OF < ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BiaOPv HFC 161996 3ARD ECONOMY THESIS TNHOI.SVTOILTSL.E LEAFATTACH. I D ECONOMY A~UTH.1ST D OR TITLEI.D. ^ffiCOLTOR^W 20706 IS ISSN. BIX 3 SYSID - S^NCY*"* -BL- EW SERIES Systematic Review of Southeast Asian M&cm Longtail Macaca fascicular™ (Raffles, [11 590 5 . Confabs :d for puti FIELDIANA Zoology NEW SERIES, NO. 81 Systematic Review of Southeast Asian Longtail Macaques, Macaca fascicularis (Raffles, [1821]) Jack Fooden ResearchAssociate DivisionofMammals DepartmentofZoology FieldMuseumofNaturalHistory Chicago, Illinois60605-2496 Accepted January 13, 1995 Published November 30, 1995 Publication 1470 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1995 Field Museum ofNatural History LibraryofCongress Catalog CardNumber: 95-61092 ISSN 0015-0754 PRINTED IN THE UNITED STATES OFAMERICA Table ofContents Subspecies Accounts 70 Macaca fascicularisfascicularis (Raffles, [1821]) 70 Abstract 1 Macacafascicularis aurea I. Geoffroy, Introduction 1 [1831] 77 Geographic Distribution 2 Macacafascicularisphilippinensis I. Pelage 3 Geoffroy, [1843] 79 General Characterization 3 Macacafascicularis umbrosa Miller, Sex and Age Variation 11 1902b 79 Geographic Variation 11 Macacafascicularisfusca Miller, Dorsal Pelage Color Saturation 11 1903a 81 Dorsal Pelage Erythrism 13 Macacafascicularis lasiae (Lyon, Crown Color 15 1916) 81 Lateral Facial Crest Pattern 18 Macacafascicularis atriceps Kloss, External Measurements and Propor- 1919c 82 tions 21 Macacafascicularis condorensis Kloss, Sex and Age Variation 21 1926 84 Geographic Variation 22 Macacafascicularis tua Kellogg, Head and Body Length 22 1944 86 Tail Length 23 Macacafascicularis karimondjawae Cranial Characters 30 Sody, 1949 86 Sex and Age Variation 30 Macacafascicularis subspecies undeter- Geographic Variation 33 mined 87 Molecular Biology and Genetics 35 Evolution and Dispersal 87 Mitochondrial DNA 35 Stage I, > 1 Ma 88 Nuclear DNA 39 Stage II, ca. 160 Ka 88 Multiple Alpha-Globin Genes 39 Stage III, > 18 Ka 90 Highly Repeated Restriction Patterns ... 41 Stage IV, ca. 18 Ka 95 Blood Proteins 42 Stage V, < 18 Ka 95 Blood Groups 45 Stage VI, ca. 4.5 Ka 96 Serum Cholesterol Response 46 Acknowledgments 97 Karyology 47 Literature Cited 98 Diseases 47 Appendix 1: Specimens Examined 120 Malaria 47 Appendix 2: Gazetteer ofNon-Philippine Simian T-Lymphotrophic Retrovirus, Macaca fascicularis Localities 128 Type 1 49 Appendix 3: Dorsal Pelage Color Satur- Susceptibilityto Attenuated Polio Virus ... 49 ation in Fringing-Island Samples of Natural History 49 Macaca fascicularis Compared with Habitats 49 Saturation in Core-Area Reference Arboreality/Terrestriality 50 Samples 181 Swimming 51 Appendix 4: Dorsal Pelage Color Troop Size and Composition 51 Erythrism in Fringing-Island Home Range, Day Range 51 Samples of Macaca fascicularis Density 53 Compared with Erythrism in Diet 54 Core-Area Reference Samples 184 Predators 54 Appendix 5: Crown Color Pattern Intertroop Behavior 56 Frequency in Samples of Macaca Interspecific Behavior 57 fascicularis That Include Reproduction 58 Dark-Crowned Specimens 187 Fossils and Subfossils 63 Appendix 6: Geographic Variation of Systematics 65 Lateral Facial Crest Pattern in Subspecific Taxonomy 65 Samples of Macaca fascicularis .... 