garden— mnhn 1962.1443. Supplementary information has been derived from 1 2 unexamined specimens of M. f. fascicu- laris collected at the following localities: INDO- NESIA, Borneo: Kalimantan, Kaboerau (Gyld- enstolpe, 1920, pp. 14-15); Loa Bambam (Coll. No. 10, H. C. Raven field catalog, usnm); Long Pangian (Gyldenstolpe, 1920, pp. 13-14); Pelai- hari (Kohlbrugge, 1 896a, p. 185); Java, Kediri dis- trict— zsbs 191 1/2363, 191 1/2364 (specimens not seen, cranial measurements from G. H. Albrecht, pers. comm., Oct. 1991); Tengger, Pegunungan (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. Albrecht, pers. comm., Oct. 1991). MALAYSIA: SABAH, Borneo, Abai— mcz 35768/492 (not seen, external measurements from expedition record cards). PHILIPPINES, Mindanao, Zamboanga- 2 specimens (Martens, 1876, p. 206). Macaco fascicularis aurea, Total 55 Skins and Skulls, 44 BURMA, 33. Mainland (23), Ban Sadein— amnh 54968; Haungtharaw-zsi 1 1989; Tagoot, bnhs 5073, 5075; Taungbyauk Chaung-BM(NH) 1936.9.10.8, 1936.9.10.9, 1936.9.10.12, 1936.9.10.13; Tavoy R.-bm(nh) 1936.9.10.10, 1936.9.10.14; Tenas- serim-BM(NH) 1914.12.8.15-1914.12.8.18, bnhs 5070, 5076, 5079, fmnh 82804, zrc 4-02 1-4-023; Wimpong— bm(nh) 1885.8.1.15; Ye Forest— bm(nh) 1910.12.24.1; Kadan (1), no locality— bm(nh) 1925.7.2.1; Kathema (4), no locality— bm(nh) 1936.9.10.4-1936.9.10.7; Lanbi (1), no locality— usnm 104442; Letsok-aw (1), no locali- ty— usnm 124176 (cranium missing); Mibya (2), no locality-BM(NH) 1936.9.10.3, 1910.9.10.11; Zadetkyi (1), no locality— usnm 1 1 1898. THAILAND, 11. Mainland (1 1), Ban Tamrong Phato— ctnrc (catalog number unknown), fmnh 99641, 99642, 99644-99648; Wong, Nam Mae, 65 km E of Um Pang— amnh 54677, bm(nh) 1924.9.2.8; Wong, Nam Mae, 85 km E of Um Pang— amnh 54679. Skins Only, 8 BURMA, 6. Mainland (6), Arakan Div.— zsi 1 1990; Haungtharaw-zsi 4389, 4390, 4392 (skulls inside all three); Mergui, bnhs 5078, zsi 5238. COUNTRY UNKNOWN, 2. "Bengale"-MNHN 362/234 (lectotype), 368/247. Skulls Only, 3. BURMA, 3.