UNIVERSITY OF

ILLINOIS LIBRARY

AT URBANA-CHAMPAIGN

BIOLOGY

MAR 2 6 1985

<u^i_ V

r.cLDIANA Zoology

Published by Field Museum of Natural History New Series, No. 10

TAXONOMY AND EVOLUTION OF THE SINICA GROUP OF MACAQUES: 3. SPECIES AND SUBSPECIES ACCOUNTS OF MACACA ASSAMENSIS

JACK FOODEN

JUL 2 6 1982

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January 29, 1982 Publication 1329

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TAXONOMY AND EVOLUTION

OF THE SINICA GROUP OF MACAQUES:

3. SPECIES AND SUBSPECIES ACCOUNTS

OF MACACA ASSAMENSIS

FIELDIANA Zoology

Published by Field Museum of Natural History

New Series, No. 10

TAXONOMY AND EVOLUTION OF THE SINICA GROUP OF MACAQUES: 3. SPECIES AND SUBSPECIES ACCOUNTS OF MACACA ASSAMENSIS

JACK FOODEN

Research Associate

Field Museum of Natural History

Professor of Zoology Chicago State University

Accepted for publication June 24, 1981 January 29, 1982 Publication 1329

Library of Congress Catalog Card No.: 81-69575

ISSN 0015-0754

PRINTED IN THE UNITED STATES OF AMERICA

CONTENTS

List of Illustrations vi

List of Tables vii

Introduction 1

Species and Subspecies Accounts 2

Macaca assamensis (McClelland in Horsfield, 1840) 2

Distribution 2

Pelage 6

External measurements and proportions 8

Cranial characters 12

Natural history 16

Ecogeographic relations with other non-human primates 21

Relationships with non-macaque primate genera 21

Relationship with Macaca arctoides 21

Relationship with Macaca nemestrina 23

Relationship with Macaca mulatto 23

Macaca assamensis pelops 31

Synonymy 31

Type 32

Type-locality 32

Distribution 32

Diagnostic tail measurements and proportions 33

Specimens examined 33

Macaca assamensis assamensis 33

Synonymy 33

Type 34

Type-locality 34

Distribution 34

Diagnostic tail measurements and proportions 35

Specimens examined 35

Gazetteer of Macaca assamensis Localities 35

Macaca assamensis pelops 35

Macaca assamensis assamensis 38

Literature Cited 44

Appendix: Macaca mulatto Localities 51

LIST OF ILLUSTRATIONS

1. Distribution of subtropical mountain evergreen forest in southern Asia and lo- cality records of Macaca assamensis pelops 4

2. Locality records of Macaca assamensis assamensis 5

3. External characters in Macaca assamensis adult males 7

4. Tail length vs. head and body length in immature and adult specimens of Macaca assamensis pelops and M. a. assamensis 9

5. Cranial characters in Macaca assamensis 13

6. Allometry of rostral length vs. postrostral length in immature and adult specimens

of Macaca assamensis 15

7. Geographic relationship between known localities of Macaca assamensis assamensis and nearby known localities of M. nemestrina leonina 24

8. Limits of distribution of Macaca assamensis and M. mulatta and geographic rela- tionship between known localities of M. a. pelops and nearby known localities of

M. mulatta 26

9. Geographic relationship between known localities of Macaca assamensis assamensis and nearby known localities of M. mulatta 27

10. External characters of Macaca assamensis pelops compared with those of M. mulatta in hitherto unpublished drawing commissioned in Nepal by B. H. Hodgson and sent to British Museum in 1845 31

LIST OF TABLES

1. Crown hair arrangement in Macaca assamensis specimens 8

2. Latitudinal variation of external measurements and external proportions in adult Macaca assamensis 10

3. Cranial dimensions and ratios in various age classes of Macaca assamensis 14

4. Latitudinal variation of greatest skull length in adult Macaca assamensis 14

5. Altitudinal distribution of Macaca assamensis localities 16

6. Individuals per troop reported in Macaca assamensis 18

7. Age-sex composition in three troops of Macaca assamensis 18

8. Evidence of probable reproductive seasonality in Macaca assamensis pregnant fe- males and infants 20

9. Genera and species of other primates reported in vicinity of Macaca assamensis 22

10. Collection details and field observations concerning 11 local areas inhabited by Macaca assamensis assamensis, but apparently not inhabited by Macaca mulatto . . 28

11. Collection details and field observations concerning 10 localities where both Ma- caca assamensis assamensis and Macaca mulatto have been collected or observed . 29

vii

INTRODUCTION

Macaca assamensis is a short-tailed monkey that inhabits the foothills of the Himalayas and adjoining mountain chains in Southeast Asia. This species — together with M. sinica, M. radiata, and M. thibetana — is assigned to the sinica group of macaques, a species group defined by shared derived characters of male and female reproductive anatomy (Fooden, 1980, p. 1). Although M. arc- toides and M. mulatto also have been postulated as close relatives of M. assamensis (Delson, 1980, p. 22; Cronin et al., 1980, p. 46), neither of these two species resembles recognized members of the sinica group in critical characters of male or female reproductive anatomy. Species accounts of M. sinica and M. radiata are published (Fooden, 1979, p. 109; 1981, p. 2); a species account of M. thibetana and a comprehensive comparative overview of the sinica group are in prepa- ration.

In the present account of Macaca assamensis, species-level discussions of dis- tribution, morphology, and natural history are presented first. These are fol- lowed by taxonomic reviews of the two subspecies now recognized, sub-Hi- malayan M. a. pelops and Southeast Asian M. a. assamensis. The basis for recognition of these subspecies is a clearly defined difference in relative tail length in adult males; the longer tail in M. a. pelops presumably is more primitive than the shorter tail in M. a. assamensis.

This account is based on study of 112 museum specimens, review of relevant literature, and field study of M. a. assamensis in western Thailand. Specimens examined are preserved in the following institutions, which hereafter are cited by means of the indicated abbreviations:

AMNH American Museum of Natural History, New York

BM British Museum (Natural History), London

BNHS Bombay Natural History Society, Bombay

CTNRC Centre for Thai National Reference Collections, Applied Sci-

entific Research Corporation of Thailand, Bangkok FMNH Field Museum of Natural History, Chicago

MCZ Museum of Comparative Zoology, Harvard University, Cam-

bridge, Mass. MNHN Museum National d'Histoire Naturelle, Paris

USNM U.S. National Museum of Natural History, Washington, D.C.

ZMB Zoologisches Museum des Humboldt-Universitat, Berlin

ZSBS Zoologisches Sammlung des Bayerischen Staates, Munich

ZSI Zoological Survey of India, National Zoological Collection,

Calcutta

I thank officials of these institutions for permission to study collections in their custody and for their generous assistance and courtesy.

2 FIELDIANA: ZOOLOGY

SPECIES AND SUBSPECIES ACCOUNTS

Macaca assamensis McClelland in Horsfield [1840]. Synonymies under subspe- cies headings.

Distribution (Figs. 1, 2)

Lower and middle ranges (ca. 150-2,750 m) of Himalayas and adjoining moun- tain chains in Southeast Asia, from central Nepal eastward through Bhutan, northeastern India (Sikkim, northern West Bengal, Nagaland, probably Arun- achal Pradesh), northern and eastern Burma, southern China (Xizang, Yunnan, Guangxi), northern and western Thailand, Laos, and northern Vietnam, and one widely disjunct record (see below) near sea level in the Sundarbans coastal swamps, southwestern Bangladesh. The main range is broken by an apparent gap of about 400 km between the easternmost known locality of M. a. pelops in Bhutan and the westernmost known locality of M. a. assamensis in Nagaland; however, this intervening area has not been adequately surveyed and the hiatus is probably an artifact. Somewhat incongruously, the range of M. assamensis apparently does not extend into Assam as this Indian state is now delimited; when the species was named, "Assam" denoted practically all of northeastern India.

The isolated Sundarbans record of M. assamensis is peculiar but apparently valid. This record is based on one or two specimens collected in 1870 for J. Anderson, then Curator of the Indian Museum; one of these specimens (ZSI 11999, skin only) is extant. A few details concerning collection of M. assamensis in the Sundarbans are included in Anderson's (1872, p. 529) report on monkeys of this region:

The natives of the [Sundarbans] assert that two Monkeys occur in it, viz. the red- faced Inuus rhesus [ = M. mulatta] and another Monkey [M. assamensis], which they assert has no red about the face or on the hinder quarters. Acting on this information, I sent a collector to procure for me specimens of the two forms; and he returned with a number of undoubted examples of /. rhesus [six specimens now in ZSI], and with two fresh skins [M. assamensis, one now in ZSI] which appear to me to be very different from any adult of J. rhesus that I have examined. The specimens in question were shot about 50 miles east of Calcutta; but as they only reached me as skins I can only give the measurements of these, and of the bones of the limbs and the characters of the skull. . . .

Places outside of the distribution specified above frequently are included in the range of M. assamensis, usually as a result of misidentification of specimens of M. mulatta, which is superficially similar to M. assamensis. Eight of these questionable locality indications are briefly discussed below.

1. Kashmir area, India and Pakistan (ca. 34°-35°N, 73°-75°E). Blanford (1898, p. 361) misinterpreted True's (1894, p. 2) characterization of five M. mulatta specimens (USNM 20120-24) collected at Lolab (Jammu and Kashmir, India) and incorrectly identified these specimens as M. assamensis. Based on this error, Blanford also tentatively identified as M. assamensis macaques that inhabit the vicinity of Murree and Abbottabad in adjacent Pakistan; these also are M. mulatta (cf. BM 23.11.4.1 6 and BNHS 51139, Patriata, Murree).

2. Mussoorie vicinity, Uttar Pradesh, India, northwest of Nepal (30°30'N, 78°00'E). The original and sole basis for current routine inclusion of this place in the distribution of M. assamensis is Hutton's (1865, p. xiii) assertion that the range of Inuus pelops (=M. assamensis pelops) is "Nipal, Kumaon, Mussooree and

FOODEN: MAC AC A ASSAMENSIS 3

Simla." Hutton, who was a British army officer stationed at Mussoorie, observed macaques nearby and first identified them as M. mulatto (1837, p. 935). Later, purely on a priori grounds ("on the score of climate"), Hutton (1868, pp. 945-51) decided that these macaques — in the cool, hilly Kumaon-Mussoorie-Simla re- gion— should be considered specifically distinct from M. mulatto of the warm lowlands. Accordingly, he changed his identification of the upland macaques to Inuus pelops, a nominal species described by Hodgson (1841, p. 1213) in upland east-central Nepal, 750 km east of Mussoorie. Kumaon and Simla, however, are known to be inhabited by M. mulatta and evidently are not inhabited by M. assamensis (Dodsworth, 1914, p. 730; Wroughton, 1914, p. 285; Hingston, 1920, p. 244; Southwick et al., 1961a, p. 540); presumably this also is true of Mussoorie, since Hutton indicates that the same macaque species is common to all three places (see quotation above). Hutton collected, but did not preserve, two living young macaques in the Mussoorie area (Blyth, 1865, p. 192); unfortunately, the only character given for these specimens is "wavy fur" (Jerdon, 1867, p. 12), which does not distinguish M. assamensis from long-haired upland M. mulatta. No other observer or collector has recorded M. assamensis in the Kumaon-Mussoorie-Simla region; the macaques observed and collected there by Hutton almost certainly were M. mulatta, as he originally identified them.

3. Garo Hills, Meghalaya, eastern India (25°30'N, 90°30'E). In an influential paper, Hinton & Wroughton (1921, p. 667) remarked that they were prepared "to accept, with confidence, the specimens collected by Mr. J. P. Mills in the Garo Hills as being practically topotypes of assamensis." Based on this comment, Allen (1938, p. 283) and Carter (1943, p. 106) designated the Garo Hills as probable or definite type-locality for this species. There is no evidence, however, that J. P. Mills or anyone else ever collected specimens of M. assamensis in the Garo Hills (cf. Spence, 1920, map between pp. 372-73). Hinton & Wroughton's reference to this place apparently is a lapsus for Naga Hills, 400 km east of Garo Hills, where Mills (1922, p. v; 1923, p. 221) was stationed as an assistant sub- division officer and collected M. assamensis in 1919 (BM 21.8.2.3-4; ZSI 12093), 1920 (BNHS 5116), and 1923 (BNHS 5115).

4. Imphal, about 4 mi north (24°25'N, 93°57'E), Manipur, eastern India. A juvenile male rhesus macaque (ZSI 11187: M. mulatta; lower back brighter than upper back, relative tail length 0.49) collected at this place by M. L. Roonwal (1949, p. 83; 1950, p. 16) in November 1945 was misidentified as M. assamensis (cf. Mukherjee, 1978a, p. 279).

5. Neghereting (26°45'N, 94°00'E) and Dibru Reserved Forest (27°40'N, 95°15'E), left (south) bank of Brahmaputra River, Assam, northeastern India. Pilleri (1975, pp. 20, 43) reported that he observed troops of M. assamensis at these places during the course of an expedition to study river dolphins [Plan- tanista gangetica) in the Brahmaputra River. Color slides of these monkeys kindly sent to me by Professor Pilleri (letter, 15 December 1978) permit the species to be identified as M. mulatta.

6. Myitkina ( = Myitkyina; 25°23'N, 97°24'E), northern Burma. A living sub- adult female macaque purchased at this place, probably in 1945, was examined by M. L. Roonwal (1950, p. 16) and identified as M. assamensis. Judging from pelage color characters given ("hind-parts deep rusty"), the monkey probably was a specimen of M. mulatta (cf. BM 37.12.3.75 9, 37.12.3.77 2, M. mulatta, collected at Pidaung Reserve, Myitkyina District, by R. Kaulback, 25 May 1936).

FIELDIANA: ZOOLOGY

' r — =p

+ + +

M. a. pe/ops

■ Specimens examined

a Literature records

BANGLADESH

Fig. 1. A, Distribution of subtropical mountain evergreen forest in southern Asia (So- chava and Lukicheva, 1964, p. Ill, vegetation type 114). B, Locality records of Macaca assamensis pelops; inset (upper right) depicts position of localities in Sikkim-northern West Bengal area more accurately than possible in main map. Key to localities (for details, see Gazetteer): 1. Tipling. 2. Jiri. 3. Iswa Khola. 4. Sabaya Khola. 5. Lachen. 6. Manshitang. 7. Chuntang. 8. Dikchu. 9. Singtam. 10. Rhenok. 11. Rongli. 12. Palmajua. 13. Batasia. 14. Gopaldhara. 15. Kurseong. 16. Sookia Pokhari. 17. Birch Hill; Darjeeling, near. 18. Takdah. 19. Pashok. 20. Tarkhola. 21. Singsir. 22. Daling. 23. Shamgong. 24. Manas River. 25. Sundarbans.

7. Nine Burmese localities reported for M. assamensis by Wroughton, 1915-21, as follows: Mingun, Mt. Popa and Yenangyaung (1915, p. 464); Yin, Tatkon and Homalin (1916a, p. 296); "Atechg." (1916c, p. 763); Shan States (1918, p. 555); and Theme (1921, p. 553). All of these records are based on misidentified M. mulatta, as subsequently recognized by Wroughton (in Hinton & Wroughton, 1921, p. 666). Wroughton (1916a, p. 296) also identified as M. assamensis two macaques collected at Hisweht, Burma; of these, one is M. mulatta (BM 31.1.11.24 9) and the other is M. assamensis (BM 1937.3.24.13 8, skin/BNHS 5118, skull).

8. Vinh Linh region (17°N, 107°E), east central Vietnam. A juvenile male macaque collected in this region in August 1956 was identified as M. assamensis by Dao Van Tien (1960, p. 228; 1962, p. 724). Characters given for the specimen (croup reddish, relative tail length 0.52) indicate that it probably is M. mulatta.

Evidence concerning occurrence of M. assamensis in the Khasi Hills (ca. 25°30'N, 91°30'E), Meghalaya, eastern India, is ambiguous. One possible record here is Hill's (1974, p. 733) report, based on personal correspondence, that Dr. M. Bertrand (1969, p. 16) and Dr. R. K. Lahiri observed one or more large troops of M. assamensis in January 1965 at Tukreshwari Temple (ca. 26°00'N, 90°30'E) near the town of Abia (? Agia); because of the possibility of confusing M. as- samensis with M. mulatta (see above), the specific identification of these troops requires reconfirmation. Another possible record here is provided by an isolated skull (BM 1937.3.24.16, adult male) collected by H. W Wells, date unknown, at

FOODEN: MACACA ASSAMENSIS

U + + + + + T

M. a. assomensis

• Specimens examined

° Literature records

Fig. 2. Locality records of Macaca assamensis assamensis. Key to localities (for details, see Gazetteer): 26. Bomi. 27. Medog. 28. Tebang River. 29. Adung Valley. 30. Adung Long. 31. Tamai, Nam. 32. Goletu. 33. Taron, Wang. 34. Mokokchung. 35. Hisweht. 36. Nampuk. 37. Haibum. 38. Jantang-Dagung Hka. 39. Htingnan Triangle. 40. Mahtum. 41. Irrawady River, some miles below 2nd defile. 42. Irrawady River, 2nd defile. 43. Taping River. 44. Bhamo, near. 45. Nanding He. 46. Gengma. 47. Yado. 48. Luang Chiang Dao, Doi. 49. Puan, Nam Mae. 50. Inthanon, Doi. 51. Ban Mae Lamao. 52. Ban Pong Nam Ron, 25 km. W. 53. Ban Pong Nam Ron, 8 km. W. 54. Ban Pong Nam Ron, 5 km. W. 55. Ko Keow. 56. Khao Nang Rum, NW. 57. Chang Tai, Huai. 58. Khao Nang Rum, ranger station. 59. Ban Muang Baw Ngam. 60. Chongkrong. 61. Lancang Jiang. 62. Lo Tiao. 63. Mekong River 64. Ou, Nam. 65. Muang Ngoy. 66. Muong Moun. 67. Chapa. 68. Longzhou Xian. 69. Hoi Xuan. 70. Muang Thateng.

