UNIVtRSlTY ^y « umSS -chTmpwgn *"• BIOLOGY ■'..■■.'. .-• ■ .-..■. ■ .-,.'" ■ ..- ■■■■'.■■■■ FIKLDIANA Zoology NEW SERIES, NO. 29 9A-2S -.} I*70i BIOLOGY LIBRARY 101 . HALL Taxonomy and Evolution of the Sinica Group of Macaques: . Overview of Natural History Fooden A Contribution in Celebration of the Distinguished Scholarship of Robert F. Inger on the Occasion of His Sixty-Fifth Birthday July 31, 1986 Publication 1367 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY Information for Contributors to Fieldiana General: Fieldiana is primarily a journal for Field Museum staff members and research associates, although manuscripts from nonaffiliated authors may be considered as space permits. The Journal carries a page charge of $65 per printed page or fraction thereof. 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All desired corrections of type must be made on the single set of page proofs. Changes in page proofs (a> opposed to corrections) are very expensive. Author-generated changes in page proofs can only be mi the authoi idvance to pay for them. FIELDIANA Zoology NEW SERIES, NO. 29 Taxonomy and Evolution of the Sinica Group of Macaques: 5. Overview of Natural History Jack Fooden Research Associate Division of Mammals Department of Zoology Field Museum of Natural History Chicago, Illinois 60605-2496 A Contribution in Celebration of the Distinguished Scholarship of Robert F. Inger on the Occasion of His Sixty-Fifth Birthday Accepted for publication June 25, 1985 July 31, 1986 Publication 1367 PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY © 1986 Field Museum of Natural History Library of Congress Catalog Card Number: 81-65061 ISSN 0015-0754 PRINTED IN THE UNITED STATES OF AMERICA Table of Contents List of Illustrations Abstract 1 Introduction 1 Natural History of the Sinica Group ... 1 Habitats 1 Arboreality/Terrestriality 3 Food 4 Predators and Parasites 4 Seasonal Migration 5 Troop Size and Composition 5 Home Range 7 Male Emigration 8 Seasonal Breeding 8 Mating Behavior 10 Life History 11 Geographic and Ecological Relationships with Other Primates 14 Summary 16 Acknowledgments 17 Literature Cited 17 Appendix: Macaca assamensis Locality Rec- ord Supplement 22 1 . Locality records and inferred limits of natural distribution of sinica group species 2 2. Macaca radiata young adult male follow- ing M. mulatta adult female, with another M. mulatta individual sitting nearby 15 List of Tables 1 . Troop size in forest populations of sinica group species 5 2. Macaca radiata: troop size in forest troops compared with that in nonforest troops 6 3. Macaca radiata: ratio of sexually mature males (subadults and adults) to sexually mature females (adults) in 4 1 8 troops . . 6 4. Home range area in Macaca sinica and Macaca radiata 7 in Taxonomy and Evolution of the Sinica Group of Macaques: 5. Overview of Natural History Abstract The sinica group of macaques includes four species: Macaca sinica, M. radiata, M. assamen- sis, and M. thibetana. The geographic ranges of these species extend from tropical Sri Lanka (M. sinica) to subtropical east-central China (M. thi- betana). In the present review of the natural his- tory of the sinica group, these species are com- pared with respect to habitat, arboreality/ terrestriality, food, predators and parasites, sea- sonal migration, troop size and composition, home range, male emigration, seasonal breeding, mating behavior, life history, and geographic and ecolog- ical relationships with other primates. of other aspects of the systematics and evolution- ary biology of these species is in preparation. In citations of unpublished natural history notes recorded on specimen tags or in collectors' field notebooks in Chinese museums, the following in- stitutional abbreviations are used: izas— Institute of Zoology, Academia Sinica, Beijing; kiz— Kunming Institute of Zoology, Academia Sinica, Kunming. I thank officials of these institutions for generously providing access to specimens and field notebooks in their custody. I am happy to dedicate this paper to Robert F. Inger, distinguished zoologist and valued col- league, on the occasion of his 65th birthday an- niversary. Introduction The sinica group is one of four species groups of macaques defined by the morphology of male and female reproductive organs (Fooden, 1 97 1 , p. 72). This group includes four species: Macaca sin- ica, in Sri Lanka; M. radiata, in southern penin- sular India; M. assamensis, in the central and east- ern Himalayan foothills and adjacent mountains of Southeast Asia; and M. thibetana, in east-cen- tral China (fig. 1). Taxonomic accounts of these four species have been published recently (Foo- den, 1979, 1981, 1982a, 1983). This paper is a comparative review of the nat- ural history of sinica group species. Information available for these species varies greatly in com- pleteness. In addition to works cited here, those cited in previously published species accounts should also be consulted. A comprehensive review Natural History of the Sinica Group Habitats Habitats of species in the sinica group differ climatically according to the varying latitudes of the geographic ranges of these species (fig. 1). The latitudinal effect on climate is further accentuated by an altitudinal effect, since the more northern species tend to inhabit higher altitudes. In the range of M. sinica (latitude 06°-09°N, altitude 0-2100 m), climate varies from tropical wet to tropical arid; in the range of M. radiata (08°-2 1 °N, 0-2 1 00 m), climate also varies from tropical wet to trop- ical arid, but arid habitats constitute a greater pro- portion of the total; in the range of M. assamensis (15°-30°N, 0-3100 m; see kiz 780118, 780144, Bilushui Shan, Yunnan, China), climate is sub- tropical wet; and in the range of M. thibetana (25°- FOODEN: NATURAL HISTORY OF SINICA MACAQUES .. * c •rt o i~ Q >,*£> ^ 00 •5&V a- Baf- jf £ .2 — S 8* * I •s x "a « 3 a 2 w (« 00 B On ^T s-a- be p . i * <- -a o <3 tn _E — ^< g£ So a rf '8 9 •C ■* .« _f (I! £ fi § — ' a s 300 2 35 arcr00 •a c oo § "so - U Ov 1.9 S la|K-g _&" '< *■ £ <« ! *»> r* "> i — — < a. ?3? o •* — 1 * S« cd lags i _r o « J* o c ! Ov -c C/3 g u, o "1 « ££* « § .2 caoo^g - S-S" a S . s-' 3 ** g 2 2 PriS U, = C oo T3 •5 ^ <* -a FIELDIANA: ZOOLOGY 33°N, 1000-2500 m), climate varies from sub- tropical wet to moderate continental moist, with snow in winter. Species in the sinica group differ in the kinds and diversity of forest habitats that they occupy within their geographic ranges. M. sinica success- fully inhabits all three major forest types— ever- green, semievergreen, and arid— within its range (Dittus, 1977b, p. 256; Fooden, 1979, p. 121). Arid zone forest habitats of M. sinica are restricted to the immediate vicinity of streams and ponds. Dittus (1982, p. 276; 1985, p. 4) has documented the important effects of recurrent tropical cyclones on the habitats of M. sinica. Although M. radiata also has been observed in all forest types within its geographic range, it is rarely encountered in evergreen forests and instead is much more common in semievergreen, decid- uous, and disturbed habitats (Prasad et al., 1979, p. 738; Fooden, 1981, p. 15; Singh et al., 1984, p. 8). The preference of M. radiata for disturbed hab- itats has been reconfirmed by a recent field survey (Pirta et al., 1981, pp. 431, 433): uous and bamboo forests and in cultivated fields, but only in the immediate vicinity of mountain broadleaf evergreen forest (Dao, 1978, p. 382; Fooden, 1982a, p. 17). One anomalous, possibly relict, record of this species is in the Sundarbans tidal mangrove swamps at the mouths of the Ganges River (Anderson, 1872, p. 529; Fooden, 1982a, p. 2; Zoological Survey of India, National Zoological Collection, Calcutta, Specimen No. 11999). The preferred habitat of M. thibetana also ev- idently is midelevation mountain broodleaf ev- ergreen rainforest (Fooden, 1983, p. 11; Xiong, 1 984, p. 3; Fooden et al., 1 985, p. 24). This species, like M. assamensis, occasionally moves opportun- istically from evergreen forest into adjacent bam- boo groves, natural clearings, and cultivated fields (Mell in Matschie, 1912, p. 309; Xiong, 1984, p. 5; Quan Guoqiang, 1960, unpublished field notes, izas 17966-69, Sujiapo, Guizhou). Arboreality/Terrestriality Most of the bonnet groups were found outside the forest inhabiting roadsides near agricul- tural areas covered with a thick vegetation of banyan (Ficus bengalensis), pipal (Ficus reli- giosa) and imli (Tamarindus indica) trees. . . . It was also observed that bonnet monkeys are very rare in the interior forest, and some groups which do live in jungle are found near tribal huts or the forest bungalows. Although M. radiata apparently prefers nonev- ergreen and disturbed habitats to evergreen forest, in its generally nonevergreen habitats this species intensively exploits scattered or clumped ever- green trees, such as fig trees, for food and roosting places (Singh et al., 1980, p. 102). At Elephanta Island, near Bombay, Kuruvilla (1980, p. 979) re- ports: During the dry seasons most of the trees shed their leaves. At this time the macaques [M. radiata] select roosts where evergreen trees are numerous. The preferred habitat of M. assamensis evi- dently is subtropical