Skip to main content

Full text of "Report on primates collected in western Thailand, January-April, 1967"

See other formats


BIOLOGY 



The person charging this material is re- 
sponsible for its return to the library from 
which It was withdrawn on or before the 
Latest Date stamped below. 

Theft, mutllotlon, and underilning of books or. reasons 
for dlsciplinory octlon and may result In dismissal from 
tlie University. 
To renew coll Telephone Center, 333-8400 

UNIVERSITY OF ILIINOIS IIBRARY AT URBANA-CHAMPAIGN 




L161— O1096 



r9 




FIELDIANA 
Zoology 

Published by Field Museum of Natural History 



VOLUME 59, NUMBER 1 

REPORT ON PRIMATES 

COLLECTED IN WESTERN THAILAND 
JANUARY-APRIL, 1967 

JACK FOODEN 



MARCH 81, 1971 



MAY 25 1971 






FIELDIANA: ZOOLOGY 

A Continuation of the 

ZOOLOGICAL SERIES 

of 

FIELD MUSEUM OF NATURAL HISTORY 



VOLUME 59 




FIELD MUSEUM OF NATURAL HISTORY 
CHICAGO, U.S.A. 



THE LIBRARY OF THE 

MAYS 1972 



TABLE OF CONTENTS 

PAGE 

1. Report on Primates Collected in Western Thailand January-April, 1967. 
By Jack Fooden 1 

2. The Viperid Snake Azemiops: Its Comparative Cephalic Anatomy and 
Phylogenetic Position in Relation to Viperinae and Crotalinae. By Karel 

F. Liem, Hymen Marx, and George B. Rabb 63 



\. 



FIELDIANA 
Zoology 

Published by Field Museum of Natural History 



VOLUME 59, NUMBER 1 

REPORT ON PRIMATES 

COLLECTED IN WESTERN THAILAND 
JANUARY- APRIL, 1967 

JACK FOODEN 

Research Associate, Field Museum of Natural History 

and 

Professor of Zoology, Chicago State College 



MARCH 31, 1971 



PUBLICATION 1123 



Patricia M. Williams 
Editor 



Library of Congress Catalog Card Number: 76-155312 



PRINTED IN THE UNITED STATES OF AMERICA 
BY FIELD MUSEUM PRESS 






CONTENTS 

PAGE 

Introduction 7 

Zoogeography and Ecology 11 

Itinerary and Locality Notes 14 

Species Accounts 21 

Nycticebus coucang 21 

Macaca fascicularis 22 

Macaca mulatta 32 

Macaca nemestrina 32 

Macaca assamensis 36 

Macaca arctoides 38 

Presbytis cristatus 39 

Presbytia phayrei 42 

Hylobates lar 43 

Plates 48 

References 59 



ABSTRACT 

During four months of field work 152 specimens of nine species 
of primates were collected in western Thailand. Three species — 
Macaca fascicularis, M. nemestrina, Presbytis cristatus — are Indo- 
Malayan; three — M. mulatta, M. assamensis, P. phayrei — are In- 
do-Chinese; and three — Nycticebus coucang, M. arctoides, Hylobates 
lar — inhabit both faunal and subregions. Specimens collected indi- 
cate that the range of M. nemestrina is marginally sympatric with 
that of M. assamensis, whereas the ranges of M. fascicularis and 
P. cristatus appear to be allopatric with those of M. mulatta and 
P. phayrei. Analysis of new evidence concerning the zoogeographical 
and morphological inter-relationships of M. fascicularis and M. mu- 
latta suggests that these macaques should be regarded as species 
instead of subspecies as previously indicated. Most encounters with 
the nine primate species collected were in evergreen forest, but 
N. coucang, M. fascicularis, and M. mulatta also were taken in 
groves of bamboo. Only M. arctoides was encountered out of the 
trees on the forest floor. For each species collected external measure- 
ments are given and observations are recorded on habitats, group 
size, stomach contents, and reproductive condition. 



INTRODUCTION 

During the first four months of 1967 I collected primates in 
Western Thailand for Field Museum of Natural History as part of 
a continuing program of taxonomic research that is primarily fo- 
cussed on the genus Macaca. The expedition period in Thailand 
coincided with the final two-thirds of the annual dry season (north- 
east monsoon). Nineteen localities were visited along the Dawna 
Range, a mountain chain on the Thai-Burmese border (fig. 1), and 
a total of 152 primates of nine species were collected (Table 1). 
Standard dry skins and skulls were preserved routinely, and, when- 
ever possible, stomachs with contained food material and external 
and internal reproductive organs were preserved in fluid. A few 
skeletons and undissected infants in fluid also were preserved. Notes 
on habitats and behavior were recorded for most primate troops 
observed in the course of the expedition. 

The expedition party consisted of myself and two Thai associates, 
Mr, Pong Leng-EE and Mr. Wirot Meangmongkoon, plus local 
hunters, guides, and assistants temporarily employed at each col- 
lecting site as required. Until recently, the hill country in which 
we worked was sparsely inhabited and minimally disturbed. How- 
ever, as a result of rapid expansion of the Thai population and econ- 
omy, the region currently is in the process of being settled, deforested, 
and cultivated. Consequently, we frequently had to travel fairly 
long distances from main roads in order to reach suitable collecting 
sites. Our transportation was by jeep. Land Rover, lumber truck, 
motor canoe, and motor launch, depending on what was locally avail- 
able and appropriate. Usually we camped in or near isolated vil- 
lages, where we invariably were received with the generous hospi- 
tality for which Thai people are famous. 

The expedition was financed by U. S. Public Health Service 
Grant No. GM 13113. Valuable advice and assistance was provided 
by the U. S. Department of State. In Thailand the expedition re- 
ceived indispensable support and assistance from the Royal Forest 
Department and the Applied Scientific Research Corporation, to 
whose officials and staff members I am deeply grateful. Special 
thanks are due my field companions Mr. Pong Leng-EE of the Royal 



iKW7,:r 



lOO" 101" 




Fig. 1. Map of western Thailand showing location of collecting localities. 
CHANGWAT (=province) MAE HONG SONG: 1— Mae Sariang, about 30 km. 
E (collected by Dr. G. Berkson); CHANGWAT TAK: 2— Huai Ap Nang, 
3 — Huai Kwang Pah (left bank Mae Nam Ping) and Huai Wang Kwao (right 
bank Mae Nam Ping), 4— Ban Mae Lamao; CHANGWAT KAMPHAENG 
PHET: 5— Ban Pong Nam Ron, 6— Ban Mae Na Ree, 7— Ban Nam Lai Tai, 
8— Ko Keow; CHANGWAT NAKHON SAWAN: 9— Khao Naw; CHANGWAT 
UTHAI THANI: 10— Kata Taek, 11— Samnak Rabam, 10 km. SE, 12— Khao 
Phatowee; CHANGWAT KANCHANABURI: 13— Ban Kerng Chada, 14— Ban 
Tamrong Phato, 15— Ban Muang Baw Ngam, 16 — Chongkrong, 17 — Ban Phu 
Toei, 18— Ban Huai Maenam Noi; CHANGWAT LOP BURI: 19— Lop Buri. 



8 



FOODEN: PRIMATES FROM THAILAND 9 

Forest Department, who made important contributions to the plan- 
ning of all phases of the expedition, and Mr. Wirot Meangmongkoon, 
who served diligently as chief field assistant, hunter, and interpreter. 
In species accounts in this report some specimens collected in 
Thailand are compared with specimens preserved in the museums 
listed below; names of museums are abbreviated as indicated. I 
thank curators of these institutions for permission to study material 
in their custody. 

AMNH. — American Museum of Natural History, New York 
BMNH. — British Museum (Natural History), London 
FMNH. — Field Museum of Natural History, Chicago 
MCZ. — Museum of Comparative Zoology, Harvard University, 
Cambridge 
MNHN.— Museum National d'Histoire Naturelle, Paris 
USNM. — United States National Museum, Washington, D. C. 






^ ^ 



rH C<1 CO CO •^ 



•<^ I— I lo CO 



-^ l-H 



« O T-l 

^ Is 



rH ■<* I tH 






ini (M OS 



C9 



n 

Eh 



o C 






.2 
'S 



§ s i 



£. 



e -S V2 



gi 03 






o 



10 



ZOOGEOGRAPHY AND ECOLOGY 

Western Thailand is an area of special zoogeographic importance 
in the Oriental faunal region because it is in or near the zone of tran- 
sition between the subtropical Indo-Chinese subregion and the trop- 
ical Indo-Malayan subregion (Pocock, 1939, p. xxiii; Chasen, 1940, 
p. x). This subregional faunal transition is manifest in the distri- 
bution of six of nine primate species collected by the expedition. 
Macaca mulatta, M. assamensis, and Presbytis phayrei are Indo- 
Chinese species, and M. fascicularis, M. nemestrina, and P. cristatus 
are the respective Indo-Malayan counterparts. In the zone of tran- 
sition in western Thailand, expedition collecting results indicate that 
M. mulatta and M. fascicularis are allopatric, M. assamensis and 
M. nemestrina are marginally sympatric, and P. phayrei and P. cris- 
tatus are allopatric (Table 1). 

Although borders between the ranges of these three pairs of spe- 
cies are in the same general area in western Thailand, these borders 
apparently do not coincide precisely. In Indo-Chinese species, the 
known ranges of M. assamensis and P. phayrei extend south as far as 
Chongkrong (14°41'N), whereas the range of M. mulatta apparently 
does not extend south beyond the vicinity of Ban Umphang (approx. 
16°00'N; Fooden, 1964, p. 363). In Indo-Malayan species, the 
known range of P. cristatus extends north to Ban Kerng Chada 
(15°08'N), that of M. fascicularis extends to Ban Mae Na Ree 



Table 2. — Forest types in which primate troops were encountered; 
for details see Species Accounts. 



Species 




Forest type 






Deciduous 


Bamboo 


Evergreen 


Nycticebus coucang 





1 


1 


Macaca fascicularis 


1-2 


5 


4-6 


M. mulatta 





2 


1 


M. nemestrina 


1 





8 


M. assamensis 





2 


8 


M. arctoides 








2 


Presbytis cristatus 








5 


P. phayrei 








9 


Hylobates lar 








19 



11 



12 FIELDIANA: ZOOLOGY, VOLUME 59 

Table 3. — Food material in stomachs and cheek pouches of primate species 
collected; + +indicates major component, +indicates minor component; 
for details see Species Accounts. 

Contents of stomachs and cheek pouches 



Stomachs Fruit, 

Species examined Leaves seeds Invertebrates Vertebrates 

Nycticebus coucang 2 + + + 

Macaca fascicularis 28 ++ + 

M. mulatta 3 + + + 

M. nemestrina 7 + + + 

M. assamensis 10 + + + + 

M. arctoides 4 + + 

Presbytis cristatus 4 + + 

P. phayrei 5 + + 

Hylobates lar 21 + + 

(16°25'N), and that of M. nemesirma extends to Khun Tan (18°35'N; 
Gyldenstolpe, 1917, p. 7). 

Forest types in which the nine primate species collected were ob- 
served are indicated in Table 2. All species occur in evergreen trees, 
and four were observed in no other habitat. Bear macaques (Macaca 
arctoides), two species of langurs (Presbytis cristatus, P. phayrei), and 
gibbons (Hylobates lar) were encountered exclusively in evergreen 
forest. Almost all encounters with Assamese and pigtail macaques 
(M. assamensis, M. nemestrina) also were in evergreen forest; how- 
ever, M. assamensis was encountered twice in groves of bamboo, and 
M. nemestrina was encountered once in deciduous trees. Slow lorises 
(Nycticebus coucang), crab-eating macaques (M. fascicularis), and 
rhesus macaques (M. mulatta) appear to be least restricted to ever- 
green forest; all were encountered in bamboo forest approximately 
as often as in evergreen forest, and M. fascicularis also was encoun- 
tered twice in deciduous forest. Botanical characterizations of habi- 



Table 4. — Probable month of birth estimated for fetuses (F) and infants (I) 
of primate species collected; for details see Species Accounts. 

Estimated month of birth 
Species Aug. Sept. Oct. Nov. Dec. Jan. Feb. Mar. Apr. May June July 

Macaca 

fascicularis F,I F,F F 

M. nemestrina 1,1 F F 

M. assamensis F F F,F 

Presbytis 

cristatus I F F 

P. phayrei 1,1 I I 

Hylobates lar I 1,1 



FOODEN: PRIMATES FROM THAILAND 13 

tats in some areas visited during the present expedition and in similar 
areas in neighboring parts of Thailand have been published by Larsen 
(1962, p. 110) and Tern Smitinand (1968, p. 289). 

Specimens of most sympatric species were collected — frequently 
on the same day — at points so close together that it seems highly 
probable that these species pass through the same trees in the course 
of their normal activity. Although M. fascicularis and M. nemestrina 
are broadly sympatric, these two macaques may be ecologically seg- 
regated, judging from the rarity of their concurrent collection (Ta- 
ble 1). Of primate species collected, only M. arctoides appears to 
spend much time on the ground (see Species Accounts and McCann, 
1933, p. 807). 

