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Full text of "Taxonomy and evolution of liontail and pigtail macaques (Primates : Cercopithecidae)"

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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 

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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. 



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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. 




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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) 



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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). This indicates that 
there probably is a constant allometric relationship between rostral 



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Fig. 8. Allometry of rostral length v. postrostral length in immature and adult 
lion tails and pigtails. Log-log scales; slope - 3.543. 



14 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 15 

length (y) and post -rostral length (x) in these macaques. The power 
function equation for this relationship computed (with generous 
assistance of Dr. Gary E. Eertmoed, Chicago State University) by 
the method of principal axes is log y =3.543 log x- 5.222; 95 per cent 
confidence limits for the slope are Li = 3.381 and L,2 = 3.720. 
Numerical values in this power function equation are comparable to 
those in equations for the relationship between muzzle length and 
calvaria length in species of baboons studied by Freedman (1962, pp. 
121-125); as in the baboon scattergrams, points in Figure 8 for very 
young specimens tend to lie to the right of the regression line 
(relative rostral lengths less than expected). The conspicuous sexual 
and taxonomic differentiation of relative rostral length in adult 
liontails and pigtails appears, as in baboons (Freedman, 1962, p. 
127), to be a result of differential rates of ontogenetic development 
of a constant allometric relationship between growth of rostrum 
(facial skeleton) and growth of postrostrum (braincase). Since 
cranial dimensions are known to be approximately equal in 
immature male and female M. n. nemestrina up to age three years 
(Swindler et al., 1973, p. 229), the rate differential that results in 
adult sexual dimorphism of relative rostral length apparently 
develops (at least in this subspecies) sometime between age three 
years and the age of physical maturity (probably about age six or 
seven years). 

Rostral length and postrostral length, although allometrically 
related to each other, are each linearly related to length of 
component parts of the adult maxillary dental arch (fig. 9). 
Relevant measurements taken of the dental arch are: (1) total 
length of arch— perpendicular distance from most anterior point on 
premaxilla to line passing through most posterior points on alveoli 
of third molars; (2) length of premolar-molar field — alveolar 
distance from front of first premolar to rear of last molar (measured 
on right side, wherever possible); (3) length of incisor-canine field — 
difference between total length of arch and length of premolar- 
molar field. 

Length of the premolar-molar field typically is 35-40 per cent of 
postrostral length in both sexes of all four subspecies and species of 
adult liontails and pigtails (fig. 9a); similarly, length of the incisor- 
canine field typically is 30-40 per cent of rostral length (fig. 9b). 
From the allometric relationship that exists between rostral length 
and postrostral length, it follows that length of the incisor-canine 
field is allometrically related to length of the premolar-molar field. 



16 



FIELDIANA: ZOOLOGY, VOLUME 67 




Rostral length (mm) 



Fig. 9. Relative length of dental fields in adult liontails and pigtails; a, length of 
premolar- molar field v. postrostral length (linear scales); b, length of incisor-canine 
field v. rostral length (linear scales). 



In specimens examined of M. silenus, relative length of the incisor- 
canine field tends to be slightly greater (fig. 9b) and relative length 
of the premolar-molar field tends to be slightly less (fig. 9a) than in 
the three subspecies of M. nemestrina. 

Supraorbital ridges (fig. 3) in M. silenus are strongly projecting, 
although relatively thin (about 5 mm. thick in adult males). In M. 
n. leonina and M. n. pagensis these ridges are somewhat thicker (7- 
8 mm.), but they tend to recede laterally. In M. n. nemestrina 
supraorbital ridges are relatively thin (5 mm.), especially dorsolat- 
eral^, and strongly receding. In M. n. leonina, M. n. nemestrina, 
and M. n. pagensis the malar region is flat to weakly convex; in M. 
silenus well-developed intraorbital concavities are present in this 
region. In none of these macaques do the temporal ridges meet in 
the midline to form a sagittal crest. 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 17 

Size, relative rostral length, and relative dental field 
proportions evidently constitute a complex of interrelated elements 
in the cranial evolution of liontail and pigtail macaques. Greatest 
length of skull is trigonometrically related to rostral length and 
postrostral length; rostral length is allometrically related to 
postrostral length; and rostral length and postrostral length are 
linearly related to length of incisor-canine field and premolar-molar 
field. It is not clear which of these interrelated cranial elements are 
primary factors in evolution and which are secondary. Cranial 
differentiation in these macaques may be the result of selective 
forces that act primarily on overall size, size of the braincase, size of 
the facial skeleton, size of the incisors and canines, size of the 
premolars and molars, on some combination of these, or on none. 
Present evidence does not appear adequate to indicate a clear choice 
among these alternative evolutionary possibilities. 



TAIL - PROPORTIONS, PELAGE, VERTEBRAE, CARRIAGE 

Relative tail length (tail /head and body) in adult male and 
female specimens examined varies from 0.24 - 0.34 in M. n. 
pagensis, from 0.28 - 0.45 in M. n. nemestrina, from 0.30 - 0.46 in 
M. n. leonina, and from 0.57 - 0.75 in M. silenus (fig. 2). (The 
denominator used in computing these ratios is the standard linear 
distance from tip of nose to base of tail, as measured by 
mammalogists, not the crown-rump height customarily employed 
by anthropologists.) The short, thin tail in the three subspecies of 
M. nemestrina is, of course, the basis for their collective designation 
as "pigtail" macaques. Judging from the few known specimens of M. 
n. pagensis, relative tail length in this subspecies averages slightly 
less than in the other two pigtail subspecies, which are essentially 
similar in tail proportions, although different in overall size. In M. 
n. leonina and M. n. nemestrina, the two subspecies for which 
reasonably adequate samples are available, relative tail length tends 
to be greater in infants (head and body length less than 300 mm.; 
probably less than one year old) than in juveniles and adults (cf. 
Lumer and Schultz, 1941, p. 297). The rate of growth of the tail 
relative to that of the head and body also is known to decline with 
age, at least until age two or three years, in M. mulatto (Schultz, 
1933, p. 18) and M. fascicularis (Karrer, 1970, p. 174). 

In M. silenus the fur of the tail is uniformly blackish, like that 
of the trunk; guard hairs at the tip of the tail usually are elongated 
to form a well-defined terminal tuft— hence, the "liontail" (fig. 21). 
In M. n. leonina and M. n. nemestrina the fur of the tail is sharply 
defined bicolor, dark brown to blackish dorsally and buffy 
ventrally; in M. n. nemestrina the hairs at the tip frequently are 
slightly elongated and reddish-brown to ochraceous, forming a 
weakly defined tuft. In M. n. pagensis the tail is virtually naked, 
superficially resembling that of an opossum. The dark chocolate- 
brown fur of the lower back in this subspecies extends only to the 
base of the dorsal surface of the tail; more distally, for 80-90 per 
cent of its length, the tail is sparsely covered by inconspicuous short 

18 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 19 

TABLE 2. Reported caudal vertebrae counts in M. n. leonina and M. n. 
nemestrina. 

Descriptions of Number of caudal vertebrae/frequency 

specimens Sex 14 15 16 17 18 19 20 

M. n. leonina 

Skeletons* $ 111 

Skeletons 1 & 11 

Cadaver S 2 5 1 2 

Cadavers2 S 2 13 



Skeletons 1 $ 1 

Skeletons 1 £ 3 2 

(?) M. n. leonina orM. n. nemestrina^ 

Living ? Ill 

Living <5 112 

Skeletons 5 2 2 1 

Skeletons 3 112 2 11 

1- See Table 3. 

2- In collection of Dr. Russell H. Tuttle, Department of Anthropology, The 
University of Chicago; vertebral counts reported by Wilson (1970, p. 186). 

3 - Reported by Wilson (1970, p. 186; 1972, p. 245), who does not distinguish 
between M. n. leonina and M. n. nemestrina. 

buffy hairs (2-3 mm. long) through which the corrugated surface of 
the epidermis is plainly visible. 