midelevation mountain broadleaf evergreen rainforest (Fooden, 1982a, p. 1 7; Li Zhixiang, 1973, unpublished field notes, kiz: Coll. No. 73231, Satun, Yunnan, China). This species occasionally has been recorded in decid- Macaca sinica, M. radiata, and M. assamensis evidently are much more arboreal than M. thi- betana. In well-studied troops of M. sinica, day- time activity at one locality was 76% arboreal (Dit- tus, 1977b, pp. 245, 257). In M. radiata, daytime activity at various localities is estimated to range from 70% to 90% arboreal (Sugiyama, 1972, p. 251; Rahaman & Parthasarathy, 1979, p. 415; cf. Clarke, 1978, p. 518). Percentage estimates of ar- boreal activity are not available for M. assamensis, but this species is rarely observed on the ground and therefore probably is at least as arboreal as M. sinica and M. radiata (Fooden, 1982a, p. 17; Quan Guoqiang, 1960, unpublished field notes, izas 17942-47, 17955-58, Zhung Shan, Yunnan, China). M. thibetana, by contrast, is often ob- served on the ground and appears to be more ter- restrial than arboreal (David, 1874, p. 9; 1875, p. 256; Mell in Matschie, 1912, p. 309; Steward, 1925, p. 80; Fooden et al., 1 985, p. 24); this species often leaves marks on the ground where it travels and forages (Xiong, 1984, p. 4; Fooden et al., 1985, p. 24). Nighttime sleeping is exclusively arboreal in M. sinica (Dittus, 1977b, p. 245) and in M. radiata, except in treeless urban areas (Kuruvilla, 1 980, p. 979; Fooden, 1981, p. 16). Although nighttime sleeping habits have not been reported for M. as- samensis, this species probably also sleeps in tall trees. M. thibetana, however, does not sleep in FOODEN: NATURAL HISTORY OF SINICA MACAQUES trees, but instead sleeps on cliffs or in caves (Xiong, 1984, pp. 3, 8; Fooden et al., 1985, p. 24). M. sinica, M. radiata, and M. thibetana are known to enter streams freely (Fooden, 1979, p. 122; 1981, p. 16; Xiong, 1984, p. 5). This probably also applies to M. assamensis. Food Like other macaques, species in the sinica group are predominantly vegetarian, with animal food constituting less than 5% of the total dietary in- take. Macaca sinica, M. radiata, and M. thibetana are known to eat fruit, flowers, seeds, shoots, leaves, resin, and other parts of a wide variety of plants in their environment; this probably also is true of M. assamensis. Macaca sinica is recorded in one area as eating parts of 4 1 species of trees and 22 or 23 species of shrubs (Dittus, 1977b, p. 247); M. radiata is recorded as eating parts of 36 plant species in one area and about 40 plant species in another (Makwana, 1980, p. 10; Kuruvilla, 1980, p. 981); and M. thibetana is recorded as eating parts of at least 1 9 plant species in one area (Xiong, 1984, p. 4). Seasonal changes in plant foods have been carefully documented for M. sinica by Dittus (1974, ch. 2, pp. 91-102) and for M. radiata by Kuruvilla (1980, pp. 982-985), who also notes age and sex differences in plant feeding patterns. Of all plant parts eaten, fruits generally are the main food of M. sinica, M. radiata, and M. assamensis (Dittus, 1977b, p. 248; Kuruvilla, 1980, p. 981; Fooden, 1981, p. 17; 1982a, p. 17); these three species all eat figs, which constitute more than half the diet in some populations of M. sinica. Bamboo shoots are an important food of M. radiata, M. assamensis, and M. thibetana (Fooden, 1981, p. 17; Xiong, 1984, p. 5; Li Zhixiang, 1973, unpub- lished field notes, kiz: Coll. No. 73231, M. assa- mensis, Satun, Yunnan, China). Leaves may con- stitute a major part of the diet of M. thibetana; of 1 7 food plants tabulated for this species, 1 1 are exploited for their leaves and only 5 for their fruit (Xiong, 1984, p. 4). Leaves also may be important in the diet of some populations of M. assamensis; according to Li Zhixiang (1973, unpublished field notes, kiz: Coll. No. 73231, see above), the diet of one specimen of this species collected in May in Yunnan was 60% leaves and 40% bamboo shoots. All four species in the sinica group are notorious raiders of cultivated fields; among other crops pilfered, rice is habitually taken by M. sinica and M. radiata, and maize is habitually taken by M. assamensis and M. thibetana. Animal food, including both invertebrates and vertebrates, is reported as part of the diet of all four species (M. sinica: Dittus, 1977b, p. 248; Fooden, 1979, p. 123; Farook, 1979, p. 76. M. radiata:. Rahaman & Parthasarathy, 1979, pp. 415, 420-421; Kuruvilla, 1980, pp. 986-987; Fooden, 1981, p. 17. M. assamensis: Fooden, 1982a, p. 17. M. thibetana: Xiong, 1984, p. 4; Fooden et al., 1985, p. 25). Insects, including larvae and adults, probably are the most important prey of these macaques. Also recorded as prey are spiders and spiders' eggs, snails, small fish, lizards, snakes, birds and birds' eggs, palm squirrels, and tree mice. Al- though animal food constitutes less than 5% of the diet of these species, it apparently is nutritionally important: M. sinica spends about 20% of daily foraging time searching for animal prey (Dittus, 1977b, p. 248); M. radiata has been observed to swim across a fast-flowing stream to obtain birds' eggs at a nesting ground (Krishnan, 1972, p. 539); and M. thibetana troop members have been ob- served fighting over the remains of a lizard that one of them had killed (Xiong, 1984, p. 4). Xiong (1984, p. 5) reports that starvation is a frequent cause of death in M. thibetana in winter when food is in short supply for this species. In- tratroop fights over food also are common in this species. During the rainy season, M. sinica and M. ra- diata obtain adequate water from food and from tree holes and wet surfaces (Dittus, 1977b, p. 247; Fooden, 1979, p. 123; 1981, p. 18; Kuruvilla, 1980, p. 980). During the dry season these species make long daily trips to drink at permanent water sources. No information is available concerning how M. assamensis and M. thibetana satisfy their water requirements. Predators and Parasites Known nonhuman predators of M. sinica are crocodiles (Crocodylus palustris), leopards (Pan- thera pardus), and domestic dogs (Canis familiar- is) (Dittus, 1977b, p. 251; Fooden, 1979, p. 122; Farook, 1979, p. 76; cfi Seidensticker, 1983, p. 325); probable additional predators of M. sinica are pythons (Python molurus), cobras (Naja naja), Russell's viper (Vipera russelli), and large eagles and hawks. The same species and, in addition, tigers (Panthera tigris) and smaller felids, probably FIELDIANA: ZOOLOGY also are predatory to M. radiata (Rahaman & Par- thasarathy, 1979, p. 407; Makwana, 1980, p. 11; Fooden, 1981, p. 16). Although predators of M. assamensis and M. thibetana have not been spe- cifically reported, M. thibetana is habitually vig- ilant while foraging (Xiong, 1984, p. 2), which suggests that predation, possibly nonhuman, is a substantial threat. Macaca thibetana troop mem- bers have been observed in a protracted skirmish with an attacking kite (Milvus korschun) (Xiong, 1984, p. 2), which, however, is too small to be a serious predator of these macaques. Parasitological surveys of sinica group species have mainly concentrated on M. sinica and M. radiata (Reardon & Rininger, 1968, p. 578; Fiennes, 1972, symposium; Wong & Conrad, 1978, p. 413; McSwain, 1983, pp. 62, 89, 91). Parasitic organisms reported for these two species include viruses, bacteria, fungi, protozoa, trematodes, ces- todes, nematodes, and arthropods. A tickborne virus, the agent of epidemic Kyasanur Forest dis- ease, is often fatal to M. radiata in Shimoga Dis- trict, India (Trapido et al., 1964, p. 763). Parasitic protozoa and nematodes have been reported in wild-collected M. assamensis in Vietnam and China (Sandground, 1933, pp. 564, 575; Yin, 1973, p. 355; 1980, p. 23; Long et al., 1979, p. 320). Seasonal Migration 11 March, ca. 2500 m (David, 1874, p. 9). At Huang Shan, north-facing slopes are favored in summer and south-facing slopes are favored in winter; this suggests that the migration of M. thi- betana in this area probably is an adaptation to seasonal temperature extremes as well as to local availability of plant foods. Although M. sinica and M. radiata are not known to migrate seasonally, some troops of these two species extend their home ranges considerably during the dry season in order to reach distant permanent sources of drinking water (Koyama, 1973, p. 229; Dittus, 1977b, p. 247). Such seasonal range extension may provide a model for the origin of true seasonal migration, as occurs in M. assamensis and M. thibetana (see Darlington, 1957, p. 243). Troop Size and Composition Judging from available data, troop size is rough- ly similar in all four species in the sinica group (table 1). In forest populations that are relatively independent of humans, mean troop size in these species is about 20 to 25 individuals and extreme troop sizes are about 7 and 70 individuals. In M. radiata, nonforest troops that are more or less closely associated with humans generally tend to be larger than forest troops (table 2; Pirta et al., 1981, p. 433; Singh et al., 1984, p. 10); however, In some areas M. assamensis and M. thibetana are known to migrate seasonally between higher elevations in summer and lower elevations in win- ter. Crump (in Wroughton, 1916, p. 476) indicates that M. assamensis descends to 600-1200 m in winter in the Sikkim-Darjeeling area (cf. Fooden, 1982a, p. 17), and Shou et al. (1964, p. 62) report that this species also habitually descends from the mountains in winter in Guangxi and Yunnan. Xiong (1984, p. 3) provides greater detail con- cerning the seasonal migration of M. thibetana at Huang Shan in southeastern Anhwei: the summer range in this area is at 1 300-1 700 m; the fall range is at 500-700 m; the winter range is at 400-600 m; and the spring range is at 570-1 700 m. In some other areas, M. thibetana evidently remains at higher elevations in fall and winter: NE Jiangxi— Qianshan Xian, Jan., 1000-1200 m (izas 17970); NE Guizhou-Jiangko, Oct., 1200 m (kiz 3193); Fanjing Shan, Dec, 1200 m, 2350 m (Fooden et al., 1985, p. 24); central Sichuan— Emei Shan, Oct., ca. 1 500 m (Wilson, 1 9 1 3, p. 224); Baoxing, NNW, Table 1 . Troop size in forest populations of sinica group species.