Biology 590.5 Fl N.S. no. 104(2005) AUG ^ 3 2005 HI ^[yr\ Zoology NEW SERIES, NO. 104 Systematic Review of Japanese Macaques, Macaca fuscata (Gray, 1870) Jack Fooden Mitsuru Aimi March 14, 2005 Publication 1533 PUBLISHED BY FIELD MUSEUM OF NATURAT HISTORY Information for Contributors to Fieldiana Fieldianu ... .. ^ ... ..,^...^..... ,.....s published b} .... . ..... . ... .,....., publishes the research of staff members and our research associates. The page charge currently is S65 per page. This figure is subject to change. All authors are encouraged to provide fundin ^,^ cn.^n,^rt \h.- '-"'xluction of tHclr works. Submission procedures: A submission procedures document is available from the scientific -ditor of the journal and on the Museum's web site (by 2004). Manuscripts submitted for re\ ic should be prepared using standard electronic software and submitted to the scientific editor in three photocopies. 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The publication of long works should be arranged well in advance with the scientific editor. Please bear in mind: Changes on page proofs are very expensive. Author-generated chanin s . ,i page proofs can only be made if the author agrees in advance to pay for them. UNIVERSITY OF ILLINOIS LIBRARY AT URBANA-CHAMPAIGN BIOLOGY FIELDIANA Zoology NEW SERIES, NO. 104 Systematic Review of Japanese Macaques, Macaca fuscata (Gray, 1870) Jack Fooden Mitsuru Aimi Division of Mammals Department of Evolution and Phylogeny Department of Zoology Primate Research Institute Field Museum of Natural History Kyoto University 1400 South Lake Shore Drive Inuyama, Aichi 484-8506 Chicago, Illinois 60605-2496 Japan U.S.A. Accepted August 3, 2004 Published March 14, 2005 Publication 1533 BIOLOGY LIBRARY 101 BURRILL HALL AUG 0 3 2005 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 2005 Field Museum of Natural History ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents Abstract 1 Introduction 1 Geographic Distribution, Total Popula- tion Estimate 3 External Characters 5 Pelage 5 General Characterization 5 Early Development 5 Seasonal Variation 5 Geographic Variation 6 Albinism 8 External Measurements and Proportions .. 8 Linear Measurements of Body Size .... 8 Tail Length 11 Hind Foot Length 12 Body Weight 13 Summary 13 Dermatoglyphics 15 Congenital Malformation of Limbs 15 Cranial Characters 17 Effect of Provisioning 17 Sex and Age Variation 17 Geographic Variation 19 Interspecific Comparisons 21 Molecular Biology and Genetics 22 DNA: Intraspecific Studies 22 Mitochondrial DNA 22 Minisatellite DNA and Microsatellite DNA 33 DNA: Interspecific Studies 33 Mitochondrial DNA 33 Nuclear DNA: Noncoding Intron 33 Nuclear DNA: Y-Chromosome 35 Nuclear DNA: Random Amplified Poly- morphism 35 Mitochondrial DNA-Nuclear DNA: Multiple Loci 35 Blood Proteins 37 Hemoglobin Amino- Acid Sequence 42 Antibodies to Cedar Pollen Antigens 43 Karyology 45 Parasites 46 Protistans 46 Trematodes 47 Cestodes 47 Nematodes 47 Mite 49 Lice 49 Tick 49 Q Fever Bacterium 49 Viruses 49 Natural History 50 Habitats 50 Terrestriality/Arboreality 51 Group Size and Composition 51 Home Range Area and Group Density ... 52 Diet 52 Predators 54 Intergroup Dispersal 54 Reproduction 54 Seasonality 54 Sexual Maturation 57 Sexual Skin 57 Menstrual Cycle 58 Estrus 58 Consortship 59 Copulatory Behavior 60 Dominance Rank and Reproductive Suc- cess 60 Inbreeding 61 Hybridization 61 Nonreproductive Sexual Behavior 62 Gestation 63 Parturition 64 Neonatal Sex Ratio; Birth Weight 65 Twinning 66 Birthrate 66 Infant Mortality 68 Nursing; Weaning 68 Longevity; Reproductive Senescence 71 Population Growth Rate 72 Fossils 72 S ystematics 72 Geographic Variation and Subspecific Recognition 72 Pelage Color 75 Body Size 75 Cranial Morphology 75 Mitochondrial DNA; Blood Proteins ... 75 Key to External Characters of Recog- nized Subspecies 77 Japanese Vernacular Names Applied to Macaca fuscata 77 Subspecies Accounts: Macaca fuscata fuscata (Gray, 1870) 77 Synonymy 77 Type Series 79 Type Locality 80 Distribution 81 Diagnosis 81 Remarks 81 Specimens Examined 82 Macaca fuscata yakui Kuroda, 1 940 82 Synonymy 82 Type Series 82 ui Type Locality 83 Distribution 83 Diagnosis 83 Specimens Examined 83 Evolution and Dispersal 83 Acknowledgments 86 Literature Cited 86 Appendix L Specimens Examined 105 Appendix 2: Macaca fuscata Localities Reported in Postal Survey' Conduct- ed BY K. Hasebe in 1923 108 Appendix 3. Locality Records (Specimens Examined and Literature Records) OF Macaca fuscata 113 Appendix 4: Gazetteer of Macaca fuscata Collection and Observation Locali- ties 119 Appendix 5: Local Variation in Sitting Height in Macaca fuscata Adults on Honshu 191 Appendix 6: Local Variation in Anterior Trunk Length in Macaca fuscata Adults on Honshu 192 Appendix 7: Local Variation in Tail Length and Tail Length/Anterior Trunk Length ratio in Macaca fusca- ta Adults on Honshu 194 Appendix 8: Local Variation in Hind Foot Length and Hind Foot/Anteri- or Trunk Length Ratio in Macaca fuscata Adults on Honshu 196 Appendix 9: Local Variation in Body Weight in Macaca fuscata Adults on Honshu 197 Appendix 10: Local Variation in Great- est Length of Skull in Macaca fus- cata Adults on Honshu 198 Index 199 Note Added in Proof 200 List of Illustrations 1. Macaca fuscata depicted in first scroll (detail) of four-scroll Choju-jinbutsu- giga set 2 2. Survey reports of geographic distribu- tion of Macaca fuscata A. Survey conducted in 1923 by K. Hasebe 4 B. Survey conducted in 1992 by Envi- ronmental Agency of Japan 4 3. Macaca fuscata — adult male, adult fe- male, and infant — photographed at Takasakiyama, Oita Prefecture, Kyushu 6 4. Latitudinal variation in head and body length in Honshu and Shikoku samples of Macaca fuscata aduhs, 10 5. Latitudinal variation in sitting height in Macaca fuscata adults 12 6. Latitudinal variation in anterior trunk length in Macaca fuscata adults 14 7. Latitudinal variation in tail length in Macaca fuscata adults 16 8. Tail length vs. head and body length in Macaca fuscata immatures and adults ' 18 9. Latitudinal variation in hind foot length m Macaca fuscata aduXis 20 10. Latitudinal variation in body weight in Macaca fuscata adults 22 1 1 . Skull of adult female Macaca fuscata fuscata 26 12. Skull of adult female Macaca fuscata yakui 27 13. Skull of adult male Macaca fuscata fuscata 28 14. Skull of adult male Macaca fuscata yakui 29 15. Latitudinal variation in greatest length of skull in Macaca fuscata adults 32 16. Greatest length of skull vs. estimated age in Honshu and Yakushima speci- mens of Macaca fuscata 34 17. Orbital height vs. orbital breadth in in- sular samples of Macaca fuscata adults 36 18. Geographic variation in mtDNA se- quence haplotypes in Macaca fuscata (Hayasaka et al., 1991) 38 19. Geographic variation in mtDNA se- quence haplotypes in Macaca fuscata (Kawamoto, 2002) 41 20. Geographic distribution and genetic in- terrelationships of blood-protein groups in Macaca fuscata 43 21. Distribution oi Macaca fuscata troops in Japan for which birth seasonality data are available 55 22. Latitudinal variation in birth seasonali- ty in in-situ troops of Macaca fuscata 56 23. Age-specific birthrates in two nonpro- visioned troops and two provisioned troops of Macaca fuscata 70 IV 24. Distribution of fossil macaque locali- ties in Japan 76 25. Bathymetric map of Yakushima-Sakha- lin island-chain area, illustrating proba- ble land bridges exposed by sea-level depression of 140 m 84 List of Tables 1 . Estimates of total extant population of Macaca fuscata 5 2. Geographic variation in mean paleness index values of pelage on dorsal sur- face of trunk, dorsal surface of hands, and dorsal surface of feet in Macaca fuscata 7 3. Months during which molting was ob- served in nine Macaca fuscata captives maintained outdoors at the Japan Mon- key Centre, Inuyama, Aichi Prefecture, Honshu 7 4. Dorsal hair density in infant and adult Macaca fuscata 8 5. Prefectural variation in head and body length and related external measure- ments in Macaca fuscata 9 6. Insular variation in sitting height in Macaca fuscata adults 11 7. Insular variation in anterior trunk length in Macaca fuscata adults 13 8. Insular variation in tail length and tail length/anterior trunk length ratio in Macaca fuscata adults 15 9. External measurements in seven Maca- ca fuscata adults with unusually short tails 17 10. Insular variation in hind foot length and hind foot length/anterior trunk length ratio in Macaca fuscata adults .. 19 1 1 . Insular variation in body weight in Macaca fuscata adults 21 12. Frequency of dermatoglyphic patterns on both hands in five local samples of Macaca fuscata 23 13. Provisioning status and occurrence of congenital limb malformations in 65 Macaca fuscata groups 23 14. Neonatal incidence of congenital limb malformations in nine Macaca fuscata groups 24 15. Sexual variation in incidence of con- genital limb malformations in Macaca fuscata neonates 24 16. Greatest length of skull in sample of provisioned Macaca fuscata population collected at Takagoyama, Chiba Prefec- ture, Honshu, compared with that in sample of nonprovisioned populations collected elsewhere in Chiba Prefecture 30 17. Cranial measurements and proportions in age/sex classes of nonprovisioned wild-collected Macaca /Micara 30 18. Dental emergence chronology in Maca- ca fuscata 31 19. Insular variation in greatest length of skull (mm) in wild-collected Macaca fuscata 31 20. Insular variation in head length (great- est midsagittal length from glabella to occiput) in living Macaca fuscata 33 21. Insular variation in orbital measure- ments and orbital height/breadth index in adult wild-collected Macaca fuscata 35 22. Greatest length of skull and rostral- postrostral ratio \n fascicularis-gxoxyp species 37 23. Local variation in mtDNA sequence haplotypes in Macaca fuscata (Hayasa- kaet al., 1991) 39 24. Mitochondrial DNA haplotypes detect- ed in Macaca fuscata at localities sam- pled by Kawamoto (2002) 40 25. Mean mtDNA divergence estimates re- ported between Macaca fuscata and M. mulatta, M. cyclopis, and M. fascicu- laris 42 26. Sister-group relationships of Macaca fuscata indicated by sequencing of pa- ternal, maternal, and biparental markers 42 27. Mean frequency of major alleles at 2 loci in blood-protein groups of Macaca fuscata and frequency of corresponding alleles in southern Chinese M. mulatta 44 28. Incidence of serum specimens positive for antibodies to crude cedar pollen an- tigen and incidence of serum speci- mens with positive reactions to purified cedar pollen antigens in prefectural samples of Macaca fuscata 46 29. Incidence of protistan parasite species detected in fecal samples of Macaca fuscata on Shimokita Peninsula, Ao- mori Prefecture 46 30. Incidence of cestode parasites in local samples of Macaco fuscata 47 3 1 . Incidence of nematode parasites in lo- cal samples of Macaco fiiscata 48 32. Incidence of antibodies to eight viruses in three local groups of Macaco fusca- ta ^ 50 33. Relationship between housing system and incidence of antibodies to hepatitis A in captive Macaco fiiscata 51 34. Geographic variation in group size in wild Mococa fuscata populations 52 35. Group composition and ratios in wild Macaco fuscata 53 36. Menstrual cycle length in Macaco fus- cata 58 37. Frequency and duration of estrous peri- ods in mature Macaco fuscata females 59 38. Reported results concerning relation- ship between male dominance rank and reproductive success in natural and seminatural groups of Macaco fuscato 60 39. References concerning nonreproductive sexual behavior reported in natural and seminatural groups of Macaco fuscata 62 40. Gestation period duration in Macaco fuscata 64 41. Neonatal sex ratio in Macaco fuscata .. 65 42. Birth weight in Afacaca yzY5cara 66 43. Annual birthrate in Macaca yM5cara .... 67 44. Effect of Macaco fuscata infant death on interbirth interval of mother 68 45. Relationship between birthrate and fe- male dominance rank in nonprovi- sioned and provisoned groups of Ma- coca fuscato 68 46. Infant mortality rate in nonprovisioned and provisioned groups of Macaco fus- cato 69 47. Mean life span in Macaco fuscato fe- males 71 48. Population growth rate in Mococa fiis- cata groups 73 49. Fossil macaque localities in Japan, with indication of associated fossil pro- boscideans 74 50. Mammal stages, biozones. and stratig- raphy of Quaternary deposits in Japan 75 51. Dental measurements in four Korean fossil macaque specimens collected at Turubong Cave and one Japanese fossil macaque specimen collected at Ando Quarry 77 52. Dental measurements in samples of ex- tant Mococa fuscata, for comparison with corresponding measurements in Korean and Japanese fossil macaques 78 53. Distribution relative to Tsugaru Strait (between Honshu and Hokkaido) of Yakushima-Sakhalin species of nonvo- lant terrestrial mammals 83 VI Systematic Review of Japanese Macaques, Macaca fuscata (Gray, 1870) Jack Fooden Mitsuru Aimi Abstract Japanese macaques, Macaca fuscata (Gray, 1 870), are systematically reviewed, based on examination of 1264 specimens, survey of relevant literature, and observation of natural pop- ulations. This review includes analyses of external and cranial characters, molecular biology and genetics, and parasites. Information also is presented concerning natural history, reproduc- tion, fossils, and taxonomic history. Concerning the evolutionary history of M. fuscata, avail- able evidence suggests that this species is derived from a M. mulatta-like population that dispersed to the Japanese archipelago during one or two glacial intervals of sea-level depres- sion, ca. 0.63-0.43 million years ago (Ma), via a now-submerged dry-land connection between the Korean Peninsula and the Kyushu-West Honshu area. In an appendix, an annotated gaz- etteer lists 1347 localities at which M. fuscata has been collected or observed. Introduction Macaca fuscata is a stump-tailed monkey that is unique to the Japanese archipelago and is the only nonhuman primate that inhabits these is- lands. Its geographic distribution, which extends from ca. 30°15'N to 4r30'N, is the northernmost of any extant nonhuman primate. Although Japanese artists have been fascinated by the morphology and behavior of M. fuscata for at least eight centuries (Fig. 1; cf. Ohnuki-Tier- ney, 1987, p. 27), Western science apparently did not become aware of the existence of this species until 1727, when it was briefly mentioned by E. Kaempfer (1727, p. 126; cf. Bodart-Bailey, 1999, p. 72) in his book The History of Japan . . . , Kaempfer, a German physician, encountered M. fuscata while serving with a Dutch trading mis- sion in Japan during the period 1690-1692. The first scientific specimens of M. fuscata were sent to Europe in 1821-1822 by C. G. Reinwardt (let- ters, RMNH archives) and in 1823-1829 by R E von Siebold (Holthuis & Sakai, 1970, p. 25). The first scientific account of the species was pub- lished in 1825 by E Cuvier (1824-1829, livraison 47, p. 1), based on a captive exhibited in a zoo in India. The present review of M. fuscata is based on study of 1264 specimens (Appendix 1), survey of relevant literature, and observation of natural pop- ulations. Specimens examined are preserved in in- stitutions listed below, which hereafter are cited by means of the indicated abbreviations; the num- ber of M. fuscata specimens studied at each in- stitution is indicated by a parenthetical notation. AIUM Anthropologisches Institut der Univer- sitat Miinchen (32) AIUZ Anthropologisches Institut der Univer- sitat Zurich (2) AMNH American Museum of Natural History, New York (3) ANSP Academy of Natural Sciences, Phila- delphia (3) BM(NH) The Natural History Museum, London (12) JMC Japan Monkey Centre, Inuyama, Japan (39) KFL Koshima Eield Laboratory, Kushima, Japan (48) MCZ Museum of Comparative Zoology, Har- vard University, Cambridge, Massachu- setts (2) FIELDIANA: ZOOLOGY, N.S., NO. 104, MARCH 14, 2005, PP. 1-200 f X ^* Ji ^ ^ mf ' "^ — \ V iw*-' T-^ Fig. 1. Macaca fuscata depicted in first scroll (detail) of four-scroll Choju-jinbutsu-giga set, in collection of Kozanji monastery. Kyoto (cf. Ohnuki-Tierney, 1987, p. 28); attributed to Toba Sojo, also known as Kakuyu (1053-1140), noted priest and artist. (Reproduced by permission of Kozanji monastery.) FIELDIANA: ZOOLOGY MNHN Museum National d'Histoire Naturelle, Paris (5) MUE Miyagi University of Education, Sen- dai, Japan (15) NHR Naturhistoriska Riksmuseet, Stockholm (2) PRIKU Primate Research Institute, Kyoto Uni- versity, Inuyama, Japan (895) RMNH Nationaal Naturhistorisch Museum, Leiden (14) SU Saitama University, Urawa, Japan (96) TPM Tochigi Prefectural Museum, Utsuno- miya, Japan (32) USNM National Museum of Natural History, Washington, D.C. (12) ZMB Museum fiir Naturkunde, Berlin (28) ZMUZ Zoologisches Museum der Universitat Zurich (2) ZSBS Zoologisches Staatssammlung, Munich (22) Geographic Distribution, Total Population Estimate The most recent comprehensive survey of the geographic distribution of M. fuscata was con- ducted in 1992 by the Environmental Agency of Japan (now Ministry of the Environment, Japan). In this survey, the land area of the Japanese ar- chipelago was subdivided into a grid of 10 X 10- km cells, for each of which the presence or ab- sence of monkeys was reported by local officials (Fig. 2B); M. fuscata was reported present in ca. 1000 of the 10 X 10-km cells. This survey con- firms that M. fuscata is relatively widely distrib- uted in upland areas on three of the four large Japanese islands — Honshu, Shikoku, and Kyu- shu— and that it also inhabits six close-lying small islands — Kinkazan (off northeastern Honshu), Awajishima and Shodoshima (in the Inland Sea, between Honshu and Shikoku), Kashima (off southwestern Shikoku), Kojima {- Koshima, off southeastern Kyushu), and Yakushima (ca. 60 km south of Kyushu); all six of the small islands in- habited by M. fuscata are within the 100-m bathy- metric line (Iwano, 1974, figs. 1-5). M. fuscata formerly inhabited three additional small islands where it is now extinct (Appendix 3; Kawanaka, 1973, p. 115); these three islands are Omishima (north of westernmost Honshu), where M. fuscata survived until ca. 1965 (Tanaka, 1991, p. 19); Hi- radoshima (northwest of Kyushu), where this spe- cies survived until ca. 1925 (Sakura, 1976, p. 151); and Tanegashima (northeast of Yakushima), where it survived until ca. 1955 (Azuma, 1972, p. 261) and subsequently became extinct as a result of deforestation. The earliest comprehensive survey of the dis- tribution of M. fuscata was conducted in 1 923 by K. Hasebe, but the results of this survey were not published until 1974 (Iwano, 1974, p. 5; cf. Ama- gasa & Ito, 1978, p. 96); somewhat less rigorous surveys were conducted by K. Kishida in 1953 and by H. Takeshita in 1964a,b (Iwano, 1974, p. 8). Hasebe mailed 563 survey questionnaires to heads of each prefecture and county in Japan and received 535 replies that identified a total of 818 monkey localities (Fig. 2A). Although Hasebe's survey preceded the Environmental Agency sur- vey (Fig. 2B) by nearly seven decades, the two surveys reveal basically similar patterns of geo- graphic distribution; in particular, both surveys in- dicate relatively few records of monkeys in three areas that are within the geographic range of M. fuscata — northern Honshu (ca. 39-40. 5°N), north of Boso Peninsula (ca. 35.5-36.5°N), and north- western Kyushu (ca. >33°N, <131°E). A map of localities of M. fuscata specimens and literature records examined for the present study (Appendix 3; cf. Baldwin et al., 1980, p. 269) also reveals a pattern of geographic distribution generally simi- lar to that revealed by the 1 992 survey map. Fac- tors responsible for the sparse distribution of M. fuscata in northern Honshu (ca. 39-40. 5°N) have been analyzed historically by Mito (1992, p. 143; cf. Oi et al., 1997, p. 9; 2000, p. 11). Although M. fuscata is now absent in most lowland areas of Japan (Amagasa & Ito, 1978, p. 102), it presumably inhabited lowland areas be- fore their intensive exploitation by humans (cf. Tokita et al., 1978, p. 88). The greatest elevation at which M. fuscata has been reported is 3 1 80 m (see below. Natural History: Habitats). M. fuscata is absent on Hokkaido (the north- ernmost of the four large Japanese islands), on the Ryukyu archipelago (southwest of Kyushu and Yakushima), and on most small islands that sur- round Honshu, Shikoku, and Kyushu — including most islands within the 100-m bathymetric line (Iwano, 1974, figs. 1-5). One small island — Mi- yajima — is inhabited by an artificially introduced population of M. fuscata, and two other small is- lands— Okinoshima, and Tomogashima — former- ly were inhabited by introduced populations (for details, see Gazetteer, Appendix 4). Fragmentary skeletal evidence indicates that M. fuscata was FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 2?o O o^ U) . c r^ •"" -o II >% u ?^ . 0 3 > * * o ri^ j . ' " U. ^ HELDIANA: ZOOLOGY Table 1 . Estimates of total extant population of Macaca fuscata. Estimated Estimated total number of References population groups Basis of estimates Kishida(1953, p. 65) 15,614 313 Questionnaire survey, 1950 Takeshita (1964b, p. 20) 22,000-34,000 425 Questionnaire survey, 1961-1962 Kawamura (1973, p. 456) 43.161 — Questionnaire survey,' 1970 Kawamura (1975, p. 502)^ 50,000-100,000 — Field survey (incomplete) Hashiba(1989, p. 30) 114,431 — Prefectural survey of broadleaf forest areas Survey conducted by H. Takeshita. Cf. Kawamura (1973, p. 455). translocated from Yakushima to Okinawa (Shuri Castle) several centuries ago (Mouri et al., 2000, p. 87); artificially modified monkey bones discov- ered at an archaeological site on Rebun Island, northwest of Hokkaido, may have been transport- ed to Rebun Island as artifacts (Nishimoto, 1981, p. 430). Prior to ca. 1945, the geographic distribution of M. fuscata apparently was contracting (K. Watan- abe and Y. Muroyama, PRIKU, pers. comm.). Subsequently— despite recent annual extermina- tion of ca. 10,000 monkeys as agricultural pests— the distribution generally has been expanding; this contraction and expansion apparently are related to changing patterns of Japanese human demog- raphy and behavior (cf. Sprague, 2002, p. 255). The extant total population of M. fuscata is esti- mated to be approximately 100,000 individuals (Table 1). External Characters Pelage (Fig. 3) General Characterization — In M. fuscata adults in prime pelage, the color of the crown and dorsal surface of the trunk varies from pale yel- lowish brown to grayish brown to dark golden brown. Fur on the outer surface of the limbs and tail tends to be slightly paler than that on the ad- jacent surface of the trunk (cf. Hamada et al., 1992, p. 6); fur on the dorsal surface of the hands varies from pale yellowish brown to blackish, and that on the feet varies from pale yellowish brown to dark brown. Fur on the ventral surface of the trunk and the inner surface of the limbs varies from whitish to pale gray to pale ochraceous buff. The prominent side-whiskers are somewhat paler than the crown, and the lateral facial crest pattern is infrazygomatic (Fooden, 1995, p. 18). The thin- ly haired facial skin is pinkish to reddish (for dis- cussion of sexual skin, see below. Reproduction). Dorsal pelage color in adult females averages slightly paler than in adult males (Table 2; Ha- mada et al., 1992, p. 5). In a very old free-ranging male (estimated age 38 years), the fur on the haunches and dorsal surface of the trunk report- edly had become "quite white" (Itani, 1967, p. 90). Dorsal pelage is relatively long in M. fuscata. Although mean length of hairs on the back in adults is ca. 40 mm, length of the longest hairs in this region is ca. 70-90 mm (Inagaki, 1985, p. 335; 1986, p. 117; cf. Hori et al., 1977, p. 315). The proximal half of individual dorsal hairs is pale gray; the distal half is marked by one to three pairs of alternating pale and dark annulations (Temminck, 1842, p. 9; Inagaki, 1986, p. 116; 1996, p. 92; Inagaki & Nigi, 1988b, p. 506); pale annulations are yellow to golden, and dark an- nulations are brown to blackish. Early Development — Natal pelage in M. fus- cata is unannulated, grayish brown to dark brown (Forbes, 1894, p. 14; Pocock, 1906, p. 566; Hir- aiwa, 1981, p. 313; Hamada et al., 1992, p. 7; Pavelka, 1993, p. 61). Facial skin in newborns is pale pink, paler than in adults. Replacement of the unannulated pelage of newborns by annulated pel- age, generally similar to that of older monkeys, commences about age 2 months (Takada, 1966, p. 73; Maruhashi, 1982, p. 320), and this replace- ment is complete by age 4-5 months. Seasonal Variation— M/M5cato undergoes an annual molt in late spring/early summer (Table 3). In nine immature and adult captives maintained outdoors at the Japan Monkey Centre, Inuyama, Aichi Prefecture, molting occurred from April to August, and the duration of the molting process FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES Fig. 3. Macaca fuscata — adult male, adult female, and infant — photographed 13 March 2002 at Takasakiyama, Oita Prefecture, Kyushu. (Photo courtesy of Minoru Kinoshita, PRIKU.) in individual monkeys varied from less than 1 month to approximately 4 months. In all five par- turient females in this group, molting occurred af- ter parturition. As a result of the molting cycle, dorsal pelage is relatively short and dark during the summer (distal dark annulations exposed, proximal pale gray region not yet fully exposed) and relatively long and pale during the winter (Mori, 1979b, p. 373; Inagaki and Nigi, 1988a, p. 83; Hamada et al., 1992, p. 19). During late winter/early spring, the annulated tips of the hairs become progres- sively abraded, and the pelage appears faded and scraggly. Geographic Variation — Thomas (1906 ["1905"], p. 361) and Kuroda (1940, p. 273) ap- parently were the first authors to indicate that dor- sal pelage color in M. fuscata tends to be paler in the northern part of its range than in the southern part. Although few localized museum study skins are available to document this cline (see Appen- dix 1), requisite data have been provided by Ha- mada et al. (1992, p. 1), who conducted an inten- sive photometric study of living monkeys of known geographic origin (Table 2). Hamada et al. (1992, p. 9) have shown that, in females and males from Yakushima, Kojima (= Koshima), Kyushu (Takasakiyama), and Honshu, paleness of dorsal pelage color in M. fuscata generally in- creases with increasing latitude and decreasing winter temperature; three museum skins collected on Shikoku (BM(NH) 1906.1.4.1-1906.1.4.3) are relatively dark and therefore suggest that pelage color on this island also is compatible with the cline documented by Hamada et al. (cf. Thomas, 1906 ["1905"], p. 361). As noted by Hamada et FIELDIANA: ZOOLOGY Table 2. Geographic variation in mean paleness index values of pelage on dorsal surface of trunk, dorsal surface of hands, and dorsal surface of feet in Macaca fuscata; greater index values indicate paler pelage.' Locality of group Latitude (N) Longitude (E) Mean January tempera- ture (»C) Mean paleness ; index values- [or original locality of group progenitors]' Females Males Hands, Back dorsum^ Feet, dorsum** Back Hands, dorsum^ Feet, dorsum** Honshu Shiga Nikko [Wakasa] [Takagoyama] [Arashiyama] Hagachi 36°44' 36°44' 35°20' 35° 13' 35°00' 34°4r 138°26' 139°30' 134°24' 139°59' 135°4r 138°44' -4.7 -4.9 1.85 3.7 2.8 5.6 39 45(1.15) 37 41(1.11) 32 38(1.19) 38 40(1.05) 33 39(1.18) 32 31 (0.97) Kyushu 50(1.28) 45(1.22) 43(1.34) 48(1.26) 46(1.39) 37(1.16) 37 33 29 37 29 30 42(1.14) 37(1.12) 38(1.31) 37(1.00) 34(1.17) 32(1.07) 47(1.27) 42(1.27) 45(1.22) 42(1.45) 37(1.23) Takasakiyama 33°10' 131°27' 4.9 30 31 (1.03) Kojima 36(1.20) 27 27(1.00) 32(1.19) Kojima 31°27' 131°23' 6.7 31 31(1.00) Yakushima 36(1.16) 29 32(1.10) 35(1.21) [Yakushima*] Nagata 30°34' 30°34' 130°26' 130°26' 11.2 11.2 31 27(0.87) 31 27(0.87) 33(1.06) 29 (0.94) 29 25 24 (0.83) 23 (0.92) 35(1.21) 26(1.04) ' Reference: Hamada et al. (1992, pp. 4, 10). - Standard deviation, extremes, and sample size not published in cited reference. ' Brackets indicate original localities of progenitors of four sampled groups that had been translocated one or two generations previous to the cited study, as follows: [Wakasa] and [Arashiyama] group progenitors translocated to PRIKU, Inuyama, Aichi Prefecture; [Takagoyama] group progenitors translocated to Higashi-Tsukuba, Utopia Zoo, Yasato-cho, Ibaraki Prefecture; and [Yakushima] group progenitors translocated to JMC, Ohirayama, Inuyama, Aichi Prefecture (cf. Kawai, 1960, p. 183). ■* Hands/back paleness index ratio in parentheses. ^ Feet/back paleness index ratio in parentheses. ^ Hamada et al. (1992, p. 3) suspect that pelage color in this translocated group (Yakushima to Inuyama) may have changed in response to its new environmental conditions. Table 3. Months during which molting (M) was observed in nine Macaca fuscata captives' maintained outdoors at the Japan Monkey Centre, Inuyama, Aichi Prefecture, Honshu; for parturient females, the month of parturition (P) also is indicated. - Age (yr) Sex Spec. no. March April May June July August 2 $ 58 3 6 46 5 9 3 5 9 4 >6 9 43 >6 9 38 >6 9 73 >6 3 5 >6 S 42 M M M M M M P M M P M M M M P M M P/M M M M M M M Origin: Kyoto Prefecture, Honshu. Reference: Inagaki and Nigi (1988a, pp. 82, 83). FOODEN AND AI1V1I: SYSTEMATIC REVIEW OF JAPANESE MACAQUES Table 4. Dorsal hair density (number of hairs per 1 -cm-diameter circle) in infant and adult Macaca fuscata: studv conducted on open-air groups during winter 1983-1984.' Locality Latitude (N) Longitude (E) Mean annual tempera- . ture (°C) Hair density- Infants Adults Island Mean ± SD N Mean ± SD N Honshu Honshu Mivajima Shimokita' Hagachi-* Miyajima-cho^ 41-08' 34-~4r 34-17' 140-50' 138-44' 132-18' 9.1' ca. 16" ca. 16^^ 1625.0 = 134.4 1116.3 r 202.8 3 0 1028.9 = 309.4 793.5 ± 96.6 745.0 i 226.4 12 4 4 ■ Reference: Inagaki and Hamada (1985. p. 87). - Data for females and males combined: no significant difference between sexes. ' Studied in November 1983. ■•Studied in December 1983. ^ Studied in January 1984: progenitors of this group were translocated from Shodoshima (ca. 34''30'N. 134-18'E) to Miyajima-cho in 1962. al. (1992. p. 18). the pigmentation cline in M. fus- cata accords well with Gloger's rule (cf. Mayr. 1963. p. 324). Neonatal pelage color apparently follows a sim- ilar latitudinal cline (Hamada et al.. 1992. p. 7: cf. Kuroda. 1984. p. 14). In four captive neonates (age <4 months) that originated on Yakushima. the mean dorsal pelage paleness index \ alue was 27.9. and in five captive neonates that originated on Honshu, this value was 30.8. Pelage on the dorsal surface of the hands and feet also generally a\erages paler with increasing latitude and decreasing winter temperature on Yakushima. Kojima. Kyushu (Takasakiyama), and Honshu (Table 2); at all localities studied, the dor- sal surface of the feet averages paler than the dor- sal surface of the hands in both sexes. The dorsal surface of the hands is particularly dark (blackish) in Yakushima M. fuscata. On Kojima. Kyushu. Shikoku (BMfNHJ 1906.1.4.1-3). and Honshu, the dorsal surface of the hands averages paler than, or approximately as pale as. the dorsal sur- face of the trunk (hands/back paleness index ratio. 0.97-1.31); on Yakushima. the dorsal surface of the hands averages distinctly darker than the dor- sal surface of the trunk (hands/back paleness in- dex ratio. 0.83-0.92). Available infonnation per- mits no inference concerning whether variation of the hands/back paleness index ratio in the Yaku- shima population of M. fuscata is completely dis- junct from that in non- Yakushima populations of this species. Latitudinal variation of dorsal hair density in M. fitscata has been studied at three localities b\ Inagaki and Hamada (1985. p. 85). In accord with thermoregulatory expectations, adult hair density at these localities increases with increasing lati- tude and decreasing mean annual temperature (Ta- ble 4: cf. Hori et al.. 1977. p. 316). Hair density in infants (age <1 year), which has been studied at two of these localities, also apparently increases with increasing latitude and decreasing mean an- nual temperature; this suggests that the thermo- regulatory adaptation is genetic, not merely phys- iological. Albinism — Albino M. fitscata individuals have been captured in Tochigi and Yamagata Prefec- tures, northern Honshu (Temminck. 1842. p. 10; Kuroda. 1940. p. 271). Al Takasakiyama. Oita Prefecture. Kyushu, during the period 1950-1963. Itani et al. (1964. p. 30; cf. Itani & Mizuhara. 1957. p. 106) observed abnormal, sharply defined patches of white fur. usually on the hands and rarely on the feet, in 27 monkeys (10 females. 17 males) of a total population of 771 individuals; on the hands, the smallest of these white patches were restricted to the fingertips, whereas the larg- est extended as far as the distal forearms (cf. Tro- isi et al.. 1981. p. 6. who report patches of white fur on the fingers in one of 54 monkeys in a cap- tive group translocated from Takasakiyama to It- aly). Similar white patches also have been report- ed on the limbs of "more than six" monkeys at Hagachi. Shizuoka Prefecture. Honshu, and on the limbs of "a few" monkeys on Kojima. off the east coast of Kyushu (Nishida. 1966. p. 199; cf. Yamagiwa. 1979. p. 492). External Measurements and Proportions LiNE.AR Measurements of Body Size — The standard mammalogical measurement of body size is head and bodv length, which is the distance FIELDIANA: ZOOLOGY Table 5. Fig. 4).'-2 Prefectural variation in head and body length and related external measurements in Macaca fuscata (cf. Mean coordinates Adult females Adult males Head and Relative Body Head and Relative Body of sample body length tail length weight body length tail length weight Prefecture localities (mm) (T/HB X 100) (kg) (mm) (T/HB X 100) (kg) Honshu Niigata 38°18'N 532.0 15.4 10.1 561 16.0 12.2 139°36'E U) (/) {1) {1) {1) {1) Nagano 35°58'N 528.2 ± 25.82 15.2 ± 3.08 8.9 ± 1.51 580.1 ± 33.08 14.9 ± 4.14 12.6 ± 2.17 137°55'E 477-572 4.4-20.8 5.5-13.8 499-630 3.5-21.7 8.8-18.4 {42) {42) {42) {31) {30) {31) Shimane 34°52'N 488.5 ± 20.80 17.8 ± 3.25 7.9 ± 1.33^ 511.9 ± 27.93 16.2 ± 4.17 8.6 ± 1.17 132°38'E 464-517 13.0-22.8 6.0-9.2 472-545 8.8-19.9 7.6-10.8 {6) (6) (5) (7) (7) (7) Mie^ 34°20'N 136°30'E — — — 650 (i) 13.8 (i) — Shikoku Tokushima 33°50'N 515.8 ± 58.68 16.3 ± 2.21 8.2 — — — 134°18'E 472-601 {4) 13.3-18.6 {4) 7.8-8.6 (2) Kochi 33°24'N — — — 597 12.4 11.2 133°03'E U) (i) {1) Totals — — 522.8 ± 30.50 15.6 ± 3.09 8.8 ± 1.49 570.1 ± 42.30 15.1 ± 3.99 11.8 ± 2.49 464-601 4.4-22.8 5.5-13.8 472-650 3.5-21.7 7.6-18.4 {53) {53) {50) {41) {40) {40) ' Within each cell, figures indicate mean ± SD (where n > 2), extremes, and sample size (italicized, in parentheses). ^ This table excludes measurements of ""Head and body length" reported by Napier and Napier (1967, p. 406), who cite "Iwamoto et al. (1966) personal communication" as source; the published measurements actually are measurements of sitting height, not head and body length (M. Iwamoto, letter, 3 Feb. 1998). Also excluded are two measurements reported as head and body length by Aizawa and Hagiwara (2001, p. 5); these are measurements of anterior trunk length (M. Hagiwara, pers. comm., 3 Oct. 2002). ' Excludes one pregnant adult female, body weight = 9.4 kg. 4 Data from Kuroda (1940, p. 271). from the tip of the nose to the base of the tail (Imaizumi, 1970, p. 89). This measurement is available for 94 wild-collected adult specimens of M. fuscata — 53 females collected in three prefec- tures on Honshu and one prefecture on Shikoku and 41 males collected in four prefectures on Honshu and one prefecture on Shikoku (Table 5); nearly 80% of these specimens were collected in Nagano Prefecture, Honshu. Mean (±SD) head and body length is 522.8 ± 30.50 mm (extremes, 464-601 mm) in the available female sample and 570.1 ± 42.30 mm (extremes, 472-650 mm) in the available male sample. Although the unbal- anced geographic distribution of the available samples hinders inference concerning geographic variation of head and body length in M. fuscata, this measurement apparently averages less in both sexes in Shimane Prefecture, southwestern Hon- shu, than in the more northeasterly prefectures on Honshu for which data are available (Table 5; Fig. 4). Sitting height, the distance from the apex of the crown to the plane of the ischial callosities (Schultz, 1929, pp. 218, 233), is an anthropolog- ical measurement of body size that is similar but not identical to head and body length; this mea- surement is available for 168 adult females (mean = 537.4 mm) and 126 adult males (mean = 590.7 mm) that originated on five Japanese islands (Ta- ble 6). Inter-island comparison indicates that sit- ting height in M. fuscata is smallest on Yakushi- ma, as previously noted by Iwamoto (1971, p. 154); somewhat greater on Shodoshima; and greatest on Kyushu and Honshu (Fig. 5; Table 6); evidence provided by the small Shikoku sample is equivocal. On Honshu, no general pattern of latitudinal or longitudinal variation in sitting height is evident (Appendix 5); however, this FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 650 600 550 500 450 • • • • • • Adult males o • • • • • • • • • • • • 0 • 600 0 - - ^ <■ • 550 ••:. • Adult females o • • * •• • • • 500 « • • • ^ o ( , • • • • • Honshu o Shikoku 450 33 34 35 36 Latitude ("N) 37 38 39 Fig. 4. Latitudinal variation in head and body length in Honshu and Shikoku samples of Macaca fuscata adults (cf. Table 5). measurement apparently averages less on Boso Peninsula (Appendix 5: 35-36°N. 139-14rE) than elsewhere on Honshu (Boso Peninsula — fe- males, 525.7 mm, n = 6; males, 579.5 mm, n = 1 1; elsewhere on Honshu — females, 552.1 mm, n = 95; males, 604.6 mm, n = 58), and it also may be relatively small in males in southwestern Hon- shu (south of 35°N, west of 134°E; Appendix 5). The linear measurement of body size in M. fus- cata for which the largest samples are available is anterior trunk length, an anthropological measure- ment of the distance from the suprasternal notch to the cranial margin of the pubic symphysis (Schultz, 1929, pp. 232, 234); this measurement is available for 783 adult females (mean = 365.2 mm) and 350 adult males (mean = 392.5 mm) that originated on FIELDIANA: ZOOLOGY Table 6. Insular variation in sitting height (mm) in Macaco fuscata adi Lilts (cf. Fig. 5).' Adult females Adult males Mean latitude Island of localities (N) Mean ± SD N Mean ± SD N Honshu-' 35°37' 555.5 ± 29.21^ 75 602.5 ± 34.395 61 Honshu- 34°30' 536.3 ± NA* 26 586.1 ± NA" 8 Shodoshima^ 34°30' 518.8 ± 14.47 20 574.9 ± 22.58 21 Shikoku^ 33°34' 527.5** 2 609 1 Kyushu^ 33°10' 537.1 ± 19.08 11 621.2 ± 19.54 8 Yakushima^ 30°34' 510.1 ± NA* 34 568.1 ± NA* 27 Totals — 537.4 ± NA" 168 590.7 ± NA" 126 ' Reference: Iwamoto (1971, table opp. p. 156), except as otherwise indicated in footnote 3. ^ For details concerning these samples, see Appendix 5. -'' Data in this row were derived from specimens examined, not from Iwamoto (1971). '' Extremes: 487-625 mm. 5 Extremes: 504-682 mm. ^ NA = not available. ' Captive and/or provisioned populations. ** Extremes: 512-543. seven Japanese islands (Table 7). In accord with Bergmann's rule (cf. Mayr, 1963, p. 320), anterior trunk length generally increases with latitude from Yakushima and Kojima to Honshu (Fig. 6), as pre- viously noted by Hamada et al. (1996a, p. 106). As indicated above for sitting height, no general pattern of latitudinal or longitudinal variation in an- terior trunk length is evident in Honshu samples (Appendix 6). Also as indicated for sitting height, anterior trunk length apparently averages less on Boso Peninsula (Appendix 6, 35-36°N, 139- 14 TE) than elsewhere on Honshu (Boso Peninsu- la— females, 351.5 mm, n = 23; males, 394.7 mm, n = 16; elsewhere on Honshu — females, 374.7 mm, n = 338; males, 403.1 mm, n = 194), and anterior trunk length tends to be relatively small in a small sample of adult males collected in south- western Honshu (south of 35°N, west of 133°E; Appendix 6). Tail Length — Mean tail length in M. fuscata is 82.6 mm in 665 adult females and 91.1 mm in 312 adult males (Table 8). The tail in both sexes tends to be relatively long on Shodoshima and Yakushima (Table 8, Fig. 8), as previously noted by Hamada et al. (1996a, p. 105), and it tends to be relatively short on Kojima and probably on Shikoku. The apparent difference in tail length be- tween the Yakushima and Kojima samples is per- haps unanticipated because the distance between these two small islands is less than 140 km. Among females on Honshu, tail length may tend to decrease with latitude (Fig. 7; Appendix 7). The ratio of tail length to head and body length (T/HB) provides a useful index value for com- paring individuals or species of different body siz- es (cf. Fooden & Albrecht, 1999, p. 433). In avail- able samples collected on Honshu and Shikoku, the mean value of T/HB is similar in adults of both sexes — 15.6% in 53 females and 15.1% in 40 males (Table 5); this ratio averages greater in young immatures (T/HB = ca. 25%) (Fig. 8), which indicates that the postnatal growth rate of head and body length exceeds that of tail length. The ratio of tail length to anterior trunk length (T/ATL), which is available for samples that are much larger and geographically more broadly dis- persed than those for T/HB, provides a supple- mentary basis for appraisal of insular variation of tail length relative to body size (Table 8; Appen- dix 7). Judging from these samples, T/ATL tends to be relatively large in both sexes on Yakushima and Shodoshima and tends to be relatively small in both sexes on Shikoku. Although T/ATL in the total sample of females (22.7%) is close to T/ATL in the total sample of males (23.3%), this ratio in males exceeds that in females in seven of the eight available insular samples (Table 8). The tail is unusually short (<50 mm, T/HB <10%) in three adult females and three adult males collected in Nagano Prefecture, Honshu, and in one adult male collected in Shimane Pre- fecture, Honshu (Table 9; Fig. 8; cf. Table 8); three of these short-tailed specimens (one female, two males) make up the entire sample that was collected at Shimashima, Nagano Prefecture. Monkeys with similarly short tails (reportedly <30 mm) have been observed but not measured at two additional M. fuscata localities (Itani et al.. FOODEN AND AlMl: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 11 700 • Honshu D Kyushu • • m __ ..__ _ o Shodoshima A Yakushima D * " • 1 • •• • • • 1 o • • V 1 ■ 1' ^ ( 1 1 « • • •• • • 550 X • ; • • • ? — 500 o • • Ajdult males c 5 in • 600 • • • • s • 550 • • • • • i - I o 1 o ( 1 ( • • • 500 J i- •' • 1 ! \dii\t females 450 ! 1 30 32 33 34 35 36 37 38 39 Latitude (°N) Fig. 5. Latitudinal variation in sitting height in Macaca fuscata adults (cf. Table 6). Data points with standard- deviation error bars are mean values derived from Iwamoto (1971, table opp. p. 156); data points without error bars are individual values derived from specimens examined (except for one male data point with coordinates x = 34.47 and y = 595, which is an individual value from Iwamoto, 1971). 1964, p. 35): at Takasakiyama, Kyushu, seven ab- normally short-tailed adults or subadults (three fe- males, four males) were seen in a total population of 771, and on Kojima, off the eastern coast of Kyushu, one abnormally short-tailed adult male was seen in a total population of ca. 50 (cf. Ka- wai, 1963, p. 113). Hind Foot Length — Mean hind foot length in Macaca fuscata is 153.6 min in 657 adult females and 170.4 mm in 273 adult males (Table 10). Hind foot length generally tends to increase with lati- tude from Yakushima to Honshu (Fig. 9). How- ever, in female and male samples available from Shodoshima, hind foot length is notably less than expected for the latitude of this island. In Honshu samples, hind foot length is not closely correlated with latitude (Appendix 8). The ratio of hind foot length to anterior trunk length (HF/ATL), which has been reported for several M. fuscata samples by Hamada et al. 12 FIELDIANA: ZOOLOGY Table 7. Insular variation in anterior trunk length (mm) in Macaca fiiscata adults (cf. Fig. 6).' Adult females Adult males Mean latitude Island of localities (N) Mean ± SD N Mean ± SD N Honshu-"* 35°43' 376.6 ± 23.71" 88 398.2 ± 28.975 68 Honshu- 35°37' 373.3 ± NA* 273 404.6 ± NA" 142^ Shodoshima** 34°30' 355.0 ± 19.48 19 392.8 ± 21.93 20 Awajishima 34°14' 353.3 ± 8.03 9 387.4 ± 21.12 19 Kyushu** 33°10' 363.7 ± 18.03 278 392.4 ± 23.70 22 Shikoku 33°04' 347.3 ± NA^ 179 398 110 Kojima" 31°27' 346.2 ±20.13 55 366.0 ± 26.89 43 Yakushima^ 30°34' 338.7 ± 15.84 44 367.8 ± 22.83 35 Totals — 365.2 ± NA^ 783 392.5 ± NA" 350 ' Reference: Hamada et al. (1996a, pp. 101, 102), except as otherwise indicated in footnotes 3, 7, 9, and 10. ^ For details concerning these samples, see Appendix 6. -^ Data in this row were derived from specimens examined, not from Hamada et al. (1996a). 4 Extremes: 329-470 m. ■''Extremes: 315-450 mm. * NA = not available. ^ Includes two measurements (400 mm, 404 mm) that were originally reported as head and body length by Aizawa and Hagiwara (2001, pp. 5-6); see Table 5, footnote 2. ^ Captive and/or provisioned populations. ' Includes two specimens examined (anterior trunk length 337 and 360 mm). '° Specimen examined. " Semiprovisioned population (ca. 12 kg of wheat distributed weekly to ca. 70 monkeys). (1996a, pp. 104, 105), provides a useful index val- ue that facilitates comparison of hind foot pro- portions in individuals of varying body size. As noted above for hind foot length, HF/ATL is no- tably low in the Shodoshima female and male samples; this ratio is notably high in the Kojima female and male samples. Although mean HF/ ATL in the total sample of females (42.2%) is close to that in males (43.1%), this ratio in males exceeds that in females in all seven available pairs of insular samples (Table 10) (cf. above. Tail Length). Body Weight — Body weight in M. fuscata av- erages 8.4 kg in 832 adult females and 1 1.3 kg in 368 adult males (Table 11; cf. Hazama, 1964, p. 94; Hiraiwa, 1981, p. 314). As previously noted by Hamada et al. (1996a, p. 100), body weight in this species generally tends to increase with lati- tude (Fig. 10), in accord with Bergmann's rule; however, a small sample of two low-weight adult males (5.6 kg and 6.5 kg) collected on Kinkazan, an islet off northeastern Honshu, is a conspicuous exception to this generalization. Body weight also averages exceptionally small in 36 adult females (7.0 kg) and 25 adult males (9.5 kg) studied on Boso Peninsula, southeastern Honshu (Appendix 9, 35-36°N, 139-14rE). Seasonal changes in body weight have been re- ported in M. fuscata. At Shiga and Hakusan, which are relatively cool habitats, body weight tends to decrease by up to ca. 2 kg during the winter (Hamada et al., 1996b, p. 320; cf. Hiraiwa, 1981, p. 314). Conversely, on Kojima, a warm habitat, body weight tends to decrease by approx- imately 2 kg during the summer (Mori, 1979b, p. 387; Horii et al., 1982, p. 427). In a group of laboratory captives of unknown origin that was housed in an air-conditioned room and fed a stan- dardized diet, mean body weight decreased by ca. 1 .5 kg during the winter months (Matsubayashi & Enomoto, 1983, p. 523; cf. Hamada et al., 2003, p. 85), approximately as in free-ranging monkeys living in cool habitats (see above). Translocation of M. fuscata to cooler or warmer climates apparently modifies body weight in a manner predicted by Bergmann's rule (Paterson, 1996, p. 589). Twenty-seven years after translo- cation of a group from Mihara, Hiroshima Prefec- ture, to a cooler climate in Oregon (U.S.A.), fe- male and male growth curves for body weight in the descendant group had become significantly higher than those in the original group. Converse- ly, 20 years after translocation from Arashiyama, Kyoto Prefecture, to a warmer climate in Texas (U.S.A.), growth curves for body weight in the descendant group had become lower than those in the original group. Summary — Available data concerning sitting FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 13 475 450 425 400 375 350 325 ! .0. I ■ Awajishima • Honshu A Kojima p Kyushu o Shikoku * Shodoshima A Yakushima — 1 j •■ Adult males r 1 .J. tI r4 *• • i 1 [ r 1 P 'r 1 .11 J 1 1 i ' 1 j - — J i L i^ • • • 1 j ! j ■ » i i 1 ! ^ • ' ■ill 300 30 31 32 33 34 35 36 Latitude (°N) 37 38 39 40 41 42 Fig. 6. Latitudinal variation in anterior trunk length in Maccica fuscata adults (cf. Table 7). Data points with standard-deviation error bars are mean values derived from Hamada et al. (1996a, pp. 101-102); data points without error bars are individual values derived from specimens examined. height, anterior trunk length, and body weight in- dicate that body size in M. fuscata generally tends to increase with latitude from Yakushima to Hon- shu, in accord with Bergmann's rule. On Honshu, sitting height and anterior trunk length generally do not increase with latitude, but body weight does. Data concerning sitting height, anterior trunk length, and body weight all indicate that body size is unusually small on Boso Peninsula, southeastern Honshu. Body size probably also is unusually small on Kinkazan island, off north- eastern Honshu, but this inference is based exclu- sively on body weight data available for two adult males. 14 FIELDIANA: ZOOLOGY Table 8. Insular variation in tail length (mm) and tail length/anterior trunk length ratio (T/ATL) in Macaco fuscata adults (cf. Figs. 7-8).' Mean latitude of " localities _ (N) Adult females Adult males Tail length Mean ± SD N T/ATL Tail length Island Mean ± SD N T/ATL Honshu-'' 35°5r 81.7 ± 15.00 54 21.7% 88.3 ± 19.79 49 22.2% Honshu^ 35°37' 79.3 ± NA^ 213 21.2% 91.1 ± NA-» 125= 22.5% Shodoshima^ 34°30' 87.2 ± 9.14 19 24.6% 102.1 ± 9.24 21 26.0% Awajishima 34° 14' 77.3 ± 3.81 9 21.9% 88.0 ± 8.65 19 22.7% Shikoku 33°ir 73.5 ± NA4 17^ 20.8% 74 1« 18.6% Kyushu"" 33°10' 86.5 ± 11.16 260 23.8% 94.0 ± 14.76 21 24.0% Kojima'o 3r27' 73.3 ± 8.69 49 21.2% 80.4 ± 12.77 41 22.0% Yakushima" 30°34' 90.0 ± 8.25 44 26.6% 101.4 ± 10.61 35 27.6% Totals — 82.6 ± NA^ 665 22.7% 91.1 ± NA^ 312 23.3% ' Reference: Hamada et al. (1996a, pp. 101-102, 104-105), except as otherwise indicated in footnotes 2, 5, 7, and 8. - Specimens examined. '' For details concerning these samples, see Appendix 7. ■* NA = not available. ^ Includes one specimen reported by Aizawa and Hagiwara (2001, p. 5; cf. Table 5, footnote 2); cf. Kuroda (1940, p. 271). '' Captive and/or provisioned populations. '' Includes two specimens examined (cf. Table 7). •* Specimen examined (cf. Table 7). ** For tail length measurements in a Macaca fuscata group that was translocated from Kyushu (Takasakiyama) to Italy, see Masali and Scarsini (1981, p. 12). '" Semiprovisioned population (ca. 12 kg of wheat distributed weekly to ca. 70 monkeys). Tail length and T/ATL, which are exceptionally high on Yakushima and Shodoshima and excep- tionally low on Kojima, apparently are not closely correlated with latitude. Conversely, hind foot length, like body size, tends to increase with lat- itude in M. fuscata from Yakushima to Honshu, but it does not tend to increase with latitude on Honshu. Hind foot length and HF/ATL are nota- bly low on Shodoshima. Dermatoglyphics Dermatoglyphic patterns on the hands of M. fuscata were studied by Iwamoto and Suryobroto (1994, p. 78; cf. Iwamoto, 1964, p. 54; 1975a, p. 73; De Stefano et al., 1981b, p. 34; Iwamoto & Suryobroto, 1990, p. 433) in five samples that originated on Honshu, Kyushu (two samples), Ko- jima, and Yakushima. Although no absolute dif- ferences in palmar patterns or palmar ridge direc- tions were found among these samples, several frequency differences were discovered in these traits (Table 12); frequencies in the Shimokita Peninsula (Honshu) and Yakushima samples, re- spectively, at the northern and southern limits of distribution of M. fuscata, were the most deviant among the five samples studied. On the feet of M. fuscata, frequency differences in plantar derma- toglyphic patterns have been discovered in Taka- sakiyama (Kyushu) and Yakushima samples (Iwa- moto, 1967b, p. 156; cf. De Stefano et al., 1981a, p. 22). Intraspecific variation in palmar dermatoglyph- ic pattern frequencies within M. fuscata apparent- ly may exceed interspecific variation in pattern frequencies among M. fuscata, M. fascicularis, and M. nemestrina (Cauble & Mavalwala, 1973, p. 151; Morris & Kerr, 1974, p. 229). Congenital Malformation of Limbs In 1954 and 1955, two or three years after reg- ular provisioning of several free-ranging M. fus- cata groups had been initiated, primatologists in Japan began to notice the occurrence of monkeys with congenitally malformed appendages (Itani & Mizuhara, 1957, p. 106; Itani et al., 1964, p. 30; Furuya, 1966, p. 488; Shidei et al., 1981, p. 1). The severity of observed malformations varied from syndactyly, to deformed interphalangeal FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 15 130 120 110 100 90 80 70 60 50 40 I 30 £" 20 o = 110 n I- 100 90 80 70 60 50 40 30 20 ■r * i ■ Awajishima • Honshu A Kojima D Kyushu o Shikoi i 1 i : j II 1 -l-T '"J-. • 1. 1 1 i ir • ; A Yakushima 1 ! o ■ J '• i ' 1 1 ! ! i • i • 1 i ' • i j i 1 ; j • ! Adult males i « • ■ . lu d'- 1 ' , .1 > T T II 1) li • <• • 1 , < » ( 1 1 1 1 j [^ J i ■ 1 " !• !■•■•; * • » ■'■ 1 1 1 ■■ 1 ! i i ! • ! * • ! ; 1 Adult females • 30 34 36 Latitude (°N) 37 38 39 40 42 Fig. 7. Latitudinal variation in tail length in Macaca fuscata adults (cf. Table 8). Data points with standard-deviation error bars are mean values derived from Hamada et al. (1996a, pp. 101-102); data points without error bars are individual values derived from specimens examined or from literature records cited in Table 8. joints, to missing digits (usually middle digits) on hands and/or feet, to absence of hands or feet, to virtually complete absence of pelvic limbs; only one instance of Polydactyly has been reported (Homma, 1980, p. 26). The geographic distribution of M. fuscata groups with malformed members extends broadly from Shiga, north-central Honshu, to Yakushima, at the southern limit of the specific range (Itani et al., 1964, p. 30; Furuya, 1966, p. 489; Nishida, 1966, p. 199; Iwamoto, 1967b, p. 248; Tanaka & Nigi, 1967, pp. 93, 96; Iwamoto & Hirai, 1970, p. 395; Iwamoto et al., 1975, p. 78; Yoshihiro et al., 1979, p. 459; Homma, 1980, p. 21; Shidei et al., 1981, p. 17; Nakamichi et al., 1983, p. 53; Minezawa et al., 1990, p. 573). Of 65 groups sur- veyed for congenital limb malformations, 22 (33.8%) included malformed individuals (Table 16 FIELDIANA: ZOOLOGY Table 9. External measurements in seven Macaco fuscata adults with unusually short tails (cf. Figs. 7-8). Head and body Tail PRIKU length length Locality field no. Sex (mm) (mm) T/HB Nagano Prefecture !, Honshu Aoki 87 ? 529 46 8.7% Nishiharuchika 120 9 505 42 8.3% Shimashima 39 9 502 22 4.4% Kamimachi 41 6 600 21 3.5% Shimashima 38 6 537 45 8.4% Shimashima 40 S 557 38 6.8% Shimane Prefecture, Honshu Ushiroyama (H-9906) 6 545 48 8.8% pie autosomal dominant or as a simple autosomal recessive (Iwamoto, 1967b, p. 261; Iwamoto & Hirai, 1970, p. 396; Yoshihiro et al., 1979, p. 468; Shidei et al., 1981, p. 28; Minezawa et al., 1990, p. 573). Inbreeding, pesticide exposure, and con- tamination of provisioned food have all been sug- gested as possible etiological factors (Nishida, 1966, p. 199; Yoshihiro et al., 1979, p. 469; Shidei et al., 1981, p. 35; Minezawa, 1990, p. 575). Al- though elevated organochlorine pesticide residues have been detected in cadavers of malformed monkeys and in cadavers of mothers of mal- formed monkeys, this evidence is not yet conclu- sive as to the cause of malformations (Ito et al., 1988, pp. 109, 111, 112). 13). The occurrence of malformed monkeys in provisioned groups appears to be more common than in nonprovisioned groups. The incidence of congenital limb malforma- tions in newborn infants apparently varies locally and varies over time at a single locality. At nine localities where affected groups have been studied for 2-26 years, the mean neonatal incidence of malformations varies from ca. 2.0% to 17.3% (Ta- ble 14); at one of these localities (Awajishima), the annual incidence varied from 71.4% in 1971 to 0% in 1972 to ca. 40% in 1977 (Yoshihiro et al., 1979, p. 461). The incidence of malformations in female infants apparently is similar to that in male infants (Table 15); results of one study in- dicate that primiparous mothers are less likely to produce malformed infants than are multiparous mothers (Nakamichi et al., 1997, p. 229). During the 1980s, the incidence of malformations is said to have declined rapidly (Minezawa et al., 1990, p. 575). The cause or causes of the congenital limb mal- formations are unclear (Iwamoto, 1967b, p. 261; Morris, 1971, p. 338; Yoshihiro et al., 1979, p. 469; Shidei et al., 1981, p. 39; Ito et al., 1988, p. 1 12; Minezawa et al., 1990, p. 576; Nakamichi et al., 1997, p. 231). Some mothers with malforma- tions have produced infants with malformations, and some affected individuals are known to be siblings (Itani et al., 1964, p. 34; Iwamoto, 1967b, p. 248; Nakamichi et al., 1997, p. 232; cf. Hase- gawa & Iwasaki, 1984, p. 399), which may sug- gest a genetic influence. However, malformations are not associated with detectible karyological ab- normalities, and pedigree analysis tends to rule out inheritance of malformations either as a sim- Cranial Characters (Figs. 11-14) E^ect of Provisioning Some free-ranging groups of M. fuscata — such as those inhabiting Kojima, Shodoshima, Taka- goyama, and Takasakiyama — are artificially pro- visioned to varying degrees (see Gazetteer, Ap- pendix 4). To investigate the effect of artificial provisioning on skull size, greatest length of skull in a sample of provisioned adults collected at Ta- kagoyama, Chiba Prefecture, has been compared with that in a sample of nonprovisioned adults collected elsewhere in the same prefecture (Table 16). Mean greatest length of skull in the provi- sioned females (1 18.1 mm) and males (134.4 mm) significantly exceeds that in the nonprovisioned females (115.8 mm) and males (129.5 mm). For this reason, where provisioned samples are in- cluded in subsequent analyses, they are individu- ally identified. Data from skulls of monkeys raised in captivity are excluded from subsequent analyses. Sex and Age Variation In the available sample of nonprovisioned wild- collected adult specimens of M. fuscata, greatest length of the skull in males (133.2 ± 5.72 mm, n = 100) averages about 12% greater than in fe- males (119.1 ± 4.94 mm, n = 141) (Table 17); size differences between M. fuscata males and fe- males reportedly are already evident in the skulls of early juveniles (Mouri, 1994, p. 46). Although rostral-postrostral ratio (XlOO) in adult males FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 17 in CM o in o CM T- T- '\ •*• of ■ • • tv . , o '. • . • ■ • . •. • 1 1 '. •• . ■ ■ o Females, adult • Males, adult o Females, immature or ac • Males, immature or age o in CD o o CO o in in o ti o r^ in ^ ,^*^ .