"Laiterai, Khasi Hills" (ca. 25°30'N, 91°30'E). Although this skull was originally identified as M. speciosa ( = M. arctoides) by Pocock (1939, p. 76), it may instead be M. assamensis (cf. Pocock, 1939, p. 70). To be conservative, these equivocal records are not included in the distribution given above for M. assamensis or in the distribution maps (figs. 1, 2).

6 FIELDIANA: ZOOLOGY

Pelage (Fig. 3)

Dorsal and lateral surface of trunk brown, varying individually from golden brown to dark chocolate brown, usually with scapular region somewhat brighter, paler, and longer haired than lumbosacral region; individual dorsal hairs usually annulated on distal third (alternating bands of pale golden and dark brown), unannulated grayish brown on proximal two-thirds; outer surface of limbs col- ored approximately like adjacent surface of trunk, or slightly paler; dorsal surface of tail colored approximately like adjacent surface of trunk, occasionally becom- ing paler distally, ventral surface of tail paler than dorsal surface; ventral surface of trunk and limbs thinly haired, pale buffy to whitish, exposed ventral skin pigmented pale bluish; crown colored approximately like back, with hair variably smooth, tufted, or forming a rudimentary cap; cheek tufts and chin whiskers usually prominent in adults, buffy to whitish; circumfacial marginal hairs black- ish; face thinly haired, supraorbital skin clearly defined pale pinkish to whitish, skin of muzzle darker, brownish to purplish.

Pelage color in 96 M. assamensis adult, subadult, and juvenile (post-infant) skins examined shows no consistent pattern of local, subspecific, sexual, or age variation. The full range of prime pelage color variation in M. assamensis assa- mensis is exhibited by a series of three subadult and adult males collected at Chapa, northern Vietnam, in November-December 1929 (FMNH 39164, sub- adult, medium golden brown; FMNH 39162, adult, dark golden brown; FMNH 39163, subadult, dark brown). These eastern specimens of M. a. assamensis are matched by western adult male specimens of the same subspecies collected in eastern India and northern Burma (FMNH 82808, Mokokchung, medium golden brown; AMNH 112736, Jantang-Dagung Hka, dark golden brown; BM 1937.3.24. 10, Tebang River, dark brown). Coloration in M. assamensis pelops adult males (FMNH 35454, Singtam, Sikkim, medium golden brown; FMNH 82809, Dikchu, Sikkim, dark brown) also matches that in the Chapa series of M. a. assamensis; these two subspecies are distinguished by tail length (see next section), not color. Adult females in both subspecies are colored approximately like adult males, as are juveniles of both sexes. Early infantile pelage is distinctively drab grayish brown (FMNH 39161, Chapa). One unusual adult male specimen (FMNH 31765, Muong Moun) has the dorsal surface marked by a broadly distributed sprinkling of white hairs; judging from dental wear, this specimen is not excessively old.

A spring molt in M. assamensis has been postulated, without documentation, by Pocock (1939, p. 52). Evidence for such a seasonal pelage change in M. a. assamensis in western Thailand is now provided by a series of 11 adult and subadult specimens collected during the dry season in 1967; two were collected about midway through the dry season (17-19 January; FMNH 99622 6 , 99623 9 ) and nine were collected near the end of the same season (8 March-23 April; FMNH 99627 6\ 99629 9, 99631-32 6 6, 99633-37 9 9). In the two specimens taken in mid-dry season, pelage is in prime condition and coloration is as de- scribed above. By contrast, in eight of nine specimens taken late in the dry season (all except FMNH 99634, subadult 9, 14 April), the dorsal pelage is distinctively short, coarse, and drab, with hair banding weak or absent. Micro- scopic examination of dorsal hairs in drab specimens reveals that the distal ends of many or most of these hairs are abruptly truncated, as though the brightly banded tips (ca. 2 cm) were broken off or abraded. Presumably, this drab, worn pelage would have been replaced by bright new pelage early in the rainy season,

Fig. 3. External characters in Macaca assamensis adult males. Above, Head and fore- quarters in M. assamensis subsp., captive (courtesy Dr. Clifford J. Jolly); below, body proportions in wild-collected M. a. assamensis (THAILAND: Ban Muang Baw Ngam, Loc. No. 59).

8 FIELDIANA: ZOOLOGY

as evidently occurs in the Indian bonnet macaque, M. r. radiata (Fooden 1981, p. 9). Available evidence is inadequate to ascertain how widespread spring molting may be in M. assamensis; there is no indication of molting in an adult male specimen of M. a. assamensis (FMNH 82802) collected 26 March 1920 at Mokokchung, eastern India.

Erythrism, in which dorsal fur coloration tends to bright burnt orange, is sporadic in M. assamensis; among specimens examined, three M. a. pelops skins are exceptionally brightly colored (BNHS 5119 9, Rongli; BNHS 5121 8, Sookia Pokhari; ZMB 91098 8 , Manshitang), as are two M. a. assamensis skins (BNHS 5115 8 , Mokokchung; ZSI 11924 8 , Bhamo vicinity, captive). A bright patch of deep chestnut fur is present on the ventral surface of the root of the tail in two adult females, one M. a. pelops (BM 25.1.1.1, Gopaldhara; cf. Hill, 1974, p. 735) and the other M. a. assamensis (FMNH 38018, Thateng). Pigment reduction and pigment absence (albinism), occasionally observed in M. sinica and M. radiata (Fooden, 1979, p. 114; 1981, p. 11), have not been reported in M. assamensis.

Annulation of dorsal hairs typically is conspicuous in M. assamensis, as cor- rectly noted by Hill (1974, p. 732). Pocock's (1939, p. 52) erroneous assertion that hair annulation is only barely perceptible in this species is inexplicable, considering the usual acuity of this author's observations.

Hair arrangement on the crown of M. assamensis is highly variable (table 1), as previously indicated by Pocock (1939, p. 52; 1941, p. 470). In some specimens a well-defined whorl centered at the vertex, suggestive of a vestige or rudiment of the cap that occurs in M. sinica and M. radiata, is present (FMNH 35454 8 , Singtam); in other specimens an irregular tuft or cowlick is present on the crown; and in still others the crown hairs lie flat and are smoothly directed posteriorly (FMNH 39163 8, Chapa). Judging from specimens examined, the whorl ar- rangement tends to be slightly more frequent in M. a. pelops than in M. a. assamensis. Also variably present is a median frontal hair parting (Hill & Bern- stein, 1969, p. 4; Hill, 1974, p. 726).

Compared with M. mulatta, with which M. assamensis frequently is confused because of similar tail proportions, dorsal pelage color is distinctly brighter on the scapular area and drabber on the lumbosacral area. In M. assamensis the dorsal surface is golden brown to reddish brown anteriorly and drab brown posteriorly; in M. mulatta this surface is drab grayish brown anteriorly and bright reddish brown posteriorly. The anterior-posterior reversal of dorsal pelage brightness in these two species was first noted by Hodgson (1841, p. 1212), who also depicted the contrast in a hitherto unpublished colored drawing (fig. 10).

External Measurements and Proportions

External measurements geographically and subspecifically variable (see be- low), head and body length (HB) 437-587 mm in 13 adult females, 538-730 mm

Table 1. Crown hair arrangement in Macaca assamensis specimens examined, where determinable.

Crown hair arrangement Subspecies N Smooth Tuft/cowlick Whorl

M. a. pelops 31 9 (29%) 7 (23%) 15 (48%)

M. a. assamensis 62 16 (26%) 27 (43%) 19 (31%)

Totals 93 25(26%) 34(37%) 34(37%)

FOODEN: MACACA ASSAMENSIS 9

in 21 adult males; relative tail length (T/HB) .39-. 55 in 13 adult females, .26-. 63 in 20 adult males; relative ear length (E/HB x 100) 5.0-7.8 in 11 adult females, 5.3-7.3 in 18 adult males; weight 4.86-8.6 kg in seven adult females, 7.9-15.0 kg in 12 adult males (table 2, fig. 4).

Available collectors' measurements of head and body length permit a few tentative generalizations concerning possible geographic, subspecific, and sexual variation of adult body size in M. assamensis (table 2, fig. 4): (1) Within M. a. assamensis, HB (and weight) in 13 adult males tends to decrease from north to south (table 2, footnote 8) in accordance with Bergmann's rule (cf. Albrecht,

400 h

300

200

100 -

100

200

300

400

500 600 700

Head and body length (mm) Fig. 4. Tail length ys. head and body length in immature and adult specimens of Macaca assamensis pelops and M. a. assamensis (log-log plot, ratio scale provided for interval 0.3 to 0.6); for details concerning measurements of adults (dashed line envelopes), see Table 2. Excluded from this figure are questionable measurements of four adults (table 2) and one subadult (MCZ 35920 9, M. a. assamensis, Doi Inthanon, Loc. No. 50, HB 492 mm, T/HB 0.31); also excluded are measurements of one abnormal subadult (BM 55.1507 9, M. a. assamensis, Puan, Nam Mae, Loc. No. 49, HB 465 mm, T/HB 0.09, discussed in text). Head and body length of fetuses as shown here differs slightly from sitting height given in Table 8.

10

FIELDIANA: ZOOLOGY

Table 2. Latitudinal variation of external measurements1 and external proportions in adult Macaca assamensis.

Lo-

cality

No.

Head and

Relative

Relative

Latitude

(figs.

body length

tail length

ear length

Weight

(°N)

1,2)

(mm)

(T/HB)

(E/HB x 100)

(kg)

Adult females, M. a. pelops

27°14'

9

537,572

.47, .51

6.1, 6.3

7.9, 8.6

27°07'

20

530,533,587

.55, .44, .44

6.3, 6.7, 5.0

7.0, -, -

26°55'

14

[?] 6202

[?] .612

[?] 4.72

X±S.D. (N)

552 ±25(5)

.48 ±.05(5)

6.1 ±.6(5)

7.8 ±.8(3)

Adult females

>, M. a. assamensis

24°08'

423

437

.44

23°30'

453

480, 500

.47,.42

6.9, 7.0

20°43'

65

[?] 5804

[?] .264

16°48'

51

523

.39

6.7

6.8

16°20'

53

482,492

.42, .43

6.8, 7.5

6.2, 6.6

15°26'

70

484

.43

14°55'

59

463

.47

7.8

4.86

X±S.D.(N)

482 ±25(8)

.43 ±.03(8)

6. 6 ±.7(6)

6.9 ±1.2(4)

Adult males, M. a. pelops

27°37'

7

560

.59

7.0

12.7

27°25'

8

550

.63

7.3

10.4

27°14'

9

635

.57

6.0

12.4

27°03'

19

555

.60

7.2

27°02'

16

575, 600, 610

.56, .50, .52

7.0, 7.0, 6.4

11.3, 10.9, -

27°00'

13

565

.50

6.9

X±S.D.(N)

581 ±30(8)

.56 ±.05(8)

6.8 ±.4(8)

11.5 ±1.0(5)

Adult males,

M. a. assamensis

29°50'

265

630,665

.37, .30

6.0, 6.0

14.5, 13.0

29°19'

275

620

.26

[?] 3.2

15.0

28°00'

28

640,640

.30, .31

6.3, 6.3

10.4, -

26°36'

39

597

.36

5.3

26°08'

38

610

.39

26°06'

40

730

[?] .184

5.6

22°21'

67b

600

.36

21°42'

66

[?] 4954

[?] .504

.

•

20°22'

69

575

.40

6.8

19°14'

477

190mm

18°32'

50

602

.36

6.8

7.9

16°20'

52

538

.44

7.2

8.7

14°55'

59

555

.36

7.0

8.5

X±S.D.(N)

616±49(23)8

.35 ±.05(12)

6.3 ±.6(20)

11. Id

t 3.0(7)

1980, p. 146); however, no such tendency is apparent in HB measurements available for eight adult females of this subspecies. (2) Comparing specimens of M. a. assamensis with those of M. a. pelops collected at approximately the same latitude, 26°-30°N, adult males in the former subspecies tend to be larger (HB 642 ± 41 mm, n = 8) than those in the latter subspecies (HB 581 ± 30, n = 8); HB in adult females cannot be similarly compared in these two subspecies be- cause no measurements are available for females of M. a. assamensis collected at the appropriate latitude. (3) Finally, sexual dimorphism of HB in M. a. assa-

FOODEN: MACACA ASSAMENSIS 11

FOOTNOTES TO TABLE 2

1 Flesh measurements recorded by collectors; included in this table are data for 12 wild- collected adult specimens that also are listed by Hill (1974, p. 727), who gives incorrect measurements for four of them (Locality Nos. 8, 16, 28 [two specimens]).

2 Measurements probably inaccurate, judging from size of dry skin (cf. Pocock, 1939, p. 56).

3 Measurements reported by Anderson (1879, p. 66).

* Measurements probably inaccurate, judging from size of dry skin.

5 Measurements courtesy Dr. Feng Zuo-jian, Beijing Institute of Zoology (letter 15 Oc- tober 1980).

6 Measurements reported by Osgood (1932, p. 209).

7 Measurement reported by De Beaux (1923, p. 27), who also reports that " Vertice capo- ano" length in this specimen is 510 mm; this latter measurement, which is equivalent to crown-rump length, is not directly comparable with head and body length (tip of nose to root of tail) as reported for other specimens listed.

8 Specimens collected N of 26°00', 642 ±41(8); specimens collected S of 26°00', 574 ±28 (5).

mensis seems to exceed that in M. a. pelops; although females in both subspecies average smaller than males, in M. a. pelops specimens HB in females broadly overlaps that in males, whereas in M. a. assamensis specimens it does not.

Tail length in adult male specimens of M. a. assamensis is clearly less than in M. a. pelops, which is the basis for current taxonomic recognition of these two subspecies (table 2, fig. 4); this tail length difference was first noted by Pocock (1939, p. 55). In 13 M. a. assamensis adult males tail length is 160-240 mm (x = 209 ± 23 mm) as compared with 283-360 mm (x = 324 ± 24 mm) in eight M. a. pelops adult males; relative tail length (T/HB) in adult males in these subspecies is similarly disjunct, .26-.44 (x = .35 ± .05) in M. a. assamensis and .50-. 63 (x = .56 ± .05) in M. a. pelops. Tail length in adult females is less differentiated in these subspecies, 193-225 mm (x = 208 ± 10 mm) in eight M. a. assamensis specimens and 236-293 mm (x = 265 ± 25 mm) in five M. a. pelops specimens; relative tail length in females — unlike that in adult males — overlaps slightly in M. a. assamensis (.39-.47; x = .43 ± .03) and M. a. pelops (.44-.55; x = .48 ± .05). Since the geographic range of M. a. assamensis extends farther south than that of M. a. pelops, the shorter tail in M. a. assamensis is a subspecies- level departure from Allen's rule, which otherwise applies to tail length reduction in the sinica group of macaques (Fooden, 1971b, p. 70).

In M. a. pelops relative tail length in adult males averages greater than in adult females, whereas in M. a. assamensis relative tail length in adult females averages greater than in adult males (table 2). Judging from relative tail length in immature specimens examined (fig. 4), this reversal apparently results from subspecific and sexual differences in the growth curve of tail length relative to head and body length. Adult males in M. a. pelops and adult females in M. a. assamensis retain approximately the same tail proportions as immatures of both sexes in their respective subspecies, whereas adult females in M. a. pelops and adult males in M. a. assamensis have tails that are relatively shorter than immatures in their subspecies. This seems to imply that tail growth is coordinate with head and body growth throughout postnatal development in M. a. pelops males and

12 FIELDIANA: ZOOLOGY

M. a. assamensis females. However, in M. a. pelops females and M. a. assamensis males the coordinate growth of tail length relative to head and body length apparently ceases at or near adolescence, subsequent to which tail growth ap- parently decelerates or stops during the final late adolescent phase of head and body growth.

Relative tail length in three subadult M. a. assamensis specimens examined is apparently greater than normal in this subspecies (fig. 4). Two of these specimens were collected in northern Burma (AMNH 112738 6, Nampuk: HB 521 mm, T/ HB .56; AMNH 112737 9, Haibum: HB 483 mm, T/HB .53) and one in western Thailand (FMNH 99632 6, Ban Pong Nam Ron, 25 km W: HB 485 mm, T/HB .52). The significance of unusually long tails in these subadult specimens is unclear.

The tail is abnormally short in one subadult female specimen of M. a. assamensis (BM 55.1507: HB 465 mm, T/HB .09) collected at Nam Mae Puan, northern Thailand, as previously noted by Kloss (1919, p. 50); the form and pelage of the tail do not suggest artificial mutilation (Kloss, loc. cit.). The same specimen also has a malformed right hand with only two fingers present; perhaps the tail and hand abnormalities are developmentally interrelated.

Cranial Characters (Fig. 5; Tables 3, 4)

Skull moderately large, greatest length (GL) 120.2 ± 5.2 mm in 20 adult females, 143.3 ± 6.7 mm in 29 adult males; relative zygomatic breadth (ZB/GL) moderately great, 0.66 ± .02 in 20 adult females, 0.66 ± .03 in 29 adult males; rostrum moderately projecting, rostral-postrostral ratio 0.48 ± .03 in 17 adult females, 0.57 ± .04 in 23 adult males; supramaxillary ridges well defined, par- ticularly in adult males, arching posterosuperiorly from canine alveolus to in- fraorbital rim; supraorbital ridges thick (7-13 mm in adult males); temporal lines well defined, convergent in adult males, frequently uniting to form a prominent sagittal crest (18 mm high in FMNH 35454, Singtam); nuchal crest prominent in adult males (19 mm high in FMNH 35454).