Major categories of food material contained in the stomachs and 
cheek pouches of primate species collected are summarized in Table 3. 
Preliminary analysis of preserved stomach contents indicates that all 
seven frugivorous species frequently fed on the same kinds of fruit. 

Evidence of seasonal breeding that is provided by the estimated 
ages of fetuses and infants collected by the expedition is summarized 
in Table 4. 



ITINERARY AND LOCALITY NOTES 

December 26, 1966 — Left Chicago. 

December 29, 1966-January 9, 1967 — Bangkok (=Krung Thep). 

January 11-23— Ban Muang Baw Ngam, 14°55'N, 98°55'E, alt. 
approx. 1,100 m. (fig. 1, Loc. No. 15). This lead and zinc mining 
camp in the highlands between Mae Nam Khwae Yai (=Mae Nam 
Mae Klong) and Mae Nam Khwae Noi has a resident population 
of about 100, including miners and their families. The shallow sub- 
surface ore deposits are worked by means of small manually-exca- 
vated open pits. The terrain is hilly with numerous steep rocky 
outcrops. Dense evergreen forest interspersed with groves of bamboo 
covers the hills (Plate I, a). All specimens were collected within a 
half day's walk of camp. The temperature at Ban Muang Baw 
Ngam was markedly cooler than at any other collecting locality 
visited (Table 5). 

January 26-29— Chongkrong, 14°41'N, 98°52'E, alt. 600-900 m. 
(fig. 1, Loc. No. 16). This is a small intermittently occupied lumber 
camp about 17 km. west of Si Sawat (see below) and about 25 km. 
south of Ban Muang Baw Ngam (see above). The terrain is some- 
what less rugged than at Ban Muang Baw Ngam, but the evergreen 
and bamboo forest is essentially similar. All specimens were taken 
within two or three hours' walk of camp. 

January 30-February 1— Si Sawat, 14°41'N, 99°02'E, alt. ap- 
approx. 50 m. This village of about 100 houses is on the right bank 
of Mae Nam Khwae Yai about 90 km. upstream from Kanchanaburi, 
the provincial capital. The village is the district government head- 
quarters. It is surrounded by cultivated fields and low hills covered 
with deciduous and bamboo forests. Local residents informed us 
that long-tailed macaques (M. fascicularis) in the area are restricted 
to the east (left) bank of the river. Accordingly, we sent one of our 
hunters on an overnight collecting trip to Ban Phu Toei (fig. 1, Loc. 
No. 17), a small settlement about 8 km. northeast of Si Sawat, where 

14 



FOODEN: PRIMATES FROM THAILAND 15 

Table 5. — Temperature extremes at collecting localities visited. 



Collecting locality 


Date (1967) 


Night-time lows 


Day-time high 


Ban Muang Baw Ngam 


Jan. 11-22 


44-57°F 


74-79°F' 


Chongkrong 


Jan. 26-29 


63-66 


88-91 


Si Sawat 


Jan. 30-Feb. 1 


65-66 


91-97 


Ban Kerng Chada 


Feb. 4-8 


53-63 


86-89 


Ban Tamrong Phato 


Feb. 9-13 


60-63 


92-93 


Ban Huai Maenam Noi 


Feb. 14-18 


58-62 


86-88 


Kata Taek 


Feb. 27-March 3 


65-68 


84-93 


Ko Keow 


March 5-10 


65-71 


88-92 


BanMaeNaRee 


March 13-15 


66-68 


96 


Ban Mae Lamao 


March 17-26 


62-66 


92-95 


Huai Kwang Pah 


March 28-30 


62-76 


92-93 


Ban Nam Lai Tai 


April 1-6 


68-73 


97-99 


Ban Pong Nam Ron 


April 8-15 


69-74 


95-103 


Khlong Suan Mak 


April 20-24 


73-76 


88-90 



1 Excludes exceptional high of 87°F recorded on Jan. 22. 

the hunter obtained two specimens of M. fascicularis in bamboo 
forest along a nearby creek (Huai Ong Sit). 

February 2-3— Ban Wang Kalang (=Sangkhla Buri), 15°06'N, 
98°28'E, alt. approx. 100 m. This large village near the headwaters 
of Mae Nam Khwae Noi is the district government headquarters. 
On a hunting trip about 10 km. downstream from Ban Wang Kalang 
we encountered a troop of about 20 M. fascicularis in bamboo forest 
along the bank of a small western (right) affluent of Mae Nam Khwae 
Noi, but we were unable to collect specimens. During the return 
trip upstream, from our boat we observed a second troop of monkeys 
(probably also M. fascicularis) in another bamboo grove on the right 
bank of Mae Nam Khwae Noi. 

February 4-8— Ban Kerng Chada, 15°08'N, 98°31'E, alt. ap- 
approx. 150 m. (fig. 1, Loc. No. 13). This Karen village has a pop- 
ulation of about 150 and is about 10 km. east of Ban Wang Kalang 
(see above) on the right bank of Mae Nam Ran Ti, one of the head- 
waters of Mae Nam Khwae Noi. Ban Kerng Chada is in the foot- 
hills of a mountain range that reaches a peak of 1,805 m. about 25 
km. to the northeast of the village. The hills are covered with ever- 
green forest interspersed with occasional groves of bamboo. All 
specimens were taken within a half day's walk of the village. 

February 9-13 — Ban Tamrong Phato (=Ban Wang Phato), 
14°54'N, 98°31'E, alt. approx, 100 m. (fig. 1, Loc. No. 14). This 
village of about 80 houses is on the west (right) bank of Mae Nam 
Khwae Noi about 25 km. downstream from Ban Wang Kalang (see 



16 FIELDIANA: ZOOLOGY, VOLUME 59 

above). The village is surrounded by cultivated fields, with ever- 
green and bamboo forests on the hills beyond. Seven crab-eaters 
(M. fascicularis) were taken within two hours' walk of the village in 
evergreen trees near the river; two gibbons and one crab-eater were 
taken about a half day's walk from the village; and three langurs 
were taken on an overnight hunting trip to Phapung, about 10 km. 
from Ban Tamrong Phato. 

February 9-13— Ban Huai Maenam Noi, 14°25'N, 98°51'E, alt. 
approx. 75 m. (fig. 1, Loc. No. 18). This village consists of about 
10 houses built on rafts moored to both banks of Mae Nam Kwai 
Noi about 75 km. northwest of Kanchanaburi, the provincial capital. 
The flat land near the river banks is under cultivation. Beyond the 
fields are low hills covered with evergreen and bamboo forest. Our 
hunters made several half-day trips by boat and lumber truck in 
order to collect on both banks of the river upstream and downstream 
from the village. Local residents informed us that the forests in 
which we hunted around Ban Huai Maenam Noi are the southern- 
most remaining monkey habitats on Mae Nam Khwae Noi. 

February 19-23 — Bangkok. 

February 24— Lop Buri, 14°48'N, 100°37'E, alt. approx. 25 m. 
(fig. 1, Loc. No. 19) . This town is a provincial capital about 120 km. 
north of Bangkok. It is traversed by the Bangkok-Chiang Mai 
highway. A colony of 100-200 long-tailed macaques (M. fascicu- 
laris) inhabits the ruins of an old Buddhist temple that are preserved 
on a traffic island in the center of town (Plate II, a). The monkeys 
are fearless and well fed by resident monks and crowds of devout 
visitors. Temple custodians believe that this colony is descended 
from wild monkey populations that inhabited the Lop Buri area 50 
or 60 years ago when the region was densely forested. I know of 
no reason to doubt this explanation of the colony's origin. Today, 
the nearest populations of wild monkeys probably are at Khao Yai 
National Park, about 100 km. southeast of Lop Buri. 

February 25— Khao Phatowee, 15°28'N, 99°45'E, alt. approx. 
50 m. (fig. 1, Loc. No. 12). This sacred hill is an isolated limestone 
crag that rises about 100 m. above the rice fields of the surrounding 
plain (Plate I,b,c). The hill, which is riddled with caves and over- 
grown with shrubs and trees, is inhabited by a colony of 100-200 
long- tailed macaques (M. fascicularis). As at Lop Buri (see above), 
local residents believe that the Khao Phatowee monkey colony is a 



FOODEN: PRIMATES FROM THAILAND 17 

relict of wild populations that inhabited extensive forests that ex- 
isted in the area up to about 40 years ago, before the onset of inten- 
sive cultivation. 

February 26— Samnak Rabam, 15°31'N, 99°29'E. alt. approx. 
150 m. Enroute to Kata Taek (see below), about noon from our 
truck we spotted a troop of pigtail macaques (M. nemestrina) in a 
stretch of evergreen forest along the trail approximately 10 km. 
southeast of the village of Samnak Rabam. We dismounted and 
pursued the troop into the forest, and succeeded in collecting two 
specimens (fig. 1, Loc. No. 11). 

February 27-March 3— Kata Taek, 15°28'N, 99°23'E, alt. ap- 
prox. 200 m. (fig. 1, Loc. No. 10). This is a small seasonally occupied 
dipterocarp resin-collecting camp on the left bank of a narrow creek 
about 10 km. southwest of Samnak Rabam (see above). The camp 
is surrounded by hills covered with mixed evergreen-deciduous-bam- 
boo forest in which relatively low trees, up to about 20 m. tall, pre- 
dominate. All specimens were collected within three hours' walk 
of camp. 

March 5-10— Ko Keow, 15°57'N, 99°26'E, alt. approx. 200 m. 
(fig. 1, Loc. No. 8). This large lumber camp is a few kilometers 
north of the upper Nam Mae Wong in the foothills of the Dawna 
Range. Approximately 100 workers are employed in intensive log- 
ging operations that began here about 10 years ago. The immediate 
vicinity of the camp is now farmland with a few scattered relicts of 
evergreen forest. In 1924 crab-eating macaques (M. fascicularis) 
were collected here for the American Museum of Natural History 
by A. S, Vernay. Although local residents report that M. fascicu- 
laris was common along the river up to about 20 years ago, these 
monkeys apparently no longer exist in the vicinity. Our hunters 
traveled by lumber truck within a radius of 10-15 km. of Ko Keow 
and collected four gibbons, one langur, one pigtail macaque, and one 
Assamese macaque, 

March 11— Khao Naw, 15°59'N, 99°51'E, alt. approx. 75 m. 
(fig. 1, Loc. No. 9). This sacred hill is an isolated peak surrounded 
by farmland. The hill, which is 42 km. northwest of Nakhon Sawan 
and about 1 km. east of the Bangkok-Chiang Mai highway, is cov- 
ered with evergreen forest and is inhabited by a semi-domesticated 
colony of long-tailed macaques (M. fascicularis) estimated to number 
300 individuals. The monkeys are fed by Buddhist monks who 
maintain a temple at the foot of the hill. I observed the monkeys 



18 FIELDIANA: ZOOLOGY, VOLUME 59 

assemble for food (boiled rice) at the sound of a gong that was rung 
by one of the monks. A Buddhist shrine in a cave half-way up the 
hill has been in existence for at least 100 years, and, according to 
tradition, monkeys have inhabited the hill throughout this period. 

March 13-15— Ban Mae Na Ree, 16°25'N, 99°23'E, alt. approx. 
150 m. (fig. 1, Loc. No. 6). This village of approximately 50 houses 
on the right bank of Khlong Suan Mak was established about two 
years ago by migrants from a northern area flooded by the Yan Hee 
Dam. Formerly the Ban Mae Na Ree region was covered by ever- 
green forest, but now most of the land is under cultivation and the 
forest is reduced to a 2 X 5 km. tract along the stream. All specimens 
were collected in this small relict tract of evergreen forest. 

March 17-26— Ban Mae Lamao, 16°48.5'N, 98°45.0'E, alt. ap- 
prox. 350 m. (fig. 1, Loc. No. 4). This large village of about 200 
houses is in the mountains between Tak and Mae Sot on the right 
bank of Huai Mae Lamao, a stream that flows northwestward to 
join the Sal ween River on the border between Thailand and Burma. 
The village is surrounded by cultivated fields. About three hours' 
walking distance (10-15 km.) to the south and west of the village are 
forested hills where our hunters collected five species of primates. 
At lower elevations in these hills deciduous dipterocarp forest pre- 
dominates; at higher elevations this is largely replaced by broadleaf 
evergreen forest; and on some of the highest ridges there are stands 
of pine. Interspersed at all elevations are restricted or extensive 
groves of bamboo. In April-July, 1924 K, G. Gairdner collected 
gibbons, langurs, and macaques about 20 km. northeast of Ban Mae 
Lamao (Chasen and Kloss, 1930, p. 63). 