The number of caudal vertebrae evidently averages about 21 or 
22 in M. silenus and about 18 or 19 in M. n. leonina and M. n. 
nemestrina (tables 2, 3; Schlegel, 1876, p. Ill; Anderson, 1879, p. 
55). Caudal vertebrae counts in specimens examined of M. n. 
leonina and M. n. nemestrina apparently agree with those in 
specimens studied by Schultz (1938, p. 6), who reports an average of 
18.7 caudal vertebrae in six pigtail macaques, all or most of which 
probably are M. n. nemestrina (G. M. Allen and Coolidge, 1940, p. 
147). A group of aberrantly low caudal vertebrae counts of 
uncertain validity is reported by Wilson (1970, p. 187; 1972, p. 245) 
in inadequately identified skeletal material (table 2). 

The length of each of the proximal three caudal vertebrae (Cal, 
Ca2, Ca3) is approximately 10.5 mm. in adult male M. n. leonina 
and M. n. nemestrina and approximately 12.5 mm. in adult male M. 
silenus (table 3). The next four vertebrae (Ca4-Ca7) usually become 
successively longer in all three of these macaques. Vertebra Ca7, 



— 

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23 



24 FIELDIANA: ZOOLOGY, VOLUME 67 

which tends to be the longest member of the series, averages about 
19 mm. long in adult male M. n. leonina and M. n. nemestrina and 
about 28.5 mm. long in adult male M. silenus. More distal vertebrae 
typically become successively shorter, about 2 mm. per vertebra, to 
a terminal vertebral length of 1-5 mm. The pattern of caudal 
vertebral length variation presented here for adult males is 
essentially similar to that in a series of 10 males and females, many 
of which apparently are immature, that is reported by Wilson (1970, 
p. 198). Comparing M. silenus with M. n. leonina and M. n. 
nemestrina, the greatest reduction in vertebral length is at the 
longest vertebra in the series, which, as noted, is usually Ca7. This 
vertebra also appears to be at or near the focus of evolutionary tail 
reduction in Celebesian stumptail macaques (Fooden, 1969a, p. 17) 
and in crab-eating and rhesus macaques (Wilson, 1970, p. 60). 

In adult male M. silenus the relaxed tail is carried hanging 
posteriorly (fig. 21; Karr, 1973, p. 191), in what apparently is the 
typical manner in most long-tailed monkeys. In adult male M. n. 
leonina the base of the tail normally is inflected anteriorly, and the 
tail is carried arched over the back with the tip directed upward and 
forward. Judging from published illustrations (Sclater, [1870], pi. 
35; Yerkes Newsletter, 1971, p. 14) and observation of captive 
specimens (fig. 29), the tail in adult male M. n. leonina is seldom, if 
ever, dropped posteriorly (Tickell, 1863-1875, MS., p. 139; Heck, 
1916, p. 552). In adult male M. n. nemestrina the tail is carried 
arched rearward, with the base inflected dorsally and the distal part 
inflected downward (fig. 26; Wilson, 1970, p. 115). The manner of 
tail carriage in M . n. pagensis has not been recorded. 

I have attempted to investigate the anatomical basis for the 
difference of tail carriage in M. n. leonina and M. n. nemestrina. 
Material available for detailed study consists of embalmed cadavers 
of three adult males of M. n. leonina (kindly lent by Dr. Russell H. 
Tuttle, Department of Anthropology, The University of Chicago; 
spec. nos. NM6, PMl, PM2), caudal vertebrae of three wild-caught 
adults of M. n. leonina (FMNH 99673 6", 99688?, 99091c?), and 
caudal vertebrae of two captive subadults of M. n. nemestrina 
(FMNH 54252 6\ 54305$). Judging from cadavers examined, the 
first two or three postsacral vertebrae, although osteologically 
"caudal," are, in fact, located in the sacral region of the trunk, 
dorsal to the rectum and anterior to the anus (fig. 10). The vertebra 
at the base of the external tail is Ca3 or Ca4 (Wilson, 1970, p. 48). 
The upward inclination of the tail base that is evident in living M. 



•o::z> u;:~oi t ?::- - -:i •■..-: - v ". l- 



_; 







?jjmn££ wi r msrniiamL nf ""'i"' "snenras it 



jl fcoiuna <%. 29 » and Jf. jl aenaestfrina «*%. : 
:-ff_.: :: i:-«..i. - :t:- :'r -?.;- :: _- :~r t^t-: :-i_. 
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 






54262 1 


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.). It is also generally similar to that in M. 



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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. 



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36 FIELDIANA: ZOOLOGY, VOLUME 67 

67); in M. arctoides, with glans lanceolate, cervical colliculi are 
absent and columnar epithelium extends unusually far distally 
(Fooden, 1967, p. 940; Demers et al., 1972, p. 532; Hafez and 
Jaszczak, 1972, p. 304). Information on cervical structure in other 
species of the genus has not been recorded. 




Fig. 13. Penis in adult M. n. leonina (FMNH 99691), left lateral view and dorsal 
view, x 1.5. (Photo FMNH) 

Patterns of copulatory behavior apparently also are distinctive 
in each of the three subdivisions of macaques that are defined by 
male and female genital morphology (Michael et al., 1973, p. 248; 
Nadler and Rosenblum, 1973a, p. 18; 1973b, p. 217). In M. n. 
nemestrina, M. mulatto., and M. fuscata (glans rounded, cervix 
moderately large), copulation usually consists of a series of non- 
ejaculatory mounts that precede the ejaculatory mount (median 7- 
11 mounts per ejaculation); although evidence concerning M. 
fascicularis is ambiguous, copulatory behavior in this species seems 
to be basically similar to that in M. mulatto (Goustard, 1968, p. 468; 
Kanagawa et al., 1972, p. 453; Kanagawa and Hafez, 1973, p. 234). 
In M. radiata (glans apically elongate, cervix inflated) copulation 
usually is completed in one mount (maximum four mounts) with 5- 
30 intromissive thrusts to ejaculation. In M. arctoides (glans 
lanceolate, cervix simplified) copulation also usually is completed in 
one or a few mounts, but with about 60-70 thrusts to ejaculation. 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 



37 



The correlated variation of male genitalia, female genitalia, and 
copulatory behavior in these three macaque subgroups undoubtedly 
is related, in a way not yet understood, to the history of 
evolutionary differentiation and reproductive isolation in the genus. 

Although the bluntly rounded form of the glans penis in 
liontail and pigtail macaques is generally similar to that in the 
mulatta-group, and length of the glans is broadly overlapping in the 
two groups, diameter of the glans in the liontail-pigtail group 
averages nearly twice as great as in the mulatta-group. In specimens 
examined, the ratio of glans breadth to glans length is 0.60-0.89 in 
liontail and pigtail macaques (tables 6, 9) as contrasted with 0.36- 
0.55 in the mulatta-group (table 7). Relative glans breadth in 
Celebesian macaques appears to be about the same as in liontail 

TABLE 7. Relative breadth of glans penis in fluid-preserved specimens 1 of 
M. fascicularis, M. mulatto, and M. fuscata. 





Estimated 


Dental 


age^ 


stage^ 


(years) 


dec. nr»2 


1 


(Mi) 


1.5 


dl) 


2.5 


(P) 


4 


(C, M3) 


6 


M3 


>6 


Ml 


2 


(12, M2) 


3 


(C) 


4.5 


M3 


>6 


— 


3 


(C) 


5 


— 


>5 


— 


>6 


— 


>7 



Glans length 4 (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 


ita 6 


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 



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FOODEN: LIONTAIL & PIGTAIL MACAQUES 



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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 \ -antitrypsin 4 

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« 




PA f s 






100.0 











97.2 




2.8 






1 


6-1 Unidentified 






100.0 












94.5 


1.4 






A 




AC 






100.0 











94.3 




5.7 






AB B 


AC BC C CD 


BD 


D 


1.6 





7.8 68.7 14.1 


1.6 


6.2 


0.7 6.6 


1. 


