ILli,.^.- LIBRARY AT UR3ANA-CHAMPAIGM - ft JjlL,l AJ.-UW jQ^A.^ W FIELDIANA Zoology Published by Field Museum of Natural History Volume 67 TAXONOMY AND EVOLUTION OF LIONTAIL AND PIGTAIL MACAQUES (PRIMATES: CERCOPITHECIDAE) JACK FOODEN WML HISTORY S8WH j AN -2 9 '^76 LIBRARY November 25, 1975 FIELDIANA: ZOOLOGY A Continuation of the ZOOLOGICAL SERIES of FIELD MUSEUM OF NATURAL HISTORY VOLUME 67 '& FIELD MUSEUM OF NATURAL HISTORY CHICAGO, U.S.A. TAXONOMY AND EVOLUTION OF LIONTAIL AND PIGTAIL MACAQUES (PRIMATES: CERCOPITHECIDAE) FIELDIANA Zoology Published by Field Museum of Natural History Volume 67 TAXONOMY AND EVOLUTION OF LIONTAIL AND PIGTAIL MACAQUES (PRIMATES: CERCOPITHECIDAE) JACK FOODEN Research Associate Field Museum of Natural History and Professor of Zoology Chicago State University November 25, 1975 Publication 1216 Patricia M. Williams Managing Editor. Scientific Publications Library of Congress Catalog Card Number: 753914 US ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Dedicated to the memory of Kitti Thonglongya (1930-1974), zoologist, friend, colleague and field companion during 1973 collecting trip to peninsular Thailand. CONTENTS PAGE Introduction l Generic names applied to liontail and pigtail macaques 4 key to external characters 6 Cranial comparisons 7 Tail-proportions, pelage, vertebrae, carriage 18 Female genitalia 28 Glans penis and baculum 33 Blood proteins 45 Comparative ecology and behavior 51 Hybridization 54 Fossil record 60 Evolution and dispersal 66 Species accounts 73 Macaca silenus (Linnaeus, 1758) Synonymy 73 Type 75 Type-locality 77 Distribution 78 External characters 78 Cranial characters 78 Habits and habitats 79 Specimens examined 84 Macaca nemestrina (Linnaeus, 1766) Taxonomy 85 Macaca nemestrina nemestrina (Linnaeus, 1766) Synonymy 91 Type 93 Type-locality 95 Distribution 95 External characters 95 Cranial characters 100 Habits and habitats 100 Remarks 104 Specimens examined 105 Macaca nemestrina leonina (Blyth, 1863) Synonymy 107 Type 109 Type-locality 110 Distribution 110 External characters 112 I II Cranial characters 115 Habits and habitats 115 Remarks 117 Specimens examined 118 Macaca nemestrina pagensis Miller, 1903 Synonymy 118 Type 119 Type-locality 119 Distribution 119 External characters 119 Cranial characters 120 Habits and habitats 121 Specimens examined 121 Gazetteer of collecting localities 122 Macaca silenus 123 Macaca nemestrina nemestrina 125 Macaca nemestrina leonina 137 Macaca nemestrina pagensis 142 References 143 Addenda 168 INTRODUCTION Liontail macaques inhabit southwestern India, and three well- defined forms of pigtail macaques inhabit Southeast Asia from eastern Assam to Borneo (fig. 1). No two of these macaques are sympatric. The geographic ranges of liontail macaques and pigtail macaques are separated by a 2000 km. gap; the ranges of recognized forms of pigtail macaques are adjacent but not overlapping. Current taxonomic practice, following Kloss (1919b, p. 138), is to regard liontails as one species (Macaca silenus Linnaeus, 1758) and various forms of pigtail macaques as subspecies of another species (M. nemestrina Linnaeus, 1766). Although available evidence is not as conclusive as generally assumed (see below), this usage is retained in the present revision. Liontails (M. silenus) and Sundaic pigtails (M. nemestrina nemestrina), both described in the eighteenth century, are the best known members of this group. Indochinese pigtails (M. n. leonina), described in 1863, are less widely known and are frequently confused with M. n. nemestrina. Kepulauan Mentawai pigtails (M. n. pagensis), described in 1903 and represented in collections by a total of nine museum specimens, are known only to a few specialists. The present revision is based on study of 462 museum specimens and review of relevant literature. Specimens examined are preserved in the museums listed below. I am deeply grateful to officals of these institutions for permission to study material in their custody. AIUZ Anthropologisches Institut der Universitat Zurich AMNH American Museum of Natural History, including Archbold Collection (AMNH-AC), New York BM British Museum (Natural History), London BNHS Bombay Natural History Society, Bombay CTNRC Centre for Thai National Reference Collections, Applied Scientific Research Corporation of Thai- land, Bangkok FIELDIANA: ZOOLOGY. VOLUME 67 FMNH Field Museum of Natural History, Chicago IRSN Institut Royal des Sciences Naturelles de Belgique, Brussels Museum of Comparative Zoology, Harvard Univer- sity, Cambridge Museum National d'Histoire Naturelle, Paris Museum Zoologicum Bogoriense, Bogor Naturhistorisches Museum, Basel NHMBe Naturhistorisches Museum, Bern NHRM Naturhistoriska Riksmuseet, Stockholm Natur-Museum Senckenberg, Frankfurt Rijksmuseum van Natuurlijke Historie, Leiden Sarawak Museum, Kuching Staatliches Museum fur Tierkunde, Dresden United States National Museum of Natural History, Washington, D.C. Zoologisches Museum der Humboldt-Universitat, Ber- lin Zoologisches Museum der Universitat, Zurich Zoological Reference Collection, University of Sing- apore (This collection was formerly housed in the Singapore National Museum, also known as Raffles Museum.) Zoologisches Sammlung des Bayerischen Staates, Munich Zoological Survey of India, National Zoological Collection, Calcutta Early stages in this research were supported by U.S. Public Health Service Grant No. GM-13113. Final stages were partially supported by Minorities Schools Biomedical Support Grant No. RR-08043 (National Institutes of Health) to Chicago State University. For skillful assistance I thank the following persons: Mr. Kevin Royt, illustrations; Mr. John Bayalis and Mr. Fred Huysmans, Division of Photography, Field Museum of Natural History, photographs; Mrs. Nancy Shortall, typing. NOTE ADDED IN PROOF: Volume 7 of Primates: Comparative Anatomy and Taxonomy by W. C. Osman Hill (John Wiley and Sons, 1974), which includes a discussion of liontail and pigtail macaques (pp. 652-687), was published after the present work had been submitted for publication; no evidence presented by Hill requires modification of the present work. MCZ MNHN MZB NHMB NMS RMNH SMK SMTD USNM ZMB ZMUZ ZRCS ZSBS ZSI 3 •c I 0 g E IS 15 :§3 3 •c B U a a S'E I 8 8" GENERIC NAMES APPLIED TO LIONTAIL AND PIGTAIL MACAQUES Macaca Lac6pede, 1799, p. 4 — first applied to pigtail macaques by Daudin, [1802], p. 148; synonyms listed below are those applicable to liontail and pigtail macaques only. Silenus Goldfuss, 1820, pp. xxiv, 479 — subgenus of Cynocephalus; included species, C. silenus ( = Macaca silenus), type-species by monotypy and tautonymy. Lesson, [1830], p. 101 — subgenus of Macacus; included species, Macacus silenus, type-species by original designation. Lesson, 1839, p. 70 — genus. J. A. Allen, 1916, p. 51 — genus; selected to replace Macaca. Fjedler, 1956, p. 177 — subgenus; external characters. Les Maimons: I. Geoffroy, 1826, p. 588 — one of three subdivisions ("petite groups") recognized within Macacus; included species, M. silenus, M. erythraeus ( = Macaca mulatto), M. nemestrinus ("Le Maimon"), M. libidinosus ( = Macaca n. nemestrina), M. sped- osus (=M. fuscata), M. maurus. Maimon Wagner, [1839], p. 141 — subgenus of Inuus (=Macaca); included species, /. silenus, I. erythraeus (=Af. mulatto), I. nemestrinus (type-species by tautonymy; Maimon of Buffon cited in synonymy, p. 144), /. arctoides, I. speciosus (=Af. fuscata), I. niger. J. A. Allen, 1916, p. 50 — invalid designation of /. erythraeus as type-species. Vetulus Reichenbach, 1862, p. 125 — genus; included species, V. silenus (type-species by tautonymy, synonym), V. nestor, V. ursinus, V. priamus, V. thersites. Nemestrinus Reichenbach, 1862, p. 139 (not Latreille, 1802), subgenus of Macacus; included species, M. nemestrinus (type- species by tautonymy), M. libidinosus (= Macaca n. nemestrina); preoccupied by Nemestrinus Latreille, 1802, p. 437, a genus of Diptera. (UJUJ) L|JBU9| | IDJ^ KEY TO EXTERNAL CHARACTERS Conspicuous external characters of adult male liontail and pigtail macaques (fig. 1) are contrasted in the following key. Details of individual, sexual and ontogenetic variation are discussed in species and subspecies accounts. Although length of head and body averages about 10-20 per cent greater in adult M. n. nemestrina than in other members of this group, this character is not absolutely diagnostic because of species overlap (figs. 2, 22). 1. Tail length greater than 50 per cent of combined length of head and trunk. Trunk and limbs blackish; crown patch blackish, narrow anteriorly; cheek ruff and beard prominent, pale grayish-brown; muzzle relatively short Macaca silenus Tail length less than 50 per cent of combined length of head and trunk. Macaca nemestrina, 2 2. Tail thinly furred, skin of tail clearly visible through sparse downy pelage. Trunk chocolate-brown non-agouti dorsally, becoming pale brown ventrally; hind limbs brown, forelimbs reddish-brown; crown patch brown, broad anteriorly; cheek hairs short, lateral surface of throat region sharply defined pale ochraceous-buff; muzzle relatively short M. n. pagensis Tail densely furred, skin of tail completely covered by hair 3 3. Mid-dorsal region of trunk golden-brown agouti, similar in color to lateral surface of trunk and limbs. Crown patch dark brown, narrow anteriorly; cheek ruff moderately long, pale ochraceous-buff; reddish streaks extending laterally from external angles of eyes; muzzle relatively short; tail carried arched forward over back, tip of tail directed upward and forward M. n. leonina Mid-dorsal region of trunk blackish, contrasting with golden-brown agouti color of lateral surface of trunk and limbs. Crown patch blackish, broad anteriorly; cheek hairs short, pale basally, blackish at tips; muzzle long; tail carried arched rearward, tip of tail directed downward M. n. nemestrina CRANIAL COMPARISONS (Figures 3-6) Cranial characters that appear to be of greatest taxonomic importance in lion tail and pigtail macaques are (1) greatest length of skull (excluding incisors), (2) relative zygomatic breadth, (3) relative length of rostrum, (4) length of incisor-canine field and premolar-molar field, (5) morphology of supraorbital ridges, and (6) morphology of malar region. These characters are discussed in the following paragraphs and their variation is graphed in Figures 7-9, 22. The method of measurement, except for dental -field measure- ments defined below, is described and illustrated elsewhere (Fooden, 1969a, p. 40). As usual in catarrhines, taxonomic differences in liontails and pigtails are more distinct in skulls of adult males than in skulls of females and immatures. Since only a few adult specimens of M. n. pagensis and wild-collected M. silenus are available, no attempt has been made to construct a formal key to cranial characters. Two male and three female skulls of M. silenus have previously been compared with an unspecified number of skulls of M. n. nemestrina (Kurup, 1963, p. 249). Greatest length of skull averages about 15 per cent larger in M. n. nemestrina than in M. n. leonina, M. n. pagensis, and M. silenus, which are approximately equal in this measurement (figs. 7, 22). Overlap between M. n. nemestrina and the three smaller subspecies or species is minimal. In specimens examined of M. n. leonina, M. n. pagensis, and M. silenus, the range of greatest skull length is 102.3- 121.1 mm. in adult females and 124.1-140.1 mm. in adult males; in M. n. nemestrina these ranges are 115.4-138.9 mm. in adult females (only slightly smaller than in adult male M. n. leonina, M. n. pagensis, and M. silenus) and 139.5-174.5 mm. in adult males. Predictably, variation of skull length generally parallels variation of length of head and body in these subspecies and species (figs. 2, 7, 22). Fig. 3. Anterior and lateral views of adult skulls: M. silenus — a, female, RMNH 1314, zoo specimen; b, c, male, FMNH 47322, zoo specimen; M. n. leonina — d, female, FMNH 99677, Loc. No. 37; e, f, male, FMNH 99691, Loc. No. 21. Approx. Vt nat. size. (Photos FMNH) For locality numbers see Figures 20, 25, and 28. 8 Fig. 4. Anterior and lateral views of adult skulls: M. n. nemestrina — a, female, USNM 114502, Loc. No. 30; b, c, male, USNM 154367, Loc. No. 69; M. n. pagensis - d, female, AMNH-AC 103398, Loc. No. 93; e, f, male, AMNH-AC 103394, Loc. No. 92. Approx. Vi nat. size. (Photos FMNH) 9 •' : ■ --;;: ".'•"'.;*. j *^wZl^^J^^^QPhS^^^^'^/^-^v. ^VjTV H 1" 8 1 ^r x p^HBEeB^ v^& ' II 1 1 '■'-" * 1 B^a^lM bW^mP^bI ov^'-^&vtS. ^^B ^^^^^^ ,^H Fig. 5. Dorsal and ventral views of adult male skulls: a, b, M. silenus, FMNH 47322, zoo specimen; c, d, M. n. leonina, FMNH 99691, Loc. No. 21. Approx. Vi nat. size. (Photos FMNH) 10 Fig. 6. Dorsal and ventral views of adult male skulls: a, b, M. n. nemestrina, USNM 154367, Loc. No. 69; c, d, M. n. pagensis, AMNH-AC 103394, Loc. No. 92. Approx. Vi nat. size. (Photos FMNH) 11 12 FIELDIANA: ZOOLOGY, VOLUME 67 Relative zygomatic breadth (zygomatic breadth /greatest length excluding incisors) in M. silenus and M. n. leonina tends to be somewhat greater than in M. n. nemestrina and M. n. pagensis (fig. 7, table 1). Within each subspecies or species this ratio is about the TABLE 1. Relative zygomatic breadth (zygomatic breadth/greatest skull length x 100) in liontaii and pigtail macaques. Mean — standard error; sample size indicated by figure in parenthesis Species or Infants and subspecies juveniles Adult females Adult males M. silenus 66.43 ± 0.65 (6) 68.40± 0.88 (7) 68.84 ± 0.68 (6) M. n. leonina 64.88 ± 0.43 (23) 67. 39± 0.60 (19) 70.15 ± 0.58 (17) M. n. nemestrina 63.26 ± 0.36 (50) 63.82± 0.39 (35) 64.49± 0.32 (76) M. n. pagensis 62.89 ± 1.07 (2) 64.24 ± 0.68 (3) 63.11(1) same in both sexes. In adult specimens examined relative zygomatic breadth varies from 0.58 to 0.71 in M. n. nemestrina and M. n. pagensis and from 0.63 to 0.75 in M. silenus and M. n. leonina. In infant and juvenile specimens of all four subspecies and species, zygomatic breadth is relatively narrow, approximately as in adult M. n. nemestrina and M. n. pagensis. Zygomatic proportions in 12 M. n. nemestrina specimens (without locality data) measured by Khajuria (1954, p. 118) fall within the range of variation in specimens examined of that subspecies (fig. 7). In liontaii and pigtail macaques, as in many other catarrhines (Vogel, 1966, p. 21) relative length of rostrum (rostral length/ postrostral length) is strongly correlated with skull size (figs. 7, 8). In adult female skulls examined of M. silenus, M. n. leonina, and M. n. pagensis, which are the smallest adult skulls measured, relative rostral length is 0.40-0.55; in adult female skulls of M. n. nemestrina, the next larger size class, relative rostral length is 0.47-0.61; in adult male skulls of M. silenus, M. n. leonina, and M. n. pagensis, which average slightly larger than those of female M. n. nemestrina, relative rostral length is 0.55-0.61; in adult male skulls of M. n. nemestrina, the largest size class in this group of subspecies and species, relative rostral length is 0.62-0.78. Although relative rostral length in liontails and pigtails varies ontogenetically, sexually, and taxonomically, the composite log-log plot of all measurements available for immature and adult males and females is approximately linear (fig. 8). 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Ca3-4. In Jf. a. letwaaa intervertebral joints Ca4~; 7 also are hyperextended to form the anteriorly «i in this subspecies «fig_ 101. In Jf . a. aenresfn intervertebral joints Ca4-a. Ca5-& and Ca€-' ventriflexed to form the proximal put of the d tail flexure, and intervertebral joints Ca7-£ an flexed to complete the tail arch kb%. 101. T difference of tail carriage in these tmo sul consequently, to depend on habitual dorsal a inteivertebral joints Ca4~5. Ca5-€L and CaS-7 in ::-::is:ei =::r. r.£r.:_£. .f:.:r.:.ri.::. ;: :r.f^ . i:-T. i-Tl _•£.> Although mtervertebral joints Ca*-4. €*>-£. Ca6-7 are opposne- ly inflected in Jf . a. Jeoasna and Jf. a. 26 FIELDIANA: ZOOLOGY, VOLUME 67 examined there seems to be no consistent difference between these subspecies in the structure of vertebrae Ca4-Ca7, which are the vertebrae directly affected. However, in vertebra Ca3 the transverse processes in M. n. leonina are consistently broader, relatively and TABLE 4. Breadth (mm.) of transverse processes of first seven caudal ver- tebrae in subadult and adult specimens of M. silenus, M. n. leonina, and M. n. nemestrina; figure in parenthesis is ratio of breadth of transverse processes to length of vertebral centrum (see Table 3). FMNH No.; Vertebra No. sex 1 2 3 4 M. silenus 5 6 7 47322 cJ — ca. 21.2 20.0 16.3 13.8 10.3 8.7 47387 ? 17.4 (1.67) 18.4 (1.57) 15.7 (1.23) 12.2 (0.79) 8.7 (0.41) 8.0 (0.29) 6.8 (1.53) (1.67) (1.32) (0.75) (0.41) (0.34) (0.28) M n. leonina 99673 S ca. 24.0 ca. 24.0 23.0 16.2 13.8 8.1 7.1 99691 i (2.12) 23.5 (2.31) 25.4 (2.19) 21.7 (1.54) 15.0 (1.08) 9.3 (0.55) 6.9 (0.39) 5.5 99688 $ (2.18) 19.4 (2.47) 18.2 (2.05) 15.7 (1.28) 10.8 (0.62) 7.6 (0.35) 5.8 (0.27) 5.1 (2.09) (2.04) (1.85) (1.17) (0.71) (0.55) (0.35) Af. n. nemestrina 542621 6 19.8 19.0 13.2 11.8 10.0 6.8 5.7 543051 ? (1.80) 17.4 (2.00) 15.8 (1.65) 11.8 (1.19) 11.1 (0.74) 7.3 (0.39) 5.9 (0.27) 5.0 (2.38) (2.25) (1.44) (1.08) (0.57) (0.38) (0.32) 1 Subadult (see Table 3). usually absolutely (table 4), than in M. n. nemestrina (and M. silenus). Reduction of these transverse processes in specimens of M. n. nemestrina has been noted previously by Wilson (1970, p. 53, fig. 3). These transverse processes receive the tendons of insertion of m. ischiocaudalis, which acts to abduct and depress the base of the tail (Wilson, 1970, p. 78; Trevor-Jones, 1970, pp. 320, 323). Enlargement of these transverse processes in M. n. leonina may be a function of increased abductor and ventriflexor tension. This, in turn, may stabilize the base of the tail against increased extensor tension (of m. extensor caudae medialis) that apparently is exerted more distally to effect the characteristic dorsal hyperextension of joints Ca4-5, Ca5-6, and Caf>7. FOODEN: LIONTAIL & PIGTAIL MACAQUES 27 A more complete explanation of the structural basis for the difference of tail carriage in M. n. leonina and M. n. nemestrina probably will require further comparative study employing more adequate skeletal and cadaver material of M. n. nemestrina. The evolutionary significance of the tail carriage difference between these subspecies, like that of many conspicuous and taxonomically important differences between primates subspecies and species, remains obscure. FEMALE GENITALIA Cyclical estrous swelling of the circumanal sexual skin is well known in M. n. nemestrina (F. Cuvier, 1816-1817, pi. 5, fig. 2; Pocock, 1926, fig. 68b; Zuckerman, 1937, p. 319; Kuehn et al., 1965, figs. 1, 2; White et al., 1973, p. 189). This swelling typically develops gradually, beginning during or immediately after the menstrual period and reaching maximal tumescence usually in about 15 days. At its maximum, the engorged pinkish hairless area extends dorsoventrally from the base of the tail to the ventral border of the ischial callosities, which are partly buried in the swelling, and extends laterally over an area twice the breadth of both callosities (fig. 11). Ventral to the callosities the thinly haired skin anterior to the vulva (mons pubis) is less conspicuously swollen. Following ovulation, usually within one or two days after maximal tumes- cence, the swelling rapidly subsides. Detumescence persists for about 15 days, until the next menstrual period. In laboratory mating tests, maximal female perineal tumescence in M. n. nemestrina is directly correlated with maximal sexual behavior in females (Goldfoot, 1971, p. 330) and with maximal incidence and rate of ejaculation in males (Bullock et al., 1972, p. 230). Sexual skin in M. silenus has been cursorily described in an unspecified number of specimens by Pocock (1926, p. 1547) and more precisely characterized in one specimen by W. C. O. Hill (1937, p. 210, figs. 2a, 2b). Judging from these accounts, the general pattern of sexual skin morphology is similar in M. silenus and M. n. nemestrina, although swelling is much less distended in estrous M. silenus (fig. 11). In this species the swelling does not form a continuous pillowlike mass, as in M. n. nemestrina, but instead is subdivided into five separate swollen areas— one large subcaudal swelling, a pair of smaller swellings dorsolateral to the anus, and a pair of broad swellings that surround the ischial callosities, lateral to the vulva. (The ventral margin of the vulva in one M. silenus adult female that I observed at Lincoln Park Zoo, Chicago, is approximately in line with the lower border of the ischial callosities, 28 FOODEN: LIONTAIL & PIGTAIL MACAQUES 29 M. silenus M. n. nemestrina Fig. 11. Female sexual swellings in M. silenus (modified from W. C. O. Hill, 1937, p. 214) and M. n. nemestrina (Kuehn et aL, 1965, p. 253): 1, subcaudal swelling; 2, para-anal swellings; 3, para-vulval swellings. as in other species of macaques; the vulva is not especially high between the callosities, as described and figured by W. C. O. Hill, loc. cit.) The paired swellings dorsolateral to the anus in M. silenus are particularly less developed than in M. n. nemestrina, where these para-anal swellings fuse with the subcaudal swellings dorsally, extend over the gluteal region dorsolateral^, and overlap the ischial callosities ventrally. Morphology of sexual swelling has not been reported in estrous specimens of M. n. leonina and M. n. pagensis. In non-estrous M. n. leonina specimens examined (FMNH 99674, 99687, subadults; 99685, 99688, pregnant adults; 99677, 99678, 99679, lactating adults), the hairless area of sexual skin seems to be less extensive than in corresponding non-estrous specimens of M. n. nemestrina, more nearly resembling that in M. silenus. Internal female reproductive structures have been previously investigated in three specimens of M. n. nemestrina (Hafez and Jaszczak, 1972, pp. 297, 300). The present account is based on study of 12 alcohol-preserved reproductive tracts of M. n. leonina immatures and adults (table 5). The female tract in M. n. leonina (fig. 12) is essentially similar to that in M. n. nemestrina (Hafez and Jaszczak, op. cit.). 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I- cs Q IE U o CM zi 01 o> £ o .£ .