ILli,.^.- LIBRARY
AT UR3ANA-CHAMPAIGM
-
ft
JjlL,l AJ.-UW jQ^A.^
W
FIELDIANA
Zoology
Published by Field Museum of Natural History
Volume 67
TAXONOMY AND EVOLUTION OF
LIONTAIL AND PIGTAIL MACAQUES
(PRIMATES: CERCOPITHECIDAE)
JACK FOODEN
WML HISTORY S8WH
j AN -2 9 '^76
LIBRARY
November 25, 1975
FIELDIANA: ZOOLOGY
A Continuation of the
ZOOLOGICAL SERIES
of
FIELD MUSEUM OF NATURAL HISTORY
VOLUME 67
'&
FIELD MUSEUM OF NATURAL HISTORY
CHICAGO, U.S.A.
TAXONOMY AND EVOLUTION OF
LIONTAIL AND PIGTAIL MACAQUES
(PRIMATES: CERCOPITHECIDAE)
FIELDIANA
Zoology
Published by Field Museum of Natural History
Volume 67
TAXONOMY AND EVOLUTION OF
LIONTAIL AND PIGTAIL MACAQUES
(PRIMATES: CERCOPITHECIDAE)
JACK FOODEN
Research Associate
Field Museum of Natural History
and
Professor of Zoology
Chicago State University
November 25, 1975
Publication 1216
Patricia M. Williams
Managing Editor. Scientific Publications
Library of Congress Catalog Card Number: 753914
US ISSN 0015-0754
PRINTED IN THE UNITED STATES OF AMERICA
Dedicated to the memory of Kitti Thonglongya (1930-1974), zoologist, friend,
colleague and field companion during 1973 collecting trip to peninsular Thailand.
CONTENTS
PAGE
Introduction l
Generic names applied to liontail and pigtail macaques 4
key to external characters 6
Cranial comparisons 7
Tail-proportions, pelage, vertebrae, carriage 18
Female genitalia 28
Glans penis and baculum 33
Blood proteins 45
Comparative ecology and behavior 51
Hybridization 54
Fossil record 60
Evolution and dispersal 66
Species accounts 73
Macaca silenus (Linnaeus, 1758)
Synonymy 73
Type 75
Type-locality 77
Distribution 78
External characters 78
Cranial characters 78
Habits and habitats 79
Specimens examined 84
Macaca nemestrina (Linnaeus, 1766)
Taxonomy 85
Macaca nemestrina nemestrina (Linnaeus, 1766)
Synonymy 91
Type 93
Type-locality 95
Distribution 95
External characters 95
Cranial characters 100
Habits and habitats 100
Remarks 104
Specimens examined 105
Macaca nemestrina leonina (Blyth, 1863)
Synonymy 107
Type 109
Type-locality 110
Distribution 110
External characters 112
I
II
Cranial characters 115
Habits and habitats 115
Remarks 117
Specimens examined 118
Macaca nemestrina pagensis Miller, 1903
Synonymy 118
Type 119
Type-locality 119
Distribution 119
External characters 119
Cranial characters 120
Habits and habitats 121
Specimens examined 121
Gazetteer of collecting localities 122
Macaca silenus 123
Macaca nemestrina nemestrina 125
Macaca nemestrina leonina 137
Macaca nemestrina pagensis 142
References 143
Addenda 168
INTRODUCTION
Liontail macaques inhabit southwestern India, and three well-
defined forms of pigtail macaques inhabit Southeast Asia from
eastern Assam to Borneo (fig. 1). No two of these macaques are
sympatric. The geographic ranges of liontail macaques and pigtail
macaques are separated by a 2000 km. gap; the ranges of recognized
forms of pigtail macaques are adjacent but not overlapping. Current
taxonomic practice, following Kloss (1919b, p. 138), is to regard
liontails as one species (Macaca silenus Linnaeus, 1758) and various
forms of pigtail macaques as subspecies of another species (M.
nemestrina Linnaeus, 1766). Although available evidence is not as
conclusive as generally assumed (see below), this usage is retained
in the present revision.
Liontails (M. silenus) and Sundaic pigtails (M. nemestrina
nemestrina), both described in the eighteenth century, are the best
known members of this group. Indochinese pigtails (M. n. leonina),
described in 1863, are less widely known and are frequently
confused with M. n. nemestrina. Kepulauan Mentawai pigtails (M.
n. pagensis), described in 1903 and represented in collections by a
total of nine museum specimens, are known only to a few
specialists.
The present revision is based on study of 462 museum
specimens and review of relevant literature. Specimens examined
are preserved in the museums listed below. I am deeply grateful to
officals of these institutions for permission to study material in
their custody.
AIUZ Anthropologisches Institut der Universitat Zurich
AMNH American Museum of Natural History, including
Archbold Collection (AMNH-AC), New York
BM British Museum (Natural History), London
BNHS Bombay Natural History Society, Bombay
CTNRC Centre for Thai National Reference Collections,
Applied Scientific Research Corporation of Thai-
land, Bangkok
FIELDIANA: ZOOLOGY. VOLUME 67
FMNH Field Museum of Natural History, Chicago
IRSN Institut Royal des Sciences Naturelles de Belgique,
Brussels
Museum of Comparative Zoology, Harvard Univer-
sity, Cambridge
Museum National d'Histoire Naturelle, Paris
Museum Zoologicum Bogoriense, Bogor
Naturhistorisches Museum, Basel
NHMBe Naturhistorisches Museum, Bern
NHRM Naturhistoriska Riksmuseet, Stockholm
Natur-Museum Senckenberg, Frankfurt
Rijksmuseum van Natuurlijke Historie, Leiden
Sarawak Museum, Kuching
Staatliches Museum fur Tierkunde, Dresden
United States National Museum of Natural History,
Washington, D.C.
Zoologisches Museum der Humboldt-Universitat, Ber-
lin
Zoologisches Museum der Universitat, Zurich
Zoological Reference Collection, University of Sing-
apore (This collection was formerly housed in the
Singapore National Museum, also known as Raffles
Museum.)
Zoologisches Sammlung des Bayerischen Staates,
Munich
Zoological Survey of India, National Zoological
Collection, Calcutta
Early stages in this research were supported by U.S. Public
Health Service Grant No. GM-13113. Final stages were partially
supported by Minorities Schools Biomedical Support Grant No.
RR-08043 (National Institutes of Health) to Chicago State
University. For skillful assistance I thank the following persons:
Mr. Kevin Royt, illustrations; Mr. John Bayalis and Mr. Fred
Huysmans, Division of Photography, Field Museum of Natural
History, photographs; Mrs. Nancy Shortall, typing.
NOTE ADDED IN PROOF: Volume 7 of Primates: Comparative
Anatomy and Taxonomy by W. C. Osman Hill (John Wiley and
Sons, 1974), which includes a discussion of liontail and pigtail
macaques (pp. 652-687), was published after the present work had
been submitted for publication; no evidence presented by Hill
requires modification of the present work.
MCZ
MNHN
MZB
NHMB
NMS
RMNH
SMK
SMTD
USNM
ZMB
ZMUZ
ZRCS
ZSBS
ZSI
3
•c
I
0
g
E
IS
15
:§3
3
•c
B
U
a
a
S'E
I 8
8"
GENERIC NAMES APPLIED TO
LIONTAIL AND PIGTAIL MACAQUES
Macaca Lac6pede, 1799, p. 4 — first applied to pigtail macaques by
Daudin, [1802], p. 148; synonyms listed below are those applicable
to liontail and pigtail macaques only.
Silenus Goldfuss, 1820, pp. xxiv, 479 — subgenus of Cynocephalus;
included species, C. silenus ( = Macaca silenus), type-species by
monotypy and tautonymy. Lesson, [1830], p. 101 — subgenus of
Macacus; included species, Macacus silenus, type-species by
original designation. Lesson, 1839, p. 70 — genus. J. A. Allen, 1916,
p. 51 — genus; selected to replace Macaca. Fjedler, 1956, p. 177 —
subgenus; external characters.
Les Maimons: I. Geoffroy, 1826, p. 588 — one of three subdivisions
("petite groups") recognized within Macacus; included species, M.
silenus, M. erythraeus ( = Macaca mulatto), M. nemestrinus ("Le
Maimon"), M. libidinosus ( = Macaca n. nemestrina), M. sped-
osus (=M. fuscata), M. maurus.
Maimon Wagner, [1839], p. 141 — subgenus of Inuus (=Macaca);
included species, /. silenus, I. erythraeus (=Af. mulatto), I.
nemestrinus (type-species by tautonymy; Maimon of Buffon cited
in synonymy, p. 144), /. arctoides, I. speciosus (=Af. fuscata), I.
niger. J. A. Allen, 1916, p. 50 — invalid designation of /.
erythraeus as type-species.
Vetulus Reichenbach, 1862, p. 125 — genus; included species, V.
silenus (type-species by tautonymy, synonym), V. nestor, V.
ursinus, V. priamus, V. thersites.
Nemestrinus Reichenbach, 1862, p. 139 (not Latreille, 1802),
subgenus of Macacus; included species, M. nemestrinus (type-
species by tautonymy), M. libidinosus (= Macaca n. nemestrina);
preoccupied by Nemestrinus Latreille, 1802, p. 437, a genus of
Diptera.
(UJUJ) L|JBU9| | IDJ^
KEY TO EXTERNAL CHARACTERS
Conspicuous external characters of adult male liontail and
pigtail macaques (fig. 1) are contrasted in the following key. Details
of individual, sexual and ontogenetic variation are discussed in
species and subspecies accounts. Although length of head and body
averages about 10-20 per cent greater in adult M. n. nemestrina
than in other members of this group, this character is not
absolutely diagnostic because of species overlap (figs. 2, 22).
1. Tail length greater than 50 per cent of combined length of head and trunk. Trunk
and limbs blackish; crown patch blackish, narrow anteriorly; cheek ruff and beard
prominent, pale grayish-brown; muzzle relatively short Macaca silenus
Tail length less than 50 per cent of combined length of head and trunk.
Macaca nemestrina, 2
2. Tail thinly furred, skin of tail clearly visible through sparse downy pelage. Trunk
chocolate-brown non-agouti dorsally, becoming pale brown ventrally; hind limbs
brown, forelimbs reddish-brown; crown patch brown, broad anteriorly; cheek hairs
short, lateral surface of throat region sharply defined pale ochraceous-buff; muzzle
relatively short M. n. pagensis
Tail densely furred, skin of tail completely covered by hair 3
3. Mid-dorsal region of trunk golden-brown agouti, similar in color to lateral surface
of trunk and limbs. Crown patch dark brown, narrow anteriorly; cheek ruff
moderately long, pale ochraceous-buff; reddish streaks extending laterally from
external angles of eyes; muzzle relatively short; tail carried arched forward over
back, tip of tail directed upward and forward M. n. leonina
Mid-dorsal region of trunk blackish, contrasting with golden-brown agouti color of
lateral surface of trunk and limbs. Crown patch blackish, broad anteriorly; cheek
hairs short, pale basally, blackish at tips; muzzle long; tail carried arched rearward,
tip of tail directed downward M. n. nemestrina
CRANIAL COMPARISONS
(Figures 3-6)
Cranial characters that appear to be of greatest taxonomic
importance in lion tail and pigtail macaques are (1) greatest length
of skull (excluding incisors), (2) relative zygomatic breadth, (3)
relative length of rostrum, (4) length of incisor-canine field and
premolar-molar field, (5) morphology of supraorbital ridges, and (6)
morphology of malar region. These characters are discussed in the
following paragraphs and their variation is graphed in Figures 7-9,
22. The method of measurement, except for dental -field measure-
ments defined below, is described and illustrated elsewhere (Fooden,
1969a, p. 40).
As usual in catarrhines, taxonomic differences in liontails and
pigtails are more distinct in skulls of adult males than in skulls of
females and immatures. Since only a few adult specimens of M. n.
pagensis and wild-collected M. silenus are available, no attempt has
been made to construct a formal key to cranial characters. Two
male and three female skulls of M. silenus have previously been
compared with an unspecified number of skulls of M. n. nemestrina
(Kurup, 1963, p. 249).
Greatest length of skull averages about 15 per cent larger in M.
n. nemestrina than in M. n. leonina, M. n. pagensis, and M. silenus,
which are approximately equal in this measurement (figs. 7, 22).
Overlap between M. n. nemestrina and the three smaller subspecies
or species is minimal. In specimens examined of M. n. leonina, M. n.
pagensis, and M. silenus, the range of greatest skull length is 102.3-
121.1 mm. in adult females and 124.1-140.1 mm. in adult males; in
M. n. nemestrina these ranges are 115.4-138.9 mm. in adult females
(only slightly smaller than in adult male M. n. leonina, M. n.
pagensis, and M. silenus) and 139.5-174.5 mm. in adult males.
Predictably, variation of skull length generally parallels variation of
length of head and body in these subspecies and species (figs. 2, 7,
22).
Fig. 3. Anterior and lateral views of adult skulls: M. silenus — a, female, RMNH
1314, zoo specimen; b, c, male, FMNH 47322, zoo specimen; M. n. leonina — d,
female, FMNH 99677, Loc. No. 37; e, f, male, FMNH 99691, Loc. No. 21. Approx. Vt
nat. size. (Photos FMNH)
For locality numbers see Figures 20, 25, and 28.
8
Fig. 4. Anterior and lateral views of adult skulls: M. n. nemestrina — a, female,
USNM 114502, Loc. No. 30; b, c, male, USNM 154367, Loc. No. 69; M. n. pagensis -
d, female, AMNH-AC 103398, Loc. No. 93; e, f, male, AMNH-AC 103394, Loc. No.
92. Approx. Vi nat. size. (Photos FMNH)
9
•' : ■ --;;:
".'•"'.;*. j *^wZl^^J^^^QPhS^^^^'^/^-^v.
^VjTV
H 1" 8 1 ^r x p^HBEeB^ v^&
' II 1
1 '■'-" * 1
B^a^lM
bW^mP^bI
ov^'-^&vtS. ^^B ^^^^^^ ,^H
Fig. 5. Dorsal and ventral views of adult male skulls: a, b, M. silenus, FMNH
47322, zoo specimen; c, d, M. n. leonina, FMNH 99691, Loc. No. 21. Approx. Vi nat.
size. (Photos FMNH)
10
Fig. 6. Dorsal and ventral views of adult male skulls: a, b, M. n. nemestrina,
USNM 154367, Loc. No. 69; c, d, M. n. pagensis, AMNH-AC 103394, Loc. No. 92.
Approx. Vi nat. size. (Photos FMNH)
11
12 FIELDIANA: ZOOLOGY, VOLUME 67
Relative zygomatic breadth (zygomatic breadth /greatest length
excluding incisors) in M. silenus and M. n. leonina tends to be
somewhat greater than in M. n. nemestrina and M. n. pagensis (fig.
7, table 1). Within each subspecies or species this ratio is about the
TABLE 1. Relative zygomatic breadth (zygomatic breadth/greatest skull
length x 100) in liontaii and pigtail macaques.
Mean — standard error; sample size indicated by figure in
parenthesis
Species or Infants and
subspecies juveniles Adult females Adult males
M. silenus 66.43 ± 0.65 (6) 68.40± 0.88 (7) 68.84 ± 0.68 (6)
M. n. leonina 64.88 ± 0.43 (23) 67. 39± 0.60 (19) 70.15 ± 0.58 (17)
M. n. nemestrina 63.26 ± 0.36 (50) 63.82± 0.39 (35) 64.49± 0.32 (76)
M. n. pagensis 62.89 ± 1.07 (2) 64.24 ± 0.68 (3) 63.11(1)
same in both sexes. In adult specimens examined relative zygomatic
breadth varies from 0.58 to 0.71 in M. n. nemestrina and M. n.
pagensis and from 0.63 to 0.75 in M. silenus and M. n. leonina. In
infant and juvenile specimens of all four subspecies and species,
zygomatic breadth is relatively narrow, approximately as in adult
M. n. nemestrina and M. n. pagensis. Zygomatic proportions in 12
M. n. nemestrina specimens (without locality data) measured by
Khajuria (1954, p. 118) fall within the range of variation in
specimens examined of that subspecies (fig. 7).
In liontaii and pigtail macaques, as in many other catarrhines
(Vogel, 1966, p. 21) relative length of rostrum (rostral
length/ postrostral length) is strongly correlated with skull size
(figs. 7, 8). In adult female skulls examined of M. silenus, M. n.
leonina, and M. n. pagensis, which are the smallest adult skulls
measured, relative rostral length is 0.40-0.55; in adult female skulls
of M. n. nemestrina, the next larger size class, relative rostral length
is 0.47-0.61; in adult male skulls of M. silenus, M. n. leonina, and
M. n. pagensis, which average slightly larger than those of female
M. n. nemestrina, relative rostral length is 0.55-0.61; in adult male
skulls of M. n. nemestrina, the largest size class in this group of
subspecies and species, relative rostral length is 0.62-0.78.
Although relative rostral length in liontails and pigtails varies
ontogenetically, sexually, and taxonomically, the composite log-log
plot of all measurements available for immature and adult males
and females is approximately linear (fig. 8). This indicates that
there probably is a constant allometric relationship between rostral
8 s
IS
i 5
3 «
(UHU) L((pD8Jq DUDLUOBX2
I
C C
E 4)
*, £
3 to
■O 0)
3
of "y
|.s
.§<rf
a cm
•a „,
■3 5,
« .2
1S
I*
.£ ^3
'■0
5*
I J
08 -
3 S
^ a -S
fa « a
S.2
13
Immoturat Adults
<f % <f
tiltnut
0 m laonma
O • ntmiilnno
v ▼ pagtntit
00
^6 15
PoilroJlral Ungth (mm)
no
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
Skeletons1 & 11
CadaverS2 5 1 2
Cadavers2 S 2 13
Skeletons1 $ 1
Skeletons1 £ 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,
—
0)
oo t"- os co co
tN OS t^ C^ 00
CN H rt .-H t-l
» 91 P3 O t>; N
•-< i — < cn cn i-< oo
a in n n oq
in cn © © ~
CN CN tN CN CN
CO to CN OS to
tO tT CN ^ f!
N N (N (N N
OS tN 00 © r~ ©
eo *f to «* cn ©
m h oo o » <r
m to in to ^r ^
o oo to to o^
x I in i cn ci co
CN tN <N CN (N
t- <— CO r- -rr o
to oo oo r- m' in
t^ m oo oo o
00
(C
re
CN
—
—
00
i/a
~i
CN
CN
CN
cn
•-
3
tN
a
-
o
3
o
"M
>C
•c
00
CO
■<r
u
CQ
Z
T5
o
t-
—
CO
re
W5
t^
(9
PJ
tN
CN
tN
CN
b
tN
3
r
la
o>
>
C
-Si
35
m
ee
CN
CO
tO
3
C
tv
©
CO
O
CO
n
•M
N
CN
CN
'JJ
tN
c
S
5J
si
tt
X
W
oo
O
o
t-
o
r-l
i-
C»
00
m
T-l
-1
Ol
—
T-t
r- •
CO Tf | »H CN CO t*
oc os © os m in
-» —i tN —• •» —
co — « co ic co t*»
00 6 6 O)' TT «5
>-. CM tN "-. t— i •«.
t> in co in in o
Tf os x C"- o in
x iH os os c- ^r
tN in co co o ^
CO ■— * CO r— * OS
co in co co eo'
m r- O — " CN CN
o ^ wi i-i OS OS
—
9
I-
in
os
-c
t>
^r
tN
CN
—
o
o
CO
|H
—
|H
"
i— i
1-1
I-
m
O
c-
■<*
o
C-l
CN
■ —
OS
o
H
"
1 '
~
""" '
"^
t-
^r
w
to
o
CN
—
-
tH
o
1
—
—
—
I— "
I-1
in co co co in os
© co d © x r~
Tf CO CO <— OS CN
O O OS O X X
co x in cn co Tf
h O <— i <-" OS X
o
z
*
a
*o *o o» o* o*
CM
CN C-
NhOO
CO CO CO
C~ OS c- «
fli CO V T) f
~ X ~ = -
ZS z z z
s x s s s
b Z d< B! B!
co
—
in
X
Z
•<3 *0 *0 »0 O* 0»
CO «-> X
t~ OS X
CO CO J3 CD W
OS OS ~- M OS -~r
OS OS -O £ OS «
in g ac x
Z Z Z 5 Z z
-i. ^- - - -
20
CO
©
1-H
CM
co'
ITS
1-H
©
t»
CO
d
o
CM
CO
1-H
1-t
1-H
CO
m
43
3
C8
1-H
CO
Z
a
u
.o.
z.
CD
1-H
5
3
f-
co
co
Ifl
m
g
•5
c
o
<N
—
«?
co
CO
rH
CN
CN
cu
CM
CM
kfl
0)
>
CO
1-H
3
C
■Si
3
CD
CO
t»
CO
t>
c
si
CN
CN
CO
£
X
t-;
CO
CO
o
c^
o
CO
CM O
1-H
CO
CO
CO
t^
CD
CO
CO
•*'
tJ<' CN
o
Ifl
!M
1-H
00
CD
CO
CD
1-H 1/2
CO
c-i
o
1-H
o
1-H
o
1-H
©
1-H
■«t
•*
■f*
l« CN
1-H
t-
CN
o
c-
©
o
in
t- CO
in
I-H
lO
CN
~
—
OJ
CO
t^
CO
cd in
CM
en
CD
rH
CO
"*.
CO
CO
Tf CO
I-H
T— 1
1-H
cc
—
CO
<—
-#
t>
t^
t>
CO CO
CN
en
CN
CO
CD
1-H Tf
CO
-<1«
O CO
CO
O
CN
ifl
CO
1-H
1-H
m
1-H
CD CD
o
CD
o c-
1-H
PD
0
Z
c
«
47322
3931
473872
d
4a
4)
a
CO
FMNH
NHMB
FMNH
RMNH
RMNH
CO rH
X
C~- CD
X
CO CD .O.
CD
w
CD CD --
05
CD CD .O
B
99
M
SIS
eg
S
K
z z z
5
X
/.
222
2
>.
— — -
u-
K
21
OlONOXNifin °. **.
cNOOtNOtNrHr-
x i co r- in co eo in It*
—
a
CD
^r
>H
as
pH
■~
TT
CN
-
in
n
as
<-i
o
FJ
CO
C
o
•«r
—
M
rn
'-
rH
(0
i-i eo
o
m
o
t^
X
t-
t-
\a
-
O
—
Ifl
CO
X
eo ih
rH
1-1
1-1
|H
*"*
r4
rH
*"*
—
ci
~~
— 1
?-*
*"*
i-l rH
o
as
Tl
o
—
o
a
O
^
CD
t^
CN
CO
O t-
m
rH
X
99
1-
co
(0
X
-~
m
-
1-
en
«T O
as
f->
t— 1
'"",
'~l
,H
—
■~~
,—
M.
**
—
,"H
f-H
1-H 1— »
cn
m
tfi
as
ac
t^
c
•o
CC
CO
^
C»
US
■-
co
O co
0)
X
VI
eo
—
r.
CO
CO
r»
ac
r- 1
CO
X
C-
r.
in cn
CO
3
"*
3
*"*
"""
'""'
— -
-~-
*""'
**"
IN
—
'""'
*"*
3
«->
CO
*"*
i— i ■»•«
d
ca
£
OS
•u
to
'C
OS
CO
©
CN
_
O
,_
ts.
^-
E
E
eo
> t-
Z
CO
c
m
O
©
CO
l-
t^
CO
—
CO
CO
CO
•X,
iO o
«
r~
T-H
'-'
O)
C-|
■— .
rH
*""
"-1
N
—
»H
<3
tN
— . r-
i-
a
VJ
0)
•S
to
<Sj
c
•4->
1-
11
o
©■
Cu
c
m
to
—
o
OS
co
o
Ifi
m
CO
o
CO
OS
CO
to
cv
s
CO
> CN
in o
«>
**"*
CN
H
T)
rH
—
—
»— 1
•—
i-^
I— 1
—
1-<
1-H
~H .— c
c
c
c
5
t>
5
co
T
O
CO
co
N
n
t-
W
1 —
CO
c
tO
c^ o
temtionHHiijH-n ^ cocnw
i— li— li— I.— <i— (i— ( rH t— < i— I f— c t— ( ^ rHrH
chcoini-H-^tocoojinineo
cbrHwocnaicdeoairHrH
os co co
os d t>
eoOOcoeoc— rH-rj-ascoO
rHOi-iOososososoboo
O CN X
co cd in
HOONnhnoionw
oioHoaiddriodo'
in o x
os c-^ in
rHifiosineO'TinocO'^'Os
oso-— "OososcNinosi—io
O co OS
r- C^ CD
5
X-
-
8
3
2
I
0
Z
c
1
I
5
s
X
*o*o*o*o*o*o»o*o*o*o*o
CD
tj< eo Os
CO CN X
O n on hi *
ZZ2(o2Z22ZD5
S5£Saio-o-D-o.N
in ^
cn in
m o
cn eo <~
-r -r l"
m in hi
FMNH
FMNH
RMNH
22
<N
CO
en
rH CO
OS
in
<N
CO
CO <N
^*
t^
%o
00
— i
X
X
CO
</3
q rH
00
t— t
co
OJ
—
CO
CO
CO
0)
Tt ^r
«-"
3
o
j-
5
ca
2
3
3
rH
£
«Q
«
00
T3
m co
CO
Oi
CO
05
X
s
i> m
t^
£
ri
eg
a
rr rf
m"
"*
^p
^r
TT
^t CO
_C
•S
>
CO
c
o
00
CO
us
CD CO
CO
in
00
CO
r-
lri
CO
cu
T— 1
X
in in
PQ
<
a
x
X
CO
x'xxr-r-x'oSt^x ■«:
cocoas— 'HOh^nn
inooaJoioioooocri
COCXN(Cn«0>NHCO
tCNNHHOHTfOH
CO i-t
CO(NX20-(MCO^
-TCOXq^-,*-,^h^
ZZZNZZZZD5
x w x —
co co co
ai cd in
o .->
rH 00
CO CO rH
<N oi X
54252
54305
15f
FMNH
FMNH
RMNH
o
c
• c
so o
c • -
1?
•— —
Xi 3
<D -a
_ X)
« «
.S £
£ =>
H a-
CO >,
o> —
rv C
3 ©
u O
_: at c
t« a*
0—0)
3 — -»->
s £ 2*
o o 2
0> O ai
• — CO
<N O 2
* 3 3
'£ -a T3
Oi W £8
Z- -- —
3 3 3
~S X X
H CN CO •* ifi
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££ wir 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
542621
6
19.8
19.0
13.2
11.8
10.0
6.8
5.7
543051
?
(1.80)
17.4
(2.00)
15.8
(1.65)
11.8
(1.19)
11.1
(0.74)
7.3
(0.39)
5.9
(0.27)
5.0
(2.38)
(2.25)
(1.44)
(1.08)
(0.57)
(0.38)
(0.32)
1 Subadult (see Table 3).
usually absolutely (table 4), than in M. n. nemestrina (and M.
silenus). Reduction of these transverse processes in specimens of M.
n. nemestrina has been noted previously by Wilson (1970, p. 53, fig.
3). These transverse processes receive the tendons of insertion of m.
ischiocaudalis, which acts to abduct and depress the base of the tail
(Wilson, 1970, p. 78; Trevor-Jones, 1970, pp. 320, 323). Enlargement
of these transverse processes in M. n. leonina may be a function of
increased abductor and ventriflexor tension. This, in turn, may
stabilize the base of the tail against increased extensor tension (of
m. extensor caudae medialis) that apparently is exerted more
distally to effect the characteristic dorsal hyperextension of joints
Ca4-5, Ca5-6, and Caf>7.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 27
A more complete explanation of the structural basis for the
difference of tail carriage in M. n. leonina and M. n. nemestrina
probably will require further comparative study employing more
adequate skeletal and cadaver material of M. n. nemestrina. The
evolutionary significance of the tail carriage difference between
these subspecies, like that of many conspicuous and taxonomically
important differences between primates subspecies and species,
remains obscure.
FEMALE GENITALIA
Cyclical estrous swelling of the circumanal sexual skin is well
known in M. n. nemestrina (F. Cuvier, 1816-1817, pi. 5, fig. 2;
Pocock, 1926, fig. 68b; Zuckerman, 1937, p. 319; Kuehn et al., 1965,
figs. 1, 2; White et al., 1973, p. 189). This swelling typically develops
gradually, beginning during or immediately after the menstrual
period and reaching maximal tumescence usually in about 15 days.
At its maximum, the engorged pinkish hairless area extends
dorsoventrally from the base of the tail to the ventral border of the
ischial callosities, which are partly buried in the swelling, and
extends laterally over an area twice the breadth of both callosities
(fig. 11). Ventral to the callosities the thinly haired skin anterior to
the vulva (mons pubis) is less conspicuously swollen. Following
ovulation, usually within one or two days after maximal tumes-
cence, the swelling rapidly subsides. Detumescence persists for
about 15 days, until the next menstrual period. In laboratory
mating tests, maximal female perineal tumescence in M. n.
nemestrina is directly correlated with maximal sexual behavior in
females (Goldfoot, 1971, p. 330) and with maximal incidence and
rate of ejaculation in males (Bullock et al., 1972, p. 230).
Sexual skin in M. silenus has been cursorily described in an
unspecified number of specimens by Pocock (1926, p. 1547) and
more precisely characterized in one specimen by W. C. O. Hill (1937,
p. 210, figs. 2a, 2b). Judging from these accounts, the general
pattern of sexual skin morphology is similar in M. silenus and M. n.
nemestrina, although swelling is much less distended in estrous M.
silenus (fig. 11). In this species the swelling does not form a
continuous pillowlike mass, as in M. n. nemestrina, but instead is
subdivided into five separate swollen areas— one large subcaudal
swelling, a pair of smaller swellings dorsolateral to the anus, and a
pair of broad swellings that surround the ischial callosities, lateral
to the vulva. (The ventral margin of the vulva in one M. silenus
adult female that I observed at Lincoln Park Zoo, Chicago, is
approximately in line with the lower border of the ischial callosities,
28
FOODEN: LIONTAIL & PIGTAIL MACAQUES
29
M. silenus
M. n. nemestrina
Fig. 11. Female sexual swellings in M. silenus (modified from W. C. O. Hill, 1937,
p. 214) and M. n. nemestrina (Kuehn et aL, 1965, p. 253): 1, subcaudal swelling; 2,
para-anal swellings; 3, para-vulval swellings.
as in other species of macaques; the vulva is not especially high
between the callosities, as described and figured by W. C. O. Hill,
loc. cit.) The paired swellings dorsolateral to the anus in M. silenus
are particularly less developed than in M. n. nemestrina, where
these para-anal swellings fuse with the subcaudal swellings dorsally,
extend over the gluteal region dorsolateral^, and overlap the ischial
callosities ventrally.
Morphology of sexual swelling has not been reported in estrous
specimens of M. n. leonina and M. n. pagensis. In non-estrous M. n.
leonina specimens examined (FMNH 99674, 99687, subadults;
99685, 99688, pregnant adults; 99677, 99678, 99679, lactating adults),
the hairless area of sexual skin seems to be less extensive than in
corresponding non-estrous specimens of M. n. nemestrina, more
nearly resembling that in M. silenus.
Internal female reproductive structures have been previously
investigated in three specimens of M. n. nemestrina (Hafez and
Jaszczak, 1972, pp. 297, 300). The present account is based on study
of 12 alcohol-preserved reproductive tracts of M. n. leonina
immatures and adults (table 5). The female tract in M. n. leonina
(fig. 12) is essentially similar to that in M. n. nemestrina (Hafez and
Jaszczak, op. cit.). It is also generally similar to that in M.
>
£
lA lO
a
.2
00
q
CO
i— <
U3
t» co
CO CO
«-J
q q
•o
1-1
cn
CO
cn
CO
c- co
cn r-
t^
d t^
>>
"8
at
•
0
CO
00 00
q oq
00
q »-<
01
0>
C
in
«5
N.
if
CN
CN ©
r-i C35
d
CO CO
c
PQ
d
c-^
00
00
l-H
i— • rl
CO <-
CN
f-H 1-H
'C
i— i
3
X
D
c
•
m
m
q
t-; C"^
© q
CN
CN r«
in
CO
d
6
■<r
00 CO
C-^ 00
oo'
CN d
J
d
t~
»-<
rH
rl
1-t 1—1
CO O
i— <
CN i-
3
U
3
|
I
I
(d «
o> 0 *
2 O CJ J=
OS
■ao
M
c
v
J
00 co
^ m g
>
3
q I-;
CO CO
O C-;
d d
CO T
<J> CN
.-h CN
00 CN
CO CO
q q
oo oo
co -«r
q
d
o cn
d -
CN O
co ;i
3
T3
cd
X)
3
CO
d
2
-c
co q
rr d
q q
CN iH
CO <N
d d
■«r CN
i-> CN
CO CN
d d
00(0
d d cn
^
=
Tf lO oo
cn d d
00 CO 0"i
d d d
q
d
d oo
i-i in
m c- o>
CN CO
CO CO
CO CN CN
Cd «
« T3 TJ J*
*cgg
— - — —
—
m
'•"
c
i—i
CN
d
^r
CO CO
CO
CO
CO
~
~
A/\
A A
CN
CO CO
<N
E
SS
W
u
l
CN
>— K
CN CN
CO
CO
CO
m
re
-
o>
!"^.
2
P-
2S
ss
sss
-r
■*r
oo
00
o
CO
CN
CO
00
00
•t
t-
00
t»
^"
Oi
N 1
CO
t~
00
TT
00
ph
CN
co
CO
CO
-*
1-
T
■*"
—
—
in
s
pa
2
X
Z d
S Z
CN <N
00 Oi
CO CO
C7> Ol
© ©
© r^
00 00
CO CO
© ©
© ©
00
r.
S
■fco
© Z
*r in
co r-
m oo
o> 00 t»
00 00
r- t- t-
CO CO
CO CO CO
© ©
OS © ©
© ©
05 CJ> CJ5
30
t-
Oi
t>-
00
a •
*3
. >>
. C
o 50
J= o
co °
c £
Oi o
'-• a
-- o
01 >-
_■ o*
rH CO
(90 g
« .2
5 a,
.2 *
09
—
o °
JS C
jd
■*»* _
-•-> cj
1 c
o £
3 g
a c
H
0) o
3 £
o "«
£ in
I— 1 I-H
* &
« £
■*-> _
« ■>->
c ">
._ ^
CO CN
3
00 00
CO CO
Oi OS
£ S
° £h
8 £r
£r£
£ m
X X
z z
a
0
fa fa
c
c —
tx
>w* ^-^
o
o> a*
rs
o **
-*J -»J
C
CI UJ
U
c c
cs
U
■3
>- «J
£ «
_. I-
cs Q
IE
U
o
CM zi
01 o>
£ o
.£ .£
0 0
a
*j ™
« ._>
0) o>
(0
o
0) 0
'8
0)
2*
vi 0)
be be
CO CQ
T3 -O
a>
C 0
>
>> X,
o> o>
e
o s
» 0>
C
2 c
CQ cs
£ £
0)
01 —
e/5 —
0)
«j *^3
'■W 4J
08
cs O
.c
CA (O
0*
PQUh
— -7
fa fa
i-J
N CO
*tf lO
o t>
CN
q
CO CN
r-J ©
i-H
r-^
°°.
°°.
m
q
CO
CN
CN CN
T-i <n
CO
T-H
CN
CN
CN
rf'
X
X
X X
X X
X
X
X
X
X
X
q
q
rH O
00 CN
CO
q
CN
q
q
q
t~
t^-
CO' CT)
CN ■<*
lO
Tf
Tf
CO
iC
c-^
X
X
X X
X X
X
X
X
X
X
X
I-H
t~
q q
CO CO
q
I-H
-*
q
q
eN
CN
d
CO <-H
in m
00
C^
<T>
t^
CO
00
I-H
q
I-H
I-H
I-H
co q
q ©
q
CO
q
t-H
■*
CN
CO
CN CO
i-i CN
CN
CO
CO
CN
CN
X
X
X X
X X
X
X
X
X
X
■<f
q
q q
© t~-
q
iq
CN
I-H
q
05
00
in cd
CO CO
1
rr
cd
CO
co'
CO
X
X
X X
X X
X
X
X
X
X
q
CO
Tf Tf
00 CO
c^
q
q
q
q
<N
d
00 CN
T? to
CO
oi
00
lO
r^
I-H
I-H
i-H
o
7.
