m^ ■ 1 1 ^^^^^^ ^ FI N . S . fV9.98 Biology -»=L- ....,..,:.-.:.. " NF^\ NO. 98 Systematic Review of the Taiwanese Macaque, Macaca cyclopis Swinhoe, 1863 Jack Fooden Hai-Yin Wu November 30, 2001 Publication 1515 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY {JOB for rorstribiitors to Fieldmna iiacn!)fi uicifoi. raymc;ii oi at least 50% ol the publication time. Contributions from staff, diess of ability to pay page charges. ;nuscripts. Three complete copies of iiiouid be submitted (one original copy pUis two '-,]] be considered for publication or submitted to ntific Editor. Field Museum of Natural nistorv, rhica-o •uter using MS-DOS, also submit text on 5V4-inch ' " ' "■' '"^' "^^amna, Micro.soft Word, \'olks- ,o .iubmii a " fable of Contents," a "List ol' 0. In most cases, the text should be preceded il any) and "Literature Cited." arc not used after abbreviated measurements). The { ; .vl; c'l.) rv The lJni»c;rsiiv lU DC l^lVCil 111 Ivill. \\ llfu: di-'i>ie\ ilU.^>i!^ iin: '1 follow BotanicO'Peyiodiciini-liuntianum and il papers) or Serial Sources for .cs of botanical authors should '84 edition, or 77.-2, V Press, Stanford, Cahf , 943 pp. on of montane and lowland rain '^^i-r,' nristics. Journal of Ecology, 51: 567-601. n;it1crns in -virions, pp. 63-80. In Browiv • hers. The Hagn. ._^ ard, J. H.. ed., \ •■ -uth •zations. Builetni 143, Bureau of American Ethnology, atemala. Part fL Polypodiaceae. Fieldiana: Botany. ^ ') the text (not as "plates") Figures must be accompanied its in figure captions alone, such as "XO.8," are not . jtively. See receitt issues oi Fieldiana for details of rnc. figure nutnbert's), and "top." _, i 1 inches (22 X' 28 cm) and may not exceed i 1 Vi >n boards in the arrangement to be obtained in the ) the printer as follows: Pen and ink drawings ■0 originals, but within the size limitation; and ■ mis. (Jriginal illustrations will be returned to the cor, ■- color reproduction must make prior tie corresponding author will normally rece , .• - can be made ar.. answered. Only oe made on the of page proofs. .'enerated cv fage proofs can quirements Of ANSI/NISO Z39.48-1992 (Permanence of Paper). FIELDIANA Zoology NEW SERIES, NO. 98 Systematic Review of the Taiwanese Macaque, Macaca cyclopis Swinhoe, 1863 Jack Fooden Adjunct Curator Division of Mammals Department of Zoology Field Museum of Natural History 1400 South Lake Shore Drive Chicago, Illinois 60605-2496 U.S.A. Hai-Yin Wu Assistant Professor HdJMARyQFTHE Institute of Natural Resources '"^ UWVWf VJT !<«> National Dong Hwa University Hualien, Taiwan J^U ? 0 2002 R.O.C. 974 Accepted February 16, 2001 Published November 30, 2001 Publication 1515 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 2001 Field Museum of Natural History ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents Abstract 1 Introduction 1 Geographic Distribution; Island-wide Population Estimate 2 Pelage 2 General Characterization 2 Age and Sex Variation 2 Seasonal and Geographic Variation 3 Albinism 3 External Measurements and Proportions 3 Cranial Characters 3 Molecular B iolog y and Genetics 9 Mitochondrial DNA 9 Nuclear DNA 12 Blood Proteins 14 Karyology 15 Parasites 17 Protistans 17 Flagellates 17 Amebas 17 Malaria 17 Ciliate 18 Trematodes 18 Cestodes 19 Nematodes 19 Leech 20 Lice 20 Viruses 20 B Virus 20 Reovirus 20 Simian Foamy Virus 20 Simian Virus 40 21 Natural History 21 Habitats 21 Terrestriality/Arboreality 22 Group Size and Composition 22 Home Range Area and Population Density ... 23 Diet 23 Predators 26 Intergroup Dispersal 26 Reproduction 26 Seasonality 26 Sexual Skin 26 Polythelia 28 Menstrual Cycles 28 Sexual Maturation 29 Copulation 29 Gestation and Parturition 29 Reproductive Rate 29 Neonatal Sex Ratio, Twinning, Birth Weight 30 Infant Mortality 30 Nursing; Longevity 31 Population Growth Rate 31 Fossils 31 Taxonomy 32 Synonymy 32 Type Series 32 Type Locality 33 Remarks 34 Evolution and Dispersal 36 Acknowledgments 38 Literature Cited 38 Note Added in Proof 45 Appendix 1 : Specimens Examined 46 Appendix 2: Gazetteer of Macaca cyclopis Localities 47 List of Illustrations 1 . Locality records of Macaca cyclopis 4-5 2. External characters in Macaca cyclopis 10 3. Tail length vs. head and body length in Macaca cyclopis 12 4. Skull of adult female Macaca c>'c/o;j/5 14 5. Skull of adult male Macaca cyclopis 15 6. Map of southwestern Taiwan showing col- lecting localities of fossil monkey teeth 34 7. Fossil monkey teeth collected in southwest- em Taiwan 35 List of Tables 1. External measurements and proportions in Macaca cyclopis 11 2. Mean birth weight reported in laboratory- born Macaca cyclopis 12 3. Cranial measurements and proportions in Macaca cyclopis 13 4. Local variation in greatest length of skull in wild-collected Macaca cjc/op/5 16 5. Sister groups and divergence ages of Ma- caca cyclopis as determined in five stud- ies of mtDNA in fascicularis-group spe- cies 16 6. Allele frequencies at blood-protein loci in samples of Macaca cyclopis 17 m 7. Incidence of trypanosome infections (Trypanosoma cf. conorhini) detected in samples of natural populations of Macaca cyclopis 17 8. Incidence of ameba species detected in unlocalized fecal samples of Macaca cy- clopis 18 9. Incidence of natural malarial infections detected in surveys of Macaca cyclopis populations 18 10. Incidence of trematode parasite infections detected in Macaca cyclopis samples 19 1 1 . Incidence of cestode parasite infections detected in fecal samples of Macaca cy- clopis 19 12. Incidence of nematode parasite infections detected in samples of natural populations of Macaca cyclopis 20 13. Incidence of anti-B virus antibodies de- tected in /aidcM/am-group macaques 21 14. Incidence of simian foamy virus, type 1 and type 2, detected in samples of natural populations of Macaca cyclopis 21 15. Incidence of simian virus 40 infections detected in Macaca cyclopis 21 16. Elevational distribution of Macaca cyclo- pis locality records 22 17. Geographic variation in group size in Ma- caca cyclopis populations 23 18. Ratio of sexually mature males to sexual- ly mature females reported in Macaca cy- clopis groups 24 19. Home range area and population density reported in Macaca cyclopis 24 20. Foods reported eaten by Macaca cyclopis .. 25 21. Observations concerning reproductive sea- sonality in Macaca cyclopis 27 22. Menstrual cycle duration in captive Maca- ca cyclopis 28 23. Annual reproductive rate in natural popu- lations of Macaca cyclopis 30 24. Neonatal sex ratio in natural populations of Macaca cyclopis 31 25. Taiwanese fossil monkey teeth: measure- ments and collection information 33 26. Dental measurements in wild-collected specimens of Macaca cyclopis 36 IV Systematic Review of the Taiwanese Macaque, Macaca cyclopis Swinhoe, 1863 Jack Fooden and Hai-Yin Wu Abstract The Taiwanese macaque, Macaca cyclopis Swinhoe, 1863 ("1862"), is systematically re- viewed, based on examination of 237 museum specimens, survey of relevant literature, and observation of natural populations. This review includes analyses of pelage characters, external measurements and proportions, cranial characters, molecular biology and genetics, and para- sites. Information also is presented concerning natural history, reproduction, fossils, and tax- onomic history. A hypothetical interpretation of the evolutionary history of this species is proposed. In an appendix, an annotated gazetteer lists 429 localities at which M. cyclopis has been collected or observed. Introduction Macaca cyclopis is endemic to Taiwan and is the only species of nonhuman primate native to this subtropical island, which is situated 140 km off the coast of mainland China. The existence of monkeys on Taiwan became known to Western science as a result of a visit there in 1715 by the Jesuit missionary Pere J.-A.-M. de Moyria de Mailla (Du Halde, 1735, pp. xxxiv, 162; Camp- bell, 1903, pp. 504, 506). The first scientific spec- imens of M. cyclopis were collected ca. 1 862 by R. Swinhoe (1863, p. 350), author of the technical name of the species, who was then the British vice-consul assigned to the island. Relatively few museum specimens have subsequently been col- lected from natural populations (Appendix 1), and, probably as a consequence, relatively few systematic studies of this species have been pub- lished (see Synonymy, below). The present review of M. cyclopis is based on the study of 237 museum specimens (62 of which were collected at known localities), survey of rel- evant literature, and observation of natural popu- lations. Specimens examined are preserved in in- stitutions listed below, which hereafter are cited by means of the indicated abbreviations; the num- ber of M. cyclopis specimens in each institution is indicated by a parenthetical notation. AMNH American Museum of Natural History, New York (3) ANSP Academy of Natural Sciences, Philadel- phia (1) bm(nh) British Museum (Natural History), Lon- don (3) JMC Japan Monkey Centre, Inuyama, Japan (45) KUPRi Kyoto University Primate Research In- stitute, Inuyama, Japan (53) Mcz Museum of Comparative Zoology, Har- vard University, Cambridge, Massachu- setts (1) NMNS National Museum of Natural Science, Taichung, Taiwan (16) NTUZ National Taiwan University, Depart- ment of Zoology. Taipei (7) SMTD Staatliches Museum fiir Tierkunde, Dresden (1) TESRi Taiwan Endemic Species Research In- stitute, Chichi, Taiwan (14) TM Taiwan Museum, Taipei (12) UMMZ Museum of Zoology, University of Michigan, Ann Arbor (2) USNM National Museum of Natural History, Washington, D.C. (41) Yio Yamashina Institute for Ornithology, Abiko, Japan (2) FIELDIANA: ZOOLOGY, N.S., NO. 98, NOVEMBER 30, 2001, PP. 1-70 ZMB Zoologisches Museum des Humboldt- Universitiit. Berlin (25) — Private collections (11) Geographic Distribution, Island-wide Population Estimate Locality records of M. cyclopis are widely dis- tributed in the mountainous eastern two-thirds of Taiwan Island (Fig. 1): questionnaire and field surveys indicate that this species currently inhab- its 111 of 270 townships (41.17f) in Taiwan (Hai- Yin Wu. Ling-Ling Lee. and Shih-Wei Chang, 2001. unpublished report). Prior to intensive hu- man settlement and agricultural exploitation of the western lowlands, macaques probably also were fairly common there, judging from Pere de Mail- la's observations in western Taiwan in 1715 (Du Halde. 1735. pp. xxxiv. 162). Local disappearance of macaques in western Taiwan was noted in 1 862 by Swinhoe (1863. p. 351). The natural geograph- ic distribution of M. cyclopis apparently does not extend to any of the small islands that surround Taiwan Island (Fig. 1). Introduced feral populations of M. cyclopis have become well established at four localities in Japan: (1) Oshima, a small island south of Tokyo, M. cyclopis population introduced sometime be- tween 1942 and 1953: (2) Nojima, a small island south of Nagoya, population introduced in 1958: (3) Wakayama Prefecture, south of Osaka, popu- lation introduced ca. 1960: and (4) Shimokita Hanto (= Peninsula), at the northern tip of Hon- shu (near the northern limit of distribution of M. fuscata), population introduced in 1971. For fur- ther details concerning these feral populations, see Appendix 2: Gazetteer. Based on an ongoing survey conducted since 1996 by Ling-Ling Lee. Shih-Wei Chang. G. Agoramoorthy. and Hai-Yin Wu. the total extant population of M. cyclopis in Taiwan has been es- timated to be ca. 250.000 ± 100.000 individuals in ca. 10,000 ± 5.000 groups. ivaceous) to golden brown; more caudally, on the lumbosacral region, the color gradually brightens to a richer golden brown. In individual hairs on the dorsal thoracic region, the basal two-thirds is grayish brown and the distal one-third is conspic- uously marked by 1-3 bands of pale golden (cf. Inagaki. 1996. p. 91). On the lumbosacral region, the base of individual hairs is paler than in hairs on the dorsal thoracic region, and the distal bands are more erythristic. The crown generally is somewhat paler than the dorsal thoracic region, but it is not as brightly colored as the lumbosacral region. The skin of the thinly haired face is pink- ish to reddish. Surrounding the face, on the fore- head and cheeks, is an indistinct fringe of some- what elongated blackish hairs. The lateral facial crest, which is pale grayish brown, is restricted to the posterior mandibular region (infrazygomatic crest; Fooden, 1995, p. 19). The proximodorsal surface of the limbs is approximately the same color as the adjacent dorsal surface of the trunk; more distally, the dorsal surface of the limbs grad- ually becomes somewhat more grayish. The prox- imal one-third of the dorsal surface of the tail is approximately the same color as the lumbosacral region; the distal two-thirds of the dorsal surface of the tail is dark grayish brown to clearly defined blackish. The ventral surface of the trunk, limbs, and tail is pale grayish. For a discussion of sexual skin coloration, see Reproduction below. Dorsal pelage color in M. cyclopis is generally similar to that in many specimens of M. fascicu- laris (Fooden, 1995. p. 3), but the color contrast between the dorsal thoracic region and the lum- bosacral region is greater in M. cyclopis. Dorsal pelage color in M. cyclopis is generally darker than that in M. mulatta (Fooden, 2000, p. 7), and the color contrast between the dorsal thoracic re- gion and the lumbosacral region is less in M. cy- clopis. Hair stream patterns on the trunk and extremi- ties of M. cyclopis have been studied in detail by Yasugawa (1960, p. 330). Age and Sex Variation Pelage General Characterization (Fig. 2) The dorsal surface of the thoracic region in postinfantile M. cyclopis is yellowish brown (ol- Neonatal pelage is dark gray to blackish (Hor- ikawa, 1932. p. 69; Kuroda, 1940, p. 269; Hsu. 1990. p. 66). In older infants (usnm 294177, Ml not erupted, age ca. 9 months), pelage color is approximately the same as that in adults, but hairs are shorter and finer than in adults. Pelage color is generally similar in females and males, but the FIELDIANA: ZOOLOGY dark dorsal surface of the distal two-thirds of the tail is more blackish in fully adult males than in fully adult females (H.-Y. Wu, pers. obs.)- Seasonal and Geographic Variation Although evidence of molting was not detected in M. cyclopis skins examined, Namiye (1914, p. 214) has reported that winter pelage is darker than summer pelage. Based on observation at several localities in Taiwan, Jui-Hua Chu (ntuz; pers. comm., 16 Apr. 2000) confirms that annual molt- ing occurs in M. cyclopis during the period May- August. The available sample of wild-collected skins is insufficient to determine whether or not pelage color varies geographically in M. cyclopis. Albinism Two or three albino captives of M. cyclopis re- portedly originated in Hualien Hsien during the period 1978-1980 (Poirier & Davidson, 1979, pp. 141, 142; Anonymous, 1981, p. 47), and an albino juvenile was captured by local residents at Wu- shihkeng, Taichung Hsien, in the early 1990s (photograph in possession of H.-Y. Wu). Aber- rantly pale specimens of M. cyclopis previously had been reported elsewhere in Taiwan by Kuroda (1940, p. 269) and Jones (1973, p. 370). Jachowski (1968, p. 1 1) asserted that body size is greater in northern M. cyclopis than in southern M. cyclopis, no external measurement data are available to test this statement (cf. Cranial Char- acters below). A brief summary of crown-rump length measurements in 116 wild-collected but unlocalized specimens has been reported by Satoh et al. (1956, p. 42; cf. Setoguti & Sakuma, 1959, p. 158; Lin, 1976, p. 75). Relative tail length (T/HB) apparently is similar in adult females and males, varying from approx- imately 0.65 to 0.95 (Fig. 3). In pooled samples (Table 1), mean relative tail length is 0.82 ± 0.058 (SD) in 24 adult females and 0.85 ± 0.089 in 8 adult males. This ratio apparently remains rela- tively constant from late fetal life to adulthood. The hind foot apparently grows more slowly than the head and body from infancy to adulthood (Table 1). The same probably also is true of the ear, but data are ambiguous on this point. Body weight (Table 1) is 8.12 kg in one wild- collected adult male (cf. Setoguti & Sakuma, 1959, p. 158) and varies from 6.30 kg to 18.50 kg in nine captive adult males (cf. Lin, 1976, p. 75; Poirier & Davidson, 1979, p. 139; Makita et al., 1984, p. 386; Smith & Jungers, 1997, p. 544). The female/male weight ratio is 0.66 in 36 captive adults. Mean birth weight is 402 g in 89 captives born in colonies in Taipei and 430 g in 22 captives born in a colony in Southboro, Massachusetts (Ta- ble 2). External Measurements and Proportions External measurements are available for only ten wild-collected specimens of M. cyclopis (Ta- ble 1); of these, two are adults — one female and one male — and eight are immatures. To supple- ment these meager data, measurements of captive specimens and specimens of unknown history also are presented; however, these measurements probably are not directly comparable with those of wild-collected specimens. The following infer- ences are based on the limited available data. Head and body length is 440 mm in one wild- collected adult female and 650 mm in one wild- collected adult male (Table 1). The female/male length ratio is 0.68 in the two wild-collected adults, 0.86 in 25 captive adults, and 0.97 in six adults of unknown history. Although Bergner and Cranial Characters (Figs. 4, 5) In wild-collected specimens examined of M. cyclopis, the greatest length of the skull in adult males (125.2 mm, n = 15) averages approximate- ly 13% greater than in adult females (1 10.4 mm, n = 12) (Table 3). Skull length in both sexes of M. cyclopis tends to exceed that in M. fascicularis and M. mulatta (cf. Fooden, 1995, p. 38; 2000, p. 38). Available measurements of wild-collected specimens of M. cyclopis are inadequate to indi- cate whether or not skull length varies geograph- ically in this species (Table 4). In captive speci- mens of M. cyclopis, greatest length of the skull tends to exceed that in wild-collected specimens (Table 3; P < 0.02; cf. Komatsu, 1944. p. 18; Mouri, 1995, p. 189; Koppe et al., 1999, p. 78). The rostrum is moderately projecting in M. cy- clopis, more similar to that in M. fascicularis than to that in M. mulatta (cf. Murie, 1873, p. 779; FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 0) o Z FIELDIANA: ZOOLOGY CM a -a - (U u o ^ -J O '. >^ - ^ c C CS O J= EU "O ■^ 't aj T3 Os c OJ On 'g 3 ■ - C3 T3 ^ X c m .— ' e OS 'o u 3 w Q. ^ — C/D ^ C a. ■*- -4 Cu < •0 T3 c u u 0 C3 > ^ 3 c 3 0 J3 S 3 O o 0 > a 1— > C •o aj ,_o fl3 c ii 00 u, .;^ OC u 5. ■^ oa "^ XI 0 -a 2 c - [L 1) >. . s ,^ -^ n3 -^ ^ C -JO 0 ^ C 3 -J c P « 53 -1 •0 S>; _3 FOODEN AND WU: SYSTEMATIC REVIEW OF MAC AC A CYCLOPIS Key to Locality Numbers Northern Taiwan (N) 1. Hsi Lin. 2. Shihting Hsiang vicinity. 3. Mucha. 4. Sanhsia. 5. Shihtoushan. 6. Wulai; Wulai area. 7. Hsiaoi. 8. Tunghou. 9. Ilan (=Giran) region. 10. Ayu Shan. 1 1. Fu.shan Forest Station: Fushan Forest Station: Bo- tanic Garden. 12. Fushan ( = Rimogan). 13. Northern Cross-Island Highway. Fuhsing Hsiang. 550 m. 14. Fuhsing Hsiang 15. Ha-pen, Wulai Hsiang, 530 m. 16. Ha-pen. Wulai Hsiang. 535-570 m. 17. Ha-pen. Wulai Hsiang, 650 m. 18. Anpingkenghsi Mining Trail, Suao-Nanao Hsiang, 175 m. 19. Suao vicinity. 20. Wulaokenghsi Mining Trail. 21. Suhua road. 22. Lupishan Logging Trail, Nanao Hsiang, 245 m; 595-798 m. 23. Anpingkenghsi Mining Trail, Suao-Nanao Hsiang. 690 m: 1000 m. 24. Anpingkenghsi Mining Trail, Suao-Nanao Hsiang. 685 m"; 810-930 m. 25. Taipingshan. 26. Jentse: Taipingshan, northwestern foothill area. 27. Northern Cross-Island Highway, Tatung Hsiang, 1258 m. 28. Northern Cross-Island Hiahway, Tatung Hsiang, 460 m. 29. Northern Cross-Island Highway, Tatuna Hsiang, 720 m. 30. Northern Cross-Island Highway, Fuhsins Hsiana, 1050 m. 31. Loshan Logsina Trail, Wufens-Chienshih Hsiana, 1 270 m. " " 32. Loshan Logging Trail, Wufeng-Chienshih Hsiang, 1050 m: 1210 m. 33. Tunghotsun. 34. Fayun Ssu. 35. Huoyenshan. 36. Hsiaoponko. 37. Tawan: Dikanhsi-Hsuehchenhsi; Pipitaishan. 38. Tahsueh Shan Losainc Trail, Taian Hsiang, 1750 m. 39. Ssumahsien Logging Road: Tahsueh Shan, Hoping Hsiang, 1940-2100 m: Tahsueh Shan Logging Trail, Taian Hsiang. 1995 m. 40. Tahsueh Shan Logging Trail. Taian Hsians. 2540 m. 41. Taanhsi. 42. Tahsueh Shan Logging Road 230. Hoping Hsiang, 2545 m: 2565 m: Tahsueh Shan Logging Road 230, Taian Hsiang, 2515 m. 43. Tahsueh Shan area, Taian Hsiang, 940-2775 ni: Tahsueh Shan Hiking Trail, Taian Hsiang, 2385- 2765 m. 44. Hsueh Shan. 45. Wuling (Taoshan Pupu). 46. Nanhu Tashan. 47. Chutsun, Hsiulin Hsiang. 48. Chutsun, Hsiulin Hsiang, 1100 ni. 49. Chiumei Bridge: Lienhuachih, Hsiulin Hsiang, 800-1 100 m: 1 100-1200 m: Meiyuan; Taroko Na- tional Park. 50. Hsipao. 51. Tienhsiang. 52. Yen Hai Logging Trail. 53. Ta Ching Shui. 54. Chingshui Cliff: Hsiao Ching Shui. 55. Taroko National Park Headquarters. 56. Chingmei. 57. Hualien, mountains near: Hualien vicinity. 58. Mu-Kua-Chau: Shuilien Bridge No. 14. 59. Nengkao-Tungmen Powerline Right-of-way, Hsiu- lin Hsiang, 650 m. 60. Nengkao-Tungmen Powerline Right-of-way, Hsiu- lin Hsiang, 750-1200 m: Nengkao-Tungmen Pow- erline Right-of-way, Hsiulin Hsiang, 900-1200 m. 61. Nengkao-Tungmen Powerline Right-of-way, Hsiu- lin Hsiang, 1200-1310 m: Nengkao-Tungmen Powerline Right-of-way. Hsiulin Hsiang, 1300 m (24°02'N, 12r22'E). 62. Nengkao-Tungmen Powerline Right-of-way. Hsiu- lin Hsiang, 1300 m (24°0rN, 12r2rE). 63. Nengkao-Tungmen Powerline Right-of-way, Hsiu- lin Hsiang, 1300-1340 m: Nengkao-Tungmen Powerline Right-of-way, Hsiulin Hsiang, 1345 m. 64. Nengkao-Tungmen Powerline Right-of-way, Hsiu- lin Hsiang, 1 840-2085 m. 65. Peitunyenshan. Jenai Hsiang, 1900 m. 66. Juiyen, Jenai Hsiang: Juiyenhsi, Jenai Hsiang, 2190-2205 m. 67. Peitunyenshan, Jenai Hsiang: Peitunyenshan, Jenai Hsiang, 1700-1800 m. 68. Chientai Logging Trail, Jenai Hsiang, 1250- 1300 m. 69. Wanta Logging Trail. 70. Nanshanhsi, Jenai Hsiang: Nanshanhsi, Jenai Hsi- ang, 1300-1580 m. 71. Tungfeng. 72. Hui-Sun Experimental Forest, 73. Chunghsing Tahsueh. 74. Pahsienshan Forest Recreation Area. 75. Kukuan. 76. Shangnankeng; Tahsueh Shan Logging Road 220, Hoping Hsiang, 1925-1980 m: 2020 m: Tahsueh Shan Logging Road 221, Hoping Hsiang. 2030 m; Tahsueh Shan Logging Road 222, Hoping Hsiang, 2040 m: Wuling Farm. 77. Tahsueh Shan Logging Road 230, Taian Hsiang, 2250-2525 m. 78. Tahsueh Shan Logging Trail 230: Taian Hsiang, 2515 m. 79. Anmashan. 80. Hsuehshankeng. Hoping Hsiang, 750-880 m; Hsuehshankeng Logging Road, Hoping Hsiang, 1200 m. FIELDIANA: ZOOLOGY 81. 82. 83. 84. 85. 86. 87. 89. 90. 91. 92. 93. 94. 95. 96. 97. 98. 99. 100. 101. 102. 103. 104. 105. 106. 107. 108. 109. 110. 111. 112. 113. 114. 115. 116. 117. 118. 119. 120. 121 122. 123. 124. 125. 126. 127. Wushihkeng, ca. 1200 m; Wushihkeng vicinity. Wushihkeng, Hoping Hsiang; Wushihkeng, Hop- ing Hsiang, 600-1250 m; Wushihkeng, Hoping Hsiang, 635-1305 m. Shentao No. 8. Chingjen Ku. Takeng. 460-510 m. Maanliao. Tienleng. Paimao Shan Logging Road, Hoping Hsiang, 1140 m. Paimao Shan Logging Road, Hoping Hsiang, 850- 1040 m. Pingting, Taiping Hsiang, 270 m. Toupienkeng. Hsiangtao No. 136, Kuohsing Hsiang, 600 m. Kuohsing Hsiang. Kangou. Chiuchiu Feng. Shihchotzu, Tsaotun Chen, 300-400 m; Shuangtung. Peishankeng. Lienhuachih. Puli ( = Posia; Polisia); Puli vicinity. Linshoudien. Luliu-Chepinglun. Jihyueh Tan ( = Sun-Moon Lake, Tsuisia Lake) vi- cinity. Erpingshan. Hsiangtao No. 131. Chichi Tashan, Chichi Chen; Chichi Tashan, Chi- chi Chen, 700-1250 m. Tienchung Park. Chungliao Hsiang. Chinshuikou. Fuchouli; Luku Shan. Pakua Shan. Ershui Monkey Protection Area, Ershui Hsiang, 170-210 m; Puchung. Pingting, Linnei Hsiang, 150 m. Hupen. Kukeng. Kankenghsi. Changhu, Kukeng Hsiang, 310-350 m. Tsaoling Road. Juilung Pupu. Taichi Hsiaku (Gorge). Hsitou, Luku Hsiang, 1270-1360 m; Hsitou (=Chi Tou), 1000 m; Hsitou ( = Hsiton), Fenghuangshan, Luku Hsiang. Chunta Logging Trail, Hsini Hsiang (23°36-40'N, 120°52-56'E); Chunta Logging Trail, Hsini Hsi- ang, 1355-2515 m. Chunta Logging Trail, Hsini Hsiang, 810 m; Jenlun Logging Trail, Hsini Hsiang, 1300-1400 m. Chunta Logging Trail, Hsini Hsiang, 1580 m. Jenlun Logging Trail, Hsini Hsiang, 1400 m. Chunta Logging Trail, Hsini Hsiang (23°44'N, 120°56'E); Chunta Logging Trail, Hsini Hsiang, 2705 m; Chunta Logging Trail, Hsini Hsiang, 2790-2855 m; Jenlun Logging Trail, Hsini Hsiang, 1750 m. Chunta Logging Trail, Hsini Hsiang, 2495 m. Chunta Logging Trail, Hsini Hsiang, 2450-2455 m; Chunta Logging Trail, Hsini Hsiang, 2470 m. 128. Chunta Logging Trail, Hsini Hsiang, 2485 m. 129. Chunta Logging Trail, Hsini Hsiang, 1910- 1920 m. 130. Chunta Logging Trail, Hsini Hsiang, 1425 m. 131. Chitsaihu, Hsini Hsiang, 2600-2730 m. 132. Wanjung Logging Trail, Wanjung Hsiang, 1700 m. 133. Wanjung Logging Trail. Wanjung Hsiang, 820 m; Wanjung Logging Trail, Wanjung Hsiang, 960 m. 134. Wanjung Logging Trail, Wanjung Hsiang, 1410- 1600 m. 135. Wanjung Logging Trail, Wanjung Hsiang, 820 m; Wanjung Logging Trail, Wanjung Hsiang, 1085- 1165 m. 136. Wanjung Logging Trail, Wanjung Hsiang, 565 m; Wanjung Logging Trail, Wanjung Hsiang, 800 m. 137. Kuangfu vicinity. 138. Tahoshan. 139. Fuyuan Forest Recreation Area. 140. Shalihsien Hsi, Hsini Hsiang, 1350-2100 m; Sha- lihsien Hsi Logging Road, Hsini Hsiang; Shalih- sien Hsi Logging Road, Hsini Hsiang, 1405-1600 m; Shalihsien Hsi Logging Road, Hsini Hsiang, 1485-1800 m. 141. New Central Cross-Island Highway; New Central Cross-Island Highway, Hsini Hsiang, 2000 m (23°32'N, 120°53'E); Shenmu Logging Road. Hsi- ni Hsiang, 1260-2500 m; Shenmu Tsun. 142. Chushuihsi. 143. Alishan zone, Alishan Hsiang, 2260 m; Alishan zone, Alishan Hsiang, 2310 m. 144. Fengshan. 145. Shihpi. Kukeng Hsiang. 1010-1065 m. 146. Juifeng Bridge; Tashiaku. 