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APR 91992
FIELDIANA
Zoology
NEW SERIES, NO. 45
Jack Fooden
Taxonomy and Evolution
of the Sinica Group of Macaques:
6. Interspecific Comparisons and Synthesis
%
^
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June 30, 1988
Publication 1389
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Croat, T. B. 1978. Flora of Barro Colorado Island. Stanford University Press, Stanford, Calif, 943 pp.
Grubb, P. J., J. R. Lloyd, and T. D. Pennington. 1963. A comparison of montane and lowland rain forest
in Ecuador. I. The forest structure, physiognomy, and floristics. Journal of Ecology, 51: 567-601.
Langdon, E. J. M. 1979. Yage among the Siona: Cultural patterns in visions, pp. 63-80. In Browman, D. L.,
and R. A. Schwarz, eds., Spirits, Shamans, and Stars. Mouton Publishers, The Hague, Netherlands.
Murra, J. 1946. The historic tribes of Ecuador, pp. 785-821. In Steward, J. H., ed., Handbook of South
American Indians. Vol. 2, The Andean Civilizations. Bulletin 143, Bureau of American Ethnology, Smithsonian
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FIELDIANA
Zoology
NEW SERIES, NO. 45
Taxonomy and Evolution
of the Sinica Group of Macaques:
6. Interspecific Comparisons and Synthesis
Jack Fooden
Research Associate
Division of Mammals
Field Museum of Natural History
Chicago, Illinois 60605-2496
Accepted for publication September 23, 1987
June 30, 1988
Publication 1389
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
© 1988 Field Museum of Natural History
Library of Congress Catalog Card Number: 87-83733
ISSN 0015-0754
PRINTED IN THE UNITED STATES OF AMERICA
For Elizabeth
Table of Contents
Abstract 1
Introduction 1
Comparisons 3
Pelage 3
External Measurements 5
Cranial Characters 5
Caudal Vertebrae 15
Glans Penis and Baculum 19
Female Reproductive Tract 24
Blood Proteins 25
Electrophoresis 25
Agglutination 28
Karyology 28
Hybridization 29
Intergeneric Hybridization 29
Intergroup Hybridization 29
Intragroup Hybridization 31
Phenotypes of Hybrids 31
Evolution and Dispersal 31
1. Origin and Early Dispersal of sinica
Group: Macaca sinica/ M. radiata ... 32
2. Origin of Macaca assamensis pelops ... 33
3. Origin of Macaca assamensis assamen-
sis 33
4. Origin of Macaca thibetana 33
5. Late Pleistocene 33
6. Holocene 34
Acknowledgments 34
Gazetteer 34
Literature Cited 39
List of Illustrations
1 . Locality records and inferred limits of
natural distribution of sinica-group ma-
caques 2
2. External characters and distribution of
5/wca-group macaques 4
3. Latitudinal variation of head and body
length in adult 5/wca-group macaques . . 7
4. Latitudinal variation of tail length in
adult 5/wca-group macaques 9
5. Tail length vs. head and body length in
immature and adult 5/wca-group ma-
caques 10
6. Skulls of sinica-group macaques, adult
males 12
7. Latitudinal variation of greatest skull
length in adult sinica-group macaques . . 13
8. Ontogenetic allometry of rostral length
vs. postrostral length in sinica-group
macaques 14
9. Mean length of vertebral centrum in
successive caudal vertebrae of sinica-
group macaques 18
10. Mean length of vertebral centrum in
successive caudal vertebrae of adult
male silenus-group macaques 18
1 1 . Male external genitalia of M. a. assa-
mensis 19
12. Radiographs of penis of M. a. assamen-
sis, dorsal and lateral views, showing
position of baculum 19
13. Bacula of subadult and adult sinica-
group macaques 21
14. Sagittal section of female reproductive
tract of M. a. assamensis 24
15. Hypothetical reconstruction of principal
stages in evolution and dispersal of sini-
ca-group macaques 32
16. Phylogenetic relationships inferred
among sinica-group macaques 34
List of Tables
1 . External measurements and proportions
in 5/wca-group macaques 6
2. Regression statistics for latitudinal vari-
ation of external measurements in sini-
ca-group macaques 8
3. Cranial measurements and proportions
in 5/wca-group macaques 11
4. Regression statistics for latitudinal vari-
ation of greatest length of skull in sini-
ca-group macaques 11
5. Ontogenetic and interspecific allometry
of rostral length relative to postrostral
length in 5/wca-group macaques 15
6. Length of centrum of caudal vertebrae
in sinica -group macaques 16
7. Male external genitalia: specimens exam-
ined and measurements of baculum .... 22
8. Blood protein electrophoresis: mono-
morphism in Macaca spp., including
5/wca-group species 25
9. Blood protein electrophoresis: monomor-
phism in sinica-group species, polymor-
phism in other species of macaques .... 26
10. Blood protein electrophoresis: dimor-
phism in 5/wca-group species 27
1 1. Blood protein electrophoresis: trimor- 13. Blood protein agglutination: human-
phism in sinica-group species 27 type blood groups in M. radiata 28
12. Blood protein electrophoresis: polymor- 14. Hybridizations reported for sinica-group
phism of plasma transferrin in sinica- species 30
group species 28
VI
Taxonomy and Evolution
of the Sinica Group of Macaques:
6. Interspecific Comparisons and Synthesis
Abstract
The sinica group of macaques comprises four
species and six subspecies: Macaca sinica (with
subspecies M. s. sinica and M. s. aurifrons), M.
radiata (M. r. radiata, M. r. diluta), M. assamensis
(M. a. assamensis, M. a. pelops), and M. thibetana.
The geographic ranges of these species are allo-
patric or parapatric and extend from Sri Lanka to
east-central China. In this paper, sinica-group
species are compared with respect to pelage, ex-
ternal measurements, cranial characters, caudal
vertebrae, glans penis and baculum, female repro-
ductive tract, blood proteins, karyology, and hy-
bridization. A hypothetical reconstruction of ma-
jor developments in the evolutionary history of
this group is proposed. New locality records of
sinica-group macaques are documented in a gaz-
etteer.
Introduction
This is the concluding part in a series of papers
that systematically review the sinica group of ma-
caques. Five previous publications in this series
present accounts of the four recognized species in
the sinica group and an overview of the natural
history of these species (Fooden, 1979, 1981, 1982,
1983, 1986). The present paper provides compar-
ative studies of external characters, skeletal char-
acters, genital characters, blood proteins, karyol-
ogy, and hybridization, and a hypothetical
reconstruction of major developments in the evo-
lutionary history of this group. A gazetteer pre-
sents details of sinica-group locality records dis-
covered subsequent to publication of previous
species accounts.
Four species and six subspecies are recognized
in the sinica group:
1. Macaca sinica (Linnaeus, 1771)
M. s. sinica (Linnaeus, 1771)
M. s. aurifrons Pocock, 1931
2. Macaca radiata (E. Geoffroy, 1812)
M. r. radiata (E. Geoffroy, 1812)
M. r. diluta Pocock, 1931
3. Macaca assamensis McClelland in Horsfield,
[1840]
M. a. assamensis McClelland in Horsfield,
[1840]
M. a. pelops Hodgson, 1841
4. Macaca thibetana A. Milne-Edwards, 1870
Taxa in the sinica group, as in other species groups
of macaques, are allopatric or parapatric (fig. 1 ;
Fooden, 1980, p. 4). Allocation of sinica-group
taxa to specific or subspecific rank therefore is
somewhat arbitrary. Plausible arguments can be
made, for example, for regarding M. a. pelops as
specifically distinct from M. a. assamensis or, con-
versely, for regarding M. sinica and M. radiata as
conspecific. However, because available evidence
is equivocal, the classification given above is re-
tained as reasonable and widely accepted.
In references to specimens cited in this paper,
institutional names are abbreviated as indicated
below:
aiuz Anthropologisches Institut der Uni-
versitat Zurich, Zurich, Switzerland
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
FIELDIANA: ZOOLOGY
amnh American Museum of Natural His-
tory, New York
bjmnh Beijing Museum of Natural History,
Beijing
bm British Museum (Natural History),
London
ecnu East China Normal University,
Shanghai
Field Museum of Natural History,
Chicago
Guangxi Institute of Forest Investi-
gation and Design, Nanning,
Guangxi Province, China
Ganzhou Zoo, Ganzhou, Jiangxi
Province, China
Hangzhou Zoo, Hangzhou, Zhejiang
Province, China
Institute of Medical Microbiology,
Zhejiang Academy of Medicine,
Hangzhou, Zhejiang Province,
China
Institut Royal des Sciences Naturelles
de Belgique, Brussels
Institute of Zoology, Chinese Acade-
my of Sciences, Beijing
Jinggangshan Nature Reserve Bureau,
Jinggangshan, Jiangxi Province,
China
Jiangxi University, Biology Depart-
ment, Nanchang, Jiangxi Province,
China
Kunming Institute of Zoology, Chinese
Academy of Sciences, Kunming,
Yunnan Province, China
Museum of Comparative Zoology,
Harvard University, Cambridge,
Mass.
Museum National d'Histoire Natu-
relle (Mammiferes), Paris
Naturhistorisches Museum, Basel,
Switzerland
National Museum, Sri Lanka, Colom-
bo, Sri Lanka
Northwest Plateau Institute of Biolo-
gy, Xining, Qinghai Province, China
Nanchang Zoo, Nanchang, Jiangxi
Province, China
Qinxidong Nature Reserve, Ruyuan,
Guangdong Province, China
Ruyuan County Forest Bureau, Ru-
yuan, Guangdong Province, China
Rijksmuseum van Natuurlijke His-
toric, Leiden, Netherlands
South China Institute of Endangered
Animals, Guangzhou, Guangdong
Province, China
smnh Shanghai Museum of Natural Histo-
ry, Shanghai
szg Shanghai Zoological Garden, Shang-
hai
usnm National Museum of Natural History,
Washington, D.C.
zmb Zoologisches Museum des Humboldt-
Universitat, Berlin
zmnh Zhejiang Museum of Natural History,
Hangzhou, Zhejiang Province,
China
zrcnus Zoological Reference Collection, Na-
tional University of Singapore, Sin-
gapore
Comparisons
Pelage
Dorsal pelage color in adults is variably brown-
ish in .szwca-group macaques (fig. 2; Fooden, 1 979,
p. 1 1 0; 1 98 1 , p. 2; 1 982, p. 6; 1 983, p. 7; Fooden
et al., 1985, p. 15). Pelage color is relatively pale
and bright in Macaca sinica and M. radiata diluta
(yellowish brown to golden brown), darker in M.
assamensis (golden brown to dark brown), and
darkest in M. thibetana (dark brown to blackish).
The grayish brown dorsal pelage in M. r. radiata
is distinctly drabber than in other s/«/ca-group
species and subspecies; this may be related to the
relative dryness of the habitat of M. r. radiata.
Erythrism occurs sporadically in M. sinica and,
apparently less commonly, in M. assamensis. Al-
binism has been reported as a rare anomaly in M.
sinica, M. radiata, and M. thibetana. Seasonal
molting has been locally documented in late spring,
at the beginning of the rainy season, in M. radiata
and M. assamensis, and in late summer, near the
end of the rainy season, in M. thibetana; seasonal
molting has not been reported in M. sinica. In-
terscapular hair length varies from about 50 mm
in M. sinica to 90 mm in M. thibetana.
Crown hairs in Macaca sinica are elongated and
radiate from a central whorl to form a conspicuous
oval cap that extends anteriorly as far as the brow
ridges; in M. s. sinica the entire cap is golden brown,
whereas in M. s. aurifrons the anterior part of the
cap is clearly defined yellowish. A conspicuous cap
also is present in M. radiata, but in this species
the anterior hairs of the cap are much shorter than
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
i
UJ
FIELDIANA: ZOOLOGY
the posterior hairs, so that the cap extends ante-
riorly only to midway between the vertex and the
brow ridges; the exposed frontal area in M. radiata
is covered with short hairs that diverge laterally
to form a median part. Crown hair arrangement
in M. assamensis is variable; in some specimens
there is a rudimentary cap centered at the vertex,
in others there is an irregular tuft or cowlick, and
in still others a whorl is absent and crown hairs
are smoothly directed posteriorly. In M. thibetana
specimens examined, a rudimentary cap is con-
sistently present. Side-whiskers and beard are rel-
atively inconspicuous in M. sinica and M. radiata,
moderately developed in M. assamensis, and
prominent in M. thibetana.
Facial skin color in 5/mctf-group adults is buffy
in adult males and variably buffy to pinkish to red
in adult females. In M. sinica, ears and lips are
blackish; in other sinica-group species, they are
buffy.
External Measurements
Body size is sexually dimorphic in sinica-group
species (table 1 ; fig. 3), as in other macaques. Length
of head and body in adult males averages 5°/o-23%
greater than in adult females, and weight averages
47%-70% greater. Sexual dimorphism in Macaca
thibetana and M. a. assamensis apparently exceeds
that in M. radiata and M. sinica; this generally
accords with previous indications that sexual di-
morphism increases with body size (Rensch, 1 960,
p. 157; Clutton-Brock et al., 1977, p. 798; Al-
brecht, 1980, p. 148). Macaca assamensis pelops,
however, is represented by a sample of five ex-
ceptionally large adult females and apparently is
the least dimorphic taxon in the sinica group.
Mean length of head and body and mean body
weight of species in the sinica group increase with
increasing latitude of the ranges of these species
(fig. 3; table 1; Fooden, 1971, p. 72). In Macaca
thibetana, the northernmost species in the group,
mean length of head and body in adults is about
30% greater than in M. sinica, the southernmost
species, and mean weight is more than 200% great-
er. This relationship between body size and lati-
tude conforms to Bergmann's rule (Mayr, 1963,
p. 320) and probably indicates that size in these
species is adapted to temperature of habitat. The
progressive increase of head and body length of
sinica-group species is gradual, with measure-
ments broadly overlapping in neighboring species.
