Population Ecology

, Volume 54, Issue 2, pp 239–250 | Cite as

Elevational gradient in the cyclicity of a forest-defoliating insect

  • Kyle J. HaynesEmail author
  • Andrew M. Liebhold
  • Derek M. Johnson
Original article


Observed changes in the cyclicity of herbivore populations along latitudinal gradients and the hypothesis that shifts in the importance of generalist versus specialist predators explain such gradients has long been a matter of intense interest. In contrast, elevational gradients in population cyclicity are largely unexplored. We quantified the cyclicity of gypsy moth populations along an elevational gradient by applying wavelet analysis to spatially referenced 31-year records (1975–2005) of defoliation. Based on geographically weighted regression and nonlinear regression, we found either a hump-shaped or plateauing relationship between elevation and the cyclicity of gypsy moth populations and a positive relationship between cyclicity and the density of the gypsy moth’s preferred host-tree species. The potential effects of elevational gradients in the density of generalist predators and preferred host-tree species on the cyclicity of gypsy moth populations were evaluated with mechanistic simulation models. The models suggested that an elevational gradient in the densities of preferred host tree species could partially explain elevational patterns of gypsy moth cyclicity. Results from a model assuming a type-III functional response of generalist predators to changes in gypsy moth density were inconsistent with the observed elevational gradient in gypsy moth cyclicity. However, a model with a more realistic type-II functional response gave results roughly consistent with the empirical findings. In contrast to classical studies on the effects of generalist predators on prey population cycles, our model with a type-II functional response predicts a unimodal relationship between generalist-predator density and the cyclicity of gypsy moth populations.


Gypsy moth Lymantria dispar Peromyscus leucopus Population cycle Periodicity 



Gino Luzader provided valuable assistance with the gypsy moth defoliation database. Jonathan Walter provided useful comments on an earlier draft of this publication. Funding for this project was provided by a USDA-NRI Grant (2006-35306-17264) to D.M. Johnson.

Supplementary material

10144_2012_305_MOESM1_ESM.pdf (99 kb)
Supplementary material 1 (PDF 99 kb)