X
z
rf iO
co r~
<N CN
OJ I-H
r^
■*eo
m 00
OS 00 t>
00 Ol
00 00
00
I> rt
00 00
t~ c~ t~
CO CO
CO CO
CO
CO 5
CO CO
CO CO CO
OS Oi
O <T>
Oi
CT> g
<j> a>
CJi CJ5 C7>
CJ5 O
OJ <3>
05
o> Z
C31 o>
CJ> CTl (Ji
31
32
FIELDIANA: ZOOLOGY, VOLUME 67
Fig. 12. Sagittal section of female reproductive tract in subadult M. ru leonina,
FMNH 99674; x 2. UB: uterine body, UC: uterine cervix, V: vagina (distortion of
wall is artifact of preservation). (Photo FMNH)
fascicularis (Jainudeen et al., 1972, p. 473) and M. mulatto
(Wislocki, 1933, p. 239; Eckstein and Zuckerman, 1956, p. 138;
Cuadros, 1971, p. 139). However, the uterine cervix in M. n. leonina
evidently is somewhat smaller than at corresponding stages in M.
mulatto. Judging from specimens examined (table 5), the cervix in
M. n. leonina enlarges markedly approximately at puberty (FMNH
99687), regresses during adolescence (FMNH 99674; NM4), reaches
maximal size during pregnancy (FMNH 99685, 99688), and regresses
during lactation (FMNH 99677, 99679).
Structure of the female tract in M. n. leonina is distinctly
different from that in the M. sinica-group (M. sinica, M. radiata,
M. assamensis, M. thibetana), which is characterized by presence of
cornified vaginal spines and a hypertrophied endocervical chamber
(Fooden, 1971b, p. 67). It also differs from that in M. arctoides, in
which a large dorsal vestibular collide is present and cervical
colliculi are absent (Fooden, 1967, p. 939; Demers et al., 1972, p.
529; Hafez and Jaszczak, 1972, p. 302).
NOTE ADDED IN PROOF: Cf. Eaton, G. G. and J. A. Resko, 1974,
Ovarian hormones and sexual behavior in Macaca nemestrina, J.
Comp. Physiol. Psychol., 86, pp. 919-925.
GLANS PENIS AND BACULUM
This account is based on study of 30 specimens of penises or
bacula of liontail and pigtail macaques (M. silenus, 3; M. n. leonina,
19; M. n. nemestrina, 8; tables 6, 9) and review of relevant literature
(Daubenton, in Buffon and Daubenton, 1766, p. 179; F. Cuvier,
1820, p. 1; F. Cuvier, 1822a, p. 2; DeBeaux, 1917, p. 10; 1923, p. 34;
Pocock, 1921, p. 227; 1926, p. 1559; Pohl, 1928, p. 103; W. C. O. Hill,
1937, p. 213; 1958, p. 655).
In liontail and pigtail macaques the glans penis is helmet
shaped and bluntly bilobed (fig. 13), closely resembling that in
Celebesian macaques (Fooden, 1969a, p. 20) and M. sylvanus
(Daubenton, in Buffon and Daubenton, 1766, p. 119; W. C. O. Hill,
1958, p. 655, fig. 18a) and generally similar to that in the mulatta-
group (M. fascicularis, M. mulatto,, M. cyclopis, M. fuscata; Fooden,
1972, p. 310). The bluntly rounded glans in macaque groups listed
above is distinctly different from the apically elongate subpyramid-
al glans in the smica-group (M. sinica, M. radiata, M. assamensis,
M. thibetana; Fooden, 1971b, p. 72) and the even more elongate
lanceolate glans in M. arctoides (Fooden, 1967, p. 939). These three
sharply defined glans types in the genus Macaca (fig. 14) were first
recognized by Pocock (1921, p. 229); subsequent authors, including
Pocock (1926, p. 1557), have tended to obscure the basic penial
trichotomy of macaques. The bluntly rounded glans type seems to
be the most common form in macaques and other cercopithecids
(Pocock, 1926, p. 1557) and, therefore, probably is primitive in the
genus and family.
The three subdivisions of the genus Macaca that are defined by
structure of the glans penis exactly correspond, as far as presently
known, to three subdivisions defined by structure of the uterine
cervix. In M. n. leonina, M. fascicularis, and M. mulatto, all with
glans bluntly rounded, the cervix and cervical colliculi are
moderately large (see above); in M. sinica, M. radiata, and M.
assamensis, with glans apically elongate, the cervix is enormously
inflated and cervical colliculi are hypertrophied (Fooden, 1971b, p.
Q a*
inn h
in ■<? m
© <n «-i ^* © cn cn © « ©
»-( l-H CN 1-HC^»-Hl-Hl-l»-l'-<»-l'-'C^
s
H
3
_3
O"
CO
3
u
CJ
ea
CO
£
-
>
(m
—
£
o
1
a
B
Ifl (O t«;
CN CN CN
c
3
5
1
c
1
i
o
s
c
**
".
"5
09
•o
i-H (N <N
in © in in © CN CN © 00 CN
CM* co cn cn oi cn co cn CN CO
8 JP
v. C
»-l f-H -<t
(N (N (N
t-r-©©©©"'}'©^^?
t-iCNCNOJCNCN<N<NCNCN
3
U
1
&
'.3 "Q
cu .a
00 ©
r- ©
tN CO 00 «N CO ^f
r- t- © it- © r^
d © © © © ©
© 00
00 ©
o iS
00
c
m ©
o ^r
I s
© CN Tf © © 00
in m
*-< oo
oo t» oo to t* (>■ . com
!-( rH ,-H i-H i-h i-H I rl «-H
O
Z
I
CO
o *o
m 2
in X
m 2
§2
CN W
CN ©
© ©
^ ©
tt in
CN CI
KlO © © © <r> 2 S
fcZZZZ^DD
34
2 ° c
Q a*
00 00 O Tf N t» 00
00 H 0> ^ N H N
tfJoJ-^iriui^ioioo
S
3
3
o
C8
22
m oj i-i <o c~ m cm
<N <N CN i .-< r-i <N <N (N
v. c
BJ .S
_5
-
c
0
-
PQ
2
4
c
-
o
Z
c
£
o
4)
a
en
(OH HOHD03 H
i-tCNr-lr-li-li-l01?-l
I I I I
OO) H
CO 00 t>
I
u
IS
o
«4-l
o
1
>
c
D
a»
e
I
c
<:
I I I
in iflm
, CO O) Tf o
4) fc
1 *
0 3 =
r- <-> 0/
•= 3 !2
_^ /• _
w c
- 0 Oh
CO
©CN
CI
2oo
CO
sill
1/3 ITS X
. CO
£ a
12
i2v .§ "S i»'
5 g
* o
« - o
in m ^
CQ CO E
CO <N JC
N CQ Z
OSS
Sz2
0 3
N CO CO Q CJ CQ
a. o
e
o
35
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 specimens1 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 length4 (mm.) Relative breadth
M. fascicularis
6.2, 7.7, 8.5
0.44, 0.45, 0.52
7.4, 7.7
0.41, 0.42
7.0
0.43
7.4, 12.3
0.47, 0.53
12.9
0.49
14.4, 15.2, 16.6
0.55, 0.46, 0.41
M.
mulatto
11.8 (10.6-13. 1)5
0.42(0.36-0.50)5
13.5
0.36
9.5
0.47
18.7
0.54
M.
fusct
ita6
17.0, 17.1
0.38, 0.40
18.3
0.40
15.9
0.39
20.0
0.40
20.0
0.48
1- Specimens in FMNH.
2- Parentheses indicate incompleted erupted teeth.
3- Based on dental emergence norms for M . mulatto (Hurme, 1960, p. 797).
4- Measured parallel to long axis of glans; differs slightly from greatest length
reported previously (Fooden, 1972, p. 306).
5- Mean and extremes in eight specimens.
6- For age estimates see Fooden, 1972, p. 306.
38 FIELDIANA: ZOOLOGY, VOLUME 67
and pigtail macaques, judging from measurements of dry specimens
reconstituted in fluid (table 8).
TABLE 8. Relative breadth of glans penis in reconstituted dry specimens
of Celebesian macaques.
Glans Relative
Species Specimen No. length (mm.) breadth
M brunnescens NMS 1017, 1020 13.8,15.2 0.59,0.63
M hecki RMNH 1099, 3801 14.9, 17.3 0.66, 0.51
Af maura RMNH 751 (2363, 2484) 8.4, 13.3 0.83, 0.66
M. nigra RMNH 13, MCZ 34459 7.9,9.8 0.87,0.72
Af ochreata NMS 1022 13.2 0.61
Af tonkeana NMS 16838 13.2 0.64
In liontail and pigtail macaques (fig. 13), as in M. mulatto, the
dorsal part of the glans extends back over the shaft about twice as
far as the ventral part, the glans is bilobed, the urethral orifice
opens between the lobes, and the baculum protrudes into the left
lobe, which is somewhat larger than the right. Also as in Af.
mulatto, the glans and distal part of the shaft are densely studded
with tiny epidermal papillae about 0.05 mm. in diameter; these are
much smaller than the epidermal spines (0.3 mm. diameter) present
on corresponding penial surfaces in the sinica -group (fig. 14). In
early accounts (Daubenton, in Buffon and Daubenton, 1766, p. 179;
F. Cuvier, 1822a, p. 2) the glans in M. n. nemestrina is
characterized, somewhat confusingly, as three-lobed; this evidently
is a result of counting as a lobe a small variably defined V-shaped
dorsomedian eminence that is situated between the bases of the
right and left lobes, proximal to the urethral cleft (fig. 13).
In living specimens of M. silenus, color of the detumescent
glans is dark gray ( W. C. O. Hill, 1937, p. 214; personal observation,
one adult, Lincoln Park Zoo, Chicago) and color of the tumescent
glans is "bright purple" (W. C. O. Hill, loc. cit). In M. n. leonina,
the glans is pinkish (DeBeaux, 1923, p. 35; personal observation, one
adult, National Zoological Park, Washington, D.C.). In M. n.
nemestrina the color of the "penis" (?glans) is recorded as "pale
crimson" (field tag of BM 55.1503, Sungai Renggam, West
Malaysia).
The external similarity in structure of the glans in liontail and
pigtail macaques conceals unexpected internal differences in size
and structure of the baculum (fig. 15; table 6). Comparing species
for which reasonably adequate samples are available, length of the
Fig. 14. Basic glans types in genus Macaco, left lateral and dorsal views: a, b,
bluntly rounded type, M. fascicularis, FMNH 99642; c, d, subpyramidal type, M.
h, 99e91,> 6 yr.; i, 99673, > > 6 yr. Natural size. (Courtesy Chicago Zoological Park,
Brookfield, 111.)
39
40 FIELDIANA: ZOOLOGY, VOLUME 67
baculum in M. n. leonina (21.0-26.7 mm. in 10 adults) averages
about 25 per cent greater than in M. n. nemestrina (16.1-21.1 mm.
in seven adults; cf. Kinzey, 1971, p. 102). This is somewhat
surprising, considering that head and body length in M. n. leonina
averages about 15 per cent less than in M. n. nemestrina (fig. 2).
The bacular length difference in M. n. leonina and M. n. nemestrina
is due principally to a difference in length of the downward
inflected distal process (fig. 15). In M. n. leonina this process is
sharply defined, and its length is 29-51 per cent of that of the
proximal bacular shaft (10 specimens); in M. n. nemestrina the
inflected distal process is more weakly defined, and its length is 20-
30 per cent of that of the proximal shaft (seven specimens). The
dorsoventral diameter of the bacular shaft also is notably greater in
M. n. leonina than in M. n. nemestrina; as a result, the shaft is
bilaterally flattened in M. n. leonina and more nearly cylindrical in
M. n. nemestrina.
One baculum of M. n. leonina and one of M. n. nemestrina are
anomalous. In the anomalous M. n. leonina specimen (NM6; fig.
15k) the shaft is unusually slender, the basal end is studded with
small tuberosities, and the distal process, which is weakly inflected,
terminates in three vertically arrayed lobules, in place of a simple
ventrally directed tip. In the anomalous M. n. nemestrina baculum
(AMNH 119514, zoo animal) the bone is unusually thickened and
porous (fig. 15s). As indicated previously (Fooden, 1969a, p. 25), the
abnormal structure of this baculum may be a result of captivity-
induced bone pathology. However, bacular structure in two other
captive specimens examined (NMS 2629; RMNH 2000; figs. 15p,
15q) is essentially similar to that in wild-collected specimens.
Although examination of three available specimens of M.
silenus (fig. 15a-c, table 6) suggests that bacular structure is
Opposite:
FlG. 15. Bacula in adult liontail and pigtail macaques; left lateral view, dorsal
surface of baculum at top, distal end at left; italicized figures below indicate locality
numbers as in distribution maps, Figures 25, 28. a, b, c, M. silenus (zoo specimens):
BM 41.1.15.1 (subadult), NMS 2630 (subadult or adult), RMNH 1308 (subadult or
adult); d-k, M. n. leonina: BM 55.1505 47.NM 5 ("Thailand"), NM 7 ('Thailand"),
USNM 241022 25, NM2 ("Thailand"), PMl ('Thailand"), USNM 258868 36, NM6
("Thailand"); !-•, M. n. nemestrina: BM 55.1503 12, MCZ 35670 89, FMNH 68703 90
(subadult), NMS 2629 (zoo), RMNH 2000 (subadult or adult, zoo), BM 55.707 58,
FMNH 105676 4 (subadult), AMNH 119514 (captive, deformed), x 2. (Photos
FMNH)
41
a
0
o
a
9
es
s
!
3
-
£
£
w) O
co i »■« o» oe i »ft © «r ^ cn
Q S. *
3=
CO
_ J=
t— i •— t— i t—c i— i CM
2 A -2 - x
ff-mtttO'-rtioo
a
-
OH
-
a
Si
F^
rH
i-
o
r-
(0
1-
as
r»
zt
1 —
O
X
pH
m
o>
as
~]
-
V
1 —
—
iH
fH
1-1
i-t
?\
n
o>
>
— '
—
X
X
pm
^r
O
o
X
re
N
i
•—
o
O
X
o
o
o
t-
-
o
o
O
• JS
OhOtONfflMNO)
» t« C« X N ^' 1(5 t« X
£
M
-c
1-
*■» 01
.<« BO
— CO
.8
c
8.
■a
4 c
,—. <-^ ^^ > i_
—
NN CO CO . 0
H
V
E~~ SS » *
c
0)
ft
—
e
o> c
-M
K3
tt
as r~
tN CO
3
o
c
c-j
«-« r-l
OS CD
CO CO
— >,3 r~ to <o © t «-" <fi
(5«otte m B(OX h ^ t-
01 q C CN P3 P5 CO ^ ^ ^
S £ i!
O
Z
X
S
E
ooiemioiNnfj
<J>Xt~r~XXr~c35r-
lD<0<0(OtO<0(C<ClO
C35C7>05C7><35C75<75C7>05
C7>CTlC75CT>C75C75C75CT>C75
42
FOODEN: LIONTAIL & PIGTAIL MACAQUES
43
~'m
k
^ mLm •
H^H^^Bi
fc
jpm**zi
Fig. 16. Radiographs of glans penis in immature and adult M. n. leonina showing
ontogenetic development of baculum: a, FMNH 99690, estimated age 1 yr.; b, 99680,
2.5 yr.; c, 99676, 3 yr.; d, 99675, 3.5 yr.; e, 99686, 4.5 yr.; f, 99684, 6 yr.; g, 99672, 6 yr.;
h, 99691, > 6 yr.; i, 99673, >> 6 yr. Natural size. (Courtesy Chicago Zoological
Park, Brookfield, 111.)
taxonomically distinctive in this macaque, as in M. n. leonina and
M. n. nemestrina, the evidence is imperfect and should be
interpreted with caution. All three bacula are from zoo animals,
which introduces the possibility of captivity-induced deformation.
In addition, one of these specimens is from a known subadult, and
the other two are from animals of unknown age. In these three M.
silenus bacula, the bases are exceptionally thick, the proximal
shafts are stout and cylindrical, and the distal processes are short
and weakly defined. Bacular form in these specimens is more like
that in M. n. nemestrina than in M. n. leonina; however, overall
size exceeds that in M. n. nemestrina, and diameter of the base is
disproportionally greater than in M. n. nemestrina.
A series of nine M. n. leonina specimens collected in a
restricted area in western Thailand in 1967 (Fooden, 1971a, p. 32)
provides an opportunity for investigation of ontogenetic devel-
44 FIELDIANA: ZOOLOGY, VOLUME 67
opment of penial structure in this subspecies. Estimated ages of
these nine specimens are: 1 year (infant), 1; 3 years (juvenile), 3; 4-5
years (late juvenile), 1; 6 years (subadult), 2; > 6 years (adult), 2
(table 9). Judging from available evidence (table 9, fig. 16), size of
glans and baculum remains approximately constant from about age
one year to age three years, while length of head and body is
increasing by nearly 50 per cent. Beginning about age 4.5 years,
when permanent canines emerge and puberty begins, size of glans
and baculum increases rapidly, at a rate about five times that of
head and body (linear dimensions). This pattern of development
suggests that penial growth in M. n. leonina is controlled primarily
by sex hormones, rather than by pituitary growth hormones.
Similar indications for M. fuscata were reported previously (Fooden,
1972, p. 309). The characteristic prolongation of the downward
inflected distal process of the baculum appears relatively late in
ontogenetic development of M. n. leonina (fig. 16; table 9). As a
result, bacular form in juvenile M. n. leonina (fig. 16), resembles
that in adult M. n. nemestrina (fig. 15), which therefore is
paedomorphic for this character.
BLOOD PROTEINS
Data suitable for serological comparison of M. n. leonina and
M. n. nemestrina are provided by a recent series of studies of blood
protein polymorphism in various macaques (table 10). Although
authors of the studies do not explicitly recognize these two
subspecies of pigtails, adequate information is given on the
geographic origin of specimens to permit subspecific identification.
No blood protein data are available for M. n. pagensis or for M.
silenus.
Blood protein phenotypes are variably differentiated in M. n.
leonina and M. n. nemestrina (table 10), although none of these
phenotypes are absolutely diagnostic and characteristic in these or
other macaque subspecies and species. M. n. leonina and M. n.
nemestrina are essentially similar in frequency distributions of
phenotypes of haptoglobin, acid phosphatase, albumin, thyroxine-
binding prealbumin, alkaline phosphatase, plasma phosphohexose
isomerase, 6-phosphogluconate dehydrogenase, and phosphogluco-
mutase. These two subspecies differ slightly to moderately in
frequency distributions of phenotypes of alpha l-antitrypsin, carbo-
nic anhydrase II, erythrocyte phosphohexose isomerase, tranferrin,
and carbonic anhydrase I. These subspecies differ strikingly in
frequency distributions of phenotypes of plasma agglutinin and
hemoglobin. The ratio of anti-A to anti-A + B aglutinin phenotypes
is about 3:1 in M . n. leonina, and about 1:9 in M. n. nemestrina', the
ratio of FS to F hemoglobin phenotypes (determined elec-
trophoretically) is about 3:1 in M. n. leonina, while in M. n.
nemestrina F is the only hemoglobin so far detected elec-
trophoretically. Since all studies in this series evidently are based
on blood samples from many or most of the same monkeys,
differences in the frequency distributions presumably indicated that
at least some of these blood protein phenotypes are inherited
independently; however, the loci for carbonic anhydrase I and
carbonic anhydrase II probably are linked in these macaques
45
TABLE 10. Frequency distribution (per cent) of polymorphic blood pro-
tein phenotypes in Af. n. leonina and Af. n. nemestrina.
Plasma proteins
Haptoglobin *
Af. n. leonina (85 specimens)
Af. n. nemestrina (158 specimens
Albumin*
Af. n. leonina (85)
Af. n. nemestrina (158)
Thyroxine-binding prealbumin^
Af. n. leonina (36)
Af. n. nemestrina (128)
Phospohexose isomerase^
Af. n. leonina (25)
Af. n. nemestrina (73)
Alkaline phosphatase*
Af. n. leonina (85)
Af. n. nemestrina (158)
Alpha \ -antitrypsin4
Af. n. leonina (64)
Af. n. nemestrina (151)
Transferrin^
Af. n. leonina (85)
Af. n. nemestrina (158)
Af. n. leonina (cont.)
Af. n. nemestrina (cont.)
Agglutinin^
Af. n. leonina (82)
Af. n. nemestrina (144)
Erythrocyte proteins
NADH diaphorase?
Af. n. nemestrina (25)
Acid phosphatase^
Af. n. leonina (40)
Af. n. nemestrina (127)
6-phosphogluconate dehydrogenase^
Af. n. leonina (36)
Af. n. nemestrina (110)
Phosphoglucomutase^
Af. n. leonina (23)
Af. n. nemestrina (111)
Carbonic anhydrase 11*0
Af. n. leonina (17)
Af. n. nemestrina ( 12)
Phenotypes
\ and frequencies
Hpl-1
100.0
) 100.0
Single
Double
98.8
1.2
98.1
1.8
PA«
PAfs
100.0
0
97.2
2.8
1
6-1 Unidentified
100.0
0
0
94.5
1.4
A
AC
100.0
0
94.3
5.7
AB B
AC BC C CD
BD
D
0 1.6
0
7.8 68.7 14.1
1.6
6.2
0.7 6.6
1.
3 32.4 58.3 0.7
0
0
BG CC CD CF CG D"D DD DF
1.2 1.2 3.5 4.7 12.9
0 0.6 3.2 4.5 2.5
DG F'GFF' F'G FF
17.7 0 0 0 3.5
23.4 0.6 1.3 0.6 7.6
1.2 8.2 3.5
0 13.9 17.7
FG GG
10.6 31.8
17.7 6.4
Anti-A
Anti-B
Anti-A+B
74.4
1.2
24.4
9.0
2.1
88.9
Phenotypes and frequencies
A AB AC
92.0 4.0 4.0
A
100.0
100.0
AS
A
AF
2.8
97.2
0
0
97.3
2.7
I
V
95.7
4.2
1
92.8
7.5
»
b
32
a2b
52.9
17.7
29.4
66.6
16.7
16.7
46
1 5-1
6-1
81.8 18.2
0
97.1 0
2.9
a b ab
c d o
7.9 57.1 1.6
0 6.4 27.0
32.1 23.7 9.1
1.2 1.2 32.7
F FS
S
24.7 72.9
2.4
100.0 0
0
FOODEN: LIONTAIL & PIGTAIL MACAQUES 47
TABLE 10. (continued)
Erythrocyte proteins Phenotypes and frequencies
Phosphohexose isomerase^
M. n. leonina (11)
M. n. nemestrina (34)
Carbonic anhydrase 1*0
M. n. leonina (63)
M. n. nemestrina (165)
Hemoglobin^1
M. n. leonina (85)
M. n. nemestrina (160)
1- Ishimoto, 1972a, pp. 260, 273.
2. Weiss etal., 1971, p. 78.
3- Ishimoto and Kuwata, 1972, p. 161; enzyme activity stronger in red cells
than in plasma.
4- Omoto et al., 1970, p. 220.
5- Ishimoto, 1972a, p. 255; cf. Prychodko et al., 1971, p. 180.
6- Nakajima et al., 1970, p. 246.
7- Ishimoto, 1971, p. 385; M. leonina specimens not included in this study.
8- Ishimoto, 1972b, pp. 339, 341.
9- Ishimoto, 1972b, p. 342; cf. Prychodko et al., 1971, p. 176.
10. Tashian et al., 1971, p. 190.
!!• Ishimoto et al., 1970, p. 235, phenotypes determined electrophoretically;
hemoglobin polymorphism in M. n. nemestrina has subsequently been
detected chromatographically (Nute and Pataryas, 1974a, p. 21).
(DeSimone, Linde, and Tashian, 1973, p. 55; Magid et al., 1973, p.
157; DeSimone, Magid, and Tashian, 1973, p. 165).
Blood protein data published for subspecifically indeterminate
pigtails of unknown geographic origin are generally concordant with
data summarized above for subspecifically identifiable specimens. In
40 adult pigtails of unknown geographic origin studied by Nute and
Stamatoyannopoulos (1971a, p. 145; 1971b, p. 109), hemoglobin
phenotype is monomorphic, which indicates that these specimens
probably were M. n. nemestrina (see table 10). In 75 pigtails studied
by Crawford (1966, p. 398), the frequency distribution of hemoglobin
phenotypes is 57.3 per cent F (type 2 of Crawford), 26.7 per cent FS
(types 3 and 4), and 16.0 per cent S (type 1), which suggests that
this may have been a mixed group of M. n. leonina and M. n.
nemestrina, approximately as later suspected by Crawford (1971, p.
706; cf. Nute and Pataryas, 1974a, p. 17). In a general review of
ABO blood groups in primates, Moor-Jankowski and Wiener (1971,
p. 240) report that B and O are characteristic of pigtails
(subspecifically indeterminate), which is compatible with the
48 FIELDIANA: ZOOLOGY, VOLUME 67
observed preponderance of agglutinin anti-A in M. n. leonina and
anti-A + B in M. n. nemestrina (table 10). Transferrin poly-
morphism reported in subspecifically indeterminate pigtails
(Goodman et al., 1965, p. 885; Goodman, 1967, p. 8; Prychodko et
al., 1969, p. 104) is likewise compatible with transferrin poly-
morphism in identifiable M. n. leonina and M. n. nemestrina (table
10).
In a captive multi-male colony of 16 pigtails of unknown
geographic orgin, nine blood proteins were studied in an attempt to
determine paternity of five infants (Simons and Crawford, 1969, p.
255). Since hemoglobin phenotype was monomorphic, the colony
probably consisted exclusively of M. n. nemestrina (see table 10).
This colony also was monomorphic for phenotypes of phosphogluco-
mutase (PGM-M), acid phosphatase (pm), erythrocyte glucose-6-
phosphate dehydrogenase (GdA), erythrocyte 6-phosphogluconate
dehydrogenase (6PGDA; cf. table 10) and haptoglobin (Hpl). The
same colony was polymorphic for phenotypes of serum pseudocholi-
nesterase (E«; E.s, Eiu, hypothesized), transferrin (TfA, TfB, TfD),
and blood groups (14 reagents). In a mixed group of M. n. leonina
and M. n. nemestrina, MN and Lewis blood groups were studied by
Nakajima et al. (1970, p. 248); 39 individuals tested were positive for
M or M-like antigens, 236 tested were negative for Lea antigens, and
12 of 123 tested were positive for Leb antigens. In 13 pigtails of
unknown geographic origin, erythrocyte isoantigen polymorphism
was studied with taxonomically inconclusive results (LaSalle, 1969,
p. 120). The amino acid composition of the f$ - hemoglobin chain in
M. n. nemestrina has been compared with that in M. fascicularis,
M. mulatto, and M. fuscata (Nute and Pataryas, 1974b, p. 79); the
composition of this chain in M. n. leonina is unknown.
In Indochinese populations of M. n. leonina, M. mulatto, and
M. fascicularis, similarities in frequency distributions of transferrin
phenotypes and 6-phosphogluconate dehydrogenase phenotypes
have been construed as evidence of interspecific gene exchange
(Goodman, 1967, p. 9; Prychodko et al., 1969, p. 108; Prychodko et
al., 1971, p. 181). M. arctoides (-M. speciosa of authors) formerly
was also included in this hypothetical group of interbreeding
"semispecies" (Goodman, 1965, p. 255; Goodman et al. 1965, p. 886),
but in recent reports this macaque apparently is regarded as a well-
isolated species (Prychodko et al., 1969, p. 107; Prychodko et al.,
1971, p. 179). However, regardless of the disposition of M. arctoides,
available evidence does not support the assumption that similarity
FOODEN: LIONTAIL & PIGTAIL MACAQUES 49
of blood protein phenotype frequencies necessarily implies gene
exchange. Transferrin phenotypes, for example, are more similar in
Thai M. n. leonina and Thai M. mulatto (index of dissimilarity
0.360; Prychodko et al., 1969, p. 107), which are members of two
different species, than in Thai M. mulatto and northeast Indian M.
mulatta (index of dissimilarity 0.546), which are local populations in
one continuously distributed species. Since gene exchange usually is
assumed to be greater within species than between species, it
follows that similar blood protein phenotypic frequencies are not a
reliable indicator of gene exchange.
If interspecific gene exchange has occurred in Indochinese
macaques with similar transferrin and 6-phosphogluconate dehy-
drogenase phenotypes, this probably should be also be evident in
non-hematological characters of these specimens. However, no such
non-hematological evidence of gene exchange has been observed or
reported in these critical specimens. In fact, there is no indication
that other blood protein phenotypes in these specimens support the
hypothesis of gene exchange, and one published analysis of their
phenotypic frequencies of alphai - antitrypsin definitely fails to
demonstrate such gene exchange (Omoto et al., 1970, p. 226). As
previously indicated by Jolly (1971, p. 197), it seems likely that
reported similarities in blood protein phenotype frequency dis-
tributions in Indochinese macaque species probably are the result of
common inheritance and/or convergent evolution, not gene ex-
change (cf. Weiss et al, 1973, p. 225).
Frequency distributions of blood protein phenotypes also have
provided the basis for a computer-generated dendrogram of
hypothetical interrelationships among macaque species and in-
fraspecific populations (Goodman, 1971, p. 83; cf. Weiss et al., 1973,
pp. 220-223 and Ishimoto, 1973, p. 9). (It should be noted that this
dendrogram is based on the implicit assumption that similar blood
protein phenotype frequencies are the result of common inher-
itance, not of gene exchange.) Judging from the dendrogram,
hematological evidence apparently indicates that M. n. leonina and
M. n. nemestrina diverged from the common ancestral macaque
lineage subsequent to the divergence of Celebesian macaques and
that M. mulatta diverged subsequent to the divergence of M.
fuscata. However, these phylogenetic inferences are contrary to
those derived from non-hematological characters, which indicate
that short-tailed Indochinese M. n. leonina and Sundaic M. n.
nemestrina probably represent the stock that is ancestral to (hence
50 FIELDIANA: ZOOLOGY, VOLUME 67
diverged prior to) stump -tailed Celebesian macaques (fig. 18c;
Fooden, 1969a, p. 61) and that short-tailed continental M. mulatta
probably represents the stock ancestral to stump -tailed insular M.
fuscata (Fooden, 1972, p. 310). Judging from non-hematological
characters, it is likewise difficult to accept the hematologically
supported assertion (Goodman, 1971, p. 86) that "human beings are
very generalized mammals"; compared with Mesozoic stem mam-
mals and with primitive marsupials and insectivores (Romer, 1966,
pp. 193-211), humans obviously are highly specialized, except for
retention of pentadactyly (cf. Clark, 1972, p. 473).
COMPARATIVE ECOLOGY AND BEHAVIOR
As discussed in detail in species and subspecies accounts (see
below), liontail macaques and pigtail macaque subspecies appear to
be essentially similar in habitat, diet, and troop size. The usual or
exclusive habitat of liontails and pigtails is dense evergreen rain
forest, and their geographic distribution closely coincides with the
distribution of this habitat in tropical Asia (fig. 17). The food of
these macaques evidently consists chiefly of fruits and seeds,
occasionally augmented by young tender leaves, insects (larvae and
adults), spiders, and, probably, birds eggs and small vertebrates.
Troop size in these macaques typically is about 5 to 20. Most troops
probably are composed of 1-3 adult males, 2-10 adult females, and
2-10 immatures. Solitary adult males are fairly common; this
presumably is correlated with the unbalanced ratio of male and
female adults within troops.
Altitudinal distribution is not uniform in liontails and pigtail
subspecies (table 11). Of the three relatively well-known species or
subspecies, M. silenus, with known altitudinal range 300-1300 m., is
restricted to upper elevations; M. n. leonina, with altitudinal range
75-1250 m., is limited to the vicinity of foothills of major mountain
chains; and M. n. nemestrina, with altitudinal range 0-1700 m., is
relatively widespread and inhabits lowland areas as far as 150 km.
distant from the nearest mountains. These differences in altitudinal
range probably are determined by regional temperature and rainfall
patterns (Koeppe and DeLong, 1958, pi. 2), which, in turn,
determine rain forest distribution (fig. 17).
Liontails and pigtail subspecies also appear to differ in
population density and arboreal-terrestrial preference. Population
density in M. silenus seems to be much lower than in M. n. leonina
and M. n. nemestrina, judging from frequency of reported
encounters. M. silenus and M. n. leonina apparently are much more
arboreal than M. n. nemestrina; M. silenus and M. n. leonina are
rarely encountered on the forest floor, whereas such sightings of M.
n. nemestrina are commonplace. When alarmed, M. silenus and M.
51
-■'
"*>,
£"8 §
« "§'5
8-S8
O a)
.9
3!
~ j= jj
u C 3
.2 * -
— = =
SO"2
g i =
•p >. ^
« -I
3- — "*
08 J a
O 00 ♦<
.sa:
|8
c «2
U
^ 00
g -
<*. 5"
o «
So
8
9
.a o. «.
-o
-a
~Z c
e «
4) —
0
o
St>
52
FOODEN: LIONTAIL & PIGTAIL MACAQUES 53
TABLE 11. Altitudinal distribution of accurately known localities of lion-
tail and pigtail macaques. Locality numbers (italicized) as in Figures 20, 25,
28; for details, see Gazetteer of Collecting Localities.
Species or subspecies; locality numbers
Altitude (m.)
M. silenus
M. n. leonina
M. n. nemestrina
M.
n. pagensis
1600-1799
—
—
82
-
1400-1599
—
—
—
—
1200-1399
—
6
—
—
1000-1199
7, 10,
13,
17
—
16, 75
—
800-999
—
8, 32
66
—
600-799
—
30
16, 17, 25
—
400-599
8
—
43, 82
—
200-399
1
21, 22, :
23,
25
50, 56
—
0-199
—
1, 2, 29,
31
, 46
11, 44, 46,
88
92
, 93
n. leonina usually flee into the canopy, while M. n. nemestrina
typically descends from the trees and flees on the ground.
Except for M. n. pagensis in Kepulauan Mentawai, the
geographic ranges of liontails and pigtail subspecies overlap those of
several other species of macaques. M. silenus inhabits forest tracts
in southwestern India that also are inhabited by M. radiata. The
combined range of M. n. leonina and M. n. nemestrina broadly
overlaps that of M. fascicularis; however, these species are at least
partly segregated ecologically by the preference of M. n. leonina
and M. n. nemestrina for inland evergreen rain forest and that of M.
fascicularis for marginal mixed secondary growth habitats. The
same ecological segregation may also apply to the relationship
between M. n. leonina and M. mulatto where their ranges narrowly
overlap (cf. below, fig. 28, and Fooden, 1971a, fig. 2). The combined
range of M. n. leonina and M. n. nemestrina narrowly overlaps that
of M . arctoides, which has been collected in the same forests as M.
n. leonina. Although the range of M. n. leonina is almost
completely allopatric with that of M. assamensis, these species have
been collected together in a few forest tracts along the frontier
between their ranges.
HYBRIDIZATION
Information available concerning hybrid matings that involve
lion tails or pigtail subspecies indicates a relatively low level of
hybrid viability and fertility (table 12). Of five reported intergeneric
matings, one failed to produce conceptions, three produced hybrids
that died in infancy, and one produced a hybrid that survived less
than two years. Of 48 interspecific and intersubspecific hybrid
conceptions reported, at least nine resulted in stillbirths or
produced young that died in infancy; 11 of the hybrids are known
to have reached the age of sexual maturity, and only three of these
are known to have produced living offspring.
All intergeneric crosses that involve liontail or pigtail macaques
are restricted to members of the 42 chromosome group of
cercopithecine genera (Chiarelli, 1967, p. 162). All interspecific
crosses are restricted to macaque species in which the glans penis is
bluntly rounded and the uterine cervix and cervical colliculi are
moderately large (see above). Because of the size difference between
sexes in catarrhine monkeys, it is not surprising that most hybrid
matings of M. n. nemestrina with smaller-sized monkeys {M.
silenus, M. n. leonina, M. fascicularis, Cercocebus sp.) are between
a female of M. n. nemestrina and a male of the smaller species or
subspecies; conversely, all four crosses with larger-sized species
(Papio spp., Mandrillus sp.) involve a male M. n. nemestrina and a
female of the larger species. The high incidence of hybrid matings
reported between pigtail macaques and either M. fascicularis or M.
mulatto may merely reflect the fact that M. fascicularis and M.
mulatto are the macaque species most frequently kept in captivity.
Available evidence concerning Fi phenotypes is limited and of
varying precision and reliability (table 13). This deficiency is
unfortunate because such evidence would be useful in interpreting
the genetics and evolution of taxonomic characters in liontail and
pigtail macaques. Although relevant data are somewhat ambiguous,
the following genetic inferences are probably valid: (1) dorsal pelage
color in M. fascicularis is dominant to that in M. silenus and M. n.
54
co s
I
I
-
co
o
•-1 _«'
o
I— 1
rH D" m
a
lO ._- .
CO
CO
^ 32 i* <->
<7>
-' - 3J _
rH
oo .S
si oo -w -3
52
0 S
c o
m n * m
^ ; h A A 2
OS C
a> . a
«J ^cvf oo
So*0?
_ Oi ^ <N
•3 f-H * •
_ „ >> m
a
CM* DO
Oi CO
--1 CN
«~ o
~ CO
3 rH CN
CN 00 tN *
in <? m CN
C75 CO DTi CD
>-• CN --1 CN
& a
S2
>> in
.S 3
t/i
"£ "5 ? DM
MSNh
>> in >> in
ig<N-*CN
£ CN S CN
0 CN O CN
.* , -X .