147. Taiping, Meishan Hsiang. 920 m. 148. Chuchi. 149. Juili, Meishan Hsiang. 150. Juili, Meishan Hsiang, 700 m. 151. Niupi. 152. New Central Cross-Island Highway, Hsini Hsiang, 2300 m (23°30'N, 120°52'E). Absence Report A. Penghu Chuntao: Penghu Tao. Southern Taiwan (S) 1. Chia-I. 2. Kungtientsun. 3. Danayee. 4. New Central Cross-Island Highway, Alishan Hsi- ang, 1395 m. 5. Tapang. 6. Tefuyeh. 7. New Central Cross-Island Highway, Alishan Hsi- ang, 1655 m. 8. New Central Cross-Island Highway, Alishan Hsi- ang, 1 860-2700 m. 9. New Central Cross-Island Highway, Hsini Hsiang, 2300 m (23°28'N, 120°49'E); New Central Cross- Island Highway, Hsini Hsiang, 2300 m (23°28'N, 120°50'E); New Central Cross-Island Highway, Hsini Hsiang. 2300 m; (23°29'N, 120°49'E); Tzu- chung; Yushan National Park; Yushan ( = Yuishan), 2200 m. 10. New Central Cross-Island Highway, Hsini Hsiang, 2300 m (23°28'N, 120°5rE). (continued on the folowing page) FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 11. Lulin Shan Preserve. Alishan Hsiang, 2300-2420 m; Lulin Shan Preserve, Alishan Hsiang, 2700 m; New Central Cross-Island Highway, Hsini Hsiang, 2300 m (23°28'N, 120°52'E): Tatachia. 12. Paiyun Hiking Trail. 13. Nantzuhsien Hsi Logging Road. Alishan Hsiang, 1700-2120 m. 14. Nanshi Logging Road. 15. Patungkuan. 16. Lakulakuhsi Hiking Trail, Chohsi Hsiang, 1300 m (23°24'N, 121°05'E). 17. Lakulakuhsi Hiking Trail, Chohsi Hsiang, 1420- 1570 m. 18. Lakulakuhsi Hiking Trail. Chohsi Hsiang, 1455- 1735 m. 19. Lakulakuhsi Hiking Trail, Chohsi Hsiang, 1200 m. 20. Lakulakuhsi Hiking Trail, Chohsi Hsiang, 1300 m (23°24'N. 121°07'E): Lakulakuhsi Hiking Trail. Chohsi Hsiang. 1300-2205 m. 21. Lakulakuhsi Hiking Trail, Chohsi Hsiang, 1478 m. 22. A-Sun-Lai-Ga. 23. Lakulakuhsi Hiking Trail. Chohsi Hsiang, 1630 m. 24. Yulishan. 25. Lakulakuhsi Hiking Trail. Chohsi Hsiang, 800 m. 26. Taiping, Chohsi Hsiang. 27. Haianshanmo. 28. Takangkou. 29. Chunfu. 30. Antungwenchuan. 31. Tungli. 32. Chingshui Hsi Power Station; Choching. 33. Lakulakuhsi Hiking Trail. Chohsi Hsiang, 250- 420 m. 34. Changliang Logging Trail, Chohsi Hsiang, 980 m. 35. Changliang Logging Trail, Chohsi Hsiang, 825 m. 36. Lakulakuhsi Hiking Trail, Chohsi Hsiang, 710 m. 37. Lakulakuhsi Hiking Trail, Chohsi Hsiang, 950- 1030 m: Lakulakuhsi Hiking Trail, Chohsi Hsiang. 1200 m. 38. Lakulakuhsi Hiking Trail, Chohsi Hsiang, 1620 m. 39. Lakulakuhsi Hiking Trail, Chohsi Hsiang, 1690 m. 40. Tafen slope. 41. Lakulakuhsi Hiking Trail. Chohsi Hsiang, 1220- 1320 m; Lakulakuhsi Hiking Trail, Chohsi Hsiang, 1490-1520 m. 42. Southern Cross-Island Highway, Haituan Hsiang, 2065-2090 m. 43. Southern Cross-Island Highway, Taoyuan Hsiang, 2185 m; Southern Cross-Island Highway. Taoyuan Hsiang. 2310-2455 m; Southern Cross-Island Highway, Taoyuan Hsiang. 2585 m. 44. Southern Cross-Island Highway, Taoyuan Hsiang, 2015-2110 m; Southern Cross-Island Highway, Taoyuan Hsiang, 2250-2415 m. 45. Southern Cross-Island Highway, Taoyuan Hsiang, 1635-1750 m; Southern Cross-Island Highway, Taoyuan Hsiang, 1825-1975 m. 46. Southern Cross-Island Highway, Taoyuan Hsiang, 975-1050 m: Southern Cross-Island Highway, Taoyuan Hsiang, 1445-1565 m. 47. Meishan; Southern Cross-Island Highway, Tao- yuan Hsiang. 995-1235 m; Southern Cross-Island Highway, Taoyuan Hsiang, 1250-1445 m. 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. 85. 87. 90. 91. Minsheng. Tsengwen Shuiku. Tapu Hsiang. Kueitan. Nanhua Hsiang. Liukuei Hsiang. Chuyunshan Logging Trail, 2300-2320 m. Chuyunshan Logging Trail. 2260-2275 m. Chuyunshan Logging Trail. 2220 m. Southern Cross-Island Highway, Haituan Hsiang. 1855 m. Southern Cross-Island Highway, Haituan Hsiang, 1665 m. Southern Cross-Island Highway. Haituan Hsiang, 930-943 m; Southern Cross-Island Highway. Hai- tuan Hsiang, 1040-1125 m. Southern Cross-Island Highway. Haituan Hsiang. 1170 m. Wulu Logging Trail, Haituan Hsiang, 2240 m. Wulu Logging Trail, Haituan Hsiang, 2060- 2215 m. Wulu Logging Trail. Haituan Hsiang, 1925- 2005 m. Luliaoshan. Wulu Logging Trail, Haituan Hsiang, 860 m. Southern Cross-Island Highway, Haituan Hsiang. Hungshih Logging Trail. Kuanshan Chen vicinity; Kuanshan ( = Kuang Shan Working Circle). Southern Cross-Island Highway, Haituan Hsiang, 390 m. Southern Cross-Island Highway, Haituan Hsiang. 435 m. Southern Cross-Island Highway, Haituan Hsiang. 515 m. Changliang Logging Trail. Chohsi Hsiang. 1920 m. Changliang Logging Trail. Chohsi Hsiang, 1965 m. Chungan. Iwan. Chengkung Chen; Chungan. Poai- li. Chengkung Chen. Hsinkangshan. Sanminli. Sanhsienli. Hoping. Hsini. Tousienchiao. Peiyuan. Tayuan, Haianshanmo; Tayuan (=Tai Yuan). Tungho township. Tulanshan. Shanli. Peinan Hsiang. Yenping Logging Trail. Yenping Hsiang. 1 150 m. Yenping Logging Trail, Yenping Hsiang; Yenping Logging Trail, Yenping Hsiang, 1215-1360 m. Tungshing. Yenping Logging Trail, Yenping Hsiang, 1330- 1525 m; Yenping Logging Trail, Yenping Hsiang, 1555-1650 m; Yenping Logging Trail, Yenping Hsi- ang, 1600 m. Yenping Logging Trail, Yenping Hsiang, 1705-1730 m; Yenping Logging Trail. Yenping Hsiang. 1740 m. Yenping Logging Trail, Yenping Hsiang, 1665- 1835 m. Yenping Logging Trail. Yenping Hsiang. 1820- 2060 m. FIELDIANA: ZOOLOGY Fooden, 1995, p. 38; 2000, p. 38). From infancy to adulthood, the rostral-postrostral ratio more than doubles in M. cyclopis males (Table 3); dur- ing the same period, relative zygomatic breadth increases by less than 10%. Development of a sagittal crest apparently is relatively rare in wild- collected M. cyclopis adult males (3 of 15 speci- mens examined), as in M. mulatta. Mean ages (in days) of emergence of deciduous teeth in 15 M. cyclopis captives studied by Yang et al. (1968, p. 543) are as follows: i„ 5.4; i', 1 1.2; i2, 19.4; 12, 30.4; c„ 54.6; c', 53.6; m„ 63.3; m', 63.4; m., 125.2 (n = 14); m^, 139.2. These ages are less than corresponding emergence ages re- ported for captive M. fascicularis and M. mulatta (Fooden, 1995, p. 38; 2000, p. 39). Morphological details of permanent teeth in captive M. cyclopis have been studied by Saheki et al. (1962, p. 101; cf. Table 26 below). Molecular Biology and Genetics Mitochondrial DNA Mitochondrial DNA has been investigated in six studies of three M. cyclopis captives without locality information (Table 5). These studies agree that, among living fascicularis-group species, mtDNA in M. cyclopis is most similar to that in M. mulatta and M. fuscata. In fact, in one of these 500 m (23°00'N, 1 92. Yenping Logging Trail, Yenping Hsiang, 2085 m. 93. Yenping Hsiang. 94. Lichia Logging Trail, Peinan-Yenping Hsiang, 1420 m. 95. Chuyunshan Logging Trail, 1360 m; Chuyunshan Logging Trail, 1400 m. 96. Chuyunshan Logging Trail, 1135-1300 m. 97. Chuyunshan Logging Trail, 1350 m. 98. Chuyunshan Logging Trail, 930-960 m. 99. Chuyunshan Logging Trail, 1290 m. 100. Shanping. 101. Wushan, Nanhua Hsiang, 500 m (23°0rN, 120°30'E). 102. Wushan, Nanhua Hsiang, 500 m (23°01'N, 120°29'E). 103. Wushan, Nanhua Hsiang, 120°29'E). 104. Chishan vicinity, Chishan-Meinon, 200 m (22°54'N, 120°30'E). 105. Chishan vicinity, Chishan-Meinon, 200 m (22°53'N, 120°29'E). 106. Kaohsiung (=Takao) vicinity; Shou Shan. 107. Kuanyinshan. 108. Pingtung vicinity. 109. Santimen. 110. Wutao Shan. 111. Hsiao Kueihu. 112. Lichia Logging Trail, Peinan-Yenping Hsiang, 1400-1475 m; Lichia Logging Trail, Peinan-Yenp- ing Hsiang, 1435-1565 m. 113. Lichia Logging Trail, Peinan-Yenping Hsiang, 1 1 10-1 130 m; Lichia Logging Trail, Peinan-Yenp- ing Hsiang, 1 175-1370 m. 1 14. Lichia, Peinan Hsiang. 115. Lichia Logging Trail, Peinan-Yenping Hsiang, 1055-1 100 m. 116. Lichia Logging Trail, Peinan-Yenping Hsiang, 930-950 m; Lichia Logging Trail, Peinan-Yenping Hsiang, 970-1045 m. 1 17. Peinan. 118. Taitung vicinity. 119. Chienho. 120. 121. 122. 123. 124. 125. 126. 127. 128. 129. 130. 131. 132. 133. 34. 135. 136. 137. 138. 139. 140. 141. 142. 143. 144. 145. 146. 147. 148. 149. 150. 151. 152. 153. Chihpen Wenchuan. Chihpen Hsi, right bank. Chihpen Hsi, left bank. Huayuan. Chihpen Logging Trail. Chiniunshan. Chinlun, Taimali Hsiang. Toliang. Tawu Mountain Nature Preserve, Taimali Hsi, 300-400 m. Taimali Hsi; Tawu Mountain Nature Preserve. Chaochou. Wanglung. Linpien Hsiang. Fangliao vicinity; Shuitiliao. Fangshan Hsi. Taku. Tachu. Tawu Hsi. Shouchia. Lilungshan. Mutan. Lungkeaou. Hengchun (=Koshun; Koshiun). Renting Kungyuan. Lanjen Hsi; Liteshan. Laofoshan. Pailuz; Sabkun Hsi. Payao. Chushe Hsi; Kaoshihfoshan; Shihmenshe. Forest Section No. 15; Forest Section No. 27; Kao- shih, Mutan Hsiang; Tataan. Chiupeng; Kangtzu; Nanjenlu-Kangtzu. Chufengshan. Chulaoshu Shan; Manchoutsun. Hengchun Nature Preserve; Renting, 225 m; She- ting. Absence reports B. Liuchiu Yu. C. Lu Tao. D. Lan Yu. FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS Fig. 2. External characters in Macaco cyclopis: adult male photographed at Yushan National Park by Tsung-Su Ding. studies (Melnick et al.. 1993. pp. 286. 289). the interspecific mtDNA difference between M. cy- clopis and Burmese-South Chinese M. mulotta was found to be less than the intraspecific mtDNA difference between Burmese-South Chinese M. midatta and Pakastani-lndian M. mulotta. Esti- mates of the age of di\ ergence between M. cyclo- pis and M. mulotta vary in these studies from 0.25 Ma to 1.8 Ma (Table 5). The 464-bp sequence of the mtDNA D-loop of a captive specimen of M. cyclopis (without local- ity information) has been published by Kawamoto et al. (1999. p. 57). Based on a comparison of this sequence with that of a wild-collected specimen of M. fuscata and that of an apparent cyclopis X fuscata hybrid. Kawamoto et al. determined that the mother of the hybrid was M. cyclopis. In this study, the genetic distance between the M. cyclo- pis sample and the putative hybrid sample was less than 10% of the aenetic distance between the M. fuscata sample and the hybrid and M. cyclopis samples. In an unpublished series of studies of the 550- bp sequence of the mtDNA D-loop in natural pop- ulations of M. cyclopis, Jui-Hua Chu (ntuz; pers. comm.. 1997-2000: cf. Wu et al.. 2001b. p. 414) has investigated local and regional mtDNA vari- ation within this species. The three most impor- tant findings of this research are: 1. At Yushan National Park, central Taiwan, haplotypes are identical in all samples from seven groups studied. 2. At Shou Shan, southwestern Taiwan, four haplotypes were identified in samples taken from females in 13 groups. Ten of these groups have the same haplotype: each of the remaining three groups has a different haplotype. Three haplo- types at this locality — including the haplotype common to ten of the groups — are more similar to each other (sequence divergence = O.S9c) than 10 FIELDIANA: ZOOLOGY Table 1 . External measurements and proportions in Macaca cyclopis. Relative hind Relative Age/sex class- Head and body length (mm) Relative tail length (T/HB X 100) foot length (HF/HB X 100) ear length (E/HB X 100) Weight (kg) Infant 289 Wild-collected : 73.4 specimens^^ 34.3 11.1 Juveniles 374.7 ± 30.01 87.2 ± 7.84 34.9 ± 2.54 8.9 ± 1.37 3.60 ± 1.545 335-405 78.3-95.8 30.2-37.3 7.5-11.3 1.89-5.25 Subadult male (6) 490 {5f 79.6 {6) (6) 6.3 {4) Adult female 440 88.6 — 6.8 — Adult male 650 69.2 25.4 7.4 8.12 Infants Captive specimens* 1.15^ 0.60-1.70 Juveniles 434.9 ± 69.02 83.3 ± 7.63 31.9 7.4 ± 1.28 (2) 4.47 ± 1.699 338-550 72.2-93.9 5.9-8.2 1.60-7.00 Subadult females (//) 446.7 ± 15.27 ill) 85.4 ± 4.54 (/) (i) {14) 4.40 ± 0.400 430-460 80.2-88.9 4.00-4.80 Subadult males (i) 522.5 (-?) 88.8 {3) 7.10 ± 1.268 485-560 86.6-91.1 5.60-8.70 Adult females (2) 466.9 ± 29.56 (2) 82.2 ± 4.10 4.7 {4) 6.69 ± 1.497 420-530 75.0-94.3 2.50-9.50 Adult males {18) 544.3 ± 67.85 {18) 90.0 ± 4.38 21.2 (/) 6.6 {27) 10.21 ± 3.679 475-680 84.9-94.7 6.30-18.50 (7) (6) (/) il) (9) Fetus 151 Specimens of unknovcn history^'^ 82.1 — Juvenile 391 88.7 37.3 8.1 — Subadult males 574.5 529-620 74.2 66.1-82.2 30.6 7.4 — Adult females (2) 501.4 ± 63.36 (2) 79.0 ± 10.54 31.9 ± 4.14 8.0 ± 1.66 446-600 65.0-91.0 25.8-35.0 5.5-10.0 Adult male (5) 519 (5) 77.3 {4) 29.5 (5) 8.1 — ' Mean ± SD (where n > 2), extremes, and sample size (italicized figures in parentheses); cf. Horikawa (1932, p. 69). ^ 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. ^ Sample size = 1 unless otherwise indicated. ^ Infant: usnm 394177, Maanliao. Juveniles: amnh 184955, Hualien, mountains near; nmns T1726, Ta Ching Shui, Yio 733'' and 734, Hengchun; Shui-Wei Chang (pers. comm., 27 Apr. 1999), measurements of two specimens, Luku Shan. Subadult male: usnm 296795, Kaohsiung vicinity. Adult female: usnm 296794, Kaohsiung vicinity. Adult male: Dien (1958, p. 346), Wulai. ■'' Excludes questionable relative tail length = 1 14.0% (456 mm/400 mm) in yio 733, Hengchun. " Infants: kupri 3658, 6082. Juveniles: kupri 2795, 2796, 3652, 3653, 3660; nmns 000546; usnm 296796; Hill (1974, p. 589) (PM. 16); Hai-Yin Wu, measurements of eight zoo animals. Subadult females: Hai-Yin Wu, measure- ments of three zoo animals. Subadult males: kupri 3654, 3655; Hai-Yin Wu, measurements of two zoo animals. Adult females: kupri 1745, 2913, 3056, 3224, 3651, 3656, 3657, 3659, 3852, 4238, 5228, 5843; usnm 296797; Hai-Yin Wu, measurements of 17 zoo animals. Adult males: kupri 2514, 4233, 6431; ntuz Aoki 3484; Hai-Yin Wu, mea- surements of six zoo animals. ' Mean birth weight is 0.408 kg in 109 laboratory-born captives (see Table 2). ^ Fetus: tm MA-0378. Juvenile: tesri ace. no. T 0177. Subadult males: nmns T 3911; ntuz uncatalogued. Adult females: nmns 000547, T 1883, T 3909; ntuz Aoki 94; tesri ace. no. T 01 15. Adult male: nmns T 1725. FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS Table 2. Mean birth weight (g) reported in laboratory-born Macaca cyclopis. Taiwanese laboratories U.S. laboratory Yang et al. Peng (1968, p. 538) (1973, et al. p. 209) Price et al. (1972, p. 304) Sex N Mean weight N Mean weight^ N Mean weight- Females Males Totals 22 13 35 408 30 393 22 402 (300-530)^ 52* 398 ± 67 407 ± 55 402 ± 61 7 7 22 430 ± 45 ' Both studies were conducted at the College of Medicine, National Taiwan University, and the U.S. Naval Medical Research Unit No. 2. - SD also indicated. ^ Extremes. * Inadvertently recorded as "54" in cited reference. to the fourth haplotype (sequence divergence = 3.0%). 3. Based on paired comparison of sequence di- vergence, samples from localities in various parts of Taiwan were divisible into four regional groups: southern Taiwan (Kaohsiung and Ping- tung counties), central Taiwan (Chia-I and Nantou counties), eastern Taiwan (Hualien county), and northeastern Taiwan (Ilan county). The within- group sequence divergence is <4.0%, and the be- tween-group sequence divergence is 4.5-9.0%. Nuclear DNA The hinge region of the immunoglobulin C„ gene was studied by PCR amplification and elec- trophoresis of three M. cyclopis samples (Sumi- yama et al., 1998, p. 33). In these samples, the size and sequence of the PCR products were found to be individually variable. Similar poly- morphism also was detected in six other species of macaques, in baboons, and in colobines. This polymorphism is hypothesized to be the result of 550 Relative tail length (T/HB): 1 .00 0.90 0.80 0.70 0.60 100 150 200 250 300 350 400 450 500 550 600 650 Head and body length (mm) Fig. 3. Tail length vs. head and body length in Macaca cyclopis. 12 HELDIANA: ZOOLOGY Table 3. Cranial measurements' and proportions in Macaca cyclopis. - Relative zygomatic Rostral-postrostral Greatest length breadth Postrostral length ratio Age/sex class' (mm) (ZB/GL X 100) (mm) (R/PR X 100) Wild-collected specimens Infants 81.0 ± 1.46 63.5 ± 2.07 69.9 ± 1.47 25.6 ± 4.68 79.8-83.5 60.8-65.5 69.0-71.6 20.8-30.1 (5) (5) (3) (3) Juveniles 100.9 ± 7.00 67.0 ± 2.32 77.5 ± 3.67 36.8 ± 1.61 92.4-113.4 63.7-71.2 74.1-83.7 35.6-39.3 (10) (W) (5) (5) Subadult female* 104.2 66.3 — — Subadult males 120.1 ± 4.59 70.0 ± 1.93 86.0 ± 2.58 50.6 ± 2.92 114.0-125.9 66.6-72.2 82.9-90.3 45.6-55.1 (8) (8) (7) (7) Adult females^ 110.4 ± 2.40 67.6 ± 1.75 81.4 ± 2.02 48.2 ± 3.04 107.7-1 14.5 64.7-70.4 78.7-83.4 44.7 ± 51.1 (12) (12) (4) (4) Adult males' 125.2 ± 5.20 69.2 ± 2.58 88.5 ± 4.85 55.2 ± 2.10 114.6-133.4 64.4-73.6 77.8-93.1 52.6-58.2 U5) (14) Captive specimens (8) (8) Adult females 113.6 ± 6.16 68.4 ± 2.56 83.6 ± 3.58 47.8 ± 3.65 100.0-127.3 64.0-77.0 75.6 ± 95.1 41.1-56.4 (44) (44) (44) (44) Adult males 131.0 ± 7.47 70.8 ± 2.89 91.2 ± 3.90 54.3 ±3.13 113.0-142.6 65.4-76.1 82.9-98.2 49.0-60.6 (23) (23) (22) (22) Specimens of unknown history Adult females 116.2 ± 6.22 67.9 ± 0.65 84.2 ± 2.68 50.7 ± 4.01 107.9-122.9 67.1-68.4 80.3-86.5 46.8-56.3 (4) (4) (4) (4) Adult males 122.6 ± 6.02 69.8 ± 2.36 87.6 ± 2.60 52.8 ± 5.36 113.7-129.3 67.1-73.0 84.4-90.3 45.6-60.3 (6) (6) (6) (6) ' For definition of measurements, see Fooden (1969, p. 40); cf. Horikawa (1932, p. 69). - Mean ± SD (where n > 2), 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. '' Sample size - 1 . 5 Cf. Table 4. positive selection for maintenance of the activity of the immunoglobulin IgA antibody. Polymorphic microsatellite loci in captive and provisioned natural populations of M. cyclopis were studied by Chu et al. (1999, p. 575). These loci were found to be useful in paternity deter- mination. In addition, at one highly polymorphic locus, one group of alleles in M. cyclopis was found to be notably larger (269-285 bp) than al- leles at the same locus in M. fascicularis (<206 bp) and M. mulatta (<250 bp). Tosi et al. (2000, p. 138), who sequenced two Y-chromosome genes (TSPY and SRY) in a M. cyclopis captive without locality information and compared these sequences with those in samples of various populations of 17 other macaque spe- cies, found that M. cyclopis sequences were most similar to those in o\hcr fascicularis-grou^ spe- cies. Within the fascicularis group, M. cyclopis clustered most closely with M. mulatta (south- eastern China, Burma, India) and Vietnamese M. fascicularis (unresolved polychotomy); next inost closely with M. fuscata; and least closely with non-Vietnamese M. fascicularis (Thailand, Java, Borneo, Philippines). The somewhat incon- gruous Y-chromosomal sequence similarity of M. cyclopis to Vietnamese M. fascicularis was attri- buted by Tosi et al. to possible hybridization be- tween Vietnamese M. fascicularis and M. mulat- ta. FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 13 Fig. 4. Skull of adult female Macaca cyclopis — usnm 296794, Kaohsiung vicinity, Kaohsiung Hsien. (Photographs by John Weinstein, The Field Museum.) Blood Proteins Variation of blood-protein allele frequencies is relatively low in available samples of M. cyclopis (Table 6). Of 2 1 loci that are polymorphic in Ma- caca. 1 1 are monomorphic in M. cyclopis and eight have major alleles with frequencies of at least 0.90. At one of the remaining loci (TBPA), the major allele has a frequency of 0.75; at the other (Pa), the major allele has a frequency of 0.50. Within ihe fasciciilaris group, blood proteins in M. cyclopis are more similar to those in M. mulatto than to those in M. fascicularis, and least similar to those in M. fuscata (Fooden & Lanyon, 1989, p. 224); based on the amount of blood-pro- tein difference between M. cyclopis and M. mu- latta, Nozawa et al. (1977, p. 26) have estimated that these two species began to diverge ca. 70 Ka. Blood-protein analysis — particularly of the ADA. Dia. and Tf loci — indicates the occurrence of hybridization in Japan between a recently in- troduced population of M. cyclopis and a native population of M. fuscata (Kawamoto et al.. 1999, p. 55). 14 FIELDIANA: ZOOLOGY Fig. 5. Skull of adult male Macaca cyclopis — usnm 3991 1 1, Hsiaoi, Wulai District, Taipei Hsien. (Photographs by John Weinstein, The Field Museum.) Karyology The karyotypes of one M. cyclopis female and one M. cyclopis male have been published by Chen et al. (1981, p. 73); precise geographic or- igins of the specimens are not specified. These M. cyclopis karyotypes appear similar to those in oth- er species of Macaca (cf. Makino, 1952, p. 290; FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 15 Table 4. Local variation in greatest length of skull in wild-collected Macaca cyclopis (cf. Table 3). Latitude Longitude Locality' (N) (E) Mean ± SD Extremes N Adult females Chingshui Cliff 24° 12' 121°40' 110.9 1 Hui-Sun Experimental Forest 24°03' i2i°or 108.0 1 A-Sun-Lai-Ga 23°22' i2i°ir 114.5 1 Kaohsiuno vicinitv 22°39' 120°15' 107.8 1 Tawu Mountain Nature Preserve 22°35' 120°49' 108.1 1 Manchoutsun 22°or 120°5r 110.9 ± 2.12 107.7-113.9 7 All Adult males- 110.4 ± 2.40 107.7-114.5 12 Wulai' 24°52' 121°33' 121.0 1 Ayu Shan 24°47' 121°36' 125.7 1 Puli 23°58' 120°57' 129.6 1 Mu-Kua-Chau 23°50' 121=33' 128.0 126.5-129.5 2 Manchoutsun 22°or 120°5r 124.6 ± 5.52 114.6-131.6 7 Kentina 21°58' 120°49' 127.8 122.2-133.4 2 ? (UMMZ 85349) — — 117.5 1 All 125.2 ± 5.20 114.6-133.4 15 ' In this and subsequent tables, localities are listed in north-south order, where sufficient information is available. - Excludes LSNM 3991 1 1 (Coll. No. 392), adult male, trapped 6 Jun. 1963 at Hsiaoi (24°5rN, 121°35'E), greatest length of skull 131.1 mm. This monkey was maintained as a laboratory captive for approximately 1 year, until 22 May 1964 (Bergner & Jachowski, 1968, pp. 22, 30); its cranial measurements have been included in the calculation of statistics for captive adult males presented in Table 3. ' Measurement from Dien (1958. p. 346). T.\BLE 5. Sister groups and divergence ages of Macaca cyclopis as determined in five studies of mtDNA in fascicularis-gToup species; in these studies, M. cyclopis data were derived from a total of three captive specimens. References Analytical method Origin of M. mulatta samples studied Sister groups' of Af. cyclopis Tree construction method Neighbor-joining OtheH -Divergence ages (Ma) Studies in which M. cyclopis data were derived from one captive specimen' Hayasaka et al. (1988, p. 278) Melnick et al. (1993, p. 289)^ Zhang and Shi (1993a, p. 13)^ Hayasaka et al. (1996, p. 1050) Restriction sites, 17 enzymes Restriction sites. 15 enzymes Restriction sites. 16 enzymes Sequencing. 896- bp region India Pakistan, India, Burma, S China India.*^ S China India, unknown MUL (India) + MUL (India) + 0.9-1.8 FUS MUL (Burma & S China) + FUS MUL (India) + FUS MUL (India) + FUS FUS MUL (Burma & S China) + FUS MUL (S China) MUL (India)^ 1.8« 0.63-0.70 Study in which M. cyclopis data were derived from two captive specimens Morales and Melnick Restriction sites, N India. S China MUL (S China) MUL (N India) 0.25 (1998, p. 15) 22 enzymes ' Abbrexiations: FUS. M. fuscata: MUL, M. mulatta. -Maximum parsimony: Hayasaka et al. (1996). PAUP: Melnick et al. (1993): Morales and Melnick (1998). Un- weighted pair group: Hayasaka et al. ( 1988): Zhang and Shi (1993a). ' The same M. cyclopis captive provided mtDNA data for all four studies. ■•The source of M. cyclopis data used in this study is Hayasaka et al. (1988, p. 273). ' The source of the Indian M. mulatta data used in this study is Hayasaka et al. (1988, p. 273). *- See Zhang and Shi (1993b, p. 602). This is the sister group of M. cyclopis in a tree constructed from data for 896 sites: in a tree constructed from data for 664 sites, the sister group of M. cyclopis is MUL (India) + FUS. 16 FIELDIANA: ZOOLOGY Table 6. Allele frequencies (%) at blood-protein loci in samples of Macaco cyclopis.^ Locus Major allele: frequency Minor alleles: frequencies Acp 121 A 100 ADA 67 2 100 Alb 167 A : >99 B: <1 CA-I 139 A 99 D2: 1 CA-IC 92 X+ 100 CellEs 46 1 100 ChEs 50 1 100 Dia 47 c 93 A: 4; D: 3 Go 99 SI 100 HbA2 151 1 100 HbB^ 102 1 100 IDH 44 2 92 I: 8 LDHB 50 1 100 Pa 99 D 50 FI: 28; F2: 22 PGD 137 A : >99 S: <1 PGM-I 39 1 100 PGM-II 39 1 100 PHI 64 I 92 4: 8 PI 139 C 90 B: 7; D: 2; F: 1 TBPA 305 F 75 S: 25 Tf 267 C 91 D: <1; E: 4; F: <1; F": 4; G: < 1 ' For references and key to locus abbreviations, see Fooden and Lanyon (1989, p. 214) and Chao (1990, pp. 52, 60, 66, 68, 73). - Cf. Chao (1990, p. 60), who similarly reports that a sample of M. cyclopis (n = 99) was monomorphic for hemoglobin (Hb, allele S). Ardito, 1979, p. 251; Cao et al., 1981, p. 120). The diploid chromosome number in M. cyclopis is 42. Parasites Protistans at least four species of Entamoeba and one spe- cies each of Endolimax and lodamoeba (Table 8). The incidence of Entamoeba species in these un- localized samples varies from 11.8% to 41.2%, the incidence of Endolimax nana is 16.2%, and the incidence of lodamoeba buetschlii is 14.2%. Malaria — Macaca cyclopis is naturally infect- ed by Plasmodium inui and P. cynomolgi, which are the two most widely distributed malarial par- Flagellates — Fifty-three of 594 blood sam- ples (8.9%) collected from natural populations of M. cyclopis in seven counties in eastern and cen- tral Taiwan were positive for a trypanosome sim- ilar to Trypanosoma conorhini (Table 7). Among counties, the incidence of positive samples varied from 5.4% to 40.0%. The incidence of positive samples in Taipei, Ilan, and Hualien counties — adjacent counties in northeastern Taiwan — tends to exceed that in the remaining four counties; however, sample sizes are small for Taipei, Ilan, and Hualien counties. Three of 38 unlocalized fecal samples (7.9%) of M. cyclopis were positive for Trichomonas hominis (Kuntz et al., 1968, p. 123). Amebas — Endoparasitic amebas that have been detected in fecal samples of M. cyclopis include Table 7. Incidence of trypanosome infections {Try- panosoma cf. conorhini) detected in samples of natural populations of Macaca cyclopis. ' N Infected County No. % Taipei 18 4 22.2 Ilan 7 1 14.3 Hualien 15 6 40.0 Nantou 13 I 7.7 Kaohsiung 75 5 6.7 Taitung 135 18 13.3 Pingtung 331 18 5.4 Totals 594 53 8.9 ' References: Cross et al. (1970, p. 15; 1983, p. 536). Cf. Kuntz et al. (1968, p. 126), who report Trypanosoma sp. in two unlocalized M. cyclopis individuals. FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 17 Table 8. Incidence of anieba species detected in un- localized fecal samples of Macaco cyclopis (N - 136).' Incidence Species No. % Entamoeba coli 56 41.2 E. hartmanni 16 11.8 E. histolytica 43 31.6 E. polecki 41 30.1 Entamoeba sp.