— E := E > C JC ~~- ■*^ ^ o O) :; u; -^ c 1 >x r- •a •~ o — ^ tl "O JJ o o c re _>1 •D ^ re -o o X ij o ,^- in > CO o o o in o o CO (lUiu) i{;6u3| ijei FIELDIANA: ZOOLOGY Table 10. Insular variation in hind foot length (mm) and hind foot length/anterior trunk length ratio (HF/ATL) in Macaca fuscata adults (cf. Fig. 9).' Mean Adult females Adult males latitude of localities _ (N) Hind foot Mean ± SD length N HF/ATL Hind foot length Island Mean ± SD N HF/ATL Honshu- 35°37' 159.0 ± NA^ 238 42.5% 177.3 ± NA-^ 135^ 44.0% Shodoshima^ 34°30' 139.0 ± 4.45 19 39.2% 159.6 ± 5.37 21 40.6% Awajishima 34° 14' 147.6 ± 5.01 8 41.8% 168.6 ± 4.98 18 43.5% Kyushu'^ 33°10' 155.1 ± 5.80 276 42.6% 170.3 ± 8.25 21 43.4% Shikoku 33°06' 150.8 ± NA^ 176 43.4% — 0 — Kojima' 31°27' 152.6 ± 6.01 55 44.1% 164.1 ± 8.21 43 44.8% Yakushima^ 30°34' 137.0 ± 7.03 44 40.5% 155.7 ± 6.48 35 42.3% Totals — 153.6 ± NA-^ 657 42.5% 170.4 ± NA^ 273 43.1% ' Reference: Hamada et al. (1996a, pp. 101-103), except as otherwise indicated in footnotes 4 and 6. ^ For details concerning these samples, see Appendix 8. ^ NA = not available. ■* Includes one specimen reported by Kuroda (1940, p. 271) and one specimen reported by Aizawa and Hagiwara (2001, p. 5; cf. Table 5, footnote 2); the specimen reported by Kuroda lacks ATL information and therefore is not included in the HF/ATL calculation. '^ Captive and/or provisioned populations. ^ Includes two BM(NH) specimens examined; these specimens lack ATL information and therefore are not included in the HF/ATL calculation. '' Semiprovisioned population (ca. 12 kg of wheat distributed weekly to ca. 70 monkeys). (57.8 ± 3.65. n = 100) averages about 13% great- er than in adult females (51.2 ± 3.03, n = 137), relative zygomatic breadth (XlOO) in males (7L7 ± 2.22, n = 98) is approximately the same as that in females (70.9 ± 2.16, n = 139). From infancy to adulthood, rostral-postrostral ratio in males in- creases 105%, whereas relative zygomatic breadth increases only 10%. Dental emergence norms in M. fuscata are pre- sented in Table 18. For further details concerning sex and age variation in skulls of this species, see Schweyer (1909, p. 12), Ikeda and Watanabe (1966, p. 272), Koike and Shimamura (1988, p. 77), and Mouri (1994, p. 46; 1996, p. 283). Geographic Variation Judging from available data, mean greatest length of skull in M. fuscata increases successive- ly from south to north on Yakushima, Kojima, and Kyushu, the three southernmost islands represent- ed by samples (Table 19; Fig. 15); this is in accord with Bergmann's rule (cf. Mayr, 1963, p. 320). Farther northward, however, mean skull length apparently does not increase from south to north on Shikoku, Shodoshima, Honshu, and Kinkazan; in fact, mean skull length apparently decreases on Kinkazan, an islet off the northeastern coast of Honshu. Intra-island variation of skull length in the relatively large sample of Honshu specimens is analyzed in Appendix 10; as previously noted by Kuroda (1989, p. 7), the most conspicuous fea- ture of this variation is the aberrantly small skull size in females and males collected on Boso Pen- insula, southeastern Honshu (Appendix 10, cell 35-36°N, 140-141°E; cf. Fig. 15). A somewhat related series of measurements of "head length" (greatest midsagittal length from glabella to occiput) in living M. fuscata from sev- en islands has been published by Hamada et al. (1996a, pp. 102-104) (Table 20); the head length measurement employed in this study, which is standard in anthropology, excludes the rostrum of the skull and therefore pertains only to the cra- nium. The study of Hamada et al. further differs from the present one in including some specimens from captive and/or translocated groups. Despite these methodological differences, the pattern of insular variation indicated by measurement of head length has many similarities to that indicated by measurement of greatest length of skull (cf. Tables 19, 20). Ontogenetic development of greatest length of skull in Yakushima M. fuscata is compared with that in Honshu M. fuscata in Figure 1 6. A mean size difference between specimens from these two islands is already apparent in skulls with newly FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 19 lyo 190 185 180 175 170 Adult males ■ Awajishima • Honshu A Kojima D Kyushu o Shikoku o Shodoshima A Yakushima - - J M. •• ( 1 ( ) n fr ■ ( ( ) - - - r"' II 1 ■'■ ( 1 ■tRe^ 1 1Rn ' T'' ^ k ■■ ■'■ i I i 155 — ■1- 1) 145 iJ 175 o •2 170 ■o c I 165 160 lies T Ad ult femj r ( 1 ( ■ • 1 1 • I 1 i< 1 i ( 1 ■ •• ^'^^ 1 I 1 J- ( ■- J i 145 140 135 130 125 [ C 6 0 II T ( 1 I i ■■ ■■ ■• ■■ I <' "" i ] ' i 30 31 32 33 34 35 36 Latitude (°N) 37 38 39 40 41 42 Fig. 9. Latitudinal variation in hind foot length in Macaca fuscata adults (cf. Table 10). Data points with standard- deviation error bars are mean values derived from Hamada et al. (1996a, pp. 101), data points without error bars are individual values derived from specimens examined or from literature records cited in Table 10. erupted third molars (age ca. 6 years); available evidence is inadequate to determine the age of onset of this size difference. Ikeda and Watanabe (1966, p. 274; cf. Kuroda, 1984, p. 15) have shown that orbital index (orbital height/orbital breadth) averages greater in M. fus- cata collected on Yakushima than in M. fuscata collected on other Japanese islands (cf. Table 21). The average insular difference in this index is pri- marily the result of lesser orbital breadth in Yak- ushima M. fuscata — not of greater orbital height (Fig. 17). Computed tomographic imaging of a sample of 46 M. fuscata skulls that originated at six locali- ties— extending from Yakushima to Shimokita Peninsula — has revealed that nasal cavity area in this species tends to increase with increasing lat- itude, whereas maxillary sinus volume tends to decrease with increasing latitude (Rae et al., 2003, p. 155). Enlargement of the nasal cavity at higher latitudes has been interpreted by Rae et al. as an adaptation that functions to augment the warming of inspired air in cooler climates; reduction of the maxillary sinuses at higher latitudes is interpreted 20 FIELDIANA: ZOOLOGY Table 11. Insular variation in body weight (kg) in Macaca fuscata adults (cf. Fig. 10).' Mean Adult females Adult males latitude of localities (N) Island Mean ± SD N Mean ± SD N Kinkazan- 38°17' _ 0 6.0^ 2 Honshu"* 35°54' 9.6 ± NA^ 276 12.9 ± NA5 143^ Honshu^" 35°38' 8.7 ± 1.56^ 125 11.4 ± 2.44» 81 Shodoshima^ 34°30' 8.9 ± 1.15 18 10.7 ± 2.73 20 Awajishima 34°14' 7.1 ± 1.26 10 10.8 ± 1.58 19 Kyushu'^ 33°10' 7.8 ± 1.10 281'« 10.8 ± 2.15" 22 Shikoku 33°05' 8.2 ± NA5 1712 11.2 1'^ Kojima'^ 31°27' 6.3 ± 0.70 55 8.0 ± 1.83 43 Yakushima* 30°34' 8.0 ± 1.60 44 10.2 ± 2.06 35 Yakushima- 30°20' 5.3 ± 0.95'5 616 9.5'^ 29 Totals — 8.4 ± NA' 832 11.3 ± NA^ 368 ' Reference: Hamada et al. (1996a, pp. 101, 102), except as otherwise noted in footnotes 2, 6, 10, 12, and 13. ^ Data in this row were derived from specimens examined, not from Hamada et al. (1996a). ^ Individual values: 5.6 kg, 6.5 kg. •* For details concerning these samples, see Appendix 9. ^ NA = not available. 6 Includes two specimens recorded by Aizawa and Hagiwara (2001, p. 5). ^ Extremes: 4.8-13.8 kg; includes one known pregnant female, weight = 9.4 kg. ^Extremes: 7.6-18.4 kg. ' Captive and/or provisioned individuals. '" Includes one specimen examined, 6.3 kg. "Cf. Nigi et al. (1980, p. 236). '- Includes two specimens examined (see Table 5). ''' Specimen examined (see Table 5). '* Semiprovisioned population (ca. 12 kg of wheat distributed weekly to ca. 70 monkeys). '^ Extremes: 4.0-6.5 kg. '* Includes four captives. '^ Individual values: 9.2 kg, 9.8 kg. as a passive result of crowding by the adjacent enlarged nasal cavity. Based on frequency of occurrence of selected nonmetric cranial characters, Kuroda (2002b, p. 125) has reported that skulls of the Boso Penin- sula, Shodoshima, and Yakushima populations of M. fuscata tend to differ from skulls of other stud- ied populations of this species; relevant frequen- cies are not specified in this report. In another publication, Kuroda (2002a, p. 115) has compared metric cranial characters of M. fuscata subfossils, dated Early Jomon to Late Jomon (ca. 10,000- 300 B.C.), with those of modern populations from various Japanese localities; discriminant functions indicate that measurements of most of the subfos- sils are generally similar to those of modern pop- ulations. Dental morphology and measurements in M. fuscata skulls originating on Yakushima are com- pared in detail with those in skulls originating on other unspecified Japanese islands by Saheki (1966, p. 412). In another dental study, Yoshika- wa and Deguchi (1992, p. 123) found that per- manent maxillary canines were double-rooted in 9 of 22 female M. fuscata specimens examined (40.9%) and in 0 of 22 male specimens (all single- rooted); collecting localities of the specimens were not reported. Dental enamel thickness in M. fuscata has been studied by Shimizu (2002, p. 613). Interspecific Comparisons M. fuscata has the largest skull and the most projecting rostrum of any of the four species in the fascicularis group (Table 22; cf. Ehara, 1974, p. 92). Development of a well-defined median sagittal crest is relatively uncommon in adult males of M. fuscata (23 of 158 specimens ex- amined; cf. Ikeda & Watanabe, 1966, p. 275), as in M. mulatta and M. cyclopis (Fooden, 2000, p. 44; Fooden & Wu, 2001, p. 9). FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 21 18 16 12 i 4 O 14 OQ 12 • Adult males • i* (, • T • T « T' T-" „{■ ^ 1l 1 T T i-J lb • o II <> w ■> • • ■ Awajishima • Honshu ♦ Kinkazan A Kojima D Kyushu o Shikoku o Shodoshima A Yakushima — A ■■ -1- • • < •• 4 • . B • ♦ j ♦ j I Adult f emales • 1 • 1 • • » -•I • tv * • < 1 ( 1 T i" ^t!' ft ^ 1 I ^Z . V iS- o ^ Jl II D i r ! • i i A • • 1 1 i 30 35 36 Latitude (°N) Fig. 10. Latitudinal variation in body weight in Macaca fuscata adults (cf. Table 11). Data points with standard- deviation error bars are mean values derived from Hamada et al. (1996a, pp. 101-102); data points without error bars are individual values derived from specimens examined or from literature records cited in Table 1 1 . Molecular Biology and Genetics DNA: Intraspecific Studies Mitochondrial DNA — The most important studies of individual and local variation in mtDNA in M. fuscata have been published by Hayasaka et al. (1986, p. 348; 1991, p. 401) and Kawamoto (1997, p. 32; 1998, p. 53; 1999, p. 302; 2002, p. 61; cf. Hayaishi & Kawamoto, 2002, p. 164). Hayasaka et al. sequenced the ma- jor noncoding region of mtDNA in a sample of 100 monkeys; Kawamoto analyzed mtDNA re- striction site patterns in a composite sample of 134 monkeys, sequenced 412 base pairs in the D- loop in a sample of 119 monkeys, and (with co- author Hayaishi) sequenced 203 base pairs in the D-loop in another sample of 38 monkeys. Local- ities sampled Kawamoto include those sampled by Hayasaka et al. 22 FIELDIANA: ZOOLOGY Table 12. Frequency (%) of dermatoglyphic patterns on both hands in five local samples of Macaca fuscata.^ Localities Takasaki- Shimokita, Kawaradake, yama, Dermatoglyphic Pattern Honshu Kyushu Kyushu Kojima Yakushima pattern type (N = 70) (N = 106) (N = 236) (N = 234) (N = 276) Thenar O 11.4 63.7 59.6 67.2 22.5 L 88.6** 36.3 40.0 32.8 77.5** W 0 0 0.4 0 0 First interdigital O 5.7 0 1.3 0 0 L 92.9 92.1 68.4 100.0 80.7 W 1.4 7.9 30.3** 0 19.32 Second interdigital L 21.4** 2.0 0.9 5.6 1.1 W 78.6 98.0 99.1 94.4 98.9 Third interdigital O 1.4 3.0 2.6 0 0.7 L 17.4 37.6^ 19.7 49.1** 13.5 W 81.2 59.4 77.7 50.9 85.8 Fourth interdigital L 1.4 0 1.3 4.7 8.0 W 98.6 100.0 98.7 95.3 92.0 Distal hypothenar o 71.4 61.0 69.9 75.6 100.0** L 28.6 37.0 23.3 21.4 0 W 0 2.0 6.8 3.0 0 Proximal hypothenar L 1.4 37.8 35.7 47.6 97.1** W 98.6** 62.2 64.3 52.4 2.9 Ending of ridge Prox 0 85.1 82.7 91.4 5.1 direction II Furc 0 5.0 6.9 0.9 1.8 Ulna 100.0** 9.9 10.3 7.7 92.7** X 0 0 0 0 0.4 Ending of ridge Prox 0 10.0 8.6 3.9 1.1 direction III Furc 0 3.0 3.0 1.7 0 Ulna 100.0 81.0 82.8 92.7 95.6 Other 0 6.0 5.6 1.7 3.3 ' Reference: Iwamoto and Suryobroto (1994, pp. 78, 82). Double asterisks (**) indicate values that are significantly different from others in the same row (P < 0.01), except as otherwise noted. - Not significantly different from Takasakiyama value (P > 0.05). ^ Not significantly different from Kojima value (P > 0.05). Table 13. Provisioning status and occurrence of congenital limb malformations in 65 Macaco fuscata groups.' Provisioning status of group Groups with malformed members Number % . Number of groups Free ranging Nonprovisioned Provisioned 5 14 Captive 23.8 40.0 21 35 Provisioned 3 33.3 9 Totals 22 33.8 65 ' Reference: Yoshihiro et al. (1979, p. 460); cf. Shidei et al. (1981. p. 16). Hayasaka et al.'s (1986, p. 348; 1991, p. 401) sample of 100 monkeys originated at 12 localities that span entire geographic range of the species from northernmost Honshu to Yakushima (Fig. 18). Sequencing of the major noncoding region revealed that eight mtDNA haplotypes (designat- ed types I-VIII) are present in these 100 samples (Table 23). Based on sequence similarities, three of the eight mtDNA haplotypes may be allocated to group II/IA^II, three other haplotypes may be allocated to group III/IVA^III, and the remaining two haplotypes (V and VI) are of uncertain rela- tionships. In group \y\fW\\, mtDNA haplotype II includes one copy of a repeatable 158-bp sequence, hap- lotype I includes two copies of the same se- quence, and haplotype VII includes three copies (Hayasaka et al., 1991, p. 406). Similarly, in group III/IV/VIII, mtDNA haplotypes III, IV, and VIII, respectively, include one, two and three cop- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 23 Table 14. Neonatal incidence of congenital limb malformations in nine Macaca fuscata groups.' Incidence of congenital limb malformations Observation Locality Island period Number % N Provisioned groups Shiga Honshu 1963-1977 19 12.4 153 Otoumi Honshu 1972-1974 2 5.4 37 Ohirayama Honshu- -1956-1966 3 -2.0 -150 Gagyusan Honshu 1955-1977 66 12.2 542 Hagachizaki Honshu 1972-1975 8 12.1 66 Miyajima-cho Miyajima^ 1961-1978 17 9.0 188 Awajishima Awajishima 1969-1995 132 17.3 762 Takasakiyama Kyushu 1965-1978 Captive group 126 3.7 3395 Takaosan Honshu"* 1970-1977 2 8.0 25 Totals — — 375 -7.1 -5318 ' References: Iwamoto (1967b, p. 249; Ohirayama group only); Yoshihiro et al. (1979, p. 460); Nakamichi et al. (1997, p. 227; Awajishima group only); cf. Shidei et al. (1981, p. 16). - Group studied after translocation from Yakushima to Ohirayama. ^ Group studied after translocation from Shodoshima to Miyajima. "* Group progenitors captured at Hasumi (Honshu) and Shodoshima. ies of a repeatable 167/168-bp sequence. Haplo- types V and VI do not include repeated sequenc- es. Haplotypes I and VII probably were derived from haplotype II by segmental duplication and triplication, respectively (Hayasaka et al., 1991, p. 411), and haplotypes IV and VIII probably were similarly derived from haplotype III. In a se- quenced 896-bp HindWl fragment, the number of pairwise nucleotide differences reported within group II/I/VII is nine, and the number of differ- ences reported between group II/IA^II and group III/IVA^III is 14-17 (Hayasaka et al., 1996, p. 1048). Of the 1 2 localities studied, nine are represent- ed by multiple specimens (2-32 specimens), and three are represented by one specimen each (Table 23). Mitochondrial DNA haplotypes are mono- Table 15. Sexual variation in incidence of congen- ital limb malformations in Macaca fuscata neonates.' Incidence of congenital limb malformations Sex Number % N Females Males Totals 71 95 166 A.V 6.02 5.3 1520 1585 3105 ' Reference: Yoshihiro et al. (1979, p. 462); cf. Shidei et al. (1981, p. 30); Nakamichi et al. (1997, p. 227). 2P = 0.10. morphic at each of six localities represented by multiple specimens, and they are polymorphic at the remaining three localities represented by mul- tiple specimens; because sample size averages smaller at monomorphic localities (6.5 specimens per locality; extremes, 2-15) than at polymorphic localities (19.3 specimens per locality; extremes, 8-32), it is possible that larger sample sizes would reveal a higher incidence of local mtDNA hap- lotype polymorphism. At one of the polymorphic localities (Nikko), seven specimens are haplotype II, and one is haplotype VI (dimorphism); these two haplotypes are not members of the same hap- lotype group. At the other two polymorphic lo- calities, most specimens included two distinguish- able mtDNA haplotypes (heteroplasmy)-haplo- types 1 and VII (both members of group II/I/VII) at Takahama and haplotypes IV and VIII (both members of group III/IV/VIII) on Yakushima. Judging from available data, the most common and widespread mtDNA haplotype in M. fuscata is haplotype II, a member of group II/I/VII. Hap- lotype II occurs at seven localities over a span of ca. 900 km, from Shimokita Peninsula in northern Honshu to Wakasa (Tottori Prefecture) in west- central Honshu and to Shodoshima, an islet in the Inland Sea off the southern coast of west-central Honshu. At Takahama, which is within the area of distribution of haplotype II, 12 of 18 specimens are heteroplasmic for haplotypes I and VII; as in- dicated above, haplotypes I and VII probably 24 FIELDIANA: ZOOLOGY were derived from haplotype II by segmental du- plication and triplication. Also within the area of distribution of haplotype II is Nikko, where seven of eight specimens are haplotype II and one is haplotype VI, and Izu, where the single available specimen is haplotype V; the relationship of hap- lotype VI and haplotype V to other haplotypes is uncertain. Limited data available from three southern lo- calities suggest that the III/IVA^III group of mtDNA haplotypes replaces the II/I/VII group west and south of Wakasa and Shodoshima. At Gagyusan (Okayama Prefecture), which is on Honshu and is only 60-90 km west of Wakasa and Shodoshima, all five specimens examined are mtDNA haplotype III. And on Yakushima, which is ca. 570 km southwest of Gagyusan, all 32 spec- imens examined are either haplotype IV or hap- lotype VIII or both (heteroplasmic); haplotypes IV and VIII probably were derived from haplo- type III by segmental duplication and triplication. However, at Takasakiyama, which is on Kyushu and is between Gagyusan and Yakushima, all three specimens examined are haplotype V, a mtDNA haplotype of uncertain relationships, oth- erwise known in M. fuscata only in the single specimen from Izu, which is on Honshu ca. 700 km east-northeast of Takasakiyama. In summary, the most important systematic im- plications of Hayasaka et al.'s (1991, p. 406) se- quencing study of the major noncoding region of mtDNA in 100 M. fuscata samples representing 12 Japanese localities are as follows: 1. Eight mtDNA sequence haplotypes (I-VIII) are known in M. fuscata. 2. Mitochondrial DNA haplotypes I and VII probably were derived from mtDNA haplotype II by segmental duplication and triplication, respec- tively (group II/I/VII). Similarly, mtDNA haplo- types IV and VIII probably were derived from mtDNA haplotype III by segmental duplication and triplication (group III/IV/VIII). The relation- ship of haplotype V and haplotype VI to other haplotypes is uncertain. 3. Mitochondrial DNA haplotype is monomor- phic at six of nine Japanese localities represented by multiple samples, it is dimorphic at one local- ity, and it is heteroplasmic at two localities. 4. Mitochondrial DNA haplotype II occurs at 7 of 12 sampled localities, and mtDNA haplotype V occurs at two localities. Each of six other mtDNA haplotypes occurs at only one locality. 5. The II/I/VII group of haplotypes is wide- spread in northern and central Honshu, and this group also occurs on nearby Shodoshima. 6. The III/IV/VIII group apparently replaces the II/I/VII group in southwestern Honshu and on Yakushima. 7. Mitochondrial DNA haplotype V is disjunct- ly distributed in south-central Honshu and Kyu- shu. 8. Mitochondrial DNA haplotype VI is known only in one of eight specimens obtained at Nikko, north-central Honshu. Kawamoto's (1997, p. 32; 1998, p. 53; 1999, p. 302) restriction-site studies yielded results that are largely compatible with results of his geographi- cally more comprehensive sequencing study (2002, p. 65). For this reason, only the sequencing study will be discussed here. Kawamoto's (2002, p. 57) sequencing study of 119 monkeys included samples collected at 106 localities spanning the entire geographic range of M. fuscata, from Shimokita Peninsula in the north to Yakushima in the south. Of 49 haplotypes de- tected in this study, 31 were restricted to one lo- cality, six occurred at two localities, four at three localities, three at four localities, one at five lo- calities, two at seven localities, one at 12 locali- ties, and one at 18 localities (Table 24, Fig. 19). Localities that share the same haplotype tend to be fairly tightly clustered geographically; how- ever, this does not apply to the 18 localities that share haplotype no. 1, which span ca. 400 km in northern Honshu, and also does not apply to the seven localities that share haplotype no. 20, which span ca. 400 km in central Honshu; unlike hap- lotype no. 20, haplotype no. 1 appears to be the only haplotype present in most of its broad area of distributions. Haplotypes are polymorphic at 10 of 1 1 localities represented by multiple samples (monomorphic at one locality). Based on sequence divergence values, with M. cyclopis taken as an out-group, the 49 sequence haplotypes that Kawamoto (2002, p. 65) detected in M. fuscata are clearly separable into two groups — group I, 17 haplotypes (nos. 7-11, 35- 43 [including 38 + 1, 41 - 1, 41 + 1]), mean pairwise difference = 8.81 ± 4.19 (SD), and group II, 32 haplotypes (nos. 1-6, 12-23, 26-34 [including 16 + 1, 17 + 1, 17 + 2, 20 + 1, 26 + 1]), mean pairwise difference = 5.98 ± 2.88. In groups I and II combined, with a total of 49 haplotypes, mean pairwise difference = 9.10 ± 4.22. Although the known geographic distribution of group I haplotypes is mainly in southern Japan FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 25 Fig. 11. Skull of adult female Macaca fuscata fuscata- Honshu. Scale in cm. (Photographs by Minoru Kinoshita.) -PRIKU 6492, Kuradama. Kimitsu. Chiba Prefecture, (Fig. 19), extending from Yakushima to Awajish- ima and southwestern Honshu (Gagyusan, Oka- yama Prefecture), group I haplotypes also occur disjunctly in a relatively small area in east-central Honshu (eight specimens in Yamanashi, Tokyo, and Saitama Prefectures; one specimen in Niigata Prefecture). Group II haplotypes are widely and apparently continuously distributed from Sho- doshima and west-central Honshu (Wakasa, Tot- tori Prefecture) to northernmost Honshu (includ- ing Kinkazan islet). Although group II localities virtually surround the disjunct group I localities in east-central Honshu, group I haplotypes and group II haplotypes are not known to occur at the same locality. Because group I haplotypes are more differen- tiated than group II haplotypes, Kawamoto (2002, p. 68) has inferred that the group I population originated earlier than the group II population. Kawamoto (2002, p. 69) has further suggested 26 FIELDIANA: ZOOLOGY Fig. 12. Skull of adult female Macaca fuscata yakui — PRIKU 6638, Hanyama, Kamiyaku-cho, Kagoshima Pre- fecture, Yakushima. Scale in cm. (Photographs by Minoru Kinoshita.) that the divergence of group I and group II pop- ulations and the broad dispersal of haplotype no. 1 (group II) in northern Honshu are consequences of changes in climate and vegetation that were induced by Pleistocene glacial advance and re- treat. In a supplementary publication, Hayaishi and Kawamoto (2002, p. 164) reported on D-loop se- quence haplotypes (203 bp) in 38 M. fuscata sam- ples that were collected at 21 localities widely dis- persed on Yakushima. Analysis of these samples revealed four haplotypes (Y1-Y4) that differed at only three of 203 sites. Thirty-three samples col- lected at 18 localities were identified as Yl (cf. Domingo-Roura et al., 2004, p. 37); two samples collected at two close-lying localities were iden- tified as Y2; two samples collected at one locality were identified as Y3; and one sample — collected at a locality that also yielded a sample identified as Yl — was identified as Y4. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 27 Fig. 13. Skull of adult male Macaca fuscata fiiscata- Scale in cm. (Photographs by Minora Kinoshita.) -PRIKU 5414. Naka, Natashou, Fukui Prefecture, Honshu. Following is a brief summary of the results of Kawamoto's (2002, p. 61) sequencing study of 119 A/, fiiscata samples representing 106 locali- ties from Shimokita Peninsula to Yakushinia: 1. Of 49 haplotypes detected. 31 were restrict- ed to a single locality. 2. Haplotypes shared by multiple localities tend to cluster in local areas. However, each of two exceptional haplotypes (nos. 1 and 20) is dis- tributed over a span of ca. 400 km; haplotype no. 1 apparently is the only haplotyjje present in most of its broad area of distribution. 28 FIELDIANA: ZOOLOGY Fig. 14. Skull of adult male Macaca Juscata yakiii — PRIKU 1571, Hanyama, Kamiyaku-cho, Kagoshima Prefecture, Yakushima. Scale in cm. (Photographs by Minoru Kinoshita.) 3. Haplotypes are polymorphic at 10 of 1 1 lo- calities represented by multiple specimens. 4. Sequence divergence separates the 49 de- tected haplotypes into two groups — group 1 (17 haplotypes) and group II (32 haplotypes). 5. Group I haplotypes are distributed from Yakushima to southwestern Honshu and disjunct- ly in east-central Honshu; group II haplotypes are continuously distributed from west-central Hon- shu to northernmost Honshu. Group I haplotypes and group II haplotypes are not known to occur together at the same locality. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 29 Table 16. Greatest length of skull (mm) in sample of provisioned Macaca fuscata population collected at Tak- agoyama,' Chiba Prefecture, Honshu, compared with that in sample of nonprovisioned populations collected elsewhere in Chiba Prefecture. Sample statistics Provisioned Nonprovisioned Mean ± SD Extremes N Mean ± SD Extremes N 118.1 ± 2.85 113.7-125.5 20 134.4 ± 3.26 128.6-138.1 Adult females 115.8 ± 3.35 109.2-120.9 25 Adult males 129.5 ± 4.22 121.3-139.0 21 2.44 2.95 <0.022 <0.012 ' Cf. Koike and Shimamura (1988, p. 75). - Two-tailed test. -* Excludes one Takagoyama specimen, collected ca. 1961, for which provisioning history is unknown (JMC 2588, greatest length of skull = 134.1). 6. Greater genetic differentiation of group I haplotypes suggests that the group I population may be older than the group II population. 7. Divergence of group I and group II popu- lations and broad dispersal of haplotype no. 1 (group II) may be related to Pleistocene glacial advance and retreat. The most salient point of agreement in the re- sults published by Hayasaka et al. (1986, p. 348; 1991, p. 401) and Kawamoto (1997, p. 32; 1998, p. 53; 1999, p. 302; 2002, p. 61) is that, based on sequence data, mtDNA haplotypes in M. fuscata are broadly subdivisible into two geographic groups — a northeastern group, distributed in northern and central Honshu, and a mainly south- western group, distributed primarily in western Table 17. Cranial measurements' and proportions in age/sex classes of nonprovisioned wild-collected Macaca fuscata.- Greatest length Relative zygomatic Postrostral Rostral-postrostral of skull breadth length ratio Age/sex class"* (mm) (ZB/GL X 100) (mm) (R/PR X 100) Infants 90.4 ± 5.14 65.1 ± 1.98 75.9 ± 3.82 28.2 ± 2.70 77.5-104.4 60.1-69.1 66.5-86.2 22.3-33.2 {51) {51) (52) (57) Juveniles 106.8 ± 9.57 68.7 ± 2.19 82.9 ± 4.62 40.6 ± 6.23 86.9-132.8 63.3-76.4 72.1-98.1 28.7-58.3 {283) {280) {284) {282) Subadult females 113.5 ± 3.99 69.9 ± 1.79 83.9 ± 2.96 A1.6 ± 1.83 107.2-121.4 67.1-73.9 78.1-88.2 44.0-52.4 {27) (26) {27) {27) Subadult males 128.9 ± 6.03 69.7 ± 2.05 91.6 ± 3.85 54.3 ± 2.71 113.8-143.1 66.2-73.8 82.9-100.0 49.0-61.1 {40) {39) {39) {39) Adult females^ 119.1 ± 4.94 70.9 ± 2.16 86.5 ±3.18 51.2 ± 3.03 109.2-133.7 64.3-77.9 79.1-94.8 43.5-57.5 U41) {139) {138) {137) Adult male.s^ 133.2 ± 5.72 71.7 ± 2.22 92.0 ± 3.05 57.8 ± 3.65 120.6-148.3 65.5-77.3 83.0-98.8 48.3-66.2 {100) {98) {105) {100) ' For definition of measurements, see Fooden (1969, p. 40). - Mean ± SD, extremes, and sample size (italicized figures in parentheses). ' 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. "Cf. Fig. 15. 30 FIELDIANA: ZOOLOGY Table 18. Dental emergence-'' chronology in Macaca fuscata. (References: Iwamoto et al., 1984, p. 277; 1987, p. 24; cf. Nass, 1977, p. 309; Passarello, 1980, p. 54). Tooth Sex Dental emergence age (yr) Dispersion (ca. 10th percentile- Mean ca. 90th percentile) h m,, m' c,, c' nij m2 M, M' I2 P M2 M2 C, P3. P' C P,, P' P4, P4 c, c M3, M-^ F + M F + M F + M F + M F + M F + M F + M F + M F + M F + M F + M F + M F + M F + M F F M F + M M M F + M Deciduous teeth* 0.019 0.000-0.038 0.058 0.019-0.096 0.077 0.038-0.12 0.21 (0.15-0.27)^ 0.21 (0.15-0.29)^ 0.46 (0.38-0.56)^ 0.50 (0.42-0.60)7 Permanent teeth 8 1.50 1.00-2.00 1.75 1.25-2.25 2.50 2.00-3.00 2.75 2.25-3.25 3.25 2.75-3.75 3.50 3.00-4.00 3.75 3.25-4.25 3.75 3.25-4.50 4.00 3.50-4.50 4.00 3.50-4.75 4.25 3.75-4.75 4.25 3.75-4.75 4.50 4.00-5.25 4.75 4.25-5.50 5.75 5.00-6.75 39 + 37 39 + 37 39 + 37 39 + 37 39 + 37 39 + 37 39 + 37 101 + 106 101 + 106 101 + 106 101 + 106 101 + 106 101 + 106 101 + 106 101 101 101 106 101 + 106 106 106 101 + 106 ' "* Tooth position indicators. ^ Defined as initial penetration of gingiva by crown of tooth. * Samples include 31 captives (16 females, 15 males) and 45 free-ranging monkeys (23 females, 22 males). '' Dispersion values reported for these teeth are extreme emergence ages. ^ Samples include 39 captives (17 females, 22 males) and 168 free-ranging monkeys (84 females, 84 males). Samples of free-ranging populations at Shiga (31 females, 24 males) and Kojima (46 females, 35 males) were excluded from calculation of the tabulated statistics by Iwamoto et al. (1987, pp. 19, 24); dental emergence ages in these two populations apparently are ca. 25-45% greater than in other populations of M. fuscata. Table 19. Insular variation in greatest length of skull (mm) in wild-collected Macaca fuscata (cf. Fig. 15). Mean latitude of localities . (N) Adult females Adult males Island Mean ± SD Extremes N Mean ± SD Extremes N Kinkazan 38°17' 118.1 ± 1.74 114.8-119.8 7 133.9 ± 2.18 131.4-137.1 7 Honshu' 35°35' 121.0 ± 4.64 109.2-133.7 152 134.7 ± 5.25 121.3-147.7 74 Shodoshima^ 34°31' — — 0 139.8 ± 3.23 134.4-146.3 9 Shikoku 34°00' 122.6 119.7-125.4 2 — — 0 Kyushu^ 32°53' 121.3 ± 1.20 119.6-122.4 4 142.5 ± 4.92 134.9-148.3 5 Kojima"* 31°27' 119.1 ± 4.71 109.1-129.2 19 135.2 ± 5.88 122.6-148.3 13 Yakushima 30°20' 113.4 ± 2.44 109.6-118.5 20 128.9 ± 5.23 116.2-137.4 29 Totals — 120.0 ± 4.90 109.1-133.7 204 134.1 ± 5.99 116.2-148.3 137 ' See Appendix 10. ' Provisioned population. ^ Includes three females and four males provisioned at Takasakiyama. '' Semiprovisioned population (ca. 12 kg of wheat grain distributed weekly to ca. 70 monkeys). FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 31 150 145 140 135 130 125 E E. 3 120 ® 135 130 125 120 115 110 105 — ▲ D • ! Adult males B I-**- * i A AA 4 A A A A A ▲ A * • • • • ; 1 1 1 i 1 A '< ♦ t 1 • A j 1 ! • 1% i i i i i A • • i i ! : i i 1 1 ^ i 1 ! i : i 1 • • Adult females I " i 1 ^ i • i B 1 : 9 • Honshu A A A ▲ A ' 1 1 ♦ ■ ♦ Kinkazan 1 A Kojima A ▲ : • D Kyushu ' o Shikoku ▲ ■,9 : ! I i A Yakushima 30 31 32 33 34 38 39 40 35 36 37 Latitude (°N) Fig. 15. Latitudinal variation in greatest length of skull in Macaca fuscata adults (cf. Table 19). 41 42 Honshu. Shikoku. Kyushu, and Yakushima (Figs. 18, 19). The zone of transition between these two groups apparently is in west-central Honshu, be- tween Wakasa (35°20'N. 134''24'E) and Gagyusan (34°47'N. 133°37'E). Haplotypes on the islet Sho- doshima (ca. 34°30'N, 134°15'E) apparently be- long to the northeastern group, and haplotypes on the islet Awajishima (ca. 34°15'N. 134°50'E) ap- parently belong to the southwestern group. In a recent study, Marmi et al. (2004, p. 677) sequenced a 392-bp fragment of the mtDNA con- trol region in samples obtained from 50 M. fiis- cata specimens that originated at six localities in central and southern Japan (Hakusan. n = 8; Tak- ahama. 9; Awajishima, 8: Takasakiyama, 6; Ko- jima. 8; Yakushima, 11); these sequences were compared with corresponding sequences — ob- tained from the GenBank/EMBL database — of 89 eastern M. mulatta specimens (82 from six local- ities in China; seven from one locality in Viet- 32 FIELDIANA: ZOOLOGY Table 20. Insular variation in head length (greatest midsagittal length from glabella to occiput, mm) in living Macaca fuscata (Hamada et al., 1996a, pp. 102-104). Adult females Adult males Island Mean ± SD' N Mean ± SD' N Honshu- 96.6 ± 4.01 238 103.8 ±4.11 130 Shodoshima 89.6 ± 2.22 19 94.6 ±3.17 21 Awajishima 96.9 ± 4.60 9 103.4 ± 3.58 19 Shikoku 92.3 ± 3.37 15 — 0 Kyushu 94.3 ± 3.34 277 99.4 ± 4.09 21 Kojima 89.2 ± 2.71 55 92.3 ± 5.04 43 Yakushima^ 89.2 ± 3.12 44 96.3 ± 4.37 35 Means 94.2 ± 3.35 657 99.9 ±4.17 269 ' Extreme values not published. 2 Includes captive and/or translocated groups. nam). Mitochondrial DNA haplotypes are mono- morphic at each of four of the M. fuscata locaH- ties and polymorphic at the remaining two (Tak- ahama; Kojima). The results of neighbor-joining and maximum-likelihood analysis in this study al- locate five of the six M. fuscata localities to two groups (Hakusan/Takahama and Awajishima/Tak- asakiyama/Kojima), which coincides with a major result of the studies of Hayasaka et al. (1986, p. 348; 1991, p. 401) and Kawamoto (1997, p. 32; 1998, p. 53; 1999, p. 302; 2002, p. 61) (see above); evidence in Marmi et al.'s study concern- ing the group relationship of the sixth locality (Yakushima) is equivocal. Compared with M. mu- latta haplotypes, the number of nucleotide differ- ences averages less in Yakushima haplotypes (28.3 ± 2.39; extremes, 24.8-32.0) than in hap- lotypes at o\hev M. fuscata localities (41.4 ± 4.12; 34.7-51.6). In other research projects, mtDNA variation has been employed in studies of hybridization, sub- fossils, and male dispersal. Kawamoto et al. (1999, p. 57; 2001, p. 17) used restriction-site analysis and sequencing of the mtDNA D-loop to compare M. fuscata, M. cyclopis, and putative hy- brids in Wakayama Prefecture. Mouri et al. (2000, p. 87) and Agatsuma and Iwagami (2002, p. 75) compared the D-loop sequence in several M. fus- cata subfossils with that in modem M. fuscata; among their findings is the discovery that, ca. 400 years ago, M. fuscata had been transported by hu- mans from Yakushima to Okinawa, where the species does not naturally occur. Yoshimi and Ta- kasaki (2003, p. 71) sequenced the D-loop in 16 monkeys in Okayama Prefecture and concluded that two local males had dispersed >100 km from their natal troop. MiNISATELLITE DNA AND MiCROSATELLITE DNA — Analysis of minisatellite DNA and micro- satellite DNA in blood samples has been used successfully to determine paternity in captive groups of M. fuscata (Weiss et al., 1988, p. 75; Inoue et al., 1990, p. 564; 1991, p. 204; Soltis et al., 1997a, p. 739). Analysis of microsatellite DNA is feasible in smaller and/or more degraded samples, including samples of semen, urine, and feces, and therefore is suitable for noninvasive de- termination of parentage in natural groups (Inoue & Takenaka, 1993, p. 41; Domingo-Roura et al., 1997, p. 358; 2004, p. 37; Hayakawa & Takenaka, 1999, p. 301). Local variation in microsatellite al- lele frequencies has been reported for two M. fus- cata localities (Wakasa, n = 28; Arashiyama, n = 34) by Inoue & Takenaka (1993, p. 41). DNA: Interspecific Studies Mitochondrial DNA — Within the fascicularis group, mtDNA in M. fuscata apparently is much more similar to that in M. mulatta and M. cyclopis than to that in M. fasciculahs (Table 25; cf. Mouri et al., 2000, p. 92; Domingo-Roura et al., 2004, p. 37). Within M. mulatta, mtDNA varies geo- graphically (Melnick et al., 1993, p. 289; Morales & Melnick, 1998, p. 16; Tosi et al., 2002, p. 168); mtDNA in M. fuscata is more similar to that in eastern M. mulatta (China, Burma) than to that in western M. mulatta (India, Pakistan). Nuclear DNA: Noncoding Intron — Deinard and Smith (2001, p. 47) sequenced a fragment of intron 4 (—410 bp) and the entire length of intron 5 (—545 bp) of the natural resistance-associated macrophage protein 1 (NRAMPl) gene in 59 FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 33 150 140 130 120 - 110 100 90 « 80 o 140 130 Males 8 • t 120 110 100 90 80 Females • • • 4 i! • Honshu A Yakushima 10 12 Estimated age (yr) 14 16 20 Fig. 16. Greatest length of skull vs. estimated age in Honshu and Yakushima specimens of Macaca fuscata. Age estimates to age ca. 6 years are based on dental emergence chronology (cf. Table 18): age estimates beyond age 6 years are based on degree of tooth wear (relative to tooth wear in known-age Kojima specimens). 34 FIELDIANA: ZOOLOGY Table 21. Insular variation in orbital measurements and orbital height/breadth index in adult wild-collected Macaca fuscata (cf. Fig. 17). Orbital breadth (mm) Orbital height (mm) Orbital index (H/Br) Island Mean ± SD Extremes Mean ± SD Extremes Mean ± SD Extremes N Adult females Honshu 24.4 ± 1.02 21.9-27.2 24.1 ± 1.70 20.5-28.0 0.99 ± 0.056 0.89-1.14 88 Shodoshima — — — — — — 0 Kyushu 23.5 — 22.4 — 0.96 — 1' Kojima 24.3 ± 0.79 22.5-25.9 24.8 ± 1.09 23.1-26.4 1.02 ± 0.039 0.95-1.10 192 Yakushima 22.1 ± 0.81 20.4-23.3 23.9 ± 1.51 19.1-25.8 1.08 ± 0.065 0.94-1.18 20 Adult males Honshu 26.1 ± 1.06 23.9-28.2 24.6 ± 1.43 21.7-28.2 0.94 ± 0.054 0.80-1.06 60 Shodoshima 25.0 ± 0.62 24.5-26.2 23.8 ± 0.61 23.2-25.1 0.95 ± 0.031 0.90-1.00 11' Kyushu 28.2 ± 0.78 27.2-29.2 25.6 ± 0.57 25.0-26.5 0.91 ± 0.042 0.86-0.97 53 Kojima 26.1 ± 1.09 24.2-27.8 25.7 ± 1.30 22.2-27.5 0.98 ± 0.036 0.91-1.03 142 Yakushima 23.6 ± 0.93 21.9-25.3 25.0 ± 1.68 22.6-29.0 1.06 ± 0.066 0.96-1.20 25 ' Provisioned population. ^ Semiprovisioned population (ca. 12 kg of wheat grain distributed weekly to ca. 70 monkeys). ^ Includes four provisioned specimens. samples of 1 1 species of macaques, including two samples of M. fuscata captives of unknown geo- graphic origin. Both M. fuscata samples share a single NRAMPl haplotype, which clusters closely with the haplotypes of M. mulatta, M. cyclopis, and M. fascicularis samples; at the center of this fascicularis-group cluster is a Chinese M. mulatta haplotype, from which the M. fuscata haplotype differs by one mutation. Nuclear DNA: Y-Chromosome — Tosi et al. (2000, p. 138; 2002, p. 168) sequenced 3.1 kb of Y-chromosome DNA (SRY and TSPY loci) in three M. fuscata captives of unknown geographic origin and compared these sequences with those in other species of macaques and other catar- rhines. Y-chromosome DNA in M. fuscata was found to be generally more similar to that in M. mulatta and M. cyclopis than to that in M. fasci- cularis. However, this generalization does not ap- ply to Indochinese populations of M. fascicularis, which are postulated to have received M. mulatta Y-chromosome DNA as a result of hybridization between dispersing M. mulatta males and philo- patric M. fascicularis females. Nuclear DNA: Random Amplified Polymor- phism— Vernesi et al. (2000, p. 186) have studied Random Amplified Polymorphic DNA (RAPD) in samples from nine M. fuscata specimens of un- known geographic origin and compared these samples with those from specimens of nine other macaque species. Within each of the 10 species studied, including M. fuscata, the RAPD pattern was identical in all samples. Interspecific cluster analysis of RAPD patterns suggests that M. fus- cata is the sister group of a hitherto unrecognized species group that includes M. nemestrina, M. fas- cicularis, M. tonkeana, M. arctoides, and M. mu- latta. However, Vernesi et al. (2000, p. 191) con- clude their report by cautioning that the uncertain genetic significance of RAPD variation dictates that "[p]rudence should guide inferences about nucleotide divergence, population structure and phylogeny based on RAPD markers." Mitochondrial DNA-Nuclear DNA: Multi- ple Loci— Tosi et al. (2003, p. 1421) have se- quenced one mitochondrial locus and three nucle- ar loci in three specimens of M. fuscata (precise origins unspecified) and 60 specimens of 18 other macaque species (Table 26). For three of the four loci (one Y-chromosomal, one mitochondrial, one autosomal), the sister group o^ M. fuscata was de- termined to consist variously of one or more fas- cicularis-group macaque species; for the fourth locus (autosomal), the sister group of M. fuscata was determined to consist of four sinica-group species. Tosi et al. (2003, p. 1425) suggest that the variation in sister groups that is indicated by these four loci probably is a result either of in- complete lineage sorting or of hybridization; for further discussion of incongruence in molecular phylogenies that are based on different genes, see Rokas et al. (2003, p. 798). Based on Y-chromo- some and mtDNA haplotype data, the divergence date of M. fuscata, M. mulatta, and M. cyclopis is estimated to be 1.0-1.2 million years ago (Tosi et al., 2003, p. 1431). FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 35 \ — D x: w c O X • A Kojima □ Kyushu o Shodoshima A Yakushima • • • ' 4 < • < < < << • I males < 1 o^ < < < Adult !] ^ o CO 00 CNJ CD CN CSJ 1 t # * 4 • •• • smales # ^ 'A 1. %• • i I* • Adult f< < ^ < < < < < < • < o o CO CO CM CD CN CNJ CNJ O CNI s ^ o ■o 1) eg ra o o CO E 4-* SI c k_ O c CO C3 CO ^ CM o Tt" CSJ i (uiuj) ^MBjaq |e)!qjo 36 FIELDIANA: ZOOLOGY Table 22. Greatest length of skull and rostral-postrostral ratio in fascicularis-gmup species. (References: Table 17; Fooden, 1995, p. 38; 2000, p. 44; Fooden & Wu, 2001, p. 13). Species Adult females Adult males Mean ± SD Extremes N Mean ± SD Extremes Greatest length of skull (mm) M. fascicularis 100.4 + 5.63 84.0-119.2 439 118.7 M. mulatta 107.8 ± 7.07 92.9-126.5 120 121.8 M. cvclopis 110.4 ±2.40 107.7-114.5 12 125.2 M.fiiscata 119.1 ±4.94 109.2-133.7 141 133.2 Rostral-postrostral ratio (%) M. fascicularis 47.6 ± 4.21 36.9-61.0 241 56.7 M. mulatta 44.1 ± 3.68 37.6-59.4 111 50.5 M. cyclopis 48.2 ± 3.04 44.7-51.1 4 55.2 M.fiiscata 51.2 ± 3.03 43.5-57.5 137 57.8 7.29 97.4-140.1 454 8.28 107.1-143.1 80 5.20 114.6-133.4 15 5.72 120.6-148.3 100 3.94 41.9-66.3 316 3.04 41.3-58.2 69 2.10 52.6-58.2 8 3.65 48.3-66.2 100 Blood Proteins The most comprehensive survey of blood-pro- tein variation in M. fuscata has been pubHshed by Nozawa et al. (1991, p. 415; 1996, p. 22), who studied 32 genetic loci in 3409 blood specimens that originated at 38 localities, spanning the entire specific range from Shimokita Peninsula to Yak- ushima (Fig. 20); in addition, these authors com- pared allele frequencies in M. fuscata with cor- responding frequencies in a sample of southern Chinese M. mulatta (n = 150), selected as a close phylogenetic relative of M. fuscata. A more lim- ited review of blood-protein variation in M. fus- cata and M. mulatta was published by Fooden and Lanyon (1989, p. 215). Although blood-protein variability apparently is less in M. fuscata than in many other macaques (Nozawa et al., 1996, p. 7), intraspecific analysis of Nei's standard genetic distance (D) has re- vealed a clear pattern of geographic differentia- tion in the Japanese species (Fig. 20). Based on this analysis, Nozawa et al. (1991, p. 425; 1996, p. 27) allocated the 38 sample localities of M. fuscata to seven groups; within each of these groups, interlocality D is less than 0.005. The geographic distribution of group 1 , which comprises 28 of the 38 localities, extends broadly from north-central Honshu to southernmost Kyu- shu, including the neighboring large island Shi- koku and small islands Awajishima and Kojima. The remaining six groups, to which 10 localities are allocated, are geographically peripheral to group 1 and are progressively more genetically distant from it. Group 2 comprises only one locality (Hakusan) in central Honshu, slightly marginal to group 1 localities (Fig. 20); between group 1 and group 2, D ~ 0.01 1. Group 3 comprises three localities on Izu Peninsula, south-central Honshu; between group 3 and the two preceding groups, D ~ 0.013. Group 4 comprises two localities, 220 km apart, on two small islands (Shodoshima and Kashima) off the coast of Shikoku; between this group and groups 1-3, D ~ 0.016. Blood proteins in group 5 — one locality on Boso Peninsula, south-central Honshu — are somewhat similar (D ~ 0.008) to those in group 6 — two localities on Yakushima, ca. 1000 km southwest of Boso Peninsula; be- tween groups 1-4 and groups 5-6, D ~ 0.025. Group 7, finally, includes only one locality on Shimokita Peninsula, northernmost Honshu; be- tween groups 1-6 and group 7, D ~ 0.035. Al- though localities in groups 3-7 are restricted to peninsulas or small islands, this pattern of distri- bution does not entirely distinguish them from lo- calities in group 1, some of which are similarly distributed on peninsulas and small islands (Fig. 20). Of the 32 blood-protein loci studied by Nozawa et al., eight are monomorphic and 24 are poly- morphic, either in M. fuscata or in the Chinese M. mulatta sample or in both (Table 27). At 19 of the 24 polymorphic loci, the seven M. fuscata groups are homogeneous with respect to their ma- jor alleles, and at 15 of these 19 loci, the M. mu- latta sample also shares the same major alleles. The four loci at which the common major alleles in M. fuscata groups differ from the major alleles in the M. mulatta sample are ADA, Dia, TBPA, and Tf; high frequencies of the major alleles at these four loci may be shared derived characters in M. fuscata. At 5 of the 24 polymorphic loci, the seven M. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 37 Fig. 18. Geographic variation in mtDNA sequence haplotypes in Macaca fuscata (Hayasaka et al., 1991, p. 401), sample size in parentheses; for details, see Table 23. fuscata groups are heterogeneous with respect to their major alleles, and at all five of these loci, the major alleles in group 1 are the same as the major alleles in the M. mulatto sample (Table 27); this suggests that high frequencies of these alleles may represent the primitive character state in M. fuscata. If so, the primary blood-protein diver- gence of group 7 (the most genetically divergent of the groups) from the ancestral M. fuscata stock was an increase in the frequency of allele Hb-(3: 2 (and probably also of allele PGM-II:2), the pri- mary divergence of groups 5 and 6 was an in- crease in the frequency of allele PGM-II:2, the primary divergence of group 4 was an increase in the frequency of allele CA-I;J„ the primary di- vergence of group 3 was an increase in the fre- quency of allele Gc:b, and the primary divergence of group 2 was an increase in the frequency of 38 FIELDIANA: ZOOLOGY Table 23. cf. Fig. 18. Local variation in mtDNA sequence haplotypes in Macaca fuscata (Hayasaka et al., 1991, p. 401); Approx. Reported mtDNA haplotypes distance MtDNA from ] Interrelated mtDNA haplotypes pre- haplotypes of uncer- Locality (listed in northeast- ceding locality Repeatable sequence status tain - relation- Island Prefecture to-southwest order) (km) Single Double Triple ships N Group II/IA'II Honshu Aomori Shimokita Peninsula — II 1 Honshu Tochigi Nikko 510 11(7)' VI (1)' 8 Honshu Nagano Shiga 100 II 15 Honshu Kanagawa Izu Peninsula 230 V 1 Honshu Fukui Takahama 320 P VIP 18 Honshu Kyoto Arashiyama 60 II 6 Honshu Osaka Minoo-city 20 II 1 Honshu Tottori Wakasa 120 II 8 Shodoshima Kagawa Shodoshima 90 II Group III/IVA^III 2 Honshu Okayama Gagyusan 60 111 5 Kyushu Oita Takasakiyama 250 V 3 Yakushima Kagoshima Yakushima 340 IV3 VHP 32 ' Dimorphic. ' Heteroplasmic (12/18). ^ Heteroplasmic (29/32). allele MDH:2. Changes in frequency of minor al- leles apparently have played a lesser role in the genetic differentiation of M. fuscata groups (No- zawa et al., 1991, p. 427; 1996, p. 27). Independent random genetic drift in semi-iso- lated local populations probably accounts for much of the observed blood-protein variation in M. fuscata (Nozawa et al., 1991, p. 431; 1996, p. 32). Following are supplementary comments con- cerning two striking disjunctions in blood-protein group distribution: 1. Group 4 populations, characterized by rela- tively high frequency of allele CA-l-d^, apparently are restricted to Shodoshima and Kashima, two small islands that are more than 200 km apart (Fig. 20); between these two islets is Shikoku, a large island apparently inhabited by group 1 pop- ulations. Two alternative hypotheses may apply here: (a) high frequencies of CA-I-ti, evolved in- dependently on Shodoshima and Kashima, or (b) CA-I-^2 populations formerly were widespread on Shodoshima, Shikoku, and Kashima but subse- quently were replaced on Shikoku by group 1 populations with high frequency of CA-I-a (cf. Nozawa et al., 1991, p. 431; 1996, p. 31). 2. Groups 5 and 6 populations, characterized by high frequencies of allele PGM-II:2, are dis- tributed ca. 1000 km apart on Boso Peninsula (south-central Honshu) and Yakushima (southern- most island inhabited by M. fuscata), respectively (Fig. 20); PGM-II:2 also is relatively frequent (41.1%) in group 7 populations on Shimokita Pen- insula (northernmost Honshu). As above, two al- ternative explanatory hypotheses may apply here: (a) high frequencies of PGM-II:2 evolved inde- pendently on Shimokita Peninsula, Boso Penin- sula, and Yakushima, or (b) PGM-II:2 formerly was widespread on the Japanese archipelago but subsequently has virtually disappeared every- where except on Shimokita Peninsula, Boso Pen- insula, and Yakushima (cf. Hayasaka et al., 1987, p. 513; Nozawa et al., 1991, p. 433; 1996, p. 32). The pattern of geographic variation of blood- protein groups in M. fuscata (Fig. 20) differs strikingly from the pattern of geographic variation of mtDNA types in this species (Figs. 18-20). Three major differences may be noted: (1) unlike blood-protein groups, mtDNA types are abruptly discontinuous in west-central Honshu; (2) unlike blood-protein groups, mtDNA type on Shimokita Peninsula is not strongly divergent from that else- where on Honshu; and (3) also unlike blood-pro- tein groups, mtDNA type on Shodoshima is not divergent from that in neighboring central Hon- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 39 8 2 9 4 10 11 35 36 37 38 3 38 + 1 1 39 2 40 2 41 2 41 - 1 2 41 + 1 1 42 1 43 1 Table 24. Mitochondrial DNA haplotypes detected in Macaca fiiscata at localities sampled by Kawamoto (2002, p. 60); cf. Fig. 19. Haplotype no. N Localities' Group I haplotypes Kurokawa-mura Arakawa-mura; Yokoze-machi Ashigakubo, Hinohara-mura; Itsukaichi; Tabayama-mura 1 Okutama-machi Tabayama-mura 2 Awajishima Gagyusan Anan-city Hiromi-cho; Nakamura-city; Nametoko Matsuno-cho Kisuki-cho; Ooda-city Kugino-mura; Takasakiyama Kanoya-city; Kojima Sagara-mura; Satsuma-cho Kushima-city Yakushima 1 Yakushima 2 Group II haplotypes 21 Ajigasawa-machi; Asahi-mura; Fukushima-city; Genbikei; Goyosan 1; Hachimori-machi; Higashine-city; Kaminoyama-city; Kanita-machi; Kashima-machi; Kinkazan; Ogachi-ma- chi; Shichigashuku-machi; Shimokita 1; Shimokita 2; Shimokita 3; Shimokita 4; Soma- city; Takahatamachi; Tsugaru-touge; Yamadera Goyosan 2 Nikko 1 Nikko 2 Boso Kyonan-machi Tsukui-machi Nishiizu-cho Atami-city; Odawara-city Aikawa-machi; Fujiyoshida-city; Kiyokawa-mura 1; Nishikatsura-cho Sanada-machi; Shiga-kogen; Sugadaira Karuizawa-machi Hodaka 1 ; Omachi-city; Outaki-mura Kamiichi-machi Oosawano-machi Hodaka 2 Unazuki-machi 1 Arashiyama 1; Asahi; Hosoiri-mura; Mihama-cho, Mikata-gun; Nishiazai-cho; Sodoshima; Unazuki-machi 2 Neo-mura Inuyama-city; Kiyokawa-mura 2; Tatsuyama-mura; Shinshiro-city; Toei-cho Mukawa-mura Shichisou-cho Takahama-cho Adogawa-cho 3^ Hatasho-cho Eigenji-cho; Hino-cho; Kameyama-city; Tsuchiyama-cho Hokusei-cho; Inabe-cho; Taga-cho Kozagawa-cho Hongu-cho Kanaya-cho; Koka-cho 1 ; Kumano-city; Mihama-cho, Minamimuro-gun; Owase-city; Oya- mada-mura; Tsubaki Adogawa-cho 1 ; Adogawa-cho 2; Arashiyama 2; Hieizan; Ibuki-cho; Kohoku-cho; Koka- cho 2; Minoo-city; Natasho-mura; Otsu-city; Sasayama-cho; Takatsuki-cho; Yogo-cho Wakasa-cho ' For details, see Gazetteer, Appendix 4; two specimens were sampled at each of nine localities (indicated by number "1" or "2" following a locality name), three specimens were sampled at one locality (Adogawa-cho 1-3), and four specimens were sampled at one locality (Shimokita 1-4). - Y. Kawamoto (pers. comm., Nov. 2003). 40 FIELDIANA: ZOOLOGY 2 1 3 1 4 1 5 1 6 1 12 1 13 1 14 2 15 4 16 3 16 + 1 1 17 3 17 + 1 1 17 + 2 1 18 1 19 1 20 7 20 + 1 1 21 5 22 1 23 1 26 1 26 + 1 1 27 1 28 4 29 3 30 1 31 1 32 7 33 13 34 1 130° 135' Fig. 19. Geographic variation in mtDNA sequence haplotypes in Macaca fuscata (Kawamoto, 2002, p. 60); for details, see Table 24. shu. The discrepancies between the distributions of these two types of genetic characters presum- ably are related to their respective modes of in- heritance— biparental for blood proteins and uni- parental (maternal) for mtDNA types (cf. Melnick et al., 1993, p. 290; Tosi et al., 2000, p. 138). Because female macaques are much more philo- patric than males, mtDNA types would be ex- pected to disperse more slowly than blood pro- teins; the distribution of mtDNA types therefore would be expected to preserve evidence of more ancient zoogeographic barriers or dispersal routes. Except for the distinctiveness of the Yakushima population, neither blood-protein group distribu- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 41 Table 25. Mean mtDNA divergence estimates reported between Macaca fuscata and M. mulatta, M. cyclopis, and M. fascicularis. Mean divergence (Div.) estimates (%) References Analytical techniques Intra- specific M. fuscata M. mulatta Divergence from M. fuscata M. cyclopis M. fasci- cularis Div. N Div. N Div. N Div. N Hayasaka et al. (1988b. p. 277) Hayasaka et al. (1988a. p. 635) Hayasaka et al. (1996, p. 1049) Zhang & Shi (1989, p. 335) Zhang & Shi (1993. p. 12) Melnick et al. (1993, p. 286) Morales & Melnick (1998, p. 13)^ Kawamoto et al. (1999. p. 57) LP 17 enzymes 1.6 10 3.6 1 3.7 1 6.2 1 uencing: 0.9 kb' — 1- 3.7 1 — 0 8.8 1 uencing: 0.896 kb' 1.6 2"* 3.6 1 3.2 1 9.4 1 LP 10 enzymes — 35 2.8 4 — 0 — 0 LP 16 enzymes — P 3.8 2 — 0 6.5 1 LP 15 enzymes 1.7 10^ 3.7 19" 3.9 V 6.0 P LP 22 enzymes 0 2 0.6 3 1.0 2 3.2 3 ue ncing: 0.464 kb* — 1 — 0 2.4 1 — 0 ' Includes genes for tRNA"'\ tRNA^", tRNA^*^", 3' region of ND4, and 5' region of ND5. - One of 10 specimens previously studied by Hayasaka et al. (1988b, p. 277). ' Hindlll fragment. ■* Includes one of 10 specimens previously studied by Hayasaka et al. (1988b. p. 277). ' Data from Hayasaka et al. (1986, p. 352: 1988b, p. 277). * Includes data from one specimen reported by Hayasaka et al. (1988b, p. 277). ■^ Cf. Tosi et al. (2002. p. 168). who sequenced a 1.5-kb fragment of mtDNA (3' end of 12S ribosomal gene; tRNA^'^'; 5' end of 16S ribosomal gene) in M. fuscata (n = 3), M. mulatta (n = 5), M. cyclopis (n = 2), and M. fascicularis (n - 17); although divergence estimates are not specified in this work, the authors' bootstrap consensus tree (fig. 2a) indicates that mtDNA in M. fuscata is more similar to that in M. mulatta and M. cyclopis than to that in M. fascicularis. ^ D loop. tions nor mtDNA type distributions bear much re- lationship to the present configuration of land and water in the Japanese archipelago (Figs. 18-20). Instead, as indicated above, these distributions may reflect the climate, vegetation, land bridges, and water gaps that existed during previous stages in the evolutionary history of M. fuscata. Hemoglobin Amino-Acid Sequence The amino-acid sequence of hemoglobin in M. fuscata has been compared with that in M. mu- latta (Matsuda, 1982, p. 293). In these species, the a-chains of hemoglobin are identical, and the P-chains differ only in a single amino acid. Table 26. Sister-group relationships of Macaca fuscata indicated by sequencing of paternal, maternal, and bi- parental markers (Tosi et al.. 2003. pp. 1426-1429). Loci studied M. fuscata specimens' Indicated sister group- '-chromosome: Nos. 29- -31 SRY, TSPY, 3. 1 kb ItDNA: Nos. 29- -31 12s ribosomal gene, 1.5 kb kUtosome: Nos. 29- -31 C4 long intron 9, 3.3 kb LUto.some: No. 29' IRBP intron 3, 1.6 kb No. 31' (CYC (MUL India (MUL Burma/China, FAS Thailand/ Cambodia/Vietnam))) (MUL Burma) ((SIN)(RAD)(ASS. THD) (FAS West Malaysia) (FAS Thailand) ' Origin "Japan," precise localities unspecified. - Key to species abbreviations: /a.vc/cM/am group — FAS = M. fascicularis. CYC = M. cyclopis. MUL = M. mulatta: sinica group — SIN - M. sinica, RAD = M. radiata. ASS = M. assamensis, THI = M. thibetana. -' Sister-group relationships of M. fuscata specimen no. 30 are not reported for this locus. 