Within szwca-group species, the relationship
between latitude and head and body length may
be analyzed by least squares linear regression (Al-
brecht, 1980, p. 144). Available data are adequate
to establish that regression of head and body length
on latitude is statistically significant for male spec-
imens of Macaca a. assamensis and for female
specimens of M. radiata (table 2). This intra taxon
trend is particularly evident in M. a. assamensis
males, known from a sample of 24 adults that span
1 5 degrees of latitude. Head and body length in
M. a. assamensis males collected in the northern
part of the subspecific range apparently exceeds
that in males of M. a. pelops and M. thibetana
collected at the same latitude.
Mean tail length of species in the sinica group
generally decreases with increase jn latitude of the
specific range (figs. 4-5; table 1), in broad agree-
ment with Allen's rule (Mayr, 1963, p. 323). The
pattern of tail length decrease, however, is not
symmetrical with the pattern of head and body
length increase. Mean tail length is approximately
equal in Macaca sinica and M. radiata, despite the
difference in latitude of their ranges. Mean tail
length then decreases successively in M. a. pelops,
M. a. assamensis, and M. thibetana, with little or
no overlap of this measurement in neighboring
species or subspecies. Within species or subspe-
cies, there is no significant tendency for tail length
to decrease with latitude (table 2; the only signif-
icant regression is in M. radiata males, where the
slope is positive). Relative tail length in immatures
in sinica-group species apparently is approxi-
mately the same as in adults (fig. 5). Although tail
reduction is sometimes associated with increased
terrestriality, this does not apply to M. assamensis,
which apparently is at least as arboreal as M. ra-
diata (Fooden, 1986, p. 3).
In Macaca sinica, M. radiata, and M. assamen-
sis, mean ear length, like mean tail length, tends
to decrease with increase in latitude of specific
ranges (table 1). Macaca thibetana, however, has
ears that are relatively large and thus departs from
the general pattern of the other three species.
Cranial Characters
Species in the sinica group differ markedly in
skull size (figs. 6-7; table 3). In available samples
of adults, mean skull length varies from 97.1 mm
in female Macaca sinica to 130.2 mm in female
M. thibetana and from 1 13.0 mm in male M. sin-
ica to 156.2 mm in male M. thibetana. Sexual
dimorphism of skull length in sinica-group species
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
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FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
Table 2.
figs. 3-4).
Regression statistics for latitudinal variation of external measurements in sinica-group macaques (cf.
Species or
subspecies
Sex
No. of No. of
specimens localities
y-intercept
Slope
se slope
P slope
M. sinica
6
9
M. radiata
6
9
M. a. pelops
6
9
M. a. assamensis
6
9
M. thibetana
6
9
M. sinica
S
9
M. radiata
6
9
M. a. pelops
6
9
M. a. assamensis
$
$
M. thibetana
6
9
21
12
12
11
8
5
24
17
6
5
22
13
12
10
8
5
22
17
5
5
Head and Body Length
10
379.6
11.69
7.85
.10-.25
9
343.4
10.29
11.46
.25-.50
9
451.8
5.52
5.51
.25-.50
8
334.0
9.75
3.50
.025-.05*
5
1,609.4
-37.83
59.85
.50-.75
2
-559.5
40.91
18
426.9
7.70
2.03
.001-.005
12
448.7
2.51
2.46
.25-.50
5
275.0
12.56
10.10
.25-.50
5
-307.1
Tail Length
29.29
14.97
.10-.25
11
676.4
-14.92
11.51
.10-.25
9
504.3
2.40
24.32
> .75
9
348.5
15.61
6.36
.025-.05*
8
491.5
0.87
11.82
> .75
6
-1,123.8
53.26
43.37
.25-.50
2
-2,162.7
89.39
16
228.2
-0.66
1.07
.50-.75
13
236.2
-1.88
1.24
.10-.25
5
164.7
-3.41
3.30
.25-.50
5
55.4
0.45
7.31
> .75
* = .05 > P > .01. ** = .01 > P > .001.
apparently is greater than sexual dimorphism of
head and body length (cf. figs. 3, 7). The progres-
sive increase of mean skull length in sinica-group
species, like the corresponding increase of mean
head and body length, is correlated with increasing
latitude of the specific ranges. Unlike head and
body length variation, however, skull length vari-
ation is not continuous between all four species.
Skull length variation in both sexes of the two
smaller species (M. sinica, M. radiata) is discon-
tinuous from that in the two larger species (M.
assamensis, M. thibetana); even at the same lati-
tude, the largest M. radiata skull in each sex is
smaller than the smallest M. assamensis skull. Skull
length tends to increase with latitude within species
as well as between species; within species or sub-
species, regression of skull length on latitude is
statistically significant in males of M. radiata, M.
a. assamensis, and M. thibetana and in females of
M. a. assamensis (table 4). Relative to head and
body length, skull length in M. thibetana is ex-
ceptionally large (cf. figs. 3, 7).
Although species in the sinica group differ in
skull size, they are remarkably similar in general
proportions (fig. 6; table 3; Albrecht, 1978, p. 76).
Relative zygomatic breadth averages approxi-
mately 0.67 in both sexes of all four species. Ros-
tral/postrostral ratio, a measure of the ratio of fa-
cial length to cranial length, increases only slightly
with increasing skull size, from 0.47 in female M.
sinica to 0.51 in female M. thibetana and from
0.55 in male M. sinica to 0.59 in male M. thibet-
ana. The two smaller species (M. sinica, M. ra-
diata) tend to differ from the two larger species
(M. assamensis, M. thibetana) in morphology of
the temporal lines and sagittal crest in adult males;
in the smaller species the temporal lines usually
are separate, whereas in the larger species the tem-
poral lines often converge to produce a prominent
sagittal crest in adult males (cf. Pocock, 1939, pp.
35, 40, 53; Kurup, 1966, p. 74). Width of the
rostrum tends to be relatively smaller in M. sinica
and M. radiata than in M. assamensis and M.
thibetana (fig. 6). No known cranial character
uniquely distinguishes sinica-group species from
those in other species groups.
Ontogenetic allometry of rostral length relative
to postrostral length apparently differs among
species in the sinica group (fig. 8; table 5). In a
composite log-log plot of rostral length against
postrostral length, data points for immature and
mature specimens are approximately collinear
FIELDIANA: ZOOLOGY
I
^<jj
a.
_o
a
(iuuj) msu»i n»i
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
700
Open symbols — females; solid symbols — males.
Larger symbols — adults; smaller symbols — immatures.
400 -
100
500 600 700
200 300 400
Head and body length (mm)
Fig. 5. Tail length vs. head and body length in immature and adult sinica -group macaques.
10
FIELDIANA: ZOOLOGY
Table 3. Cranial measurements and proportions in sinica-group macaques.
Species or
subspecies
GL
(mm)
zb/gl
Postrostral length
(mm)
rl/pl
Adult Males
M. sinica1
113.0 ± 4.2(21)
106.7-119.8
.69 ± .02 (20)
.65-.71
77.8 ± 3.0(21)
72.7-83.1
.55 ± .03(21)
.50-.60
M. radiata1
120.0 ± 4.0(12)
114.6-127.9
.67 ± .02(12)
.65-. 71
83.1 ± 2.6(12)
80.2-86.5
.54 ± .02(12)
.51-.57
M. a. pelops1
142.1 ± 6.5(11)
131.9-154.1
.66 ± .03(11)
.63-. 70
97.3 ± 3.6(10)
89.9-101.1
.57 ± .03(10)
.51-.62
M. a. assamensis2
146.9 ± 5.8 (28)
138.1-160.3
.66 ± .03 (28)
.62-.70
99.4 ± 4.0 (26)
93.9-107.8
.58 ± .04 (25)
.50-.65
M. thibetana?
156.2 ± 6.1(18)
146.1-167.5
.67 ± .02(18)
.62-.71
105.7 ± 3.2(15)
102.0-112.1
.59 ± .02(15)
.53-.62
Adult Females
M. sinica1
97.1 ± 3.6(15)
93.0-104.9
.66 ± .02(13)
.63-.69
71.3 ± 2.2(15)
68.6-75.9
.47 ± .03(15)
.43-.53
M. radial a'
104.4 ± 2.4 (10)
99.0-107.7
.65 ± .03(10)
.60-.69
76.9 ± 2.0(10)
73.4-78.6
.46 ± .02(10)
.43-.50
M. a. pelops*
121.6 ± 5.5(8)
116.0-131.5
.67 ± .03 (8)
.63-.71
88.8 ± 2.9 (7)
85.5-93.6
.49 ± .04 (7)
.45-.58
M. a. assamensis*
121.2 ± 6.2(23)
113.1-138.5
.66 ± .02 (23)
.63-.71
88.1 ± 4.3(21)
81.0-97.9
.49 ± .03(21)
.41-.55
M. thibetana*'
130.2 ± 5.3(10)
120.7-140.0
.67 ± .01 (10)
.65-.68
93.8 ± 2.6 (8)
91.0-97.3
.51 ± .02(8)
.46-.S3
ol = Greatest length of skull, excluding incisors; zb/gl = relative zygomatic breadth; rl/pl = rostral/postrostral
ratio. Mean, standard deviation, and sample size (in parentheses) reported in first line of each entry; extremes, in
second line. For explanation of measurements, see Fooden, 1969, p. 41.
1 References: Fooden, 1979, p. 1 14; 1981, p. 14; 1982, p. 14. 2 References: Fooden, 1982, p. 14 (excluding gl =
129.6, subadult); izcas, 6 specimens; kjz, 5 specimens; nwpib, 2 specimens. 3 References: Fooden et al., 1985, p. 19
(excluding 1 amnh specimen of unknown origin); izcas, 2 newly acquired specimens; sciea, 3 specimens.
4 References: Fooden, 1982, p. 14; fmnh 94089, Jiri, Nepal. 5 References: Fooden, 1982, p. 14; izcas, 2 specimens;
kiz, 8 specimens. 6 References: Fooden et al., 1985, p. 19; sciea, 1 specimen; zmnh, 1 specimen.
within each species, but data points for larger
species generally are shifted to the right of those
for smaller species. Evolutionary changes of size
in 5/mca-group species evidently have been ac-
companied by compensating transformations of
the allometric growth curves, with the result that
these species differ only slightly in rostral/post-
rostral ratios of adults. In this respect, sinica-group
Table 4. Regression statistics for latitudinal variation of greatest length of skull in sinica-group species or sub-
species (cf. fig. 7).
Species or
No. of
No. of
Y-
P
subspecies
Sex
specimens
localities
intercept
Slope
se slope
slope
M. sinica
6
21
12
104.8
1.11
1.40
.25-.50
9
15
12
84.2
1.77
0.80
.05-. 10
M. radiata
6
12
9
104.3
1.12
0.41
.025-.05*
9
10
8
108.8
-0.37
0.31
.25-.50
M. a. pelops
6
11
8
100.8
1.54
1.65
.25-.50
9
8
5
495.3
-13.72
7.22
.10-.25
M. a. assamensis
6
28
21
129.1
0.73
0.23
.005-.01**
9
23
17
103.7
0.81
0.36
.025-.05*
M. thibetana
8
18
15
115.0
1.49
0.58
.01-.025*
9
10
10
82.8
1.68
1.02
.10-.25
* - .05 > P > .01. ** = .01 > P > .001.
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
11
12
FIELDIANA: ZOOLOGY
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FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
13
60
50
40
30
20 -
Open symbols — females; solid symbols —
adults
(symbols enclosed in envelopes)
'IT" i
immatures
50
60
70 80
Postrostral length (mm)
90
100
110
Fig. 8. Ontogenetic allometry of rostral length (y) vs. postrostral length (x) in sinica-group macaques. Principal
axis equations: M. sinica, log y = 4.317 log x - 6.507; M. radiata, log y = 3.395 log x - 4.860; M. assamensis, log
y = 3.71 1 log x - 5.610; M. thibetana, log y = 3.297 log x - 4.851 (table 5).
14
FIELDIANA: ZOOLOGY
Table 5. Ontogenetic and interspecific allometry of rostral length (y) relative to postrostral length (x) in sinica-
group macaques (cf. fig. 8).
Species/sex
N
Line-fitting
technique
Slope
95% confidence
limits
y-intercept
M. sinica1
M. radiata*
M. thibetana*
Adult males
Adult females
692
452
M. assamensis3 1 1 72
432
4s
45
Ontogenetic Allometry
ma
4.317
3.83CM.937
-6.507
rma
3.861
3.417^*.305
-5.661
lsr
3.405
2.961-3.849
-4.816
ma
3.395
3.019-3.865
-4.860
rma
3.181
2.809-3.553
-4.456
lsr
2.942
2.570-3.314
-4.005
ma
3.711
3.457^.003
-5.610
rma
3.482
3.243-3.721
-5.165
lsr
3.232
2.993-3.471
-4.679
ma
3.297
3.040-3.598
-4.851
rma
3.200
2.950-3.450
-4.659
lsr
3.086
2.819-3.353
-4.434
Interspecific Allometry
ma
1.291
1.182-1.412
-0.816
rma
1.290
1.040-1.540
-0.815
lsr
1.287
1.037-1.537
-0.809
ma
1.312
1.119-1.549
-0.914
rma
1.309
0.847-1.771
-0.909
lsr
1.301
0.839-1.763
-0.893
0.882
0.925
0.928
0.964
0.998
0.993
ma = Major axis; rma = reduced major axis; lsr = least squares regression. For discussion of these techniques,
see Sokal and Rohlf (1981, p. 549) and Steudel (1985, p. 462).