  1. Berryman AA (1991) The gypsy moth in North America: a case of successful biological control? Trends Ecol Evol 6:110–111CrossRefGoogle Scholar
  2. Bess HAS, Spurr H, Littlefield EW (1947) Forest site conditions and the gypsy moth. Harvard forest bulletin no. 22. Harvard University Press, CambridgeGoogle Scholar
  3. Bjørnstad ON, Wilhelm F, Stenseth NC (1995) Geographic gradient in small rodent density fluctuations: a statistical modeling approach. Proc R Soc B 262:127–133PubMedCrossRefGoogle Scholar
  4. Bjørnstad ON, Robinet C, Liebhold AM (2010) Geographic variation in North-American gypsy moth population cycles: sub-harmonics, generalist predators and spatial coupling. Ecology 91:106–118PubMedCrossRefGoogle Scholar
  5. Brooks RT, Smith HR, Healy WM (1998) Small-mammal abundance at three elevations on a mountain in central Vermont, USA: a sixteen-year record. For Ecol Manage 110:181–193CrossRefGoogle Scholar
  6. Brown JH (2001) Mammals on mountainsides: elevational patterns of diversity. Global Ecol Biogeogr 10:101–109CrossRefGoogle Scholar
  7. Brundson CV, Fotheringham AS, Charlton ME (1996) Geographically weighted regression: a method for exploring spatial nonstationarity. Geogr Anal 28:281–298Google Scholar
  8. Burnham KP, Anderson DR (2002) Model selection and multimodel inference: a practical information—theoretic approach. Springer, New YorkGoogle Scholar
  9. Campbell RW (1981) Population dynamics. In: Doane CC, McManus ML (eds) The gypsy moth: research toward integrated pest management. Technical bulletin 1584, USDA Forest Service, Washington, DC, pp 65–216Google Scholar
  10. Cattadori IM, Hudson PJ (1999) Temporal dynamics of grouse populations at the southern edge of their distribution. Ecography 22:374–383CrossRefGoogle Scholar
  11. Christenson LM, Lovett GM, Mitchell MJ, Groffmanet PM (2002) The fate of nitrogen in gypsy moth frass deposited to an oak forest floor. Oecologia 131:444–452CrossRefGoogle Scholar
  12. Ciesla WM (2000) Remote sensing in forest health protection. Forest Health Technology Enterprise Team and Remote Sensing Applications Center report no. 00-03. USDA Forest Service, Remote Sensing Applications Center, Salt Lake CityGoogle Scholar
  13. Coulson TP, Rohani P, Pascual M (2004) Skeletons, noise and population growth: the end of an old debate? Trends Ecol Evol 19:359–364PubMedCrossRefGoogle Scholar
  14. Davidson CB, Johnson JE, Gottschalk KW, Amateis RL (2001) Prediction of stand susceptibility and gypsy moth defoliation in Coastal Plain mixed pine-hardwoods. Can J For Res 31:1914–1921Google Scholar
  15. Dwyer G, Elkinton JS (1993) Using simple models to predict virus epizootics in gypsy moth populations. J Anim Ecol 62:1–11CrossRefGoogle Scholar
  16. Dwyer G, Elkinton JS, Buonaccorsi JP (1997) Host heterogeneity in susceptibility and disease dynamics: tests of a mathematical model. Am Nat 150:685–707PubMedCrossRefGoogle Scholar
  17. Dwyer G, Dushoff J, Elkinton JS, Levin SA (2000) Pathogen-driven outbreaks in forest defoliators revisited: building models from experimental data. Am Nat 156:105–120PubMedCrossRefGoogle Scholar
  18. Dwyer G, Dushoff J, Yee SH (2004) The combined effects of pathogens and predators on insect outbreaks. Nature 430:341–345PubMedCrossRefGoogle Scholar
  19. Elassal AAC, Caruso VM (1983) Digital elevation models. Geological Survey Circular 895-B. US Geological Survey, RestonGoogle Scholar
  20. Elkinton JS, Liebhold AM (1990) Population dynamics of gypsy moth in North America. Ann Rev Entomol 35:571–596CrossRefGoogle Scholar
  21. Elkinton JS, Gould JR, Liebhold AM, Smith HR, Wallner WE (1989) Are gypsy moth populations regulated at low density? In: Wallner WE, McManus KA (eds) Lymantriidae: a comparison of features of new and old world tussock moths. General technical report NE-123. USDA Forest Service, Northeastern Forest Experiment Station, Broomall, pp 233–249Google Scholar
  22. Elkinton JS, Healy WM, Buonaccorsi JP, Boettner GH, Hazzard A, Liebhold AM, Smith HR (1996) Interactions among gypsy moths, white-footed mice, and acorns. Ecology 77:2332–2342CrossRefGoogle Scholar
  23. Elkinton JS, Liebhold AM, Muzika R (2004) Effects of alternative prey on predation by small mammals on gypsy moth pupae. Popul Ecol 46:171–178CrossRefGoogle Scholar
  24. Elton C (1924) Periodic fluctuations in the number of animals: their causes and effects. Brit J Exp Biol 2:119–163Google Scholar
  25. Erelli MC, Ayres MP, Eaton GK (1998) Altitudinal patterns in host suitability for forest insects. Oecologia 117:133–142CrossRefGoogle Scholar
  26. Farge M (1992) Wavelet transforms and their applications to turbulence. Ann Rev Fluid Mech 24:395–457CrossRefGoogle Scholar
