mmmMmMmm'ifmp FIELD \i ± K Zoology II NO. 96 r-t-i Systematic Review of the Rhesus Macaque, Macaca mulatta (Zimmermann, 1780) Jack Fooden C3 O Jane 30^ 2000 Publication 1S09 PUBL1SHL;D by field MDSI UN4 of N \ 1 URAL Hi- lnii,im:a.ws\ for ( oiitrlhufors to FieJdiana us should be suhiniUcd (one <>:\ , y plus two > will be considered for public, > ;l>mined (o ' lent i fie Editor. ■./, Field Musciiii jvi vYiiij wuie If typed on an IBM-compatible computer using MS-DOS, also submit text on SVi-inch 1,4.2, or 5.0, MiiltiMp!.- n.^r,iov.,..rit.» "> i x- 1 w-.^^.- '>f "^"--^im M^ i , |"i V.',.,' '• ', ". •;u- programs or ASCI i ,„..r mn manuscript i^uiics, muliuis are requested lo submit a "lable ot Coiiients," a "List of '"" '■ 'f Tables" immediately following title page. In most cases, the text should be preceded Hiuid conclude with "Acknowledgments'" (if any) and "Literature Cited." I. should be in the metric system (periods are not u.sed after abbreviated measurements). Tlic 1 ■ ■ Mi and style ol headings should follow that of recent issues of Fie/ ^or more detailed style information, see 77ie Chicago Manual of ■ >t Chicago Press, and also recent issues of Fieldiana. References: In "Literature Cited," book and jouraal titles should be givcii ii. inii. Wl«ere abbrcManons are desirable (eg , in citation of synonymies), authors consistently should follow Botanico-Periodicum-Huntianum and 77- ' r.n,u,.„.,, I ;,..,.,„,,..,, K,. p A. Stafleu & R. S. Cowan (1976 et seq.) (botanical papers) or Serial Sources for nlished by the BioSciences Information Service. Names of botanical authors should i>iaii index oi .Auiiior Abbreviations, Royal n..iini, r;,,i.!, n-, F^.v ;"v:,,'i',,: ., ■/ > nces should be typed in the following form T. B. 1978. Flora of Barro Colo- mJ. Stanford University Press, Stanford, Calif., 943 pp P J., J. R. Lloyd, and T. D. Pi n. 1963. A comparison of montane and lowland rain Ecuador. I. The fores-t structure, physiognomy, and floristics. Journal of Ecology, 51: 567-601 ON, E. J. M. 1979. Yag6 among the Siona: Cultural patterns in visions, pp. 63-80. Jn Browniau, i> I and R. A. Schwarz, eds., Spirits, Shamans, and Stars. Mouton Publishers, The Hague, Netherlands M. RRA, J 1946. The historic tribes of Ecuador, pp. 785-821. In Steward, J. H.. ed.. Handbook of South American Indians. Vol. 2, The Andean Civilizations. Bulletin ir 1 thnology, Smithsonian Institution, Washington, DC. '■ I'... '!-■•. ■■ ■ [• " .lypodiaceae. Fieldiana i. illustrations: Illustrations are referred to as "figures" in the text (not as bv s^.nie indication of scale, normaUy a reference bar. Statements in figure taptiuns alone, such as xu «, are not f aptions should be typed double-spaced and consecutively. See recent issues of Fieldiana for details of ons should be marked ' -'I, ,' hould, whenever practicable, be Wi by 11 niches (22 X 28 cm) and may not exceed llVi in). Illustrations should be mounted on boards in the arrangement to be obtained in the -anal set should I ' le for tran.smission to the printer as follows: Pen and ink drawings I'Z-ned) or photos, ,. ,. ,lcd drawings mu.st be originals, but within the .size limitation; and ' V, black and white prn s otherwise specified »{;e Proofs: i . , , jan be made aiul icd. Only ? made on the m ■e proofs. jp©r meets the requirements of ANSI/NISO Z39.48- 1992 (Permanence of Paper). FIELDIANA Zoology NEW SERIES, NO. 96 Systematic Review of the Rhesus Macaque, Macaca mulatta (Zimmermann, 1780) oo r Jack Fooden — < Division of Mammals ^^ Department of Zoology :X3 Field Museum of Natural History '^ 1400 South Lake Shore Drive Chicago. Illinois 60605-2496 U.S.A. Accepted June 10, 1999 Published June 30, 2000 Publication 1509 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 2000 Field Museum of Natural History ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents Abstract 1 Introduction 1 Geographic Distribution and Current Population Estimates 2 Pelage 7 General Characterization 7 Early Development 7 Seasonal Variation 11 Geographic Variation 16 Nepalese Standards 16 Survey of Sample Areas 16 Summary 25 External Measurements and Proportions 26 Sex and Age Variation 26 Geographic Variation 26 Head and Body Length 26 Tail Length 29 Relative Tail Length 29 Body Weight 30 Cranial Characters 38 Sex and Age Variation 38 Geographic Variation 39 Skull Length 39 Cranial and Dental Morphology 42 Comparison with Macaca fascicularis 44 Molecular Biology and Genetics 44 Mitochondrial DNA 44 Nuclear DNA 52 Blood Proteins 52 Karyology 52 Physiology and Disease 53 Blood, Cerebrospinal Fluid, and Temper- ament 53 Lead Content of Molars 53 Malaria 53 Viral Infections 54 Natural History 54 Habitats 54 Arboreality/Terrestriality 54 Swimming 56 Group Size and Composition 56 Home Range, Day Range 57 Population Density 57 Diet 57 Predators 61 Intergroup Behavior 62 Interspecific Behavior 66 Intrageneric 66 Intergeneric 68 Reproduction , 68 Seasonality 68 Sexual Maturation 70 Sexual Skin 71 Menstrual Cycle 71 Estrus 71 Consortship 72 Copulatory Behavior 72 Dominance Rank and Reproductive Suc- cess 73 Inbreeding 73 Nonreproductive Sexual Behavior 74 Gestation Length 74 Parturition 74 Birth Weight, Infant Sex Ratio 75 Birth Rate, Infant Mortality Rate 75 Nursing, Weaning 76 Menopause 77 Annual Mortality Rate 78 Population Growth Rate 78 Fossils and Subfossils 79 System ATics 79 Geographic Variation and Subspecific Recognition 79 Size 79 Tail Length 81 Pelage 81 Molecular Diversity 82 Synonymy 83 Type 86 Type Locality 86 Evolution and Dispersal 87 Acknowledgments 89 Literature Cited 90 Note Added in Proof 120 Appendix 1: Specimens Examined 121 Appendix 2: Gazetteer of Macaca MULATTA Localities 124 Index 179 List of Illustrations 1 . Known locality records of Macaca mii- latta 2 2. Detail maps of Macaca mulatta locali- ties A. Western section 4 B. Central section 8 C. Eastern section 12 3. Dorsal pelage color in Macaca mulatta A. Topotype 17 B. Contrasting pelage color in two adult males collected 4 days apart at Rajapara, India 17 m 4. Monthly incidence of prime, faded, and molting pelage stages in wild-col- lected specimens of Macaca mulatta 18 5. Sample areas cited in Macaca mulatta pelage comparisons 20 6. Latitudinal variation in interscapular hair length in Macaca mulatta adults .... 27 7. Latitudinal variation in midtail hair length in Macaca mulatta adults 28 8. Latitudinal variation in head and body length in Macaca mulatta adult non- captives 32 9. Latitudinal variation in tail length in Macaca mulatta adult noncaptives 34 10. Longitudinal variation in tail length in Macaca mulatta adult noncaptives 35 1 1 . Latitudinal variation in relative tail length in Macaca mulatta adult non- captives 36 12. Longitudinal variation in relative tail length in Macaca mulatta adult non- captives 38 13. Latitudinal variation in body weight in Macaca mulatta adult noncaptives 41 14. Skull of adult female Macaca mulat- ta— FMNH 99668, Thailand: Ban Mae Lamao 42 15. Skull of adult male Macaca mulatta — FMNH 99669, Thailand: Huai Ap Nang 43 16. Dental emergence chronology in Maca- ca mulatta 45 17. Latitudinal variation in greatest length of skull in Macaca mulatta adult non- captives 47 18. Longitudinal variation in greatest length of skull in Macaca mulatta adult noncaptives 48 19. Sample areas cited in mitochondrial DNA study of Zhang and Shi 50 20. Consensus dendrogram of mitochondri- al DNA relationships in Macaca mulat- ta samples studied by Zhang and Shi .... 51 21. Type localities of nominal species or subspecies allocated to Macaca mulat- ta; known limits of natural distribution of Macaca w»/arto also indicated 82 22. Hypothetical stages in the evolution and dispersal of A/acaca mii/arra 88 List of Tables 1 . Estimated population of Macaca mulat- ta in five countries for which data are available 3 2. External measurements and proportions in age/sex classes of wild-collected Macaca mulatta 30 3. Geographic variation in head and body length in adult Macaca mulatta 31 4. Geographic variation in tail length in adult Macaca mulatta 33 5. Geographic variation in relative tail length in adult Macaca mulatta 37 6. Dry-skin measurements of 14 Macaca mulatta specimens collected at Xing- long Xian (= Eastern Tombs), Hebei Province, northeastern China 39 7. Geographic variation of body weight in adult Macaca mulatta 40 8. Cranial measurements and proportions in age/sex classes of wild-collected Macaca mulatta 44 9. Geographic variation in greatest length of skull in adult Macaca mulatta 46 10. Frequencies of major alleles at poly- morphic blood-protein loci in samples of Macaca w«/arra from six countries .... 53 11. Habitats reported for Macaca mulatta ... 55 12. Frequency distribution of elevation rec- ords of Macaca mulatta 56 13. Arboreal/terrestrial behavior recorded during daylight hours in samples of Macaca mulatta 56 14. Group size reported for Macaca mulat- ta 58 15. Ratio of sexually mature males to sex- ually mature females reported in groups of Macaca mulatta 60 16. Home range area reported for groups of Macaca mulatta 61 17. Day range reported for groups of Ma- caca mulatta 62 18. Population density reported for Macaca mulatta 63 19. Foods reported eaten by Macaca mu- latta. Dietary proportions are indicated where data are available 64 20. Daily waking-hour time budget esti- mates for Macaca mulatta 66 21. Intergroup contact behavior reported for Macaca mulatta 67 22. Mating and birth periods reported for natural populations of Macaca mulatta 69 23. Age of sexual maturity reported for natural populations of Macaca mulatta 70 24. Birth weight in laboratory-housed Ma- caca mulatta 75 25. Infant sex ratio in Macaca mulatta 76 26. Annual birth rate in natural populations of Macaca mulatta 77 27. Infant mortality rate in natural popula- tions of Macaca mulatta 78 28. Population growth rate in Macaca mu- latta 80 29. Localities and ages of Macaca mulatta fossils or subfossils 81 30. Subspecies recognized in published classifications of Macaca mulatta, 1932-95 83 31. Latitudinal range and relative tail length in fascicularis-groxvp species of macaques 89 Systematic Review of the Rhesus Macaque, Macaca mulatta (Zimmermann, 1780) Jack Fooden Abstract The rhesus macaque, Macaca mulatta (Zimmermann, 1780), is systematically reviewed, based on examination of 638 museum specimens, observation of natural populations, and sur- vey of relevant literature. The natural distribution of M. mulatta extends from eastern Afghan- istan and western India to eastern China and northern Vietnam. This review includes analyses of geographic variation in pelage characters, external measurements and proportions, cranial characters, molecular biology and genetics, and physiology and disease. Information also is presented concerning natural history, reproduction, and paleontology. Taxonomically, local and regional populations are regarded as inadequately differentiated to warrant recognition of sub- species in M. mulatta. A hypothesis is proposed to explain the evolution and dispersal of this species. In an appendix, an annotated gazetteer lists 1,239 localities where M. mulatta has been collected or observed. Introduction bnhs The rhesus macaque, Macaca mulatta (Zim- ctnrc mermann, 1780), probably is the most intensively studied species of non-human primate (see, e.g.. Primate Information Center, 1998, p. 27). The fcxm present systematic review of M. mulatta is based on examination of 638 museum specimens (Ap- fdcg pendix 1); observation of natural populations in India, Thailand, and China; and survey of relevant fmnh literature. The principal subjects covered in this review are geographic variation in characters; nat- fubd ural history, reproduction, and paleontology; tax- onomy; and evolution and dispersal. Specimens hubd examined are preserved in the following institu- tions (number of specimens in parentheses), jggj^ which hereafter are cited by means of the indi- cated abbreviations: ^^^^ AMNH American Museum of Natural History, New York (53) izcas ANSP Academy of Natural Sciences, Philadel- phia (8) KIZ BMNH Beijing Museum of Natural History, Beijing (7) mcz bm(nh) British Museum (Natural History), Lon- don (122) Bombay Natural History Society, Mum- bai (39) Centre for Thai National Reference Col- lections, Thailand Institute of Scientific and Technological Research, Bangkok ( 1 ) Forestry College of Vietnam, Xuan Mai (5) Forestry Designing Centre of Guangxi, Nanning (8) Field Museum of Natural History, Chi- cago (23) Fudan University, Biology Department, Shanghai (2) Hangzhou University, Biology Depart- ment, Hangzhou (1) Institute of Ecology and Biological Re- sources, Hanoi (38) Institut Royal des Sciences Naturelles de Belgique, Brussels (3) Institute of Zoology, Chinese Academy of Sciences, Beijing (47) Kunming Institute of Zoology, Chinese Academy of Sciences, Kunming (28) Museum of Comparative Zoology, Har- vard University, Cambridge, Massachu- setts (8) FIELDIANA: ZOOLOGY, N.S., NO. 96, JUNE 30, 2000, PP. 1-180 Fig. 1 . Known locality records of Macaca mulatta; specimens examined include living monkeys personally ob- served in the field. For details, see Figures 2A-C. MZB MNHN NHMB NWPIB P-CM RMNH SCIEA .SIZ SMNH UPS USNM ZMB Museum Zoologicum Bogoriense, Bo- gor (1) Museum National d'Histoire Naturelle (Mammiferes), Paiis (15) Naturhistorisches Museum, Basel (2) Northwest Plateau Institute of Biology, Chinese Academy of Sciences, Xining (6) Powell-Cotton Museum, Birchington, Kent (2) National Museum of Natural History, Leiden (2) South China Institute of Endangered An- imals, Guangzhou (23) Shaanxi Institute of Zoology. Xi'an (4) Shanghai Museum of Natural History, Shanghai (54) University of Puget Sound, Tacoma (1) National Museum of Natural History, Washington, D.C. (39) Zoologisches Museum des Humboldt- Universitat, Berlin (6) ZMNH Zhejiang Museum of Natural History, Hangzhou (2) ZMVNU Zoological Museum, Vietnam National University, Hanoi (47) ZRC Zoological Reference Collection, De- partment of Zoology, National Univer- sity of Singapore (6) zsBS Zoologisches Sanrmilung des Bayerisch- en Staates, Munich (1) zsi Zoological Survey of India, National Zoological Collection, Calcutta (25) — Private collections (9) Geographic Distribution and Current Population Estimates Macaca mulatta inhabits parts of 1 1 countries in southern and southeastern Asia (Figs. 1, 2, 21), from ca. 15°N (in India, Thailand, Laos, and Viet- nam) to ca. 36°N (in Afghanistan, Pakistan, India, FIELDL\NA: ZOOLOGY Table 1. Estimated population of Macaca inulatta in five countries for which data are available.' Country Population estimate (X 1,000) References Afghanistan >4 Bangladesh 190-276 China 205-218 India >500 Vietnam 20 Total >919 Puget. 1971. p. 199 Giltins & Akonda, 1982. p. 277; Feeroz et al.. 1995. p. 75 Wang & Jiang. 1995. p. 9 Southwick & Siddiqi. 1995. p. 19 Dang. 1983. p. 1284 Cf. MacKinnon & MacKinnon. 1987. p. 189. and China) and from ca. 70°E (in Afghanistan and India) to ca. 120°E (in China). An isolated pop- ulation that formerly occurred at 40°24'N in northeastern China apparently was extirpated in 1987 (Zhang et al., 1989, p. 380). The natural range of Af. mulatta is known to include 12 shal- low-water islands — one off the coast of south- eastern Bangladesh (Maishkhal), four off the coast of northeastern Vietnam (Cat Ba, Quan Lan, Van Canh, and Van Hai), and seven off the coast of southeastern China (Hainan, Dahao, Dangan, Erzhou, Xianggang [= Hong Kong], Neilingding, and Shangchuan; the last six of these islands are near the mouth of the Zhujiang [= Pearl River]). Archaeological evidence suggests that ca. 4000 B.P. the range of M. mulatta may have extended as far west as Moenjo Daro, Pakistan (27°19'N, 68°07'E) (Mackay, 1931, p. 349; Iyer, 1977, p. 15). Although the northern limit of natural distri- bution of M. mulatta apparently is determined pri- marily by physiographic or climatological factors, the southern limit apparently is determined pri- marily by interspecific competition. The north- western limit of distribution is defined by the Great Indian Desert, the north-central limit by the Himalayas and Xizang-Qinghai (Tibetan) Plateau, and the northeastern limit in China by the transi- tion from mesothermal to microthermal climate (Trewartha, 1978, p. 9). Contrastingly, the south- western limit probably is related to competition with neighboring M. radiata (Fooden et al., 1981, p. 464), and the southeastern limit probably is re- lated to competition with neighboring M. fasci- cularis (Fooden, 1997, p. 226). The actual distri- bution of M. mulatta is of course much more re- stricted than indicated above, primarily as a result of human activity (cf. McNeely, 1992. p. 374). Two rejected locality records — one in Rajas- than, India, and the other in Xizang (= Tibet), China — are outside the limits of distribution of M. mulatta, indicated in Figures 1 and 21. In western Rajasthan, Bhargava (1982, p. 7) cited a postal survey report of M. mulatta at Babuwali, Jasalmer District (26°47'N, 69°44'E); subsequently this re- port was found to be spurious (Bhargava, 1984, p. 43). In north-central Xizang, south of a pass through the "Dupleix" mountain chain (ca. 33°38'N, 89°43'E), Bonvalot (1891, vol. 1, p. 210; cf. 1892, p. 218) reported a sighting of "monkeys" with external characters suggestive of M. mulatta: Today [ 1 8 January 1 890] we have seen monkeys crossing the frozen river and playing on the rocks which form its banks. But we cannot kill one of these animals, which are very short with red hair, small head, and an almost imperceptible tail. According to Professor Feng Zuojian, IZCAS (pers. comm., 27 September 1985), who is an au- thority on Tibetan mammals, the occurrence of monkeys in this part of the Tibetan Plateau is highly improbable. Professor Feng suspects that Bonvalot actually observed a small group of mar- mots, presumably Marmota himalayana (Zhang et al., 1997, p. 170), which he misidentified as mon- keys. Although populations of M. mulatta now in- habit the Mumbai (= Bombay) region of India, this region is outside the natural range of the spe- cies; the existing populations in this region (Raj Bhavan compound, ca. 18°56'N, 72°55'E; Borivli National Park, ca. 19°10'N, 72°55'E) are the result of artificial introductions that occurred during Worid War II (Serrao & Amladi, 1979, p. 30). The populations that now inhabit the vicinity of Kow- loon, China (ca. 22°20'N, 1 14°10'E), probably are the result of an artificial introduction that occurred during Worid War I (Herklots, 1951, p. 83); how- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA Fig. 2A. Detail map of Macaca mulatta localities, western section; for documentation, see Gazetteer. Appendix 2. Abbreviations in parentheses are those used in gazetteer locality codes; specimens examined include livmg monkeys personally observed in the field. FIELDIANA: ZOOLOGY Afghanistan (A) 27. 1 . Asmar, east of; Asmar, northwest of: Barikowt, south- east of; Kamu Valley; Kouchlaus, south of; Landay Sind, left bank, near confluence with Kunar R.; Lan- day Sind. left bank, near Pule Saret; Landay Sind, 28. near Chascoup; Landay Sind, near Merdech; Landay 29. Sind, right bank, between Mandagal and Omiul; Lan- 30. day Sind. right bank, near Sang e Safed; Landay Sind 31. Valley, near Kamu Valley; Landay Sind Valley, 32. southern slope; Mandagal, northwest of; Nurestan, eastern. 33. 2. Wama. north of. 34. 3. Gusalek, north of. 35. 4. Nurestan vicinity. 36. 5. Alicheng River, east of. 37. 6. Alingar, east of. 7. Outapour, south of. 38. 8. Chigha Sarai, north of. 39. 9. Kotgay (= Cotgai), northeast of. 40. 10. Kotgay (= Cotgai), east of. 41. 11. Chamkani, northeast of. 12. Chamkani, southeast of. 42. China (C) 43. 1. Gyirong Subcounty. 44. India (I) 45. Lolab; Lolab Valley. 46. Ovra Sanctuary, proposed. 47. Kotihar India. 48. Ramban Township vicinity. 49. Akhnoor; Chowkichora; Jammu; Nandini Wildlife 50. Sanctuary; Narota-Bun, highway between. 51. Jammu & Kashmir, southern border with Pakistan. 52. Surinsar. Kathua vicinity. 53. Ramnagar. 54. Dumel; Jhajjarkotli; Udhampur. 55. Dunwein. 56. Dharmsala; Kangra; Kangra Fort; Samayala. 57. Manali. 58. Jagatsukh. 59. Kasol; Kulu District; Kulu valley; Pulga. 60. Sainj; Sarahan; Tirthan. 61. 17. Narkanda, ca. 1 km north of; Narkanda, ca. 4 km 62. south of; Narkanda, ca. 5 km north of; Rampur, 63. northwest of; Sungri, ca. 2 km south of; Sungri, ca. 64. 4 km north of; Sungri, ca. 6 km northwest of; Sun- 65. gri, ca. 10 km northwest of; Sungri, ca. 10 km 66. southwest of; Sungri, ca. 15 km southwest of; Sun- 67. gri, ca. 20 km south-southwest of; Sungri, north of. 68. 18. Boileauganj; Cecil: Chail Sanctuary; Jakko Hill; 69. Kasauli; Kufri, ca. 0.5 km southeast of; Kufri, ca. 1.5 km southeast of; Kufri, ca. 3 km southeast of; NheraTTara Devi: Simla, western suburb; Simla vi- cinity; Simla Water Catchment Reserve; Solon Dis- trict; View, Simla vicinity. 70 Chandigarh, outskirts of. 71 Ambala District. 72 Patiala District. 73 Saraswati Forests. 74 Kamal District. 75 Kurukshetra District; Veer Sontri forests. Saharanpur vicinity. 76 1. 2. 3. 4. 5. 6. 7. 8. 9. 10 II 12 13 14. 15. 16. 26. West Timli. Aglar River; Asarori Forest; Dehra Dun; Dehra Dun vicinity; Dehra Dun vicinity, 600 m; Dhaulkot Forest; Hardwar: Kansrao; Mohan; Mussoorie vi- cinity; Rajaji Wildlife Sanctuary. Tehri-Garhwal District. Kedarnath Sanctuary. Molta. Bageshwar. Haldwani: Hanumangarhi Hill; Kumaun Hills; Naini Tal; Ratighat Dela; Jhirna; Ramganga River; Ramnagar. Bijnor; Sita Bani. Meerut District. Sonepat District. Kheri Sahd; Panipat-Rhotak. highway between; Rhohtak District. Jind District. Hissar District. Bhiwani District. Delhi; Delhi vicinity: Delhi-Mathura Road; Ghazia- bad District; Lai Kuan; Meetha Pur; Tughlaqabad. Rewari-Patudi-Gurgaon, highway between. Delhi-Agra. Khair Delhi-Hathras. Bulandshahr District; Delhi-Aligarh. Rampur-Ghaziabad. Bareilly-Agra, highway between. Makhena. Moradabad vicinity. Pilibhit-Tanakpur. Bareilly; Kareilly, west of; Karghena, west of; Ramganga South Station. Shahjahanpur-Bareilly, highway between. Sitapur-Bareilly. Haripur. Nishangara; Nishangara vicinity. Shahjahanpur-Lucknow. Lucknow-Sitapur. Gonda vicinity. Ayodhya; Faizabad-Ajodhya; Faizabad vicinity. Balrampur forest. Ayodhya-Gorakpur. Azamgarh vicinity. Sultanpur vicinity. Lucknow-Faizabad. Halwapura: Kakori: Lucknow vicinity. Sitapur-Shahjahanpur, highway between. Delhi-Kanpur, highway between. Achal Tank: Aligarh; Aligarh vicinity: Baj Garhi Bridge: Barauli Bridge: Barotha; Chaunpur; Chha- tari-do-Raha; Harduaganj; Hathras; Jawan; Nanau; Qasimpur Canal: Sasni; Satha: Sindholi: Sumera; Sumera Fall Jungle. Agra. Mathura; Vrindavan. Bharatpur. Alwar District Bandipul; Sariska Tiger Reserve. Ambagarh Reserve Forest; Amer; Barri Chopal; Galta; Jaipur. Umri Devi. (continued on following page) FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 77. 78. 79. 80. 81. 82. 83. 84. 85. 86. 87. 88. 89. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 1 17, 118. 119. 120, 121. 122. 123. 124. 125. 126. 127, 128, 129, 130 131 132 Marot. Bikaner. Sheo. Jodhpur. Kota. Sawai Madhopur. Jhansi. Banda vicinity. Chitrakut, Jagvedi and Bara Math Temples. Saklesgarh. Varanasi. Chakia Forest Range. Karkatgarh. Shahabad District. Sahebgunj. Palamau. Bandhavgarh National Park. Kakara. Sohagpur. Bhopal, east of. Tal Vraksh. Kherwada Forest. Ghori Hill: Hadya; Mahal, 2-5 km northwest of; Sadard Devi, 1 km west of; Vasunia. 3 km west of. Ajanta Caves. Bhim Kund Point; Punch Bol. Nagpur. Malua. Deogarh. Luia. Harchandi Sahai; Puri. Berhampur. Gudari. Balimila vicinity. Malkangiri. Orcha, ca. 1 km northwest of. Gurjal. Gundi. Kausa Gutta; Khanapur, 3-5 km west of; Nirmal, 16 km east of. Ankapur; Balkonda. Ali Sagar; Rudrur Agricultural Station. Hanmajipet; Ibrahimpet; Komlancha; Konapur; Magi; Mallur; Narva; Waddepalle. Mustapur, 0.5 km east of. Kondegattu temple. Bussapuram; Dumpallagudem; Govindaraopeta; Jakaram, 4 km southwest of; Laxmidevipeta; Pa- lampeta. Warangal. Gangupahad; Narayanapur; Raghunathapalle. Mulug; Pamulaparthi; Tunki; Wargel. Hyderabad. Raigir; Yadagiri Gutta. Velkicharla. Charkonda. Munipamula. Siddeldar Hill. Dachepalle. Jaggayyapet; Jaggayyapet, 4 km north of. Ballapet; Khammam; Mudigonda, Pallepadu; Tal- lada; Tanikella; Wira, 1 km south of. 133. Ashoknagar; Pakhal Lake, west side. 134. Yellandu. 135. Kothagudem; Kothagudem, 4 km north of. 136. Muttagudem. 137. Makkimarigudem; Tirumaladevipeta; Venketeswa- ra Swami Temple. 138. Dommeru; Rajahmundry, 3 km northeast of; Ra- jahmundry, 13 km northeast of. 139. Dharmajigudem. 140. Hanuman Junction; Mudhalaparava; Yeppuru. 141. Gokavaram. 142. Velatur 143. Kuchipudi; Tenali, 5.5 km west of. 144. Kondapalle; Vijayawada. 145. Anapalam; Kotanemalipuri; Narasaraopet, 2 km northwest of; Sattenapalle. 146. Darsi. 147. Angaluru; Kondra Mutla; Mellavagu; Vinukonda. Supplementary Indian localities (received too late for inclusion in map; listed in Gazetteer): Dudhwa National Park; Kanha National Park. Nepal (N) 1. Chaur. 2. Barmdeo Mandi: Sukla Phanta. 3. Bilauri. 4. Dhangarhi. 5. Aum River, ca. 20 km above mouth; Babai River vicinity; Karnali River, ca. 30 km above mouth; Kamali River/Aurn River, ca. 10 km above conflu- ence. 6. Hutu Forest; Pina, forests above. 7. Bheri River 8. Dudurhani; Simri, Narayani River. 9. Chengli. 10. Hitaura. 11. Trisuli Bazar, 4 mi (= 6.5 km) southeast of. 12. Bouzini; Gaushalla; Gokama; Katmandu; Nagarkot; Pashupati; Swayambhunath. 13. Chandikhola; Hazaria Patherghatta; Russian Camp; Simri, Birganj Forest District; Singaul. Supplementary Nepalese localities (received too late for inclusion in map; listed in Gazetteer): Balthali. Ka- vre; GhodaGhodi Tal; Pokhara; Ramnagar; Sankhu- Bajrajogini; Tripureswor, Thapathali. Pakistan (P) 1. Kanti vicinity; Kaotai; Kunar River; Mirkhani; Shi- shi Koh vicinity; Utzun vicinity. 2. Dokdusra; Gwaldri Valley: Landrai Valley. 3. Swat Kohistan region. 4. Khyber Pass vicinity. 5. Swat River. 6. Bar Chanrai Hill. 7. Pajja Hill. 8. Hazara District, southern. 9. Neelum Valley. 10. Paia; Paras vicinity; Shogran vicinity. 11. Margalla Hills. 12. Dunga Gali vicinity; Ghora Dhaka, 1 mi (= 1.6 km) east of; Kazinag; Machayara Game Reserve; Mur- ree, outskirts; Nathia Gali: Patriata; Phala/Kutbor Game Reserve. FIELDIANA: ZOOLOGY ever, this area probably had previously been in- habited by a natural population of M. mulatto. Population estimates are available for M. mu- latta in Afghanistan, Bangladesh, China, India, and Vietnam — five of the 1 1 countries inhabited by this species (Table 1). In these countries, the total estimated population is >9 19,000. Although comparable estimates are not available for Bhu- tan, Laos, Myanmar, Nepal, Pakistan, and Thai- land, the combined area of M. mulatta habitat in these six countries is less than in the preceding five (Fig. 1), which probably indicates that the total living population of M. mulatta is less than two million. According to reports published in 1995, the population of M. mulatta recently has been increasing in India (Southwick & M. F. Sid- diqi, 1995. p. 18) and has been decreasing in Chi- na (Jiang Haisheng et al., 1995, p. 178). Pelage General Characterization Dorsal pelage coloration is a key character for species identification of M. mulatta. In M. mulat- ta, the fur of the lower back is conspicuously more erythristic than that of the upper back (Fig. 3). In specimens in prime pelage (see "Seasonal Variation," below), the color of the upper back varies from yellowish gray to golden brown to burnt orange, and the color of the lower back varies correspondingly from golden brown to burnt orange to intense burnt orange ("almost fi- ery red"; Pocock, 1932, p. 531; cf. Sikorska-Pi- wowska, 1959, p. 272). On the upper back, the proximal two-thirds of individual dorsal hairs is gray, and the distal one-third is annulated with alternating pale and dark bands, yellowish to golden and blackish (cf. Koppikar & Sabnis, 1976, p. 7); on the lower back, the grayish color of the proximal two-thirds of individual hairs is paler, the pale distal annulations are more ery- thristic, and the dark distal annulations are more dilute. The crown, nape, and sides of the head are approximately the same color as the adjacent up- per back; the anterior edge of the crown is marked by a blackish superciliary streak, and the cheeks often are also fringed with blackish hairs. Crown hairs usually are smoothly directed posteriorly. Hairs on the side of the head usually form a small crest or whorl near the angle of the jaw (infra- zygomatic crest; Fooden, 1995, p. 19); occasion- ally, this crest is elongated and extends upward between the eye and the ear as far as the side of the crown (transzygomatic crest; 44 of 240 spec- imens examined). The thinly haired facial skin is buffy to reddish, except for the upper eyelids, which are whitish (unpigmented). On the proxi- mal part of the limbs, dorsal pelage is similar in color to that on the adjacent trunk; more distally, the pelage color of the limbs becomes less ery- thristic and more dilute. The basal one-fourth of the tail is approximately the same color as the adjacent lower back; the distal three-fourths is bi- color, dark brown dorsally and buffy ventrally. The ventral surface of the trunk and limbs is thin- ly haired, pale buffy to whitish. Broadly distrib- uted areas of sexual skin undergo cyclical changes in color and swelling (see "Reproduction." be- low); in adult males, the glans is blue-black (Wil- son & Vessey, 1968, p. 5). The pelage in M. mulatta populations at upper elevations (ca. 2400 m) in India apparently is lon- ger and sleeker than in nearby populations at low- er elevations (Dodsworth, 1914, p. 730). In cap- tivity, the pelage of dominant individuals, partic- ularly high-ranking males, reportedly is sleeker than that of subordinate individuals (Chance, 1956, p. 5; Waterhouse & Waterhouse, 1971, p. 19). The large amount of grooming received by dominant individuals may contribute to their sleekness. Abnormally pale "golden" M. mulatta individ- uals reportedly occur in various parts of India and Pakistan at an estimated frequency of 1/10.000 (Pickering & van Wagenen. 1969. p. 161; cf. Anonymous, 1978, p. 12; Kessler et al.. 1986, p. 264). Dorsal pelage color in these aberrant indi- viduals is pale yellowish anteriorly and pale red- dish-golden posteriorly; the skin is virtually un- pigmented, and retinal pigmentation also is re- duced. Experimental breeding indicates that the "golden" condition probably is inherited as an autosomal recessive trait. Among specimens ex- amined, a fluid-preserved late fetus or neonate with umbilical cord still attached (bm(nh) 1972.836), collected at Pyaunggaung, Myanmar, 14 May 1915, appears to be of the "golden" phe- notype. Early Development Pelage color in M. mulatta neonates is pale brown to dark brown, generally somewhat darker than in adults (Tinklepaugh & Hartman, 1932, p. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA Fig. 2B. Detail map of Macaca mulatta localities, central section: for documentation, see Gazetteer, Appendix 2. Abbreviations in parentheses are those used in gazetteer locality codes; specimens examined include living monkeys personally observed in the field. FIELDIANA: ZOOLOGY <— Bangladesh (Ba) 17. Yushu Xian. 1. Dinajpur District. 18. Jegu Xiang; Maluling. 2. Rangpur District. 19. Dainkog. 3. Nawabganj vicinity. 20. Routoumdo. 4. Naogaon. 21. Qamdo. 5. Madhupur, ca. 100 km west of. 22. Dege vicinity. 6. Garo Hills, foot. 23. Baima. 7. Madhupur National Park; Rasulpur vicinity. 24. Jigzhi. 8. Madhupur National Park, southern portion. 25. Zoige Xian. 9. Mymensingh, northern. 26. Danba. 10. Kalabokhani. 27. Kangding; Moshemien. 1 1 . Patharia. 28. Olongche. 12. Fechugang. 29. Yajiang. 13. Harargaj; Rajkandi; Rema-Kalenga; Srimangal Tea 30. Xi Golog. Estate; Srimangal vicinity; Sylhet Forest Division; 31. Litang-Batang. Tarap; West Bhanugach. 32. Batang vicinity; Kiang-ka. 14. Raghunandan. 33. Dzo La, southeast of. 15. Barmi; Borme. 34. Zayii Xian. 16. Dhamrai. 35. Yanjing vicinity. 17. Meherpur 36. Deqen. 18. Chuadanga. 37. Xiangcheng. 19. Jessore District. 38. Zhongdian. 20. Satkhira. 39. Muli. 21. Sundarbans, ca. 50 mi (= 80 km) east of Calcutta. 40. Yanyuan. 22. Satkhira, southern. 41. Weixi. 23. Sundarbans. 42. Fugong. 24. Wazipur 43. Biloxue Shan. 25. Char mugoria; Madaripur Township. 44. Ashi. 26. Faridpur District. 45. Lijiang. 27. Bondor; Dhaka; Narayanganj; Sonargaon. 46. Yong.sheng [Xian]. 28. Matlab Bazar; Chandpur Bazar, Old; Puran Bazar. 47. Huaping [Xian]. 29. Comilla District. 48. Miyi. 30. Matlab, southeast of. 49. Yuanmou. 31. Noakhali. 50. Luquan. 32. Sitakunda. 51. Shuangbai. 33. Chittagong, northern. 52. Chuxiong. 34. Chittagong Hill Tracts, northern; Pablakhali. 53. Nanhua. 35. Chittagong, eastern; Hazarikhil; Kaptai; Kaptai 54. Nanjian. Lake; Satghar; Sitapahar/Rampahar 55. Yunlong [Xian]. 36. Chittagong Hill Tracts, eastern. 56. Baoshan. 37. Cox's Bazar, northern; Sangu/Malamuhari. 57. Datang. 38. Chittagong, southern; Chokoria Sunderbans; Chun- 58. Hui-yao. ati Wildlife Sanctuary; Cox's Bazar; Himchari; Kap- 59. Tengchong (= Momien); Tengchong Xian. tai, south of; Maiskhal Island; Padua. 60. Yingjiang. 39. Ukhia. 61. Hotha Valley. 40. Teknaf Peninsula. 62. Santaishan. Bhutan (Bh) 63. Cala Shan. 1. Royal Manas National Park. 64. Changning [Xian]. China (C) 65. Nu Jiang, above Changlung. 1. Yadong Xian. 66. Minglang; Yongde vicinity. 2. Maizhokunggar Xian. 67. Gengma. 3. Qusum Xian. 68. Nanding He. 4. Lhiinze Xian. 69. Lincang Prefecture. 5. Tsari Chu. 70. Lancang Jiang. 6. Nang Xian. 71. Jingdong Xian. 7. Gyaca Xian. 72. Ailao Shan (= Mountain) Reserve. 8. Gongbo'gyamda Xian. 73. Wuliang Shan (= Mountain) Reserve; Zhenyuar 1 Xian. 9. Dowoka. 74. Jinggu Xian. 10. Kongbo. 75. Mojiang Xian. 11. Yigong; Yigong Forest Reserve. 76. Liichun. 12. Gyala, above. 77. Pu'er Xian. 13. Medog Xian. 78. Simao Xian. 14. Ngamda. 79. Lancang Xian. 15. Baizha Plantation. 80. Meng-ban. 16. Makehe Plantation. (continued on following page) FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 81. Menghai; Menghun; Shanman. 82. Menglong. 83. Mengyang. 84. Mcnglun. 85. Manpa. 86. Nonglin. India (I) 1. Harinbhanga. 2. Jhilla. 3. Calcutta, Hastings Road; Calcutta. Indian Museum Compound: Calcutta, northern. 4. Basirhat Reserve Forest. 5. Bhagalpur District. 6. Mangpu; Narbong; Panighatta; Siliguri; Simulbari- Pankhabari; Sivok; Sivok, ca. 3 km east of; Sivok, ca. 5 km east of; Sivok, ca. 6 km east of; Sukna; Sukna-Kurseong. 7. Sikkim; Tarkhola. 8. Gorubathan forest. 9. Bharnabhari; Hasimara; Jaldapara Wildlife Sanctu- ary. 10. Jamduar vicinity; Maure, near; Raimona vicinity. 1 1 . West Garo Hills District. 12. Balphakram region. 13. Garo Hills; Rongrenggiri vicinity. 14. United Khasi-Jaintia Hills. 15. Kulsi [River]; Rajapara. 16. Bogra Nadi. 17. Nongpoh. 18. Narpuh Reserved Forest. 19. Hot Springs. 20. Lamsakhang. 21. Baguri Block; Haldhibari Block. 22. Golaghat. 23. Neghereting. 24. Dafla Hills. 25. Adupuria; Akhoiphutia; Bahgara; Bezogaon; Chere- kapar, near; Chetia; Dichialgaon; Dihajan habi; Ha- tighuli; Japisojia; Jayrapar; Jhanji; Judi-Jatakia; Kath- par; Khanamukh; Lunpuriagaon; Mesogarh; Meteka Misajan; Moduri; Nanglamora; Nimaijan-Bahdhora Rajmai tea garden, east of National Highway No. 37 Rajmai tea garden, west of National Highway No. 37; Sala Reserve Forest; Saraguri; Tipomia. 26. Dangori Nadi, near; Tinsukia District (Bherjan, Bo- rajan, Podumoni R. F). 27. Yongyap Chu. 28. Dening. 29. Margherita. 30. Dhuniopathar; Dillighat; Diroi (Rangoli) Reserve Forest; Sapekhati Reserve Forest. 31. Changchang Pani. 32. Chungtia. 33. Mikir Hills. 34. Samaguting. 35. Cachar District. 36. North Cachar Hills. 37. Imphal, ca. 4 mi {= 6.5 km) north of; Imphal, Ma- habali temple. 38. Bishenpur 39. Gharmur, ca. 1 km south of; Nagorhgena; North District, northeastern; Paach piror mukam. 40. Ampi Bazar, ca. 3 km southeast of resthouse; Char- ilam resthouse; Gumti Sanctuary; North District, north-central; North District, northwestern; North District, western; South District, north-central; South District, south-central; South District, south- eastern; South District, southwestern; South District, west-central; West District, east-central; West Dis- trict, south-central; West District, southern; West District, southwestern; West District, western. 41. North District, ea,stern. 42. North District, .southeastern; North District, south- western; South District, northeastern. 43. Phawngpui Wildlife Sanctuary. Supplementary Indian locality (received too late for inclusion in map; listed in Gazetteer): Manas National Park. Loos (L) 1. Ou, Nam, between Muang Khoua and Muang Ngoy. 2. Ou, Nam (= Nam hou). 3. Louangphrabang. 4. Xaignabouri. 5. Mekong River, 90 km above Viangchan. 6. Ban Kuai, several km south of; Ban Napo vicinity; Ban So vicinity; Ban Wangma vicinity. 7. Viangchan. 8. Ban Mak Nao. Myanma r (M) 1. Bawmwang; Htingnan Triangle. 2. Hkandau. 3. N'Changyang. 4. Tanga-Shingaw, road between. 5. Karen Chaung; Myitkyina; Tang Hpre. 6. Nanyaseik. 7. Taga Hka; Taro. 8. Singkaling Hkamti, northern Chindwin River; Sing- kaling Hkamti, right (west) bank; Singkaling Hkam- ti, upper Chindwin River, east bank. 9. Heinsun. 10. Moklok. 11. Tamanthi Wildlife Sanctuary. 12. Hisweht. 13. Maungkan. 14. Homalin. 15. Bhamo. 16. Kindat, 20 mi (= 32 km) northwest of. 17. Ali Cha. 18. Kindat; Tatkon, near Kindat, east bank of Chindwin River; Tatkon, near Kindat, west bank of Chindwin River. 19. Mansam Falls; Se-eng. 20. Pyaunggaung. 21. Lethan Hka; Maymyo. 22. Madaya. 23. Mingun. 24. Kin; Yin, east bank of lower Chindwin River; Yin, lower Chindwin River. 25. Chittagong Hill Tracts. 26. Irrawaddy River, left bank, below Yenangyaung. 27. Popa Hill. 28. Kokkoaing. 29. Toungoo, 30 mi (= 48 km) northwest of. 30. Toungoo, 15 mi (= 24 km) north of. 10 FIELDIANA: ZOOLOGY 259; Hill, 1974, p. 565; Roonwal & Mohnot. 1977, p. 147; Kessler et al., 1986, p. 264; Higley et al., 1987, p. 9); the crown hair of neonates is parted by a midline bare area. At birth, the bare skin of the face, hands, and feet is dull purple, but within about 5 minutes it gradually becomes pale pink (Rawlins, 1979, p. 432). Skin color changes from pinkish to buffy by age ca. 2 months, and the dark neonatal pelage is gradually replaced by paler pelage, similar in color to that in adults and without a coronal part, by age ca. 6 months (Hinde et al., 1964, p. 611; Roonwal & Mohnot, 1977, p. 147; Higley et al., 1987, p. 9). In two young infant specimens examined (fmnh 82806, 82807, Yin, Myanmar. 18 June 1915), with deciduous first incisors emerging (age <7 weeks; cf. Fig. 16), the dorsal pelage is relatively thin and fine-textured — brown on the crown, pale yellow- ish brown on the upper back, pale burnt orange on the lumbosacral region, and pale ochraceous on the tail and limbs; the face is nearly naked, and the underparts are thinly covered with short pale ochraceous hairs. In two older infants (amnh 57108. Eastern Tombs, China, winter 1929; fmnh 99671, Huai Kwang Pah, Thailand, 29 March 1967), with deciduous second upper molars erupt- ing (age ca. 5 months), the pelage is essentially similar in color and texture to that in adults. How- ever, fine-textured pelage may be retained by some young juveniles, even after the permanent first molars have begun to erupt (usnm 20122, Lolab, India, 9 September 1891, age >1 year). The reaction of captive adult females to 6- month-old infants with either naturally colored buffy faces or artificially colored pink faces (sim- ilar in color to the faces of neonates) has been investigated experimentally (Higley et al., 1987, p. 16). In this study, the adult females paid more attention to pink-faced (neonate-like) infants than to buffy-faced infants. Seasonal Variation Macaca miilatta undergoes seasonal molting, both in natural populations (Pocock, 1932, p. 531; Pearl et al.. 1987, p. 38) and in captivity (Hart- man, 1931, p. 141; Stewart, 1933, p. 30; Rowell, 1963, p. 195; Vessey & Morrison, 1970, p. 90; Morrison & Menzel, 1972, p. 63; Mac Arthur et al., 1978, p. 155; Wolfe, 1985, p. 243; O'Neill- Wagner, 1997, p. 138). In natural populations, most postinfantile specimens collected during the period September-February are in prime pelage (Fig. 4; cf. Pocock, 1932. p. 531; Pearl et al., 1987, p. 38). In this condition, the fur on the dor- sal surface is richly colored, lustrous, long, soft, and smooth; the pale distal annulations on indi- vidual dorsal hairs are bright, crisp, and conspic- uous (e.g., FMNH 35448, adult male, Mangpu, West Bengal, India, 5 December 1930). A slight fading of the pelage may become evident as early as October, but this early-stage fading is most commonly seen in March; compared with prime pelage, the dorsal fur is more grayish, less lus- trous, and somewhat disheveled, and the contrast between the pale distal annulations and adjacent darker regions of individual hairs is reduced (FMNH 99668, adult female. Ban Mae Lamao, Thailand, 21 March 1967). In subsequent months fading continues, with the result that dorsal fur in most specimens collected from April through June is dull colored, weakly annulated, streaky, harsh textured, and scraggly (late-stage fading); at this 31. Toungoo. 13 mi (= 21 km) east of. 32. Toungoo, east side of Sittang River. 33. Toungoo, 20 mi (= 32 km) west of. 34. Pye (= Prome). 30 mi (= 50 km) southeast of; Pye (= Prome), 35 mi (= 55 km) southeast of. Nepal (N) 1. Sankhuwa Khola. 2. Sabaya Khola. 3. Kosi River. 4. Morang region. Supplementary Nepalese localities (received too late for inclusion in map; listed in Gazetteer): Heluwabeshi; Lakuwa; Sagarmatha (= Mount Everest) National Park; Siva (= Shiva). Thailand (T) 1. Chiang Dao. 2. Chiang Mai, near 3. Pang Nam Un. 4. Kaeng Mae Hat (rapids). 5. Huai Ap Nang; Huai Kwang Pah. 6. Ban Mac Lamao. 7. Tha Chang Tai. 8. Ban Umphang, 28 mi (= 45 km) southeast of. 9. Khao Nang Rum. western slope; Khao Nang Rum Research Station. 10. Dan Sai district. 11. Nong Khai. Vietnam (V) 1. Muong Bourn. 2. Muong Mo. 3. Muong Cha. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA II 105° 110° 115" 120° Fig. 2C. Detail map of Macaca mulatta localities, eastern section; for documentation, see Gazetteer. Appendix 2. Abbreviations in parentheses are those used in gazetteer locality codes; specimens examined include living monkeys personally observed in the field. 12 FIELDIANA: ZOOLOGY China (C) 1. Huashi. north of; Liulipenshan: Xianglong Xian, southern. 2. Yicheng. 3. Li Shan National Nature Reserve. 4. Jiyuan, ca. 30 km northwest of: Manghe Nature Reserve; Taihang Shan. 5. Yangcheng. 6. Jincheng; Shanxi, southeastern. 7. Lingchuan, southeast of. 8. Huixian. 9. Xinxiang Xian. 10. Xiuwu Xian. 1 1. Boai; Qinyang. 12. Jiyuan, ca. 20 km northeast of. 13. Jiyuan. ca. 80 km west-northwest of. 14. Yuanqu; Zhongtiao Shan. 15. Ruicheng. 16. Liangdang. 17. Huixan. 18. Baishuijiang Natural Reserve. 19. Kangxian. 20. Chengxian. 21. Wudu. 22. Nanping. 23. Qingchuan. 24. Pingwu. 25. Maowen. 26. Wenchuan. 27. Wassuland. 28. Wanglang Natural Reserve. 29. Qionglai. 30. Guanxian. 31. Tonojiang Xian. 32. Dashuping. 33. Nanjiang. 34. Chenojaba. 35. Dahe; Longchi. 36. Zhengba Xian. 37. Wanyuan. 38. Wanxian. 39. Enshi. 40. Wushan. 41. Zhenping. 42. Zhushan. 43. Shennongjia Forestry Region. 44. Huangliangping. 45. Zigui. 46. Yichang; Yichang (= Ichang), Chang Jiang ( = Yangtze) gorges above. 47. Zhaotan. 48. Guniujiang; Liukou, Qimen Xian. 49. Gegong. 50. Jiuhua Shan; Tangxi. 51. Jiuhua Shan, ca. 30 km east of. 52. Jiuhua Shan, ca. 30 km northeast of. 53. Biyun. ca. 30 km north of. 54. Yangliupu. 55. Daoshiwu; Hule; Qingliangfeng. 56. Anji Xian. 57. Laodian. 58. Xindeng. 59. Changhua vicinity. 60. Jixi. 61. Shexian; Tunxi. 62. Biyun; Dalingxia; Fuxi; Huangshan; Jilian; Rucun; Shimen; Tong Kou; Yixian. 63. Liukou, Xiuning Xian; Qihong. 64. Huangtankou; Quxian. 65. Xinluwan. 66. Julongshan Nature Reserve; Zhidaikou; Zhuxi. Zhoucun: 67. Kucun. 68. Huangqiao. 69. Zhouning Xian. 70. Pingnan Xian. 71. Jiufeng Shan. 72. Fangdao Nature Reserve; Jian'ou Xian. 73. Jianyang Xian. 74. Songxi Xian. 75. Pucheng. 76. Chong'an Xian; Kuatun; Shangang Nature Re- serve; Wuyi Shan Nature Reserve. 77. Fengxingshan. 78. Zixi Xian. 79. Guangze Xian. 80. Shaowu Xian. 81. Taining; Taining Xian. 82. Sha Xian. 83. Youxi. 84. Yongtai Xian. 85. Fuqing vicinity. 86. Daiyun Shan. 87. Yongchun. 88. Sanming Xian. 89. Yong'an. 90. Liancheng Xian. 91. Meihua Shan. 92. Longyan Xian. 93. Longxi. 94. Shanghang Xian. 95. Xingning. 96. Lianping. 97. Bachi. 98. Anyuan. 99. Shuanyuan. 100. Ningdu. 101. Chongyi. 102. Guidong. 103. Ninggang. 104. Luyuan. 105. Taihe. 106. Pinxiang vicinity. 107. Qiasui. 108. Changde. 109. Cili. 110. Suoxi Valley. 111. Sangzhi. 112. Yongshun. 113. Jishou. 114. Songlao. 115. Jiangkou. 116. Fanjingshan. 117. Youyang. 118. Yinjiang. 119. Shiqian. 120. Yuqing. 121. Getou. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 13 122. Sandu. 174. 123. Dushan. 175. 124. Huishui. 176. 125. Guiyang. 126. Weng'an. 177. 127. Kaiyang. 128. Mcitan, near. 129. Ziinyi. 178. 1 30. Suiyang. 179. 131. Gulin. 180. 132. Zheng'an. 181. 133. Nanchuan. 182. 134. Xishui. 183. 135. Tongzi. 136. Hejiang. 184. 137. Yibin. 185. 138. Tseo-Jia-Geo; Yunnan border; Yunnan border. 186. south of Yibin. 187. 139. Emei Shan; Leshan (= Kia-ting), mountains 30 mi 188. southwest of; Shihshahshu Temple. 189. 140. Gin Keo Ho, cliff above; Wa Shan. 190. 141. Meigu. 191. 142. Leibo; Yongshan. 192. 143. Yiliang. 193. 144. Shuicheng Xian vicinity. 194. 145. Qingzhen. 146. Changshun. 147. Fuyuan. 195. 148. Mile. 149. Gejiu. 196. 150. Jinping. 197. 151. Pingbian. 198. 152. Hekou. 199. 153. Guangnan. 200. 154. Maojie Bird Reserve. 201. 155. Fameng; Jinzhong Shan Bird Reserve. 202. 156. Xingyi. 203. 157. Anlong. 204. 158. Wangmo. 205. 159. Ceheng. 206. 160. Dahongbao Nature Reserve. 207. 161. Huagong Water Regulation Forest Reserve. 208. 162. Nazuo Water Regulation Forest Reserve. 209. 163. Funing. 210. 164. Defu Water Regulation Forest Reserve; Longhua Water Regulation Forest Reserve. 165. Nongxin Water Regulation Forest Reserve. 166. Dizhou Water Regulation Forest Reserve. 211. 167. Ditin; Gulong Shan Water Regulation Forest Re- 212. serve; Motianling. 213. 168. Dawanglin Water Regulation Forest Re iserve; 214. Huanglian Shan Water Regulation Forest Reserve. 215. 169. Baidonghe Water Regulation Forest Reserve; 216. Chengbihe Water Regulation Forest Reserve. 217. 170. Batu. 218. 171. Cenwanglao Shan Nature Reserve; Yuhun. 219. 172. Gao Lo Shan. 220. 173. Bamo village, near; Buliuhe Water Regulation For- 221. est Reserve; Chuan Dong; Chuandonghe Water 222. Regulation Forest Reserve; Guangli, 50 m above; 223. Hongshui He, between Tian'e and Hai Zhou; 224. Hongshui He, left bank, 9 km and 10 km below Heke; Hongshui He. right bank, 500 m below Heke; Koditan; Sanpihu Water Regulation Forest Reserve; Tian'e. Liuzhai vicinity; Lungli vicinity. Libo. Xunle vicinity; Xunle Water Regulation Forest Re- serve. Dongmen; Dongshan; Jenli, 2-3 km north of; Mu- lun Nature Reserve; Pochuan, 6-7 km west of; Xianan; Xianan-Mulun. Cioupu Shan; Hechi Prefecture; Jinchengjiang. Luocheng. Sijian Shan Water Regulation Forest Reserve. Jiuwanshan Water Regulation Forest Reserve. Yuanbao Shan Nature Reserve. Shoucheng { = Shocheng) Water Regulation Forest Reserve. Huaping Nature Reserve. Longsheng. Chengbu. Xinning Xian, Southern. Quanzhou. Mao'er Shan Nature Reserve. Yindian Shan Water Regulation Forest Reserve. Qianjiandong Water Regulation Forest Reserve. Haiyang Shan Water Regulation Forest Reserve. Qingshi Tan Water Regulation Forest Reserve. Jiaqiaolin Water Regulation Forest Reserve; Lagou Bird Reserve; Xilin Shan Water Regulation Forest Reserve. Dayao Shan Nature Reserve; Laoshan; Piangzu; Puquan Road Maintenance Station. Daping Shan Nature Reserve. Wuzhou; Xi Jiang. Nan Ling. Gupo Shan Water Regulation Forest Reserve. Wangjuanshan. Lianzhou. Yangshan. Shanmoji. Yingde. Boluo. Qigong. Huaiji. Luofu Shan. Huidong. Dahao Dao; Eagle's Nest Trail; Kam Shan En- trance; Sam Shui Wan Valley; Shing Man Country Park; Tai Po Kau Nature Reserve; Tai Tam Res- ervoir; [Victoria] Peak. Dangan Dao; Erzhou Dao. Neilingding Dao. Beichuanshan; Miwan. Tiantang Shan Water Regulation Forest Reserve. Linwan Shan Water Regulation Forest Reserve. Guixi. Shiwan Dashan Water Regulation Forest Reserve. Shangsi. Bapon. Nanning. Darning Shan Nature Reserve. Longjun (= Lingjun) Hsienmu Reserve. Daxin Rare Animal Reserve; Zhongzhou. Chunxiu Water Regulation Forest Reserve; Qing- long Shan Water Regulation Forest Reserve; Xialei Water Regulation Forest Reserve. 14 FIELDIANA: ZOOLOGY stage, the tips of individual hairs frequently are abraded (fmnh 99669, adult male, Huai Ap Nang, Thailand, 29 March 1967). From June to October, but most commonly in July and August — which is the peak of the rainy season in most parts of the range of M. mulatta — molting occurs, and the faded old fur is replaced by short bright new fur, first on the crown and tail, next on the arms and legs, and finally on the back and flanks (bm(nh) 1915.5.5.3, adult male, Homalin, Myanmar, 14 July 1914). Although the process of fading, molt- ing, and hair replacement is gradual, and the as- signment of marginal specimens to particular stages in the process is therefore somewhat arbi- trary, the general pattern is reasonably clear. Captive populations in England and in Maryland and Florida, U.S.A., reportedly molt in the spring or summer (Hartman, 1931, p. 141; Rowell, 1963, p. 195; MacArthur et al., 1978, p. 155; Wolfe, 1985, p. 243; O'Neill-Wagner, 1997, p. 138); this is similar to the timing of molting in natural pop- ulations (Fig. 4). In individual members of a cap- tive free-ranging population (n = 156-186), the duration of the molting stage varied from 4 weeks to 16 weeks and generally was longest in adult males (Vessey & Morrison, 1970, p. 90). Pocock (1932, p. 532; 1939, p. 47) suggested that the molting season in M. mulatto may vary geographically, depending on local environmental conditions. This suggestion is supported by evi- dence from two free-ranging colonies studied by Vessey and Morrison (1970, p. 92) and evidence from one translocated free-ranging group studied by Morrison and Menzel (1972, p. 63). Vessey and Morrison found that the molting period in a colony at Cayo Santiago, Puerto Rico (January- May), is about 3 months earlier than in a colony at La Parguera, Puerto Rico (April-August). The La Parguera colony inhabits a pair of relatively dry islands ca. 160 km southwest of Cayo Santi- ago, which is relatively wet; the timing of molting in the La Parguera colony is fairly close to that in natural populations of M. mulatta (Fig. 4). The group studied by Morrison and Menzel was trans- located in July 1966 from Cayo Santiago to De- secheo I., a relatively dry island ca. 60 km north- 225. Chongzuo Rare Animal Reserve; Fusui Rare Ani- 7. Ly Bon. mal Reserve; Longhu Shan Nature Reserve; Xid- 8. Ban Vay; Po Lu. aming Shan Water Regulation Forest Reserve. 9. Trung Khanh District. 226. Longan; Longrui (= Lingrui) Nature Reserve. 10. Bac Can; Linh Thong. 227. Longgang Nature Reserve; Ningming. 11. Dinh Ca. 228. Danzhou. 12. Lang Son Province. 229. Nada; Nanfeng (= Nam Fong) Shi. 13. Quan Lan. Dao; Van Canh, Dao; Van Hai. 230. Tunchang Xian. 14. Ang Co; Cat Ba Dao. 231. Wenchang Xian. 15. Thai Nguyen. 232. Pisui; Wuzhi Shan; Zhayun. 16. Kien Thiet vicinity. 233. Baisha. 17. Thuong Bang La. 234. Bawangling; Mihouling; Yiajia. 18. Thanh Son. 235. Changtian; Dongfang; Xi Shia. 19. Tin Toe Forest. 236. Jianfengling. 20. Hoa Binh. 237. Nychow (?= Yai-ch'eng) vicinity. 21. Phu Vach. 238. Sanya. 22. Hoi Xuan. 239. Nanwan; Nanwan Nature Reserve. 23. Cue Phuong. 240. Xinlong. 24. Nghia Dan. Laos (L) 25. Nghia Dung. 1. Xiangkhoang. 26. Ban Bu. 2. Muang Pakxan. 27. Huong Son (= Muong Son) District. 3. Muang Thateng. 28. Trai Tru. Thailand (T) 29. Ky Son. 1. Kumpawapi Park. 30. Bo Trach District. Vietnam (V) 31. Xuan Ninh. 1. Bac Tan Trai. 32. Vinh Linh region. 2. Muong Muon; Muong Pon. 33. Hue. 3. Muong Moun. 34. Song-Ta-Voy. 4. Nam Ngap. 35. Bach Ma National Park. 5. Cham Chu; Chiem Hoa. 36. Son Tra. Mt.; Son Tra. Mt., 3.9 km west and 0.3 km 6. Ban Bung vicinity; Ban Thi; Cho Don District; south of. Gam, Song, left bank; Tat Ke vicinity; Thanh 37. Dak Sut. Tuong. 38. Mom Ray (= Mon Ray) Nature Reserve. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 15 west of La Parguera; after a transitional molting season in 1967, the onset of the molting season in the Desecheo group apparently shifted to May (1968-1971), close to the onset at La Parguera, instead of January, when the molting season be- gins at Cayo Santiago. In natural populations, the peak of the molting season usually coincides with the end of the birth season, and most individuals are in prime pelage during the mating season (cf. Fig. 4, Table 22). A similar relationship between molting and birth seasons also prevails in the free-ranging Cayo Santiago and La Parguera populations (Vessey & Morrison, 1970, p. 92); in the Desecheo group, the molting peak apparently occurs after the birth season (Morrison & Menzel, 1972, p. 75). In the La Parguera colony, molting tended to occur ear- lier in mature males and mature nonpregnant fe- males than in immatures and pregnant females, which usually molted after parturition (cf. O'Neill-Wagner, 1997, p. 138); this suggests that molting may be controlled, at least in part, by sex hormones (cf. Dietz et al., 1995, p. 282). Anom- alously, one 4-year-old pregnant female in the La Parguera colony molted twice in the same year, once after the mating season and once after the birth season. Other species of macaques in which seasonal molting has been reported are M. fuscata, M. ra- diatch M. assamensis, and M. thibetana (Pocock, 1931, p. 276; 1939, p. 52; Hill, 1974, p. 765; Fooden, 1981, p. 9; 1982a, p. 6; 1983, p. 8; Ina- gaki & Nigi, 1988, p. 82). Molting in M. fiiscata, M. radiata, and M. assamensis apparently occurs in May or June, which probably is slightly earlier than usual in natural populations of M. mulatta. Molting in M. thibetana apparently occurs in late summer (ca. August), which is about the same as in natural populations of M. mulatta. Geographic Variation Pelage characters have served as the primary basis for recognition of subspecies in M. mulatta. For this reason, detailed analysis of geographic variation of these characters is required. As indi- cated by Pocock (1932, p. 533), the preferred standard of comparison for such an analysis would be the type specimen, which unfortunately has not been preserved. Lacking the type speci- men, specimens collected at or near the type lo- cality will be used as standards of comparison. Nepalese Standards — The type locality of M. mulatta, originally given as "India," subsequently was restricted by Pocock (1932, p. 533) to "Nepal Tarai" (now spelled "Terai"), which is the narrow plain that extends along the southern border of Nepal (Karan, 1960, p. 1). One specimen from the Nepal Terai is now available; this topotype is bm(nh) 1922.5.16.2, an adult male in prime pel- age, collected 17 February 1921 at Hazaria Path- erghatta (ca. 27°00'N, 85°15'E), 180 m (Fig. 3A). Dorsal pelage in this specimen is bright golden brown anteriorly, becoming intensely burnt or- ange on the lumbosacral region (cf. Pocock, 1932, p. 534). Interscapular hair length (ISHL) is ca. 50 mm. The tail is not particularly bushy; midtail hair length (MTHL) is ca. 25 mm. Two near topotypes, also in prime pelage, are available from higher elevations in Nepal — BM(NH) 1921.5.1.1, adult female, 15 October 1920, Nagarkot (ca. 85 km northeast of Hazaria Patherghatta), 2400 m, and bm(nh) 1931.1.11.11, late juvenile male, 7 December 1922, Chengli (ca. 130 km northwest of Hazaria Patherghatta), alti- tude unspecified. Dorsal pelage color in these two skins is slightly more erythristic than in the topo- type. Hair length (female, ISHL = 40 mm, MTHL = 10 mm; male, ISHL = 60 mm, MTHL = 30 mm) is similar to that in the topotype. Ten additional Nepalese specimens examined are less useful for pelage comparisons. These are bm(nh) 1921.5.1.2, infant female; fmnh 104164, young juvenile male (Ml, II), early-stage faded pelage; bm(nh) 1931.1.11.10, juvenile female col- lected in June, beginning to molt; four badly de- teriorated skins collected before 1846 (bm(nh) 1845.1.8.222-1845.1.8.224, 1972.1015); and three juveniles preserved in fluid (fmnh 135427- 135429). Survey of Sample Areas — In subsequent par- agraphs, postinfantile M. mulatta specimens col- lected in 30 sample areas within the specific range (Fig. 5) are compared with the three Nepalese skins that are in prime pelage (bm(nh) 1921.5.1.1, 1922.5.16.2, 1931.1.11.11). Museum catalog numbers are indicated where the number of spec- imens in a cited sample subset is less than five. 1. India: northern Uttar Pradesh, ca. 600 km west-northwest of Hazaria Patherghatta; 12 skins, seven localities. Eleven specimens, collected Oc- tober-March, are in prime pelage, and one (BM(NH) 1931.1.11.3), collected in January, is ear- ly-stage faded. In seven of the specimens in prime pelage, collected October-March, dorsal pelage color is essentially similar to that in the three Nep- alese standards; in the remaining four (bm(nh) 16 FIELDIANA: ZOOLOGY Fig. 3. Dorsal pelage color in Macaco mulatta. A. Topotype — bm(nh) 1922.5.16.2. adult male, collected 17 Feb. 1921 at Hazaria Patherghatta. 600 ft (= 180 m). Nepal. B. Contrasting pelage color in two adult males collected 4 days apart at Rajapara. 600 ft (= 180 m). Assam. India— bm(nh) 1931.1.11.7. 21 Nov. 1920 (left), and bm(nh) 1921.7.9.3, 25 Nov. 1920 (right). Scale bar = 15 cm. 1914.7.10.3-5; bnhs 5108), collected October- January, the color is distinctly less erythristic — pale yellowish brown anteriorly becoming washed with burnt orange on the lumbosacral region. Bright and dull specimens have been collected in the same month at each of two localities (Bagesh- war and Ratighat). ISHL is 51.4 ± 2.9 mm (mean ± SD) in three adult females and 61.0 ± 6.5 mm in five adult males. MTHL is 20 mm and 30 mm in two adult females and 27.5 ±5.0 mm in four adult males; in two of these males (bm(nh) 1914.7.10.1 and 1914.7.10.2, both collected at Bageshwar), the tail is somewhat bushy. 2. India: northwestern Himachal Pradesh, ca. 1.000 km northwest of Hazaria Patherghatta; six skins, three localities. Of three specimens in prime pelage, two (bm(nh) 1933.12.1.2: bnhs 5112), col- lected in February and March, are brightly col- ored— similar to the Nepalese standards, and one (BNHS 5114), collected in March, is slightly less erythristic. Three specimens (bm(nh) 1923.9.1.1 18, 1931.1.11 .34-35), collected in March and May, are late-stage faded — pale yellowish gray anteriorly, faintly washed with burnt orange posteriorly. ISHL is 65 mm in one adult female and 60 and 80 mm in two adult males; MTHL is 20 and 35 mm in two adult males. 3. India: southwestern Jammu and Kashmir, ca. 1.300 km northwest of Hazaria Patherghatta; 11 skins, four localities (including type locality of Macacus rhesus villosus True, 1894). Ten speci- mens are in prime pelage, and one (usnm 173814), collected in February, is early-stage faded. Five of the specimens in prime pelage, collected in September and month unspecified (one specimen), are brightly colored — similar to the Nepalese FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 17 39 25 25 Sample size 23 22 17 11 17 34 18 15 Jun Jul Month Prime Early-stage faded Q Late-stage faded ^ 4Molting Fig. 4. Monthly incidence of prime, faded, and molting pelage stages in wild-collected specimens of Macaca mulatto. standards; the remaining five, collected in Octo- ber, November, and month unspecified (one spec- imen), are slightly less erythristic. The holotype of Macacus rhesus villosus (usnm 20120; Septem- ber) is one of the brightly colored specimens in prime pelage — golden brown anteriorly, becom- ing burnt orange on the lumbosacral region. In all these specimens, the pelage is notably dense. ISHL is 65 and 70 mm in two adult females and 75 and 85 mm in two adult males (cf. Pocock, 1932, p. 539); MTHL is 40 and 45 mm in two adult females and 40 mm in two adult males. Tails are bushy in all 11. 4. Northeastern Pakistan, ca. 1,400 km north- west of Hazaria Patherghatta; five skins, four lo- calities. One skin (usnm 353186), collected in September, is in prime pelage and is slightly less erythristic than the Nepalese standards. Two skins (bm(nh) 1923.11.4.1; bnhs 5113), collected in June, are late-stage faded. One skin (usnm 353187), collected in August, is in the process of molting; most of the pelage consists of short (30 mm) bright new hairs, but scattered among these are a few long (90 mm) faded old hairs. In the fifth skin (USNM 326332), collected in September, the molt apparently had just been completed; the fur is short (35 mm) and bright, similar in color to that in the Nepalese standards. ISHL is 65 mm in one adult female and 50 and 70 nrmi in two adult males; MTHL is 30 and 40 in two adult males. In the male in prime pelage, the tail is bushy. 5. Northwestern Pakistan and eastern Afghani- stan, ca. 1,600 km northwest of Hazaria Pather- ghatta; three skins, three localities (including type locality of Macaca mulatta mcmahoni Pocock, 18 FIELDL\NA: ZOOLOGY 1932). These three specimens — one wild-collect- ed and two captives — are the only preserved skins known to have originated at the northwestern lim- it of distribution of M. mulatta. The wild-collected specimen (bm(nh) 1920.6.11.1), an adult male, was taken in early February 1914 in Pakistan near the border with Afghanistan. One of the captives (bm(nh) 1931.1.9.1. skin only) apparently origi- nated in eastern Afghanistan (Pocock, 1932, p. 543); it was received at the Regent's Park Zoo- logical Gardens, London, on 3 April 1906, died there on 19 January 1910, and was cataloged at bm(nh) in 1931. Available information concerning the sex of this specimen is contradictory. Al- though zoo records and the zoo tag on the skin indicate that the specimen was a male (P. Jenkins, bm(nh), letter, 21 June 1995), bm(nh) records and the museum tag on the skin indicate that it was a female (Pocock, 1932, p. 543); the skull has not been preserved, and my examination of the skin for evidence of sexual characters was inconclu- sive. Pending further information, the zoo records are regarded as more reliable, and the specimen is considered to be a male; it probably was an adult (cf. Pocock, 1932, p. 543). The other captive (FMNH 102839), a juvenile female that reportedly also originated in eastern Afghanistan, was pur- chased on 2 November 1965 near Gandahar, Af- ghanistan, ca. 600 km southwest of the reported place of original capture (Hassinger, 1968, p. 72). Pelage color is diverse in these three skins. The captive male, which died in January, is in prime pelage — dark golden brown on the dorsal thoracic region and bright burnt orange on the lumbosacral region; the color of the lumbosacral region is sim- ilar to that in the Nepalese standards, but the color of the dorsal thoracic region is much darker than in most specimens of M. mulatta. The wild-col- lected male (bm(nh) 1920.6.11.1, holotype of Ma - caca mulatta mcmahoni), taken in February, is yellowish gray anteriorly, becoming washed with burnt orange posteriorly. The shaggy, dull-col- ored, weakly annulated fur probably indicates that this specimen is early-stage faded; this interpre- tation differs from that of Pocock (1932, p. 545), who noted that the pelage of this male is similar to that of a late-stage faded specimen (bm(nh) 1923.11.4.1) collected at Patriata, Murree, Paki- stan, but concluded that, because bm(nh) 1920.6.11.1 was collected in February, it should be in prime pelage (cf. late-stage faded bm(nh) 1923.9.1.118, India: Kangra Fort, Himachal Pra- desh, collected 18 March). The captive juvenile female, obtained in November, is relatively dull- colored — yellowish brown anteriorly, becoming golden brown on the sacral region. This skin is more uniformly brown than most specimens of M mulatta; the texture of the pelage does not suggest that the specimen is seasonally faded. In the adult males, ISHL is 50 and 90 mm, and MTHL is 30 and 50 mm. The tails of the adult males are bushy. 6. India: central Madhya Pradesh and southern Gujarat, ca. 600 to 1 ,400 km southwest of Hazaria Patherghatta; six skins, four localities. One skin from Gujarat (bm(nh) 1931.1.11.3), collected in 1922-1923, is in prime pelage and is brightly col- ored but is slightly paler than the Nepalese stan- dards. Another skin from Gujarat (bm(nh) 1931.1.11.2), also collected in 1922-1923 and brightly colored, appears from its disheveled pel- age to be early-stage faded. Two skins from Mad- ya Pradesh (bm(nh) 1931.1.11.4-5), collected in April and May, are late-stage faded. The remain- ing two skins (bm(nh) 1931.1.11.1; bnhs 5107), collected in 1922-1923 and date unknown, are pale juveniles. ISHL is 46.7 ± 2.9 mm in three adult females and 50 mm in one adult male; MTHL is 20.0 ± 0.0 mm in the females and 20 mm in the male. 7. India: southwestern Bihar, western and southern Orissa, and eastern Andhra Pradesh, ca. 600 to 1,300 km south-southwest of Hazaria Path- erghatta; seven skins, five localities. One skin (bm(nh) 1928.3.7.4), collected in Orissa in Sep- tember, is in prime pelage and is similar to the Nepalese standards. Two skins (zsi Coll. No. OM/ DD/30; ZSI, Siddeldar Hill, unnumbered), collect- ed in Orissa and Andhra Pradesh in November and December, are in prime pelage but are dis- tinctly less erythristic than the Nepalese stan- dards. The remaining four skins were collected in Bihar and Orissa in August; of these, three (bm(nh) 1915.4.3.2, 1928.3.7.3; bnhs 5089) were in the process of molting, and one (bm(nh) 1915.4.3.1) is newly molted. The bright new fur in all four is similar in color to that in the Nep- alese standards. ISHL is 15 mm in one adult fe- male (molting, new fur), 30 mm in a subadult or adult female (skin only), 50 mm in a subadult or adult male (skin only), and 65 mm in an adult male; MTHL is 20 mm in the subadult or adult female and 30 mm in the subadult or adult male. 8. Bangladesh: Sundarbans and India: Tripura, ca. 700 km southeast of Hazaria Patherghatta; nine skins, three localities. The two specimens from Tripura, an adult male (zsi Coll. No. TM18) collected in November and a subadult male (zsi Coll. No. TM4) collected in January, are in prime FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 19 40* 20, Pelage color sample 30% 2(f- ^(f-~ 21 • r 22 23 110° Fig. 5. Sample areas cited in Macaco mulatta pelage comparisons. Key to included localities (for details, see Gazetteer, Appendix 2): Nepalese standards — Hazaria Patherghatta; Nagarkot; Chengli. 