- 34 25.0 Endolimx nana 22 16.2 lodamoeba buefsclilii 19 14.0 ' Reference: Kuntz et al. (1968. pp. 123. 124). - Includes one or more unidentified species. asites of macaques; these two parasites also nat- urally infect M. fascicularis, M. mulatta, and sev- eral other macaque species (Hsieh, 1960, p. 483; Garnham, 1966, pp. 207, 286, 340; Coatney et al., 1971, pp. 90. 255, 318; Collins, 1988. pp. 1477, 1484, 1485; cf. Fooden, 1994, p. 576). Maccica cyclopis probably also is naturally infected by P. knowlesi, which is otherwise restricted to M. fas- cicularis and M. nemestrina in Southeast Asia, but evidence for the M. cyclopis-P. knowlesi associ- ation is not conclusive (see references above). The length of the asexual erythrocytic cycle is 72 hr (quartan) in P. inui, 48 hr (tertian) in P. cynom- olgi, and 24 hr (quotidian) in P. knowlesi. The incidence of natural malarial infections in M. cyclopis apparently is greater than 25% (Table 9); in one survey in which the sex of monkeys was reported, the incidence of infections in males was approximately one-half of that in females. As expected, the incidence of malarial infections in M. cyclopis tends to be greater in wet areas of Taiwan than in drier areas (Bergner & Jachowski. 1968, p. 27). The natural vector of malaria in M. cyclopis probably is Anopheles takasagoensis, which is the only Taiwanese member of the Leu- cosphyrus Group (Peyton & Harrison, 1980, pp. 336, 343); this group of mosquitoes apparently is essential in the transmission of simian malaria in Asia (Coatney et al., 1971, p. 24). CiLiATE — Bakmtidiiim coli was detected in 45 of 136 unlocalized fecal samples (33.1%) of M. cyclopis that were studied by Kuntz et al. (1968, pp. 123, 124). and a similar or identical parasitic ciliate was also frequently detected in M. cyclopis fecal samples collected at Yushan National Park, central Taiwan (Wu & H.-Y. Lin, 1996, p. 24). At Yushan National Park, the incidence of this par- asite in M. cyclopis samples was highest in sum- mer and autumn. Trematodes Two species of flukes have been reported as natural parasites of M. cyclopis (Table 10). One of these, an unidentified species of Lecithoden- driidae. has been detected in M. cyclopis only once. The more common fluke parasite of M. cy- clopis is Ogmocotyle indica (Family Notocotyli- dae); in M. cyclopis samples examined, the inci- dence of this fluke varies from 0% to 66.7%. The geographic pattern of this variation is not readily interpretable; Yu-Li, where the reported incidence of O. indica is 66.7% (n = 3). is less than 50 km east of Yushan National Park, where the reported incidence is 0% (n = 134). The human liver fluke, Clonorchis sinensis, has been detected in one M. cyclopis individual by Kuntz et al. (1968, p. 128). However, these au- thors suspect that the infection may have been ac- Table 9. Incidence of natural malarial infections detected in surveys of Macaco cyclopis populations. Survey area N Infected Survey date No. % References' ca. 1940 Taitung, Hualien 28 11 39.3 / 1949 "southern Taiwan" 35 182 51.4 2 1962-1964 "throug! hout Taiwan" 277 713 25.6^ 3 ca. 1965 Taitung •7 7 71.9 4 Total >340 >1()0 >29.4 ' Key to references: /. Hsieh (1960. p. 479). 2. Hsieh (1960, p. 482). 3. Bergner and Jachowski (1968, p. 27). 4. Peyton and Harrison (1980, p. 342). - Possibly includes two Hepatocystis infections. ' Possibly includes Hepatocystis infections. •* 13.7% in 102 males, 27.2% in 136 females (.sex undetermined in 39 host individuals). 18 FIELDIANA: ZOOLOGY Table 10. Incidence of trematode parasite infections detected in Macaco cyclopis samples. N Incidence Locality No. % References^ Notocotylidae: Ogmocotyle indica^ Unknown 98 28 28.6 1 Unknown 18 2 11.1 2 Yu-Li, Taitung Hsien 3 2 66.7 3 Yushan National park 134 0 0 3 Shou Shan 131 0 0 3 Lecithodendriidae: Gen. sp. ? Unknown 18 1 5.6 2 ' Excludes one infection with Clonorchis sinensis reported by Kuntz et al. (Ref. /), who suspect that this infection was acquired in captivity. - Key to references: /. Kuntz et al. (1968, pp. 124, 128). 2. Yoshimura et al. (1968, p. 292; 1969, p. 460). 3. Hai- Yin Wu. Huey-Yu Lin, and Yao-Sung Lin (unpublished data). ^ The species reported by Yoshimura et al. (Ref. 2) is O. ailuri, which may be a synonym of O. indica (see Ref. 1). cidentally acquired while the monkey was being held in captivity. Cestodes Two species of tapeworms have been detected at relatively low frequencies (<5%) in fecal sam- ples of M. cyclopis (Table 11). Nematodes Eleven species of nematodes have been report- ed as parasites in natural populations of M. cyclo- pis (Table 12). Of these, the species most fre- quently detected in M. cyclopis are Trichuris tri- chiiira (68.3% incidence), Oesophagostomum bi- Table 11. Incidence of cestode parasite infections detected in fecal samples of Macaca cyclopis. Incidence Refer- Locality N No. % ences' Bertiella studeri Unknown 110 4 3.6 / Yushan National Park 134 2 1.5 2 Shou Shan 131 3 2.2 2 Totals 375 9 2.4 Taenia hydatigena Unknown 57 2 3.5 3 ' Key to references: /. Kuntz et al. (1968, p. 124); Jessee et al. (1970. p. 87). 2. Hai-Yin Wu, Huey-Yu Lin, and Yao-Sung Lin (unpublished data). 3. Kuntz and Meyers (1967, p. 84). furcum (29.6%), Strongyloides fuelleborni (26.7%), and Ternidens sp. (11.5%); the reported incidence in M. cyclopis of each of the other sev- en parasitic nematode species is less than 10%. The incidence of nematode parasitism varies geographically in M. cyclopis. Although 42% of 55 M. cyclopis individuals trapped in Taipei and Taoyuan counties — the two northernmost counties in Taiwan — were parasitized by Macacanema for- mosana, none of 222 individuals trapped in ten other counties were positive for this parasite (Bergner & Jachowski, 1968, p. 24); the area of Macacanema parasitism in Taipei and Taoyuan counties is in a region of especially heavy rainfall (Bergner & Jachowski, 1968, p. 15). Similarly, the incidence of nematode parasitism in M. cyclo- pis at Yushan National Park, central Taiwan, dif- fers from that at Shou Shan, southwestern Taiwan (Table 12). The incidence of nematode parasitism in M. cy- clopis also varies seasonally and sexually. The re- ported incidence of parasitism by Oesophagosto- mum, Strongyloides, and Trichuris is greatest in summer and autumn (Lin et al., 1996, abstract no. 712; Wu & H.-Y. Lin, 1996, p. 24). Analyzed ac- cording to sex, the incidence of Macacanema par- asitism in Taipei and Taoyuan counties is 28.0% in 25 M. cyclopis males and 53.3% in 30 M. cy- clopis females (Bergner & Jachowski, 1968, p. 28). The same species of Enterobius that parasitizes M. cyclopis in Taiwan (£. macaci) also is known to parasitize M. mulatta in China (Hugot, 1999, p. 544). FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 19 Table 12. clopis. ' Incidence of nematode parasite infections detected in samples of natural populations of Macaca cy- Yushan 1 Locality National Park Shou Shan unknown Total R( jfei Parasite- No. % No. % No. % No. % rences^ Class Adenophorea Order Stichosomida Capillariu sp. 134 2.2 131 2.2 12 0 277 2.2 /, 2 Trichiiris trick ii t ra 134 84.3 131 38.2 310 74.2 575 68.3 /, 2, 3, 4, 5 Class Secernentea Order Rhabditida Strongyloides fuellebomi 134 17.2 131 38.2 12 8.3 277 26.7 /, 2 Order Strongylida Oesophai^ostomwn bifurcitm 134 42.5 131 12.2 110 34.5 375 29.6 /. 2, 3, 4 Ternidens sp. 134 6.7 131 1.5 110 29.1 375 11.5 /, 2. 4 Trichostrongylus sp. 134 6.0 131 4.6 110 ?4.5^ 375 ?5.1 7, 2_ 4 Order Ascaridida Ascaris sp. 134 4.5 131 14.5 265 9.4 2, 6 Enterobiiis macoci 134 2.2 131 0 98 ?1.0^ 363 ?1.1 2, 4, 7 Order Spirurida Macacanema formosana 277 8.3 277 8.3 3, 8 Physaloptera twnefaciens 134 4.5 131 0 110 14.5 375 5.9 1, 2. 3, 4 Streptopharagiis piginentatits 134 2.2 131 0 110 16.4 375 5.6 1. 2, 3, 4 ' Excludes Ogmocotxle indica, a trematode cited as a nematode by Bergner (1967, p. 107; cf. Yoshimura et al., 1968. p.292: 1969. p.460). -Classification follows Maggenti (1983. p. 29). ''Key to references: /. Jessee et al. (1970. p. 87). 2. Hai-Yin Wu. Huey-Yu Lin, and Yao-Sung Lin (unpublished data: cf. Lin et al.. 1996. abstract no. 712; Wu & Lin. 1996. p. 24). 3. Bergner (1967. p. 107). 4. Kuntz et al. (1968. p. 124; cf. Myers & Kuntz. 1964. p. 864). 5. Pryor et al. ( 1970b. p. 1 120). 6. Lin et al. (1996. abstract no. 712). 7. Hugot (1999. p. 544). 8. Bergner and Jachowski (1968. p. 28: cf. Hsieh. 1961. p. 99; Schad & Anderson, 1963, p. 797; Kim & Bergner. 1964. p. 81; Kuntz et al.. 1968. p. 128). ""Cited as "Trichostrongylid." ' Cited as "Oxyurid." Leech Kuntz et al. (1968. p. 129) accidentally discov- ered leeches. Dinobdella ferox, in the nasal cavi- ties of at least three M. cyclopis skulls that were being prepared as museum specimens; no infor- mation is available concerning sample size. Sub- sequently, Pryor et al. (1970a, p. 1926) detected the same species of leech in the nasal cavities of 6 of 150 laboratory captives of M. cyclopis (4%) that had been acquired from animal brokers in various parts of Taiwan: in lots of 20-30 mon- keys, the incidence of leeches reportedly varied from 0% to 69c. Part of an unidentified leech was observed pro- truding from the nostril of a living adult male ma- caque at Renting, extreme southern Taiwan (H.- Y. Wu, pers. obs.). Lice Two species of Pedicinus have been detected on M. cyclopis specimens — P. eurygaster on 1 2- 15 specimens of M. cyclopis and P. obtusus on one specimen (Kuhn & Ludwig, 1967a, pp. 173, 252; 1967b, p. 266; Kuntz et al., 1968, p. 129). Viruses B Virus — Seven of 43 M. cyclopis individuals (16.3%) were positive for anti-B virus antibodies (Table 13); locality data are not available for the monkeys tested. In other fascicularis-group spe- cies, the incidence of positive samples varied from 0% to 34.99c. Reovirus — Of 15 M. cyclopis individuals test- ed, ten (66.7%) were positive for antibodies to at least one type of reovirus; the other five individ- uals were negative for antibodies to all three types of reovirus (Yang et al., 1976, p. 2). Simian Foamy Virus — Of 120 M. cyclopis samples tested, 37.5% were positive for either simian foamy virus type 1 or simian foamy virus type 2 (Table 14); in these M. cyclopis samples. 20 FIELDIANA: ZOOLOGY Table 13. Incidence of anti-B virus antibodies de- tected in foscicularis-gvoup macaques.' Positive Species N No. % Macaco fasciculahs- 86 30 34.9 Macaca cyclopis^ 43 7 16.3 Macaca mulatta'^ 6 0 0 Macaca fiiscata^ 61 7 11.5 Totals 196 44 22.4 ' Reference: Endo et al. (1959, p. 229). ' Cited as M. irus; origin of specimens: Thailand, Ma- laysia, Indonesia, and Philippines. "* Origin of specimens: "Formose" (^Taiwan). ■* Origin of specimens: Pakistan. ■ Origin of specimens: Yakushima, southern Japan. virus type 1 tended to be somewhat more frequent than virus type 2 (P < 0.025). No strong geo- graphic pattern is evident in the incidence of these virus types in Taiwanese counties. Simian Virus 40 — Infections with simian virus 40 (SV4()) were detected in 45 of 197 serum sam- ples (22.8%) collected from natural populations of M. cyclopis (Table 15). SV40 infections were much more common in samples from southern Taiwan (42/103 = 40.8%) than in those from northern and eastern Taiwan (1/89 = 1.1%). Natural History Habitats The primary habitat of M. cyclopis is broadleaf evergreen forest (Bergner & Jachowski, 1968, p. 11; Poirier & Davidson, 1979, p. 132; Masui et al., 1986, p. 390; Tanaka, 1986, p 100; Lee & Lin, 1994, p. 19). At upper elevations, as at Yushan (1730-2670 m), this species also inhabits mixed broadleaf-needleleaf forest (Lu et al., 1991, p. 91 ; Norikoshi, 1991, p. 138), and occasionally it may be found in needleleaf forest and bamboo forest (McCullough, 1974, p. 11; Poirier & Davidson, 1979, p. 131). Although M. cyclopis exploits all strata of the forest, from ground level to the crowns of the tallest trees, it is observed most frequently in lower forest strata (Kano, 1940, p. 63; Lu et al., 1991, p. 95). Swinhoe (in Blyth, 1860, p. 87; 1863, p. 350) misleadingly indicated that rocky, relatively tree- less areas were the preferred habitat of M. cyclo- pis. This misapprehension presumably was the re- Table 14. Incidence of simian foamy virus, type 1 and type 2, detected in samples of natural populations of Macaca cyclopis. ' Viral incidence (%) County N Type 1 Type 2 Total Taipei- Ilan^ Hualien Chia-I Taitung Pingtung Unknown Totals 3 12 74 2 7 6 16 120 33.3 16.7 -18.9 0 42.9 33.3 -56.3 -25.8 0 16.7 12.2 0 0 16.7 •12.5 •11.7 33.3 33.3 31.1 0 42.9 50.0 68.8 37.5 Reference: Johnston (1961, p. 4). Wulai vicinity. Ilan vicinity and Suao vicinity. suit of Swinhoe's observation of these monkeys at Shou Shan (= Ape's Hill), an uplifted coral reef formation in southwestern Taiwan, where mem- bers of the local population of M. cyclopis often rest on exposed rocky surfaces (Hai-Yin Wu, pers. obs.). Bergner (1967, p. 33) observed that many shallow caves on the west side of Shou Shan were stained with monkey urine, and he therefore in- ferred that M. cyclopis frequently enters these caves; M. cyclopis occasionally has been ob- served to enter a cave, apparently to keep cool, at Hengchun Nature Preserve, Kenting National Park, in extreme southern Taiwan (Wu & Lin, 1993, p. 250). The elevational range of 617 M. cyclopis lo- calities is 100-3400 m (Table 16; Gazetteer, Ap- pendix 2); approximately one-third of these lo- calities are at 1000-1500 m elevation, and only about 5% are above 2500 m. Generally similar elevational distributions have been reported by Kano (1940, p. 33), Poirier and Davidson (1979, p. 131), Masui et al. (1986, p. 390), Tanaka (1986, p. 100), and Lee and Lin (1994, p. 19); reported Table 15. Incidence of simian virus 40 infections detected in Macaca cyclopis. ' Sample area Counties N Infected No. % North Taipei, Ilan 50 1 2.0 East Hualien, Taitung 39 0 0 Central Nantou, Chia-1 5 2 40.0 South Pingtung 103 42 40.8 Totals 197 45 22.8 Reference: Yang et al. (1967, p. 146). FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 21 Table 16. Elevational distribution of Mwrara rvc7<>- pis locality records: for details see Gazetteer. Appendix 2. Records Elevation (m) No. ^r 0-499 73 11.8 500-999 105 17.0 1000-1499 207 33.6 1500-1999 113 18.3 2000-2499 88 14.3 2500-2999 27 4.4 3000-3499 4 0.6 Totals 617 100.0 crop raiding by M. cyciopis tends to occur at low- er elevations, usually below 1000 m (Chang, 2000. p. 3). The highest elevation at which M. cyciopis has been recorded (3400 m) is on one of the peaks in Chung Yang Shan Mo (= Central Mountain Range), where McCullough (1974. p. 1 1 ) observed monkey droppings in a bamboo grassland. Terrestrialit y/ A rborealit y Detailed information concerning the amount of time that M. cyciopis spends on the ground and in trees is available for only one population, ob- served Nov. 1987-Oct. 1988 in Yushan National Park, central Taiwan (Lu et al.. 1991. p. 95: cf. Kano. 1940. pp. 59. 62: Bergner. 1967. p. 33: McCullough. 1974. p. 10). In this population, the average proportion of time that monkeys were ob- served on the ground each hour between 0700 and 1900 varied from 09c to 50*^: the composite daily averages were 16.89^ terrestrial behavior and 83.2% arboreal behavior. Local circumstances apparently influence the degree of terrestriality or arboreality of M. cycio- pis. In well-habituated groups at Kenting. Shou Shan, and Fushan. members of this species are more likely to move on the ground when traveling in disturbed areas and more likely to move through the trees when traveling in mature forests (Hai-Yin Wu. pers. obs.). In northern Taiwan, ac- cording to Bergner (1967. p. 33). this species ha- bitually remains on the ground, except in response to danger, when it flees into the trees. McCullough (1974. p. 10). by contrast, indicates that M. cycio- pis may respond to danger by fleeing either on the ground or into the trees: no locality information is provided concerning this variation. Although M. cyciopis probably usually sleeps in trees, sleeping on the ground and in caves also has been reported (Bergner. 1967. p. 33; Poirier & Davidson, 1979. p. 164). This species report- edly swims well and sometimes swims in the ocean (Imaizumi. 1970. p. 287: Poirier & David- son. 1979. p. 165: cf. Azuma & Norikoshi. 1990. p. 47). Group Size and Composition In an island-wide survey, Lee and Lin (1991, p. 34) found that most M. cyciopis groups were relatively small. Group size was 2-10 individuals in 57. 99^ of 121 groups reported. 11-20 individ- uals in 19.89^ of the groups, and >20 individuals in 223% of the groups (cf. Kano. 1929, p. 333 Kuroda. 1940, pr 270: Imaizumi, 1970. p. 287 McCullough. 1974. p. 10: Tanaka. 1986, p. 101) a rough estimate of mean size in these 1 2 1 groups is 12.2 individuals per group. Thirteen solitary monkeys — presumably males — also were report- ed by Lee and Lin (1991, p. 34: cf. Poirier & Davidson, 1979, p. 158: Masui et al.. 1986, p. 391: Lu et al.. 1991. p. 94: Hsu & Agoramoorthy. 1999, p. 64). Larger groups in this survey tended to be restricted to elevations below 1500 m: above 3000 m, only small groups (2-5 individuals) and solitary monkeys were reported (cf. Norikoshi, 1991, p. 140). All-male groups, including adults and juveniles in the same group, have been observed at Taip- ingshan. northeastern Taiwan (Kawamura et al., 1988. p. 3: 1991. p. 99: Azuma & Norikoshi. 1990. p. 47). and Kenting, extreme southern Tai- wan (Jon-Ne Yo, pers. comm.). At both localities, the all-male groups were loosely associated with neighboring mixed-sexed groups (cf. Hsu & Lin, 2o6\. p. 55). Where group size has been reported for speci- fied localities (Table 17). it is relatively low at Kenting and in the introduced population on Osh- ima (island). Japan. At Kenting. a group that had increased from 10 individuals in 1985 to 19 in- dividuals in 1987 was observed in 1987 to split into two smaller daughter groups (Wu et al.. 1991. p. 30: Wu & Lin. 1992. p^ 88): similar group fis- sion has been reported at Shou Shan (Hsu & Ago- ramoorthy. 1999. p. 64: Hsu & Lin. 2001. pp. 53. 56). Although local informants have reported groups of > 100 individuals (Lee & Lin. 1991. p. 35) and even 500 individuals (Poirier & David- son. 1979. p. 157). the largest verified group sizes 22 FIELDIANA: ZOOLOGY Table 17. Geographic variation in group size in Macaco cyclopis populations. Locality Group size N County Mean Minimum Maximum References' Taipei Hsi Lin 30 ! / Taipei/Ilan Fushan Forest Station ? ? >25 5 2 Ilan Fushan 23 21 25 2 3 Taipingshan 20.1 12 27 6 4 Changhua Pakua Shan -15 — — 1 1 Nantou Chichi Tashan 75 — — 1 5 Nantou/Chia-I Yushan National Park^ -29 14 43 7 6 Kaohsiung Shanping 20 — — 1 1 Shou Shan (1963) 24 — — 1 7 Shou Shan (1995- 1997) 46.8 9 86 13 8 Taitung Kuanshan -30 -12 <50 2 1 Chihpen Hsi >45 — — 1 9 Pingtung Wanlung Shan -15 — — 1 1 Kenting (1985) 10 — — 1 10 Renting (1990) 14.5 13 16 2 10 Kenting (1997) 27 22 32 2 11 Introduced feral population (Japan) Oshima 12.4 6 27 7 12 ' Key to references: 7. Poirier and Davidson (1979, p. 157). 2. Lee (1994, p. 438). 3. K.-Y. Chang (1999, pp. 7- 8). 4. Azuma and Norikoshi (1990, p. 47); Kawamura et al. (1988, table 1; 1991, p. 99; cf. Poirier & Davidson, 1979, p. 157). 5. S.-W. Chang (1999, p. 13). 6. Hai-Yin Wu (pers. obs., 1991-1996). 7. Bergner (1967, p. 33). 8. Hsu and Agoramoorthy (1999, p. 64); Hsu and Lin (2001, p. 52, Aug. 1997 census). 9. Masui et al. (1986, p. 391). 10. Wu and Lin (1992, p. 88; cf. Poirier & Davidson, 1979, p. 157). 11. W. Birky (Rutgers University, pers. comm., 26 Apr. 2000). 12. Kazama and Norikoshi (1991, p. 54). - Cf. Lu et al. (1991, p. 94), whose group-size estimates for a nearby area in Yushan National Park are mean = 7.8; minimum < 5; maximum = 14; n ^ 11. are 86 individuals at Shou Shan and 75 individ- uals at Chichi Tashan; both of these large groups inhabit isolated relict forest patches surrounded by broad expanses of unsuitable agricultural and ur- ban areas. In eight natural populations of M. cyclopis, the pooled ratio of sexually mature males to sexually mature females varies from 0.15 to 0.55 (Table 18). The composite pooled sex ratio in these eight populations — with the Kenting population repre- sented by the 1996-1998 sample— is 0.49 (172 sexually mature males, 349 sexually mature fe- males). The pooled sex ratio of the introduced population in Oshima exceeds that of all eight nat- ural populations. Home Range Area and Population Density Mean home range area in six natural popula- tions of M. cyclopis is approximately 130 ha, and mean population density is approximately 0.25 in- dividuals per ha (Table 19); home range area and population density in the isolated Chichi Tashan group (37 ha, 2.03 individuals per hectare) are aberrant. Home ranges of neighboring groups may partly overlap (Kawamura et al., 1988, p. 2; 1991, p. 98). In the introduced Oshima population, mean home range area is approximately one-fourth that in natural populations, and population density is approximately twice as great as in natural popu- lations (Table 19); Azuma and Norikoshi (1990, p. 53) suggest that heavy hunting pressure in for- mer years in Taiwan may be partly responsible for these differences. Poirier and Davidson (1979, pp. 137, 173) sug- gested that groups of M. cyclopis that inhabit up- per elevations may shift their home ranges up- ward in summer and downward in winter. How- ever, Kawamura et al. (1991, p. 98) detected no such shifting in this species at Taipingshan during a period of frequent snowfall at 1000 m. Local seasonal movements probably occur in lowland populations as well as in upland populations (Hai- Yin Wu, pers. obs.). Diet Although leaves and other vegetative plant parts are included in the diet of M. cyclopis, fruits FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 23 Table 18. Ratio of sexually mature males to sexually mature females reported in Mucaca cyclopis groups. No. of No. of sexually mature Group sex ratios Pooled Refer- County Locality groups individuals Minimum Maximum sex ratio ences' Ilan Fushan 2 17 0.13 0.33 0.21 / Taipingshan 4- 32 0.33 1.50 0.52 2 Nantou Chichi Tashan 1 23 — — 0.44 3 Nantou/Chia-I Yushan N. P. 5 66 0.29 0.54 0.40 4 Kaohsiung Yushan N. R 6^ 19 0.33 0.50 0.46 5 Shou Shan 16 328 0.30 0.82 0.55 6 Taitung Chihpen Hsi 1 15 — — 0.15 7 Pingtung Kenting (1985) 1 4 — — 0.33 8 Renting (1990) 2 9 0.25 0.33 0.29 8 Kenting (1996-1998) 2 21 0.50 0.50 0.50 9 Kenting (1999) 1 11 — — 0.38 10 Introduced feral population (Japan) Oshima 6 21 0.33 1.00 0.75 11 ' Key to references: /. K.-Y.Chang (1999, p. 7). 2. Azuma and Norikoshi (1990. p. 47); Kawamura et al. (1988, table 1; 1991. p. 99). 3. S.-W. Chang (1999. p. 13). 4. Hai-Yin Wu (pers. obs.. 1992-1994). 5. Lu et al. (1991, p. 94). 6. Hsu et al. (2000b. p.201); cf. Hsu and Lin (2001. p. 53). 7. Masui et al. (1986, p. 391). 8. Wu and Lin (1992, p. 88). 9. W. Birky (Rutgers University, pers. comm.. 26 Apr. 2000). 10. Jon-Ne Yo (ntuz, pers. comm.). 11. Kazama and Norikoshi (1991. p. 54). - Excludes two groups in which sex was not determined for all adults. ' Excludes four groups in which sex was not determined for all adults. and seeds are the primary food of this species (Table 20); known plant food sources include trees, shrubs, and herbs (Lee, 1991, p. 297). Lo- cally, food sources include 38 plant species iden- tified at Yushan National Park (Wu & Lin, 1996, p. 24), 51 species at Jentse (Su, 1993, p. 16), 55 species at Kenting (Yo, 2000, p. 18), and 132 spe- cies at Fushan (K.-Y Chang, 1999, p. 17; Chen, pp. 70-73). In addition to wild plants, at least 33 species of cultivated crops are opportunistically Table 19. Home range area and population density reported in Macaca cyclopis. Mean home Mean population No. of No. of range area density Refer- County Locality groups individuals (ha) (ind./ha) ences' Ilan Fushan 1 21 97 0.22 1 Taipingshan 5 119 140 0.17 2 Nantou Chichi Tashan 1 75 37 2.03 3 Nantou/Chia-1 Yushan N. R 7 -200 -86 -0.33 4 Kaohsiung Shou Shan 7-15 171-652 167 0.26 5 P -40 -120 -0.33 6 Taitung Chihpen 1 >45 -200 -0.22 7 various 39 ■7 '1 (-0.001 groups/ha) 7 Pingtung Kenting 1 -30 -100^ -0.30 8 Introduced feral population (Japan) Oshima 6-8 -100 -30 0.48^ 9 ' Key to references: /. K.-Y.Chang (1999, p. 27). 2. Kawamura et al. (1988. p. 2); Azuma and Norikoshi (1990. p. 44). 3. S.-W. Chang (1999, p. 13). 4. Hai-Yin Wu (pers. obs., 1992-1994). 5. Hsu and Agoramoorthy (1999, p. 64); Hsu and Lin (2001. pp. 51, 53). 6. Shu-Mei Chen (ntuz. pers. comm., 17 Apr. 2000). 7. Masui et al. (1986. pp. 390, 391). 8. Jon-Ne Yo (ntuz. pers. comm). 9. Kazama and Norikoshi (1991, p. 54); Noriko.shi (1991, p. 140). - Provisioned group. 3 Underestimated as -20 ha by Wu and Lin (1993. p. 250). ^ 0.26 in less favorable part of study area. 0.74 in more favorable part of study area. 24 FIELDIANA: ZOOLOGY Table 20. Foods reported eaten by Macaco cyclopis. Locality Fruits, berries, seeds Shoots, leaves, stems Cultivated Crustaceans, plants insects Mollusks Vertebrates Other References' Fushan X X x2 Fushan Jentse X X X X X X Taipingshan X x5 MiaoH-Nantou'^ x' Yushan N. P. X X X Shou Shan X X X X8 Kenting X X X9 Renting X X X2 x'o x' Unspecified X X X Xl3 x X X X x''* x' X X X X X X X Xl6 X3 1 x* 2 3 4 5 6, 7 7 8 X'2 9 10 11 12 x'7 13 14 15 (Japan) Introduced feral population 16 ' Key to references: 1. K.-Y. Chang (1999, pp. 17-20). 2. Chen (1999, pp. 13-15). 3. Su (1993, p. 10). 4. Kawamura et al. (1988, pp. 3, 4; 1991, p. 98); Azuma and Norikoshi (1990, p. 44). 5. Chang (2000, p. 5). 6. Wu and Lin (1996, p. 24). 