Journal of Insect Behavior

, Volume 28, Issue 4, pp 403–416 | Cite as

Interspecific Variation in Aggressive Fighting Behavior of Shelter-Building Caterpillars



Herbivorous insects rarely directly compete over food resources, except in the case of concealed feeders when the food source also serves as shelter and territory. The objective of this study was to characterize the competitive behaviors of four common, co-occurring caterpillar species that tie together overlapping leaves on white oak – Psilocorsis quercicella (Oecophoridae), P. cryptolechiella, P. reflexella, and Pseudotelphusa quercinigracella (Gelechiidae). Artificial leaf ties were created by clipping a piece of transparency paper to a white oak leaf. An ‘occupant’ caterpillar was allowed to build a shelter between the transparency paper and leaf, after which an ‘intruder’ caterpillar was introduced into the leaf tie. Caterpillars were observed pushing and hitting one another to gain or maintain access to the shelter. The occupant caterpillar maintained possession of the shelter in 52 % of the interactions, the intruder usurped the shelter from the occupant in 24 % of the interactions, and the caterpillars shared the shelter in 24 % of the interactions. The four species examined differed significantly in their behavior toward other caterpillars, and ordered most to least aggressive are P. reflexella, P. cryptolechiella, P. quercicella, and Pse. quercinigracella. There were also behavioral differences within and between species when acting as an occupant or an intruder. Psilocorsis cryptolechiella was more aggressive and Pse. quercinigracella was less aggressive when defending a shelter than when attempting to usurp a shelter. It was also found that the four focal caterpillar species, when collected from naturally colonized leaf ties, co-occur significantly less often than expected when compared to a null model. It appears that direct competition for shelters that serve as a territory as well as a food source is frequent and influences the composition of the leaf-tying caterpillar community.


Leaf-tying caterpillar leaf shelter direct competition territoriality physically aggressive behavior 



Many thanks go to John T. Lill for guidance throughout this research. Thanks to all members of the Lill lab for assistance in caterpillar collection. Thank you DC Plant-Insect Group (DC PIG) and anonymous reviewers for helpful comments. Funding provided by GWU Mortensen Fund, GWU Harlan Trust, and Washington Biologist Field Club Research Award.

Supplementary material

10905_2015_9511_MOESM1_ESM.pdf (1.8 mb)
ESM 1 (PDF 1881 kb)
10905_2015_9511_MOESM2_ESM.pdf (41 kb)
Supplemental Video 2 Video shows representative examples of the behaviors and outcomes of physically aggressive encounters between leaf-tying caterpillars. Caterpillars pushed by placing their head against the other caterpillar and contracting body segments to walk forward. They hit each other by rapidly moving the head laterally. Trials ended with caterpillars sharing the shelter (both residing in shelter with no further aggressive interactions), occupant caterpillar ‘winning’ (maintaining possession of the shelter), or intruder caterpillar ‘winning’ (usurping the shelter). (PDF 40 kb)

(MPG 8330 kb)


