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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 measurements 1 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 


[?] 620 2 


[?] .61 2 


[?] 4.7 2 




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' 


42 3 


437 


.44 






23°30' 


45 3 


480, 500 


.47,.42 


6.9, 7.0 




20°43' 


65 


[?] 580 4 


[?] .26 4 






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' 


26 5 


630,665 


.37, .30 


6.0, 6.0 


14.5, 13.0 


29°19' 


27 5 


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 


[?] .18 4 


5.6 






22°21' 


67 b 


600 


.36 








21°42' 


66 


[?] 495 4 


[?] .50 4 




. 


• 


20°22' 


69 


575 


.40 


6.8 






19°14' 


47 7 




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 


116 1 , 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 


... 




134.6 2 


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 






146 3 


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 L 2 = 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) 



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 
Adults 1 



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. pelops 2 

3 .60 5 

M. a. assamensis 3 


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. 


infant 1 


month 2 




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,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; 26 o 00'-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). 




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