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Functional response: rigorous estimation and sensitivity to genetic variation in prey


  • Thomas Tully,

  • Phillip Cassey,

  • Régis Ferrière

T. Tully, P. Cassey and R. Ferrière, Lab. d’Écologie, CNRS UMR 7625, École Normale Supérieure, 46 rue d'Ulm, FR-75005 Paris, France ( PC also at: School of Biosciences, Univ. of Birmingham, Edgbaston, UK, B15 2TT. RF also at: Dept of Ecology and Evolutionary Biology, Univ. of Arizona, Tucson, AZ 85721, USA.


Holling's type II functional response is a cornerstone of community ecology and coevolutionary theory. The so-called disc equation is the most widely used model of the type II response, yet thus far no robust experimental assessment has been achieved in any single system. Fundamental issues that remain to be assessed include whether the assumptions of the disc equation are fulfilled, whether the disc equation yields accurate estimates of predation-related individual traits, and whether differences in disc equation parameters can capture genetic variation in prey behaviour. This paper provides a rigorous approach to all of these questions. The functional response of the predatory mite Pergamasus crassipes on three genetically distinct clones of the springtail Folsomia candida was measured at six levels of prey density in controlled conditions where prey number and arena size were concomitantly manipulated. A crucial assumption of Holling's disc equation was fulfilled by maintaining a constant prey density for the entire experimental period of predation. The timing of each attack and capture, as well as the duration of the handling time, were recorded by constant observation. We contrasted three different methods to calculate functional response curves: (1) indirect estimation of the disc equation's parameters from the number of prey killed by the end of each experimental run; (2) direct estimation of the parameters via a unique protocol of constant observation; and (3) independently deriving a function based on direct measurements of encounter rate and attack success. The basic assumptions of the disk equation were globally fulfilled. Estimations of the functional response's parameters (type II) were remarkably congruent across approach (1) and (2). A single genetic effect was detected – the relationship between the encounter rate and prey density differed significantly between clones – whereas a direct comparison of functional response across clones failed to reveal genetic variation.

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