UNIVERSITY OF

ILLINOIS LIBRARY

AT URBANA-CHAMPAIGN

BIOLOGY

MAR 2 6 198S

^1 c

FIELDIANA
Zoology

Published by Field Museum of Natural History

New Series, No. 9

TAXONOMY AND EVOLUTION

OF THE SINICA GROUP OF MACAQUES:

2. SPECIES AND SUBSPECIES ACCOUNTS

OF THE INDIAN BONNET MACAQUE, MACACA RADIATA

JACK FOODEN

m i o mi

3Ji5 LIBRARY OF THS

ffSM&ttsa

October 15, 1981
Publication 1325

TAXONOMY AND EVOLUTION

OF THE SINICA GROUP OF MACAQUES:

2. SPECIES AND SUBSPECIES ACCOUNTS

OF THE INDIAN BONNET MACAQUE, MACACA RADIATA

FIELDIANA
Zoology

Published by Field Museum of Natural History

New Series, No. 9

TAXONOMY AND EVOLUTION

OF THE S/MG4 GROUP OF MACAQUES:

2. SPECIES AND SUBSPECIES ACCOUNTS

OF THE INDIAN BONNET MACAQUE, MACACA RADIATA

JACK FOODEN

Research Associate

Field Museum of Natural History

Professor of Zoology
Chicago State Unix>ersity

Accepted for publication February 2, 1979
October 15, 1981
Publication 1325

Library of Congress Catalog Card No.: 81-65061

ISSN 0015-0754

PRINTED IN THE UNITED STATES OF AMERICA

CONTENTS

List of Illustrations vi

List of Tables vii

Abstract 1

Introduction 1

Species and Subspecies Accounts 2

Macaca radiata (E. Geoffroy, 1812)

Distribution 2

External characters 2

Pelage color variation: geographic, ontogenetic, seasonal 7

Cranial characters 11

Natural history 14

Addendum 34

Macaca radiata radiata (E. Geoffroy, 1812)

Synonymy 34

Types 35

Type-locality 35

Distribution 35

Diagnostic pelage color characters (prime pelage) 35

Specimens examined 35

Macaca radiata diluta Pocock, 1931

Synonymy 35

Types 36

Type-locality 36

Distribution 36

Diagnostic pelage color characters (prime pelage) 37

Specimens examined 37

Gazetteer of Macaca radiata Localities

Macaca radiata radiata 37

Macaca radiata diluta 45

Literature Cited 47

LIST OF ILLUSTRATIONS

1. Locality records of Macaca radiata and neighboring species of macaques 4

2. Crown pelage in Indian bonnet macaque, Macaca radiata, compared with that in

Sri Lanka toque macaque, M. sinica 6

3. Seasonal fading and molting of pelage in Macaca radiata radiata 7

4. Cranial characters in Macaca radiata radiata 12

5. Allometry of rostral length vs. postrostral length in immature and adult specimens

of Macaca radiata compared with corresponding allometry in M. sinica 13

6. Vegetation zones and typical annual rainfall curves in peninsular Indian habitat

of Macaca radiata 15

7. Troop of Macaca radiata radiata near Dharwar 20

8. Bivariate plots of age-sex composition in troops of Macaca radiata radiata 25

9. Reproductive seasonality in Macaca radiata radiata 27

VI

LIST OF TABLES

1. External measurements and ratios in adult Macaca radiata 3

2. Seasonal distribution of prime and faded pelage in Macaca radiata specimens . . 10

3. Latitudinal analysis of greatest skull length in adult specimens of Macaca
radiata 14

4. Cranial dimensions and ratios in immature and adult specimens of Macaca
radiata 14

5. Altitudinal distribution of accurately known Macaca radiata localities 16

6. Food plants reported for Macaca radiata 17

7. Interspecific dominance and subordination in reported natural encounters be-
tween bonnet macaques (Macaca radiata) and liontail macaques (Macaca silenus) 19

8. Troop size in Macaca radiata radiata 21

9. Home range area and population density in troops of Macaca radiata radiata ... 22

10. Age-sex composition in observed troops of Macaca radiata radiata 24

11. Dominance shifts of 10 adult males in Somanathapur troop of Macaca radiata
radiata 26

12. Intertroop movements reported in Macaca radiata 30

13. Reproductive success and neonatal sex ratio in troops of Macaca radiata observed
during birth season 31

14. Age distribution in two troops of Macaca radiata radiata studied near Dharwar,
northwestern Karnataka 32

15. External measurements of type-series of Macaca radiata diluta Pocock, 1931. ... 36

ABSTRACT

The present species account of Macaco, radiata (E. Geoffroy, 1812), the Indian
bonnet macaque, is based on study of all known museum specimens (128) and
on review of relevant literature. The northern limit of distribution of this species
is not the Godavari River, as generally assumed, but instead is 100-250 km south
of the Godavari River for most of this river's length. External and cranial char-
acters of M. radiata are analyzed, with particular emphasis on geographic, on-
togenetic, and seasonal variation of pelage color and on allometry of craniofacial
proportions. An extensive review of the natural history of this species focuses
on habitats, predators, diet, relations with other primate species, troop size and
composition, home range area, intertroop behavior patterns, breeding behavior
and seasonality, life- table survivorship probabilities, and mortality factors. Two
subspecies are recognized: M. r. radiata, which is widely distributed, and M. r.
diluta Pocock, 1931, which is restricted to a relatively narrow southeastern coastal
zone. For each subspecies, basic information is provided on synonyms, types,
type-locality, distribution, and diagnostic external characters. A gazetter of M.
radiata localities includes information on available museum specimens and on
published field notes by collectors or observers.

INTRODUCTION

This account of Macaca radiata, the South Indian bonnet macaque, is the second
part of a planned comprehensive monographic revision of the four species that
constitute the sinica group of macaques (M. sinica, M. radiata, M. assamensis, M.
thibetana). The first part of this monograph, an account of M. sinica, was pub-
lished elsewhere (Fooden, 1979, pp. 109-140). For the present work, 128 museum
specimens of M. radiata were examined; about half of these were wild-collected,
and the other half were captives (see list of specimens examined). Specimens
examined are preserved in the following institutions, which are subsequently
cited by means of the indicated abbreviations:

AIUZ Anthropologisches Institut der Universitat Zurich

AMNH American Museum of Natural History, New York

ANSP Academy of Natural Sciences, Philadelphia

BM British Museum (Natural History), London

BNHS Bombay Natural History Society, Bombay

FMNH Field Museum of Natural History, Chicago

IRSN Institut Royal des Sciences Naturelles de Belgique, Brussels

MNHN Museum National d'Histoire Naturelle, Paris

NHMB Naturhistorisches Museum, Basel

NHRM Naturhistoriska Riksmuseet, Stockholm

2 FIELDIANA: ZOOLOGY

RMNH Rijksmuseum van Natuurlijke Historie, Leiden
USNM U.S. National Museum of Natural History, Washington, D.C.
ZMUZ Zoologisches Museum der Universitat, Zurich
ZSBS Zoologisches Sammlung des Bayerischen Staates, Munich

ZSI Zoological Survey of India, National Zoological Collection, Cal-

cutta

SPECIES AND SUBSPECIES ACCOUNTS

Macaca radiata (E. Geoffroy, 1812). Synonymies under subspecies headings.

Distribution (fig. 1; Fooden et al., 1981, fig. 1). — Peninsular India from Cape
Comorin (8°13'N, 77°34'E) northward to the northern end of the Western Ghats
(20°47'N, 73°55'E), the Manjra Plateau (18°17'N, 75°57'E) and the northern end
of the Velikonda Range, Eastern Ghats (15°35'N, 79°08'E), with an isolated outlier
population in the Vijayawada Hills (16°32'-16°45'N, 80°33'-80°38'E). This de-
marcation of the northern limit of distribution of M. radiata, which also approx-
imately coincides with the southern limit of distribution of M. mulatta, is based
on results of a recent survey (Fooden et al., 1981, fig. 1) undertaken after the
present work had been submitted and accepted for publication. The boundary
between M. radiata and M. mulatta that is specified here is 100-250 km south of
the Godavari River, which, following Blyth (1863, p. 9) and Blandford (1889, p.
23), routinely has been cited as the interspecific boundary (Pocock, 1939, p. 38;
Prater, 1971, p. 35; Hill, 1974, p. 705; Roonwal & Mohnot, 1977, p. 192). Evidence
that the Godavari is not the interspecific boundary previously was provided by
Rae (in Southwick et al., 1961, p. 538), Spillett (1968a, p. 8), and Krishnan (1972,
p. 540).

E. Geoffroy (1812, p. 98) originally gave the range of this species merely as
"l'lnde." The first reasonably precise locality record was provided by F. Cuvier
(1820, p. 1), who reported on specimens collected along the Malabar Coast by
M. Housard, a French naval officer. Jerdon (1867, p. 12) evidently was the first
zoologist specifically to note the allopatric relationship between M. radiata and
M. mulatta.

Macaca radiata was successfully introduced in the Mascarene Islands, east of
Madagascar, apparently by sixteenth-century Portuguese mariners (Leguat,
[1708] 1891, p. 204; LaCaille, 1763, pp. 216, 230; Newton, 1893, p. 217; de Sornay,
1949, p. 65). During the nineteenth century, museum specimens were collected
both in Mauritius (I. Geoffroy, 1851, p. 26) and Reunion (Schlegel, 1876, p. 99),
which are the two largest islands in the Mascarene group. Another species of
long-tailed macaque, M. fascicularis, also was introduced by Europeans in the
Mascarene Islands (I. Geoffroy, 1851, p. 29); published reports of monkeys in
these islands (all originally introduced) usually are not sufficiently precise to
permit unambiguous species determination (cf. Pridham, 1849, p. 226).

An early report of M. radiata in "Formosa" (Taiwan) evidently is based on an
introduced or captive specimen (ZSI 11826; Blyth, 1860, p. 88; 1863, p. 8; 1875,
p. 8). An erroneous report of this species in Nepal (Hodgson, 1834, p. 96) is
based on misidentified M. mulatta (cf. Hodgson, 1832, p. 339).

External characters (figs. 2, 3, 7; table 1). — Head and body length 455 ± 31 mm
in 11 adult females, 527 ± 38 mm in 12 adult males; relative tail length (T/HB)
1.10 ± 13 in 10 adult females, 1.07 ± .09 in 12 adult males; weight 3.85 ± .50
kg in 14 adult females, 6.67 ± .85 kg in 13 adult males. Dorsal pelage color

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA
Table 1. External measurements and ratios in adult Macaca radiata.

Head and body Relative

Relative

Locality

length

tail length

ear length

Weight

No.1

(mm)

(T/HB)
Adult females, M.

(E/HB x 100)
r. radiata

(kg)

10

480, 5152

1.16, 1.10

7.7, 7 A

15

470, 470

1.02, 1.05

8.1, 7.7

3.63, 3.63

33

455

0.95

7.7

3.86

38

4.992

45

450

1.13

8.2

. . .

59

400

0.88

9.3

. . .

62

445

1.18

7.9

. . .

71

470

1.16

8.1

?3

375, 430

?10.0, 8.4

4.16, 4.16

T4

2.93 - 4.42 (N = 8

X±S.D.

(N)

458 ±31 (10)

1.07 ±.10 (9)
Adult females, M.

8.0 ±.5 (9)
r. diluta

3.85 ±.50 (14)

87

?328, 420

?1.60, 1.36
Adult males, M. r

?15.2, 10.7
radiata

7

5072

1.26

7.9

7.26

10

570, 570

0.96, 1.09

7.5, 7.9

. . .

12

590

1.14

8.1

8.85

15

530

1.06

7.7

6.58

26

520

1.08

7.7

5.44

33

495, 508, 530

0.98, 1.13, 0.94

7.7, 7.5, 7.4

, 5.90

59

450

1.06

8.9

63

510

1.10

6.9

1*

5.67-7.00 (N = 8)

X±S.D.

(N)

525 ±40(11)

1.07±.09 (11)
Adult male, M. r

7.7±.5(11)
diluta

6.67±. 85 (13)

82

5452

1.00

7.3

. . .

'For key to locality numbers, see distribution map (fig. 1).
^kin only; adult status inferred from external measurements.
'External measurements from Hill, 1974, p. 708.
♦Weights from Hartman, 1938, p. 468.
5X = 3.69, S.D. = 0.45.
*X = 6.60, S.D. = 0.44.

subspecifically and seasonally variable (see next section), prime pelage drab
grayish brown to golden brown, becoming drabber and variably streaked with
blackish on the lower back, hair banding variably conspicuous; outer surface of
thighs same color as back or drabber, outer surface of arms and shanks somewhat
paler, ochraceous gray to pale grayish brown; tail bicolor, dark brown to blackish
dorsally (slightly paler distally), pale grayish brown to ochraceous ventrally;
pelage of ventral surface of trunk and limbs pale ochraceous buff to whitish,
skin of chest and belly pigmented dark bluish gray or mostly unpigmented
whitish; crown hairs colored approximately like back (sometimes with blackish
tips), radiating from a central whorl to form a conspicuous cap (fig. 2), with
posterior crown hairs longer (extending to occiput) than anterior crown hairs
(extending to midway between vertex and brow ridges); anterior crown area
(between anterior cap hairs and brow ridges) covered with short hairs that
diverge laterally to form a median part; facial skin, including that of ears and
lips, pinkish buff to tan, occasionally scarlet in females (Anderson, 1879, p. 90;
Simonds, 1965, p. 175).

CLOSED SYMBOLS = SPECIMENS EXAMINED

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA 5

Head and body length in adult males averages about 15% greater than in adult
females, and weight in adult males averages about 75% greater than in adult
females. Conversely, relative tail length and relative ear length in adult males
average about 4%-7% smaller than in adult females.

Geographically, there seems to be a tendency for head and body length to
increase from south to north in M radiata (table 1); this tendency is more apparent
in female specimens than in male specimens examined. Although external di-
mensions in M. radiata overlap those in Sri Lanka M. sinica (Fooden, 1979, table
1), there is a consistent pattern of mean differences between these two species.
Head and body length averages about 10%-13% greater in M. radiata than in M.
sinica, weight averages about 9%-26% greater in M. radiata, relative tail length

Opposite:

Fig. 1. Locality records of Macaca radiata (circles) and neighboring species of macaques
(M. mulatto, triangles; M. silenus, stars; M. sinica, squares). Closed symbols indicate spec-
imens examined, open symbols indicate literature or sight records; heavy dashed line
indicates inferred border between subspecific ranges of M. r. radiata and M. r. diluta. For
details and references, see Gazetteer. New marginal locality records of M. radiata and M.
mulatto recently have been reported by Fooden et al. (1981, fig. 1).

Macaca r. radiata localities are as follows: 1, Yeur. 2, Salsette Island. 3, Trombay Hills.
4, Matheran. 5, Khandala. 6, Konkan. 7, Ghatmatha. 8, Dharwar, 18.4-29.5 km SW. 9,
Dharwar, 4.4-17.5 km SW. 10, Dharwar, 2,300 ft. 11, Gadag. 12, Vijayanagar. 13, Karwar.
14, Devikop. 15, Samasgi. 16, Sorab-Banavasi. 17, Halgalali; Kummuru; Nadhalli; Sorab;
Sorab-Shiralkoppa. 18, Karchikoppa; Malalagadde; Shigga. 19, Gersoppa. 20, Avalgodu;
Dodderi; Haravadike; Hosabale; Kuppe; Kuruvari; Kyasanur Forest; Nishrani; Tavanandi.
21, Keladi. 22, Marasa. 23, Yalgalali. 24, Sagar-Shimoga. 25, Srisailam. 26, "Malakonda-
penta." 27, Eastern Ghats. 28, Coromandel Coast. 29, Tirupati. 30, Tada. 31, Sholinghur.
32, Chiknayakanhalli. 33, Haleri. 34, Mysore-Mangalore. 35, Ranganathittoo Sanctuary.
36, Mysore-Bangalore, milestone 28, 30, 32. 37, Mysore-Bangalore, milestone 34, 36. 38,
Biligiri Rangan Hills. 39, Mysore-Bangalore, milestone 83.40, Bangalore. 41, Bangalore-
Krishnagiri. 42, Malur. 43, Kolar. 44, Jalarpet. 45, Wotekolli. 46, Nagarhole Wildlife Sanc-
tuary. 47, Byrankuppe. 48, Mysore; Ootacamund-Mysore, milestone 117; Ootacamund-
Mysore, milestone 121. 49, Ootacamund-Mysore, milestone 114. 50, Ootacamund-Mysore,
milestone 107, 108. 51, Ootacamund-Mysore, milestone 92. 52, Gundlupet-Chamrajnagar,
milestone 1, 2. 53, Hangala; Somanathapur. 54, Bandipur; Bandipur, 1 mile NE; Bandipur,
1 mile W; Bandipur Wildlife Sanctuary; Kakkenahala; Mangata. 55, Gudalur-Theppakadu;
Manali Theppakadu; Mudumalai Wildlife Sanctuary; Padamara; Theppakadu-Kargudi;
Veduru Guta. 56, Nilgiri Hills, 5,000-8,600 ft, 7,000 ft; Nilgiri Hills, N; Nilgiri Hills, W 57,
Nilgiri Hills. 58, Sigur Range. 59, Kotagiri; Nilgiri Hills, N and E. 60, Coonoor-Mettupa-
laiyam, milestone 12; Coonoor-Mettupalaiyam, milestone 17. 61, Siddharkovil. 62, Ku-
rumbapatti. 63, Shevaroy Hills. 64, "Tenmali." 65, Mattathur. 66, Sharnelli Estate. 67,
Nelliampathy Hills, N. 68, Nelliampathy Hills, S. 69, Anaimalai Hills. 70, Panniar. 71,
Palni Hills, N. 72, Machchur; Palni Hills, 5,500 ft.