* Species Mean troop size** Extreme No. of troop troops sizes observed M. sinica 21.6 8-51 28f M. radiata 24.9 ± 12.3 7-50 13 M. assamensis >20 10-50 >25 M. thibetana >20 10-70 >9 * References: M. sinica— Eisenberg & Lockart, 1972, p. 76; Eisenberg et al., 1972, p. 870; Dittus, 1974, ch. 2, p. 66; 1975, p. 137; 1977b, p. 243; 1980, p. 267; cf. Farook, 1979, p. 74. M. radiata— see Table 2. M. as- samensis—Fooden, 1982a, p. 18;QuanGuoqiang, 1960, unpublished field notes, izas \19A2-A1, 1 7955-58, Zhung Shan, Yunnan, China, 10 to 20 individuals in each of three troops. M. thibetana— David, 1875, p. 256; Xiong, 1984, p. 1; Fooden et al., 1985, p. 26. ** Data inadequate for computation of standard de- viations, except in M. radiata. t At least three of these troops are partly dependent on food sources generated by humans (Dittus, 1974, ch. 2, p. 71) and therefore are not typical forest troops. FOODEN: NATURAL HISTORY OF SINICA MACAQUES Table 2. Macaca radiata: troop size in forest troops compared with that in nonforest troops. Forest troops Nonforest troops (urban, rural, roadside, temple) Mean ± SD Ex- tremes N Ex- Mean ± SD tremes N Reference 36.0 ± 2.8 33.7 17.0 3.5 14.1 4V 25.5 14.6 24.9 14.5 4.9 34-38 30-37 7-27 12-50 11-23 1 5* 5§ 8.9 7.4 21.1 28.9 29.5 ± 15.4 23 39.7 ± 5.5 48.5t 36.9 ± 26.2 22.7 ± 9.6 10-43 16-44 15-59 34-45 10-100 7-44 14 10 6* 1 3 1 22$ 28§ ± 12.3 7-50 18 28.2 ± 16.8 7-100 85 17.7 ± 1.40 5-59 308 Nolte, 1955, p. 78 Rahaman & Parthasarathy, 1967, p. 253 Sugiyama, 1972, p. 255; Koyama, 1973, p. 228 Simonds, 1973, p. 601; 1974a, p. 152 Singh & Sachdeva, 1977, p. 606 Rahaman & Parthasarathy, 1979, p. 408 Kuruvilla, 1980, p. 978 Foodenetal., 1981, p. 469 Singh et al., 1984, p. 10 Summary Kurup, 1984b, p. 59 * For more recent census data on some of these troops, see Makwana (1980, p. 10). ** Mean size of troop whose membership varied between 38 and 44 individuals, October 1974-September 1975. t Mean size of troop whose membership varied between 45 and 52 individuals, October 1974-September 1975. t Excludes Elephanta Island troops, which are more accurately reported by Kuruvilla (1980, p. 978) and are tabulated separately. § Some of these troops apparently are also included in the population study of Pirta et al., 1981, p. 433. recently published data on 308 nonforest troops in four Indian states indicate that mean size in such troops probably is smaller than previously inferred (table 2; Kurup, 1984b, p. 59). Provi- sioned temple troops of M. radiata and M. thibe- tana sometimes reach extraordinary sizes of 100 to 300 individuals (Li, 1960, p. 202; Fooden et al., 1981,p.469;Guo, 1981, p. 65). Solitary males, unknown in M. sinica, M. radiata, and M. assa- mensis, have been reported in M. thibetana by three observers (David, 1874, p. 9; Mell in Mat- schie, 1912, p. 309; Steward, 1925, p. 80). In an average troop, about 50 to 60% of the members are sexually mature (Fooden, 1979, p. 125; 1981, p. 25; 1982a, p. 18; Xiong, 1984, p. 2; Singh et al., 1984, p. 12; Kurup, 1984b, p. 63). The ratio of sexually mature males (subadults and adults) to sexually mature females (adults) is 0.60 Table 3, Macaca radiata: ratio of sexually mature males (subadults and adults) to sexually mature females (adults) in 4 1 8 troops. Sexually mature individuals Pooled sex ratios Extreme sex No. of (males : ratios in No. of indi- troops females) troops observed viduals observed observed Reference 0.88 0.12-2.00 330-331 26* Simonds, 1973, p. 600 0.52 0.50-0.56 64 3 Rahaman & Parthasarathy, 1979, p. 409 0.71 0.71-0.71 47-55 2 Kuruvilla, 1980, p. 978 0.79 0.27-1.67 418 27** Fooden, 1981, p. 24 0.61 0.36-1.75 92 9 Koyama & Shekar, 1981, p. 248 0.68 0.25-2.00 181 iot Foodenetal., 1981, p. 470 0.93 [425] 33* Singh etal., 1984, pp. 10, 11 0.69§ [3,028] 308 Kurup, 1984b, p. 59 0.73 0.12-2.00 4,585-4,594 418 Summary * Cf. Makwana (1980, p. 10), who reports more recent data for some of the same troops. ** Excludes troops reported by Simonds ( 1 973, p. 600), which are tabulated separately; also excludes troops reported by Nolte (1955, p. 78), for which age-sex categories probably are not coordinate with those in other studies. t Age-sex data recorded in unpublished field notes on troops reported previously. $ Some of these troops apparently are also included in the population study of Pirta et al., 1981, p. 432. § Excludes subadults, which are not separated according to sex in this study. FIELDIANA: ZOOLOGY Table 4. Home range area in Macaca sinica and Macaca radiata. Mean home range area (hectares) Extremes (hectares) Habitat No. of troops observed Reference >100 ca. 41.3 ± 26.7 >200 26 ca. 100 240 ca. 212 ca. 195 ca. 100 17-115 ca. 130-260 ca. 40-260 ca. 100 Macaca sinica Forest 2 Various 1 3 Forest 1 Macaca radiata Garden 1 Plantation 3 Various 2 Forest 5 Roadside 4 Garden 2 Eisenberg & Lockart, 1972, p. 76; Eisen- bergetal., 1972, p. 870 Dittus, 1974, ch. 2, p. 70; 1977b, p. 250 Hladik & Hladik, 1972, p. 205 Singh &Pirta, 1978, p. 265 Rahaman & Parthasarathy, 1979, p.415 Kuruvilla, 1980, p. 979 Fooden, 1981, p. 22 Fooden, 1981, p. 22 Fooden, 1981, p. 22 in 26 troops of M. sinica (276 mature individuals; Dittus, 1975, pp. 133, 137; cf. Farook, 1979, p. 74), 0.73 in 418 troops of M. radiata (ca. 4,590 individuals; table 3), and 0.43 in 3 troops of M. assamensis (30 individuals; Fooden, 1 982a, p. 1 8). These ratios are not significantly different from 0.72, which is the weighted mean ratio for all three species (chi square, 3.75; df, 2; 0.20 > P > 0.10); the low ratio in M. assamensis suggests possible divergence, but available data for this species are insufficient to establish this. In M. thibetana, al- though no data are available concerning the ratio of sexually mature males to sexually mature fe- males, Xiong (1984, p. 2) reports that the ratio of all males to all females among 115 individuals in 5 troops is 1.80; if confirmed as a general rule in M. thibetana, the great excess of males over fe- males in this species would set it apart from all other studied species of macaques. Two small isolated all-male subgroups have been reported in the vicinity of larger mixed-sex troops. In M. sinica, Dittus (1975, p. 139) observed an all- male group of fluctuating size, maximum 9 to 1 0 (2 adults, 1 subadult, 6 to 7 juveniles), that re- mained isolated for at least 3.5 years peripheral to a troop of 33 to 34 members; 3 nonpermanent juvenile members of the all-male group are known to have detached from the main troop, and another nonpermanent juvenile member detached from a neighboring troop. In M. radiata, Simonds (1973, p. 600) observed a group of 3 males that remained isolated for at least several weeks within the home range of a mixed-sex troop of 28 members; 1 of the males in this isolated group is known to have detached from the main troop, and the other 2 may have also. Home Range Home range area is known from fairly complete data for several troops of M. sinica and M. radiata (table 4) and from fragmentary data for a few troops of M. thibetana; home range area in M. assamensis is unknown. In M. sinica, home range area varies from 17 to >200 hectares, and, in M. radiata, home range area varies from 26 to ca. 260 hect- ares; available data suggest that home ranges may tend to average somewhat smaller in M. sinica than in M. radiata. In M. sinica, home range area is positively correlated with troop size (Dittus, 1977b, p. 250); this probably also applies to M. radiata. In M. thibetana, seasonally migratory troops at Huang Shan, southeastern Anhui, have very large home ranges, each of which encompasses several mountains (Xiong, 1984, p. 5). When adequate food is locally available, a troop at Huang Shan forages within an area less than 2.5 km in diameter (less than 500 hectares). In winter when food is scarce, by contrast, a foraging troop may travel 30 km in one day, at a rate of 6 to 7 km/hr, searching for seeds as it moves. At Fanjing Shan, north- eastern Ghizhou, in late November-early Decem- ber, a troop of M. thibetana remained within an area of less than 200 hectares for four days (Food- en et al., 1985, p. 25). Troop home ranges are known to be relatively stable over many years in M. sinica (3'/2 years; Dittus, 1974, ch. 2, p. 68), M. radiata (16 years; Makwana, 1980, p. 9; Fooden, 1981, p. 22), and M. thibetana (4 years; Xiong, 