護巢行為是一項提升繁殖成功率的重要策略，不同鳥種面對不同威脅程度的天敵時，可能會表現出不同的護巢行為。灰頭鷦鶯（Prinia flaviventris）和褐頭鷦鶯（P. inornata）是兩種共域、且親緣關係相當接近的物種，牠們的形態和生態習性相似，也棲息在同樣的環境中，不過在某些繁殖特性上有所差異。在本研究樣區中，這兩種鷦鶯受到來自相同掠食者的捕食風險，但可能面臨不同的托卵寄生壓力。本研究欲探討牠們在不同繁殖階段對不同天敵的反應。在巢邊擺放掠食者（棕背伯勞 Lanius schach）、托卵性鳥類（中杜鵑 Cuculus Saturatus）和控制組（白腹鶇 Turdus pallidus）等鳥類標本，並記錄親鳥的行為表現。結果顯示兩種鷦鶯皆對掠食者有顯著的躲避行為，牠們不攻擊、也鮮少對棕背伯勞標本表現威嚇行為，且不因繁殖階段而有差異。然而，這兩種鷦鶯的行為不盡相同：灰頭鷦鶯會表現較多的警戒聲、接近和躲藏行為；而褐頭鷦鶯則傾向遠離掠食者，並以警戒聲吸引其注意力。灰頭鷦鶯在卵期對中杜鵑標本表現顯著的攻擊行為，在雛期則否。雖然灰頭鷦鶯對中杜鵑的其他行為（威嚇和接近）和對控制組（白腹鶇）的行為無顯著差異，但在兩個繁殖階段都有相當強烈的行為表現。相較之下，褐頭鷦鶯則無顯著的防禦托卵性鳥類行為。強烈的護巢行為和卵色的高度模擬，都顯示灰頭鷦鶯可能是中杜鵑在台灣的主要宿主。本研究結果指出兩種共域鷦鶯各自具有不同的護巢行為：當面對相同的掠食者時，牠們會使用不同的防禦策略；對於托卵性鳥類的反應也截然不同，這可能反映了兩種鷦鶯受到的寄生壓力有所差異。

Nest-guarding is an important strategy for successful reproduction. Birds may adopt different nest defense behavior according to the degree of threat from the intruders. The yellow-bellied prinia (Prinia flaviventris) and plain prinia (P. inornata) are two sympatric, phylogenetically closely related species. They are similar in size and share the same feeding ecology, but differ in some breeding traits. They suffer predation risk from the same avian predator, and may have different pressures from brood parasites. In this study, we tested their responses to the intruders during different breeding stages. We presented mounted specimens of an avian predator (long-tailed shrike, Lanius schach), brood parasite (oriental cuckoo, Cuculus optatus) and a control (pale thrush, Turdus pallidus) near the nest and recorded the nest owners’ responses. Both species showed significant avoidance to the predator regardless of breeding stages. They did not attack and performed very few mobbing behaviors toward the shrike model. However, they differed in other behaviors: the yellow-bellied prinias were more alarmed by the shrike, with more approachings and hidings, while the plain prinia tended to be away from the shrike and used alarm calls to distract the predator. For the responses to the brood parasites, the yellow-bellied prinia displayed significant attacking behavior toward the cuckoo model at the egg stage, but not in the nestling stage. The plain prinias showed no significant anti-cuckoo behavior at either breeding stage. The strong aggression toward the cuckoo and high similarity in their egg coloration indicates that the yellow-bellied prinia may be the main host for the oriental cuckoo. Our results highlighted the different nest guarding strategies of the two species: When facing the same predator, they used different nest defense behaviors; the responses to the brood parasites also differed, possibly reflecting their susceptibility to being brood-parasitized.