188 Key to Recognized Subspecies 69 Appendix 7: Headand Body Length in in Fringing-Island Samples of Macaca 7. Crown color pattern in dark-crowned fascicularis compared with head Macacafascicularis 18 and Body Length in Core-Area 8. Infrazygomatic and transzygomatic lat- Reference Samples Collected at eral facial crest patterns in Macacafas- Similar Latitudes 190 cicularis 19 Appendix 8: Tail Length in Fringing- 9. Geographicvariationoflateralfacialcrest Island Samples of Macaca fascicu- patterninsamplesofMacacafascicularis laris Compared with Tail Length collected in Indochinese Peninsula, Isth- in Core-Area Reference Samples mus ofKra, Malay Peninsula, and neigh- Collected at Similar Latitudes 193 boring islands 21 Appendix9:RelativeTailLengthinFring- 10. Latitudinal variation of head and body ing-Island Samples of Macaca fas- length in adult core-area specimens of cicularis Compared with Relative Macacafascicularis 24 Tail Length in Core-Area Reference 11. Latitudinal variation of head and body Samples Collected at Similar length in adult shallow-water fringing-is- Latitudes 196 land specimens of Macaca fascicularis Appendix 10: Blood-Protein Allele Fre- compared with that in adult core-area quencies at Polymorphic Loci in 1 1 specimens 26 Geographic Samples of Macaca 12. Latitudinal variation of head and body fascicularis 198 length inadultdeep-waterfringing-island Appendix 11: Monthly Distribution of specimens of Macaca fascicularis com- Recorded Reproductive Events in pared with that in adult core-area speci- Natural Populations of Macaca mens 27 fascicularis 200 13. Latitudinalvariationoftaillengthinadult Appendix 12: Subspecific Variation of core-area specimens of Macacafascicu- Head and Body Length, Relative laris 29 Tail Length, and Greatest Length 14. Latitudinal variationofrelativetaillength of Skull in Macaca fascicularis .... 203 in adult core-area specimens ofMacaca Subject Index 204 fascicularis 30 Index to Scientific Names 206 15. Latitudinalvariationoftaillengthinadult shallow-water fringing-island specimens ofMacacafasciculariscomparedwiththat in adult core-area specimens 31 16. Latitudinalvariationoftaillengthinadult List of Illustrations deep-water fringing-island specimens of Macacafascicularis compared with that in adult core-area specimens 32 1. Known locality records ofMacacafasci- 17. Latitudinal variationofrelativetaillength cularis 3 in adult shallow-water fringing-island 2. Detail maps ofnon-Philippine localities specimens of Macaca fascicularis com- ofMacacafascicularis pared with that in adult core-area speci- A. Northwestern quadrant 4 mens 34 B. Southwestern quadrant 6 18. Latitudinalvariationofrelativetaillength C. Southeastern quadrant 9 in adult deep-water fringing-island speci- 3. ApproximateextentofdrylandinSouth- mensofMacacafasciculariscomparedwith eastAsiaduringlastglacialmaximum,ca. that in adult core-area specimens 35 18 Ka 12 19. SkullofadultfemaleMacacafascicularis, 4. Core area, shallow-water fringing islands near topotype 36 anddeep-waterfringingislandsinhabited 20. Skull ofadult male Macacafascicularis, by Macacafascicularis 13 near topotype 37 5. External characters in Macaca fascicu- 21. Latitudinal variation ofgreatest length of laris 14 skullinadultcore-areaspecimensofMa- 6. Dorsal pelage color saturation extremes cacafascicularis 40 in Macacafascicularis 15 22. Latitudinal variation ofgreatestlength of