1984, p. 5); this presumably also applies to M. assamensis. A troop of M. radiata in Mysore City returned to its orig- FOODEN: NATURAL HISTORY OF SINICA MACAQUES inal home range after an unnatural move that last- ed 37 days and took the troop 2 km away from its home range (Singh & Pirta, 1978, p. 265). Home ranges of adjacent troops often overlap, at least in M. sinica and M. radiata (Dittus, 1 977b, p. 250; Rahaman & Parthasarathy, 1979, p. 415; Kuruvilla, 1980, p. 979; Fooden, 1981, p. 22). In overlap areas, intertroop tension is high (Dittus, 1974, ch. 2, p. 68; Rahaman & Parthasarathy, 1979, p. 415; Kuruvilla, 1980, p. 979; Fooden, 1981, p. 23; Xiong, 1984, p. 2). In these areas, larger troops are frequently, but not always, dom- inant to smaller troops. In M. radiata and M. thibetana, fierce intertroop fights have been ob- served in overlap areas. After confrontations in overlap areas, both troops generally withdraw to more remote parts of their respective home ranges. Male Emigration Intertroop transfer of males has been docu- mented by field observations in M. sinica and M. radiata and is suggested in M. thibetana. Although relevant observations are lacking in M. assamen- sis, it seems likely that male transfer is widespread or universal in the sinica group. The pattern in M. sinica, where male transfer has been most inten- sively studied (Dittus, 1975, p. 140), may be typ- ical. In M. sinica, all males probably leave their natal troop as adolescents, sometime between age 3.5 years and age 7.0 years, before their first breed- ing season. Adult males also may emigrate, on average about once every 5 years. Emigration usu- ally occurs during the breeding season. Emigrating males, both adolescents and adults, are seldom or never dominant in the troops they leave, but an emigrating adult sometimes becomes dominant in the troop that he enters. The hardships and com- bats involved in emigration probably are a major cause of mortality in males, particularly in ado- lescent males, in which the annual mortality rate reaches approximately 30%. An apparent genetic consequence of male emigration in M. sinica, as in other macaques, is that the coefficient of rela- tionship within troops, estimated from blood pro- tein polymorphism, is relatively low (0.191 ± 0.054; Aoki & Nozawa, 1984, p. 179) and average heterozygosity of blood protein alleles is relatively high (0.078; Shotake & Santiapillai, 1982, p. 93). Although early field observations of M. radiata suggested that male emigration was rare or absent in this species (Simonds, 1965, p. 186; Rahaman & Parthasarathy, 1969a, p. 275), subsequent stud- ies revealed that it is fairly common (Simonds, 1973, p. 600; Makwana, 1980, p. 9; Pirta et al., 1981, p. 432; Fooden, 1981, p. 29; Ali, 1984, p. 319; cf Wade, 1979, p. 363; Small, 1982, p. 3; Caldecott, 1984, p. 12). In M. radiata, as in M. sinica, none of the emigrants observed was the dominant male in the troop that he left, although one became dominant in the troop that he entered. Further study of emigration in M. radiata is re- quired to clarify similarities and differences be- tween male emigration in this species and M. sin- ica. In M. thibetana, new arrivals from another troop (number and sex of these individuals not specified) have been identified among troop mem- bers of the lowest dominance rank (Xiong, 1984, p. 7). Female emigration evidently is relatively rare in the sinica group. Two instances (of which one is equivocal) are reported in M. sinica (Dittus, 1975, p. 139; Farook, 1979, p. 77), and four in- stances are reported in M. radiata (Rahaman & Parthasarathy, 1969a, p. 268; Moore & Ali, 1984, p. 101; Moore, 1984, p. 579). Seasonal Breeding Breeding seasonality is well documented in M. sinica and M. radiata and less well documented in M. assamensis and M. thibetana (see references below). In all four species, apparently, a well-de- fined period of maximum copulations precedes a well-defined birth season by 5 to 6 months, which is the typical gestation length in macaques (Ardito, 1 976, p. 2 1 5). The frequency of copulations during the peak mating period probably is more than 1 0 times greater than during the rest of the year (Fooden, 1981, p. 27). Occasional reversed peri- odicity of annual mating and birth seasons evi- dently may occur in all four species. Although an- nual reproductive cycles generally are approximately synchronous within each of these four species, they are only partially synchronous between species. In 1 8 troops of M. sinica in the Polonnaruwa area, mating frequency peaks in July-September, and birth frequency peaks in December-February (Dittus, 1975, p. 134; 1977a, p. 312); a few births also are reported in September and October, which may indicate reversed periodicity of mating and birth seasons. At Anuradhapura, a locality about 80 km from Polonnaruwa, the birth season in 5 to 6 troops reportedly is in the spring, slightly later than at Polonnaruwa, and the peak mating period FIELDIANA: ZOOLOGY therefore presumably is in the fall; 2 to 3 troops at this locality had reversed periodicity, with the birth season reportedly in the fall and the peak mating period presumably in the spring. In hi. radiata, at numerous localities from Bom- bay to Mysore, the peak mating period is in Au- gust-November, although matings at low frequen- cy also have been observed in other months (Parthasarathy, 1977, p. 49; Rahaman & Partha- sarathy, 1979, p. 413; Makwana, 1980, p. 1 1; Ku- ruvilla, 1980, p. 977; Fooden, 1981, p. 27). In accord with the well-defined mating frequency peak, 86 of 88 recorded births at these same lo- calities occurred during the period December-June, with 8 1 of these births more narrowly restricted within the period January-April. The two excep- tional birth records, one in September (Rahaman & Parthasarathy, 1979, p. 422) and the other in October (Kuruvilla, 1980, p. 979), suggest re- versed periodicity. The peak mating period (August-November) and birth season (January-April) in M. radiata ap- proximately coincide with the corresponding re- productive seasons in most troops of hi. sinica at Anuradhapura (peak mating period, fall; birth sea- son, spring) and are only one or two months later than the corresponding seasons in M. sinica at Polonnaruwa (peak mating period, July-Septem- ber; birth season, December-February). Although known annual reproductive cycles are approxi- mately synchronous in these two species, these cycles are inversely correlated with local wet and dry seasons. In most parts of the range of M. ra- diata, the peak mating period is at the end of the rainy season, and the birth season is at the end of the dry season (Fooden, 1981, p. 27); in hi. sinica, however, matings at Polonnaruwa and Anurad- hapura peak at the end of the dry season, and births peak at the end of the rainy season (Lancaster & Lee, 1965, p. 509; Dittus, 1977c, p. 269). The significance of synchronism with respect to the calendar and of divergence with respect to local wet-dry seasons in the annual reproductive cycles of these two species is unclear. In hi. assamensis, 8 of 10 records available from eastern India to northern Vietnam (Fooden, 1982a, p. 20) indicate that births probably peak in the summer (rainy season) and that matings therefore probably peak in the winter (dry season). The re- maining two records from natural populations of this species indicate births in winter, which sug- gests reversed periodicity. In a zoo in Germany, four infants that were produced in nine years by a captive female hi. assamensis were all born in the spring— successively in May, March, May, and April (Dathe, 1983, p. 126). In a laboratory pop- ulation of 1 0 hi. assamensis females, reproductive seasonality was individually variable, and re- sponse to experimental changes in photoperiod also was individually variable (Wehrenberg & Dy- renfurth, 1983, p. 120). Evidence concerning breeding seasonality in M. thibetana is ambiguous. Reports from three widely separated localities, one in the western part of the range, one in the central part, and one in the east- ern part, suggest that matings in this species peak in the spring and that births peak in the fall (Food- en et al, 1985, p. 26). However, data from two additional localities in the eastern part of the range, one of them particularly well studied, indicate re- versed periodicity, with matings reportedly most frequent during the fall (late September-early No- vember; Xiong, 1984, p. 6) and one birth record apparently in the spring (Fooden, 1983, p. 13; C. Pope, 1926, unpublished field note, American Museum of Natural History, N.Y., Specimen No. 84475, Chong'an Xian, Fujian). Numerous observations are available concern- ing breeding seasonality in captive 'hi. radiata. A colony of this species kept outdoors at Davis, Cal- if, has maintained annual reproductive season- ality for more than 10 years (Judge & Rodman, 1976, p. 535;Glick, 1979, p. 272; Silk etal., 1981, p. 1 109). The mating season in this colony starts in August and peaks in October and November, as in India; matings in the Davis colony then con- tinue at a moderately high level for another three months, December-February, which are not in- cluded in the peak mating season in India (Fooden, 1981, p. 