The subspecies M. a. pelops and M. a. assamensis broadly overlap in cranial characters. In M. a. assamensis, skull size tends to be greater in northern spec- imens collected between 26° and 28° North Latitude, which is the latitude of the main range of M. a. pelops, than in more southern specimens (table 4, footnotes 4-5; cf. Albrecht, 1980, p. 146). Individual cranial variation in adult M. assamensis specimens includes two specimens with healed fractures of the zygomatic arch (BM 15.9.1.3 6, Sookia Pokhari; BM 1937.3.24.8 S, Batasia), four specimens with premolars or incisors absent congeni tally or lost early in life (BM 15.9.1.3 S , Sookia Pokhari; BM 1937.3.24.10 6, Tebang River; BM 1937.3.24.11 6, Tebang River; FMNH 35452 9, Singtam), and one specimen with two seemingly un- related cranial anomalies (FMNH 39163 6 , Chapa: right brow ridge conspicu- ously deficient as a result of unilateral deformity of frontal bone; two lower incisors absent, apparently congenitally).

Ontogenetically, relative zygomatic breadth in M. assamensis remains approx- imately constant from infancy to adulthood (table 3); this differs from the pattern of zygomatic arch development in previously studied M. sinica and M. radiata, where relative zygomatic breadth increases with age (Fooden, 1979, table 2; 1981, table 4). Rostral length in M. assamensis increases allometrically relative to post- rostral length (fig. 6). The power function equation for the ontogenetic relation-

13

Table 3. Cranial dimensions and ratios (X±S.D.) in various age classes of Macaca as- samensis; sample size indicated by italicized figure in parentheses.

Skull,

Relative

greatest

zygomatic

Postrostral

Rostral-

length

breadth

length

postrostral

Age class

(mm)

(ZB/GL)

(mm)

ratio

Infants

81.3 ±3.6(9)

.65 ±.04(8)

70.2 ±2.3(9)

.24 ±.03(8)

Juveniles

102.8 ±8.6(25)

.66 ±.02(15)

80.8 ±4.8(14)

.39 ±.05(14)

Subadult females

111.5±3.4(6)

.66 ±.03(6)

83.0 ±1.5(6)

.44 ±.04(5)

Adult females

120.2 ±5.2(20)

.66 ±.02(20)

87.5 ±3.5(1 7)

.49 ±.03(2 7)

Subadult males

129.9 ±6.0(9)

.66 ±.02(9)

90.3±5.1(7)

.54 ±.04(7)

Adult males

143.3 ±6.7(29)

.66 ±.03(29)

98.1 ±3.9(23)

.57 ±.04(23)

Table 4. Latitudinal variation of greatest skull length in adult Macaca assamensis.

Locality

No.

Greatest skull

length (mm)

Latitude

(figs. 1, 2)

Adult females M. a. pelops

Adult males

ca. 27°30'

"Nepal"

119.5

131.9

27°37'

7

143.3

27°25'

8

139.8

27°14'

9

119.6, 123.7

154.1

27°07'

20

1161, 117.9, 127.4

27°05'

19

136.5

27°02'

16

. . .

140.6, 143.9, 144.0, 149.9

27°00'

13

144.5

26°55'

14

131.5

22°35'

25

...

134.62

Mean±S.D.

122.2 ± 5.6 (N

= 7)

142.1 ±6.5 (N = 11)

M. a. assamensis

ca. 28°00'

28

138.1, 156.1

27°40'

33

ca. 139.5

27°37'

32

. . .

144.6

26°36'

39

160.3

26°20'

34

124.6, 126.8

144.6, 145.8, 150.4

26°08'

38

146.1

24°08'

42

114.3

23°30'

45

116.4, 117.6

• ■ • ^->

22°21'

67

124.4

129.6, 142.8

21°42'

66

139.9

20°43'

65

118.9

20°22'

69

ca. 141.9

20°20'

62

139.6

19°23'

48

113.6

19°14'

47

1463

18°32'

50

142.1

16°48'

51

117.9

16°20'

52,53

120.4, 126.0

138.9

15°26'

70

115.3

14°55'

59

113.1

145.0

Mean±S.D.

119.2±4.8 (N

= 13)4

144.0 ± 6.9 (N = 18)5

1 Measurement from Khajuria & Ghose, 1970, p. 23.

2 Measurement from Anderson, 1872, p. 532.

3 Measurement from De Beaux, 1923, p. 27. Specimens collected N of 26°00',125.7±1.6 (N = 2);

118.0±4.2(N = 11).

5 Specimens collected N of 26°00', 147.3 ±7.2 (N = 9); 140.6 ± 4.8 (N = 9).

specimens collected S of 26°00' specimens collected S of 26°00'

14

ADULT MALES

M 100

Postrostral length (mm)

Fig. 6. Allometry of rostral length (y) vs. postrostral length (x) in immature and adult specimens of Macaca assamensis (log-log plot; open circles = females, closed circles = males; smaller circles = immatures, larger circles = matures; solid line = principal axis, log y = 3.675 log x-5.531), compared with corresponding allometry (dashed lines = principal axes) in M. sinica (log y = 4.317 log x-6.507; Fooden, 1979, p. 115) and M. radiata (log y = 3.396 log x- 4.860; Fooden, 1981, fig. 5).

15

16

FIELDIANA: ZOOLOGY

ship in this species between rostral length (y) and postrostral length (x) deter- mined by the method of principal axes is:

log y = 3.675 log x - 5.531 ; 95 percent confidence limits for the slope are Lj = 3.362 and L2 = 4.046.

Comparing adult M. assamensis (table 3) with adult M. sinica and M. radiata (Fooden, 1979, p. 114; 1981, table 3), skull size in both sexes of M. assamensis is, invariably and usually, conspicuously greater than in the latter two species, which overlap in size. The known range of greatest skull length in M. sinica and M. radiata is 92.8-107.7 mm in adult females and 106.7-127.8 mm in adult males; the corresponding range in M. assamensis is 113.1-131.5 mm in adult females and 131.9-160.3 mm in adult males. Although skull size differs in M. sinica, M. radiata and M. assamensis, skull shape, including relative zygomatic breadth and rostral-postrostral ratio, is remarkably similar in these species (cf. Kurup, 1966, p. 74; Albrecht, 1978, p. 106).

Natural History

Both subspecies of M. assamensis are mainly restricted to a relatively narrow altitudinal zone between about 150 and 1,900 m above sea level (table 5; cf. Zhang et al., 1981, p. 222). Only five of 58 known altitudinal records are above this zone, to a maximum of 2,750 m, and only one record (widely disjunct) is below this zone, near sea level in the Sundarbans tidal mangrove swamps near

Table 5. Altitudinal distribution of Macaco assamensis localities for which altitude is specified by collector or observer; for details, see Gazetteer.

Altitude (m)

0

100

200

300

400

500

600

700

800

900 1,000 1,100 1,200 1,300 1,400 1,500 1,600 1,700 1,800 1,900 2,000 2,100 2,200 2,300 2,400 2,500 2,600 2,700

M. a. pelops

25

20,21 20

8

11

13, 19 19

1, 15

17

9

9, 14, 16

7, 9, 18

17 12

3,5

Locality Nos. (figs. 1,2)

M. a. assamensis

35

53, 54, 55

51, 66

36, 37, 56, 57

49, 56

28, 56

38, 52, 56, 60, 65

27, 31, 32, 70

39, 47 59

31,40 31,34

30, 31, 34, 62 29, 33, 50

67, Guangxi

26

FOODEN: MAC AC A ASSAMENSIS 17

the mouths of the Ganges (see above, Distribution). Upper elevation Himalayan populations are reported to migrate to lower altitudes in winter (C.A. Crump in Wroughton, 1916b, p. 476; Shou Chen-huang et al., 1964, p. 62). The preferred winter altitudinal range in Sikkim is said by Crump to be 600-1,200 m, but Sikkimese specimens have been collected above 1,600 m in November-February (Chuntang, 1,630 m; Singtam, 1,645 m; Manshitang, 1,830 m; for details, see Gazetteer).

The geographic and altitudinal distribution of M. assamensis closely corre- sponds to the similarly restricted distribution of subtropical mid-elevation moun- tain broadleaf evergreen rain forest (figs. 1, 2). The close association of both subspecies of M. assamensis with dense primary evergreen forest has been noted repeatedly by collectors and observers (C.A. Crump in Wroughton, 1916b, p. 476; Andrews, 1918, pp. 244, 253; Mills, 1923, p. 222; Lord Cranbrook, 1931, BM specimen tag, Adung Valley; Kaulback, 1938, pp. 17, 29; Kaulback, 1938-1939, BM specimen tags, Nam Tamai Valley, Adung Long, Taron Valley, Goletu, Hting- nan Triangle; R. L. Fernandez & H. Khajuria, 1958, ZSI specimen tags, Tarkhola; Shou et al., 1964, p. 62; Khajuria, 1966, p. 284; Fooden, 1971a, p. 36; Chakraborty, 1975, p. 18; Eudey, 1979, p. 95). A few troops of M. assamensis have been recorded in deciduous or bamboo forest and in cultivated fields, but these records are all in the immediate vicinity of evergreen forest, which almost certainly was the primary habitat of these troops (Jerdon, 1867, p. 12; Kaulback, 1939, BM 50.385 specimen tag, Htingnan Triangle; Khajuria, 1966, p. 284; Fooden, 1971a, p. 36; Eudey, 1979, p. 94).

Behavioral observations of natural populations of M. assamensis are scanty, undoubtedly because of the restricted and relatively inaccessible rain forest hab- itat of this species. Judging from stomach contents in 10 specimens of M. a. assamensis collected in western Thailand, fruit is the principal natural food of this species (Fooden, 1971a, p. 36), as is true of other species of macaques. However, stomachs in three of these specimens also contained a few insect fragments, indicating that insectivory is fairly frequent, and cheek pouches of one of the three insect eaters contained the head of a lizard (Japalura sp.), indicating that small vertebrates also are sometimes eaten. Shou et al. (1964, p. 62) report a similar mixed diet, with fruit as the main component, for the same subspecies in Guangxi, southeastern China. In western Thailand, Eudey (1979, pp. 94, 97) observed this subspecies feeding on figs (Ficus spp.) and other fruits, and in Yunnan, southwestern China, Andrews (1918, p. 255) saw a troop in the canopy about 50 m above ground "feeding upon some large green beans" (pre- sumably Leguminosae). Both M. a. assamensis and M. a. pelops are known to raid fields of maize planted in forest clearings (Jerdon, 1867, p. 12; Stevens, 1923, BM specimen tag, Gopaldhara; Kaulback, 1938, p. 29; Kaulback, 1939, BM spec- imen tag, Mahtum; ZSI Darjeeling District Expedition, ZSI specimen tag, Takdah; Khajuria, 1966, p. 284). When feeding in trees, M. assamensis gathers food either by breaking off terminal branches that bear several fruits or by plucking indi- vidual fruits (Eudey, 1979, p. 97).

Except for troops that were raiding maize fields, all reported troops of M. assamensis were in trees, usually high in the canopy, when originally observed (Blanford, 1871, p. 375; Andrews, 1918, p. 253; Fooden, 1971a, p. 36; Eudey, 1979, pp. 94-98). Judging from available evidence, M. assamensis is much more strongly arboreal than its near relative M. radiata (Fooden, 1981, p. 16); this is somewhat unexpected, because the shorter tail of M. assamensis usually would

18

FIELDIANA: ZOOLOGY

be assumed to indicate terrestriality. Troops of M. assamensis in the canopy have twice been observed in association with hornbills (Bucerotidae) in lower branches of the same or nearby trees (Andrews, 1918, p. 252; Fooden, 21 January 1967, Ban Muang Baw Ngam, unpublished notes). When frightened or disturbed, as by gunfire, M. assamensis apparently exhibits variable escape behavior: the mon- keys may flee through the branches of trees (Andrews, 1918, p. 253), descend to the ground and withdraw quietly through the undergrowth (C.A. Crump in Wroughton, 1916b, p. 476; Eudey, 1979, p. 95), or flee first through the trees and then on the ground (Shou et al., 1964, p. 62). The movements of M. assamensis are described as slower and more deliberate than those of broadly sympatric M. mulatto (Blanford, 1888, p. 15).

Recorded troop size in M. assamensis averages about 20 individuals per troop, ranging from about 12 to 50 (table 6). No solitary individuals of this species have been observed or reported. The mean ratio of immatures (non-reproductives) to subadults and adults (reproductives) in three troops is 0.66; among repro- ductives, the mean ratio of males to females is 0.44 (table 7).

Notes on social behavior within troops of M. assamensis in their natural habitat appear to have been recorded by only five observers. These few notes on in- tratroop social behavior, quoted below, generally accord with comparable ob-

Table 6. Individuals per troop reported in Macaca assamensis.

Mean No. troop

Sample troops No. individuals member-

area observed in each troop ship Reference

M. a. pelops

W Bengal

2

13-20, 10-25

17

Southwick et al., 1964, p. 446

W Bengal

?

3=12

12

Khajuria, 1966, p. 284

Bhutan

1

ca. 20

M. a. assamensis

ca. 20

Chakraborty, 1975, p. 18

N Thailand

2

12, 26

19

Carpenter, 1942, p. 185

W Thailand

11

10, 10, 12, 20, 20, 20, 24, 25, 30, 30, 50

23

Fooden, 1971a, p. 36

W Thailand

6

>10, >20, >20, >20, >20, >20

>20

Eudey, 1979, pp. 94-97

Table 7. Age-sex composition in three troops of Macaca assamensis Adults1

Total

troop

membership

Total

9$

Sex

ratio

6 6 (6 6:9 9) Total

Immatures Juv.

Inf.

Age

ratio

(Im.:Ad.)

13

8

5

Af. a. pelops2

3 .60 5

M. a. assamensis3

3

2

.63

12 26

8 14

6 10

2 .33 4 4 .40 12

2 8

2 4

.50 .86

1 Probably also includes some subadults.

2 West Bengal; Southwick et al., 1964, p. 446.

3 Northern Thailand; Carpenter, 1942, p. 185.

FOODEN: MAC AC A ASSAMENSIS 19

servations, particularly of troop control by large adult males, of M. sinica and M. radiata (Fooden, 1979, p. 125; 1981, p. 26).

1. C. A. Crump in Wroughton (1916b, p. 476), M. a. pelops, northern West Bengal and Sikkim: 'The call is a loud 'pio', rather a musical note, repeated frequently. The warning cry is 'pio', uttered once by the sentry, who is generally on the lookout high up in a tree; the whole band then descends to the ground and moves away in absolute silence, concealed by the dense undergrowth."

2. G. C. Shortridge (1920, FMNH 82808, specimen tag), M. a. assamensis, Mokokchung, Nagaland: "Very old uncle. Leader of troupe. About limit of size reached by this species here."

3. H. Khajuria (1966, p. 284), M. a. pelops, northern West Bengal: "A lonely sick (?) infant was observed moving slowly under thick bushes in the forest. When approached it gave out a few squeals. The mother immediately appeared on the spot from a nearby bush with a threatening disposition. The other mem- bers of the troupe could be seen away on the trees. The infant was shot but the mother would not allow its body to be removed even when the gun was pointed at her. She climbed a nearby tree and jumped from branch to branch in great restlessness. She would come down to attack if a nearer approach to the dead young was attempted. She did not pay any heed to the other members of the troupe which at once disappeared into the deep forest at the sight of the gun. She had to be shot down to end her agony."

4. J. Fooden (1967, unpublished field notes), M. a. assamensis, Ban Muang Baw Ngam, western Thailand: "There were about 20 individuals moving in a fairly organized band. . . . The last one in the band to cross a particular gap between branches was a half-grown young one. ... A big individual may have been left behind the troop to the right of the trail."

5. A. A. Eudey (1979, pp. 94-98), M. a. assamensis, western Thailand: "The group crossed huay Chang Tai from west to east and moved south along the stream about 0.7 km to enter feeding and sleeping trees. Continuous vocaliza- tions accompanied this progress. . . .

". . . The monkeys {another troop, 20 members] were dispersed over ap- proximately 365 m in the trees at a height of 15 m. The monkeys when contacted crossed the stream to the east and ascended a mountain ridge covered with bamboo, traveling about 0.8 km. At the crest of the ridge clear calls began to the north at 11:55. At 12:00 calling also occurred to the east. My guide and I appeared to be in the middle of a dispersed group. At 12:06 two large adult males moved slowly and deliberately from N/NE to E in the lower story of mixed deciduous vegetation, perhaps 6 m above ground. The larger of the two monkeys went first and was immediately followed by the somewhat smaller [adult] male. Both monkeys were heavy set ... . Both monkeys vocalized as they moved through the trees, making a clear 'uh uh uh' sound. The somewhat smaller male vocalized more than the larger. By 12:10 the males had passed from view although calling continued from the east. From 12:15 until 12:56 a number of small monkeys . . . moved from the south and west to the east during continuous vocalizations. 'Who' and shrill 'aho' sounds seemed to be made by the small monkeys.

". . . No vocalizations accompanied . . . feeding behavior [in another troop].

". . . Subsequent to the encounter [with M. mulatto], the group of M. assamensis remained in the trees in a ravine, keeping the [human] observers under sur- veillance. A monotonous call was made by one adult male which assumed a

20 FIELDIANA: ZOOLOGY

posture with penis exposed. Sometimes this male was joined by other, unseen monkeys. Other monkeys were observed to look, scratch, and move around in the trees but not to participate in the calling. One female which from the con- dition of her nipples appeared to be [nulliparous] was groomed by a larger female with elongated nipples for over 5 minutes at a distance of no more than 3 m from the calling male."

Judging from evidence of five pregnant females and three infants, births in M. assamensis apparently are concentrated in the period April-August (table 8), which generally coincides with the first two-thirds of the rainy season in the range of this species (Walter et al., 1975, map 5); an exceptional record is of an infant (BM 50.378) that seems to have been born in the interval December-March. Assuming a gestation period of approximately 5.5 months, as in other macaques (Napier & Napier, 1967, p. 406), it may be inferred that most copulations probably occur in the period November-March, which generally would be during the dry season. Of nine sexually mature females collected in western Thailand in Jan- uary-April 1967, seven (including four subadults) were either pregnant or lac- tating (Fooden, 1971a, p. 37; 1971b, p. 68); this suggests that most sexually mature females probably bear one young each year. Mean length of the menstrual cycle in 14 captive females was 32 ± 2 days (Wehrenberg et al., 1980, p. 523).