1 References: Fooden, 1979, p. 115; 1981, p. II.2 Both sexes, all ages. 3 Cf. Fooden, 1982, p. 16, N = 74. 4 Cf.
Fooden, 1983, p. 11,N = 20.
5 Bivariate means for each species; see table 3.
species contrast with silenus-group species Ma-
caco, silenus and M. nemestrina, which apparently
follow a common allometric growth curve and, as
adults, exhibit conspicuous size-related differen-
tiation of rostral/postrostral ratio (Fooden, 1975,
p. 1 2). In M. silenus and M. nemestrina, interspe-
cific allometry of adults is an extension of intra-
specific growth allometry; in sinica -group species,
interspecific allometry follows a trajectory differ-
ent from that of intraspecific allometry. Macaque
species groups evidently are not isomorphic in their
patterns of rostral-postrostral evolution.
Caudal Vertebrae
Interspecific variation of caudal (Cd) vertebral
morphology is of particular interest in macaques
because tail reduction is a conspicuous evolution-
ary trend in this genus. Sets of caudal vertebrae of
sinica-group species are available for long-tailed
Macaca sinica and M. radiata, for short-tailed M.
a. assamensis, and for stump-tailed M. thibetana
(table 6). No caudal vertebral specimens are avail-
able for M. a. pelops, in which tail length is inter-
mediate between that in M. radiata and M. a.
assamensis.
In Macaca sinica and M. radiata, tail length is
approximately equal (table 1); the number of cau-
dal vertebrae is similar, averaging about 25 or 26
in both species; and lengths of corresponding cau-
dal vertebrae also are similar (table 6; Schultz &
Straus, 1 945, p. 623). In adult males of both species,
caudal vertebral length increases rapidly from
about 12 mm in Cdl to about 30 mm in Cd5,
reaches a peak of about 36 mm in Cd6-9, and then
decreases somewhat more gradually to about 5
mm in the terminal vertebra (fig. 9). In six adult
males, Cd7 is the longest caudal vertebra in three
specimens, Cd8 in two specimens, and Cd9 in one
specimen. Neural arches are present in Cdl-5,
which thus constitute the proximal caudal region
as defined by Ankel ( 1 972, p. 232); all other caudal
vertebrae lack neural arches and constitute the dis-
tal caudal region. Vertebral length reaches its max-
imum in the first section of the distal caudal region
in M. sinica and M. radiata, as in most long-tailed
mammals (Lessertisseur & Saban, 1967, p. 632).
In adult females and in immatures of both sexes,
the number of caudal vertebrae and the vertebral
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
15
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FIELDIANA: ZOOLOGY
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FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
17
M. radiata
5 10 15
Caudal vertebra number
Fig. 9. Mean length of vertebral centrum in succes-
sive caudal vertebrae of 5/n/ca-group macaques: adult
male M. sinica, M. radiata, M. a. assamensis, and M.
thibetana; fetal male M. a. assamensis (table 6).
length gradient apparently are similar to those in
adult males.
In Macaca a. assamensis, caudal vertebrae are
reduced both in number and length relative to
those in M. sinica and M. radiata; the morphology
of corresponding vertebrae is generally similar,
however, in all three species. The number of cau-
dal vertebrae in five specimens of M. a. assamensis
is 17-19 (table 6; Schultz, 1938, p. 6). In adult
males, vertebral length decreases slightly from
about 12 mm in Cdl to about 10 mm in Cd2 and
Cd3, increases rapidly to about 18 mm in Cd6,
reaches a peak of about 2 1 mm in Cd7-9, and
decreases somewhat more gradually to about 2
mm in the terminal vertebra. In a near-term fetus
of M. a. assamensis, the vertebral length gradient
characteristic of this subspecies is already apparent
(fig. 9).
In Macaca thibetana, the number of caudal ver-
tebrae is reduced to 10-12 (table 6). Vertebral
35
30
E
£ 25
g 20
15
10
M. silenus \
/ X
/ ^
_' n
S-s_s /**. nemestrina »
IV n
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: *&&\
— M. nigra \ L
\ \
brunnescens n s
n \
V" Ns-s
\ M. maurus
m-m
\
■ ■ i i I i i i i i ■ ■ i i i i i i i i i i
5 10 15 20
Caudal vertebra number
Fig. 10. Mean length of vertebral centrum in suc-
cessive caudal vertebrae of adult male silenus-group ma-
caques (Fooden, 1969, p. 18; 1975, p. 20).
length in adult males decreases irregularly from
about 13 mm in Cdl to about 10 mm in Cd7 and
then decreases more abruptly to about 3 mm in
the terminal vertebra.
The caudal vertebral length gradient in Macaca
sinica and M. radiata probably is primitive in this
species group. In M. a. assamensis, peak vertebral
length in the first section of the distal caudal region
is reduced by about one-third; in M. a. pelops, the
reduction in this region presumably is somewhat
less. Although the length of Cdl is approximately
the same in M. a. assamensis as in M. sinica and
M. radiata, the length of Cd2 and Cd3 in M. a.
assamensis is reduced. Beginning at Cd4, each cau-
dal vertebra in adult male M. a. assamensis av-
erages 10-16 mm shorter than the corresponding
vertebra in adult male M. sinica and M. radiata.
In M. thibetana, the length of vertebrae in the first
section of the distal caudal region is reduced by
about one-half, relative to M. a. assamensis; this
reduction completely eliminates the length peak
that characterizes this caudal vertebral section in
M. sinica, M. radiata, and M. a. assamensis. The
FIELDIANA: ZOOLOGY
Fig. 1 1. Male external genitalia of M. a. assamensis (fmnh 99622, adult; Ban Muang Baw Ngam, Kanchanaburi
Province, Thailand).
length of Cdl in M. thibetana remains about the
same as in other species in the group. Beginning
at Cd7, each caudal vertebra in adult male M.
thibetana averages 1 0- 1 6 mm shorter than the cor-
responding vertebra in adult male M. a. assamen-
sis; this parallels the relationship noted above be-
tween M. sinicalM. radiata and M. a. assamensis.
The caudal vertebral length gradient in stump-
tailed M. thibetana clearly is not a paedomorphic
retention of the fetal gradient in short-tailed M. a.
assamensis (fig. 9). Caudal vertebral length gra-
dients in the sinica group are generally similar to
those in the silenus group (fig. 10) and, judging
from data available for M. fascicularis and M. mu-
latto, also to those in the fascicularis group (Ankel,
1962, p. 156; Wilson, 1970, pp. 196-197).
Glans Penis and Baculum
The form of the glans penis in the sinica group
is highly distinctive (fig. 11; Cuvier, 1820, p. 1;
Cuvier, 1846, p. 220; De Beaux, 1917, p. 6; Po-
cock, 1921, p. 228; Hill & Bernstein, 1969, p. 6;
Fooden, 1971, p. 72). In this group, the glans is
strongly inflected relative to the shaft of the penis,
the dorsal margin of the corona is thickened and
reflected anteriorly, the subterminal urethral mea-
tus opens anterodorsally, and the apex of the glans
is subacute. In other macaques, except Macaca
arctoides, the glans is only slightly inflected rela-
tive to the shaft, the corona is relatively simple,
the urethral meatus is terminal, and the apex of
the glans is bilobed and bluntly rounded, approx-
imately as in humans and most catarrhine mon-
keys (Pocock [1926], p. 1557; Hill, 1958, p. 650;
Fooden, 1975, p. 33); in aberrant M. arctoides, the
glans is more than twice as long as in the sinica
group and the urethral meatus opens ventral to
the apex of the glans (Fooden et al., 1985, p. 18).
Although difficult to measure, the glans and shaft
of the penis in the sinica group also seem to be
relatively larger than in most other macaques;
judging from specimens examined, the do rso ven-
tral diameter of the distal part of the shaft is about
50% greater in 5/mca-group species than in other
Fig. 1 2. Radiographs of penis of M. a. assamensis,
dorsal and lateral views, showing position of baculum
(fmnh 99632, subadult; Ban Pong Nam Ron, ca. 25 km
W, Kamphaeng Phet Province, Thailand). (Radiographs
courtesy Chicago Zoological Park, Brookfield, Illinois.)
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
19
macaque species of corresponding head and body
length. Glans morphology in the sinica group ev-
idently is derived relative to that in most ma-
caques.
The thickened dorsal margin of the corona in
the sinica group forms a horseshoe-shaped swell-
ing that surrounds almost half of the glans. Be-
tween this swelling and the dorsal end of the ure-
thral meatus is a well-defined semicircular
concavity. The urethral meatus is a dorsoventrally
oriented slit that extends about one-third the length
of the glans and terminates dorsal to the apex of
the glans. The left lip of the meatus is about twice
as thick as the right lip and contains the distal
inflected process of the baculum (fig. 1 2). Proximal
to the glans, the skin of the distal part of the shaft
of the penis is densely studded with prominent
spines, the tips of which are recurved toward the
base of the shaft. The length of these spines ranges
up to about 0.5 mm in fluid-preserved adult spec-
imens of Macaca sinica and up to about 0.7 mm
in similarly preserved adult specimens of M. a.
assamensis; the basal diameter of the spines is
about half of their length. Spines also are present
on the margin of the corona, particularly dorsally,
where they cover about half of the horseshoe-
shaped swelling. The color of the glans is pinkish
in living M. sinica, M. radiata, and M. a. assa-
mensis and buffy in living M. thibetana.
Size of the glans in sinica-group species appar-
ently is approximately proportional to head and
body length. Measurements in millimeters of glans
length (apex to middorsal margin of corona) and
breadth are 17.5 x 10.5 and 18.5 x 11.0 in two
fluid-preserved adult specimens of Macaca sinica
(fmnh 57720, 57721), 22.5 x 16.0 and 22.5 x
16.5 in two fluid-preserved adult specimens of M.
a. assamensis (fmnh 99622, 99631) (cf. Hill &
Bernstein, 1969, p. 7); measurements in one living
adult specimen of M. thibetana are 25 x 20 (Food-
en et al., 1 985, p. 1 9). These measurements suggest
that relative breadth of the glans may be greater
in the larger species.
Variation of form of the glans within and be-
tween species is relatively minor. The distinctive
form is readily recognized even in infants less than
one year old (prior to eruption of the first per-
manent teeth). However, in one specimen of M.
sinica (fmnh 57723, ?adult), the glans is abnormal.
The meatal cleft in this specimen is prolonged
ventrally and extends through the ventral border
of the corona of the glans to the right of the apex.
This extension of the meatal cleft subdivides the
distal end of the glans into two lobes, the left lobe
larger than the right, which brings the form of this
part of the glans somewhat closer to that in most
non-s/m'ctf-group species of macaques. However,
the horseshoe-shaped dorsal swelling and semi-
circular concavity in this specimen are as in typical
sinica-group specimens.
The baculum, which provides skeletal support
for the glans, is stocking-shaped and bilaterally
flattened in sinica-group macaques (figs. 12-13;
Daubenton, 1766, p. 306; De Beaux, 1917, p. 6;
Chaine [1927], p. 15; Pohl, 1928, p. 102; Fooden
[1966], p. 160). The shaft of the baculum is rooted
in the corpora cavernosa of the penis. The distal
inflected process, variably subdivided into a dorsal
and ventral lobe, projects into the left lip of the
urethral meatus, where it terminates to the left of
the navicular fossa near the ventral end of the
meatal cleft; the baculum does not extend into the
apex of the glans (fig. 1 2).
Bacular size in sinica-group species is roughly
proportional to body size (tables 1, 7). Bacular
length averages 12.2% of mean skull length in four
adult specimens of Macaca sinica, 16.6% of skull
length in four adult specimens of At. a. assamensis,
and 16.0% of skull length in two adult specimens
of At. thibetana. Length of the distal inflected pro-
cess relative to length of the shaft averages greater
in M. sinica and At. radiata than in M. a. assa-
mensis and At. thibetana. Size and form of the
baculum in At. a. assamensis and M. thibetana
tend toward those in M. nemestrina leonina
(Fooden, 1975, p. 41).
Bacular variation within and between species in
the sinica group appears to be greater than vari-
ation of glans morphology (see above). A parallel
situation previously was noted in the silenus group,
where subspecies Macaca n. nemestrina and M.
n. leonina are similar in glans morphology but
differ in bacular morphology (Fooden, 1975, p.
38). Three bacula examined exhibit special pe-
culiarities: M. radiata zmb 1 24 (?adult) has a large
fossa on the left side of the shaft of the baculum
immediately proximal to the distal inflected pro-
cess (fig. 1 3, M. radiata: b); M. a. assamensis fmnh
99622 (adult), has an exceptionally short shaft (fig.
13, M. assamensis: a); and At. thibetana amnh
84475 (infant) has a distal inflected process which
curves gradually into the shaft instead of being set
off at an abrupt angle.
The functional relationship between specialized
morphology of the penis in the sinica group and
specialized morphology of the female tract (see
below) is unknown. Copulatory behavior in this
group has been reported in detail only for Macaca
20
FIELDIANA: ZOOLOGY
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
21
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FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
23
Fig. 14. Sagittal section of female reproductive tract of M. a. assamensis (fmnh 99628, adult; Ban Mae Lamao,
Tak Province, Thailand). (Photo by John Bayalis, Division of Photography, Field Museum of Natural History.)
radiata, in which ejaculation usually is accom-
plished in a single mount with an average of 17
thrusts and a mean total duration of 10 seconds
(Shively et al., 1982, p. 376). The greater variation
of bacular morphology than of glans morphology
suggests that glans morphology is subject to more
rigorous selection pressure.