March 28-30 — Huai Kwang Pah and Huai Wang Kwao, 
17°28'N, 98°50'E, alt. approx. 300 m. (fig. 1, Loc. No. 3). Huai 
Kwang Pah is a southward flowing creek that joins Mae Nam Ping 
(left bank) about 25 km. above the Yan Hee Dam. At the mouth 
of Huai Kwang Pah is a small settlement consisting of two or three 
bamboo huts. Huai Wang Kwao is a seasonally dry creek on the 
opposite (right) bank of Mae Nam Ping 1 or 2 km. upstream from 
Huai Kwang Pah. As a result of recent construction of the Yan Hee 
Dam, Mae Nam Ping is artificially broadened in this area, and the 
hills now rise abruptly on both sides of the river. The hill forests 
near the river are mixed bamboo and deciduous; higher on the hills, 



FOODEN: PRIMATES FROM THAILAND 19 

1 or 2 km. inland, tall evergreen trees predominate. In the vicinity 
of Huai Kwang Pah we collected two gibbons, two langurs, and two 
rhesus macaques, and near Huai Wang Kwao we obtained three 
langurs. Another party of our expedition hunters collected one rhe- 
sus macaque at Huai Ap Nang (fig. 1, Loc. No. 2), a creek about 
10 km. upstream (right bank) from Huai Kwang Pah. 

April 1-6— Ban Nam Lai Tai, 16°10'N, 99°20'E, alt. approx. 
300 m. (fig. 1, Loc. No. 7) . This village of about 75 houses is 1-2 km. 
west of Nam Lai (right bank) one of the headwaters of Khlong Klung. 
The village, which was established about five years ago, is surrounded 
by a cleared and cultivated area that formerly was covered with ever- 
green forest (Plate II, b). Expedition hunters collected one gibbon 
and eight crab-eating macaques (M . fascicularis) in evergreen forest 
on the foothills of the Dawna Range about 5 km. west of Ban Nam 
Lai Tai. In July, 1949 Colin Sanborn (1952, p. 7) collected one pig- 
tail macaque and two langurs in this same general area along the 
upper Khlong Klung. 

April 8-15— Ban Pong Nam Ron, 16°20'N, 99°18'E, alt. approx. 
200 m. (fig. 1, Loc. No. 5). This village of about 100 houses is on 
the left bank of Khlong Suan Mak a few kilometers east of the foot- 
hills of the Dawna Range. The foothill forests, where all of our speci- 
mens were taken, are bamboo and evergreen at lower elevations and 
predominantly evergreen higher up. Three gibbons, one langur, and 
one pigtail macaque (M. nemestrina) were taken at elevations of 
200-300 m. within five hours' walk west of the village. On a long 
overnight hunting trip four specimens of M. assamensis were taken 
about 25 km. west of the village at approximately 750 m. elevation. 

April 16-17— Bangkok. 

April 18 — Lop Buri; repeat visit to temple ruins to re-examine 
resident colony of M. fascicularis (see above) . 

April 20-24— Khlong Suan Mak (2 km. west of Ban Pong Nam 
Ron), 16°20'N, 99°17'E, alt. approx. 200 m. On our return visit to 
the Ban Pong Nam Ron area (see above) we camped about 2 km. 
west of the village on the left bank of Khlong Suan Mak immedi- 
ately at the base of two 100 m. high foothills of the Dawna Range 
(Plate III,a,b). One M. assamensis specimen was collected about 
3 km. west of camp; three M. nemestrina specimens and three M. 
assamensis specimens were collected on successive days about 6 km. 
west of camp, and two M. nemestrina specimens were collected about 



20 FIELDIANA: ZOOLOGY, VOLUME 59 

10 km. west of camp. The altitudes of these collection points range 
up to about 250 m. 

April 25-28 — Bangkok. Before leaving Thailand I received from 
Dr. Gershon Berkson two gibbon specimens that he collected on 
March 12, 1967 in evergreen forest about 30 km. east of Mae Sariang 
(18°19'N, 98°07'E; fig. 1, Loc. No. 1). 

April 30, 1967— Arrived Chicago. 



SPECIES ACCOUNTS 
Nycticebus coucang (Boddaert). Slow Loris. 

Tardigradus Coucang Boddaert, 1784 [1785], p. 67 — lectotype, the tailless 
Maucauco: Pennant, 1781, tab. 26, designated by Thomas, 1922, p. 433. 

Four slow lorises were taken at two localities incidental to the 
collection of other primates. There was no systematic program of 
night hunting for slow lorises. Each of the two localities is repre- 
sented by a different nominal subspecies of N. coucang. 

Measurements. — See Table 6. 

Nycticebus coucang bengalensis (Fischer). 

Loris bengalensis Fischer, 1804, Anat. Maki, p. 30 (work not seen; citation from 
Pocock, 1939, p. 166). 

Specimens collected. — TAK: Ban Mae Lamao, 1. 

Habits and habitats. — This solitary adult female was collected 
alive about 10 m. above the ground in an evergreen tree. The ani- 
mal was not pregnant, judging from gross examination of the opened 
uterus. The stomach contained whitish vegetable matter and hairs 
which match those of the animal's own fur. The hairs presumably 
were ingested as a result of self-grooming with the mandibular dental 
comb. Ingested hair that is inferred to be the result of using the 
teeth for grooming also has been found in the stomachs of African 
prosimians, Euoticus elegantulus and Galago demidovii, by Jewell and 
Gates (1969, p. 245). The intestine of this slow loris contained para- 
sitic nematodes. 

Nycticebus coucang tenasserimensis Elliot. 

Nycticebus tenasserimensis Elliot, 1912 [1913], vol. 1, p. 25. 

Specimens coi^ec^erf.— KAMPHAENGPHET: Ban Mae Na Ree, 
3 (1 infant in alcohol). 

Habits and habitats. — These three specimens (adult male, adult 
female, infant female with complete deciduous dentition) constituted 
a family group collected together about 5 m. above the ground in a 
clump of bamboo. The adult female was pregnant with an early 
embryo. The stomach of the adult male contained insect fragments 
and fragments of purplish fruit. 

21 



22 FIELDIANA: ZOOLOGY, VOLUME 59 

Table 6. — External measurements in adult specimens of Nycticebus coucang. 



Females 


Head and body 
(mm.) 


Relative 
tail length* 


Weight 
(kg.) 


Locality:' 








Mae Na Ree (6) 


335 


0.05 


0.90 


Males 








Locality: 

Mae Lamao (4) 
Mae Na Ree (6) 


345 
344 


0.04 
0.10 


0.85 
0.92 


> Figures in boldface type are locality numbers shown on map, 
2 Tail length ^ head and body length. 


Figure 1. 



Remarks. — This weakly defined subspecies is geographically and 
morphologically intermediate between Indo-Malayan N. c. coucang 
and Indo-Chinese N. c. bengalensis, as previously noted by Pocock 
(1939, p. 170). In. N. c. coucang (FMNH 98478, Malaya) a dark 
mid-dorsal band extends from the lumbar region to the crown and 
continues anteriorly and laterally as four dark streaks which connect 
with dark rings around the eyes and ears. In. N. c. bengalensis (Ban 
Mae Lamao; see above) the mid-dorsal band is paler and it extends 
forward only to the occiput, so that the crown is whitish and the 
dark rings around the eyes and ears are isolated. In. N. c. tenasseri- 
mensis, which is the intermediate subspecies, the mid-dorsal band 
narrows anteriorly and the four streaks leading to the eyes and ears 
are indistinct. Future revisory study may demonstrate that the 
range of variation in N. c. coucang should be extended to include 
diagnostic characters assigned to N. c. tenasserimensis, which would 
sink the latter nominal subspecies into the synonymy of the former. 
In this case, the geographic border between the Indo-Chinese range 
of N. c. bengalensis and the extended Indo-Malayan range of rede- 
fined N. c. coucang would coincide almost precisely with the border 
between the ranges of Macaca mulatta and M. fascicularis (see below) . 

Macaca fascicularis (Raffles). Crab-eating, Kra or Long-tailed 
Macaque. 

Simia fascicularis Raffles, 1821, p. 246 — for validity of this name, see Fooden, 
1964, p. 365, footnote 2. 

Specimens collected. — KANCHANABURI: Ban Tamrong Phato, 
8; Ban Huai Maenam Noi, 2; Ban Phu Toei, 2. UTHAI THANI: 
Kata Taek, 6 (1 skeleton). KAMPHAENG PHET: Ban Nam Lai 
Tai, 8 (1 in alcohol); Ban Mae Na Ree, 3 (1 skeleton). 



e 

5 



^e 



"^. '"1 







03 








• 




■^^M 










$ 


OJ-S 


00 


\a 


iti Tf 




-*. 1 


00 


o o 






o ' 


o 


' 1-J .-J 









« e 

w o 



t- t- 

o ^. ^ 

00 eo >o '- 

^ /vC t— 00 

"^ ^ CO 00 

<o co' eo «" 



(M 



OS ^'. 

t- o 



O lO 






bo ^ 

J las 



•2 flj .— "* 5* 

s a> (S "-' J3 

•H Pi H ^ P^ 

§ « § f« ^ 

"^-^ g hJ f^ o 

S cu S 5 g 

C ^ c« eS 5 



n. P 



w C ^ 

o (i> a 

"^ '"' ««-( 

S >» a> 

aS ^ 

a, « C 

w T) to 

CQ cU 3) 

Eti E-i M 



a s 

.S fl 

m m 

e £ 



3 3 

c<] CIS 

J3 X> 

3 3 

CO M 



23 



24 FIELDIANA: ZOOLOGY, VOLUME 59 

Sight records (see Itinerary and Locality Notes). — KANCHANA- 
BURI: Mae Nam Khwae Noi, right bank, about 10 km. below Ban 
Wang Kalang, 2 troops, Feb. 3, 1967. LOP BURI: Lop Buri, pro- 
tected colony, Feb. 24 and April, 18, 1967 (Plate II,a). UTHAI 
THANI: Khao Phatowee, protected colony, Feb. 25, 1967. NAK- 
HON SAWAN: Khao Naw, protected colony, March 10, 1967. 

Measurements. — See Table 7. 

Habits and habitats. — Troops of M. fascicularis were encountered 
with approximately equal frequency in bamboo and evergreen trees, 
and one or two troops also were encountered in deciduous diptero- 
carp trees (Table 8). Troop size varied from about seven to 100. 
Solitary individuals were encountered twice (one pregnant female, 
FMNH 99641; one adult male, FMNH 99651). In all specimens 
examined recognizable stomach contents consist exclusively of fruit 
pulp and seeds. However, the cheek pouches of one adult female 
(FMNH 99645) were stuffed with small dark unidentified snails. 

Five females collected between February 10 and March 3 were 
in various stages of pregnancy (estimated one to three months before 
term), and no visibly pregnant females of M. fascicularis were col- 
lected thereafter (Table 9) . The uteri of three females collected on 
April 4-5 apparently were undergoing post-partum involution; two 
of these were lactating and one was collected with a very young in- 
fant. A total of four lactating nonpregnant females, including two 
with young infants, was collected between March 14 and April 5. 
These observations suggest that M. fascicularis in Thailand has a 
birth peak in the period March-May. 

Pregnancy in lactating females evidently is not uncommon in 
M. fascicularis living under natural conditions (Table 9, FMNH 
99644, 99645), as predicted by Spiegel (1954, p. 261), who observed 
continuous lactation through successive births in captive animals. 
The precocious pregnancy of a juvenile female (Table 9, FMNH 
99646) estimated from dental evidence to be about three years old 
appears to be unusual in M. fascicularis (Spiegel, 1954, p. 230). 

Remarks. — The geographical and morphological inter-relation- 
ships of crab-eating macaques and rhesus macaques in the Indo- 
Chinese Peninsula are of special taxonomic and evolutionary interest. 
Previously, I presented evidence of morphological intergradation 
that tended to show that these two macaques are conspecific (Fooden, 
1964, p. 363) . This was in contrast to earlier usual allocation of these 
monkeys to separate subgenera. Further information is now avail- 



bo 


s. 


0) 

JS 


TS 

c 


-s 


s 


es 


^ 


S 


a> 


rn 


> 
o 


H 


•s 



•5 


.S-S 


5 


^t 


s 


»< SJ 


u 


+j eg 




-§1 




SI 



C C c 

o) (V S 

S (U $ 





.2* 


03 

3 


m 

3 



o o a) 



X!,o,Q boM-S-5'^'O'O boMbo 
^ ,n ^ 0) 0) oJ'C^'O'O 0) 0) 0) 



5 

s 

ei 

(V 

.a 

DO 
O, 

o 
o 
t3 



00 

pa 

■< 



.2 
S 

3 
(3 

CIS 
g- 
'■P 
so 



bfl M CQ .^ .^ 
C J5 J3 O O 



9^ 9^ rt rt c8 



I 0. C^ Z Z ^ ^ ;S CS ^ ^ ^ 

St CO bfl be — — !S "i _ .^ .^ .^ 



O fctj C C S £ 
C^ bo ^« u CJ 



S rt tS ni n> g g g 



,-1 - O 1-1 «o 
CO CO tH 



C<I CO l-H rH 

O U O y — . — . — 






25 



X 

^ d 

tn (-• ^—^ 
3 J3 . 

feXS 



3 
O 



tf 



c4ai 



o 

w 



X 


X 


X 


o 


o 


o 


00 


00 


00 


X 


X 


X 


lO 


o 


o 



X 



X X 



X X 



ift 00 »-i 

■^OOOOOOi-Hi-HrHin 

xxxxxxxx 


00 
X 


M 
tJ 

^ 


osoot-incoooiM 

^ ^ T-H CO CO IM '-H 

xxxxxxxx 


CO 
X 


is 


OOOWOIOOOO 
CgrJ<^eo;o;0>0(N 


o 

CO 


^ 



T3 






t-H r-( 1— 1 

^ ^ s 

O O £ 


1—1 


















s 
S 


s 

Cli 




s 




o 


I 


















fl '4 




CO 




i?i?l 


00 


J 
















B 


.2^ 

o ^ 




3 




<u c a) c 3 


3 
0) 




















"iJ 






bfl"^ bfl"^ «*-! 




