本研究以三斑虎灰蝶Spindasis syama（Horsfield, 1829）為題材，探討其與阿美舉尾家蟻Crematogaster amia Forel, 1913的共生關係。研究範圍為共生蟻對雌蝶產卵行為、幼生期成長發育表現與幼蟲野外存活的影響，進而討論三斑虎灰蝶在共生蟻與天敵共同作用下，所表現的食性利用範圍。
　　研究結果發現三斑虎灰蝶在產卵率及產卵量表現皆因共生蟻的存在而顯著提升，且在共生蟻種存在狀況下，植物種類在產卵選擇偏好上並無顯著差別；幼蟲透過與共生蟻的互動，可達到減少幼蟲期發育時間與提升自然環境中的存活表現，以上特性顯示三斑虎灰蝶是屬於一種絕對型共生（obligate mutualism）喜蟻性灰蝶（myrmecophilous lycaenid butterfly）。喜蟻性灰蝶幼蟲為降低在自然環境遭受的高捕食壓力，雌蝶產卵往往不若多數植食性昆蟲以植物種類與品質做為選擇依據，取而代之的是以共生蟻存在與否做為最為主要考量條件，以提升整體的適存度（fitness），這種用來降低天敵捕食壓力的策略被稱為enemy free space（EFS）。這樣受到天敵捕時壓力影響的產卵偏好，雖可能將卵產於幼蟲無法利用的植物，卻也同時間增加幼蟲利用多種植物的機會，進而使喜蟻性灰蝶寄主植物利用範圍的擴張。

The present study examines the effects associated with the presence of valentine ant Crematogaster amia (Formicidae) on lycaenid butterfly Spindasis syama (Lycaenidae) with regards to its egg laying behavior, larval growth performance, and survivoship in the field.
In a series of laboratory choice test, the butterfly exhibited ant-dependent egg laying behavior and did not discriminate between host and non-host plant species. When reared in the presence of ants, the butterfly larvae underwent shorter larval development time and lost more pupal weight. The adult size and weight were independent wheather a larva had been ant-tended or not. A field experiment revealed that ant attendance had significant effect on larval survivorship. I concluded that Spindasis syama is an obligate myrmecophilous lycaenid butterfly based on the ant-dependent oviposition behavior and the high mortality of larvae in the wild without ants.
In order to gain protection in the larval stage, the myrmecophilous lycaenids may use ants as cues in oviposition rather than plant species. The female occasionally oviposits on wrong plant species which could not be utilized by the larvae. This oviposition preference offer a choice that myrmecophilous lycaenids use a wider range of plant species than non-myrmecophilous lycaenids. Under the influence of the enemy free space (EFS) mechanism, how does the ant-dependent egg laying behavior affect range of host plants will be discussed in this study.

白水 隆。1960。原色台灣蝶類大圖鑑。保育社。481pp。
濱野榮次。1987。台灣蝶類生態大圖鑑。牛頓出版社。474pp。
內田春男。1991。常夏の島フォルモサは招く。自行出版。216pp。
－ 1995。麗しき蝴蝶の島よ永えに -台灣の蝶と自然と人と-。自行出版。
208pp。
周堯(主編)。1994。中國蝶類志。河南科學技術出版社。853pp。
李俊延、王效岳。1997。台灣蝶類圖說(四)。台灣省立博物館。317pp。
詹家龍。1997。兩種喜蟻性雀斑小灰蝶生態之研究（鱗翅目：小灰蝶
　科）。國立台灣大學植物病蟲害學研究所碩士論文。63pp。
林宗岐。1998。台灣產家蟻亞科系統分類學與動物地理學研究（膜翅　
　目：蟻科）。國立台灣大學植物病蟲害學研究所博士論文。748 pp。
林春吉。2008。台灣蝴蝶食草與蜜源植物大圖鑑(上)。天下文化。
　302pp。
王俊凱。2010。從生活史不同階段初探虎灰蝶與樹棲舉尾蟻的共生關
　係。國立台灣師範大學生命科學系碩士論文。98pp。
Ackery P. R. 1988. Host plants and classification: a review of nymphalid buttrtflies. Biol. J. Linn. Soc. 33:95-203.
Atsatt P. R. 1981a. Ant-dependent food plant selection by the mistletoe butterfly Ogyris amaryllis (Lycaenidae). Oecologia 48:60-63.
－ 1981b. Lycaenid butterflies and ants: selection for enemy-free space. Am. Nat. 118:638-654.
Awmack C. S. and S. R. Leather. 2002. Host plant quality and fecundity in herbivorous insects. Annu. Rev. Entomol. 47:817-844.
