UNr RSITY OF
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NATURAL HIST. SURVEY

FIELDIANA ■»■«

Zoology aug 6 1985

Published by Field Museum of Natural History

New Series, No. 17

tffiOT

TAXONOMY AND EVOLUTION

OF THE S/MC4 GROUP OF MACAQUES:

4. SPECIES ACCOUNT OF MACACA THIBETANA

JACK FOODEN

April 29, 1983
Publication 1345

TAXONOMY AND EVOLUTION

OF THE SINICA GROUP OF MACAQUES:

4. SPECIES ACCOUNT OF MACACA THIBETAN A

FIELDIANA
Zoology

Published by Field Museum of Natural History

New Series, No. 17

TAXONOMY AND EVOLUTION

OF THE SINICA GROUP OF MACAQUES:

4. SPECIES ACCOUNT OF MACACA THIBETAN A

JACK FOODEN

Research Associate

Division of Mammals

Department of Zoology

Field Museum of Natural History

Professor of Zoology
Chicago State University

Accepted for publication October 28, 1982
April 29, 1983
Publication 1345

Library of Congress Catalog Card Number: 82-84381

ISSN 0015-0754

PRINTED IN THE UNITED STATES OF AMERICA

CONTENTS

Introduction 1

Species Account 2

Synonymy 2

Types 3

Type-locality 3

Distribution 5

External measurements 5

Pelage 7

Cranial characters 8

Natural history 11

Specimens examined 14

Gazetteer 14

Literature Cited 17

LIST OF ILLUSTRATIONS

1. Macaca thibetana, lectotype skin and skull 4

2. Macaca thibetana, distribution of known locality records 6

3. Pelage variation in Macaca thibetana, adult males 7

4. Cranial morphology in Macaca thibetana 9

5. Allometry of rostral length vs. postrostral length in 20 immature and adult speci-
mens of Macaca thibetana 12

LIST OF TABLES

1. External measurements recorded by collectors for 10 specimens of Macaca

thibetana 5

2. Cranial measurements and ratios in 25 specimens of Macaca thibetana 10

3. Field notes recorded by collectors concerning Macaca thibetana 13

INTRODUCTION

Macaca thibetana is one of two species of stumptail macaques that are native
to China; the other is M. arctoides ( = M. speciosa of many authors; Fooden, [1966],
p. 153). These two macaques differ strikingly in form and size of the glans penis
and baculum, which are about half as long in M. thibetana as in M. arctoides, and
presumably also in structure of the female reproductive tract (Fooden, 1967, p.
939; 1971b, p. 72). Vernacular names proposed for these species are "Pere David's
stumptail macaque" for M. thibetana, in reference to the collector of the type-
specimen, and "bear macaque" for M. arctoides (Fooden, 1976, p. 226). Despite
conspicuous differences between M. thibetana and M. arctoides, they frequently
are confused and often have erroneously been assumed to be conspecific because
both species are large, stumptailed, and generally brownish in dorsal pelage
color.

Based on reproductive anatomy, M. thibetana is assigned to the sinica group
of macaques (Fooden, 1971b, p. 72), which also includes M. sinica, M. radiata,
and M. assamensis. The tail length morphocline and pattern of geographic dis-
tribution in this group suggests that M. thibetana is the most derived of the four
species. Accounts of the other three species in the sinica group have been pub-
lished recently (Fooden, 1979, p. 109; 1981, p. 1; 1982, p. 1). A comprehensive
comparative overview of the group is in preparation.

The present account of M. thibetana is based on a study of 29 available museum
specimens and review of relevant literature. The most important previous works
on the taxonomy of this species are those of Milne-Edwards ([1872], p. 244),
Matschie (1912, p. 305), Allen (1930, p. 3; 1938, p. 289), Dobroruka (1967, p.
305) and Hill (1974, pp. 760, 763). Specimens examined by me are preserved in
the following institutions, whose officials and staffs I thank for their generous
cooperation:

AMNH American Museum of Natural History, New York (7 spec-

imens)

FMNH Field Museum of Natural History, Chicago (4 specimens)

MCZ Museum of Comparative Zoology, Harvard University,

Cambridge (1 specimen)

MNHN Museum National d'Histoire Naturelle (Mammiferes), Paris

(5 specimens)

USNM U.S. National Museum of Natural History, Washington,

D.C. (11 specimens)

ZMB Zoologisches Museum der Humboldt-Universitat, Berlin

(1 specimen)

2 FIELDIANA: ZOOLOGY

Only 13 localities are recorded for M. thibetana, and little is known of its natural
history. Interest in wildlife and conservation, however, is growing in China
(Zhang et al., 1981, p. 223), and we may therefore anticipate further research
that ultimately will lead to better understanding of this unique, rare, and intrigu-
ing element of the Chinese primate fauna.

SPECIES ACCOUNT
Macaca thibetana A. Milne-Edwards, 1870

singe d'assez grande taille: David, 1869, p. 11 — announcement of discovery; assigned to
genus Macacus ( = Macaca); inhabits cold mountainous region, "principaut£ thibetaine
. . . de Mou-pin."

M[acacus] Thibetanus: A. Milne-Edwards, 1870, p. 341 — external characters, taxonomic com-
parisons; provenance, "Thibet oriental." A. Milne-Edwards, 1892, p. 671 — distribution,
"la vallee de Moupin."

Macacus thibetanus: Blanchard, 1871, p. 619 — external characters and habitat of specimen
reported by David (1869, p. 11).

[Macacus (Macacus)] thibetanus: Trouessart, 1878, p. 126 — distribution, Mouping.

M[acaca] thibetanum: Pocock, [1926], p. 1571 — tentatively allocated to Lyssodes group. Rode,
1938, p. 223 — type-specimens catalogued.

Macaca thibetana: Fooden, [1966], p. 160, fig. 3 (bacula) — external characters; bacular anat-
omy; taxonomic comparisons; zoogeography. Fooden, 1971a, p. 38 — taxonomic com-
parison with M. arctoides. Fooden, 1971b, p. 72, fig. 5 (glans penis) — penial anatomy;
external measurements ex Allen, 1938, p. 292; taxonomic comparisons.

Pithecus thibetanum: Elliot, [1913], p. 196, cranial pi. 21 (skull) — external and cranial char-
acters of type-specimen.

Pithecus thibetanus: Howell, 1929, p. 34 — external characters of three specimens; locality,
Emei Shan: Si Gi Pin.

Lyssodes speciosus thibetanus: Allen, 1938, p. 290 — external and cranial characters ex litera-
ture; taxonomy; distribution, "western Yunnan northwestward into central Szechwan."
Sowerby, 1941, p. 263, fig. opp. (photo of adult and young) — external characters; dis-
tribution, "Moupin . . . Mount Omei ... no specimens . . . from Tibet Proper."

Lyssodes s[peciosa] thibetanus: Tate, 1947, p. 135 — distibution, "Tibet and Szechwan."

Macaca speciosa thibetana: Kellogg, 1945, p. 124 — distribution ex Allen, 1938, p. 291. Bertrand,
1969, p. 5 — taxonomy; distribution, "high mountains of Szechwan."

[Macaca speciosa] thibetanus: Buettner-Janusch, 1963, p. 52 — listed.