27). As an apparent consequence of the more protracted mating season, the peak birth sea- son in the Davis colony (March-June) is two months later than the peak birth season in India (January-April; Fooden, 1 98 1 , p. 27). Glick (1979, p. 272) notes that the mating season of hi. radiata at Davis generally coincides with months of high humidity and precipitation, and the birth season coincides with months of low humidity and pre- cipitation; the same relationship between repro- ductive seasons and wet-dry seasons also applies to hi. radiata in India (see above). Captives in two other hi. radiata colonies also maintain breeding seasonality, approximately as in natural populations. An indoor colony kept at Lucknow, India, apparently has a peak mating pe- riod in September-October (Kamboj et al., 1984, p. 493), and an outdoor colony of hi. radiata at Lawrenceville, Ga., has a birth season in April- FOODEN: NATURAL HISTORY OF SINICA MACAQUES August, with a strong peak in April (Hadidian & Bernstein, 1979, p. 433), approximately as in the Davis colony. In two indoor colonies, one in Vir- ginia and one in California, however, conceptions and births evidently occur through most of the year (Valerio et al., 1969, p. 291; Redman & Schneider, 1979, p. 7); the Virginia colony appar- ently exhibited breeding seasonality during the first year of captivity, but lost seasonality in the second year. In the outdoor Davis colony and the indoor Lucknow colony, serum testosterone level, mea- sured in the morning, reaches an annual peak dur- ing the mating season; this peak is approximately double the nonmating season minimum (Glick, 1979, p. 277; Kamboj et al., 1984, p. 492). Serum testosterone level in M. radiata also exhibits a daily cycle, with a nighttime maximum that is 6 to 1 1 times greater than the daytime minimum (Mukkuetal., 1976, p. 779; Kholkuteetal., 1981, p. 428; Kamboj et al., 1984, p. 492); the nighttime increase has been experimentally abolished by ex- posure to constant light (Mukku et al., 1976, p. 779). Correlated with the mating season testos- terone peak in the Davis colony is a seasonal in- crease in testis size (length and width increase by about 30%) and a seasonal increase in male body weight (about 5% increase) (Glick, 1979, pp. 271, 274). Seasonal testosterone cycles apparently dis- appear in some indoor colonies of M. radiata (Murty et al., 1979, p. 158; Munshi, 1980, p. 13; Murty et al., 1980, p. 52); this may be related to different conditions of captivity. Semen volume and sperm number, studied by electroejaculation, show no seasonal change in one indoor colony of M. radiata (Sharma et al., 1984, p. 235). Captive females of M. radiata kept in two in- door colonies in India are reported to become an- ovulatory and exhibit amenorrhea from April to July in Bangalore (Murty et al., 1979, p. 153; Sri- nath, 1980, p. 19) and from July to September in Bombay (Munshi, 1980, p. 11); most of these months are outside of the peak mating period in natural populations (Fooden, 1981, p. 27). In the Bangalore colony, ovarian and menstrual cycles resume during the period September-December, which approximately corresponds to the peak mating period in natural populations; no infor- mation is available concerning the date of cycle resumption in the Bombay colony. In Bangalore, newly captured females, obtained at any time of year, were found to exhibit amenorrhea for six or more months after introduction into the colony; these females began to cycle regularly when they were kept for two or three months within sight of a separately caged mature male in the same room (Madhwa et al., 1975, p. 394; Srinath, 1980, p. 1 7). In a colony of M. radiata in Bombay, Jay- araman et al. (1979, p. 683) report that newly in- troduced females began to cycle regularly after al- falfa was added to their diet. Mating Behavior Several accounts, partly divergent, are available concerning mating behavior in natural popula- tions of M. radiata (Simonds, 1965, p. 193; 1977, p. 161; Rajagopal, 1965, p. 232; Rahaman & Par- thasarathy, 1969a, p. 275; 1969b, p. 154; 1971, p. 99; 1979, p. 422; Sugiyama, 1972, p. 252; Par- thasarathy, 1977, p. 49; Ali, 1984, p. 319); differ- ences between these accounts may indicate indi- vidual or local differences in the behavior of this species. Although observers agree that subordinate males in M. radiata are not excluded from access to estrous females during the peak mating period, most reports suggest that dominant males engage in proportionally more copulations than subordi- nate males. Some studies indicate that dominant males may form relatively stable consort relation- ships with dominant females, whereas subordinate males and females tend to be more promiscuous in their matings. Intratroop social tensions appar- ently increase during the peak mating period. Dominant males sometimes interfere with copu- lations of subordinate males (4 of 308 attempted copulations in one study), but peaceful serial mountings of one female by two or three males, including dominant and subordinate males, also have been reported. Subordinate males and es- trous females sometimes withdraw 500 m from other troop members before copulation. Most copulations in M. radiata are initiated by males (55 of 59 attempted copulations in one study; 24 1 of 308 attempted copulations in another). Sex- ual skin is relatively inconspicuous in females of M. radiata, and, unlike females in some other ma- caque species, they rarely solicit copulation by ap- proaching males and presenting their hindquar- ters. Males apparently recognize estrous females by olfactory cues, probably associated with cycli- cal changes in cervical mucus (Ovadia et al., 1971, p. 132;McArthuretal., 1972, p. 109; Sheth et al., 1975, p. 134; Nasir-ud-Din et al., 1979, p. 397). Before mounting, a male often examines a female's perineum, frequently with digital probing of the vagina, and smells or tastes the vaginal contents, 10 FIELDIANA: ZOOLOGY which probably consists primarily of cervical mu- cus. During the peak mating season, some males examine the perineum of most or all sexually ma- ture females every day. Copulation apparently is completed in less than one minute and usually is accomplished in a single mounting (26 of 28 cop- ulations in one study; 294 of 308 in another), with 5 to 30 pelvic thrusts to ejaculation. Following copulation, excess ejaculate on the female's peri- neum is sometimes eaten by one or both partners or by another male. Copulations may occur either on the ground (197 of 362 observed copulations in one study; 54%) or in trees (46%). Males may attempt to copulate with unreceptive females, who express their unwillingness by vocalizing and fleeing (3 1 of 308 attempted copulations in one study). Homosexual mountings, not consistently corre- lated with social dominance, have often been ob- served in At. radiata. Mating behavior in a long-established captive colony of M. radiata is generally within the range of variation observed in natural populations (Glick, 1980, p. 350; Shively et al., 1982, p. 375; Samuels et al., 1984, p. 998), despite the unnatural preclu- sion of male emigration in the captive colony. In this colony, Small (1983, p. 127) reports that fe- males without infants enter estrus earlier and begin copulations earlier than females with infants. In an experimental group of males and ovariectom- ized females, sexual behavior was reduced relative to that in a control group of males and intact fe- males (Rasmussen, 1984, p. 353). Limited information is available concerning mating behavior in natural populations of M. sin- ica and hi. thibetana (Jay, 1965, p. 570; Dittus, 1974, ch. 2, p. 109, ch. 3, p. 35; 1977a, p. 311; Xiong, 1984, pp. 5, 7), and none is available for M. assamensis. Mating behavior in captive M. sinica has been reported by Hill (1939, pp. 25, 3 1; 1 974, p. 7 1 7). In M. sinica and M. thibetana, threats and fighting between males over access to estrous females may be more frequent and violent than in M. radiata (but see Singh et al., 1984, p. 15); serious injuries reportedly are inflicted in such fights in M. sinica and M. thibetana. In M. sinica, a consort pair typically withdraws up to 300 m away from its troop, apparently to reduce contact with competing males; somewhat similar behavior also has been reported in M. radiata (see above). In M. thibetana, high-ranking males reportedly intimi- date lower-ranking males and thereby monopolize most of the estrous females. In M. sinica, as in M. radiata, sexual skin in females is inconspicuous, and copulation apparently is usually initiated by males (Hill, 1974, p. 717— captives); in M. thi- betana, by contrast, sexual skin is reported to swell conspicuously during estrus, and females have been observed to initiate copulations by approaching and presenting to males. In both M. sinica and M. thibetana, a profuse vaginal discharge— presum- ably cervical mucus— has been noted during es- trus. As in M. radiata, males in M. sinica and M. thibetana have been observed to inspect the female perineum before copulation; this includes digital examination and smelling or tasting the vaginal discharge. Copulation in M. thibetana reportedly is completed in 40 to 60 seconds, which presum- ably indicates that