Female Reproductive Tract
Cyclical estrous swelling of the sexual skin in
sinica-group females is relatively modest (Zuck-
erman, 1930, p. 705; Hartman, 1938, p. 468; Hill,
1939, p. 25; Fooden, 1971, p. 63; Dittus, 1974,
chap. 1, p. 52; McArthur et al., 1972, p. 1 18; Hill,
1974, pp. 697, 729; Fooden et al., 1985, p. 23),
compared with that in silenus-group females
(Fooden, 1969, p. 13; 1975, p. 28). In Macaca
sinica and M. radiata, slight swelling of the sub-
caudal, circumanal, or labial area has been ob-
served occasionally; in M. thibetana, swelling and
reddening of the perineal region apparently are
common in estrous females (Xiong, 1984, p. 6);
no information is available concerning estrous
swelling in M. assamensis. During pregnancy, a
subcaudal swelling has been noted in M. radiata
and M. assamensis. Gray, blue, or purple color-
ation of the sexual skin has been noted in all four
species in this group: in M. sinica, a dark gray or
bright purple color that extends from tail root to
labia develops with age in adult females and may
persist through the entire menstrual cycle; in M.
radiata, the circumanal area is dark purple in non-
pregnant females and becomes even darker during
pregnancy; in M. assamensis, a bluish perineal
streak has been reported in a pregnant female, and
a dark blue circumanal triangle has been reported
in a lactating female; and in M. thibetana, a bluish
perineal streak has been reported in one nonpreg-
nant female.
The vaginal lining is distinctively spinose in two
nonpregnant nonlactating female specimens of
Macaca a. assamensis (fig. 14; Fooden, 1971, p.
67). Similar coarse spines are present in the vaginal
lining of one nonpregnant female specimen of M.
thibetana (Fooden et al., 1985, p. 22). No infor-
mation is available concerning presence or ab-
sence of vaginal spines in M. sinica or M. radiata.
Cyclical desquamation of cells from the vaginal
lining in M. radiata is only about 10% of that in
M. mulatta (Hartman, 1938, p. 473).
The uterine cervix in sinica -group species is
greatly enlarged (fig. 14; Zuckerman, 1930, p. 704;
Hartman, 1938, p. 473; Hill, 1939, p. 28; Fooden,
1971, p. 67; Ovadia et al., 1971, p. 128; Jainudeen
et al., 1972, p. 471; Fooden et al., 1985, p. 23). In
nonpregnant nonlactating adults, the interdigitat-
ing dorsal and ventral colliculi, which partly ob-
struct the cervical canal, are more than twice as
large in sinica-group species than in other ma-
caque species groups (Fooden, 1980, p. 3). The
24
FIELDIANA: ZOOLOGY
Table 8. Blood protein electrophoresis: monomorphism in Macaca spp., including s/'mca-group species (not
studied in M. thibetana).
No.
of specimens studied
Monomorphic
protein locus
M. sinica
M. radiata
M.
assamensis
Other species of Macaca
Ref. nos.
:
Plasma Proteins
Alp
131
22
28
2,106'
(13 spp.)
2-9
Amy
131
19
28
2,062
(13 spp.)
3-9
Cat
131
19
28
1,397
(5 spp.)
3-4,7
LAP
131
19
28
2,127'°
(13 spp.)
2-9
PA
0
19
28
1,807
(6 spp.)
3-6
a2
131
19
28
2,062"
(13 spp.)
3-9
Erythrocyte Proteins
g6pd
131
19
28
2,128
(13 spp.)
2-9
PGI
0
1
0
28
(5 spp.)
2
TO
131
19
28
2,062
(13 spp.)
3-9
Alp = Alkaline phosphatase; Amy = amylase; Cat = catalase; lap = leucine aminopeptidase; pa prealbumin; a2 =
slow a,-macroglobulin; g6pd - glucose-6-phosphate dehydrogenase; pgi = phosphoglucoisomerase; to = tetrazolium
oxidase.
1 Variant allele in 3 of 477 M. fascicularis specimens and in 2 of 255 Sulawesi macaque specimens.
2 Bruce, 1977, pp. 144, 152, 157, 162. 3 Nozawa et al., 1977, pp. 16, 18, 22. 4 Shotake, 1979, pp. 444, 447^*48.
5 Kawamoto et al., 1981, pp. 18, 20. 6 Kawamoto et al., 1982, p. 58. 7 Shotake and Santiapillai, 1982, pp. 81-82.
8 Kawamoto and Suryobroto, 1985, p. 35. " Kawamoto et al., 1985, pp. 46, 49.
10 Variant allele in 1 of 29 M. arctoides specimens and in 1 of 255 Sulawesi macaque specimens. " Protein absent
in 2 of 1 ,063 M. fuscata specimens.
highly developed endocervical epithelium in sin-
/ca-group species is richly glandular; regulated by
ovarian hormones, this epithelium secretes large
quantities of mucus (Percy, 1844, p. 83; Xiong,
1984, p. 6) that apparently functions as a sex pher-
omone (Rahaman & Parthasarathy, 1971, p. 98;
Fooden, 1981, p. 29).
Recent histological study of the ovary of Ma-
caca radiata has revealed that the preovulatory
Graafian follicle in this species is remarkably dif-
ferent from that in most mammals, including M.
fascicularis and M. mulatta (Barnes et al., 1978,
p. 538). Unlike the smooth-walled spherical pre-
ovulatory follicle that is typical of mammals, the
follicle in M. radiata has walls that are deeply
folded, giving the follicle a collapsed appearance.
The form of the preovulatory follicle in other sin-
/ca-group species has not been reported.
Blood Proteins
Electrophoresis— Thirty-seven blood protein
loci have been investigated electrophoretically in
three 5/mca-group species and in other macaques
(tables 8-12). The three sinica-group species that
have been studied are Macaca sinica (32 loci), M.
radiata (35 loci), and M. assamensis (30 loci). Blood
proteins in M. thibetana have not been studied.
Comprehensive analyses of electrophoretic evi-
dence from adequate samples of sinica-gxoup
specimens agree with previous determinations,
originally based on reproductive tract morpholo-
gy, that species in the sinica group are more closely
related to each other than to other species of ma-
caques (Darga et al., 1975, p. 803; Shotake, 1979,
p. 447; Melnick & Kidd, 1985, p. 138). Although
preliminary study of one specimen of Macaca as-
samensis appeared to indicate that this species was
serologically closer to M. mulatta than to M. sinica
and M. radiata (Cronin & Meikle, 1979, p. 262;
Cronin et al., 1980, pp. 44, 46; Cann et al., 1979,
p. 425; Pope & Cronin, 1984, p. 384), subsequent
study of 28 specimens of M. assamensis has es-
tablished that the serologic distance of this species
from M. mulatta is about three times as great as
its distance from M. radiata (Shotake, 1979, p.
447). Because of the possibility of convergent evo-
lution of blood proteins, study of a small number
of loci may not detect the close interrelationship
of 5/moz-group species that is revealed by more
comprehensive analysis. A recent study of trans-
ferrin allele frequencies in macaques indicates that,
with respect to alleles at this locus, M. sinica is
convergently similar to M. mulatta, and M. ra-
diata is convergently similar to M. cyclopis (Ha-
zoutetal., 1986, p. 245).
Of the 37 blood protein loci that have been stud-
ied in s/rt/ca-group species and in other macaques,
nine loci (6 plasma proteins, 3 erythrocyte pro-
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
25
Table 9. Blood protein electrophoresis: monomor-
phism in sinica-group species (not studied in M. thibet-
ana), polymorphism in other species of macaques.1
No. of sinica-group
specimens studied
M. M.
M. radi- assa-
sinica ata mensis Ref. nos.
Monomorphic
protein locus
CP
GC1
Hp
pi
AcP
CA-II
ca-i control
Cell Es9
EsD1
GOT
Hb-0
LDH-A
MDH
6-PGD
Plasma Proteins
131 22
131 0
131 28
131 196
28
0
28
28
Erythrocyte Proteins
131
0
55
131
131
0
202
131
131
188
19
22
52
19
0
26
6810
35
44
89"
28
0
0
28
0
0
28
28
28
28
2-5
5
2-5
3-5
3-5
7
8
3-5
5
2
4-5,8
2-5
2-5
2-5,8
cp = Ceruloplasmin; gc = group-specific component;
Hp = haptoglobin; pi = protease inhibitor; AcP = acid
phosphatase; ca-ii = carbonic anhydrase II; ca-i con-
trol = carbonic anhydrase I control; Cell Es = cell es-
terase; EsD = esterase D; got = glutamate oxalate
transaminase; Hb-/3 = hemoglobin beta; ldh-a =
lactate dehydrogenase A; mdh = malate dehydrogenase;
6-pgd = 6-phosphogluconate dehydrogenase.
1 Two proteins listed here, gc and EsD, have been
studied only in M. sinica.
2 Bruce, 1977, pp. 145, 149-150, 155, 158, 161.
3 Nozawaetal., 1977, pp. 16, 19, 22-23. 4 Shotake, 1979,
pp. 444, 447-448. 5 Shotake and Santiapillai, 1982, pp.
81-82.
6 Nozawa (ref. 3) and Shotake (ref. 4) indicate that the
pi allele frequency in M. radiata is 1.00 C (N = 19).
However, Lucotte et al. (1984, p. 340) indicate that the
pi allele frequency in this species is 0.96 B (N = 96). The
explanation for this discrepancy is unclear.
7Weissetal., 1973, pp. 214, 219. 8 Dargaetal., 1975,
pp. 800, 803-804. 9 Cf. Bruce, 1977, pp. 33, 159-160.
10 Cf. Bruce (1977, pp. 25, 146) and Ahaley et al. (1978,
p. 52).
" Excludes 1 specimen with variant allele (Bruce, 1977,
pp. 31, 155).
teins) are monomorphic in all 1 6 macaque species
investigated, including hi. sinica, M. radiata, and
hi. assamensis (table 8). Some or all of these
monomorphic alleles may be genus-level (or higher
category) taxonomic characters. Fourteen blood
protein loci (4 plasma proteins, 10 erythrocyte
proteins) are monomorphic in all three sinica-group
species that have been studied, but are polymor-
phic between or within other macaque species (ta-
ble 9). Some of these alleles may be species-group
characters. Fourteen blood protein loci (4 plasma
proteins, 10 erythrocyte proteins) are polymorphic
within the sinica group; 1 0 of these loci are di-
morphic (table 10), three are trimorphic (table 1 1),
and one (transferrin) exhibits seven alleles in sin-
ica-group species (table 12).
Judging from evidence available for sinica-group
macaques, it appears that plasma proteins may
vary more at higher taxonomic levels and eryth-
rocyte proteins may vary more at lower taxonomic
levels (cf. Palmour et al., 1 980, p. 806). Local vari-
ation of polymorphic blood protein allele fre-
quencies in natural populations of Macaca sinica
has been investigated in detail by Shotake and
Santiapillai (1982, p. 82).
The pattern of interspecific variation of allele
frequencies at polymorphic blood protein loci in
Macaca sinica, M. radiata, and hi. assamensis de-
viates from the pattern of variation of external
and cranial morphology in these species. Mor-
phological variation (head and body length, tail
length, skull length; figs. ?>-$, 7) follows a consis-
tent gradient from hi. sinica through hi. radiata
to hi. assamensis, and the morphological distance
from M. sinica to hi. radiata is consistently less
than the distance from hi. radiata to hi. assa-
mensis. The sequence of species and the relative
interspecific distances in these morphological gra-
dients exactly parallel the geographic interrela-
tionships of these species (geographic range of hi.
sinica at one extreme and geographic range of hi.
assamensis at the other extreme; range of hi. ra-
diata nearer that of hi. sinica than that of hi. as-
samensis; fig. 1). Variation of blood protein allele
frequencies does not conform to this pattern, either
with respect to the sequence of species or with
respect to relative interspecific distances.
There is no tendency for blood protein allele
frequencies in hi. radiata to be intermediate be-
tween those in hi. sinica and hi. assamensis (com-
parisons and abbreviations in tables 10-12). For
example, of five dimorphic loci at which allele
frequencies in hi. radiata differ from those in hi.
sinica and M. assamensis, the frequency in hi.
radiata is intermediate between that in hi. sinica
and hi. assamensis at two loci (tbpa, Hb-a) and
is not intermediate at three loci (ada, idh, pgm-i).
Allele frequency differences between hi. sinica and
hi. radiata may be compared with those between
hi. radiata and hi. assamensis for nine of the di-
morphic loci (all except ak, which has not been
studied in hi. assamensis): at three loci (Dia, idh,
ldh-b), the allele frequency difference between hi.
26
FIELDIANA: ZOOLOGY
Table 1 0. Blood protein electrophoresis: dimorphism in sinica-group species (not studied in M. thibetana).
Dimorphic
protein locus
Major
allele
Minor
allele
Frequency of major allele (sample size in parentheses)
M. sinica
M. radiata
M. assamensis
Ref. nos.
Plasma Protein
0.287' (196) 0.6612 (171)
Erythrocyte Proteins
0.966 (28)
3-6
ADA7
1
3
0.996 (131)
0.526 (19)
0.639(18)
4-5
AK
1
3
0.965 (131)
1.000 (14)
... (0)
5,8
CA-I
c
A
0.6679 (186)
1.000 (71)
1.000(28)
3-5
Dia
A
C
1.000 (131)
1.000 (21)
0.722(28)
4-5,8
Hb-a10
1"
2"
0.947'2(202)
0.736l3(68)
0.618(28)
3-5
IDH
1
2
0.902 (131)
0.963,4(27)
0.471(28)
4-5,8
LDH-B
1
3
1.000 (131)
1.000 (35)
0.912(28)
4-5,8
PGM-I7
1
5
1.000 (131)
0.773,5(22)
1.000(28)
4-5
PGM-H7
1
7
0.677 (131)
1.000 (22)
1.000(28)
4-5,8
tbpa = Thyroxine-binding prealbumin; ada = adenosine deaminase; ak = adenylate kinase; ca-i = carbonic
anhydrase I; Dia = nadh diaphorase; Hb-a = hemoglobin alpha; idh = isocitrate dehydrogenase; ldh-b = lactate
dehydrogenase B; pgm-i = phosphoglucomutase I; pgm-ii = phosphoglucomutase II.