4-> 

o 
















> 


c S 








C o G 0) *t- 
•^ bOtiS bo O 


















Oi 


.« o 


'5 


o 

-4-> 




O a; 


0) C 
















T3 


3 5 


o 


^ 






-C 3 


^ +^ 








4-9 








Is 






03 

o 


bio S ti S '§ 2 


boS £ 


i-s 








(3 
cU 

c 








■1^ 
cS 

c 


ti5 

O. w 


o 

3 


be 


s 


•+3 • '^ • bo g 
















Q. 


-M ^ 


'■2 


:^ 


cs S « S CxT, 

o S w S ij-^ 
PS ^ cs .t: (M 








-t-> 


+j 






-tJ 


-4J 




o 


■+J 


CO 


.t^T}< C 


c ^ 




Ci 


c 


'$ 




C 


c 


>-l 


'm 




— ' O ^^ O M g. 


"» ^^ « 


OS 




rt 


CD 






cS 


bS 


>> 


*4-t 




a 




rt 2 tS 2 cS . 


^ ^ M 

C8 . f- 




bo 


C 
bo 


C 

bo 


bO 

-4-3 


bo 


c 

bO 


c 


3 
to 


° "2 

bo C8 

is g 




s 


+-> 


c5 c 5 c-S 


C"S o< 


D, c3 


C8 


a 


a 


ca 


n! 


a 


a 






be 


s. 


^^ ^a ^i- 

U (-. u 

&H Ph Cl, 




o ^ ^ o o ^ ^ o 
;? J h:5 iz; :z; h:j J 2; 


o 







« 

bo 



g 

^ bO 

« f 



Q :§ :§ 1^ 



i-< en 

(V O) 

a-- 

.« bo 



'-I a 
Q a, 



bo 
cd 



s ^ 



_• \N- _• ? 3 3 w 0) to 



g s 

a) "^ 

aeo 



g T3 g : 



o<^" 



^-'^^^^^^^i'^^ 



O i-H .-H rH 







-<ll< -^ 




r^ 


■«-> ec 


CO 


CO 


CO i-H "-H 




fr 


is.c 


^ 


,c 


^jSJS'^'^^^"^ 


in 


> 


a ? 


cU 






a 

< 


3 

CO 
0) 

cfl 

C 


01-5 



cS 
O; g 

C g 

?^ "5 

r»> 50 

Xi JS 
O bo 

a '^ 

_ J3 
>^ bfl 

> +3 
'« 

-^ -S S 

GO Oi 

0) .*j> bo 
§ . 3 .C 

<u 'r^ .5 a 
o s o a> 

CO 'r^ CO '^ 
■" "~ ■> l-H 

^ CO 

i? 3 

CO O 
CO 3 

2.1 

bo u 

CO ~ 

>> J= 

o^ 

CO ^ 

CIS 



C "3 bO i5 



lo «Dooa>Oi-ieokrt«D 
to ^(ococo^co^co 



<1> Crt 

bO Oj3 

-NO) 

g 

o 



2 g 



26 



FOODEN: PRIMATES FROM THAILAND 27 

able from study of additional museum specimens and from study of 
specimens that were collected in Thailand with this specific problem 
in mind. As discussed below, the available evidence no longer ap- 
pears adequate to establish that crab-eating and rhesus macaques 
are conspecific. 

The distribution of crab-eating macaques is broadly Indo-Ma- 
layan, extending from Timor and the Philippine Islands westward 
and northward to southern Burma. The complementary, more 
northern distribution of rhesus macaques is broadly sub-Himalayan, 
extending from southern China westward to easternmost Afghanistan. 
The border between the ranges of these monkeys extends irregularly 
across the Indo-Chinese Peninsula approximately 1,500 km. from 
Akyab, Burma, to Hiae, South Vietnam (fig. 2) . Along this irregular 
front, which coincides with the margin of the mountain chains that 
extend southeastward from the Himalayas, crab-eating macaques 
inhabit low elevations up to about 500 m., and rhesus macaques in- 
habit higher altitudes. Although both monkeys are now absent from 
large parts of their former ranges as a result of progressively intensi- 
fying human activity, the pattern of complementarity remains clear 
(fig. 2). 

The most conveniently analyzed taxonomic character that dis- 
tinguishes crab-eating macaques from rhesus macaques is relative 
tail length (tail length-;- head and body length; flesh measurements). 
This proportion is comparable in adults and subadults with perma- 
nent premolars. It is not directly comparable in younger monkeys 
because relative tail length decreases ontogenetically up to about age 
four years, when permanent premolars erupt (Schultz, 1933, pp. 18, 
23). Individual and local variation of relative tail length in crab- 
eating macaques and rhesus macaques is graphed in Figure 3, which 
includes all proportions known from flesh measurements for adults 
and subadults collected north of the Strait of Malacca and east of 
the Brahmaputra River. 

There is no apparent overlap in the relative tail length of rhesus 
macaques and crab-eating macaques (fig. 3) . In the broad area shown 
in Figure 2, excluding three marginal localities discussed below, rela- 
tive tail length varies from 0.34 to 0.61 in rhesus macaques and from 
0.79 to 1.48 in crab-eating macaques. Similar limits of variation also 
apply in specimens examined from more distant parts of the ranges 
of these monkeys. For example, in eight rhesus macaques collected 
in Kashmir, about 2,000 km. west of the area studied, relative tail 
length varies from 0.32 (USNM 173812 9 ) to 0.48 (USNM 353187 9 ) ; 




Figs. 2, 3. Map showing distribution of Macaca mulatto and M. fascicularis 
in Southeast Asia, and graph showing variation of relative tail length. Sample 
areas of M. mulatta are designated by capital letters on map and graph; those of 
M. fascicularis are designated by small letters. 

Closed circles indicate localities from which adult and subadult specimens 
with external measurements are available, as follows: M. mulatta: A — Chungan 
Hsien (AMNH); 30 km. SW of Kiating (BMNH). B— Dening, Bawmwang, 
Htingnan, N'changyang (BMNH); Teng Yueh (MCZ). C— Dalu-Taga Hka, 
Nanyaseik, Hkampti-Heinsum, Moklok, Maungkan (AMNH); Homalin(BMNH); 
Changchang Pani (AMNH). D — Lamsokhang, Nangpoh, Rajapara (BMNH). 
E — Bishenpur, Kindat area, Yin (BMNH). F — Nam Yao, Hsipaw area 
(BMNH); Mandalay area, Mt. Popa (AMNH, BMNH). G— Backan (BMNH, 
MNHN); Muong Poun (AMNH); Phuqui (BMNH). H— NE of Toungoo, E of 
Toungoo, SE of Prome (BMNH). I— Huai Ap Nang (FMNH); Mae Nam Ping 
rapids (Kloss, 1917, p. 247); Pang Nam Un, Dan Sai district (USNM). J— Ban 
Mae Lamao (FMNH); 28 miles SE of Ban Umphang (AMNH). K— Dak Sut 
(USNM). 



28 



-I I I 



A ■ 
B 

















■ ■ OD 










■oK KB ■ oB 


mul a 


ffa 


G 


o 


CB ODO 

□ OQ 

■■ ao o 

K) 

a 


am 










a m ' a' 










a 


aa a a 








am a 










a 


m m 








a 


wo a m 
am 

■■■■■ am 








cm aimma m 








a 


o 






fascicular is 


m Jo 

OD 


m am 
a . 8 


m 








o 


B8BUUn8. 

CB ■ 
■ ■ ■ 
■ ■ ■ 

K»«B 



' O FEMAII 



m ..bcUg.B.a. 



.40 



.50 .60 .70 .80 .90 1.00 1.20 1.40 

REIATIVS TAll lEIIOTH 



M. fascicularis: a — Ban Mae Na Ree (FMNH). b — Ban Nam Lai Tai 
(FMNH). c-40-53 miles E of Ban Umphang (AMNH, BMNH). d— Kata 
Taek (FMNH). e— Ban Tamrong Phato (FMNH). f— Phu Phan (USNM); 
Phu Quoi (BMNH). g— Con Son (BMNH, USNM). h— Ko Chang, Ko Kram, 
Ko Kut (USNM). i— Pak Khlong Pran (Kloss, 1917, p. 289). j— Tenasserim 
area. King Island (BMNH). k— Sullivan Island, St. Matthew Island (BMNH); 
Pakchan River (AMNH, BMNH); Pak Nam Chumphon (Kloss, 1917, p. 289). 
I— Ko Phangan (Robinson and Kloss, 1914, p. 131); Ko Samui (BMNH; Robin- 
son and Kloss, 1914, p. 131). m— Trang, Ko Libong (USNM); Ko Tarutao 
(BMNH, USNM); Pulau Langkawi, Biserat, Penang Island, Ulu Ijok, Telom 
River, Changkat Mentri (BMNH). n— Pulau Redang (Kloss, 1911, p. 183). 
— Pulau Pintu, Kuala Lumpur vicinity (BMNH). p — Pulau Tioman (BMNH, 
USNM). q— Tanjong Panjair, Pulau Kaban (BMNH); Pulau Pemanggii 
(USNM); Pulau Awr (BMNH). r— Pulau Tinggi (BMNH). s— Punggol- 
Changi (BMNH); Pulau Karimon (BMNH, USNM); Pulau Durian-Pulau Sugi 
(Sody, 1949, Table 1; USNM); Pulau Bulan, Pulau Batam (BMNH); Pulau 
Garlang-Pulau Nguwal (Chasen, 1925, p. 93); Pulau Bintan (BMNH, USNM); 
Pulau Mapur (Robinson, 1916, p. 62). 

Open circles indicate marginal locality records without external measure- 
ments of adults or subadult, as follows: M. mulatta: BURMA — NE of Akyab 
(Anderson, 1878, p. xviii); NORTH VIETNAM— Vinh Linh vicinity (Dao Van 
Tien, 1962, p. 724). M. fascicularis: BURMA— Arakan (Khajuria, 1954, p. 109), 

Continued on p. SO 
29 



30 FIELDIANA: ZOOLOGY, VOLUME 59 

in nine crab-eating macaques collected in Mindanao, Philippine Is- 
lands, about 2,000 km. east of the area studied, relative tail length 
varies from 1.02 (USNM 125322 9 ) to 1.32 (FMNH 56491 cf). 

Conspicuous exceptions to the general discontinuity of relative 
tail length in rhesus and crab-eating macaques are evident in three 
series of three specimens each collected at (1) Dak Sut, South Viet- 
nam, (2) Ban Mae Na Ree (Plate IV, a) , and (3) Nam Mae Wong, 
Thailand (fig. 3, Table 10), all on the border between the ranges of 
these two monkeys. Pelage color in the Dak Sut series is rhesus-like, 
but somewhat drabber than usual. In two Dak Sut females the dor- 
sal surface is gray washed with yellowish anteriorly, becoming more 
golden posteriorly; in one subadult male the dorsal surface is darker 
grayish-brown washed anteriorly with yellowish and becoming al- 
most imperceptibly brighter posteriorly. Pelage color in the Ban 
Mae Na Ree series and in the Nam Mae Wong series is more nearly 
as in typical crab-eating macaques, ranging from golden brown an- 
teriorly, becoming slightly brighter posteriorly (FMNH 99658), to 
more or less uniformly pale yellowish brown (FMNH 99659) or uni- 
formly darker yellowish brown (FMNH 99657). Pelage color in two 
of the Nam Mae Wong specimens was described in detail previously 
(Fooden, 1964, p. 363) . Cranially, all three of these series have the 
rostrum relatively long and narrow as in typical M. fascicularis. 

In Thailand, crab-eating macaques with normal tail proportions 
have been collected very near to and interspersed between the two 
localities where aberrantly short-tailed series have been taken. These 
two localities, Mae Na Ree (16°25'N) and Nam Mae Wong (15° 
55 'N), are only about 50 km. apart, yet at Ban Nam Lai Tai (16° 
10 'N), almost exactly midway between them, five normal subadult 
and adult crab-eating macaques (relative tail length 0.96-1.22) were 
collected (Plate IV,b). Similarly, three subadults and adults taken 
at Kata Taek (15°28'N), about 50 km. south of Nam Mae Wong, 
also have normal tail proportions (relative tail length 0.94-1.18). 