1. Northern Uttar Pradesh, India — Bageshwar: Dela: Haripur; Jhirna: Rammagar: Ratighat; Sita Bani. 2. Northwestern Himachal Pradesh. India — Dharmsala; Kangra; Samayala. 3. Southwestern Jammu and Kashmir, India — Dunwein; Kashmir: Kotihar; Lolab. 4. Northeastern Pakistan — Dunga Gali vicinity, 2470 m; Ghora Dhaka; Paia; Patriata. 5. Northwestern Pakistan and eastern Afghanistan — Chigha Sarai: Kaotai; Nurestan, eastern. 6. Central Madhya Pradesh and southern Gujarat, India — Dangs: Kakara; Malua: Sohagpur. 7. Southwestern Bihar, western and southern Orissa, and eastern Andhra Pradesh, India — Deogarh; Gudari; Luia: Malkangiri: Siddeldar Hill. 8. Sundarbans, Bangladesh, and Tripura, India — Ampi Bazar; Charilam; Sundarbans, ca. 50 mi east of Calcutta. 9. Sikkim and northern West Bengal, India — Bhar- nabhari; Hasimara; Mangpu; Narbong; Sikkim; Sivok; Sukna. 10. Assam, Meghalaya, Manipur. and Nagaland, India — Bishenpur; Bogra Nadi; Changchang Pani; Golaghat; Hot Springs; Imphal, ca. 4 mi north of; Kulsi; Lamsakhang; Nangpoh; Samaguting. 1 1. Anmachal Pradesh, India — Dening; Margherita. 12. Eastern Xizang, China — Yigong. 2250 m. 13. South-central Qinghai. China — Jegu Xiang. 14. Northern Myanmar — Bawmwang; Bhamo; Heinsun; Hiswehl; Hkandau; Homalin; Htingnan Triangle; Karen Chaung; Maungkan; Moklok; Nanyaseik; N"Changyang; Singkaling Hkamti (24 July. 5 Aug., and June-Aug. 1914); Taga Hka; Tanga-Shingaw; Tang Hpre; Taro. 15. Western Yunnan, China — Ashi; Biloxue Shan; Datang; Hotha Valley; Hui-yao; Santaishan; Tengchong. 16. West-central Sichuan, Chi- na— Gin Keo Ho; Kangding; Leshan; Olongchc; Wa Shan. 17. Southeastern Sichuan, China — Tseo-Jia-Geo; Yibin; Yunnan border, south of Yibin. 18. Guizhou, China — Getou; Zunyi. 19. Northeastern Sichuan, China — Tonojiang Xian. 20. Northeastern Hebei, China — Xinglong Xian. 21. Northern Fujian, China — Chong'an Xian; Kuatun. 22. Dawanshan Islands, China — Dangan Dao; Neilingding Dao; Shangchuan Dao (Miwan). 23. Hainan Dao, China — Bawangling; Changtian; Dongfang; Hainan; Henron; Jianfengling; Mihouling; Nada; Nanfeng Shi; Nanwan. Xingcun- . gang; Nychow; Pisui; Wuzhi Shan; Xinlong; Yiajia; Zhayun. 24. Northeastern Vietnam and southern Guangxi, Chi- na— Bac Can; Ban Thi; Chiem Hoa; Linh Thong; Ly Bon; Nanning; Nghia Dan; Nghia Dung; Thanh Tuong; Van Hai; Yen Bai. 25. Northern Laos, northwestern Vietnam, and southern Yunnan, China — Lai Chau; Mengla Xian; Menglun; Muong Bourn; Muong Muon; Muong Pon; Ou, Nam; Shanman; Xishuangbanna. 26. Central Myanmar — Ali Cha; Kin; Kindat. 20 mi northwest of; Kokkoaing; Lethan Hka; Madaya; Mansam Falls; Maymyo; Mingun; Popa Hill (1000 m; 1512 m); Pyaunggaung; Se-eng; Tatkon, east bank of Chindwin River; Tatkon, west bank of Chindwin River; Yin, east bank of lower Chindwin River; Yin. lower Chindwin River. 27. Southwestern Myanmar — Pye, 30 mi southeast of; Pye 35 mi southeast of; Toungoo, 13 mi east of; Toungoo, 15 mi north of; Toungoo, 20 mi west of; Toungoo, 30 mi northwest of; Toungoo, east side of Sittang River. 28. Northwestern Thailand — Ban Mae Lamao; 20 FIELDIANA: ZOOLOGY pelage but are darker and less erythristic than the Nepalese standards. The adult is dark golden brown anteriorly, becoming strongly washed with burnt orange posteriorly; the subadult is grayish tipped with yellowish anteriorly, becoming faintly washed with burnt orange posteriorly. ISHL is 55 mm in the subadult and 65 mm in the adult. The seven Sundarbans skins, collected in April 1870. are difficult to interpret. The pelage, which is harsh in texture, is grayish brown to golden brown anteriorly, becoming variably washed with burnt orange posteriorly. The unusual pelage con- dition may be a result of seasonal fading, deteri- oration in storage, or both. Feeroz et al. (1995. p. 75) report that pigmen- tation of the face and ventral surface in living M. midatta observed at Sitakunda. southeastern Bangladesh, appeared to differ from that in M. mulatta observed elsewhere in Bangladesh. No further details concerning this color difference are specified. 9. India: Sikkim and northern West Bengal, ca. 400 km east of Hazaria Patherghatta; 10 skins, seven localities. Eight specimens are in prime pel- age. Four of these (bm(nh) 1916.7.29.2. 1931.1.11.9; FMNH 35447, 35448), collected in November-January, are brightly colored — similar to the Nepalese standards; one (fmnh 35449). col- lected in December, is slightly less erythristic; two (BM(NH) 1891.10.7.4, 1916.7.29.1), collected in January and month unknown, are slightly darker; and one (bm(nh) 1931.1.11.8), collected in Feb- ruary but apparently in prime pelage, is much darker and less erythristic — grayish brown ante- riorly, becoming faintly washed with burnt orange on the sacral region. In the remaining two speci- mens, seasonal fading may have begun. One of these (bm(nh) 1915.9.1.1). collected in March, is paler and has weaker agouti hair banding than the preceding eight specimens; the other (zsi 7294), collected in April, is drab yellowish brown ante- riorly and bright burnt orange posteriorly. ISHL is 45.0 ± 8.7 mm in three adult females and 55.0 ±5.0 mm in five adult males; MTHL is 20.0 ± 0.0 mm in three adult females and 28.8 ± 6.3 mm in four adult males. The tail is bushy in one adult male (bm(nh) 1891.10.7.4, the only postinfant specimen available from Sikkim). 10. India: Assam, Meghalaya, Manipur, and Nagaland. ca. 600 to 1.000 km east-southeast of Hazaria Patherghatta; 15 skins. 12 localities. Nine specimens in prime pelage, collected August-Feb- ruary, are brightly colored — similar to the Nep- alese standards. Two others in prime pelage, a male (zsi 11187) collected in November and an- other male (bm(nh) 1943.60) collected in Febru- ary, are somewhat darker and less erythristic — dark golden brown anteriorly, faintly washed with burnt orange posteriorly; although both of these relatively dull specimens were collected in Man- ipur, one of the bright specimens (bm(nh) 1943.61) also was collected in that state. One skin (bm(nh) 1931.1.11.15), collected in May, is late- stage faded. Two specimens (bm(nh) 1921.7.9.4; BNHS 5087), collected in July and September, were in the process of molting; the short bright new fur is sparsely covered by long faded old hairs. The most aberrant specimen in this group is a very dark and dull-colored adult male (bm(nh) 1921.7.9.3) collected on 25 November 1920 at Rajapara, Assam (Fig. 3B). This specimen, which is in prime pelage, is dark yellowish gray anteri- orly, becoming faintly washed with burnt orange on the lumbosacral region. Pocock (1932. p. 530) has drawn attention to the striking contrast be- tween this adult male and a brightly colored adult male (bm(nh) 1931.1.11.7; included among nine bright specimens cited above) that was collected 4 days earlier at the same place (Fig. 3B); Pocock cited these two specimens as an illustration of the broad range of individual variation to which pel- age color is subject in M. mulatta. ISHL is 45 and 60 mm in two adult females and 50.0 ± 8.7 mm in three adult males; MTHL is 20 and 25 mm in two adult females and 20 and 25 mm in two adult males. 1 1. India: Arunachal Pradesh, ca. 1.100 km east of Hazaria Patherghatta; four skins, two localities. These four skins (bm(nh) 1931.1.11.13-14; bnhs 5086; ZSI 12090), collected in November-April, are similar to the Nepalese standards. In two adult Ban Umphang; Chiang Mai; Huai Ap Nang; Huai Kwang Pah; Kaeng Mae Hat. 29. Northeastern Thailand and west- central Laos — Ban Mak Nao; Dan Sal District; Mekong River; Nong Khai: Pang Nam Un. 30. Southern Laos and Central Vietnam — Dak Sut: Ky Son; Song-Ta-Voy; Son Tra Ml., 3.9 km west and 0.3 km south of; Thateng, Muang; Xuan Ninh. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 21 females, ISHL is 50 and 60 mm and MTHL is 20 and 30 mm. 12. China: eastern Xizang, ca. 1,000 km north- east of Hazaria Patherghatta; one skin, one local- ity. This skin (nwpib Coll. No. 73066, juvenile female), collected in June, is late-stage faded; the dorsal pelage is gray tipped with golden anteri- orly, becoming washed with pale burnt orange posteriorly. The specimen's faded condition hin- ders comparison with the Nepalese standards. ISHL is 55 mm (juvenile). 13. China: south-central Qinghai, ca. 1,300 km northeast of Hazaria Patherghatta; two skins, one locality. One skin (nwpib Coll. No. 63167), col- lected in June, is early-stage faded — gray tipped with golden anteriorly, becoming reddish brown posteriorly; the other (nwpib 00033), collected in May, is late-stage faded — pale gray tipped with gold anteriorly, becoming weakly washed with burnt orange posteriorly. Seasonal fading in these skins hinders comparison with the Nepalese stan- dards. Both specimens are juvenile males with long hair and bushy tails; ISHL is 65 and 70 mm, and MTHL is 40 and 50 mm. 14. Northern Myanmar, ca. 1,000 to 1,300 km east-southeast of Hazaria Patherghatta; 27 skins, 17 localities. Ten specimens collected in October- March are in prime pelage; four collected in Jan- uary-May are early-stage faded; five collected in January-April are late-stage faded; and eight col- lected in July-August were in the process of molt- ing. Of the 10 specimens in prime pelage, seven are brightly colored and generally similar to the Nepalese standards. The remaining three in prime pelage are somewhat deviant: one (bm(nh) 1950.373) is brightly colored but has unusually dark gray hair bases, one (amnh 112734) is no- tably paler than the other nine, and one (usnm 279191) has the burnt orange coloration narrowly restricted to an area near the ischial callosities. Of the four early-stage faded specimens, two (bm(nh) 1937. 12.3.75, 1937.12.3.77), collected in May, are slightly more erythristic than the Nepalese standards; one (bm(nh) 1950.372), collected in January, is brightly colored but has unusually dark gray hair bases; and one (amnh 1 12739), col- lected in March, is less erythristic. Of the five late-stage faded specimens, three (amnh 112740, 112741, 114547) were collected in March or April, which is unremarkable; the other two (amnh 112722, 112723), somewhat incongruous- ly, were collected in January. ISHL is 50.0 ± 6.0 mm in eight adult females and 50 and 65 mm in two adult males (excludes molting specimens); MTHL is 19.3 ± 6.7 mm in seven adult females and 30 and 35 mm in two adult males. 15. China: western Yunnan, ca. 1,500 east- southeast of Hazaria Patherghatta; 1 3 skins, seven localities. Of nine specimens in prime pelage, eight collected in December (five skins), April (one skin), and month unknown (two skins) are brightly colored — similar to the Nepalese stan- dards; the other specimen (zsi 11986), collected in July, is somewhat less erythristic. Two skins (AMNH 43084, 43086), collected in April, are late- stage faded, and a hunter's flat skin (kiz Coll. No. 780417), reportedly taken in October, is molting. One skin (zsi 619), collected in March-July 1868, has become severely deteriorated in storage. In one adult female, ISHL is 60 mm, and MTHL is 15 mm. 16. China: west-central Sichuan, ca. 1,600 to 1,800 km east-northeast of Hazaria Patherghatta; six skins, five localities (including type locality of Macacus vestitus Milne-Edwards, 1892). Four skins collected in February, March, and June are in prime pelage; one (mnhn 1891/388, holotype of Macacus vestitus), collected in June or July, is early-stage faded; and one (usnm 241160), col- lected in July, is late-stage faded. Two of the spec- imens in prime pelage (rmnh 4585/V67, 4585/ W50, March) are similar in color to the Nepalese standards, one (mnhn 1891/387; June) is slightly less erythristic, and one (bm(nh) 1911.9.8.1; Feb- ruary) is darker and more erythristic — rich golden brown anteriorly, becoming dark burnt orange on the lumbosacral region; the last of these is very slightly paler than bm(nh) 1950.373, collected at Bawmwang, northern Myanmar. In the early-stage faded holotype of Macacus vestitus, dorsal pelage is pale grayish tipped with golden anteriorly, be- coming pale burnt orange on the lumbosacral re- gion. In two adult females, ISHL is 60 and 80 mm, and MTHL is 40 and 50 mm; in one of these (bm(nh) 1911.9.8.1), the tail is bushy, particularly distally. Three additional specimens, with vague locality information, may also have been collected in or near west-central Sichuan. One of these is bm(nh) 1871.4.21.4, the holotype of Macacus lasiotus Gray, 1868. This tailless adult male was a captive that reportedly originated in "Szechuen"; it reached London, via Shanghai, shortly before 15 January 1868 and died on 25 May 1870. It is in prime pelage, dark and strongly erythristic; the dorsal pelage is dark burnt orange anteriorly, be- coming intensely burnt orange on the sacral re- gion; ISHL is 65 mm. The nearest match is 22 FIELDIANA: ZOOLOGY BM(NH) 1927.12.1.18 (Bac Can. northeastern Viet- nam), which is slightly paler and less erythristic. The other two specimens possibly collected in west-central Sichuan (mnhn 1892/315, 1894/ 1432) are of limited value for pelage comparison. The former is late-stage faded, and the latter has suffered severe postmortem deterioration and dis- coloration. These two specimens were taken in May-July 1890 by the collectors of mnhn 1891/ 387 and 1891/388 cited above; the only locality information available is "Tibet," which, for the collectors, included part of what is now Sichuan. 17. China: southeastern Sichuan, ca. 1,900 km east of Hazaria Patherghatta; four skins, three lo- calities. Two specimens collected in October and January are in prime pelage, and two (usnm 258183. 258184). collected in February and March, are late-stage faded. Dorsal pelage color- ation in the two in prime pelage, both juveniles, is dissimilar and unusual. The younger specimen (LiSNM 239133; Ml), collected in October, is pale but erythristic — pale golden brown anteriorly, be- coming intensely burnt orange on the lumbosacral region; the older juvenile (usnm 256669; 11-2, M2), collected in January, is much darker and less erythristic — yellowish brown anteriorly, becom- ing golden brown on the lumbosacral region. In one adult female, ISHL is 55 mm, and MTHL is 40 mm. 18. China: Guizhou, ca. 2,100 to 2,300 km east of Hazaria Patherghatta; six skins, two localities. Four specimens (bm(nh) 5.66.150-153), collected in May, are late-stage faded, and two (Kiz 03179, 03181), collected in early September, were in the process of molting. Because of fading or molting, these specimens cannot be compared with the Nepalese standards. MTHL is 40 and 45 mm in two adult females; in the latter specimen, the tail is bushy. 19. China: northeastern Sichuan, ca. 2,200 km northeast of Hazaria Patherghatta; two skins, one locality. These two adult females, collected in July, had just begun to molt on their crowns and tails; the dorsal pelage is late-stage faded. In one specimen (siz 00001), the upper back is grayish, and the lower back is pale golden brown; in the other {.SIZ 00002), the upper back is pale golden brown, and the lower back is golden brown. No comparison can be made with the Nepalese stan- dards. ISHL is 45 and 50 mm. 20. China: northeastern Hebei, ca. 3,200 km northeast of Hazaria Patherghatta; 10 skins, one locality (type locality of Macacus tcheliensis Milne-Edwards, [1872]). Five specimens are in prime pelage, four (fmnh 39376, 39377; mnhn 335/381A/1867:557; usnm 240705) appear to be early-stage faded, and one (amnh 57039) is late- stage faded; the month of collection or death is reliably known for only one of these 10 specimens (bm(nh) 1931.1.7.2, male in prime pelage, re- ceived at Regent's Park Zoo 17 August 1880. died 6 March 1881). Of the specimens in prime pelage. bm(nh) 1931.1.7.2 is long-haired and very bright- ly colored — pale burnt orange anteriorly, becom- ing intensely burnt orange on the lumbosacral re- gion; this skin is similar in color to bm(nh) 1937.12.3.7.7 (Karen Chaung. northern Myan- mar) and is somewhat more erythristic than the Nepalese standards. Another specimen in prime pelage (amnh 57040) also is strongly erythristic but is much darker — dark golden brown anteri- orly, becoming burnt orange posteriorly. The re- maining three specimens in prime pelage (amnh 57042; fmnh 39378; usnm 240704) are darker and less erythristic than the Nepalese standards. The holotype of Macacus tcheliensis (mnhn 335/ 381 A/1 867:557). which apparently is early-stage faded, is less erythristic than the Nepalese stan- dards— pale golden brown anteriorly, becoming washed with burnt orange on the lumbosacral re- gion. In two adult males. ISHL is 70 mm (both specimens), and MTHL is 30 and 35 mm. 21. China: northern Fujian, ca. 3.300 km east of Hazaria Patherghatta; six skins, two localities (including type locality of Pithecus littoralis El- liot. 1909). Three specimens collected in Novem- ber and May are in prime pelage; one (amnh 84476), collected in August, was in the process of molting, and two (mnhn 1874/480, 1874/481, Kuatun), collected in November 1873, are severe- ly faded as a result of extended postmortem ex- posure to light. Of the three specimens in prime pelage, the holotype oi Pithecus littoralis (bm(nh) 1900.5.8.1; November) is the brightest — pale golden brown anteriorly, becoming washed with burnt orange on the sacral region; the coloration of this specimen, which almost perfectly matches that of bm(nh) 1931.1.11.7 (India: Rajapara, As- sam State; Fig. 3B), is similar to that of the Nep- alese standards. Another of the specimens in prime pelage (amnh 84474; May) is slightly less erythristic, and the third (bm(nh) 1898.11.1.29; May) is somewhat darker and less erythristic. ISHL is 40 mm in one adult female (holotype of Pithecus littoralis) and 80 mm in one adult male; MTHL is 25 mm in the female and 40 mm in the male; the tail of the female is relatively bushy. 22. China: Dawanshan Islands, South China Sea, FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 23 ca. 3,000 km east-southeast of Hazaria Patherghat- ta; five skins, three localities (including type local- ity o\' Iniius scimtijohannis Swinhoe. [1867]). One skin (sciEA Coll. No. 2155), collected in October, is in prime pelage; three (.scika Coll. Nos. 2150, 2151, 2153), collected in March and April, are late- stage faded; and one (bm(nh) 1868.12.29.10, ho- lotype of Inuus sanctijohannis), collected in month unknown, was in the process of molting. The spec- imen in prime pelage is brightly colored, similar to the Nepalese standards. Hair length measurements of adults are not available. 23. China: Hainan Dao, ca. 2,700 km southeast of Hazaria Patherghatta; 30 skins, 16 localities (including type locality of Pithecus brevicaudus Elliot, 1913). Twenty-one specimens, collected in October-April, are in prime pelage; four (amnh 27569, 27575, 60038; zmb A1904.09), collected in October and March, are early-stage faded; and five, collected in March, May, and July, are late- stage faded. Eleven of the 21 specimens in prime pelage (collected October-February) are brightly colored — similar to the Nepalese standards; these bright specimens include amnh 27577, the holo- type of Pithecus brevicaudus, which closely matches amnh 1 12733 (northern Myanmar: Taro). Seven of the specimens in prime pelage, collected in October-April, are slightly less erythristic than the Nepalese standards. The remaining three (AMNH 27570, October; bm(nh) 1909.7.11.1, Oc- tober; sciEA Coll. No. 0089, December) are no- tably less erythristic than the Nepalese standards. ISHL is 46.2 ± 4.8 mm in four adult females and 45 and 70 mm in two adult males; MTHL is 27.5 ± 2.9 mm in four adult females and 20 and 30 mm in two adult males. 24. Northeastern Vietnam, and China: southern Guangxi, ca. 2,200 to 2,400 km southeast of Ha- zaria Patherghatta; 20 skins, 1 1 localities. Six specimens collected in November-January are in prime pelage, eight collected in December-June (7) and month unknown ( 1 ) are early-stage faded, one collected in June is late-stage faded, and four collected in October-January have short bright fur and apparently had just completed the molt; a cap- tive (Kiz Coll. No. 631425), obtained in Nanning, China, and kept alive for an unknown period at KIZ, is long-haired and appears somewhat faded. All six specimens in prime pelage were collected in Vietnam; of these, two closely match the Nep- alese standards (bm(nh) 1927.12.1.19, Bac Can; ZMVNU 06/3.16.4, Van Hai), two are slightly dark- er (bm(nh) 1927.12.1.18, Bac Can; iebr 733 (833)7560/175, Nghia Dung), one is slightly less erythristic (iebr 33, Ly Bon), and one is notably less erythristic (zmvnu 167/3.18.14, Chiem Hoa). In adult females, ISHL is 45.0 ± 8.9 mm (n = 8), and MTHL is 24.3 ± 7.9 mm (n = 7); in one adult male, ISHL is 45 mm, and MTHL is 25 mm. The tail is moderately bushy to bushy in four of the females. 25. China: southern Yunnan, northern Laos, and northwestern Vietnam, ca. 1,600 to 1,900 km southeast of Hazaria Patherghatta; 12 skins, nine localities. Three skins (amnh 87264; fmnh 31766; KIZ Coll. No. 75840), collected in November- May, and two skins (kiz 03172, 03180), collected in month unknown, are in prime pelage and are brightly colored — similar to the Nepalese stan- dards; one skin (iebr D3/M37), collected in month unknown, is early-stage faded; one skin (fmnh 31763), collected in May, and two (kiz 03173, 03174), collected in month unknown, are late- stage faded; and a captive (amnh 87278). obtained in November 1931, was in the process of molting when it died on 15 June 1932. The remaining two specimens are somewhat aberrant in their dorsal pelage coloration: a juvenile male (kiz 000150), collected in October, is much less erythristic than the Nepalese standards — grayish tipped with golden anteriorly, becoming pale golden brown posteriorly; and a ?subadult female (kiz 000153, skin only), collected in November, is more brown- ish and less erythristic than usual in M. mulatta. Hair length measurements of adults are not avail- able. 26. Central Myanmar, ca. 1,000 to 1,300 km southeast of Hazaria Patherghatta; 33 skins, 16 lo- calities. Fifteen specimens collected in September- May are in prime pelage, 15 collected in May- August are late-stage faded, and three (bm(nh) 1914.7.19.1, 1931.1.11.23, 1931.1.11.27). collected in June-July, were in the process of molting. Of the specimens in prime pelage, seven were col- lected at various localities, and eight were collected at Popa Hill. The seven collected at various local- ities average slightly paler and less erythristic than the Nepalese standards; one of these (bm(nh) 1931.1.11.26; January) is similar to the Nepalese standards, five are slightly paler and/or less eryth- ristic, and one (bm(nh) 1931.1.11.22; January) is much darker and less erythristic — yellowish gray anteriorly, washed with burnt orange posteriorly. The eight collected at Popa Hill average notably less erythristic than the Nepalese standards; one (bnhs 5106; September) is similar to the Nepalese standards, two (amnh 16361 1, 163613; September- October) are slightly paler and less erythristic. 24 FIELDIANA: ZOOLOGY three (amnh 163612. 163614, bm(nh) 1914.7.19.2; September-October) are more grayish, and two (amnh 16310, 16315; October-November) are un- usual in their strongly mottled (coarsely agouti) pelage. ISHL is 44.6 ± 5.6 mm in 13 adult females and 46.2 ± 9.5 mm in four adult males; MTHL is 21.9 ± 5.9 mm in eight adult females and 23.3 ± 2.9 mm in three adult males. 27. Southwestern Myanmar, ca. 1.400 km southeast of Hazaria Patherghatta; 10 skins, seven localities. Of seven skins in prime pelage, two (BM(NH) 1931.I.1I.20-21), collected in September and October, are similar to the Nepalese stan- dards, and five, collected in November-February, are somewhat less erythristic. One of two skins collected at Toungoo in May (bm(nh) 1931.1.11.18) is early-stage faded — somewhat paler than the Nepalese standards, and the other (bm(nh) 1931.1.11.19) is late-stage faded. Unac- countably, the skin of an early juvenile (bm(nh) 1931.1.11.17), collected near Toungoo in Decem- ber, also is faded — pale yellowish brown anteri- orly, becoming faintly washed with burnt orange on the sacral region (cf. Pocock, 1932. p. 533). ISHL is 43.0 ± 5.7 mm in five adult females; MTHL is 18.3 ± 2.9 mm in three adult females. 28. Northwestern Thailand, ca. 1.700 to 2.000 km southeast of Hazaria Patherghatta; six skins, six localities (including type locality of Macaco siamica Kloss, 1917). Only one of these skins (ZRC 4-154), month of collection unknown, is in prime pelage; this bright skin is similar to the Nepalese standards. Three skins (ctnrc, catalog number unknown; fmnh 99668; zrc 4-188, ho- lotype of Macaca siamica). collected in March- April, are early-stage faded, and one (fmnh 99669). collected in March, is late-stage faded. One skin (amnh 54816, subadult male), collected in early February, appears to have been in the process of molting; this does not accord with the late summer molting schedule that generally ap- plies in M. mulatto. ISHL is 50 mm in one adult female and 50 and 70 mm in two adult males; MTHL is 25 mm in one adult female and 35 mm in one adult male. 29. Northeastern Thailand and west-central Laos, ca. 1,800 to 2,100 km southeast of Hazaria Patherghatta; eight skins, five localities. Five skins, collected in January-March and month un- known (one specimen), are in prime pelage; of these, three (usnm 300017; zrc 4-150. 4-151). collected in February-March, are brightly col- ored— similar to the Nepalese standards, and two (usnm 296917. 307716), collected in January and month unknown, are somewhat less erythristic. One skin (zrc 4-152), collected in February, is early-stage faded; one (usnm 307715), collected in month unknown, is late-stage faded; and one (USNM 240488), collected in July, was molting. ISHL is 45 mm in two adult females and 45.0 ± 5.0 mm in three adult males; MTHL is 20 mm in one adult female and 25 mm in one adult male. 30. Southern Laos (one locality) and central Vi- etnam, ca. 2,800 km southeast of Hazaria Path- erghatta; nine skins, six localities. Dorsal pelage color is highly variable in these specimens, all of which appear to be in prime pelage. One skin (iebr 560/3, ?adult male, Xuan Ninh) is golden brown anteriorly and burnt orange posteriorly, similar to the Nepalese standards. Two others (usnm 356968, subadult female. Mt. Sontra; usnm 320780. adult female. Dak Sut) are almost uni- formly golden brown anteriorly and posteriorly, approximately as in M. fosciciilaris (cf. Fooden, 1995, p. 25; 1997, p. 227); these two specimens are now allocated to M. mulatto solely on the ba- sis of their relative tail length (usnm 356968, 59.7%; usnm 320780, 64.5%). In the remaining six skins, dorsal pelage color is slightly brighter posteriorly than anteriorly, variably intermediate between that in typical M. mulatto and M. fasci- cularis (iebr 40. ?adult male. Ky Son; mnhn 1899/ 54, adult female. Song Ta-Voy; ansp 15135, ju- venile female, and ansp 15138, juvenile, Muang Thateng, Laos; usnm 320781, adult female, and usnm 320782, juvenile male. Dak Sut). In two adult females, ISHL is 65 mm, and MTHL is 10 and 15 mm; in two ?adult males, ISHL is 50 mm, and MTHL is 30 and 40 mm. Summary — Judging from specimens examined, there is no general pattern of geographic variation in dorsal pelage color in M. mulatto. Of 166 postin- fantile specimens in prime pelage from 30 sample areas, 78 are similar in color to the Nepalese stan- dards, 55 are variably less erythristic, and 33 from scattered sample areas are either more erythristic, darker, paler, browner, or more mottled. Individual variation is great among specimens from the same sample area and even from the same locality; this is vividly demonstrated by the two contrastingly colored adult males collected 4 days apart at Ra- japara, Assam, India (Fig. 3B). Conversely, speci- mens from widely separated parts of the specific range may be nearly identical in color, as illustrated by the following examples of matching pairs: usnm 326332, Gora Dhaka, Pakistan, and usnm 240175, Ashi, Yunnan, China; fmnh 35448, Mangpu, Sik- kim. India, and fmnh 31766, Muong Boum. Viet- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 25 nam; bm(nh) 1931.1.11.7. Rajapara, Assam, India, and bm(nh) 1900.5.8.1, Kuatun, Fujian, China; A.viNH 112733, Taro, Myanmar, and amnh 27577, Wuzhi Shan, Hainan, China; amnh 112732, Taro. Myanmar, and amnh 57043, Xinglong Xian ( = Eastern Tombs), Hebei, China; and bm(nh) 1931.1.11.17, Toungoo, 30 mi northwest. Myan- mar. and BM(NH) 1870.7.18.19. Nychow. Hainan. China. Limited available evidence suggests that spec- imens from northwestern Pakistan and eastern Af- ghanistan (Sample Area No. 5). Tripura, India (No. 8), and China, northeastern Hebei (No. 20) may tend to average somewhat darker than usual in M. midatta and that specimens from central Myanmar (No. 26) and southwestern Myanmar (No. 27) may tend to average somewhat less er- ythristic. Dorsal pelage color in southern Laos and central Vietnam (No. 30) is transitional between that in typical M. mulatta and neighboring M. fas- cicularis. ISHL averages greater in the northern part of the geographic range of M. mulatta. north of ca. 28°N latitude, than in the southern part of the range (Fig. 6). MTHL, which presumably is cor- related with perceived bushiness of the tail, also averages greater in the northern part of the range (Fig. 7; cf. Roonwal & Tak. 1981. p. 96; Tak & Kumar. 1984. p. 203). In the original description of Macacus lasiotus Gray, 1868 (p. 61), hairiness of the ears is casually cited as a diagnostic character of rhesus macaques in Sichuan, China (cf. Jiang Xuelong et al., 1991, p. 244). This appears to be invalid, as indicated by the following hst of bm(nh) specimens of M. mu- latta in which hairiness of the ears equals or exceeds that in the holotype of Macacus lasiotus (bmnh 1871.4.21.4): Afghanistan— 1931.1.9.1; Myan- mar—1931.1.1 1.21, 1931.1.11.24; China. Fujian— 1900.5.8.1; China, Hubei— 1931.1.7.2; India. As- sam— 1921.7.9.4; India. Jammu and Kashmir — 1871.3.3.5; hidia. Sikkim— 1891.10.7.4; India. Uttar Pradesh— 1914.7.10.2, 1914.7.10.4; India. West Bengal— 1916.7.29.1; Nepal— 1921.5.1.2; Paki- stan—1920.6.1 1.1, 1923.11.4.1; and Vietnam— 1927.12.1.19. 1927.12.1.20. 1928.7.1.11. External Measurements and Proportions Sex and Age Variation In wild-collected adult M. mulatta specimens examined, mean head and body length in 48 males (531.8 mm) is 13% greater than in 72 fe- males (468.8 mm), and mean body weight in 25 males (7.70 kg) is 44% greater than in 33 females (5.34 kg) (Table 2). Relative length of the tail, hindfoot. and ear in adult females is similar to relative length of these appendages in adult males. From infancy to adulthood, relative length of these appendages declines, indicating that the postnatal growth rate of the appendages is less than that of the head and body (cf. Schultz, 1933, p. 12; Lumer & Schultz, 1941, p. 284). The 33% decline in relative ear length from infancy to adulthood is particularly striking. Abundant and detailed data are available con- cerning age variation of external measurements in captive M. mulatta (Hartman. 1932. p. 23; Schultz. 1933, p. 12; 1937. p. 75; van Wagenen & Catchpole. 1956. p. 248; Pickering & Kontaxis. 1961. p. 270; Kirk. 1972. p. 573; Gavan & Hutch- inson, 1973. p. 71; Kerr et al., 1974, p. 224; Cupp & Uemura, 1981, p. 113; Gribnau & Geijsberts, 1981. p. 6; Rawlins et al.. 1984, p. 254; Riopelle et al., 1986, p. 910; Sharma & Lai, 1986, p. 143; Tumquist & Kessler. 1989, p. 8; DeRousseau, 1990, p. 288; Saxton & Lotz, 1990, p. 128; Gav- an. 1991. p. 583; Zeng. 1992, p. 18; Zlamalova et al., 1994, p. 198; 1995, p. 43; Johnson & Kap- sahs, 1995a, p. 346; Vancata et al., 1995, p. 32; Blackwelder & Golub, 1996, p. 451; Clarke & Snyder, 1996, p. 86; Champ et al., 1996, p. 487; Hudson et al.. 1996. p. 198; Maity & Rathore, 1998, p. 247; Clarke & O'Neil. 1999. p. 340). In captives, sitting height — which is comparable to head and body length as measured in wild-col- lected specimens — increases from infancy to age ca. 7 to 9 years, remains fairly constant for the next 10 to 20 years and tends to decline slightly in old age; body weight similarly increases to a plateau and ultimately declines in old age. Nurs- ery-reared infants gain weight more rapidly than mother-reared infants (van Wagenen & Catchpole. 1956. p. 249; Champoux et al.. 1989. p. 115; Kriete et al., 1995. p. 16). and captive adults gen- erally weigh more than wild-collected adults (cf. Table 2; Rawlins et al., 1984, p. 253). Smith (1994c. p. 282) reports that, in captivity, weight gain from age 1 year to age 4 years is more rapid in hybrid Chinese-Indian M. mulatta than in nonhybrid Indian M. mulatta. Geographic Variation Head and Body Length — Collectors' measure- ments of head and body length in M. mulatta are 26 FIELDIANA: ZOOLOGY 90- 80- 70- 60- 50- 40- 30- E E, - 20- « 90- Q. CO B BO- ZO- 60- 50- 40- 30- 20- Males ■^- -■■^ •V- X f ■e « -X-XK- X X >0{ - -X •X- - Females n -)K -H3Df-X-»-t--^- /T\ /Ts, U-IT 1 J i-'f* ■f^t '^ ^ ffl-f— *- -t- + i 1- t=l ■«■ -Vj- x-;- X 3? • P— X- "X --£3-^ +-t-x n X + X- -X>i Q- D +■• -x-to— ■ ■h-t- 10 -r 15 20 25 30 Latitude (°N) O Bangladesh d China x India )K Thailand t^ Vietnam 35 40 45 + Myanmar A Nepal v Pakistan Fig. 6. Latitudinal variation in interscapular hair length in Macaca mulatta adults. In this graph, the measurement for one Afghan male (50 mm) is included with Pakistani data, and the measurement for one Laotian female (45 mm) is included with Thai data; measurements of interscapular hair length are not available for Bangladeshi adults. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE. MACACA MULATTA 27 60- 50- Males 40-- f- X -X-R- 30- .Q...^. -VK- XX- -^57" 20-- E E c TO 50- ■g 40- 30- 20- 10- )K « -t- X A •x-x- Females 10 {Sd-i- ^ [3=1 -**- -QQ--Q- -I- D -- t-WK— X— X a»^ + x-^ 15 20 35 25 30 Latitude (°N) O Bangladesh n China x India + Myanmar )t^ Thailand tt Vietnam 40 45 A Nepal 7 Pakistan Fig. 7. Laliludinal variation in midtail hair length in Macaco mukitta adults. In this graph, the measurement for one Afghan male (30 mm) is included with Pakistani data; measurements of midtail hair length are not available for Bangladeshi adults. available for 120 adult specimens — 72 females and 48 males — collected at 93 localities (Table 3). Although these data are invaluable, some inter- collector variation in measurement techniques is inevitable and must be borne in mind. Head and body length in both sexes tends to increase with latitude (Fig. 8. Table 3), in accord with Bergmann's rule (cf. Mayr, 1963, p. 319). However, the relationship between head and body length and latitude is not as close as that between 28 FIELDIANA: ZOOLOGY greatest length of skull and latitude (Fig. 17, Table 9). In particular, head and body length is aber- rantly large, relative to latitude, in one male col- lected in peninsular India (15°-20°N, 75°-80°E), near the southwestern limit of the specific range, and in 12 females and 6 males collected in south- eastern China and northern Vietnam (15°-25°N, 105°-110°E), near the southeastern limit of the specific range; the aberrant group of northern Vietnamese specimens includes one large female and two large males collected on Dao Cat Ba, an island in the Gulf of Tonkin, South China Sea (Fig. 8). Head and body length is small, relative to latitude, in one female collected on Neilingding Dao and one male collected on Dangan Dao, two Chinese islands in the South China Sea (Fig. 8). Jiang Haisheng et al. (1991, p. 210) compared "body length" in samples of Chinese M. mulatta from Hainan Dao (ca. 18°30'N, island) and Guangxi (ca. 23°N, mainland). Although body length in the Hainan Dao sample is less than in the Guangxi sample, the significance of this find- ing is acknowledged by the authors to be ques- tionable because no information is available con- cerning whether measured specimens were im- matures or adults. In one adult female collected on Hainan Dao that is included in the present study, the relationship between head and body length and latitude does not appear unusual (Fig. 8). Krishnan (1972, p. 541) indicates, without documentation, that body size is reduced in a pop- ulation of N. mulatta at Jaldapara Wildlife Sanc- tuary, northeastern India. Crown-rump length in M. mulatta captives im- ported from China has been compared with crown-rump length in M. mulatta captives de- scended from monkeys imported from India (Clarke & O'Neil, 1999, pp. 340, 341). In males, crown-rump length in Chinese-origin adults equaled that in Indian-derived adults; in females, crown-rump length in Chinese-origin adults was less than that in Indian-derived adults. No infor- mation is available concerning the region of ori- gin of these monkeys within India or China. Tail Length — Collectors' measurements of tail length are available for 120 adult specimens of M. mulatta (Table 4; see "Head and Body Length," p. 26). Mean tail length ( + SD) is 207.6 ± 32.72 mm in 72 adult females and 228.9 ± 35.78 mm in 48 adult males. Tail length is aberrantly large (289 mm, 298 mm) in two adult females collected in central Vi- etnam (10°-15°N, 105°-110°E; Dak Sut), at the southeastern border of the species geographic range. These two outlier specimens have previ- ously been interpreted as evidence of hybridiza- tion between M. mulatta and M. fascicularis (Fooden. 1996, p. 859; 1997, p. 228). Excluding the two aberrant Dak Sut females, tail length apparently tends to increase slightly with latitude (Fig. 9; cf. Roonwal & Tak, 1981, p. 98). More conspicuously, tail length tends to decrease with longitude, particularly east of ca. 95°E (Fig. 10); marking the eastern end of this west-east cline are six short-tailed specimens col- lected on four shallow-water islands — Cat Ba, Dangan Dao, Hainan Dao, Neilingding Dao — in the South China Sea (105°-1 15°E) and two short- tailed specimens collected in Fujian Province on the Chinese mainland (115°-120°E). The short- ness of the tail in specimens collected on Hainan Dao and in Fujian was previously noted by Elliot (1909, p. 250) and Jiang Haisheng et al. (1991, p. 210). Although the tail in M. mulatta is shorter than in most other monkeys, it retains an important function in intraspecific communication. Tail car- riage serves as a signal of dominance status both in natural populations (Neville, 1968c, p. 15; Lindburg, 1971, p. 60; Ojha, 1974, p. 164; Roon- wal & Tak. 1981, p. 96; Wada, 1984, p. 492) and in captive colonies (Altmann, 1962, p. 378; Sade, 1967, p. 101; 1971, p. 294; Waterhouse & Water- house, 1976, p. 87; cf. Rodriguez, 1998, abstract no. 307). In captivity, the tail may also function as a rudimentary prehensile organ (Erwin, 1974, p. 130). Preliminary observations in northern In- dia suggest that tail carriage may vary geograph- ically (Roonwal & Tak, 1981, p. 96; Tak & Ku- mar, 1984, p. 203). Relative Tail Length — Relative tail length, the ratio of tail length to head and body length (T/HB), is a measure of the functional and per- ceived length of the tail. Geographic variation in this ratio is given separate treatment here because variation in tail length in M. mulatta is not en- tirely congruent with variation in head and body length (see above). This ratio is available for 120 wild-collected adult specimens (see "Head and Body Length," p. 26). Because relative tail length is similar in females and males (Table 2; Fooden, 1997, p. 223), mixed-sex samples are used in the present analysis. Latitudinal variation in relative tail length in M. mulatta is relatively minor (Fig. 11, Table 5; Fooden, 1997, p. 225), excluding two aberrant specimens collected in central Vietnam (ca. 15°N; see "Tail Length," above). Longitudinal variation FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 29 Table 2. External measurcmenls and proportions in age/sex classes of wild-collected Macaco muUitta. Relative tail Relative Relative Age/sex Head and body length hindfoot length ear length Weight' class- length (mm) (T/HB X 100) (HF/HB X 100) (K/HB X 100) (kg) Infants 284.2 ± 58.6 46.8 ± 9.9 34.2 ± 5.0 12.1 ± 2.1 1.24 ± 0.75 170-387 28.7-62.3 25.5-45.3 9.4-17.6 0.43-2.30 {25) (24) (24) (19) (8) Juveniles 406.2 ± 67.3 46.4 ± 8.9 32.7 ± 5.3 9.1 ± 1.9 3.19 ± 1.51 265-620 27.8-74.0 23.1-65.0 4.9-14.7 1.30-8.35 (88) (86) (80) (77) (47) Subadiilts — 45.4 ± 7.1 30.1 ±3.4 7.9 ± 1.3 — 32.1-59.7 21.2-36.9 5.4-10.3 (34) (M) (26) Siibaduit females 453.5 ± 50.6 48.0 ± 8.9 30.9 ± 3.5 8.6 ± 1.2 4.66 ± 0.28 385-530 32.8-59.7 26.3-36.9 7.2-10.3 4.50-4.99 (10) (10) (9) (8) (3) Subadult males 525.6 ± 43.2 44.3 ± 6.1 29.7 ± 3.3 1.1 ± 1.4 7.69 ± 2.75 440-597 32.1-56.0 21.2-34.3 5.4-10.1 4.76-15.42 (24) (24) (22) (18) (17) Adults — AAA ± 8.9 30.0 ± 3.1 8.0 ± 1.2 — 20.0-72.1 19.3-35.7 4.8-11.9 (120) (108) (103) Adult females 468.8 ± 49.1 45.0 ± 9.6^ 30.0 ± 3.2 8.1 ± 1.1 5.34 ± 1.34 370-580 27.8-72.1 19.3-35.5 5.2-10.3 3.00-9.98 (72) (72) (65) (61) (33) Adult males 531.8 ± 55.2 43.5 ± 7.9^ 29.9 ± 3.1 7.9 ± 1.3 7.10 ± 2.33 410-660 20.0-62.0 21.6-35.7 4.8-11.9 4.01-14.06 (4H) (4H) (43) (42) (25) ' Mean ± SD, extremes, and sample si/e (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; adults, all permanent teeth completely erupted. ' These data are uncontrolled for seasonal weight variation, which has been reported both in females and in males in natural populations (Lindburg, 1977b, p. 247; Pearl et al., 1987, p. 36; cf. Small, 1981, p. 93; Zeng, 1992, p. 22). ^ These values differ slightly from those published previously (Fooden, 1997, p. 224) because of inclusion here of data for specimens with intermediate relative tail length (607f-75%) and exclusion here of data for specimens that had been held in captivity prior to collection. in relative tail length in the western half of the geographic range also is relatively minor (Fig. 12, Table 5); from Pakistan (ca. 73°E) to Myanmar (ca. 95°E), mean relative tail length is 46.7 ± 7.0% (SD; extremes, 31.9%-62.0%; n = 68). East of Myanmar. in the northeastern part of the range (northern Vietnam, China), relative tail length tends to decline from west to east, reaching ca. 30% at 120°E; six specimens collected on four islands in the South China Sea are included in this cline. Conversely, relative tail length increases in the extreme southeastern part of the range; in Thailand and Laos (ca. 100°E), mean relative tail length is 53.0% ± 4.9% (extremes, 47.5%- 60.8%; n = 8), and in the two aberrant specimens collected in central Vietnam, relative tail length is 64.5% and 72.1%. High relative tail length in these southeastern specimens may be interpreted as further evidence of hybridization between M. mulatto and M. fascicularis (see "Tail Length," p. 29). Judging from dry-skin measurements and a published illustration (Table 6; Milne-Edwards, [1870], pi. 32; Pocock, 1932, p. 550), relative tail length apparently was low (?30% in adults) in the now-extinct population that formerly inhabited Xinglong Xian (= Eastern Tombs; ca. 40°24'N, 1 17°30'E), northeastern China (cf. Zhang Yongzu et al., 1989, p. 380). This would be in accord with the pattern of longitudinal variation in relative tail length indicated above (cf. Fig. 1 1 ). Body WI'IGHT — Body weight data are available for 33 wild-collected adult females and 25 wild- collected adult males (Table 7). Weight, like head and body length, tends to increase with latitude (Fig. 13); at 30° to 35°N, mean weight (females, 7.45 kg, n = 5; males, 12.48 kg, n = 2) is ap- proximately twice that at 15° to 20°N (females. 30 FIELDIANA: ZOOLOGY 8 5 C CJ O ::= 7 3 -a ■C N x: '~ « c S -a 53 O .s o _ o E '^^ E - w o •£ B oil" c 5 >^ z. ■^ ou C3 V, ■" a. .2 « C3 4j u H-o U o o ^ — sO ^ Ol "^ fN) >r~, I — +1 I — +1 I —r.-i I — ■rr r) IT) r^j Tt — r<~i On t^ r<^ ir-, o ON '^t O IT) 00 — . iri NO -^ ON /-^ C ■* <^J ir, -^ (N ro n 00 O ir-, o r<-i — — ri n — n '^ ^ in f^ lO On ■^ no +1 I ~- +1 I — +1 I ^ in — oj o — O 00 c-j ^ t^ Tf >0 C^, -—V (^ r~t ^ ^ '^ 1/-, n ^ iy~i IT) vH "* vT IT) +1 O o in 'T ^-v Tt I^ ^-v «s O Vi fN ir> \}- r^ ^ 1 +1 1 ^ V) 1/-! O O ^ -* 00 c« IT) m ^ o <^. NO <^J '-. '-v in "^ o "-I + 1 I — r-> ^ o o o 2 Tl- -xT ^ in ^ + 1 I -^ o ON NO ON rt 00 ■^ fN in •—< Tt o « T3 O x> c 03 T3 OJ CD I 400- Males CB « .«. •♦-" □ -o h CB ■»: U VX M- • X A « D O )l^ 5K DD 550- 500- 450- Females i tt CB HD n + □« 4=1 D .u.i. ^ D^ U ^ D n X : 400- 350 ^ X; ■^- + • i /^. ^ )K )K 10 15 20 Latitude (°N) O Bangladesh n China x India )K Thailand l=t Vietnam 25 — r 30 + Myanmar A Nepal 35 V Pakistan Fig 8 Latitudinal variation in head and body length in Maccica mulatui adult noncaptives: data points for insular specimens are indicated by two-letter abbreviations (CB = Cat Ba; DD = Dangan Dao: HD = Hainan Dao; ND - Neilingding Dao). In this graph, the measurement for one Laotian female (395 mm) is included with Thai data; measurements of head and body length are not available for Bangladeshi adults. 32 FIELDIANA: ZOOLOGY ^.5 "a , 3 -o 00 N T « ■- ^ ~" OJ OJ — oij E •73 ^ >o O o % c 2 .^ •o c \^ C3 E ^ c« 2 E _a P "S >< t aj ' OJ c •a c ^.^ o E o § c« ^»^ ^ o M ^ C 1) c C3 — . JS ■c > ■^ o _a) IE D. Q. 2 E Of) o u ?J a U4 jr ^ ^' ^^ -J 9 U -J 5°- 00 — ^ ri (^) + 1 1 ^rj I — Ov — r4 oo >ri oo O ri r<-, 00 ^ — r4 -- — Tf — (N fV) r-, oi — OI (N OI m NO O r<-) O ,,— V r<~, 'N-l + 1 1 On ON OJ NO OI o OJ 1 1 i/~. O FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 33 300- 250- 200- 150- E E I- X X -1- V X X X D a + t=t X /CB X X X X X a X X V D D DD / a D Males CB / D 300- 250- 200- 150- 100- Females ^ )^^^- ^ « HD H ^ X 1- «x 4- / CB -^■ "H H □ d ND / ^ X a >< ■4 D 10 15 20 25 Latitude (°N) O Bangladesh n China x India -f- Myanmar )K Thailand ^ Vietnam 30 35 A Nepal v Pakistan Fig. 9. Latitudinal variation in tail length in Macaca mulatto adult noncaptives; data points for insular specimens are indicated by two-letter abbreviations (CB = Cat Ba; DD = Dangan Dao; HD = Hainan Dao; ND = Neilingding Dao). In this graph, the measurement for one Laotian female (240 mm) is included with Thai data; measurements of tail length are not available for Bangladeshi adults. 34 FIELDIANA: ZOOLOGY 300- 250- 200- 150- - 300 H 03 250- X i V > X X X X : A ■j- ■1- > V X X X X -f- ^ X X ^ CB ° *^ Q D D DD / D Males CB / D 200- 150- 100- 70 Females 80 xH- -i-x )|^ + +=«= + )i( ° tt tt- a tt DO tt ^D HD I* CB E3- ND 110 120 90 100 Longitude (°E) O Bangladesh n China x India -i- Myanmar b. Nepal v Pakistan )K Thailand ^ Vietnam Fig. 10. Longitudinal variation in tail length in Macaca mulatta adult noncaptives. For detailed comments, see Figure 9. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 35 20 25 Latitude (°N) O Bangladesh d China x India -t- Myanmar A Nepal )K Thailand tt Vietnam Fig. 1 1 . Latitudinal variation in relative tail length (tail length/head and body length) in Macaca mulatta adult noncaptives; data points for insular specimens are indicated by two-letter abbreviations (CB = Cat Ba; DD = Dangan Dao; HD = Hainan Dao; ND = Neilingding Dao). In this graph, the value for one Laotian female (0.61) is included with Thai data; values are not available for Bangladeshi adults. 4.27 kg, n = 4; males, 6.14 kg, n = 3). Data points for specimens collected on four shallow- water islands (Cat Ba, Dangan, Hainan, Neiling- ding) in the South China Sea fit well within the latitudinal body weight cline. Weights previously reported for M. mulatta in Hainan Dao (18°23'N, 110°00'E) and northern Pakistan (34°03'N, 73°22'E) are similar to those of specimens examined from the same latitudes (Fig. 13). For Hainan Dao specimens of unknown maturity, Jiang Haisheng et al. (1991, p. 210) re- ported that mean weight was 3.88 ± 0.20 kg in 33 females and 5.08 ± 0.72 kg in 16 males. In northern Pakistan, Pearl et al. (1987, p. 36) re- ported that mean weight of adult females was 7.3 kg and that the weight of a large male was 11.9 kg. Body weight in M. mulatta captives imported from China has been compared with body weight in M. mulatta captives descended from monkeys imported from India (Clarke & O'Neil. 1999, pp. 340, 341; see "Head and Body Length," p. 26). In males, body weight in Chinese-origin adults exceeded that in Indian-derived adults; in females, body weight in Chinese-origin adults was less than that in Indian-derived adults. As previously indicated, no information is available concerning the region of origin of these monkeys within India or China. 36 FIELDIANA: ZOOLOGY — -a w c O o C TD "^ •£ fc Uh «J 5 5j OJj Q § - c n. if ^ •t: oj 4J C- U 1 1 s +1 1 "^ + 1 1 (N o ^ , 00 :> rr-, vO r) 1^ r^l *0 00 I fN o6 00 r<-] t^ i/-i o + 1 1 5+1 1 ::+! 1 ^ w O 00 r- m ~-> en ^^ in ^ ^ •^ 1/-1 \o r-; +1 I '^ +1 I Ov ro T+ 00 ro -J 5 w +1 I Tt (^ ^ +1 I "O FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MAC AC A MULATTA 37 c CD i5 cc 0.8- 0.7- 0.6- 0.5- 0.4- 0.3- 0.2- 0.1 Both sexes 70 x^"^ V % .^ : ■X- A n « ^X... ( ^x -.f- — -i)!^""^" +• ^ • X .^^ DJa "*" + ; ^ ! ^ + + + a tn° « ■=■ lb a « pi 4-1 ™ « ^ D : □ : "«-e r- CB ■Pi- CB i DD ND 80 110 120 90 100 Longitude (°E) O Bangladesh n China x India -t- Myanmar A Nepal v Pakistan ^ Thailand t=t Vietnam Fig. 12. Longitudinal variation in relative tail length (tail length/head and body length) in Macaco mulatto adult noncaptives. For detailed comments, see Figure 11. Cranial Characters Sex and Age Variation In wild-collected adult specimens of M. mulatta (Figs. 14, 15, Table 8), greatest length of skull (excluding incisors) in 80 males (121.8 ± 8.3 mm) averages about 13% greater than in 120 fe- males (107.8 ± 7.1), and rostral-postrostral ratio in 69 males (50.5 ± 3.0%) averages about 15% greater than in 1 1 1 females (44.1 ± 3.7%) (cf. Cochard. 1985, p. 237; Cheverud & Richtsmeier. 1986. p. 392; Mouri, 1995. p. 189). Relative zy- gomatic breadth (ZB/GL) in 79 males (70.7 ± 2.6%) averages only 2% greater than in 118 fe- males (69.0 ± 2.2%). From infancy to adulthood, rostral length in- creases much faster than postrostral length, where- as zygomatic breadth increases only slightly faster than greatest skull length (Table 8). In males, ros- tral-postrostral ratio in adults is more than 100% greater than in infants (cf. Bhatia, 1978. p. 66), and relative zygomatic breadth in adults is about 10% greater than in infants. Fluctuating asymmetry of the skull and teeth in M. mulatta tends to increase ontogenetically to age 6 years (Halgrimsson. 1999, p. 139). An intensive study of maxillomandibular growth in captive M. mulatta has been published by Schneiderman (1993, p. 75), and craniofacial growth in laboratory-colony captives has been compared with that in free-ranging captives by King and Schneiderman (1991, p. 105). Dental emergence norms have been carefully studied in the Yale University laboratory colony 38 FIELDIANA: ZOOLOGY Table 6. Dry-skin measurements of 14 Macacti mulatta specimens collected at Xinglong Xian (= Eastern Tombs), Hebei Province, northeastern China. Relative Head and body Tail length tail length Museum No. Sex Age length (mm) (mm) (T/HB; %) AMNH 57038 M Infant 280 95 33.9 57039 M Adult >390 135 <34.6 57040 M Juvenile 385 140 36.4 57042 F Juvenile 440 160 36.4 57043 ? Infant 380 130 34.2 57108 ? Infant 300 135 45.0 57110 ? Infant 300 110 36.7 BM(NH) 1931.1.7.2' M Adult 730 140 19.2 FMNH 39376 M Juvenile 420 130 31.0 39377 ? Juvenile 360 160 44.4 39378 ? Juvenile 290 120 41.4 MNHN 335- F Juvenile 545 125 22.9 USNM 240704 F Subadult 420 140 33.3 240705 F Juvenile 280 140 50.0 ' Captive. - Cr. Milne -Edwards. [1872]. p. 228. of A/, mulatta (Fig. 16); in this study, emergence was defined as initial penetration of the gingiva by each tooth. In the Yale colony, deciduous teeth apparently emerged in four major waves. Teeth in the first wave (i,. i'. i.) emerged at median age ca. 0.05 year (18 days), those in the second wave (i-) at age ca. 0.1 year (36 days), those in the third wave (C|, c', m,, m') at age ca. 0.2 year (73 days), and those in the fourth wave (m^, m-) at age ca. 0.4 year (156 days). Following a diapause of ap- proximately 1 year, permanent teeth also appar- ently emerged in four major waves. Teeth in the first wave of permanent teeth (M,, M') emerged at median age ca. 1.4 years, those in the second wave (I|, I', L, I-) at age ca. 2.5 years, those in the third wave (M., M-, P„ P\ C,, C\ P4, P^) at age ca. 3.5 years, and those in the fourth wave (M„ M') at age ca. 5.7 years. Mandibular teeth, particularly those in the permanent set, usually emerged slightly earlier than their maxillary coun- terparts. Ages of dental emergence in females were generally similar to those in males, except for C| and C, which emerged ca. 0.6 year earlier in females (3.13 years, 3.46 years) than in males (3.84 years. 4.04 years), and M, and M\ which emerged ca. 0.6 year earlier in males (5.30 years, 5.40 years) than in females (5.74 years, 6.23 years). The elapsed time between initial emer- gence of a tooth and its complete eruption to full height apparently is a few months for most teeth but probably is ca. 2 years for the permanent ca- nines of males (Cheverud. 1981, pp. 158, 163). In the free-ranging colony of M. mulatta introduced in Cayo Santiago, the second and third molars re- portedly emerged as much as 1 year later than in some laboratory colonies (Tumquist & Kessler, 1990. p. 309); this suggests that dental emergence in natural populations of M. mulatta may also be retarded relative to that in laboratory colonies. Geographic Variation Skull Length — Greatest length of skull pro- vides the most comprehensive and reliable indi- cation of geographic size variation in M. mulatta (Table 9). This measurement is available for 170 well-localized adult specimens — 104 females and 66 males — collected at 126 localities. Greatest length of skull in both sexes generally increases with increasing latitude (Fig. 17, Table 9). South of 20°N, mean greatest length is 101.1 ± 3.32 mm in 18 adult females and 113.7 ± 3.50 mm in 10 adult males; north of 30°N, mean great- est length is 119.0 ± 4.42 mm in nine adult fe- males and 131.2 ± 6.02 mm in 12 adult males (cf. Albrecht, 1978, p. 129; Gelvin & Albrecht, 1996, p. 1 1 1). In 13 adult specimens collected on five islands in the South China Sea, greatest length is similar to that in continental specimens collected at the same latitude (Fig. 17). In the sin- gle adult skull (AMNH 57039. male) available from northeastern China, at the northeastern limit of the specific range, greatest length is surprisingly small FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 39 .^ CJ ■o ■yj c OJ ^ ^ o o r3 s D. (U C 13 3 •a CJ CD N C O I 2 OJ •a ^.^ 3 (U N i2 'c^ 1) _o jj D. oij e n > « O " .S S c H o =* ca ^ u 4J 03 N > CJ a; a. D. « 2 c* •^ Q OJ u o o J= ^ t~-^ ^' LU c _1 o CO ■^ < C3 H ■> u P "^ — 9 ^ iji <^i uS oo 00 •^^ ^.00 ^ ^ +1 1 ir 2q + 1 I in oo ^ u-5 -J 5^ +1 •3 r- 00 r-^ in 'N VD _L, (N ^ d in ^^ O ^ ^ o m O 40 FIELDIANA: ZOOLOGY 15 Males « ^ ■m- H n a Dn/DD ■" i o -q>- 20 25 Latitude (°N) O Bangladesh a China x India )K Thailand 1=1 Vietnam 30 Females a ■ 1 ° H D V HD S"^ x-t- \ a D X X": D )K D D ■ ' i ' i 35 -(- Myanmar A Nepal v Pakistan Fig. 13. Latitudinal variation in body weight in Macaca mulatta adult noncaptives; data points for insular spec- imens are indicated by two-letter abbreviations (CB = Cat Ba; DD = Dangan Dao; HD = Hainan Dao; ND = Neilingding Dao). Weight data are not available for Bangladeshi and Nepalese adults. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 41 Fig. 14. Skull of adult female Macaca mulatta—FMNH 99668, Thailand: Ban Mae Lamao. (Photographs by John Weinstein, the Field Museum, negative Nos. Z 94270.1-4.) (123.8 mm), considering the latitude (40°24'N) of collection of this specimen (Fig. 17); in a subadult female (usnm 240704) collected in the same area, greatest length also is small (100.6 mm). Longitudinal variation in greatest length of skull is relatively minor (Fig. 18). For example, mean greatest length in a sample of adults col- lected between 30°N and 35°N in Pakistan and India (females, 120.9 ± 3.26 mm, n = 5; males, 129.3 ± 3.23 mm, n = 8) is similar to that in a sample collected 2,500 km to the east, across the Xizang-Qinghai (Tibetan) Plateau, at the same lat- itude in China (females. 116.7 ± 4.98 mm, n = 4; males, 136.8 mm [132.6-141.0 mm], n = 2). Cranial and Dental Morphology — Variation in suites of cranial and dental measurements in >150 specimens that were collected in various sample areas in China and in six specimens that originated in India (localities unspecified) has been studied by two groups of Chinese investi- 42 FIELDIANA: ZOOLOGY Fig. 15. Skull of adult male Macaca miilalta — FMNH 99669. Thailand: Huai Ap Nang. (Photographs by John Weinstein, the Field Museum, negative Nos. Z 94271.1-4.) FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 43 Table 8. Cranial measurcmenls and proportions in age/sex classes of wild-collected Macaca midatta. Relative zygomatic Rostral-postrostral Age/sex Greatest breadth Postrostral ratio class' length (mm) (ZB/GL X 100) length (mm) (R/PR X 100) Infants 78.6 ± 6.8 64.3 ± 2.6 67.3 ± 4.6 24.2 ± 3.1 65.8-93.6 59.1-69.9 59.7-79.2 19.8-34.3 {37) (37) (33) (33) Juveniles 97.8 ± 10.0 61.3 ± 2.2 76.4 ± 5.4 36.1 ± 6.0 79.1-126.7 61.8-73.5 60.9-91.5 26.8-53.6 048) (146) (133) (132) Subadult females 101.0 ± 4.3 68.1 ± 1.8 77.8 ± 3.0 39.6 ± 2.8 95.0-112.0 64.7-71.6 72.1-85.0 34.6-46.8 (20) (20) (19) (19) Subadult males 118.4 ± 7.5 68.9 ± 2.6 85.8 ± 4.7 49.7 ± 3.0 105.9-136.8 64.1-73.7 78.8-96.3 43.6-56.4 (36) (36) (33) (32) Adult females 107.8 ± 7.1 69.0 ± 2.2 80.7 ± 4.5 44.1 ± 3.7 92.9-126.5 60.1-73.9 71.3-92.6 37.6-59.4 (120) (118) (111) (111) Adult males 121.8 ± 8.3 70.7 ± 2.6 86.4 ± 4.2 50.5 ± 3.0 107.1-143.1 65.0-76.1 77.8-96.9 41.3-58.2 (80) (78) (69) (69) Note: Mean ± SD, extremes, and sample size (italicized figures in parentheses). For definition of cranial measure- ments, see Fooden (1969, p. 40). ' Dental specifications: infants, deciduous teeth only; juveniles, some permanent teeth erupted; subadults, third molars in females or canines in males incompletely erupted; adults, all permanent teeth completely erupted. gators (Jiang Xuelong et al., 1991, p. 242; 1995, p. 44; Pan et al., 1992, p. 40; Peng et al., 1993, p. 2; Yao et al., 1995, p. 113; Yu et al., 1996, p. 152; cf. Sikorska-Piwowska, 1970, p. 9). Al- though distiibutions of cranial and dental mea- surements overlap in these samples, statistically significant differences were discovered among the Chinese samples and between the Chinese and the Indian samples. The two groups of investigators disagree concerning the morphological relation- ships of samples of M. mulatta collected in south- central China; one group found its south-central Chinese sample to be most similar to a south- eastern Chinese sample (Jiang Xuelong et al., 1995, p. 46), whereas the other group found its south-central Chinese sample to be most similar to a southwestern Chinese sample (Yu et al., 1996, p. 153). Mandibular measurements in specimens of M. mulatta collected in three regions of China have been analyzed by Zhao et al. (1999. p. 63). Comparison with Macaca fascicularis Skull length in both sexes averages greater in M. mulatta than in its southern relative. M. fas- cicularis (cf. Fooden. 1995, p. 38). Conversely, the rostrum in M. mulatta protrudes less than in M. fascicularis (cf. Mouri, 1996, p. 296; Pan et al., 1998, p. 525). Width of the braincase and ros- trum appear to be greater in M. mulatta than in M. fascicularis. A median sagittal crest, formed by ontogenetic convergence of the temporal lines, is rare in M. mulatta adult males (well-defined crest in three of 7 1 specimens examined, incipient crest in one specimen), whereas a sagittal crest is common in M. fascicularis adult males. Molecular Biology and Genetics Mitochondrial DNA Data concerning the geographic variation in mi- tochondrial DNA (mtDNA) in M. mulatta have been published by three research groups: Haya- saka et al. (1988, p. 271; 1996, p. 1044), Melnick et al. (1993, p. 284; cf. Morales & Melnick, 1998, p. 7). and Zhang and Shi (1989, p. 334; 1993a, p. 8; 1993b, p. 591). In addition, Disotell et al. (1992, p. 6) have published the nucleotide se- 44 FIELDIANA: ZOOLOGY M3 C, p3 P3 M2 1^ nig m, 2 TTT Dental Emergence Age Median, 2nd Percentile, 98th Percentile O Female n Male °-^1? 000 O .D— rB o D- D- 00 -B — D -B- ^ C? O -B- ^ oip' 'd =^#5 ■^^ ^ # t ^|=«> ^ O X I I I I I " IS I I I 0.05 (18 d) 0.1 (36 d) 1.0 0.5 (182 d) Age (yr), log scale 5.0 M3 P^ P4 c, p3 P3 M^ Ms 1^ >2 r I1 M^ M, m2 10.0 Fig. 16. Dental emergence chronology in Macaca mulatto, laboratory colony .sample (Hurme & van Wagenen, 1953, pp. 297, 299; 1961, pp. 1 11-112, 128; Hurme, 1960, pp. 796-797; cf. Maity & Rathore, 1998, p. 250): median, 2nd percentile, and 98th percentile values are indicated for age at initial penetration of gingiva by deciduous and permanent teeth in females and males. Abbreviations: i/I = incisor, c/C = canine, P = premolar, m/M = molar; lowercase letters indicate deciduous teeth, uppercase letters indicate permanent teeth; subscripts indicate mandibular teeth, superscripts indicate maxillary teeth. Arrowheads (lower left in graph) indicate six off-scale 2nd percentile values: i,, females and males, 0 years (i.e., tooth already erupted at birth); i' and i,, males, 0 years; i' and i,, females. 0.0082 years. Sample sizes: i,, i', i,, i-— females (n = 53), males (44); c,, c'— females (51 ), males (41-42): m,, m'— females (50), males. (41); m., m-— females (4.3-44), males (32-33): M,, M'— females (42), males (30); I,, I', L, F— females (41-42), males (22-25); M,, M-— females (39-40), males (18-19); P„ P\ C,, C, P„ P^— females (35-39), males (13-17); M„ M'— females (30-31), males (10-12). For additional dental emergence age data collected by various procedures, see Schultz, 1935, p. 499; Eckstein, 1949, p. 367; Gavan, 1967, p. 985; McNamara et al., 1977, p. 701; Trotter et al., 1977, p. Ill; Cheverud, 1981, p. 163: Zeng et al., 1984, p. 83; Sharma & Lai, 1986, p. 145; Turnquist & Kessler, 1990a, p. 309; 1990b, p. 239; Zeng, 1992, pp. 20, 22; Smith et al., 1994, pp. 215, 226. quence of the mitochondrial COII gene in one M. mulatta individual from an unspecified locality. Hayasaka et al. (1988, p. 271) used 17 endo- nucleases to study mtDNA restriction sites in one M. mulatta specimen of Indian origin (no further locality information available) and subsequently (1996, p. 1046) determined the nucleotide se- quence of an 896-bp region of mtDNA in this FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 45 — c. 3 -a 3. I •- "P ^ .£ ■y. i-J n o ^s Cij c o ~ c o < ■''• .i 4» — U-) <^ ir, — fV" __ Qv I/-, _ (N) + 1 I -- +1 1—3 I -- ^ 3; — r^ — 3; r- _ ir, rj ^ — — — — \o — -t +1 I — +1 I — D ^^ r<-, r-l ^^ ir, — ^^ ^^ O — IT) ; oo +1 90 P OC ^ r^, ■*^ — -^ — '^ + 1 1 — +1 1 — vD vO [^ O 00 O a\ r^i 1-^ — ir-j r^, — . -■ y. — • ■r c d 2 c u .5 ^ ~" c OJ ^ y Qj 3-a ^ 46 FIELDIANA: ZOOLOGY 150 140- 130- 120- 110- Males ! b V ■ X i D ? ^ • CB- yc d xi ^ D ■ n-r-yx t, ^ a^^X i ^ X; CB ^DD i ^"^ i X"n + i X A ; « O : ; Z ISO- CO CO o 120- 110- 1 GO- GO- Females 8 HD HD ''n % Cfe ■■HD^J" HD tt i....D. « « D xP VC ^D XD S" a Dv ■^ .-Eij- ^ X- B< X a^^xA f^Q X ; . o- ■fi-x 10 15 20 25 Latitude 30 35 40 I 45 'N) O Bangladesh d China x India "^ Thailand tt Vietnam ■t- Myanmar A Nepal v Pakistan Fig. 17. Latitudinal variation in greatest length of skull in Macaca midatta adult noncaptives. Data points for insular specimens are indicated by two-letter abbreviations (CB = Cat Ba; DD = Dangan Dao; HD = Hainan Dao; SD = Shan2chuan Dao; VC = Van Canh). Indian specimen and in two M. mulatto specimens of unknown country of origin; the mtDNA data of these three M. mulatto specimens were com- pared with data derived from M. cyclopis (Taiwan; n = 1), M. fiiscoto (Japan; n = 3). and other ma- caque species. Although the nucleotide sequences of the two M. mulatto specimens of unknown country of origin differ from each other at only FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 47 \ •^--^-x- Males ^)K □ « ^^-^^VC « ^ tt tt il- CBs^« .,'HD D •HD /DD r >^ X X X Females X X ftp- X .Nfc 4: D « □ 1=1 ^ VC i HD tt a-HD HD ' 1 ' \ ' \ ^ 1 ' 70 80 90 100 110 120 Longitude (°E) O Bangladesh a China x India -t- Myanmar A Nepal V Pakistan ^ Thailand U Vietnam Fig. 18. Longitudinal variation in greatest length of skull in Macaco miilatta adult noncaptives. For key to abbreviations, see Figure 17. 