, Volume 161, Issue 2, pp 227–239 | Cite as

Relative importance of density-dependent regulation and environmental stochasticity for butterfly population dynamics

  • Piotr NowickiEmail author
  • Simona Bonelli
  • Francesca Barbero
  • Emilio Balletto
Population Ecology - Original Paper


The relative contribution of density-dependent regulation and environmental stochasticity to the temporal dynamics of animal populations is one of the central issues of ecology. In insects, the primary role of the latter factor, typically represented by weather patterns, is widely accepted. We have evaluated the impact of density dependence as well as density-independent factors, including weather and mowing regime, on annual fluctuations of butterfly populations. As model species, we used Maculinea alcon and M. teleius living in sympatry and, consequently, we also analysed the effect of their potential competition. Density dependence alone explained 62 and 42% of the variation in the year-to-year trends of M. alcon and M. teleius, respectively. The cumulative Akaike weight of models with density dependence, which can be interpreted as the probability that this factor should be contained in the most appropriate population dynamics model, exceeded 0.97 for both species. In contrast, the impacts of inter-specific competition, mowing regime and weather were much weaker, with their cumulative weights being in the range of 0.08–0.21; in addition, each of these factors explained only 2–5% of additional variation in Maculinea population trends. Our results provide strong evidence for density-dependent regulation in Maculinea, while the influence of environmental stochasticity is rather minor. In the light of several recent studies on other butterflies that detected significant density-dependent effects, it would appear that density-dependent regulation may be more widespread in this group than previously thought, while the role of environmental stochasticity has probably been overestimated. We suggest that this misconception is the result of deficiencies in the design of most butterfly population studies in the past, including (1) a strong focus on adults and a neglect of the larval stage in which density-dependent effects are most likely to occur; (2) an almost exclusive reliance on transect count results that may confound the impact of environmental stochasticity on butterfly numbers with its impact on adult longevity.


Inter-specific competition Maculinea Mark-recapture Mowing regime Weather 



Butterfly sampling was performed with the permission of the Italian Ministry for Environment, Territory and Sea, and was partly funded by the Parks Planning Department for the Piedmont Region. Silvia Ferretti, Andrea Crocetta, Camilla Colombo, Alessandra Rosso, Luca Pietro Casacci, Lisa Camerin and Marco Gherlenda helped in the fieldwork. The data analysis was conducted within the financial framework of the European Commission projects EuMon (FP6 contract no. 006463) and SCALES (FP7 contract no. 226852). We are grateful to Christine Richards for improving the English of the manuscript.

Supplementary material

442_2009_1373_MOESM1_ESM.pdf (107 kb)
Supplementary Table S1 (PDF 107 kb)


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Copyright information

© Springer-Verlag 2009

Authors and Affiliations

  • Piotr Nowicki
    • 1
    Email author
  • Simona Bonelli
    • 2
  • Francesca Barbero
    • 2
  • Emilio Balletto
    • 2
  1. 1.Institute of Environmental SciencesJagiellonian UniversityKrakówPoland
  2. 2.Department of Animal and Human BiologyUniversity of TurinTurinItaly

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