3 32.4 58.3 0.7 









BG CC CD CF CG D"D DD DF 



1.2 1.2 3.5 4.7 12.9 

0.6 3.2 4.5 2.5 

DG F'GFF' F'G FF 

17.7 3.5 

23.4 0.6 1.3 0.6 7.6 



1.2 8.2 3.5 
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 










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 





97.1 


2.9 


a b ab 


c d o 


7.9 57.1 1.6 


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 






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 (Gd A ), erythrocyte 6-phosphogluconate 
dehydrogenase (6PGD A ; cf. table 10) and haptoglobin (Hpl). The 
same colony was polymorphic for phenotypes of serum pseudocholi- 
nesterase (E«; E. s , Ei u , hypothesized), transferrin (Tf A , Tf B , Tf D ), 
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 Le a antigens, and 
12 of 123 tested were positive for Le b 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 



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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 



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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 -10,000 4(16%) 21(84%) 

25- 49 10,000-20,000 37(100%) 

50- 74 20,000-30,000 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 fX n ) 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. 



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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. 16 1 
Blanford, 1888°, p. 16l 



AMNH 548372 
Anderson, 1879, p. 93 
Blanford, 1888b, p. 16* 
Blanford, 1888b, p. 16 1 
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 - 



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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 





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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. <J, ad. ?, 
Muarabeliti). For a detailed study of activity cycles in a captive 
colony of M . n. nemestrina, see Bernstein (1972, p. 390) and works 
cited therein. 

Observers agree that the principal habitat of M. n. nemestrina 
is dense evergreen rainforest (Be-a, Be-b, D, Hg, Hss/D, K/M, L, 
McC, Mii, So/C; Menden specimens). The known alitudinal range 
of this subspecies extends from near sea level (Sumatra: Sungai 
Kateman, Sungai Siak) to upland elevations of 550-750 m. in 
Sumatra (Pulau Munteh), 1000 m. in West Malaysia (Me), and 1700 
m. in Borneo (Lumu Lumu). In West Malaysia M. n. nemestrina is 
rare in coastal areas (C, L, So/C). In East Malaysia one author (Ba) 
has reported that coastal areas are a preferred habitat; but this has 
not been confirmed by more recent observers (K/M). M. n. 
nemestrina seems to be a rare visitor in some suitable habitats; one 
tract of inland forest in West Malaysia evidently was visited by this 
subepeciee only twice in two years (McC). Recent persecution by 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 103 

man has completely eliminated M. n. nemestrina from some areas 
that it formerly inhabited (Be-b, Hss/M). 

Reported counts or estimates of troop size in this subspecies 
are: 3, 5 (Me); 6, 9 (McC); seldom more than a dozen (Hg); 6-15 
(Sch); 5-20 (L); 20-30 (Ku); 30, 47 (Be-a). The two largest troops 
reported (30 and 47 individuals) were confined to a relatively small 
and isolated forest tract and therefore may not be typical. Solitary 
males are encountered frequently (Ba, Be-b, K, Me, St; Boogaarts 
specimen). 

The population density of M. n. nemestrina in suitable habitats 
in West Malaysia has been estimated at 0.13 troops per square mile 
(So/C). Throughout its range this macaque is distinctly less 
abundant than sympatric M. fascicularis (D, Hg, Hrt, K/M, L, 
Me). In West Malaysia, the reported ratio of M. n. nemestrina to 
M. fascicularis in three local population surveys (Hs, Me/W, So/C) 
is 19:148 individuals (0.13), 3:22 individuals (0.14), and 1:13 troops 
(0.08). In eastern Kalimantan, the reported ratio of M. n. 
nemestrina to M. fascicularis in one study (R) is 40:268 groups 
(0.15). 

M. n. nemestrina is often encountered on the forest floor (Ba, 
Be-b, Hg, Hrt/T, Hss/D, Mii, R, Sch, St). However, the subspecies 
also has frequently been observed to move through the branches of 
trees (Be-a, D, Hg, McC; also see Chivers, 1973, p. 130). In East 
Malaysia, two young males (FMNH 85923, 85924; Sungai Tibas) 
were collected 50 ft. (15 m.) above the ground in secondary growth. 
In domestication M. n. nemestrina's skillfulness as a climber has 
long been exploited for harvesting coconuts (Raffles, 1821, p. 243; 
Bertrand, 1967, p. 484) and recently also for collecting botanical 
specimens (Corner, 1946, p. 89). One observer (Be-b) in West 
Malaysia estimates that free-ranging M. n. nemestrina spends about 
15 per cent of its waking hours on the ground; another observer (R) 
in Kalimantan reports that 67 per cent of 40 contacts with M. n. 
nemestrina were with groups that were on the ground. 

In fleeing from danger, M. n. nemestrina habitually descends to 
the forest floor (Be-a, C, Mii, R), as described below: 

The usual response to humans was flight, and the troop typically 
descended to the ground and fled as a unit. In descending from the trees, 
animals leapt to the ground from as high as 10 meters and then succeeded in 
fleeing silently into good cover. The few successful efforts to track troops 
fleeing on the ground revealed the troop to be in a compact mass with 
individuals often shoulder to shoulder moving quietly and rapidly along the 
forest floor (Be-b). 



104 FIELDIANA: ZOOLOGY, VOLUME 67 

Fruits and seeds evidently are the principal food of this 
subspecies (Be-b, D, McC; Spiller and Rahlman, 1970. p. 496). 
Other reported natural foods are young leaves, leaf stems, fungi, 
insects, spiders, and lizards, including geckos (Be-b, Hrt/T; 
Harrison, 1961, p. 7). Under semi -natural conditions, M. n. 
nemestrina also is reported (Corner, 1941, p. 12) to eat the 
following: plant shoots, buds, flowers, rotan-palm hearts, plant sap, 
snails (except giant garden snails), slugs, caterpillars, maggots, 
insect nests, honey, frogs eggs, birds eggs, baby birds, baby squirrels; 
items apparently not eaten under these conditions are: poisonous 
mushrooms (which are distinguished from non -poisonous), gallfigs, 
earthworms, centipedes, millipedes, dragonflies, and cockroaches. In 
an experimental study of food preference in captive M. n. 
nemestrina, insect larvae and raw birds eggs were found to be 
highly acceptable, while live earthworms, raw meat, raw fish, and 
raw shellfish were completely rejected (Spiller and Rahlman, 1970, 
p. 496). In Sarawak M. n. nemestrina often raids rice fields (Hss/D), 
and in Sumatra it raids maize fields (Hg). 

Data concerning possible breeding seasonality are minimal. One 
author (Be-b) notes a concentration of births in the period 
November -March and mentions isolated births in other months. 
Another observer (McC) reports two females with suckling young in 
March and one female with a small infant in May, which is 
compatible with a birth peak in the period November -March. 

Observed contacts between neighboring troops of M. n. 
nemestrina reportedly are peaceful (Be-b). In West Malaysia 
peaceful contacts also have been reported with sympatric M. 
fascicularis, Presbytis obscurus, P. melalophus, and Hylobates lar 
(Be -a; also see Chivers, 1973, p. 122). However, in eastern 
Kalimantan, M. n. nemestrina and M. fascicularis were found to be 
highly segregated, with the former species occupying inland habitats 
and the latter restricted to the vicinity of a major river (R). In one 
reported contact with a giant squirrel, Ratufa affinis, a pigtail 
troop drove the squirrel out of a food tree (McC). 

Remarks. — Macaca nemestrina nucifera Sody, 1936 (p. 42) is 
based on the skull and part of the skin (head only) of a specimen 
collected at Pulau Bangka (between Sumatra and Borneo) by H. J. 
V. Sody (Coll. No. Bk 75, date unspecified). The present 
whereabouts of the type skull and skin fragment is unknown. 
Although some of Sody's specimens were deposited in the Museum 
Zoologicum Bogoriense and others in the Rijksmuseum van 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 105 

Natuurlijke Historie, Leiden, the type material of nucifera has not 
been found in either of these institutions (RMNH collection 
searched by Dr. A. M. Husson, Curator of Mammals, and myself in 
June, 1965; MZB collection searched by Dr. P. F. D. Van Peenen, 
U. S. Naval Medical Research Unit No. 2, Djakarta Detachment, in 
March 1971 and by me in August, 1973). 

According to Sody (1936, p. 42), the type of nucifera is an adult 
male, and the greatest length of the skull is 136.5 mm. (table 22). 
This would be well outside the range of cranial measurements in 
southern Sumatran adult male M. n. nemestrina (fig. 22) and 
comparable to those in adult male M. silenus, M. n. leonina, and M. 
n. pagensis (fig. 7). Pelage of the head of nucifera evidently is 
indistinguishable from that in Bornean and Sumatran M. n. 
nemestrina (Sody, 1936, p. 42). 