Macaca (lyssodes) speciosa thibetana: Dobroruka, 1967, p. 312, fig. 1 (photo of female), fig.
2 (photo of head of male), fig. 5 (map of distribution) — external characters; taxonomic
comparisons and history; natural history; distribution, "Von Nan-Ling-Gebirge nach
Szetschwan."

Macacus tibetanus: A. Milne-Edwards, [1872], p. 244, pi. 34 (animal, in color), pi. 35 (skull) —
unjustified emendation; external and cranial characters of two specimens collected by
David (1869, p. 11); natural history; taxonomic comparisons; measurements; provenance,
'Tibet oriental . . . principautd de Moupin." David, 1874, p. 9 — details of collection of
type-specimen. David, 1875, p. 267 — first record in Fujian: "Koaten."

[Macacus (Macacus) arctoides] tibetanus: Trouessart, 1897, p. 27 — distribution, Mouping.

Macacus arctoides tibetanus: Lydekker, [1907], p. 997 — zoogeography.

Macaca arctoides: Forbes, 1894, p. 8 (not. I. Geoffroy, 1831, p. 61)— part, included in syn-
onymy.

Macacus (Magus) arctoides esau: Matschie, 1912, pp. 305, 309 — holotype, adult male, skin
and skull (ZMB 16179), collected alive at "Yao-tze Berge" W of Lo-ch'ang (border area
between Guangdong, Hunan, and Guangxi) by R. Mell some time before 9 Feb. 1911,
transported to Berlin Zoological Garden, died in captivity 16 March 1912; subspecific
name based on proper name applied to captive monkey; external and cranial characters;
taxonomic comparisons. Allen, 1930, p. 4 — regarded as a synonym of Macacus (Magus)
arctoides melli Matschie, 1912. Fooden, [1966], p. 160 — regarded as a synonym of Macaca
thibetana Milne-Edwards, 1870. Dobroruka, 1967, p. 312 — regarded as a synonym of
Macaca thibetana Milne-Edwards, 1870.

FOODEN: MACACA TH1BETANA 3

Simia arctoides esau: Heck, 1916, p. 547, pi. "Affen III," fig. 9 (captive, photograph; type-
specimen according to Dobroruka, 1967, p. 309, footnote 2) — natural history ex Matschie
(1912, p. 309). Petzold, 1964, p. 330, fig. 21 (captives, photograph) — tentative identifi-
cation of zoo specimens.

Macacus arctoides esau: Mell, 1922, pp. 4, 10, pi. 1, fig. 2 (type-specimen photographed
alive), fig. 3 (referred specimen photographed alive) — type history; temperament in
captivity.

Pithecus pullus: Howell, 1928, p. 41 — holotype, subadult male, skin and skull (USNM
252157), collected at Kuatun by F. T. Smith (Coll. No. 850), 16 May-24 June 1926; external
and cranial characters; taxonomic comparisons. Howell, 1929, p. 34, pi. 5 (skull of type) —
pelage comparison with Pithecus thibetanus. Allen, 1930, p. 4— said to be a synonym of
Macacus (Magus) arctoides melli Matschie, 1912.

Lyssodes speciosus melli: Allen, 1930, p. 3 (not Matschie, 1912, p. 308) — misidentification of
specimens collected at Chong'an Xian near Kuatun; external characters; taxonomy. Allen,
1938, p. 291 (not Matschie, 1912, p. 308) — external and cranial characters; measurements;
taxonomy; natural history; distribution, "northwestern Fukien southward along the
coast region of China, . . . Kwangtung and Kwangsi."

Macaca speciosa melli: Kellogg, 1945, p. 124 (not Matschie, 1912, p. 308) — part, Macacus
(Magus) arctoides esau Matschie, 1912, and Pithecus pullus Howell, 1928, included in
synonymy; distribution ex Allen, 1938, p. 292.

Macaca (Lyssodes) speciosa melli: Dobroruka, 1967, p. 312 (not Matschie, 1912, p. 308) — part,
Pithecus pullus Howell, 1928, included in synonymy and distribution.

Macaca arctoides melli: Hill, 1974, p. 760 (not Matschie, 1912, p. 308) — external and cranial
characters; measurements, most ex Allen, 1938, p. 292.

Macaca [(Lyssodes)] speciosa: Fiedler, 1956, p. 179 (not F. Cuvier, 1825, p. 2, or I. Geoffroy,
1826, p. 589) — part, Macacus thibetanus Milne-Edwards, 1870, included in synonymy.

Macaca speciosa speciosa: Pocock, 1939, p. 71 (not F. Cuvier, 1825, p. 2, or I. Geoffroy, 1826,
p. 589) — part, [Macacus (Magus) arctoides] esau Matschie, 1912, and Pithecus pullus Howell,
1928, included in synonymy.

Mlacaca] speciosa: Zhang et al. 1981, pp. 215, 221 (not I. Geoffroy, 1826, p. 589)— part,
included in distribution. Mao Jiangsen et al., 1981, p. 1590 — specimens used in hepatitis
research, collected in Zhejiang Province.

Types

The type-series available to Milne-Edwards (1870, p 341; [1872], p. 244) evi-
dently consisted of two mounted skins with skulls: an adult male (MNHN 329/
282A, 1870 No. 1, Type Cat. No. 60a), collected by A. David (1874, p. 9), 11
March 1869 (fig. 1); and an adult female (MNHN 330/282B, 1870 No. 527, Type
Cat. No. 60b, Coll. No. 109), acquired from a local hunter by A. David (1874,
p. 19), 31 March 1869. Both skins are preserved, but the skulls are missing, as
previously noted by Rode (1938, p. 223). The male skull is last known to have
been examined by Elliot ([1913], p. 197).

Rode (1938, p. 223) designated the male specimen as lectotype ("Holotype")
and the female specimen as paralectotype ("Allotype"). The lectotype skin and
skull were figured by Milne-Edwards ([1872], pis. 34, 35), who also published
measurements of the skulls and dry skins.

Type-Locality

The lectotype of M. thibetana was collected ca. 20 km NNW of Baoxing ( = Mou-
pin or Mu-ping), central Sichuan Province, China (David, 1874, p. 9). In the
original description (Milne-Edwards, 1870, p. 341), the geographic origin is im-
precisely given as "Thibet oriental." The paralectotype (female) also was collected
in the vicinity of Baoxing, where David (1874, p. 19) received it from a local
hunter.

Fig. 1. Macaca thibetana, lectotype skin and skull (A. Milne-Edwards, [1872J, pis. 34-35;
skull ca. 0.45 natural size), adult male collected by A. David, ca. 20 km NNW of Baoxing
(= Moupin), central Sichuan, China, 11 March 1869.

FOODEN: MACACA THIBETANA 5

Distribution (Fig. 2)

East-central China, ca. 25°00'N-33°00'N/ 102°30'E-119°30'E. The 13 known
locality records are restricted to upland areas in central Sichuan, southern Shaanxi,
western Hubei, northwestern Guangdong, eastern Jiangxi, northern and eastern
Fujian, and southwestern Zhejiang. The known altitudinal range is from ca.
1,000 m (Fuqing Xian; Hsiao Yang Chi; "Yao-tze Berge") to ca. 2,500 m (Baoxing,
NNW; David, 1871, p. 84, 1874, p. 9).