Macaca r. diluta localities are as follows: 73, Pondicherry. 74, Kumbakonam. 75, Kodai-
kanal Road. 76, Kodaikanal Road, milestone 23. 77, Kambam. 78, Alleppey, S. 79, Periyar
Lake, N; Periyar Wildlife Sanctuary. 80, Varushnaad Valley. 81, Srivilliputtur-Mudaliar
Ootu. 82, Travancore. 83, Trivandrum. 84, Kuttalam. 85, Papanasam. 86, Naraikkadu
Estate. 87, Bhutapandi. 88, Aramboli, 5 miles S.

Macaca mulatto localities are as follows: A, Dangs (BM 31.1.11.1-3). B, Nagpur vicinity
(Jerdon, 1867, p. 11). C, Orcha Gay 1965a, pp. 210, 212). D, Malkangiri (BM 28.3.7.3). E,
Pakhal Lake Wildlife Sanctuary (Spillett, 1968a, p. 8). F, Hyderabad vicinity (Rae in South-
wick et al., 1961, p. 538). G, Nagarjunakonda Valley, "Siddeldar Hill" (ZSI, uncatalogued);
previously, this record was considered anomalous and therefore was thought to be the
result of human introduction (Agrawal & Bhattacharyya, 1976, p. 214), but the locality is
now known to be within the natural range of M. mulatto (Fooden et al., 1981, fig. 1).

Macaca silenus localities are shown here as in Fooden (1975, pp. 79, 168) and Green &
Minkowski (1977, p. 305); M. sinica localities are shown as in Fooden (1979, p. 131).

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA

Fig. 3. Seasonal fading and molting of pelage in Macaca radiata radiata: A, late dry season,
strongly faded specimen with large irregular pale buffy lateral blotches (BNHS 50479 01
5049 9 , Samasgi, Loc. No. 15, 12 March 1912); B, early rainy season, newly molted specimen
with small scattered tufts of unshed pale fur, two tufts of which are indicated by arrows
(BNHS 50529, Sharnelli Estate, Loc. No. 66, 28 May 1921); C, late rainy season, specimen
in prime pelage (BNHS 50489, Devikop, Loc. No. 14, 21 Nov. 1911).

averages about 13%-14% less in M. radiata, and relative ear length in males
averages about 16% less in M. radiata. All of these mean differences are in accord
with Bergmann's and Allen's rules, as noted previously (Fooden, 1971, p. 72).
Pelage color variation: geographic, ontogenetic, seasonal. — Pelage color in M. radiata
varies geographically (subspecifically), ontogenetically, and seasonally. In M. r.
radiata, which is the more broadly distributed of the two recognized subspecies,
the dorsal fur in adult specimens in prime pelage is drab grayish-brown ("brun-
verdatre," E. Geoffroy, 1812, p. 98) with relatively inconspicuous hair banding,
and the ventral fur is pale buffy to whitish; crown hairs in this subspecies
frequently are tipped with blackish, the upper back is variably washed with pale
yellowish, and the lower back is marked with a variably defined dark brown to
blackish median streak. In M. r. diluta, which is restricted to a narrow south-
eastern coastal zone (fig. 1), the dorsal fur is more brightly colored, pale yellowish
brown to golden brown, with hairs conspicuously banded dark brown basally
and bright yellowish distally, and the ventral fur is whitish; cap hairs in this
subspecies lack blackish tips, and the lower back, although slightly drabber than

8 FIELDIANA: ZOOLOGY

the upper back, lacks the blackish median streak that is variably developed in
M. r. radiata.

Within the range of each of the two recognized subspecies, dorsal pelage color
is relatively constant, with little or no tendency toward geographic intergradation
between subspecies. For example, a fairly dark grayish brown male specimen
of M. r. radiata (BM 30.11.1.29) collected at Gathmatha, near the northwestern
limit of the subspecific range, closely matches three males (BM 30.11.1.18-20)
collected at Haleri (600 km SSE of Gathmatha) and also matches another male
(BM 30.11.1.25) collected at Vijayanagar (375 km SE of Gatthmatha). A second,
slightly paler male (BM 30.11.1.24) collected at Vijayanagar matches a male (BM
21.11.5.2) collected at Sharnelli Estate (525 km S of Vijayanagar) and also matches
a female (BM 30.11.1.30) collected at Palni Hills (600 km SSE of Vijayanagar),
both localities near the southern limit of distribution of M. r. radiata. Similarly,
three seasonally faded (see below) specimens of M. r. radiata collected at Mar-
lakonda (BM 30.11.1.26-27 6 6, BM 51.608 9), near the northeastern limit of
the subspecific range, match a faded female specimen (BM 30.5.24.2) collected
at Kurumbapati (450 km S of Marlakonda), at the southeastern limit of the range.
Comparable networks of matching specimens interconnect other collecting lo-
calities within the range of M. r. radiata.

Five M. r. diluta skins collected at four localities (Travancore, ZSI 8495; Bhu-
tapandi, BM 30.11.1.31, BM 1937.5.26.1; Kambam, BM 30.11.1.28; Pondicherry,
RMNH 6f ) in extreme southeastern peninsular India are readily distinguishable
by their bright color and conspicuous hair banding from all M. r. radiata speci-
mens examined. The abrupt geographic transition from M. r. radiata coloration
to M. r. diluta coloration is indicated by the striking difference between a juvenile
male (BM 21.11.5.2) of the former subspecies collected at Sharnelli Estate and
a juvenile female (BM 30.11.1.28) of the latter subspecies collected at Kambam,
which is only about 120 km southeast of Sharnelli Estate. The Sharnelli specimen
shares the general drabness of other M. r. radiata specimens and, as previously
indicated, perfectly matches a specimen (BM 30.11.1.25) of M. r. radiata collected
at Vijayanagar, which is 525 km north of Sharnelli; the brightly colored Kambam
M. r. diluta specimen closely matches all four other specimens available of this
subspecies. The only specimen examined that shows some evidence of pelage
coloration intermediate between that of M. r. radiata and M. r. diluta is a subadult
male (ZSI 12007) collected in South Arcot (probably at "Tenmali"), near the
border between the subspecific ranges; the pale yellowish brown South Arcot
specimen, which is now somewhat arbitrarily assigned to M. r. radiata, tends
toward the brightness of M. r. diluta, but lacks the conspicuous hair banding
characteristic of this latter subspecies.

In addition to differing from each other in dorsal pelage color, M. r. radiata
and M. r. diluta also differ in pigmentation of the ventral skin (not hair), as noted
by Pocock (1931b, p. 278). In M. r. radiata, the skin of the chest and belly is
broadly and deeply pigmented dark bluish gray. By contrast, in four M. r. diluta
specimens examined (Bhutapandi, BM, 2; Kambam, BM, 1; Travancore, ZSI, 1),
the skin of this region is unpigmented whitish with, at most, only small irreg-
ularly dispersed blotches of pale bluish pigment; ventral skin pigmentation was
not examined in one M. r. diluta specimen (Pondicherry, RMNH). Predictably,
the geographically intermediate South Arcot specimen (ZSI), which is somewhat
intermediate in dorsal pelage color between typical M. r. radiata and M. r. diluta
(see above), also is approximately intermediate in extent and intensity of ventral

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA 9

skin pigmentation. (I thank Mr. P. K. Das, Zoological Survey of India, Calcutta,
for reexamining ventral skin pigmentation in two critical M. radiata specimens
in the ZSI collection.)

The evolutionary factors responsible for differentiation of M. r. radiata and M.
r. diluta are unclear. Although there is no apparent physiographic barrier between
the ranges of these two subspecies, these areas do differ in timing of the annual
monsoonal rainy season (fig. 6). In the range of M. r. radiata, the months of
maximum rainfall are June or July, and the dry season begins in October or
November; in the range of M. r. diluta, as a result of frontal convergence between
the northeast monsoon and the retreating Indian Ocean westerlies (Kendrew,
1953, p. 181), the months of maximum rainfall are October or November, and
the dry season begins in January. The adaptive significance, if any, of the re-
lationship between local rainfall patterns and subspecific differentiation in M.
radiata is not known.

The bright color of the upper back in M. r. diluta is similar to the color of the
same region in the toque macaque (M. sinica), which inhabits Sri Lanka across
the Palk Strait from the coastal range of M. r. diluta; however, M. sinica is readily
distinguishable from both subspecies of M. radiata by the long anterior crown
hairs (fig. 2) and the rufous to tawny color of the outer surface of the thighs in
the former species (Fooden, 1979, p. 110). Considered interspecifically, pelage
color in subspecies of M. radiata and M. sinica follows a step cline of increasing
brightness with decreasing latitude. In M. r. radiata, which is the most northerly
subspecies in these two species, the entire dorsal surface is drab, and the cap
and lower back are variably washed with blackish; in M. r. diluta, which ranges
to the southeast of M. r. radiata, the upper back is brightly colored, and the cap
and lower back lack the blackish wash that is frequently present in M. radiata;
in M. s. sinica, which ranges farther to the southeast in Sri Lanka across the Palk
Strait, the upper back, lower back, and outer surface of thighs are all brightly
colored; finally, in M. s. aurifrons, which is restricted to southwestern Sri Lanka,
the cap is marked anteriorly by a patch of bright yellow to golden, and there
is a strong tendency to general erythrism (Fooden, 1979, p. 114). Although this
unusual clinal pattern that encompasses four subspecies in two species attests
to the close relationship between M. radiata and M. sinica, its evolutionary sig-
nificance is otherwise obscure.

Neonatal pelage in M. radiata is dark (dark brown to near blackish), short,
fine, and sparse (Nolte, 1955, p. 83; Simonds, 1965, p. 181; 1974a, p. 155; Ra-
haman & Parthasarathy, 1969b, p. 150; Krishnan, 1972, p. 545; Hill, 1974, p. 706).
At age approximately one month, this dark neonatal pelage is rapidly replaced
by a second-stage infantile pelage that is similar in color and pattern to adult
pelage but is finer in texture (cf. BM 30.11.1.11, deciduous i 1; Gersoppa). Finally,
at age approximately one year, the second-stage infantile pelage is replaced by
a juvenile pelage that is essentially indistinguishable from that in adults (cf. BM
30.11.1.28, deciduous m2, permanent M2 erupting; Kambam). Males and fe-
males in M. radiata appear not to differ appreciably in pelage color or texture.

Conspicuous seasonal pelage variation (molting), which seems to be relatively
rare in macaque species, was first noted in bonnet macaques by Pocock (1931b,
p. 276). In M. r. radiata, molting evidently occurs in late May or early June (table
2), which coincides with the onset of monsoonal rains over most of the range
of this subspecies (fig. 6). Judging from specimens examined, molting is preceded
by fading of the dorsal pelage, which is first evident in mid-March (Samasgi,

10 FIELDIANA: ZOOLOGY

Table 2. Seasonal distribution of prime (P) and faded (F) pelage in Macaca radiata spec-
imens examined,1 showing that faded pelage prevails in M. r. radiata during March-June
(hot dry season preceding season of monsoonal rains).

Pelage condition in month collected3
Lo-
cality

No. N Jan. Feb. March April May June July Aug. Sep. Oct. Nov. Dec.

M. r. radiata

2P
4P
2P

IP

7

2

10

4

12

2

14

1

15

6

19

1

26

3

33

4

38

1

40

2

43

1

45

1

59

2

62

1

63

3

66

2

729

2

72io

1

Total

39

77

1

82

1

87

2

IP,4 5F

4P

1F(?)
2F

1P(?)5
3F

IF

IP

IP
1P(?),6 IF

2F

IP/ 2F
IP8 IP

IP

4P ... 1P,10F IF 4P, 5F 2P, IF . . . 2P IP . . . 5P 3P
M. r. diluta
IP

IP
2P

'These specimens were examined at different times in various museums; under these
circumstances, some variation in standards for judging pelage condition is unavoidable.
Despite this, the main pattern of seasonal pelage change seems reasonably clear.

2For key to locality numbers, see distribution map (fig. 1).

Specimens lacking date of collection are not included in this table.

4BM 30.11.1.15, adult male, collected 12 March 1912.

5BM 30.11.1.12, subadult female with infant (age approximately three months, infantile
pelage), collected 29 May 1912.

6BM 19.6.2.1, adult male, collected 25 June 1918.

7BNHS 5051, adult male, collected 22 May 1929.

8BNHS 5052, subadult female, collected 28 May 1921, newly molted (few scattered tufts
of faded hair still present; fig. 3).

9Palni Hills, 5,500 ft.

10Machchur.

5 specimens, 12 March) — the fifth month of the seven-month dry season. In
faded specimens, the dorsal pelage appears rough-textured and disheveled and
is conspicuously blotched with irregular patches of pale buffy hairs (fig. 3),
especially anteriorly and laterally. When the rainy season begins, shedding of
the faded pelage and concomitant exposure of the prime new pelage evidently
occurs fairly rapidly, as indicated by two specimens collected in May-June 1921
at Sharnelli Estate (southwestern peninsular India). In a subadult female (BNHS
5052) collected at this locality on 28 May, the dorsal pelage, which evidently is
in the final phase of molting, is mostly prime grayish brown washed with

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA 11

yellowish, but a few small tufts of longer faded buffy hairs remain unshed
laterally on the upper back (fig. 3); in a juvenile male (BM 21.11.5.2) collected
nine days later (6 June) at the same locality, the dorsal pelage is entirely prime
grayish brown, lacking any trace of faded hairs. The physiological or ecological
significance of dry season fading and rainy season molting in M. r. radiata is
unknown. Fading or molting is not evident in the few specimens available of
M. r. diluta.

Pigment reduction or absence has been noted rarely in captive M. radiata. An
albino male with pink irises was observed in 1936 at the Trivandrum Zoo, South
India, by Hill (1937, p. 212). A captive male with white fur and skin, but with
brown irises, was observed in 1836 in the London Zoo by Ogilby (1838, p. 312).
A pale golden brown subadult female museum specimen (USNM 122171) of
unknown history also appears to have abnormally reduced pigmentation.

Cranial characters (fig. 4; tables 3, 4). — Skull moderately small, greatest length
104.4 ± 2.4 mm in 10 wild-collected adult females, 120.0 ± 4.0 mm in 12 wild-
collected adult males; relative zygomatic breadth (ZB/GL) moderately great, 0.65
± .03 in 10 adult females, 0.67 ± .02 in 12 adult males; rostrum moderately
projecting, rostral-postrostral ratio 0.46 ± .02 in 10 adult females, 0.54 ± .02
in 12 adult males; supramaxillary ridges well defined, arching superioposteriorly
from canine alveolus to infraorbital rim; supraorbital ridges moderately thick
(ca. 6 mm in adult males); temporal lines weakly to moderately defined, usually
separate, rarely converging in old males to form a low (ca. 1 mm), posteriorly
restricted sagittal crest (FMNH 82802, Dharwar); nuchal crest moderately de-
veloped (1-7 mm high) in adult males.

In adult males (table 3), greatest skull length averages significantly larger in
six specimens collected north of 14°N lat. (122.9 ± 3.3 mm; extremes, 117.7-127.9
mm) than in six specimens collected south of 13°N lat. (117.2 ± 2.0 mm;
114.6-120.3 mm). In 10 adult female specimens examined, however, there is no
indication that greatest skull length increases from south to north. This contrasts
with latitudinal increase in head and body length, which is more evident in
female specimens than in male specimens examined (table 1).