42 FIELDIANA: ZOOLOGY 130° 135° 140° Fig. 20. Geographic distribution and genetic interrelationships of blood-protein groups in Macaca fuscata (Nozawa et al., 1991, p. 418; 1996, p. 4); for details, see Table 27. Antibodies to Cedar Pollen Antigens Natural allergy to the pollen of Japanese cedar (Cryptomeria japonica) has been detected in M. fuscata (Nakamura et al., 1991, p. 643). In six provisioned free-ranging groups and three captive groups studied in nine prefectures (n = 276), the incidence of antibodies to crude cedar pollen an- tigen averaged 16.3% and varied from 0% to 31.3% (Table 28). The 45 positive sera were fur- ther tested for reactivity to two purified cedar pol- len antigens. Cry j I and Cry j II; of the positive sera, 23 reacted to both antigens, 21 reacted only to Cry j I, and one reacted only to Cry j II. The factor or factors responsible for prefectural vari- ation in M. fuscata reactivity to cedar pollen an- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 43 Table 27. Mean frequency' (%) of major alleles at 24 loci in blood-protein groups of Macaca fuscata (cf. Fig. 20) and frequency of corresponding alleles in southern Chinese M. mulatta sample;^ for details, see Nozawa et al. (1991, p. 418; 1996, p. 4). Fre- Blood-protein groups^ and mean frequencie « quency (%) in M. - of major alleles in M. fuscata Group mulatta Group 2: Group 7: Shi- sample: Group 1: central Group 3: Group 4: 5: Boso mokita southern widely Honshu Izu Kashima and Penin- Group 6: Penin- China Major distributed (1 local- Peninsula Shodoshima sula (1 Yakushima sula (1 (n = Locus allele (28 localities) ity) (3 localities) (2 localities) locality) (2 localities) locality) 150) Loci at which M. . fuscata \ groups are heterogeneous with I respect to major alleles CA-I a 97.0 ± 4.5 85.9-100.0 100.0 100.0 34.6 17.6-51.6 100.0 100.0 100.0 93.0 di 2.3 ± 4.0 0-14.1 0 0 65.4 48.4-82.4 0 0 0 5.3 Gc a 95.6 ± 5.8 78.6-100.0 76.9 45.7 ± 13.9 35.7-61.5 100.0 100.0 100.0 78.8 98.0 h 2.1 ± 5.3 0-21.4 1.9 54.1 ± 14.3 37.7-64.3 0 0 0 21.2 0 Hb-3 1 99.9 ± 0.4 98.4-100.0 100.0 85.6 ± 8.6 78.5-95.1 100.0 100.0 100.0 14.3 100.0 2 0.1 ± 0.4 0-1.6 0 14.4 ± 8.6 4.9-21.5 0 0 0 85.7 0 MDH 1 97.3 ± 4.5 84.4-100.0 48.1 95.1 ± 3.5 91.8-98.7 100.0 100.0 100.0 91.4 100.0 2 1.5 ± 3.4 0-14.3 51.9 3.9 ± 3.7 1 .3-8.2 0 0 0 0 0 PGM-II 1 99.7 ± 0.9 96.4-100.0 100.0 100.0 97.7 95.4-100.0 44.2 8.3 2.6-13.9 52.9 96.3 2 0.3 ± 0.9 0-3.6 0 0 0 55.8 91.7 86.1-97.4 47.1 3.7 Loci at which M . fuscata : groups are homogeneous with respect to major alleles Acp A 99.1 ± 2.3 89.3-100.0 100.0 100.0 100.0 100.0 100.0 100.0 98.7 ADA^ I 100.0 100.0 100.0 100.0 100.0 100.0 100.0 0 AK I 98.1 ± 3.8 85.0-100.0 100.0 98.6 ± 1.3 97.4-100.0 99.4 98.9-100.0 100.0 100.0 100.0 99.0 Alb A 100.0 ± 0.01 99.9-100.0 88.5 100.0 100.0 100.0 100.0 100.0 97.7 Alp A 100.0 100.0 100.0 100.0 99.5 100.0 100.0 100.0 CA-II a 99.2 ± 2.2 90.5-100.0 82.7 100.0 100.0 71.8 100.0 85.7 97.3 Cell Es 1 98.4 ± 3.5 87.5-100.0 100.0 83.1 ± 10.4 76.1-95.1 100.0 100.0 100.0 100.0 99.3 ChEs I 99.9 ± 0.2 99.2-100.0 100.0 100.0 99.9 99.8-100.0 100.0 100.0 100.0 100.0 Dia^ A 99.8 ± 1.0 94.6-100.0 100.0 100.0 100.0 100.0 100.0 100.0 13.4 Es-D 1 99.7 ± 1.2 95.0-100.0 100.0 98.4 ± 2.8 95.1-100.0 100.0 100.0 100.0 100.0 99.7 IDH 2 100.0 ± 0.01 99.9-100.0 100.0 100.0 100.0 100.0 100.0 100.0 82.9 LDH-A 1 98.1 ± 3.6 84.2-100.0 96.2 100.0 100.0 100.0 100.0 100.0 100.0 LDH-B I 99.9 ± 0.3 98.2-100.0 100.0 100.0 93.4 86.8-100.0 100.0 100.0 100.0 100.0 44 FIELDIANA: ZOOLOGY Table 27. Continued. Fre- Blood l-protein groups' and mean frequencies quency (%) In M. - of major alleles in M. fuscata Group mulatto Group 2: Group 7: Shi- sample: Group 1: central Group 3: Group 4: 5: Boso mokita southern widely Honshu Izu Kashima and Penin- Group 6: Penin- China Major distributed (1 local- Peninsula Shodoshima sula (1 Yakushima sula (1 (n = Locus allele (28 localities) ity) (3 localities) (2 localities) locality) (2 localities) locality) 150) PGM-1 1 99.8 ± 0.8 95.8-100.0 100.0 91.5 ± 4.1 86.9-94.9 100.0 100.0 100.0 100.0 98.7 PHI 1 94.2 ± 7.9 73.8-100.0 100.0 100.0 99.9 99.7-100.0 81.4 100.0 82.9 97.3 Pi C 97.7 ± 4.7 80.4-100.0 100.0 100.0 99.7 99.5-100.0 94.2 100.0 100.0 98.7 6PGD A 100.0 100.0 100.0 100.0 100.0 100.0 100.0 97.6 TBPA S 100.0 100.0 100.0 100.0 100.0 100.0 100.0 25.0 TP F 96.3 ± 5.3 77.9-100.0 92.3 81.3 ± 4.9 78.3-86.9 100.0 100.0 100.0 100.0 8.7 ' Unweighted mean of local allele frequencies within each M. fuscata group; standard deviation and extremes also specified, where pertinent. - Excludes eight loci monomorphic in M. fuscata and M. mulatta samples: a,; Amy; Cat; Cp; G6PD; Hb-a; Hp; To. ' Group localities (island name in capitals, prefecture name in italics, locality name in parentheses, sample size in square brackets): Group 1 [2566 samples]— AWAJISHIMA, Hyogo (Awajishima [150]); HONSHU, Aichi (Oshiro [42]), Fukui (Mikata [59], Takahama [49]), Fukushima (Fukushima-city [31]), Hiroshima (Kochi [34]), Hyogo (Ka- suga-cho [70]), Kyoto (Arashiyama [228], Hiyoshi-cho [49], Ine-cho [30], Wachi-cho [31]), Nagano (Shigakogen [208]), Nara (Muroji [19]), Okaxama (Gagyusan [172], Katsuyama-cho [28]), Osaka (Minoo-city [19]), Shiga (Ryo- zenyama [120]), Shizuoka (Tenryu-city [12]), Tochigi (Nikko [21]), Tottori (Wakasa [25]); KOJIMA, Miyazaki (Koji- ma [88]); KYUSHU, Fukuoka (Kawaradake [52]), Kumamoto (Itsuki-mura [21]), Miyazaki (Kushima-city [28]), Oita (Kamae-machi [30], Takasakiyama [866]); SHIKOKU, Ehime (Nametoko [26]), Kochi (Odo [58]). Group 2 [26 samples] — HONSHU, Ishikawwa (Hakusan [26]). Group 3 [297 samples] — HONSHU, Kanagawa (Yugawara-machi T [61]), Shizuoka (Hagachi [158], Ihama [78]). Group 4 [215 samples]— KASHIMA, Ehime (Kashima [17]); SHO- DOSHIMA, Kagawa (Shodoshima [198]). Group 5 [94 samples]— HONSHU, Chiha (Takagoyama [94]). Group 6 [176 samples]— YAKUSHIMA, Kagoshima (Kamiyaku-cho [54], Yaku [122]). Group 7 [35 samples]— HONSHU, Aomori (Wakinosawa [35]). Total samples, 3409. ^ Allele frequencies at these three loci recently have been studied in 20 blood samples collected at four localities in Mie Prefecture (HONSHU: Aoyama; Inabe-cho; Kiwa; Omiya) and four localities in Wakayama Prefecture (HON- SHU: Nakatsu; Kanaya-cho; Kozagawa-cho; Motomia) (Kawamoto et al., 1999, p. 55; 2001, p. 15); reported fre- quencies are: ADA, allele /—100.07c; Dia, allele A— 100.0%; and Tf, allele F— 92.5%, allele G— 7.5%. tigens have not been determined (Hashimoto et al., 1994, p. 396). Karyology Classical staining techniques revealed that the diploid chromosome number in M. fuscata is 42 and that chromosome morphology in this species is generally similar to that in other macaques and also to that in other papionins (Chiarelli, 1962, p. 410). Although banding techniques subsequently confirmed these general similarities, one species- specific pericentric inversion was discovered on chromosome 15 oi M. fuscata (n == 10) (Stanyon et al., 1983, p. 142; 1988, p. 6); banding tech- niques also revealed that M. fuscata, like some other papionin species, is polymorphic with re- spect to size of the nucleolar organizer region on chromosome 1 3 and size of the C band on the Y chromosome. In a survey to determine whether congenital limb malformation in M. fuscata is related to chromosomal abnormality, Minezawa et al. (1990, p. 572) studied the karyotypes of 257 monkeys representing seven local populations on five Jap- anese islands (Awajishima, n = 28; Honshu, n = 95, three populations; Kojima, n = 60; Kyushu, n = 53; Shodoshima, n = 21). No relationship was found between limb malformation and chro- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 45 Table 28. Incidence of serum specimens positive for antibodies to crude cedar pollen antigen and incidence of serum specimens with positive reactions to purified cedar pollen antigens (Cry j I, Cry j II) in prefectural samples of Macaca fuscata. ' Positive for antibodies to crude antigen Positive reactions to 1 purified antigens Prefecture Cryj I and Cry j II Cryj I only Cryj II only N Honshu Nagano Gunma Tokyo Aichi Kyoto Okayama Osaka 2(9.1%) 8 (27.6%) 1 (5.0%) 5(16.1%) 5(31.3%) 0 I (6.3%) 0 5 1 2 1 0 0 2 2 0 3 4 0 1 0 1 0 0 0 0 0 22 29 20 31 16 25 16 Awajishima Hyogo 13(25.5%) 10 Kojinia 3 0 51 Miyazaki 10(15.1%) 4 6 0 66 Totals 45(16.3%) 23(51.1%) 21 (46.7%) 1 (2.2%) 276 Reference: Hashimoto et al. (1994, p. 395; cf. Nakamura et al.. 1991. p. 645: Sakaguchi et al.. 1992. p. 325). mosomal abnormality (cf. Iwamoto & Hirai, 1970, p. 396), and no geographic variation in kar- yotype was reported. Parasites Protistans One species of flagellate and four species of amebas have been detected at an incidence of 20- 75% in fecal samples of M. fuscata at Shimokita Peninsula, which is at the northern limit of the monkey's distribution (Table 29). The ciliate Bal- antidium coli has been recorded in a M. fuscata captive by Tschemer (1985, p. 226). Malarial parasites (sporozoans) apparently are naturally absent from M. fuscata, judging from 547 blood specimens collected on Honshu, Sho- doshima, Miyajima, Kyushu, and Yakushima (O- tsuru & Sekikawa, 1979, p. 347). The absence of malaria in M. fuscata presumably is a conse- quence of Japan's geographic location, which is north of the range of the Leucosphyrus group of anopheline mosquitoes; this group of mosquitoes apparently is essential in the transmission of sim- ian malaria in Asia (Coatney et al., 1971; Fooden. 1994, p. 582). M. fuscata is susceptible to exper- imental infection with Plasmodium coatneyi (Ka- wai et al., 1998, p. 101), which is a natural ma- larial parasite of M. fascicularis in Southeast Asia (Fooden, 1994, p. 576). The sporozoan Babesia sp. has been detected in 4 of 93 blood specimens (4.3%) collected from members of a M. fuscata group that had been translocated in 1957 from Yakushima to Honshu (Ohirayama) (Otsuru & Sekikawa, 1979, p. 347). This sporozoan has not been detected in nontran- slocated groups of M. fuscata, either on Yakushi- ma (n = 39) or on Honshu, Shodoshima, Miya- jima, and Kyushu (n = 415). Table 29. Incidence of protistan parasite species de- tected in fecal samples of Macaca fuscata on Shimokita Peninsula, Aomori Prefecture (n = 20).' Incidence Species No. % Flagellate Giardia sp. 10.0 Amoebas- Eiuamoeba coli 5 25.0 E. histolytica 15 75.0 Endolimax nana 14 70.0 lodamoeba buetschlii 5 25.0 ' Reference: Takada et al. (1981, p. 70). - The same four species of amoebas have been detect- ed in fecal samples of a laboratory colony of M. fuscata (Koyama et al.. 1978. p. 60). 46 FIELDIANA: ZOOLOGY Table 30. Incidence of cestode parasites (Bertiella studeri, possibly also other species) in local samples' of Macaca fuscata. Cestode Latitude incidence Locality Island (N) (%) N References^ Shimokita' Honshu 41°10' 0 35 1,2 Kinkazan Kinkasan 38°17' 0 23 2 Nikko Honshu 36°44' 0 23 2 Shiga Honshu 36°44' 0 141 2,3,4 Hakusan"* Honshu 36°15' 0 25 2 Mikata Honshu 35°33' 3.2 63 4 Kiso Honshu 35°31' 0 7 2 Takagoyama Honshu 35°13' 0 24 2 Sakamoto Honshu 35°08' 12.5 8 2 Arashiyama Honshu 35°00' 20.5 156 2,4,5 Hasumi Honshu 34°53' 0 47 4 Hagachizaki Honshu 34°4r 0 38 2 Shodoshima Shodoshima 34°30' 0 95 2,4 Kochi Honshu 34°28' 0 34 4 Mihara-city Honshu 34°24' 0 119 4 Awajishima Awajishima 34°14' 0 20 2 Takasakiyama Kyushu 33°10' 42.6 155 4,6 Misho/Nishiumi Shikoku 32°58' 0 36 7 Kojima Kojima 31°27' >8.05 286 2, 4, 8, 9 Yakushima'' Yakushima 30°30' 3.8 157 2,4 Totals — — -8.7 1492 — ' Fecal samples, except Sakamoto and Misho/Nishiumi, which are necropsy samples. 2 Key to references: /. Takada et al. (1981, p. 70). 2. Gotoh (2000, p. 294). 3. Nigi et al. (1975, p. 36). 4. Nigi (1983, p. 38). 5. Machida and Sano (1974, p. 77). 6. Kagei and Hasegawa (1974, p. 236). 7. Itoh et al. (1988, p. 236). 8. Kawai (1974, p. 17). 9. Horii et al. (1982, pp. 416, 418). ^ Includes Wakinosawa. ^ Includes Kamuri. ' Cestode eggs reported in 23 of 73 samples (Reference 8), and "few cestode segments" reported in 74 samples (reference 9); 139 samples negative for cestodes. ^ Cf. Tanaka et al. (1962, p. 115) and Hayama and Nigi (1963, p. 103), who report B. studeri in two of 66 Yakushima captives at the Japan Monkey Centre, Inuyama (Honshu), and Nigi (1982, p. 248), who reports no cestodes in 57 Yakushima monkeys from a population that had been translocated to Ohirayama (Honshu). Trematodes No flukes were detected in parasitological nec- ropsy of 36 M. fuscata specimens collected at Misho/Nishiumi, Ehime Prefecture, Shikoku (Itoh et al., 1988, p. 236). Cestodes north of ca. 36°N. The incidence of tapeworm par- asites in necropsy samples (1/44 = 2.3%) appar- ently is somewhat less than that in fecal samples (129/1448 = 8.9%). Tscherner (1985, p. 226) has reported that a M. fuscata captive was infected with the tapeworm Hymenolepis diminuta, normally a parasite of ro- dents. Tapeworm infections are relatively rare — 8.7% incidence — in natural populations of M. fuscata (Table 30), and only one species of tapeworm, Bertiella studeri, has been positively identified as a natural parasite in these monkeys. The geo- graphic distribution of tapeworms in M. fuscata generally appears sporadic, although it may be significant that no infection has been recorded in the cool, northern part of the monkey's range. Nematodes Six species of nematodes are known to parasit- ize natural populations of M. fuscata (Table 31); this excludes two nematodes {Ternidens deminu- tus and Oxyuridae gen. sp.) reported in captive M. fuscata by Tscherner (1985, p. 227). Of the nematode parasites in natural populations, Trich- uris trichiura is the most prevalent (60.1% inci- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 47 Table 31. Incidence {%) of nematode parasites in local samples' of Mocaca fuscata. Strep- Oeso- Gongylo- to- Latitude Stron- phago- nenma phara- Locality Island- (N) Trichuris gyloides stomiim spp. gus' N References^ Shimokita" H 41°10' 60.0 48.6^ 0 0 0 35 1.^ 2 Kinkazan Ki 38°I7' 91.3 13.0 8.7 0 0 23 2 Nikko H 36°44' 100.0 0 0 0 0 23 2 Shiga H 36°44' 65.7 0 0 0 0 216 2, i," 4, 5'o Hakusan" H 36°15' 63.1 0 23.1 0 1.5 65 2, 4 Mikata H 35°33' 52.4 15.9 44.4 — 22.2 63 5'o Kiso H 35°3r 100.0 71.4 0 0 0 7 2 Takagoyama H 35° 13' 20.8 16.7 8.3 0 0 24 2 Sakamoto H 35°08' 87.5 25.0 100.0 0 75.0 8 2 Arashiyama H 35°00' 75.6 41.7 3.8 0 69.9 156 2. 5:^' 6 Hasumi H 34°53' 76.6 46.8 0 — 23.4 47 5'o Hagachizaki H 34°4r 92.1 42.1 0 0 78.9 38 2 Shodoshima Sho 34°30' 67.4 16.8 20.0 0 67.4 95 2, 5,'° 7 Kochi H 34°28' 58.8 38.2 2.9 — 76.5 34 J'o Mihara-city H 34°24' 81.5 44.5 5.9 — 42.0 119 5'o Awajishima A 34° 14' 95.0 55.0 0 0 20.0 20 2 Takasakiyama Ky 33° 10' 67.9 30.5 86.2 — 85.6 305 4. 5.'<'S Misho/Nishiumi Shi 32°58' 36.1'- 36.1'- 5.6 — 69.4 36 9 Aira Ky 31°44' — — — >0" — •7 W Kojima'^ Ko 31°27' 49.6 35.1 31.9 0 87.0 276 2. 4, J.'o 77'5 Toimisaki Ky 31°22' 0 43.8 69.6 — 72.3 112 4 Yakushima'^ Y 30°30' 45.2 42.0 29.9 10.2 43.9 157'^ 2, 5'o Totals — — 60.1 29.9 30.4 <1.0 53.3 1859'^ — ' Fecal samples, except Sakamoto. Misho/Nishiumi. and Aira. which are necropsy samples. - Abbreviations: A = Awajishima; H = Honshu; Ki = Kinkasan; Ko = Kojima; Ky = Kyushu; Shi = Shikoku; Sho = Shodoshima; Y = Yakushima. ' Goiigylonema macrogubernacuhim and/or G. pukhrum (see reference 2). Excluded are Gongylonema infections reported in translocated captives by Uni et al. (1994. p. 128); as indicated by Gotoh (2000. p. 296). some or all of these infections may have been acquired in captivity. ■* Supplementary record of this species (Yamaguti. 1935. p. 453): "in the small intestine of a Japanese monkey from Nara Prefecture," 33°5r-34°44'N. '^Key to references: /. Takada et al. (1981. p. 70). 2. Gotoh (2000, pp. 294. 296. cf. 1989, p. 9). 3. Nigi et al. (1975. p. 36). 4. Usui and Horii (1982. p. 272). 5. Nigi (1982. p. 248; 1983. p. 38). 6. Machida and Sano (1974. p. 77); Machida et al. (1978. p. 1). 7. Tanaka et al. (1963. p. 93). 7. Kagei and Hasegawa (1974. p. 236). 9. Itoh et al. (1988. p. 236). 10. Uni et al. (1992, p. 221). 11. Kawai (1974, p. 17). *' Includes Wakinosawa. ^ Cited in reference / as Strongyloides sp. near fulleborni. ** Absence of O. aculeatum. Gongylonema sp., and S. pigmentotus inferred. '* In this reference, parasites, except T. irichiiira, are cited by generic names only. '° In this reference, Trichuris trichiura is cited as Trichocephalus trichiiirus. " Includes Kamuri. '- Corrected value; apparent miscalculation in cited reference. '■' One monkey reported positive for G. pukhrum. '■*Cf. Horii et al. (1982, pp. 419-421). '■* In this reference, parasites are cited by generic names only. "'Cf. Hayama and Nigi (1963. p. 103). Tanaka and Nigi (1967. p. 99). and Usui and Horii (1982. p. 272). who have investigated the incidence of nematode parasites in captive and translocated populations of Yakushima M. fuscata. ' '' For Gongylonema spp.. n = 59. '** For Gongylonema spp., n = 1761. dence) and is the only nematode parasite that is widely distributed in northern populations of M. fuscata; Streptopharagus pigmentatus is next most prevalent (53.6%). but this species is rare north of 35°08'N; Oesophagostomum aculeatum and Strongyloides fuelleborni are relatively un- common (30.47r and 29.9%. respectively); and Gongylonema macroguhernaculum and G. pul- 48 FIELDIANA: ZOOLOGY chrum apparently are restricted to M. fuscata pop- ulations that inhabit the southern islands Kyushu and Yakushima (10.2% incidence of Gongyione- ma spp. in a sample of 59 Yakushima M. fuscata). Heavy snowfall and low winter temperatures in the northern part of the range of M. fuscata ap- parently inhibit intermonkey transmission of most nematode parasites, except T. trichiura, which is transmitted by means of a resistant thick-walled egg (Nigi et al., 1975, p. 45; Usui & Horii, 1982, p. 273; Horii, 1983, p. 14; Gotoh, 1989, p. 9; 2000, pp. 292, 296). Although Hayama and Nigi (1963, p. 103) have tabulated Physaloptera caucasica and Physalop- tera sp. as nematode parasites of captive Yaku- shima M. fuscata, this appears to be a typograph- ical error. In their text (p. 102), these authors clearly specify that P. caucasica and Physalop- tera sp. were detected only in M fascicularis and Papio cynocephalus, not in M. fuscata, which di- rectly contradicts the corresponding tabular en- tries (cf. Tanaka & Nigi, 1967, p. 103). Seasonal variation of nematode parasite inci- dence in M. fuscata on Kojima has been studied by Imada et al. (1977, p. 15), Horii et al. (1982, p. 420), and Nigi (1982, p. 248; 1983, p. 39). On Kojima and possibly elsewhere, the incidence of T. trichiura, Streptopharagus pigmentatus, and O. aculeatum tends to decrease in summer, reaching a minimum in August, and tends to increase in winter, reaching a maximum in January-February (cf. Usui & Horii, 1982, p. 270). Evidence con- cerning seasonal variation in the incidence of Strongyloides fuelleborni is equivocal. The incidence of at least some nematode par- asites apparently also varies with the age of the host monkey (Imada et al., 1977, p. 15; Horii et al., 1982, p. 419; Gotoh, 2000, p. 295). In M. fuscata on Kojima, for example, the incidence of Strongyloides fuelleborni in samples of immatures (age <5 years; combined n = 187) is 75-100%, whereas the incidence in samples of old adults (age >13 years; combined n = ca. 120) is ca. 19- 30%. Sexual variation of the host does not appear to be an important factor in the incidence of nema- tode parasites in M. fuscata (Horii et al., 1982, p. 418; Gotoh, 2000, p. 295). Mite A "mite egg?" has been reported in one of 20 M. fuscata fecal samples collected on Shimokita Peninsula, northern Honshu (Takada et al., 1981, p. 70). Lice Pedicinus (Parapedicinus) obtusus japonicus has been recorded as an ectoparasite of M. fuscata at Kamikochi (Nagano Prefecture) and "Province of Shinano" (= Nagano Prefecture) by Kuhn and Ludwig (1967, p. 248), and P. eurygaster has been briefly mentioned as another louse ectopar- asite of this monkey by Tanaka and Takefushi (1993, p. 192); unidentified immature lice have been reported as ectoparasites of M. fuscata at Jigokudani Monkey Park, Shiga Heights, Nagano Prefecture (Tanaka & Takefushi, 1993, p. 192). Removal and ingestion of louse eggs apparently are major motivations underlying the grooming behavior of M. fuscata (Tanaka & Takefushi, 1993, p. 192; cf. Tanaka, 1995, p. 200; 1998, p. 1234). Tick At Jigokudani Monkey Park, Shiga Heights, Nagano Prefecture, during the course of 4475 ob- served grooming sequences, a single tick {Hae- maphysalis longicornis) was removed from the fur of a M. fuscata infant by its mother (Tanaka & Takefushi, 1993, p. 189). The tick was crushed to death by the mother but not eaten. Q Fever Bacterium Antibodies to Coxiella burnetii, the bacterium that causes Q fever, were detected in sera collect- ed from 15 of 54 M. fuscata specimens (28%) that were live-trapped in seven prefectures in northern and central Honshu (Ejercito et al., 1993, p. 483). Detailed results for each prefecture are unavail- able. Viruses Local variation in the incidence of antibodies to eight viruses has been studied in three groups of M. fuscata by Matsubayashi et al. (1992, p. 393) (Table 32). The incidence of antibodies to two of these viruses (SA-6, Cyno-EBV) is high in all three groups, the incidence of antibodies to FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 49 Table 32. Incidence {9c) of antibodies to eight vi- ruses in three local groups of Macaca fuscata. ' Translocated groups In-situ Arashi- group yama,- Wakasa,' Takasaki- Kyoto Tottori yama. Oita Prefecture. Prefecture. Prefecture. Honshu Honshu Kvushu Virus (n = 45) (n = 53) (n"= 30) SA-6 100.0 100.0 100.0 Cyno-EBV 100.0 96.2 100.0 Measles 93.0 32.1 66.7 STLV-1 60.0 64.2 16.7 Jap. Enc. 70.5^ 54.7 3.4-^ HSV-1 75.0^ 1.9 0 SIVmac 4.4 0 0 SV-5 0 0 0 ' Reference: Matsubayashi et al. (1992. p. 393). - Original locality of group: virological study con- ducted after translocation of group to outdoor enclosure at PRIKU (Inuyama. Aichi Prefecture. Honshu) in 1981. " Original locality of group: virological study con- ducted after translocation of group to outdoor enclosure at PRIKU (Inuyama, Aichi Prefecture. Honshu) in 1971. ^N = 44. ■^ N = 29. two other viruses (SIV^,;^^. SV-5) is low in all three groups, and the incidence of antibodies to four viruses (measles. STLV-1. Japanese enceph- alitis, HSV-1) is strongly variable in the three groups. The incidence of antibodies to STLV-1 also is strongly variable (0%-ca. 9Q9c in adults) in 41-42 local groups of M. fuscata that were studied by Hayami et al. (1984. p. 180: cf. Hay- ami, 1986. p. 36). The incidence of antibodies to another virus — SV 40 — that was studied in seven local groups of M. fuscata by Ichikawa et al. (1985. p. 326) varied from 50% to 100% (ex- cludes one small sample, n = 4. with 0% inci- dence). The explanation for local variation in the incidence of antibodies to certain viruses in M. fuscata groups is unclear. Antibodies to monkey B virus were absent in 8 1 M. fuscata specimens examined on Shodoshi- ma by Tanaka et al. (1963. p. 93) and in 46 M. fuscata specimens examined at Takasakiyama by Endo et al. (1964. p. 174): however, they were present in 7 of 61 captives (11.5%) originally from Yakushima that were studied by Endo et al. (1959, p. 229) and in 21 1 of 629 captives (33.57r) of unknown origin that were studied by Sato et al. (1998. p. 2(X)). Restriction fragment length polymorphism analysis and DNA sequencing in- dicate that the B virus that infects M. fuscata is genetically distinct from the B viruses that infect other macaque species (Ohsawa et al.. 2002, p. 559). The incidence of antibodies to six viruses — hepatitis A. HSV-1. measles. SIVm^c STLV-1, and SV-40 — apparently tends to increase with age in M. fuscata (Hayami et al.. 1984. p. 180: Ichi- kawa et al.. 1985. p. 326: Hayami. 1986. p. 36: Abe et al.. 1989. p. 88: Matsubayashi et al.. 1992, p. 395): in two groups of monkeys tested for hep- atitis A antibodies, antibody incidence increased from 0% to 30% at age 1 year to 100% at age 5 years (Abe et al., 1989, p. 89). Contrastingly, the incidence of antibodies to Cyno-EBV and SA-6 in two groups was 100% at age 1 year and re- mained high to age ^21 year (Matsubayashi et al.. 1992. p. 394).^ In captive groups of M. fuscata, the system of housing apparently affects the incidence of anti- bodies to hepatitis A (Table 33). In 43 individu- ally housed monkeys, the antibody incidence was 2.3%. and in 48 group-housed monkeys, the in- cidence was 70.8%. Tanaka et al. (1963, p. 93) have reported that M. fuscata on Shodoshima tested negative for an- tibodies to adenoviruses (n = 26) and influenza viruses (n = 14). Natural History Habitats The two primary habitats of M. fuscata are warm temperate evergreen broadleaf forest in the southern part of its geographic range and cool temperate deciduous broadleaf forest in the north- em part of its range (Suzuki, 1965. p. 31: Uehara, 1975. p. 392: Maekawa. 1978. p. 18). The ele- vational distribution of this species is known to extend to 3180 m on the summit of Yarigatake (mountain). Nagano Prefecture (S. Izumiyama, pers. comm.. 29 Dec. 2003). During relatively mild weather, M. fuscata has been observed in subalpine needleleaf forest at 3050 m on Ohami- dake (mountain), Nagano Prefecture (Izumiyama, 1987a. p. 8: 1987b. p. 42: cf. 1994. p. 478: Izu- miyama et al.. 2003. p. 466): during snowy winter months. M. fuscata in this part of central Honshu moves down to the montane broadleaf forest zone at elevations that do not exceed 1800 m (Izumi- yama. 1987b. p. 41: cf. Ashizawa. 1983, p. 13). A comprehensive bibliography of field studies 50 FIELDIANA: ZOOLOGY Table 33. Relationship fuscata. between housing system and incidence of antibodies to hepatitis A in captive Macaca Housing system N Hepatitis A antibody positive Incidence (%) References Individually housed Group housed Unspecified, possibly mixed 43 48 294 1 34 167 2.3 70.8 56.8 Chiba et al. (1980, p. 165) Chibaetal. (1980, p. 165) Abeet al. (1989, p. 88) of M. fuscata (1975-1992), most of which include habitat information, has been published by Wa- tanabe (1993, p. 33). Terrestriality/Arboreality Detailed information concerning the amount of time that M. fuscata spends on the ground and in trees is available for only one population, which was observed June 1995-January 1997 at Tsubaki Wild Monkey Park, Shirahama-cho, Wakayama Prefecture (Chatani, 2003, p. 19). In this popula- tion, the average proportion of time that monkeys were observed on the ground during daytime hours was 68.3% for males and 39.8% for fe- males. Local circumstances presumably influence the degree of terrestriality or arboreality in M. fus- cata. Based on research at Tsubaki Park, Chatani (2003, p. 21) has characterized M. fuscata as semiterrestrial — behaviorally intermediate be- tween arboreal M. fascicularis and terrestrial M. nemestrina. Macaca fuscata usually sleeps in trees (Wada, 1975, p. 12; Wada & Tokida, 1985, p. 3). How- ever, sleeping on the ground has been reported on two small islands — Kinkazan (Takahashi, 1997, p. 60) and Kojima (Mori, 1979a, p. 43; pers. comm.); no predators of monkeys inhabit either of these islands. This species reportedly swims well. In 1970, a young male monkey swam more than 600 m from Kojima to Torishima, an island that lies beyond Kojima (Mito, 1980, p. 29). Group Size and Composition Mean group size in 117 groups of wild Japa- nese macaques is 40.8 ± 28.95 individuals (ex- treme values, 10-161 individuals) (Table 34; cf. Izumiyama et al., 2003, p. 471). Groups with as few as three or four members have been reported. but such small groups probably are ephemeral: on Yakushima, a group comprised of three individ- uals— adult male, adult female, 4-year-old fe- male— became extinct after less than 1 year of existence (Takahata, 1991, p. 101; Maruhashi, 1992, p. 49), and at Ryozenyama, Shiga Prefec- ture, a group comprised of four individuals, which was observed shortly after fission of a larger group, was not confirmed to be stable (Sugiyama & Ohsawa, 1982a, p. 32). Provisioned groups tend to become larger than wild groups because favorable nutritional condi- tions result in high birthrates and low infant mor- tality rates. The maximum group size reported in a provisioned group — at Takasakiyama, Oita Pre- fecture— is 1255 individuals (Sugiyama & Ohsa- wa, 1988). Groups in M. fuscata are multimale and mul- tifemale (Itani, 1954, p. 273; Takasaki & Masui, 1984, p. 310). Mean composition in 35 natural groups is ca. 18% adult males, 32% adult females, 35% juveniles, and 15% infants (Table 35). The mean ratio of number of infants to number of adult females is near 0.50, which suggests that biennial parturition is usual (see below. Repro- duction: Birthrate). Group size may change as a result of fission, takeover, or fusion/extinction (Maruhashi, 1992, p. 47). Groups that become very large often split into two or three groups (Koyama, 1970, p. 366; Maruhashi, 1992, p. 49); such fission reportedly occurs between matrilines (Koyama, 1970, p. 377). Conversely, on Yakushima small groups oc- casionally have been observed to fuse with neigh- boring larger groups (Takahata et al., 1994, p. 320). Mass mortality in a cluster of five neighboring troops (65 of 1 17 troop members), including com- plete extinction of two of the five troops, was ob- served in western Yakushima in 1998-1999 (Han- ya et al., 2004, p. 182). FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 51 Table 34. Geographic variation in group size in wild Macaca fuscata populations. Locality (latitude) Group size N Prefecture Mean Minimum Maximum References' Aomori Shimokita. NW (4 1=32 'N) 33 31 34 3 / Shimokita. SW (4r09'N) 29.5 13 46 2 2, 3 Shimokita 24.5 11 42 6 4 Miyagi Narusegawa (38°30'N) 41 — — 1 5 Kinkazan (38'18'N) 29.3 19 55 6 5 Kinkazan (38=1 8'N) 48.7 31 79 6 4 Futakuchi (38=1 6'N) 55 — — 1 5 Tarogawa (38°14'N) 26 — — 1 5 Nagano Shiga Heights (36°42'N) 25.3 15 37 6 4, 6, 7 Toyama Kurobe (36°39'N) 23.8 10 60 18 8 Ishikawa Hakusan (36"16'N) 41 16 86 17 4, 9, 10 Fukui Otoumi (35°32'N) 75 — — 1 11 Shiga Makino-cho (35°30'N) 37.5 26 49 2 4 Ryozenyama (35°2rN) 44 — — 1 12, 13 Hino-cho (35=01 'N) 108.3 83 134 3 4 Kanagawa Hakone-machi (35°11'N) 32 18 46 2 4 Okayama Katsuyama-cho (35°05'N) 112 — — 1 4 Gagyusan (34°49'N) 120 — — 1 4 Chiba Takagoyama (35=10-15'N) 82.5 56 109 2 4, 14 Aichi Shinshiro-city (34°54'N) 41.8 28 47 6 15 Osaka Minoo-city (34°5rN) 51.5 17 86 2 4 Hiroshima Taishakukyo (34°52'N) 29 — — 1 4 Kagawa Shodoshima(34°3rN) 114 52 161 3 4 Mie Nansei-cho (34=20'N) 42 — — 1 16 Fukuoka Kawaradake (33=28'N) 99.5 97 102 1 4, 17 Miyazaki Kojima(3r27'N) 22 — — 1 18 Toi-misaki (31°22'N) 76 — — 1 19 Kagoshima Yakushima (30=20'N) 28.5 11 57 24 4, 20. 21 ' Key to references: 1. Azuma and Ashizawa (1980). 2. Izawa and Nishida (1963). 3. Ashizawa (1979). 4. Yamagiwa and Hill (1998). 5. Izawa et al. (2003). 6. Suzuki et al. (1975). 7. Wada and Ichiki (1980). 8. Akaza (1988). 9. Hayashi (1970). 10. Kawai et al. (1970). 11. Watanabe (1978). 12. Sugiyama and Ohsawa (1974). 13. Sugiyama and Ohsawa (1982b). 14. Iwano et al. (1971). 75. Kawai (1994). 16. Takasaki and Masui (1984). 17. Ikeda et al. (1973). 18. Itani and Tokuda (1958). 19. Azuma (1972). 20. Maruhashi (1980). 21. Maruhashi (1982). Home Range Area and Group Density Based on a survey of 32 widely distributed M. fuscata groups. Takasaki (1981. p. 278) has shown that home range area in this species is strongly influenced by habitat forest type and is positively correlated with group size (cf. Izumi- yama et al., 2003. p. 471). In evergreen broadleaf forest, home range area is relatively small — vary- ing from 1.4 to 6.4 ha per group member; in de- ciduous broadleaf forest, home range area is much larger — varying from 9 to 79 ha per group mem- ber. Along the margins of home range areas, inter- group contact behavior apparently is variable. Ag- gression between neighboring groups reportedly is frequent on Yakushima, less frequent on Kink- azan. and rare or absent at Hakusan (Izawa, 1982, p. 270; Saito et al., 1998, p. 309). Group density on western Yakushima has been estimated as 0.70 ± 0.43 groups/km- in undis- turbed evergreen broadleaf forest and 1.48 ± 0.61 groups/km- in disturbed evergreen broadleaf for- est (Hanya et al.. 2003b. p. 50). Diet Forest fruits, seeds, nuts, leaves, flowers, shoots, buds, and bark generally constitute the pri- mary natural foods of M. fuscata (Murata & Ha- zama. 1968. pp. 5-48; Uehara, 1975, p. 395; 1977, p. 212; Maruhashi, 1980, p. 155; Iwamoto, 1982, p. 167; Koganezawa, 1983, p. 85; Watanuki & Nakayama. 1993. p. 421; Agetsuma & Naka- gawa. 1998. p. 279; Hanya, 2003, p. 336; Hanya et al.. 2003a. p. 54). Other known, relatively mi- nor dietary components include roots, grasses, herbs, fungi, insects, crabs, spiders, mollusks, fish, and bird's eggs (Suzuki, 1965, p. 46; Watan- 52 FIELDIANA: ZOOLOGY Table 35. Group composition and ratios in wild Macaca fuscata (Takasaki & Masui, 1984). Group composition Ratine Group Adult males Adult females Juveniles Infants Totals Locality AdM/AdF Inf/AdF Shimokita M 5 13 13 5 36 0.385 0.385 Z 7 18 10 7 42 0.389 0.389 O 4 4 3 2 13 1.000 0.500 A2 6 16 21 3 46 0.375 0.188 Shiga B2 8 6 4 1 19 1.333 0.167 YBl 6 19 15 5 45 0.316 0.263 Hakusan KA 9 16 14 6 45 0.563 0.375 TA 20 22 14 8 64 0.909 0.364 TB 10 20 12 4 46 0.500 0.200 O 12 18 16 7 53 0.667 0.389 Otoumi A2 6 23 35 11 75 0.261 0.478 Takagoyama T-Ib 6 15 28 7 56 0.400 0.467 Yugawara-machi T 13 19 12 2 46 0.684 0.105 P 3 10 3 2 18 0.300 0.200 Minoo-city A 6 22 41 17 86 0.273 0.773 B 1 5 8 3 17 0.200 0.600 Gagyusan — 10 35 50 25 120 0.286 0.714 Shodoshima I 48 37 35 9 129 1.297 0.243 KA 17 41 72 31 161 0.415 0.756 O 13 17 13 9 52 0.765 0.529 T 11 15 17 9 52 0.733 0.600 Gokashowan — 4 13 16 9 42 0.308 0.692 Kawaradake — 7 33 41 21 102 0.212 0.636 Kojima — 4 6 9 3 22 0.667 0.500 Yakushima Kojiba' 12 18 11 6 47 0.667 0.333 Nagata-todai^ 7 7 9 4 27 1.000 0.571 Kannonzaki 4 4 3 2 13 1.000 0.500 Hanyama 12 12 15 6 45 1.000 0.500 Nina 7 10 5 3 25 0.700 0.300 Kawara 4 6 5 2 17 0.667 0.333 Shikamizawa 8 8 9 3 28 1.000 0.375 M 5 7 6 4 22 0.714 0.571 A 7 8 11 5 31 0.875 0.625 H 3 3 3 2 11 1.000 0.667 M 7 8 8 4 27 0.875 0.500 Totals 35 312 534 587 247 1680 — — Means — 8.9 15.3 16.8 7.1 48 0.650 0.451 Group proportions — 0.186 0.318 0.349 0.147 — — — ' Ko troop. ' = Todai. abe, 1989, p. 126; Aimi & Takahata, 1994, p. 148; Hanya, 2004, p. 61). Monkeys on Yakushima have been observed eating frogs and lizards (Su- zuki et al., 1990, p. 422), and on Kojima, an 11- year-old solitary male was observed to eat a dead shrike (Lanius bucephalus) (Kimura, 1974, p. 25). In an upland mixed coniferous-broadleaf forest (1000-1200 m) on Yakushima, mature leaves of broadleaf trees were a major dietary component, accounting for 38% of observed feeding time (Hanya, 2004a, p. 61); consumption of leaves at this locality apparendy increased substantially fol- lowing a season of unusually poor fruit produc- tion (Hanya et al., 2004, p. 184). Geophagy has been observed on Yakushima (Maruhashi, 1980, p. 148), on Kojima (Iwamoto, 1982, p. 157), and at Arashiyama (Inoue, 1987, p. 104); the func- tional significance of geophagy is unclear, but it may serve as a buffer against gastric disorders (Inoue, 1987, p. 104; Wakibara et al., 2001, p. 515). Feeding occupied 38% of daytime hours in a troop studies in western Yakushima (Hanya, 2004b, p. 171). A 5-year-old M. fuscata female at Kanbataki Natural Park, Okayama Prefecture, reportedly ate her dead infant a few days after the infant had FOODEN AND AlMl: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 53 died (Itoigawa, 1964, p. 13). On Yakushima, sika deer {Cervus nippon) frequently congregate under trees where monkeys are foraging in order to feed on leaves, seeds, or fruit that are dropped or dis- lodged by the monkeys (Majolo & Ventura, 2004, p. 35). Predators Predation on M. fuscata by mountain hawk-ea- gles, Spizaetus nipalensis, has been reported in Hiroshima Prefecture and Nara Prefecture (Ha- mada & lida, 1999, p. 18; lida, 1999, p. 125); in the Hiroshima Prefecture attack, which occurred on 13 July 1997, the victim was a female monkey estimated to be 7 or 8 years old and to weigh 9.4 kg. Feral dogs formerly were abundant in Japan and frequently preyed on monkeys (Miyamoto, 1976, p. 7), but feral dogs are now rare because they have been severely controlled by government action since the 1970s (Mizuno, 1995, p. 54). Raccoon dogs {Nyctereutes procyonoides) may have preyed on M. fuscata infants on Kojima in 1964 (Iwamoto, 1974, p. 257). The Japanese wolf, Canis lupus hodophilax, which became extinct in the early 1900s, may also have been a former predator on M. fuscata. Since 1998, more than 10,000 Japanese mon- keys per year have been exterminated as agricul- tural pests (Oi, 2003, p. 31). Intergroup Dispersal Like other macaques, M. fuscata lives in ma- trilineal, female bonded groups (Itani, 1985, p. 602). Females remain in their natal groups, except in cases of group fission (cf. Fukuda, 2004, p. 56). Males over 5 years of age emigrate from their natal groups and immigrate into neighboring or distant groups (Norikoshi & Koyama, 1975, p. 48; Sprague et al., 1998, p. 353); male migrations of up to 60-100 km have been reported (Fukuda, 1982, p. 494; 1988, p. 479; Yoshimi & Takasaki, 2003, p. 71). Because spermatogenesis in M. fus- cata begins ca. age 4-6 years (Nigi, 1975a, p. 51; Nigi et al., 1980, p. 237; see below. Reproduc- tion), close inbreeding is avoided. Group tenure of males varies from 0.2 to 5.3 years (mean = 2.9 years) on Kinkazan, from 0.2 to 5.7 years at Arashiyama (Norikoshi & Koyama, 1975, p. 55), and from 1.0 to 9.5 years (mean = 2.8 years) on Yakushima (Sprague et al., 1998, p. 352). Reproduction Seasonality Reproduction is strongly seasonal in M. fusca- ta, with matings restricted to autumn and winter and births restricted to spring and summer (Kawai et al., 1967, p. 40; Nozaki et al., 1992, p. 301). Within the spring-summer period, the timing of births apparently varies geographically (Fooden & Aimi, 2003, p. Ill), and this presumably also ap- plies to the timing of matings within the autumn- winter period. In in-situ troops that inhabit the lat- itudinal zone between ca. 31°20'N and 38°20'N, the mean birth date tends to become earlier as latitude increases (Figs. 21, 22); this pattern of variation may function to provide more time for infants born at higher latitudes to achieve an ad- equate level of development before the onset of subsequent poor winter feeding conditions, which generally begin earlier at higher latitudes than at lower latitudes. The above relationship between mean birth date and latitude apparently does not apply to troops that inhabit localities at the geographic and eco- logical extremes of the distribution of M. fusca- ta— namely, Yakushima, ca. 30°20'N, and Shi- mokita Peninsula, ca. 4riO'N (Fig. 22); on Yak- ushima, the mean birth date is earlier than ex- pected, and on Shimokita Peninsula, it is later than expected. The mild winters on Yakushima may be less of a factor in infant survival than elsewhere in Japan. Conversely, winters on Shi- mokita Peninsula last so long that earlier births might result in infants being born before anteced- ent poor winter feeding conditions have ended. Troops that have been translocated to new lat- itudes in the northern hemisphere generally retain approximately the same mean birth date that they manifested at their original localities (Fooden & Aimi, 2003, p. 111). Contrastingly, in one group that was translocated to the southern hemisphere (Tasmania), the mean birth date shifted approxi- mately 6 months forward; this presumably was in response to the reversed seasons. Laboratory cap- tives kept under conditions of constant artificial day length and temperature retained their original ovulatory and birth cycles for at least 12 years (Nozaki, 1991, p. 104; Nozaki et al., 1992, p. 54 FIELDIANA: ZOOLOGY In-situ troops Translocated troops 130° 135' 140*' Fig. 21. Distribution of Macaca fuscata troops in Japan for which birth seasonality data are available (for details, see Fooden & Aimi, 2003, p. 1 15). In-situ troops. Localities with complete birth seasonality data: 1. Yakushima: Nina-A. 2. Toimisaki; 3. Kojima; 4. Takasakiyama; 5. Nametoko; 7. Kochi; 8. Taishakukyo; 9. Gagyusan; 10. Kanbanotaki; 13. Funakoshiyama; 14. Rosando; 15. Choshikei; 17. Minoo-city-A; 18. Arashiyama; 23. Hagachi; 25. Takagoyama-S; 26. Takagoyama-A; 29. Jigokudani; 30. Kinkazan A; 31. Shimokita Peninsula: Wakinosawa. Localities with incomplete birth seasonality data (not plotted in Fig. 21): 11. Katsuyama-cho; 16. Kaminada; 19. Otoumi; 20. Ryozen-A; 24. Hakone-machi. Translocated troops. 6. Miyajima-cho; 12. Takeno-cho; 21. Ohirayama; 22. Okinoshima; 27. Takaosan; 28. Hou- tosan. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 55 1 1 1 1 1 1 1 1 ■po das 6nv inp unp Ab|^ Jdy JBI/^ qej Inside Toimisaki- Outside Toimisaki- Kinkazan Kinkazan latitudinal zone latitudinal zone Earliest birth date • o Latest birth date ■ d Mean birth date ♦ O ■ A . A ■ * / • _ ■ ^ 1 m m. ^ 1 . ■ I » ^^^ ^n^ . 1 ■ iV • ▼ / • _ ■ — • / * ^ — _ m ■ ■ ^ 1 . 7^ ■ A. / A CM -"it O CO CO CO co ^^ Z M 0> CD CO 2. a> XJ 3 (S in CO CO CO CO CM CO O O CO o 00 CNI O CD CNI O CM O CNJ CM O O CM O CO O CO O CO O O CNI O O O O OO CD O •ou Abq 56 FIELDIANA: ZOOLOGY 302). Experimental manipulation of photoperiod had little or no effect on ovulatory cycles of cap- tives, but simultaneous manipulation of photope- riod and temperature resulted in reversed cycles in two of four experimental monkeys. Sexual Maturation Although M. fuscata males as young as age 1 .5 years have been observed to mount females (Tak- ahata, 1980, p. 325; Eaton et al., 1986, p. 144; Mehlman & Chapais, 1988, p. 202), ejaculation in this species normally does not occur until age 4.5 years (Hanby et al., 1971, p. 125; Wolfe, 1978, p. 62; Takahata, 1980, p. 325; Inoue et al., 1991, p. 203); one exceptional case of ejaculation at age ca. 3.5 years has been reported by Chapais and Mignault (1991, p. 173). Descent of the testes into the scrotum in various troops of free-ranging M. fuscata has been reported to occur at age ca. 4.5 years (Hiraiwa, 1981, p. 315), at age ca. 5.5 years (Nigi et al., 1989, p. 572), or even after age 6 years (Mori, 1979b, p. 381); Nigi et al. (1989, p. 574) have suggested that sexual maturation in males may be accelerated by artificial feeding. Histologically, sperm are rarely detected in the testes of monkeys at age ca. 3.5 years (Inoue & Hayama, 1963, p. 76; Nigi, 1975a, p. 51; Matsu- bayashi & Mochizuki, 1982, p. 895), but they are commonly seen in males at age 4.5 years (Nigi, 1975a, p. 51; Nigi et al., 1980, p. 233; Matsubay- ashi & Mochizuki, 1982, p. 897). By age 5 years, almost all males have emigrated from their natal troops (Sugawara, 1975, p. 354; Hasegawa & Hir- aiwa, 1980, p. 136; Fukuda, 1982, p. 493; Sugi- yama & Ohsawa, 1982b, p. 255; Matsumura, 1993; Sprague et al., 1998, p. 353; Yoshimi & Takasaki, 2003, p. 72) (see above. Natural His- tory: Intergroup Dispersal). Full sociosexual ma- turity does not occur until age >8.5 years, when the frequency of mating activity is approximately five times as great as at younger ages (Takahata, 1980, p. 325); by this age, males have developed mature or near-mature sexual skin coloration (Ma- sui et al., 1973, p. 237; Mori, 1979b, p. 381; Hir- aiwa, 1981, p. 315; Maruhashi, 1982, p. 320). Menarche (first menstruation) in M. fuscata fe- males usually occurs at age ca. 3.5 years (Nigi, 1975b, p. 208; Mori, 1979b, p. 381; Hasegawa «& Hiraiwa, 1980, p. 136; Hamada et al., 1999, pp. 441, 449; cf. Hazama, 1964, p. 89). At about the same age, characteristic para-anal pubertal swell- ings become evident (Wolfe, 1979a, p. 412; Tak- ahata, 1980, p. 304; Maruhashi, 1982, p. 320; Oi, 1988, p. 13; cf. Pocock, 1906, p. 558; Mori et al., 1997, p. 562); these are succeeded by subcaudal or paravulval pubertal swellings during the mating seasons of the next 2-3 years. Females typically begin to display estrous behavior at age ca. 3.5 years or ca. 4.5 years (Wolfe, 1979a, p. 412; Hir- aiwa, 1981, p. 311; Chapais & Mignault, 1991, p. 173; Pavelka, 1993, p. 70). In provisioned troops, females generally give birth to their first infants at age ca. 5 years or age ca. 6 years (cf. Fedigan, 1991b, p. 145; Itoigawa et al., 1992, p. 57; Koya- ma et al., 1992, p. 37); known extreme ages of primiparous females are ca. 4 years and ca. 16 years (Watanabe et al., 1992, p. 8). Provisioning of troops apparently tends to reduce the mean age of primiparity of troop members (Sugiyama & Ohsawa, 1982a, p. 37; Fukuda, 1988, p. 481; Watanabe et al., 1992, p. 8). Sexual Skin During the mating season, the bare skin on the face, nipples, and perineum of M. fuscata be- comes intensely reddish in adults and, to a lesser degree, also in adolescents. In females, this red- dening varies cyclically during the mating season, increasing during estrus intervals and decreasing during interestrus intervals (Tokuda, 1963, p. 14; Nigi, 1975b, p. 21 1; Wolfe, 1984c, p. 146; Taka- hata et al., 1995, p. 170); estrus intervals are also marked by vaginal discharge of a distinctively odoriferous secretion (Enomoto, 1974, p. 353; Takahata et al., 1995, p. 170). Unlike adolescent females (see above, Sexual Maturation), adult fe- FiG. 22. Latitudinal variation in birth seasonality in in-situ troops oi Macaca fuscata (for documentation, see Fooden & Aimi, 2003, p. 1 15). Seasonality in 18 troops distributed within the Toimisaki-Kinkazan latitudinal zone (ca. 3r20'- 38°20'N) is compared with seasonality in troops on Yakushima (Nina-A, 30°2rN) and Shimokita Peninsula (Waki- nosawa, 41°08'N). Regression equation: y = -10.07x + 493.4; r = -0.764, F = 16, P < O.OI. (Note: In 2003, data for which are not included in this graph, the mean date of 16 births at Wakinosawa was 30 April, and extreme birth dates were 10 April and 28 May [S. Matsuoka, Working Group for Monkeys in the Shimokita Peninsula, pers. comm.]). FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 57 Table 36. Menstrual cycle length in Macaca fuscata samples (mean ~ 27.1 days, n ~ 593 cycles). Publication date Mean cych length (days) i Standard Extreme deviation cycle lengths Number of (days) (days) females Number of cycles Reference' 1980 26.52 Arashiyama-B: free-ranging provisioned troop 9.1 8-53' — 73 ; 1989 27.5 Ehime University 7.8 19-47 26 94 2 1975 1977 1977 Japan Monkey Centre (mean ~ 26.2 days, n ~ 315 cycles) 26.3 5.4 < 15-35^ 10 -275 29.5 1.7 28-32 4 4 25.3 2.8 19-31 — 36 3 4 5 1977 1992 Primate Research Institute, Kyoto University (mean = 30.2 days, n 28.5 3.0 23-36 7 31.7 3.2"^ — — = 76 cycles) 36 40 6 7 1979 28.1 Tokai University 4.2 21-40 6 35 8 ' Key to references: 7. Takahata (1980, p. 311). 2. Hamada et al. (1989, p. 188). 3. Nigi (1975b, p. 210). 4. Ando et al. (1977, p. 274). 5. Nigi (1977, p. 246). 6. Oshima et al. (1977, p. 101; cf. Aso et al., 1977, p. 747). 7. Nozaki et al. (1992, p. 304). 8. Enomoto et al. (1979, p. 564). ' Mode = 26 days. -^ Excludes cycles >53 days; intervening bleeding possibly missed. '• Excludes 14 cycles, duration >40 days. ^ Standard error = 0.5. males do not exhibit perineal swellings (Nigi, 1975b, p. 211; Wolfe, 1984c, p. 146), although they do undergo cyclical reddening of the peri- neum. In ovariectomized females, reddening of the sexual skin can be induced by injection of estrogen (Nigi & Ando, 1975, p. 266). In adult males, the intensely red coloration of the sexual skin — including the face, perianal area, and scrotum — persists more or less continuously throughout the mating season (Alexander, 1970, p. 279; Enomoto, 1974, p. 353; Wolfe, 1979a, p. 413; Maruhashi, 1982, p. 319). Concurrently dur- ing the mating season, adult males carry their short tails erect, which increases the visibility of the bright red perineal sexual skin (Wolfe, 1981a, p. 26). Near the beginning of the mating season, increased redness of the sexual skin and seasonal erection of the tail are known to be associated with increased testosterone levels (Rostal et al., 1986, p. 459). Menstrual Cycle During the mating season, menstrual cycle length averages ca. 27.1 days (Table 36), and the duration of menstrual flow averages 3-5 days (Nigi, 1975b, p. 210; Aso et al., 1977, p. 747; Oshima et al., 1977, p. 101; Hamada et al., 1989, p. 188). A female's peak estradiol value, which is followed approximately 1 day later by ovulation, occurs near the middle of the menstrual cycle (Nomura et al., 1972, p. 475; Ando et al., 1976, p. 92; Ando et al., 1977, p. 274; Nigi & Torii, 1983, p. 413) During the nonmating season, men- strual cycle length becomes irregular, and men- struation may be completely suppressed (Tokuda, 1963, p. 11; Nomura et al., 1972, p. 474; Nigi, 1975b, p. 210; Hamada et al.. 1989, p. 188). Estrus Available information indicates that, during the mating season, a mature M. fuscata female will exhibit 0-6 (mean, ca. 2) discrete periods of sex- ual proceptivity and receptivity — i.e., estrus (Ta- ble 37). The known duration of these estrous pe- riods, which are not synchronized among female troop members, varies from 1 to 92 days (mean, ca. 10 days); the intervals between estrous periods also are highly variable (Wolfe, 1984c, p. 150; Pavelka, 1993, p. 96). In a recent study of the relationship between hormonal cycles and sexual behavior in a seminatural population of M. fus- cata, female attractivity and proceptivity were 58 FIELDIANA: ZOOLOGY Table 37. Frequency and duration of estrous periods in mature Macaca fuscata females. Number of estrous periods Estrous period duration (days) Observation . per female per matmg season Refer- Locality period Mean ± SD Extremes N Mean ± SD Extremes N ences' Arashiyama 1975-77 2.6 ± 1.5 0-6 87 13.6 ± 12.2 1-92 87 1 Arashiyama 1977-78 2.8 ± 1.5 — 71 -8.5 — 198 2 Arashiyama 1979-85 1.9 ± 0.9 1-4 156 8.9 ± 8.4 1-42 156 3 Kojima 1950s — — — 9.3 ± 3.1 6-13 26 4 Shiga A 1967-73 -1.7 1-5 -77 -10.7 1—75 -264 5 South Texas- 1973-74 2.5 ± 1.1 — — 8.3 ± 9.8 — — 6 ' Key to references: 7. Takahata (1980, pp. 311, 314). 2. Wolfe (1984c, p. 151). 3. Huffman (1991b, p. 104). 4. Tokuda (1961-62. p. 13). 5. Tokita and Hara (1975, p. 30). 6. Wolfe (1979b, p. 527). - Translocated from Arashiyama. found to increase during the follicular and peri- ovulatory phases of the hormonal cycles (O'Neill et al., 2004b, p. 359). At Arashiyama, estrus in females with infants (born ca. 6 months before the mating season) was delayed by approximately 1 month relative to es- trus in females without infants (Takahata, 1980, p. 317), and on Kinkazan, estrus was rare in fe- males with infants (one estrous season in 19 fe- male-years) (Takahashi, 2002, p. 145). In the Shi- ga A troop, most females reportedly exhibited clear estrus only in alternate years (Enomoto, 1975. p. 275; 1978, p. 284). On Kinkazan, fruit abundance during the mat- ing season strongly affected the observed estrus rate (Takahashi, 2002, p. 148). At Arashiyama, dominance rank influenced estrus duration; the to- tal annual duration of estrus in high-ranking fe- males (mean = 58.8 days, n = 22) was approxi- mately twice as great as that in lower-ranking fe- males (mean = 28.2 days, n = 32) (Takahata, 1980, p. 324). Most conceptions in M. fuscata occur during a female's first estrous period of the mating season (Takahata, 1980, p. 311; Huffman, 1993, p. 80; Maruhashi & Takasaki, 1996, p. 154). However, even after conception, females usually exhibit one or more additional estrous periods — with postcon- ception copulations — during the remainder of the mating season (Hanby et al., 1971, p. 137; Taka- hata, 1980, p. 311; 1999, p. 148; Wolfe, 1979b, p. 527; 1984c, p. 151; Huffman, 1993, p. 80). Male sexual activity in M. fuscata is more or less continual during the mating season and does not exhibit estrus-like periodicity. Two cases of unseasonably late estrous periods followed by un- seasonably late births suggest that the mating ac- tivity of males may be induced by the presence of estrous females (Kawai, 1966, p. 392). Consortship During the mating season, males and estrous females in natural and seminatural groups form consortships — temporary associations of varying duration in the course of which copulations usu- ally occur (Imanishi, 1963, p. 77; Itani, 1963, p. 51; Eaton, 1976, p. 103; D'Amato et al., 1982, p. 228; Gouzoules & Goy, 1983, p. 41; Yamagiwa, 1985, p. 114; Huffman, 1991c, p. 200; cf. Perloe, 1992, p. 296). Consortships may be initiated by either sex (Itani, 1963, p. 51; Tokuda, 1963, p. 16; Stephenson, 1973, pp. 67-71; 1975, p. 74; Wolfe, 1979b, p. 527; 1984c, p. 148; Xiong & Wang, 1991, p. 14; Soltis et al., 1997b, p. 733; 1997a, p. 744; 1999, p. 272); however, females and males that form consortship pairs usually are not individuals that are closely affiliated during the nonmating season (Enomoto, 1975, p. 278; Fedigan & Gouzoules, 1978, p. 493; Baxter & Fedigan, 1979, p. 451; Takahata, 1982b, p. 100). In the Arashiyama B troop, the mean (±SD) du- ration of 168 uninterrupted consortships was 1.6 ±1.1 days (extremes, 1-7 days) (Huffman, 1993, p. 80). Troop females often form consortships with nontroop solitary males, which tend to approach troops during the mating season (Nishida, 1966, p. 171; Kawai et al., 1968, p. 10; Kawanaka, 1973, p. 149; numerous subsequent authors). Dominant males frequently attempt to disrupt the consortships of subordinate males (Tokuda, 1963, p. 29; Furuichi, 1985, p. 230; Huffman, 1987, pp. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 59 Table 38. Reported results concerning relationship between male dominance rank and reproductive success in natural and seminatural groups of Macaca fitscata. Reports that variables are positively correlated Reports that variables are not positively correlated -male dominance rank and copulation success Enomoto (1974. p. 367) Stephenson (1975. p. 92) Takahata (1982b. p. 105) Xiong and Wang (1991. p 16) Natural groups: Variables Nishida(1966. p. 169) Stephenson (1975. p. 85) Huffman (1987. p. 231) Soltis et al. (2001. p. 488) Matsubara (2003. p. 1062) Natural groups: Variables — male dominance rank and fertilization success Soltis et al. (2001. p. 488) Huffman (1987. p. 231 ) Seminatural groups: Variables — male dominance rank and copulation success Eaton (1971, p. 9) Eaton (1974. p. 292) Hanbv et al. (1971. p. 132) Soltis et al. (1997a. p. 741) Stephenson (1975. p. 98) Inoue et al. (1991. p. 204) Seminatural groups: Variables — male dominance rank and fertilization success Inoue et al. (1990. p. 567) Inoue et al. (1991. p. 204) Soltis et al. (1997a. p. 742) 223-224: 1991b. p. 106: 1992, p. 83): presumably as a consequence, consortships involving domi- nant males usually occur near the center of a troop, and those involving lower-ranking males, or nontroop males, usually occur at or beyond the periphery of a troop (Imanishi. 1963, p. 77: Fu- ruichi. 1985. p. 230: Sprague. 1994. p. 185). Each sexually mature male or female is likely to have more than one consortship partner during the course of a single mating season (Tokuda. 1963. p. 36: Eaton, 1976. p. 103: Wolfe. 1986. p. 271: Inoue et al.. 1993. p. 507: Soltis et al.. 1999. p. 270: 2001. p. 489: Matsubara & Sprague. 2004. p. 904): in a seminatural group that included 79 sexually mature females, the mean (±SD) number of male partners per female during one mating season was 4.78 ± 3.68 (extremes. 1-15) (Gou- zoules & Goy. 1983. p. 41). Polyandry may ben- efit females by reducing infant harassment and in- fanticide (Soltis et al.. 2000. p. 197: Soltis. 2002. p. 192: see below. Infant Mortality). Copulatory Behavior M. fuscata generally is a multimount ejacula- tor — i.e.. each ejaculatory copulation usually con- sists of a series of mounts separated by brief (ca. 30-second) dismounts (Tokuda. 1963, p. 28: Han- by et al.. 1971, p. 127: Hanby & Brown. 1974, p. 162: Enomoto, 1974, p. 363: Stephenson, 1975. p. 74: Wolfe. 1978, p. 60: Xiong & Wang. 1991. p. 14: Pavelka. 1993. p. 101: Troisi & Carosi, 1998. p. 1263). The mean number of mounts per copulatory sequence reported in various natural and seminatural troops vaiies from 8.8 to 18.9 (extremes. 1-59), and the mean duration of cop- ulatory sequences varies from 6.2 to 13.0 minutes (extremes. l->60 minutes): the mean number of intromissive thrusts per mount vaines from 1.7 to 3.7. As in M. nmlatta. a M. fitscata male usually mounts a female dorsoventrally by gripping the female's waist and shanks with his hands and feet, respectively (Tokuda. 1963. p. 26: Hanby et al., 1971, p. 126: Hanby & Brown. 1974, p. 156: Wolfe, 1978, p. 58). Copulation occurs both in trees and on the ground (Yotsumoto. 