Axén A. H. and N. E. Pierce. 1998. Aggregation as a cost-reducing strategy for lycaenid larvae. Behav. Ecol. 9:109-115.
Ballmer G. R. and G. F. Pratt. 1988. A survey of the last instar larvae of the Lycaenidae of California. J. Res. Lepid. 27:1-81.
－ and G. F. Pratt. 1992. Quantification of ant attendance (myrmecophily) of lycaenid larvae. J. Res. Lepid. 30:95-112.
Barbero F., J. A. Thomas, S. Bonelli, E. Balletto and K. Schönrogge. 2009. Queen ants make distinctive sounds that are mimicked by a butterfly social parasite. Science 323:782-785.
Bascombe M. J., G. Johnston and F. S. Bascombe. 1999.The butterflies of Hong Kong. Natural World. 422pp.
Benrey B. and R. F. Denno. 1997. The slow-growth-high-mortality hypothesis: a test using the cabbage butterfly. Ecology 78:987-999.
Bernays E. A. and R. F. Chapman. 1994. Host-plant selection by phytophagous insects. New York: Chapman & Hall. 312pp.
Boggs C. L. 1992. Resource allocation－exploring connections between foraging and life history. Funct. Ecol. 6:508-518.
Brower L. P. and J. V. Z. Brower. 1964. Birds, butterflies and plant poisons: a study in ecological chemistry. Zoologica 49:137-159.
－ J. V. Z. Brower and J. M. Corvino. 1967. Plant posions in a terrestrial food chain. Proc. Natl. Acad. Sci. 57:893-898.
Chapman R. F. 1982. Chemoreception: the significance of receptor numbers. Adv. Insect Physiol. 16:247-356.
Clark G. C. and C. G. C. Dickson. 1971. Life history of the South African lycaenid butterflies. Cape Town: Purnell. 272pp.
Cottrell C. B. 1984. Aphytophagry in butterflies: its relationship to myrmecophily. Zool. J. Linn. Soc. 79:1-57.
Cushman J. H., V. K. Rashbrook and A. J. Beattie. 1994. Assessing benefits to both participants in a lycaenid-ant association. Ecology 75:1031-1041.
Davies N. B. 1978. Territorial defense in the speckled wood butterfly (Parage aegeria): the resident always wins. Anim. Behav. 26:138-147.
Denno R. F., S. Larsson and K. L. Olmstead. 1990. Role of enemy-free space and plant quality in host-plant selection by willow beetles. Ecology. 71(1):124-137.
DeVries P. J., D. J. Harvey and I. J. Kitching. 1986. The ant associated epidermal organs on the larvae of the lycaenid butterfly Curetis regula Evans. J. Nat. Hist. 20:621-633.
－ 1988. The larval ant-organs of Thisbe irenea (Lepidoptera: Riodinidae) and their effects upon attending ants. Zool. J. Linn. Soc. 94:379-393
－ and I. Baker. 1989. Butterfly exploitation of an ant-plant mutualism: adding insult to herbivory. J. NY Entomol. Soc. 97:332-340.
－ 1991. Mutualism between Thisbe irenea butterflies and ants, and the role of ant ecology in the evolution of larval-ant association. Biol. J. Linn. Soc. 43:179-195.
－ and C. M. Penz. 2000. Entomophagy, behavior, and elongated thoracic legs in the myrmecophilous neotropical butterfly Alesa amesis (Riodinidae). Biotropica 32:712-721.
Downey J. C. 1962. Myrmecophily in Plebejus (Icaricia) icarioides (Lepidoptera: Lycaenidae). Entomol. News 73:57-66.
Ehrlich P. R. and P. H. R. Raven. 1964. Butterflies and plants: a study in coevolution. Evolution 18:586-608.
Elgar M. A. and N. M. Pierce. 1988. Mating success and fecundity in an ant-tended lycaenid butterfly. In Reproductive Success: Studies of selection and adaptation in Contrasting Breeding Systems, TH Clutton-Brock（ed.）. Univ. of Chicago press. pp. 59-75.
Feeny P, E.Städler, I. Åhman and M. Carter. 1989.Effects of plant odor on oviposition by the black swallowtail butterfly, Papilio polyxenes (Lepidoptera: Papilionidae). J. Ins. Behav. 2:803-827.
－ 1991. Chemical constraints on the evolution of swallowtail butterflies. In: Plant-animal interactions: evolutionary ecology in tropical and temperate regions. John Wiley, New York. 639pp.
Fiedler K. and U. Maschwitz. 1989. The symbiosis between the weaver ant, Oecophylla smaragdina, and Anthene emolus, an obligate myrmecophilous lycaenid butterfly. J. Nat. Hist. 23:833-846.
