On the geographical distribution of primates in China

Yong-Zu Zhang Institute of Geography, Academia Sinica, Peking, China

On the Geographical Distribution of Primates in China

This paper deals with the geographical distribution of primates in relation to environmental conditions and threatening factors in different zoogeographical regions. Sung Wang South China region: most species of primates inhabit the tropical forest stretching along the southern border of the country. Habitat destruction have caused Nycticebus coucang, Pygathrix nemaeus, Presbytis spp. and Hylobates spp. to be threatened with extinction. Through Institute of Zoology, intensive exploitation Macaca spp. have been reduced to some Academia Siniea, Peking, China secondary forest areas. South-west China region: Presbytis entellus inhabits the area below Received 4 July 1980 and the moist temperature belt in the southern flank of the Himalayas. accepted 4 December 1980 Tupaia glis live at 2000 m elevation in evergreen broadleaf forests. Some of the Macaca spp. population are forced to live at even higher Keywords : private species, China, elevations, over 3000 m, due to deforestation. Rhinopithecus roxellanae bieti still remains in Yunnan. conservation, natural reserves. Central China region: Macaca mulatta and M. speeiosa have disappeared in most parts of this region except for some isolated groups occurring in the rugged mountains. R . r . roxellanae and R. r. brelichi exist under protection, the latter being one of the most endangered mammals in China. North China region: there are only two small relict populations of M . mulatta in Shansi and Hopei reaching as far as lat. 41~ It once had a wide distribution in northern China as indicated in the historical literature.

Guo-Qiang Quan

1. I n t r o d u c t i o n So f a r as is k n o w n , 15 species o f p r i m a t e s a n d t u p a i i d s a r e t b u n d i n C h i n a , b e l o n g i n g to f o u r f a m i l i e s a n d s e v e n g e n e r a a n d o c c u p y i n g 14 ~ o f t h e t o t a l n u m b e r o f species o f t h e s e f o u r t'amilies i n t h e w o r l d ( ' F a b l e 1: A l l e n , 1 9 3 8 ; E l l e r m a n & M o r r i s o n - S c o t t , 1 9 6 6 ; N a p i e r & N a p i e r , 1967). Table 1

P r i m a t e species in China

Number of species f

Family

Species

Lorisidae Cercopithecidae

Nycticebus coucang Macaca mulatta 21~r. assamensis M . cyclopis M . speciosa* M . nemestrina Rhinopithecus roxellanae Pygathrix nemaeus Presbytis entellus P. phayrei P. francoisi Hylobates concolor H. hoolock H. lar

Hylobatidae

Total

World

China

~,

11

1

9" 1

72

10

13.9

7

3

9

90

I4

15-5

* It has recently been treated as two separate species: M . arctoides I. Geoffroy, 1831 and A4. thibetana Milne-Edwards, 1870 (see Fooden, 1976, 1980). Journal of Human Evolution (1981) I0, 215-226 0047-2484/8 l/030215 -I 14 $02.00/0

9 1981 Academic Press Inc. (London) Limited

2 t6

Y.z. ZHANGE T

AL.

Most of the primate species (Figure 1) are concentrated in the zoogeographical subregion of South Yunnan (Zhang & Zhao, 1978). As regards to the other subregions of the c o u n t r y , p r i m a t e species a r e less a b u n d a n t . T h e r h e s u s m o n k e y , M a c a c a niulatta, is t h e o n l y species w h i c h r a n g e s as f a r as n o r t h e r n C h i n a , i.e. t h e L o e s s P l a t e a u s u b r e g i o n ( F i g u r e

1). L i m i t i n g f a c t o r s to t h e n o r t h w a r d d i s p e r s a l o f p r i m a t e s i n C h i n a a r e d u e p a r t l y to t h e t e m p e r a t u r e ( F i g u r e 2), a n d also to t h e f o o d s u p p l y i n r e l a t i o n to t h e v e g e t a t i o n t y p e s a n d Table 2