Crab-eating macaques with relatively short tails also have been 
collected at two other localities along the border between the ranges 
of M. mulatta and M. fascicularis. In a subadult female (USNM 



Figures 2 and 3 continued. 

Elephant Point (Anderson, 1881, p. 63), Pegu district (Blanford, 1891, p. 22), 
Haungtharaw (Khajuria, 1954, p. 108); THAILAND— Ban Phu Toei (FMNH); 
Lat Bua Khao (USNM); LAOS— Plateau Bolovens (AMNH); SOUTH VIET- 
NAM— Ban Me Thuot (FMNH); Sontra Peak (USNM). 



FOODEN: PRIMATES FROM THAILAND 31 

Table 10. — External measurements in three series of macaques with relative 
tail lengths intermediate between those in typical rhesus and crab-eating 
macaques. 

Head and body Tail Relative 

Specimen No. Sex Age length length tail length 

(mm.) (mm.) 

Dat Sut (700 m.), January, 1961, Collected by B. Feinstein 

USNM 320780 9 adult 462 298 0.64 

USNM 320781 9 adult 401 289 0.74 

USNM 320782 rf- subadult' 435 322 0.74 

Ban Mae Na Ree (about 150 m.), March, 1967, Collected by J. Fooden 

FMNH 99657 d" adult 448 381 0.85 

FMNH 99658 9 adult 424 334 0.79 

FMNH 99659 9 adult 407 390 0.96 

Nam Mae Wong (1,000 ft., 800 ft.), February, 1924, Collected by A. S. Vernay 

AMNH 54677' 9 adult 400 385 0.96 

AMNH 54679' 9 adult 460 350 . 76 

BMNH 24.9.2.8* cf adult 470 380 0.81 

' Incompletely erupted canines and third molars. 

* Collected 40 miles east of Ban Umphang. 

* Collected 53 miles east of Ban Umphang. 

356968) collected in September, 1967 at Sontra Peak (about 240 m.), 
South Vietnam, relative tail length is about 0.80, judging from the 
dry skin; although field measurements of the specimen indicate a rel- 
ative tail length of 0.60 (total length 615 mm., tail length 230 mm.), 
these measurements appear to be unreliable. In two infants col- 
lected in January, 1932 at Plateau Bolovens, Laos, relative tail 
length is 0.76 (AMNH 87266 9 ; total length 492 mm., tail length 
212 mm.) and 1.17 (AMNH 87270 d'; total length 530 mm., tail 
length 286 mm.). 

It now seems clear that macaque populations morphologically in- 
termediate between typical mulatta and fascicularis are restricted to 
a very narrow altitudinal and latitudinal zone in the Indo-Chinese 
Peninsula. In this narrow zone the known occurrence of morpho- 
logical intermediates is further restricted to small, discontinuously 
distributed local areas. This thin and spotty distribution of inter- 
mediates suggests that rhesus macaques and crab-eating macaques 
are not fully or freely interbreeding, hence not conspecific. The most 
reasonable interpretation of local morphological intergradation be- 
tween M. viulatta and M. fascicularis now seems to be that the inter- 
grade populations are hybrids that have resulted from secondary 
contact and occasional fertile breeding between two species that pre- 
viously differentiated in geographic isolation (cf. Mayr, 1963, p. 369). 



32 FIELDIANA: ZOOLOGY, VOLUME 59 

Speculation that M. mulatta-M. fascicularis intermediates represent 
interspecific hybrids was previously offered by Hill (1964, p. 8), 
Bernstein (1966, p. 1,559), and Kuhn (1967, p. 30). 

Macaca mulatta (Zimmermann) . Rhesus Macaque. 

Cercopitheciis (Mulatta) Zimmermann, 1780, p. 195, 
Specimens collected. — TAK : Ban Mae Lamao, 1 (skeleton) ; Huai 
Ap Nang, 1 (skeleton); Huai Kwang Pah, 2 (1 in alcohol). 

Measurements. — See Table 7. 

Habits and habitats. — The specimen collected at Ban Mae Lamao, 
a mature lactating female, was in a troop of about 20 individuals 
about 20 m. above the ground in a grove of bamboo. The Huai Ap 
Nang specimen was a solitary male — mature but not senile, judging 
from tooth wear — encountered about 50 m. above the ground in an 
evergreen tree. Two immature specimens collected at Huai Kwang 
Pah were in a large troop estimated to contain 50 individuals. When 
first encountered this troop was less than 20 m. above the ground in 
a grove of bamboo; it subsequently fled to about 40 m. above the 
ground in the leafless canopy of nearby deciduous trees. 

Stomach contents in all three specimens examined appears to con- 
sist exclusively of fruit pulp and seeds. No insect parts or other ani- 
mal remains are discernible. However, the cheek pouches of one of 
these monkeys (Ban Mae Lamao) contained fragments of the egg- 
shell of a small passerine bird, in addition to several large flat bean- 
like seeds. Six large parasitic nematodes, up to 5 cm. long and 3 mm. 
in diameter, were extracted from the stomach of one specimen (Huai 
Kwang Pah). 

A large pinkish tumor-like skin growth was present on the head 
and shoulder of the female collected at Ban Mae Lamao. The thick, 
hairless growth extended from the right cheek to the right axilla, pass- 
ing over the right side of the neck and in front of the right shoulder. 

Remarks. — -See under M. fascicularis (above). 

Macaca nemestrina (Linnaeus). Pigtail Macaque. 
[Simia] Nemestrina Linnaeus, 1766, p. 35. 
Specimens collected. — KANCHANABURI: Ban Kerng Chada, 4; 
Chongkrong, 1; Ban Huai Maenam Noi, 7 (1 in alcohol). UTHAI 
THANI: Samnak Rabam (10 km. SE), 2. KAMPHAENG PHET: 
Ko Keow, 1; Ban Pong Nam Ron (8 km. W), 3 (2 skeletons); Ban 
Pong Nam Ron (10 km. NW), 1; Ban Pong Nam Ron (12 km. W), 2. 



J3-r 


op 


MM 




S-M 


, r^ ia CO 






^"^ 


1 (O 00 (C 



5 

'^ be 

0) — 



I I 






a 

< 



ffi o 



^- 



at 



OJ 



in la to 

00 U5 t- 
•^ iO Tf 



«o 






(M 

«o "- 

"« 00 «o 
oi lo 00 

lO •>*' Tf 



o 

OS N 



.-I "5 U5 

t- 00 ;o 

rj< CO Tj< 



bfi 

a 

OS 

S 



«g 


(M 00 

la Tf 


TJ^-" 


'^fc "•* 




o w ec 


Tj< Tf «o 


Tj< Tl" Tj< 


^ 






34 



FIELDIANA: ZOOLOGY, VOLUME 59 





Table 12. — Field observations of troop 


size and habitats 


in 




Macaca nemestrina. 








Estimated 


Kind of tree 


Estimated 


Date 


number 


in which 


height above 


(1967) 


Locality in 


troop 


observed 


ground (m.) 


Jan. 28 


Chongkrong 


1 


evergreen 


45 


Feb. 5 


Kerng Chada 


12 


evergreen 


15 


Feb. 5 


Kerng Chada 


20 + 


evergreen 


40 


Feb. 5 


Kerng Chada 


30 + 


evergreen 


35 


Feb. 16 


Maenam Noi 


20 


deciduous 


20 








{Xylia kerrii) 


Feb. 26 


Samnak Rabam (10 km. SE) 


20 


evergreen 


20 


March 9 


Ko Keow 


— 


— 


— 


April 12 


Pong Nam Ron, 10 km. NW 


40 


evergreen 


40 


April 20 


Pong Nam Ron, 8 km. W 


15 


evergreen 


40 


April 21 


Pong Nam Ron, 12 km. W 


— 


— 


— 


April 21 


Pong Nam Ron, 8 km. W 


20 


evergreen 


30 


April 23 


Pong Nam Ron, 8 km. W 


— 


— 


— 



Measurements. — See Table 11. 

Habits and habitats. — Eight of nine troops of M. nemestrina for 
which habitat notes were recorded were in evergreen trees about 
15-45 m. above the ground; one troop encountered was in deciduous 
trees about 20 m. above the ground (Table 12). Observed troop size 
varied from about 12 to 40, and one subadult male (canines and third 
molars erupting) was solitary. Although stomach contents consists 
predominantly of fruit pulp and seeds, fragments of caterpillars and 
adult insects were present in the stomachs of six of seven specimens 
examined. 

Five of seven sexually mature females collected were either preg- 
nant or lactating (Table 13) . Three females collected on February 16 
were lactating; two of these were taken with infants that probably 
were born in the preceding December or January. Two observed 
pregnancies — an early pregnancy in February and a late pregnancy 
in April — ^probably would have terminated in June or July. Two 
non-lactating, non-pregnant females with apparently mature repro- 
ductive organs (Table 13, FMNH 99674, 99687) were dentally im- 
mature. 

In a mature female collected southeast of Samnak Rabam (fig. 1, 
Loc. No. 11) the nose is asymmetrically cleft (Plate V,a) . The under- 
lying nasal bone in this specimen (FMNH 99685) also is asymmetri- 
cally deformed. 

Remarks. — M. nemestrina and M. assameyisis (see below) are 
superficially similar in size, relative tail length, and general color 



FOODEN: PRIMATES FROM THAILAND 



35 



Table 13. — Reproductive data in sexually mature females of 
Macaca nemestrina. 



FMNH Date Dental 

No. (1967) development 



99674 
99677 
99678 
99679 
99685 
99687 
99688 



Feb. 5 
Feb. 16 
Feb. 16 
Feb. 16 
Feb. 26 
April 12 
April 20 



M 3 erupting 

M3 

M3 

M3 

M3 

M2 

M3 



Reproductive 
condition 

Not pregnant 

Lactating 

Lactating, with infant' 

Lactating, with infant' 

Pregnant 

Not pregnant 

Pregnant 



Uterus, 

length X bread thX 

d-v diam. (mm.) 

35X13X7 
30X15X7 
30X14X8 
40X20X9 
65X30X23' 
30X11X7 
140X95X65* 



' Infant with deciduous second incisors; estimated age 1"^ months (Hurme, 
1960, p. 796). 

* Infant with deciduous first incisors; estimated age 1 month (Hurme, 1960, 

p. 796). 
' Blastocyst diameter 20 mm. 

* Sitting height of fetus 150 mm.; estimated conception age 4 months. 



(Plate V,b,c; Table 11). These two species may be readily distin- 
guished by the color of the dorsal surface of the tail, which is con- 
trastingly darker than the back in M. nemestrina (not in M. assa- 
mensis) and the clearly defined crown cap present in M. nemestrina 
(absent in M. assamensis). In addition, the tail in M. nemestrina is 
less thickly furred than in M. assamensis. The glans penis in M. 
nemestrina is bluntly bilobed, approximately as in humans; in M. as- 
samensis the distinctive arrowhead-shaped glans penis tapers distally 
to a narrow vertical crest. 

The geographic relationship between M. nemestrina in lowland 
areas of the Indo-Chinese Peninsula and M. assamensis in upland 
areas is generally similar to the relationship between lowland M. fas- 
cicularis and upland M. mulatta (fig. 2). However, the range of 
M. nemestrina extends farther northward than the range of M. fas- 
cicularis, and in the northern part of its range M. nemestrina, unlike 
M. fascicularis, is marginally sympatric with its upland counterpart. 
There is no evidence of hybridization between M. nemestrina and 
M. assamensis at localities where the two species occur together 
(Plate V,b,c). The sympatry and apparently complete reproductive 
isolation of M. nemestrina and M. assainensis indicates that the 
phylogenetic divergence of this pair of species from one another 
began earlier or proceeded more rapidly than the divergence of M. 
fascicularis and M. mulatta. 

The name Inuus leoninus Blyth (1863, p, 7; work not seen, cita- 
tion from Khajuria, 1954, p. 116) with type-locality restricted by 



36 



FIELDIANA: ZOOLOGY, VOLUME 59 



Pocock (1939, p. 60) to N[orth] Arakan, Burma, is available for pig- 
tail macaques in the Indo-Chinese Peninsula. Because they differ 
externally, cranially, and penially from pigtails in Malaya and Indo- 
nesia, subspecific recognition of pigtails in the Indo-Chinese Penin- 
sula is undoubtedly justified and specific recognition is a possibility, 

Macaca assamensis McClelland. Assamese Macaque. 
Macacus Assamensis McClelland in Horsfield [1840], p. 148. 

Specimens collected. — KANCHANABURI: Chongkrong, 2; Ban 
Muang Baw Ngam, 4 (1 skeleton); KAMPHAENG PHET: Ko 
Keow, 1; Ban Pong Nam Ron (5 km. W), 1; Ban Pong Nam Ron 
(8 km. W), 3; Ban Pong Nam Ron (25 km. W), 4 (2 skeletons); 
TAK: Ban Mae Lamao, 3. 

