夜行性的動物傳達訊息時，氣味或聲音通常較視覺更為重要。白面鼯鼠為一種台灣夜行性哺乳動物，會頻繁地使用叫聲，但是其聲音與行為的關係尚不明確。本研究目的是藉由錄音分析配合行為觀察，探討白面鼯鼠的叫聲類型、使用狀況及可能功能。研究樣區位在阿里山光華村的次生林，於2012年3月至2013年2月，每月連續兩夜，針對樹間活動的白面鼯鼠錄音並紀錄其行為。結果顯示，白面鼯鼠全年皆頻繁的發出叫聲，晚上7-9點與清晨3-5點是兩段高峰期。鳴叫聲依據頻譜特徵可以區分為A、B、C、D四種類型。將周遭環境以及伴隨行為皆納入考慮後，發現每種叫聲類型皆可能發生在不同情況下，但根據出現的頻率，大致上可將不同類型聲音區別出不同功能，分為:單音型A叫聲為警戒音，多用於警戒人類(非同種類入侵者)；連續型A叫聲為社交叫聲，用於相伴活動的個體彼此聯繫；叫聲C為競爭叫聲，多用於同種間的競爭，像是雄鼠間競爭交配機會時；叫聲B與D功能則因樣本數不足無法推斷。叫聲類型A及B在繁殖季出現的次數高於非繁殖季。類型A的叫聲在雌鼠與雄鼠間無顯著差異，但在部分個體間有顯著差異，推測可能因為白面鼯鼠為獨居性之物種，故聲音中所涵蓋資訊不如社會性高之物種一般詳細。

Olfactory or vocal communication is usually more important than visual communication in exchanging information for nocturnal animals. The white-faced flying squirrel (Petaurista lena) is a nocturnal animal and frequently use calls to communicate. However, the relationship between their calls and behavior is not clear. The goals of this study were to describe their calls produced by the white-faced flying squirrel, understand the calling pattern and infer their functions by combining the data collected by call recording and context observation. The field works was conducted in a secondary broad-leaved forest in Guanghua Village, Chiayi County from March 2012 to February 2013. Their calls and accompanied behavior were recorded for two nights per month. The results showed that they called most frequently during 7-9 pm and 3-5 am. The calls were divided into four call types based on their acoustic characteristics. Some call types were associated with one or two contexts more frequency than the other. However, the calls have different functions, namely alarm, affiliative interaction and agonistic interaction. The call rates of call type A and B were significantly higher in reproductive season than non-reproductive season. The call type A were not significantly different between male and female but different among some individuals which possibly due to their low degree of sociality.

論文審定書 i
誌謝 ii
摘要 iii
Abstract iv
Contents v
Figure Legend vi
Table Legend vii
Introduction 1
Materials and Methods 5
Results 12
Discussion 18
Literature Cited 25
Figures 33
Tables 40

Airapetyants, A. E., and I. M. Fokin. 2003. Biology of European flying squirrel Pteromys volans L. (Rodentia: Pteromyidae) in the North-West of Russia. Russian Journal of Theriology 2: 105-113.
Aschoff, J. 1966. Circadian Activity Pattern with Two Peaks. Ecology 47: 657-662.
Babu, S., and E. A. Jayson. 2009. Anti-predator behaviour of Large brown flying squirrel (Petaurista philippensis): is this an effective census method to survey the species? Current Science 96: 772-773.
Blumstein, D. T., and K. B. Armitage. 1997. Does sociality drive the evolution of communicative complexity? A comparative test with ground-dwelling sciurid alarm calls. The American Naturalist 150: 179-200.
Blumstein, D. T., and O. Munos. 2005. Individual, age and sex-specific information is contained in yellow-bellied marmot alarm calls. Animal Behavior 69: 353-361.
Blumstein, D. T., D. T. Richardson, L. Cooley, J. Winternitz, and J. C. Daniel. 2008. The structure, meaning and function of yellow-bellied marmot pup screams. Animal Behavior 76: 1055-1064.
Bohn, K. M., G. S. Wilkinson, and C. F. Moss. 2007. Discrimination of infant isolation calls by female greater spear-nosed bats, Phyllostomus hastatus. Animal Behavior 73: 423-432.