7. Hai-Yin Wu (pers. obs.). 8. Wu and Lin (1992, p. 86; 1993, p. 249). 9. Yo (2000, p. 18-21). 10. Swinhoe (1863, p. 351). //. Bergner (1967, p. 35). 12. McCullough (1974, p. 11). 13. Poirier and Davidson (1979, pp. 147, 159). 14. Lee and Lin (1991, p. 35). 15. Hsu and Agoramoorthy (1997, p. 835). 16. Shirai (1988, p. 22); Kazama and Norikoshi (1991, p. 53). ^ Insects, wasp nests. 3 Gum, plant galls. ■• Clay, tree bark, plant galls. ' Insects. * Includes five counties: Miaoli, Taichung, Changhua, Yunlin, Nantou. ^ Thirty-three species listed. ** Insects, termite mound. ' Cicadas, caterpillars. '0 Snails. " Frog tadpoles (in tree hollow), birds eggs. '2 Clay. " Crustaceans, grasshoppers. '■* Crayfish, cockroaches, flies. '"^ Lizards. '^ Crustaceans. '^ Mushrooms, lichens, tree bark. exploited (Chang, 2000, p. 5). The diet of the in- troduced feral population of M. cyclopis in Oshi- ma (island), Japan, apparently is similar to that of natural populations (Norikoshi, 1991, p. 140). Reported animal food consumed by M. cyclopis includes crayfish, grasshoppers, cockroaches, ter- mite mounds, cicadas, beetles, caterpillars, flies, wasp nests, snails, tadpoles, lizards, and birds eggs (Table 20). In captivity, fish and beef are eaten (Yang et al., 1968, p. 538; Peng et al., 1973, p. 202). Like M. mulatta (Fooden, 2000, p. 59), natural populations of M. cyclopis have been ob- served to eat clay. The peak feeding time for M. cyclopis is in the early morning, shortly after a group awakens, and a secondary feeding peak occurs in the afternoon (Bergner, 1967, p. 34; Poirier & Davidson, 1979, p. 161; Lu et al., 1991, p. 96; Wu & Lin, 1993, p. 245). The recorded proportion of daylight hours devoted to foraging was 30-35% at Hengchun Nature Preserve, Kenting National Park (Wu & Lin, 1993, p. 246) and approximately 60% at Yu- shan National Park (Lu et al., 1991, p. 96). Although Poirier and Davidson (1979, p. 162) postulate that plant foods supply much of the wa- ter requirements of M. cyclopis, they report that a monkey sometimes will lick water from tree trunks or branches and from its own body or the FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 25 body of another monkey. This species also has been observed to lick water droplets from wet rocks and to drink from streams, ponds, and tree holes (Hai-Yin Wu, pers. obs.). Food washing by captive M. cyclopis has been reported by Tateishi (1958, p. 90). Predators The two reported predators of M. cyclopis are an eagle (Poirier & Davidson, 1979, p. 151) — probably the hawk eagle, Spizaetus nipalensis — and the clouded leopard, Neofelis nebulosa (Kano, 1930, p. 165; Imaizumi, 1970, p. 287; Mc- CuUough, 1974, p. 16; cf. Wozencraft, 1993, p. 297). An encounter between M. cyclopis and S. nipalensis, which is the largest bird of prey in Taiwan, has been observed in Kenting National Park (Hai-Yin Wu, pers. obs.): when an eagle flew into the tree in which a group of monkeys was resting, the monkeys sounded an alarm call and moved out of the tree; subsequently, an adult male monkey returned to the tree and shook a branch opposite the eagle's perch. Human predation on M. cyclopis apparently has declined during the last two or three decades, primarily as a result of recent conservation legislation (Steere, 1876. p. 310; Campbell, 1903, p. 551; Kano, 1929, p. 333; 1930, p. 165; McCuUough, 1974. p. 1 1; Poirier & Davidson, 1979, p. 176; Lee & Lin. 1991, p. 36; Agoramoorthy, 1998, p. 73). Intergroup Dispersal At Kenting National Park, during a five-year period (1985-1990), three adult males and one subadult male emigrated from two M. cyclopis study groups, of which the total population varied from 10 to 29 individuals (Wu & Lin, 1992, p. 92). During the same period, four adult males im- migrated into these two study groups; although one of the immigrant males had been artificially released into the area by local residents, the other three immigrants presumably came from neigh- boring groups without human intervention. Group tenure of five adult males in the study groups var- ied from 5 months to >28 months; mean group tenure was >17.5 months. No direct interaction was observed between the two study groups and three neighboring groups (Wu & Lin, 1992, p. 87). At Shou Shan, during a 2.75-year period (1995-1997), males (n = 54) reportedly emigrat- ed from their natal groups at age 5-6 years (Hsu & Agoramoorthy, 1999, p. 64; Hsu & Lin, 2001, p. 55); subsequently, these males either joined a neighboring group or became solitary. The mean tenure of 34 males as alpha dominant individuals in Shou Shan groups was 16.8 ± 18.9 months (extremes 1 week-6 years). A study of mtDNA in M. cyclopis groups at Shou Shan indicated that several pairs of males with the same haplotype had dispersed into resi- dent groups with a different haplotype (Jui-Hua Chu, pers. comm., 15 Apr. 2000; cf. Wu et al., 2001b, p. 414); however, it cannot be ascertained whether or not individuals in the dispersing pairs came from the same natal group. All-male groups that have been observed at Taipingshan and Kent- ing (see above. Group Size and Composition) probably were temporary aggregations of dispers- ing males. Females of M. cyclopis apparently usually re- main in their natal groups. However, during the 1995-1997 study period at Shou Shan, six adult females were observed to successfully immigrate into new groups (Hsu & Lin, 2001, p. 55). Reproduction Seasonality Reproduction is strongly seasonal in M. cyclo- pis (Table 21). In natural populations of this spe- cies, copulation frequency peaks during the period November-January and birth frequency peaks during the period March-June. Copulations may occur at a low frequency outside of the peak mat- ing season, but these probably are mostly infertile; births apparently are rare or absent during the pe- riod September-January. Climatologically, the peak mating season generally is a period of de- creasing temperature and rainfall, and, conversely, the peak birth season generally is a period of in- creasing temperature and rainfall (Wu & Lin, 1992, p. 86; Hsu et al., 2001, p. 17). In laboratory captives, annual mating and birth peaks apparent- ly are attenuated but not extinguished. Sexual Skin Cyclical changes of the sexual skin, correlated with ovulatory cycles, are conspicuous in M. cy- 26 FIELDIANA: ZOOLOGY Table 21. Observations concerning reproductive seasonality in Macaca cydopis.^ Sample area Observation period Mating/birth observations References^ Matings Free-ranging groups Shou Shan Jan. 1995-Oct. 1997 Most successful matings: Sep.-Feb. 1 Kenting Oct. 1987-Apr. 1990 304 copulations observed: Nov.-Jan., 274; Feb.-Oct., 30 2 Taiwan 7 Uncertain, usually Oct.-Nov. 3 Nojima I., Japan Aug. & Oct. 1962 6 copulations reported, Aug. and/or 4 Oct. Captive groups Ueno Zoo. Japan ca. 1958 Summer amennorhea observed, Jun.-Aug. 5 Taipei, lab 1966-1972 13 pregnancies, Aug.-Jun. 6 Taipei Zoo Sep. 1989-Mar. 1990 970 copulations observed, most Dec.-Feb. 7 Taipei Zoo ca. 1995, 1 yr Sperm count peaks Oct.-Dec.-* Births 8 Free-ranging groups Taipingshan ? May-Jun 9 NE Taiwan ? Jun. 10 E Taiwan ? Mar.-Oct. 9 S Taiwan ? Mar.-Sep. 9 Shou Shan Jul. 1963 6 births, Jun. or Jul. 11 Shou Shan 1996-1999 479 births, Feb.-Aug. 12 Kenting Mar. 1985-Aug . 1990 28 infants observed: 25 born Mar. -Jun.; 2 born Feb.; 1 born Aug. 2 Taiwan 1965-1969 48 births,* Mar-Jun. 6 Taiwan 7 Mar. -May 13 Captive groups Kurabayashi Zoo, Japan 1954-1958 7 births. Mar. -Jun. 14 Taipei, lab 1965-ca. 1968 54 births,"^ Mar-Jul. 15 U.S.A., lab 1967-1992 143 infants: 59 born Mar.-May; 84 born Jun.-Feb. 16 Taipei Zoo 1989-1990 25 births, Apr.-Jun.; 1 birth, Jul.; 1 birth, Aug. 17 ' Cf. Kano (1929, p. 333), who indicates that matings and births are nonseasonal in M. cyclopis. 2 Key to references: /. Hsu and Agoramoorthy (1999, p. 64); Hsu et al. (2001. p. 18); Hsu and Lin (2001. p. 54). 2. Wu and Lin (1992, pp. 89, 90). 3. Dien (1958, p. 346). 4. Ni.shida (1963, p. 121). 5. A.sakura (1958, p. 162). 6. Peng et al. (1973, pp. 206, 209). 7. Wu (1996, abstract no. 117). 8. Chi et al. (1998, p. 34). 9. Poirier and Davidson (1979, p. 153). 10. Swinhoe (1863, p. 351). //. Bergner (1967, p. 33). 12. Hsu et al. (20()0a, p. 154; 2001, p. 18); Hsu and Lin (2001, p. 54). 13. McCullough (1974, p. 10). 14. Tateishi (1958, p. 89). 15. Yang et al. (1968, p. 540). 16. Petto et al. (1995, p. 339). 17. Chen et al. (1990, p. 24). -* Studied by electroejaculation. ** Wild-trapped pregnant females; infants born in two Taipei laboratories. ^ Probably includes some or all of 48 births cited above (see Taiwan, 1965-1969). clopis females (Sclater, 1865, p. 710; Murie, 1873, p. 772; Hsu, 1990, pp. 52, 60; cf. Nunn, 1999, pp. 232, 245; van Schaik et al., 1999, pp. 219, 234). The affected area extends broadly over the hindquarters from the root of the tail to the back of the thighs, almost as far as the popliteal fossae. This area becomes prominently red and swollen in subadult and young adult females; in older females, the sexual skin undergoes cyclical changes in redness but not in tumescence (Hai- Yin Wu, pers. obs.). In three young females in a carefully studied natural population, the first signs of reddening and swelling of the sexual skin were detected at age 2.5-2.75 years (Wu & Lin, 1992, p. 89); approximately 1 year later, during the mat- ing season, the sexual skin in these females was prominently developed. The timing of sexual skin development follows a similar course in labora- tory captives, in which it has been further noted that a female's first menstruation (menarche) oc- FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 27 Table 22. Menstrual c\cle duration in captive Macaca cyclopis. Location of Menstrual cycle duration N colony Mean (days) Extremes (days) Females Cycles References' Determined by observation of menstrual bleeding Taipei, lab U.S.A.. lab 29.4 = 15.5 (SD) 6-109 54 27.9 = 0.5 (SE) <22->40 31 398 555 I 2 Determined by hormonal assay and/or laparoscopic examination Taipei Zoo Taipei Zoo 27.8 = 2.9 (?SE) — 3 27.3 r 8.9 (SD) 22-35 6 8 18 3 4 ' Kev to references: 1. Peng et al. (1973. p. 203). 2. Petto et al. (1995. p. 340). 3. Chen and Lin (1991. p. 42). 4. Chi (1997. p. 43). curs a few months before or after onset of her sexual skin development (Peng et al.. 1973. p. 210: Petto et al.. 1995. p. 344).^ The relationship between color changes of the sexual skin and stages of the menstrual cycle has been studied in older M. cyclopis females at the Taipei Zoo by Chi (1997. p. 45). The sexual skin is pink during the period of menstrual bleeding; it becomes red by day 6 of the cycle and is deep reddish purple by day 14 (typical day of ovula- tion): after ovulation, the color rapidly fades and is pink by day 18 and pale pink by day 23 (4 days before onset of the next menstrual bleeding). The annual cycle of sexual skin changes was monitored during 1997-1998 in 15 adult females in a natural group at Shou Shan (Shu-Mei Chen. NTUZ. pers. comm.. 17 Apr. 20(X)). In this group, color changes were restricted to the period from mid-October to late January. The number of col- or-change cycles per adult female varied from zero to five, and the duration of the color-change cycles varied from 10 days to 48 days. In males, an annual change in the color of facial skin and scrotal skin has been noted at Shou Shan by Shu-Mei Chen (ntuz. pers. comm.. 17 Apr. 2000). During the mating season, the facial and scrotal skin becomes more intensely red: this is particularly evident in the seasonal appearance in males of a pair of narrow red streaks that extend laterally from the outer comer of each eye (cf. Fooden. 1975. p. 114). Polythelia Supernumerary nipples and/or areolae have been observed in 89 of 21 1 adult females (42. 29^) and in 20 of 1 17 adult males (17.19^) in 16 groups of M. cyclopis that were studied at Shou Shan. southwestern Taiwan (Hsu et al.. 2000b. p. 201; Wu et al.. 2001a. p. 387). The frequency of su- pernumerary nipples apparently is greater in fe- males that produced twins than in females that produced singletons. Possible causes suggested for the high incidence of polythelia in this isolated population are inbreeding and/or artificial provi- sioning. Menstrual Cycles In two large studies of captive M. cyclopis, based on observations of menstrual bleeding, mean length of the menstrual cycle was deter- mined as 29.4 days and 27.9 days (Table 22): the discrepancy between these mean values may be related to the authors" divergent treatment of out- lier cycle lengths and summer amenorrhea. In two smaller studies, based on hormonal assays and laparoscopic examinations, mean menstrual cycle length was determined as 27.8 days and 27.3 days. Mean duration of the menstrual bleeding pe- riod was determined as 4.2 ± 3.2 (SD) days (ex- tremes. 1-17 days, n = 44 periods) in one of the larger studies (Peng et al.. 1973. p. 204) and as 3.4 ± 0.6 (SD) in one of the smaller studies (Chi, 1997. p. 43). The same small study also deter- mined that ovulation occurs two days after the peak concentration of circulating estradiol or 14.1 ± 1.3 (SD) days after the first day of menstrual bleeding. In 15 adult females studied during the 1997- 1998 breeding season at Shou Shan, the frequency of menstrual bleeding cycles varied from zero per female to two per female (Shu-Mei Chen, ntuz, pers. comm.. 17 Apr. 2000). Of 20 menstrual cy- cles observed, five cycles (of five females) oc- curred during the period early August to mid-Oc- 28 FIELDIANA: ZOOLOGY tober and 15 occurred during the period mid-Oc- tober to early January. Sexual Maturation In natural populations, females generally pro- duce their first young either at age 4 years or at age 5 years (Wu & Lin, 1992, p. 89; Hsu & Ago- ramoorthy, 1999, p. 64; Hsu et al., 2001, p. 20; Hsu & Lin, 2001, p. 54), which indicates that their first fertile copulations occur at age ca. 3.5 years or ca. 4.5 years. The age of sexual maturation of males in natural populations has not been report- ed, but it probably is at least 5-6 years, at which age males emigrate from their natal troops (Hsu & Agoramoorthy, 1999, p. 64). Although the mean age of sexual maturation in captive females and males apparently is similar to that in natural populations (Peng et al., 1973, p. 210; Petto et al., 1995, p. 336), semen with normal sperm count has been obtained by electroejaculation of males weighing as little as 5.0 kg (age <4.0 years) (Peng et al., 1973, p. 211). In one laboratory colony, the mean age at first reproduction tended to increase in successive generations of captives, reaching 6.75 years in the third captive generation of fe- males and 9.0 years in the third captive generation of males (Petto et al., 1995, p. 336). Copulation The copulatory posture of M. cyclopis (Hsu, 1990, pp. 50, 55) apparently is similar to that of M. mulatta (Fooden, 2000, p. 72); copulations in M. cyclopis occur both on the ground and in trees. In the Taipei Zoo population of M. cyclopis, both single-mount ejaculation (SME) and multimount ejaculation (MME) have been observed (Wu, 1996, abstract no. 1 17); in dominant males, MME tends to be more frequent than SSE. In another captive group, copulation was often preceded by rhythmic lip movements performed with male and female facing one another (Akitsu, 1959, p. 85). During the peak mating season, nontroop males have been observed to approach a troop and to copulate with troop females (Wu & Lin, 1992, p. 93; Hsu & Lin, 2001, pp. 51, 55); on at least three occasions during the mating season, nontroop males have ascended to dominant status in troops that they entered (Wu et al., 1991, p. 31; cf. Hsu & Agoramoorthy, 1999, p. 64). Conversely, some subordinate females at Yushan and Shou Shan have been observed to spend several days during the mating season away from their troops in the company of nontroop males (Hai-Yin Wu, pers. obs.; Shu-Mei Chen, ntuz, pers. comm., 17 Apr. 2000). At the Taipei Zoo, DNA testing revealed that all three juveniles present in a captive group of M. cyclopis were sired by the dominant male (Chu et al., 1999, p. 376); in addition to the dom- inant male, this group included two subordinate adult males. In an introduced M. cyclopis population in Ja- pan, copulations between members of neighboring troops were observed six times in 12 days (Nish- ida, 1963, p. 121). In the same introduced popu- lation in Japan, female activity during estrus has been intensively studied by Kawai and Mito (1973, pp. 183, 185): estrous females become ac- tive 1 hour earlier than nonestrous females, and, during the day, the frequency of movement of es- trous females was approximately twice that of nonestrous females. Gestation and Parturition Mean length of the gestation period was 162.0 ± 9.6 (SD) days in nine laboratory colony preg- nancies (Peng et al., 1973, p. 208; cf. Hsu et al., 2001, pp. 17, 18); extreme lengths were 142 days and 175 days. Of 54 laboratory births, 40 (74%) occurred between the hours of 1700 and 0800 (Yang et al., 1968, p. 540). One parturition was observed in the field at Shou Shan on 16 Jun. 1998 (Shu-Mei Chen, ntuz, pers. comm., 17 Apr. 2000). Contractions of the mother were first detected at 1535, and the infant was born at 1646; during the entire interval, the mother was on the ground. Following birth of the infant, the mother ate the placenta. During the same birth season at Shou Shan, four additional infants apparently were born at night; all four of these infants, which were first seen early in the morning, were born to mothers who had been ob- served to be pregnant the preceding evening. Reproductive Rate The known reproductive rate in natural popu- lations of M. cyclopis varies from 0.45 to 0.95 births per sexually mature female per year (Table 23); although data for these populations are not directly comparable, the mean annual reproduc- tive rate probably is approximately 0.70. The FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 29 Table 23. Annual reproductive rate (number of births/number of sexually mature females) in natural populations of Macaca cyciopis. Annual Observation No. of No. of reproductive County Locality period groups individuals rate References' llan Taipingshan 1987-1989 1 ca. 25 0.45 / Kaohsiung Shou Shan 1995 7 171 0.47 2 1996 7-15 171-652 0.70 1997 15 652 0.83 Pingtung Kenting 1985-1987 1 10-14 0.95 3 1987-1990 2 9-16 0.7 P 1996 1 18-30 0.67 4 1997 1 18-30 0.67 1998 1 18-30 0.67 5 1999 1 18-30 0.75 Mean 0.69 ' Key to references: /. Azuma and Norikoshi (1990, p. 44); Norikoshi (1991, p. 136). 2. Hsu and Agoramoorthy (1999, p. 64); Hsu et al. (2001, p. 18); Hsu & Lin (2001. p. 54). 3. Wu and Lin (1992, p. 89). 4. W. Birky (Rutgers University, pers. comm., 26 Apr 2000). 5. Jon-Ne Yo (ntuz, pers. comm.). -0.44 in one group (4 infants/9 mature female-years); 0.92 in second group (11 infants/12 mature female-years). mean interbirth interval for eight sexually mature females at Kenting was 15.4 months (extremes 10.5-24.0 months, n = 19 intervals; Wu & Lin, 1992, p. 91), and that for 144 sexually mature females at Shou Shan was 13.4 months (extremes 8.9-34.2 months, 288 intervals; Hsu et al., 2001, p. 19). In a laboratory colony, the mean interbirth interval was ca. 20 months (n = 108 intervals) (Petto et al., 1995, p. 337). At Yushan National Park, during 1992-1993, the mean proportion of adult females with infants was only ca. 0.39 in six groups ranging in size from 14 to 36 members (Hai-Yin Wu, pers. obs.); proportions in individual groups were 0.20, 0.31, 0.36, <0.43, 0.50, and 0.60. It is unknown wheth- er this low mean infant/adult female proportion is the result of a low birth rate or of a low infant survival rate. Neonatal Sex Ratio, Twinning, Birth Weight Neonatal sex ratios are available for two natural populations of M. cyciopis (Table 24). Although these ratios appear to vary annually and locally, the composite ratio is 1.19 males: 1.00 females, which is not significantly different from 1 : 1 (X^ = 0.545, P > 0.10). Six pairs of twins were included among 596 infants born in 7-16 groups observed during five birth seasons at Shou Shan, southwestern Taiwan (twinning rate = 1.0%) (Hsu & Agoramoorthy, 1999, p. 64; Hsu et al., 2000a, p. 155; 2000b, p. 203). Both members of two pairs of twins sur- vived for more than 3 years; one member of two other pairs died within 30 days of birth; both members of one pair died within 7 days of birth; and one member of one pair was aborted ca. 8 weeks before live birth of the second member. As previously indicated, mean birth weight was 402 g in 89 captives born in Taipei and 430 g in 22 captives born in the U.S.A. (Table 2). Al- though overall mean values were identical in two studies of captives in Taipei, one study indicated that female neonates are slightly heavier than male neonates and the other indicated the con- verse. Infant Mortality At Shou Shan, 105 of 475 infants (22.1%) born during the period Jan. 1996-Dec. 1999 died be- fore reaching age 1 year (Hsu et al., 2001, p. 21; cf. Hsu & Agoramoorthy, 1999, p. 64). Of these 105 infant deaths, 32 (30.5%) were stillbirths, 16 ( 1 5.2%) occurred during the first week of life, and 18 (17.1%) occurred between age 1 week and 1 month. Two deaths at ages 42 days and 59 days were the result of accidental falls from trees, and seven infant deaths may have been the result of infanticide by immigrant males (Hsu & Agora- moorthy, 1999, p. 64). At Kenting during the period 1985-1990, 20% of infants in one to two groups died before reach- ing age 1.5 years (Wu & Lin, 1992, p. 92). At 30 FIELDIANA: ZOOLOGY Table 24. Neonatal sex ratio in natural populations of Macaca cyclopis. County No. of infants Locality Observation period Male Female M/F ratio References' Kaohsiung Pingtung Totals Shou Shan 1995 3 1996 4 1997 2 1998 9 Shou Shan, totals 1995-1998 18 Renting 1985-1990 12 1996 2 1997 3 1998 0 1999 3 Kenting, totals 1985-1999 20 38 2 1 4 1 8 14 2 1 4 3 24 32 1.50 4.00 0.50 9.00 2.25 0.86 1.00 3.00 0.00 1.00 0.83 1.19 ' Key to references: 7. Shu-Mei Chen (ntuz, pers. comm., 17 Apr. 2000). 2. Wu and Lin (1992, p. 89). 3. W. Birky (Rutgers University, pers. comm, 26 Apr. 2000). 4. Jon-Ne Yo (ntuz, pers. comm.). Taipingshan, by contrast, all four infants born in one group in 1987 survived to at least age 1 year (Azuma & Norikoshi, 1990, p. 54; Kawamura et al., 1988, p. 8; 1991, p. 99). In laboratories in Taipei, 22 abortions or still- births occurred among 72 wild-trapped pregnant captives (31%) and two abortions or stillbirths oc- curred among 12 laboratory-bred captives (17%) (Peng et al., 1973, p. 210). In a U.S. laboratory, 32 of 173 pregnancies resulted in stillbirths (18%), and five infants (3%) died within 30 days after birth (Johnson et al., 1986, p. 773; cf. Price et al., 1972, p. 307). Nursing; Longevity In natural populations of M. cyclopis, females apparently nurse their infants for 6-12 months (Bergner, 1967, p. 35). The greatest age reported to have been reached by an individual of this spe- cies— a captive male at Uchide Park, Ashiya City, Hyogo Prefecture, Japan — is 39 years (Anony- mous, 2000, p. 9). Population Growth Rate In seven groups studied at Shou Shan, south- western Taiwan, the average annual population growth rate from Dec. 1994 to Dec. 1997 was 24.4% (Hsu & Lin, 2001, p. 53). During the course of this study, the annual growth rate de- clined from 29.0% in 1994-1995 to 18.7% in 1996-1997. Fossils Fossils identified as Macaca sp. have been col- lected, associated with numerous other fossil ver- tebrates, in the vicinity of Tsochen (= Chochen), Tainan Hsien {- county), southwestern Taiwan (Otsuka, 1984, pp. 37, 42, 44). The age of the Tsochen vertebrate fossil fauna is estimated to ex- tend from Early Pleistocene to Middle Pleistocene (ca. 1.2-0.3 Ma) (Otsuka, 1984, pp. 50, 51; Qi et al., 1999, p. 39; Chen, 2000, p. 238); the fauna of the beds (KU4, KU5) that have yielded the monkey fossils belongs to the later part of this interval (Otsuka, 1984, p. 44; Lai, 1989, p. 45). Other families of terrestrial mammals that have been collected in the vicinity of Tsochen include Stegodontidae (5 spp.), Elephantidae (4 spp.), Fe- lidae (1 sp.), Hyaenidae (1 sp.), Rhinocerotidae (1 sp.), Tapiridae (1 sp.), Equidae (1 sp.), Hippopo- tamidae (1 sp.), Suidae (2 spp.), Cervidae (9 spp.), and Bovidae (4 spp.) (Lai, 1989, p. 39). Another iinportant Taiwanese fossil vertebrate fauna — lacking monkeys, however — has been dredged by fisherman from the floor of the Taiwan Strait, west-northwest of Tsochen (Lai, 1989, p. 45; Chen, 2000, p. 238); this fauna dates to the last glacial age. Late Pleistocene, ca. 0.12-0.01 Ma (cf. Liew et al., 1998, p. 92). The genera (one species each, except Cervus) of terrestrial mam- mals that are included in the Penghu fauna are Palaeoloxodon, Hyaena, Equus, Sus, Cervus (2 spp.), Elaphurus, Buhalus, and Capricornis. As expected from the age difference between the Tsochen and Penghu faunas, they have no mam- malian species in common (Lai, 1989, p. 45). FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 101 BURH^LL HALL 31 Holocene subfossil macaques have been re- ported from archeological sites dated to the late NeoHthic period (3.5-2.0 Ka) and the "Metal Age" (2.0-0.4 Ka) (Chen, 2000, p. 240). The original monkey fossils reported from Tsochen by Otsuka (1984, p. 37) apparently con- sisted of one or more teeth, but this material has been lost (Kin-Yang Lai, tm, retired, pers. comm., 22 Apr 1999). However, we have been able to examine eight fossil monkey teeth subsequently collected near Tsochen from the same beds that yielded the original monkey fossils (Table 25; Figs. 6, 7); seven of these fossils were collected between 1992 and 1998 by Liang-Jeh Wang, who generously provided access to them, and one was collected in 1992 by Ji-Nan Yeh. We have also examined two additional fossil monkey teeth col- lected in neighboring counties; one of these was collected by Mr Wang in 1992-1996 in Kao- hsiung Hsien, about 20 km south of Tsochen, and the other was collected by Wen-Chi Xue in 1998 in Chia-I Hsien, about 50 km north of Tsochen. In collaboration with Eric Delson (amnh), we are currently undertaking a detailed study of these ten fossil teeth. Our preliminary findings are: (I) all ten fossil teeth may be allocated to the genus Macaca, as previously determined for other Tsochen specimens by Otsuka (1984, pp. 37, 42, 44); (2) morphologically, these teeth are generally similar to those of extant M. cyclopis; (3) mea- surements of seven fossil teeth are within the range of variation of a sample of extant M. cyclo- pis teeth (Tables 25, 26); (4) measurements of three fossil teeth (specimen nos. 4, 6, and 12) are < 1 mm beyond the range of variation of the sam- ple of extant M. cyclopis teeth. Taxonomy Synonymy Macaca cyclopis Swinhoe, 1863 ("1862"), p. 350 [Sjinges: Du Halde, 1735, p. 162 — reported in west- ern Taiwan, based on visit in 1715 by Jesuit mis- sionary Pere J.