0.2% of the sites, their sequences differ from that of the Indian specimen at ca. 6.2% of the sites; this intraspecific sequence divergence greatly ex- ceeds the interspecific divergence between the In- dian M. miilatta specimen and M. cyclopis (3.2%) and between the Indian M. mulatto specimen and M. fuscata (ca. 3.6%). Sequence divergence be- tween the two M. mulatto specimens of unknown 48 FIELDIANA: ZOOLOGY country of origin and M. cyclopis and M. fiiscata is ca. 5.1% and 5.8%, respectively. Hayasaka et al. (1996, p. 1052) suggest that the taxonomically incongruous intraspecific and interspecific diver- gences in their M. mulatta, M. cyclopis, and M. fiiscata sequence data may be the result of either retention of ancestral polymorphism or interspe- cific hybridization. Melnick et al. (1993, p. 284) used 15 endonu- cleases to study mtDNA restriction sites in 18 in- dividuals representing five M. miilatta popula- tions— one each from Pakistan (n = 3), northern India (n = 4), and Myanmar (n = 4) and two from southern China (n = 5, n = 2); the exact prove- nance of these five samples and the number of localities represented by each are unspecified (cf. Melnick et al., 1984, p. 342; Morin et al., 1997, p. 201). Of the 10 haplotypes that were identified in this study, one is unique to the smaller Chinese sample, and three sets of three haplotypes each are unique to the Indian, Myanmar, and larger Chinese samples, respectively; the single haplo- type common to the three individuals in the Pak- istani sample is the same as that of one individual in the Indian sample. Distance-based and charac- ter-based trees reveal similar patterns of relation- ships among the M. mulatta samples studied. In both kinds of trees, the primary divergence is be- tween the Pakistani/Indian (western) samples and the Myanmar/Chinese (eastern) samples (diver- gence = 3.9% ± 0.45%); on the basis of the mag- nitude of this divergence, Melnick et al. (1993, p. 287) suggest that western and eastern populations of M. mulatta formerly were separated by a major barrier. Divergences among the seven individuals in the Pakistani/Indian samples are very small (d = 0.4% ± 0.31%). The divergence between the Myanmar sample and the two Chinese samples (d = 2.3% ± 0.58%) is approximately twice as great as that between the two Chinese samples (d = 1.2% ± 0.15%). As a supplement to their investigation, Melnick et al. (1993, pp. 283, 286) compared their data with Hayasaka et al.'s (1988, p. 271) data for one Indian specimen of M. mulatta, one specimen of M. cyclopis, and three specimens of M. fuscata (see above). The haplotype of Hayasaka et al.'s Indian M. mulatta specimen is most divergent from haplotypes of Melnick et al.'s Chinese sam- ples (d = 3.3% ± 0.16%), next most divergent from the Pakistani/Indian samples (d = 3.0% ± 0.25%), and least divergent from the Myanmar sample (d = 2.1% ± 0.12%); this suggests that Hayasaka et al.'s Indian M. mulatta specimen may have originated in eastern India, nearer to Myan- mar than to Pakistan. Distance- and character- based trees indicate that haplotype similarity among eastern M. mulatta, M. cyclopis, and M. fuscata is greater than haplotype similarity be- tween eastern and western M. mulatta; this dis- crepancy between the gene tree revealed by mtDNA and the species tree revealed by mor- phology and allozymes (see "Blood Proteins," p. 52) parallels the taxonomically incongruous find- ings reported by Hayasaka et al. (see above). Mel- nick et al. (1993, p. 290; cf. Hoelzer, 1997, p. 624) interpret the pattern of mtDNA haplotype rela- tionships as a retention of ancestral mtDNA sim- ilarity by eastern M. mulatta, M. cyclopis, and M. fuscata. In a geographically detailed study, Zhang and Shi (1993b, p. 591) used 20 endonucleases to study mtDNA haplotypes in 36 M. mulatta indi- viduals collected at 23 localities in China, Myan- mar, and Vietnam (Fig. 19); their analysis also includes Hayasaka et al.'s haplotype data for the Indian M. mulatta specimen cited above. Restric- tion fragment length analysis revealed that each of the 24 localities sampled by Zhang and Shi is characterized by a distinctive mtDNA haplotype; at each of the nine localities represented by more than one individual, haplotypes were uniform in all individuals sampled (n = 2-4). Two trees — one based on the unweighted pair group (UPG) method and the other based on the neighbor-join- ing (NJ) method (cf. Melnick et al., 1992, p. 196) — were constructed by Zhang and Shi to in- vestigate the pattern of resemblance among the 24 local haplotypes. Although Zhang and Shi favored the UPG tree because of its general congruence with one of several previously proposed subspe- cific classifications of M. mulatta (Jiang Xuelong et al., 1991, p. 242; cf. 1995, p. 44), a consensus tree of stable clusters common to both the UPG tree and the NJ tree provides an independent es- timate of geographic variation in mtDNA haplo- types that is not bia.sed by taxonomic preconcep- tions (Fig. 20). As indicated by the consensus tree, the largest mtDNA haplotype divergence is between Zhang and Shi's insular Hainan M. mulatta sample (20, n = 2) and 20 mainland locality samples (d = 3.5% ± 0.49%); the positions of three mainland locality samples (E Guangxi, 6; Henan, 72; Fu- jian, 15) are unresolved relative to this dichotomy. The second-largest mtDNA haplotype divergence is between Hayasaka et al.'s Indian sample (24, n = 1) and the remaining 19 locality samples (d = FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 49 35° Mitochondrial DNA sample areas 30° zsfu 20° 13 15 105' 110° 115° 120° Fig. 19. Sample areas cited in mitochondrial DNA study of Zhang and Shi (1993b, p. 590); sample area no. 24 ("India") is not mapped. For key to sample area numbers, see Figure 20. 2.8% ± 0.35%). The third-largest divergence is between the northwestern Sichuan sample (7, n = 3) and 17 locality samples from Myanmar, China, and Vietnam (d = 1.6% ± 0.29%); the position of one Chinese locality sample (northwestern Yunnan, 19) is unresolved relative to this dichot- omy. The fourth-largest divergence is between the eastern Myanmar sample (23, n = 2) and the re- maining 16 locality samples (d = 1.3% ± 0.23%). Although the branching pattern of the residual 16 locality samples is not completely resolved, 15 of these samples are positioned within three stable clusters (Fig. 18) as follows: 1. Southwestern Yunnan (22, n = 2); west-cen- tral Yunnan (4, n = 1); southeastern Hubei (77, n = 1). Within this cluster, the close resemblance between the west-central Yunnan sample and the southeastern Hubei sample is particularly note- worthy; although these two localities are separat- ed by ca. 1,100 km, the haplotype divergence be- tween the samples is only 0.17%. 2. Central Yunnan (27, n = 1); northeastern Yunnan: Yongshan (8, n = 1); northeastern Yun- nan: Yillang (9, n = 1); southern Sichuan (14, n = 2); Hunan (5, n = 1). This and the preceding cluster overlap geographically (Fig. 17). 3. West-central Sichuan (7, n = 1); eastern Sichuan (2, n = 2); Anhui (13, n = 1); Guizhou (3, n = 3); northern Guangxi (76. n = 1); northern Vietnam (10. n = 1); northern Vietnam (77, n = 1). On the basis of available information (see above), the following hypotheses may be pro- posed concerning geographic variation in mtDNA in M. miilatta: 1 . Haplotypes usually are uniform in each local population of M. mulatta. 2. Haplotypes in each local population of M. mulatta usually differ from those in other local populations. 3. Intraspecific haplotype variation may exceed interspecific haplotype variation. 50 FIELDIANA: ZOOLOGY rC h: € rC rC India (24), 1 NW Sichuan (1), 3 NW Yunnan (19), 2 E Myanmar (23), 2 W Cent. Sichuan (7), 1 E Sichuan (2), 2 Guizhou: Dushan (3), 3 N Guangxi: Jinchengjiang (16), 1 Anhui: Huangshan (13), 1 N Vietnam (10), 1 N Vietnam (1 1), 1 NE Hubei: Zhushan (18), 1 SE Hubei: Enshi (17), 1 W Cent. Yunnan: Baoshan (4), 1 SW Yunnan: Simao (22), 2 NE Yunnan: Yiliang (9), 1 NE Yunnan: Yongshan (8), 1 S Sichuan: Muli (14), 2 Cent. Yunnan: Chuxiong (21), 1 Hunan: Jishou (5), 1 Henan: Huixian (12), 1 CFujian (15), 4 E Guangxi: Wuzhou (6), 1 Hainan (20), 2 Fig. 20. Consensus dendrogram of mitochondrial DNA relationships in Macaca miliaria samples studied by Zhang and Shi (1993b, p. 597). In each line, the italicized number in parentheses is the sample area number (see Fig. 19), and the number following the comma is the sample size. 4. Haplotypes in eastern populations of M. mii- latta (Myanmar. China, Vietnam) are more diver- gent from those in western populations of M. mu- latta (Pakistan, northern India) than they are from those in M. cyclopis (Taiwan) and M. fusccita (Ja- pan). To determine whether haplotype variation between eastern and western populations of M. mulatto is gradual or abrupt, data from geograph- ically intermediate populations would be required. 5. The known haplotype in the insular Hainan population of M. mulatto diverges strongly from haplotypes in eastern mainland populations of M. mulatto. 6. The known haplotype in the northwestern Sichuan population of M. mulatto diverges strong- ly from haplotypes in other mainland Chinese populations. Vietnamese populations, and eastern Myanmar populations. 7. Haplotypes in Myanmar populations of M. mulatto are divergent from those in Chinese and Vietnamese populations. Authors cited above differ in their interpreta- tion of the chronological significance of mtDNA haplotype variation. Hayasaka et al. (1996. p. 1052) and Zhang and Shi (1993b. p. 594) assume a constant rate of mtDNA nucleotide substitution FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 51 in Macaca, whereas Melnick et al. (1993, p. 290; cf. Melnick and Hoelzer, 1993, p. 6; Hoelzer et al., 1998, p. 29) present evidence that this rate may be viiriable. Nuclear DNA Using nine enzymes (Alu\, Bam\\\, EcoRl. Hindm, HinW Hpa\, Psth PvuW. and Xmn\), Crovella et al. (1994, p. 66) studied highly re- peated nuclear DNA restriction patterns in two M. miilatia captives. One captive, identified as M. m. mulatto, presumably originated in India, and the other, identified as M. m. lasiotus, presumably originated in China. The restriction patterns of these two captives were indistinguishable. M. mulatto is polymorphic for the chemokine receptor CXCR4 (a coreceptor for human immu- nodeficiency virus), but this polymorphism is not known to vary geographically (Chen et al., 1997, p. 2707; Pretet et al.. 1998, p. 639). Of 17 rhesus monkeys tested for CXCR4 alleles, allele 1 (ad- enine at nucleotide 641) was detected in two In- dian samples, and allele 2 (thymidine at nucleo- tide 641) was detected in six Indian samples, sev- en Chinese samples, and two samples of unknown geographic origin. Twenty M. mulatto captives — 10 of Chinese or- igin and 10 of Indian origin — were included in two studies of restriction fragment length poly- morphism at four loci in the p-globin gene cluster (Shimizu & Takenaka, 1991a, p. 178; 1991b, p. 191). Although M. mulatto is reported to be poly- morphic at all four loci, no data are available con- cerning the possibility of a relationship between polymorphism and country of origin in this spe- cies. Morin et al. (1997, p. 206; cf. Smith, 1994a, p. 205; Kanthaswamy & Smith, 1998, p. 141) report that mean gene diversity for 15 simple sequence repeat loci is greater in a Chinese M. mulatto sam- ple (0.78) than in Indian (0.66) and Thai (0.61) M. mulatto samples. Watanabe et al. (1997, p. 351) have compared the nucleotide sequence at the HPRT locus in one M. mulatto specimen of unspecified geographic origin with that of other macaque species and nonmacaque catarrhines. Blood Proteins Judging from available information, geographic variation in blood-protein allele frequencies in M. mulatto is relatively minor (Table 10; Fsj = 0.0253); the most variable locus is Tf (transfer- rin). Melnick (1988, p. 207; cf. Su et al., 1997, p. 112) studied 25 to 37 loci in samples of this spe- cies from Pakistan, India, China, and Thailand; the geographic span of these samples exceeds 3,000 km. For the loci studied, Melnick estimates that the total blood-protein gene diversity in M. mulatto is 0.0814; he allocates ca. 86.5% of this diversity to individual differences among mem- bers of the same troop, ca. 3.8% to differences among neighboring troops, ca. 1.0% to differenc- es among local populations within the same coun- try, and ca. 8.7% to differences among popula- tions in different countries. In another study, Mel- nick et al. (1986, p. 136) found that intercountry variation is weakly clinal. In this cline, the Pak- istani and Indian samples form one cluster and the Chinese and Thai samples form another; when a Bangladeshi sample was included in the analysis, it clustered with the Pakistani and Indian samples. Smith et al. (1987, p. 204). in a brief summary comment, indicate that Indian and Chinese sam- ples from unspecified localities approach fixation for opposite alleles at genetic loci for carbonic anhydrase II, properdin factor B, and albumin. Ding et al. (1998, p. 172) studied blood-protein variation in M. mulatto samples collected at six localities in western Yunnan, China. Samples from three localities north of 25°30'N tended to differ from those from three localities south of 25°30'N. Within the northern and southern groups of samples, blood-protein divergence was not re- lated to the distance between localities. Schmitt and Tomiuk (1995, p. 126) have shown that the distribution of blood-protein allele fre- quencies per locus in M. mulatto closely approx- imates that predicted by the neutral mutation hy- pothesis. Three recent studies have investigated the use of blood-protein data in monitoring and maintaining genetic variability in captive research colonies of this species (Gill et al., 1992, p. 89; Smith, 1994a, p. 204; Ely et al., 1994, p. 212). Karyology Although chromosomal polymorphism has been reported in M. mulatto (Sharma & Seth, 1984, p. 380; Small et al., 1985, p. 66), no infor- mation is available concerning possible geograph- ic variation in karyotype. The diploid chromo- some number in this species is 42. 52 FIELDIANA: ZOOLOGY Table 10. Frequencies (%) of major alleles at polymorphic blood-protein loci in samples of Macaco mttlalta from six countries (sample sizes indicated by italicized figures in parentheses). Because variability at the Tf locus is exceptionally large, frequency data are provided for six alleles at this locus. For key to locus abbreviations, additional frequency details, and references, see Fooden and Lanyon (1989. p. 214). Locus Allele Pakistan India Bangladesh China Thailand Vietnam Acp A 100 (279) 100 (214) (0) 98 (76) 100 (31) — (0) ADA 2 97 (216) 99 (214) — (0) 92 (76) 94 (31) — (0) Alb A 98 {32} 40 (214) — (0) 98 (76) 95 (31) — (0) CA-I A 100 (2 J 9) 100 (238) 100 (26) 96 (76) 85 (46) — (0) CA-II B 71 {186) — (0) — (0) — (0) 40 (5) — (0) Dia C 67 {32) 77 (214) — (0) 81 (76) 46 (31) — (0) IDH 2 100 {219) 99 (214) — (0) 83 (76) 75 (31) — (0) PGD A 99 (219) 84 (238) 84 (25) 97 (76) 99 (46) — (0) PGM-1 1 100 (219) 100 (214) — (0) 97 (76) 96 (31) — (0) PGM-II 1 100 (219) 100 (214) — (0) 97 (76) 100 (31) — (0) PHI 1 92 (216) 93 (214) — (0) 96 (76) 96 {31) — (0) PI C 100 (32) 100 (314) 94 (25) 100 (76) 99 (46) — (0) TBPA F 83 (32) 88 (342) 88 (39) 80 (76) 83 (44) — (0) Tf C 26 (219) 45 (688) 35 (55) 23 (106) 28 (59) 30 (67) D 7 22 12 20 1 13 E 1 2 10 9 14 3 F 14 4 9 9 19 15 F' 0 3 0 3 0 13 G 26 16 11 28 24 17 Other 26 8 23 8 8 9 Physiology and Disease Blood, Cerebrospinal Fluid, and Temperament Champoux et al. (1994, p. 352; 1996, p. 81; 1997, p. 56) compared M. mulatto infants of In- dian ancestry (n = 29) and M. mulatto infants of mixed Indian-Chinese ancestry (n = 13) with re- spect to hematology, serum biochemistry, cere- brospinal 5-HIAA values, and temperament; all infants were nursery reared, and no infants of pure Chinese ancestry were available for this study. The two groups of infants differed significantly (P < 0.05) in hematocrit, hemoglobin, mean cor- puscular hemoglobin concentration, mean corpus- cular hemoglobin, erythrocyte count, gamma-glu- tamyltransferase, phosphorus, and total protein. The authors were uncertain whether the same dif- ferences would be found in mother-reared infants, and they also were uncertain whether to attribute the hematological and serum biochemical differ- ences to environmental adaptation or random ge- netic drift. Compared with the purebred Indian infants, the Chinese-Indian hybrids exhibited lower orienta- tion ability and higiier irritability. In separation- reunion experiments, the Chinese-Indian hybrids tended to exhibit greater locomotor stereotypy, less vocalization, and less social play. Lead Content of Molars Gl?ib and Szostek (1996, p. 216) compared the lead content of the lower first molar in 16 M. mu- latto immatures collected in western India with that in 29 immatures collected in Thailand. The mean lead content in the Indian sample (7.12 ± 2.18 fJLg/g) was nearly five times that in the Thai sample (1.51 ± 0.48 |xg/g; P < 0.001). The au- thors indicate that the difference probably was the result of exposure to different levels of lead in the environment. Malaria The geographic range of M. mulatto is mainly north of the geographic range of the Leucosphy- rus group of Anopheles mosquitoes, which is the group that includes all known natural vectors of macaque malaria (Fooden, 1994, p. 575). Report- ed natural malaria infections in M. mulatto are restricted to Bangladesh, Thailand, and Vietnam, all in the area where the range of M. mulatto over- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 53 laps that of the Leucosphyrus group. In this area, the known incidence of natural infection is 9.3% (n = 290); contrastingly, in northern India, out- side of this area, the incidence of infection is 0% (n > 24,000). The two species of malarial parasites that are known to naturally infect M. mukitta are Plas- nuuiium cynomolgi and P. initi, the most widely distributed of the seven species of Plasmodium that infect macaques (Fooden, 1994, p. 578). Nat- ural and experimental infections with P. cyno- moli^i and P. initi are relatively benign in M. mu- kitta. However, experimental infections of M. mu- latta with P. knowlesi, a macaque parasite that does not occur within the geographic range of M. mulatta. usually are fatal (n > 90). This suggests that M. mulatta has evolved partial resistance to the malarial parasites with which it is sympatric. Viral Infections In a field study, 24 M. mulatta individuals in Bangladesh and six M. mulatta individuals in Thailand were tested for antibodies to reveal in- fections with simian T-lymphotropic retrovirus, type 1 (Ishida et al.. 1985. p. 841; Ishida & Var- avudhi, 1992, p. 163). Of these 30 individuals, only one from Thailand was seropositive. In other species of macaques studied in Thailand, four of 367 M. fascicularis individuals and two of 137 M. arctoides individuals were seropositive for this vi- rus; no species of macaque other than M. mulatta was tested in Bangladesh. In a laboratory study, five M. mulatta individ- uals imported from India and six imported from China were compared with respect to susceptibil- ity to experimental infection with simian immu- nodeficiency virus of macaques (SIVni,,,239) (Joag et al., 1994, p. 439). Plasma virus titer, infectious cell frequency, and virus burden in spleen and lymph nodes all indicated that the rhesus monkeys imported from India were significantly less resis- tant to infection with SIV,„^^,239 than were those imported from China. Natural History Habitats Judging from the geographic distribution of M. mulatta, which is centered at ca. 25°N (Fig. 1), the primary adaptation of this species probably is to the seasonal climate of the subtropical zone (cf. Darlington, 1957, p. 413); captives kept in warm climates apparently are highly susceptible to heat- stroke (Vickers, 1986, p. 522). However, the nat- ural range of M. mulatta does extend to temperate or subalpine habitats in the north (Table 11). The range of humidity extremes tolerated by M. mu- latta includes arid areas in western India and tidal swamps in eastern India and Bangladesh. Al- though most elevational records of M. mulatta are below 2000 m (Table 12), this species has been observed or collected as high as ca. 3200 m in Nepal (Hutu Forest) and ca. 4000 m in Qinghai Province, China (Baizha Plantation, Yushu Xian). Among vegetation types, broadleaf forest is the most common habitat of M. mulatta, but this spe- cies also occurs in mixed broadleaf-needleleaf forests and, least frequently, in needleleaf forests. M. mulatta often inhabits disturbed areas (Blan- ford, 1888b, p. 14; Mills, 1923, p. 222; McCann, 1933b, p. 810; Fooden, 1982b, p. 574; Richard et al., 1989, p. 569; ChaUse, 1997, p. 31; Ruggeri & Timmins, 1997, p. 2), where it raids adjacent cul- tivated fields, and in India it frequently lives in populated areas as a commensal with humans (Southwick et al., 1961. p. 705; Prakash, 1962, p. 83). Habitat variables apparently are related to geo- graphic variation in the scream call of M. mulatta (Feng et al., 1997, p. 27). Arboreality/Terrestriality Judging from the few samples of M. mulatta for which daily arboreality/terrestriality activity patterns have been estimated, this species spends, on average, about 72% of its daylight hours on the ground and about 28% in trees (Table 13; cf. Blanford, 1888b. p. 14; McCann, 1933b, p. 810). Unsurprisingly, forest groups may tend to be somewhat more arboreal than nonforest groups, and in the Sundarbans tidal swamp forests, M. mulatta reportedly rarely descends from the trees (Mandal, 1964, p. 154; Mukherjee & Gupta, 1965, p. 145). In response to sudden danger, M. mulatta flees either on the ground (K. G. Gairdner, 14 April 1916, ZRC 4-188, field tag; Hingston. [1920], p. 244; Green, 1978, p. 154; Mukherjee, 1978b, p. 741; Dang, 1983, p. 1283; Poirier, 1985, p. 298) or into the trees (Mandal, 1964, p. 154; Mukher- jee, 1969, p. 53; Fooden, 1971, p. 32; Lindburg, 1971, p. 45; Pirta & Singh, 1978, p. 277; Wada, 54 FIELDIANA: ZOOLOGY Tablh 1 i. Habitats reported for Macaca nuiUilti . Climate Forest type Culti- Village, temple. Trop- Sub- Tem- Broad Needle- vated Misc. other Refer- Sample area ical tropical perate leaf Mixed leaf field etc. habitat types ences' Afghanistan + + + + 1 Pakistan + + + + 2 India. northern- + + + + + + + + 3 India, western' + + + + Arid 4 India, central + + + + 5 India, eastern' + + + + + Swamp 6 Nepal + + + + + + 7 Bhutan -H + 8 Bangladesh + + + + Swamp 9 Myanmar + + + + + 10 Thailand + + -h n Vietnam + + 12 China, southern* + + + + 13 China, western' + + + + + Subalpine 14 China, central'* + + + + 15 China, northern'' -1- + + + + 16 China: Xianggang (= Hong Kong)'" + + + 17 ' Key to references: /. Puget, 1971, p. 200; Naumann & Nogge. 1973. p. 92. 2. Hingston, [1920], p. 244; M. Iqbal 5 Aug. 1964. USNM 353187. specimen tag; Roberts. 1977. pp. 6, 86; Melnick et al.. 1984, p. 342; Pearl et al., 1987 p. 34. 3. Wells in Lindsay, 1926, p. 599; Southwick et al., 1961a. p. 539; 1961b. p. 703; Neville. 1968b. p. 115 Mukherjee. 1969. p. 47; Lindburg. 1971. pp. 6. 8; Malhotra & Sahi, 1982, p. 25; Tak & Kumar. 1984. p. 203; Wada 1984. p. 470; Seth & Seth. 1985. p. 53; Pirta et al., 1997, p. 102. 4. Prakash. 1962, p. 83; Seth & Seth. 1985. p. 53 Fooden et al., 1981, p. 466. 5. Krishnan. 1972, p. 541; Mukherjee. 1984, p. 260; Fooden et al., 1981, p. 466; Koyama 6 Shekar. 1981, p. 252; Kurup. 1992. p. 16. 6. Mills. 1923. p. 222; McCann. 1933b, p. 810; Roonwal. 1950. p. 16 Mandal, 1964. p. 154; Southwick et al.. 1964. p. 443; Mukherjee & Gupta. 1965. p. 145; Lahan & Sonowal. 1974 pp. 246. 275; Mukherjee. 1978a. p. 275; 1982. p. 77; Tilson. 1983. p. 399; Choudhury. [1991a]. p. 32; 1996, p. 15 1997. p. 10; Gupta, 1994, p. 104; Mukherjee et al.. 1995. p. 25. 7. N. A. Baptista in Hinton & Fry, 1923. p. 403 Chesemore. 1970, p. 164; Richie et al.. 1978. p. 443; Teas et al., 1980. p. 249; Teas. 1983, pp. 212. 215; Chalise, 1997. p. 31. 8. Choudhury. 1990. p. 125; Subba & Santiapillai. 1991-92, p. 32. 9. Green, 1978, p. 154; Gittins & Akonda. 1982. p. 277; Khan. 1985, p. 31; Stanford. 1992. p. 188; Ahsan. 1994. p. 82; Feeroz et al.. 1995. p. 75. 10. G. C. Shortridge in Ryley. 1914, p. 715; H. C. Smith. 14 Jan. 1937. bm(nh) 1937.12.3.76, specimen tag; R. Kaulback. 28 Jan. 1939, bm(nh) 1950.372. specimen tag; Southwick & Southwick. 1985. p. 35. 11. Coolidge, 1940. p. 129 (cf. Allen & Coolidge. 1940. p. 147); Fooden. 1971. p. 32; Aggimarangsee, 1992. p. 119; Sriko.samatara. 1993. p. .34. 12. Pham Nhat. FCXM, pers. comm.. 30 Oct. 1995 (collec'ted at Ban Bu, 1 1 Dec. 1992). 13. Jiang Haisheng et al., 1991, p. 208; pers. obs., Guangxi: Tian'e County, 19-30 Oct. 1992; Zhang & Quan. 1996. p. 14. 14. Weigold. 1935, p. 233; Kaulback. 1938. p. 91; Shen, 1963, p. 140; Zhang & Quan, 1996,^p. 14. 15. Wang Yingxiang, 11 July 1963, Kiz Coll. No. 631094. .specimen tag; Poirier & Hu. 1983. p. 387; Tang Zieying. pers. comm., 19 Oct. 1985 (ob.served at Fujian: Fangdao Nature Reserve. July 1985; Zhang Minhua. zmnh. pers. comm.. 24 Oct. 1985 (ob.served at Zhejiang: Laodian. Aug. 1983); Kang Xiniin. pers. comm.. 24 Oct. 1985 (observed at Zhejiang: Zhidaikou. Aug. 1985); Wada et al.. 1986. p. 81; Zhang & Quan. 1996. p. 14. 16. Yao Jianchu. siz, pers. comm.. 10 Oct. 198^5 (observed at Shaanxi: Dahe and Dashuping. July 1974); Zhang et al.. 1989. p. 376; Qu et al.. 1993. p. 609; Southwick et al.. 1996. p. 97; Zhang & Quan. 1996. p. 14. 17. Southwick & Southwick. 1983. p. 18. - Jammu and Kashmir, Himachal Pradesh. Punjab. Haryana. Delhi. Uttar Pradesh. ' Rajasthan. Gujarat. Maharashtra. •* Madhya Pradesh. Andhra Pradesh. Bihar. Orissa. ' West Bengal. Sikkim. Arunachal Pradesh. Assam. Nagaland. Meghalaya, Manipur, Tripura, Mizoram. ''Yunnan, Guangxi, Guangdong (including Hainan). ' Sichuan, Xizang, Qinghai. ** Fujian, Jiangxi. Hunan. Guizhou. Hubei. Anhui. Zhejiang. ' Henan. Shaanxi. Shanxi. Hebei (provincial population apparently extirpated in 1987). '"Population probably introduced (Herklots. 1951, p. 83; Marshall. 1967. p. 45). FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 55 Table 12. Frequency distribution of elevation records oi Macaca muhitta (see Gazetteer, Appendix 2). PLlevation (m) Number of records 0-500 500-1000 1000-1500 1500-2000 2()0()-25()0 2500-3000 3000-3500 3500-4000 4000-4500 Total 155 92 34 13 20 7 3 1 2 327 1984, p. 494; Choudhury, [1991b). p. 123; Chopra et al., 1992, p. 81). Nighttime sleeping sites gen- erally are in trees (Hingston, [1920], p. 244; Ko- ford, 1963. p. 143; Mandal, 1964, p. 154; Shou et al., 1964, p. 60; Lindburg, 1971, p. 29; Vessey, 1973, p. 614; Makwana. 1978, p. 486; 1979b, p. 919), but groups in Afghanistan and northern Chi- na have been reported to sleep on the ground (Pu- get. 1971, p. 200; Qu et al., 1993, p. 616), and urban groups often sleep on the roofs of buildings (Ojha, 1977, p. 519; Mukherjee, 1969, p. 49; 1978a, p. 278). Two parturitions that have been observed in wild populations apparently occurred on the ground (Lindburg, 1971, p. 77; Mathur, 1994, p. 132). Geographic variation in the relative frequency of terrestrial locomotion in M. mulatta reportedly is correlated with variation in morphology of the scapula, clavicle, and humerus (Yu et al., 1993, p. 87; Xue et al., 1998a, p. 147; 1998b, p. 29; 1999, p. 140). Swimming M. mulatta is capable of swimming across a water gap ca. 1 km wide (Drickamer & Vessey, 1974, p. 362; Varley & Vessey, 1977, p. 54; Sade, 1985, p. 28; Rawlins & Kessler, 1986b, p. 26). This species reportedly swims to search for food (Mukherjee & Gupta, 1965, p. 145; Dang, 1983, p. 1283), to escape from danger (Muir, 1916, p. 353; Southwick et al., 1974, p. 198; Berman, 1977, p. 763), and apparently also for pleasure and/or thermoregulation (McCann, 1933b, p. 810; Pilleri & Filled, 1982, p. 158; Malik & Menon, 1992, p. 39). In captivity, 2-day-old infants are capable of swimming (Riopelle, 1980, p. 262), and juveniles have been trained to swim under- water and to open a food box underwater (An- derson et al., 1992, p. 2; 1994, p. 356). Group Size and Composition The mean size of ca. 1,182 nonprovisioned or minimally provisioned groups for which data are available is ca. 32.2 individuals (Table 14); re- ported extremes are two and ca. 250 individuals. The size of nonprovisioned groups apparently tends to average largest (86.1-ca. 105.0 individ- Tabi.f. 13. Arboreal/terrestrial behavior recorded during daylight hours in samples of Macaca mulatta. % A rboreality - Sample size Habitat type Mean ± SD Extremes References' Northwestern Pakistan Forest 34 Northern India 1 group 1 Ancient fort area Temple Urban Pond area Roadside Canal bank Forest 20 27.8 ± 2.3 17.1 ± 3.4 34.1 ± 13.9 21.0 21.8 40.2 ±11.7 25.2-29.5 13.2-19.4 10.2-46.5 20.1-23.6 29.3-52.6 1 locality 3 localities 3 groups (2 localities) 5 localities 1 locality 2 localities 3 localities 2 3 3 3 3 3 3 Island colony All samples Cayo 22.5 ± 5.6 28.4 ±11.1 Santiago, Puerto Rico 16.5-27.7 10.2-52.6 (provisioned) 3 individuals 20 groups or localities 4 ' Key to references: 1. Goldstein & Richard. 1989, p. 555. 2. Malik. 1986. p. Ill; Malik & Southwick. 1988a. p. 346. 3. Chopra et al., 1992. p. 92. 4. Fisler. 1967, p. 74. 56 FIELDIANA: ZOOLOGY uals) at the northern extremes of the species' geo- graphic range — in Afghanistan and in the Chinese provinces of Qinghai and Henan (cf. Southwick et al., 1996. p. 102). Solitary males, living inde- pendently of nearby troops, have been observed in all parts of the species range (references cited in Table 14). Provisioned groups — one of which reportedly included 1.045 members — tend to av- erage larger than nonprovisioned groups. Nonprovisioned and provisioned groups of var- ious sizes have been observed to split perma- nently into two autonomous daughter groups (Southwick & Beg. 1961, p. 390; Koford, 1966. p. 2; Missakian, 1973b. p. 622; Malik et al.. 1984. p. 315; 1985. p. 417; Seth et al.. 1986. p. 115; Malik, 1992. p. 8; Wang et al., 1996. p. 265). The smallest group known to have undergone fission included 28 individuals (daughter groups. 10 and 18 individuals) (Melnick & Kidd. 1983. p. 230). Fission probably usually occurs between matri- lines (Chepko-Sade & Sade. 1979, p. 70). In nonprovisioned and provisioned groups, the average sex ratio is approximately one sexually mature male to three sexually mature females (Ta- ble 15). The reported minimum ratio is approxi- mately one male to 12 females, and the reported maximum ratio is approximately three males to two females. Home Range, Day Range Home range averages approximately 65 ha in 323 nonforest groups of M. miilatta and 196 ha in 129 forest groups (Table 16). Overlap of home ranges of adjacent troops is extensive (Lindburg, 1971, p. 32; Southwick et al., 1982. p. 623; Jiang Haisheng et al., 1991. p. 212) and may reach 100% (Makwana, 1979b, p. 919). Depending on local food and water sources and snow cover, different parts of a group's home range may be used in different seasons (Kurup, 1965, p. 193; Neville, 1968b, p. 113; Lindburg, 1977b. p. 241; Wada. 1984. p. 487; cf. Pearl et al., 1987, p. 36). Day ranges average 1.15 km in nine nonforest groups and 1.91 km in >16 forest groups (Table 17). Population Density In areas inhabited by M. mulatto, the mean re- ported population density is 37.2 individuals/km- in forest habitats and 201.1 individuals/km- in nonforest habitats (Table 18). Among the nine for- est habitat areas surveyed, mean population den- sity was unusually high — 120.0 individuals/km- — on Hainan Dao. a tropical island off the south- eastern coast of China. In the provisioned free- ranging population of M. mulatto that was intro- duced in 1938 on Cayo Santiago (area 0.152 km-), Puerto Rico, the mean population density in 1983 had reached 7638.2 individuals/km-. Diet The natural diet of M. mulatto is primarily vegetarian and includes fruits, seeds, flowers, leaves, buds, shoots, twigs, stems, roots, bark, pith, and resin of hundreds of species of angio- sperms, gymnosperms, and fungi (Table 19). An- giosperm plants consumed include trees, shrubs, climbers, grasses, and other herbs. In six care- fully surveyed geographic areas, the following minimum numbers of species of wild plants were discovered to be exploited for food by M. mu- latto: northern Pakistan. 35 species (Goldstein & Richard, 1989, p. 552); Himachal Pradesh, north- ern India. 121 species (Pirta et al.. 1997, p. 103); Uttar Pradesh, northern India, 150 species (Lind- burg, 1977a, pp. 263-268; Makwana, 1979a, p. 244); central Nepal, 61 species (Marriott, 1978a, p. 759); Bangladesh, 41 species (Ahsan, 1994, p. 82); and Henan. east-central China. 73 species (Qu et al., 1993. p. 612). On Cayo Santiago, Puerto Rico, the provisioned introduced popula- tion of M. mulatto supplements its diet of mon- key chow by feeding on 73 of the 163 plant spe- cies that grow on the island (Marriott et al., 1993, p. 332). In addition to exploiting wild plants, natural populations of M. mulatto oppor- tunistically raid numerous species of cultivated crop plants (Makwana, 1979a. p. 247; Siddiqi & Southwick. 1980. p. 55; Poirier & Hu. 1983. p. 387; Malik & Southwick. 1988a. p. 339; Lai, 1990, pp. 113. 123; Gupta & Kumar, 1992. p. 227; Datta, 1996. p. 941). Seasonal variation has been noted in the species and parts of plants that are consumed (Lindburg, 1977a, p. 263; Wada, 1984, p. 480; Goldstein & Richard. 1989, p. 554; Fellowes, 1992, p. 132; Gupta & Kumar, 1992, p. 227; Qu et al., 1993, p. 611). Plant consumption by M. mulatto apparently also varies geographically (Table 19). Judging from available data, grasses and other herbs pro- vide most of the natural food for this species in FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 57 Tablh 14. Group size reported for Maccica miilatta. (Jroup size No. of groups Sample area Mean Minimum Maximum References' Nonprovisioned 1 or minimally provisioned groups Afghanistan ca. 105.0 ca. 30 ca. 250 >12 7 Pakistan ca. 40.0 ca. 12 ca. 78 >9 2 India, northern Jammu and Kashmir. Himachal Pradesh ca. 30.8 4 50 78 3 Punjab, Haryana. Rajasthan 24.0 <11 >73 35 4 Uttar Pradesh ca. 28.9 5 127 ca. 77 5 Various states- Urban 33.5 4 116 134 6 Rural 29.4 3 165 318 6 Forest 33.3 4 ca. 100 111 6 India, peninsular Gujarat, Maharashtra 58.4 ca. 20 ca. 100 8 7 Andhra Pradesh ca. 37.8 6 ca. 250 36 8 India, eastern Orissa. Bihar 44.0 19 69 2 9 West Bengal ca. 24.1 2 ca. 100 47 10 Assam ca. 19.9 ca. 9 ca. 60 12 U Tripura 26.6 <5 >76 162 12 Nepal 37.9 20 84 10 13 Bangladesh 16.1 2-3 84 68 14 Myanmar 15.3 9 26 3 15 Thailand 35.0 20 50 2 16 Vietnam — ca. 15 ca. 100 — 17 China Hainan Dao 58.9 20 110 18 18 Guangdong 21 — — 1 19 Guangxi 33.8 11 73 6 20 Hunan 24 — — 1 19 Hubei ca. 35.0 — — 3-4 21 Henan 86.1 35 125 23 22 Qinghai ca. 100.0 — — 12 23 Mean ca. 32.3 2 Provisioned groups ca. 250 1.188 India, northern Jammu and Kashmir, Himachal Pradesh 54.4 20 129 5 24 Delhi 51.2 19. 123 6 25 Rajasthan 66 — — I 26 Uttar Pradesh 51.9 16 128 14 27 Various states- 135.2 20 1,045 16 28 India, peninsular Andhra Pradesh ca. 43.8 8 ca. 100 4 29 India, eastern Manipur 128 — — 1 30 Nepal' 54.6 <29 138 12 31 China Hainan Dao 59.5 52 67 2 32 Henan 103.7 83 123 3 33 Mean ca. 76.9 8 1 .045 64 Provisioned introduced population (Puerto Rico) Cayo Santiago 191.7 100 306 6 34 ' Key to references: /. Puget. 1971, p. 199: Naumann & Nogge, 1973, p. 92. 2. Roberts, 1977. p. 87: Igbal & Rub, 1980. p. 214; Melnick et al., 1984, p. 344. 3. Malhotra & Sahi. 1982, p. 27: Tak & Kumar, 1984. p. 203: Wada, 1984, p. 481: Pirta et al.. 1997. p. 100. 4. R. Singh, 1984, p. 50: Boonratana & Edwin, 1986. p. 110; Wolfe & Mathur, 1988. p. 537; Mathur & Manohar, 1990, p. 356; Chandel, 1992. p. 121; Gupta & Kumar. 1992, p. 226; Mathur, 1994, 58 FIELDIANA: ZOOLOGY northern Pakistan, whereas fruits and leaves of trees, shrubs, and cUmbers provide most of the food in Bangladesh and in the Indian states of Uttar Pradesh and Rajasthan. Similarly, gymno- sperm seeds and needles are frequently consumed in northern parts of the geographic range (Af- ghanistan, Pakistan. Himachal Pradesh. Punjab, Henan, Hubei) but apparently are less frequently eaten elsewhere; this obviously is correlated with the geographic distribution of gymnosperms (Kuchler, 1978, p. 17). Larval and adult insects (Orthoptera, Isoptera, Hemiptera, Coleoptera, Lepidoptera, Hymenop- tera) apparently are the most common animal food of M. mulatto (Lindburg, 1971, pp. 23, 33; Makwana, 1979a, pp. 243-247). Other known an- imal food includes spiders, crayfish, crabs, shell- fish (?bivalves), fish, birds' eggs, and honey- combs (Table 19). In Vietnam, animal food is es- timated to constitute 5-7% of the diet of M. mu- latto (Dang, 1983, p. 1283). In parts of India, however, consumption of animal foods may be less frequent; at three Indian localities, feeding on insects was never observed (Siddiqi & Southwick, 1980, p. 55; Malik & Southwick, 1988a, p. 340; Gupta & Kumar, 1992, p. 227), and at another locality, M. mulatto individuals appeared reluctant to eat hen's eggs that were provided for them (Lindburg, 1971, p. 33). Ingestion of soil (geophagy) by M. mulatto has been observed at three localities in India (Delhi, Rajasthan, Asarori), one locality in Nepal, and one locality in China (Table 19). At one of the Indian localities (Asarori), the soil was specifical- ly identified as termite mound soil; monkeys at this locality also occasionally licked whitewash off painted walls. Geophagy also has been re- ported in the provisioned introduced population of M. mulatto on Cayo Santiago, Puerto Rico (Sul- tana & Marriott, 1982, p. 338); in these monkeys, selective ingestion of clay may function to pre- vent or ameliorate gastrointestinal disorders, in- cluding endoparasitism (Mahaney et al., 1995, p. 331; Knezevich, 1997, p. 73; cf. Bolton et al., 1998, p. 204). For captive, relatively sedentary adult female M. mulatto (mean weight = 7.7 kg), the daily maintenance energy requirement is estimated to be 430 kcal (Henderson et al.. 1993, p. 10; cf. Bourne, 1975, p. 99). This can be supplied by 150 g of monkey chow (10 large biscuits), supple- mented by small amounts of fresh fruit and mul- tivitamin tablets; such a diet has been shown to maintain body weight in singly housed monkeys for at least 1 1 weeks. In the free-ranging provi- sioned group on Key Lois. Florida, adults con- sume a daily average of ca. 225 g of chow in addition to naturally available food (Pucak et al., 1982. p. 207). Preliminary experimental evidence suggests that calorie-restricted diets may retard age-related pathology in captive M. mulatto (Couzin, 1998, p. 1018). During the rainy season, water requirements of M. mulatto are met primarily by consumption of succulent plant food (Mukherjee & Gupta, 1965, p. 146; Lindburg, 1971. p. 35; Malik & South- p. L^2. 5. Nolle. 1956. p. 180; Neville. 1968b, pp. 114, 117; 1968c, p. 15: Mukherjee. 1969. p. 48; Lindburg, 1971, p. 13; Singh. 1975, p. 472: Southwick & Siddiqi. 1977b. p. 342: Pirta. 1977-78. p. 126: 1982. p. 401: Singh. 1982, p. 8: Laws & Laws. 1984. p. 35; Imam & Yahya. 1995. p. 4. 6. Dolinow & Lindburg. 1980. p. 211: Seth & Seth, 1993, p. 55: cf. Southwick et al.. 1965, p. 120, and Mukherjee & Mukherjee, 1972, p.^67. 7. Fooden et al.. 1981, p. 466. 8. Fooden et al.. 1981, p. 466: Kurup, 1992, p. 15. 9. Krishnan. 1972. p. 541: Mukherjee. 1984. p. 260. 10. Mandal, 1964, p. 164: Southwick et al.. 1964. p. 446: Saha, 1974, p. 21 1: Southwick et al., 1974. p. 187: Mukherjee et al., 1995, p. 27: cf. de Poncins, 1935, p. 846, and Mukherjee & Gupta, 1965, p. 145. //. Mukherjee, 1978b, p. 741: Pilleri & Pilleri, 1982, p. 158: Choudhury. [199 la), p. 32: [1991bl. p. 123. 12. Mukherjee. 1982, pp. 73, 80; Gupta, 1994, p. 104. 13. Richie et al., 1978, p. 443: Teas et al.. 1980. p. 250: Marriott, 1988, p. 128. 14. Green, 1978. p. 153: Oppenheimer et al., 1983, pp. 195, 198: Khan, 1986. p. 38: Stanford. 1991, p. 17: Feeroz et al., 1995, p. 75: cf. Hendrichs. 1975, p. 171, and Ahsan, 1994, p. 83. 15. Southwick & Southwick, 1985. p. 35. 16. Fooden, 1971, p. 32. 17. Dang. 1983. p. 1283. 18. Southwick et al., 1996. p. 100. 19. Feng et al.. 1997. p. 27. 20. Wang et al., 1996, p. 266. 21. Poirier. 1983. p. 128. 22. Qu et al.. 1993, p. 614; Southwick et al.. 1996. p. 100. 23. Tan & Poirier, [1991]. p. 130. 24. Malhotra & Sahi, 1982. p. 27: Wada, 1984, p. 481. 25. Southwick et al.. 1961a, p. 543; Malik et al., 1984, p. 315. 26. Singh, 1992, p. 192. 27. Southwick et al.. 1961a. p. 543: Pirta. 1984. p. 281. 28. Seth & Seth. 1993, p. 55. 29. Fooden et al.. 1981, p. 467. 30. Mukherjee, 1978a, p. 277. 31. Southwick et al., 1980, p. 165: 1982, p. 624: Teas et al., 1982. p. 65. 32. Southwick et al., 1996, p. 100. 33. Qu et al., 1993, pp. 610. 614. 34. Rawlins & Kessler, 1986c, p. 55. - Pooled data reported for groups observed in several northern Indian states: some of these groups probably also are included in reports cited in preceding entries for northern Indian states. ' Cf. Chalise & Ghimire, 1998, pp. 1 r-15. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 59 Table 15. Ratio of sexually mature males to sexually mature females reported in groups of Macaca mulatta. Number of Pooled Group ses : ratios sexually mature No. of Sample area sex ratio Minimum Maximum individuals groups References' Nonprovisioned or minimally provisioned groups- Pakistan 0.54 7 9 126 7 1 India, northern Jammu and Kashmir, Himachal Pradesh 0.50 0.29 1.43 1,029 68 2 Haryana, Rajasthan 0.26 0.12 0.50 103 4 3 Uttar Pradesh 0.39 0.18 1.00 691 43 4 Various states' Urban 0.34 <0.16 >1.17 1,790 134 5 Rural 0.39 <0.21 >0.71 4,053 318 5 Forest 0.34 <0.18 >0.60 1,422 111 5 India, peninsular Andhra Pradesh 0.33 0.08 0.64 129 11 6 India, eastern Orissa, West Bengal 0.37 0.09 1. 00 629 44 7 Tripura 0.47 <0.25 >0.67 1,986 174 8 NepaP 0.35 0.18 0.50 131 6 9 Bangladesh 0.29 0.14 1.00 208 16 10 Myanmar 0.50 9 9 46 3 11 Mean 0.39 0.08 Provisioned 1.43 groups 12,343 939 India, northern Jammu and Kashmir, Himachal Pradesh 0.47 0.30 0.75 109 3 12 Delhi 0.38 0.25 0.75 95 6 13 Uttar Pradesh 0.48 0.29 0.85 375 14 14 Various states^ 0.26 <0.21 S0.47 961 16 15 India, eastern Manipur 0.47 — — 56 1 16 Nepal 0.26 9 9 289 12 17 China: Henan 0.30 0.21 0.48 82 2 18 Mean 0.35 <0.21 0.85 1,967 54 ' Key to references: 1. Melnick & Pearl, 1987, p. 128; cf. Igbal & Rub, 1980, p. 214: Goldstein & Richard, 1989, p. 532. and Rab et al., 1991, p. 220. 2. Camperio Ciani. 1984, p. 373; Malhotra & Sahi. 1984, p. 78; Pirta et al., 1997, p. 100. 3. Prakash, 1962. p. 83; Singh et al., 1984, p. 265; Chandel, 1992. p. 121; Mathur. 1994, p. 132; cf. Mathur & Manohar. 1990, p. 356. 4. Neville, 1968b. p. 114; Mukherjee. 1969, p. 48; Lindburg, 1971, p. 13; Singh, 1975, p. 472: Pirta, 1977-78, p. 126; Pirta & Singh, 1982, p. 20; Imam & Yahya, 1995, p. 4. 5. Dolinow & Lindburg, Seth et al., 1983. p. 40; Chopra et al.. 1992, p. 86; cf. Southwick 1972, p. 67. 6. Kurup, 1992. p. 15. 7. Mandal, 1964. p. 164; 1974, p. 211; Mukherjee, 1984, p. 260; Mukherjee et al., 1995, 104. 9. Richie et al., 1978. p. 443; Teas. 1983. pp. 217. 221; Marriott, 1988. p. 128. 10. Oppenheimer et al.. 1983, pp. 195, 198. 11. Southwick & Southwick. 1985. p. 35. 12. Camperio Ciani. 1984. p. 373; Malhotra & Sahi. 1984, p. 78. 13. Southwick et al.. 1961a. p. 543; Malik et al., 1984. p. 315. 14. Southwick et al., 1961a, p. 543; Pirta & Singh, 1982. p. 20. 15. Seth & Seth, 1993. p. 55; Chopra et al., 1992, p. 86. 16. Mukherjee. 1978a, p. 277. 17. Teas et al., 1981a, p. 119; 1982, p. 65; cf. Johnson et al.. 1988. p. 179; Chalise & Ghimire, 1998, pp. 14, 15. 18. Qu et al., 1993, pp. 613. 615. -Cf. Wang et al.. 1996. pp. 267. 268. ^ Pooled data reported for groups observed in several northern Indian states; some of these groups probably also are included in reports cited in preceding entries for northern Indian states. 'Cf. Chalise, 1997, p. 31; Chalise & Ghimire, 1998, p. 15. 1980, p. 211; Seth & Seth, 1983, p. 65; 1993, p. 55: et al., 1965, p. 120, and Mukherjee & Mukherjee Southwick et al., 1964, p. 446; 1974. p. 187; Saha, p. 27. 8. Mukherjee, 1982, p. 80; Gupta, 1994, p. wick, 1988a, p. 345). During the dry season, these monkeys move as a group to drink at streams or other open water sources, often in the morning and evening (N. A. Baptista in Hinton & Fry, 1923, p. 403; Lindburg. 1971, p. 36; Paget, 1971, p. 200; Naumann & Nogge, 1973, p. 92; Mak- wana, 1979b, p. 919). Drinking apparently is usu- ally by oral suction, but monkeys also sometimes 60 FIELDLANA: ZOOLOGY Table 16. Home range area reported for g roups of Moccua miilatta. Home ranse area (ha) Mean No. of Refer- Sample area Habitat Mean Minimum Maximum group size groups ences' Nonprovisioned or minimally pro> isioned groups India, northern Himachal Pradesh Forest 178 60 260 42.2 4 / Haryana Rural 5 — — 26 1 2 Uttar Pradesh Urban ca. 32 ca. 5 60 36.2 4 3 Rural ca. 10 ca. 10 ca. 10 36.0 2 4 Forest ca. 449 56 ca 1.560 42.3 2\ 5 Various stales- Urban 95 <6 >100 34.7 116 6 Rural 49 <6 >162 35.7 199 6 Forest 129 <129 >2.020 35.4 79 6 India, eastern Orissa Urban 3 — 69 1 7 Nepal Forest 70 — — 84 1 8 Bangladesh Forest >50 ■1 0 37.0 2 9 Vietnam Forest ca. 250 100 400 •7 0 10 China Hainan Dao Forest 37 16 72 ca. 60 19' U Henan Forest 1.333 1.100 1,500 60.0 3 12 Mean Nonforest ca. 64.8 3 >162 35.4 323 Mean Forest ca. 195.9 16 >2.020 ca. 41.3 129 Provisioned groups India, northern Rajasthan Forest* 400 — — 68 1 13 Uttar Pradesh Temple 1.7 1.5 2 98.5 -) 14 Various states- Temple 76 <70 >230 135.2 16 6 India, eastern Manipur Temple 8 — — 128 1 15 Nepal Temple ? 2.5 24 54.6 12 16 China: Henan Forest 2,000 1,800 2.200 103.0 2 12 Mean Temple 64.6 1.5 >230 131.0 19 Mean Forest 1466.7 400 2.200 91.3 3 ' Key to references: I. Wada, 1984. p. 482. 2. Singh el al.. 1984, p. 264. 3. Neville. 1968b. p. 1 18: Pirta & Sinah. 1980, p. 516. 4. Mukherjee, 1969, p. 51. 5. Neville.^ 1968b. p. 118: Lindburg, 1971, p. 28: Pirta & Singh. 1982,^p. 20. 6. Seth et al., 1983. p. 40: Chopra el al.. 1992, pp. 86, 92: Seth & Seth, 1993. p. 55. 7. Mukherjee. 1984. p. 260. S. Man-iott. 1988. p. 129. 9. Stanford, 1991, p. 17: cf. Ahsan. 1994, p. 83. 10. Dang. 1983. p. 1283. //. Jiang Haisheng et al.. 1991. p. 212. 12. Qu et al.. 1993. p. 614. 13. Singh. 1992, p. 192. 14. Pirta & Singh. 1982. p. 20. 75. Mukherjee, 1978a, p. 278. 16. Southwick et al.. 1982. p. 623: Teas et al.. 1980. p. 253. - Pooled data reported for groups observed in several northern Indian states: some of these groups may also be included in reports cited in preceding entries for northern Indian states. ' Includes two provisioned groups. ^ Near temples. dip their hands into a water source and lick the water from their hands (Lindburg, 1971. p. 36 Morrison & Menzel. 1972, p. 54; Makwana 1979a, p. 251; Malik & Menon. 1992, p. 39) Adults in the provisioned group on Key Lois Florida, drink ca. 500 ml of water per day (Pucak et al.. 1982, p. 207). In indigenous M. miilatta populations studied, the average proportion of waking hours that are spent in feeding varies from 15.8% to 45.0% (Ta- ble 20). Feeding activity generally peaks twice a day — in the morning, shortly after waking, and in the afternoon, before roosting for the night (Siddiqi & Southwick. 1980, p. 60; Jiang et al., 1988b, p. 295; Malik & Southwick, 1988b, p. 101; Gupta & Kumar, 1992, p. 229). In winter, the length of the midday feeding pause tends to be reduced (Lind- burg, 1977b, p. 231; Pearl et al.. 1987, p. 38). Predators Observers have recorded attacks on M. mulatta by raptorial birds, dogs, weasels, leopards, and ti- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 61 Tabm; 17. Day range rcporlcd for groups oi Mcicacd iiiiildira. Day range (km) Mean Number of groups Sample area Habitat Mean M inimum Maximum group size observed References' India Himachal Pradesh Urban 1.11 0 3.1 27.8 2 / Forest 1.22 0 4.6 41.2 ca. 5 y Rajasllian Urban 1.75 •7 >2.0 46.0 2 2 Rural 0.90 9 >1.0 49.4 2 2 Forest 4.00 7 >4.0 104 1 2 llttar Pradesh Urban 0.93 7 >1.0 56.3 3 2, 3 Forest 2.04 <0.4 >3.5 >50.4 >10 2. 3. 4 Bangladesh •1 2.00 7 ■-> 15.0 ') 5 China: Hainan Dao Forest ■) <1.0 2.3 -60 9 6 Mean Nonforest 1.15 0 3.1 46.1 9 Mean Forest 1.91 0 4.6 >50.9 >16 ' Key to ret'erenees: /. Wada, 1984, pp. 481, 488, 491. 2. Seth & Seth, 1983, p. 64 (day range units reported as kin-). 3. Neville, 1968b, p. 118. 4. Lindburg, 1971, p. 25; Makwana, 1978, p. 488. 5. Ahsan, 1994, p. 83. 6. Jiang et al.. 1988b. p. 296. gers (see below); additional reported predators are sharks and crocodiles (in the Sundarbans tidal swamps) and snakes (Mukherjee & Gupta. 1965, p. 145; Seth et al., 1983, p. 42). In Uttar Pradesh, India, hawks were observed to kill young rhesus monkeys (Pirta & Singh, 1978, p. 275; cf. Lind- burg, 1971, p. 43), and in Henan, China, an eagle was observed to injure two juveniles (Qu et al., 1993, p. 610). The sight of raptorial birds is known to provoke alarm responses in M. miilatta, both in India (Roonwal & Mohnot, 1977, p. 134; Singh & Pirta, 1983, p. 85) and in the introduced population on Cayo Santiago, Puerto Rico (Cha- pais & Schulman, 1980, p. 740). At a temple in India, rhesus monkeys attacked and killed two hawks (Pirta, 1984, pp. 271, 276). Attacks by dogs that resulted in injury or death to M. mulatta have been documented in Uttar Pra- desh and Delhi, India (Lindburg, 1971, p. 73; Makwana, 1978, p. 485; Johnson & Southwick, 1984, p. 211; Malik et al., 1985, p. 418); Cham- pion (1929, p. 424) noted that M. mulatta indi- viduals invariably called in the direction of his dog when he and the dog walked together in the forest. Fatal attacks by weasels on M. mulatta in- fants have been reported by Pirta and Singh ( 1978, p. 275; cf. Singh & Pirta, 1983, p. 85), who also report an alarm response to a weasel by these monkeys. Leopards are a major predator of M. mulatta (Champion, 1934, p. 120; Roberts, 1977, p. 88; Roonwal & Mohnot, 1977, p. 134). Pre- dation by tigers, alarm responses, and the appar- ent mobbing of a tiger by M. mulatta have been reported in India and Bangladesh (Mandal, 1964, p. 157; Hendrichs, 1975, p. 184; Lindburg, 1977b, p. 242). Experimental study of captive M. mulatta in- dicates that fearful avoidance of snakes is much stronger in wild-reared individuals than in labo- ratory-reared individuals (Joslin et al., 1964, p. 349; Mineka et al., 1980, p. 655). Naive, fearless laboratory-reared monkeys may become fearful of snakes as a result of exposure to the fearful be- havior of other monkeys (Mineka et al., 1984, p. 363; Cook et al., 1985, p. 595). In captive M. mulatta, fear of snakes seems to be greater in dominant individuals than in subordinate individ- uals (Brennan & Anderson, 1988, p. 357; Peugeot et al., 1994, p. 88). Intergroup Behavior The behavioral interactions of adjacent groups of M. mulatta appear to be highly variable (Table 21). Some observers report frequent intergroup contacts, whereas others report that intergroup contacts are rare and apparently avoided (Malik et al., 1984, p. 315; Makwana, 1979b, p. 920). Although harmonious intergroup encounters, in- cluding intergroup matings, have been observed, most contacts .seem to arouse antagonism, which may range from tense behavior or relatively mild branch shaking to violent physical combat (South- wick et al., 1965, p. 143; Neville, 1968c. p. 15; Lindburg, 1971, p. 37). At some localities, both peaceful and ho.stile contacts have been observed between the same groups. During contacts, larger 62 FIELDIANA: ZOOLOGY Tablf. 18. Popul ation density re ported for Macaca mulatto. ] Population density (individuals/km-) Number of groups Refer- Sample area Habitat Mean Minimum Maximum surveyed ences' India, northern Himachal Pradesh Mixed- 109.0 21.2 217.0 >5 / Delhi Ancient fort area' 51.0 32.0 70.2 6 2 Rajasthan Urban 385.4 3.1 418.4 14 3 Temple^ 947.6 882.7 1,012.6 6 3 Uttar Pradesh Urban 752.0 — — >2 4 Rural 44.8 — — 5 5 Forest 27.0 18.2 57.1 >16 6 Various states^* Urban 154.2 — — 116 7 Rural 192.5 <85.9 >356.4 199 7 Forest 34.2 — — 79 7 Temple^ 243.6 — — 16 7 India, eastern Assam Forest 28 — — — 8 Nepal Forest 9.3 — — 8 9 Bangladesh Forest 21.1 5.0 52.0 >33 10 China Hainan Dao Forest' 120.0 <50.0 <250 20 n Guangdong Forest 48.0 — — 1 12 Guangxi Forest 55.6 — — 14 13 Hunan Forest 12.0 — — 1 12 Henan Forest' 7.2 — — ca. 25 14 Mean Nonforest 201.1 3.1 1,012.6 >364 Mean Forest 36.2 <5.0 <250 >197 Introduced provisioned populations (U.S.A.) South Carolina Morgan Island Island 2.320 — — 6 15 Florida Key Lois Island 1,270 — — 7 16 Racoon Key Island 1,790 — — 8 16 Puerto Rico Cayo Santiago Island 7.638.2 — — 6 17 ' Key to references: /. Camperio Ciani. 1984, p. 373; Ross et al., 1993, p. 161: cf. Pirta et al.. 1997, p. 100. 2. Malik, 1989b, p. 581. 3. Mathur & Manohar. 1990, pp. 353, 355. 4. Neville, 1968b, p. 120. 5. Lindburg, 1969, p. 1176. 6. Neville, 1968b, p. 120; Pirta, 1977-78, p. 128. 7. Seth & Seth, 1993, p. 56. 50% + + + + + + + 20 + + + 67% + 21 5%_7%24 + 26 + + +1' + 15 / 2 3 4 5 6 7 8 9 10 11 12 13 13 14 15 16 17 18 19 20 21 22 23 24 25 26 27 28 '"* Termite mound soil with high iron content. '■* Includes grass. '■^ Termite mound soil; whitewash licked off wall. "■ Total angiosperm nonherb dietary proportion, 5.9%. '^ Crow killed by monkeys at Chhatari-do-Raha or Sumera Fall, not eaten; unsuccessful attempt to catch another bird also observed. '* Total angiosperm nonherb dietary proportion, 47.0%. '** Plant parts eaten by monkeys are not specified. 2" Fish. -' Brood nest section was removed from wild bee nest by monkeys. -- Bird's egg. -^ Fruits constitute 80% of diet in winter; leaves constitute most of diet in summer. -"^ Includes insects, crayfish, shellfish, and fish. 2'^ Population probably introduced (Herklots, 1951, p. 83). -^ Unspecified invertebrates, presumably arthropods. -'' Population now extinct. ther with the spring birth peak (Neville, 1968a, p. 772; Melnick et al., 1984, p. 238) or with the fall mating peak (Lindburg, 1969, p. 1177). One ju- venile female was observed to transfer to a new group (Makwana, 1978, p. 485); this transfer oc- curred in the spring. In provisioned introduced populations, male transfer is approximately four times more frequent during the peak mating season than during other seasons (Vandenbergh, 1967, p. 189; Boelkins & Wilson. 1972, p. 132). A recent study indicates that the age at which a male transfers to another FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 65 Tablk 20. Daily waking-hour' lime budget cslimalcs C/r ) for Macaca miilalta.- Sample area Feeding' Locomotion Resting^ Social behavior^ Other References'' Pakistan: forest India, northern Delhi: fort site Uttar Pradesh: forest^ Various slates Temple Urban Pond Roadside Canal bank Forest Nepal Temple* Forest Cayo Santiago 45 10 35 10 — 1 22.8 18.6 33.7 20.8 4.1 2 32.3 44.8 22.9 — — 3 26.7 15.8 36.7 18.6 2 2 4 21.9 19.3 36.7 19.7 2.4 4 28.0 11.2 39.3 20.5 1.0 4 31.3 14.3 34.7 19.3 0.4 4 25.0 19.9 40.0 13.0 2.1 4 33.6 24.2 35.1 5.6 1.5 4 27 25 28 21 1 5 15.8 39.4 27.0 15.1 2.7 6 Introduced provisioned population (U.S.A.) 13.3 23.1 36.3 25.0 2.3 7 ' Ca. 13 hr/day (Marriott. 1978b. p. [27]; Malik & Southwick. 1988b. p. 101). ^Cf. Wolfe. 1992. p. 48. ' Includes foraging and drinking. ■^ Includes self-grooming and category "Look." ^ Includes allogrooming and playing. " Key to references; 1. Goldstein & Richard. 1989. p. 547. 2. Malik & Southwick. 1988b. p. 101; Malik & Mennon, 1992. p. 35. 3. Neville. 1968b. p. 120. 4. Chopra et al., 1992. p. 92. 5. Teas et al.. 1980. p. 254. 6. Marriott. 1988. p. 134. 7. Marriott. 1988. p. 134; cf. Fisler, 1967, p. 72. ■'Means for five groups (extremes: feeding. 10.6%-47.8%; locomotion, 32.8%-71.2%; resting. 15.6%-36.5%). * Percentages reported as rounded whole numbers (sum 102%). group is positively correlated with the level of central nervous system serotonin activity (Mehl- man et al., 1995, p. 909). Interspecific Behavior Intrageneric — M. mulatto is broadly sympat- ric with M. cissamensis, M. thihetana, and M. arc- toides; it is more narrowly sympatric with M. ne- mestrina; and it is marginally sympatric or para- patric with M. radiata and M. fascicularis (Food- en. 1980, p. 4). Ecologically, M. assamensis, M. thihetana, M. arctoides, and M. nemesthna gen- erally prefer primary broadleaf evergreen forest, whereas M. mulatto, M. radiata, and M. fascicu- laris generally prefer secondary or deciduous for- est and disturbed habitats (Fooden, 1982b, p. 574). Groups of M. mulatto and M. assamensis have been observed in close proximity in eastern Nepal (one locality; J. A. McNeely in Fooden, 1982a, p. 26), West Bengal. India (two localities; Khajuria, 1966, p. 284; Mukherjee et al., 1995, p. 30), and western Thailand (one locality; Eudy, 1979, pp. 92, 97, 199). However, these apparently were ca- sual contacts, and no mixing of M. mulatto and M. assamensis groups was observed at any of these localities. M. mulatto and M. assamensis also reportedly coexist at one locality in Bangla- desh (Feeroz et al., 1995, p. 76); no further infor- mation is available concerning this interspecific association. In China, the local distribution of M. mulatto tends to be negatively correlated with that of M. thihetana, which generally inhabits higher eleva- tions (Wada et al., 1986, p. 93). However, three mixed groups have been reported in Zhejiang Province (Kang Ximin. zmnh, pers. comm., 24 October 1985), and another possibly mixed group has been reported in Jiangxi province (David, 1875, vol. 2, p. 256). In the Zhejiang groups (Zhoucun, January 1985, two groups; Zhidaikou, August 1985, one group), most members were M. mulatto. In the possibly mixed group in Jiangxi (Kuatun, ca. 20 km northwest, 8 October 1873), 10 individuals were M. thihetana, and one may have been M. mulatto. In Tripura, eastern India, a group of M. mulatto and a group of M. arctoides have been observed 66 FIELDIANA: ZOOLOGY Table 21. Intcrgroup conlacl behavior reported in Mucaca iiiiilattci. Larger Frequency of group displaces contacts Mood of contacts Habitat . smaller group Refer- Sample area Common Rare Peaceful Hostile ences' Pakistan Forest + + + 1 India Himaehal Pradesh Temple + 2 Rural + + 3 Delhi Fort site + + 4 Uttar Pradesh Aligarh Temple + + + + 5 Asarori Forest Forest + + + + 6 Bareilly Roadside + 7 Dehra Dun Rural + + 4- + 8 Haldwani area Forest + + 9 Kaukori- Rural + 10 West Bengal Swamp + 11 China: Henan Forest + 12 Provisioned introduced populations (U.S.A.) Puerto Rico Cayo Santiago Island + + + + 13 La Cueva I. Island + + 4- + 14 South Carolina Morgan 1. Island + + + 15 ' Key to references: /. Melnick et al., 1984, p. 237: Goldstein & Richard, 1989, p. 563. 2. Edwin & Chopra, 1984, p. 312. 3. Wada, 1984, p. 489. 4. Malik et al.. 1984, p. 315. 5. Southwick & Beg. 1961, p. 391: Southwick. 1962, p. 438; Southwick el al.. 1965. pp. 137. 142. 6. Lindburg. 1971, p. 37; 1977b. p. 241; Makwana. 1979b. p. 920. 7. Mukherjee, 1969. p. 49. 8. Lindburg. 1969. p. 1177; 1971. p. 38. 9. Neville. 1968b. p. 118; 1968c. p. 15. 10. Jay, 1963. p. 278. //. Mandal. 1964. p. 161. 12. Qu et al.. 1993. p. 616. 13. Altmann. 1962. p. 371: Boelkins & Wilson, 1972, p. 130; Hausfater, 1972, p. 81; Sade et al., 1977, p. 260; Lauer, 1980. p. 477. 14. Marsden. 1968. p. 241; 1973. p. 248; Vessey. 1968. p. 230: Drickamer. 1975. p. 28. 15. Judge & de Waal. 1994, p. 65. - Includes one mixed rhesus-langur group. 10 to 150 m from one another, peacefully feeding on the ground (Mukherjee, 1977. p. Ill; 1982, p. 75). In Bangladesh, M. mulatta M. and nemestrina are listed as sympatric at five localities (Feeroz et al., 1995, p. 76). M. mulatta and M. radiata have been reported in mixed-species groups in peninsular India at three close-lying localities near the 1,000-km-long boundary between the geographic ranges of these two species (Fooden et al., 1981, p. 465; Koyama & Shekar, 1981, p. 248). One of these mixed-spe- cies groups included one M. mulatta female and three M. radiata males; these four monkeys were ca. 50 m from a larger group of M. mulatta. An- other mixed group included at least 22 M. mulatta individuals and one M. radiata male; two addi- tional unseen M. radiata individuals reportedly were also associated with this group. The third mixed group included 18 M. mulatta individuals and one M. radiata male. At another nearby lo- cality, a group of 20 M. mulatta individuals re- mained within 10 to 50 m of a group of 22 M. radiata individuals for ca. 45 min (Fooden et al., 1981, p. 472); neither mixing nor overt interaction occurred between these two groups. The boundary between the geographic ranges of M. mulatta and M. fascicularis extends ap- proximately 2,000 km across the Indochinese pen- insula. Although direct contact between these two species has not been reported, a few morpholog- ically intermediate specimens have been collected near the interspecific boundary (Fooden. 1997, fig. 3). This probably indicates that limited hy- bridization has occurred between M. mulatta and M. fascicularis. Hybridization between M. mulat- ta and M. fascicularis has been reported in mixed- species groups in Xianggang (= Hong Kong) that are the result of human introduction (Southwick & Southwick, 1983, p. 19; Southwick &. Manry, 1987, p. 48; Burton & Chan, 1996, p. 395). FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 67 In summary, divergent habitat preferences ap- parently limit contact between M. mulatto and M. assamensis, M. thihetana, M. antoides, and M. nemestrina. In marginal habitats, M. miilatta may encounter these four species; such contacts ap- parently are not hostile, and they may result in the formation of mixed groups. The habitat prefer- ences of M. mulatto are similar to those of M. radiata and M. fosciculahs. Along the boundary between the geographic ranges of M. mulatto and M. radiata, nonhostile interspecific contact and mixed-species groups have been observed; along the boundary between the ranges of M. mulatto and M. fosciculahs, morphological evidence in- dicates the occurrence of occasional hybridiza- tion. iNTiiRGiiNERic — In northern and peninsular In- dia, the geographic range of M. mulatto broadly overlaps that of the Hanuman langur, Semnopithe- cus entellus, and frequent contacts between these two species have been observed. In antagonistic encounters, M. mulatto usually is more aggressive and displaces S. entellus (Makwana, 1979b, p. 920; Pirta, 1984, pp. 274-279; Lindburg, 1971, p. 43; Ross et al., 1993, p. 162; Mathur, 1996, p. 360); however, at one locality groups of each spe- cies defended territories against incursion by the other species (Mathur, 1982, p. 12), and at another locality a male S. entellus successfully drove off a male M. mulatto (Neville, 1968c, p. 16). Peace- ful encounters that have been reported include groups of M. mulatto and S. entellus feeding to- gether in the same tree and in the same cultivated field (Oboussier & von Maydell, 1960, p. 144; Jay, 1965, p. 212; Lindburg, 1971, p. 43; Prater, 1980, p. 37; Mathur, 1982, p. 12) and interspecific play by infants and juveniles (Manohar & Mathur, 1992, p. 114). Mixed-species groups also have been reported; these usually are composed of one or two M. mulatto individuals in a group of S. entellus (Jay, 1963, p. 274; 1965, pp. 200, 212, 249; Roonwal & Mohnot, 1977, p. 264; Singh & Sen, 1977-78, p. 136; Mathur & Lobo, 1990, p. 308); in at least two of these groups, the M. mu- latto individuals were dominant to the S. entellus individuals (Jay, 1963, p. 274; Roonwal & Mohn- ot, 1977, p. 264). In a mixed-species group that included 40 S. entellus individuals and eight M. mulatto individuals, an adult female 5. entellus nursed an infant M. mulatto, and a young male S. entellus carried an infant M. mulatto on his back (Das & Sharma, 1981, p. 496). In northeastern India. M. mulatto is sympatric with the golden langur, Tracliypithecus geei. In- terspecific behavior between these monkeys ap- parently is variable, as they have been reported to maintain distance from one another (Khajuria, 1962b, p. 128); to casually encounter one another peacefully (Mukherjee & Saha, 1974, p. 337); to feed in close proximity, but with M. mulatto on the ground and T. geei in the branches overhead (Mukherjee, 1978b, p. 741); to feed together har- moniously in favored food trees (Oboussier & von Maydell, 1959, p. 106); and, in two instances, to contest for possession of a feeding site, with M. mulatto prevailing on both occasions (Mukherjee, 1978b, p. 742). In Madhupur National Park, Bangladesh, M. mulatto reportedly coexists peacefully with the capped langur, Tracliypithecus pileatus (Islam & Husain, 1982, p. 157; cf. Gittins & Akonda, 1982, p. 278). At this locality, M. mulatto generally is observed lower in the trees than T. pileatus. A commensal relationship between M. mulatto and two species of deer. Axis axis and Muntiacus muntjok, has frequently been reported in India (Champion, 1927, p. 201; de Poncins, 1935, p. 846; Khajuria, 1962a, p. 122; Mandal, 1964, p. 157; Mukherjee & Gupta, 1965, p. 146; Hen- drichs, 1975, p. 171, Sanyal, 1983, p. 3; cf. Lind- burg, 1971, p. 42). The deer move in close asso- ciation with M. mulatto and feed on fruits, leaves, and twigs dislodged by activity of the monkeys in branches overhead. The deer apparently also ex- ploit the monkeys' alarm calls to avoid approach- ing predators (Mukherjee & Gupta, 1965, p. 146; Sanyal, 1983, p. 3). One observer reports that M. mulatto sometimes leaps down from branches and rides on the back of deer (Mandal, 1964, p. 158). In northern India, M. mulatto individuals were observed to chase a jackal, Canis aureus, and fre- quently to chase away crows, Corvus sp., that were competing for provisioned food (Lindburg, 1971, p. 43). Reproduction Seasonality Matings and births are strongly seasonal in nat- ural populations of M. mulatto (Table 22). At lo- calities widely dispersed across the geographic range of this species, reported matings consis- tently peak in the fall and winter, and births peak in the spring and summer. At four sample areas in India, a second, minor birth peak in the fall 68 FIELDIANA: ZOOLOGY Table 22. Mating and birth periods reported for natural populations of Maccica miilatta. Approximate Mating Refer- Birth Refer- Sample area latitude (N) period ences' period ences' Afghanistan 35° (No data) Apr.-Nov. / Pakistan 34° Aug.-Nov. 2. 3 Mar.-May 2. 4 India Himachal Pradesh 31° Aug.-Nov. 5, 6 Mar.-May- 5 Delhi 28° Sept.-Feb. 7 Mar.-July (major) Oct. (minor) H Rajasthan 27° Oct. 9 Mar. -July (major) Sept. -Oct. (minor) 9. 10 Uttar Pradesh 29° Sept.-Feb.' 11, 12 Mar. -July (major) Sept. (minor) II. 13 Calcutta 23° Dec. 14 (No data) — Sundarbans 22° (No data) — Apr.-May (major) Sept. -Oct. (minor) 15 Nepal 28° Oct.-Feb. 16 Apr.-Aug. 16. 17 Vietnam 20° (No data) — Summer, primarily^ IS China Hainan 18° Nov.-Mar. 19 Apr.-Aug. 19 Xianggang (= Hong Kong)^ 22° (No data) — June-July, peak'' 20 Guangxi 23° Nov. -Jan. 21 Apr.-Aug. 21 Yunnan' 22° Sept.-Jan. 22 Mar.-June 22 Hubei 31° ?Jan.-Mar.« 23 July-Aug., peak 23. 24 Henan 35° Sept. -Nov. 25 Mar.-May 25 ' Key to references: /. Puget. 1971, p. 200. 