Bouchet, H., A. S. Pellier, C. Blois-Heulin, and A. Lemasson. 2010. Sex differences in the vocal repertoire of adult red-capped mangabeys (Cercocebus torquatus): A multi-level acoustic analysis. American Journal of Primatology 72: 360-375.
Digweed, S. M., and D. Rendall. 2009. Predator-associated vocalizations in North American red squirrels, Tamiasciurus hudsonicus: are alarm calls predator specific? Animal Behavior 78: 1135-1144.
Dinets, V. 2011. Observations of the woolly flying squirrel Eupetaurus cinereus in Pakistan. Mammalia 75: 277-280.
Gilley, L. M. 2013. Discovery and Characterization of High-frequency Calls in North American Flying Squirrels (Glaucomys sabrinus and G. volans): Implications for Ecology, Behavior, and Conservation, Ph.D. Dissertation, Auburn University, Auburn, Alabama.
Greene, E., and T. Meagher. 1998. Red squirrels, Tamiasciurus hudsonicus, produce predator-class specific alarm calls. Animal Behavior 55: 511-518.
Hanski, I. K., and V. Selonen. 2009. Female-biased natal dispersal in the Siberian flying squirrel. Behavioral Ecology 20: 60-67.
Howell, A. H. 1918. Revision of the American flying squirrels. North American Fauna: 1-64.
Katona, K., O. Váczi, and V. Altbäcker. 2002. Topographic distribution and daily activity of the European ground squirrel population in Bugacpuszta, Hungary. Acta Theriologica 47: 45-54.
Kawamichi, T. 1997. Seasonal changes in the diet of Japanese giant flying squirrels in relation to reproduction. Journal of Mammalogy 78: 204-212.
Kawamichi, T. 2010. Biannual reproductive cycles in the Japanese giant flying squirrel (Petaurista leucogenys). Journal of Mammalogy 91: 905-913.
Kawamichi, T., M. Kawamichi, and R. Kishimoto. 1987. Social organization of solitary mammals. pp. 173-188 in Y. Ito, J. L. Brown, and J. Kikkawa, eds. Animal Societies: Theories and Facts. Japan Science Society Press, Tokyo, Japan.
Kondo, N., and S. Watanabe. 2009. Contact calls: Information and social function. Japanese Psychological Research 51: 197-208.
Krebs, J. R., and N. B. Davies. 1993. The design of signals: ecology and evolution. pp. 349-374. in J. R. Krebs, and N. B. Davies, eds. An Introduction to Behavioural Ecology. 3rd ed., Wiley-Blackwell, Oxford, England.
Krenz, M. C. 1977. Vocalizations of the Rock Squirrel (Spermophilus variegatus), Master thesis, Texas Tech University, Lubbock, TX.
Kuo, C. C., and L. L. Lee. 2003. Food availability and food habits of Indian giant glying squirrels (Petaurista philippensis) in Taiwan. Journal of Mammalogy 84: 1330-1340.
Lee, P. F., Y. S. Lin, and D. R. Progulske. 1993. Reproductive Biology of the Red-Giant Flying Squirrel, Petaurista petaurista, in Taiwan. Journal of Mammalogy 74: 982-989.
Lee, P. F., D. R. Progulske, and Y. S. Lin. 1993. Spotlight counts of giant flying squirrels (Petaurista petaurista and P. alborufus) in Taiwan. Bulletin of the Institute of Zoology Academia Sinica 32: 54-61.
Leger, D. W. 1993. Contextual sources of information and responses to animal communication signals. Psychological Bulletin 113: 295-304.
Lurz, P. W. W., J. Gurnell, and L. Magris. 2005. Sciurus vulgaris. Mammalian Species. American Society of Mammalogists 769: 1-10.
Manno, T. G. et al. 2007. Why do male Columbian ground squirrels give a mating call? Animal Behavior 74: 1319-1327.
Matrosova, V., D. Blumstein, I. Volodin, and E. Volodina. 2011. The potential to encode sex, age, and individual identity in the alarm calls of three species of Marmotinae. Naturwissenschaften 98: 181-192.