3 Oi 3 CT5
N >-" N rH
^ t-
_; a
o
0
N
0 rH
-I
.8*3
Si a>
O DC
CO
0)
rH 0
, o
J Jh
O O rH CN
Q Q
jr. —
co co
CO CO
a a
"o "o
o o
N N
z z
o
3
a
a*
T3
-a
c
CO
oe
e
C8
0)
3*
c
(0
c
<M
Ih
'*>
0
'•2
'5
o
>>
0)
c
C8
E
c
0
>
B
*->
C
V
o
B
n
'C
4)
CX
a
3
0
E
M
e
je
O"
CJ
^
B
c
V
0
•—
c
>
-
0
B
a
>
e
(0
a
0
■—
H
B
CD
a
3
0)
T3
B
o
E
o £
£« ^ a
c: -3 <2 >, g
o
o
£
£
o
o
6
E
rH
rH
*->
C
fca
rH
Iw
C
>>
>>
— >>
— "*-
m
r}<
>>
5 ^
3 .E
A
A
c8 O- ' N C^-
ca A
^ w o
^
>.
J=
0
a
91
X
Oh
CN
to
o
(0
•5
CO
rJ
-
<
a
to
"3
B
CD
>-
5
0)
a
co
—
3
CO
CM
E
CU
C
B
B
a
C
5,
■ ■_ -c
°* ^^ s
S o r- ft)
•S c : jj
i; 2 c ^
a, a, 5 O
k.l.k V. t. k ^ ^- -i
" Q) Q) ^>
co C C C
3
c c c c
co ^ I? ^ ^ ^ ^
co S "^ f.
Cv c« '3
O « W W
•:;
••
a
a
r
B
B
B
C
C
U
«
ai P aj ai c
alal
a o
oj
CN
CO CO -i
*•
OJ tO <JD
a a„ ^
<n qVcn
CO* Q. "* CO* Q. ^
l-H 1— 1
^H . „•■« . . •■«
Qfi &C
oo' oo* n «
(OCC5 a
oj a 5
- OJ -
CO 0* ^h ■ OC ,- •
C t>- 0. c ^ n.
2 . oj * . oj *
9, ft
9, fi
S S oo '-1
971
li, 1
971
°* CO . »*• CO . »
w <N "5 0 ^h — OJ ^, — OJ
m cn m cn
tf> ^ *n ,_
.—i •— <
1. Anver
1. Anver
34, p. 4
B, p. 141
Gray, 1
<c n H il n — i t-i — - .— 1
CO . oi • ti
^ ac ac
1- _c >_
aao'ia »^ 2 £ ^ 2
a to* ^r 50* ph*
OOO
SE;2o2»h c c ^ c c
- 50 CN "g CN
OJ OJ _- OJ _h
0 0 oj © c
c c c
CN ^H ^H
0 0 '-' OJ '**
'*• '*• **
OJ 00 *^ 0> OJ ~ „•«••■• . •- •-
oj - d _r d
N N -r -• C
c c c
** '-> g- ^h --1 <jj c c c c c c
. E - a*
S S 5
j" * \ac - • h "ai 'ai a* 3 S 01
o-£c.±£.*ccceec
qoi^.s ,j;o>a>o>o>o>o>o>
a,UStf&.aCCO.CQCQCQCQCQ
i«* '3 00" '3 00
01
Be OS ^ ^* "ti
■****« j^ 4) C«
— — —
c c c
o> *i to ti to
1 g* Soj
0 5 « - »
1- )- u
— - - -
Ct, OQ co CO to
e
o>
la
V
<—
0) 0> 0>
— — —
o>
TfTf lOIOf
C- 00
00 h n n f m m
to t^ r~'
pes
«-t »-H 1-H .-H 1— •
•-H H
H N CN (N N (N CN <N
CN CN CN
■•-> 0*
> .s
o> ■-
on a
c J5
o fc
J o
£ E
5; c £ « * -
-s ^ — :z 3 « «
■a m 3 * £ 1> >
o
£
X
•5,
CN -o
h CO •-» e^- c- to
0 **
= «2 >»
*» c
</i
M
C
*o '*"
01 <*-
CO o
6
S
*o c- *o *o C«- C-- °*
S3
2
.5-
S 2
*o o» »o *o *o *o *o
'< « it it it d m
1
r^
v.
_2
1^
41
41
4j
r
^
E
«
*o
^^
o»
3
"3
^
"5
"3
"2
~5
c
c
a
,_
_.
_
_
.tj
u
i;
s;
,jj
u
^
fi
i
3
E
c
5
e
CJ
5
z
s
C
C
0
E
^t
.^5
rf
•S5
<0
■r.
■r-
M
a
<A
M
a
a
C3
C3
3
a
a
e
tj
r •
c
c
c
c
«»i
<
«s
•fi;
«fi;
v~
*;
55
55
55
5555555
CO CO
CO <n .*
■2 a a
3 u .0
ai 5
56
ti
<
.2^
U CA
— 'G
CA j)
CO 1*
- o
. <*>
co
:S
<N '"' 0<
o> Q« cL
(N - "•
'a
C
4rf
1963, p.
, p. 304
4; 1971
106
44
968, p.
Hi, 1971
V, 1971,
,9
T3
CA
eg
t- in •
• *-> a* ca
■ CO <N t> O,
a . i- u
x
£> <JJ . i-t -
~r jj -2 o
->->
g .- . .in
°> *- J= ~
0
* «n,S
c
Sh 00 O a 35
■^ 00 t- .»H
rt >H .C C
0
r co a> <n •>
- . !! 'S
Zool
6, p.
20; 1
; 197
onius
turkh
Zool
nsteir
lernst
CA
h
0
n3 iH a r: < Z — — -
3
ca
g
e
00 a>
O i-i (N
i
tN CN
CO CO CO
CO
M
a
"C
e
CO
0
tN
-
0
00
c
O*
•—
'la
IfJ
o
a
„
X
CA
«o
CO
Cm
0
-1
o
<->00 05
._ eO CO o-
t>- *o t>- o*
o a a a
0005
"3 "3 "3 3
ESSE
5-
<D
X>
>>
<o
ECA
0)
ti "5
2 a
&!
— «j
OB jS "O
a .2" o
■£ o >-
« £ 2
e* -> O
C cu n
- g -o
2 j= 2
E g-§
8 * |
J=J5d
o
W
>>
c c
«V . O
0) > c
0 $ a
- .O J5
•5 - O
.!s O a,
£ c «
3 -3 E
CM
a £
£ C hi
-c
o
v.
a
c
0 _a>
1 "
c a
• o s
1 Ql "-*
t ^
, >- a
&*<
5 x
o
Hi
J2 3
£ E
1^
CO CO ^j >t ^
r> > Xi CO
i s
© 2
3 t H t
CO «
2- E
x £ *"-
W i ri
X J3
X X
2 E
c ^
o >>
1-1 tNCO^ lO (C t» 00
O "U
o o
S2 &
ca ca
JZ ea
3 ?
Us
£•*
CA X
57
0
Z
•m
0>
a:
»-< —i i-h OJ CM CN <N
- I
c ss
S
0>
2
c
.0
CO
c
j>
o
i
s
o
•3
m
.a
tj
"E
X-
>>
X
I
>.
—
0
c
o>
—
a
2C
<
a
>>
o
c
HI
a
— 00
0)
2 «
2 c
■S'i
C 0)
c a
c
K I I
e 'g.S
°1
u 0)
01
01
I I I
*-> c -> c
W ce W eo
SO c <0 c
>- .— «- ._
I l-S 1
sj 0 O C
. TJ . "O
e j e j
c a c a
I I I
oc —
t A O
2 "3 "o
Q a o
I I I
0
01
ex
CO
■o
CO
•
0>
a
JS
c
e
■•J
G
5
e
c
JS
Js
js
O
o
-
'—
<w
SO
CO
CO
9
a
a
—
U
c
c
e
«
■*
S
*
S
0
0
o
I I I
c
Z
0
(J
■z
° a
.5 5
* o
a 'u
« a
CO
0)
«J c
3
s *
"55
1-5
5-f
•i c
i» —
cu 9
I I I
.*» .2 IT
IT C _ej o»
P -Si a £
c c . c
.0 J3 "*»
55^5
CO ~
:= c
.§«
o >,
>> 01
o£
c <~
0» O
*i
- w
•D O
'Z c
.Q 0)
x a
4) 0)
0> -
1.1
c :
M c
0) CO
a'>
3 X
oi n,
a 2
</> CO
X5 «
OJ -o
C X
c o
a
>.
o
^ c
9 %
2 a
a iS
c
a oi
?]
x Oi
o>
a —
>> 2
o c
c £ * x
Ut*C
c c
'C X
a >>
o"S
5 E
*- 3
>.£
x a
o S
b. H
a o 5
58
FOODEN: LIONTAIL & PIGTAIL MACAQUES 59
nemestrina; (2) tail length and carriage in M. n. nemestrina is
dominant to that in M. silenus; (3) tail length in M. fascicularis is
not dominant to that in M. silenus and M. n. nemestrina (hybrids
intermediate). The number of genetic loci or alleles that underlie
each of these characters is uncertain.
FOSSIL RECORD
All known fossils and subfossils that are referable to the
liontail-pigtail group have been collected in Quaternary deposits in
the Malay Peninsula and Greater Sunda Islands, in or near the
range of living M. n. nemestrina. In the Malay Peninsula, macaque
subfossils are part of human food remains collected in floor deposits
in a rock shelter (Gua Madu) in southern Kelantan, West Malaysia.
According to Chasen (in Tweedie, 1940, p. 7), these food remains,
which are not individually described or listed, include bones and
teeth of M. n. nemestrina, langurs (2 spp.), bamboo rat, Malay bear,
wild pigs (2 spp.), barking deer, sambar deer, and wild ox; no
specimens of M. fascicularis are reported. Associated Hoabinhian
artifacts indicate that the floor deposit in this shelter was formed
about 4000-10,000 B.P.
In the Greater Sunda Islands, identified macaque subfossils and
fossils consist exclusively of fragments of jaws and teeth. Specific
determination of these specimens depends essentially on tooth size.
Based on macaque species and subspecies that now inhabit the area,
subfossils or fossils with large teeth are identified as M. n.
nemestrina, and those with small teeth are identified as M.
fascicularis. Most specimens with intermediate size teeth are
indeterminable.
In west-central Sumatra, 124 isolated subfossil teeth have been
collected in three caves (Djamboe, Lida Ajer, Sibrambang) in the
Padang highlands. Of these, 102 are identified as M. n. nemestrina
(82 per cent), 7 are identified as M. fascicularis (6 per cent), and 15
are indeterminable (12 per cent) (Hooijer, 1962a, p. 57); species
proportions are reasonably consistent in all three caves. The age of
these cave deposits is estimated to be prehistoric Holocene (Hooijer,
1962a, p. 4). The agency responsible for assembling the large
accumulations of teeth of macaques and other mammals in these
caves is unknown (Hooijer, 1948, p. 187).
In Borneo (Niah Caves, northern Sarawak) 76 identifiable
subfossil macaque jaws and teeth have been collected in strata with
60
FOODEN: LIONTAIL & PIGTAIL MACAQUES 61
TABLE 14. Stratification of M. n. nemestrina and M. fascicularis subfossils
collected in Niah Caves, northern Sarawak (Hooijer, 1962b, pp. 441-443).
Depth below Approximate Number of subfossils
surface (inches)l C14 age (B.P.)2 M. n. nemestrina M. fascicularis
0-24 0 -10,000 4(16%) 21(84%)
25- 49 10,000-20,000 0 37(100%)
50- 74 20,000-30,000 0 7(100%)
75-102 30,000-40,000 1(17%) 5(83%)
Totals 53 (7%) 70 (93%)
1- Specimens tabulated according to midpoint value of depth interval reported.
2- See Harrisson, 1958, p. 563.
3- Excludes one specimen without stratigraphic data.
Carbon 14 dates extending back to about 40,000 B. P. (Harrisson,
1958, p. 563). Of these subfossils, six are identified as M. n.
nemestrina (8 per cent) and 70 as M. fascicularis (92 per cent)
(Hooijer, 1962b, p. 440). M. fascicularis specimens strongly
preponderate at all stratigraphic levels (table 14). All of these
specimens apparently are human food remains (Medway, 1964, p.
34).
In eastern Java 28 fragments of macaque jaws and teeth have
been collected in early Holocene deposits in three caves (Gua
Djimbe, Gua Ketjil, cave near Wadjak; Hooijer, 1962a, pp. 41, 54).
All tooth dimensions in these specimens closely correspond to those
in M. fascicularis mordax, the living Javan subspecies of crab-
eating macaque (tables 16, 17). The agency responsible for
accumulation of these subfossils is unknown.
The composition of subfossil macaque faunules in the Sunda
area is compared with that of modern faunas in Table 15. In
TABLE 15. Relative abundance of M. n. nemestrina (n) and M. fascicularis
(f) in modern faunas and in subfossil faunules (see text).
Age of fauna Malay Peninsula Sumatra Borneo Java
Modern /">n , On f>n fXn)1
0-10,000 B.P.2 ny(f)1 n>/" f>n
10,000-20,000 B.P.3 fXn)1 f>(")1
20,000-30,000 B.P. fXn)l ~
30,000-40,000 B.P. - />„
1- Species absent.
2- Includes prehistoric Holocene
3- Includes early Holocene.
I
8.
r-* op op
<n co co ifi
oS /\ to tb
3
^ co
JO
rr CD
CO
00
'^
00 00
OS
OS
<N
I'lfiMf
^H
CO
>■
ClXhW
00
r^
3
c
^ r^
oo
** m
rf O
I CO CO
e
cd
B
0
C
DC
i
4>
I
gg
'3
a.
c
41
a
e
is
a
3
0
■—
E
o
-*
^-s
4>
la
E
£
_
</>
a
c
01
-
£
£
jj
>-
a.
3
M
•r
«
0
^
II
-C
(C
£
=
u
=
~
(A
X.
D
<
M
H
o
~
B
OS o>
-h
OS
CO CO
t^
t^
OIOhiO
|H
CO
co t- co m
CO
CO
— <N
•-*
in
m
CO CO
CO
00
OS
m o
CO
in
m
in in
m
in
CO CO OS ^* C*f t^ CN
*<5 in t~- in c^ co oJ
o oT os t~- in tj> as co
I CO tT CO in rf CO tT CO
Ot ot*o »>o o» »o
CN ^, CS CO
CO ^ <N rf O ^
CO r- CO OS 00 *H
oocor~int~ds'^oo
os r^OTrcbint--cb'-'
A — -h
»o o» *o o» *o »► *o
> CO ^,"^
« JC > H
Of*
*j rrt •*-»
4) O 0 0 5
E 2 .2 I ?
8-§.SJ
SQS
a
5
m
a
.c
c
o
CO
a
c
* «
5^5 5
2 w
35
-r
OS
OS
OS*
SO
in
x
■x
~
■o
eg
C!
ft*
Cl
t*
0
(0
-x
99
99
OJ
pm
9J
4a
s
a
0
>-
la
'5
0
ac
—
OS
OS
til
41
B
in
iri
X
x
X
iC
2
m
a
a
X
a
a.
a
08
»
(9
• ■
es
M
~j
99
C>1
—
tN
~
-x
iC
CC
re
te
0
99
99
~
*r
99
—
pH
n
g
^
X
>
la
X
o
*
JO
.O
'5
"3
0
0
0
< ac oa i < x
^h CN CO TT m CO
62
0)
I
CO
a
•v
c
«
V
c
<u
o
O
O
X
v
c
HI
CJ
O
I I I „ I
c
n
C
u
-—
B
o
-*
. — .
d)
R
i-
6
|H
en
a
c
e
8
E
E
CJ
D
CD
V
a
a, O
£
t-
S
3
UJ
B
hJ
co
PQ
t)
<
09
h
73
B
C2 £x a
© „j CO ^ N 2 6 2
x — x — 9x"T x
ii ^ o t^ i ^ i (,j
t^cir-ait~ — x —
_ _ — rt
— ~ ~ t- cm 5T cm cr. x
•r ifi io M cp f ifl v t*
is in* tr: to en r* « ? i c t^
^. _ « _ v: m
cyio r~ © ^ crs ~ *r ^j
too*' Cs r~ — tf: CJs X ~
tC.t*'w*-wt*nocax
irtXCOt^-i^t—ifiCJ^incys
* O O 0 O
£ Z Z Z .2
- x j: X '-
£ 9 3 3 J
5DQOS
ai ? o
<u 0
as
O 'C
w 4)
CD »j
•*■> w
c o
CO Q,
II o
o
</i '«
C -
O *
'■3 x
n oi
J "o>
* I
< o>
^ cn co -<r in co
F
—
£
-.
tN
co
Cfl
(0
x
M
m
x
in
z
m
D.
Q,
a,
d
cs
O
eg
■ »
eg
tN
OI
-r
N
<D
Oi
co
W
CO
T.
05
•*■
01
P3
01
o>
01
00
01
:z?
;^
:^
q
c
:;
0
0
i)
-
0
I Q K < S
63
64 FIELDIANA: ZOOLOGY, VOLUME 67
modern Malayan, Sumatran, and Bornean faunas, M. fascicularis is
approximately 10 times more abundant than M. n. nemestrina
(below p. 103). In Java M. fascicularis is the only macaque now
present (Chasen, 1940a, p. 65). Subfossil faunules collected in
Borneo (late Pleistocene-subrecent) and Java (early Holocene) are
similar in composition to the respective modern faunas; those
collected in the Malay Peninsula and Sumatra (prehistoric
Holocene) are not. Taken at face value, the Malayan and Sumatran
subfossil data would indicate that, within the past 10,000 years, M.
n. nemestrina was more abundant than M. fascicularis in the
western part of the Sunda area, which is the reverse of the present
situation in this area. However, this indication of recent local
reversal of relative abundance may be misleading, if the accumula-
tion of macaque remains in the Malay Peninsula and Sumatra has
been subject to some unknown selective bias (cf. Hooijer, 1962b, p.
447). Hopefully, further collecting in Sunda area caves and rock
shelters may help to resolve this intriguing and important problem
in macaque paleozoogeography.
In addition to Holocene subfossils, mid-Pleistocene fossils that
are referable to Macaca also have been collected in eastern Java
(Bangle, Punung, Sangiran, Saradan, Trinil; Deninger, 1910, p. 1;
Stremme, 1911, p. 140; Badoux, 1959, p. 88; Hooijer, 1962a, p. 50;
Hooijer, 1964, p. 76). These Pleistocene fossils, some of which
originate in fissure deposits with those of Homo erectus, are among
the oldest known macaque remains in Southeast Asia. Dental
measurements in 14 recorded specimens (tables 16, 17) are not
completely consonant with measurements in either living Javan M.
fascicularis mordax or living Sumatran and Bornean M. n.
nemestrina (and still less with those in living Sumatran and
Bornean M. fascicularis; Hooijer, 1962a, p. 58). Although dimen-
sions of anterior teeth in these Pleistocene Javan fossils are
approximately the same as in living M. f mordax (and in Holocene
subfossil M. f mordax', tables 16, 17), breadth of the last premolar
and all three molars is nearer to that in M. n. nemestrina. In the
best preserved Pleistocene specimen (Dub. No. 3785, see table 16),
breadth of the last two molars is 8-9 per cent greater than in the
largest measured specimens of M. f. mordax and well within the
range of molar breadths in M. n. nemestrina.
The relationship of the Pleistocene Javan fossil macaque to
living M. n. nemestrina and M. f. mordax has been carefully
reviewed by Hooijer (1962a, p. 51). In interpreting these data,
FOODEN: LIONTAIL & PIGTAIL MACAQUES 65
Hooijer notes that the ratio of premolar breadth to molar breadth
in the fossil macaque is less than in living M. n. nemestrina and
postulates that this would not be true in the Pleistocene ancestor of
M. n. nemestrina. Accordingly, he rejects a relationship to M. n.
nemestrina and concludes that the fossil macaque probably is an
ancestral form of M. f. mordax, in which the premolar-molar
breadth ratio was greater than in the living form.
However, the a priori assumption that the ratio of premolar
breadth to molar breadth in the Pleistocene ancestor of M. n.
nemestrina would not be less than in living M. n. nemestrina may
not be valid. The relatively broad premolars in living M. n.
nemestrina may be a recently evolved condition that is related to
allometric increase in size of the rostrum and anterior part of the
dental arch (figs. 8, 9). Cranial and dental characters in the stock
ancestral to M. n. nemestrina may have been similar to those in M.
n. leonina, which has tooth proportions that are fairly close to
those in the Pleistocene Javan fossil macaque (tables 16, 17).
Available evidence permits two plausible interpretations of the
Javan fossil macaque. This Pleistocene macaque may be ancestral
to Holocene and living M. f. mordax, in which case relative
premolar breadth in this line has undergone evolutionary decrease;
this interpretation implies that the pigtail stock was absent from
Java during the Pleistocene, as well as during the Holocene and
present. Alternatively, the Pleistocene macaque may be ancestral to
living M. n. nemestrina, in which case relative premolar breadth has
undergone evolutionary increase from proportions similar to those
in living M. n. leonina; this interpretation implies that the pigtail
stock inhabited Java during the Pleistocene and subsequently
became locally extinct, as did the siamang, orangutan, Malay bear,
elephant, and tapir (Badoux, 1959, p. 129).
65
EVOLUTION AND DISPERSAL
Liontails, pigtail subspecies, and Celebesian stumptail species
are allopatric and apparently constitute a closely interrelated
phylogenetic unit, the silenus-group. (If subgeneric recognition of
this group is warranted, the tautonym Silenus Goldfuss, 1820, is
available and valid.) The close relationship between liontails and
pigtails originally was inferred from similarities in pattern of hair
growth, morphology of female sexual swelling, and structure of tail
(Pocock, 1926, p. 1571). The relationship between Sundaic pigtails
(M. n. nemestrina) and Celebesian stumptails was inferred from
evidence of external proportions, structure of baculum, and
geographic distribution (Fooden, 1969a, p. 61). The present study
further indicates that species and subspecies in the silenus-group
are similar in form and proportions of the glans penis (above, p. 33);
these macaques also are similar in general form of estrous sexual
skin swelling (above, fig. 11; Pocock, 1926, p. 1547; Fooden, 1969a, p.
13). Judging from penial and cervical morphology, the silenus-group
probably is more closely related to M. sylvanus and to the
fascicularis-group (M. fascicularis, M. mulatto, M. cyclopis, M.
fuscata) than to either the sim'ca-group (M. sinica, M. radiata, M.
assamensis, M. thibetana) or M. arctoides (above, pp. 32, 33).
The close relationship of species in the silenus-group evidently
is not revealed by their chromosome morphology. Karyologically,
Af. silenus and M. nemestrina are less similar to each other than
each is to other macaque species (Chiarelli, 1962, pp. 408-412;
Schmager, 1972, p. 481).
Geographic distribution of species and subspecies in the silenus-
group (fig. 180 closely corresponds to major subdivisions of the
Oriental faunal region (Sclater and Sclater, 1899, p. 131, pi. 5). The
range of M. silenus is in the Indian Subregion, that of M. n. leonina
is in the Indochinese Subregion (extended to include the Isthmus of
Kra), and that of M. n. nemestrina is in the Sundaic Subregion (for
name of this subregion, see Johnson, 1964, p. 69; Bullock, 1964, p.
212). The insular range of M. n. pagensis in Kepulauan Mentawai is
66
FOODEN: LIONTAIL & PIGTAIL MACAQUES
67
Present group
Fig. 18. Hypothetical stages in evolution and dispersal of species and subspecies
in the silenus-group.
at the edge of the Sunda Shelf, marginal to the Sundaic Subregion.
The range of Celebesian stumptail species is in the transition zone
between the Oriental Region and the Australian Region.
The zoogeographic hypothesis presented below (schematically
illustrated in fig. 18) is proposed as a first approximation of the
Quaternary history of the silenus-group. Obvious problems in
zoogeographic interpretation are posed by deficiencies in the fossil
68 FIELDIANA: ZOOLOGY, VOLUME 67
record of this group (see above, p. 60) and by incomplete
information concerning Pleistocene changes in the geography,
climate, and vegetation of the Oriental Region. Major topics
considered in the following discussion are:
1. Center of origin of si/enus-group; main pattern of dispersal.
2. Disjunct distribution of liontails and pigtails.
3. Dispersal of silenus-group to zoogeographically peripheral
Kepulauan Mentawai and Celebes (Sulawesi).
4. Geographic contiguity and morphological discontinuity
between M. n. leonina and M. n. nemestrina.
5. Geographic isolation and morphological similarity within M.
n. nemestrina.
In species and subspecies of the silenus-group there is a clear
geographic pattern of progressive decrease of tail length from west
to east. Relative tail length is 0.57-0.75 in M. silenus, 0.28-0.46 in M.
n. leonina and M. n. nemestrina, 0.24-0.34 in M. n. pagensis, 0.05-
0.12 in southern and central Celebesian species, and 0.03-0.07 in
northern and northeastern Celebesian species (fig. 2; Fooden, 1969a,
p. 15). Assuming that a long tail is primitive in macaques, as in
other monkeys, this west-to-east trend may be interpreted as a
trend from primitive to derived. Similarly, the relatively short
rostrum in skulls of M. silenus and M. n. leonina probably is more
primitive than the long rostrum in M. n. nemestrina (fig. 8); the
most specialized skulls in this group, or in any species of macaque,
are those of M. nigrescens and M. nigra in northeastern Celebes
(Fooden, 1969a, p. 30). The geographic pattern of morphological
variation of skull and, especially, of tail length tends to suggest
that, among living species and subspecies, Indian M. silenus is
morphologically most similar to the ancestral stock of the silenus-
group and that Indochinese and Sunda area species and subspecies
represent successive stages in evolutionary development of the
group. Accordingly, the starting point of the present hypothesis is
that the silenus-group originated in the Indian Peninsula (fig. 18a)
and subsequently dispersed to the Indochinese Peninsula and the
Sunda area (fig. 18b).
The initial direction of dispersal from the Indian Peninsula
presumably was northward (fig. 18b). This may have given impetus
to tail reduction in the silenus-group, since higher latitudes are
known to be correlated with shorter tails in two previously studied
groups of macaques (Fooden, 1971b, p. 72; 1972, p. 310). However,
FOODEN: LIONTAIL & PIGTAIL MACAQUES 69
relative tail length in the unknown original progenitor of the
silenus-group probably was greater than in M. silenus (0.57-0.75),
judging from external proportions in most monkeys (Napier and
Napier, 1967, p. 413). M. fascicularis, which is the only living
macaque with a long tail (relative length 0.79-1.48; Fooden, 1971a,
p. 27) and a rounded glans penis (see above) may serve as a
morphological model of the unknown pre-silenus ancestor of the
group.
The present range of M. silenus in southwestern India is
separated from the western border of the range of M. n. leonina in
Assam and Burma by a gap that is about 2000 km. broad. At
present, natural vegetation in the intervening area is deciduous or
semi-deciduous forest (Kuchler, 1964, pp. 17, 141), which evidently is
not suitable habitat for the si/enws-group (fig. 17). Past dispersal of
the silenus-group across this area, as assumed in the present
hypothesis (fig. 18b), probably implies that the area formerly
supported evergreen forest, which, in turn, implies more continuous
rainfall than at present.
It seems likely, therefore, that dispersal of the silenus-group
from the Indian Peninsula to the Indochinese Peninsula occurred
during a Pleistocene pluvial interval (fig. 18b), and that subsequent
disjunction of Indian and Indochinese sections of the group dates
from a succeeding arid interval (fig. 18c). The same series of
Pleistocene climate changes probably also accounts for similar
disjunct distributions in Indian and Indochinese species of langurs,
lorises, giant squirrels, civets, and mouse deer (Pocock, 1939, p. xxvi;
cf. Kurup, 1966, p. 19). Analogous zoogeographic interpretations
based on Pleistocene rainfall variation previously have been
proposed to account for distributions of tropical bird species in
Southeast Asia (Stresemann, 1939, p. 416) and South America
(Haffer, 1969, p. 131). Recent human interference with natural
vegetation patterns probably has played a relatively minor role in
exaggerating disjunction of the ranges of M. silenus and M. n.
leonina.
Isolation of Indian and Indochinese sections of the silenus-
group presumably led to morphological differentiation of ancestral
liontails and pigtails. Populations of the ancestral pigtail stock
evidently dispersed to Kepulauan Mentawai and to Celebes, where
subsequent isolation resulted in differentiation of a stumptail stock
(fig. 18c). In Kepulauan Mentawai, M. n. pagensis is part of a
strikingly peculiar mammal fauna (Kloss, [1928], p. 806) that also
70 FIELDIANA: ZOOLOGY, VOLUME 67
includes an endemic species of gibbon, Hylobates klossii. Because
dispersal of gibbons evidently is strongly inhibited by water barriers
(Groves, 1967, p. 281; Fooden, 1969b, p. 640), it seems likely that
ancestral populations of the distinctive Kepulauan Mentawai fauna,
including ancestors of M. n. pagensis and H. klossii, reached this
island group by way of a transitory land connection with the
adjacent Sunda Shelf. Conversely, absence of gibbons from Celebes,
and the generally depauperate character of the Celebesian mammal
fauna (Darlington, 1957, p. 519), suggests that original dispersal of
the silenus-group to this island was across a water gap, probably by
rafting (Fooden, 1969a, p. 64).
Dispersal of the silenus-group to Kepulauan Mentawai and
Celebes probably occurred during maximal Pleistocene glacial
lowering of sea level, to about 200 m. below present sea level
(Jongsma, 1970, p. 151), when the Sunda Shelf was most extensively
exposed. Relative richness of mammal faunas in Kepulauan
Mentawai and Celebes evidently has been controlled by depth of
the straits that separate these insular areas from the Sunda Shelf.
Minimum depth of the strait between the Sunda Shelf and
Kepulauan Mentawai is about 200 m., and that between the Sunda
Shelf and Celebes is about 600 m. (Atlas van Tropisch Nederland,
1938, pi. 3). The absence of otherwise widespread M. fascicularis
from both Kepulauan Mentawai and Celebes suggests that this
macaque had not yet arrived in the Sunda area at the time that
maximal Pleistocene lowering of sea level made possible dispersal of
the silenus-group to these outlying islands (cf. Dorst, 1953, p. 312).
The border between the ranges of M. n. leonina and M. n.
nemestrina almost exactly coincides with the location of the
Khlong Marui Fault (Garson and Mitchell, 1970, p. 45), which cuts
across the sharp bend in the Thai-Malay Peninsula. Significantly,
the Khlong Marui Fault area is a border zone for many groups of
animals and plants. Zoological evidence places the subregional
boundary between the Indochinese and Sundaic vetebrate faunas
about 250 km. north of the Khlong Marui Fault (Chasen, 1940a, p.
ix), and botanical evidence places the corresponding boundary
between the Continental Southeast Asiatic and Malesian floras
about 250 km. south (Kloss, 1920, p. 79; van Steenis, 1950, p. lxxii;
Keng, 1970, p. 51).
The broad pattern of faunal and floral differentiation, including
differentiation of M. n. leonina and M. n. nemestrina, that centers
around the Khlong Marui Fault probably implies the former
FOODEN: LIONTAIL & PIGTAIL MACAQUES 71
existence in this area of a physical barrier that blocked genetic
exchange between populations on either side of the barrier.
Geographic considerations suggest that this barrier may have been a
local marine transgression that transected the Thai-Malay Penin-
sula; this transgression may have been either eustatic or tectonic in
origin. Geological evidence pertaining to such a transgression is
ambiguous (Scrivenor, 1911, p. 9; 1949, p. 114; Viji Stresthaputra et
al., 1951, p. 40; Ho, 1960, p. 18; Kobayashi, 1960, p. 142).
Judging from degree of morphological differentiation, isolation
of the M. n. pagensis pigtail stock in Kepulauan Mentawai
probably dates from about the same time as disjunction and
isolation of the M. n. leonina and M. n. nemestrina pigtail stocks.
Dispersal of the ancestral Celebesian stumptail stock to peripheral
islands in the hypothetical Pleistocene Celebesian archipelago
(Fooden, 1969a, p. 64) may have occurred during this same stage in
the evolution of the silenus-group (fig. 18d).
Subsequent to the morphological differentiation of M. n.
leonina and M. n. nemestrina, the postulated former barrier
between their ranges evidently disappeared, bringing these two
pigtail stocks into secondary contact (fig. 18e). Judging from the
present restricted distribution of morphological intermediates (table
20), it appears that only limited gene exchange has subsequently
occurred between these two subspecies. The geographic change that
brought M. n. leonina and M. n. nemestrina into secondary contact
may also have provided an opportunity for the apparently recent
entry of M. fascicularis into the Sunda region (see above).
About 20,000 years ago, during the latest Wisconsin/Wurm
glacial advance, sea level in the Sunda area evidently was at least
120 m. lower than at present (Van Andel et al., 1967, p. 743; Emery,
1969, p. 114; Jongsma, 1970, p. 150). As a result of this most recent
(but not most extensive) exposure of the Sunda Shelf, the Malay
Peninsula, Sumatra, Bangka, and Borneo, which are now isolated
parts of the range of M. n. nemestrina, were interconnected by dry
land (fig. 18e). This relatively recent continuity of range presum-
ably promoted partial homogenization of the gene pool of M. n.
nemestrina, which is evident in the morphological similarity of now
disjunct insular and peninsular population of the subspecies (fig.
22).
With the postglacial rise of sea level, ranges of species and
subspecies in the silenus-group attained their present configuration
72 FIELDIANA: ZOOLOGY, VOLUME 67
(fig. 180- Two negative features of these ranges are somewhat
difficult to interpret. These two enigmatic features are (1) the
absence of M. n. leonina from apparently suitable evergreen forest
habitats (fig. 17) in central and northern Laos and North Vietnam
(Osgood, 1932, pp. 208-211) and (2) the absence of M. n. nemestrina
from similar suitable habitats in Java, which is on the Sunda Shelf.
These absences may indicate that unknown past physical or
biological barriers prevented the si/enus-group from dispersing to
the northeastern part of the Indochinese Peninsula and Java or,
alternatively, that populations of the group formerly inhabited
either or both of these areas and subsequently became locally
extinct. Problematical fossils that may record the presence in
Pleistocene Java of a population ancestral to M. n. nemestrina are
discussed above (p. 64).
SPECIES ACCOUNTS
Macaca silenus (Linnaeus, 1758)
[Simia] Silenus Linnaeus, 1758, p. 26 — part, subsequently restricted by Buffon
(1766, p. 169); external characters; distribution, Asia: Ceylon, Java, etc; the
specific name is a nominative singular masculine noun (Latin name of bearded
woodland diety) standing in apposition to the feminine generic name.
Simia Silenus [variety A]: Schreber, 1774, pp. 87, 186, pi. 11 (animal ex Buffon) -
part (also includes references to Presbytis sp.); external characters; habits;
distribution, Ceylon, Coromandel Coast, possibly East Indies.
Sim [ia] silenus: G. Cuvier, 1817, p. 108 — species included among "Les Macaques";
distribution, Ceylon.
Simia silanus [sic]: F. Cuvier, 1822b, p. 2, pi. (animal) — external characters of
captive, origin unknown; Indian name, Nil-Bandar.
Cercopithecus silenus: Kerr, 1792, pp. 32d, 64-part, lion tail macaque assigned to
subspecies Cercopithecus silenus albibarbatus Kerr, 1792.
Papio silenus: E. Geoffrey, 1803, p. 27 — zoo specimen listed; distribution, Ceylon.
[Ogilby], 1838, pp.364, 386, fig. (animal) — external charcters; taxonomic
relationships.
Pithecus silenus: Desmarest, 1817, p. 321 — external characters; distribution,
Ceylon and neighboring lands.
P[ithecus] silenus: Blainville, 1839, p. 30 — citation of possible reference in ancient
Greek literature.
Macacus silenus: Desmarest, 1820, p. 63, pi. 10 (animal ex Buffon) — external
characters; distribution, Ceylon. Bennett, 1829, p. 20, fig. (animals) — external
characters; habits in captivity; distribution, Malabar Coast, Ceylon. Blyth, 1851,
p. 154 — vernacular name not Wanderoo, which instead applies to Presbytis sp.
Anderson, 1879, p. 93 — external and cranial characters; distribution, Western
Ghats from near Goa to Cape Comorin. Blanford, 1888b, p. 16, fig. 5 (head) —
external characters; nomenclature; distribution, Western Ghats northward to
14°N.
[Macaca] Silenus: Thomas, 1911, p. 126 — nomenclature.
[Macaca] silenus-group: Miller, 1933, p. 5 — key to external characters.
Macaca silenus: Pocock, 1939, p. 66, pi. 4 (animals), figs. 16 (head ex Blanford,
1888b), 17 (skull) — external and cranial characters; taxonomy, closely related to
M. nemestrina leonina: Pocock, 1939; ecology; distribution, Western Ghats in
Travancore and Cochin, reportedly from Cape Comorin northward to 14°N.