1976. p. 202). During the daytime, copulation is most fre- quently observed early in the morning (Hanby et al.. 1971, pp. 127-131: Oda & Masataka, 1995, p. 133: Vasey, 1998b, p. 583). but optically as- sisted night viewing and fresh copulatory plugs observed at dawn indicate that copulations also occur at night (Inoue et al., 1991, p. 206: Soltis et al.. 1997a, p. 744). Dominance Rank and Reproductive Success Equivocal results have been reported concerning the relationship between male dominance rank and reproductive success in M. fitscata (Table 38). This may indicate variation in this relationship among 60 FIELDIANA: ZOOLOGY the groups studied and/or variation in the meth- odology of observers (cf. Takahashi, 2004, p. 101). In females, dominance rank and birthrate were found to be positively correlated in two studies (see below. Birthrate) and were found to be uncorrelat- ed in four studies. In a small captive group of M. fuscata, the dominant female successfully restricted the copulation and fertilization of a subordinate fe- male (Rendall & Taylor, 1991, pp. 323, 325). Inbreeding In natural populations of M. fuscata, the op- portunity for close inbreeding is minimized by the strong tendency for young males to leave their natal troops before they reach full sociosexual maturity (see above. Sexual Maturation). Some adolescent males that have not yet emigrated may exhibit positive sexual behavior toward closely re- lated females, but these advances generally are rebuffed by the females (Enomoto, 1974, p. 369). Aunt-nephew matings are the most closely inbred pairings with ejaculatory copulations that have been reported in natural troops (Koyama, 1970, p. 357; Enomoto, 1978, p. 290; cf. Imanishi, 1961, p. 119; Takahata et al., 2002, p. 406). Although male emigration is impossible in con- fined seminatural populations, close matrilineal inbreeding also is rare in such groups (Eaton, 1971, p. 9; Hanby & Brown, 1974, p. 163; Wolfe, 1977, p. 168; Baxter & Fedigan, 1979, p. 454; Pavelka, 1993, p. 109; Soltis et al., 1997a, p. 739); father-daughter matings and matings between pa- ternal half siblings have been observed in semi- natural groups (Inoue et al., 1990, p. 568; 1992, p. 134). As in natural troops, rejection by females probably is a major factor in the inhibition of ma- trilineal inbreeding in seminatural groups (Soltis et al., 1999, p. 270). Homosexual behavior ap- parently also is rare between close matrilineal rel- atives, at least in seminatural groups (Fedigan & Gouzoules, 1978, p. 493; Mehlman & Chapais, 1988, p. 202; also see below, Nonreproductive Sexual Behavior). No serial mounts were observed between a M. fuscata mother and her adult son that were ex- perimentally caged together during three mating seasons (Itoigawa et al., 1981, p. 502). Hybridization Breeding between M. fuscata and other ma- caque species has occurred both in captivity and under free-ranging conditions. In zoos, M. fuscata reportedly has hybridized with M. cyclopis and M. silenus (International Zoo Yearbook, 1963, p. 224; 1 97 1, p. 265). Under free-ranging conditions, hy- bridization with an introduced population of M. mulatta has been observed, and hybridization with an introduced population of M. cyclopis has been inferred (see below). Judging from available evi- dence, females and males in free-ranging M. fus- cata troops spontaneously interbreed with other macaque species when given the opportunity. At Arashiyama, near Kyoto, two estrous fe- males— members of a local free-ranging troop of M. fuscata — climbed over an electrified fence that enclosed a small imported group of M. mulatta, copulated with M. mulatta males, climbed out of the enclosure, and subsequently gave birth to hy- brid infants (Wolfe, 1981b, p. 131). One of the hybrid infants, a female bom in 1968, was per- mitted to remain in the M. fuscata troop; when she reached adulthood, her size and dorsal pelage color apparently were intermediate between that in M. fuscata and M. mulatta, her tail length ap- parently was similar to that in M. mulatta (cf. Fooden, 2000, p. 30), and her estrous perineal sexual skin apparently was similar to that in M. fuscata. In 1972 and 1973, this hybrid gave birth to two daughters; as adults, the daughters (0.75 M. fuscata, 0.25 M. mulatta) resembled their mother in size and dorsal pelage color, but their tail length was intermediate between that of their mother and that of M. fuscata. In Wakayama Prefecture, south of Osaka, an imported group of M. cyclopis escaped from a lo- cal menagerie in 1955 and became established in a nearby forest south of Ooike village (Kawamoto et al., 1999, p. 53; 2001, p. 13). A recent study of blood proteins and mtDNA in the Ooike pop- ulation indicates that M. fuscata males from troops in adjacent areas have freely hybridized with females in the introduced M. cyclopis pop- ulation; approximately 25% of the current Ooike population gene pool is estimated to be of M. fus- cata origin. The tail length reported in a young male F, hybrid (age 4-5 years) is 290 mm, which is approximately intermediate between tail length in M. fuscata and that in M. cyclopis (see above. External Measurements and Proportions; Fooden & Wu, 2001, p. 13). At the southern tip of Boso Peninsula, Chiba Prefecture, M. fuscata males apparently have hy- bridized with introduced individuals or groups of M. fascicularis, M. cyclopis, and M. mulatta (Ai- zawa & Hagiwara, 2001, p. 4). FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 61 Table 39. References' concerning nonreproductive sexual behavior reported in natural and seminatural groups of Macaca fuscata. Nonreproductive mounts- Masturbation Locality ;/ ^ -j; z/z ^ ' Natural groups Arashiyama 7.2.. ^,4 5 6 3 Kojima •) 1,5 7 Minoo-ciiy 2 8 Miyajima-cho" 5 Shiga 9 9 9 9 9 Takasakiyama 9 9 9 9 9. 10 Yakushima 11 Seminatural groups Arashiyama West. U.S. .A. 12, 13. 14 14. 15 12 15 Beaverton. U.S. .A. 16,17 16 16, 17, 18 16.19 Calgary Zoo. Canada 20 20 Ca\ riglia Park. Italy 21 St. Hyacinthe. Canada^ 22 Universite de Montreal, Canada 23,24 23 ' Ke\ to references: /. Takahata (1980. pp. 306. 315. 323). 2. Takahata (1982b. p. 104). 3. Wolfe (1984c. pp. 150, 151). 4. Wolfe (1986. pp. 268. 271 ). 5. Stephenson (1973. pp. 63. 66. 70). 6. Takenoshita (1998. p. 365). 7. Stephenson (1975. p. 92). 8. Perloe (1992. p. 300). 9. Eonomoto (1974. pp. 354. 364). 10. Itani (1959. p. 86). //. Thomsen (2000. p. 213); Soltis et al. (2001. p. 487): Domingo-Roura et al. (2004. p. 34). 12. Fedigan and Gouzoules (1978, p. 493). 13. Baxter and Fedisan (1979. p. 454): Wolfe (1979b. pp. 526. 531: 1984a, p. 23^5); Yoshida (1988. p. 58); Pavelka (1993. p. 108). 14. Gouzoules and Gov (1983. p. 42); O'Neill et al. (2004a. p. 29). 15 Wolfe (1978, pp. 58, 63). 16. Hanby et al. (1971. pp. 126. 132. 133. 137). 17. Eaton (1978, p. 49). 18. Hanbv (1974. pp. 838. 843). 19. .■\lexander (1970. p. 279). 20. Rendall and Tavlor (1991, p. 324). 21. Lunardini (1989. p.'l83); Corradino (1990, pp. 356. 358). 22. Vasey (1996, pp. 543. 550; 1998b. pp. 582. 585. 590): Chapais et al. (1997. p. 1096); Vasev et al. (1998, p. 392); Vasev and Gauthier (2000. pp. 20, 22). 23. Mehlman and Chapais (988. p. 202). 24. Chapais and Mignauh (1991. pp. 174. 175). - Tabulated references to homosexual mounts pertain exclusively to mount series; references to single-mount events, which usually are not sexually motivated, are excluded. 'This population was translocated from Shodoshima in 1962 (Kawai et al.. 1967. p. 47). - This population was translocated from the Universite de Montreal in 1992 (Chapais, 1986, p. 409; Chapais et al., 1997. p. 1091; Vasey. 1998b. p. 583). Nonreproductive Sexual Behavior Homosexual and autosexual behaviors have been observed in natural and seminatural groups of A/, fuscata (Table 39); these behaviors include female/female mounts, female/male mounts, male/male mounts, and masturbation by females and males. As indicated below, published data sets concerning these behaviors often are frag- mentary or incongruent and therefore are some- v^hat difficult to compare. In natural groups (Table 39), female/female consortships. including reciprocal mounts, have been reported at Arashiyama (47*!^ of 94 sexually mature females in 1976^1977: TT^ of 88 sexually mature females in 1977-1978). Kojima (rarely observed). Minoo. Shiga, and Takasakiyama (no frequency data for the last three localities); at Ara- shiyama. the homosexual estrous p>eriods of a fe- male usually did not overlap with her heterosex- ual estrous periods (ca. 679^ of observed estrous periods). In addition to mounting other females, females also have been observed to mount males during the course of heterosexual consortships; this behavior has been reported at Arashiyama (lO'v^ of 87 observed heterosexual pairs), on Mi- yajima (14% of 49 observed heterosexual pairs), at Shiga and Takasakiyama. but never on Kojima (>58 observed heterosexual pairs). Male/male consortships and ejaculatory mounts, including reciprocal mounts, have been reported at Arashi- yama (six males in two large troops [>300 indi- viduals]). Shiga, and Takasakiyama: at Arashiya- ma. two of six males that participated in homo- sexual mounts also were observed to engage in heterosexual activity during the same mating sea- son. Female masturbation (stimulation of clitoris manually or with inanimate object) has been re- ported at Arashiyama (167r of 111 females). Shi- ga, and Takasakiyama. Male masturbation, includ- 62 FIELDIANA: ZOOLOGY ing masturbation by troop leaders, has been re- ported at Kojima, Shiga, and Takasakiyama and on Yakushima. In seminatural groups (Table 39), at Arashiya- ma West, Calgary Zoo, Cavriglia Park, Universite de Montreal, and St. Hyacinthe, homosexual con- sortships and mounts were reciprocally performed by 51-100% of sexually mature females, whereas at Beaverton the reported mean frequency of fe- male homosexual mounters was only 9.7%; al- most all females that participated in homosexual mounts also participated in heterosexual mounts. At Arashiyama West, O'Neill et al. (2004a, p. 29) have discovered that female homosexual mounts — like heterosexual mounts — are restricted to the follicular and periovulatory phases of the ovarian cycle. Females that were engaged in ho- mosexual consortships have been observed to re- ject the sexual solicitations of intruding males (Vasey, 1998b, p. 590); homosexual consortships apparently are rare or absent between closely re- lated females (Wolfe, 1979b, p. 531; Chapais & Mignault, 1991. p. 175; Chapais et al., 1997, p. 1096; Vasey & Gauthier, 2000, p. 20). During the course of heterosexual consortships, females have been observed to mount males at Arashiyama West (31% of 1 14 copulations in 1973-1974; 42% of 79 sexually active females in 1977-1978), Bea- verton (ca. 40% of 37-58 sexually active fe- males), and Universite de Montreal (25% of eight sexually active females). Male/male mount series have been reported at Arashiyama West (ca. 1% of 702 consortships during three mating seasons) and Beaverton (ca. 15% of 22-59 sexually active males during four mating seasons; extremes, 4- 127 male/male mount series per mating season). Female masturbation in seminatural groups has been reported only at Calgary Zoo (three of three sexually mature females). Male masturbation, in- cluding manual and oral stimulation of penis, has been reported at Arashiyama West (2 of ca. 15 sexually mature males) and Beaverton (13 of 26 sexually active males; 27 masturbations, 12 ejac- ulatory). In individually caged M. fuscata males at PRIKU, the mean daily frequency of mastur- bation during sampled mating-season months of October and December (0.63 masturbations/day) was nearly four times as great as during sampled birth-season months of April and June (0.17 mas- turbations/day) (Matsubayashi, 1974, p. 254). Two tentative generalizations are suggested by the available data: (1) The frequency of some or all of the above nonreproductive sexual behaviors probably varies locally and annually in M. fus- cata. (2) The frequency of female homosexual be- havior in M. fuscata probably exceeds the fre- quency of male homosexual behavior (cf. Vasey, 1995, p. 179). Various interpretations of nonreproductive sex- ual behaviors have been published. Eaton (1978, p. 48) and Wolfe (1979b, p. 532; 1984a, p. 235; 1984c, p. 152; 1986, p. 273) have noted that var- iation in the frequency of female and/or male ho- mosexual behavior apparently is related to varia- tion in the sex ratio of mature troop members; this is interpreted to indicate that homosexual behav- ior in M. fuscata may be a response to an inade- quate number of suitable heterosexual partners (cf. Fedigan & Gouzoules, 1978, p. 494; Rendall & Taylor, 1991, p. 326; Takenoshita, 1998, p. 366; Vasey & Gauthier, 2000, p. 22). Gouzoules and Goy (1983, p. 47) have suggested that mounting by females may be a learned behavior that is de- pendent on parturition or pregnancy status. Vasey (1996, p. 550; 1998a, p. 417; 1998b, p. 595; Va- sey et al., 1998, p. 395; Vasey & Gauthier, 2000, p. 23) and coworkers have proposed that female/ female consortships are motivated by sexual at- traction and gratification and that this behavior persists because it is selectively neutral. Wolfe (1984c, p. 152) has suggested that female masturbation may be a learned behavior that is so- cially transmitted within matrilines, whereas Ren- dall and Taylor (1991, p. 326) indicated that this behavior may be a result of restricted access to a suitable mating partner Thomsen (2000, p. 213) has proposed that male masturbation may function to improve ejaculate quality by removing sperm that have begun to deteriorate; this interpretation would not be applicable to female masturbation. Gestation In a composite sample of 73 M. fuscata preg- nancies, mean duration of the gestation period is 171.7 days (Table 40); although the standard de- viation of this composite mean is indeterminable, extreme durations are 157 and 188.5 days. For infants of known gestation length, sex has been reported for only two females and 10 males (Osh- ima et al., 1977, p. 100; Negayama et al., 1986, p. 367; cf. Table 40); in the unsatisfactory sample of two infant females, mean duration of the ges- tation period is 170.5 days (extremes, 169.5- 171.5 days), and in the more satisfactory sample of 10 infant males, the mean (±SD) duration is 175.4 ± 7.10 days (extremes, 161-184.5 days). More data are required to ascertain whether the mean gestation period of male infants exceeds that of female infants. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 63 Table 40. Gestation period duration in Macaca fuscata. Publi- Gestation period duration (days) Refer- cation date Mean SD Extremes N Basis of conception date estimate ences- 1966 175.5-^ _ _ 1 Estrous period 1 1976 173.0 6.9 161-188 17 48-hour controlled mating period 2 1977 176.9^ 5.5 169.5-184.5 5 Copulatory period 3 1986 173.0^ 7.4 161-179.5 V Unspecified 4 1988 167.4 7.2 — 14 Ovulation'' (n = 6); 11 -day con- trolled mating period (n = 8) 5 1990 162.8 3.6 157-180 6 5-day controlled mating period 6 1992 173.5 1.7 161-183 11 Ovulation^ 7 1998 169.5 — 168-171 2 Artificial insemination 8 2000 166 — — 1 In vitro fertilization'' 9 2003 176.3 7.1 170-188.5 9 Ovulation'* 10 Totals 171.7 — 157-188.5 73 — — ' Excludes two imprecise estimates: Tokuda (1961-62, p. 1 1) — 150-170 days; Hazama (1964, p. 89) — 5-6 months. - Key to references: 7. Kawai (1966, p. 391). 2. Nigi (1976, p. 83). 3. Oshima et al. (1977, p. 100). 4. Negayama et al. (1986, p. 367). 5. Shimizu (1988, p. 252). 6. Nozaki et al. (1990, p. 446). 7. Mitsunaga et al. (1992, p. 29). 8. Torii and Nigi (1998, p. 401). 9. Torii et al. (2000, p. 44). W. Fujita (2003, p. 72). ' Mean based on midpoint(s) of duration range(s) specified in reference. '' Excludes one stillbirth and one imprecise estimate (135-176 days). -'' Determined by hormonal analysis. ^ Embryo subsequently implanted in fallopian tube. During most pregnancies at Primate Research Institute, Kyoto University, implantation bleeding was detected for 3-7 days, beginning ca. 25-30 days after conception (Oshima, 1980, p. 4). At Arashiyama West, Texas, females pregnant with female fetuses reportedly gave and received more aggression than females pregnant with male fe- tuses, particularly during the first 2-3 months of pregnancy (Noyes, 1981, p. 317). Parturition The time of parturition in M. fuscata is best known, as would be expected, in caged popula- tions (Itoigawa & Tanaka, 1963, p. 76; Oshima, 1980, p. 7; Negayama et al., p. 367; Kanazawa & Nakamichi, 1991, p. 424). Of 16 reported partu- ritions in caged females, 12 (including one or two stillbirths) occurred near midnight during the in- terval 2300-0300, three occurred during the in- terval 2000-2200, and one occurred at 0727. In one seminatural troop, a parturition occurred at ca. 1800 (Fedigan & Fedigan, 1977, p. 205), and in two natural troops, parturitions occurred at 1655 and ca. 0745 (Nakamichi et al., 1992, p. 414; Thomsen, 1997, p. 339). Other, less precise reports concerning seminatural and natural troops similarly indicate that parturition in these troops usually occurs at night or during early morning hours (Hazama, 1964, p. 83; Murray & Murdoch, 1977, p. 818; Pavelka, 1993, p. 61; Thomsen & Solis, 2000, p. 690). Shortly before parturition, prepartum females in seminatural or natural troops apparently move some distance from other troop members, fre- quently into concealing underbrush (Fedigan & Fedigan, 1977, p. 205; Nakamichi et al., 1992, p. 414; Fedigan & Zohar, 1997, p. 172; Thomsen, 1997, p. 339); all three or four reported parturi- tions in seminatural or natural groups occurred on the ground, not in trees. In three parturitions, con- tractions were first observed 35, 44, and 51 min- utes before birth of the infants (Itoigawa & Ta- naka, 1963, p. 76; Nakamichi et al., 1992, p. 414; cf. Kanazawa & Nakamichi, 1991, p. 424); in two of these parturitions, rupture of the amniotic sac reportedly occurred 4 minutes and 9 minutes be- fore birth (Itoigawa & Tanaka, 1963, p. 76), whereas in the third parturition, rupture of the sac reportedly occurred 50 minutes before birth (O- shima, 1980, p. 7). In caged, seminatural, and nat- ural populations, the placenta apparently is almost always completely consumed by the mother (Itoi- gawa & Tanaka, 1963, p. 76; Hazama, 1964, p. 89; Fedigan & Fedigan, 1977, p. 206; Murray & Murdoch, 1977, p. 818; Oshima, 1980, p. 7; Hirai- wa, 1981, p. 315; Negayama etal., 1986, pp. 372- 373; Nakamichi et al., 1992, p. 414; Thomsen, 1997, p. 339; Thomsen & Soltis, 2000, p. 690). 64 FIELDIANA: ZOOLOGY Table 41. Neonatal sex ratio in Macaca fuscata. Number of births Ratio Sex Observation unspeci- males/ Refer- Locality period Females Males fied Total females ences' Arashiyama 1957-66 95 81 44 220 0.85 / Arashiyama 1972-83 469 486 20 975 1.04 2 Awajishima 1978-96 281 311 — 592 1.11 3 Beaverton, U.S.A. 1966-75 -103 -74 — -177 -0.72 4 Berlin, Germany 1977-95 55 58 4 117 1.05 5 Choshikei 1958-66 34 47 68 149 1.38 1 Funakoshiyama 1965-66 31 41 — 72 1.32 I Gagyusan 1956-66 64 72 107 243 1.12 1 Hagachi 1966 5 2 2 9 0.40 1 Hakone-machi 1971-77 69 66 14 149 0.96 6 Houtosan 1963-66 33 28 — 61 0.85 1 Kanbanotaki 1962 15 12 — 27 0.80 I Katsuyama-cho 1958-86 482 437 — 919 0.91 7 Kinkazan 1983-94 40 56 1 97 1.40 8 Kochi 1958-59 9 15 — 24 1.67 I Kojima 1952-86 140 1602 21 321 1.14 9 Mihara-city 1964-65 5 9 — 14 1.80 I Minoo-city 1957-66 117 148 — 265 1.26 1 Miyajima-cho 1962-66 20 25 6 51 1.25 1 Nametoko 1963-66 9 13 24 46 1.44 1 Ohirayama 1957-66 70 92 — 162 1.31 1 Okinoshima 1958-66 29 27 1 57 0.93 1 Rome, Italy 1977-88 37 48 — 85 1.30 10 Rosando 1958-62 36 44 — 80 1.22 1 Shiga' 1963-75 56 58 4 118 1.04 11 Taishakukyo 1958-62 19 20 — 39 1.05 1 Takagoyama (A, S) 1959-66 28 31 26 85 1.11 I Takagoyama (I) 1974-78 53 36 — 89 0.68 12 Takaosan 1958-66 28 34 — 62 1.21 1 Takasakiyama 1956-62 260 289 118 667 1.11 13 Takasakiyama 1963-66 263 283 38 584 1.08 1 Takeno-ho 1965-66 7 14 4 25 2.00 1 Toimisaki 1958-65 40 43 22 105 1.08 1 Yaku.shima 1974-93 37 33 6 76 0.89 8 Totals — -3039 -3193 530 -6762 -1.05 — ' Key to references: /. Kawai et al. (1967, pp. 60-70). 2. Huffman (1991a, p. 33). 3. Nakamichi et al. (1997, p. 230). 4. Murray and Murdoch (1977, p. 818). 5. Harigel (1996b, p. 16). 6. Fukuda (1988, p. 481). 7. Itoigawa et al. (1992, p. 57). 8. Takahata et al. (1988a, p. 341). 9. Watanabe et al. (1992, p. 8; cf. Iwamoto, 1974, p. 259). 10. Aureli et al. (1990, p. 177). //. Tokita and Hara (1975, p. 27). 12. Hiraiwa (1981, p. 310). 13. Itani et al. (1964, p. 15). - Includes one pair of male twins. '' Shiga-A = Jigokudani. Neonatal Sex Ratio; Birth Weight In ca. 6232 M. fuscata births where sex of the infant has been determined, the ratio of males to females is ca. 1.05 (Table 41); this ratio is not quite significantly different from 1.00 (x" = 3.806, 0.10 > f > 0.05). Neonatal sex ratios in M. fus- cata apparently are not consistently correlated with the mother's dominance rank (Wolfe, 1984b, p. 141; Schino et al., 1999, p. 257; Brown & Silk, 2002, p. 11254). Judging from the small samples of birth weight that are available for M. fuscata (Table 42), mean weight in female neonates (546.8 g, n = 11) ap- pears to be fairly close to that in male neonates (538.7 g, n - 10). Twinning In a translocated M. f. yakui population at Ohi- rayama, one pair of twins was recorded among FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 65 Table 42. Birth weight (g) in Macaco fuscataJ Females Mean ± SD Males Sex unspecified Mean ± SD Mean ± SD Refer- ences- 600 536.75 600.0» 1 46 0 2 517 ± 90 4 — 0 546.8 11 500 Arashiyama 1 '450 Primate Research Institute, Kyoto University^ 543.0 ±21.0^ 4 — 558.3' 3^ — — 0 501.0 ± 52.9 460 1 — Medical Lake Primate Field Station, U.S.A. — 0 — Tierpark Sababurg, Germany 580 1 — Totals 538.7 10 -498.6 0 0 20 0 0 0 21 ' Cf. Ohno et al. (1980, p. 936); Kurita et al. (2002, pp. 419-420). - Key to references: 1. Hazama (1964, p. 91). 2. Oshima et al. (1977, p. 100). 3. Hamada (1982, p. 547). 4. Shimizu (1988, p. 252). 5. Ota et al. (1991, p. 36). 6. Sirianni et al. (1975, p. 20). 7. Harigel (1998, p. 231). ' Supplementary information reported by Oshima (1980, p. 7): "'The birth weight of the offspring delivered naturally ranged from 430 g to 610 g" (mean, sample size, and sex of infants not specified). ^Extremes, 518-570 g. 5 SD unspecified. ^ Age, in days, of infants when weighed: <1, 1, 5, 7. ^ Age, in days, of infants when weighed: <1, 1,6. ** Extremes, 545-655 g. 234 births that occurred during 13 reproductive seasons (1957-1969); in a natural M. f. fuscata population at Takasakiyama, one pair of twins was recorded among 488 births that occurred dur- ing 13 reproductive seasons (1977-1989); and in another natural M. f. fuscata population at Koji- ma, one pair of twins was recorded among 320 births that occurred during 35 reproductive sea- sons (1952-1986) (Tanaka et al., 1970, p. 114; Soumah & Yokota, 1992, p. 16; Watanabe et al., 1992, p. 8); the composite twinning rate for these three populations is 3/1042 = 0.29% (cf. Geiss- mann, 1990, p. 391). Two additional pairs of twins in two natural M. f. fuscata populations (sample sizes not specified) have been reported by Nakamichi (1983, p. 576), and one additional pair of twins in a seminatural population (sample size not specified) has been reported by Pavelka (1993, p. 18). At Arashiyama, a female nurtured a pair of "ar- tificial twin" infants for at least 8 months (Ogawa, 1998, p. 101); this pair of infants consisted of one male bom to the female on 13 May 1987 and a second male neonate adopted by her 1 day later. Birthrate The mean annual birthrate (births/sexually ma- ture females/year) in provisioned M. fuscata troops (ca. 0.57) apparently exceeds that in non- provisioned troops (ca. 0.38) (Table 43; see above. Natural History: Group Size and Compo- sition); corresponding mean interbirth intervals are 1.75 years and 2.63 years, respectively (cf. Ross, 1992, p. 209; Takahashi, 2002, p. 145). In provisioned troops, variations in the level of pro- visioning are positively correlated with variations in the birthrate (Mori, 1979b, p. 375; Watanabe et al., 1992, p. 7), and in nonprovisioned troops, an- nual variations in fruiting rates during mating and pregnancy seasons are positively correlated with variations in birthrates during subsequent birth seasons (Suzuki et al., 1998, p. 318). In the pro- visioned natural group at Arashiyama, the mean number of lifetime births per adult female report- edly was approximately eight (maximum, 13) (Koyama et al., 1975, p. 416). In females whose infants have survived for at least 1 year, intervals to the next birth are great- 66 FIELDIANA: ZOOLOGY Table 43. Annual birthrate (births/sexually mature females/year) in Macaca fuscata. Locality Observation period Number of births Annual birth rate References' Natural groups, nonprovisioned Kawaradake 1971-78 80 0.59 / Kinkazan 1983-92 249 0.36 2 Kojima- 1952-86 320 0.36 3 Ryozen 1974-80 45 0.34 4 Shimokita, SW 1965-74 10 0.31^ 5 Yokoyugawa Valley'' 1970-75 -11 -0.25 6 Yakushima (M. / \akui) Kojiba^ 1974-79 45 0.45 7 Mt. Kuniwari^ 1975-78 27 0.46 7 Nine troops^ 1989-95 46 0.28 2 Totals Natural groups, -833 provisioned -0.38 — Arashiyama 1954-83 973 0.62 8 Hakone-machi 1971-77 149 0.55 9 Katsuyama-cho 1958-86 919 0.44 10 Ryozen 1969-73 48 0.59 4 Shiga-A 1963-75 118 0.51 11 Takagoyama 1974-78* 87 0.53 12 Takasakiyama 1971-75 1406 0.62 6 Totals — 3700 0.57 — Seminaturai groups, provisioned Aarashiyama West, U.S.A. 1972-79 99 0.51" 13 Beaverton, U.S.A. 1966-67 15 -0.79 14 Berlin, Germany 1977-95 117 0.46 15 PRIKU 1989-90 19 0.50 16 Miyajima-cho'° 1969 13 0.50 17 Rome, Italy 1977-81 46 0.86 18 Totals — 210 -0.58 — ' Key to references: 1. Ikeda (1982, p. 344). 2. Suzuki et al. (1998, pp. 315-316; cf. Takahata et al., 1998b, p. 247; 1998a, p. 341); Sugiura et al. (2002, p. 76). 3. Watanabe et al. (1992, p. 8). 4. Sugiyama and Ohsawa (1982b, p. 248). 5. Azuma (1985, pp. 3-4). 6. Yoshihiro (1985, p. 55). 7. Maruhashi (1982, pp. 321-322; cf. Azuma, 1985, p. 2). 8. Koyama et al. (1992, p. 42). 9. Fukuda (1988, p. 481). 10. Itoigawa et al. (1992, p. 56). //. Suzuki et al. (1975, p. 18). 12. Hasegawa and Hiraiwa (1980, p. 130). 13. Gouzoules and Goy (1983, p. 42). 14. Alexander and Bowers (1967, p. 335). 15. Harigel (1996, p. 16). 16. Nozaki et al. (1995, p. 1251). 17. Stephenson (1975, p. 66). 18. Scucchi (1984, p. 204). - Population variably provisioned. ^ 10 births/32 females. ■* Yokoyugawa Valley = Shiga-B (see Shiga-kogen). "^ Includes daughter troops M, N, A, and H (products of successive fissioning); during the period 1984-89, the annual birth rate of M troop decreased to 0.15 (Takahata et al., 1994, p. 318). * Seven troops, excluding Kojiba troop. 7 Troops B, CC, G, H. M, NNA, P, S, T * Takagoyama. provisioning discontinued April 1976. ■* Omitted from totals. '° Translocated from Shodoshima in 1962. er — and birthrates are correspondingly lower — than in females whose infants have not survived (Table 44; cf. Tanaka et al., 1970. p. 115; Nomura et al., 1972, p. 478; Mitsunaga et al., 1992, p. 25; Takahashi, 2002, p. 145). Lactation apparently tends to suppress the ovarian endocrine function of females (Hiraiwa, 1981, p. 312; Yoshida et al., 2001, p. 371). The age-specific birthrate in M. fuscata appar- ently tends to increase during the first few repro- ductive years, tends to maintain a relatively high level during the middle reproductive years, and probably tends to decline during the last few re- productive years (Fig. 23; cf. Wolfe & Noyes, 1981, p. 699; Gouzoules et al., 1982, p. 1140; Sugiyama & Ohsawa, 1982b, p. 246; Fukuda, FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 67 Table 44. Effect of Macaca fuscata infant death on interbirth interval of mother. Mean interbirth interval (yr) Locality Following survival of infant to age 1 yr Following death of infant before age 1 yr Refer- ences' Natural groups, nonprovisioned Kinkazan 2.37 1 .59 Yakushima 2.24 1.50 Natural groups, provisioned Arashiyama i.46 1.15 Katsuyama-cho 1.58 1.29 Seminatural group, provisioned Rome 1.22- 1.06^ ' Key to references: /. Takahata et al. (1998a, p. 341). 2. Scucchi (1984, p. 205). ^ Survival of infant to age 6 months. ^ Death of infant before age 6 months; includes still- births. 1988, p. 482; Watanabe et al., 1992, p. 17; Hari- gel, 1996, p. 16; Shimizu, 1998, p. 1 15). In seven groups where the relationship between birthrate and female dominance rank has been studied (Ta- ble 45), birthrate is positively correlated with dominance rank in two groups, birthrate is not correlated with female dominance rank in four groups, and evidence is equivocal in one group. Infant Mortality The infant mortality rate (deaths before age 1 year/births) in nonprovisioned groups of M. fus- cata (mean = 28.4%) apparently is more than twice as great as that in most provisioned natural groups (9.4%) and seminatural groups (8.5%) (Ta- ble 46); the high infant mortality rate reported in the Hakone T provisioned natural group (49.7%) is exceptional and remains unexplained (Fukuda, 1988, p. 483). Approximately half of all infant deaths probably occur before age 1 month (Itoi- gawa et al., 1992, p. 63; Koyama et al., 1992, p. 41 ; Watanabe et al., 1992, p. 22; Nakamichi et al., 1997, p. 233). Known violent causes of infant deaths include attacks by dogs or raccoon dogs (Iwamoto, 1974, p. 257) and attacks by adult male monkeys (Kur- land, 1977, p. 83; Soltis et al., 2000, p. 197; Taka- hashi, 2002, p. 151). Mothers have been observed to carry the bodies of stillbirths or dead infants for several days, even after the bodies have begun to decompose (Green, 1975, p. 15; Nakamichi et al., 1983, p. 64; Pavelka, 1993, p. 63; Takahashi, 2002, p. 144). Nursing; Weaning Three observed newborn M. fuscata infants be- gan to suckle 23 minutes, ca. 90 minutes, and 1 10 minutes after parturition (Itoigawa & Tanaka, 1963, p. 76; Thomsen, 1997, p. 339). During the Table 45. Relationship between birthrate and female dominance rank in nonprovisioned and provisioned groups of Macaca fuscata. Locality Birthrate significantly Observation Number correlated with female period of births dominance rank? References' Kinkazan Kojima- Yakushima Arashiyama Arashiyama West"" Katsuyama-cho Takasakiyama Nonprovisioned groups 1983-94 97 No 1 1952-86 258 Evidence equivocal 2 1974-93 76 Provisioned groups No 1 1954-83 975 No 3 1976-78 >82 No 4 1958-84 919 Yes 5 1977-89 488 Yes 6 ' Key to references: /. Takahata et al. (1998a, p. 345). 2. Mori (1979b, p. 388); Watanabe et al. (1992, p. 11). 3. Wolfe (1984b, p. 139); Koyama et al. (1992, p. 42); cf. Fedigan et al. (1986, p. 151). 4. Gouzoules et al. (1982, p. 1141); Fedigan et al. (1986, p. 151). 5. Itoigawa et al. (1992, p. 65). 6. Soumah and Yokota (1992, p. 15). - Population variably provisioned. ^ South Texas, U.S.A. 68 FIELDIANA: ZOOLOGY Table 46. Infant mortality rate (infant deaths before age 1 yr/births) in nonprovisioned and provisioned groups of Macaca fuscata. ' Locality Observation years Number of births Infant mortality rate (%) References^ Kinkazan Kojima' Shiga B-2 Shimoicita SW Yakushima Totals Arashiyama Awajishima Hakone T Katsuyama-cho Shiga A Takagoyama-I Takasakiyama Totals Arashiyama West'' Natural groups, nonprovisioned 1983-94 97 22.7 1 1952-86 320 31.2 2 1970-75 15 40.0^ 3 1965-74 9 0 4 1974-93 76 25.0 1,4,5 — 517 28.4 — Natural groups, provisioned 1954-83 953 10.3 6 1978-95 543 12.3 7 1971-77 149 49.7 8 1958-85 949 10.2 9 1963-75 118 6.85 3, 10 1975-78 68 16.2 11 1956-62 667 4.5 12 1987-89 215 882 8.8 5.6 13 — 3513* 9.46 — Seminatural group, provisioned 1972-93 1230 8.5 14 ' For mortality rate to age 2 years at Ryozenyama, see Sugiyama and Ohsawa (1982b, p. 250) and Takahata et al. (1998a, p. 341). - Key to references: 1. Takahata et al. (1998a, p. 341). 2. Mori (1979b, p. 375); Watanabe et al. (1992, p. 8). 3. Suzuki et al. (1975, pp. 19, 21; cf. Yoshihiro, 1985, p. 55). 4. Azuma (1985, pp. 3, 4). 5. Maruhashi (1982, p. 322). 6. Koyama et al. (1992, p. 41). 7. Nakamichi et al. (1997, p. 233). 8. Fukuda (1988, p. 481). 9. Itoigawa et al. (1992, p. 63). 10. Tokita and Hara (1975, p. 27). //. Hiraiwa (1981, p. 319). 12. Itani et al. (1964, p. 15). 13. Soumah and Yokota (1992, p. 17). 14. Fedigan and Zohar (1997, p. 168). ^ Population variably provisioned. ^ Excludes two miscarriages and/or stillbirths. ■* Excludes seven miscarriages and/or stillbirths. *> Excludes aberrant Hakone T data. ^ South Texas, U.S.A. first 2 weeks of life, 33 infants in a seminatural group spent an average of 87% of observed in- tervals in contact with their mothers' ventrum; 69% of these intervals included infants' oral con- tact with their mothers' nipples (Murray & Mur- doch, 1977, p. 819). Infants reportedly begin to taste solid food at age 2 weeks, and while con- tinuing to nurse, they begin to eat solid food at age 4-6 weeks (Pocock, 1906, p. 567; Hasegawa & Hiraiwa, 1980, p. 135; Hiraiwa, 1981, p. 313). When an infant is approximately 6 months old — which is near the beginning of the troop's next mating season — its mother's milk secretion abruptly decreases (Ota et al., 1991, pp. 41, 47; Tanaka, 1992, p. 133), and when an infant is 7- 10 months old, its mother may reject the infant's attempts to suckle (Fedigan & Fedigan, 1977, p. 217; Hasegawa & Hiraiwa, 1980, p. 135; Hiraiwa, 1981, p. 318); such rejection reportedly is more frequent in estrous females than in nonestrous fe- males (CoUinge, 1987, p. 142; 1991, p. 163; Pav- elka, 1993, p. 66). Although some infants may be fully weaned by age 6-8 months (Pocock, 1906, p. 567; Tanaka et al., 1970, p. 116; Ota et al., 1991, p. 47; Pavelka, 1993, p. 66), some continue to nurse to age ca. 1 year — until just before their mother's next parturition (Tanaka, 1992, p. 136; Inoue et al., 1993, p. 505), and others may con- tinue to nurse to age ca. 2.5 years — if their moth- ers have had no intervening parturitions (Fedigan & Fedigan, 1977, p. 217; Hiraiwa, 1981, p. 315; Sugiyama & Ohsawa, 1982b, p. 250; Nakamichi, 1989, p. 739; Tanaka, 1992, p. 136; Pavelka, 1993, p. 66; Tanaka et a!., 1993, p. 171). Adult males oi M. fuscata, including high-rank- ing males, occasionally have been observed to FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 69 ^^ T3 9" ® -- 2 .52 c ^ 55 > .2 o i > 2 « « 1 ..I onpro (nonp (prov (provi 1 i <-'' inkazan (n akushima rashiyama atsuyama o:::['< . . .<' ■ ;:< ) '^ > < ^ i _'<::.'■- - o ' o ^1 0 ^ 1 _\ K'''7r ''''''' <>'-'-. \<' ■••.■.'--:.. \ ..<■[ ^■■■■■' o - - - -^' - / o::;-- < / \ 1 "*■*•'- / <'\o / _ . . . -o / 1 , 1 .--'' ""■--. / ' o---'/. , .".*-<:< ^^ ^^^. ^ — ^^^ <:;" ""'"-o ^^""^--v. ^ j o^ . . . '<. ^-^""^^^^^^J* "''■-. ^^ — "^ / <-'/;o • — A •»'**• f < "k:^ i .-•' " / ! *«■'.-'-. ^""^■♦^^1 "'•""■*-. *L '"o """-< V^^ 0--- < o o CD O in o o CO o CN (%) aiBJ mjjg 70 FIELDIANA: ZOOLOGY Table 47. Mean life span in Macaca fuscata females. Locality Number of Birth period of deceased females studied females Mean (±SD) female life span (yr) References' Yakushima: M. X P Natural population, nonprovisioned ca. 1974-93 36 6.3 ± 5.2 / Takasakiyama Natural population, provisioned ca. 1952-73 — 8.4 ± ? 2 Arashiyama AAVest Natural/seminatural- population, provisioned 1950-67 72 13.6 ± 9.4^ 3 Key to references: /. Takahata et al. (1998a, p. 342). 2. Masui et al. (1975, p. 405). 3. Fedigan (1991b, p. 142). Natural provisioned population prior to February 1972; seminatural provisioned population subsequently. For this calculation, reported deaths at unspecified ages < 1 year were arbitrarily set at 0.5 years. protect and groom infants (Itani, 1959, p. 89; Ta- kahata, 1982a, p. 14; Yoshimura, 1984, p. 25). Longevity; Reproductive Senescence The greatest accurately known life spans in M. fuscata are 33 years in a female in a provisioned natural group (Koyama et al., 1992, p. 37; of. Pav- elka & Fedigan, 1999, p. 458; Fedigan & Pavelka, 2001, p. 113) and 28 years in a male in another provisioned natural group (Nakamichi et al., 1995, p. 386); greater estimated ages of 30-40 years have been reported for four males, but the actual birth dates of these males are unknown (Itani et al., 1964, p. 18; Tasumi, 1969, p. 263; Itoigawa, 1982, p. 380). Mean female life spans apparently are much less than half of the greatest known female life span; reported mean female life spans are 6.3 years in a nonprovisioned natural population, 8.4 years in a provisioned natural pop- ulation, and 13.6 years in a provisioned natural/ seminatural population (Table 47). Because males usually emigrate from their natal troops, deter- mination of mean male life spans in unconfined populations is impractical; in a confined popula- tion, the mean male life span apparently was ca. 60% of the mean female life span (Fedigan & Zohar, 1997, p. 165). The greatest age at which a female M. fuscata is known to have borne an infant is 26 years (Itoi- gawa et al., 1992, p. 56; Shimizu, 1998, p. 115); this occurred in a provisioned natural group at Katsuyama-cho. In a provisoned natural group at Arashiyama, the mean (±SD) age at last parturi- tion in seven females that survived to age 20 years was 21.7 ± 2.0 years (extremes, 19-25 years) (Takahata et al., 1995, p. 172); similarly, at Ara- shiyama West, South Texas, the mean age at last parturition in 16 females that survived to age 20 years was 21.3 ± 3.2 years (extremes, 11-25 years) (Pavelka & Fedigan, 1999, p. 458). The mean postreproductive interval between a fe- male's last parturition and her disappearance (probable death) from the Arashiyama group was 6.0 ±3.5 years (1-10 years, n = 8) (Takahata et al., 1995, p. 172), and the corresponding mean postreproductive interval between a female's last parturition and her death in the Arashiyama West group was 5.0 ± 2.5 years (2-11 years, n = 20) (Pavelka & Fedigan, 1999, p. 458). Postreprod- uctive females may contribute to the fertility of their daughters, and they probably contribute to the survival of their daughters' offspring (Pavelka et al., 2002, p. 412). Although the sexual activity of females appar- ently tends to decline during the postreproductive interval, females at this stage often continue to have estrous cycles and to engage in copulation (Koyama et al., 1992, p. 37; Mitsunaga et al.. Fig. 23. Age-specific birthrates in two nonprovisioned troops (solid symbols) and two provisioned troops (open symbols) oi Macaca fuscata: nonprovisioned troops — Kinkazan, 1983-1994, 97 births, and Yakushima, 1974-1993, 76 births (Takahata et al., 1998, pp. 343-344); provisioned troops — Arashiyama, 1954-1983, 975 births (Koyama et a!., 1992, p. 37); Katsuyama-cho, 1958-1986, 919 births (Itoigawa et al., 1992, p. 56). FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 71 1992. p. 28: Takahata et al.. 1995. p. 174): cop- ulation by a 32-year-old female has been reported (Oda & Masataka. 1995. p. 135). The ovaries of postreproductive females reportedh are reduced in size, impaired histologically, and impaired en- docrinologically (Mitsunaga et al.. 1992. p. 26: Nozaki et^al.. 1995. p. 1253: 1997. p. 92: Shimizu. 1998. pp. 115. 117: Yoshida et al.. 2001. p. 370). A male M. fitscata at KatsuN ama-cho \\ as ob- served to copulate at age ca. 27 years, and another male at Takasakiyama was observed to chase an estrous female at age ca. 30 years (Itoiga\\a. 1982. p. 382). At Arashiyama West, however, a male of estimated age >20 years rarely main- tained an erection, although he consorted with five estrous females. The testes and accessory glands of males of estimated age ^26 % ears \\ ere found to be degenerate relati\e to those of males age 20-2 1 \ ears, although a few sperm \\ ere noted in the seminiferous tubules of males near age 30 years (Matsubayashi et al.. 1994. pp. 201-204). Population Growth Rate Predictably, the mean population growth rate in nonprovisioned natural M. fuscoto groups (ca. 2.5^^) is considerably less than that in provisioned natural groups (ca. 9.0*^) and in provisioned seminatural groups (ca. 12.3^) (Table 48). Fossils Fossils referred to Macaco Jiiscata. associated w ith numerous other fossil mammals, have been unearthed at various localities on Honshu. Shi- koku and Kyushu (Tables 49. 50: Fig. 24). Based on tephrostratigraphy and oxygen isotope strati- graphy of Japanese Pleistocene proboscidean spe- cies, it has been concluded that dry-land connec- tions ("land bridges") between the Japanese is- lands and the adjacent Asian continent existed twice during the Middle Pleistocene (Kawamura. 1998. p. 252: Dobson & Kawamura. 1998. p. 387: Konishi and Yoshikawa, 1999. p. 131). These land bridges apparently occurred at oxygen isotope stage 16 (0.63 million years ago: Ma) and at ox- ygen isotope stage 12 (0.43 Ma), when Stegodon ohentalis and Palaeoloxodon naunianni. resp>ec- tively, immigrated into the Japanese islands. S. or- ientalis survi\ed for some time but apparently was ultimately replaced by P. naunianni: these two proboscideans were not coexistent. To date, the purported oldest macaque fossil in Japan is an isolated left lower third molar (mesio- distal length. 12.1 mm: buccolingual width. 7.0 mm), comparable in size and shaf)e to correspond- ing molars of the extant Japanese macaque, that was obtained from a fissure deposit at Ando Quar- ry. Yamaguchi Prefecture (Iwamoto and Hasega- wa. 1972. p. 78: Aimi. 2002. p. 242). The asso- ciated fossil proboscideans are S. orientalis and P. naunianni, which indicates that the fossil as- semblage at Ando Quarry is a mixed one. The age of the fossil macaque molar remains elusive: if it was originally associated with 5. orientalis, its age is ca. 0.63 Ma. but if it was originally associated with P. naunianni. its age is ca. 0.43 Ma. Another early macaque fossil in Japan is a right humerus, similar to that in extant male Japanese macaques, that was unearthed at Yarimizu, Jizodo Formation or Kamiizumi-Kiyokawa Formation, Chiba Prefecture (Iwamoto and Hasegawa. 1991, p. 100). The associated proboscidean fossils have been identified as P. naunianni. A\ailable evidence suggests that the ancestors of M. fuscata dispersed from mainland Asia to the Japanese Islands either ca. 0.63 Ma or ca. 0.43 Ma. No evidence exists concerning the possibility of disp>ersal of M. fuscata ancestors to Japan be- fore 0.63 Ma. Several macaque fossils ha\e been obtained from late Middle Pleistocene to Late Pleistocene deposits on the Korean Peninsula, where no ma- caques occur at present (Lee. 1983, p. 40: Park and Lee, 1998, p. 56). Although some of these Korean fossils haN e been referred to Macaca cf. robitsta based on their molar size (Table 51), the measurements of these fossil molars actually are within the range of variation show n by Japanese macaques (Table 52). Further comparative studies of the Korean fossils and extant Japanese ma- caques are required to in\estigate the possibility of an evolutionary relationship. Systematics Geographic Variation and Subspecific Recognition Based on geographic variation of pelage color and body size. Kuroda (1940, p. 273) proposed that the population of M. fuscata that inhabits 72 FIELDIANA: ZOOLOGY Table 48. Population growth rate in Macac a fuscata groups. Number Annualized of population groups Census Initial Final growth rate' Refer- Locality observed interval census census (%) ences- Natural groups, i nonprovisioned Kinkazan 5 1983-93 -270 -260 — 0.3^ 1 Kojima^ 1 1952-86 21 -100 -4.7 2 Shimokita M 1 1971-85 -24 -79 -8.9 3 Shimokita O 1 1963-81 -12 -20 -2.9 4 Takagoyama I' 1 1976-78 107 100 -3.3 5 Yakushima'' 1-4 1974-90 42 -60 -2.2 6 Mean ± SD — Natural groups i, provisioned — -2.5 ± 4.2 — Arashiyama A and B 2 1954-72 -28 301 -14.1 7 Arashiyama B 1975-78 185 253 11.0 8 Awajishima 1978-95 -100 >250 -5.5 9 Gagyusan 1955-58 110 160 13.2 10 Hagachizaki 1955-59 >70 >100 -9.3 U Hakone T 1956-67^ -35 -75 -7.2 12 Katsuyama-cho 1-2 1958-86 112 236 2.7 13 Shiga A 1963-75 23 95 12.5 14 Shiga A-l 1990-97 108 252 12.9 15 Takagoyama I' 1974-76 98 107 4.5 5 Takasakiyama 1-3 1951-92 166 1963 6.2 16 Mean ± SD — — — — -9.0 ± 4.0 — Seminatural groups, provisioned Arashiyama West, U.S.A. 1972-94 150 >600 >6.5 17 Beaverton, U.S.A. 1964-78 46 316 14.8 18 Berlin, Germany 1977-95 9 77 12.7 19 Arnhem. Netherlands 1973-85 8 26 10.3 20 Montreal, Canada 1984-93 14 40 12.4 21 Nojima 1969-71 51 113 48.9« 22 Rome. Italy 1977-81 27 51 17.2 23 Mean ± SD — — — — -12.3 ± 3.7" — ' Annualized growth rate = [(C^/C,)"^] - 1, where Y = years in census interval, C, = initial census, and C2 - final census. - Key to references: 1. Takahata et al. (1998a, p. 340). 2. Watanabe et al. (1992, p. 4). 3. Watanuki et al. (1994, p. 16). 4. Izawa (1982. p. 13); Azuma (1985, p. 4). 5. Hasegawa and Hiraiwa (1980, p. 130). 6. Takahata et al. (1994, p. 319; cf. Maruhashi, 1982, p. 330). 7. Koyama et al. (1975, p. 412). 8. Grewal (1980, p. 331; cf. Takahata, 1980, p. 304). 9. Nakamichi et al. (1997, p. 226). 10. Furuya (1960, p. 149). //. Nishida (1966, p. 151). 12. Fukuda (1988, pp. 480, 481). 13. Itoigawa et al. (1992, pp. 52, 53). 14. Suzuki et al. (1975, p. 18; cf. Tokita & Hara, 1975, p. 27). 15. Tanaka (1998. p. 1230). 16. Ohsawa and Sugiyama (1996. pp. 163, 167). 17. Jack and Pavelka (1997, p. 370). 18. Eaton (1976, p. 105); Rostal et al. (1986. p. 453). 19. Harigel (1996. p. 16). 20. Scheurer and Thierry (1985, p. 491). 21. Prud'homme and Chapais (1996, p. 432). 22. Nomura et al. (1972, p. 481). 23. Scucchi (1984, p. 204). ^ During the winter of 1983-84, an unusually heavy snowfall at this locality resulted in the deaths of ca. 90 monkeys (Takahata et al., 1998a, p. 340). ■* Variably provisioned. '' Provisioning of this group was discontinued in March 1976. ^ Kojiba troop and its daughter troops. ^ Excludes period after 1 967, when uncensused daughter troops separated from the main troop. •* Birthrates in this group were artificially enhanced by removing infants from their mothers 3-6 months after birth. ' Excludes Nojima mean (see footnote 8). Yakushima, a small island at the southern limit of distribution of the species, should be recognized as a distinct subspecies, M. f. yakui. Comparing the Yakushima population with more northern populations, Kuroda noted that pelage color is darker — particularly on the back, limbs, and ex- tremities— in the Yakushima population and that body size is smaller in this population. Although FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 73 e « at ■a ■'r^ n i/i £> «h! O u 60 oa r\i^"^'oM-^f\'^ooooooC)"^-~.NO I I i + i I++I + I+ + +I + I I I I I I I I I I I I I I I + + ^ — ^^^.£^^ ,(U ^ ,^-E S >- 00 00 < (3 O oI.E E CO ^. =5 •- tJ- o o- U c. .2 .-^ S '5 cox: c .i: 3 ^ r3 r- 1/5 3 0 U S „ 3 C/0 h- 3 X) = c b35°N), broadleaf forest habi- tats— suitable for M. fuscata — apparently were less abundant than at present (Tsukada, 1982, p. 1092; Takahara et al., 2000, p. 673; Harrison et al., 2001, p. 130). During glacial sea-level lowering, M. fuscata presumably was able to disperse to most or all of the small shallow-water islands that surround Kyushu, Shikoku, and Honshu. Subsequently, however, this species apparently became extinct on all except six of these small islands (Awajish- ima, Kashima, Kinkazan, Kojima, Shodoshima, and Yakushima). Such extinctions on small is- lands have recently been recorded on Omishima, Hiradoshima, and Tanegashima (see above. Geo- graphic Distribution). Subsequent to the initial arrival of M. fuscata on the Japanese archipelago, its body size appar- ently has tended to increase as the species dis- persed northward, in accord with Bergmann's rule (Figs. 5, 6, 9, 10, 15; Mayr, 1963, p. 320). Four measurements of body size — sitting height, ante- rior trunk length, hind foot length, and greatest length of skull — generally tend to increase with latitude from Yakushima to southern Honshu (ca. 35°N) but do not tend to increase with latitude farther north on Honshu; one measurement of body size — body weight — tends to increase with latitude from Yakushima to northern Honshu. A hypothesis to account for present body-size vari- ation in M. fuscata is suggested by Kawamoto's (2002, p. 68; see above. Molecular Biology and Genetics) interpretation of mtDNA variation in this species: body-size variation in M. fuscata may have been completely Bergmannian prior to the last glacial maximum, but northern popula- tions (large-bodied) may have become locally ex- tinct because they were unable to survive unfa- vorable glacial climate conditions; during the sub- sequent postglacial period, northern Honshu may have been reoccupied by immigrants from south- ern Honshu (smaller-bodied). Body weight may be more plastic than the other four body-size at- tributes, and it therefore may have more rapidly regained a Bergmannian pattern of latitudinal var- iation. The apparent reduction of body size in Boso Peninsula and Kinkazan populations of M. fuscata is unexplained. In accord with Allen's rule (Mayr, 1963, p. 323), tail length in M. fuscata apparently has de- creased relative to that in its presumed M. mulat- r«-like progenitors (Tables 5, 8; Fooden, 2000, p. 29). Tail length in M. fuscata is approximately as predicted by the tail length vs. latitude regression determined by other fascicularis-group species (Fooden & Albrecht, 1999, p. 433). Pelage color in available samples of M. fuscata generally becomes paler with increasing latitude, in accord with Gloger's rule (Table 2; Mayr, 1963, p. 324). This apparently is another example of ad- aptation to local climate that has occurred since monkeys initially arrived on the Japanese archi- pelago. As indicated above (see Molecular Biology and Genetics), the history of dispersal of M. fuscata within the Japanese archipelago that is implied by mtDNA evidence differs from the history of dis- persal that is implied by blood-protein evidence. MtDNA evidence suggests that, since the last gla- cial maximum, a relatively new M. fuscata pop- ulation— with group II haplotypes — has rapidly expanded its distribution in northern Honshu and replaced a now disjunctly distributed older pop- ulation— with group I haplotypes (Kawamoto, 2002, p. 68). Contrastingly, blood-protein evi- dence suggests that extant monkey populations on Yakushima, Boso Peninsula, and Shimokita Pen- insula are relicts of a formerly widespread first wave of immigrants to the Japanese archipelago that subsequently were displaced elsewhere on the archipelago by a possibly better adapted second wave of immigrants (Nozawa et al., 1991, p. 433; 1996, p. 33). Further research is required to rec- oncile these two interpretations. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 85 In summary, the progenitor of M. fuscata prob- ably was a M. mulatta-Uke population that dis- persed via a now-submerged land bridge from the Korean Peninsula area to Kyushu-West Honshu; this dispersal probably occurred during one or two glacial intervals of sea-level depression ca. 0.43- 0.63 Ma. In addition to dispersing to Kyushu, Honshu, and Shikoku, M. fuscata probably also dispersed to most or all of the surrounding small shallow-water islands, but not to Hokkaido; sub- sequently, the species apparently became extinct on all but six of the surrounding shallow-water islands. Since the arrival of M. fuscata on the Jap- anese archipelago, its body size, tail length, and pelage color apparently have tended to become adapted to local climate conditions. As a result of unsuitable climate and vegetation conditions dur- ing the last glacial maximum (ca. 1 8 Ka), M. fus- cata may have become locally extinct in northern Honshu; during the subsequent postglacial period, this area may have been reoccupied by a popu- lation from southern Honshu. Acknowledgments For access to specimens and generous assis- tance, we are deeply grateful to officials and staff members of institutions cited above (see Intro- duction). For valuable consultation, we thank Eric Delson (AMNH), Siho Fujita (PRIKU), Goro Hanya (PRIKU), Takashi Hiraoka (Yamashina In- stitute for Ornithology), Mitsuo Iwamoto (JMC), Paula Jenkins (BM[NH]), Yoshi Kawamoto (PRI- KU), Naoki Koyama (Kyoto University), Shiro Matsuoka (Working Group for Monkeys on Shi- mokita Peninsula), Yukihisa Mito (JMC), Akio Mori (PRIKU), Yasuyuki Muroyama (PRIKU), Toyohiro Nishimoto (National Museum of Japa- nese History), Hideyuki Ohsawa (PRIKU), Chris Smeenk (RMNH), Shigeru Suzuki (Kyoto Uni- versity), Kazuo Wada (Inuyama), Kunio Watana- be (PRIKU), and Takao Yamaguchi (Kumamoto University). Officials at Kozanji Temple gener- ously permitted us to reproduce part of "Choju- jinbutsu-giga" (Frolicking Animals and Figures Scrolls). Photographs reproduced in Figures 3 and 1 1-14 were taken by Minoru Kinoshita (PRIKU). W Burger, S. Drasner, M. Pannell, C. Simpson, and H. Voris (all at Field Museum of Natural His- tory) facilitated publication, and K. H. 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Kanagawa (2): Fuji-san— MNHN 1887/15, 1887/571 (skulls inside). Prefec- ture unknown (7): USNM 13830 (female skin mismatched with male skull; skin may belong with skull USNM 13831), ZSBS 1901/101, 1904/ 1718-1904/1722. KOJIMA, 1. Miyazaki (1): KFL "Hiyoshi" (name of monkey). Island unknown, 12. ANSP 4952 (skull inside), BM(NH) 1861.8.20.1, MCZ 6703, RMNH 2-b, 3- C/D20G, 4-d/D20L, 6-f (skull inside), ZMB 12956/RexuCo, 22203, 32430, 34244, ZMUZ 11603. Skulls or Skeletons Only, 1035 HONSHU, 863. Akita (1): Hachimori-machi— PRIKU 6651 (mandible and postcranials only). Aomori (4): Bushidomari — JMC 2577; Kusoudo- mari — JMC 1757; Nishimeya-mura — JMC 1506; Wakinosawa— PRIKU 2153. Chiba (284): Aika- wa— PRIKU 6580, 6586, 6645, 6758, 6762; Fu- kurogura,— PRIKU 6786; Futairi— PRIKU 6644 (cranial fragments); Hagiu — PRIKU 6588; Hina- batake— PRIKU 6217, 6265, 6516, 6524, 6585, 6767; Hirooka— PRIKU 6283; Hongou— PRIKU 6761; Ishizuka— PRIKU 6493, 6507, 6522, 6774; Kaisho— PRIKU 6258, 6287; Kamou— PRIKU 6242, 6280, 6282, 6290 (cranial fragments, man- dible, postcranials), 6302, 6566, 6590; Kitako- machi— PRIKU 6260, 6783; Kiwadahata— PRI- KU 6293 (cranial fragments, mandible, postcran- ials), 6300, 6495, 6593, 6765, 6773; Kokurano— PRIKU 6256, 6276, 6284; Koshikiya— PRIKU 6776; Kozawamata— PRIKU 6294 (cranial frag- ments, mandible, postcranials), 6624, 6756 (cra- nial fragments), 6789; Kuradama— PRIKU 6235, 6236, 6241, 6245, 6274, 6291 (cranial fragments, mandible, postcranials), 6292 (cranial fragments, mandible, postcranials), 6296 (cranial fragments, mandible, postcranials), 6298, 6299 (cranium re- numbered "6239"), 6301, 6491, 6492, 6494, 6500, 6502, 6503, 651 1, 6519, 6573, 6587, 6591, 6592, 6594, 6607, 6608, 6752, 6757, 6768, 6769, 6778; Mera— PRIKU 6253; Kayano-Nanamaga- ri— PRIKU 6251, 6501, 6625 (cranial fragments); Nishihikasa— PRIKU 6497; Nutazawa— PRIKU 6760 (cranial fragments), 6779; Okugome — PRI- KU 6505, 6510 (cranial fragments), 6576, 6599, 6600, 6643 (cranial fragments), 6754 (cranial fragments), 6763, 6780, 6792; Okutani— PRIKU 6295 (cranial fragments, mandible, postcranials); Okuyama— PRIKU 6513; Ooiwa— PRIKU 6496, 6499; Ootashiro— PRIKU 6243 (female, misla- beled as male); Sanda— PRIKU 6575; Shigumi— PRIKU 6244, 6271, 6787; Shimosakuma— PRI- KU 6249, 6252, 6262, 6269, 6281, 6286, 6288, 6289, 6571, 6595, 6610; Shippara— PRIKU 6263, 6267, 6270, 6273, 6521, 6577, 6609, 6771, 6784; Sunada — PRIKU 6237 (female, mislabeled as male), 6266, 6272, 6297 (cranial fragments, man- dible, postcranials), 6498, 6506, 6508, 6509, 6514, 6515, 6517, 6572, 6606, 6759, 6766, 6785, 6788, 6791, 6793; Tagura— PRIKU 6504, 6581, 6753, 6775, 6781; Takagoyama— JMC 2588 (mandible lacking), SU [1], [2], [3], [4], 5, 6, 7, 8, 9, 10, H-1, H-2, H-3, H-4 (postcranials only), H-5, H-?, HK-34, I-l, 1-2, 1-3, 1-4, 1-5 (postcran- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 105 ials only), I-6A, I-6B, I-6C, 1-9, I- 1 1, 1-12, 1-13 (postcranials only), 1-13, 1-14, I-? [1], I-? [2], K- 1 (postcranials only), K-2, K-3, K-4, K-5, K-11 or K-14 (postcranials only), K-12, K-13, K-15, K- 15?, Ko-1, Ko-2, Ko-3, Ko-4, Ko-5, Ko-6, Ko-7, Ko-8, Ko-9, Ko-10, M-1, M-2A, M-2B, M-3, M- 4, M-5 (postcranials only), M-7, M-9, M-10, M- 11, M-1 2, M-1 3 (postcranials only), M-1 5, N-1, N-2, N-3 (postcranials only), N-4, N-5, N-6, N- 6?, N-7 (postcranials only), N-8, N-9, N-10, N- 11, N-12B, N-12[A], N-13, N-14, N-16, N-17, W- 1, Y-1, Y-2, Y-2?, Y-3 (postcranials only), Y-4, Y- 5, Y-5?, Y-6 (postcranials only), Z-1, Z-2, "Jiro- chou" (name of monkey); Tashiro — PRIKU 6257; Tokuji— PRIKU 6770, 6790; Tsukagoshi— PRIKU 6238, 6254 (cranial fragments, mandible lacking, postcranials), 6255, 6259, 6279; Tsuki- zaki— PRIKU 6247, 6250, 6264, 6268, 6518, 6523; Tsutsumori— PRIKU 6275, 6285; Utsutsu- mi— PRIKU 6239, 6240, 6246, 6261, 6338, 6512, 6520, 6574, 6578, 6579, 6583, 6589, 6596-6598, 6755, 6764, 6777, 6782; Yamanaka— PRIKU 6582, 6772; Yokose— PRIKU 6248, 6277, 6278 (mandible fragmented); Yonezawa — PRIKU 6584. Fukui (15): Doumoto— PRIKU 5192, 5826 (cranial fragments); Hotokedani — PRIKU 5415 (postcranials only); Kaminaka-cho — PRIKU 5184, 5193, 5194; Kidani— PRIKU 5183, 5824 (cranial fragments, mandible, postcranials); Mi- kata-cho— JMC 2012, 2034, 2035; Naka— PRI- KU 5414, 5736 5737; Otomi— PRIKU 1869 (mandible lacking). Hiroshima (1): Miyajima- cho — PRIKU 6647 (cranial fragments). Hyogo (14): Kasuga-cho— JMC 3486, 3517, 3525, 3556, 3566, 3567, 3570; Sasayama-cho— JMC 3642, 3643, 3653, 3658, 3666, 3667; Takeno-cho— JMC 2502. Kyoto {\A): Hiyoshi-cho— JMC 3488, 3503, 3505, 3511, 3520-3522; Ine-cho— JMC 3501, 3504, 3515, 3519; Kamigamo— PRIKU 6129; Mizuho-cho— JMC 3198; Wachi-cho— JMC 4214. Mie (4): Ilga-cho— PRIKU 257, 2111, 21 12; Oomiya-cho— PRIKU 6530. Nagano (261): Aoki— PRIKU Coll. Nos. 86, 87; Atosugiyama— PRIKU Coll. No. 49; Azumabasi— PRIKU 6317; Buna — PRIKU 6106 (cranial fragments, postcran- ials), 6189, 6200 (cranial fragments, mandible, postcranials) , 6203, 6209; Fukagasawa— PRIKU 6477; Fukasawa— PRIKU Coll. Nos. 46, 47, 51; Furukaidou— PRIKU 6488, 6568; Higashiina— PRIKU 6461; Higashino— PRIKU 6098, 6113, 6474, 6480, 6800; Hinata— PRIKU 6099; Hina- tayama— PRIKU 6163. 