Table 8. Evidence of probable reproductive seasonality in Macaca assamensis pregnant females and infants.

Locality

Sitting height

No.

Date of

of fetus;

Estimated

(figs.

collection or

Specimen

dentition of

birth

1,2)

observation

No.

infant1

month2

M. a

pelops, northern West

Bengal

20

Aug. 1958

"Several unweaned in- fants . . . were ob- served."3

"Possibly born in the preceding spring"3

M.a

. assamensis, northern Burma

31

28 March 1938

BM 50.378 (inf.)

(c, ml)

Dec-March

31

9 Sept. 1938

BM 50.379 (inf.)

(c, ml)

June-Aug.

31

23 Sept. 1938

BM 50.380 (inf.)

(c, ml)

June-Sept.

42

3 March 1875

ZSI 11909, "gravid"4

April-June

M. a.

assamensis, western Thailand

51

25 March 1967

FMNH 99629 (incl. fetus)

170 mm

April

52

14 April 1967

FMNH 99633 (incl. fetus)

140 mm

June

52

14 April 1967

FMNH 99634 (incl. fetus)

140 mm

June

54

20 April 1967

FMNH 99635 (incl. fetus)

170 mm

June

M.a.

assamensis, northern Vietnam

67

6 Nov. 1929

FMNH 39161 (inf.)

(m2)

May-Sept.

1 Parenthetical notation indicates deciduous teeth in process of eruption.

2 Infant age estimates based on dental emergence norms for M. mulatta (Hurme, 1960, p. 796); fetal conception age estimates based sitting height norms for M. mulatta (Wagenen, 1972, p. 10).

3 Khajuria, 1966, p. 284.

4 Anderson, 1879, p. 65.

FOODEN: MACACA ASSAMENSIS 21

Relations between humans and M. assamensis vary locally, from veneration of the monkeys to persecution. At two Buddhist pagodas along the Irrawady River below Bhamo, northern Burma, resident troops of M. assamensis are or were maintained and fed by local worshippers (Anderson, 1871, p. 213; 1879, p. 65). More usually, including elsewhere in northern Burma, this monkey is regarded as an agricultural pest because of its frequent raiding of maize fields (see above). In local areas in Sikkim and adjacent West Bengal, M. assamensis is hunted and eaten (C. A. Crump in Wroughton, 1916b, p. 476; Khajuria, 1966, p. 284). In parts of China, and undoubtedly elsewhere, the species has been hunted to near extinction (Shou et al., 1964, p. 62).

ECOGEOGRAPHIC RELATIONS WITH OTHER NON-HUMAN PRIMATES

Macaca assamensis has been collected or observed in the vicinity of four non- macaque primate genera — Hylobates (gibbon), Presbytis and Pygathrix (leaf mon- keys), and Nycticebus (slow loris) — and also in the vicinity of three other species of macaques, M. arctoides (bear macaque), M. nemestrina (pigtail macaque), and M. mulatto (rhesus macaque) (table 9). One additional species of macaque, M. fascicularis (crab-eating macaque), is marginally associated with M. assamensis along the extreme southern edge of the range of M. assamensis (Fooden, 1971a, p. 28). All of these associated genera or species occur southeast of the Brah- maputra River (Assam Valley) in or near the subspecific range of M. a. assamensis; northwest of the Brahmaputra River, in or near the range of M. a. pelops, the only associated primate genera or species are Presbytis spp. and M. mulatta.

Relationships with non-macaque primate genera. — The geographic ranges of three of the associated genera — Hylobates, Presbytis, Nycticebus — broadly overlap the range of M. assamensis (Pocock, 1939, p. 165; Fooden, 1969, p. 640; 1975b, p. 108); the range of Pygathrix narrowly overlaps that of M. assamensis only in northern Vietnam (Groves, 1970, p. 585). Ecologically, Hylobates spp., Presbytis spp., and M. assamensis are known to share the same upland evergreen habitat; Hylobates spp., Presbytis spp., and M. assamensis frequently have been collected or observed within a period of a few days in the same tracts of forest (Andrews, 1918, p. 253; Mills, 1923, p. 221; Kingdon Ward, 1937, p. 337; Kaulback, 1938, pp. 17, 29; Bannikov, 1958, p. 68; Fooden, 1971a, p. 10, Ban Muang Baw Ngam, Chongkrong; Eudey, 1979, p. 129). In fact, Andrews (1918, pp. 253-55) observed H. lar, P. phayrei, and M. assamensis feeding together in the same tree at the same time. At least in northern Burma, however, the range of M. assamensis apparently extends to higher elevations than that of either Hylobates or Presbytis (Kingdon Ward, 1937, p. 337). Pygathrix and Nycticebus probably also inhabit some of the same forest tracts as M. assamensis, but evidence for this is not conclusive (Os- good, 1932, pp. 204, 210, 213; Carter, 1943, p. 106; Fooden, 1971a, p. 21).

Presumably differences in dietary preferences partly mitigate competition be- tween M. assamensis and the more folivorous Presbytis and Pygathrix on the one hand and the more insectivorous (and nocturnal) Nycticebus on the other (Mac- kenzie in Fry, 1929, p. 641; Fooden, 1971a, p. 12). Such differences in dietary preferences are unknown and unlikely for M. assamensis and Hylobates, both of which apparently feed mostly on the same species of fruits (Fooden, 1971a, p. 12); however, Macaca and Hylobates are known to differ in their technique and efficiency of arboreal harvesting of fruit (Grand, 1972, p. 198).

Relationship with Macaca arctoides. — Of the closely associated species of ma-

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FOODEN: MACACA ASSAMENS1S 23

caques (M. arctoides, M. nemestrina, M. mulatto), M. arctoides broadly overlaps M. assamensis both geographically (Fooden, 1980, p. 5) and ecologically. In western Thailand these two species were collected a few kilometers apart in the same forest in a single morning (Fooden, 1971a, p. 36, Chongkrong, 28 January 1967). Eudey (1979, p. 89) also may have observed M. arctoides twice in western Thailand within a few kilometers of places where she observed M. assamensis; however, these identifications of M. arctoides are provisional because Eudey caught only fleeting glimpses of the monkeys and therefore was unable to discern either the distinctive red face or the stump tail. Although diets of M. arctoides and M. assamensis are essentially similar (Fooden, 1971a, p. 12), competition between these macaques probably is not severe: the former species usually forages on the forest floor (McCann, 1933, p. 807; Bertrand, 1969, p. 145; Fooden, 1971a, p. 38) and the latter usually forages high in the canopy (see above, Natural history).

Relationship with Macaca nemestrina. — Macaca nemestrina, like M. assamensis and M. arctoides, apparently is mainly restricted to evergreen rain forest (Fooden, 1975a, p. 51). Although the geographic ranges of M. nemestrina and M. assamensis closely interdigitate along the foothills of Southeast Asian mountain chains (fig. 7), these specific ranges appear to be almost perfectly complementary and non- overlapping (parapatric). The range of M. n. leonina, the local subspecies of M. nemestrina, generally is more southern and/or at lower elevations than that of M. assamensis. The complementary distributions of M. n. leonina and M. assa- mensis, together with the similarity of their diets and arboreal habits (Fooden, 1971a, p. 12), suggest a relationship of competitive exclusion in this area between these two species; this would explain the absence, previously noted as enigmatic (Fooden, 1975a, p. 72), of M. n. leonina from apparently suitable rain forest habitats in central and northern Laos and northern Vietnam, within the range of M. assamensis. If competitive exclusion is operating, M. n. leonina evidently is the more successful competitor in tropical forest, and M. assamensis is the more successful competitor in subtropical forest (cf. Trewartha in Espenshade & Morrison, 1978, p. 9, climate map). Body size in M. n. leonina (Fooden, 1975a, pp. 86, 114) averages smaller than in M. assamensis (table 2), and this difference may confer a thermoregulatory advantage on each of these species in their respective tropical and subtropical ranges. (Although the subspecies M. n. ne- mestrina is larger in body size than either M. n. leonina or M. assamensis and has a more equatorial distribution, it also differs behaviorally, being more strongly terrestrial; Fooden, 1975a, pp. 86, 99, 103.) M. n. leonina and M. assamensis have been collected within a few kilometers of each other, sometimes on the same day, along the frontier between their respective ranges (table 9: NE India; NW Burma; south-central Burma; W Thailand), but direct contact between these species has not been observed or reported.

Relationship with Macaca mulatto. — The geographic and ecological relationships between M. assamensis and M. mulatto, the rhesus macaque, are particularly close and complex. These two species are broadly sympatric (fig. 8A), with the rel- atively small range of M. assamensis almost completely contained asymmetrically within the much larger range of M. mulatto. The latitudinal correspondence of these two specific ranges, especially the similar southeastern margins that ob- viously are associated with the relatively cool Indo-Chinese uplands (within nine degrees of the Tropic of Cancer), strongly suggests that both of these macaques

24

FIELDIANA: ZOOLOGY

M. a. ossomensis

M. nemestrina I e o n i n a

inferred interspecific frontier

Fig. 7. Geographic relationship between known localities of Macaca assamensis assamensis (fig. 2) and nearby known localities of M. nemestrina leonina (Fooden, 1975a, fig. 28; Biswas & Diengdoh, 1978, p. 344).

are primarily adapted to a subtropical climate. This also is indicated by the similar degree of tail reduction in these two species (Fooden, 1980, p. 7).

Although M. assamensis and M. mulatta are broadly sympatric and apparently similar in climatic adaptation, these two species usually are not encountered at exactly the same localities (figs. 8B, 9), which suggests that they probably are divergent in specific habitat preferences. This is most clearly evident in the ecogeographic relationship between M. a. pelops and M. mulatta along the south- ern flank of the Himalayas, where the ranges of these two species obviously are altitudinally stratified (fig. 8B). In Sikkim and adjacent northern West Bengal, the best studied portion of this sub-Himalayan area, the known distributions of these two macaques are almost perfectly nonoverlapping (fig. 8B, inset), with M. mulatta restricted to lower elevations (100-1,100 m) and M. a. pelops inhabiting higher elevations (250-2,700 m).

FOODEN: MACACA ASSAMENSIS 25

The altitudinal stratification of M. a. pelops and M. mulatto in the Sikkim-West Bengal area corresponds to local vegetational stratification (Schweinfurth, 1957, Vegetationskarte, Beilage). The range of M. a. pelops coincides with the zone of subtropical mid-elevation mountain broadleaf evergreen rain forest, as previ- ously noted (see above, Natural history). By contrast, the lower altitudinal range of M. mulatta in this area coincides with the zone of tropical deciduous wet sal forest, known locally as "terai." The association of M. mulatta and "terai" forest that is implied by the altitudinal correspondence of their ranges is confirmed by available field observations:

1. C. A. Crump (in Wroughton, 1916b, p. 475) — "It [M. mulatta] occurs in the Terai and foot hills below Darjeeling."

2. H. Stevens (1930, FMNH 35447, Sivok, M. mulatta, tag)— "Terai."

3. C. H. Southwick et al. (1964, p. 443) — "All of the rhesus [M. mulatta] groups were seen in a narrow band of terai forest at the base of the Himalayan foothills in northern Bengal."

4. H. Khajuria (1966, p. 284) — see quotation in next paragraph.

M. mulatta has never been recorded in the mid-elevation broadleaf evergreen forest that is the prime habitat of M. a. pelops. However, both macaques have been observed together once in the Sikkim-West Bengal area at a marginal locality near the lower limit of distribution of M. a. pelops:

"At Tarkhola 325 m . . . [M. a. pelops] frequented villages surrounded by forest consisting mainly of the sal (Shorea robusta) and the teak (Tectona grandis) and fearlessly raided maize fields. ... It occurred side by side with the rhesus Macaque [M. mulatta] . . . but was not seen to mix with it."

This is the only reported instance of joint occurrence of M. a. pelops and M. mulatta in 23 macaque locality records known for the Sikkim-West Bengal area (M. a. pelops, 16 localities; M. mulatta, six localities; both species, one locality).

Although M. mulatta in the Sikkim-West Bengal area is absent from cool moist mid-altitude broadleaf evergreen forests that are inhabited by M. a. pelops, evi- dence from other areas shows that M. mulatta is not inherently less tolerant of upper altitudes and low temperatures. In western Nepal (Hutu forest, 3,050 m) and in southwestern China (Dzo La ca. 4,000 m; Rama Pass vicinity, ca. 5,000 m), M. mulatta inhabits cool to cold upland coniferous forests (Richie et al., 1978, p. 443; Kaulback, 1938, p. 91; Schafer, 1933, p. 280) that typically are not inhabited by M. assamensis and that are well above the upper altitudinal limit of M. assa- mensis. This suggests that the distributions of M. mulatta and M. a. pelops in the Sikkim-West Bengal area are determined more directly by forest type than by temperature.

In Nepal, west of the Sikkim-West Bengal area, rainfall decreases from east to west and the broadleaf evergreen habitat of M. a. pelops terminates near Pokhara, central Nepal (Schweinfurth, 1957, p. 298; Vegetationskarte, Beilage). Correspondingly, in western Nepal, M. mulatta completely replaces M. a. pelops, the westernmost known report of which is a sight record at Tipling (1,220 m), central Nepal. (For discussion of a supposed record of M. assamensis at Mus- soorie, India, west of Nepal, see above, Distribution). In central Nepal, M. mulatta inhabits higher elevations (Gokarna, ca. 1,375 m, "deciduous forest"; Nagarkot, 2,450 m, forest type unspecified) than in the Sikkim-West Bengal area; presum- ably this is related to progressive westerly disappearance of broadleaf evergreen habitats and hence of M. a. pelops. The altitudinal and vegetational stratification of M. mulatta and M. a. pelops in eastern Nepal was noted by Hodgson (1841,

26

FIELDIANA: ZOOLOGY

/ M. a. /" ',~(^Cm JI,0*'0,I!£n", Jfr'

Fig. 8. A, Limits of distribution of Macaca assamensis (figs. 1, 2) and M. mulatto (Fooden, 1980, p. 5). B, Geographic relationship between known localities of M. a. pelops (fig. 1) and nearby known localities of M. mulatta (see Appendix).

p. 1213) in his original description of M. a. pelops, in which the habitat of M. mulatta is given as "Tarai [sic] and lower hills" and that of M. a. pelops is given as "Northern region of hills exclusively." Chesemore (1970, p. 164), who studied M. mulatta widely in southern Nepal, notes: "This species was usually seen at the edge of farm land and jungle and seldom in dense jungle."

Of the three mapped locality records (ca. 500-2,750 m) of M. a. pelops in eastern Nepal (fig. 8B), one, apparently a marginal locality (ca. 500 m) for this macaque, is a joint or proximate occurrence with M. mulatta (J. A. McNeely, letter, 29 July 1973):

M. assamensis is found in our area [Arun Valley, eastern Nepal], from 1000 to 10,000 feet [300-3,000 m], while M. mulatta is found from [300-1,800 m], based on preliminary examination of sight records. The interesting thing is that at the lower altitudes, particularly in the Sabaya Khola [ca. 500 m], both species are present; in the space of 2-3 kilometers on June 3, 1973, I saw two groups of each in the same habitat type, Shorea robusta [sal, deciduous] forest, heavily grazed. I have seen no evidence of intergradation, nor have I observed any interaction between groups.

East of the Sikkim-West Bengal area there are only two known records (ca. 150 m, 1,500 m) of M. a. pelops in south-central Bhutan (fig. 8B). The six nearest records of M. mulatta are restricted to an altitudinal range of 60-600 m.

M. a. pelops has been collected with or near M. mulatta 500 kilometers south of the Himalayas, near sea level along the Bay of Bengal in the Sundarbans tidal mangrove swamps (see above, Distribution), but there is no direct information concerning the nature of the association between these two species in the Sun- darbans. M. a. pelops does not now inhabit the western (Indian) part of the Sundarbans (Khajuria, 1955, p. 114), although M. mulatta does occur there (Mandal, 1964, p. 155; Mukherjee & Gupta, 1965, p. 145). If M. a. pelops is still

FOODEN: MACACA ASSAMENSIS

27

Sp«ciH>«i>i Lit*raiu>«

M. a. atsamtntit M. mulatto

Fig. 9. Geographic relationship between known localities of Macaca assamensis assamensis (fig. 2) and nearby known localities of M. mulatto (see Appendix).

extant anywhere in these swamps, it presumably is restricted to the eastern part, in what is now Bangladesh. This part of the Sundarbans receives the greatest rainfall (Imp. Gaz., 23: 141) and, judging from density trends of M. mulatto in the western Sundarbans (references above), the eastern part probably also has the highest density of rhesus macaques. Unfortunately, the interesting problem of possible joint occurrence of M. a. pelops and M. mulatta in the eastern Sun- darbans could not be studied during a recent primate survey of Bangladesh (Green, 1978, p. 157).

The irregular lobulated configuration of the range of M. a. assamensis in South- east Asia complicates the distributional relationship between this subspecies and M. mulatta (fig. 9). The range of M. a. assamensis follows the configuration of the north-south trending mountain chains in Southeast Asia, which in turn deter- mines the distribution of the mid-elevation broadleaf evergreen forest habitat of this species (fig. 1).

Table 10. Collection details and field observations concerning 11 local areas inhabited by Macaca assamensis assamensis (multiple specimens collected or observed), but apparently not inhabited by Macaca mulatta (fig. 9); for key to locality numbers, see figure 2.

1. Loc. Nos. 29-33 (10 km from nearest known M. mulatta locality) BURMA: upper Irrawady R., 900-1,800 m, 8 April 1931-7 Dec. 1938

Eight M. assamensis specimens collected; "only monkey we saw in the Adung Valley" (Kingdon Ward, 1937, p. 337). Specimen tag notes: "dense hill jungle" (BM 50.378-50.380); "dense jungle" (BM 50.381); "thick hill forest" (BM 50.383); "thick forest" (BM 50.384); "dense hill forest" (BM 50.386).