Contrary to the implication of the specific name, M. thibetana does not occur
within the currently recognized boundaries of the Tibetan Autonomous Republic
(now officially known as Xizang), as previously noted by Sowerby (1941, p. 263).
The type-locality of this species is 350 km east of the present border of Tibet
(Xizang), but this place was close to the Tibetan border as understood by David
(1874, p. 4) when he collected the type specimens.

External Measurements (Table 1)

Although it is clear from the evidence of dry skins that M. thibetana is a large
macaque, probably the largest species in the genus, and that tail length is about
10 percent of head and body length, actual flesh measurements recorded by
collectors are available for only 10 specimens, and of these only six are adults;
at least one set of adult measurements probably is invalid (Table 1). Recorded

Table 1.
thibetana.

External measurements recorded by collectors for 10

specimens of Macaca

Locality
No.1

Province

Head and

body length

(mm)

Tail
length
(mm)

Relative

tail length

(T/HB)

Adult males2

1

5
103
11

Sichuan
Sichuan
Fujian
Fujian

710

[?]950

ca. 700

6134

Adult females5

65

[?]90

ca. 50

55

0.09

[?]0.09

ca. 0.07

0.09

1
11

Sichuan
Fujian

630

507*

Subadult male

80
56

0.13
0.11

10

Fujian

607
Subadult females

67

0.11

1

5

Sichuan
Sichuan

650
600

Juvenile male

60
55

0.09
0.09

5

Sichuan

584

75

0.13

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

*Cf. dry skin measurements recorded by Milne-Edwards ([1872], p. 244) for lectotype
male, Sichuan: Loc. No. 4, HB 800 mm, T 100 mm, T/HB 0.13.

'Measurements from Sowerby, 1941, p. 263.

4Ear length 46 mm.

5Cf. dry skin measurements recorded by Milne-Edwards ([1872], p. 245) for paralectotype
female, Sichuan: Loc. No. 3, HB 540 mm, T 60 mm, T/HB 0.11.

"Ear length 42 mm.

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FOODEN: MACACA THIBETANA 7

head and body lengths are 507 mm and 630 mm in two adult females (650 mm
reported in one subadult) and 613 mm, ca. 700 mm, and 710 mm in three adult
males. Relative tail lengths (T/HB) are 0.11 and 0.13 in the adult females and
ca. 0.07, 0.09, and 0.09 in the adult males. Ear length is 42 mm in one adult
female (E/HB x 100 = 8.3) and 46 mm in one adult male (E/HB x 100 - 7.5).
Body weight information has not been recorded for museum specimens
examined, but Hu Xiaosu et al. (1982, p. 200) report that two large labora-
tory specimens (presumably adult males) obtained in Sichuan weighed 15 kg and
17.5 kg.

Pelage (Figs. 1, 3)

Dorsal surface of trunk dark brown, frequently with a diffuse blackish wash
caudally, hairs dense and long in prime pelage (5-10 cm long on scapular region
in adult males), hair banding indistinct or absent; dorsal surface of limbs colored
like back proximally, becoming paler distally, to pale brown on dorsum of hands
and feet; crown brown, paler than back, with hairs radiating to form a small
whorl (5 cm diameter) centered at vertex; skin of muzzle region pale brownish;
circumorbital skin color possibly sexually dimorphic, whitish in immatures and
adult males, pinkish in adult females (judging from color slides of Emei Shan
troop members kindly provided by Dr. J. D. Lazell, Jr., The Conservation Agency,
22 Dec. 1982); side-whiskers buffy, prominent (hairs 3-6 cm long), concealing
ears in anterior view; beard usually prominent, buffy to brown; hairs on ventral
surface of trunk and limbs pale buffy to pale gray, contrastingly paler and less
dense than on back.

Geographic variation of pelage color is negligible in specimens of M. thibetana
examined; adult specimens collected in Sichuan Province closely match those
collected in Fujian Province, 1,500 km to the east. Adult males and females
generally are similar in color, but the side-whiskers and beard are more prom-
inent in males. In one adult female (FMNH 39500, Hsiao Yang Chi), the ventral
surface of the tail is marked by a small bright patch of chestnut-colored fur. An

Fig. 3. Pelage variation in Macaca thibetana, adult males (left — photographed by G. Four-
nier at Emei Shan, August 1982, transmitted by Dr. J. D. Lazell, Jr., The Conservation
Agency; right — reproduced from Heck, 1916, pi. Affen III, fig. 9).

8 FIELDIANA: ZOOLOGY

albino captive specimen of this species reportedly was kept in a village in Guang-
dong Province, close to a natural habitat (Mell, 1922, p. 10).

Pelage color apparently darkens with age in M. thibetana. The smallest spec-
imen examined (AMNH 84475 6 , Chong'an Xian, skin only), an infant probably
only a few months old, is pale grayish brown on crown and back, buffy on the
sides and virtually hairless on the ventral surface. A larger infant (AMNH 60161
9, same locality, Ml visible in alveoli), probably about one year old, is buffy
brown on crown and back, paler on the sides and pale buffy on the ventral
surface. A young juvenile (USNM 259030 6 , Si Gi Pin, Ml erupting), probably
about 18 months old, is golden brown (near reddish brown) on crown and back.
Two older juveniles (USNM 241161 6, Si Gi Pin, 12 and M2 erupting; USNM
258687 6 , Yin Shiu Wan, skin only), probably both about three years old, are
respectively pale medium brown and dark medium brown on crown and back.
Older juveniles and subadults are only slightly paler than adults.

Seasonal molting in M. thibetana in late summer is suggested by pelage con-
dition in a series of four specimens collected in August (1925, 1929) at Si Gi Pin,
central Sichuan, as previously noted by Hill (1974, p. 765). In an adult female
(USNM 241162) collected 16 August 1925 and a subadult male (USNM 254800)
collected 6 August 1929, most of the dorsal hairs are only 2-3 cm long, but a
scattering of longer hairs (6-7 cm long) also is present. It appears that the longer
hairs in these specimens were in the process of being shed, thus revealing the
short (? new) fur. In a juvenile male (USNM 241161) collected 6 August 1925,
only short (? new) dorsal hairs are present. An adult male (USNM 241163)
collected 17 August 1925 is in prime pelage.

As previously indicated, M. thibetana frequently is confused with M. arctoides
because of the similar brown color, large size, and stump tail of these two species.
Living specimens of these species may be distinguished by conspicuous genital
characters (see Introduction). Dry museum specimens may be distinguished by
pelage characters, particularly around the face and on the ventral surface. In M.
thibetana, the forehead (to the brow ridges) and cheeks are thickly covered with
dense fur and the buffy side- whiskers are prominent; in M. arctoides, the forehead
and cheeks are only thinly covered by very short hairs (superficially, forehead
and cheeks appear bald) and hairs of the side-whisker region are indistinguish-
able in color and length from those of adjacent regions on the head. Fur on the
ventral surface in M. thibetana is pale buffy to pale gray, distinctly paler than
that on the back; fur on the ventral surface in M. arctoides is dark brown, similar
in color to that on the back. Texture of the dorsal pelage also seems to differ in
these two species; dorsal hairs appear coarser in M. thibetana (woolly) than in
M. arctoides (silky).