Ontogenetically, relative zygomatic breadth (ZB/GL) increases slightly from
0.62 in one available infant to 0.67 ± 0.02 in 12 adult males (table 4). By contrast,
the ratio of rostral length to postrostral length, which is a measure of relative
size of splanchnocranium and neurocranium (Fooden, 1969, p. 41), increases
strikingly from 0.22 to 0.54 ± 0.02 in the same specimens (fig. 5). The power
function equation for the postnatal allometric relationship between rostral length
(y) and postrostral length (x) determined by the method of principal axes is:

log y = 3.395 log x - 4.860.

The 95% confidence limits for the slope are L, = 3.027 and L2 = 3.853.

Metric and nonmetric cranial characters in M. radiata overlap those in M. sinica,
the Sri Lanka toque macaque, which probably is the closest living relative of M.
radiata. However, greatest skull length in M. radiata specimens examined aver-
ages 6%-7% greater than in M. sinica (Fooden, 1979, p. 114). Conversely, relative
zygomatic breadth averages about 2%-3% less in adult M. radiata than in adult
M. sinica, and rostral-postrostral ratio may also average slightly less in adult M.
radiata than in adult M. sinica. These comparative observations generally agree
with those of Pocock (1939, p. 40), who characterizes the skull of M. radiata as
larger but relatively less robust than that of M. sinica (cf. Fooden, 1979, fig. 4).

*o °

12

ADULT MALES

60 70 80

Postrostral length (mm)

100

Fig. 5. Allometry of rostral length (y) vs. postrostral length (x) in immature and adult
specimens of Macaca radiata (open circles = females, closed circles = males; smaller circles
= immatures, larger circles = adults; solid line = principal axis) compared with corre-
sponding allometry in M. sinica (dashed line = principal axis). Macaca radiata principal
axis, log y = 3.395 log x - 4.860; M. sinica principal axis, log y = 4.317 log x - 6.507
(Fooden, 1979, p. 115).

13

14

FIELDIANA: ZOOLOGY

Table 3. Latitudinal analysis of greatest skull length in adult specimens of Macaca radiata;
means and standard deviations indicated for northern (xN) and southern (xs) halves of
female and male samples and for total (xTot) of each sample.

Latitude

Locality

No.1

l ICllgll

i viiiiii;

(°N)

Adult females

Adult males

18°59'

4

. . .

123.3

15°48'

26

117.7

15°28'

10

104.1

121.5, 127.8

15°19'

12

124.0

14°40'

15

99.0, 104.1

XN

123.0
= 122.9 ± 3.3 (n = 6)

12°27'

33

102.6

114.6, 117.1, 118.6

12°08'

45

xN

105.5
= 103.1±2.5(n =

5)

11°50'

63

120.3

11°45'

62

107.2

11°26'

59

107.7

116.3

11°20'

57

116.1

10°22'

71

104.9

8°16'

87

104.0,105.0

*s

= 105.8 ± 1.6 (n =

5)

xs

= 117.2±2.0(n = 6)

xTot

= 104.4 ±2.4 (n =

10)

XTot

= 120.0 ±4.0 (n = 12)

'For key to locality numbers, see distribution map (fig. 1).

Table 4. Cranial dimensions and ratios (x
of Macaca radiata.

S.D.) in immature and adult specimens

Skull,

Relative

Age class

N

greatest
length
(mm)

zygomatic
breadth
(ZB/GL)

Postrostral
length
(mm)

Rostral-

postrostral

ratio

Infants

1

71.0

.62

59.8

.22

Juveniles

7

93.0 ±10.1

.65 ±.02

72.2 ±5.1

.39 ±.07

Subadult females

8

99.5±6.7

.66 ±.02

74.6 ±2.7

.44 ±.03

Adult females

10

104.4±2.4

.65 ±.03

76.9 ±2.0

.46 ±.02

Subadult males

7

113.3±5.5

.66 ±.01

81.6±4.5

.50 ±.02

Adult males

12

120.0 ±4.0

.67 ±.02

83.1 ±2.6

.54 ±.02

Fifteen adult male M. radiata specimens without locality data, most or all of
which probably are zoo specimens, average significantly larger (greatest skull
length, 123.9 ± 4.5 mm; extremes 113.6-129.7 mm) than wild-collected spec-
imens reported above (table 3). This difference presumably is a result of artificial
conditions of captivity. Because of this difference, measurements of specimens
without locality data have not been included in the present analysis of cranial
characters in M. radiata; in previous revisions of macaque species, I have included
(injudiciously, it now appears) some measurements of known or probable zoo
specimens.

Natural history. — Macaca radiata inhabits monsoonal peninsular India (fig. 6),
where it occupies a broad range of wooded and partly wooded habitats from
sea level to 2,100 m (table 5). Above 2,100 m, sheltered upland forest (shola)
gradually yields to open grassland, and monkeys are absent; the highest peak
in peninsular India is 2,695 m. Forest types in which troops of M. radiata have
been encountered include deciduous (Sugiyama, 1968, p. 289; Kurup, 1971, p.
17), bamboo (Kinloch, 1923, p. 552; Nolte, 1955, p. 84), evergreen (Webb-Peploe,

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA

15

1947, p. 630; Hutton, 1949, p. 689; Sugiyama, 1972, p. 247), and shola (McCann
in Lindsay, 1926, p. 592; Khan, 1977, p. 517). Sightings of this species seem to
be more common in lowland deciduous and bamboo forests than in upland
evergreen and shola forests ( J. R. O'Brien, specimen tag, BNHS 5052, Sharnelli
Estate; Kinloch, 1923, p. 552; Hutton, 1949, p. 689; Krishnan, 1972, p. 536).
Troops of M. radiata often occur in heavily disturbed or manmade habitats,
including cultivated areas where roadside trees are nearby (Simonds, 1965, p.
175); village outskirts (Nolte, 1955, p. 79); temple compounds (Shortridge in
Wroughton & Ryley, 1913, p. 60); and suburbs, urban parks, and urban market
areas (W. Elliot, 1839, p. 95; Rahaman & Parthasarathy, 1967, p. 253).

80°

+ 20°

VEGETATION ZONES
TROPICAL FOREST

Evergreen: Wet (W), Dry (D)

Semi-Eve rgreen

Deciduous: Moist (M), Dry (D)

Thorn
SUB-TROPICAL FOREST, WET (S-Tr)
TEMPERATE FOREST, WET (Tm)

• MANGAIORE
3275 MM/YR

Fig. 6. Vegetation zones (Roonwal & Mohnot, 1977, p. 27) and typical annual rainfall
curves (Walter et al., 1975, map 5) in peninsular Indian habitat of Macaca radiata.

16 FIELDIANA: ZOOLOGY

Table 5. Altitudinal distribution of accurately known Macaca radiata localities; for details
see Figure 1 and Gazetteer.

Altitude (m) Locality Nos.

0-499 12, 19, 66

500-999 10, 14, 15, 20,1 45, 53, 54,2 69, 71

1,000-1,499 33, 40, 43, 55,3 56,4 63,5 70, 726

1,500-1,999 56/ 59,8 639 72, 10 80

2,000-2,499 56"

!Kyasanur Forest.

^andipur, 1 mile NE; Bandipur, 1 mile W; Mangata.

3Mudumalai Wildlife Sanctuary.

4Nilgiri Hills, N; Nilgiri Hills, W.

^hevaroy Hills, 4,500 ft.

6Machchur.

7Nilgiri Hills, N & E.

8Kotagiri.

9Shevaroy Hills, 5,400 ft.

10Palni Hills, 5,500 ft.

"Nilgiri Hills, 7,000 ft.

Daytime activity (0600-1800 hr) in M. radiata probably is about 90% arboreal
(10% terrestrial) in forest troops and about 70% arboreal in troops in cultivated
areas (Sugiyama, 1972, p. 251). Nighttime sleeping is exclusively arboreal, except
in treeless urban areas where M. radiata sleeps on roofs or ledges of buildings
(R. S. N. Pillay, specimen tag, BM 1937, 5.26.1, Bhutapandi; Rahaman & Par-
thasarathy, 1967, p. 254). Forest troops probably change sleeping trees nearly
every night (Nolte, 1955, p. 79). When moving through trees, M. radiata runs
nimbly along the upper surface of branches, like other macaques, but it does
not bound, as do sympatric langurs (Presbytis entellus) (Krishnan, 1972, p. 539).
Macaca radiata also is adept at scaling cliffs. On the ground, these macaques
often stop and stand bipedally to survey their surroundings (Sugiyama, 1972,
p. 250). When frightened on the ground, they usually flee into the canopy
(Simonds, 1965, p. 196; Rajagopal, 1965, p. 229), but they also have been ob-
served to flee on the ground in bamboo forest (Nolte, 1955, p. 84). Strong
sunlight is avoided when possible (Rahaman & Parthasarathy, 1969a, p. 273).
These monkeys swim well and appear to enjoy water (Stonor, 1944, p. 591); near
a temple at Sholinghur, northeastern Tamil Nadu, a mother bonnet macaque
was observed swimming with her infant on her back (Krishnan, 1972, pi. 3).

Judging from alarm and flight reactions observed in M. radiata, probable nat-
ural predators are tigers, leopards, smaller wild felids, and large predatory birds
(Sugiyama, 1972, p. 261). Crocodiles and large snakes are other possible natural
predators. Domestic dogs are greatly feared by M. radiata (Nolte, 1955, p. 84)
and have been observed to kill an infant monkey (Simonds, 1965, p. 179). How-
ever, a jackal did not provoke a flight response, but instead was mobbed and
driven off by a troop of bonnet macaques (Nolte, 1955, p. 84).

The natural diet of M. radiata, like that of other macaques, is predominantly
vegetarian and probably consists mostly of fruit (table 6); however, flowers,
young leaves, shoots, plant pith, and sap also are eaten. Judging from the known
sample of diverse plant species that have been observed to provide food for M.
radiata, it seems likely that parts of most plant species in the environment are
eaten at least occasionally; however, strychnine fruit and cultivated castor beans
are said to be avoided (Dixon, 1894, p. 104; Simonds, 1965, p. 175). Wild figs

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA
Table 6. Food plants (wild or semi-wild) reported for Macaca radiata.

17

Reference

Food plant:

Part

Preference

(footnote

family/species1

consumed

level

No.)

ANACARDIACEAE

[Mangifera indica] (mango)

Fruit, flowers, leaves

Favored

2, 3

APOCYNACEAE

Carissa conject (karwanda)

Fruit, young leaves

Main food
(seasonal)

4

CANNACEAE

[Carina sp.]

Pith

. . .

5

ELAEOCARPACEAE

[Elaeocarpus] tuberculatus

Fruit

Relished

6

GRAMINAE

Bambusa arundinacea (bamboo)

Seeds, shoots

Major food

7

Dendrocalamus strictus (bamboo)

Seeds, shoots

Major food

7

giant bamboo

Leaf buds

8

bamboo spp.

Shoots

Preferred

2, 5

grasses

Seeds, blades

Main food

4, 5, 8

LEGUMINOSAE

[Acacia spp.]

Fruit, young leaves

Main food

4

Butea frondosa

Flowers

9

Dalbergia sisoo

Fruit, shoots

9

Pongamia glabra (pongam)

Fruit, young leaves

Main food
(seasonal)

4

Tamarindus indica (tamarind)

Fruit, young leaves

Main food

4, 5, 7, I

MELIACEAE

Azadirachta indica (nihm)

Fruit, young leaves

Main food

4

MORACEAE

Ficus bengalensis (banyan)

Fruit

Major food

7

F. benjamina (banyan)

Fruit

Major food

7

[Ficus spp.] (figs)

Fruit, young leaves

Main food

2,4, 5

[Morus sp.] (mulberry)

Fruit

Preferred

5

ROSACEAE

[Pygeum] gardneri

Fruit

Relished

6

VERBENACEAE

Lantana aculeata

Fruit, flowers, young
leaves

Main food

2,4

'Names cited as in original source, except where bracketed.

^olte, 1955, p. 79.

3Rahaman & Parthasarathy, 1967, p. 253.

<Sugiyama, 1972, pp. 249, 252.

5Rahaman & Parthasarathy, 1969b, p. 152.

6McCann in Lindsay, 1926, p. 592.

7Simonds, 1965, p. 177.

"Krishnan, 1972, p. 538.

'McCann, 1933, p. 810.

may constitute the most important natural food for M. radiata (cf. Simonds, 1965,
p. 179), as apparently is true for M. sinica (Fooden, 1979, p. 123). Animal food
items known to be eaten by M. radiata include adult and immature insects
(notably flies and grasshoppers), spiders, lizards (one observation), and birds'
eggs (Hutton, 1949, p. 689; Nolte, 1955, p. 79; Rahaman & Parthasarathy, 1969b,
p. 152; Sugiyama, 1972, p. 252; Krishnan, 1972, p. 539). Much of the terrestrial
foraging of M. radiata apparently is in search of arthropod prey. This species also

18 FIELDIANA: ZOOLOGY

expends considerable effort in obtaining birds' eggs and has been observed
swimming across a fast- flowing stream to raid a nesting ground. Troops that
inhabit agricultural areas probably obtain about 40% of their food by raiding
cultivated crops (Simonds in Lancaster & Lee, 1965, p. 493); because this species
is regarded as semi-sacred, its crop raiding engenders a tense, ambivalent re-
lationship between bonnet monkeys and local farmers (Buchanan, 1807, p. 55;
Baker in Blyth, 1859, p. 283; Shortridge in Ryley, 1913b, p. 492; Rajagopal, 1965,
p. 234). Troops in temple and market areas often are deliberately provisioned,
or they may obtain their food by raiding houses, shops, and refuse heaps (Ra-
haman & Parthasarathy, 1967, p. 254; Singh & Sachdeva, 1977, p. 606).

Depending on seasonal abundance, feeding occupies about 40% of daylight
hours (Rahaman & Parthasarathy, 1969b, p. 154). Particularly intense feeding
periods occur in the morning shortly after awakening and in late afternoon
before retiring (Nolte, 1955, p. 80). Like other macaques, M. radiata habitually
feeds by first stuffing food in the cheek pouches and later chewing and swal-
lowing it (Ogilby, 1838, p. 310; Giebel, 1859, p. 66; Rajagopal, 1965, p. 229). The
moisture content of foods evidently is not sufficient to supply the metabolic
needs of M. radiata, at least during the dry season, and these macaques therefore
drink daily at streams or other open water (Nolte, 1955, p. 80). Drinking typically
occurs in late afternoon but may also occur at other times; water is gulped
directly by mouth. At the height of the dry season, long daily excursions may
be necessary to find water (Koyama, 1973, p. 229).

Macaca radiata is partly sympatric with one other species of macaque, M. silenus,
and with two species of langurs, Presbytis entellus and P. johnii. However, eco-
logical preferences apparently are not identical in M. radiata and the other three
sympatric monkey species. Macaca silenus and P. johnii are restricted to upland
evergreen forest (Poirier, 1970, p. 257; Fooden, 1975, p. 81; Khan, 1977, p. 517;
Green & Minkowski, 1977, p. 290; Prasad et al., 1979, p. 738), whereas M. radiata
is more common in deciduous forest (see above). Presbytis entellus ranges farther
into dry scrubby areas than does M. radiata (Simonds, 1965, p. 176). The diet
of M. radiata is very similar to that of M. silenus (Fooden, 1975, p. 82; Green &
Minkowski, 1977, p. 295); the diets of P. johnii and P. entellus include more leaves
and less fruit and insects than that of M. radiata (Sugiyama, 1967, p. 223; Poirier,
1970, p. 270).

Interspecific encounters between M. radiata and M. silenus occasionally have
been observed in the upland evergreen habitat of the latter species. Judging
from the few available published reports (table 7), M. radiata is dominant in
some of these encounters, M. silenus is dominant in some, and neither species
is dominant in some. Relative to the distribution of M. radiata, however, the
distribution of M. silenus obviously is highly restricted (fig. 1). This, together
with the broad discontinuity that exists between the range of M. silenus and that
of its nearest relative, M. nemestrina leonina (Fooden, 1975, p. 69), suggests that
expansion of the range of M. radiata has been correlated with reduction of the
range of M. silenus (Fooden, 1976, p. 228). Regardless of which of these two
macaque species is dominant in direct interspecific contacts, it seems clear that,
in the long-term zoogeographic view, M. radiata generally has gained territory
at the expense of M. silenus; this probably is a secondary result of changes in
climate and vegetation. Recently observed interspecific encounters between M.
radiata and neighboring M. mulatta were not overtly antagonistic (Fooden et al.,
1981, in press).