73
74 FIELDIANA: ZOOLOGY, VOLUME 67
Cynoceph [alus] Silenus: Schinz, 1821, p. 114 — new combination; taxonomy;
distribution, Ceylon.
I [nuus] Silenus: Wagner, [1839], p. 141 — part (also includes references to Presbytia
sp. ); external characters taxonomy; distribution, Ceylon. Blyth, 1847, p. 731 —
not native to Ceylon according to Colombo resident Dr. R. Templeton; said to
be abundant in Travancore and Cochin.
I nuus silenus: Jerdon, 1867, p. 10 - external characters; taxonomy; field
observations; distribution, Western Ghats northward to 14°N.
V[etulus] Silenus: Reichenbach, 1862, p. 125, figs. 321-325 (animals ex authors) -
external characters; habits in captivity; distribution, Malabar.
Ouanderou: Buffon and Daubenton, 1766, p. 169, pi. 18 — part (also includes
references to Presbytis sp.); restriction of species and virtual designation of
neotype; external characters and measurements; habits; distribution, Ceylon,
Malabar Coast (implied). F. Cuvier, 1837, liv. 70, pi. (animal) — taxonomic
history; distribution, penisular India.
Lion tailed monkey [variety a ]: Pennant, 1771, p. 109, pi. 13a (animal) —
characters and habits of captive exhibited in London; compared with neotype of
M. silenus (Linnaeus, 1758).
[Cercopithecus] Vetulus Erxleben, 1777, p. 25 — new name based on (1) Simia
silenus Linnaeus, 1758, (2) Ouanderou: Buffon, 1766, (3) Lion tailed monkey:
Pennant, 1771, (4) various characterizations of Presbytis sp. and Colobus sp.;
type-locality, "Zeylona, Asia, Africa." Boddaert, 1785, p. 57 — restricted to
liontail macaque; distribution, Ceylon, Coromandel Coast. Fischer, 1829, p. 28 —
a synonym of Simia silenus Linnaeus, 1758.
Simia ferox Shaw, 1792, p. 69, pi. opposite p. 71 (animals) — new name based on (1)
Simia silenus Linnaeus, 1758, (2) Ouanderou: Buffon, 1766, (3) Lion tailed
monkey; Pennant; 1771, (4) characterization by Ray (1693, p. 158) of Presbytis
senex; type locality, "East Indies, ... Ceylon.. .also said to be found in the interior
parts of Africa." Shaw, 1800, p. 30, pi. 16 (animal ex Buffon) — external
characters; taxonomy. Blanford, 1888b, p. 16 — a synonym of Macacus silenus.
[Simia] Ferox: Griffith, 1821, p. 105, pi. (animal) — a synonym of Simia silenus
Linnaeus, 1758.
M[acaca] ferox: Wroughton, 1918, p. 554 — nomenclature; external characters; type-
locality, Ceylon.
Cercopithecus silenus albibarbatus Kerr, 1792, pp. 32d, 64 — new subspecific name
based on (1) Simia silenus Linnaeus, 1758, (2) Ouanderou: Buffon, 1766, (3) Lion
tailed monkey: Pennant, 1771, (4) characterization by Ray (1693, p. 158) of
Presbytis senex; type- locality, "Ceylon and the rest of India."
Pithecus albibarbatus: Elliot, [1913], p. 218, pi. 8 (animal; not cranial pi. 24,
misidentified skull of Macaca radiata; see Pocock, 1939, p. 68) — external
characters; taxonomic history.
Silenus albibarbatus: DeBeaux, 1923 p. 34 — external characters.
[Simia] Veter: Audebert, 1798-1800, Fam. 4, p. 4 (not Linnaeus, 1766) - said to be a
variety of Ouanderou: Buffon, 1766.
Silenus veter: J. E. Gray, 1843 P- 8 (not Linnaeus, 1766) —specimens listed.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 75
[Simla] leonina: G. Cuvier, 1817, p. 108 (not Shaw, 1800) — external characters;
name incorrectly attributed to Linnaeus and Gmelin; distribution, Ceylon.
Type. — None preserved. A captive specimen (fig. 19) described
and figured by first reviser Buffon (in Buffon and Daubenton, 1766,
p. 169, pi. 18) is now regarded as neotype (see discussion below).
As previously noted (Blanford, [1888a], p. 620; Thomas, 1911, p.
126; Elliot, 1913, p. 219; J. A. Allen, 1916, p. 50; Pocock, 1939, p. 66),
it is difficult or impossible to identify with certainty the original
objective basis of Simia silenus Linnaeus, 1758 as described in the
tenth edition of Systema Naturae (p. 26). Linnaeus' concept of this
taxon apparently was derived solely from published accounts of
earlier naturalists, not from direct examination of a specimen. In
the original description, quoted in full below, Linnaeus (1758, p. 26)
explicitly acknowledges that the species is obscure and that most of
its characters are unknown:
Silenus. 5. S[imia] caudata barbata, corpore nigro, barba nivea prolixa.
Simia Callitriches magnitudine cynocephalorum. Alp. aegypt. 242?
Habitat in Asia: Zeylona, Java & c.
Species obscurior, ignotis Pedum unguibus aliisque plurimis attributis.
In this account the skimpy diagnosis (tailed monkey with black
body and full white beard) is generally applicable to the Indian
liontail macaque. However, the bibliographic citation and the
assigned geographic range refer to at least two other species of
cercopithecids. The doubtful synonym cited (with a question mark)
from Alpinus (1735, Kistoriae Aegypti naturalis) is based on a figure
of a 6/ac/s-bearded monkey or baboon that reportedly originated in
Egypt. The indefinite geographic distribution given by Linnaeus
(Asia: Ceylon, Java, etc.) may have been partly influenced by Ray's
(1693, p. 158) characterization of a monkey in Ceylon as
"Cercopithecus niger barba incana promissa" ( = Presbytis senex); it
should be noted, however, that Ray's account of this Ceylonese
monkey is not explicitly cited by Linnaeus, although Ray's work is
cited in synonymies of eight other monkey species described in the
tenth edition of Systema Naturae.
The earliest known unequivocal association of Simia silenus
Linnaeus, 1758, with an actual specimen is in Buffon 's (in Buffon
and Daubenton, 1766, p. 169, pi. 18) account of the "Ouanderou" or
"Lowando." This report is based on a captive liontail (fig. 19) that
was exhibited at a fair in France. Although the owner of the captive
/'/ XI III /'.
fit .Iff /w
Ol'ANDHHOr.
HvMi
Fig. 19. Ouanderou: Buffon (in Buffon and Daubenton, 1766, pi. 18), neotype of
M. silenus Linnaeus, 1758.
76
FOODEN: LIONTAIL & PIGTAIL MACAQUES 77
asserted that it originated in South America, Buffon dismissed this
assertion as mere showmanship. He surmised instead that the
monkey probably was native to Ceylon. The sources of this
erroneous geographic conjecture evidently are the faulty account of
Simia silenus given by Linnaeus (1758, p. 26) and misinterpreted
characterizations of Presbytia senex published by Knox (1681, p. 41,
[1911 ed.]) and Ray (1693, p. 158), all of which are cited by Buffon
in the synonymy of the "Ouanderou"; Buffon derived the name
"Ouanderou" from the Ceylonese vernacular name cited by Knox
for Presbytis senex. Along with the mistaken reference to Ceylon,
Buffon (in Buffon and Daubenton, 1766, p. 171) also refers to the
true geographic range of the liontail ("Malabar") in a footnote
abstract of field notes published by P&re Vincent Marie, who
observed this species while serving as a missionary in southwestern
India.
In the same year that Buffon published his account of the
"Ouanderou," Linnaeus (1766, p. 36), in the twelfth edition of
Systema Naturae, published a radically modified characterization of
Simia silenus. In an obvious attempt to eliminate discrepancies
between the characters and range of Simia silenus and those of the
monkey cited from Alpinus, Linnaeus emended the diagnosis of
Simia silenus in the twelfth edition to "S[imia] caudata barbata
nigra, barba nigra prolixa" (beard black, not white, as in the tenth
edition) and changed the distribution to "Habitat in Egypto" (not
Asia: Ceylon, Java etc., as in the tenth edition).
However, despite ambiguity in the original characterization of
this species, and despite Linnaeus' subsequent contradictory
redefinition, the name silenus has been applied to the Indian
liontail macaque by virtually all authors since Buffon. To preserve
stability of the specific name, Buffon (in Buffon and Daubenton,
1766, p. 169, pi. 18) may be regarded as first reviser of the
imprecisely characterized composite nominal species Simia silenus
Linnaeus, 1758, and Buffon 's restriction of this species to the Indian
liontail macaque, represented by a described and figured captive
specimen (neotype, fig. 19), may be regarded as decisive. According
to this interpretation, Simia silenus Linnaeus, 1766, which is based
on a black-bearded African monkey, is an unavailable junior
homonym of Simia silenus Linnaeus, 1758, as restricted to the
Indian liontail macaque by first reviser Buffon (1766).
Type -locality. — Geographic origin initially given as "Asia:
Zeylona, Java & c." (Linnaeus, 1758, p. 26); restricted to "Ceylan"
78 FIELDIANA: ZOOLOGY, VOLUME 67
and "Malabar" by first reviser Buffon (in Buffon and Daubenton,
1766, pp. 169-171); "No exact typical locality available," according
to Thomas (1911, p. 126); type-locality designated "Egypt" by
Elliot ([1913], p. 219); now corrected to Western Ghats, inland from
Malabar Coast, southwestern India (see above).
Distribution. — Restricted to the Western Ghats, a low
mountain chain in southwestern peninsular India (fig. 20); known
altitudinal range 300-1300 m. (Loci, 12). The northernmost known
locality is Anshi Ghat (14°55'N), near Goa, and the southernmost is
Kalakkadu Hills (8°25'N), near Cape Comorin. The range of this
species evidently has been narrowly restricted at least since the
middle of the nineteenth century (see below). Recently the range
has been further reduced, almost to nil, as a result of local
deforestation caused by man (Sugiyama, 1968, p. 284; C. A. Hill,
1971, p. 35).
External characters (fig. 21). — Head and body length 457-481
mm. in 3 adult females, 508-584 mm. in 4 adult males; relative tail
length (T/HB) 0.56-0.69 in 3 adult females, 0.57-0.75 in 4 adult
males (table 18; fig. 2); weight 6.75 kg. in one male, presumably
adult (Napier and Napier, 1967, p. 406). Fur blackish on crown,
dorsal surface of trunk, arms, legs, hands, feet, and tail; side
whiskers and beard long, pale ochraceous-gray, forming a
conspicuous facial ruff; crown patch narrow anteriorly; skin of
muzzle blackish, thinly covered with inconspicuous short whitish
hairs; ventral surface of trunk thinly haired, grayish brown to
brown.
In adults the dorsal pelage is long and dense, 8-10 cm. long on
the scapular region and 4 cm. long on the lumbosacral region. The
fur at the tip of the tail in adult specimens examined consists
exclusively of coarse guard hairs, which are sparsely distributed
elsewhere in the pelage. These hairs usually are elongated and form
a more or less distinct terminal caudal tuft (fig. 21), on which
Pennant (1781, p. 183) based the name "lion tailed monkey." Hairs
on the crown are 2-3 cm. long and form a whorl radiating from the
vertex. More anteriorly, the dark frontal hairs are about 2 cm. long,
posteriorly directed, and form a sharply defined wedge-shaped patch
that is bordered anteriorly and laterally by short pale hairs of the
lateral supraorbital region and long pale hairs of the facial ruff. In
adults, pale hairs of the facial ruff may attain a length of 10-15 cm.
Cranial characters (figs. 3, 5). — Skull relatively small (greatest
length excluding incisors 102.3-112.6 mm. in 7 adult females, 128.5-
FOODEN: LIONTAIL & PIGTAIL MACAQUES
79
Fig. 20. Locality records of Macaca silenus: closed triangles = specimens
examined, open triangles = literature records. For details, see Gazetteer of Collecting
Localities.
Key to localities: 1. Anshi Ghat; Goa, near. 2. North Kanara. 3. South Kanara.
4. Coorg. 5. Kuttyadi Pass. 6. Wynaad. 7. Nilgiri Hills, 800-1300 m; Nilgiri Hills,
western slopes. 8. Parambikulam Valley. 9. Cochin. 10. Cotengady Estate;
Nelliampathy Hills; Palagapandy; Seetagundy Estate. 11. Anaimalai Hills, northern.
12. Anaimalai Hills, 800-1300 m.; Grass Hills; Varagaliyar. 13. Panniar. 14. High
Wavy Mountain; Varushnaad Valley. 16. Periyar Lake, northern shore; Periyar
Lake, south. 16. Manjolai Tea Estate. 17. Kalakkadu Hills; Naraikkadu Estate.
138.5 mm. in 7 adult males), rostrum relatively short, zygomatic
arches broad (table 19; figs. 7, 8); supraorbital ridges thick and
prominent; anterior surface of malar with a clearly defined
infraorbital concavity.
Habits and habitats. — This account is based on field
observations published by authors listed below. Authors are cited
TABLE 18. External variation in Af . silenus adults; collecting localities un-
known except as indicated.
Specimen No.
or reference
FMNH 98163
ZSBS 1959/264
Blanford, 1888°, p. 161
Blanford, 1888°, p. 16l
AMNH 548372
Anderson, 1879, p. 93
Blanford, 1888b, p. 16*
Blanford, 1888b, p. 161
Pocock, 1939, p. 67
1- Collected at Travancore.
2 Collected at Nelliampathy Hills.
Head and
Relative tail
body (mm.)
length (T/HB)
Females
481
0.57
?610
? 0.39
457
0.69
457
0.56
Males
575
0.57
?610
? 0.42
533
0.64
508
0.75
ca. 584
0.63
TABLE 19. Cranial variation in Af. silenus adults (permanent dentition
completely erupted); collecting localities unknown except as indicated.
Relative
Greatest
zygomatic
Rostral-postrostral
Specimen No.
length (mm.)
breadth 1
Females
ratio
BM 10b
111.1
0.67
0.51
BNHS 52832
ca. 106.2
ca. 0.69
ca. 0.46
NHRM Z4288
108.4
0.71
—
RMNH 1314
110.0
0.67
0.49
ZSBS 1959/264
112.6
0.65
0.49
ZSI 7752
102.3
0.71
0.40
ZSI7753
105.7
0.62
0.45
Means
108.0
0.67
Males
0.47
BNHS 5059/50643
134.8
0.69
0.61
BNHS 50622
128.5
0.68
0.57
FMNH 47322
135.4
0.69
0.60
MNHN 1925/8
136.6
0.67
—
NHMB 1610/3931
138.5
0.68
—
RMNH 2122
131.3
0.72
0.57
Means
134.2
0.69
0.59
1 Zygomatic breadth/greatest length.
2- Collected at North Kanara.
3- Collected at Palagapandy.
80
FOODEN: LIONTAIL & PIGTAIL MACAQUES 81
by means of the indicated abbreviations. Of these authors,
Sugiyama (1968) provides the most detailed and useful information.
B Baker in Blyth, 1859, p. 283
H Hutton, 1949, p. 690
J Jerdon, 1867, p. 10
Ka Karr, 1973, pp. 191-192
Ki Kinloch in Pocock, 1939, p. 68
Kr Krishnan, 1972, pp. 542-543
O'B O'Brien in Pocock, 1939, p. 69
P Poirier, 1970, p. 261
Sp Spence, 1921, p. 970
St Stonor, 1944, p. 591
Su Sugiyama, 1968, pp. 283-292
W-P Webb-Peploe, 1947, p. 629
M. silenus is a hill species restricted to dense evergreen forest
(B, H, J, Ki, Kr, P, Sp, Su, W-P) or semi -evergreen forest in which
trees are more than 20 m. tall (Su) at elevations between 300 m.
(Anshi Ghat) and 1300 m. (Su). Within this limited habitat, the
distribution of troops of M. silenus is sparse (H, Kr, O'B, St, Su, W-
P). As early as 1867, an observer (J) noted that this monkey
"frequents the most dense and unfrequented parts of the forest, . . .
and I had often traversed the Malabar forests before I first fell in
with it." In 1968 the total living wild population was estimated at
fewer than 1000 individuals (Su) in a maximum habitat area of 50
km. X 750 km.
The maximum number of individuals reported in a troop of M.
silenus is 22 (Su). Other reports of troop size are: "four or five" (St),
6+ (Ka), "about a dozen" (Ki), 16 (Su), "twelve to twenty or more"
(J), and "about twenty" (W-P). In two carefully studied troops, one
with 16 members included 2 adult males and 7 adult females, and
another with 22 members included 2 adult males and 10 adult
females (Su). Solitary adult males have been observed fairly
frequently (H, St, Su).
Although M. silenus is primarily arboreal (Su), it occasionally
descends to the ground:
Once several were seen climbing slowly down one tree and walking along
tbe ground to the next tree. This they climbed, investigated and came down to
earth to repeat the process. (W-P)
Similar terresterial excursions also are reported by two other
observers (H, Su). M. silenus may occasionally venture into
streams, since captive specimens are said to be fond of water and to
swim well (H). In the trees M. silenus evidently is somewhat
82 FIELDIANA: ZOOLOGY, VOLUME 67
deliberate in its locomotor behavior (Ka), moving more leisurely
and with less leaping than sympatric M. radiata, for example (H,
Kr, W-P). When disturbed, M. silenus flees high into the canopy
(Su).
Like other macaques, M. silenus feeds chiefly on fruit, but
augments its diet with a variety of other plant and animal foods.
The species has been observed to eat wild figs (St). Another
observer (Su) reports:
The only important food.. .during the study period was a.. .chestnut-like
fruit.. .In addition... they ate other fruits, nuts, flowers and young buds of many
kinds of trees, insect larvae living under tree-bark, pith of cardamon stems and
so on.
In one troop that also fed on fruit, behavior apparently directed
at locating arthropod food is described (Ka) as follows:
Another adult climbed to the uppermost branches of a tall emergent tree
where several dead branches projected above the surrounding canopy. Pieces
up to a foot long were broken off and torn apart, presumably in search of
arthropods in the rotting wood. The macaque moved lower in the tree before
dismantling the branch and then moved to the upper branches again, peered
into the broken stub and tore off another dead branch.
During its morning feeding session, this troop evidently was
dispersed over several hectares of forest (Ka).
Information on breeding periodicity is ambiguous. A birth peak
in January is indicated by observation (Su) in that month of three
new-born infants and also by observation (Ka) in early March of
one or two young about two months old. However, another observer
(W-P) asserts that females carrying young are seen regularly in
September. Likewise difficult to interpret is a report (Su) of 12
instances of sexual behavior, including four copulations, observed in
January and February; if these were fertile matings, the young
presumably would have been born in June or July.
Encounters between adjacent troops of M. silenus apparently
are usually hostile and characterized by aggressive vocal challenges:
As the overlapping part of the ranges of the two troops had many food
trees. ..both troops frequently came to them, and contact between the two
troops was observed many times. They were antagonistic and troop 2 was
dominant. When they came near, large adult males in each troop displayed by
whooping loudly but no direct fighting was seen. Usually, troop 1 moved away
after a short time of vocalization. (Su)
Contradictory reports concerning interactions of M. silenus
with sympatric primate species are discussed below. These reports
Fig. 21. Macaca silenus, external characters. (Photos courtesy San Diego Zoo)
84 FIELDIANA: ZOOLOGY, VOLUME 67
are not readily harmonized. According to one observer (Su), M.
silenus is tolerant of and subordinate to M. radiata:
A troop of bonnet macaque [M. radiata] moved among liontailed
macaques and sometimes even travelled following the latter. Liontailed
macaques usually did not react to the appearance of bonnet macaques but
sometimes moved away slowly from them. Although smaller, bonnet macaques
were dominant.
However, another observer (W-P) reports that M. radiata "will
keep out of the way when the Lion-tailed Monkey is about." This
report agrees with an early anecdotal account by Vincent Marie (in
Buffon, 1766, p. 171), which also implies that M. silenus is dominant
to M. radiata.
Similarly, three observers (Ka, St, Su) indicate that M. silenus
associates amicably with the Nilgiri langur, Presbytis johnii, while
two others (H, W-P) report that contacts between these species are
hostile:
In the Animalais there are about as many Lion-tailed as Black monkeys
[P. johnii] and some trespassing on each others' territory is inevitable. On
these occasions some terrific battles take place in the trees, and there are
casualities on both sides (W-P).
These contradictory reports may signify that interspecific
behavior is variable in these monkeys.
Like other macaques (Fooden, 1969a, p. 52), M. silenus may
form symbiotic associations with some species of birds. One observer
(St) noted a troop of these monkeys "in a Ficus tree, oddly enough
in company with the Great Indian Hornbill (Dichoceros bicornis)."
A giant squirrel, Ratufa indica, that approached a female M.
silenus with infant was responded to aggressively (Ka).
Specimens examined — Total 59. INDIA: Cochin, 1 (skull only,
BM); Cotengady Estate, 1 (BM); Malabar, 5 (1, NMS; 4, skins only,
RMNH); Malabar Coast ?, 1 (skin only, ZSI); Nelliampathy Hills, 2
(skins only, AMNH); North Kanara, 2 (skull only, BNHS);
Palagapandy, 1 (BNHS); Parambikulam, 1 (skin only, BM);
Seetagundy Estate, 1 (skin only, BNHS). Imprecise localities: 5 (2
skins only, AMNH), 6 (3 skins only, BM), 3 (1 skin and postcranial
skeleton only, 1 skin only, FMNH), 2 (skulls in skins, IRSN), 5 (2
skulls in skins, 1 skull only, MNHN), 1 (skull only, NHRM), 1 (skin
only, NMS), 7 (3 skins only, RMNH), 3 (1 skin only, USNM), 1
(ZSBS), 10 (4 skins only, 1 skull only, ZSI).
FOODEN: LIONTAIL & PIGTAIL MACAQUES 85
Macaca nemestrina (Linnaeus, 1766). (Synonymies under sub-
species headings)
Taxonomy. — Compared with liontail macaques, the three
recognized allopatric forms of pigtail macaques (Inuus leoninus
Blyth, Simia nemestrina Linnaeus, Macaca pagensis Miller) are
obviously similar in tail proportions and general brownish pelage
hue. These three forms of pigtails are dissimilar in pelage pattern,
average size, cranial characters, bacular structure, arboreal-terrest-
rial preference, facial markings, tail carriage, and plasma protein
frequencies. Whether to regard these three pigtails (leonina,
nemestrina, pagensis) as species or as subspecies depends ultimately
on analysis of geographic variation of the characters that
distinguish them (Mayr, 1963, p. 194). In the following paragraphs,
overall geographic variation of adequately known characters in
parapatric leonina and nemestrina is considered first. This is
followed by detailed discussion of diagnostic characters in critical
specimens of leonina and nemestrina that have been collected near
the common border between their ranges. The taxonomic status of
insular pagensis is considered last.
Pelage pattern is relatively constant and distinctive in leonina
and nemestrina. The pale dorsal surface and anteriorly narrow
crown patch in all leonina specimens examined is readily
distinguishable from the dark dorsal surface and broad crown patch
in all but four geographically marginal specimens of nemestrina (see
below). In leonina specimens, preauricular sidewhisker hairs are
elongate and pale from base to tip; in nemestrina specimens, these
hairs are short and dark tipped. Although pelage pattern varies
individually in leonina and nemestrina, this variation is not
geographic. There is no apparent overall geographic tendency for
pelage pattern in either of these pigtails to approach that of the
other.
External size, measured as the combined length of head and
body, apparently follows a continuous latitudinal cline in adult
leonina and nemestrina (fig. 22), with geographic variation parallel
in males and females. Judging from specimens examined, external
size is relatively constant in Indochinese populations of leonina
south to about 10° N in peninsular Thailand. Farther southward,
length of head and body in local populations of leonina and
nemestrina gradually increases, by about 100 mm. over 15 degrees
of latitude, to reach a maximum in Sumatran nemestrina. It is
significant that geographically isolated populations of nemestrina in
Sumatra, Borneo, and the Malay Peninsula apparently all conform
86
FIELDIANA: ZOOLOGY, VOLUME 67
20«N -
'.= 10° N
O
-
0
o ■ /eon/no
o • nemestrina
v » pagensis
-
o
0
o
a
A
•.
m
-u
%o
§.'
•
■
■"
°o
•
■
■
•
n
ooo
>
•
M
•
•
•
o
•••
•
9?
o
•
•
•
1
1
0
oo
o
1 1
o
- _1_
o
• •
• • •
iiil.
_ 0 T) O ^
■
■ • ■
° ° ° o-
<• •
OOOOQO
•* / .
o % oooo
O 0
•a
_ o
o
oo
o •
ooo
1 1 1 1
o
• •• •
•
1111
400 500 600 700
Head and body length (mm)
120 140 160
Skull length (mm)
Fig. 22. Latitudinal variation of head and body length and skull length in adult
specimens examined of leonina, nemestrina, and pagensis; specimen locality numbers
(figs. 25, 28) as follows: leonina (fig. 28), 1, 21-26, 29, 31, 32, 37, 39, 42, 44 (for
measurements of one female skull, see Pocock, 1932, p. 308), 45-47, Chamchay
(unnumbered locality, see "Gazetteer of Collecting Localities"); nemestrina (fig. 25),
14, 6, 9, 10, 12, 18, 24, 26, 28-30, 33, 34, 36-39, 42-52, 55, 56, 58, 64, 66, 68, 69, 76, 77, 82-
84, 87, 89, Banang, Trang (unnumbered locahties, see Gazetteer); pagensis (fig. 25),
92, 93. Small symbols represent measurements in two subadult nemestrina males
(canines incompletely erupted).
to the same latitudinal size cline. Over the entire range, external
size is broadly overlapping in leonina and nemestrina.
Cranial size, measured as greatest length of skull excluding
incisors, evidently follows a step cline in leonina and nemestrina
(fig. 22). The leonina curve and the nemestrina curve appear to be
abruptly discontinuous about 8°N, and known overlap of greatest
skull length in these two pigtails is mimimal. In adult males, skull
length exceeds 142 mm. in 56 of 58 nemestrina specimens examined
and in none of 15 leonina specimens; in adult females, skull length
exceeds 120 mm. in 24 of 27 nemestrina specimens and in only one
of 19 leonina specimens. Other aspects of cranial morphology
(zygomatic breadth, rostral-postrostral proportions, relative length
FOODEN: LIONTAIL & PIGTAIL MACAQUES 87
of dental fields; see above) are directly or allometrically related to
skull length and therefore have a similar pattern of geographic
variation.
The geographic ranges of leonina and nemestrina are
contiguous in peninsular Thailand (8-9° N) at the southern end of
the Isthmus of Kra (fig. 23). Available from this area are 13 pigtail
specimens (table 20), of which six are referred to leonina and seven
to nemestrina. In five insular specimens collected at Klong Tung
Sai and Telok Poh, virtually all known external and cranial
characters are those of leonina; the only exception is external size
in one adult male (570 mm.), which is in the range of overlap of
leonina and nemestrina. A baculum of the Telok Poh specimen also
is available (fig. 15d, table 6); although this is the shortest adult
leonina baculum examined (greatest length 19.7 mm.), its form is
clearly that of leonina (distal process length/shaft length = 0.47).
One mainland specimen collected at Ban Nong Kok (skin only,
subadult or adult male) likewise is a typical example of leonina in
all available characters, as previously noted by Chasen (1935, p. 38,
"Ghirbi" specimen).
The northernmost pigtail specimen that clearly exhibits pelage
characters of nemestrina is an adult female recently collected by
me at Khao Si Kaeo, which is about 10 km. north of Ban Nong Kok
(mainland leonina locality mentioned above). Critical characters in
this specimen are a mixture of those of nemestrina and leonina,
with nemestrina characters considered to predominate. The dorsal
surface is dark brown as in nemestrina, distinctly darker than in the
Ban Nong Kok leonina specimen and in all other leonina specimens
examined. Skull length (119.9 mm.) is in the range of overlap of
adult female nemestrina and leonina. The crown patch is narrow
anteriorly, as in leonina, and length of head and body (434 mm.) is
within the range of female leonina (less than in any other adult
female nemestrina). When encountered by hunters who took the
specimen, the troop to which this female belonged was 20-25 m.
above the ground in dipterocarp trees; in response to gunfire, some
members of the troop descended and fled on the ground, as in
nemestrina, and others fled in the treetops, as in leonina.
In one adult female and one adult male collected at Sathani
Lam Phura, about 100 km. southeast of Khao Si Kaeo, the dorsal
surface is dark and the crown patch is broad, both as in typical
nemestrina. Skull length in the female (127.7 mm.) also is typical of
— 13
o
(0
« -2
•z.
o> i^
•r
uo
*- ti
la
01
.a 2:
*
0
1- u
u
S3
< a
a
c
«
u
o
la
3
.c
3
u
c
u
p
0
tu
CQ
~
I I I
I I I I - I
c II I --
I I
CO
01
3
Zl
•-
_£
-
CO
X
-
1-
01
—
1-
0
—
z
B
—
—
CO
2
Q£
—
>>
r
c
S3
~
o>
0>
d
J
JS
■fl
v.
-
s
B
a.
0
JZ
a
u
a
a
5
0
_^
—
o>
c
M
ti
i_
eg
1
a
"3
i
0
~
o
S
=
~
01
CO
E
o>
01
"5
o
'5
2
o<
a-o
ta
o>
—
£
—
■-
o*
X
ti
o>
c
X
a
..
e
II
o>
■—
c
<
— C E —
t» t— C- t- X
<J> Oi Gi O C5
ot *0 o» "O
c- *- —
o c
Oi
cc n
— *-*
o
r~ x-
en a;
CI
U5 Cft
U3 IJ) Vi Ui U' W
"O jtt
i/: 0
CJ 01
o 5
(N ~
_:
<
0
Z
. B
II
CO -_
o
c
3 in
E- *
c m
a
><
0
CQ
>• -
0 © -£
- * m 3 o
*r cj -* .-« * t~
Cfl CO CO 0
o o u ft« _
a: qs a. .* S z o-
NNN 0 CO c N
X; c w
15 o O
X »-
m 3
*° £
in »> en w
22£
3
-
1- 't
U3 CO
5 z •- u u
Q S « os o;
a fc ■£ N N
lO
m' c
in 2
bS?
0 03 w
.c 3
to x 0
f- *"- Z-
to to -»
in in in
0 o —
•—• »-< tN
1 I . CO
ZZ'MU
S «> c OC
tv U- 2 N
FOODEN: LIONTAIL & PIGTAIL MACAQUES
Fig. 23. Map of peninsular Thailand showing marginal collecting localities of
Indochinese pigtails (leonina, squares) and Sundaic pigtails (nemestrina, circles).
nemestrina. (Length of head and body given on the tag of this
specimen is 590 mm., which is highly improbable; this may be a
lapsus for 490 mm.) Skull length in the male (139.6 mm.) is at the
lower limit of the nemestrina range (upper limit of leonina range)
and length of head and body (532 mm.) also is at the lower limit of
the nemestrina range (well within leonina range).
In one subadult male (canines not quite fully erupted)
collected at Chong, 25 km. southeast of Sathani Lam Phura, the
crown patch is narrow anteriorly, as in leonina, and the dorsal
surface is pale golden-brown agouti with a weakly defined median
dark streak (about 5 cm. broad), approximately intermediate
between typical leonina and nemestrina. Hairs in the side- whisker
region are short and dark-tipped, as in nemestrina. Skull length in
this subadult (147.7 mm.) is within the range of adult male
nemestrina, and length of head and body (648 mm.) approaches
90 FIELDIANA: ZOOLOGY, VOLUME 67
that in the largest known adult male specimens of nemestrina
collected in southern Sumatra. This Chong subadult appears to
have been the specimen on which Kloss (1919b, p. 344) based his
influential decision to regard leonina and nemestrina as conspecific.
In one subadult male (upper canines about 75 per cent erupted)
and one juvenile female that I collected near Huai Lian, 17 km. east
of Chong, the dorsal surface is dark brown to blackish, approximate-
ly as in typical nemestrina, and the crown patch is narrow, as in
leonina. In the subadult male, skull length (139.6 mm.) is about
equal to the maximum known in adult male leonina and to the
minimum known in adult male nemestrina. Length of head and
body in this subadult (548 mm.) is in the range of overlap of adult
male leonina and nemestrina. In the baculum of this subadult,
relative length of the distal process (0.30; fig. 15r) is slightly greater
than the maximum known in adult nemestrina (0.29; table 6, NMS
2629) and slightly smaller than in a less mature subadult leonina
(0.31; table 9, FMNH 99672); on balance, bacular structure in this
subadult seems slightly closer to that in nemestrina than in
leonina. These Huai Lian specimens were collected on successive
days and belonged to different troops; in response to gunfire, both
troops fled in the treetops, as in leonina.
Also collected in the general area of Sathani Lam Phura,
Chong, and Huai Lian, is an isolated adult male skull with locality
given merely as "Trang" (province). This is a typical, large
nemestrina skull (greatest length 154.3 mm.). Another male
specimen of nemestrina collected in this area is briefly mentioned in
passing by Kloss (1919b, p. 344); this may be the Sathani Lam
Phura male discussed above.
In summary, typical leonina specimens have been collected as
far south as Klong Tung Sai, Telok Poh, and Ban Nong Kok.
Within 10-150 km. of Ban Nong Kok, at Khao Si Kaeo, and in the
Trang area (Sathani Lam Phura, Chong, Huai Lian), seven pigtails
have been collected in which nemestrina characters— most
conspicuously, dark dorsal pelage color— are variably mixed with
leonina characters, especially the narrow crown patch. This accords
well with Kloss' (1919b, p. 344) observation, based on two specimens
available to him, that "Trang may therefore be regarded as the
locality where intermediates between the two forms occur." About
400 km. farther south, Malayan pigtail specimens exhibit characters
that are unambiguously those of typical nemestrina.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 91
The mixture of characters in geographically marginal speci-
mens referred above to nemestrina, taken together with the
continuous clinal distribution of external size in leonina and
nemestrina tends to support classification of these two pigtails as
conspecific subspecies, which, following Kloss (1919b, p. 344), has
been the usual recent taxonomic practice. However, the narrow
geographic distribution of known specimens with mixed or inter-
mediate pelage and bacular characters and the step cline in cranial
size probably indicate that genetic exchange between leonina and
nemestrina is relatively restricted. On balance, despite some
evidence of partial genetic discontinuity, it seems appropriate to
follow a conservative course and continue to regard these two
pigtails as subspecies of the species M. nemestrina.
The geographic isolation of Kepulauan Mentawai pagensis
precludes the possibility of interbreeding or intergradation with
neighboring Sumatran nemestrina. Since the degree of morphologi-
cal differentiation between pagensis and nemestrina is comparable
to that between nemestrina and leonina, it seems reasonable to
accord equivalent taxonomic status to all three of these forms.
Accordingly, pagensis is also treated in this revision as a subspecies
of M. nemestrina.
Macaca nemestrina nemestrina (Linnaeus, 1766)
Pig-tailed Monkey: Edwards, 1758, p. 8, pi. 214 (animal in color) — external
characters of two captives; distribution, Sumatra.
Le Maimon: Buffon and Daubenton, 1766, p. 176, pi. 19 (animal) — external
characters and measurements of captive, origin unknown; internal anatomy;
osteology; taxonomy; distribution, Sumatra and probably elsewhere in East
Indies.
[Simia] Nemestrina Linnaeus, 1766, p. 35 - external characters ex Edwards (1758);
distribution, Sumatra; the specific name is an adjective (derived from Latin
name of god of groves) agreeing in gender with the feminine generic name.
Simia Nemestrina: Schreber, 1774, p. 79, pi. 9 (animal ex Edwards, 1758) -
external characters; distribution, Sumatra.
[Papio] Nemestrina: Erxleben, 1777, p. 20 - external characters; distribution,
Sumatra, Japan.
Papio nemestrinus: [Olgiby], 1838, pp. 363, 373 - taxonomy. Cantor, 1846, p. 176 -
first record of pigtail macaque in Malay Peninsula (Pulau Pinang).
Macaca nemestrina: Daudin, [1802], p. 148 - listed; new combination. Miller, 1906,
p. 556, pis. 13-20 (skin, skulls) — external and cranial characters; range restricted
to Sumatra. Grimm, 1967, p. 361 - vocalizations. Blakley, Morrow, and Morton,
1973, p. 120 — serum cholesterol higher in Sumatran specimens than in West
Malaysian specimens.
92 FIELDIANA: ZOOLOGY, VOLUME 67
Macacus nemestrinus: Desmarest, 1820, p. 66, pi. 10 (animals ex Edwards and
Buffon) — external characters; distribution, Java Sumatra. Anon., 1851, p. 444 —
distribution, Sumatra, Borneo, Pinang, Malacca. Bleeker, 1851, p. 527 — first
record of pigtail macaque in Pulau Bangka. Hagen, 1890, p. 82 — field
observations. Sclater, 1893, p. 325 — report of aberrantly colored Bornean
specimen. Thomas and Hartert, 1894, p. 654 — field observations; introduced in
Kepulauan Natuna.