ejaculation is accomplished in a single mount. Attempts by males to force cop- ulation on unreceptive females have been ob- served in M. thibetana, as in M. radiata. Life History Available life history information is relatively good for M. sinica and M. radiata and relatively poor for M. assamensis and M. thibetana. Length of the gestation period is best known in M. radiata. Valerio et al. (1969, p. 295) report that the ges- tation period in a captive colony of this species is 159 ± 6 days (mean ± SD) for 5 female births, 163 ± 3 days for 11 male births, and 162 ± 4 days for both sexes combined. Similar gestation periods in M. radiata also are reported by Hart- man (1938, p. 470)— 166 days for one female and 169 days for one male (cf. 153 days for one still- born male); Souri and Swani (1962, p. 83)— ca. 165 days for one birth, sex unspecified; Stabenfeldt and Hendrickx (1972, p. 619)— ca. 167 days for four births, sex unspecified; Jayaraman et al. (1979)— 1 79 ± 4 days for four males; and Srinath (1980, p. 20)— 166 days for 27 births, sex un- specified. In M. sinica. Hill (1966, p. 138) reports a gestation period of 1 80 days, without specifying the number or sex of births observed; the mean gestation period in this species probably is less than 1 80 days, judging from known mean gesta- tion periods for small macaques (cf. Ardito, 1976, p. 2 1 5). In M. thibetana, Xiong (1984, p. 6) reports that the gestation period for two captive births (sex unspecified) was about eight months; if confirmed as typical for this species, this would be about as long as the gestation period in mandrills (Ardito, 1976, p. 216). Prenatal mortality information is available for captive M. radiata maintained at Davis, Calif. (Hendrickx & Nelson, 1971, p. 414; Hendrickx & FOODEN: NATURAL HISTORY OF SINICA MACAQUES 11 Binkerd, 1980, p. 48). In an indoor group of this species, the prenatal mortality rate was 26.0% of 227 pregnancies during nine years; 16.8% of these pregnancies resulted in spontaneous abortions, and 9.2% resulted in stillbirths. In an outdoor group, the prenatal mortality rate was markedly higher, 41.0% during a two-year period (number of preg- nancies unspecified). The sex ratio at birth in a few sampled natural populations of M. sinica and At. radiata is close to 1:1. In two local populations of At. sinica, this ratio is 57 males : 54 females (1.06), which is not significantly different from 1.00 (Dittus, 1975, p. 132). In a smaller sample of three local popula- tions of At. radiata, the sex ratio at birth is 1 7 males: 15 females (1.13) (Fooden, 1981, p. 31). In captive populations of At. radiata, the sex ratio at birth appears to be highly variable. In a colony of this species maintained in Virginia, Valerio et al. (1969, p. 295) report that the sex ratio at birth during a two-year period was 1 1 males : 5 females (2.20), and, in a colony maintained in California, Redman and Schneider (1979, p. 7) report that this ratio during a seven-year period was 118 males: 140 females (0.84). In a subsequent study of a separately confined part of the California col- ony, Silk et al. (1981, p. 1113) report that the sex ratio at birth during a 1 0-year period was 1 1 0 males : 7 1 females (1.55). This ratio fluctuated an- nually from a low of 0.72 to a high of 3.60; annual fluctuations in the sex ratio at birth of this captive group were found to be correlated with differential survival of male and female infants during the preceding year and with dominance rank-order of mothers. No information is available concerning sex ratio at birth in M. assamensis and M. thi- betana. Birth weight in captive At. radiata reported by Valerio et al. (1969, p. 295) is 388 ± 34 g (mean ± SD) for 5 females and 411 ± 56 g for 1 1 males. Similar captive birth weights in this species also have been reported by Hartman (1938, p. 470), 330 g for one female and 370 g for one male, and by Leutenegger ( 1 977, p. 1 54), mean 368 g for four newborn, sex unspecified. Jayaraman et al. (1979, p. 683) note that weight in four captive-born male infants (361 ± 40 g) averages greater than in one wild-born male infant (316 g). Composite mean birth weights in captivity are 378 g for females (N = 6) and 396 g for males (N = 16). The ratio of mean birth weight to mean adult weight in captive At. radiata is 0. 1 0 for females and 0.038 for males. These ratios are based on mean adult weights of 3.753 kg for 10 captive females (Leutenegger, 1977, p. 154) and 10.550 kg for 5 captive males (Glick, 1979, p. 271; cf. Rosenblum & Smiley, 1980, p. 250); captive adult males apparently weigh about 50% more than adult males in natural populations (see Fooden, 1981, p. 3). In At. thibetana, weight of one newborn or young infant (sex unspecified) recorded by Xiong (1984, p. 6) is 550 g; the ratio of this infant's weight to mean adult weight (14.8 ± 4.22 kg, both sexes combined; Xiong, 1984, p. 2) is 0.037. Infant mortality, from birth to age one year, apparently averages about 45% in natural popu- lations of At. sinica (Dittus, 1980, p. 268) and about 50% in natural populations of At. radiata (Fooden, 1981, p. 32). In M. sinica, Dittus (1975, pp. 135, 137) reports that the mortality rate of female infants significantly exceeds that of male infants at one locality and vice versa at another locality; in At. radiata, Singh et al. (1984, p. 12) report that the mortality of female infants exceeds that of male infants in nonurban habitats and vice versa in urban habitats. In a captive population of At. radiata, Redman and Schneider (1979, p. 6) report that infant mortality during the first six months of life varied from 1 1.8% in indoor in- dividual cages to 4 1 % in an outdoor group enclo- sure. In a subsequent 10-year study of the same outdoor group, Silk et al. (1981, p. 1111) report that infant mortality during the first six months of life was 48% (61 deaths/ 126 infants) and was particularly intense during the first month of life (57/61 deaths). In this study, infant mortality was highest in female offspring of low- ranking mothers and also tended to be high in offspring of young mothers (less than age six years). Spontaneous abortions, stillbirths, and neonatal deaths in this colony were not correlated with either primagra- vidity or gravidity order (Small & Rodman, 1981, p. 165). Weaning in natural and captive populations of At. radiata occurs at approximately age six months (Kaufman & Rosenblum, 1966, p. 240; 1969, p. 51; Rahaman & Parthasarathy, 1969b, p. 156; Singh etal., 1980, p. 105; Cantwell & Reite, 1984, p. 429; cf. Simonds, 1965, p. 193); the weaning period generally coincides with the beginning of the next mating season for the mother. Weaning in At. thibetana also occurs about age six months (Xiong, 1984, p. 6). In captive At. sinica, menstruation apparently begins about age 2.5 to 3.0 years, and length of the normal menstrual cycle is estimated to be 29 12 FIELDIANA: ZOOLOGY days (Hill, 1939, pp. 23, 34). In M. radiata, mean length of the menstrual cycle is 28 ± 4.3 days (SD; 1 ,262 cycles, 1 30 females), modal length is 28 days (ca. 650 of 1 ,260 cycles), and extreme cycle lengths are 15 and 42 days (Srinath, 1980, p. 20); similar cycle lengths are reported for smaller samples of M. radiata by Hartman (1938, p. 468), Valerio et al. (1969, p. 287), McArthur et al. (1972, p. 108), Stabenfeldt and Hendrickx (1972, p. 614), Madh- waRaj etal. (1975, p. 395), Jayaraman et al. (1979, p. 683), and Munshi (1980, p. 11). The mean du- ration of menstrual flow in captive M. radiata is reported as 3.8 days (N = 17 females) by Valerio et al. (1969, p. 286) and as 4 ± 1.05 days (SD; N= 1,262 cycles) by Srinath (1980, p. 20; cf. Madhwa Raj et al., 1975, p. 395). During the peak mating season, females in natural populations of M. radiata probably are in estrus for four or five days during each cycle (Rahaman & Parthasara- thy, 1971, p. 100). InM. assamensis, Wehrenberg et al. (1980, p. 523) report that length of the men- strual cycle is 32 ± 2 days (mean ± SD; 14 captive females). Sexual activity begins prior to completion of physical development (Phillips, 1926, p. 267). In natural populations of M. sinica, Dittus (1977a, pp. 283, 308) reports that late juvenile females and late juvenile males (age ca. 3.5 years) engage in sexual approaches and consortships with adults; at this age, males begin to emigrate from their natal troops and suffer a concomitant high rate of mor- tality (Dittus, 1975, p. 136). In natural populations of M. radiata, the youngest females and males that were observed to participate in copulations were age ca. 4.5 years (Rahaman & Parthasarathy, 1969b, p. 155; Sugiyama, 1972, pp. 252, 259); Simonds (1974b, p. 91) indicates that both sexes in this species may be capable of copulation at age ca. 3.5 years. In captive M. radiata, females engage in copulation as early as age 2.5 to 3.0 years (Silk et al., 198 1, p. 1110) and males, as early as age 2 years (Nadler & Rosenblum, 1971, p. 101; Glick, 1980, p. 350). Sperm production in captive M. radiata begins about age 3 years (Nadler & Ro- senblum, 1971, p. 101). In captive M. thibetana, infertile copulations occur as early as age 2 to 3 years (Xiong, 1 984, p. 5). The age at first pregnancy in natural populations of M. sinica is estimated