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA 19

Integeneric encounters between M. radiata and Presbytis entellus have been
frequently observed. In these encounters, M. radiata tends to be dominant and,
as an initial response, P. entellus tends to withdraw from contact (Simonds, 1965,
p. 195; Sugiyama, 1968, p. 289, and 1972, p. 253; Poirier, 1970, p. 261; Makwana
& Majumdar, 1980, p. 126). Later, however, once the dominant status of M.
radiata has been established, these two species may coexist peacefully in close
proximity or even in temporarily intermixed troops (fig. 7; Shortridge in Wrough-
ton, 1912, p. 1176; Jay, 1965b, p. 541; Sugiyama, 1967, p. 226; Krishnan, 1972,
p. 550; Koyama, 1973, p. 227). Intergeneric encounters between M. radiata and
P. johnii are relatively poorly reported; Hutton (1949, p. 689) indicates that P.
johnii is dominant in these encounters, whereas Webb-Peploe (1947, p. 629),
Simonds (1965, p. 195), and Poirier (1970, p. 261) indicate that these encounters
are peaceful or that M. radiata is slightly dominant.

Troop member counts are available for 38 troops of M. r. radiata studied in
two areas in western Karnataka (table 8; NW Karnataka, Dharwar-Haliyal, 13
troops; SW Karnataka, all other localities listed, 25 troops). Forest and roadside
troops studied in this subspecies average about 29 individuals per troop (range

Table 7. Interspecific dominance and subordination in reported natural encounters
between bonnet macaques (Macaca radiata) and liontail macaques (Macaca silenus).

a. M. radiata dominant

1. Sugiyama (1968, p. 289), Panniar (ca. 10°00'N, 77°00'E):

In the [evergreen forest] study area bonnet macaques (Macaca radiata) lived within
the range of lion-tailed macaque troops. Smaller in size but with a larger troop size
(about 30 head) the bonnet macaque is more terrestrial and is commoner in dry
deciduous forests and in the vicinity of the villages in drier areas. In the study area
it was as arboreal as the lion-tailed macaques. Troops of both species were not overtly
antagonistic. A troop of bonnet macaques moved among lion-tailed macaques and
sometimes even travelled following the latter. Lion-tailed macaques usually did not
react to the appearance of bonnet macaques but sometimes moved away slowly from
them. Although smaller, bonnet macaques were dominant.

b. Interspecific tolerance, or dominance not observed

2. Poirier (1970, p. 261), Nilgiri Hills, W (ca. 11°25'N, 76°30'E):

At the border of tracts 2 [moist evergreen] and 3 [sholas] . . . overlapped two species
of Indian macaques, M. radiata and M. silenus.

3. Green and Minkowski (1977, p. 300), locality unspecified:

Bonnet macaques (M. radiata) typically occur in the drier deciduous forests and
scrub jungles at much lower elevations, but sometimes seasonally move up into the
wet forests inhabited by lion-tailed monkeys. The few times we saw bonnets with M.
silenus, no antagonism occurred. . . . However, resource competition with M. radiata
may have prevented M. silenus from successfully occupying the lower elevation forests
in which M. radiata is so well established.

c. M. silenus dominant

4. Vincent Marie in Buffon & Daubenton (1766, p. 171), Malabar district (ca. 11°15'N,
76°00'E):

. . . the other [three species of] monkeys [presumably including M. radiata] have
so much respect for [M. silenus] that they humble themselves in its presence, as if
they were able to recognize its [M. silenus] superiority, (translated from French)

5. Webb-Peploe (1947, pp. 629-630), Naraikkadu Estate (ca. 8°30'N, 77°30'E):

... in evergreen forest . . . [M. radiata] will keep out of the way when the Lion-
tailed Monkey is about. . . . Occasionally [they] meet on the [arboreal] bridge. The
Bonnet Monkey . . . disappears when the Lion-tailed Monkey is in the vicinity.

si

20

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA

21

Table 8. Troop size in Macaca radiata radiata.

Sample area (Loc. No.)1

Dharwar-Haliyal (8)
Byrankuppe (47)
Moyar R.-Theppakadu (54)
Subtotal

Dharwar-Haliyal (9)

Mysore vicinity-Somanathapur
(36, 48, 50, 51, 53)
Subtotal

Bangalore (40)

Mysore (48)

Subtotal
All troops5

No. Individuals per troop

of

troops x±S.D. Extremes

Reference

Forest troops

3 33.7±3.5
2 36.0±2.83
_23 17.0±14.1
7

30-37 Sugiyama, 1972, p.255

34, 382 Nolte, 1955, p. 78

7, 27 Simonds, 1974a, p. 152

29.6 ±10.7 7-37

Roadside troops

10 28.9 ±7.4 16-44 Sugiyama, 1972, p. 255;

Koyama,4 1973, p. 228

6 29.5 ±15.4 15-59 Simonds, 1973, p. 601;

1974a, p. 152

16 29.1 ±10.6 15-59

Urban troops

14 21.1±8.9

23

10-13

23

10-43

15 21.3±8.6

38 26.1 ±10.4 7-59

Rahaman &

Parthasarathy, 1967,

p. 253
Singh & Sachdeva,

1977, p. 606

'For key to locality numbers, see distribution map (fig. 1).
including, respectively, two and five infants in arms.

'These two troops had been recently formed as a result of fission of a larger troop.
*Troop E, 30 individuals, including seven infants.

*Cf. estimated troop size recently recorded in 29 additional troops, x = 33.2 ± 24.5,
extremes 10-100 (Fooden et al., 1981, table 1).

7-59). Urban troops of the same subspecies studied in Bangalore and Mysore
average about 21 individuals per troop (range 10-43) and therefore generally
seem to be somewhat smaller than forest and roadside troops (cf. Krishnan,
1972, p. 538). In the subspecies M. r. diluta, in extreme southeastern peninsular
India, troop size may average larger than in M. r. radiata, judging from a few
rough troop census estimates published by Hill (1937, p. 212) and Hutton (1949,
p. 689); the former reports that two temple troops of M. r. diluta each included
at least 50 individuals, and the latter indicates that an unspecified number of
forest troops of the same subspecies also each included more than 50 individuals.

Solitary individuals, living independently of any troop, have never been ob-
served in M. radiata (Rahaman & Parthasarathy, 1967, p. 254; Sugiyama, 1972,
p. 262; Simonds, 1973, p. 600), although such solitary individuals are well known
in other species of macaques (Fooden, 1975, p. 51). One small detached all-male
group (two adults, one subadult) has been recorded; this group lived within the
home range of a troop of 28 members and evidently was composed of troop
outcasts (Simonds, 1973, p. 600).

Home range area in M. r. radiata averages about 200 hectares in nine forest
and roadside troops and about 100 hectares per troop in two urban botanical
garden troops (table 9). Daily foraging range is highly variable but may average
about 50% of the total home range (Nolte, 1955, p. 79; Rahaman & Parthasarathy,
1969a, p. 274; Sugiyama, 1972, p. 249). Population density averages about one
monkey per five hectares. Assuming that an average troop member weighs

22

FIELDIANA: ZOOLOGY

Table 9. Home range area and population density in troops of Macaca radiata radiata.

Home

No. of

range

Density

Reference

troop

area

(ind./

(footnote

Locality1

members

Forest

(hectares)
troops

hectare)

No.)

Byrankuppe (47)

34

150

0.23

2

Bandipur A, 1 mile NE (54)

±260

3

Bandipur B, 1 mile W (54)

±260

3

Kakkenahala (54)

6

±130

±0.05

3

Mangata (54)

28 +

±260

±0.11

3

Means

±212

±0.13

Roadside troops

Dharwar D (9)

32

40

0.80

4

Dharwar E (9)

30

220

0.14

5

Hangala (53)

46

±260

±0.18

3

Somanathapur (54)

58

±2606

±0.22

3

Means

Urban

±195
troops

±0.33

(in botanical garden)

Bangalore 1 (40)

22

±100

±0.22

7

Bangalore 2 (40)

20

±100

±0.20

7

Means

±100

±0.21

locality numbers (in parentheses) as in distribution map, Figure 1.
2Nolte, 1955, p. 78.
Simonds, 1965, pp. 178, 180.
Sugiyama, 1972, p. 250.
5Koyama, 1973, pp. 225, 228.

6Area estimated from map of home range (Simonds, 1965, p. 178, fig. 6-2); this estimate
differs from home range area of 2 miles2 (520 hectares) specified in text (ibid., p. 177).
7Rahaman & Parthasarathy, 1969a, p. 269.

about 3 kg (cf. M. sinica, studied by Dittus, 1975, p. 144), the estimated average
biomass density in M. radiata is approximately 0.6 kg per hectare. One excep-
tional troop that occupies an unusually small home range (table 9, Dharwar
Troop D, 40 hectares) has a population density of nearly one monkey per hectare.
Long-term studies indicate that, despite occasionally observed fissions and mi-
grations (Rajagopal, 1965, p. 231; Kawamura in Sugiyama, 1972, p. 263; Simonds,
1974a, p. 153; Singh & Pirta, 1978, p. 265), troops generally tend to remain
relatively stable in their home range and total number of members (Sugiyama,

1972, p. 247, two years' observation; Rahaman & Parthasarathy, 1969a, p. 268,
three years' observation; Simonds, 1965, p. 176, and 1974a, p. 153, nine years'
observation).

In suitable habitats, troops of M. radiata generally are closely packed, with
about 5%-25% overlap between home ranges of adjacent troops (Simonds, 1965,
p. 178; Rahaman & Parthasarathy, 1969a, p. 270; Sugiyama, 1972, p. 248; Koyama,

1973, p. 229). Two of 38 troops studied (table 8) have home ranges that lie
completely within the home ranges of two other troops (Simonds, 1974a, p. 153;
Koyama, 1974, p. 227); such ranges that overlap completely may be characteristic
of newly formed sister troops that have recently developed by fission of a single
large ancestral troop (cf. Simonds, loc. cit.). By contrast, some areas of apparently
suitable habitat are only sparsely occupied by M. radiata, and home ranges of

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA 23

neighboring troops may be separated by up to 4 km of unoccupied territory
(Simonds, 1965, p. 178).

The long-term stability of home ranges in M. radiata probably is a result, at
least in part, of competitive social interactions between adjacent troops. These
interactions are known to include intertroop fights, vocal and visual threat be-
haviors, and mutual avoidance (see below). It seems likely that the wide range
of observed intertroop behaviors may represent successive stages in develop-
ment of a relationship of peaceful coexistence along intertroop borders. At a
presumed early stage in development of an intertroop relationship, incursions
of one troop into the home range of an adjacent troop are known frequently to
result in violent arboreal battles with serious injuries inflicted on some of the
combatants (Rajagopal, 1965, p. 230; Rahaman & Parthasarathy, 1969a, p. 271;
Yoshiba in Sugiyama, 1972, p. 263). In subsequent encounters, threatening vo-
calizations may replace direct physical conflict, as observed by Rahaman &
Parthasarathy (loc. cit.):

... on three occasions we noticed troupe 1 trying to invade the core area of
troupe 2 . . . , which resulted in serious fights on the first two occasions and in
just an aggressive exchange of overtones on the third.

Aggressive staring may represent a later stage in ritualization of intertroop
conflict behavior, as suggested by the following observation of Simonds (1965,
p. 195):

When the two [adjacent] groups met the subadult and adult males would move
toward the other group, and then sit and look at each other with approximately
20 feet between them. Then the males in one of the groups would begin to drift
back in the opposite direction and the groups would separate.

Ultimately, in the presumed final stage of intertroop accommodation, one
troop may immediately and automatically withdraw in response to advance into
its range of an adjacent, more dominant troop (Rahaman & Parthasarathy, 1969b,
p. 151; Sugiyama, 1972, p. 263; Kawamura in Koyama, 1973, p. 229; Singh &
Sachdeva, 1977, p. 605). In all reported instances of automatic withdrawal, the
troop that withdrew contained fewer members than the troop that advanced.
Occasionally a small subordinate troop may move temporarily into the home
range of a larger dominant troop, but only when the larger troop is absent from
that part of its range (Sugiyama, 1972, p. 263). Adjacent troops, however, ap-
parently never commingle (cf. Nolte, 1955, p. 79), and even large dominant
troops that have advanced into the range of an adjacent subordinate troop
apparently usually return to their own home range (Sugiyama, 1972, p. 263).

Age-sex composition data are available for 35 troops of M. r. radiata, and sex
ratio data only are available for 18 additional troops (table 10, fig. 8). Observers
have distinguished four age-sex classes in these troops: (1) infants, (2) juveniles,
(3) subadult and adult females, and (4) subadult and adult males. Because of
the annual birth peak in M. radiata (fig. 9), age cohorts of immatures are rea-
sonably discrete. Infants are less than one year old and usually are still nursing
or are otherwise closely dependent on their mothers. Juveniles, one year old to
less than four years old, are more or less independent of their mothers but have
not entered the breeding population; although laboratory studies have shown
that juvenile males as young as age two and one-half years are physiologically
capable of copulation and ejaculation (Rosenblum & Nadler, 1971, p. 394), such
sexual behavior by juveniles has not been observed in free-ranging natural troops
(see below). Subadults and adults, age four years and older, are actual or po-
tential breeders (reproductives); sexual maturity in males and females usually

24 FIELDIANA: ZOOLOGY

Table 10. Age-sex composition in observed troops of Macaca radiata radiata.

Total

Refrence

Total

Reference

troop

Age-sex

(footnote

troop

Age-sex

(footnote

members

composition1

No.)

members

composition1

No.)

7

2:1:3:1

2

27

9:9:5:4

5

10

4:3:1:2

3

27

9:6:6:6

2

11

4:5:1:1

3

28

12:7:5:4

3

14

5:2:5:2

3

28

9:8:7:4

5

14

5:5:4:0

3

28

8:9:6:5

4

15

5:1:4:5

4

30

10:10:6:4

5

16

7:6:1:2

3

30

8:11:4:7

4

16

5:5:4:2

5

30

11:4:8:7

6

17

7:3:6:1

3

31

16:10:3:2

3

20

6:3:5:6

3

32

9:7:8:8

5

20

6:5:5:4

3

32

9:7:10:6

5

20

6:6:3:5

4

33

11:8:8:6

5

22

11:3:5:3

3

34

5:3:17:97

8

23

5:4:7:7

3

34

13:8:8:5

5

23

7:5:6:5

5

37

13:11:9:4

5

24

8:7:6:3

5

38

8:5:12:137

8

25

8-9:1:7:8-9

4

43

12:20:4:7

3

27

7:8:5:7

3

44

13:11:13:7

5

59

17:18:10:14

2

:First entry = No. of adult or subadult females (age >4 yr.); second entry = No. of adult
or subadult males (age 3=4 yr.); third entry = No. of juveniles (age 1-3 yr.); fourth
entry = No. of infants (age <1 yr.).

2Simonds, 1974a, p. 152.

3Rahaman & Parthasarathy, 1967, p. 253.

Simonds, 1973, p. 601.

^ugiyama, 1972, p. 255.

6Koyama, 1973, p. 228.

7Age-sex categories probably not coordinate with those in other studies cited (Simonds,
1965, p. 181); data not included in Figure 8.

8Nolte, 1955, p. 78.

is achieved during the breeding season following the fourth birthday, which is
about age four and one-half years. At the onset of sexual maturity, females are
close to their adult size (head and body length ca. 450 mm, weight 4 kg), and
males the same age are only slightly larger than females; however, males con-
tinue to grow at least two or three more years after achieving sexual maturity,
and they ultimately reach an adult head and body length that is 10%-15% greater
than in adult females (weight more than 50% greater than in females).

In troops for which data are available (fig. 8), infants and juveniles (nonre-
productives) generally are outnumbered by subadults and adults (reproduc-
tives). The mean ratio of infants and juveniles to subadults and adults in 35
troops is 0.75 (S.D. = 0.36, extremes 0.19-1.63); this ratio seems to be somewhat
lower in urban troops in Bangalore than in roadside troops in the Dharwar and
Mysore areas. Among reproductives, the mean sex ratio of subadult and adult
males to subadult and adult females in 53 troops is 0.84 (S.D. = 0.34, extremes
0.12-2.00); in only 10 of these 53 troops does the number of breeding males
exceed the number of breeding females (fig. 8). The number of infants generally
averages about 80% of the number of juveniles. Mean composition in an average
size troop of 26 individuals would be eight subadult or adult females, seven
subadult or adult males, six juveniles, and five infants. Part of the observed high

AGE COMPOSITION OF TROOPS
NON-REPRODUCTIVES (Y)/REPRODUCTIVES (X)
N = 35 ©/

10 IS 20

NO. SUBADULTS AND ADULTS

(REPRODUCTIVES)

25

IN TROOP

SEX RATIO OF REPRODUCTIVES
MALES (Y)/FEMALES (X)
N =53

•V/

V

NO

S 10 IS 20

FEMALES (SUBADULT, ADULT) IN TROOP

• Bangalore
14 troops

■ Dharwar area
13 troops

a Mysore area
8, 26 troops

Fig. 8. Bivariate plots of age-sex composition in troops of Macaca radiata radiata: A, non-
reproductives vs. reproductives in 35 troops (table 10); B, male reproductives vs. female
reproductives in 53 troops (table 10; Simonds, 1973, p. 600).