－ 1991. Systematic, evolutionary, and ecological implications of myrmecophily within the Lycaenidae (Insecta: Lepidoptera: Papilionoidea). Bonn. Zool. Monogr. 31:5-157.［An electronic version of Table 17 of this work with MS additions and corrections was provided by Dr Fiedler in February 1996.］
－ and B. Hölldoobler. 1992. Ants and Polyommatus icarus immatures (Lycaenidae) – sex-related developmental benefits and costs of ant attendance. Oecologia 91:468-473.
－ and C. Saam. 1995. Ant benefit from attending facultatively myrmecophilous Lycaenidae caterpillars: evidence from a survival study. Oecologia 104:316-322.
－ 1996. Host-plant relationships of lycaenid butterflies: large-scale patterns, interactions with plant chemistry, and mutualism with ants. Entomol. Exp. Appl. 80:259-267.
－ 2001. Ants that associate with Lycaeninae butterfly larvae: diversity, ecology and biogeography. Divers. Distrib. 7:45-60.
Forister M. L., Z. Gompert, C. C. Nice, G. W. Forister and J. A. Fordyce. 2010. Ant association facilitates the evolution of diet breadth in a lycaenid butterfly. Proc. R. Soc. B. doi: 10.1098/rspb.2010.1959.
Fraser A. M., A. H. Axen and N. E. Pierce. 2001. Assessing the quality of different ant species as partners of myrmecophilous butterfly. Oecologia 129:452-460.
－ T. Tregenza, N. Wedell, M. A. Elgar and N. E. Pierce. 2002. Oviposition tests of ant preference in a myrmecophilous butterfly. J. Evol. Biol. 15:861-870.
Heath A. 1997. A review of African genera of the tribe Aphnaeini (Lepidoptera: Lycaenidae). Metamorph. Occas. Suppl. 2:1-60.
－ and A. J. M. Claassens. 2003. Ant-association among southern Aferican Lycaenidae. J. Lepid. Soc. 57(1):1-16.
Henning S. F. 1983. Chemical communication between lycaenid larvae (Lepidoptera: Lycaenidae) and ants (Hymenoptera: Formicidae). J. Entomol. Soc. S. Afr. 46:341-366.
Hölldoobler B. and E. O. Wilson 1990. The Ants. Cambridge, MA: Belknap press. Harvard Univ. 732pp.
Inbar M., H. Doostdar and R. T. Mayer. 2001. Suitability of stressed and vigorous plants to various insect herbivores. Oikos 94:228-235.
Janz N. and S. Nylin. 1998. Butterflies and plants: a phylogenetic study. Evolution 52:486-502.
Jeffries M. J. and J. H. Lawton. 1984. Enemy free space and the structure of ecological communities. Biol. J. Linn. Soc. 23:269-286.
Jermy T., A. Szentesi and J. Horváth. 1988. Host plant finding in phytophagous insects: the case of the Colorado potato beetle. Entomol. Exp. Appl. 49:83-98.
Johnson S. J. and P. S. Valentine. 1986. Observations on Liphyra brassolis Westwood (Lepidoptera: Lycaenidae) in North Queensland. Aust. Entomol. Mag. 13:22-26.
Judd G. T. R. and J. H. Borden. 1989. Distant olfactory response of the onion fly, Delia antique, to host-plant odour in the field. Physiol. Entomol. 14:429-441.
Kalberer N. M., T. C. J. Turlings and M. Rahier. 2001. Attraction of a leaf beetle (Oreina cacaliae) to damaged host plants. J. Chem. Ecol. 27:647-661.
Kitching R. L. 1987. Aspects of the natural history of the lycaenid butterfly Allotinus major in Sulawesi. J. Nat. Hist. 21:535-544.
Leather S. R. and A. F. G. Dixon.1981. The effect of cereal growth stage and feeding site on the reproductive activity of the bird cherry aphid Rhopalosiphum padi. Ann. Appl.Biol. 97:135-141.
Lederhouse R. C. 1982. Territorial defense and lek behavior of the black swallowtail butterfly, Papilio polyxenes. Behav. Ecol. Sociobiol. 10:109-118.
Lee L. H. Y. and Winney R. 1981. Check list of agricultural insects of Hong Kong. Hong Kong Agriculture and Fisheries Department Bulletin. 2. 164pp.
Loon J. J. A. van. 1996. Chemosensory basis of feeding and oviposition behaviour in herbivorous insects: a glance at the periphery. Entomol. Exp. Appl. 80:7-13.
Maschwitz U., M. Wüst and K. Schurian. 1975. Bläulingsraupen als Zuckerlidferanten für Ameisen. Oecologia 18:17-21.