The only other known museum specimen of a pigtail collected 
in Bangka is an isolated skull (MZB 6501, adult 6\ fig. 27b), for 
which, unfortunately, the collector, date of collection, and exact 
locality of origin in Bangka are unknown. Judging from the 
abruptly truncated canines and unworn molars, the skull may be 
that of a captive animal. Greatest length of the skull is 157.5 mm., 
which is within the size range of M. n. nemestrina in southern 
Sumatra and southern Borneo (fig. 22). 

Although available evidence is not conclusive, M. n. nucifera 
Sody is now provisionally regarded as indistinguishable from M. n. 
nemestrina (Linnaeus). The missing type of nucifera may have been 
mis-sexed or incorrectly measured (but see table 22). Additional 
properly documented specimens of pigtails collected in Bangka are 
required for definitive resolution of this problem. 

Specimens examined. - Total 306. INDONESIA (Kaliman- 
tan): Badang, 4 (AMNH-AC); Bahau, 1 (skull only, MNHN); 
Banang, 1 (ZSBS); Djembajan, Sungai, 1 (USNM); Kapuas, Sungai, 
1 (skull in skin, RMNH); Karangan, Sungai, 2 (USNM); Kend- 
awangan, Sungai, 1, skull only (USNM); Landak, Sungai, 1 (skull 
only, USNM); Pamukan, Teluk, 1 (USNM); Peleben, 3 (2, AMNH- 
AC; 1, MZB); Pontianak, 1 (skin only, RMNH); Puruktjahu, 1 
(skin only, BM); Riam, 2 (AMNH-AC); Sekajam, Sungai, 2 
(USNM): Sintang, 1 (skull only, ZSBS); Sukadana, 1 (USNM); 
Tabang, 2 (MZB); Tandjung, 1 (skull only, mandible lacking, 
NHMB); southwestern Kalimantan, 5 (3 skins only, 2 skulls only, 
ZSBS); no locality, 1 (AMNH), 1 (NHMB), 1 (skull only, ZMB); 2 



106 FIELDIANA: ZOOLOGY, VOLUME 67 



TABLE 22. Cranial measurements (mm.) reported by Sody (1936, p. 42) 
for type-specimen of Ma caca nemestrina nucifera. 

Greatest length of skull J 36.5 

Basal length 98.5 

Postrostral length 90 

Palatal length 60 

Zygomatic breadth 89 

Braincase breadth 63.5 

Interorbital constriction 8 

Postorbital constriction 48 

Upper toothrow (P-M3) 44.5 

Canine, alveolar diameter 10.5 

Crown of upper M3 8.5 x 8.3 

Mandibular length 99.5 

Lower toothrow 50.5 



(1 skull only, MZB). INDONESIA (Sumatra); Aru, Teluk, 3 
(USNM); Babat, 2 (MZB); Bangka, 1 (skull only, MZB); 
Batangkuwis, 3 (1 skins only, ZSBS); Bengkulu, 2 (skulls only 
(RMNH); Brandan, 2 (skins only, ZMB); Deli, 1 (skeleton only, 
NMS); 1 (skin only, ZSBS); "Dolok- Oelve," 1 (skull only, NMS); 
Kajutanam, 2 (skulls only, MZB); Kateman, Sungai, 5 (1 skull only, 
USNM); Kluang, 3 (2 skulls only, MZB); Lampoengsche Distrikten, 
1 (MZB); Langsa, 1 (skull only, MZB); Loeboek Karet, 1 (MZB); 
Lubuklinggau, 1 (AMNH-AC); Medan vicinity, 8 (2 skins only, 
ZSBS); Muarabeliti, 2 (AMNH-AC); Ogan, Air, 3 (FMNH); Padang, 
1 (NMS), 3 (skulls in skins, RMNH), 1 (ZMUZ); Pagaralam, 2 
(skulls only, MZB); Pajakumbuh, 1 (skull and limb bones only, 
RMNH); Palembang, 2 (skulls only, NHMBe), 1 (RMNH); 
Palembang district, 1 (BM); Pasi, 1 (skull only, MZB); Pulau 
Munteh, 1 (MZB); Pulutelang, 1 (skeleton only, NHMB); Sanggul, 
Bukit, 8 (6, AMNH-AC; 2 MZB); Siak, Sungai, 2 (USNM); Sibolga, 
Teluk, 2 (USNM); Solok, 1 (RMNH); Tandjungmorawa, 1 (skull 
only, RMNH), 2 (ZSBS); Tarusan, Teluk, 2 (USNM); Tebingtinggi, 
3 (1 skin only, 1 skull only MZB); no locality, 2 (skulls only, BM), 1 
(FMNH), 2 (skins only, MNHN), 3 (1 skull in skin, RMNH), 7 (4 
skulls only, AMNH), 1 (MZB), 4 (2 skulls only, ZMB), 3 (skulls 
only, ZSBS). MALAYSIA, EAST: Abai, 10 (1 skin only, 4 skulls 
only, MCZ); Bareo, 1 (FMNH); Betotan, 2 (ZRCS); Entawa- 
Samarahan, 1 (BM); Kalulong, Bukit, 1 (BM); Kretam Kechil, 
Sungai, 1 (FMNH); Kuching, 3 (1 skull in skin, SMK) Labuan, 
Pulau, mainland opposite, 1 (skin only, BM); Lumu Lumu, 7 (5 
skulls only, MCZ); Madihit, Sungai, 1 (skull in skin, SMK); Papar, 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 107 

1 (USNM); Pa Umor, 1 (FMNH); Pulo Paku, 2 (BM); Ranau, 1 
(AMNH); Salai, Sungai, 4 (1, BM; 3, skins only, SMK); Sandakan, 
1 (skeleton only, MCZ); Sapagaya, Sungai, 1 (type of broca, 
USNM); Segama, Sungai, 1 (skin only, ZMB); Sidong, Gunong, 1 
(skin only, SMK); Sut, Sungai, 1 (FMNH); Tibas, Sungai, 2 
(FMNH). MALAYSIA, WEST: Bentong, 1 (BM); Hantu, Tanjong, 
(ZRCS); Jeram, 1 (ZRCS); Kelang, 1 (skin only, ZRCS); Kepong, 2 
(ZRCS); "?Kuala Lumpur," 1 (ZRCS); Kuala Lumpur, Cheras 
Road, 1 (skin only, ZRCS); Kuala Lumpur, Weld's Hill, 2 (ZRCS); 
Pinang, Pulau, 1 (BM), 2 (skulls only, NHMB), 1 (skin only, 
NHRM), 2 (skeletons only, USNM); Simpang Empat Rungkup, 1 
(BM); Sungai Renggam, 1 (BM). THAILAND: Chong, 1 (BM), 
Huai Lian, 2 (FMNH); Khao Si Kaeo, 1 (FMNH); Lam Phura, 
Sathani, 2 (ZRCS); "Trang," 1 (skull only, ZRCS). Imprecise 
localities: 9 (1 skin only, 1 skeleton only, 1 skull only, AMNH), 6 (3 
skins only, BM), 1 (FMNH), 8 (3 skins only, 5 skeletons only, 
IRSN), 10 (3 skins only, 3 skulls only, MNHN), 16 (skulls only, 
NHMBe), 7 (3 skins only, 3 skulls only, NMS), 26 (2 skulls in skins, 
6 skins only, 15 skulls only, RMNH), 3 (skins only, USNM), 18 (16 
skins only, ZMB), 5 (1 skin only, 1 skeleton only, 1 skull only, 
ZMUZ), 4 (2 skulls only, ZRCS), 3 (skulls only, ZSBS), 16 (9 skins 
only, 5 skulls only, 2 skeletons only, ZSI). 

Macaca nemestrina leonina (Blyth, 1863) 

Another species of Cercopithecus: Heifer, 1838, p. 858 - reported to inhabit 
limestone buttes in northern Tenasserim. 

Macacus nemestrinus (?): Blyth, 1844, p. 473 — external characters of specimens 
collected in Arakan district, Burma, by A. P. Phayre; probable distribution, 
Arakan, Tenasserim. 