2. Hingston, [1920], p. 245; Pearl et al., 1987, p. 36. 3. Igbal & Rub. 1980, p. 214; Rab et al., 1991, p. 221. 4. Roberts, 1977, p. 87, 5. Dodsworth, 1914, p. 730. 6. Heape, 1894, p. 414. 7. Malik & Johnson, 1992. pp. 26-27. 8. Malik et al., 1984. p. 314; John.son et al., 1993, p. 68. 9. Wolfe & Mathur, 1988, p. 538. 10. Prakash, 1962, p. 85; Ojha, 1983, p. 75: Singh. 1989. p. 140; Mathur, 1994. p. 132. //. Neville. 1968a, p. 772; 1968c, p. 15. 12. Southwick et al., 1965, p. 151; Siddiqi & Southwick. 1980, p. 54 Lindburg, 1969, p. 1177; 1983, p. 46. 13. Heape, 1897. p. 137; Nolle, 1956, p. 181; Southwick et al., 1961b, p 705; Mukherjee. 1969. p. 54; Lindburg. 1971. p. 77; Pirta & Singh. 1981, p. 342; Strum & Southwick, 1986, p 953. 14. Saha. 1974, p. 211. 15. Mandal. 1964, p. 159. 16. Teas, 1984, p. 242. 17. Marriott, 1988, p. 129. 18 Dang, 1983, p. 1283. 19. Jiang et al., 1988a, pp. 106-107. 20. Burton & Chan, 1996, p. 402. 21. Wang et al. 1996, pp. 267-268, 271. 22. Editors, 1989, p. 144. 23. Poirier, 1985, p. 298. 24. Poirier & Hu, 1983, p. 387. 25 Qu et al., 1993. p. 614. - Excludes questionable report: Aug.-Sept. (J. E. T. Aitchison in Heape, 1894, p. 414; 1897. p. 137). ' Excludes rare matings observed in April (Southwick et al., 1965. p. 151; Lindburg, 1983, p. 46). ■" Some neonates reportedly were also observed in other seasons. ■^Population probably introduced (Herklots, 1951, p. 83); possibly includes some interspecific hybrids. *> "[Blirths range from mid-winter to early autumn." ' Data probably were derived mainly from a captive population; also see Zeng et al., 1983, p. 155. ■* Inferred from unspecified evidence. also has been reported; this presumably implies an unreported minor mating peak in the spring. The occurrence of a minor birth peak in the fall may account for the somewhat protracted birth seasons that have been reported in Afghanistan and Vietnam. There is no evidence that births in natural populations living at higher latitudes occur later than in those living at lower latitudes (Table 22). as previously suggested by data derived pri- marily from translocated captive populations (Van Horn, 1980, p. 192; cf. Lindburg, 1987, p. 197). In captive groups of M. niulatta housed outdoors in the northern hemisphere, reproductive season- ality is generally similar to that in natural popu- lations (Harrison, 1980. p. 271; Van Horn, 1980, p. 183; Curie-Cohen et al., 1983, p. 129; Goo & Fugate, 1984, p. 67; Small & Smith, 1986, p. 293; Taub & Mehlman. 1989. p. 164; Bercovitch, 1992, p. 275; Ouyang & Ma. 1992, p. 14; Lehman et al., 1994, p. 120; Johnson & Kapsalis, 1995b, p. 272); captives in the southern hemisphere ex- hibit reversed cycles, generally mating in March- August and giving birth in September-February, in accord with southern hemisphere seasons (Hartman, 1931, p. 135; Strahan et al., 1973, p. 385; Coimbra-Filho & Maia, 1977, p. 75; Bielert & Vandenbergh, 1981, p. 231; de Faaria & Guer- ra. 1985, p. 187). Captives housed indoors tend to FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 69 Tabi.h 23. Age of sexual maturity reported for natural populations of Macaca mulatla. Estimated age (vr) at Approxi- mate latitude (N) Provi.sioned Adolescent _ sample size first fertile copulation' Sample area Youngest age Typical age References- Females Pakistan. N 34° No 6 4.5 5.5 1 India: Delhi 28° Yes 13 2.5 3.5 2 Vietnam 20° 7 7 (1.5, puberty) — 3 China: Hainan 18° Yes 7 3.5 4.5 4 China: Henan 35° Yes <24 Males 4.5' 5.5' 5 Pakistan, N 34° No 7 — 6.5 1 Vietnam 20° 7 7 — >1.5 3 China: Henan 35= Yes <24 4.5^ 6.5^ 5 ' Durins the mating season, all individuals are approximately midway between their birth anniversaries. - Kev to references: /. Melnick et al.. 1984. p. 230. 2. Mahk & Johnson. 1992, p. 27. 3. Dang, 1983, p. 1283. 4. Jiang et al.. 1988a. p. 112: 1991. p. 211. 5. Qu et al., 1993, p. 614. ' Reported as 4 to 5 years. ^ Reported as 4.5 to 6 years. lose reproductive seasonality (Valerio et al., 1969a, p. 66; Michael & Zumpe, 1976. p. 308; Hemdon et al.. 1985. p. 735). Sexual Maturation In natural populations, females may become sex- ually mature (i.e.. capable of engaging in fertile copulations) as early as age 2.5 years (Table 23); more commonly, however, female sexual maturity is not achieved until age 3.5 to 5.5 years. Males in natural populations apparently become sexually mature later than females, perhaps usually at age 6.5 years. In captive colonies, most females be- come sexually mature at age 3.5 years (Vanden- bergh. 1973. p. 7; Rawlins & Kessler, 1986c. p. 52; Ouyang & Ma. 1992. p. 14; Bercovitch & Ber- ard, 1993. p. 105); reproductive capability in cap- tive females probably requires a minimum body weight of ca. 4 kg (Bercovitch et al., 1998, p. 137). Although captive males are capable of fertilizing females at age 3.5 years (Catchpole & van Wage- nen. 1975, p. 133; Stem & Smith, 1984, p. 24; Bernstein et al., 1991, p. 33), males in the free- ranging Cayo Santiago colony usually do not par- ticipate in breeding activity before age 4.5 or 5.5 years (Conaway & Koford. 1964. p. 586; Sade, 1968, p. 25; cf. Ouyang & Ma, 1992. p. 14). In captivity, sexual maturation is influenced by diet, housing, group density, and social rank (Zimmer- mann et al., 1975, p. 298; Wilen & Naftolm, 1976. p. 358; Schwartz et al., 1988, p. 240; Wilson et al.. 1988, p. 2655; Bercovitch & Berard, 1993. p. 105; Mann et al., 1998, p. 497). In 59 captive females, the first menstrual bleeding (menarche) preceded sexual maturity by ca. 1 year (Bercovitch & Goy. 1990. p. 64). The mean age at menarche in this group was 29.8 ± 0.6 months (SEM; extremes, 20.1-42.4 months); in another colony, the mean age at menarche was 9 months less (20.6 months, n = 25; extremes, 13.9-25.3 months) (van Wagenen. 1972. p. 25). In males, the testes usually are scrotal at birth (Wislocki, 1933a, p. 134; 1933b, p. 234). Within a few months they ascend to the inguinal region and remain there until age 3 to 4 years, when they return to the scrotum (Schultz, 1933, p. 26; Sade, 1964, p. 175; Goy et al., 1982, p. 288). During the next year or two, the testes may temporarily reascend to the inguinal region, but about age 5.5 years they become permanently scrotal. At or be- fore the age of permanent testis descent, males generally emigrate from their natal group, usually near the beginning of a mating season (Koford, 1966, p. 5; Sade, 1968, p. 26; Missakian. 1973a, p. 228; Drickamer & Vessey, 1974, p. 361; Sade et al., 1977, p. 256; Colvin, 1983, p. 161; Jiang et al.. 1988a. p. 1 10; Kaplan et al.. 1995. p. 231). Subsequently, males also generally emigrate again as adults (Lindburg, 1969. p. 1177); in one cap- tive free-ranging colony, the average group tenure of postjuvenile males was ca. 2 years (Vessey & Meikle. 1987. p. 289). Females apparently rarely emigrate from their natal group; in one captive free-ranging colony, fewer than 3% of females 70 FIELDIANA: ZOOLOGY emigrated during a 3-year observation period (Koford. 1966. p. 6). Sexual Skin In postjuvenile females and males, regions of the skin undergo intermittent swelling and/or red- dening (Darwin. 1871. p. 279: 1876. p. 19; An- derson, 1879, p. 58; Langley & Shenington, 1891, p. 284; Heape, 1896, p. 202; 1897, p. 139; Corner, 1923, p. 82; Allen, 1926, p. 226; 1927, p. 9; Coll- ings, 1926, p. 272; Zuckerman, 1930, p. 702; Stewart, 1933, p. 29). In captive females, the ear- liest manifestation of this "sexual skin" (Langley & Sherrington. 1891. p. 290) is the development during the second or third year of life of a pair of pinkish pubic swellings (Haitman. 1928c, p. 182; 1932, p. 20; Zuckerman et al., 1938, p. 385; van Wagenen, 1950, p. 26; Eckstein & Zuckerman, 1956, p. 139; Hadidian & Bernstein, 1979, p. 436); the location of these swellings is similar to that of the scrotum in a male. After approximately 2 weeks, these pubic swellings subside and the first menstrual bleeding occurs. Within the next few months, the fully developed sexual skin of puberty appears; this consists of a large bilobed blister-like pubic swelling, from which a less prominent edematous midline swelling extends posteriorly as far as the sides of the vulva. During the subsequent menstrual cycles of adolescence, a period that may extend up to 2 years, the sexual skin of puberty is gradually transformed, becom- ing less acutely swollen, redder, and more exten- sive; the cyclically edematous area of sexual skin often extends beyond the vulva and anus to the root of the tail, over the buttock area and posterior surface of the thighs, and over the iliac region. In sexually mature females (beginning at age ca. 3.5-5.5 years), the primary cyclical manifestation of sexual skin is reddening rather than swelling; in addition to the buttocks and adjacent regions, the areas of redness frequently include the face and nipples. During the menstrual cycle, the red color of the sexual skin generally reaches maxi- mum intensity near the day of ovulation (Anon- ymous, 1973, p. 9; Czaja & Bielert, 1975, p. 587; Czaja et al.. 1975, p. 1681; cf. Zuckerman, 1930, p. 728; Zuckerman et al., 1938, p. 389; Carpenter. 1942a, p. 131); this cyclic variation apparently may be diminished in older females (Hartman, 1932, p. 21; Stewart, 1933. p. 29; Valerio et al.. 1969b, p. 284). Sexual skin color reportedly is brighter during the mating season than during the nonmating season (Gordon & Bernstein, 1973, p. 223; Baulu. 1976. p. 487; Rab et al.. 1991, p. 221; cf. McCann, 1933b, p. 810). The bright red color of the sexual skin is retained during pregnancy and frequently during lactation (Heape. 1894. p. 456; 1897. p. 140; Hartman. 1928b. p. 539; 1928c. p. 187; 1932. p. 21; Tinklepaugh & Hart- man. 1930, p. 66; Bielert et al., 1976. p. 182). In pubertal and adolescent males, red sexual skin develops in approximately the same posterior and facial regions as in females, but there is no pubertal swelling of the sexual skin in males (Zuckerman, 1937, p. 327). As in females, the red color in males is brighter during the mating sea- son than during the nonmating season (Sade, 1964, p. 179; Koford, 1965, p. 165; Lindburg, 1971, p. 91; 1983. p. 47; Gordon & Bernstein, 1973, p. 223; Baulu, 1976, p. 485; Bielert & Van- denbergh. 1981. p. 231; Rab et al., 1991. p. 221). Subcutaneous fat deposits also vary seasonally in sexually mature males, increasing prior to the mating season and decreasing during the mating season (Lindburg, 1977b. p. 247; Bernstein et al.. 1989, p. 253; Bercovitch, 1992, p. 277; Zeng, 1992, p. 22; Bercovitch & Nurnberg, 1996, p. 63). Menstrual Cycle In three studies of 4.626 menstrual cycles (ca. 600 females), modal cycle lengths were 27, 28, and 28/30 (bimodal) days (Valerio et al., 1969b, p. 286; Anand Kumar et al.. 1980. p. 38; Dailey & Neill, 1981, p. 562); extreme cycle lengths were 6 and 237 days, but few cycles were longer than 50 days, and longer cycles (?anovulatory pe- riods) generally occuned during the summer .sea- son of infrequent matings. The modal durations of menstrual flow reported in two of these studies were 3 and 4 days (3,370 cycles, 490 females); extreme durations in both studies were 1 and 1 1 days, and mean durations were 3.5 and 3.9 days. Estrus During the mating season, female sexual activ- ity (estrus) is cyclical in natural and seminatural groups of A/, mulatta (Carpenter, 1942a, p. 117; Conaway & Koford, 1964, p. 584; Kaufmann, 1965. p. 501; Loy, 1971. p. 2; Lindburg. 1971. p. 94). Within groups, the estrous periods of individ- ual females are asynchronous, and male sexual activity apparently is not cyclical. In five studies FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 71 of ca. 160 females, length of the estrous period averaged 8 to 12 days (Wilson & Gordon, 1980, p. 639; Lindburg, 1983. p. 50; Berman et al., 1993, p. 393; cf. Loy, 1970, p. 287), and length oi' the intervening period of sexual inactivity av- eraged 19 to 22 days (not reported in two studies). The midpoint of a female's estrous period approx- imately coincides with her day of ovulation (ca. 11-14 days after onset of previous menstruation), which is the day on which she generally partici- pates in the maximum number of ejaculatory cop- ulations (Ball & Hartman. 1935, p. 118; Michael, 1965, p. 596; Valerio et al., 1969b, p. 292; Wallen, 1990, p. 236; Michael & Zumpe, 1993, p. 226). During three years of observation, 34 to 40 fe- males in a seminatural group averaged 2.2 estrous periods per mating season (extremes, 0-5 periods per season) (Kaufmann, 1965, p. 504), and a sim- ilar frequency of estrous periods per season was observed during a 10-month study of 35 females in a natural population (Lindburg, 1971, p. 95). In two groups, 63% and 77% of the females ap- parently were impregnated during their first es- trous periods of the mating season (Conaway & Koford. 1964, p. 585; Lindburg, 1971, p. 95); most of these females apparently also had subse- quent postconception estrous periods during the same mating season (cf. Altmann, 1962, p. 389; Lindburg, 1983, p. 50). In captive females, estrous cycles may be attenuated (Tinklepaugh, 1933, p. 336; Rowell, 1963, p. 198; Kuehn & Young, 1965, p. 688; Johnson & Phoenix, 1978, p. 167; Kev- eme, 1981, p. 119). Consortship In natural and seminatural groups, temporary copulatory associations (consortships) are formed between estrous females and their male partners (Carpenter, 1942a, p. 118; Altmann, 1962, p. 393; Kaufmann, 1965, p. 502; Southwick et al., 1965, p. 151; Vandenbergh & Vessey, 1968, p. 73; Lind- burg, 1971, p. 91; Brereton, 1981, p. 419; Hill. 1987, p. 443; Rab et al., 1991. p. 222; Berard et al., 1993, p. 483; cf. Manson, 1996b, p. 156; 1997, p. 353). The duration of these consortships reportedly varies from <25 minutes to 1 1 days; a duration of 1 to 2 days probably is typical (South- wick et al., 1965, p. 152; Hill, 1987, p. 446). A female usually consorts with more than one male dunng each of her estrous periods; in a natural group, females changed consort partners in 24 of 38 estrous periods observed (Lindburg, 1971, p. 94; cf. Carpenter, 1942a, p. 138). Over the course of one mating season, one female was observed to consort with 11 of 12 group males, and one male was observed to consort with 1 8 of 34 group females (Conaway & Koford, 1964, p. 582); dur- ing a single day, a female may consort with more than one male, and a male may consort with more than one female (Southwick et al., 1965, p. 152; Vandenbergh & Vessey, 1968, p. 73). Although either sex may initiate a consort relationship (Kaufmann, 1965, p. 502; Lindburg, 1971, p. 92; 1983, p. 54; Rab et al., 1991. p. 222), in one study of 53 consortships, male partners were responsi- ble for maintaining proximity approximately twice as frequently as females (Hill, 1987, p. 446). The consortships of high-ranking individu- als usually are conducted near the center of their group; those of low-ranking individuals may be conducted several hundred meters from other group members (Lindburg, 1971, p. 92; Rab et al., 1991, p. 222). In a seminatural population, fe- males occasionally have been observed to move temporarily into another group and to form a con- sortship there (Carpenter, 1942b, p. 154; Brereton, 1981, p. 419). Copulatory Behavior Copulations, like consortships. may be initiated by either sex (Hinde & Rowell. 1962. p. 16; Har- low, 1965, p. 235); in laboratory pair tests with multiple copulations, the frequency of female ini- tiations in second copulations (ca. 35%) tends to exceed that in first copulations (ca. 15%) (Michael & Zumpe, 1970, p. 176; Dixson et al., 1973, p. 42). Near the beginning of a copulation, the fe- male typically presents by turning her perineal re- gion toward the male, and the male mounts dor- soventrally by grasping the female's waist and shanks with his hands and feet, respectively (Car- penter, 1942a, p. 132; Altmann. 1962, p. 374; Southwick et al., 1965, p. 152). Although ejacu- lation in M. mukitta occasionally is accomplished in a single mount (Kaufmann, 1965, p. 502; Mi- chael et al., 1973, p. 249; Shively et al., 1982, p. 376; Curie-Cohen et al.. 1983, p. 129; Manson, 1996a, p. 1225), usually a series of mounts, sep- arated by dismounts, is required to complete a copulation. Each mount includes one to 15 intro- missive thrusts and usually lasts less than 1 min (Prakash, 1962, p. 84; Southwick et al., 1965, p. 152; Michael & Saayman, 1967b, p. 462; Shively et al., 1982, p. 376; Lindburg, 1983, p. 51). A 72 FIELDIANA: ZOOLOGY copulation may include as many as ca. 100 mounts and dismounts (Carpenter, 1942a, p. 133) and may last up to 30 min or, rarely, 1 hr (Prak- ash, 1962, p. 84; Kaufmann, 1965, p. 502; South- wick et al., 1965, p. 152; Lindburg, 1971, p. 94; Manson, 1996a, p. 1223); in laboratory pair tests with multiple copulations, the number of mounts per copulation and the duration of copulations tend to increase in successive copulations (Kuehn & Young, 1965, p. 688; Michael & Saayman, 1967b, p. 463; Missakian et al., 1969, p. 234). The copulation rate of estrous females in a seminatural group has been estimated to be ca. 0.4/hr (Man- son, 1992, p. 412). In caged animals, the rate of copulatory behaviors apparently is greater at 0900 than at 2100 (Chambers et al., 1982, p. 38), and copulation rarely occurs late at night (Erffmeyer, 1982, p. 246). Following copulation, coagulated semen often is visible on the perineum of the fe- male ("vaginal plug," Hartman, 1932, p. 39; cf. Carpenter, 1942a, p. 119; Altmann, 1962, p. 375; Mastroianni & Manson, 1963, p. 1026; Vanden- bergh & Vessey, 1968, p. 73; Blandau, 1973, p. 295; Lindburg, 1983, p. 47). Dominance Rank and Reproductive Success The relationship between male copulation fre- quency, fertilization success, and dominance rank in M. mulatto has been studied in natural groups, free-ranging seminatural groups, and compound- housed groups (cf. Bercovitch, 1997, p. 248). Ev- idence from natural groups suggests that male copulation frequency and fertilization success are positively correlated with dominance rank, and evidence from seminatural and compound-housed groups equivocally tends in the same direction (cf. Paul, 1997, p. 345; Bercovitch & Numberg, 1997, p. 1703; Rifkin et al., 1999, p. 93). Field studies at two localities in India and one locality in Pak- istan indicate that a male's copulation frequency and number of female partners are correlated with his dominance rank (Southwick et al., 1965, p. 152; Igbal & Rub, 1980, p. 214; Lindburg, 1983, p. 52; Rab et al., 1991, p. 222); preliminary anal- ysis of blood protein data from two groups at the Pakistan locality suggests that most of the infants bom during three years were fathered by the alpha male of each group (Melnick & Hoelzer, 1 996, p. 430). Although most studies of the seminatural Cayo Santiago population similarly indicate that male copulation frequency is positively correlated with dominance rank (Carpenter, 1942b, p. 156; Koford, 1963, p. 150; Conaway & Koford, 1964, p. 582; Kaufmann, 1965, p. 507; Sade, 1980, p. 182; McMillan, 1982b, p. 312; Chapais, 1983, p. 219; Hill, 1987, p. 445; Manson, 1992, p. 412; 1996a, p. 1223; Berard et al., 1993, p. 485), a few studies of this population have failed to find such a correlation (Loy, 1971, p. 8; McMillan, 1982a, p. 207; Berard, 1993, p. 298; 1999, p. 163); fac- tors that may tend to bias these observations are the relatively low visibility of low-ranking males (Drickamer, 1974a, p. 119) and the relatively low mating success of low-ranking subadult males (McMillan, 1982a, p. 207). Available results of genetic paternity tests in the Cayo Santiago pop- ulation are inconclusive concerning a correlation between fertilization success and dominance rank (Sade, 1980, p. 182; Berard et al.. 1993, p. 484; 1994, p. 184; Berard & Schmidtke, 1996, abstract no. 459). Male reproductive success in the Cayo Santiago population may be affected by the ab- normally large group size in this provisioned pop- ulation and the high ratio of sexually mature males to sexually mature females (Tables 14,15; Manson, 1992, pp. 407, 414). Most, but not all, evidence from studies of compound-housed groups suggests that copulation frequency may be correlated with dominance rank (Gordon et al., 1976, p. 239; Ruiz de Elvira et al., 1982, p. 829; Shively, 1982, p. 377; Wilson et al., 1982, p. 24; Curie-Cohen et al., 1983, p. 130; Stem & Smith, 1984, p. 29; Jebavy et al., 1994, p. 202; Bercov- itch & Niimberg, 1996, p. 61) and that fertiliza- tion success also may be correlated with domi- nance rank (Duvall et al., 1976, p. 29; Smith, 1980, p. 248; 1981, p. 87; 1993. p. 474; 1994b, p. 234; Berenstain et al., 1981, p. 1058; Curie- Cohen et al., 1983, p. 133; Stem & Smith, 1984, p. 27; Smith & Smith, 1988, p. 557; Bercovitch & Numberg, 1996, p. 61; Bercovitch, 1997, p. 248). In seminatural and compound-housed groups, dominant males and females often harass and dismpt the copulations of subordinate group members (Hemdon et al., 1986, p. 95; Manson, 1994, p. 136; 1996a, p. 1224). Inbreeding Blood-protein evidence from five groups in Pakistan indicates that close inbreeding is rare in natural populations of M. mulatta (Melnick et al., 1984, p. 235); the opportunity for such inbreeding is obviously limited by the tendency for pubertal males to emigrate from their natal groups (see FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 73 above). In the Cayo Santiago free-ranging popu- lation, relatively few mother-son and brother-sis- ter copulations have been observed (Kaufmann, 1965, p. 508; Sade, 1968, pp. 18, 33, 36; 1972, p. 392; Loy, 1971, p. 12; Missakian, 1973a, p. 230; Sade et al., 1984, p. 213; Manson & Perry. 1993, p. 340; Berard & Schmidtke, 1996, abstract no. 459), and the same is true of the Yerkes com- pound-housed population (Wilson, 1981, p. 475). Genetic paternity tests of compound-housed groups at Davis. California, indicate that matrilin- eal inbreeding is rare (two of 132 infants) but that patrilineal inbreeding, except for father-daughter mating, is common (Smith, 1982, p. 448; 1986a, p. 219; 1986b, p. 864; 1995, p. 34). (Silk et al., 1993, p. 97). Gestation length aver- ages greater in young primaparous mothers (ca. 169 days) than in old multiparous mothers (ca. 162 days) (Hartman, 1932, p. 53; Silk et al., 1993, p. 98). In an experimental study of nutritional ef- fects, gestation length was ca. 8.5 days less in pregnant females fed a high-protein diet than in those fed a low-protein diet (Riopelle & Hale, 1975, p. 1173); this suggests that gestation length in unprovisioned natural populations may be greater than gestation length in provisioned cap- tive populations. Parturition Nonreproductive Sexual Behavior Sexual behavior in M. mulatta may serve non- reproductive functions (Carpenter. 1942a, p. 132). Male and female homosexual mounting has been observed in natural, seminatural, compound- housed, and caged groups (Kempf, 1917, p. 134; Carpenter, 1942b. p. 150; Altmann, 1962. p. 383; Gordon & Bernstein, 1973, p. 224; Gordon et al.. 1978, p. 626; Akers & Conaway, 1979, p. 66; Ig- bal & Rub, 1980, p. 214; Loy & Loy, 1982, p. 308). Mounting of males by females also has been observed in seminatural and caged groups (Car- penter, 1942b, p. 152; Altmann, 1962, p. 383; Freedman & Rosvold, 1962, p. 26; Michael et al., 1974. p. 401). Male masturbation has been re- ported in natural groups (Prakash. 1962, p. 85; Lindburg. 1973. p. 146; Igbal & Rub, 1980, p. 214) and in seminatural groups, compound- housed groups, and caged individuals (Carpenter, 1942b. p. 152; Altmann, 1962, p. 375; Rowell, 1963, p. 196; Michael & Saayman, 1967a, p. 217). Gestation Length In a large laboratory colony of M. mulatta, the mean (±SD) gestation length was 165.4 ± 6.1 days for 1,067 viable female births and 166.2 ± 6.7 days for 1,115 viable male births (Shaughnes- sy et al., 1978, p. 132; cf. Hartman, 1928a, p. 15); in two other large colonies, mean gestation length was 166.5 days (n = 709; Silk et al., 1993, p. 97) and 168.4 days (n = 311; van Wagenen, 1972, p. 9). Reported minimum and maximum lengths of viable pregnancies are 133 days and 200 days Parturition is rarely observed in natural popu- lations of M. mulatta, even in those that have been closely studied (Southwick et al., 1965, p. 155; Lindburg, 1971, p. 77; Teas et al., 1981b, p. 580; Mathur, 1994, p. 132). Most newborn infants in closely studied natural populations are first seen early in the morning, which presumably implies that births usually occur at night. Of the five births and one stillbirth that have been partially or com- pletely observed during daylight hours (early morning to late afternoon), all apparently oc- curred while the mother was on the ground — in undergrowth in at least three cases (Lindburg, 1971, p. 77; Teas et al., 1981b, p. 581; Mathur, 1994, p. 132). From the first observed contraction to delivery of the afterbirth, the approximate du- ration of three of the live births was 15 min, 1 hr, and 0.5 to 2.5 hr. The placenta was eaten by at least two of the five mothers. Observed births in captive populations apparently are generally sim- ilar to those in natural populations with respect to timing, duration, and consumption of the placenta (Pocock, 1906. p. 562; Hartman. 1932. p. 52; Hin- de et al.. 1964, p. 613; Valerio et al.. 1969a, p. 72; Brandt & Mitchell, 1971. p. 199; Shaughnessy et al., 1978. p. 130; Rawlins, 1979, p. 432; Adachi et al., 1982, p. 585; Gibber. 1986. p. 121). The frequency of stillbirths was 5.5% (259 still- births/4.711 births) in four large captive colonies (Shaughnessy et al., 1978. p. 130; Scanlan et al., 1985. p. 363; Rawlins & Kessler. 1986c, p. 52; Small & Smith, 1986, p. 293). The frequency of twinning was 0.23% (13 twin pairs/5,561 live births) in three large captive colonies (Geissmann, 1990, p. 392; cf. Chalise & Ghimire, 1998, p. 1 1). 74 FIELDIANA: ZOOLOGY Table 24. Birth ' kveight (g) in laboratory-hou ised Macaca midatta. Female infant* Male infants Labora- Pregnancy information Refer- tory' Mean ± SD N Mean ± SD N ences- DOGYU 457.7 161 479.1 150 1 China' 430 ± 56.4 40 460 ± 58.8 40 2 CRPRC 470.4 ± 70.0 356 485.5 ± 72.0 353 3 LBI 476.0 ± 76.1 1.067 502.8 ± 73.4 1 1,115 4 LBI 474 ± 74 -250 504 ± 23 -250 Bred in laboratory 5 LBI 402 ± 87 -250 440 ± 69 -250 Received pregnant 5 LPP 46 L5 ± 82.6 116 484.1 ± 76.3 119 — 4 NLAC 414.6 ± 12.7 11 416.4 ± 17.5 7 6 WRPRC 464 ± 63 255 498 ± 66 255 — 7 ' DOGYU = Department of Obstetrics and Gynecology, Yale University School of Medicine. New Haven, CT; CRPRC = California Regional Primate Research Center, University of California, Davis, CA; LBI = Litton Bionetics. Inc.. Kensington. MD; LPP = Laboratory of Perinatal Physiology. National Institute of Neurological Diseases and Stroke. San Juan. PR; NLAC = National Laboratory Animal Centre. Central Drug Research Institute. Lucknow. India: WRPRC - Wisconsin Regional Primate Research Center. Madison. WI. - Key to references: 1. van Wagenen. 1972. p. 9. 2. Zeng, 1992. p. 18. 3. Silk et al.. 1993. p. 98. 4. Shaughnessy et al.. 1978. p. 131.5. Valerio et al.. 1969a. pp. 48. 67. 6. Maity & Rathore, 1998, p. 247. 7. Kemnitz, 1994, p. 220. ' Laboratory unspecified. Birth Weight, Infant Sex Ratio Birth weight in one large laboratory colony av- eraged 476.0 g in 1.067 female infants and 502.8 g in 1,115 male infants (Table 24); in five other large colonies, birth weight averaged somewhat less. The birth weight of inbred infants averages less than that of non-inbred infants (Smith, 1986b, p. 869). Judging from the relatively low weight of infants born to females that were pregnant when imported (Table 24), birth weight in natural populations may be less than in laboratory colo- nies. The male/female infant sex ratio was 1.16 in three natural populations of M. mulatta (n = 212 sexed infants) and 1.03 in 10 captive populations (n = 7,445 sexed infants) (Table 25; cf. Debyser, 1995, p. 955); neither of these ratios differs sig- nificantly from 1.00 (P > 0.10). The possibility that infant sex ratio in this species may be influ- enced by maternal dominance rank has been in- vestigated with equivocal results in four captive colonies. In two compound-housed colonies, high-ranking females tended to have female-bi- ased infant sex ratios (Simpson & Simpson, 1982, p. 440; 1985, p. 85; Small & Smith, 1985, p. 356; Small & Hrdy, 1986, p. 362; Gomendio, 1990, p. 369; Nevison et al., 1996, p. 127; Nevison, 1997, p. 287); the skew was statistically significant in one colony — at least in smaller matrilines — but not significant in the other colony. Conversely, in one large seminatural colony, high-ranking fe- males had significantly male-biased infant sex ra- tios (MeikJe et al., 1984, p. 179), and in another large seminatural colony, maternal rank and infant sex ratio were not significantly correlated (Ber- man & Rawlins, 1985, p. 332; Rawlins & Kessler, 1986a, p. 12; Berman, 1988, p. 313). Paternal dominance rank had no apparent effect on infant sex ratio in a compound-housed colony (Small & Smith, 1985, p. 359). Birth Rate, Infant Mortality Rate The mean annual birth rate (birthsAsexually ma- ture females) in natural populations of M. mulatta varies from 42.9% to 90.8% (Table 26); only three of 17 reported means are less than 63%. One of the three low outlier values is based on observa- tions of 14 Indian temple groups; the other two are based on observations of the two northernmost populations for which birth rate data are available (Pakistan. 34°03'N; China, 35°10'-35°17'N). At both northern localities, females usually produce infants only in alternate years (Qu et al., 1993, p. 619). In captivity, the mean annual birth rate varies from 65.0% to 76.7% in four free-ranging colonies (Vandenbergh. 1973, p. 7; Rawlins & Kessler, 1986c, p. 52, live births to females age ^4 years; Johnson & Kapsalis, 1995b, p. 273), and is 65.8% and 71.8% in two compound-housed colonies (Casebolt et al., 1985, p. 291; Goo, 1986, p. 75; cf. Litton & Izard. 1999, p. 74). The reported mean annual infant mortality rate, during the first year of life, was 7.0% in southern FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 75 Table 25. Infant sex ratio in Macaca mulatta. Infants Maleifemale Sex Locality or colony' N Females Males unknown ratio References- Natural populations India: Dehra Dun 23 10 10 3 1.00 / China: Longhushan 56 23 33 0 1.43 2 China: Nanwan 136 65 71 0 1.09 3 Totals 215 98 114 3 1.16 Captive populations^ Cayo Santiago, PR 1,407 665 720 22 1.08 4 CRPRC 815 399 398 18 1.00 5 DOGYU 311 161 150 0 0.93 6 La Parguera, PR 362 183 179 0 0.98 7 LBI 2,182 1,067 1,115 0 1.04 8 LPP 235 116 119 0 1.03 8 UCM 299 151 148 0 0.98 9 UMCG 169 83 83 3 1.00 10 WRPRC 1.616 808 808 0 1.00 11 YRPRCFS 97 37 55 5 1.49 12 Totals 7,493 3,670 3,775 48 1.03 ' CRPRC = California Regional Primate Research Center, Davis, CA; DOGYU = Department of Obstetrics and Gynecology, Yale University School of Medicine, New Haven, CT; LBI = Litton Bionetics, Inc., Kensington, MD; LPP = Laboratory of Perinatal Physiology, National Institute of Neurological Diseases and Stroke, San Juan, PR; UCM = University of Cambridge, Madingley, Cambridge, England; UMCG = University of Miami, Coral Gables, FL; WRPRC = Wisconsin Regional Primate Research Center, Madison, WI; YRPRCFS = Yerkes Regional Primate Research Center Field Station, Lawrenceville, GA. - Key to references: 1. Lindburg, 1971, p. 21. 2. Wang et al.. 1996, p. 268. 3. Jiang et al., 1988a, p. 111.4. Rawlins & Kessler. 1986a, p. 12. 5. Small & Smith, 1986, p. 295. 6. van Wagenen, 1972, p. 9. 7. Drickamer, 1974b, p. 76. 8. Shaughnessy et al., 1978, p. 131. 9. Nevison et al., 1996. p. 127. 10. Taylor, 1994, p. 244. //. Dyke et al.. 1986, p. 264. 12. Bourne & Bourne, 1975, p. 271. ' Cf. Arnold & Hay ward, 1998, p. 452. China, 15.5% to 18.0% in northern India, 16.0% to 29.0% in Nepal, and 28.5% to 31.5% in north- eastern China (Table 27). In a group of 19 pri- maparous captive females, the mortality rate (to age 6 months) of male infants (5/11) exceeded that of female infants (1/8) (Bercovitch et al., 1998, p. 138; cf. Dai et al., 1998, p. 30). When an infant dies, its mother often carries the corpse for several days, even after it has decayed; this has been observed in natural populations (Dods- worth, 1914. p. 730; Prakash, 1962, p. 85; Ne- ville. 1968c. p. 18; Mukherjee, 1969, p. 55; Lind- burg, 1971, p. 17; Taylor et al., 1978, p. 346; Shukla et al., 1984, p. 20) and in captive popu- lations (Yerkes, 1915, p. 404; Carpenter, 1942c, p. 199; Koford, 1965, p. 169; Sade, 1968, p. 21; Rawlins & Kessler, 1983, p. 170). Nursing, Weaning In two captive groups. 41 infants (age 0-6 weeks) nursed from the left nipple approximately 50% more than from the right nipple (Tomaszycki et al., 1998, p. 308). In natural populations, in- fants sometimes begin to take small amounts of solid food at age 2 weeks, and mothers begin to resist nursing attempts about 3 months later (Lind- burg, 1971, p. 84). The weaning process gradually intensifies over the next several months, but it may extend through the subsequent mating season and not be completed until the birth of the next infant, when the first infant is ca. 1 year old (Prak- ash, 1962, p. 85; Southwick et al., 1965, p. 156; Lindburg, 1971, pp. 63, 84). The weaning process in captive populations is generally similar to that in natural populations (Pocock, 1906, p. 569; Hartman, 1932, p. 22; Hinde et al., 1964, p. 637; Sade, 1968, p. 24; Vandenbergh & Vessey, 1968, p. 75; Gomendio. 1989, p. 452; Simpson & Tar- tabini, 1992, p. 31). An experimental study indi- cates that prolongation of the nursing period de- lays subsequent conception (Goo & Fugate, 1984, p. 67); the mean (±SD) interbirth interval of mothers of infants artificially weaned at age 6 76 FIELDIANA: ZOOLOGY Table 26. Annual birth rate (births/sexually mature females X 100) in natural populations of Macaca mitlotta. Study period No. of _ groups Annual birth rate (%) Refer- Sample area Mean Extremes ences' Pakistan Dunga Gali 1978-80 1 -50 — / India Delhi; Tughlaqabad- 1980-87 2-5 78.8 70.7-86.7 2 Uttar Pradesh Aligarh District 1961-85 8-21 83.7 75.4-90.4 3 1990-91 8 82.2 60.0-90.2 4 Chhatari-do-Raha- 1959-79 2 90.3 72.7-100.0 5 1990-91 2 80.7 80.0-80.8 4 Dehra Dun 1965-66 5 90.8 90.3-91.2 6 Northern India' Temple- 1981-91 14 42.9 — 7 Urban 1981-91 103 64.7 — 7 Village 1981-91 42 68.8 — 7 Village/pond 1981-91 60 64.2 — 7 Pond 1981-91 28 66.2 — 7 Roadside 1981-91 33 71.1 — 7 Canal side 1981-91 22 64.8 — 7 Forest 1981-91 70 64.1 — 7 Nepal Kathmandu Valley- 1975-78 12 63.0 51.0-73.0 8 China Hainan: Nanwan- 1978-90 2 -75 50.0-100.0 9 Guangxi: Longhushan 1988-95 6 75.4 45.0-100.0 10 Xianggang (= Hong Kong)-"* 1981 3 75.0 — 11 Henan: Jiyuan- 1987-88 2 46.9 30.1-66.7 12 ' Key to references: 1. Melnick et al., 1984, p. 343; Pearl & Goldstein, 1984. p. 203; Pearl et al., 1987, p. 38. 2. Malik, 1989a, p. 118. 3. Southwick & Siddiqi, 1988, p. 190; cf. 1994b, p. 56. 4. Imam & Yahya, 1995, p. 6. 5. Southwick & Siddiqi, 1983, p. 230. 6. Lindburg, 1971, p. 20. 7. Seth et al., 1992, p. 65. 8. Southwick et al., 1980, p. 166; Teas. 1983, p. 224. 9. Jiang Haisheng et al.. 1991, p. 213; 1994, p. 168. 10. Wang et al., 1996, p. 268. //. Southwick & Southwick, 1983, p. 22. J 2. Qu et al.. 1993, pp. 613, 616; cf. Southwick et al., 1996, p. 101. - Provisioned population. ' Seven states. ^Mixed-species groups; M. mulatta probably introduced (Herklots, 1951, p. 83; Marshall. 1967. p. 45). months was 382 ± 34 days (n = 153), whereas that of mothers of infants artificially weaned at age 12 months was 403 ± 63 days (n = 147). If a mother fails to produce a second infant during the following birth season, she may suckle the first infant for 2 years (Hartman, 1929, p. 157; Fleischman, 1963, p. 706). Menopause In a natural population in India, one female ap- parently became menopausal and ceased mating activity at an estimated age of >20 years (Malik & Johnson, 1992, p. 28); subsequent to meno- pause, this female survived for 4 years. In three closely observed captives, menopause occurred at ages 26.9, 27.6, and 29.0 years (van Wagenen, 1972, p. 26; van Wagenen & Simpson, 1973, p. 24; cf. Hodgen et al., 1977, p. 582; Davis, 1985, p. 79; Ouyang & Ma, 1992, p. 14). One female in each of two other colonies apparently became menopausal as early as age ca. 17 years (Tilford, 1981, p. 638; Vancatova et al., 1986, p. 263). The frequency of menopause in old females reported in three studies of captive colonies is as follows: 49 females, ages 25 to 27 years, 43% menopausal (Johnson & Kapsalis, 1998, p. 757); seven fe- males, ages 26 to 34 years, 71% menopausal (Walker, 1995, p. 61); 10 females, ages 28 to 34.6 years, 100% menopausal (Champ et al., 1996, p. 486). Based on hormone profiles of 26 captive females aged 21-29 years, 1 1 were determined to be premenopausal (mean age 22.5 years), 13 were determined to be perimenopausal (mean age 24.0 years), and 2 were determined to be postmeno- pausal (mean age 29.5 years)(Gilardi et al., 1997, p. 337). The greatest age at which a captive fe- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 11 Table 27. Inlam mortality rate in natural populations of Mcicaca mulatta. No. of Infant mortality rate' (%) No. of Study years Refer- Sample area groups period of data Mean ± SD Extremes ences- India Uttar Pradesh Aligarh District 8-21 1961-76' 15 18.0 ± 10.0 3.8-32.1 ; 8 1990-91 1 2.3 — 2 Chhatari-do-Raha 2 1961-76 16 15.5 ± 14.5 0-44.4 1 2 1990-91 1 2.2 — 2 Nepal Kathmandu Valley Pashupati ^-6 1975-78 3 29.0 ± 2.6 27-32 3 Swayambhu 7-10 1975-78 3 16.0 ± 7.0 11-24 3 China Guangxi Longhu Shan 1-2 1988-95^ 3 7.0 ± 3.8 3.7-11.1 4 Henan: Jiyuan Doudin 1 1987 1 31.5 — 5 Shagon 1 1987 1 28.5 — 5 ' Annual infant deaths during first year of life/annual births. - Key to references: I. Southwick et al., 1980, pp. 161-162. 2. Imam & Yahya. 1995. p. 6. 3. Teas et al., 1981. p. 1 19. 4. Wang et al.. 1996, p. 268. 5. Qu et al., 1993, p. 616. ' Data for 1975 missing. ' Data available for 1988, 1989, and 1995. male is known to have produced a living infant is ca. 28.5 years (Dyke et al., 1986, p. 264). Males are known to remain capable of copu- latory ejaculation to age ca. 30 years (Phoenix & Chambers, 1988, p. 159). However, in another se- ries of copulatory tests of six healthy males age ca. 21 to 31 years (mean = 25 years), four (67%) failed to ejaculate (Chambers & Phoenix, 1992, p. 15; cf. Ouyang & Ma, 1992, p. 14). The maximum reported longevity in captive M. mulatta is 37 years (Uno et al., 1998, p. 21; cf. Davis, 1985, p. 57; Erschler et al., 1988, p. 182). the annual mortality rate was 6.8% (7-year aver- age; Rawlins & Kessler, 1986c, p. 51). Observed causes of death in natural populations include infanticide, intraspecific fighting, preda- tion (dog, tiger), and an accidental fall from a cliff (Lindburg, 1971, p. 17; Camperio Ciani, 1984, p. 373; Singh, 1986, p. 607). In a provisioned Nep- alese population, ca. 40 monkeys were acciden- tally electrocuted in 1996 (Shrestha, 1997, p. 31; Chalise & Ghimire, 1998, p. 11). During winter, members of northern populations apparently die of exposure or malnutrition (Pearl et al., 1987, p. 37; Qu et al., 1993, p. 613). Annual Mortality Rate Annual mortality rates apparently are locally variable in natural populations of M. mulatta. In northern India, the annual mortality rate was ap- proximately 2.2% in Aligarh District (Imam & Yahya, 1995, p. 7), 3.6% at Asarori Forest (Lind- burg, 1971, p. 17), 3.7% at Tughlaqabad (3-year average; Malik et al., 1984, p. 317), and 9.6% at Dehra Dun (Lindburg, 1971, p. 17; cf. Southwick & M. F. Siddiqi, 1988, p. 190); in Nepal, the mor- tality rate was 19.0% at Swayambhu and 22.3% at Pashupati (3-year averages; Teas et al., 1981a, p. 1 19). In the free-ranging Cayo Santiago colony, Population Growth Rate Population growth rate estimates are available for 28 natural populations of M. mulatta (Table 28; cf. Ross, 1988. p. 218); for 14 of these pop- ulations, the census interval is 1 year, and for the remaining 14 the census interval is 3 to 32 years. In 12 of 14 populations with 1-year census inter- vals, annual growth rates vary from 3.8% to 26.9% (mean ± SD = 14.8% ± 6.8%); growth rates of the remaining two of these populations are outlier negative values (—9.5%, -15.9%). In the 14 populations with multiyear census inter- 78 FIELDIANA: ZOOLOGY vals, annualized growth rate tends to be greater in populations with shorter census intervals; in eight populations with census intervals less than 7 years, the mean annualized growth rate is 12.3% ± 5.9% (extremes. 0.9% and 19.1%). and in six populations with census intervals greater than 22 years, the mean annualized growth rate is 3.5% ± 4.5% (extremes, -0.4% and 9.1%). In three Nep- alese populations, annualized growth rate tends to be low regardless of census interval length (3 years, 0.9%; 23 years, -0.3% and -0.4%); the explanation for this low rate of population in- crease in Nepal is unclear (Teas et al., 1981a. p. 120; Johnson et al.. 1988. p. 179). In two areas on Hainan Dao. China, population growth rates apparently began to decline after optimum popu- lation densities were reached (Jiang et al., 1998. p. 101). In two free-ranging colonies introduced in Puerto Rico, the combination of relatively high population growth rates (13.5%, 16.5%) and rel- atively short census intervals (7 years. 9 years) is concordant with the norm for natural populations (see above). Fossils and Subfossils Published data concerning M. mulatto fossils and subfossils are meager and fragmentary (Table 29; cf. Szalay & Delson, 1979, pp. 356, 363; Del- son. 1980. p. 20; Pan & Jablonski, 1987, p. 63; Jablonski & Pan, 1988. p. 859; Jablonski. 1990. p. 39; Xue & Zhang, 1991, p. 357). The most important implication of these data probably is that a macaque similar or identical to M. mulotta had become established within the present range of M. mulatto in China and Vietnam during or prior to Late Pleistocene (>40 Ka). Systematics Geographic Variation and Subspecific Recognition Given the broad distribution and diverse habi- tats of M. mulatto (see above), it is not surprising that this species exhibits great variation in nu- merous characters. Based on individual or geo- graphic variation, 15 species-group names have been proposed for application to this taxon (Fig. 21). In the first comprehensive subspecific revi- sion of M. mulatto, six subspecies were recog- nized (Pocock, 1932, p. 533); in subsequent clas- sifications, various combinations of 10 subspecies have been recognized (Table 30). Although geographic variation in M. mulatto is clearly evident (see above), the differentiation of local and regional populations is now regarded as inadequate to warrant formal recognition of sub- species (cf. Fooden. 1995. p. 65). Where charac- ter-state transitions are gradual or irregular, as in M. mulatto, the delimitation of subspecies is ar- bitrary (cf. Mayr et al., 1953, p. 147), and un- ambiguous diagnosis of subspecies is virtually im- possible. Problems concerning the delimitation of subspecies in M. mulatto have been discussed by Pocock (1932, p. 530), Napier (1981, p. 20), Cor- bet (1992, p. 170), Jiang Xuelong et al. (1995, p. 46). and Yu et al. (1996, p. 153). Unless future research reveals a pattern of geographic differ- entiation of characters that is much more coherent than is now known in M. mulatto, it appears un- likely that taxonomically useful subspecies can be defined in this species. The principal characters that previously have been used in defining subspecies of M. mulatto are overall size, tail length, pelage color and length, and molecular diversity. The pattern of geographic variation in these characters has been discussed in detail above and is briefly summa- rized here. Size — Overall size usually is measured as the combined length of head and body (based on flesh measurements recorded by the collector). Overall size presumably may also be inferred from great- est length of skull, excluding incisors; although skull length is not a direct measure of overall size, compared with head and body length, skull length usually is available for more specimens, and it is subject to less interobserver variability. Meaning- ful comparisons of size require measurements of fully adult specimens, segregated according to sex. In M. mulatto, both head and body length and skull length tend to increase from south to north (Figs. 8, 17; Tables 3, 9). The large size of spec- imens collected north of 30°N in Afghanistan. Pakistan, and India in the west and disjunctly in China in the east is particularly striking. However, the latitudinal size gradient is gradual, and size distributions in local samples generally overlap (cf. Jiang Xuelong et al.. 1991. p. 242). Size in insular samples is generally similar to that in con- tinental samples collected at the same latitude (Figs. 8. 17). FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 79 Table 28. Population growth rate in Macacu mulatto. Annual/ annualized' population No. of groups Initial Final growth Refer- Sample area observed Census interval census census rate ( % ) ences- One- year census intervals Pakistan Dunga Gali 3 1978-79 105 95 -9.5 / India Jamniu and Kashmir Jammu 1 1983-84 123 156 26.9 2 Himachal Pradesh Simla 2 1983-84 154 186 20.8 2 Solan 2 1983-84 89 108 21.3 2 Haryana Bhiwani 18 1983-84 523 621 18.7 2 Hissar 29 1983-84 910 1.045 14.8 2 Jind 30 1983-84 1,016 854 -15.9 2 Rohtak 28 1983-84 857 932 8.8 2 Delhi 22 1983-84 744 772 3.8 2 Rajasthan Alwan 8 1983-84 1,067 1.219 14.2 2 Jaipur 5 1983-84 335 353 5.4 2 Uttar Pradesh AUgarh vicinity 14 1990-91 651 669 2.8 3 Dehra Dun 18 1983-84 597 688 15.2 2 Ghaziabad 17 1983-84 705 785 11.3 2 Saharanpur 12 1983-84 448 524 17.0 2 Multiyear census intervals India Delhi Tughlaqabad' 1-5 1965-90 (25 yr) 68 -600 -9.1 4 Rajasthan Bandipul 1 1976-80 (4 yr) 86 140 13.0 5 Jaipur 2 1976-80 (4 yr) 74 113 11.2 5 Marot 2 1976-80 (4 yr) 87 128 10.1 5 Uttar Pradesh Aligarh District 8-21 1959-91 (32 yr) -287 267 — 0.2 6 Chhatari-do-Raha^ 2 1959-91 (32 yr) -50 178 -4.0 6 Dehra Dun 2 197^80 (4 yr) 57 113 18.7 5 Khair 2 197^80 (4 yr) 96 171 15.5 5 Nepal Pashupati'-' 6-10 1975-98 (23 yr) 358 330 -0.4 7. 8 Swayambhu' 5-7 1975-98 (23 yr) 328 308 -0.3 7. 8 Tripureswor l-?2 1995-98 (3 yr) 37 38 0.9 8 China Hainan: Nanwan-* 5-20 1965-94 (29 yr) -115 -1,300 -8.7 9 Guangxi: Longhu Shan 3-6 1985-90 (5 yr) 128 203 9.7 10 Henan: Jiyuan^ -26 1982-88 (6 yr) -700 -2,000 -19.1 U U.S.A.: Puerto Rico Cayo Santiago' 5-6 1976-83 (7 yr) 479 1,161 13.5 12 La Parguera' 4-7 1963 '-72 (9 yr) 106 418 16.5 13 ' Annualized growth rate = [(C^/C,)"''] - 1. where Y = years in census interval, C, = initial census, and C, = final census. - Key to references: /. Melnick et al.. 1984, p. 344. 2. Seth et al.. 1992. p. 67. 3. Imam & Yahya. 1995. p. 4. 4. Malik et al.. 1984. p. 314; Malik. 1989a. p. 118; Southwick & Siddiqi. 1995. p. 19. 5. Seth & Seth. 1983. p. 63. 6. Southwick & Siddiqi. 1988. p. 189; 1994b. p. 57. 7. Teas et al.. 1980. p. 252; 1981a. p. 119; Johnson et al.. 1988, p. 179. 8. Chalise & Ghimire, 1998. p. 14. 15. 9. Jiang Haisheng et al., 1991. p. 213; 1998. p. 101. 10. Wang et al., 1996. p. 266. //. Anonymous, 1985, p. 241; Qu et al.. 1993. p. 608. 12. Rawlins et al., 1984, p. 249; cf. Koford, 1965, p. 160; Boelkins & Wilson, 1972. p. 136. 13. Drickamer. 1974b. p. 65. ' Provisioned. 80 FIELDIANA: ZOOLOGY Table 29. Localities and ages of Macaca nnilatta fossils or subfossils. Approx- imate Latitude Longitude age Refer- Locality Province/State (N) (E) Epoch (Ka') ences- China Yin ruins Henan 36°07' 114°19' Holocene 2.5 /. 2. 3 Xiawanggang Henan ~33°15' ~lir27' Holocene 4.5-5.2 2, 4. 5 Zhengpiyan Guangxi ~25°17' ~110°17' Holocene 6.6 2. 5. 6 Hemuda Zhejiang ~30°03' ~121°09' Holocene 6-7 2. 7 Baoshan' Yunnan 25°05' 99°05' Holocene 7 8 Shuanglong Cave^ Zhejiang 29°12' 119°37' Holocene 7.x 9 Xianren Cave Jiangxi 28°45' 117°09' Holocene 8.5 2. 3. 10 Shenxian Cave^ Jiangsu ~31°38' ~119°02' Holocene/PIeistocene 11.2 2, 5, // Luoding^ Guangdong 22°40' 111°30' Pleistocene'* 10-120 12 Jiandaoqian Shan Fujian -24°30' ~117°30' Pleistocene^ 40-120 13 Vietnam Keo Leng^ Lang Son ~2r57' ~106°23' Pleistocene 20-30 14. 15 Lang Trang' Than Hoa 20°2r 105°13' Pleistocene 7 15. 16 India Goalpara (= Giilpara) Assam 26°10' 90°37' Holocene 7 17 Madras region ?Andhra Pradesh •7 ■7 Holocene ■) 17 ' Ka = thousands of years ago. -Key to references: 1. Teilhard de Chardin & Young, 1936, p. 53. 2. Zhang, 1985, pp. 164, 165. 3. Chang, 1986, pp. 100. 317. 4. Chia & Chang, 1977, p. 49. 5. Xue & Zhang, 1991, pp. 330, 331. 6. Li & Han, 1978, p. 249. 7. Chekiang Provincial Museum, 1978, p. 95; Wu, 1983. p. 165; Han, 1988, p. 869. 8. Jablonski et al., 1994, p. 307. 9. Ma & Tang, 1992, p. 310. 10. Huang & Chi, 1963, p. 266. 11. Li & Lei. 1980, p. 60. 12. Gu et al., 1996, p. 247; Jablonski & Gu, 1996, p. 130. 13. You & Cai. 1996. p. 341. 14. Nguyen. 1985, p. 97; Olscn & Ciochon, 1990, pp. 764, 766. 75. Nisbett & Ciochon. 1993. p. 774. 16. Ciochon et al.. 1990, pp. 1 12, 240. 17. Lydekker. 1880. pp. 32. 33. ' Species identification provisional. "* Xiashan Cave and Shanbeiyan Cave. ^ '"[Plrobable Late Pleistocene." '' "[E]arly Late Pleistocene." Tail Length — Like head and body length, tail length in adults of both sexes of M. mulatta gen- erally tends to increase from south to north (Fig. 9; Table 4). However, relatively long-tailed spec- imens collected in the Indochinese peninsula, near the southeastern limit of distribution of M. mulat- ta, constitute an important exception to this gen- eralization; these aberrant specimens have been interpreted as the result of hybridization between M. mulatta and M. fasciculah.s (Fooden, 1997, p. 228). Because of the generally parallel latitudinal variation of tail length and head and body length in M. mulatta, relative tail length (tail length/head and body length) is relatively constant latitudi- nally, except in the extreme southeast, where it increases at lower latitudes (Fig. 11; Table 5). Longitudinally, tail length and relative tail length in M. mulatta tend to decrease east of ca. 105°E, again except in the extreme southeast (Fig. 12; Table 5). Both latitudinally and longitudinally, variation in tail length and relative tail length is gradual (Figs. 11, 12). In insular samples, tail length and relative tail length are similar to cor- responding lengths in nearby continental samples (Figs. 9, 11). Pfxagf- — Analysis of geographic variation in pelage color and pelage length in M. mulatta is complicated by age variation and particularly by seasonal variation (see above). However, when comparisons are restricted to postinfantile speci- ■* In 1996. ca. 40 monkeys were accidentally electrocuted in this group. "^ Includes two provisioned groups. *" In 1975-83. 189 monkeys were removed from this population for research purposes. ' Infants born in 1963 are included in a census of this population that was conducted in early 1964. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 4o»-: ^(/2 mcmahoni- Type localities of proposed species and subspecies .villosus 3ori rhesus 20" m 70° littoralis erythraea sanctijohannis brevicaudus brachyurus 110° A°' Fig. 21. Type localities of nominal species or subspecies allocated to Macaco mulatto: known limits of natural distribution of Macaco mulatto also indicated. Type locality references: Cercopithecus (Mulatto) Zimmermann, 1780, p. 120 — "India": restricted to "Nepal Tarai" by Pocock (1932. p. 533). Cercopithecus ful\ us Kerr, 1792, pp. 32e. 73 — objective synonym of Cercopithecus {Mulatto) Zimmermann. Simla rhesus Audebert, [1799], p. 5 — provenance unknown: designated as "ITnde . . . des bords du Gange" by F. Cuvier (1819, p. 2). Simla Erythraea Schreber, [18()()|, suppl. pi. 8c — objective synonym of Simla rhesus Audebert. Mocacus Olnops Hodgson, 1841, p. 1212 — "Tarai and lower Hills", Nepal: restricted to "Nepal Tarai" by Pocock ( 1932, p. 533). Inuus sonctl-johannls Swinhoe, [1867], p. 556— "North Lena Island" (= Dangan Dao), Guangdong Province, China. Mocacus loslotus Gray, 1868, p. 60 — "S/echwan" (= Sichuan Province), China. Mocacus Tchellensls Milne-Edwards, [1870], pis. 32, 33 — "la cordilierc de Test de la province du Tche-ly [= Hebei]", China. Mocacus vestltus Milne-Edwards. 1892, p. 671 — "Du Tengri-Nor a Batang," China: specified as "Tasin Lou" (= Kangding), Sichuan Province, China, on stand of mounted holotype. Mocacus rhesus vUlosus True, 1894, p. 2 — "Lolab. Kashmir, . . . 7,500 feet" {= Lolab. Jammu & Kashmir, 2300 m), India. Plthecus littoralis Elliot, 1909, p. 250 — "Kuatun," Fujian Province, China. Plthecus brachyurus Elliot. 1909. p. 251 — "Island of Hainan." China: specified as "Mt. Wuchi" (= Wuzhi Shan). Hainan Dao. China, on specimen tag of holotype: name is a permanently invalid junior primary homonym. Plthecus brevi- caudus Elliot. 1913, p. 216 — replacement name for Plthecus brachyurus Elliot. Macaco slamlca Kloss, 1917, p. 247 — "Me Ping rapids below Chiengmai, . . . 850 ft" (= Kaeng Mae Hat, Mae Nam Ping, 260 m). Thailand. Macaco mulatto mcmahoni Pocock, 1932, p. 544 — "Kootai in Lower Chitral. between the Bashgal Valley in Kafiristan and the Chitral Valley: 3,600 ft." (= Kaotai, lower Kunar River, 1 100 m), Pakistan. men.s in prime pelage, individual variation in pel- age color apparently exceeds geographic variation in this species. Pelage color frequently differs strongly in individuals collected at the same lo- cality (Fig. 3B), and pelage color frequently is similar or identical in individuals collected at lo- calities separated by thousands of kilometers (see discussion above, "Pelage — Geographic Varia- tion— Summary," p. 25; cf. Jiang Xuelong et al.. 1991. p. 242). Although pelage length, both on body and appendages, tends to increase from south to north, the latitudinal transition is gradual (Figs. 6, 7). Molecular Diversity — Although mtDNA haplotypes in M. mulatta generally are unique at each locality sampled, haplotypes in neighboring 82 FIELDIANA: ZOOLOGY Table 30. Subspecies recognized (x) in published classilicalions of Mcicaca miilatta. 1932-95.' Italicized code numbers indicate recognized subspecies to which unrecognized subspecies are referred in cited classifications: dash ( — ) indicates that proposed subspecies is not considered in cited classification. Prof >osed .subspecies Authors- and dates of classifications Code Poc. Kel. EMS Hill QWZ MW Peng JWM WJ no. Name' Date 1932 1945 1951 1974 1981 1988 1990 1991 1995 1 initlatta 1780 X X X X X X X X X 2 sanclijohanius 1867 X X V 1' r 8' — 3 lasiotHS 1868 X X r 5' r X^ X" 4 tclu'liensis 1870 X X r 5' X x^ X 5 vestitus 1892 — 1 X X X X X 6 villosus 1894 X X — — — — — 7 littoralis 1909 2 2 r 1' }' X X 8 biTvicaudus 1913 / /" X x" x" X X 9 siainica 1917 / / — — — J' X 10 incinahoni 1932 X X X X — — — — — ' Allen ( 1930. p. 1 : 1938, p. 284), who did not publish a formal classification of subspecies of M. mulatto, expressed doubt concerning the distinctness of all propo.sed subspecies, with the possible exception of M. m. vestita and M. m. villosa: Corbet's ( 1992, p. 170) opinion is similar ("it is unlikely that discrete subspecies can be recognized""). - Key to abbreviations: Poc. = Pocock, p. 533: Kel. = Kellogg, p. 212 (similar classification subsequently published by Napier. 1981, p. 21); EM-S = Ellerman & Morrison-Scott, p. 197 (similar classifications subsequently published by Sanderson. 1957. p. 127: Napier & Napier. 1967. p. 404: Fa. 1989. p. 54: and Zhang et al.. 1991. p. 177): Hill, p. 565: QWZ = Quan. Wang. & Zhang, p. 8 (cla.ssification repeated in Zhang et al.. 1997. p. 58): MW = Ma & Wang. p. 252: Peng. 1990. p. 21: JWM = Jiang. Wang. & Ma, p. 242: WJ = Wang & Jiang, 1995, pp. 4. 9 (cf. Peng et al., 1993, p. 4: Zhang & Shi, 1993b. p. 600: Yao et al., 1995, p. 1 16). ' Termination as in original spelling of name. "* Allocation of unrecognized subspecies inferred from geographic distributions given for recognized subspecies. ^ Unrecognized subspecies allocated provisionally. '' Cited by synonym or incorrectly spelled name. ■ Recognized provisionally. populations tend to be more similar than those in widely separated populations (see "Mitochondrial DNA" above); somewhat unexpectedly, known haplotypes in eastern M. mulatto are more similar to those in M. cyclopis and M. fiiscata than they are to those in western M. mulatto. Further eval- uation of the bearing of mtDNA data on subspe- cies determination in M. mulatto will require ad- ditional samples from precisely known, geograph- ically intermediate localities. Few data are avail- able concerning geographic variation of nuclear DNA in M. mulatto (see above). Available blood- protein allele frequency data (see above) also are inadequate for delimitation of subspecies in M. mulatto. Synonymy Macaca mulatta (Zimmermann, 1780, p. 195) Tawny [Monkey]: Pennant, 1771, p. 120 — based on menagerie animal, not preserved: "Inhabits India."" Cercopithecus [sp.]: Erxleben, 1777, p. 43 — generic allocation of Tawny Monkey: Pennant, 1771. Cercopithecus (Mulatto) Zimmermann, 1780, p. 195 — based solely on Tawny Monkey: Pennant, 1771: origin "O.stindien."" Cercopithecus molatta: Anderson. 1879. p. 56 — in- correct spelling, not an available name. Simio Mulatto: Goldfuss. 1809. p. 61 — new combi- nation. Macaco mulotto: Hinton & Wroughton. 1921. p. 668 — new combination: laxonomic history. Macaco mulato: Tate. 1947. p. 132 — incorrect spell- ing, not an available name. Macaca imillatto: Khajuria. 11955], p. 1 14 — incorrect spelling, not an available name. M[ocacus\ mulatto: Dover. 1932. p. 244 — new com- bination. Rhesus mulatto: Furuya. 1962. p. 377-new combina- tion. Macaca mulatto mulatto: Pocock. 1932. p. 533 — new rank: type locality restriction. Macaco mulloto mullato: Khajuria, [1955|. p. 113 — incorrect spelling, not an available name. Macaque a queue courte: Buffon. 1789. p. 56. pi. 13 — based on captive feinale. died 7 February 1778. skin formerly preserved "au cabinet du Roi"; provenance unspecified. Cuvier. 1819. p. 2 — distribution. "Flndc . . . des bords du Gange." Hinton & Wroughton. 1921. p. 668 — synonym of Cercopithecus mulatto Zimmermann, 1 780. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 83 Cercopitheciis fiilvus Kerr. 1792. pp. 32e, 73 — based solely on Tawny Monkey: Pennant, 1771; habitat, India. Hinlon & Wroughton. 1921, p. 668 — syno- nym of Cenopithecus muUitta Zinimermann. 1780. Pocock. 1932. p. 533 — synonym of Macaca mu- Uitta muUitta (Zimmcrmann. 1780). Simici fiilva: Shaw. 18()(), p. 57 — new combination. Anderson. 1879. p. 56 — possible synonym of Cer- copitheciis m\ii\latta Zinimermann. 1780. Simla rhesus Audebert, [1799]. p. 5 — based on (1) captive female, not preserved, provenance un- known, and (2) Macaque a queue courte: Buffon. 1789. Cuvier. 1819. p. 2 — distribution, -rinde . . . des bords du Gauge." Fischer. 1829. p. 29 — hardly separable from Cenopithecus mulatta Zinimer- mann. 1780. Anderson. 1879. pp. 55. 56 — possible synonym of Cenopithecus m[u]latta Zimmermann. 1780. Hinton & Wroughton, 1921. p. 668— syno- nym of Cercopithecus mulatta Zimmermann. 1780. Pocock. 1932. p. 533 — synonym of Macaca mu- latta mulatta (Zimmermann. 1780). Macaca rhesus: Daudin. 1802. p. 148 — new combi- nation. Pithecus rhesus: E. Geoffroy. 1803, p. 25 — new com- bination. Cyn[ocephalus] rhesus: Latreille, 1804, p. 292 — new combination. Inuus rhesus: E. Geoffroy, 1812, p. 101 — new com- bination. Innuus Rhesus: Schinz, 1821, p. 1 13 — incorrect spell- ing of generic name. [M]acacus rhesus: Desmarest, 1820, p. 66 — new combination. Papio Rhesus: Ogilby, [1840], p. lix — new combina- tion. Papio Rhaesus: Percy. 1844. p. 83 — incorrect spell- ing, not an available name. [Silenus] rhesus: Stiles & Nolan. 1929. p. 533-new combination. Simla Eiythraea Schreber. [1800]. suppl. pi. 8c — name proposed in caption of figure copied from illustration of Macaque a queue courte: Buffon. 1789. pi. 13: text published subsequently by Wag- ner. [1839]. p. 142 (cf. Sherborn. 1892. pp. 590, 591). Goldfuss. 1809, p. 54 — synonym of Simla rhesus Audebert, 1799. Cuvier, 1819, p. 2 — distri- bution, "Tlnde . . . des bords du Gange." Ander- son. 1879. p. 55 — possible synonym of Cercopi- thecus ni[u]latta Zimmermann. 1780. Blanford. [1888a|. p. 625 — taxonomic history. Hinton & Wroughton. 1921. p. 668 — synonym of Cercopi- thecus mulatta Zimmermann, 1780. Pocock. 1932. p. 533 — synonym of Macaca mulatta mulatta (Zimmermann, 1780). Macacus erythraeus: 1. Geoffroy, 1826, p. 588 — new combination. I\nuus\ erythraeus: Wagner, [1839], p. 142 — new combination. Innuus {Maimon) erythraeus: Anderson, 1879, p. 56 — incorrect spelling of generic name. Pith[ecus] {Mac[acus\) erythraeus: Dahlbom, 1856, p. 1 16 — new combination. [Silenus] erythraeus: Stiles & Nolan, 1929, p. 529— new combination. Wrinkled Baboon: Shaw, 1800. p. 33— based on Ma- caque a queue courte: Buffon. 1789. and Simla er- ythraea Schreber. [1800]. Macacus radiatus: Hodgson, 1834, p. 96 (not E. Geof- froy. 1812) — misidentification. Ogilby. [1840], p. Ix — .synonym of \Simia\ rhesus Audebert. [1799]. Macacus [(Pithex)] Oiuops Hodg.son. 1841. p. 1212, fig. p. 1213 — type series not specified, probably in- cluded among seven specimens (including three skulls only) collected in Nepal by B. H. Hodgson (date unknown): bm(nh) 1845.1.8.5 (skull only), 1845.1.8.222-224, 1858.6.24.144 (skull only), 1972.1013 (skull only, not seen). 1972.1015 (cf. Napier. 1981, p. 24); type locality, Nepal: "Tarai [= plain] and lower hills." Blyth, 1844, p. 475— probable synonym of [Simla] rhesus Audebert, [1799]. Gray, 1846, p. 2 — synonym of [Simla] rhe- sus Audebert, [1799]. Wagner, [1851-55], p. 56— synonym of Inuus erythraeus (Schreber, [1800]). Wroughton, 1918. p. 555 — lectotype designated. [BM(NH)] No. [18] 43.1.12.5 [No specimen num- bered 1843.1.12.5 is now present in the bm(nh) col- lection; however, a note on the label of bm(nh) 1972.1015 indicates that this renumbered specimen may be the missing lectotype of Macacus oinops.] Hinton & Wroughton. 1921. p. 668 — synonym of Cercopithecus mulatta Zimmermann, 1780. Po- cock, 1932, p. 533 — synonym of Macaca mulatta mulatta (Zimmermann, 1780). Napier, 1981, p. 24 — probable type series cataloged. Macaca cinops: Chiarelli. 1972, p. 208 — incorrect spelling, not an available name. [Macacus] Nipalensis Hodgson, 1841, p. 1212 — man- uscript name cited as a synonym of Macacus [{Pithex)] Oinops Hodgson, 1841, not an available name. Gray. 1846, p. 2 — synonym of [Simla] rhe- sus Audebert, [1799]. Pocock, 1939, p. 45 — syno- nym of Macaca mulatta mulatta (Zimmermann, 1780). M[acacus] Pelops: Gray. 1843, p. 8 (not Hodgson, 1841) — misidentification. Hinton & Wroughton, 1921, p. 668 — synonym of Cercopithecus mulatta Zimmermann, 1780. Inuus pelops: Hutton, 1865, p. xiii (not Hodgson, 1 84 1 ) — misidentification. Inuus sanctl-johannls Swinhoe, [1867], p. 556 — ho- lotype, BM(NH) 1868.12.29.10. juvenile female, skin and skull, collected alive at North Lena Island ( = Dangan Dao), Guangdong, China, by Commander St. John, ca. June 1866, died in captivity ca. De- cember 1868. Swinhoe, [1871], p. 615— type his- tory. Sclater, 1871, p. 222 — possible synonym of [Simla] rhesus Audebert, [1799]. Allen, 1930, p. 2 — synonym of Cercopithecus mulatta Zimmer- mann, 1780. Kellogg, 1945. p. 121 — synonym of Macaca mulatta mulatta (Zimmermann. 1780). Na- pier. 1981. p. 22 — holotype cataloged. Jiang Xue- long et al.. 1991. p. 243 — provisional synonym of M[acaca] mulatta hrevlcaudus (Elliot. 1913). Innus scmctl-johannis: Matschie. 1912. p. 306 — in- correct spelling oi generic name. Innuus sanctl-johannls: Pocock. 1932. p. 546 — incor- rect spelling of generic name. Macacus sanctl-johannls: Swinhoe. 1870. p. 615 — new combination. Pithecus sanctl-johannls: Elliot. 1913. p. 198 — new combination. 84 FIELDIANA: ZOOLOGY [Silenus] sancti-johannis: Stiles & Nolan. 1929. p. 535 — new combination. Macaca mulalta sancti-joliannis: Pocock, 1932. p. 546 — new rank. Macacus lasiotus Gray. 1868, p. 60 — holotype. BM(NH) 1871.4.21.4. adult male, skin, skull, and skeleton (No. 1561a): bobtailed captive shipped from Shanghai by relative of C. A. Winkworth. who presented captive to Zoological Society of London, 15 January 1868: captive died 25 May 1870: reported origin, "Szechwan" (= Sichuan Province), China. Sclater. 1871. p. 221 — type his- tory: possible synonym of [5//;/;^] rhesus Audebert. [1799]. Allen, 1930, p. 2 — synonym of Cercopi- thecus iniilatta ZimmerimLnn. 1780. Kellogg. 1945. p. 121 — synonym of Macaco inulatta mulatta (Zimmermann. 1780). Napier. 1981, p. 22 — holo- type cataloged. Macacus lasiods: Gray. 1868, figure caption — inad- vertent misspelling (cf. text), not an available name. Macacus lariotis: Mollendorf. 1889. p. 9 — incorrect spelling, not an available name. I[nuus] lasiotus: Blyth, 1875, p. 5 — new combination. Pithecus kisiotis: Elliot, 1913, p. 198 — new combi- nation. [Silenus] lasiotis: Stiles & Nolan. 1929. p. 531 — new combination. Macaca mulatta lasiotus: Pocock. 1932, p. 548 — new rank. Macacus Tcheliensis Milne-Edwards, [1870], captions for plates 32 and 33 — holotype. mnhn 335/281 A/ 1867-557 (Type Cat. No. 61). juvenile female, skin and skull (lacking mandible), collected in moun- tains of eastern "Tche-ly"' (= Hebei) Province by M. Fontanier, 1867. Milne-Edwards. [1872], pp. 227. 229 — external and cranial characters: taxono- my, possibly a synonym of M[acacus] lasiotus Gray, 1868. Anderson, 1879. p. 83 — synonym of Macacus lasiotus Gray. 1868. Osgood, 1932. p. 209 — provisional synonym of Cercopithecus mu- latta Zimmermann. 1780. Ho. 1935. p. 139 — syn- onym of Cercopithecus mulatta Zimmermann. 1780. Rode, 1938, p. 224— holotype cataloged. Kellogg, 1945, p. 121 — synonym of Macaca mu- latta mulatta (Zimmermann. 1780). M[acacus] tchiliensis: Blyth. 1875. p. 6 — incorrect spelling, not an available name. Anderson in Blyth, 1875, p. 6 — probable synonym of M\acacus\ lasi- otus Gray. 1868. Macaca tcheliensis: Flower. 1931. p. 154 — new com- bination. [Macacus lasiotis] tcheliensis: Trouessart. 