1. 前言 1
1.1 護巢行為 1
1.2 鳥類掠食者（avian predators）的威脅 2
1.3 托卵性鳥類（brood parasites）的威脅 3
1.4 兩種共域鷦鶯 6
1.5 兩種鷦鶯的天敵 6
1.6 研究目的 10
2. 研究材料與方法 11
2.1 研究樣區與鳥巢搜尋 11
2.2 標本實驗 11
2.3 研究樣區與鳥巢搜尋 14
3. 結果 15
3.1 灰頭鷦鶯對三種鳥類標本的反應 15
3.2 褐頭鷦鶯對三種鳥類標本的反應 21
3.3 兩種鷦鶯行為反應的比較 22
4. 討論 29
4.1 兩種鷦鶯對托卵性鳥類的護巢行為比較 29
4.2 兩種鷦鶯對掠食者的護巢行為比較 31
5. 引用文獻 35

林文宏。2010。杜鵑科。出自：劉小如，丁宗蘇，方偉宏，林文宏，蔡牧起，顏重威，台灣鳥類誌（中），442-469頁。行政院農業委員會林務局。台北。
蔡牧起。2010a。扇尾鶯科。出自：劉小如，丁宗蘇，方偉宏，林文宏，蔡牧起，顏重威，台灣鳥類誌（下），161-169頁。行政院農業委員會林務局。台北。
蔡牧起。2010b。鶇科。出自：劉小如，丁宗蘇，方偉宏，林文宏，蔡牧起，顏重威，台灣鳥類誌（下），418-420頁。行政院農業委員會林務局。台北。
Abrams, P. A. 2000. The evolution of predator-prey interactions: theory and evidence. Annual Review of Ecology and Systematics 31:79-105.
Altmann, S. A. 1956. Avian mobbing behavior and predator recognition. Condor 58:241-253.
Antonov, A., B. G. Stokke, J. R. Vikan, F. Fossøy, P. S. Ranke, E. Røskaft, A. Moksnes, A. P. Møller and J. A. Shykoff. 2010. Egg phenotype differentiation in sympatric cuckoo Cuculus canorus gentes. Journal of Evolutionary Biology 23:1170-1182.
Boncoraglio, G., N. Saino and L. Z. Garamszegi. 2008. Begging and cowbirds: brood parasites make host scream louder. Behavioral Ecology 20:215-221.
Brooke, M. de L. and N. B. Davies. 1988. Egg mimicry by cuckoos Cuculus canorus in relation to discrimination by hosts. Nature 335:630-632.
Campobello D. and S. G. Seally. 2010. Enemy recognition of reed warblers (Acrocephalus scirpaceus): threats and reproductive value act independently in nest defence modulation. Ethology 116:498-508.
Chen, W. J., P. F. Lee and R. S. Lin. 2009. The first record of a cuckoo (Cuculus saturatus) killing nestlings in a potential host’s nest. Endemic Species Research 11:63-67.
Colombelli-Négrel, D., R. Jeremy. and K. Sleindorfer. 2010. Nestling presence affects the anti-predator response of adult superb fairy-wrens (Malurus cyaneus). Acta Ethologica 13:69-74.
Davies, N. B. 2000. Cuckoos, cowbirds and other cheats. T. & AD Poyser. London.
Davies N. B., S. H. M. Butchart, T. A. Burke, N. Chaline and I. R. K. Stewart. 2003. Reed warblers guard against cuckoos and cuckoldry. Animal Behaviour 65:285-295.
Dearborn, D. C. 1998. Begging behavior and food acquisition by brown-headed cowbird nestlings. Behavioral Ecology and Sociobiology 43:259-270.
Delm, M. M. 1990. Vigilance for predators: detection and dilution effects. Behavioral Ecology and Sociobiology 26:337-342.
Evans, C. S., L. Evans and P. Marler. 1993. On the meaning of alarm calls: functional reference in an avian vocal system. Animal Behaviour 46:23-38.
Fernández-Juricic, E., J. Jokimäki, J. C. McDonald, F. Melado, A. Toledano, C. Mayo, B. Martín, I. Fresneda and V. Martín. 2004. Effects of opportunistic predation on anti-predator behavioural responses in a guild of ground foragers. Behavioral Ecology 140:183-190.
Forbes, M. R. L., R. G. Clark, P. J. Weatherhead and T. Armstrong. 1994. Risk-taking by female ducks: intra- and interspecific tests of nest defense theory. Behavioral Ecology and Sociobiology 34:79-85.
Gill, S. A. and S. G. Seally. 1996. Nest deffence by yellow warbler: recognition of a brood parasite and an avian nest predator. Behaviour 133:263-282.