25

26

FIELDIANA: ZOOLOGY

variability of troop composition may be attributable to variation in the season
of observation, which fails to control for seasonal birth and death peaks.

Social relations within troops of M. radiata generally conform to the sex-age
dominance system that is usual in macaques (Nolte, 1955, p. 78; Simonds, 1965,
p. 182; Rahaman & Parthasarathy, 1969b, p. 151; Sugiyama, 1972, p. 255; Koyama,
1973, p. 228). Adult males play a key role in leading the troop during movement,
in controlling fighting between troop members, in giving alarm calls, and in
defending the troop against external dangers. Among adult males, dominance
rank is closely related to body size and, especially, to canine size (Gee, 1964, p.
20; Simonds, 1965; p. 185). Dominance shifts in one carefully studied troop
usually span only one or two ranks per year, but three adult males in this troop
shifted three or four ranks within one year (table 11). Dominance rank is man-
ifested in the feeding and drinking order of individuals in a troop (Sugiyama,
1972, p. 256; Koyama, 1973, p. 228) and therefore presumably is related to
survival when food or water is scarce.

Although the dominance hierarchy in M. radiata is generally similar to that
in other macaques, observers agree that display of dominance and subordination
behaviors in M. radiata appears to be somewhat more flexible than in other
species, particularly M. mulatta and M. fuscata (Simonds, 1965, p. 186; Sugiyama,
1972, p. 264). Observers also indicate that social relations in troops of M. radiata
are characterized by an unusually high incidence of certain social behaviors:
namely, intense grooming; clasped resting and sleeping, often in clusters (cf.
Kaufman & Rosenblum, 1966, p. 219); mixed-age playing; and homosexual
mounting. Individual relations in all of these high-incidence social behaviors
frequently fail to conform to expected patterns of dominance and subordination.

Breeding is strongly seasonal in M. radiata (fig. 9). More than 80% of 308
copulations observed in two carefully studied troops in Bangalore occurred dur-
ing September, October, and November (Rahaman & Parthasarathy, 1969b, p.
156), which approximately coincides with the postmonsoon (northeast monsoon)
season. These same three months also constitute the peak mating season ob-
served in two other carefully studied troops, one near Mysore and the second
near Dharwar (Simonds in Lancaster & Lee, 1965, p. 494; Sugiyama, 1972, p.
252). In the troop studied near Dharwar, most of the relatively few copulations
that occurred outside of the September-November mating season involved the
alpha male (nine of 14 copulations) and his favorite female partner; of the re-

Table 11. Dominance shifts of 10 adult males in Somanathapur troop of Macaca radiata
radiata (Simonds, 1965, p. 191; 1974a, p. 158).

Male

Dan

Zeb

Pirn

Hala

One-Eye

Kink

Andy

Rock

Butch

Shorty

Estimated

age in
1961-1962

Prime
Prime
Late prime
Early prime
Early prime
Prime
Very old
Late prime
Old
Old

1
2
3
4
5
6
7
8
9
10

Dominance rank on date

Oct. 1961 June 1962

4
1
5
3
2
6
Deceased
7
8
9

July-Sept.
1963

?
1
4
?
?

Deceased
?
?

?

Jan. -May

1970

High ranking

Absent

Absent

Rank unclear

Absent

Absent

Deceased

?

?

?

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA

Rainfall (mm)

27

50

o

15 -

Cop ulat io ns or

Cop u la tion Days

n = 353

10

Births
n = 47

JULY AUG SEPT OCT NOV DEC JAN FEB MAR APR MAY JUNE

Fig. 9. Reproductive seasonality in Macaca radiata radiata. References: Rainfall — Walter
et al., 1975, map 5 (monthly means for Belgaum, Mysore, and Bangalore, which are
weather stations near or at localities where reproduction observations were made). Cop-
ulation activity — Rahaman & Parthasarathy, 1969b, p. 156 (copulations); and Sugiyama,
1972, p. 253 (copulation days). Births — Rahaman & Parthasarathy, 1969b, p. 156; Sugiyama,
1972, p. 253; Koyama, 1973, p. 228; Simonds, 1974a, p. 154; for five additional births
observed on unspecified dates between 1 Dec. 1974 and 15 May 1975, see Singh & Sachdeva
(1977, p. 606); and for two more births recorded in March-May 1977, see Singh & Pirta
(1978, p. 267).

maining five copulations observed outside of the regular mating season in this
troop, four involved a young subadult that was not seen to copulate during the
regular mating season; the other copulation outside of the regular mating season
involved another subadult male; three adult males in this troop were never
observed to copulate outside of the regular mating season.

28 FIELDIANA: ZOOLOGY

All births (N = 54) that have been recorded in wild troops of M. radiata are
restricted to the period December-June (fig. 9; 47 births in specified months,
seven births in unspecified months within the December-May interval). This
broadly overlaps the dry season in most of peninsular India. Significantly, it is
the mating season (see above), not the birth season, that is timed to occur during
the rainy months when food and water are most abundant. About 80% of the
accurately dated births occurred during February, March, and April, which is
approximately five months after the peak mating period; this is as expected
because the gestation period in M. radiata is five to six months. The complete
absence of births during the period July-November suggests that the relatively
few copulations that occur five months earlier (February-June) are inherently
infertile.

The timing of natural breeding seasonality in M. radiata apparently depends
on environmental cues, presumably correlated with rainfall. In a laboratory
colony of this species that was maintained for about three years in Davis, Cal-
ifornia, the annual birth peak shifted to April-July Qudge & Rodman, 1976, p.
535), which is about three months later than in wild troops in India.

Males and females apparently enter the breeding population about age four
and one-half years (Sugiyama, 1972, pp. 253, 259). Each breeding female prob-
ably is in estrus about four or five days of each approximately 30-day menstrual
cycle (cf. Hartman, 1938, p. 470). Because cycles of different females in the same
troop usually are not in phase, normally only one or two females are in estrus
on any given day during the mating season (Rahaman & Parthasarathy, 1971,
p. 100). While in estrus, each female copulates eight to 12 times (total) with two
to four male partners. Females usually copulate with more than one male on
each day of their estrus period; one female was observed to copulate successively
with three different males within a four-minute time span (Simonds, 1965, p.
194).

Subordinate males are not excluded from access to estrous females, but dom-
inant males probably engage in significantly more copulations. In one troop that
included six sexually mature males and nine sexually mature females, the alpha
male and beta male each copulated on eight days during 30 days of observation
during the mating season, two less dominant adult males copulated on six and
five days, a lower ranking subadult male copulated on five days, and the lowest
ranking subadult male did not copulate at all (Sugiyama, 1972, p. 252). In a
troop that included three sexually mature males and six sexually mature females,
one male (presumably the alpha male) copulated 26 times during the observation
period, whereas the other two males copulated 17 times and 13 times (Rahaman
& Parthasarathy, 1971, p. 100). The correlation between dominance status and
sexual activity in male M. radiata also has been demonstrated in laboratory
experiments (Coe, 1977, p. 4208). Although exclusive consort relationships are
not maintained, there apparently is a tendency for most copulations to occur
between preferred partner pairs.

Social tensions are high during the mating season (Simonds, 1977, p. 161).
Although dominant males usually tolerate copulations between estrous females
and lower ranking males, dominant males occasionally have been observed to
interrupt such matings by an attack on the lower ranking male (Rahaman &
Parthasarathy, 1969b, p. 155; Sugiyama, 1972, p. 258). Perhaps to avoid such
attacks, a lower ranking male accompanied by an estrous female sometimes

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA 29

moves more than 500 m away from other troop members before copulating
(Sugiyama, loc. cit.).

Copula tory behavior in M. radiata usually is initiated by the male partner (93%
of 59 copulations observed by Simonds, 1965, p. 194; 78% of 308 copulations
observed by Rahaman & Parthasarathy, 1969b, p. 155). Experimental evidence
indicates that males recognize estrous females at a distance by an unidentified
olfactory cue or pheromone (Rahaman & Parthasarathy, 1971, p. 98); this phero-
mone presumably is a component of secretions produced by the specially en-
larged cervix that characterizes M. radiata and other members of the M. sinica
species group (Hill, 1932, p. 314; Fooden, 1971, p. 71; Kanagawa et al., 1973, p.
211). Having perceived this olfactory signal, a male typically approaches the
posterior end of an estrous female, pushes the tail aside, visually inspects the
perineal area, smells or tastes secretions in the vagina, and proceeds to mount
and copulate. Close examination of the perineum seems to be a ritual, because
practically all females that are approached are subsequently mounted (Rahaman
& Parthasarathy, 1969b, p. 155); the pheromone that is perceived at a distance
and that induces the male to approach probably is the main effective stimulus
in male sexual arousal and copulatory behavior. Macaco radiata typically is a
single mount ejaculator, with five to 30 intromissive thrusts to ejaculation and
a total intromissive mount interval of six to 23 seconds (Simonds, 1965, p. 194;
Rajagopal, 1965, p. 232; Nadler & Rosenblum, 1969, p. 484).

Occasional transfer of individuals between troops of M. radiata is an obvious
requisite for maintenance of generic continuity within the species. Such transfers
are not often observed, however, and hence, are only poorly documented (table
12). The rarity or absence of solitary individuals in M. radiata (see above) indicates
that a bonnet macaque that leaves one troop normally either rapidly joins another
troop or fails to survive. Five of six individuals observed in intertroop shifts
were males, all either subadult or adult; of these, two were high-ranking (beta,
gamma), but none was an alpha male. All four shifts that are reasonably ac-
curately dated occurred during the rainy season (fig. 9); only one of these cer-
tainly occurred during the mating season. Based on these skimpy data, the
following series of provisional hypotheses may be proposed for future investi-
gation: (1) Under natural conditions, all males of M. radiata probably leave their
natal troop before reaching their first breeding season, most likely as late ju-
veniles or early subadults; Koyama (1973, p. 228) provides evidence that a male
may remain in his natal troop with his mother as late as age four years (cf.
Simonds, 1974a, p. 157). (2) Most males probably switch troops at least once
more during their life, but not as an alpha male. (3) Most intertroop shifts
probably occur during the rainy season, often during the first three months of
the rainy season and therefore before onset of the peak mating season. It should
be noted that intertroop shifts may be geographically impossible in some widely
separated urban troops (Rahaman & Parthasarathy, 1967, p. 251).

Troop reproductive success, measured as the average number of live births
per year per sexually mature female is 0.88 in eight M. radiata troops that were
observed through a complete breeding season (table 13). (Although single births
are the rule in M. radiata and other macaques, twinning may occasionally occur
[Webb-Peploe, 1947, p. 630].) High reproductive success in M. radiata also is
indicated by incomplete data on 19 additional troops that may not have been
observed through a complete breeding season (Rahaman & Parthasarathy, 1967,

30

FIELDIANA: ZOOLOGY

Table 12. Intertroop movements reported in Macaca radiata.
Intertroop mover

Date

29-30 May 1962

1-7 June 1962

4 Sept. 1962 ,
1962

Summer 1963

July-Sept. 1963 or
Jan.-May 1970

Sex

Age

Original

dominance

rank

6 14-17 yr.

6-7 yr.
[ad.]

ad.

subad.

6-10

4-7

'Rahaman & Parthasarathy, 1969a, p. 268.
2Sugiyama, 1972, p. 262.
3Kawamura in Koyama, 1973, p. 229.
Simonds, 1974a, p. 158; cf. Simonds, 1965, p.
5Simonds, 1973, p. 600.

Intertroop
movement
observed

Refer-
ence
(footnote
No.)

Shift from Troop 2 to
Troop 1 (Bangalore)

Permanent disappearance
from Troop D (near
Dharwar)

{Temporary disappearance
from Troop D
Reentry into Troop D

Shift from Troop E to
Troop C after death of
leader of Troop C (near
Dharwar); repeated
shifting between troops;
finally, permanent entry
into Troop C as new
leader

Shift from Somanathapur
troop to Hangala South
troop; shift completed in
one afternoon, rank
unchanged

Shift from Moyar R. troop
to detached group of two
isolated males

185.

p. 253; Simonds, 1973, p. 601); of these, four include as many infants as sexually
mature females (reproductive success 1.00), and one reportedly includes two
more infants than sexually mature females (reproductive success 1.40). The
neonatal sex ratio in four carefully studied troops averages 1.1 male infant to
1.0 female infant (table 13), which is not significantly different from 1:1 (X2 =
0.125; 0.9 > P > 0.5). Obviously, there is no indication in these data that the sex
ratio at birth directly determines the sex ratio among sexually mature adults,
where females generally outnumber males (fig. 8; cf . Rahaman & Parthasarathy,
1967, p. 254).

Available cross-sectional census data on age structure in two carefully studied
troops of M. radiata permit a first-approximation life-table analysis of survivor-
ship probability in this species (table 14). Assuming that the two troops studied
are reasonably typical of the species and are relatively stable in composition, it
may be inferred that there probably are three peaks in the age-specific mortality
curve. Not surprisingly, these three inferred mortality peaks are (1) during the
first year of life; (2) during the period between age four and six years, when
individuals enter the breeding population; and (3) during the period of old age,
starting about age 15 years.

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FIELDIANA: ZOOLOGY

Table 14. Age distribution in two troops of Macaca radiata radiata studied near Dharwar
(Loc. No. 9), northwestern Karnataka.

No. of troop members in each age class

^ge1 (yr.)

Troop D2

Neonate

6(2dd/49 9)

1

3 (3/0)

2

4 (1/3)

3

3 (3/0)

4

3(2/1)

0-4

19 (11/8)

5-9

5 (1/4)]

10-14

4 (2/2) \

15-(19)

4 (2/2) J

Total

32 (16/16)

Troop E3

7(4<J6739?;

3(2/1)

4(2/2)

I (0/1)
4 (1/3)
19 (1/10)

II (3/8)
30 (12/18)

Total

13(66 6/79 9)

6 (5/1)
8 (3/5)
4 (3/1)

7 (3/4)

38 (20/18)

24 (8/16)
62 (28/34)

Estimated during birth season. 2Sugiyama, 1972, pp. 253, 259. 3Koyama, 1973, p. 228.

Survivorship of neonates to age one year probably is only about 50% (table
14). Sugiyama (1972, p. 253) records that one of six neonates in one troop
disappeared within eight days of birth, and Simonds (1974a, p. 154) records that
two of five neonates in another troop died within one month after birth. Singh
& Pirta (1978, p. 267) report that two infants born during an unusual troop
migration disappeared within five days. Other authors (Hutton, 1949, p. 689;
Rajagopal, 1965, p. 233; Rahaman & Parthasarathy, 1969b, p. 157) report seeing
near-dead neonates that were abandoned by their mothers and dead neonates
that were carried by their mothers for several days. Nolte (1955, p. 85) observed
that a helpless infant was abandoned (?temporarily) by its mother when the
mother joined in a troop attack on an unidentified predator; such abandoned
infants sometimes may be retrieved by juvenile or subadult males (Sugiyama,
1972, p. 261). The fact that most births occur at the end of the dry season (fig.
9), when food and water are relatively scarce, may be a factor that contributes
to the high infant mortality rate.

The annual age-specific mortality rate from age one year to at least age four
years evidently is much lower than during the first year of life, with the result
that most one-year-olds probably survive to age four years (table 14). This gen-
eralization is based on detailed age data available for Dharwar area Troops D
and E (table 14), but it is also supported by less precise data available for 33
additional troops listed in Table 10; the ratio of infants to juveniles age one to
three years in Troops D and E (13 infants: 18 juveniles = 0.72) is reasonably
close to the corresponding ratio in 33 additional troops listed in Table 10 (149.5
infants: 182 juveniles = 0.82).

Evidence from Dharwar area Troop D, supported by comparable but less
complete evidence from Troop E, indicates that age four to five years, like age
birth to one year, is a period of unusually high mortality risk (table 14). In Troop
D, the number of individuals in the infant cohort (age one year) is six; the
average number of individuals in each cohort from age one through four years
is about three; and the average in each cohort from age five years to approxi-
mately age 19 years is about one. Age four to five years, as previously indicated,
is the age of puberty, and the high mortality rate that apparently characterizes
this age presumably is a result of hazards related to entering the breeding
population. For males, these probably are hazards associated with dominance

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA 33

fights and with shifting from one troop to another; for females, the nature of
the hazards of puberty are less clear, but they may be obstetrical hazards of
primaparity. Higher male mortality risk during this period of adolescence prob-
ably is the main reason why troop breeding populations usually include more
females than males (fig. 8).