£ 0 0 a *j ™ « ._> 0) o> (0 o 0) 0 '8 0) 2* vi 0) be be CO CQ T3 -O a> C 0 > >> X, o> o> e o s » 0> C 2 c CQ cs £ £ 0) 01 — e/5 — 0) «j *^3 '■W 4J 08 cs O .c CA (O 0* PQUh — -7 fa fa i-J N CO *tf lO o t> CN q CO CN r-J © i-H r-^ °°. °°. m q CO CN CN CN T-i t^ CO 00 I-H q I-H I-H I-H co q q © q CO q t-H ■* CN CO CN CO i-i CN CN CO CO CN CN X X X X X X X X X X X ■ 00 Ol 00 00 00 I> rt 00 00 t~ c~ t~ CO CO CO CO CO CO 5 CO CO CO CO CO OS Oi O Oi CT> g a> CJi CJ5 C7> CJ5 O OJ <3> 05 o> Z C31 o> CJ> CTl (Ji 31 32 FIELDIANA: ZOOLOGY, VOLUME 67 Fig. 12. Sagittal section of female reproductive tract in subadult M. ru leonina, FMNH 99674; x 2. UB: uterine body, UC: uterine cervix, V: vagina (distortion of wall is artifact of preservation). (Photo FMNH) fascicularis (Jainudeen et al., 1972, p. 473) and M. mulatto (Wislocki, 1933, p. 239; Eckstein and Zuckerman, 1956, p. 138; Cuadros, 1971, p. 139). However, the uterine cervix in M. n. leonina evidently is somewhat smaller than at corresponding stages in M. mulatto. Judging from specimens examined (table 5), the cervix in M. n. leonina enlarges markedly approximately at puberty (FMNH 99687), regresses during adolescence (FMNH 99674; NM4), reaches maximal size during pregnancy (FMNH 99685, 99688), and regresses during lactation (FMNH 99677, 99679). Structure of the female tract in M. n. leonina is distinctly different from that in the M. sinica-group (M. sinica, M. radiata, M. assamensis, M. thibetana), which is characterized by presence of cornified vaginal spines and a hypertrophied endocervical chamber (Fooden, 1971b, p. 67). It also differs from that in M. arctoides, in which a large dorsal vestibular collide is present and cervical colliculi are absent (Fooden, 1967, p. 939; Demers et al., 1972, p. 529; Hafez and Jaszczak, 1972, p. 302). NOTE ADDED IN PROOF: Cf. Eaton, G. G. and J. A. Resko, 1974, Ovarian hormones and sexual behavior in Macaca nemestrina, J. Comp. Physiol. Psychol., 86, pp. 919-925. GLANS PENIS AND BACULUM This account is based on study of 30 specimens of penises or bacula of liontail and pigtail macaques (M. silenus, 3; M. n. leonina, 19; M. n. nemestrina, 8; tables 6, 9) and review of relevant literature (Daubenton, in Buffon and Daubenton, 1766, p. 179; F. Cuvier, 1820, p. 1; F. Cuvier, 1822a, p. 2; DeBeaux, 1917, p. 10; 1923, p. 34; Pocock, 1921, p. 227; 1926, p. 1559; Pohl, 1928, p. 103; W. C. O. Hill, 1937, p. 213; 1958, p. 655). In liontail and pigtail macaques the glans penis is helmet shaped and bluntly bilobed (fig. 13), closely resembling that in Celebesian macaques (Fooden, 1969a, p. 20) and M. sylvanus (Daubenton, in Buffon and Daubenton, 1766, p. 119; W. C. O. Hill, 1958, p. 655, fig. 18a) and generally similar to that in the mulatta- group (M. fascicularis, M. mulatto,, M. cyclopis, M. fuscata; Fooden, 1972, p. 310). The bluntly rounded glans in macaque groups listed above is distinctly different from the apically elongate subpyramid- al glans in the smica-group (M. sinica, M. radiata, M. assamensis, M. thibetana; Fooden, 1971b, p. 72) and the even more elongate lanceolate glans in M. arctoides (Fooden, 1967, p. 939). These three sharply defined glans types in the genus Macaca (fig. 14) were first recognized by Pocock (1921, p. 229); subsequent authors, including Pocock (1926, p. 1557), have tended to obscure the basic penial trichotomy of macaques. The bluntly rounded glans type seems to be the most common form in macaques and other cercopithecids (Pocock, 1926, p. 1557) and, therefore, probably is primitive in the genus and family. The three subdivisions of the genus Macaca that are defined by structure of the glans penis exactly correspond, as far as presently known, to three subdivisions defined by structure of the uterine cervix. In M. n. leonina, M. fascicularis, and M. mulatto, all with glans bluntly rounded, the cervix and cervical colliculi are moderately large (see above); in M. sinica, M. radiata, and M. assamensis, with glans apically elongate, the cervix is enormously inflated and cervical colliculi are hypertrophied (Fooden, 1971b, p. Q a* inn h in ■ (m — £ o 1 a B Ifl (O t«; CN CN CN c 3 5 1 c 1 i o s c ** ". "5 09 •o i-H (N ■ . com !-( rH ,-H i-H i-h i-H I rl «-H O Z I CO o *o m 2 in X m 2 §2 CN W CN © © © ^ © tt in CN CI KlO © © © 2 S fcZZZZ^DD 34 2 ° c Q a* 00 00 O Tf N t» 00 00 H 0> ^ N H N tfJoJ-^iriui^ioioo S 3 3 o C8 22 m oj i-i I u IS o «4-l o 1 > c D a» e I c <: I I I in iflm , CO O) Tf o 4) fc 1 * 0 3 = r- <-> 0/ •= 3 !2 _^ /• _ w c - 0 Oh CO ©CN CI 2oo CO sill 1/3 ITS X . CO £ a 12 i2v .§ "S i»' 5 g * o « - o in m ^ CQ CO E CO 6 Ml 2 (12, M2) 3 (C) 4.5 M3 >6 — 3 (C) 5 — >5 — >6 — >7 Glans length4 (mm.) Relative breadth M. fascicularis 6.2, 7.7, 8.5 0.44, 0.45, 0.52 7.4, 7.7 0.41, 0.42 7.0 0.43 7.4, 12.3 0.47, 0.53 12.9 0.49 14.4, 15.2, 16.6 0.55, 0.46, 0.41 M. mulatto 11.8 (10.6-13. 1)5 0.42(0.36-0.50)5 13.5 0.36 9.5 0.47 18.7 0.54 M. fusct ita6 17.0, 17.1 0.38, 0.40 18.3 0.40 15.9 0.39 20.0 0.40 20.0 0.48 1- Specimens in FMNH. 2- Parentheses indicate incompleted erupted teeth. 3- Based on dental emergence norms for M . mulatto (Hurme, 1960, p. 797). 4- Measured parallel to long axis of glans; differs slightly from greatest length reported previously (Fooden, 1972, p. 306). 5- Mean and extremes in eight specimens. 6- For age estimates see Fooden, 1972, p. 306. 38 FIELDIANA: ZOOLOGY, VOLUME 67 and pigtail macaques, judging from measurements of dry specimens reconstituted in fluid (table 8). TABLE 8. Relative breadth of glans penis in reconstituted dry specimens of Celebesian macaques. Glans Relative Species Specimen No. length (mm.) breadth M brunnescens NMS 1017, 1020 13.8,15.2 0.59,0.63 M hecki RMNH 1099, 3801 14.9, 17.3 0.66, 0.51 Af maura RMNH 751 (2363, 2484) 8.4, 13.3 0.83, 0.66 M. nigra RMNH 13, MCZ 34459 7.9,9.8 0.87,0.72 Af ochreata NMS 1022 13.2 0.61 Af tonkeana NMS 16838 13.2 0.64 In liontail and pigtail macaques (fig. 13), as in M. mulatto, the dorsal part of the glans extends back over the shaft about twice as far as the ventral part, the glans is bilobed, the urethral orifice opens between the lobes, and the baculum protrudes into the left lobe, which is somewhat larger than the right. Also as in Af. mulatto, the glans and distal part of the shaft are densely studded with tiny epidermal papillae about 0.05 mm. in diameter; these are much smaller than the epidermal spines (0.3 mm. diameter) present on corresponding penial surfaces in the sinica -group (fig. 14). In early accounts (Daubenton, in Buffon and Daubenton, 1766, p. 179; F. Cuvier, 1822a, p. 2) the glans in M. n. nemestrina is characterized, somewhat confusingly, as three-lobed; this evidently is a result of counting as a lobe a small variably defined V-shaped dorsomedian eminence that is situated between the bases of the right and left lobes, proximal to the urethral cleft (fig. 13). In living specimens of M. silenus, color of the detumescent glans is dark gray ( W. C. O. Hill, 1937, p. 214; personal observation, one adult, Lincoln Park Zoo, Chicago) and color of the tumescent glans is "bright purple" (W. C. O. Hill, loc. cit). In M. n. leonina, the glans is pinkish (DeBeaux, 1923, p. 35; personal observation, one adult, National Zoological Park, Washington, D.C.). In M. n. nemestrina the color of the "penis" (?glans) is recorded as "pale crimson" (field tag of BM 55.1503, Sungai Renggam, West Malaysia). The external similarity in structure of the glans in liontail and pigtail macaques conceals unexpected internal differences in size and structure of the baculum (fig. 15; table 6). Comparing species for which reasonably adequate samples are available, length of the Fig. 14. Basic glans types in genus Macaco, left lateral and dorsal views: a, b, bluntly rounded type, M. fascicularis, FMNH 99642; c, d, subpyramidal type, M. h, 99e91,> 6 yr.; i, 99673, > > 6 yr. Natural size. (Courtesy Chicago Zoological Park, Brookfield, 111.) 39 40 FIELDIANA: ZOOLOGY, VOLUME 67 baculum in M. n. leonina (21.0-26.7 mm. in 10 adults) averages about 25 per cent greater than in M. n. nemestrina (16.1-21.1 mm. in seven adults; cf. Kinzey, 1971, p. 102). This is somewhat surprising, considering that head and body length in M. n. leonina averages about 15 per cent less than in M. n. nemestrina (fig. 2). The bacular length difference in M. n. leonina and M. n. nemestrina is due principally to a difference in length of the downward inflected distal process (fig. 15). In M. n. leonina this process is sharply defined, and its length is 29-51 per cent of that of the proximal bacular shaft (10 specimens); in M. n. nemestrina the inflected distal process is more weakly defined, and its length is 20- 30 per cent of that of the proximal shaft (seven specimens). The dorsoventral diameter of the bacular shaft also is notably greater in M. n. leonina than in M. n. nemestrina; as a result, the shaft is bilaterally flattened in M. n. leonina and more nearly cylindrical in M. n. nemestrina. One baculum of M. n. leonina and one of M. n. nemestrina are anomalous. In the anomalous M. n. leonina specimen (NM6; fig. 15k) the shaft is unusually slender, the basal end is studded with small tuberosities, and the distal process, which is weakly inflected, terminates in three vertically arrayed lobules, in place of a simple ventrally directed tip. In the anomalous M. n. nemestrina baculum (AMNH 119514, zoo animal) the bone is unusually thickened and porous (fig. 15s). As indicated previously (Fooden, 1969a, p. 25), the abnormal structure of this baculum may be a result of captivity- induced bone pathology. However, bacular structure in two other captive specimens examined (NMS 2629; RMNH 2000; figs. 15p, 15q) is essentially similar to that in wild-collected specimens. Although examination of three available specimens of M. silenus (fig. 15a-c, table 6) suggests that bacular structure is Opposite: FlG. 15. Bacula in adult liontail and pigtail macaques; left lateral view, dorsal surface of baculum at top, distal end at left; italicized figures below indicate locality numbers as in distribution maps, Figures 25, 28. a, b, c, M. silenus (zoo specimens): BM 41.1.15.1 (subadult), NMS 2630 (subadult or adult), RMNH 1308 (subadult or adult); d-k, M. n. leonina: BM 55.1505 47.NM 5 ("Thailand"), NM 7 ('Thailand"), USNM 241022 25, NM2 ("Thailand"), PMl ('Thailand"), USNM 258868 36, NM6 ("Thailand"); !-•, M. n. nemestrina: BM 55.1503 12, MCZ 35670 89, FMNH 68703 90 (subadult), NMS 2629 (zoo), RMNH 2000 (subadult or adult, zoo), BM 55.707 58, FMNH 105676 4 (subadult), AMNH 119514 (captive, deformed), x 2. (Photos FMNH) 41 a 0 o a 9 es s ! 3 - £ £ w) O co i »■« o» oe i »ft © «r ^ cn Q S. * 3= CO _ J= t— i •— t— i t—c i— i CM 2 A -2 - x ff-mtttO'-rtioo a - OH - a Si F^ rH i- o r- (0 1- as r» zt 1 — O X pH m o> as ~] - V 1 — — iH fH 1-1 i-t ?\ n o> > — ' — X X pm ^r O o X re N i •— o O X o o o t- - o o O • JS OhOtONfflMNO) » t« C« X N ^' 1(5 t« X £ M -c 1- *■» 01 .<« BO — CO .8 c 8. ■a 4 c ,—. <-^ ^^ > i_ — NN CO CO . 0 H V E~~ SS » * c 0) ft — e o> c -M K3 tt as r~ tN CO 3 o c c-j «-« r-l OS CD CO CO — >,3 r~ to Xt~r~XXr~c35r- lD<0<0(OtO<0(C05C7><35C75<75C7>05 C7>CTlC75CT>C75C75C75CT>C75 42 FOODEN: LIONTAIL & PIGTAIL MACAQUES 43 ~'m k ^ mLm • H^H^^Bi fc jpm**zi Fig. 16. Radiographs of glans penis in immature and adult M. n. leonina showing ontogenetic development of baculum: a, FMNH 99690, estimated age 1 yr.; b, 99680, 2.5 yr.; c, 99676, 3 yr.; d, 99675, 3.5 yr.; e, 99686, 4.5 yr.; f, 99684, 6 yr.; g, 99672, 6 yr.; h, 99691, > 6 yr.; i, 99673, >> 6 yr. Natural size. (Courtesy Chicago Zoological Park, Brookfield, 111.) taxonomically distinctive in this macaque, as in M. n. leonina and M. n. nemestrina, the evidence is imperfect and should be interpreted with caution. All three bacula are from zoo animals, which introduces the possibility of captivity-induced deformation. In addition, one of these specimens is from a known subadult, and the other two are from animals of unknown age. In these three M. silenus bacula, the bases are exceptionally thick, the proximal shafts are stout and cylindrical, and the distal processes are short and weakly defined. Bacular form in these specimens is more like that in M. n. nemestrina than in M. n. leonina; however, overall size exceeds that in M. n. nemestrina, and diameter of the base is disproportionally greater than in M. n. nemestrina. A series of nine M. n. leonina specimens collected in a restricted area in western Thailand in 1967 (Fooden, 1971a, p. 32) provides an opportunity for investigation of ontogenetic devel- 44 FIELDIANA: ZOOLOGY, VOLUME 67 opment of penial structure in this subspecies. Estimated ages of these nine specimens are: 1 year (infant), 1; 3 years (juvenile), 3; 4-5 years (late juvenile), 1; 6 years (subadult), 2; > 6 years (adult), 2 (table 9). Judging from available evidence (table 9, fig. 16), size of glans and baculum remains approximately constant from about age one year to age three years, while length of head and body is increasing by nearly 50 per cent. Beginning about age 4.5 years, when permanent canines emerge and puberty begins, size of glans and baculum increases rapidly, at a rate about five times that of head and body (linear dimensions). This pattern of development suggests that penial growth in M. n. leonina is controlled primarily by sex hormones, rather than by pituitary growth hormones. Similar indications for M. fuscata were reported previously (Fooden, 1972, p. 309). The characteristic prolongation of the downward inflected distal process of the baculum appears relatively late in ontogenetic development of M. n. leonina (fig. 16; table 9). As a result, bacular form in juvenile M. n. leonina (fig. 16), resembles that in adult M. n. nemestrina (fig. 15), which therefore is paedomorphic for this character. BLOOD PROTEINS Data suitable for serological comparison of M. n. leonina and M. n. nemestrina are provided by a recent series of studies of blood protein polymorphism in various macaques (table 10). Although authors of the studies do not explicitly recognize these two subspecies of pigtails, adequate information is given on the geographic origin of specimens to permit subspecific identification. No blood protein data are available for M. n. pagensis or for M. silenus. Blood protein phenotypes are variably differentiated in M. n. leonina and M. n. nemestrina (table 10), although none of these phenotypes are absolutely diagnostic and characteristic in these or other macaque subspecies and species. M. n. leonina and M. n. nemestrina are essentially similar in frequency distributions of phenotypes of haptoglobin, acid phosphatase, albumin, thyroxine- binding prealbumin, alkaline phosphatase, plasma phosphohexose isomerase, 6-phosphogluconate dehydrogenase, and phosphogluco- mutase. These two subspecies differ slightly to moderately in frequency distributions of phenotypes of alpha l-antitrypsin, carbo- nic anhydrase II, erythrocyte phosphohexose isomerase, tranferrin, and carbonic anhydrase I. These subspecies differ strikingly in frequency distributions of phenotypes of plasma agglutinin and hemoglobin. The ratio of anti-A to anti-A + B aglutinin phenotypes is about 3:1 in M . n. leonina, and about 1:9 in M. n. nemestrina', the ratio of FS to F hemoglobin phenotypes (determined elec- trophoretically) is about 3:1 in M. n. leonina, while in M. n. nemestrina F is the only hemoglobin so far detected elec- trophoretically. Since all studies in this series evidently are based on blood samples from many or most of the same monkeys, differences in the frequency distributions presumably indicated that at least some of these blood protein phenotypes are inherited independently; however, the loci for carbonic anhydrase I and carbonic anhydrase II probably are linked in these macaques 45 TABLE 10. Frequency distribution (per cent) of polymorphic blood pro- tein phenotypes in Af. n. leonina and Af. n. nemestrina. Plasma proteins Haptoglobin * Af. n. leonina (85 specimens) Af. n. nemestrina (158 specimens Albumin* Af. n. leonina (85) Af. n. nemestrina (158) Thyroxine-binding prealbumin^ Af. n. leonina (36) Af. n. nemestrina (128) Phospohexose isomerase^ Af. n. leonina (25) Af. n. nemestrina (73) Alkaline phosphatase* Af. n. leonina (85) Af. n. nemestrina (158) Alpha \ -antitrypsin4 Af. n. leonina (64) Af. n. nemestrina (151) Transferrin^ Af. n. leonina (85) Af. n. nemestrina (158) Af. n. leonina (cont.) Af. n. nemestrina (cont.) Agglutinin^ Af. n. leonina (82) Af. n. nemestrina (144) Erythrocyte proteins NADH diaphorase? Af. n. nemestrina (25) Acid phosphatase^ Af. n. leonina (40) Af. n. nemestrina (127) 6-phosphogluconate dehydrogenase^ Af. n. leonina (36) Af. n. nemestrina (110) Phosphoglucomutase^ Af. n. leonina (23) Af. n. nemestrina (111) Carbonic anhydrase 11*0 Af. n. leonina (17) Af. n. nemestrina ( 12) Phenotypes \ and frequencies Hpl-1 100.0 ) 100.0 Single Double 98.8 1.2 98.1 1.8 PA« PAfs 100.0 0 97.2 2.8 1 6-1 Unidentified 100.0 0 0 94.5 1.4 A AC 100.0 0 94.3 5.7 AB B AC BC C CD BD D 0 1.6 0 7.8 68.7 14.1 1.6 6.2 0.7 6.6 1. 3 32.4 58.3 0.7 0 0 BG CC CD CF CG D"D DD DF 1.2 1.2 3.5 4.7 12.9 0 0.6 3.2 4.5 2.5 DG F'GFF' F'G FF 17.7 0 0 0 3.5 23.4 0.6 1.3 0.6 7.6 1.2 8.2 3.5 0 13.9 17.7 FG GG 10.6 31.8 17.7 6.4 Anti-A Anti-B Anti-A+B 74.4 1.2 24.4 9.0 2.1 88.9 Phenotypes and frequencies A AB AC 92.0 4.0 4.0 A 100.0 100.0 AS A AF 2.8 97.2 0 0 97.3 2.7 I V 95.7 4.2 1 92.8 7.5 » b 32 a2b 52.9 17.7 29.4 66.6 16.7 16.7 46 1 5-1 6-1 81.8 18.2 0 97.1 0 2.9 a b ab c d o 7.9 57.1 1.6 0 6.4 27.0 32.1 23.7 9.1 1.2 1.2 32.7 F FS S 24.7 72.9 2.4 100.0 0 0 FOODEN: LIONTAIL & PIGTAIL MACAQUES 47 TABLE 10. (continued) Erythrocyte proteins Phenotypes and frequencies Phosphohexose isomerase^ M. n. leonina (11) M. n. nemestrina (34) Carbonic anhydrase 1*0 M. n. leonina (63) M. n. nemestrina (165) Hemoglobin^1 M. n. leonina (85) M. n. nemestrina (160) 1- Ishimoto, 1972a, pp. 260, 273. 2. Weiss etal., 1971, p. 78. 3- Ishimoto and Kuwata, 1972, p. 161; enzyme activity stronger in red cells than in plasma. 4- Omoto et al., 1970, p. 220. 5- Ishimoto, 1972a, p. 255; cf. Prychodko et al., 1971, p. 180. 6- Nakajima et al., 1970, p. 246. 7- Ishimoto, 1971, p. 385; M. leonina specimens not included in this study. 8- Ishimoto, 1972b, pp. 339, 341. 9- Ishimoto, 1972b, p. 342; cf. Prychodko et al., 1971, p. 176. 10. Tashian et al., 1971, p. 190. !!• Ishimoto et al., 1970, p. 235, phenotypes determined electrophoretically; hemoglobin polymorphism in M. n. nemestrina has subsequently been detected chromatographically (Nute and Pataryas, 1974a, p. 21). (DeSimone, Linde, and Tashian, 1973, p. 55; Magid et al., 1973, p. 157; DeSimone, Magid, and Tashian, 1973, p. 165). Blood protein data published for subspecifically indeterminate pigtails of unknown geographic origin are generally concordant with data summarized above for subspecifically identifiable specimens. In 40 adult pigtails of unknown geographic origin studied by Nute and Stamatoyannopoulos (1971a, p. 145; 1971b, p. 109), hemoglobin phenotype is monomorphic, which indicates that these specimens probably were M. n. nemestrina (see table 10). In 75 pigtails studied by Crawford (1966, p. 398), the frequency distribution of hemoglobin phenotypes is 57.3 per cent F (type 2 of Crawford), 26.7 per cent FS (types 3 and 4), and 16.0 per cent S (type 1), which suggests that this may have been a mixed group of M. n. leonina and M. n. nemestrina, approximately as later suspected by Crawford (1971, p. 706; cf. Nute and Pataryas, 1974a, p. 17). In a general review of ABO blood groups in primates, Moor-Jankowski and Wiener (1971, p. 240) report that B and O are characteristic of pigtails (subspecifically indeterminate), which is compatible with the 48 FIELDIANA: ZOOLOGY, VOLUME 67 observed preponderance of agglutinin anti-A in M. n. leonina and anti-A + B in M. n. nemestrina (table 10). Transferrin poly- morphism reported in subspecifically indeterminate pigtails (Goodman et al., 1965, p. 885; Goodman, 1967, p. 8; Prychodko et al., 1969, p. 104) is likewise compatible with transferrin poly- morphism in identifiable M. n. leonina and M. n. nemestrina (table 10). In a captive multi-male colony of 16 pigtails of unknown geographic orgin, nine blood proteins were studied in an attempt to determine paternity of five infants (Simons and Crawford, 1969, p. 255). Since hemoglobin phenotype was monomorphic, the colony probably consisted exclusively of M. n. nemestrina (see table 10). This colony also was monomorphic for phenotypes of phosphogluco- mutase (PGM-M), acid phosphatase (pm), erythrocyte glucose-6- phosphate dehydrogenase (GdA), erythrocyte 6-phosphogluconate dehydrogenase (6PGDA; cf. table 10) and haptoglobin (Hpl). The same colony was polymorphic for phenotypes of serum pseudocholi- nesterase (E«; E.s, Eiu, hypothesized), transferrin (TfA, TfB, TfD), and blood groups (14 reagents). In a mixed group of M. n. leonina and M. n. nemestrina, MN and Lewis blood groups were studied by Nakajima et al. (1970, p. 248); 39 individuals tested were positive for M or M-like antigens, 236 tested were negative for Lea antigens, and 12 of 123 tested were positive for Leb antigens. In 13 pigtails of unknown geographic origin, erythrocyte isoantigen polymorphism was studied with taxonomically inconclusive results (LaSalle, 1969, p. 120). The amino acid composition of the f$ - hemoglobin chain in M. n. nemestrina has been compared with that in M. fascicularis, M. mulatto, and M. fuscata (Nute and Pataryas, 1974b, p. 79); the composition of this chain in M. n. leonina is unknown. In Indochinese populations of M. n. leonina, M. mulatto, and M. fascicularis, similarities in frequency distributions of transferrin phenotypes and 6-phosphogluconate dehydrogenase phenotypes have been construed as evidence of interspecific gene exchange (Goodman, 1967, p. 9; Prychodko et al., 1969, p. 108; Prychodko et al., 1971, p. 181). M. arctoides (-M. speciosa of authors) formerly was also included in this hypothetical group of interbreeding "semispecies" (Goodman, 1965, p. 255; Goodman et al. 1965, p. 886), but in recent reports this macaque apparently is regarded as a well- isolated species (Prychodko et al., 1969, p. 107; Prychodko et al., 1971, p. 179). However, regardless of the disposition of M. arctoides, available evidence does not support the assumption that similarity FOODEN: LIONTAIL & PIGTAIL MACAQUES 49 of blood protein phenotype frequencies necessarily implies gene exchange. Transferrin phenotypes, for example, are more similar in Thai M. n. leonina and Thai M. mulatto (index of dissimilarity 0.360; Prychodko et al., 1969, p. 107), which are members of two different species, than in Thai M. mulatto and northeast Indian M. mulatta (index of dissimilarity 0.546), which are local populations in one continuously distributed species. Since gene exchange usually is assumed to be greater within species than between species, it follows that similar blood protein phenotypic frequencies are not a reliable indicator of gene exchange. If interspecific gene exchange has occurred in Indochinese macaques with similar transferrin and 6-phosphogluconate dehy- drogenase phenotypes, this probably should be also be evident in non-hematological characters of these specimens. However, no such non-hematological evidence of gene exchange has been observed or reported in these critical specimens. In fact, there is no indication that other blood protein phenotypes in these specimens support the hypothesis of gene exchange, and one published analysis of their phenotypic frequencies of alphai - antitrypsin definitely fails to demonstrate such gene exchange (Omoto et al., 1970, p. 226). As previously indicated by Jolly (1971, p. 197), it seems likely that reported similarities in blood protein phenotype frequency dis- tributions in Indochinese macaque species probably are the result of common inheritance and/or convergent evolution, not gene ex- change (cf. Weiss et al, 1973, p. 225). Frequency distributions of blood protein phenotypes also have provided the basis for a computer-generated dendrogram of hypothetical interrelationships among macaque species and in- fraspecific