Gill, S. A., D. L. Neudorf and S. G. Sealy. 1997. Host responses to cowbirds near the nest: cues for recognition. Animal Behaviour 53:1287-1293.
Greig-Smith, P. W. 1980. Parental investment in nest defence by stonechats（Saxicola torquata）. Animal Behaviour 28:604-619.
Grim, T. 2005. Host recognition of brood parasites: implications for methodology in studies of enemy recognition. Auk 122:530-543.
Hauber, M. E. and C. K. Ramsey. 2003. Honesty in host-parasite communication signals: the case for begging by fledgling brown-headed cowbirds Molothrus ater. Journal of Avian Biology 34:339-344.
Hauber, M. E. and C. Moskát. 2007. Shared parental care is costly for nestling of common cuckoos and their great reed warbler hosts. Behavioral Ecology 1093:79-86.
Honza, M., T. Grim, M. Jr Čapek, A. Moksnes and E. Røskaft. 2004. Nest defence, enemy recognition and nest in speciation behavior of experimentally parasitized reed warblers Acrocephalus scirpaceus. Bird Study 51:256-263.
Honza, M., V. Šicha, P. Procházka and R. Ležalová. 2006. Host nest defense against a color-dimorphic brood parasite: great reed warblers（Acrocephalus arundinaceus）versus common cuckoos（Cuculus canorus）. Journal of Ornithology 147:629-637.
Hromada, M., P. Tryjanowski and M. Antczak. 2002. Presence of the great grey shrike Lanius excubitor affects breeding passerine assemblage. Annales Zoologici Fennici 39:125-130.
Hudson, P. J. and D. Newborn. 1990. Brood defence in a precocial species: variations in the distraction displays of red grouse, Lagopus lagopus scoticus. Animal Behaviour 40:254-261.
Johnsgard, P. A. 1997. The avian brood parasites. Oxford University Press. New York.
Knight, R. L. and S. A. Temple. 1986. Nest defence in the American goldfinch. Animal Behavior 34:887-897.
Lahti, D. C. 2005. Evolution of bird eggs in the absence of cuckoo parasitism. Proceedings of the National Academy of Sciences of the United States of America 102:18057-18062.
Langmore, N. E., S. Hunt and R. M. Kilner. 2003. Escalation of a coevolutionary arms race through host rejection of brood parasitic young. Nature 422:157-160.
Langmore, N. E., G. Maurer, G. J. Adcock and R. M. Kilner. 2008. Socially acquired host-specific mimicry and the evolution of host races in Horsfield's bronze-cuckoo Chalcites basalis. Evolution 62:1689-1699.
Lima, S. L. 1998. Nonlethal effects in the ecology of predator-prey interactions. BioScience 48:25-34.
Lin, L. S., P. S. Alexander and L. L. Severinghaus. 1988. Breeding ecology of tawny and yellow-bellied wren warblers（Prinia subflava and P. flaviventris）. Bulletin of the Institute of Zoology, Academia Sinica 27:57-66.
Lorek, G., P. Tryjanowski and J. Lorek. 2000. Birds as prey of the Great Grey Shrike (Lanius excubitor). Ring 22:37-44.
Lyon, B. E. 2003. Egg recognition and counting reduce costs of avian conspecific brood parasitism. Nature 422:495-499.
Lyon, B. 2007. Mechanism of egg recognition in defenses against conspecific brood parasitism: American coots（Fulica americana） know their own eggs. Behavioral Ecology Sociobiology 61:453-463.
Macedonia, J. M. and C. S. Evans. 1993. Variation among mammalian alarm call systems and the problem of meaning in animal signs. Ethology 93:177-197.
Marchetti, K. 1992. Costs to host defence and the persistence of parasitic cuckoos. Biological Science 248:41-45.
Martin, T. E. 1993a. Nest predation among vegetation layers and habitat types: revising the dogmas. The American Naturalist 141:897-913.
Martin, T. E. 1993b. Nest predation and net sites. BioScience 43:523-532.
Moksnes, A., E. Røskaft, A. T. Braa, L. Korsnes, H. M. Lampe and H. C. Pedersen. 1991. Behavioural responses of potential hosts towards artificial cuckoo eggs and dummies. Behaviour 116:64-89.
