UNIVERSITY OF ILLINOIS LIBRARY AT URBANACHAMPAIGN BIOLOGY APR 91992 FIELDIANA Zoology NEW SERIES, NO. 45 Jack Fooden Taxonomy and Evolution of the Sinica Group of Macaques: 6. Interspecific Comparisons and Synthesis % ^ * f.* June 30, 1988 Publication 1389 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Information for Contributors to Fieldiana General: » is primarily a journal for Field Museum staff members and research associates, although manu m nonaffiliated authors may be considered as space permits. The Journal carries a page charge of $65 per printed page or fraction thereof. Contributions from staff, research associates, and invited authors will be idered for publication regardless of ability to pay page charges, but the full charge is mandatory for nonaffiliated authors of unsolicited manuscripts. Payment of at least 50% of page charges qualifies a paper for expedited processing, which reduces the publication time. Manuscripts should be submitted to Dr. James S. 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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 — i — i u-i vo m © vo (N oo ^ • • — . • 00 ~ m +1 00 +| -. +| vo r~ tJ- in tj- m I r-. os ^0Tt —Jo — < °^ Tf rn vd m — ■> — > — r~ +1 *p o r- in Tt (N •oo m Tt oo m ,-H _ ; • os • m • m • vo +1 - +1 op +| r-' +| r-' +| r^" -* — r~ os oo o o * Tt os o\t^ r^vd vd\o vd Tt r-'vo —c — 00 in m ^H Tt CO (N 00 fS O +1 "j> m t~~ 23 O 00 23 +1 "j> Tt m Tt m Tt m m m Tt TT a 00 in CM o ■* — OS O^Ci O <*"> o^ • m • (N • VO • Tt - — +1 -* +1 ©' +1 O +1 © t^ Tt VO O ION (N Os (NO OON in in M(S -^ © — * — _' 0' do do do 23 E - (NO — hO — vo m O +| m +| (N +| 00 I J I Tt in (N m Tt m (N © — m *oov vo r- n oo — os r- m in Tt m Tt m (N (N — so +| so O 00 o w oo m © m ""> TO m O (N OS +1 «j> +1 "A +| oooo r-o — © vo (N o_ so — (N m oo in — m n oo Tt Tf in Tt vi in vo •/"> vo m in — « m — " oo +| d oo © r^ oo --; in +1 ^ +\^ os in m -* Tt 00 ON mn ciri f» r- in — • oo os r- so " o +1 -r -:- *i +1 cp Tf V© od r-' ooV Tt +1 oo +| oo !"«• m f» — _ vo m vd Tt r-' r* 323 +1 .n +1 ^ +1 rs +1 ^ +1 ^ •n-H t^in tj-os Tfin os m mm mm m (N m (N mm — -* in rs _^ m m _ Tj- _ vo o • Tf ■ m 00 (N O 00 00 ^ +1 t-j- +| vo +| Tt O— ovofNoo mo o . . 8 82 B T3 ii u O II S.Oi, >< Uh Z w .. w 00 5c« iu 4> vo — »2 os a aos , « 2 m 0 Tf .. SO +1 « O o 8 vi 1 ^ « — ■f -_) c c c 1> «J 1) „ i i j «« o 5 u 00 £ m ,S II ,3 0 c — w ea HELDIANA: ZOOLOGY (ujuj) mBuei Apoq pus pbsh 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 ^ P > .01. ** = .01 > P > .001. FOODEN: COMPARISONS AND SYNTHESIS IN SINICA MACAQUES 11 12 FIELDIANA: ZOOLOGY ^^~% *\ /t^sA r-»-^ 0 o / 1^ \ - o (•J ■ / \\^ \ 0 Q\ c ■ / * \ \. \ ° \ 0 / ■ / ■ \ \\ o 0 \ • ■ / *"r \ - /* v (► *■ \^ \ , — / A 7 \ o w ■ \ m \ X ■*_ ■).. f \ 1 \ ' {> > k* > \ \ \ >" \ \ t* > K» \ ► V w \ ► \ > > >\ c 0) \ * \ "* E 0 \ \ ** m _ 0 0 o CN4 \ ► \ 0 6 \ ► \ . 