-A.-M. de Moyria de Mailla (cf. Du Halde. 1735. p. xxxiv; Weiss, 1870-1873. p. 120; Campbell. 1903. pp. 504, 506; Davidson. 1903, p. 68). M\acaciis] speciosiis: Swinhoe in Blyth, 1860. p. 88 (not I. Geoffroy, 1826, p. 589) — "large Formosa Monkey . . . frequents the rocks on the coast of Formosa, especially in the neighborhood of Sakow or 'Ape's hill' " (p. 87); external characters. Macacus cyclopis Swinhoe, 1863 ("1862"). p. 350, pi. 42 — proposed provisionally; for details con- cerning type series, see below. Sclater, 1865 ("1864"), p. 710, fig. p. 711 — type history; exter- nal characters; probably specifically distinct from Macaca mulatta. Gray. 1 870, pp. 31.1 28 — type se- ries catalogued. Swinhoe. 1871 ("1870"). p. 615— type history. Sclater, 1871, p. 222 — unproved as specifically distinct from Macaca mulatta. Murie, 1873 ("1872"). p. 771 — type history; external and skeletal characters; specific status supported. Schle- gel, 1876, p. 1 14 — listed as an inadequately known geographic variant of Macacus enthraeus [= Ma- caca mulatta]. M[acaca] cyclopis: Pocock, 1926 ("1925"). p. 1547 — sexual skin. Macaca cyclopis: Kuroda, 1935, p. 291 — collection report. Kuroda. 1938, p. Ill — locality records. Kellogg, 1945. p. 119 — type-locality restriction. "Jusan (Apes' Hill), Takao prefecture. Formosa." Jones. 1975, p. 189 — type locality questionable, "?Apes Hill." Napier, 1981. p. 26 — type series cat- alogued. [[luius] cyclopis: Blyth. 1875. p. 5 — taxonomic rela- tionship. [Silenu.s] cyclopi.s: Stiles & Nolan. 1929, p. 528— parasites. Parasilenus cyclopis: Kishida. 1939 (work not seen); 1953, pp. 7, 24, 79— taxonomy. Pithecus cyclopsis: Elliot, 1913, pp. 181. 202 — incor- rect subsequent spelling, not an available name; type locality, "Island of Formosa"; lectotype des- ignated, "The type is a young male" (p. 203); ex- ternal characters; external and cranial measure- ments of lectotype. Macaca cyclopsis: Kishida, 1924. pp. 15 (index), 215 — allocated to subgenus Nemestrinus. Aoki, 1930, p. 505— habitat. Horikawa, 1932, p. 69— species account. Carter et al.. 1945, pp. 68, 203 — taxonomic relationship. Tate. 1947. p. 134 — exter- nal characters. M[acaca\ cyclopsis: Saheki et al., 1962, p. 101 — den- tal morphology. Macacus cyclopsis: Satoh et al., 1956, p. 41 — external measurements and proportions. Rhesus cyclopsis: Furuya. 1962, p. 377 — dermato- glyphics. Macacus Cyclapis: Horikawa, 1925, p. 237 — incor- rect subsequent spelling, not an available name; geographic distribution. Macacus cyclapis: Kano, 1929, p. 333 — geographic distribution. Kano. 1930, p. 165 — ethnozoology. Macaca coclopis: Kuroda, 1952, p. 288 — incorrect subsequent spelling, not an available name; listed. Macacus Sancti-Johannis: Gray, 1870, p. 129 (not Swinhoe. 1867 ("1866"], p. 556) — misidentifica- tion of BM(NH) 1868.4.14.4 (cf. Anderson, 1879, p. 87). Macacus (radiatus) affiiiis: Elliot. 1913. p. 202 (not Blyth. 1863. p. 8) — misidentification (see Remarks below). Type Series Macacus cyclopis Swinhoe, 1863 ("1862"), p. 350, is based on a captive male and a captive 32 FIELDIANA: ZOOLOGY Table 25. Taiwanese fossil monkey teeth: measurements and collection information Tentative identifica- Measurements'' (mm) Collection information Fossil Collec- no.^ tion APL A Br PBr tor^ Year County Locality** Custodian'' p3_p4 5 LP* 5.1 6.1 — Wang 1992-1996 M'-M^ Tainan Ihsindian THNHEC 1 R M^ (?) 7.5 7.2 6.2 Yeh 1992 Tainan Sanchunghsi TFM 2 R M2 (?) 8.7 7.7 7.3 Wang 1992-1996 Tainan Sankanshan TFM 4 R M^ 9.1 7.9 6.6 Wang 1992-1996 Tainan Ihsindian THNHEC 7 L M' (?) 6.5 6.0 6.0 Wang 1992-1996 Kaohsiung Hsiaokunshui L-J W 11 L MVM^ 7.3 7.7 6.8 Wang 1992-1996 M.-M^ Tainan Sankanshan TFM 6 R M,/M, 9.5 7.3 7.3 Xue 1998 Chia-I Shihtzulu W-C X 8 L M, (?) 8.4 6.8 6.5 Wang 1992-1996 M, 1998 Tainan Sanchunghsi L-J W 3 R M, 11.1 7.1 7.1 Wang Tainan Kuoling L-J W Deciduous premolar 12 P^/P4 6.8 4.2 4.0 Wang 1992-1996 Tainan Kuoling L-J W '^ Tooth-row position indicators. ' In the absence of formal catalog numbers, we have assigned informal interim numbers to these fossils. Fossil nos. 9 and 10 are problematic and are omitted here; these specimens will be discussed in a subsequent publication. *> Measurement abbreviations: A? L = anteroposterior length; A Br = anterior breadth; P Br = posterior breadth (inapplicable to permanent premolars). ^ Complete names of collectors: Liang-Jeh Wang; Wen-Chi Xue; Jinan Yeh. ** See map, Figure 6. ** Key to abbreviations: L-J W - Liang-Jeh Wang, Guanmiao, Tainan Hsien, private collection; TFM = Tsailiao Fossil Museum, Tsochen, Tainan Hsien; THNHEC = Tainan Hsien Natural History Education Center, Tsochen, Tainan Hsien; W-C X = Wen-Chi Xue, Chia-I Shih, private collection. female presented to the menagerie of the Zoolog- ical Society of London in 1862 by R. Swinhoe, then British vice-consul in Taiwan. The two syn- types died in captivity sometime between Decem- ber 1 864, when they were reported alive by Scla- ter (1865, p. 710), and April 1866, when the skins, skulls, and postcranial skeletons were accessioned by bm(nh) (male, no. 1866.4.25.8, original no. 1486A; female, no. 1866.4.25.9, original no. 1486B; cf. Swinhoe, 1871, p. 615). At the time of death, the male was a late juvenile — with ca- nines and third molars incompletely erupted, and the female was a subadult — with third molars in- completely erupted; when described by Swinhoe in 1862, both syntypes were at least 2 years youn- ger. The male was designated as lectotype by El- liot (1913, p. 203), and the female thereby became the paralectotype (International Code of Zoologi- cal Nomenclature, 1999, Article 74.1.3). A col- ored illustration of what appears to be the lecto- type male was published by Swinhoe (1863, pi. 42), and a black-and-white illustration of the par- alectotype female (with tumescent sexual skin) was published by Sclater (1865, p. 711). The pel- vis and adjacent lumbar and caudal vertebrae of the lectotype and paralectotype were figured by Murie (1873, pp. 773, 775). Type Locality In Swinhoe's (1863, p. 350) original description of Macacus cyclopis, the sole reference to the geographic origin of the syntypes is their desig- nation as "Formosan [^Taiwanese] Monkeys." The type locality was restricted by Kellogg (1945, p. 119) to "Jusan (Ape's Hill), Takao prefecture, Formosa" (=Shou Shan, Kaohsiung Hsien, Tai- wan). Swinhoe (1863, p. 350) apparently had no in- formation concerning the place of capture within Taiwan of the two syntypes of Macacus cyclopis. However, in his general discussion of the natural history of the species, Swinhoe mentioned that its habitats included "Apes' Hill" (=Shou Shan) in southwestern Taiwan (cf. Swinhoe in BIyth, 1860, FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 33 Sanchunghsi , jsochen Sankanshan \^ Ihsindian Kuoling Hsiaokunshui 100 Kilometers pf 25^ + 24- 20 Kilometers t / / Fig. 6. Map of southwestern Taiwan showing collecting localities of fossil monkey teeth. p. 87) and "mountains of the north and east"" of Taiwan. The first habitat mentioned by Swinhoe was selected by Kellogg (1945. p. 11 9) as the ba- sis for his type-locality restriction (with the spell- ing "Apes" Hill'" changed to "Ape"s Hill""). Remarks The name cyclopis (from "kyklos"" = circle, Greek) presumably was based on Swinhoe's (1863. p. 350) belief that what he perceived as the "round head"" of the immature syntypes was a primary character that distinguished this species from M. miilatta. Although Swinhoe proposed Macaciis cyclopis as a "provisional name"" (p. 350). such a conditional proposal does not impair the availability of scientific names proposed be- fore 1961 (International Code of Zoological No- menclature. 1999. Article 15.1). The name "Macaciis {radiatiis) affinis," attri- buted to Blyth (1863. p. 8). is routinely included in the synonymy of M. cyclopis (Elliot. 1913. p. 202: Kellogg. 1945. p. 119; Ellerman & Morri- son-Scott. 1951. p. 198: Imaizumi. 1960, p. 105; 1970. p. 320: Hill. 1974. p. 583; Napier, 1981, p. 26; Corbet. 1992. p. 171; Groves. 1993, p. 266); however, this is an error. At the root of this error is a second pair (juvenile male and female) of captive macaques — not the type series of M. cy- clopis (see above) — that was acquired ca. 1859 by R. Swinhoe (in Blyth. 1860. p. 87). In 1859, the skull of the male and the living captive female were sent by Swinhoe to the Asiatic Society of Bengal, Calcutta (Blyth, 1860, p. 87; 1863, p. 8; 1875, p. 8; Anderson. 1881. p. 60). Although the present whereabouts of the male skull is un- known, the female skin and skull are preserved in the collection of the Zoological Survey of India (National Zoological Collection. Calcutta, no. 11826: cf. Khajuria. 1955. p. 112): this skin and skull are unambiguously identifiable as those of an Indian bonnet macaque. M. radiata (E. Geof- froy. 1812. p. 98), the range of which is approx- imately 4,000 km west of Taiwan. However, this identification was not made by Swinhoe (in Blyth, 1860, p. 87; cf. Swinhoe, 1863, p. 350). who in- stead mistakenly asserted that his pair of M. ra- 34 FIELDIANA: ZOOLOGY Fig. 7. Fossil monkey teeth eollected in soutiiwestern Taiwan. Fossil No. 2 (Sankanshan), right upper second molar (?), length 8.7 mm: A, buccal view; B, occlusal view. Fossil No. 3 (Kuoling), right lower third molar, length 11.1 mm: C, lingual view; D, occlusobuccal view. diata captives were examples of "the small For- mosa Monkey" (italics in original; note: for Swin- hoe, the large Formosa Monkey was M. cyclopis), which allegedly inhabited "the camphor forests of the Formosa mountains"; the source of this geo- graphic misinformation presumably was the un- known person who supplied the M. radiata cap- tives to Swinhoe. The spurious locality informa- tion misled Blyth (1863, p. 8; cf. 1860, p. 88, footnote), zoological curator of the Asiatic Soci- ety of Bengal, who incorrectly concluded that a slightly variant form of M. radiata inhabited Tai- wan; Blyth cited this form as "M[acacus] radia- TUS (affinis),'' indicating that he regarded the un- named form as related to ("affinis," Latin) M. radiata, not that he was proposing affinis as a new subspecific or specific name. Anderson (1879, p. 91, footnote), without explanation or justification, changed the italicization and shifted the parenthe- sis in Blyth's citation to produce ''Macacus (ra- diatus) affinis,'' which is the orthography em- ployed for this citation in current checklists. Elliot (1913, p. 202) adopted Anderson's orthography and, based on the spurious locality "Formosa," he included ''Macacus (radiatus) affinis'' in the synonymy of his Pithecus cyclopsis (=M. cyclo- pis), thereby establishing the pattern that has been followed by subsequent authors. However, M[acacus] radiatus {affinis): Blyth, 1863, clear- ly is a synonym of Macaca radiata (E. Geoffroy, 1812) (cf. Fooden, 1981, p. 34), not of Macaca cyclopis Swinhoe, 1863 ("1862"). FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 35 Table 26. Dental measurements (mm) in wild-collected specimens'^ of Macaca cyclopis. Females Males Dental . dimen- Mini- Maxi- Mini- Maxi- Tooth sion'' N Mean SD mum mum N Mean SD mum mum Permanent teeth P3 APL 5 5.0 0.23 4.8 5.4 10 5.2 0.35 4.8 5.9 Br 5 5.7 0.30 5.4 6.2 10 6.0 0.34 5.5 6.6 F APL 5 4.9 0.21 4.8 5.3 5.1 0.31 4.7 5.7 Br 5 6.3 0.41 5.9 6.8 6.3 0.26 5.9 6.6 M' APL 6 7.1 0.26 6.8 7.4 7.2 0.42 6.5 7.9 A Br 6 6.8 0.52 6.1 7.5 7.0 0.46 6.3 7.7 PBr 6 6.3 0.55 5.7 7.1 6.6 0.55 5.8 7.7 M2 APL 5 7.8 0.20 7.6 8.1 8.1 0.35 7.6 8.7 A Br 5 7.6 0.53 7.0 8.2 8.2 0.42 7.5 8.9 PBr 5 7.1 0.49 6.5 7.5 7.6 0.40 6.8 8.0 M3 APL 5 7.6 0.39 7.1 8.0 8.0 0.54 7.3 8.8 A Br 5 7.4 0.26 7.0 7.6 8.0 0.39 7.4 8.4 PBr 5 6.6 0.45 6.2 7.3 6.8 0.40 5.8 7.2 M, APL 4 6.8 0.35 6.4 7.1 10 6.9 0.48 6.1 7.6 A Br 4 5.5 0.48 5.1 6.1 10 5.7 0.37 5.0 6.3 PBr 4 5.3 0.27 5.0 5.6 10 5.6 0.32 5.1 5.9 M2 APL 4 7.8 0.30 7.4 8.0 11 8.1 0.35 7.5 8.7 A Br 4 6.6 0.53 6.0 7.0 11 6.9 0.39 5.9 7.3 PBr 4 6.1 0.48 5.6 6.7 11 6.4 0.33 5.9 7.0 M3 APL 4 10.0 0.62 9.5 10.9 10 10.9 0.60 9.9 11.9 A Br 4 6.9 0.18 6.7 7.1 10 7.2 0.53 6.2 8.2 PBr 4 5.9 0.36 5.4 6.2 9 6.2 0.34 5.4 6.6 Deciduous teeth P^ APL A Br PBr 5.7 4.9 4.6 V V V 5.5 4.8 4.9 P* APL A Br P Br 5.9 5.8 5.1 V V 6.2 5.6 5.4 Pj APL A Br PBr 5.8 3.9 3.8 P4 APL A Br P Br 6.0 4.8 5.0 '"^ Tooth-row position indicators. "^ AMNH. USNM. private collections. ^ Abbreviations: A Br = anterior breadth; AP L = anteroposterior length: Br ^ Sex of specimen uncertain; mandible missing. breadth; P Br = posterior breadth. Evolution and Dispersal In this section, we will attempt to address four main questions concerning the evolutionary his- tory of Taiwanese macaques: 1 . Where did the progenitors of M. cyclopis come from? 2. When did these progenitors disperse to Tai- wan? 3. Why is M. cyclopis absent from nearby islands surrounding Taiwan? 4. Why is M. miilatta absent from Taiwan? Taiwan is situated on the Asian continental shelf, separated from the mainland by the Taiwan Haixia ( = Formosa Strait), which is ca. 140 km wide and less than 120 m deep (U.S. Defense Mapping Agency, 1981); rising from the floor of the strait is a sinuous submarine ridge that con- nects Taiwan with mainland Asia at a depth of less than 60 m below present sea level. Therefore, during Pleistocene glacial advances and correlated sea-level depressions, a dry-land connection in- termittently existed between Taiwan and the mainland (Fooden, 1995, p. 87). The evidence of pollen studies indicates that. 36 FIELDIANA: ZOOLOGY during Pleistocene glacial advances, the climate of Taiwan was colder and drier than at present (Liew, 1991, p. 30). During the Last Glacial Max- imum, ca. 1 8 Ka, upland forests were more open, lowland grasslands were much more extensive, and annual precipitation was at least 800 mm less than at present (Liew et al., 1998, p. 93); at this time, vegetation that is now restricted to 1500- 2500 m elevation occurred as low as 650 m (Huang et al., 1997, p. 67). The terrestrial mammal fauna of Taiwan is closely related to that of the Asian mainland to the west, not to that of the Philippine Islands to the south or the Ryukyu Islands to the east (Swin- hoe, 1863, p. 350; Wallace, 1876, p. 334; 1881, p. 367; Kuroda, 1939, p. 45; 1952, p. 285; Kano, 1940, p. 107; Corbet & Hill, 1992, p. 4). Zoogeo- graphic analysis indicates that Taiwan has accu- mulated its vertebrate and invertebrate fauna as a result of multiple waves of dispersal from main- land Asia to the island (Kano, 1940, p. 122; Ja- meson & Jones, 1977, p. 479; Yu, 1995, p. 80; Hsu et al., 1997, p. 154; Wang & Tu, 1997, p. 103); some or all of these dispersal waves pre- sumably were correlated with successive glacial advances. Subsequent to dispersal, the immediate ancestors of many extant Taiwanese species ap- parently have become extinct on the Asian main- land opposite Taiwan, as originally noted by Swinhoe (1863, p. 349) and Wallace (1881, p. 404). Reproductive anatomy, DNA evidence, and blood-protein evidence indicate that M. cyclopis is closely related to M. mulatta (see Molecular Biology and Genetics, above; Fooden, 1972, p. 310; Fooden & Lanyon, 1989, p. 227), which sug- gests two alternative evolutionary hypotheses: (1) long-tailed M. cyclopis is descended from a short- tailed M. mulatta-Vike ancestor (which itself was previously descended from a long-tailed ances- tor), or (2) short-tailed M. mulatta is descended from a long-tailed M. cyclopis-like ancestor. Com- parative study of tail-length variation in fascicu- taris-group species tends to support the second hypothesis (Fooden & Albrecht, 1999, p. 437; Fooden, 2000, p. 87). According to this interpre- tation, M. cyclopis is an insular relict of a rela- tively long-tailed population of macaques that for- merly inhabited mainland Asia between 15°N and 25°N; this extinct mainland long-tailed popula- tion, which northward was ancestral to M. mulat- ta, has now been replaced between 15°N and 25°N by short-tailed M. mulatta. Such an evolutionary history would parallel that of many other Taiwa- nese species (see above). Macaques were present on Taiwan before or during the Middle Pleistocene, at least 300 Ka (see Fossils, above). The fossilized teeth of these macaques, the only parts that have been pre- served, are similar in size and morphology to the teeth of living M. cyclopis. The simplest hypoth- esis, as previously implied by Shen (1994, p. 6), is that the Pleistocene Taiwanese macaques rep- resented by the fossil teeth were directly ancestral to extant M. cyclopis. Based on the age of the Taiwanese macaque fossils, the dispersal of the postulated relatively long-tailed population from mainland Asia to Tai- wan apparently occurred before 300 Ka — well be- fore the onset of the most recent glacial advance (ca. 1 15 Ka). The physiological ability of M. cy- clopis to survive subsequent cold glacial intervals in Taiwan is demonstrated by its present occur- rence at relatively high elevations in Taiwan (Ta- ble 16) and by the survival of introduced popu- lations in temperate Japan, 10°- 15° north of Tai- wan (see Distribution, above). Macaca cyclopis is absent from nearby islands that surround Taiwan, including the relatively large islands Lan Yu, Lu Tao, and Penghu Tao (Fig. 1). Lan Yu and Lu Tao lie beyond the con- tinental shelf and are separated from Taiwan by deep channels (>1500 m and >500 m, respec- tively), which would account for the apparent in- ability of M. cyclopis to disperse eastward from Taiwan to these two islands (U.S. Defense Map- ping Agency, 1981). A similar explanation would apply, with even greater force, to the absence of M. cyclopis from the Ryukyu Islands, 1 10 km east of Taiwan and the continental shelf. Penghu Tao lies on the continental shelf with Taiwan, within the 120-m bathymetric contour line, less than 50 km west of Taiwan. However, Penghu Tao currently is relatively dry (977 mm annual precipitation) and minimally forested (Hsieh, 1964, pp. 103, 282; Hsieh & Shen, 1994, p. 3), and it may have been totally unforested dur- ing the Last Glacial Maximum, ca. 18 Ka (see below). The absence of M. cyclopis from Penghu Tao probably is explained by the absence there of suitable habitat, at least since the Last Glacial Maximum. Although the rhesus macaque, M. mulatta, ranges widely in mainland Asia and also inhabits some continental islands, it is absent from Taiwan (Fooden, 2000, p. 2). Macaca mulatta probably dispersed from mainland Asia to Hainan Dao, an FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 37 island on the continental shelf ca. 1000 km south- west of Taiwan, during the Last Glacial Maximum (Fooden, 2000, p. 89). Unsuitable habitat condi- tions on the contemporaneous land bridge to Tai- wan may account for the apparent failure of M. midatta to disperse to Taiwan during the Last Gla- cial Maximum. Palynological evidence indicates that most of the exposed Late Pleistocene conti- nental shelf near Taiwan was grassland or barren (Liew et al., 1998, p. 93). The composition of the Taiwan Strait fossil fauna (see Fossils, above) is compatible with this interpretation. To summarize, tentative answers to questions concerning the evolutionary history of M. cyclo- pis that were posed in the first paragraph of this section are as follows: 1. The progenitors of M. cyclopis probably dis- persed to Taiwan from nearby mainland Asia. The now-extinct mainland Asian ancestors probably were long-tailed, like M. cyclopis it- self. 2. The dispersal of M. cyclopis from mainland Asia to Taiwan apparently occurred before 300 Ka. 3. Persistent water barriers probably have pre- vented M. cyclopis from dispersing from Tai- wan to many of the surrounding islands. Al- though Penghu Tao was joined to Taiwan by dry land during Pleistocene sea-level regres- sions, forest habitats suitable for the survival of M. cyclopis apparently have been absent from Penghu Tao, at least since the Last Gla- cial Maximum. 4. Macaco miilatta probably was unable to dis- perse to Taiwan during the Last Glacial Max- imum because the land bridge available at that time was unforested. Acknowledgments For access to specimens and generous cooper- ation, we are deeply grateful to officials and staff members of institutions cited above (see Intro- duction). For supplementary advice, information, and assistance, we also thank M. Aimi (kupri), W. Birky (Rutgers University), D. Brandon-Jones (bm(nh)), Shih-Wei Chang (tesri), Shu-Mei Chen (ntuz), Jui-Hua Chu (ntuz), E. Delson (amnh), M. Iwamoto (jmc), Y. Kawamoto (kuprO, Kin- Yang Lai (tm), Ling-Ling Lee (ntuz), Si-Ming Lin (ntuz), p. Lowther (Field Museum), M. Pan- nell (Field Museum), C. Simpson (Field Muse- um), R. W. Thorington, Jr. (usnm), Liang-Jeh Wang (Guanmiao, Tainan Hsien), J. Weinstein (Field Museum), Wen-Chi Xue (Chia-I Hsien), and Jon-Ne Yo (ntuz). K. H. Hamilton (Chicago) kindly assisted with translation of Japanese pri- matological literature. This research was partly supported by the Barbara E. Brown Fund for Mammal Research. 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Journal of Human Evolution, 41: 45-59 Hamada, Y, T Watanabe, and M. Iwamoto. 1992. Variation of body color within macaques, especially in the Japanese macaques. Primate Research, 8: 1-23. (In Japanese, English summary) Su, Hsiu-Hui, AND Ling-Ling Lee. 2001. Food habits of Formosan rock macaques {Macaca cyclopis) in Jentse, Northeastern Taiwan, assessed by fecal analysis and behavioral observation. International Journal of Pri- matology, 22: 359-377. FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 45 Appendix 1: Specimens Examined (Total 237) Skins and Skulls, 36 Hengchun — yio 733, 734; Hualien, mountains near — amnh 184955; Ilan region — zmb 43048; Kaohsiung vicinity — usnm 296794; 296795; Maanliao — usnm 294177; Manchoutsun — smtd B4845/26383, zmb 26472-26477, 26479; Mu- Kua-Chau— AMNH 184956, 184957; Pinam— zmb A54.04; Puli — ummz 85348. No locality — bm(nh) 1866.4.25.8 (lectotype), 1866.4.25.9 (paralecto- type), 1868.4.14.4, mcz 37679, ummz 85349, usnm 296796, 308427-308429, 308722, 396927, 397401, 397444, 397645-397647, 397707. Skins Only, 28 Kuohsing Subcounty — tesri Accession No. T 01 15 (cadaver); Nanhua Subcounty — tesri Ac- cession No. T 0177 (cadaver); Puli vicinity — TM MA-0272; Ta Ching Shui— nmns T 1726 (cadav- er); Taroko National Park— nmns T 1883, T 391 1. No locality— NMNS T 1725, T 1731, T 3909 (3 cadavers), ntuz uncatalogued specimen, tesri MC 0078, MC 0091, TM MA-OOlO-MA-0012, MA- 0092, MA-0094-MA-0097, MA-0236, MA-0265, MA-0378, usnm 296797, 308717, 308721, zmb 42135, 42546. Preserve — private collection, unnumbered skull; Eastern Taiwan — ntuz 063, 064, uncatalogued specimen; Hsiaoi — usnm 3991 1 1; Hui-Sun Exper- imental Forest — private collection, Mk-7; Jentse — ntuz uncatalogued specimen; Renting — private collection, Mk-1, Mk-2, Mk-6; Manchou- tsun—zmb 26366, 26478, 26553-26557, 26568- 26573; Oshima (introduced) — kupri 4486; Shao Hsing Shui — nmns 001396; Shenmu Tsun — tesri MC 0012, MC 0017; Shoulin Shan— tesri MC 0084-MC 0086 (mandibles only), Shou Shan— usnm 399110; Taimali Hsi— tesri MC 0014 (mandible only), MC 0114; Wushihkeng — tesri MC 0016; Wutao Shan— tesri MC 0081; Yen Hai logging trail — tesri MC 0043. No locality — ansp 11884, JMC 415, 910, 946, 1024, 1203, 1232, 1469, 1551, 1593, 2069, 2070, 2112, 2128, 2181, 2245, 2288, 2500, 3104, 3153, 3208, 3224, 3232, 3329, 3368, 3371, 3374, 3381, 3384, 3390, 3401, 3465, 3625, 3634, 3764, 4169, 4514, 4532, 4553, 4641, 4665, 4891, 5040, 5047, 5142, 5150, kupri 302, 303. 305, 526, 528, 559, 560, 586-588, 1315, 1358, 1511, 1682, 1745, 1832, 1883,2289, 2290, 2452, 2514, 2795, 2796, 2846, 2913, 3056, 3224, 3225, 3651-3660, 3852, 4201, 4217, 4233, 4238, 4508, 5018, 5228, 5843, 5847, 6082, 6431, nmns 000088, 000102, 000103, 000148, 000546- 000548, 001264, ntuz Aoki 3484, private collec- tion (Mk-3, Mk-4, Mk-5), usnm 308718-308720, 332888-333895, 358612-358618, 396928, 397975, zmb 23245. Skulls Only, 169 A-Sun-Lai-Ga — private collection, two unnum- bered skulls; Ayu Shan — usnm 399112; Ching- shui Cliff — private collection, unnumbered skull; Chuchi — ntuz Aoki 94; Tawa Mountain Nature Postcranials Only, 4 No locality— kupri 304, 2288, 4721, nmns 001996. 