BIOLOGY The person charging this material is re- sponsible for its return to the library from which It was withdrawn on or before the Latest Date stamped below.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 0 1 1 Macaca fascicularis 1-2 5 4-6 M. mulatta 0 2 1 M. nemestrina 1 0 8 M. assamensis 0 2 8 M. arctoides 0 0 2 Presbytis cristatus 0 0 5 P. phayrei 0 0 9 Hylobates lar 0 0 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 (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. 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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 □ 0 OQ 0 ■■ 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 0 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). 0 — 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 0 . 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 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) =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. 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