本研究首次發現鱗翅目 (Lepidoptera) 幼蟲在糞鏈 (frass chain) 頂端的振動行為，以及證明「振動」可大幅增加幼蟲的存活率。獵物如何在捕食中存活下來是很重要的課題，經選汰下的獵物已發展出多樣化的防禦策略。建造糞鏈是鱗翅目幼蟲行為防禦的一種。糞鏈與葉主脈共同組成幼蟲停棲的「蟲巢」。過去研究已證明幼蟲藉由躲避於糞鏈上而防止螞蟻捕食。幼蟲保持靜止不動，而且螞蟻也不能爬上蟲巢。然而懸巢舉尾蟻 (Crematogaster rogenhoferi) 可在蟲巢上爬行。當螞蟻快碰到糞鏈上的異紋帶蛺蝶 (Athyma selenophora) 幼蟲，幼蟲會藉由振動迫使螞蟻撤離。而這種振動行為目前尚未被報導。本研究探討異紋帶蛺蝶的「蟲巢」、「糞鏈」以及「振動」對於懸巢舉尾蟻的防禦效果。結果證實「振動」是高效率的防禦策略。在防禦入侵蟲巢的天敵方面，「振動」可能比「糞鏈」與「蟲巢」扮演著更重要的角色。而多了「糞鏈」的蟲巢比起單純的葉主脈蟲巢，有更高的幼蟲存活率。本研究對於節肢動物的防禦行為，提供了新的資訊。

This study first discovered lepidopteran larvae could vibrate on frass chains hence increasing survival rate by vibration. How to survive during predation events is very important for prey. Under selection pressure, prey had developed a variety of defensing strategies. Constructing frass chain is one of the caterpillars’ behavioral defenses. Frass chain and mid-vein of a leaf formed a “refuge” for caterpillars. In previous studies, it had been demonstrated that caterpillars on frass chains could reduce predation rate of ants and keep motionless upon the approach of ants. The ants couldn’t climb on refuges. By contrast, Crematogaster rogenhoferi could climb on frass chains where the larvae of Athyma selenophora rested. When the ants were about to touch the larva on the frass chain, the larva would vibrate immediately. This might make the ants fall down from the frass chain. This vibration behavior had not been previously reported. Our study would examine how “refuge”, “frass chain”, and “vibration” would lower the predation rate respectively. We proved that “vibration” was an efficient defensing strategy. Compared to “frass chain” and “refuge”, “vibration” might play an important role on defensing. Refuge with frass chain had higher survival rate than mid-vein of a leaf. This study provided new information about defensive behavior of arthropod.

Aiello A. 1984. Adelpha (Nymphalidae): deception on the wing. Psyche. 91:1-45.
Aiello A. 1991. Adelpha ixia leucas: immature stages and position within Adelpha (Nymphalidae). J Lepid Soc. 45:181-187.
Barbosa EP, Kaminski LA, Freitas AVL. 2010. Immature stages of the butterfly Diaethria clymena janeira (Lepidoptera: Nymphalidae: Biblidinae). Zoologia (Curitiba). 27:696-702.
Barton AM. 1986. Spatial variation in the effect of ants on extrafloral nectary plant. Ecology. 67:495-504.
Berenbaum MR, Green ES, Zangerl AR. 1993. Web costs and web defense in the parsnip webworm (Lepidoptera: Oecophoridae). Environ Entomol. 22:791-795.
Bergelson JM, Lawton JH. 1988. Does foliage damage influence predation on the insect herbivores of birch? Ecology. 69:434-445.
Bernays EA. 1997. Feeding by lepidopteran larvae is dangerous. Ecol Entomol. 22:121-123.
Córdoba-Alfaro J, Murillo-Hiller LR. 2014. Early stages and natural history of Memphis perenna lankesteri butterfly (Nymphalidae, Charaxinae) from Costa Rica with the first published female pictures. Annu Res Rev Biol. 4:154-162.
Caldas A. 1994. Biology of Anaea ryphea (Nymphalidae) in Campinas, Brazil. J Lepid Soc. 48:248-257.