populations (Goodman, 1971, p. 83; cf. Weiss et al., 1973, pp. 220-223 and Ishimoto, 1973, p. 9). (It should be noted that this dendrogram is based on the implicit assumption that similar blood protein phenotype frequencies are the result of common inher- itance, not of gene exchange.) Judging from the dendrogram, hematological evidence apparently indicates that M. n. leonina and M. n. nemestrina diverged from the common ancestral macaque lineage subsequent to the divergence of Celebesian macaques and that M. mulatta diverged subsequent to the divergence of M. fuscata. However, these phylogenetic inferences are contrary to those derived from non-hematological characters, which indicate that short-tailed Indochinese M. n. leonina and Sundaic M. n. nemestrina probably represent the stock that is ancestral to (hence 50 FIELDIANA: ZOOLOGY, VOLUME 67 diverged prior to) stump -tailed Celebesian macaques (fig. 18c; Fooden, 1969a, p. 61) and that short-tailed continental M. mulatta probably represents the stock ancestral to stump -tailed insular M. fuscata (Fooden, 1972, p. 310). Judging from non-hematological characters, it is likewise difficult to accept the hematologically supported assertion (Goodman, 1971, p. 86) that "human beings are very generalized mammals"; compared with Mesozoic stem mam- mals and with primitive marsupials and insectivores (Romer, 1966, pp. 193-211), humans obviously are highly specialized, except for retention of pentadactyly (cf. Clark, 1972, p. 473). COMPARATIVE ECOLOGY AND BEHAVIOR As discussed in detail in species and subspecies accounts (see below), liontail macaques and pigtail macaque subspecies appear to be essentially similar in habitat, diet, and troop size. The usual or exclusive habitat of liontails and pigtails is dense evergreen rain forest, and their geographic distribution closely coincides with the distribution of this habitat in tropical Asia (fig. 17). The food of these macaques evidently consists chiefly of fruits and seeds, occasionally augmented by young tender leaves, insects (larvae and adults), spiders, and, probably, birds eggs and small vertebrates. Troop size in these macaques typically is about 5 to 20. Most troops probably are composed of 1-3 adult males, 2-10 adult females, and 2-10 immatures. Solitary adult males are fairly common; this presumably is correlated with the unbalanced ratio of male and female adults within troops. Altitudinal distribution is not uniform in liontails and pigtail subspecies (table 11). Of the three relatively well-known species or subspecies, M. silenus, with known altitudinal range 300-1300 m., is restricted to upper elevations; M. n. leonina, with altitudinal range 75-1250 m., is limited to the vicinity of foothills of major mountain chains; and M. n. nemestrina, with altitudinal range 0-1700 m., is relatively widespread and inhabits lowland areas as far as 150 km. distant from the nearest mountains. These differences in altitudinal range probably are determined by regional temperature and rainfall patterns (Koeppe and DeLong, 1958, pi. 2), which, in turn, determine rain forest distribution (fig. 17). Liontails and pigtail subspecies also appear to differ in population density and arboreal-terrestrial preference. Population density in M. silenus seems to be much lower than in M. n. leonina and M. n. nemestrina, judging from frequency of reported encounters. M. silenus and M. n. leonina apparently are much more arboreal than M. n. nemestrina; M. silenus and M. n. leonina are rarely encountered on the forest floor, whereas such sightings of M. n. nemestrina are commonplace. When alarmed, M. silenus and M. 51 -■' "*>, £"8 § « "§'5 8-S8 O a) .9 3! ~ j= jj u C 3 .2 * - — = = SO"2 g i = •p >. ^ « -I 3- — "* 08 J a O 00 ♦< .sa: |8 c «2 U ^ 00 g - <*. 5" o « So 8 9 .a o. «. -o -a ~Z c e « 4) — 0 o St> 52 FOODEN: LIONTAIL & PIGTAIL MACAQUES 53 TABLE 11. Altitudinal distribution of accurately known localities of lion- tail and pigtail macaques. Locality numbers (italicized) as in Figures 20, 25, 28; for details, see Gazetteer of Collecting Localities. Species or subspecies; locality numbers Altitude (m.) M. silenus M. n. leonina M. n. nemestrina M. n. pagensis 1600-1799 — — 82 - 1400-1599 — — — — 1200-1399 — 6 — — 1000-1199 7, 10, 13, 17 — 16, 75 — 800-999 — 8, 32 66 — 600-799 — 30 16, 17, 25 — 400-599 8 — 43, 82 — 200-399 1 21, 22, : 23, 25 50, 56 — 0-199 — 1, 2, 29, 31 , 46 11, 44, 46, 88 92 , 93 n. leonina usually flee into the canopy, while M. n. nemestrina typically descends from the trees and flees on the ground. Except for M. n. pagensis in Kepulauan Mentawai, the geographic ranges of liontails and pigtail subspecies overlap those of several other species of macaques. M. silenus inhabits forest tracts in southwestern India that also are inhabited by M. radiata. The combined range of M. n. leonina and M. n. nemestrina broadly overlaps that of M. fascicularis; however, these species are at least partly segregated ecologically by the preference of M. n. leonina and M. n. nemestrina for inland evergreen rain forest and that of M. fascicularis for marginal mixed secondary growth habitats. The same ecological segregation may also apply to the relationship between M. n. leonina and M. mulatto where their ranges narrowly overlap (cf. below, fig. 28, and Fooden, 1971a, fig. 2). The combined range of M. n. leonina and M. n. nemestrina narrowly overlaps that of M . arctoides, which has been collected in the same forests as M. n. leonina. Although the range of M. n. leonina is almost completely allopatric with that of M. assamensis, these species have been collected together in a few forest tracts along the frontier between their ranges. HYBRIDIZATION Information available concerning hybrid matings that involve lion tails or pigtail subspecies indicates a relatively low level of hybrid viability and fertility (table 12). Of five reported intergeneric matings, one failed to produce conceptions, three produced hybrids that died in infancy, and one produced a hybrid that survived less than two years. Of 48 interspecific and intersubspecific hybrid conceptions reported, at least nine resulted in stillbirths or produced young that died in infancy; 11 of the hybrids are known to have reached the age of sexual maturity, and only three of these are known to have produced living offspring. All intergeneric crosses that involve liontail or pigtail macaques are restricted to members of the 42 chromosome group of cercopithecine genera (Chiarelli, 1967, p. 162). All interspecific crosses are restricted to macaque species in which the glans penis is bluntly rounded and the uterine cervix and cervical colliculi are moderately large (see above). Because of the size difference between sexes in catarrhine monkeys, it is not surprising that most hybrid matings of M. n. nemestrina with smaller-sized monkeys {M. silenus, M. n. leonina, M. fascicularis, Cercocebus sp.) are between a female of M. n. nemestrina and a male of the smaller species or subspecies; conversely, all four crosses with larger-sized species (Papio spp., Mandrillus sp.) involve a male M. n. nemestrina and a female of the larger species. The high incidence of hybrid matings reported between pigtail macaques and either M. fascicularis or M. mulatto may merely reflect the fact that M. fascicularis and M. mulatto are the macaque species most frequently kept in captivity. Available evidence concerning Fi phenotypes is limited and of varying precision and reliability (table 13). This deficiency is unfortunate because such evidence would be useful in interpreting the genetics and evolution of taxonomic characters in liontail and pigtail macaques. Although relevant data are somewhat ambiguous, the following genetic inferences are probably valid: (1) dorsal pelage color in M. fascicularis is dominant to that in M. silenus and M. n. 54 co s I I - co o •-1 _«' o I— 1 rH D" m a lO ._- . CO CO ^ 32 i* <-> <7> -' - 3J _ rH oo .S si oo -w -3 52 0 S c o m n * m ^ ; h A A 2 OS C a> . a «J ^cvf oo So*0? _ Oi ^ > m a CM* DO Oi CO --1 CN «~ o ~ CO 3 rH CN CN 00 tN * in -• CN --1 CN & a S2 >> in .S 3 t/i "£ "5 ? 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Anver 34, p. 4 B, p. 141 Gray, 1 _ aao'ia »^ 2 £ ^ 2 a to* ^r 50* ph* OOO SE;2o2»h c c ^ c c - 50 CN "g CN OJ OJ _- OJ _h 0 0 oj © c c c c CN ^H ^H 0 0 '-' OJ '** '*• '*• ** OJ 00 *^ 0> OJ ~ „•«••■• . •- •- oj - d _r d N N -r -• C c c c ** '-> g- ^h --1 a>o>o>o>o>o> a,UStf&.aCCO.CQCQCQCQCQ i«* '3 00" '3 00 01 Be OS ^ ^* "ti ■****« j^ 4) C« — — — c c c o> *i to ti to 1 g* Soj 0 5 « - » 1- )- u — - - - Ct, OQ co CO to e o> la V <— 0) 0> 0> — — — o> TfTf lOIOf C- 00 00 h n n f m m to t^ r~' pes «-t »-H 1-H .-H 1— • •-H H H N CN (N N (N CN 0* > .s o> ■- on a c J5 o fc J o £ E 5; c £ « * - -s ^ — :z 3 « « ■a m 3 * £ 1> > o £ X •5, CN -o h CO •-» e^- c- to 0 ** = «2 >» *» c co :S Q« cL (N - "• 'a C 4rf 1963, p. , p. 304 4; 1971 106 44 968, p. Hi, 1971 V, 1971, ,9 T3 CA eg t- in • • *-> a* ca ■ CO O, a . i- u x £> -> g .- . .in °> *- J= ~ 0 * «n,S c Sh 00 O a 35 ■^ 00 t- .»H rt >H .C C 0 r co a> - . !! 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"O e j e j c a c a I I I oc — t A O 2 "3 "o Q a o I I I 0 01 ex CO ■o CO • 0> a JS c e ■•J G 5 e c JS Js js O o - '— , >> 01 o£ c <~ 0» O *i - w •D O 'Z c .Q 0) x a 4) 0) 0> - 1.1 c : M c 0) CO a'> 3 X oi n, a 2 CO X5 « OJ -o C X c o a >. o ^ c 9 % 2 a a iS c a oi ?] x Oi o> a — >> 2 o c c £ * x Ut*C c c 'C X a >> o"S 5 E *- 3 >.£ x a o S b. H a o 5 58 FOODEN: LIONTAIL & PIGTAIL MACAQUES 59 nemestrina; (2) tail length and carriage in M. n. nemestrina is dominant to that in M. silenus; (3) tail length in M. fascicularis is not dominant to that in M. silenus and M. n. nemestrina (hybrids intermediate). The number of genetic loci or alleles that underlie each of these characters is uncertain. FOSSIL RECORD All known fossils and subfossils that are referable to the liontail-pigtail group have been collected in Quaternary deposits in the Malay Peninsula and Greater Sunda Islands, in or near the range of living M. n. nemestrina. In the Malay Peninsula, macaque subfossils are part of human food remains collected in floor deposits in a rock shelter (Gua Madu) in southern Kelantan, West Malaysia. According to Chasen (in Tweedie, 1940, p. 7), these food remains, which are not individually described or listed, include bones and teeth of M. n. nemestrina, langurs (2 spp.), bamboo rat, Malay bear, wild pigs (2 spp.), barking deer, sambar deer, and wild ox; no specimens of M. fascicularis are reported. Associated Hoabinhian artifacts indicate that the floor deposit in this shelter was formed about 4000-10,000 B.P. In the Greater Sunda Islands, identified macaque subfossils and fossils consist exclusively of fragments of jaws and teeth. Specific determination of these specimens depends essentially on tooth size. Based on macaque species and subspecies that now inhabit the area, subfossils or fossils with large teeth are identified as M. n. nemestrina, and those with small teeth are identified as M. fascicularis. Most specimens with intermediate size teeth are indeterminable. In west-central Sumatra, 124 isolated subfossil teeth have been collected in three caves (Djamboe, Lida Ajer, Sibrambang) in the Padang highlands. Of these, 102 are identified as M. n. nemestrina (82 per cent), 7 are identified as M. fascicularis (6 per cent), and 15 are indeterminable (12 per cent) (Hooijer, 1962a, p. 57); species proportions are reasonably consistent in all three caves. The age of these cave deposits is estimated to be prehistoric Holocene (Hooijer, 1962a, p. 4). The agency responsible for assembling the large accumulations of teeth of macaques and other mammals in these caves is unknown (Hooijer, 1948, p. 187). In Borneo (Niah Caves, northern Sarawak) 76 identifiable subfossil macaque jaws and teeth have been collected in strata with 60 FOODEN: LIONTAIL & PIGTAIL MACAQUES 61 TABLE 14. Stratification of M. n. nemestrina and M. fascicularis subfossils collected in Niah Caves, northern Sarawak (Hooijer, 1962b, pp. 441-443). Depth below Approximate Number of subfossils surface (inches)l C14 age (B.P.)2 M. n. nemestrina M. fascicularis 0-24 0 -10,000 4(16%) 21(84%) 25- 49 10,000-20,000 0 37(100%) 50- 74 20,000-30,000 0 7(100%) 75-102 30,000-40,000 1(17%) 5(83%) Totals 53 (7%) 70 (93%) 1- Specimens tabulated according to midpoint value of depth interval reported. 2- See Harrisson, 1958, p. 563. 3- Excludes one specimen without stratigraphic data. Carbon 14 dates extending back to about 40,000 B. P. (Harrisson, 1958, p. 563). Of these subfossils, six are identified as M. n. nemestrina (8 per cent) and 70 as M. fascicularis (92 per cent) (Hooijer, 1962b, p. 440). M. fascicularis specimens strongly preponderate at all stratigraphic levels (table 14). All of these specimens apparently are human food remains (Medway, 1964, p. 34). In eastern Java 28 fragments of macaque jaws and teeth have been collected in early Holocene deposits in three caves (Gua Djimbe, Gua Ketjil, cave near Wadjak; Hooijer, 1962a, pp. 41, 54). All tooth dimensions in these specimens closely correspond to those in M. fascicularis mordax, the living Javan subspecies of crab- eating macaque (tables 16, 17). The agency responsible for accumulation of these subfossils is unknown. The composition of subfossil macaque faunules in the Sunda area is compared with that of modern faunas in Table 15. In TABLE 15. Relative abundance of M. n. nemestrina (n) and M. fascicularis (f) in modern faunas and in subfossil faunules (see text). Age of fauna Malay Peninsula Sumatra Borneo Java Modern /">n , On f>n fXn)1 0-10,000 B.P.2 ny(f)1 n>/" f>n 10,000-20,000 B.P.3 fXn)1 f>(")1 20,000-30,000 B.P. fXn)l ~ 30,000-40,000 B.P. - />„ 1- Species absent. 2- Includes prehistoric Holocene 3- Includes early Holocene. I 8. r-* op op ■ ClXhW 00 r^ 3 c ^ r^ oo ** m rf O I CO CO e cd B 0 C DC i 4> I gg '3 a. c 41 a e is a 3 0 ■— E o -* ^-s 4> la E £ _ a c 01 - £ £ jj >- a. 3 M •r « 0 ^ II -C (C £ = u = ~ (A X. D < M H o ~ B OS o> -h OS CO CO t^ t^ OIOhiO |H CO co t- co m CO CO — as co I CO tT CO in rf CO tT CO Ot ot*o »>o o» »o CN ^, CS CO CO ^ CO ^,"^ « JC > H Of* *j rrt •*-» 4) O 0 0 5 E 2 .2 I ? 8-§.SJ SQS a 5 m a .c c o CO a c * « 5^5 5 2 w 35 -r OS OS OS* SO in x ■x ~ ■o eg C! ft* Cl t* 0 (0 -x 99 99 OJ pm 9J 4a s a 0 >- la '5 0 ac — OS OS til 41 B in iri X x X iC 2 m a a X a a. a 08 » (9 • ■ es M ~j 99 C>1 — tN ~ -x iC CC re te 0 99 99 ~ *r 99 — pH n g ^ X > la X o * JO .O '5 "3 0 0 0 < ac oa i < x ^h CN CO TT m CO 62 0) I CO a •v c « V c w c o CO Q, II o o * I < o> ^ cn co - 01 00 01 :z? ;^ :^ q c :; 0 0 i) - 0 I Q K < S 63 64 FIELDIANA: ZOOLOGY, VOLUME 67 modern Malayan, Sumatran, and Bornean faunas, M. fascicularis is approximately 10 times more abundant than M. n. nemestrina (below p. 103). In Java M. fascicularis is the only macaque now present (Chasen, 1940a, p. 65). Subfossil faunules collected in Borneo (late Pleistocene-subrecent) and Java (early Holocene) are similar in composition to the respective modern faunas; those collected in the Malay Peninsula and Sumatra (prehistoric Holocene) are not. Taken at face value, the Malayan and Sumatran subfossil data would indicate that, within the past 10,000 years, M. n. nemestrina was more abundant than M. fascicularis in the western part of the Sunda area, which is the reverse of the present situation in this area. However, this indication of recent local reversal of relative abundance may be misleading, if the accumula- tion of macaque remains in the Malay Peninsula and Sumatra has been subject to some unknown selective bias (cf. Hooijer, 1962b, p. 447). Hopefully, further collecting in Sunda area caves and rock shelters may help to resolve this intriguing and important problem in macaque paleozoogeography. In addition to Holocene subfossils, mid-Pleistocene fossils that are referable to Macaca also have been collected in eastern Java (Bangle, Punung, Sangiran, Saradan, Trinil; Deninger, 1910, p. 1; Stremme, 1911, p. 140; Badoux, 1959, p. 88; Hooijer, 1962a, p. 50; Hooijer, 1964, p. 76). These Pleistocene fossils, some of which originate in fissure deposits with those of Homo erectus, are among the oldest known macaque remains in Southeast Asia. Dental measurements in 14 recorded specimens (tables 16, 17) are not completely consonant with measurements in either living Javan M. fascicularis mordax or living Sumatran and Bornean M. n. nemestrina (and still less with those in living Sumatran and Bornean M. fascicularis; Hooijer, 1962a, p. 58). Although dimen- sions of anterior teeth in these Pleistocene Javan fossils are approximately the same as in living M. f mordax (and in Holocene subfossil M. f mordax', tables 16, 17), breadth of the last premolar and all three molars is nearer to that in M. n. nemestrina. In the best preserved Pleistocene specimen (Dub. No. 3785, see table 16), breadth of the last two molars is 8-9 per cent greater than in the largest measured specimens of M. f. mordax and well within the range of molar breadths in M. n. nemestrina. The relationship of the Pleistocene Javan fossil macaque to living M. n. nemestrina and M. f. mordax has been carefully reviewed by Hooijer (1962a, p. 51). In interpreting these data, FOODEN: LIONTAIL & PIGTAIL MACAQUES 65 Hooijer notes that the ratio of premolar breadth to molar breadth in the fossil macaque is less than in living M. n. nemestrina and postulates that this would not be true in the Pleistocene ancestor of M. n. nemestrina. Accordingly, he rejects a relationship to M. n. nemestrina and concludes that the fossil macaque probably is an ancestral form of M. f. mordax, in which the premolar-molar breadth ratio was greater than in the living form. However, the a priori assumption that the ratio of premolar breadth to molar breadth in the Pleistocene ancestor of M. n. nemestrina would not be less than in living M. n. nemestrina may not be valid. The relatively broad premolars in living M. n. nemestrina may be a recently evolved condition that is related to allometric increase in size of the rostrum and anterior part of the dental arch (figs. 8, 9). Cranial and dental characters in the stock ancestral to M. n. nemestrina may have been similar to those in M. n. leonina, which has tooth proportions that are fairly close to those in the Pleistocene Javan fossil macaque (tables 16, 17). Available evidence permits two plausible interpretations of the Javan fossil macaque. This Pleistocene macaque may be ancestral to Holocene and living M. f. mordax, in which case relative premolar breadth in this line has undergone evolutionary decrease; this interpretation implies that the pigtail stock was absent from Java during the Pleistocene, as well as during the Holocene and present. Alternatively, the Pleistocene macaque may be ancestral to living M. n. nemestrina, in which case relative premolar breadth has undergone evolutionary increase from proportions similar to those in living M. n. leonina; this interpretation implies that the pigtail stock inhabited Java during the Pleistocene and subsequently became locally extinct, as did the siamang, orangutan, Malay bear, elephant, and tapir (Badoux, 1959, p. 129). 65 EVOLUTION AND DISPERSAL Liontails, pigtail subspecies, and Celebesian stumptail species are allopatric and apparently constitute a closely interrelated phylogenetic unit, the silenus-group. (If subgeneric recognition of this group is warranted, the tautonym Silenus Goldfuss, 1820, is available and valid.) The close relationship between liontails and pigtails originally was inferred from similarities in pattern of hair growth, morphology of female sexual swelling, and structure of tail (Pocock, 1926, p. 1571). The relationship between Sundaic pigtails (M. n. nemestrina) and Celebesian stumptails was inferred from evidence of external proportions, structure of baculum, and geographic distribution (Fooden, 1969a, p. 61). The present study further indicates that species and subspecies in the silenus-group are similar in form and proportions of the glans penis (above, p. 33); these macaques also are similar in general form of estrous sexual skin swelling (above, fig. 11; Pocock, 1926, p. 1547; Fooden, 1969a, p. 13). Judging from penial and cervical morphology, the silenus-group probably is more closely related to M. sylvanus and to the fascicularis-group (M. fascicularis, M. mulatto, M. cyclopis, M. fuscata) than to either the sim'ca-group (M. sinica, M. radiata, M. assamensis, M. thibetana) or M. arctoides (above, pp. 32, 33). The close relationship of species in the silenus-group evidently is not revealed by their chromosome morphology. Karyologically, Af. silenus and M. nemestrina are less similar to each other than each is to other macaque species (Chiarelli, 1962, pp. 408-412; Schmager, 1972, p. 481). Geographic distribution of species and subspecies in the silenus- group (fig. 180 closely corresponds to major subdivisions of the Oriental faunal region (Sclater and Sclater, 1899, p. 131, pi. 5). The range of M. silenus is in the Indian Subregion, that of M. n. leonina is in the Indochinese Subregion (extended to include the Isthmus of Kra), and that of M. n. nemestrina is in the Sundaic Subregion (for name of this subregion, see Johnson, 1964, p. 69; Bullock, 1964, p. 212). The insular range of M. n. pagensis in Kepulauan Mentawai is 66 FOODEN: LIONTAIL & PIGTAIL MACAQUES 67 Present group Fig. 18. Hypothetical stages in evolution and dispersal of species and subspecies in the silenus-group. at the edge of the Sunda Shelf, marginal to the Sundaic Subregion. The range of Celebesian stumptail species is in the transition zone between the Oriental Region and the Australian Region. The zoogeographic hypothesis presented below (schematically illustrated in fig. 18) is proposed as a first approximation of the Quaternary history of the silenus-group. Obvious problems in zoogeographic interpretation are posed by deficiencies in the fossil 68 FIELDIANA: ZOOLOGY, VOLUME 67 record of this group (see above, p. 60) and by incomplete information concerning Pleistocene changes in the geography, climate, and vegetation of the Oriental Region. Major topics considered in the following discussion are: 1. Center of origin of si/enus-group; main pattern of dispersal. 2. Disjunct distribution of liontails and pigtails. 3. Dispersal of silenus-group to zoogeographically peripheral Kepulauan Mentawai and Celebes (Sulawesi). 4. Geographic contiguity and morphological discontinuity between M. n. leonina and M. n. nemestrina. 