6817; Hodaka Golf Course— PRIKU 6469; Horane— PRIKU 6463, Coll. No. 178; Inokamireien— PRIKU 6820; Iri- sawai — PRIKU 6799 (cranial fragments); Kaki- zore— PRIKU 6470, 6619; Kamidaira— PRIKU 6100, 6105, 6115, 6167, 6168, 6177, 6330, Coll. Nos. 65, 73, 78-81, 88-90, 108-115, 124, 141- 145; Kamimachi — PRIKU 6466; Kamiseinaiji — PRIKU 6483, 6823 (cranial and mandibular frag- ments); Karitayama — PRIKU 6490; Kashio — PRIKU 6123, 6165; Kasugadaira— PRIKU 6124 (cranial fragments), 6180 (cranial fragments, post- cranials), 6197 (cranial fragments, mandible, post- cranials), 6208, 6214, 6487 (cranial and mandib- ular fragments); Katsurashima — PRIKU 6806; Kawanishi— PRIKU Coll. Nos. 55, 56; Kawasu- so— PRIKU 6603; Kiso— PRIKU 1486, 2210; Ki- tamura— PRIKU 6176/6454, 6192 (cranial frag- ments, mandible, postcranials), 6194 (cranial frag- ments, mandible, postcranials), 6199, 6201 (cra- nial fragments, mandible, postcranials), 6202 (cranial fragments, mandible, postcranials), 6204, 6206, 621 1, 6454; Coll. No. 28; Konishi— PRIKU 6117; Kurouchi— PRIKU 6540; Kuwasononan- bu— PRIKU Coll. Nos. 32-35; Maki— PRIKU Coll. No. 118; Masugata— PRIKU 6464, Coll. Nos. 173, 184; Metaka— PRIKU 6460; Minami- machi— PRIKU Coll. No. 123; Misokawa— PRI- KU 6564, 6565; Miyamoto— PRIKU Coll. Nos. 92, 97; Miyamura— PRIKU 6485, 6570, 6611, Coll. Nos. 174, 185; Miyashiro— PRIKU 6121, 6179, 6339, 6458 (cranial and mandibular frag- ments, postcranials), 6617 (cranial and mandibu- lar fragments), 6673; Miyashita— PRIKU 6207; Motomura— PRIKU 6459; Nakadachi— PRIKU 6615; Nakahora— PRIKU Coll. No. 94; Nakasu- PRIKU 6322 (mandibular fragments, postcrani- als); Nakayama— PRIKU 6796, Coll. No. 176; Nezumiana— PRIKU 6104, 6344; Nishi— PRIKU 6801; Nishiharuchika— PRIKU 6122, 6174, 6182 (cranial fragments, postcranials), 6186 (cranial fragments, mandible, postcranials), 6341 (cranial fragments), 6455, 6468, 6484, 6671 (cranial frag- ments), 6805, Coll. No. 134; Ochiai— PRIKU 6465, 6804; Ogikubo— PRIKU 6111, 6563, 6616 (cranial fragments); Oobara — PRIKU 6824; Ooji- masan— PRIKU 61 16, 6210; Ookawara— PRIKU 6213; Oosaki— PRIKU Coll. Nos. 188, 189; Sa- musawa — PRIKU 61 10, 6172 (cranial fragments), 6173, 6181 (cranial fragments, mandible, post- cranials), 6187 (cranial fragments, mandible, post- cranials), 6195 (cranial fragments, mandible, post- cranials), 6196 (cranial fragments, mandible, post- cranials), 6467, 6569 (note: external measure- ments recorded for PRIKU 6187 and 6467 [Coll. Nos. 98 and 99] appear to be mismatched with the respective skeletons); Seto — PRIKU 6102, 106 FIELDIANA: ZOOLOGY 6212, 6456, 6618; Shiga-kogen— PRIKU 2141, 2142 (mandible lacking); Shimashima— PRIKU 6171, 6175, 6332, 6476, 6486; Shimizu— PRIKU 6166 (cranial and mandibular fragments), 6471, 6812 (cranial fragments), 6814; Shimoichiba — PRIKU Coll. No. 71; Shimoseinaiji— PRIKU 6472, 6601, Coll. Nos. 175, 177; Shinyashiki— PRIKU Coll. No. 23; Shiono— PRIKU 6614, Coll. No. 182; Shouzawa— PRIKU 6479; Suza- wa— PRIKU 6482 (cranial fragments); Takai— PRIKU 6125, 6604, 6818, Coll. No. 180; Taki- goshi— PRIKU 6479, 6922; Takinozawa— PRIKU 6794; Tatai— PRIKU 6531, 6648, 6649; Taza- wa— PRIKU 6184, 6190, 6191 (cranial fragments, mandible, postcranials). Coll. Nos. 190-193; Ten- jinbara— PRIKU 6119, 6178, 6473, 6567, 6813; Tokiwa— PRIKU 6331, 6612, 6613; Tsukiyo- daira— PRIKU 6097, 6103, 6107, 6109, 6112, 6118, 6120, 6126, 6164, 6169, 6185, 6342, 6352, 6353; Ushimaki— PRIKU 6096, 6101, 6108, 6114, 6170, 6183 (cranial fragments, mandible, postcranials), 6188 (cranial fragments, mandible, postcranials), 6193, 6198, 6205, 6340, 6350, 6457, 6462, 6475, 6539, 6672 (cranial fragments), 6795, 6802, 6803, 6807-681 1, 6815, 6819, 6822; Wakahowatauchi— PRIKU 6825; Wakamiya- Hachimanjinja— PRIKU Coll. No. 130; Warabi- taira— PRIKU 6489; Yakeyama— PRIKU 6797 (cranial fragments), 6798 (cranial fragments), 6816 (cranial fragments), 6821 (cranial frag- ments), 6826 (cranial fragments); Yamabuki — PRIKU 6481 (skull damaged); Yanagisawa— PRI- KU Coll. No. 103; locality unknown, "Shinshiu" (= Nagano Prefecture)— USNM 21116. Niigata (5): Komogawa— PRIKU 6541, 6602, Coll. No. Niigata 1 ; Takane— PRIKU Coll. Nos. Niigata 2- 3. Osaka (3): Minoo-city— PRIKU 277, 616, 617. Shiga (1): Tsuchiyama-cho, Koka-gun [2] — TPM (unnumbered). Shimane (156): Akashi — PRIKU 5721; Fukano— PRIKU 5149, 5179 (cranial frag- ments, postcranials); Hasumi — PRIKU Coll. No. Shimane 7; Higashihinobori— PRIKU 5154, 5176, 5178 (postcranials only); Hinagawa — PRI- KU 6525, 6620, 6621, Coll. No. Shimane 10; Hi- rasa— 561 1, 571 1 (postcranials only), 5713-5715; Houin— PRIKU 5142 (cranial fragments), 5177, 5180; Imanishi— 5871, 5875, 5877, 5884 (cranial fragments), 6131, 6138; Coll. No. Shimane 8; Koujiro — PRIKU 5141 (cranial fragments, post- cranials), 5143 and 5148 (postcranials only); Mik- usu— PRIKU 5152, 5153, 5155, 5610: Mitoya- cho— PRIKU 5612, 5710 (cranial fragments), 5739, 5867, 5880; Nishihinobori— PRIKU 5145, 5735; Okuyama— PRIKU 5741 (female, misla- beled as male); Shimogou — PRIKU 5136 (po- stcranials only), 5137, 5138, 5139 (postcranials only), 5140, 5144, 5147, 5156, and 5238 (4 spec- imens, postcranials only), 5239 (cranial frag- ments, postcranials), 5240 (postcranials only), 5712, 5716, 5720, 5827 (postcranials only), 5859-5863, 5865, 5868-5870, 5878, 5883, 5885, 5886, 5889, 5890, 5892, 5893, 5895, 5980 (cra- nial fragments), 6142, 6152, 6160, 6351; Taku- wa— PRIKU 5151, 5708 (mandible broken), 5709, 5722, 5881 (cranial and mandible frag- ments), 6132, 6143, 6529; Tsukinoya— PRIKU 5146, 5150 (mandible and postcranials only), 5241 (cranial fragments, postcranials); Ueda — PRIKU 5864, 5872-5874, 5882, 5887, 5888, 5891, 6127, 6128, 6130, 6133-6137, 6139, 6146, 6148-6150, 6153-6155, 6158, 6161, 6438, 6526, 6527; Ushiroyama— 5738, 5876, 5894, 5979, 6140, 6141, 6144, 6145, 6147, 6151, 6156, 6157, 6159, 6162, 6528, Coll. Nos. Shimane 1-6, 9, 1 1- 24; Yoshida-mura— PRIKU 5866. Shizuoka (5): Hagachizaki— PRIKU 1381; Hamakita-city— PRIKU 1376, 1377, 1380; locality unknown— PRIKU 6670. Tochigi (39): Imaichi-city— TPM 59/171, 61/074, 61/118-61/121, 61/123, 61/132; Kuriyama-mura — TPM 61/090, unnumbered skull (mandible lacking); Nikko-cityTPM 56/083-56/ 086, 57/095, 57/178, 58/067, 58/071, 59/135, 59/ 170, 60/101, 60/102 (mandible lacking), 60/106, 60/107, 60/246, 61/067, 61/072, 61/080, 61/091, 61/092, 61/1 10, ZMB Coll. Nos. III-X. Prefecture unknown (56): "Jedo'V'Yedo" = Tokyo — ZMB A3469, A3470, A3476, A3481, 5982, unnum- bered female skeleton acquired 13 Feb. 1874; "Jokohama" = Yokohama-city (cf. Schweyer, 1909, p. 8)— AIUM 3472-3480, 3482-3487, 3489, 3490, 3492-3506; "Tokyo Market"— USNM 13830, 13831, 20937; "Yokohama"— ZSBS 1902/97; locality unknown— ZSBS 1901/ 69, 1901/71-1901/83. KINKAZAN, 15. Miyagi (15): MUE K/1, K/2, K/6, K/lO-K/15, K/20, K/24, K/28, K/33, K/Yl, K/Y6. KOJIMA, 47. Miyazaki (47): KFL 1, 2 (crani- um broken), 3-9, 10 (mandible lacking), 11, 12, 13-15 (mandibles lacking), 16, 17 (mandible lacking), 18, 19 (mandible lacking), 20, 21, 22 (mandible lacking), 23 (cranial fragments), 24, 25 (mandible lacking), 26, 28 (cranial fragments), 29-34, 35 (cranial fragments), 36-45, "Usagi" (name of monkey), unnumbered skull (mandible lacking), unnumbered mandible only. KYUSHU, 18. Kagoshima (5): Kanayama— PRIKU, five uncatalogued skulls. Miyazaki (1): FOODEN AND AIMI: SYSTEMATIC REVIEW OF J7\PANESE MACAQUES 107 Toimisaki— JMC 1747. Oita (12): Takasakiya- ma.— JMC 1949. 3971. PRIKU 2131. 2133. 2143. 2146. 2152. 2154 (mandible and postcranials only). 2155-2158. SHODOSHIMA. 47. Kagawa (47): JMC 925. PRIKU 1405-1409. 1410 (mandible only). 1411 (mandible and ulna only). 1412 (mandible and postcranial fragments only). 1413-1421. 1429 (mandible and postcranials only). 1431, 1432 (cranial fragments). 1433 [1]. 1433 [2]. 1434. 1438 [1]. 1438 [2]. 1438 [3]. 1444. 1446-1448. 1450 [1], 1450 [2], 1455, 1456 (mandible and po- stcranials only). 1457 (mandible and postcranials only). 1458-1460. 1464-1466. 1468. 1470. 1472- 1474. Island unknown, 45. AIUZ 2063. 6064. AMNH 35640. BM(NH) 1850.8.15.2. NHR Z2266. Z2267. PRIKU 4715. 4717. and 4718 (3 speci- mens, postcranials only). 5879 (juvenile skull mistakenly associated with data pertaining to a large male). 6215. 6216. 6218. 6303-6305^ 6306 (cranium broken). 6307-6312. 6467 (adult female mistakenly associated with data pertaining to an infant male), unnumbered skull (with note refer- ring to PRIKU 5139 and PRIKU 5140). RMNH 8-e. 9-f. 10-g. 11-h. 12-a. 13-b. USNM 259982. 260229, 344995. ZMB 12957. 25547. 25548. 25550. 25552-25554, RexuCo, unnumbered male skeleton acquired in 1874. ZMUZ 1 1603a. ZSBS. unnumbered male skull. Macaca fuscata yakui, Total 182 Skins and Skulls, 5 YAKUSHIMA. 5. Kagoshima (5): locality un- known—BM(NH) 1905.1 1.3.1-1905.1 1.3.5. Skulls or Skeletons Only, 177 YAKUSHIMA, 177. Kagoshima (177): Hany- ama— PRIKU 1571. 1572. 1729. 1730. 1731 (mandible lacking). 1732 (mandible lacking). 1733. 1749. 3430,^3957-3959, 4395 (cranial frag- ment). 4490. 4491. 5424 (skull broken). 5425, 6626. 6627 (mandible and postcranials only), 6628-6630. 6631 (mandible lacking). 6632, 6633. 6634 (mandible and postcranials only). 6635, 6636 (mandible fragment only), 6637-6642. 6751; Hinokuchi— PRIKU 3153 (male skeleton with mismatched female mandible). 3154-3157; Issou— PRIKU 3158; Kamiyaku-cho (30°16'N)— PRIKU 2148, 3433-3446; Kojima— PRIKU 2610 (cranial fragments, mandible, postcranials) ; Kun- iwaridake — PRIKU 2167 (cranial fragments, mandible, postcranials; Kusugawa — PRIKU 2595: Miyanoura— PRIKU 2147 (mandible lack- ing); Mugio— PRIKU 2588. 2591. 2594. 2607, 2609. and 2611-2615 (7 specimens, postcranials only); Nagata — PRIKU 2168 (mandible lacking). 2585-2587. 2608 (postcranials only). 3159-3162; Nakama— PRIKU 2593. 2597-2605. 2616-2618 (3 specimens, postcranials only); Nakase — PRI- KU 2606 (postcranials only); Nakasegawa — PRI- KU 2584. 2589. 2590; Seibu-rindou— PRIKU 3351 (mandible lacking). 3352 (cranial and man- dibular fragments): Shichigodake— PRIKU 2592. 2596; Shirakoyama— PRIKU 3141-3144. 3145 (postcranials only). 3146. 3147. 3148 [1] (infant skeleton with mismatched juvenile mandible), 3148 [2] (aduh skeleton, skull damaged). 3149, 3150 [1], 3150 [2], 3150 [3]. 3151 (cranial frag- ments). 3152 [1] (skull damaged). 3152 [2], 3152 [3]. 3152 [4], 3152 [5], 3451-3453; Yudomari— PRIKU 3123-3131. 3132 (postcranials only), 3133-3140; locality unknown— JMC 601, PRIKU 3447-3450. 3454-3456. 3457 (cranial fragments), 4120 (cranial fragment and mandible only). 41 21- 4124, 4459 (cranial fragment), 4460 (skull dam- aged), 4461, 4462, 4463. and 4720 (3 specimens, postcranials only). 4941. 5009. 5227, 5247-5249, 5373 (postcranials only). 5416. 6083-6086. 6951, 6952, 6953 (cranial fragments, postcranials), 6954. 6955. Appendix 2: Macaca fuscata Localities Reported in Postal Survey Conducted by K. Hasebe in 1923 (Fig. 2A; cf. Iwano, 1974, p. 5; Uehara & Koganezawa, 1976, p. 742; Amagasa & Ito, 1978, p.96); for Locality Details, see Gazetteer, Appendix 4. YAKUSHIMA: Kagoshima Prefecture— Yakxi- shimayama. TANEGASHIMA: Kagoshima Prefecture— Fu- rutayama et al.; Masudayama. KOJIMA: Miyazald Prefecture — Kojima. KYUSHU: Kagoshima Prefecture — Bono-mi- saki; Shimoyama; Shibisan; Shiraki; Harimochi; Kirishimasanrin; Onomi and Otaniyama; Takaku- madake, Kanoye-cho; Kamimobiki vicinity; Taka- kumadake; Takakuma; Kunimidaira vicinity; Ina- 108 FIELDIANA: ZOOLOGY odake vicinity; Sarutsubo vicinity. Miyazaki Pre- fecture— Misaki-Makinouchi vicinity; Ariki Na- tional Forest vicinity; Kirishimayama, Nishidake-son; Kirishimayama vicinity, Takahara- son; Ohira National Forest vicinity; National For- est, Suki-son; Ichifusa vicinity; Tatsufusayama vi- cinity; Fukiyama National Forest; Kanasumi Na- tional Forest, Kijo-son; Osuzu National Forest; Tedayama vicinity; Nishinoyakaeyama; Totoro- gauchi vicinity, Kadokawason; Okumiyama; To- torogauchi, Kitagouson; Mukabakiyama; Hinoki- yama; Ookueyama vicinity; Sokujitsunomine; Na- misegawara; Tsuriganeyama; Higakureyama vi- cinity; Oshioi; Yumigiyama. Oita Prefecture — Mumeinoyama, Onoichi-mura; Mumeinoyama, Shigeoka-mura; Odairayama vicinity; Katamuki- yama; Mitakeyama; Honjoyama; Mumeinoyama, Imaichi-mura; Mumeinoyama, Kawanobori-mura; Taninoshiriyama; Inouesanrin; Onagarayama For- est; Shihatsuyama; Shikakumadake, Shimogou- mura; Shikakumadake, Yamautsurui-mura; Shigi- ra Shinyabakei; Hikosanroku; Ota; Tano; Futago- yama; Higashi- and Nishi-Kurokiyama. Kuma- moto Prefecture — Nishiohira; Gongenyama vicinity; Shiroiwayama; Yamate vicinity; Shira- kiyama vicinity; Sakasegawa vicinity; Kosadake; Fukabayama, Fukuoka Prefecture — Kusenbuya- ma vicinity; Kawaradake, Kawara-machi; Egawa vicinity. KASHIMA: Ehime Prefecture — Kashima. SHIKOKU: Ehime Prefecture — Aikawayama; Shigekiyama; Masakiyama; Sasagou vicinity; Shi- monaru; Kurase vicinity; Tataki vicinity; Higashi- mikatagamori; Nakao; Kitamine; Amagoitaki wa- terfall vicinity; Kuragariyama; Tatsukawayama; Daieiyama; Tanegawayama vicinity; Fujigaishi vi- cinity; Higashinokawa vicinity; Somakawa-mura; Gayatani vicinity; Ibushikanyama vicinity; Koya- yama; Ondayama vicinity; Akago; Umegatani vi- cinity; Ryuoka-mura National Forest; Okuyama. Kochi Prefecture — Kashiwajima; Souro vicinity; Onozumi vicinity; Kurokawa; Kunimi vicinity; Nakahama vicinity; Tsuro vicinity; Shimonokae vicinity; Kujuu vicinity; Yamada; Kyouhou vicin- ity; Fujikawa vicinity; Nagaoi; Tsuneroku vicini- ty; Nakatsugawa vicinity; Nonogawa vicinity; Kannonyama; Nadanishiyama vicinity; Nadayama vicinity; Matsubakawayama vicinity; Amagoiya- ma; Tachibanaseiryuu National Forest; Tsubaya- ma; Yasui National Forest; Nakakurayama; Taka- bagamori; Kirakuyama vicinity; Seto; Kiyano vi- cinity; Minamikawa vicinity; Tochu vicinity; Shi- mokawaba vicinity; Shiragayama; Ouyama vicinity; Nishitoyonaga-mura; Makitaniyama vi- cinity; Okugataniyama; Chaen; Nonesankei; Mo- toyama; Oozakurayama; Yokootsuzukiyama vi- cinity; Sugawayama vicinity, Iketaniyama vicini- ty; Kanegaryumorisankei; Iwataniyama. Tokushi- ma Prefecture — Kushigawa vicinity; Hirai vicinity; Kitousanrin, Okukitou-son; Kitousanrin, Kamikitou-son; Shindenyama; Kigiura vicinity; Senbagatake vicinity; Akamatsu; Kawachi vicin- ity; Nadayama; Gongendaki vicinity; Kakeban vi- cinity; Fukuhara-son; Yanagitani; Okutachikawa vicinity; Shikakubidani; Ooboke; Tokuzen-nishi vicinity; Motoi vicinity; Kashiji vicinity; Kuchi- yama-son; Iwaya; Handairayama-son; Yuumaya- ma; Kawatayama vicinity; Koyadaira; Kidooku; Nagafuka vicinity; Hanto-cho; Kitanada-son; Gosho-son. Kagawa Prefecture — Yamawakiya- ma; Higashiyama National Forest; Asan Mountain Range. SHODOSHIMA: Kagawa Prefecture— Tmma- san; Kensozan, Oonude-mura; Kensozan, Kitaura- mura; Kensozan, Oobe-mura; Kensozan, Kusaka- be-mura; Kensozan, Yasuda-mura. OMISHIMA: Yamaguchi Prefecture — Omishi- ma, Senzaki-cho. AWAJISHIMA: Hyogo Prefecture— Yuzuru- hayama; Kashiwabarayama foothills. HONSHU: Yamaguchi Prefecture — Onigajo; Gezan; Sarugababa; Shirataki; Nagatokyo vicini- ty, Kawakami-son; Nagatokyo vicinity, Ikumo- son; Nagatokyo vicinity, Shinobu-son; Hagi-ma- chi; Northern villages, Tsuno-gun; Kuratani vicin- ity; Kuroda vicinity. Hiroshima Prefecture — Go- kurakujisanrin; Nenbutsudani National Forest; Onosanrin; Masakiyama; Yokogawaburaku vicin- ity; Karioyama vicinity; Takiyamagawa vicinity; Ryuuzuyama; Samuhikiyama; Kabetouge vicinity, Honji-son; Kabetouge vicinity, Minamikata-son; Doutokoyama; Hongushisan; Shirokiyama vicini- ty; Takayama; National Forest, Nishishiwa-son; Shirokiyama; Takanosuyama, Kuba-son; Takano- suyama, Takeni-son; Takamurayama, Nyuno-son; Youkurayama, Zen-nyuuji-son; Youkurayama, Kamikitakata-son; Youkurayama, Funaki-son; Youkurayama, Ookawa-son; Buttsujisan; Koni- shioku; Ootaniyama vicinity; Ryuozan, Miyauchi; Kugayama National Forest vicinity; Dakeyama; Tarekaya Natioanl Forest; Iwamiyama vicinity; Hizaoyama; Taishakugawa vicinity; Inyou Moun- tain Range vicinity, Kimita-son; Inyou Mountain Range vicinity, Funo-son; Inyou Mountain Range vicinity, Sakugi-son. Okayama Prefecture — You- zenotaki; Abeyama; Kanehirayama; Tenjinyama; Sarugamiyama; Tsuchiosanrin vicinity; Furutani National Forest; Douyama; Kamikumatani, Ku- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 109 matani-son; Karamatsu; Akoumayama; Kanba; Yumoto; Gagyusan, Tsugawa-son; Gagyusan, Takahashi-cho; Gagyusan, Matsuyama-son; Ita- mayama; Kuniyama National Forest; Takiyama; Okutsugawa National Forest. Shimane Prefec- ture— Kaneyamadani; Tachigouchi; Tadeno; Ka- mitakashiri; Kabatani; Shiroyama, Hatagasako- mura; Shiroyama, Tsuwano-cho; Tadachiokuya- ma; Yokomichi vicinity; Takitani; Sumikawa vi- cinity; Rakanyama, Hiromi; Kametaniyama vicinity; Itotaniyama; Kuigautsu, Hinui-mura; Maruseyama; Kirigataniyama vicinity; Iwaidani- yama; Kanmuriyama; Kurokagara vicinity; Ooe Takayama; Soratani; Hamahara-mura; Akutaya- ma; Tsunotani; Kogarashigataniyama vicinity; Ooyahongou; Yashiroyama, Yashiro-mura; Yata- kiyama vicinity, Nishida, Yusato-mura; Ginzan vicinity; Oshihara-mura; Hata-mura; Matsukasa- mura; Yoshida-mura; Tai-mura. Tottori Prefec- ture— Okinoyama; Momobayama; Iwayadou, Ike- da-son. Hyogo Prefecture — Kusaki-mura; Miu- chisanrin, Ishii-mura; Funakoshiyama; Kawaraya- ma; Nabegatani; Akanishi vicinity; Rengeiwa vicinity; Kayano vicinity; Kishida; Yukihikoya- ma; Nagakura vicinity, Hase-mura; Kurataniyama, Tataragi, Nakagawa-mura; Kanatokoyama, Itoi- mura; Myomisan; Nishitokoo vicinity; Kanato- koyama, Aihashi-mura; Mikuniyama; Houtaku- san; Egasayama; Sasagamine; Atagoyama vicini- ty; Akaioku, Nunuki-mura; Ishidosan; Mitsudake; Ikuno, Doujou-mura; Kitayama, Shioze-mura; Mayasan. Osaka Prefecture — Minoosan National Forest; Hattasan vicinity; Katsuojisan; Shakaga- dake; Ushitakisan; Inunakisan. Wakayama Prefec- ture— Onoueyama; Ui, Ishigaki-mura; Otayama; Nikkosan; Nakaodani vicinity, Go-mura; Muro- kawayama; Kashiyama vicinity; Kawanaka-mura; Hidaka Mountain Range; Kawamatasanrin; Shi- mosanji-mura; Kamisanji-mura; Higashiyama vi- cinity, Kamihaya-mura; Ryujinyama vicinity; Takaosan vicinty, Nagano-mura; Toragamine Mountain Range vicinity; Mochihirasan; Takao- san vicinity, Chikano-mura; Ryujinyama vicinity, Inari-mura; Ryujinyama vicinity, Kamiakitsu- mura; Jujotouge vicinity; Mugikomoriyama vicin- ity; Damasanchi, Ichinose-mura; Ashiyama vicin- ity, Ikuma-mura; Shokawagoe vicinity; Hirutani vicinity; Kongoumine vicinity; Endani vicinity; Mimaisan vicinity; Futagoyama vicinity; Nakano- goudani vicinity; Ebitadani vicinity; Shimizuyama vicinity; Mureyama vicinity; Shigatakiyama vi- cinity; Arida-mura/Higashimuro-gun boundary; Fujihashi-mura/Higashimuro-gun boundary; Nyu- douzan vicinity; Miokawa-mura; Shichikawa- mura; Ogawa-mura; Takaike-mura; Hachirouya- ma; Kamioota-mura; Mizunoshitayama; Irokawa- mura; Takata-mura; Kitanokawa vicinity; Yo- mura; Sunomata vicinity, Ukegawa-mura; Karekidani, Shikiya-mura; Mikoshi vicinity, Mi- sato-mura; Kujuu-mura; Tamakikuchi-mura. Nara Prefecture — Kitamatasanrin, Nosegawa-mura; Akataniyama; Hirosesanrin, Tenkawa-mura; To- tsukawa-mura; Shimokitayama-mura; Kamikita- yama-mura; Kawakami-mura; Mitsue-mura; Beni- gadake vicinity; Taishiyama, Murou-mura; Taka- toriyama National Forest, Takatori-cho; Takatori- yama National Forest, Takaichi-mura. Kyoto Prefecture — Konoshiro, Takano-mura; Oyama vi- cinity; Choenzan; Taikoyama; Ookaneyama; Houzawayama; Koryujiyama; Akaiwayama, Oka- danaka-mura; Akaiwa Mountain Range, Maruyae- mura; Kagamisakadani, Oono-mura; Sumugitani, Miyajima-mura; Horatani vicinity, Tsurugaoka- mura; Ashiu vicinity, Chii-mura; Hacchoyama; Kumamidani vicinity, Kuroda-mura; Takitani vi- cinity, Kuda-mura; Chishorodani; Urayama, Shi- no-mura; Arashiyama vicinity; Yanagitani, Kaiin- ji-mura; Makiosan vicinity; Ujitahara-mura. Mie Prefecture — Tadosan vicinity, Komi-mura; Tado- san vicinity, Tado-mura; Tashida vicinity. Hatta- mura; Toriidoyama; Yunoyama; Kanmuriyama; Nyudougatake; Sengatake; Nonoboriyama vicini- ty; Nakatsugawayama; Kabutoyama; Shakujou- gadake; Hiraki vicinity, Nagano-mura; Takarajou vicinity, Awa-mura; Shorenjisan vicinity; Naga- saka-mukaiyama; Iwayama, Tarou-mura; Ohara- dake; Idodani vicinity; Hirakura, Yahata-mura; Kuchisubosanrin; Awano, Kawamata-mura; Hachisusanrin vicinity, Mori-mura; Miyama Na- tional Forest, Ookawachi-mura; Shimatani vicin- ity, Ryounai-mura; Aritani, Hagiwara-mura; Oo- sugitani-mura; Oouchiyama-mura; Kasagi, Kashi- wazaki-mura; Kurosu vicinity, Shimazu-mura; Fujikiya forest, Nanaho-mura; Ishigagawa, Naka- gawa-mura; Goriyama, Ogawagyou-mura; Higa- shimiyamata vicinity, Ichinose-mura; Oshibuchi vicinity, Hobara-mura; Michikata vicinity, Naka- jima-mura; Oitsuge vicinity, Nishiki-mura; Nisa- kasanrin vicinity; Tochiyama vicinity; Shimaka- chisanrin vicinity; Ooike beach; Ookawachiyama vicinity; Iwaidani vicinity; Wataka vicinity, Owa- se-cho; Higashimisaki vicinity; Morimatsu vicin- ity, Kitawauchi-mura; Sone vicinity, Minamiwau- chi-mura; Komataoku vicinity, Asuka-mura; Mo- mozaki vicinity, Isato-mura; Ogawaoku vicinity, Kamikawa-mura; Komori, Nishiyama-mura; Oomine vicinity; Wake vicinity, Kamikawa-mura; Oosato, Onodani-mura; Asaoki, Mifune-mura; 110 FIELDIANA: ZOOLOGY Matani, Agira-mura. Shiga Prefecture — Iwamaya- ma; Hieizan; Hirasan vicinity, Kido-mura; Hira- san vicinity, Katsuragawa-mura; Komatsu-mura; Tsunogawa-Akana vicinity; Fukashimizu vicinity, Kawakami-mura; Hidarimaeyama vicinity; Cho- shigatani; Kataoka-mura; Niu-mura; Sugino- mura; Ibukiyama, Ibuki-mura; Ibukiyama, Suijou- mura; Ryozenyama, Kashiwabara-mura; Ibukiya- ma, Samegai-mura; Ryozenyama, Toriimoto- mura; Ryozen, Seritani-mura; Wakigahata-mura; Takidani vicinity, Ootaki-mura; Higashiogura- mura; Akasugiha vicinity, Yamakami-mura; Oo- gawara, Aiga-mura; Kurotaki, Yamauchi-mura; Tomikawayama. Fukui Prefecture — Aonogou- mura; Takahama-cho; Saburi-mura; Wada-mura; Hongou-mura; Ooshima-mura; Kado-mura; Hitot- sudai, Okunata-mura; Nishiogawa vicinity, Uchi- tomi-mura; Ayuhanasanrin, Nishida-mura; Mika- tasanrin, Ya-mura; Shinjou vicinity, Mimi-mura; Kurokawayama; Okuaso vicinity, Arachi-mura; Takasuyama; Ashimi-mura; Ohara vicinity, Kita- tani-mura; Gokairiaiyama, Kamishou-mura; Ni- gure, Kamianama-mura. Ishikawa Prefecture — Kuratani vicinity, Saigawa-mura; Okuike vicinity, Kawachi-mura; Jadani vicinity, Yoshinotani- mura; Hidaritsubute vicinity, Torigoe-mura; Ozoe- chinai, Okuchi-mura; Kuwashimachinai, Shira- mine-mura. Aichi Prefecture — Hongusan, Ichino- miya-mura; Toyamagoryouchi vicinity, Nagashi- no-mura; Ebi-cho; Kandayama; Myojinyama; Ooiriyama, Sono-mura; Yatsudake vicinity; Ha- guro-mura; Kurisuyama; Hongusan, Gakuden- mura; Owarifuji vicinity. Gifu Prefecture — Tado Mountain Range, Ishizu-mura; Tado Mountain Range, Shiroyama-mura; Tokiyama; Oobora-Sa- tani, Tara-mura; Shogatake vicinity; Ootani, Ma- kita-mura; Shimonotani vicinity, Sakauchi-mura; Shiratani vicinity; Kouchi, Yokokura-mura; Ooka- wara vicinity, Neo-mura; Kinbaradani, Toyama- mura; Kamagatani, Kamiijira-mura; Okuookura vicinity, Kuzuhara-mura; Natsusaka vicinity, Ki- tayama-mura; Tarudaira vicinity, Horado-mura; Kawaurayama, Itadori-mura; Shimoyama vicinity, Shimomaki-mura; Kouwayama vicinity. Suhara- mura; Akakuzureyama, Shimonoho-mura; Otsu- kamebora, Nakanoho-mura; Hatofukiyama; Fu- kubegadake; Nahisanrin, Aioi-mura; Konahisan- rin vicinity, Nishiwara-mura; Negitani, Nishika- wa-mura; Inunaki, Kuchimyougata-mura; Gozenyama; Sangenyama; Senchougahara; Hige- tayama vicinity, Takane-mura; Byobuiwa, Kado- hara; Kanakidoyama; Fukatani vicinity, Yama- nokuchi-mura; Jigokudani vicinity, Siioukawa- mura; Aratani vicinity, Shirakawa-mura. Toyama Prefecture — Mizunashiyama vicinity; Nagato- okuyama, Fukusawa-mura; Takasugi vicinity, Ooyama-mura; Onigajo vicinity; Nabemashi vi- cinity, Shirahagi-mura; Kakumahachi, Matsukura- mura; Sangayama; Kurobedani; Aimotosanchi vi- cinity, Aimoto-mura; Ogawadani, Yamazaki- mura; Taiheisanrin, Sakai-mura. Shizuoka Prefec- ture— Senzu-goryourin, Kamikawane-mura; Fujikawayama; Hakkouzan, Shimokawane-mura; Hakkouzan, Goka-mura; Sekinosawa, Ikawa- mura; Yakeyama vicinity; Utougi, Ookawauchi- mura; Yunodake; Uchiyama, Yoshinaga-mura; Sudoyama vicinity, Sudo-mura; Aitakayama; Kin- tokiyama; Ita, Heda-mura; Amagisan, Kitou- mura; Amagisan, Inatori-cho; Amagisan, Kami- kawatsu-mura; Amagisan, Nishina-mura; Iwashi- na-mura; Mihama-mura. Nagano Prefecture — Na- gisoyama, Azuma-mura; Nagisoyama vicinity, Yomikaki-mura; Tadachi-goryorin; Inagawa-go- ryorin; Komagatake Mountain Range; Miurayama vicinity; Mitakeyama-goryorin; Komagane Moun- tain Range, Hiyoshi-mura; Misogawa-goryorin vi- cinity, Kiso-mura; Azumi National Forest; Mina- mikurosawa National Forest, Azusa-mura; Kara- sugawa National Forest; Karasugawa, Nishihoda- ka-mura; Kurosawa National Forest, Ogura-mura; Nakabusa National Forest, Ariake-mura; National Forest, Matsukawa-mura; National Forest, Toki- wa-mura; National Forest, Taira-mura; National Forest, Kamishiro-mura; National Forest, Hokujo- mura; National Forest, Nakatsuchi-mura; Seto, Kinasa-mura; Togakushiyama vicinity; Hotokei- wa, Hirao-mura; Yamada-mura; Shiranesan vicin- ity; Chichiyama, Toyooka-mura; Nireyama vicin- ity, Nire-mura; Kakuma National Forest, Osa- mura; Yatsugatake National Forest, Kitamaki- mura; National Forest, Minamiaiki-mura; Yatsugatake National Forest, Minamimaki-mura; Chichibu Mountain Range; Kamanashiyama; Ni- shikomagadake, Ina-cho; Fudoutaki; Minamiko- magadake; Mitsuminegawa, upstream, Inasato- mura; Akaishisan vicinity, Ooshika-mura; Nishi- fujiyama, Kami-mura; Kamisawayama, Kisawa- mura; Nishiyama, Wada-mura; Nishiyama, Minamiwada-mura; Kumabushiyama; Enasan; Ai- kawautsubo, Namiai-mura. Niigata Prefecture — Komagatake vicinity; Koide, Kuramata-mura; Mukaiyama, Mitsumata-mura; Mantaroyama vi- cinity, Tsuchidaru-mura; Ushigadake, Ikazawa- mura; Hakkaisan, Jounai-mura; Hakkaisan, Higa- shi-mura; Daikokuyama vicinity; Akagawaomote, Yunotani-mura; Asakusayama vicinity, Irihirose- mura; Kurohimeyama, Ukawa-mura; Komeyama, Noda-mura; Kurohimeyama, Takayanagi-mura; FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 111 Awagatake; Kawachiyama, Kawachi-mura; Mika- gura, Nishikawa-mura; Zoukiyama, Higashikawa- mura; lidesan vicinity, Toyomi-mura; Nagahashiri vicinity, Hideya-mura; Akataniyama, Akatani- mura; Ooishifukagawairi vicinity, Sekitani-mura; Kamikaifu-mura; Komagatake vicinity, Takane- mura; Wasedasuirin vicinity, Shionomachi-mura; Fujisawa vicinity, Shimokaifu-mura; Kumadaya- ma, Nakamata-mura. Yamanashi Prefecture — Ji- zoutouge vicinity; Tsukiyonodan; Furuyajo, To- yooka-mura; Takatoriyama; Shichimensanroku, Motodate-mura; Zarugadake vicinity; Daikoku- yama; Bettou, Misato-mura; Daitouzan vicinity; Okusenju, Hirabayashi-mura; Norogawairi, Ashi- yasu-mura; Kinpusan vicinity, Miyamoto-mura; Kanegatake vicinity, Kiyokawa-mura; Gozaishi- yama vicinity, Seitetsu-mura; Gozaishi, Maruno- mura; Ohsawakanrin, Hourai-mura; Komagatake Mountain Range; Obi, Masutomi-mura; Namesa- wayama, Mitomi-mura; Shougenyama; Tokusa. Hagiwara, Oofuji-mura; Ushiokuyama vicinity, Okunoda-mura; Ootsuneki, Tabayama-mura; Ko- ganezawa vicinity, Nanaho-mura; Saihara-mura; Shimohatsugari, Hatsugari-mura; Okuno, Sasago- mura; Takara-mura; Mitsutouge, Nishikatsura- mura; Doushi-mura; Junigatouge. Nagahama- mura; Oudake; Amegatake; Tojiroyama; Tsuba- kiyama vicinity, Ookouchi-mura; Sano-goryorin, Sakae-mura. Tokyo Metropolis — Kanotoyama vi- cinity; Iriokuyama vicinity; Tarusawayama vicin- ity; Bonhoriyama. Kanagawa Prefecture — Ashi- gara Mountain Range; Kintokiyama vicinity; Bukkazan; Kanazawayama vicinity; Hirugadake. Chiba Prefecture — Takagoyama; Shimizusawa, Otaki, Toyooka-mura; National Forest vicinity, Kameyama-mura; Tsutsumori National Forest, Oi- kawa-mura. Saitama Prefecture — Ryogamisan; Ooborayama; Hinoyama; Arimayama; Bukouzan, Kagemori-mura; Bukouzan, Yokoze-mura. Ihar- aki Prefecture — Tsukubasan. Tochigi Prefecture — Konan National Forest vicinity, Ashio-machi; Fu- kazawa National Forest, Nikko-machi; Fujimiya- ma vicinity; Tsukiyama vicinity; Ogadake; Taka- harasan vicinity; Oosabiyama vicinity. Gunma Prefecture — Hondani National Forest vicinity, Ueno-mura; Hoshio National Forest vicinity, Oza- wa-mura; Nishinomaki, Saimoku-mura; Myogisan National Forest; Nakakimura; Maruyama, Ubu- chi-mura; Manzayama, Tsumagoi-mura; Yokoka- be National Forest vicinity, Naganohara-mura; Iri- yamagawaura National Forest, Kuni-mura; Dai- gentasan; Bunajurin, Minakami-mura; Hotakaya- ma vicinity, Kawaba-mura; Hotakayama vicinity, Katashina-mura; Hirakawa vicinity, Azuma-mura; Akagiyama vicinity, Akagine-mura; Akagiyama, Kurohone-mura. Fukushima Prefecture — Futama- tayama, Yumoto-mura; Kasshizan, Nishigou- mura; Yaheishiroburaku, Okugawa-mura; Jinku- rousanrin-oku, Hibara-mura; Azumasan, Azuma- mura; Shinzan, Mizuho-mura; Moniwa-mura; Ryozen, Ryozen-mura; Asahidake, Yamakiya- mura; Hiyama, Asahi-mura. Yamagata Prefec- ture— Azumayama, Minamihara-mura; Nishine, Kitaokuni-mura; Fukuei-mura; Rokujuri. Hongou- mura; Rokujuri, Azuma-mura; Shinzan adjacent to Mogami-gun, Oosawa-mura; Shinzan adjacent to Mogami-gun, Nikko-mura; Kuromoriyama vi- cinity, Nishiokuni-mura; Maemoriyama vicinity, Higashiokuni-mura; Kamitakarazawa, Higashisa- wa-mura. Miyagi Prefecture — Ryozen, Hippo- mura; Oouchi-mura; Oofukasawayama, Shichiga- shuku-mura; Kuraishidake, Miya-mura; Kawane- dake National Forest vicinity; Daitoudake vicini- ty, Akiu-mura; Takakurayama, Ohsawa-mura; Ooshibayama vicinity, Onikoube-mura; Funako- shi, Jugohama-mura; Makinosaki vicinity, Ooh- ara-mura. Akita Prefecture — Ookawamae Nation- al Forest, Funaoka-mura; Iwamiyama National Forest vicinity, Iwamisannai-mura; Hatta, Taihei- mura; Shirayama National Forest, Kamishinjou- mura; Okuyamasawa National Forest, Kamikoani- mura; Iwase National Forest, Yamase-mura; Ma- sezawa, Hachimori-mura; Ozawayama, Sawame- mura; Kasuge-mura; Fujikotozawa National Forest vicinity. Fujikoto-mura; Towadako Lake vicinity. Iwate Prefecture — Goyosan, Hikoroichi- mura; Tsuchikurayama, Kamiarusu-mura; Utou, Otsukirai-mura; Shobuzaki, Yoshihama-mura; Kirigatana, Touni-mura; Komagatake, Kanega- saki-mura; Samusawayama, Oota-mura; Minami- toyosawayama vicinity, Yukuchi-mura; Wainai, Kariya-mura; Fuefuki National Forest, Aozasa- mura; Kataiwa National Forest, Kamigou-mura; Matsukurayama, Kasshi-mura; Katahayama, Ku- rihashi-mura; Tsunagi, Yamagata-mura. Aomori Prefecture — National Forest, Iwasaki-mura; Na- tional Forest, Fukaura-mura; National Forest, Oodose-mura; National Forest, Akaishi-mura; Ka- waratai vicinity, Nishimeya-mura; Tanosawa vi- cinity, Uchigata-mura; Yotsugadake vicinity, Aiu- chi-mura; Nakakodomariyama, Kodomari-mura; Sanyoushi, Minyama-mura; National Forest, Ima- betsu-mura; National Forest, Tairadate-mura; Ya- bitsuyama National Forest, Kanita-mura; Azuma- dake vicinity, Azumadake-mura; Towada, Houo- kuzawa-mura; Yaichirousawa vicinity, Oohata- mura; Hachimoriyama, Kazamaura-mura; 112 FIELDIANA: ZOOLOGY Futamatayama, Oo-oku-mura; Nuidouishiyama; Gendoushiro National Forest, Wakinosawa-mura. KINKAZAN (= KINK AS AN): Miyagi Prefec- ture— Kinkazan vicinity, Ayukawa-mura. Appendix 3. Locality Records (Specimens Examined and Literature Records) of Macaca fuscata, Excluding Localities Recorded in Questionnaire Surveys Conducted by K. Hasebe in 1923 (Fig. 2A, Appendix 2), K. Kishida in 1953, and H. Takeshita in 1964 (see above, Geographic Distribution and Total Population Estimate); for Documentation, see Gazetteer, Appendix 4. Key to Locality Numbers Northeastern section (NE) 1. Oma-machi 2. Sai-mura 3. Okoppe River 4. Bushidomari 5. Kusodomari; Shimokita Peninsula; Wakino- sawa-mura vicinity 6. Kanita-machi 7. Okuradake 8. Ajigasawa-machi 9. Tsugaru-touge 10. Nishimeya-mura 11. Hachimori-machi 12. Yamagata-mura 13. Akita-city 14. Kawabe-machi 15. Nangai-mura 16. Chokai-machi 17. Ogachi-machi 18. Genbikei 19. Sumita-cho 20. Goyosan 21. Ofunato-city 22. Kinkazan, Locs. 1, 2, 6, 10-15, 20, 24, 28, 33, Y6 23. Narusegawa 24. Tagajo-ato 25. Tarogawa 26. Futakuchi 27. Higashine-city 28. Yamadera 29. Kaminoyama-city 30. Kaisho 3 1 . Komogawa; Takane 32. 33. 34. 35. 36. 37. 38. 39. 40. 41. 42. 43. 44. 45. 46. 47. 48. 49. 50. 51. 52. 53. 54. 55. 56. 57. 58. 59. 60. 61. 62. 63. 64. 65. 66. 67. 68. 69. 70. 71. 72. 73. 74. 75. 76. 77. 78. 79. 80. 81. 82. 83. 84. Asahi-mura Kurokawa-mura Takahata-machi Shichigashuku-machi Azumayama Fukushima-city Soma-city Kashima-machi Shiobara-mura Kawajionsen Kuriyama-mura Imaichi-city; Nikko-city Ashio-machi Karuizawa-machi Sanada-machi Sugadaira Shiga-kogen (= Shiga Heights) Jigokudani; Karitayama; Maki; Masugata; Mi- yamura; Nakayama; Ogikubo; Samusawa; Shiono; Takai; Tenjinbara; Wakamiya-Hachi- manjinja; Warabitaira Wakahowatauchi Atosugiyama Asahi-machi Unazuki-machi Kurobe Kamiichi-machi Oosawano-machi Hosoiri-mura Hakusan Neo-mura Shichisou-cho Inuyama-city; Ohirayama Okinoshima Oshiro Shinshiro-city Hamakita-city Tatsuyama-mura; Tenryu-city Toei-cho Motomura; Suzawa Kamimachi; Nakadachi Aoki; Fukagasawa; Irisawai; Kashio; Metaka; Nakahora; Nishi; Ochiai; Ookawara; Shimizu; Shimoichiba Buna; Katsurashima; Yanagisawa Horane Kamiseinaiji; Kawasuso; Shimoseinaiji; Taki- nozawa Azumabashi Kakisore Kamidaira; Kuwasononanbu; Miyamoto; Mi- yashita; Oojimasan; Tazawa; Tsukiyodaira; Ushimaki; Yamabuki Kasugadaira; Kitamura; Shinyashiki Higashiina Nishiharuchika Higashino; Kiso Oobara; Shouzawa Hinata Outaki-mura; Seto, Outaki-mura Takigoshi FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 113 130° 135° 85. Furukaidou; Konishi 86. Fukasawa; Kurouchi; Misokawa 87. Kamikochi 88. Ohamidake; Yarigatake 89. Hida-sanmyaku 90. Hinatayama; Omachi-city; Oosaki; Tokiwa 91. Omachi, hills near 92. Inokamireien; Kawanishi; Miyashiro: Nezumiana 93. Hodaka; Hodaka Golf Course; Tatai 94. Shimashima; Yakeyama 95. Minamimachi: Nakasu 96. Mukawa-mura 97. Kawaguchi-ko 98. Fujiyoshida-city: Nishikatsura-cho 99. Fuji-san 100. Nishiizu-cho 101. Hagachizaki; Ihama 102. Atami-city 114 FIELDIANA: ZOOLOGY a Northeastern section (NE) Specimens examined Literature records Introduced populations Extinct populations Subfossils 56^55 57 1 92 *393 40° 34 35 43/ 84.83828180 ^>7 75. 77J , <• f 71 m VJ°9o8o107 • ~ 99 1045 103%/ ,67 rf^ Q 140° 35° FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 115 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. Hakone-machi; Yugawara-machi Odawara-city Ninomiya-machi Isehara-city Atsugi-city Aikawa-machi; Kiyokawa-mura; Tsukui-machi Tanzawayama Fujino-machi; Takaosan Hachioji-city Hinohara-mura; Itsukaichi Ome-city Tabayama-mura Okutama-machi Arakawa-mura Ashigakubo; Yokoze-machi Houtosan Kimitsu-city; Otsubo Aikawa, Futtsu-city; Boso Peninsula (ca. 35°16'N); Futairi; Futtsu-city; Hagiu; Hinaba- take; Hirooka; Hongou; Kiwadahata; Kyonan- machi; Nishihikasa; Nutazawa; Okugome; Okuyama, Kyonan-machi; Ooiwa; Shigumi; Shimosakuma; Shippara; Tagura; Takagoyama A; Takagoyama S; Takagoyama Tl; Yamanaka Yonezawa Mera Shirahama-machi Boso Peninsula, (ca. 35°00'N). Fukurogura; Kitakomachi; Kiyosumiyama; Okutani; Sunada; Utsutsumi Kamou; Kokurano; Kozawamata; Kuradama; Ootashiro; Tashiro; Tsukagoshi; Tsutsumori; Yokose Ishizuka; Koshikiya; Tokuji; Tsukizaki Kayano-Nanamagari; Sanda Mobara-city Southwestern section (SW) 1. Mikata-cho 2. Mihama-cho, Fukui 3. Makino-cho 4. Nishiazai-cho 5. Yogo-cho 6. Kohoku-cho; Takatsuki-cho 7. Ibuki-cho 8. Ryozenyama 9. Taga-cho 10. Hatasho-cho 11. Eigenji-cho 12. Hokusei-cho 13. Inabe-cho 14. Yunoyama, Komono-cho 15. Tsuchiyama-cho [1]; Tsuchiyama-cho [2] 16. Hino-cho 17. Koka-cho 18. Kameyama-city 19. Oyamada-mura 20. Iga-cho 21. Muroji 22. Oomiya-cho 23. Gokashowan; Nansei-cho 24. Owa.se-city 25. Kumano-city 26. Mihama-cho, Mie 27. Hongu-cho 28. Kozagawa-cho 29. Tsubaki Wild Monkey Park 30. Shirahama-cho 31. Kanaya-cho 32. Tomogashima 33. Minoo-city 34. Arashiyama 35. Otsu-city 36. Hieizan 37. Kamigamo 38. Hiyoshi-cho 39. Mizuho-cho; Sasayama-cho 40. Kasuga-cho 41. Wachi-cho 42. Doumoto; Kidani; Naka; Natasho-mura 43. Adogawa-cho 44. Kaminaka-cho 45. Hotokedani 46. Takahama 47. Otoumi 48. Ine-cho 49. Takeno-cho 50. Wakasa-cho 51. Funakoshiyama 52. Kumayama-cho 53. Kanbanotaki 54. Katsuyama-cho 55. Gagyusan 56. Tenjinkyo 57. Enmeikyo 58. Taishakukyo 59. Akashi; Fukano; Higashihinobori; Houin; Kisuki-cho; Koujiro; Mitoya-cho; Nishihi- nobori; Okuyama, Mitoya-cho; Takuwa; Tsukinoya; Yoshida-mura 60. Ooda-city 61. Mikusu 62. Hasumi-mura; Hinagawa; Hirasa; Imanishi; Shimogou; Ueda; Ushiroyama 63. Kochi-cho 64. Mihara-city 65. Miyajima-cho 66. Akiyoshi Cave 67. Omishima 68. Choshikei; Rosando; Shodoshima [1]; Sho- doshima [2] 116 FIELDIANA: ZOOLOGY 69. Awajishima; Kaminada 70. Jinrio (= Jinryo) 71. Anan-city 72. Tachibana 73. Kushigawa 74. Aki-city 75. Chikaraishi 76. Hiromi-cho 77. Nametoko 78. Matsuno-cho 79. Nakamura-city 80. Odo; Odomisaki; Okuuchi-mura 81. Kashima, Ehime 82. Kamae-machi 83. Takasakiyama 84. Kawaradake 85. Gongenyama 86. Hiradoshima 87. Kyogatake; Tara-dake 88. Kugino-mura 89. Higo 90. Itsuki-mura 91. Sagara-mura 92. Kanayama 93. Satsuma-cho 94. Ichiki-cho 95. Kanoya-city 96. Kushima-city 97. Kojima; Torishima 98. Toi-misaki 99. Tanegashima 100. Issou; Kamiyaku-cho [1]; Kusugawa; Mi- yanoura; Shirakoyama; Yakushima, Loc. P; Yakushima, Loc. Q 101. Hanyama; Kamiyaku-cho [2]; Kannonzaki; Kawara; Kojiba; Kuniwaridake, Kamiyaku- cho; Kuniwaridake, southeastern slopes; Kuniwaridake, summit area; Nagata; Naga- ta-todai; Nina-A; Seibu-rindou; Shikamiza- wa; Yaku vicinity; Yakushima, Loc. A-Loc. O; Yakushima, southwestern; Yakushima, western, road vicinity, high zone; Yakushi- ma, western, road vicinity, low zone; Yaku- shima, western, road vicinity, middle zone 102. Hinokuchi, Yaku-cho; Koshima; Mugio; Nakama; Nakase; Nakasegawa; Shichigo- dake; Yakushima, Loc. R-Loc. U; Yudomari FOODEN AND AIML SYSTEMATIC REVIEW OF JAPANESE MACAQUES 117 Appendix 4: Gazetteer of Macaca fuscata Collection and Observation Localities The sequence of information presented in this gazetteer is as follows: 1. LocaHty name, in roman letters and Kanji/Kana characters. 2. Name of prefecture (italics) and island (capi- tal letters). 3. Coordinates of locality. 4. Date of collection or observation. 5. Name of collector or observer. 6. Bibliographic reference (in parentheses) to published or unpublished field notes, if any, 7. Abbreviated name of museum (see Introduc- tion) where specimens are preserved. 8. Number of specimens available (with indi- cation of part preserved, if skin and skull are not both present). 9. Locality number (italicized) as indicated in distribution maps (Appendix 3); this does not apply to Hasebe's 1923 survey records, which are mapped in Figure 2A. Abeyama, Naka-son, Kawakami-gun (JI|_t?P4' WPrI^Oj); Okayama, HONSHU; 34°48'N, 133°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV-2). See Fig. 2A. ^ Adogawa-cho, Takashima-gun (iliS?P^^JI| mi); Shiga, HONSHU; ca. 35°19'N, 136°01'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW43. Aikawa, Futtsu-city ( S /$ T^ ffi Jll ); Chiba, HONSHU; 35°11'N, 139°53'E; collected 18 Dec. 1998-30 Mar. 1999 by PRIKU staff; PRIKU, 5 (skeletons only). NE120. Aikawa-machi, Aikou-gun ( S ¥ ?P ^ Jl| fflJ ); Kanagawa, HONSHU; ca. 35°29'N, I39°17'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33). NEWS. Aikawautsubo, Namiai-mura, Shimoina-gun (T ^Mmi^^¥i^ )\\0'y -^^y, Nagano, HONSHU; 35°22'N, 137°41'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXI-53). See Fig. 2A. Aikawayama, Misho-cho, Minamiuwa-gun (1^^ ^?P=!®SfflI*IJIiai); Ehime, SHIKOKU; 32°59'N, 132°34'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18: IX-2). See Fig. 2A. Aimotosanchi vicinity, Aimoto-mura, Shimonii- kawa-gun (Tiff JH IPM^^^^ Ol Hfe^); Toyama, HONSHU; 36°52'N, 137°35'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 42: XXIX-10). SeeFig. 2A. Aitakayama, Takane-mura, Sunto-gun (^^^ fll^^SliUJ); Shizuoka, HONSHU; 35°09'N, 138°49'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p.45:XXX-14). SeeFig.2A. Ajigasawa-machi, Nishitsugaru-gun (H/^fe?P 0<7-)Rarr); Aomori, HONSHU; ca. 40°47'N, 140°12'E; mtDNA sample collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NE8. Akagawaomote, Yunotani-mura, Kitauonuma- gun (db^)S?P^;^^^#JI|^); Niigata, HONSHU; 37°11'N, 139°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII-9). See Fig. 2A. Akagiyama, Kurohone-mura, Seta-gun {^^W> H ft ^ ^ # ^ UJ ); Gunma, HONSHU; 36°3rN, 139°16'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 52: XL- 17). See Fig. 2A. Akagiyama vicinity, Akagine-mura, Tone-gun ( ^IJ a IP # ^ a ^ '* ^ OJ ffe ); Gunma, HONSHU; 36°35'N, 139°13'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 52: XL- 16). See Fig. 2A. Akago, Kitayoshii-mura, Onsen-gun (/m^lPdb B#W'^^^); Ehime, SHIKOKU; 33°50'N, 132°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21:IX-24). SeeFig. 2A. Akaioku, Nunuki-mura, Hikami-gun (7KJllP>iS a:^##^); Hyogo, HONSHU; 35°08'N, 135°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-25). See Fig. 2A. Akaishisan vicinity, Ooshika-mura, Shi- moina-gun (T#IP^A^^#^ajfte); Na- gano, HONSHU; 35°33'N, 138°06'E; reported FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 119 in questionnaire suney conducted in 1923 by K. Hasebe (Iwano. 1974, p. 47: XXXI-46). See Fig. 2A. Akaiwa Mountain Range, Maruyae-mura, Kasa-gun (iPfelPAA>I^#^Ujfl^): Kyoto, HONSHU; coordinates unknown; reported in questionnaire sur\ey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXII-9). Not mapped. Akaiwayama, Okadanaka-mura, Kasa-gun (iP-te iP fS^^ffl 4^ ^ # ^ UJ ); Kyoto, HONSHU; 35°27'N, 135°15'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXII-8). See Fig. 2 A. Akakuzureyama. Shimonoho-mura, Mu2i-2un HONSHU; 35°33'N, 136°59'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIII- 19). See Fig. 2A. Akamatsu, Akagaw achi-son, Kaifu-gun (^nP?P # )! 1*1 ^ # V^ ); Tokushima. ^SHIKOKU; 33°44'N. 134°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-8). See Fig. 2 A. Akanishi vicinity, Okutani-mura. Shisou-gun (5^ ^lPa^^#ffi^); Hyogo, HONSHU; 35°13'N, 134°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-8). See Fig. 2 A. Akashi, Mitoya-cho, lishi-gun (IS51PH TJMfflJ a^ S ); Shimane. HONSHU; 35°14'N, 132°50'E; collected 25 Jul 1995 by PRIKU staff; PRIKU, 1 (skeleton only). SW59. Akasugiha vicinity, Yamakami-mura, Kan- zaki-gun (#if AOJ ±^#^!^^itfe); Shiga, HONSHU; 35°04'N, 136°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV-24). See Fig. 2A. Akataniyama, Akatani-mura, Kitakanbara-gun (db«'jiiP#^*\r##aj); Migata, HONSHU; 37°50'N, 139°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII-20). See Fig. 2A. Akataniyama, Ootou-mura, Yoshino-gun ("a© lPA^^#^aj); Nara. HONSHU; 34°09'N, 135°46'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35:XXI-2). SeeFig. 2A. Aki-cit>' (^STtT); KochL SHIKOKU; 33°30'N, 133°54'E; reported present before 1830 by P. F. von Siebold (MS., 1830-1842, p. I; cf. Tem- minck. 1842. p. 10; Kuroda, 1938, p. 112; 1940, p. 270). SW74. Akita-city {^X.^1^): Akita. HONSHU; 39°43'N, 140°07'E; mtDNA extracted from hunting tro- phy before 2003 by T. Agatsuma and M. Ishi- gami (2002, p. 79). NEIS^ Akiyoshi Cave. Shuhou-cho, Mine-gun (II^\1P 4li^ ^ ^K^ M ); YamagiichL HONSHU; 34°14'N, 131°18'E; subfossils, possibly older than Early Jomon (possibly >12 Ka), reported by M. Kuroda (2002a, p. 115); Kyoto Univer- sity Anthropology Laboratory. 2 (skulls only, not seen). SW66. Akoumayama. Toyonaga-son. Atetsu-gun (P5f|^ IP M ^ ^ # .1 UJ V rt ^ 111 ); Okayama, HONSHU; 34°58'N. 133°35'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 27: XV-12). See Fig. 2A. Akutayama. Tsugayuki-mura, Oochi-gun (e^ ?P tP M ^ ^^^ \h ); Shimane. HONSHU; 35°00'N, 132°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 28: XVI-24). See Fig. 2A. Amagisan, Inatori-cho, Kamo-gun (Mj^lPHUi IHI^i^aj); Shizuoka, HONSHU; 34°46'N, 139°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974, p. 45: XXX- 19). SeeFig. 2 A. Amagisan. Kamikavvatsu-mura, Kamo-gun (l£^ iP±)p[^^;f^t^^UJ); Shizuoka, HONSHU; 34°49'N, 138°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX-20). See Fig. 2 A. Amagisan. Kitou-mura. Kamo-gun (MS?P^^ :H"^^^aj); Shizuoka. HONSHU; 34°50'N, 139°04'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p.45:XXX-18). SeeFig.2A. Amagisan, Nishina-mura, Kamo-gun (SM^Pil- ^i^t^^LiJ); Shizuoka, HONSHU; 34°47'N, 138°46'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p.45:XXX-21). SeeFig. 2A. Amagoitaki waterfall vicinity. Shinritsu-mura, 120 FIELDIANA: ZOOLOGY Uma-gun (^JSl5|/f II;f^)■M'£)tftfe); Ehime, SHIKOKU; 33°56'N, 133°40'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18: IX- 12). See Fig. 2A. _ Ambo-Onoaida( ^M—M'i^f^). See Yakushima, Census Area 4. Amegadake, Furuseki-mura, Nishiyatsushiro-gun (ffi A ft ^ * H ^ M X!tMW^M-[hi^y, Niigata, HONSHU; 37°22'N, 139°05'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII-10). SeeFig. 2A. Asan Mountain Range, Tawa-mura, Ookawa-gun ( A Jll IP ^ ^ ^ PrI M LiJ M ); Kagawa, SHIKOKU; 34°10'N, 134°11'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 24: XII-4). See Fig. 2A. Asaoki, Mifune-mura, Minamimuro-gun (^#ft ^^^^^'^Wy, Mie, HONSHU; 33°44'N, I35°58'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIII-57). See Fig. 2A. Ashigakubo, Yokoze-machi, Chichibu-gun (^;^ ^^M^P^X^y, Saitama, HONSHU; ca. 35°58'N, 139°08'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE117. Ashigara Mountain Range, Miyagino-mura, Ashigarashimo-gun (£ffi"F?P^^i?^S:M 111 M ); Kanagawa, HONSHU; 35°16'N, 139°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXV-3). See Fig. 2A. Ashimi-mura, Oono-gun (A©?P^^^); Fukui, HONSHU; 36°03'N, 136°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV- 19). See Fig. 2A. Ashio-machi, Kamitsuga-gun (Ji^MlPSMffll); Tochigi, HONSHU; 36°38'N, 139°27'E; col- lected 20 Aug. 1986 by TPM staff; TPM, 1 (not seen, data from specimen list). NE44. Ashiu vicinity, Chii-mura, Kitakuwada-gun (db ^ffllPife^^^r^^ffe); Kyoto, HONSHU; 35°18'N, 135°40'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXII- 17). See Fig. 2A. Ashiyama vicinity, Ikuma-mura, Nishimuro-gun (H^«?P^.l^Fajffe); Wakayama, HONSHU; 33°41'N, 135°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-24). See Fig. 2A. Atagoyama vicinity, Sachiyo-mura, Hikami-gun (7K±?P^tM:WS^Ujfte); Hyogo, HONSHU; 35°irN, 135°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-24). See Fig. 2A. Atami-city (l^^rfT); Shizuoka, HONSHU; ca. 35°05'N, 139°04'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NE102. Atosugiyama, Otani-mura, Kitaazumi-gun (db$ *iP'h^W^^^Uj); Nagano, HONSHU; 36°52'N, 137°54'E; collected 14 Jan. 1998 by M. Aimi; PRIKU, 1 (skeleton only). NE51. Atsugi-city (jpT^ r{]); Kanagawa, HONSHU; ca. 35°27'N, 139°22'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33). NE107. Awagatake, Morimachi-mura, Minamikan- bara-gun {^MW-^W>^^ ¥iMT Uy Niigata, HONSHU; 37°28'N, 139°09'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXIl-14). See Fig. 2A. Awajishima C^^^y Hyogo, AWAJISHIMA; ca. 34° 14' N, 134°52'E; provisioning initiated in 1967 (Shidei et al., 1981, p. 18). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). Microsatellite DNA analyzed before 1996 by Domingo-Roura et al. (1997, p. 358). External measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW69. Awano, Kawamata-mura, linan-gun (tSl^fPJII ^ ^ ^ S ); Mie, HONSHU; 34°26'N, 136°18'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-23). See Fig. 2A. Ayuhanasanrin, Nishida-mura, Mikata-gun (H^ ^'mm^TJ.)\-i-\h^y Fukui, HONSHU; 35°32'N, 135°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV- 12). See Fig. 2A. Azumabashi, Nagiso-machi, Kiso-gun (^KelP ^TK^fflJl^SM); Nagano, HONSHU; 35°35'N, I37°35'E; collected 28 Mar. 1998 by M. Aimi; PRIKU, 1 (skeleton only). NE74. Ill FIELD! AN A: ZOOLOGY Azumadake vicinity, Azumadake-mura, Higa- shitsugaru-gun (K/$^15:^^:H"K^ftfe); Aomori, HONSHU); 40°50'N, 140°54'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-17). SeeFig. 2A. Azumasan, Azuma-mura, Yama-gun (?P^1Pr- SWl^gllj); Fukushima, HONSHU; 37°37'N, 140°irE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 52: XLI-8). See Fig. 2A. Azumayama (^SUJ); Yamagata, HONSHU; 37°44'N, 140°12'E; capture of albino monkey 13 Mar. 1938 reported by N. Kuroda (1940, p. 21\).NE26. Azumayama, Minamihara-mura, Minamioki- tama-gun (j^Sl^lPl^J^^^SUj); Yama- gata, HONSHU; 37°48'N, 140°08'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLII-2). See Fig. 2A. Azumi National Forest, Azumi-mura, Minamia- zumi-gun (^^ftl^^ft^^^ftBW^); Nagano, HONSHU; 36°13'N, 137°42'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-12). SeeFig. 2A. Beaverton; Oregon, U.S.A.; 45°29'N, 122°48'W; captive provisioned group translocated from Mihara ca. 1965 (Van Horn, 1980, p. 198; Rostal et al., 1986, p. 453). Not mapped. Benigadake vicinity, Soni-mura, Uda-gun (^PS W>^M^%LT ^^); Nam, HONSHU; 34°3rN, 136°08'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXI-9). See Fig. 2 A. Bettou, Misato-mura, Minamikoma-gun (I^^JS IP H M ^^t Sy ^ ); Yamanashi, HONSHU; 35°30'N, 138°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-8). See Fig. 2A. Bonhoriyama, Togura-mura, Nishitama-gun (S ^SiPP^^^^iSUj); Tokyo Metropolis, HONSHU; coordinates unknown; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXIV-4). Not mapped. Bono-misaki, Nishiminamikata-mura, Ka- wanabe-gun ( Jll ffl^ffi l^7^^±& y ffllllt^); Kagoshima, KYUSHU; 31°20'N, 130°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-4). See Fig. 2A. Boso Peninsula (M-lvt^ft); Chiba, HONSHU; ca. 35°00'N, 139°55'E; observed Jan.- Dec. 1997 by K. Aizawa and M. Hagiwara (2001, p. 6). NE124. Boso Peninsula (M,1vg¥ft); Chiha; HONSHU; ca. 35°16'N, 139°59'E; field studies conducted in 1972-1973 reported by K. Fukuda (1975, p. 386) and T. Iwano (1977, p. 46; cf. Study Group on the Present Status of Japanese Mon- keys, 1977b, p. 26). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE120. Bukkazan, Susugaya-mura, Aikou-gun (^^IP ^ ^ # ^ ^A ^ Oj ); Kanagawa, HONSHU; 35°3rN, 139°15'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 50: XXXV-6). See Fig. 2A. Bukouzan, Kagemori-mura, Chichibu-gun (^;5<^ ?P ^ ^ ^^ E ¥ liJ ); Saitama, HONSHU; 35°58'N, 139°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 50: XXXVII-7). See Fig. 2A. Bukouzan, Yokoze-mura, Chichibu-gun (l>^^?P #)i^^E¥UJ); Saitama, HONSHU; 35°58'N, 139°08'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 50: XXXVII-8). See Fig. 2A. Buna, Matsukawa-machi, Shimoina-gun (T'^IP IP^iJIIWJE^); Nagano, HONSHU; 35°36'N, 137°57'E; collected 23-Jan.-25 Feb. 1998 by M. Aimi; PRIKU, 5 (3 skeletons only, 2 cranial fragments and postcranials only). NE71. Bunajurin, Minakami-mura, Tone-gun (^'J^^ yY.h^\l\^^^^); Gunma, HONSHU; 36°49'N, 139°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XL- 12). See Fig. 2 A. Bushidomari, Wakinosawa-mura, Shimokita-gun iTitmJ&m)R^^±^^y, Aomori, HONSHU; 41°12'N, 140°47'E; collected in 1965-1966 by S. Azuma; JMC, 1 (skull only, found on sea- shore). NE4. Buttsujisan, Takasaka-son, Toyota-gun (MBB^ m^^il^M^ihy, Hiroshima, HONSHU; 34°28'N, 133°02'E; reported in questionnaire FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 123 survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-25). See Fig. 2 A. Byobuiwa, Kadohara, Nakahara-mura, Ma- suda-gun (MBa?P4^i^WP^i^JPm^); Gifu, HONSHU; 35°44'N, 137°13'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIII-35). See Fig. 2A. Cape Toi, Kushima. See Toi-misaki, Kushima. Chaen, None-mura, Aki-gun ($S?Pi?^^l^ S); Kochi, SHIKOKU; 33°30'N, 134°16'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-42). See Fig. 2A. Chichibu Mountain Range, Kawakami-mura, Minamisaku-gun (^fejXlPJI|±^^5<:UjM); Nagano, HONSHU; 35°56'N, 138°36'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-39). See Fig. 2A. Chichiyama, Toyooka-mura, Kamitakai-gun (_h i5#?PfiEW?Lai); Nagano, HONSHU; 36°36'N, 138°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-30). See Fig. 2A. Chikaraishi, Higashitsuno-mura, Takaoka-gun {m^^Mni^ti^); Kochi, SHIKOKU; 33°24'N, 133°03' E; collected 10 Aug. 2002 by M. Aimi; PRIKU, \.SW75. Chishorodani vicinity, Hanase-mura, Atago-gun (SS?P^W^»tti:S&#ffe); Kyoto, HONSHU; 35°09'N, 135°48'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXII-22). See Fig. 2A. Choenzan, Honjou-mura, Yosa-gun (%^lP^j£ ^ :U^ OJ ); Kyoto, HONSHU; 35°44'N, 135°15'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35:XXII-3). SeeFig. 2A. Chokai-machi, Yuri-gun (S^JIPB^^W); Akita, HONSHU; 39°11'N, 140°I1'E; mtDNA ex- tracted from hunting trophy before 2003 by T. Agatsuma and M. Ishigami (2002, p. 79). NEI6. Choshigatani, Makino, Nishinosho-mura, Taka- shima-gun (S S^PBH+^ife^^^ ^ ^); Shiga, HONSHU; 35°28'N, 136°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV- 10). See Fig. 2A. Choshikei, Tonosho-cho, Shozu-gun {^\^3.^:t. Ji fflj ^k ^ M ); Kagawa, SHODOSHIMA; 34°3rN, 134°15'E; provisioned group; birth season, 1958-1966, reported by Kawai et al. (1967, pp. 37, 38). 5^65. Daieiyama, Nakahagi-mura, Nii-gun (^M^P't' i^^A^Uj); Ehime, SHIKOKU; 33°55'N, 133°2rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21:1X-15). SeeFig. 2A. Daigentasan, Niiharu-mura, Tone-gun (^J1S?P^ >p^A3f^;*;aJ); Gunma, HONSHU; 36°47'N, 138°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51:XL-I1). SeeFig. 2A. Daikokuyama, Miyakogawa-mura, Mi- namikoma-gun (^ ^S^PUP Jll ^ A M UJ ); Yamanashi, HONSHU; 35°28'N, 138°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-7). See Fig. 2A. Daikokuyama vicinity, Mikuni-mura, Mi- namiuonuma-gun (^^>SiPHH:H"AMUjflfe); Niigata, HONSHU; coordinates unknown; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXlI-8). Not mapped. Daitoudake vicinity, Akiu-mura, Natori-gun (=S IJ^iPfJ^^WA^^fte); Miyagi, HONSHU; 38°17'N, 140°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLIII-6). See Fig. 2A. Daitouzan vicinity, Nishiyama-mura, Mi- namikoma-gun (^i^S^PHlllWA^Lil'te); Yamanashi, HONSHU; 35°33'N, 138°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-9). See Fig. 2A. Dakeyama, Kiyodake-son, Kounu-gun (¥^?P m^^^\h); Hiroshima, HONSHU; 34°44'N, 133°10'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-32). See Fig. 2A. Damasanchi, Ichinose-mura, Nishimuro-gun (M #ftfPTfTy^;^^^ajitk); Wakayama, HONSHU; 33°42'N, 135°28'E; reported in questionnaire survey conducted in 1923 by K. 124 FIELDIANA: ZOOLOGY Hasebe (Iwano, 1974, p. 33: XX-23). See Fig. 2A. Doumoto, Natashou-mura, Onyu-gun (^^IP^ fflJ±4^M;z|s^); FukuU HONSHU; ca. 35°23'N, 135°40'E; collected Aug.-Nov. 1994 by PRIKU staff; PRIKU, 2 (skeletons only). SW42. Doushi-mura, Minamitsuru-gun (i^tPlilPJil^ ^ ); Yamanashi, HONSHU; 35°32'N, 139°02'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXIIl-31). See Fig. 2A. Doutokoyama, Suzuhari-son, Asa-gun (^fe^P ip ?I ^ ^ ^ UJ ); Hiroshima, HONSHU; 34°37'N, 132°34'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV- 12). See Fig. 2A. Douyama, Hata, Tajibe-son, Atetsu-gun (P5[^?P :l^ >p pP ^ ^ M OJ ); Okayama, HONSHU; 34°59'N, 133°33'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV-9). See Fig. 2A. Ebi-cho, Minamishitara-gun (l^lx^lP/S^fflJ); Aichi, HONSHU; 35°01'N, 137°35'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVII-3). See Fig. 2A. Ebitadani vicinity, Ootsuka-mura, Nishi- muro-gun (S^«?P AtP)Rl^I ti'^^fe); Wakayama, HONSHU; 33°34'N, 135°35'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-32). See Fig. 2A. Egasayama, Shimo-mura, Izushi-gun (t±JS?PS §^^t>lSUJ); Hyogo, HONSHU; 35°30'N, 135°02'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-22). See Fig. 2A. Egawa vicinity, Kamiakizuki-mura, Asakura-gun ( ^ 5t ^ ± 1^ >^ ^ >I Jll ffe ); Fukiwka, KYUSHU; 33°41'N, 130°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 17: Vll-3). See Fig. 2A. Eigenji-cho, Kanzaki-gun ( tt llif IP ^ /Jl ^ BJ ); Shiga, HONSHU; ca. 35°05'N, 136° 18' E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SWII. Enasan, Chisato-mura, Shimoina-gun (~F'^3P^ ^S^SSPOJ); Nagano, HONSHU; 35°26'N, 137°42'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXI-52). See Fig. 2A. Endani vicinity, Kawazoe-mura, Nishimuro-gun (®#S?PJI|^WR^ffe); Wakayama, HONSHU; 33°38'N, 135°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-28). See Fig. 2A. Enmeikyo, Yamano-cho, Fukuyama-city (^ ill TtT Ol i? ffll M II l^^ ); Hiroshima, HONSHU; 34°42', 133°22'E; mtDNA samples collected 1998-2000 by I. Yoshimi and H. Takaski (2003, p. 1\).SW57. Figo. See Higo. Fudoutaki, Nishikomagadake, Akaho-mura, Kamiina-gun (±^3P?P#B^M|6lS); Nagano, HONSHU; 35°45'N, 137°58'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXl-42). See Fig. 2A. Fuefuki National Forest, Aozasa-mura, Kami- hei-gun (Jim^mm^^^^m^^y, Iwate, HONSHU; 39°20'N, 141°40'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Mito, 1989, p. 25). See Fig. 2A. Fuji-san (Silll) (= Fuji-yama); Kanagawa, HONSHU; 35°22'N, 138°44'E; collected be- fore 1888 by M. Seller; MNHN, 3 (including 2 with skulls in skins). NE99. Fujigaishi vicinity, Kamo-mura, Nii-gun (Iff M^ iP j^ W ^ 1^Maj^): Mie. HONSHU: 34°23'N. 136°29'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 37: XXIIl-32). See Fig. 2A. Fujikotozawa National Forest vicinity. Fuji- koto-mura. Yamamoto-gun (lll^fP^^^M #)RSW^fte): .-lA7/t7r HONSHU: 40=20TsI, 140°47'E: reported in questionnaire survey conducted in 1 923 by K.. Hasebe ( Iwano. 1 974. p. 56: XLI V- 12). See Fig. 2 A. Fujimiyama vicinitv. Kuriyama-mura, Shio'ya-gun (i^^lPliaj^S±Maj^): 7b- chigi. HONSHU: 36=52'N. 139°32'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 51: XXXIX-4). See Fig. 2A. Fujino-machi. Tsukui-gun (^^^IPMSW"): kanagawa. HONSHU": ca. 35°38'N, 139°irE: mtDNA samples collected before 1998 by Y. Kawamoto (1997. p. 33). SEllO. Fujisawa vicinitv. Shimokaifu-mura, Iwa- fline-gun (^ISiPT^it^S)R^): Migata. HONSHU: 38°25'N. 139°3rE: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 48: XXXII-25). See Fig. 2A. Fujiyoshida-cit>- ( S ± n^ B9 rfT ); Yamanashi. HONSHU": ca. 35=29'N. 138°48*E: mtDNA samples collected before 2003 by Y. Kawa- moto (2002. p. 60). \E98. Fukabayama. Suigen-mura. Kikuchi-gun (^^ ?P7K)g^)^^Uj): Kumamoro. k\Tj"SHU: 33°00'N. 130°54'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 17: IV- 1 1). See Fig. 2 A. Fukagasawa. Ooshika-mura. Shimoina-2un (T ^m^Am^^^'rn): Xagano. HONSHU: 35°30'N. 138°01'E: collected 14 Sep. 1998 by M. Aimi: PRIKU. 1 (skeleton only). \E70. Fukano. Yoshida-mura, lishi-gun (t5^1P"affl ^ ^^ S ): Shimane, HONSHU: 35=1 l-N. 132°55'E: collected 13-15 Dec. 1994 by PRIKU staff: PRIKU. 2 (skeletons only). SW59. Fukasawa, Kiso-mura. Kiso-gun (AelP^fi^ }S)R): Sagano. HONSHU: 35°58' N, 137°46'E; collected 30 Dec. 1997 and 28 Jan. 1998 by M. .\imi: PRIKU. 3 (skeletons only). NE86. Fukashimizu vicinity. Kawakami-mura. Taka- shima-gun (i^SIPJI|±^;^^7Kffe): Shiga, HONSHU: 35°25'N. 136°01'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 38: XXIV-8). See Fig. 2A. Fukatani vicinity. Yamanokuchi-mura, Oono-gun (AgiPaj;^P^}^^ffe): Gifu, HONSHU; 35°59'N. 137°10'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 41: XXVIII-37). See Fig. 2 A. Fukazawa National Forest. Nikko-machi, Kamitsuga-gun (±l?MiPB3feB0"}^>RSW ^): Tochigi. HONSHU: 36°44'N. 139°35'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XXXIX-3). SeeFig. 2A. Fukiyama National Forest. Minoo-son, Kov-u-gun (JPu^^iPZlrt^q^LijSW^); Mi- xazakL KYUSHU: 32°07'N. 131°19'E: re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 15: 11-17). See Fig. 2A. Fukubegadake. Takeda-mura, Gujo-gun (^JilP * ^^ MS): G/f//, HONSHU; 35°39'N, 136°56'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974, p. 41: XXVIII-22). See Fig. 2A. Fukuei-mura. Nishitagavva-gun (IIBQ JII?P^3^ ^): Yamagata. HONSHU: 38°34'N. 139°44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974, p. 55: XLII-5). See Fig. 2A. Fukiihara-son. Katsuura-gun (0/i^?PISi^^); Tokushima. SHIKOKU: 33°53'N. 134°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 23: XI-15). SeeFig. 2A. Fukurogura, Kamogawa-citv- ( fli jll rfT ^ j& ); Chiba, HONSHU: ca. 35°08'N. 140°07'E; col- lected Sep. 1997 by PRIKU staff: PRIKU, I (skeleton only). \EI25. Fukushima-cit> (IS firfJ): Fukushima. HONSHU; ca. 37°45'N. 140°28'E: blood samples col- lected before 1992 by Nozawa et al. (1991, p. 126 HELD! AN A: ZOOLOGY 414; 1996, p. 6). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE37. Funakoshi, Jugohama-mura, Monou-gun (^fe^ ?P + £ ^ ^^ fa M ); MiyagU HONSHU; 38°30'N, 141°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLIII-10). See Fig. 2A. Funakoshiyama, Mikawa-mura, Shisou-gun (7^ H ?P H M ;f^t fq^ M liJ ); Hyogo, HONSHU; 35°07'N, 134°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-5). See Fig. 2A. Funakoshiyama, Nankou-cho, Sayo-gun ("feffl^ ]^)tefflTfp'®aj); Hyogo. HONSHU; 35°07'N, 134°25'E; provisioned group; birth season, 1965-1966, reported by Kawai et al. (1967, pp. 37, 38). SW51. Furukaidou, Kaida-mura, Kiso-gun (T^e^^ffl ^*^^jI); Nagano, HONSHU; 35°58'N, 137°35'E; collected 4 Sep. 1998 by M. Aimi; PRIKU, 2 (skeletons only). NE85. Furutani National Forest vicinity, Warabihara, Kamiosakabe-son, Atetsu-gun (PR[@^±ffJoP ^MJ^*#lW^ffe); Okayama, HONSHU; 35°08'N, 133°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV-8). See Fig. 2A. Furutayama vicinity, Kitatane-mura, Kumage-gun (fl^?Pdb«^^* BQ OJ^); Kagoshima, TANEGASHIMA; 30°44'N, 131°02'E; reported as extinct in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-2). See Fig. 2A. Furuyajo, Toyooka-mura, Minamikoma-gun (1^ ^SiPMI^W^^^); Yamanashi, HONSHU; 35°20'N, 138°24'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIIl-3). See Fig. 2A. Futagoyama, Nishimusashi-mura, Higashikuni- saki-gun (:^S^?PffiEMWi35^Li|); Oita, KYUSHU; 33°34'N, 13r36'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: 111-20). See Fig. 2A. Futagoyama vicinity, Susami-mura, Nishi- muro-gun (lS#«lPMl#m^)5(^ UJ ffe); Wakayama, HONSHU; 33°33'N, 135°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-30). See Fig. 2A. Futairi, Kimitsu-city ( ^ /$ rtT ~ A ); Chiba, HONSHU; 35°13'N, 140°00'E; collected 23 Aug. 1998 by PRIKU staff; PRIKU, 1 (cranial fragments only). NE120. Futakuchi, Sendai-city ( fllj n T^ — P ), Miyagi, HONSHU; 38°16'N, 140°33'E; observed Jan.-Mar. 2003 by K. Izawa, T. Uno, and H. Fujita (2003, p. \1).NE26. Futamatayama, Oo-oku-mura, Shimokita-gun (T dblPAH^-XUj); Aomori, HONSHU; 41°26'N, 140°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-21). See Fig. 2A. Futamatayama, Yumoto-mura, Iwase-gun {^Wi. iPM^^-MliJ); Fukmhima, HONSHU; 37°16'N, 140°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 52: XLI-4). See Fig. 2A. Futtsu-city (s)iTU); Chiba, HONSHU; ca. 35=127^^, 139°53'E; mtDNA samples collected before 1999 by Y. Kawamoto (1998, p. 54). NE120. Gagyusan ( El 4 OJ ); Okayama, HONSHU; 34°49'N, 133°37'E; provisioning initiated in 1955 (Shidei et al., 1981, p. 18). Birth season, 1956-1966, reported by Kawai et al. (1967, p. 39). Liver samples collected from translocated captives before 1987 by Hayasaka et al. (1986, p. 346). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW55. Gagyusan, Matsuyama-son, Joubou-gun (_hj^?P ^i^ \h ^ ^ ^ \h ); Okayama, HONSHU; 34°49'N, 133°37'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV- 17). See Fig. 2A. Gagyusan, Takahashi-cho, Joubou-gun (JiMlP i^ ^ fflj ei 4^ Uj ); Okayama, HONSHU; 34°49'N, 133°37'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV- 16). See Fig. 2A. Gagyusan, Tsugawa-son, Joubou-gun (_hj^?Py$ JHWEil^Uj); Okayama, HONSHU; 34°49'N, 133°37'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 127 p. 27: XV- 15). See Fig. 2A. Gayatani vicinity, Nakatsu-mura, Kamiukena- gun (±>?7v:^4'^^:^'-\'#^); Ehime. SHIKOKU; 33°37'N, 133°04'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: IX-20). See Fig. 2A. Genbikei, Genbi-cho, Ichinoseki-city ( — ^TtJJ^ II WJ^ 11)1); hxate. HONSHU; 38°58'N, 141°02'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NE18. Gendoushiro National Forest, Wakinosawa-mura. Shimokita-gun (Tdb^^if)R;^)lK^lW ^); Aomoh, HONSHU; 41°25'N, 140°54'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-23). See Fig. 2A. Gezan, Toyotashimo-son, Toyoura-gun (sM?P S ra T W ^ OJ ); Yamagiichi, HONSHU; 34°10'N, 131°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII-2). See Fig. 2 A. Ginzan vicinity, Oomori-cho, Nima-gun (SRIP A^BTr^Ujftfe); Shimane, HONSHU; 35°08'N, 132°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-30). See Fig. 2A. Gokairiaiyama, Kamishou-mura, Oono-gun (A i^lP±Ji^E'r A^ULl); Fukui. HONSHU; 35°57'N, 136°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV-21). See Fig. 2A. Gokashowan {^TpfiMy, Mie. HONSHU; ca. 34°20'N, 136°41'E; observed Aug. 1979 by K. Masui (Takasaki & Masui, 1984, p. 310). SW23. Gokurakujisanrin, Hara-son, Saeki-gun (fe'ffilP ii^S^#lll^); Hiroshima', HONSHU; 34°22'N, 132°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV- 1). See Fig. 2A. Gongendaki vicinity. Sakashukitou-son, Naka-gun (i9m?PJS'>H^IIW=I>y >^ + ^ ); Tokushima, SHIKOKU; 33°50'N, 134°20'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 23:XI-13). SeeFig. 2A. Gongenyama, Higashisefuri-son, Kanzaki-gun (ftif^SW^^^mUj); Saga, KYUSHU; 33°22'N, 130°23'E; reported in 1973 by H. Ikeda and K. Eguchi (1978, p. 59). SW85. Gongenyama \icinity, Konose-mura, Kuma-gun (SSfPtt^^ft^iajfte); Kumamoto, KYUSHU; 32°18'N, 130°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: IV-3). See Fig. 2A. Goriyama. Kawaguchi, Ogawagyou-mura, Wa- tarai-gun (Jg^lP^hJHiP^JH PSMOj); Mie, HONSHU; 34°24'N, 136°37'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-34). See Fig. 2A. Gosho-son, Itano-gun (^i?fP1Slp|f^); Toku- shima, SHIKOKU; 34°09'N, 134°23'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 24: XI-35). See Fig. 2A. Goyosan ( £ ^ Oj ); hvate, HONSHU; ca. 39°12'N, 14r44'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60; cf. Study Group on the Present Status of Japanese Monkeys, 1977a, p. \{)).NE20. Goyosan, Hikoroichi-mura, Kesen-gun (^^fllj^P B tiltT;f>tE^Uj); hvate, HONSHU; 39°09'N, 141°4rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56:XLV-1). SeeFig. 2A. Gozaishi, Maruno-mura, Kitakoma-gun (db^0 fPlil©^^^^); Yamanashi, HONSHU; 35°44'N, 138°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII- 15). See Fig. 2A. Gozaishiyama vicinity. Seitetsu-mura, Kita- koma-gun (db&0?P^^^^^^Uj^); Yamanashi, HONSHU; 35°43'N, 138°21'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-14). SeeFig. 2A. Gozenyama, Hagiwara-cho, Masuda-gun (MB9 iPi^J^fflJ^lfUj); Gifu, HONSHU; 35°52'N, 137°16'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p.41:XXVIII-31). SeeFig. 2A. Hacchoyama, Yuge-mura, Kitakuwada-gun (db HBaiP^ijtfATOj); Kyoto, HONSHU; 35°12'N, 135°39'E; reported in questionnaire 128 FIELDIANA: ZOOLOGY survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXII- 18). See Fig. 2 A. Hachimori-machi, Yamamoto-gun (Lil^t^^PAl^ fflj); Akita, HONSHU; ca. 40°19'N, 140°02'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60; cf. Study Group on the Present Status of Japanese Mon- keys, 1977a, p. 10). Collected before 2002 by PRIKU staff; PRIKU, 1 (mandible and post- cranials only). NEl 1. Hachimoriyama, Kazamaura-mura, Shi- mokita-gun (TdbiPEPB^/i^A^lij); Ao- mori, HONSHU; 41°26'N, 141°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-20). See Fig. 2A. Hachioji-city (Ai-frfT); Tokyo Metropolis, HONSHU; ca. 35°39'N, 139°20'E; mtDNA samples collected before 1999 by Y. Kawa- moto (1998, p. 54). ^£777. Hachirouyama, Tahara-mura, Higashimuro-gun ( m # « ?P ffl J^ W A HP OJ ); Wakayama, HONSHU; 33°32'N, 135°54'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-43). See Fig. 2A. Hachisusanrin vicinity, Mori-mura, linan-gun ( IS it IP 1^ ^ a OJ # ); Mie, HONSHU; 34°2rN, 136°08'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-24). See Fig. 2A. Hagachizaki, Minamiizu-cho, Kamo-gun (M^ ^M^s:fflI):^Blli^) (= Hashozaki); Shizuoka, HONSHU; 34°41'N, 138°45'E; reported pre- sent in 1934 by N. Kuroda (1938, p. 1 12; 1940, p. 270). Provisioning initiated 1953-1955 (Akabori & Suzuki, 1977, p. 73; Study Group on the Present Status of Japanese Monkeys, 1977b, p. 23; Shidei et al., 1981, p. 17). Birth season, 1966, reported by Kawai et al. (1967, p. 39). Collected 19 Nov. 1978 by PRIKU staff; PRIKU, 1 (skeleton only). MtDNA samples collected ca. 1986-1991 by Hayasaka et al. (1991, p. 400). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). Microsatellite DNA analyzed before 1996 by Domingo-Roura et al. (1997, p. 358). Ex- ternal measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). MtDNA samples collected before 1998 by Y. Kawa- moto (1997, p. 33). A^£707. Hagi-machi, Abu-gun (PRf^lPMBJ); Yamaguchi, HONSHU; coordinates unknown; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII- 10). Not mapped. Hagiu, Futtsu-city ( ^ /$ TfT f^ ^ ); Chiba, HONSHU; 35°10'N, 139°50'E; collected 30 Mar. 1999 by PRIKU staff; PRIKU, 1 (skele- ton only). NE120. Hakkaisan, Higashi-mura, Minamiuonuma-gun (itl^)S^^WA^aj); Niigata, HONSHU; 37°10'N, 138°59'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII-7). See Fig. 2A. Hakkaisan. Jounai-mura, Minamiuonuma-gun (^^)SlP^rt^A^^aj); Niigata, HONSHU; 37°08'N, 138°59'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII-6). See Fig. 2 A. Hakkouzan, Goka-mura, Haibara-gun (^i^?P£ W^e3feUj); Shizuoka, HONSHU; 34°52'N, 138°06'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p.45:XXX-6). SeeFig. 2A. Hakkouzan, Shimokawane-mura, Haibara-gun ( ^ J^ ?P T Jll ^ ^ e Tfe UJ ); Shizuoka, HONSHU; 34°58'N, 138°04'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX-5). See Fig. 2A. Hakone-machi, Ashigarashimo-gun (S.ffiT?P^ ^ BO" ); Kanagawa, HONSHU; 35°11'N, 139°02'E; provisioning initiated in 1956 (Ta- naka & Masui, 1977, p. 62). Variable provi- sioning reported 1971-1977 by F. Fukuda (1988, p. 480). Intergroup dispersal reported 1966-1980 by F. Fukuda (2004, p. 54). NE103. Hakusan (fiOj); Ishikawa, HONSHU; 36°16'N, 136°46'E; field study begun in 1962, provi- sioning attempted in 1964 (Masui, 1977, p. 81). Blood samples collected before 1992 by No- zawa et al. (1991, p. 414; 1996, p. 6). Mi- crosatellite DNA analyzed before 1996 by Domingo-Roura et al. (1997, p. 358). External measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). NE58. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 129 Hamahara-mura, Oochi-gun (e^^P^J^^); Shimane, HONSHU; 35°03'N, 132°36'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-23). See Fig. 2A. Hamakita-city (^JbTti ); Shizuoka, HONSHU; ca. 34°48'N, 137°48'E; collected before 1991 by PRIKU staff; PRIKU, 3 (skeletons only). NE65. Handairayama-son. See Nagafuka vicinity. Hanto-cho, Itano-gun (^iJ^P^^fflJ); Tokii- shima, SHIKOKU; 34°09'N, 134°31'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-33). See Fig. 2A. Hanyama, Kamiyaku-cho, Kumage-gun (^^IP ±M:^fflI^Uj); Kagoshima, YAKUSHIMA ; 30°23'N, 130°23'E; collected Mar. 1983 by T. Maruhashi; PRIKU, 1 (skeleton only). Col- lected ca. 1983-1999 by PRIKU staff; PRIKU, 30 (skeletons only, including 7 incomplete). Collected 10 May 1984 by f. Oi; PRIKU, 1 (skull only). Collected Jun.-Sep. 1991 by N. Agetsuma; PRIKU, 3 (skeletons only). Mass mortality reported in 1998-1999 by G. Hanya et al. (2004, p. \^2)SW101. Hanyama-Segire, Kamiyaku-cho, Kumage-gun (^^^±M^fflI¥ajM^). See Yaku'^hima, Census Area 7. Harimochi, Nishitara-mura, Isa-gun ("^felPM ;*;^^t1-J^); Kagoshima, KYUSHU; 32°02'N, 130°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12:1-9). See Fig. 2A. Hashozaki. See Hagachizaki. Hasumi, Hasumi-mura, Oochi-gun (e^?P^^ M^^^H); Shimane, HONSHU; 34°51'N. 132°40'E; collected 6 Dec. 2000 by M. Aimi; PRIKU, 1 (skeleton only). SW62. Hata-gun. See Okuuchi-mura. Hata-mura, lishi-gun (ISS?P/^^W); Shimane, HONSHU; 35°08'N, 132°39'E; reported (as possibly temporary habitat) in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-32). See Fig. 2 A. Hatasho-cho, Echi-gun (^^IP^Sfflf); Shiga, HONSHU; ca. 35° 10' N, 136°16'E; mtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). SWIO. Hatofukiyama, Dota-mura, Kani-gun (RTJ/^lPih ffl ^ ii R^ UJ ); Gifu, HONSHU; 35=247^, 137°02'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 41: XXVIII-21). See Fig. 2A. Hatta, Taihei-mura, Minamiakita-gun (^^i^ffllP ;l!^¥:^Affl); Akita, HONSHU; 39°45'N, 140°irE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLIV-4). See Fig. 2A. Hattasan vicinity, Todoromi-mura, Toyono-gun {^m^±^ SM^ABBOJte); Osaka, HONSHU; 34°53'N, 135°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XIX-2). See Fig. 2A. Hida-sanmyaku (mountain range) (M^lllM); Nagano, HONSHU; ca. 36°25'N, 137°40'E; observed before 2004 by J. Ackerman (2003, p. 97). NE89. Hidaka Mountain Range, Kawakami-mura, Hi- daka-gun ( S SlPJlfi^ S S UjM); Waka- yama, HONSHU; 33°58'N, 135°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-9). See Fig. 2A. Hidarimaeyama vicinity, Tsuji, Momose-mura, Takashima-gun (l§E^W^Wit:£Bif Ujffe); Shiga, HONSHU; 35°26'N, 136°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXI V-9). See Fig. 2A. Hidaritsubute \'icinity. Torigoe-mura, Nomi-gun {^^^MU^&Mi^y, Ishikawa, HONSHU; 36°18'N, 136°37'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVI-4). See Fig. 2A. Hieizan (J±0Uj); Shiga, HONSHU; ca. 35°05'N, 135°50'E; >12 km troop movement, 26 Oct.-21 Nov. 1996, reported by G. Hanya et al. (2002, p. 417). MtDNA samples collected be- fore 2003 by Y. Kawamoto (2002, p. 60). SW56. Hieizan. Sakamoto-mura, Shiga-gun (^MIP^ ^WJtMOj); Shiga, HONSHU; 35°05'N, 135°5rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV-2). See Fig. 2A. Higakureyama vicinity, Iwato-son, Ni- 130 nELDIANA: ZOOLOGY shiusuki-gun (ffi Q tt?P^P ^ S H UJ ffe); Miyazaki, KYUSHU; 32°45'N, 131°25'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-30). See Fig. 2A. Higashi- and Nishi-Kurokiyama, Kamiimi-mura, Higashikunisaki-gun (!^P^?P±^II^^- S H ?K 111 ); Oita, KYUSHU; 33°37'N, 131°36'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16:111-21). See Fig. 2A. Higashihinobori, Kisuki-cho, Oohara-gun (AJ^ ^ ^ ;^ fflj K B ^ ); Shimane, HONSHU; 35°16'N, 132°55'E; collected 16 Sep.-22 Dec. 1994 by PRIKU staff; PRIKU, 3 (skeletons only, 1 without skull). SW59. Higashiina, Komagane-city {^^^l^Mi^M); Nagano, HONSHU; 35°45'N, 137°58'E; col- lected 23 Jul. 1998 by M. Aimi; PRIKU, 1 (skeleton only). NE78. Higashimikatagamori, Tokuda-mura/Nakagawa- mura/Sekiya-mura border area, Shusou- gun(j^^^t§Ba^^Ba)t. ^mmm^tm% E.y5 r By, Ehime, SHIKOKU; 33°54'N, 132°57'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 18:IX-9). SeeFig. 2A. Higashimisaki vicinity, Kuki-mura, Kita- muro-gun {it^U^%%.^'^%^^^)\ Mie, HONSHU; 34°01'N, 136°15'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-47). See Fig. 2A. Higashimiyamata vicinity, Kawakami, Ichi- nose-mura, Watarai-gun (J^^IP — ^i^^^Jil JiM'BU^); Mie, HONSHU; 34°20'N, I36°34'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-35). See Fig. 2A. Higashine-city (M^'J^); Yamagata, HONSHU; ca. 38°24'N, 140°25'E; mtDNA samples col- lected before 2003 by Y. Kawamoto (2002, p. 60). NE27. Higashino, Agematsu-machi, Kiso-gun {^mW> ±4i^ffll^i?); Nagano, HONSHU; 35°45'N, 137°43'E; collected 25 Mar. and 22 Jun. 1998 by M. Aimi; PRIKU, 5 (skeletons only). NE80. Higashinokawa vicinity, Oohuki-mura, Nii-gun (ff^^PA^^^myJIIftfe); Ehime, SHIKOKU; 33°51'N, 133°11'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: IX- 18). See Fig. 2A. Higashiogura-mura, Echi-gun (M^^^^^J^^^^); Shiga, HONSHU; 35°06'N, 136°24'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV-23). See Fig. 2A. Higashiyama National Forest, Yasuharakamin- ishi-mura, Kagawa-gun (#JI|?P$J^-hM^ ^lUm^^y, Kagawa, SHIKOKU; 34°08'N, 134°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 24: Xll-3). See Fig. 2A. Higashiyama vicinity, Kamihaya-mura, Nishi- muro-gun (H#ft?P±^*^^tKajfte); Wa- kayama, HONSHU; 33°49'N, 135°25'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-13). SeeFig. 2A. Higetayama vicinity, Takane-mura, Masuda-gun {^m^mm^^f^\hWy, a/u, honshu; 36°06'N, 137°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIII-34). See Fig. 2A. Higo (flE^) (= Figo); Kumamoto, KYUSHU; ca. 32°27'N, 130°46'E; reported present before 1830 by P. F. von Siebold (MS., 1830-1842, p. 1; cf Temminck, 1842, p. 10; Kuroda, 1938, p. 112; 1940,p. 271).5^<^9. Hikosanroku, Tsukinoki-mura, Shimoge-gun (T ^^P^tK^^^UjM); Oita, KYUSHU; 33°28'N, 130°59'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: 111-17). See Fig. 2A. Hinabatake, Kyonan-machi, Awa-gun (^^?PIS l^ffllBfSjjffl); Chiba, HONSHU; 35°08'N, 139°52'E; collected 17 Dec. 1997-2 Dec. 1998 by PRIKU staff; PRIKU, 6 (skeletons only). NE120. Hinagawa, Hasumi-mura, Oochi-gun (e^^^ MMWS^JII); Shimane, HONSHU; 34°51'N, 132°40'E; collected 18 Jun. 1998-15 Jan. 1999 by PRIKU staff; PRIKU, 3 (skeletons only). Collected 8 Dec. 2000 by M. Aimi; PRIKU, 1 (skeleton only). SW62. Hinata, Mitake-mura, Kiso-gun (TKe^PH^^t^ B fS]); Nagano, HONSHU; 35°50'N, 137°39'E; FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 131 collected 30 Jul. 1998 by M. Aimi; PRIKU, 1 (skeleton only). NE82. Hinatayama, Oomachi-city (AffljTfJ B [d]i1|); Nagano, HONSHU; 36°32'N, 137°47'E; col- lected 19 Sep. and 19 Dec. 1997 by M. Aimi; PRIKU, 2 (skeletons only). NE9Q. Hino-cho, Gamou-gun (S^?PS©fflI); Shiga, HONSHU; ca. 35°01'N, 136°15'E; mtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). SW16. Hinohara-mura, Nishitama-gun (B^BIP^J^ ^); Tokyo Metropolis, HONSHU; ca. 35°43'N, 139°10'E; mtDNA samples collected before 1999 by Y. Kawamoto (1998, p. 54; 2002, p. 6Q).NE112. Hinokiyama, Toumi-son, Higashiusuki-gun (^ Q^IPK^W^UJ); Miyazaki, KYUSHU; 32°38'N, 131°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-26). See Fig. 2 A. Hinokuchi, Yaku-cho, Kumage-gun (^^?PM :^ HI ffi y P ); Kagoshima, YAKUSHIMA; 30°17'N, 130°38'E; collected 25 Feb. 1989 by PRIKU staff; PRIKU, 5 (skeletons only, in- cluding 1 with mismatched mandible). SIV102. Hinoyama, Shirakawa-mura, Chichibu-gun {^1>t ?P S Jll ^ S © UJ ); Saitaina, HONSHU; 35°56'N, 138°59'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 50: XXXVII-5). See Fig. 2A. Hippo-mura, Igu-gun (#M^^^^ : Hfil^M Uj MX^); Toyama, HONSHU; ca. 36°32'N, 137°14'E; mtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). NE57. Hotakayama vicinity, Katashina-mura, Tone-gun (^IJ^lP>^pp^^^li|ffe); Gunma, HONSHU; 36°47'N, 138°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XL- 14). See Fig. 2 A. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 133 Hotakayama vicinity. Kavvaba-mura, Tone-gun {mWi^}\\^^^M\Ui^): Gimma. HONSHU; 36°47'N. 138°50'E; reported in questionnaire sur\ey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XL- 13). See Fig. 2A. Hotokedani, Obama-city (d^MTtTi^^); Fiikui. HONSHU; 35°31'N, 135°43'E; collected Jul. 1994 by PRIKU staff; PRIKU, 1 (postcranials only). 'SW45. Hotokeiwa, Hirao-mura, Shimotakai-gun (TM ^ ?P ¥ ^ ^ ^iA ^ ): Nagano, HONSHU; 36°44'N. 138°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974. p. 46: XXXI-26). See Fig. 2 A. Houin, Kisuki-cho, Oohara-gun (AJ^^^/^fflJ ^ ep ); Shimane, HONSHU; 35°13'N, 132°57'E; collected 27 May-12 Dec. 1994 by PRIKU staff; PRIKU, 3 (skeletons only). SW59. Houtakusan. Kamiyoshi-mura, Izushi-gun (ttjS IP tt H ^ ^ k OJ ); Hyogo. HONSHU; 35°30'N, 134°55'E; reported in questionnaire sur\ey conducted in 1923 by K. Hasebe (hvano. 1974. p. 30: XVlII-21). See Fig. 2A. Houtosan. Nagatoro-machi, Chichibu-gun (^^ ^:i:5^fflTi^Uj) (= Hodosan; Nagatoro): Saitama. HONSHU; 36°04'N, 139°05'E; mon- key park population, translocated in 1962 from Gagyusan. Okayama Prefecture (Kanaizuka, 1977, p. 43; cf. Study Group on the Present Status of Japanese Monkeys, 1977b, p. 23). Birth season. 1963-1966. reported by Kawai et al. (1967, p. 39). External measurements taken before 1997 by Hamada et al. (1996a. pp. 98. 99). NE118. Houzaw ayama. Kaw akami-mura, Kumano-gun (^Pipjl|±;^i;RUj); Kyoto, HONSHU; 35°33'N. 134°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXII-6). See Fig. 2A. Ibuki-cho. Sakata-gun (J^BaiP^R^BO"); Shiga, HONSHU; ca. 35°23'N. 136°23'E; mtDNA samples collected before 2003 by Y. Kawa- moto (2002. p. 60). SW7. Ibukiyama. Ibuki-mura, Sakata-gun (^ffllP'^RX ^ # R^ OJ ); Shiga, HONSHU; 35°25'N, 136°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV- 15). See Fig. 2 A. Ibukiyama, Samegai-mura, Sakata-gun (^ffl^ M^^^R^OJ)^; Shiga, HONSHU; 35°19'N, 136°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974, p. 38: XXIV- 18). See Fig. 2A. Ibukiyama. Suijou-mura. Sakata-gun (^B91P# Mk^D^Ul); Shiga. HONSHU; 35°23'N, 136°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974, p. 38: XXIV- 16). See Fig. 2 A. Ibushikanyama vicinity. Yanadani-mura. Kamiukena-gun (±>?:iX?P*JP^^-r7^5>t 04^); Ehime, SHIKOKU; 33°31'N, 132°59'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 21: IX-2I). SeeFig. 2A. Ichifusa vicinit}'. Nishimera-son. Ko\ii-gun {% ^^Pe^l^M^TfJMffe); Miyazaki, KYUSHU; 32°I4'N. 131°09'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-15). See Fig. 2 A. Ichiki-cho, Hioki-gun ( B a?PTtT3ifflT); Kago- shinia. KYUSHU; 31°35'N, 130°21'E; Jomon subfossils (age, 2.3-12 Ka) reported by K. Kawanaka (1973, p. 1 18). SW94. Idodani vicinit\. Shimonokawa-mura, Ichishi-gun (-^x^T;^JII^#P^ftfe); Mie, HONSHU; 34°33'N. 136°2rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 37: XXIII-20). See Fig. 2A. Iga-cho. Ayama-gun ( PrI Oj IP # M fiJ ); Mie, HONSHU; ca. 34°45'N. 136°10'E; collected 23 Oct. 1979-14 Dec. 1983 by PRIKU staff; PRIKU, 3 (skeletons only). SW20. Ihama. Minamiizu-cho. Kamo-gun (MMIPM"^ SfflJ^^); Shizuoka, HONSHU; 34°4nsJ, 138°46'E; group fission in 1965 and 1967 re- ported by Study Group on the Present Status of Japanese Monkeys (1977b, p. 23). Blood sam- ples collected before 1992 by Nozawa et al. (1991. p. 414; 1996.p. 6)..V£/07. lidesan vicinit>. Toyomi-mura. Higashikan- bara-gun (^MI^iPMH^tSWOj^); Mi- gata, HONSHU; 37=41'N, 139=35'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 47: XXXII- 18). See Fig. 2A. Iketaniyama vicinit>'. Kitagawa-mura, Aki-gun 134 FIELDIANA: ZOOLOGY ($S?PdbJI|^)til#a4fte); Kochi, SHIKOKU; 33°29'N, 134°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-48). See Fig. 2 A. Ikuno, Doujo-mura, Arima-gun (W/^^PjM^^ ^i?); Hyogo, HONSHU; 34°50'N, 135°16'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-28). See Fig. 2A. Imaichi-city (^T^TTl:); Tochigi, HONSHU; ca. 36°47'N, 139°39'E; collected 1 Dec. 1984-9 Nov. 1986 by TPM staff; TPM, 8 (skulls only; excluding 2 skulls not seen, data from speci- men list). NE43. Imanishi, Hasumi-mura, Oochi-gun (e^^P^M W ); Shimane, HONSHU; 34°52'N, 132°37'E; collected 7 Dec. 1996-8 May 1997 by PRIKU staff; KUPR, 6 (skeletons only). Collected 1 Mar. 2000 by M. Aimi; PRIKU, 1 (skeleton only). SW62. Inabe-cho, Inabe-gun (M^lPM^fflJ); Mie, HONSHU; ca. 35°07'N, 136°35'E; mtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). SW13. Inagawa-goryorin, Ookuwa-mura, Nishichi- kuma-gun (ffim»?PA^^t^i9PJII^3^«); Nagano, HONSHU; 35°40'N, 137°42'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXXI-5). See Fig. 2A. Inaodake vicinity, Tashiro-mura, Kimotsuki-gun ( if Jl IP ffl ft W ^ m S ftfe ); Kagoshima, KYUSHU; 3r07'N, 130°53'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-17). See Fig. 2A. Ine-cho, Yosa-gun {^Wi^^^^); Kyoto, HONSHU; 35°43'N, 135°17'E; captured alive 5-1 1 Mar. 1980 by JMC staff, died in captivity 6-25 Apr. 1980; JMC, 4 (skulls only). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; \996,^.6).SW48. Inokamireien, Matsukawa-mura, Kitaazumi-gun (db^m?P^AMII^^(D#Sg); Nagano, HONSHU; 36°24'N, 137°49'E; collected 20 Nov. 1998 by PRIKU staff; PRIKU, 1 (skele- ton only). NE92. Inouesanrin, Inbi-mura, Minamiamabe-gun (]^ ^^P?PHM^#y±Uj^); Oita, KYUSHU; 32°57'N, 131°44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: III- 11). See Fig. 2A. Inunaki, Kuchimyougata-mura, Gujo-gun (^Ji ^U^^l5¥iii^%); Gifu, HONSHU; 35°46'N, 137°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41 : XXVlII-26). See Fig. 2A. Inunakisan, Ootsuchi-mura, Sennan-gun (:^^^ A±^;'^lliaj); Osaka, HONSHU; 34°20'N, 135°22'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XIX-6). See Fig. 2A. Inuyama-city (;^lJL|Tt]); Aichi, HONSHU; ca. 35°23'N, 136°56'N; mtDNA sample collected before 2003 by Y. Kawamoto (2002, p. 60). NE6L Inyou Mountain Range vicinity, Funo-son, Fu- tami-gun ()5lHlJ^^^^^|iajM-^); Hi- roshima, HONSHU; 34°52'N, 132°47'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XlV-38). See Fig. 2A. Inyou Mountain Range vicinity, Kimita-son, Futami-gun ()5(H?P^Ba^ltliajM- w); Hiroshima, HONSHU; 34°54'N, 132°51'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XIV-37). See Fig. 2A. Inyou Mountain Range vicinity, Sakugi-son, Futami-gun {7^^W^^i^^^m\hU-w); Hiroshima, HONSHU; 34°54'N, 132°44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XIV-39). See Fig. 2A. Iriokuyama vicinity, Okouchi-mura, Nishi- tama-gun (H^ft?P^J^)RlI*l^^^AllLilffe); 7b- l^o Metropolis, HONSHU; 35°46'N, 139°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXIV-2). See Fig. 2A. Irisawai, Ooshika-mura, Shimoina-gun {Y-^M IP ;^ J^ ^ A )R # ); Nagano, HONSHU; 35°35'N, 138°04'E: collected 7 Dec. 1998 by PRIKU staff; PRIKU, 1 (fragmented skull only). NE70. Iriyamagawaura National Forest, Kuni-mura, Agatsuma-gun (■ >/N a nx\h}\mmm ^); Gunma, HONSHU; 36°39'N, 138°40'E; FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 135 reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XL-10). SeeFig. 2A. Irokawa-mura, Higashimuro-gun (^^SlPfeJH ^ ); Wakayama, HONSHU; 33°40'N, 135°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-46). See Fig. 2A. Isehara-city (^^J^l^J); Kanagawa, HONSHU; ca. 35°24'N, 139°18'E; mtDNA samples col- lected before 1998 by Y. Kawamoto (1997, p. 33). NE106. Ishidosan, Nii-mura, Hikami-gun (7KJL?Plff^ ^^P \1\); Hyogo, HONSHU; 35°06'N, 135°02'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-26). See Fig. 2A. Ishigagawa, Shimesasu, Nakagawa-mura, Wata- rai-gun (Jg^lPt JH^jijiJaS ^)R[); Mie, HONSHU; 34°24'N, 136°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-33). See Fig. 2A. Ishizuka, Ichihara-city ( Tfi IP! rfT ^ i^ ); Chiba, HONSHU; 35°16'N, 140°07'E; collected 1 Sep. 1997-30 Mar. 1999 by PRIKU staff; PRIKU, 4 (skeletons only). NE127. Isokoen. See Onoaida. Issou, Kamiyaku-cho, Kumage-gun (^^IPJiM X^ - My, Kagoshima, YAKUSHIMA; 30°26'N, 130°28'E; collected 25 Feb. 1989 by PRIKU staff; PRIKU, 1 (skeleton only). SWIOO. Issou-Miyanoura. See Yakushima, Census Area 2. Ita, Heda-mura, Tagata-gun {Wil5WF^¥i^ ffl); Shizuoka, HONSHU; 34°59'N, 138°47'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX-17). SeeFig. 2A. Italy. See Rome. Itamayama, Saikihon-son, Akaiwa-gun (#^^ feffi^^^W^iUj); Okayama, HONSHU; 34°52'N, 134°07'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV- 19). See Fig. 2A. Itotaniyama, Ichiyama-mura, Oochi-gun (e-^^ rtT|l|;^^#Uj); Shimane, HONSHU; 34°57'N, 132°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28:XVI-14). SeeFig. 2A. Itsukaichi, Akiruno-city ($$^i?rfT£Srfi"); Tokyo Metropolis, HONSHU; ca. 35°44'N, 139°13'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 6Q).NEU2. Itsuki-mura, Kuma-gun (^H^S^K^); Ku- mamoto KYUSHU; 32°24'N, 130°50'E; blood samples collected before 1992 by Nozawa et al. (1991, p. 414; \996,^. 6). SW90. Iwaidani vicinity, Aiga-mura, Kitamuro-gun (db ^«lP*IS^^##ffe); Mie, HONSHU; 34°06'N, 136°13'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-45). See Fig. 2 A. Iwaidaniyama, Ibara-mura, Oochi-gun (b^^ #i^^^^^Uj); Shimane, HONSHU; 34°55'N, 132°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI- 18). See Fig. 2A. Iwamayama, Ishiyama-mura, Shiga-gun (^M?P S ill ^^ Pel 111); Shiga, HONSHU; 34°57'N, 135°53'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38:XXIV-1). SeeFig. 2A. Iwamiyama National Forest vicinity, Iwami- sannai-mura, Kawabe-gun (/Rl32.?P^^Hrt ^^.^OlSWWffe); Akita, HONSHU; 39°44'N, 140°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLIV-3). See Fig. 2A. Iwamiyama vicinity, Toyomatsu-son, Jinseki-gun ( 4$ ^ iP W ^A^ ^ ^ m UJ ffe ); Hiroshima, HONSHU; 34°48'N, 133°21'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-34). See Fig. 2A. Iwase National Forest, Yamase-mura, Kita- akita-gun {it^km^\^M^^MmM^); Akita, HONSHU; 40°20'N, 140°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLIV-7). See Fig. 2A. Iwashina-mura, Kamo-gun ( S j^ ^ ^ ^ ^ ); Shizuoka, HONSHU; 34°44'N, 138°48'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX-23). See Fig. 2A. 136 FIELDIANA: ZOOLOGY Iwataniyama, Tsuro-mura, Aki-gun ($S^/$S ^^^Uj); Kochi, SHIKOKU; coordinates unknown; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-50). Not mapped. Iwaya, Anabuki-son, Mima-gun (M/^^TvlIl^^ ^ M ); Tokushima, SHIKOKU; 34°02'N, 134°10'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-25). See Fig. 2A. Iwayado, Ikeda-son, Yazu-gun (AH^ylfeffl^ ^ ^ ^ ); Tottori, HONSHU; 35°19'N, 134°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 29: XVII-3). See Fig. 2A. Iwayama, Tarou-mura, Ichishi-gun (— ^IPAfiP ^ ^ ^ Uj ); Mie, HONSHU; 34°32'N, 136°10'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36:XXIII-18). SeeFig. 2A. Izu Peninsula. See Hagachizaki. Jadani vicinity, Yoshinotani-mura, Ishikawa-gun ( ^ JIIIP ^ if ^ ^ !te # ^ ); Ishikawa, HONSHU; 36°21'N, 136°39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVI-3). See Fig. 2A. Japan ( 0 ^ ); probably HONSHU; 33°28'- 41°34'E, 130°53'-142°04'E; collected 1823-1829 by P. F. von Siebold (MS., 1830-1842, p. 1; cf. Temminck, 1842, p. 10; Holthuis & Sakai, 1970, p. 25); RMNH, 11 (including 4 skins only and 5 skulls only). Collected 1830-1832 by H. Burger (cf. Holthuis & Sakai, 1970, p. 37); RMNH, 1 (skeleton only). Specimen received by BM(NH) from RMNH before 1843, probably collected 1823-1832 by P. F. von Siebold or H. Burger (cf Gray, 1843, pp. x, 8; 1870, p. 31); BM(NH), Collected in 1874 by Mr. v. Hilgendorf; ZMB, 1 (skeleton only). Pur- chased 1900-1901 by K. A. Haberer (cf Schweyer, 1909, p. 8); ZSBS, 20 (6 skins only, 14 skulls only). Collected before 1887 by M. Soller; MNHN, 1. Collected before 1966 by J. Itani, AIUZ, 2 (skulls only). Not mapped. Jedo (>IP). See Tokyo. Jigokudani, Yamanouchi-machi, Shimotakai-gun (Tit^^POjy rtBTTitki^^) (= Shiga-A); Nagano, HONSHU; 36°44'N, 138°25'E; birth season, 1962-1966, reported by Kawai et al. (1967, p. 38). Provisioning initiated in 1963 (Shidei et al., 1981, p. 16). Field study re- ported by E. Tokita and S. Hara (1975, p. 24). Field study conducted 1994-1996 by M. Iwago and H. Iwago (1999, pp. 154, 160). NE49. Jigokudani vicinity, Shoukawa-mura, Oono-gun (Ai?^SJIIW±tk3i^ffe); Gifu, HONSHU; 35°59'N, 137°10'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIII-38). See Fig. 2 A. Jinkurousanrin-oku, Hibara-mura, Yama-gun (IP ^iP^J^vWSABPOj^ll); Fukushima, HONSHU; 37°42'N, 140°02'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 52: XLI-7). See Fig. 2A. Jinrio, Kamiyama-cho, Myouzai-gun (^ffi^P-f^ lijffll^^) (= Jinryo; Tuino), 150 m elevation; Tokushima, SHIKOKU; ca. 34°00'N, 134°15'E; collected 15 and 20 Feb. 1905 by M. P. Anderson (in Thomas, 1906 ["1905"], p. 336; cf Kuroda, 1940, p. 272; Napier, 1981, p. 27); BM(NH), 3. SW70. Jizoutouge vicinity, Tomikawa-mura, Mi- namikoma-gun (^^MWmM^ii'^MW^); Yamanashi, HONSHU; 35°13'N, 138°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-1). SeeFig. 2A. Jokohama. See Yokohama. Jujotouge vicinity, Tomisato-mura, Nishi- muro-gun (S^«^SM^t + 3tllt#ftfe); Wa- kayama, HONSHU; 33°45'N, 135°37'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-21). SeeFig. 2A. Junigatouge, Nagahama-mura, Minamitsuru-gun (itl[PmPM^^+-IfflT); Oita, KYUSHU; 32°48'N, 131°56'E; blood samples collected before 1992 by No- zawa et al. ( 1 99 1 , p. 4 1 4; 1 996, p. 6). SW82. Kamagatani, Kamiijira-mura, Yamagata-gun (ill m?P±#gm^llR Oj ); Nagano, HONSHU; 35°21'N, 137°58'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXI-48). See Fig. 2A. Kamiseinaiji, Seinaiji-mura, Shimoina-gun (T# MWm\k^^ ±M\H^); Nagano, HONSHU; 35°30'N, 137°4rE; collected 28 Oct. 1998 by M. Aimi; PRIKU, 1 (skeleton only). Collected 25 Nov. 1998 by PRIKU staff; PRIKU, 1 (fragmented skull only). NE73. Kamitakarazawa, Higashisawa-mura, Minami- murayama-gun (]^^ Uj ?P^}R:14±S)H); Yamagata, HONSHU; 38°12'N, 140°27'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLII-13). SeeFig. 2A. Kamitakashiri, Nanukaichi-mura, Kanoashi-gun H^^m^Z^l^^hM^y, Shimane, HONSHU; 34°23'N, 131°55'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-4). See Fig. 2A. Kamiyaku-cho [1], Kumage-gun (^^lP_hM^ fflj); Kagoshima, YAKUSHIMA; ca. 30°25'N, I30°35'E; blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 7). SWIOO. Kamiyaku-cho [2], Kumage-gun (rI^^PJiM^ HTT); Kagoshima, YAKUSHIMA; 30°16'N, 130°29'E; collected 26 Nov. 1977 by PRIKU staff; PRIKU, 1 (skeleton only). Collected 9 Jun. 1989 by S. Azuma; PRIKU, 14 (skeletons only). SWIOI. Kamou, Kimitsu-city ( ^ >$ r|T H ^ ); Chiba, HONSHU; 35°13'N, 140°07'E; collected 30 Jul. 1996-30 Mar. 1999 by PRIKU staff; PRIKU, 7 (skeletons only, including 1 with fragmented skull). NE126. Kanakidoyama, Kamitakara-mura, Yoshiki-gun (^^?P±S^^7KFlij); Gifu, HONSHU; 36°20'N, 137°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIII-36). See Fig. 2A. Kanasumi National Forest, Kijo-son, Koyu-gun (Jft^iPTK^^J^J^aW^); Miyazaki, KYUSHU; 32°14'N, 131°24'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-18). See Fig. 2A. Kanatokoyama, Aihashi-mura, Izushi-gun (tij^ IP-^M^^iiUj); Hyogo, HONSHU; FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 139 35°24'N, 134°56'E; reported in questionnaire sur\ey conducted in 1923 by K. Hasebe (hvano, 1974, p. 30: XVIII- 19). See Fig. 2 A. Kanatokoyama, Itoi-mura, Yabu-gun (M5^1P^ ^^^iiUl); Hyogo. HONSHU; 35°2rN, 134°53'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30:XVIII-16). SeeFig. 2A. Kanaya-cho. Arida-gun (WBaiP^MfflJ): Wa- kayama. HONSHU; ca. 34°04'N, 135°15'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002. p. 60). SW31. Kanayama. Satsuma-cho. Satsuma-gun (SS?P SSfflJ^Uj): Kagoshima. KYUSHU; 31°55'N. 130°36'E; collected 24 Jan. 2000 by M. Aimi; PRIKU. 5 (skulls only). SW92. Kanazavvayama \icinity. Miyagase-mura. Aiko-gun ( g ¥ ?P ^ V ^ ;^ ±^R Uj ^ ); Kanagaua, HONSHU; 35°31'N, 139°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 50: XXXV-7). See Fig. 2A. Kanba. Katsuyama-cho. Maniwa-gun (XfilPM Lijffllttfi); Okayama. HONSHU; 35°05'N. 133°42'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 27:XV-13). SeeFig. 2A. Kanbanotaki. Katsuyama-cho. Maniwa-gun (Ml JS?PBlijfflIttJS(7)>t); Okayama. HONSHU; 35°07'N. 133°4rE; provisoned group; birth season. 1962. reported by Kawai et al. (1967. pp. 37. 38). SW55. Kandayama. Hongou-cho. Kitashitara-gun (dbix ^A^iPfflTttBaOJ); AichL FIONSHU; 35°05'N. 137°42'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 40: XXVII-4). See Fig. 2 A. Kanegaryumorisankei. Nakayama-mura. Aki-gun ( $^^' ?P ^ 111 ;^ M Md^MlU): Shimane, HONSHU; 34°55'N, 132°34'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI- 19). SeeFig. 2A. Kannon. See Kannonzaki. Kannonyama. Yotsu-mura. Takaoka-gun (Si^ IP ^')$ ^ ^ # OJ ); KochL SHIKOKU; 33°irN, 133°13'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974. p. 22: X-16). See Fig. 2A. Kannonzaki. Kamiyaku-cho. Kumage-gun {M.^ lP±M^ffltia^*f); ^Kagoshima. YAKUSHIMA; ca. 30°23'N, 130°23'E; ob- served Oct. 1977 by T. Maruhashi (1982, p. 318; cf. Takasaki & Masui. 1984, p. 311). SWIOI. Kanotoyama vicinity. Hinohara-mura, Nishi- tama-gun (S^0iP^JI^#P Ujffe); Tokyo Metropolis. HONSHU; 35°43'N, 139°07'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXIV-1). SeeFig. 2A. Kanoya-city (^MTfT); Kagoshima. KYUSHU; ca. 31°22'N, 130°46'E; mtDNA samples col- lected before 2003 by Y. Kawamoto (2002, p. 60). SW95. Karamatsu, Miyoshi-son, Atetsu-gun (Po[^^|| ^^0^i); Okayama, HONSHU; 34°58'N, 140 FIELDIANA: ZOOLOGY 133°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27:XV-11). SeeFig. 2A. Karasugawa, Nishihodaka-mura, Minamia- zumi-gun (]^$ft?PffiM>S^,^JI|); Nagano, HONSHU; 36°19'N, 137°51'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI- 15). See Fig. 2A. Karasugawa National Forest, Karasugawa-mura, Minamiazumi-gun ( j% ^ S IP B J i I W .i J 1 1 S #^); Nagano, HONSHU; 36°16'N, 137°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-14). SeeFig. 2A. Karekidani, Higashiyashiki, Shikiya-mura, Hi- gashimuro-gun (^^W:^WiE.^%M.Wi^^ ^ ); Wakayama, HONSHU; 33°50'N, 135°49'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34:XX-51). SeeFig. 2A. Karioyama vicinity, Yahata-son, Yamagata-gun ( Ol m ?P A 1i ^ XIJ M UJ ffe ); Hiroshima, HONSHU; 34°41'N, 132°11'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-6). See Fig. 2A. Karitayama, Takayama-mura, Kamitakai-gun (_h m^W>m\^¥iM^\h); Nagano, HONSHU; 36°41'N, 138°20'E; collected 1 Nov. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE49. Karuizawa-machi, Kitasaku-gun (db'fe^^S^ >RlflT); Nagano, HONSHU; ca. 36°21'N, 138°38'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE45. Kasagi, Kashiwazaki-mura, Watarai-gun (S^ ^^fillit^^TK); Mie, HONSHU; 34°18'N, 136°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-30). See Fig. 2A. Kashiji vicinity, Ichiu-son, Mima-gun (H/^IP — ^ t^ @ itk fife ); Tokushima, SHIKOKU; 33°56'N, 134°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-23). See Fig. 2A. Kashima (Eft); Ehime, KASHIMA; 32°57'N, 132°27'E; onset of provisioning in 1955 re- ported by S. Yoshihiro (1983, p. 41; cf Edito- rial Committee of Nihonzaru, 1977, p. 112). Blood samples collected before 1992 by No- zawa et al. (1991, p. 414; 1996, p. 6). SW81. Kashima, Nishisotoumi-mura, Minamiuwa-gun (l^^^^lPS^^^Jift); Ehime, KASHIMA 32°57'N, 132°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18: IX- 1). See Fig. 2A. Kashima-machi, Soma-gun ( ^1 M IP E ft BJ ); Fukushima, HONSHU; ca. 37°42'N, 140°58'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE59. Kashio, Ooshika-mura, Shimoina-gun (T#i§P?P AMWMi^); Nagano, HONSHU; 35°36'N, 138°03'E; collected 18 Dec. 1997 by M. Aimi; PRIKU, 1 (skeleton only). Collected 14 Mar. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE70. Kashiwabarayama foothills, Kaminada-mura, Tsuna-gun (^=glP±yllWttJ^ajM); Hyogo, AWAJISHIMA; 34°16'N, 134°53'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-2). See Fig. 2A. Kashiwajima, Okuuchi-mura, Hata-gun ("l^^^ Urt^ttft); Kochi, SHIKOKU; 32°47'N, 132°40'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21:X-1). SeeFig. 2A. Kashiyama vicinity, Funatsuki-mura, Hidaka-gun ( B S IP ^q' * ^ a Ol ftfe ); Wakayama, HONSHU; 33°57'N, 135°17'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-7). See Fig. 2A. Kasshizan, Nishigou-mura, Nishishirakawa-gun O a )Rl IP a ^ ;^^f ¥ ^ UJ ); Fukushima, HONSHU; 37°irN, 140°09'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 52: XLl-5). See Fig. 2A. Kasuga-cho, Hikami-gun ( tR ± IP # 0 fflj ) Hyogo, HONSHU; ca. 35°09'N, 135°10'E collected 23 and 27 Feb. 1980 by JMC staff; JMC, 7 (5 skulls only, 2 skeletons only). Blood samples collected before 1992 by Nozawa et al (1991, p. 414; 1996, p. 6). SW40. Kasugadaira, lijima-machi, Kamiina-gun (_h'^ 3P?PI5ftfflT#B¥); Nagano, HONSHU; 35°42'N, 137°54'E; collected 30 Oct. and 2 FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 141 Dec. 1997 by M. Aimi; PRIKU, 4 (skeletons only, including 2 with fragmented skulls). Collected 12 Nov. and 2 Dec. 1997 by PRIKU staff; PRIKU, 2 (skeletons only, fragmented skulls). NE77. Kasuge-mura, Yamamoto-gun (|1|^^^^^); Akita, HONSHU; 40°17'N, 140°18'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLlV-1 1). See Fig. 2A. Kataiwa National forest, Kamigou-mura, Kami- hei-gun (±r^#?P±^^>^^aWW); Iwate, HONSHU; 39°15'N, 141°36'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Mito, 1989, p. 25). See Fig. 2A. Katamukiyama, Hakusan-mura, Oono-gun (Ai? ^PfiOj^^Oj); Oita, KYUSHU; 32°49'N, 131°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16:111-4). See Fig. 2A. Kataoka-mura, Ika-gun ('^^^jnf^^^); Shiga, HONSHU; 35°33'N, 136°12'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV- 11). See Fig. 2A. Katsuojisan, Toyokawa-mura, Mishima-gun (H IIPWJH^SM^UJ); Osaka, HONSHU; 34°50'N, 135°33'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XIX-3). See Fig. 2A. Katsurashima, Nakagawa-mura, Kamiina-gun (±#i9P?P4'JII^Sft); Nagano, HONSHU; 35°36'N, 137°56'E; collected 7 Jul. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE71. Katsuyama-cho, Maniwa-gun (M^lPMlilfflJ); Okayama, HONSHU; 35°05'N, 133°38'E; blood samples collected before 1992 by No- zawa et al. (1991, p. 414; 1996, p. 6). Field study reported by Itoigawa et al. (1992, p. 58). External measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). SW54. Kawabe-machi, Kawabe-gun (yRlaS^PyRjiSfflJ); Akita, HONSHU; ca. 39°38'N, 140°13'E; mtDNA extracted from hunting trophies before 2003 by T. Agatsuma and M. Ishigami (2002, p. 79). NE14. Kawachi vicinity, Mugi-cho, Kaifu-gun (^pP?P # tt fflj )Rr rt flk ); Tokushima, SHIKOKU; 33°40'N, 134°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-9). See Fig. 2 A. Kawachiyama, Kawachi-mura, Nakakanbara-gun (4^aj^^JI||*l^JllrtUj); Niigata, HONSHU; 37°4rN, 139°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXIl-15). See Fig. 2A. Kawaguchi-ko ()RrP)^); Yamanashi, HONSHU; 35°31'N, 138°45'E; two groups, enclosed and provisioned before 1978, reported by Study Group on the Present Status of Japanese Mon- keys (1977b, p. 23). Translocation from Kamae and Shodoshima reported by T. Shidei etal. (1981,p. \1).NE97. Kawajionsen, Fujiwara-machi, Shioya-gun {^^ ?P^J^fflIJI|>^;U;7^); Tochigi, HONSHU; 36°53'N, 139°42'E; collected 8 Dec. 1986 by TPM staff; TPM, 1 (not seen, data from specimen list). NE41. Kawakami-mura, Yoshino-gun CRiJ^PJH-h^); Nara, HONSHU; 34°18'N, 136°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXI-7). See Fig. 2A. Kawamatasanrin, Mazuma-mura, Hidaka-gun ( B ift ?P M S W Jll 3104 ^ ); Wakayama, HONSHU; 33°51'N, 135°19'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX- 10). See Fig. 2A. Kawanaka-mura, Hidaka-gun (0S^JII4'W); Wakayama, HONSHU; 33°56'N, 135°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-8). See Fig. 2A. Kawanedake National Forest vicinity, Kawa- saki-mura, Shibata-gun (^Ba?PJI|lli^WJII a"^ aWWffe); Miyagi, HONSHU; 38°11'N, 140°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLlII-5). See Fig. 2A. Kawanishi, Matsukawa-mura, Kitaazumi-gun (db ^•?P;^iJIIWJI|ffi); Nagano, HONSHU; 36°26'N, 137°50'E; collected 22 Feb. 1998 by M. Aimi; PRIKU, 2 (skeletons only). NE92. Kawara, Kamiyaku-cho, Kumage-gun (^^^-h M:^fflTJI|J^); Kagoshima, YAKUSHIMA; ca. 30°21'N, 130°23'E; observed Jul. 1975 by T. Maruhashi (1982, p. 318; cf. Takasaki & Ma- 142 FIELDIANA: ZOOLOGY sui, 1984, p. 311). 5^7 r?/. Kawaradake (##S); Fukuoka, KYUSHU; 33°40'N, 130°51'E; blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996,p. 6).5Pf<^^. Kawaradake, Kawara-machi, Tagawa-gun (B9JII iP##fflT##|5); Fukuoka, KYUSHU; 33°28'N, 130°44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 17: VII-2). See Fig. 2 A. Kawaratai vicinity, Nishimeya-mura, Na- katsugaru-gun (4^)*felPffii Jl^Jlli^¥ffe); Aomori, HONSHU; 40°3rN, 140°16'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-7). See Fig. 2A. Kawarayama, Tsutasawa-mura, Shisou-gun (5^ ^ 15 M )R W )Rl Jl Lij ); Hyogo, HONSHU; 35°02'N, 134°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-6). See Fig. 2A. Kawasuso, Seinaiji-mura, Shimoina-gun {~^^M ^m\H^^]\\^); Nagano, HONSHU; 35°28'N, 137°42'E; collected 19 Oct. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE73. Kawatayama vicinity, Miyama-son, Oe-gun (^ ffl^HLl4;f4Jllfflajffe); Tokushima, SHIKOKU; 34°00'N, 134°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-27). See Fig. 2A. Kawaurayama, Itadori-mura, Mugi-gun (^'ffi^P ^IX^^JIhHOj); Gifu, HONSHU; 35°44'N, 136°48'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41:XXV1II-16). SeeFig. 2A. Kayano vicinity, Oshiro-mura, Mikata-gun (H^ ^ dMt ^ ^ i? -te ); Hyogo, HONSHU; 35°25'N, 134°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVllI-10). See Fig. 2A. Kayano-Nanamagari, Kisarazu-city (^M/$rfi^ i? -b te ); Chiba, HONSHU; 35°20'N, 140°05'E; collected 6 Nov. 1996-12 Jan. 1999 by PRIKU staff; PRIKU, 3 (skeletons only, in- cluding 1 with fragmented skull). NE128. Kensozan, Kitaura-mura, Shozu-gun (d'^S.lPdb )i ^^t lll^ llil 111 ); Kagawa, SHODOSHIMA; 34°32'N, 134°16'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 24: XII-7). See Fig. 2A. Kensozan, Kusakabe-mura, Shozu-gun (d'^SlP ^Mt^lllJ^lliflOj); Kagawa, SHODOSHIMA; 34°30'N, 134°19'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 24: XII-9). See Fig. 2A. Kensozan, Oobe-mura, Shozu-gun ()- ll!^ Ilil Ol ); Kagawa, SHODOSHIMA; 34°29'N, 134°21'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 24: XII- 10). See Fig. 2A. Kidani, Natasho-mura, Onyu-gun {^%M>^ B9 Ji ^ ^ # ); Fukui, HONSHU; 35°23'N, 135°43'E; collected Jul. 1994 by PRIKU staff; PRIKU, 2 (skeletons only, including 1 with fragmented skull). SW42. Kidooku, Ebara-son, Mima-gun (H/^IP/IJ^W ?KFII); Tokushima, SHIKOKU; 34°08'N, 134°irE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23:XI-31). SeeFig. 2A. Kigiura vicinity, Mikita-cho, Kaifu-gun (^nPlP Hllij^BafflT7KlliJ^)ifte); Tokushima, SHIKOKU; 33°47'N, 134°36'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-6). See Fig. 2 A. Kimitsu-city (^^Tfi); Chiba, HONSHU; ca. 35°19'N, 139°53'E; provisioning 1969-1976 reported by Study Group on the Present Status of Japanese Monkeys (1977b, p. 26). MtDNA samples collected before 1999 by Y. Kawa- moto (1998, p. 54). ^£7/ 9. Kinbaratani, Toyama-mura, Motosu-gun (^^?P ^KUJ^^J^^); Gifu, HONSHU; 35°34'N, 136°39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41:XXV111-11). SeeFig. 2A. Kinkazan (^llOj); Miyagi, KINKAZAN (= FOODEN AND AIMI: SYSTEMATIC REVIEW OE JAPANESE MACAQUES 143 KINKASAN); 38°16'-38°19'N, 141°33'- 141°36'E; birth season, 1966, reported by Kawai et al. (1967, p. 39; cf. Study Group on the Present Status of Japanese Monkeys, 1977a, p. 10). Collected 11 Aug. 1982 by K. Izawa; MUE, 1 (skull only). MtDNA samples col- lected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). Not mapped. Kinkazan vicinity, Ayukavva-mura, Oshika-gun ( tt E IP l£ Jll W ^ H OJ fte ); Mvfl^/, KINKAZAN; 38°18'N, 141°33'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLIlI-12). See Fig. 2A. Kinkazan, Loc. 1 ( ^ H ill ); Miyagi. KINKAZAN; 38°17.5'N, 141°35.5'E; collected 3 Dec. 1982 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 2 ( :^ H |JL| ); Miyagi. KINKAZAN; 38°18.25'N, 141°35'E; collected 18 Jul. 1983 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 6 ( :^ ¥ ill ); Miyagi, KINKAZAN; 38°17.5'N, 141°34'E; collected 24 Feb. 1984 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 10 ( ^ H |JL| ); Miyagi. KINKAZAN; 38°17.25'N, 141°34'E; collected 31 Mar. 1984 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 11 ( :^ H ill ); Miyagi. KINKAZAN; 38°18.25'N, 141°35'E; collected 7 Apr. 1984 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 12 ( :^ ¥ ill ); Miyagi. KINKAZAN; 38°17.25'N, 141°34'E; collected 14 Apr. 1984 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 13 ( ^ H ill ); Miyagi. KINKAZAN; 38°18'N, 141°34'E; collected 23 Apr. 1984 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 14 ( :^ ¥ ill ); Miyagi, KINKAZAN; 38°18'N, 141°34'E; collected 29 Apr. 1984 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 15 ( :^ ^ ill ); Miyagi. KINKAZAN; 38°17'N, 141°35'E; collected 17 May 1984 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 20 ( :#: H ill ); Miyagi, KINKAZAN; 38°17.5'N, 141°35'E; collected 20 Jan. 1985 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 24 ( ^ H |lj ); Miyagi, KINKAZAN; 38°16.5'N, 141°35.5'E; collected 15 May 1985 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 28 ( ^ H ill ); Miyagi, KINKAZAN; 38°17.25'N, 141°34'E; collected 12 Apr. 1986 by K. Izawa; MUE, 1 (skull only). NE22. Kinkazan, Loc. 33 ( :^ ? ill ); Miyagi, KINKAZAN; 38°17'N, 141°35.5'E; collected 3 Oct. 1988 by K. Izawa; MUE, I (skull only). NE22. Kinkazan, Loc. Y6 ( :^ ? ill ); Miyagi, KINKAZAN; 38°17.25'N, 141°35'E; collected 26 May 1984 by K. Izawa; MUE, 1 (skull only). hE22. Kinpusan vicinity, Miyamoto-mura, Naka- koma-gun {^^W.^'M^^±^\^^); Ya- manashi, HONSHU; 35°51'N, 138°37'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-12). SeeFig. 2A. Kinsenji. See Kyogatake. Kintokiyama, Ashigara-mura, Sunto-gun (f^M ?P £ ffi ^ ^ B#'UJ ); Shizuoka, HONSHU; 35°19'N, 139°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX- 15). See Fig. 2A. Kintokiyama vicinity, Sengokuhara-mura, Ashi- garashimo-gun (£ffiT#fllj^I^W^Btlljflfe); Kanagawa, HONSHU; 35°17'N, 139°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXV-4). See Fig. 2A. Kirakuyama vicinity, Kiyomizu-mura, Agawa-gun (#JlliP^7K;^^Maj^); Kochi, SHIKOKU; 33°44N, 133°22'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-26). See Fig. 2A. Kirigatana, Touni-mura, Kesen-gun (^"(llj^^ :^^^ fl^ -Jr il ); luate, HONSHU; 39°12'N, 141°5rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, 144 FIELDIANA: ZOOLOGY p. 56: XLV-6). See Fig. 2A. Kirigataniyama vicinity, Kawagoe-mura, Oochi-gun {G^m}\m^m'T'^aii\k): Shi- mane, HONSHU; 34°57'N, 132°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI- 17). See Fig. 2A. Kirishimasanrin, Higashisonoyama-mura, Aira- gun (Itp^^P^ftOJ^^tSftUj^); Kagoshima, KYUSHU; 31°50'N, 130°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: I- 10). See Fig. 2A. Kirishimayama, Nishidake-son, Kitamoro- kata-gun (itMW^^^^^MMiU); Miyazaki, KYUSHU; 31°52'N, 130°58'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: II-8). See Fig. 2 A. Kirishimayama vicinity, Takahara-son, Ni- shimorokata-gun {'^WW.^mW-^^BMiUifky, Miyazaki, KYUSHU; 3r56'N, 131°04'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-12). See Fig. 2A. Kishida, Hatta-mura, Mikata-gun (M^^fPABB ^^f # ffl ); Hyogo, HONSHU; 35°31'N, 134°26'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 30:XVIII-11). SeeFig. 2A. Kiso ( tK ^ ); Nagano, HONSHU; 35°45'N, 137°43'E; field studies 1970-1975 reported by F. Fukuda (1977, p. 76). Collected 27 Apr. 1982 by N. Agetsuma; PRIKU, 1 (skeleton only). Collected 24 May 1982 by PRIKU staff; PRIKU, 1 (skeleton only). NE80. Kisuki-cho, Ohara-gun ( Ai^lP7K;^fflI); Shi- mane, HONSHU; ca. 35°17'N, 132°54'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW59. Kitakomachi, Kamogawa-city (fIJII rf]db''J''fflI); Chiba, HONSHU; 35°07'N, 140°03'E; col- lected 7 and 8 Feb. 1998 by PRIKU staff; PRIKU, 2 (skeletons only). NE125. Kitamatasanrin, Nosegawa-mura, Yoshino-gun (^i?IPi?MJIIWdbflS:ajW); Nara, HONSHU; 34°07'N, 135°39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXI- 1). See Fig. 2 A. Kitamine, Kinsha-mura, Uma-gun {^MW>^^ ^ it ^ ); Ehime, SHIKOKU; 33°54'N, 133°34'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18:IX-11). SeeFig. 2A. Kitamura, lijima-machi, Kamiina-gun (_b#BP?P tSSfflJdb^); Nagano, HONSHU; 35°40'N, 137°54'E; collected 31 Oct. and 2 Dec. 1997 by M. Aimi; PRIKU, 1 1 (skeletons only, in- cluding 4 with fragmented skulls and 1 with postcranials only). NE77. Kitanada-son, Itano-gun (^if^Pdbil^^); Toku- shima, SHIKOKU; 34°12'N, 134°31'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 24: XI-34). See Fig. 2A. Kitanokawa vicinity, Koguchi-mura, Higashi- muro-gun (^^ftlPd^ P^dby JHftfe); Wa- kayama, HONSHU; 33°45'N, 135°52'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-48). See Fig. 2A. Kitayama, Shioze-mura, Arima-gun (W/^?P^ )i ^ Jb UJ ); Hyogo, HONSHU; 34°48'N, 135°18'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-29). See Fig. 2A. Kitousanrin, Kamikitou-son, Kaifu-gun (^qP?P ±7KM^^7K1IUj^); Tokushima, SHIKOKU; 33°46'N, 134°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: Xl-4). See Fig. 2A. Kitousanrin, Okukitou-son, Kaifij-gun (^nP?P ||7KlI;^7KIIli|#); Tokushima, SHIKOKU; 33°45'N, 134°10'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-3). See Fig. 2A. Kiwadahata, Kimitsu-city (^/$ Tti H^ BB !lffl); Chiba, HONSHU; 35°11'N, 140°0rE; col- lected Jul. 1996-Dec. 1998 by PRIKU staff; PRIKU, 6 (skeletons only, including 1 with fragmented skull). NE120. Kiyano vicinity, Ookawa-mura, Tosa-gun (zhfe IPAJIl^TKMifflfe); Kochi, SHIKOKU; 33°48'N, 133°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-29). See Fig. 2A. Kiyokawa-mura, Aikou-gun ( S ¥ ?P /ff J 1 1 W ); Kanagawa, HONSHU; ca. 35°28'N, 139°17'E; reported present before 1978 by Study Group FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 145 on the Present Status of Japanese Monkeys (1977b, p. 23). MtDNA samples collected be- fore 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NEWS. Kiyosumiyama (/^Mlll) (= Motokiyosumiyama; Mt. Kiyosumi); Chiba, HONSHU; ca. 35°08'N, 140°12'E; reported before 1965 by K. Ka- wanaka (1973, p. 115). Troop distribution mapped by A. Suzuki (1972, p. 334). NE125. Ko. See Kojiba. Kobe-city (ttpTtT); Hyogo, HONSHU; 34°41'N, 135°10'E, reported origin of monkey obtained 2 Aug. 1904 by National Zoological Park, Washington, D.C.; USNM (specimen received 12 Jun. 1912), 1. Not mapped. Kochi-cho, Kamo-gun ( M j^ ?P M 1*1 fflj ); Hi- roshima, HONSHU; 34°28'N, 132°53'E; pro- visoned group; birth season, 1958-1959, re- ported by Kawai et al. (1967, pp. 37, 39). Blood samples collected before 1992 by No- zawa et al. (1991, p. 414; 1996, p. 6). SW63. Koganezawa vicinity, Nanaho-mura, Kita- tsuru-gun (db|[PlgiP-b^^d^^)Rftfe); Ya- manashi, HONSHU; 35°41'N, 138°57'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-25). See Fig. 2A. Kogarashigataniyama vicinity, Sawatani-mura, Oochi-gun (E^?P)i#W7Ktt#ajffe); Shi- mane, HONSHU; 35°05'N, 132°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-26). See Fig. 2A. Kohoku-cho, Higashiazai-gun (^/^#?P/Sdb BU); Shiga, HONSHU; ca. 35°26'N, 136°15'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW6. Koide, Kuramata-mura, Nakauonuma-gun (4^^ >S iP 5t ^ W ^J^ lii ); Niigata, HONSHU; 36°58'N, 138°44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXlI-2). See Fig. 2A. Kojiba, Kamiyaku-cho, Kumage-gun (^^?P_h WX^HM^y, Kagoshima, YAKUSHIMA; ca. 30°22'N, 130°23'E; observed 1975-1979 by T. Maruhashi (1982, p. 318; cf. Takasaki & Masui, 1984, p. 311). SfF/^y. Kojima, Ichiki-son, Minaminaka-gun (j^SPlRliP TfT^KW^fi); Miyazaki, KOJIMA; 31°27'N, 131°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15:11-3). See Fig. 2A. Kojima, Kushima-city ($f^r(i^A) (= Ko- shima); Miyazaki, KOJIMA; 3r27'N, 131°23'E; reported present before 1939 by N. Kuroda (1938, p. 112; 1940, p. 271). Provi- sioning history, 1952-1977, reported by A. Mori (1979b, p. 372). Birth season, 1956-1966, reported by Kawai et al. (1967, p. 39). Col- lected 11 Sep. 1969-29 Sep. 1996 by KFL staff; KFL, 48 ( 1 skin only; 47 skeletons only, including 3 with fragmented skulls, 8 with missing mandibles, and 1 with missing cra- nium). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). Mi- crosatellite DNA analyzed before 1996 by Domingo-Roura et al. (1997, p. 358). External measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW97. Koka-cho, Koka-gun (¥M^¥MfflI); Shiga, HONSHU; ca. 34°54'N, 136°14'E; mtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). 5^/7. Kokurano, Ootaki-machi, Isumi-gun (MPfI^A ^^fflJ/JN^tif); Chiba, HONSHU; 35°12'N, 140°09'E; collected 10 Aug. and 8 Sep. 1997 by PRIKU staff; PRIKU, 3 (skeletons only). NE126. Komagane Mountain Range, Hiyoshi-mura, Ni- shichikuma-gun (M^^iPSM^ffil'^-^fgUj M); Nagano, HONSHU; 35°51'N, 137°47'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-10). SeeFig. 2A. Komagatake, Kanegasaki-mura, Isawa-gun (ll/R ^^T^^^T^); hvate, HONSHU; 39°irN, 141°06'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLV-7). See Fig. 2A. Komagatake Mountain Range, Agematsu-machi, Nishichikuma-gun (ffimSIP±^A^fflJ|fi|J\W.nP^^m^\^); Nagano, HONSHU; 35°15'N, 137°53'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXI-51). SeeFig. 2A. Kumadayama, Nakamata-mura, Iwafline-gun (^ ^p' ^ 4^ ^M ^ ^ ea UJ ); Niigata, HONSHU; 38°3rN, 139°36'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXII-26). See Fig. 2A. Kumamidani vicinity, Kuroda-mura, Kitaku- wada-gun (db^ffllPHBa^l^m^ffe); Kyoto, HONSHU; 35°13'N, 135°44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXII- 19). See Fig. 2A. Kumano-city {MWt'^); Mie, HONSHU; ca. 33°53'N, 136°07'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW25. Kumayama-cho, Akaiwa-gun (55^^?Pfl^li| fff); Okayama, HONSHU; 34°47'N, 134°05E; 148 FIELDIANA: ZOOLOGY mtDNA samples collected 1998-2000 by I. Yoshimi and H. Takasaki (2003, p. 71). SW52. Kunimi vicinity, Higashinakasuji-mura, Hata-gun i^^mM^M^mM.i\k); Kochi, SHIKOKU; 32°57'N, 132°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: X-5). See Fig. 2A. Kunimidaira vicinity, Uchinoura-mura, Ki- motsuki-gun (flf M ^ 1*1 ;^M^ @ .^^ftfe); Kagoshima, KYUSHU; 31°15'N, 131°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-16). See Fig. 2A. Kuniwaridake (= Mt. Kuniwari), Kamiyaku-cho, Kumage-gun (^.^^±M^fflT@SlJS); Ka- goshima, YAKUSHIMA; 30°20'N, 130°25'E; collected 27 Jul. 1981 by PRIKU staff; PRIKU, 1 (skeleton only, with fragmented skull). SWIOL Kuniwaridake (= Mt. Kuniwari), southeastern slopes, Kamiyaku-cho, Kumage-gun (SI^^P ±M^ffliaflJ^I^K#4®), ca. 900-1200 m elevation; Kagoshima, YAKUSHIMA; ca. 30°20'N, 130°26'E; ca. 11 troops censused Jul.-Aug. 1995 by S. Yoshihiro et al. (1999, p. 4\\).SWW1. Kuniwaridake (= Mt. Kuniwari), summit area, Kamiyaku-cho, Kumage-gun (^l^^-hM^ fflJSitJS]I±), ca. 1200-1323 m elevation; Kagoshima, YAKUSHIMA; ca. 30°21'N, 130°25'E; 4 troops censused Jul.-Aug. 1995 by S. Yoshihiro et al. (1999, p. 41 1). SWIOI. Kuniwaridake (= Mt. Kuniwari), western slopes, Kamiyaku-cho, Kumage-gun (^^^-hM^ ffllSilJSffif^®), ca. 300-900 m elevation; Kagoshima, YAKUSHIMA; ca. 30°20'N, 130°24'E; ca. 13 troops censused Jul.-Aug. 1995 by S. Yoshihiro et al. (1999, p. 411). DNA samples collected before 2005 by X. Domingo-Roura et al. (2004, p. 33). Not mapped. Kuniyama National Forest, Iwanashi-son, Akaiwa-gun ('^^^S^WSUjSW^); Okayama, HONSHU; 34°50'N, 134°09'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV-20). See Fig. 2A. Kuradama, Kimitsu-city (^/^rtTMi); Chiba, HONSHU; 35°13'N, 140°07'E; collected 2 Jul. 1996-5 Mar. 1999 by PRIKU staff; PRIKU, 31 (skeletons only, including 3 with frag- mented skulls). NE126. Kuragariyama, Kamiyama-mura, Uma-gun (^S W±lh^'y'7:^U[Uy, Ehime, SHIKOKU; 33°58'N, 133°4rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: IX- 13). See Fig. 2A. Kuraishidake, Miya-mura, Katsuta-gun (XlJBQ^P ^^^^11); Miyagi, HONSHU; 38°0rN, 140°37'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55:XLIII-4). SeeFig. 