by Dittus (1975, p. 129) to be 4.5 to 5.5 years. This probably also is the age at first pregnancy in natural populations of M. radiata (Sugiyama, 1972, p. 253; Koyama, 1973, p. 228) and M. thibetana (Xiong, 1984, p. 5). In captivity, M. radiata females may become preg- nant as early as age 2.8 years, and viable infants have been produced by females as young as age 4 years (Silk et al., 1981, p. 11 10). The mean fertility rate in a natural population of M. sinica studied for four years is 0.688 infants/ adult female/year (Dittus, 1975, p. 131). In five natural populations of M. radiata studied for 1 to 2 years, the corresponding composite mean is 0.794 infants/adult female/year (Kuruvilla, 1980, pp. 978, 979; Fooden, 1981, p. 31); this agrees fairly well with a preliminary report by Kurup (1984b, p. 61), who indicates that the mean fertility rate in an unspecified number of troops of M. radiata is 0.73 infants/adult female/year. In captive pop- ulations of M. radiata, the reported mean fertility rate varies from 0.630 to 0.734 (Valerio et al., 1969, p. 293; Silk et al., 1981, p. 1 108). Adult size in M. sinica in natural populations is achieved at age 4.5 to 5.0 years in females and at age 7.0 to 8.0 years in males (Dittus, 1975, p. 129). During growth in this species, females are almost the same size as males until age ca. 5 years when growth in females ceases; males subsequent- ly grow (as subadults) for an additional 2 to 3 years and ultimately achieve an adult weight (5.72 kg) that is about 60% more than adult weight in fe- males (3.59 kg) (Dittus, 1974, ch. 1, p. 36). Adult size in M. radiata females and males in natural populations apparently is attained about the same age as in M. sinica (Simonds, 1974b, p. 91), and this evidently also is true in captive populations of M. radiata (Glick, 1979, p. 271; Rosenblum & Smiley, 1980, p. 250). The same growth pattern probably also applies to M. thibetana (Xiong, 1 984, p. 2) and, inferentially, to M. assamensis. Menopause in natural populations of M. sinica is estimated to occur at age 25 to 30 years (Dittus, 1975, p. 132); fertility apparently is unimpaired until menopause. Postmenopausal females were estimated to constitute about 6% of adult females in this population of M. sinica. Postmenopausal females also have been reported in natural pop- ulations of M. thibetana (Xiong, 1984, p. 7). In captive groups of M. radiata, copulations have been reported in females at age 16.5 years, and a viable infant was born to a female about age 16 years (Jensen et al., 1980, p. 400; Jensen & Blan- ton, 1 98 1 , p. 1 90). In another captive population of this species, females became pregnant until at least age 16 years, but no female older than age 1 3 years produced an infant that survived 6 months (Silk et al., 1981, pp. 1 1 10, 1 1 12). An experimen- FOODEN: NATURAL HISTORY OF SINICA MACAQUES 13 tal comparative study of older (age ca. 16.5 years) and younger (age ca. 1 0 years) mature M. radiata females indicates that older females evoke fewer copulations, that their mating behavior is less cor- related with estrous cycle phase, and that during estrus their peak plasma estradiol level is dimin- ished (Jensen et al., 1982, p. 451); a 19-year-old M. radiata male used in this study apparently had unimpaired virility. A captive female M. assa- mensis that lived to age ca. 28 years conceived until age ca. 18 years; this female produced her last viable infant when she was age ca. 14 years, and she produced two stillbirths at ages ca. 17 years and ca. 18 years (Dathe, 1983, p. 126). Assuming that a female's first pregnancy occurs about age 5 years and that her last successful preg- nancy occurs sometime between age 1 5 years and age 27 years and assuming a fertility rate of 0.70, an average female that reaches sexual maturity would be expected to produce 8 to 1 6 infants dur- ing her reproductive lifetime. If infant mortality rate is 0.50, 4 to 8 of these offspring may survive to age 1 year. Longevity records in captivity are 33 to 35 years for M. sinica (Hill, 1937, p. 255), ca. 30 years for M. radiata (Prater, 1980, p. 36), and ca. 28 years forM assamensis (Dathe, 1983, p. 1 29). Although aged individuals have been observed in natural populations of all four species in the sinica group (Simonds, 1965, p. 180; Dittus, 1975, p. 130; Fooden, 1982a, p. 19; Xiong, 1984, p. 2), esti- mated average life expectancy at birth is only about 15% of the maximum potential life-span, ca. 4.7 years in natural populations of M. sinica (Dittus, 1975, p. 135) and ca. 5.0 years in M. radiata (Fooden, 1981, p. 33). Geographic and Ecological Relationships with Other Primates Four levels of geographic and ecological rela- tionships of sinica group macaques are discussed in the following paragraphs. These are: (1) allo- patry and parapatry between species in the sinica group; (2) broad sympatry between sinica group macaques and non-sinica group macaques; (3) marginal sympatry and parapatry between sinica group macaques and non-sinica group macaques; (4) sympatry between sinica group macaques and gibbons, leaf monkeys, and lorises. Within the sinica group, only M. assamensis and M. thibetana are in geographic contact (fig. 1). Macaca sinica in Sri Lanka is separated from M. radiata in peninsular India by the Palk Strait, and M. radiata is separated from M. assamensis in the Himalayan foothills by a 1,200-km gap in central India. Macaca assamensis and M. thibetana, by contrast, are parapatric in southeastern China. The latitudinal relationship between the ranges of M. assamensis and M. thibetana suggests that these two species probably are differentially adapted to warmer and cooler climates, respectively. No in- formation is available concerning ecological or be- havioral relationships between M. assamensis and M. thibetana in the interspecific contact zone. Although three sinica group species— M. radia- ta, M. assamensis, and M. thibetana— zxt broadly sympatric with macaques in other species groups, the sinica group macaques and sympatric non- sinica group macaques are segregated ecologically (Fooden, 1982b, p. 574). In peninsular India, where M. radiata is sympatric with M. silenus, M. ra- diata typically inhabits deciduous forests or dis- turbed habitats, whereas M. silenus is restricted to upland broadleaf evergreen rainforest (Fooden, 1981, p. 18); these two species sometimes meet in transitional habitat areas (Groombridge, 1984, p. 146), but the pattern of interspecific interaction in such meetings remains unclear. In the Indochinese Peninsula, M. assamensis is sympatric with both M. mulatta (see Dao, 1978, p. 382) and M. arc- toides; M. assamensis, however, inhabits broad- leaf evergreen forest and is mainly arboreal and, therefore, is segregated by habitat from M. mu- latta, which inhabits secondary and deciduous for- ests, and by ecological niche from M. arctoides, which inhabits broadleaf evergreen forest but is mainly terrestrial (Fooden, 1982a, p. 21). Macaca thibetana is sympatric with M. mulatta in east- central China, where M. thibetana apparently in- habits broadleaf evergreen forest, and M. mulatta inhabits secondary and disturbed habitats (Foo- den, 1983, p. 14; Fooden et al., 1985, p. 24). Macaca radiata, M. assamensis, and M. thibe- tana are marginally sympatric or parapatric with other non-sinica group species of macaques. In peninsular India, M. radiata is marginally sym- patric with the ecologically similar species M. mu- latta along the 1,000-km border between their ranges (Fooden et al., 1981, p. 465; Saha, 1984, p. 163). At the southeastern end of this border area, a small population of M. radiata is isolated ca. 1 50 km within the range of M. mulatta. At a few localities where M. radiata and M. mulatta have been observed in contact, there is no overt aggression between these species, and, in fact, har- monious mixed-species troops have been recorded 14 FIELDIANA: ZOOLOGY Fig. 2. Macaca radiata young adult male (right, long tail) following M. mulatta adult female (left, short tail), with another M. mulatta individual sitting nearby; these monkeys are part of a mixed-species troop (1 to 3 M radiata members, ca. 25 M. mulatta members) that was observed at Kondra Mutla (16°08'N, 79°46'E), Andhra Pradesh, India, 30 April 1980 (Fooden et al., 1981, p. 465). (fig. 2). In the Indochinese Peninsula, M. assa- mensis is parapatric or marginally sympatric with M. nemestrina and M. fascicularis (Fooden, 1 982a, p. 23); M. nemestrina, like M. assamensis, inhab- its broadleaf evergreen rainforest, whereas M. fas- cicularis inhabits secondary, deciduous, and dis- turbed habitats. The latitudinal replacement of M. radiata by M. mulatta in the Indian Peninsula and the corresponding replacement of M. assamensis by M. nemestrina at approximately the same lat- itude in the Indochinese Peninsula suggest parallel climatic differentiation in these otherwise ecolog- ically similar pairs of species. The relationship be- tween M. assamensis pelops and M. mulatta in the central and eastern Himalayan foothills also seems to be one of marginal sympatry or parapatry (Fooden, 1982a, p. 24); this probably is a conse- quence of the relatively sharp altitudinal zonation in this area of midaltitude broadleaf evergreen