P r i m a t e species in different zoogeographical regions

Zoogeographical region

Zoogeographical subregion

South China

Southern Yunnan

Min-Guang

Hainan

Taiwan

Central China

Hilly-Plain

Mountains

South-western China

South-west

Himalaya

North China

Loess plateau

Natural conditions

Primates species inhabited

Macaca mulatla M . assamensis M . speciosa M. nemestrina Presbytis phyrei Hylobates hoolock H. concolor H. lar Tupaia glis Nycticebus coucang Tropical monsoon forest. Tupaia glis Temperatures of 10~ Macaca mulatta for more than 300 days M . assamensis M. speciosa Presbytis francoisi Tropical rain tbrest and Tupaia glis monsoon forest Macaca mulatta Temperatures of 10~ for Pygathrix nemaeus more than 300 days Hylobates concolor Tropical rain forest and monsoon forest Macaca cyclopis Temperatures of 10~ for more than 300 days

Species number

Tropical rain forest. Temperatures of 10~ for more than 300 days

Evergreen subtropical forest with temperatures of 10~ for between 225 and 280 days Evergreen subtropical forest with temperatures of 10~ for between 225 and 280 days

Macaca mulatta M. speciosa

Subtropical mountain forest with temperate conifers. Summer less than 90 days

Tupaia glis Macaca mulatta M. speciosa M. assamensis Rhinopithecus roxellanae Tupaia glis macaca mulalta M . assamensis Presbytis entellus

Subtropical mountain forest with temperate conifers. Summer less than 90 days Deciduous broad-leaf forest with temperatures of 10~ for less than 175 days. Summer less than 90 days; 4200 daylight hours annually

Macaca mulatla M. speciosa Rhinopithecus roxellanae

Macaca mulatta

10

2-5

4 1

2

3

5

4

1

DISTRIBUTION OF PRIMATESIN CHINA

217

growing season. I t is also necessary to consider the amount of daylight during the winter as a limiting factor. In fact, daylight lasts for only nine hours at the winter solstice at Xinglung, which is the northernmost locality inhabited by the rhesus monkey. The total amount of daylight throughout the year in this region is 4200 h. Thus, the abundance of the primate fauna in different zoogeographical regions or subregions depends upon a combination of these factors (Table 2). In addition, h u m a n activities in such an ancient country must also be considered as an important factor influencing primate distribution. Two species of primates, RhinoDithecus roxellanae (Plate 1) and Macaca cyclopis, are confined to China. The former is distributed discontinuously in Sichuan, Kansu, Hupei, Shensi, Yunnan and Guizhou (Figure 3) and recognized as three subspecies. A tbssil (Rhinopithecus avunculus) has been found in Pleistocene deposits in South China of subtropical area. iUIacaca cyclopis is an insular species found only in Taiwan. O n account of the present status of primates in China, it is necessary to deal with the geographical distribution together with the problem of primate conservation from a zoogeographical point of view. Figure 1. The number of primate species in different zoogeographical subregions. [Zoogeographical division from Zhang & Zhao (1978).] Palaearctic realm. 1, North-east China region: Ia, Mt. Xingan subregion; Ib, Mt. Changbai subregion; Ic, Song-liao subregion; II, North China region : IIa, Huang-huai subregion; IIb, Loess plateau subregion; III, Neimeng-Sinjiang ~'egion; IIIa, East steppe subregion; IIIb, West desert subregion; IIIc, Mt. Tianshan subregion; IV: Qinghai-Xizang region; IVa, Qiang-Tang subregion; IVb, Qinghai-S. Xizang subregion. Oriental realm: V, South-west China region: Va, Southwest mountain subregion; Vb, Himalaya subregion; VI, Central China region: Via, Eastern hilly-plain subregion; VIb, Western mountains subregion; VII, South China region: VIIa, Min-Guang subregion; VIIb, Southern Yunnan subregion; VIIc, Hainan subregion ; VIId, Taiwan subregion; VIIe, Southsea Islands subregion.

%

218

Y.Z.

ZHANG ETAL.

Figurc 2. The northern limits of some climatic factors in China, ( ~ B i n ) Mean day temperature ~>10~ for more than 200 days; (. . . . . ) mean day temperature :>10~ for 225-280 days; (. . . . . . ) mean day temperature ~10~ for more than 300 days; ( ) more than a 90 day period with temperatures ~22~ (....... ) areas dominated by tropical rain forest.