5. Geographic isolation and morphological similarity within M. n. nemestrina. In species and subspecies of the silenus-group there is a clear geographic pattern of progressive decrease of tail length from west to east. Relative tail length is 0.57-0.75 in M. silenus, 0.28-0.46 in M. n. leonina and M. n. nemestrina, 0.24-0.34 in M. n. pagensis, 0.05- 0.12 in southern and central Celebesian species, and 0.03-0.07 in northern and northeastern Celebesian species (fig. 2; Fooden, 1969a, p. 15). Assuming that a long tail is primitive in macaques, as in other monkeys, this west-to-east trend may be interpreted as a trend from primitive to derived. Similarly, the relatively short rostrum in skulls of M. silenus and M. n. leonina probably is more primitive than the long rostrum in M. n. nemestrina (fig. 8); the most specialized skulls in this group, or in any species of macaque, are those of M. nigrescens and M. nigra in northeastern Celebes (Fooden, 1969a, p. 30). The geographic pattern of morphological variation of skull and, especially, of tail length tends to suggest that, among living species and subspecies, Indian M. silenus is morphologically most similar to the ancestral stock of the silenus- group and that Indochinese and Sunda area species and subspecies represent successive stages in evolutionary development of the group. Accordingly, the starting point of the present hypothesis is that the silenus-group originated in the Indian Peninsula (fig. 18a) and subsequently dispersed to the Indochinese Peninsula and the Sunda area (fig. 18b). The initial direction of dispersal from the Indian Peninsula presumably was northward (fig. 18b). This may have given impetus to tail reduction in the silenus-group, since higher latitudes are known to be correlated with shorter tails in two previously studied groups of macaques (Fooden, 1971b, p. 72; 1972, p. 310). However, FOODEN: LIONTAIL & PIGTAIL MACAQUES 69 relative tail length in the unknown original progenitor of the silenus-group probably was greater than in M. silenus (0.57-0.75), judging from external proportions in most monkeys (Napier and Napier, 1967, p. 413). M. fascicularis, which is the only living macaque with a long tail (relative length 0.79-1.48; Fooden, 1971a, p. 27) and a rounded glans penis (see above) may serve as a morphological model of the unknown pre-silenus ancestor of the group. The present range of M. silenus in southwestern India is separated from the western border of the range of M. n. leonina in Assam and Burma by a gap that is about 2000 km. broad. At present, natural vegetation in the intervening area is deciduous or semi-deciduous forest (Kuchler, 1964, pp. 17, 141), which evidently is not suitable habitat for the si/enws-group (fig. 17). Past dispersal of the silenus-group across this area, as assumed in the present hypothesis (fig. 18b), probably implies that the area formerly supported evergreen forest, which, in turn, implies more continuous rainfall than at present. It seems likely, therefore, that dispersal of the silenus-group from the Indian Peninsula to the Indochinese Peninsula occurred during a Pleistocene pluvial interval (fig. 18b), and that subsequent disjunction of Indian and Indochinese sections of the group dates from a succeeding arid interval (fig. 18c). The same series of Pleistocene climate changes probably also accounts for similar disjunct distributions in Indian and Indochinese species of langurs, lorises, giant squirrels, civets, and mouse deer (Pocock, 1939, p. xxvi; cf. Kurup, 1966, p. 19). Analogous zoogeographic interpretations based on Pleistocene rainfall variation previously have been proposed to account for distributions of tropical bird species in Southeast Asia (Stresemann, 1939, p. 416) and South America (Haffer, 1969, p. 131). Recent human interference with natural vegetation patterns probably has played a relatively minor role in exaggerating disjunction of the ranges of M. silenus and M. n. leonina. Isolation of Indian and Indochinese sections of the silenus- group presumably led to morphological differentiation of ancestral liontails and pigtails. Populations of the ancestral pigtail stock evidently dispersed to Kepulauan Mentawai and to Celebes, where subsequent isolation resulted in differentiation of a stumptail stock (fig. 18c). In Kepulauan Mentawai, M. n. pagensis is part of a strikingly peculiar mammal fauna (Kloss, [1928], p. 806) that also 70 FIELDIANA: ZOOLOGY, VOLUME 67 includes an endemic species of gibbon, Hylobates klossii. Because dispersal of gibbons evidently is strongly inhibited by water barriers (Groves, 1967, p. 281; Fooden, 1969b, p. 640), it seems likely that ancestral populations of the distinctive Kepulauan Mentawai fauna, including ancestors of M. n. pagensis and H. klossii, reached this island group by way of a transitory land connection with the adjacent Sunda Shelf. Conversely, absence of gibbons from Celebes, and the generally depauperate character of the Celebesian mammal fauna (Darlington, 1957, p. 519), suggests that original dispersal of the silenus-group to this island was across a water gap, probably by rafting (Fooden, 1969a, p. 64). Dispersal of the silenus-group to Kepulauan Mentawai and Celebes probably occurred during maximal Pleistocene glacial lowering of sea level, to about 200 m. below present sea level (Jongsma, 1970, p. 151), when the Sunda Shelf was most extensively exposed. Relative richness of mammal faunas in Kepulauan Mentawai and Celebes evidently has been controlled by depth of the straits that separate these insular areas from the Sunda Shelf. Minimum depth of the strait between the Sunda Shelf and Kepulauan Mentawai is about 200 m., and that between the Sunda Shelf and Celebes is about 600 m. (Atlas van Tropisch Nederland, 1938, pi. 3). The absence of otherwise widespread M. fascicularis from both Kepulauan Mentawai and Celebes suggests that this macaque had not yet arrived in the Sunda area at the time that maximal Pleistocene lowering of sea level made possible dispersal of the silenus-group to these outlying islands (cf. Dorst, 1953, p. 312). The border between the ranges of M. n. leonina and M. n. nemestrina almost exactly coincides with the location of the Khlong Marui Fault (Garson and Mitchell, 1970, p. 45), which cuts across the sharp bend in the Thai-Malay Peninsula. Significantly, the Khlong Marui Fault area is a border zone for many groups of animals and plants. Zoological evidence places the subregional boundary between the Indochinese and Sundaic vetebrate faunas about 250 km. north of the Khlong Marui Fault (Chasen, 1940a, p. ix), and botanical evidence places the corresponding boundary between the Continental Southeast Asiatic and Malesian floras about 250 km. south (Kloss, 1920, p. 79; van Steenis, 1950, p. lxxii; Keng, 1970, p. 51). The broad pattern of faunal and floral differentiation, including differentiation of M. n. leonina and M. n. nemestrina, that centers around the Khlong Marui Fault probably implies the former FOODEN: LIONTAIL & PIGTAIL MACAQUES 71 existence in this area of a physical barrier that blocked genetic exchange between populations on either side of the barrier. Geographic considerations suggest that this barrier may have been a local marine transgression that transected the Thai-Malay Penin- sula; this transgression may have been either eustatic or tectonic in origin. Geological evidence pertaining to such a transgression is ambiguous (Scrivenor, 1911, p. 9; 1949, p. 114; Viji Stresthaputra et al., 1951, p. 40; Ho, 1960, p. 18; Kobayashi, 1960, p. 142). Judging from degree of morphological differentiation, isolation of the M. n. pagensis pigtail stock in Kepulauan Mentawai probably dates from about the same time as disjunction and isolation of the M. n. leonina and M. n. nemestrina pigtail stocks. Dispersal of the ancestral Celebesian stumptail stock to peripheral islands in the hypothetical Pleistocene Celebesian archipelago (Fooden, 1969a, p. 64) may have occurred during this same stage in the evolution of the silenus-group (fig. 18d). Subsequent to the morphological differentiation of M. n. leonina and M. n. nemestrina, the postulated former barrier between their ranges evidently disappeared, bringing these two pigtail stocks into secondary contact (fig. 18e). Judging from the present restricted distribution of morphological intermediates (table 20), it appears that only limited gene exchange has subsequently occurred between these two subspecies. The geographic change that brought M. n. leonina and M. n. nemestrina into secondary contact may also have provided an opportunity for the apparently recent entry of M. fascicularis into the Sunda region (see above). About 20,000 years ago, during the latest Wisconsin/Wurm glacial advance, sea level in the Sunda area evidently was at least 120 m. lower than at present (Van Andel et al., 1967, p. 743; Emery, 1969, p. 114; Jongsma, 1970, p. 150). As a result of this most recent (but not most extensive) exposure of the Sunda Shelf, the Malay Peninsula, Sumatra, Bangka, and Borneo, which are now isolated parts of the range of M. n. nemestrina, were interconnected by dry land (fig. 18e). This relatively recent continuity of range presum- ably promoted partial homogenization of the gene pool of M. n. nemestrina, which is evident in the morphological similarity of now disjunct insular and peninsular population of the subspecies (fig. 22). With the postglacial rise of sea level, ranges of species and subspecies in the silenus-group attained their present configuration 72 FIELDIANA: ZOOLOGY, VOLUME 67 (fig. 180- Two negative features of these ranges are somewhat difficult to interpret. These two enigmatic features are (1) the absence of M. n. leonina from apparently suitable evergreen forest habitats (fig. 17) in central and northern Laos and North Vietnam (Osgood, 1932, pp. 208-211) and (2) the absence of M. n. nemestrina from similar suitable habitats in Java, which is on the Sunda Shelf. These absences may indicate that unknown past physical or biological barriers prevented the si/enus-group from dispersing to the northeastern part of the Indochinese Peninsula and Java or, alternatively, that populations of the group formerly inhabited either or both of these areas and subsequently became locally extinct. Problematical fossils that may record the presence in Pleistocene Java of a population ancestral to M. n. nemestrina are discussed above (p. 64). SPECIES ACCOUNTS Macaca silenus (Linnaeus, 1758) [Simia] Silenus Linnaeus, 1758, p. 26 — part, subsequently restricted by Buffon (1766, p. 169); external characters; distribution, Asia: Ceylon, Java, etc; the specific name is a nominative singular masculine noun (Latin name of bearded woodland diety) standing in apposition to the feminine generic name. Simia Silenus [variety A]: Schreber, 1774, pp. 87, 186, pi. 11 (animal ex Buffon) - part (also includes references to Presbytis sp.); external characters; habits; distribution, Ceylon, Coromandel Coast, possibly East Indies. Sim [ia] silenus: G. Cuvier, 1817, p. 108 — species included among "Les Macaques"; distribution, Ceylon. Simia silanus [sic]: F. Cuvier, 1822b, p. 2, pi. (animal) — external characters of captive, origin unknown; Indian name, Nil-Bandar. Cercopithecus silenus: Kerr, 1792, pp. 32d, 64-part, lion tail macaque assigned to subspecies Cercopithecus silenus albibarbatus Kerr, 1792. Papio silenus: E. Geoffrey, 1803, p. 27 — zoo specimen listed; distribution, Ceylon. [Ogilby], 1838, pp.364, 386, fig. (animal) — external charcters; taxonomic relationships. Pithecus silenus: Desmarest, 1817, p. 321 — external characters; distribution, Ceylon and neighboring lands. P[ithecus] silenus: Blainville, 1839, p. 30 — citation of possible reference in ancient Greek literature. Macacus silenus: Desmarest, 1820, p. 63, pi. 10 (animal ex Buffon) — external characters; distribution, Ceylon. Bennett, 1829, p. 20, fig. (animals) — external characters; habits in captivity; distribution, Malabar Coast, Ceylon. Blyth, 1851, p. 154 — vernacular name not Wanderoo, which instead applies to Presbytis sp. Anderson, 1879, p. 93 — external and cranial characters; distribution, Western Ghats from near Goa to Cape Comorin. Blanford, 1888b, p. 16, fig. 5 (head) — external characters; nomenclature; distribution, Western Ghats northward to 14°N. [Macaca] Silenus: Thomas, 1911, p. 126 — nomenclature. [Macaca] silenus-group: Miller, 1933, p. 5 — key to external characters. Macaca silenus: Pocock, 1939, p. 66, pi. 4 (animals), figs. 16 (head ex Blanford, 1888b), 17 (skull) — external and cranial characters; taxonomy, closely related to M. nemestrina leonina: Pocock, 1939; ecology; distribution, Western Ghats in Travancore and Cochin, reportedly from Cape Comorin northward to 14°N. 73 74 FIELDIANA: ZOOLOGY, VOLUME 67 Cynoceph [alus] Silenus: Schinz, 1821, p. 114 — new combination; taxonomy; distribution, Ceylon. I [nuus] Silenus: Wagner, [1839], p. 141 — part (also includes references to Presbytia sp. ); external characters taxonomy; distribution, Ceylon. Blyth, 1847, p. 731 — not native to Ceylon according to Colombo resident Dr. R. Templeton; said to be abundant in Travancore and Cochin. I nuus silenus: Jerdon, 1867, p. 10 - external characters; taxonomy; field observations; distribution, Western Ghats northward to 14°N. V[etulus] Silenus: Reichenbach, 1862, p. 125, figs. 321-325 (animals ex authors) - external characters; habits in captivity; distribution, Malabar. Ouanderou: Buffon and Daubenton, 1766, p. 169, pi. 18 — part (also includes references to Presbytis sp.); restriction of species and virtual designation of neotype; external characters and measurements; habits; distribution, Ceylon, Malabar Coast (implied). F. Cuvier, 1837, liv. 70, pi. (animal) — taxonomic history; distribution, penisular India. Lion tailed monkey [variety a ]: Pennant, 1771, p. 109, pi. 13a (animal) — characters and habits of captive exhibited in London; compared with neotype of M. silenus (Linnaeus, 1758). [Cercopithecus] Vetulus Erxleben, 1777, p. 25 — new name based on (1) Simia silenus Linnaeus, 1758, (2) Ouanderou: Buffon, 1766, (3) Lion tailed monkey: Pennant, 1771, (4) various characterizations of Presbytis sp. and Colobus sp.; type-locality, "Zeylona, Asia, Africa." Boddaert, 1785, p. 57 — restricted to liontail macaque; distribution, Ceylon, Coromandel Coast. Fischer, 1829, p. 28 — a synonym of Simia silenus Linnaeus, 1758. Simia ferox Shaw, 1792, p. 69, pi. opposite p. 71 (animals) — new name based on (1) Simia silenus Linnaeus, 1758, (2) Ouanderou: Buffon, 1766, (3) Lion tailed monkey; Pennant; 1771, (4) characterization by Ray (1693, p. 158) of Presbytis senex; type locality, "East Indies, ... Ceylon.. .also said to be found in the interior parts of Africa." Shaw, 1800, p. 30, pi. 16 (animal ex Buffon) — external characters; taxonomy. Blanford, 1888b, p. 16 — a synonym of Macacus silenus. [Simia] Ferox: Griffith, 1821, p. 105, pi. (animal) — a synonym of Simia silenus Linnaeus, 1758. M[acaca] ferox: Wroughton, 1918, p. 554 — nomenclature; external characters; type- locality, Ceylon. Cercopithecus silenus albibarbatus Kerr, 1792, pp. 32d, 64 — new subspecific name based on (1) Simia silenus Linnaeus, 1758, (2) Ouanderou: Buffon, 1766, (3) Lion tailed monkey: Pennant, 1771, (4) characterization by Ray (1693, p. 158) of Presbytis senex; type- locality, "Ceylon and the rest of India." Pithecus albibarbatus: Elliot, [1913], p. 218, pi. 8 (animal; not cranial pi. 24, misidentified skull of Macaca radiata; see Pocock, 1939, p. 68) — external characters; taxonomic history. Silenus albibarbatus: DeBeaux, 1923 p. 34 — external characters. [Simia] Veter: Audebert, 1798-1800, Fam. 4, p. 4 (not Linnaeus, 1766) - said to be a variety of Ouanderou: Buffon, 1766. Silenus veter: J. E. Gray, 1843 P- 8 (not Linnaeus, 1766) —specimens listed. FOODEN: LIONTAIL & PIGTAIL MACAQUES 75 [Simla] leonina: G. Cuvier, 1817, p. 108 (not Shaw, 1800) — external characters; name incorrectly attributed to Linnaeus and Gmelin; distribution, Ceylon. Type. — None preserved. A captive specimen (fig. 19) described and figured by first reviser Buffon (in Buffon and Daubenton, 1766, p. 169, pi. 18) is now regarded as neotype (see discussion below). As previously noted (Blanford, [1888a], p. 620; Thomas, 1911, p. 126; Elliot, 1913, p. 219; J. A. Allen, 1916, p. 50; Pocock, 1939, p. 66), it is difficult or impossible to identify with certainty the original objective basis of Simia silenus Linnaeus, 1758 as described in the tenth edition of Systema Naturae (p. 26). Linnaeus' concept of this taxon apparently was derived solely from published accounts of earlier naturalists, not from direct examination of a specimen. In the original description, quoted in full below, Linnaeus (1758, p. 26) explicitly acknowledges that the species is obscure and that most of its characters are unknown: Silenus. 5. S[imia] caudata barbata, corpore nigro, barba nivea prolixa. Simia Callitriches magnitudine cynocephalorum. Alp. aegypt. 242? Habitat in Asia: Zeylona, Java & c. Species obscurior, ignotis Pedum unguibus aliisque plurimis attributis. In this account the skimpy diagnosis (tailed monkey with black body and full white beard) is generally applicable to the Indian liontail macaque. However, the bibliographic citation and the assigned geographic range refer to at least two other species of cercopithecids. The doubtful synonym cited (with a question mark) from Alpinus (1735, Kistoriae Aegypti naturalis) is based on a figure of a 6/ac/s-bearded monkey or baboon that reportedly originated in Egypt. The indefinite geographic distribution given by Linnaeus (Asia: Ceylon, Java, etc.) may have been partly influenced by Ray's (1693, p. 158) characterization of a monkey in Ceylon as "Cercopithecus niger barba incana promissa" ( = Presbytis senex); it should be noted, however, that Ray's account of this Ceylonese monkey is not explicitly cited by Linnaeus, although Ray's work is cited in synonymies of eight other monkey species described in the tenth edition of Systema Naturae. The earliest known unequivocal association of Simia silenus Linnaeus, 1758, with an actual specimen is in Buffon 's (in Buffon and Daubenton, 1766, p. 169, pi. 18) account of the "Ouanderou" or "Lowando." This report is based on a captive liontail (fig. 19) that was exhibited at a fair in France. Although the owner of the captive /'/ XI III /'. fit .Iff /w Ol'ANDHHOr. HvMi Fig. 19. Ouanderou: Buffon (in Buffon and Daubenton, 1766, pi. 18), neotype of M. silenus Linnaeus, 1758. 76 FOODEN: LIONTAIL & PIGTAIL MACAQUES 77 asserted that it originated in South America, Buffon dismissed this assertion as mere showmanship. He surmised instead that the monkey probably was native to Ceylon. The sources of this erroneous geographic conjecture evidently are the faulty account of Simia silenus given by Linnaeus (1758, p. 26) and misinterpreted characterizations of Presbytia senex published by Knox (1681, p. 41, [1911 ed.]) and Ray (1693, p. 158), all of which are cited by Buffon in the synonymy of the "Ouanderou"; Buffon derived the name "Ouanderou" from the Ceylonese vernacular name cited by Knox for Presbytis senex. Along with the mistaken reference to Ceylon, Buffon (in Buffon and Daubenton, 1766, p. 171) also refers to the true geographic range of the liontail ("Malabar") in a footnote abstract of field notes published by P&re Vincent Marie, who observed this species while serving as a missionary in southwestern India. In the same year that Buffon published his account of the "Ouanderou," Linnaeus (1766, p. 36), in the twelfth edition of Systema Naturae, published a radically modified characterization of Simia silenus. In an obvious attempt to eliminate discrepancies between the characters and range of Simia silenus and those of the monkey cited from Alpinus, Linnaeus emended the diagnosis of Simia silenus in the twelfth edition to "S[imia] caudata barbata nigra, barba nigra prolixa" (beard black, not white, as in the tenth edition) and changed the distribution to "Habitat in Egypto" (not Asia: Ceylon, Java etc., as in the tenth edition). However, despite ambiguity in the original characterization of this species, and despite Linnaeus' subsequent contradictory redefinition, the name silenus has been applied to the Indian liontail macaque by virtually all authors since Buffon. To preserve stability of the specific name, Buffon (in Buffon and Daubenton, 1766, p. 169, pi. 18) may be regarded as first reviser of the imprecisely characterized composite nominal species Simia silenus Linnaeus, 1758, and Buffon 's restriction of this species to the Indian liontail macaque, represented by a described and figured captive specimen (neotype, fig. 19), may be regarded as decisive. According to this interpretation, Simia silenus Linnaeus, 1766, which is based on a black-bearded African monkey, is an unavailable junior homonym of Simia silenus Linnaeus, 1758, as restricted to the Indian liontail macaque by first reviser Buffon (1766). Type -locality. — Geographic origin initially given as "Asia: Zeylona, Java & c." (Linnaeus, 1758, p. 26); restricted to "Ceylan" 78 FIELDIANA: ZOOLOGY, VOLUME 67 and "Malabar" by first reviser Buffon (in Buffon and Daubenton, 1766, pp. 169-171); "No exact typical locality available," according to Thomas (1911, p. 126); type-locality designated "Egypt" by Elliot ([1913], p. 219); now corrected to Western Ghats, inland from Malabar Coast, southwestern India (see above). Distribution. — Restricted to the Western Ghats, a low mountain chain in southwestern peninsular India (fig. 20); known altitudinal range 300-1300 m. (Loci, 12). The northernmost known locality is Anshi Ghat (14°55'N), near Goa, and the southernmost is Kalakkadu Hills (8°25'N), near Cape Comorin. The range of this species evidently has been narrowly restricted at least since the middle of the nineteenth century (see below). Recently the range has been further reduced, almost to nil, as a result of local deforestation caused by man (Sugiyama, 1968, p. 284; C. A. Hill, 1971, p. 35). External characters (fig. 21). — Head and body length 457-481 mm. in 3 adult females, 508-584 mm. in 4 adult males; relative tail length (T/HB) 0.56-0.69 in 3 adult females, 0.57-0.75 in 4 adult males (table 18; fig. 2); weight 6.75 kg. in one male, presumably adult (Napier and Napier, 1967, p. 406). Fur blackish on crown, dorsal surface of trunk, arms, legs, hands, feet, and tail; side whiskers and beard long, pale ochraceous-gray, forming a conspicuous facial ruff; crown patch narrow anteriorly; skin of muzzle blackish, thinly covered with inconspicuous short whitish hairs; ventral surface of trunk thinly haired, grayish brown to brown. In adults the dorsal pelage is long and dense, 8-10 cm. long on the scapular region and 4 cm. long on the lumbosacral region. The fur at the tip of the tail in adult specimens examined consists exclusively of coarse guard hairs, which are sparsely distributed elsewhere in the pelage. These hairs usually are elongated and form a more or less distinct terminal caudal tuft (fig. 21), on which Pennant (1781, p. 183) based the name "lion tailed monkey." Hairs on the crown are 2-3 cm. long and form a whorl radiating from the vertex. More anteriorly, the dark frontal hairs are about 2 cm. long, posteriorly directed, and form a sharply defined wedge-shaped patch that is bordered anteriorly and laterally by short pale hairs of the lateral supraorbital region and long pale hairs of the facial ruff. In adults, pale hairs of the facial ruff may attain a length of 10-15 cm. Cranial characters (figs. 3, 5). — Skull relatively small (greatest length excluding incisors 102.3-112.6 mm. in 7 adult females, 128.5- FOODEN: LIONTAIL & PIGTAIL MACAQUES 79 Fig. 20. Locality records of Macaca silenus: closed triangles = specimens examined, open triangles = literature records. For details, see Gazetteer of Collecting Localities. Key to localities: 1. Anshi Ghat; Goa, near. 2. North Kanara. 3. South Kanara. 4. Coorg. 