48 (6) : , ( Grooming) 1991 ; Chiarello, 1995) (Allogrooming), (Autogrooming) ( Perez et al. )

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1 48 (6) :707715, 2002 A cta Zoologica S inica 3 33 (, , ) (, , ) (, , ), 113 ( 351 ), 293 (bout), ( Rhinopithecus roxellana), ( Hygienic functional hypothesis) 17,,, :,,,,, ( Grooming) 1991 ; Chiarello, 1995),, (Allogrooming), (Autogrooming) ( Perez et al. ), (, ) ;, ( Goodall, 1968 ; Ghiglieri, 1984) ( Struhsaker, : (1) ( Hygienic hy2 pothesis) : ( Hladik, 1975), ( Hladik, 1975 ; Smit h, 1977 ; Ghiglieri, 1984 ; Perez et al., 1999) ; (2) (Distensive hypothesis) :,, ( Terry, 1970 ; Goosen, 1974 ; Boccia, 1983, 1986 ; Schino et al., 1990 ; De Waal et al., 1989) ; (3) (Affiliative hypothesis) :, ( M acaca ( Kummer, nemest rina) 1982 ; Smuts, 1983, 1985 ; Idani, 1991 ; Dunbar, 1967 ; Smith, 1977) Buettner2Janusch et al. (1962), ( Prosimian) Boccia (1989) ( M acaca radiata) Borries (1992) ( Presbytis entel , (No ) ( ) (2001) ( ) ( ) 33 E2mail : edu. cn,, 40, :

2 l us),, 351, 293, Nikon,, (2040 ) Nikon (10 ), Reichard et al. bates lar) (1994) ( Hylo2 ( Focal2animal sam2, pling),,, (, 2000),, 30 s,, 30 s (, 1 m, 2002), , (bout),,, 1 min, Altmann (1974) ( E, 33, N),, Nekaris (2001), ( ( Focal2animal instantaneous point2sampling) ) ( ) 113 (2 600 ) 2, 4, 90, :,, 110,, (,, ),, ; (1) :,, , (Long et al., 1998) : 1), ; 2),,,, ; :,, (,, (1) 113 ),, (, 10 4 ), ; :, 1 Table 1 A tally of observation period and troop composition Observation period Tracking days Troop composition (March 17 May 3, 2000) ( February 19 March 20, 2001) (September 3 December 8, 2001) 37 d (37 days) 22 d (22 days) 54 d (54 days) 20, 27, 17, 27, 91 (20 adult males, 27 adult females, 17 juveniles, 27 infants, 91 individuals in total) 4, 9, 2, 3, 18 ( 4 adult males, 9 adult females, 2 juveniles, 3 infants, 18 individuals in total) 8, 36, 7, 7, 58 ( 8 adult males, 36 adult females, 7 juveniles, 7 infants, 58 individuals in total),,,,

3 6 : :, ( ) Table 2 Percentages of body surface occupied by each site ( ) and percentages of allogrooming time received by Sichuan snub2nosed monkeys :, : : : : : : : 1314 : :, (Anogenital area) :, :, :, :,, :, : (Distal tail), Perez et al. (1999), 12, ( n = 3) cm 2 ( n = 3) (Leg) cm 2 ( n = 3) ( Foot) cm 2 ( n = 3 ) cm 2, :, : ( ), :, :,,, Body site :, :, :,,,,, Surface (Mean SD) Allogrooming time (Mean SD) ( Face) ( Head) (Neck) (Upper back) (Lower back) ( Inaccessible by autogrooming) ( Tail base) (Chest) (Arm) (Belly) ( Flank) (Difficult to reach by autogrooming) ( Hand) ( Forearm) ( Thigh) ( Easy to reach by autogrooming) Total ( ) ( ) ( df = 2, 2 = 12118, P < 01001) [ The 2 ( df = 2) comparing observed ( allogrooming) and expected ( surface) percentages represented by each accessibility category was 12118, P < 01001) ],,