2. Loc. No. 34 (10 km from nearest known M. mulatta locality) INDIA: Nagaland, 1,400 m, 5 Sept. 1919-17 Dec. 1923

Six M. assamensis specimens collected; "common in . . .jungle. . . rarely approaches villages"; cf. M. mulatta (10 km distant), "swarms round villages" (Mills, 1923, p. 222). Also cf. M. mulatta (20 km distant), "inhabit the margin of the forest and the open plain, but do not enter the densely forested areas" (McCann, 1933, p. 807).

3. Loc. Nos. 36-38 (10 km from nearest known M. mulatta locality) BURMA: upper Chindwin R., 230-490 m, 21 Feb.-l March 1935

Four M. assamensis specimens collected: "heavy evergreen jungle . . . many species of Ficus and Dipterocarpus;" "Fine specimens of . . . Pterospermus and of Elaeocarpus also abounded. . . . Many and large . . . Upas trees (Atiaris toxicaria)" (Morris, 1936, pp. 657, 661); "tropical rain-forest" (Raven in Mayr, 1938, p. 279).

4. Loc. Nos. 41, 42 (30 km from nearest known M. mulatta locality) BURMA: middle Irrawady R., [ca. 200m], 21 Jan. 1865-3 March 1875

Two M. assamensis troops observed (one specimen collected); "The only wild monkey . . . observed;" "splendid mantle of dark green forest" (Anderson, 1871, p. 213; 1879, pp. 57, 65).

5. Loc. No. 45 (45 km from nearest known M. mulatta locality) CHINA: SW Yunnan, [ca. 500 m], 3-4 March 1917

Four M. assamensis specimens collected, evidently only macaque in area; evergreen forest, trees ca. 45 m tall (Andrews, 1918, p. 253).

6. Loc. No. 48 (55 km from nearest known M. mulatta locality) THAILAND, NW: 1,800 m, 6-12 March 1937

Two M. assamensis specimens collected; "tall tropical evergreen forest," "primary forest . . . [little] underbrush" (Coolidge, 1940, pp. 125, 128).

7. Loc. Nos. 52-55 (50 km from nearest known M. mulatta locality) THAILAND, west-central: ca. 250-750 m, 14-23 April 1967

Eight M. assamensis specimens collected; M. mulatta actively sought, evidently not present in area; semi-evergreen forest, teak, bamboo (Fooden, 1971a, p. 36).

8. Loc. No. 58 (5 km from nearest known M. mulatta locality) THAILAND, west-central: 580-800 m, 18 March 1974-6 March 1975

Two M. assamensis troops observed; one troop in "dry evergreen forest," one troop from "dry evergreen forest" into "bamboo" (Eudey, 1979, pp. 95, 96, table 7).

9. Loc. Nos. 59, 60 (70 km from nearest known M. mulatta locality) THAILAND, west-central: 600-1,100 m, 15-28 Jan. 1967

Six M. assamensis specimens collected; M. mulatta actively sought, evidently not present in area; evergreen forest (Fooden, 1971a, p. 36).

10. Loc. Nos. 64, 65 (30 km from nearest known M. mulatta locality) LAOS, N: [ca. 450-700 m], 21 June 1924-28 Nov. 1931

Two M. assamensis specimens collected; one taken "half way up the mountain" (Legendre, 1936, p. 158).

11. Loc. No. 67 (85 km from nearest known M. mulatta locality) VIETNAM, NW: 1,500 m, 4 Nov. 1929-2 Feb. 1939

Eight M. assamensis specimens collected; "forets . . . toujourverts" (Delacour, 1930, p. 457), "dense forest" (Lowe, 1947, p. 60).

28

FOODEN: MAC AC A ASSAMENSIS 29

c

M. a. assamensis has been collected or observed at approximately 45 Southeast Asian localities that range in altitude from about 150 to 2,750 m (fig. 9). In the same general area, M. mulatto has been collected or observed at approximately 115 localities that range in altitude from about 75 to 4,000 m. The greater alti- tudinal range of M. mulatto is obviously correlated with its broader geographic distribution. Despite the broader altitudinal and geographic distribution of M. mulatto, several local areas are inhabited by M. a. assamensis and not by M. mulatto (table 10). Predictably, these exclusive M a. assamensis areas all are in mid-elevation broadleaf evergreen forest. By contrast, at 10 known localities of joint or proximate occurrence of M. a. pelops and M. mulatta, the vegetation generally is mixed deciduous and evergreen forest (table 11), and there are no records of M. mulatta in the evergreen forest components at any of these local- ities. Available evidence thus indicates that the ecological relationship between M. a. assamensis and M. mulatta in Southeast Asia is essentially the same as between M. a. pelops and M. mulatta south of the Himalayas. The primary habitat of M. a. assamensis in Southeast Asia apparently is mid-elevation evergreen forest and the primary habitats of M. mulatta are low elevation deciduous forest or high elevation coniferous forest. At intermediate altitudes with intermediate rainfall, where low elevation deciduous forest intergrades with mid-elevation evergreen forest, the two species may occur close together or in direct contact.

A probable encounter between M. a. pelops and M. mulatta has been recorded in western Thailand near Khao Nang Rum by Eudey (1979, pp. 92, 97, 199), who selected this area for study because both macaques were known to inhabit the vicinity. The encounter was in mixed vegetation at an intermediate altitude (480-540 m). Difficult field conditions prevented close or complete observation of the encounter:

On 24 July 1977 [11:45 a.m.] a group of about six M. mulatta was contacted near the summit of Khao Nang Rum [540 m] in mixed deciduous-bamboo forest. ... At 11:55 this group of [rhesus] macaques probably encountered a large group [>20 individuals] of M. assamensis which subsequently came under observation [480 m]. Although the encounter was not observed visually, the squawking, squealing, and thudding of bamboo which accompanied it was recorded on tape but remains to be analyzed, [p. 92] Subsequent to the encounter, until 12:50, the group of M. assamensis remained in the trees in a ravine, keeping the [human] observers under surveillance, [p. 97]

These notes seem to indicate that the larger troop of M. a. assamensis displaced and dominated the smaller troop of M. mulatta in this encounter.

Table 11. Collection details and field observations concerning 10 localities where both Macaca assamensis assamensis and Macaca mulatta have been collected or observed (fig. 9); for key to locality numbers, see fig. 2.

1. Loc. No. 35; BURMA, upper Chindwin R.

M. assamensis (one specimen, 25 Aug. 1914, 140 m): "Forest partly evergreen and partly mixed jungle, but more scrubby than further North" (Shortridge in Wroughton, 1916a, p. 293).

M. mulatta (one specimen, 24 Aug. 1914, 140 m): see note above.

2. Loc. No. 39; BURMA, upper Irrawady R.

M. assamensis (one specimen, 7 Jan. 1939, 1,100 m): "edge of clearing in thick forest" (BM 50.385, tag).

M. mulatta (one specimen, 28 Jan. 1939, 1,100 m): "in yam field in forest" (BM 50.372, tag).

TABLE 11. Continued.

3. Loc. Nos. 43, 44; BURMA, middle Irrawady R.

M. assamensis (three specimens, Jan. 1875-Aug. 1886, altitude unknown): "Bhamo, near" (ZSI 11924, tag); "in the neighborhood of Bhamo a young male was brought to me" (Anderson, 1879, p. 68); "In the forest 2 miles N. of Bhamo" (Thomas, 1892, p. 916).

M. mulatto (one specimen, 15 Feb. 1936, 180 m): "Bhamo, 600 ft" (BM 36.12.26.4).

4. Loc. No. 46; CHINA, SW Yunnan

M. assamensis (no. specimens unknown, 1964, [ca. 1,000 m]); nematode parasites reported.

M. mulatta (no. specimens unknown, 1964, [ca. 1,000 m]); nematode parasites re- ported, same species as in M. assamensis (Yen Wen-chen, 1973, p. 362).

5. Loc. No. 48; THAILAND, NW.

M. assamensis (four specimens collected, 8-11 June 1937, 400-2,100 m): collected "in the Doi Dao region;" vegetation: "false teak, fruit bearing trees . . . bamboo . . . broken by clearings . . . many trees lost their leaves . . . islands of evergreen" (Car- penter, 1940, pp. 20-24), "first 1,800 feet [550 m] . . . dry deciduous forest of oak and bamboo . . . [550-1, 400m] pine and oak forests . . . occasional second growth . . . evergreen along streams . . . summit . . . open and grass covered" (Coolidge, 1940, p. 125).

M. mulatta (one specimen collected, 14 June 1937, 400-2,100 m); see notes above.

6. Loc. No. 51; THAILAND, west-central.

M. assamensis (three specimens, 25 March 1967, ca. 350 m): in teak tree, semi-ev- ergreen forest.

M. mulatta (one specimen, 21 March 1967, ca. 350 m): in bamboo (Fooden, 1971a, pp. 32, 36).

7. Loc. No. 56; THAILAND, west-central.

M. assamensis (two troops observed): one troop (13 Feb. 1974, 500 m), "feeding in the crown of Ficus sp. tree," "mixed deciduous;" one troop (22-24 July 1977, 460-480 m), "mixed deciduous-bamboo forest."

M. mulatta (three troops observed): one troop (3 Dec. 1973, 320 m), "feeding on the seeds of flowering bamboo, Bambusia sp.;" one troop (3 Dec. 1973, 340 m), "mixed deciduous forest;" one troop (24 July 1977, 540 m), "mixed deciduous-bamboo forest," probable encounter with M. assamensis troop (Eudey, 1979, pp. 91, 92, 94, 97, table 7).

8. Loc. No. 57; THAILAND, west-central.

M. assamensis (one troop observed, 23 Jan. 1974, 400 m): vegetation "mixed de- ciduous."

M. mulatta (one troop observed, 21 Dec. 1973, 500 m): "mixed deciduous" (Eudey, 1979, table 7).

9. Loc. No. 61; CHINA, SW Yunnan.

M. assamensis (no. troops unknown, Aug. -Sept. 1957, altitude unknown): encoun- tered infrequently; dense tropical forest, characteristic elements Ficus, magnolia, bam- boo, acacia, camphor, wild banana (Bannikov, 1958, p. 67).

M. mulatta (Feb. 1957-Aug. 1958, altitude unknown): troops observed (no. un- known), common (Bannikov, 1958, p. 67); four specimens collected, vegetation: broad- leaf evergreen forest to 1,000 m, Albizzia chinensis, Ficus spp., Gossampinus malabarica, Duabanga grandiflora, Dipterocarpus yunnanensis, Hopea millissima, Antiaris toxicaria, Po- metia tomentosa, Chukrasia tabularis, Canarium sp., Sinocalamus strictus (Kao Yueh-ting et al., 1962, pp. 181, 188).

10. Loc. No. 66; VIETNAM, NW.

M. assamensis (one specimen collected, 11 March 1929, 370 m): "Country diversified and much changed by man but some large forest left. The Thais and the Meos from the surrounding hills brought in a large number of the specimens preserved here" (Bangs & Van Tyne, 1931, p. 37).

M. mulatta (one specimen collected, 15 March 1929, 370 m): see note above.

30

FOODEN: MAC AC A ASSAMENS1S

31

Fig. 10. External characters of Macaca assamensis pelops (below) compared with those of M. mulatto (above) in hitherto unpublished drawing commissioned in Nepal by B. H. Hodgson and sent to British Museum in 1845. In the colored original of this drawing (BM), the grayish posterodorsal pelage in M. a. pelops is effectively contrasted with the reddish posterodorsal pelage in M. mulatto.

Macaca assamensis pelops Hodgson, 1841.

[Macacus {Pithex)] Pelops: Hodgson, 1841, p. 1213 — external characters compared with those of M. mulatto; habitat, Nepal, "Northern region of hills exclusively"; species name is derived from Greek mythology, name of son of Tantalus. Gray, 1846, p. 2 — possibly a synonym of M. assamensis.

M[acacus] Pelops: Schinz, 1844, p. 60 — characters and distribution ex Hodgson, 1841. Blyth, 1847, p. 313 — taxonomy.

l[nuus (Rhesus)] pelops: Wagner, 1855, p. 56 — doubtfully distinct from M. mulatto.

Pithecus pelops: Wroughton, 1916b, p. 475 — taxonomic comparisons.

Macaca pelops: Wroughton, 1916c, p. 776 — specimen recorded, Pashok. Wroughton, 1918, p. 554 — taxonomy.

Macaca assamensis pelops: Pocock, 1939, p. 55 — external and cranial characters; natural his- tory; taxonomy.

Macacus Oinops: Gray, 1843, p. 8 (not Hodgson, 1841, p. 1212) — misidentified specimen (cf. Gray, 1846, p. 2).

M[acacus] MacClellandii: Gray, 1846, p. 2 — nomen nudum, in synonymy, with citation "Horsf. Ogilby, Royle, H."; name does not appear in either Horsfield (1840, pp. 146-167) or Ogilby in Royle (1840, pp. lvi-lxxiv).

Macacus sikimensis: Hodgson in Jerdon, 1867, p. 12 — nomen nudum cited by Jerdon from unpublished manuscript of B. H. Hodgson.

M[acacus] sikkimensis: Anderson, [1879], p. 72 — modified spelling of Macacus sikimensis, Hodgson in Jerdon, 1867, nomen nudum.

32 FIELDIANA: ZOOLOGY

Macacus assamensis: Sclater, [1869], p. 566 — captive collected at Dalamcote Fort ( = Daling), presented by Major C. Richards to Menagerie of Zoological Society of London; taxonomic comparisons.

Macaca assamensis: Wroughton, 1918, p. 555 — part, type-specimen of Macacus problematicus, Gray, 1870, listed.

Macacus problematicus: Gray, 1870, p. 128 — holotype, stuffed skin with skeleton, BM 69.3.5.15, juvenile female, captured alive December 1863 at Dalamcote Fort ( = Daling), northern West Bengal, presented to Menagerie of Zoological Society of London by Major C. Richards, 9 November 1868; died March 1869. Sclater, 1871, p. 222— a synonym of M. assamensis. Wroughton, 1916b, p. 475 — a synonym of Pithecus pelops.

Supposed new monkey from the Sunderbunds: Anderson, 1872, p. 529 — compared with M. mulatta; measurements of dry skin, head, and body 22 inches [559 mm], tail 12 Vz inches [318 mm; relative tail length 0.57]; not named. Sclater in Anderson, 1872, p. 532, footnote — probably M. assamensis.

M[acacus] rhesus: Anderson, 1879, p. 62, figs. 5-6 (skull) (part, figure caption; not Audebert, 1797-99, p. 5)— misidentified skull, BM 45.1.8.4 8, collected in "Nepal" by B. H. Hodg- son (cf. Hinton & Wroughton, 1921, p. 667). Anderson, 1881, p. 67 (part of synonymy and Specimen No. 41e)— characterization of specimen collected at Sundarbans. Blanford, 1888, p. 12, figs. 3-4 (skull) — illustration and erroneous caption copied from Anderson, 1879, p. 62.

Type

Juvenile male, stuffed skin with damaged skull (BM 43.1.12.4, Coll. No. 14), collected by B. H. Hodgson (1841, p. 1213) sometime during the period 1834-40 (cf. Hodgson, 1832, p. 339; 1834, p. 95). External measurements given by Hodg- son (1841, p. 1213) for Macacus pelops are: "Twenty inches [508 mm] long. Tail less hair, nine and a half [241 mm]." External measurements of the dry, juvenile type-specimen are: head and body 470 mm, relative tail length 0.66.

The type-specimen was received and catalogued at the British Museum (Nat- ural History) early in 1843 (Gray, 1843, pp. xv, 8; Thomas, 1906, p. 6). In 1845 the British Museum received from Hodgson another Nepalese specimen of M. a. pelops (BM 45.1.8.225, Coll. No. 54, adult female) and a colored drawing (fig. 10) of a living or recently dead specimen (Gray, 1846, p. 2). In addition, at least four more specimens of M. a. pelops also were sent by Hodgson to the British Museum (Natural History) (BM 45.1.8.4 8, skull only; BM 58.6.24.67 6; BM 79.11.21.302 8; BM 79.11.21.303 9) (cf. Gray, 1863, pp. vi, ix, 1; Scully, 1888, p. 234).

Type-Locality Nepal: "Northern region of hills exclusively" (Hodgson, 1841, p. 1213).

Distribution (Fig. 1)

Lower and middle ranges of Himalayas (300-2,750 m) from central Nepal; eastward through Sikkim (India); adjacent northern West Bengal (India); Bhutan; and, probably, Arunachal Pradesh (India) as far east as the great bend of the Brahmaputra River. One widely disjunct record is near sea level in the Sundar- bans coastal swamps, southwestern Bangladesh. The known western and east- ern limits of distribution of this subspecies, both based on reliable sight records, are Tipling, Nepal (83°36'E), and Manas River, Bhutan (90°58'E). Although M. a. pelops has not been recorded east of south-central Bhutan, suitable habitats unbroken by intervening barriers apparently exist in adjacent Arunachal Pradesh

FOODEN: MAC AC A ASS AMEN SIS 33

(Schweinfurth, 1957, Vegetationskarte, Beilage); accordingly, as previously sug- gested by Hill (1974, p. 735), it seems likely that the subspecific range extends eastward as far as the Brahmaputra River (95°E), which is a major zoogeographic boundary (cf. Hylobates; Fooden, 1969, fig. 3). The widely disjunct Sundarbans record is based on a reasonably well documented museum specimen (see above, p. 2) and, therefore, in the absence of contradictory evidence, it must be accepted as authentic; presumably, the Sundarbans population of M. a. pelops is or was a zoogeographic relict.

(Note Added in Proof: For probable new records of M. a. pelops in north central Nepal [ca. 28°15'N, 85°30'E], see M. J. B. Green, 1981, J. Bombay Nat. Hist. Soc., 78: 80.)

Diagnostic Tail Measurements and Proportions

Tail length 236-293 mm (265 ± 25 mm) in five adult females, 283-382 mm (330 ± 32 mm) in eight adult males; relative tail length (T/HB) .44-.55 (.48 ± .05) in five adult females, .50-.69 (.56 ± .05) in eight adult males (table 2, fig. 4).