2. South China Region

South Yunnan Subregion T h e northern margin of the tropical rain forest extends from I n d o c h i n a northwards to this region. It has often been regarded as the most convenient b o u n d a r y for the faunal characteristics of tropical and subtropical. T h e Old W o r l d tropical primates, such as Nycticebus eoucang, Hylobates hoolock, H. concolor, H. lar, Presbytis phayrei, reach northern limits which roughly coincide with that of the rain forest (Figures 4-6). T h e temperatures here, especially in the higher mountains, are unlike the southern parts of the rain forest. T h e annual temperature is 16-18~ with occasional cold weather below 10~ during J a n u a r y . However, favourable conditions of vegetation and sufficient food supply are available tbr tropical primates t h r o u g h o u t the year. Relatively, this subregion is provided with the best natural environment for primates in China. Besides the species mentioned above, four species of 2klacaca and the tupaiid Tupaia glis are also found here (Figures 6-7). Most of them are represented by distinct geographical subspecies within this subregion, including T o n k i n and adjacent areas: Tupaia glis modesta (Southern Y u n n a n , Hainan), T. g. chinensis (Western Y u n n a n ) , Nycticebus coucang bengaIensis (Western Y u n n a n ) , Maeaea nemestrina leonina, Macaca mulatta mulatta (common in China proper), M. speciosa melanotus, M. assamensis assamensis, Presbytis phayrei crepusulus, Hylobates concolor concolor (South-western Y u n n a n ) and H. concolor leueogenys (Southern Y u n n a n ) .

Plate 1. R/linopilhecus roxellanae brdichi Thornas (~, in Beijing Zoo), photo by G.-Q. Quan.

218

Plate 2. 'r

Natural reserve, Sichuan, photo by S. "Wang.

219

DISTRIBUTION OF PRIMATES IN CHINA

)

Figure 3. Geographical distribution ofRhinopithecus roxellanae ( and R. avunculus ( . . . . . . . . . . . ).

,

0

250

500

0

750

Figure 4.

Geographical distribution of Hylobates concolor ( and H . lar ( ...... ) in China.

H . hoolock ( m ~ ~ )

250 5 0 0

/

....

),

220

Y . z . ZHANG ET AL. Figure 5. Geographical distribution of Presbytis entellus (-----), P. francoisi ( .... ), P. phayrei ( ) and Pygathrix nemaeus ( ....... ) in China.

7

0 250 L__J

500 750 J --. J km

Figure 6. Geographical distribution of Tupaiaglis ( bus coucang (-- -- --) in China.

) and Nyctice-

f~

#

o

~to ~oo ~tom

221

D I S T R I B U T I O N OF P R I M A T E S IN C H I N A

Figure 7. Geographical distribution of Macaca mulatta ( m), M. speciosa (. . . . ), M. assamensis ( ~ ~ m), M. nemestrina (. . . . . . ) and M. cyclopsis ( ...... ) in China.

,..#

i f - - f \

L~ ~

,J

f

0 I

250 J

500 ~

750 km

3/Iin-Guang Subregion Long periods of intensive farming, hunting and deforesting have led to extinction of some species of land vertebrates in most parts of this subregion. However, it is difficult to know the changes in primate fauna from the old Chinese literature, in which the species identifications are not clear. At present, three species of primates distributed mainly in Indochina, are found in the extreme southern border of the country where the natural forests are well preserved. A distinct subspecies of Presbytis francoisi leucocephalus (Tan, 1957) is found in a local area near Nanlin. Macaea mulatta mulatta and M . speciosa speciosa, which are rather common in most parts of eastern and southern China, occur in the entire subregion. Hainan Subregion The isolated insular environment may account for the sparse mammalian fauna of Hainan. In spite of the favourable tropical climate, only four species of primates are found on the island. Tupaia glis and Hylobates concolor, similar to T. g. modesta and H. e. concolor respectively of the South Yunnan subregion, are regarded as the same subspecies. Macaca mulatta brevicaudus is considered as a valid subspecies of the island. Pygathrix nemaeus of Indochina had been recorded from Hainan but has not been reported in recent years. Taiwan Subregion The indigenous monkey, A/iacaca cyclopis, is the only representative of primates found in Taiwan. It occurs in at least five or six places in the Yu-Shan Mountains (Figure 7).

222

v.z.

ZHANG E T A L.