1897. p. 27 — new rank. Macaca lasiotis tscheliensis: de Beaux, 1923. p. 28 — incorrect spelling, not an available name. [Silenus] lasiotis tcheliensis: Stiles & Nolan, 1929, p. 531 — new combination. Macaca mulatta tcheliensis: Pocock. 1932, p. 550 — new combination. Jiang Xuelong et al.. 1991, p. 245 — validity of subspecies questionable. M[acaca] m[ulatta] tcheliensis: Jiang Xuelong et al.. 1991, p. 241 — incorrect spelling, not an available name. Macacus vestitus Milne-Edwards. 1892, p. 671 — ho- lotype, MNHN 334/282F/ 189 1-388 (Type Cat. No. 59: Coll. No. 52), adult male, skin and skull, col- lected at "Tasin-Lou"" (= Kangding). Sichuan Province, China, by H. d"Orleans, June-July 1890: paratype, young female, purchased alive at "Houmda"7'"Kian Tatie" (= Ngamda), Xizang Province, China, by H. dOrleans. 7 May 1890. liv- ing in menagerie of mnhn 22 August 1892. prob- ably MNHN C.G.I 892-3 1 5 or C.G.I 894- 1432 (both skins only. "Tibet"). Osgood, 1932, p. 209 — pro- visional synonym of Cercopithecus mulatta Zim- mermann, 1780. Rode. 1938, p. 223 — holotype cat- aloged. Kellogg, 1945, p. 121 — synonym of Ma- caca mulatta mulatto (Zimmermann, 1780). Pithecus vestitus: Elliot, 1913 — new combination. Macaco vestitus: Rode. 1938, p. 223 — new combi- nation. Macaco mulatto vestito: EUerman & Morrison-Scott, 1951. p. 198— new rank. Macacus rhesus villosus True. 1894, p. 2 — holotype. USNM 20120/35485 (Coll. No. IV). adult male, skin and skull, collected at Lolab. Jammu & Kashmir, India, by W. L. Abbott, 8 September 1891: para- types. USNM 20121/35486 (Coll. No. 5. juvenile male. 9 September), lsnm 20122/38172 (Coll. No. 6. juvenile male. 9 September). L'.snm 20123/35488 (Coll. No. 7, subadult male. 8 September), usnm 20124/35489 (Coll. No. 8. subadult male. 9 Sep- tember), skins and skulls, collected at Lolab. Jam- mu & Kashmir, India, by W. L. Abbott. 1891. Blan- ford. 1891. p. 361 — synonym oi Macaco ossamen- sis McClelland in Horsfield. [1840]. Lyon & Os- good, 1909, p. 285 — holotype cataloged. Allen, 1930, p. 1 — "doubtfully distinct race." Poole & Schantz, 1942, p. 245 — holotype catalogued. M[acaco] rhesus villosus: Wroughton, 1918, p. 544 — new combination. Macaco mulatto villoso: Pocock. 1932. p. 539 — new combination. [Macacus] villosus: Trouessart. 1897. p. 27 — new rank. Pithecus villosus: Elliot, 1913, p. 200 — new combi- nation. M[ocacus] ossamensis: Blanford, 1898. p. 361 (not McClelland in Horsfield. [ 1840]— misidentifica- tion. Pithecus ossamensis: Wroughton. 1915a. p. 464 (not McClelland in Horsfield. [1840]— misidentifica- tion. Hinton & Wroughton. 1921. p. 668 — syno- nym of Cercopithecus mulatta Zimmermann, 1780. Macaca ossamensis: Wroughton, 1916c, p. 763 (not McClelland in Horsfield. [1840]— misidentifica- tion. Pithecus littorolis Elliot. 1909. p. 250 — holotype, BM(NH) 1900.5.8.1 (Coll. No. a), adult female, skin and skull, collected at Kuatun. Fujian Province, China, by C. B. Rickett. November 1898: para- types, BM(NH) 1898.11.1.29 (juvenile male, skin and skull, purchased at Kuatun. Fujian Province, China, by J. de la Touchc. 12 May 1898), bm(nh) 1871.3.3.5 (juvenile female, menagerie animal, skin and skull, obtained from Zoological Society of London before 1872. provenance reportedly "Kashmir"). Allen. 1930. p. 2 — synonym of Cer- copithecus mulatto Zimmermann, 1780. Pocock, 1932, p. 546 — synonym of Macaca mulatto sancti- johannis (Swinhoe. [1867]). Kellogg, 1945, p. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 85 121 — synonym of Maccica muUitta muUitta (Zim- mcrmann. 1780). Napier. 1981, pp. 22. 25— type series cataloged. M[acaca\ inulcitici lituinilis: Jiang Xuelong el al.. 1991, p. 244— new rank. Pithecus brachyurus Elliot, 1909, p. 251 (not Maca- cus brachyurus Smith, 1842, p. 103) — holotype, AMNH 21511. adult male, skin and skull, collected at Mt. Wuchi (= Wu/.hi Shan), Hainan Dao, China, by A. Owston. 10 October 1905: paralypes (inex- plicitly cited in original description), amnh 26646 (Coll. No. 13/16, adult male, skin and skull, col- lected at Henron, Hainan Dao, China by A. Ows- ton, 10 May 1904), amnh 27568 (infant male, 1 October), 27569 (juvenile female, 2 October), 27570 (juvenile female, 3 October), 27571 (adult female, 4 October), 27572 (subadult male, 5 Oc- tober), 27573 (adult female, 5 October), 27574 (adult female. 6 October). 27575 (juvenile female. 7 October), 27578 (subadult female, 30 October), BM(NH) 1909.7.11.1 (Coll. No. 13, AMNH 27576. ju- venile male, 10 October), collected at Wuzhi Shan, Hainan Dao, China, by A. Owston. 1905. Elliot, 1913. p. 217 — homonym of Macacus brachyurus Smith, 1842; replaced by Pithecus brevicaudus El- liot, 1913. Macaca mulatto brachyurus: Xu et al., 1983, p. 312 — new rank. Pithecus brevicauilus Elliot. 1913, p. 216 — replace- ment name for Pithecus brachyurus Elliot. 1909. junior homonym of Macacus brachyurus Smith, 1842. Pocock, 1932, p. 533 — synonym of Macaca mulatta mulatta (Zimmermann, 1780). Pithecus brevicaudatus: Fiedler, 1956, p. 179 — incor- rect spelling, not an available name. Macacus brevicaudus: Meli. 1922, p. 11 — new com- bination. M{acaca] brevicaudatus: Tate, 1947, p. 134 — new combination; incorrect spelling, not an available name; "doubtfully valid form." M[acaca] m[ulatta\ brevicaudus: Quan et al., 1981, p. 8 — new rank. M[acaca] m[ulatta] brovicaudata: Peng, 1990, p. 21 — incorrect spelling, not an available name. Macaca siamica Kloss, 1917, p. 247 — holotype, ZRC 4-188/2530 (Coll. No. 320), adult male, skin and skull, collected at Me Ping rapids, 850 ft (= Kaeng Mae Hat. Mae Nam Ping. 260 m). below Chiang Mai. Thailand, by K. G.^Gairdner, 14 April 1916. Pocock, 1932, p. 533 — synonym of Macaca mu- latta mulatta (Zimmermann, 1780). Weitzel et al., 1988, p. 1 16 — holotype cataloged. Macaca simica: Yang & Chou, 1984, p. 56 — incorrect spelling, not an available name. M[acaca] m\ulatta\ siamica: Tate, 1947, p. 134 — new rank. Macaca mulatta mcmahoni Pocock, 1932, p. 544 — holotype, bm(nh) 1920.6.1 1.1, adult male, skin and skull, collected al Kootai, lower Chitral, 3600 ft ( = Kaotai, lower Kunar River, 1 100 m), Pakistan, by F. D. Stirling, early February 1914; paratype, BM(NH) 1931.1.9.1, [adult male], skin only, ob- tained alive in eastern Nurestan. Pakistan, by H. McMahon, received at Regents Park Zoo 3 April 1906, died 19 January 191(\ Napier, 1981, p. 24— type series cataloged. \Macaca mulatta] momahoni: Buettner-Janusch, 1963, p. 52 — incorrect spelling, not an available name. \Macacci mulatta] momahori: Peng et al., 1993. p. 4 — incorrect spelling, not an available name. M[acaca\ m\ulatta\ momachori: Peng et al.. 1993, p. 5 — incorrect spelling, not an available name. Type Cercopithecus mulatta Zimmermann, 1780 (p. 195), is expressly based on Pennant's (1771, p. 120) brief characterization of a menagerie captive that he observed, presumably in London, "in Mr. Brooks's exhibition." No part of the captive, which Pennant designated as the Tawny Monkey, is known to have been preserved. In an addendum. Pennant (1771, p. xxiii, pi. XIII.A, fig. II) characterized and figured a second monkey (unpreserved, species unidentified) that he regarded as a "variety" of the Tawny Monkey. This second monkey also is cited as a "Spielart" in Zimmermann's (1780, p. 195) original descrip- tion of Cercopithecus mulatta. As an acknowl- edged variant, the second monkey does not qual- ify as a syntype of Cercopithecus mulatta (Inter- national Code of Zoological Nomenclature, 1985, Article 72[b]:i). Although the holotype of Cer- copithecus mulatta Zimmermann appears not to have been preserved, designation of a neotype is not appropriate here. Article 75(a) of the Inter- national Code of Zoological Nomenclature (1985) specifies that a neotype is to be designated "only in exceptional circumstances when a neotype is necessary in the interests of stability of nomen- clature." Because such circumstances do not ap- ply to M. mulatta, designation of a neotype for this species would have no standing (Article 75[c]). Type Locality Concerning the geographic origin of the Tawny Monkey, Pennant (1771, p. 120) merely com- mented, "Inhabits India"" (italics in original). Zimmermann (1780, p. 195) rephrased this com- ment as "Er kam aus Ostindien." Based on the type locality of Macacus oinops Hodgson, 1841, a subjective synonym of Cercopithecus mulatta Zimmermann, 1780. Pocock (1932, p. 533) re- stricted the type locality of M. mulatta to "Nepal Tarai" — that is, the belt of Nepalese lowlands 86 FIELDIANA: ZOOLOGY (Terai) that extends along the border between Ne- pal and India (Fig. 21). Evolution and Dispersal Fossil evidence indicates that a macaque simi- lar or identical to M. mulatta inhabited Vietnam ca. 20 to 30 Ka and inhabited eastern China ca. 40 to 120 Ka (Table 29). This evidence estabishes minimum dates for the existence of M. mulatta, or a close relative, in eastern Asia; the actual date of first appearance of this species in this region may of course be much earlier. Following is a hypothetical interpretation of the evolutionary and geographical history of this species, based pri- marily on variation in relative tail length (Fig. 22). M. mulatta is a member of the fascicularis group of macaques, which also includes M. fas- cicularis in peninsular and insular Southeast Asia, M. cyclopis in Taiwan, and M. fuscata in Japan (cf. Fooden & Albrecht, 1999, p. 432; Morales & Melnick, 1998, p. 17). In these species, relative tail length generally decreases as latitude increas- es, in accord with Allen's rule (Table 31). Assuming that reduced relative tail length is a shared derived character state in the fascicularis group, M. fascicularis may be regarded as the primitive sister group of M. cyclopis, M. mulatta, and M. fuscata. Initial reduction of relative tail length and splitting of the derived species from M. fascicularis presumably occurred before ca. 40 Ka, in or near the northern part of the Indochinese peninsula (Fig. 22A); reduction of relative tail length in this area probably was an evolutionary response to lower ambient temperatures encoun- tered by a fascicularis group population as it dis- persed northward from its ancestral tropical hab- itat. The latitudinal range of the northward-dis- persing fascicularis group population eventually extended to at least ca. 40°N in eastern Asia. Although relative tail length in the fascicularis group generally decreases with increasing lati- tude, this correlation does not apply to southern populations of M. mulatta (ca. 15-25°N), in which relative tail length is similar to that in conspecific northern populations (Fig. 11) and therefore is less than expected according to Allen's rule. This suggests that southern populations of M. mulatta did not originate within their present latitudinal zone but instead dispersed there relatively recent- ly from farther northward (cf. Fooden & Albrecht, 1999. p. 438). Judging from the general relation- ship between latitude and relative tail length in the fascicularis group, relative tail length in the population of this group that originally inhabited the 15 to 25°N latitudinal zone probably was sim- ilar to that in M. cyclopis. This interpretation as- sumes that evolutionary shortening of a long tail in response to cooler climate occurs more readily than evolutionary lengthening of a short tail in response to warmer climate (cf. M. fuscata. Figs. 22 A. B; M. a. assamensis, Fooden, 1988, pp. 4, 9). Mean relative tail length in the northward-dis- persing fascicularis group population apparently varied from ca. 0.90 at ca. 20°N to ca. 0.30 at ca. 40°N; conversely, body size and pelage length and density presumably increased at higher latitudes (Figs. 6, 8). At this evolutionary stage, the pro- genitors of M. cyclopis (mean relative tail length ca. 0.90) presumably dispersed from the mainland to Taiwan, and the progenitors, of M. fuscata (mean relative tail length ca. 0.30) dispersed from the mainland to the Japanese islands. There is no evidence that the so\x\hevx\ fascicularis group pop- ulation (15-25°N) with mean relative tail length ca. 0.90 dispersed northwest of the Indochinese peninsula; this suggests that a barrier to north- westward dispersal may have exi.sted during this stage in the evolution of the fascicularis group. As indicated above, the evidence of relative tail length suggests that an M. mulatta population (mean relative tail length <0.50) dispersed south- ward and replaced the cyclopis-\\kQ population (mean relative tail length ca. 0.90) that is postu- lated to have originally inhabited the 15°-25°N latitudinal zone in mainland eastern Asia (Fig. 22B); as a result of this replacement, M. cyclopis became relictual in Taiwan. The southward dis- persal of M. mulatta and the correlated replace- ment of the aboriginal mainland cyclopis-\\ke population may have occurred during the last gla- cial maximum (ca. 18 Ka), when climatic deteri- oration rendered the northern part of the present range of M. mulatta unsuitable for habitation by this species (Xu, 1988. p. 875; Tong & Shao, 1991, p. 65; Tong & Zhang, 1991, p. 389; Winkler & Wang, 1993, p. 245; Zheng & Lei, 1999, p. 357). Before or during the southward shift of the range of M. mulatta in eastern Asia, a west-east gradient of declining relative tail length apparent- ly had become established in this species (100°E, mean relative tail length ca. 0.45; 120°E, mean relative tail length ca. 0.30) (Fig. 12); the factors FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 87 A: >40 Ka C: <18 Ka FIELDIANA: ZOOLOGY Table 31. Latitudinal range and relative tail length (tail length/head and body length) in/c/.vcvV^/rtm-group species of macaques (Fooden & Albrecht. 1999. tig. 2). Relative tail length ( % ) in adults Species Latitudinal range Mean ± SD Extremes N M. fascicularis M. cyclopis M. mulatta M. fuscata 10°S-21°N 22°N-25°N 15°N-41°N 30°N-41°N 116.9 ± 14.4 82.9 ± 6.3 44.4 ± 8.9 13.5 ± 2.7 69.2-149.5 69.2-94.7 20.0-72.1 9.5-18.6 393 26 120 7 responsible for establishing this west-east gradient in relative tail length in eastern Asia are obscure. In addition to displacing the cyclopis-\\V.e pop- ulation that formerly inhabited mainland eastern Asia, the southward dispersal of M. mulatto to the I5°-25°N latitudinal zone apparently had three further zoogeographic consequences: (1) Eastern members of the M. mulatta population (mean rel- ative tail length ca. 0.30) apparently dispersed to Hainan and two groups of smaller shallow-water islands in the South China Sea (Fig. 22B; cf. Pan et al., 1992, p. 42); during the last glacial maxi- mum, these present-day islands were connected to continental Asia by dry land (Fooden, 1995, p. 12). (2) Western members of the M. mulatta pop- ulation (mean relative tail length ca. 0.45) may have dispersed westward, perhaps marking the en- trance of M. mulatta into the Indian subregion, which ultimately may have contributed to local disappearance of sinica-group species of ma- caques (Fooden, 1989, p. 42); a Holocene fossil of M. mulatta has been reported in the Madras region (ca. 80°E; Table 29), and archaeological evidence indicates that this species was known at Moenjo Daro (27°19'N, 68°07'E) ca. 4 Ka (see "Geographic Distribution," pp. 2 ff.). (3) In the south, M. mulatta contacted M. fascicularis; this contact apparently has resulted in limited hybrid- ization between M. mulatta and M. fascicularis (Fooden, 1997, p. 226). Subsequent to the last glacial maximum, as the climate at higher latitudes ameliorated. M. mulatta in eastern Asia apparently redispersed northward and reoccupied the northern area it had previously vacated (Fig. 22C). The western population of M. mulatta also dispersed to higher latitudes in the Indian subregion, apparently retaining the relative tail length (ca. 0.45) of its putative founders (Ta- ble 5). Eastern and western populations of M. mu- latta at higher latitudes are similar in their en- hanced body size, pelage length, and pelage den- sity (Figs. 6, 8; Table 3); this similarity presum- ably is the result of parallel adaptation to cool temperature. The above scenario is compatible with major findings concerning geographic variation in mtDNA haplotypes in M. mulatta and other /a^v- cicularis group species (see above). The similarity of haplotypes in eastern M. mulatta, M. cyclopis, and M. fuscata may be attributable to the common origin of these three groups from M. fascicularis (Fig. 22A). The divergence of haplotypes in west- em M. mulatta and. independently, in Hainanese M. mulatta from those in eastern M. mulatta may be attributable to the presence of distinctive hap- lotypes in founders of the western and Hainanese populations (Fig. 22B). Acknowledgments For access to specimens and generous cooper- ation, I am deeply grateful to curators and staff members of institutions cited above (see "Intro- duction"). For helpful discussions, useful supple- mentary information, and other assistance. I also thank D. Brandon-Jones and P. D. Jenkins (British Museum [Natural History]); D. J. Melnick and J. C. Morales (Columbia University); J. Bates, L. R. Heaney, J. C. Kerbis Peterhans. B. D. Patterson, and J. Weinstein (Field Museum of Natural His- tory); Zhang Yongzu (Institute of Geography, Chinese Academy of Sciences); Liu Wanfu (For- estry Department of Guangxi); Wu Mingchuan (Forestry Designing Center of Guangxi); Feng Fig. 22. 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Wong. C. L., and I.-H. Ni. 1999. Population dynamics of the feral macaques in the Kowloon Hills of Hong Kong. American Journal of Primatology. 50: 53-66, (For date of publication, see outside back cover.) 120 FIELDIANA: ZOOLOGY Appendix 1: Specimens Examined (Total 638) Skins and Skulls, 342 AFGHANISTAN, 1. Konarha (\\ Chigha Sarai. north of— FMNH 102839. BANGLADESH, 6. Satkhiro (6): Sundarbans. ca. 50 mi east of Calcutta— zsi 11905-11907. 11925, 11931, 11985. CHINA, 137. Anhiii (1): Tong Kou— izcas 17588. Fujian (7): Chong'an Xian- — amnh 84474, 84476; Fuqing vicinity— izcas 23020, 23021; Kuatun BM(NH) 1898.11.1.29, 1900.5.8.1, mnhn 1874/481. Guangdong (41): Bawangling — .sciea Coll. No. 0409; Changtian— SCIEA Coll. Nos. 0470-0473; Dangan Dao— bm(nh) 1868.12.29.10, sciea Coll. Nos. 2150, 2151; Dongfang— izcas 12742; Hen- ron — AMNH 26646; Jianfengling — sciea Coll. No. 0089; Mihouling— sciea Coll. No. 0736; Miwan— sciea Coll. No. 2153; Nada— amnh 59987-59989, 60067, 39375; Nanfeng Shi— amnh 60038, mcz 26475; Nanwan, Xingcungang — sciea Coll. Nos. 0755, 0776; Neilingding Dao— sciea Coll. No. 2155; Nychow vicinity— bm(nh) 1870.7.18.19; Pi- sui— zmb a 194.09; Wuzhi Shan— amnh 27568- 27575, 27577, 27578, bm(nh) 1909.7.11.1; Xi Shia— izcas 17962; Xinlong— sciea Coll. No. 036; Yiajia — sciea Coll. No. 718; Zhayun — sciea Coll. No. 0251; locality unknown — mcz 19991. Guang- dong or Guangxi (1): locality unknown — mcz 20017. Guangxi (11): Lungli vicinity — private col- lection, Lungli; Nanning — kiz Coll. No. 631425; Pochuan, 6-7 km west of — izcas unnumbered; Xi Jiang, near Wuzhou — zmb A73.12; locality un- known—smnh 1126, 1127, 2106, 2108, usnm 240008, 240010. 240011. Guizhou (6): Getou— kiz 03178. 03179, 03181; Meitan, near— izcas 17959; locality unknown— kiz 000179, Coll. No. 610051. Hebei (13): Xinglong Xian, southern (= Eastern Tombs)— amnh 57038-57040, 57042. 57043, 57108, 57110, FMNH 39376-39378, mnhn 335 (281 A, 1867-557), usnm 240704, 240705. Hunan (3): locality unknown — sciea Coll. Nos. 013-015. Qingluii (2): Baima — izcas 19186; Jegu Xiang — NWPiB 00033. Sichuan (18): Gin Keo Ho, chff above — usnm 241160; Kangding — mnhn 1891/388; Leshan, mountains 30 mi southwest of — bm(nh) 1911.9.8.1; Olongche— MNHN 1891/387; Tongjiang Xian— siz 00001-00004; Tseo-Jia-Geo— usnm 256669; Wa Shan, near Dong He— rmnh 4585/ W50, 4585/V67; Xi Golog (= Singolo)— ansp 15126 (skin)/MCZ 30384 (skull); Yajiang— mcz 7922; Yibin (= Suifu)— usnm 239133; Yunnan bor- der, south of Yibin— USNM 258183. 258184; locality unknown— bm(nh) 1871.4.21.4; zmb 28919. Xizang (3): Yigong— NWPiB Coll. No. 73066; Zayii Xian— IZCAS Coll. No. 20 (external measurements question- able, provided by kx:al hunter [Quan Guogiang, iz- cas, letter, 30 October 1995]), 21. Yunnan (25): Ashi— USNM 240175, 240176; Cala Shan— izcas 17960, 17963; Datang— kiz Coll. Nos. 76318, 76320, 76321. 76324, 76325, 76344; Gengma— iz- cas 25227,25233. 25260; Hotha Valley— zsi 1 1986; Hui-Yao— AMNH 43084. 43086, mcz 26478; Man- pa— izcas 19554, 20218; Meng-ban— izcas 15054; Mengla Xian — kiz 000150; Mengyang — izcas 10303, 10304.15053; Santaishan— kiz OO0n\.Zhe- jiang ( 1 ): Huangqiao — hubd unnumbered. Province unknown (5): CHINA— bmsh 79; zsbs 17/1943; CHINA, northern— zmb 5811; CHINA, South— mcz 19988, 19990. INDIA, 60. Ammichal Pradesh (2): Dening— BM(NH) 1931.1.11.13. \9?>\.\.\\.\A. Assam (ly.Eo- gra Nadi — bm(nh) 1931.1.11.6; Golaghat — bm(nh) 1931.1.11.12; Hot Springs— bnhs 5087; Kulsi [Riv- er]— bnhs 5088; Lamsakhang — bm(nh) 1921.7.9.4; Rajapara — bm(nh) 1921.7.9.3; locality unknown — zsi 11928. Bihar (3): Luia— bm(nh) 1915.4.3.1, 1915.4.3.2, bnhs 5089. Gujarat (3): Dangs Dis- trict—bm(nh) 1931.1.11.1-1931.1.11.3. Himachal Pradesh (6): Dharmsala— bm(nh) 1933.12.1.2; Kan- gra — bnhs 5112, 5114; Kangra Fort — bm(nh) 1923.9.1.118; Samayala— bn (nh) 1931.1.11.34, 1931.1.11.35. Jammu & Kashmir (11): Dunwein — p-CM T4/2(skin), T4/8 (skull); Kashmir— bm(nh) 1871.3.3.5, usnm 63471; Kotihar— usnm 173812, 173813; Lolab— usnm 20120-20124; Lolab Val- ley— USNM 173814. Manipur (3): Bishenpur — bm(nh) 1943.60, 1943.61; Imphal, ca. 4 mi north of — ZSI 11187. Meghalaya (1): Nangpoh — bm(nh) 1931.1.11.15. Nagaland (2): Changchang Pani— AMNH 83431, 83432. Tripura (2): Ampi Bazar, ca. 3 km southeast of resthouse — zsi Coll. No. TM4; Charilam resthouse — zsi Coll. No. TM18. Uttar Pradesh (10): Bageshwar— bm(nh) 1914.7.10.1- 1914.7.10.3; Jhima— /si 12091; Ramnagar— bnhs 5108; Ratighat— bm(nh) 1914.7.10.4. 1914.7.10.5, BNHS 5109; Sita Bani— bm(nh) 1931.1.11.31, bm(nh) 1931.1.11.32. West Bengal (8): Bharnab- hari — bm(nh) 1931.1.11.8; Hasimara— bm(nh) 1916.7.29.1, 1916.7.29.2, 1931.1.1 1.9; Mangpu— FMNH 35448, 35449; Narbong— bm(nh) 1915.9.1.1; Sukna— ZSI 7294. State unknown (2): INDIA— BM(NH) 1841.12.25.1, bm(nh) 1842.4.29.55. INDIA or BANGLADESH, 2. Bengalen— nhmb 29, 31. LAOS, 8. Louangphrabang or Phongsali (1): FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE. MACACA MULATTA 121 Ou. Nam {- Nam U), between Muang Khoua and Muang Ngoy — fmnh 31763. Saravan (5): Muang Thateng— ANSP 15134-15137, 15138 (external measurements in amnh catalog, no. 87277). Vien- tiane (2): Ban Mak Nao. Camp No. 34 — zrc 4- 152; Mekong River, 90 km above Viangchan — USNM 240488. MYANMAR, 72. Chin (2): Ali Cha— bm(nh) 1931.1.1 1.22; Kindat, 20 mi northwest of— bm(nh) 1931.1.11.26. Irmwaddy (1): Pye (= Prome), 30 mi southeast of — bmhs 5081. Kachin (19): Bawmwang — bm(nh) 1950.373; Bhamo — bm(nh) 1936.12.26.4; Htingnan Triangle; — bm(nh) 1950.372; Karen Chaung— bm(nh) 1937.12.3.75; N'Changyang — bm(nh) 1950.374; Nanyaseik — AMNH 112722-112725; Singkaling Hkamti — bm(nh) 1931.1.11.25; Singkaling Hkamti, 500 ft— bnhs 5093; Singkaling Hkamti. left (east) bank — zsi 12088; Singkaling Hkamti, right (west) bank — AMNH 112988; Taga Hka— amnh 112734, 112971; Tang Hpre — usnm 279191; Tanga-Shingaw — amnh 114547; Taro— AMNH 112732, 112733. Karen (3): Toungoo. 13 mi east of — bm(nh) 1931.1.11.20, BNHS 5105; Toungoo, 15 mi north of — bm(nh) 1927.11.18.1. Mandalay (14): Kokkoaing— bm(nh) 1937.12.3.76; Lethan Hka— bm(nh) 1936.12.26.3; Maymyo — amnh 163616; Popa Hill, 1000 m — AMNH 163610-163615; Popa Hill, 4961 ft— bm(nh) 1914.7.19.2, BNHS 5102-5104, 5106. Pegu (6): Pye (= Prome), 35 mi southeast of — bm(nh) 1931.1.11.21; Toungoo, 30 mi northwest of — bm(nh) 1931.1.11.17, BNHS 5101; Toungoo, east side of Sittang River — bm(nh) 1931.1.11.16, 1931.1.11.18, 1931.1.11.19. Sagaing (20): Hein- sun— AMNH 112739, 112972; Hisweht— bm(nh) 1931.1.11.24; Homalin— bm(nh) 1915.5.5.3, 1915.5.5.4, bnhs 5094 Kin— bnhs 5095; Maung- kan — amnh 112741; Mingun — bm(nh) 1914.7.19.1; Moklok— amnh 112740; Tatkon, east bank of Chindwin River — bnhs 509 1 ; Tatkon, west bank of Chindwin River — bm(nh) 1915.5.5.6, bnhs 5090; Yin, east bank of lower Chindwin River — bm(nh) 1915.5.5.7; bnhs 5097; fmnh 82806, 82807; Yin, lower Chindwin River— bnhs 5096, 5098, 5099. Shan O): Mansam Falls— bm(nh) 1931.1.11.28, bnhs 5082, 5083, 5085; Pyaung- gaung— bm(nh) 1931.1.11.30, bnhs 5084; Se- eng— bm(nh) 1931.1.11.29. NEPAL, 10. Bagmati (4): Bouzini — bm(nh) 1931.1.11.10; Gokama— fmnh 104164; Nagarkot bm(nh) 1921.5.1.1, 1921.5.1.2. Gandaki (1): Chen- gli — bm(nh) 1931.1.11.11. Narayani (1): Hazaria patherghatta — bm(nh) 1922.5.16.2. Zone unknown (4): A^£PAZ^BM(NH) 1845.1.8.222-1845.1.8.224, 1972.1015. PAKISTAN, 9. North-west Frontier (4): Dunga Gali — USNM 353186; Ghora Dhaka, 1 mi east of — usnm 326332; Kaotai— bm(nh) 1920.6.11.1; Paia — USNM 353187. Punjab (2): Patriata — bm(nh) 1923.11.4.1, BNHS 5113. Province un- known (3): "Karrachi" — irsn 6857-6859. THAILAND, 14. Chiang Mai (3): Chiang Dao— Mcz 37706; Chiang Mai, near — zrc 4-154; Kaeng mae Hat (= Me Ping rapids) — zrc 4-188. Kam- phaeng Phet (1): Ban Umphang, 28 mi southeast of — AMNH 54816. Loei (3): Dan Sai District — usnm 300017, 307715 307716. Nan (1): Pang Nam Un— usnm 296917. Nong Khai (2): Nong Khai, Camp No. 28— ZRC 4-150, 4-151. Tak (4): Ban Mae La- mao — FMNH 99668: Huai Ap Nang — fmnh 99669; Huai Kwang Pah — ctnrc (formerly fmnh 99670); Tha Chang Tai— zrc 4-153. VIETNAM, 23. Bac Thai (3): Bac Can— bm(nh) 1927.12.1.18, 1927.12.1.20, mnhn 1929/456. Hai Phong (2): Ang Co— zmvnu 01/3.61. 40/Pc 40; Cat Ba, Dao— zmvnu 05/3.163.0. Hoa Binh (1): Hoa Binh— lEBR 70/1238/11. Kon Turn (3): Dak Sut— usnm 320780-320782. Lai Chau (5): Bac Tan Trai — ansp15133; Muong Boum — fmnh 31766; Muong Moun — fmnh 31764; Muong Muon-AMNH 87278; Muong Pon— amnh 87264. Nghe An (3): Nghia Dan — bm(nh) 1928.7.1.11; Nighia Dung — lEBR 520/145/695, 733(833)/560/175. Quang Nam- Da Nang (2): Son Tra. Mt., 3.9 km west and 0.3 km south of — usnm 356968; Song-Ta-Voy — mnhn 1899/54. Tuyen Quang (3): Chiem Hoa — zmvnu 167/3.18.14; Thanh Tuong-IEBR 48, 62 (external measurements of both in Dao, 1985, p. 36). Prov- ince unknown (1): VIETNAM— \ebr 2241. Skins only. 154 AFGHANISTAN, 1. Konarha (1): Nurestan ( = Kafirstan). eastern — bm(nh) 1931.1.9.1. BANGLADESH, 1. Satkhira (1): Sundarbans, ca. 50 mi east of Calcutta — zsi 11984. CHINA, 84. Anhui (9): Tong Kou— izcas 18068; Tunxi — smnh 23; locality unknown — SMNH 14, 25, 26, 32, 33, 35, 463. Fujian (4): Kua- tun — MNHN 1874/480; Pucheng Xian — sciea Coll. No. Min 008; Sha Xian— sciea Coll. No. Min 01 1; locality unknown — sciea Coll. no. Min 009. Guangdong (2): Dongfang — fubd 156; Hainan Dao — zmb 43500. Guangxi (24): Batu — fdcg A 005; Dongmen — izcas unumbered; Longman — FDCG unnumbered (skull inside); Xunle vicinity — 122 FIELDIANA: ZOOLOGY private collection, Xunle, three unnumbered skins; Zhongzhou — fdcg C 0014 (skull inside); locality unknown — smnh 15, 19, 20. 24. 28. 29. 367, 368, 484, 856. 915, 916, 1002, 1168, 1425. 2107, usNM 240012. Guizhou (5): Zunyi vicini- ty—bmnh 5.66.150—5.66.153, izcas 20770. He- bei (1): Xinglong Xian (= Yungling), southern — bm(nh) 1931.1.7.2. Qinghai (1): Jegu Xiang — NWPiB Coll. No. 63167. Sichuan (2): Batang vi- cinity— IZCAS 20219; Dege vicinity — izcas 25940. Sichuan or Xizang (2): Tibet — mnhn 1892/315, 1894/1432. Xizang (1): Zayu Xian— izcas 73248. Yunnan (12): Biloxue Shan — Kiz Coll. no. 780417; Menghai— izcas 15052; Mengla Xian— Kiz 03174, 03180; Menglun— kiz Coll. No. 75840, unnumbered; Shanman — kiz 000153: Tengchong (= Momien) — izcas 17586, zsi 619; Xishuangbanna — Kiz 03172, 03173; Yongde vi- cinity— IZCAS 21513. Zhejiong (1): Xindeng — ZMNH Coll. No. 633 (skull inside). Province un- known (20): C///A^A— BM(NH) 1931.1.7.1, izcas 17954, 18121, 18123, 24914, sciea Coll. No. 0003. SMNH 16, 17, 21, 22, 27 (on exhibit). 30. 31. 38, 39, 458, 479, 914, 1162; ?CHINA—MN\\n unnumbered. INDIA, 22. Andhra Pradesh (1): Siddeldar Hill — ZSI unnumbered. Arunachal Pradesh (2): Dening — bnhs 5086; Margherita — zsi 12090. As- sam (1): Rajapara — bm(nh) 1931.1.11.7. Jammu & Kashmir (1): Dunwein — p-cm T4/1 (on exhibit, skull inside). Madhya Pradesh (3): Kakara — bm(nh) 1931.1.11.5; Malua — bnhs 5107; Sohag- pur — bm(nh) 193 1 . 1 . 1 1 .4. Nagaland ( 1 ): Samagut- ing — zsi 11987. Orissa (3): Deogarh — zsi Coll. No. OM/D/30; Gudari— bm(nh) 1928.3.7.4; Mal- kangiri— bm(nh) 1928.3.7.3. Sikkim (2): locality unknown— BM(NH) 1891.10.7.4, 1891.10.7.5 (skull inside). Uttar Pradesh (3): Dela — bm(nh) 1931.1.11.33; Haripur — bnhs 5111; Ramnagar — bm(nh) 1951.609. West Bengal (2): Hasimara — BNHS unnumbered (on exhibit); Sivok — fmnh 35447. State unknown (3): INDIA— bm(nh) 1842.12.19.14. 1926.2.9.1; 7INDIA—bm(nh) 1851.8.29.4 (skull inside). INDIA or BANGLADESH, 1. "(Bengal)"— ansp 3950. MYANMAR, 1 1 . Kachin (3): Hkandau— bm(nh) 1950.376; Karen Chaung— bn (nh) 1937.12.3.77; N'Changyang — bm(nh) 1950.375. Mandalay (1): Madaya— BM(NH) 1936.12.26.5. Pegu (2): Toun- goo, 20 mi west of — bnhs 5100; Toungoo, 30 mi northwest of — zsi 12089. Sagaing (3): Tatkon, near Kindat, west bank of Chindwin River — bm(nh) 1931.1.11.23, bnhs 5092; Yin, east bank of lower Chindwin River — zsi 12094. Shan (2): Mansam Falls— BM(NH) 1931.1.11.27, 1972.836 (in alco- hol). NEPAL, 4. Bagmati (4); Gokarna— i PS Coll. No. 529 (formerly i mnh 104163); Trisuli Bazar, 4 mi southeast of^iMNH 135427-135429 (all three in alcohol). THAILAND, 1. Tak (1): Huai Kwang Pah— FMNH 99671 (in alcohol). VIETNAM, 29. Bac Thai (4): Bac Can— bm(nh) 1927.12.1.19; Ban Thi— zmvnu 07; Linh Thong— ZMVNU 08/3.20.72, 26/3.19.71. Cao Bang (3): Ly Bon — lEBR 32, 33, 34 (external measurements in Dao, 1985, p. 46). Ha Tinh (1): Ky Son— ihbr 40/ 441. Hai Phong (3): Cat Ba, Dao— zmvnu 02/ 3.66.11, 03/3.62.9, 04/3.64.0. Hoa Binh (2): Phu Vach — IFBR 1275/M44; locality unknown — zmvnu 28/3.81.Pc7 (mismatched with M. arcto- ides skull). Lai Chau (1): locality unknown — if.br D. 3/M37. Quang Binh (1): Xuan Ninh— ihbr 560. Quang Ninh (2): Van Hai, Dao — zmvnu 06/ 3.16.4, 537/3.17.5. Yen Bai (1): locality un- known— iebr D1 (possibly belongs with iebr 835/ 36/199, skull only, Thuong Bang La). Province unknown (11): Tonkin— mnhn 1887/11; VIET- NAM— iebr unnumbered (6), zmvnu 09/3.15.3, 10/3.22, 731, 732. Skulls only, 142 CHINA, 52. Anhui (1): Locality unknown — smnh 1514. Fujian (1): Kuatun— bm(nh) 1897.6.5.2. Guangdong (3): Dongfang — ihbd 156; Xi Shia — izcas 17961, 17964. Guangxi (19): Bamo village, near — izcas unnumbered; Bapon — iixx; P 037; Batu— fdcg a 004; Ditin— fikg A 008; Hechi Pre- fecture— IZCAS unnumbered (3); Jenli. 2-3 km north of — private collection, Jenli; Liuzhai vicinity-pri- vate collection, Liuzhai; Piangzu, 3.5 km northeast of Banli — private collection, Banli; Tian'e Xian — IZCAS unnumbered (3); Xianan. Huanjiang Xian — private collection, Huanjiang; Xianan-Mulun — pri- vate collection. Mulun; Yuhun — Fixo unnumbered (2); locality unknown — smnh 1186. 1515. Guizhou (1): Fameng— KIZ Coll. No. 631094. Hebei (1): Xinglong Xian. southern — usnm 240703. Sichuan (2): Olongche — mnhn 1891/389; Yunnan border — usnm 253780. Xizang (3) Zayu Xian — nwpib Coll. Nos. 73230. 73232, 73233. Yunnan (9): Jinping— IZCAS 04990; Manpa— kiz Coll. Nos. 80853, 80854; Menghai — izcas 10305; Mengla Xian — Kiz Col. No. 00; Nonglin— KIZ Coll. No. 592005; Tengchong Xian-Kiz Coll. No. 76323; locality unknown — izcas FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE. MACACA MULATTA 123 17947, 25953. Zhejkmg (1): Zhoucun— zmnh Coll. No. 85002 (skin unavailable in October 1985). Province unknown (11): China — bmnh 49, smnh 1516-1519, 1775 2091, 2096, 2099-2101. INDIA, 16. Assam (2): Cachar District — bnhs 6246; locality unknown — liM(NH) 1858.5.4.247. Himaclwl Pradesh (1): Simla vicinity — bnh.s 5110. Jammii & Kashmir (1): Kashmir — bm(nh) 1856.5.6.12 (identification questionable). Uttar Pradesh (6): Bageshwar — bm(nh) 1914.7.10.6, 1914.7.10.7, Bijnor— bm(nh) 1926.10.8.8; Ram- nagar— bm(nh) 1914.7.10.8, 1914.7.10.9; Sak- tesgarh— BM(NH) 1848.2.1.26. West Bengal (3): Calcutta, Indian Museum Compound — zsi 15293; locality unknown — bm(nh) 1856.5.6.18, BNHS Coll. No. 5321. State unknown (3): IN- DIA— bm(nh) 1843.5.27.2, zsi unnumbered (two, including female skull mismatched with male skin collected at Siddeldar Hill). INDIA or BANGLADESH, 1. Bengalen — mzb 6732. LAOS, 1. Louanqphrabanq (1): Ou, Nam ( = Nam hou)— MNHN 1892/1357. MYANMAR, 9. Chin (1): Chittagong Hill Tracts— BM(NH) 1926.10.8.7. Kachin (3): MYANMAR, up per — bm(nh) 1950.377; Singkaling Hkamti — BM(NH) 1915.5.5.5, 1972.1333. Mandalay (1): Popa Hill — BNHS 6249. Sagaing (2): Chindwin River — BNHS 6247; Kindat— bm(nh) 1910.10.19.5. Shan(\): Pyaunggaung — bnhs 6248 (locality and date in- ferred from collector's number). Tenasserim (1): locality information probably inaccurate — zmb A161.12(2). NEPAL, 5. Bagmati (3): Trisuli Bazar, 4 mi south- east of — FMNH 104165-1041 167. Zone unknown (2): A^£PAZ^BM(NH) 1845.1.8.5, 1858.6.24.144. VIETNAM, 58. Bac Thai (4): Ban Thi— zmvnu 185/3.162.0 (identification tentative), Cho Don District — fcxm Oil; Thai Nguyen zmvnu — 172/ 3.151.Pc23, 196/Pc22. Cao Bang (4): Ban Vay— zmvnu 186/3.169.0; Ly Bon— iebr T33; Po Lu— zmvnu 184/3.165.61; Trung Khanh District— FCXM 43 (with mismatched mandible). Ha Trinh (2): Huong Son — fcxm 05; Trai Tru — ihbr 63/ 622 (external measurements in Dao, 1985, p. 244). Hai Phong (4): Ang Co-zmvnu 174/3.161. Pc52/38; Cat Ba, Dao— zmvnu 175/3.1 58. Pc49, 176/3.159.Pc50, 177/3.160.Pc51. Lang Son (1): locality unknown — zmvnu 180/Pc25. Nghe An (4): Ban Bu — fcxm 015 (external measurements in card catalog); locality unknown — zmvnu 168/ 3.153.Pc31, 169/3. 155.Pc33, 170/3.154.Pc32. Ninh Binh ay. Cue Phuong— zmvnu 64/3.148.Pc35, 173/ 3.149.Pcl9 (both identifications tentative). Qiiang Binh (1): Bo Trach District — iebr 1431. Quang Nam-Da Nang (2); Son Tra, Mt— usnm 356978 (mandible missing), 356979 (external measurements in collector's fieldbook). Quang Ninh (4): Quan Lan. Dao— iebr T15; Van Canh, Dao— iebr T.13, 14 (identification tentative); Van Hai, Dao — ^zmvnu 178/3. 152.Pc27. Vinh Phii (1); Thanh Son— zmvnu 171/33.150.PC.21. Yen Bai (3): Nam Ngap— iebr 18, T19; Thuong Bang La— iebr 835/36/199. Prov- ince unknown (26): Tu Chi— zmvnu 189/34; VI- ETNAM— FCXM 028 (identification tentative), iebr 440, 2311, 2335, 2358, 2359, unnumbered (4), ZMVNH 179/3. 157.Pc46, 181/3.164.0 (identification tentative), 182/3.166.0, 183/3.167.0, 187/3.168,0, 190/35, 191/36, 192/37, 193/38, 194/39, 195/40, 541/3.156. Pc45, unnumbered; [V/£77VAM]— mnhn 1962/1439, 1962/1448. Appendix 2: Gazetteer of Macaca mulatta Localities Locality names listed as primary entries in this gazetteer preferentially are official names ap- proved in gazetteers published by the U.S. Board on Geographic Names (USBGN; Afghanistan, 1971; Bangladesh, 1976; Burma [= Myanmar], 1966a; China, 1979; India [includes Bhutan and Nepal], 1952; Laos, 1973; Pakistan, 1983; Thai- land, 1966b; Vietnam, 1986). In addition, supple- mentary references have been consulted for lo- calities in China (Administrative Divisions of the People's Republic of China— 1980, 1981; Zong- hua Renmin Gongheguo Fen Sheng Dituji, 1983), India (National Atlas of India, 1979; Census of India 1981 — various district census handbooks), and Vietnam (Cue Ban Do-Bo Tong Tham Muu, Quan Doi Nhan Dan Viet Nam, 1980-88). Lo- calities of M. mulatta that were not found in USBGN gazetteers or supplementary references are spelled here as in the original sources. Sec- ondary entries, with cross references to corre- sponding primary entries, indicate variant spell- ings or alternate locality names that appear on specimen tags, in published literature, or in un- published notes concerning M. mulatta. The sequence of information presented in pri- mary entries is as follows: 1. Locality name. (Note: Chinese locality names frequently include the following generic geo- graphic terms: Dao - Island; Shan = Moun- tain; Xian = County.) 124 FIELDIANA: ZOOLOGY 2. Altitude, if reported by collector or observer. 3. Name of province, state, or other first-order administrative division, in italics. 4. Name of country, in capital letters. 5. Coordinates of locality (principal sources: USBGN gazetteers, supplementary references indicated above, published or unpublished field notes of collectors or observers). 6. Date of collection or observation. 7. Name of collector or observer. 8. Bibliographic reference (in parentheses) to published or unpublished field notes, if any. 9. Abbreviated name of museum (see "Intro- duction") where specimens are preserved. 10. Number of specimens available (with indi- cation of part preserved, if skin and skull are not both present). 1 1 . Locality code as indicated in distribution maps (Figs. 2A-C). Achaltal. See Achal Tank. Achal Tank, Aligarh; Uttar Pradesh, INDIA; 27°53'N, 78°05'E; observed Sept. 1959-June 1960 and Dec. 1970-July 1972 by C. H. South- wick, M. A. Beg, and M. R. Siddiqi (1961a, p. 543; Southwick, 1962, p. 437; Southwick et al., 1976, p. 13). Observed Jan. 1990-Mar. 1991 by E. Imam and H. S. A. Yahya (1995, p. 2). A:I- 69. Adupuria; Assam, INDIA; ca. 26°48'N, 94°45'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. AFGHANISTAN, eastern; 33°-37°N, 69°-72°E; reported before 1971 by E. Kullmann (1970, p. 23). Not mapped. AFGHANISTAN, northeastern; 33°-37°N, 69°- 72°E; reported Oct. 1808-Aug. 1809 by M. El- phinstone (1842, p. 188). Not mapped. Agartala. See Charilam. Aglar River; Uttar Pradesh, INDIA; ca. 30°35'N. 78°05'E; observed 30 Apr. 1944 by H. Harrer (1982, p. 40). A:I-27. Agra; Uttar Pradesh, INDIA; 27°irN. 78°0rE; trapped before 1993 by K. Waheeda (1992, p. 111). A:I-70. Agra District; Uttar Pradesh, INDIA; 26°50'- 27°20'N, 77°30'-78°30'E; observed 1981-1983 by R K. Seth. S. Seth, G. L. Reddy, and R K. Chopra (1992, p. 62). Not mapped. Ailao Shan Reserve; Yunnan, CHINA; ca. 24°15'N, 101°18'E; observed in 1990 by L. K. Sheeran and F E. Poirier (1994. p. 21). B:C-72. Ajanta Caves vicinity; Maharashtra, INDIA; ca. 20°33'N, 75°42'E; occurrence before 500 A.D. inferred from cave painting (Fooden et al., 1981. p. 465; Marathe and Mahabal, 1984, p. 74). A:I-100. Ajodhya. See Ayodhya. Akhnoor; Jammii & Kashmir, INDIA; 32°54'N, 74°44'E; observed before 1983 by Y. R. Mal- hotra and D. N. Sahi (1982. p. 27). A:I-5. Akhoiphutia; Assam. INDIA; ca. 27°03'N, 94°39'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991aJ. p. 31). B:I-25. Ale Chaung. See Ali Cha. Ali Cha, Chin Hills, 20 mi (= 32 km) southwest of Kindat, 1000 ft (= 300 m); Chin, MYAN- MAR (= BURMA); ca. 23°40'N. 94°05'E; col- lected 3 Jan. 1915 by J. M. Mackenzie (Wroughton. 1916c. p. 759); bm(nh). 1. B:M- 17. Alicheng River, east of; Laghman, AFGHANI- STAN; ca. 34°55'N, 70°05'E; reported before 1972 by A. Puget (1971, p. 201). A:A-5. Aligarh; Uttar Pradesh, INDIA; 27°53'N, 78°05'E; observed Dec. 1970-July 1972 by C. H. Southwick, M. F Siddiqi, M. Y Farooqui, and B. C. Pal (1976, p. 13). Observed Jan. 1990-Mar. 1991 by E. Imam and H. S. A. Yah- ya (1995, p. 2) A:I-69. Aligarh District; Uttar Pradesh. INDIA; 27°30'- 28°10'N, 77°30'-78°35'E; observed 1959-1984 by C. H. Southwick and M. F Siddiqi (1983, p. 229; Southwick, 1985. p. 191). Not mapped. Aligarh vicinity; Uttar Pradesh, INDIA; ca. 27°53N. 78°05'E; blood samples obtained 16- 27 Apr. 1964 by K. V. Shah and C. H. South- wick (1965, p. 489). A:I-69. Alingar. east of; Laghman, AFGHANISTAN; ca. 34°55'N. 70°30'E; reported before 1972 by A. Puget (1971, p. 201). A:A-6. Ali Sagar. Nizamabad District, 440 m; Andhra Pradesh, INDIA; 18°42'N, 78°00'E; observed 16 Apr. 1980 by J. Fooden. A. Mahabal. and S. S. Saha (1981, p. 466). A:I-116. Alwar District; Rajasthan, INDIA; ca. 27°38'N, 76°35'E; observed 1981-1983 by P K. Seth, S. Seth, G. L. Reddy, and P K. Chopra (1992, p. 62). A:I-73. Ambagarh Reserve Forest, 9 km northeast of Jai- pur; Rajasthan, INDIA; ca. 27°00'N, 75°53'E; observed 1985-1993 by A. Lobo. B. R. Ma- nohar, and R. Mathur (Mathur & Lobo, 1990, p. 308; Manohar & Mathur, 1992. p. 114; Ma- thur. 1994, p. 132). A:I-75. Ambala District; Haryana, INDIA; ca. 3()°5rN, 76°30'E; observed 1981-1983 by P K. Seth, S. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 125 Seth, G. L. Reddy, and P. K. Chopra (1992, p. 62). A:l-20. Amber. See Amer. Amen Jaipur District; Rajasthan, INDIA; 26°59'N, 75°52'E; observed 1975-1980 by P. K. Seth and S. Seth (1983, p. 63). A:I-75. Ampi Bazar, ca. 3 km southeast of resthouse; Tri- pura. INDIA; ca. 26°40'N, 9r38'E; collected 18 Jan. 1971 by V. C. Agrawal (Agrawal & Bhattacharyya, 1977, p. 137); zsi, 1. 8:1-40. Anapalam, Rajupalem Taluk, Guntur District; An- dhra Pradesh, INDIA; ca. I6°20'N, 80°00'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). A:I-145. Angaluru, Guntur District, 125 m; Andhra Pra- desh, INDIA; 16°12'N, 79°47'E; observed 30 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 468). A:I-147. Anganganj. See Khair, Tahsil. Ang Co, Cat Ba Island; Hai Phong, VIETNAM; 20°43'N, 107°03'E; purchased in market 13 July 1964 by unknown collector (Dang, 1983, p. 1282; Dao, 1985, p. 82; Nisbitt & Ciochon, 1993, p. 772); zmvnu, 1. Collected 11 Aug. 1964 by Hien Hao (Dao, 1985, p. 82; Nisbitt & Ciochon, 1993, p. 772); zmvnu, 1 (skull only). C:V-14. Anhui, CHINA; 29°-35°N, 115°-120°E; collected 1959-1960 by museum collectors; smnh, 8 (7 skins only, 1 skull only). Not mapped. Anji Xian; Zhejiang, CHINA; ca. 30°40'N, 119°40'E; ca. 50 captives acquired 1950-1959 from Local Products Supply Co. by Hangzhou Zoo (Fu Yiyuan, Director, pers. comm., 25 Oct. 1985). C:C-56. Ankapur, Armur Taluk, Nizamabad District; An- dhra Pradesh, INDIA; 18°45'N, 78°16'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-115 Anlong; Guizhou, CHINA; 25°06'N, 105°3rE; reported before 1998 (Zhang et al., 1997, p. 58). C:C-157. Annapurna Conservation Area. See Pokhara. Annavangal, Banswada Taluk, Nizamabad Dis- trict; Andhra Pradesh, INDIA; not precisely lo- cated, 18°05'-18°35'N, 77°45'-78°05'E; report- ed Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). Not mapped. Anyuan; Jiangxi, CHINA; 25°09'N, 1 15°2rE; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-98. Arbesi. See Basirhat Reserve Forest. Anmachal Pradesh, INDIA; 26°55'-29°30'N, 91°35'-97°25'E; reported during "past few years" by A. Borang and G. S. Thapliyal (1993, p. 839). Not mapped. Asarori Forest, 425-950 m; Vttar Pradesh, IN- DIA; ca. 30°15'N, 78°00'E; ob.served June 1965-May 1966 by D. G. Lindburg (1971, p 5). Observed June 1973-Oct. 1976 by S. C Makwana, R. S. Pirta, and M. Singh (Pirta 1977-78, p. 125; 1984, p. 272; Makwana, 1978 p. 483; 1979a, p. 242; Makwana & Pirta, 1978 p. 164; 1983, p. 301; Pirta & Singh, 1981, p 340). A:I-27. Ashi; Yunnan, CHINA; ca. 26°53'N, 100°00'E; collected in 1921 (see usnm catalog) by J. F. Rock (1925, p. 447; 1926, p. 139; Chock, 1963, p. 93); USNM, 2. B:C-44. Ashoknagar, Warangal District, 340 m; Andhra Pradesh, INDIA; 17°55'N, 79°57'E; observed 19 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A:I-133. Asmar, east of; Konarha, AFGHANISTAN; ca. 35°00'N, 71°30'E; reported before 1972 by A. Puget (1971, p. 201). A:A-1. Asmar, northwest of; Konarha, AFGHANISTAN; ca. 35°10'N, 71°20'E; reported before 1972 by A. Puget (1971, p. 201). A:A-1. Assam, INDIA; 24°10'-28°00'N, 89°40'-96°00'E; acquired before 1859 by Zoological Society of London; bm(nh), 1 (skull only). Acquired be- fore 1956 by Calcutta Zoological Garden (Kha- juria, [1955], pp. 113, 114); zsi, 1. Not mapped. Assembly. See Simla. Aum River, ca. 20 km above mouth, Karnali Bar- dia Game Reserve; Bardia, NEPAL; ca. 28°30'N, 81°19'E; observed 20 Feb.-lO Mar. 1976 by J. Teas (1983, p. 214). A:N-5. Ayodhya; Uttar Pradesh, INDIA; 26°48'N, 82°12'E; reported before 1978 by M. L. Roon- wal and S. M. Mohnot (1977. p. 100). A:I-60. Ayodhya-Gorakpur, highway between; Uttar Pra- desh, INDIA; ca. 26°45'N, 82°45'E; observed Sept. 1959-June 1960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961b, p. 702). A: 1-62. Azamgarh vicinity; Uttar Pradesh, INDIA; ca. 26°04'N, 83°irE; observed Sept. 1959-Feb. 1960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961a, p. 540). A:I-63. Babai River vicinity, Karnali Bardia Game Re- serve; Bardia, NEPAL; ca. 28°26'N, 8I°2rE; observed 20 Feb.-IO Mar. 1976 by J. Teas (1983, p. 214). A:N-5. Babuwali, Jasalmer District; Rajasthan, INDIA; ca. 26°47'N, 69°44'E; falsely reported 4 Sept. 126 FIELDIANA: ZOOLOGY 1980 by K. Singh (Bhargava. 1982. p. 7; 1984. p. 43). Not mapped. Bac Can. Tonkin region. 500 ft (= 150 m); Bac Thai, VIETNAM; 22°08'N. 105°49'E; collected 13 Dec. 1926-14 Jan. 1927 by J. Delacour and W. P. Lowe (Delacour & Jabouille, 1927, p. 302); BM(NH). 3 (including 1 skin only): mnhn, 1. C:V-10. Bachepalli. See Dachepalle. Bachi, Pingyuan [Xian]; Guangdong, CHINA; 24°46'N. 115°49'E; reported before 1998 (Zhang et al.. 1997. p. 58). C:C-97. Bach Ma National Park; Thua Thien-Hue, VIET- NAM; ca. 16°12'N. 107°52'E; reported ca. 1990-1995 by L. K. Lippold (1995. p. 199). C: V-35. Bac Tan Trai (= Bac Tan Tray). Tonkin region; Lai Chan, VIETNAM; 22°24'N, 103°12'E; col- lected 5 Nov. 1931 by T. D. Carter (Legendre. 1936. p. 83); ansp. 1. C:V-1. Badu. See Batu. Bageshwar (= Bageswar). Kumaun region, 3200 ft (= 975 m); Uttar Pradesh, INDIA; 29°51'N, 79°46'E; collected Aug. 1913-Mar. 1914 by C. A. Crump (in Wroughton. 1914. p. 283); bm(nh), 5 (including 2 skulls only). A:I-31. Baguri Block, Kaziranga Wild Life Sanctuary; As- sam, INDIA; ca. 26°37'N. 93°15'E; reported Jan.-June 1966 by J. J. Spillett (1967. p. 496). B:I-21. Bahgara; Assam, INDIA; ca. 26°53'N, 94°45'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a]. p. 31). B:I-25. Baidonghe Water Regulation Forest Reserve; Guangxi, CHINA; ca. 23°55'N, 106°44'E; ob- served in 1976. 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 127; letter. Aug. 1996). C:C-169. Baima; Qinghai, CHINA; ca. 33°02'N. 100°04'E; collected 1 1 Sept. 1967 by Wang Zongyi; izcas. 1. B:C-23. Bairaglia. See Hazaria Patherghatta. Baisha. Hainan Dao; Hainan, CHINA; 19°13'N. 109°26'E; reported before 1998 (Zhang et al.. 1997. p. 58). C:C-233. Baishuijiang Natural Reserve, 1700-2900 m and 3000-3400 m; Shaanxi, CHINA; ca. 33°35'N, 105°54'E; reported before 1989 by Ma Guoyao (1988, p. 27). C:C-18. Baizha Plantation, Nangqen Xian, 3600-4400 m; Qinghai, CHINA; 32°04'N, 96°21'E; observed in early 1960s by Zheng Changlin (pers. comm.. 7 Oct. 1985). B:C-15. Baj Garhi Bridge; Uttar Pradesh, INDIA; 28°03'N. 78°03'E; observed 1959-1975 by C. H. Southwick and M. F Siddiqi (1977. p. 342). A:I-69. Bak shoi mun. See Luofu Shan. Balimila (= Balimela) vicinity; Orissa, INDIA; ca. 18°15'N. 82°08'E; observed 1959-1970 by M. Krishnan (1972. p. 540). A:I-109. Balkonda. Armur Taluk. Nizamabad District; An- dhra Pradesh, INDIA; 18°52'N. 78°2rE; ob- served 1972-1973 by N. Koyama and P B. Shekar (1981. p. 248). Reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-115. Ballapet. Khammam Taluk. Khammam District; Andhra Pradesh, INDIA; 17°16'N, 80°12'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984. p. 58; 1992, p. 18). A:I-132. Balphakram region; Meghalaya INDIA; ca. 25°10'N, 90°15'E; observed ca. 1977 by A. K. Ghosh and S. Biswas (1977, p. 24). B:I-12. Balrampur forest; Uttar Pradesh, INDIA; ca. 27°26'N, 82°irE; observed 1973-1974 by R. K. Singh and N. N. Sen (1977-78, p. 136). A: 1-61. Balthali, Kavre; East No. 1, NEPAL; 27°30'N, 85°30'E; observed May 1998 by M. K. Chalise and M. Ghimire (1998, p. 12). A:N-supplemen- tary. Bamo village, near, Bamo Subcounty. Tiane Xian; Guangxi, CHINA; 24°55'N. 107°21'E; collected in 1987 by Yang Changbi, subsequently pur- chased by Huang Runqiang (pers. comm., 24 Oct. 1992); IZCAS, 1 (skull only). C:C-173. Ban Bu, Khe Choang (river). Con Cuong District, 500 m; Nghe An, VIETNAM ca. 19°03'N, 104°45'E; collected 11 Dec. 1992 by Pham Nhat and Mr. Hung; fcxm, 1 (skull only; exter- nal measurements recorded in card catalog). C: V-26. Ban Bung vicinity, Na Hang District; Tuyen Quang, VIETNAM; ca. 22°20'N, 105°20'E; reported 25 Feb.-5 Apr 1992 by R. Ratajszczak, Ngoc Can, and Pham Nhat (1992, p. 16). C:V-6. Banda District; Uttar Pradesh, INDIA; 24°55'- 25°55'N, 80°05'-81°35'E; observed before 1916 by G. B. F Muir (1916, p. 353). Observed 1960-1980 by M. F Siddiqi (Southwick, 1985, p. 191 ). Not mapped. Banda vicinity; Uttar Pradesh, INDIA; ca. 25°29'N, 80°20'E; observed Sept. 1959-Feb. 1960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961a. p. 540). A:I-84. Bandhavgarh National Park; Madhya Pradesh, INDIA; ca. 23°40'N, 81°02'E; reported Feb.- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 111 Mar. 1976 by N. K. Sinha (1977, p. 5). Re- ported before 1997 by K. K. Gurung and R. Singh (1996. p. 80). A:I-93. Bandipul, Tahsil Thana Ghazi; Rajasthan, INDIA; ca. 27°25'N. 76°19'E; observed 1975-1980 by R K. Seth and S. Seth (1983, p. 63). A:I-74. Bangma; Sichuan, CHINA; not located, 26°00'- 34°10'N, 97°40'-110°05'E; reported before 1992 by Jiang Xuelong. Wang Yingxiang, and Ma Shilai (1991, p. 243). Not mapped. Bangram. See Dhaka. Bangsal. See Dhaka. Ban Kuai, Several km south of, 170-609 m; Vien- tiane. LAOS; ca. 18°16'N, 102°08'E; observed Feb.-July 1996 by J. W. Duckworth (1996, p. 229). B:L-6. Ban Mae Lamao, 350 m; Tak, THAILAND; 16°48'N. 98°45'E; collected 21 Mar. 1967 by J. Fooden (1971, p. 18). fmnh, 1. B:T-6. Ban Mak Nao, Camp No. 34; Vientiane, LAOS; 18°00'N, 102°58'E; collected 16 Feb. 1920 by J. Bangassar (Weitzel et al., 1988, p. 116); zrc, 1. B:L-8. Ban Manao. See Ban Mak Nao. Bannabari. See Bhamabhari. Ban Napo vicinity, 170-609 m; Vientiane, LAOS; ca. 18°17'N, 102°irE; observed Feb.-July 1996 by J. W. Duckworth (1996, p. 229). B:L-6. Ban So vicinity, 170-609 m; Vientiane, LAOS; ca. 18°19'N. 102°06'E; observed Feb.-July 1996 by J. W. Duckworth (1996, p. 229). B:L-6. Ban Thi, Cho Don District; Bac Thai, VIETNAM; 22°14'N, 105°3rE; collected 2 May 1970 by Nguyen Trong Tien; zmvnu, 1 (skull only, spe- cies identification tentative). Collected 20 May 1970 by Mr. Nhe; zmvnu, 1 (skin only). C:V-6. Ban Umphang, 28 mi (= 45 km) southeast of, 1750 ft (= 530 m); Kamphaeng Phet, THAI- LAND; ca. 15°28'N, 99°04'E; collected 4 Feb. 1924 by A. S. Vemay (Lowe, 1932, p. 197; 1933, p. 260); amnh, 1. B:T-8. Ban Vay (= Ban Vai). Khang Ninh vicinity; Cao Bang, VIETNAM; 22°27'N, 105°39'E; collect- ed 12 Aug. 1971 by Nguyen Van Chau; zmvnu, 1. C:V-8. Ban Wangma vicinity, 170-609 m; Vientiane, LAOS; ca. 18°23'N, 102°06'E; observed Feb.- July 1996 by J. W. Duckworth (1996, p. 229). B:L-6. Baoshan; Yunnan, CHINA; 25°07'N, 99°09'E; tis- sue sample obtained ca. 1991 (Zhang & Shi, 1993b, p. 589). Immunological survey con- ducted before 1996 by Duan Xingsheng, Liu Yuanwei, Wu Jing, Dao Weiying, and Liu Jianghai (1995, p. 411). B:C-56. Bapon, Fusui Xian; Guangxi, CHINA; ca. 22°35'N, 107°57'E; collected Aug. 1986 by Wu Mingchuan (pers. comm., 27 Nov. 1992); fdcg, 1 (skull only). C:C-219. Bara Math Temple. See Chitrakut. Barami. See Barmi. Barauli Bridge; Uttar Pradesh, INDIA; ca. 28°05'N, 78°03'E; observed 1959-1975 by C. H. Southwick and M. F Siddiqi (1977, p. 342). A:I-69. Barautha. See Barotha. Bar Chanrai Hill, lower Swat Valley opposite Ma- lakand; North-West Frontier, PAKISTAN; ca. 34°34'N, 71°56'E; reported before 1902 by A. H. McMahon (1901b, p. 9). A:P-6. Bardia National Park. See Babai River vicinity. Bareilly; Uttar Pradesh, INDIA; 28°21'N, 79°25'E; trapped for psychological study before 1993 by K. Waheeda (1992, p. 111). A:I-52. Bareilly-Agra, highway between; Uttar Pradesh, INDIA; ca. 28°50'N, 78°30'E; observed 1964- 1965 by R. P. Mukherjee and G. D. Mukherjee (1972, p. 67). A:I-48. Barikowt (= Baricot), southeast of; Konarha, AF- GHANISTAN; ca. 35°10'N, 71°35'E; reported before 1972 by A. Puget (1971. p. 201). A:A-I. Barmdeo Mandi; Kanchanpur, NEPAL; 28°52'N, 80°09'E; observed June 1964-Dec. 1965 by D. L. Chesemore (1970, p. 164). A:N-2. Barmi; Gazipur, BANGLADESH; ca. 24°08'N, 90°22'E; observed Sept. 1975 by J. R. Oppen- heimer, A. K. Akonda, and K. Z. Husain (1983, p. 194). B:Ba-15. Barotha; Uttar Pradesh, INDIA; 27°57'N, 78°10'E; observed 1959-1975 by C. H. South- wick and M. F Siddiqi (1977, p. 342). A:I-69. Barri Chopal, Jaipur District; Rajasthan, INDIA; ca. 26°50'N. 75°50'E; observed 1975-1980 by R K. Seth, S. Seth, and A. K. Shukla (1983, p. 38). A:I-75. Bashgal Valley. See Kaotai. Basirhat (= Basinhat) Reserve Forest; West Ben- gal, INDIA; ca. 22°40'N, 88°53'E; observed 1955-1960 by A. K. Mukherjee (Mukherjee & Gupta, 1965, p. 145). B:I-4. Baska Nadi. See Bogra Nadi. Batang vicinity, 2200-3000 m; Sichuan, CHINA; ca. 30°00'N, 99°00'E; reported 1914-1916 by H. Weigold (1924, p. 71). Purchased at Batang market in 1961 by unknown collector; izcas, 1 (skin only). B:C-32. Batu, Pinglang District, Tianlin Xian; Guangxi, 128 FIELDL\NA: ZOOLOGY CHINA; ca. 24°18'N, 106°13'E; collected 31 July 1978 and ca. Oct. 1978 by Neuong Shihua (Quan Guoqiang. pers. comm.. 29 Nov. 1985); FDCG, 2 (1 skin only. 1 skull only). C:C-17(). Bawangling. Changjiang Xian, Hainan Dao, 500- 600 m; Hainan, CHINA; 19°07'N. 109°05'E; collected 1 Jan. 1964 by Liu Zhenhe; sciha, 1. C:C-234. Bawmwang. 3200 ft ( = 975 m); Kachin, MYAN- MAR (= BURMA); 26°39'N. 97°50'E; collect- ed 6 Feb. 1939 by R. Kaulback; bm(nh). 1. B: M-1. Beichuanshan, Taishan Xian, Shangchuan Dao, 0-500 m; Guangdong. CHINA; 21°45'N, 112°50'E; observed Apr.-May 1981 by Liu Zhenhe, sciea (pers. comm.. 25 Nov. 1985). C: C-213. Beijing (= Pekin), monastery outside of; Beijing, CHINA; 39°56'N, 116°24'E; erroneous locality information (Morris & Morris, 1966, p. 18; Hill, 1974. p. 583) for monkeys (presumably either M. midatta or M. thibetana) observed ca. 1325 at monastery in Hangzhou vicinity by Odoric of Pordenone (1928, pp. 232, 234). Not mapped. Benaras. See Varanasi. Benares. See Saktesgarh; Varanasi. Bengal. See West Bengal. "(Bengal)", INDIA or BANGLADESH; 22°- 27°N, 86°-93°E; date and collector unknown; captive. Zoological Society of Philadelphia; ANSP. 1 (skin only). Not mapped. Bengalen; INDIA or BANGLADESH; 22°-27°N. 86°-93°E; collected in 1859 by unknown col- lector; NHMBa. 2. Captive obtained (date un- known) in Sumatra by Prof. Neisser, reportedly imported from "Bengalen"; mzb, I (skull only). Not mapped. Berhampur. Ganjam District, 25 m; Orissa, IN- DIA; 19°19'N, 84°47'E; observed 20 May 1980 by J. Fooden. A:I-107. Bezogaon; Assam, INDIA; ca. 26°56'N, 94°33'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. Bhagalpur District; Bihar, INDIA; ca. 25°15'N, 87°00'E; reported 1810-1811 by F Buchanan (1939, p. 285; posthumous publication). B:I-5. Bhamo, 600 ft (= 180 m); Kachin, MYANMAR (= BURMA); 24°16'N, 97°14'E; collected 15 Feb. 1936 by R F Garthwaite; bm(nh), 1. B:M- 15. Bharatpur; Rajasthan, INDIA; 27°13'N, 77°29'E; reported before 1965 by I. Prakash (letter, 25 Aug. 1964). Reported before 1997 by K. K. Gu- rung and R. Singh (1996, p. 100). A:I-72. Bhamabhari (= Bharnabhavi), Bhutan Duars, 6(X) ft (= 180 m); West Bengal, INDIA; ca. 26°45'N, 89°23'E; collected 21 Feb. 1916 by N. A. Baptista (H. V. O'Donel in Wroughton, 1917, p. 63); BM(NH), 1. B:I-9. Bheri River; Sallyana, NEPAL; ca. 28°30'N, 82°00'E; observed Feb. 1977 by R Byrne (1979, p. 70). A:N-7. Bherjan. See Tinsukia. Bhim Kund Point, Amravati District, 875 m; Ma- harashtra, INDIA; 21°24'N, 77°20'E; observed 31 Jan. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 466). A:I-101. Bhiwani District; Haryana, INDIA; ca. 28°47'N, 76°08'E; observed 1981-1983 by R K. Seth, S. Seth, G. L. Reddy. and R K. Chopra (1992. p. 62). A:I-40. Bhopal, east of; Madhya Pradesh, INDIA; ca. 23°20'N, 77°40'E; reported ca. 1922 by B. C. Ellison (1922, p. 1 100; cf. Fooden, 1989, p. 44). A:I-96. Bhowal area. See Naini Tal. Bhotan Duars. See Bhamabhari; Hasimara. Bhutan Duars. See Bhamabhari; Hasimara. Bialibazar Rubber Plantation; Sylhet, BANGLA- DESH; not precisely located, 24°08'-24°50'N, 9r37'-92°17'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). Not mapped. Bijnor, 1000 ft (= 300 m); Uttar Pradesh, INDIA; 29°22'N, 78°08'E; collected before 1927 by B. B. Osmaston (Napier, 1981, p. 23); bm(nh), I (skull only). A:I-34. Bikaner; Rajasthan, INDIA; 28°0rN, 73°18'E; observed I953-I956 by I. Prakash (1956, p. 3; 1959, p. 39). Reported before 1982 by R. N. Bhargava (1984. pp. 41. 42). A:I-78. Bilauri; Kanchanpur, NEPAL; 28°35'N. 80°22'E; observed June 1964-Dec. 1965 by D. L. Chese- more (1970, p. 164). A:N-3. Biloxue Shan, near Nujiang (= Salween River), Bijiang Xian; 3000 m; Yunnan, CHINA; ca. 26°35'N, 99°05'E; collected 19 Oct. 1978 by Ma Shilai (pers. comm., 1 Sept. 1983); kiz, 1 (skin only). B:C-43. Bishenpur (= Bistenpur). 3000 ft (= 900 m); Manipur, INDIA; 24°38'N, 93°46'E; collected 20 Feb. 1940 by W. J. C. Frost; bm(nh), 2. B: 1-38. Biyun, ca. 30 km north of; Anhui, CHINA; ca. 30°25'N, 118°20'E; observed before 1973 by FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 129 Xiong Chenpei (Wada et al., 1986. p. 82). C:C- 53. Biyun. Jingde Xian, 300-400 m; Anhui, CHINA; ca. 30°10'N, 1 18°20'E; observed 1973-1986 by Xiong Chenpei (Wada et al., 1986, p. 83). C:C- 62. Boai; Hemuu CHINA; 35°10'N, 113°04'E; re- ported before 199.8 (Zhang et al., 1997, p. 58). C:C-11. Boga Juli. See Bogra Nadi. Bogra Nadi (?= Boga Juli), North Kamrup, 2000 ft (= 600m); Assam INDIA; ca. 26°48'N, 9r35'E; collected 5 Jan. 1921 by H. W. Wells (Hinton & Lindsay, 1926, p. 385); hm(nh), 1. B:I-16. Bohea Mts. See Kuatun. Boileauganj. Simla vicinity; Himochal Pradesh, INDIA; ca. 31°06'N, 77°08'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984, p. 477). A: 1-18. Boluo, Yingde Xian; Guangdong, CHINA; 24°25'N, 113°00'E; reported in 1970 by local residents to Liu Zhenhe, sciea (pers. comm., 25 Nov. 1985). C:C-205. Bolovens. See Muang Thateng. Bombay. See Mumbai. Bondor (= Bondar); Narayanganj, BANGLA- DESH; ca. 23°35'N, 90°35'E; reported before 1986 by M. A. R. Khan (1981, p. 13; 1985, p. 31). Observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-27. Borajan. See Tinsukia. Borivli National Park; Maharashtra, INDIA; 19°10'N, 72°55'E; population artificially intro- duced in 1940s (Serrao & Amladi, 1979, p. 29). Observed 1972-1973 by N. Koyama and P. B. Shekar (1981, p. 248). Observed 25 Dec. 1979 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 469). Not mapped. Borme; Gazipur, BANGLADESH; ca. 24°05'N, 90°25'E; reported before 1986 by M. A. R. Khan (1985, p. 31). Observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-15. Bormibazan See Borme. Boska Nadi. See Bogra Nadi. Bo Trach District; Quang Binh, VIETNAM; ca. 17°35'N, 106°33'E; collected 25 Mar. 1976 by unknown collector; iebr, 1 (skull only). C:V-30. Bouzini. Katmandu Valley; Bagmati, NEPAL; ca. 27°4rN, 85°irE; collected 18 June 1922 by N. A. Baptista (Hinton & Fry, 1923, p. 403); BM(NH), 1. A:N-12. Brindaban. See Vrindavan. Bulandshahr District; Uttar Pradesh, INDIA; ca. 28°23'N, 77°52'E; observed 1981-1983 by P K. Seth, S. Seth, G. L. Reddy, and P K. Chopra (1992, p. 62). A:I-46. Buliuhe Water Regulation Forest Reserve; Guangxi, CHINA; ca. 24°57'N, 106°54'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127; letter, Aug. 1996). C:C-173. Bulun. See Sungri. ca. 2 km south of. BURMA. See MYANMAR. Busapuri. See Bussapuram. Bussapuram, Mulug Taluk, Warangal District; An- dhra Pradesh, INDIA; 18°10'N, 80°05'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-120. Cachar District; Assam, INDIA; ca. 25°30'N, 93°00'E; collected Aug.-Sept. 1920 by H. W. Wells (Hinton & Lindsay, 1926, p. 385); bnhs, 1 (skull only). B:I-35. Cala Shan, Zhungshan Township. Luxi Xian, 1500-2000 m; Yunnan, CHINA; ca. 24°30'N, 98°40'E; collected 15 Sept. 1960 by Quan Guo- qiang (pers. comm., 25 Aug. 1983); izcAS, 2. B:C-63. Calcutta, Hastings Road; West Bengal, INDIA; 22°32'N, 88°22'E; observed in 1962 and Dec. 1970-July 1972 by C. H. Southwick, A. Ghosh, and C. D. Louch (1964, p. 444; Southwick et al., 1976, p. 13). B:I-3. Calcutta, Indian Museum compound; West Ben- gal, INDIA; 22°32'N, 88°22'E; collected 25 July 1962 by B. S. Lamba; zsi, 1 (skull only). Observed Dec. 1968-Nov. 1969 by S. S. Saha (1974. p. 211). B:I-3. Calcutta, northern; West Bengal, INDIA; 22°32'N. 88°22'E; observed ca. 1991-1995 by J. Datta (1996, p. 941). B:I-3. Calcutta vicinity; West Bengal, INDIA; ca. 22°32'N, 88°22'E; reportedly "not obtainable [by collectors] near Calcutta," Dec. 1890-Jan. 1891 (Heape. 1894, p. 412). Not mapped. Camp No. 28. See Nong Khai. Camp No. 34. See Ban Mak Nao. Cao Bang, VIETNAM; 22°15'-23°05'N, 105°15'- 106°50'E; reported 1892-1893 by A. Billet (1896, p. 61). Not mapped. Cat Ba, Dao, Vung (= Bay) Ha Long; Hai Phong, VIETNAM; ca. 20°45'N, 107°00'E; purchased in market 28 Jan. 1957 by unknown collector; ZMVNU. 3 (including 2 skins only). Collected in 1958 by unknown collector (Dang, 1983, p. 1282); ZMVNU, 1 (skin only). Collected 11 Aug. 130 FIELDIANA: ZOOLOGY I 1964 by Hien Hao; zmvnu, 3 (skulls only). C: V-14. Cecil, Simla; Himachal Pradesh. INDIA; ca. 31°06'N, 77°09'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984. p. 477). A:I-18. Ceheng; Guizhou, CHINA; 24°58'N. 105°49'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-I59. Cenwanglao Shan Nature Reserve; Giiang.xi. CHINA; ca. 24°19'N. 106°35'E; observed in 1976, 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 126; letter, Aug. 1996). C:C- 171. Chaibasa. See Luia. Chail Sanctuary; Himachal Pradesh, INDIA; ca. 3I°00'N, 77°15'E; reported 1978-1980 by A. J. Gaston. P. J. Garson, and M. L. Hunter, Jr. (1983. p. 300). A:I-18. Chakia Forest Range; Uttar Pradesh. INDIA; ca. 25°03'N. 83°13'E; observed Aug. 1977-July 1978 by R. S. Pirta (1982, p. 401; Pirta & Singh. 1982, p. 15). A:I-88. Chakmani. See Chamkani. Cham Chu, Chiem Hoa District; Tiiyen Qiiang. VIETNAM; 22°12'N. 105°07'E; reported FeK- Apr. 1992 by R. Ratajszczak, Ngoc Can, and Pham Nhat (1992, p. 18). C:V-5. Chamkani, northeast of; Paktia, AFGHANI- STAN; ca. 33°55'N, 69°50'E; reported before 1972 by A. Puget (1971. p. 201). A:A-11. Chamkani. southeast of; Paktia. AFGHANI- STAN; ca. 33°45'N. 70°05'E; reported before 1972 by A. Puget (1971. p. 201). A:A-12. Chandigarh, outskirts of; Punjab. INDIA; ca. 30°44'N. 76°55'E; observed 1985-1986 by R. Boonratana and C. J. Edwin (1986, p. 110). A: 1-19. Chandikhola. Birganj Forest District; Raiitahat, NEPAL; 27°04'N. 85°22'E; observed June 1964-Dec. 1965 by D. L. Chesemore (1970, p. 164). A:N-13. Chandpur Bazar, Old; Chandpur. BANGLA- DESH; 23°13'N, 90°39'E; observed Feb. 1990- June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-28. Changchang Pani; Nagaland. INDIA; 26°36'N, 94°26'E; collected 15-18 Feb. 1930 by C. McCann (1933a, p. 395); amnh, 2. B:I-31. Changde; Hunan. CHINA; 29°02'N, 1 1 l°4rE; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-108. Changhua vicinity, Lin" an Xian; Zhejiang. CHI- NA; ca. 30°10'N, 119°13'E; skin collected lo- cally in 1979 observed Apr. 1980 by Tang Ziey- ing, n'BD (pers. comm., 19 Oct. 1985). C:C-59. Chang Jiang. See Yichang. Changlung. See Nu Jiang. Changnmg [Xian]; Yunnan. CHINA; ca. 24°50'N, 99°36'E; blood sample obtained before 1999 by Ding Bo, Zhang Yaping. and Hou Yidi (1998, p. 172). B:C-64. Changshun; Guizhou. CHINA; 25°59'N, 106°25'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-I46. Changtian. Dongfang Xian, Hainan Dao, 400-500 m; Hainan. CHINA; 19°02'N, 108°54'E; col- lected 23-24 Jan. 1964 by Liu Zhenhe, sciea (pers. comm.. 26 Nov. 1985); sciea, 4. C:C- 235. Chapai. See Nawabganj vicinity. Charilam resthouse, ca. 25 km south of Agartala; Tripura. INDIA; 23°38'N, 9I°18'E; collected 16 Nov. 1969 by V. C. Agrawal (Agrawal & Bhattacharyya, 1977, p. 137); zsi, 1. B:I-40. Charkonda, Mahbubnagar District, 420 m; An- dhra Pradesh, INDIA; 16°42'N. 78°43'E; ob- served 28 and 29 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 468). A:I- 127. Char mugoria; Madaripur. BANGLADESH; ca. 23°10'N. 90°12'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, pp. 76, 79). B:Ba-25. Chaunpur; Uttar Pradesh. INDIA; 27°57'N, 78°08'E; observed Jan. 1990-Mar. 1991 by E. Imam and H. S. A. Yahya (1995. p. 2). A:I-69 Chaur; Dandeldhura, NEPAL; 29°I7'N, 80°2rE; observed June 1964-Dec. 1965 by D. L. Chese- more (1970, p. 164). A:N-1. Chengbihe Water Regulation Forest Reserve; Guangxi. CHINA; ca. 24°02'N, 106°36'E; ob- served in 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127, letter. Aug. 1996). C:C-169. Chengbu; Hunan. CHINA; 26°26'N, 1 10°2rE; re- ported before 1998 (Zhang et al.. 1997. p. 58). C:C-186. Chengjaba, Nanzheng Xian, 900 m; Shaanxi, CHINA; ca. 32°30'N, 107°15'E; observed June 1981 by Yao Jianchu, siz (pers. comm., 10 Oct. 1985). C:C-34. Chengli. 2 mi (= 3 km) west of Ghurkha; Gan- daki. NEPAL; ca. 28°00'N, 84°35'E; collected 7 Dec. 1922 by N. A. Baptista; bm(nh). 1. A: N-9. Chengxian; Gansu. CHINA; 33°42'N. 105°36'E; FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE. MACACA MULATTA 131 reported before 1998 (Zhang et al., 1997, p. 58). C:C-20. Cherekapar, near; Assam, INDIA; ca. 26°59'N, 94°40'E; observed 9 Mar.- 16 Feb. 1988 by A. Choudhury ([1991a], p. 32). B:I-25. Chetia; Assam, INDIA; ca. 27°0rN, 94°44'E; re- ported 9 Mar. 1987-16 Feb. 1988 by A. Choud- hury ([1991a], p. 31). B:I-25. Chhatari-do-Raha; Uttar Pradesh, INDIA; ca. 28°07'N, 78°09'E; observed 1959-1984 by R. L. Johnson, M. F. Siddiqi, and C. H. Southwick (Southwick et al., 1976, p. 13; 1986, p. 433; Southwick & M. F Siddiqi, 1977, p. 342; Sid- diqi & Southwick 1980, p. 54; 1988, p. 121; Johnson & Southwick, 1984, p. 201; South- wick, 1985, p. 191; Johnson, 1986, p. 193). Ob- served Jan. 1990-Mar. 1991 by E. Imam and H. S. A. Yahya (1995, p. 2). A:I-69. Chhota Bangahal, upper Beas River catchment area; Himachal Pradesh, INDIA; not precisely located, 31°30'-32°30'N, 77°00'-77°30'E; re- ported 1978-1980 by A. J. Gaston, R J. Garson, and M. L. Hunter, Jr. (1983, p. 300). Not mapped. Chiang Dao; Chiang Mai, THAILAND; 19°22'N, 98°58'E; collected 14 June 1937 by C. R. Car- penter (1940, p. 20; Coolidge, 1940, p. 129); Mcz, 1. B:T-1. Chiang Mai, near; Chiang Mai, THAILAND; ca. 18°47'N, 98°59'E; collected in 1931 by H. G. Deignan (Chasen, 1935, p. 38); zrc, 1. B:T-2. Chiem Hoa; Tuyen Qiiang, VIETNAM; 22°09'N, 105°17'E; collected in 1962 by unknown col- lector; ZMVNU, 1. C:V-5. Chieng Dao. See Chiang Dao. Chiengmai. See Chiang Mai, near. Chiengmai, below. See Kaeng Mae Hat. Chigha Sarai, north of; Konarha, AFGHANI- STAN; ca. 34°52'N, 71°10'E; collector and date unknown, captive purchased 4 mi (= 6.5 km) north of Kandahar by J. Hassinger (1968, p. 72), 2 Nov. 1965; fmnh, 1. A:A-8. Chihli. See Xinglong Xian, southern. CHINA; 18°-41°N, 85°-120°E; captive shipped from Shanghai before 24 Jan. 1880 by unknown collector; bm(nh), 1 (skin only). Captive pur- chased in Lhasa (extralimital) 1938-1939 by E. Schafer; zsbs, 1. Collected 1950-1978 by mu- seum collectors; smnh, 23 (13 skins only, 10 skulls only). Collected in 1960 by unknown collectors; izcas, 4 (skins only). Date and col- lector unknown; bmnh, 2 (including 1 skull only); sciea, 1 (skin only). Not mapped. 