2 behavior of the bears changed with three phases of the cycle: the proestrus, the estrus, and the metestrus. Each phase lasted about a week. During t

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1 7(1) 111, Breeding Behavior of the Formosan Black Bear (Ursus thibetanus formosanus) in Captivity 1 ChiehChung Yang, WenHui Chan and FuHsiung Hsu 1 Endemic Species Research Institute, Chichi, Nantou, Taiwan (Ursus thibetanus formosanus) (estrous cycle) (proestrus) (estrus) (metestrus) 47 (p<0.001) min (estradiol) (progesterone) Abstract A pair of the Formosan black bears (Ursus thibetanus formosanus) in captivity was housed together during the breeding seasons of , and their breeding behavior associated with estrous cycle was continuously recorded with a video recorder. The estrous cycle lasted three weeks for each season, and 1 300

6 min (estradiol) (progesterone) Abstract A pair of the Formosan black bears (Ursus thibetanus formosanus) in captivity was housed together during the breeding seasons

2 2 behavior of the bears changed with three phases of the cycle: the proestrus, the estrus, and the metestrus. Each phase lasted about a week. During the proestrus the male actively pursued (interacted with) the female but was ignored. During the estrus both male and female interacted frequently and mating (mounting and copulation) occurred frequently for four to seven days. During the metestrus the female still followed the male, but the male showed less interest on the female and no mating behavior occurred. The percentages of time periods of resting, of activity and of stereotypical behavior among the three phases were not significantly different, but the interactive behavior differed significantly among the phases (KruskalWallis test h=16.557, p<0.001). The copulation usually took place in early morning and evening, and lasted from 0.2 to 26.6 min. The fecal estradiol and progesterone concentrations in the estrus were higher those in the proestrus and metestrus. Probably it was a main reason that higher progesterone and estradiol concentrations, and their synergistic effect brought about the female bear to admit for mounting. Key words: Ursus thibetanus formosanus, estrous cycle, breeding behavior Received: August 11, 2004 Accepted: November 9, 2004 (Hafez 1993) (estrous cycle) (proestrus) (estrus) (metestrus) (diestrus) (McDonald 1980) (steroid sex hormone) (estrogen) (follicular phase) (progesterone ) (luteal phase) (McDonald 1980; Flowerdew 1987) (Ursus thibetanus formosanus) (Yang et al. 2003) (pseudopregnancy) (delayed implantation) 5 (Ursus americanus) (Wimsatt 1963; Flowerdew 1987; Hunt and León 1995) (Ursus thibetanus ) (Domico and Newman 1988)

During the estrus both male and female interacted frequently and mating (mounting and copulation) occurred frequently for four to seven days.

3 7(1) 111, (Helarctos malayanus) ( 1991) (mount) 35 ( 1998) ( 1991) (1991) kg kg 105 kg kg CCD 24 (rest) (action) (stereotype) (interaction) (mating) ( ) ( ) ( ) ( ) ( ) ( ) StatView KruskalWallis MannWhitney (noninvasion) g 3 ml 10 min min 20 min 3,500 rpm 30 min (1994)

5 79 3 2004 2 79 5 2 CCD 24 (rest) (action) (stereotype) (interaction) (mating) ( ) ( ) ( ) ( ) ( ) 20022004

4 4 (enzyme immunoassay, EIA) (estradiol) (Sigma P9776) anthos 2001 (Austria ) Boone et al. (2003) (courtship) (refractory period) Herrero and Hamer (1977) (Ursus arctos) Goodwin et al. (1979) Methyl phydrobezoate (oestrogen) (Curtis et al. 1971; Michael and Keverne 1970) (1991) (1991) (U. arctos yesoensis) (Ishikawa et al. 2003) (Diskin and Sreenan 2000) 510 sec ( 1998; Tsubota and Kanagawa 1991; Ishikawa et al. 2003) (1992) Boone et al. (2003)

(1979) Methyl phydrobezoate (oestrogen) (Curtis et al. 1971; Michael and Keverne 1970) (1991) (1991) (U.

5 7(1) 111, (pelvic thrust) (flutter) Mundy and Flook (1964) ( 1991) (U. a. horribilis) (Herrero and Hamer 1977) 1436 (Ishikawa et al. 2003) 20 ( 1992) (Craighead et al. 1969) (1998) % % (Cortney et al. 2002; Cortney and Mark 2003) (p<0.0001) (p=0.0005) (p=0.1467) (1991) (1998) 35

horribilis) 27 16 (Herrero and Hamer 1977) 1436 (Ishikawa et al. 2003) 20 ( 1992) (Craighead et al.

6 (A) (B) (C) Fig. 1. The % profiles of breeding behaviors in a day for the female Formosan black bear in the proestrus (A), the estrus (B), and the metestrus (C) for the years of 2002 to 2004.