1 Weighted mean of 0.385 (N = 65; ref. 3) and 0.2385 (N = 131; ref. 5). 2 Weighted mean of 0.705 (N = 56; ref.
3), 0.6842 (N = 19; ref. 4), and 0.63 (N = 96; ref. 6).
3 Darga et al., 1975, pp. 800, 802, 804. 4 Shotake, 1979, pp. 444, 448. 5 Shotake and Santiapillai, 1982, pp. 83,
89, 91.6Lucotteetal., 1984, p. 340. 7 Cf. Palmour et al., 1980, pp. 800, 805. 8 Bruce, 1977, pp. 141, 147-148, 151,
153-154.
• Weighted mean of 0.510 (N = 55; ref. 3) and 0.7323 (N = 131; ref. 5). ,0 Cf. Bruce (1977, pp. 25, 146), Ahaley
et al. (1978, p. 52), and Matsuda (1985, p. 360).
1 ' Major allele designated H and minor allele designated M by Darga et al. (ref. 3).
12 Weighted mean of 0.951 (N = 71; ref. 3) and 0.9449 (N = 131; ref. 5). ,3 Weighted mean of 0.725 (N = 49; ref.
3) and 0.7632 (N = 19; ref. 4). M Weighted mean of 1.000 (N = 19; ref. 4) and 0.875 (N = 8; ref. 8). ,5 Weighted
mean of 0.737 (N = 19; ref. 4) and 1.000 (N = 3; ref. 8).
sinica and M. radiata is less than that between M.
radiata and M. assamensis; at five loci (tbpa, ada,
ca-i, Hb-a, pgm-ii), the difference between M. sin-
ica and M. radiata is greater; and at one locus
(pgm-i), the difference is equal. For these nine di-
morphic loci, the mean allele frequency difference
between M. sinica and M. radiata is 0.222 ± 0. 1 70
(sd) and that between M. radiata and M. assa-
mensis is 0. 1 80 ± 0. 1 6 1 . The sequence and dis-
tance of allele frequency variations at the tri-
morphic loci (Alb, Ch-Es, phi) and the polymorphic
transferrin locus exhibit the same lack of con-
cordance with the sequence and distance of mor-
phological variation (tables 11-12).
Blood protein allele frequencies in the sinica
group evidently have evolved independently of
external and cranial morphology (cf. King & Wil-
son, 1975, p. 114). Hazout et al. (1984, p. 346)
have suggested that blood protein allele frequen-
cies are partly determined by natural selection in
response to climatic and geographic factors. Part
of the allele frequency divergence of M. sinica may
be a consequence of insular genetic drift (cf. Pry-
chodko et al., 1969, p. 105; Nozawa et al., 1977,
p. 26).
Table 1 1 . Blood protein electrophoresis: trimor-
phism in sinica-gjroup species (not studied in M. thibe-
tana).
Allele frequencies
Alleles
M. sinica'
(N = 131)
M. radiata1
(N = 19)
M.
assamensis2
(N = 28)
Alb
A
B
D'
0
0.980
0.020
0
1.0003
0
Ch-Es
0.161
0.839
0
1
4
5
0.988
0
0.012
0.808
0.192
0
PHI
1.000
0
0
1
15
16
0.752
0.228
0.020
1.000
0
0
1.000
0
0
Alb = Plasma albumin; Ch-Es = plasma cholinester-
ase; phi = cell phosphohexoseisomerase.
1 Reference: Shotake and Santiapillai, 1982, p. 83.
2 Reference: Shotake, 1979, p. 448. 3 N = ca. 50; Shotake
(ref. 2)— 19 specimens; Bruce, 1977, p. 156—5 speci-
mens; Weiss et al., 1973, p. 214—20-30 specimens (es-
timate).
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
27
Table 12. Blood protein electrophoresis: polymorphism of plasma transferrin (Tf) in sinica-group species (not
studied in M. thibetana).
N
Tf allele frequencies
Ref. nos.
B
C
D
E
F
F"
G
M. sinica
1
69
.058
.246
.051
.283
.022
.007
.333
2
131
.181
.185
0
.291
0
0
.343
3
39
.08
.30
.06
.19
0
0
.37
Means
239
.129
.221
.025
M. radiata?
.272
.006
.002
.345
1
59
0
.568
0
0
.407
0
.025
3
51
.14
.59
0
.01
.26
0
.01
5
19
0
.921
0
.079
0
0
0
Means
129
.054
.628
0
M. assamensis
.015
.288
0
.015
5
28
0
.146
.146
.708
0
0
0
6
7
.071
.214
.571
0
.071
0
.071
Means
35
.014
.160
.231
.567
.014
0
.014
1 Darga et al., 1975, p. 801. 2 Shotake and Santiapillai, 1982, p. 83; note that allele E is designated Di in this
study (see p. 82). 3 Hazout et al., 1986, p. 244; cf. Lucotte et al., 1984, p. 340. 4 Cf. Devor, 1977, p. 127. 5 Shotake,
1979, pp. 444, 448. 6 Annenkov, 1974, pp. 60, 62; in this work, allele F of other authors apparently is designated as
F.
Agglutination— Macaca radiata is the only
sinica-group species in which blood group agglu-
tination has been investigated. Two studies of hu-
man-type blood groups indicate that groups A, B,
and AB are all fairly common in M. radiata and
that group O is rare or absent. One study suggests
that M. radiata is monomorphic for group M in
the M-N series (table 1 3).
In a study of simian-type blood groups, eryth-
rocytes of 52 specimens of Macaca radiata were
tested for agglutinogens by using isoimmune sera
of 10 rhesus monkeys (M. mulatto) (Socha et al.,
1976, p. 489; Moor-Jankowski & Socha, 1978, p.
139). Unlike erythrocytes of some other macaque
species, erythrocytes of M. radiata were either uni-
formly positive (5 sera) or uniformly negative (5
sera) when tested with the isoimmune rhesus sera.
A similar monomorphic response previously had
been obtained when erythrocytes of six M. radiata
specimens were tested with one rhesus isoimmune
serum (LaSalle, 1969, p. 127). Intraspecific cross-
testing of erythrocytes and sera from a series of
M. radiata specimens yielded results that were
mostly negative, but responses to two sera were
polymorphic (Socha & Ruffle, 1983, p. 168). Ad-
ditional agglutination studies of other species in
the sinica group will be required in order to eval-
uate the possible systematic significance of hu-
man-type and simian-type blood group characters
in this species group.
Karyology
Classically stained karyotypes are known for
Macaca sinica, M. radiata, and M. assamensis
(Ardito, 1979, pp. 255-258). Banded karyotypes
are known for M. radiata (Stanyon, 1982, p. 72;
Table 1 3. Blood protein agglutination: human-type blood groups in Macaca radiata (not studied in other sinica-
group species).
References
N
Blood group
frequencies
52
25
77
25
O
A
B
AB
Socha & Ruffle, 1983, p. 47
More & Banerjee, 1979, p. 1331
Means
0
0
0
M
0.40
0.44
0.42
N
0.27
0.56
0.36
MN
0.33
0
0.22
More & Banerjee, 1979, p. 1331
1.00
0
0
28
FIELDIANA: ZOOLOGY
1983, p. 58; Brown et al., 1984, p. si 4; 1986, p.
168;Krishna-Murthyetal., 1984a, p. 195; 1984b,
p. 179), M. assamensis (Chen et al., 1980, p. 92;
1981, p. 37; Cao et al., 1981, p. 120), and M.
thibetana (Chen et al., 1981, pp. 92-1 15; Chen &
Luo, 1985, p. 83). The diploid chromosome num-
ber is 42 in all macaques, including sinica-group
species. Chromosome number and morphology are
remarkably similar in Macaca, Papio, Theropi-
thecus, and Cercocebus (Chiarelli, 1966, p. 168;
Dutrillauxetal., 1982, p. 100; Muleris et al., 1986,
p. 40).
Based on classically stained karyotypes, Schma-
ger (1972, p. 481) analyzed chromosome lengths
in sinica-group species and other macaques. The
reported morphometric karyological similarities
generally do not agree with relationships indicated
by nonkaryological evidence; for example, chro-
mosome length unites Macaca sinica, M. radiata,
and M. silenus in one group and separates these
species from another group that includes M. as-
samensis and M. nemestrina (cf. Fooden, 1980, p.
7). Banded karyotypes of M. radiata, M. assa-
mensis, and M. thibetana reportedly are generally
similar to those of other macaque species. No di-
rect comparison has been made of the banded
karyotypes of these three sinica-group species.
Hybridization
Species in the sinica group have been reported
as participants in 15 hybrid matings, all in cap-
tivity (table 14). Of these matings, one— of ques-
tionable reliability— is intergeneric, nine are with
macaques in other species groups (intergroup), and
five are with other species in the sinica group (in-
tragroup).
Intergeneric Hybridization— The question-
able intergeneric record is based on ambiguous
evidence of infantile pelage and skin color in a
male offspring born to a Cercopithecus aethiops
female (Gunning, 1910, p. 54; Gray, 1972, pp. 6,
1 1 , listed four times under various specific names;
Chiarelli, 1973, p. 301, listed twice; Hill, 1974, p.
470, listed four times). More than six months prior
to birth of the infant, the C aethiops female had
been caged with a M. radiata male. No informa-
tion is available as to whether other male monkeys
may also have had access to this female. Paternity
of the infant is suspect.
Successful hybridization between Cercopithecus
and Macaca would be surprising because these
genera belong to karyologically divergent subgroups
in the subfamily Cercopithecinae (Ardito, 1979,
p. 25 1 ; Chiarelli, 1 979, p. 28; Bernstein & Gordon,
1980, pp. 138, 145). In one of these subgroups
(Cercopithecus, Erythrocebus) the chromosome
number is 2n = 48-72, whereas in the other (Ma-
caca, Cercocebus, Papio, Theropithecus) the chro-
mosome number is 2n = 42.
Two other reports of hybridization between the
karyologically divergent subgroups, in addition to
the questionable Cercopithecus aethiops x Ma-
caca radiata record cited above, are listed in Gray's
(1972) checklist of mammalian hybrids, but both
of these reports also are suspect. Gray's tentative
record of hybridization ("presumed hybrid") be-
tween Cercopithecus sabaeus and Macaca mulatto
(p. 1 1 ; also listed as C aethiops x M. mulatta, p.
6) is cited from Zuckerman (1931, p. 338; 1933,
p. 96; 1953, p. 942); Zuckerman himself charac-
terizes this record as "supposed" (1931), "doubt-
ful" (1933), and "uncertain" (1953). Gray's record
of hybridization between a Cercocebus torquatus
female and a Cercopithecus mitis male (p. 5) is
cited from Montagu (1950, p. 150) and Chiarelli
(1961, table 1 ; secondary source). No such inter-
generic cross is listed by Montagu. Gray and Chi-
arelli appear to have misinterpreted a hybridiza-
tion record, explicitly labeled "Interspecific", that
Montagu lists as "Cercocebus aethiops 9 x Cer-
cocebus mitas <5"; this evidently is a lapsus for
Cercopithecus aethiops 9 x Cercopithecus mitis 6
(interspecific not intergeneric). No known record
reliably documents hybridization between the 48-
72-chromosome cercopithecine subgroup and the
42-chromosome subgroup.
Intergroup Hybridization— Nine hybridiza-
tions are reported between species in the sinica
group (Macaca radiata, 5 hybridizations; M. as-
samensis, 4) and species in the fascicularis (M.
fascicular is, 1 , inferred; M. mulatta, 4), silenus (M.
nemestrina, 1), and arctoides (M. arctoides, 3)
groups (table 14). Male and female reproductive
organs in the sinica group are strikingly different
from those in the fascicularis, silenus, and arc-
toides groups (Fooden, 1 980, p. 2), but these an-
atomical differences evidently do not prevent in-
tergroup copulation and fertilization, at least in
captivity.
Attempts to form mixed-species social groups
by confining together members of sinica-group
species (Macaca radiata, M. assamensis) with
members of fascicularis-group and silenus-group
species generally have been unsuccessful (Bern-
stein & Gordon, 1980, pp. 135, 137). However,
Stynes et al. (1975, p. 822, abstract only) report a
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
29
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FIELDIANA: ZOOLOGY
tendency toward increased social interaction and
sexual behavior (details unspecified) between M.
radiata and M. nemestrina after several individ-
uals of these species had been kept together for
more than 1 2 weeks.
Natural intergroup contacts between Macaca
radiata (sinica group) and M. mulatta (fascicularis
group) have been observed in India at four local-
ities along the border between the ranges of these
two species (Fooden et al., 1981, p. 465; Fooden,
1986, p. 14). In one village, a troop of M. radiata
remained within 1 0-50 m of a troop of M. mulatta
for about an hour without overt social interaction
between the troops. At three other localities, M.
radiata males apparently were closely integrated
into M. mulatta troops and interacted amicably
with M. mulatta females. No matings— interspe-
cific or intraspecific— were seen during the course
of these observations, which were made outside
of the peak breeding season (September-Novem-
ber) of both M. radiata and M. mulatta (Roonwal
& Mohnot, 1977, p. 110; Fooden, 1981, p. 27).
Recognizable hybrids of M. radiata and M. mu-
latta were not observed in this border area. Al-
though M. radiata and M. mulatta hybridize in
captivity (table 14) and apparently are compatible
in naturally occurring mixed-species troops, an
unknown behavioral or physiological barrier, pre-
viously also postulated by Bernstein and Gordon
(1979, p. 271; 1980, p. 146), evidently restricts
gene flow between these species in the interspecific
contact zone.