Caldas A. 1996. Intraseasonal variation in a population of Fountainea ryphea (Cramer) (Lepidoptera, Nymphalidae). Rev Bras Zool. 13:399-404.
Cushing PE. 1989. Possible eggsac defense behaviors in the spider Uloborus Glomosus (Araneae: Uloboridae). Psyche. 96:269-277.
Cushing PE, Opell BD. 1990. Disturbance behaviors in the spider Uloborus glomosus (Araneae, Uloboridae): possible predator avoidance strategies. Can J Zool. 68:1090-1097.
DeVries PJ. 1987. The butterflies of Costa Rica and their natural history, Volume I: Papilionidae, Pieridae, Nymphalidae. Princeton (NJ): Princeton University Press.
Edmunds M. 1974. Defence in animals: a survey of anti-predator defences. Harlow (UK): Longman.
Feeny P, Blau WS, Kareiva PM. 1985. Larval growth and survivorship of the black swallowtail butterfly in central New York. Ecol Monogr. 55:167-187.
Floren A, Biun A, Linsenmair KE. 2002. Arboreal ants as key predators in tropical lowland rainforest trees. Oecologia. 131:137-144.
Freitas AVL. 1999. An anti-predator behavior in larvae of Libytheana carinenta (Nymphalidae, Libytheinae). J Lepid Soc. 53:130-131.
Freitas AVL. 2006. Immature stages of Adelpha malea goyama Schaus (Lepidoptera: Nymphalidae, Limenitidinae). Neotrop Entomol. 35:625-628.
Freitas AVL, Brown KSJ, Aiello A. 2001. Biology of Adelpha mythra feeding on Asteraceae, a novel plant family for the Neotropical Limenitidinae (Nymphalidae), and new data on Adelpha ‘species-group VII’. J Lepid Soc. 54:97-100.
Freitas AVL, Oliveira PS. 1992. Biology and behavior of the Neotropical butterfly Eunica bechina(Nymphalidae) with special reference to larval defence against ant predation. J Res Lepid. 31:1-11.
Freitas AVL, Oliveira PS. 1996. Ants as selective agents on herbivore biology: effects on the behaviour of a non-myrmecophilous butterfly. J Anim Ecol. 65:205-210.
Gomes-Filho A. 2003. Seasonal fluctuation and mortality schedule for immatures of Hypna clytemnestra (Butler), an uncommon Neotropical butterfly (Nymphalidae: Charaxinae). J Res Lepid. 37:37-45.
Greeney HF, DeVries PJ, Penz CM, Granizo-T RB, Connahs H, Stireman JO, Walla TR, Dyer LA. 2009. The early stages and natural history of Antirrhea adoptiva porphyrosticta (Watkins, 1928) in eastern Ecuador (Lepidoptera: Nymphalidae: Morphinae). J Insect Sci. 9:1-10.
Greeney HF, Dyer LA, DeVries PJ, Walla TR, Vega LS, Simbaña W, Salagaje L. 2010. Early stages and natural history of Perisama oppelii (Latreille, 1811) (Nymphalidae, Lepidoptera) in Eastern Ecuador. Kempffiana. 6:16-30.
Greeney HF, Dyer LA, Smilanich AM. 2012. Feeding by lepidopteran larvae is dangerous: a review of caterpillars’ chemical, physiological, morphological, and behavioral defenses against natural enemies. Invert Surviv J. 9:7-34.
Greeney HF, Gerardo NM. 2001. Descriptions of the immature stages and oviposition behavior of Pyrrhogyra otolais (Nymphalidae). J Lepid Soc. 54:88-90.
Greeney HF, Whitfield JB, Stireman JO, Penz CM, Dyer LA. 2011. Natural history of Eryphanis greeneyi (Lepidoptera: Nymphalidae) and its enemies, with a description of a new species of Braconid parasitoid and notes on its Tachinid parasitoid. Ann Entomol Soc Am. 104:1078-1090.
Gross P. 1993. Insect behavioral and morphological defenses against parasitoids. Annu Rev Entomol. 38:251-273.
Hölldobler B, Wilson EO. 1990. The ants. Cambridge (MA): Harvard University Press.
Jeanne RL. 1979. A latitudinal gradient in rates of ant predation. Ecology. 60:1211-1224.