Macacus nemestrinus: Morice, 1875, p. 41 — recorded in South Vietnam. 

Macaca nemestrina sub. sp.: McCann, 1933b, p. 808 - external characters; field 
observations. 

Macaca nemestrina: Yin, 1954, p. 269 - sight record in Burma. Fooden, 1971a, p. 
32, pi. 5a (head), pi. 5b (animal) — report on specimens collected in Thailand; 
possibly specifically distinct from Sundaic M. nemestrina; nasal deformity in 
adult female. 

I[nuus] arctoides (?. Is. Geoff.): Blyth, 1847, p. 731 - revised identification of 
specimens earlier identified as Macacus nemestrinus (?): Blyth, 1844. 

Inuus pileolatus Tickell, 1854-1863, MS., pp. 109, 111 - unavailable manuscript 
name based on captive immature male (pi. 23) collected by R. S. Tickell in forest 
near Ye, Burma, Nov. 1854; subsequently regarded as a synonym of Silenus 
nemestrinus (Tickell, 1863-1875, MS., p. 139). 



108 FIELDIANA: ZOOLOGY, VOLUME 67 

I[nuus] leoninus Blyth, 1863, p. 7 — new species based on specimens previously 
identified as Macacus nemestrinus (?): Blyth (1844, p. 473) and Inuus arctoides: 
Blyth (1847, p. 731); specific name is an adjective, presumably alluding to the 
lionlike mane and cheek ruff; subsequently considered identical with Inuus 
nemestrinus (Blyth, 1863, appendix, p. [189] ). Blyth, 1875, p. 4-redescription of 
syntypes; zoogeography. 

Macacus leoninus Sclater, [1871], p. 663, pi. 35 (animals) — taxonomic history, 
Anderson, 1879, p. 52, figs. 1, 2 (skull of type of M. andamanensis Bartlett, 1869) 
— external characters; cranial characters and measurements; osteology; 
relationships; distribution, southern Arakan, Irrawaddy Valley. Anderson, 1881, 
p. 71 — part (also includes reference to misidentified female specimen of Macaca 
assamensis collected near Bhamo); redescription of holotype. Sclater, 1898, p. 
280 - first record in Thailand. Elliot, [1913], pp. 208, 209 - incorrectly rejected 
as a junior homonym of Simia leonina Shaw, 1800, which is a langur. not a 
macaque. 

Macacus coininus: Kloss, 1903, pp. 322, 325 — lapsus for Macacus leoninus. 

M[acaca] leonina: Wroughton, 1918, p. 555 — external characters; type-locality, 
Arakan. Kloss, 1919, p. 344 — redescription of holotype; regarded as a synonym 
of Macacus andamanensis Bartlett, 1869. 

Macacus nemestrinus, var. leonina: Schlegel, 1876, p. 356 — specimen received from 
"?Arracan." 

Macaca nemestrina leonina: Pocock, 1939, p. 59 — external and cranial characters; 
taxonomy; field observations; distribution, upper Burma to Tenasserim, Mergui 
Archipelago, Thailand. Fiedler, 1956, p. 177 — external characters. J. R. Napier 
and P. H. Napier, 1967, p. 403 - listed. 

Pithecus leoninus: Wroughton, 1916a, p. 296 — external characters; taxonomic 
comparisons. 

Nemestrinus leoninus: Heck, 1916, p. 551 — external characters; taxonomic 
comparisons; facial expressions of captive; distribution, Arakan, Irrawaddy 
region, upper Burma, possibly Thailand. 

Rhesus (Nemestrinus) leoninus: De Beaux, 1923, p. 34 — external and cranial 
characters. 

Macacus andamanensis Bartlett, 1869, p. 57, fig. (animal) — holotype B.M. 
71.5.19.2 (skin, skull), subadult female obtained alive in 1864 by Capt. T. A. 
Brown, R.N., at Port Blair, South Andaman Island, presented to Menagerie of 
Zoological Society of London on July 8, 1869. Sclater. [1870], p. 467, fig. (captive 
holotype) — type history. Hamilton, 1870, p. 220 — holotype said to have been 
transported to Andaman Islands from Burma. Sclater, [1871], p. 663 — a 
synonym of Inuus leonina Blyth. 

Pithecus andamanensis: Elliot, [1913], p. 208, pi. 2 (animals ex Sclater, 1870) - 
redescription of holotype of Inuus Lioninus Blyth, 1863; distribution, Arakan, 
Irrawaddy Valley, possibly Thailand, introduced in Andaman Islands. 

Macaca andamanensis: Kloss, 1916a, p. 30 — external and cranial characters; 
taxonomic comparisons; nomenclature. Kloss, 1916b, p. 2 — external and cranial 
characters; taxonomic comparisons. 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 109 

[Macaca nemestrina] andamanensis: Kloss, 1919b, p. 345 — key to external 
characters. Pocock, 1931, p. 302 — external characters and variation; cranial 
characters; distributions, "Arakan and Upper Burma to Tenasserim, including 
the Mergui Archipelago, and the adjoining districts of Siam." 

Macaca nemestrina andamanensis: Pocock, 1939, p. 60 — junior synonym of 
Macaca nemestrina leonina: Pocock, 1939. 

Macaca adusta Miller, 1906, p. 559, pis. 13-17 (skin and skull of holotype) — 
external and cranial characters; holotype, adult male skin and skull (USNM 
124023) collected at Champang, Tenasserim Div., Burma, by W. L. Abbott, Dec. 
22, 1903; distribution: "Malay Peninsula. Limits of range unknown." Pocock, 
1931, p. 303 — a synonym of Macacus leoninus Blyth, 1863. 

Macaca nemestrina adusta: Kloss, 1908, p. 147 — included in provisional list of 
mammals of southern Malay Peninsula. Gyldenstolpe, 1919, p. 130 — reported in 
peninsular Thailand. Kloss, 1919b, p. 343 — key to external characters; 
distribution, southern Tenasserim. 

Pithecus adusta: Elliot, [1913], p. 206 — redescription of holotype; zoogeography; 
distribution, Tenasserim. Robinson and Kloss, 1914a, p. 392 — doubtfully 
distinguishable from Sumatran Macaca nemestrina. 

Macaca insulana Miller, 1906, p. 560 — external and cranial characters; holotype, 
adult male, skin and skull (USNM 104441, Coll. No. 199) collected at Chance 
Island ( = Ko Chan), Mergui Archipelago, Thailand, by W. L. Abbott, Jan. 1, 
1900; distribution, Ko Chan. Pocock, 1931, p. 303 — a synonym of Macacus 
leoninus Blyth, 1863. 

Pithecus insulanus: Elliot, [1913], p. 207 — redescription of type; zoogeography. 

Macaca nemestrina indochinensis Kloss, 1919b, p. 343 — external and cranial 
characters; holotype, adult male, skin and skull (USNM 236623 Coll. No. 2148) 
collected at Lat Bua Khao, southeastern Thailand, by C. B. Kloss, Oct. 12, 1916. 
Pocock, 1931, p. 303 — a synonym of Macacus leoninus Blyth, 1863. Chasen, 
1935, p. 38 — external characters; taxonomic comparisons. 

[Macaca nemestrina] blythii Pocock, 1931, p. 305 — external and cranial 
characters; holotype, adult male, skin and skull (BM 72.11.18.2; Orig. No. 
1552B), captive specimen of unknown origin, originally identified as Macacus 
leoninus (Sclater, [1871], p. 663, pi. 35), acquired by British Museum (Natural 
History) in 1872; distribution unknown. 

Macaca nemestrina blythii: Pocock, 1939, p. 63, fig. 15 (animal ex Sclater, [1871]) 

— external and cranial characters of holotype; type history; hypothetical 
distribution, "?Naga Hills in Assam." J. R. Napier and P. H. Napier, 1967, p. 403 

— listed. 

Macaca fascicularis subsp.: Delacour, 1940, p. 24 — said to be common in vicinity 
of Hue and Sai Gon, South Vietnam. 