  1. Abarca M, Boege K (2011) Fitness costs and benefits of shelter building and leaf trenching behaviour in a pyralid caterpillar. Ecol Entomol 36:564–573. doi: 10.1111/j.1365-2311.2011.01299.x CrossRefGoogle Scholar
  2. Atlegrim O (1992) Mechanisms regulating bird predation on a herbivorous larva guild in boreal coniferous forests. Ecography 15:19–24CrossRefGoogle Scholar
  3. Berenbaum MR, Green ES, Zangerl AR (1993) Web costs and web defense in the parsnip webworm (Lepidoptera: Oecophoridae). Environ Entomol 22:791–795CrossRefGoogle Scholar
  4. Bowen JL, Mahony SJ, Mason AC, Yack JE (2008) Vibration-mediated territoriality in the warty birch caterpillar Drepana bilineata. Physiol Entomol 33:238–250. doi: 10.1111/j.1365-3032.2008.00627.x CrossRefGoogle Scholar
  5. Cappuccino N (1993) Mutual use of leaf-shelters by lepidopteran larvae on paper birch. Ecol Entomol 18:287–292CrossRefGoogle Scholar
  6. Carroll MR, Kearby WH (1978) Microlepidopterous oak leaftiers (Lepidoptera:Gelechioidea) in central Missouri. J Kans Entomol Soc 51:457–471Google Scholar
  7. Chesson P (2000) Mechanisms of maintenance of species diversity. Annu Rev Ecol Syst 31:343. doi: 10.1146/annurev.ecolsys.31.1.343 CrossRefGoogle Scholar
  8. Damman H (1987) Leaf quality and enemy avoidance by the larvae of a pyralid moth. Ecology 68:88–97CrossRefGoogle Scholar
  9. Denno RF, McClure MS, Ott JR (1995) Interspecific interactions in phytophagous insects - competition re-examined and resurrected. Annu Rev Entomol 40:297–331. doi: 10.1146/annurev.ento.40.1.297 CrossRefGoogle Scholar
  10. Fletcher LE, Yack JE, Fitzgerald TD, Hoy RR (2000) Vibrational communication in the cherry leaf roller caterpillar Caloptilia serotinella (Gracillarioidea : Gracillariidae). J Insect Behav 19:1–18. doi: 10.1007/s10905-005-9007-y CrossRefGoogle Scholar
  11. Fox J (2003) Effect displays in R for generalised linear models. J Stat Softw 8:1–27CrossRefGoogle Scholar
  12. Fox J, Weisberg S (2011) An {R} companion to applied regression, 2nd edn. Sage, Thousand OaksGoogle Scholar
  13. Fukui A, Murakami M, Konno K, Nakamura M, Ohgushi T (2002) A leaf-rolling caterpillar improves leaf quality. Entomol Sci 5:263–266Google Scholar
  14. Gotelli NJ (2000) Null model analysis of species co-occurrence patterns. Ecology 81:2606–2621CrossRefGoogle Scholar
  15. Gotelli NJ, Entsminger GL (2011) EcoSim: null models software for ecology. 7 edn. Acquired Intelligence Inc. & Kesey-Bear, Jericho, VT 05465Google Scholar
  16. Hairston N, Smith F, Slobodkin L (1960) Community structure, population control and competition. Am Nat 94:421–425CrossRefGoogle Scholar
  17. Hunter MD, Willmer PG (1989) The potential for interspecific competition between 2 abundant defoliators on oak-leaf damage and habitat quality. Ecol Entomol 14:267–277CrossRefGoogle Scholar
  18. Janzen DH (1973) Host plants as islands. II. Competition in evolutionary and contemporary time. Am Nat 107:786–790. doi: 10.1086/282876 CrossRefGoogle Scholar
  19. Jones MT, Castellanos I, Weiss MR (2002) Do leaf shelters always protect caterpillars from invertebrate predators? Ecol Entomol 27:753–757. doi: 10.1046/j.1365-2311.2002.00465.x CrossRefGoogle Scholar
  20. Joos B, Casey TM, Fitzgerald TD, Buttemer WA (1988) Roles of the tent in behavioral thermoregulation of eastern tent caterpillars. Ecology 69:2004–2011CrossRefGoogle Scholar
  21. Kaplan I, Denno RF (2007) Interspecific interactions in phytophagous insects revisited: a quantitative assessment of competition theory. Ecol Lett 10:977–994. doi: 10.1111/j.1461-0248.2007.01093.x CrossRefPubMedGoogle Scholar
  22. Lill JT (2001) Selection on herbivore life-history traits by the first and third trophic levels: the devil and the deep blue sea revisited. Evolution 55:2236–2247CrossRefPubMedGoogle Scholar
  23. Lill JT, Marquis RJ (2003) Ecosystem engineering by caterpillars increases insect herbivore diversity on white oak. Ecology 84:682–690CrossRefGoogle Scholar
  24. Lill JT, Marquis RJ (2004) Leaf ties as colonization sites for forest arthropods: an experimental study. Ecol Entomol 29:300–308CrossRefGoogle Scholar
  25. Lill JT, Marquis RJ, Walker MA, Peterson L (2007) Ecological consequences of shelter sharing by leaf-tying caterpillars. Entomol Exp Appl 124:45–53. doi: 10.1111/j.1570-7458.2007.00546.x CrossRefGoogle Scholar
  26. Loeffler CC (1996) Adaptive trade-offs of leaf folding in Dichomeris caterpillars on goldenrods. Ecol Entomol 21:34–40CrossRefGoogle Scholar
  27. LoPresti EF, Morse DH (2013) Costly leaf shelters protect moth pupae from parasitoids. Arthropod Plant Interact 7:445–453. doi: 10.1007/s11829-013-9261-4 CrossRefGoogle Scholar
  28. Marquis RJ, Lill JT (2007) Effects of arthropods as physical ecosystem engineers on plant-based trophic interaction webs. In: Ohgushi T, Craig TP, Price PW (eds) Ecological communities: plant mediation in indirect interaction webs. Cambridge University Press, New York, pp 246–274CrossRefGoogle Scholar