46 FIELDIANA: ZOOLOGY Appendix 2: Gazetteer of Macaca cyclopis Localities The romanization of Taiwanese locality names that are listed as primary entries in this gazetteer preferentially follows that in "List of Locality Names in Taiwan Province" (Zhuang, 1992). In addition, the following supplementary geo- graphic references have been consulted: Na- tional Atlas of China, Volume 1 — Taiwan (1967); Chu and Yamanaka (1973); U.S. Board on Geographic Names, Republic of China — Official Standard Names Gazetteer (1974); Taiwan Atlas— Republic of China (1981); Kyoda (1989); Chen (1993); Huang et al. (1993). Localities of M cyclopis that were not found in "List of Locality Names in Taiwan Province" or supplementary references are spelled here as in the original sources. Secondary entries, with cross references to corresponding primary en- tries, indicate variant spellings or alternate lo- cality names that appear on specimen tags, in published literature, or in unpublished notes concerning M cyclopis. The sequence of information presented in pri- mary entries is as follows: 1 . Locality name, in roman letters and Chinese characters. 2. Elevation, if reported by collector or ob- server. 3. Name of county (="hsien"), in capital let- ters. 4. Coordinates of locality; the principal sources of coordinates are references indi- cated above and published or unpublished field notes of collectors or observers. 5. Date of collection or observation. 6. Name of collector or observer. 7. Bibliographic reference (in parentheses) to published or unpublished field notes, if any. 8. Abbreviated name of museum (see Intro- duction) where specimens are preserved. 9. Number of specimens available (with indi- cation of part preserved, if skin and skull are not both present). 10. Locality number (italicized) as indicated in distribution map (Fig. 1). Alishan zone (HSUjM), Alishan Hsiang, 2260 m; CHlA-1; 23°32'N, 120°48'E; droppings observed 30 Jul. 1997 by TESRI survey team. N143 Alishan zone (HMUj^), Alishan Hsiang, 2310 m; CHlA-1; 23°3rN, 120°48'E; droppings observed 1 Aug. 1997 by tesri survey team. N143 Ali village. See Wutao Shan. Amma Shan. See Anmashan. Anmashan { ^ % \U ) (=Amma Shan), 1500-2900 m; TAICHUNG; 24''16'N, 12r00'E; observed Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145). Reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386; Tanaka, 1986, p. 101). A^79 Anpingkenghsi Mining Trail {'^'^inMMWjB. 1^), Suao-Nanao Hsiang, 175 m; ILAN; 24°37'N, 121°46'E; 3 monkeys observed 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N18 Anpingkenghsi Mining Trail {"^^^ijVi^WvBM l^), Suao-Nanao Hsiang, 685 m; ILAN; 24°35'N, 121°44'E; observed 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N24 Anpingkenghsi Mining Trail {"t^^inMMWM {^), Suao-Nanao Hsiang, 690 m; ILAN; 24°33'N, 121°43'E; vocalization heard 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N23 Anpingkenghsi Mining Trail (S^^ijiW:W^^ l^), Suao-Nanao Hsiang, 810-930 m; ILAN; 24°35'N, 121°44'E; observed at 810 m and 930 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N24 Anpingkenghsi Mining Trail Q^^Wi^W^M jg#), Suao-Nanao Hsiang, 1000 m; ILAN; 24°33'N, 12r43'E; vocalization heard 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N23 Antungwenchuan ( ^ jS ^m :^ ), Yuli Chen; HUALIEN; 23°17'N, 121°20'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386).5i0 Apenberg. See Shou Shan. Ape's Hill. See Shou Shan. A-Sun-Lai-Ga ( R # ^ DM ), Lakulaku Hsi, 800-1000 m; HUALIEN; 23°22'N, 12ril'E; collected in 1999 by H.-C. Chu; private col- lection, 2 (skulls only, including 1 lacking mandible). S22 FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS Al Ayu Shan (R5[i4); ILAN; 24°47'N, 121°36'E; collected 12 Apr. 1963. probably by U.S. Naval Medical Research Unit No. 2; usnm, 1 (skull only). NIO Central Range. See Chung Yang Shan Mo. Chai Shan. See Shou Shan. Changhu {WM\ Kukeng Hsiang, 310-350 m; YUNLIN; 23''36'N, 120°37'E; 3 monkeys ob- served at 340 m 9 Dec. 1994 by TESRI survey team. Observed at 310 m and 340 m 13 Dec. 1994 by TESRi survey team. Observed at 320 m and 350 m 16 Dec. 1994 by TESRI survey team. Droppings observed at 350 m 3 Mar. 1995 by TESRl survey team. Droppings ob- served at 340 m 12 Apr. 1995 by tesri survey team. Nl 16 Changliang Logging Trail (:M^#xS), Chohsi Hsiang, 825 m; HUALIEN; 23°15'N. 12ri2'E; observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. S35 Changliang Logging Trail (:^^#3M), Chohsi Hsiang, 980 m; HUALIEN; 23°15'N, 12ri3'E; observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. S34 Changliang Logging Trail (:M:K^xS). Chohsi Hsiang, 1920 m; HUALIEN; 23°12'N, 12r08'E; 2 monkeys observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. S72 Changliang Logging Trail (:^^#jM), Chohsi Hsiang, 1965 m; HUALIEN; 23°12'N, 121°09'E; observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. S73 Chaochou ( ^ jtl ) (=Choshu); PINGTUNG; 22°33'N, 120°32'E; observed before 1930 by T. Kano(1929, p. 333).57iO Chia-I (MS); CHIA-I; 23°29'N, 120°27'E; col- lected for parasitological research before 1964 by R. E. Kuntz (Schad and Anderson, 1963, p. 799). Serum samples obtained 1964-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967. p. 146). SI CHIA-I (MS); 23°13-38'N, 120°07-57'E; col- lected 12 Jan. 1937 probably by B. Aoki; NTUZ, 1 (skull only; external measurements on skull tag). Not mapped. Chichi Tashan ( ^ ^ ;*v [ll ), Chichi Chen; NANTOU; 23''51'N, 120°50'E; reported in 1980 by S. Tanaka (1986. p. 98), subsequent extinction at localit\ suspected. Observed in 1996 by S.-W. Chang (1999. p. 13). N105 Chichi Tashan ( ^ ^ y>v Uj ), Chichi Chen, 700-1250 m; NANTOU; 23°5rN, 120°49'E; reported at 800 m 15 Nov. 1992 to tesri sur- vey team. Capture reported at 700 m 10 Dec. 1992 to TESRI survey team. Droppings ob- served at 1250 m 8 Mar. 1993 by tesri survey team. N105 Chienho (^^), Peinan Hsiang; TAITUNG; 22''44'N, 121°03'E; reported in 1982 by K. Masui, Y. Narita. and S. Tanaka (1986, p. 386). 5/79 Chientai Logging Trail ( ^ ■□ # IM ), Jenai Hsiang, 12^50-1300 m; NANTOU; 23°56'N, 121°07'E; droppings observed at 1250 m and 12 monkeys observed at 1300 m 29 Nov. 1996 by S.-W. Chang (tesri). N68 Chihpen Hsi (^p^^^), left bank, Peinan Hsiang; TAITUNG; ca. 22°40'N, 120°57'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 5/22 Chihpen Hsi (^0^?^), right bank, Peinan Hsiang; TAITUNG; ca. 22°4rN, 120°59'E; observed 7-18 Aug. 1982 by K. Masui, Y. Narita. and S. Tanaka (1986, pp. 386. 391). S121 Chihpen Logging Trail (^^^IM), Peinan Hsiang. 280^80 m; TAITUNG; 22°38'N, 12r00'E; observed at 200 m and 480 m 31 May 1996 by S.-W. Chang (tesri). S124 Chihpen Wenchuan (^D^^m:^), Peinan Hsiang; TAITUNG; 22°42'N, \2\°0\'E; observed 1967-1971 and 1982 by K. Masui, Y. Narita, and S. Tanaka (1986. p. 386). SI 20 Chingjen Ku (fWAS). Fengyuan; 480 m; TAICHUNG; 24°14'N, 120°46'E; 3 monkeys observed 13 Oct. 1995 by tesri survey team. N84 Chingmei (if ^), Hsiulin Hsiang; HUALIEN; 24''05'N, 121°36'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). N56 Chingshui Cliff (ifTKifr^). Hsiulin Hsiang, 100 m; HUALIEN; 24°12'N, 121^40^; col- lected 27 Jan. 2000 by J.-H. Chu; private col- lection (skull only). N54 Chingshui Hsi Power Station (^"raTK^^fl;)^), Chohsi Hsiang; HUALIEN; 23°17'N, 12ri6'E; reported in 1982 by K. Masui, Y. 48 HELDIANA: ZOOLOGY Narita, and S. Tanaka (1986, p. 386). S32 Ching Sway Go. See Chinshuikou. Chinlun (^^), Taimali Hsiang; TAITUNG; 22°32'N, 120°57'E; reported 1967-1971 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S126 Chinlunshan (^^[JLj), Tahsi, Chinfeng Hsiang; TAITUNG; 22°34'N, 120''56'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). 5725 Chinshuikou (vf tRS) (=Ching Sway Go), 1000 m; NANTOU; ca. 23°49'N, 120°45'E; three captives reported Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145; Masui et al., 1986, p. 386). N108 Chishan vicinity (:K|1(%1^), Chishan-Meinon, 200 m; KAOHSIUNG; 22°53'N, 120°29'E; observed 15 Feb. 2000 by H.-Y. Wu. SI 05 Chishan vicinity (iKUjil^lm), Chishan-Meinon, 200 m; KAOHSIUNG; 22°54'N, 120°30'E; 3 groups observed 15 Feb. 2000 by H.-Y. Wu. SI 04 Chi Tou. See Hsitou. Chitsaihu (-^7^^/^), Hsini Hsiang, 2600-2730 m; NANTOU; 23°45'N, 121°13'E; droppings observed at 2730 m 17 Nov. 1994 by TESRI survey team. Droppings observed at 2600 m 18 Nov. 1994 by TESRi survey team. N131 Chiuchiu Feng {j\f\M\ Tsaotun Chen, 200 m; NANTOU; 24°00'N, 120°45'E; droppings observed 3 Feb. 1993 by tesri survey team. N95 Chiumei Bridge (A^i^^), Hsiulin Hsiang, 800 m; HUALIEN; 24°13'N, 121°28'E; ob- served 2 Aug. 1997 by H.-Y. Wu. N49 Chiupeng ( A ffl ), Manchou Hsiang; PINGTUNG; 22°07'N, 120°52'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). 5/50 Choching (J^/f ), Chohsi Hsiang; HUALIEN; 23°16'N, 121°16'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S32 Choshu. See Chaochou. Chuchi (ttllff); CHIA-I; 23°31'N, 120*'33'E; collected 3 Jan. 1930, probably by B. Aoki; NTUZ, 1 (skeleton only, external measure- ments on specimen tag). N148 Chufengshan ( tB JH lJL| ), Manchou Hsiang; PINGTUNG; 22"03'N, 120°52'E; reported Jul.-Nov. 1985 by S. Tanaka (1986, p. 98). S151 Chulaoshu Shan {M \^^\i\) (=Teraso); PINGTUNG; 22°01'N, 120°51'E; collected ca. 1897 by I. Namiye (1914, p. 213; Huang et al., 1993, p. 167); photo of mounted specimen published, museum unknown. 5752 Chunfu (?), Changpin (=Chanpin) Hsiang; TAITUNG; ca. 23''20'N, 121°25'E; observed in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 529 Chungan (M^), Iwan (^^), Chengkung Chen; TAITUNG; ca. 23°12'N, 121°23'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 57^ Chungan (M^), Poaili (t#^M), Chengkung Chen; TAITUNG; ca. 23°I2'N, 12 1^23 'E; re- ported in 1982 by K. Masui, Y. Narita, and S. Tanaka(1986, p. 386). 57^ Chunghsing Tahsueh (=University), experimen- tal forest (cfnffly^^^,^#i#), Jenai Hsiang; NANTOU; ca. 24°04'N, Hl^OO'E; reported Jul.-Nov. 1985 by S. Tanaka (1986, p. 98). N73 Chungliao Hsiang {^'^M\ 1000 m; NANTOU; ca. 23°52'N, 120°45'E; reported 14 Jan. 1993 to tesri survey team. A'^707 Chung Yang Shan Mo (4^^UjM) (=Central Range), 3400 m; 22°15'-24°35'N, 120°45'-121°35'E; monkey droppings ob- served Jun.-Jul. 1973 by D. R. McCullough (1974, p. 11). Not mapped. Chunta Logging Trail (^A#?S), Hsini Hsiang; NANTOU; 23°36^0'N, 120°52-56'E; re- ported 1 Jan. 1994 to tesri survey team. A^727 Chunta Logging Trail (^:;^#jiS), Hsini Hsiang; NANTOU; 23''44'N, 120°56'E; reported 29 May 1997 to S.-W. Chang (tesri). A^725 Chunta Logging Trail (^[5;;*C#xS), Hsini Hsiang, 810 m; NANTOU; 23''41'N, 120''53'E; drop- pings observed 25 Jun. 1997 by S.-W. Chang (tesri). A^7 22 Chunta Logging Trail (^P:;^^?!), Hsini Hsiang, 1355-2515 m; NANTOU; 23°33-40'N, I20°54'E; 5 droppings observed at 1355 m and 1360 m 1 Sep. 1993 by TESRI survey team. Two monkeys observed at 2050 m and drop- FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 49 pings observed at 2280 m 3 Sep. 1993 by TESRI survey team. Vocalization heard at 1380 m and droppings observed at 1470 m 3 Feb. 1994 by TESRI survey team. Droppings ob- served at 1750 m and 2040 m 10 Mar. 1994 by TESRJ survey team. Droppings observed at 1725 m 11 Mar. 1994 by TESRi survey team. Droppings observed at 2515 m 8 Jan. 1995 by TESRI survey team. N121 Chunta Logging Trail (^A#xl), Hsini Hsiang, 1425 m; NANTOU; 23°41'N, 120°54'E; 4 monkeys observed 24 June 1997 by S.-W. Chang (TESRI). N130 Chunta Logging Trail (^::A;#3g), Hsini Hsiang, 1580 m; NANTOU; 23°42'N, 120°54'E; vo- calization heard 24 June 1997 by S.-W. Chang (TESRI). N123 Chunta Logging Trail (?[5:;'v#xS), Hsini Hsiang, 1910-1920 m; NANTOU; 23°42'N, 120°55'E; 3 monkeys observed at 1910 m 9 Mar. 1994 by TESRI survey team. Three monkeys ob- served at 1920 m by S.-W. Chang (tesri). N129 Chunta Logging Trail (^:;^#jM), Hsini Hsiang, 2450-2455 m; NANTOU; 23°45'N, 120°58'E; droppings observed at 2450 m and 2455 m 28 May 1997 by S.-W. Chang (TESRi). N127 Chunta Logging Trail (?|j;*v#iS), Hsini Hsiang, 2470 m; NANTOU; 23°45'N, 120°59'E; 3 monkeys observed 28 May 1997 by S.-W. Chang (TESRI). A^/2 7 Chunta Logging Trail (^f^;*v#xS), Hsini Hsiang, 2485 m; NANTOU; 23"44'N, 120^58^; droppings observed 28 May 1997 by S.-W. Chang (TESRI). N128 Chunta Logging Trail (^:;'v#3fi), Hsini Hsiang, 2495 m; NANTOU; 23°44'N, 120°57'E; drop- pings observed 28 May 1997 by S.-W. Chang (TESRI). N126 Chunta Logging Trail (^A^M), Hsini Hsiang, 2705 m; NANTOU; 23°43'N, 120°56'E; food remains and droppings observed 29 May 1997 by S.-W. Chang (tesri). N125 Chunta Logging Trail (^y*C#ii), Hsini Hsiang, 2790-2855 m; NANTOU; 23°44'N, 120°56'E; food remains and droppings observed at 2790 m and droppings observed at 2840 m and 2855 m 29 May 1997 by S.-W. Chang (tesri). N125 Chushe Hsi ( tl" li ^ ), Mutan Hsiang; PINGTUNG; ca. 22°08'N, 120°48'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka(1986, p. 386). 5/^5 Chushuihsi (^tR]^), Hsini Hsiang, 1300 m; NANTOU; 23°31'N, 120°51'E; 3 monkeys observed 20 June 1995 by tesri survey team. N142 Chutsun (t-tt^), Hsiulin Hsiang; HUALIEN; 24°16'N, 121°30'E; collected in 1989 by C.-T. Yao; tesri, 3 (mandibles only). N47 Chutsun (t^t^), Hsiulin Hsiang, 1100 m; HUALIEN; 24°15'N, 12r29'E; observed 8 Aug. 1997 by H.-Y. Wu. N48 Chuyunshan Logging Trail (tBSLl4#xi), 930-960 m; KAOHSIUNG; 22°57'N, 120°45'E; group observed at 930 m and drop- pings observed at 960 m 21 Oct. 1998 by S.-W. Chang (tesri). S98 Chuyunshan Logging Trail (tBff|ll#^), 1135-1300 m; KAOHSIUNG; 22°58'N, 120°47'E; vocalization heard at 1135 m and droppings observed at 1300 m 21 Oct. 1998 by S.-W. Chang (TESRi). S96 Chuyunshan Logging Trail (ttiS[JL|#jI)< 1290 m; KAOHSIUNG; 22°57'N, 120°44'E; 5 monkeys observed 22 Oct. 1998 by S.-W. Chang (TESRI). S99 Chuyunshan Logging Trail (tb®Uj#7l), 1350 m; KAOHSIUNG; 22''56'N, 120°45'E; 5 monkeys observed 22 Oct. 1998 by S.-W. Chang (TESRI). S97 Chuyunshan Logging Trail (thSUjt-l^jI), 1360 m; KAOHSIUNG; 22°58'N, 120°48'E; dead monkey observed 21 Oct. 1998 by S.-W. Chang (TESRI). S95 Chuyunshan Logging Trail (tBff Uj^M), 1400 m; KAOHSIUNG; 22''59'N, 120°48'E; drop- pings observed 21 Oct. 1998 by S.-W. Chang (TESRI). S95 Chuyunshan Logging Trail (tBgLlj#^), 2220 m; KAOHSIUNG; 23°01'N, 120°49'E; drop- pings observed 20 Oct. 1998 by S.-W. Chang (TESRI). S56 Chuyunshan Logging Trail (Ht4ffU4#?I), 2260-2275 m; KAOHSIUNG; 23^0 1'N, 120°48'E; 2 monkeys observed at 2260 m and droppings observed at 2275 m 20 Oct. 1998 by S.-W. Chang (tesri). S55 50 FIELDIANA: ZOOLOGY Chuyunshan Logging Trail ( tlj ff [1( # jI ), 2300-2320 m; KAOHSIUNG; 23°00'N, 120°48'E; vocalization heard at 2300 m and droppings observed at 2320 m 20 Oct. 1998 by S.-W. Chang (TESRi). S54 Danayee ( ^ ^ f^ ^ ) (Shanmei), ?Alishan Hsiang; 690-930 m; CHIA-I; 23°22'N, 120°39^1'E; droppings observed at 690 m, 720 m, 730 m, 770 m, and 820 m 1 Apr. 1998 by TESRI survey team. Droppings observed at 930 m 5 Apr. 1998 by TESRI survey team. S3 Dikanhsi-Hsuehchenhsi (^^M — SM^), Taian Hsiang, 820-860 m; MIAOU; ca. 24°26'N, 120°54'E; droppings observed 3 Dec. 1996 by TESRI survey team. N37 Eastern coast, mountains ( ^ pi! ^ ^ [Jj M ), "principally in the central district"; imprecise locality; collected in 1953 by local Ami resi- dents (Satoh et al., 1956, p. 42; Setoguti & Sakuma, 1959, p. 158); Formerly in Depart- ment of Anatomy, Nagasaki University, 116 preserved cadavers (not seen); present whereabouts unknown. Not mapped. Eastern Taiwan {^M.W), <2000 m; imprecise locality; collected before 1999 by unknown collector; NTUZ, 3 (skulls only). Not mapped. Erpingshan (Zlt^[JL(), Shuili Hsiang, 390 m; NANTOU; 23°49'N, 120°51'E; 6 monkeys observed 12 Mar. 1994 by TESRI survey team. N103 Ershui Monkey Protection Area (— 7K^f^f:^|M M. ), Ershui Hsiang, 170-210 m; CHANGHUA; 23°49'N, 120°38'E; 20 drop- pings observed at 170 m 5 Jan. 1995 by TESRi survey team. Food remains observed at 170 m and 2 1 0 m 5 Jan. 1 995 by TESRI survey team. Ten monkeys observed at 170 m 6 Jan. 1995 by TESRI survey team. Droppings observed at 200 m 8 Mar. 1995 by TESRI survey team. Nlll Fangliao vicinity (|:5^%M); PINGTUNG; ca. 22°22'N, 120°35'E; serum samples obtained 1962-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967, p. 146). S133 Fangshan Hsi (1^5 [Ij M ), Fangshan Hsiang; PINGTUNG; ca. 22°25'N, 120''37'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka(1986, p. 386). 5/5^ Fayun Ssu (t^g^), Tahu Hsiang, 310-390 m; MIAOLI; 24°27'N, 120^5 1'E; 10 monkeys observed at 310 m 29 Jan. 1997 by tesri sur- vey team. Droppings observed at 390 m 30 Jan. 1997 by tesri survey team. N34 Fengshan ( S (JU ), Alishan Hsiang, 780 m; CHIA-I; 23°34'N, 120°44'E; 2 groups ob- served 4 Dec. 1997 by TESRI survey team. N144 Forest Section No. 15 (-hS#BI), Mutan Hsiang; PINGTUNG; ca. 22°07'N, 120°49'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). SI 49 Forest Section No. 27 (:^-h-t:#S£), Mutan Hsiang; PINGTUNG; ca. 22°07'N, 120°49'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). SI 49 Fuchouli (s'JtIM), Chushan Chen, 250 m; NANTOU; 23°48'N, 120°45'E; droppings observed 5 Jan. 1993 by tesri survey team. N109 Fuhsing Hsiang CiKMM); TAOYUAN; ca. 24°43'N, 12r22'E; reported Jul.-Nov. 1985 by S. Tanaka (1986, p. 98). N14 Fushan Forest Station ( # git P/f li [ll 5^ ^If ); ILAN/TAIPEI; ca. 24°46'N, 12r35'E; ob- served Nov. 1992-Jul. 1993 by L.-L. Lee (1994, p. 437). TV// Fushan Forest Station: Botanic Garden (^fitiP/r iJfgUjtfif^llI), west of Hapen Creek (D^^r^); TAIPEI; 24°46'N, 121°35'E; observed 25 Apr 1999 by H.-H. Su and J. Fooden. Nil Fushan (;|gUj) (=Rimogan); TAIPEI; 24°47'N, 121°30'E; reported before 1933 by Y. Hori- kawa(1932, p. 69).W2 Fuyuan Forest Recreation Area iWM^WM^ IS), Wanjung Hsiang, 200 m; HU ALIEN; 23°36'N, 121°20'E; reported 1 Oct. 1999 to H.-Y. Wu. N139 Giran. See Ilan. Haianshanmo C^S^UjM), Tungho Hsiang. See Tayuan, Haianshanmo. Haianshanmo {M ^ \U M ), Yuli Chen; HUALIEN; ca. 23°25'N, I21''25'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 5*27 Ha-pen (fl^^), Wulai Hsiang, 530 m; TAIPEI; 24°45'N, 121°32'E; 4 monkeys observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N15 Ha-pen (B^^), Wulai Hsiang, 535-570 m; FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 51 TAIPEI; 24°45'N, 12r33'E; groups observed at 535 m, 550 m, and 570 m 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N16 Ha-pen (P^^), Wulai Hsiang, 650 m; TAIPEI; 24°45'N, 121°34'E; 4 monkeys observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N17 Hapen Creek (Dn^M)- See Fushan Forest Sta- tion. Hengchun ( [fi # ) (=Koshun; Koshiun); PINGTUNG; 22°00'N, 120°44'E; reported before 1930 by T. Kano (1929, p. 333). Col- lected 4-6 Jun. 1932 by H. Orii (Kuroda, 1935, p. 291; 1940, p. 269); YIO, 2. Serum samples obtained 1962-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967, p. 146). SI 42 Hengchun Nature Preserve (1fi#^llf:^^M), Renting National Park, 200-250 m; PINGTUNG; 21°58'N, 120^49^; observed 1985-1990 by H.-Y. Wu and Y.-S. Lin (1992, p. 86; Wu, Lin & Lee, 1991, p. 29). SI 53 Hoko I. See Penghu Chuntao. Hokuzanko. See Peishankeng. Hori. See Peishankeng. Hoping (fn^), Papien, Chiapin, Chengkung Chen; TAITUNG; ca. 23*^04 'N, 121°20'E; observed in 1982 by K. Masui, Y. Narita, and S. Tanaka(1986, p. 386).57<^ Hsiangtao No. 131 {^i^U\WMM\ Shuili Hsiang; NANTOU; ca. 23°48'N, 120°51'E; observed 14 Apr. 1994 by tesri survey team. N104 Hsiangtao No. 136 ((^1^1365^1,^), Kuohsing Hsiang, 600 m; NANTOU; 24°03'N, 120°51'E; reported 14 Oct. 1993 to TESRi survey team. N92 Hsiao Ching Shui ( /Jn /f tR ); HUALIEN; 24°12'N, 121°40'E; collected 15 Feb. 1997 by K.-Y. Chung; NMNS, 1 (skull only). N54 Hsiaoi (#ft), Wulai District; TAIPEI; 24°51'N, 121°35'E; trapped for parasitological research 6 Jun. 1963 by J. F. Bergner Jr., autopsied 22 May 1964 (Bergner & Jachowski, 1968, pp. 22, 30); USNM (skull only). N7 Hsiao Kueihu {^h%M)\ TAITUNG; 22°42'N, 120°53'E; reported Jul.-Nov. 1985 by S. Ta- naka(1986, pp. 97, 98).57/7 Hsiaoponko (/Jn#[5P), ?Tahu Hsiang; MIAOLl; 24°23'N, 120°52'E; reported 13 Aug. 1996 to TESRI survey team. N36 Hsi Lin (±#); TAIPEI; ca. 25°06'N, 12r32'E; reported Jun.-Dec. 1978 by informant to F. E. Poirier (Poirier & Davidson, 1979, p. 157). NI Hsini (fs^), Tousienchiao, Chengkung Chen; TAITUNG; 23°02'N, 121°19'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). 579 Hsinkangshan ( 0f ^ I-U ), Chengkung Chen; TAITUNG; ca. 23°08'N, 121°19'E; reported 1967-1971 by K. Masui, Y. Narita, and S. Tanaka(1986, p. 386). 575 Hsipao ( H ff ), Hsiulin Hsiang, 1000 m; HUALIEN; 24°12'N, 121°28'E; observed I Aug. 1997 by H.-Y. Wu. N50 Hsiton. See Hsitou. Hsitou (i^Hi), Luku Hsiang, 1270-1360 m; NANTOU; 23°40'N, 120°47'E; droppings observed at 1360 m 26 Apr. 1995 by tesri survey team. Droppings observed at 1270 m 27 Apr. 1995 by TESRI survey team. Four monkeys observed at 1320 m 28 Nov. 1994 by TESRI survey team. Droppings observed at 1280 m 30 Nov. 1994 by TESRi survey team. N120 Hsitou iWM) (=Chi Tou), 1000 m; NANTOU; 23°41'N, 120M7'E; observed Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145; Masui et al., 1986, p. 386). N120 Hsitou (?^Si) (=Hsiton), Fenghuangshan, Luku Hsiang; NANTOU; 23°4rN, 120°47'E; ob- served in 1982 by K. Masui, Y. Narita, and S. Tanaka(1986, p. 386). 7V720 Hsiungshan (?), "Chishan ?" Chen; KAOHSIUNG; ca. 22°50'N, 120°25'E; albino infant reported in 1982 by K. Masui, Y. Na- rita. and S. Tanaka (1986, p. 387); report subsequently contradicted (Tanaka, 1986, p. 100). Not mapped. Hsuehshankeng ( § [1] $t ), Hoping Hsiang, 750-880 m; TAICHUNG; 24°19'N, 120°57'E; vocalization heard at 750 m 26 Oct. 1995 by TESRI survey team. Droppings observed at 880 m 7 Feb. 1996 by TESRI survey team. N80 Hsuehshankeng Logging Road {W-\U^J\M^W), Hoping Hsiang, 1200 m; TAICHUNG; 24°19'N, 120°57'E; droppings observed 10 Jan. 1996 by TESRi survey team. N80 52 FIELDIANA: ZOOLOGY Hsueh Shan (MlM) (^Tsugitaka Mountains), 0-3000 m, mostly <2100 m; TAICHUNG; ca. 24°23'N, 12n3'E; reported 1925-1933 by T. Kano(1940, pp. 10, 33). TV^^ Hualien ( fg ^ ); HUALIEN; ca. 23°59'N, 121''36'E; semm samples obtained 1962-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967, p. 146). Reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). N57 Hualien (^'g^), mountains near; HUALIEN; ca. 23°59'N, 12r36'E; collected 1 Apr. 1960 by R. B. White; AMNH, 1 (external measure- ments in field catalogue). N57 HUALIEN forests ( ft S % M ^ # ); 23°06'-24''22'N, 120°59'-121°46'E; reported before 1959 by Z.-M. Dien (1958, p. 346). Not mapped. Huayuan (WM), Taimali Hsiang; TAITUNG; 22"40'N, 12r01'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). SI 23 Hui-Sun Experimental Forest (MWM^^^M)', NANTOU; 24°03'N, 12r01'E; monkey acci- dentally poisoned in 1994, collected by H.-Y. Wu; private collection, 1 (skull with partial postcranials). N72 Hungshih Logging Trail (IQIS^^jM), Haituan Hsiang, 1160-1170 m; TAITUNG; 23°04'N, 121°07'E; vocalization heard at 1160 m and droppings observed at 1 170 m 4 Dec. 1996 by S.-W. Chang (tesri). S67 Huoyenshan (ikj^lU), Sani Hsiang; MIAOLl; 24°22'N, 120°43'E; reported Jul.-Nov. 1985 by S. Tanaka (1986, p. 98). N35 Hupen (M^X Linnei Hsiang, 210 m; YUNLIN; 23°43'N, 120°37'E; droppings observed 12 Jan. 1995 by TESRi survey team.NJJS Ilan (g|0) (=Giran) region; ILAN; ca. 24''46'N, 12r45'E; collected 25 Nov. 1930 by B. Aoki; ZMB, 1. Serum samples obtained 1959-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967, p. \46).N9 Jenlun Logging Trail (Afw^f^iS), Hsini Hsiang, 1300-1400 m; NANTOU; 23°40'N, 120°53'E; reported 27 June 1997 to S.-W. Chang (TESRi). A^/22 Jenlun Logging Trail (Afra^sM), Hsini Hsiang, 1400 m; NANTOU; 23°44'N, 120°54'E; re- ported 26 June 1997 to S.-W. Chang (tesri). NJ24 Jenlun Logging Trail (Af##3l), Hsini Hsiang, 1750 m; NANTOU; 23°43'N, 120°56'E; drop- pings observed 11 Jan. 1994 by S.-W. Chang (TESRI). N125 Jentse (iZW); ILAN; 24°33'N, 121°30'E; re- ported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). Collected in 1991 by H.-H. Su; NTUZ, 1 (skeleton only). N26 Jentse Wenchuan. See Jentse. Jihyueh Tan (S^~M) (=Jihyuhtan, Sun-Moon Lake, Tsuisia Lake) vicinity; NANTOU; ca. 23°5rN, 120°55'E; monkey-skin clothing and strings of monkey skulls observed ca. Oct. 1873 by J. B. Steere (1876, pp. 310, 319). Two captives reported Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145; Masui et al., 1986, p. 387). N102 Juifeng Bridge (^lllt^fg), Meishan Hsiang, 420 m; CHlA-1; 23''34'N, 120°38'E; >30 monkeys observed 17 Oct. 1997 by TESRI survey team. N146 Juili (JroM), Meishan Hsiang; CHIA-I; ca. 23°33'N, 120''38'E; reported in 1975 by S. Tanaka (1986, p. 98); confirmed as locally extinct in \9S6.NI49 Juili (^M), Meishan Hsiang, 700 m; CHIA-I; 23°31'N, 120°40'E; 5 monkeys observed 4 Dec. 1997 by TESRI survey team. N150 Juilung Pupu (imWM'ft^), Chushan Chen, 320 m; NANTOU; 23°39'N, 120''41'E; 2 monkeys observed 10 Mar. 1993 by TESRI survey team. Droppings observed 11 Mar. 1993 by TESRI survey team. NI 18 Juiyen (51©^), Jenai Hsiang, 2200 m; NANTOU; 24°07'N, 121°irE; >2 monkeys observed 26 Oct. 1993 by tesri survey team. N66 Juiyenhsi (^^M), Jenai Hsiang, 2190-2205 m; NANTOU; 24°09'N, 12n2'E; droppings or tracks observed at 2190 m and droppings ob- served at 2205 m 28 Feb. 1997 by S.-W. Chang (TESRI). N66 Kainan City. See Moushi. Kangou (^M), Kuohsing Hsiang, 630 m; NANTOU; 24°01-02'N, 120°49-50'E; drop- pings observed 14 Oct. 1993 by TESRI survey team. N94 Kangtzu (^f?), Manchou Hsiang; PINGTUNG; 22°08'N, 120°52'E; reported in 1982 by K. FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 53 Masui. Y. Narita, and S. Tanaka (1986. p. 386). SI 50 Kankenghsi (^^Ml Chushan Chen. 260 m; NANTOU: 23''40'N. 120°39'E; droppings observed 10 Mar. 1993 by tesri survey team. Vocalization heard and droppings observed 11 Mar. 1993 by tesri survey team. One monkey observed 23 Apr. 1993 by tesri sur- vey team. NI 15 Kaohsiung (j^S^) (=Takao) vicinity, probably obtained on Shou Shan: KAOHSIUNG: ca. 22''39'N. nO'-'lS'E; collected 7 Mar. 1953 by R. M. Dasch: USNM, 2. S106 KAOHSIUNG forests ( ffi ^ ife ^ ^ # ): 22''29'-23''28'N. 120"'10'-121°02'E: reported before 1959 by Z.-M. Dien (1958. p. 346). Not mapped. Kaoshih {M±l Mutan Hsiang; PINGTUNG: 22°07'N, 120''50'E: reported in 1982 by K. Masui, Y. Narita. and S. Tanaka (1986. p. 386). S149 Kaoshihfoshan ( r^ dr f^ Uj ). Mutan Hsiang: PINGTUNG: 22°08'N. 120°48'E: reported in 1982 by K. Masui. Y. Narita. and S. Tanaka (1986. p. 386). 