3. Central China R e g i o n The history of human civilization in this region of the Yangtze river basin has been continuous longer than in the South China region. Two species of monkeys, i.e. Macaca mulatta and M . speciosa thibetana, inhabit this region. Their terrestrial and omnivorous habits enable them to live in more or less disturbed forest and around farmer's habitations. The evergreen vegetation with its growing season more than 225-280 days provides them with an abundant food supply.

Hilly-Plain Subregiolz More intensive farming has reduced considerably the areas of natural forest which now are restricted to the hills with poor soil or rocky, steep slopes, usually at the borderland between provinces or distant f)om areas of dense settlements. The populations of M. m. mulatta and M . s. thibetana in this subregion are distributed discontinuously with wide gaps. In the areas along tile middle and lower reaches of the river Yangtze and its tributaries, river Han, Gan and Yuan, the two species became extinct long ago.

Mountain Subregion Topographically, this subregion is dominated by a big basin, the Sichuan Basin, and the surrounding mountains. The former has been farmed intensively and contains a dense human population. The mountains provide more or less suitable habitats for primates, because the farms and the wildland are interspersed with each other. Numerous collecting places recorded recently show a geographical distribution along the ridges of mountains. The ranges of Rhinopithecus roxdlanae roxellanae and R. r. brelichi have been restricted to quite small areas, and, compared with those of other primates in China, there has been no great change since they were first recorded about a century ago. R. r. roxellanae is distributed along the ranges of the Qionglai, Min-Shan and Qingling Mountains in this subregion. It was also recently discovered in the Shennonjia Mountains of Hupeh Province. It inhabits an altitude from 1400 to about 3000 m or more, and migrates vertically twice per year in different seasons. R. r. brelichi is found in the Fanjing Mountains in northeastern Guizhou. It is already very rare and threatened to become extinct.

4. South-west China R e g i o n South-west 2V[ountain Subregion The Meridiana Mountains run in a north south direction throughout the whole subregion. Topographically and climatically, this subregion has a complex environment with considerable vertical change of vegetational zones including the subtropical forest, dry savanna, temperate mixed forest, cool temperate conifers, and alpine shrub. The primates and typaiids have been observed as follows: (1) Tupaia glis. Northern limit about 29~ ranges from 860 to 2200 m altitude. (2) Macaca muIatta. Northern limit about 32~ ranges from 1700 to 3400 m altitude. (3) M . speciosa. Northern limit about 32~ ranges from 1700 to 3400 m altitude. (4) M . assamensis. Northern limit about 30~ ranges from 2500 to 3100 m, restricted to the southwestern border. (5) Rhinopithecus r. roxellanae. Northern limit about 34~ ranges from 2000 to 3000 m altitude.

DISTRIBUTION

223

OF PRIMATES IN CHINA

(6) R. r. bieti. Restricted in divided ranges of the river Mekong and Jinsa (DiChu), between 29 ~ to 30~ Their seasonal migration has been known to be not only due to the climatic change but also to the food supply in various types of forest. Himalaya Subregion

There are four species of primates and tupaiids discovered in the southern flank within Chinese territory. T. glis versurae has been recorded in Mishmi area of tropical forest. M . m. mulatta and M . a. assamensis are common in the eastern and middle Himalayas, and can be found frequently below 3400 m in subtropical and tropical forest. Presbytis entellus sehistaceus extends along the valley up to the highest limit of its range, and inhabits the tropical moist forest and the subtropical mixed forest in the Middle Himalayas (Jilon valley) below 2700 m altitude, whereas P. e. lania had been described by Elliot (1913) on the basis of one incomplete specimen from Chumbi Valley, but further information has been obtained in recent years (Oppenheimer, 1977). 5. N o r t h C h i n a R e g i o n Loess Plateau Subregion

H u m a n agricultural activity, including detbrestation, has lasted tbr about 5000 years and has led the primates of the Yellow river basin to become extinct in such a vast area. Fossils of A4acaca have been recorded in many places in this subregion from Pleistocene deposits (Pei, 1957; Zhou, 1963). Ancient Chinese literature indicates that some species of primates existed in the Loess Plateau during the seventeenth or eighteenth centuries. In recent years only the widely distributed M . m. mulatta still exists with two isolated populations, in the Zhongtiao Mountains, southern Shansi and in the Xinglon Mountains, northern Hopei. The latter reaches 41~ and is the northern limit of temperate deciduous forest, with about 90 days of summer and 180 days of winter. The monkey suffers more here than in other places of its range with regard to food supply. 6. C o n s e r v a t i o n o f C h i n e s e P r i m a t e s