Cranial Characters (Fig. 4; Table 2)

Skull large, greatest length 130.9 ± 7.6 mm in five adult females, 157.1 ± 5.4
mm in nine adult males (cf. Albrecht, 1978, p. 129); relative zygomatic breadth
(ZB/GL) 0.67 ± .01 and 0.67 ± .02 in the same males and females; rostral-
postrostral ratio 0.52 ± .01 in three adult females, 0.58 ± .03 in seven adult
males; supramaxillary ridges weakly developed in adult males; supraorbital ridges
thick in adult males (10-13 mm); sagittal crest and nuchal crest prominent (6-8
mm) in old adult males.

Table 2. Cranial measurements' and ratios in 25 specimens of Macaco thibetana2.

Skull,

Relative

'

greatest

zygomatic

Piistrostr.il

Rostral-

Locality

length

breadth

length

postrostr.il

No.'

Province

(mm)

(ZB/GL)

(mm)

ratio

Adult males4

1

Sichuan

161.3

0.68

109.5

0.58

2

Sichuan

161.7

0.68

107.4

0.61

45

Sichuan

164.0

0.65

5

Sichuan

151.7,158.5

0.68,0.67

102.1,105.9

0.59,0.59

_6

?Sichuan

157.8

0.65

107.0

0.61

8

Guangdong

; 152.2

0.65

11

Fujian

159.1

0.65

110.1

0.55

12

Fujian

147.7

0.69

103.7

0.53

x ± S.D.(n)

157.1 ± 5.4(9)7

0.67 ± .02(9)

106.5 ± 2.7(7)

0.58 ± .03(7)

Adult females

1

Sichuan

136.4

0.68

97.3

0.53

2

Sichuan

127.6

0.66

91.3

0.51

35

Sichuan

140.0

0.67

. . .

. . .

11

Fujian

120.7

0.68

. . .

. . .

_»

?Fujian

129.9

0.65

92.8

0.52

x±S.D.(n)

130.9 ± 7.6(5)9

0.67 ± .01(5)

93.8 ±3.1(3)

0.52 ± .01(3)

Subadult males

2

Sichuan

142.1

0.61

95.9

0.58

10

Fujian

140.3

0.64

98.8

0.51

ll10

Fujian

143.9

0.69

98.1

0.58

x±S.D.(n)

142.1 ± 1.8(3)

0.65 ± .04(3)

97.6 ± 1.5(3)

0.56 ± .04(3)

Subadult females

1

Sichuan

125.1

0.68

90.3

0.50

5

Sichuan

127.8

0.66

93.7

0.48

x±S.D.(n)

126.4 ± 1.9(2)

0.67 ± .01(2)

92.0 ± 2.4(2)

0.49 ± .01(2)

Juveniles: 4 males, 1 female (Loc. No. 10)

2

Sichuan

98.2,110.2

0.67,0.66

80.8,86.5

0.32,0.37

5

Sichuan

138.2

0.65

96.9

0.53

10

Fujian

120.0

0.63

. . .

no"

Fujian

125.8

0.67

91.2

0.5

x±S.D.(n)

118.4 ± 15.2(5)

0.66 ± .02(5)
Infant male

88.8 ± 6.8(4)

0.43 ± .10(4)

11

Fujian

86.0

0.68

74.1

0.28

'For detailed explanation of measurements, see Fooden, 1969, p. 41.

Includes all specimens tabulated by Allen (1938, p. 292), Hill (1974, p. 761) and Albrecht
(1978, p. 114).

'For key to locality numbers, see distribution map (Fig. 2).

4 Age class criteria: adults — last molars and permanent canines completely erupted; sub-
adults — last molars in females or permanent canines in males emergent but not completely
erupted; juveniles — one or more permanent teeth erupted, but prior to subadult stage;
infants — deciduous teeth only.

'Measurements from Milne-Edwards ([1872), p. 247).

locality: "Chinese Thibet."

"Sichuan specimens, 159.2 ± 4.3(6); Guangdong and Fujian specimens, 153.0 ± 5.7(3).

"Locality: "China? No data."

"Sichuan specimens, 134.7 ± 6.4(3); Fujian specimens, 125.3 ± 6.5(2).

'"Specimen reported as adult by Albrecht (1978, p. 114); locality listed as "Fu-ching",
based on label error.

"Locality: "probably Kuatun."

10

FOODEN: MACACA THIBETAN A 11

In the western part of the range (Sichuan), skull length in M. thibetana may
average somewhat greater than in the eastern part (Guangxi, Fujian), judging
from the few adult specimens available (Table 2). Dental anomalies occur in two
of five skulls (USNM 258649, USNM 258651, both adult males) collected at Yin
Shiu Wan, Central Sichuan: both of these specimens lack right and left lower
lateral incisors, and one (USNM 258651) also lacks left upper and lower third
molars; no trace of alveoli for these teeth is evident.

Ontogenetically, relative zygomatic breadth remains approximately constant
from infancy to adulthood, while rostral-postrostral ratio increases allometrically
(Table 2; fig. 5). The power function equation for the ontogenetic relationship
between rostral length (y) and postrostral length (x) determined by the method
of principal axes is:

logy = 3.103 log x - 4.468.

The 95 percent confidence limits for the slope are L, = 2.723 and L2 = 3.591.

Natural History

Available information concerning the natural history of M. thibetana is meager.
Brief field notes have been recorded by four collectors (Table 3). Two other
observers have published notes that may refer either to M. thibetana or to M.
mulatta, or to both (La Touche in Thomas, [1899], p. 770; Wilson, 1913, vol. 1,
p. 198).

The known limits of distribution of M. thibetana roughly correspond to the
boundaries of the zoogeographic area designated by Zhang et al. (1981, p. 217)
as the "Central China region." In this area the temperature remains above 10°
C for 225-280 days per year, and the climax vegetation generally is subtropical
broadleaf evergreen forest (Zhang et al., 1981, p. 216); Quercus, Castanopsis and
Pasania are the dominant forest genera (Wang, 1961, p. 133; Sochava & Lukicheva,
1964, p. 110). Long-continued intensive agriculture has reduced the natural forest
to scattered remnants on unarable uplands (Zhang et al., 1981, p. 222).

Most observers agree that M. thibetana is restricted to forested areas within its
range (David, 1871, p. 88; David, 1875, p. 324; Wilson, 1913, vol. 1, p. 224; Smith
in Sowerby, 1929, p. 315; Pope, 1932, p. 495). Only Mell (in Matschie, 1912, p.
309), who observed this species in northwestern Guangdong, reports that it
inhabits rocky cliffs, not forests; perhaps the monkeys encountered by Mell were
at the edge of a forested area. Within forests, this species has been sighted both
in trees and on the ground (David, 1875, p. 256; Smith in Sowerby, 1929, p.
315); available evidence is inadequate to assess the relative degree of arboreality
or terrestriality. Footprints in the snow followed by David (1874, p. 9) indicate
that one old male walked on the ground for some distance; this individual
subsequently hid in a rocky hollow.

The diet of M. thibetana has not been recorded, but presumably it consists
primarily of fruit supplemented by some animal food, as in other species of
macaques. At Emei Shan, central Sichuan, Wilson (1913, vol. 1, p. 224) reports
that these monkeys "are fond of the blue pod-like fruit of Decaisnea, the shining
black flattened seeds of which, however, I noticed they cannot digest."