5745 Kasho I. See Lu Tao. Kenting (^T). 225 m: PINGTUNG: 21'58'N. 120''49'E: collected in 1987 by H.-Y. Wu: private collection. 1 (skull only). SI 53 Kenting Kung\-uan (^Ti^^) (=Park). Heng- chun Chen:^PrNGTUNG: 2r58'N. 120'48'E: reported in 1982 by K. Masui. Y. Narita. and S. Tanaka (1986. p. 386). Observed 1987-1989 by L.-L. Lee and Y. S. Lin (1991. p. 36: 1994. p. 19). Collected in 1989 by H.-Y. Wu and collected in 1998 by J.-N. Yue: pri- vate collection. 2 (skulls only). SI 43 Kenting Monke> Rock. See Kenting Kung\nan. Koshiun. See Hengchun. Koshun. See Hengchun. Koto I. See Lan Yu. Kuangfti vicinit> (Ttllife^); HUALIEN: ca. 23'40'N. 12r'25'E. reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386).A^/i7 Kuang Shan Working Circle. See Kuanshan. Kuanshan (HlJj) (=Kuang Shan Working Cir- cle): TAITUNG: 23°03'N, 12I''09'E: reported Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145; Masui et al., 1986, p. 386). S68 Kuanshan Chen vicinity (M iU M ^1k^ )'• TAITUNG: ca. 23°03'N, 12I°09'E; reported in 1982 by K. Masui. Y. Narita. and S. Ta- naka (1986, p. 386). 565 Kuanyinshan (^#[14), Tashe Hsiang, 200 m: KAOHSIUNG: 22°43'N, 120^2 1'E^ reported 15 Feb. 2000 to H.-Y. Wu. SI 07 Kueihu. See Hsiao Kueihu. Kueitan ( ^ :^ ). Nanhsi Hsiang: TAINAN: 23'08'N. 120*'29'E: Jul.-Nov. 1985 by S. Ta- naka (1986. p. 98). 557 Kukeng (^i/C): YUNLIN: 23''39'N. 120°33'E: reported origin of captive obtained before 1998 (Chang 1997. p. 3). N114 Kukuan (-gfli). Hoping Hsiang: TAICHUNG: 24°12'N. 12r00'E: reported in 1982 by K. Masui. Y. Narita. and S. Tanaka (1986. p. 386). N-5 Kungtientsun (:S-B3)S). Fanlu Hsiang. 590 m; CHIA-I: 23°22'N, 120°36'E; >10 monkeys observed 10 Feb. 1998 by tesri survey team. One dead monkey observed 18 June 1998 by TESRI survey team. S2 Kuohsing Hsiang (mttiM): NANTOU: ca. 24''0rN, 120^^5 1'E: collected before 1999 by TESRI recoven. center (S.-W. Chang, TESRI, personal communication, 27 Apr. 1999); TESRI. 1 (frozen carcass). N93 Kuwabarusha (?): KAOHSIUNG (=Takao); not located; observed before 1930 by T. Kano (1929, p. 333). Not mapped. Kwang Fu. See Yulishan. Lakulakuhsi Hiking Trail (^W-tiL^M^ih^ B)- Chohsi Hsiang, 250-420 m; HUALIEN; 23^19'^ 12ri4'E: vocalizations heard at 250 m and 420 m 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S33 Lakulakuhsi Hiking Trail (U^^^M^lUB ^). Chohsi Hsiang. 710 m; HUALIEN; 23''20'N. 12n2'E:^2 monkeys observed 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S36 Lakulakuhsi Hiking Trail i^mt^MM^iU^ 1^). Chohsi Hsiang. 800 m; HUALIEN: 23°21'N. 12ri6'E:^4 monkeys observed 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S25 Lakulakuhsi Fliking Trail (Sl^Sffr^^lll^ H). Chohsi Hsiang. 850 m; HUALIEN: 54 FIELDIANA: ZOOLOGY 23°20'N, 121°11'E; 1 monkey observed 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S37 Lakulakuhsi Hiking Trail {W^-'&W-U^ll^^ H), Chohsi Hsiang, 950-1030 m; HUALIEN; 23°20'N, 12riO'E; 1 monkey observed at 950 m and 1 monkey observed at 1030 m 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S37 Lakulakuhsi Hiking Trail {^WMlWM^IU^ U), Chohsi Hsiang, 1200 m; HUALIEN; 23°25'N, 121°07'E; droppings observed 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. SI 9 Lakulakuhsi Hiking Trail {t^iW-'&MM^iUl^ U. ), Chohsi Hsiang, 1220-1320 m; HUALIEN; 23°22'N, 121°05'E; droppings observed at 1220 m, 1250 m, and 1320 m 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S41 Lakulakuhsi Hiking Trail {W$-'&MM^\U^ H), Chohsi Hsiang, 1300 m; HUALIEN; 23°24'N, 12r05'E; droppings observed 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S16 Lakulakuhsi Hiking Trail {WKlW^M^l^M 1^), Chohsi Hsiang, 1300 m; HUALIEN; 23°24'N, 12r07'E; droppings observed 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S20 Lakulakuhsi Hiking Trail {^WMlM^^M^ ^$. ), Chohsi Hsiang, 1300-2205 m; HUALIEN; 23°24'N, 121°08'E; droppings observed at 1300 m and vocalization heard at 2205 m 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S20 Lakulakuhsi Hiking Trail {^WMll '^^ m^ U. ), Chohsi Hsiang, 1420-1570 m; HUALIEN; 23°25'N, 12r05'E; droppings observed at 1420 m and 1570 m 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. SI 7 Lakulakuhsi Hiking Trail (S0fci)¥r^^[l(g^ U. ), Chohsi Hsiang, 1455-1735 m; HUALIEN; 23°24'N, 121°06'E; vocalization heard at 1455 m and 1 monky observed at 1735 m 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S18 Lakulakuhsi Hiking Trail (fczl¥fcil^rMg[ll£^ H), Chohsi Hsiang, 1478 m; HUALIEN; 23°24'N, 121^09^; 2 monkeys observed 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S21 Lakulakuhsi Hiking Trail {^W&WM^M^ 1^ ), Chohsi Hsiang, 1490-1520 m; HUALIEN; 23°23'N, 12r05'E; droppings observed at 1490 m, 1510 m, and 1520 m 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. Vo- calization heard at 1490 m 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S41 Lakulakuhsi Hiking Trail {tsLWMlWM^lU^ 1^), Chohsi Hsiang, 1620 m; HUALIEN; 23°2rN, 121°09'E; vocalization heard 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S38 Lakulakuhsi Hiking Trail {t^LW-'&MM^\U^ 1^), Chohsi Hsiang, 1630 m; HUALIEN; 23°21'N, 121°10'E; 1 monkey observed 1 Mar 1997 by L.-L. Lee and H.-Y. Wu. S23 Lakulakuhsi Hiking Trail (MlMWMM^\U^ ID, Chohsi Hsiang, 1690 m; HUALIEN; 23°2rN, 121°08'E; vocalization heard 1 Mar. 1997 by L.-L. Lee and H.-Y. Wu. S39 Lamhongo. See Suao vicinity. Langkeaou. See Lungkeaou. Lanjen Hsi ( j^ {H ^ ), Manchou Hsiang; PINGTUNG; 22°02'N, 120°50'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). 5/^^ Lan Yu (M^) (=Koto 1.); TAITUNG; ca. 22°03'N, 121°32'E; reported absent before 1931 byT. Kano(1930, p. 165). D Laofoshan ( ^ f^ [Jj ), Manchou Hsiang; PINGTUNG; 22°02'N, 120°47'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 387). 5/^5 Lichia ( ^(J M ), Peinan Hsiang; TAITUNG; 22°47'N, 121°03'E; observed in 1967-1971 and 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 57/^ Lichia Logging Trail ( ^fj ^ # il ), Peinan- Yenping Hsiang, 930-950 m; TAITUNG; 22°48'N, 121°02'E; droppings or food remains observed 5 Mar. 1998 by S.-W. Chang(TESRl). S'/Zd Lichia Logging Trail ( ^'J M # ?1 ), Peinan- Yenping Hsiang, 970-1045 m; TAITUNG; 22°48'N, 12r01'E; 2 monkeys observed at 970 m 3 Dec. 1996 by S.-W. Chang (TESRi). Droppings or food remains observed at 1010-1045 m 5 Mar. 1998 by S.-W. Chang (tesri). SI 16 Lichia Logging Trail ( ^IJ ^ # M ), Peinan- Yenping Hsiang, 1055-1100 m; TAITUNG; 22°49'N, 121°01'E; 5 monkeys observed at 1075 m 3 Dec. 1996 by S.-W. Chang (tesri). Droppings or food remains FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 55 observed at 1080-1100 m 4 Mar. 1998 by S.-W. Chang (tesri). Two monkeys observed at 1055 m and droppings observed at 1070 m 5 Mar. 1998 by S.-W. Chang (tesri). SI 15 Lichia Logging Trail ( ^IJ M # il ), Peinan-Yenping Hsiang, 1110-1130 m; TAITUNG; 22°49'N, 121°00'E; droppings or food remains observed 4 Mar. 1998 by S.-W. Chang (TESRI). 5/75 Lichia Logging Trail ( ^0 ^ # il ), Peinan-Yenping Hsiang, 1175-1370 m; TAITUNG; 22''50'N, 121°00'E; one monkey observed at 1290 m 3 Dec. 1996 by S.-W. Chang (tesri). Droppings or food remains observed at 1175-1200 m 4 Mar. 1998 by S.-W. Chang (TESRi). Groups observed at 1210 m, 1320 m, and 1370 m 4 Mar. 1998 by S.-W. Chang (TESRi). Tracks observed at 1220 m and 1355 m 4 Mar. 1998 by S.-W. Chang (TESRI). 57/5 Lichia Logging Trail ( ^0 ^ # H ), Peinan-Yenping Hsiang, 1400-1475 m; TAITUNG: 22°50'N, 120''59'E; groups ob- served at 1400 m and 1475 m 4 Mar. 1998 by S.-W. Chang (TESRI). 5772 Lichia Logging Trail ( ^IJ M # il ), Peinan-Yenping Hsiang. 1420 m; TAITUNG; 22°5rN, 120°59'E; vocalization heard 4 Mar. 1998 by S.-W. Chang (TESRi). S94 Lichia Logging Trail ( f 0 M # jI ), Peinan-Yenping Hsiang, 1435-1565 m; TAITUNG; 22''50'N, 120''58'E; groups ob- served at 1435 m and 1525 m 3 Dec. 1996 by S.-W. Chang (tesri). Vocalization heard at 1550 m and droppings observed at 1550 m and 1565 m 3 Mar. 1998 by S.-W. Chang (TESRI). 5772 Lienhuachih { M ^ 1^ ), Hsiulin Hsiang, 800-1100 m; HUALIEN; 24°13'N, 121°28'E; observed at 800 m 3 Aug. 1997 and at 1 100 m 4 Aug. 1997 by H.-Y. Wu. N49 Lienhuachih ( S ffc ilfe ), Hsiulin Hsiang, 1 100-1200 m; HUALIEN; 24°13'N, 121°29'E; observed at 1200 m 5 Aug. 1997 and at 1 100 m 6 Aug. 1997 by H.-Y. Wu. N49 Lienhuachih ( ^ 20 monkeys observed 1 1 Aug. 1 993 by TESRI survey team. NlOl Lungkeaou (?) (=Langkeaou); PINGTUNG; ca. 22°05'N, 120°43'E; observed Jul. 1864 by R. Swinhoe (1865, p. 380; 1866, p. 127). S141 Lupishan Logging Trail (J^^lJ4#xi), Nanao Hsiang, 245 m; ILAN; 24°27'N, 12r44'E; 2 monkeys observed 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N22 Lupishan Logging Trail {]^J^\i\WM), Nanao Hsiang, 595-798 m; ILAN; 24''27'N, 121°43'E; 1 monkey observed at 595 m and droppings observed at 798 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N22 Lu Tao (It ft) (=Kasho L); TAITUNG; ca. 22''40'N, 121°29'E; reported absent before 1931 byT. Kano(1930, p. 165). C Maanliao {MMiW:) (=Ma-an Lio), 6 mi (=10 km) south of Tungshih (=Tung Shi), 490 m; TAICHUNG; 24°10'N, 120°49'E; collected 16 Apr. 1952 by D. H. Johnson; USNM, 1. N86 Ma-an Lio. See Maanliao. Mali River Reserve (?), contiguous with Shan- ping; KAOHSIUNG; not located; reported Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145; Masui et al., 1986, p. 386). Not mapped. Manchoutsun (ri'jfO (=Teraso); PINGTUNG; 22°0rN, 120°5rE; collected Oct. 1908 and Dec. 1908-Feb. 1909 by H. Sauter (cf Huang et a!., 1993, p. 167); SMTD, 1; ZMB, 20 (in- cluding 7 skulls only and 6 skeletons only). S152 Meishan (^|JL|), Taoyuan Hsiang, >1500 m; KAOHSIUNG; 23°17'N, 120°49'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386; Tanaka, 1986, p. 101). S47 Meiyuan (^H), Hsiulin Hsiang, 1000 m; HUALIEN; 24°14'N, 12r28'E; observed 7 Aug. 1997 by H.-Y. Wu. N49 Mikawa-Wan. SeeNojima. Minsheng ( .K ^ ), Sanmin Hsiang; KAOHSIUNG; 23°17'N, 120°43'E) reported Jul.-Nov. 1985 by S. Tanaka (1986, p. 98). S48 Mount Shaushan. See Shou Shan. Moushi, Kainan City, Wakayama Prefecture, Honshu Island, Japan; ca. 34''09'N, 135°12'E; introduced population, escaped from local menagerie ca. 1960, feral individuals report- edly shot by hunters (Wakayama Prefecture, 1978, p. 2; Maekawa, 1982, p. 102; Kawa- moto et al., 1999, p. 53). Not mapped. Mt. Longevity. See Shou Shan. Mucha (^i); TAIPEI; 24°59'N, 121''34'E; reported Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145; Masui et al., 1986, p. 386). ^5 Mu-Kua-Chau (?); HUALIEN; ca. 23°50'N, 121°33'E; collected 3 Apr. 1960 by R. B. White (see field catalogue); AMNH, 2. N58 Mutan ( f± :f^ ) (=Sekimon; Shihmen); PINGTUNG; 22°08'N, 120°46'E; reported before 1933 by Y. Horikawa (1932, p. 69). S140 Mutsu. See Shimokita Hanto. Nakatsu, Wakayama Prefecture, Honshu Island, Japan; ca. 33°57'N, 135°16'E; probable hy- bridization reported between introduced feral M. cyclopis and native M. fuscata (Kawamoto et al. 1999, p. 57). Not mapped. Nanhua Hsiang {"^itM); TAINAN; ca. 23°07'N, 120''33'E; reported in 1957 by S. Tanaka (1986, p. 98); confirmed as locally extinct in 1986. Collected 11 Sep. 1996 by TESRI recovery center; TESRI, 1 (frozen car- cass). S52 Nanhu Tashan {'^Mi^iUX Hoping Hsiang, 3095-3165 m; TAICHUNG; 24°22'N, 121°25'E; droppings observed at 3095 m and 3130-3165 m 23 Nov. 1995 by TESRI survey team. N46 Nanjenlu-Kangtzu (f^{r^#-i#f?), Manchou Hsiang; PINGTUNG; ca. 22°07'N, 120°53'E; reported in 1982 by K. Masui, Y. Narita, and FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 57 S. Tanaka (1986, p. 386). SI 50 Nanshanhsi (]^[jL|i^), Jenai Hsiang; NANTOU; 24°00'N, 12r03'E; reported 11 May 1994 to TESRl survey team. N70 Nanshanhsi (hlMM), Jenai Hsiang, 1300-1580 m; NANTOU; 24°00'N, 121°03'E; vocaliza- tion heard at 1300 m and 1580 m 29 Dec. 1994 by TESRI survey team. Droppings ob- served at 1420 m 29 Dec. 1994 by tesri sur- vey team. Vocalization heard at 1530 m 31 Dec. 1994 by TESRI survey team. N70 Nanshi Logging Road (^l^^jS), Yushan Na- tional Park, 1730-2670 m; KAOHSIUNG; 23''24-28'N, 120°53-54'E; observed Mar. 1987-Oct. 1988 by J.-F. Lu, Y.-S. Lin, and L.-L. Lee(1991,p. 93). 5/^ 7NANT0U (j^lx) (county uncertain); acquired before Dec. 1992 by Phoenix Valley Aviary; NMNS, 1 (skeleton only; external measure- ments in computerized catalog). Not mapped. Nantzuhsien Hsi Logging Road (^#fLliM#xl), Alishan Hsiang, 1700-2120 m; CHIA-I; 23°27'N, 120°53'E; food remains observed at 1940 m 7 Oct. 1997 by tesri survey team. Droppings observed at 2010 m and 10 mon- keys observed at 2120 m 7 Oct. 1997 by TESRI survey team. Droppings observed at 2120 m 8 Oct. 1997 by TESRi survey team. One monkey observed at 1700 m 24 June 1998 by TESRi survey team. SJ3 Nengkao-Tungmen Powerline Right-of-way (ft mmn-^nm^M^), HsiuUn Hsiang, 650 m; HU ALIEN; 24°00'N, 12r24'E; droppings observed 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N59 Nengkao-Tungmen Powerline Right-of-way (ft mmn^nU^^mmi Hsiulin Hsiang, 750-1200 m; HUALIEN; 24°00'N. 121°23'E; droppings observed at 750 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. Two groups ob- served at 820 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. Vocalization heard at 910 m and 1200 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N60 Nengkao-Tungmen Powerline Right-of-way (fg Mmn-^n^mmm), Hsiulin Hsiang, 900-1200 m; HUALIEN; 24°00'N, 121°22'E; vocalization heard at 900 m and 1200 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N60 Nengkao-Tungmen Powerline Right-of-way (ft Mmr'^-^M^W^m.mi HsiuHn Hsiang, 1200-1310 m; HUALIEN; 24°0rN, 121°22'E; vocalization heard at 1200 m, 1300 m. and 1310 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. Groups observed at 1220 m and 1250 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. Droppings observed at 1300 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N6I Nengkao-Tungmen Powerline Right-of-way (ft Mmn-^nm^M^l Hsiulin Hsiang, 1300 m; HUALIEN; 24°01'N, 121°21'E; droppings observed 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N62 Nengkao-Tungmen Powerline Right-of-way (tt i^Sir^-^'Bl^liMl^), Hsiulin Hsiang, 1300 m; HUALIEN; 24°02'N, 12r22'E; 4 monkeys observed 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N61 Nengkao-Tungmen Powerline Right-of-way (fb mmn^nm^MBX HsiuHn Hsiang, 1300-1340 m; HUALIEN; 24°02'N, 121°20'E; groups observed at 1300 m, 1310 m, and 1340 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. Droppings observed at 1320 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N63 Nengkao-Tungmen Powerline Right-of-way (ft mmnt^nm^M^), HsiuUn Hsiang, 1345 m; HUALIEN; 24°01'N, 121°20'E; 4 monkeys observed 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N63 Nengkao-Tungmen Powerline Right-of-way (ft mmn-^m^K^mmi HsiuUn Hsiang, 1840-2085 m; HUALIEN; 24°01'N, 12ri8'E; droppings observed at 1840 m and 2085 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. Vo- calization heard at 1930 m and 1975 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N64 New Central Cross-Island Highway (^f^'tS^ £^ ), Alishan Hsiang, 1395 m; CHIA-I; 23°28'N, 120°42'E; >6 monkeys observed 16 May 1998 by TESRI survey team. S4 New Central Cross-Island Highway (0f4^tM^ g^ ), Alishan Hsiang. 1655 m; CHIA-I; 23°29'N, 120^45^; 12 monkeys observed 8 Jan. 1998 by tesri survey team. S7 New Central Cross-Island Highway (iff^^t^^i^ f#), Alishan Hsiang, 1860-2700 m; CHIA-I; 23°29'N, 120°48'E; droppings observed at 58 FIELDIANA: ZOOLOGY 2250 m 26 Sep. 1997 by tesri survey team. Groups observed at 1860 m, 2650 m, and 2700 m 30 Apr. 1998 by TESRI survey team. S8 New Central Cross-Island Highway (|fr4^fM^ {?§), Hsini Hsiang, 2000 m; NANTOU; 23°32'N, nO^SS'E; group observed 1 Aug. 1999byH.-Y. Wu. A^/¥7 New Central Cross-Island Highway (iff^'tM^ g§), Hsini Hsiang, 2300 m; NANTOU; 23°28'N, 120''49'E; 2 groups observed 1 Aug. 1999byH.-Y. Wu.^-P New Central Cross-Island Highway (^f^^tS^ B), Hsini Hsiang, 2300 m; NANTOU; 23''28'N, 120"50'E; 2 groups observed 1 Aug. 1999byH.-Y. Wu. 59 New Central Cross-Island Highway (§]t^^Wj^ g§), Hsini Hsiang, 2300 m; NANTOU; 23°28'N, 120°5rE; troop observed 1 Aug. 1999byH.-Y. Wu. 570 New Central Cross-Island Highway (§t^W.^/^ B), Hsini Hsiang, 2300 m; NANTOU; 23°28'N, 120''52'E; group observed 1 Aug. 1999byH.-Y. Wu. 5// New Central Cross-Island Highway (0f4^IM^ B), Hsini Hsiang, 2300 m; NANTOU; 23°29'N, 120°49'E; group observed 1 Aug. 1999byH.-Y. Wu. 5P New Central Cross-Island Highway (iFr4^tS:2r g§), Hsini Hsiang, 2300 m; NANTOU; 23°30'N, 120°52'E; group observed 1 Aug. 1999byH.-Y.Wu.yV/ 52 New Central Cross-Island Highway (0r4^tM:2^ g#), Hsini Hsiang, 2470 m; NANTOU; 23°32'N, 120^53^; 1 monkey observed 18 Oct. 1993 by TESRI survey team. N14] Niupi i^i^), Shuili Hsiang, 710-720 m; NANTOU; 23°30'N, 120°50'E; 14 monkeys observed at 710 m and droppings observed at 720 m 12 Mar. 1994 by TESRI survey team. N15] Noheji. See Shimokita Hanto. Nojima (island), Mikawa-Wan (bay), Aichi Pre- fecture, Japan; 34°39'N, 137°01'E; introduced by Japan Monkey Centre in 1958, free-ranging population observed Aug. and Oct. 1962 by T. Nishida (1963, p. 121). Ob- served 26 Feb.-20 Mar. 1970 by M. Kawai and U. Mito (1973, p. 180). Not mapped. Northern and eastern Taiwan, mountains (M^ Ma^Aita^lMM); imprecise locality; re- ported as abundant before 1863 by R. Swin- hoe (1863, p. 351). Not mapped. Northern Cross-Island Road (^tpPtSJ /jJ:\ r), Fuhsing Hsiang, 550 m; TAOYUAN; 24°45'N, 12r20'E; droppings observed 1 Sep. 1998 by L.-L. Lee and H.-Y. Wu. N13 Northern Cross-Island Highway (^bpBt^M^^A jg§), Fuhsing Hsiang, 1050 m; TAOYUAN; 24''39'N, 121°25'E; 1 monkey observed 1 Sep. 1998 by L.-L. Lee and H.-Y. Wu. N30 Northern Cross-Island Highway (4toWSM^ }?#), Tatung Hsiang, 460 m; ILAN; 24''37'N, 121°29'E; 6 monkeys observed 1 Sep. 1998 by L.-L. Lee and H.-Y. Wu. N28 Northern Cross-Island Highway (^tnl^t^M^ B), Tatung Hsiang, 720 m; ILAN; 24''36'N, 121°29'E; droppings observed 1 Sep. 1998 by L.-L. Lee and H.-Y. Wu. N29 Northern Cross-Island Highway (^bpPtMM^ B), Tatung Hsiang, 1258 m; ILAN; 24°36'N, 12r30'E; droppings observed 1 Sep. 1998 by L.-L. Lee and H.-Y. Wu. N27 Ohata. See Shimokita Hanto. Ohshima. See Oshima. Oshima (island; =Ohshima), eastern coast, Izu-Shoto (island group), Tokyo Prefecture, Japan; ca. 34''45'N, 139°26'E; introduced feral population, escaped from menagerie ca. 1940, reported before 1954 by K. Kishida (1953, pp. 24, 83; cf de Vos et al., 1956, p. 166). Ob- served 1988-1989 by K. Kazama (Kazama & Norikoshi, 1991, p. 51; Norikoshi, 1991, p. 138). Collected before 2000 by unknown col- lector; KUPRI, 1 (skull only). Not mapped. Otake (?); not located; reported before 1933 by Y. Horikawa (1932, p. 69). Not mapped. Pahsienshan Forest Recreation Area (AftllUj^ # 3l ^ M ), Hoping Hsiang, 890 m; TAICHUNG; 24°11'N, 121''00'E; droppings observed 19 Jul. 1995 by TESRI survey team. N74 Pailuz (?), Manchou Hsiang; PINGTUNG; ca. 22°04'N, 120°50'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S146 Paimao Shan Logging Road (S^Uj^jI), Hoping Hsiang, 850-1040 m; TAICHUNG; FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 59 24''09'N, 120''54'E; 3 monkeys observed at 870 m 20 Sep. 1995 by tesri survey team. Vocalization heard at 920 m 20 Sep. 1995 by TESRI survey team. Droppings observed at 850 m 21 Sep. 1995 by TESRI survey team. Mon- keys photographed with automatic camera at 1010 m 29 Dec. 1995 by tesri survey team. Droppings observed at 1010 m 6 Feb. 1996 by TESRI survey team Food remains observed at 1040 m 23 Apr. 1997 by S.-W. Chang (tesri). N89 Paimao Shan Logging Road ( fz3 ^IJl|#xS), Hoping Hsiang, 1140 m; TAICHUNG; 24°09'N, 120''55'E; droppings observed 23 Apr. 1997 by S.-W, Chang (tesri). N88 Paishanhang. See Peishankeng. Paiyun Hiking Trail {W9^\\^^W), Alishan Hsiang, 2820 m; CHIA-1; 23°28'N, 120°54'E; droppings observed 25 Sep. 1997 by TESRI survey team. SI 2 Pakhongo. See Suao vicinity. Pakuasan. See Pakua Shan. Pakua Shan { J\ ^Y [i\ ) (=Pakuasan); CHANGHUA; 23°54'N, 120°38'E; two cap- tives reported Jun.-Dec. 1978 by F. E. Poirier (Poirier &. Davidson, 1979, p. 145; Masui et al., 1986, p. 386). yV7 70 Patungkuan (AiSHfl), Hsini Hsiang, 2800 m; NANTOU; 23''29'N, 120''59'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386; Tanaka, 1986, p. 101). S15 Payao (y\JS), Manchou Hsiang; PINGTUNG; 22''06'N, 120°50'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). 57^7 Peinan (^|^) (=P'i-nan); TAITUNG; 22°46'N, 121°07'E; collected [ca. 1900-1901] by [K. A.]. Haberer (cf. Schweyer, 1909, p. 7); ZMB, 1.5777 Peishankeng (4 till la,) (=Hokuzanko; Paishan- hang), Puli (=Hori) district; NANTOU; 23°59'N, 120°52'E; collected before 1939 by Mr. Goto (Kuroda, 1938, p. 1 12; 1940, p. 269; cf. Huang et al., 1993, p. 135; Hill, 1974, p. 589); formerly in personal collection of Mar- quis N. Kuroda, 1 (probably destroyed by bombardment during World War II; cf Austin etal., 1948, p. 4). A^97 Peitunyenshan ( 4b ^ SM lij ), Jenai Hsiang; NANTOU; 24°04'N, 12r09'E; vocalization heard 10 May 1994 by tesri survey team. N67 Peitunyenshan ( :1b ^ IS ill ), Jenai Hsiang, 1700-1800 m; NANTOU; 24°04'N, 121°09'E; food remains observed at 1700 m 28 Jul. 1993 by TESRI survey team. Tracks and droppings observed at 1730 m 28 Jul. 1993 by TESRI sur- vey team. Monkeys observed at 1 800 m 3 Mar 1994 by TESRI survey team. N67 Peitunyenshan (^b^BSLU), Jenai Hsiang, 1900 m; NANTOU; 24°04'N, 121°15'E; 5 monkeys observed 15 Aug. 1993 by TESRI survey team. N65 Peiyuan {MtM\ Tungho Hsiang; TAITUNG; 23°01'N, 12ri7'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S80 Penghu Chuntao CM>M^Bj): Penghu Tao (^Pescadores Islands: Hoko I.); PENGHU; 23°11^5'N, 119''24^3'E; reported absent before 1931 by T. Kano (1930, p. 165; cf Lee &Lin,. 1991, p. 36; 1994, p. \9). A Pescadores Islands. See Penghu Chuntao. P'i-nan. See Peinan. Pingting ( i^ li ), Linnei Hsiang, 150 m; YUNLIN; 23°45'N, 120°38'E; droppings ob- served 1 1 Jan. 1 995 by TESRI survey team. 7V772 Pingting ( i^ IM ), Taiping Hsiang, 270 m; TAICHUNG; ca. 24°'07'N, 120°46'E; drop- pings observed 18 Jan. 1996 by tesri survey team. N90 Pingtung vicinity (P^ti^lS); PINGTUNG; ca. 22°40'N, 120°29'E; serum samples obtained 1962-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967, p. 146). S108 Pipitaishan (H^llj^llf), ?Taian Hsiang, 3215 m; MlAOLl; ca. 24°26'N, 120°54'E; droppings observed 30 Apr. 1997 by TESRI survey team. N37 Polisia. See Puli. Posia. See Puli. Puchung (igc^), Mingchien Hsiang, 200 m; NANTOU; 23°49'N, 120°38'E; droppings observed 5 Feb. 1993 by tesri survey team. 7V777 Puli (ifM) (=Posia; Polisia); NANTOU; 23°58'N, 120°57'E; collected Oct. 1873 by J. 60 FIELDIANA: ZOOLOGY B. Steere and Mr. Beal (Steere, 1876, p. 319); UMMZ, 1. Reported origin of two captives obtained before 1998 (Chang 1997, p. 3). N99 Puli vicinity ( if M ii 1^ ); NANTOU; ca. 23°58'N, 120°57'E; serum samples obtained 1964-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967, p. 146). Collected 14 Dec. 1987 by unknown collector; TM, 1 (mounted skin with skull fragment inside). N99 Rimogan. See Fushan. Sabkun Hsi (?), Manchou Hsiang; PINGTUNG; ca. 22°04'N, 120°50'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S146 Sakow. See Shou Shan. Sanhsia ( H ll^l^ ) (=Sung Sha); 500-800 m; TAIPEI; 24''56'N, 121°22'E; reported Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145; Masui et al., 1986, p. 386). N4 Sanhsienli ( H ftij M ), Chengkung Chen; TAITUNG; 23°07'N, 121°23'E; observed in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). 577 Sanminli ( H K M ), Chengkung Chen; TAITUNG; ca. 23°07'N, 12r21'E; observed in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 579 monkeys observed at 300 m 22 Feb. 1994 by tesri survey team. Seven or more monkeys ob- served at 300 m 23 Feb. 1994 by TESRI survey team. Droppings observed at 400 m 23 Feb. 1994 by TESRI survey team. N96 Shihmen. See Mutan. Shihmenshe ( S T^ (jth ), Tuanshushe, Mutan Hsiang; PINGTUNG; ca. 22°08'N, 120°48'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). 8148 Shihpi (BW), Kukeng Hsiang, 1010-1065 m; YUNLIN; 23°35'N, 120°4rE; droppings ob- served at 1010 m 9 Nov. 1994 by tesri sur- vey team. Droppings observed at 1030 m 11 Nov. 1994 by TESRi survey team. Droppings observed at 1065 m 14 Dec. 1994 by tesri survey team. N145 Shihting Hsiang vicinity (BW^UtM); TAIPEI; ca. 24°59'N, 12r39'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). N2 Shihtoushan (JiSlUj), Hsintien Chen, 700-800 m; TAIPEI; 24°54'N, 121°29'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). ^5 Shimokita Hanto (=Peninsula), Aomori Prefec- ture, Honshu Island, Japan; 40°55'-4r25'N, 140°45'-141°30'E; introduced feral popula- tion, multiple escapes from animal park ca. 1979-1991 (Shirai, 1988, p. 3; Mori, 1991, p. 98; Matsubayashi, 1992, p. 2). Not mapped. Shouchia ( ^ ilf|^), Anshuo, Tajen Hsiang; TAITUNG; 22°15'N, 120°50'E; reported in 1955 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386).57i5 Shou Shan (^[Jj) (=Apenberg; Ape's Hill; Chai Shan; Mount Shaushan; Mt. Longevity; Sa- kow; Shao-shan; Takow); KAOHSIUNG; 22°39'N, 120°15'E; geographic account of "Apenberg" (Valentyn, 1726, p. 36). Monkeys reported 12 Jun. 1858 by R. Swinhoe (1859, p. 149; cf Swinhoe in BIyth, 1860, p. 87). Population reported as "almost entirely dis- appeared" by R. Swinhoe (1863, p. 351; cf Collingwood, 1868, p. 39; Namiye 1914, p. 214; Horikawa, 1932, p. 69; Dien, 1958, p. 347). Observed during July 1963 and trapped 62 FIELDIANA: ZOOLOGY for parasitological research 31 Jul. 1963 by J. F. Bergner, Jr. (1967, p. 32); USNM, 1 (skull only). Parasitological samples collected be- fore 1997 by H.-Y. Wu and H. Y. Lin (1996, p. 24). Observed 1995-1997 by M.-J. Hsu and G. Agoramoorthy (1999, p. 64). DNA samples obtained before 1999 by J.-H. Chu, H.-Y. Wu, Y.-J. Yang, O. Takenaka, and Y.-S. Lin (1999, p. 574). Observed 21 Apr. 2000 by H.