Matrosova, V., I. Schneiderová, I. Volodin, and E. Volodina. 2012. Species-specific and shared features in vocal repertoires of three Eurasian ground squirrels (genus Spermophilus). Acta Theriologica 57: 65-78.
Matrosova, V. A., I. A. Volodin, E. V. Volodina, and N. A. Vasilieva. 2010. Stability of acoustic individuality in the alarm calls of wild yellow ground squirrels Spermophilus fulvus and contrasting calls from trapped and free-ranging callers. Naturwissenschaften 97: 707-715.
McCowan, B., and S. L. Hooper. 2002. Individual acoustic variation in Belding's ground squirrel alarm chirps in the High Sierra Nevada. Journal of the Acoustical Society of America 111: 1157-1160.
McComb, K., D. Reby, L. Baker, C. Moss, and S. Sayialel. 2003. Long-distance communication of acoustic cues to social identity in African elephants. Animal Behavior 65: 317-329.
Mougeot, F., and V. V. Bretagnolle. 2000. Predation as a cost of sexual communication in nocturnal seabirds: an experimental approach using acoustic signals. Animal Behavior 60: 647-656.
Oshida, T., L. K. Lin, S. W. Chang, Y.-J. Chen, and J.-K. Lin. 2011. Phylogeography of two sympatric giant flying squirrel subspecies, Petaurista alborufus lena and P. philippensis grandis (Rodentia: Sciuridae), in Taiwan. Biological Journal of the Linnean Society 102: 404-419.
Owren, M. J., and D. Rendall. 1997. An affect-conditioning model of nonhuman primate vocal signaling. pp. 299-346 in D. H. Owings, M. D. Beecher, and N. S. Thompson, eds. Perspectives in Ethology 12: Communication. Plenum Press, New York, USA.
Patel, E. R., and M. J. Owren. 2012. Silky sifaka (Propithecus candidus) "zzuss" vocalizations: sexual dimorphism, individuality, and function in the alarm call of a monomorphic lemur. Journal of the Acoustical Society of America 132: 1799-1810.
Rendall, D., and M. J. Owren. 2002. Animal Vocal Communication: Say What? pp. 307-313 in M. Bekoff, C. Allen, and G. M. Burghardt, eds. The Cognitive Animal: Empirical and Theoretical Perspectives on Animal Cognition. MIT Press, Cambridge, MA.
Schneiderova, I., and R. Policht. 2011. Alarm calls of the european ground squirrel Spermophilus citellus and the taurus ground squirrel S. taurensis encode information about caller identity. Bioacoustics 20: 29-43.
Schneiderova, I., and R. Policht. 2012. Acoustic analysis of the alarm call of the Anatolian ground squirrel Spermophilus xanthoprymnus: a description and comparison with alarm calls of the Taurus S. taurensis and European S. citellus ground squirrels. Naturwissenschaften 99: 55-64.
Seyfarth, R. M., D. L. Cheney, and P. Marler. 1980. Vervet monkey alarm calls: semantic communication in a free-ranging primate. Animal Behavior 28: 1070-1094.
Soltis, J., K. Leong, and A. Savage. 2005. African elephant vocal communication II: rumble variation reflects the individual identity and emotional state of callers. Animal Behavior 70: 589-599.
Tamura, N. 1995. Postcopulatory mate guarding by vocalization in the Formosan squirrel. Behavioral Ecology and Sociobiology 36: 377-386.
Tyack, P. L. 1997. Development and social functions of signature whistles in bottlenose dolphins Tursiops truncatus. Bioacoustics 8: 21-46.
Vignal, C., N. Mathevon, and S. Mottin. 2004. Audience drives male songbird response to partner's voice. Nature 430: 448-451.
Vignal, C., N. Mathevon, and S. Mottin. 2008. Mate recognition by female zebra finch: analysis of individuality in male call and first investigations on female decoding process. Behavioural Processes 77: 191-198.
Wilson, D. R., and C. S. Evans. 2008. Mating success increases alarm-calling effort in male fowl, Gallus gallus. Animal Behavior 76: 2029-2035.
Yeon, S. C. 2012. Acoustic communication in the domestic horse (Equus caballus). Journal of Veterinary Behavior-Clinical Applications and Research 7: 179-185.