Until 50-100 years ago, M. thibetana apparently was fairly common within its
restricted habitat (David, 1873, collector's note, MNHN 1874/479, Kuatun; Mell
in Matschie, 1912, p. 309; Wilson, 1913, vol. 1, p. 224; Smith in Sowerby, 1929,
p. 315; Guo Zhusong, 1981, p. 65). David (1871, p. 88) encountered an old hunter

40 70 10 90 100 110 120

Postrostral length (mm)

Fie. 5. Allometry of rostral length (y) vs. postrostral length (x) in 20 immature and adult
specimens of Macaca thibetana (log-log plot; open circles = females, closed circles = males;
smaller circles = immatures, larger circles = adults); principal axis, log y = 3. 103 log x
- 4.468.

12

FOODEN: MACACA THIBETANA 13

in central Sichuan who claimed to have taken nearly 800 M. thibetana individuals
during his lifetime. Troop size has been recorded only once (David, 1875, p.
256), in a troop that contained 10 members; among these was one that was
reddish, probably a young juvenile (see above, Pelage). Solitary individuals have
been reported by David (1874, p. 9) in central Sichuan and by Mell (in Matschie,
1912, p. 309) in northwestern Guangdong. Concerning possible reproductive
seasonality, all that is known is that an "infant in arms" (AMNH 84475 6 , skin
only, Chong'an Xian), probably only a few months old, was collected by C. H.
Pope in northwestern Fujian in mid-May 1926.

Three other species of nonhuman primates — Rhinopithecus (or Pygathrix) rox-
ellanae, Macaca arctoides, M. mulatta — occur in or near the range of M. thibetana.
R. roxellanae has been collected or observed near M. thibetana, along the north-
western edge of the range of M. thibetana, from central Sichuan (David, 1871,
p. 88) to western Hubei (Xiao Zhi, 1979, p. 31); according to David (loc. cit.),
R. roxellanae occurs at higher altitudes than M. thibetana. M. arctoides, which
generally ranges to the southwest of M. thibetana, has been collected once (ZMB

Table 3. Field notes recorded by collectors concerning Macaca thibetana.

1(a). A. David (1874, p. 9), 11 March 1869, Loc. No. 4, Baoxing, 20 km NNW (translated
from French) — Having walked to the upper end of our valley 5 leagues [20 km] from my
living quarters, I discovered in the snow the tracks of a fairly large monkey mixed with
the tracks of a leopard. By following the tracks for some time, we were able to locate in
a rocky hollow an old monkey with long fur and a short tail [identified in footnote as
"Macacus Tibetanus (A. Edw.)"]; a fortunate shot brought it down dead at our feet. . . .
This monkey lives in mountains that are quite cold where there is an abundance of large
trees, firs and other gigantic conifers, of which a great number of trunks lie piled up and
rotting in the mountain streams.

1(b). A. David (1875, pp. 255-6), 8 Oct. 1873, Loc. No. 9, Kuatun, ca. 20 km NW
(translated from French)-— In the morning we travel in a long cool valley that has its
bordering slopes well forested; I am all eyes (as far as my illness permits) to perceive some
trace of the monkeys that I am told abound in these parts. In fact, we pass some houses
where people keep two or three monkeys that were captured in this region. ... A little
farther, I myself see a group of 10 of these brown-haired macaques in the trees and on
the rocks; I am very surprised to see among them one individual that is pale reddish and
appears to belong to a different species [possibly a juvenile; see above, Pelage]. In fact,
the mountain people with me say that there are two or three different species of monkeys
here [M. thibetana, M. mulatto].

1(c). A. David (1875, p. 324), summary of observations (translated from French) — Ma-
cacus tcheliensis [ = Macaca mulatta] lives in small numbers in rocky places from the cold
mountains near Beijing to Sichuan and Fujian. Macacus tibetanus lives in the center of the
[Chinese] empire, from west to east, in forested regions.

2. R. Mell (in Matschie, 1912, p. 309), 1908-11, Loc. No. 8, "Yao-tze Berge" (translated
from German) — In this mountainous region, 2000 m estimated altitude [subsequently
corrected to 800-1,200 m; Mell, 1922, p. 10], this monkey is said to be not uncommon. I
have only seen it solitary and, as indeed would be inferred from its short limbs, only on
rocky slopes, not in forest. The same sorts of places also are inhabited by the red-faced
monkey [= M. arctoides].

3. F. T Smith (in Sowerby, 1929, p. 315), 16 May-24 June 1926, Loc. No. 10, Kuatun—
. . . while in spots these monkeys were fairly numerous, they were exceedingly shy and
difficult to approach, since they kept well hidden in the thick foliage of the trees, and
could only be located by their calls. Another much smaller macaque [M. mulatta] was also
secured at the same place. . . .

4. C H. Pope (in Allen, 1938, p. 293), July 1925-May 1926, Loc. No. 11, Chong'an
Xian — . . . they prefer the more rugged precipitous sections.

14 FIELDIANA: ZOOLOGY

15925, Lien Shan [ = Lianshan], NW) about 135 km southwest of a M. thibetana
locality ("Yao-tze Berge"), near the southeastern limit of distribution of M. thi-
betana; the collector (Mell in Matschie, 1912, p. 309) indicates that M. arctoides
(= M. arctoides melli Matschie, 1912) and M. thibetana (= M. arctoides esau Mat-
schie, 1912) occupy the same kind of habitat in this area.

M. mulatto is broadly sympatric with M. thibetana, but these two species may
differ in their ecological preferences. M. mulatto and M. thibetana have been
collected or observed within 25 km of each other in central Sichuan, southern
Shaanxi, western Hubei, and northwestern Fujian. References to the occurrence
of M. mulatto in these areas are: Baoxing vicinity, David, 1871, p. 88; Emei Shan
vicinity, USNM 241160 (Gin K'eo Ho), BM 11.9.8.1 (Kia-ting, SW), RMNH (Lei-
den) 4585/V67-W50 (Wa-shan); southern Shaanxi, David, 1875, p. 42; western
Hubei, Xiao Zhi, 1979, p. 31; Kuatun vicinity, David, 1875, pp. 281, 285; Kuatun,
BM 97.6.5.2, BM 98.11.1.29, MNHN 336/480, 337/481; Chong'an Xian, AMNH
84474, 84476. At Kuatun, northwestern Fujian, La Touche (in Thomas, [1899],
p. 770) reports that M. mulatto is more abundant than M. thibetana (not specifically,
identified by La Touche), but David's (1875, p. 281) hunters at this place required
almost one month longer to obtain M. mulatto than to obtain M. thibetana.
Throughout the range of M. thibetana, David (1875, p. 324) asserts that it favors
forested areas while M. mulatto favors more open rocky areas.

Specimens Examined

Total, 29.

Skins and skulls, 20— SICHUAN: Loc. Nos. 1, Hsiao Yang Chi (3— FMNH 39499 ad. 6.
39500 ad. 9, 39501 subad. 9); 2, Si Gi Pin (5— USNM 241161 juv. d, 241162 ad. 9, 241163
ad. 6. 254800 subad. 6, 259030 juv. d); 5, Yin Shiu Wan (4— USNM 258649 ad. 6, 258650
subad. 9, 258651 ad. 6, 258686 juv. <J). GUANGDONG: Loc. No. 8, "Yao-tze Berge" (1—
ZMB 16179 ad. 6). FUJIAN: Loc. Nos. 10, Kuatun (2— MNHN 1874/479 juv. 9, USNM
252157 subad. 6); 11, Chong'an Xian (4— AMNH 60161 inf. 9, 84472 ad. 6, FMNH 39383
subad. 6, MCZ 26477 ad. 9); 12, Fuqing Xian (1— AMNH 60160 ad. 6).