7CHINA; 18°-41°N, 85°-120°E; no data, possibly collected May-July 1890 at Ngamda or Roue- toundo (see below) by G. Bonvalot and H. d'Orleans (Bonvalot, 1891, vol. 2, p. 210; 1892, p. 505); MNHN, 1 (skin only). Not mapped. CHINA, northern; 30°-45°N, 100°-125°E; ob- tained date unknown by Mr. Gerrard; zmb, 1 . Not mapped. CHINA, South; 20°-30°N, 100°-120°E; collected 1923-1924 by F R. Wulsin (letter, 9 Jan. 1925; USNM archives, no. 85377); mcz, 2. Not mapped. Chindwin River; Sagaing, MYANMAR (= BUR- MA); 21°35'-27°00'N, 94°20'-97°10'E; collect- ed ca. 1 9 14 by G. C. Shortridge; bnhs, 1 (skull only). Not mapped. Chin Hills. See Kindat, 20 mi (= 32 km) north- west of. Chitrakut, Jagvedi and Bara Math Temples; Uttar Pradesh, INDIA; 25°12'N, 81°00'E; observed 1959-1960 and 1964-1965 by C. H. South- wick, M. A. Beg, and M. R. Siddiqi (1961a, p. 543; Southwick and M. R. Siddiqi, 1966, p. 312). A:I-85. Chitral. See Nurestan, eastern. Chitral, lower. See Kaotai; Kunar River; Mirk- hani. Chittagong, eastern; Chittagong, BANGLA- DESH; ca. 22°20'N, 92°10'E; reported early in 1980 by S. R Gittins and A. W. Akonda (1982, p. 278). B:Ba-35. Chittagong, northern; Chittagong, BANGLA- DESH; ca. 22°50'N, 9r40'E; observed early in 1980 by S. R Gittins and A. W. Akonda (1982, p. 278). Observed before 1982 by M. A. R. Khan (1981, p. 13). B:Ba-33. Chittagong, southern; Cox's Bazar, BANGLA- DESH; ca. 21°50'N, 92°00'E; observed early in 1980 by S. R Gittins and A. W. Akonda (1982, p. 278). B:Ba-38. Chittagong Hill Tracts, BANGLADESH; 2n5'- 22°20'N, 92°25'-92°45'E; reported July-Nov. 1976 by K. M. Green (1978, p. 146). Not mapped. Chittagong Hill Tracts; Chin, MYANMAR ( = BURMA); ca. 2r40'N, 92°40'E; collected be- fore 1927 by B. B. Osmaston (Napier, 1981, p. 21); BM(NH), 1 (skull only). B:M-25. Chittagong Hill Tracts, eastern, BANGLADESH; ca. 22°15'N, 92°30'E; reported early in 1980 by S. R Gittins and A. W. Akonda (1982, p. 278). B:Ba-36. Chittagong Hill Tracts, northern, BANGLA- DESH; ca. 23°20'N, 92°10'E; Observed early in 1980 by S. R Gittins and A. W. Akonda (1982, 132 FIELDIANA: ZOOLOGY p. 278). Observed before 1982 by M. A. R. Khan (1981. p. 13). B:Ba-34. Chitwan National Park. See Dudurhani; Siniri. Narayani River. Cho Don District; Bac Thai, VIETNAM; ca. 22°10'N. 105°36'E; collected Oct. 1989 by un- known collector; fcxm. 1 (skull only). C:V-6. Chokoria Sunderbans; Cox's Bazar. BANGLA- DESH; ca. 21°45'N, 92°00'E; reported early in 1980 by S. R Gittins and A. W. Akonda (1982. p. 278). B:Ba-38. Chong'an Xian; Fiijian. CHINA; ca. 27°5rN. 117°48'E; collected 4 May and 18 Aug. 1926 by C. H. Pope (1929, p. 345; 1932c, pp. 491, 495; 1935, pp. 493. 499; 1940, p. 72); amnh, 2. Reported Aug. 1981 by Zheng Xueqing (1984, p. 145). C:C-76. Chongyi; Jiangxi, CHINA; 25°42'N, 114°19'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-101. Chongzuo Rare Animal Reserve; GuangxL CHI- NA; ca. 22°35'N, 107°28'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter, Aug. 1996). C:C-225. Chowkichora, Akhnoor Subdistrict; Jammu & Kashmir. INDIA; ca. 32°54'N, 1A°AA"E; ob- served before 1983 by Y. R. Malhotra and D. N. Sahi (1982, p. 27). A:I-5. Chuadanga, Chuadanga, BANGLADESH; 23°38'N, 88°51'E; reported before 1982 by M. A. R. Khan (1981, p. 13). B:Ba-18. Chuan Dong, Tian'e Xian; Guangxi. CHINA; 25°09'N, 107°05'E; captured 9 Oct. 1992 by Qing Kailon (pers. comm., 16 Oct. 1992); cap- tive observed at Chuan Dong by J. Fooden. 16 Oct. 1992. C:C-173. Chuandonghe Water Regulation Forest Reserve; Guangxi. CHINA; ca. 25°10'N. 107°03'E; ob- served 1976. 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C-I73. Chunabati. See Sukna-Kurseong. Chunar. See Saktesgarh. Chunati Wildlife Sanctuary; Chittagong. BANG- LADESH; ca. 21°58'N, 92°04'E; observed Feb. 1990-June 1993 by M. M. Feeroz. M. A. Islam, and M. Kabir (1995. p. 76). B:Ba-38. Chungan Hsien. See Chong'an Xian. Chungtia, 4000 ft (= 1200 m); Nagakmd. INDIA; 26°24'N, 94°28'E; collected ca. Sept. 1919 by J. R Mills (1923, p. 222); museum unknown, 2 (not seen). B:I-32. Chunxiu Water Regulation Forest Reserve; Guangxi, CHINA; ca. 22°27'N, 106°34'E; ob- served 1976, 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 129; letter. Aug. 1996). C:C-224. Chuxiong; Yunnan. CHINA; 25°02'N. 101°33'E; tissue sample obtained ca. 1991 by Zhang Yap- ing and Shi Liming (1993b. p. 589). B:C-52. Cih; Hunan, CHINA; 29°24'N. 111°04'E; report- ed before 1998 (Zhang et al.. 1997. p. 58). C: C-109. Cioupu Shan. Hechi town. Hechi Xian; Guangxi, CHINA; 24°42'N. 108°02'E; collected June 1992 by Tan Qin (pers. comm.. 13 Oct. 1992); not preserved. C:C-178. Comilla. BANGLADESH; ca. 23°27'N. 9n2'E; reported before 1982 by M. A. R. Khan (1981, p. 13). B:Ba-29. Corbett National Park; Uttar Pradesh. INDIA; 29°25'-29°35'N, 78°40'-79°05'E; observed Apr. 1954 by E. P Gee (1975. p. 101). Reported before 1997 by K. K. Gurung and R. Singh (1996. p. 86). Not mapped (see A:I-33). Cotgai. See Kotgay. Cox's Bazar. BANGLADESH; 20°48'-2I°10'N, 92°05'-92°20'E; observed early in 1980 by S. P Gittins and A. W. Akonda (1982. p. 278). Not mapped (see B:Ba-38). Cox's Bazar; Cox's Bazar. BANGLADESH; 21°26'N. 91°59'E; observed before 1982 by M. A. R. Khan (1981. p. 13). B:Ba-38. Cox's Bazar, Forest Division (South); Cox's Ba- zar. BANGLADESH; 20°45'-21°30'N, 92°00'- 92°20'E; reported May 1982-Dec. 1983 by S. M. A. Rashid, A. Khan, and M. A. R. Khan (1990, p. 64). Not mapped (see B:Ba-38). Cox's Bazar, northern, BANGLADESH; ca. 21°35'N, 92°10'E; observed early in 1980 by S. P Gittins and A. W. Akonda (1982, p. 278). B: Ba-37. Cue Phuong; Ninh Binh, VIETNAM; 20°19'N, 105°38'E; collected 13 Apr. and 26 Aug. 1963 by unknown collector; zmvnu, 2 (skulls only, species identification tentative). C:V-23. Dacca. See Dhaka. Dachepalle. Piduguralla Taluk. Guntur District; Andhra Pradesh, INDIA; 16°36'N, 79°44'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, pp. 17, 19). A:I-130. Dafla Hills; Arunachal Pradesh. INDIA; ca. 27°20'N, 93°30'E; reported Dec. 1972-Feb. 1973 by R. L. Tilson (1983, p. 399). B:I-24. Dahao Dao; Xianggang (= Hong Kong). CHINA; ca. 22°15'N, 114°00'E; reported 1908-1921 by R. Mell (1922, p. 10). C:C-210. Dahe. Xixiang Xian, 800 m; Shaanxi. CHINA; FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 133 32°37'N, 107°27'E; observed July 1974 by Yao Jianchu, siz (pers. comm.. 10 Oct. 1985). C:C- 35. Dahongbao Nature Reserve; Guangxi, CHINA; 24°52'N. 105°18'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127; letter, Aug. 1996). C:C-160. Dainkog (= Dengke); Sichuan. CHINA; 32°32'N, 97°55'E; reported before 1998 (Zhang et al., 1997, p. 58). B:C-19. Daiyun Shan; Fiijian, CHINA; ca. 25°50'N, 1 18° 15 'E; reported Nov. 1983 by Zhen Xueqing (1984. p. 145). C:C-86. Dak Sut, 700 m; Kon Turn, VIETNAM; 14°56'N, 107°44'E; collected 13-16 Jan. 1961 by B. Feinstein (Van Peenen et al., 1969, p. 286; Fooden, 1997, p. 227); usnm, 3. C:V-37. Dalimkhola. See Gorubathan Forest. Dalingxia. Gangcun. She Xian, 400-700 m; An- Imi. CHINA; ca. 30°00'N, 118°10'E; observed 1973-1986 by Xiong Chenpei, K. Wada, and Wang Qishan (Wada et al., 1986, pp. 83, 88). C:C-62. Dalu. See Taro. Darning Shan Nature Reserve; Guang.xi, CHINA; ca. 23°27'N, 108°22'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 126; letter, Aug. 1996). C:C-221. Damoh. See Kakara. Danba; Sichuan. CHINA; 30°57'N, 10r55'E; col- lected before 1992 by unknown collector (Jiang Xuelong et al., 1991, p. 243); museum un- known, 4 (not seen). B:C-26. Dangan Dao (- North Lena Island); Guangdong, CHINA; ca. 22°02'N, 114°18'E; observed 5 May 1854 by W. Stimpson (unpublished jour- nal, p. 131; R. Vasile, letters, 2 and 13 May 1987). Collected in 1866 by Commander St. John (in Swinhoe, [1867], p. 556); bm(nh), 1 (holotype of Inuus sancti-johannis Swinhoe, [1867]). Reported ca. 1951 by G. A. C. Herk- lots (1951, p. 83). C:C-211. Dangan Dao, Zhuhai Xian, 150 m; Guangdong, CHINA; 22°02'N, 114°18'E; collected 2 and 25 Mar. 1981 by Liu Zhenhe and Xu Longhuei (Liu Zhenhe, sciea, pers. comm., 26 Nov. 1985); SCIEA, 2. C:C-211. Dangen Island. See Dangan Dao. Dangori Nadi. near; Assam, INDIA; ca. 27°36'N, 95°16'E; observed 6-8 Feb. 1974 by G.Pilleri (1975, p. 20; Pilleri & Pilleri, 1982, p. 158; let- ter, 15 Dec. 1978); misidentified as Macaca as- samensis. B:I-26. Dangs District; Gujarat, INDIA; ca. 20°35'- 2rOO'N, 73°30'-73°55'E; collected 1922-1923 (date discrepancy between original tags and museum tags) by A. C. Miller; bm(nh), 3. Not mapped (see A:I-99). Dan Sai District; U)ei, THAILAND; ca. 17°17'N, 101°09'E; collected 31 Mar. 1954 by R. E. El- bel; LSNM. 3. B:T-10. Danzhou, Hainan Dao; Hainan, CHINA; 19°43'N, 109°17'E; reported before 1998 (Zhang et al., 1997. p. 58). C:C-228. Daoshiwu, Lin' an Xian; Zhejiang, CHINA; ca. 30°18'N, 118°57'E; observed Feb. 1985 by Wu Fuhai. Hangzhou Zoo (pers. comm., 25 Oct. 1985). C:C-55. Daping Shan Nature Reserve; Guangxi, CHINA; ca. 23°33'N, 109°58'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127; letter. Aug. 1996). C:C-196. Dareh Nur. See Khyber Pass vicinity. Darjeeling. See Narbong; Sukna. Darsi, 100 m; Andhra Pradesh, INDIA; 15°46'N, 79°41'E; reported Feb. 1977-JuIy, 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). Observed 6 May 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 469). A:I-146. Dashashamedh Ghat. See Varanasi. Dashuping, 400 m; Shaanxi, CHINA; 32°26'N, 107°27'E; observed July 1974 by Yao Jianchu, SIZ (pers. comm.. 10 Oct. 1985). C:C-32 Datang. Lunshuihe (river). Tengchong Xian; Yun- nan, CHINA; ca. 25°30'N, 98°45'E; collected Dec. 1976 by Ma Shilai (pers. comm., 1 Sept. 1983); Kiz, 6. B:C-57. Dauthara, Aligarh vicinity; Uttar Pradesh, IN- DIA; not precisely located, 27°40'-28°10'N, 77°50'-78°20'E; observed 1959-1975 by C. H. Southwick and M. E Siddiqi (1977, p. 342). Not mapped. Dawanglin Water Regulation Forest Reserve; Guangxi, CHINA; ca. 23°32'N. 106°20'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 127; letter, Aug. 1996). C:C-168. Daxin Rare Animal Reserve; Guangxi. CHINA; ca. 22°45'N, 107°08'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter. Aug. 1996). C:C-223. Dayao Shan Nature Reserve; Guangxi, CHINA; ca. 24°02'N, 110°08'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 126; letter, Aug. 1996). C:C-195. Deep Water Bay. See Sam Shui Wan Valley. Defu Water Regulation Forest Reserve; Guangxi, CHINA; 23°17'N, 105°47'E; observed 1976, 134 FIELDIANA: ZOOLOGY 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C-164. Dege vicinity; Sichuan, CHINA; 31°49'N, 98°40'E; purchased in Dege by unknown col- lector. 15 July 1961; izcas. 1 (skin only). B:C- 22. Dehra Dun; Una?- Pradesh. INDIA; 30°19'N, 78°02'E; observed June 1965-May 1966 by D. G. Lindburg (1971. p. 5). A:I-27. Dehra Dun vicinity; Uttar Pradesh. INDIA; ca. 30°19'N, 78°02'E; observed Sept. 1959-Feb. 1960 by C. H. Southwick, M. A. Beg. and M. R. Siddiqi (1961a, p. 540). Observed 1975- 1980 by P. K. Seth. S. Seth. and A. Shukla (1983. p. 38; Seth and Seth. 1983. p. 63). Ob- served 1981-1983 by P. K. Seth. S. Seth, G. L. Reddy, and R K. Chopra (1992, p. 62). Trapped for psychological study before 1993 by K. Wa- heeda (1992, p. 111). A:I-27. Dehra Dun vicinity, 600 m; Uttar Pradesh, IN- DIA; ca. 30°19'N, 78°02'E; observed Aug. 1955 by M. L. Roonwal (1956, p. 171). A:I-27. Dela, ca. 8 mi ( = 13 km) west of Ramnagar, Ku- maun region, 1500 ft (= 460 m); Uttar Pra- desh, INDIA; 29°26'N, 79°00'E; collected 7 Jan. 1914 by C. A. Crump (in Wroughton, 1914, p. 284; Napier, 1981, p. 24); bm(nh), 1 (skin only). A:I-33. Delhi; Delhi, INDIA; 28°40'N, 77°13'E; observed Sept. 1959-Feb. 1960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961a, p. 543). Ob- served 1964-1965 by R. P Mukherjee and G. D. Mukherjee (1972, p. 67). A:I-41. Delhi-Agra, highway between; Haryana, INDIA; ca. 28°00'N. 77°20'E; observed Sept. 1961 by 1. Krumbiegel (1965. p. 32). Observed 1964- 1965 by R. P. Mukherjee and G. D. Mukherjee (1972. p. 67). A:I-43. Delhi-Aligarh. highway between; Uttar Pradesh, INDIA; ca. 28°20'N, 77°50'E; observed 2-5 Jan. 1965 by C. H. Southwick, D. Lindburg, M. R. Siddiqi. R. P. Mukherjee, and B. Singh (Southwick & M. R. Siddiqi, 1966, p. 306). A: 1-46. Delhi-Hathras, highway between; Uttar Pradesh, INDIA; ca. 28°10'N, 77°40'E; observed Sept. 1959-June 1960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961b, p. 702). A:I- 45. Delhi-Kanpur, highway between; Uttar Pradesh, INDIA; ca. 27°30'N, 78°45'E; observed 1964- 1965 by R. P. Mukherjee and G. D. Mukherjee (1972, p. 67). A:l-68. Delhi-Mathura, highway between; Delhi, INDIA; ca. 28°40'N, 77°13'E; observed Dec. 1970-July 1972 by C. H. Southwick, M. E Siddiqi, M. Y. Farooqui, and B. C. Pal (1976, p. 13). A:I-4I. Delhi vicinity; Delhi, INDIA; ca. 28°40'N, 77°13'E; autopsied ca. 1966 by K. K. Chawla, C. D. S. Murthy, R. N. Chakravarti. and R N. Chhuttani (1967, p. 85). Observed 1981-1983 by R K. Seth, S. Seth, G. L. Reddy, and P K. Chopra (1992, p. 62). A:I-41. Dengke. See Dainkog. Dening. Mishmi Hills. 2240-2250 ft (= 685 m); Anmachal Pradesh, INDIA; 28°0rN, 96°14'E; collected 29 Mar. and 6 Apr. 1921 by H. W. Wells (Hinton & Lindsay, 1926, p. 385); BM(NH), 3 (including 1 skin only). B:I-28. Deogarh, Sambalpur District; Orissa, INDIA; 2r32'N, 84°44'E; collected 30 Dec. 1972 by A. K. Mandal; zsi, 1 (skin only). A:I-104. Deqen; Yunnan, CHINA; 28°30'N, 98°52'E; re- ported before 1998 (Zhang et al., 1997, p. 58). B:C-36. Dhaka; Dhaka, BANGLADESH; 23°43'N, 90°25'E; observed 1975-1977 by J. R. Oppen- heimer, A. W. Akonda, and K. Z. Husain (1983, p. 194). Observed July-Nov. 1976 by K. M. Green (1978, p. 154). Reported before 1982 by M. A. R. Khan (1981, p. 13; 1985, p. 31). Ob- served Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995. p. 76). B:Ba- 27. Dhalar. See Narkanda. ca. 1 km north of. Dhaleswari River. See Nagorhgena. Dhamrai (= Dhamrei); Dhaka, BANGLADESH; 23°55'N. 90°13'E; observed Jan. 1976 by J. R. Oppenheimer. A. K. Akonda, and K. Z. Husain (1983, p. 194). Observed July-Nov. 1976 by K. M. Green (1978, p. 154). Reported before 1982 by M. A. R. Khan (1981. p. 13; 1985. p. 31). Observed Feb. 1990-June 1993 by M. M. Fee- roz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-16. Dhangarhi; Kailali, NEPAL; 28°34'N, 80°36'E; observed June 1964-Dec. 1965 by D. L. Chese- more (1970, p. 164). A:N-4. Dharmajigudem (= Dharmajidudem), Chintala- pudi Taluk, West Godavari District; Andhra Pradesh, INDIA; 16°53'N, 81°00'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). A:I-I39. Dharmsala, 4500 ft (= 1370 m); Himachal Pra- desh, INDIA; 32°13'N, 76°19'E; collected 5 Feb. 1922 by H. W. Wells (Lindsay. 1926. p. 599); BM(NH). 1. A:I-12. Dhaulkot Forest. Dehra Dun region; Uttar Pra- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE. MACACA MULATTA 135 desh, INDIA; ca. 30°25'N. 77°55'E; observed Oct.-Dec. 1977 by S. C. Makwana (1979a, p. 242). A:I-27. Dhela. See Dela. Dhikala. See Ramganga River. Dhulkot. See Dhaulkot Forest. Dhuniopathar; Assam, INDIA; ca. 27°07'N, 95°10'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991aJ. p. 31). B:I-30. Dibru-Saikhowa Wildlife Sanctuary; Assam. IN- DIA; 27°35'-27°47'N. 95°10'-95°40'E; ob- served 1 992- 1 996 by A. Choudhury (1998, pp. 194, 197). Not mapped (see B:I-26). Dichialgaon; Assam, INDIA; ca. 26°58'N, 94°3rE; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991aJ, p. 31). B:I-25. Diggi Road. See Aligarh. Dihajan habi; Assam, INDIA; 27°08'N, 94°46'E; observed 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a]. p. 32). 8:1-25. Dillighat; Assam, INDIA; ca. 27°10'N, 95°17'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-30. Dimapur. See Imphal, ca. 4 mi (= 6.5 km) north of. Dinajpur District; Dinajpur, BANGLADESH; ca. 25°50'N, 88°35'E; reported before 1982 by M. A. R. Khan (1981, p. 13; 1985, p. 31). B:Ba-l. Dinh Ca, Thai Nguyen region, 300 m; Bac Thai, VIETNAM; 21°45'N, 106°03'E; collected 20 Dec. 1956 by unknown collector (Dao, 1961, p. 302; 1985, p. 64); museum unknown, 1 (skull only; cranial measurements cited by Dao). C:V- 11. Dir District. See Dokdusra; Gwaldri Valley; Lan- drai Valley. Diroi (Rangoli) Reserve Forest; Assam, INDIA; ca. 27°08'N, 95°01'E; observed 9 Mar. 1987- 16 Feb. 1988 by A. Choudhury ([1991a], p. 32). B:I-30. Ditin, Jingxi Xian; Guangxi, CHINA; ca. 23°10'N, 106°30'E; collected Nov. 1982 by Wei Zhanyi; fdcg, 1 (Skull only). C:C-167. Dizhou Water Regulation Forest Reserve; Guang- xi, CHINA; ca. 23°02'N, 106°13'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C- 166. Dokdusra, northern Dir District; North-West Frontier, PAKISTAN; 35°32'N, 72°I3'E; re- ported before 1978 by T. J. Roberts (1977, p. 86). A:P-2. Dommeru, West Godavari District, 40 m; Andhra Pradesh, INDIA; 17°02'N, 8r4rE; observed 17 May 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A:I-138. Dongfang, Hainan Dao; Hainan, CHINA; ca. 19°04'N, 108°5rE; collected 7 Feb. 1957 by Tang Ziying; fubd, 2 (1 skin only, 1 skull only). Collected 9 May 1957 by Wang Sung; izcas, 1. C:C-235. Dong He. See Wa Shan. Dongmen, Xianan Subcounty, Huanjiang Xian, 460 m; Guangxi, CHINA; ca. 25°09'N, 107°53'E; collected 21 Oct. 1992 by Tan Nenrui (pers. comm. 5 Nov. 1992); izcas, 1 (skin only). C:C-177. Dongshan, 5 km south of Xianan, Huanjiang Xian, 440 m; Guangxi, CHINA; ca. 24°52'N, 107°57'E; captured ca. 15 Oct. 1992 by Gung Lao (pers. comm., 5 Nov. 1992); captive ob- served 5 Nov. 1992 at Xianan. C:C-177. Dowoka, above 9500 ft (>2900 m); Xizang ( = Tibet), CHINA; ca. 29°22'N, 94°18'E; reported July-Aug. 1913 by F M. Bailey (1914, map; 1915, p. 74). B:C-9. Doza. See Narkanda, ca. 1 km north of. Dudhwa National Park; Uttar Pradesh, INDIA; ca. 28°25'N, 80°45'E; reported before 1997 by K. K. Gurung and R. Singh (1996, p. 90). A:I- supplementary. Dudurhani, Narayani River, Rapti Valley; Chita- wan, NEPAL; 27°34'N, 84°14'E; observed June 1964-Dec. 1965 by D. L. Chesemore (1970, p. 164). Reported at Chitwan National Park before 1997 by K. K. Gurung and R. Singh (1996, p. 84). A:N-8. Dumel, Udhampur Subdistrict; Jammu & Kashmir INDIA; ca. 32°56'N, 75°08'E; observed before 1983 by Y. R. Malhotra and D. N. Sahi (1982, p. 27). A:I-10. Dumpallagudem, Mulug Taluk, Warangal District; Andhra Pradesh, INDIA; 18°09'N, 80°09'E; re- ported Feb. 1977-JuIy 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-120. Dunga Gali vicinity, Hazara District; North-West Frontier, PAKISTAN; ca. 34°03'N, 73°22'E; reported summer 1968 by T J. Roberts (1977, p. 87). Blood samples collected 1978-1979 by D. J. Melnick, C. J. Jolly, and K. K. Kidd (1984, p. 342). Observed 15 Mar. 1980-15 Dec. 1981 by A. Rab, S. S. Sahibzada, and M. Afzal (1991, p. 219). A:P-12. Dunga Gali vicinity, Hazara District, 2000-2800 m; North-West Frontier, PAKISTAN; 34°03'N, 73°22'E; collected 12 Sept. 1963 at 8100 ft ( = 2470 m) by R. L. Amouraux; usnm, 1. Ob- served 1978-1981 at 2000-2800 m by S. J. 136 FffiLDLANA: ZOOLOGY Goldstein and A. F. Richard (1989. p. 532). Ob- served 1979-1980 at 8200 ft (= 2500 m) by S. S. Sahibzada, Q. A. Iqbal, and A. Rab (1985. p. 198). A:P-12. Dunwein. Kishtwar District. 7600-7800 ft ( = 2320-2380 m); Jammu & Kashmir. INDIA; 33°20'N. 75°49'E; collected 21 Nov. 1897 by P. H. G. Powell-Cotton (Napier, 1981, p. 25); p- CM, 2 (including 1 mounted skin with skull in- side). A:I-11. Dupleix Mountains, south of; Xizang. CHINA; ca. 33°38'N, 89°43'E; improbable record (Bonva- lot, 1891, vol. 1, p. 210; 1892, p. 218; See "Geographic Distribution and Current Popula- tion Estimates" above). Not mapped. Durga Temple. See Varanasi. Dushan; Guizhou CHINA; 25°50'N, 107°32'E; tissue samples obtained ca. 1991 by Zhang Tap- ing and Shi Liming (1993b, p. 589). C:C-123. Dzo La, southeast of; Xizang (= Tibet), CHINA; ca. 29°I3'N, 97°07'E; observed 12-24 July 1935 by R. Kaulback (1938, p. 91). B:C-33. Eagle's Nest Trail, Kowloon Reservoir Area; Xianggang (= Hong Kong), CHINA; ca. 22°21'N, 114°09'E; population probably artifi- cially introduced (Herklots, 1951, p. 83). Ob- served July-Aug. 1980 and July-Aug. 1981 by C. H. Southwick and K. L. Southwick (1983, p. 19). Observed 1984-1988 by E D. Burton and L. Chan (1996, p. 396). Observed 10 Jan.- 3 Feb. 1987 by C. H. Southwick and D. Manry (1987, p. 48). C:C=210. East BURMA. See MYANMAR (= BURMA), eastern. Eastern Mausleum. See Xinglong Xian, south- em. Eastern Mausoleum. See Xinglong Xian, south- em. Eastem Tombs. See Xinglong Xian, southem. East Sichuan. See Sichuan, eastem. Emei Shan (= Mount Omei); Sichuan, CHINA; ca. 29°32'N, 103°21'E; collected before 1930 by unknown collector (Howell, 1929, p. 35); USNM, 1 (specimen not located). Reported be- fore 1942 by A. de C. Sowerby (1941, p. 261). C:C-139. Enshi; Hubei, CHINA; 30°18'N, 109°29'E; tissue sample obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). C:C-39. Erzhou Dao; Guangdong, CHINA; ca. 22°00'N, 114°11'E; reported in 1981 by Liu Zhenhe and Yuan Xicai (Zhang et al., 1991, p. 177; 1997, p. 58). Population possibly artificially intro- duced > 1 00 years ago (Zhang Yongzu, letter. 3 July 1996). C:C-211. Faizabad vicinity; Uttar Pradesh, INDIA; ca. 26°47'N, 82°08'E; blood samples collected 16- 27 Apr. 1964 by K. V. Shah and C. H. South- wick (1965, p. 489). A:l-60. Faizabad-Ajodhya, highway between; Uttar Pra- desh. INDIA; ca. 26°45'N, 82°10'E; observed Sept. 1959-June 1960 by C. H. Southwick, M. A. Beg. and M. R. Siddiqi (1961b. p. 702). A: 1-60. Fameng, Xingyi Xian, 1580 m; Guizhou. CHINA; ca. 24°45'N, 104°45'E; collected 11 July 1963 by Wang Yingxiang (pers. comm.. 1 Sept. 1983); Kiz, 1 (skull only). C:C-155. Fangdao Nature Reserve, Jian'ou Xian; Fujian, CHINA; 27°0rN, 118°08'E; observed July 1985 by Tang Ziying, fubd (pers. comm., 19 Oct. 1985). C:C-72. Fanjingshan; Guizhou, CHINA; 27°57'N, 108°50'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-116. Faridpur, BANGLADESH; ca. 23°15'N, 90°00'E; reported before 1982 by M. A. R. Khan (1981, p. 13). B:Ba-26. Fechugang; Sylhet, BANGLADESH; 24°42'N, 91°57'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-12. Fengxingshan, Jing'an Xian; Jiangxi, CHINA ca. 8°52'N, 115°22'E; trapped and released May 1984 (Tan, 1985, p. 73). C:C-77. Fokien Occid. See Kuatun. Forest Research Institute. See Dehra Dun. Fugong; Yunnan, CHINA; 26°58'N, 98°54'E; re- ported before 1998 (Zhang et al., 1997, p. 58). B:C-42. Fuhai. See Menghai. Fujian, CHINA; 24°-28°N, 116°-120°E; collected before 1986 by unknown collector; .sciha, 1 (skin only). Tissue samples obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). Not mapped. Funing; Yunnan, CHINA; 23°37'N, 105°36'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-163. Fuqing vicinity; Fujian, CHINA; ca. 25°43'N, 119°22'E; purchased Aug. 1963 by unknown collector; IZCAS, 2. C:C-85. Fusui Rare Animal Reserve; Guangxi, CHINA; ca. 22°30'N, 107°32'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter, Aug. 1996). C:C-225. Fuxi, She Xian, 400-700 m; Anhui, CHINA; ca. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 137 30°05'N. 118°15'E; observed 1973-1986 by Xiong Chenpei, K. Wada, and Wang Qishan (Wada et al., 1986, pp. 83, 88, 91). C:C-62. Fuyuan; Yimnatu CHINA; 25°40'N, 104°14'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-147. Galta, ca. 10 km from Jaipur; Rajasthau, INDIA; ca. 26°53'N, 75°52'E; observed Mar. 1956-Oct. I960 by I. Prakash (1958, p. 154; 1962, p. 83). Observed 1975-1980 by P. K. Seth and S. Seth (1983. p. 63). Observed Nov. 1979-Oct. 1980 by R. Singh (1986, p. 607; 1989, p. 139; 1992, p. 192). Observed ca. 1982-1988 by L. D. Wolfe and R. Mathur (1988, p. 535; Mathur, 1982, p. 12; Wolfe, 1992, p. 45). Observed be- fore 1993 by N. K. Chandel (1992, p. 121). A: 1-75. Gam, Song, Left bank; Tuyen Qiiang, VIETNAM; ca. 22°25'N, 105°22'E; reported Feb.-Apr. 1992 by R. Ratajszczak, Ngoc Can, and Pham Nhat (1992, p. 17). C:V-6. Gange. See Ganges River. Ganges River, forests along banks ("forets des bords du Gange"); Bihar. Uttar Pradesh, or West Bengal INDIA; 23°-31°N, 78°-88°E; re- ported before 1820 by E Cuvier (1819, p. 2). Not mapped. Gangupahad (= Ganukapahad), Jangaon Taluk, Warangal District; Andhra Pradesh, INDIA; 17°47'N, 79°08'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-122. Gao Lo Shan; Guangxi, CHINA; ca. 24°10'N, 106°47'E; captured Nov.-Dec. 1990 by Liu Jinhwei (pers. comm., 15 Oct. 1992); captive observed at Tian'e, 15 Oct. 1992. C:C-172. Garampani. See Hot Springs. Garidhwa. See Simulbari-Pankhabari. Garo Hills; Meghalaya INDIA ca. 25°30'N, 90°30'E; reported Dec. 1972-Feb. 1973 by R. L. Tilson (1983, p. 399). B:I-13. Garo Hills, foot; Sherpur, BANGLADESH; ca. 25°00'N, 90°00'E; observed before 1982 by M. A. R. Khan (1981, p. 13). B:Ba-6. Garubathan. See Gorubathan Forest. Gaushalla, 1330 m; Katmandu Valley, NEPAL; 27°42'N, 85°21'E; observed 1977-1982 by B. M. Marriott (1988, p. 126). Observed summer 1983 by R. L. Johnson and C. H. Southwick (1984, p. 201). A:N-12. Gegong, Dongzhi Xian, 400-600 m; Anhid, CHI- NA; 3()°05'N, 117°irE; observed 1973-1986 by Xiong Chenpei (Wada et al., 1986, p. 83). C:C-49. Gejiu; Yunnan, CHINA; 23°23'N, 103°09'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-149. Gengma; Yunnan, CHINA; ca, 23°3rN, 99°24'E; collected 18-20 May 1964 by Quan Guoqiang (pers. comm., 25 Aug. 1983; Yen Wenchen, 1973, p. 356); izcas, 3. B:C-67. Getou, Leishan Xian, 1200 m; Guizhou, CHINA; ca. 26°20'N, 108°10'E; collected 4 Sept. 1963 by unknown collector; Kiz, 3. C:C-121. Gharmur (= Gharmura), ca. 1 km south of ; As- sam, INDIA; ca. 24°17'N, 92°3rE; observed 21-25 Mar. 1986 and 1987-1988 by A. Choud- hury (1983, p. 14; 1989, p. 491; [1991b], p. 124; 1994, p. 207). B:I-39. Ghaziabad District; Uttar Pradesh, INDIA; ca. 28°40'N, 77°26'E; observed 1981-1983 by R K. Seth, S. Seth, G. L. Reddy, and R K. Chopra (1992, p. 62). A:I-41. Ghazipur; Camilla, BANGLADESH; 23°32'N, 91°08'E; introduced population, reported before 1986 by M. A. R. Khan (1985, p. 31). Not mapped. GhodaGhodi Tal; Kailali, NEPAL; 28°41'N, 80°57'E; observed in 1998 by M. K. Chalise and M. Ghimire (1998, p. 12). A:N-supplemen- tary. Ghora Dhaka, 1 mi (= 1.6 km) east of, Hazara District; North-West Frontier, PAKISTAN; 34°02'N, 73°26'E; collected 18 Sept. 1962 by H. W Setzer; usnm, I. A:P-12. Ghori Hill, Dangs District, 325 m; Gujarat, IN- DIA; 20°51'N, 73°33'E; observed 8 Jan. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 466). A:I-99. Gin Keo Ho (= Giakeoho), cliff above; Sichuan, CHINA; 29°20'N, 103°05'E; collected 18 July 1925 by D. G. Graham (1926-29, p. 31; un- published map, USNM, archives, no. 89413); USNM, 1. C:C-140. Gokama, King's Forest, 5 mi (= 8 km) northeast of Katmandu, 4500 ft (= 1370 m); Bagmati, NEPAL; ca. 27°43'N, 85°23'E; collected 24 Dec. 1966 by C. O. Maser; fmnh, 1; ups, 1 (skin only). A:N-12. Gokavaram, Krishna District, 1 m; Andhra Pra- desh, INDIA; 16°16'N, 81°13'E; observed 15 May 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 468). A:M41. Gokteik. See Pyaunggaung. Golaghat, Sibsagar District, 300 ft (= 90 m); A^- sam, INDIA; 26°3rN, 93°58'E; collected 4 Jan. 1920 by H. W. Wells (Hinton & Lindsay, 1926, p. 385); BM(NH), 1. B:l-22. 138 FIELDIANA: ZOOLOGY I Golog Zangzu Zizhizhou (prefecture); Qinghai. CHINA: ^32 30'-35^^0'N. 96"50'-102'30'; re- ported 1959-1961 by Chang Chieh and Wang Tsung-yi (1963. p. 126). Not mapped. Gonda vicinity; Uttar Pradesh. INDIA; ca. 27°08'N. 81°56'E; blood samples collected 16- 27 Apr. 1964 by K. V. Shah and C. H. South- wick (1965, p. 489). A:I-59. Gongbo'gyamda Xian; Xizang (= Tibet). CHINA; ca. 29°55'N. 93°15'E; observed 1979-1982 by Zhang Cizu. Director. Shanghai Zoo (pers. comm.. 18 Oct. 1985). B:C-8. Gorkha. See Chengli. Gorubathan Forest; West Bengal. INDIA; ca. 26°58'N. 88°42'E; observed Mar.-Apr. 1985 by R. P. Mukherjee. S. Chaudhuri. and A. Murmu (1995. p. 27). A:I-8. Government Press. See Aligarh. Govindaraopeta. Mulug Taluk. Warangal District; Andhra Pradesh. INDIA; 18°1 1 'N. 80°08'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984. p. 58; 1992. p. 18). A:I-120. Guangdong. CHINA; 20°-26°N. 1 10°-1 17°E; cap- tive purchased in Shanghai 1923-1924 by F R. Wulsin (letter. 9 Jan. 1925; lsnm archives, no. 85377); mcz. 1. Not mapped. Guangdong or Guangxi. CHINA; 20°-27°N. I04°-l 17°E; collected 1923-1924 by F R. Wul- sin (letter. 9 Jan. 1925; lsnm archives, no. 85377); mcz, 1. Not mapped. Guangli, 50 m above, Hongshui He. right bank. Tian'e Xian. 260 m; Guangxi. CHINA; 25°12'N. 106°56'E; observed 30 Oct. 1992 by J. Fooden (cf. Fooden et al.. 1994. p. 623). C: C-173. Guangnan; Yunnan, CHINA; 24°03'N, 105°03'E; reported before 1998 (Zhang et al.. 1997. p. 58). C:C-153. Guangxi. CHINA; 21°-27°N. 104°-112°E; pur- chased in Shanghai Nov. 1922 by F R. Wulsin; USNM, 4 (including 1 skin only). Collected 1951-1980 by museum collectors; smnh. 22 (including 16 skins only. 2 skulls only). Not mapped. Guangze Xian; Fujian. CHINA; ca. 27°30'N. 117°24'E; reported Sept. 1981 by Zheng Xue- qing (1984. p. 145). C:C-79. Guanxian; Sichuan. CHINA; 3rOO'N, 103°37'E; reported before 1998 (Zhang et al., 1997. p. 58). C:C-30. Gudari; Orissa, INDIA; 19°21'N. 83°47'E; col- lected 10 Sept. 1927 by A. V. Sundaram and A. H. Bishop; bm(nh). 1 (skin only). A:I-108. Guidong; Hunan. CHINA; 26°12'N. 114°00'E; re- ported before 1998 (Zhang et al.. 1997. p. 58). C:C-102. Guixi; Guangxi. China ca. 21°50'N. 109°40'E; re- ported before 1998 (Zhang et al.. 1997. p. 58). C:C-216. Guiyang; Guizhou. CHINA; 26°35'N. 106°43'E; reported before 1998 (Zhang et al.. 1997. p. 58). C:C-I25. Guizhou. CHINA; 25°-29°N. 104°-109°E; col- lected Jan. 1962 by unknown collectors; Kiz, 2. Immunological survey conducted before 1996 by Duan Xiangsheng. Liu Yuanwei. Wu Jing. Dao Weiying. and Liu Jianghai (1995. p. 411). Not mapped. Gulin; Sichuan. CHINA; 28°07'N, 105°5rE; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-131. Gulong Shan Water Regulation Forest Reserve; Guangxi. CHINA; ca. 23°05'N. 106°40'E; ob- served 1976. 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 128; letter. Aug. 1996). C:C-167. Gumti Sanctuary; Tripura. INDIA; ca. 23°30'N. 9r40'E; reported before 1990 by Ranjitsinh (1990. p. 435; Gupta. 1994. p. 102). B:I-40. Gundi. Asifabad Taluk. Adilabad District; Andhra Pradesh. INDIA; 19°22'N. 79°20'E; reported Feb. 1977-July 1980 by G. U. Kump (1984, p. 58; 1992, p. 18). A:I-I13. Guniujiang. Shitai-Qimen Xian; <700 m; Anhui. CHINA; ca. 30°05'N, 117°30'E; observed 1973-1986 by Xiong Chenpei (Wada et al., 1986. p. 83). C:C-48. Guolo Prefecture. See Golog Zangzu Zizhizhou. Gupo Shan Water Regulation Forest Reserve; Guangxi. CHINA; ca. 24^39'N, lir33'E; ob- served 1976. 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 128; letter. Aug. 1996). C:C-199. ' Gurjal. Asifabad Taluk. Adilabad District; Andhra Pradesh. INDIA; 19°02'N, 79°30'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984. p. 58; 1992. p. 18). A:-112. Gurkha. See Chengli. Gusalek, north of; Konarha. AFGHANISTAN; ca. 35°05'N, 70°40'E; reported before 1972 by A. Puget (1971. p. 201). A:A-3. Gwaldri Valley, northern Dir District; North-West Frontier. PAKISTAN; ca. 35°25'N. 72°04'E; reported before 1978 by T. J. Roberts (1977. p. 86). A:N-2. Gyaca Xian; Xizang (= Tibet). CHINA; ca. 29°25'N. 92°40'E; observed 1979-1982 by FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE. MACACA MULATTA 139 Zhang Cizu, Director, Shanghai Zoo (pers. comm., 18 Oct. 1985). B:C-7. Gyala, above, 2800 m; Xizang (= Tibet), CHINA; ca. 29°38'N, 94°56'E; observed 17 July 1913 by F. M. Bailey (1914. map; 1915, p. 74; 1957, p. 122). 3:0-12. Gyirong Subcounty; Xizang (= Tibet), CHINA; ca. 28°30'N, 85°15'E; Observed summer 1945 by H. Harrer (1982, p. 85). Observed in 1982 by Zhang Cizu, Shanghai Zoo (Zhang et al., 1991, p. 177; letter, 3 July 1996). A:C-1. Hadya, Sural District, 250 m; Gujarat, INDIA; 21°05'N, 73°38'E; observed 24 Jan. 1980 by J. Fooden, A. Mahabal. and S. S. Saha (1981, p. 466). A:I-99. Hainan Dao; Hainan, CHINA; 18°-20°N, 109°- 111°E; reported before 1736 by J. B. du Halde (1735, p. 233). Purchased 20 Dec. 1919 in Guangzhou by R. Mell (1922, p. 11); zmb, 1 (skin only). Tissue samples obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). Not mapped. Haiyang Shan Water Regulation Forest Reserve Giiangxi, CHINA; ca. 25°14'N, 110°42'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127; letter, Aug. 1996). C:C-192. Haldhibari Block, Kaziranga Wild life Sancturary. 75 m; Assam, INDIA; ca. 26°37'N, 93°22'E; ob- served 24-25 Mar. 1972 by R Lahan and R. N. Sonowal (1974, pp. 261, 278; Spillett, 1967, p. 503). B:I-21. Haldwani; Uttar Pradesh, INDIA; 29°13'N, 79°31'E; observed Jan.- Dec. 1965 by M. K. Neville (1968b. p. 114). A:I-32. Haldwani vicinity, bhabar forest, subtropical pine belt, and tropical moist deciduous forest; Uttar Pradesh, INDIA; ca. 29°13'N, 79°31'E; ob- served Jan.-Dec. 1965 by M. K. Neville (1968b. p. 114). A:I-32. Halwapura, 5 mi (= 8 km) from Kaukori; Uttar Pradesh, INDIA; ca. 26°54'N, 80°48'E; ob- served Feb. 1963 by R Jay (1965, pp. 200, 249). A:I-66. Hanmajipet (= Hanumajipet), Banswada Taluk. Nizamabad District Andhra Pradesh, INDIA; 18°25'N. 78°00'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992. p. 18). A:I-117. Hanumangarhi Hill. near. Naini Tal District, 2000 m; Uttar Pradesh, INDIA; ca. 29°23'N. 79°27'E; observed Sept. 1978 by S. M. Das and B. D. Sharma (1981, p. 496). A:I-32. Hanuman Junction. Krishna District. 20 m; An- 140 dhra Pradesh, INDIA; 16°38'N. 80°58'E; ob- served 16 May 1980 by J. Fooden. A. Mahabal. and S. S. Saha (1981, p. 468). A:I-140. Harargaj; Moulvi Bazar, BANGLADESH; ca. 24°25'N. 92°00'E; observed eariy in 1980 by S. R Gittins and A. W. Akonda (1982, p. 278). B: Ba-I3. Harbhajwala. See Dehra Dun vicintiy. Harchandi Sahai, southwest of Puri; Orissa, IN- DIA; ca. 19°48'N, 85°50'E; observed 15-17 Feb. 1980 by R. R Mukherjee (1984, p. 260). A:I-106. Harduaganj; Uttar Pradesh, INDIA; ca. 27°56'N, 78°10'E; observed 1959-1975 by C. H. South- wick and M. F Siddiqi (1977, p. 342). A:I-69. Hardwar; Uttar Pradesh, INDIA; 29°58'N, 78°10'E; observed Sept. 1959-Feb. 1960 and 1964-1965 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961a, p. 543; Southwick and M. R. Siddiqi, 1966, p. 309). A:I-27. Harinbhanga; West Bengal, INDIA; ca. 21°45'N, 89°00'E; observed 1955-1960 by A. K. Mu- kherjee (Mukherjee & Gupta, 1965, p. 145). B: I-l. Haripur, Kheri District; Uttar Pradesh, INDIA; ca. 28°07'N, 80°43'E; collected Mar. 1932 by C. McCann; bnhs, 1 (skin only). A:I-55. Hamarayan Mohalla. See Khair, Tahsil. Hasimara, Bhutan Duars, 550 ft (= 170 m); West Bengal, INDIA; ca. 26°45'N, 89°20'E; collected 1-10 Jan. 1916 by N. A. Baptista (H. V. O'Donel in Wroughton, 1917, p. 63); bm(nh), 3; BNHS, mounted skin, on exhibit. A:I-9. Hastings Road. See Calcutta. Hathras; Uttar Pradesh, INDIA; 27°36'N, 78°03'E; observed Sept. 1959-Feb. I960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961a, p. 543). A:I-69. Hatighuli; Assam, INDIA; ca. 26°56'N, 94°31'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. Hazara District, southern; North-West Frontier, PAKISTAN; ca. 34°00'N, 73°00'E; observed 1914-1916 by R. W. G. Hingston ([1920], p. 243). A:P-8. Hazaria Patherghatta, 600 ft (= 180 m); Narayani, NEPAL; ca. 27°00'N, 85°15'E; collected 17 Feb. 1921 by N. A. Baptista (Hinton & Fry, 1923, p. 402); bm(nh), I (topotype of Cerco- pithecus (Miilatta) Zimmerman, 1780; see Po- cock, 1932. p. 533). A:N-13. Hazarikhil; Chittagong, BANGLADESH; ca. 22°20'N. 92°00'E; observed Feb. 1990-June FIELDIANA: ZOOLOGY J 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-35. Hechi Prefecture; Guangxi. CHINA; ca. 24°42'N, 108°02'E; acquired in 1992 from Chinese Med- icine Division, Hechi; izcas, 3 (skulls only). B: C-178 Heinsun (= Heinsum; Heinzun), east bank of Chindwin River; Sagaing, MYANMAR ( = BURMA); 25°52'N, 95°35'E; collected 11 Mar. 1935 by H. C. Raven (in Carter, 1943, p. 1(X); Morris, 1936. p. 665); amnh, 2. B:M-9. Hejiang; Sichuan, CHINA; 28°50'N, 105°46'E; reported before 1992 by Jiang Xuelong. Wang Yingxiang. and Ma Shilai (1991, p. 244). C:C- 136. Hekou; Yunnan, CHINA; 22°36'N. 103°58'E; re- ported before 1998 (Zhang et al., 1997. p. 58). C:C-152. Heluwabeshi; Dhankuta, NEPAL; 27°26'N, 87°08'E; observed in 1998 by M. K. Chalise and M. Ghimire (1998. p. 12). B:N-supplemen- tary. Henron; Hainan Dao; Hainan, CHINA; not pre- cisely located, 18-20°N, 109-1 11°E; collected 10 May 1904 by A. Owston (Allen, 1906, pp. 465, 488); amnh, 1. Not mapped. Himchari; Cox's Bazar, BANGLADESH; ca. 21°45'N, 92°00'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-38. Hindan Bridge Temple. See Delhi. Hissar District; Haryana, INDIA; ca. 29°26'N. 75°18'E; observed 1981-1983 by R K. Seth, S. Seth, G. L. Reddy, and P K. Chopra (1992, p. 62). A:I-39. Hisweht (= Hiswet), west bank of Nantaleik Chaung, above Tamanthi, upper Chindwin Riv- er, 460 ft (= 140 m); Sagaing, MYANMAR ( = BURMA); ca. 25°22'N, 95°16'E; collected 25 Aug. 1914 by G. C. Shortridge and S. A. Mac- millan (Shortridge in Wroughton, 1916a, p. 293); BM(NH), 1. B:M-12. Hitaura, 0.5 mi (= 1 km) west, 1450 m; Chisa- pani, NEPAL; 27°27'N, 85°02'E; recorded 12 Feb. 1967 by C. O. Maser (field notebook, FMNH, p. 39); specimen not located. A:N-10. Hkamti. See Singkaling Hkamti. Hkandau, 2000 ft (= 610 m); Kachin, MYAN- MAR (= BURMA); 26°0rN, 97°50'E; collect- ed 8 Aug. 1939 by R. Kaulback; bm(nh). 1 (skin only). B:M-2. Hoa Binh; Hoa Binh, VIETNAM; ca. 20°49'N, 105°22'E; collected 1 June 1972 by Dang Huy Huynh and Truong Van Le; iebr, 1. C:V-20. Hoa Binh, VIETNAM; 20°20'-21°00'N, 104°50'- 105°50'E; collected 19 July 1960 by unknown collector; zmynh, 1 (skin only). Not mapped. Hoi Xuan, Quan Hoa District; Thanh Hoa, VI- ETNAM; 20°23'N, 105°()6'E; collected Mar. 1964 by unknown collector (Dao, 1985. p. 196); museum unknown (not seen). 2. C:V-22. Homalin, west bank of upper Chindwin River. 400 ft (= 120 m); Sagaing, MYANMAR (= BUR- MA); 24°52'N, 94°55'E; collected 14-15 July 1914 by G. C. Shortridge and S. A. Macmillan (Shortridge in Wroughton, 1916a, p. 293); bm(nh), 2: BNHS, 1. B:M-14. Hong Kong. See Xianggang. Hongshui He, between Tian'e and Hal Zhou, 4- 5 km northwest of Tian'e. Tian'e Xian; Guang- xi, CHINA; ca. 25°01'N. 107°07'E; captured Apr. 1990 by Luo Mingfei (pers. comm.. 15 Oct. 1992); captive observed 15 Oct. 1992 at Tian'e. C:C-173. Hongshui He, left bank, 9 km below Heke and 10 km below Heke, Tian'e Xian, 260-360 m; Guangxi, CHINA; 25°10'N, 106°58'E observed 29-30 Oct. 1992 by J. Fooden (cf. Fooden et al., 1994, p. 623). C:C-173. Hongshui He, right bank, 500 m below Heke, Tian'e Xian, 260 m; Guangxi, CHINA; 25°14'N, I06°58'E; observed 19 Oct. 1992 by J. Fooden (cf. Fooden et al., 1994, p. 623). C: C-173. Hopeh. See Xinglong Xian, southern. Hoshangabad. See Sohagpur. Hot Mix Plant. See Aligarh. Hot Springs (= Garampani). Jaintia Hills, 2400 ft (= 730 m); Assam, INDIA; 25°3rN, 92°34'E; collected 18 July 1920 by H. W. Wells (Hinton 6 Lindsay, 1926, p. 385); bnhs, 1. B:I-19. Hotha Valley; Yunnan, CHINA; ca. 24°25'N, 97°55'E; captive purchased July 1868 by J. An- derson (1876, p. 275; 1879, pp. xvi, 56); zsi, 1. B:C-61. Houmda. See Ngamda. Hsignolo. See Xi Golog. Hsi-kiang. See Xi Jiang. Hsi-o-lo. See Xi Golog. Htingnan Triangle, 3500 ft (= 1070 m); Kachin, MYANMAR (= BURMA); 26°36'N. 97°53'E; collected 28 Jan. 1939 by R. Kaulback; bm(nh), 1. B:M-1. Huagong Water Regulation Forest Reserve; Guangxi, CHINA; ca. 24°3rN. 104°58'E; ob- served 1976. 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C-16I. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 141 Huai Ap Nang, right bank of Mae Nam Ping, 350 m; Tak, THAILAND; 17°25'N, 98°43'E; col- lected 29 Mar. 1967 by J. Fooden ( 1971, p. 18); FMNH, 1. B:T-5. Huaiji; Guangdong, CHINA; 23°55'N, 1I2°10'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-2()7. Huai Kwang Pah, left bank of Mae Nam Ping, 350 m; Tak, THAILAND; 17°28'N, 98°50'E; collected 29 Mar. 1967 by J. Fooden (1971, p. 18); CTNRC, 1; fmnh, 1 (in alcohol). B:T-5. Huanglian Shan Water Regulation Forest Reserve; Giiangxi, CHINA; ca. 23°35'N, 106°25'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C-168. Huangliangping, Xingshan Xian; Hubei. CHINA; ca. 31°22'N, 110°53'E; observed Mar.-July 1982 by E E. Poirier and Hu Hongxhin (1983, p. 387; Poirier, 1983, p. 124). C:C-44. Huangqiao, Wuyanlin Nature Reserve, Taishun Xian. 650 m; Zhejiang, CHINA; 27°42'N, 119°48'E; collected Apr. 1981 by Zuge Yang (pers. comm., 24 Oct. 1985); hubd, 1. C:C-68. Huangshan; Anhiii, CHINA; 30°10'N, 118°07'E; tissue sample obtained ca. 1991 by Zhang Ya- ping and Shi Liming (1993b, p. 589). C:C-62. Huangtankou, Suichang Xian; Zhejiang, CHINA; 28°50'N, 118°54'E; captives acquired ca. Aug. 1981 by Fu Yiyuan and Wu Fuhai, Hangzhou Zoo (pers. comm., 25 Oct. 1985). C:C-64. Huaping Nature Reserve; Guangxi, CHINA; ca. 25°36'N, 109°54'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 126; letter, Aug. 1996). C:C-184. Huaping [Xian]; Yunnan, CHINA; ca. 26°37'N, 101°I3'E; blood sample obtained before 1999 by Ding Bo, Zhang Yaping, and Hou Yidi (1998, p. 172). B:C-47. Huashi, north of; Hebei CHINA; ca. 40°24'N, 117°30'E; reported 1940-1980 by local resi- dents (Zhang et al., 1989, p. 379). C:C-1. Hubei, CHINA; 29°-33°N, 109°-1I6°E; reported before 1992 by Jiang Xuelong, Wang Yingxiang, and Ma Shilai (1991, p. 244). Not mapped. Hue, 0 m; Thua Thien-Hue, VIETNAM; 16°28'N, 107°36'E; collected Oct.-Dec. 1925 by J. De- lacour, P. Jabouille, and W. P. Lowe (Delacour et al., 1927, p. 132; Delacour, 1940, pp. 21, 24); ?MNHN, 1-2 (not seen). C:V-33. Huidong; Guangdong, CHINA; 22°58'N, 1 14°44'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-209. Huishui; Guizhou, CHINA; 26°08'N, 106°36'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-124. Huixan; Gansu, CHINA; 33°46'N, 106°06'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-17. Huixan; Henan, CHINA; 35°32'N, 113°54'E; tis- sue sample obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). C:C-8. Hui-yao; Yunnan, CHINA; ca. 25°15'N, 98°30'E; collected 26 Apr. 1917 by R. C. Andrews (in Andrews & Andrews, 1918, pp. 298, 305); AMNH, 2; Mcz, 1. B:C-58. Hule (= Huleu), Ningguo Xian, 200-500 m; An- hui, CHINA; ca. 30°2rN, 118°47'E; observed 1973-1986 by Xiong Chenpei (Wada et al., 1986, p. 83). C:C-55. Hunan, CHINA; 25°-30°N, 110°-114°E; captives obtained Oct.-Nov. 1973 and May 1976 by mu- seum collectors; sciea, 3. Not mapped. Huong Binh. See Song-Ta-Voy. Huong Him. See Song-Ta-Voy. Huong Son (= Muong Son) District; Ha Tinh, VI- ETNAM; ca. 18°31'N, 105°28'E; collected 19 June 1985 by Nguyen Van Dung; fcxm, 1 (skull only). C:V-27. Hutu Forest, above Rara Daha (= Lake), 3200 m; Jumla, NEPAL; ca. 29°35'N, 82°05'E; observed Oct. 1975 by T Richie and R. Shrestha (Richie et al., 1978, p. 443; cf. Fooden, 1989, p. 44). A:N-6. Hwang Liang Commune. See Huangliangping. Hyderabad, 560 m; Andhra Pradesh, INDIA; 17°23'N, 78°29'E; observed 31 Mar. and 22 Apr. 1980 by J. Fooden. A. Mahabal, and S. S. Saha (1981, p. 467). A:I-124. Ibrahimpet, Banswada Taluk, Nizamabad District; Andhra Pradesh, INDIA; 18°23'N, 77°55'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-117. Ichang. See Yichang. Imphal, ca. 4 mi (= 6.5 km) north of milestone 129 [km] on Dimapur Road, 3000 ft (= 900 m); Manipur, INDIA; 24°52'N, 93°56'E; col- lected 6 Nov. 1945 by M. L. Roonwal (1949, p. 84; 1950, p. 16 [misidentified as M. assa- mensis]); zsi, 1. B:I-37. Imphal, Mahabali Temple, 762 m; Manipur, IN- DIA; 24°49'N, 93°57'E; observed May-June 1974 and Feb. 1975 by R. R Mukherjee (1978a, p. 276). B:I-37. Indian Museum compound. See Calcutta. Imperial Hunting Grounds. See Xinglong Xian, southern. 142 FIELDIANA: ZOOLOGY INDIA; 15°-35°N, 71°-97°E; collected before 1843 by Mr. Cross; bm(nh), 1 (skin only). Col- lected before 1844 by B. H. Hodgson; bm(nh). 1 (skull only). Collected before 1852 by un- known collectors; bm(nh), 2. Collected 15 Dec. 1925 by R. P. Page; bm(nh), 1 (skin only). Date and collector unknown; zsi. 2 (skull only). Not mapped. 7INDIA; 15°-35°N, 71°-97°E; collected before 1852 by unknown collector; bm(nh), 1 (skin with skull inside). Not mapped. Irrawaddy River, left bank, below Yenangyaung; Magwe, MYANMAR (= BURMA); ca. 20°27'N, 94°52'E; observed before 1879 by J. Anderson (1879, p. 57). B:M-26. Jagannath Temple. See Puri. Jagatsukh; Himachal Pradesh, INDIA; ca. 32°10'N, 77°15'E; reported 1978-1980 by A. J. Gaston, P. J. Garson, and M. L. Hunter, Jr. (1983, p. 300). A:I-14. Jaggayyapet, Guntur District, 50 m; Andhra Pra- desh, INDIA; 16°53'N, 80°06'E; observed 21 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 468). A:I-131. Jaggayyapet, 4 km north of, Guntur District, 75 m; Andhra Pradesh, INDIA; 15°55'N, 80°07'E; observed 21 Apr. 1980 by J. Fooden, A. Ma- habal, and S. S. Saha (1981, p. 468). A:I-131. Jagvedi Temple. See Chitrakut. Jaintia Hills. See Hot Springs; Narpuh Reserved Forest. Jaipur; Rajasthan, INDIA; 26°55'N, 75°49'E; ob- served Mar. 1956-Oct. 1960 by I. Prakash (1958, p. 154; 1962, p. 83). Observed ca. 1982- 1988 by R. Mathur and L. D. Wolfe (Mathur, 1982, p. 11; Wolfe & Mathur, 1988, p. 535; Wolfe, 1992, p. 44). A:I-75. Jaipur District; Rajasthan, INDIA; ca. 26°20'- 27°50'N, 74°55'-76°50'E; observed 1981-1983 by R K. Seth, S. Seth, G. L. Reddy, and R K. Chopra (1992, p. 62). Not mapped. Jakaram, 4 km southwest of, Warangal District, 325 m; Andhra Pradesh, INDIA; 18°08'N, 79°53'E; observed 19 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A: I- 1 20. Jakka. See Sungri, ca. 2 km south of. Jakko (= Jako; Jakoo; Jaku) Hill, Simla, 8500 ft (= 2600 m); Himachal Pradesh, INDIA; 31°06'N, 77°10'E; observed Feb. 71836 by T Hutton (1837, p. 935). Observed before 1888 by W T Blanford (1888b, p. 14). Observed Aug. 1972-Feb. 1973 by K. Wada (1984, p. 477). A:I-18. Jakkpom, Banswada Taluk, Nizamabad District; Andhra Pradesh, INDIA; not precisely located, 18°05'-18°35'N, 77°45'-78°05'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). Not mapped. Jaldapara Wildlife Sanctuary; West Bengal, IN- DIA; ca. 26°38'N. 89°18'E; observed 23 Oct. 1965 by M. Krishnan (1972, p. 540). Reported Jan.-June 1966 by J. J. Spillett (1967, p. 549). B:I-9. Jalpaiguri. See Bharnabhari; Hasimara. Jamduar vicinity, <270 m; Assam, INDIA; ca. 26°43'N, 89°53'E; observed in 1957 by G. A. von Maydell (Oboussier & von Maydell, 1959, p. 106). Observed May-June 1973 by R. R Mu- kherjee and S. S. Saha (1974, p. 337; Mukher- jee, 1978b, p. 742). B:I-10. Jammu; Jammii & Kashmir, INDIA; 32°44'N, 74°52'E; observed ca. 1982 by Y. R. Malhotra and D. N. Sahi (1982, p. 27). A:I-5. Jammu District; Jammu & Kashmir, INDIA; 32°20'-32°55'N, 74°40'-75°20'E; observed 1981-1983 by R K. Seth. S. Seth, G. L. Reddy, and R K. Chopra (1992, p. 62). Not mapped. Jammu & Kashmir, southern border with Paki- stan, INDIA; ca. 32°30'N, 74°40'E; trapped Nov. 1976 by animal dealer (Remfry, 1982, p. 144). A:I-6. Jantschin. See Yanjing. Japisojia; Assam, INDIA; ca. 27°03'N, 94°48'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. Jawan; Uttar Pradesh, INDIA; 28°02'N, 78°06'E; observed 1959-1975 by C. H. Southwick and M. F Siddiqi (1977, p. 342). Observed Jan. 1990-Mar. 1991 by E. Imam and H. S. A. Yah- ya (1995, p. 2). A:I-69. Jaypore Agency. See Malkangiri. Jayrapar; Assam, INDIA; ca. 27°02'N, 94°39'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury (11991a], p. 31). B:I-25. Jegu Xiang, Yushu Xian; Qinghai, CHINA; 32°42'N, 97°15'E; collected 30 May and 10 June 1963 by Shou Zhongchan; nwpib, 2 (in- cluding 1 skin only). B:C-18. Jenli, 2-3 km north of, Mulun Subcounty, Huan- jiang Xian, 670 m; Guangxi, CHINA; ca. 25°07'N, 108°0rE; collected fall 1991 by Hu Huguan (pers. comm., 5 Nov. 1992); skeleton examined 5 Nov. 1992 at Jenli. C:C-177. Jessore, BANGLADESH; ca. 23°15'N, 89°15'E; reported before 1982 by M. A. R. Khan (1981, p. 13). B:Ba-19. Jeypore Agency. See Malkangiri. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 143 Jhajjarkotli, Udhampur Subdistrict; Jammii & Kashmir. INDIA; ca. 32°56'N, 75°08'E; ob- served by Y. R. Malhotra and D. N. Sahi (1982, p. 27). A:I-10. Jhanji; Assam, INDIA; ca. 26°52'N, 94°30'E; re- ported 9 Mar. 1987-16 Feb. 1988 by A. Choud- hury ([1991a], p. 31). B:I-25. Jhansi; Uttar Pradesh, INDIA; 25°26'N, 78°35'E; bacteriological survey conducted before 1985 by J. P. Tiwari and A. K. Shukla (1984. p. 498). A:I-83. Jhilla; West Bengal, INDIA; ca. 22°00'N, 89°00'E; observed 1955-1960 by A. K. Mu- kherjee (Mukherjee & Gupta, 1965, p. 145). B: 1-2. Jhima. ca. 17 mi (= 27 km) west of Ramnagar, Kumaun region, 1500 ft (= 460 m); Uttar Pra- desh, INDIA; 29°27'N, 78°54'E; collected 17 Jan. 1914 by C. A. Crump (in Wroughton. 1914, p. 284); zsi, 1. A:I-33. Jian'ou Xian; Fiijian, CHINA; ca. 27°03'N, 1I8°19'E; reported Aug. 1980 by Zheng Xue- qing (1984, p. 145). C:C-72. Jianfengling, Ledong Xian, Hainan Dao, 800-900 m; Hainan. CHINA; 18°43'N, 108°53'E; col- lected 5 Dec. 1962 by Liu Zhenhe, sciea (pers. comm., 26 Nov. 1985); sciea, I. Reported be- fore 1982 by Zeng Qingsong (1982, p. 69). C: C-236. Jiangkou; Guizhou, CHINA; 27°42'N, 108°50'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-1I5. Jianyang Xian; Fujian. CHINA; ca. 27°20'N. 118°07'E; reported July 1982 by Zheng Xue- qing (1984, p. 145). C:C-73. Jiaqiaolin Water Regulation Forest Reserve; Guangxi, CHINA; ca. 24°46'N, 110°06'E; ob- served 1976, 1986, 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127; letter. Aug. 1996). C:C- 194. Jigzhi (= Jiuzhi); Qinghai, CHINA; 33°28'N, 10r29'E; reported before 1998 (Zhang et al., 1997, p. 58). B:C-24. Jilian, Yi Xian, 400-600 m; Anhiii, CHINA; ca. 30°00'N, 118°00'E; observed 1973-1986 by Xiong Chenpei, K. Wada, and Wang Qishan (Wada et al., 1986, pp. 83. 88). C:C-62. Jincheng; Shanxi, CHINA; 35°30'N. 1 12°50'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-6. Jinchengjiang; Guangxi, CHINA; 24°42'N, 108°02'E; tissue sample obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). C:C-178. Jind District; Haryana, INDIA; ca. 29°19'N, 76°19'E; ob.served 1981-1983 by R K. Seth, S. Seth, G. L. Reddy, and R K. Chopra (1992. p. 62). A:I-38. Jingdong Xian; Yunnan, CHINA; 24°28'N, 100°54'E; reported before 1992 by Jiang Xue- long, Wang Yingxiang, and Ma Shilai (1991, p. 243). Immunological survey conducted before 1996 by Duan Xingsheng, Liu Yuanwei, Wu Jing. Dao Weiying, and Liu Jianghai (1995, p. 411). Blood sample obtained before 1999 by Ding Bo, Zhang Yaping, and Hou Yidi (1998, p. 172). B:C-71. Jinggangshan; Jiangxi, CHINA; 26°42'N, 114°07'E; reported before 1998 (Zhang et al., 1997, p. 58). Report unverified; M. thibetana only macaque verified at this locality (Li Xiongshan, Jinggangshan Nature Reserve Bu- reau, pers. comm., 4 Nov. 1985; Liu Zhenhe, SCIEA, pers. comm., 25 Nov. 1985). Not mapped. Jinggu Xian; Yunnan, CHINA; ca. 23°28'N, 100°42'E; immunological survey conducted be- fore 1996 by Duan Xingsheng, Liu Yuanwei, Wu Jing, Dao Weiying, and Liu Jianghai (1995, p. 411). B:C-74. Jinping; Yunnan, CHINA; ca. 22°50'N, 103°15'E; collected date unknown by Ye Zongyao (Quan Guoqiang, pers. comm., 25 Aug. 1983); izcas, 1 (skull only). C:C-150. Jinzhong Shan Bird Reserve; Guangxi, CHINA; 24°45'N, 104°55'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter, Aug. 1996). C:C-155. Jirna. See Jhima. Jishou; Hunan, CHINA; 28°19'N. 109°43'E; tis- sue sample obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). C:C-113. Jiufeng Shan; Fujian, CHINA; ca. 27°00'N, 118°48'E; reported Nov. 1983 by Zheng Xue- qing (1984), p. 145). C:C-71. Jiuhua Shan. ca. 30 km east of; Anhui, CHINA; ca. 30°25'N, 118°05'E; observed 1973-1978 by Xiong Chenpei (Wada et al., 1986, p. 82). C:C- 51. Jiuhua Shan, ca. 30 km northeast of; Anhui, CHI- NA; ca. 30°40'N, 118°00'E; observed 1973- 1982 by Xiong Chenpei (Wada et al.. 1986, p. 82). C:C-52. Jiuhua Shan, Qingyang Xian, 400-800 m; Anhui, CHINA; ca. 30°27'N, 117°48'E; observed 1973-1986 Xiong Chenpei (Wada et al., 1986, p. 83). C:C-50. Jiuwanshan Water Regulation Forest Reserve; 144 HELDIANA: ZOOLOGY Giiangxi, CHINA; ca. 25°14'N. 108°43'E; ob- served 1976, 1986. 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127, letter, Aug. 1996). C:C- 181. Jiuzhi. See Jigzhi. Jixi; Anhui, CHINA; 30°05'N, 1 18°36'E; reported before 1988 (Zhang et al., 1997, p. 58). C:C- 60. Jiyuan, ca. 20 km northeast of; 750 m; Henon, CHINA; ca. 35°11'N, II2°41'E; observed 1981-1988 by Qu Wenyuan, Zhang Yongzu, D. Manry, and C. H. Southwick (1993, p. 617; cf. Southwick et al., 1991, p. 25). C:C-12. Jiyuan, ca. 30 km northwest of; 750 m; Hencin, CHINA; ca. 35°11'N, 1I2°23'E; observed 1981-1988 by Qu Wenyuan, Zhang Yongzu, D. Manry, and C. H. Southwick (1993, p. 617; cf. Southwick et al., 1991. p. 25). C:C-4. Jiyuan, ca. 80 km west-northwest of, 750 m; He- nan, CHINA; ca. 35°12'N, I12°05'E; observed 1981-1988 by Qu Wenyuan, Zhang Yongzu. D. Manry, and C. H. Southwick (1993, p. 617; cf. Southwick et al., 1991, p. 25). C:C-13. Jodhpur; Rajasthan, INDIA; 26°17'N, 73°02'E; observed before 1960 by I. Prakash (1959, p. 39). Observed May 1979 by M. L. Roonwal (Bhargava, 1984, p. 43). A:I-80. Judi-Jatakia, near Hologuri-Kalugaon; Assam, IN- DIA; ca. 26°56'N, 94°39'E; observed 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 32). B:I-25. Julongshan Nature Reserve, Suichang Xian; Zhe- jiang, CHINA; 28°21'N, 118°53'E; captives and skins observed in local shops, 1980-1981, by Tang Ziying, fubd (pers. comm., 19 Oct. 1985). Observed 1982 by Sheng Helin, ECNU (pers. comm., 19 Oct. 1985). C:C-66. Kaeng Mae Hat (rapids), Mae Nam Ping (river), below Chiang Mai. 850 ft (= 260 m); Chiang Mai, THAILAND; 17°51'N. 98°41'E; collected 14 Apr. 1916 by K. G. Gairdner (Kloss. 1917, p. 247); ZRC, 1 (holotype of Macaca siamica). B:T-4. Kafiristan. See Nurestan. Kaiyang; Guizhou, CHINA; 27°04'N, 106°58'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-127. Kakara, Damoh District, 1200 ft (= 370 m); Madhya Pradesh, INDIA; ca. 23°50'N, 79°27'E; collected 12 May 1912 by C. A. Crump (in Wroughton & Ryley. 1913, p. 46); bm(nh). 1 (skin only). A:I-94. Kakhyen Hills. See Tengchong. Kakkaraparu, Peravalli Block, East Godavari Dis- trict; Andhra Pradesh, INDIA; not precisely lo- cated, 16°40'-17°50'N, 8r30'-82°35'E; report- ed Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). Not mapped. Kakori; Uttar Pradesh, INDIA; 26°54'N, 80°48'E; observed 1 Dec. 1959-30 Mar. 1960 by P Jay (1963, p. 274; 1965, p. 212). A:I-66. Kalabokhani, Sylhet; Sylhet, BANGLADESH; ca. 24°54'N, 9r52'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-10. Kali Bari Temple. See Simla. Kam Shan Entrance, Kowloon Reservoir Area; Xianggang (= Hong Kong), CHINA; ca. 22°22'N, 114°09'E; population probably artifi- cially introduced (Herklots, 1951, p. 83). Ob- served July-Aug. 1980 and July-Aug. 1981 by C. H. Southwick and K. L. Southwick (1983, p. 19). Observed 10 Jan.-3 Feb. 1987 by C. H. Southwick and D. Manry (1987, p. 48). Ob- served Feb. 1995 by K. A. Bolton, V. M. Camp- bell, and F D. Burton (1998, p. 197). C:C-210. Kamdech. See Landay Sind. Kamta, Mukhrbind Temple, Banda District; Uttar Pradesh, INDIA; not precisely located, 24°50'- 25°50'N, 80°10'-81°35'E; observed Sept. 1959-Feb. 1960, 1964-1965. and 1979-1980 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961a, p. 543; Southwick and Siddiqi, 1966, p. 312; Southwick et al., 1983, p. 437). Not mapped. Kamu Valley; Konarha, AFGHANISTAN; ca. 35°25'N, 71°25'E; observed July-Aug. 1970 by C. Naumann and G. Nogge (1973, p. 92). A: A-1. Kangding (= K'ang-ting; Tasin Lou); Sichuan, CHINA; 30°03'N, 102°02'E; collected June- July 1890 by H. d' Orleans (Bonvalot, 1891, vol. 2, p. 210; 1892, p. 506); mnhn, 1 (holotype of Macacus vestitus). B:C-27. Kangra, 2500 ft (= 760 m); Himachal Pradesh, INDIA; 32°()6'N, 76°16'E; collected 20 Mar. 1921 by H. Whistler; bnhs, 2. Trapped in 1976 by R. V. Henrickson (Karr et al., 1979, p. 789; 1980, p. 201). A:I-12. Kangra Fort, 2450 ft (= 750 m); Himachal Pra- desh, INDIA; ca. 32°05'N, 76°16'E; collected 18 Mar. 1921 by H. W. Wells (Lindsay, 1926, p. 599); BM(NH). 1. A:I-12. Kanha National Park; Madhya Pradesh, INDIA; ca. 22°20'N, 80°40'E; reported before 1997 by K. K. Gurung and R. Singh (1996, p. 96). A:I- supplementary. Kangxian; Gansii, CHINA; 33°26'N, 105°37'E; FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 145 reported before 1998 (Zhang et al., 1997, p. 58). C:C-19. Kansrao; Uttar Pradesh, INDIA; 30°05'N, 78°08'E; observed spring 1953 by A. Nolle (1956, p. 180). A:I-27. Kanti vicinity, Chitral District; North-West Fron- tier, PAKISTAN; ca. 35°35'N, 71°4rE; report- ed before 1978 by T. J. Roberts (1977, p. 86). A:P-1. Kaotai (= Kootai), lower Kunar (= Chitral) River, 3600 ft (= 1100 m); North-West Frontier. PAKISTAN; ca. 35°20'N, 71°35'E; collected early Feb. 1914 by F. D. Stirling (Wroughton, 1918, p. 553); bm(nh), 1 (holotype of Macaca mulatto mcmahoni). A:P-1. Kao Tien. See Kuatun. Kaptai; Rangamati, BANGLADESH; 22°2rN, 92°17'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-35. Kaptai, south of; Chittagong, BANGLADESH; ca. 22°00'N, 92°00'E; tentatively reported July- Nov. 1976 by K. M. Green (1978, p. 146). B: Ba-38. Kaptai Lake; Rangamati, BANGLADESH; ca. 22°30'N, 92°14'E; observed 1975-1976 by J. R. Oppenheimer, A. W. Akonda, and K. Z. Husain (1983. p. 193). B:Ba-35. Karampodu, Ipur Taluk, Guntur District; Andhra Pradesh, INDIA; not precisely located. 16°05'- 16°25'N. 79°35'-79°55'E; reported Feb. 1977- July 1980 by G. U. Kurup (1984, p. 58; 1992. p. 19). Not mapped. Kareilly, west of; Uttar Pradesh, INDIA; ca. 28°16'N, 79°22'E; observed Apr.-June 1965 by R. P Mukherjee (1969, p. 47). A:l-52. Karen Chaung, Pidaung Reserve, Myitkyina Dis- trict, 500 ft (= 150 m); Kachin, MYANMAR (= BURMA); ca. 25°25'N, 97°15'E; collected 25 May 1936 by H. C. Smith (Napier. 1981, p. 21); BM(NH), 2 (including 1 skin only). B:M-5. Karghena, west of; Uttar Pradesh, INDIA; ca. 28°16'N, 79°22'E; observed Apr.-June 1965 by R. P Mukherjee (1969, p. 47). A:I-52. Karkara. See Kakara. Karkatgarh (= Karkatnagar); Bihar, INDIA; ca. 24°53'N, 83°22'E; observed 1959-1970 by M. Krishnan (1972, p. 540). A:I-89. Karnal District; Haryana, INDIA; ca. 29°44'N, 76°44'E; observed 1981-1983 by P K. Seth, S. Seth, G. L. Reddy, and P K. Chopra (1992, p. 62). A:I-23. Kamali River, ca. 30 km above mouth, Karnali Bardia Game Reserve; Bardia, NEPAL; ca. 28°37'N, 81°19'E; observed 20 Feb.-lO Mar. 1976 by J. Teas (1983, p. 214). A:N-5. Karnali River- Aurn River, ca. 10 km above con- fluence, Karnali Bardia Game Reserve; Bardia, NEPAL; ca. 28°26'N, 8n5'E; observed 20 Feb.- 10 Mar. 1976 by J. Teas (1983, p. 214). A:N-5. Karrachi (= Karachi). See PAKISTAN. Kasauli, Siwalik Hills; Himachal Pradesh, IN- DIA; 30°55'N. 76°57'E; observed before 1984 by M. Singh and R. S. Pirta (1983, p. 81). A:I- 18. Kashmir (region); Jammii & Kashmir, INDIA; ca. 34°N, 75°E; obtained before 1857 by Theobald Collection/ Prof. Oldham; bm(nh), 1 (skull only, species identification questionable; cf. Napier, 1981, p. 26). Acquired before 1872 by Zoolog- ical Society of London (Anderson, 1879, p. 63 Elliot, 1913, p. 202; Pocock, 1932, p. 540) BM(NH), 1. Collected in 1985 by W. L. Abbott USNM, 1. Not mapped. Kasol; Himachal Pradesh, INDIA; ca. 32°00'N, 77°20'E; reported 1978-1980 by A. J. Gaston, R J. Garson, and M. L. Hunter, Jr. (1983, p. 300). A:I-15. Kathmandu. See Katmandu. Kathpar; Assam, INDIA; ca. 27°00'N, 94°37'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. Kathua vicinity; Jammii & Kashmir, INDIA; ca. 32°22'N, 75°3rE; trapped in 1976 by R. V. Henrickson (Karr et al., 1979, p. 789; 1980, p. 201). A:I-8. Katmandu; Katmandu Valley, NEPAL; 27°43'N, 85°18'E; observed before 1979 by B. Marriott (1978b, p. [27]). A:N-12. [Katmandu Valley], NEPAL; 27.5°-28°N, 85°- 85.5°E; collected before 1845 by B. H. Hodg- son (Scully, 1888, p. 234; Napier, 1981, p. 24); bm(nh), 7 (including 3 skulls only [1 not seen]); probably includes part of type series of Maca- cus Oinops. Not mapped. Kaukori. See Kakori. Kausa Gutta, Adilabad District, 300 m; Andhra Pradesh, INDIA; 19°07'N, 78°43'E; observed 25 Mar. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 466). A:I-I14. Kazinag; Azad Kashmir, PAKISTAN; ca. 34°00'N, 73°35'E; reported before 1984 by M. Nawaz (1983, p. 6). A:P-12. Kaziranga National Park; Assam, INDIA; 26°35'- 26°45'N, 93°I0'-93°30'E; reported before 1988 by A. Choudhury (1987, p. 162). Not mapped. K. B. Road. See Dhaka. 146 FIELDIANA: ZOOLOGY Kedarnath Sanctuary. 1400-2100 m; Urtar Pra- desh. INDIA: ca. 30=25 W. 79M5E: observed 1979-1982 by A. N. Singh (1982. p. 8). A:I- 29. Kengma. See Gengma. Kerwada Forest. See Kherwada Forest. Khair. Aligarh District: L'nar Pradesh. INDIA: 27°57'Nr irSO'E: observed Sept. 1959-Feb. 1960 by C. H. Southwick. M. A. Beg. and M. R. Siddiqi ( 1961a. p. 542). Observed May-June 1976 by R. S. Pina (1984. p. 542). Observed May-June 1976 by R. S. Pina (1984. p. 272). Observed Jan. I99'0-Mar. 1991 b> E. Imam and H. S. A. Yahya (1995. p. 2). A:I-44. Khair. Tahsil: Unar Pradesh. INDIA: 27°50'- 28'10'N. 77=30'-78=05'E: observed 1975-1980 by R K. Seth. S. Seth. and A. K. Shukia (1983. p. 38; Seth & Seth. 1983. p. 63). Not mapped. Khair Inter College. See Khair. Tahsil. Khammam: Andhra Pradesh. LNDIA: 17'15'N. 80-09 E: reponed Feb. 1977-JuIy 1980 by G. U. Kurup (1984. p. 58: 1992. p. 18). A:I-132. Khanamukh: Assam, INDIA: ca. 26"57'N. 94=28 E: reponed 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a]. p. 31). B:I-25. Khanapur. 3-5 km west of. .-Xdilabad District. 350 m: Andhra Pradesh. INDIA: 19=04N. 78=37 E: observed 24 Mar. 1980 by J. Fooden. A. Ma- habal. and S. S. Saha (1981. p. 466). A:I-114. Khao Nang Rum. western slope: Uthai Thani. THAILAND: ca. 15=30'N. 99^17'E: observed 1973-1977 by A. A. Eudey (1979. pp. 91-97. table 7). B:T-9. Khao Nang Rum Research Station. 4(X)-7(X) m: Ujhai Vumi. TFLA1L.\NT): ca. 15=28'N. 99"18'E: observed Mar. -May 1988 by S. Srikosamatara (1993. p. 37). B:T-9. Khasi Hills. See Nongpoh. Khatuajhuri: West Bengal. INDIA: not precisely located. 2r30'-22=15N. 88=40-89=10E: ob- served 1955-1960 by A. K. Mukherjee (Mu- kherjee & Gupta. 1965. p. 145). Not mapped. Kheiber. See Khyber Pass vicinity. Kheo Ting-Ta Ke. See Tat Ke vicinity. Khen Sahd (= Kheri Saad). Rohtak District: Har- yana. INDIA; ca. 28"56'N. 76°34'E; observed July-Oct. 1979 by R. Singh (1984. p. 50: Singh et al.. 1984. p. 264). A:I-^37. Kherwada Forest. Surat District: Gujarat. INDIA: ca. 21=20'N. 73=30E: observed 1972-1973 by N. Koyama and R B. Shekar (1981. p. 248). A: 1-98. Khirganga. See Pulga. Khyber Pass vicinity: North-West Frontier. PAKI- STAN: ca. 34=05 'N. 7nO'E: reported ca. 1525 by Z. M. Babur ( 1921 [translation], p. 218). A: P-4. Kiang-ka: Sichuan. CHINA: ca. 30°00'N. 99°00'E: reported 31 Aug. 1877 by local residents (Gill, 1883. p. 230). B:C-32. Kiangsu. See Giiangxi. Kian Tatie. See Ngamda. Kia-ting. See Leshan. Kien Thiet vicinity. Yen Son District: Tuyen Quang. VIETNAM: ca. 21°58'N. 105"22'E: re- poned Feb.-Apr. 1992 by R. Ratajszczak. Ngoc Can. and Pham Nhat (1992. p. 20). C:V-16. Kin. west bank of lower Chindwin River: 5a- gaing. MYANMAR (= BURMA): 22=46'N. 94=42'E: collected 9 June I9I4 by G. C. Short- ridge and S. A. Macmillan (Shonridge in Wroughton. 1916a. p. 294): bnhs. 1. B:M-24. Kindat: Sagaing. MYANMAR (= BURMA); 23=44'N. 94=26'E; collected before 1911 by C. H. Hobart: bm(nh). 1. (skull only). B:M-18. Kindat. 20 mi ( = 32 km) northwest of. Chin Hills. 600 ft (= 180 m): Chin. MYANMAR ( = BUR- MA): ca. 23°50'N. 94"10'E: collected 20 Jan. 1915 by J. M. Mackenzie (Wroughton. 1916c. p. 759): BM(NH). 1. B:M-16. King's Forest. See Gokama. Kintachie. See Ngamda. Kistawar. See Dunwein. Kloster Nam miu. See Luofu Shan. Koditan. Hongshui He. right bank. Tian"e Xian. 260 m: Guangxi. CHINA: 25^05 'N. 1 06^59 'E; observed 28 Oct. 1992 by J. Fooden (cf. Food- en et al.. 1994. p. 623). C:C-173. Koh-e-Sefid. See Khyber Pass vicinity. Kokara. See Kakara. Kokkoaing (= Kokhoanig). 500 ft (= 150 m); Mandalay. MYANM.\R ( = BURMA): 20^47 'N, 95=56'E: collected 14 June 1937 by H. C. Smith (cf. Moore & Tate. 1965. p. 323: Napier. 1981, p. 21): BM(NH). 1. B:M-28. Keoladeo Ghana National Park. See Bharatpur. Komlancha (Komallancha). Banswada Taluk. Ni- zamabad District: Andhra Pradesh. INDIA; 18'16'N. 77°55'E: reported Feb. 1977-July 1980 by G. U. Kurup (1984. p. 58; 1992. p. 18). A:I-117. Konapur. Banswada Taluk. Nizamabad District; Andhra Pradesh. INDIA; 18=23N. 77=59 E: re- ported Feb. 1977-July 1980 by G. U. Kurup (1984. p. 58: 1992. p. 18). A:I-117. Kondapalle. Krishna District. 