Following the demographic crisis of puberty, the mortality rate in M. radiata
apparently declines, with the result that most monkeys that reach age six to
seven years probably survive at least 10 years more, to old age. This span, from
age six to seven years to about age 15 years, probably is the prime of life, the
period when individuals reach their highest level in the dominance hierarchy.
In one long-term study, it was observed that two of six high-ranking males
survived at least nine years after reaching adulthood (table 11).

Finally, some time after age 15 years, the cumulative effects of old age, among
which worn teeth may be especially important, result in another period of high
mortality. A 20-year-old bonnet macaque probably is near the upper age limit
for survival in the wild. Under artificial conditions of captivity, however, indi-
viduals may survive to about age 30 years (Prater, 1971, p. 36). Average life
expectancy at birth under natural conditions probably is about five years (table
14).

Causes of morbidity and mortality that have been directly observed in M.
radiata include fights within and between troops, attacks by predators, infectious
diseases, and old age. Of these, wounds sustained in the course of intra troop
fights have been most frequently reported (Simonds, 1965, pp. 179-180; Raha-
man & Parthasarathy, 1968, p. 261; 1969b, p. 151; Krishnan, 1972, pp. 540, 550;
Sugiyama, 1972, p. 256; Simonds, 1974b, p. 178); these fights usually are related
to competition for food and/or dominance position and often result in serious
injury or death. Intertroop fights are less frequently reported, but these also
may be extremely violent and produce serious casualties (Rajagopal, 1965, p.
230; Yoshiba in Sugiyama, 1972, p. 263). A predatory attack by domestic dogs
that resulted in the death of an infant bonnet macaque has been reported by
Simonds (1965, p. 179). Fatal diseases that have been reported or identified in
M. radiata include bubonic plague, which is said to have killed large numbers
of bonnet macaques near Gadag, 50 km east of Dharwar (Shortridge in Wrough-
ton, 1912, p. 1176), and Kyasanur Forest disease, a tick-borne virus infection
that has been isolated in individuals of this species that were found dead in
forests in Shimoga District, 100 km south of Dharwar (Work & Trapido, 1957,
p. 342; Work, 1958, p. 250). Old age is suspected as the cause of death of a
senescent male that disappeared from the Somanathapur troop, south of Mysore,
between January and April 1962 (Simonds, 1965, p. 179).

Individual survival and population size under natural conditions apparently
are strongly influenced by predation pressure and by carrying capacity of the
habitat. Population growth may be explosive in bonnet macaque groups that are
artificially introduced in areas where there are no predators and no nearby troops
to compete for resources. Krishnan (1972, p. 538) reports on such a group that
was introduced in the Kodikadu Reserved Forest, Point Calimere. This group
increased in five or six years from 30 individuals to about 100 individuals, which
implies that 10 or 15 infants were born each year and that nearly all of these
infants survived to the end of the five- to six-year period of observation (cf . table
14). A similar explosive population increase also may have occurred when M.
radiata was introduced in the Mascarene Islands (see under Distribution).

34 FIELDIANA: ZOOLOGY

Addendum. — Six important works, listed below, that relate to the natural
history of M. radiata were published while the present paper was in press.

Glick, B. B. 1980. Ontogenetic and psychobiological aspects of the mating activities of
male Macaca radiata, pp. 345-369. In Lindburg, D. G., ed., The Macaques: Studies in
Ecology, Behavior and Evolution. Van Nostrand Reinhold, New York.

Handen, C. E., and P. S. Rodman. 1980. Social development of bonnet macaques from
6 months to 3 years of age: A longitudinal study. Primates, 21 (3): 350-356.

Kuruvilla, G. P. 1980. Ecology of the bonnet macaque (Macaca radiata Geoffroy) with
special reference to feeding habits. J. Bombay Nat. Hist. Soc, 75 (suppl): 976-988.

Makwana, S. C. 1980. Observations on population and behavior of the bonnet monkey,
Macaca radiata. Comp. Physiol. Ecol., 5: 9-12.

Pirta, R. S., P. Prakash, and M. Singh. 1981. A population study of two species of non-
human primates: Macaca mulatto and Macaca radiata. J. Bombay Nat. Hist. Soc, 77 (3):
429-434.

Rahaman, H., and M. D. Parthasarathy. 1979. Behavioral variants of bonnet macaque
(Macaca radiata) inhabiting cultivated gardens. J. Bombay Nat. Hist. Soc, 75 (2): 406-425.

Macaca radiata radiata (E. Geoffroy, 1812)

Cercocebus radiatus: E. Geoffroy, 1812, p. 98 — external characters; cranial comparisons
with M. silenus; distribution, India; specific name is an adjective referring to the radiate
arrangement of crown hairs.

[P]ithecus radiatus: Desmarest, 1817, p. 325 — external characters; taxonomic history; dis-
tribution, India.

[M]acacus radiatus: Desmarest, 1820, p. 64 — taxonomy; distribution, India.

Cercopithecus radiatus: Kuhl, 1820, p. 13 — external characters; distribution, eastern India.

S[imia] M[acacus] radiatus: Griffith, 1827, p. 17 — external characters; taxonomy; distri-
bution, India.

S[imia] radiata: G. Cuvier, 1829, p. 95 — external comparison with M. sinica.

I[nuus] radiatus: Wagner, [1839], p. 140 — external characters; distribution, Malabar Coast.

M[acacus] radiatus (affinis): Blyth, 1863, p. 8 — distribution, "Formosa."

M[acaca] radiata: Hinton and Wroughton, 1921, p. 815 — nomenclature.

Z[ati; genus or subgenus] radiata: Pocock, [1926], p. 1572 — taxonomy; distribution, south-
ern India.

M[acaca] radiata radiata: Pocock, 1931b, p. 276 — part, external, and cranial characters;
taxonomy.

Macaca (Zati) radiata: Hill, 1932, p. 314 — external characters. Hill, 1974, pp. xiv, xv, 694 —
taxonomy.

Toque: F. Cuvier, 1820, p. 1 (part), plate (animal, color) — taxonomy; penial characters;
distribution, Malabar Coast.

Simia sinica . . . var.: Griffith, 1821, p. 110 (not Linnaeus, 1771) — taxonomy.

M[acacus] sinicus: I. Geoffroy, 1851, p. 26 (not L.) — taxonomic history; type history;
distribution, India, Mauritius ("vraisemblablement importee").

l[nuus] sinicus: Wagner, 1855, p. 56 (not L.) — external characters; distribution, peninsular
India.

Pith[ecus] (Mac[acus\) sinicus: Dahlbom, 1856, p. 117 (not L.) — external characters; dis-
tribution, India.

[Cynamolgus] Z[ati; subgenus] sinicus: Reichenbach, 1862, p. 130, pi. 23 (figs. 27-29;
animals in color) (not L.) — taxonomic history; distribution, India, Mauritius (intro-
duced).

Cerococebus sinicus: Schlegel, 1876, p. 99 (not L.) — part, external characters; distribution,
Deccan.

Macacus (sinicus), affinis: Anderson, 1879, p. 91 (not L.) — doubtful species; distribution,
Formosa (probably introduced).

Simia sinica: Wroughton, 1912, p. 1176 (not L.) — locality records; vernacular names; field
notes. Hinton and Wroughton, 1921, p. 813 — name not properly applicable to this
species.

Pithecus sinicus: D. G. Elliot, [1913], p. 221 (not L.) — external and cranial characters;
distribution, southern India.

M[acaca] sinica: Wroughton, 1918, p. 555 (not L.) — external characters; locality records.

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA 35

Zati sinicus: Pocock, 1921, p. 227, figs, g-k (penis) (not L.) — penial comparisons; tax-
onomy.

Silenus . . . sinicus: Stiles and Nolan, 1929, p. 535 (not L.) — parasites.

Macacus pileatus: Newton, 1893, p. 217 (not Kerr, 1792, p. 69)— introduced in Mauritius.

Pithecus albibarbatus: D. G. Elliot, [1913], cranial pi. xxiv (skull) (not Kerr, 1792)— mis-
identification (see Pocock, 1939, p. 68).

Types. — Of the type series examined by E. Geoffroy (1812, p. 98), two speci-
mens have been preserved. The lectotype, designated by I. Geoffroy (1851, p.
26), is a mounted skin (MNHN 353/223, Type Cat. No. 56a; skull inside) of an
adult male that was presented alive to the Paris Menagerie by Mme. Regnault
de Saint Jean d'Angely in 1801; the monkey died in captivity in December 1803
or January 1804 (Nivose, an XII, French Republican calendar). The paralectotype
is a mounted skin (MNHN 356/226, Type Cat. No. 56b; no skull) of a captive
immature male that died in the Paris Menagerie in 1815 (cf. Rode, 1938, p. 222);
when examined by E. Geoffroy (1812, p. 98), this monkey evidently was still
alive. Another mounted specimen (MNHN 355/227) in the Paris Museum also
is tagged "Type," but this is the skin of a juvenile that died in captivity in 1837,
and therefore it obviously was not available to E. Geoffroy (1812, p. 98) when
he described this species.

Type-locality. — Peninsular India. E. Geoffroy's (1812, p. 98) statement concern-
ing the provenance of this species is "Habite l'lnde," which is the locality notation
on the tag of the lectotype (MNHN 353/223). The spurious notation "Bengale"
is written on the underside of the stand of the paralectotype (MNHN 356/226).

Distribution (fig. 1). — Peninsular India from the northern limit of the species
(above, p. 2) southward as far as the Palni Hills and the Shevaroy Hills.

Diagnostic pelage color characters (prime pelage). — Crown grayish brown, cap hairs
frequently tipped with blackish; upper back drab grayish brown, variably
washed with pale yellowish, hair banding relatively inconspicuous; lower back
with a variably defined dark brown to blackish median streak; outer surface of
arms and legs ochraceous gray; skin of ventral surface broadly and deeply pig-
mented dark bluish gray.

Specimens examined. — Total, 122.

Skins and skulls, 53— Localities Nos. 10 (3), 12(2), 14 (1), 15 (6), 19 (2), 26 (3), 33 (4), 40
(2), 43 (1), 45 (1), 59 (2), 62 (1), 63 (2), 64 (1), 66 (2), 71 (1), 72 (3), "Ceylon" (2: ANSP, 1;
USNM, 1), "Formosa" (1), "Madras" (1), "Southeastern Asia," (2: AMNH), "Travancore"
(1), no locality data (9: FMNH, 1; IRSN, 1; MNHN, 1; NHMB, 1; RMNH, 5).

Skins only, 26— Localities Nos. 7 (2), 10 (1), 38 (1), 63 (1), "Bengale" (1), "Decan" (4,
including 2 with skull inside), "India" or equivalent (4: IRSN, 1, skull inside; MNHN, 2,
including 1 with skull inside; ZSBS, 1), "Malaya Peninsula" (1: AMNH), "le Maurice" (1,
skull inside), Reunion (1), no locality data (9: IRSN, 1; MNHN, 1; RMNH, 5; ZMUZ, 1;
ZSI, 1).

Skulls only, 43— Localities Nos. 1 (1), 2 (2), 57 (1), "India" (2: BM, 1; NHRM, 1), no
locality data (37: AIUZ, 17; BM, 10; FMNH, 1; IRSN, 3; USNM, 1; ZSI, 5, including 1
mandible only).

For details, except where museum is specified here, see Figure 1 and Gazetteer.

Macaca radiata diluta Pocock, 1931

Cercocebus sinicus: Schlegel, 1876, p. 100 (not Linnaeus, 1771)— part, locality record,

Pondicherry.
Macaca sinica: Wroughton, 1921, pp. 546, 549 (not L.)— locality records, Kambam, Bhu-

tapandi.

36 FIELDIANA: ZOOLOGY

M[acaca) radiata radiata: Pocock, 1931b, p. 278 — part, external characters of specimen
collected at "Cumbum" [Kambam].

M[acaca] radiata diluta Pocock, 1931b, p. 278 — external characters, distribution, "Booth-
apaundy" [Bhutapandi]; subspecific name is an adjective alluding to the coloration,
which is characterized as paler than in the nominate subspecies.

Macaca (Zati) radiata diluta: Hill, 1937, pp. 205, 212 — external characters; field notes.

Types. — The holotype designated by Pocock (1931b, p. 279) is the stuffed skin,
with skull, of an adult female (BM 1937.5.26.1, Coll. No. 158), collected at Bhu-
tapandi by R. S. N. Pillay, 18 Nov. 1915 (cf. Wroughton, 1921, p. 549). This
specimen, however, is not currently tagged as type; instead, the paratype (BM
30.11.1.31, Coll. No. 171, adult female, skin and skull, same locality and collector,
20 Nov. 1915) erroneously is so tagged (cf. Hill, 1974, p. 708). The basis for
determination of the authentic holotype (Int. Code Zool. Nomencl., Art 73b) is
explained below.

Pocock (1931b, p. 278) based this subspecies solely on two adult female spec-
imens (table 15). Although no specimen numbers are mentioned, each of the
two specimens is identifiable by its recorded external measurements, which are
specified by Pocock. (Following his standard practice, Pocock converted mea-
surements recorded by the collector in millimeters to inches by dividing by 25.)
Of the two specimens, the one with the smaller measurements (BM 1937.5.26.1)
is designated by Pocock (p. 279, table) as "Type of diluta." This same type
designation ("the smaller specimen, the type") is repeated by Pocock (1939, p.
42) in a subsequent account of M. r. diluta, thereby unequivocally confirming
his original selection. These two published designations of the smaller specimen
as holotype clearly establish that tagging of the larger specimen as "Type," which
probably was done by Pocock himself, was an unfortunate curatorial lapsus.

Type-locality. — Bhutapandi, Tamil Nadu (state), extreme southern India; cited
by Pocock (1931b, p. 278) as "Boothapaundy in Travancore."

Distribution (fig. 1). — Southeastern peninsular India; known northern marginal
records of this subspecies are Alleppey (few miles S; ca. 9°25'N, 76°20'E), Kam-
bam (9°44'N, 77°18'E), and Pondicherry (11°56'N, 79°49'E).

Table 15. External measurements of type-series of Macaca radiata diluta Pocock, 1931.

_ Length

Source _

of data Head and body Tail

Pocock, 1931b, p. 279:

"Type of diluta," $ 1' 1 1/5"1 [330 mm]2 1' 9" [525 mm]2

Paratype, $ 1' 4 4/5" [420 mm]2 1' 10 4/5" [570 mm]2

Specimen tags:

BM 1937.5.26. I3 ? 328 mm4 524 mm

BM 30.11. 1.31s $ 420 mm 571 mm

'This is the measurement given in Pocock's (1931b, p. 279) table; the slightly different
measurement, 1' 1 3/5", given in the text on the same page evidently is the result of a
typographical error.

Pocock's conversion factor: 1 inch = 25 mm.

Collector's No. 158, collected by R. S. N. Pillay at "Bhoothapaundi, Travancore," 18
Nov. 1915.

4This measurement, which was taken and recorded by the collector, probably is in error
(see table 1); however, the error was not noticed by Pocock and consequently does not
affect his type designation. In any case, measured as dry skins, BM 1937.5.26.1 (HB =
510 mm) is clearly smaller than BM 30.11.1.31 (HB = 565 mm).

Collector's No. 171, collected by R. S. N. Pillay at "Bhoothapaundy, Travancore," 20
Nov. 1915.

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA 37

Diagnostic pelage color characters (prime pelage). — Crown pale yellowish brown;
upper back brightly colored, pale yellowish brown to golden brown, hairs con-
spicuously banded dark brown basally and bright yellowish distally; lower back
slightly drabber than upper back; outer surface of arms and legs pale grayish
brown; skin of ventral surface mostly unpigmented whitish in four specimens
examined (Bhutapandi, BM, 2; Kambam, BM, 1; Travancore, ZSI, 1; not ex-
amined in Pondicherry specimen, RMNH).

Specimens examined. — Total, 6.

Skins and skulls, 3— Locality Nos. 77 (1), 87 (2).

Skins only, 3— Locality Nos. 73 (1, skull inside), 82 (1), "India" (1: BM).

For details, except where museum is specified here, see Figure 1 and Gazetteer.

GAZETTEER OF MACACA RAD1ATA LOCALITIES

Locality names listed as primary entries in this gazetteer preferentially are the
standard names approved in U.S. Board on Geographic Names (USBGN) Gaz-
etteer: India (1952), with diacritical marks omitted. Primary entries for macaque
localities that are not included in USBGN: India are spelled here either as in U.S.
Defense Mapping Agency world map sheets, Series 1301, scale 1:1,000,000, or
as in the original source. Secondary entries, with cross references to correspond-
ing primary entries, give variant spellings or alternate locality names that appear
on specimen tags or in published literature on M. radiata.