Skins only, 6— SICHUAN: Loc. Nos. 3, Baoxing, vie. (1— MNHN 1870/527 ad. 9); 4,
Baoxing, ca. 20 km NNW (1— MNHN 1870/1 ad. 6); 5, Yin Shiu Wan (1— USNM 258687
juv. 6). FUJIAN: Loc. Nos. 10, Kuatun (2— MNHN 1874/477 ad. 6, 1874/478 juv. 9); 11,
Chong'an Xian (1— AMNH 84475 inf. 6).

Skulls only, 3— "China? No data" (1— AMNH 83994 ad. 9); "Chinese Thibet" (1—
AMNH 129 ad. 6), "probably Kuatun" (1— AMNH 60162 juv. d).

For further details, see Gazetteer.

GAZETTEER

Locality names listed in this gazetteer preferentially are Pinyin names ap-
proved by the U.S. Board on Geographic Names (USBGN) in Gazetteer of the
People's Republic of China (1979). Primary entries for macaque localities that
are not included in the USBGN gazetteer are spelled here as in the original
source. Secondary entries, with cross references to corresponding primary en-
tries, give variant spellings or alternate locality names that appear on specimen
tags or in published literature on M. thibetana.

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

1. locality name

2. altitude, if reported by collector or observer

3. name of province (capitalized)

4. coordinates of locality

FOODEN: MACACA THIBETANA 15

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 kept 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. 2).

Baoxing, ca. 20 km NNW; SICHUAN; ca. 30°35'N, 102°35'E; collected by A.

David (1874, p. 9), 11 Mar. 1869 (MNHN, 1 [skin only]). 4
Baoxing, vicinity; SICHUAN; ca. 30°23'N, 102°50'E; obtained from local hunter

by A. David (1874, p. 19), 31 Mar. 1869 (MNHN, 1 [skin only]). 3

Chensi. See Shaanxi.

"China? No data"; [probably FUJIAN]; collected by R. C. Andrews' Third Asiatic

Exp., date unknown (AMNH, 1 [skull only]). Not mapped.
"Chinese Thibet"; [possibly SICHUAN]; Verreaux Coll., 1844 (AMNH, 1 [skull

only]). Not mapped.
Chong'an Xian [county]; FUJIAN; ca. 27°51'N, 117°49'E (Pope, 1929, p. 345;

1932, pp. 491, 495; 1935, p. 499); collected by C. H. Pope (citations above, and

1940, p. 72), 7 July 1925, 13-16 May 1926 (AMNH, 3 [1 skin only]; FMNH, 1;

MCZ, 1). 11
Chungan Hsien. See Chong'an Xian.

Daba mountain range. See Shennongjia Mountains.

Emei Shan, ca. 1,900 m; SICHUAN; ca. 29°32'N, 103°21'E; sight record, E. H.
Wilson (1913, vol. 1, p. 224, vol. 2, p. 192), Oct. 1903 (also see Heim, 1932,
p. 2); photograph published (Guo Zhusong, 1981, p. 62), date unknown;
observed by J. D. Lazell, Jr. (letter 22 Dec. 1982), August 1982. 2

Fokien occidental. See Kuatun.

Fu-ch'ing Hsien. See Fuqing Xian.

Fukien. See Fuqing Xian.

Fuqing Xian [county]; FUJIAN; ca. 25°44'N, 119°24'E (Pope, 1935, p. 494); col-
lected by C. H. Pope (1929, p. 345; 1932, p. 493; 1935, p. 11), 6 Oct. 1925
(AMNH, 1). 12

Futsien. See Fuqing Xian.

Futsing Hsien. See Fuqing Xian.

Hokow. See Yalong Jiang.

Hong-chan-tin. See Baoxing, ca. 20 km NNW.

Hsiao Yang Chi; SICHUAN; 29°07'N, 103°18'E (unpublished map, Division of

Birds, FMNH); collected by F. T. Smith, 10 June 1932 (FMNH, 3). 1
Hung Shan Ting. See Baoxing, ca. 20 km NNW

Jalungfluss. See Yalong Jiang.
Jao-tze Bergen. See "Yao-tze Berge."
Ju-juen. See "Yao-tze Berge."
Ju-juen, village near. See Ruyuan.

16 FIELDIANA: ZOOLOGY

Kaoten. See Kuatun.

Koaten. See Kuatun. ,

"Kuangsi" ( = GUANGXI); province uncertain; unreliable record based on al-
leged origin of captive specimen observed in Canton ( = Guangzhou) market
by R. Mell (1922, p. 10), 1908-21. Not mapped.

Kuangtung, Hunan, Kuangsai: Grenzgebiete zw. See "Yao-tze Berge."

Kuatun; FUJI AN; ca. 27°51'N, 117°48'E (La Touche, 1892, p. 402; Pope, 1929, p.
343, 1935, p. 496); collected by A. David (1875, p. 267), 20 Oct. 1873 (MNHN,
3 [2 skins only]); F. T. Smith (1926, p. 131), [16 May-24 June 1926] (USNM, 1;
another specimen retained in Shanghai according to Sowerby, 1929, p. 315).
For additional locality notes see La Touche (1899, p. 173; in Thomas, [1899],
p. 769 and Pope (1932, p. 490). 10

Kuatun, [ca. 20 km NW]; JIANGXI; ca. 28°00'N, 117°40'E; observed by A. David
(1875, p. 256), 8 Oct. 1873. 9

Kuatun, "probably"; FUJIAN; ca. 27°51'N, 117°48'E; collected by C. H. Pope
(1932, p. 490), prior to 1 Feb. 1926 (AMNH, 1 [skull only]). Not mapped.

Kuei-yung, S; SICHUAN; ca. 30°30'N, 102°00'E; questionable record (Hill, 1974,
p. 766), based on inconclusive field note of E. H. Wilson (1913, vol. 1, p. 198:
"The men who were in front of me saw several troupes of monkeys [species
unreported] . . . , but I saw no animals. . . ."), 1908. Not mapped.

Lien-dsau. See "Yao-tze Berge."
Lihn-dsau. See "Yao-tze Berge."
Log-dsong. See "Yao-tze Berge."
Lo-tschang-ho, W. See "Yao-tze Berge."

Mapienting, [WNW]. See Hsiao Yang Chi.
Mount Emei. See Emei Shan.
Mount Omei. See Emei Shan.
Moupin. See Baoxing.
Muping. See Baoxing.

Nan-Ling-Gebirge. See "Yao-tze Berge."

Omei, Mt. See Si Gi Pin.

Ra-ma-la. See Yalong Jiang.

Ruyuan, village near; GUANGDONG; ca. 24°46'N, 113°16'E; captive observed
by R. Mell (1922, p. 10), 1908-21. Not mapped.

San Chiang. See Chong'an Xian.

Shaanxi, southern; SHAANXI; ca. 33°N, 107°E; observed by A. David (1875, p.
42), Oct. 1872-Apr. 1873. 6

Shennongjia Mountains; HUBEI; ca. 31°45'N, 110°45'E; reported by Xiao Zhi
(1979, p. 31), date unknown. 7

Shensi. See Shaanxi.