Intragroup Hybridization— Three reported
intragroup hybridizations are between Macaca
sinica and M. radiata, and two are between M.
radiata and M. assamensis (table 14). Not sur-
prisingly, these two hybridizing species pairs are
composed of species that are near each other in
body size (table 1).
Intragroup hybridization may occur more readi-
ly than intergroup hybridization. A captive M. as-
samensis female who had easy access to M. ne-
mestrina and M. arctoides males and more difficult
access to a M. radiata male preferentially associ-
ated with the M. radiata male, despite repeated
efforts by keepers to separate them (Acharjyo &
Misra, 1977, p. 521; 1982, p. 376); this pair ul-
timately produced two hybrid offspring. For
another account of compatibility of captive M.
assamensis and M. radiata, see Dathe (1983, p.
127).
Natural intragroup hybridization is now im-
possible between Macaca sinica and M. radiata,
the ranges of which are separated by the Palk Strait,
and between M. radiata and M. assamensis, sep-
arated by a 1,300-km gap. Natural hybridization
may occasionally occur between M. assamensis
and M. thibetana in northern Guangxi Province,
China, where the ranges of these two species are
in close proximity (fig. 1).
Phenotypes of Hybrids— Phenotypic data are
available for six intrageneric hybrids (table 14).
Based on these limited data, tentative inferences
may be drawn concerning relative dominance of
certain taxonomic character states in Macaca. ( 1 )
Crown hair growth pattern: The pattern in M. ra-
diata (large whorl with short anterior hairs, ex-
posed forehead hairs parted) apparently is domi-
nant to that in M. sinica (large whorl with long
anterior hairs), M. assamensis (whorl small or ab-
sent), M. fascicularis (whorl small and irregular or
absent), and M. mulatta (whorl absent). (2) Tail
length: The long tail in M. radiata apparently is
dominant to the shorter tail in M. assamensis, but
it is incompletely dominant (hybrids intermediate)
to the short tail in M. mulatta; tail length domi-
nance in intragroup hybridization may differ from
that in intergroup hybridization. (3) Facial skin
color: Pale lips and ears in M. radiata apparently
are dominant to blackish lips and ears in M. sinica.
(4) Dorsal pelage color: The saturate brown color
in M. assamensis apparently is dominant to the
drab brown color in M. radiata. Judging from
available evidence, character states in M. radiata
generally tend to be dominant over those in other
macaque species; this was previously indicated by
Hill (1937, p. 384), based on study of one M.
sinica x M. radiata hybrid.
Evolution and Dispersal
The following reconstruction of the evolution-
ary history of the sinica group is based mainly on
inferences from morphology, distribution, and
natural history of living species and subspecies.
Only one known fossil has been unequivocally re-
ferred to the sinica group (Delson, 1980, p. 19;
Ha, 1985, p. 82).
Macaques probably reached southern Asia about
Late Pliocene (Delson, 1980, p. 25), which implies
that evolution of the sinica group occurred mainly
during the Pleistocene. This was an epoch of great
changes in the topography, climate, sea level, and
plant distribution of southern and eastern Asia
(Liu & Ding, 1984, p. 14; Sharma, 1984, p. 58;
Vishnu-Mittre, 1984, p. 499), and undoubtedly
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
31
Fig. 15.
80° 90° 100° 110°
Hypothetical reconstruction of principal stages in evolution and dispersal of sm/ca-group macaques.
these changes strongly influenced the evolutionary
history of the sinica group. Unfortunately, knowl-
edge of the details of these environmental changes
generally is not sufficiently precise to permit spe-
cific environmental changes to be associated with
specific evolutionary events in the history of the
sinica group. Such association is attempted here
only for Late Pleistocene and Holocene, the last
two of six evolutionary stages discussed below.
1. Origin and Early Dispersal
of sinica Group: Macaco sinica/ M. radiata
Species and subspecies in the sinica group con-
stitute an orderly morphological and geographic
series that extends from small-bodied, long-tailed
M. sinica and M. radiata in Sri Lanka and pen-
insular India at one extreme, to large-bodied, short-
tailed M. thibetana in east-central China at the
other extreme (fig. 2). The regularity of this series
suggests that these species and subspecies origi-
nated sequentially as a result of successive epi-
sodes of dispersal, isolation, and differentiation.
The evolutionary polarity in this series presum-
ably is from longer-tailed species with many cau-
dal vertebrae to shorter-tailed species with few
caudal vertebrae, since a long tail generally is the
primitive condition in monkeys. This implies that
M. sinica and M. radiata probably are closest to
the ancestral stock of the sinica group and that the
center of origin of the group probably was in the
area of Sri Lanka and peninsular India (fig. 15; cf.
Hill & Bernstein, 1969, p. 13; Delson, 1980, p. 25;
Eudey, 1980, p. 64; Wada, 1985, p. 38).
The silenus group of macaques apparently also
originated in the area of Sri Lanka and peninsular
India (Fooden, 1975, p. 68). The morphology of
male and female genitalia is more derived in the
sinica group (see pp. 19, 24) than in the silenus
group (Fooden, 1 975, p. 28). The sinica group may
have originated as an offshoot of the silenus group
in the Sri Lanka-peninsular India area. If so, the
silenus group ancestor presumably was an un-
known, extinct species in which the tail was longer
than in living M. silenus. The origin of the sinica
group probably occurred fairly early in the Pleis-
tocene, judging from the number of subsequent
speciation events that are inferred to have oc-
curred in this group. The underlying cause of the
32
FIELDIANA: ZOOLOGY
original splitting, which presumably inaugurated
the distinctive genital specializations of the sinica
group, is unclear.
From the Sri Lanka-peninsular India area, the
sinica group, at the stage of ancestral Macaca sin-
ica or M. radiata, evidently spread northward and
ultimately reached the foothills of the Himalayas
(which were then lower than at present). Whether
this northward dispersal occurred simultaneously
with the parallel dispersal of the silenus group
(Fooden, 1975, p. 68) is unknown; at present,
species in these two groups are almost completely
segregated from one another, either ecologically or
geographically (Fooden, 1986, p. 1 4), and they may
have been similarly segregated in the past. As the
sinica- radiata stock moved northward, its body
size apparently increased (fig. 3), in accord with
Bergmann's rule, but its tail length apparently re-
mained approximately constant— about 550 mm
(fig. 4).
parently was obstructed by the high north-south
ranges of Hengduan Shan. Blocked from north-
ward and eastward spread by high mountains, this
stock evidently spread southward via mideleva-
tion forest accessible on the relatively low moun-
tain chains that extend into the Indochinese Pen-
insula. As the M. a. assamensis stock spread
southward, its head and body length apparently
decreased, in accord with Bergmann's rule, but its
tail length remained approximately constant (figs.
3—4). Ultimately, southward spread of the M. a.
assamensis stock in the Indochinese Peninsula ap-
parently was stopped by competition with M. ne-
mestrina leonina, which has similar habitat re-
quirements and is almost perfectly parapatric with
M. a. assamensis (Fooden, 1982, p. 24). Fossil
evidence indicates that M. a. assamensis reached
northern Vietnam (Vo Nhai District, ca. 21°45'N,
106°00'E) before 18,600 ybp (Ha, 1985, p. 82).
2. Origin of Macaca assamensis pelops
A major evolutionary discontinuity evidently
occurred when an offshoot of the sinica- radiata
stock colonized midelevation evergreen forest on
the slopes of the east-west trending Himalayas
(Fooden, 1982, p. 17). Tail length in the Hima-
layan population shortened, apparently abruptly,
from about 550 to 300 mm (fig. 4). This shortening
of the tail, which marked the origin of M. assa-
mensis pelops, may have been an adaptation to
the cooler climate of the new habitat, as predicated
by Allen's rule. The M. a. pelops stock apparently
spread from west to east through the belt of Hi-
malayan midelevation evergreen forest.
3. Origin of Macaca assamensis assamensis
4. Origin of Macaca thibetana
An offshoot of the M. a. assamensis stock ap-
parently dispersed around the southern end of
Hengduan Shan and became isolated in the region
of upper Chang Jiang (Yangtze River). This iso-
lation may have been caused by a glacial advance
at the divide between the drainage basins of upper
Lancang Jiang and Yuan Jiang (Mekong and Red
rivers) and the drainage basin of upper Chang Jiang.
Tail length in the isolated upper Chang Jiang pop-
ulation decreased, again abruptly, from about 200
mm to less than 100 mm (fig. 4), marking the
origin of M. thibetana. Head and body length in
the M. thibetana stock evidently has remained ap-
proximately the same as in the northern popula-
tion of M. a. assamensis from which it was derived
(fig. 3).
The next important change in the morphology
of the sinica group evidently occurred when an
offshoot of the M. a. pelops stock gained access to
the foothills of the north-south trending moun-
tains in Southeast Asia (Hengduan Shan) and be-
came isolated there; this isolation may have been
caused by a glacial advance in the region of the
Brahmaputra gap at the eastern end of the Hi-
malayan chain. Tail length in the isolated Heng-
duan Shan population decreased, again apparently
abruptly, from about 300 to 200 mm (fig. 4), mark-
ing the origin of M. a. assamensis. Eastward spread
of the newly evolved M. a. assamensis stock ap-
5. Late Pleistocene
During the period of the most recent glaciation,
climaxing about 1 8,000 years ago, air temperature
was reduced and sea level was lowered. Both of
these environmental changes presumably affected
species and subspecies of the sinica group: (1)
Northern species and subspecies were forced
southward or to lower altitudes along with their
forest habitats (see Liu & Ding, 1984, p. 34). Dur-
ing this period, the altitudinal range of Macaca a.
pelops presumably was lower on the slopes of the
Himalayas than at present; the northern limit of
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
33
— M. sinica
— M. radiata
— M. a. pelops
M. a. assamensis
M. t hi be tana
Fig. 16. Phylogenetic relationships inferred among
sinica-group macaques. Note that M. a. assamensis and
M. thibetana are shown as sharing a common ancestor
more recently than M. a. assamensis and M. a. pelops;
this is incongruous but unavoidable when a particular
subspecies of one species is identified as the probable
ancestor of another species.
the range of M. thibetana was farther south; the
boundary between M. thibetana and M. a. assa-
mensis was farther south; and the boundary be-
tween M. a. assamensis and M. nemestrina leonina
was farther south and/or at lower altitudes. (2) As
a consequence of glacially induced sea-level regres-
sion, the present range of M. sinica in Sri Lanka
was connected to the present range of M. radiata
in peninsular India (Jacob, 1949, p. 341; Sahni &
Mitra, 1980, p. 56). The step-cline color gradient
that now extends through both recognized sub-
species of M. sinica and both recognized subspe-
cies of M. radiata (Fooden, 1 98 1 , p. 9), transcend-
ing the specific boundary, suggests that the M. sinica
and M. radiata stocks may have been genetically
continuous— hence not specifically distinct— when
their ranges were geographically continuous dur-
ing the most recent glaciation. Hainan and Taiwan
also were connected to the mainland during the
same glaciation (Liu & Ding, 1984, p. 16), but
neither of these islands is now inhabited by sinica-
group macaques, although both are inhabited by
macaques belonging to the fascicularis group
(Fooden, 1980, p. 5). If M. a. assamensis or M.
thibetana colonized Hainan or Taiwan during the
late Pleistocene sea-level regression, they evi-
dently subsequently became locally extinct.
Lanka from peninsular India, thereby isolating the
sinica stock from the radiata stock and presum-
ably promoting their genetic divergence. The Ho-
locene may also be the epoch when M. mulatta
dispersed westward into northern peninsular India
and disrupted the presumed former contiguity of
the ranges of M. radiata and M. a. pelops (Fooden,
in press). An isolated population of M. radiata
within the range of M. mulatta in east-central pen-
insular India suggests that the advance of M. mu-
latta and disappearance of M. radiata in this area
have occurred relatively recently (Fooden et al.,
1981, p. 472; Saha, 1984, p. 164). The isolated
Sundarbans population of M. a. pelops (Fooden,
1982, p. 2) may be another indication of recent
contraction of the range of the sinica group in this
area.
Phylogenetic relationships among sinica-group
macaques that are implied by the proposed evo-
lutionary reconstruction are depicted in Figure 1 6.
Acknowledgments
For facilitating this research, I am deeply grate-
ful to officials and staff members of the institutions
listed in the Introduction. I am also grateful to the
Committee on Scholarly Communication with the
People's Republic of China for supporting my study
of Macaca assamensis and M. thibetana in China
in 1985. Valued collaborators in the research proj-
ect in China were Quan Guoqiang and Luo Yining,
Institute of Zoology, Chinese Academy of Sci-
ences, Beijing. I also thank James W. Koeppl and
Peter Lowther, Field Museum of Natural History,
for statistical advice and assistance, and Bruce D.
Patterson, Field Museum of Natural History, for
helpful comments on parts of the manuscript.
6. Holocene
During the Holocene, as a consequence of post-
glacial warming, vegetation zones have shifted
northward (with some oscillations), and the ranges
of Macaca a. pelops, M. a. assamensis, and M.
thibetana have correspondingly shifted northward
and upward to their present latitudes and altitudes.
Holocene sea-level elevation has separated Sri
Gazetteer
This list of sinica-group macaque localities sup-
plements previously published lists, as specified
below for each species or subspecies. For speci-
mens examined, a parenthetical notation indicates
the abbreviated name of the institution where
specimens are preserved (see Introduction), the\
number of specimens available, and the part that I
is preserved, if skin and skull are not both present
34
FIELDIANA: ZOOLOGY
Macaca sinica
(supplement to Fooden, 1979, p. 133; 1986, p. 2)
Sri Lanka
Ruhunu National Park; Southern Prov.; 06°21'N,
8 1°27'E; observed 1 968-1 975 by W. P. J. Dittus
(1977, p. 242).