4 Ghiglieri (1984) Boccia (1983),, ( ) A1 :,, %,, ; B : 3, %,, ; C1, :,, %, ( 1), 117,, 1 %, 1 % ( Perez et, al., 1999), (Sign test) %, +,, -, 1 % ( 011 %(019 %111 %), ), N 1 ( ) Fig. 1 Percentages of autogrooming corresponding to grooming times on various body sites ( from diff icult to easy) in Sichuan snub2nosed monkeys ARM :,, [ Area extending from the shoulder to the elbow (both included) ; the armpit is also included in this cat2 egory] CHE :, (Chest, vertral surface of the thorax, between the neck and the belly) FLA :, ( Flank, lateral region extending from the chest and belly to the back) ANO :, (Anogenital area, area embracing the buttocks, ischial callosities, sexual skin, and anus) TA2B :, ( Tail base, tail region extending from the lower back to the middle of the tail) BE :, (Belly, ventral surface of the abdomen, between the chest and the anogenital area) FOOT :,, [ Terminal part of both hind limbs, from the ankle to the tip of the toes (both included) ] hind limbs extending from the knee (included) to the ankle (not included) ] L EG:,,, [ Area of both D2TA :, (Distal tail, tail region ex2 tending from the tail base to the terminal end of the tail) FOR :,, [ Forearm, area extending from the wrist to the elbow (excluding the latter) ] ing from the hips (included) to the knees (not included) ] extending from the wrist (not included) to the finger tips] THI :,, [ Thigh, proximal segments of both hind limbs, extend2 HAND :,, [ Hand, terminal part of both front limbs

5 6 : ,,,,, ( Sign2test) ( n, = 17, P < 0105) (3), (0127 %) (0151 %) (0162 %) 2,, 1314 % % 213, 1 % 3157 % ;,, % %, 1 % 1118 % ; < 0105), 12/ 16 ( P > 0105), %, 14/ 15 ( P < 0101), 16/ %, 1 % ( P < 0101) (4), 0128 %, ( df = 2, 2 = , P < 01001) 2,,,,,, ; ( Furura, 1957 ;, Sparks, 1967 ; Goodall, 1968 ; Barton, 1983 ; (1 %) (0185 %) ; ;,, 14/ 17 ( P 3 Ghiglieri, 1984 ; Borries, 1992) Reichard et al. 2 Fig. 2 Percentages of autogrooming and allogrooming of different body sites relative to their accessibility in the Sichuan snub2nosed monkey

6 % 4 Table 3 Rates of allogrooming time per 1 % of body surface area and f it of the data for each site to the allogrooming prediction derived from the compensation hypothesis in Sichuan snub2nosed monkeys 1 % Body Body site Rates of allogrooming Fit of the data for each site time per 1 % of body sur2 site to the prediction face area (Mean SD) derived from the com2 pensation hypothesis ( Face) ( Head) (Neck) (Upper back) (Lower back) ( Tail base) (Anogenital area) (Chest) (Arm) (Belly) ( Flank) (Distal tail) ( Forearm) ( Hand) ( Thigh) (Leg) ( Food) / ( + - ) 13/ (Consitent with prediction) - ferences between percentage of allogrooming received and percentage of ( Contrary to prediction), ( Sign2test, n = 17, 3 P < 0105) Table 4 Fit of data from each body site to that predicted by the compensation hypothesis in different sex2age classes in Sichuan snub2nosed monkeys Adult Adult Infant Total J uvenile male female ( Face) ( Head) (Neck) + + N + (Upper back) (Lower back) ( Tail base) (Anogenital area) (Chest) - - N + (Arm) + N + + (Belly) ( Flank) (Distal tail) ( Forearm 33 ) N ( Hand) ( Thigh) (Leg) ( Food) / / 16 + / ( + - ) All sites + / ( + - ) 14/ / / (Sign2test) : 3 P < 0105 ( P < 0105 indi2 cates a result significantly different to the predicted value) N, 1 % 0101 % ( N indicates no dif2 body surface area) 2 (Symbol interpretation as in Table 2). (1994),,,,, ( Me Kenna, 1978 ; Barton, 1983 ; Boccia, 1983,, 1986, 1989) (2),, ( Terry,, 1970) (3),,, (, 2000),,, Boccia