Shin Kai Si. See Si Gi Pin.

Sichuan Province; SICHUAN; ca. 26°-34° N, 98°-110° E; imprecise origin of lab-
oratory specimens, reported by Hu Xiaosu et al., p. 200. Not mapped.

FOODEN: MACACA TH1BETANA 17

Si Gi Pin, 6,000 ft [1,800 m]; SICHUAN; ca. 29°30'N, 103°30'E (unpublished map
in Smithsonian Institution archives, Accession No. 89413, 1925); collected by
D. C. Graham and Yang Fong Tsang, 11-17 Aug. 1925, 6 Aug. 1929, 7 July
1934 (USNM, 5). 2

Singolo. See Yalong Jiang.

Suichang Xian [county], "mountainous country"; ZHEJIANG; ca. 28°35'N,
119°15'E; origin of captive laboratory specimens, reported by Mao Jiangsen et
al., 1981, p. 1590 (county data in letter 4 June 1982). 13

Wa Shan, region; SICHUAN; ca. 29°20'N, 103°05'E; tentative report, E. H. Wilson

(1913, vol. 2, p. 192), July 1903. Not mapped.
Wen Chuan, near. See Yin Shiu Wan.

Yalong Jiang; SICHUAN; ca. 30°N, 101°E; misleading report (Schafer, 1942, pp.
208, 257; cf. Schafer, 1933, pp. 192, 280), "Tibetmakaken" ( = M. mulatto, not
M. thibetana), collected by E. Schafer, 31 Oct. 1931 (Academy of Natural Sci-
ences, Philadelphia, 1 [skin], MCZ [skull]). Not mapped.

"Yao-tze Berge", 800-1, 200m; GUANGDONG; ca. 25°15'N, 113°15'E; collected
by R. Mell (in Matschie, 1912, p. 309; 1922, pp. 4, 10), 26 Sep. 1908-9 Feb.
1911 (ZMB, 1). 8

Yin Shin Wan. See Yin Shiu Wan.

Yin Shiu Wan, ca. 6,000 ft [1,800m]; SICHUAN; ca. 31°05'N, 103°31'E (Helde,
1923, map); collected by D. C. Graham and Yang Fong Tsang, 10 Apr.-4 Nov.
1933 (USNM, 5 [1 skin only]). 5

Yin Shu Wan. See Yin Shiu Wan.

Zhejiang Province. See Suichang Xian.

LITERATURE CITED

Albrecht, G. H. 1978. The craniofacial morphology of the Sulawesi macaques: multivariate

approaches to biological problems. Contrib. Primat., 13: 1-151.
Allen, G. M. 1930. Primates and pangolins from the Asiatic Expeditions. Amer. Mus.

Nov., no. 429: 1-7.
. 1938. The mammals of China and Mongolia. Part 1. American Museum of Natural

History, New York.
Bertrand, M. 1969. The behavioral repertoire of the stump-tail macaque: a descriptive

and comparative study. Bibl. Primat., 11: 1-273.
Blanchard, E. 1871. Les recentes explorations de la Chine: IV. Voyage dans les provinces

centrales de la Chine et au Thibet oriental. Rev. Deux Mondes, 93: 611-32.
Buettner-Janusch, J. 1963. An introduction to the primates, pp. 1-64. In Buettner-Ja-

nusch, J., ed., Evolutionary and Genetic Biology of Primates, vol. 1. Academic Press,

New York and London.
Cuvier, F. 1825. Macaque a face rouge, liv. 47, pp. 1-2. In Geoffroy Saint-Hilaire, E., and

F. Cuvier, Histoire naturelle des mammiferes, vol. 5. A. Belin, Paris.
David, A. 1869. Voyage en Chine fait sous les auspices de S. Exc. le Ministre de lTnstruction

publique, pt. 3. Nouv. Arch. Mus. Hist. Nat. Paris, 5 (Bull.): 3-13.
. 1871. Rapport adresse a MM. le professeurs-administrateurs du Museum d'Histoire

Naturelle. Nouv. Arch. Mus. Hist. Nat. Paris, 7(Bull.): 75-100.

1874. Journal d'un voyage dans le centre de la Chine et dans le Thibet oriental.

Troisieme partie. Sejour dans la principaute de Moupin. Nouv. Arch. Mus. Hist. Nat.
Paris, 10(Bull.): 3-82.

18 FIELDIANA: ZOOLOGY

. 1875. Journal de mon troisieme voyage d'exploration dans 1'empire Chinois, vol.

2. Hachette, Paris.
Dobroruka, L. j. 1967. Uber den "Blassgesichtsmakak" nebst einer Ubersicht iiber die

Art Macaca speciosa (F. Cuvier 1825). Milu, 2(3): 305-12.
Elliot, D. G. (1913). A review of the primates, vol. 2. American Museum of Natural

History, New York. For date of publication, see Zool. Rec: Mammalia, 1913: 9.
Fiedler, W. 1956. Ubersicht iiber das System der Primates, pp. 1-266. In Hofer, H., A. H.

Schultz, and D. Starck, eds., Primatologia, vol. 1. Karger, Basel.
Fooden, J. (1966). Identification of the stump-tailed monkey, Macaca speciosa. I. Geoffroy,

1826. Fol. Primat., 5(3): 153-64.
. 1967. Complementary specialization of male and female reproductive structures

in the bear macaque, Macaca arctoides. Nature, 214(5091): 939-41.

1969. Taxonomy and evolution of the monkeys of Celebes (Primates : Cercopi-

thecidae). Bibl. Primat., 10: 1-148.

1971a. Report on primates collected in western Thailand January-April, 1967.

Fieldiana: Zool., 59(1): 1-62.

1971b. Female genitalia and taxonomic relationships of Macaca assamensis. Primates,

12(1): 63-73.

1976. Provisional classification and key to living species of macaques (Primates:

Macaca). Fol. Primat., 25(2-3): 225-36.

1979. Taxonomy and evolution of the sinica group of macaques: 1. Species and

subspecies accounts of Macaca sinica, Primates, 20(1): 109—40.

1981. Taxonomy and evolution of the sinica group of macaques: 2. Species and

subspecies accounts of the Indian bonnet macaque, Macaca radiata. Fieldiana: Zool., n.s.,
no. 9: 1-52.

-. 1982. Taxonomy and evolution of the sinica group of macaques: 3. Species and

subspecies accounts of Macaca assamensis. Fieldiana: Zool., n.s., no. 10: 1-52.
Forbes, H. O. 1894. A hand-book to the primates, vol. 2. Allen, London.
Gazetteer of the People's Republic of China: Pinyin to Wade-Giles, Wade-Giles to

Pinyin. 1979. Defense Mapping Agency, Washington, D.C.
Geoffroy Saint-Hilaire, I. 1826. Macaque, pp. 584-90. In Bory de Saint-Vincent, J.-B.,

G.-M.,ed., Dictionnaire classique d'histoire naturelle, vol. 9. Rey et Gravier and Baudoin

Freres, Paris.
. 1831. Mammiferes, pp. 1-160. In Belanger, C, Voyage aux Indes-Orientales . . .