5. Kuttyadi Pass. 6. Wynaad. 7. Nilgiri Hills, 800-1300 m; Nilgiri Hills, western slopes. 8. Parambikulam Valley. 9. Cochin. 10. Cotengady Estate; Nelliampathy Hills; Palagapandy; Seetagundy Estate. 11. Anaimalai Hills, northern. 12. Anaimalai Hills, 800-1300 m.; Grass Hills; Varagaliyar. 13. Panniar. 14. High Wavy Mountain; Varushnaad Valley. 16. Periyar Lake, northern shore; Periyar Lake, south. 16. Manjolai Tea Estate. 17. Kalakkadu Hills; Naraikkadu Estate. 138.5 mm. in 7 adult males), rostrum relatively short, zygomatic arches broad (table 19; figs. 7, 8); supraorbital ridges thick and prominent; anterior surface of malar with a clearly defined infraorbital concavity. Habits and habitats. — This account is based on field observations published by authors listed below. Authors are cited TABLE 18. External variation in Af . silenus adults; collecting localities un- known except as indicated. Specimen No. or reference FMNH 98163 ZSBS 1959/264 Blanford, 1888°, p. 161 Blanford, 1888°, p. 16l AMNH 548372 Anderson, 1879, p. 93 Blanford, 1888b, p. 16* Blanford, 1888b, p. 161 Pocock, 1939, p. 67 1- Collected at Travancore. 2 Collected at Nelliampathy Hills. Head and Relative tail body (mm.) length (T/HB) Females 481 0.57 ?610 ? 0.39 457 0.69 457 0.56 Males 575 0.57 ?610 ? 0.42 533 0.64 508 0.75 ca. 584 0.63 TABLE 19. Cranial variation in Af. silenus adults (permanent dentition completely erupted); collecting localities unknown except as indicated. Relative Greatest zygomatic Rostral-postrostral Specimen No. length (mm.) breadth 1 Females ratio BM 10b 111.1 0.67 0.51 BNHS 52832 ca. 106.2 ca. 0.69 ca. 0.46 NHRM Z4288 108.4 0.71 — RMNH 1314 110.0 0.67 0.49 ZSBS 1959/264 112.6 0.65 0.49 ZSI 7752 102.3 0.71 0.40 ZSI7753 105.7 0.62 0.45 Means 108.0 0.67 Males 0.47 BNHS 5059/50643 134.8 0.69 0.61 BNHS 50622 128.5 0.68 0.57 FMNH 47322 135.4 0.69 0.60 MNHN 1925/8 136.6 0.67 — NHMB 1610/3931 138.5 0.68 — RMNH 2122 131.3 0.72 0.57 Means 134.2 0.69 0.59 1 Zygomatic breadth/greatest length. 2- Collected at North Kanara. 3- Collected at Palagapandy. 80 FOODEN: LIONTAIL & PIGTAIL MACAQUES 81 by means of the indicated abbreviations. Of these authors, Sugiyama (1968) provides the most detailed and useful information. B Baker in Blyth, 1859, p. 283 H Hutton, 1949, p. 690 J Jerdon, 1867, p. 10 Ka Karr, 1973, pp. 191-192 Ki Kinloch in Pocock, 1939, p. 68 Kr Krishnan, 1972, pp. 542-543 O'B O'Brien in Pocock, 1939, p. 69 P Poirier, 1970, p. 261 Sp Spence, 1921, p. 970 St Stonor, 1944, p. 591 Su Sugiyama, 1968, pp. 283-292 W-P Webb-Peploe, 1947, p. 629 M. silenus is a hill species restricted to dense evergreen forest (B, H, J, Ki, Kr, P, Sp, Su, W-P) or semi -evergreen forest in which trees are more than 20 m. tall (Su) at elevations between 300 m. (Anshi Ghat) and 1300 m. (Su). Within this limited habitat, the distribution of troops of M. silenus is sparse (H, Kr, O'B, St, Su, W- P). As early as 1867, an observer (J) noted that this monkey "frequents the most dense and unfrequented parts of the forest, . . . and I had often traversed the Malabar forests before I first fell in with it." In 1968 the total living wild population was estimated at fewer than 1000 individuals (Su) in a maximum habitat area of 50 km. X 750 km. The maximum number of individuals reported in a troop of M. silenus is 22 (Su). Other reports of troop size are: "four or five" (St), 6+ (Ka), "about a dozen" (Ki), 16 (Su), "twelve to twenty or more" (J), and "about twenty" (W-P). In two carefully studied troops, one with 16 members included 2 adult males and 7 adult females, and another with 22 members included 2 adult males and 10 adult females (Su). Solitary adult males have been observed fairly frequently (H, St, Su). Although M. silenus is primarily arboreal (Su), it occasionally descends to the ground: Once several were seen climbing slowly down one tree and walking along tbe ground to the next tree. This they climbed, investigated and came down to earth to repeat the process. (W-P) Similar terresterial excursions also are reported by two other observers (H, Su). M. silenus may occasionally venture into streams, since captive specimens are said to be fond of water and to swim well (H). In the trees M. silenus evidently is somewhat 82 FIELDIANA: ZOOLOGY, VOLUME 67 deliberate in its locomotor behavior (Ka), moving more leisurely and with less leaping than sympatric M. radiata, for example (H, Kr, W-P). When disturbed, M. silenus flees high into the canopy (Su). Like other macaques, M. silenus feeds chiefly on fruit, but augments its diet with a variety of other plant and animal foods. The species has been observed to eat wild figs (St). Another observer (Su) reports: The only important food.. .during the study period was a.. .chestnut-like fruit.. .In addition... they ate other fruits, nuts, flowers and young buds of many kinds of trees, insect larvae living under tree-bark, pith of cardamon stems and so on. In one troop that also fed on fruit, behavior apparently directed at locating arthropod food is described (Ka) as follows: Another adult climbed to the uppermost branches of a tall emergent tree where several dead branches projected above the surrounding canopy. Pieces up to a foot long were broken off and torn apart, presumably in search of arthropods in the rotting wood. The macaque moved lower in the tree before dismantling the branch and then moved to the upper branches again, peered into the broken stub and tore off another dead branch. During its morning feeding session, this troop evidently was dispersed over several hectares of forest (Ka). Information on breeding periodicity is ambiguous. A birth peak in January is indicated by observation (Su) in that month of three new-born infants and also by observation (Ka) in early March of one or two young about two months old. However, another observer (W-P) asserts that females carrying young are seen regularly in September. Likewise difficult to interpret is a report (Su) of 12 instances of sexual behavior, including four copulations, observed in January and February; if these were fertile matings, the young presumably would have been born in June or July. Encounters between adjacent troops of M. silenus apparently are usually hostile and characterized by aggressive vocal challenges: As the overlapping part of the ranges of the two troops had many food trees. ..both troops frequently came to them, and contact between the two troops was observed many times. They were antagonistic and troop 2 was dominant. When they came near, large adult males in each troop displayed by whooping loudly but no direct fighting was seen. Usually, troop 1 moved away after a short time of vocalization. (Su) Contradictory reports concerning interactions of M. silenus with sympatric primate species are discussed below. These reports Fig. 21. Macaca silenus, external characters. (Photos courtesy San Diego Zoo) 84 FIELDIANA: ZOOLOGY, VOLUME 67 are not readily harmonized. According to one observer (Su), M. silenus is tolerant of and subordinate to M. radiata: A troop of bonnet macaque [M. radiata] moved among liontailed macaques and sometimes even travelled following the latter. Liontailed macaques usually did not react to the appearance of bonnet macaques but sometimes moved away slowly from them. Although smaller, bonnet macaques were dominant. However, another observer (W-P) reports that M. radiata "will keep out of the way when the Lion-tailed Monkey is about." This report agrees with an early anecdotal account by Vincent Marie (in Buffon, 1766, p. 171), which also implies that M. silenus is dominant to M. radiata. Similarly, three observers (Ka, St, Su) indicate that M. silenus associates amicably with the Nilgiri langur, Presbytis johnii, while two others (H, W-P) report that contacts between these species are hostile: In the Animalais there are about as many Lion-tailed as Black monkeys [P. johnii] and some trespassing on each others' territory is inevitable. On these occasions some terrific battles take place in the trees, and there are casualities on both sides (W-P). These contradictory reports may signify that interspecific behavior is variable in these monkeys. Like other macaques (Fooden, 1969a, p. 52), M. silenus may form symbiotic associations with some species of birds. One observer (St) noted a troop of these monkeys "in a Ficus tree, oddly enough in company with the Great Indian Hornbill (Dichoceros bicornis)." A giant squirrel, Ratufa indica, that approached a female M. silenus with infant was responded to aggressively (Ka). Specimens examined — Total 59. INDIA: Cochin, 1 (skull only, BM); Cotengady Estate, 1 (BM); Malabar, 5 (1, NMS; 4, skins only, RMNH); Malabar Coast ?, 1 (skin only, ZSI); Nelliampathy Hills, 2 (skins only, AMNH); North Kanara, 2 (skull only, BNHS); Palagapandy, 1 (BNHS); Parambikulam, 1 (skin only, BM); Seetagundy Estate, 1 (skin only, BNHS). Imprecise localities: 5 (2 skins only, AMNH), 6 (3 skins only, BM), 3 (1 skin and postcranial skeleton only, 1 skin only, FMNH), 2 (skulls in skins, IRSN), 5 (2 skulls in skins, 1 skull only, MNHN), 1 (skull only, NHRM), 1 (skin only, NMS), 7 (3 skins only, RMNH), 3 (1 skin only, USNM), 1 (ZSBS), 10 (4 skins only, 1 skull only, ZSI). FOODEN: LIONTAIL & PIGTAIL MACAQUES 85 Macaca nemestrina (Linnaeus, 1766). (Synonymies under sub- species headings) Taxonomy. — Compared with liontail macaques, the three recognized allopatric forms of pigtail macaques (Inuus leoninus Blyth, Simia nemestrina Linnaeus, Macaca pagensis Miller) are obviously similar in tail proportions and general brownish pelage hue. These three forms of pigtails are dissimilar in pelage pattern, average size, cranial characters, bacular structure, arboreal-terrest- rial preference, facial markings, tail carriage, and plasma protein frequencies. Whether to regard these three pigtails (leonina, nemestrina, pagensis) as species or as subspecies depends ultimately on analysis of geographic variation of the characters that distinguish them (Mayr, 1963, p. 194). In the following paragraphs, overall geographic variation of adequately known characters in parapatric leonina and nemestrina is considered first. This is followed by detailed discussion of diagnostic characters in critical specimens of leonina and nemestrina that have been collected near the common border between their ranges. The taxonomic status of insular pagensis is considered last. Pelage pattern is relatively constant and distinctive in leonina and nemestrina. The pale dorsal surface and anteriorly narrow crown patch in all leonina specimens examined is readily distinguishable from the dark dorsal surface and broad crown patch in all but four geographically marginal specimens of nemestrina (see below). In leonina specimens, preauricular sidewhisker hairs are elongate and pale from base to tip; in nemestrina specimens, these hairs are short and dark tipped. Although pelage pattern varies individually in leonina and nemestrina, this variation is not geographic. There is no apparent overall geographic tendency for pelage pattern in either of these pigtails to approach that of the other. External size, measured as the combined length of head and body, apparently follows a continuous latitudinal cline in adult leonina and nemestrina (fig. 22), with geographic variation parallel in males and females. Judging from specimens examined, external size is relatively constant in Indochinese populations of leonina south to about 10° N in peninsular Thailand. Farther southward, length of head and body in local populations of leonina and nemestrina gradually increases, by about 100 mm. over 15 degrees of latitude, to reach a maximum in Sumatran nemestrina. It is significant that geographically isolated populations of nemestrina in Sumatra, Borneo, and the Malay Peninsula apparently all conform 86 FIELDIANA: ZOOLOGY, VOLUME 67 20«N - '.= 10° N O - 0 o ■ /eon/no o • nemestrina v » pagensis - o 0 o a A •. m -u %o §.' • ■ ■" °o • ■ ■ • n ooo > • M • • • o ••• • 9? o • • • 1 1 0 oo o 1 1 o - _1_ o • • • • • iiil. _ 0 T) O ^ ■ ■ • ■ ° ° ° o- <• • OOOOQO •* / . o % oooo O 0 •a _ o o oo o • ooo 1 1 1 1 o • •• • • 1111 400 500 600 700 Head and body length (mm) 120 140 160 Skull length (mm) Fig. 22. Latitudinal variation of head and body length and skull length in adult specimens examined of leonina, nemestrina, and pagensis; specimen locality numbers (figs. 25, 28) as follows: leonina (fig. 28), 1, 21-26, 29, 31, 32, 37, 39, 42, 44 (for measurements of one female skull, see Pocock, 1932, p. 308), 45-47, Chamchay (unnumbered locality, see "Gazetteer of Collecting Localities"); nemestrina (fig. 25), 14, 6, 9, 10, 12, 18, 24, 26, 28-30, 33, 34, 36-39, 42-52, 55, 56, 58, 64, 66, 68, 69, 76, 77, 82- 84, 87, 89, Banang, Trang (unnumbered locahties, see Gazetteer); pagensis (fig. 25), 92, 93. Small symbols represent measurements in two subadult nemestrina males (canines incompletely erupted). to the same latitudinal size cline. Over the entire range, external size is broadly overlapping in leonina and nemestrina. Cranial size, measured as greatest length of skull excluding incisors, evidently follows a step cline in leonina and nemestrina (fig. 22). The leonina curve and the nemestrina curve appear to be abruptly discontinuous about 8°N, and known overlap of greatest skull length in these two pigtails is mimimal. In adult males, skull length exceeds 142 mm. in 56 of 58 nemestrina specimens examined and in none of 15 leonina specimens; in adult females, skull length exceeds 120 mm. in 24 of 27 nemestrina specimens and in only one of 19 leonina specimens. Other aspects of cranial morphology (zygomatic breadth, rostral-postrostral proportions, relative length FOODEN: LIONTAIL & PIGTAIL MACAQUES 87 of dental fields; see above) are directly or allometrically related to skull length and therefore have a similar pattern of geographic variation. The geographic ranges of leonina and nemestrina are contiguous in peninsular Thailand (8-9° N) at the southern end of the Isthmus of Kra (fig. 23). Available from this area are 13 pigtail specimens (table 20), of which six are referred to leonina and seven to nemestrina. In five insular specimens collected at Klong Tung Sai and Telok Poh, virtually all known external and cranial characters are those of leonina; the only exception is external size in one adult male (570 mm.), which is in the range of overlap of leonina and nemestrina. A baculum of the Telok Poh specimen also is available (fig. 15d, table 6); although this is the shortest adult leonina baculum examined (greatest length 19.7 mm.), its form is clearly that of leonina (distal process length/shaft length = 0.47). One mainland specimen collected at Ban Nong Kok (skin only, subadult or adult male) likewise is a typical example of leonina in all available characters, as previously noted by Chasen (1935, p. 38, "Ghirbi" specimen). The northernmost pigtail specimen that clearly exhibits pelage characters of nemestrina is an adult female recently collected by me at Khao Si Kaeo, which is about 10 km. north of Ban Nong Kok (mainland leonina locality mentioned above). Critical characters in this specimen are a mixture of those of nemestrina and leonina, with nemestrina characters considered to predominate. The dorsal surface is dark brown as in nemestrina, distinctly darker than in the Ban Nong Kok leonina specimen and in all other leonina specimens examined. Skull length (119.9 mm.) is in the range of overlap of adult female nemestrina and leonina. The crown patch is narrow anteriorly, as in leonina, and length of head and body (434 mm.) is within the range of female leonina (less than in any other adult female nemestrina). When encountered by hunters who took the specimen, the troop to which this female belonged was 20-25 m. above the ground in dipterocarp trees; in response to gunfire, some members of the troop descended and fled on the ground, as in nemestrina, and others fled in the treetops, as in leonina. In one adult female and one adult male collected at Sathani Lam Phura, about 100 km. southeast of Khao Si Kaeo, the dorsal surface is dark and the crown patch is broad, both as in typical nemestrina. Skull length in the female (127.7 mm.) also is typical of — 13 o (0 « -2 •z. o> i^ •r uo *- ti la 01 .a 2: * 0 1- u u S3 < a a c « u o la 3 .c 3 u c u p 0 tu CQ ~ I I I I I I I - I c II I -- I I CO 01 3 Zl •- _£ - CO X - 1- 01 — 1- 0 — z B — — CO 2 Q£ — >> r c S3 ~ o> 0> d J JS ■fl v. - s B a. 0 JZ a u a a 5 0 _^ — o> c M ti i_ eg 1 a "3 i 0 ~ o S = ~ 01 CO E o> 01 "5 o '5 2 o< a-o ta o> — £ — ■- o* X ti o> c X a .. e II o> ■— c < — C E — t» t— C- t- X Oi Gi O C5 ot *0 o» "O c- *- — o c Oi cc n — *-* o r~ x- en a; CI U5 Cft U3 IJ) Vi Ui U' W "O jtt i/: 0 CJ 01 o 5 (N ~ _: < 0 Z . B II CO -_ o c 3 in E- * c m a >< 0 CQ >• - 0 © -£ - * m 3 o *r cj -* .-« * t~ Cfl CO CO 0 o o u ft« _ a: qs a. .* S z o- NNN 0 CO c N X; c w 15 o O X »- m 3 *° £ in »> en w 22£ 3 - 1- 't U3 CO 5 z •- u u Q S « os o; a fc ■£ N N lO m' c in 2 bS? 0 03 w .c 3 to x 0 f- *"- Z- to to -» in in in 0 o — •—• »-< tN 1 I . CO ZZ'MU S «> c OC tv U- 2 N FOODEN: LIONTAIL & PIGTAIL MACAQUES Fig. 23. Map of peninsular Thailand showing marginal collecting localities of Indochinese pigtails (leonina, squares) and Sundaic pigtails (nemestrina, circles). nemestrina. (Length of head and body given on the tag of this specimen is 590 mm., which is highly improbable; this may be a lapsus for 490 mm.) Skull length in the male (139.6 mm.) is at the lower limit of the nemestrina range (upper limit of leonina range) and length of head and body (532 mm.) also is at the lower limit of the nemestrina range (well within leonina range). In one subadult male (canines not quite fully erupted) collected at Chong, 25 km. southeast of Sathani Lam Phura, the crown patch is narrow anteriorly, as in leonina, and the dorsal surface is pale golden-brown agouti with a weakly defined median dark streak (about 5 cm. broad), approximately intermediate between typical leonina and nemestrina. Hairs in the side- whisker region are short and dark-tipped, as in nemestrina. Skull length in this subadult (147.7 mm.) is within the range of adult male nemestrina, and length of head and body (648 mm.) approaches 90 FIELDIANA: ZOOLOGY, VOLUME 67 that in the largest known adult male specimens of nemestrina collected in southern Sumatra. This Chong subadult appears to have been the specimen on which Kloss (1919b, p. 344) based his influential decision to regard leonina and nemestrina as conspecific. In one subadult male (upper canines about 75 per cent erupted) and one juvenile female that I collected near Huai Lian, 17 km. east of Chong, the dorsal surface is dark brown to blackish, approximate- ly as in typical nemestrina, and the crown patch is narrow, as in leonina. In the subadult male, skull length (139.6 mm.) is about equal to the maximum known in adult male leonina and to the minimum known in adult male nemestrina. Length of head and body in this subadult (548 mm.) is in the range of overlap of adult male leonina and nemestrina. In the baculum of this subadult, relative length of the distal process (0.30; fig. 15r) is slightly greater than the maximum known in adult nemestrina (0.29; table 6, NMS 2629) and slightly smaller than in a less mature subadult leonina (0.31; table 9, FMNH 99672); on balance, bacular structure in this subadult seems slightly closer to that in nemestrina than in leonina. These Huai Lian specimens were collected on successive days and belonged to different troops; in response to gunfire, both troops fled in the treetops, as in leonina. Also collected in the general area of Sathani Lam Phura, Chong, and Huai Lian, is an isolated adult male skull with locality given merely as "Trang" (province). This is a typical, large nemestrina skull (greatest length 154.3 mm.). Another male specimen of nemestrina collected in this area is briefly mentioned in passing by Kloss (1919b, p. 344); this may be the Sathani Lam Phura male discussed above. In summary, typical leonina specimens have been collected as far south as Klong Tung Sai, Telok Poh, and Ban Nong Kok. Within 10-150 km. of Ban Nong Kok, at Khao Si Kaeo, and in the Trang area (Sathani Lam Phura, Chong, Huai Lian), seven pigtails have been collected in which nemestrina characters— most conspicuously, dark dorsal pelage color— are variably mixed with leonina characters, especially the narrow crown patch. This accords well with Kloss' (1919b, p. 344) observation, based on two specimens available to him, that "Trang may therefore be regarded as the locality where intermediates between the two forms occur." About 400 km. farther south, Malayan pigtail specimens exhibit characters that are unambiguously those of typical nemestrina. FOODEN: LIONTAIL & PIGTAIL MACAQUES 91 The mixture of characters in geographically marginal speci- mens referred above to nemestrina, taken together with the continuous clinal distribution of external size in leonina and nemestrina tends to support classification of these two pigtails as conspecific subspecies, which, following Kloss (1919b, p. 344), has been the usual recent taxonomic practice. However, the narrow geographic distribution of known specimens with mixed or inter- mediate pelage and bacular characters and the step cline in cranial size probably indicate that genetic exchange between leonina and nemestrina is relatively restricted. On balance, despite some evidence of partial genetic discontinuity, it seems appropriate to follow a conservative course and continue to regard these two pigtails as subspecies of the species M. nemestrina. The geographic isolation of Kepulauan Mentawai pagensis precludes the possibility of interbreeding or intergradation with neighboring Sumatran nemestrina. Since the degree of morphologi- cal differentiation between pagensis and nemestrina is comparable to that between nemestrina and leonina, it seems reasonable to accord equivalent taxonomic status to all three of these forms. Accordingly, pagensis is also treated in this revision as a subspecies of M. nemestrina. Macaca nemestrina nemestrina (Linnaeus, 1766) Pig-tailed Monkey: Edwards, 1758, p. 8, pi. 214 (animal in color) — external characters of two captives; distribution, Sumatra. Le Maimon: Buffon and Daubenton, 1766, p. 176, pi. 19 (animal) — external characters and measurements of captive, origin unknown; internal anatomy; osteology; taxonomy; distribution, Sumatra and probably elsewhere in East Indies. [Simia] Nemestrina Linnaeus, 1766, p. 35 - external characters ex Edwards (1758); distribution, Sumatra; the specific name is an adjective (derived from Latin name of god of groves) agreeing in gender with the feminine generic name. Simia Nemestrina: Schreber, 1774, p. 79, pi. 9 (animal ex Edwards, 1758) - external characters; distribution, Sumatra. [Papio] Nemestrina: Erxleben, 1777, p. 20 - external characters; distribution, Sumatra, Japan. Papio nemestrinus: [Olgiby], 1838, pp. 363, 373 - taxonomy. Cantor, 1846, p. 176 - first record of pigtail macaque in Malay Peninsula (Pulau Pinang). Macaca nemestrina: Daudin, [1802], p. 148 - listed; new combination. Miller, 1906, p. 556, pis. 13-20 (skin, skulls) — external and cranial characters; range restricted to Sumatra. Grimm, 1967, p. 361 - vocalizations. Blakley, Morrow, and Morton, 1973, p. 120 — serum cholesterol higher in Sumatran specimens than in West Malaysian specimens. 