7 6 : 713 (1989) ( M acaca radiata),,, ( M acaca nemest rina) ;,,,, Boccia (1989), Li et al. (1994) ; ( M acaca thibetana),,, ( Idani, 1991 ;, Dunbar, 1991 ; Chiarello, 1995), Borries ( 1998) Presbytis entell us,,, 1 % 1 %, 0185 % ; 1 %,, 3,,, 1 %,,, ;, ( Perez et al., 1999),,,,, ( 3) (4),, Kazuo Wada 12/ 16 ( P > 0105), 14/ 15 ( P < 0101), 16/ 16 ( P < 0101),,,,, 14/ 17 ( P < 0105), Fumio Fukuda ( References) Altmann, J Observational study of behavior : sampling method. Behavior 49 : Barton, R. A A comparative study of grooming interactions in primates. Pri mates 24 : Boccia, M. L A functional analysis of social grooming patterns through direct comparison with self2grooming in rhesus monkeys. Int. J. Pri matol. 6 : Boccia, M. L Grooming site preferences as a form of tactile communication and their role in the social relations of rhesus monkeys. In : Taub, D. M and F. A. King ed. Current Perspectives in Social Dynamics. New York : Van Nostrand Reinhold, Boccia, M. L Comparison of the physical characteristics of grooming in two species of macaques ( Macaca nemest ri na and Macaca radia2 ta). J. Com p. Psychol. 103 : Borries, C Grooming site preferences in female langurs ( Presbytis entell us). Int. J. Pri matol. 13 : Buettner2Janusch, J. and R. K. Andrew 1962 Use of incisors by primates in grooming. A m. J. Phys. A nthropol. 20 : Chiarello, A. G Grooming in brown howler monkeys, A louatta f usca. A m. J. Pri matol. 37 : De Waal, F. B. M. and L. M. Luttrell 1989 Towards a comparative socioecology of the genus Macaca : different dominance styles in rhesus and stumptail monkeys. A m. J. Pri matol. 19 : Dunbar, R. I. M Functional significance of social grooming in primates. Folia Pri matol. 57 : Furuya, Y Grooming behavior in wild Japanese monkeys. Pri mates 1 : Ghiglieri, M. P The chimpanzees of Kibale Forest : a field study of ecology and social structure. New York : Columbia University Press.

8 Goodall, J The behavior of free2living chimpanzees in the Combe Stream reserve. A nti. Behav. Monogr. 1 : Goosen, C Immediate effects of allogrooming in adult stumptailed macaques ( Macaca arctoi des). Behavior 48 : Idani, G Social relationships between immigrant and resident bonobo ( Pan paniscus) females at Wamba. Folia Pri matol. 57 : Li, B. G., B. P. Ren, Y. F. Gao, S. Y. Zhang and B. Liang 2000 Home range shift of Sichuan golden monkeys in summer caused by defor2 estation in the Qinling Mountains. Acta Zoologica Si na 46 (4) : (4) : ] [,,,, 2000 Li, J. H. and Q. S. Wang 1994 Allogrooming in wild Tibetan macaques ( Macaca thibetana) with special reference to social structure. In : Roed2 er, J. J., B. Thierry, J. R. Anderson and N. Herrenschmidt ed. Current Primatology. Vol. Social Development, Learning and Behavior. Strasbourg, France : University Louis Pasteur, Long, Y. C, R. C. Kirkpatrick, L. Xiao and T. Zhong 1998 Time budgets of the Yunnan snub2nosed monkey ( Rhi nopithecus [ Rhi nopithecus ] bieti). In : Jablonski, N. G., ed. The Natural History of the Doucs and Snub2noesd Monkeys. Singapore : World Scientific Publishing, Kummer, H Social knowledge in free2ranging primates. In : Griffen, D. R. ed. Animal Mind2Human Mind. Berlin : Springer2Verlag, Mckenna, J. J Biosocial functions of grooming behavior among common Indian langur monkeys ( Presbytis entell us). A m. J. Phys. A n2 thropol. 48 : Nekaris, K. A. I Activity budget and positional behavior of the Mysore slender loris ( L oris tardigradus lydekkerianus) : implications for slow climbing locomotion. Folia Pri matol. 72 : Perez, A. P. and J. J. Vea Baro 1999 Does allogrooming serve a hygienic function in Cercocebus torguat us l unulat us? A m. J. Pri matol. 49 : Reichard, U. and V. Sommer 1994 Grooming site preferences in white2handed gibbons ( Hylobates lar). Pri mates 35 : Ren, B. P., S. Z. Xia, Q. F. Li, S. Y. Zhang, B. Liang and J. H. Qiu 2002 Male coplulatory patlems in captive sichun snub2nosed monkeys ( Rhi nopithecus roxellana). Acta Zoologica Si nica 48 (5) : [,,,,, (5) : ] Schino, G., D. Maestripieri, S. D. Scucchi and P. G. Turillazzi 1990 Social tension in familiar and unfamiliar pairs of long2tailed macaques. Be2 havior 113 : Smith, C. C Feeding behavior and social organization in howling monkeys. In : Clutton2Brock, T. H. ed. Primate Ecology : studies of feeding and ranging behaviors in lemurs, monkeys and ape. London : Academic Press. Smutts, B. B Special relationship between adult male and female baboons : selective advantages. In : Hinde, R. A. ed. Primate Social Re2 lationships. Oxford : Blackwell, Smutts, B. B Sex and Friendship in Baboons. New York : Aldine Publishing Company. Sparks, J Allogrooming in primates : a review. In : Morris, D. ed. Primate Ethology. London : Weidenfieldland Nicholson, Struhsaker, T. T Social structure among vervet monkeys. Behavior 29 : Terry, R. L Primate grooming as a tension reduction mechanism. J. Psychol. 76 :