Measurements. — See Table 11. 

Habits and habitats. — Of 11 M. assamensis troops encountered, 
eight were in evergreen trees 15-50 m. above the ground, two were 
in bamboo trees 20-25 m. above the ground, and one, for which 
forest type was not recorded, was about 10 m. above the ground 
(Table 14) . Observed troop size varies from about 10 to 50 individ- 
uals. Stomach contents in ten specimens examined consists pre- 
dominantly of fruit pulp and seeds, although a few insect fragments 
are present in each of three specimens. The cheek pouch of one adult 
female (Ban Pong Nam Ron, 8 km, W) contained the severed head 
of an agamid lizard (Japalura sp.). 

Seven of nine sexually mature females collected were either preg- 
nant or lactating (Table 15). Of these, four specimens (all just ac- 





Table 14. — Field observations of troop 


size and habitats 


in 




Macaca 


assamensis. 










Estimated 


Kind of tree 


Estimated 


Date 






number 


in which 


height above 


(1967) 


Locality 




in troop 


observed 


ground (m.) 


Jan. 15 


Muang Baw Ngam 




12 


evergreen 


— 


Jan. 17 


Muang Baw Ngam 




24 


evergreen 


40-50 


Jan. 19 


Muang Baw Ngam 




20 


evergreen 


20 


Jan. 201 


Muang Baw Ngam 




20 


evergreen 


30 


Jan. 27 


Chongkrong 




20 


evergreen 


15 


Jan. 28 


Chongkrong 




50 


evergreen 


40 


March 8 


Ko Keow 




30 


— 


10 


March 25 


Mae Lamao 




25 


evergreen 


30 


April 14 


Pong Nam Ron, 25 kn 


i.W 


10 


bamboo 


25 


April 20 


Pong Nam Ron, 5 km. 


W 


10 


evergreen (teak) 


30 


April 23 


Pong Nam Ron, 8 km. 


w 


30 


bamboo 


20 



Sight record; no specimens collected. 



>> 


F 


-gx 


H 


.OJ3 










e 

sS 


0) ti 




■<-> J3 




P 


> 

1 



»-l O lO O O 03 

X I xxxxxxx 

I-H rHOOOit-Oi'— <1-t 









s s 


s e 












g g 


S 6 












o o 


o o 












t- ■'f 


■^ t> 












T-l f-H 


1— I I-H 












3 § 


§ 3 












-U -kJ 


-<^ -M 












<U (V 


(V 0) 








bfi 




(JO v^ •« 


««-l «H 








fl 




fl V-. <» 


«<-l Vl 








'S 




•^ o o 


o o 








ci 




rt -M -u 


+J +J 






c 


■M 


>j 


t; J3 J3 


J3 J3 








« 




« bO M 

J2 -s -s 


.5? .S? 

'S "53 






-S 


-tJ 


g 


*^ xi js 


J3 J3 






c 


o 




2 M bO 

.s .s 


b£ bfi 






o 


C 


X 

-4^ 

•? 


.S .S 






> 


-t-T 


IJ' '-3 '-M 








'■4^ 


CD 




rt CO w 


'm 'm 






u 


b£ 


1^'^' 


-M 4-> 


bfl 


bO 


3 


.s 


C fl 


.g 


,a 


T3 


u 


'•X3 


ti c4 c4 


ca cs 


+3 


t^ 


O 


O. 


cd 


a c c 


s c 


c3 


03 




-i-> 


■4-J 


^j M M 


b£ be 


■u 


-!-> 



3 



-2 



s .s '^ 









0.0.0.0. 
3 3 3 3 

^ ^ ^ >H 

0) <U (V ID 



C c^cococccococccooo 



in m 



cu 
Q 



;-; ^ J3 -fl 



C C c« CD 



a a a a o. 






o 






87 



38 FIELDIANA: ZOOLOGY, VOLUME 59 

quiring last molars) collected between March 25 and April 20 were 
in late stages of pregnancy. This may indicate an annual birth peak 
in the period April-June. 

Remarks. — See under M. nemestrina (above) . 

Macaca arctoides I. Geoffroy. Bear Macaque. 

Macacus arctoides I. GeofTroy [1831], p. 61 — for nomenclatural discussion, see 
Fooden, 1967a, p. 153; 1967b, p. 250. 

Specimens collected. — KANCHANABURI: Ban Muang Baw 
Ngam, 1; Chongkrong, 3. 

Measurements. — External measurements of an adult male col- 
lected at Ban Muang Baw Ngam are: head and body 524 mm.; tail 
51 mm.; weight 10.1 kg. 

Habits and habitats. — Two specimens taken at Chongkrong were 
members of a large band of about 50 individuals that was encoun- 
tered on the ground in an evergreen forest. In response to gunfire 
this band fled high into the trees. Another specimen collected at 
Chongkrong was in a troop of five individuals that also was encoun- 
tered on the ground in evergreen forest. No habitat notes were re- 
corded for the specimen collected at Ban Muang Baw Ngam. 
Stomach contents in all four specimens appears to consist exclusively 
of fruit pulp and seeds. Parasitic nematodes were present in the 
stomachs of three of the four specimens collected. 

Expedition hunters made a careful search for bear macaques at 
all localities visited. The small number of specimens collected re- 
flects a low incidence of encounters with the species. M. arctoides 
evidently is rarer than other species of primates in the area visited. 

Remarks. — The geographic range of M. arctoides generally con- 
forms to the upland pattern of distribution previously noted in 
M. mulatta and M. assamensis. However, the range of M. arctoides 
extends farther southward — as far as northern Malaya (Medway, 
1963, p. 63) — than that of either of the other two upland macaques. 
This may be related to the fact that M. arctoides, unlike M. mulatta 
and M. assamensis, has no lowland counterpart. 

In a recently published comprehensive study of behavior in M. 
arctoides, Bertrand (1969, p. 3) surmises that this macaque is con- 
specific with the Chinese stumptail, M. thibetana. No evidence is 
cited in support of this conclusion, which is not based on first-hand 
study of the Chinese stumptail and which erroneously assumes that 
M. thibetana is restricted to the high mountains of Szechwan (Ber- 



FOODEN: PRIMATES FROM THAILAND 39 

trand, 1969, p. 5), although its range actually extends 1,500 km. 
farther eastward to Kwangtung and Fukien (Mell, 1922, p. 10; 
AMNH 84472, Chungan Hsien, Fukien; Fooden, 1967a, Fig. 3). 
The only character, other than those common to the genus, that is 
known to be shared by M. arctoides and M. thihetana is the vestigial 
tail, a character that has evolved several times in macaques (M. arc- 
toides, M. fuscata, M. maura, M. sylvana, M. thihetana). M. arctoides 
and M. thihetana are much more strongly differentiated than M. 
arctoides and M. fuscata (the Japanese stumptail), the latter two of 
which Bertrand (1969, p. 6) treats as separate species. M. arctoides 
and M. thihetana differ strikingly in 1) color and texture of the pelage, 
2) color of the face, 3) extent of the naked facial skin, 4) morphology 
of the skull, and 5) structure of the glans penis and baculum (Fooden, 
1967a, p. 160; Dobroruka, 1967, figs. 1-4; Bertrand, 1969, p. 3). 
There is no indication of intergradation or interbreeding in specimens 
of M. arctoides and M. thihetana collected in Kwangtung, China, 
where their ranges are contiguous or marginally overlapping (Mell, 
1922, p. 10, pi. 1-2). Unless evidence of intergradation is discovered, 
it seems appropriate to continue to regard these two sharply differ- 
entiated stumptail macaques as separate species. 

Presbytis cristatus (Raffles). Silvered Leaf Monkey. 

Simia cristata Raffles, 1821, p. 244. 

Specimens collected. — KANCHANABURI: Ban Kerng Chada, 4; 
Ban Tamrong Phato, 3; Ban Huai Maenam Noi, 4. 

Measurements. — See Table 16. 

Hahits and hahitats. — All five troops encountered of P. cristatus 
were in evergreen forest. Four troops were high in the trees, about 
40-50 m. above the ground, and the fifth was about 15 m. above the 
gi'ound. The estimated number of individuals in each of these five 
troops was: 9, 10, 12, 24, 30. All stomachs examined appear to con- 
tain only leaf pulp. The volume of stomach contents in mature 
specimens is impressive, ranging from about 500 to 775 ml. 

Of four sexually mature females examined in the period Febru- 
ary 12-15, one was lactating and collected with an infant probably 
born in the preceding December, two were pregnant and prob- 
ably would have delivered in April or May, and one was neither 
pregnant nor lactating (Table 17). The non-pregnant, non-lactat- 
ing female was dentally immature. 



.£ M 



rt 



e<3 i-< 



1 



« S 

►2 'CI 
ffi o 



1—1 «o 

o o 



0=1 



03 a 



>=i G 










P 00 


00 


rr 


to «£> 

1 1 

00 o 


oo" 


to" 




«3 


lO 


;o to 'O 



55 ^ 






0)^ Tj< «0 



o o 



•§s 






rtg 


OS 


o 


(M 


(M 


TS""^ 


lO 


"^ 


OS >> 


00 

o 





OS «D 



-5 S g 



P5 



2^^ 



;! .2 1 ^ ?: E: 



■o tS cS 



O 






W5 ^ 00 

-^ O -N 

■^ 4J ^ 

"« n' ►£ 

§ O c S 

.2 £ S 2J 

« W H S 



40 



, FOODEN: PRIMATES FROM THAILAND 41 

Table 17. — Reproductive data in sexually mature females of 
Presbytis cristatus. 

Uterus, 
Date FMNH Dental Reproductive length X breadth X 

(1967) No. development condition d-v diam. (mm.) 

Feb. 12 99705 M3 Pregnant' 150X75x55 

Feb. 12 99706 M3 Pregnant' 120X55X20 

Feb. 12 99707 M 3 and C erupting Not pregnant 30X15X6 

Feb. 15 99708 M3 Lactating; with infant' 45X18x6 

'Sitting height of fetus 140 mm., tail length 260 mm.; estimated conception 

age 3 months. 
J Sitting height of fetus 85 mm., tail length 130 mm.; estimated conception 

age 2 months. 
» Female infant, bufTy pelage, deciduous M 1 erupting; estimated age 2 

months. 

Remarks. — Dorsal pelage color is dark brownish-gray to blackish 
in adult specimens collected of P. cristatus and pale buflfy-gray in 
adult specimens of P. phayrei (see below). P. cristatus in western 
Thailand lacks the large sharply defined whitish mouth patch that 
is conspicuous both in P. phayrei (Plate VI, a), its neighbor to the 
north, and P. ohscurus, its neighbor to the south. 

Judging from known collecting localities, the range of P. cristatus 
in western Thailand and eastern Burma extends northeast only to the 
edge of the Dawna Range. Three localities recorded in the present 
report, all along Mae Nam Khwae Noi between the Dawna Range 
and the Bilauktaung Range, define the eastern boundary of the known 
range of the species. Thai specimens of P. cristatus collected by 
K. G. Gairdner that are listed by Pocock (1935, p. 953) were ob- 
tained in the immediate vicinity of Ban Huai Maenam Noi, one of 
the localities included in the present report. Burmese specimens of 
P. cristatus listed by Pocock (1935, p. 953) were collected at Ye Forest 
(approx. 15°29'N, 97°55'E) and Nwalabo Taung (13°54'N, 98°30'E), 
both of which are southwest of the Dawna Range. 

In the interior of the Dawna Range P. cristatus evidently is re- 
placed by P. phayrei (see below). Judging from 43 specimens col- 
lected at ten localities (Table 1), these two species of langurs are 
allopatric, and no other species of langur apparently occurs at any 
of the localities visited. Present evidence does not support earlier 
suggestions (Pocock, 1939, p. 146; Ellerman and Morrison-Scott, 
1951, p. 204) that as many as four species of langurs — P. cristatus 
{= Tra^hypithecus pyrrhus atrior: Pocock), P. phayrei, P. ohscurus, 
P. jemoralis — are sympatric in this area. 



42 



FIELDIANA: ZOOLOGY, VOLUME 59 



If subspecific distinction of P. cristatus in western Thailand and 
eastern Burma from eastern and southern populations of the species 
is justified, the name [Pithecus pyrrhus] atrior Pocock (1928, p. 673), 
with type-locality Ye Forest, Burma, is available. 



Table 18.- 



Date 

(1967) 

Jan. 13 
Jan. 13 
Jan. 13 
Jan. 14 
Jan. 27 
March 18 
March 21 
March 29 



FMNH 

No. 

99694 
99695 
99696 
99697 
99700 
99725 
99729 
99733 



-Reproductive data in sexually mature females of 
Presbytis phayrei. 

Uterus, 
Dental Reproductive length X bread thX 

development condition d-v diam. (mm.) 



M 3 Lactating 

M 3 Not pregnant 

M 3 Not pregnant 

M 3 Lactating 

M 3 Not pregnant 

M 3 Not pregnant 

M 3 Lactating 

M 3 Lactating; with infant^ 



30X19X7 

30X15X6 

35X21X9 

80X40X261 

40X19X9 

35X18X8 

35X17X8 



1 No blastocyst grossly visible in uterine cavity; probably an early stage in 

postpartum involution. 