The sequence of information presented in each primary entry is as follows:

1. locality name

2. altitude, if available

3. name of state (capitalized)

4. coordinates of locality, usually taken from USBGN: India

5. name of collector or observer followed by parenthetical reference to pub-
lished locality notes, if any

6. date of collection or observation

7. abbreviated name of museum where specimens are stored and number of
specimens available (with indication of part preserved, if skin and skull are
not both present) or bibliographic reference to literature record

8. italicized locality number as shown in distribution map (fig. 1)

Macaca radiata radiata

Anamalai Hills. See Anaimalai Hills.

Anaimalai Hills, N, 1,500-3,000 ft; KERALA; ca. 10°20'N, 76°50'E; observed by
C. R. Stonor (1944, p. 591), 12 Feb.-lO March 1944. 69

Avalgodu; KARNATAKA; not precisely located, ca. 14°20'N, 75°00'E; collected
by Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & An-
derson, 1971, p. 849), specimen not preserved. 20

Bairankuppe. See Byrankuppe.

Bandipur; KARNATAKA; 11°40'N, 76°38'E; observed by M. Krishnan (1972, p.

536), date unknown. 54
Bandipur, 1 mile NE, 3,000-3,500 ft; KARNATAKA; 11°40'N, 76°38'E; observed

by P. E. Simonds (1965, p. 178), Oct. 1961-June 1962. 54

38 FIELDIANA: ZOOLOGY

Bandipur, 1 mile W, 3,000-3,500 ft; KARNATAKA: 11°40'N, 76°38'E; observed
by P. E. Simonds (1965, p. 178), Oct. 1961-June 1962. 54

Bandipur National Park. See Bandipur Wildlife Sanctuary.

Bandipur Wildlife Sanctuary; KARNATAKA; ca. 11°40'N, 76°38'E; observed by
H. C. Sharatchandra and M. Gadgil (1976, p. 646; Nair et al., 1978, p. 420),
1974-75, and M. A. R. Khan (1977, p. 517), 1975-76. 54

Bangalore, 4,000 ft; KARNATAKA; ca. 13°00'N, 77°35'E; collected by Ripley In-
dian Expedition, 18 March 1947 (AMNH, 2); observed in Bangalore City by
A. S. Rajagopal (1965, p. 228), date unknown, and H. Rahaman and M. D.
Parthasarathy (1967, p. 252; 14 troops), Aug. 1964-Jan. 1965. 40

Bangalore-Krishnagiri road; KARNATAKA or TAMIL NADU; ca. 12°45'N,
77°45'E; several troops observed by A. S. Rajagopal (1965, p. 228), date un-
known. 41

Bellary. See Vijayanagar.

"Bengale"; ca. 22°-27°N, 84°-90°E; spurious locality notation (MNHN, 1 [skin
only]).

Biligiri Rangan Hills: KARNATAKA: 11°54'N, 77°14'E; collected by "H. C. Raven,
Hopwood Chindwin Exp.," 26 March 1935 (AMNH, 1 skin only). For basis of
doubt concerning identity of collector, see Morris, 1936, p. 668. 38

Billigirirangan Hills. See Biligiri Rangan Hills.

Bombay. See Yeur.

Byrankuppe, near; KARNATAKA; 11°52'N, 76°12'E; two troops observed by A.
Nolte (1955, p. 77; 1956, p. 177), April-May 1953. 47

Cauvery River. See Ranganathittoo Sanctuary.

Channapatna. See Malur.

Chica Nayakana Hully. See Chiknayakanhalli, vicinity.

Chiknayakanhalli, vicinity; KARNATAKA; 13°26'N, 76°37'E; sight record (iden-
tification equivocal), reported by F. Buchanan (1807, p. 55), 21 Aug. 1800. 32

Cochin. See Sharnelli Estate.

Coonoor-Mettupalaiyam road, milestone 12 [km]; TAMIL NADU; 11°19'N,
76°56'E; observed by P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 60

Coonoor-Mettupalaiyam road, milestone 17 [km]; TAMIL NADU; 11°18'N,
76°56'E; observed by P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 60

Coromandel coast; ANDHRA PRADESH; ca. 14°00'N, 80°10'E; collected by F.
Eydoux and L. F. A. Souleyet (1841, p. xiv), 5 April-12 June 1837 (MNHN,
2; not seen, reported by I. Geoffroy, 1851, p. 26). 28

Dakhan. See Deccan.

Decan. See Deccan.

Deccan; MAHARASHTRA, ANDHRA PRADESH and KARNATAKA; ca. 10°-15°N,

75°-80°E; reported by W H. Sykes (1831, p. 99); collector unknown, 1862-75

(RMNH, 4 [2 skulls in skins, 2 skins only], including at least two captives).

Not mapped.
Devikop, 2,000 ft; KARNATAKA; 15°08'N, 74°56'E; collected by G. C. Shortridge

(in Wroughton, 1912, p. 1176), 21 Nov. 1911 (BNHS, 1). 14
Dharwar, 2,300 ft; KARNATAKA; ca. 15°28'N, 75°02'E; collected by G. C. Short-

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA 39

ridge (in Wroughton, 1912, p. 1176), 14 Nov. 1911 (BM, 2; FMNH, 1; ZSI, 1
[skin only]). 10

Dharwar, 4.4-17.5 km SW; KARNATAKA; ca. 15°25'N, 74°55'E; nine open-land
troops observed by Y. Sugiyama (1972, p. 255), June-Sept. 1961; six troops
observed by N. Koyama (1973, p. 225), 2 Jan.-14 March 1963. 9

Dharwar, 18.4-29.5 km SW; KARNATAKA; ca. 15°20'N, 74°50'E; three forest
troops observed by Y. Sugiyama (1972, p. 255), June-Sept. 1961. 8

Dodderi; KARNATAKA; not precisely located, ca. 14°20'N, 75°00'E; collected by
Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Ander-
son, 1971, p. 849), specimen not preserved. 20

Dukhun. See Deccan.

Eastern Ghats, near Nellore; ANDHRA PRADESH; ca. 14°45'N, 79°10'E; spec-
imen examined by T. C. Jerdon (1867, p. 13), date unknown. 27

"Formosa"; TAIWAN; ca. 22°-25°N, 120°-122°E; living captive sent by R. Swin-
hoe, 1859 (ZSI, 1; see Blyth, 1860, p. 87, and 1875, p. 8). Not mapped.

Gadag; KARNATAKA; 15°25'N, 75°37'E; reported by G. C. Shortridge (in
Wroughton, 1912, p. 1176), Nov.-March 1911. 11

Gaday. See Gadag.

Gersoppa, sea level; KARNATAKA; 14°15'N, 74°39'E; collected by G. C. Short-
ridge (in Wroughton, 1913, p. 30), 27-29 May 1912 (BM, 2 [1 skin only]). 19

Ghatmatha; MAHARASHTRA; 17°25'N, 73°40'E (Moore & Tate, 1965, p. 320);
collected by S. H. Prater (in Wroughton, 1916, p. 311), 17 Dec. 1914 (BM, 1
[skin only]); collector unknown, 15 Dec. 1914 (BNHS, 1 [mounted specimen
on exhibit]). 7

Gudalur-Theppakadu, forest country between; TAMIL NADU; ca. 11°33'N,
76°32'E; collected by C. Donovan (1920, p. 718), May-June 1919 (specimens
not preserved). 55

Gundalapettu-Chamarajnagar. See Gundlupet-Chamrajnagar road.

Gundlupet-Chamrajnagar road, milestone 1, 2 [km]; KARNATAKA; 11°49'N,
76°41'E; two troops observed by P. E. Simonds (1974a, p. 157), 1961-63 or
1970. 52

Gudulur. See Gudalur-Theppakadu.

Haleri, 3,555 ft; KARNATAKA; ca. 12°27'N, 75°48'E; collected by G. C. Shortridge
(in Ryley, 1913b, p. 489), 20 Dec. 1912, 16 Jan. 1913 (BM, 4). 33

Halery. See Haleri.

Halgalali; KARNATAKA; 14°21'N, 75°06'E; (Work, 1958, p. 273); collected by
Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Ander-
son, 1971, p. 849) specimen not preserved. 17

Hampi. See Vijayanagar.

Hangala, 2,500-3,000 ft; KARNATAKA; 11°45'N, 76°39'E; observed by P. E. Si-
monds (1965, p. 178), Oct. 1961-June 1962. 53

Haravadike; KARNATAKA; not precisely located, ca. 14°20'N, 75°00'E; collected
by Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & An-
derson, 1971, p. 849), specimen not preserved. 20

Hosabale; KARNATAKA; 14°19'N, 75°02'E (Work, 1958, p. 272); collected by

40 FIELDIANA: ZOOLOGY

Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Ander-
son, 1971, p. 849), specimen not preserved. 20

Jalarpet; TAMIL NADU; 12°34'N, 78°35'E; observed by M. Krishnan (1972, pp.
509, 537), date unknown. 44

Kabbanir-Fluss. See Byrankuppe.

"Kachh" [Kutch]; GUJARAT; ca. 23°-24°N, 68°-72°E; imported captive observed
by F. Stoliczka (1872, p. 220), 1871-72. Not mapped.

Kakkenahala; KARNATAKA; 11°37'N, 76°35'E; observed by P. E. Simonds (1965,
p. 178), Oct. 1961-June 1962. 54

Kanara [district]. See Gersoppa.

Kanara Forest. See Devikop.

Karchikoppa; KARNATAKA; 14°18'N, 75°08'E (Work, 1958, p. 273); collected by
Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Ander-
son, 1971, p. 849), specimen not preserved. 18

Kargudi. See Theppakadu-Kargudi area.

Karjikoppa. See Karchikoppa.

Karumbapatti. See Kurumbapatti.

Karwar, forest; KARNATAKA; ca. 14°48'N, 74°08'E; observed by M. Krishnan
(1972, p. 536), date unknown. 13

Kauveri River. See Ranganathittoo Sanctuary.

Keladi; KARNATAKA; 14°13'N, 75°00'E (Work, 1958, p. 272); collected by Virus
Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Anderson, 1971,
p. 849), specimen not preserved. 21

Khandala, rocky cliffs and forests; MAHARASHTRA; 18°45'N, 73°23'E; reported
by C. McCann (1933, p. 810). 5

Kilkotagiri. See Kotagiri.

Kodaikanal. See Palni Hills, 5,500 ft.

Kolar, 2,786-4,026 ft; KARNATAKA; 13°08'N, 78°08'E; collected by G. C. Short-
ridge (in Ryley, 1913a, p. 283) 29 Sept. 1912 (BM, 1). 43

Konkan; MAHARASHTRA; ca. 18°05'N, 73°25'E; reported by R. M. Dixon (1894,
p. 104). 6

Kotagiri, 5,120 ft; TAMIL NADU; 11°26'N, 76°53'E; collected by P. Gosse (in
Wroughton & Davidson, 1920, p. 1031), 25 June 1918 (BM, 2). 59

Koyna Valley. See Ghatmatha.

Kummur. See Kummuru.

Kummuru; KARNATAKA; 14°25'N, 75°06'E (Work, 1958, p. 273); collected by
Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Ander-
son, 1971, p. 849), specimen not preserved. 17

Kuppe; KARNATAKA; not precisely located, ca. 14°20'N, 75°00'E; collected by
Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Ander-
son, 1971, p. 849), specimen not preserved. 20

Kurnool. See "Malakondapenta, Kurnool District."

Kurumbapatti; TAMIL NADU; 11°45'N, 78°15'E (Moore & Tate, 1965, p. 324);
collected by N. A. Baptista, 21 April 1929 (BM, 1). 62

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA 41

Kuruvari; KARNATAKA; not precisely located, ca. 14°20'N, 75°00'E; collected
by Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & An-
derson, 1971, p. 849), specimen not preserved. 20

Kyasanur Forest, 580-650 m; KARNATAKA; 14°17'N, 75°02'E; diseased specimen
reported by T. H. Work (1958, pp. 250, 272), 28-29 March 1957; 172 specimens
collected in general area by Virus Research Centre, Poona (Trapido et al.,
1964, pp. 765, 768, 769), 1957-61, examined for ticks, monkeys not preserved.
20

Machchur, Palni Hills, 4,000 ft; TAMIL NADU; 10°16'N, 77°35'E; (Survey of India,
1:126,720, Sheet 58 F/SE); collected by C. McCann (in Lindsay, 1926, p. 592),
5 April 1922 (ZSI, 1). 72

Machur. See Machchur.

Maddathoray. See Mattathur.

"Madras" [formerly a state]; ANDRHA PRADESH or TAMIL NADU; ca. 8°-20°N,
75°-80°E; collected by T. C. Jerdon, date unknown (BM, 1; possibly same
specimen as that reported by Jerdon [1867, p. 13] from Eastern Ghats, near
Nellore). Not mapped.

Malabar coast; KERALA; ca. 10°15'-12°20'N, 75°00'-76°00'E; two specimens col-
lected by M. Housard, date unknown (apparently not preserved; see F. Cuvier,
1820, p. 1). Not mapped.

"Malakondapenta, Kurnool District," probably equivalent to Maria Konda-Mar-
lapenta, Kurnool District (cf. Pocock, 1931a, p. 51); ANDHRA PRADESH;
15°48'N, 79°01'E (Survey of India, 1:126,720, Sheet 57 M/NW); collected by
N. A. Baptista, 14 May 1930 (BM, 3 [including 1 skull in BNHS, No. 5053]).
26

Malalagadde; KARNATAKA; 14°18'N, 75°05'E (Work, 1958, p. 272); collected by
Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Ander-
son, 1971, p. 849), specimen not preserved. 18

Malur, Channaparna [subdistrict]; KARNATAKA: 13°00'N, 77°55'E; observed by
A. S. Rajagopal (1965, p. 228), date unknown. 42

Manali Theppakadu, forest; TAMIL NADU; ca. 11°35'N, 76°34'E; observed by
P. E. Simonds (1973, p. 600), 1961-63 or 1970. 55

Mangata, 3,000-3,500 ft; KARNATAKA; 11°39'N, 76°40'E; observed by P. E. Si-
monds (1965, p. 178), Oct. 1961-June 1962. 54

Marasa; KARNATAKA; 14°12'N, 75°03'E (Work, 1958, p. 272); collected by Virus
Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Anderson, 1971,
p. 849), specimen not preserved. 22

Maria Konda. See "Malakondapenta, Kurnool District."

Marlapenta. See "Malakondapenta, Kurnool District."

Matheran; MAHARASHTRA: 18°59'N, 73°16'E; collected by H. M. Phipson,
April 1888 (BNHS, 1 [skull only]); collected by J. Banty, May 1892 (BNHS, 1
[skull only]). 4

Mattathur; KERALA; 10°23'N, 76°20'E (USDMA, series 1301, Sheet NC43) re-
ported as host of pseudoscorpion by E. Ellingsen (1914, p. 10), 17 Nov. 1908;
host of tick, Haemaphysalis bispinosa (Nuttall & Warburton, 1915, p. 432). 65

Maurice. See Mauritius.

42 FIELDIANA: ZOOLOGY

Mauritius; MASCARENE ISLANDS: 20°18'S, 57°36'E; M. r. radiata introduced

(see above, p. 2); presented by M. l'abbe Bascou, 1851 or earlier) (MNHN,

1 [skin only]). Not mapped.
Moyar River. See Manga ta.

Moyar Valley. See Nilgiri Hills, N, overlooking Moyar Valley.
Mudumalai Wild Life Sanctuary, 3,000-3,800 ft; TAMIL NADU; ca. 11°37'N,

76°31'E; reported by J. J. Spillett (1969, p. 640), 1965-66; observed by M.

Krishnan (1972, p. 536, pi. 4), 27 Sept. 1966, and M. A. R. Khan (1977, p.

517), 1974-76, and S. S. C. Nair et al. (1978, p. 420), July-Oct. 1975. 55
Mysore, Horticulture Department field; KARNATAKA; 12°18'N, 76°39'E; ob-
served by M. Singh and R. Sachdeva (1977, p. 605) and M. Singh and R. S.

Pirta (1978, p. 265), 1974-77, 48
Mysore-Bandipur. See Somanathapur, 1 mi. W.
Mysore-Bandipur Sanctuary; KARNATAKA; 12°18'N, 76°39'E-11°40'N, 76°38'E;

reported by J. J. Spillett (1968b, p. 318), 27-29 Nov. 1966. 48-53
Mysore-Bangalore road, milestone 28, 30, 32 [km]; KARNATAKA; 12°30'N,

76°50'E; three troops observed by P. E. Simonds (1974a, p. 157), 1961-63 or

1970. 36
Mysore-Bangalore road, milestone 34, 36 [km]; KARNATAKA; 12°32'N, 76°52'E;

two troops observed by P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 37
Mysore-Bangalore road, milestone 83 [km]; KARNATAKA; 12°44'N, 77°17'E;

observed by P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 39
Mysore-Mangalore road, milestone 94, 96 [km]; KARNATAKA; 12°23'N, 75°50'E;

two troops observed by P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 34

Nadahalli. See Nadhalli.