The major problem on the conservation of primates in China is that the cultural, industrial and agricultural constructions have developed rapidly after liberation. The natural woodlands which most primates inhabit are disturbed or destroyed vastly. On the other hand, primates are subjected to over-trapping pressure. Statistics of purchase number of primate skins show the critical situation on this problem (Table 3). In addition, the common species of Maeaca living near to farms are reputed to raid crops and are hunted regularly. Table 3

Purchase number of primate skins (1974-76)

Year 1974 1975 1976

Purchase no. of primate and tupaiid skins* 8974 6355 10,885

* Tupaia glis, Presbytis entellus, P. nemaeus and Macaca cyclopis are not

counted.

224

Y . z . ZHANG1,:TAL.

Recently, the g o v e r n m e n t has paid more a t t e n t i o n to wildlife conservation, principally to the most famous rare species such as the g i a n t p a n d a a n d the gold monkey. P r i m a t e species are listed in the list of protected species of rare animals. T h e conservation for primates in different zoogeographical regions is discussed below. South China Region

I n southern Y u n n a n , for a long time, the s l a s h - b u r n cultivation a n d regular h u n t i n g carried out by the m o u n t a i n - l i v i n g natives has r e d u c e d the n a t u r a l habitats of primates as well as in some extent the p r i m a t e p o p u l a t i o n n u m b e r . I n the last decade, the noticeable influence has b e e n the r u b b e r p l a n t a t i o n which occupies a vast area a n d erodes the wilderness of S o u t h e r n Y u n n a n . E v e n d u r i n g the 1960s m a n y species such as Nycticebus coucang, Presbytis spp., Hylobates spp, a n d Macaca spp. could be found in the same patch of n a t u r a l forest not far from settlements, b u t these have b e e n t h r e a t e n e d to extinction except Macaca spp. a n d are now reduced to a few individuals in the rugged m o u n t a i n s . O v e r t r a p p i n g of the rhesus m o n k e y in South C h i n a in the 1950s for export caused m u c h reduction in its p o p u l a t i o n size. A l t h o u g h the exportation of primates has now basically ceased, a series of actions must be taken to conserve these v a l u a b l e a n i m a l resources efficiently. U p to now, seven n a t u r a l reserves or sanctuaries have been established particularly for p r i m a t e conservation (Table 4). Table 4

Natural reserves or sanctuaries mainly for primate conservation

Location

Vegetation

Primate specie

South China region

Mengluen, southern Yunnan

Tropical monsoon forest

6267

1958

tluaping, northern Guangxi Long-gang, southwestern Guangxi Jianfcngling, Hainan Nanwan, Hainan Bangxi, Hainan Bawangling, Hainan

Subtropical forest Tropical forest

Nycticebus coucang Presbytis phayrei Macaca spp. Macaca spp.

13,918

1961

Presbytisfrancoisi

7997

1979

Tropicalrain forest Hylobates concolor

1635

1960

Tropical rain forest Macaca mulatta Tropical rain forest iVIacacamulatta Tropicalrain forest IIylobates concolor

933 333 2000

1965 1976 1980

35,000

1978

95,292

1978

36,700

1978

20,000

1963

.

60,003 27,700 40,000

1978 1965 1978

. .

10,000 12,000

1979 1974

40,000 30,000 182,431

1975 1978 1963

Central China Fupin, southern region Shensi Baishuijiang, Shensi Fanjingshan, Guizhou

Subtropical forest

Rhinopithecus r. roxellanae Subtropical forest Rhinopithecus r. roxellanae Subtropical forest Rhinopithecus r. brelichi

South-western Baihe, Sichuan Subtropical forest region Jiuzhaigou, Sichuan . . . Wanglang, Sichuan ,, Tangjiahe, Sichuan . . . Xiaozhaizigou, Sichuan . . . Labahe, Sichuan . . . Fongtongzhai, Sichuan ,, Mabian, Sichuan ,, Baishuaijiang, Gansu . . .

Rhin@ithecus r. roxellanae

. ,,

,, ,, .