  29. Marquis RJ, Lill JT, Piccinni A (2002) Effect of plant architecture on colonization and damage by leaftying caterpillars of Quercus alba. Oikos 99:531–537CrossRefGoogle Scholar
  30. Martinsen GD, Floate KD, Waltz AM, Wimp GM, Whitham TG (2000) Positive interactions between leafrollers and other arthropods enhance biodiversity on hybrid cottonwoods. Oecologia 123:82–89CrossRefGoogle Scholar
  31. Morin PJ (1999) Community ecology. Blackwell Publishing, MaldenGoogle Scholar
  32. Nakamura M, Ohgushi T (2003) Positive and negative effects of leaf shelters on herbivorous insects: linking multiple herbivore species on a willow. Oecologia 136:445–449. doi: 10.1007/s00442-003-1285-5 CrossRefPubMedGoogle Scholar
  33. Okuda T (1989) Aggressive characteristics of diapausing larvae of a stem borer, Busseola fusca Fuller (Lepidoptera: Noctuidae) in artificially crowded conditions. Appl Entomol Zool 24:238–239Google Scholar
  34. Rathcke BJ (1976) Competition and coexistence within a guild of herbivorous insects. Ecology 57:76–87. doi: 10.2307/1936399 CrossRefGoogle Scholar
  35. Robson DS (1972) Appendix: statistical tests of significance. J Theor Biol 34:350–352Google Scholar
  36. Sagers CL (1992) Manipulation of host plant-quality - herbivores keep leaves in the dark. Funct Ecol 6:741–743CrossRefGoogle Scholar
  37. Sandberg SL, Berenbaum MR (1989) Leaf-tying by tortricid larvae as an adaptation for feeding on phototoxic Hypercium perforatum. J Chem Ecol 15:875–885CrossRefPubMedGoogle Scholar
  38. Schluter D (1984) A variance test for detecting species associations, with some example applications. Ecology 65:998–1005CrossRefGoogle Scholar
  39. Scott JL, Yack JE (2012) Vibratory territorial signals in caterpillars of the poplar lutestring, Tethea or (Lepidoptera: Drepanidae). Eur J Entomol 109:411–417CrossRefGoogle Scholar
  40. Scott JL, Kawahara AY, Skevington JH, Yen SH, Sami A, Smith ML, Yack JE (2010a) The evolutionary origins of ritualized acoustic signals in caterpillars. Nat Commun 1:9. doi: 10.1038/ncomms1002 CrossRefGoogle Scholar
  41. Scott JL, Matheson SM, Yack JE (2010b) Variation on a theme: vibrational signaling in caterpillars of the rose hook-tip moth, Oreta rosea. J Insect Sci 10:15CrossRefGoogle Scholar
  42. Sigmon E, Lill JT (2013) Phenological variation in the composition of a temperate forest leaf tie community. Environ Entomol 42:29–37CrossRefPubMedGoogle Scholar
  43. Sipura M (1999) Tritrophic interactions: willows, herbivorous insects and insectivorous birds. Oecologia 121:537–545CrossRefGoogle Scholar
  44. Sliwinski M, Sigmon E (2013) Why do leaf-tying caterpillars abandon their leaf ties? PeerJ 1:e173PubMedCentralCrossRefPubMedGoogle Scholar
  45. Stiling PD, Strong DR (1984) Experimental density manipulation of stem-boring insects - some evidence for interspecific competition. Ecology 65:1683–1685. doi: 10.2307/1939147 CrossRefGoogle Scholar
  46. Stone L, Roberts A (1990) The checkerboard score and species distributions. Oecologia 85:74–79CrossRefGoogle Scholar
  47. Tack AJM, Ovaskainen O, Harrison PJ, Roslin T (2009) Competition as a structuring force in leaf miner communities. Oikos 118:809–818. doi: 10.1111/j.1600-0706.2008.17397.x CrossRefGoogle Scholar
  48. Team RC (2012) R: a language and environment for statistical computing. R Foundation for Statistical Computing, ViennaGoogle Scholar
  49. Tilman D (1982) Resource competition and community structure. Princeton University Press, PrincetonGoogle Scholar
  50. Wang HG, Marquis RJ, Baer CS (2012) Both host plant and ecosystem engineer identity influence leaf-tie impacts on the arthropod community of Quercus. Ecology 93:2186–2197CrossRefPubMedGoogle Scholar
  51. Weiss MR (2003) Good housekeeping: why do shelter-dwelling caterpillars fling their frass? Ecol Lett 6:361–370. doi: 10.1046/j.1461-0248.2003.00442.x CrossRefGoogle Scholar
  52. Weiss MR, Wilson EE, Castellanos I (2004) Predatory wasps learn to overcome the shelter defences of their larval prey. Anim Behav 68:45–54. doi: 10.1016/j.anbehav.2003.07.010 CrossRefGoogle Scholar
  53. Willmer PG (1980) The effects of fluctuating environment on the water relations of larval Lepidoptera. Ecol Entomol 5:271–292CrossRefGoogle Scholar
  54. Yack JE, Smith ML, Weatherhead PJ (2001) Caterpillar talk: acoustically mediated territoriality in larval Lepidoptera. Proc Natl Acad Sci U S A 98:11371–11375PubMedCentralCrossRefPubMedGoogle Scholar

Copyright information

© Springer Science+Business Media New York 2015

Authors and Affiliations

  1. 1.Department of Biological SciencesGeorge Washington UniversityWashingtonUSA

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