Specimens Examined

Total 35.

Skins and skulls, 23— Localities Nos. 7 (1), 8 (1), 9 (3), 11 (2), 13 (1), 14 (1), 16 (4), 19 (2), 20 (4), 21 (1), 22 (1), "Nepal" (2:BM). Skins only, 10— Localities Nos. 2 (1), 6 (2), 18 (1), 25 (1), "Gangtok" (1:ZMB), "Lhasa" (1:ZSBS), "Nepal" (3, incl. 2 with skulls inside: BM). Skulls only, 2— Locality No. 16 (1), "Nepal" (1:BM).

For further details, see Figure 1 and Gazetteer.

Macaca assamensis assamensis McClelland in Horsfield, [1840].

Macacus Assamensis: McClelland in Horsfield, 1839 [March 1840], p. 148 — characterization in Latin and English; no exact locality. Horsfield, 1851, p. 21 — type-specimen listed. Anderson, 1879, p. 64 — type-specimen characterized. Blanford, 1888, p. 15 — type-spec- imen "from Assam, possibly from the hills to the northward."

Pttpio Assamensis: Ogilby in Royle, 1839 [1840], p. Ix — taxonomic relationships; probable distribution, "more eastern hills."

/. [lnuus (Rhesus)] assamensis: Wagner, 1855, p. 57 — taxonomy.

Pithecus assamensis: Elliot, [1913], p. 209 — type-locality, "Assam"; type history.

Macaca assamensis: Wroughton, 1918, p. 553 (part) — modified spelling of generic name. Hinton & Wroughton, 1921, p. 669— taxonomic history. Allen, 1938, p. 282 — type-lo- cality, "Assam, probably in the Garo Hills region."

Macaca assamensis assamensis: Pocock, 1939, p. 53 — distribution; external characters.

Rhesus assamensis: Furuya, 1962, p. 377 — dermatoglyphics.

l[nuus) rhesus: Blyth, 1865, p. 192 (not Audebert, 1797-99, p. 5)— type-specimen of M. assamensis probably "merely an individual variety of colour of the common animal of Lower Bengal."

Macacus rheso-similis: Sclater, 1872, p. 495, pi. 25 (animal) — "temporary designation"; ho- lotype, stuffed skin without skull, BM 76.1.31.23, immature male (originally recorded as female), living captive acquired in Calcutta market by W Jamrach, November 1871, purchased by Menagerie of Zoological Society of London 15 February 1872, died 9 December 1872, skin and skull transferred to British Museum May-June 1875; habitat given on specimen tag, "E. Indies"; dry skin measurements, head and body 450 mm, relative tail length 0.45. Sclater in Anderson, 1872, p. 532, footnote — probably a synonym of M. assamensis. Sclater, 1875, p. 418 — type history. Sclater, 1883, p. 21 — type-specimen listed.

Macaca assamensis coolidgei: Osgood, 1932, p. 209 — holotype, flat skin with skull, BM 32.4.19.1, adult male, collected by J. Delacour & W Lowe (No. 1824), 19 January 1930, Hoi Xuan, Annam, northern Vietnam; 11 specimens in type-series. Pocock, 1939, pp. 53-55 — a synonym of M. a. assamensis. Allen & Coolidge, 1940, p. 146 — six specimens recorded, Doi Inthanon, Doi Luang Chiang Dao; taxonomic comparison, possibly an undescribed subspecies.

34 FIELDIANA: ZOOLOGY

Macaca nemestrina: Dollman, 1932, p. 9 (not Linnaeus, 1766, p. 35) — misidentification of BM 32.11.1.2 <?, Adung Valley. Kingdon Ward, 1937, p. 337— same misidentification. Kaulback, 1938, p. 313 — same misidentification.

Type

Adult male, stuffed skin with skull, collected by J. McClelland, September 1835-February 1837 (McClelland in Horsfield, 1840, pp. 147, 148; Griffith, 1847, pp. ii, 1-59). The type-specimen formerly was preserved in the Zoological Mu- seum of the East India Company, London, where it was examined and char- acterized by Horsfield (1851, p. 64), who noted that the skull was then missing. Subsequently, Gray (1870, p. 31) was unable to locate the specimen, and Sclater (1871, p. 222) inquired concerning its whereabouts and was informed that it was "boxed up in the cellars of the new Indian Office." In 1879 the collections of the Zoological Museum of the East India Company were transferred to the British Museum (Natural History); the transferred collections included 695 mammal specimens, among which should have been McClelland's type-specimen (Thomas, 1906, p. 40). However, there is no evidence that this type-specimen was received by the British Museum, and it has not been seen since sometime before 1888 (Blanford, 1888, p. 16; Elliot, 1913, p. 211).

External measurements of the dry skin were recorded by Anderson (1879, p. 65), as follows: "The length of the animal along the curve of the head and back is 26.75 inches [679 mm], the tail measuring 9Vi inches [235 mm; relative tail length 0.35]."

Type-Locality

Not precisely known; originally given as "Assam" (Horsfield, 1840, p. 147), which at that time designated practically all of northeastern India. The type- specimen was collected during the course of a 17-month expedition to explore tea-producing areas in northeastern India. The route of this expedition covered approximately 1,000 km from near the mouth of the Ganges River (24°N, 90°E), upward along the Brahmaputra River and its affluents, as far as the Mishmi Hills (28°N, 97°E) (Griffith, 1847, pp. ii, 1-59; 1848, fold-out map). No member of the expedition party seems to have recorded the exact or approximate point along this route where the type-specimen of M. assamensis was obtained.

Blanford (1888, p. 15) speculated that the type-specimen may have been col- lected in "the hills to the northward" of Assam — that is, in the Mishmi Hills — which is possible but not certain. Hinton & Wroughton (1921, p. 667) implied that the type-specimen was collected in or near the Garo Hills (25°30'N, 90°30'E), but there is no known evidence that M. assamensis inhabits this vicinity (see above, p. 3).

Distribution (Fig. 2)

Lower and middle ranges of mountain chains in continental Southeast Asia (200-2,750 m), from great bend of Brahmaputra River in southeastern Xizang (China) and northeastern India (eastern Arunachal Pradesh, Nagaland), south- ward and eastward through northern and eastern Burma, southwestern Yunnan (China), northern and western Thailand, Laos, northern Vietnam, to south- western Guangxi (China). The northern limit of distribution of M. a. assamensis

FOODEN: MACACA ASSAMENSIS 35

is similar to that of the genus Hylobates (Fooden, 1969, fig. 3) and of the Tra- chypithecus group in the genus Presbytis (Fooden, 1975b, p. 108).

Diagnostic Tail Measurements and Proportions

Tail length 193-225 mm (208 ± 10 mm) in eight adult females, 160-240 mm (209 ± 23 mm) in 13 adult males; relative tail length (T/HB) .39-.47 (.43 ± .03) in eight adult females, .26-.44 (.35 ± .05) in 12 adult males (table 2, fig. 4).

Specimens Examined

Total 77.

Skins and skulls, 65— Localities Nos. 28 (2), 29 (1), 30 (1), 31 (4), 32 (1), 33 (1), 34 (5), 35 (1), 36 (1), 37 (1), 38 (2), 39 (1), 42 (1), 44 (1), 45 (2), 48 (3), 49 (1), 50 (2), 51 (3), 52 (4), 53 (3), 54 (1), 55 (1), 59 (4), 60 (2), 62 (1), 63 (1), 64 (1), 65 (1), 66 (1), 67 (6), 69 (1), 70 (1); "Laos Mts." (1:BM); no locality data (2:AMNH, ZSI). Skins only, 10— Localities Nos. 34 (2), 40 (1), 48 (1), 67 (2, ind. 1 with skull inside), "Mishmi Hills," (1:ZSI), "E. Indies" (1:BM), no locality data (2:BNHS, ZSI). Skulls only, 2— Locality No. 45.

For further details, see Figure 2 and Gazetteer.

GAZETTEER OF MACACA ASSAMENSIS LOCALITIES

The form and spelling of primary entries in this gazetteer follow, where pos- sible, those in U.S. Board on Geographic Names (USBGN) gazetteers (Bhutan, India, Nepal, 1952; Vietnam, 1964; Burma, 1966; Thailand, 1966; Laos, 1973; China, 1979). Primary entries for macaque localities that are not listed in USBGN gazetteers are spelled here either as in the original source or as in an indicated reference. Secondary entries, with cross references to corresponding primary entries, give variant spellings or alternate locality names that appear on specimen tags or in the published literature on Macaca assamensis.

The sequence of information presented in each primary entry is:

1. locality name

2. altitude, if available

3. name of country (capital letters) and state, province or other first-order administrative unit (capital and lower case letters)

4. coordinates of locality, usually taken from USBGN gazetteer

5. name of collector or observer followed by parenthetical reference to pub- lished locality notes, if any

6. date of collection or observation

7. abbreviated name of museum where specimens are preserved and number of specimens available (with indication of part preserved, if skin and skull are not both present) or bibliographic reference to literature record

8. italicized locality number as indicated in distribution maps (figs. 1, 2).

Macaca assamensis pelops (fig. 1)

Abbottabad vicinity; PAKISTAN; Peshawar; 34°09'N, 73°13'E; erroneous record (Blanford, 1898, p. 362), apparently based on observation of misidentified M. mulatta.

Arun Valley, 1,000-10,000 ft. [300-3,000 m]; NEPAL; East No. 4; ca. 27°10'N, 87°10'E; reported by J. A. McNeely (letter, 29 July 1973). Not mapped.

"Assam." See Daling.

36 FIELDIANA: ZOOLOGY

Batasia, 4,000 ft. [900 m]; INDIA; West Bengal; ca. 27°00'N, 88°08'E; collected by C. A. Crump (in Wroughton, 1916b, p. 472), 28 Feb. 1915 (BM, 1). 13

"Bhootan." See Daling.

Birch Hill; INDIA; West Bengal; 27°04'N, 88°17'E; observed by C. H. Southwick et al. (1964, p. 444), Oct.-Nov. 1962. 17

Chuntang, 5,350 ft. [1,630 m]; INDIA; Sikkim; 27°38'N, 88°38'E; collected by C. A. Crump (in Wroughton, 1916b, p. 470), 16 Dec. 1914 (BM, 1). 7

Dalamcote Fort. See Daling.

Daling; INDIA; West Bengal; 27°01'N, 88°42'E; captured alive Dec. 1863; pre- sented by C. Richards, 9 Nov. 1868, to Menagerie of Zoological Society of London (Sclater, 1869, p. 566; 1871, p. 222); died Mar. 1869 (BM, 1). For locality notes, see Eden (1865, pp. 59-66) and Rennie (1866, pp. 178, 342). 22

Darjeeling, near, 4,000-5,000 ft. [1,200-1,500 m]; INDIA; West Bengal; ca. 27°02'N, 88°16'E; observed by T. C. Jerdon (1867, p. 12), date unknown. 27

Darling. See Daling.

Dhalimkote. See Daling.

Dhaling. See Daling.

Dikchu, 2,000 ft. [600 m]; INDIA; Sikkim; 27°25'N, 88°35'E; collected by C. A. Crump (in Wroughton, 1916b, p. 470), 25 Jan. 1915 (FMNH, 1). 8

"Gangtok"; INDIA; Sikkim; 27°20'N, 88°30'E; skin purchased by E. Schafer, date

unknown (ZMB, 1 [skin only]). Not mapped. Gopaldhara, 5,200 ft. [1,575 mj; INDIA; West Bengal; 26°55'N, 88°12'E; (Moore

& Tate, 1965, p. 320); collected by H. Stevens, 2 June 1923 (BM, 1). 14

Iswa Khola, 9,000 ft. [2,700 m]; NEPAL; Dhankuta; 27°37'N, 87°08'E; observed by J. A. McNeely (letter, 4 Feb. 1973), date unknown. 3

Jalpaiguri district; INDIA; West Bengal; ca. 26°30'N, 88°45'E; imprecise locality, captive reported to have been caught locally (Inglis et al., 1919, p. 822). Not mapped.

Jiri; NEPAL; East No. 2; 27°38'N, 86°14'E; collected by R. L. Fleming (in Fleming & Traylor, 1964, p. 502), 10 Nov. 1960 (FMNH, 1 [skin only]). 2

Kali Gandak River. See Tipling.

Kashmir. See Lolab.

Kumaon (Division); INDIA; Uttar Pradesh; ca. 30°N, 80°E (Wroughton, 1914, p.

282); probably erroneous record (Hutton, 1865, p. xiii), apparently based on

misidentified M. mulatta (see above, p. 2). Kurseong, near [N], 4,000 ft. [1,200 m]; INDIA; West Bengal; ca. 26°53'N, 88°17'E;

observed by C. H. Southwick et al. (1964, p. 444), Oct.-Nov. 1962. 15

Lachen, 8,900 ft. [2,700 m]; INDIA; Sikkim; 27°44'N, 88°33'E; observed by J. D. Hooker (1854, p. 37), 27 May 1849. 5

Lamteng. See Lachen.

"Lhasa"; CHINA; Xizang ( = Tibet); 29°42'N, 91°06'E; captive (presumably arti- ficially transported), purchased by E. Schafer, 1938-39, died in Cologne Zoo, Dec. 1943 (ZSBS). Not mapped.

FOODEN: MACACA ASSAMENS1S 37

Lingtam. See Singtam (Moore & Tate, 1965, p. 325).

Lolab; INDIA; Jammu and Kashmir; ca. 34°30'N, 74°20'E; erroneous record (Blan- ford, 1898, p. 361), based on M. mulatta (USNM 20120/35485-20124/35489).

Manas River, west bank; BHUTAN; Kenga Dzong; 26°47'N, 90°58'E; observed

by R. P. Mukherjee (1978b, pp. 739, 742), Jan. 1976. 24 Manshitang, [6,000 ft. = 1,800 m]; INDIA; Sikkim, ca. 27°42'N, 88°34'E; collected

by E. Schafer (1943, p. 131), 10 Nov. 1938 (ZMB, 2 [skins only]). For notes on

locality see Pohle (1944, p. 185) and Ali (1962, p. xxviii). 6 Masuri. See Mussooree. Murree vicinity; PAKISTAN; Rawalpindi; 33°54'N, 73°24'E; probably erroneous

record (Blanford, 1898, p. 362), apparently based on misidentified M. mulatta

(see above, p. 2). Mussooree ( = Mussoorie); INDIA; Uttar Pradesh; 39°27'N, 78°05'E; probably

erroneous record (Hutton, 1865, p. xiii), apparently based on misidentified M.

mulatta (see above, p. 2).

Nepal, "Northern region of hills exclusively"; NEPAL; imprecise locality; col- lected by B. H. Hodgson (1832, p. 335; 1841, p. 1213), 1832-41 (BM, 6 [two skulls in skins, one skull only]). Not mapped.

Palmajua, 2,250 m; INDIA; West Bengal; 27°04'N, 88°06'E (US Defense Mapping

Agency, 1:253,440; Sheet G-45E); tentative sight record, observed at distance

by H. Khajuria (1966, p. 284), June-Aug. 1958. 12 Pashok, 3,000 ft. [900 m], 3,500 ft. [1,050 m]; INDIA; Sikkim; 27°03'N, 88°25'E;

coUected by C. A. Crump (in Wroughton, 1916b, p. 472), 3 Feb. 1915 (ZSI, 1);

N. A. Baptista (in Wroughton, 1916c, p. 776), 4 July 1915 (BM, 1). 19

Rhenok; INDIA; Sikkim; 27°11'N, 88°39'E (US Defense Mapping Agency,

1:1,000,000, Sheet N.G.-45); observed by W T. Blanford (1871, p. 375), 18

Aug. 1870. 10 Rhinok. See Rhenok. Rongli, 2,700 ft. [800 m]; INDIA; Sikkim; 27°13'N, 88°44'E; collected by C. A.

Crump (in Wroughton, 1916b, p. 470), 21 Nov. 1914, 21 Apr. 1915 (BNHS, 2

[one skull only; skin in BM]). 11

Sabaya Khola; NEPAL; Dhankuta; ca. 27°20'N, 87°15'E; observed by J. A.

McNeely (letter, 29 July 1973), 3 June 1973. 4 Sangsir. See Singsir.

Shamgong; BHUTAN; Shamgong Dzong; 27°14'N, 90°39'E; observed by Zoo- logical Survey of India Party, 1966-69 (Chakraborty, 1975, p. 18). 23 Simla; INDIA; Himachal Pradesh; 31°06'N, 77°10'E; probably erroneous record

(Hutton, 1865, p. xiii), apparently based on misidentified M. mulatta (see

above, p. 3). Singsir, 1,000 ft. [300 m]; INDIA; West Bengal; 27°04'N, 88°35'E (Moore & Tate,

1965, p. 332); collected by H. Stevens, 7 Jan. 1931 (FMNH, 1). For collector's

itinerary, see Sanborn, 1932, p. 181. 21 Singtam, ca. 4,750 ft. [1,450 m], 5,000 ft. [1,500 m], 5,400 ft. [1,650 m]; INDIA;

Sikkim; 27°14'N, 88°33'E (Stevens, 1923, map between pp. 502-3; Moore &

38 FIELDIANA: ZOOLOGY

Tate, 1965, p. 332); collected by H. Stevens, 28 Jan., 5 and 24 Feb. 1931 (FMNH,

3). 9 Sookia Pokhari, 5,000 ft. [1,500 m]; INDIA; West Bengal; 27°02'N, 88°14'E (Moore

& Tate, 1965, p. 332); collected by C. A. Crump (in Wroughton, 1916b, p.

472), 18, 22 Feb. 1915 (BNHS, 2). 16 Sukiapokhri. See Sookia Pokhari. Sukiapokri. See Sookia Pokhari. Sundarbans, ca. 50 mi. [80 km] E of Calcutta [near sea level]; BANGLADESH;

Khulna; ca. 22°35'N, 89°15'E; collected by museum collector (Anderson, 1872,

p. 529), 26 Apr. 1870 (ZSI, 1 [skin only]; collection originally included two

skins and at least one skull, according to Anderson, loc. cit.). 25 Sunderbunds. See Sundarbans.

