mmmMmMmm'ifmp \i FIELD ± K Zoology II NO. 96 r-t-i Systematic Review of the Rhesus Macaque, Macaca mulatta (Zimmermann, 1780) Jack Fooden OC3 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 j<c4|>vi vYiiij wuie If typed on an IBM-compatible computer using MS-DOS, also submit text on SVi-inch "> i 1,4.2, or 5.0, MiiltiMp!.- n.^r,iov.,..rit.» 1 '>f '• x- w-.^^.- ', i ". "^"--^im M^ , |"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 ofrecent 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, 63-80. pp. 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 Sexual Maturation 70 Sexual Skin 71 Menstrual Cycle 71 Abstract 1 Estrus 71 Introduction 1 Consortship 72 Geographic Distribution and Current Copulatory Behavior 72 Population Estimates 2 Dominance Rank and Reproductive Suc- Pelage 7 cess 73 General Characterization 7 Inbreeding 73 Early Development 7 Nonreproductive Sexual Behavior 74 Seasonal Variation 11 Gestation Length 74 Geographic Variation 16 Parturition 74 Nepalese Standards 16 Birth Weight, Infant Sex Ratio 75 Survey of Sample Areas 16 Birth Rate, Infant Mortality Rate 75 Summary 25 Nursing, Weaning 76 External Measurements and Proportions 26 Menopause 77 Sex and Age Variation 26 Annual Mortality Rate 78 Geographic Variation 26 Population Growth Rate 78 Head and Body Length 26 Fossils and Subfossils 79 Tail Length 29 System ATics 79 Relative Tail Length 29 Geographic Variation and Subspecific Body Weight 30 Recognition 79 Cranial Characters 38 Size 79 Sex and Age Variation 38 Tail Length 81 Geographic Variation 39 Pelage 81 Skull Length 39 Molecular Diversity 82 Cranial and Dental Morphology 42 Synonymy 83 Comparison with Macaca fascicularis 44 Type 86 Molecular Biology and Genetics 44 Type Locality 86 Mitochondrial DNA 44 Evolution and Dispersal 87 Nuclear DNA 52 Acknowledgments 89 Blood Proteins 52 Literature Cited 90 Karyology 52 Note Added in Proof 120 Physiology and Disease 53 Appendix 1: Specimens Examined 121 Blood, Cerebrospinal Fluid, and Temper- Appendix 2: Gazetteer of Macaca ament 53 MULATTA Localities 124 Lead Content of Molars 53 Index 179 Malaria 53 Viral Infections 54 Natural History 54 List of Illustrations Habitats 54 Arboreality/Terrestriality 54 Swimming 56 1 Known locality records of Macaca mii- Group Size and Composition 56 latta 2 Home Range, Day Range 57 2. Detail maps of Macaca mulatta locali- Population Density 57 ties Diet 57 A. Western section 4 Predators 61 B. Central section 8 Intergroup Behavior 62 C. Eastern section 12 Interspecific Behavior 66 3. Dorsal pelage color in Macaca mulatta Intrageneric 66 A. Topotype 17 Intergeneric 68 B. Contrasting pelage color in two Reproduction , 68 adult males collected 4 days apart Seasonality 68 at Rajapara, India 17 m . 4. Monthly incidence of prime, faded, List of Tables and molting pelage stages in wild-col- lected specimens of Macaca mulatta 18 1 Estimated population of Macaca mulat- 5. areas cited in Macaca mulatta Sample ta in five countries for which data are 20 pelage comparisons available 3 6. Latitudinal variation in interscapular 2. External measurements and proportions length in Macaca mulatta adults .... 27 hair in age/sex classes of wild-collected 7. Latitudinal variation in midtail hair Macaca mulatta 30 length in Macaca mulatta adults 28 3. Geographic variation in head and body 8. Latitudinal variation in head and body length in adult Macaca mulatta 31 length in Macaca mulatta adult non- 4. Geographic variation in tail length in captives 32 adult Macaca mulatta 33 9. Latitudinal variation in tail length in 5. Geographic variation in relative tail Macaca mulatta adult noncaptives 34 length in adult Macaca mulatta 37 10. Longitudinal variation in tail length in 6. Dry-skin measurements of 14 Macaca Macaca mulatta adult noncaptives 35 mulatta specimens collected at Xing- relative tail 1 1 Latitudinal variation in long Xian (= Eastern Tombs), Hebei in mulatta adult non- length Macaca Province, northeastern China 39 captives 36 7. Geographic variation of body weight in 12. Longitudinal variation in relative tail adult Macaca mulatta 40 length in Macaca mulatta adult non- 8. Cranial measurements and proportions captives 38 in age/sex classes of wild-collected Latitudinal variation in body weight in 13. Macaca mulatta 44 Macaca mulatta adult noncaptives 41 9. Geographic variation in greatest length 14. Skull of adult female Macaca mulat- of skull in adult Macaca mulatta 46 ta FMNH 99668, Thailand: Ban Mae — 10. Frequencies of major alleles at poly- Lamao 42 morphic blood-protein loci in samples 15. Skull of adult male Macaca mulatta— of Macaca w«/arra from six countries .... 53 FMNH 99669, Thailand: Huai Ap Nang 43 11. Habitats reported for Macaca mulatta ... 55 16. Dental emergence chronology in Maca- 12. Frequency distribution of elevation rec- mulatta 45 ca ords of Macaca mulatta 56 17. Latitudinal variation in greatest length 13. Arboreal/terrestrial behavior recorded of skull in mulatta adult non- Macaca during daylight hours in samples of 47 captives Macaca mulatta 56 18. Longitudinal variation in greatest 14. Group size reported for Macaca mulat- length of skull in Macaca mulatta ta 58 adult noncaptives 48 15. Ratio of sexually mature males to sex- 19. Sample areas cited in mitochondrial ually mature females reported in and Shi 50 DNA study of Zhang groups of Macaca mulatta 60 20. Consensus dendrogram of mitochondri- 16. Home range area reported for groups al relationships in Macaca mulat- DNA of Macaca mulatta 61 ta samples studied by Zhang and Shi .... 51 17. Day range reported for groups of Ma- 21. Type localities of nominal species or caca mulatta 62 subspecies allocated to Macaca mulat- 18. Population density reported for Macaca ta; limits of natural distribution known mulatta 63 of Macaca w»/arto also indicated 82 19. Foods reported eaten by Macaca mu- 22. Hypothetical stages in the evolution latta. Dietary proportions are indicated and dispersal of A/acaca mii/arra 88 where data are available 64 20. Daily waking-hour time budget esti- 27. Infant mortality rate in natural popula- mates for Macaca mulatta 66 tions of Macaca mulatta 78 21. Intergroup contact behavior reported 28. Population growth rate in Macaca mu- for Macaca mulatta 67 latta 80 22. Mating and birth periods reported for 29. Localities and ages of Macaca mulatta natural populations of Macaca mulatta 69 fossils or subfossils 81 23. Age of sexual maturity reported for 30. Subspecies recognized in published natural populations of Macaca mulatta 70 classifications of Macaca mulatta, 24.