5 \ 7 \£ > \ / ^-- > o ID 1 i , i v i i i , i ~>" i . i i r 0) D 5 • a O J_ o _ o o\ .; s •v w ** c z — ■ o • w 5 3 JO Sb (wui) giBuei jseieejO '||n)|S 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 — i Tf 00 r-' no' no fN fN fN in © fN co co co ON NO fN Tf f*> Tf VOOON -h in co Tf -h le c/5 r-1 no r~" On rt in fN fN fN tsmoo — © © fN NO O fN noo « o\ on e M ^, CO I I o NMNt NO fN C~- fN On On -^ O r»" Tf co »-> co On d Tf co co co r*S -h r- co >n Tf p oo no no 00 O Tf fN fN fN fN fN oo v© r~ 0\ — © © r» oo oo oo m r-^ — < oo 1 "*. °. on fN no On "o CO fN n o — < fN gTfTf in co Tf in in in Tf NO O rs p -h on r^ oo > ^ rs ^ On On On •O ~^ tv . fN NO Tf o oo 0\ co in o o on r- CS [ON p Tf CO Tf CO no P-_ 00 Tf NO fN Tf CO _ Tf m o CO fN —I CM fN fN fN fN "5 CM 41 od in d no O CN -<" on co in £2zir fN in p on E CM 2 ss m r~ Tf oo ■<* oo no r-» NO —J iriiflNr- r-' r-' no no £ NO Tf mi fN © ©do od ro co © O Tf -h' if — i p Tt ; in § CO Tf CO co Tf fN CO 0 -^ CO Tf* CO >> ON 1 On o e o "S 1 "J M 00 J? 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S c 3 « 2 u ' y. -$, 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 2 (N P — Oh e o a * >••= tf! •g -o n S ? B R 1,3 5° < W WW Os A AH 2^- ■* so M 5 "* 00 oo oo 55 e _5 >> >> >> S in MN .. S All All All S a o C ill*! o o o a JUhffl «-S a O O «j ** *P ^ o. «5 W h^ rt — i-, o» c^« *o t>~ e>- tr- *o so r- os os o «n r- 5 SO I SO *^ oo r- os os — } -Si -& ^2 ^2 •!2 § -3 * $ S 2 2 co -« ? a P ts tj ^3 ♦o a a a « , •i •£; •£; <3 v. h. v» v. l. q q <3 a Co Co Co V. S^^^ ^ 3*3*1 -h' fN co - g 2 S •»■§ ob o*^ ± i3 « w- ^ 3 S C (3£ o "S « S <£ c u 2 s o g o-> ,9 ■acd ^ 1-1 ^j a-? S 5 5U 30 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). Literature Cited Acharjyo, L. N., and R. Misra. 1977. Notes on a young hybrid macaque. Journal of the Bombay Nat- ural History Society, 73: 521-522. . 1982. Further notes on the birth of hybrid macaque Macaca radiata x Macaca assamensis at Nandankanan Biological Park, Orissa. The Indian Forester, 108: 376. (Reprinted in The Indian Forester, 108: 464.) Ahaley, S. K., M. 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Species and Subspecies Accounts of the Indian Bonnet Macaque, Macaco, radiata. By Jack Fooden. Fieldiana: Zoology, n.s. 3, 1981. 52 pages, 9 illus., 15 tables. Publication 1325, $5.50 raxonomy and Evolution of the Sinica Group of Macaques: 3. Species and Subspecies Accounts of By Jack Fooden. Fieldiana: Zoology, n.s. 9, 1982. 52 pages, 10 illus., 1 1 tables. Publication 1329, $5.25 raxonomy and Evolution of the Sinica Group of Macaques: 4. Species Account of Macaca thibetana. By Jack Fooden. Fieldiana: Zoology, n.s. 17, 1983. 20 pages, 5 illus., 3 tables. Publication 1345, $3.50 raxonomy and Evolution of the Sinica Group of Macaques: 5. Overview of Natural History. By Jack