Area (ha)

Time established

Region

DISTRIBUTION OF PRIMATES IN CHINA

225

Central China Region Macaca mulatta and M. speciosa have disappeared in most parts of this region, especially in the Hilly-Plain subregion. The most endangered primate is Rhinopithecus roxellanae brelichi, which inhabits the higher mountains in Guizhou. Tremendous deforestation in recent decades within a small area has caused the reduction of forest habitat available for this valuable subspecies. It is believed that its population size has been reduced to no more than 200 individuals in several groups. On the other hand, R. r. roxellanae is in less danger of extinction because of its larger population size and wider distributional range. Three natural reserves (Plate 2) are settled particularly for conserving these two valuable subspecies in this region (Table 4). South-west China Region The destruction of forests in the steep mountains of this region has altered the habitat of primates. In general, the primates inhabiting this region face less trapping pressures than in other regions. Nine natural reserves are established, particularly for the giant panda, together with Rhinopitheeus r. roxellanae in this region.

North China Region In the 1940s, the population of Macaca mulatta occurring in Hopei still maintained a considerable number and a wide distribution. Subsequent deforesting and over-hunting caused a rapid decrease of its population size to about 200 individuals by the end of that decade. In recent years, the wild population of this monkey has remained at less than 10 individuals. They inhabit the rocky, shrubby areas at an altitude of about 1000 m in the Xinlon Mountains, about 100 km north-east of Peking. The present status of this monkey in the Zhontiao Mountains of Shansi is not clear. Generally speaking, great efforts should be made urgently to protect the vanishing primates in China. For this purpose, filrther detailed investigation of the present status of Chinese primates is essential. References

Allen, G. M. (1938). The Mammals of China and Mongolia, part 1, pp. 279-311. New York: American Museum of Natural History. Chang, C. & Wang, T. Y. (1963). Faunistic studies of mammals of the Chingbai province. Acta Zoologiea Sinica, 15, 125-138. Cheng, J. Z. (1969). VertebrateFauna of Taiwan. Taiwan Zanwo Jinshukun. Chivers, D . J . (1977). The lesser apes. In Primates Conservation, pp. 539-594. Chou, M. Z. (1963). The mammalian fossils and the climate of Pleistocene. Acta PalaeovertebrataSinica 7, 362-367. Ellerman, J. R. & Morrison-Scott, T. C. S. (1966). Checklist of Palaearctic and Indian Mammals. London: British Museum (Natural History). Elliot, 12). G. (1913). A Review of the Primates. 3 vols. New York: American Museum of NaturaI History. Fooden, J. (1976). Provisional classification and key to living species of macaques (Primates: Maeaca). Folia Primatologica, 25, 225-236. Fooden, J. (1980). Classification and distribution of living macaques (Macaca Lae6p6de, 1799). In The Macaques, pp. 1-9. Kao, Y. T., Lu, C. K., Chang, C. & Wang, S. (1962). Mammals of the Hsi-shuan-pan-na area in southern Yunnan. Acta Zoologica Siniea 14, 180-196. Lu, C. K., Wang, T. Y., Qyan, G. J., Gin, S. K., Ma, T. W. & Yang, T. I-I. (1965). On the mammals from the Lin-tsang area, west Yunnan. Acta Zootaxonomica Sinica 2~ 279-295. Napier, J. R. & Napier, P. H. (1967). A Handbook of Living Primates. London: Academic Press.

226

Y.z.

ZHANG ET AL.

Oppenheimer, J. R. (1977). Presbytis entellus. In Primates Conservation, pp. 469-512. Pei, W. Z. (1957). Distribution of the mammalian faunain Pleistocene of China. ActaPalaeovertebrataSinica 1, 9-24. .' Qian, Y. W. (1974). On ayes and mammals li~omJumulongma area. In Scientific Reports of Jumulongma area-Biology and Alpine Physiology, pp. 1-23. Tan, P. C. (1957). Rare catches by Chinese animal collectors. Zoo Life 12, 61-63. Wang, S., Lu, C. K., Kao, Y. T. & Loo, T. C. (1962). On the mammals from southwestern Kwangsi, China. Acta Zoologica Sinica 14, 555-568. Zhang, Y. Z. & Zhao, K. T. (1978). On the zoogeographical regions of China. Acta Zoologica Sinica 24~ 196-202.