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< OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BiaOPv HFC 161996 NO. VOLS. 3ARD ECONOMY THESIS LEAF ATTACH. I THIS TITLE D ECONOMY AUTH. 1ST D ~ OR TITLE I.D. COLOR 20706 ^ffiT^W IS ISSN. BIX 3 - SYSID S^NCY*"* -BL- EW SERIES Systematic Review of Southeast Asian Longtail M&cm 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 Research Associate Division of Mammals Department of Zoology Field Museum of Natural History Chicago, Illinois 60605-2496 Accepted January 13, 1995 Published November 30, 1995 Publication 1470 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1995 Field Museum of Natural History Library of Congress Catalog Card Number: 95-61092 ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents Subspecies Accounts 70 Macaca fascicularis fascicularis (Raffles, [1821]) 70 Abstract 1 Macaca fascicularis aurea I. Geoffroy, Introduction 1 [1831] 77 Geographic Distribution 2 Macaca fascicularis philippinensis I. Pelage 3 Geoffroy, [1843] 79 General Characterization 3 Macaca fascicularis umbrosa Miller, Sex and Age Variation 11 1902b 79 Geographic Variation 11 Macaca fascicularis fusca Miller, Dorsal Pelage Color Saturation 11 1903a 81 Dorsal Pelage Erythrism 13 Macaca fascicularis lasiae (Lyon, Crown Color 15 1916) 81 Lateral Facial Crest Pattern 18 Macaca fascicularis atriceps Kloss, External Measurements and Propor- 1919c 82 tions 21 Macaca fascicularis condorensis Kloss, Sex and Age Variation 21 1926 84 Geographic Variation 22 Macaca fascicularis tua Kellogg, Head and Body Length 22 1944 86 Tail Length 23 Macaca fascicularis karimondjawae Cranial Characters 30 Sody, 1949 86 Sex and Age Variation 30 Macaca fascicularis subspecies undeter- Geographic Variation 33 mined 87 Molecular Biology and Genetics 35 Evolution and Dispersal 87 > Mitochondrial DNA 35 Stage I, 1 Ma 88 Nuclear DNA 39 Stage II, ca. 160 Ka 88 Multiple Alpha-Globin Genes 39 Stage III, > 18 Ka 90 Highly Repeated Restriction Patterns ... 41 Stage IV, ca. 18 Ka 95 Blood Proteins 42 Stage V, < 18 Ka 95 Blood Groups 45 Stage VI, ca. 4.5 Ka 96 Serum Cholesterol Response 46 Acknowledgments 97 Karyology 47 Literature Cited 98 Diseases 47 Appendix 1 : Specimens Examined 1 20 Malaria 47 Appendix 2: Gazetteer of Non-Philippine Simian T-Lymphotrophic Retrovirus, Macaca fascicularis Localities 1 28 Type 1 49 Appendix 3: Dorsal Pelage Color Satur- Susceptibility to Attenuated Polio Virus ... 49 ation in Fringing-Island Samples of Natural History 49 Macaca fascicularis Compared with Habitats 49 Saturation in Core-Area Reference Arboreality/Terrestriality 50 Samples 181 Swimming 51 Appendix 4: Dorsal Pelage Color Troop Size and Composition 51 Erythrism in Fringing-Island Home Range, Day Range 51 Samples of Macaca fascicularis Density 53 Compared with Erythrism in Diet 54 Core-Area Reference Samples 1 84 Predators 54 Appendix 5: Crown Color Pattern Intertroop Behavior 56 Frequency in Samples of Macaca Interspecific Behavior 57 fascicularis That Include Reproduction 58 Dark-Crowned Specimens 187 Fossils and Subfossils 63 Appendix 6: Geographic Variation of Systematics 65 Lateral Facial Crest Pattern in Subspecific Taxonomy 65 Samples of Macaca fascicularis .... 188 Key to Recognized Subspecies 69 Appendix 7: Head and Body Length in in Fringing-Island Samples of Macaca 7. Crown color pattern in dark-crowned fascicularis compared with head Macaca fascicularis 18 and Body Length in Core-Area 8. Infrazygomatic and transzygomatic lat- Reference Samples Collected at eral facial crest patterns in Macaca fas- Similar Latitudes 1 90 cicularis 19 Appendix 8: Tail Length in Fringing- 9. Geographic variation of lateral facial crest Island Samples of Macaca fascicu- pattern in samples of Macaca fascicularis laris Compared with Tail Length collected in Indochinese Peninsula, Isth- in Core-Area Reference Samples mus of Kra, Malay Peninsula, and neigh- Collected at Similar Latitudes 193 boring islands 21 Appendix 9: Relative Tail Length in Fring- 10. Latitudinal variation of head and body ing-Island Samples of Macaca fas- length in adult core-area specimens of cicularis Compared with Relative Macaca fascicularis 24 Tail Length in Core-Area Reference 1 1 . Latitudinal variation of head and body Samples Collected at Similar length in adult shallow-water fringing-is- Latitudes 196 land specimens of Macaca fascicularis Appendix 10: Blood-Protein Allele Fre- compared with that in adult core-area quencies at Polymorphic Loci in 1 1 specimens 26 Geographic Samples of Macaca 12. Latitudinal variation of head and body fascicularis 198 length in adult deep-water fringing-island Appendix 1 1 : Monthly Distribution of specimens of Macaca fascicularis com- Recorded Reproductive Events in pared with that in adult core-area speci- Natural Populations of Macaca mens 27 fascicularis 200 1 3. Latitudinal variation of tail length in adult Appendix 12: Subspecific Variation of core-area specimens of Macaca fascicu- Head and Body Length, Relative laris 29 Tail Length, and Greatest Length 14. Latitudinal variation of relative tail length of Skull in Macaca fascicularis .... 