Takdah, 1,650 m; INDIA; West Bengal; 27°02'N, 88°22'E; collected by ZSI party, Harvard- Yale Exped., 7 Aug. 1958 (ZSI, 1 [skin only]). For locality note, see Khajuria & Ghose, 1970, p. 17. 18

Tarkhola, 325 m, 1,400 ft. [430 m]; INDIA; West Bengal; 27°07'N, 88°33'E (Stevens, 1923, map between pp. 502-3; Moore & Tate, 1965, p. 333); collected by H. Stevens, 15 Jan. 1931 (USNM, 1); collected by H. Khajuria (1966, p. 284; Khajuria & Ghose, 1970, p. 17) and R. L. Fernandez, 17-22 Aug. 1958 (ZSI, 3 [1 skin only]). 20

Tipling, Kali Gandak River, left bank, ca. 4,000 ft. [1,200 m]; NEPAL; Baglung; ca. 28°27'N, 83°36'E; observed by R. L. Fleming, Nov. 1949 (personal com- munication). For itinerary notes, see Rand & Fleming, 1957, pp. 10, 15. 1

Tonglu. See Batasia.

Macaca assamensis assamensis (fig. 2)

Abia. See Tukreshwari Temple.

Adung Long, 5,000 ft. [1,500 m]; BURMA; Kachin, State; "28°04'N, 97°40'E"; collected by R. Kaulback, 31 Oct. 1938 (BM, 1). 30

Adung Valley, 6,000 ft. [1,800 m]; BURMA; Kachin State; 28°10'N, 97°40'E; col- lected by Lord Cranbrook (in Kinnear, 1934, p. 348; Kingdon Ward, 1937, p. 337), 8 Apr. 1913 (BM, 1). 29

Agia. See Tukreshwari Temple.

Ale Chaung. See "Atechg."

Ali Cha. See "Atechg."

Angka, Mt. See Inthanon, Doi.

Ao Naga tribal area; INDIA; Nagaland; 26°12'-26°45'N, 94°18'-94°50'E; reported as agricultural pest (Smith, 1925, p. 5). Not mapped (but see Mokokchung, which is in this tribal area).

"Assam"; INDIA; state uncertain; 24°-28°N, 90°-97°E; collected by J. McClelland (in Horsfield, 1840, p. 148; Griffith, 1847, pp. ii, 1-59), Sep. 1835-Feb. 1837; specimen formerly in Zoological Museum of East India Company, London (Horsfield, 1851, p. 21), subsequently lost (Blanford, 1888, p. 16). Not mapped.

"Atechg." ( = Ale Chaung), 1,000 ft. [300 m]; BURMA; Chin Hills Special Divi- sion; ca. 23°37'N, 94°22'E; erroneous record (Wroughton, 1916c, p. 763), based on misidentified M. mulatto (BM 31.1.11.22 6).

FOODEN: MAC AC A ASSAMENSIS 39

Ban Mae Lamao, 350 m; THAILAND; Tak; 16°48'N, 98°45'E; collected by J.

Fooden (1971a, p. 18), 25 Mar. 1967 (CTNRC, 1; FMNH, 2). 51 Ban Muang Baw Ngam, 1,100 m; THAILAND; Kanchanaburi; 14°55'N, 98°55'E;

collected by J. Fooden (1971a, p. 14), 15-19 Jan. 1967 (FMNH, 4). 59 Ban Pong Nam Ron, 5 km W, 250 m; THAILAND; Kamphaeng Phet; 16°20'N,

99°15'E; collected by J. Fooden (1971a, p. 19), 20 Apr. 1967 (FMNH, 1). 54 Ban Pong Nam Ron, 8 km W, 250 m; THAILAND; Kamphaeng Phet; 16°20'N,

99°13'E; collected by J. Fooden (1971a, p. 19), 23 Apr. 1967 (FMNH, 3). 53 Ban Pong Nam Ron, 25 km W, 750 m; THAILAND; Kamphaeng Phet; 16°20'N,

99°04'E; collected by J. Fooden (1971a, p. 19), 14 Apr. 1967 (FMNH, 4). 52 Bhamo, near; BURMA; Kachin State; ca. 24°16'N, 97°14'E; obtained alive by J.

Anderson (1881, p. 71) in 1875, died in Zoological Gardens, Calcutta, 19 Jan.

1881 (ZSI, 1); another specimen presented by J. Anderson (1879, p. 68) to

Regent's Park Zoo, London, apparently not preserved. 44 Bolovens Plateau. See Muang Thateng. Bomi, 2,750 m; CHINA; Xizang ( = Tibet); 29°50'N, 95°45'E; collected by Feng

Zuo-jian et al. (1980, p. 97; letter 15 Oct. 1980), May-Sep. 1973 (Institute of

Zoology, Beijing, 2, not seen). 26

Chang Tai, Huai, 400 m; THAILAND; Uthai Thani; 15°27'N, 99°15'E; observed

by A. A. Eudey (1979, pp. 94, 197, 208), 23 Jan. 1974. 57 Chapa, 5,000 ft. [1,500 m]; VIETNAM; Lao Cai; 22°21'N, 103°50'E; collected by

J. Delacour (1930, p. 457) and W. P. Lowe (1947, p. 60) 6 Nov.-14 Dec. 1929

(BM, 2 [one skin only]; FMNH, 4; MNHN, 1 [skull in skin]); collected by B.

Bjorkegren, 2 Feb. 1939 (MCZ, 1). For locality note, see Osgood, 1932, p. 197.

67 Chieng Dao. See Luang Chiang Dao, Doi. Chongklong. See Chongkrong. Chongkrong, 600-900 m; THAILAND; Kanchanaburi; 14°41'N, 98°52'E; collected

by J. Fooden (1971a, p. 14), 27-28 Jan. 1967 (CTNRC, 1; FMNH, 1). 60

Dagung Hka. See Jantang-Dagung Hka.

Damda; INDIA; Arunachal Pradesh; ca. 28°16'N, 94°55'E; fur haversack, possibly

made of skin of M. a. assamensis, collected by S. Kemp (1912, p. 2, pi. 1), 23

Jan.-3 Feb. 1912 (Robinson, 1913, p. 85). Not mapped. Debuk Damda. See Damda. Dibru Reserved Forest; INDIA; Assam; ca. 27°40'N, 95°15'E; erroneous report

(Pilleri, 1975, pp. 20, 43) based on observation of M. mulatto (see above, p.

3). Dimapur road. See Imphal Valley. Dinapur road. See Imphal Valley. Doi Inthanon. See Inthanon, Doi.

Garo Hills; INDIA; Meghalaya; ca. 25°30'N, 90°30'E; erroneous report (Hinton & Wroughton, 1921, p. 667), probably a lapsus for Naga Hills (see above, p. 3).

Gauhati. See Tukreshwari Temple.

Gengma; CHINA; Yunnan; 23°31'N, 99°24'E; collected by Institute of Zoology,

40 FIELDIANA: ZOOLOGY

Academia Sinica (Yen Wen-chen, 1973, p. 362), date unknown (presumably

in collection of Institute of Zoology, Beijing). 46 Goalpara. See Tukreshwari Temple. Goletu, 3,000 ft. [900 m]; BURMA; Kachin State; "27°37'N, 97°54'E"; collected

by R. Kaulback, 7 Dec. 1938 (BM, 1). 32 Guangxi [province], 1,000-2,000 m; CHINA; Guangxi; 21°30'-23°30'N,

105°25'-108°50'E; reported by Wang Sung et al. (1962, p. 556) and Shou Chen-

huang et al. (1964, p. 62). Not mapped.

Haibum, West bank [1,600 ft = 490 m]; BURMA; Sagaing Division; 26°02'N,

95°47'E; collected by H. C. Raven (in Mayr, 1938, p. 278; Morris, 1936, p. 660),

27 Feb. 1935 (AMNH, 1). 37 Hisweht, 460 ft. [140 m]; BURMA; Sagaing Division; ca. 25°22'N, 95°16'E; col- lected by G. C. Shortridge & S. A. MacMillan (in Wroughton, 1916a, p. 293),

24 Aug. 1914 (BM [skin], BNHS [skull], 1). 35 Hkamti ( = Singkaling Hkamti); BURMA; Sagaing Division; 26°00'N, 95°42'E;

erroneous record (Wroughton, 1916a, p. 296), based on M. mulatta (BM

15.5.5.5, BM 31.1.11.25, BNHS 5093, ZSI 12088). H. Me Kom. See Puan, Nam Mae. Hoi Xuan; Vietnam; Thanh Hoa; 20°22'N, 105WE; collected by J. Delacour

(1930, p. 459) and W P. Lowe (1947, p. 66), 19 Jan. 1930 (BM, 1). 69 Homalin; BURMA; Sagaing Division; 24°52'N, 94°55'E; erroneous record

(Wroughton, 1916a, p. 296), based onM. mulatta (BM 15.5.5.3-4, BNHS 5094). Htingnan Triangle, 3,500 ft. [1,050 m]; BURMA; Kachin State; "26°36'N, 97°52'E";

collected by R. Kaulback, 7 Jan. 1939 (BM, 1). 39 Huay Chang Tai. See Chang Tai, Huai. Huay Kha Kaeng Game Sanctuary. See Chang Tai, Huai; Khao Nang Rum, NW;

and Khao Nang Rum, ranger station, vie.

Imphal Valley, ca. 3,000 ft. [900 m]; INDIA; Manipur; ca. 24°52'N, 93°57'E; er- roneous record (Roonwall, 1949, p. 83), based on misidentified M. mulatta (ZSI 11187 6).

Inthanon, Doi, 6,000 ft. [1,800 m]; THAILAND; Chiang Mai; 18°32'N, 98°32'E; collected by H. J. Coolidge, Jr. (1940, p. 128), 6, 12 Mar. 1937 (MCZ, 2). 50

Irrawady River, 2nd defile, 20-25 mi below Bhamo, right bank; BURMA; Kachin State; ca. 24°08'N, 97°00'E; collected by J. Anderson (1879, p. 65), 3 Mar. 1875 (ZSI, 1). 42

Irrawady River, some miles below 2nd defile, right bank; BURMA; Kachin State; ca. 24°12'N, 96°49'E; observed by J. Anderson (1879, p. 65), 1875. 41

Jaddo. See Yado.

Jantang-Dagung Hka [750 ft. = 225 m]; BURMA; Sagaing Division; ca. 26°08'N,

95°57'E; collected by H. C. Raven (in Mayr, 1938, p. 278; Morris, 1935, p. 657),

21 Feb. 1935 (AMNH, 2). 38

Kengma. See Gengma.

Khao Nang Rum, NW, 460-480 m, 500 m; THAILAND; Uthai Thani; 15°30'N,

99°16'-99°18'E; observed by A. A. Eudey (1979, pp. 94, 97, 197-99, 208), 13

Feb. 1974, 22-24 July 1977. 56

FOODEN: MACACA ASSAMENSIS 41

Khao Nang Rum, ranger station, vie, 580-800 m, 600-660 m; THAILAND; Uthai Thani; 15°28'N, 99°18'-99°19'E; observed by A. A. Eudey (1979, pp. 95-96, 197-98, 208), 18-19 Mar. 1974, 6 Mar. 1975. 58

Kin. See Yin.

Ko Keow, ca. 200 m; THAILAND; Kamphaeng Phet; ca. 15°55'N, 99°25'E; col- lected by J. Fooden (1971a, p. 36), 8 Mar. 1967 (FMNH, 1). 55

Kom. See Puan, Nam Mae.

Kwangsi. See Guangxi.

Laiterai, Khasi Hills; INDIA; Meghalaya; not precisely located, ca. 25°30'N, 91°30'E; questionable record (see above, p. 4), based on isolated skull (BM 1937.3.24.16 8, "M. speciosa"), collected by H. W. Wells, date unknown. Not mapped.

Laitura. See Laiterai.

Lakon Lampang. See Puan, Nam Mae.

Lancang Jiang; CHINA; Yunnan; ca. 22°00'N, 100°50'E; observed by A. G. Ban- nikov (1958, p. 68), Aug-Sept. 1957. 61

"Lao Mounts"; THAILAND or LAOS; 15°-20°N, 101°-102°E; collected by H. Mouhot (1864, pp. 119-160), 10 Apr.-19 Oct. 1861 (probably before 27 July 1861; Mouhot, 1864, pp. 269-71, 294) (BM, 1). Not mapped.

Lhota Naga tribal area; INDIA; Nagaland; 26o00'-26°30:N, 94°00'-94°25'E; ques- tionable report (Mills, 1922, p. 67), hunted with crossbow, possibly based on misidentified M. mulatta. Not mapped.

Longzhou Xian; CHINA; Guangxi; ca. 22°25'N, 106°50'E; acquired by Long Zupei et al. (1979, p. 320), Sep. 1975. 68

Lo Tiao, 1,500 m; LAOS; Houakhong; 20°20'N, 100°25'E; collected by J. C. Green- way (in Delacour & Greenway, 1940, p. 19), 5 Jan. 1939 (MCZ, 1). For altitude, see Delacour, 1940, p. 21. 62

Luang Chiang Dao, Doi; THAILAND; Chiang Mai; 19°23'N, 98°54'E; collected by C. R. Carpenter (1940, p. 20), 8-11 June 1937 (MCZ, 4 [one skin only]). For additional locality note, see Coolidge, 1940, p. 129. 48

Lul worth Cove. See Imphal Valley.

Mahtum, 4,000 ft. [1,200 m]; BURMA; Kachin State; "26°05'N, 97°58'E"; collected

by R. Kaulback, 31 Aug. 1939 (BM, 1 [skin only]). 40 Manipur. See Imphal Valley. Man-k'a. See Nanding He. Medog, 950 m; CHINA; Xizang ( = Tibet); 29°19'N, 95°19'E; collected by Feng

Zuo-jian et al. (1980, p. 97; letter 15 Oct. 1980), May-Sep. 1973 (Institute of

Zoology, Beijing, 1, not seen). 27 Mekong. See Ou, Nam. Mekong R., 110 km WNW of Vientiane; LAOS; Xaignabouri or Vientiane; ca.

18°28'N, 101°40'E; collected by F. R. Wulsin, 4 July 1924 (USNM, 1). 63 Mekong River, Yunnan. See Lancang Jiang. Me Pooan. See Puan, Nam Mae. Me Puan. See Puan, Nam Mae.

42 FIELDIANA: ZOOLOGY

Mingun; BURMA; Sagaing Division; 22°01'N/ 96°01'E; erroneous record (Wroughton, 1915, p. 464), based on M. mulatta (BM 14.7.19.1 6).

Mishmi Hills; INDIA; Arunachal Pradesh; 28°00'-29°30'N, 95°00'-97°00'E; col- lected by J. Anderson (1881, p. 70), 6 June 1876 (ZSI, 1 [skin only]). Not mapped.

Mokochung. See Mokokchung.

Mokokchung, 4,500 ft. [1,375 m], 5,000 ft. [1,500 m]; INDIA; Nagaland; ca. 26°20'N, 94°32'E; collected by J. P. Mills (1923, p. 222), 5-19 Sep. 1919, 11 Oct. 1919, 17 Dec. 1923 (BM, 3 [1 skin only]; BNHS, 1; ZSI, 1 [skin only]); collected by G. C. Shortridge, 26 Mar. 1920 (BNHS, 1; FMNH, 1). 34

Monkey Hill. See Tukreshwari Temple.

Monti Carin. See Yado.

Moung Moun. See Muong Moun.

Mt. Angka. See Inthanon, Doi.

Mt. Popa; BURMA; Mandalay Division; 20°55'N, 95°15'E; erroneous record (Wroughton, 1915, p. 464), based on M. mulatta (BM 14.7.19.2, BNHS 5102-4, 5106).

Muang Ngoy, [400-1,000 m]; LAOS; Louangphrabang; 20°43'N, 102°41'E; col- lected by T. D. Carter, 28 Nov. 1931 (AMNH, 1). For altitude, see Legendre, 1936, p. 158. 65

Muang Thateng, 3,000 ft. [900 m]; LAOS; Saravan; 15°26'N, 106°23'E; collected by J. Delacour (1932, p. 419; in Osgood, 1932, p. 198), 14 Dec. 1931. For additional notes on locality, see Engelbach, 1932, pp. 441, 444. 70

Muong Moun, 1,200 ft. [350 m]; VIETNAM; Lai Chau; 21°42'N, 103°22'E; col- lected by R. E. Wheeler, 11 Mar. 1929 (FMNH, 1). For locality note, see Bangs & Van Tyne, 1931, p. 37. 66

Muong Ngoi. See Muang Ngoy.

Myitkina ( = Myitkyina); BURMA; Kachin State; 25°23'N, 97°24'E; probably er- roneous record (Roonwal, 1950, p. 16), apparently based on captive specimen of M. mulatta.

Naga Hills. See Mokokchung.

Nam Hou. See Ou, Nam.

Nam-ka. See Nanding He.

Nam Mae Puan. See Puan, Nam Mae.

Nampuk, west bank, [ca. 490 m]; BURMA; Sagaing Division; ca. 26°02'N, 95°45'E; collected by H. C. Raven (in Mayr, 1938, p. 278), 1 Mar. 1935 (AMNH, 1). 36

Nam Tamai Valley. See Tamai, Nam.

Nam-ting River. See Nanding He.

Nanding He; CHINA; Yunnan; 23°30'N, 98°55'E; collected by R. C. Andrews (1918, p. 245), 3 Mar. 1917 (AMNH, 3 [two skulls only]; MCZ, 1). 45

Negheretting ( = Neghereting); INDIA; Assam; 26°44'N, 94°07'E; erroneous rec- ord (Pilleri, 1975, p. 43), based on temple troop of M. mulatta (see above, p. 3).