9 6 : 715 ( Abstract) DOES ALLOGROOMING SERVE A HY GIENIC FUNCTION IN THE SICHUAN SNUB2NOSED MONKEY ( R HINO PI TH ECUS RO XELL ANA)? 3 L I Bao2Guo 33 ZHAN G Peng Kunio WA TANAB E Chia L. TAN ( Depart ment of Biology, College of L if e Science, Northwest U niversity, Xiπan , Chi na) ( Pri mate Research Instit ute, Kyoto U niversity, Inuyama , Japan) ( Center f or Reproduction of Endangered S pecies, The Zoological Society of S an Diego, P. O. Box , San Diego, CA , USA ) To test the hygienic functional hypothesis of allogrooming in the Sichuan snub2nosed monkey, we analyzed the distribution of such behavior over the body surface of individuals in the West Ridge Troop in the Yuhuang2 miao region of t he Qinling Mountains, Shaanxi Province, China. Based on 113 days of observations of t he troop, 293 allogrooming bouts were recorded using focal2animal sampling. We divided the body surface of the monkeys into 17 areas. Before data analyses, 12 specimens were measured in order to calculate the size of each area. Each area was then ranked according to its level of accessibility to the groomer : easy to reach, difficult to reach, and inaccessible. We performed t hree successive analyses to determine : 1) t he overall dist ribution of al2 logrooming behavior on body surface areas ; 2) the overall distribution of allogroomed areas based on the three categories of accessibility ; and 3) age and sex class differences in allogrooming f requencies based on bot h t he de2 fined body surface areas and t he accessibility categories. Our preliminary result s show t hat allogrooming behavior concent rates primarily on t he dorsal and anogenital regions. Accordingly, t he post ures of grooming dyads pre2 cludes them from facing each other, a potential social strategy adopted by these monkeys to reduce tension and decrease conflict. Allogrooming also occurred most f requently on body surface areas in t he inaccessible category. We found that the frequency of allogrooming of difficult to reach areas was proportional to the size these areas covered on t he monkeyπs body. Furt hermore, areas t hat were easy to reach were less f requently allogroomed t han expected. Finally, t he findings on age and sex class differences showed t hat allogrooming were influenced by social ranking. The dist ribution of allogrooming behavior on body surface areas st rongly suggest s t hat cleaning is an important motivation for such activity ; therefore, the results support the hygienic functional hypothesis. Key words Sichuan snub2nosed monkey ( Rhinopithecus roxellana), Allogrooming, Autogrooming, Hygienic function 3 This work was supported by National Natural Science Foundation of China ( No ), The Zoological Society of San Diego of USA ( ), Shaanxi Education Department Foundation of China (2001), Nippon Life Insurance Fund of Japan ( ) and Daiko Foundation of Japan ( ) 33 Corresponding author. edu. cn