92 FIELDIANA: ZOOLOGY, VOLUME 67 Macacus nemestrinus: Desmarest, 1820, p. 66, pi. 10 (animals ex Edwards and Buffon) — external characters; distribution, Java Sumatra. Anon., 1851, p. 444 — distribution, Sumatra, Borneo, Pinang, Malacca. Bleeker, 1851, p. 527 — first record of pigtail macaque in Pulau Bangka. Hagen, 1890, p. 82 — field observations. Sclater, 1893, p. 325 — report of aberrantly colored Bornean specimen. Thomas and Hartert, 1894, p. 654 — field observations; introduced in Kepulauan Natuna. Macaca nemestrina nemestrina: Kloss, 1919b, p. 345 — external characters; said to intergrade with Indochinese pigtail macaque at Trang, peninsular Thailand. J. R. Napier and P. H. Napier, 1967, p. 403 - listed. Macacus nemestrinus nemestrinus: Davis, 1958, p. 126 — external characters; field notes. Pithecus nemestrinus: E. Geoffroy, 1803, p. 25 — external characters of two museum specimens; distribution, "L'Inde meridionale." Cynocephalus nemestrinus: Latreille, 1804, p. 291 — external characters. Inuus nemestrinus: E. Geoffroy, 1812, p. 101 — external characters, distribution, Java, Sumatra. I{nuus] nemestrinus: Wagner, [1839], p. 143 — external characters; distribution, Sumatra, Borneo. Macacus (Innus) nemestrinus: Muller, 1839-1840, p. 14 — field observations; distribution, Sumatra and Borneo, transported from Sumatra to Java. S[almacis] nemestrina: Gloger, [1841], p. 36 — new combination. Nemestrinus nemestrinus: Schneider, 1905, p. 73 — field observations. Rhesus nemestrinus: DeBeaux, 1923, p. 36 — external and cranial characters. Pig-tail [Baboon]: Pennant, 1771, p. 105 — part (also includes references to Macaca fuscata); external characters; distribution, Sumatra, Japan. New Baboon: Pennant, 1771, p. xxiii, pi. 13 (animal) — external characters, based on figure of specimen of unknown origin. Simia platypygos Schreber, 1774, p. 74, pi. 5b (animal, head) — syntypes, male and female of unknown origin observed in captivity by D. Hermann; external characters; habits; characteristic pigtail grimace noted. Erxleben, 1777, p. 629 — external characters. Zimmermann, 1780, p. 181 — possibly a synonym of Papio nemestrina Erxleben, 1777. E. Geoffroy, 1812, p. 101 - a synonym of Simia nemestrina Linnaeus, 1766. Papio platypygos: Kerr, 1792, pp. 32d, 62 - listed. Brown Baboon: Pennant, 1781, pi. 20 (animal) - replacement name for New Baboon: Pennant, 1771; Simia platypygos Schreber, 1774, cited in synonymy. Babouin a longues jambes: Buffon, 1789, p. 41, pi. 8 (ex Pennant, 1781) - external characters and habits of captive specimen, origin unknown; Simia platypygos Schreber, 1774, cited in synonymy. S[i/rn'a] Longicruris Link, 1795, p. 62 — based on Babouin a longues jambes: Buffon, 1789. J. E. Gray, 1843, p. 7 — synonym of Macacus nemestrinus Desmarest, 1820. FOODEN: LIONTAIL & PIGTAIL MACAQUES 93 S[imia] Fusca Shaw, 1800, p. 24, pi. 13 (animal ex Pennant, 1781) - type, Brown Baboon: Pennant, 1781; Simia platypygos Schreber, 1774, cited as a possible synonym. E. Geoffroy, 1812, p. 101 — a synonym of Simia nemestrina Linnaeus, 1766. Macaca sphinx [Variete C]: Daudin, [1802], p. 149 (not Linnaeus, 1758) - applied to Babouin a longues jambes: Buffon, 1789. Macaque, Espece inedite: F. Cuvier, 1816-1817, pi. 5 (animal) — figure of tumescent female, origin unknown. Singe a queue de cochon: F. Cuvier, 1820, liv. 19, p. 1, pi. (animal) - taxonomy, referred to Simia nemestrina Linnaeus; external characters; reproductive biology, sexual swelling, gestation 7 months 20 days; distribution, Sumatra. F. Cuvier, 1822a, liv. 36, p. 1, pi. (animal, head) — taxonomy; distribution, Sumatra. Simia carpolegus Raffles, [1821], p. 243 — based on specimens collected and observed in Sumatra by T. S. Raffles between 1818 and 1820; syntypes formerly preserved in collections of Zoological Society of London (Waterhouse, 1838, p. 8) and East India Company (Horsfield, 1851, p. 19); external characters and variation; M3 5-cusped; trained to pick coconuts; Malay name, Bruh; type-locality, Bencoolen ( = Bengkulu), southwestern Sumatra. Fischer, 1829, p. 30 — a synonym of Simia nemestrina Linnaeus, 1766. Cynocephalus carpolegus: [Vigors], 1829, p. 13 — specimen listed. Macacus libidinosus I. Geoffroy, 1826, p. 589 — based on illustration of Macacque, Espece inedite: F. Cuvier, 1816-1817, origin unknown. S[imia] libidinosa: Fischer, 1829, p. 30 — doubtfully distinguishable from Simia nemestrina Linnaeus, 1766. P[ithecus] Maimon Blainville, 1839, pi. 10 (dentition of adult male) — new name used in figure caption. Macaca nemestrina var. leoninus: Oudemans, 1890, p. 266 — captive juvenile received from Sumatra. Macaca broca Miller, 1906, p. 558 — external and cranial characters; holotype, adult male, skin and skull (USNM 34930, Coll. No. 19211), collected at Sungai Sapagaya, Sabah by C. F. Adams, Nov. 21, 1887; distribution, Borneo. Lyon, 1907b, p. 566 — cranial comparison with Bornean M. nemestrina. Elliot, [1913], p. 205 — a synonym of Simia nemestrina Linnaeus, 1766; synonymy said to be concurred in by G. S. Miller, original author of broca. P[ithecus] broca: Lyon, 1911, p. 136 — cranial comparison with Bornean M. nemestrina. Macaca nemestrina broca: Myers and Kuntz, 1969, p. 420 — nematode parasite. Macaca nemestrina nucifera Sody, 1936, p. 42 — based on skin (head only) and skull of adult male collected at Pulau Bangka by H. J. V. Sody (Coll. No. Bk 75), date and museum unknown; external and cranial characters; taxonomic comparisons. Sody, 1937, p. 248 — taxonomic comparison. Chasen, 1940a, p. 65 — a synonym of Macaca nemestrina nemestrina. Type. — None preserved. Linnaeus' (1766, p. 35) description of this species is based solely on Edwards' (1758, p. 8) illustrated FIG. 24. Pig- tailed Monkey: Edwards (1758, p. 8), holotype of M. nemestrina nemestrina (Linnaeus, 1766). 94 FOODEN: LIONTAIL & PIGTAIL MACAQUES 95 account of a captive juvenile male (fig. 24) that was brought from Sumatra to England on a British warship in 1752 and was subsequently purchased by Edwards. In Edwards' account, the holotype juvenile is compared with an older captive female (paratype) that was then on exhibit at a fair in London. Neither holotype nor paratype is known to have been preserved. Type -locality. — Sumatra (Edwards, 1758, p. 8). Distribution (fig. 25). — Sunda area from Surat Thani-Krabi depression in peninsular Thailand (8-9°N) southeastward through West Malaysia, Sumatra, Bangka, and Borneo; known altitudinal range 0-1700 m. (table 11). M. n. nemestrina apparently is native to the offshore islets Pulau Pinang (west coast of West Malaysia), Pulau Tioman (east coast of West Malaysia), and Pulau Batam (Riau Archipelago, southern tip of West Malaysia) (see Gazetteer). Other offshore islets generally are excluded from the natural range of the subspecies, although it often is present as an introduced element. A captive specimen said to have been obtained in the Nicobar Islands (Corner, 1941, p. 14) probably represents such an introduction (see Kloss, 1928, p. 802). Anomalous locality information is recorded on the museum tag of one captive adult male specimen of M. n. nemestrina (AMNH 119514, skin and skeleton; blackish dorsal streak, greatest length of skull 144.8 mm.) that died in the New York Zoological Park on June 18, 1937 and was catalogued at the museum on the same day. According to the tag, this monkey was born about December, 1931 and was obtained by the Legendre Indo-China Expedition (date unspecified) at Plateau des Bolovens, Laos, which, however, is far outside the indicated range of M. n. nemestrina (fig. 25). It is known that the Legendre expedition collected a series of M. n. leonina skins (AMNH 87268 6, 87269 8, 87273?, 87274$; immatures) at Plateau des Bolovens in February 1932 and also obtained a living specimen that was donated to the New York Zoological Park later the same year (Berger, 1932, p. 329; Legendre, 1932, p. 495; New York Zoological Society Annual Report, 1933, p. 109). It seems likely that geographic information properly applicable to Legendre's captive Indochinese pigtail (whereabouts now unknown) was inadvertently transferred to a captive Sundaic specimen (AMNH 119514); this may have occurred while both were being kept at the New York Zoological Park. External characters (fig. 26). - Head and body length 434-576 mm. in 17 adult females, 532-738 mm. in 23 adult males (excluding Fig. 25. Locality records of M. n. nemestrina (circles) and M. n. pagensis (inverted triangles); closed symbols = specimens examined, open symbols = literature records; squares - marginal records of M. n. leonina; letters and heavy broken lines indicate negative reports. For details, see "Gazetteer of Collecting Localities." M. n. nemestrina localities: 1. Khao Si Kaeo. 2. Lam Phura. 3. Chong. 4. Huai Lian. 5. Biserat. 6. Pinang. 7. Larut; Perak, Upper. 8. Gedong; Keroh Forest Reserve. 9. Hantu, Tanjong; Simpang Empat Rungkup. 10. Jeram; Lima Belas Estate. 11. Sungai Buloh. 12. Kelang; Meru; Sungai Renggam. 13. Galas, Sungai. 14. Gua Madu. 15. Bukit Fraser, 5 km. S; Bukit Fraser, northern slopes. 16. Kuala Lompat: G. Benom. 17 Bentong; Bentong, 6-9 miles W. 18. Ampang; Kepang; ? Kuala Lumpur; Kuala Lumpur, Cheras Road, 7th mile; Ulu Gombak Forest Reserve. 19. Melaka. 20. Janing, Gunong. 21. Johor, southeast. 22. Tioman, Pulau. 23. Batam, Pulau. 24. Pasi. 25. Leuser Reserve; Pulau Munteh. 26. Aru, Teluk; Bukit Tinggi; Langsa; Pangkalanberandan; Pulutelang. 27. Borohok; Langkat; Serapit. 28. Batangkuwis; Deli; Dolok-Oelve; Medan; Tandjungmorawa. 29. Bedagi; Lauttador; Padang [district]; Paguruan, Sungai; Serdang; Tebingtinggi; Tandjung. 30. Sibolga, Teluk. 31. Sirambas. 32. Lubukbasung. 33. Padang. 34. Lida Ajer cave; Pajakumbuh. 35. Djamboe Cave; Sibrambang Cave. 36. Kajutanam; Solok. 37. Tarusan, Teluk. 38. Siak, Sungai. 39. Kateman, Sungai. 40. Baru, Danau. 41. Kota, Danau; Rengat. 42. Bengkulu. 43. Sanggul, Bukit. 44. Lubuklinggau; Muarabeliti. 5. Pagaralam. 46. Babat; Kluang; Loeboek Karet; Palembang [district]. 47. Palembang. 48. Ogan, Air. 49. Bangka, Pulau. 50. Kuching; Sidong, Gunong. 51. Sekajam, Sungai. 52. Landak, Sungai. 53. Kapuas, Sungai. 54. Pontianak. 55. Sukadana. 56. Riam. 57. Kendawangan, Sungai. 58. Pulo Paku. 59. Entawa-Samarahan. 60. Roema Manoeal. 61. Semitau. 62. Sintang. 63. Niah Caves. 64. Salai, Sungai. 65. Kalulong, Bukit. 66. Sut. 67. Puruktjahu. 68. Tandjung. 69. Pamukan, Teluk. 70. Klumpeng, Teluk. 71. Pelaihari. 72. Papar. 73. Labuan, Pulau, mainland opposite. 74. Madihit, Sungai. 75 96 FOODEN: LIONTAIL & PIGTAIL MACAQUES 97 improbable measurements: adult female, 590 mm., ZRSC 1214/10, Loc. No. 2; adult male, 469 mm., ZRSC 2592/10, Loc. No. 10) (fig. 22) relative tail length (T/HB) 0.30-0.44 in 16 adult females (0.16 in one aberrant adult female, AMNH-AC 106562, Loc. No. 43, largest female specimen examined), 0.28-0.42 in 22 adult males (fig. 2); weight 5.4-7.6 kg. in 10 adult females, 10.0-13.6 kg. in 8 adult males (table 21). Pelage of trunk yellowish-brown agouti to golden-brown agouti laterally, with a variably developed blackish mid-dorsal streak or patch (about 4-20 cm. broad) that becomes more or less indistinct anteriorly on the scapular region; crown hairs short (1 cm.), blackish, radiating to form a whorl centered at the vertex; crown patch broad anteriorly, extending laterally on supraorbital region approximately as far as lateral angle of each eye; side whiskers relatively short (2-3 cm.), with buffy bases and blackish tips that collectively form a pair of dark preauricular streaks or sideburns; skin of muzzle pale brownish, thinly covered with inconspicuous short whitish hairs; outer surface of arms, legs, hands and feet yellowish-brown to golden-brown, approximately like flanks; posterior surface of shanks frequently with indistinct dark streaks; tail thin, bicolor, sharply defined blackish dorsally, pale ochraeceous-buff ventrally, frequently with terminal hairs elongate and weakly defined ochraceous to dark reddish-brown, forming an inconspicuous tuft; underparts thinly haired whitish to ochraceous- buff anteriorly, often becoming darker (buffy to pale brown) on the epigastric region. Fig. 25. continued Bareo; Pa Umor. 76. Bahau, Sungai. 77. Badang; Peleben. 78. Karangan, Sungai. 79. Kin.il Reserve. 80. Tabang. 81. Djembajan, Sungai. 82. Lumu Lumu; Ranau. 83. Betotan; Lungmanis Station. 84. Sapagaya, Sungai. 85. Ulu Segama Forest Reserve. 86. Tibas, Sungai. 87. Sandakan. 88. Kinabatangan, Sungai. 89. Abai. 90. Kretam Kechil, Sungai; Segama, Sungai. M. pagensis localities: 91. Sioban. 92. Pagai Utara, Pulau. 93. Pagai Selatan, Pulau. Negative reports: C. Nicobar Islands, northern. D. Nicobar Islands, southern. E. Klet Kaeo, Ko (fig. 28). F. Simeulue, Pulau. G. Banjak, Kepulauan. H. Bangkaru, Pulau. I. Nias, Pulau. J. Musala, Pulau. K. Enggano, Pulau. L. Sebesi, Pulau. M. Panaitan, Pulau. N. Krakatau, Kepulauan. O. Pangkor, Pulau. P. Berhala, Pulau. Q. Jarak, Pulau. R Sembilan, Kepulauan. S. Perhentian Besar, Pulau. T. Redang, Pulau. U. Bidong Laut, Pulau. V. Kundur, Pulau. W. Karimun, Pulau. X. Tinggi, Pulau. Y. Durian, Pulau. Z. Sugi, Pulau. a. Bulan, Pulau. b. Bintan, Pulau. c. Sebangka, Pulau. d. Bakung, Pulau. e. Lingga, Pulau. f. Singkep, Pulau. g. Natuna, Kepulauan. h. Lemukutan, Pulau. i. Temadju, Pulau. j. Datuk, Pulau. k. Masalembo-Besar. 1. Laut, Pulau. m. Sebuku. FlG. 2& Mid in a nemestrina nemestrina, external characters. (Photo courtesy Japan Monkey Centre) 98 Females Males 5.45 — — 10.9, 11.8 7.3, 7.6 — 7.3 13.6 6.4 10.9, 11.0 10.9, 10.9 5.4, 5.7, 6.4, 6.4, 6.8 10.0 6.5 11.2 FOODEN: LIONTAIL & PIGTAIL MACAQUES 99 TABLE 21. Weight in wild collected adult specimens of M. n. nemestrina. Locality Weight (kg.) Nos.l / 26 30 37 38. 39 55, 69 89 Means 1- See map, Figure 25. In adult males dorsal hairs are longest (7-9 cm. long) on the scapular region, where the blackish median streak is least distinct. The basal quarter of each of these scapular hairs is pale buffy, and the distal three-quarters is annulated with about ten alternating bands of yellowish or golden and blackish, with the blackish bands generally broader than the yellowish bands. More posteriorly, hairs in the blackish median streak are 3-4 cm. long, with the basal half of each hair pale buffy and the distal half blackish, rarely with an indistinct narrow pale annulation. In three subadult or adult males collected at localities less than 150 km. apart in west-central West Malaysia, width of the blackish mid-dorsal streak varies from about 8 cm. (BM 55.1502, subadult, Bentong) to 12 cm. (BM 55.1503, adult, Sungai Renggam) to 18 cm. (BM 50.1501, adult, Rungkup). Anterior extent of the dorsal streak is roughly correlated with its width. In the West Malaysian specimen with the narrowest dorsal streak (BM 55.1502), the streak is separated from the blackish patch on the crown and nape by a clearly defined zone of golden-brown on the scapular region; in the specimen with the broadest dorsal streak (BM 55.1501), a virtually uninterrupted blackish band extends from the crown to the dorsal surface of the tail, with only a slight tinge of brownish on the scapular region. The darkest of these West Malaysian specimens (BM 55.1501) is closely matched by an adult specimen collected in southeastern Sumatra (BM 1938.11.30.6, Palembang), near the southern limit of the range of the subspecies, and by another collected in northeastern Sarawak (BM 93.6.2.2, Bukit Kalulong), near the northeastern limit of the range. In a very old male collected in Sabah (USNM 34930, Sungai Sapagaya, type of broca 100 FIELDIANA. ZOOLOGY, VOLUME 67 Miller), the dorsal ground color is drab-yellowish-brown, with unusually indistinct annulations on individual hairs. In adult females and in immatures of both sexes the dark dorsal streak is generally somewhat more restricted and dilute than in adult males, the hair on the flanks is paler and drabber than in adult males, and the agouti pattern is less conspicuous. In one adult female (BM 93.4.29.1, Sungai Baram, Sarawak) that reportedly lived in captivity in Sarawak for about five years (Sclater, 1893, p. 325), the dorsal fur, which is unusually fine and silky, is pale golden- brown non-agouti laterally, with a narrow and indistinct darker brown streak medially. Pocock (1931, p. 310) has speculated that this exceptionally pale specimen is a captive-bred M. mulata-M. n. nemestrina hybrid; alternatively, it may merely be a albinistic specimen of M. n. nemestrina. Two albinistic Bornean specimens have been reported by Banks (1931, p. 111). As previously noted (fig. 22), length of head and body generally increases from north to south in M. n. nemestrina, with the same clinal pattern of increase evident in all three isolated components of the range (Malay Peninsula, Sumatra, Borneo). However, one exceptionally large subadult male (BM 55.1504) has been collected at Chong, peninsular Thailand, near the northern limit of the range (fig. 22). Relative tail length tends to be somewhat less in the largest specimens examined than in smaller ones (fig. 2). Cranial characters (figs. 4, 6, 27). - Skull relatively large, greatest length excluding incisors 115.4-138.9 mm. in 35 adult females, 139.5-174.5 mm. in 77 adult males (fig. 7); rostrum long (fig. 8); zygomatic arches relatively narrow (fig. 7); supraorbital ridges relatively thin, especially laterally, and somewhat receding; anterior surface of malar flat to weakly convex. Skull length in specimens collected in the northern part of the range of M. n. nemestrina (Malay Peninsula; Sumatra and Borneo north of 3°N) averages about 10 mm. less than in those collected in the southern part of the range (Bangka; Sumatra and Borneo south of 3°N) (fig. 22). However, one large specimen (NMS 16640, skull only, greatest length 170.7 mm.) reportedly collected at "Dolok- Oelve," Deli, northern Sumatra, appears to be an exception to this generalization (fig. 22). Habits and habitats. — This account is based primarily on field observations published by authors listed below; authors are cited by means of indicated abbreviations. Fig. 27. Cranial variation in adult male M. n. nemestrina: a, AMNH-AC 106037, Badang; b, MZB 6501, Pulau Bangka (possible captive): c, AMNH-AC 106563, Bukit Sanggul. xtt (Photos FMNH) 101 102 FIELDIANA: ZOOLOGY, VOLUME 67 A Abbott in Lyon, 1907b, p. 566 Ba Banks, 1931, p. 110 Be-a Bernstein, 1967a, pp. 198-207 Be-b Bernstein, 1967b, pp. 217-228 C Chivers, 1971, pp. 78, 84 D Davis, 1962, p. 58 Hg Hagen, 1890, p. 82 Hrt Hartert, 1902, p. 193 Hrt/T Hartert in Thomas and Hartert, 1894, pp. 654-655 Hs Harrison, 1969, p. 175 Hss/D Harrisson in Davis, 1958, p. 126 Hss/M Harrisson in Medway, 1965, p. 85 K Kawabe, 1970, p. 286 K/M Kawabe and Mano, 1972, p. 216 Ku Kurt, 1973, pp. 64-65 L Lim Boo Liat, 1969, p. 127 McC McClure, [1965], p. 57 Me Medway, 1970, p. 527 Me/W Medway and Wells, 1971, p. 247 Mii Muller, 1839-1840, pp. 17-18 R Rodman, 1973, pp. 655-659 Sch Schneider, 1905, p. 73 So/C Southwick and Cadigan, 1972, p. 8 St Stott, 1964, p. 12 These published field observations have been supplemented by brief unpublished notes recorded on field tags of four museum specimens collected in Sumatra by M. Boogaarts in 1929 (SMF 16640, ad 6\ "Dolok-Oelve") and J. J. Menden in 1933 (AMNH-AC 102203, ad $, Lubuklinggau; AMNH-AC 102201-02, juv. W Fig. 30. Type specimen of Macaca nemestrina pagensis Miller (1903b) (center; USNM 121653, adult female) compared with adult female specimens of M. ru leonina (above) and M. n. nemestrina (below). (Photo by J. Fooden) brown distally; dorsal surface of basal 10-20 per cent of tail covered with dark brown fur, balance of tail virtually naked, thinly covered with inconspicuous short buffy hairs; underparts thinly haired, buffy to pale brown. The non-agouti dorsal pelage in M. n. pagensis is strikingly different from the agouti pelage in M. n. leonina and M. n. nemestrina, particularly in adult males (figs. 30, 31). Crown hairs in M. n. pagensis are relatively long (3-4 cm.) and radiate to form a broad whorl centered at the vertex; this cap of radiating hairs extends notably farther anteriorly and laterally than in M. n. leonina and M. n. nemestrina. The large pale patches on the sides of the neck in M. n. pagensis constrict the dark area on the nape to a relatively narrow band (about 5 cm. broad) which joins the broader dark area on the crown to that on the dorsal surface of the trunk. The nearly hairless condition of the distal 80-90 per cent of the tail, which superficially resembles that of an opossum, is consistent in all eight specimens examined and therefore apparently is normal in the subspecies. The color of the dorsal surface of the hands and feet is about the same as that of the forearms and FOODEN: LIONTAIL & PIGTAIL MACAQUES 121 Fig. 31. Dorsal pelage in adult male M. n. pagensis (above; AMNH-AC 103394, Pulau Pagai Utara) compared with that in adult male M. n. nemestrina (below; FMNH 33643, Sumatra: Air Ogan); in M. n. pagensis note broad crown whorl, pale lateral throat patches, dark non-agouti dorsum, contrastingly colored (reddish-brown) forearms, and thinly haired tail. (Photo FMNH) shanks. Miller (1903b, p. 62) indicates that the hands and feet are contrastingly darker than the arms and legs, but this seems to be based on misinterpretation of grease stains on the hands and feet of the type-specimen (USNM 121653). Cranial characters (figs. 4, 6). — Skull relatively small, greatest length excluding incisors 109.1-112.1 mm. in 3 adult females, 134.2 mm. in 1 adult male (fig. 22); rostrum short (fig. 8); zygomatic arches narrow (fig. 7); supraorbital ridges thick, somewhat receding; anterior surface of malar weakly convex. Habits and habitats. — The only information available concerning behavior or ecology in M. n. pagensis is that three specimens collected at Pulau Pagai Utara and three collected at Pulau Pagai Selatan were taken in primary forest at sea level (J. J. Menden, notes on field tags of AMNH-AC 103394-103399). Specimens examined. — Total 8. INDONESIA, Sumatra: Pulau Pagai Selatan, 3 (AMNH-AC), 1 (type of pagensis, USNM); Pulau Pagai Utara, 3 (AMNH-AC). Imprecise locality: l.