2 Infant FMNH 99734, estimated age 1 month; see Table 19. 

Presbytis phayrei Blyth. Phayre's Leaf Monkey. 
Presbytis Phayrei Blyth, 1847, p. 733. 

Specimens collected. — KANCHANABURI: Ban Muang Baw 
Ngam, 5 (1 skeleton) ; Chongkrong, 3. UTHAI THANI: Kata Taek, 
11. KAMPHAENG PHET: Ko Keow, 1; Ban Pong Nam Ron, 1. 
TAK: Ban Mae Lamao, 6; Huai Wang Kwao, 3; Huai Kwang Pah, 2. 

Measurements. — See Table 16. 

Habits and habitats. — All ten troops of P. phayrei for which field 
notes were recorded were encountered in evergreen forest. Four 
troops were very high in the canopy, about 40-50 m. above the 
ground; six troops were about 15-30 m. above the ground. The esti- 
mated number of individuals in each of these ten troops is: 3, 5, 6, 
10, 10, 10, 20, 20, 30, 30. Stomach contents in all specimens exam- 
ined appear to consist exclusively of leaf pulp. In mature specimens 
the volume of stomach contents varies from about 500 to 775 ml. 



Table 19. — Developmental data in infants of Presbytis phayrei. 



Date 

(1967) 



FMNH 
No. 



Feb. 28 99721 

Feb. 28 99722 

March 18 99728 

March 29 99734 



Dental 
Sex development 

cf Dec. M 2 erupting 

9 Dec. M 2 erupting 

cf Dec. M 2 erupting 

d^ Dec. I 2 erupting 



Estimated age 
Pelage in months 



Buffy-^gray 
Buffy-»gray 
Buffy-^gray 
Buffy 



FOODEN: PRIMATES FROM THAILAND 43 

Reproductive tracts were examined in 7 of 14 sexually mature 
females collected (Table 18). None of these seven was pregnant. 
Four infant specimens of P. phayrei lack permanent molars and 
therefore presumably are less than one year old (Table 19). These 
infants probably were born in the preceding September (2 infants), 
October (1 infant), and February (1 infant). Although the data are 
meager, the absence of pregnancies in seven female specimens exam- 
ined of P. phayrei contrasts strikingly with the 50 per cent incidence 
of pregnancies in four specimens of P. cristatus collected during the 
same period (Table 17) . This and the apparent discrepancy between 
these species in probable months of birth inferred for fetuses and in- 
fants may indicate that P. phayrei and P. cristatus have different 
breeding seasons in western Thailand (Table 4) . 

Remarks. — Examination of specimens collected of P. phayrei re- 
veals that Pocock's (1939) otherwise excellent key and discussion 
are misleading in two respects concerning this species. The white 
eye ring may be incomplete medially in P. phayrei (Plate VI, a) ; this 
condition is not restricted to P. obscurus, as indicated by Pocock 
(1939, p. 138). Color of the pubic region is sexually dimorphic in 
P. phayrei (Plate VI, b), as in P. cristatus (Trachypithecus pyrrhus 
atrior: Pocock); Pocock's (1939, p. 121) key incorrectly implies that 
females of P. phayrei lack the distinctive pale pubic patch. 

Geographic interrelationships of P. phayrei and P. cristatus are 
discussed in the account of P. cristatus (above). If subspecific dis- 
tinction of western Thai P. phayrei from northern populations of the 
species is justified, the name Presbytis crepuscula Elliot (1909, p. 271), 
with type-locality Mulayit Taung, Burma (near the Thai-Burmese 
border), is available. 

Hylobates lar (Linnaeus). White-handed Gibbon. 
Homo lar Linnaeus, 1771, p. 521. 

Specimens collected. — KANCHANABURI: Ban Tamrong Phato, 
2; Chongkrong, 4 (1 in alcohol, 1 skeleton); Ban Muang Baw Ngam, 
4 (1 in alcohol); Ban Kerng Chada, 1. UTHAI THANI: Kata 
Taek, 4. KAMPHAENG PHET: Ko Keow, 1; Khlong Tawai 
(approx. 10 km. NW of Ko Keow), 2; Ban Nam Lai Tai, 1; Ban 
Pong Nam Ron, 3. TAK: Ban Mae Lamao, 3; Huai Kwang Pah, 2 
(1 in alcohol). MAE HONG SONG: Mae Sariang (30 km. E), 2. 

Measurements. — See Table 20. 

Habits and habitats. — All 19 gibbon groups encountered were in 
evergreen forest. Of these, 13 groups were about 20 m. above the 



44 



FIELDIANA: ZOOLOGY, VOLUME 59 



Table 20. — External measurements 


in adult specimens of Hylobates lar. 




Females 

A 




Males 




Head and 


Weight 


Head and Weight 


Locality! 


body (mm.) 


(kg.) 


body (mm.) (kg.) 


Kwang Pah (3) 


442 


4.50 


— — 


MaeLamao (4) 


— 


— 


462 6.0 


Pong Nam Ron (5) 


442 


5.55 


411* 3.53* 


Nam Lai Tai (7) 


— 


— 


440 5.2 


Ko Keow (8) 


377,' 438 


3.62,^5.7 


447,467 6.1,5.8 


KataTaek(lO) 


438 


4.5 


435, 450 4.97, 5.65 


Kerng Chada (13) 


— 


— 


416* 4.47* 


Tamrong Phato (14) 


428 


4.60 


438 5.88 


Muang Baw Ngam (15) 


420 


4.4 


432 — 


Chongkrong (16) 


440 


4.6 


446, 447 5.4, 6.2 



^ Figures in boldface type are locality numbers shown in map. Figure 1. 
* Subadult, canines not quite fully erupted. 
3 Subadult, canines half erupted. 



ground, three groups were 30-40 m. above the ground, and one group 
was about 60 m. above the ground; height observations were not re- 
corded for two groups. In 14 troops the number of individuals per 
troop ranged from two to five, as follows: five troops of two; three 
troops of three; four troops of four; two troops of five. Three larger 
gibbon groups (one of seven individuals; one of about 10 individuals; 
one of about 15 individuals) also were observed, but these probably 
represent transient encounters between two or more troops with 
partly overlapping territories (Ellefson, 1967, p. 30). The color com- 
position of gibbon troops observed during the expedition is reported 
in detail in a general article on color-phase in gibbons (F'ooden, 1969, 
p. 632). 

Stomach contents in specimens examined appears to consist ex- 
clusively of fruit pulp and seeds. Food fragments in the stomachs 
of gibbons are coarser than in other primates collected. Some pieces 
of fruit in gibbon stomachs measure 3x2x1.5 cm. (FMNH 99760), 
about twice the dimensions of the largest fragments noted in other 
primates. Gibbons evidently chew their food less thoroughly than 
macaques and langurs (cf. Carpenter, 1940, p. 86; Ellefson, 1967, 
p. 103). 

Six of eight sexually mature females examined were lactating 
(Table 21). Five of these were observed with young infants: four 
infants were collected. These ranged in age from about one to 12 
months. None of these eight sexually mature females was pregnant, 
judging from gross examination of the uteri. 



FOODEN: PRIMATES FROM THAILAND 



45 



Table 21. — Reproductive data in sexually mature females of Hylobales lar. 



Date FMNH Dental 

(1967) No. development 

Jan. 15 99736 M 3, C 

Jan. 27 99741 M 3, C 

Feb. 10 99745 M 3, C 

Feb. 28 99747 M 3, C 

March 7 99751 M 3, C 

March 8 99752 M 3; C erupting 

March 29 99757 M 3, C 

April 10 99760 M 3, C 



Reproductive 
condition 



Uterus, 
length X bread thx 
d-v diam. (mm.) 



Lactating, with infant' 26 X 17 X 10 

Lactating, with infant' 28 X 17 X9 

Not lactating, not pregnant 24 X 19 X9 
Lactating with infant' (> 14) X 18 X6 

Lactating 25X20X5 

Not lactating, not pregnant 24 X 17 X 10 
Lactating, with infant^ — 

Lactating, with infant* 28 X 18 X 12 



' Infant FMNH 99737, deciduous I 1-2, estimated age 2 months. 

' Infant FMNH 99742, deciduous I 1 erupting, estimated age 1 month. 

* Infant FMNH 99746, deciduous M 2, estimated age 1 year. 

* Infant FMNH 99758, deciduous M 1, estimated age 3 months. 

* Infant not collected. 



In pale-phase gibbon skins collected, the terminal third of most 
hairs is speckled with minute black spots. Under the microscope 
these dark bodies appear to be colonies of the fungal infection black 
piedra. 

Remarks. — Gibbons in northern Thailand, hitherto included in 
H. lar entelloides I. Geoffroy, 1842 (p. 717; type-locality Malay Pen- 
insula, 12°N) recently have been allocated to a newly proposed sub- 
species, H. lar carpenteri Groves, 1968 (p. 625; type-locality Doi 
Inthanon, Thailand). The geographic range of the proposed new 
subspecies in northern Thailand is said to extend southwest as far as 
Doi Inthanon (IS'^SS'N, 98°28'E), and the range of redefined en- 
telloides in southern Burma and Thailand is said to extend north as 
far as Myawadi, Burma (16°41'N, 98°31'E). Accordingly, of 11 gib- 
bon localities reported in the present paper, eight are in the range of 
redefined H. lar entelloides (fig. 1, Loc. No. 5, 7, 8, 10, 13-16), and 
three (Loc. No. 1, 2, 4) are in the postulated zone of intergradation of 
carpenteri and redefined entelloides (Groves, 1968, p. 626). 

In the original description of carpenteri, this subspecies is distin- 
guished from redefined entelloides on the basis of three pelage char- 
acters, as follows: 1) dark-phase darker in carpenteri ("very dark 
chocolate brown") than in entelloides ("dark brown"), pale-phase 
paler in carpenteri ("creamy-white") than in entelloides ("honey- 
coloured"); 2) base of dorsal hairs paler than tip in carpenteri (both 
color phases), base and tip approximately uniformly colored in en- 
telloides; 3) interscapular hair length 79-103 mm. in carpenteri, 
29-56 mm. in entelloides. These diagnostic characters were derived 



46 



FIELDIANA: ZOOLOGY, VOLUME 59 



Table 22. — Color and hair length variation in 27 Hylobates lar 
specimens collected. 



Rank in 

pale-dark 

series FMNH No.; sex 



1 (whitish) 

2 

3 

4 

5 

6 

7 

8 

9 
10 
11 
12 
13 
14 
15 
16 



99759 cf 
99741 9 
99761 d^ 

99763 d" 

99764 9 
99756 d" 

99752 9 

99744 d^ 

99745 9 
99743 d^ 

99739 d' 
99749 d" 

99753 d^ 
99748 cf 
99738 d' 

99740 d' 



17 (cinnamon) 99760 9 



1 (pale 

chocolate) 99754 d" 

2 99755 d" 

3 99747 9 

4 99757 9 

5 99751 9 

6 99750 d" 

7 99762 9 

8 99746 d" 

9 (blackish) 99736 9 



Age 



Interscapular 

hair length 

(mm.) 



Pale-phase 



adult 

adult 

subadult 

adult 

adult 

subadult 

subadult 

adult 

adult 

subadult 

adult 

adult 

adult 

adult 

adult 

adult 

adult 



65 
80 
75 
80 
85 
75 
65 
85 
50 
60 
80 
80 
60 
70 
50 
55 
80 



Dark-phase 



infant 

adult 

adult 

adult 

adult 

adult 

juvenile 

infant 

adult 



85 
70 
70 
60 
70 
60 
85 
35 
75 



Loc. 
No.> 



7 

16 

5 

1 

1 

4 

8 

14 

14 

13 

16 

10 

8 

10 

15 

16 

5 



4 

4 

10 

3 

8 

4 

5 

10 

15 



Hypothetical 
identification 
based on 
locality* 



entelloides 

entelloides 

entelloides 

car.-ent. 

car.-ent. 

car.-ent. 

entelloides 

entelloides 

entelloides 

entelloides 

entelloides 

entelloides 

entelloides 

entelloides 

entelloides 

entelloides 

entelloides 



car.-ent. 

car.-ent. 

entelloides 

car.-ent. 

entelloides 

car.-ent. 

entelloides 

entelloides 

entelloides 



^ Locality numbers as in map, Figure 1. 

' Notation entelloides indicates that collecting locality is in postulated range 
of redefined H. I. entelloides; notation car.-ent. indicates that collecting 
locality is in postulated zone of intergradation of H. I. carpenteri and rede- 
fined H. I. entelloides (Groves, 1968, p. 626). 



from study of 163 skins assigned to carpenteri and an unspecified 
number assigned to entelloides (Groves, 1968, p. 626). 