  27. Fotheringham AS, Brundson C, Charlton M (2002) Geographically weighted regression: the analysis of spatially varying relationships. Wiley, West SussexGoogle Scholar
  28. Gaston KJ, Williams PH (1996) Spatial patterns in taxonomic diversity. In: Gaston KJ (ed) Biodiversity: a biology of numbers and difference. Blackwell Science, Oxford, pp 77–113Google Scholar
  29. Gray DR, Ravlin FW, Braine JA (2001) Diapause in the gypsy moth: a model of inhibition and development. J Insect Physiol 47:173–184PubMedCrossRefGoogle Scholar
  30. Grenfell BT, Bjørnstad ON, Kappey J (2001) Travelling waves and spatial hierarchies in measles epidemics. Nature 414:716–723PubMedCrossRefGoogle Scholar
  31. Grushecky ST, Liebhold AM, Greer R, Smith RL (1998) Does forest thinning affect predation on gypsy moth (Lepidoptera: Lymantriidae) larvae and pupae? Environ Entomol 27:268–276Google Scholar
  32. Hajek AE, Humber R, Elkinton JS (1995) The mysterious origin of Entomophaga maimaiga in North America. Am Entomol 41:31–42Google Scholar
  33. Hamilton DJ, Lechowicz MJ (1991) Host effects on the development and fecundity of gypsy-moth, Lymantria dispar, reared under field conditions. Can J Zool 69:2217–2224CrossRefGoogle Scholar
  34. Hanski I, Hansson L, Henttonen H (1991) Specialist predators, generalist predators, and the microtine rodent cycle. J Anim Ecol 60:353–367CrossRefGoogle Scholar
  35. Hansson L (1987) An interpretation of rodent dynamics as due to trophic interactions. Oikos 50:308–318CrossRefGoogle Scholar
  36. Haynes KJ, Liebhold AM, Johnson DM (2009) Spatial analysis of harmonic oscillation of gypsy moth outbreak intensity. Oecologia 159:249–256PubMedCrossRefGoogle Scholar
  37. Holling CS (1959) Some characteristics of simple types of predation and parasitism. Can Entomol 91:385–398CrossRefGoogle Scholar
  38. Holling CS (1965) The functional response of predators to prey density and its role in mimicry and population regulation. Mem Ent Soc Can 45:1–60CrossRefGoogle Scholar
  39. Houston DR, Valentine HT (1977) Comparing and predicting forest stand susceptibility to gypsy moth. Can J For Res 7:447–461CrossRefGoogle Scholar
  40. Hunter AF, Elkinton JS (2000) Effects of synchrony with host plant on populations of a spring-feeding lepidopteran. Ecology 81:1248–1261CrossRefGoogle Scholar
  41. Hutchinson MF, Gessler PE (1994) Splines—more than just a smooth interpolator. Geoderma 62:45–67CrossRefGoogle Scholar
  42. Ims RA, Henden J, Killengreen ST (2008) Collapsing population cycles. Trends Ecol Evol 23:79–86PubMedCrossRefGoogle Scholar
  43. Johnson DR, Swanson BJ, Eger JL (2000) Cyclic dynamics of eastern Canadian ermine populations. Can J Zool 78:835–839CrossRefGoogle Scholar
  44. Johnson DM, Liebhold AM, Bjørnstad ON (2006) Geographical variation in the periodicity of gypsy moth outbreaks. Ecography 29:367–374CrossRefGoogle Scholar
  45. Keith LB, Cary JR, Rongstad OJ, Brittingham MC (1984) Demography and ecology of a declining snowshoe hare population. Wildl Monogr 90:1–43Google Scholar
  46. Kendal BE, Prendergast J, Bjørnstad ON (1998) The macroecology of population dynamics: taxonomic and biogeographic patterns in population cycles. Ecol Lett 1:160–164CrossRefGoogle Scholar
  47. Kingsley NP (1985) A forester’s atlas of the northeast. General technical report NE-95. USDA Forest Service, Northeastern Forest Experiment Station, BroomallGoogle Scholar
  48. Klemola T, Tanhuanpää M, Korpimäki E, Ruohomäki K (2002) Specialist and generalist natural enemies as an explanation for geographical gradients in population cycles of northern herbivores. Oikos 99:83–94CrossRefGoogle Scholar
  49. Körner C, Basler D (2010) Phenology under global warming. Science 327:1461–1462PubMedCrossRefGoogle Scholar
  50. Lambin X, Bretagnolle V, Yoccoz N (2006) Vole population cycles in Northern and Southern Europe: is there a need for different explanations for a single pattern? J Anim Ecol 75:340–349PubMedCrossRefGoogle Scholar
  51. Liebhold AM, Bascompte J (2003) The Allee effect, stochastic dynamics and the eradication of alien species. Ecol Lett 6:133–140CrossRefGoogle Scholar
  52. Liebhold AM, Halverson JA, Elmes GA (1992) Gypsy moth invasion in North America: a quantitative analysis. J Biogeogr 19:513–520CrossRefGoogle Scholar
  53. Liebhold AM, Elmes GA, Halverson JA, Quimby J (1994) Landscape characterization of forest susceptibility to gypsy-moth defoliation. For Sci 40:18–29Google Scholar
  54. Liebhold AM, Gottschalk KW, Muzika R, Montgomery ME, Young R, O’Day K, Kelley B (1995) Suitability of North American tree species to the gypsy moth: a summary of field and laboratory tests. General technical report NE-221. USDA Forest Service, Northeastern Forest Experiment Station, RadnorGoogle Scholar