for- est, which is the habitat of M. assamensis, and lower altitude deciduous forest, which is the hab- itat of M. mulatta. Other genera of primates with which sinica group species are sympatric are Hylobates (gibbons), Presbytis and Pygathrix (leaf monkeys), and Loris and Nycticebus (lorises); the taxonomy used here for nonmacaque genera and species follows Hon- acki et al. (1982, p. 238). Macaca assamensis is sympatric with Hylobates spp. in the Indochinese Peninsula, where M. assamensis and gibbons in- habit the same broadleaf evergreen forests and have similar, mainly frugivorous diets (Fooden, 1982a, p. 2 1). Macaca assamensis and sympatric gibbons differ, however, in their techniques and efficiency of harvesting arboreal fruit. Dietary differences separate sinica group species, which are primarily frugivorous, from sympatric leaf monkeys (Presbytis spp. and Pygathrix spp.). In Sri Lanka and peninsular India, M. sinica and M. radiata are sympatric with Presbytis spp. FOODEN: NATURAL HISTORY OF SINICA MACAQUES 15 (Fooden, 1979, p. 123; 1981, p. 19; Herzog & Hohmann, 1984, p. 195). Macaca radiata and Presbytis entellus are often closely associated, sometimes in mixed-species bands. In interspe- cific encounters, M. radiata generally appears to be dominant to P. entellus. In southern peninsular India, south of ca. 15°N, M. radiata apparently is more widely distributed in disturbed areas than is P. entellus, which tends to be restricted to forested tracts in this region (Kurup, 1 984a, p. 253); farther north, however, in northern Karnataka and central Maharashtra, P. entellus is widely distributed in disturbed areas (Fooden et al., 1981, p. 465; Ku- rup, 1984a, p. 253; cf. Prasad et al., 1979, p. 726). In the Indochinese Peninsula, M. assamensis as- samensis is sympatric with other species of Pres- bytis, sometimes occurring together in the same tree (Fooden, 1982a, p. 21). Sub-Himalayan M. assamensis pelops is sympatric with P. entellus in the western part of its range (west of the Tista River and Ch'u-mu-pi Shan-ku) and with P. geei and P. pileata in the eastern part (east of the San- kosh River) (see Fooden, 1975, p. 109). In China and northern Vietnam, M. assamensis and M. thibetana evidently are partly sympatric with Pygathrix spp. (subgenus Rhinopithecus). In southwestern China, Pygathrix roxellanae (in- cluding subspecies roxellanae and bieti) probably inhabits higher elevations than M. assamensis and M. thibetana (Li et al., 1981, p. 9; Fooden, 1983, p. 1 3). In northern Guizhou, P. brelichi apparently occurs within the range of M. thibetana, and, in northern Vietnam, P. avunculus apparently occurs within the range of M. assamensis, but little is known of the natural history of these latter two species of Pygathrix (Groves, 1970, p. 569; Quan & Xie, 1981, p. 113; Happel, 1982, p. 292). In Sri Lanka and peninsular India, M. sinica and M. radiata are sympatric with Loris tardigra- rf«5(Pocock, 1939, pp. 38, 177; Fooden, 1979, p. 123), and, in the Indochinese Peninsula, M. as- samensis is sympatric with Nycticebus spp. (Foo- den, 1982a, p. 21). Because both genera of lorises are nocturnal and mainly insectivorous, there probably is little direct ecological interaction be- tween sympatric macaques and lorises. Summary 1 . In the ranges of M. sinica and M. radiata, climate varies from tropical wet to tropical arid; in the range of M. assamensis, climate is subtrop- ical moist; and in the range of M. thibetana, cli- mate is subtropical moist to moderate continental moist. 2. Macaca sinica inhabits deciduous forests and broadleaf evergreen forests, M. radiata prefers de- ciduous forests and disturbed habitats, and M. as- samensis and M. thibetana are mainly restricted to broadleaf evergreen forests. 3. Macaca sinica, M. radiata, and M. assamen- sis are predominantly arboreal, whereas M. thi- betana apparently is predominantly terrestrial. 4. All four species are mainly vegetarian, con- suming various parts of a wide variety of plants, but they also all eat invertebrates and small ver- tebrates. The diet of M. thibetana may include a greater proportion of leaves than that of the other three species. 5. Nonhuman predators of these monkeys in- clude large reptiles, raptorial birds, and carnivo- rous mammals. Known parasites include viruses, bacteria, fungi, protozoa, trematodes, cestodes, nematodes, and arthropods. 6. At some localities, the two northern species, M. assamensis and M. thibetana, migrate season- ally between higher altitudes in summer and lower altitudes in winter. 7. Troop size in forest populations of all four species generally averages about 20 to 25 individ- uals and varies from about 7 to 70 individuals, but troops provisioned by humans may be larger. Solitary males are known only in M. thibetana. 8. The weighted mean ratio of sexually mature males to sexually mature females is 0.72 in troops of M. sinica, M. radiata, and M. assamensis; males may outnumber females in troops of M. thibetana. 9. Troop home range area in M. sinica and M. radiata varies from 1 7 to > 200 hectares. 10. Males in M. sinica and M. radiata are known to emigrate from their natal troop before their first reproductive season. 1 1 . Matings and births exhibit well-defined an- nual peaks in all four species; breeding seasonality sometimes disappears in captive populations. 12. In all four species, extruded cervical mucus probably serves as an important signal of estrus, and all four species probably are single- mount ejaculators. Species in this group apparently differ in intensity of overt male competition for mates and in precopulatory behavior of estrous females. 13. Available life history information indicates that gestation period and birth weight are greater in male infants than in female infants; sex ratio at birth is near unity in natural populations, but is highly variable in captive populations; infant mor- 16 FIELDIANA: ZOOLOGY tality during the first year of life is about 50%; weaning occurs at about age 6 months; males and females in natural populations begin mating at about age 3.5 to 4.5 years; the age of first pregnancy in females in natural populations is about 4.5 to 5.5 years; mean fertility rate is about 0.70 infants/ adult female/year; physical growth ceases at about age 4.5 to 5.0 years in females and at about age 7.0 to 8.0 years in males; maximum longevity in captivity averages about 30 years. 14. Species in this group are segregated from each other geographically, and they are segregated from macaques in other species groups ecologi- cally or geographically. 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On the nematodes of mammals from the Dehong area, Yunnan, China. Acta Zootaxonomica Sinica, 5: 22-29. FOODEN: NATURAL HISTORY OF SINICA MACAQUES 21 Appendix Macaca assamensis Locality Record Supplement In addition to localities shown in previously published distribution maps (Fooden, 1982a, pp. 4, 5), Figure 1 includes the following supplemen- tary records of M. assamensis. All coordinates are approximate. NEPAL: Bhargu vicinity, 28°08'N, 85°18'E; Manigaon-Ramche, 28°02'N, 85°13'E (Green, 1981, p. 80). BANGLADESH: Sylhet Forest, N, 25°10'N, 92°00'E; Sylhet Forest, SE, 24°35'N, 92°10'E; Chittagong Hill Tracts Forest Division, N, 23°35'N, 92°15'E(Khan, 1981, pp. 13, 14). CHINA: Xizang, Nyalam vicinity, 28°12'N, 85°58'E (izas; Feng Zuojian, 1983, personal com- munication). Yunnan, Santun/Shuanglawa/Qi- qing, 27°45'N, 98°35'E (kiz); Gongshan vicinity, 27°4 1 'N, 98°37'E (izas); Bijiang Xian, NE, 26°40'N, 99°05'E (kiz); Bilushui Shan, 26°35'N, 99°05'E (kiz); Yao Jia Ping, 26°00'N, 98°50'E (kiz); Xima Nabang, 24°45'N, 97°45'E (kiz); Yingjiang, 24°45'N, 98°00'E (izas); Xiaoheshan, 24°30'N, 98°35'E (kiz); Zhung Shan 24°30'N, 98°40'E (izas); Jingdong Xian, 24°25'N, 100°50'E(kiz); Linchang Xian, 23°50'N, 100°05'E (kiz); Dahongshan, 23WN, 102°20'E (kiz); Dawei Shan, 22°55'N, 103°40'E(kiz); Menglun, 21°55'N, 101°15'E(kiz); Menghan, 21°50'N, 100°55'E (kiz); Xiangming, 21°45'N, 101°25'E (kiz); Manpa (izAs)/Mengla Xian (kiz), 21°30'N, 101°35'E. Guangxi, Tian'e, 25°00'N, 107°10'E; Huanjiang, 24°50'N, 108°15'E; Napo, 23°23'N, 105°48'E; Debao, 23°20'N, 106°37'E; Jiangxi, 23°08'N, 106°25'E (Wu, 1983, p. 16). VIETNAM: Luc-yen, 22°05'N, 104°43'E (Dao, 1 967, p. 1 1 7); Chieng ve, 20°46'N, 1 04°34'E (Dao, 1978, p. 382). 22 FIELDIANA: ZOOLOGY Other Fieldiana: Zoology Titles Available in This Series Taxonomy and Evolution of the Sinica Group of Macaques: 2. Species and Subspecies Accounts of the Indian Bonnet Macaque, Macaca radiata. By Jack Fooden. 1981. 52 pages, 9 illus., 15 tables. Publication 1325, $5.50 Taxonomy and Evolution of the Sinica Group of Macaques: 3. Species and Subspecies Accounts of Macaca assamensis. By Jack Fooden. 1982. 52 pages, 10 illus., 11 tables. Publication 1329, $5.25 Taxonomy and Evolution of the Sinica Group of Macaques: 4. Species Account of Macaca thibetana. By Jack Fooden. 1983. 20 pages, 5 illus., 3 tables. Publication 1345, $3.50 Order by publication number and/or ask for a free copy of our price list. Address all requests to: FIELD MUSEUM OF NATURAL HISTORY Division of Publications Roosevelt Road at Lake Shore Drive Chicago, Illinois 60605-2496, U.S.A. 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