Macaca nemestrina nemestrina: Kloss, 1919b, p. 345 — external characters; said to
intergrade with Indochinese pigtail macaque at Trang, peninsular Thailand. J. R.
Napier and P. H. Napier, 1967, p. 403 - listed.
Macacus nemestrinus nemestrinus: Davis, 1958, p. 126 — external characters; field
notes.
Pithecus nemestrinus: E. Geoffroy, 1803, p. 25 — external characters of two
museum specimens; distribution, "L'Inde meridionale."
Cynocephalus nemestrinus: Latreille, 1804, p. 291 — external characters.
Inuus nemestrinus: E. Geoffroy, 1812, p. 101 — external characters, distribution,
Java, Sumatra.
I{nuus] nemestrinus: Wagner, [1839], p. 143 — external characters; distribution,
Sumatra, Borneo.
Macacus (Innus) nemestrinus: Muller, 1839-1840, p. 14 — field observations;
distribution, Sumatra and Borneo, transported from Sumatra to Java.
S[almacis] nemestrina: Gloger, [1841], p. 36 — new combination.
Nemestrinus nemestrinus: Schneider, 1905, p. 73 — field observations.
Rhesus nemestrinus: DeBeaux, 1923, p. 36 — external and cranial characters.
Pig-tail [Baboon]: Pennant, 1771, p. 105 — part (also includes references to Macaca
fuscata); external characters; distribution, Sumatra, Japan.
New Baboon: Pennant, 1771, p. xxiii, pi. 13 (animal) — external characters, based
on figure of specimen of unknown origin.
Simia platypygos Schreber, 1774, p. 74, pi. 5b (animal, head) — syntypes, male and
female of unknown origin observed in captivity by D. Hermann; external
characters; habits; characteristic pigtail grimace noted. Erxleben, 1777, p. 629 —
external characters. Zimmermann, 1780, p. 181 — possibly a synonym of Papio
nemestrina Erxleben, 1777. E. Geoffroy, 1812, p. 101 - a synonym of Simia
nemestrina Linnaeus, 1766.
Papio platypygos: Kerr, 1792, pp. 32d, 62 - listed.
Brown Baboon: Pennant, 1781, pi. 20 (animal) - replacement name for New
Baboon: Pennant, 1771; Simia platypygos Schreber, 1774, cited in synonymy.
Babouin a longues jambes: Buffon, 1789, p. 41, pi. 8 (ex Pennant, 1781) - external
characters and habits of captive specimen, origin unknown; Simia platypygos
Schreber, 1774, cited in synonymy.
S[i/rn'a] Longicruris Link, 1795, p. 62 — based on Babouin a longues jambes:
Buffon, 1789. J. E. Gray, 1843, p. 7 — synonym of Macacus nemestrinus
Desmarest, 1820.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 93
S[imia] Fusca Shaw, 1800, p. 24, pi. 13 (animal ex Pennant, 1781) - type, Brown
Baboon: Pennant, 1781; Simia platypygos Schreber, 1774, cited as a possible
synonym. E. Geoffroy, 1812, p. 101 — a synonym of Simia nemestrina Linnaeus,
1766.
Macaca sphinx [Variete C]: Daudin, [1802], p. 149 (not Linnaeus, 1758) - applied
to Babouin a longues jambes: Buffon, 1789.
Macaque, Espece inedite: F. Cuvier, 1816-1817, pi. 5 (animal) — figure of tumescent
female, origin unknown.
Singe a queue de cochon: F. Cuvier, 1820, liv. 19, p. 1, pi. (animal) - taxonomy,
referred to Simia nemestrina Linnaeus; external characters; reproductive
biology, sexual swelling, gestation 7 months 20 days; distribution, Sumatra. F.
Cuvier, 1822a, liv. 36, p. 1, pi. (animal, head) — taxonomy; distribution, Sumatra.
Simia carpolegus Raffles, [1821], p. 243 — based on specimens collected and observed
in Sumatra by T. S. Raffles between 1818 and 1820; syntypes formerly preserved in
collections of Zoological Society of London (Waterhouse, 1838, p. 8) and East India
Company (Horsfield, 1851, p. 19); external characters and variation; M3 5-cusped;
trained to pick coconuts; Malay name, Bruh; type-locality, Bencoolen
( = Bengkulu), southwestern Sumatra. Fischer, 1829, p. 30 — a synonym of Simia
nemestrina Linnaeus, 1766.
Cynocephalus carpolegus: [Vigors], 1829, p. 13 — specimen listed.
Macacus libidinosus I. Geoffroy, 1826, p. 589 — based on illustration of Macacque,
Espece inedite: F. Cuvier, 1816-1817, origin unknown.
S[imia] libidinosa: Fischer, 1829, p. 30 — doubtfully distinguishable from Simia
nemestrina Linnaeus, 1766.
P[ithecus] Maimon Blainville, 1839, pi. 10 (dentition of adult male) — new name
used in figure caption.
Macaca nemestrina var. leoninus: Oudemans, 1890, p. 266 — captive juvenile
received from Sumatra.
Macaca broca Miller, 1906, p. 558 — external and cranial characters; holotype,
adult male, skin and skull (USNM 34930, Coll. No. 19211), collected at Sungai
Sapagaya, Sabah by C. F. Adams, Nov. 21, 1887; distribution, Borneo. Lyon,
1907b, p. 566 — cranial comparison with Bornean M. nemestrina. Elliot, [1913], p.
205 — a synonym of Simia nemestrina Linnaeus, 1766; synonymy said to be
concurred in by G. S. Miller, original author of broca.
P[ithecus] broca: Lyon, 1911, p. 136 — cranial comparison with Bornean M.
nemestrina.
Macaca nemestrina broca: Myers and Kuntz, 1969, p. 420 — nematode parasite.
Macaca nemestrina nucifera Sody, 1936, p. 42 — based on skin (head only) and
skull of adult male collected at Pulau Bangka by H. J. V. Sody (Coll. No. Bk
75), date and museum unknown; external and cranial characters; taxonomic
comparisons. Sody, 1937, p. 248 — taxonomic comparison. Chasen, 1940a, p. 65 —
a synonym of Macaca nemestrina nemestrina.
Type. — None preserved. Linnaeus' (1766, p. 35) description of
this species is based solely on Edwards' (1758, p. 8) illustrated
FIG. 24. Pig- tailed Monkey: Edwards (1758, p. 8), holotype of M. nemestrina
nemestrina (Linnaeus, 1766).
94
FOODEN: LIONTAIL & PIGTAIL MACAQUES 95
account of a captive juvenile male (fig. 24) that was brought from
Sumatra to England on a British warship in 1752 and was
subsequently purchased by Edwards. In Edwards' account, the
holotype juvenile is compared with an older captive female
(paratype) that was then on exhibit at a fair in London. Neither
holotype nor paratype is known to have been preserved.
Type -locality. — Sumatra (Edwards, 1758, p. 8).
Distribution (fig. 25). — Sunda area from Surat Thani-Krabi
depression in peninsular Thailand (8-9°N) southeastward through
West Malaysia, Sumatra, Bangka, and Borneo; known altitudinal
range 0-1700 m. (table 11). M. n. nemestrina apparently is native to
the offshore islets Pulau Pinang (west coast of West Malaysia),
Pulau Tioman (east coast of West Malaysia), and Pulau Batam
(Riau Archipelago, southern tip of West Malaysia) (see Gazetteer).
Other offshore islets generally are excluded from the natural range
of the subspecies, although it often is present as an introduced
element. A captive specimen said to have been obtained in the
Nicobar Islands (Corner, 1941, p. 14) probably represents such an
introduction (see Kloss, 1928, p. 802).
Anomalous locality information is recorded on the museum tag
of one captive adult male specimen of M. n. nemestrina (AMNH
119514, skin and skeleton; blackish dorsal streak, greatest length of
skull 144.8 mm.) that died in the New York Zoological Park on
June 18, 1937 and was catalogued at the museum on the same day.
According to the tag, this monkey was born about December, 1931
and was obtained by the Legendre Indo-China Expedition (date
unspecified) at Plateau des Bolovens, Laos, which, however, is far
outside the indicated range of M. n. nemestrina (fig. 25). It is known
that the Legendre expedition collected a series of M. n. leonina
skins (AMNH 87268 6, 87269 8, 87273?, 87274$; immatures) at
Plateau des Bolovens in February 1932 and also obtained a living
specimen that was donated to the New York Zoological Park later
the same year (Berger, 1932, p. 329; Legendre, 1932, p. 495; New
York Zoological Society Annual Report, 1933, p. 109). It seems
likely that geographic information properly applicable to Legendre's
captive Indochinese pigtail (whereabouts now unknown) was
inadvertently transferred to a captive Sundaic specimen (AMNH
119514); this may have occurred while both were being kept at the
New York Zoological Park.
External characters (fig. 26). - Head and body length 434-576
mm. in 17 adult females, 532-738 mm. in 23 adult males (excluding
Fig. 25. Locality records of M. n. nemestrina (circles) and M. n. pagensis
(inverted triangles); closed symbols = specimens examined, open symbols =
literature records; squares - marginal records of M. n. leonina; letters and heavy
broken lines indicate negative reports. For details, see "Gazetteer of Collecting
Localities."
M. n. nemestrina localities: 1. Khao Si Kaeo. 2. Lam Phura. 3. Chong. 4. Huai
Lian. 5. Biserat. 6. Pinang. 7. Larut; Perak, Upper. 8. Gedong; Keroh Forest
Reserve. 9. Hantu, Tanjong; Simpang Empat Rungkup. 10. Jeram; Lima Belas
Estate. 11. Sungai Buloh. 12. Kelang; Meru; Sungai Renggam. 13. Galas, Sungai. 14.
Gua Madu. 15. Bukit Fraser, 5 km. S; Bukit Fraser, northern slopes. 16. Kuala
Lompat: G. Benom. 17 Bentong; Bentong, 6-9 miles W. 18. Ampang; Kepang; ? Kuala
Lumpur; Kuala Lumpur, Cheras Road, 7th mile; Ulu Gombak Forest Reserve. 19.
Melaka. 20. Janing, Gunong. 21. Johor, southeast. 22. Tioman, Pulau. 23. Batam,
Pulau. 24. Pasi. 25. Leuser Reserve; Pulau Munteh. 26. Aru, Teluk; Bukit Tinggi;
Langsa; Pangkalanberandan; Pulutelang. 27. Borohok; Langkat; Serapit. 28.
Batangkuwis; Deli; Dolok-Oelve; Medan; Tandjungmorawa. 29. Bedagi; Lauttador;
Padang [district]; Paguruan, Sungai; Serdang; Tebingtinggi; Tandjung. 30. Sibolga,
Teluk. 31. Sirambas. 32. Lubukbasung. 33. Padang. 34. Lida Ajer cave; Pajakumbuh.
35. Djamboe Cave; Sibrambang Cave. 36. Kajutanam; Solok. 37. Tarusan, Teluk.
38. Siak, Sungai. 39. Kateman, Sungai. 40. Baru, Danau. 41. Kota, Danau; Rengat.
42. Bengkulu. 43. Sanggul, Bukit. 44. Lubuklinggau; Muarabeliti. 5. Pagaralam. 46.
Babat; Kluang; Loeboek Karet; Palembang [district]. 47. Palembang. 48. Ogan, Air.
49. Bangka, Pulau. 50. Kuching; Sidong, Gunong. 51. Sekajam, Sungai. 52. Landak,
Sungai. 53. Kapuas, Sungai. 54. Pontianak. 55. Sukadana. 56. Riam. 57.
Kendawangan, Sungai. 58. Pulo Paku. 59. Entawa-Samarahan. 60. Roema Manoeal.
61. Semitau. 62. Sintang. 63. Niah Caves. 64. Salai, Sungai. 65. Kalulong, Bukit. 66.
Sut. 67. Puruktjahu. 68. Tandjung. 69. Pamukan, Teluk. 70. Klumpeng, Teluk. 71.
Pelaihari. 72. Papar. 73. Labuan, Pulau, mainland opposite. 74. Madihit, Sungai. 75
96
FOODEN: LIONTAIL & PIGTAIL MACAQUES 97
improbable measurements: adult female, 590 mm., ZRSC 1214/10,
Loc. No. 2; adult male, 469 mm., ZRSC 2592/10, Loc. No. 10) (fig.
22) relative tail length (T/HB) 0.30-0.44 in 16 adult females (0.16 in
one aberrant adult female, AMNH-AC 106562, Loc. No. 43, largest
female specimen examined), 0.28-0.42 in 22 adult males (fig. 2);
weight 5.4-7.6 kg. in 10 adult females, 10.0-13.6 kg. in 8 adult males
(table 21). Pelage of trunk yellowish-brown agouti to golden-brown
agouti laterally, with a variably developed blackish mid-dorsal
streak or patch (about 4-20 cm. broad) that becomes more or less
indistinct anteriorly on the scapular region; crown hairs short (1
cm.), blackish, radiating to form a whorl centered at the vertex;
crown patch broad anteriorly, extending laterally on supraorbital
region approximately as far as lateral angle of each eye; side
whiskers relatively short (2-3 cm.), with buffy bases and blackish
tips that collectively form a pair of dark preauricular streaks or
sideburns; skin of muzzle pale brownish, thinly covered with
inconspicuous short whitish hairs; outer surface of arms, legs, hands
and feet yellowish-brown to golden-brown, approximately like
flanks; posterior surface of shanks frequently with indistinct dark
streaks; tail thin, bicolor, sharply defined blackish dorsally, pale
ochraeceous-buff ventrally, frequently with terminal hairs elongate
and weakly defined ochraceous to dark reddish-brown, forming an
inconspicuous tuft; underparts thinly haired whitish to ochraceous-
buff anteriorly, often becoming darker (buffy to pale brown) on the
epigastric region.
Fig. 25. continued
Bareo; Pa Umor. 76. Bahau, Sungai. 77. Badang; Peleben. 78. Karangan, Sungai. 79.
Kin.il Reserve. 80. Tabang. 81. Djembajan, Sungai. 82. Lumu Lumu; Ranau. 83.
Betotan; Lungmanis Station. 84. Sapagaya, Sungai. 85. Ulu Segama Forest Reserve.
86. Tibas, Sungai. 87. Sandakan. 88. Kinabatangan, Sungai. 89. Abai. 90. Kretam
Kechil, Sungai; Segama, Sungai.
M. pagensis localities: 91. Sioban. 92. Pagai Utara, Pulau. 93. Pagai Selatan,
Pulau.
Negative reports: C. Nicobar Islands, northern. D. Nicobar Islands, southern. E.
Klet Kaeo, Ko (fig. 28). F. Simeulue, Pulau. G. Banjak, Kepulauan. H.
Bangkaru, Pulau. I. Nias, Pulau. J. Musala, Pulau. K. Enggano, Pulau. L. Sebesi,
Pulau. M. Panaitan, Pulau. N. Krakatau, Kepulauan. O. Pangkor, Pulau. P. Berhala,
Pulau. Q. Jarak, Pulau. R Sembilan, Kepulauan. S. Perhentian Besar, Pulau. T.
Redang, Pulau. U. Bidong Laut, Pulau. V. Kundur, Pulau. W. Karimun, Pulau. X.
Tinggi, Pulau. Y. Durian, Pulau. Z. Sugi, Pulau. a. Bulan, Pulau. b. Bintan, Pulau. c.
Sebangka, Pulau. d. Bakung, Pulau. e. Lingga, Pulau. f. Singkep, Pulau. g. Natuna,
Kepulauan. h. Lemukutan, Pulau. i. Temadju, Pulau. j. Datuk, Pulau. k.
Masalembo-Besar. 1. Laut, Pulau. m. Sebuku.
FlG. 2& Mid in a nemestrina nemestrina, external characters. (Photo courtesy
Japan Monkey Centre)
98
Females
Males
5.45
—
—
10.9, 11.8
7.3, 7.6
—
7.3
13.6
6.4
10.9, 11.0
10.9, 10.9
5.4, 5.7,
6.4,
6.4,
6.8
10.0
6.5
11.2
FOODEN: LIONTAIL & PIGTAIL MACAQUES 99
TABLE 21. Weight in wild collected adult specimens of M. n. nemestrina.
Locality Weight (kg.)
Nos.l
/
26
30
37
38. 39
55, 69
89
Means
1- See map, Figure 25.
In adult males dorsal hairs are longest (7-9 cm. long) on the
scapular region, where the blackish median streak is least distinct.
The basal quarter of each of these scapular hairs is pale buffy, and
the distal three-quarters is annulated with about ten alternating
bands of yellowish or golden and blackish, with the blackish bands
generally broader than the yellowish bands. More posteriorly, hairs
in the blackish median streak are 3-4 cm. long, with the basal half
of each hair pale buffy and the distal half blackish, rarely with an
indistinct narrow pale annulation.
In three subadult or adult males collected at localities less than
150 km. apart in west-central West Malaysia, width of the blackish
mid-dorsal streak varies from about 8 cm. (BM 55.1502, subadult,
Bentong) to 12 cm. (BM 55.1503, adult, Sungai Renggam) to 18 cm.
(BM 50.1501, adult, Rungkup). Anterior extent of the dorsal streak
is roughly correlated with its width. In the West Malaysian
specimen with the narrowest dorsal streak (BM 55.1502), the streak
is separated from the blackish patch on the crown and nape by a
clearly defined zone of golden-brown on the scapular region; in the
specimen with the broadest dorsal streak (BM 55.1501), a virtually
uninterrupted blackish band extends from the crown to the dorsal
surface of the tail, with only a slight tinge of brownish on the
scapular region. The darkest of these West Malaysian specimens
(BM 55.1501) is closely matched by an adult specimen collected in
southeastern Sumatra (BM 1938.11.30.6, Palembang), near the
southern limit of the range of the subspecies, and by another
collected in northeastern Sarawak (BM 93.6.2.2, Bukit Kalulong),
near the northeastern limit of the range. In a very old male
collected in Sabah (USNM 34930, Sungai Sapagaya, type of broca
100 FIELDIANA. ZOOLOGY, VOLUME 67
Miller), the dorsal ground color is drab-yellowish-brown, with
unusually indistinct annulations on individual hairs.
In adult females and in immatures of both sexes the dark
dorsal streak is generally somewhat more restricted and dilute than
in adult males, the hair on the flanks is paler and drabber than in
adult males, and the agouti pattern is less conspicuous. In one adult
female (BM 93.4.29.1, Sungai Baram, Sarawak) that reportedly lived
in captivity in Sarawak for about five years (Sclater, 1893, p. 325),
the dorsal fur, which is unusually fine and silky, is pale golden-
brown non-agouti laterally, with a narrow and indistinct darker
brown streak medially. Pocock (1931, p. 310) has speculated that
this exceptionally pale specimen is a captive-bred M. mulata-M. n.
nemestrina hybrid; alternatively, it may merely be a albinistic
specimen of M. n. nemestrina. Two albinistic Bornean specimens
have been reported by Banks (1931, p. 111).
As previously noted (fig. 22), length of head and body generally
increases from north to south in M. n. nemestrina, with the same
clinal pattern of increase evident in all three isolated components of
the range (Malay Peninsula, Sumatra, Borneo). However, one
exceptionally large subadult male (BM 55.1504) has been collected
at Chong, peninsular Thailand, near the northern limit of the range
(fig. 22). Relative tail length tends to be somewhat less in the
largest specimens examined than in smaller ones (fig. 2).
Cranial characters (figs. 4, 6, 27). - Skull relatively large,
greatest length excluding incisors 115.4-138.9 mm. in 35 adult
females, 139.5-174.5 mm. in 77 adult males (fig. 7); rostrum long
(fig. 8); zygomatic arches relatively narrow (fig. 7); supraorbital
ridges relatively thin, especially laterally, and somewhat receding;
anterior surface of malar flat to weakly convex.
Skull length in specimens collected in the northern part of the
range of M. n. nemestrina (Malay Peninsula; Sumatra and Borneo
north of 3°N) averages about 10 mm. less than in those collected in
the southern part of the range (Bangka; Sumatra and Borneo south
of 3°N) (fig. 22). However, one large specimen (NMS 16640, skull
only, greatest length 170.7 mm.) reportedly collected at "Dolok-
Oelve," Deli, northern Sumatra, appears to be an exception to this
generalization (fig. 22).
Habits and habitats. — This account is based primarily on field
observations published by authors listed below; authors are cited by
means of indicated abbreviations.
Fig. 27. Cranial variation in adult male M. n. nemestrina: a, AMNH-AC 106037,
Badang; b, MZB 6501, Pulau Bangka (possible captive): c, AMNH-AC 106563, Bukit
Sanggul. xtt (Photos FMNH)
101
102
FIELDIANA: ZOOLOGY, VOLUME 67
A Abbott in Lyon, 1907b, p. 566
Ba Banks, 1931, p. 110
Be-a Bernstein, 1967a, pp. 198-207
Be-b Bernstein, 1967b, pp. 217-228
C Chivers, 1971, pp. 78, 84
D Davis, 1962, p. 58
Hg Hagen, 1890, p. 82
Hrt Hartert, 1902, p. 193
Hrt/T Hartert in Thomas and Hartert, 1894, pp. 654-655
Hs Harrison, 1969, p. 175
Hss/D Harrisson in Davis, 1958, p. 126
Hss/M Harrisson in Medway, 1965, p. 85
K Kawabe, 1970, p. 286
K/M Kawabe and Mano, 1972, p. 216
Ku Kurt, 1973, pp. 64-65
L Lim Boo Liat, 1969, p. 127
McC McClure, [1965], p. 57
Me Medway, 1970, p. 527
Me/W Medway and Wells, 1971, p. 247
Mii Muller, 1839-1840, pp. 17-18
R Rodman, 1973, pp. 655-659
Sch Schneider, 1905, p. 73
So/C Southwick and Cadigan, 1972, p. 8
St Stott, 1964, p. 12
These published field observations have been supplemented by
brief unpublished notes recorded on field tags of four museum
specimens collected in Sumatra by M. Boogaarts in 1929 (SMF
16640, ad 6\ "Dolok-Oelve") and J. J. Menden in 1933 (AMNH-AC
102203, ad $, Lubuklinggau; AMNH-AC 102201-02, juv. <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; llo12'-ll°40'N, 76°14'-
77°0O'E; observed by Y. Sugiyama (1968, p. 284) Sept., 1961-Feb.. 1962. 7
Nilgiri Hills, western slopes, 3000-4000 ft.; INDIA: Tamil Nadu; ca. 11°25'N, 76°30'E;
observed by F. E. Poirier (1970, pp. 258, 260), Sept., 1965-Aug., 1966. 7
North Kanara [district]; INDIA: Mysore; ca. 14°45'N, 74°30'E; collected by W. F.
Jardine, March 7, 1906 (BNHS). 2
Palghat. See Cotengady Estate.
Palagapandy; Nelliampathy Hills; INDIA: Kerala; ca. 10°35'N, 76°45'E; collected by
A. M. Kinloch, March 25, 1920 (BNHS; also see Anon., 1921, p. 411). 10
Panniar, 1000-1250 m., High Range, Cardamon Hills; INDIA: Kerala; ca. 10°00'N,
77°00'E; observed by Y. Sugiyama (1968, p. 284), Jan. 5-Feb. 27, 1963. 13
Parambikulam Valley, 1600 ft., near Kuriarkutti; INDIA: Kerala; 10°25'N, 76°43'E;
collected by C. H. Biddulph, catalogued 1953 (BM). 8
Periyar Lake, northern shore; INDIA: Kerala; 9°36'N, 77°11'E; reported by J.
Tanaka (1965, p. Ill), Jan. 7-Feb. 5, 1963. 15
Periyar Lake, south of, 800-1300 m.; INDIA: Kerala; ca. 9°25'N, 77°20'E; observed
by Y. Sugiyama (1968, p. 284), Sept., 1961-Feb., 1962. 15
Seetagundy Estate, Kollangode district, Nelliampathy Plateau (Lindsay, 1926, p.
592); INDIA: Kerala; ca. 10°35'N, 76°45'E; collected by A. M. Kinloch. Nov. 29,
1920 (BNHS; also see Spence. 1921, p. 971). 10
South Kanara [district], northern border; INDIA: Mysore; ca. 13°55'N, 74°40'E;
reported by G. C. Shortridge (in Ryley, 1913, p. 490). 3
Tinnevelly. See Naraikkadu Estate.
Travancore district; INDIA: Kerala; 8°04'-10°21'N, 76°14'-77°37'E; collected by F.
W. Muu nli 1 Ion. date and museum unknown (Blanford, 1888b, p. 17). (not mapped)
Vaigai River. See High Wavy Mountain.
Varagaliyar, Anaimalais; INDIA: Kerala or Tamil Nadu; not precisely located,
10°15'-10°31'N, 76°51'-77020'E; observed by M. Krishnan ([1972], p. 542), April 30.
1960. 12
FOODEN: LIONTAIL & PIGTAIL MACAQUES 125
Varushnaad Valley, south of, 3000-6000 ft.; INDIA: Tamil Nadu; ca. 9°35'N,
77°30'E; observed by A. F. Hutton (1949, p. 690). 14
Wynaad district; INDIA: Kerala; 11°27'-H058'N, 75°47'-76°27'E; reported by A. F.
Hutton (1949, p. 690). 6
Macaca nemestrina nemestrina
Map, Figure 25
Abai; EAST MALAYSIA: Sabah; 5°42'N, 118°23'E (Coolidge, 1940, pp. 123, 129);
collected by S. L. Washburn and A. H. Schultz, June 27-July 26, 1937 (MCZ). 89
Ampang; WEST MALAYSIA: Selangor; 3°09'N, 101°46'E; observed by I. S.
Bernstein (1967b, p. 220), May 4-14, 1965. 18
Aru, Teluk; INDONESIA: Sumatra; 4°09'N, 98°12'E; collected by W. L. Abbott (in
Lyon, 1908, p. 620), Nov. 29-Dec. 11, 1905 (USNM). 26
Atjeh. See Langsa, Atjeh.
Babat; INDONESIA: Sumatra; 2°46'S, 104°06'E; collected by Soekarno, Oct. 17-
Nov. 2, 1933 (MZB). 46
Badang; INDONESIA: Kalimantan; not precisely located; ca. 3°N, 117°E; collected
by V. von Plessen, May 12-25, 1935 (AMNH-AC). 77
Bagan Datoh. See Simpang Empat Rungkup.
Bahau, Sungai; INDONESIA: Kalimantan; 2°50'N, 115°57'E; collected by P. Pfeffer
(letter June 22, 1972), May 21, 1957 (MNHN). 76
Bakong. See Bakung, Pulau.
Bakung, Pulau; INDONESIA: Sumatra; 0°04'N, 104°27'E; M. n. nemestrina absent
(Dammerman, 1926, p. 316). d
Banang, Sungai; INDONESIA: Kalimantan; not precisely located, on west coast
between 2°N and 3°S; collected by C. Bruegel, Feb. 27, 1909 (ZSBS). (not mapped)
Bangka, Pulau; INDONESIA: Sumatra; 2°S, 106°E; reported by Bleeker (1851, p.
527); collected by H. J. V. Sody (1936, p. 45), date and museum unknown; collector
and date unknown (MZB). 49
Bangka, Pulau, north coast or west coast; INDONESIA: Sumatra; 1°30'-2°40'S,
105°20'-106°00'E; observed by W. L. Abbott (in Lyon, 1906, p. 611), May 20-July 4,
1904.49
Bangkaru, Pulau; INDONESIA: Sumatra; 2°04'N, 97°07'E; pigtail macaques
reportedly absent (Abbott in Miller, 1903a, p. 480). H
Banjak, Kepulauan; INDONESIA: Sumatra; 2°10'N, 97°15'E pigtail macaques
absent (Kloss, [1928], p. 802). G
Banka. See Bangka, Pulau.
Baram River. See Salai, Sungai.
Bareo, 3700 ft.; EAST MALAYSIA: Sarawak; 3°45'N, 115°27'E: collected by T.
Harrisson, Nov. 28, 1947 (FMNH). 75
Bario. See Bareo.
126 FIELDIANA: ZOOLOGY, VOLUME 67
Barito River. See Puruktjahu.
Baru, Danau; INDONESIA: Sumatra; 0°22'S, 102°23'E; collected by G. Schneider
(1905, p. 73), 1897-1899 (museum unknown). 40
Batam, Pulau; INDONESIA: Sumatra; 1°05'N, 104°03'E; reported by C. B. Kloss
i in Lyon, 1907a, p. 657), based on information supplied by local residents. 23
Batangkuwis; INDONESIA: Sumatra; 3°36'N, 98°47'E (Atlas Trop. Ned, pi. 12a);
collected by Widnmann, April, 1907-Aug., 1908 (ZSBS). 28
Bedagi [district]; INDONESIA: Sumatra; ca. 3°20'N, 99°05'E; collected by G.
Schneider ( 1905, p. 73), 1897-1899 (museum unknown). 29
Bedung, Pulau. See Bidong Laut, Pulau.
Bencoolen. See Bengkulu.
Benkoelen. See Bengkulu.
Bengkulu, vicinity; INDONESIA: Sumatra; ca. 3°48'S, 102°16'E; observed by T. S.
Raffles (1821, p. 243), 1817-1820; collected by E. Dubois, date unknown (RMNH),
and H. J. V. Sody, date unknown (RMNH). 42
Benom, Gunong, northeast slope, 2400 ft. and 3600 ft.; WEST MALAYSIA: Pahang;
3°51'N, 102°10'E; reported by Lord Medway (1972, p. 120), 1967-1968. 16
Bentong; WEST MALAYSIA: Pahang; 3°22'N, 101°55'E; collector unknown, Dec.
16, 1912 (BM). 17
Bentong, 6-9 miles W; WEST MALAYSIA: Pahang; 3°32'N, 101°50'E; collected by
C. B. Kloss (1911a, p. 146), June, 1910 (museum unknown). 17
Berhala, Pulau; INDONESIA: Sumatra; 3°46'N, 99°31'E; M. n. nemestrina absent
(Harrison and Hendrickson, 1963, p. 548). P
Bernam, Sungei. See Lima Belas Estate.
Betong. See Pulo Paku.
Betotan; EAST MALAYSIA: Sabah; 5°47'N, 117°52'E; collected by F. N. Chasen
and C. B. Kloss (1931, p. 50), Aug. 14-15, 1927 (ZRCS). 83
Bettotan. See Betotan.
Bidong Laut, Pulau; WEST MALAYSIA: Terengganu; 5°37'N, 103°04'E; M. n.
nemestrina introduced (Kloss, 1911b, p. 176). U
Bintan, Pulau; INDONESIA: Sumatra, 1°05'N, 104°30'E; M. n. nemestrina absent
(Dammerman, 1926, p. 316). 6
Biserat, vicinity; THAILAND: Yala; ca. 6°35'N, 101°20'E; observed by H. C.
Robinson {in Bonhote, 1903, p. 3), 1901-1902. 5
Bohorok, vicinity; INDONESIA: Sumatra; 3°30'N, 98°12'E; collected by G.
Schneider (1905, p. 73), 1897-1899 (museum unknown). 27
Brandan. See Pangkalanberandan.
Btg. Kwis. See Batangkuwis.
Bukit Cheraka Klang. See Jeram.
Bukit Fraser. 5 km. S; WEST MALAYSIA: Selangor; 3041'N, 101°45'E; observed by
M. Kawabe (1970, p. 286), Aug., 1966-March, 1967. 15
FOODEN: LIONTAIL & PIGTAIL MACAQUES 127
Bukit Fraser, northern slopes; WEST MALAYSIA: Pahang; ca. 3°45'N, 101°45'E;
observed by D. Chivers (1971, p. 80; 1973, p. 130) 1968-1970. 15
Bukit Tinggi; INDONESIA: Sumatra; 3°57'N, 98°18'E; collected by G. Schneider
(1905, p. 73), 1897-1899 (museum unknown). 26
Bulan, Pulau; INDONESIA: Sumatra; 0°58'N, 103°55'E; M. n. nemestrina absent
(Dammerman, 1926, p. 316). a
Bulu Telang. See Pulutelang.
Charas Road. See Kuala Lumpur, Cheras Road, 7th mile.
Cheras Road. See Kuala Lumpur, Cheras Road, 7th mile.
Chong; THAILAND: Trang; 7°33'N, 99°47'E; collected by H. C. Robinson and C. B.
Kloss (1910, p. 669), Dec. 14, 1909 (BM). 3
Datu, Pulo. See Datuk, Pulau.
Datuk, Pulau; INDONESIA: Kalimantan; 0°10'N, 108°32'E; monkeys absent
(Abbott in Lyon, 1911, p. 59). j
Deli district; INDONESIA: Sumatra; ca. 3°30'N,98°3O'E; collected by G. Schneider
(1905, p. 73), 1897-1899 (museum unknown); Geryun, 1904 (ZSBS); collector
unknown, 1952 (RMNH); observed by E. Hartert, date unknown (Thomas and
Hartert, 1894, p. 655). 28
Dewhurst Bay. See Kretam Kechil, Sungai.
Dindings. See Hantu, Tanjong.
Djambajan, Sungai. See Djembajan, Sungai.
Djamboe cave, near Tapisello, Padang highlands (Hooijer, 1946, p. 17);
INDONESIA: Sumatra; ca. 0°25'S, 100°30'E; subfossils, prehistoric Holocene;
collected by E. Dubois, 1888-1890 (presumably in RMNH; Hooijer, 1962a, p. 58). 35
Djembajan, Sungai; INDONESIA: Kalimantan; ca. 0°35'S, 116°45'E (Deignan, 1959,
p. 269); collected by H. C. Raven, May 13, 1914 (USNM). 81
"Dolok-Oelve, Deli"; INDONESIA: Sumatra; not precisely located, ca. 3°30'N,
98°30'E; collected by M. Boogaarts, March 3, 1929 (NMS). 28
Durian, Pulau; INDONESIA: Sumatra; 0°42'N, 103°43'E; M. n. nemestrina absent
(Dammerman, 1926, pp. 285, 302). Y
East Perhentian Island. See Perhentian Besar, Pulau.
Engano Id. See Enggano, Pulau.
Enggano, Pulau; INDONESIA: Sumatra; 5°24'S, 102°16'E; pigtail macaques absent
(Kloss, [1928], p. 802). K
Entawa-Samarahan; EAST MALAYSIA: Sarawak; ca. 1°15'N, 111°00'E; collected
by H. C. Robinson, Nov. 23, 1919 (BM). 59
Etawa. See Entawa-Samarahan.
Fraser's Hill. See Bukit Fraser.
Galas, Sungai, west of; WEST MALAYSIA: Kelantan; ca. 5°15'N, 102°00'E;
reported by D. Chivers (1971, p. 80). 13
Gasip, Sungai. See Siak, Sungai.
128 FIELDIANA: ZOOLOGY, VOLUME 67
Gedong; WEST MALAYSIA: Perak; 4°07'N, 101°18'E (Annandale and Robinson,
1903, map); observed by N. Annandale (in Bonhote, 1903, p. 3). 8
Geram. See Jeram.
Great Redang Group. See Redang, Pulau.
Gua Madu; WEST MALAYSIA: Kelantan; ca. 4°50'N, 101°57'E; remains of
monkeys eaten by humans, ca. 8000-2000 B. C; collected by M. W. F. Tweedie
(1940, p. 7), July-Aug, 1939 (probably in National Museum, Singapore). 14
Hantu, Tanjong; WEST MALAYSIA: Perak; 4°19'N, 100°34'E; collected by Langgi,
Aug. 2, 1918 (ZRCS). 9
Huai Lian; THAILAND: Phatthalung; ca. 7°30'N, lOOWE; collected by J. Fooden,
June 28-29, 1973 (FMNH). 4
Jalan Kebun. See Kelang, Jalan Kebun, 7th mile.
Janeng, Gunong. See Janing, Gunong.
Janing, Gunong; WEST MALAYSIA: Johor; 2°31'N, 103°25'E; observed by H. J.
Kelsall (1894, pp. 4, 16), Oct. 20, 1892. 20
Jarak, Pulau; WEST MALAYSIA: Perak; 3°59'N, 100°06'E; M. n. nemestrina
absent (Harrison and Hendrickson, 1963, p. 548). Q
Jeram; WEST MALAYSIA: Selangor; 3°13'N, 101°19'E; collector unknown, Nov. 16,
1910 (ZRCS). 10
Johor, southwest; WEST MALAYSIA: Johor; ca. l°3(m, 103°3O'E; reported by D.
Chivers (1971, p. 80). 21
Ka jan. Sungai. See Peleben.
Kajutanam; INDONESIA: Sumatra; 0°33'S, 100°20'E; collected by P. Ouruans, 1911
(MZB). 36
Kalabakan. See Tibas, Sungai.
Kalulong, Bukit; EAST MALAYSIA: Sarawak; 3°14'N, 114°39'E; collected by C.
