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Waiting Time as a New Component in Functional Response Models

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For more content, see Archives Néerlandaises de Zoologie (Vol 1-17) and Animal Biology (Vol 53 and onwards).

(1) The functional response of the three-spined stickleback to changes in prey density was determined by presenting individual sticklebacks (for 15 seconds) with different densities of Daphnia. (2) The observed response, at first sight a type II, could not be adequately described by the existing type II models, i.e. Holling's "disc equation" and Rogers's "random predator equation". (3) The above mentioned models assume that the predator is unable to search for new prey while handling. This is claimed not to be true in the case of the stickleback-Daphnia interaction. (4) A modification of the random predator equation has been developed, which is based upon a biological explanation of the predation process, and gives an adequate description of the data. Two components of handling are distinguished: external and internal handling. Searching can again take place during the internal handling time. From this the possibility arises that the predator finds new prey before the handling of the previous prey has finished. This would result in a "waiting time". (5) The new model has been compared with a model in which the attack-rate increases as prey density increases, which also gives a good fit to the data. Analysis of video-tape recordings of the feeding of sticklebacks shows however, that handling time in this "variable attack rate model" is not in accordance with the actual behaviour. (6) Similarities between the feeding behaviour of the sticklebacks and that of mites and damselflies have been discussed. The "waiting time" resembles the "bottleneck-effect" found in the feeding behaviour of the latter two.

Affiliations: 1: Animal Ecology Research Group, Department of Zoology, South Parks Road, Oxford; 2: Theory Division, Rutherford Laboratory, Chilton, Didcot, Oxon

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