2A. Kurase vicinity, Sakuragi-mura, Shusou-gun (M mW^^^^Mi\^); Ehime, SHIKOKU; 33°49'N, 133°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18: IX-7). See Fig. 2A. Kuratani vicinity, Kitagouchi-son, Kuga-gun (^ iRl ?P db )rI rt W ^ ^ ffe ); Yamaguchi, HONSHU; 34°09'N, 132°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII- 13). See Fig. 2A. Kuratani vicinity, Saigawa-mura, Ishikawa-gun (^JIIiPJlJIW5t#ffe); Ishikawa, HONSHU; 36°28'N, 136°42'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXVI- 1). See Fig. 2A. Kurataniyama, Tataragi, Nakagawa-mura, Asago-gun im^W>^)\m^^ ^^^t'^lh); Hyogo, HONSHU; 35°15'N, 134°49'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-15). SeeFig. 2A. Kurio-Segire. See Yakushima, Census Area 6. Kurisuyama, Joto-mura, Niwa-gun (^^?Pi^K ^ ^ ffi Uj ); Aichi, HONSHU; 35°21'N, 137°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVII-9). See Fig. 2A. Kuriyama-mura, Shioya-gun (^#^^li|^); Tochigi, HONSHU; ca. 36°54'N, 139°35'E; collected 2 Aug. 1972 and 18 Sep. 1986 by TPM staff; TPM, 2 (skulls only, including 1 with mandible missing). NE42. Kurobe (H^); Toyama, HONSHU; 36°39'N, 137°41'E; observed Nov. 1979-Jan. 1987 by H. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 149 Akaza(1988,p. \2).NE54. Kurobedani,Uchiyama-mura, Shimoniikawa-aun ( T Sf Jll ?P rt OJ ^ H ^ ^ ); Toyama, HONSHU; 36°51'N, 137°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 42: XXIX-9). See Fig. 2A. Kuroda vicinity, Takane-son, Kuga-gun (3Eft.iRllP R S ^ H ea -te ); YamaguchC HONSHU; 34°24'N, 132°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII- 14). See Fig. 2 A. Kurohimeyama, Takayanagi-mura, Kariwa-gun (XlJ^?PS^^IIieaj);^M/ga/fl, HONSHU; 37°11'N, 138°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII- 13). See Fig. 2A. Kurohimeyama, Ukawa-mura, Kariwa-gun (X'J^ ^ 56IJII ^ II ie UJ ); Niigata, HONSHU: 37°13'N, I38°34'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII- 1 1). See Fig. 2 A. Kurokagara vicinity, Asuna-mura, Oochi-gun (e ^W>n^M^W.ti1i^ ^)\ Shimam. HONSHU; 34°53'N, 132°37'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-20). See Fig. 2A. Kurokawa, Hirata-mura, Hata-gun (IS^IP^BQ ^ H Jll ); Kochi, SHIKOKU; 32°56'N. 132°49'E; reported in questionnaire sur\ey conducted in 1 923 by K. Hasebe (Iwano, 1 974. p. 21:X-4). SeeFig. 2A. Kurokawa-mura, Kitakanbara-gun (dbSJ^lPM Jll ^ ); Niigata, HONSHU; ca. 38°04'N. I39°26'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002. p. 60). NE33. Kurokawayama, Awano-mura, Tsuruga-gun (^ MIPHS^H^rTOJ); FukuL HONSHU; 35°34'N, 136°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV- 16). See Fig. 2 A. Kuromoriyama vicinity. Nishiokuni-mura, Mogami-gun ( S ± IP ® ^M ^hTH ^ Oj ffe ); Yamagata, HONSHU; 38°48'N, 140°29'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLII-10). See Fig. 2 A. Kurosawa National Forest. Ogura-mura, Mi- namiazumi-gun (]^$^iP'J^li"^M/RB W ^); Nagano, HONSHU; 36°20'N, 137°50'E; reported in questionnaire sur\'ey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-16). SeeFig. 2A. Kurosu vicinity. Shimazu-mura, Watarai-gun (]^ ^^Pft^^H^-te); Mie, HONSHU; 34°15'N, 136°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37:XXIII-31). SeeFig. 2A. Kurotaki. Yamauchi-mura, Koka-gun (^MlPill 1*1 ^ II )i ); Shiga, HONSHU; 34°55'N, 136°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV-26). See Fig. 2A. Kurouchi, Kiso-mura, Kiso-gun (^e^^ffi^ Urt); Nagano, HONSHu'J 35°58'N. 137°45'E; collected 12 Dec. 1997 by M. Aimi; PRIKU, I (skeleton only). NE86. Kusaki-mura, Sayo-gun (■feffl^P^i'^W); Hyogo, HONSHU; 34°56'N. 134°18'E; reported in questionnaire sur\'ey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-3). See Fig. 2A. Kusenbuyama vicinity. Minamihata-mura. Chi- kushi-gun {m^^^m^h=f^\h^): Fu- kuoka, KYUSHU; 33°24'N, 130°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 17: VII- 1). See Fig. 2A. Kushigawa. Kaifu-cho, Kaifu-gun (^pPlP^qP urr^iJII); Tokushima, SHIKOKU; 33°36'N, 134°17'E; collected 19 Sep. 2001 by M. Aimi; PRIKU, \.SW73. Kushigawa vicinity. Kawanishi-son, Kaifu-gun ( m^ ^ Jll S ^ # Jll ^ ); Tokushima, SHIKOKU; 33°35'N. 134°17'E; reported in questionnaire sur\ey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI- 1). See Fig. 2A. Kushima-city (^ralrfJ); Miyazaki. KYUSHU; ca. 3^287^, 131°12'E; blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW96. Kusoudomari, Wakinosawa-mura. Shi- mokita-gun {Ti^^^WR^XB.^); Ao- mori, HONSHU; 41°09'N, 140°47'E; found 150 FIELDIANA: ZOOLOGY and donated in 1964 by unknown collector (cf. Study Group on the Present Status of Japanese Monkeys, 1977a, p. 10, locality Aomori-2); JMC, 1 (skull only). NE5. Kusugawa, Kamiyaku-cho, Kumage-gun (^^ IP ± Jl ^ ffll ffi Jll ); Kagoshima, YAKUSHIMA; 30°23'N, 130°35'E; collected 25 Jul. 1987 by PRIKU staff; PRIKU, 1 (skeleton only). SWIOO. Kuwashimachinai, Shiramine-mura, Nomi-gun ( tg H ?P a III* ^ # ft itfe I*] ); Ishikawa, HONSHU; 36°10'N, 136°39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVI-6). See Fig. 2A. Kuwasononanbu, Matsukawa-machi, Shi- moina-gun (T^IPlP^AMIIffll^Ult^); Na- gano, HONSHU; 35°36'N, 137°52'E; collected 11 Dec. 1997 by M. Aimi; PRIKU, 4 (skele- tons only). NE76. Kyogatake ( i^ "^ S ); Nagasaki-Saga border, KYUSHU; ca. 32°59'N, 130°04'E; observed in 1940s by local residents (Sakura, 1976, p. 151); subsequently extinct at this locality (Ikeda and Eguchi, 1978, p. 59). 5^(97. Kyonan-machi, Awa-gun (^J^^PISl^fflJ); Chiba, HONSHU; ca. 35°06'N, 139°52'E; mtDNA samples collected before 1999 by Y. Kawamoto (1998, p. 54; 2002, p. 60). NEI20. Kyouhou vicinity, Hajikami-mura, Hata-gun (♦# ^?P^±^^t:^)iffe); Kochi, SHIKOKU; 33°00'N, 132°43'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: X-1 1). See Fig. 2 A. Launceston, Tasmania; 4r35'S, 147°22'E; translocation in 1980 from Kasuga, Hyogo Prefecture, reported by M. Nozaki (1993, p. j97). Not mapped. Maemoriyama vicinity, Higashiokuni-mura, Mogami-gun (ft ±1^^^^ S ^lu I^Ul ffe); Yamagata, HONSHU; 38°46'N, 140°33'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLIl-11). SeeFig. 2A. ^ Maki, Takayama-mura, Kamitakai-gun (Jiisi^ ^PilUj;^!"^); Nagano, HONSHU; 36°40'N, 138°25'E; collected 19 Jul. 1998 by M. Aimi; PRIKU, 1 (skeleton only). NE49. Makino-cho, Takashima-gun (i^ S?P'7 ^ y BJ), Shiga, HONSHU; 35°30'N, 136°03'E; ob- served Aug.-Sept. 1989 by M. Nakamichi et al. (Yamagiwa & Hill, 1998, p. 267). SW3. Makinosaki vicinity, Oohara-mura, Oshika-gun (ttElPAJ^^^(7)llif ffe); Miyagi, HONSHU; 38°2rN, 141°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLIII-1 1). See Fig. 2A. Makiosan vicinity, Makinoshima-mura, Kuze-gun (r^^iP^ft^^mUjftfe); Kyoto, HONSHU; 34°53'N, 135°46'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXII-28). See Fig. 2A. Makitaniyama vicinity, Umaji-mura, Aki-gun (^ ^^m^^W^\h^); Kochi, SHIKOKU; 33°32'N, 134°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-39). See Fig. 2A. Mantaroyama vicinity, Tsuchidaru-mura, Mi- namiuonuma-gun (]^^5S?P±#^ll:^l!Plil fte); Niigata, HONSHU; 36°53'N, 138°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII-4). See Fig. 2A. Manzayama, Tsumagoi-mura, Agatsuma-gun (^ ^ IP If i2^x ^ 77 0 111 ); Gunma, HONSHU; 36°38'N, 138°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XL-8). See Fig. 2 A. Maruseyama, Ichigi-mura, Oochi-gun (e^^Tfi" :^¥iSlM\^); Shimane, HONSHU; 34°51'N, 132°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI- 16). SeeFig. 2A. Maruyama, Ubuchi-mura, Usui-gun (fltt^XlP/^ ^J ^ ^ UJ ); Gunma, HONSHU; 36°25'N, 138°46'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51:XL-6). SeeFig. 2A. Masakiyama, Ipponmatsu-mura, Minamiuwa-gun (jt^^^P-^^i^IETKUj); Ehime, SHIKOKU; 32°59'N, 132°40'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18: IX-4). See Fig. 2A. Masakiyama, Tsutsuga-son, Yamagata-gun (|JL| !f;15MM^IE7KUj); Hiroshima, HONSHU; 34°35'N, 132°15'E; reported in questionnaire FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 151 survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-4). See Fig. 2A. Masezawa, Hachimori-mura, Yamamoto-gun (llj ^W>J\M^MM'yR); Akita, HONSHU; 40°22'N, 140°07'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLIV-9). See Fig. 2A. Masudayama, Nakatane-mura, Kumage-gun {M. ^W^U^^i^miU); Kagoshima, TANEGASHIMA; 30°35'N, 130°59'E; re- ported as extinct in questionnaire survey con- ducted in 1923 by K. Hasebe (Iwano, 1974, p. 12:1-3). See Fig. 2A. Masugata, Takayama-mura, Kamitakai-gun (Ji mk^mih^mM); Nagano, HONSHU; 36°4rN, 138°22'E; collected 7 Oct. and 1 Nov. 1998 by M. Aimi; PRIKU, 3 (skeletons only). NE49. Matani, Agira-mura, Kawage-gun (5nJS^Q^:|4 M-^y, Mie, HONSHU; coordinates unknown; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIII-58). Not mapped. Matsubakawayama vicinity, Matsubakawa-mura, Takaoka-gun (I^I?^?P^A^SJI|^^iSJI| UUffe); Kochi, SHIKOKU; 33°16'N, 133°04'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-19). See Fig. 2A. Matsukasa-mura, lishi-gun ( tS S ?P ^ S ^ ); Shimane, HONSHU; 35°11'N, 132°42'E; re- ported (as possibly temporary habitat) in ques- tionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-33). See Fig. 2A. Matsuno-cho, Kitauwa-gun (db^^?P^ii?fflI); Ehime, SHIKOKU; ca. 33°12'N, 132°44'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW78. Mayasan, Nishinada-mura, Muko-gun (K0^S li;^^t0gP^J); Hyogo, HONSHU; 34°44'N, 135°12'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-30). See Fig. 2A. Mera, Tateyama-city ( tl ill TtT ^ ^ ); Chiba, HONSHU; 34°54'N, 139°49'E; collected 12 Jul. 1997 by PRIKU staff; PRIKU, 1 (skeleton only). NE122. Metaka, Ooshika-mura, Shimoina-gun (T#3P ^XU¥it:m)\ Nagano, HONSHU; 35°38'N, 138°03'E; collected 31 Dec. 1997 by M. Aimi; PRIKU, 1 (skeleton only). NE70. Michikata vicinity, Nakajima-mura, Watarai-gun (Jt^^4^ft^]l:^'te); Mie, HONSHU; 34°18'N, 136°35'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-37). See Fig. 2A. Mie ( H M ) (= Miye), HONSHU; 33°44'-35°15'N, 135°52'-136°54'E; collected. 4 Jan. 1939 by T. Barbour; MCZ, 1. Not mapped. Mihama-cho, Mikata-gun {=.^5^^)^^); Fu- kiti, HONSHU; ca. 35°36'N, 135°56'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW2. Mihama-cho, Minamimuro-gun (l^-^ft^P^^ fflj); Mie, HONSHU; ca. 33°48'N, 136°02'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW26. Mihama-mura, Kamo-gun (Mj^^PH^^); Shi- zuoka, HONSHU; 34°43'N, 138°46'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX-24). See Fig. 2A. Mihara-city (Hj^TtT); Hiroshima, HONSHU; 34°24'N, 133°05'E; external measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). SW64. Mikagura, Nishikawa-mura, Higashikanbara-gun (^aj^?PMJI|^1itt2$<); Niigata, HONSHU; 37°37'N, 139°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII-16). See Fig. 2A. Mikata-cho, Mikata-gun (H;?31PH;J3 fflJ); Fukui, HONSHU; 35°37'N, 135°51'E; captured 1 May 1968 by JMC staff, died in captivity 24 May-5 June 1968; JMC, 3 (skulls only). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6).5Pr7. Mikatasanrin, Ya-mura, Mikata-gun (H^?PA ;^^t H :i^ Oj :|9^ ); Fukui, HONSHU; 35°33'N, 135°55'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39:XXV-13). SeeFig. 2A. Mikoshi vicinity, Misato-mura, Higashimuro-gun (K^«?PHM^H^^); Wakayama, HONSHU; 33°51'N, 135°43'E; reported in questionnaire survey conducted in 1923 by K. 152 FIELDIANA: ZOOLOGY Hasebe (Iwano, 1974, p. 34: XX-52). See Fig. 2A. Mikuniyama, Takahashi-mura, Izushi-gun (t±i5 iPilM^HSUj); Hyogo, HONSHU; 35°26'N, 135°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-20). See Fig. 2 A. Mikusu, Minakami-cho, Ooda-city (AB9 rfT7K_L ffllH:^^); Shimane, HONSHU; 35°04'N, 132°26'E; collected 15 Aug. 1994 by PRIKU staff; PRIKU, 4 (skeletons only). SW6L Mimaisan vicinity, Mimai-mura, Nishimuro-gun (ffi^ftfPH^^^H^Ojffe); Wakayama, HONSHU; 33°36'N, 135°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-29). See Fig. 2A. Minamikawa vicinity, Mori-mura, Tosa-gun (i te ^ 1^ ^ it Jll ^ ); Kochi, SHIKOKU; 33°45'N, 133°33'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-30). See Fig. 2A. Minamikomagadake. Nanakubo-mura, Kami- ina-gun {^L^M^^zXU^'^^'r ^); Na- gano, HONSHU; 35°39'N, 137°53'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-43). See Fig. 2A. Minamikurosawa National Forest, Azusa-mura, Minamiazumi-gun (it$S?P#^^]tM/H@ W^); Nagano, HONSHU; 36°14'N, 137°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-13). SeeFig. 2A. Minamimachi, Suwa-city (Ml^rfTj^fflJ); Nagano, HONSHU; 36°00'N, 138°08'E; collected 26 Jul. 1998 by M. Aimi; PRIKU, 1 (skeleton only). NE95. Minamitoyosawayama vicinity, Yukuchi-mura, Hienuki-gun (Pml^^^P^lt^/RLljffe); Iwate, HONSHU; 39°25'N, 141°06'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLV-9). See Fig. 2A. Minoo-city ( ^ ® rf] ) (= Minou); Osaka, HONSHU; ca. 34°51'N, 135°29'E; provisioned group; birth season, 1957-1966, reported by Kawai et al. (1967, pp. 37, 39). Collected 10 Apr. 1979 and 19 Jun. 1980 by PRIKU staff; PRIKU, 3 (skeletons only). MtDNA samples collected ca. 1986-1991 by Hayasaka et al. (1991, p. 400). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW53. Minoosan National Forest, Minoo-mura, Toyono-gun (MtelP^®^^® ill HW^); Osaka, HONSHU; 34°49'N, 135°30'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XIX-1). SeeFig. 2A. Miokawa-mura, Higashimuro-gun (!^-^SlPH MJII^); Wakayama, HONSHU; 33°34'N, 135°4rE; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 34: XX-39). See Fig. 2A. Misaki-Makinouchi vicinity, Toi-son, Minami- naka-gun {"^MM^n^^imm^cn]^^)- Miyazaki, KYUSHU; 31°23'N, 13r20'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: II- 1). See Fig. 2A. Misogawa-goryorin vicinity, Kiso-mura, Ni- shichikuma-gun (SmSlPTKia^D^PgJIIiai ^ W ftfe ); Nagano, HONSHU; 35°59'N, 137°46'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46:XXXI-11). SeeFig. 2A. Misokawa, Kiso-mura, Kiso-gun (^a^^ffi^t ^ Pi Jll ); Nagano, HONSHU; 35°58'N, 137°46'E; collected 30 Dec. 1997 by PRIKU staff; PRIKU, 2 (skeletons only). NE86. Mitakeyama, Aikawa-mura, Oono-gun (Ai??P -^JHWISli^Lil); Oita, KYUSHU; 32°54'N, 131°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16:111-5). SeeFig. 2A. Mitakeyama-goryorin, Mitake-mura, Nishichi- kuma-gun (ffimJSiPH|&;14|EPi^aj|i^#); Nagano, HONSHU; 35°53'N, 137°36'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXXI-9). See Fig. 2A. Mitoya-cho, lishi-gun (ISSlPHTJMffll); Shi- mane, HONSHU; 35°17'N, 132°52'E; col- lected 7 Feb. 1995-13 Mar. 1997 by PRIKU staff; PRIKU, 5 (skeletons only, including 1 with fragmented skull; excluding 1 juvenile FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 153 skull mistakenly associated with data pertain- ing to large adult male). SW59. Mitsudake, Hata-mura, Taki-gun (^iS^^^H it); Hyogo, HONSHU; 35°07'N, 135°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-27). See Fig. 2A. Mitsue-mura, Uda-gun (^PSIP^^^); Nara, HONSHU; 34°30'N, 136°11'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXI-8). See Fig. 2A. Mitsuminegawa upstream, Inasato-mura, Kami- ina-gun (±^3PIP#SPM^H** Jl| ±)^); Nagano, HONSHU; 35°41'N, 138°07'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-44). See Fig. 2A. Mitsutouge, Nishikatsura-mura, Mina- mitsuru-gun (^tPlilPM#^H7 llf^); Ya- manashi, HONSHU; 35°31'N, 138°49'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXIII-30). See Fig. 2A. Miuchisanrin, Ishii-mura, Sayo-gun (fcffilP^ ^:|4^/il*mj^); Hyogo, HONSHU; 35°05'N, 134°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-4). See Fig. 2A. Miurayama vicinity, Outaki-mura, Nishichi- kuma-gun O^S^PiJt^ftHMUjffe); Na- gano, HONSHU; 35°48'N, 137°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXXI-8). See Fig. 2A. Miyagi CM^), HONSHU; 37°47'-39°00'N, 140°16'-141°4rE; studied 1982-2002 at various localities by Miyagi Monkey Investi- gation Group (2003, p. 37). See Fig. 2B. Miyajima-cho, Saeki-gun (fe-fSlP^SBlT); Hi- roshima, MIYAJIMA; 34°16'N, 132°20'E; population translocated from Shodoshima, Kagawa Prefecture, in 1962; provisioned on Miyajima (Iwamoto & Hirai, 1970, p. 395; Shidei et al., 1981, p. 18). Birth season, 1962-1966, reported by Kawai et al. (1967, p. 38). Collected before 2002 by PRIKU staff; PRJKU, 1 (skeleton only, with fragmented skull). SW65. Miyama National Forest, Ookawachi-mura, li- nan-gun (ISl^?PAyRlrt^;^UjSWW); Mie, HONSHU; 34°31'N, 136°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-25). See Fig. 2A. Miyamoto, Takamori-machi, Shimoina-gun (T PMW>mMViJ'B^y, Nagano, HONSHU; 35°34'N, 137°51'E; collected 6 Apr. and 14 May 1998 by M. Aimi; PRIKU, 2 (skeletons only). NE76. Miyamura, Takayama-mura, Kamitakai-gun (_h it^lPSlil^EW); Nagano, HONSHU; 36°41'N, 138°24'E; collected 7 Nov. 1997-2 Nov. 1998 by M. Aimi; 4, (skeletons only). Collected 19 Oct. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE49. Miyanoura, Kamiyaku-cho, Kumage-gun (^^ W ± M X Vil 'B (D M ); Kagoshima YAKUSHIMA 30°25'N, 130°34'E; collected 3 Dec. 1977 by PRIKU staff; PRIKU, 1 (skele- ton only). SWIOO. Miyanoura-Ambo. See Yakushima, Census Area J. Miyashiro, Hodaka-machi, Minamiazumi-gun (^$*lP^SfflT^^); Nagano, HONSHU; 36°23'N, 137°49'E; collected 24 Aug. 1998 by M. Aimi; PRIKU, 6 (skeletons only, including 2 with fragmented skulls). NE92. Miyashita, Matsukawa-mura, Kitaazumi-gun (db ^•iP^AMIIW^T); Nagano, HONSHU; 35°34'N, 137°51'E; collected 22 Feb. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE76. Miye. See Mie. Mizuho-cho, Funai-gun (^n^lP^^fflJ); Kyoto, HONSHU; ca. 35°10'N, 135°21'E; captured 26 Mar. 1977 by JMC staff, accidental death in captivity 4 Apr. 1977; JMC, 1 (skull only). SW39. Mizunashiyama vicinity, Toga-mura, Higashito- nami-gun (mfll):^lP^JM^7KMUjflfe); To- yama, HONSHU; 36°17'N, 137°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 42: XXIX-2). See Fig. 2A. Mizunoshitayama, Iseki, Nachi-mura, Higashi- muro-gun (m#«^i9mW#Pl7Ky TUJ); Wakayama, HONSHU; 33°39'N, 135°57'E; reported in questionnaire survey conducted in 154 FIELDIANA: ZOOLOGY 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-45). See Fig. 2A. Mobara-city ( j5S i^ TtT ); Chiba, HONSHU; 35°25'N, 140°18'E; Jomon subfossils (age, 2.3-12 Ka) reported by K. Kawanaka (1973, p. \\5).NE129. Mochihirasan vicinity, Futagawa-mura, Nishi- muro-gun (ffi^ftlP-Jll^^^ajffe); Wa- kayama, HONSHU; 33°52'N, 135°22'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-17). SeeFig.2A. Momobayama, Itoshiromi, Wakasa-cho, Yazu-gun (A^^P^^fflJ.^ 6 B^^yXOj); Tottori, HONSHU; 35°19'N, 134°25'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 29: XVII-2). See Fig. 2A. Momozaki vcinity, Isato-mura, Minamimuro-gun im^mW^m^mm^); Mle, HONSHU; 34°00'N, 136°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-51). See Fig. 2A. Moniwa-mura, Date-gun ('^ji^^j^SW); Fu- kushima, HONSHU; 37°55'N, 140°27'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 52: XLI-10). SeeFig. 2A. Morimatsu vicinity, Kitawauchi-mura, Minami- muro-gun (]^#ft?Pdblfflrt^^^A^fte); Me, HONSHU; 34°00'N, 136°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-48). See Fig. 2A. Motoi vicinity, Higashiiyayama-son, Mima-gun (H^liPmrn^Oj^TU^fte); Tokushima, SHIKOKU; 33°54'N, 133°57'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-22). See Fig. 2A. Motokiyosumiyama. See Kiyosumiyama. Motomura, Minamishinano-mura, Shimoina-gun ( T ^ 3P ?P 1^ if )I ^^ ^ ^ ); Nagano, HONSHU; 35°18'N, 137°55'E; collected 20 Sep. 1998 by M. Aimi; PRIKU, 1 (skeleton only). NE68. _ Motoyama, Muroto-cho, Aki-gun ($S^^P ffll Tt UJ ); Kochi, SHIKOKU; 33°18'N, 134°10'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-44). See Fig. 2A. Mt. Kiyosumi. See Kiyosumiyama. Mt. Kuniwari. See Kuniwaridake; Nina- A. Mt. Takago. See Takagoyama. Mugikomoriyama vicinity, Ayukawa-mura, Ni- shimuro-gun (M#«IPIiJIIW^3^^ajffe); Wakayama, HONSHU; 33°41'N, 135°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-22). See Fig. 2A. Mugio, Yaku-cho, Kumage-gun (^^^MA.fflI U^y, Kagoshima, YAKUSHIMA; 30°16'N, 130°36'E; collected 21 Jul. 1987 by PRIKU staff; 3 (skeletons only). Collected 21-25 Jul. 1987 by PRIKU staff; PRIKU, 7 (postcranials only). SW102. Mukabakiyama, Minamikata-son, Higa- shiusuki-gun {1^B^^ny5Wlffllliajfte); Kochi, SHIKOKU: 33°16'N, 133°16'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-18). See Fig. 2A. Nagaflika vicinity, Handairayama-son, Mima-gun (ll.l?P¥¥UJ^^)^^); Toku- shima, SHIKOKU; 33°58'N, 134°08'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-32; Mito, 1989, p. 25). See Fig. 2A. Nagahashiri vicinity. Hideya-mura, Higashikan- bara-gun (m^i^lP B tli^^^^'^fe); Nii- gata, HONSHU; 37°43'N, 139°32'E; reported in questiormaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXII- 19). See Fig. 2A. Nagakura vicinity, Hase-mura, Kanzaki-gun (tt ^^^"^m-llO^^Y Hyogo, HONSHU; 35°07'N, 134°44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974. p. 30: XVIII- 14). See Fig. 2A. Nagano (MS) (= Shinshiu), HONSHU; 35°12'-37°01'N, 137°13'-138°48'E; collected Sep. 1882 by P. L. Jouy; USNM, 1 (skull only). Questionnaire survey conducted June 1972 by E. Tokita and S. Hara (1977, p. 27). Not mapped. Nagaoi. Egawasaki-mura. Hata-gun (♦S^^PxIl jl|*f^k^); Kochi, SHIKOKU; 33°irN, 132°47'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: X- 13). See Fig. 2 A. Nagasaka-mukaiyama. Takigawa-mura, Naga-gun {4%^M}\\-^^^^\Uy, Mie, HONSHU; 34°34'N, 136°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII- 17). See Fig. 2A. 156 FIELD! AN A: ZOOLOGY Nagata, Kamiyaku-cho, Kumage-gun (^^?P_h MrXfflJ^ffl); Kagoshima, YAKUSHIMA; ca. 30°23'N, 130°25'E; collected 7 Jul. 1977 and 22 Jul. 1987 by PRIKU staff; PRIKU, 2 (1 skeleton without mandible, 1 postcranials only). Collected 20 Jul. 1987 by Mr. Imaegawa; PRIKU, 2 (skeletons only). Collected 25 Jul. 1987 by Mr. Sinmachi; PRIKU, 1 (skeleton only). Collected 25 Feb. 1989 by PRIKU staff; PRIKU, 4 (skeletons only). SWIOI. Nagata-Issou. See Yakushima, Census Area 1 . Nagata-todai, Kamiyaku-cho, Kumage-gun (^ ^?P±Jl:^fflT*Baff^); Kagoshima, YAKUSHIMA; ca. 30°23'N, 130°23'E; ob- served Aug. 1978 by T. Maruhashi (1982, p. 318; cf. Takasaki & Masui, 1984, p. 311). SWlOl. Nagato-okuyama, Fukusawa-mura, Kaminii- kawa-gun (±ifJII^II)K^m^f MUJ); To- yama, HONSHU; 36°25'N, 137°19'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 42: XXIX-3). See Fig. 2A. Nagatokyo vicinity, Ikumo-son, Abu-gun (PrI^ ^ ^m^ ^f^^ - "^ )\ Yamaguchi, HONSHU; 34°23'N, 131°37'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII-8). See Fig. 2A. Nagatokyo vicinity, Kawakami-son, Abu-gun {n^W>\\\:t^^f^^-w)\ Yamaguchi, HONSHU; 34°21'N, 131°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII-7). See Fig. 2A. Nagatokyo vicinity, Shinobu-son, Abu-gun (Pr[ ^W^U^^^cf^m-^y, Yamaguchi, HONSHU; 34°20'N, 131°36'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII-9). See Fig. 2A. Nagatoro. See Houtosan. Nagisoyama, Azuma-mura, Nishichikuma-gun (SmiSlPl^g^^TK^Uj); Nagano, HONSHU; 35°35'N, 137°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXXI- 1). See Fig. 2A. Nagis(wama vicinity, Yomikaki-mura, Nishichi- kuma-gun (S^SiPlTcH^lt^^ajfe); Nagano, HONSHU; 35°37'N, 137°38'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXXI-2). See Fig. 2A. Nahisanrin, Aioi-mura, Gujo-gun (?P_hlP1S^ ^ SP it UJ ^ ); Gifu, HONSHU; 35°42'N, 136°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p.41:XXVIII-23). SeeFig. 2A. Naka, Natashou-mura, Onyu-gun (]ai![?P^B9 Ji^ct'); Fukui, HONSHU; 35°23'N, 135°38'E; collected Aug.-Dec. 1994 by PRIKU staff; PRIKU, 3 (skeletons only). SW42. Nakabusa National Forest, Ariake-mura, Mi- namiazumi-gun (]^$#IPWB^^4'MSW ^); Nagano, HONSHU; 36°21'N, 137°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-17). SeeFig.2A. Nakadachi, Minamishinano-mura, Shimoina-gun ( T # IP ?P ^ € ^M ^ 4^ it ); Nagano, HONSHU; 35°22'N, 137°58'E; collected 9 Sep. 1998 by M. Aimi; PRIKU, 1 (skeleton only). NE69. Nakahama vicinity, Shimizu-mura, Hata-gun (Wl ^?P)t7K;t4 4'^^); Kochi, SHIKOKU; 32°46'N, 132°58'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: X-6). See Fig. 2 A. Nakahora, Ooshika-mura, Shimoina-gun (T#SP W>XU^^M); Nagano, HONSHU; 35°33'N, 138°02'E; collected 12 Apr. 1998 by M. Aimi; PRIKU, 1 (skeleton only). NE70. Nakakimura, Usui-machi, Usui-gun (fift:^?PS #fflI4^^^); Gunma, HONSHU; 36°18'N, 138044'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 51:XL-5). SeeFig. 2A. Nakakodomariyama, Kodomari-mura, Ki- tatsugaru-gun ( db ^ ^ ?P 'J^ )& ^ 4^ ^^ )^ Ol ); Aomori, HONSHU; 41°07'N, 140°20'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-12). SeeFig. 2A. Nakakurayama, Shimoyakawa-mura, Aga- wagun ( f^ Jll iP T A Jll^ 4^^111); Kochi, SHIKOKU; 33°36'N, 133°20'E; reported in questionnaire survey conducted in 1923 by K. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 157 Hasebe (Iwano, 1974, p. 22: X-24). See Fig. 2A. Nakama, Yaku-cho, Kumage-gun (^^?PM^ BTTft^fB^); Kagoshima, YAKUSHIMA; 30°16'N, 130°29'E; collected 20 Jul. 1987 by Mr. Imaegawa; PRIKU, 1 (skeleton only). Col- lected 26-27 Jul. 1987 by PRIKU staff; PRIKU, 3 (postcranials only). Collected 26-27 Jul. 1987 by Mr.Yasumaki; PRIKU, 7 (3 skeletons only, 4 skulls only). Collected 27 Jul. 1987 by PRIKU staff; PRIKU, 1 (skeleton only). Collected 27 Jul. 1987 by Mr. Mura- kami; PRIKU, 1 (skull only). SW102. Nakamura-city (4^^TtJ); Kochi, SHIKOKU; ca. 32°59'N, 132°56'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW79. Nakanogoudani vicinity, Samoto-mura, Nishi- muro-gun (ffi#«lPfe^^^f >f ^ ^P#ffe); Wakayama, HONSHU; 33°35'N, 135°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-31). SeeFig. 2A. Nakao, Tomisato-mura, Uma-gun (^JSIPS^P W 4^ M ); Ehime, SHIKOKU; 33°53'N, 133°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18:IX-10). SeeFig. 2A. Nakaodani vicinity, Go-mura, Arida-gun (Wffl W^^^W^^y, Wakayama, HONSHU; 34°03'N, 135°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-5). See Fig. 2A. Nakase, Yaku-cho, Kumage-gun (^^^M^fflJ 4^^); Kagoshima, YAKUSHIMA; 30°16'N, 130°36'E; collected 23 Jul. 1987 by PRIKU staff; PRIKU, 3 (2 skeletons only, 1 postcrani- als only). SW102. Nakasegawa, Yaku-cho, Kumage-gun (^^^M X^^M]\\); Kagoshima, YAKUSHIMA; 30°16'N, 130°36'E; collected 21 Jul. 1987 by PRIKU staff: PRIKU, 1 (skeleton only). SW102. Nakasu, Suwa-city ( M IS rfT 4^ /jtl ); Nagano, HONSHU; 35°59'N, 138°07'E; collected 26 Jul. 1998 by PRIKU staff; PRIKU, 1 (man- dibular fragments and postcranials only). NE95. Nakatsugawa vicinity, Taisho-mura, Hata-gun (M^aPAlE^^4^}tJI|'te); Kochi, SHIKOKU; 33°14'N, 132°59'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: X-15). See Fig. 2A. Nakatsugawayama, Sakanoshita-mura, Su- zuka-gun (Ip El^^feT W 4^ /$)rI Uj ); Mie, HONSHU; 34°53'N, 136°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII- 10). See Fig. 2A. Nakayama, Takayama-mura, Kamitakai-gun (_h K#lPI^Uj^4^LiJ); Nagano, HONSHU; 36°4rN, 138°23'E; collected 19 Oct. 1998 by M. Aimi, PRIKU, 1 (skeleton only). Collected Nov. 1998 by PRIKU staff; PRIKU, 1 (skele- ton only). NE49. Namesawayama, Mitomi-mura, Higashiyamana- shi-gun (^UJ^^PHSW^/RUJ); Yamanashi, HONSHU; 35°51'N, 138°45'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-20). See Fig. 2A. Nametoko, Nogawa, Uwajima-city (^^PfiTtTi? JIlTiJ^); Ehime, SHIKOKU; ca. 33°12'N, 132°38'E; provisioned group; birth season, 1963-1966, reported by Kawai et al. (1967, pp. 37, 39). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). External measurements taken by Hamada et al. (1996a, pp. 98, 99). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SIV77. Namisegawara, Kitaura-son, Higashiusuki-gun {MB^Witm^^&mniW^y, MiyazaU, KYUSHU; coordinates unknown; reported in questionnaire survey conducted in 1923 by K. Hasebe (Mito,1989, p. 25). Not mapped. Nangai-mura, Senboku-gun (-(llldb^P I^^KW); Akita, HONSHU; 39°24'N, 140°24'E; mtDNA extracted from umayazaru (talismanic hunting trophy) before 2003 by T. Agatsuma and M. Ishigami (2002, p. 79). NEI5 . Nansei-cho, Watarai-gun (^^1^1^^ BJ); Mie, HONSHU; ca. 34°20'N, 136°41'E; observed Aug. 1979 by K. Masui (Takasaki & Masui, 1984, pp. 310, 317). 5^2i. Naruse. See Narusegawa. Narusegawa, ( 1l 0 Jl| ); Miyagi, HONSHU; 38°30'N, 141°04'E; observed Jul. 2002 by K. 158 FIELDIANA: ZOOLOGY Izawa, T. Uno, and H. Fujita (2003, p. 27). NE23. Natasho-mura, Onyu-gun (3si!(?P=SfflJiW); Fukiii, HONSHU; ca. 35°24'N, 135°41'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW42. National Forest, Akaishi-mura, Nishitsugaru-gun (ffi)$giP#;5;|^S#;i9^); Aomori, HONSHU; 40°44'N, ]40°irE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-6). See Fig. 2 A. National Forest, Fukaura-mura, Nishitsugaru-gun 0)*fe?P;^)t^^aW^); Aomori, HONSHU; 40°37'N, 139°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-2). See Fig. 2A. National Forest, Hokujo-mura, Kitaazumi-gun ( Jb S « ^ db ^ ^ S W ^ ); Nagano, HONSHU; 36°40'N, 137°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-22). See Fig. 2A. National Forest, Imabetsu-mura, Higashit- sugaru-gun (^^/^^^^SIJ^SW^); Aomori, HONSHU; 41°10'N, 140°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-14). See Fig. 2A. National Forest, Iwasaki-mura, Nishitsugaru-gun (M}$^IP^llit^SIW^); Aomori, HONSHU; 40°34'N, 139°57'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-1). See Fig. 2A. National Forest, Kamishiro-mura, Kitaazumi-gun ( db ^ » ?P tt J^ ^^r il W ^ ); Nagano, HONSHU; 36°38'N, 137°49'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-21). See Fig. 2A. National Forest, Matsukawa-mura, Kitaa- zumi-gun (db^ft^P^iJII^SW^I^^); Nagano, HONSHU; 36°25'N, 137°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI- 18). See Fig. 2A. National Forest, Minamiaiki-mura, Mina- misaku-gun (l^fe^^Pi^^lTK^ttH^^); Nagano, HONSHU; 36°00'N, 138°35'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-37). See Fig. 2A. National Forest, Nakatsuchi-mura, Kitaa- zumi-gun (db$#?P'^±^g#^); Nagano, HONSHU; 36°50'N, 137°57'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-23). See Fig. 2A. National Forest, Nishishiwa-son, Kamo-gun (I? j^?PIS.^^^^S##); Hiroshima, HONSHU; 34°29'N, 132°36'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV- 16). See Fig. 2A. National Forest, Oodose-mura, Nishitsugaru-gun (S/$^1PAF^^SW#); Aomori, HONSHU; 40°43'N, 140°04'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-5). See Fig. 2A. National Forest, Taira-mura, Kitaazumi-gun (db $#1P¥^1#^); Nagano, HONSHU; 36°33'N, 137°49'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-20). See Fig. 2 A. National Forest, Tairadate-mura, Higashit- sugaru-gun (K/$^1P¥^^@'^#); Aomori, HONSHU; 41°07'N, 140°37'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-1 5). See Fig. 2A. National Forest, Tokiwa-mura, Kitaazumi-gun (db^ftlPSM^lW*^); Nagano, HONSHU; 36°28'N, 137°49'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-19). See Fig. 2A. National Forest, Yuzono, Suki-son, Nishimoro- kata-gun (IS^miP^^Wtt^SW^); Mi- yazaki, KYUSHU; 32°05'N, 131°05'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-14). See Fig. 2A. National Forest vicinity, Kameyama-mura, Ki- mitsu-gun (^)*lPmiljWSW^^); Chiba, HONSHU; 35°13'N, 140°07'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 50: XXXVI-4). See Fig. 2A. Natsusaka vicinity, Kitayama-mura, Yama- FOODEN AND AIM!: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 159 gata-gun (OjmiPdbOJ^MJS^); Gifu, HONSHU: 35°40'N, 136°47'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIlI-14). See Fig. 2A. Negitani. Nishikawa-mura, Gujo-gun (?P_hlPS ]\\ ^ :^ =^' ^ ): Gifu, HONSHU: 35°48'N. 136°53'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 41 : XXVIII-25). See Fig. 2 A. Nenbutsudani National Forest, Minochi-son. Saeki-gun {^\^^:^P^^±\U"^mM^Y Hiroshima. HONSHU: 34°31'N. 132°18'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 26: XIV-2). See Fig. 2A. Neo-mura, Motosu-gun (^^IP^^^): Gifu. HONSHU: ca. 35°38'N, 136°37'E: mtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). NE59. Nezumiana, Matsukawa-mura. Kitaazumi-gun (db^^^P^A^JII^ETv:): Nagano, HONSHU: 36°24'N, 137°50'E: collected 22 Feb. 1998 by M. Aimi: PRIKU, 2 (skeletons only). NE92. Nigure, Kamianama-mura. Oono-gun (ASlP-h 7^,l^if •): Fukui, HONSHU: 35°51'N. 136°46'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 39: XXV-22). See Fig. 2A. Nikko-city (B^fellT): Tochigi. HONSHU: ca. 36°45'N, 139°37'E: albino monkey capuired before 1826 (von Siebold, MS., 1830-1842, p. 1: cf. Temminck. 1842. p. 10): specimen not preserved. Collected in 1875 by Mr. v. Hilgendorf (Schweyer, 1909. p. 8): ZMB. 8 (1 skeleton only. 7 skulls only). Eight skulls pur- chased in 1900-1901 by K. A. Haberer (Schweyer. 1909. p. 8): some or ail of these skulls may be included among unlocalized specimens examined in AIUM. Reported pre- sent before 1939 by N. Kuroda (1938. p. 112: cf. Study Group on the Present Status of Japa- nese Monkeys, 1977b, p. 26). Collected 8 Sep. 1981-24 Sep. 1986 by TPM staff: TPM. 21 (skulls only, excluding 10 specimens not seen, data from specimen list). MtDNA samples collected ca. 1986-1991 by Hayasaka et al. (1991, p. 400). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). External measurements taken by Hamada et al. (1996a, pp. 98, 99). MtDNA samples col- lected before 1998 by Y. Kawamoto (1997, p. 33:2002.p. 60).A'£-^i. Nikkosan. Yahata-mura. Arida-gun (WB9?PANI ^S3feaj): Wakayawa, HONSHU: 34°05'N, 135°27'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974, p. 33:XX-4). SeeFig. 2A. Nina-A. Hanyama. Kamiyaku-cho. Kumage-gun (^^lP±M:XfflI¥aj~:)-A||): Kagoshiiiw, YAKUSHIMA: 30°21'N. 130°23'E: field study conducted Aug.-Dec. 1976 by T. Maruhashi (1981, p. 2: cf.^982, p. 323; Soltis et al., 2000, p. 196). Feeding association with Cenus nip- pon observed Jan.-May 2002 by B. Majolo and R. Ventura (2004. p. 35). SWIOI. Ninomiya-machi. Naka-gun ( "^ ?P — §' fflj ): Kanaga^va, HONSHU: ca. 35°18'N, 139°16'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997. p. 33). NE105. Nireyama vicinity. Nire-mura, Kamitakai-gun ( ± it ^ IP t ^L ^ t ^L OJ fife ): Nagano. HONSHU: 36°36'N. 138°21'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 46: XXXI-31). See Fig. 2A. Nisakasanrin vicinity. Nigou-mura, Kita- muro-gun {i\L^UW-n^n^\^^^)\ Mie. HONSHU: 34°13'N. 136°20'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974. p. 37: XXIII-40). See Fig. 2A. Nishi. Ooshika-mura, Shimoina-gun (T'^SP^P AE^ffi): Nagano. HONSHU: 35°35'N, 138°02'E: collected 4 Dec. 1998 by PRIKU staff: PRIKU. 1 (skeleton only). NE70. Nishiazai-cho. Ika-gun (##?PS^#fflI): Shiga. HONSHU: ca. 35°29'N. 136°08'E: mtDNA samples collected before 2003 by Y. Kawamoto (2002. p. 60). SW4. Nishifijjiyama. Kami-mura, Shimoina-gun (T*^ IP ?P ± :14 S S Oj ): Nagano. HONSHU; 35°24'N, 137°58'E: reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974. p. 47: XXXI-47). See Fig. 2A. Nishiharuchika, Ina-city ("^IP rf5"ffi#]fi); Na- gano, HONSHU; 35°47'N, 137°56'E; collected 17 Mar.-I3 Aug. 1998 by M. Aimi; PRIKU, 160 FELDIANA: ZOOLOGY 1 1 (skeletons only, including 4 with frag- mented skulls). NE79. Nishihikasa, Kimitsu-city ( ^ /$ TtT g 0 ^ ); Chiba, HONSHU; 35°13'N, 139°59'E; col- lected 22 Aug. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE120. Nishihinobori, Kisuki-cho, Ohara-gun ( AJ^^^ ;^fflIMS^); Shimane, HONSHU; 35°15'N, 132°54'E; collected. 4 Dec. 1994 and 12 Feb. 1995 by PRIKU staff; PRIKU, 2 (skeletons only). SW59. Nishiizu-cho, Kamo-gun (Mj^^PM^aBI); Shizuoka, HONSHU; 34°46'N, 138°47'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NEIOO. Nishikatsura-cho, Minamitsuru-gun (j^li^li^ S^fflJ); Yamanashi, HONSHU; ca. 35°30'N, 138°5rE; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE98. Nishikomagadake, Ina-machi, Kamiina-gun (_b #ilP?P^ilWM|6)^l&); Nagano, HONSHU; 35°50'N, 137°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-41). See Fig. 2A. Nishikurokiyama. See Higashi- and Ni- shi-Kurokiyama. Nishimeya-mura, Nakatsugaru-gun (^'/♦S^PM iM^); Aomori, HONSHU; ca. 40°30'N, 140°15'E; captured alive, briefly held in cap- tivity, donated to JMC 22 Jul. 1966; JMC, 1 (skull only). NEW. Nishine, Kitaokuni-mura, Nishiokitama-gun (ffi amPdb^hS^ffi^); Yamagata, HONSHU; 38°10'N, 139°46'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLII-4). See Fig. 2A. Nishinomaki, Saimoku-mura, Kitakanra-gun (db ^^^'^'^^'Em^y, Gimma, HONSHU; 36°14'N, 138°40'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XL-3). See Fig. 2A. Nishinoyakaeyama, Saigou-son, Higa- shiusuki-gun {1^B^WmM¥^'^ )\^\\^); Miyazaki, KYUSHU; 32°24'N, 131°28'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-21). See Fig. 2A. Nishiogawa vicinity, Uchitomi-mura, Onyu-gun ( ':&m. ?P 1*1 ^Wi ^ H d^ Jll fife ); Eukui, HONSHU; 35°31'N, 135°47'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV- 1 1). See Fig. 2A. Nishiohira, Ichibu-mura, Kuma-gun (^^IP — E^ffiA^); Kumamoto, KYUSHU; 32°10'N, 130°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16:IV-2). SeeFig. 2A. Nishitokoo vicinity, Murohani-mura, Izushi-gun (tbSlP^il^S^Mffe); Hyogo, HONSHU; 35°26'N, 134°5rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII- 18). See Fig. 2 A. Nishitoyonaga-mura, Nagaoka-gun (:M[5??PMM ^W); Kochi, SHIKOKU; 33°49'N, 133°45'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-34). See Fig. 2A. Nishiyama, Minamiwada-mura, Shimoina-gun (T^jiPiP^^ffl^ffiOj); Nagano, HONSHU; 35°19'N, 137°55'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXI-50). See Fig. 2A. Nishiyama, Wada-mura, Shimoina-gun (T#3P iP^ffl^ffiOj); Nagano, HONSHU; 35°19'N, 137°57'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 47: XXXI-49). See Fig. 2A. Niu-mura, Ika-gun (##IP:I^^^); Shiga, HONSHU; 35°34'N, 136°15'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV- 12). See Fig. 2A. Nonesankei, Sakihama-mura, Aki-gun ($S?P feSMWS'aaj.^); Kochi, SHIKOKU; 33°28'N, 134°09'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-43). See Fig. 2A. Nonoboriyama vicinity, Nonobori-mura, Su- zuka-gun ($^SiPi?^^i?^aJ^); Mie, HONSHU; 34°56'N, 136°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII-9). See Fig. 2A. Nonogawa vicinity, Nishikamiyama-mura, Hata-gun (^i^iPS±aj:^i?P^^m^)\\X); Yamanashi, HONSHU; 35°40'N, 138°17'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-11). See Fig. 2A. Northern Japan Alps, Kitaazumi-gun and Mi- namiazumi-gun {i\:T )l7°7s), 600-3000 m elevation; Nagano, HONSHU; ca. 36°10-30'N, 137°40-50'E; 22 troops studied 26 Dec. 1996-21 Feb. 2000 by S. Izumiyama et al. (2003, p. 465). Not mapped (see Tatai and nearby localities). Northern villages, Tsuno-gun (tPii^Pdb^^^); Yamaguchi, HONSHU; 34°14'N, 131°51'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII-12). SeeFig. 2A. Nozokinosawa vicinity, Kasshi-mura, Kami- hei-gun (±r^#lP¥^Wy V^y)Rffe); Iwate, HONSHU; 39°14'N, 14r49'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Mito, 1989, p. 27). See Fig. 2A . Nuidouishiyama, Sai-mura, Shimokita-gun (Tdb fPfe^^i^jiSOj); Aomori, HONSHU; 41°23'N, 140°53'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-22). See Fig. 2A. Nutazawa, Kimitsu-city (:^/$rfT.^.ffl)H); Chiba, HONSHU; 35°12'N, 140°01'E; collected 20 Aug. and 16 Dec. 1998 by PRIKU staff; PRIKU, 2 (skeletons only). NE120. Nyudougatake, Tsubaki-mura, Suzuka-gun (IpM fP^^AM;!)'"*^); Me, HONSHU; 34°58'N, 136°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII-7). See Fig. 2A. Nyudouzan vicinity, Toyohara-mura, Nishi- muro-gun (ffi^SlPMJ^^Ajllijffe); Wa- kayama, HONSHU; coordinates unknown; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-38). Not mapped. Obi, Masutomi-mura, Kitakoma-gun (Jb^S^ iis^^J^M); Yamanashi, HONSHU; 35°52'N, 138°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIlI-18). See Fig. 2A. Ochiai, Ooshika-mura, Shimoina-gun (T'^SPIP AEW^H-); Nagano, HONSHU; 35°34'N, 138°02'E; collected 14 Dec. 1998 by PRIKU staff; PRIKU, 2 (skeletons only). NE70. Odairayama vicinity, Hasegawa-mura, Oono-gun (Ai??P^#JI|^m¥ajffe); Oita, KYUSHU; 32°52'N, 131°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: III-3). See Fig. 2A. Odake, Kurihashi-mura, Kamihei-gun (_h^'(^?P ^M^^^); Iwate, HONSHU; 39°20'N, 141°43'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Mito, 1989, p. 25). See Fig. 2A. Odawara-city ( ^J"* ffl J^ rfT ); Kanagawa, HONSHU; ca. 35°15'N, 139°10'E; mtDNA samples collected before 1998 by Y. Kawa- moto (1997, p. 33; 2002, p. 60). NEJ04. Odo, Ohtsuki-cho, Hata-gun (<#^?PA>^ fflJA ^); Kochi, SHIKOKU; ca. 32°47'N, 132°38'E; blood samples collected before 1992 by No- zawa et al. (1991, p. 414; 1996, p. 6). SW80. Odomisaki, Ohtsuki-cho, Hata-gun (^^^1PA>^ ffll A ^ llif ); Kochi, SHIKOKU; 32°46'N, 132°38'E; external measurements taken by Hamada et al. (1996a, pp. 98, 99). SW80. Ofunato-city (A^p^y^gTfT); Iwate, HONSHU; ca. 39°04'N, 14r43'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33). NE21. Ogachi-machi, Ogachi-gun ( ^ 0 ?P 4i 0 fflj ); Akita, HONSHU; ca. 39°00'N, 140°19'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NEI7. Ogadake, Miyori-mura, Shioya-gun (^^IPH i^^^^^y, Tochigi, HONSHU; 36°58'N, 139°4rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51:XXXIX-6). SeeFig. 2A. Ogawa-mura, Higashimuro-gun (^^ftlPd'^JH ^ ); Wakayama, HONSHU; 33°36'N, 135°48'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34:XX-41). SeeFig.2A. Ogawadani, Yamazaki-mura, Shimoniikawa-gun ( T ^ Jll ?P UJ ^ ^ ^h Jll # ); Toyama, HONSHU; 36°55'N, 137°34'E; reported in 162 FIELDIANA: ZOOLOGY questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 42: XXIX- 11). See Fig. 2A. Ogawaoku vicinity, Kamikawa-mura, Minami- muro-gun (^#■glP#JI|^^tmJllllfte); Mie, HONSHU; 33°56'N, 136°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-52). See Fig. 2A. Ogikubo, Takayama-mura, Kamitakai-gun (-hi§ ^M>m\h¥i^^)\ Nagano, HONSHU; 36°41'N, 138°25'E; collected 26 Sep.-12 Nov. 1997 by M. Aimi; PRIKU, 2 (skeletons only). Collected 7 Aug. 1998 by PRIKU staff; PRIKU, 1 (skeleton only, with fragmented skull). NE49. Ohamidake ( ADtlS), 3050 m elevation; 36°20'N, 137°39'E; Nagano, HONSHU; reported pre- sent before 1988 by S. Izumiyama (1987a, p. 8; \9^1h,p.A2).NE88. Ohara vicinity, Kitatani-mura, Oono-gun (Ai? IPdk^Wd^Jiffe); Fukid, HONSHU; 36°05'N, 136°35'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV-20). See Fig. 2A. Oharadake, Yachi-mura, Ichishi-gun (— ^n^A ^ ^ d^ J^ H ); Mie, HONSHU; 34°33'N, 136°15'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37:XXIII-19). SeeFig. 2A. Ohira National Forest vicinity, Kobayashi-cho, Nishimorokata-gun (SMli^^d^^fflTA^S M ^^); Miyazaki, KYUSHU; 32°00'N, 131°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15:11-13). SeeFig. 2A. Ohirayama, Inuyama-city (i^lilTfiA^^lll); Aichi, HONSHU; 35°23'N, 136°56'E; variably provi- sioned population, translocated from Yaku- shima in Mar. 1957 (Kawai, 1960, pp. 204, 222; cf. Fooden & Aimi, 2003, p. 111). Birth season, 1957-1966, reported by Kawai et al. (1967, p. 39). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 7). NE6L Ohsawakanrin, Hourai-mura, Kitakoma-gun (db &0?Pm5lt^A)HtW); Yamanashi, HONSHU; 35°50'N, 138°16'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII- 16). See Fig. 2A. Oide, hills near. See Omachi, hills near. Oitsuge vicinity, Nishiki-mura, Kitamuro-gun (db^SiPII^7t->r7PqIt^liaj)H SW^); Akita, HONSHU; 40°00'N, 140°19'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLIV-6). See Fig. 2A. Oma-machi. Shimokita-gun ( T db IP A Pal fflJ ); Aomori. HONSHU; ca. 41°32'N. 140°55'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997. p. 33). NEl. Omachi. hills near (Affllrfi) (= Oide, hills near), HONSHU; ca. 36°30'N, 137°52'E; collected 12 Nov. 1882 by P. L. Jouy; USNM, 1 (specimen not located, information from USNM field catalog). 7V£97. Omachi-city (AffllTtJ); Nagano, HONSHU; ca. 36°30'N, 137°52'E; mtDNA samples collected before 1999 by Y. Kawamoto (1998, p. 54; 2002, p. 60). NE90. Ome-city ( ff ^S rfi ); Tokyo Metropolis, HONSHU; ca. 35°47'N, 139°15'E; mtDNA samples collected before 1998 by Y. Kawa- moto (1997. p. 33). 7V£77i. Omishima. Senzaki-cho, Otsu-gun (A^lP'flljil'^ fflJff^S) (= Oomishima; Oumishima); Ya- maguchi, OMISHIMA; ca. 34°25'N. 131°11'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974, p. 25: XIII-5). Occurrence until 1950 reported by K. Kawanaka (1973, p. 115). population subse- 164 FIELDIANA: ZOOLOGY quently extinct. SW67. Onagarayama Forest, Nakano-mura, Minamia- mabe-gun (]^)§^?P4^if;f^r^^¥aJ^); Oita, KYUSHU; 32°59'N, 131°48'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: III- 12). See Fig. 2A. Ondayama vicinity, Yunoyama-mura, Onsen-gun (S^^iP^ULl^i^fflUjffe); Ehime, SHIKOKU; 33°53'N, 132°5rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: IX-23). See Fig. 2A. Onigajo, Utsui-son, Toyoura-gun (sM^rt 0 ^%Ti0.); Yamaguchi, HONSHU; 34°07'N, 130°58'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII- 1). See Fig. 2 A. Onigajo vicinity, Tateyama-mura, Nakanii- kawa-gun (4^if JlllPilLil^^^r^ffe); To- yama, HONSHU; 36°38'N, 137°2rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 42: XXIX-5). See Fig. 2A. Onoaida, Yaku-cho, Kumage-gun ( M.^W>MP^ ^ M ^f^ ); Kagoshima, YAKUSHIMA; 30°15'N, 130°33'E; reported origin of monkeys kept at Isokoen {W^%'^^''j^M), a park in Kagoshima City; blood samples were collected from these captives before 1 992 by Nozawa et al. (1991, p. 414; 1996, p. 7; T. Shotake, PRIKU, pers. comm., Nov. 2003). Not mapped. Onoaida-Kurio. See Yakushima, Census Area 5. Onomi and Otaniyama, Matsuyama-mura, Soo-gun (^J^^^iOl^^rMifm- A^UJ); Kagoshima, KYUSHU; 31°36'N, 13r06'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: I-U). See Fig. 2A. Onosanrin, Miwa-son, Saeki-gun ('fe'ffi^H^^ Ai?lil#); Hiroshima, HONSHU; coordinates unknown; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-3). Not mapped. Onoueyama, Sashiki-mura, Arida-gun (Wffl^S M >^ ^^ ^ i? ± Lil ); Wakayama, HONSHU; 34°04'N, 135°19'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX- 1). See Fig. 2A. Onozumi vicinity, Misaki-mura, Hata-gun (ifil^ IPHllit^^^^ffe); Kochi, SHIKOKU; 32°48'N, 132°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: X-3). See Fig. 2A. Oobara, Kisofukushima-machi, Kiso-gun (^e ^^Kt^fifflJAJ^); Nagano, HONSHU; 35°50'N, 137°45'E; collected Nov. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE81. Ooboke, Sanmyou-son, Miyoshi-gun (H^^H ^ ^ A ^ fe ); Tokushima, SHIKOKU; 33°54'N, 133°45'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-20). See Fig. 2 A. Oobora-Satani, Tara-mura, Yourou-gun (ft^lP ^ ^^ ±MR^"Sy, Gifu, HONSHU; 35°16'N, 136°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVIIl-4). See Fig. 2 A. Ooborayama, Ootaki-mura, Chichibu-gun (^^ fP A )t ^^t A )Iql OJ ); Saitama, HONSHU; 35°50'N, 138°53'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 50: XXXVII-2). See Fig. 2A. Ooda-city (ABSTtT); Shimane, HONSHU; ca. 35°08'N, 132°36'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW60. Ooe Takayama, Soshiki-mura, Oochi-gun (b^ lPfi5t^A>IlfUj); Shimane, HONSHU; 35°05'N, 132°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-21). See Fig. 2A. Ooflikasawayama, Shichigashuku-mura, Ka- tsuta-gun ( XIJ Ba IP ^= ^ =fS ^ A )f )K liJ ); Mi- yagi, HONSHU; 38°02'N, 140°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLIII-3). See Fig. 2A. Oogawara, Aiga-mura, Koka-gun (¥M?PI(&/R[ ^ A M M ); Shiga, HONSHU; 34°57'N, 136°22'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV-25). See Fig. 2 A. Ooike beach, Sugari-mura, Kitamuro-gun (it^ mw>mnm^Ammmy, Mie, honshu; 34°06'N, 136°16'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-43). See Fig. 2A. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 165 Ooiriyama. Sonomura, Kitashitara-gun (db^?^ IPi^AAOj): AichL HONSHU; 35°06'N. 137^44'E; reported in questionnaire sur\ey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 40: XXVII-6). See Fig. 2A. Ooishifukaoawairi \icinit\. Sekitani-mura. Iwa- t\ine-gun^(^^p^iPil#^AS^JI|Affe): Migata. HONSHU: 38°04'N. 139°34'E: re- ported in questionnaire sur\ey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 47: XXXII-'21). SeeFig. 2A. Ooiwa. Kimitsu-cit> ( ^ ^ TfT A qs ): Chiba, HONSHU: 35=12'N. 140°0rE: collected 18 Sep. 1998 and Mar. 1999 by PRIKU staff: PRIKU. 2 (skeletons only). \E120. Oojimasan. Takamori-machi. Shimoina-gun (T ^IPiPi^^BrrAlLij): Xagano. HONSHU: 35°34'N. 137^5rE: collected 6 Apr. and 14 May 1998 by PRIKU staff: PRIKU. 2 (skele- tons only). XE76. Ooka. Haguro-mura. Niwa-gun (^^?P^M^ 'J^E): Aic/iL HONSHU"; ^35°20'N. 136^58'E; reported absent in questionnaire sur\ey con- ducted in 1923 by K. Hasebe (Iwano. 1974. p. 40: XXVII-8). OokancNama. Noma-mura, Yosa-gun (-^^^PS ral^A^LiJ): Kyoto. HONSHU; 35=40'N. I35°10'E; reported in questionnaire surxey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 35: XXII-5). See Fig. 2 A. Ookawachiyama \icinit\. Funatsu-mura, Kita- muro-gun (db^«?P^a'^^^A>R[rt Ojffe); Uie. HONSHU; 34=13'N. 136°12'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 37: XXIII-44). ' See Fig. 2A. Ookawamae National Forest, Funaoka-mura. Kawabe-gun ()qIffllP^p'|5?^AJIIIil SW^); Akita, HONSHU; 39=38'N. 140=20'E; reponed in questionnaire sur\ey conducted in 1923 b\ K. Hasebe (Iwano, 1974, p. 56: XLIV-2). See Fig. 2A. Ooka\\ara. Ooshika-mura, Shimoina-gun (T# M^AB^AM^): Xagano. HONSHU; 35=33'N. 138=02'E; collected 12 Apr. 1998 by PRIKU statT; PRIKU. 1 (skeleton only). XEJO. Ookawara %icinirv. Neo-mura. Motosu-gun (^ m?P«M^A)qI^ffe); GifiL HONSHU; 35"44'N, 136"34'E; reported in questionnaire sur\ey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 41: XXVIII- 10). See Fig. 2 A. Ookueyama \icinit\. Kitagawa-son, Higa- shiusuki-gun (^ Q^SPdb'jH*^ AUb UU^); MiyazakL KYUSHU; 32°44TnJ, I31°30'E; re- ported in questionnaire sur\ey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 15: 11-27). See Fig. 2A. Oomine vicinit}.. Kogurusu. Iruka-mura. Mi- namimuro-gun (^^-SSPAJ^^d^^^A^ ffe); Mie. HONSHU; 33=52'N. 135°55'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 37: XXIII-54). See Fig. 2A. Oomishima. See Omishima. Oomiya-cho. Watarai-gun (S^IPA^BJ); Mie, HONSHU; 34°2rN. 136°27E; collected 11 No\. 1997 by PRIKU staff; PRIKU. 1 (skele- ton only). Slh2. Oosabiyama vicinitv. Takabayashi-mura, Nasu-gun (M^mM^^^^MOi^): To- chigL HONSHU; 37=03'N. 139°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 51: XXXIX-8). See Fig. 2A. Oosaki. Omachi-cit> ( A WJ rfJ A ^ ); Xagano, HONSHU; 36°29'N. 137°50'E; collected 2 Nov. 1998 by M. Aimi; PRIKU, 2 (skeletons only). XE90. Oosato. Onodani-mura, Minamimuro-gun (^^ $?PffiS^^AM); Mie. HONSHU: 33=47?^, 135°59'E; reported in questionnaire survey conducted in 1923 bv K. Hasebe (Iwano, 1974, p. 38: XXIII-56). See Fig. 2.\. Oosawano-machi, Kaminiikawa-gun (_L^JI|?P A)RSirr); Toyama. HONSHU; ca. 36=34?%', 137°12'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). XE56. Ooshibayama vicinity. Onikoube-mura, Tama- zukuri'-gun (3Ei^?P^M^A^aj^); Miyagi, HONSHU; 38°47'N. 140=40'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 55: XLIII-8). See Fig. 2A. Ooshima-mura. Ooi-gun ( AIS?PA^^); Fukiii, HONSHU; 35=32^, 135=39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974. p. 39: XXV-7). See Fig. 2A. 166 nELDI.\NA: ZOOLOGY Oosugitani-mura, Taki-gun (^"^^A^^^^); Mie, HONSHU; 34°16'N, 136°13'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXlII-28). See Fig. 2A. Ootani, Makita-mura, Yourou-gun (ft^lP4$![ffl ^A#); Gifu, HONSHU; 35°20'N, 136°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVIII-6). See Fig. 2A. Ootaniyama vicinity, Yamanaka-son, Mit- sugi-gun (^H^OJ^^^A^OJ^); Hi- roshima, HONSHU; 34°26'N, 133°06'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-27). See Fig. 2A. Ootashiro, Ootaki-machi, Isumi-gun (MP^^A ^* HI AS -ft); Chiba, HONSHU; 35°14'N, 140°10'E; collected 9 Aug. 1997 by PRIKU staff; PRIKU, 1 (skeleton only). NE126. Ootsuneki, Tabayama-mura, Kitatsuru-gun (dblrt^ ^ m ¥^ 1^\^^ ±n :^); Yamanashi, HONSHU; 35°48'N, 138°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-24). See Fig. 2A. Oouchi-mura, Igu-gun (PM.W>A\^^ : ^B^ MUL|c^U<^); Miyagi, HONSHU; 37°52'N, 140°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55:XLIII-2). SeeFig. 2A. Ooyahongou, Ooya-mura, Nima-gun (MB^A E^AE^^); Shimam, HONSHU; 35°02'N, 132°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-27). See Fig. 2A. Oozakurayama, Kiragawa-mura, Aki-gun ($S ^P^MJII^A^OJ); Kochi, SHIKOKU; 33°20'N, 134°08'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-45). See Fig. 2A. Oregon. See Beaverton. Oshibuchi vicinity, Hobara-mura, Watarai-gun (Jt^?PaJl^^^)^Jffe); Mie, HONSHU; 34°2rN, 136°4rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXlll-36). See Fig. 2A. Oshihara-mura, Nima-gun ( S B ?P ^n J^ ^ ); Shimane, HONSHU; coordinates unknown; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-31). Not mapped. Oshioi, Takachiho-cho, Nishiusuki-gun (ffiQ^ Wm^%^'Ml^^); Miyazaki, KYUSHU; 32°42'N, 131°18'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-31). See Fig. 2 A. Oshiro, Nukata-cho, Nukata-gun (MBg^P^BaBJ A -ft ); Aichi, HONSHU; ca. 35°00'N, 137°25'E; blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). NESS. Osuzu National Forest, Tsuno-cho, Koyu-gun HPumm^m^m^m^^y, Miyazaki, KYUSHU; 32°17'N, 131°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-19). See Fig. 2A. Ota, Yahata-mura, Kusu-gun (3B^S^1PA<^^A B9); Oita, KYUSHU; 33°22'N, 13r08'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: III-18). See Fig. 2A. Otayama, Iwakura-mura, Arida-gun (WffllP^ ^;l^ABgiiJ); Wakayama, HONSHU; 34°03'N, 135°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33:XX-3). SeeFig. 2A. _^ Otoumi, Takahama-cho, Ooi-gun (At5?Pi=i^ ^=^M); Fiikiii, HONSHU; ca. 35°32'N, 135°29'E; collected winter 1967 by K. Wata- nabe (1978, p. 37); PRIKU, 1 (skull only, mandible missing). SW47. Otsu-city ( A /* TtT ); Shiga, HONSHU; ca. 35°00'N, 135°52'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW35. Otsubo, Kameda, Futtsu-city (S/$rtT%ffl Aif); Chiba, HONSHU; 35°15'N, 139°52'E; Early Jomon subfossil (age, ca. 12 Ka) reported be- fore 2003 by M. Kuroda (2002a, p. 1 15); Wa- seda University, 1 (mandible only; not seen). NE119. Otsukamebora, Nakanoho-mura, Mugi-gun (it m^^^^^Z.%m); Gifu, HONSHU; 35°34'N, 137°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIII-20). See Fig. 2A. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 167 Ouchiyama-mura, Watarai-gun ( i^ ^ ?P A rt ill ^y, Mie, HONSHU; 34°17'N, 136°20'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-29). See Fig. 2A. Oudake, Kamikuishiki-mura, Nishiyatsu- shiro-gun (ffiA^t?P± A-fe^ill); Ya- manashi. HONSHU; 35°31'N, 138°37'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXIII-33). See Fig. 2A. Oumishima. See Omishima. Outaki-mura, Kiso-gun (^H?Pi/^^); Na- gano, HONSHU^ ca. 35°48'N, 137°33'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NESS. Ouyama vicinity, Oosugi-mura, Nagaoka-gun {^^^±Y^¥iM^\hh); KochU SHIKOKU; 33°48'N, 133°43'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-33). See Fig. 2A. Owarifuji vicinity, Ikeno-mura, Niwa-gun (^^ iP>fei?WM?is±ffe); Aichi, HONSHU; 35°20'N, 137°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVII- 1 1). See Fig. 2A. Owase-city ( M M TtJ ); Mie, HONSHU; ca. 34°04'N, 136°12'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW24. Oyama vicinity, Shimoukawa-mura, Takeno-gun (ttg^PT^JII^majfte); Kyoto, HONSHU; 35°44'N, 135°13'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXII-2). See Fig. 2A. Oyamada-mura, Ayama-gun (PRllijfPAl-ljffl^); Mie, HONSHU; caT 34°46'N, 136°14'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW19. Oza\\ayama, Sawame-mura, Yamamoto-gun (ill ^ ?P )R g ^ d^ )K LiJ ); Akita, HONSHU; 40°17'N, 140°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLIV-10). See Fig. 2 A. Ozoechinai. Okuchi-mura, Nomi-gun (BbH^PM U^MW^^P^); Ishikawa, HONSHU; 36°17'N, 136°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVI-5). See Fig. 2A. Rakanyama, Hiromi, Hikimikami-mura, Mino-gun (ll^^lPE5B±WiAB^:^ > Ul); Shimam, HONSHU; 34°31'N, 132°00'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-12). SeeFig. 2A. Rebunto (^L:^ft); Hokkaido, REBUNTO; ca. 45°23'N, 141°02E; 2 humeri (artificially modi- fied) and 1 ulna — all 3 presumably imported to Rebunto — collected at archaeological site be- fore 1982 (Nishimoto, 1981, p. 430). Not mapped. Rengeiwa vicinity, Mikata-mura, Shisou-gun (5^ M^^y^^W.^^^); Hyogo, HONSHU; 35°irN, 134°36'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-9). See Fig. 2 A. Rokujuri, Azuma-mura, Higashitagawa-gun (M fflJIIiP^^/N + M); Yamagata, HONSHU; 38°3rN, 140°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLII-7). See Fig. 2A. Rokujuri. Hongou-mura, Higashitagawa-gun {M BaJH^^M/N+S); Ya/nagata, HONSHU; 38°34'N, 139°48'E; reported in questionnaire sur\ey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLII-6). See Fig. 2A. Rome, Italy; 4r56'N, 12°30'E; population trans- located from Takasakiyama. Oita Prefecture, ca. 1977 (Cozzolino et al., 1992, p. 331; Coz- zolino & Schino, 1998, p. 860). Not mapped. Rosando, Uchinomi-cho, Shozu-gun ('J^'S.lPrt M^ ^^^ M ); Kagawa. SHODOSHIMA; 34°3rN, 134°18'E; provisioned group; birth season, 1958-1962, reported by Kawai et al. {\967,pp.31,3S).SW68. Ryogamisan, Ryogami-mura, Chichibu-gun (^ 5»(^$ Jl| ^fltt Uj ftfe); Wakayama, HONSHU; 33°49'N, 135°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-14). SeeFig. 2A. Ryujinyama vicinity, Inari-mura, Nishimuro-gun (S^ftlP^^WffittUjffe); Wakayama, HONSHU; 33°45'N, 135°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX- 19). See Fig. 2A. Ryujinyama vicinity, Kamiakitsu-mura, Nishi- muro-gun (®^«lP±ti:)tWil4^U4flfe); Wakayama, HONSHU; 33°45'N, 135°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-20). See Fig. 2A. Ryuoka-mura National Forest, Ochi-gun (j^^?P ^^¥iUM^); Ehime, SHIKOKU; 33°57'N, 132°58'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21:IX-26). SeeFig. 2A. Ryuozan, Miyauchi, Yawata-son, Mitsugi-gun (ffllUlPAIf ^^rtfEiOj); Hiroshima, HONSHU; 34°28'N, 133°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-28). See Fig. 2A. Ryuuzuyama, Tsudani-son, Yamagata-gun (ULj!^ IPIS^^^HMUj); Hiroshima, HONSHU; 34°39'N, 132°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-8). See Fig. 2A. Sabiyama. See Shiobara-mura. Saburi-mura, Ooi-gun {i^WM^'^%\\^); Fukui, HONSHU; 35°26'N, 135°33'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV-4). See Fig. 2A. Sagara-mura, Kuma-gun (^^?P^I.^W); Ku- mamoto, KYUSHU; ca. 32°14'N, 130°48'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW91. Sai-mura, Shimokita-gun (TdblP'fc^W); Ao- mori, HONSHU; 41°26'N, 140°52'E; reported present before 1939 by N. Kuroda (1938, p. 112; 1940,p. 270).A^£'2. Saihara-mura, Kitatsuru-gun (dbtPH^PffiJ^^); Yamanashi, HONSHU; 35°41'N, 139°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-26). See Fig. 2A. Sakasegawa vicinity, Itsuki-mura, Kuma-gun (^ BW>^^^J^M}\\i^y, Kimamoto, KYUSHU; 32°24'N, 130°5rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: IV-7). See Fig. 2 A. Samuhikiyama, Ooasa-son, Yamagata-gun (|JL|!