Hose, Feb. 1893 (BM). 65
Kampong Durian bei Medan. See Medan.
Kapuas, Sungai; INDONESIA: Kalimantan; ca. 0°, 110°E; collected by L. A. C. M.
Schwaner, May, 1845 (RMNH). 53
Karagan, Sungai. See Karangan, Sungai.
Karangan. Sungai; INDONESIA: Kalimantan; ca. 1°19'N, 117°42'E (Deignan, 1959,
p. 269); collected by H. C. Raven, Dec. 2, 1913 (USNM). 78
Karimon. See Karimun, Pulau.
Karimun. Pulau; INDONESIA: Sumatra; 1°03'N, 103°22'E; M. n. nemestrina absent
(Dammerman, 1926, p. 316). W
Kateman, Sungai; INDONESIA: Sumatra: ca. 0°12'N, 103°20'E; collected by W. L.
Abbott, Aug. 15-27, 1903, Dec. 11. 1905 (USNM). 39
Kelabit Plateau. See Bareo and Pa Umor.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 129
Kelang, Jalan Kebun, 7th mile; WEST MALAYSIA: Selangor; ca. 3°02'N, 101°27'E;
collector unknown, July 22, 1935 (ZRCS). 12
Kendawangan, Sungai; INDONESIA: Kalimantan; ca. 2°15'S, 110°30'E; collected
by W. L. Abbott (in Lyon, 1911, p. 54), June 17-Sept. 29, 1908. 57
Kenepai, Gunung. See Roema Manoeal.
Kepong; WEST MALAYSIA: Selangor; 3°13'N, 101°38'E; collected by A. L. Butler,
date unknown (ZRCS). 18
Keroh Forest Reserve; WEST MALAYSIA: Perak; 4°13'N, 101°07'E; collected by R.
C. Morris (1936, p. 443), June 1-12, 1935 (museum unknown). 8
Khao Si Kaeo; THAILAND: Krabi; ca. 8°12'N, 98°52'E; collected by J. Fooden,
June 6, 1973 (FMNH). /
Kinabalu, Mount. See Lumu Lumu.
Kinabatangan district. See Kretam Kechil, Sungai.
Kinabatangan R[iver]. See Abai.
Kinabatangan, Sungai, 0-245 m; EAST MALAYSIA: Sabah; ca. 5°30'N, 118°0O'E;
observed by M. Kawabe, Oct., 1968-March, 1969 (Kawabe and Mano, 1972, p. 216).
88
Klang. See Kelang, Jalan Kebun, 7th mile.
Kluang; INDONESIA: Sumatra; 2°41'S, 103°54'E; collected by Soekarno, Aug. 21-
29, 1933 (MZB). 46
Klumpeng, Teluk, NW; INDONESIA: Kalimantan; ca. 2°45'S, 116°00'E; observed
by W. L. Abbott (in Lyon, 1911, p. 58), Jan. 8-March 13, 1908. 70
Kota, Danau; INDONESIA: Sumatra; 0°23'S, 102°25'E; collected by G. Schneider
(1905, p. 73), 1897-1899 (museum unknown). 41
Kotawaringin. See Riam
Krakatau, Kepulauan; INDONESIA: Sumatra; 6°07'S, 105°25'E; primates absent
(Dammerman, 1922, p. 65). N
Kretam Kechil, Sungai; EAST MALAYSIA: Sabah; 5°31'N, 118°33'E; collected by
D. D. Davis (1962, p. 58), June 3, 1950 (FMNH). 90
Kroh Reserve. See Keroh Forest Reserve.
Kuala Lompat Post, vicinity; WEST MALAYSIA: Pahang; ca. 3°4(yN, 102°20'E;
observed by Lord Medway and D. R. Wells (1971, p. 246), Feb. 22-26, 1970;
observed by D. Chivers (1971, p. 80; 1973, p. 122), 1968-1970. 16
Kuala Lumpur, Cheras Road, 7th mile; WEST MALAYSIA: Selangor; 3°05'N,
101°45'E; collector unknown, Feb. 8, 1929 (ZRCS). 18
Kuala Lumpur, 22 miles E. See Ulu Gombak Forest Reserve.
?Kuala Lumpur; WEST MALAYSIA: Selangor; ca. 3°10'N, 101°42'E; collector and
date unknown (ZRCS). 18
Kuching, vicinity; EAST MALAYSIA: Sarawak; ca. 1°33'N, HO^CE; collected by
O. Beccari (1904, p. 30) and G. Doria, 1865 (Museo Civico di Storia Naturale
130 FIELDIANA: ZOOLOGY. VOLUME 67
"Giacomo Dona," Genova [DeBeaux, 1923, p. 36]); collected by Bukong, Nov. 23,
1891, March 31, 1892 (SMK); collector unknown, Oct. 2, 1931 (SMK). 50
Kundur, Pulau; INDONESIA: Sumatra; 0°45'N, 103°26'E; M. n. nemestrina absent
(Dammerman, 1926, p. 316). V
Kutai Reserve; INDONESIA: Kalimantan; ca. 0°40'N, 117°40'E; observed by P. S.
Rodman (1973, p. 655), May, 1970-July, 1971. 79
Labuan, Pulau, mainland opposite; EAST MALAYSIA: Sabah; ca. 5°30'N,
115°30'E; collected by H. Low, 1848-1877 (BM). 73
Lamukotan, Pulo. See Lemukutan, Pulau.
Landak, Sungai; INDONESIA: Kalimantan; ca. 0°15'N, 109°45'E; collected by W.
L Abbott (in Lyon, 1907b, p. 548), June-Sept., 1905 (USNM). 52
Lam Phura, Sathani; THAILAND: Trang; 7°40'N, 99°35'E; collected by museum
collectors (Robinson and Kloss, 1910, p. 670), Jan. 20, 1910 (ZRCS). 2
Lampoengsche Distrikten; INDONESIA: Sumatra 4°-6°S, 104°-106°E; zoo specimen,
Aug. 17, 1937 (MZB). (not mapped)
Lam-ra. See Lam Phura, Sathani (Moore and Tate, 1965, p. 324).
Lang Island. See Krakatau, Kepulauan.
Langkat district; INDONESIA: Sumatra; 3°20'-4o20'N, 98o00-98°40,E; observed by
E. Hartert, date unknown (Thomas and Hartert, 1894, p. 655). 27
Langsa, Atjeh; INDONESIA: Sumatra; 4°28'N, 97°58'E; collected by Berthold, Feb.
15, 1935 (MZB). 26
Lankat. See Langkat district.
Larut; WEST MALAYSIA: Perak; 4°48'N, 100°45'E; collector and date unknown
(Taiping Museum [Flower, 1900, p. 315]). 7
Laut. See Lauttador.
Laut, Pulau; INDONESIA: Kalimantan; 3°40'S, 116°10'E; M. n. nemestrina absent
(Abbott in Lyon, 1911, p. 62). /
Lauttador; INDONESIA: Sumatra; 3°18'N, 99°15'E; collected by G. Schneider
(1905, p. 73) 1897-1899 (museum unknown). 29
Lemukutan, Pulau; INDONESIA: Kalimantan; 0°45'N, 108°43'E; M. n. nemestrina
absent (Abbott in Lyon, 1911, p. 59). h
Leuser Reserve; INDONESIA: Sumatra; 2°10'-4o00'N, 97°10'-97°25'E; observed by
F. Kurt (1973, p. 64), April-Aug., 1970. 25
Lida Ajer cave, near Pajakumbuh, Padang highlands (Hooijer, 1946, p. 17);
INDONESIA: Sumatra; ca. 0°14'S, 100°38'E; subfossils, prehistoric Holocene;
collected by E. Dubois, 1888 (presumably in RMNH; Hooijer, 1962a, p. 58). 34
Lima Belas Estate. Sungei Bernam; MALAYSIA: Selangor; 3°46'N, 101°21'E;
observed by I. S. Bernstein (1967a, p. 198), 1965-1966. 10
Lima Bias Estate. See Lima Belas Estate.
Lingga, Pulau; INDONESIA: Sumatra; O^S, 104°35'E; M. n. nemestrina absent
(Dammerman, 1926, p. 316). e
FOODEN: LIONTAIL & PIGTAIL MACAQUES 131
Little Kretam River. See Kretam Kechil, Sungai.
Little Redang Island. See Bidong Laut, Pulau.
Loeboe Basong. See Lubukbasung.
Loeboek Karet; INDONESIA: Sumatra; 2°52'S, 104°18'E (Atlas van Nederlandsch
Oost-Indie. 1901. p. 8); collected by Soekarno, May 5, 1933 (MZB). 46
Loeboek Linggun. See Lubuklinggau.
Long Salai. See Salai, Sungai.
Lubukbasung, vicinity; INDONESIA: Sumatra; ca. 0°20'S, 100°04'E; collected by F.
von Feber, date unknown, (International Colonial Exhibition, Amsterdam
[Jentink, 1883, p. 171]). 32
Lubukbnggau, sea level; INDONESIA: Sumatra; 3°18'S, 102°52'E; collected by J. J.
Menden, Nov. 30, 1933 (AMNH-AC). 44
Lumu Lumu, 5500 ft; EAST MALAYSIA: Sabah; ca. 6°02'N, 116°34'E; collected by
J. A. Griswold, Jr. (1939, pp. 403, 506), July 20, 1937 (MCZ). 82
Lungmanis Station; EAST MALAYSIA: Sabah; ca. 5°40'N, 117°45'E; observed by
K. Stott, Jr. (1964, p. 12), date unknown. 83
Madihit, Sungai; EAST MALAYSIA: Sarawak; 4°11'N, 115°08'E; museum collector,
June, 1911 (SMK). 74
Malacca. See Melaka.
Mansalar Id. See Musala, Pulau.
Mara. See Peleben.
Masalembo-Besar; INDONESIA: Kalimantan; 5°34'S, 114°26'E; monkeys absent
(Abbott in Lyon, 1911, p. 61). k
Masolombo Besar. See Masalembo-Besar.
Medan, vicinity; INDONESIA: Sumatra; ca. 3°35'N, 98°40'E; collected by H. Durk.
Dec. 29, 1904-Jan. 5, 1905 (ZSBS); Widnmann, 1908 (ZSBS). 28
Melaka; WEST MALAYSIA: Melaka; ca. 2°12'N, 102°15'E; reported by N.
Annandale {in Bonhote, 1903, p. 3) and E. J. H. Corner (1941, p. 14). 19
Meru; WEST MALAYSIA: Selangor; 3°08'N, 101°26'E; observed by I. S. Bernstein
(1967b, p. 220), May 4, 1965- Feb. 28, 1966. 12
Moyen Bahau. See Bahau, Sungai.
Muarabehti, sea level; INDONESIA: Sumatra; 3°15'S, 103°02'E; collected by J. J.
Menden, Nov. 26, 1933 (AMNH-AC). 44
Muarabliti. See Muarabeliti.
Musala, Pulau; INDONESIA: Sumatra; 1°38'N, 98°32'E; pigtail macaques absent
(Kloss, [1928], p. 802). J
Natuna, Kepulauan; INDONESIA: Sumatra; 4°00'N, 108°15'E; M. n. nemestrina
introduced (Thomas and Hartert, 1894, p. 654). g
Niah Caves; EAST MALAYSIA: Sarawak; 3°49'N, 113°47'E; subfossils, late
Pleistocene-subrecent; collected by Sarawak Museum, 1954-1962 (Hooijer, 1962b, p.
445). 63
132 FIELDIANA: ZOOLOGY, VOLUME 67
Nias, Pulau; INDONESIA: Sumatra; 1°05'N, 97°35'E; pigtail macaques absent
(Kloss, [1928], p. 802). /
Nicobar Islands, northern. See under Macaca n. leonina. C
Nicobar Islands, southern. See under Macaca n. leonina. D
Ogan, Air, upper; INDONESIA: Sumatra; ca. 3°30'S, 104°40'E; collected by
Chancellor Stuart Exp., 1929 (FMNH). 48
Padang; INDONESIA: Sumatra; 0°57'S, 100o21'E; collected by S. Muller, 1836
(RMNH); H. Meyer, 1890 (ZMUZ); I. Hagenbeck, May 2, 1925 (NMS). 33
Padang [district]; INDONESIA: Sumatra; ca. 3°15'N, 99°15'E; collected by G.
Schneider (1905, p. 73), 1897-1899 (museum unknown). 29
Padang highlands. See Djamboe cave, Lida Ajer cave, and Sibrambang cave.
Pagar, Tanjong. See Singapore Island.
Pagaralam; INDONESIA: Sumatra; 4°01'S, 103°16'E; collected by A. V. A. Coulin,
June 19, 1941 (MZB). 45
Pagurawan. See Paguruan, Sungai.
Paguruan, Sungai; INDONESIA: Sumatra; 3°26'N, 99°20'E; collected by G.
Schneider (1905, p. 73), 1897-1899 (museum unknown). 29
Pajakombo. See Pajakumbuh.
Pajakumbuh; INDONESIA: Sumatra; 0°14'S, 100°38'E; collected by E. Dubois, date
unknown (RMNH). 34
Pajakumbuh, near. See Lida Ajer cave.
Palembang; INDONESIA: Sumatra; 2°55'S, 104°45'E; collector unknown, Feb. 4,
1921 (RMNH); collector unknown, March 12, 1924 (NHMBe). 47
Palembang [district], sea level; INDONESIA: Sumatra; "3°S, 104°E(appr; collected
by W. J. Frost, April 20, 1938 (BM). 46
Pamarkang Bay. See Pamukan, Teluk.
Pamukan, Teluk; INDONESIA: Kalimantan; ca. 2°30'S, 116°15'E; collected by W.
L. Abbott (in Lyon, 1911, p. 58), March 25, 1909 (USNM). 69
Panaitan, Pulau; INDONESIA: Java; 6°32'-6°40'S, 105°04'-105°14'E; M. n.
nemestrina apparently absent (Hoogerwerf, 1953, p. 492). M
Pangkalanberandan; INDONESIA: Sumatra; 4°01'N, 98°17'E; collected by Wilhelm
Th. A. H. Volz (1912, p. 22), Oct. 14- Nov. 10, 1904 (ZMB). 26
Pangkor, Pulau; WEST MALAYSIA: Perak; 4°13'N, 100°34'E; M. n. nemestrina
absent (Harrison and Hendrickson, 1963, p. 548). 0
Papar; EAST MALAYSIA: Sabah; 5°44'N, 115°56'E; collected by R. E. Kuntz,
Sept. 12, 1960 (USNM). 72
Pasi; INDONESIA: Sumatra; 5°12'N, 95°17'E; collected by Lt. v. d. Rest, date
unknown (MZB). 24
Pa Umor, 3400 ft.; EAST MALAYSIA: Sarawak; 3°44'N, 115031'E; collected by T.
Harrisson, Jan. 3, 1948 (FMNH). 75
FOODEN: LIONTAIL & PIGTAIL MACAQUES 133
Pa Umur. See Pa Umor.
Pelaihari; INDONESIA: Kalimantan; 3°48'S, 114°45'E; collected by J. H. F.
Kohlbrugge (1896, p. 181), Dec 15-21, 1894 (museum unknown). 71
Peleben [on Sungai Kajan near Mara; MS notes in AMNH, reported by Dr. Joseph
Curtis Moore, personal communication]; INDONESIA: Kalimantan; ca. 2°45'N,
117°15'E; collected by V. von Plessen, June 24-July 21, 1935 (AMNH-AC; ZMB).
77
Pendeng. See Pulau Munteh.
Penang. See Pinang, Pulau.
Pennan, Koh. See Phangan, Ko.
Perak, Upper; WEST MALAYSIA: Perak; ca. 5°00'N, 101°0CE; reported by N.
Annandale (in Bonhote, 1903, p. 3). 7
Perhentian Besar, Pulau; WEST MALAYSIA: Terengganu; 5°54'N, 102°46'E;
macaques absent (Kloss, 1911b, p. 180). S
Phangan, Ko; THAILAND: Surat Thani; 9°45'N, lOOWE; "M. n. nemestrina is
stated on native authority to be found. ..Captive specimens were seen but their
provenance was uncertain and they had not improbably been brought from the
mainland" (Robinson and Kloss, 1914b, p. 130). (not mapped)
Pinang, Pulau; WEST MALAYSIA: Pinang; 5°24'N, 100°14'E; collected by T.
Cantor (1846, p. 177; museum unknown); v. Duben, 1846 (NHRM); W. Riitimeyer,
1880 (NHMB); Wistar Inst, of Anat., 1900 (USNM); collector and date unknown
(BM). 6
Pleihari. See Pelaihari.
Pohorok. See Bohorok.
Pontianak; INDONESIA: Kalimantan; 0°02'S, 109°2O'E; collected by P. Diard,
[1826], (RMNH). 54
Priaman [district]. See Lubukbasung.
Pulau Munteh, 550-750 m.; INDONESIA: Sumatra, 4°05'N, 97°30'E (Chasen, 1941,
map opp. p. 1); collected by A. Hoogerwerf (in Chasen, 1940b, p. 485), May 1, 1937
(MZB). 25
Pulo Paku, Betong Saribas; EAST MALAYSIA: Sarawak; ca. 1°30'N, 111°15'E;
collected by H. C. Robinson, Oct. 15-Nov. 25, 1916 (BM). 58
Pulutelang; INDONESIA: Sumatra; 3°53'N, 98°20'E; collected by G. Schneider
(1905, p. 73), 1897-1899 (NHMB). 26
Puruktjahu; INDONESIA: Kalimantan; 0°35'S, 114°35'E; collected by G. C.
Shortridge, Sept. 3, 1909 (BM). 67
Ranau, 1350 ft.; EAST MALAYSIA: Sabah; 5°58'N, 116°41'E; collected by R. E.
Kuntz, Sept., 1960 (USNM). 82
Redang, Pulau, group; WEST MALAYSIA: Terengganu; 5°4TN, 103°01'E; M. n.
nemestrina absent (Kloss, 1911b, p. 177). T
Rengam. See Sungai Renggam.
134 FIELDIANA: ZOOLOGY, VOLUME 67
Rengat, inland; INDONESIA: Sumatra; ca. 0°24'S, 102°33'E; collected by G.
Schneider (1905, p. 73), 1897-1899 (museum unknown). 41
Riam, Kotawaringin district, 300 m.; INDONESIA: Kalimantan; 1°50'S, 111°54'E;
collected by J. J. Menden, Nov. 6, 1935 (AMNH-AC). 56
Ringat. See Rengat.
Roema Manoeal; INDONESIA: Kalimantan; 0°46'N, 111°39'E; collected by J.
Biittikofer (1897, p. 12), Dec. 26, 1893 (RMNH; specimen not seen, data from
Jentink, 1897, p. 39). 60
Rungkup. See Simpang Empat Rungkup.
Sagamafluss. See Segama, Sungai.
Sakaiam River. See Sekajam, Sungai.
Salai, Sungai; EAST MALAYSIA: Sarawak; ca. 3°30'N, 114°30'E; captive presented
to London Zoo by F. Day in 1892, originally obtained about 1888 (Sclater, 1893, p.
325) (BM); collected by C. Hose, Feb.-June, 1891 (SMK). 64
Samarahan. See Entawa-Samarahan.
Sandakan, vicinity; EAST MALAYSIA: Sabah; ca. 5°50'N, 118°07'E; collected by
H. G. Deignan, June 10, 1937 (MCZ). For note on locality, see Coolidge (1940. p.
130). 87
Sanggul, Bukit, 500 m.; INDONESIA: Sumatra; 3°50'S, 102°37'E; collected by J. J.
Menden, Aug. 20-Sept. 2, 1936 (AMNH-AC; MZB). 43
Sapagaya, Sungai; EAST MALAYSIA: Sabah; ca. 5°39'N, 118°06'E; collected by C.
F. Adams, Nov. 21, 1887 (USNM). 84
Saribas. See Pulo Paku.
Sebang. See Sebangka Pulau.
Sebangka, Pulau; INDONESIA: Sumatra; 0°07'N, 104°36'E; M. n. nemestrina
absent (Dammerman, 1926, p. 316). c
Sebesi, Pulau; INDONESIA: Sumatra; 5°57'S, 105°30'E; primates absent
(Dammerman, 1922, p. 75). L
Sebesy. See Sebesi, Pulau.
Sebuku, Pulau; INDONESIA: Kalimantan; 3°30'S. 116°22'E; M. n. nemestrina
absent (Abbott in Lyon, 1911, p. 62). m
Segama, Sungai; EAST MALAYSIA: Sabah; ca. 5°30'N. 118°30'E; collected by Dr.
Pagel, Aug. 22, 1907 (ZMB). 90
Sekajam, Sungai; INDONESIA: Kalimantan; probably about 0°50'N, 110°25'E;
collected by W. L. Abbott (in Lyon. 1907b, pp. 548, 566). June-Sept., 1905 (USNM).
51
Sembilan, Kepulauan; WEST MALAYSIA: Perak; 4°02'N. 100°33'E; At. n.
nemestrina absent (Harrison and Hendrickson. 1963. p. 548). R
Semitau, vicinity; INDONESIA: Kalimantan; ca. 0°33'N. 111°58'E; observed by J.
Biittikofer (1897, p. 11; in Jentink, 1897, p. 39) Nov.- Dec., 1893. 61
FOODEN: LIONTAIL & PIGTAIL MACAQUES 135
Serapit; INDONESIA: Sumatra 3°34'N, ^l^E; collected by G. Schneider (1905, p.
73), 1897-1899 (museum unknown). 27
Serdang [district]; INDONESIA: Sumatra; ca. 3°30'N, 98°50'E; observed by E.
Hartert, date unknown (Thomas and Hartert, 1894, p. 655). 29
Siak, Sungai, lower, at mouth of Sungai Gasip; INDONESIA: Sumatra; 0°43'N,
101°42'E; collected by W. L. Abbott, Jan. 1, 1907 (USNM). 38
Sibolga, Teluk; INDONESIA: Sumatra; 1°38'N, 98°45'E; collected by W. L. Abbott
(in Miller, 1903a, p. 482), Feb. 19-March 25, 1902 (USNM). 30
Sibrambang cave, Padang highlands; INDONESIA: Sumatra; not precisely located,
ca. 0°30'S, 100°30'E; subfossils, prehistoric Holocene; collected by E. Dubois, 1888-
1890 (presumably in RMNH; Hooijer, 1962a, p. 58). 35
Sidong, Gunong, 1200 ft.; EAST MALAYSIA: Sarawak; ca. 1°05'N, 110°25'E
(Medway, 1965, map); collector unknown, Oct. 10, 1910 (ZRCS). 50
Simeulue, Pulau; INDONESIA: Sumatra; 2°33'N, 95°55'E; pigtail macaques absent
(Abbott in Miller, 1903a, p. 479). F
Simpang Empat Rungkup; WEST MALAYSIA: Perak; 3°57'N, 100°43'E; collected
by E. S[eimund], Nov. 28, 1913 (BM). 9
Singapore Island; SINGAPORE: 1°20'N, 103°50'E; M. n. nemestrina introduced
(Ridley, 1895, p. 26; Annandale in Bonhote, 1903, p. 3). (not mapped)
Singkep, Pulau; INDONESIA: Sumatra; 0°30'S, 104°25'E; M. n. nemestrina absent
(Dammerman, 1926, p. 316). /
Sintang; INDONESIA: Kalimantan; 0°04'N, 111°30'E; collected by C. Bruegel,
March 18, 1907 (ZSBS). 62
Sim ass. See Sekajam.
Sirambas; INDONESIA: Sumatra; 0°49'N, 99°32'E; collected by E. Modigliani, date
unknown (Museo Civico di Storia Naturale "Giacomo Doria," Genova [DeBeaux,
1923, p. 36]). 31
Si-Rambe. See Sirambas.
Smitau. See Semitau.
Solok; INDONESIA: Sumatra; 0°48'S, 100°39'E; collected by P. O. Stolz, July 17,
1915 (RMNH). 36
Solombo. See Masalembo-Besar.
Sugi, Pulau; INDONESIA: Sumatra; 0°50'N, 103°47'E; M. n. nemestrina absent
(Dammerman, 1926, p. 316). Z
Sukadana; INDONESIA: Kalimantan; 1°15'S, 109°57'E; collected by W. L. Abbott
(in Lyon, 1911, p. 54), June 10, 1907 (USNM). 55
Sungai Buloh, 150-300 ft.; WEST MALAYSIA: Selangor; 3°12'N, 101°35'E; collected
by Institute for Medical Research, 1947-1957 (Harrison and Hendrickson, 1963, p.
548; Harrison, 1969, p. 176). 11
Sungai Renggam; WEST MALAYSIA: Selangor; 3°03'N, 101°32'E; collector
unknown, May 28, 1923 (BM). 12
136 FIELDIANA: ZOOLOGY, VOLUME 67
Sungei Rengam. See Sungai Renggam.
Sut, [Sungai], 3000 ft.; EAST MALAYSIA: Sarawak; 2°42'N, 114°39'E; coUected by
T. A. Chavasue, Sept. 27, 1955 (FMNH). 66
Tabong; INDONESIA: Kalimantan; 0°34'N, 116°02'E; collected by A. M. R.
Wegner, Sept. 13, 1956 (MZB). 80
Tandjong. See Tandjung.
Tandjung; INDONESIA: Kalimantan; 2°11'S, 115°23'E; coUected by A. Buxtorf,
1905 (NHMB). 68
Tandjung; INDONESIA: Sumatra; 3°17'N, 99°19'E; collected by G. Schneider (1905,
p. 73), 1897-1899 (museum unknown). 29
Tandjungmorawa; INDONESIA: Sumatra; 3°30'N, 98°47'E (Atlas Trop. Ned., pi.
12a); collected by B. Hagen (1890 pp. 80, 82), 1882-1883 (RMNH; also see Jentink,
1887, p. 26); C. Bruegel, 1906 (ZSBS). 28
Tanjong Antu. See Hantu, Tanjong.
Tapanuli Bay. See Sibolga, Teluk.
Tapisello. See Djamboe cave.
Tarusan, Teluk; INDONESIA: Sumatra; 1°13'S, 100°25'E; collected by W. L.
Abbott, Dec. 31, 1904 (USNM). 37
Tawau district. See Tibas, Sungai.
Tebingtinggi, Deli; INDONESIA: Sumatra; 3°20'N, 99°09'E; coUected by J. A.
Coenraad, Nov., 1931, Feb., 1932 (MZB). 29
Temadju, Pulau; INDONESIA: Kalimantan; 0°29'N, 108°52'E; monkeys absent
(Abbott in Lyon, 1911, p. 59). i
Temaju, Pulo. See Temadju, Pulau.
Tibas, Sungai; EAST MALAYSIA: Sabah; 4°26'N, 117°29'E (Davis, 1962, p. 127);
collected by R. F. Inger, June 9, 1956 (FMNH). 86
Tinggi, Pulau; WEST MALAYSIA: Johor; 2°18'N, 104°07'E; M. n. nemestrina
absent (Robinson, 1919, p. 325). X
Tioman, Pulau; WEST MALAYSIA: Pahang; 2°48'N, 104°11'E; M. n. nemestrina
said to be absent (Abbott in Miller, 1900, p. 246); collected by J. L. Harrison and J.
R. Hendrickson (1963, p. 548), May, 1958 (museum unknown). 22
Trang. See Chong.
Trang [province]; THAILAND: Trang; ca. 7°30'N, 99°30'E; collector and date
unknown (ZRCS). (not mapped)
Ulu Gombak Forest Reserve, 2000 ft.; WEST MALAYSIA: Selangor; ca. 3°20'N,
lOrsCE; observed by H. E. McClure ([1965], p. 57), May 20, 1961, March 10, 1962;
reported by D. Chivers (1971, p. 80). 18
Ulu Madihih. See Madihit, Sungai.
Ulu Segama Forest Reserve, 800-5000 ft.; EAST MALAYSIA: Sabah; ca. 5°11'N,
117°53'E; observed by J. MacKinnon (1971, p. 153), 1968-1970. 85
Ulu Sempan, Raub. See Bukit Fraser.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 137
Verlaten Island. See Krakatau, Kepulauan.
Macaca nemestrina leonina
Map, Figure 28
Adung Valley. See Nam Tamai.
Akyab. See Sittwe.
Andaman Islands; INDIA: Andaman and Nicobar Islands; 10°-14°N, 92°-93°E;
monkeys not indigenous, M. n. leonina introduced (Hamilton, 1870, p. 220). A
Annam. See Da Ban.
Arakan district, northern hills; BURMA: Arakan Div.; ca. 21°30'N, 93°00'E;
collected by A. P. Phayre, 1844 (ZSI); Darling, date unknown (ZMB); collector
and date unknown (RMNH); reported by Sclater ([1881], p. 537). 4
Arracan. See Arakan.
Assam [state]; INDIA: Assam; 25°-27°N, 90°-95°E; obtained by Mr. Gerrard,
purchased March 17, 1885 (IRSN). (not mapped)
Bang Hue Horn. See Ban Hue Horn.
Ban Huai Maenam Noi, vicinity, ca. 75 m.; THAILAND: Kanchanburi; ca. 14°25'N,
98°51'E; collected by J. Fooden (1971a, p. 32), Feb. 16, 1967 (CTNRC, FMNH). 31
Ban Huang Som; THAILAND: Trat; 11°51'N, 102°5O'E; collected by C. B. Kloss,
(1916a, pp. 30, 66), Jan. 11, 1915 (BM). 37
Ban Hue Horn, hill forests north of; THAILAND: Phrae; ca. 17°55'N, 100°10'E;
reported by N. Gyldenstolpe (1914, p. 5). 10
Bankachon, Victoria Point; BURMA: Tenasserim Div.; 10°09'N, 98°36'E; collected
by G. C. Shortridge (in Wroughton, 1915b, p. 696), Nov. 11-Dec. 29, 1913 (BM;
BNHS;ZSI). 44
Bankasun. See Bankachon.
Ban Kerng Chada, ca. 150 m.; THAILAND: Kanchanaburi; 15°08'N, 98°31'E;
collected by J. Fooden (1971a, p. 32), Feb. 5, 1967 (FMNH). 29
Ban Khlong Wan; THAILAND: Ranong; lO'WN, 98°49'E; collected by H. C.
Robinson and C. B. Kloss (1921, p. 9), March 16, 1919 (ZRCS). 46
Ban Nong Kho; THAILAND: Chon Buri 13°09'N, 101°04'E; collected by H. M.
Smith, Nov. 13, 1926 (USNM). 35
Ban Nong Kok; THAILAND: Krabi; 8°06'N, 98°52'E; collected by H. C. Robinson
and C. B. Kloss (1919, p. 88), Jan. 7, 1918 (ZRCS). 48
Ban Pong Nam Ron, 8-12 km. W and 10 km. NW, ca. 250-300 m.; THAILAND:
Kamphaeng Phet; ca. W^N, 99°15'E; collected by J. Fooden (1971a, p. 32), April
12-23, 1967 (FMNH). 21
Ban Tha San, 220 ft.; THAILAND: Chumphon; 10°29/N, 98°55'E; collected by H. C.
Robinson and C. B. Kloss (1921, p. 10), March 14, 1919 (ZRCS). 46
Barren Island; INDIA: Andaman and Nicobar Islands; 12°15'N, 93°50'E; monkeys
absent (Kloss, 1903, p. 14). B
138 FIELDIANA: ZOOLOGY, VOLUME 67
Bhamo, 20-25 miles below; BURMA: Kachin State; ca. 24°10'N, 97°00'E; erroneous
record (Anderson, 1881, p. 72) of M. n. leonina (see above, p. 110).
Bien Hoa. See Trang Bom.
Bolovens, Plateau des; LAOS: 14°45'-15°30'N, 106°00'-106o50'E; collected by T. D.
Carter, Legendre Indo-China Expedition, Feb. 3-13, 1932 (AMNH). For locality
note, see Legendre (1932, p. 495). 17
Chamchay, Cochinchine; SOUTH VIETNAM: not precisely located, 8°30'-12°30'N,
104°30'-107°30'E; collected by M. Pierre, April 1874 (MNHN). (not mapped)
Champang; BURMA: Tenasserim Div.; 10°13'N, 98°31'E; collected by W. L. Abbott,
Dec. 21-22, 1903 (USNM). 44
Chan, Ko; THAILAND: Ranong, Mergui Archipelago; 9°25'N, 97°50'E (Moore and
Tate, 1965, p. 323); collected by W. L. Abbott, Dec. 30, 1899-Jan. 1, 1900 (USNM ).
45
Chance Island. See Chan, Ko (Moore and Tate, 1965, p. 317).
Changchang Pani, 500 ft.; INDIA: Nagaland; 26°35'N, 94°25'E; collected by C.
McCann (1933a, p. 395), Feb. 7-19, 1930 (AMNH; BNHS). 1
Chantabun. See Chantaburi.
Chantaburi; THAILAND: Chantaburi; 12°36'N, 102°09'E; collected by C. J.
Aagaard, Dec. 16-17, 1926 (ZRCS). 36
Chongkrong, 600-900 m.; THAILAND: Kanchanaburi; 14°41'N, 98°52'E; coUected by
J. Fooden (1971a, p. 32), Jan. 28, 1967. 30
Cochinchine. See Chamchay.
Da Ban; SOUTH VIETNAM: Ninh Thuan; ca. 11°45'N, 108°45'E, collected by C. B.
Kloss (1919a, p. 395), March 24, 1918 (ZRCS). 41
Ghirbi. See Ban Nong Kok.
Hin Lap, Sathani; THAILAND: Sara Bun; 14°40,N, 101°09'E; coUected by H. M.
Smith, Oct. 2, 1932 (USNM). 24
Hkamti. See Singkaling Hkamti.
Hue\ vicinity; SOUTH VIETNAM: Thua Thien; 16°28'N, 107°36'E; reported by J.
Delacour (1940, p. 24). 19
Junk Seylon. See Klong Tung Sai.
Kao Sabab. See Sa Bap, Khao.
Kawkareik; BURMA: Kawthule State; 16°33'N, 98°14'E; collected by L. Fea, May
1887 (Museo Civico di Storia Naturale "Giacomo Doria," Genova [Thomas, 1892,
p. 916; DeBeaux, 1923, p. 34]). 20
Khlung, Khlong, upper; THAILAND: Kamphaeng Phet; ca. 16°05'N, 99°20'E;
collected by C. C. Sanborn (1952, p. 2) and F. C. Wonder, July 2, 1949 (FMNH). 22
Khun Tan, mountains; THAILAND: Lamphun; ca. 18°30'N, 99°15'E; coUected by
N. Gyldenstolpe ([1917], p. 7), June 7, 1914 (NHRM). 9
Klet Kaeo, Ko; THAILAND: Chon Bun; 12°46'N, 100°51'E; pigtail macaques absent
(Berkson et al., 1971, p. 237). F
FOODEN: LIONTAIL & PIGTAIL MACAQUES 139
Klong Klung. See Khlung, Khlong.
Klong Menao. See Ban Huang Som (Moore and Tate, 1965, p. 323).
Klong Tundai. See Klong Tung Sai.
Klong Tung Sai, Ko Phuket; THAILAND: Phuket; 7°98'N, ^^E (Robinson and
Kloss, 1919, p. 88); collected by C. B. Kloss, Dec. 20-26, 1917 (BM; ZRCS). For
comment on locality, see Chasen, 1935, p. 38. 47
Klong Tun Sai. See Klong Tung Sai.
Klong Wan. See Ban Khlong Wan.
Kokareet. See Kawkareik.
Ko Keow, vicinity, ca. 200 m.; THAILAND: Kamphaeng Phet; ca. 15°55'N, 99°25'E;
J. Fooden (1971a, p. 32), March 9, 1967 (FMNH).22
Koon Tan. See Khun Tan.
Krabi. See Ban Nong Kok.
Lampi Island. See Lanbi Kyun.
Lanbi Kyun; BURMA: Tenasserim Div., Mergui Archipelago; 10°50'N, 98°15'E;
collected by C. Primrose, March 3, 1922 (BM). 42
Lat Bua Kao. See Lat Bua Khao.
Lat Bua Khao, Sathani, north of; THAILAND: Nakhon Ratchasima; 14°52'N,
101°36'E; collected by C. B. Kloss (1919b, p. 343), Oct. 12, 1916 (USNM). 26
Meh Lem river, north of; THAILAND: Phrae; ca. 18°25'N, 100°20'E (Gyldenstolpe,
1913, p. 4; 1916 pl. 1); observed and tentatively identified by N. Gyldenstolpe
(1914, p. 4), March 5-27, 1912. ;/
Meng-hai; CHINA: Yunnan; 21°52'N, 100°28'E; collected by Kao Yuen-ting et al.
(1962, p. 188), 1957-1958 (probably in collection of Institute of Zoology, Academia
Sinica, Peking). 8
Mong Nai; BURMA: Shan State; 20°31'N, 97°52'E; collected by Dr. Griggs, in or
before April, 1894 (Brown, 1896, p. 485). 7
Muang Leip. See Muang Liap.