The evidence of 26 dry skins in the present collection (Table 22) 
indicates that carpenteri and entelloides are less distinct than origi- 
nally postulated. Dorsal pelage color in 17 pale-phase specimens 
varies more or less continuously from whitish to cinnamon, and dor- 
sal color in nine dark-phase specimens varies from pale chocolate to 
blackish. In specimens of both color-phases collected at localities 



FOODEN: PRIMATES FROM THAILAND 47 

within the geographic range assigned to redefined entelloides, the 
range of individual variation includes diagnostic pelage colors of both 
carpenteri and redefined entelloides. In the postulated zone of inter- 
gradation of carpenteri and entelloides, pale-phase and dark-phase 
specimens average slightly paler than those collected in the range of 
redefined entelloides. 

Basal and distal portions of the dorsal hairs are approximately 
uniformly colored in all 17 pale-phase specimens collected. In two 
of nine dark-phase specimens the bases of dorsal hairs are pale gray- 
ish and the distal portions are contrastingly dark brown; one of these 
specimens (FMNH 99751 9 ) was collected within the range of rede- 
fined entelloides, and the other (FMNH 99754 cT) was collected in 
the presumptive carpenteri-entelloides intergradation zone. In two 
dark-phase specimens (FMNH 99736 9, 99746 cT) collected in the 
range of redefined entelloides the dorsal hairs are approximately uni- 
formly dark from base to tip. In the remaining five dark-phase 
specimens the basal portions of dorsal hairs are slightly paler (brown- 
ish-gray) than the distal portions (brown). 

Interscapular hair length in all but three specimens in the present 
collection is greater than the maximum postulated for redefined en- 
telloides. In nine specimens, collected both in the range of entelloides 
and in the presumptive carpenteri-entelloides intergradation zone, in- 
terscapular hair length is within the range postulated for carpenteri. 
This character also appears to be highly variable within the geo- 
graphic range allocated to carpenteri; Groves (1968, p. 627) notes that 
interscapular hair length in one lowland specimen assigned to car- 
penteri is 24 mm. less than the minimum specified for that nominal 
subspecies. The discrepancies noted above indicate that final deter- 
mination of the validity and significance of the distinction between 
carpenteri and redefined entelloides will require further detailed com- 
parison of more extensive series of specimens. 




Plate I, a. Ban Muang Baw Ngm, evergreen forest on hillside, miners' 
quarters at base of slope (foreground). 



48 




Clj 

'S 

Eh 



M 

o 
43 



49 




Plate I, c. Khao Phatowee, detail view showing vegetation growing on 
precipices. 



50 




c 

O 

a 



3 

pq 

O 






61 




> 

IS 



52 




a 



58 




Plate III, b. Khlong Suan Mak, mixed evergreen and bamboo forest near 
camp. 



54 




Plate IV, a. Macaco fascicularis, short-tailed series collected at Ban Mae 
Na Ree. 



66 




Plate IV, b. Macaca fascicularis, typical long-tailed series collected at Ban 
Nam Lai Tai, about 50 km. south of Ban Mae Na Ree. 



-56 




Plate V, a. Macaca nemestrina, cleft nose in female collected 10 km. south- 
east of Samnak Rabam. 




Plate V, b. Macaca nemestrina female collecitd un mi 8 km. west of Ban 
Pong Nam Ron, April 20, 1967. 




Plate V, c. Macaca assamensis, female collected about 5 km. west of Ban 
Pong Nam Ron, April 20, 1967. 

67 




Plate VI, a. Presbytis phayrei, male (left) and female (right) collected at 
Chongkrong; note medially incomplete eye rings. 




Plate VI, b. Sexual dimorphism in Presbytis phayrei, female (left, with 
whitish pubic patch) and male (right) collected at Chongkrong. 



58 



REFERENCES 



Anderson, J. 

1878 (1879). Anatomical and zoological researches. . . . Bernard Quaritch, 
London. 

Bernstein, I. S. 

1966. Naturally occurring primate hybrid. Science, 154, pp. 1,559-1,560. 

Bertrand, M. 

1969. The behavioral repertoire of the stumptail macaque. Bibl. Primat., 
No. 11. S. Karger, Basel. 

BODDAERT, p. 

1784 (1785). Elenchus animalium, vol. 1. C. R. Hake, Rotterdam. 

Blyth, E. 

1847. Supplementary report of the Curator of the Zoological Department. 
Jour. Asiatic Soc. Beng., 16, pp. 728-737. 

Carpenter, C. R. 

1940. A field study in Siam of the behavior and social relations of the gibbon 
{Hylobates lar). Comp. Psychol. Monogr., 16, pp. 1-212. 

Chasen, F. N. 
1925. Notes on the fauna of Pulau Galang, Rhio Archipelago. Jour. Malay. 
Br. Roy. Asiatic Soc, 3, pp. 92-97. 

1940. A handlist of Malaysian mammals. Bull. Raffles Mus., 15, i-xx, pp. 
1-209. 

Chasen, F. N. and C. B. Kloss 

1930. On mammals from the Raheng District, western Siam. Jour. Siam Soc. 
Nat. Hist. Suppl., 8, pp. 61-78. 

Dao Van Tien 

1962. Materiaux sur la faune des v6rtebr6s de Vietnam. Zool. Z., 41 , pp. 724- 
735. 

DOBRORUKA, L. J. 

1967. tlber den "Blassgesichtsmakak" nebst einer tibersicht iiber die Art Ma- 
caca speciosa (F. Cuvier 1825). Milu, 2, pp. 305-312. 

Ellefson, J. O. 

1967. A natural history of gibbons in the Malay Peninsula. Ph.D. Thesis. 
University of California, Berkeley. 

Ellerman, J. R. and T. C. S. Morrison-Scott 

1951. Checklist of Palaearctic and Indian mammals. British Museum (Natural 
History), London. 

59 



60 FIELDIANA: ZOOLOGY, VOLUME 59 

Elliot, D. G. 

1909. Descriptions of apparently new species and subspecies of the genera 

Callicebus, Lagoihrix, Papio, Pithecus, Ceropithectis, Erythrocebtis, and Pres- 

bytis. Ann. Mag. Nat. Hist., (8) 4, pp. 244-274. 
1912 [1913]. A review of the primates, vol. 1. American Museum of Natural 

History, New York. 

FOODEN, J. 

1964. Rhesus and crab-eating macaques: intergradation in Thailand. Science, 
143, pp. 363-365. 

1967a [1966]. Identification of the stumptailed monkey, Macaca speciosa 
I. Geoffroy, 1826. Folia primat., 5, pp. 153-164. 

1967b. Macaca fiiscata (Blyth, 1875): proposed conservation as the name for 
the Japanese macaque (Mammalia). Bull. zool. Nomencl., 24, pp. 250-251. 

1969. Color-phase in gibbons. Evolution, 23, pp. 627-644. 

Geoffroy Saint-Hilaire, I. 

1831. Mammiferes. In Belanger, C, ed. Voyage aux Indes-Orientales . . . 

Zoologie, pp. 1-160. Arthus Bertrand, Paris. 
1842. Sur les singes de I'ancien monde, sp^cialement sur les genres Gibbon et 

Semnopitheque. Compt. Rend. Acad. Sci., Paris, 15, pp. 716-720. 

Groves, C. P. 

1968. A new subspecies of white-handed gibbon from northern Thailand, Hylo- 
bates lar carpenteri new subspecies. Proc. Biol. Soc. Wash., 81, pp. 625-627. 

Gyldenstolpe, N. 

1916 (1917). Zoological results of the Swedish expedition to Siam 1911-1912 
and 1914-1915. V. Mammals II. K. Sven. Vet. Akad. Handl. (n.f.), 57 (2), 
pp. 1-59, 

Hartman, C. G. 

1932. Studies in the reproduction of the monkey Macacus (Pithecus) rhesus, 
with special reference to menstruation and pregnancy. Contr. Embryol., 
Carnegie Inst. Wash., 23, pp. 1-161. 

Hill, W. C. O. 

1964. On the identification of monkeys. Lab. Prim. Newsl., 3, pp. 8-9. 

Horsfield, T. 

1840. Communication to the Society of Mr. McClelland's list of Mammalia, 
and birds collected in Assam. Proc. Zool. Soc. London, 1839, pp. 146-167. 

Hurme, V. O. 

1960. Estimation of monkey age by dental formula. Ann. N. Y. Acad. Sci., 
85, pp. 795-802. 

Jewell, P. A. and J. F. Gates 

1969. Ecological observations on the lorisoid primates of African lowland 
forest. Zool. Afr., 4, pp. 231-248. 

Khajuria, H. 

1954. Catalogue of mammals in the Indian Museum (Zool. Surv.) II. Pri- 
mates: Cercopithecidae. Rec. Indian Mus., 52, pp. 101-127. 

Kloss, C. B. 

1911. On a collection of mammals and other vertebrates from the Trengganu 
Archipelago. Jour. Fed. Malay States Mus., 4, pp. 175-212. 



FOODEN: PRIMATES FROM THAILAND 61 

KUHN, H.-J. 

1967. Zur Systematik der Cercopithecidae. In Starck, D., R. Schneider, and 
H.-J. Kuhn, eds. Progress in primatology, 1, pp. 25-46. Gustav Fischer 
Verlag, Stuttgart. 

Larsen, K. 

1962. Preliminary report on the Thai-Danish botanical expedition to the Kan- 
chanaburi Province 1961-62. Nat. Hist. Bull. Siam Soc, 20, pp. 109-119. 

Linnaeus, C. 

1766. Systema naturae, 12th ed., vol. 1. L. Salvii, Holmiae. 
1771. Mantissa plantarum altera. L. Salvii, Holmiae. 

Mayr, E. 

1963. Animal species and evolution. Harvard University Press, Cambridge. 

McCann, C. 

1933. Notes on some Indian macaques. Jour. Bombay Nat. Hist. Soc, 36, 
pp. 796-810. 

Medway, Lord 

1963. Review of "Malayan Museum Pamphlets, No. 9: The apes and monkeys 
of Malaya (including the slow loris), by J. L. Harrison . . ." Jour. Malay 
Nat., 17, pp. 62-64. 

Mell, R. 

1922. Beitrage zur Fauna sinica. I. Die Vertebraten Siidchinas; Feldlisten 
und Feldnoten der Sauger, Vogel, Reptilien, Batrachier. Arch. Natur., 88, 
pp. 1-146. 

Pennant, T. 

1781. History of quadrupeds, vol. 1. B. White, London. 

POCOCK, R. I. 

1928. The langurs, or leaf monkeys, of British India, pt. 2. Jour. Bombay 
Nat. Hist. Soc, 32, pp. 660-677. 

(1935). The monkeys of the genera Pithecus (or Presbytia) and Pygathrix found 
to the east of the Bay of Bengal. Proc Zool. Soc. London, 1934, pp. 895-961. 

1939. The fauna of British India, including Ceylon and Burma. Mammalia, 
Vol.1. Primates and Camivora (in part). Taylor and Frances, Ltd., London. 

Raffles, T. S. 

(1821). Descriptive catalogue of a zoological collection, made on account of the 
Honourable East India Company, in the island of Sumatra and its vicinity. 
Trans. Linn. Soc. London, 13, pp. 239-274. 

Robinson, H. C. 

1916. A collection of mammals and birds from Pulau Panjang or Pulau Mapor, 
Rhio-Lingga Archipelago. Jour. Fed. Malay States Mus., 7, pp. 59-72. 

Robinson, H. C. and C. B. Kloss 

1914. The zoology of Koh Samui and Koh Pennan. Jour. Fed. Malay States 
Mus., 5, pp. 128-155. 

Sanborn, C. C. 

1952. The mammals of the Rush Watkins Zoological Expedition to Siam. 
Jour. Siam Soc, 15, pp. 1-20. 



62 FIELDIANA: ZOOLOGY, VOLUME 59 

SCHULTZ, A. H. 

1933. Growth and development. In Hartman, C. G. and W. L. Straus, Jr., ed. 
The anatomy of the rhesus monkey (Macaca mulatta), pp. 10-27. Williams 
Wilkins Company, Baltimore. 

1937. Fetal growth and development of the rhesus monkey. Contr. Embryol., 
Carnegie Inst. Wash., 26, pp. 71-97. 

SODY, H. J. V. 

1949. Notes on some Primates, Carnivora and the Babirusa from the Indo- 
Malayan and Indo-Australian regions. Treubia, 20, pp. 121-190. 

Spiegel, A. 

1954. Beobachtungen und Untersuchungen an Javamakaken. Zool. Gart. 
(N.F.), 20, pp. 227-270. 

Tem Smitinand 

1968. Vegetation of Khao Yai National Park. Nat. Hist. Bull. Siam Soc, 22, 
pp. 289-305. 

Thomas, O. 

1922. Note on the nomenclature of the northern slow loris. Jour. Bomb. Nat* 
Hist. Soc, 28, p. 433. 

ZiMMERMANN, E. A. W. 

1780. Geographische Geschichte des Menschen und der vierfiissigen Thiere, 
vol. 2. Weygandschen Buchhandlung, Leipzig. 



Publication 1123