-Y. Wu and J. Fooden. S106 Shuangtung (^^), ?Tsaotun Chen, 400 m; NANTOU; 23°58'N, 120°46'E; reported 15 Jan. 1993 to tesri survey team. N96 Shuilien Bridge No. 14 {7\(S^\A^M), Shoufeng Hsiang, 100 m; HUALIEN; 23°49'N, 12r33'E; reported 1 Jan. 2000 to H.-Y. Wu. N58 Shuitiliao ( tR ^ ^ ), Fangshan Hsiang; PINGTUNG; 22''23'N, 120°35'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). 5755 Southern Cross-Island Highway {'^a^WM'J^ g#), Haituan Hsiang; TAITUNG; 23°08'N, 12r06'E; 3 monkeys observed 9 Nov. 1998 by S.-W. Chang (TESRi). S66 Southern Cross-Island Highway (l^nl^tMM^ gt), Haituan Hsiang, 390 m; TAITUNG; 23°08'N, 12r07'E; vocalization heard 9 Nov. 1998 by S.-W. Chang (TESRi). S69 Southern Cross-Island Highway (|^p[5|^M^ g§), Haituan Hsiang, 435 m; TAITUNG; 23°09'N, 12r06'E; 1 monkey observed 9 Nov. 1998 by S.-W. Chang (tesri). S70 Southern Cross-Island Highway (I^nPtMM^ jg#), Haituan Hsiang, 515 m; TAITUNG; 23°09'N, 121°05'E; 10 monkeys observed 9 Nov. 1998 by S.-W. Chang (TESRi). S71 Southern Cross-Island Highway (j^nPtSM"^ S#), Haituan Hsiang, 930-945 m; TAITUNG; 23^1 1'N, 121°02'E; 5 monkeys observed at 930 m 8 Nov. 1998 by S.-W. Chang (tesri). Droppings observed at 935-945 m 8 Nov. 1998 by S.-W. Chang (TESRi). Six monkeys observed at 943 m 8 Nov. 1998 by S.-W. Chang (TESRi). S59 Southern Cross-Island Highway (|^p|5|^M^ l^ ), Haituan Hsiang, 1040-1125 m; TAITUNG; 23°11'N, 12 TOTE; droppings observed at 1040 m and 1065 m 8 Nov. 1998 by S.-W. Chang (tesri). Five monkeys ob- served at 1125 m 8 Nov. 1998 by S.-W. Chang (TESRI). S59 Southern Cross-Island Highway (l^pBt^M^ l^), Haituan Hsiang, 1170 m; TAITUNG; 23°I2'N, 121°00'E; 10 monkeys observed 8 Nov. 1998 by S.-W. Chang (tesri). S60 Southern Cross-Island Highway (f^pBt^M"^ £#), Haituan Hsiang, 1665 m; TAITUNG; 23°13'N, 12r00'E; vocalization heard 7 Nov. 1998 by S.-W. Chang (TESRi). S58 Southern Cross-Island Highway ([^p[5tSH:2r f^), Haituan Hsiang, 1855 m; TAITUNG; 23°14'N, 120°59'E; droppings observed 7 Nov. 1998 by S.-W. Chang (tesri). S57 Southern Cross-Island Highway (|^r|^|MM:2^ l^ ), Haituan Hsiang, 2065-2090 m; TAITUNG; 23°I5'N, I20°59'E; vocalization heard at 2065 m 7 Nov. 1998 by S.-W. Chang (TESRi). Five monkeys observed at 2090 m 7 Nov. 1998 by S.-W. Chang (tesri). S42 Southern Cross-Island Highway (MpI^IMM^a l^ ), Taoyuan Hsiang, 975-1050 m; KAOHSIUNG; 23°16'N, 120°51'E; 8 mon- keys observed at 975 m 5 Nov. 1998 by S.-W. Chang (TESRI). Droppings observed at 1050 m 5 Nov. 1998 by S.-W. Chang (tesri). S46 Southern Cross-Island Highway (I^pPtSM^:2^ g# ), Taoyuan Hsiang, 995-1235 m; KAOHSIUNG; 23°16'N, 120°50'E; 4 mon- keys observed at 995 m 5 Nov. 1998 by S.-W. Chang (tesri). Droppings observed at 1235 m 5 Nov. 1998 by S.-W. Chang (tesri). S47 Southern Cross-Island Highway (I^pBt^M^ l^ ), Taoyuan Hsiang, 1250-1445 m; KAOHSIUNG; 23*'17'N, 120°50'E; droppings observed at 1250 m, 1255 m, 1353 m, 1390 m, and 1405-1445 m 5 Nov. 1998 by S.-W. Chang (TESRI). S47 Southern Cross-Island Highway (Ma^^WW^i^ l^ ), Taoyuan Hsiang, 1445-1565 m; KAOHSIUNG; 23°17'N, 120°51'E; droppings observed at 1445-1485 m, 1485-1515 m, 1520-1550 m, and 1555 m 5 Nov. 1998 by S.-W. Chang (TESRi). Vocalization heard at 1565 m 5 Nov. 1998 by S.-W. Chang (TESRi). S46 Southern Cross-Island Highway (l^pf^tl /A PIOa ), Taoyuan Hsiang, 1635-1750 m; FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 63 N147 Taipingshan ( ic zp ^ij ); ILAN; 24''30'N, 12r31'E; observed at 1920 m in Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145). Reported above 1500 m in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386; Tanaka, 1986, p. 101). Ob- served 1987-1989 by L.-L. Lee and Y.-S. Lin (1991, p. 36; 1994, p. 19).A^25 Taipingshan (:::^^lJ4), northwestern foothill area, 400-1350 m; ILAN; ca. 24°33'N, 12r30'E; observed 1987-1989 by S. Kawa- mura, N. Azuma, and K. Norikoshi (1988, p. 1; Kawamura et al., 1991, p. 97; Azuma & Norikoshi, 1990, p. 43; Norikoshi, 1991, p. 136).A^26 Taitocho. See TAITUNG. TAITUNG (=Taitocho) forests (ft^i:t^M^#), mountains; 22°14'-23°27'N, 120°44'- 121°29'E; reported before 1930 by T. Kano (1929, p. 333). Reported before 1959 by Z.-M. Dien (1958, p. 346). Not mapped. Taitung vicinity (^^%M); TAITUNG; ca. 22°45'N, 121°09'E; serum samples obtained 1962-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967, p. 146). SI 18 Taiwan; imprecise locality; collected before 9 Jan. 1958 by unknown collector (R. E. Kuntz, shipping list, 9 Jan. 1958, USNM archives); USNM 308721, 1 (skin only). Not mapped. Tai Yuan. See Tayuan. Takangkou (A^P), Jusui Pass, Fengpin dis- trict; HUALIEN; 23''28'N, 121°29'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 5'2<^ Takao. See Kaohsiung. Takeng (y^i/C), 460-510 m; TAICHUNG; 24°10'N, 120°47'E; groups observed at 460 m and 510 m 23 Dec. 1994 by TESRI survey team. N85 Takow. See Shou Shan. Taku ( ^ ^ ) (=Taririku, Taimari Subdist.); TAITUNG; ca. 22°27'N, 120°50'E; observed before 1930 by T. Kano (1929, p. 333; cf Kuroda, 1938, p. 112; Huang et al., 1993, p. 166). SI 35 Taoshan Pupu. See Wuling. Tapang (^^[5), Alishan Hsiang, 850 m; CHIA-I; ca. 23°27'N, 120°43'E; 1 monkey observed 1 May 1998 by L.-L. Lee and H.-Y. Wu. S5 Tapu Hsiang {-Xi^), 380 m; CHIA-I; ca. 23°18'N, 120°35'E; droppings observed 12 Sep. 1997 by TESR] survey team. S50 Taririku. See Taku. Taroko National Park (^feg-IS S^^Bl); HUALIEN; ca. 24°03-23'N, 12n3-42'E; observed 1987-1989 by L.-L. Lee and Y.-S. Lin (1991, p. 36; 1994, p. 19). Collected ca. 9 Apr. 1997 and 19 Jan. 1998 by unknown col- lector; NMNS, 2 (frozen carcasses). N49 Taroko National Park Headquarters (yfe-^^^S ^^Sl'JiJS), Hsiulin Hsiang, 200 m; HUALIEN; 24°09'N, 121°37'E; reported 1 Oct. 1999toH.-Y. Wu. A^55 Tashiaku {'XMi^), Meishan Hsiang, 510 m; CHIA-I; 23°34'N, 120°38'E; 10 monkeys ob- served 3 Dec. 1997 by tesri survey team. N146 Tataan (?), forestry branch office, Mutan Hsiang; PINGTUNG; ca. 22°07'N, 120°49'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 5/49 Tatachia {i^i^j]\\\ Hsini Hsiang; NANTOU; 23°29'N, 120°53'E; 13 monkeys observed 25 Sep. 1994 by TESRI survey team. Sll Tawan {X'M), Taian Hsiang, 990-1030 m; MIAOLI; ca. 24°26'N, 120°54'E; droppings observed 4 Dec. 1996 by TESRi survey team. N37 Tawu Hsi Ok^^), right bank, Tawu Hsiang; TAITUNG; ca. 22°22'N, 120^54^; reported 1967-1971 by K. Masui, Y. Narita, and S. Tanaka(1986, p. 386).5/i7 Tawu Mountain Nature Preserve {'XMaU^'^. fMiiM); TAITUNG; ca. 22°35'N, 120''46'E; observed 1987-1989 by L.-L. Lee and Y.-S. Lin (1991, p. 36; 1994, p. 19). S129 Tawu Mountain Nature Preserve O^MMj^^^. i^^M), Taimali Hsi, 300^00 m; TAITUNG; 22°35'N, 120°49'E; collected 19 Feb. 2000 by S.-M. Lin; private collection, 1 (skull only). S128 Tayuan (#/J^), Haianshanmo, Tungho Hsiang; TAITUNG; ca. 23°00'N, 121°18'E; reported in 1967-1971 and 1982 by K. Masui, Y. Na- rita, and S. Tanaka (1986, p. 386). S81 Tayuan (MW (= Tai Yuan); TAITUNG; ca. 23°00'N, 12ri8'E; reported Jun.-Dec. 1978 66 FIELDIANA: ZOOLOGY by F. E. Poirier (Poirier & Davidson, 1 979, p. 145). S81 Teftiyeh (#gi?), Alishan Hsiang, 1520 m; CHIA-I; 23''27'N, 120°45'E; observed 6 Apr. 1998 by TESRI survey team. S6 Teraso. See Chulaoshu Shan and Manchoutsun. Tienchung Park (E94^^§), Tienchung Chen, 130 m; CHANGHUA; 23°5rN, \20°41'E; 3 monkeys observed 20 Jan. 1 995 by TESRI sur- vey team. N106 Tienhsiang (^;#), Hsiulin Hsiang; HU ALIEN; 24°11'N, 121°29'E; reported Jul.-Nov. 1985 by S. Tanaka (1986, p. 98). N51 Tienleng (%}v), Tungshih (=Tungshin) Chen; TAICHUNG; 24°10'N, 120°51'E; reported Jul.-Nov. 1985 by S. Tanaka (1986, p. 98). N87 Toliang (#^), Taimali Hsiang; TAITUNG; 22°30'N, 120°57'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S127 Totsu. See Shimokita Hanto. Toupienkeng iW.ftin.i[h^M), Taiping Hsiang, 300 m; TAICHUNG; 24°06'N, 120°46'E; re- ported 14 Dec, 1995 to TESRI survey team. N91 Tsaoling Road {^^^^), Chushan Chen; NANTOU; 23''36'N, 120^40^; 10 monkeys observed 23 Dec. 1992 by tesri survey team. Nil? Tsengwen Shuiku (^Reservoir) ("^^tKI^), Tapu Hsiang, 300 m; CHIA-I; 23°16'N, 120°36'E; 10 monkeys observed 22 Dec. 1996 by TESRI survey team. Ten monkeys observed 2 Nov. 1997 by TESRI survey team. Six mon- keys observed 1 Feb. 1998 by tesri survey team. S49 Tsugitaka Mountains. See Hsueh Shan. Tsuisia Lake. See Jihyueh Tan. Tulanshan ( ^ M [JL[ ), Tungho Hsiang; TAITUNG; 22''54'N, 12ni'E; observed in 1967-1971 and 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S83 Tungfeng (Mi^M), Jenai Hsiang, 905-1010 m; NANTOU; 24°03'N, 121°04'E; droppings observed at 905 m 29 Dec. 1994 by tesri sur- vey team. Droppings observed at 1010 m 30 Dec. 1994 by tesri survey team. N71 Tungho township ( M M ); TAITUNG; ca. 22°58'N, 121''18'E; collected for parasi- tological research before 1966 by C. H. Chuang, J. C. Lien, and S. Y. Lin (Peyton & Harrison, 1980, p. 342). S82 Tunghotsun ( ^ M i^^ ), Nanchuang Hsiang; MIAOLI; 24°36'N, 121°0rE; reported 23 Oct 1996 to TESRI survey team. N33 Tunghou (ffitt), Wulai, 480 m; TAIPEI; 24°50'N, 121°38'E; vocalization heard 1 May 1997 by L.-L. Lee and H.-Y. Wu. N8 Tungli ( ^ M ), Fuli Hsiang; HUALIEN; 23''16'N, 12ri8'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S31 Tungshih. See Maanliao. Tungshing (?), Hungyeh, Yenping Hsiang; TAITUNG; ca. 22°54'N, 12r03'E; observed in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 5<5 7 Tzuchung (§>^x), Alishan Hsiang, 2400 m; CHIA-I; 23°29'N, 120°49'E; 13 monkeys ob- served 1 May 1998 by TESRI survey team. S9 Wakayama Prefecture. See Moushi; Nakatsu. Wakinosawa. See Shimokita Hanto. Wang Lung Shan. See Wanglung. Wanglung (?) (?=Wang Lung Shan); PINGTUNG; ca. 22°30'N, 120°34'E; reported Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 157). 5/57 Wanjung Logging Trail (M^WM), Wanjung Hsiang, 565 m; HUALIEN; 23^43 'N, 12r2rE; 9 monkeys observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N136 Wanjung Logging Trail (M^^M), Wanjung Hsiang, 800 m; HUALIEN; 23°44'N, 12r21'E; 1 monkey observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. Ml 36 Wanjung Logging Trail (H^^xt), Wanjung Hsiang, 820 m; HUALIEN; 23°44'N, 121°20'E; 1 monkey observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N135 Wanjung Logging Trail (M^WM), Wanjung Hsiang, 820 m; HUALIEN; 23°45'N, 121°20'E; 1 monkey observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N133 Wanjung Logging Trail (M^^^M), Wanjung Hsiang, 960 m; HUALIEN; 23°45'N, 121''19'E; 8 monkeys observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N133 FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 67 N147 Taipingshan ( i: 2p Oj ); ILAN; 24°30'N, 121°31'E; observed at 1920 m in Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145). Reported above 1500 m in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386; Tanaka, 1986, p. 101). Ob- served 1987-1989 by L.-L. Lee and Y.-S. Lin (1991, p. 36; 1994, p. \9).N25 Taipingshan (:;^2p^J)^ northwestern foothill area, 400-1350 m; ILAN; ca. 24°33'N, 12r30'E; observed 1987-1989 by S. Kawa- mura, N. Azuma, and K. Norikoshi (1988, p. 1; Kawamura et al., 1991, p. 97; Azuma & Norikoshi, 1990, p. 43; Norikoshi, 1991, p. U6).N26 Taitocho. See TAITUNG. TAITUNG (=Taitocho) forests (ft^iiblE^#), mountains; 22°14'-23°27'N, 120°44'- 12r29'E; reported before 1930 by T. Kano (1929, p. 333). Reported before 1959 by Z.-M. Dien (1958, p. 346). Not mapped. Taitung vicinity (^^ii^lS); TAITUNG; ca. 22°45'N, 121°09'E; serum samples obtained 1962-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967, p. 146). SI 18 Taiwan; imprecise locality; collected before 9 Jan. 1958 by unknown collector (R. E. Kuntz, shipping list, 9 Jan. 1958, USNM archives); USNM 308721, 1 (skin only). Not mapped. Tai Yuan. See Tayuan. Takangkou (A"i#P), Jusui Pass, Fengpin dis- trict; HU ALIEN; 23°28'N, 12r29'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 525 Takao. See Kaohsiung. Takeng ( A i/L ), 460-510 m; TAICHUNG; 24°10'N, 120''47'E; groups observed at 460 m and 5 10 m 23 Dec. 1994 by tesri survey team. N85 Takow. See Shou Shan. Taku ( ;^ ^ ) (=Taririku, Taimari Subdist.); TAITUNG; ca. 22°27'N, 120°50'E: observed before 1930 by T. Kano (1929, p. 333; cf Kuroda, 1938, p. 112; Huang et al., 1993, p. \66).S135 Taoshan Pupu. See Wuling. Tapang (^^ji), Alishan Hsiang, 850 m; CHIA-I; ca. 23''27'N, 120°43'E; 1 monkey observed 1 May 1998 by L.-L. Lee and H.-Y. Wu. S5 Tapu Hsiang (;^ii), 380 m; CHIA-I; ca. 23°18'N, 120°35'E; droppings observed 12 Sep. 1997 by tesri survey team. S50 Taririku. See Taku. Taroko National Park (:^''i^K S^^P); HUALIEN; ca. 24°03-23'N. 12ri3^2'E; observed 1987-1989 by L.-L. Lee and Y.-S. Lin (1991, p. 36; 1994, p. 19). Collected ca. 9 Apr. 1997 and 19 Jan. 1998 by unknown col- lector; NMNS, 2 (frozen carcasses). N49 Taroko National Park Headquarters (:;^'§-^S ^^^^S^), Hsiulin Hsiang, 200 m; HUALIEN; 24°09'N, 121°37'E; reported 1 Oct. 1999 to H.-Y. Wu.A^55 Tashiaku (Afl^^), Meishan Hsiang, 510 m; CHIA-1; 23°34'N, 120°38'E; 10 monkeys ob- served 3 Dec. 1997 by TESRI survey team. N146 Tataan (?), forestry branch office, Mutan Hsiang; PINGTUNG; ca. 22°07'N, 120°49'E; reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 5/^P Tatachia {i^i^M), Hsini Hsiang; NANTOU; 23°29'N, 120°53'E; 13 monkeys observed 25 Sep. 1994 by tesri survey team. Sll Tawan (Ai^), Taian Hsiang, 990-1030 m; MIAOLI; ca. 24°26'N, 120°54'E; droppings observed 4 Dec. 1996 by tesri survey team. N37 Tawu Hsi {i\^Wi), right bank, Tawu Hsiang; TAITUNG; ca. 22°22'N, 120°54'E; reported 1967-1971 by K. Masui, Y. Narita, and S. Tanaka(1986, p. 386). 5/57 Tawu Mountain Nature Preserve {i\.^\\-\^^. ^HM); TAITUNG; ca. 22°35'N, 120°46'E; observed 1987-1989 by L.-L. Lee and Y.-S. Lin (1991, p. 36; 1994, p. 19). SI 29 Tawu Mountain Nature Preserve (AiKlijS^ ^^1^), Taimali Hsi, 300^00 m; TAITUNG; 22°35'N, 120°49'E; collected 19 Feb. 2000 by S.-M. Lin; private collection, 1 (skull only). S128 Tayuan {^W), Haianshanmo, Tungho Hsiang; TAITUNG; ca. 23°00'N, 12ri8'E; reported in 1967-1971 and 1982 by K. Masui, Y. Na- rita. and S. Tanaka (1986, p. 386). S81 Tayuan {WM) (= Tai Yuan); TAITUNG; ca. 23°00'N, 12n8'E; reported Jun.-Dec. 1978 66 FIELD! AN A: ZOOLOGY by F. E. Poirier (Poirier & Davidson, 1979, p. 145). S8] Teftiyeh (#si?), Alishan Hsiang, 1520 m; CHIA-I; 23°27'N, 120°45'E; observed 6 Apr. 1998 by TESR] survey team. S6 Teraso. See Chulaoshu Shan and Manchoutsun. Tienchung Park (EH 4^:2^3), Tienchung Chen, 130 m; CHANGHUA; 23°51'N, OOMT'E; 3 monkeys observed 20 Jan. 1995 by tesri sur- vey team. N106 Tienhsiang (^|^), Hsiulin Hsiang; HUALIEN; 24°11'N, 121°29'E; reported Jul.-Nov. 1985 by S. Tanaka (1986, p. 98). N51 Tienleng (^i^), Tungshih (=Tungshin) Chen; TAICHUNG; 24''10'N, 120°51'E; reported Jul.-Nov. 1985 by S. Tanaka (1986, p. 98). N87 Toliang (#^), Taimali Hsiang; TAITUNG; 22°30'N, 120°57'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). SI 27 Totsu. See Shimokita Hanto. Toupienkeng (0^fK^{Lll"f'/l5]), Taiping Hsiang, 300 m; TAICHUNG; 24°06'N, 120°46'E; re- ported 14 Dec. 1995 to TESRi survey team. N9I Tsaoling Road ( ^ ^ S# ), Chushan Chen; NANTOU; 23°36'N, 120°40'E; 10 monkeys observed 23 Dec. 1992 by TESR] survey team. Nil? Tsengwen Shuiku (=Reservoir) ('^^tR)^), Tapu Hsiang, 300 m; CHIA-I; 23°16'N, 120°36'E; 10 monkeys observed 22 Dec. 1996 by TESRI survey team. Ten monkeys observed 2 Nov. 1997 by TESRI survey team. Six mon- keys observed 1 Feb. 1998 by tesri survey team. S49 Tsugitaka Mountains. See Hsueh Shan. Tsuisia Lake. See Jihyueh Tan. Tulanshan ( W> M lU ), Tungho Hsiang; TAITUNG; 22°54'N, 12ril'E; observed in 1967-1971 and 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S83 Tungfeng (M^'M), Jenai Hsiang, 905-1010 m; NANTOU; 24°03'N, 12r04'E; droppings observed at 905 m 29 Dec. 1994 by tesri sur- vey team. Droppings observed at 1010 m 30 Dec. 1994 by tesri survey team. N71 Tungho township ( ^ M ); TAITUNG; ca. 22°58'N, 12n8'E; collected for parasi- tological research before 1966 by C. H. Chuang, J. C. Lien, and S. Y. Lin (Peyton & Harrison, 1 980, p. 342). 552 Tunghotsun ( M M 14 ), Nanchuang Hsiang; MIAOLI; 24°36'N, 121°01'E; reported 23 Oct 1996 to TESRI survey team. N33 Tunghou (ffift), Wulai, 480 m; TAIPEI; 24''50'N, 121°38'E; vocalization heard 1 May 1997 by L.-L. Lee and H.-Y. Wu. N8 Tungli ( ^ M ), Fuli Hsiang; HUALIEN; 23°16'N, 121°18'E; reported in 1982 by K. Masui, Y. Narita, and S. Tanaka (1986, p. 386). S3] Tungshih. See Maanliao. Tungshing (?), Hungyeh, Yenping Hsiang; TAITUNG; ca. 22°54'N, 121°03'E; observed in 1982 by K. Masui, Y. Narita, and S. Ta- naka (1986, p. 386). 5<^7 Tzuchung (g)^^), Alishan Hsiang, 2400 m; CHIA-I; 23°29'N, 120°49'E; 13 monkeys ob- served 1 May 1998 by tesri survey team. S9 Wakayama Prefecture. See Moushi; Nakatsu. Wakinosawa. See Shimokita Hanto. Wang Lung Shan. See Wanglung. Wanglung (?) (?=Wang Lung Shan); PINGTUNG; ca. 22°30'N, 120°34'E; reported Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 151). SI 31 Wanjung Logging Trail (M^WM), Wanjung Hsiang, 565 m; HUALIEN; 23°43'N, 12r21'E; 9 monkeys observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. NI36 Wanjung Logging Trail (M^^^M), Wanjung Hsiang, 800 m; HUALIEN; 23°44'N, 121°2rE; 1 monkey observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N136 Wanjung Logging Trail (M^^M), Wanjung Hsiang, 820 m; HUALIEN; 23°44'N, 121°20'E; 1 monkey observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. NI35 Wanjung Logging Trail (il|^#jl), Wanjung Hsiang, 820 m; HUALIEN; 23''45'N, 121°20'E; 1 monkey observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N133 Wanjung Logging Trail (||^#jI), Wanjung Hsiang, 960 m; HUALIEN; 23°45'N, 121°19'E; 8 monkeys observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N133 FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 67 Wanjung Logging Trail (^^^M), Wanjung Hsiang, 1085-1165 m; HUALIEN; 23''44'N, 121°19'E; monkeys observed at 1085 m and 1165 m 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N135 Wanjung Logging Trail (H^^iS), Wanjung Hsiang, 1410-1600 m; HUALIEN; 23''44'N, 121°18'E; monkeys observed at 1410 m and 1600 m 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N134 Wanjung Logging Trail (^^#jM), Wanjung Hsiang, 1700 m; HUALIEN; 23°45'N, 121°16'E; 3 monkeys observed 1 Feb. 1997 by L.-L. Lee and H.-Y. Wu. N1S2 Wanta Logging Trail (||;*C#iM), Jenai Hsiang, 1590 m; NANTOU; 23''56'N, 12r06'E; vo- calization heard 28 Nov. 1996 by S.-W. Chang (TESRI). N69 Western Taiwan (^mM)\ imprecise locality; reported in 1715 by Pere J.-A.-M. de Moyria de Mailla (Du Halde, 1735, pp. xxxiv, 162; Weiss, 1870-1873, p, 120; Campbell, 1903, pp. 504, 506; Davidson, 1903, p. 68). Not mapped. Wulai (B5}5); TAIPEI; 24^52'N, 12r33'E; col- lected before 1959 by unknowTi collector (Dien, 1958, p. 345); museum unknown. Col- lected for parasitological research before 1 965 probably by R. E. Kuntz (Schad & Anderson, 1963, p. 799); museum unknown. Observed at 1000 m in Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145). Reported in 1982 by K. Masui, Y. Narita, and S. Ta- naka(1986,p. 386).7V6 Wulai area (.^J^it^M); TAIPEI; ca. 24°52'N, 121°33'E; serum samples obtained 1959-1966 by C.-S. Yang, C.-H. Kuo, and C.-Y. Chen (1967, p. 146). Trapped for parasitological research before 1965 (Kim &. Bergner. 1964. p. 81).7V6 Wulaokenghsi Mining Trail (:K^JKt^^^jS g§), Nanao Hsiang, 224-625 m; ILAN; 24°33'N, 12r47'E; vocalization heard at 224 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. Three monkeys observed at 610 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. Droppings observed at 625 m 1 Dec. 1996 by L.-L. Lee and H.-Y. Wu. N20 Wuling Farm (^|^0i#), Hoping Hsiang, 1987-2010 m; TAICHUNG; 24°15'N, 121°0rE; droppings observed at 1987 m and 2010 m 11 Apr. 1996 by TESRI survey team. N76 Wuling (:^|^) (Taoshan Pupu), Hoping Hsiang, 2140 m; TAICHUNG; 24°26'N, 12n7'E; 1 monkey observed 1 6 Nov. 1 995 by TESRI sur- vey team. N45 Wulu Logging Trail ( S ^ # iM ), Haituan Hsiang, 860 m; TAITUNG; 23°09'N, 121°03'E; 2 monkeys observed 21 Nov. 1997 by S.-W. Chang (TESRl). S65 Wulu Logging Trail ( ^ ^ 1^ xi ). Haituan Hsiang, 1925-2005 m; TAITUNG; 23''08'N, I2I°00'E; droppings observed at 1925 m and 1930 m 19 Nov. 1997 by S.-W. Chang (TESRi). Vocalization heard at 1930 m 19 Nov. 1997 by S.-W. Chang (TESRi). Three monkeys ob- served at 2005 m 20 Nov. 1997 by S.-W. Chang (TESRJ). S63 Wulu Logging Trail ( ^ i^ ^ xi ), Haituan Hsiang, 2060-2215 m: TAITUNG; 23°08'N. 120°59'E; groups observed at 2060 m and 2215 m 20 Nov. 1997 by S.-W. Chang (tesri). Dead monkey observed at 2145 m 20 Nov. 1997 by S.-W. Chang (tesri). Vocalization heard at 2165 m 20 Nov. 1997 by S.-W. Chang (tesri). S62 Wulu Logging Trail (^i^^jl), Haituan Hsiang, 2240 m; TAITUNG; 23°08'N, 120°58'E; vocalization heard 20 Nov. 1997 by S.-W. Chang (tesri). S61 Wushan ( .^ [J_| ), Nanhua Hsiang, 500 m; TAINAN; 23°00'N, 120°29'E; 2 monkeys ob- served 1 Jan. 1997 by L.-L. Lee and H.-Y. Wu. SI 03 Wushan ( ,^ !_Ll ). Nanhua Hsiang. 500 m: TAFNAN; 23°0rN, 120^^29^; 50 monkeys observed 1 Jan. 1997 by L.-L. Lee and H.-Y. Wu. SI 02 Wushan ( ,% lJ_| ), Nanhua Hsiang, 500 m; TAINAN; 23^0 1'N, 120°30'E; droppings ob- served 1 Jan. 1997 by L.-L. Lee and H.-Y. Wu. SlOl Wushihkeng { % ^ iK X ca. 1200 m; TAICHUNG; 24°18'N. 120°55'E; collected before 1999 by K.-L. Huang and S.-W. Chang; TESRI, 1 (skeleton only). N81 Wushihkeng ( ,% H ^ ), Hoping Hsiang; 68 RELDIANA: ZOOLOGY TAICHUNG; 24°16'N, 120°56'E; reported 30 Dec. 1992 to tesrj survey team. Droppings observed 19 Feb., 7 Jul., 8 Jul., and 17 Sep. 1993 by TESRJ survey team. One monkey ob- served 9 Jul. 1993 by tesri survey team. N82 Wushihkeng ( .^ S Itl ), Hoping Hsiang, 600-1250 m; TAICHUNG; 24°16'N, 120°56'E; 14 monkeys observed at 700 m 15 Oct. 1992 by TESRJ survey team (same team also is source of following observations at this locality). Capture reported at 1000 m 15 Dec. 1992. Droppings observed at 600 m 19 Jan. 1993. Droppings observed at 700 m 26 Mar. 1993. Six monkeys observed at 920 m 7 Jul. 1993. Three monkeys observed at 810 m 4 Aug. 1993. Vocalization heard at 1010 m 17 Sep. 1993. Vocalization heard at 1080 m 19 Oct. 1993. One monkey observed at 800 m 20 Oct. 1993. Food remains observed at 860 m 20 Oct. 1993. Droppings observed at 870 m 20 Oct. 1993. Vocalization heard at 1100 m 21 Oct. 1993. Droppings observed at 700 m 18 Nov. 1993. Fifteen monkeys observed at 930 m 18 Nov. 1993. Droppings observed at 680 m 14 Dec. 1993. Food remains observed at 1200 m 14 Dec. 1993. Droppings observed at 1250 m 14 Dec. 1993. Droppings observed at 820 m and 1245 m 15 Dec. 1993. Ten monkeys observed at 1080 m 16 Feb. 1994. Dead monkey observed at 1070 m 17 Feb. 1994. Vocalization heard at 1230 m 19 Apr. 1994. Thirteen monkeys observed at 1250 m 20 Apr. 1994. Two monkeys observed at 1090 m 18 May 1994. Food remains observed at 1120 m 18 May 1994. Droppings observed at 1080 m 19 May 1994. Droppings observed at 1150 m 27 Oct. 1995. Vocalization heard at 1200 m 27 Oct. 1995. Eight monkeys ob- served at 1000 m 15 Nov. 1995. Droppings observed at 920 m 10 Jan. 1996. N82 Wushihkeng ( ,% H ^ ), Hoping Hsiang, 635-1305 m; TAICHUNG; 24''16'N, 120°56'E; observed 26 Feb. 1997 by S.-W. Chang (TESRI). N82 Wushihkeng i%^iK) vicinity; TAICHUNG; ca. 24°18'N, 120°55'E; albino juvenile captured in early 1990s by local residents (see above. Albinism). N81 Wutao Shan (^Slllj), Ali village, 6 km along logging trail; PINGTUNG; 22°43'N, 120°47'E; collected 23 Jan. 1995 by S.-Y. Huang; TESRi, 1 (skull only). 5770 Yen Hai Logging Trail {WM'W^M.), Hsiulin Hsiang; HUALIEN; ca. 24°09'N, 12r30'E; collected 28 Oct. 1994 by Y.-C. Sun; TESRI, 1 (skull only). N52 Yeng Ping Working Circle. See Yenping Hsiang. Yenping Hsiang (Mr^M) (=Yeng Ping Work- ing Circle), 1500 m; TAITUNG; ca. 22°54'N, 120°59'E; observed Jun.-Dec. 1978 by F. E. Poirier (Poirier & Davidson, 1979, p. 145; Masui et al., 1986, p. 386). S93 Yenping Logging Trail (Jl^^xi), Yenping Hsiang; TAITUNG; 22°54'N, 121''02'E; vo- calization heard and droppings observed 5 Dec. 1996 by S.-W. Chang (tesri). S86 Yenping Logging Trail (MJ^^'M.), Yenping Hsiang, 1150 m; TAITUNG; 22°53'N, 12r02'E; 2 monkeys observed 1 May 1998 by S.-W. Chang (TESRi). S85 Yenping Logging Trail (^^#jS), Yenping Hsiang, 1215-1360 m; TAITUNG; 22°54'N, 121°02'E; droppings observed at 1360 m 30 Apr. 1998 by S.-W. Chang (tesri). Droppings observed at 1215 m 1 May 1998 by S.-W. Chang (TESRI). S86 Yenping Logging Trail (^^#jM), Yenping Hsiang, 1330-1525 m; TAITUNG; 22°55'N, 121°02'E; droppings observed at 1330 m, 1385 m, and 1430 m 30 Apr. 1998 by S.-W. Chang (tesri). Vocalization heard at 1435 m 30 Apr. 1998 by S.-W. Chang (TESRi). Two monkeys observed at 1525 m 30 Apr. 1998 by S.-W. Chang (TESRi). S88 Yenping Logging Trail (^^#3!), Yenping Hsiang, 1555-1650 m; TAITUNG; 22°56'N, 121°02'E; 5 monkeys observed at 1645 m 5 Dec. 1996 by S.-W. Chang (TESRi). Droppings observed at 1555 m, 1635 m, and 1650 m 30 Apr. 1998 by S.-W. Chang (TESRi). Vocaliza- tion heard at 1635 m 30 Apr. 1998 by S.-W. Chang (TESRI). S88 Yenping Logging Trail (M/^WM.), Yenping Hsiang, 1600 m; TAITUNG; 22°56'N, 121°01'E; droppings observed 30 Apr. 1998 by S.-W. Chang (tesri). S88 Yenping Logging Trail (M^^jI), Yenping FOODEN AND WU: SYSTEMATIC REVIEW OF MACACA CYCLOPIS 69 Fielaian hiiippiiie Island*! r P- ; 44 pages. 60 'u -I '- i^oira^o^Srouyan Isianci, Ronibion i*rovincc, f'hjiipp.i Distnbution and Biogeographic Affinities. By Stephen M is.<\\/^ ,L /<■<:!. n'V. II. s 1)1!^ o[ Vietnam; A \iv ')1. 1^)99. 46 pay.-,, ' i Oi! ^t•\^ t A ! i'.Hitl and (lUinal In\c!i*u.>ry oi ihc Faic .Naaouui !.k-\a!!onai Xariaiion. i-.ditcd by Stephen M. Goodm ini^/cs. wiih U!ii>. anc! falilc-. To order l-'icliliuiia. please ! et.. 1 i -•; ..:,a:- Fortsas Books. Ltd. 5415 North I (iveuv- UNIVERSITY OF ILUN0I9-URBANA 3 0112 040058098 Field Museum of Natural Histor 1400 South Lake Shore Drive Chicago, Illinois 60605-2496 Telephone: (312) 665-7055