48 (6) :707715, 2002 A cta Zoologica S inica 3 33 (, 710069, ) (, 484-8506, ) (, 120551, ), 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 entel2 2002202207, 2002207220 3 (No. 39970116) (200122003) (2001) (199922000) (200022002) 33 E2mail : baoguoli @nwu. edu. cn,, 40, :
08 7 48 l us),, 351, 293, 4 251 Nikon,, (2040 ) Nikon (10 ), 20300 Reichard et al. bates lar) (1994) ( Hylo2 ( Focal2animal sam2, pling),,, (, 2000),, 30 s,, 30 s (, 1 m, 2002), 2000 3 2001 12, (bout),,, 1 min, Altmann (1974) 1 111 (108 14 108 18E, 33, 45 33 50N),, Nekaris (2001), ( 1 400 2 200 ( Focal2animal instantaneous point2sampling) ) (2 200 2 600 ) 113 (2 600 ) 2, 4, 90, :,, 110,, (,, ),, ; (1) :,, 112 2000 3 2001 12, (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 2000 3 17 5 3 (March 17 May 3, 2000) 2001 2 19 3 20 ( February 19 March 20, 2001) 2001 9 3 12 8 (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),,,,
6 : 709 2 114 :, ( ) 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) 12 23713 cm 2 ( n = 3) (Leg) 9 54110 cm 2 ( n = 3) ( Foot) 6 92419 cm 2 ( n = 3 ) 1 18516 cm 2, 2 115 :, : ( ), :, :,,, Body site :, :, :,,,,, Surface (Mean SD) Allogrooming time (Mean SD) ( Face) 1194 0116 1164 0118 ( Head) 3186 0124 9137 0112 (Neck) (Upper back) (Lower back) ( Inaccessible by autogrooming) 1127 0102 4103 0111 3142 0106 14177 0126 2191 011 18101 0112 0124 47182 0151 ( Tail base) 4114 0106 6133 0107 (Chest) (Arm) (Belly) 1127 0102 10174 0113 4177 0114 1128 0106 10157 0106 6157 01002 5140 0166 9123 0109 ( Flank) 5112 0103 2168 0105 31127 0156 (Difficult to reach by autogrooming) ( Hand) 36183 0128 3109 0112 3117 0104 ( Forearm) 9168 0115 11328 0103 5130 0103 0129 0101 ( Thigh) 15121 0110 8111 0104 55133 1154 ( Easy to reach by autogrooming) 14115 0107 2119 0104 7190 0106 01311 0107 15135 0120 Total 100 100 2 ( ) ( ) ( 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) ],,
10 7 48 116 2 Ghiglieri (1984) Boccia (1983),, ( ) A1 :,, 13140 %,, ; B1 11712 : 3, 31127 %,, ; C1, :,, 55133 %, ( 1), 117,, 1 %, 1 % ( Perez et, al., 1999), (Sign test) 11711 1 %, +,, -, 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
6 : 711 11713 212,,,,, ( Sign2test) 17 13 ( n, = 17, P < 0105) (3), 2 211 (0127 %) (0151 %) (0162 %) 2,, 1314 % 47182 % 213, 1 % 3157 % ;,, 31127 % 36183 %, 1 % 1118 % ; < 0105), 12/ 16 ( P > 0105), 55133 %, 14/ 15 ( P < 0101), 16/ 16 15135 %, 1 % ( P < 0101) (4), 0128 %, ( df = 2, 2 = 121186, 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
12 7 48 3 1 % 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) 0185 0110 - ( Head) 2143 0116 + (Neck) (Upper back) (Lower back) 3117 0105 + 5178 0103 + 4147 0113 + ( Tail base) 1153 0104 + (Anogenital area) (Chest) (Arm) (Belly) 8146 0108 + 0127 0102-0162 0100-1171 0106 + ( Flank) 0151 0102 - (Distal tail) 0138 0104 + ( Forearm) 0134 0100 + ( Hand) 0105 0101 + ( Thigh) 0153 0101 + (Leg) ( Food) 0115 0101 + 0104 0101 + + / ( + - ) 13/ 17 3 + (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) + + + + 14/ 17 3 12/ 16 + / ( + - ) All sites + / ( + - ) 14/ 15 3 16/ 16 3 894/ 1 020 3 (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
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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 710069, Chi na) ( Pri mate Research Instit ute, Kyoto U niversity, Inuyama 48428506, Japan) ( Center f or Reproduction of Endangered S pecies, The Zoological Society of S an Diego, P. O. Box 120551, San Diego, CA 9211220551, 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. 39970116), The Zoological Society of San Diego of USA (20012003), Shaanxi Education Department Foundation of China (2001), Nippon Life Insurance Fund of Japan (19992000) and Daiko Foundation of Japan (20002002) 33 Corresponding author. baoguoli @nwu. edu. cn