Type. - Adult male, skin only (ZSI 11823/I.M. 43a), preserved 
in the National Zoological Collection, Zoological Survey of India, 
Calcutta; collected by A. P. Phayre, 1844. The type-series originally 
also included an immature specimen, which was also collected by 



1 10 FIELDI ANA: ZOOLOGY, VOLUME 67 

Phayre (Blyth, 1863, p. 8), but the whereabouts of this syntype is 
now unknown. 

Type -locality. — "Mountainous and rocky situation," Arakan 
district, southeastern Burma (Blyth, 1844, p. 473). 

Distribution (fig. 28). — Indochinese Peninsula, from eastern 
Assam (India), extending southeastward through Burma, south- 
western Yunnan (China), Thailand, southern Laos, South Vietnam, 
Cambodia (inferred), south to Surat Thani-Krabi depression in 
peninsular Thailand (8-9°N); known altitudinal range about 75- 
1250 m. (table 11). The lack of records of M . n. leonina in central 
Burma, an area reasonably well collected by the Bombay Natural 
History Society's Mammal Survey (Ryley, 1914, p. 710; Wroughton, 
1915a, p. 460; 1916a, p. 291; 1916b, p. 758; 1921, p. 549; Fry, 1928, p. 
545; 1929, p. 636), suggests that this area may constitute a natural 
hiatus within the range of the subspecies; Tickell (1854-1863, MS., 
p. 112) likewise indicates that this subspecies is rare or absent in 
central Burma. Known insular records of M. n. leonina are 
restricted to Lanbi Kyun, Ko Chan, Ko Phuket, and Ko Yao Yai, 
all off the west coast of peninsular Burma and Thailand. 

Two erroneous locality records of M. n. leonina have been 
published by Anderson (1881). A female specimen (ZSI 11909/I.M. 
43c) collected 20-25 miles below Bhamo in northeastern Burma was 
referred to M. n. leonina (Anderson, 1881, p. 72) after having been 
originally identified as M. assamensis (Anderson, 1879, p. 65); the 
correct identification of this specimen is M. assamensis, as 
previously indicated by Khajuria (1954, p. 115). Anderson (1881, p. 
72) also records a male specimen of M. n. leonina (ZSI 
11926/I.M.43e) that was presented to the Indian Museum by O. L. 
Fraser, Jan. 6, 1878, and was said to have originated in "Perak," 
which presumably refers to the state of that name in West 

Fig. 28 - continued: 

15. Thakhek. 16. Savannahket. 17. Pakse; Plateau des Bolovens. 18. Saravane. 19. 
Hue. 20. Kawkareik. 21. Ban Pong Nam Ron. 22. Khlung. Khlong; Ko Keow. 23. 
Samnak Rabam. 24. Hin Lap, Sathani. 26. Pak Chong, Sathani. 26. Lat Bua Khao, 
Sathani. 27. Phu Kheo Forest Reserve. 28. Ye. 29. Ban Kerng Chada. 30. 
Chongkrong. 31. Ban Huai Maenam Noi. 32. Paungdaw power station. 33. Rat Buri. 
34. Phet Buri. 36. Ban Nong Kho. 36. Chantaburi; Sa Bap, Khao. 37. Ban Huang 
Som. 38. Tay Ninh. 39. Trang Bom. 40. Sai Gon. 41. Da Ban. 42. Lanbi Kyun. 43. 
Red Point. 44. Bankachon; Champang; Telok Besar. 45. Chan, Ko. 46. Ban Khlong 
Wan; Ban Tha San. 47. Klong Tung Sai; Telok Poh. 48. Ban Nong Kok. 

Negative Reports: A. Andaman Islands. B. Barren Island. C. Nicobar 
Islands, northern. D. Nicobar Islands, southern. E. Klet Kaeo, Ko. 




600 MILES 



KILOMETtBS 



Fig. 28. Locality records of M. n. leonina; closed squares = specimens examined; 
open squares = literature records; circles = marginal records of M. n. nemestrina; 
letters and heavy broken lines indicate negative reports. For details, see "Gazetteer of 
Collecting Localities." 

Key to localities: 1. Changchang Pani. 2. Singkaling Hkamti. 3. Pidaung Game 
Sanctuary. 4. Arakan. 5. Sittwe. 6. Taho. 7. Mong Nai. 8. Meng-hai. 9. Khun Tan. 
10. Ban Hue Horn. 11. Meh Lem river. 12. Muang Liap. 13. Vientiane. 14. Paksane. 



Ill 



112 FIELDIANA: ZOOLOGY, VOLUME 67 

Malaysia. However, this state is more than 300 km. beyond the 
apparent range of M. n. leonina (fig. 28). The tail of this specimen is 
mutilated and the fur seems to have been worn off in life, 
suggesting that the monkey was a captive; the Malaysian locality is 
undoubtedly spurious (Blanford, 1888b, p. 20; cf. Chasen, 1940a, p. 
65). 

According to Dollman (1932, p. 9), Lord Cranbrook supposedly 
collected a pigtail macaque at Nam Tamai, a tributary of the upper 
Irrawady River in northernmost Burma. However, this report 
probably is based on misidentification of a juvenile male specimen 
of M. assamensis (BM 32.11.1.2; Pocock, 1939, p. 54) obtained by 
Lord Cranbrook on April 8, 1931 at Adung Valley, which is another 
name for the same tributary of the Irrawady River (Cranbrook in 
Kinnear, 1934, p. 348). 

External characters (fig. 29). — Head and body length 400-490 
mm. in 15 adult females (excluding improbable measurement of 555 
mm. recorded for BNHS 5068, Loc. No. 1; McCann, 1933b, p. 809), 
500-595 mm. in 13 adult males; relative tail length (T/HB) 0.28-0.45 
in 14 adult females, 0.30-0.48 in 13 adult males (fig. 2); weight 4.4- 
5.7 kg. in 7 adult females, 6.2-9.1 kg. in 6 adult males (table 23). 
Dorsal pelage yellowish-brown agouti to golden-brown agouti 
anteriorly, becoming slightly drabber posteriorly, with an indistinct 
median dark streak (about 2 cm. broad) on the lumbosacral region; 
crown short-haired, usually somewhat darker golden-brown than 
back; anterior part of crown patch narrow, extending laterally 
approximately as far as middle of each eye, sharply defined 
anterolateral^ by contrast with pale ochraceous-buff lateral 
supraorbital patches; side whiskers long, pale ochraceous-buff, 
forming a well-defined facial ruff that tends to conceal the ears 
from anterior view; skin of muzzle pale brownish, densely covered 
with short buffy hairs; outer surface of arms above elbows colored 
approximately like back, outer surface of forearms, hands, thighs, 
shanks and feet slightly paler and drabber; tail thin, bicolor, 
sharply defined blackish dorsally, pale ochraceous-buff ventrally; 
base of tail set off by a pair of dorsolateral^ projecting tufts of 
whitish fur; throat, chest and anterior surface of shoulders pale 
ochraceous-buff; underparts thinly haired pale yellowish-brown to 
whitish. (See Addenda, p. 168.) 

In adult males hairs on the scapular region are 7-10 cm. long, 
the distal two-thirds of each of these hairs is conspicuously 
annulated with about 15 alternating bands (each 3-4 mm. broad) of 






Fig. 29. Macaca nemestrina leonina, external characters. (Photos by J. Fooden 
at Calcutta Zoo) 



113 



114 FIELDIANA: ZOOLOGY, VOLUME 67 

TABLE 23. Weight in wild-collected adult specimens of M. n. leonina. 

Locality Weight (kg.) 

Nos. 1 Females Males 

21.23 4.71,5.48 8.5 

29. 31 4.40,4.65,4.96 8.1 

44 4.6,5.7 8.2,9.1 

45 - 6.2, 6.2 
Means 4.9 7.7 

1 See map. Figure 28. 

yellowish or golden and dark brown, and the basal third of each 
hair is pale ochraceous-buff. Laterally on the lumbosacral region, 
where individual hairs are 3-4 cm. long, the number of pale and 
dark annulations is reduced to 3-5, and the pale ochraceous-buff 
unannulated basal portion is about 1 cm. long. Medially on the 
lumbosacral region, individual hairs in the faintly defined median 
dark streak lack the pale basal portion and are about 1 cm. shorter 
than hairs lateral to the streak. In two living adult male specimens 
examined (National Zoological Park, Washington, D.C.; Calcutta 
Zoological Gardens), the skin of the perineal area is marked with a 
clearly defined red streak that extends ventrally along the median 
raphe from the anus to the prepuce, both of which are red. 