Moon D. C. and P. Stiling. 2006. Trade-off in oviposition strategy: choosing poor quality host plants reduces mortality from natural enemies for a salt marsh planthopper. Ecol. Entomol. 31:236-241.
Murphy S. M. 2004. Enemy-free space maintains swallowtail butterfly host shift. Proc. Natl. Acad. Sci. 101:18048-18052.
Pierce, N. E. and P. S. Mead. 1981. Parasitoids as selective agents in the symbiosis between lycaenid butterfly larvae and ants. Science 211:1185-1187.
－ and M. A. Elgar. 1985. The influence of ants on host plat selection by Jalmenus evagoras, a myrmecophilous lycaenid butterfly. Behav. Ecol. Sociobiol. 16:209-222.
－ and S. Easteal. 1986. The selective advantage of attendant ants for the larvae of a lycaenid butterfly, Glaucopsyche lygdamus. J. of Am. Ecol. 55(2):451-462
－ 1987. The evolution biogeography of associations between lycaenid butterflies and ants. In: Harvey PH, Partridge L, eds. Oxford surveys in evolutionary biology.Oxford Univ. press. 4:89-116.
－ R. L. Kitching, R. C. Buckley, M. F. J. Taylor and K. F. Benbow. 1987. The costs and benefits of cooperation between the Australian lycaenid butterfly, Jalmenus evagoras, and its attendant ants. Behav. Ecol. Sociobiol. 21:237-248.
－ 1989. Butterfly-ant mutualisms. In Toward a More Exact Ecology. Oxford: Blackwell Sci. 447pp.
－ and D. R. Nash. 1999. The Imperial Blue: Jalmenus evagoras (Lycaenidae). In The Biology of Australian Butterflies. Sydney: CSIRO Press. 401pp.
－ 1995. Predatory and parasitic Lepidoptera: carnivores living on plants. J. Lepid. Soc. 49:412-453.
－ M. F. Braby, A. Heath, D. J. Lohman, J. Mathew, D. B. Rand and M. A. Travassos. 2002. The ecology and evolution of ant association in the Lycaenidae (Lepidoptera). Annu. Rev. Entomol. 47:733-771.
Price P. W., C. E. Bouton, P. Gross, B. A. McPheron, J. N. Thompson and A. E. Weis. Interactions among three tropic levels: influence of plants on interactions between insect herbivores and natural enemies. Annu. Rev. Ecol. Sys. 11:41-65.
－ 1991. The plant vigor hypothesis and herbivore attack. Oikos 62:244-251.
Robbins R. K. 1991. Cost and evolution of a facultative mutualism between ants and lycaenid larvae (Lepidoptera). Oikos 62:363-369.
Robinson G. S., P. R. Ackery, I. J. Kitching, G. W. Becaloni and L. M. Hernandez. 2001. Hostplants of the moth and butterfly caterpillars of the Oriental Region. The Natural History Museum and Southdene SDN BHD. 744pp.
Rossiter M. C., D. L. Cox-Foster and M. A. Briggs. 1993. Initiation of maternal effects in Lymantria dispar－genetic and ecological components of egg provisioning. J. Evol. Biol. 6:577-590.
－ 1996. Incidence and consequences of inherited environmental dffects. Annu. Rev. Ecol. Syst. 27:451-476.
Rothschild M., T. Reichstein, J. von Euw, R. Alpin and R. P. M. Harman. 1970. Toxic Lepidoptera. Toxicon 8:293-299.
Schroth M. and U. Maschwitz. 1984. Zur larvalbiologie und wirtsfindung von Maculinea teleius (Lepidoptera: Lycaenidae), eines parasite von Myrmica laevinoids (Hymenoptera: Formicidae). Entomol. Gener. 9(4):225-230.
Thomas J. 1980. Why did the Large Blue become extinct in Britain? Oryx 15(3):243-247.
Van-Wright R. I. 1978. Ecological and behavioural origins of diversity in butterflies. In: The diversity of insect faunas. Symposia of the Royal Entomological Society of London 9:56-70.
Wagner D. and del Rio C. Martinez. 1997. Experimental tests of the mechanism for ant-enhanced growth in an ant-tend lycaenid butterfly. Oecologia 112:424-429.
Weeks J. A. 2003. Parasitism and ant protection alter the survival of the lycaenid Hemiargus isola. Ecol. Entomol. 28:228-232.
Wickman P-O. 1985. Territorial defense and mating success in males of the small heath butterfly, Coenonympha pamphilus L. (Lepidoptera: Satyridae). Anim. Behav. 33:1162-1168.
Wilson E. O. 1971. The insect societies. Belknap press of Harvard Univ. press. Cambridge. Mass. 548pp.