Udawatakelle Sanctuary; Central Prov.; 07°18'N,
80°39'E; observed 1968-1975 by W. P. J. Dittus
(1977, pp. 239, 257).
Macaca radiata
(supplement to Fooden, 1981, p. 37; 1986, p. 2;
Fooden et al., 1981, p. 469)
India
Parambikulam Wildlife Sanctuary, ca. 600 m;
Kerala State; ca. 10°25'N, 76°43'E; observed
1972-1978 by V. S. Vijayan (1979, p. 890); ob-
served 1981-1983 by M. Balakrishnan and P.
S. Easa(1986, p. 196).
Thambraparni and Servalar rivers, Mundanthu-
rai Sanctuary, 180 m; Tamil Nadu State; ca.
08°40'N, 77°28'E; observed Feb. 1977-Apr.
1978 by R. Ali (1986, p. 98).
Udevara, NE, 960 m; Hassan District, Karnataka
State; 13°01'N, 75°50'E; observed Apr. 1972-
Aug. 1973 by H. Rahaman and M. D. Parthas-
arathy (1979, p. 406).
Macaca assamensis pelops
(supplement to Fooden, 1982, p. 35; 1986, p. 22)
China
Xizang
Zhangmu; Nyalam Co.; 28°02'N, 85°55'E; col-
lected by Scientific Mountaineering Team of
China, 1974 (nwpib, 1, skin only).
Macaca assamensis assamensis
(supplement to Fooden, 1982, p. 35; 1986, p. 22)
China
Guangxi
Chongzuo Co., ca. 22°24'N, 107°2 1 'E; reported by
Tan (1985, p. 73).
Darning Shan (mt.); probably Shanglin Co.; ca.
23°23'N, 108°30'E; reported by Shen Lantian
(in Tan, 1985, p. 73).
Daxin Co.; ca. 22°50'N, 107°12'E; reported by Wu
(1983, p. 16).
Jingxi Co.; ca. 23°08'N, 106°25'E; reported by Wu
(1983, p. 16). Comment: misspelled "Jiangxi"
by Fooden (1986, p. 22).
Ningming Co.; ca. 22°07'N, 107°02'E; reported by
Tan (1985, p. 73).
Guizhou
Jiangkou Co.; ca. 27°41'N, 108°49'E; apparently
erroneous report (Editorial Committee of
Guizhou Fauna, 1979, p. 1 10), probably based
on misidentified M. thibetana (see Fooden et
al., 1985, p. 15). Not mapped in Figure 1.
Xizang
Beibeng, 900 m; Medog Co.; 29°15'N, 95°30'E;
collected by Cai Guiquan and Feng Zuojian, 3
Aug. 1977 (nwpib, 1). Comment: locality pre-
viously recorded as "Medog" (Fooden, 1982, p.
41).
Yigong, 2250 m and 2750 m; Bomi Co.; 30°08'N,
95°02'E; collected by Feng Zuojian and Zheng
Changlin, 21 June and 9 Sep. 1973 (izcas, 2
[including 1 skull at nwpib]). Comment: locality
previously recorded as "Bomi" (Fooden, 1982,
p. 39).
Yunnan
Lengsuihe; Datang Dist., Tenchong Co.; 25°39'N,
98°38'E; collected by Fang Lixiang, Apr. 1960
(bjmnh, 2, skins only).
Lijiang Co.; 26°51'N, 100°13'E; apparently erro-
neous report (Tan, 1985, p. 73). Comment: ac-
cording to Wang Yingxiang, kjz, the only species
of macaque in Lijiang Co. is M. mulatta (pers.
comm., 1 1 Dec. 1985). Not mapped in Figure 1.
Longling Co. (Li & Lin, 1983, p. 113). See Xiao-
heshan (Fooden, 1986, p. 22).
Luchun Co. (Li & Lin, 1983, p. 113). See Da-
hongshan (Fooden, 1986, p. 22).
Menglian Co.; ca. 22°21'N, 99°36'E; reported by
Tan (1985, p. 73).
Pingbian Co. (Li & Lin, 1983, p. 113). See Dawei
Shan (Fooden, 1986, p. 22).
Xishuangbanna Prefecture (Li & Lin, 1 983, p. 1 1 3).
See Lancang Jiang, Menglun, Menghan, Xiang-
ming, Manpa, and Mengla Xian (Fooden, 1986,
P. 22).
India
Proposed Dhaleswari Wildlife Sanctuary; Assam
State; 24°10'-24°40'N, 92°20'-93°10'E; report-
ed by Choudhury (1983, p. 14).
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
35
Thailand
Huai Nua Pla, 2500 ft [760 m]; Tak Prov.; 1 6°54'N,
98°48'E; collected by J. H. Chambai, 9 May
1924 (zrcnus, 1). Comments: for locality notes
and coordinates, see Chasen and Kloss (1930,
p. 62) and Moore and Tate (1965, p. 321); spec-
imen previously misidentified as M. mulatta
(Fooden, 1982, p. 52).
Hue Nya Pla. See Huai Nua Pla.
Hue Yah Pla. See Huai Nua Pla.
Macaca thibetana
(supplement to Fooden, 1983, p. 14; Fooden et
al., 1985, p. 15)
China
Anhui
Banqiao, 700-1000 m; Ningguo Co.; ca. 30°38'N,
1 18°58'E; hunter's pelt observed in farmhouse,
1973-1985 (Wada et al., 1986, pp. 81, 83). Not
mapped in Figure 1 .
Chimen Co. See Qimen Co.
Gegong, 600-800 m; Dongzhi Co.; 30°05'N,
1 17°1 l'E; reported 1973-1985 by Xiong Cheng-
pei (Wada et al., 1986, p. 83).
Guimenguan, 500 m; Huang Shan, Shexian Co.;
ca. 30°03'N, 1 1 8°09'E; troop captured Nov. 1972
(Wadaetal., 1986, p. 89).
Guniujiang, 1000-1 500 m; Shitai-Qimen Cos.; ca.
30°05'N, 117°30'E; reported 1973-1985 by
Xiong Chengpei (Wada et al., 1986, p. 83).
Huanghuajian, 600-1200 m; Shitai Co.; ca.
30°08'N, 117°20'E; reported 1973-1985 by
Xiong Chengpei (Wada et al., 1986, p. 83).
Jilian, 600-800 m; Yixian Co.; ca. 30°00'N,
1 18°00'E; reported 1 973-1985 by Xiong Cheng-
pei (Wada et al., 1986, p. 83).
Jiuhua Shan, 1000-1200 m; Qingyang Co.; ca.
30°27'N, 1 17°48'E; 6 troops reported 1973-1985
by Xiong Chengpei (Wada et al., 1986, pp. 83,
90).
Lianhuafeng, 800-1600 m; Huang Shan, Shexian
Co.; ca. 30°07'N, 118°10'E; observed 1976 by
Xiong Chengpei (Wada et al., 1986, p. 89).
Pailou, 600 m; Guichi Co.; 30°21'N, 117°18'E;
reported 1973-1985 by Xiong Chengpei (Wada
etal., 1986, p. 83).
Qihong, 200-600 m; Qimen Co.; ca. 29°35'N,
117°40'E; one monkey captured 1964, appar-
ently now extinct at locality (Wada et al., 1986,
p. 83).
Qimen Co.; ca. 29°53'N, 117°43'E; reported by
Tan (1985, p. 75).
Quliting, 1000-1400 m; Huang Shan, Shexian Co.;
ca. 30°08'N, 1 18°1 l'E; observed 1976 by Xiong
Chengpei (Wada et al., 1986, p. 89).
Rucun, 500-1000 m; Xiuning Co.; ca. 29°55'N,
1 18°07'E; observed 1960-1965, apparently now
extinct at locality (Wada et al., 1986, p. 83).
Shangyangjian, 800-1 200 m; Jixi Co.; ca. 30°05'N,
1 1 8°20'E; hunter's pelt observed in farmhouse,
1973-1985 (Wada et al., 1986, pp. 81, 83). Not
mapped in Figure 1 .
Songguan, 890-1700 m; Huang Shan, Shexian Co.;
ca. 30°1 l'N, 1 18°10'E; observed 1976 and 1977
by Xiong Chengpei (Wada et al., 1986, p. 89).
Tanglingguan, 800-1350 m; Huang Shan, Shexian
Co.; ca. 30°07'N, 118°09'E; observed 1977 by
Xiong Chengpei (Wada et al., 1986, p. 89).
Tianbangshi, 700-1 100 m; Huang Shan, Shexian
Co.; ca. 30°07'N, 1 18°09'E; one troop captured
Nov. 1972; another troop observed 1975-1977
by Xiong Chengpei, 1985 by Wada et al. (1986,
p. 89).
Xiancun, 600-900 m; Taiping Co.; ca. 30°08'N,
1 18°05'E; reported 1973-1 985 by Xiong Cheng-
pei (Wada et al., 1986, p. 83).
Xiangrupeng, ca. 800 m; Huang Shan, Shexian
Co.; ca. 30°08'N, 118°06'E; 15 monkeys cap-
tured 1980 (Wada et al., 1986, p. 89).
Xinglong, 600-800 m; Jingde Co.; ca. 30°19'N,
1 1 8°3 1 'E; hunter's pelt observed in farmhouse,
1973-1985 (Wada et al., 1986, pp. 81, 83). Not
mapped in Figure 1 .
Yixian Co.; ca. 29°55'N, 117°55'E; reported by
Tan (1985, p. 75).
Yulingkeng, 800-1100 m; Huang Shan, Shexian
Co.; ca. 30°04'N, 1 18°08'E; observed 1973-1977
and 1980 by Xiong Chengpei; 27 monkeys cap-
tured 1974 and 1977; observed 1985 by Wada
etal. (1986, p. 89).
Yungusi, 570-1000 m; Huang Shan, Shexian Co.;
ca. 30°07'N, 1 18°13'E; observed 1973 and 1975
by Xiong Chengpei (Wada et al., 1986, p. 89).
Yunwaifeng, ca. 1000 m; Huang Shan, Shexian
Co.; ca. 30°08'N, 118°09'E; observed 1977 by
Xiong Chengpei (Wada et al., 1986, p. 89).
Fujian
Dadongken; Shangang Dist., Chong'an Co.;
27°50'N, 117°48'E; collected by Qin Yaoling,
1960(sciea, 1).
Guangze Co.; ca. 27°3 1 'N, 1 1 7°1 9'E; observed Sep.
1981 (Zheng, 1984, p. 145), cited as At. arc-
toides.
36
FIELDIANA: ZOOLOGY
Longyan Co.; ca. 25°06'N, 117°00'E; tentative
identification; observed Oct. 1982, cited as M.
arctoides by Zheng ( 1 984, p. 1 46), who applies
the same name to the stumptail macaque of
Chong'an Co. (= M. thibetana; amnh, fmnh,
MCZ, MNHN, SCIEA, USNM).
Meihua Shan (mts.); ca. 25°15'N, 116°45'E; ten-
tative identification; reported as M. arctoides by
Zheng ( 1 984, p. 1 46), who applies the same name
to the stumptail macaque of Chong'an Co. (=
M. thibetana; amnh, fmnh, mcz, mnhn, sciea,
usnm).
Pucheng Co.; ca. 27°54'N, 1 1 8°3 1 'E; observed Aug.
1980 (Zheng, 1984, p. 145), cited as M. arc-
toides.
Shanghang Co.; ca. 25°02'N, 116°23'E; tentative
identification; observed Sep. 1982, cited as M.
arctoides by Zheng ( 1 984, p. 1 46), who applies
the same name to the stumptail macaque of
Chong'an Co. (= M. thibetana; amnh, fmnh,
MCZ, MNHN, SCIEA, USNM).
Shaowu Co.; ca. 27°1 9'N, 1 1 7°29'E; observed June
1983 (Zheng, 1984, p. 145), cited as M. arc-
toides.
Gansu
Southern Gansu; ca. 32°50'N, 104°40'E; reported
by Tan (1985, pp. 75,80).
Guangdong
Bibei Qu, 100-200 m; Ruyuan Co.; ca. 25°01'N,
1 13°17'E; collected by unknown Yao hunter, 9
Nov. 1985, not preserved (Ling Wenfeng, rcfb,
pers. comm., 10 Nov. 1985).
Da'ao, 500-600 m; Luoyang Dist., Ruyuan Co.;
24°43'N, 113°05'E; traces observed Feb. 1983
by Ling Wenfeng, rcfb (pers. comm., 10 Nov.
1985).
Dapingding, ca. 1000 m; Longnan Dist., Ruyuan
Co.; 24°48'N, 113°06'E; observed 2 Oct. 1985
by Ling Wengfeng, rcfb (pers. comm., 10 Nov.
1985).
Goujiken, 700 m; Ruyuan Co.; 24°56'N, 1 1 3°04'E;
observed 8 Nov. 1985 by Huang Mingyan (Ling
Wengfeng, rcfb, pers. comm., 10 Nov. 1985).
Gouweizhang, < 1684 m; Dongpin Dist., Ruyuan
Co.; 24°57'N, 1 13°14'E; observed 15 Oct. 1985
by vice-director of district (Ling Wengfeng, rcfb,
pers. comm., 10 Nov. 1985).
Gumudong, 600-700 m; Gumushui Dist., Ruyuan
Co.; 24°36'N, 113°03'E; crop raid June 1985
reported by local farmers (Ling Wengfeng, rcfb,
pers. comm., 10 Nov. 1985).
Julongpin, ca. 1400 m; Lianxian Co.; 24°52'N,
1 12°41'E; living captive observed 25 Oct. 1985
by Cheng Xinzhou (pers. comm., 1 3 Nov. 1 985).