Muang Liap; LAOS: Xaignabouri; 18°29'N, 101 °40'E; collected by J. Bangga, Jan.
15, 1920 (ZRCS). 12
Nam Tamai; BURMA: Kachin State; 27°42'N, 97°54"E (Moore and Tate, 1965, p.
328); probably erroneous record (Dollman, 1932, p. 9; see above p. 112).
Nicobar Islands, northern; INDIA: Andaman and Nicobar Islands; 8°-10°N, 92°-
94°E; monkeys absent (Kloss, 1903, p. 114). C
Nicobar Islands, southern; INDIA: Andaman and Nicobar Islands; 7°-8°N, 93°-94°;
pigtail macaques absent (Kloss, [1928], p. 802). D
Nong Khor. See Ban Nong Kho (Chasen, 1935, p. 32; Moore and Tate, 1965 p. 329).
Nong Kok. See Ban Nong Kok.
Pak Chong, Sathani, 900 ft.; THAILAND: Nakhon Ratchasima: 14°42'N, 101°25'E;
140 FIELDIANA: ZOOLOGY, VOLUME 67
collected by E. G. Herbert and M. Smith, Dec. 4, 1915 (BM; Kloss, 1916b, p. 2) and
H. M. Smith, March 2, 1924 (USNM). 25
Pak Jong. See Pak Chong, Sathani.
Paksane (province); LAOS: ca. WSCN, 104°00'E; collected by J. Deuve and M.
Deuve (1963, p. 60), date and museum unknown. 14
Pakse (province); LAOS: ca. 15°00'N, 106°00'E; collected by J. Deuve and M. Deuve
(1963, p. 60), date and museum unknown. 17
Panjang, Pulau. See Telok Poh.
Pan-na-meng-hai. See Meng-hai.
Paungdaw power station, hill NW, 3000 ft.; BURMA: Tenasserim Div.; ca. 14°00'N,
98°30'E; collected by J. Keenan, July 12, 1961 (BM). 32
"Perak"; WEST MALAYSIA: Perak; ca. 5°00'N, 101°00'E; presented by O. L.
Fraser, Jan. 6, 1878 (ZSI). Erroneous locality (see above, p. 110).
Petchaburi. See Phet Buri.
Phet Buri [province]; THAILAND: Phet Buri; 12°35'-13°10'N, 99°15'-100°05'E;
reported by K. G. Gairdner (1914, p. 36). 34
Phu Kheo Forest Reserve; THAILAND: Chaiyaphum; ca. 16°15'N, 101°30'E;
remains of monkeys eaten by humans, reported by J. A. McNeely and E. W.
Cronin (1972, p. 459), Jan. 1972. 27
Phuket, Ko. See Klong Tung Sai.
Pidaung Game Sanctuary; BURMA: Kachin State; 25°25'N, 97°09'E; observed by U
Tun Yin (1954, p. 269), March, 1951-Feb., 1952. 3
Plateau des Bolovens. See Bolovens, Plateau des.
Puket. See Klong Tung Sai.
Rat Buri [province]; THAILAND: Rat Buri; 13°10'-13°45'N, 99°20'-100°05'E;
reported by K. G. Gairdner (1914, p. 36). 33
Red Point; BURMA: Tenasserim Div.; 10°40'N, 98°30'E (Moore and Tate, 1965, p.
331); collected by W. L. Abbott, Feb. 20, 1904 (USNM). 43
Ross Island. See Andaman Islands.
Sa Bap, Khao; THAILAND: Chanthaburi; 12°31'N, HE'lffE; collected by H. M.
Smith, Nov. 5, 1933 (USNM). 36
Sai Gon; SOUTH VIETNAM: Gai Dinh; 10°45'N, 106°40'E; reported by J. Delacour
(1940, p. 24). 40
Salanga. See Klong Tung Sai.
Samnak Rabam, 10 km. SE, ca. 200 m.; THAILAND: Uthai Thani; 15°25'N,
99°30'E; collected by J. Fooden (1971a, p. 32), Feb. 26, 1967 (CTNRC, FMNH). 23
Saravane (province); LAOS: ca. 15°45'N, 106°45'E; collected by J. Deuve and M.
Deuve (1963, p. 60), date and museum unknown. 18
Savannahket (province); LAOS: ca. 16°30'N, 104°45'E; collected by J. Deuve and M.
Deuve (1963, p. 60), date and museum unknown. 16
FOODEN: LIONTAIL & PIGTAIL MACAQUES 141
Selok Poh. See Telok Poh.
"Siam"; THAILAND: no exact locality; collected by R. C. Bulkeley, date unknown
(AMNH). (not mapped)
Singkaling Hkamti, about 20 miles north [east], W bank of Chindwin River, 500 ft.;
BURMA: Sagaing Div.; ca. 26°10'N, 96°00'E; collected by G. C. Shortridge and S.
A. Macmillan, July 26, 1914 (BM). For locality note, see Wroughton, 1916a, p. 293.
2
Sittwe, hills inland; BURMA: Arakan Div.; ca. 20°00'N, 93°30'E; reported by S. R.
Tickell (1854-1863, MS., p. 112; Pocock, 1939, p. 60) and E. Blyth (1863, appendix p.
[189]; in Dunn, 1864, p. 370). 5
Sullivan's Island. See Lanbi Kyun.
Taho, 1200-1300 m.; BURMA: Shan State; 19°27'N, 96°53'E; collected by L. Fea
(1888, pp. 855, 859), Feb. 29, 1888 (Museo Civico di Storia Naturale "Giacomo
Doria," Genova [Thomas, 1892, p. 916; DeBeaux, 1923, p. 34]). 6
Tapli. See Ban Khlong Wan.
Tasan. See Ban Tha San.
Tavoy. See Ye.
Tay Ninh; SOUTH VIETNAM: Tay Ninh; 11°18'N, 106°05'E, reported by A.
Morice(1875, p. 41). 38
Telok Besar; BURMA: Tenasserim Div.; 10°22'N, 98°35'E (Moore and Tate, 1965, p.
333); collected by W. L. Abbott, March 17, 1904 (USNM). 44
Telok Poh, Ko Yao Yai; THAILAND: Phangnga; 8°00'N, 98°38'E (Robinson and
Kloss, 1919, p. 88); collected by C. B. Kloss, Jan. 22, 1918 (BM). For comment on
locality, see Chasen, 1935, p. 38. 47
Thakhek (province); LAOS: ca. 17°30'N, 105°00'E; collected by J. Deuve and M.
Deuve (1963, p. 60), date and museum unknown. 15
Thad. See Taho.
Tongha. See Klong Tung Sai.
Tongka. See Klong Tung Sai.
Trang Bom; SOUTH VIETNAM: Bien Hoa, 10°57'N, W7°WE; collected by C. B.
Kloss (1919a, p. 401), June 5, 1918 (ZRCS); M. Poilane, Sept. 4, 1932 (USNM; for
locality restriction, see Van Peenen et al., 1969, p. 102). 39
Victoria Point. See Bankachon.
Vientiane (province); LAOS: ca. 18°00'N, 102°3O'E; collected by J. Deuve and M.
Deuve (1963, p. 60); date and museum unknown. 13
Yao Yai, Ko. See Telok Poh.
Ye, forest near; BURMA: Tenasserim Div.; ca. 15°15'N, 98°00'E; collected by S. R.
Tickell (1854-1875, MS., pp. 112, 137, 139), Nov., 1854 and Jan.-Feb., 1860 (museum
unknown). 28
Yey. See Ye.
Zingkalling. See Singkaling Hkamti.
142 FIELDIANA: ZOOLOGY, VOLUME 67
Macaca nemestrina pagensis
Map, Figure 25
Mentawai, Kepulauan. See Sipura, Pulau; Pagai Utara, Pulau; Pagai Selatan, Pulau.
North Pagi. See Pagai Utara, Pulau.
Pagai Selatan, Pulau, sea level; INDONESIA: Sumatra; ca. 3°00'S, 100°20'E;
collected by W. L. Abbott, Nov. 17, 1902 (USNM) and J. J. 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.
REFERENCES
Ali, S. and H. Santapau
1956. Distribution of the liontailed monkey, Macaco, silenus (Linnaeus). J. Bombay
Nat. Hist. Soc, 53, p. 687.
Allen, G. M. and H. J., Coolidge, jr.
1940. Mammals of the Asiatic Primate Expedition. Bull. Mus. Comp. Zool.
Harvard., 87 (3), pp. 131-166
Allen, J. A.
1916. The proper generic name of the macaques. Bull. Amer. Mus. Nat. Hist., 35,
pp. 49-52.
Anderson, J.
1879. Anatomical and zoological researches: comprising an account of the
zoological results of the two expeditions to Western Yunnan in 1868 and 1875.
Vol. 1. Bernard Quaritch, London. For date of publication, see Corrigenda,
between p. xii and p. xiii.
1881. Catalogue of Mammalia in the Indian Museum, Calcutta, pt. 1, Primates,
Prosimiae, Chiroptera, and Insectivora. Indian Museum, Calcutta.
Annandale, N. and H. C. Robinson
1903. Fasciculi Malayenses. Supplement. Map and itinerary. Univ. Press Liverpool,
Longmans, Green & Co., London.
Anonymous
1851. Opsomming der thans bekende zoogdieren van den indischen archipel
getrokken uit de zoology of the Voyage of H. M. Ship Samarg (Lond. 1850).
Natuurk. Tijdschr. Ned. Ind., 2, pp. 443-455.
1921. Proceedings of the meeting held on June 24, 1920. Additions to the Society's
Museum. J. Bombay Nat. Hist. Soc, 27, pp. 410-412.
Antonius, O.
1951. Uber Artbastarde bei Saugetieren. Verh. ZooL-Bot. Gesellsch. Wien, 92, pp.
106-115.
Atlas van Nederland Oost-Indie
1898-1907. Department van Kolonien.'s Gravenhage.
Atlas van Tropisch Nederland
1938. Koninklijk Nederlandsch Aardrijkskundig Genootschap, Amsterdam.
143
144 FIELDIANA: ZOOLOGY, VOLUME 67
AUDEBERT, J. B.
1797-1800. Histoire naturelle des singes et des makis. Desray, Paris.
Badoux, D. M.
1959. Fossil mammals from two fissure deposits at Punung (Java)...Thesis. Univ.
Utrecht. Kemink en Zoon N. V., Utrecht.
Banks, E.
1931. A popular account of the mammals of Borneo. J. Malay. Br. R. Asiatic Soc.,
9(2), pp. 1-139.
Bartlett, A. D.
1869. Andaman monkey (Macacus andamanensis). Land and Water, 8, p. 57.
Beccari, O.
1904. Wanderings in the great forests of Borneo. Translated by E. H. Giglioli;
revised and edited by F. H. H. Guillemard. Archibald Constable & Co. Ltd.,
London.
Bennett, E. T.
1829-1830 The garden and menagerie of the Zoological Society delineated, voL 1,
Quadrupeds. Charles Tilt, London. For date of publication, see Mag. Nat. Hist,
London, 3, p. 81 (1830); 4, pp. 101, 200 (1831).
Berger, A. K.
1932. American Museum expeditions and notes. Nat. Hist., N. Y. 32, pp. 327-336
Berkson, G., R. Ross, and Samrit Jatinandana
1971. The social behavior of gibbons in relation to a conservation program. Prim.
Behav. Dev. Field Lab. Res., 2, pp. 225-255.
Bernstein, I. S.
1966. Naturally occurring primate hybrid. Science, 154, pp. 1559-1560.
1967a. Intertaxa interactions in a Malayan primate community. Folia prima t, 7,
pp. 198-207.
1967b. A field study of the pigtail monkey (Macaca nemestrina). Primates, 8, pp.
217-228,
1968. Social status of two hybrids in a wild troop of Macaca irus. Folia primat., 8,
pp. 121-131.
1972. Daily activity cycles and weather influences on a pigtail monkey group. Folia
primat., 18, pp. 390-415.
Bertrand, M.
1967. Training without reward: traditional training of pig-tailed macaques as
coconut harvesters. Science, 155, pp. 484-486.
Blainville, H. M. D. de
1839. Osteographie des primates. Sur les primates en general et sur les singes
(Pithecus) en particulier. In Blainville, H. M. D. de, Osteographie... .des
FOODEN: LIONTAIL & PIGTAIL MACAQUES 145
Mammiferes, vol. 1, pp. 1-52. J. B. Bailliere et Fils, Paris. For date of publication,
see Ann. Mag. Nat. Hist, (7) 2, p. 76 (1898).
Blakley, G. A., A. C. Morrow, and W. R. Morton
1973. Intraspecies variation in serum cholesteral levels in imported Macaca
nemestrina. Lab. Anim. Sci., 23 (1), pp. 119-121.
Blanford, W. T.
1887 [1888a]. Critical notes on the nomenclature of Indian mammals. Proc. Zool.
Soc. London, 41, pp. 620-638. For date of publication, see Proc. Zool. Soc.
London (Ser. A), 107, p. 74 (1937).
1888b. The fauna of British India, including Ceylon and Burma. Mammalia
(Introduction, Primates, Carnivora, Insectivora). Taylor and Francis, London.
Bleeker, P.
1851. Zoogdieren van Banka. Natuurk. Tijdschr Ned.-Ind., 2, pp. 527-528.
Blyth, E.
1844. Notices of various Mammalia, with descriptions of many new species. Part
1-The primates. Lin. J. Asiatic Soc. Bengal, 13 (1), pp. 463-494.
1847. Supplementary report of the Curator of the Zoological Department. Proc.
Asiatic Soc. Bengal, 16, pp. 728-737.
1851. Report on the Mammalia and more remarkable species of birds inhabiting
Ceylon. J. Asiatic Soc. Bengal, 20, pp. 153-185.
1859. Report of Curator, Zoological Department, for February to May meetings,
1859. Proc. Asiatic Soc. Bengal, 28, pp. 271-298.
1863. Catalogue of the Mammalia in the Museum of the Asiatic Society. Savielle &
Cranenburgh, Bengal Printing Co. Ltd., Calcutta.
1875. Catalogue of Mammals and birds of Burma. J. Asiatic Soc. Bengal, 44 (2),
extra number, pp. 1-167.
Boddaert, P.
1784 [1785]. Elenchus animalium, vol. 1. C. R. Hake, Rotterdam.
Bonhote, J. L.
1903. Report on the mammals, pp. 1-45. In Annandale, N. and H. C. Robinson,
Fasciculi Malayenses: Zoology, pt. 1, Univ. Press Liverpool, Longmans, Green &
Co., London.
Brown, A. E.
1896. The occurrence of Macacus leoninus (Blyth) in eastern Burmah. Proc. Acad.
Nat. Sci. Philad., 1896, p. 485.
Bufpon, G. L. L. DE
1789. Histoire naturelle generate et particultere, servant de suite a 1' histoire des
animaux quadrepedes, suppl. vol. 7. I .' Imprimerie Royale, Paris.
Bufpon, G. L. L. de and L. J. M. Daubenton
1766. Histoire naturelle, generate et particuliere, avec la description du cabinet du
roi, voL 14. L' Imprimerie Royale, Paris.
146 FIELDIANA: ZOOLOGY, VOLUME 67
Bullock, D. W., C. A. Paris, and R. W. Goy
1972. Sexual behaviour, swelling of the sex skin and plasma progesterone in the
pigtail macaque. J. Reprod. Fert, 31, pp. 225-236
Bullock, J. A.
1964. Editor's note. Malayan Nat. J., 18 (4), p. 212
BCttikofer, J.
1897. Zoological results of the Dutch Scientific Expedition to Central Borneo:
Introduction. Notes Ley den Mus. 19, pp. 1-25.
Cantor, T.
1846. Catalogue of Mammalia inhabiting the Malayan Peninsula and islands. J.
Asiatic Soc. Bengal, 15, pp. 171-203.
Chasen, F. N.
1935. On mammals from Siam. J. Siam Soc., Nat. Hist. Suppl., 10 (1), pp. 31-57.
1940a. A handlist of Malaysian mammals. Bull. Raffles Mus. Singapore, 15,
xx + 209 pp.
1940b. The mammals of the Netherlands Indian Mt. Leuser Expedition 1937 to
North Sumatra. Treubia, 17, pp. 479-840.
1941. The birds of the Netherlands Indian Mt. Leuser Expedition 1937 to North
Sumatra: 1. Introduction. Treubia, 18 (suppl.), pp. 1-3.
Chasen, F. N. and C. B. Kloss
1928. Spolia Mentawiensia. Mammals. Proc. Zool. Soc., London, 1927, pp. 808-840.
For date of publication, see Proc. Zool. Soc. London, 1928, p. 268.
1931. On a collection of mammals from the lowlands and islands of North Borneo.
Bull. Raffles Mus., 6, pp. 1-82
Chiarelli, B.
1962. Comparative morphometric analysis of primate chomosomes. II. The
chromosomes of the genera Macaca, Papio, Theropithecus and Cercocebus.
Caryologia, 15 (2), pp. 401-420.
1967. Caryological and hybridological data available for a taxonomic revision of
the Old Word primates at a suprageneric level, pp. 160-163. In Starck, D., R.
Schneider, and H. J. Kuhn, eds., Neue Ergebnisse der Primatologie Gustav
Fischer Verlag, Stuttgart.
1971. Checklist of primate hybrids. Lab. Prim. Newsl., 10 (4), pp. 1-11.
Chivers, D.
1971. The Malayan siamang. Malay. Nat. J., 24, pp. 78-86.
1973. An introduction to the socioecology of Malayan forest primates, pp. 101-146.
In Michael, R.P. and J.H. Crook, eds., Comparative ecology and behaviour of
primates, Academic Press, London and New York.
Clarke, B.
1972. Review of: Chiarelli, A.B., eds., Comparative genetics in monkeys, apes and
men. Nature, 238, p. 473
FOODEN: LIONTAIL & PIGTAIL MACAQUES 147
Coolidge, H. J., Jr.
1940. Mammal and bird collections of the Asiatic Primate Expedition: In-
troduction. Bull. Mus. Comp. Zool. Harvard, 87 (3), pp. 121-130.
Corner, E. J. H.
1941. A naturalist's companion. Malay Nat. J., 2 (1), pp. 11-14
1946. The pig-tailed monkey as a plant collector. Zoo Life, London, 1, pp. 89-92.
Crawford, M. H.
1966. Hemoglobin polymorphism in Macaca nemestrina. Science, 154, pp. 398-399.
1971. Hemoglobin polymorphism. Science, 171, p. 706.
Cuadros, A.
1971. New findings relating to the gross and microscopic morphology of the uterine
cervix in the rhesus monkey. Fertility and Sterility, 22 (2), pp. 138-143.
Cuvier, F.
1816-1817. Macaque, espece ihedite. In Dictionnaire des science naturelles.
Planches. 2e partie: Regne organise. Zoologie. Mammiferes pi. [5], fig. 2. F. G.
Levrault, Paris.
1820. Singe a queue de cochon. In Geoffroy Saint- Hillaire, E. and Cuvier, F.,
Histoire naturelle des mammiferes, vol. 1, liv. 19, pp. 1-3 Belin, Paris.
1822a. Le singe a queue de cochon et le rhesus, males adultes. In Geoffroy Saint -
Hillaire, E. and Cuvier, F., Histoire naturelle des mammiferes, vol. 3, liv. 36, pp.
1-3. Belin, Paris.
1822b. L' Ouanderou. In Geoffroy Saint-Hillaire, E. and Cuvier, F., Histoire
naturelles de mammiferes, vol. 3, liv. 36, pp. 1-2 [bis]. Belin, Paris.
1837. Ouanderou male. In Geoffroy Saint-Hillaire, E. and Cuvier, F., Histoire
naturelle des mammiferes, vol. 7, liv. 70, pp. 1-2 Belin, Paris.
Cuvier, G.
1817. Le regne animal.., vol. 1. Deterville, Paris.
Dammerman, K. W.
1922. The fauna of Krakatau, Verlaten Island and Sebesy. Treubia, 3, pp. 61-112.
1926. The fauna of Durian and the Rhio-Lingga Archipelago. Treubia, 8, pp. 281-
326.
Daniel, J. C.
1970 [1971; see p. 646]. The Nilgiri tahr, Hemitragus hylocrius Ogilby, in the High
Range, Kerala and the southern hills of the Western Ghats. J. Bombay Nat.
Hist. Soc, 67 (3), pp. 534-542.
Darlington, P. J., Jr.
1957. Zoogeography, the geographical distribution of animals. Wiley, New York.
Daudin, F. M.
1802. Indication de toutes les especes decrites par Buffon, et leur distribution dans
chacun des genres. In Lac^pede, B. G. E., Tableau des divisions, sous-divisions,
ordres et genres, [appended to] Histoire naturelle par Buffon: Quadrupedes, vol.
148 FIELDIANA: ZOOLOGY, VOLUME 67
14, pp. 143-346. [Saugrain], Paris. For publisher and date of publication, see Zool.
Med. Rijksmus. Nat. Hist. Leiden, 32, p. 215 (1953).
Davidar, E. R. C.
1971. A note on the status of the Nilgiri Tahr (Hemitragus hylocrius) on the Grass
Hills in the Anamallais. J. Bombay Nat. Hist. Soc, 68 (2), pp. 347-354.
Davis, D. D.
1958. Mammals of the Kelabit Plateau, northern Sarawak. Fieldiana: Zool., 39, pp.
119-147.
1962. Mammals of the lowland rain-forest of North Borneo. Bull. Nat. Mus.
Singapore, no. 31, pp. 1-129.
De Beaux, 0.
1917. Osservazioni morfologiche e sistematiche sul penis de Macacus arctoides Is.
Geoffr. e di cinque altre specie di macachi. Giorn. Morf. Uomo Primati, 1(1), pp.
1-12.
1923. Contributo alio studio dei macachi. Atti Soc. Lig. Sci. Lett. II, 1, pp. 22-39.
Deignan, H. G.
1959. Harry Cusheir Raven's travels in eastern Borneo. Sarawak Mus. J., (n.s.) 9
(13-14): 267-269.
Delacour, J.
1940. Liste provisoire des mammife'res de 1' Indochine franchise. Mammalia, 4 (1),
pp. 20-29.
Demers, L. M., J. MacDonald, A. T. Hertig, N. W. King, and J. J. Mackey
1972. The cervix uteri in Macaca arctoides and Macaca fascicularis—a
comparative anatomic study with special reference to Macaca arctoides as a
unique model for endometrial study. Fertility and Sterility, 33 (8), pp. 529-534.
Deninger, K.
1910. Ueber einen Affenkiefer aus den Kendengschichten von Java. Centralbl. f.
Mineralogie, 1910, pp. 1-3.
DeSimone, J., M. Linde, and R. E. Tashian
1973. Evidence for linkage of carbonic anhydrase isozyme genes in the pig-tailed
macaque, Macaca nemestrina. Nature New Biol., 242, pp. 55-56.
DeSimone, J., E. Magid, and R. E. Tashian
1973. Genetic variation in the carbonic anhydrase isozymes of macaque monkeys.
II. Inheritance of red cell carbonic anhydrase levels in different carbonic
anhydrase I genotypes of the pig-tailed macaque, Macaca nemestrina. Biochem.
Genetics, 8 (2), pp. 165-174.
Desmarest, A. G.
1817. Macaque, pp. 319-329. In Nouveau dictionnaire d'histoire naturelle..., vol. 18,
Deterville, Paris.
1820. Mammalogie ou description des especes de mammiferes, pt. 1. Agasse, Paris.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 149
Deuve, J. and M. Deuve
1963. Contribution a la connaissance des mammiferes du Laos. Liste des especes
actuellement indentifiees. Bull. Soc. R. Sci. Nat. Laos, no. 8, pp. 49-62.
DOLLMAN, G.
1932. Mammals collected by Lord Cranbrook and Captain F. Kingdon Ward in
upper Burma. Proc. Linn. Soc. London, 145, pp. 9-11.
Dorst, J.
1953. A propos de la repartition de quelques mammiferes dans l'archipel malayo-
papou. Mammalia, 17 (4), pp. 306-317.
Dunn, W.
1864. Extract from a letter to Mr. Blyth. Proc. Zool. Soc. London, 1863, pp. 370-
371. For date of publication, see Proc. Zool. Soc. London, (ser. A), 107, p. 72
(1937).
Eckstein, P. and S. Zuckerman
1956. Morphology of the reproductive tract, pp. 43-155. In Parks, A. S., ed.,
Marshall's physiology of reproduction, vol. 1, pt. 1, Longmans, Green, & Co.,
London.
Edwards, G.
1758. Gleanings of natural history. Privately published, London.
Elliot, D. G.
[1913]. A review of the primates, vol. 2. Amer. Mus. Nat. Hist., New York.
Emery, K. O.
1969. The continental shelves. Sci. Amer., 221 (3), pp. 106-122.
Erxleben, J. C. P.
1777. Systema regni animalis...Classis I Mammalia. Impensis Weygandianis,
Lipsiae.
Fea, L.
1888. Leonardo Fea nei Carin indipendenti. Boll. Soc. Geog. Italiana (Ser. 3), 1, pp.
854-868.
Fiedler, W.
1956. Ubersicht uber das System der Primates. In Hofer, H., A. H. Schultz, and D.
Starck, eds., Primatologia, vol. 1, pp. 1-266 Karger, Basel.
Fischer, J. B.
1829. Synopsis mammalium. J. G. Cotta, Stuttgart.
Flower, S. S.
1900. On the Mammalia of Siam and the Malay Peninsula. Proc. Zool. Soc.
London, 1900, pp. 306-379.
1929. A list of the vertebrated animals exhibited in the Gardens of the Zoological
Society of London, 1818-1927. I. Mammalia. Zoological Society, London.
150 FIELDIANA: ZOOLOGY. VOLUME 67
FOODEN, J.
1967. Complementary specialization of male and female reproductive structures in
the bear macaque, Macaca arctoides. Nature, 214(5091), pp. 939-941.
1969a. Taxonomy and evolution of the monkeys of Celebes (Primates: Cercopithe-
cidae). Bibl. Primat., 10, pp. 1-14&
1969b. Color-phase in gibbons. Evolution, 23 (4), pp. 627-644.
1971a. Report on primates collected in western Thailand January-April, 1967.
Fieldiana: Zool., 59(1), pp. 1-62.
1971b. Female genitalia and taxonomic relationships of Macaca assamensis.
Primates, 12(1), pp. 63-73.
1972. Male external genitalia and systematic relationships of the Japanese macaque
( Macaca fuscata Blyth, 1875). Primates, 12 (3-4), pp. 305-311.
Freedman, L.
1962. Growth of muzzle length relative to calvaria length in Papio. Growth, 26, pp.
11 7- 12a
Fry, T. B.
1928. Bombay Natural History Society's Mammal Survey of India, Burma and
Ceylon. Report No. 46. J. Bombay Nat. Hist. Soc., 32, pp. 545-547.
1929. Bombay Natural History Society's Mammal Survey of India, Burmah and
Ceylon. Report No. 46 (Supplementary). J. Bombay Nat. Hist. Soc, 33, pp. 636-
652.
Gairdner, K. G.
1914. Notes on the fauna and flora of Ratburi and Petchburi Districts. J. Nat.
Hist. Soc. Siam., 1, pp. 27-40.
Garson, M. S. and H. G. Mitchell
1970. Transform faulting in the Thai Peninsula. Nature, 228(5266), pp. 45-47.
Gentry, T. G
1872. Note on a hybrid monkey. Proc. Acad. Nat. Sri. Philad., 1872, p. 122.
GeoffroySaint-Hilaire, E.
1803. Catalogue des mammifdres du Museum National d' Histoire Naturelle. Mus.
Nat. Hist. Nat., Paris.
1812. Tableau des quadrumanes, ou des animaux composant le premier ordre de la
classe des mammiferes. Ann. Mus. Hist. Nat., Paris, 19, pp. 85-122.
GeoffroySaint-Hilaire, I.
1826. Macaque, pp. 584-590. In Bory de Saint-Vincent, ed., Dictionnaire classique d'
histoire naturelle, vol. 9, Rey et Gravier and Baudoin Freres, Paris.
Gloger, C. W. L
1841. Gemeinnutziges Hand- und Hilfsbuch der Naturgeschichte, vol. 1. Aug. Schulz
& Comp., Breslau.
Goldfoot, D. A.
1971. Hormonal and social determinants of sexual behavior in the pigtail monkey
FOODEN: LIONTAIL & PIGTAIL MACAQUES 151
(Macaca nemestrina) pp. 325-342. In Stoelinga, G. B. A. and J. J. van der Werff
ten Bosch, eds., Normal and abnormal development of brain and behavior,
Leiden University Press, Leiden.
Goldfuss, G. A.
1820. Handbuch der Zoologie. Johann Leonhard Schrag, Numberg.
Goodman, M.
1965. Reply to Macaque "semispecies." Science, 148, p. 255.
1967. Effects of evolution on primate macromolecules. Primates, 8, pp. 1-22.
1971. Molecular evolution at the population level in higher primates, pp. 81-87. In
Proc. 3 Int. Congr. Primat., Zurich 1970, vol. 2, Karger, Basel.
Goodman, M., A. Kulkarni, M. D. Poulik, and E. Reklys
1965. Species and geographic differences in the transferrin polymorphism of
macaques. Science, 147, pp. 884-886.
GOUSTARD, M.
1968. La sequence d'activites liees a la copulation chez le macaque crabier Macaca
irus. Ann. Sci. Nat. Zool., (12) 10, pp. 463-474.
Gray, A. P.
1971. Mammalian hybrids, 2nd ed. Commonwealth Agricultural Bureaux, Famham
Royal, Slough, England.
Gray, J. E.
1843. List of the specimens of Mammalia in the collection of the British Museum.
British Museum, London.
Griffith, E.
1821. Class Quadrimembria, Order Quadrumana. General and particular descrip-
tions of monkeys and lemurs. Baldwin, Craddock & Joy and Rodwell & Martin,
London.
Grimm, R. J.
1967. Catalogue of sounds of the pigtailed macaque (Macaca nemestrina). J. Zool.
London, 152, pp. 361-373.
Griswold, J. A., Jr.
1939. Up Mount Kinabalu. Sci. Monthly, 48, pp. 401-414, 504-518.
Groves, C. P.
1967. Geographic variation in the hoolock or white-browed gibbon (Hylobates
hoolock Harlan, 1834). Folia primat., 7, pp. 276-283.
Gyldenstolpe, N.
1913. Birds collected by the Swedish Zoological Expedition to Siam 1911-1912. K.
Sven. Vet. Akad. Handl. (N. F.), 50 (8), pp. 1-76
1914. Mammals collected, or observed by the Swedish Zoological Expedition to
Siam 1911-1912. Ark. Zool., 8 (23), pp. 1-36.
152 FIELDIANA: ZOOLOGY, VOLUME 67
1916. Zoological results of the Swedish Zoological Expeditions to Siam 1911-1912 &
1914-1915. IV. Birds II. K. Sven. Vet. Akad Handl. (N.F.) 56 (2), pp. 1-160, 5 pi.
1916 [1917]. Zoological results of the Swedish Zoological Expeditions to Siam 1911-
1912 & 1914-1915. V. Mammals II. K. Sven. Vet. Akad. Handl.. (N.F.) 57 (2), pp.
1-60.
1919. A list of the mammals at present known to inhabit Siam. J. Nat. Hist. Soc.
Siam, S (3), pp. 127-175.
Hafez, E. S. E. and S. Jaszczak
1972. Comparative anatomy and histology of the cervix uteri in non-human
primates. Primates, 13 (3), pp. 297-316.
Haffer, J.
1969. Speciation in Amazonian forest birds. Science, 165 (3889), pp. 131-137.
Hagen, B.
1890. Die Pflanzen-und Thierwelt von Deli auf der Ostkuste Sumatra's. Tijdschr.
K. Ned. Aardr. Genootsch., (Ser. 2) 7, pp. 1-240.
Hamilton, F.
1870. Extract from a letter to Dr. E. Hamilton concerning the Andaman monkey.
Proc. ZooL Soc. London, 1870, p. 220.
Harrison, J. L.
1961. The natural food of some Malayan mammals. Bull. Nat. Mus. Singapore, no.
30, pp. 5-18.
1969. The abundance and population density of mammals in Malayan lowland
forests. Malay Nat. J., 22, pp. 174-178.
Harrison, J. L. and J. R. Hendrickson
1963. The fauna of the islands of the Straits of Malacca, pp. 543-555. In Gressitt, J.
L., ed., Pacific Basin biogeography. A symposium, Tenth Pacific Science
Congress, Honolulu, Hawaii, 1961, Bishop Museum Press, Honolulu.
Harrisson, T.
1958. The caves of Niah: a history of prehistory. Sarawak Mus. J., 8, pp. 549-595.
Hartert, E.
1902. Aus den Wanderjahren eines Naturforschere. II. Abschitt. Reise nach
Sumatra, Malakka und Indien. Kapitel 2-5. Novitat. zool., 9, pp. 193-272.
Heck, L.
1916. Herren-oder Hochtiere (Primates), pp. 380-699. In Strassen, O. zur ed.,
Brehms Tierleben, Saugetiere, 4th ed., vol. 4, Bibliographisches Institut,
Leipzig /Wien.
Helfer, J. W.
1838. Notes on the animal productions of the Tenaaserim provinces. J. Asiatic Soc.
Bengal, 7 (2), pp. 855-86a
Hill, C. A.
1971. Zoo's help for a rare monkey. Oryx, 11 (1), pp. 35-38.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 153
Hill, W. C. O.
1937. The primates of Travancore. Proc. Zool. Soc. London (Ser. B), 107, pp. 205-
216
1958. External genitalia, pp. 630-704. In Hofer, H., A. H. Schultz, and D. Starck,
eds., Primatologia, vol. 3, pt. 1, Karger, Basel/New York.
Ho, R.
1960. The evolution of the Indo-Malayan region, pp. 7-20. In Purchon, R. D., ed.,
Proc. Cent. Bicent. Congr. Biol., Singapore, December 2-9, 1958, Univ. Malaya
Press, Singapore.
HOOGERWERF, A.
1953. Some notes about the Nature Reserve Pulau Panaitan (Prinseneiland) in
Strait Sunda, with special reference to the avifauna. Treubia, 21 (3), pp. 481-505.
HOOIJER, D. A.
1946. Prehistoric and fossil rhinoceroses from the Malay Archipelago and India.
Zool. Med. Rijksmus. Nat. Hist. Leiden, 26, pp. 1-138.
1948. Prehistoric teeth of man and of the orang-utan from central Sumatra, with
notes on the fossil orang-utan from Java and southern China. Zool. Med.
Rijksmus. Nat. Hist. Leiden, 29, pp. 175-301.
1962a. Quaternary langurs and macaques from the Malay Archipelago. Zool. Verh.
Rijksmus. Nat. Hist. Leiden, no. 55, pp. 1-64.
1962b. Prehistoric bone: the gibbons and monkeys of Niah Great Cave. Sarawak
Mus. J. (n. s.), 11, pp. 428-449.
1964. New records of mammals from the Middle Pleistocene of Sangiran, central
Java. Zool. Med. Rijksmus. Nat. Hist. Leiden, 40, (10), pp. 73-88.
Horsfield, T.
1851. A catalogue of the Mammalia in the museum of the Hon. East-India
Company. East India Co., London.
Hurme, V. O.
1960. Estimation of monkey age by dental formula. Ann. N. Y. Acad. Sci., 85, pp.
795-799.
Hutton, A. F.
1949. Notes on the snakes and mammals of the High Wavy Mountains, Madura
District, South India. Part II-Mammals. J. Bombay Nat. Hist. Soc., 48, pp. 681-
694.
The Imperial Gazetteer of India,
1907-1909. new ed., 26 vols. Clarendon Press, Oxford.
International Zoo Yearbook
1963. Species of animals bred in zoos and aquaria during 1962. Int. Zoo Yearb., 4,
pp. 220-261.
1966. Species of animals bred in zoos and aquaria during 1964. Int. Zoo Yearb., 6,
pp. 385-431.
154 FIELDIANA: ZOOLOGY, VOLUME 67
1967. Species of wild animals bred in captivity during 1965. Int. Zoo Yearb., 7, pp.
300-356.
1968. Species of wild animals bred in captivity during 1966. Int. Zoo. Yearb., 8, pp.
288- 34a
1969. Species of wild animals bred in captivity during 1967. Int. Zoo Yearb., 9, pp.
215-273
1970. Species of wild animals bred in captivity during 1968. Int. Zoo Yearb. 10, pp.
249-310
1971. Species of wild animals bred in captivity during 1969. Int. Zoo. Yearb., 11, pp.
259-320.
1972. Species of wild animals bred in captivity during 1970. Int. Zoo. Yearb., 12, pp.
311-375.
ISHIMOTO, G.
1971. Electrophoretic patterns of red cell NADH diaphorase in macaque monkeys.
J. Anthropol. Soc. Nippon, 79 (4), pp. 382-387.
1972a. Blood protein variations in Asian macaques. I. Serum proteins and
hemoglobin. J. Anthropol. Soc. Nippon, 80 (3), pp. 250-274.