Nadhalli; KARNATAKA; 14°22'N, 75°04'E (Work, 1958, p. 272); collected by Virus

Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Anderson, 1971,

p. 849), specimen not preserved. 17
Nagarhole Wildlife Sanctuary; KARNATAKA; 12°05'N, 76°05'E (Spillett, 1968b,

map 2); observed by M. A. R. Khan (1977, p. 517), 1975-76. 46
Nagarjunakonda valley; ANDHRA PRADESH; 16°31'N, 79°14'E; erroneously

reported by R. Subrahmanyam and V. V Rao (1975, p. 6), who also erroneously

report the liontail macaque, "Innus silenus" ( = Macaca silenus), at the same

place (cf. Fooden et al., 1981, fig. 1). Not mapped.
Nelliampathy Hills, northern cliffs; KERALA; ca. 10°35'N, 76°45'E; reported by

A. P. Kinloch (1921, p. 939; 1923, p. 552), 1911-1923. 67
Nelliampathy Hills, southern slopes; KERALA; ca. 10°25'N, 76°45'E; reported

by A. P. Kinloch (1921, p. 939; 1923, p. 552), 1911-1923. 68
Nellore. See Eastern Ghats, near Nellore.
"Nepal"; NEPAL; ca. 27°-30°N, 80°-88°E; spurious report by B. H. Hodgson

(1834, p. 96), based on misidentified M. mulatta (cf. Hodgson, 1832, p. 339).

Not mapped.
Nilghiris. See Nilgiri Hills.
Nilgiris. See Nilgiri Hills.
Nilgiri HiUs; KERALA or TAMIL NADU; ca. 11°20'N, 76°30'E; collected by W.

Davison, date unknown (BM, 1 [skull only]). 57

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA 43

Nilgiri Hills, 5,000-8,600 ft; TAMIL NADU; ca. 11°29'N, 76°34'E; "seen occa-
sionally" by M. A. R. Khan (1977, p. 517), 1974-76. 56

Nilgiri Hills, 7,000 ft; TAMIL NADU; ca. 11°29'N, 76°34'E; observed by P. E.
Simonds (1965, pp. 175, 177), Oct. 1961-June 1962. 56

Nilgiri Hills, N, overlooking Moyar Valley, 4,000 ft; TAMIL NADU; ca. 11°30'N,
76°33'E; collector unknown, Dec. 1919-Jan. 1920 (specimen not preserved;
reported by C. Donovan, 1920, p. 721). 56

Nilgiri Hills, northern and eastern slopes, ca. 6,000 ft; TAMIL NADU; ca. 11°25'N,
76°55'E; observed by F. E. Poirier (1970, p. 260), Sept. 1965-Aug. 1966. 59

Nilgiri Hills, western slopes, 3,000-4,000 ft; TAMIL NADU; ca. 11°25'N, 76°30'E;
observed by F. E. Poirier (1970, pp. 258-260), Sept. 1965-Aug. 1966. 56

Nishrani; KARNATAKA; not precisely located, ca. 14°20'N, 75°00'E; collected by
Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Ander-
son, 1971, p. 849), specimen not preserved. 20

North Coorg. See Haleri.

Ootacamund-Mysore road, milestone 92 [km]; KARNATAKA; 12°04'N, 76°40'E;

observed by P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 51
Ootacamund-Mysore road, milestone 107, 108 [km]; KARNATAKA; 12°10'N,

76°40'E; two troops observed by P. E. Simonds (1974a, p. 157), 1961-63 or

1970. 50
Ootacamund-Mysore road, milestone 114 [km]; KARNATAKA; 12°15'N, 76°40'E;

observed by P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 49
Ootacamund-Mysore road, milestone 117 [km]; KARNATAKA; 12°17'N, 76°40'E;

observed by P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 48

Ootacamund-Mysore road, milestone 121 [km]; 12°18'N, 76°40'E; observed by
P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 48

Padamara, forest; TAMIL NADU; not precisely located, ca. 11°35'N, 76°35'E;

observed by P. E. Simonds (1973, p. 600), 1961-63 or 1970. 55
Palni Hills, 5,500 ft; TAMIL NADU; 10°16'N, 77°33'E; collected by C. McCann

(in Lindsay, 1926, p. 592), 22-23 March 1922 (BM, 1; BNHS, 1). 72
Palni Hills, northern slopes, 3,000 ft; TAMIL NADU; ca. 10°22'N, 77°35'E (Survey

of India, 1:126,720, Sheet 58 F/SE); collected by J. Riley O'Brien (in Lindsay,

1926, p. 592), 1921-22 (BM, 1). 71
Panniar, 1,000-1,250 m; KERALA; ca. 10°00'N, 77°00'E; observed by Y. Sugiyama

(1968, p. 289), 5 Jan.-27 Feb. 1963. 70
Perumel. See Palni Hills, 5,500 ft.

Ranganathittoo Sanctuary, Cauvery River; KARNATAKA; 12°25'N, 76°40'E (Spil-
lett, 1968b, p. 299); observed by M. Krishnan (1972, p. 536), date unknown.
35

Reunion; MASCARENE ISLANDS: 20°06'S, 55°36'E; M. r. radiata introduced (see
above, p. 2); collected by M. Pollen, 1865 (RMNH, 1 [skin only]). Not
mapped.

Sagar-Shimoga road; KARNATAKA; ca. 14°07'N, 75°09'E; collected by Virus
Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Anderson, 1971,
p. 849), specimen not preserved. 24

44 FIELDIANA: ZOOLOGY

Salem District. See Kurumbapatti.

Salsette Island, hills; MAHARASHTRA; ca. 19°12'N, 72°54'E; observed by C.

McCann (1933, p. 810), date unknown. 2
Samasgi, 2,000 ft; KARNATAKA; 14°40'N, 75°00'E (Moore & Tate, 1965, p. 320);

collected by G. C. Shortridge (in Wroughton, 1912, p. 1174), 12 March 1912

(BM, 3; BNHS, 2; FMNH, 1). 15
Satara. See Ghatmatha.
Segur Range. See Sigur Range.
Sharnelli Estate, 1,500 ft; KERALA; 10°32'N, 76°40'E (Survey of India, 1:126,720,

Sheet 58 B/NE); collected by J. Riley O'Brien (in Lindsay, 1926, p. 592), 28 May,

6 June 1921 (BM, 1; BNHS, 1). 66
Shernelly. See Sharnelli Estate.
Shevaroy Hills, 4,500 ft; 5,400 ft; TAMIL NADU; ca. 11°50'N, 78°17'E; collected

by N. A. Baptista, 19-25 May 1929 (BM, 1; BNHS, 1; ZSI, 1 [skin only]). 63
Shigga; KARNATAKA; 14°19'N, 75°10'E; diseased specimen reported by T. H.

Work (1958, pp. 250, 273), 28-29 March 1957. 18
Shimoga District. See Kyasanur Forest.
Sholinghur, temple; TAMIL NADU; 13°07'N, 79°25'E; observed by M. Krishnan

(1972, p. 536, pi. 3), 24 May 1960. 31
Siddharkoil. see Siddharkovil.

Siddharkovil; TAMIL NADU; 11°37'N, 78°04'E (Imp. Gaz. India, 14:401); ob-
served by M. Krishnan (1972, p. 509), date unknown. 61
Sigur Range; TAMIL NADU; ca. 11°30'N, 76°42'E (USDMA, Series 1301, Sheet

NC43); observed by M. A. R. Khan (1977, p. 517), 1975-76. 58
Somanathapur, 1 mile W, 2,500-3,000 ft; KARNATAKA; 11°43'N, 76°40'E; ob-
served by E. P. Gee (1964, p. 20, pi. 8b) and P. E. Simonds (1965, p. 178), Oct.

1961-June 1962. 53
Sorab; KARNATAKA; 14°23'N, 75°06'E; collected by Virus Research Centre,

Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Anderson, 1971, p. 849), speci-
men not preserved. 17
Sorab-Banavasi road; KARNATAKA; ca. 14°28'N, 75°04'E; collected by Virus

Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Anderson, 1971,

p. 849), specimen not preserved. 16
Sorab-Shiralkoppa road; KARNATAKA; ca. 14°23'N, 75°10'E (Work, 1958, p. 273):

collected by Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan

& Anderson, 1971, p. 849), specimen not preserved. 17
South Arcot. See "Tenmali, S. Arcot."
"Southern Mahratta Country"; MAHARASHTRA and KARNATAKA; ca. 14°-19°N,

74°-77°E; reported by W Elliot (1839, pp. 94, 95). Not mapped.
Srisailam; ANDHRA PRADESH; 16°05'N, 78°52'E; collected by Zoological Survey

of India, 11 Feb. 1970 (ZSI, 1; not seen, reported by Agrawal & Bhattacharyya,

1976, p. 213). 25

Tada; ANDHRA PRADESH; 13°35'N, 80°02'E; observed by G. U. Kurup and J.

Fooden, 24 May 1973. 30
Tavanadi; KARNATAKA; not precisely located, ca. 14°20'N, 75°00'E; collected

FOODEN: INDIAN BONNET MACAQUE, MACACA RADIATA 45

by Virus Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & An-
derson, 1971, p. 849), specimen not preserved. 20
"Tenmali, S. Arcot," possibly equivalent to Timmalai, South Arcot District;

TAMIL NADU; ?11°44'N, 79°08'E (Survey of India, 1:126,720, Sheet 58M/NW);

collected by W. P. Howell and museum collector, date unknown (ZSI, 1 [skin,

No. 12007; skull, No. 7089]). 64
Teppikadu. See Gudalur-Theppakadu.
Thana. See Yeur.
Theppakadu-Kargudi area; TAMIL NADU; ca. 11°35'N, 76°35'E (Spillett, 1969,

p. 663); observed by M. Krishnan (1972, p. 537, pi. 3), 21 Sept. 1970; P. E.

Simonds (1973, p. 600), 1961-63 or 1970. 55
Theppakkadu. See Theppakadu-Kargudi area.
Timmalai. See "Tenmali, S. Arcot."
Tirupathi. See Tirupati.
Tirupati; ANDHRA PRADESH; 13°39'N, 79°25'E; observed by A. S. Rajagopal

(1965, p. 228) and M. Krishnan (1972, pp. 509, 537), dates unknown. 29
"Travancore" [formerly a state]; KERALA; specimen probably from Regent's

Park Zoo (Hill, 1937, p. 212), obtained from Mr. Bartlett, date unknown (BM,

1). Not mapped.
Trombay Hills; MAHARASHTRA; 19°02'N, 72°56'E; reported by C. McCann

(1933, p. 810). 3

Veduru Guta, forest; TAMIL NADU; not precisely located, ca. 11°35'N, 76°35'E;
observed by P. E. Simonds (1973, p. 600), 1961-63 or 1970. 55

Venu Gopal Wild Life Park. See Mysore-Bandipur Sanctuary.

Vijayanagar, 1,500 ft; KARNATAKA; 15°19'N, 76°28'E; collected by G. C. Short-
ridge (in Wroughton & Ryley, 1913, p. 60), 4 Aug. 1912 (BM, 2). 12

"Western Ghauts." See Deccan.

Wotekolli, 2,000 ft; KARNATAKA; ca. 12°08'N, 75°47'E; collected by G. C. Short-
ridge (in Ryley, 1913b, p. 489), 27 Dec. 1912 (BM, 1). 45
Wottekolle. See Wotekolli.

Yalgalali; KARNATAKA; 14°08'N, 75°02'E (Work, 1958, p. 272); collected by Virus
Research Centre, Poona, Nov. 1965-Nov. 1967 (Rajagopalan & Anderson, 1971,
p. 849), specimen not preserved. 23

Yelagalale. See Yalgalali.

Yeur, jungle near; MAHARASHTRA; 19°14'N, 72°57'E (Survey of India, 1:63,
360, Sheet 47 A/16); collected by R. E. Hawkins, 10 Aug. 1958 (BNHS, 1 [skull
only], picked up along roadside; collector's statement provided by H. Abdulali,
letter, 23 May 1978). 1

Macaca radiata diluta

Alleppey, temple few miles S, on road to Quilon; KERALA; ca. 9°25'N, 76°20'E;

observed by W. C. O. Hill (1937, p. 212), June 1936. 78
Aramboli, temple 5 miles S; TAMIL NADU; 8°13'N, 77°34'E; observed by W C.

O. Hill (1937, p. 212), June 1936. 88

46 FIELDIANA: ZOOLOGY

Ashamboo Hills. See Papanasam.
Ashambu Hills. See Papanasam.

Bhoothapaundy. See Bhutapandi.

Bhutapandi; TAMIL NADU; 8°16'N, 77°26'E; collected by R. S. N. Pillay (in

Wroughton, 1921, p. 549), 18, 20 Nov. 1915 (BM, 2). 87
Boothaundy. See Bhutapandi.

Courtallam. See Kuttalam.
Cumbum. See Kambam.
Cumbun. See Kambam.

Kambam; TAMIL NADU; 9°44'N, 77°18'E; collected by S. H. Prater (in Wrough-
ton, 1921, p. 546), 30 May 1917 (BM, 1). 77

Kodaicanal Road, milestone 23. See Kodaikanal Road, milestone 23 [km].

Kodaikanal Road; TAMIL NADU; 10°11'N, 77°56'E; observed by M. Krishnan
(1972, p. 509), date unknown. 75

Kodaikanal Road, milestone 23 [km]; TAMIL NADU; ca. 10°10'N, 77°40'E; ob-
served by P. E. Simonds (1974a, p. 157), 1961-63 or 1970. 76

Kodikadu Reserved Forest; TAMIL NADU; 10°17'N, 79°46'E; M. radiata intro-
duced from Kambakonam vicinity, April 1965 (Daniel, 1968, p. 519; Krishnan,
1972, pp. 516, 538). Not mapped.

Kumbakonam, near; TAMIL NADU; ca. 10°58'N, 79°23'E; collector unknown,
April 1965, ca. 30 living specimens obtained and subsequently transported to
Kodikadu Reserved Forest (Krishnan, 1972, p. 538). 74

Kuttalam, forest; TAMIL NADU; ca. 8°55'N, 77°17'E; observed by K. Krishna-
moorthy and M. Krishnan (1972, pp. 509, 536, 539), April 1967 and other dates
unspecified. 84

Madura. See Kambam.

Naraikkadu Estate, vicinity; TAMIL NADU; ca. 8°30'N, 77°30'E; reported by
C. G. Webb-Peploe (1947, p. 630). 86

Papanasam; TAMIL NADU; 8°42'N, 77°23'E; observed by M. Krishnan (1972,

pp. 509, 537), date unknown; observed by H. Abdulali (letters, 23 May 1978,

26 June 1978) and R. Ali, April 1978. 85
Papavinasam. See Papanasam.
Periyar Lake, N; KERALA; ca. 9°34'N, 77°10'E; reported by J. Tanaka (1965, p.

Ill), 7 Jan.-5 Feb. 1963. 79
Periyar Wildlife Sanctuary; KERALA; ca. 9°28'N, 77°04'E; reported by G. U.

Kurup (1971, p. 17). 79
Point Calimere Sanctuary. See Kodikadu Reserved Forest.
Pondicherry; PONDICHERRY; ca. 11°56'N, 79°49'E; collector and date unknown

(RMNH, 1 [skull in skin]). 73
Pondichery. See Pondicherry.

Quilon. See Alleppey.

Srivilliputhur. See Srivilliputtur-Mudaliar Ootu.

FOODEN: INDIAN BONNET MACAQUE, MACACA RAD1ATA 47

Srivilliputtur-Mudaliar Ootu, hill track between; TAMIL NADU; ca. 9°35'N,
77°34'E; reported by A. J. T. John Singh (1974, p. 376). 81

Travancore; KERALA; ca. 9°00'N, 77°00'E; collected by Dr. N. Annandale, 28

Nov. 1908 (ZSI, 1 [skin only]). 82
Trivandrum, vicinity; KERALA; ca. 8°29'N, 76°55'E; locally obtained living and

preserved specimens examined in Trivandrum Zoo and Trivandrum Museum

by W. C. O. Hill (1937, p. 212), June 1936. 83

Varushnaad Valley, ca. 5,000 ft (one observation, more common at lower ele-
vations); TAMIL NADU; ca. 9°35'N, 77°30'E; reported by A. F. Hutton (1949,
p. 689). 80

Vedaranyam. See Kodikadu Reserved Forest.

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