70 m: Andhra Pra- desh. INDIA: 16"37'N. 80=33 E: observed 13 FOODEN: SYSTEMATIC RE\ lEW OF THE RHESUS MACAQUE. MACACA MULATTA 147 May 1980 by J. Fooden, A. Mahabal. and S. S. Saha (1981, p. 468). A:I-144. Kondegattu Temple, Mallial Taluk. Karimnagar District; Andhra Pradesh, INDIA; ca. 18°42'N, 78°58'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-1 19. Kondra Mutla (= Kondramadla), Ipur Taluk, Gun- tur District, 125 m; Andhra Pradesh, INDIA; 16°08'N, 79°46'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). Observed 30 Apr 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 469). A:l- 147. Kongbo; Xizang (= Tibet), CHINA; ca. 29°30'N, 94°45'E; captive observed at Lu, 19 Aug. 1913, by F M. Bailey (1914, map; 1957, p. 171; cf. Fooden, 1982a, p. 52). B:C-10. Kootai. See Kaotai. Kosi River, left bank; Saptari, NEPAL; ca. 26°35'N, 86°55'E; collected Mar. 1887 by H. d'Orleans (1889, pp. 225, 379); museum un- known (not seen). B:N-3. Kota; Rajasthan, INDIA; 25°irN, 75°50'E; re- ported before 1965 by I. Prakash (letter, 25 Aug. 1964). A:I-81. Kotanemalipuri, Guntur District, 100 m; Andhra Pradesh, INDIA; 16°28'N, 79°56'E; observed 9 May 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 468). A:I-145. Kotgay (= Cotgai), east of; Nangarhar, AF- GHANISTAN; ca. 34°00'N, 70°20'E; reported before 1972 by A. Puget (1971, p. 201) A:A- 10. Kotgay (= Cotgai), northeast of; Nangarhar, AF- GHANISTAN; ca. 34°04'N, 70°00'E; reported before 1972 by A. Puget (1971, p. 201). A:A-9. Kothagudem, Khammam District, 100 m; Andhra Pradesh, INDIA; 17°32'N, 80°38'E; observed 20 Apr 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A:I-135. Kothagudem, 4 km north of, Khammam District, 100 m; Andhra Pradesh, INDIA; 17°34'N, 80°38'E; observed 20 Apr 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A: 1-135. Kotihar, 7000 ft (= 2100 m); Jammu & Kashmir, INDIA ca. 33°45'N, 75°10'E; collected 3 and 18 Oct. 1910 by W. L. Abbott; usnm, 2. A:I-3. Kouchlaus, south of; Konarha, AFGHANISTAN; ca. 35°20'N, 7n5'E; reported before 1972 by A. Puget (1971, p. 201). A:A-1. Kounar See Landay Sind, left bank, near conflu- ence with Kunar Riven Kowloon; Xianggang {= Hong Kong), CHINA; ca. 22°15'-22°30'N, 1 14°10'-1 14°25'E; report- ed in 1992 by J. R. Fellowes (Southwick & M. F Siddiqi, 1994b, p. 52). Not mapped. Kowloon Reservoir vicinity. See Eagle's Nest Trail; Kam Shan Entrance. Kuatun; Fujian, CHINA; ca. 27°5rN, 117°48'E, collected 15 and 28 Nov. 1873 by A. David (1875, vol. 2, pp. 260, 281, 288); mnhn, 2 (in- cluding 1 skin only). Collected before 1898 by C. B. Rickett (Rickett & de La Touche, 1896, p. 489; Napier, 1981, p. 22; Fooden, 1983, p. 16); BM(NH), 1 (skull only). Collected 12 May 1898 by J. de la Touhe (in Thomas, 1899, p. 769; Fooden, 1983, p. 16); bm(nh), 1. Collected Nov. 1898 by C. B. Rickett (Napier, 1981, p. 22; Fooden, 1983, p. 16); bm(nh), 1 (holotype of Pithecus littoralis). Collected 16 May-24 June 1926 by E T Smith (1926, p. 131; Sow- erby, 1929, p. 315); museum unknown (not seen). C:C-76. Kuchipudi, Tenali Taluk, Guntur District; Andhra Pradesh, INDIA; 16°10'N, 80°40'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). A:I-143. Kucun, Longquan Xian; Zhejiang, CHINA; ca. 28°05'N, 119°07'E; captives acquired 1960- 1970 by Fu Yiyuan and Wu Fuhai, Hangzhou Zoo (pers. comm., 25 Oct. 1985). C:C-67. Kufri, ca. 0.5 km southeast of; Himachal Pradesh, INDIA; ca. 31°06'N, 77°17'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984, p. 474). A: M8. Kufri, ca. 1.5 km southeast of; Himachal Pradesh, INDIA; ca. 31°05'N, 77°18'E; reported Aug. 1972-Feb. 1973 by K. Wada (1984, p. 474). A: M8. Kufri, ca. 3 km southeast of; Himachal Pradesh, INDIA; ca. 31°04'N, 77°18'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984, p. 474). A: 1-18. Kullu Valley. See Kulu Valley. Kulsi [River], South Kamrup, 750 ft (= 230 m); Assam, INDIA; ca. 26°00'N, 91°23'E; collected 25 Aug. 1920 by H. W. Wells (Hinton & Lind- say, 1926, p. 385); bnhs, 1. B:I-15. Kulu District; Himachal Pradesh, INDIA; 31°58'N, 77°06'E; observed before 1982 by M. L. Roonwal and P C. Tak (1981, p. 96). A:I- 15. Kulu Valley, 1150-2500 m; Himachal Pradesh, INDIA; ca. 32°00'N, 77°10'E; reported before 1991 by O. R Lai (1990, p. 123). A:I-15. Kumaun (= Kumaon) Hills; Uttar Pradesh, IN- DIA; ca. 29°20'N, 79°30'E; observed before 148 FIELDIANA: ZOOLOGY 1982 by M. L. Roonwal and P. C. Tak (1981, p. 96). A:l-32. Kumpawapi Park, 170 m; Udon Thani, THAI- LAND; 17°07'N, 103°02'E; observed July 1989 and Jan. 1991 by N. Aggimarangsee (1992, p. 118; pers. comm., Oct. 1993). C:T-1. Kumtatchie. See Ngamda. Kunar River, lower; North-West Frontier, PAKI- STAN ca. 35°25'N, 71°40'E; reported before 1902 by A. H. McMahon (1901a, p. 4). A:P-1. Kuo-Lo. See Golog Zangzu Zizhuzhou. Kurukshetra District; Haryana, INDIA; ca. 29°59'N, 76°51'E; observed 1981-1983 by P K. Seth, S. Seth, G. L. Reddy, and P K. Chopra (1992, p. 62). A:I-24. Kyirong. See Gyirong. Ky Son, Ky Anh District; Ha Tinh, VIETNAM; 17°57'N. 106°06'E; collected 26 Jan. 1964 by Nguyen Lien (Dao, 1985, p. 233, 242, misiden- tified as M. assamensis; Dang et al., 1994, p. 165; cf. Nisbitt & Ciochon, 1993, p. 772); iebr, 1 (skin only). C:V-29. Lagou Bird Reserve; Guangxi, CHINA; ca. 24°38'N, 110°04'E; observed 1976, 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter Aug. 1996). C:C-194. Lai Chau, VIETNAM; 20°50'-22°50'N, 102°10'- 103°50'E; collected in 1963 by unknown col- lector; IEBR, 1 (skin only). Not mapped. Lakhimpur. See Margherita. Lakuwa, 980-2300 m; Dhankuta, NEPAL; 27°28'N, 87°10'E; observed in 1997 and 1999 by M. K. Chalise and M. Ghimire (1998, p. 12). B:N-supplementary. Lai Kuan; Delhi, INDIA; ca. 28°30'N, 77°20'E; reported Feb. 1990 by I. Malik and R. L. John- son (1994, p. 237). A:I-41. Lamsakhang, Cachar District, 200 ft (= 60 m); Assam, INDIA; 25°48'N, 93°06'E; collected 10 Sept. 1920 by H. W. Wells (Hinton & Lindsay, 1926, p. 385); bm(nh), 1. 3:1-20. Lancang Jiang (= Mekong River), Lincang Dis- trict; Yiimtan, CHINA; ca. 24°00'N, 100°23'E; reported before 1996 by Lan Daoying and Guo Guang (1995, p. 6). B:C-70. Lancang Xian; Yunnan, CHINA; ca. 22°32'N, 99°56'E; immunological survey conducted be- fore 1996 by Duan Xingsheng, Liu Yuanwei, Wu Jing, Dao Weiying, and Liu Jianghai (1995, p. 411). B:C-79. Landay Sind, left bank, near confluence with Ku- nar River; Konarha, AFGHANISTAN; 35°2rN, 71°33'E; observed before 1972 by A. Puget (1971, p. 200). A:A-I. Landay Sind, left bank, near Pule Saret; Konarha, AFGHANISTAN; 35°22'N, 71°33'E; observed before 1972 by A. Puget (1971, p. 200). A:A-1. Landay Sind, near Chascoup; Konarha, AF- GHANISTAN; 35°3rN, 7r22'E; observed be- fore 1972 by A. Puget (1971. p. 199). A:A-1. Landay Sind, near Merdech; Konarha, AFGHAN- ISTAN; 35°23'N, 7I°32'E; observed before 1972 by A. Puget (1971, p. 200). A:A-1. Landay Sind, right bank, between Mandagal and Ormul; Konarha, AFGHANISTAN; 35°28'N, 7r20'E; observed before 1972 by A. Puget (1971, p. 199). A:A-1. Landay Sind, right bank, near Sang e Safed; Kon- arha, AFGHANISTAN; 35°22'N, 71°32'E, ob- served before 1972 by A. Puget ( 1971, p. 200). A:A-1. Landay Sind Valley, near Kamu Valley; Konarha, AFGHANISTAN; ca. 35°25'N, 7r25'E; ob- served July 1970 by C. Naumann and G. Nogge (1973, p. 92). A:A-1. Landay Sind Valley, southern slope; Konarha, AFGHANISTAN; ca. 35°25'N, 71°25'E; ob- served 26 Dec. 1971 by C. Naumann and G. Nogge (1973, p. 92). A:A-1. Landrai Valley, northern Dir District; North-West Frontier, PAKISTAN; ca. 35°30'N, 72°00'E; reported before 1978 by T J. Roberts (1977, p. 86). A:P-2. Lang Son, VIETNAM; 21°25'-22°50'N, 106°05'- 107°20'E; collected 1962 by unknown collec- tor; ZMVNU, 1 (skull only). C:V-12. Lan-tao. See Dahao Dao. Laodian, Lin'an Xian, 1000 m; Zhejiang, CHINA; ca. 30°20'N, 119°25'E; observed Aug. 1983 by Zhang Minhua, zmnh (pers. comm., 24 Oct. 1985). C:C-57. Laoshan, Jinxiu Subcounty, Jinxiu Xian; Guangxi, CHINA; ca. 24°07'N. 110°12'E; captured Sept. 1992 by Shu Jenyung (pers. comm., 16 Nov. 1992); captive observed 16 Nov. 1992 at Jinxiu. C:C-195. Laxmidevipeta (= Laxmideirpet), Mulug Taluk, Warangal District; Andhra Pradesh, INDIA; 18°18'N, 79°58'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-120. Ledhan hla. See Lethan Hka. Leibo; Sichuan, CHINA; 28°15'N, 103''34'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-142. Lema Islands. See Dangan Dao. Lena Island. See Dangan Dao. Leshan (= Kia-ting), mountains 30 mi southwest FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 149 of, 2900 ft (= 880 m); Sichuan, CHINA; ca. 29'28'N. 103'18'E; collected 6 Feb. 1911 by M. P. Anderson; bm(nh). 1. C:C-139. Lethan Hka, Maymyo E D.. 300 ft (= 90 m); Mandalay, MYANMAR (= BURMA); ca. 22°00'N. 96°30'E; collected 22 Dec. 1935 by H. C. Smith; bm(nh). 1. B:M-21. Le Thuy. See Xuan Ninh. Lhasa. See CHINA. Lhunze Xian; Xizang (= Tibet), CHINA; ca. 28^30'N. 92"25'E; observed 1979-1982 by Zhang Cizu. Director, Shanghai Zoo (pers. comm.. 18 Oct. 1985). B:C-4. Liancheng Xian; Fujiaiu CHINA; ca. 25°47'N, 116'48T; reported Aug. 1982 by Zheng Xue- qing (1984. p. 146). C:C-90. Liangdang; Gansii. CHINA; 33'56'N, 106n2'E; reported before 1998 (Zhang et al., 1997. p. 58). C:C-I6. Liangping; Guangdong. CHINA; 24'22'N, 114=30'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-96. Lian Xian. See Lianzhou. Lianzhou, Lian Xian; Guangdong, CHINA; 24=48'N, 112"25'E; reported before 1998 (Zhang et al., 1997. p. 58). C:C-201. Libo; Guizhou, CHINA; 25°25'N, 107=53'E; re- ported before 1998 (Zhang et al.. 1997, p. 58). C:C-175. Lijiang; Yunnan. CHINA: 26=48'N, 100M6'E; re- ported before 1998 (Zhang et al., 1997, p. 58). B:C-45. Lina. See Dangan Dao. Lincang Prefecture; Yunnan. CHINA; ca. 23°54'N, 100'02'E; immunological survey con- ducted before 1996 by Duan Xingsheng. Liu Yuanwei, Wu Jing, Dao Weiying, and Liu Jian- ghai (1995. p. 411). B:C-69. Lingchuan. southeast of, 900-1400 m: Shanxi. CHINA; ca. 35'38'N. 113=28'E; observed 1985-1994 by Zhu Jun, Zhao Yishan, and Fan Longsuo (1995, p. 134). C:C-7. Lingtin Island. See Neilingding Dao. Lin^un. See Lon^un. Lingrui. See Longrui. Linh Thong, Dinh Hoa; Bac Thai, VIETNAM; 22°00'N. 105°42'E; collected 22 June 1967 by Truong Van La; zmvnl. 2 (skins only). C:V-10. Lintin Dao. See Neilingding Dao. Linwan Shan Water Regulation Forest Reserve; Guangxi. CHINA; 22'25'N, 109"55'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter, Aug. 1996). C:C-215. Li Shan National Nature Reserve. 700-1500 m; Shanxi, CHINA; ca. 35'22'N, lir59'E; ob- served 1985-1994 by Zhu Jun. Zhao Yishan, and Fan Longsuo (1995, p. 134). C:C-3. Litang-Batang; Sichuan. CHINA; ca. 30"00'N, 100'00'E; reported 23 Aug. 1877 by local res- idents (Gill 1883. p. 212). B:C-31. Liukou, Qimen Xian. 300 m; Anhui. CHINA; ca. 29°55'N, 117=30'E; observed 1973-1986 by Xiong Chenpei (Wada et al., 1986, p. 83). C:C- 48. Liukou, Xiuning Xian. 400 m; Anhui. CHINA; 29'34'N. 117^'49'E; observed 1973-1986 by Xiong Chenpei (Wada et al.. 1986, p. 83). C:C- 63. Liulipenshan; Hebei CHINA; ca. 40°24'N, 117'^30'E; reponed fall 1987 by local hunters (Zhang et al.. 1989. p. 380). C:C-1. Liulipenshan. southern slope; Hebei CHINA; ca. 40'24'N. 117'30E: reported 1985-1986 by lo- cal resident Wang Chaori (Zhang et al.. 1989. p. 379). C:C-1. Liuzhai vicinity. Nandan Xian; Guangxi. CHINA; ca. 25°18'N. 107"24'E; collected June 1992 by local farmer (Lu Gwangyan. manager of local products station, pers. comm.. 27 Oct. 1992); skeleton examined 27 Oct. 1992 at Liuzhai. C: C-174. Lofau. See Luofu Shan. Lolab. 7500 ft (= 2300 m); Jammu & Kashmir. INDIA; ca. 34=30'N. 74=35 'E; collected 8-9 Sept. 1891 by W. L. Abbott (True. 1894. p. 3; Blanford. 1898. p. 361): lsnm. 5 including ho- lotype of Macacus rhesus villosus). A:I-1. Lolab Valley; Jammu & Kashmir. INDIA; ca. 34=30'N, 74=35 'E; collected 1 1 Feb. 191 1 by W L. Abbott; lsnm, 1. A:I-1. Longan. Xiangshui District. Longzhou Xian; Guangxi. CHINA; ca. 22=24'N. 107=10'E; col- lected Dec. 1980 by Wu Mingchuan (pers. comm., 27 Nov. 1992); fdcg, 1 (skin with skull mside). C:C-226. Longchi. Dahe District. Xixiang Xian. 1000-1500 m- Shaanxi. CHINA; 32=42'''N. 1 07=28 'E: ob- served June-July 1966 by Chen Fugan. North- west University. Xi'an (pers. comm.. 14 Oct. 1985). C:C-35. Longgang Nature Reserve; Guangxi. CHINA; ca. 22'23'N. 106=53'E; observed 1976. 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 126; letter. Aug. 1996). C:C-227. Longhua Water Regulation Forest Reserve; Guangxi. CHINA; ca. 23=17'N. 105=34'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and 150 FIELDL\NA: ZOOLOGY Wei Zhenyi (1995, p. 128; letter. Aug. 1996). C:C-164. Longhu Shan Nature Reserve. 490 m; Guang.xi, CHINA: 22"42'N. 107°30'E; observed 1976. 1986. 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 127; letter, Aug. 1996). Observed 1988-1990 by Feng Min, Jiang Haisheng. and Wang Jun (1997. p. 27). C:C-225. Longjun (= Lingjun) Hsienmu Reserve; Guang.xi. CHINA; ca. 23°14'N. 107°54'E; observed 1976. 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter, Aug. 1996). C:C-222. Longrui (= Lingrui) Nature Reserve; Guang.xi, CHINA; ca. 22°28'N. 107°12'E; observed 1976. 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 126; letter, Aug. 1996). C:C-226. Loshan. See Leshan. Longsheng; Guang.xi, CHINA; 25°48'N. 1 10°00'E; reported before 1998 (Zhang et al.. 1997, p. 58). C:C-185. Longxi (prefecture); Fujian, CHINA; ca. 24°3rN, 117°40'E; reported before 1998 (Zhang et al.. 1997, p. 58). C:C-93. Longyan Xian; Fujian, CHINA; ca. 25°11'N, 117°00'E; reported Oct. 1982 by Zheng Xue- qing (1984, p. 146). C:C-92. Louangphrabang, downstream; Louangphrabang, LAOS; ca. 19°52'N, 102°08'E; reported before 1964 by J. Deuve and M. Deuve (1963, p. 59). B:L-3. Lower Bazar. See Simla. Lu. See Kongbo. Luang-Prabang. See Louangphrabang. Luchun; Yunnan, CHINA; 23°19'N. 102°10'E; re- ported before 1998 (Zhang et al.. 1997, p. 58). B:C-76. Lucknow vicinity; Uttar Pradesh, INDIA; ca. 26°5rN, 80°55'E; captured in 1938 by C. R. Carpenter (Rawlins & Kessler, 1986b, pp. 17, 21). Observed Sept. 1959-Feb. 1960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961a, p. 543). Blood samples obtained 16-27 Apr. 1964 by K. V. Shah and C. H. Southwick (1965, p. 489). Reported ca. 1971 by D. Wein- man (1974, p. 345). A:I-66. Lucknow-Faizabad, highway between; Uttar Pra- desh, INDIA; ca. 26°50'N, 81°30'E; observed Sept. 1959-June 1960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961b, p. 702). Ob- served 24-25 Oct. 1964 by C. H. Southwick, R. K. Lahiri. M. Bertrand. D. Lindburg, and R Jay (Southwick & M. R. Siddiqi, 1966, p. 306). A:I-65. Lucknow -Sitapur, highway between; Uttar Pra- desh, INDIA; ca. 27°10'N, 80°50'E; observed 21-29 Oct. 1964 by C. H. Southwick. D. Lind- burg. M. Neville, R Jay, M. R. Siddiqi, and R. R Mukherjee (Southwick & M. R. Siddiqi, 1966, p. 306). Observed 1964-65 by R. R Mu- kherjee and G. D. Mukherjee (1972, p. 67). A: 1-58. Luia, Chaibasa vicinity, Singhbhum District, 1000 ft (= 300 m); Bihar, INDIA; 22°23'N, 85°32'E; collected 1 Aug. 1914 by C. A. Crump (in Wroughton, 1915b, p. 99); bm (nh), 3. A-I:105. Lungli vicinity, southeast of Liuzhai, Nandan Xian; Guang.xi, CHINA; ca. 25°17'N, 1()7°25'E; collected Dec. 1991 by He Kean (pers. conim., 27 Oct. 1992); skin and skeleton examined 27 Oct. 1992 at Lungh. C:C-174 Lunpuriagaon, Assam, INDIA; ca. 26°57'N, 94°47'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. Luocheng; Guang.xi, CHINA; 24°47'N, 108°54'E; reported before 1998 (Zhang et al., 1997. p. 58). C:C-179. Luofu Shan. 800-1100 m; Guangdong, CHINA; ca. 23°17'N, 114°03'E; reported 1908-1921 by R. Mell (1922, pp. 10, 11). C:C-208. Luquan; Yunnan, CHINA; 25°35'N, 102°30'E; re- ported before 1998 (Zhang et al., 1997, p. 58). B:C-50. Luyuan, Taihe Xian; Jiangxi, CHINA; ca. 26°50'N, 1 14°40'E; reported Oct. 1979 by local residents (Liu Zhenhe, scika, pers. comm., 25 Nov. 1985). C:C-104. Ly Bon, Bao Lac District; Cao Bang, VIETNAM; ca. 22°57'N, 105°4rE; collected 5 June 1965 by unknown collector (Dao, 1985, p. 38); lEBR, 4 (3 skins only, 1 skull only [possibly belongs with one of the skins]). C:V-7. Machayara Game Reserve; Azad Kashmir, PAKI- STAN; ca. 34°00'N, 73°35'E; reported before 1984 by M. Nawaz (1983, p. 6). A:P-12. Madaripur Township; Madaripur, BANGLA- DESH; ca. 23°10'N. 90°12'E; reported before 1986 by M. A. R. Khan (1981, p. 13; 1985. p. 31). B:Ba-25. Madaya. Maymyo Reserve; Mandalay, MYAN- MAR (= BURMA); 22°13'N, 96°07'E; collect- ed 15 Feb. 1936 by R F Garthwaite; bm (nh), 1 (skin only). B:M-22. Madhupur, ca. 100 km west of; Natore, BANG- LADESH; ca. 24°30'N, 89°00'E; tentatively re- ported July-Nov. 1976 by K. M. Green (1978, p. 146). B:Ba-5. Madhupur National Park; Tangail, BANGLA- DESH; ca. 24°45'N, 90°08'E; observed July- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MAC AC A MULATTA 151 Nov. 1976 by K. M. Green (1978, p. 154). Ob- served Dec. 1977-July 1978 by M. A. Islam and K. Z. Husain (1982. p. 157). Observed ear- ly in 1980 by S. P. Gittins and A. W. Akonda (1982, p. 278). Observed before 1982 by M. A. R. Khan (1981, p. 13). Observed Feb. 1990- June 1993 by M. M. Feeroz. M. A. Islam, and M. Kabir (1995. p. 76). B:Ba-7. Madhpur National Park, southern portion; Tan- gciil BANGLADESH; ca. 24°30'N, 90°10'E; observed Sept.-Dec. 1986 and Dec. 1987-Dec. 1988 by C. B. Stanford (1991. p. 17; 1992. p. 188). B:Ba-8. Magi, Banswada Taluk. Nizamabad District; An- dhra Pradesh. INDIA; 18°12'N, 77°55'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-117. Mahabali Temple. See Imphal. Mahabali Temple. Mahal, 2-5 km northwest of. Dangs District. 200- 275 m. Gujarat, INDIA; 20°56'N, 73°37'E; ob- served 12 Mar. 1980 by J. Fooden, A. Mahabal. and S. S. Saha (1981, p. 466). A:I-99. Maiskhal Island; Cox's Bazar. BANGLADESH; ca. 21°35'N, 91°55'E; reported early in 1980 by S. P Gittins and A. W. Akonda (1982, p. 278). B:Ba-38. Maizhokunggar Xian; Xizhang (= Tibet), CHI- NA; ca. 29°50'N, 91°45'E; observed 1979- 1982 by Zhang Cizu, Director, Shanghai Zoo (pers. comm.. 18 Oct. 1985). B:C-2. Makalu-Barun Conservation Area. See Sankhuwa Khola. Makehe Plantation. 3100-4000 m; Qinghai. CHI- NA; ca. 33°00'N, 96°20'E; purchased ca. 1982 at Baima Xian by Liao Yianfa. Director of Xin- ing Zoo (pers. comm., 6 Oct. 1985); captive observed 6 Oct. 1985. B:C-16. Makhena, near Anupshahr; Uttar Pradesh, IN- DIA; ca. 28°22'N, 78°16'E; reported ca. 1980 by C. H. Southwick and M. F Siddiqi (1984, p. 559). A:I-49. Makkimarigudem (= Makkinavarigudem), Chin- talapudi Taluk. West Godavari District; Andhra Pradesh, INDIA; 17°09'N. 81°05'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984. p. 58; 1992. p. 19). A:I-137. Malakand. See Bar Chanrai Hill. Malipura. See Khair, Tahsil. Malkangiri (= Malkanagiri). Jeypore Agency, Vi- sakhapatnam vicinity; Orissa, INDIA; 18°2rN. 81°54'E; collected 28 Aug. 1927 by A. V. Sun- daram and A. H. Bishop; bm(nh), 1 (skin only). A:I-110. Mall. See Simla. Mallur, Banswada Taluk, Nizamabad District; An- dhra Pradesh, INDIA; 18°16'N, 77°53'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-117. Malua. Seoni Tank; Madhya Pradesh, INDIA; ca. 2r00'N, 79°59'E; collected before 1974 by S. W. Prater; bnhs, 1 (skin only). A:I-103. Maluling; Xizang (= Tibet), CHINA; ca. 32°40'N, 97°20'E; reported before 1998 (Zhang et al., 1997, p. 58). B:C-18. Mamsam Falls. See Mansam Falls. Manali; Himachal Pradesh, INDIA; ca. 32°20'N, 77°05'E; reported 1978-1980 by A. J. Gaston. P J. Garson, and M. L. Hunter, Jr. (1983, p 300). A:I-13. Manas. See Royal Manas National Park. Manas National Park; Assam, INDIA; ca. 26°40'N, 90°55'E; reported before 1997 by K. K. Gurung and R. Singh (1996, p. 104). B:I- supplementary. Mandagal, northwest of; Konarha, AFGHANI- STAN; ca. 35°35'N, 71°15'E; reported before 1972 by A. Puget (1971, p. 201). A:A-1. Mandal. See Kedarnath Sanctuary. Manghe Nature Reserve, 680-1000 m; Shanxi, CHINA; ca. 35°15'N, 112°27'E; observed 1985-1994 by Zhu Jun, Zhao Yishan, and Fan Longsuo (1995, p. 134). C:C-4. Mangpu, 3000 ft and 3500 ft (= 910 m and 1070 m); West Bengal, INDIA; 26°58'N, 88°24'E; collected 5 and 12 Dec. 1930 by H. Stevens; FMNH, 2. B:I-6. Manipompla. See Munipamula. Manipuri Tea Estate; Sylhet, BANGLADESH; not precisely located. 24°08'-24°50'N. 91°37'- 92°17'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). Not mapped. Manpa, Mengla Xian, 650-750 m; Yunnan, CHI- NA; ca. 2r40'N, 101°37'E; collected 14 May 1959 by Deng Xiangfu; kiz. 2 (skulls only). Collected 19 Jan. and 8 May 1962 by Quan Guoqiang (pers. comm.. 25 Aug. 1983); izcas, 2. B:C-85. Mansam Falls. Nam Yao (river). 2000 ft (= 610 m); Shan, MYANMAR (= BURMA); 22°48'N, 97°32'E; collected 6 June 1913 by G. C. Short- ridge (in Ryley. 1914, p. 713); bm(nh), 2 (in- cluding 1 skin only); bnhs, 3. B:M-19. Mansar Patwar. See Surinsar. Mao'er Shan Nature Reserve; Guangxi, CHINA; ca. 25°53'N, 110°28'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 126; letter, Aug. 1996). C:C-189. 152 FIELD! AN A: ZOOLOGY Maojie Bird Reserve; Guangxi. CHINA; ca. 24°28'N, 104°35'E; observed 1976. 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 129; letter, Aug. 1996). C:C-154. Maowen; Sichuan. CHINA; 31°41'N. 103°52'E; tentatively reported 1914-1916 by H. Weigold (1924. p. 71). C:C-25. Marco River. See Golog Zangzu Zishizhou. Mardan. See Pajja Hill. Margalla Hills National Park; Federal Capital Territory. PAKISTAN; 33°48'N. 73°10'E; re- ported ca. 1975 by S. J. Goldstein and A. F. Richard (1989, p. 563). Reported before 1978 by T. J. Roberts (1977. p. 87). Reported before 1984 by M. Nawaz (1983. p. 2). A:P-11. Margherita, Lakhimpur District. 200 ft ( = 60 m); Anmachal Pradesh, INDIA; 27°17'N. 95°4rE; collected 14 Nov. 1919 by H. W. Wells (Hinton & Lindsay. 1926. p. 385); zsi. 1 (skin only). B:I-29. Marot (= Maroth), Nagaur District; Rajasthan, INDIA; 27°05'N, 75°05'E; observed June 1971-June 1972 by R R. Ojha (1974, p. 163). Observed 1975-1980 by R K. Seth and S. Seth (1983, p. 63). A:I-77. Mar Qu. See Golog Zangzu Zishizhou. Mat, Phu; Nghe An, VIETNAM; 18°38'-20°00'N, 103°53'-105°12'E; observed ca. 1990-1995 by L. K. Lippold (1995, p. 198; cf. Cao, 1995, p. 182). Not mapped. Mathura; Uttar Pradesh, INDIA; 27°30'N, 77°41'E; observed Sept. 1959-Feb. 1960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961a, p. 543). Observed 1964-1965 by C. H. Southwick and M. R. Siddiqi 1966, p. 309). A: 1-71. Mathura District (= Muttra); Uttar Pradesh, IN- DIA; 27°15'-28°00'N, 77°15'-77°50'E; labora- tory animals obtained in 1891 by W. Heape (1897, p. 135). Not mapped. Matlab Bazar; Chandpur, BANGLADESH; 23°20'N, 90°43'E; reported before 1986 by M. A. R. Khan (1981, p. 13; 1985. p. 31). B:Ba- 28. Matlab, southeast of; Noakhali BANGLADESH; ca. 23°00'N, 91°00'E; tentatively reported July- Nov. 1976 by K. M. Green (1978, p. 146). B.Ba-30. Maungkan, east bank of Chindwin River; Sa- gaing, MYANMAR (= BURMA); 25°05'N, 95°02'E; collected 20 Mar. 1935 by H. C. Ra- ven and R. C. Morris (Raven in Carter, 1943, p. 100; Morris, 1936, p. 667); amnh, 1. B:M- 13. Maure, near, 1.6 km north of Jamduar; Assam, INDIA; ca. 26°44'N, 89°53'E; observed 19 Nov.-l Dec. 1959 by E. P Gee (1961, p. 6). B: MO. Mautschou. See Maowen. Maymo F. D. See Lethan Hka. Maymyo (= Maymo). 800 m; Mandalay, MYAN- MAR (= BURMA); 22°02' N, 96°28'E; col- lected 3 Dec. 1937 by G. Heinrich; amnh, 1. B: M-21. Maymyo Reserve. See Madaya. Medog Xian; Xizang {= Tibet), CHINA; ca. 29°15'N, 95°15'E; observed 1979-1982 by Zhang Cizu, Director, Shanghai Zoo (pers. comm. 18 Oct. 1985). B:C-13. Meerut District; Uttar Pradesh. INDIA; ca. 28°55'N, 77°4rE; observed 1981-1983 by P K. Seth, S. Seth, G. L. Reddy. and P K. Chopra (1992, p. 62), A:I-35. Meetha Pur, ca. 10 km from Tughlaqabad; Delhi, INDIA; ca. 28°30'N, 77°15'E; reported Aug. 1989 by I. Malik and R. L. Johnson (1991, p. 63; 1994, p. 237). A:I-41. Meherpur; Meherpur, BANGLADESH; 23°46'N, 88°38'E; reported before 1982 by M. A. R. Khan (1981, p. 13). B:Ba-17. Meigu; Sichuan, CHINA; 28°20'N, 103°04'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-141. Meihua Shan (= Meihaushan); Fujian, CHINA; ca. 25°15'N, 116°45'E; reported Nov. 1983 by Zheng Xueqing (1984, p. 145). Reported before 1986 by Tan Ban^ie (1985, P 73). C: C-9I. Meitan, near; Guizhou, CHINA; ca. 27°40'N, 107°30'E; captive purchased 7 Nov. 1960 by Quan Guoqiang (pers. comm. 25 Aug. 1983); izcAS, 1 C:C-128. Mekong River, 90 km above Viangchan; Vien- tiane, LAOS; 18°05'N, 10r57'E; collected 7 July 1924 by F R. Wulsin (field catalog and map, usNM archives); usnm, 1. B:L-5. Mellavagu (= Melavagu), Ipur Taluk, Guntur Dis- trict; Andhra Pradesh, INDIA; 16°20'N, 79°45'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). A:I-147. Meng-ban; Yunnan, CHINA; ca. 21°45'N, 100°10'E; collected 30 May 1958 by Ye Zong- yao (Bannikov. 1958. p. 68; Kao et al., 1962, p. 188; Quan Guoqiang, pers. comm., 25 Aug. 1983). IZCAS, 1. B:C-80. Menghai; Yunnan, CHINA; 21°58'N, 100°28'E; collected 27 Nov. 1957 and 17 Apr. 1958 by Ye Zongyao (Bannikov, 1958, p. 68; Kao et al., 1962, p. 188); izcas, 2(1 skin only; 1 skull only FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 153 [external measurements recorded on skull tag]). B:C-81. Menghun; Yunnan, CHINA; ZTSO'N, 100°23'E; reported before 1998 (Zhang et al., 1997, p. 58). B:C-81. Mengla Xian, 640 m and 730 m; Yunnan, CHI- NA; 2r-22°N, 101°-102°E; collected in 1959 by unknown collector; kiz, 3 (2 skins only. 1 skull only). Collected 30 Oct. 1961 by Li Zhi- xiang; kiz. 1. Not mapped. Menglong; Yunnan, CHINA; 21°36'N. 100°40'E; reported befored 1998 (Zhang et al.. 1997, p. 58). B:C-82. Menglun. Mengla Xian, 740 m; Yunnan, CHINA; 21°55'N. lOnS'E; collected 8 May 1959 by Deng Xiangfu (Wang Yingxiang. pers. comm., 29 Aug. 1983); kiz, 1 (skin only). Collected in 1961 by Yang Lan (Wang Yingxiang. pers. comm.. 29 Aug. 1983); kiz. 1 (skin only). B:C- 84. Mengyang; Yunnan, CHINA; ca. 22°00'N, 100°50'E; collected 10 and 12 Apr. 1957 and 18 Oct. 1958 by Ye Zongyao (Bannikov, 1958, p. 68; Kao et al.. 1962. p. 188; Quan Guoqiang. pers. comm. 25 Aug. 1983); izcas, 3. B:C-83. Me Ping rapids. See Kaeng Mae Hat. Mesogarh; Assam, INDIA; ca. 26°58'N, 94°41'E; reported 9 Mar 1987-16 Feb. 1988 by A. Choudhury ([1991a]. p. 31). B:I-25. Meteka; Assam, INDIA; ca. 26°58'N. 94°39'E; observed 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 32). B:I-25. Mg. Khua. See Ou, Nam, between Muang khoua and Muang Ngoy. Mg. Ngoi. See Ou, Nam, between Muang Khoua and Muang Ngoy. Midwest Sichuan. See Sichuan, west-central. Mihouling, Ledong Xian. 700-800 m; Hainan Dao; Hainan, CHINA; 18°55'N, 109°08'E; pur- chased 5 May 1964 by Liu Zhenhe, sciea (pers. comm., 26 Nov. 1985); sciea, 1. C:C-234. Mikir Hills. Assam, INDIA; ca. 26°10'N, 93°30'E; reported Dec. 1972-Feb. 1973 by R. L. Tilson (1983. p. 399). B:I-33. Mile; Yunnan, CHINA; 24°24'N. 103°27'E; re- ported before 1998 (Zhang et al., 1997. p. 58). C:C-148. Minglang; Yunnan, CHINA; 23°53'N, 99°irE; reported before 1998 (Zhang et al., 1997. p. 58). B:C-66. Mingun. near Sagaing, 250 ft (= 70 m); Sagaing, MYANMAR (= BURMA); 22°03'N. 96°0rE; collected 12 July 1913 by G. C. Shortridge (in Wroughton. 1915a. p. 461); bm(nh). 1. B:M-23. Mintal. See Wassuland. Mintal, ob. See Maowen. Mirkhani (= Mirkandi), 4000 ft (= 1200 m); North-West Frontier, PAKSITAN; 35°28'N. 71°44'E; observed before 1902 by Capt. B. E. M. Gurdon (McMahon, 1901a, p. 4). A:P-1. Misajan; Assam, INDIA; ca. 26°52'N, 94°39'E; reported 9 Mar 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. Mishmi Hills. See Dening. Miyi; Sichuan, CHINA; 26°50'N, 102°03'E; re- ported before 1998 (Zhang et al., 1997, p. 58). B:C-48. Miwan, Taishan Xian, Shangchuan Dao, <200 m; Guangdong, CHINA; 21°37'N, 112°45'E; col- lected 19 Apr. 1981 by Liu Zhenhe and Xu Longhuei (Liu Zhenhe, sciea, pers. comm., 26 Nov. 1985); sciea. 1. C:C-213. Moduri; Assam, INDIA; ca. 26°55'N, 94°44'E; re- ported 9 Mar 1987-16 Feb. 1988 by A. Choud- hury ([1991a], p. 31). B:I-25. Moenjo Daro: Sind, PAKISTAN; 27°19'N, 68°07'E; possible occurrence ca. 4000 B.P. in- dicated by prehistoric statuettes (Mackay, 1931, p. 349; Iyer, 1977, p. 15). Not mapped. Mohan (= Mohand); Uttar Pradesh, INDIA; 30°11'N, 77°54'E; reported 1964-1966 by D. G. Lindburg (1977a, p. 268). A:I-27. Mohenjodero. See Moenjo Daro. Mojiang Xian; Yunnan, CHINA; ca. 23°25'N, 101°44'E; immunological survey conducted be- fore 1996 by Duan Xingsheng, Liu Yuanwei, Wu Jing. Dao Weiying, and Liu Jianghai (1995. p. 411). B:C-75. Moklok. east bank of Chindwin River; Sagaing, MYANMAR (= BURMA); ca. 25°37'N. 95°25'E; collected 16 Mar. 1935 by H. C. Ra- ven (in Carter. 1943. p. 100; Morris, 1936. p. 666); AMNH. 1. B:M-10. MoUur See Mallur. Molta; Uttar Pradesh, INDIA; ca. 30°00'N, 79°00'E; observed 1955-1957 by G. A. von May dell (Oboussier & von Maydell. 1960. p. 143). A:I-30. Momien. See Tengchong. Mom Ray (= Mon Ray) Nature Reserve; Kon Turn, VIETNAM; ca. 14°27'N. 107°45'E; re- ported ca. 1990-1995 by L. K. Lippold (1995. p. 200). C:V-38. Mong Moen. See Muong Muon. Mong Moun. See Muong Pon. Monkey Bridge. See Lucknow vicinity. Monkey hill. <10 mi (<16 km) south of Jing- gangshan; Jiangxi, CHINA; ca. 26°30'N, 154 FIELDIANA: ZOOLOGY 114°10'E; reported ca. 1985 by Tan Bangjie (1985. p. 73). Report unverified; M. thibetana only macaque verified at this locality (Li Xiongshan. Jinggangshan Nature Reserve Bu- reau, pers. comm.. 4 Nov. 1985; Liu Zhenhe, sciEA, pers. comm.. 25 Nov. 1985). Not mapped. Mon Ray Nature Reserve. See Mom Ray Nature Reserve. Moradabad vicinity; Uttar Pradesh. INDIA; ca. 28°50'N. 78°47'E; autopsied ca. 1966 by K. K. Chawla. C. D. S. Murthy, R. N. Chakravarti. and P. N. Chhuttani (1967. p. 85). A:I-50. Morang region; Morang, NEPAL; ca. 26°30'N. 87°30'E; reported 1920-1921 by N. A. Baptista (in Hinton & Fry. 1923. p. 403). B:N-4. Morit forest. See Kokkoaing. Moshemien; Sichuan, CHINA; ca. 30°00'N, 102°00'E; reported 24 June-7 July 1929 by lo- cal residents (Stevens. 1934. p. 132). B:C-27. Motianling. Debao Xian; Guang.xi, CHINA; ca. 23°20'N. 106°37'E; collected ca. Oct. 1979 by unknown collector (Quan Guoqiang, pers. comm.. 13 Dec. 1985); fdcg. 1 (mounted skin with skull inside; specimen not seen). C:C-167. Moung Boum. See Muong Bourn. Moung Mouen. See Muong Muon. Moung-moun. See Muong Moun. Mount Everest. See Sagarmatha. Mount Omei. See Emei Shan. M. R. G. High School. See Dhaka. Mt. Wuchi. See Wuzhi Shan. Muang Khoua. See Ou. Nam, between Muang Khoua and Muang Ngoy. Muang Ngoy. See Ou. Nam. between Muang Khoua and Muang Ngoy. Muang Pakxan; Vientiane, LAOS; ca. 18°22'N. 103°39'E; reported before 1964 by J. Deuve and M. Deuve (1963. p. 59). C:L-2. Muang Thateng, Plateau des Bolovens; Saravan, LAOS; 15°26'N. 106°23'E; purchased 29 Jan. and 13 Feb. 1932 by T. D. Carter (Legendre, 1932. p. 495; 1936. p. 251; Fooden. 1997, pp. 227. 229); an.sp. 5. C:L-3. Mudhalaparava, Krishna District 120 m; Andhra Pradesh, INDIA; 16°56'N. 80°45'E; observed 14 May 1980 by J. Fooden. A. Mahabal, and S. S. Saha (1981. p. 468). A:I-140. Mudigonda (= Mudukonda). Khammam Taluk. Khammam District; Andhra Pradesh, INDIA; 17°10'N, 80°04'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984. p. 58; 1992. p. 18). A:I-132. Muh; Sichuan, CHINA; 27°50'N. 101°15'E; re- ported 9 Apr 1929 by H. Stevens (1934. p. 132). Ti.ssue samples obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). Reported before 1992 by Jiang Xuelong, Wang Yingxiang, and Ma Shilai (1991, p. 243). B:C- 39. Mulug (= Mulugu), Gajwel Taluk, Medak Dis- trict; Andhra Pradesh, INDIA; 17°45'N, 78°38'E; reported Feb. 1977-July 1980 by G.U. Kurup (1984, p. 58; 1992, p. 18). A:I-123. Mulun Nature Reserve; Guang.xi, CHINA; ca. 25°08'N, 107°50'E; observed 1976. 1986. and 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 127; letter. Aug. 1996). C:C-177. Mumbai (= Bombay). Raj Bhavan compound; Maharashtra, INDIA; 18°56'N. 72°48'E; pop- ulation atrificially introduced (Serrao & Amla- di, 1979, pp. 29, 32). Observed Apr-May 1973 by J. Fooden. Not mapped. Municipal Corporation Builiding. See Aligarh. Munipamula, Ramannapet Taluk. Nalgonda Dis- trict; Andhra Pradesh, INDIA; 17°19'N, 79°09'E; reported Feb. 1977-July 1980 by G.U. Kurup (1984. p. 58; 1992. p. 18). A:I-128. Muong Boum. Tonkin region; Lai Chau, VIET- NAM; 22°23'N, 102°49'E; collected 27 Mar 1929 by R. W Hendee (Bangs & Van Tyne, 1931. p. 37; Coolidge. 1933, p. 94); fmnh. 1. B:V-1. Muong Cha; Lai Chau, VIETNAM; 21°58'N, I02°5rE; collected 27 Apr 1963 by unknown collector (Dao. 1985, p. 147); museum un- known (not seen). B:V-3. Muong Mo; Lai Chau, VIETNAM; 22°13'N, 102°55'E; captives acquired 12-20 Mar 1929 by H. J. Coolidge. Jr. and R. W Hendee (Cool- idge. 1933. pp. 86. 216; Bangs & Van Tyne, 1931. p. 35). B:V-2. Muong Moun. Tonkin region; Lai Chau, VIET- NAM; 21°42'N, 103°2rE; collected 15 Mar 1929 by R. E. Wheeler (Bangs & Van Tyne. 1931. p. 34); KMNH. 1. C:V-3. Muong Muon. Tonkin region; Lai Chau, VIET- NAM; 21°40'N. 103°04'E; captive purcha.sed Nov. 1931 by T D. Carter (Legendre, 1936, p. 125); died in zoo 15 June 1932; amnh, 1. C:V-2. Muong Pon (= Muong Poun). Tonkin region; Lai Chau, VIETNAM; 21°33'N, 103°0rE; collect- ed 18 Nov. 1931 by T D. Carter (Legendre, 1936, p. 125); amnh, 1. C:V-2. Muong Son. See Huong Son. Murree, outskirts; Punjab, PAKISTAN; 33°54'N, 73°22'E; reported in 1964 by T J. Roberts (1977, p. 87). A:P-i2. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 155 Mussoorie (= Mussooree) vicinity; Uttar Pra- desh, INDIA; ca. 30°27'N, 78°05'E; reported before 1866 by T. Hutton (1865, p. xiii [mis- identified as Inuus pelops]; cf. Fooden, 1982a, p. 2). A:I-27. Mustapur. 0.5 km east of, Nizamabad District, 560 m; Andhra Pradesh, INDIA; 18°17'N, 78°10'E; observed 15 Apr. 1980 by J. Fooden, A. Ma- habal, and S. S. Saha (1981, p. 467). A:I-I18. Muttagudem. Krishna District, 80 m; Andhra Pra- desh. INDIA; 17°07'N. 80°37'E; observed 14 May 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A:I-136. Muttra. See Mathura. MYANMAR (= BURMA), eastern; 20°-24°N, 97°-10rE; tissue samples obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). Not mapped. MYANMAR (= BURMA), upper; Kachin, MYANMAR; 26°00'-26°40'N, 97°50'-97°55'E; collected ca. 1939 by R. Kaulback (Pocock, 1941, p. v); bm(nh), 1 (skull only). Not mapped (see B:M-1). Myitkyina; Kachin, MYANMAR (= BURMA); 25°23'N, 97°24'E; captive purchased June-Dec. 1945 by M. L. Roonwal (1950, p. 16 [misiden- tified as M. assamensis]). B:M-5. Mymensingh, northern, BANGLADESH; ca. 25°10'N, 90°30'E; reported early in 1980 by S. P. Gittins and A. W. Akonda (1982, p. 278). B: Ba-9. Na chaka. See Yajiang. Nachuka. See Yajiang. Nada (= Nodoa), Hainan Dao; Hainan, CHINA; 19°31'N, 109°33'E; collected 9-28 Jan. and 19 Apr. 1923 by C. H. Pope (1932b, p. 481; 1935, p. 498); AMNH, 4; fmnh, I. C:C-229. Nagarjunakonda Valley. See Siddeldar Hill. Nagarkot (= Nagarcot), 8000 ft (= 2400 m); Bag- mati, NEPAL; 27°42'N, 85°3rE; collected 15 Oct. 1920 by R. L. Kennion (Hinton & Fry, 1923, p. 403); bm(nh), 2. A:N-12. Nagchuka. See Yajiang. Nagorhgena; Assam, INDIA; ca. 24°17'N, 92°30'E; observed 21-25 Mar. 1986 by A. Choudhury (1983, p. 14; 1989, p. 491; [1991a], p. 124). B:I-39. Nagpur, 300 m; Maharashtra, INDIA; 21°10'N, 79°05'E; observed 30 Mar. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 466). A: 1-102. Nai Basti. See Khair, Tahsil. Naini Tal, Bhowal area, Kumaon region; Uttar Pradesh, INDIA; ca. 29°23'N, 79°27'E; ob- served Sept. 1959-June 1960 by C. H. South- wick, M. A. Beg, and M. R. Siddiqi (1961b, p. 703). Parasitological study conducted ca. 1980 by S. N. Arya (1981, p. 261). A:I-32. Naini Tal District; Uttar Pradesh, INDIA; 28°45'- 29°35'N, 78°40'-80°10'E; observed 1981-1983 by R K. Seth, S. Seth, G. L. Reddy, and R K. Chopra (1992, p. 62). Not mapped. Nam Co (= Tengri-Nor); Xizang, CHINA; 30°42'N, 90°35'E; erroneous record (Elliot, 1913, p. 197; cf. Milne-Edwards, 1892, p. 670). Not mapped. Nam Fong. See Nanfeng. Nam hou. See Ou, Nam. Nam miu. See Luofu Shan. Nam Ngap, Luc Yen District, Yen Bai, VIET- NAM; ca. 22°07'N, 104°47'E; collected Dec. 1971 and date unknown by unknown collector; lEBR, 2 (skulls only). C:V-4. Nam U. See Ou, Nam. Nam Yao. See Mansam Falls. Nam Yu. See Ou, Nam. Nanau; Uttar Pradesh, INDIA; 27°48'N, 78°16'E; observed 1959-1975 by C. H. Southwick and M. F Siddiqi (1977, p. 342). Observed Jan. 1990-Mar. 1991 by E. Imam and H. S. A. Yah- ya 1995, p. 2). A:I-69. Nanchuan; Sichuan, CHINA; 29°07'N, 107°16'E; reported before 1992 by Jiang Xuelong, Wang Yingxiang, and Ma Shilai (1991, p. 244). C:C- 133. Nanding He (= Nanting River), Lincang District; Yunnan, CHINA; ca. 24°00'N, 99°44'E; report- ed before 1996 by Lan Daoying and Guo Guang (1995, p. 6). B;C-68. Nandini Wildlife Sanctuary, Jammu District; Jam- mu & Kashmir INDIA; ca. 32°44'N, 74°52'E; reported before 1984 by B. K. Tikader (1983. p. 297). A:I-5. Nanfeng (= Nam Fong) Shi; Hainan Dao; Hai- nan, CHINA; 19°24'N, 109°3rE; collected 23 Mar. and 14 May 1923 by C. H. Pope (1935, p. 498); AMNH, 1; mcz, 1. C:C-229. Nanglamora; Assam, INDIA; ca. 27°00'N, 94°46'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. Nangpoh. See Nongpoh. Nang Pon. See Nongpoh. Nang Xian; Xizang (= Tibet), CHINA; ca. 29°05'N, 93°05'E; observed 1979-1982 by Zhang Cizu, Director, Shanghai Zoo (pers. comm., 18 Oct. 1985). B:C-6. Nanhua; Yunnan, CHINA; 25°13'N, 101°21'E; re- ported before 1998 (Zhang et al., 1997, p. 58). 156 FIELDIANA: ZOOLOGY Blood sample obtained before 1999 by Ding Bo, Zhang Yaping. and Hou Yidi (1998, p. 172). B:C-53. Nanjian; Yunnan, CHINA; 25°04'N, 100°32'E; re- ported before 1998 (Zhang et al., 1997, p. 58). B:C-54. Nanjiang; Sichuan, CHINA; 32°2rE; 106°50'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-33. Nan Ling; Guangdong, CHINA; ca. 24°10'N, 112°00'E; reported 1980-1981 by Fu Ting- zhang (1987, p. 37). C:C-198. Nanning; Guangxi, CHINA; ca. 22°49'N. 108°19'E; captive purchased Jan. 1964 by un- known collector; kiz, 1. C:C-220. Nanping; Sichuan, CHINA; 33°14'N, 104°06'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-22. Nantaleik Chaung. See Hisweht. Naming River. See Nanding He. Nanwan, Xingcungang, Lingshui Xian, 100-200 m; Hainan Dao; Hainan, CHINA; 18°24'N, 109°59'E; captured in 1970 by Liu Zhenhe, sciEA (pers. comm., 26 Nov. 1985); died in cap- tivity 15 May 1976; sciea, 2. Reported before 1981 (Anonymous, 1980, p. 17). Observed be- fore 1988 by Wang Zeng (1987, p. 39). Re- ported before 1989 by Tan Manni (1988, p. 14). C:C-239. Nanwan Nature Reserve, Hainan Dao, 255 m; Hainan, CHINA; 18°23'N, 110°00'E; observed 1965-1989 by Jiang Haisheng, Liu Zhenhe, Zhang Yongzu, and C. Southwick (1991, p. 208; Southwick et al., 1991, p. 25; Jiang et al., 1994, p. 166). Observed 1988-1990 by Feng Min, Jiang Haisheng, and Wang Jun (1997. p. 27). Observed ca. 1991 by D. Manry (1991, p. 10). C:C-239. Nanyaseik, 480 ft (= 145 m); Kachin, MYAN- MAR (= BURMA); ca. 25°37'N, 96°36'E; col- lected 7 and 11 Jan. 1935 by H. C. Raven (in Carter, 1943, p. 100; Monis, 1936, p. 648); AMNH, 4. B:M-6. Naogaon; Naogaon, BANGLADESH; 24°47'N, 88°56'E; reported before 1982 by M. A. R. Khan (1981, p. 13). B:Ba-4. Narasaraopet, 2 km northwest of, Guntur District, 75 m; Andhra Pradesh, INDIA; 16°14'N, 80°02'E; observed 9 May 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 468). A: M45. Narayanapur (= Narayanpur), Jangaon Taluk, Warangal District; Andhra Pradesh, INDIA; 17°44'N, 79°14'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-122. Narayanganj (= Narayangong); Naraxanganj, BANGLADESH; 23°37'N, 90°30'E; reported before 1982 by M. A. R. Khan (1981, p. 13; 1985, p. 31). B:Ba-27. Narbong, Darjeeling Disrtict, 2000 ft ( = 600 m); West Bengal, INDIA; 26°5rN, 88°20'E; col- lected 1 1 Mar. 1915 by C. A. Crump (Wrough- ton, 1916b, p. 472); bm(nh), 1. B:I-6. Narkanda, ca. 1 km north of; Himachal Pradesh, INDIA; ca. 31°17'N, ITll'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984, pp. 475. 483). A:M7. Narkanda, ca. 4 km south of; Himachal Pradesh, INDIA; ca. 3ri4'N, ITII'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984. p. 475). A: 1-17. Narkanda. ca. 5 km north of; Himachal Pradesh, INDIA; ca. 31°18'N. ITII'E; reported Aug. 1972-Feb. 1973 by K. Wada (1984. p. 475). A: M7. Narma. See Narva. Narota-Bun, highway between; Jammu & Kash- mir, INDIA; ca. 32°49'N. 74°55'E; observed before 1983 by Y R. Malhotra and D. N. Sahi (1982, p. 27). A:I-5. Narpuh Reserved Forest, Jaintia Hills; Megha- laya, INDIA; ca. 25°05'N, 92°20'E; observed ca. 1996 by A. Choudhury (1998. p. 8). B:I-18. Narva, Banswada Taluk, Nizamabad District; An- dhra Pradesh, INDIA; 18°14'N, 77°59'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992. p. 18). A:I-117. Nathia Gali; North-West Frontier, PAKISTAN; 34°04'N. 73°24'E; reported in 1964 by T J. Roberts (1977. p. 87). A:P-12. Nawabganj vicinity; Nawabganj, BANGLA- DESH; ca; 24°36'N. 88°17'E; reported before 1982 by M. A. R. Khan (1981. p. 13; 1985, p. 31). B:Ba-3. Nawakot. See Trisuli Bazar. Nazuo Water Regulation Forest Reserve; Guang- xi, CHINA; ca. 24°12'N, 105°32'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127; letter, Aug. 1996). C:C- 162. N"Changyang, 1500 ft (= 460 m); Kachin, MYANMAR (= BURMA); 25°50'N, 97°48'E; collected 4 and 19 July 1939 by R. Kaulback; BM(NH), 2 (including 1 skin only). B:M-3. Neelum Valley; Azad Kashmir, PAKISTAN; ca. 34°20'N, 73°35'E; reported before 1978 by T J. Roberts (1977, p. 86). A:P-9. FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 157 Neghereting (= Negheretting); Assam, INDIA; 26°44'N. 94°07'E; observed in 1969 and 12-15 Feb. 1974 by G. Pilleri (1975, p. 43; Pilleri & Pilleri, 1982, p. 158). B:I-23. Neilingding Dao (= Lintin Island), 50-340 m; Gmmi^Jonii, CHINA; ca. 22°25'N, 113°48'E; tentatively reported before 1863 by R. Swinhoe (1863, p. 351). Collected 25 Oct. 1981 by Liu Zhenhe and Xu Longhuei (Liu Zhenhe, sciea, pers.comm., 26 Nov. 1895); sciea, 1. Observed Dec. 1984 by Liu Zhenhe, sciea (pers. comm.. 25 Nov. 1985). Observed 1988-1990 by Feng Min. Jiang Haisheng, Wang Jun (1997. p. 27). Reported 25 Aug. 1993 by M. W. Lau (1995, p. 209). C:C-212. NEPAL. See [Katmandu Valley]. Nepal Tarai. See Terai. Newakot. See Trisuli Bazar. New Forest Estate. See Dehra Dun vicinity. Ngamda (?= Kintachie; ?= Houmda); Xizang { = Tibet), CHINA; ca. 31°05'N, 96°43'E; captive purchased 7 May 1890 by G. Bonvalot and H. d' Orleans (Bonvalot, 1891, vol. 2, pp. 149, 156; 1892, p. 505; Bonvalot et al., 1891, map); cap- tive living in menagerie of mnhn, 22 Aug. 1892 (Milne-Edwards, 1892, p. 671); skin possibly in MNHN (see below, Tibet). B:C-14. Nghe An, VIETNAM; 18°35'-20°00'N. 103°50'- 105°50'E; collected in 1959 and Nov. 1961 by unknown collectors; zmvnu, 3 (skulls only). Not mapped (see C:V-24 through C:V-26). Nghia Dan (= Nghia Hung, Phu Qui). 100 ft (= 30 m); Nghe An, VIETNAM; 19°19'N, 105°25'E; collected 28 Feb. 1928 by J. Dela- cour and W. P Lowe (Delacour, 1929, p. 198); BM(NH). 1. C:V-24. Nghia Dung, Tan Ky District, Nghe An, VIET- NAM; 19°07'N, 105°2rE; collected 4 and 9 Dec. 1964 by Lo Van Hong (Dao. 1985, p. 216; Nisbitt & Ciochon, 1993, p. 772; cf. Dang, 1983, p. 1282); iebr, 2. C:V-24. Nghia Hung. See Nghia Dan. Nghia Lo. See Yen Bcii. Nhera/Tara Devi, Simla vicinity; Himachal Pra- desh, INDIA; ca. 31°05'N, 77°09'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984, p. 474). A:M8. Nheri. See Narkanda, ca. 1 km north of. Nimaijan-Bahdhora; Assam, INDIA; ca. 27°00'N, 94°4rE; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a]. p. 31). B:I-25. Ningdu; Jiangxi, CHINA; 26°22'N, 115°48'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-100. Ninggang; Jiangxi, CHINA; 26°45'N, 113°58'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-1()3. Ningming; Guangxi, CHINA; 22°12'N, 1()7°05'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-227. Nirmal, 16 km east of, Adilabad District, 360 m; Andhra Pradesh, INDIA; 19°05'N, 78°30'E; observed 26 Mar. 1980 by J. Fooden, A. Ma- habal. and S. S. Saha (1981, p. 466). A:M14. Nishangara, Bahraich District; Uttar Pradesh, IN- DIA; 28°15'N, 81°13'E; collected 1956-1957 by Zoological Survey of India (Kurup, 1965, pp. 186, 193); museum unknown (not seen). A: 1-56. Nishangara vicinity; Uttar Pradesh, INDIA; ca. 28°15'N, 8n3'E; observed 1955-1957 by G. A. von Maydell (Oboussier & von Maydell, 1960, p. 144). A:I-56. Noakhali, BANGLADESH; ca. 22°45'N, 91°10'E; reported before 1982 by M. A. R. Khan (1981, p. 13). B:Ba-31. Nodoa. See Nada. Nong Khai (= Nong Kay; Nong Kha), Camp No. 28; Nong Khai, THAILAND; 17°52'N, 102°44'E; collected 9 Feb. 1920 by J. Bangas- sar (C. B. Kloss, unpublished itinerary notes, ZRC); ZRC. 2. B:T-11. Nonglin, Mengla Xian, 670 m; Yunnan, CHINA; ca. 21°28'N, 101°35'E; collected 19 Dec. 1959 by Deng Xiangfu; Kiz, 1 (skull only). B:C-86. Nongpoh. Khasi Hills, 1200 ft (= 370 m); Megh- alaya INDIA; 25°54'N. 91°53'E; collected 27 May 1920 by H. W. Wells (Hinton & Lindsay, 1926, p. 385); bm(nh). 1. B:I-17. Nongxin Water Regulation Forest Reserve; Guangxi, CHINA; 22°55'N, 105°53'E; ob- served 1976. 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter. Aug. 1996). C:C-165. Nordchina. See CHINA, northern. North Cachar Hills; Assam, INDIA; ca. 25°10'N, 93°00'E; reported Dec. 1972-Feb. 1973 by R. L. Tilson (1983. p. 399). B:I-36. North District, eastern; Tripura, INDIA; ca. 24°00'N. 92°25'E; observed May-June 1978 by R. P Mukherjee (1982, p. 71). Observed May- Aug. 1989 by A. K. Gupta (1994, p. 102). B:I- 41. North District, north-central; Tripura, INDIA; ca. 24°20'N. 92°00'E; observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). B:I-40. North District, northeastern; Tripura, INDIA; ca. 24°20'N, 92°25'E; reported May-June 1978 by 158 FIELDIANA: ZOOLOGY R. P. Mukherjee (1982. p. 71). Observed May- Aug. 1989 by A. K. Gupta (1994. p. 102). B:I- 39. North District, northwestern; Tripura, INDIA; ca. 24°11'N. 91°49'E; observed May-Aug. 1989 by A. K. Gupta (1994. p. 102). B:I-40. North District, southeastern; Tripura, INDIA; ca. 23°47'N, 92°14'E; observed May-Aug. 1989 by A. K. Gupta (1994. p. 102). B:I-42. North District, southwestern; Tripura, INDIA; ca. 23°42'N. 92°0rE; observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). B:I-42. North District, western; Tripura, INDIA; ca. 24°00'N. 91°49'E; observed May-June 1978 by R. R Mukherjee (1982. p. 71). Observed May- Aug. 1989 by A. K. Gupta (1994. p. 102). B:I- 40. North Kamrup. See Bogra Nadi. North Lena Island. See Dangan Dao. North VIETNAM; 17°-23°N. 102°-107°E; tissue samples obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b. p. 589). Not mapped. Nuguvedu. See Yeppuru. Nu Jiang (= Salween River), above Changlung; Yunnan, CHINA; ca. 24°15'N. 99°05'E; tenta- tively reported Mar. 1917 by R. C. Andrews (in Andrews & Andrews, 1918. p. 279). B:C-65. Nurestan (= Nuristan). eastern; Konarha, AF- GHANISTAN; ca. 35°30'N. 7r30'E; captive obtained in 1906 by H. McMahon (Pocock, 1932, p. 543); died 19 Jan 1910 in Regents Park Zoo; bm(nh). 1 (skin only). A:A-1. Nurestan (region), densely wooded districts; Kon- arha, Laghman, or Nangarhar, AFGHANI- STAN; 34°-36°N. 70°-72°E; reported before 1860 by H. G. Raverty (1859. p. 332). Not mapped. Nurestan vicinity; Laghman, AFGHANISTAN; 35°00'N. 70°20'E; reported before 1972 by A. Puget (1971. p. 201); A:A-4. Nur Valley. See Khyber Pass vicinity. Nyachuka. See Yajiang. Nyainqentanglha Sheng (region); Xizang (= Ti- bet), CHINA; ca. 29°-32°N, 90°-96°E; reported before 1964 by Shen Xiaozhou (1963, p. 140; cf. Fooden, 1982a. pp. 26. 51; 1989, p. 44). Not mapped (see B:C-1, B:C-3. and B:C-6 through B:C-13). Nychow (?= Yai-cheng) vicinity; Hainan Dao; Hainan, CHINA; ca. 18°22'N. 109°08'E; col- lected Mar. 1868 by R. Swinhoe (1870. p. 226; Napier. 1981. p. 22); bm(nh). 1. C:C-237. Old Chandpur Bazar. See Chandpur Bazar, old. Olongche (?= Wolongshi); Sichuan, CHINA; ca. 30°03'N. 101°2rE; collected [21 Jun. 1890] by G. Bonvalot and H. d'Orleans (Bonvalot. 1892, p. 506); MNHM, 2 (including 1 skull only). B:C- 28. Omei. Mount. See Emei Shan. Orcha. ca. 1 km northwest of. Bastar District; Madhya Pradesh, INDIA; ca. 19°22'N. 8ri2'E; observed Nov. 1958-Nov. 1959 by R Jay (1963. p. 281; 1965. p. 210; cf. Fooden. 1989. p. 44). B:I-111. Ou. Nam (= Nam hou); Louangphrahang, LAOS; ca. 20°30'N. 102°35'E; collected ca. Apr. 1892 by H. d'Orleans (Gagnepain, 1944, map 1. p. 45); MNHN. 1 (skull only). B:L-2. Ou. Nam. between Muang Khoua and Muang Ngoy; Louangphrahang or Phongsali, LAOS; ca. 21°00'N. 102°45'E; collected 20 May 1929 by R. W. Hendee (field catalog, fmnh, p. 49; Bangs & Van Tyne. 1931. p. 37; Osgood. 1932. p. 195) FMNH. 1. B:L-1. Ououlongtche. See Olongche. Outapour, south of; Konarha, AFGHANISTAN; ca. 34°50'N. 70°50'E; reported before 1972 by A. Puget (1971. p. 201). A:A-7. Ovra (= Overa) Sanctuary, proposed, 2135 m; Jammu & Kashmir, INDIA; ca. 34°00'N, 75°00'E; observed Apr.-May 1981 by P C. Tak and G. Kumar (1984, p. 203; Roonwal & Tak, 1981, p. 96). A:I-2. Paach piror mukam; Assam, INDIA; ca. 24°16'N. 92°30'E; observed 21-25 Mar. 1986 by A. Choudhury (1983. p. 14; 1989. p. 491; 11991b]. p. 124). B:I-39. Pablakhali; Chittagong Hill Tracts, BANGLA- DESH; 23°17'N. 92°07'E; observed Feb. 1990- June 1993 by M. M. Feeroz. M. A. Islam, and M. Kabir (1995. p. 76). B:Ba-34. Paddavaram. Guntur District; Andhra Pradesh, INDIA; 16°0rN. 79°38'E; population report- edly introduced in 1977. observed 5 May 1980 by J. Fooden. A. Mahabal. and S. S. Saha (1981. p. 471). Not mapped. Padua; Chittagong. BANGLADESH; 22°03'N. 92°07'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-38. Pahalgam vicinity. See Ovra (= Overa) Sanctu- ary. Paia. ca. 6 mi (= 10 km) east of Shogran. Hazara Division, 8700 ft (= 2650 m); North-West Frontier, PAKISTAN; 34°37'N, 73°33'E; col- lected 5 Aug. 1964 by M. Iqbal (Roberts. 1977, p. 343); USNM. 1. A:P-10. Pajja Hill, north of Mardan; North-West Frontier, FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE. MACACA MULATTA 159 PAKISTAN; ca. 34°I2'N, 72°02'E; observed 1899-1901 by unidentified British officers (Mc- Mahon, 1901b, p. 9). A:P-7. Pakhal Lake, west side, Warangal District, 340 m; Amlhra Pradesh, INDIA; 17°56'N, 79°58'E; observed 19 and 20 Apr. 1980 by J. Fooden, A. Mahabal. and S. S. Saha (1981. p. 467). A:I- 133. Pakhal Wild Life Sanctuary; Andhra Pradesh, IN- DIA; 17°50'-18°05'N, 79°55'-80°10'E; report- ed Nov. 1966 by J. J. Spillett (1968, p. 8). Not mapped (see A:I-133). PAKISTAN; 33°-36°N, 7r-74°E; obtained in Ka- rachi (extralimital) before 1949 by R. Henry; iRSN, 3. Not mapped. Paksane. See Muang Pakxan. Palamau; Bihar, INDIA; ca. 23°50'N, 84°10'E; observed 22 Feb. 1970 by M. Krishnan (1972, p. 540). A:I-92. Palampeta, Mulug Taluk, Warangal District; An- dhra Pradesh, INDIA; 18°16'N, 79°56'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984. p. 58; 1992, p. 18). A:I-120. Pallepadu (= Pallipad), Khammam Taluk, Kham- mam District, Andhra Pradesh, INDIA; 17°12'N, 80°20'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). A:I-I32. Pamulaparthi (= Pamulparthi), Gajwel Taluk, Me- dak District; Andhra Pradesh, INDIA; 17°46'N, 78°42'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-123. Pang Nam Un; Nan, THAILAND; 18°30'N, 100°33'E; collected 25 Jan. 1953 by R. E. Elbel and Prasit Seecharong (Moore & Tate, 1965, p. 329). usNM, 1. B:T-3. Panighatta; West Bengal, INDIA; 26°48'N, 88°15'E; observed in 1962 by C. H. Southwick, A. Ghosh, and C. D. Louch (1964, p. 446). B: 1-6. Panipat-Rhotak, highway between; Haryana, IN- DIA; ca. 29°05'N, 76°40'E; observed 1964- 1965 by R. R Mukherjee and G. D. Mukherjee (1972, p. 67). A:I-37. Pankhabari. See Simulbari-Pankhaburi. Paras vicinity, lower Kaghan Valley; North-West Frontier, PAKISTAN; ca. 34°39'N. 73°3rE; reported before 1978 by T J. Roberts (1977. p. 86). A:P-10. Pashok; Sikkim, INDIA; 27°03'N, 88°25'E; im- probable report (Wroughton, 1916d, p. 776; cf. Fooden, 1982a, p. 51). Not mapped. Pashupati, 4400 ft (= 1300 m); Katmandu Valley, NEPAL; ca. 27°38'N. 85°21'E; observed 1974- 1975 by H. Taylor, J. Teas, T Richie, C. South- wick, and R. Shrestha (1978, p. 344). Observed 1975-1978 by J. Teas (1983, p. 224). Reported before 1982 by T K. Shrestha (1981, p. 269). Reported 13 Aug. 1996 by N. Shrestha (1997, p. 31). Observed 1995-1998 by M.K. Chalise and M. Ghimire (1998, p. 11). A:N-12. Patang. See Batang vicinity. Patharia (= Pathalia); Moulvi Bazar, BANGLA- DESH; ca. 24°45'N, 92°15'E; reported early in 1980 by S.R Gittins and A. W. Akonda (1982, p. 278). Observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, pp. 76, 79). B:Ba-ll. Patiala District; Punjab, INDIA; ca. 30°19'N. 76°24'E; observed 1981-1983 by RK. Seth, S. Seth, G. L. Reddy. and R K. Chopra (1992, p. 62). A:I-21. Patriata, Murree region. 7150 ft (= 2180 m); Pun- jab, PAKISTAN; 33°5rN, 73°29'E; collected 14-15 June 1923 by H. W. Wells (in Lindsay, 1926, p. 608); bm(nh), 1; bnhs, 1. A:P-12. Peak. See [Victoria] Peak. Pekin. See Beijing. Phala/Kutbor Game Reserve; Azad Kashmir, PAKISTAN; ca. 34°00'N, 73°35'E; reported be- fore 1984 by M. Nawaz (1983, p. 6). A:P-12. Phatuntula; Sylhet, BANGLADESH; not precisely located, 24°08'-24°50'N, 9r37-92°17'E; ob- served Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). Not mapped. Phawngpui Wildlife Sanctuary; Mizoram, INDIA; ca. 22°40'N, 93°03'E; observed 13-18 Jan. 1994 by T R. S. Raman, C. Mishra, and A. J. T Johnsingh (1996, p. 59). B:I-43. Phu Qui. See Nghia Dan. Phu Vach, Tan Lac District; Hoa Binh, VIET- NAM; ca. 20°35'N, 105°18'E; collected 21 Jan. 1973 by Pham Trong Anh; iebr, 1 (skin only). C:V-21. Piangzu, 3.5 km northeast of Banli, Luoshan Sub- county, Jinxiu Xian; Guangxi, CHINA; ca. 24°02'N, 110°15'E; captive purchased ca. 1987 by Zhong Changwan (pers. comm., 21 Nov. 1992); died in 1991; skeleton examined 21 Nov. 1992 at Banli. C:C-195. Pidaung Reserve. See Karen Chaung. Pilibhit-Tanakpur, highway between; Uttar Pra- desh, INDIA; ca. 28°45'N, 79°50'E; observed Sept. 1959-June 1960 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961b, p. 702). A: 1-51. Pina, forests above, Rara Daha (= Lake) vicinity; 160 FIELDIANA: ZOOLOGY I Jumla, NEPAL; ca. 29°33'N, 82°05'E; observed Nov. 1979 by P. Byrne (1982, p. 115). A:N-6. Pingbian; Yunnan, CHINA; 22°54'N, 103°40'E; reported before 1992 by Jiang Xuelong, Wang Yingxiang, and Ma Shilai (1991, p. 242). C:C- 151. Pinglang. See Batu. Pingnan Xian; Fujian, CHINA; ca. 26°56'N, I19°03'E; reported Sept. 1980 by Zheng Xue- qing (1984. p. 146). C:C-70. Pingwu; Sichuan, CHINA; 32°25'N, 104°36'E; re- ported before 1998 (Zhang et al.. 1997, p. 58). C:C-24. Pinxiang vicinity; Jiangxi, CHINA; ca. 27°37'N, 113°5rE; captured in 1983 for Pingxiang Zoo; observed at zoo in summer 1984 by Sheng He- lin, ECNU (pers. comm., 19 Oct. 1985). C:C-106. Pisui, Loi-Gebiet, Hainan Dao; Hainan, CHINA; ca. 19°02'N, 109°43'E; collected 19 Mar. 1909 by H. Schoede; zmb, 1. C:C-232. Plateau des Bolovens. See Muang Thateng. Pochuan. 6-7 km west of, Xianan Subcounty, Huanjiang Xian, 460 m; Guangxi, CHINA; ca. 25°00'N, 107°52'E; collected 25 Oct. 1992 by Tan Nenrui (pers. comm., 5 Nov. 1992); IZCAS, 1. C:C-177. Podumoni. See Tinsukia. Pokhara; West No. 4, NEPAL; 28°12'N, 83°56'E; observed Mar. 1996 by M. K. Chalise and M. Ghimire (1998, p. 12; e-mail, 20 Nov. 1998). Reported at Annapuma Conservation Area be- fore 1997 by K. K. Gurung and R. Singh (1996, p. 78). A:N-supplementary. Po Lu, Ba Be vicinity; Cao Bang, VIETNAM; ca. 22°24'N, 105°38'E; collected 6 Aug. 1967 by Vo Quy; zmvnu, 1 (skull only). C:V-8. Popa Hill; Mandalay, MYANMAR (= BURMA); ca. 20°55'N, 95°i5'E; observed Feb. 1984 by C. H. Southwick and K. L. Southwick (1985, p. 35). B:M-27. Popa Hill, 1000 m; Mandalay, MYANMAR ( = BURMA); 20°55'N, 95°15'E; collected 21 Oct.-5 Nov. 1937 by G. Heinrich; amnh, 6. B: M-27. Popa Hill, 4961 ft (= 1512 m); Mandalay, MYANMAR (= BURMA); 20°55'N, 95°15'E; collected July-Oct. 1913 by G. C. Shortridge (in Wroughton, 1915a, p. 461); bm(nh), 1; bnhs, 5 (including 1 skull only). B:M-27. Prag Oil Mill. See Aligarh. Prome. See Pye. Pu'er Xian; Yunnan, CHINA; ca. 23°05'N, 101°03'E; immunological survey conducted be- fore 1996 by Duan Xingsheng, Liu Yuanwei, Wu Jing, Dao Weiying, and Liu Jianghai (1995, p. 411). B:C-77. Pucheng Xian; Fujian, CHINA; ca. 27°55'N, 118°30'E; collected 15 Aug. 1960 by unknown collector; sciea, 1 (skin only). Reported Aug. 1980 by Zheng Xueqing (1984, p. 145). C:C- 75. Pulareddi. See Siddeldar Hill. Pulga; Himachal Pradesh. INDIA; ca. 32°()()'N, 77°30'E; reported 1978-1980 by A. J. Gaston, R J. Garson. and M. L. Hunter. Jr. (1983, p. 300). A:I-15. Punch Bol, Amravati District, 825 m; Maharash- tra, INDIA; 21°26'N, 77°17'E; observed 31 Jan. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 466). A:I-101. Puquan Road Maintenance Station, Zhongliang Subcounty, Jinxiu Xian, 850 m; Guangxi, CHI- NA; 24°irN, 110°19'E; observed in 1991 by Mo Cailian and Mo Xiuzhen, road maintenance workers (pers. comm., 14 Nov. 1992). C:C-195. Puran Bazar, Chandpur Division; Chandpur, BANGLADESH; 23°20'N, 90°47'E; reported before 1986 by M. A. R. Khan (1981. p. 13; 1985, p. 31). B:Ba-28. Puri, 25 m; Orissa, INDIA; 19°48'N, 85°50'E; ob- served 16 Feb. 1980 by R. P Mukherjee (1984, p. 261). Observed 21 May 1980 by J. Fooden. A:I-106. Pyaunggaung, 2794 ft (= 852 m); Shan, MYAN- MAR (= BURMA); 22°35'N. 97°05'E; collect- ed 8 and 14 May 1913 by G. C. Shortridge (in Ryley, 1914, p. 713; Moore & Tate, 1965, p. 330); BM(NH), 2 (including 1 in alcohol); bnhs, 2 (including 1 skull only). B:M-20. Pye (= Prome), 30 mi (= 50 km) southeast of, 200 ft (- 60 m); Irrawaddy, MYANMAR ( = BURMA); ca. I8°30'N, 95°30'E; collected 2 Feb. 1917 by J. M. D. Mackenzie (Wroughton, 1921, p. 553); bnhs, 1. B:M-34. Pye (= Prome), 35 mi (= 55 km) southeast of, 800 ft (= 240 m); Pegu, MYANMAR (= BUR- MA); ca. 18°30'N, 95°35'E; collected 25 Oct. 1916 by J. M. D. Mackenzie (Wroughton, 1921, p. 553); BM(NH), 1. B:M-34. Qamdo; Xizang (= Tibet), CHINA; 3nO'N, 97°14'E; reported before 1992 by Jiang Xue- long, Wang Yingxiang, and Ma Shilai (1991. p. 245). B:C-21. Qasimpur Canal; Uttar Pradesh, INDIA; 27°59'N, 78°09'E; observed 1959-1975 by C. H. South- wick and M. F Siddiqi (1977, p. 342). A:I-69. Qianjiandong Water Regulation Forest Reserve; Guangxi, CHINA; ca. 25°26'N, 111°16'E; ob- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 161 served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995. p. 128; letter, Aug. 1996). C:C-191. Qianxian. See Yixian. Qiasui. Huaiji Xian; Guangdong, CHINA; ca. 24°06'N, 1 12°20'E; reported Apr 1982 by local residents (Liu Zhenhe, sciiiA, pers. comm., 26 Nov. 1985). C:C-107. Qigong, Yangshan Xian; Guangdong, CHINA; 24°18'N, 1 12°34'E; reported July 1982 by local residents (Liu Zhenhe, scihA. pers. comm., 25 Nov. 1985). C:C-206. Qihong, Qimen Xian, 200-600 m; Anhui, CHI- NA; ca. 29°35'N, 117°40'E; observed 1973- 1986 by Xiong Chenpei (Wada et al., 1986, p. 83). C:C-63. Qingchuan; Sichuan, CHINA; 32°36'N, 105°09'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-23. Qingliangfeng, Jixi Xian, 800 m; Anhui, CHINA; ca. 30°10'N, 118°50'E; observed 1973-1986 by Xiong Chenpei (Wada et al., 1986, p. 83). C:C- 55. Qinglong Shan Water Regulation Forest Reserve; Guangxi, CHINA; ca. 22°33'N, 106°42'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter, Aug. 1996). C:C-224. Qingshi Tan Water Regulation Forest Reserve; Guangxi, CHINA; ca. 25°36'N, llO^ll'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127; letter, Aug. 1996). C:C-193. Qingzhen; Guizhou, CHINA; 26°33'N, 106°28'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-145. Qinyang; Henan, CHINA; 35°06'N, 1 12°57'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-11. Qionglai; Sichuan, CHINA; 30°25'N, I03°29'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-29. Quan Lan, Dao; Quang Ninh, VIETNAM; ca. 20°52'N, 107°30'E; collected 3 June 1969 by unknown collector; iebr, 1 (skull only). C:V-13. Quang Ninh. See Xuan Ninh. Quanzhou; Guangxi, CHINA; 25°57'N, 1 1 1°04'E; reported before 1998 (Zhang et a)., 1997, p. 58). C:C-188. Qusum Xian; Xizang (= Tibet), CHINA; ca. 29°05'N. 92°10'E; observed 1979-1982 by Zhang Cizu, Director. Shanghai Zoo (pers. comm., 18 Oct. 1985). B:C-3. Quxian; Zhejiang, CHINA; 28°58'N, 118°52'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-64. Raghunandan; Habiganj, BANGLADESH; ca. 24°10'N, 91°20'E; observed early in 1980 by S. P. Gittins and A. W. Akonda (1982, p. 278). B: Ba-14. Raghunathapalle, Jangaon Taluk, Warangal Dis- trict; Andhra Pradesh, INDIA; 17°45'N, 79°I5'E; reported Feb. 1977-JuIy 1980 by G. U. Kurup (1984, p. 58; 1992. p. 18). A:I-122. Raigir. Hyderabad District; Andhra Pradesh, IN- DIA; 17°32'N, 78°57'E; observed 1972-1973 by N. Koyama and P. B. Shekar (1981, p. 248). A:I-125. Railway Station. See Aligarh. Raimona vicinity, Goalpara district, <270 m; As- sam, INDIA; 26°39'N, 89°58'E; collected 1956-1957 by Zoological Survey of India (Ku- rup, 1965. pp. 188, 193); museum unknown (not seen). Reported before 1963 by H. Kha- juria (1962b, p. 128). Observed May-June 1973 by R. R Mukherjee and S. S. Saha (1974, p. 337; Mukherjee, 1978b, p. 742). B:I-10. Raj Bhavan compound. See Mumbai. Rajahmundry, 3 km northeast of. East Godavari District, 50 m; Andhra Pradesh, INDIA; 17°02'N, 8r49'E; observed 18 May 1980 by J.Fooden, A. Mahabal, and S. S. Saha (1981, p 467). A:I-138. Rajahmundry, 13 km northeast of. East Godavari District, 75 m; Andhra Pradesh, INDIA 17°03'N, 8r52'E; observed 18 May 1980 by J Fooden, A. Mahabal. and S. S. Saha (1981. p 467). A:I-138. Rajaji and Corbett National Parks; Uttar Pradesh, INDIA; 29°25'-30°13'N, 77°54'-79°05'E; re- ported before 1995 by A. J. T Johnsingh and J. Joshua (1994, p. 137). Not mapped (see A:I-27 and A:I-33). Rajaji Wildhfe Sanctuary, 365-610 m; Uttar Pra- desh, INDIA; ca. 30°10'N, 78°00'E; observed Mar.-Dec. 1978 by J. W. Laws and J. V. H. Laws (1984, p. 35). A:I-27. Rajapara, South Kamrup, 600 ft (= 180 m); As- sam, INDIA; ca. 25°55'N, 9n5'E; collected 21 and 25 Nov. 1920 by H. W. Wells (Hinton & Lindsay, 1926, p. 385); bm(nh), 2 (including 1 skin only). B:I-15. Rajendrapur forest. See Ghazipur. Rajkandi; Moulvi Bazar, BANGLADESH; ca. 24°15'N, 9r55'E; reported eariy in 1980 by S.R Gittins and A. W. Akonda (1982, p. 278). Reported Oct. 1986, Apr 1988. and Nov. 1988 by C. B. Stanford (1992, p. 190). Observed 162 FIELDIANA: ZOOLOGY Feb. 1990-June 1993 by M. M. Feeroz. M. A. Islam, and M. Kabir (1995. p. 76). B:Ba-13. Rajmai tea garden, east of National Highway No. 37; Assam, INDIA; ca; 27°07'N. 94°44'E; ob- served 9 Mar. 1987-16 Feb. 1988 by A. Choud- hury ([1991aJ. p. 32). B:I-25. Rajmai tea garden, west of National Highway No. 37; Assam, INDIA; ca. 27°07'N. 94°43'E; ob- served 9 Mar. 1987-16 Feb. 1988 by A. Choud- hury ([1991a]. p. 32). B:I-25. Rama Pass. See Xi Golog. Ramban Township vicinity; Jammu & Kashmir. INDIA; ca. 33°15'N. 75°15'E; parasitological survey conducted ca. Nov. 1973 by D. Wein- man (1974. p. 345). A:I-4. Ramganga River. Corbett National Park; Uttar Pradesh, INDIA; ca. 29°35'N, 79°05'E; ob- served Sept. 1959-June 1960 and 1964-1965 by C. H. Southwick. M. A. Beg, and M. R. Siddiqi (1961b. p. 703; Southwick & M. R. Sid- diqi. 1966. p. 312). A:I-33. Ramganga South Station, near Bareilly; Uttar Pradesh, INDIA; ca. 28°21'N. 79°25'E; ob- served 1990-1991 by C. H. Southwick and M. F Siddiqi (1994a. p. 227). A:I-52. Ramnagar; Chitawan, NEPAL; 27°44'N. 84°27'E; observed 1992-1994 by M. K. Chalise and M. Ghimire (1998. p. 11). A:N-supplementary. Ramnagar; Jammu & Kashmir INDIA; 32°49'N, 75°19'E; observed before 1983 by Y. R. Mal- hotra and D. N. Sahi (1982. p. 27). A:I-9. Ramnagar. Kumaun region. 1100 ft (= 340 m); Uttar Pradesh, INDIA; ca. 29°24'N. 79°07'E; collected Aug. 1913-Mar. 1914 by C. A. Crump (in Wroughton. 1914. p. 284; Napier. 1981, p. 24); BM(NH), 3 (1 skin only. 2 skulls only); bnhs. 1. A:I-33. Rampur, northwest of; Himachal Pradesh, IN- DIA; ca. 31°27'N, 77°38'E reported Aug. 1972- Feb. 1973 by K. Wada (1984, p. 476). A:I-I7. Rampur-Ghaziabad; Uttar Pradesh, INDIA; ca. 28°50'N, 78°10'E; observed 29 Oct. 1964-9 May 1965 by C. H. Southwick, D. Lindburg, P Jay, M. Khati, and B. Singh (Southwick and M. R. Siddiqi, 1966, p. 306). A:I-47. Rangoli Reserve Forest. See Diroi (Rangoli) Re- » serve Forest. Rangpur, BANGLADESH; ca. 25°40'N. 89°20'E; reported before 1982 by M. A. R. Khan (1981. p. 13; 1985. p. 31). B:Ba-2. Rara Daha (= Lake). See Hutu Forest; Pina. Rasulpur vicinity. Madhupur Forest; Tangail, BANGLADESH; ca. 24°42'N, 90°09'E; ob- served May 1977 by J. R. Oppenheimer, A. W. Akonda. and K. Z. Husain (1983. p. 194). B: Ba-7. Ratighat. Naini Tal vicinity, Kumaun region, 3700 ft and 3800 ft (1 130 m and 1 160 m) Uttar Pra- desh, INDIA; 29°27'N. 79°29'E; collected 1-8 Nov. 1913 by CA. Crump (in Wroughton, 1914, p. 283); bm(nh), 2; bnhs, 1. A:I-32. Rehnathapalli. See Raghunathapalle. Rema-Kalenga, Moiilvi Bazar, BANGLADESH; ca. 24°06'N, 91°36'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, pp. 76, 79). B:Ba-13. Rest House. See Dehra Dun vicinity. Rewari-Patudi-Gurgaon, highway between Hary- ana, INDIA; ca. 28°20'N, 76°50'E; observed 1964-1965 by R. P Mukherjee and G. D. Mu- kherjee (1972. p. 67). A:I-42. Ridge. See Simla vicinity. Ripon Hospital. See Simla vicinty. Rohtak District; Haryami, INDIA; ca. 28°53'N. 76°44'E; observed 1981-1983 by P K. Seth, S. Seth. G. L. Reddy. and R K. Chopra (1992, p. 62). A:L37. Rongrenggiri vicinity, Garo hills; Assam, INDIA; ca. 25°33'N, 90°34'E; observed Jan.-Feb. 1957 by H. Khajuria (1962a, p. 122). B:I-13. Rongtong. See Sukna-Kurseong. Rouetoundo (= Routeoundo); Xizang (= Tibet), CHINA; ca. 31°35'N, 97°25'E; infant captured and two specimens collected 17 May 1890 by G. Bonvalot and H. d'Orleans (Bonvalot. 1891, vol. 2, p. 156; 1892. p. 505; Bonvalot et al.. 1891, map); skins possibly in mnhn (see below, Tibet). B:C-20. Royal Manas National Park; Galygpiig, BHU- TAN; ca. 26°47'N, 90°50'E; reported before 1990 by P B. Subba and C. Santiapillai (1991- 92, p. 32). Reported before 1997 by K. K. Gu- rung and R. Singh (1996, p. 104). B:Bh-l. Rucun, Xiuning Xian, 500-800 m; Anhui, CHI- NA; ca. 29°55'N, 118°07'E; observed 1973- 1986 by Xiong Chenpei, K. Wada, and Wang Qishan (Wada et al.. 1986. pp. 83. 