Ou, Nam, 80 km above Mekong R.; LAOS; Louangphrabang; ca. 20°38'N, 102°39'E; collected by F. R. Wulsin, 21 June 1924 (USNM, 1). 64

FOODEN: MAC AC A ASSAMENSIS 43

Popa. See Mt. Popa.

Puan, Nam Mae, 1,500-2,000 ft. [450-600 m]; THAILAND; Lampang; 19°03'N, 99°37'E; coUected by J. Chambai, W. J. F. Williamson & M. Smith, Sep. 1917 (BM, 1 [skin and mandible only]). For note on locality and date, see Kloss, 1919, p. 50. 49

Rengma Naga tribal area; INDIA; Nagaland; 25°40'-26°00'N, 94°00'-94°45'E; questionable report (Mills, 1937, p. 102), agricultural pest, possibly based on misidentified M. mulatto. Not mapped.

Sessoungan pagoda. See Irrawady River, 2nd defile.

"Siam" ( = THAILAND); erroneous record (Gray, 1870, p. 31), based on mis- identified specimen of M. fascicularis (probably BM 60.4.20.2 8, "Siam," col- lected by H. Mouhot).

Sisawat. See Chongkrong.

Tamai, Nam, 3,000 ft. [900 m], 4,000 ft. [1,200 m], 4,500 ft. [1,375 m], 5,000 ft.

[1,500 ml; BURMA; Kachin State; "27°42'N, 97°54'E"; collected by R. Kaulback,

28 Mar. 1938, 9-16 Sep. 1938 (BM, 4). 31 Tapaing River. See Taping River. Tapeng River. See Taping River. Tapen R. See Taping River. Taping River, 2 mi [3.7 km] N of Bhamo; BURMA; Kachin State; 24°17'N, 97°14'E;

collected by L. Fea (1886, p. 390), 21 Aug. 1886 (Museo Civico di Storia Naturale

"Giacomo Doria," Genova, 1 [skin only]; not seen, data from Thomas, 1892,

p. 916). 43 Taron Valley. See Taron, Wang. Taron, Wang, 6,000 ft. [1,800 m]; BURMA; Kachin State; "27°40'N, 98°10'E";

collected by R. Kaulback, 5 Dec. 1938 (BM, 1). 33 Tatkon; BURMA; Sagaing Division; 23°47'N, 94°29'E; erroneous record (Wrough-

ton, 1916a, p. 296), based on M. mulatto (BM 15.5.5.6, BM 31.1.11.23, BNHS

5090-92).

Tebang river, 2,000 ft. [600 m]; INDIA; Arunachal Pradesh; ca. 28°00'N, 96°10'E; collected by H. W Wells, 27-29 Apr. 1921 (BM, 2). For locality note, see Hinton & Lindsay, 1926, p. 385. 28

Tecang River. See Tebang River.

Thateng. See Muang Thateng.

Tounghoo. See Yado.

Tukreshwari Temple; INDIA; Assam or Meghalaya; ca. 26°00'N, 90°30'E; ques- tionable record (Hill, 1974, p. 733), based on provisioned troops observed by M. Bertrand (1969, p. 16) and R. K. Lahiri, Jan. 1965, possibly M. mulatto (see above, p. 4). Not mapped.

Tungtang-Dagunghka. See Jantang-Dagung Hka.

Vientiane. See Mekong R., 110 km WNW of Vientiane.

Vinh Linh region; VIETNAM; Vinh Linh Special Zone; ca. 17°04'N, 107°02'E;

probably erroneous record (Dao Van Tien, 1960, p. 228), apparently based on

M. mulatto (see above, p. 4).

44 FIELDIANA: ZOOLOGY

Yado, 1000 m.; BURMA; Pegu Division; 19°14'N, 96°55'E; collected by L. Fea (1888, p. 856), 22 Dec. 1887 (Museo Civico di Storia Naturale "Giacomo Doria," Genova, 1; not seen, data from Thomas, 1892, p. 916, and De Beaux, 1923, p. 27). 47

Yenangaung. See Yenangyaung.

Yenangyaung; BURMA; Magwe Division; 20°28'N, 94°53'E; erroneous record (Wroughton, 1915, p. 464), based on M. mulatta (cf. Hinton & Wroughton, 1921, p. 666).

Yin (and Kin); BURMA; Sagaing Division; 22°47'N, 94°42'E; erroneous record (Wroughton, 1916a, p. 296), based onM. mulatta (BM 15.5.5.7, BNHS 5095-99, FMNH 82806-7, ZSI 12094).

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APPENDIX

Macaca mulatta localities shown in figs. 8B and 9 are documented below by reference to specimen collections or literature records. These localities are grouped by country and listed in west-east sequence for each unit degree of latitude.

Figure 8B: Macaca mulatta localities

CHINA: Nyainqentanglha Sheng, ca. 29°N, 90°E (Shen Hsiao-chou, 1963, p. 140).

NEPAL: Chaur, 29°17'N, 80°21'E (Chesemore, 1970, p. 164). Hutu Forest, 29°35'N, 80°05'E (Richie et al., 1978, p. 443). Dhangarhi, 28°34'N, 80°36'E (Chesemore, 1970, p. 164). Bheri R., ca. 28°30'N, 82°00'E (Byrne, 1979, p. 70). Chengli, ca. 28°00'N, 84°35'E (BM). Trisuli, 4 mi SE, ca. 27°53'N, 84°10'E (C. O. Maser, field catalog, FMNH). Dudurhani, 27°34'N, 84°14'E; Simri, 27°36'N, 84°19'E (Chesemore, 1970, p. 164). Hitaura, W mi. W, 27°27'N, 85°02'E (C. O. Maser, field catalog, FMNH). Russian Camp, 27°12'N, 85°04'E; Simri[bis], 27°06'N/ 85°14'E; Swayambhunath, ca. 27°43'N, 85°18'E; Singaul, Tekan, 27°10'N, 85°20'E (Chesemore, 1970, p. 164). Pashupari, ca. 27°38'N, 85°21'E (Taylor et al., 1978, p. 344). Chandikhola, 27°04'N, 85°22'E (Chesemore, 1970, p. 164). Gokarna, 27°43'N, 85°23'E (FMNH). Hazaria Patherghatta, 27°05'N, 85°25'E; Nagarkot, 27°42'N, 85°31'E (BM). Sabaya Khola, ca. 27°20'N, 87°15'E (J. A. McNeeley, letter, 4 Feb. 1973). Morang district, ca. 26°30'N, 87°30'E (N. A. Baptista in Hinton & Fry, 1923, p. 403).

INDIA: Nishangara, 28°15'N, 81°13'E (Kurup, 1965, p. 193). Tarkhola, 27°07'N, 88°33'E (Khajuria, 1966, p. 284). Lucknow, 26°51'N, 80°55'E (Southwick et al., 1961a, p. 543). Lucknow-Faizabad, ca. 26°50'N, 81°30'E; Faizabad-Ajodhya, ca. 26°45'N, 82°10'E; Ajodhya- Gorakpur, ca. 26°45'N, 82°45'E (Southwick et al., 1961b, p. 702). Azamagarh, ca. 26°04'N, 83°11'E (Southwick et al., l%la, p. 541). Panighatta, 26°48'N, 88°15'E (Southwick et al., 1964, p. 444). Narbong, 26°51'N, 88°19'E (BM). Sukna, 26°47'N, 88°22'E (ZSI). Mangpu, 26°58'N, 88°24'E (FMNH). Siliguri, 26°42'N, 88°26'E (Southwick et al., 1964, p. 444). Sivok, 26°52'N, 88°29'E (FMNH). Jaldapara Wildlife Sanctuary, ca. 26°38'N, 89°18'E (Spillett, 1%7, p. 544). Hasimara, 26°44'N, 89°22'E; Bannabari, 26°44'N, 89°24'E (BM). Maure, Jamduar, 26°43'N, 89°53'E (Gee, 1961, p. 5; Mukherjee, 1978b, p. 742). Raimona, 26°39'N, 89°58'E (Mukherjee, 1978b, p. 742). Bogra Nadi, ca. 26°50'N, 91°35'E (BM). Kulsi, 25°59'N, 91°24'E (BNHS). Rajapara, 25°52'N, 91°26'E (BM). Agartala, ca. 25 km S, ca. 23°38'N, 91°18'E; Ampi, ca. 23°40'N, 91°38'E (ZSI). Calcutta, 22°32'N, 88°22'E (ZSI; Southwick et al., 1964, p. 444). Basinhat Reserve Forest, ca. 22°40'N, 88°53'E; Jhilla, ca. 22°N, 89°E; Harinbhanga, ca. 21°45'N, 89°00'E (Mukherjee & Gupta, 1965, p. 145). Note: Pashok (27°03'N, 88°25'E) is reported by Wroughton (1916c, p. 776) as the locality of the two M. mulatta specimens said to have been collected for the BNHS Mammal Survey of India, but no such specimens can now be found and the locality is therefore doubtful.

BANGLADESH: Madhupur, ca. 24°37'N, 90°02'E; Dhamrei, 23°55'N, 90°13'E; Dacca, 23°43*N, 90°25'E (Green, 1978, pp. 146, 154). Calcutta, ca. 50 mi E, ca. 22°35'N, 89°15'E (ZSI). Satkhira, S, ca. 22°N, 89°E (Mandal, 1964, p. 164).

Figure 9: Macaca mulatta localities

CHINA: Kumtatchie ( = Houmda), 31°05'N, 96°44'E (Bonvalot, 1891, pp. 149, 156). Rou- toundo, 31°35'N, 97°25'E (MNHN). Xi Golog ( = Singolo), 30°02'N, 100°45'E (MCZ). Rama

52 FIELDIANA: ZOOLOGY

Pass, 30°01'N, 100°52'E (Schafer, 1933, p. 280). Yajiang ( = Na-chu-kar), 30°02'N, 101°02'E (MCZ). Kongbo, ca. 29°00'N, 93°15'E; Dowoka, ca. 29°22'N, 94°18'E; Gyala, above, ca. 29°38'N, 94°56'E (Bailey, 1915, p. 74; 1957, pp. 122, 171). Dzo La, SE, ca. 29°13'N, 97°07'E (Kaulback, 1938, p. 91). Tsari Chu, ca. 28°45'N, 93°10'E (Bailey, 1915, p. 74). Ashi, ca. 26°56'N, 100°00'E (USNM). Kakyen Hills, ca. 25°02'N, 98°28'E (ZSI). Teng Yueh [ca. 80 mi N], 25°13'N, 98°28'E (AMNH, MCZ). Hotha Valley, ca. 24°25'N, 97°55'E (ZSI). Kengma, 23°31'N, 99°24'E (Yen Wen-chen, 1973, p. 362). Mengyang, ca. 22°01'N, 100°54'E (Kao Yueh-ting et al., 1962, p. 188; cf. Bannikov, 1958, p. 68). Menghun, 21°50'N, 100°23'E; Meng-lung, 21°47'N, 100°27'E (Kao Yueh-ting et al., 1962, p. 188). Nam Fong, 19°24'N, 109°31'E; Nada (Nodoa), 19°31'N, 109°33'E (AMNH).

INDIA: Yongyap Valley, ca. 29°10'N, 95°37'E (Bailey, 1915, p. 74). Dening, 28°01'N, 96°14'E (BM, BNHS). Dangori Nadi, ca. 27°36'N, 95°16'E (Pilleri, 1975, p. 20; letter 15 Dec. 1978). Margherita, 27°17'N, 95°41'E (ZSI). Baguri Block, ca. 26°37'N, 93°15'E (Spillett, 1967, p. 496). Haldhibari Block, 26°37'N, 93°22'E (Lahan & Sonowal, 1974, p. 261). Gohalghat, 26°31'N, 93°58'E (BM). Neghereting, 26°44'N, 94°07'E (Pilleri, 1975, p. 43). Changchang Pani, 26°25'N, 94°25'E (AMNH). Chungtia, 26°24'N, 94°28'E (Mills, 1923, p. 222). Nongpoh, 25°54'N, 91°53'E (BM). Garam Pani ( = Hot Springs), ca. 25°30'N, 92°35'E (BNHS). Lam- sakhang, ca. 25°50'N, 93°05'E (BM). Samaguting, 25°48'N, 93°48'E (ZSI). Imphal, 4 mi. N, 24°52'N, 93°56'E (ZSI). Bishenpur, 24°38'N, 93°46'E (BM).

BANGLADESH: Chittagong Hill Tracts, ca. 21°45'N, 92°25'E (BM; Green, 1978, p. 146).

BURMA: Singkaling Hkamti, ca. 26°00'N, 95°42'E (AMNH, BM, BNHS, ZSI). Taga Hka, 26°21'N, 96°09'E; Taro (Dalu) 26°21'N, 96°11'E (AMNH). Bawmwang, 26°39'N, 97°50'E; Hkandau, 26°01'N, 97°50'E (BM); Htingnan Triangle, 26°36'N, 97°52'E. Maungkan, 25°05'N, 95°02'E (AMNH). Hisweht, 25°22'N, 95°16'E (BM). Moklok, 25°37'N, 95°25'E; Heinsun, 25°52'N, 95°35'E; Nanyaseik, 25°37'N, 96°36'E (AMNH). Pidaung Reserve, ca. 25°25'N, 97°09'E (BM). Myitkyina, 25°23'N, 97°24'E (Roonwal, 1950, p. 16). N'Changyang, 25°50'N, 97°48'E (BM). Tanga-Shingaw Road, ca. 25°40'N, 97°55'E (AMNH). Homalin, 24°52'N, 94°55'E (BM, BNHS). Bhamo, ca. 24°16'N, 97°14'E; Kindat, 20 mi. NW, ca. 23°50'N, 94°10'E; Ali Cha, ca. 23°25'N, 94°15'E; Kindat, 23°44'N, 94°26'E (BM). Tatkon, W bank, 23°47'N, 94°29'E (BM, BNHS). Tatkon, E bank, ca. 23°47'N, 94°30'E; Kin, 22°46'N, 94°42'E (BNHS). Yin, 22°47'N, 94°42'E (BNHS, FMNH, ZSI). Mingun, 22°00'N, 95°57'E; Kokko- aing, 22°28'N, 96°18'E (BM). Maymo, 22°02'N, 96°28'E (AMNH). Pyaunggaung, 22°35'N, 97°05'E (BM, BNHS). Se-eng, 22°43'N, 97°31'E (BM). Mamsam Falls, ca. 22°50'N, 97°35'E (BNHS). Akyab, NE, ca. 21°00'N, 94°00'E; Yenangyaung, below, ca. 20°30'N, 94°55'E (Anderson, 1879, pp. xviii, 57). Popa Hill, 20°56'N, 95°16'E (AMNH, BM, BNHS). Toungoo, 30 mi. NW, ca. 19°10'N, 96°05'E (BNHS, ZSI). Toungoo, 20 mi. W, ca. 19°00'N, 96°10'E (BNHS). Toungoo, 15 mi. N, ca. 19°10'N, 96°25'E (BM). Toungoo, 13 mi. E, ca. 19WN, 96°40'E (BM, BNHS). Prome, 30 mi. SE, ca. 18°40'N, 95°36'E (BNHS). Theme, ca. 18°45'N, 95°45'E; Toungoo, E, ca. 18°56'N, 96°26'E (BM).

THAILAND: Doi Luang Chiang Dao, 19°23'N, 98°54'E (MCZ). Chiang Mai, vicinity, ca. 18°45'N, 99°00'E (Zool. Ref. Coll., Singapore). Pang Nam Un, Nan, ca. 18°30'N, 100°33'E (USNM). Chiang Mai, rapids below, ca. 17°52'N, 98°40'E (Zool. Ref. Coll., Singapore). Huai Ap Nang, 17°25'N, 98°43'E; Huai Kwang Pah, 17°28'N, 98°50'E (FMNH). Dan Sai district, ca. 17°17'N, 101°09'E (USNM). Ban Mae Lamao, vicinity, ca. 16°49'N, 98°45'E (FMNH). Huai Nua Pla, 16°54'N, 98°48'E; Tha Chang Tai, 16°51'N, 99°03'E (Zool. Ref. Coll., Singapore). Ban Umphang, 28 mi. SE, ca. 15°40'N, 99°13'E (AMNH). Huai Chang Tai, 15°27'N, 99°15'E; Khao Nang Rum, NW, ca. 15°30'N, 99°17'E (Eudey, 1979, pp. 91-97, table 7).

LAOS: Nam Hou, ca. 21°00'N, 102°45'E (FMNH). Xaignabouri (Sayabouri), ca. 19°15'N, 101°45'E; Louangphrabang, ca. 19°52'N, 102°08'E; Xiangkhoang, ca. 19°20'N, 103°22'E (Deuve & Deuve, 1963, p. 59). Mekong River, ca. 18°03'N, 101°57'E (USNM). Ban Mak Nao, ca. 18°00'N, 102°58'E (Zool. Ref. Coll., Singapore). Pakxan, ca. 18°22'N, 103°40'E; Vientiane, ca. 17°58'N, 102°36'E (Deuve & Deuve, 1963, p. 59). Nong Khai Camp, No. 28, ca. 17°52'N, 102°44'E (Zool. Ref. Coll., Singapore).

VIETNAM: Muong Bourn, 22°23'N, 102°50'E (FMNH). Muong Mo, 22°13'N, 102°55'E (Coolidge in Coolidge and Roosevelt, 1933, pp. 86, 216). Bac Can, 22°08'N, 105°49'E (BM, MNHN). Muong Pon, 21°33'N, 103°01'E; Muong Muon, 21°41'N, 103°02'E (AMNH). Muong Moun, 21°42'N, 103°22'E (FMNH). Phu Qui, 19°19'N, 105°25'E (BM). Vinh Linh, vicinity, 17°04'N, 107°02'E (Dao Van Tien, 1962, p. 724). Hue, 16°28'N, 107°36'E (Delacour, 1940, p. 24). Song-Ta-Voy, ca. 16°08'N, 107°45'E (MNHN). Dak Sut, 14°56'N, 107°45'E (USNM).

Field Museum of Natural History Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496 Telephone: (312) 922-9410

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