7 7(1) 111, Table 1. Percentages (mean S.E.) of different behaviors of female Formosan black bears in captivity during the breeding seasons of Item Rest Action Stereotype Interaction Mating Proestrus 68.3± ± ± ±0.841 a n=288 Estrus 68.8± ± ± ±0.734 ab 1.9±0.466 n=360 Metestrus 70.4± ± ± ±0.611 c n=264 HValue PValue a,b,c Significantly different at 1% level (KruskalWallis test h=16.557, p=0.0003) between the figures with the different superscripts a, b and c of the same column ( ) Table 2. Mating (mounting) frequency and time periods (mean; ranges in parentheses) of the Formosan black bears in captivity during the estrus cycle in Date Frequency Time period (min.) May 15Apr 17Mar ( ) 1.0(0.51.9) 1.1(0.49.0) 6May 16Apr 18Mar ( ) 5.9( ) 17.6(17.6) 8May 17Apr 20Mar ( ) 2.6( ) 2.5( ) 9May 18Apr 21Mar ( ) 8.3( ) 4.6( ) 23Mar 6 7.4( ) Total or average ( ) 4.4( ) 4.1( ) (1985) 0.1 min 41.2 min 3.8 min 51.6 sec (Boone et al. 2003) 20 min (Craighead et al. 1969; Sparrowe 1964) 530 min ( 1998)

1±0.841 a n=288 Estrus 68.8±1.965 10.5±0.953 14.8±1.588 4.0±0.734 ab 1.9±0.466 n=360 Metestrus 70.4±2.307 11.3±1.070 15.7±1.687 2.6±0.611 c n=264 HValue 0.569 3.839 1.738 16.557 PValue 0.7524 0.

8 Fig. 2. The distribution of the total hourly mating frequencies during a day for the Formosan black bear in breeding seasons from 2002 to Fig. 3. The distribution of total hourly mating period during a day for the Formosan black bear in breeding seasons from 2002 to (pheromone) (sexual receptivity) (McDonald 1980) 2003

9 7(1) 111, Fig. 4. The variance of the fecal estradiol and progesterone concentrations for the female Formosan black bear in captivity at proestrus, estrus, and metestrus of the estrous cycle in 2002, 2003 and 2004.

10 (psychic estrus) ((McDonald 1980) (blastocyst) (Hamlett 1935; Renfree and Calaby 1981) (2) : (4): Boone, W. R., M. E. Richardson and J. A. Greer Breeding behavior of the American black bear Ursus americanus. Theriogenology 60: Cortney, A. T., C. Y. Mark and H. L. Mark Envirnonmental enrichment: Effects on stereotyped behavior and reginal neuronal metabolic activity. Brain Research 938: Cortney, A. T. and H. L. Mark

Breeding behavior of the American black bear Ursus americanus. Theriogenology 60: 289297. Cortney, A. T., C. Y. Mark and H. L. Mark. 2002.

11 7(1) 111, Environment enrichment: Effects stereotyped behavior and neurotrophin levels. Physiology and Behavior 80: Craighead, J. J., M. G. Hornocker and F. C. Craighead Reproductive biology of young female grizzly bears. Jorunal of Reproduction and Fertility 6: Curtis, R. F., J. A. Bullentine, E. B. Keverne, R. W. Bonsal and R. P. Michael Identification of primate sexual pheromones and the properties of synthetic attractants. Nature 232: Diskin, M. G. and J. M. Sreenan Expression and detection of oestrus in cattle. Reproduction, Nutrition, Development 6: Domico, T. and M. Newman Bears of the world. Facts on File, New York. Flowerdew, J. R Mammals: Their reproductive biology and population ecology. Edward Arnold, Great Britain. Goodwin, M., K. M. Gooding and F. Regnier Sex pheromone in the dog. Science 203: Hafez, E. S. E Reproduction in farm animals: Reproduction behavior. Lea and Febiger, Philadelphia. Hamlett, G. W. D Delayed implantation and discontinuous development in the mammals. The Quarterly Review of Biology 10: Herrero, S. and D. Hamer Courtship and copulation of a pair of grizzly bears, with comments on reproductive plasticity and strategy. Journal of Mammalogy 58: Hunt, J. P. and V. León Bears. Silver Burdett Press, USA. Ishikawa, A., H. Sakamoto, S. Katagiri and Y. Takahashi Changes in sexual behavior and fecal steroid hormone during the breeding season in female Hokkaido Brown bears (Ursus arctos yesoensis) under captive condition. The Journal of Veterinary Medical Science 65(1): McDonald, L. E Veterinary endocrinology and reproduction. Lea and Febiger, Philadelphia. Michael, R. P. and E. B. Keverne Primate sex pheromones of vaginal origin. Nature 225: Mundy, K. R. D. and D. R. Flook Notes on the mating activity of grizzly and black bears. Journal of Mammalogy 45: Renfree, M. B. and J. H. Calaby Background to delayed implantation and embryonic diapause. Journal of Reproduction Fertility. Supplement 29: 19. Sparrowe, R. D Sexual behavior of grizzly bears. The American Midland Naturalist 80: Tsubota, T. S. and H. Kanagawa Wimsatt, W. A Delayed implantation in the Ursidae, with particular reference to the black bear (Ursus americanus Pallas). pp In Delayed Implantation. A. C. Enders (ed.). University of Chicago, Chicago. Yang, C. C., F. C. Mao, F. T. Chan and T. C. Ho Reproduction behavior and characters of the Formosan black bear in captivity. Endemic Species Research 5(1): 113.

Identification of primate sexual pheromones and the properties of synthetic attractants. Nature 232: 396398. Diskin, M. G. and J. M. Sreenan. 2000. Expression and detection of oestrus in cattle.

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1111 The Planning and Establishment of Internet on Asiatic Black Bears Conservation in Yushan National Park 理年 I II IV 論 1 行 1 1 2 2 類 7 13 16 六 19 料 22 28 連 30 流 31 論 31 參 31 錄 33 錄錄 34 錄見 36 錄見 36