88). C:C-62. Rudrur Agricultural Station. Nizamabad District, 440 m; Andhra Pradesh, INDIA; 18°35'N, 77°53'E; observed 16 Apr 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A: 1-116. Ruicheng; Shanxi, CHINA; 34°42'N. 110°42'E; reported before 1998 (Zhang et al.. 1997, p. 58). C:C-15. Russian Camp, Birganj Forest District; Bara, NE- PAL; 27°12'N, 85°04'E; observed June 1964- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 163 Dec. 1965 by D. L. Chesemore (1970, p. 164). A:N-13. S. A. Factory. See Dhaka. Sabaya Khola; Dhankiita, NEPAL; ca. 27°20'N, 87°15'E; observed ca. 1973 by J. A. McNeeley (letter, 4 Feb. 1973). B:N-2. Sadard Devi. 1 km west of, Valsad District, 75 m; Gujarat. INDIA; 20°48'N. 73°29'E; observed 9 Jan. 1980 by J. Fooden. A. Mahabal, and S. S. Saha (1981, p. 466). A:I-99. Sagarmatha (= Mount Everest) National Park; East No. i, NEPAL; ca. 27°55'N. 86°50'E'; re- ported before 1997 by K. K. Gurung and R. Singh (1996, p. 114). B:N-supplementary. Saharanpur District; Uttar Pradesh. INDIA; 29°30'-30°20'N, 77°00'-78°10'E; observed 1981-1983 by P K. Seth, S. Seth, G. L. Reddy, and R K. Chopra (1992. p. 62). Not mapped (see A:I-25). Saharanpur vicinity; Uttar Pradesh. INDIA; ca. 29°58'N, 77°33'E; autopsied ca. 1966 by K. K. Chawla, C. D. S. Murthy, R. N. Chakravarti, and R N. Chhutani (1967. p. 85). A:I-25. Sahebgunj. Gaya; Bihar, INDIA; ca. 24°50'N, 85°00'E; reported 1811-1812 by F Buchanan ([1936], p. 403, posthumous publication). A:I- 91. Sainj; Himachal Pradesh. INDIA; ca. 3r45'N, 77°25'E; reported 1978-1980 by A. J. Gaston, P J. Garson, and M. L. Hunter, Jr. (1983, p. 300). A:I-16. Saktesgarh; Uttar Pradesh. INDIA; 24°59'N, 82°49'E; collected before 1849 by unknown collector (Napier, 1981, p. 25); bm(nh), 1 (skull only). A:I-86. Sala Reserve Forest; Assam. INDIA; ca. 27°00'N, 94°54'E; observed 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a]. p. 32). B:I-25. Salween River. See Nu Jiang. Samaguting; Nagaland. INDIA; 25°47'N. 93°47'E; collected before 1873 by J. Butler (Anderson, 1881, p. 69); zsi, 1 (skin only). B: 1-34. Samayala ( = Samyala), Kangra Valley, 5000 ft ( = 1500 m); Himachal Pradesh. INDIA; ca. 32°10'N, 76°25'E; collected 9 May 1922 by H. W. Wells (Lindsay, 1926, p. 599); b.vi(nh), 2. A: 1-12. Sambalpur. See Deogarh. Sam Shui Wan Valley; Xianggang (— Hong Kong). CHINA; ca. 22°14'N, 114°10'E; report- ed before 1952 by G. A. C. Herklots (1951. p. 83). C:C-210. Sandu; Guizhou, CHINA; 25°59'N. 107°52'E; re- ported before 1998 (Zhang et al.. 1997. p. 58). C:C-122. Sangu/ Matamuhari; Bandarban, BANGLA- DESH; ca. 2r30'N, 92°30'E; reported early in 1980 by S. P Gittins and A. W. Akonda (1982, p. 278). B:Ba-37. Sangzhi; Hunan. CHINA; 29°24'N, 110°09'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-111. Sankat Mochan Temple. See Varanasi. Sankhu-Bajrajogini; Katmandu Valley, NEPAL; ca. 27°43'N, 85°27'E; observed June 1996 by M. K. Chalise and M. Ghimire (1998, pp. 12, 15). A:N-supplementary. Sankhuwa Khola, both banks. Makalu-Barun Conservation Area; Dhankuta. NEPAL; ca. 27°30'N, 87°05'E; observed ca. 1997, by M. K. Chalise (1997. p. 31; e-mail, 9 Nov. 1998). B: N-1. Sanming Xian; Fujian, CHINA; ca. 26°14'N, 117°35'E; reported Apr. 1981 by Zheng Xue- qing (1984. p. 145). C:C-88. Sanpihu Water Regulation Forest Reserve; Guangxi, CHINA; ca. 25°06'N, 107°13'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C-173. Santaishan, Luxi Xian; 1250 m; Yunnan. CHINA; ca. 24°15'N, 98°25'E; collected 11 Apr. 1962 by unknown collector (Wang Yingxiang, Kiz, pers. comm., 1 Sept. 1983); kiz 1. B:C-62. Sanya, Hainan Dao; Hainan, CHINA; 18°14'N, 109°29'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-238. Sapekhati Reserve Forest; Assam, INDIA; ca. 27°08'N. 95°11'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-30. Saraguri; Assam, INDIA; ca. 27°00'N, 94°29'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. Sarahan; Himachal Pradesh, INDIA; ca. 31°35'N, 77°30'E; reported 1978-1980 by A. J. Gaston, P J. Garson, and M. L. Hunter, Jr. (1983, p. 300). A:I-16. Saraswati Forests, Kurukshetra District; Haryana, INDIA; 29°58'N, 76°32'E; observed Oct. 1989-Sept. 1990 by R. C. Gupta and S. Kumar (1992, p. 226). A:I-22. Sariska Tiger Reserve; Rajasthan, INDIA; ca. 27°20'N, 76°25'E; reported May-Oct. 1990 by C. Ross and A. Srivastava (1994, p. 362). Re- ported before 1997 by K. K. Gurung and R. Singh (1996, p. 116). A:I-74. 164 FIELDLMSfA: ZOOLOGY Sarupduli. See Ramganga River. Sasni; Uttar Pradesh. INDIA; 27°43'N, 78°05'E; observed Jan. 1990-Mar. 1991 by E. Imam and H. S. A. Yahya (1995. p. 2). A:I-69. Satghar; Chittagong. BANGLADESH; ca. 22°20'N. 92°00'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-35. Satha. mango grove; Uttar Pradesh. INDIA; 27°58'N, 78°08'E; observed 1959-1975 by C. H. Southwick and M. F Siddiqi (1977. p. 342). A:I-69. Satkhira; Satkhira, BANGLADESH; 22°43'N. 89°06'E; reported before 1982 by M. A. R. Khan (1981. p. 13). B:Ba-20. Satkhira, southern; BANGLADESH; ca. 22°00'N, 89°10'E; observed 1951-1961 by A. K. Mandal (1964, p. 164). B:Ba-22. Sattenapalle, Guntur District, 75 m; Andhra Pra- desh, INDIA; 16°23'N, 80°09'E; observed 9 May 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 468). A:I-145. Sawai Madhopur; Rajasthan, INDIA; 25°59'N, 76°22'E; reported before 1965 by I. Prakash (letter, 25 Aug. 1964). A:I-82. Sayabouri. See Xaignabouri. School Yard. See Chhatari-do-Raha. Se-eng, Hsipaw District, 141 1 ft (= 430 m); Shan, MYANMAR (= BURMA); 22°43'N, 97°31'E; collected 25 May 1913 by G. C. Shortridge (in Ryley, 1914, p. 713); bm(nh), 1. B:M-19. Seoni. See Malua. Seri. See Sungri, ca. 2 km south of. Setschuen. See Sichuan. Sevoke. See Sivok. Shahabad District; Bihar, INDIA; ca. 25°30'N, 84°15'E; reported 1809-1810 by E Buchanan (1934, p. 227), posthumous publication). A:l- 90. Shahjahanpur-Bareilly, highway between; Uttar Pradesh, INDIA; ca. 28°00'N, 79°40'E; ob- served 1964-1965 by R. R Mukherjee and G. D. Mukherjee (1972, p. 67). A:I-53. Shahjahanpur-Lucknow, highway between, Uttar Pradesh, INDIA; ca. 27°30'N, 80°30'E; ob- served Sept. 1959-June 1960 by C. H. South- wick, M. A. Beg, and M. R. Siddiqi (1961b, p. 702). A:I-57. Shakari Bazar. See Dhaka. Shangang Nature Reserve, Chong'an Xian; Fii- jian, CHINA; 27°45'N, 1 I7°40'E; calls heard in 1982 by Tang Ziying, fubd (pers. comm.. 19 Oct. 1985). C:C-76. Shangchuan Dao. See Miwan. Shanghai. See Sichuan. Shanghang Xian; Fujian, CHINA; ca. 25°05'N, II6°30'E; reported Sept. 1982 by Zheng Xue- qing (1984. p. 146). C:C-94. Shangsi; Guangxi, CHINA; 22°10'N, 108°00'E; reported before 1998 (Zhang et al.. 1997, p. 58). C:C-218. Shangzhou Is. See Miwan. Shanman, Menghai Xian; Yunnan, CHINA; ca. 2r55'N, 100°25'E; collected Nov. 1957 by un- known collector (Wang Yingxiang, Kiz. pers. comm.. 1 Sept. 1983); kiz 1 (skin only). B:C- 81. Shanmoji. Dabu District. Ruyuan Yaozu Zizhi- xian. >1000 m; Guangdong. CHINA; 24°33'N, 1 13°12'E; captive purchased in 1979 by district purchasing agent (Ling Wenfeng, county forest officer, photos and pers. comm., 10 Nov. 1985). C:C-203. Shanxi (= Shansi), southeastern, CHINA; ca. 35°20'N, 112°50'E; reported before 1942 by A. de C. Sowerby (1941, p. 261). Reported before 1922 by Jiang Xuelong, Wang Yingxiang. and Ma Shilai (1991, pp. 242. 244). C:C-6. Shaowu Xian: Fujian, CHINA; ca. 27°I8'N, 117°30'E; reported June 1983 by Zheng Xue- qing (1984, p. 145). C:C-80. Sha Xian; Fujian, CHINA; ca. 26°25'N, 1 17°45'E; collected 22 Sept. 1960 by unknown collector; sciea, 1 (skin only). Reported Apr. 1981 by Zheng Xueqing (1984, p. 145). C:C- 82. Shennongjia Forestry Region; Hubei, CHINA; ca. 31°44'N, 110°44'E; reported before 1980 by XiaoZhi (1979. p. 31). C:C-43. Shenzhen Shi. See Neilingding Dao. Sheo. Barmer District; Rajasthan, INDIA; 26°irN, 7ri5'E; solitary individual reported ca. 1980 by T. R. Parmar (Bhargava. 1984, p. 43). A:I-79. Shexian; Anhui, CHINA; 29°52'N, 118°26'E; re- ported before 1998 (Zhang et al.. 1997. p. 58). C:C-61. Shihshahshu Temple; Sichuan, CHINA; ca. 29°32'N, 103°2rE; reported 4-8 Oct. 1929 by local residents (Stevens, 1934, p. 222; possibly misidentified M thihetana). C:C-139. Shimen. Qimen Xian. 300-500 m; Anhui, CHI- NA; ca. 29°55'N. 117°45'E; observed 1973- 1986 by Xiong Chenpei (Wada et al., 1986, p. 83). C:C-62. Shimla. See Simla. Shingaw. See Tanga-Shingaw. Shing Mun Country Park; Xianggang (= Hong FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 165 Kong), CHINA; ca. 22°23'N, 114°08'E; report- ed before 1992 by J. R. Fellowes (1992, p. 131). C:C-210. Shiqian; Guizhou, CHINA; 27°30'N, 108°I4'E; reported before 1998 (Zhang et al., 1997 p. 58). C:C-119. Shishi Koh vicinity, Chitral District; North-We.st Frontier, PAKISTAN; ca. 35°35'N, 71°48'E; reported before 1978 by T. J. Roberts (1977, p. 86). A:P-1. Shiva. See Siva. Shiwan Dashan Water Regulation Forest Reserve; Guang.xi, CHINA; ca. 2r49'N, 108°00'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C-217. Shocheng. See Shoucheng. Shogran vicinity, lower Kaghan Valley; North- west Frontier, PAKISTAN; ca. 34°37'N, 73°28'E; reported before 1978 by T J. Roberts (1977, p. 86). A:P-10. Shoucheng (= Shocheng) Water Regulation For- est Reserve; Guang.xi, CHINA; ca. 25°16'N, 109°47'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127; letter, Aug. 1996). C:C-183. Shuangbai; Yunnan, CHINA; 24°40'N, 101°38'E; reported before 1998 (Zhang et al., 1997, p. 58). B:C-51. Shuanyuan, Anyuan Xian; Jiangxi, CHINA; ca. 25°30'N, 115°15'E; reported Oct. 1979 by local residents (Liu Zhenhe, sciea, pers. comm., 25 Nov. 1985). C:C-99. Shuicheng Xian vicinity; Guizhou, CHINA; ca. 26°50'N, 105°00'E; reported before 1989 by Tan Bangjie and E E. Poirier ([1991], p. 131). C:C-144. Sibsagar. See Golaghat. Sichuan, CHINA; 26°-34°N, 98°-110°E; reported origin of captive shipped from Shanghai ca. Jan. 1868 (Gray. 1868, p. 61); bm(nh), 1 (ho- lotype of Macacus lasiotus). Reported origin of specimen obtained before 2 Oct. 1917 via Qing- dao (= Tsingtau) by Assesor Cramer; zmb, 1. Immunological survey conducted before 1996 by Duan Xingsheng, Liu Yuanwei, Wu Jing, Dao Weiying, and Liu Jianghai (1995, p. 411). Not mapped. Sichuan, eastern, CHINA; 28°-33°N, 106°-I10°E; tissue samples obtained ca. 1991 by Zhang Ya- ping and Shi Liming (1993b, p. 589). Not mapped. Sichuan, northwestern, CHINA; 26°-34°N, 98°- 103°E; tissue samples obtained ca. 1991 by Zhang Yaping and Shi Liming (1993, p. 589). Not mapped. Sichuan, west-central, CHINA; 28°-32°N, 99°- I02°E; tissue sample obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). Not mapped. Siddeldar Hill, Nagarjunakonda Valley, 2 mi ( = 3 km) northeast of Pullareddigudem; Andhra Pradesh, INDIA; 16°33'N, 79°16'E; collected 2 Nov. 1963 by B. Nath (Chaturvedi, [1973], p. 17; letter, 29 Sept. 1978; Subrahmanyam, 1975, p. xvii; Agrawal & Bhatiacharyya, 1976, p. 213); zsi, 1 (skin only). A:I-129. Sijian Shan Water Regulation Forest Reserve; Guangxi, CHINA ca. 25°12'N, 108°57'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C-180. Sikarwar. See Khair, Tahsil. Sikkim, INDIA; ca. 27°10'N, 88°30'E; collected before 1892 by [L. Mandelli] (Blanford, 1888b, p. 14); BM (NH), 2 (skins only, 1 with skull in- side). B:I-7. Siliguri; West Bengal, INDIA 26°42'N, 88°26'E; observed in 1962 by C. H. Southwick, A. Ghosh, and C. D. Louch (1964, p. 444). B:I-6. Simao Xian; Yunnan, CHINA; ca. 22°46'N, 101°05'E; tissue samples obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). Immunological survey conducted before 1996 by Duan Xingsheng, Liu Yuanwei, Wu Jing, Dao Weiying, and Liu Jianghai (1995, p. 411). B:C-78. Simla, western suburb, 1800-2200 m; Himachal Pradesh, INDIA; 31°06'N, 77°10'E; observed Aug. 1972-Jan. 1973 by Y Sugiyama (1976, p. 262). A:M8. Simla District; Himachal Pradesh, INDIA; 30°45'-31°10'N, 76°55'-77°15'E; observed 1981-1983 by R K. Seth, S. Seth, G. L. Reddy, and P K. Chopra (1992, p. 62). Not mapped (see A:I-18). Simla vicinity, 8000 ft (= 2440 m); Himachal Pradesh, INDIA; ca. 3r06'N, 77°10'E; ob- served May 1911 by P T L. Dodsworth (1914, p. 730). Collected 5 Sept. 1913 by P T L. Dod- sworth (1914, p. 730); bnhs, 1 (skull only). Ob- served Aug. 1972-Feb. 1973 by K. Wada (1984, p. 477). Observed Dec. 1981-Feb. 1982 by A. Camperio Ciani (1983, p. 275; 1984, p. 372). Observed Apr.-May 1991 by C. Ross, A. Srivastava, and R. S. Pirta (1993, p. 160). A:I- 18. Simla Water Catchment Reserve; Himachal Pra- 166 FffiLDIANA: ZOOLOGY desk, INDIA; ca. 31°05'N, 77°10'E; reported 1978-1980 by A. J. Gaston, P. J. Garson, and M. L. Hunter, Jr. (1983, p. 300). A:I-18. Simri. Birganj Forest District; Rautahat, NEPAL; 27°06'N, 85°14'E; observed June 1964-Dec. 1965 by D. L. Chesemore (1970, p. 164). A:N- 13. Simri, Narayani River, Rapti Valley; Chitawan, NEPAL; 27°36'N, 84°19'E; observed June 1964-Dec. 1965 by D. L. Chesemore (1970. p. 164). Reported at Chitwan National Park before 1997 by K. K. Gurung and R. Singh (1996, p. 84). A:N-8. Simulbari-Pankhabari, highway between; West Bengal. INDIA; ca. 26°48'N 88°18'E; observed Mar.-Apr. 1985 by R. P Mukherjee, S. Chau- dhuri. and A. Murmu (1995, p. 27). B:I-6. Sindholi (= Sindhauli); Uttar Pradesh, INDIA; 27°52'N, 78°07'E; observed 1959-1975 by C. H. Southwick and M. E Siddiqi (1977, p. 342). A:I-69. Singaul, Tekan, Birganj Forest District; Rautahat, NEPAL; 27'10'N, 85°20'E; observed June 1964-Dec. 1965 by D. L. Chesemore (1970, p. 164). A:N-13. Singaw. See Tanga-Shingaw. Singhbhum. See Luia. Singkaling Hkamti, upper Chindwin River; Ka- chin, MYANMAR (= BURMA); ca. 26°00'N, 95°42'E; collected June-Aug. 1914 by G. C. Shortridge (in Wroughton, 1916a, p. 293). BM(NH), 3 (including 2 skulls only). B:M-8. Singkaling Hkamti, upper Chindwin River, 500 ft (=150 m); Kachin, MYANMAR (= BURMA); ca. 26°00'N, 95°42'E; collected 5 Aug. 1914 by G. C. Shortridge and S. A. Macmillan (in Wroughton, 1916a, p. 293); bnhs, 1. B:M-8. Singkaling Hkamti, upper Chindwin River, left (east) bank, 500 ft (= 150 m); Kachm, MYAN- MAR (= BURMA); ca. 26°00'N, 95°42'E; col- lected 24 July 1914 by G. C. Shortridge and S. A. Macmillan (in Wroughton, 1916a, p. 293); zsi, 1. B:M-8. Singkaling Hkamti, upper Chindwin River, right (west) bank; Kachin, MYANMAR (= BUR- MA); ca. 26°00'N, 95°42'E; collected 8 Mar. 1935 by H. C. Raven (in Carter, 1943, p. 100; Morris, 1936, p, 662); amnh, 1. B:M-8. Singolo. See Xi Golog. Sita Bani, Ramnagar vicinity, Kumaun region. 2000 ft (= 600 m); Uttar Pradesh, INDIA; 29°24'N, 78°13'E; collected 22 Nov. 1913 by C. A. Crump (in Wroughton, 1914, p. 283); bm(nh), 2. A:I-34. Sitakunda; Chittagong, BANGLADESH; 22°37'N, 91°39'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-32. Sitapahar/Rampahar; Rangamati, BANGLA- DESH; ca. 22°30'N, 92°15'E; reported early in 1980 by S. P Gittins and A. W. Akonda (1982, p. 278). B:Ba-35. Sitapur-Bareilly, highway between; Uttar Pra- desh, INDIA; ca. 27°50'N, 80°00'E; observed 28 Oct.-29 Dec. 1964 by C. H. Southwick. D. Lindburg. M. Neville. P. Jay. and R. P. Mu- kherjee (Southwick and M R. Siddiqi. 1966. p. 306). A:I-54. Sitapur-Shahjahanpur. highway between; Uttar Pradesh, INDIA; ca. 27°40'N. 80°20'E; ob- served 1964-1965 by R. P Mukherjee and G. D. Mukherjee (1972, p. 67). A:I-67. Sittang River. See Toungoo, 15 mi. (= 24 km) north of. Siva (= Shiva); Dhankuta, NEPAL; 27°28'N, 87°10'E; observed in 1997 by M. K. Chalise and M. Ghimire (1998, p. 12). B:N-supplemen- tary. Sivalik Hills. See Siwalik Range. Sivok; West Bengal, INDIA; 26°52'N, 88°27'E; collected 11 Nov. 1930 by H. Stevens; fmnh, 1 (skin only). B:I-6. Sivok, ca. 3 km east of; West Bengal, INDIA; ca. 26°52'N, 88°30'E; observed Mar.-Apr. 1985 by R. P. Mukherjee, S. Chaudhuri, and A. Murmu (1995, p. 27). B:I-6. Sivok, ca. 5 km east of; West Bengal, INDIA; ca. 26°52'N, 88°31'E; observed Mar.-Apr. 1985 by R. P. Mukherjee, S. Chaudhuri, and A. Murmu (1995, p. 27). B:I-6. Sivok, ca. 6 km east of; West Bengal, INDIA; ca. 26°52'N, 88°32'E; observed Mar.-Apr. 1985 by R. P. Mukherjee. S. Chaudhuri. and A. Murmu (1995, p. 27). B:I-6. Siwalik Range; Himachal Pradesh, INDIA; 30°20'-32°10'N, 75°40'-77°40'E; observed 9 Aug.-13 Sept. 1974 by M. Singh (1975, p. 472). Not mapped. Sohagpur, Hoshangabad District, 1000 ft (= 300 m); Madhya Pradesh, INDIA; 22°42'N, 78°12'E; collected 10 Apr. 1912 by C. A. Crump (Wroughton & Ryley, 1913, p. 45); bm(nh), 1 (skin only). A:I-95. Solon (= Solan) District; Himachal Pradesh, IN- DIA; ca. 30°55'N, 77°07'E; observed 1981- 1983 by P K. Seth, S. Seth, G. L. Reddy, and P K. Chopra (1992, p. 62). A:I-I8. Sonargaon; Narayanganj, BANGLADESH; FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MAC AC A MULATTA 167 23°39'N, 90°37'E; reported in 1971 by J. R. Op- penheimer, A. K. Akonda. and K. Z. Husain (1983, p. 194). B:Ba-27. Sonepat District; Hatyana, INDIA; ca. 28°59'N, 77°0rE; observed 1981-1983 by P. K. Seth, S. Seth, G. L. Reddy, and P. K. Chopra (1992, p. 62). A:I-36. Songtao; Guizhou, CHINA; 28°12'N, 109°12'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-114. Song-Ta-Voy (= Song-Ta-Voi); Quang Nam-Da Nang, VIETNAM; ca. 16°10'N, 107°40'E; collected [?Jan. 1899] by R E S. Barthelemy (1904, p. 38); mnhn, 1. C:V-34. Songxi Xian; Fujian, CHINA; ca. 27°36'N, 118°47'E; reported Oct. 1980 by Zheng Xue- qing (1984, p. 146). C:C-74. Son La- VIETNAM; ca. 20°35'-22°05'N, 103°I5'-105°00'E; collected before 1986 by unknown collector (Dao, 1985, p. 166); muse- um unknown (not seen). Not mapped. Son Tra. Mt.; Quang Nam-Da Nang, VIETNAM; ca. 16°07'N, 108°18'E; collected 20 Aug. 1965 at >400 m by P. F. Ryan (Van Peenen et al., 1968, p. 609; 1971, pp.127, 134; Fooden, 1995, p. 25); USNM, 1 (skull only, external measure- ments in collector's fieldbook). Collected 19 May 1966 by J. T. Lowery (Van Peenen et al., 1968, p. 609; 1971, pp. 127, 134; Fooden, 1995, p. 25); usnm, 1 (skull only, mandible missing). Reported ca. 1990-1995 by L. K. Lippold (1995, p. 199; identified as M. fasci- ciilahs). C:V-36. Son Tra, Mt., 3.9 km west and 0.3 km south of; 240 m; Quang Nam-Da Nang, VIETNAM; ca. 16°07'N, 108°15'E; collected 14 Sept. 1967 by R F D. Van Peenen (Van Peenen et al., 1968, p. 609; 1971, pp. 127, 134; Fooden, 1995, p. 25); USNM, 1 (external measurements question- able). C:V-36. South China. See CHINA, South. South District, north-central; Tripura, INDIA ca. 23°45'N, 91°35'E; observed in 1976 and 1978 by R. R Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). 8:1-40. South District, northeastern; Tripura, INDIA; ca. 23°35'N, 91°55'E; observed in 1976 and 1978 by R. R Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). 3:1-42. South District, south-central; Tripura, INDIA; ca. 23°05'N, 91°40'E; observed in 1976 and 1978 by R. R Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). 3:1-40. South District, southeastern; Tripura, INDIA; ca. 23°05'N, 91°48'E; observed in 1976 and 1978 by R. R Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). B:I-40. South District, southwestern; Tripura, INDIA; ca. 23°00'N, 9r35'E; observed in 1976 and 1978 by R. R Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). 3:1-40. South District, west-central; Tripura, INDIA; ca. 23°20'N, 91°30'E; observed in 1976 and 1978 by R. R Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). 3:1-40. South Kamrup. See Kulsi; Rajapara. "Southwest" Yunnan. See Yunnan [northwestern]. Srimangal Tea Estate; Moulvi Bazar, BANGLA- DESH; ca. 24°19'N, 91°44'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). 3:3a- 13. Srimangal vicinity; Moulvi Bazar, BANGLA- DESH; ca. 24°19'N, 91°44'E; tentatively re- ported July-Nov. 1976 by K. M. Green (1978, p. 146). B:Ba-13. Srinagar, ca. 22 km southeast of. See Ovra ( = Overa) Sanctuary. Suifu. See Yibin. Suiyang; Guizhou, CHINA; 27°57'N, 107°11'E; reported before 1988 (Zhang et al., 1997, p. 58). C:C-130. Sucktaisgur. See Saktesgarh. Sukla Phanta; Kanchanpur, NEPAL; 28°50'N, 80°11'E; observed June 1964-Dec. 1965 by D. L. Chesemore (1970, p. 164). Reported before 1997 by K. K. Gurung and R. Singh (1996, p. 118). Observed in 1998 by M. K. Chalise and M. Ghimire (1998, p. 12). A:N-2. Sukna, Darjeeling District; West Bengal, INDIA; 26°47'N, 88°22'E; collected 25 Apr. 1892 by W. Partridge; zsi, 1. Observed in 1962 by C. H. Southwick, A. Ghosh, and C. D. Louch (1964, p. 446). Observed Mar.-Apr. 1985 by R. R Mu- kherjee, S. Chaudhuri. and A. Murmu (1995, p. 27). 3:1-6. Sukna-Kurseong, highway between; West Bengal, INDIA; ca. 26°48'N, 88°21'E; observed Mar.- Apr. 1985 by R. R Mukherjee, S. Chaudhuri, and A. Murmu (1995, p. 27). 3:1-6. Sultanpur vicinity; Uttar Pradesh, INDIA; ca. 26°16'N, 82°04'E; blood samples obtained 16- 168 FIELDIANA: ZOOLOGY 27 Apr. 1964 by K. V. Shah and C. H. South- wick (1965, p. 489). A:I-64. Sumera; Uttar-Pradesh. INDIA; ca. 28°02'N, 78°09'E; observed Jan. 1990-Mar. 1991 by E. Imam and H. S. A. Yahya (1995, p. 2). A:I-69. Sumera Fall Jungle; Uttar Pradesh. INDIA; ca. 28°02'N, 78°09'E; observed 1959-ca. 1980 by C. H. Southwick and M. E Siddiqi (1977, p. 342; 1984, p. 559; Siddiqi & Southwick, 1980, p. 54; 1988, p. 121). A:I-69. Sundarbans (= Sunderbans; Sunderbunds); Khiil- na, BANGLADESH; ca. 22°00'N, 89°30'E; ob- served 29 Jan.-21 Apr. 1971 by H. Hendrichs (1975, p. 177). Tentatively reported July-Nov. 1976 by K. M. Green (1978, p. 146). Reported before 1978 (Anonymous, 1977, p. 14). Ob- served early in 1980 by S. P. Gittings and A. W. Akonda (1982, p. 278). Observed in 1980 and 1982 by M. A. R. Khan and M. F Ahsan (Khan, 1985, p. 31; 1986, p. 37). Observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, pp. 76, 79). Reported be- fore 1997 by K. K. Gurung and R. Singh (1996, p. 122). B:Ba-23. Sundarbans (= Sunderbans, Sunderbunds); West BengoL INDIA; 88°00'-89°00'N, 2I°30'- 22°30'E; reported in 1892 by E. de Poncins (1935, p. 846). Reported before 1997 by K. K. Gurung and R. Singh (1996, p. 122). Not mapped (see B:I-1 and B:I-2). Sundarbans (= Sunderbunds), ca. 50 mi (= 80 km) east of Calcutta; Satkhira, BANGLA- DESH; ca. 22°35'N, 89°15'E; collected 26 Apr. 1870 by museum collector (Anderson, 1872, p. 529); zsi 7 (including 1 skin only). B:Ba-21. Sungri, ca. 2 km south of; Himachal Pradesh, IN- DIA; ca. 31°18'N, 77°42'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984, p. 476). A: 1-17. Sungri, ca. 4 km north of; Himachal Pradesh, IN- DIA; ca. 31°2rN, 77°42'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984. p. 476). A: 1-17. Sungri, ca. 6 km northwest of; Himachal Pradesh, INDIA; ca. 31°21'N, 77°39'E; reported Aug. 1972-Feb. 1973 by K. Wada (1984, p. 476). A: 1-17. Sungri. ca. 10 km northwest of; Himachal Pra- desh, INDIA; ca. 31°23'N, 77°38'E; reported Aug. 1972-Feb. 1973 by K. Wada (1984, p. 476). A:I-17. Sungri, ca. 10 km southwest of; Himachal Pra- desh, INDIA; ca. 31°14'N. 77°37'E; reported Aug. 1972-Feb. 1973 by K. Wada (1984, p. 476). A:I-17. Sungri, ca. 15 km southwest of; Himachal Pra- desh, INDIA; ca. 3I°14'N. 77°33'E; reported Aug. 1972-Feb. 1973 by K. Wada (1984. p. 476). A:I-17. Sungri, ca. 20 km south-southwest of; Himachal Pradesh, INDIA; ca. 31°08'N. 77°38'E; ob- served Aug. 1972-Feb. 1973 by K. Wada (1984, p. 476). A:I-17. Sungri, north of; Himachal Pradesh, INDIA; ca. 31°19'N, 77°42'E; observed Aug. 1972-Feb. 1973 by K. Wada (1984, p. 476). A:I-I7. Suoxi Valley (= Suoxiyu), Cili Xian, 1256 m; Hu- nan, CHINA; ca. 29°55'N, 110°50'E; reported before 1986 by Di Chen (1985. p. 33). Ob- served 1988-1990 by Feng Min, Jiang Haish- eng, and Wang Jun (1997, p. 25). C:C-II0. Surat. See Dangs District. Surinsar, Samba Subdistrict; Jammu & Kashmir INDIA; ca. 32°34'N, 75°07'E; observed before 1983 by Y. R. Malhotra and D. N. Sahi (1982, p. 27). A:I-7. Suritola. See Dhaka. Swat Khoistan region; North-West Frontier, PAKI- STAN; ca. 35°35'N. 72°30'E; reported before 1978 by T J. Roberts (1977, pp. 86. 87). A:P-3. Swat River; North-West Frontier, PAKISTAN; ca. 34°20'N, 71°35'E; blood samples collected ca. 1978-1979 by D. J. Melnick, C. J. Jolly, and K. K. Kidd (1986, p. 129). A:P-5. Swat Valley, lower. See Bar Chanrai Hill. Swayambhunath, 4400 ft (= 1300 m); Katmandu Valley, NEPAL; ca. 27°43'N, 85°18'E; ob- served June 1964-Dec. 1965 by D. L. Chese- more (1970, p. 164). Observed 1974-1975 by H. Taylor, J. Teas, T. Richie, C. Southwick. and R. Shrestha (1978, p. 344).Observed 1975- 1978 by J. Teas (1983. p. 224). Reported before 1982 by T K. Shrestha (1981, p. 269). Ob- served summer 1982 by R. L. Johnson and C. H. Southwick (1984. p. 201). Observed 1995- 1998 by M. K. Chalise and M. Ghimire (1998, p. 11). A:N-12. Sylhet Forest Division; Moulvi Bazar, BANG- LADESH; ca. 24°25'N, 92°00'E; reported be- fore 1982 by M. A. R. Khan (1981, p. 13). B: Ba-13. Szechuan. See Sichuan. Szechuen. See Sichuan. Tachienlu. See Kangding. Ta Chang Tai. See Tha Chang Tai. Taga Hka, Chindwin River, west bank; Kachin, MYANMAR (= BURMA); 26°2rN, 96°09'E; FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 169 collected 1 1 Feb. 1935 by R. C. Morris and C. McCann (Morris. 1936. pp. 653. 655; H. C. Ra- ven in Carter. 1943. p. 100); .amnh. 2. B:M-7. Taihang Shan; Hemm, CHINA; ca. 35°05'N, 112°24'E; observed 1988-1995 by Fang Bao- hua, Xu Xinjie. and Liu Bingxu (1995. p. 355; cf. Chen et al.. 1988. p. 25). C:C-4. Taihe; Jiang.xi. CHINA; 26°48'N. 114°56'E; re- ported before 1998 (Zhang et al.. 1997. p. 58). C:C-105. Taining. Wuyishan; Fiijian, CHINA; 26°55'N. 117°12'E; reported before 1998 (Zhang et al.. 1997, p. 58). C:C-81. Taining Xian; Fujian, CHINA; ca. 26°55'N. 117°12'E; reported May 1981 by Zheng Xue- qing (1984. p. 145). C:C-81. Tai Po Kau Nature Reserve: Xianggang (= Hong Kong). CHINA; ca. 22°25'N. 114°10'E; report- ed before 1992 by J. R. Fellowes (1992. p. 131). C:C-210. Tai Tarn Reservoir; Xianggang (= Hong Kong), CHINA; ca. 22°I4'N, I14°13'E; reported in 1947 by G. A. C. Herklots (1951. p. 83). C:C- 210. Takerhat. See Dhaka. Tallada, Khammam District, 100 m; Andhra Pra- desh, INDIA; 17°13'N, 80°25'E; observed 21 Apr. 1980 by J. Fooden. A. Mahabal. and S. S. Saha (1981. p. 467). A:I-132. Tai Vraksh. Tahsil Kushalgarh; Rajasthan, IN- DIA; ca. 23°10'N. 74°27'E; observed 1975- 1980 by P. K. Seth. S. Seth, and A. K. Shukla (1983. p. 38). A:I-97. Tamanthe. See Hisweht. Tamanthi Wildlife Sanctuary; Sagaing, MYAN- MAR (= BURMA); 25°26'N, 95°37'E; ob- served 2-14 Mar. 1994 by A. Rabinowitz, G. B. Schaller. and U Uga (1995. p. 125). B:M-1 1. Tanbazar. See Narayanganj. Tang Hpre (= Tang Hper); Kachin, MYANMAR (= BURMA); ca. 25°23'N. 97°14'E; collected 14 Oct. 1945 by K. E. Stager (letter, 9 Aug. 1985); LSNM. 1. B:M-5. Tanga-Shingaw (= Tang-Singaw). road between. 800 ft (= 240 m); Kachin, MYANMAR ( = BURMA); ca. 25°40'N. 97°55'E; collected 9 Apr 1939 by H. E. Anthony (1941. pp. 55. 83); AMNH, 1. B:M-4. Tangxi, Guichi Xian, 200-560 m; Anhiii, CHINA; ca. 30°20'N, 117°40'E; observed 1973-1986 by Xiong Chenpei (Wada et al.. 1986. p. 83). C:C- 50. Tanikella. Khammam Taluk, Khammam District; Andhra Pradesh, INDIA; 17°15'N, 80°15'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992. p. 19). A:l-132. Tanti Road. See Dhaka. Tara Devi. See Nhera/Tara Devi. Tarai. See Terai. Tarap; Moidvi Bazar, BANGLADESH; ca. 24°10'N, 91°35'E; observed early in 1980 by S. P Gittins and A. W. Akonda (1982. p. 278). B: Ba-13. Tarkhola. Darjeeling District. 325 m; West Ben- gal, INDIA; 27 WN, 88°33'E; observed June- Aug. 1958 by H. Khajuria (1966, p. 284; Kha- juria & Ghose. 1970. p. 17). B:I-7. Taro (?= Dalu); Kachin, MYANMAR (= BUR- MA); 26°2rN. 96°irE; collected 5 Feb. 1935 by H.C. Raven (in Carter, 1943, p. 100; Morris, 1936, p. 653); amnh, 2. B:M-7. Tasin Lou. See Kangding. Tat Ke vicinity (= Kheo Ting-Ta Ke area). Na Hang District; Tiiyen Quang, VIETNAM; ca. 22°25'N. 105°25'E; captured ca. 1992 by local resident (Ratajszczak et al.. 1992. p. 14). C:V-6. Tatkon. near Kindat. east bank of Chindwin River. 250 ft (= 75 m); Sagaing, MYANMAR ( = BURMA); ca. 23°47'N, 94°30'E; collected 5 July 1914 by G. C. Shortridge and S. A. Mac- millan (Shortridge in Wroughton. 1916a. p. 293); BNHS, 1. B:M-18. Tatkon, near Kindat, west bank of Chindwin Riv- er, 250 ft (= 75 m); Sagaing, MYANMAR ( = BURMA); 23°47'N. 94°29'E; collected 28 June and 5 July 1914 by G. C. Shortridge (in Wroughton. 1916a, p. 293); bm(nh), 2 (includ- ing 1 skin only); bnhs. 2 (including I skin only). B:M-18. Ta-tsien-lou. See Kangding. Tatura. Chandigarh; Punjab, INDIA; not precisely located, 30°38'-30°47'N, 76°43'-76°53'E; re- ported 1964-1966 by D. G. Lindburg (1977a. p. 268). Not mapped. Tay Tru. See Trai Tru. Tche-ly. eastern mountains. See Xinglong Xian. southern. Tehri-Garhwal District; Uttar Pradesh, INDIA; ca. 30°23'N, 78°29'E; observed 1981-1983 by P K. Seth, S. Seth, G. L. Reddy. and P K. Cho- pra (1992. p. 62). A:L28. Teknaf Peninsula; Cox's Bazar, BANGLADESH; 20°52'N. 92°18'E; observed Feb. 1990-June 1993 by M. M. Feeroz. M. A. Islam, and M. Kabir (1995. p. 76). B:Ba-40. Tenali. 5.5 km west of. Guntur District. 10 m; Andhra Pradesh, INDIA; 16°14'N, 80°37'E; 170 FIELDIANA: ZOOLOGY observed 10 May 1980 by J. Fooden. A. Ma- habal. and S. S. Saha (1981, p. 468). A:I-143. Tenasserinn MYANMAR (= BURMA); 10°- 15°N. 98°-100°E; obtained 4 Oct. 1912 by Sch. Med. Rat. Donitz; locality information probably inaccurate (see above. Fig. 2B); zmb, 1 (skull only). Not mapped. Tengchong (= Momien); Yunnan. CHINA; 25°02'N. 98°28'E; captive purchased Man-July 1868 by J. Anderson (1876, pp. 190, 247; 1879, pp. xvi, 56); zsi, 1 (skin only). Collected 5 Jan. 1961 by Zhou Jiadi; IZCAS, 1 (skin only). B:C- 59. Tengchong Xian; Yunnan, CHINA; ca. 25°02'N. 98°28'E; collected before 1984 by Ma Shilai; Kiz, 1 (skull only). B:C-59. Tengri-Nor. See Nam Co. Teng-yue-chow. See Tengchong. Teng-yueh. See Hui-yao. Ten Ky. See Nghia Dung. Terai (region); NEPAL; 26°-29°N, 80°-88°E; re- ported before 1842 by B. H. Hodgeson (1841, p. 1212; cf. Karan, 1960, p. 92). Not mapped. Tha Chang Tai (= Ta Chang Tai), 600 ft (= 180 m); Tak. THAILAND; 16°51'N, 99°03'E; col- lected 14 July 1924 by J. H. Chambrai (Kloss, 1930, p. 62); zrc, 1 (excludes zrc 4-822, a co- lobine: cf. Weitzel et al., 1988, p. 116). B:T-7. Thai Nguyen; Bac Thai. VIETNAM; ca. 21°36'N, 105°50'E; collected 30 Dec. 1956 and 17 June 1959 by unknown collectors; zmvnu, 2 (skulls only). C:V-15. Thana Ghazi. See Bandipul. Thanh Son; Vinh Phu, VIETNAM; 21°13'N, 105°irE; collected 27 June 1961 by unknown collector; zmvnu, 1 (skull only). C:V-18. Thanh Tuong, Na Hang District; Tuyen Quang. VIETNAM; 22°19'N, 105°24'E; collected 18 Jan. 1965 and 23 Octt. 1965 by Ma Van Dam (Dao, 1985, p. 29; external measurements pub- Ushed); iebr, 2. C:V-6. Thapathali. See Tripureswor. Tharikella. See Tanikella. Thateng. See Muang Thateng. Theme. See Pye (= Prome), 35 mi (= 55 km) southeast of. Thirumunidevipetta. See Tirumaladevipeta. Thuong Bang La, Van Chan District; Yen Bai. VI- ETNAM; 2r25'N, 104°47'E; collected Mar. 1963 by unknown collector (Dao, 1985, pp. 183, 192); lEBR, 1 (skull only). C:V-17. Tian'e Xian; Guangxi. CHINA; ca. 25°00'N, 107°10'E; purchased at traditional medicine shop 15 Oct. 1992 by Quan Guoqiang (cf. Fooden et al., 1994, p. 623); izcas, 3 (skulls only). C:C-173. Tiantang Shan Water Regulation Forest Reserve; Guangxi, CHINA; ca. 22°36'N. 11()°42'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter, Aug. 1996). C:C-214. Tibet; Sichuan or Xizang (= Tibet). CHINA; 30°- 32°N, 96.5°-102°E; collected [May-July 1890] by G. Bonvalot and H. d'Orleans (Bonvalot, 1891, vol. 2, p. 210; 1892, p. 505); possibly purchased at Ngamda ('?= Kintachie/Houmda) and/or collected at Rouetoundo (see above; Bonvalot, 1891, vol.2, pp. 149, 156); mnhn, 2 (skins only). Not mapped. Tibet Colony. See Dehra Dun vicinity. Tin Toe Forest, Moc Chau Plateau, 900 m; Son La, VIETNAM; ca. 20°50'N, 104°46'E; col- lected 20 Jan. 1970 by Le Vo Dinh Tuong (Dao, 1978, pp. 378, 382); museum unknown. 2 (not seen). C:V-19. Tinsukia District (Bherjan, Borajan, Podumoni Reserved Forests); Assam. INDIA; ca. 27°30'N, 95°22'E; observed Sept.-Nov. 1995 by A. Choudhury (1997, p. 10). B:I-26. Tipomia; Assam. INDIA; ca. 27°00'N, 94°48'E; reported 9 Mar. 1987-16 Feb. 1988 by A. Choudhury ([1991a], p. 31). B:I-25. Tipusultan Road. See Dhaka. Tirthan; Himachal Pradesh. INDIA; ca. 31°40'N. 77°30'E; reported 1978-1980 by A. J. Gaston, P J. Garson, and M. L. Hunter, Jr. (1983, p. 300). A:I-16. Tirumaladevipeta, Chintalapudi Taluk. West Go- davari District; Andhra Pradesh. INDIA; 17°05'N, 81°09'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). A:I-137. Tongjiang Xian; Sichuan. CHINA; ca. 3r55'N, I07°I3'E; collected July 1966 by Expedition of Biological and Agricultural Resources in Qin- ling, Shaanxi; siz. 4. C:C-31. Tong Kou; Anhui. CHINA; ca. 30°05'N. 1 18°10'E; collected 23 May and 21 July 1959 by Wang Zeyeh; izcas, 2 (including 1 skin only). C:C-62. Tong-lin, mountains of. See Xinglong Xian. southern. Tongzhi. See Tongzi. Tongzi (= Tongzhi); Guizhou. CHINA; 28°08'N, 106°49'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-135. Tonkin region; VIETNAM; 20°-23°N, 102°- 108°E; captive obtained in 1886 by Lt. Stahl; FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 171 died in zoo 26 Jan. 1887; mnhn, 1 (skin only). Not mapped. Toungoo. 13 mi (= 21 km) east of, 500 ft (= 150 m); Karen. MYANMAR (= BURMA); ca. 18°55'N. 96°40'E; collected 2 Sept. 1927 by J. M. D. Mackenzie (in Fry, 1929, p. 637); bm (nh), 1; BNHS. 1. B:M-31. Toungoo. 15 mi (= 24 km) north of, east side of Sittang River, 400 ft (= 120 m); Karen, MYANMAR (= BURMA); ca. 19°10'N, 96°25'E; collected 26 Jan. 1927 by J. M. D. Mackenzie (Fry, 1928, P. 545); bm(nh), 1. B: M-30. Toungoo, 20 mi (= 32 km) west of, 500 ft (= 150 m); Pei^u, MYANMAR (= BURMA); ca. 18°55'N, 96°08'E; collected 22 Feb. 1929 by J. M. D. Mackenzie; bnhs, 1 (skin only). B:M-33. Toungoo, 30 mi (= 48 km) northwest of, 500 ft (=150 m); Pegu, MYANMAR (= BURMA); ca. 19°15'N, 96°05'E; collected 26 Nov. and 6 Dec. 1928 by J. M. D. Mackenzie (Khajuria, [1955], p. 113); bm(nh), 1; bnhs, 1; zsi, 1 (skin only). B:M-29. Toungoo, east side of Sittang River, 100 ft (= 30 m); Pegu, MYANMAR (= BURMA); ca. 18°56'N, 96°27'E; collected 15 May 1927 by J. M. D. Mackenzie (in Fry, 1929, p. 637); BM(NH), 3. B:M-32. Trai Tru (= Tay Tru), Huong Khe District; Ha r/«/2, VIETNAM; 18°11'N, 105°35'E; collected 6 Feb. 1964 by unknown collector (Dao. 1985, p. 233; external measurements published); lEBR, 1 (skull only). C:V-28. Tripureswor, Thapathali; Katmandu Valley, NE- PAL; ca. 27°43'N, 85°19'E; observed Dec. 1995 and Dec. 1998 by M. K. ChaUse and M. Ghimire (1998, pp. 12, 15). A:N-supplementa- ry. Trisuli Bazar, 4 mi (= 6.5 km) southeast of, Na- wakot District, 1875 ft (= 570 m); Bagmati, NEPAL; ca. 27°55'N, 85°10'E; collected 9 and 13 Apr. 1967 by C. O. Maser; FMNH, 6 (3 skeletons only, 3 in alcohol); ups, 2 (in alcohol, not seen). A:N-1 1. Trung Khanh District, Cao Bang, VIETNAM; ca. 22°50'N, 106°3rE; collected 9 May 1967 by Hoang Tung; FCXM, 1 (skull only, mandible missing). C:V-9. Tsari Chu (= Tsari Valley); Xizang (= Tibet), CHINA; ca. 28°45'N, 93°10'E; observed Sept.- Oct. 1913 by F M. Bailey (1914, map; 1915, p. 74). B:C-5. Tseo-Jia-Geo; Sichuan, CHINA; ca. 28°18'N, 104°12'E; collected 14 Jan. 1931 by D. G. Gra- ham (Moore & Tate, 1965, p. 334); usnm, 1. C: C-138. Tsingtau. See Sichuan. Tsung he. See Luofu Shan. Tu Chi; VIETNAM; not located, 15°-23°N, 102°- 109°E; date and collector unknown; zmvnu, 1 (skull only). Not mapped. Tughlaqabad (= Tughlakabad; Tukhlabad); Delhi, INDIA; ca. 28°30'N, 77°15'E; observed 1964- 1965, Dec. 1970-July 1972. Mar. 1980-Aug 1983. and July-Aug. 1987 by I. Malik, R K Seth, and C. H. Southwick (1984, p. 312 Southwick and M. R. Siddiqi, 1966, p. 309 Southwick et al.. 1976. p. 13; Malik. 1989a. p 118). Observed May 1987-Feb. 1989 by R. L Johnson, I. Malik, and C. M. Berman (1991, p 339). A:I-41. Tunchang Xian; Hainan Dao; Hainan, CHINA; ca. 19°22'N, 1 10°05'E; reported before 1986 by Xu Longhui and Liu Zhenhe (1985, p. 148). C: C-230. Tungho. See Wa Shan. Tunki, Gajwel Taluk, Medak District; Andhra Pradesh, INDIA; 17°45'N, 78°34'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-123. Tung Ling. See Xinglong Xian. southern. Tunxi; Anhui, CHINA; 29°43'N. 118°19'E; col- lected 23 Jan. 1959 by museum collector; smnh, 1 (skin only). C:C-61. Tusu River. See Hisweht. Tuzu River. See Hisweht. Twenty-four Parganas District; West Bengal, IN- DIA; 21°30'-22°20'N, 88°00'-89°00'E; ob- served 1951-1961 by A. K. Mandal (1964, R 165). Not mapped (see B:I-1 and B:I-2). U, Nam. See Ou, Nam. Udhampur; Jammu & Kashmir, INDIA; 32°56'N. 75°08'E; observed before 1983 by Y. R. Mal- hotra and D. N. Sahi (1982. p. 27). A:I-I0. Ukhia; Cox's Bazar, BANGLADESH; 2ri5'N, 92°08'E; observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995. p. 76). B:Ba-39. Umri Devi. Tahsil Ramgarh; Rajasthan, INDIA; ca. 27°15'N. 75°irE; Observed 1975-1980 by R K. Seth, S. Seth, and A. K. Shukla (1983, p. 38). A:I-76. Undrajapuram, Peravalli Block, East Godavari District; Andhra Pradesh, INDIA; not precisely located, 16°40'-17°50'N, 81°30'-82°35'E; re- ported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 19). Not mapped. United Khasi-Jaintia Hills; Meghalaya, INDIA; 172 FIELDIANA: ZOOLOGY ca. 25°30'N, 91°30'E; reported Dec. 1972-Feb. 1973 by R. L. Tilson (1983. p. 399). B:I-14. Um Pang. See Ban Umphang. Umpilagudem. See Dumpallagudem. Uppadhyay Mohalla. See Khair, Tahsil. Utzun Vicinity. Chitral District 5000 ft (= 1500 m); North-West Frontier, PAKISTAN; ca. 35°30'N, 71°40'E; reported 1901-1902 by H. Fulton (1903. p. 758). Reported before 1978 by T. J. Roberts (1977. p. 86). A:P-1. Uzipur. See Wazipur. Vaddeppalli. See Waddepalle. Van Canh. Dao; Quang Ninh. VIETNAM; ca. 20°52'N 107°22'E; collected 3 Apr. 1969 and June 1969 by unknown collectors; ikbr, 2 (skulls only). C:V-13. Van Hai. Dao; Quang Ninh, VIETNAM; 20°54'N. 107°30'E; collected Dec. 1961 by Xin Cua Dan (C. Groves, letter, 15 Dec. 1994) and unknown collector; zmvnu. 2 (skins only). Collected Dec. 1961 by Thanh, Khoi, and Cong; zmvnu, I (skull only). C:V-13. Varanasi (= Benares); Uttar Pradesh, INDIA; 25°20'N, 83°00'E; observed Sept. 1959-Feb. 1960 and 1964-1965 by C. H. Southwick, M. A. Beg, and M. R. Siddiqi (1961a, p. 543; Southwick & M. R Siddiqi. 1966, p. 312). Ob- served Aug. 1977-July 1978 by R. S. Pirta (1982, p. 401; Pirta & Singh, 1982, p. 15). Ob- served 2 Apr.-30 July 1980 and 15 Mar.-30 July 1981 by R. K. Singh (1984. p. 430). A:I- 87. Varanasi (= Benares) District; Uttar Pradesh, IN- DIA; 24°40'-25°30'N, 82°05'-83°30'E; cap- tives obtained in 1891 by W. Heape (1897, p. 135). Not mapped (see A:I-87). Vasunia, 3 km west of, Dangs District. 420 m; Gujarat, INDIA; 20°43'N, 73°38'E; observed 12 Jan. 1980 by J. Fooden, A. Mahabal, and S. S. Saha(1981, p. 466). A:I-99. Veer Sontri Forests. Kurukshetra District; Hary- ana, INDIA; 30°00'N. 77°0rE; observed Oct. 1989-Sept. 1990 by R. C. Gupta and S. Kumar (1992, p. 226). A:I-24. Velatur, Guntur District, 5 m; Andhra Pradesh, INDIA; 16°08'N, 80°52'E; observed 10 May 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 468). A:I-142. Velkicharia, Mahbubnagar District, 500 m; An- dhra Pradesh, INDIA; 16°37'N, 78°07'E; ob- served 9 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 468). A:I-126. Venketeswara Swami Temple, Aswaraopet Taluk, Khammam District; Andhra Pradesh, INDIA; ca. 17°15'N, 81°09'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992. p. 19). A:I-I37. Vergel. See Wargel. Verigedu. Peravalli Block. East Godavari District; Andhra Pradesh, INDIA; not precisely located, 16°40'-17°50'N, 81°30'-82°35'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984. p. 58; 1992. p. 19). Not mapped. Viangchan; Vientiane. LAOS; ca. 17°58'N. 102°36'E; reported before 1964 by J. Deuve and M. Deuve (1963, p. 59). B:L-7. [VictorialJ Peak; Xianggang (= Hong Kong), CHINA; 22°16'N, 114°08'E; reported before 1952 by G. A. C. Herklots (1951, p. 83). C:C- 210. Vientiane. See Viangchan. VIETNAM; 15°-23°N, 102°-I09°E; collected 28 Feb. 1965 by unknown collector; zmvnu, 2 (skulls only). Collected in 1992 and 71994 by unknown collectors; iebr, 2(1 skin only, 1 skull only). Date and collector unknown; fcxm, 1 (skull only; identification tentative); iebr, 14 (including 5 skins only. 8 skulls only); zmvnu, 14 (3 skins only, 1 1 skulls only [one identifi- cation tentative]). Not mapped. [VIETNAM]; 15°-23°N. 102°-109°E; collected before 1963 by J. Delacour and W. P. Lowe; MNHN. 2 (skulls only). Not mapped. View. Simla vicinity; Himachal Pradesh, INDIA; ca. 31°06'N, 77°11'E; observed Aug. 1972- Feb. 1973 by K. Wada (1984. p. 477). A:I-18. Vijayawada. Krishna District, 80 m; Andhra Pra- desh, INDIA; 16°32'N, 80°38'E; observed 21 Nov. 1972 by N. Koyama and P. B. Shekar (1981. p. 248). A:I-144. Vinh Linh region. 50 m; Quang Tri, VIETNAM; ca. 17°04'N. 107°02'E; collected 28 Aug. 1956 by unknown collector (Dao, 1960, p. 228; 1962, p. 724); misidentified as M. assamensis; muse- um unknown. C:V-32. Vinukonda. Guntur District, 150 m; Andhra Pra- desh, INDIA; 16°03'N. 79°45'E; observed 30 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981. p. 468). A:I-147. Visakhapatnam. See Malkangiri. Vizag. See Malkangiri. Vrindavan (= Vrindaban); Uttar Pradesh, INDIA; 27°35'N. 77°42'E; observed 1964-1965 by C. H. Southwick and M. R. Siddiqi (1966, p. 309). Reported in 1996 by T Patel (1996, p. 10). A: 1-71. Waddepalle, Banswada Taluk, Nizamabad Dis- trict; Andhra Pradesh, INDIA; 18°I4'N, FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 173 77°54'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:l-117. Walter Road. See Dhaka. Wama. north of ; Komirha, AFGHANISTAN; ca. 35°20'N. 70°45'E; reported before 1972 by A. Puget (1971, p. 200). A:A-2. Wangjuanshan, Huaiji Xian; Guangdong. CHINA ca. 24°15'N. 112°20'E; reported July 1982 by local residents (Liu Zhenhe, sciha, pers. comm., 25 Nov. 1985). C:C-200. Wanglang Natural Reserve; Sichuan, CHINA; ca. 30°40'N, I03°20'E; reported 1968-1969 by Gi- ant Panda Expedition of the Wanglang Natural Reserve (1974, p. 163). C:C-28. Wangmo; Guizhou, CHINA; 25°14'N, 105°59'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-158 Wanhsien. See Wa Shan. Wanxian; Sichuan, CHINA; 30°49'N, 108°21'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-38. Wanyuan; Sichuan, CHINA; 32°04'N, 108°02'E; reported before 1998 (Zhang et al.. 1997, p. 58). C:C-37. Warangal, 350 m; Andhra Pradesh, INDIA; 18°00'N, 79°35'E; observed 18-19 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A:I-121. Wargel, Gajwel Taluk, Medak District; Andhra Pradesh, INDIA; 17°46'N, 78°38'E; reported Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, p. 18). A:I-123. Wa Shan (- Waschan), near Dong He (= Tun- gho), 900-2000 m; Sichuan, CHINA; ca. 29°15'N, 103°03'E; collected 29 Mar. 1915 by H. Weigold (1916, p. 74; 1922, p. iv; 1924, p. 71; 1935, p. 212; Israel, 1919. pi. 7; Jacobi, 1923, p. 1); RMNH, 2. C:C-140 Wassuland; Sichuan, CHINA; ca. 3r05'N, 103°10'E; tentatively reported 1914-1916 by H. Weigold (1924, p. 71). C:C-27. Wazipur (= Uzipur); Barisal, BANGLADESH; 22°50'N, 90°15'E; reported before 1986 by M. A. R. Khan (1981, p. 13; 1985, p. 31). Ob- served Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba- 24. Weixi; Yunnan, CHINA; 27°13'N, 99°16'E; re- ported before 1998 (Zhang et al., 1997, p. 58). B:C-41. Wenchang Xian; Hainan Dao; Hainan, CHINA; ca. 19°37'N, 110°43'E; reported before 1986 by Xu Longhui and Liu Zhenhe (1985, p. 148). C: C-23 1 . Wenchuan; Sichuan, CHINA; 31°28'N, 103°35'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-26. Weng'an; Guizhou, CHINA; 27°00'N, 107°32'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-126. West Bengal, INDIA; ca. 22°-28°N, 86°-90°E; ob- tained before 1857 by Theobald Collection; bm(nh), 1 (skull only). Collected date unknown by [C. A. Crump]; bnhs, 1 (skull only). Not mapped. West Bhanugach; Moulvi Bazar, BANGLA- DESH; ca. 24°21'N, 91°48'E; reported early in 1980 by S. R Gittins and A. W. Akonda (1982, p. 278). Observed Feb. 1990-June 1993 by M. M. Feeroz, M. A. Islam, and M. Kabir (1995, p. 76). B:Ba-13. West District, east-central; Tripura, INDIA; ca. 23°55'N, 91°45'E; observed in 1976 and 1978 by R. R Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). B:I-40. West District, south-central; Tripura, INDIA; ca. 23°37'N, 91°23'E; observed in 1976 and 1978 by R. P Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). B:I-40. West District, southern; Tripura, INDIA ca. 23°05'N, 91°23'E; observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). B:I-40. West District, southwestern; Tripura, INDIA; ca. 23°17'N, 91°22'E; observed in 1976 and 1978 by R. R Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). B:I-40. West District, western; Tripura, INDIA; ca. 23°43'N, 91°irE; observed in 1976 and 1978 by R. P Mukherjee (1982, p. 71). Observed May-Aug. 1989 by A. K. Gupta (1994, p. 102). B:I-40. Wast Garo Hills District; Meghalaya. INDIA; ca. 25°30'N, 90°00'E; reported July 1985-Mar. 1987 by J. R. B. Alfred and J. P Sati (1990, p. 300). B:I-11. West Sichuan. See Sichuan, western. West Timli; Uttar Pradesh, INDIA; ca. 30°23'N, 77°43'E; reported 1964-1966 by D. G. Lind- burg (1977a, p. 268). A:I-26. Wira, 1 km south of. Khammam District, 100 m; Andhra Pradesh, INDIA; 17°irN. 80°22'E; observed 21 Apr. 1980 by J. Fooden, A. Ma- habal, and S. S. Saha (1981, p. 467). A:I-132. Wolongshi. See Olongche. Wonglung kun. See Luofu Shan. 174 FIELDIANA: ZOOLOGY Wuchi. See Wuzhi Shan. Wudu; Gansii, CHINA; 33°24'N, 104°50'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-21. Wuliang Shan Reserve; Yunnan, CHINA; ca. 24°00'N, lOrOO'E; reported in 1990 by L. K. Sheeran and F. E. Poirier (1994, p. 21). B:C-73. Wushan; Sichuan, CHINA; 31°02'N, 109°56'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-40. Wu-tsao. See Xi Jiang. Wuyi Shan; Fujian, CHINA; ca. 27°45'N, 117°39'E; reported 1963-1990 by Wu Haohan (1995, p. 277). Reported Nov. 1983 by Zheng Xueqing (1984. p. 145). C:C-76. Wuzhi Shan, Hainan Dao; Hainan, CHINA; ca. 18°54'N, 109°40'E; collected 1-30 Oct. 1905 by A. Owston;AMNH, 10 (including holotype of Pithecus brachyurus and Pithecus brevicau- dus); BM(NH), 1. C:C-232. Wuzhou; Guangxi, CHINA; 23°29'N, lin9'E; tissue sample obtained ca. 1991 by Zhang Yap- ing and Shi Liming (1993b, p. 589). C:C-197. Xaignabouri, LAOS; ca. 19°15'N, 101°45'E; re- ported before 1964 by J. Deuve and M. Deuve (1963, p. 59). B:L-4. Xhin Xhan. See Huangliangping. Xialei Water Regulation Forest Reserve; Guangxi, CHINA; ca. 22°26'N, 106°26'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter, Aug. 1996). C:C-224. Xianan. Huanjiang Xian; Guangxi, CHINA; ca. 24°59'N, 107°59'E; collected Oct. 1992 by local farmer (Wei Yuhei, manager of medical prod- ucts station, pers. comm., 3 Nov. 1992); skel- eton examined 3 Nov. 1992 at Huanjiang. C:C- 177. Xianan-Mulun, Huanjiang Xian; Guangxi, CHI- NA; ca. 25°03'N, 107°57'E; collected ca. 1991 by local resident (Tan Yulung, pers. comm., 5 Nov. 1992); skull examined 5 Nov. 1992 at Mu- lun. C:C-177. Xiangcheng; Sichuan; CHINA; 29°00'N, 99°46'E, reported before 1998 (Zhang et al., 1997, p. 58). B:C-37. Xianggang (= Hong Kong), CHINA; 22° 10'- 22°35'N, 113°55'-114°25'E; reported 1990- 1992 by N. J. Goodyer (1992, p. 72); popula- tion apparently artificially introduced. Not mapped (see C:C-210). Xiangkhoang, LAOS; ca. 19°20'N, I03°22'E; re- ported before 1964 by J. Deuve and M. Deuve (1963, p. 59). C:L-1. Xidaming Shan Water Regulation Forest Reserve; Guangxi, CHINA ca. 2°49'N. 107°32'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 127; letter, Aug. 1996). C:C-225. Xieng-Khouang. See Xiangkhoang. Xi Golog (= Singolo); Sichuan, CHINA; 30°00'N, 100°42'E; collected 31 Oct. 1931 by Dolan West China Expedition (Schafer, 1933, p. 191; 1942, p. 257; Stone, 1933, p. 170; Do- lan, 1939, p. 178); ansp (skin) / mcz (skull), 1. B:C-30. Xi Jiang (= Hsi-kiang; river), near Wuzhou ( = Wu-tsao); Guangxi, CHINA; ca. 23°29'N, 111°19'E; collected 27 Apr. 1912 by R. Mell (Matschie, 1912, p. 305); zmb, 1. C:C-197. Xilin Shan Water Regulation Forest Reserve; Guangxi, CHINA; ca. 24°50'N. 110°10'E; ob- served 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 129; letter, Aug. 1996). C:C-194. Xindeng, Fuyang Xian, <500 m; Zhejiang, CHI- NA; 29°58'N, 119°44'E; collected 2 Feb. 1963 by museum collector; zmnh, 1 (skin with skull inside). C:C-58. Xinglong Xian, southern (= Eastern Tombs); He- bei, CHINA; ca. 40°24'N, 117°30'E; collected in 1867 by M. Fontanier (Milne-Edwards, [1870], pi. 32; [1872], p. 227; Zhang et al., 1989, p. 376); mnhn, 1 (holotype of Macacus Tcheliensis). Reported May-Oct. 1872 by A. David (1875, vol. 1 p. 42). Captive obtained 1879-1880 by S. W. Bushell (in Sclater, 1881, p. 537; Hill, 1974, pp. 581, 582); died in zoo 6 Mar. 1881; bm(nh), 1 (skin only). Reported in 1914 by local residents (Sowerby, 1925, p. 12). Collected and/or purchased 1920-1922 by R. C. Andrews, Third Asiatic Expedition (An- drews, 1932, p. 20; Pope, 1932a, p. 470); amnh, 7; FMNH, 3. Collected in [1923] by F R. Wulsin (letter, 9 Jan. 1925, usnm archives); u.snm, 3 (in- cluding 1 .skull only). Reported before 1940 by A. de C. Sowerby (1939, p. 228; I94I, p. 261). Observed in 1963 by Quan Guoqiang (Zhang et al., 1989, p. 379). Reported ca. 1964 by Shui Chingwong, Lee Yangwan, and Kan Singwun (Shou, 1964, p. 61). Reported in 1976 by local farmers to Quan Guoqiang (Zhang et al., 1989, p. 379). C:C-1. Xinglung. See Xinglong Xian, southern. Xingning; Guangdong, CHINA; 24°08'N, 115°43'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-95. Xingyi; Guizhou, CHINA; 25°03'N, 104°59'E; re- FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 175 ported before 1998 (Zhang et al., 1997, p. 58). C:C-156. Xinlong. Wanning Xian, 500-600 m; Hainan Dao; Hainan. CHINA; 18°45'N, 110°12'E; col- lected 3 July 1960 by Wu Luping (Liu Zhenhe, sciKA. pers. comm.. 26 Nov. 1985); sciea, 1. C: C-240. Xinluwan, Suichang Xian; Zhejiang. CHINA; 28°42'N, 119°15'E; captives acquired 1970- 1980 by Fu Yiyuan and Wu Fuhai, Hangzhou Zoo (pers. comm., 25 Oct. 1985). C:C-65. Xinning Xian, southern; Hunan CHINA; ca. 26°20'N. 1 10°50'E; reported Nov. 1980 by local residents (Liu Zhenhe. sciea, pers. comm.. 25 Nov. 1985). C:C-187. Xinxiang (= Xinyang) Xian; Henan, CHINA; 35°19'N, 113°52'E; reported before 1989 by Tan Bangjie and E E. Poirier ([1991], p. 131). C:C-9. Xi Shia, Dongfang Xian. Hainan Dao; Hainan. CHINA; ca. 19°00'N, 108°55'E; collected 13 and 14 Apr. 1960 by Quan Guoqiang (pers. comm.. 25 Aug. 1983); izcas, 3 (including 2 skulls only). C:C-235. Xishuangbanna (perfecture); Yunnan, CHINA; 21°-23°'N, 99°-102°'E; collected in 1960 and 1964 by unknown collectors; kiz, 2 (skins only). Not mapped (see B:C-80 through B:C- 86). Xishui; Guizhou, CHINA; 28°24'N, 106°15'E; re- ported before 1998 (Zhang et al.. 1997, p. 58). C:C-134. Xiuwu Xian; Henan. CHINA; 35°14'N, 113°26'E; reported before 1989 by Tan Bangjie and E E. Poirier ([1991], p. 131). C:C-10. Xuan Ninh; Quang Binh. VIETNAM; 17°20'N, 106°39'E; collected 17 Oct. 1964 by unknown collector (Dao, 1985, pp. 248, 225; external measurements published); iebr, 1 (skin only). C:V-31. Xunle vicinity, Huanjiang Xian; Guangxi. CHI- NA; ca. 25°22'N, 108°15'E; purchased Aug.- Oct. 1992 from local residents by Wu Xiaowu, Xunle (pers. comm., 4 Nov. 1992); three skins examined 4 Nov. 1992 at Xunle. C:C-176. Xunle Water Regulation Forest Reserve; Guangxi. CHINA; ca. 25°22'N, 108°12'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C-176. Yadagiri Gutta, Nalgonda District, 530 m; Andhra Pradesh. INDIA; 17°32'N, 78°55'E; observed 18 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A:I-125. Yadong Xian; Xizang (= Tibet). CHINA; ca. 27°30'N, 89°00'E; reported before 1964 by Shen Xiaozhou (1963, p. 140; Zhang et al., 1989, p. 379; 1991, p. 177; Zhang Yongzu, let- ter, 3 July 1996). B;C-1. Yai-cheng. See Nychow. Yajiang, 10000 ft (= 3050 m); Sichuan, CHINA; 30°02'N, 101°02'E; collected 26 Aug. 1908 by W. R. Zappey (Henshaw, 1912, p. 109; Dolan, 1939, p. 177); mcz, 1. B; C-29. Yangcheng; Shanxi. CHINA; 35°32'N, 112°36'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-5. Yangliupu (= Yangliu), Xuancheng Xian, 200- 400 m; Anhui. CHINA; ca. 30°50'N, 118°36'E; observed 1973-1986 by Xiong Chenpei (Wada et al., 1986, p. 83). C:C-54. Yangshan; Guangdong, CHINA; 24°29'N, 112°38'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-202. Yangtze Gorges. See Yichang (= Ichang). Yanjing vicinity; Xizang (= Tibet), CHINA; ca. 29°07'N, 98°33'E; reported 1914-1916 by H. Weigold (1924, p. 71). B:C-35. Yanyuan; Sichuan, CHINA; 27°25'N, 101°33'E; reported before 1998 (Zhang et al., 1997, p. 58). B:C-40. Yao, Nam. See Mansam Falls. Yellandu, Khammam District, 200 m; Andhra Pradesh, INDIA; 17°35'N, 80°20'E; observed 20 Apr. 1980 by J. Fooden, A. Mahabal, and S. S. Saha (1981, p. 467). A:I-I34. Yenangyaung. See Irrawaddy River. Yen Bai. VIETNAM; 21°15'-22°20'N, 103°55'- 105°10'E; collected in 1963 by unknown col- lector; lEBR, 1 (skin only). Not mapped. Yeppuru, Nazvid Taluk, Krishna District; Andhra Pradesh. INDIA; ca. 16°45'N, 80°50'E; report- ed Feb. 1977-July 1980 by G. U. Kurup (1984, p. 58; 1992, pp. 17, 19). A:H40. Yiajia, Bawangling District, Changjiang Xian, 1000 m; Hainan Dao; Hainan. CHINA; 19°05'N, 109°08'E; collected 24 Oct. 1964 by Liu Zhenhe, sciea (pers. comm., 26 Nov. 1985); SCIEA, 1. C:C-234. Yibin (= Suifu); Sichuan. CHINA; 28°46'N, 104°34'E; collected 10 Oct. 1922 by D. G. Gra- ham; usNM, 1. C:C-137. Yichang; Hubei, CHINA ca. 30°42'N, lin8'E; reported before 1989 by Hu Hongxing (Zhang et al., 1989, p. 379; 1991, p. 177; Zhang Yong- zu, letter, 3 July 1996). C:C-46. Yichang (= Ichang), Chang Jiang (= Yangtze) gorges above; Hubei, CHINA; ca. 30°45'N, 176 FIELDIANA: ZOOLOGY 111°15'E; reported before 1942 by A. de C. Sowerby (1941, p. 262). C:C-46. Yicheng; Shanxi, CHINA; 35°42'N. 1 1 1°40'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-2. Yigong, Bomi Xian 2250 m; Xizang (= Tibet), CHINA; 30°08'N, 95°02'E; collected 24 June 1973 by Zheng Changlin and Cai Guiquan (pers. comm., 7 Oct. 1985; Feng et al., 1984, p. 344); NwpiB, 1. B:C-11. Yigong Forest Reserve, Bomi Xian; Xizang (= Tibet), CHINA; 30°08'N. 95°02'E; captives ob- tained 1979-1982 by Zhang Cizu, Director, Shanghai Zoo (pers. comm., 18 Oct. 1985); captives observed 18 Oct. 1985. B:C-11. Yiliang; Yunnan, CHINA; 27°35'N, 104°0rE; tis- sue sample obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). C:C-143. Yin, east bank of lower Chindwin River; Sagaing, MYANMAR (= BURMA); 22°47'N, 94°42'E; collected 9-18 June 1914 by G. C. Shortridge (in Wroughton, 1916a, p. 294); bm(nh), 1; BNHS, 1; FMNH, 2; ZSI, 1 (skin only). B:M- 24. Yin, lower Chindwin River; Sagaing, MYAN- MAR (= BURMA); 22°47'N, 94°42'E; collect- ed 15 and 18 June 1914 by G. C. Shortridge and S. A. Macmillan (Shortridge in Wroughton, 1916a, p. 294); bnhs, 3. B:M-24. Yindian Shan Water Regulation Forest Reserve; Guangxi, CHINA; ca. 25°26'N, 110°30'E; ob- served 1976. 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 128; letter, Aug. 1996). C:C-190. Yingde; Guangdong, CHINA; 24°10'N, 1 13°24'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-204. Yingjiang; Yunnan, CHINA; 24°48'N, 98°05'E; reported before 1998 (Zhang et al., 1997, p. 58). B:C-60. Yinjiang; Guizhou, CHINA; 28°01'N, 108°24'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-118. Yixian (= Qianxian); Anhui, CHINA; 29°53'N, 117°57'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-62. Yong'an Xian; Fujian, CHINA; ca. 25°58'N 117°22'E; reported Apr. 1981 by Zheng Xue- qing (1984, p. 145). C:C-89. Yongchun; Fujian, CHINA; 25°19'N, 118°17'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-87. Yongde vicinity; Yunnan, CHINA; ca. 24°(X)'N. 99°15'E; purchased in market Aug. 1964 by Quan Guoqiang (pers. comm., 25 Aug. 1983); IZCAS, 1 (skin only). B:C-66. Yongshan; Yunnan, CHINA; 28°irN, 103°35'E; tissue sample obtained ca. 1991 by Zhang Yap- ing and Shi Liming (1993b. p. 589). C:C-142. Yongsheng [Xian]; Yunnan, CHINA; ca. 26°42'N, 100°45'E; blood sample obtained before 1999 by Ding Bo, Zhang Yaping, and Hou Yidi (1998. p. 172). B:C-46. Yongshun; Hunan, CHINA; 29°(X)'N. 1()9°54'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-112. Yongtai Xian; Fujian, CHINA; ca. 25°52'N, 118°55'E; reported Nov. 1983 by Zheng Xue- qing (1984, p. 146). C:C-84. Yongyap Chu (= Yongyap Valley). 9500 ft ( = 2900 m); Arunachal Pradesh, INDIA; ca. 29°10'N, 95°37'E; observed May-June 1913 by E M. Bailey (1914, map; 1915. p. 74). B:I-27. Youxi Xian; Fujian, CHINA; ca. 26°10'N. I18°II'E; reported Mar. 1981 by Zheng Xue- qing (1984. p. 145). C:C-83. Youyang; Sichuan. CHINA; 28°52'N. 108°45'E; reported before 1992 by Jiang Xuelong, Wang Yingxiang. and Ma Shilai (1991, p. 244). C:C- 117. Yu, Nam. See Ou. Nam. Yuanbao Shan Nature Reserve; Guangxi, CHINA; ca. 25°27'N, 109°10'E; observed 1976, 1986, and 1993 by Liu Wanfu and Wei Zhenyi (1995, p. 126; letter, Aug. 1996). C:C-182. Yuanmou; Yunnan, CHINA; 25°42'N, 101°52'E; reported before 1998 (Zhang et al., 1997, p. 58). B:C-49. Yuanqu; Shanxi, CHINA; 35°18'N, lll°4rE; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-14. Yuhun, Lingyun Xian; Guangxi, CHINA; ca. 24°24'N, 106°31'E; collected 10 Aug. 1978 by Neuong Shihua; fdcg, 1 (skull only). Collected Nov.-Dec. 1978 by Ling Chen; fdcg, 1 (skull only). C:C-171. Yung-ling. See Xinglong Xian, southern. Yunlong [Xian]; Yunnan, CHINA; ca. 25°50'N, 99°28'E; blood sample obtained before 1999 by Ding Bo, Zhang Yaping, and Hou Yidi (1998, p. 172). B:C-55. Yunnan, CHINA; 21°-29°N, 98°-106°E; collected in 1957 by Quan Guoqiang; izcas, 1 (skull only). Date and collector unknown; izcas, 1 (skull only). Not mapped. Yunnan, [northwestern], CHINA; 25°-29°N, 97°- 102°E; tissue samples obtained ca. 1991 by FOODEN: SYSTEMATIC REVIEW OF THE RHESUS MACAQUE, MACACA MULATTA 111 Zhang Yaping and Shi Liming (1993b, p. 589, fig. 1). Not mapped (see B:C-36). Yunnan border; Sichuan. CHINA; ca. 28°20'N. 104°20'E; collected 17 Sept. 1928 by D. G. Graham; usnm, (skull only). C:C-I38. Yunnan border, south of Yibin (= Suifu) 3000 ft (= 910 m); Sichuan. CHINA; ca. 28°20'N. I04°20'E; collected 19 Feb. and 25 Mar. 1932 by D. G. Graham; usnm, 2. C:C-138. Yuqing; Guizhou, CHINA; 27°12'N, 107°56'E; re- ported before 1998 (Zhang et al., 1997, p. 58). C:C-120. Yushu Xian, 3600-4300 m; Qinghai, CHINA; ca. 33°00'N, 96°45'E; reported 1959-1961 by Chang Chieh and Wang Tsung-yi (1963, p. 126). Captive obtained in 1981 by Liao Yianfa, Director. Xining Zoo (pers. comm., 6 Oct. 1985); captive observed 6 Oct. 1985. B:C-17. Zackala. See Yanjing vicinity. Zayu Xian; Xizang (= Tibet). CHINA; ca. 28°28'N, 97°04'E; collected in 1973 by Feng Zuojian (Feng et al., 1984, p. 344; Quan Guo- qiang, letter, 30 Oct. 1995); izcas, 3 (including I skin only). Purchased Aug. 1973 at traditional medicine shop, Qamdo, by Zheng Changlin (pers. coimn., 7 Oct. 1985); nwpib, 3 (skulls only). B:C-34. Zhangjiajie Nature Preserve. See Suoxi Valley. Zhaotan (= Zhaotang), Dongzhi Xian, 200-500 m; Anhui. CHINA; ca. 29°39'N, 116°49'E; ob- served 1973-1986 by Xiong Chenpei (Wada et al., 1986, p. 83). C:C-47. Zhayun, Qiongzhong Xian, Hainan Dao, 200 m; Hainan. CHINA; 19°00'N, 109°36'E; collected 15 Oct. 1963 by Liu Zhenhe, sciea (pers. comm., 26 Nov. 1985); sciea, I. C:C-232. Zheng'an; Guizhou. CHINA; 28°30'N, 107°30'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-132. Zhengba Xian; Shaanxi, CHINA; ca. 32°30'N, 107°50'E; captives obtained Aug. 1985 by Tian Xiaoming, Director, Xi'an Zoo (pers. comm., II Oct. 1985); captives observed 11 Oct. 1985. C:C-36. Zhenping; Shaanxi. CHINA; 31°56'N, 109°3rE; reported 1963-1966 by Wu Jiayan and Li Gui- hui (1982, p. 63). C:C-41. Zhenyuan Xian; Yunnan, CHINA; ca. 23°5rN, 100°59'E; immunological survey conducted be- fore 1996 by Duan Xingsheng, Liu Yuanwei, Wu Jing, Dao Weiying, and Liu Jianghai (1995, p. 411). B:C-73. Zhidaikou, Suichang Xian; Zhejiang. CHINA; 28°16'N, 118°46'E; observed Aug. 1985 by Kang Ximin, zmnh (pers. comm., 24 Oct. 1985). C:C-66. Zhongdian; Yunnan. CHINA; 27°50'N, 99°36'E; re- ported before 1992 by Jiang Xuelong, Wang Yingxiang, and Ma Shilai (1991, p. 243). B:C-38. Zhongtiao Shan, 1050-1500 m; Shanxi CHINA; ca. 35°15'N, 1 1 r30'E; reported before 1966 by Tang Changzhu, Ma Yong, Wang Jianjun. Wang Ziyu, and Zhou Naiwu (1965, p. 88). Reported before 1987 by Wang Sung and Quan Guoqiang (1986. p. 215). C:C-14. Zhongzhou; Guangxi. CHINA; ca. 22°40'N, 107°05'E; collected 24 Mar. 1982 by Wu Mingchuan (pers. comm., 27 Nov. 1992); fdcg, 1 (skin with skull inside). C:C-223. Zhoucun. Jiangshan Xian, 1000 m; Zhejiang, CHINA; 28°22'N, 118°37'E; collected Jan. 1985 by Kang Ximin (pers. comm., 24 Oct. 1985); ZMNH, 1 (skull examined, skin unavail- able Oct. 1985). Collected 13 May 1985 by Kang Ximin (pers. comm.. 24 Oct. 1985); mu- seum unknown. C:C-66. Zhouning Xian; Fujian. CHINA; ca. 27°15'N, 119°13'E; reported Oct. 1980 by Zheng Xue- qing (1984, p. 146). C:C-69. Zhuhai Xian. See Dangan Dao. Zhushan; Hubei. CHINA; 32°13'N, 110°24'E; tis- sue sample obtained ca. 1991 by Zhang Yaping and Shi Liming (1993b, p. 589). C:C-42. Zhuxi, Longquan Xian; Zhejiang, CHINA; 28°11'N, 118°54'E; purchased Sept. 1972 from local residents by Cai Chunmo, zmnh (pers. comm.. 24 Oct. 1985); not retained. C:C-66. Zigui; Hubei. CHINA; 31°01'N. 110°35'E; reported before 1998 (Zhang et al., 1997, p. 58). C:C-45. Zixi Xian; Jiangxi, CHINA; ca. 27°45'N, 117°00'E; trapped in 1982 for Nanchang Zoo (Huang Zhangsen, pers. comm.. 28 Oct. 1985); captives observed 28 Oct. 1985. C:C-78. Zoige Xian; Sichuan. CHINA; ca. 33°30'N, 102°54'E; reported before 1983 by Hu Jinchu and Wang Youzhi (Zhang et al., 1989, p. 379; 1991, p. 177; 1997, p. 58; Zhang Yongzu, letter, 3 July 1996). B:C-25. Zunyi vicinity; Guizhou, CHINA; ca. 27°42'N. 106°55'E; purchased in market Nov. 1960 by Quan Guoqiang (pers. comm., 25 Aug. 1983); iz- cas, 1 (skin only). Collected 30 and 31 May 1964 by Fang Lixiang; bmnh, 4 (skins only). C:C-129. 178 FIELDIANA: ZOOLOGY Index Specific and subspecific names referable to Macaco miilatta are spelled here as they were originally proposed. abbreviations, institutional names 1 Allen's rule 87 annual mortality rate 78 Anopheles 53 arboreal ity 54 archaeological evidence 3, 89, 154 Axis axis 68 Bergmann's rule 28 beta-globin gene 52 birth peaks 68 birth rate, annual 75 birth weight 75 blood proteins 52, 83 body weight adult 30 neonatal 75 brachyurus 86 brevicaudus 24. 86 Canis aureus 68 cerebrospinal fluid 53 chemokine receptor 52 chromosomes 52 climatic deterioration 87 commensalism with humans 54 competition, interspecific 3 consortship 72 copulatory behavior 72 Connts sp. 68 cranial measurements 38 geographic variation 39. 79 insular variation 39 ontogeny 38 sexual dimorphism 38 crop raiding 54, 57 day range 57 death, causes of 78 dental emergence 38 diet 57 geographic variation 57 ingestion of soil 59 nutritional requirements 59 seasonal variation 57 diploid chromosome number 52 dispersal scenario 87 distribution factors limiting 3 geographic 2 dominance rank and reproductive success 73 drinking 60 elevational range 54 eiytluaea 84 estrous cycle 71 evolutionary hypothesis 87 external measurements 26 geographic variation body weight 30 head and body length 26, 79 relative tail length 29, 81. 87 tail length 29. 81 hybrids 26 ontogeny 26 sexual dimorphism 26 extirpated population 3 fascicularis group 87 fat deposits, seasonal variation 71 feeding schedule 61 fossils 79 founder effect 89 fulvus 84 geophagy 59 geographic reference works 124 gestation length 74 glacial maximum, last 87 glans, color of 7 golden pelage phenotype 7 group fission 57 group size and composition 56 habitats 54 harassment of copulaUons 73 hematology 53 hoine range area 57 seasonal variation 57 homosexual mounting 74 HPRT locus 52 hybridization 29. 30. 89 hybrids 26, 53 inbreeding, incidence of 73 inerspecific behavior 66 intergeneric 68 intrageneric 66 infant corpse carried by mother 76 mortality rate 75 sex ratio 75 infanticide 78 infrazygomatic crest 7 insular populations 3, 79, 81, 89 intergroup behavior 62 intergroup transfer 63, 70 introduced populations 3. 67 invertebrate prey 59 karyology 52 lasiotis 22, 26, 85 last glacial maximum 87 lateral facial crest 7 lead content of molars 53 Leucosphyrus group 53 littoral is 23. 85 longevity 78 Macaca arctoides 66 assamensis 16, 66 cwiopis 87 fascicularis 3, 44. 67, 87, 89 fuscata 16, 87, 89 ueinestrina 66 radiata 3, 16, 67 thibetana 16, 66 malaria 53 Mannota hiinalayami 3 masturbation 74 mcmahoni 18, 86 menarche 70 menopause 77 menstrual cycle 71 mitochondrial DNA 44, 82. 89 molecular biology 44 molting 1 1 mortality rate, annual 78 multimount ejaculation 72 Muntiacus muntjak 68 museums 1 natural history 54 neutral mutation hypothesis 52 nipaleihsis 84 nonreproductive sexual behavior 74 nuclear DNA 52. 83 nursing 76 nutritional requirements 59 oinops 84 parturition 56. 74 pelage 7 elevational variation 7 erythrism 7 geographic variation 15. 16, 82 hair banding 7 hair length 26. 82 hairiness of ears 26 individual variation 21, 25, 82 neonatal 7 Nepalese standards 16 ontogenetic changes 7 seasonal variation 1 1 survey of sainple areas 16 Plasmodium 54 population density 57 estimates 7 growth rate 78 predators 61, 78 pregnancy 74 INDEX 179 prey 59 provisioned groups 57, 65. 74 rejected locality records 3, 129, 155 relative tail length 29. 81, 87 reproduction 68 reproductive success 73 rhesus 84 sagittal crest 44 sancti-johannis 24, 84 scream call 54 seasonal breeding 68 Semnopithecits entellits 68 serotonin activity 66 sex ratio adult 57 neonatal 75 sexual maturation 70 sexual skin 7, 71 females 71 males 71 shoulder girdle and humerus 56 siamica 25, 86 simian immunodeficiency virus 54 simian T-lymphotropic retrovirus 54 simple sequence repeat loci 52 sinica group 89 sleeping sites 56 snakes, fear of 62 solitary males 57 southern hemisphere, reversed cy- cles 69 specimens examined 121 stillbirths, frequency of 74 subfossils 79 subspecific recognition 79 swimming 56 synonymy 83 systematics 79 tail carriage 29 tcheliensis 23, 85 temperament 53 terrestriality 54 testicular descent 70 time budget 66 Trachypithecus geei 68 pileatus 68 transzygomatic crest 7 twinning, frequency of 74 type 86 type locality 86 vaginal plug 73 vectors, malaria infection 53 vegetational range 54 vertebrate prey 59 vestitus 22, 85 villosus 17, 85 viral infections 54 water requirements 59 weaning 76 weight. See body weight 180 FIELDIANA: ZOOLOGY A ^ielccled Listing of Othier tielUiaitu a liable Studies in Neotrop and Robert ^ i By Jack Fooden. Fieldiana: Zoology, r le Bats of the Philippine Islands. By Nina K. Ingl ubiuafion 1382, S45.00 \\ib\^ Publication 1440, SI 4.00 S}^^J:uauL Kc - icw of Southeasi .v Jack Fooden. Fieldiana: Zoology, ') tables. Publication i4/ii. >()>.mi» The Birds of Sibuyan Island, Romblon Province, Philippines, with Particular Reference to Elevationa! Distribution and Biogeographir AlTinifies Rv *^fonb'"p VT rjo'^Hinrin OnviM F wnirirfl -in/) p.> luc jiiiyaoit.- m !.:->. ».ii>iKu-ciit;i.K.> on any U.S. bank or the U.S. subsidiary of any foreign bank. Prices and terms subject to change w ^ ' ' ill requests to; 1400 Sontii I