  55. Liebhold AM, Gottschalk KW, Mason DA, Bush RR (1997) Forest susceptibility to the gypsy moth. J For 95:20–24Google Scholar
  56. Liebhold AM, Elkinton J, Williams D, Muzika R (2000) What causes outbreaks of the gypsy moth in North America? Popul Ecol 42:257–266CrossRefGoogle Scholar
  57. Liu YG, Liang XS, Weisberg RH (2007) Rectification of the bias in the wavelet power spectrum. J Atmos Ocean Tech 24:2093–2102CrossRefGoogle Scholar
  58. Moran PAP (1949) The statistical analysis of the sunspot and lynx cycles. J Anim Ecol 18:115–116CrossRefGoogle Scholar
  59. Morin RS, Liebhold AM, Luzader ER, Lister AJ, Gottschalk KW, Twardus DB (2005) Mapping host-species abundance of three major exotic forest pests. Research paper NE-726. USDA Forest Service, Northeastern Research Station, Newtown SquareGoogle Scholar
  60. Murdoch WW (1973) The functional response of predators. J Appl Ecol 10:335–341Google Scholar
  61. Murdoch WW, Oaten A (1975) Predation and population stability. Adv Ecol Res 9:1–131CrossRefGoogle Scholar
  62. Murray DL (2000) A geographic analysis of snowshoe hare population demography. Can J Zool 78:1207–1217CrossRefGoogle Scholar
  63. Peltonen M, Liebhold AM, Bjørnstad ON, Williams DW (2002) Spatial synchrony in forest insect outbreaks: roles of regional stochasticity and dispersal. Ecology 83:3120–3129CrossRefGoogle Scholar
  64. Rahbek C (1995) The elevational gradient of species richness: a uniform pattern. Ecography 18:200–205CrossRefGoogle Scholar
  65. Rogers DJ (1972) Random search and insect population models. J Anim Ecol 41:360–383CrossRefGoogle Scholar
  66. Royama T (1992) Analytical population dynamics. Chapman and Hall, LondonCrossRefGoogle Scholar
  67. Sardeshmukh PD, Compo GP, Penland C (2000) Changes of probability associated with El Niño. J Clim 13:4268–4286CrossRefGoogle Scholar
  68. Schauber EM (2001) Models of mast seeding and its ecological effects on gypsy moth populations and Lyme disease risk. Ph.D. dissertation. The University of Connecticut, StorrsGoogle Scholar
  69. Schauber EM, Ostfeld RS, Jones CG (2004) Type 3 functional response of mice to gypsy moth pupae: is it stabilizing? Oikos 107:592–602CrossRefGoogle Scholar
  70. Schenk D, Bacher S (2002) Functional response of a generalist insect predator to one of its prey species in the field. J Anim Ecol 71:524–531CrossRefGoogle Scholar
  71. Sergio F, Pedrini P (2007) Biodiversity gradients in the Alps: the overriding importance of elevation. Biodivers Conserv 16:3243–3254CrossRefGoogle Scholar
  72. Smith HR (1983) Wildlife and the gypsy moth. Trans Northeast Fish Wildl Conf 40:66Google Scholar
  73. Smith HR (1985) Wildlife and the gypsy moth. Wildl Soc Bull 13:166–174Google Scholar
  74. Smith HR (1989) Predation: its influence on population dynamics and adaptive in morphology and behavior of the Lymantriidae. In: Wallner NE, McManus KA (eds) The Lymantriidae: a comparison of features of new and old world tussock moths. General technical report NE-123. USDA Forest Service, Northeastern Forest Experiment Station, Broomall, pp 469–488Google Scholar
  75. Tobin PC, Whitmire SL, Johnson DM, Bjørnstad ON, Liebhold AM (2007) Invasion speed is affected by geographical variation in the strength of Allee effects. Ecol Lett 10:36–43PubMedCrossRefGoogle Scholar
  76. Torrence C, Compo GP (1998) A practical guide to wavelet analysis. Bull Am Meteorol Soc 79:61–78CrossRefGoogle Scholar
  77. Turchin P, Hanski I (1997) Empirically based model for latitudinal gradient in vole population dynamics. Am Nat 149:842–874PubMedCrossRefGoogle Scholar
  78. Whitmire SL, Tobin PC (2006) Persistence of invading gypsy moth populations in the United States. Oecologia 147:230–237PubMedCrossRefGoogle Scholar
  79. Wolff JO (1980) The role of habitat patchiness in the population dynamics of showshoe hares. Ecol Monogr 50:111–130CrossRefGoogle Scholar
  80. Yahner RH, Smith HR (1991) Small mammals abundance and habitat relationships on deciduous forested sites with different susceptibility to gypsy moth defoliation. Environ Manage 15:113–120CrossRefGoogle Scholar

Copyright information

© The Society of Population Ecology and Springer 2012

Authors and Affiliations

  • Kyle J. Haynes
    • 1
    Email author
  • Andrew M. Liebhold
    • 2
  • Derek M. Johnson
    • 3
  1. 1.The Blandy Experimental FarmUniversity of VirginiaBoyceUSA
  2. 2.USDA Forest ServiceNorthern Research StationMorgantownUSA
  3. 3.Department of BiologyVirginia Commonwealth UniversityRichmondUSA

Personalised recommendations