Zoologie. Bertrand, Paris. For date of publication, see Ann. Mag. Nat. Hist., (7)7: 390

(1901).
Guo Zhusonc. 1981. Marvelous sights on Mount Emei. China Reconstructs, 30(8): 60-65.
Heck, L. 1916. Hen-en- oder Hochtiere (Primates), pp. 380-699. In Hilzheimer, M. and L.

Heck, eds., Brehms Tierleben, 4th ed., Die Saugetiere, vol. 4. Bibliographisches Institut,

Leipzig/Wien.
Mm m. A. 1932. Tectonical study of Omei-shan Szechuan. Geol. Surv. Kwangtung & Kwangsi

(Liang-kwang Ti-chih Tiao-cha-so), Spec. Publ. No. 13. 54 pp.
Mm in G. G. 1923. Map: part of western Szechwan. J. W China Bord. Res. Soc., 1: folding

map, end of volume.
Hill, W. C. O 1974. Primates: Comparative anatomy and taxonomy. VII. Cynopithecinae:

Cercocebus, Macaca. Cynopithecus. John Wiley it Sons, New York.
Howell, A. B. 1928. New Chinese mammals. Proc. Biol. Soc. Wash., 41: 41-44.
. 1929. Mammals from China in the collections of the United States National Mu-
seum. Proc. U.S. Nat. Mus. 75: 1-82.
Hu Xiaosu, Hu Anqi, Lin Fancqinc, Yan Heping, Zhu Shenghua, and Deng Anxiao.

1982. On the infection of Macaca thibetanus with Plasmodium knowlesi. Acta Zoologica

Sinica, 28(2): 200-201.
Kellogg, R. 1945. Macaques, pp. 113-34. In Aberle, S. D., ed.. Primate Malaria. National

Research Council, Division of Medical Science, Office of Medical Information, Wash-
ington, D.C.

FOODEN: MACACA THIBETANA 19

La Touche, J. D. de. 1892. On birds collected or observed in the vicinity of Foochow and
Swatow in south-eastern China. Ibis, (6)4 (16): 400-430.

. 1899. Notes on the birds of North-west Fohfcien. Ibis, (7)5 (18): 169-210.

Lydekker, R. [1907]. On the occurrence of the bruang in the Tibetan Province. Proc. Zool.
Soc. London, 1906: 997-9. For date of publication, see Proc. Zool. Soc. London (Ser.
A), 107: 76 (1937).

Mao Jiangsen, Guo Xingying, Huang Haiying, Yu Peihua, Huang Bozhang, Ding
Zhanchu, Chen Nianliang, Yu Junhao and Xie Ruying. 1981. Studies on the trans-
mission of human hepatitis A virus to stump-tailed monkey. Scientia Sinica, 24(11):
1590-6.

Matschie, P. 1912. Zur Kenntnis der sudchinesischen Kurzschwanz-Makaken. Sitz. Ges.
naturf. Freunde Berlin, 1912(5): 305-10.

Mell, R. 1922. Beitrage zur Fauna sinica. I. Die Vertebraten Siidchinas; Feldlisten und
Feldnoten der Sauger, Vogel, Reptilien, Batrachier. Arch. Naturgesch. 88(10), Abt. A:
1-146.

Milne-Edwards, A. 1870. Note sur quelques mammiferes du Thibet oriental. C. R. Hebd.
Seances Acad. Sci. Paris, 70: 341-2.

. [1872]. Memoire sur la faune mammalogique du Tibet oriental et principalement

de la principaute de Moupin, pp. 231-386. In Milne-Edwards, H., and A. Milne-Ed-
wards, Recherches pour servir a 1'histoire naturelle des mammiferes. G. Masson, Paris.
For date of publication, see Zool. Rec: Mamm., 9: 2 (1872).

-. 1892. Observations sur les mammiferes du Thibet. Rev. Gen. Sci. Pures Appl.,

3(19): 670-2.
Petzold, H.-G. 1964. Im Tierpark Berlin 1959 erstmalig gehaltene Tierformen. Milu, 1:

317-43.
Pocock, R. I. [1926]. The external characters of the catarrhine monkeys and apes. Proc.

Zool. Soc. London, 1925 (4): 1479-1579. For date of publication, see Proc. Zool. Soc.

London, 1926(1): outside back cover.
. 1939. The fauna of British India, including Ceylon and Burma. Mammalia. Vol. 1.

Primates and Carnivora (in part). Taylor and Francis, London.
Pope, C. H. 1929. Notes on reptiles from Fukien and other Chinese provinces. Bull. Amer.

Mus. Nat. Hist., 58(8): 33S487.
. 1932. Fukien and Kiangsi Provinces, pp. 488-500. In Andrews, R. C, ed., The

New Conquest of Central Asia. American Museum of Natural History, New York.
. 1935. The reptiles of China: turtles, crocodilians, snakes, lizards. American Mu-
seum of Natural History, New York.
. 1940. China's animal frontier. Viking Press, New York.

Rode, P. 1938. Catalogue des types de mammiferes du Museum National d'Histoire Na-
turelle. I. Ordre des Primates. A. Sous-ordre des simiens. Bull. Mus. Nat. Hist. Nat.,
Paris, (2)10(3): 202-51.

Schafer, E. 1933. Berge, Buddhas und Baren: Forschung und Jagd in geheimnisvollem
Tibet. Paul Parey, Berlin.

. 1942. Tibet ruft: Forschung und Jagd in den Hochgebirgen Osttibets Tibetexpe-

dition 1931/32. Paul Parey, Berlin.

Smith, F T. 1926. The Fukien Scientific Expedition. China J. Sci. Arts, 5(3): 128-31.

Sochava, V. B. and A. N. Lukicheva. 1964. [Vegetation: Asia], pp. 110-11. In Gerasimov,
I. P., ed., [Physical-geographic atlas of the world]. Akademia Nauk SSSR, Moscow.

Sowerby, A. de C. 1929. A new monkey from China. China J., 11(6): 315.

. 1941. Apes, monkeys and lemurs of China. China J., 34(6): 259-65.

Tate, G. H. 1947. Mammals of Eastern Asia. The Macmillan Co., New York.

Thomas, O. [1899]. On mammals collected by Mr. J. D. La Touche at Kuatun, N. W. Fokien,
China. Proc. Zool. Soc. London, 1898: 769-75. For date of publication, see Proc. Zool.
Soc. London (Ser. A), 107: 75 (1937).

Trouessart, E.-L. 1878. Catalogue des mammiferes vivants et fossiles. Rev. Mag. Zool.
Pure Appl., (3)6: 108-40.

20 FIELDIANA: ZOOLOGY

. 1897. Catalogus mammalium tarn vi ventium quam fossilium, fasc. 1 . Friedlander

& Sohn, Berlin.

Wang, C.-W. 1961. The forests of China, with a survey of grassland and desert vegetation.
Maria Moors Cabot Found. Publ. No. 5, Harvard Univ. xiv + 313 pp.

Wilson, E. H. 1913. A naturalist in western China . . . , 2nd ed., 2 vol. Methuen, London.

Xiao Zhi. 1979. Shennongjia forests: home of rare species. China Reconstructs, 28(8): 28-
32.

Zhang, Y.-Z, S. Wang, and G.-Q. Quan. 1981. On the geographical distribution of pri-
mates in China. J. Hum. Evol., 10: 215-26.

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