^ fPAiS^^^^Oj); Hiroshima, HONSHU; 34°47'N, 132°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-9). See Fig. 2 A. Samusawa, Yamanouchi-machi, Shimotakai-gun (TiS#iPajyrtfflI*)R); Nagano, HONSHU; 36°43'N, 138°24'E; collected 2 Aug. 1997-21 Aug. 1998 by M. Aimi; PRIKU, 9 (skeletons only, including 5 with fragmented skulls). NE49. Samusawayama, Oota-mura, Hienuki-gun (#S iP;l!^BaW*>RUj); Iwate, HONSHU; 39°23'N, 141°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLV-8). See Fig. 2A. Sanada-machi, Chiisagata-gun (d^l^iPKEQWI); Nagano, HONSHU; ca. 36°27'N, 138°20'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE46. Sanda, Kimitsu-city ( ^ /$ T|T H BQ ); Chiba, FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 169 HONSHU; 35°19'N, 140°04'E; collected Dec. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE128. Sangayama, Katakaidani-mura, Shimonii- kawa-gun (TWf JUIPKM^^H® Uj); To- yama, HONSHU; 36°46'N, 137°31'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 42: XXIX-8). See Fig. 2A. Sangenyama, Ochiai, Osaka-cho, Masuda-gun (MBa^Pd^^fflJ^-^HrB^Oj); Gifii, HONSHU; 35°56'N, 137°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIII-32). See Fig. 2A. Sano-goryorin, Sakae-mura, Nishiyatsushiro-gun {mJ\i^W>^^i^mW^^); Yamanashi, HONSHU; 35°18'N, 138°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXIII-37). See Fig. 2A. Sanyoushi, Minmaya-mura, Higashitsugaru-gun (K)igl5HM^^^fflef); Aomori, HONSHU; 41°10'N, 140°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-13). See Fig. 2A. Sarugababa, Tonoi-son, Toyoura-gun (^y^^Pi^ E^+)-)l:^J\J\); Yamaguchi, HONSHU; 34°16'N, 13r03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25: XIII-3). See Fig. 2A. Sarugamiyama, Manzai-son, Atetsu-gun (Pr[^?P 7T M ^ ?i tt UJ ); Okayama, HONSHU; 34°53'N, 133°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV-5). See Fig. 2 A. Sarutsubo vicinity, Sada-son, Kimotsuki-gun (if ]i?P^i^^^i¥ftfe); Kagoshima, KYUSHU; 31°06'N, 130°48'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-18). See Fig. 2A. Sasagamine, Kadono-mura, Hikami-gun (7j<±^ Si?W^^ll#); Hyogo, HONSHU; 35°10'N, 135°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII-23). See Fig. 2A. Sasagou vicinity, Mimaki-mura, Kitauwa-gun (db^^iPISl^^^^ffe); Ehime, SHIKOKU; 33°03'N, 132°40'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18: IX-5). See Fig. 2A. Sasayama-cho, Taki-gun ( ^ ^ IP li Ol fflj ); Hyogo, HONSHU; ca. 35°08'N, 135°21'E; captured alive, arrived at JMC 26 Feb. 1981, died 16 Apr.-21 May 1981; JMC, 6 (skulls only). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW39. Satsuma-cho, Satsuma-gun ( M B ?P ^ JS fflj ); Kagoshima, KYUSHU; ca. 31°55'N, 130°32'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW93. Satsuma-gun (MS^); Kagoshima, KYUSHU; 3r54'N, 130°35'E; reported absent in ques- tionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-7). Not mapped. Seibu-rindou, Kamiyaku-cho, Kumage-gun {W( ^^±Jl^fflTM^^3i); Kagoshima, YAKUSHIMA; 30°23'N, 130°25'E; collected Feb.-Mar. 1990 by PRIKU staff; PRIKU, 2 (skeletons only, including 1 with mandible missing and 1 with fragmented skull). SWIOl. Sekinosawa, Ikawa-mura, Abe-gun (^io^^JH ^ll;^)R); Shizuoka, HONSHU; 35°12'N, 138°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX-7). See Fig. 2A. Senbagatake vicinity, Hiwasa-cho, Kaifu-gun (^ ^^BWtefflI=F^%^¥^W^TU)- Osaka, HONSHU; 34°52'N, 135°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XIX-4). See Fig. 2A. Shakujougadake, Kouchi-mura, Ano-gun {^M. WM^^^^T^); Mie, HONSHU; 34°49'N, 136°2rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII- 1 2). See Fig. 2 A. ^ Shibisan, Takaono-mura, Izumi-gum (tij^K^isi m if ^ ^ M UJ ); Kagoshima, KYUSHU; 32°02'N, 130°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-6). See Fig. 2A. Shichigashuku-machi, Katsuta-gun (XlJB9?P"t'^ ?i fflj ); Miyagi, HONSHU; ca. 38°00'N, 140°25'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NESS. Shichigodake, Nakama, Yaku-cho, Kumage-gun {M^^mX^^m^'^^); Kagoshima, YAKUSHIMA 30°17'N, 130°30'E; collected 27 Jul. 1987 by PRIKU staff; PRIKU, 2 (skeletons only). SfF7 02. Shichikawa-mura, Higashimuro-gun (^#ft?P -bJH^); Wakayama, HONSHU; 33°35'N, 135°44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-40). See Fig. 2A. Shichimensanroku, Motodate-mura, Mi- namikoma-gun (i^i!S?P^B^-b® lilM); Yamanashi, HONSHU; 35°24'N, 138°22'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-5). See Fig. 2A. Shichisou-cho, Kamo-gun (iPj^^P-b^fflJ); Gifu, HONSHU; ca. 35°34'N, 137°06'E; mtDNA samples collected before 1998 by Y. Kawa- moto (1997, p. 33; 2002, p. 60). NE60. Shiga Heights. See Shiga-kogen. Shiga-A. See Jigokudani. Shiga-kogen, Shimotakai-gun (Ti^^lP^xMi^ J^) (= Shiga Heights; includes Yokoyugawa Valley = Shiga-B); Nagano, HONSHU; ca. 36°42'N, 138°29'E; collected 6 Jul. 1962 and 16 Apr. 1963 by PRIKU staff; PRIKU, 2 (skulls only, including 1 with mandible miss- ing). MtDNA samples collected ca. 1986-1991 by Hayasaka et al. (1991, p. 400). Blood sam- ples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). External measure- ments taken before 1997 by Hamada et al. (1996a, pp. 98, 99). MtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NE48. Shigatakiyama vicinity, Tanami-mura, Nishi- muro-gun (ffi#«?P BQ M*^f .^^Miiaj ^); Wakayama, HONSHU; 33°30'N, 135°44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-35). See Fig. 2A. Shigekiyama, Midorisouzu-mura, Mi- namiuwa-gun (it^^^lP-^iitP^MTKUj); Ehime, SHIKOKU; 33°00'N, 132°35'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18: lX-3). See Fig. 2A. Shigira Shinyabakei, Kakiyama-mura, Shi- moge-gun (T^?P# OJ ^fimj^IP.!)^); Oita, KYUSHU; 33°23'N, Oril'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: III- 16). See Fig. 2A. Shigumi, Futtsu-city ( g /$ TfT /"^s ^ ); Chiba, HONSHU; 35°11'N, 139°57'E; collected 19 Jan.-28 Feb. 1998 by PRIKU staff; PRIKU, 3 (skeletons only). NE120. Shihatsuyama, Yahata-mura, Oita-gun (A'^^ Ri^^mUlU); Oita, KYUSHU; 33°15'N, 131°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16:111-13). See Fig. 2A. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 171 Shikakubidani, Takara-son, Katsuura-gun (0M ^^B^^M'M^y, Tokushima, SHIKOKU; 33°59'N, 134°3rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI- 19). See Fig. 2 A. Shikakumadake, Shimogou-mura, Shimoge-gun (T^^T®WS^I&); Oita, KYUSHU; 33°25'N, 131°08'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: III-14). See Fig. 2A. Shikakumadake, Yamautsuri-mura, Shimoge-gun (T^fPUj^^^ESIS); Oita, KYUSHU; 33°25'N, 131°10'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: III- 15). See Fig. 2A. Shikamizawa, Kamiyaku-cho, Kumage-gun (M. ^?P±M^fflTEB)R); Kagoshima, YAKUSHIMA; ca. 30°20'N, 130°23'E; ob- served Aug. 1978 by T. Maruhashi (1982, p. 318; cf. Takasaki & Masui, 1984, p. 311). SWIOI. Shimakachisanrin vicinity, Katsuragi-mura, Ki- tamuro-gun (db#«?P^^WftBUj^ftfe); Mie, HONSHU; 34°07'N, 136°17'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-42). See Fig. 2A. Shimashima, Azumi-mura, Minamiazumi-gun (]^$*?P$»^ftiite ); kocm, SHIKOKU; 32°53'N, 132°57'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: X-8). See Fig. 2A. Shimonotani vicinity, Sakauchi-mura, Ibi-gun (SH^Jt^rt^Ty^flfe); Gifu, HONSHU; 35°34'N, 136°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVIII-7). See Fig. 2A. Shimosakuma, Kyonan-machi, Awa-gun {^M ^iSl^fflTTfe^fBl); Chiba, HONSHU; 35°06'N, 139°51'E; collected 17 Jul. 1997-10 Feb. 1999 by PRIKU staff; PRIKU, 11 (skele- tons only). NE120. Shimosanji-mura, Hidaka-gun (Si^?PTLi-l£& ^ ); Wakayama, HONSHU; 33°53'N, 135°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX- 11). See Fig. 2A. Shimoseinaiji, Seinaiji-mura, Shimoina-gun (~F # 3P ?P )t 1*1 S& 4t T )f rt i?& ); Nagano, HONSHU; 35°30'N, 137°42'E; collected 7 Oct.-l Nov. 1998 by M. Aimi; PRIKU, 4 (skeletons only). NE73. Shimoyama (Till); Aomori. See Shimokita Pen- insula. Shimoyama, Kushikino-mura, Hioki-gun ( 0 S 15 $ tK if ;t^^t T 111 ); Kagoshima, KYUSHU; 31°43'N, 130°15'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-5). See Fig. 2A. Shimoyama vicinity, Shimomaki-mura, Mugi-gun (Stm^Ti^^^TUj^); Gifu, HONSHU; 35°36'N, 136°55'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIII- 17). See Fig. 2A. Shindenyama, Abu-son, Kaifu-gun (^pP^PPrIpP ^l/feaUj); Tokushima, SHIKOKU; 33°48'N, 134°39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23:XI-5). SeeFig. 2A. Shinjou vicinity, Mimi-mura, Mikata-gun {=.^5 ^:^WifJ±'f-)'JfiUJ#ftfe); Fukui, HONSHU; 35°35'N, 135°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV- 14). See Fig. 2A. Shinshiro-city (llf^TfT); Aichi, HONSHU; ca. 34°54'N, 137°30'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NE64. Shinshiu ("(§#1). See Nagano. Shinyashiki, lijima-machi, Kamiina-gun (_h#BP ?Pt5ftffriffMi(); Nagano, HONSHU; 35°39'N, 137°53'E; collected 12 Nov. 1997 by M. Aimi; PRIKU, 1 (skeleton only). NE77. Shinzan, adjacent to Mogami-gun, Nikko-mura, Akumi-gun {I^M^Umi^^mZ.^^^ ^ )^ Ol ); Yamagata, HONSHU; 39°02'N, 140°06'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55:XLII-9). SeeFig. 2A. Shinzan, adjacent to Mogami-gun, Oosawa-mura, Akumi-gun {^m^i^'R^rnhmzm^t ^ )^ UJ ); Yamagata, HONSHU; 38°59'N, 140°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55:XLII-8). SeeFig. 2A. Shinzan, Mizuho-mura, Shinobu-gun (M^^P^X -f^^Ht)^!!!); Fukushima, HONSHU; 37°45'N, 140°2rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 52: XLI-9). See Fig. 2A. Shiobara-mura, Nasu-gun (JSPM^^^J^W) (= Sabiyama; Shioyono); Tochigi, HONSHU; 36°58'N, 139°50'E; observed before 1939 by N. Kuroda (1938, p. 112; 1940, pp. 270, 272). NEW. Shiono, Suzaka-city ( ^ ^ Tti im i? ); Nagano, HONSHU; 36°38'N, 138°21'E; collected 28 Oct. 1998 by M. Aimi; PRIKU, 2 (skeletons only). NE49. Shioyono. See Shiobara-mura. Shippara, Futtsu-city ( S /$ rtT M i^ ); Chiba, HONSHU; 35°10'N, 139°54'E; collected 23 Dec. 1997-30 Mar. 1999 (excludes two specimens with unknown collection dates) by PRIKU staff; PRIKU, 9 (skeletons only). NEJ20. Shiragayama, Motoyama-cho, Nagaoka-gun {-B: ^iP^OjfflJSSUj); Kochi, SHIKOKU; 33°48'N, 133°35'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-32). See Fig. 2A. Shirahama-cho, Nishimuro-gun (H-^ft^PS*^ HI ); Wakayama, HONSHU; ca. 33°41'N, 135°2rE; onset of provisioning in 1952 re- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 173 ported by Editorial Committee of Nihonzaru (1977,p. 112).5^m _ Shiraiiama-machi, Awa-gun (^J^^PS^HJ); Chiba, HONSHU; ca. 34°54'N, 139°54'E; re- ported present before 1978 by Study Group on the Present Status of Japanese Monkeys, 1977b, p. 26). MtDNA samples collected be- fore 1999 by Y. Kawamoto (1998, p. 54). NE123. Shiraki, Hatsuki-mura, Isa-gun ('^'te?P^>^^ S ^ ); Kagoshima, KYUSHU; 32°04'N, 130°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12:1-8). See Fig. 2A. Shirakiyama vicinity, Mizukami-mura, Kuma-gun (^^?P7K±:|^fi7Klij'te); Kuma- moto, KYUSHU; 32°22'N, 131°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: IV-6). See Fig. 2A. Shirakoyama, Kamiyaku-cho, Kumage-gun (^ ^lP±M:^Brrajl|aj); Kagoshima, YAKUSHIMA; 30°27'N, 130°28'E; collected 25 Feb. 1989 by PRIKU staff; PRIKU, 22 (skeletons only, including 1 with fragmented skull, 1 with mismatched mandible, and 1 postcranials only). SWIOO. Shiranesan vicinity, Takai-mura, Kamitakai-gun (±S#lPS#^fi«Ujffe); Nagano, HONSHU; 36°39'N, 138°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-29). See Fig. 2A. Shirataki, Tasuki-son, Toyoura-gun (M/^lPffl^ :^fi^); Yamaguchi, HONSHU; 34°16'N, 131°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 25:XIII-4). SeeFig. 2A. Shiratani vicinity, Tokuyama-mura, Ibi-gun (S HiP^^Uj^e^fte); Gifu, HONSHU; 35°40'N, 136°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVIII-8). See Fig. 2 A. Shirayama National Forest, Kamishinjou-mura, Minamiakita-gun {mXm^±Wi^^ 6 Uj S W^); Akita, HONSHU; 39°49'N, 140°09'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLIV-5). See Fig. 2A. Shiroiwayama, Yamae-mura, Kuma-gun (^S?P Uj XL ^ S ^ UJ ); Kiimamoto, KYUSHU; 32°18'N, 130°46'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: IV-4). See Fig. 2A. Shirokiyama, Ichikawa-son, Takata-gun (i^BQlP TfT Jll ;^ fi tK Oj ); Hiroshima, HONSHU; 34°33'N, 132°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV- 17). See Fig. 2A. Shirokiyama vicinity, Miiri-son, Asa-gun ($fe ^PHA^S^Ojffe); Hiroshima, HONSHU; 34°34'N, 132°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV- 14). See Fig. 2A. Shiroyama, Hatagasako-mura, Kanoashi-gun (M S.?Ptt0ifi^J^aj); Shimane, HONSHU; 34°28'N, 13r44'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-6). See Fig. 2A. Shiroyama, Tsuwano-cho, Kanoashi-gun (MS iP)$^i?fflJ^|ij); Shimane, HONSHU; 34°28'N, 131°45'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-7). See Fig. 2A. Shizuoka, HONSHU; 34°35'-35°38'N, 137°30'-139°08'E; collected before 2002 by PRIKU staff; PRIKU, 1 (skeleton only). Not mapped. Shobuzaki, Yoshihama-mura, Kesen-gun (^"fllj iP^^^MSllif ); hvate, HONSHU; 39°09'N, 141°49'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56: XLV-5). See Fig. 2A. Shodoshima ( ^J^ S 1 ); Kagawa, SHODO- SHIMA; 34°23'-34°33'N, 134°07'-134°21'E; reported present before 1941 by N. Kuroda (1940, p. 270). MtDNA samples collected from captive-bom monkeys before 1987 by Hayasaka et al. (1986, p. 346). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). Not mapped. Shodoshima [1] ( d^ S. ft ); Kagawa, SHODOSHIMA; ca. 34°31'N, 134°16'E; non- provisioned, captured 12 Feb. 1962, died in captivity 6 Apr. 1962; JMC, 1 (skeleton only). Provisioned, collected 7 Feb. 1981 by Y. 174 FIELDIANA: ZOOLOGY Hamada; PRIKU, 46 (33 skeletons only, in- cluding 1 with fragmented skull; 3 skulls with partial postcranials; 5 skulls only; 5 mandibles with partial postcranials; and 1 mandible only). SIV68. Shodoshima [2] ( ''J'' S A ); Kagawa, SHODOSHIMA; 34°30'N, 134°18'E; external measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). SW68. Shogatake vicinity, Ichinose-mura, Yourou-gun {^^^-:^M^^tf'^i\k); Gifu, HONSHU; 35°16'N, 136°30'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVIII-5). See Fig. 2A. Shokawagoe vicinity, Kitatonda-mura, Nishi- muro-gun (M^«?Pdb S ffl ^^tiS JH Mffe); Wakayama, HONSHU; 33°39'N, 135°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-25). See Fig. 2A. Shorenjisan vicinity, Minowa-mura, Naga-gun (^Mi^Xft^^tffa^lljffe); Me, HONSHU; 34°35'N, 136°07'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIlI-16). See Fig. 2A. Shougenyama, Kamikane-mura, Higashiyama- nashi-gun (mOj^^Ptt^^^^iilil); Yama- nashi, HONSHU; 35°47'N, ]38°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-21). See Fig. 2A. Shouzawa, Kisofukushima-machi, Kiso-gun (^ MiP7K^^ftfflIIE)R); Nagano, HONSHU; 35°49'N, 137°44'E; collected 10 Dec. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE81. Shuri Castle, Shuri-Tounokura-cho, Naha-city (^SPHrtJMM^'MfflTMM^); Okinawa, OKINAWA; 26°13'N, 127°43'E; mtDNA ex- tracted in 1999 from skeletal fragments (age ca. 0.4 Ka) by T. Mouri et al. (2000, p. 87); frag- ments apparently derived from captives intro- duced from Yakushima. Not mapped. Sokujitsunomine, Kitakata-mura, Higa- shiusuki-gun (^Q^l^db^^^^TM 0 ^); Mi- yazaki, KYUSHU; 32°33'N, 131°27'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-28). See Fig. 2A. Soma-city (^l/^rfi); Fukushima, HONSHU; ca. 37°48'N, 140°55'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE38. Somakawa-mura, Kamiukena-gun (_L/?!^^tlii Jll^); Ehime, SHIKOKU; 33°46'N, 133°07'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: IX-19). SeeFig.2A. Sone vicinity, Minamiwauchi-mura, Minami- muro-gun (l^#g?Pj^li^[^^^^ffe); Mie, HONSHU; 33°58'N, 136°12'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-49). See Fig. 2A. Soratani, Kasubuchi-mura, Oochi-gun (e^?P^ M^V=P'^); SMmane, HONSHU; 35°05'N, 132°36'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 28: XVI-22). See Fig. 2A. Souro vicinity, Shimokawakuchi-mura, Hata-gun di^l^TJII P^^S-te); Kochi, SHIKOKU; 32°46'N, 132°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21 : X-2). See Fig. 2A. South Texas. See Arashiyama West. Sudoyama vicinity, Sudo-mura, Fuji-gun (sdi 15^5$;fvtMytUj'te); Shizuoka, HONSHU; 35°11'N, 138°47'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX- 13). See Fig. 2A. Sugadaira, Sanada-machi, Chiisagata-gun {^b^ ^ W g fflj ^ 5|Z ). Magano, HONSHU; ca. 36°31'N, 138°19'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE47. Sugawayama vicinity, Nahari-cho, Aki-gun (^ g^^¥^IJfflI^JIiajfte); Kochi, SHIKOKU; 33°24'N, 134°02'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-47). See Fig. 2A. Sugino-mura, Ika-gun (##^l!^i?W); Shiga, HONSHU; 35°33'N, 136°16'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV- 13). See Fig. 2A. Sumikawa vicinity, Hikimishimo-mura, Mino-gun (HJIIPEBT^^JH^); Shimane, HONSHU; 34°35'N, 131°57'E; reported in questionnaire survey conducted in 1923 by K. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 175 Hasebe (Iwano, 1974, p. 28: XVI- 1 1). See Fig. 2A. Sumita-cho, Kesen-gun {^i\hW>i^^^); hvate, HONSHU; ca. 39°08'N, 141°36'E; mtDNA samples collected before 1998 by Y. Kawa- moto (1997, p. 33). A^£7 9. Sumugitani, Miyajima-mura, Kitakuwada-gun (db#ffl?P^ft^^^#); Kyoto, HONSHU; 35°15'N, 135°32'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXII- 15). See Fig. 2A. Sunada, Amatsukominato-machi, Awa-gun ($ MlP^^d^^JIfflJ^Ba); Chiba, HONSHU; 35°07'N, 140°10'E; collected 7 Mar. 1998-30 Mar. 1999 by PRIKU staff; PRIKU, 19 (skeletons only, including 1 with fragmented skull). NE125. Sunomata vicinity, Ukegawa-mura, Higashi- muro-gun (m^ft^lfJII ^X ^ ^ ^ fife); Wakayama, HONSHU; 33°48'N, 135°48'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-50). See Fig. 2A. Suzawa, Minamishinano-mura, Shimoina-gun ( T # IP IP m ft )i ^ ^ )H ); Nagano, HONSHU; 35°17'N, 137°56'E; collected 13 Dec. 1998 by PRIKU staff; PRIKU, 1 (skele- ton only, with fragmented skull). NE68. Tabayama-mura, Kitatsuru-gun (db|[PI§?P^/^ \h^)\ YamanashU HONSHU; ca. 35°47'N, 138°56'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE114. Tachibana, Mugi-cho, Kaifti-gun {M^W^^l^ ^ ); Tokushima, SHIKOKU; 33°42'N, 134°26'E; collected 18 Mar. 2002 by M. Aimi; PRIKU, \.SW72. Tachibanaseiryuu National Forest, Nano- kawa-mura, Agawa-gun (n"JII?P^i?JIIW^ mUm^^)^ Kochi, SHIKOKU; 33°34'N, 133°08'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22:X-21). SeeFig. 2A. Tachigouchi, Muikaichi-mura, Kanoashi-gun (M £lPASTl?:ftlZ:)RlP«;i); Shimam, HONSHU; 34°2rN, 131°57'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-2). See Fig. 2A. Tadachi-goryorin, Tadachi-mura, Nishichi- kuma-gun (S^SiP BQ ll ^BQ ll^3^f^); Nagano, HONSHU; 35°36'N, 137°33'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXXI-4). See Fig. 2A. Tadachiokuyama, Ogawa-mura, Kanoashi-gun ( E S: iP ^J^ Jll ^ e ^ H Lil ); Shimane, HONSHU; 34°30'N, 131°47'E; reported in questionnaire sur\'ey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-8). See Fig. 2A. Tadeno, Asakura-mura, Kanoashi-gun (^.S:?P l^5tW#i?); Shimane, HONSHU; 34°21'N, 131°54'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-3). See Fig. 2A. Tado Mountain Range, Ishizu-mura, Kaizu-gun (^^)i?P^)i^^igLilM); Gifu, HONSHU; 35°08'N, 136°39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVIII- 1). See Fig. 2A. Tado Mountain Range, Shiroyama-mura, Kaizu-gun {M%^^\h¥i^^\^U); Gifu, HONSHU; 35°09'N, 136°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVIII-2). See Fig. 2A. Tadosan vicinity, Komi-mura, Kuwana-gun (1^ =glP*ll^^]gajfte); Mie, HONSHU; 35°08'N, 136°36'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII- 1). See Fig. 2 A. Tadosan vicinity, Tado-mura, Kuwana-gun (H ^IP^JgW^i^lijflfe); Mie, HONSHU; 35°07'N, 136°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII-2). See Fig. 2A. Taga-cho, Inukami-gun (:^_hlP^MfflI); Shiga, HONSHU; ca. 35°13'N, 136°17'E; mtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). SW9. Tagajo-ato, Tagajo-city (^M^TtT^M^Kl^); Miyagi, HONSHU; ca. 38°18'N, 141°01'E; mtDNA extracted from subfossils (age, Nara period, ca. 1.3 Ka) before 2003 by T. Agatsuma and M. Ishigami (2002, p. 84); Okumatsushima Jomon-mura History Museum, 2 (femur, tibia; specimens not seen). NE24. Tagura, Futtsu-city ( M /¥ rfT BQ ^ ); Chiba, HONSHU; 35°14'N, 139°57'E; collected 18 176 HELDIANA: ZOOLOGY Dec. 1998 and 30 Mar. 1999 by PRIKU staff; PRIKU, 5 (skeletons only). NE120. Tai-mura, lishi-gun (tS^fPBQ^^); Shimane, HONSHU; 35°12'N, 132°56'E; reported (as possibly temporary habitat) in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-35). See Fig. 2A. Taiheisanrin, Sakai-mura, Shimoniikawa-gun (T llfJlliPJtW;^¥aJ^); Toyama, HONSHU; 36°57'N, 137°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 42: XXIX- 12). See Fig. 2A. Taikoyama, Tsutsukawa-mura, Yosa-gun (%S} ?PMJII^;*!^i^aj); Kyoto, HONSHU; 35°42'N, 135°15'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35:XXII-4). SeeFig. 2A. Taimasan, Ikeda-mura, Shozu-gun (-'J'^S.^Pyfeffl ;^ A Jl^ Uj ); Kagawa, SHODOSHIMA; 34°irN, 134°irE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 24: XlI-5). See Fig. 2A. Taipei pet shop, Taiwan; 25°03'N, 121°30'E; purchased in Oct. 1957 by R. E. Kuntz; USNM, 1 . Not mapped. Taishakugawa vicinity, Nagato-son, Jinseki-gun (ttS^*/:iW^lRJI|-w); Hiroshima, HONSHU; 34°53'N, 133°15'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XIV-36). See Fig. 2A. Taishakukyo, Tojo-cho, Hiba-gun (i;b^?P^^ fflI^f^llB'5); Hiroshima, HONSHU; 34°52'N, 133°14'E; provisioned group; birth season, 1958-1962, reported by Kawai et al. (1967, pp. 37, 39). SW58. Takabagamori, Mitsuse-mura, Agawa-gun (ra'JII ^^M^M^'T^); Kochi, SHIKOKU; 33°34'N, 133°22'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 22: X-25). See Fig. 2A. Takagoyama (SSlJLl) (= Mt. Takago); Chiba, HONSHU ca. 35°10'-35°15'N, 139°55'- 1 40°00'E; provisioning of group III initiated in 1956-1957 (Suzuki, 1972, pp. 333, 334; Study Group on the Present Status of Japanese Mon- keys, 1977b, p. 26). Collected ca. 1961 by S. Azuma; JMC, 1 (skull only, mandible missing). Blood samples collected before 1992 by No- zawa et al. (1991, p. 414; 1996, p. 6). MtDNA samples collected before 1998 by Y. Kawa- moto (1997, p. 33). Not mapped. Takagoyama, Tamaki-mura, Kimitsu-gun (^/$: IP^^if SUl); Chiba, HONSHU; 35°13'N, 139°57'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 50: XXXVI-2). See Fig. 2A. Takagoyama A (i^^UjA); Chiba, HONSHU; ca. 35°14'N, 140°01'E; provisioned group; birth season, 1966, reported by Kawai et al. (1967, pp. 37, 39). NEI2(). Takagoyama S (i^gOjS); Chiba, HONSHU; ca. 35°12'N, 139°59'E; provisioned group; birth season, 1959-1966, reported by Kawai et al. (1967,pp. 37, 38).A^£/20. Takagoyama Tl (ifgOlTl); Chiba, HONSHU; 35°12'N, 139°59'E; provisioning initiated in 1960 (Suzuki, 1972, p. 333). Subgroup cap- tured in Mar. 1973, died of disease soon after capture (Koike & Shimamura, 1988, p. 74); SU, 96 (84 skeletons only, 12 postcraniais only). External measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). NE120. Takahama (S^); Fukui, HONSHU; ca. 35°29'N, 135°33'E; mtDNA samples collected from captive group tranlocated to PRIKU before 1987 by Hayasaka et al. (1986, p. 346). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). Microsatellite DNA of laboratory specimen analyzed before 1996 by Domingo-Roura et al. (1997, p. 358). Ex- ternal measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). MtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). SW46. _^ Takahama-cho, Ooi-gun (AtS^Pd^W"); Fukui, HONSHU; 35°28'N, 135°33'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV-3). See Fig. 2A. Takaharasan vicinity, Tamanyu-mura, Shioya-gun (Jt#?Pi^^Si^Ujffe); To- chigi, HONSHU; 36°46'N, 139°51'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XXXIX-7). See Fig. 2A. Takahata-machi, Higashiokitama-gun (^^II?P FOODEN AND AIM!: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 177 SfflfflJ); Yamagata, HONSHU; ca. 38°00'N, 140°13'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE34. Takai, Takayama-mura, Kamitakai-gun (JiS^ ?PSUj;^S^); Nagano, HONSHU; 36°39'N, 138°2rE; collected 16 Jul. and 27 Oct. 1998 by M. Aimi; PRIKU, 4 (skeletons only). NE49. Takaike-mura, Higashimuro-gun (^^^-ftlPS/til ^ ); Wakayama, HONSHU; 33°32'N, 135°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-42). See Fig. 2A. Takakumadake, Kanoya-cho, Kimotsuki-gun (flf MiP^MBTTSP?!^); Kagoshima, KYUSHU; 31°33'N, 130°53'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-12). See Fig. 2 A. Takakumadake, Takakuma-mura, Kimotsuki-gun i^mmm^^m^my, Kagoshima, KYUSHU; 31°23'N, 130°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 12: 1-14). See Fig. 2A. Takakurayama, Ohsawa-mura, Miyagi-gun ("S"^ ^A)R^mt[Uy, Miyagi, HONSHU; 38°18'N, 140°45'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 55: XLIII-7). See Fig. 2A. Takamurayama, Nyuno-son, Toyota-gun (Wffl ?P A ^ W M UJ ); Hiroshima, HONSHU; 34°28'N, 132°5rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-20). See Fig. 2A. Takane, Asahi-mura, Iwafune-gun (^^p?P^ B W S ^ ); Niigata, HONSHU; 38°20'N, 139°37'E; collected 15 Oct. 1998 by M. Aimi; PRIKU, 2 (skeletons only). NE31. Takanosuyama, Kuba-son, Toyota-gun (sffl^ X^^SM^'M.Shy, Hiroshima, HONSHU; 34°34'N, 132°47'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV- 18). See Fig. 2A. Takanosuyama, Takeni-son, Toyota-gun (SB9?P 1tt^fl|y^Uj); Hiroshima, HONSHU; 34°3rN, 132°45'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV- 19). See Fig. 2A. Takaosan, Hachioji-city (Ai-^Tfii^Mllj); 7b- kyo Metropolis, HONSHU; 35°37'N, 139°13'E; provisioned population, translocated from Shodoshima in Sep. 1957; birth season, 1958-1966, reported by Kawai et al. (1967, pp. 37, 38, 47; cf. Study Group on the Present Status of Japanese Monkeys, 1977b, p. 23). NEUO. Takaosan vicinity, Chikano-mura, Nishi- muro-gun (ffi#8^]ifii?Wift Mill fife); Wa- kayama, HONSHU; 33°49'N, 135°37'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-18). SeeFig. 2A. Takaosan vicinty, Nagano-mura, Nishimuro-gun 0#«?PMi?WSmajffe); Wakayama, HONSHU; 33°47'N, 135°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX- 15). See Fig. 2A. Takara-mura, Minamitsuru-gun (l^fPli^^W); Yamanashi, HONSHU; 35°34'N, 138°51'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXIII-29). See Fig. 2A. Takarajou vicinity, Awa-mura, Ayama-gun (Pr[ \hmni^¥imB.^^y Mie, HONSHU; 34°45'N, 136°18'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII- 14). S_ee Fig. 2A. Takasakiyama, Oita-city (A^'^TfTi^lli^lil); Oita, KYUSHU; 33°15'N, 13r32'E; provisioning initiated Nov.-Dec. 1952 (Itani et al., 1964, p. 2). Birth season, 1958-1966, reported by Kawai et al. (1967, pp. 38-39). B virus survey conducted before 1965 by M. Endo et al. (1964, p. 173). Captured in Jan. 1968, died 15 Feb. 1968; JMC, 1 (skeleton only). Collected in 1976 by PRIKU staff; PRIKU, 2 (1 skeleton only, 1 mandible and postcranials only). Killed by dog in 1984; JMC, 1 (skull only). MtDNA samples collected ca. 1986-1991 by Hayasaka et al. (1991, p. 400). Collected before 1991 by K. Wada; PRIKU, 3 (skeletons only). Col- lected before 1991 by PRIKU staff; PRIKU, 5 (4 skeletons only, 1 skull only). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). Microsatellite DNA of laboratory specimen analyzed before 1996 by Domingo-Roura et al. (1997, p. 358). External measurements taken before 1997 by Hamada 178 FIELDIANA: ZOOLOGY et al. (1996a, pp. 98, 99). MtDNA samples col- lected before 2003 by Y. Kawamoto (2002, p. 60). SIV83. Takasugi vicinity, Ooyama-mura, Kaminii- kawa-gun (±lffJlliPAllj^St^^); Toyama, HONSHU; 36°30'N, 137°22'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 42: XXIX-4). See Fig. 2A. Takasuyama, Takasu-mura, Sakai-gun (^#?P li^^^tli^aj); FukuU HONSHU; 36°06'N, 136°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV- 18). See Fig. 2A. Takata-mura, Higashimuro-gun (!^#-ft^i^B3 ^ ); Wakayama, HONSHU; 33°44'N, 135°54'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 34: XX-47). See Fig. 2A. Takatoriyama, Minobu-mura, Minamikoma-gun (lt^S?P#EWJKlXaj); Yamanashi, HONSHU; 35°22'N, 138°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-4). See Fig. 2A . Takatoriyama National Forest, Takaichi-mura, Takaichi-gun (i^Tt]fPi^T^:ttS15ililSW^); Nara, HONSHU; 34°28'N, 135°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXI-13). See Fig. 2A. Takatoriyama National Forest, Takatori-cho, Takaichi-gun (if TfT^Pif IJ^ffllSUUjaW^); Nara, HONSHU; 34°25'N, 135°48'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35: XXI-12). See Fig. 2A. Takatsuki-cho, Ika-gun (##?PIf >^ BJ); Shiga, HONSHU; ca. 35°28'N, 136°15'E; mtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). SW6. Takayama, Kuchi-son, Asa-gun (^jilP^Xitfe^ If llj ); Hiroshima, HONSHU; 34°3rN, 132°24'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26:X1V-15). SeeFig. 2A. Takeno-cho, Kinosaki-gun ( ^ lli$ IP t^ i? ffll ); Hyogo HONSHU; ca. 35°35'N, 134°44'E; pro- visioned group, translocated from Kochi be- fore 1966; birth season, 1965-1966, reported by Kawai et al. (1967, pp. 38, 47). Collected ca. 27 Oct. 1971 by JMC staff; JMC, 1 (skele- ton only). SW49. Takidani vicinity, Ootaki-mura, Inukami-gun (^ ±fP;'^)i^il#fe); Shiga, HONSHU; 35°18'N, 135°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV-22). See Fig. 2A. Takigoshi, Outaki-mura, Kiso-gun (^KelPi/^ ^ )t ® ); Nagano, HONSHU; 35°49'N, 137°27'E; collected 4 and 6 Sep. 1998 by M. Aimi; PRIKU, 2 (skeletons only). NE84. Takinozawa, Seinaiji-mura, Shimoina-gun (T"^ M^^\^^¥SM(D)R); Nagano, HONSHU; 35°30'N, 137°40'E; collected 5 Dec. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE73. Takitani, Sugawa-mura, Kanoashi-gun (^JE?P ^JIW)t#); Shimane, HONSHU; 34°33'N, 131°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28:XVI-10). SeeFig. 2A. Takitani vicinity, Kuda-mura, Atago-gun (^S W>X^¥iM"^^); Kyoto, HONSHU; 35°15'N, 135°49'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 36:XXII-21). SeeFig. 2A. Takiyama, Kitayoshino-son, Katsuta-gun (Mffl iP db ^ i? ^ >t Ol ); Okayama, HONSHU; 35°06'N, 134°09'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV-21). See Fig. 2A. Takiyamagawa vicinity, Kake-cho, Yama- gata-gun {\hWkWM\^M\^]\\y^^y ^^ ffe); Hiroshima, HONSHU; 34°37'N, 132°21'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-7). See Fig. 2A. Takuwa, Mitoya-cho, lishi-gun (ISS5PHZJM fflJ^^^); Shimane, HONSHU; 35°15'N, 132°52'E; collected 12 Dec. 1994-27 Jul. 1997 by PRIKU staff; PRIKU, 8 (skeletons only, in- cluding 1 with fragmented skull and 1 with fragmented mandible). SW59. Tanegashima ( S •? S ); Kagoshima, TANEGASHIMA; ca. 30°36'N, 130°59'E; oc- currence in 1905 reported by local residents to M. P. Anderson (in Thomas, 1906 ["1905"], p. 362). Occurrence in 1940s reported by K. Ka- FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 179 wanaka (1973, p. 1 15). Extinction or near ex- tinction of island population reported by local residents in April 1970 (Azuma, 1972, p. 262). SW99. Taneeawayama vicinity, Funaki-mura, Nii-gun ( ^ M ?P ^p^ ^ ^ S ^ Jll UJ fte ); Ehime, SHIKOKU; 33°55'N, 133°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: IX- 16). See Fig. 2A. Taninoshiriyama vicinity, Kamiiritsu-mura, Mi- namiamabe-gun CMM^^JiX^M^^^^ Ojfe); Oira, KYUSHU; 32°52'N. 131°56'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: I1I-9). See Fig. 2A. Tano, Handa-mura, Kusu-gun (3E^S^lPtSffl^EB W)\ Oita, KYUSHU; 33°09'N, 131°15'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 16: III- 19). See Fig. 2A. Tanosawa vicinity, Uchigata-mura, Ki- tatsugaru-gun (db^^^lP 1*1 ^^ BB ^ >R fife); Aoniori, HONSHU; 41°03'N, 140°29'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-10). SeeFig. 2A. Tanzawayama, Kiyokawa-mura, Aikou-gun (S ¥?P>f JII^:l^>RLil); Kanagawa, HcJnSHU; ca. 35°28'N, 139°10'E; mtDNA samples col- lected before 1999 by Y. Kawamoto (1998, p. 54). NE109. Tara-dake, Fujitsu-gun (R^?P^MS); Saga, KYUSHU; ca. 32°59'N, 130°06'E; observed in 1940s by local residents (Sakura, 1976, p. 151); subsequently extinct at this locality (Ikeda and Eguchi, 1978, p. 59). 5PF57. Tarekaya National Forest, Yuki-cho, Jinseki-gun {^^W/^^^^ U:^i7SW^); Hiroshima, HONSHU; 34°47'N, 133°18'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-33). See Fig. 2A. Tarogawa, Kawasaki-machi, Shibata-gun (^ffl IP Jll *$ fflj ;!!; HP Jll ); Miyagi, HONSHU; 38°14'N, 140°35'E; observed Dec. 2002-Jan. 2003 by K. Izawa, T. Uno, and H. Fujita (2003, p. \^).NE25. Tarudaira vicinity, Horado-mura, Mugi-gun (^ 'B.^Mf^^^ )l^^)\ Gifii, HONSHU; 35°37'N, 136°5rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 41: XXVIII- 15). See Fig. 2A. Tanisawayama vicinity, Hikawa-mura, Nishi- tama-gun (^^MW^]\\^M)R\U^)\ Tokyo Metropolis, HONSHU; 35°47'N, 139°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXI V-3). See Fig. 2 A. Tashida vicinity, Hatta-mura, Inabe-gun (M#?P >^Ba^^^^Baftfe); Mie, HONSHU; 35°08'N, 136°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII-3). See Fig. 2A. Tashiro, Ootaki-machi, Isumi-gun (MP^?PA^ S BTT BQ ft ); Chiba, HONSHU; 35°13'N, 140°12'E; collected 16 Aug. 1997 by PRIKU staff; PRIKU, 1 (skeleton only). NE126. Tashiyama, Murou-mura, Uda-gun (^PS?P^^ ^ A 6f Ol ); Nara, HONSHU; 34°31'N, 136°03'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 35:XXI-10). SeeFig. 2A. Tasmania. See Launceston. Tatai, Horigane-mura, Minamiazumi-gun (]^$ #iP^^WBa^#); Nagano, HONSHU; 36°18'N, 137°50'E; collected 9 Nov. 1997 by M. Aimi; PRIKU, 1 (skeleton only). Collected 20 Nov. 1997 and 27 Nov. 1999 by PRIKU staff; PRIKU. 2 (skeletons only). NE93. Tataki vicinity, Tokuda-mura. Shusou-gun (j^lil IP ^> Ba ^ ea >i ffe ); EJume, SHIKOKU; 33°54'N. 133°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 18: IX-8). See Fig. 2A. Tatsufusayama vicinity, Higashimera-son, Koyu-gun (iPo^^^m^KM^HMLUffe); M- ya-aki, KYUSHU; 32°15'N, 13ri7'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-16). See Fig. 2A. Tatsuka\\ ayama, Sumino-mura, Nii-gun (^M?P ^^^^aUIIOj); Ehime, SHIKOKU; 33=52?^, 133°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: IX- 14). SeeFig. 2 A. Tatsuyama-mura, Iwatagun (#ffl?PilU4^); Shizuoka. HONSHU; ca. 34°52'N, 137°49'E; 180 HELDIANA: ZOOLOGY mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NE66. Tazawa, Takamori-machi, Shimoina-gun (T"^ IP ?P if ^ BTT Ba )R ); Nagano, HONSHU; 35°35'N, 137°51'E; collected 11 Dec. 1997 by PRIKU staff; PRIKU, 3 (skeletons only, in- cluding 1 with fragmented skull). Collected 3 Nov. 1998 by M. Aimi; PRIKU, 4 (skeletons only). NE76. Tedayama vicinity, Togo-son Higashiusuki-gun (maff^m^^^BgOlffe); Miyazakt, KYUSHU; 32°22'N, 131°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-20). See Fig. 2A. Tenjinbara, Takayama-mura, Kamitakai-gun (_h S#^ifaj;Ht^ttJI); Nagano, HONSHU; 36°41'N, 138°24'E; collected 11 Aug.-7 Sep. 1998 by M. Aimi; PRIKU, 5 (skeletons only). NE49. Tenjinkyo, Yoshii-cho, Shituki-gun {^HW^^ ffll^#ll!^); Okayama, HONSHU; 34°39'N, 133°25'E; mtDNA samples collected 1998-2000 by I. Yoshimi and H. Takasaki (2003, p. 71). 5^56. Tenjinyama, Yuno-son, Kawakami-gun (Jl|_h^ ^ IF W ^ ^ UJ ); Okayama, HONSHU; 34°52'N, 133°25'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV-4). See Fig. 2A. Tennoji ( ^ 3E ^ ); Osaka, HONSHU; ca. 34°40'N, 135°30'E; living captive purchased at market 10 Jun. 1826 by P. F. von Siebold (Holthuis & Sakai, 1970, p. 67); specimen possibly included among those sent by von Siebold to RMNH. Not mapped. Tenryu-city (^"^Tf]); Shizuoka, HONSHU; ca. 34°53'N, 137°49'E; blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6).yV£'66. Texas. See Arashiyama West. Tochiyama vicinity, Akaba-mura, Kitamuro-gun (db#«lP#^^^Ujfte); Mie, HONSHU; 34°14'N, 136°16'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-41). See Fig. 2A. Tochu vicinity, Hongawa-mura, Tosa-gun (zhfe ?P ^ Jll ^ F 4^ -te ); Kochi, SHIKOKU; 33°44'N, 133°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Mito, 1989, p. 25). See Fig. 2A. Todai. See Nagata-todai. Toei-cho, Kitashitara-gun (dblS^^P^^BO"); Aichi, HONSHU; ca. 35°02'N, 137°45'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NE67. Togakushiyama vicinity, Togakushi-mura, Kamiminochi-gun (±7K[*l?PFH:|4p|iiiJL| fife); Nagano, HONSHU; 36°43'N, 138°05'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 46: XXXI-25). See Fig. 2A. Toi-misaki, Kushima-city ( $ Pb^ TfJ |[P ^ ill? ) (= Cape Toi, Kushima); Miyazaki, KYUSHU; 31°22'N, 131°20'E; provisioned group; birth season,1966, reported by Kawai et al. (1967. pp. 37, 39). Collected 31 Dec. 1966 by JMC staff; JMC, 1 (skeleton only). SW98. Tojiroyama, Tomisato-mura, Nishiyatsu- shiro-gun OA-ftlPsM^^'ftUj); Yama- nashi, HONSHU; 35°26'N, 138°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 49: XXXIII-35). See Fig. 2A. Tokio. See Tokyo. Tokio Market. See Tokyo market. Tokiwa, Oomachi-city (AffllTfi^^); Nagano, HONSHU; 36°29'N, 137°50'E; collected 8 Apr. 1998 by M. Aimi; PRIKU, 1 (skeleton only). Collected 2 Nov. 1998 by PRIKU staff; PRIKU, 2 (skeletons only). NEW. Tokiyama, Toki-mura, Yourou-gun (ft^iPB^^ B^lll); Gifu, HONSHU; 35°14'N, 136°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 40: XXVIII-3). See Fig. 2A. Tokuji, Ichihara-city ( TtT i^ TfT ^> E ); Chiba, HONSHU; 35°18'N, 140°09'E; collected 5 Feb. and 30 Mar. 1999 by PRIKU staff; PRIKU, 2 (skeletons only). NE127. Tokusa, Hagiwara, Oofuji-mura, Higashiyama- nashi-gun (mOJ^^P AM^i^J^^M); Ya- manashi, HONSHU; 35°40'N, 138°49'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-22). See Fig. 2A. Tokuzen-nishi vicinity, Nishiiyayama-son, Mima-gun (M^l^Pem^Oj^^^Sffiftfe); FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 181 Tokushima, SHIKOKU; 33°54'N, 133°50'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-21). SeeFig. 2A. Tokyo (^^tP) (= Jedo; Tokio; Yedo); Tokyo Metropolis, HONSHU; 35°42'N. 139°46'E; obtained 1874-1876 by Mr. v. Hilgendorf (cf. Schweyer, 1909, p. 8), ZMB, 6 (1 skeleton only, 5 skulls only). Purchased 1900-1901 by K. A. Haberer (cf. Schweyer, 1909, p. 8); mu- seum unknown, 2 (skulls only, possibly in- cluded among unlocalized specimens exam- ined). Not mapped. Tokyo market. (M^f^^) (= Tokio Market); Tokyo Metropolis, HONSHU; 35°42'N, 139°46'E; purchased 31 Dec. 1882 and 1 1 Feb. 1883 by P. L. Jouy; USNM, 3 or 4 (2 male skulls, 1 female skull, and 1 female skin [mis- matched with male skull]). Not mapped. Tomikawayama, Ooishi-mura, Kurita-gun (^^ ?PA^^sJIIUJ); Shiga, HONSHU; 34°52'N, 135°53'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXIV-28). See Fig. 2A. Tomogashima, Wakayama-city ( ^ 1^ |1| rfi" S 'It My, Wakayama, TOMOGASHIMA; 34°17'N, 135°00'E; monkey park population translo- cated from Yakushima, opened in 1963, closed in 1972 (cf Shidei et al., 1981, p. 17). SWS2. Toragamine Mountain Range vicinity, Kurusu- kawa-mura, Nishimuro-gun (ffi#S^^ffiJI| ^M.T^\\\U^); Wakayama, HONSHU; 33°5rN, 135°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XX-16). See Fig. 2A. Torihama shell mound, Mikata-cho, Mikata-gun {^15^^15^ M'ikRW); Fukui, HONSHU; coordinates unknown; mid-Jomon subfossils (age, 5-6 Ka) reported by M. Kuroda (2002a, p. 118); Jakuo Historical and Ethnographic Museum, 2 (skulls only; not seen). Not mapped. Toriidoyama, Chikusa-mura, Mie-gun (HMIP^ SWBMpUj); Mie, HONSHU; 35°02'N, 136°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII-4). See Fig. 2A. Torishima, Kushima-city (^PalrfiBM); Miya- zah, TORISHIMA; 3r27'N, 131°24'E; a young male monkey swam ca. 600 m to this island in 1970 (Mito, 1980, p. 29). SW97. Totorogauchi, Kitagou-son, Higashiusuki-gun (mQttlPdb0W±RfflJ)$g|lt#); Ao- mori, HONSHU; 40°34'N, 140°09'E; mtDNA samples collected before 2003 by Y. Kawa- moto (2002, p. 60). NE9. Tsukagoshi, Ootaki-machi, Isumi-gun (MP^lPA ^M^MM); Chiba, HONSHU; 35°14'N, 140°09'E; collected 31 Jul. 1997-23 Nov. 1997 by PRIKU staff; PRIKU, 5 (skeletons only, in- cluding 1 with fragmented skull). NE126. Tsukinoya, Kisuki-cho, Ohara-gun (AJ^IP^/^ fflJ^yM); Shimane, HONSHU; 35°13'N, 132°56'E; collected 21 Nov. 1994 by PRIKU staff; PRIKU, 3 (skeletons only, including 1 with fragmented skull and 1 with mandible and postcranials only). SW59. Tsukiyama vicinity, Fujiwara-machi, Shioya-gun (^^^SJ^fflTv^Ujffe); Tochigi, HONSHU; 36°50'N, 139°39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XXXlX-5). See Fig. 2A. Tsukiyodaira, Takamori-machi, Shimoina-gun {T ^M^nm^ n^^y, Nagano, HONSHU; 35°34'N, 137°52'E; collected 13 Mar.^ Aug. 1998 by M. Aimi; PRIKU, 14 (skeletons only). NE76. Tsukiyonodan, Mutsuai-mura, Minamikoma-gun (lt&0?PBi-^^.^^IS:); Yamanashi, HONSHU; 35°14'N, 138°23'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-2). See Fig. 2A. Tsukizaki, Ichihara-city ( rfT J^Tfi ^ lli^); Chiba, HONSHU; 35°18'N, 140°08'E; collected 21 Jan.-30 Mar. 1998 by PRIKU staff; PRIKU, 6 (skeletons only). NE127. Tsukubasan, Tsukuba-machi, Tsukuba-gun (m^^ IP ^ }:^ fflj ^ /:^ U4 ); Ibaraki, HONSHU; 36°14'N, 140°08'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 51: XXXVIII-1). See Fig. 2A. Tsukui-machi, Tsukui-gun (y^^^^/$^#ffll); Kanagawa, HONSHU; ca. 35°31'N, 139°16'E; mtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33; 2002, p. 60). NE108. Tsunagi, Yamagata-mura, Kunohe-gun {%WW^ UU ff^ ^ 1^ ); Iwate, HONSHU; 40°10'N, 141°39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56:XLV-11). SeeFig. 2A. Tsuneroku vicinity, Tomiyama-mura, Hata-gun (<#^iPaUj^^7Nffe); Kochi, SHIKOKU; 33°07'N, 133°00'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: X-14). See Fig. 2 A. Tsunogawa-Akana vicinity, Mitani-mura, Taka- shima-gun (^ft?PH^;f^r^JI^^^=Sflfe); 5%fl, HONSHU; 35°25'N, 135°55'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV-7). See Fig. 2A. Tsunotani, Tsuga-mura, Oochi-gun (e^^PfPM ;1^ ^ ^ ); Shimane, HONSHU; 34°58'N, 132°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 28: XVI-25). See Fig. 2A. Tsuriganeyama, Nanaori-son, Nishiusuki-gun (® e^^-biff^t^Mlij); Miyazaki, KYUSHU; 32°4rN, 131°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-29). See Fig. 2A. Tsuro vicinity, Kaminada-mura, Hata-gun (<#^ FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 183 ?P ± ^ W )i S fte ); Kochi, SHIKOKU; 32°45'N, 133°0rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: X-7). See Fig. 2A. Tsutsumori, Ootaki-machi, Isumi-gun (MP^^A ^gffllM0); Chiba, HONSHU; 35°13'N, 140°09'E; collected 30 Jun. 1997 and 6 Aug. 1998 by PRIKU staff; PRIKU, 2 (skeletons only). NE126. Tsutsumori National Forest, Oikawa-mura, Isumi-gun (HP^i^^JII^M^aW^); Chiba, HONSHU; 35°11'N, 140°12'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 50: XXXVI-7). See Fig. 2A. Tuino. See Jinrio. Uchiyama, Yoshinaga-mura, Fuji-gun (gdr^'o ^Wl^lLiJ); Shizuoka, HONSHU; 35°14'N, 138°46'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX- 12). See Fig. 2 A. Ueda, Hasumi-mura, Oochi-gun (b^^^MH W ± Ba ); Shimane, HONSHU; 34°51'N, 132°41'E; collected 2 Jul. 1996-12 Nov. 1997 by PRIKU staff; PRIKU, 29 (skeletons only). SW62. Ui, Ishigaki-mura, Arida-gun (WBB^^ilW^ # ); Wakayama, HONSHU; 34°00'N, 135°20'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33:XX-2). SeeFig. 2A. Ujitahara-mura, Tsuzuki-gun (^§^?P^/p BBJ^ ^); Kyoto, HONSHU; 34°50'N, 135°52'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXII-29). See Fig. 2A. Umegatani vicinity, Miuchi-mura, Onsen-gun {iB^^^^P^^^T'^^); Ehime, SHIKOKU; 33°47'N, 132°57'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 21: IX-25). See Fig. 2A. Unazuki-machi, Shimoniikawa-gun (Tl!f JII?P^ ^^^fflj); Toyama, HONSHU; ca. 36°51'N, 137°34'E; mtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). NE53. Unsen. See Kyogatake. Urayama, Shino-mura, Minamikuwada-gun (]^ ^Ba^P^^ROl); Kyoto, HONSHU; 34°59'N, 135°38'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXlI-25). See Fig. 2A. Ushigadake, Ikazawa-mura, Minamiuonuma-gun (|lg^)S?PE + )R*t4^^); Niigata, HONSHU; 37°03'N, 138°57'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 47: XXXlI-5). See Fig. 2A. Ushimaki, Takamori-machi, Shimoina-gun (T# M^M^^ ^^); Nagano, HONSHU; 35°34'N, 137°51'E; collected 11 Jan. 1998 by M. Aimi; PRIKU, 1 (skeleton only). Collected 9 Mar.-8 Dec. 1998 by PRIKU staff; PRIKU, 27 (skeletons only, including 3 with frag- mented skulls). NE76. Ushiokuyama vicinity, Okunoda-mura, Higashi- yamanashi-gun (^Ul^^PEif BgW^HOj- w ); Yamanashi, HONSHU; 35°41'N, 138°49'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXlII-23). See Fig. 2A. Ushiroyama, Hasumi-mura, Oochigun (b^?P ^^^Wik\h); Shimane, HONSHU; 34°52'N, 132°38'E; collected 19 Dec. 1995-29 Jul. 1998 by PRIKU staff; PRIKU, 15 (skeletons only). Collected 11 Mar. 1999-14 Jul. 2000 by M. Aimi; PRIKU, 21 (skeletons only). SW62. Ushitakisan, Yamataki-mura, Senboku-gun {^ db^Uj>i^^)1iaj); Osaka, HONSHU; 34°24'N, 135°28'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 33: XIX-5). See Fig. 2A. Utou, Otsukirai-mura, Kesen-gun ('^'(llj^j^S 51^ ^ B 3l ); Iwate, HONSHU; 39°07'N, 141°50'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 56: XLV-4). See Fig. 2A. Utougi, Ookawauchi-mura, Abe-gun (^"f^^PA )Rlrt^W^7K); Shizuoka, HONSHU; 35°12'N, 138°22'E; reported in questionnaire survey conducted in 1 923 by K. Hasebe (Iwano, 1 974, p. 45:XXX-9). SeeFig. 2A. Utsutsumi, Kamogawa-city (^IJIlTtTflM); Chiba, HONSHU; 35°07'N, 140°04'E; collected 11 Nov. 1997-18 Dec. 1998 by PRIKU staff; PRIKU, 19 (skeletons only). NE125. Wachi-cho, Funai-gun (^p'^^^^fflJ); Kyoto, HONSHU; ca. 35°17'N, 135°25'E; captured ca. 16 Apr. 1986, died 10 Jun. 1986; JMC, 1 (skull 184 FIELDIANA: ZOOLOGY only). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). SW4L Wada-mura, Ooi-gun (AtS?P^B9^); Fukui, HONSHU; 35°29TSf, 135°35'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 39: XXV-5). See Fig. 2A. Wainai, Kariya-mura, Shimohei-gun (TPB'^fP XlJM^W^I^); Iwate, HONSHU; 39°41'N, 141°43'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 56:XLV-10). SeeFig. 2A. Wakahowatauchi, Nagano-city (:^i?rfT^#;^ rt); Nagano, HONSHU; 36°37'N, 138°16'E; collected 17 Nov. 1998 by PRIKU staff; PRIKU, 1 (skeleton only). NE50. Wakamiya-Hachimanjinja, Takayama-mura, Kamitakai-gun {hm^^m^\l\^^'U R^^ tt); Nagano, HONSHU; 36°41'N, 138°25'E collected 5 Feb. 2000 by M. Aimi; PRIKU, 1 (skeleton only). NE49. Wakasa-cho, Yazu-gun (AHiPS^fflT); Tottori, HONSHU; ca. 35°20'N, 134°24'E; captive group translocated from Wakasa to PRIKU, mtDNA samples collected ca. 1986-1991 by Hayasaka et al. (1991, p. 400). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). Microsatellite DNA ana- lyzed before 1996 by Domingo-Roura et al. (1997, p. 358). External measurements taken before 1997 by Hamada et al. (1996a, pp. 98, 99). MtDNA samples collected before 2003 by Y. Kawamoto (2002, p. 60). SW50. Wake vicinity, Kamikawa-mura, Minami- muro-gun (^#ftlP± Jl| W^-^ffe); Mie, HONSHU; 33°49'N, 135°55'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIII-55). See Fig. 2A. Wakigahata-mura, Inukami-gun (:^-b^flS '^'"llffl ^); Shiga, HONSHU; 35°13'N, 136°22'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 38: XXIV-21). SeeFig. 2A. Wakinosawa-mura vicinity, Shimokita-gun (Tdb ?P fiS i? )H W - w ); Aomori; HONSHU; 41°09'N, 140°46'E; collected 10 Oct. 1961 by PRIKU staff; PRIKU, 1 (skull only). Reported before 1970 by C. B. Koford (1969, p. 12; cf. Azuma, 1985, p. 3; Fooden & Aimi, 2003, p. 111). Blood samples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). NE5. Warabitaira, Takayama-mura, Kamitakai-gun (Jb i^#iPi^Uj;^M¥); Nagano, HONSHU; 36°41'N, 138°24'E; collected 3 Jan. 2000 by M. Aimi; PRIKU, 1 (skeleton only). NE49. Wasedasuirin vicinity. Shionomachi-mura, Iwa- fune-gun {^li^^^m^^^mm^^^; Niigata, HONSHU; 38°25'N, 139°34'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXII-24). See Fig. 2A. Wataka vicinity, Owase-cho, Kitamuro-gun (db #«lPMiKfflI}lgEffe); Mie, HONSHU; 34°04'N, 136°I1'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 37: XXIII-46). See Fig. 2A. Watarai-gun (Jg^lP); Mie, HONSHU; ca. 34°20'N, 136°30'E; acquired before 1939 by N. Kuroda (1938, p. 112; 1940, pp. 270, 271); formerly in private collection of N. Kuroda, destroyed by fire in 1945 (Austin et al., 1948, p. 4), 1 (not seen). Not mapped (see Oo- miya-cho). Yabitsuyama National Forest, Kanita-mura, Hi- gashitsugaru-gun (^/$^IPMffl^^ffllll@ W^); Aomori, HONSHU; 41°03'N, 140°39'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-16). SeeFig. 2A. Yaheishiroburaku, Okugawa-mura, Yama-gun (IP0?PIIJI|^?^^¥Eg^P^^); Fukushima, HONSHU; 37°43'N, 139°41'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 52: XLI-6). See Fig. 2A. Yaichirousawa vicinity, Oohata-mura, Shi- mokita-gun (T:lblPA;lffl^?^^-fiP)Rflb); Aomori, HONSHU; 41°24'N, 141°08'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-19). SeeFig. 2A. Yakeyama, Azusagawa-mura, Minamiazumi-gun (lt^«?P^JI|^:^aj); Nagano, HONSHU; 36°11'N, 137°49'E; collected 7-10 Nov. 1998 by PRIKU staff; PRIKU, 5 (skeletons only, with fragmented skulls). NE94. FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 185 Yakeyama vicinity, Umegashima-mura, Abe-gun ( ^ ^ ?P ^S 2.3 Ka) reported before 2003 by M. Ku- roda (2002a, p. 115); Kyoto University An- thropology Laboratory, 1 (mandible only; not seen). Not mapped. Yotsugadake vicinity, Aiuchi-mura, Ki- tatsugaru-gun (db5*g?P*l rt^EB 7 Sftfe); Aomori, HONSHU; 41°03'N, 140°22'E; re- ported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 57: XLVI-11). SeeFig.2A. Youkurayama, Funaki-son, Toyota-gun (SBB^ ^r' ^ ^ ffl ^ UJ ); Hiroshima, HONSHU; 34°28'N, 132°59'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-23). See Fig. 2A. Youkurayama, Kamikitakata-son, Toyota-gun (^m^±ity5^mt\h); Hiroshima, HONSHU; 34°26'N, 132°58'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-22). See Fig. 2A. Youkurayama, Ookawa-son, Toyota-gun (WBB ^AM^MtiU); Hiroshima, HONSHU; co- ordinates unknown; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-24). Not mapped. Youkurayama, Zen-nyuji-son, Toyota-gun (MB9 FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 189 ?P#A#^ffl^UL|); Hiroshima, HONSHU; 34°26'N, 132°56'E; reported in questionnaire sur\'ey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 26: XIV-21). See Fig. 2A. Youzenotaki, Fuka-son, Kawakami-gun (JI|_hlP S 1^ ^ ffl ^ y >i ); Okayama^ HONSHU: 34°46'N. 133°29'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 27: XV- 1). See Fig. 2 A. Yudomari, Yaku-cho, Kumage-gun (^^IPM^ BI ^ )S ); Kagoshima, " YAKUSHIMA: 30°15'N,130°29'E; collected 25 Feb. 1989 by PRIKU staff; PRIKU, 18 (skeletons only, in- cluding 1 postcranials only). SW102. Yugawara-machi, Ashigarashimo-gun (S:ffiT^ ^ )RI J^ fflj ); Kanagmva, HONSHU; ca. 35°09'N, 139°04'E; provisioning initiated in 1956 (Study Group on the Present Status of Japanese Monkeys, 1977b, p. 23). Blood sam- ples collected before 1992 by Nozawa et al. (1991, p. 414; 1996, p. 6). MtDNA samples collected before 1998 by Y. Kawamoto (1997, p. 33). NEWS. Yukihikoyama, Kaya-mura, Shikama-gun {^M iPE^^l^Ul); Hyogo, HONSHU; 34°58'N, 134°4rE; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVIII- 13). See Fig. 2 A. Yumigiyama, Shiiba-son, Nishiusuki-gun (®Q tfiP^^^^TKlij); Miyazaki, KYUSHU; 32°29'N, nriO'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 15: 11-32). See Fig. 2 A. Yumoto, Yubara-cho, Maniwa-gun (KfilPi^J^ fflJ^^^); Okayama, HONSHU; 35°13'N, 133°45'E; reported in questionnaire sur\ey conducted in 1923 by K. Hasebe (Iwano, 1974. p. 27:XV-14). SeeFig. 2A. Yunodake, Oohira, Ryogauchi-mura, Ihara-gun (^iilPia^)Rlrt^A¥^i?^); Shizuoka, HONSHU; 35°09'N, 138°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 45: XXX- 10). See Fig. 2A. Yunoyama, Komono-cho, Mie-gun {E-M-WM.^ fflj ^^ y OJ ); Mie, HONSHU; 35°01'N, 136°27'E; reported present before 1939 by N. Kuroda(1938, p. 112; 1940, p. 270). SPF/^. Yunoyama, Komono-mura, Mie-gun (Hj i? ^ ^^ y OJ ); Me, HONSHU; 35°01'N, 136°27'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 36: XXIII-5). See Fig. 2A. Yuumayama, Kawata-son, Oe-gun (^^^JHffl ^ i^ ^ U \h ): Tokushima, SHIKOKU; 34°02'N, 134°14'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 23: XI-26). See Fig. 2 A. Yuzuruhayama, Nada-mura, Mihara-gun (HJ^ iPH^MS^LiJ); Hyogo, AWA^JISHIMA; 34°14'N, 134°49'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 30: XVlII-1). See Fig. 2A. Zarugadake vicinity, Suzurishima-mura, Mi- namikoma-gun {'h^W-^WM'¥i^T ^^); Yamanashi. HONSHU; 35°22'N, 138°19'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano, 1974, p. 48: XXXIII-6). See Fig. 2A. Zoukiyama, Higashikawa-mura, Higashikan- bara-gun (^^J^IP^JH^MtKUj); Niigata, HONSHU; 37°35'N. 139°26'E; reported in questionnaire survey conducted in 1923 by K. Hasebe (Iwano. 1974, p. 47: XXXII- 17). See Fig. 2A. 190 FIELD! AN A: ZOOLOGY Appendix 5. Local Variation in Sitting Height (mm) in Macaca fuscata Adults on Honshu (cf. Fig. 5)' Lati- tude Longitude (E)- ' (N)- 132° 133' J 137° 138° 139° 140° Adult females 38° 550.0 37° 550-550 (2) 36° 573.0 ± 42.00 557.6 ± 34.81 35° 551.3 ± 50.33 525-603 (J) 510-608(7) 564.5 ± 22.61 558.7 ± 20.65 521.0 528.0 ± 27.48 34° 515-625(4) 547.5 ± 29.68 497-590(79)-' 542.0 ± 18.81 523-604 (i7) 535-573 (J) Adult males 512-530(2) 487-545 (4) 38° 594 37° (/) 36° 603.6 ± 32.53 612.5 ± 29.02 35° 615 552-630 (5) 553-648 (S) 623.2 ± 25.66 623 565 581.0 ± 17.67 34° (/) 577.8 ± 40.23 504-625 (ISf 584.9 ± (7)' 39.91 570-682 {22) (/) U) 551-614(70) ' Data derived from specimens examined, except as otherwise indicated in footnotes 5 and 6. Within each cell, figures indicate mean ± SD (where n > 2). extremes, and sample size (italicized, in parentheses). - Column and row headings indicate lower limits of one-degree latitude-longitude blocks. ^ No data are available for specimens originating in the longitudinal zone 134-136.99°E. ^ Iwamoto (1971, table opp. p. 156) has published the following data for two additional localities situated within this latitude-longitude block: Kochi (provisioned group), 530.0 ± 28.04 mm, n = 72; Hasumi (nonprovisioned group), 541.4 ± 13.39 mm, n = 5. '' Data from Iwamoto (1971, table opp. p. 156); extreme values not published. '' Includes one adult male (sitting height = 595 mm) measured at Kochi (38°28'N, 132°53'E; provisioned group) by Iwamoto (1971, table opp. p. 156). FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 191 Appendix 6. Local Variation in Anterior Trunk Length (mm) in Macaca fuscata Adults on Honshu (cf. Fig. 6)' Lati- Longitude (E)- tude (N)- 132° 133° 134° 135° Adult females 36° 35° 381.2 ± 23.19 363-420 (5) 361.4 ± 12.26(7) 358.5 ± 16.69(27)^ 371.8 ± 19.53(47)-^ 34° 373.4 ± 26.13 332-415 {14) 364.0 ± 11.24(6)^ 372.0 ± 21.15(46)-' Adult males 36° 35° 385 (/) 397.6 ± 12.01 {Sy 411.2 ± 23.88(77)-^ 34° 364.4 ± 37.25 315-435(72) 394.5 ± 20.12(6)^ 399.1 ± 16.68(22)' ' Within each cell, figures above the dashed line indicate data derived from specimens examined — mean ± SD (where n > 2), extremes, and sample size (italicized, in parentheses): figures below the dashed line indicate data published by Hamada et al. (1996a, pp. 101. 102) — mean ± SD and sample size. To conserve space, data for three localities on Honshu north of 37°N are not included in the body of this table; these data are as follows: Takane, 38°20'N, 139°37'E, female, anterior trunk length = 371 mm, n = 1; Komogawa, 38°18'N, 139°36'E, female, anterior trunk length = 371 mm, n = 1, male, anterior trunk length = 399 mm, n = 1: Shimokita, 41°08'N, 140°50'E, females, 367.9 ± 22.47 mm. n = 10 (Hamada et al., 1996a. p. 102). - Column and row headings indicate lower limits of one-degree latitude-longitude blocks. "* Captive and/or provisioned. •» Mihara-city, 34°24'N, 133°05'E. ■'' Gagyusan. 34°49'N, 133°37'E (translocated to Nagatoro). * References: Aizawa and Hagiwara (2001, pp. 5-6); M. Hagiwara (pers. comm., 3 Oct. 2002); these measurements were originally reported as head and body length. 192 FIELDIANA: ZOOLOGY Appendix 6. Extended. Longitude (E)^ 136° 137° 138° 139° 140° Adult females 396.0 ± 53.14 369.6 ± 26.94 374.4 ± 11.19 344-470(4) 338-414(7) 355-392(77) 394.4 ± 13.42(77) 391.3 ± 20.30(42) 371.4 ± 12.07(76) 384.1 ± 20.34 380.7 ±23.16 347.5 358.6 ± 13.26 329-428 (J7) 355-400 (J) 335-360(2) 329-371(9) 369.9 ± 21.24(75)3 346.9 ± 13.82 (72)^ 374.6 ± 17.76(57) Adult males 400.4 ± 25.53 415.4 ± 21.66 406.0 ± 4.36 371-434(5) 383-446 (9) 403-411 (3) 413.6 ± 20.44(29) 403.3 ± 18.42(70) 411.5 ± 18.17 401 380 394.7 ± 23.23 375-441 (22) (1) (1) 365-450 (13) 404.2 ± 13.44(74)3 402.0(2)" 401.1 ± 19 A9 (34) FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 193 DXl ^ a o. < s + 1^ (N (N VD — a .JL, U ri J 5 w +1 ON r<-i <5 o <^ 00 O + 1 q ON ^ — + 1 ^^ r- ON o ON + i r- o o o VO IT) +1^ C^, NO + 1 o 00 in 00 ON Csl d o\ (N — (N On (N ^ NO d (N +1^ 00 r^ + 1^ t^ ON NO — o^ — 00 u. E II "S'S E^ y s •0 rt J. ij^ 00 ^"^^^ ^go =.' 3 — r^ • - 60 1 So ^ « — p^ 0 so 0 rv ca 00 <-l each 6a, p : Tak .6%; s. a ^^ "-N C ON g NO ^ ^ r- E ^ 5 3 — ^ ^-' t^ : mean ± SD (where n > ures below the dashed li of 37°N are not included gawa, 38°18'N, 139°36'E 19.4% (Hamada et al.. 1 id row headings indicate d/or provisioned. y. 34°24'N, 133°05'E. 34°49'N. 133°37'E(trans Aizawa and Hagiwara. 2 n, anterior trunk length n 3.0 ± 7.95 1.0% 9.2 ± 10.4 4.8% 00 (N Ov (N 0^ +1^ t^^E" --Si 8^ 't 't ' Tail len; examined; Honshu no 22.1%; Ko (5), T/ATL ~ Column 3 Captive '' Mihara- ' Gagyus, ^ Referen ^ Watarai 00 (N 01 n FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 195 ^ a ^ (N +1 + 1 m ^ 00 ^ 00 o '^ 00 '^ d 2; — ^ '^ 't +1 00 ^ J -^«^ "a \D ^ 00 f~^ B vO 0 (N — -* — -^ a Sm ^ 3 ■o ^ < ^ ^ +1 VO On ro + 1 + 1 + 1 o6^ in (N — '^ ■^ r^ 00 00 +1 CM ^. n — ^ — ^ '^ a ^ -rt .S 1) "ay 1/; O C Q a +IZ c^ C3 <^ E o .S -c 1) O aS "'^ c ^ o .5 _3 :^ ^■■3 1 u (u c o c o c o ^ ■ .5 rt r^ ■ o y - ;*: > ^ > fcH (U U So , ° 8b ^ o ^ 8^ . " .2 So . -i ^S; "^ i Q. C C Ov ■•- > . i-H 2i o w c = ■ Co S c ^ ,0 (U 2 -S -^ M 2 c Z .2 D-x: eoi^i: UO 196 FIELDIANA: ZOOLOGY o OS So s *o3d e o ^ rn in FN N-l o +'7 ON 00 +'7 tN in ON >o (N ^ + 1 00 OS 00 vd ^ in o cr 00 in 3- + 1 +1 oo (N r- p a\ r-' r^ r<-i >£i M- vO O ^^ +1+1- a> (N — — do o +1 -: in »n 00 »n d ON O OS OS vd +'7 R Os"^ 00 ^-^ ^^ -- in so +1 -r +'7 -? r^ in no 5- ; -= o Tt f- 1- O "^ 0 S = ^ : « =^o 1 c ^ « • t •* i- ' -^ Cl.^ uuS z « o a; FOODEN AND AIMI: SYSTEMATIC REVIEW OF JAPANESE MACAQUES 197 a IT) ON rn O + 1- >n ON CT — O CN — IT) + 1- oi >n +'7 (N r<^ ori„ d d (N — R 00 ro ^ +1 -r +1 — r^ r<^ oT rn vd S" m r^i Zl. r<~, CN c; — r~; O 00 — r^, O E^ 3 =3 •7 ?= ^ o o Z X f; E "J E f S3 J= CU NO ^ OJ ON 2 00 c P — CO ■" o