Hairs on the crown are about 1.5 cm. long posteriorly and 0.5 
cm. long anteriorly and form a whorl radiating from the vertex. On 
the midfrontal region the anteriorly directed golden-brown crown 
hairs meet the posteriorly directed pale ochraceous-buff hairs of the 
lateral supraorbital region to form a low but sharply defined wedge- 
shaped crest that marks the anterior border of the crown patch. 
The pale side whiskers in front of the ears are about 4 cm. long. The 
skin around the eyes is sharply defined pale bluish white. In most 
adult males a pair of clearly defined narrow reddish streaks extend 
laterally from the outer corner of each eye to the side whiskers 
(Tickell, [1863]-1875, MS., p. 139; Sclater, [1871], colored pi. 35, 
republished in Elliot, [1913], colored pi. 2). Brown (1896, p. 485) 
reports that the red color of these eye streaks became intensified 
during excitement in one young captive specimen. These streaks 
seem to be absent in an adult male observed at the National 
Zoological Park, Washington, D. C. 

In adult females the pelage is generally similar to that in 
males, but the fur is somewhat shorter and averages somewhat 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 115 

paler and drabber — pale yellowish-brown instead of golden-brown. 
Although the anterior margin of the crown patch is sharply defined 
in females, as in males, there usually is little contrast between the 
color of the crown and that of the upper back in females. The 
sparse hairs on the ventral surface of adult females are whitish 
instead of pale yellowish-brown agouti as in males. Immatures are 
generally paler and drabber than adult females, and the agouti 
pattern tends to be less developed in immatures. 

A series of three small adult male specimens collected at Ko 
Chan (Loc. No. 45), Mergui Archipelago, is the basis of Macaca 
insulana Miller, 1906; this may be an instance of insular dwarfism 
(fig. 22). Two adult females (BNHM 5068, 5069) collected at 
Changchang Pani are somewhat darker golden-brown than a third 
adult female (AMNH 83434) collected at the same place and also 
darker than most other specimens examined of M. n. leonina. 

Cranial characters (figs. 3, 5). — Skull relatively small, greatest 
length excluding incisors 103.0-121.1 mm. in 19 adult females, 124.1- 
140.1 mm. in 17 adult males (fig. 2); rostrum short (fig. 8); 
zygomatic arches broad (fig. 7); supraorbital ridges thick and 
prominent; anterior surface of malar flat to weakly convex. In 
specimens examined, skull length in northern populations of M. n. 
leonina (north of about 11°N) averages slightly greater than in 
southern populations (fig. 22). (See Addenda, p. 168.) 

Habits and habitats. — This account is based on field 
observations published by the following authors, who are cited by 
means of indicated abbreviations: 

F Fooden, 1971a, p. 32 

H Heifer. 183a p. 858 

McC McCann, 1933b. p. 808 

P/B Phayre in Blyth, 1844, p. 473 

S/W Shortridge in Wroughton, 1915b, p. 700 

T Tickell, 1854-186a MS., p. 112 

Information in these published reports is supplemented by brief 
notes on field tags of two museum specimens, one collected by M. 
Pierre in 1874 (MNHN 1878/1126, ad. $, Chamchay, South 
Vietnam) and the other collected by W. L. Abbott in 1903 (USNM 
124022, ad. 3, Champang, Burma). 

All known specimens of this subspecies have been collected in 
the foothills of Southeast Asian mountain ranges (fig. 28; H, P/B, 
T) at elevations ranging from about 75 m. (Ban Huai Maenam Noi, 



116 FIELDIANA: ZOOLOGY, VOLUME 67 

Thailand) to 1200-1300 m. (Taho, Burma) (table 11). All but one 
troop for which type of forest is known were observed in dense 
evergreen forest (F, McC, T); the exceptional troop was observed in 
deciduous dipterocarp forest (F). Within its range this species is 
fairly abundant, although not as common as gibbons and langurs 
(F, S/W, T). 

Reported troop size in eight troops is: 12, 15, 20, 20, 20, 20 + , 
30 + , 40 (F). Of two solitary males that have been collected (F; 
Pierre specimen), one is a subadult and the other an adult. 

M. n. leonina is almost exclusively arboreal (F, McC). However, 
in at least part of the range, troops are known to descend to the 
ground in order to raid rice fields (S/W). When frightened, this 
macaque flees into the canopy (F, McC). Although M. n. leonina is 
naturally more arboreal than M. n. nemestrina, this subspecies does 
not seem to be domesticated and used as a trained coconut 
harvester, as is frequently done with M. n. nemestrina (see above). 

The diet of M. n. leonina consists predominantly of fruit (F, 
McC), although leaves (McC), caterpillars and adult insects (F), and 
rice (S/W) also are eaten, at least occasionally. 

Information on breeding seasonality in M. n. leonina is scanty. 
A tumescent female, evidently in prime breeding condition, was 
collected in September (Abbott specimen). Pregnant females 
collected in February and in April apparently would have given 
birth in April-May (McC) or June-July (F). Young infants collected 
in February appeared to have been born in the preceding December 
(F). These meager data may indicate that there are semi-annual 
birth peaks in this subspecies, approximately in June and 
December. 

There are no known reports of encounters between M. n. 
leonina and other species of primates, although species of gibbons 
and langurs inhabit the same forests and probably pass through the 
same trees (F). M. arctoides also inhabits the same forests, but 
encounters with M. n. leonina presumably are infrequent since M. 
arctoides is primarily terrestrial and M. n. leonina is primarily 
arboreal (F, McC). M. assamensis, which seems to be very similar to 
M. n. leonina in habits and ecological requirements (F), is almost 
completely allopatric in geographic distribution; known areas of 
overlap of M. n. leonina and M. assamensis are restricted to a 
narrow zone in west-central Thailand (F) and another in the 
Assam-Burma border area (Changchang Pani, Singkaling Hkamti). 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 117 

M. fascicularis is broadly sympatric with M. n. leonina but seems 
to be ecologically segregated by its preference for lower elevations 
and non-evergreen habitats (F). 

Remarks. — M. nemestrina blythii Pocock, 1931, is now 
regarded as inseparable from M. n. leonina Blyth, 1863. This is in 
agreement with the determinations of Blyth (in Sclater, [1871], p. 
664) and Anderson (1879, p. 52), but contrary to the opinion of 
Pocock (1931, p. 304; 1939, p. 62), who is followed by authors of 
recent checklists. 

The type and only known specimen of blythii is the skin and 
skull (BM 72.11.18.2) of an adult male pigtail without locality data 
that was purchased by the Menagerie of the Zoological Society of 
London from an animal dealer on June 14, 1870 (Sclater, [1871], p. 
663; 1883, p. 19). Within three or four months of acquisition, the 
living captive was examined by Blyth (in Sclater, [1871], p. 664), 
who identified it as Macacus leoninus (=M. n. leonina), and it also 
was depicted in an excellent colored illustration by the animal 
artist J. Wolf (in Sclater, [1871], pi. 35; republished in Elliot, [1913], 
vol. 2, colored pi. 2). This illustration clearly shows the V-shaped 
crown patch, the red lateral orbital streaks and the anteriorly 
directed tail arch that are characteristic of M. n. leonina. After 
about 2V2 years of captivity in the London Zoo, the monkey died 
and its skin and skull were acquired by the British Museum 
(Natural History), probably in mid-November, 1872, judging from 
the catalog number. 

The stuffed skin is in relatively poor condition with much of 
the fur missing from the sacral and buttock regions. What fur 
remains generally resembles that in M. n. leonina, but is much 
darker and drabber — brown to golden-brown agouti dorsally, 
instead of pale golden-brown agouti. The skull is indistinguishable 
from that in adult male M. n. leonina (Anderson, 1879, figs. 1,2; 
Pocock, 1931, p. 306; 1939, p. 65); a transverse eminence on the left 
parietal lateral to the temporal ridge appears to be the result of an 
old healed fracture. 