Jushonglou, 1000 m; Fucheng Dist., Ruyuan Co.;
24°49'N, 1 13°17'E; > 300 monkeys shot by lo-
cal hunter in 1 969-1 97 1 ; observed 3 Nov. 1 985
by officials of county construction bureau (Ling
Wengfeng, rcfb, pers. comm., 10 Nov. 1985).
Laopengeyiduei, 0.7 km NNE, 1100 m; Ruyuan
Co.; 24°56'N, 1 1 3°0 1 'E; collected by Mr. Zhang,
Forest Ranger, Qinxidong Nature Reserve, 1 1
June 1 985 (qnr headquarters, living captive ob-
served 9 Nov. 1985).
Leyang, ca. 800 m; Ruyuan Co.; 24°40'N, 1 13°03'E;
collected by Liu Zhenhe and Xu Longhuei, June
1970 and 15 July 1981 (sciea, 3, including 1
skull only).
Longnan Dist.; Ruyuan Co.; ca. 24°50'N, 1 13°05'E;
collected by Quan Guoqiang, 10 Nov. 1985 (iz-
cas, 2, skulls only).
Pingxi, ca. 800 m; Ruyuan Co.; 24°45'N, 1 1 3°00'E;
collected by Xu Longhuei, 15 July 1981 (sciea,
1).
Qinxidong Nature Reserve, Tract No. 25, ca. 700
m; Ruyuan Co.; 24°58'N, 1 1 3°02'E; collected by
Ling Wenfeng, rcfb, Oct. 1983, two specimens,
not preserved (pers. comm., 10 Nov. 1985).
Qinxidong Nature Reserve, Tract No. 37, ca. 1 100
m; Ruyuan Co.; 24°57'N, 1 13°03'E; calls of two
monkeys heard 6 Nov. 1985 by Ling Wenfeng,
rcfb (pers. comm., 10 Nov. 1985).
Qinxidong Nature Reserve, Tract No. 44, ca. 1 000
m; Ruyuan Co.; 24°56'N, 113°03'E; observed
Oct. 1983 by Ling Wenfeng, rcfb (pers. comm.,
10 Nov. 1985).
Shijiaoken, 700 m; Ruyuan Co.; 24°57'N, 1 1 3°05'E;
observed Oct. 1 983 by Ling Wenfeng, rcfb (pers.
comm., 10 Nov. 1985).
Tianjinshan, 800-1000 m; Luoyang Dist., Ruyuan
Co.; 24°42'N, 1 12°53'E; observed July 1983 by
local officials (Ling Wenfeng, rcfb, pers. comm.,
10 Nov. 1985).
Yao Shan (mts.), 800-1200 m; Lechang Co.; ca.
25°15'N, 113°15'E, collected by R. Mell, Sep.
1908-Feb. 1911 (zmb, 1). Comment: type lo-
cality of M. arctoides esau Matschie, 1912; pre-
viously reported as "Yao-tze Berge" (Mell in
Matschie, 1912, p. 309; Mell, 1922, pp. 4, 10;
Fooden, 1983, pp. 2, 17).
Guangxi
Chuanzhou Co. See Quanzhou Co.
Lingui Co.; ca. 25°12'N, 110°H'E; reported by
Shen Lantian (in Tan, 1985, p. 75).
FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES
37
Lipu Co.; ca. 24°30'N, 1 10°24'E; reported by Shen
Lantian (in Tan, 1985, p. 75).
Longsheng Co.; ca. 25°43'N, 110°01'E; reported
by Shen Lantian (in Tan, 1985, p. 75).
Luoyiang; HuanjiangCo.; 24°58'N, 108°12'E; col-
lected by local people in 1981, not preserved
(reported 29 Nov. 1985 by Wu Mingchuan, gi-
fid, to Quan Guoqiang, izcas; pers. comm., 12
Dec. 1985).
Quanzhou Co.; ca. 25°56'N, 1 1 1°02'E; reported by
Shen Lantian (in Tan, 1985, p. 75).
Xunle, HuanjiangCo.; 25°25'N, 108°15'E; present
in 1981 (reported 29 Nov. 1985 by Wu Ming-
chuan, gifid, to Quan Guoqiang, izcas; pers.
comm., 12 Dec. 1985).
Yangshuo Co.; ca. 24°46'N, 1 10°29'E; reported by
Shen Lantian (in Tan, 1985, p. 75).
Yangso Co. See Yangshuo Co.
Yongfu Co.; ca. 24°57'N, 109°58'E; reported by
Shen Lantian (in Tan, 1985, p. 75).
Youngfu Co. See Yongfu Co.
Yueli; Nandan Co.; 25°25'N, 107°15'E; specimen
collected Nov. 1981 by local people, not pre-
served (reported 29 Nov. 1985 by Wu Ming-
chuan, gifid, to Quan Guoqiang, izcas; pers.
comm., 12 Dec. 1985).
Ziyaan Co. See Ziyuan Co.
Ziyuan Co.; ca. 26°01'N, 110°39'E; reported by
Shen Lantian (in Tan, 1985, p. 75).
Guizhou
Chingzhen Co. See Qingzhen Co.
Guiding Co.; ca. 26°34'N, 107°13'E; reported by
Editorial Committee of Guizhou Fauna (1979,
p. 110).
Jiangkou Co. (Editorial Committee of Guizhou
Fauna, 1979, p. 1 10). See Fooden et al. (1985,
p. 15).
Qingzhen Co.; ca. 26°33'N, 106°28'E; reported by
Editorial Committee of Guizhou Fauna (1979,
p. 110).
Sandu Co.; ca. 25°58'N, 107°5 l'E; reported by Ed-
itorial Committee of Guizhou Fauna (1979, p.
110).
Suiyang Co.; ca. 27°56'N, 107°10'E; reported by
Editorial Committee of Guizhou Fauna (1979,
p. 110).
Xingyi Co.; ca. 25°05'N, 104°53'E; reported by
Editorial Committee of Guizhou Fauna (1979,
p. 110).
Zheng'an Co.; ca. 28°33'N, 107°26'E; reported by
Editorial Committee of Guizhou Fauna (1979,
p. 110).
Zhijin Co.; ca. 26°39'N, 105°46'E; reported by Ed-
itorial Committee of Guizhou Fauna (1979, p.
110).
Hunan
Chengbu Co.; ca. 26°20'N, 1 10°19'E; reported by
local people, Nov. 1 980 (Liu Zhenhe, sciea, pers.
comm., 25 Nov. 1985).
Guidong Co., E; ca. 26°00'N, 113°53'E; reported
by local people, Oct. 1979 (Liu Zhenhe, sciea,
pers. comm., 25 Nov. 1985). Comment: cited
as M. arctoides by Tan (1985, p. 74; pers. comm.,
16 Dec. 1985).
Huangshuang Nature Reserve; Suining Co.;
26°25'N, 110°03'E; reported by local people,
Nov. 1980 (Liu Zhenhe, sciea, pers. comm., 25
Nov. 1985). Comment: cited as M. arctoides by
Tan (1985, p. 74; pers. comm., 16 Dec. 1985).
Lanshan Co.; 25°21'N, 112°10'E; living captive
obtained 1982 (observed in Lianxian, Guang-
dong Pro v., 13 Nov. 1985).
Shunhuangshan Plantation, ca. 1000 m; Xinning
Co.; ca. 26°30'N, 110°55'E; reported by local
people, Nov. 1980 (Liu Zhenhe, sciea, pers.
comm., 25 Nov. 1985). Comment: cited as M.
arctoides by Tan (1985, p. 74; pers. comm., 16
Dec. 1985).
Xinning Co.; ca. 26°31'N, 110°48'E; collected 24
Dec. 1984 by local residents (Zhou, 1986, p.
109).
Zhezhiping, 1 200 m; Mangshan Dist., Yizhang Co.;
ca. 24°56'N, 1 12°53'E; collected by Liu Zhenhe,
9 Nov. 1980 (sciea, 1). Comment: cited as M.
arctoides by Tan (1985, p. 74; pers. comm., 16
Dec. 1985).
Zhiyunshan Nature Reserve, ca. 1000 m; Xinning
Co.; ca. 26°35'N, 1 1 1°05'E; reported by local
people, Nov. 1980 (Liu Zhenhe, sciea, pers.
comm., 25 Nov. 1985). Comment: cited as M.
arctoides by Tan (1985, p. 74; pers. comm., 16
Dec. 1985).
Jiangxi
Guixi Co.; Wuyi Shan (mts.); ca. 28°18'N,
1 17° 12 'E; living captives collected, Winter 1983,
not preserved (Sheng Helin, ecnu, pers. comm.,
19 Oct. 1985).
Hexilong, 700-1000 m; Jinggangshan Co.; ca.
26°32'N, 114°09'E; collected by Zheng Xian-
huai, 25 Dec. 1982 (jnrb, 1, mounted skin with
skull inside).
Jingzhushan, 1000 m; Jinggangshan Co.; 26°31'N,
1 14°06'E; collected by Long Dizong, 1 April 1 980
(jubd, 1 , mounted skin with skull inside).
38
FIELDIANA: ZOOLOGY
NE Jiangxi, "near the Anhui border"; probably
Jingdezhen Co.; ca. 29°00'N, 1 18°00'E; reported
by Tan (1985, pp. 75, 80).
Pingxiang (town), vicinity; Pingxiang Co.; ca.
27°38'N, 113°50'E; living captive collected 1983,
observed in Pingxiang Zoo by Sheng Helin, ecnu
(pers. comm., 19 Oct. 1985) and Huang Zhang-
sen, nz (pers. comm., 28 Oct. 1985).
Shangyou Co.; ca. 25°48'N, 1 14°30'E; living cap-
tive obtained May-June 1978 by Ma Jielun,
Ganzhou Zoo (captive observed at gz, 6 Nov.
1985). Comment: probably collected at Wuz-
hifeng, Shangyou Co.
Shanbaishan; Xunwu Co.; ca. 25°00'N, 1 15°45'E;
collected in 1976 by local people, not preserved
(Liu Zhenhe, sciea, pers. comm., 25 Nov. 1 985).
Vanshan Co.; ca. 28°18'N, 117°42'E; two living
captives collected by Huang Zhangsen, 1980
(captives observed at nz, 28 Oct. 1985); living
captives collected by local people, Winter 1983
(Sheng Helin, ecnu, pers. comm., 1 9 Oct. 1 985).
Comment: misspelled "Qianshan" by Fooden
etal. (1985, p. 15).
rushan Co., probably; ca. 28°41'N, 1 18°13'E; liv-
ing captive obtained in late 1 970s by Wu Fuhai,
Hangzhou Zoo, Zhejiang Prov. (captive ob-
served at hz, 25 Oct. 1985). Comment: obtained
from local people in Jiangshan, Zhejiang; re-
portedly collected across provincial boundary in
nearby Jiangxi Prov.
Sichuan
Bao Guo Si (temple), near; Emei Shan (mt.), Emei
Co.; ca. 29°32'N, 103°21'E; collected by Quan
Guoqiang, Aug. 1959 (izcas, 1, skull only).
~hiu-lao-tung; Emei Shan (mt.), Emei Co.; ca.
29°32'N, 103°21'E; observed Aug. 1982 by J.
D. Lazell, Jr. (1983, p. 61).
Xiang Feng, ca. 1900 m; Emei Shan (mt.), Emei
Co.; ca. 29°32'N, 103°21'E; observed Aug. 1982
by J. D. Lazell, Jr. (1983, p. 62).
'Western Sichuan", 43 counties; 27°-33°N, 98°-
103°E; questionable report (Tan, 1985, pp. 75,
80). Comment: hitherto, only M. mulatto has
been reported or collected in this area (Wilson,
1913, p. 192; Weigold, 1935, p. 233). Not
mapped in Figure 1.
Xizang
"Eastern Tibet"; ca. 28°40'N, 97°00'E; improbable
report (Tan, 1985, pp. 75, 80). Comment: ap-
parently in range of M. a. assamensis (see Food-
en, 1982, p. 27). Not mapped in Figure 1.
Xizang Prov.; improbable locality datum (smnh,
1, skin only). Comment: specimen received 8
June 1962 from Shanghai Zoological Garden,
which now has no record of it (Zhang Cizu, szg,
pers. comm., 18 Oct. 1985). Not mapped in
Figure 1.
Yunnan
Yongshan Co.; ca. 28°10'N, 103°40'E; collected
Aug. 1 984 by local hunter, not preserved (Wang
Yingxiang, kiz, pers. comm., 11 Dec. 1985).
Zhejiang
Beiyandangshan; YueqingCo.; 28°23'N, 121°04'E;
collected by Chai Weixi, 1 960 (zmnh, 2, mount-
ed skins with skulls inside).
Daoshiwu; Lin'an Co.; ca. 30°13'N, 1 19°43'E; liv-
ing captives collected Feb. 1985 by local people
(Wu Fuhai, hz, pers. comm., 25 Oct. 1985).
Jiulong Shan. See Zhuanxian.
Wangcunkou; Suichang Co.; 28°24'N, 118°59'E;
collected by Mao Jiangsen, June 1 979 (immzam,
1).
Zhoucun, ca. 1000 m; Jiangshan Co.; 28°22'N,
118°37'E; collected by Kang Ximin, 23 Mar.
1985 (zmnh, 1).
Zhuanxian, near Jiulong Shan (mt.); Suichang Co.;
ca. 28°20'N, 1 19°00'E; collected by villagers, 23
May 1957 (Zhou, 1984, p. 58).
Zhidaikou, ca. 1000 m; Suichang Co.; 28°16'N,
1 18°46'E; observed Aug. 1985 by Kang Ximin,
zmnh (pers. comm., 24 Oct. 1985).
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44
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