Montgomerie, R. D. and P. J. Weatherhead. 1988. Risk and rewards of nest defense by parent birds. Quarterly Review of Biology 63:163-187.
Moskát, C. 2005. Nest defence and egg rejection in great reed warblers over the breeding cycle: are they synchronised with the risk of brood parasite. Annales Zoologici Fennici 42:579-586.
Padilla, D. P., M. Nogales, and J. P. Antonio. 2005. Seasonal diet of an insular endemic population of Southern Grey Shrike Lanius meridionalis koenigi on Tenerife, Canary Islands. Ornis Fennica 82:155-165.
Palleroni, A., C.T. Miller, M. Hauser and P. Marler. 2005. Predation: prey plumage adaptation against falcon attack. Nature 434:973-974.
Phillips, J., E. Nol, D. Burke and W. Dunford. 2005. Impacts of housing developments on Wood Thrush nesting success in hardwood forest fragments. Condor 107:97-106.
Redondo, T. 1989. Avian nest defence: theoretical models and evidence. Behaviour 111:161-195.
Ricklefs, R. E. 1969. An analysis of nesting mortality in birds. Smithsonian Contributions to Zoology 9:1-48.
Rivers, S. W. 2007. Nest mate size, but not short-term need, influences begging behavior of a generalist brood parasite. Behavioral Ecology 18:222-230.
Rohner, C., N. James, M. Joyce, F. I. Doyle and R. Boonstra. 1995. Northern hawk-owls in the nearctic boreal forest: Prey selection and population consequences of multiple prey cycles. Condor 97:208-220.
Røskaft, E., A. Moksnes, B. G. Stokke, T. C. Moská and M. Honza. 2002. The spatial habitat structure of host populations explains the pattern of rejection behavior in hosts and parasitic adaptations in cuckoos. Behavior Ecology 13:163–168.
Scott, D. M., P. J. Weatherhead and C. D. Akeney. 1992. Egg-eating by female brown-headed cowbirds. Condor 94:579-584.
Seally, S. G. 1992. Removal of yellow warbler eggs in association with cowbird parasitism. Condor 94:40-54.
Smith, N. G. 1975. “Spshing noise”: biology significance of its attraction and nonattraction by birds. Proceedings of the National Academy of Sciences of the United States of America 72:1411-1414.
Smith, J. N. M., M. J. Taitt and L. Zanette. 2002. Removing brown-headed cowbirds increases seasonal fecundity and population growth in song sparrows. Ecology 83:3037-3047.
Soler, M., J. J. Soler and J. G. Martínez. 1997. Great spotted cuckoos improve their reproductive success by damaging magpie host nests. Animal Behaviour 54:1227-1233.
Sordahl, T. A. 1990. The risks of avian mobbing and distraction behavior: an anecdotal review. Wilson Bulletin 102:349-352.
Stokke, B. G., L. PolaČiková, A. Dyrcz, I. Hafstad, A. Moksnes and E. Røskaft. 2010. Response of reed warbler Acrocephalus scirpaceus to non-mimetic eggs of different sizes in a nest parasitism experiment. Acta Ornithologica 45:98-104.
Strausberger, B. M. 1998. Temporal patterns of host availability, brown-headed cowbird brood parasitism, and parasite egg mass. Oecologia 116:267-274.
Tryjanowski P., M. K. Karg and J. Karg. 2003. Diet composition and prey choice by the red-backed shrike Lanius collurio in western Poland. Belgian Journal of Zoology 133:157-162.
Voous, K. H. 1969. Predation potential in birds of prey from Surinam. Ardea 57:117-148.
Welbergen, J. A. and N. B. Davies. 2009. Strategic variation in mobbing as a front line of defense against brood parasitism. Current Biology 19:235-240.
Welbergen, J. A. and N. B. Davies. 2011. A parasite in wolf’s clothing: hawk mimicry reduces mobbing of cuckoos by hosts. Behavioral Ecology 22:574-579.
Yoram, Y. T. 1980. Intraspecific nest parasitism in birds. Biological Reviews 55:93-108.
Yoram, Y. T. 2001. An updated list and comments on the occurrence of intraspecific nest parasitism in birds. Ibis 143:133-143.