(MZB). GAZETTEER OF COLLECTING LOCALITIES Locality names listed as primary entries in this gazetteer generally are the standard names approved in U. S. Board on Geographic Names gazetteers (No. 10: Malaysia, Singapore, and Brunei, 1970; No. 13: Indonesia and Portugese Timor, 1968; No. 22: Mainland China, 1968; No. 58: South Vietnam, 1971; No. 73: Laos, 1973; No. 96: Burma, 1966; No. 97: Thailand, 1966). For localities in India, The Imperial Gazetteer of India (1907-1909) is used as the standard reference. In primary entries for localities not included in U. S. Board on Geographic Names gazetteers or in the Imperial Gazetteer of India, spellings given below follow those in the original source. Secondary entries, with cross references to corresponding primary entries, provide a key to variant spellings and alternate names that have been applied to liontail and pigtail localities. In alphabetizing primary and secondary entries, generic geographic terms have been treated as in U. S. Board on Geographic Names gazetteers; these gazetteers provide comprehensive glossaries to local generic geographic terms. Primary entries include the following information: locality name; altitude, if known; country; state or other administrative division; coordinates (taken from standard references listed above, unless otherwise indicated); name of collector or observer; date of collection or observation; abbreviated name of museum in which specimens examined are preserved, or bibliographic reference to literature records. If field notes have been published concerning a locality, this is indicated by a bibliographic citation following the name of the collector or observer. Numbers in italics at the end of primary entries are locality numbers used in distribution maps (figs. 20, 25, 28); letters in italics at the end of primary entries indicate negative reports as shown in distribution maps. 122 FOODEN: LIONTAIL & PIGTAIL MACAQUES 123 Macaca silenus Map, Figure 20 Anaimalai Hills, 800-1300 m.; INDIA: Kerala and Tamil Nadu; 10°15'-10°31'N, 76°51'-77°20'E; observed by A. F. Hutton (1949, p. 690), date unknown, and Y. Sugiyama (1968, p. 284), Sept., 1961-Feb., 1962. 12 Anaimalai Hills, northern area, 1500-3000 ft., Cochin district; INDIA: Kerala; ca. 10°30'N, 77°00'E; observed by C. H. Stonor (1944, p. 591), Feb. 12-March 10, 1944. 11 Anshi Ghat, 1000 ft.. North Kanara district between Kadra (14°55'N, 74°20'E) and Kumbharwada (15°07'N, 74°24'E); INDIA: Mysore; ca. 15°00'N, 74°20'E; observed by R. S. Dharmakumarsinhji, Dec. 20, 1955 (Ali and Santapau, 1956, p. 687). 1 Cardamon Range. See Panniar. Cochin district; INDIA: Kerala; 9°48'-10°49'N, 76°00'-76°55'E; collected by F. Colyer, 1937 (BM). 9 Coorg district, western border; INDIA: Kerala; ca. 12°30'N, 75°30'E; reported by G. C. Shortridge (in Ryley, 1913, p. 490). 4 Cotengady Estate, 3500 ft., Nelliampathy Pleateau, 24 miles from Palghat (Lindsay, 1926, p. 592); INDIA: Kerala; ca. 10°30'N, 76°45'E; collected by F. R. O'Brien, April 28, 1921 (BM). 10 Cotiaddy Pass. See Kuttyadi Pass. Courtallam. See Kuttalam. Dohnavur. See Naraikkadu Estate. Goa district, near; INDIA: 15°-16°N, 74°E; reported by Rev. H. Baker, Jr. (in Blyth, 1859, p. 283). 1 Grass Hills; INDIA: Tamil Nadu; 10°21'N, 77°03'E; reported by E. R. C. Davidar (1971, p. 353), April 3-8, 1971. 12 High Range. See Panniar. High Wavy Mountain, 4000-5100 ft.: INDIA: Tamil Nadu; ca. 9°40'N, 77°25'E; observed by A. F. Hutton (1949, p. 690). 14 Kadra. See Anshi Ghat. Kalakkadu Hills; INDIA: Tamil Nadu; ca. 8°25'-8°32'N, 77°30'E; reported by J. C. Daniel (1970, p. 541). 17 Kellengode. See Seetagundy Estate. Kollangod. See Seetagundy Estate. Kumbharwada. See Anshi Ghat. Kuriarkutti. See Parambikulam Valley. Kuttalam, near; INDIA: Tamil Nadu; ca. 8°56'N, 77°16'E; observed by M. Krishnan ([1972], p. 542), about 1935; species subsequently exterminated at this locality. 16 Kuttyadi Pass, top of, between Malabar and Wynaad; INDIA: Kerala; ca. ll'WN, 76°(KyE; observed by T. C. Jerdon (1867, p. 10). 5 124 FIELDIANA: ZOOLOGY, VOLUME 67 Madura. See High Wavy Mountain. Malabar Coast?; INDIA: Kerala; 10°15'-12°18'N, 75°00,-76°00'E; J. McClelland, 1848 (ZSI). Malabar [district]; INDIA: Kerala; 10°15'-12°18'N, 75°11'-76°51'E; collector unknown, 1887 (NMS); collector and date unknown (RMNH); reported by Pere Vincent Marie (in Buffon and Daubenton, 1766, p. 171). Manjolai Tea Estate, ca. 1000 m.; INDIA: Tamil Nadu; ca. 8°38'N, 77°25'E; observed by J. R. Karr (1973, p. 191), March 2-3, 1972. 17 Naraikkadu Estate, vicinity, 2500-5000 ft.; INDIA: Tamil Nadu; ca. 8°30'N, 77°30'E; observed by C. G. Webb-Peploe (1947, p. 629). 17 Nelliampathy Plateau. See Cotengady Estate. Nelliampathy Hills, 3500 ft.; INDIA: Kerala; 10°26'-10°42'N, 76°31'-76°52'E; collected by A. P. Kinloch, March 19, 1923 (AMNH). 10 Nilgiri Hills, 800-1300 m; INDIA: Kerala and Tamil Nadu; llo12'-ll°40'N, 76°14'- 77°0O'E; observed by Y. Sugiyama (1968, p. 284) Sept., 1961-Feb.. 1962. 7 Nilgiri Hills, western slopes, 3000-4000 ft.; INDIA: Tamil Nadu; ca. 11°25'N, 76°30'E; observed by F. E. Poirier (1970, pp. 258, 260), Sept., 1965-Aug., 1966. 7 North Kanara [district]; INDIA: Mysore; ca. 14°45'N, 74°30'E; collected by W. F. Jardine, March 7, 1906 (BNHS). 2 Palghat. See Cotengady Estate. Palagapandy; Nelliampathy Hills; INDIA: Kerala; ca. 10°35'N, 76°45'E; collected by A. M. Kinloch, March 25, 1920 (BNHS; also see Anon., 1921, p. 411). 10 Panniar, 1000-1250 m., High Range, Cardamon Hills; INDIA: Kerala; ca. 10°00'N, 77°00'E; observed by Y. Sugiyama (1968, p. 284), Jan. 5-Feb. 27, 1963. 13 Parambikulam Valley, 1600 ft., near Kuriarkutti; INDIA: Kerala; 10°25'N, 76°43'E; collected by C. H. Biddulph, catalogued 1953 (BM). 8 Periyar Lake, northern shore; INDIA: Kerala; 9°36'N, 77°11'E; reported by J. Tanaka (1965, p. Ill), Jan. 7-Feb. 5, 1963. 15 Periyar Lake, south of, 800-1300 m.; INDIA: Kerala; ca. 9°25'N, 77°20'E; observed by Y. Sugiyama (1968, p. 284), Sept., 1961-Feb., 1962. 15 Seetagundy Estate, Kollangode district, Nelliampathy Plateau (Lindsay, 1926, p. 592); INDIA: Kerala; ca. 10°35'N, 76°45'E; collected by A. M. Kinloch. Nov. 29, 1920 (BNHS; also see Spence. 1921, p. 971). 10 South Kanara [district], northern border; INDIA: Mysore; ca. 13°55'N, 74°40'E; reported by G. C. Shortridge (in Ryley, 1913, p. 490). 3 Tinnevelly. See Naraikkadu Estate. Travancore district; INDIA: Kerala; 8°04'-10°21'N, 76°14'-77°37'E; collected by F. W. Muu nli 1 Ion. date and museum unknown (Blanford, 1888b, p. 17). (not mapped) Vaigai River. See High Wavy Mountain. Varagaliyar, Anaimalais; INDIA: Kerala or Tamil Nadu; not precisely located, 10°15'-10°31'N, 76°51'-77020'E; observed by M. Krishnan ([1972], p. 542), April 30. 1960. 12 FOODEN: LIONTAIL & PIGTAIL MACAQUES 125 Varushnaad Valley, south of, 3000-6000 ft.; INDIA: Tamil Nadu; ca. 9°35'N, 77°30'E; observed by A. F. Hutton (1949, p. 690). 14 Wynaad district; INDIA: Kerala; 11°27'-H058'N, 75°47'-76°27'E; reported by A. F. Hutton (1949, p. 690). 6 Macaca nemestrina nemestrina Map, Figure 25 Abai; EAST MALAYSIA: Sabah; 5°42'N, 118°23'E (Coolidge, 1940, pp. 123, 129); collected by S. L. Washburn and A. H. Schultz, June 27-July 26, 1937 (MCZ). 89 Ampang; WEST MALAYSIA: Selangor; 3°09'N, 101°46'E; observed by I. S. Bernstein (1967b, p. 220), May 4-14, 1965. 18 Aru, Teluk; INDONESIA: Sumatra; 4°09'N, 98°12'E; collected by W. L. Abbott (in Lyon, 1908, p. 620), Nov. 29-Dec. 11, 1905 (USNM). 26 Atjeh. See Langsa, Atjeh. Babat; INDONESIA: Sumatra; 2°46'S, 104°06'E; collected by Soekarno, Oct. 17- Nov. 2, 1933 (MZB). 46 Badang; INDONESIA: Kalimantan; not precisely located; ca. 3°N, 117°E; collected by V. von Plessen, May 12-25, 1935 (AMNH-AC). 77 Bagan Datoh. See Simpang Empat Rungkup. Bahau, Sungai; INDONESIA: Kalimantan; 2°50'N, 115°57'E; collected by P. Pfeffer (letter June 22, 1972), May 21, 1957 (MNHN). 76 Bakong. See Bakung, Pulau. Bakung, Pulau; INDONESIA: Sumatra; 0°04'N, 104°27'E; M. n. nemestrina absent (Dammerman, 1926, p. 316). d Banang, Sungai; INDONESIA: Kalimantan; not precisely located, on west coast between 2°N and 3°S; collected by C. Bruegel, Feb. 27, 1909 (ZSBS). (not mapped) Bangka, Pulau; INDONESIA: Sumatra; 2°S, 106°E; reported by Bleeker (1851, p. 527); collected by H. J. V. Sody (1936, p. 45), date and museum unknown; collector and date unknown (MZB). 49 Bangka, Pulau, north coast or west coast; INDONESIA: Sumatra; 1°30'-2°40'S, 105°20'-106°00'E; observed by W. L. Abbott (in Lyon, 1906, p. 611), May 20-July 4, 1904.49 Bangkaru, Pulau; INDONESIA: Sumatra; 2°04'N, 97°07'E; pigtail macaques reportedly absent (Abbott in Miller, 1903a, p. 480). H Banjak, Kepulauan; INDONESIA: Sumatra; 2°10'N, 97°15'E pigtail macaques absent (Kloss, [1928], p. 802). G Banka. See Bangka, Pulau. Baram River. See Salai, Sungai. Bareo, 3700 ft.; EAST MALAYSIA: Sarawak; 3°45'N, 115°27'E: collected by T. Harrisson, Nov. 28, 1947 (FMNH). 75 Bario. See Bareo. 126 FIELDIANA: ZOOLOGY, VOLUME 67 Barito River. See Puruktjahu. Baru, Danau; INDONESIA: Sumatra; 0°22'S, 102°23'E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown). 40 Batam, Pulau; INDONESIA: Sumatra; 1°05'N, 104°03'E; reported by C. B. Kloss i in Lyon, 1907a, p. 657), based on information supplied by local residents. 23 Batangkuwis; INDONESIA: Sumatra; 3°36'N, 98°47'E (Atlas Trop. Ned, pi. 12a); collected by Widnmann, April, 1907-Aug., 1908 (ZSBS). 28 Bedagi [district]; INDONESIA: Sumatra; ca. 3°20'N, 99°05'E; collected by G. Schneider ( 1905, p. 73), 1897-1899 (museum unknown). 29 Bedung, Pulau. See Bidong Laut, Pulau. Bencoolen. See Bengkulu. Benkoelen. See Bengkulu. Bengkulu, vicinity; INDONESIA: Sumatra; ca. 3°48'S, 102°16'E; observed by T. S. Raffles (1821, p. 243), 1817-1820; collected by E. Dubois, date unknown (RMNH), and H. J. V. Sody, date unknown (RMNH). 42 Benom, Gunong, northeast slope, 2400 ft. and 3600 ft.; WEST MALAYSIA: Pahang; 3°51'N, 102°10'E; reported by Lord Medway (1972, p. 120), 1967-1968. 16 Bentong; WEST MALAYSIA: Pahang; 3°22'N, 101°55'E; collector unknown, Dec. 16, 1912 (BM). 17 Bentong, 6-9 miles W; WEST MALAYSIA: Pahang; 3°32'N, 101°50'E; collected by C. B. Kloss (1911a, p. 146), June, 1910 (museum unknown). 17 Berhala, Pulau; INDONESIA: Sumatra; 3°46'N, 99°31'E; M. n. nemestrina absent (Harrison and Hendrickson, 1963, p. 548). P Bernam, Sungei. See Lima Belas Estate. Betong. See Pulo Paku. Betotan; EAST MALAYSIA: Sabah; 5°47'N, 117°52'E; collected by F. N. Chasen and C. B. Kloss (1931, p. 50), Aug. 14-15, 1927 (ZRCS). 83 Bettotan. See Betotan. Bidong Laut, Pulau; WEST MALAYSIA: Terengganu; 5°37'N, 103°04'E; M. n. nemestrina introduced (Kloss, 1911b, p. 176). U Bintan, Pulau; INDONESIA: Sumatra, 1°05'N, 104°30'E; M. n. nemestrina absent (Dammerman, 1926, p. 316). 6 Biserat, vicinity; THAILAND: Yala; ca. 6°35'N, 101°20'E; observed by H. C. Robinson {in Bonhote, 1903, p. 3), 1901-1902. 5 Bohorok, vicinity; INDONESIA: Sumatra; 3°30'N, 98°12'E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown). 27 Brandan. See Pangkalanberandan. Btg. Kwis. See Batangkuwis. Bukit Cheraka Klang. See Jeram. Bukit Fraser. 5 km. S; WEST MALAYSIA: Selangor; 3041'N, 101°45'E; observed by M. Kawabe (1970, p. 286), Aug., 1966-March, 1967. 15 FOODEN: LIONTAIL & PIGTAIL MACAQUES 127 Bukit Fraser, northern slopes; WEST MALAYSIA: Pahang; ca. 3°45'N, 101°45'E; observed by D. Chivers (1971, p. 80; 1973, p. 130) 1968-1970. 15 Bukit Tinggi; INDONESIA: Sumatra; 3°57'N, 98°18'E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown). 26 Bulan, Pulau; INDONESIA: Sumatra; 0°58'N, 103°55'E; M. n. nemestrina absent (Dammerman, 1926, p. 316). a Bulu Telang. See Pulutelang. Charas Road. See Kuala Lumpur, Cheras Road, 7th mile. Cheras Road. See Kuala Lumpur, Cheras Road, 7th mile. Chong; THAILAND: Trang; 7°33'N, 99°47'E; collected by H. C. Robinson and C. B. Kloss (1910, p. 669), Dec. 14, 1909 (BM). 3 Datu, Pulo. See Datuk, Pulau. Datuk, Pulau; INDONESIA: Kalimantan; 0°10'N, 108°32'E; monkeys absent (Abbott in Lyon, 1911, p. 59). j Deli district; INDONESIA: Sumatra; ca. 3°30'N,98°3O'E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown); Geryun, 1904 (ZSBS); collector unknown, 1952 (RMNH); observed by E. Hartert, date unknown (Thomas and Hartert, 1894, p. 655). 28 Dewhurst Bay. See Kretam Kechil, Sungai. Dindings. See Hantu, Tanjong. Djambajan, Sungai. See Djembajan, Sungai. Djamboe cave, near Tapisello, Padang highlands (Hooijer, 1946, p. 17); INDONESIA: Sumatra; ca. 0°25'S, 100°30'E; subfossils, prehistoric Holocene; collected by E. Dubois, 1888-1890 (presumably in RMNH; Hooijer, 1962a, p. 58). 35 Djembajan, Sungai; INDONESIA: Kalimantan; ca. 0°35'S, 116°45'E (Deignan, 1959, p. 269); collected by H. C. Raven, May 13, 1914 (USNM). 81 "Dolok-Oelve, Deli"; INDONESIA: Sumatra; not precisely located, ca. 3°30'N, 98°30'E; collected by M. Boogaarts, March 3, 1929 (NMS). 28 Durian, Pulau; INDONESIA: Sumatra; 0°42'N, 103°43'E; M. n. nemestrina absent (Dammerman, 1926, pp. 285, 302). Y East Perhentian Island. See Perhentian Besar, Pulau. Engano Id. See Enggano, Pulau. Enggano, Pulau; INDONESIA: Sumatra; 5°24'S, 102°16'E; pigtail macaques absent (Kloss, [1928], p. 802). K Entawa-Samarahan; EAST MALAYSIA: Sarawak; ca. 1°15'N, 111°00'E; collected by H. C. Robinson, Nov. 23, 1919 (BM). 59 Etawa. See Entawa-Samarahan. Fraser's Hill. See Bukit Fraser. Galas, Sungai, west of; WEST MALAYSIA: Kelantan; ca. 5°15'N, 102°00'E; reported by D. Chivers (1971, p. 80). 13 Gasip, Sungai. See Siak, Sungai. 128 FIELDIANA: ZOOLOGY, VOLUME 67 Gedong; WEST MALAYSIA: Perak; 4°07'N, 101°18'E (Annandale and Robinson, 1903, map); observed by N. Annandale (in Bonhote, 1903, p. 3). 8 Geram. See Jeram. Great Redang Group. See Redang, Pulau. Gua Madu; WEST MALAYSIA: Kelantan; ca. 4°50'N, 101°57'E; remains of monkeys eaten by humans, ca. 8000-2000 B. C; collected by M. W. F. Tweedie (1940, p. 7), July-Aug, 1939 (probably in National Museum, Singapore). 14 Hantu, Tanjong; WEST MALAYSIA: Perak; 4°19'N, 100°34'E; collected by Langgi, Aug. 2, 1918 (ZRCS). 9 Huai Lian; THAILAND: Phatthalung; ca. 7°30'N, lOOWE; collected by J. Fooden, June 28-29, 1973 (FMNH). 4 Jalan Kebun. See Kelang, Jalan Kebun, 7th mile. Janeng, Gunong. See Janing, Gunong. Janing, Gunong; WEST MALAYSIA: Johor; 2°31'N, 103°25'E; observed by H. J. Kelsall (1894, pp. 4, 16), Oct. 20, 1892. 20 Jarak, Pulau; WEST MALAYSIA: Perak; 3°59'N, 100°06'E; M. n. nemestrina absent (Harrison and Hendrickson, 1963, p. 548). Q Jeram; WEST MALAYSIA: Selangor; 3°13'N, 101°19'E; collector unknown, Nov. 16, 1910 (ZRCS). 10 Johor, southwest; WEST MALAYSIA: Johor; ca. l°3(m, 103°3O'E; reported by D. Chivers (1971, p. 80). 21 Ka jan. Sungai. See Peleben. Kajutanam; INDONESIA: Sumatra; 0°33'S, 100°20'E; collected by P. Ouruans, 1911 (MZB). 36 Kalabakan. See Tibas, Sungai. Kalulong, Bukit; EAST MALAYSIA: Sarawak; 3°14'N, 114°39'E; collected by C. Hose, Feb. 1893 (BM). 65 Kampong Durian bei Medan. See Medan. Kapuas, Sungai; INDONESIA: Kalimantan; ca. 0°, 110°E; collected by L. A. C. M. Schwaner, May, 1845 (RMNH). 53 Karagan, Sungai. See Karangan, Sungai. Karangan. Sungai; INDONESIA: Kalimantan; ca. 1°19'N, 117°42'E (Deignan, 1959, p. 269); collected by H. C. Raven, Dec. 2, 1913 (USNM). 78 Karimon. See Karimun, Pulau. Karimun. Pulau; INDONESIA: Sumatra; 1°03'N, 103°22'E; M. n. nemestrina absent (Dammerman, 1926, p. 316). W Kateman, Sungai; INDONESIA: Sumatra: ca. 0°12'N, 103°20'E; collected by W. L. Abbott, Aug. 15-27, 1903, Dec. 11. 1905 (USNM). 39 Kelabit Plateau. See Bareo and Pa Umor. FOODEN: LIONTAIL & PIGTAIL MACAQUES 129 Kelang, Jalan Kebun, 7th mile; WEST MALAYSIA: Selangor; ca. 3°02'N, 101°27'E; collector unknown, July 22, 1935 (ZRCS). 12 Kendawangan, Sungai; INDONESIA: Kalimantan; ca. 2°15'S, 110°30'E; collected by W. L. Abbott (in Lyon, 1911, p. 54), June 17-Sept. 29, 1908. 57 Kenepai, Gunung. See Roema Manoeal. Kepong; WEST MALAYSIA: Selangor; 3°13'N, 101°38'E; collected by A. L. Butler, date unknown (ZRCS). 18 Keroh Forest Reserve; WEST MALAYSIA: Perak; 4°13'N, 101°07'E; collected by R. C. Morris (1936, p. 443), June 1-12, 1935 (museum unknown). 8 Khao Si Kaeo; THAILAND: Krabi; ca. 8°12'N, 98°52'E; collected by J. Fooden, June 6, 1973 (FMNH). / Kinabalu, Mount. See Lumu Lumu. Kinabatangan district. See Kretam Kechil, Sungai. Kinabatangan R[iver]. See Abai. Kinabatangan, Sungai, 0-245 m; EAST MALAYSIA: Sabah; ca. 5°30'N, 118°0O'E; observed by M. Kawabe, Oct., 1968-March, 1969 (Kawabe and Mano, 1972, p. 216). 88 Klang. See Kelang, Jalan Kebun, 7th mile. Kluang; INDONESIA: Sumatra; 2°41'S, 103°54'E; collected by Soekarno, Aug. 21- 29, 1933 (MZB). 46 Klumpeng, Teluk, NW; INDONESIA: Kalimantan; ca. 2°45'S, 116°00'E; observed by W. L. Abbott (in Lyon, 1911, p. 58), Jan. 8-March 13, 1908. 70 Kota, Danau; INDONESIA: Sumatra; 0°23'S, 102°25'E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown). 41 Kotawaringin. See Riam Krakatau, Kepulauan; INDONESIA: Sumatra; 6°07'S, 105°25'E; primates absent (Dammerman, 1922, p. 65). N Kretam Kechil, Sungai; EAST MALAYSIA: Sabah; 5°31'N, 118°33'E; collected by D. D. Davis (1962, p. 58), June 3, 1950 (FMNH). 90 Kroh Reserve. See Keroh Forest Reserve. Kuala Lompat Post, vicinity; WEST MALAYSIA: Pahang; ca. 3°4(yN, 102°20'E; observed by Lord Medway and D. R. Wells (1971, p. 246), Feb. 22-26, 1970; observed by D. Chivers (1971, p. 80; 1973, p. 122), 1968-1970. 16 Kuala Lumpur, Cheras Road, 7th mile; WEST MALAYSIA: Selangor; 3°05'N, 101°45'E; collector unknown, Feb. 8, 1929 (ZRCS). 18 Kuala Lumpur, 22 miles E. See Ulu Gombak Forest Reserve. ?Kuala Lumpur; WEST MALAYSIA: Selangor; ca. 3°10'N, 101°42'E; collector and date unknown (ZRCS). 18 Kuching, vicinity; EAST MALAYSIA: Sarawak; ca. 1°33'N, HO^CE; collected by O. Beccari (1904, p. 30) and G. Doria, 1865 (Museo Civico di Storia Naturale 130 FIELDIANA: ZOOLOGY. VOLUME 67 "Giacomo Dona," Genova [DeBeaux, 1923, p. 36]); collected by Bukong, Nov. 23, 1891, March 31, 1892 (SMK); collector unknown, Oct. 2, 1931 (SMK). 50 Kundur, Pulau; INDONESIA: Sumatra; 0°45'N, 103°26'E; M. n. nemestrina absent (Dammerman, 1926, p. 316). V Kutai Reserve; INDONESIA: Kalimantan; ca. 0°40'N, 117°40'E; observed by P. S. Rodman (1973, p. 655), May, 1970-July, 1971. 79 Labuan, Pulau, mainland opposite; EAST MALAYSIA: Sabah; ca. 5°30'N, 115°30'E; collected by H. Low, 1848-1877 (BM). 73 Lamukotan, Pulo. See Lemukutan, Pulau. Landak, Sungai; INDONESIA: Kalimantan; ca. 0°15'N, 109°45'E; collected by W. L Abbott (in Lyon, 1907b, p. 548), June-Sept., 1905 (USNM). 52 Lam Phura, Sathani; THAILAND: Trang; 7°40'N, 99°35'E; collected by museum collectors (Robinson and Kloss, 1910, p. 670), Jan. 20, 1910 (ZRCS). 2 Lampoengsche Distrikten; INDONESIA: Sumatra 4°-6°S, 104°-106°E; zoo specimen, Aug. 17, 1937 (MZB). (not mapped) Lam-ra. See Lam Phura, Sathani (Moore and Tate, 1965, p. 324). Lang Island. See Krakatau, Kepulauan. Langkat district; INDONESIA: Sumatra; 3°20'-4o20'N, 98o00-98°40,E; observed by E. Hartert, date unknown (Thomas and Hartert, 1894, p. 655). 27 Langsa, Atjeh; INDONESIA: Sumatra; 4°28'N, 97°58'E; collected by Berthold, Feb. 15, 1935 (MZB). 26 Lankat. See Langkat district. Larut; WEST MALAYSIA: Perak; 4°48'N, 100°45'E; collector and date unknown (Taiping Museum [Flower, 1900, p. 315]). 7 Laut. See Lauttador. Laut, Pulau; INDONESIA: Kalimantan; 3°40'S, 116°10'E; M. n. nemestrina absent (Abbott in Lyon, 1911, p. 62). / Lauttador; INDONESIA: Sumatra; 3°18'N, 99°15'E; collected by G. Schneider (1905, p. 73) 1897-1899 (museum unknown). 29 Lemukutan, Pulau; INDONESIA: Kalimantan; 0°45'N, 108°43'E; M. n. nemestrina absent (Abbott in Lyon, 1911, p. 59). h Leuser Reserve; INDONESIA: Sumatra; 2°10'-4o00'N, 97°10'-97°25'E; observed by F. Kurt (1973, p. 64), April-Aug., 1970. 25 Lida Ajer cave, near Pajakumbuh, Padang highlands (Hooijer, 1946, p. 17); INDONESIA: Sumatra; ca. 0°14'S, 100°38'E; subfossils, prehistoric Holocene; collected by E. Dubois, 1888 (presumably in RMNH; Hooijer, 1962a, p. 58). 34 Lima Belas Estate. Sungei Bernam; MALAYSIA: Selangor; 3°46'N, 101°21'E; observed by I. S. Bernstein (1967a, p. 198), 1965-1966. 10 Lima Bias Estate. See Lima Belas Estate. Lingga, Pulau; INDONESIA: Sumatra; O^S, 104°35'E; M. n. nemestrina absent (Dammerman, 1926, p. 316). e FOODEN: LIONTAIL & PIGTAIL MACAQUES 131 Little Kretam River. See Kretam Kechil, Sungai. Little Redang Island. See Bidong Laut, Pulau. Loeboe Basong. See Lubukbasung. Loeboek Karet; INDONESIA: Sumatra; 2°52'S, 104°18'E (Atlas van Nederlandsch Oost-Indie. 1901. p. 8); collected by Soekarno, May 5, 1933 (MZB). 46 Loeboek Linggun. See Lubuklinggau. Long Salai. See Salai, Sungai. Lubukbasung, vicinity; INDONESIA: Sumatra; ca. 0°20'S, 100°04'E; collected by F. von Feber, date unknown, (International Colonial Exhibition, Amsterdam [Jentink, 1883, p. 171]). 32 Lubukbnggau, sea level; INDONESIA: Sumatra; 3°18'S, 102°52'E; collected by J. J. Menden, Nov. 30, 1933 (AMNH-AC). 44 Lumu Lumu, 5500 ft; EAST MALAYSIA: Sabah; ca. 6°02'N, 116°34'E; collected by J. A. Griswold, Jr. (1939, pp. 403, 506), July 20, 1937 (MCZ). 82 Lungmanis Station; EAST MALAYSIA: Sabah; ca. 5°40'N, 117°45'E; observed by K. Stott, Jr. (1964, p. 12), date unknown. 83 Madihit, Sungai; EAST MALAYSIA: Sarawak; 4°11'N, 115°08'E; museum collector, June, 1911 (SMK). 74 Malacca. See Melaka. Mansalar Id. See Musala, Pulau. Mara. See Peleben. Masalembo-Besar; INDONESIA: Kalimantan; 5°34'S, 114°26'E; monkeys absent (Abbott in Lyon, 1911, p. 61). k Masolombo Besar. See Masalembo-Besar. Medan, vicinity; INDONESIA: Sumatra; ca. 3°35'N, 98°40'E; collected by H. Durk. Dec. 29, 1904-Jan. 5, 1905 (ZSBS); Widnmann, 1908 (ZSBS). 28 Melaka; WEST MALAYSIA: Melaka; ca. 2°12'N, 102°15'E; reported by N. Annandale {in Bonhote, 1903, p. 3) and E. J. H. Corner (1941, p. 14). 19 Meru; WEST MALAYSIA: Selangor; 3°08'N, 101°26'E; observed by I. S. Bernstein (1967b, p. 220), May 4, 1965- Feb. 28, 1966. 12 Moyen Bahau. See Bahau, Sungai. Muarabehti, sea level; INDONESIA: Sumatra; 3°15'S, 103°02'E; collected by J. J. Menden, Nov. 26, 1933 (AMNH-AC). 44 Muarabliti. See Muarabeliti. Musala, Pulau; INDONESIA: Sumatra; 1°38'N, 98°32'E; pigtail macaques absent (Kloss, [1928], p. 802). J Natuna, Kepulauan; INDONESIA: Sumatra; 4°00'N, 108°15'E; M. n. nemestrina introduced (Thomas and Hartert, 1894, p. 654). g Niah Caves; EAST MALAYSIA: Sarawak; 3°49'N, 113°47'E; subfossils, late Pleistocene-subrecent; collected by Sarawak Museum, 1954-1962 (Hooijer, 1962b, p. 445). 63 132 FIELDIANA: ZOOLOGY, VOLUME 67 Nias, Pulau; INDONESIA: Sumatra; 1°05'N, 97°35'E; pigtail macaques absent (Kloss, [1928], p. 802). / Nicobar Islands, northern. See under Macaca n. leonina. C Nicobar Islands, southern. See under Macaca n. leonina. D Ogan, Air, upper; INDONESIA: Sumatra; ca. 3°30'S, 104°40'E; collected by Chancellor Stuart Exp., 1929 (FMNH). 48 Padang; INDONESIA: Sumatra; 0°57'S, 100o21'E; collected by S. Muller, 1836 (RMNH); H. Meyer, 1890 (ZMUZ); I. Hagenbeck, May 2, 1925 (NMS). 33 Padang [district]; INDONESIA: Sumatra; ca. 3°15'N, 99°15'E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown). 29 Padang highlands. See Djamboe cave, Lida Ajer cave, and Sibrambang cave. Pagar, Tanjong. See Singapore Island. Pagaralam; INDONESIA: Sumatra; 4°01'S, 103°16'E; collected by A. V. A. Coulin, June 19, 1941 (MZB). 45 Pagurawan. See Paguruan, Sungai. Paguruan, Sungai; INDONESIA: Sumatra; 3°26'N, 99°20'E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown). 29 Pajakombo. See Pajakumbuh. Pajakumbuh; INDONESIA: Sumatra; 0°14'S, 100°38'E; collected by E. Dubois, date unknown (RMNH). 34 Pajakumbuh, near. See Lida Ajer cave. Palembang; INDONESIA: Sumatra; 2°55'S, 104°45'E; collector unknown, Feb. 4, 1921 (RMNH); collector unknown, March 12, 1924 (NHMBe). 47 Palembang [district], sea level; INDONESIA: Sumatra; "3°S, 104°E(appr; collected by W. J. Frost, April 20, 1938 (BM). 46 Pamarkang Bay. See Pamukan, Teluk. Pamukan, Teluk; INDONESIA: Kalimantan; ca. 2°30'S, 116°15'E; collected by W. L. Abbott (in Lyon, 1911, p. 58), March 25, 1909 (USNM). 69 Panaitan, Pulau; INDONESIA: Java; 6°32'-6°40'S, 105°04'-105°14'E; M. n. nemestrina apparently absent (Hoogerwerf, 1953, p. 492). M Pangkalanberandan; INDONESIA: Sumatra; 4°01'N, 98°17'E; collected by Wilhelm Th. A. H. Volz (1912, p. 22), Oct. 14- Nov. 10, 1904 (ZMB). 26 Pangkor, Pulau; WEST MALAYSIA: Perak; 4°13'N, 100°34'E; M. n. nemestrina absent (Harrison and Hendrickson, 1963, p. 548). 