Kaminski LA, Rodrigues D. 2011. Species-specific levels of ant attendance mediate performance costs in a facultative myrmecophilous butterfly. Physiol Entomol. 36:208-214.
Lin C-C, 1998. Systematic and zoogeographic studies on the ant subfamily Myrmicinae in Taiwan (Hymenoptera: Formicidae): National Taiwan University.
Machado G, Freitas AVL. 2001. Larval defence against ant predation in the butterfly Smyrna blomfildia. Ecol Entomol. 26:436-439.
Marston NL, Schmidt GT, Biever KD, Dickerson WA. 1978. Reaction of five species of soybean caterpillars to attack, by the predator, Podisus maculiventris. Environ Entomol. 7:53-56.
Murphy SM, Leahy SM, Williams LS, Lill JT. 2010. Stinging spines protect slug caterpillars (Limacodidae) from multiple generalist predators. Behav Ecol. 21:153-160.
Myers JH, Smith JNM. 1978. Head flicking by tent caterpillars: a defensive response to parasite sounds. Can J Zool. 56:1628-1631.
Oliveira PS, da Silva AF, Martins AB. 1987. Ant foraging on extrafloral nectaries of Qualea grandiflora (Vochysiaceae) in cerrado vegetation: ants as potential antiherbivore agents. Oecologia. 74:228-230.
Oliveira PS, Freitas AVL. 1991. Hostplant record for Eunica bechina magnipunctata (Nymphalidae) and observations on oviposition sites and immature biology. J Res Lepid. 30:140-141.
Olofsson M, Jakobsson S, Wiklund C. 2012. Auditory defence in the peacock butterfly (Inachis io) against mice (Apodemus flavicollis and A. sylvaticus). Behav Ecol Sociobiol. 66:209-215.
Prop N. 1960. Protection against birds and parasites in some species of tenthredinid larvae. Arch Néerl Zool. 13:380-447.
Richard F-J, Fabre A, Dejean A. 2001. Predatory behavior in dominant arboreal ant species: the case of Crematogaster sp. (Hymenoptera: Formicidae). J Insect Behav. 14:271-282.
Robinson MH, Valerio CE. 1977. Attacks on large or heavily defended prey by tropical Salticid spiders. Psyche. 84:1-10.
Ruxton GD, Sherratt TN, Speed MP. 2004. Avoiding attack: the evolutionary ecology of crypsis, warning signals and mimicry. Oxford (UK): Oxford University Press.
Schoener TW, Spiller DA. 1992. Stabilimenta characteristics of the spider Argiope argentata on small islands: support of the predator-defense hypothesis. Behav Ecol Sociobiol. 31:309-318.
Scoble MJ. 1992. The Lepidoptera: form, function and diversity. Oxford (UK): Oxford University Press.
Stamp NE. 1982. Behavioral interactions of parasitoids and Baltimore checkerspot caterpillars (Euphydryas phaeton). Environ Entomol. 11:100-104.
Sugiura S, Yamazaki K. 2014. Caterpillar hair as a physical barrier against invertebrate predators. Behav Ecol. 25:975-983.
Teshirogi M. 2004. Life history of Agrias beatifica beata Staudinger, 1886 (Lepidoptera, Nymphalidae). Trans Lepid Soc Japan. 55:134-146.
Villas-Boas IM, Goncalves-de-Andrade RM, Pidde-Queiroz G, Assaf SLMR, Portaro FCV, Sant'Anna OA, van den Berg CW, Tambourgi DV. 2012. Premolis semirufa (Walker, 1856) envenomation, disease affecting rubber tappers of the Amazon: searching for caterpillar-bristles toxic components. PLoS neglected tropical diseases. 6:e1531.
Weiss MR. 2003. Good housekeeping: why do shelter-dwelling caterpillars fling their frass? Ecol Lett. 6:361-370.
Weiss MR. 2006. Defecation behavior and ecology of insects. Annu Rev Entomol. 51:635-661.
Zalucki MP, Clarke AR, Malcolm SB. 2002. Ecology and behavior of first instar larval Lepidoptera. Annu Rev Entomol. 47:361-393.