The dark pelage color at the type-specimen of blythii is the sole 
basis for Pocock's (1931, 1939) opinion that it represents a form 
distinct from M. n. leonina; Pocock's (1939, p. 65) unfounded 
speculations concerning the hypothetical geographic origin of the 
specimen are irrelevant. In view of the otherwise near-perfect 
identity of characters of the museum specimen with those of M. n. 



118 FIELDIANA: ZOOLOGY, VOLUME 67 

leonina, and in view of Blyth's identification of the living animal as 
M. n. leonina, the significance of the dark coloration is suspect. 
This coloration may be an individual variable, it may be a result of 
conditions of captivity, or it may be an artifact of past conditions of 
storage; a captive specimen of M. n. nemestrina (BM 42.2.20.7) is 
similarly darkened as compared with wild-collected specimens of 
that subspecies. In any event, the ambiguous coat color evidence of 
this tattered zoo specimen of unknown origin does not appear 
adequate to warrant continued recognition of blythii as a valid 
taxonomic entity. 

Specimens examined. — Total 89. BURMA: Arakan, 2 (1 skull 
only, ZMB; 1 skin only, type of leonina, ZSI); Bankachon, 8 (3 
skins only, 1 skull only, BM; 3, BNHS; 1 skin only, ZSI); 
Champang, 2 (including type of adusta, USNM); Lanbi Kyun, 1 
(BM); Paungdaw, 1 (BM); Red Point, 1 (USNM); Singkaling 
Hkamti, 1 (BM); Telok Besar, 2 (1 skull only, USNM); no locality, 
1 (ZMB). INDIA: "Assam," 1 (IRSN); Changchang Pani, 4 (2, 
AMNH; 2, BNHS); "Ross Island, Andaman Islands," 1 (type of 
andamanensis, BM). LAOS: Muang Liap, 1 (ZRCS); Plateau des 
Bolovens, 4 (skins only, AMNH). SOUTH VIETNAM: Chamchay, 
1 (skull only, MNHN); Da Ban, 1 (ZRCS); Trang Bom, 3 (2, 
USNM; 1, ZRCS). THAILAND: Ban Huai Maenam Noi, 7 (1, 
CTNRC; 6, FMNH); Ban Huang Som, 1 (BM); Ban Kerng Chada, 
4 (FMNH); Ban Khlong Wan, 1 (ZRCS); Ban Nong Kho, 2 
(USNM); Ban Nong Kok, 1 (skin only, ZRCS); Ban Pong Nam 
Ron, 7 (1 fetus, FMNH); Ban Tha San, 2 (ZRCS); Chan, Ko, 3 
(including type of insulana, USNM); Chantaburi, 2 (ZRCS); 
Chongkrong, 1 (FMNH); Hin Lap, 1 (USNM); Khlung, Khlong, 1 
(FMNH); Khun Tan, 1 (NHRM); Klong Tung Sai, 4 (1, BM; 3, 
ZRCS); Ko Keow, 1 (FMNH); Lat Bua Khao, 1 (type of 
indochinensis, USNM); Pak Chong, 2 (1, BM; 1, USNM); Sa Bap, 
Khao, 1 (USNM): Samnak Rabam, 2 (1, CTNRC; 1, FMNH); Telok 
Poh, 1 (BM); no locality, 1 (skin only, AMNH). Imprecise localities: 
1 (AMNH), 1 (type of blythii, BM), 1 (skull in skin, MNHN), 2 
(skins only, RMNH); 2 (skins only, ZSI). 

Macaca nemestrina pagensis Miller, 1903b 

Macacus nemestrinus: Thomas, 1895, pp. 661, 664 — report of specimen collected at 
Sioban, Pulau Sipura; said to be possibly introduced. 

Macaca nemestrina: Medway, 1970, pp. 527, 528 - part; distribution, Pagi Islands, 
said to be possibly introduced. 



FOODEN: LIONTAIL & PIGTAIL MACAQUES 119 

Macacus pagensis Miller, 1903b, p. 61, pis. 11-13 (skull of holotype) — external and 
cranial characters; distribution, Pulau Pagi Selatan; specific name is adjective 
derived from name of locality. 

Macaco pagensis: Miller, 1906, p. 557, pis. 18-20 (skull of holotype) - redescription 
of type; distribution, "probably confined to the Pagi Islands." 

Macaca nemestrina pagensis: Chasen, 1940a, p. 66 — distribution, Pagi Islands. J. 
R. Napier and P. H. Napier, 1967, p. 403 - listed. 

Pithecus pagensis: Elliot, [1913], p. 200 — external and cranial characters of 
holotype. 

Rhesus nemestrina mentaveensis DeBeaux, 1923, p. 37 — provisionally proposed 
subspecies; external and cranial characters; holotype, juvenile male, skin and 
skull in collection of Museo Civico di Storia Naturale "Giacomo Doria," Genova, 
collected at Sioban, Pulau Sipura, by E. Modigliani, 1894. 

Type. — The holotype is an adult female skin (fig. 30) and skull 
preserved in the United States National Museum (USNM 121653, 
Coll. No. 2053), collected by W. L. Abbott, November 17, 1902 
(Miller, 1903b, p. 61, pi. 11-13, skull). A juvenile paratype, offspring 
of the holotype (Miller, 1903b, p. 63), cannot now be located. 

Type locality. — South Pagi Island ( = Pulau Pagai Selatan), 
southwest of Sumatra, Indonesia. 

Distribution (fig. 25). — Known only from Pulau Sipura, Pulau 
Pagi Utara (North Pagi Island), and Pulau Pagai Selatan (South 
Pagi Island), all in Kepulauan Mentawei (Mentawi Islands), 
southwest of Sumatra. The range probably also includes Pulau 
Siberut, which is the largest and northernmost island in the 
Kepulauan Mentawei chain and which has a mammalian fauna 
generally similar to that in the three southern islands (Chasen and 
Kloss, 1928, pp. 808-840). (See Addenda, p. 168.) 

External characters (figs. 30, 31). — Head and body length 435- 
456 mm. in 3 adult females, 530 mm. in 1 adult male (fig. 22); 
relative tail length (T/HB) 0.25-0.33 in 3 adult females, 0.30 in 1 
adult male (fig. 2); weight 4.5 kg. in 1 adult female (USNM 121653). 
Dorsal surface chocolate-brown non-agouti, becoming pale brown 
non-agouti on flanks; crown golden-brown, slightly paler than back; 
side whiskers and beard short (about 1 cm. long), dark brown; skin 
of muzzle brownish, thinly covered with inconspicuous short buffy 
hairs; throat, sides of neck and anterior surface of shoulders clearly 
defined pale ochraceous-buff; outer surface of upper arm (deltoid 
region) drab brown, similar to adjacent region of back; outer surface 
of elbows, forearms and hands contrastingly pale reddish-brown; 
outer surface of legs chocolate-brown proximally, becoming pale 



120 



FIELDIANA: ZOOLOGY, VOLUME 67 




>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; ll o 12'-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'-77 20'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'-H 58'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; 3 41'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'-4 o 20'N, 98 o 00-98°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'-4 o 00'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, 100 o 21'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). 

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, 115 31'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'-106 o 50'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. Menden, Jan. 24, 1935 
(AMNH-AC). 93 

Pagai Utara, Pulau, sea level; INDONESIA: Sumatra; ca. 2°42'S, 100°07'E; 
collected by J. J. Menden, Jan. 13-26, 1935 (AMNH-AC). 92 

Pagi Islands. See Pagai Selatan, Pulau, and Pagai Utara, Pulau. 

Sioban; INDONESIA: Sumatra; 2°11'S, ^^E; collected by E. Modigliani, 1883 
(Museo Civico di Storia Naturale "Giacomo Doria," Genova (Thomas, 1895, p. 664; 
DeBeaux, 1923, p. 37]). 91 

Sipora. See Sioban. 

Sipura, Pulau. See Sioban. 

South Pagi. See Pagai Selatan, Pulau. 



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ADDENDA 

1. While this monograph was in press, one additional adult 
male specimen of Macaca nemestrina leonina (No. 20039) collected 
at Khao Soi Dao, Thailand (see below) was examined at the 
Academy of Natural Sciences, Philadelphia. 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. 



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