0 Papar; EAST MALAYSIA: Sabah; 5°44'N, 115°56'E; collected by R. E. Kuntz, Sept. 12, 1960 (USNM). 72 Pasi; INDONESIA: Sumatra; 5°12'N, 95°17'E; collected by Lt. v. d. Rest, date unknown (MZB). 24 Pa Umor, 3400 ft.; EAST MALAYSIA: Sarawak; 3°44'N, 115031'E; collected by T. Harrisson, Jan. 3, 1948 (FMNH). 75 FOODEN: LIONTAIL & PIGTAIL MACAQUES 133 Pa Umur. See Pa Umor. Pelaihari; INDONESIA: Kalimantan; 3°48'S, 114°45'E; collected by J. H. F. Kohlbrugge (1896, p. 181), Dec 15-21, 1894 (museum unknown). 71 Peleben [on Sungai Kajan near Mara; MS notes in AMNH, reported by Dr. Joseph Curtis Moore, personal communication]; INDONESIA: Kalimantan; ca. 2°45'N, 117°15'E; collected by V. von Plessen, June 24-July 21, 1935 (AMNH-AC; ZMB). 77 Pendeng. See Pulau Munteh. Penang. See Pinang, Pulau. Pennan, Koh. See Phangan, Ko. Perak, Upper; WEST MALAYSIA: Perak; ca. 5°00'N, 101°0CE; reported by N. Annandale (in Bonhote, 1903, p. 3). 7 Perhentian Besar, Pulau; WEST MALAYSIA: Terengganu; 5°54'N, 102°46'E; macaques absent (Kloss, 1911b, p. 180). S Phangan, Ko; THAILAND: Surat Thani; 9°45'N, lOOWE; "M. n. nemestrina is stated on native authority to be found. ..Captive specimens were seen but their provenance was uncertain and they had not improbably been brought from the mainland" (Robinson and Kloss, 1914b, p. 130). (not mapped) Pinang, Pulau; WEST MALAYSIA: Pinang; 5°24'N, 100°14'E; collected by T. Cantor (1846, p. 177; museum unknown); v. Duben, 1846 (NHRM); W. Riitimeyer, 1880 (NHMB); Wistar Inst, of Anat., 1900 (USNM); collector and date unknown (BM). 6 Pleihari. See Pelaihari. Pohorok. See Bohorok. Pontianak; INDONESIA: Kalimantan; 0°02'S, 109°2O'E; collected by P. Diard, [1826], (RMNH). 54 Priaman [district]. See Lubukbasung. Pulau Munteh, 550-750 m.; INDONESIA: Sumatra, 4°05'N, 97°30'E (Chasen, 1941, map opp. p. 1); collected by A. Hoogerwerf (in Chasen, 1940b, p. 485), May 1, 1937 (MZB). 25 Pulo Paku, Betong Saribas; EAST MALAYSIA: Sarawak; ca. 1°30'N, 111°15'E; collected by H. C. Robinson, Oct. 15-Nov. 25, 1916 (BM). 58 Pulutelang; INDONESIA: Sumatra; 3°53'N, 98°20'E; collected by G. Schneider (1905, p. 73), 1897-1899 (NHMB). 26 Puruktjahu; INDONESIA: Kalimantan; 0°35'S, 114°35'E; collected by G. C. Shortridge, Sept. 3, 1909 (BM). 67 Ranau, 1350 ft.; EAST MALAYSIA: Sabah; 5°58'N, 116°41'E; collected by R. E. Kuntz, Sept., 1960 (USNM). 82 Redang, Pulau, group; WEST MALAYSIA: Terengganu; 5°4TN, 103°01'E; M. n. nemestrina absent (Kloss, 1911b, p. 177). T Rengam. See Sungai Renggam. 134 FIELDIANA: ZOOLOGY, VOLUME 67 Rengat, inland; INDONESIA: Sumatra; ca. 0°24'S, 102°33'E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown). 41 Riam, Kotawaringin district, 300 m.; INDONESIA: Kalimantan; 1°50'S, 111°54'E; collected by J. J. Menden, Nov. 6, 1935 (AMNH-AC). 56 Ringat. See Rengat. Roema Manoeal; INDONESIA: Kalimantan; 0°46'N, 111°39'E; collected by J. Biittikofer (1897, p. 12), Dec. 26, 1893 (RMNH; specimen not seen, data from Jentink, 1897, p. 39). 60 Rungkup. See Simpang Empat Rungkup. Sagamafluss. See Segama, Sungai. Sakaiam River. See Sekajam, Sungai. Salai, Sungai; EAST MALAYSIA: Sarawak; ca. 3°30'N, 114°30'E; captive presented to London Zoo by F. Day in 1892, originally obtained about 1888 (Sclater, 1893, p. 325) (BM); collected by C. Hose, Feb.-June, 1891 (SMK). 64 Samarahan. See Entawa-Samarahan. Sandakan, vicinity; EAST MALAYSIA: Sabah; ca. 5°50'N, 118°07'E; collected by H. G. Deignan, June 10, 1937 (MCZ). For note on locality, see Coolidge (1940. p. 130). 87 Sanggul, Bukit, 500 m.; INDONESIA: Sumatra; 3°50'S, 102°37'E; collected by J. J. Menden, Aug. 20-Sept. 2, 1936 (AMNH-AC; MZB). 43 Sapagaya, Sungai; EAST MALAYSIA: Sabah; ca. 5°39'N, 118°06'E; collected by C. F. Adams, Nov. 21, 1887 (USNM). 84 Saribas. See Pulo Paku. Sebang. See Sebangka Pulau. Sebangka, Pulau; INDONESIA: Sumatra; 0°07'N, 104°36'E; M. n. nemestrina absent (Dammerman, 1926, p. 316). c Sebesi, Pulau; INDONESIA: Sumatra; 5°57'S, 105°30'E; primates absent (Dammerman, 1922, p. 75). L Sebesy. See Sebesi, Pulau. Sebuku, Pulau; INDONESIA: Kalimantan; 3°30'S. 116°22'E; M. n. nemestrina absent (Abbott in Lyon, 1911, p. 62). m Segama, Sungai; EAST MALAYSIA: Sabah; ca. 5°30'N. 118°30'E; collected by Dr. Pagel, Aug. 22, 1907 (ZMB). 90 Sekajam, Sungai; INDONESIA: Kalimantan; probably about 0°50'N, 110°25'E; collected by W. L. Abbott (in Lyon. 1907b, pp. 548, 566). June-Sept., 1905 (USNM). 51 Sembilan, Kepulauan; WEST MALAYSIA: Perak; 4°02'N. 100°33'E; At. n. nemestrina absent (Harrison and Hendrickson. 1963. p. 548). R Semitau, vicinity; INDONESIA: Kalimantan; ca. 0°33'N. 111°58'E; observed by J. Biittikofer (1897, p. 11; in Jentink, 1897, p. 39) Nov.- Dec., 1893. 61 FOODEN: LIONTAIL & PIGTAIL MACAQUES 135 Serapit; INDONESIA: Sumatra 3°34'N, ^l^E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown). 27 Serdang [district]; INDONESIA: Sumatra; ca. 3°30'N, 98°50'E; observed by E. Hartert, date unknown (Thomas and Hartert, 1894, p. 655). 29 Siak, Sungai, lower, at mouth of Sungai Gasip; INDONESIA: Sumatra; 0°43'N, 101°42'E; collected by W. L. Abbott, Jan. 1, 1907 (USNM). 38 Sibolga, Teluk; INDONESIA: Sumatra; 1°38'N, 98°45'E; collected by W. L. Abbott (in Miller, 1903a, p. 482), Feb. 19-March 25, 1902 (USNM). 30 Sibrambang cave, Padang highlands; INDONESIA: Sumatra; not precisely located, ca. 0°30'S, 100°30'E; subfossils, prehistoric Holocene; collected by E. Dubois, 1888- 1890 (presumably in RMNH; Hooijer, 1962a, p. 58). 35 Sidong, Gunong, 1200 ft.; EAST MALAYSIA: Sarawak; ca. 1°05'N, 110°25'E (Medway, 1965, map); collector unknown, Oct. 10, 1910 (ZRCS). 50 Simeulue, Pulau; INDONESIA: Sumatra; 2°33'N, 95°55'E; pigtail macaques absent (Abbott in Miller, 1903a, p. 479). F Simpang Empat Rungkup; WEST MALAYSIA: Perak; 3°57'N, 100°43'E; collected by E. S[eimund], Nov. 28, 1913 (BM). 9 Singapore Island; SINGAPORE: 1°20'N, 103°50'E; M. n. nemestrina introduced (Ridley, 1895, p. 26; Annandale in Bonhote, 1903, p. 3). (not mapped) Singkep, Pulau; INDONESIA: Sumatra; 0°30'S, 104°25'E; M. n. nemestrina absent (Dammerman, 1926, p. 316). / Sintang; INDONESIA: Kalimantan; 0°04'N, 111°30'E; collected by C. Bruegel, March 18, 1907 (ZSBS). 62 Sim ass. See Sekajam. Sirambas; INDONESIA: Sumatra; 0°49'N, 99°32'E; collected by E. Modigliani, date unknown (Museo Civico di Storia Naturale "Giacomo Doria," Genova [DeBeaux, 1923, p. 36]). 31 Si-Rambe. See Sirambas. Smitau. See Semitau. Solok; INDONESIA: Sumatra; 0°48'S, 100°39'E; collected by P. O. Stolz, July 17, 1915 (RMNH). 36 Solombo. See Masalembo-Besar. Sugi, Pulau; INDONESIA: Sumatra; 0°50'N, 103°47'E; M. n. nemestrina absent (Dammerman, 1926, p. 316). Z Sukadana; INDONESIA: Kalimantan; 1°15'S, 109°57'E; collected by W. L. Abbott (in Lyon, 1911, p. 54), June 10, 1907 (USNM). 55 Sungai Buloh, 150-300 ft.; WEST MALAYSIA: Selangor; 3°12'N, 101°35'E; collected by Institute for Medical Research, 1947-1957 (Harrison and Hendrickson, 1963, p. 548; Harrison, 1969, p. 176). 11 Sungai Renggam; WEST MALAYSIA: Selangor; 3°03'N, 101°32'E; collector unknown, May 28, 1923 (BM). 12 136 FIELDIANA: ZOOLOGY, VOLUME 67 Sungei Rengam. See Sungai Renggam. Sut, [Sungai], 3000 ft.; EAST MALAYSIA: Sarawak; 2°42'N, 114°39'E; coUected by T. A. Chavasue, Sept. 27, 1955 (FMNH). 66 Tabong; INDONESIA: Kalimantan; 0°34'N, 116°02'E; collected by A. M. R. Wegner, Sept. 13, 1956 (MZB). 80 Tandjong. See Tandjung. Tandjung; INDONESIA: Kalimantan; 2°11'S, 115°23'E; coUected by A. Buxtorf, 1905 (NHMB). 68 Tandjung; INDONESIA: Sumatra; 3°17'N, 99°19'E; collected by G. Schneider (1905, p. 73), 1897-1899 (museum unknown). 29 Tandjungmorawa; INDONESIA: Sumatra; 3°30'N, 98°47'E (Atlas Trop. Ned., pi. 12a); collected by B. Hagen (1890 pp. 80, 82), 1882-1883 (RMNH; also see Jentink, 1887, p. 26); C. Bruegel, 1906 (ZSBS). 28 Tanjong Antu. See Hantu, Tanjong. Tapanuli Bay. See Sibolga, Teluk. Tapisello. See Djamboe cave. Tarusan, Teluk; INDONESIA: Sumatra; 1°13'S, 100°25'E; collected by W. L. Abbott, Dec. 31, 1904 (USNM). 37 Tawau district. See Tibas, Sungai. Tebingtinggi, Deli; INDONESIA: Sumatra; 3°20'N, 99°09'E; coUected by J. A. Coenraad, Nov., 1931, Feb., 1932 (MZB). 29 Temadju, Pulau; INDONESIA: Kalimantan; 0°29'N, 108°52'E; monkeys absent (Abbott in Lyon, 1911, p. 59). i Temaju, Pulo. See Temadju, Pulau. Tibas, Sungai; EAST MALAYSIA: Sabah; 4°26'N, 117°29'E (Davis, 1962, p. 127); collected by R. F. Inger, June 9, 1956 (FMNH). 86 Tinggi, Pulau; WEST MALAYSIA: Johor; 2°18'N, 104°07'E; M. n. nemestrina absent (Robinson, 1919, p. 325). X Tioman, Pulau; WEST MALAYSIA: Pahang; 2°48'N, 104°11'E; M. n. nemestrina said to be absent (Abbott in Miller, 1900, p. 246); collected by J. L. Harrison and J. R. Hendrickson (1963, p. 548), May, 1958 (museum unknown). 22 Trang. See Chong. Trang [province]; THAILAND: Trang; ca. 7°30'N, 99°30'E; collector and date unknown (ZRCS). (not mapped) Ulu Gombak Forest Reserve, 2000 ft.; WEST MALAYSIA: Selangor; ca. 3°20'N, lOrsCE; observed by H. E. McClure ([1965], p. 57), May 20, 1961, March 10, 1962; reported by D. Chivers (1971, p. 80). 18 Ulu Madihih. See Madihit, Sungai. Ulu Segama Forest Reserve, 800-5000 ft.; EAST MALAYSIA: Sabah; ca. 5°11'N, 117°53'E; observed by J. MacKinnon (1971, p. 153), 1968-1970. 85 Ulu Sempan, Raub. See Bukit Fraser. FOODEN: LIONTAIL & PIGTAIL MACAQUES 137 Verlaten Island. See Krakatau, Kepulauan. Macaca nemestrina leonina Map, Figure 28 Adung Valley. See Nam Tamai. Akyab. See Sittwe. Andaman Islands; INDIA: Andaman and Nicobar Islands; 10°-14°N, 92°-93°E; monkeys not indigenous, M. n. leonina introduced (Hamilton, 1870, p. 220). A Annam. See Da Ban. Arakan district, northern hills; BURMA: Arakan Div.; ca. 21°30'N, 93°00'E; collected by A. P. Phayre, 1844 (ZSI); Darling, date unknown (ZMB); collector and date unknown (RMNH); reported by Sclater ([1881], p. 537). 4 Arracan. See Arakan. Assam [state]; INDIA: Assam; 25°-27°N, 90°-95°E; obtained by Mr. Gerrard, purchased March 17, 1885 (IRSN). (not mapped) Bang Hue Horn. See Ban Hue Horn. Ban Huai Maenam Noi, vicinity, ca. 75 m.; THAILAND: Kanchanburi; ca. 14°25'N, 98°51'E; collected by J. Fooden (1971a, p. 32), Feb. 16, 1967 (CTNRC, FMNH). 31 Ban Huang Som; THAILAND: Trat; 11°51'N, 102°5O'E; collected by C. B. Kloss, (1916a, pp. 30, 66), Jan. 11, 1915 (BM). 37 Ban Hue Horn, hill forests north of; THAILAND: Phrae; ca. 17°55'N, 100°10'E; reported by N. Gyldenstolpe (1914, p. 5). 10 Bankachon, Victoria Point; BURMA: Tenasserim Div.; 10°09'N, 98°36'E; collected by G. C. Shortridge (in Wroughton, 1915b, p. 696), Nov. 11-Dec. 29, 1913 (BM; BNHS;ZSI). 44 Bankasun. See Bankachon. Ban Kerng Chada, ca. 150 m.; THAILAND: Kanchanaburi; 15°08'N, 98°31'E; collected by J. Fooden (1971a, p. 32), Feb. 5, 1967 (FMNH). 29 Ban Khlong Wan; THAILAND: Ranong; lO'WN, 98°49'E; collected by H. C. Robinson and C. B. Kloss (1921, p. 9), March 16, 1919 (ZRCS). 46 Ban Nong Kho; THAILAND: Chon Buri 13°09'N, 101°04'E; collected by H. M. Smith, Nov. 13, 1926 (USNM). 35 Ban Nong Kok; THAILAND: Krabi; 8°06'N, 98°52'E; collected by H. C. Robinson and C. B. Kloss (1919, p. 88), Jan. 7, 1918 (ZRCS). 48 Ban Pong Nam Ron, 8-12 km. W and 10 km. NW, ca. 250-300 m.; THAILAND: Kamphaeng Phet; ca. W^N, 99°15'E; collected by J. Fooden (1971a, p. 32), April 12-23, 1967 (FMNH). 21 Ban Tha San, 220 ft.; THAILAND: Chumphon; 10°29/N, 98°55'E; collected by H. C. Robinson and C. B. Kloss (1921, p. 10), March 14, 1919 (ZRCS). 46 Barren Island; INDIA: Andaman and Nicobar Islands; 12°15'N, 93°50'E; monkeys absent (Kloss, 1903, p. 14). B 138 FIELDIANA: ZOOLOGY, VOLUME 67 Bhamo, 20-25 miles below; BURMA: Kachin State; ca. 24°10'N, 97°00'E; erroneous record (Anderson, 1881, p. 72) of M. n. leonina (see above, p. 110). Bien Hoa. See Trang Bom. Bolovens, Plateau des; LAOS: 14°45'-15°30'N, 106°00'-106o50'E; collected by T. D. Carter, Legendre Indo-China Expedition, Feb. 3-13, 1932 (AMNH). For locality note, see Legendre (1932, p. 495). 17 Chamchay, Cochinchine; SOUTH VIETNAM: not precisely located, 8°30'-12°30'N, 104°30'-107°30'E; collected by M. Pierre, April 1874 (MNHN). (not mapped) Champang; BURMA: Tenasserim Div.; 10°13'N, 98°31'E; collected by W. L. Abbott, Dec. 21-22, 1903 (USNM). 44 Chan, Ko; THAILAND: Ranong, Mergui Archipelago; 9°25'N, 97°50'E (Moore and Tate, 1965, p. 323); collected by W. L. Abbott, Dec. 30, 1899-Jan. 1, 1900 (USNM ). 45 Chance Island. See Chan, Ko (Moore and Tate, 1965, p. 317). Changchang Pani, 500 ft.; INDIA: Nagaland; 26°35'N, 94°25'E; collected by C. McCann (1933a, p. 395), Feb. 7-19, 1930 (AMNH; BNHS). 1 Chantabun. See Chantaburi. Chantaburi; THAILAND: Chantaburi; 12°36'N, 102°09'E; collected by C. J. Aagaard, Dec. 16-17, 1926 (ZRCS). 36 Chongkrong, 600-900 m.; THAILAND: Kanchanaburi; 14°41'N, 98°52'E; coUected by J. Fooden (1971a, p. 32), Jan. 28, 1967. 30 Cochinchine. See Chamchay. Da Ban; SOUTH VIETNAM: Ninh Thuan; ca. 11°45'N, 108°45'E, collected by C. B. Kloss (1919a, p. 395), March 24, 1918 (ZRCS). 41 Ghirbi. See Ban Nong Kok. Hin Lap, Sathani; THAILAND: Sara Bun; 14°40,N, 101°09'E; coUected by H. M. Smith, Oct. 2, 1932 (USNM). 24 Hkamti. See Singkaling Hkamti. Hue\ vicinity; SOUTH VIETNAM: Thua Thien; 16°28'N, 107°36'E; reported by J. Delacour (1940, p. 24). 19 Junk Seylon. See Klong Tung Sai. Kao Sabab. See Sa Bap, Khao. Kawkareik; BURMA: Kawthule State; 16°33'N, 98°14'E; collected by L. Fea, May 1887 (Museo Civico di Storia Naturale "Giacomo Doria," Genova [Thomas, 1892, p. 916; DeBeaux, 1923, p. 34]). 20 Khlung, Khlong, upper; THAILAND: Kamphaeng Phet; ca. 16°05'N, 99°20'E; collected by C. C. Sanborn (1952, p. 2) and F. C. Wonder, July 2, 1949 (FMNH). 22 Khun Tan, mountains; THAILAND: Lamphun; ca. 18°30'N, 99°15'E; coUected by N. Gyldenstolpe ([1917], p. 7), June 7, 1914 (NHRM). 9 Klet Kaeo, Ko; THAILAND: Chon Bun; 12°46'N, 100°51'E; pigtail macaques absent (Berkson et al., 1971, p. 237). F FOODEN: LIONTAIL & PIGTAIL MACAQUES 139 Klong Klung. See Khlung, Khlong. Klong Menao. See Ban Huang Som (Moore and Tate, 1965, p. 323). Klong Tundai. See Klong Tung Sai. Klong Tung Sai, Ko Phuket; THAILAND: Phuket; 7°98'N, ^^E (Robinson and Kloss, 1919, p. 88); collected by C. B. Kloss, Dec. 20-26, 1917 (BM; ZRCS). For comment on locality, see Chasen, 1935, p. 38. 47 Klong Tun Sai. See Klong Tung Sai. Klong Wan. See Ban Khlong Wan. Kokareet. See Kawkareik. Ko Keow, vicinity, ca. 200 m.; THAILAND: Kamphaeng Phet; ca. 15°55'N, 99°25'E; J. Fooden (1971a, p. 32), March 9, 1967 (FMNH).22 Koon Tan. See Khun Tan. Krabi. See Ban Nong Kok. Lampi Island. See Lanbi Kyun. Lanbi Kyun; BURMA: Tenasserim Div., Mergui Archipelago; 10°50'N, 98°15'E; collected by C. Primrose, March 3, 1922 (BM). 42 Lat Bua Kao. See Lat Bua Khao. Lat Bua Khao, Sathani, north of; THAILAND: Nakhon Ratchasima; 14°52'N, 101°36'E; collected by C. B. Kloss (1919b, p. 343), Oct. 12, 1916 (USNM). 26 Meh Lem river, north of; THAILAND: Phrae; ca. 18°25'N, 100°20'E (Gyldenstolpe, 1913, p. 4; 1916 pl. 1); observed and tentatively identified by N. Gyldenstolpe (1914, p. 4), March 5-27, 1912. ;/ Meng-hai; CHINA: Yunnan; 21°52'N, 100°28'E; collected by Kao Yuen-ting et al. (1962, p. 188), 1957-1958 (probably in collection of Institute of Zoology, Academia Sinica, Peking). 8 Mong Nai; BURMA: Shan State; 20°31'N, 97°52'E; collected by Dr. Griggs, in or before April, 1894 (Brown, 1896, p. 485). 7 Muang Leip. See Muang Liap. Muang Liap; LAOS: Xaignabouri; 18°29'N, 101 °40'E; collected by J. Bangga, Jan. 15, 1920 (ZRCS). 12 Nam Tamai; BURMA: Kachin State; 27°42'N, 97°54"E (Moore and Tate, 1965, p. 328); probably erroneous record (Dollman, 1932, p. 9; see above p. 112). Nicobar Islands, northern; INDIA: Andaman and Nicobar Islands; 8°-10°N, 92°- 94°E; monkeys absent (Kloss, 1903, p. 114). C Nicobar Islands, southern; INDIA: Andaman and Nicobar Islands; 7°-8°N, 93°-94°; pigtail macaques absent (Kloss, [1928], p. 802). D Nong Khor. See Ban Nong Kho (Chasen, 1935, p. 32; Moore and Tate, 1965 p. 329). Nong Kok. See Ban Nong Kok. Pak Chong, Sathani, 900 ft.; THAILAND: Nakhon Ratchasima: 14°42'N, 101°25'E; 140 FIELDIANA: ZOOLOGY, VOLUME 67 collected by E. G. Herbert and M. Smith, Dec. 4, 1915 (BM; Kloss, 1916b, p. 2) and H. M. Smith, March 2, 1924 (USNM). 25 Pak Jong. See Pak Chong, Sathani. Paksane (province); LAOS: ca. WSCN, 104°00'E; collected by J. Deuve and M. Deuve (1963, p. 60), date and museum unknown. 14 Pakse (province); LAOS: ca. 15°00'N, 106°00'E; collected by J. Deuve and M. Deuve (1963, p. 60), date and museum unknown. 17 Panjang, Pulau. See Telok Poh. Pan-na-meng-hai. See Meng-hai. Paungdaw power station, hill NW, 3000 ft.; BURMA: Tenasserim Div.; ca. 14°00'N, 98°30'E; collected by J. Keenan, July 12, 1961 (BM). 32 "Perak"; WEST MALAYSIA: Perak; ca. 5°00'N, 101°00'E; presented by O. L. Fraser, Jan. 6, 1878 (ZSI). Erroneous locality (see above, p. 110). Petchaburi. See Phet Buri. Phet Buri [province]; THAILAND: Phet Buri; 12°35'-13°10'N, 99°15'-100°05'E; reported by K. G. Gairdner (1914, p. 36). 34 Phu Kheo Forest Reserve; THAILAND: Chaiyaphum; ca. 16°15'N, 101°30'E; remains of monkeys eaten by humans, reported by J. A. McNeely and E. W. Cronin (1972, p. 459), Jan. 1972. 27 Phuket, Ko. See Klong Tung Sai. Pidaung Game Sanctuary; BURMA: Kachin State; 25°25'N, 97°09'E; observed by U Tun Yin (1954, p. 269), March, 1951-Feb., 1952. 3 Plateau des Bolovens. See Bolovens, Plateau des. Puket. See Klong Tung Sai. Rat Buri [province]; THAILAND: Rat Buri; 13°10'-13°45'N, 99°20'-100°05'E; reported by K. G. Gairdner (1914, p. 36). 33 Red Point; BURMA: Tenasserim Div.; 10°40'N, 98°30'E (Moore and Tate, 1965, p. 331); collected by W. L. Abbott, Feb. 20, 1904 (USNM). 43 Ross Island. See Andaman Islands. Sa Bap, Khao; THAILAND: Chanthaburi; 12°31'N, HE'lffE; collected by H. M. Smith, Nov. 5, 1933 (USNM). 36 Sai Gon; SOUTH VIETNAM: Gai Dinh; 10°45'N, 106°40'E; reported by J. Delacour (1940, p. 24). 40 Salanga. See Klong Tung Sai. Samnak Rabam, 10 km. SE, ca. 200 m.; THAILAND: Uthai Thani; 15°25'N, 99°30'E; collected by J. Fooden (1971a, p. 32), Feb. 26, 1967 (CTNRC, FMNH). 23 Saravane (province); LAOS: ca. 15°45'N, 106°45'E; collected by J. Deuve and M. Deuve (1963, p. 60), date and museum unknown. 18 Savannahket (province); LAOS: ca. 16°30'N, 104°45'E; collected by J. Deuve and M. Deuve (1963, p. 60), date and museum unknown. 16 FOODEN: LIONTAIL & PIGTAIL MACAQUES 141 Selok Poh. See Telok Poh. "Siam"; THAILAND: no exact locality; collected by R. C. Bulkeley, date unknown (AMNH). (not mapped) Singkaling Hkamti, about 20 miles north [east], W bank of Chindwin River, 500 ft.; BURMA: Sagaing Div.; ca. 26°10'N, 96°00'E; collected by G. C. Shortridge and S. A. Macmillan, July 26, 1914 (BM). For locality note, see Wroughton, 1916a, p. 293. 2 Sittwe, hills inland; BURMA: Arakan Div.; ca. 20°00'N, 93°30'E; reported by S. R. Tickell (1854-1863, MS., p. 112; Pocock, 1939, p. 60) and E. Blyth (1863, appendix p. [189]; in Dunn, 1864, p. 370). 5 Sullivan's Island. See Lanbi Kyun. Taho, 1200-1300 m.; BURMA: Shan State; 19°27'N, 96°53'E; collected by L. Fea (1888, pp. 855, 859), Feb. 29, 1888 (Museo Civico di Storia Naturale "Giacomo Doria," Genova [Thomas, 1892, p. 916; DeBeaux, 1923, p. 34]). 6 Tapli. See Ban Khlong Wan. Tasan. See Ban Tha San. Tavoy. See Ye. Tay Ninh; SOUTH VIETNAM: Tay Ninh; 11°18'N, 106°05'E, reported by A. Morice(1875, p. 41). 38 Telok Besar; BURMA: Tenasserim Div.; 10°22'N, 98°35'E (Moore and Tate, 1965, p. 333); collected by W. L. Abbott, March 17, 1904 (USNM). 44 Telok Poh, Ko Yao Yai; THAILAND: Phangnga; 8°00'N, 98°38'E (Robinson and Kloss, 1919, p. 88); collected by C. B. Kloss, Jan. 22, 1918 (BM). For comment on locality, see Chasen, 1935, p. 38. 47 Thakhek (province); LAOS: ca. 17°30'N, 105°00'E; collected by J. Deuve and M. Deuve (1963, p. 60), date and museum unknown. 15 Thad. See Taho. Tongha. See Klong Tung Sai. Tongka. See Klong Tung Sai. Trang Bom; SOUTH VIETNAM: Bien Hoa, 10°57'N, W7°WE; collected by C. B. Kloss (1919a, p. 401), June 5, 1918 (ZRCS); M. Poilane, Sept. 4, 1932 (USNM; for locality restriction, see Van Peenen et al., 1969, p. 102). 39 Victoria Point. See Bankachon. Vientiane (province); LAOS: ca. 18°00'N, 102°3O'E; collected by J. Deuve and M. Deuve (1963, p. 60); date and museum unknown. 13 Yao Yai, Ko. See Telok Poh. Ye, forest near; BURMA: Tenasserim Div.; ca. 15°15'N, 98°00'E; collected by S. R. Tickell (1854-1875, MS., pp. 112, 137, 139), Nov., 1854 and Jan.-Feb., 1860 (museum unknown). 28 Yey. See Ye. Zingkalling. See Singkaling Hkamti. 142 FIELDIANA: ZOOLOGY, VOLUME 67 Macaca nemestrina pagensis Map, Figure 25 Mentawai, Kepulauan. See Sipura, Pulau; Pagai Utara, Pulau; Pagai Selatan, Pulau. North Pagi. See Pagai Utara, Pulau. Pagai Selatan, Pulau, sea level; INDONESIA: Sumatra; ca. 3°00'S, 100°20'E; collected by W. L. Abbott, Nov. 17, 1902 (USNM) and J. J. 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Significant measure- ments (mm.) of this specimen are: head and body length, 574; tail length, 162 (relative tail length, 0.28); greatest length of skull excluding incisors, 141.5; zygomatic breadth, 99.6; rostral length, 56.3; postrostral length, 95.5; length of incisor-canine field, 18.1; length of premolar-molar field, 37.8. These new measurements slightly extend the range of relative tail length and greatest skull length specified above (pp. 7, 18, 112, 115) for this subspecies; inclusion of these measurements in graphs of dimensional charac- ters (figs. 2, 7, 8, 9, 22) would have relatively minor effects. 2. Three recent publications, listed below, provide valuable morphological information concerning growth changes and sex differences in Macaca nemestrina. Garn, S. M., D. R. Swindler, and P. E. Minnis 1975. The sex difference in dimensional communalities in Macaca nemestrina. Amer. J. Phys. Anthropol., 43, pp. 19-22. Sirianni, J. E., D. R. Swindler, and L. H. Tarrant 1975. Somatometry of newborn Macaca nemestrina. Folia primatoL, 24, pp. 16-23. Tarrant, L. H. 1975. Postnatal growth in the pig-tailed monkey (Macaca nemestrina). Ph. D. Thesis. Univ. of Washington, Seattle, ii+ 113 pp. 3. Supplementary locality records (unmapped) to be added to the Gazetteer of Collecting Localities (pp. 112-142) are documented below. Macaca silenus Shimoga Dist., 2000 ft., INDIA: Mysore; ca. 14°00'N, 75°30'E; observed by R. Whitaker, Mar., 1972 (Pruett, C, 1974, J. Bombay Nat. Hist. Soc., 70, p. 547). 168 FOODEN: LIONTAIL & PIGTAIL MACAQUES 169 Silent Valley, ca. 4200 ft.; INDIA: Kerala; ca. 11°10'N, 76°30'E; troop including 11- 12 adults (2 females with young infants) observed by C. Pruett (ibid., pp. 546-547) and D. Hayles, Mar., 1972. Singampatti Hills (ibid., p. 546). See Manjolai Tea Estate. Macaca nemestrina nemestrina Kenangan, near; INDONESIA: Kalimantan; ca. 1°05'S, 116°45'E; observed by C. C. Wilson and W. L. Wilson (1975, Folia prima toL, 23, pp. 252, 253), March, 1973. Macaca nemestrina leonina Soi Dao, Khao, 3500 ft.; THAILAND: Nakhon Sawan; lS'WN; 100°44'E; collected by R. M. de Schauensee (1946, Proc. Acad. Nat. Sci. Philad., 98, p. 5), May 25, 1938 (Academy of Natural Sciences, Philadelphia, 1, skin and skeleton). Macaca nemestrina pagensis Tei-Tei Peleigei, near; INDONESIA: Sumatra (Pulau Siberut); ca. 1°24'S, 99°01'E; reported by R. R. Tenaza (1975, Folia primatol., 24, p. 67), July 1-Oct. 7, 1972. Pub