1972b. Blood protein variation in Asian macaques. II. Red cell enzymes. J.
Anthropol. Soc. Nippon, 80 (4), pp. 337-350.
1973. Blood protein variations in Asian macaques. III. Characteristics of macaque
blood protein polymorphism. J. AnthropoL Soc. Nippon, 81 (1), pp. 1-13.
Ishimoto, G. and M. Kuwata
1972. Phosphohexose isomerase polymorphism in macaques. Jap. J. Hum. Genetics,
16(3), pp. 154-165.
Ishimoto, G., T. Tanka, N. Hideo, and W. Prychodko
1970. Hemoglobin variation in macaques. Primates, 11, pp. 229-241.
Jainudeen, M. R., E. S. E. Hafez, and L. Kothari
1972. Histomorphology of the cervix uteri of toque (M. sinica) and bonnet
macaques (M. radiata). Folia primatol., 18, pp. 469-476.
Jentink, F. A.
1883. A list of species of mammals from West-Sumatra and North-Celebes, with
descriptions of undescribed or rare species. Notes Leyden Mus., 5, pp. 170-181.
1887. Catalogue ost^ologique des mammiferes, Museum d' histoire naturelle des
Pays- Bays, vol. 9. Brill, Leiden.
1897. Zoological results of the Dutch Scientific Expedition to central Borneo. The
mammals. Notes Leyden Mus., 19, pp. 26-66.
Jerdon, T. C.
1867. The mammals of India. Privately published, Roorkee.
Johnson, D. S.
1964. A question of nomenclature. Malay. Nat. J., 18 (1), pp. 68-69.
Jolly, C. J.
1971. Protein structure and primate systematica. Medical primat 1970. Proc. 2nd.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 155
Conf. exp. Med. Surg. Primates, N. Y., 1969, pp. 190-212. Karger, Basel.
JONGSMA, D.
1970. Eustatic sea level changes in the Arafura Sea. Nature, 228, pp. 150-151.
Kanagawa, H. and E. S. E. Hafez
1973. Copulatory behavior in relation to anatomical characteristics of three
macaques. Amer. J. Phys. Anthropol., 38, pp. 233-240.
Kanagawa, H., E. S. E. Hafez, M. M. Nawar, and S. Jaszczak
1972. Patterns of sexual behavior and anatomy of copulatory organs in macaques.
Z. Tierpsychol., 31, pp. 449-460.
Kao Yueh-Ting, Lu Chang-Kwun, Chang Chieh and Wang SPng
1962. Mammals of the Hsi-Shuan-Pan-Na area in southern Yunnan. Acta Zool.
Sinica, 14 (2), pp. 180-196.
Karr, J. R.
1973. Ecological and behavioural notes on the liontailed macaque (Macaca silenus)
in South India. J. Bombay Nat. Hist. Soc, 70, pp. 191-193.
Karrer, R.
1970. The use of the tail by an old world monkey. Primates, 11, pp. 171-175.
Kawabe, M.
1970. A preliminary study of the wild siamang gibbon (Hylobates syndactylus) at
Fraser's Hill, Malaysia. Primates, 11, pp. 285-291.
Kawabe, M. and T. Mano
1972. Ecology and behavior of the wild proboscis monkey, Xasalis lariatus
(Wurmb), in Sabah, Malaysia. Primates, 13 (2), pp. 213-228.
Kelsall, H. J.
1894. A journey on the Sembrong River from Kuala Indau to Batu Pahat. Personal
account of the journey. J. Straits Br. R. Asiatic Soc, 26, pp. 1-19.
Keng, H.
1970. Size and affinities of the flora of the Malay Peninsula. J. Trop. Geog., 31, pp.
43-56.
Kerr, R.
1792. The animal kingdon... Class I. Mammalia.... A. Strahan and T. Cadell and W.
Creech, Edinburgh.
Khajuria, H.
1954 [1955]. Catalogue of mammals in the Indian Museum (Zool. Surv.) II.
Primates: Cercopithecidae. Rec. Indian Mus., 52, pp. 101-127.
Kinnear, N. B.
1934. On the birds of the Adung Valley, North-East Burma. J. Bombav Nat. Hist.
Soc, 37, pp. 347-368.
156 FIELDIANA: ZOOLOGY, VOLUME 67
Kinzey, W. G.
1971. Male reproductive system and spermatogenesis, pp. 85-114. In Hafez, E. S. E.,
ed., Comparative reproduction of nonhuman primates, Charles C. Thomas,
Springfield, Illinois.
Kloss, C. B.
1903. In the Andamans and Nicobars. John Murray, London.
1908. A provisional list of the mammals of the peninsular region. J. Fed. Malay
States Mus., 2, pp. 147-150.
1911a. On mammals and birds from the lowlands of Pahang. J. Fed. Malay States
Mus., 4, pp. 144-166
1911b. On a collection of mammals and other vertebrates from the Trengganu
Archipelago. J. Fed. Malay States Mus., 4, pp. 175-212.
1916a. On a collection of mammals from the coast and islands of South-East Siam.
Proc. Zool. Soc. London, 1916, pp. 27-75.
1916b. On a collection of mammals from Siam. J. Nat. Hist. Soc. Siam., 2(1), pp.
1-32.
1919a. On birds from South Annam and Cochin China. Part 1. Phasianidae-
Campophagidae. Ibis, (11) 1, pp. 392-453.
1919b. On mammals collected in Siam. J. Nat. Hist. Soc. Siam, 3 (4), pp. 233-407.
1920. On a collection of plants from peninsular Siam: some account of the journey
on which the plants were collected. J. Fed. Malay States Mus., 10, pp. 66-80.
1928. Spolia Mentawiensia.— Mammals. Introduction. Proc. Zool. Soc. London,
1927, pp. 797-807. For date of publication, see Proc. Zool. Soc. London, 1928, p.
268.
Knottnerus-Meyer, T.
1904. Nochmals "Uber Saugetierbastarde." Zool. Gart., 45, pp. 60-63.
Knox, R.
1681 [1911 ed.]. An historical relation of Ceylon.... James Mac Lehose and Sons,
Glasgow.
KOBAYASHI, T.
1960. Notes on the geologic history of Thailand and adjacent territories. Jap. J.
Geol. Geog., 31, pp. 129-148.
Koeppe, C. E. and G. C. De Long
1958. Weather and climate. McGraw Hill, New York.
KOHLBRUGGE, J. H. F.
1896. Bijdragen tot de natuurlijke geschiedenis van menschen en dieren. Ill
Zoogdieren van zuid-oost Borneo. Natuurk. Tijdschr. Ned.-Ind., 55, pp. 176-200.
Krishnan, M.
1971 [1972]. An ecological survey of the larger mammals of peninsular India. J.
Bombay Nat. Hist. Soc, 68 (3), pp. 503-555.
Kf'CHI.ER, A. W.
1964. Natural vegetation, pp. 16, 17. In Espenshade, E. B., Jr.. ed., Goode's world
FOODEN: LIONTAIL & PIGTAIL MACAQUES 157
atlas, 12th ed., rev., Rand McNally & Co., Chicago.
Kuehn, R. E., G. D. Jensen, and R. K. Morrill
1965. Breeding in Macaca nemestrina: a program of birth engineering. Folia
pnmat.. 3, pp. 251-262.
Kurt, F.
1973. Der Gunung Leuser Survey 1970. Z. Kolner Zoo, 16 (2), pp. 59-74.
KURUP, G. U.
1963. On the cranial characters of Macaca silenus (Linn.) (Primates: Cercopithe-
cidae). J. Bombay Nat. Hist. Soc, 60 (1), pp. 246-249.
1966. Mammals of Assam and adjoining areas. 1. An analytical study. Proc. Zool.
Soc. Calcutta, 19, pp. 1-21.
Lacepede, B. G. E.
1799. Tableau des divisions, sous-divisions, ordres et genres des mammiferes, pp. 1-
18. In Discours d' ouverture et de cloture du cours d' histoire naturelle donne
dans le Museum national d' Histoire naturelle, Tan VII de la Republique, et
Tableaux m£thodique des mammiferes et des oiseaux, [pt. 2], Plassan, Paris.
La Salle, M.
1969. Erythrocyte isoantigens of the genus Macaca. Proc. 2 Int. Congr. Primat.,
Atlanta, GA. 1968, vol. 3, pp. 120-128. Karger, Basel.
Latreille, P. A.
1802. Histoire naturelle generate et particuliere, des crustaces et des insectes (C. S.
Sonnini, ed.), vol. 3. F. Dufart, Paris.
1804. Tableau m^thodique des singes, pp. 275-298. In Sonnini, C. S. ed., Histoire
naturelle generate et particuliere, par Leclerc de Buffon, Dufart, Paris.
Legendre, S. J.
1932. Adventures on hunting trails of Indo-China. Nat. Hist., New York, 32, pp.
481-496.
Lesson, R. P.
1834 [1830]. Complement des oeuvres de Buffon..., vol. 4. Pourrat Freres and Roret,
Paris. For date of publication, see I. Geoffroy, Cat. Meth. Mamm. Mus. Hist.
Nat. Paris, p. vji (1851).
1839. Mastologie methodique. Rev. Zool., Paris 2, pp. 68-70.
LlM Boo LlAT
1969. Distribution of the primates of West Malaysia. Proc. 2 Int. Congr. Primat,
Atlanta, Ga. 1968, vol. 2, pp. 121-130. Karger, Basel.
Lindsay, H. M.
1926. Bombay Natural History Society's Mammal Survey of India, Burma and
Ceylon. Report Nos. 43-45. J. Bombay Nat. Hist. Soc., 31, pp. 591-614.
Link, H. F.
1795. Beytrage zur Naturgeschichte, vol. 2 K. C. Stiller, Rostock & Leipzig.
158 FIELDIANA: ZOOLOGY, VOLUME 67
I.INKK. K.
1971. Javaner-Wanderu-Bastard im Zoo Rostock. Zool. Gart. (N. F., Leipzig) 40
(3), pp. 163-166.
Linnaeus, C.
1758. Systema naturae, 10th ed., vol. 1. Impensis Direct. Laurentii Salvii, Holm.
1766. Systema naturae, 12th ed., vol. 1. Impensis Direct. Laurentii Salvii, Holm.
LUMER, H. and A. H. ScHULTZ
1941. Relative growth of the limb segments and tail in macaques. Hum. Biol., 13,
pp. 283-305.
Lyon, M. W., Jr.
1906. Mammals of Banka, Mendanau, and Billiton Islands between Sumatra and
Borneo. Proc. U. S. Nat. Mus., 31, pp. 575-612.
1907a. Mammals of Batam Island, Rhio Archipelago. Proc. U. S. Nat. Mus., 31, pp.
653-657.
1907b. Mammals collected in western Borneo by Dr. W. L. Abbott. Proc. U. S. Nat.
Mus., 33, pp. 547-572.
1908. Mammals collected in eastern Sumatra by Dr. W. L. Abbott during 1903,
1906, and 1907, with descriptions of new species and subspecies. Proc. U. S. Nat.
Mus., 34, pp. 619-679.
1911. Mammals collected by Dr. W. L. Abbott on Boreno and some of the small
adjacent islands. Proc. U. S. Nat. Mus., 40, pp. 53-146.
MacKinnon, J.
1971. The orang-utan in Sabah today. A study of a wild population in the Ulu
Segama Reserve. Oryx, 11 (2-3), pp. 141-191.
Magid, E., J. DeSimone, and R. E. Tashian
1973. Genetic variation in the carbonic anhydrase isozymes of macaque monkeys. I.
The radioimmunosorbent assay. Biochem. Genetics, 8 (2), pp. 157-164.
Mayr, E.
1963. Animal species and evolution. Harvard Univ. Press, Cambridge.
McCann, C.
1933a. Notes on the colouration and habits of the white-browed gibbon or hoolock
(Hylobates hoolock Harl.). J. Bombay Nat. Hist. Soc, 36, pp. 395-405.
1933b. Notes on some Indian macaques. J. Bombay Nat. Hist. Soc, 36, pp. 796-810.
McClure, H. E.
1964. [1965]. Some observations of primates in climax dipt[er]ocarp forest near
Kuala Lumpur, Malaya. Primates, 5, pp. 39-58.
Mc Neely, J. A. and E. W. Cronin
1972. Rhinos in Thailand. Oryx, 11 (6), pp. 457-460.
Medway, Lord
1964. Post- Pleistocene changes in the mammalian fauna of Borneo. Archaeological
FOODEN: LIONTAIL & PIGTAIL MACAQUES 159
evidence from the Niah caves. Stud. Speleol., 1(1), pp. 33-37.
1965. Mammals of Borneo: field keys and an annotated checklist. J. Malay. Br. R.
Asiatic Soc, 36 (3), pp. 1-193.
1970. The monkeys of Sundaland: ecology and systematics of the cercopithecids of
a humid equatorial environment, pp. 513-553 In Napier, J. R. and P. H. Napier,
eds., Old world monkeys: evolution, systematics, and behavior, Academic Press,
New York.
1972. The Gunong Benom Expedition 1967. G The distribution and altitudinal
zonation of birds and mammals on Gunong Benom. Bull. Brit. Mus. Nat. Hist,
Zool., 23 (5), pp. 103-154.
Medway, Lord and D. R. Wells
1971. Diversity and density of birds and mammals at Kuala Lompat, Pahang.
Malay Nat. J., 24, pp. 238-247.
Michael, R. P., M. Wilson, and T. M. Plant
1973. Sexual behaviour of male primates and the role of testosterone, pp. 235-313.
In Michael, R. P. and J. H. Crook, eds., Comparative ecology and behaviour of
primates.
Miller, G. S., Jr.
1900. Mammals collected by Dr. W. L. Abbott on the islands in the North China
Sea. Proc. Wash. Acad. Sci., 2, pp. 203-246.
1903a. Mammals collected by Dr. W. L. Abbott on the coast and islands of
northwest Sumatra. Proc. U.S. Nat., Mus., 26, pp. 437-484.
1903b. Seventy new Malayan mammals. Smiths. Misc. Coll., 45, pp. 1-73
1906. The monkeys of the Macaca nemestrina group. Proc. U.S. Nat. Mus., 29, pp.
555-563.
1933. The groups and names of macaques pp. 1-9. In Hartman, C. G. and W. L.
Straus, Jr., eds., The anatomy of the rhesus monkey (Macaca mulatto), Williams
& Wilkins, Baltimore.
Montagu, M. F. A.
1950. A hybrid gibbon. J. Mammal., 31, pp. 150-153.
Moore, J. C. and G. H. H. Tate
1965. A study of the diumal squirrels, Sciurinae, of the Indian and Indochineae
Subregions. Fieldiana: Zool., 48, pp. 1-351.
Moore-Jankowski, J. and A. S. Wiener
1971. Blood groups of primates: their contribution to taxonomy and phylogenetics.
Med. Primat. 1970. Proc. 2 Conf. Exp. Med. Surg. Primates, New York 1969, pp.
232-244. Karger, Basel.
Morice, A.
1875. Coup d'oeil sur la faune de la Cochinchine Francaise. H. George, Lyon.
Morris, R. C.
1936. To Malaya for a rhinoceros. J. Bombay Nat. Hist. Soc, 38, pp. 438-446.
160 FIELDIANA: ZOOLOGY, VOLUME 67
MULLER, S.
1839-1640. Over de zoogdieren van den Indischen Archipel. In: Temminck, C. J.
(ed.), Verhandelingen over de natuurilijke geschiedenis der Nederlandsche
overzeesche bezittingen, vol. 1, Zoologie, pp. l-[63]. Luchtmans and van der
Hoik, Leiden. For date of publication, see Zool. Med., Leiden 34, p. 22 (1955).
Myers, B. J. and R. E. Kuntz
1969. Nematode parasites of mammals (Dermoptera, Primates, Pholidota,
Rodentia, Carnivora, and Artiodactyla) from North Borneo (Malaysia).
Canadian J. Zool., 47 (3), pp. 419-421.
Nadler, R. D. and L. A. Rosenblum
1973a. Sexual behavior of male pigtail macaques in the laboratory. Brain, Behav.
Evol., 7, pp. 18-33.
1973b. Sexual behavior during successive ejaculations in bonnet and pigtail
macaques. Amer. J. Phys. Anthropol., 38, pp. 217-220.
Nakajima, H., T. Tanka, H. Nigi, and W. Prychodko
1970. Human-type ABO, MN, and Lewis blood groups, and Gm and Inv factors in
several species of macaques. Primates, 11, pp. 243-253.
Napier, J. R. and P. H. Napier
1967. A handbook of living primates. Academic Press, London /New York.
Nesturkh, M.
1959. The origin of man. Foreign Lang. Publ. House, Moscow.
New York Zoological Society
1933. Thirty-seventh annual report. New York.
Nute, P. E. and H. A. Pataryas
1974a. Multiple fetal and adult hemoglobins in Malaysian Macaca nemestrina:
consequences of a-chain variation. Amer. J. Phys. Anthropol., 40, pp. 17-26.
1974b. Amino acid compositions of the tryptic peptides comprising the )8-
hemoglobin chain of Macaca nemestrina. Amer. J. Phys. Anthropol, 40, pp. 75-
82.
Nute, P. E. and G. Stamatoyannopoulos
1971a. Evidence for duplication of the haemoglobin y-chain locus in Macaca
nemestrina. Nature New Biol., 229, pp. 145-147.
1971b. The ontogensis of hemoglobins in Macaca nemestrina. Blood, 38 (1), pp.
108-115.
[Ogilby, W.]
1838. The menageries. The natural history of monkeys, opossums, and lemurs, vol.
1. Knight, London. For name of author, see Proc. Zool. Soc. London, 1843, p. 11.
Omoto, K., S. Harada, T. Tanaka, H. Nigi, and W. Prychodko
1970. Distribution of the electrophoretic variants of serum alphai - antitrypsin in
six species of macaques. Primates, 11, pp. 215- 228
FOODEN: LIONTAIL & PIGTAIL MACAQUES 161
Osgood, W. H.
1932. Mammals of the Kelley-Roosevelts and Delacour Asiatic Expeditions. Field
Mus. Nat. Hist., Publ. 312, Zool. Ser., 18 (10), pp. 193-339.
OUDEMANS, A. C.
1890. Uber zwei seltene und eine neue Art Affen des zoologisches Gartens in Haag,
Holland. Zool. Gart., 31, pp. 266-269.
Pennant, T.
1771. Synopsis of quadrupeds. J. Monk, Chester.
1781. History of quadrupeds, vol. 1. B. White, London.
POCOCK, R. I.
1906. Notes upon menstruation, gestation, and parturition of some monkeys that
have lived in the Society's Gardens. Proc. Zool. Soc. London, 1906, pp. 558-570.
1921. The systematic value of the glans penis in macaque monkeys. Ann. Mag. Nat.
Hist., (Ser. 9) 7, pp. 224-229.
1926. The external characters of the catarrhine monkeys and apes. Proc. Zool. Soc.
London, 1925, pp. 1479-1579. For date of publication, see Proc. Zool. Soc.
London, (Ser. A) 107, p. 77 (1937).
1931. The pig-tailed macaques (Macaca nemestrina). J. Bombay Nat. Hist. Soc,
35, pp. 297-311.
1939. The fauna of British India, including Ceylon and Burma. Mammalia. — Vol.
1. Primates and Carnivora (in part). Taylor and Francis Ltd., London.
Pohl, L.
1928. Zur morphologie der mannlichen Kopulationsorgane der Saugetiere; in-
sbesondere der Versuch einer veigleichend-anatomischen Studie uber den Penis
der Primaten, einschlisslich des Menschen. Z. anat. Entwickl.-Gesch., 86, pp. 71-
119.
Poirier, F. E.
1970. The Nilgiri langur (Presbytia johnii) of South India, pp. 251-383. In
Rosenblum, L. A., ed., Primate behavior: developments in field and laboratory
research, Academic Press, New York.
Prychodko, W., M. Goodman, E. Poulik, T. Miki, and T. Tanaka
1969. Geographic variations of transferrin allelic frequencies in continental and
insular macaque populations. Proc. 2 Int. Congr. Primat, Atlanta, GA, vol. 2, pp.
103-108. Karger, Basel.
Prychodko, W., M. Goodman, M. Singal, M. L. Weiss, G. Ishimoto, and T.
Tanaka
1971. Starch-gel electrophoretic variants of erythrocyte 6-phosphogluconate
dehydrogenase in Asian macaques. Primates, 12 (2), pp. 175-182.
Raffles, T. S.
1821. Descriptive catalogue of a zoological collection, made on account of the
Honourable East India Company, in the island of Sumatra and its vicin-
ity-Trans. Linn. Soc. London, 13, pp. 239-274.
162 FIELDIANA: ZOOLOGY, VOLUME 67
Ray, J.
1693. Synopsis methodica animalium quadrupediutn et serpentini generis. S. Smith
& B. Walford, London.
Reich enbach, L.
1862. Die vollstandigste Naturgeschichte der Affen. Expedition der vollstandigste
Naturgeschichte, Dresden /Leipsig.
Ridley, H. N.
1895. The mammals of the Malay Peninsula. Nat. Sci., London, 6, pp. 23-29.
1906. The menagerie at the Botanic Gardens. J. Straits Br. R. Asiatic Soc, 46, pp.
133-194.
Robinson, H. C.
1919. Notes on the vertebrate fauna of the Pahang-Johore Archipelago. J. Fed.
Malay States Mus., 7, pp. 325-329.
Robinson, H. C. and C. B. Kloss
1910. On birds from the northern portion of the Malay Peninsula, including the
islands of Langkawi and Tarutau; with notes on other rare Malayan species from
the southern districts. Ibis, 4, pp. 659-675.
1914a. Some remarks on Dr. D. G. Elliot's "Review of the Primates." Ann. Mag.
Nat. Hist., (8) 13, pp. 389-399.
1914b. The zoology of Koh Samui and Koh Pennan: Mammals. J. Fed. Malay
States Mus., 5, pp. 130-139.
1919. On a collection of birds from the province of Puket, peninsular Siam. J. Nat.
Hist. Soc. Siam, 3 (2), pp. 87-119.
1921. The birds of South-west and peninsular Siam. J. Nat. Hist. Soc. Siam, 5(1),
pp. 1-88.
Rodman, P. S.
1973. Synecology of Bornean primates. I. A test for interspecific interactions in
spatial distribution of five species. Amer. J. Phys. Anthropol., 38, pp. 655-660.
Romer, A. S.
1966. Vertebrate paleontology, 3rd. ed. Univ. Chicago Press, Chicago.
Ryley, K. V.
1913. Bombay Natural History Society's Mammal Survey of India. Report Nos. 10,
11. J. Bombay Nat Hist. Soc., 22, pp. 464-513
1914. Bombay Natural History Society's Mammal Survey of India, Burma and
Ceylon. Report Nos. 12, 13, 14. J. Bombay Nat. Hist. Soc, 22, pp. 684-725.
Sanborn, C. C.
1952. The mammals of the Rush Watkins Zoological Expedition to Siam. J. Siam
Soc, 15 (1), pp. l-2(i.
Schinz, H. R.
1821. Das Thierreich..., voL 1. J. G. Cotta, Stuttgart and Tubingen.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 163
SCHLEGEL, H.
1876. Les singes: Simiae, vol. 7, monograph 40. Mus. Hist. Nat. Pays-Bas, Leiden.
SCHMAGER, J.
1972. Cytotaxonomy and geographical distribution of the Papinae. J. Hum. Evol.,
1, pp. 477-485.
Schneider, G.
1905. Ergebnisse zoologischer Forschungsreisen in Sumatra. Erster Teil. Saugetiere
(Mammalia). Zool. Jahrb., 23, pp. 1-172.
SCHREBER, J. C. D.
1774. Die Saugthiere in Abbildungen nach der Natur mit Beschreibungen, pt. 1.
Wolfgang Walther, Erlangen. For date of publication, see Proc. Zool. Soc.,
London, 1891, pp. 587-592.
Schultz, A. H.
1933. Growth and development, pp. 10-27. In Hartman, C. G. and W. L. Straus, Jr.,
eds., The anatomy of the rhesus monkey (Macaca mulatto), Williams & Wilkins,
Baltimore.
1938. The relative length of the regions of the spinal column in Old World
primates. Amer. J. Phys. Anthropol., 24, pp. 1-22
Schwarz, E.
1934. Uber einen Hybriden von Schweinsaffe (Macaca nemestrina Linnaeus) und
Mohrenmakak (Macaca nigra maura F. Cuvier). Zool. Gart. (n. s.) 7, pp. 44-46
Sclater, P. L.
[1870]. Notices of recent additions to the Menagerie. Proc. Zool. Soc. London,
1869, pp. 467-469. For date of publication, see Proc. Zool. Soc. London, (Ser. A)
107, p. 72(1937).
[1871]. Reports on additions to the Society's Menagerie in June, July, August, and
September 1870, and description of Buceros subcylindricus. Proc. Zool. Soc.
London, 1870, pp. 663-671. For date of publication, see Proc. Zool. Soc. London,
(Ser. A) 107, p. 72(1937).
[1881]. Report of additions made to the Society's Menagerie during the months of
June, July, August, September and October 1880. Proc. Zool. Soc. London, 1880,
pp. 537-539. For date of publication, see Proc. Zool. Soc. London, (Ser. A) 107, p.
73 (1937).
1883. List of the vertebrated animals now or lately living in the Gardens of the
Zoological Society of London, 8th ed. Zoological Society, London.
1893. Exhibition of, and remarks upon, a skin of a variety of the pig-tailed monkey,
Macacus nemestrinus. Proc. Zool. Soc. London, 1893, p. 325.
1898. Remarks on the principal animals observed during recent visits to the
Marseilles Garden, the menagerie of the Bey of Tunis at Marsa, and the Jardin
d'Acclimatation, Paris. Proc. Zool. Soc. London, 1898, p. 280.
Sclater, W. L. and Sclater, P. L.
1899. The geography of mammals. Kegan Paul, Trench, Triibner & Co., London.
164 FIELDIANA: ZOOLOGY, VOLUME 67
SCRIVENOR, J. B.
1911. A sketch of the geological structure of the Malay Peninsula. J. Straits Br. R.
Asiatic Soc, 59. pp. 1-1.1
1949. Geological and geographical evidence for changes in sealevel during ancient
Malayan history and late prehistory. J. Malay. Br. R. Asiatic Soc., 22 (1), pp.
107-115.
Shaw, G.
1792. Museum Leverianum..., vol. 1. James Parkinson, London.
1800. General zoology or systematic natural history, vol. 1, pt. 1 (Mammalia). G.
Kearsley, London.
Simons, R. C. and M. H. Crawford
1969. Determination of paternity in group-born pigtailed monkeys (Macaca
nemestrina). Proc. 2 Int. Congr. Prim., Atlanta, Ga. 1968, vol. 1, pp. 254-260.
Karger, Basel.
Socie'te' Royale de Zoologie d'Anvers.
1968. Inventaire des collections zoologiques, 1968. Zoo, Anvers, 34 (1 -2, suppL), pp.
1-31.
Sody, H. J. V.
1936. Seventeen new generic, specific, and subspecific names for Dutch East Indian
mammals. Natuurk. Tijdschr. Ned. Ind., 96, pp. 42-55.
1937. On the mammals of Banka. Temminckia, 2, pp. 220-250.
Southwick, C. H. and F. C. Cadigan, Jr.
1972. Population studies of Malaysian primates. Primates, 13 (1), pp. 1-18.
Spence, R. A.
1921. Proceedings of the meeting held on the 2nd March 1921. J. Bombay Nat.
Hist. Soc, 27, pp. 968-973.
Spiller, G. A. and D. F. Rahlmann
1970. Physiological effect and acceptance of various natural foodstuffs in the pig-
tailed monkey (Macaca nemestrina). Lab. Anim. Care, 20 (3), pp. 494-498.
Steenis, C. G. G. J. Van
1950. The delimitation of Malaysia and its main plant geographical divisions, pp.
lxx-lxxv. In Steenis, C. G. G. J. van, ed., Flora Malesiana, Ser. 1, Vol. 1,
Noordhoff-Kolff N. V., Djakarta.
Stonor, C. R.
1944. Observations on the elephant and other mammals in the Anamalai Hills of
Cochin. J. Bombay Nat. Hist Soc., 44, pp. 588-592.
Stott, K., Jr.
1964. Lungmanis: peace in a forest primeval. San Deigo Zoonooz, 37 (10), pp. 10-15.
Stremme, H.
1911. Die Saugetiere mit Ausnahme der Proboscidier, pp. 82-150. In Selenka, L and
FOODEN: LIONTAIL & PIGTAIL MACAQUES 165
M. Blanckenhorn, eds., Die Pithecanthropus-Schichten auf Java, Wilhelm
Engelmann, Leipzig.
Stresemann, E.
1939. Die Vogel von Celebes, pts. 1, 2. J. f. Orn., 87, pp. 299-425.
Sugiyama, Y.
1968. The ecology of the lion-tailed macaque [Macaca silenus (Linnaeus)] - A pilot
study. J. Bombay Nat. Hist. Soc, 65, (2), pp. 283-293.
Swindler. D. R., J. E. Sirianni, and L. H. Tarrant
1973. A longitudinal study of cephalofacial growth in Papio cynocephalus and
Macaca nemestrina from three months to three years. Symp. IV Int. Congr.
Primat., vol. 3: Craniofacial biology of primates, pp. 227-240. Karger, Basel.
Tanaka, J.
1965. Social structure of Nilgiri langurs. Primates, 6 (1), pp. 107-122.
Tashian, R. E., M. Goodman, V. E. Headings, J. De Simone, and R. H. Ward
1971. Genetic variation and evolution in the red cell carbonic anhydrase isozymes
of macaque monkeys. Biochem. Genet., 5, pp. 183-200.
Thomas, O.
1892. On the Mammalia collected by Signor Leonardo Fea in Burma and
Tenasserin. Ann. Mus. Civ. Stor. Nat. Genova, (ser. 2) 10, pp. 913-949.
1895. On some mammals collected by Dr. E. Modigliani in Sipora, Mentawei
Islands. Ann. Mus. Civ. Stor. Nat. Genova, (ser. 2) 14, pp. 660-672
1911. The mammals of the tenth edition of Linnaeus; an attempt to fix the types of
the genera and the exact bases and localities of the species. Proc. Zool. Soc.
London, 1911, pp. 120-158.
Thomas, 0. and E. Hartert
1894. First glimpses of the zoology of the Natuna Islands. III. List of the first
collection of mammals from the Natuna Islands. Novitat. Zool. 1, pp. 652-662.
Tickell, S. R.
1854-1875. Mammals of India. Unpublished manuscript in library of Zoological
Society of London.
Trevor-Jones, R.
1970. The musculo-skeletal anatomy of the baboon's tail. S. Afr. J. Sci., 66, pp.
316-323.
Tweedie, M. W. F.
1940. Report on excavations in Kelantan. J. Malay. Br. R. Asiatic Soc., 18, pp. 1-
22.
U. S. Board on Geographic Names
1966a. Official standard names gazetteer no. 96: Burma. Washington, D. C.
1966b. Official standard names gazetteer no. 97: Thailand. Washington, D. C.
1968a. Official standard names gazetteer no 13: Indonesia and Portugese Timor,
2nd ed. Washington, D.C.
166 FIELDIANA: ZOOLOGY, VOLUME 67
1968b. Official standard names gazetteer no. 22: Mainland China, 2nd ed., 2 vols.
Washington, D. C.
1970. Official standard names gazetteer no. 10: Malaysia, Singapore, and Brunei,
2nd ed. Washington, D. C.
1971. Official standard names gazetteer no. 58: South Vietnam, 2nd ed.
Washington, D. C.
1973. Official standard names gazetteer no. 73: Laos, 2nd ed. Washington, D. C.
Van Andel, T. H., G. R. Heath, T. C. Moore, and F. R. Mc Geary
1967. Late quaternary history, climate, and oceanography of the Timor Sea,
northwestern Australia. Amer. J. Sci., 265, pp. 737-758.
Van Peenen, P. F. D., P. F. Ryan, and R. H. Light
1969. Preliminary identification manual for mammals of South Vietnam. U. S. Nat.
Mus., Washington, D. C.
[Vigors, N. A.]
1829. Catalogue of the animals preserved in the museum of the Zoological Society.
Zoological Society, London.
Vui Sresthaputra, Saman Buravas and G. F. Brown
1951. Geologic reconnaissance of the mineral deposits of Thailand: Geology. Geol.
Surv. Bull., 984, pp. 25-50.
Vogel, C.
1966. Morphologische Studien am Gesichtsschadel catarrhiner Primaten. Bibl.
Prima tol., 4, pp. 1-226.
VOLZ, W. T. A. H.
1912. Nord -Sumatra, vol. 2. Dietrich Reimer, Berlin.
Wagner, J. A.
1840 [1839]. Schreber's Die Saugethiere, suppL vol 1. Expedition des Schreberschen
Saugthier- und des Esperschen Schmetterlingswerkes... , Erlangen. For date of
publication, see Proc. Zool. Soc. London, 1891, p. 591.
Waterhouse, G. R.
1838. Catalogue of the Mammalia preserved in the Museum of the Zoological
Society of London, 2nd ed. Richard and John E. Taylor, London.
Webb-Peploe, C. G.
1947. Field notes on the mammals of South Tinnevelly, South India. J. Bombay
Nat. Hist. Soc, 46, pp. 629-644.
Weiss, M.L., M. Goodman, W. Prychodko, G. W. Moore, and T. Tanaka
1973. An analysis of macaque systematics using gene frequency data. J. Hum.
Evol., 2 (2), pp. 213- 226.
Weiss, M.L., M. Goodman, W. Prychodko, and T. Tanaka
1971. Species and geographic distribution patterns of the macaque prealbumin
polymorphism. Primates, 11(1): 75-80.
FOODEN: LIONTAIL & PIGTAIL MACAQUES 167
White, R. J., C. R. Blaine, and G.A. Blakley
1973. Detecting ovulation in Macaca nemestrina by correlation of vaginal
cytology, body temperature and perineal tumescence with laparoscopy. Amer. J.
Phys. Anthropol., 38(2), pp. 189-194.
Wilson, D. R.
1970. A functional morphology of the tail, ischial callosities and pelvocaudal
musculature in Macaca. Ph. D. Thesis. Univ. of Chicago, 232 pp.
1972. Tail reduction in Macaca, pp. 241-261. In Tuttle, R., ed., The functional and
evolutionary biology of primates, Aldin-Atherton, Chicago.
WlSLOCKJ, G. B.
1933. The reproductive systems, pp. 231-247. In Hartman, C. G. and W. L. Straus,
Jr., eds., The anatomy of the rhesus monkey Macaca mulatto, Williams &
Wilkins, Baltimore.
Wroughton, R. C.
1915a. Bombay Natural History Society's Mammal Survey of India, Burma and
Ceylon. Report No. 16. J. Bombay Nat. Hist. Soc., 23, pp. 460-480.
1915b. Bombay Natural History Society's Mammal Survey of India, Burma and
Ceylon. Report No. 17. J. Bombay Nat. Hist. Soc., 23, pp. 695-720.
1916a. Bombay Natural History Society's Mammal Survey of India, Burma and
Ceylon. Report Nos. 20, 21, 22. J. Bombay Nat. Hist. Soc, 24, pp. 291-316.
1916b. Bombay Natural History Society's Mammal Survey of India, Burma and
Ceylon. Report Nos. 24, 25, 26 J. Bombay Nat. Hist. Soc, 24, pp. 749-782.
1918. Summary of the results from the Indian Mammal Survey of the Bombay
Natural History Society. J. Bombay Nat. Hist. Soc, 25, pp. 547-598.
1921. Summary of the results from the Indian Mammal Survey of the Bombay
Natural History Society. J. Bombay Nat. Hist. Soc, 27, pp. 520-534.
Yerkes Newsletter
1971. [Photograph]. Yerkes, Newsl., 8 (1), p. 14.
Yin, U Tun
1954. Wild life preservation and sanctuaries in the Union of Burma. J. Bombay
Nat. Hist. Soc, 52, pp. 264-284.
ZlMMERMANN, E. A. W.
1780. Geographische Geschichte des Menschen und der vierfuasigen Thiere, vol. 2,
Quadrupeden. Weygandschen Buchhandlung, Leipzig.
ZUCKERMAN, S.
1933. Functional affinities of man, monkeys, and apes. Kegan, Paul, Trench,
Trubner & Co., London.
1937. The duration and phases of the menstrual cycle in primates. Proc ZooL Soc.
London (Ser. A) 107, pp. 315-329.
Zukowsky, L.
1952. Die Wanderu - Schweinsaffen-Mischlinge des Frankfurter Tiergartens. Zool.
Gart., (N. F.), 19(1), pp. 1-26
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.
Pub
Live Music Archive
Librivox Free Audio
Metropolitan Museum
Cleveland Museum of Art
Internet Arcade
Console Living Room
Books to Borrow
Open Library
TV News
Understanding 9/11