203 in adult core-area specimens of Macaca Subject Index 204 fascicularis 30 Index to Scientific Names 206 15. Latitudinal variation of tail length in adult shallow-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens 31 16. Latitudinal variation of tail length in adult List of Illustrations deep-water fringing-island specimens of Macaca fascicularis compared with that in adult core-area specimens 32 tail 1 . Known locality records of Macaca fasci- 17. Latitudinal variation of relative length cularis 3 in adult shallow-water fringing-island 2. Detail maps of non-Philippine localities specimens of Macaca fascicularis com- of Macaca fascicularis pared with that in adult core-area speci- A. Northwestern quadrant 4 mens 34 B. Southwestern quadrant 6 1 8. Latitudinal variation of relative tail length C. Southeastern quadrant 9 in adult deep-water fringing-island speci- 3. Approximate extent of dry land in South- mens ofMacacafascicularis compared with east Asia during last glacial maximum, ca. that in adult core-area specimens 35 18 Ka 12 1 9. Skull of adult female Macacafascicularis, 4. Core area, shallow-water fringing islands near topotype 36 and deep-water fringing islands inhabited 20. Skull of adult male Macaca fascicularis, by Macaca fascicularis 13 near topotype 37 5. External characters in Macaca fascicu- 21. Latitudinal variation of greatest length of laris 14 skull in adult core-area specimens of Ma- 6. Dorsal pelage color saturation extremes caca fascicularis 40 in Macaca fascicularis 15 22. Latitudinal variation of greatest length of skull in adult shallow-water fringing- List of Tables island specimens of Macaca fascicularis compared with that in adult core-area specimens 41 23. Latitudinal variation of greatest length of 1 . Dorsal pelage color saturation in core-area skull in adult deep-water fringing-island Macaca fascicularis: summary of varia- specimens of Macaca fascicularis com- tion 16 pared with that in adult core-area speci- 2. Dorsal pelage color saturation in core-area mens 43 Macacafascicularis: geographic variation 24. Blood-protein dendrograms for geograph- of mean value in 2-degree latitude-lon- ic samples of Macaca fascicularis: Den- gitude blocks 16 drogram that excludes Mauritius sample 3. Dorsal pelage erythrism in core-area compared with dendrogram that includes Macaca fascicularis: summary of varia- Mauritius sample; consensus dendrogram tion 17 47 4. Dorsal pelage erythrism in core-area Ma- 25. Type localities and limits of distribution caca fascicularis: geographic variation of of recognized subspecies of Macaca fas- mean value in 2-degree latitude-longitude cicularis and type localities of synony- blocks 17 mous nominal species or subspecies ... 64 5. Crown color pattern in Macaca fascicu- 26. Variation ofgreatest length ofskull in adult laris: summary of variation 18 Macaca fascicularisfusca (Indonesia: Pu- 6. Lateral facial crest pattern in Macacafas- lau Simeulue) compared with variation in cicularis: summary of variation 20 adult Sumatran M. f fascicularis 83 7. External measurements and proportions 27. Tail length vs. head and body length in in age/sex classes of wild-collected Ma- Macaca fascicularis fusca and M. f la- caca fascicularis 22 siae 83 8 . Head and body length in core-area Macaca 28. Craniometric comparisons of adult Ma- fascicularis: summary of variation 23 caca fascicularis atriceps, M. f condor- 9. Summary comparison of head and body ensis, and mainland Indochinese M. fas- length in fringing-island and core-area cicularis 85 samples of Macaca fascicularis 25 29. Latitudinal variation of adult head and 10. Tail length and relative tail length in core- body length in samples of Macaca fasci- area Macaca fascicularis: summary of cularis collected in the southern part of variation 28 the Indochinese Peninsula and adjacent 1 1 . Summary comparison of tail length and Isthmus of Kra, north of 10°N, compared relative tail length in fringing-island and with that in samples of parapatric hi. mu- core-area samples of Macaca fascicularis latta collected in the northern part of the 33 Indochinese Peninsula 92 1 2. Cranial measurements and proportions in 30. Latitudinal variation of adult greatest age/sex classes of wild-collected Macaca length of skull in samples of Macaca fas- fascicularis