Benefits to satellite members in mixed-species foraging groups: an experimental analysis

Hypotheses proposed to explain the formation of mixed-species foraging groups have focused on both foraging and antipredation benefits. Mixed-species flocks of bark-foraging birds form during the winter in the eastern deciduous forests of North America. These flocks are composed of two parid nuclear species, tufted titmice, Baeolophus bicolor, and either Carolina or black-capped chickadees, Poecile carolinensis or P. atricapillus, and several satellite species including downy woodpeckers, Picoides pubescens, and white-breasted nuthatches, Sitta carolinensis. The parid nuclear species seem to act as flock leaders and are closely followed by the satellite species. To elucidate what advantages downy woodpeckers and white-breasted nuthatches gain by flocking with parids, we removed parids from eight Ohio woodlots isolated by surrounding agricultural fields and compared the woodpeckers and nuthatches in these woodlots to those in eight controls. We tested four predictions generated by group-foraging hypotheses: compared with controls, satellite birds in treatment woodlots should (1) forage more in microclimates that reduce metabolic costs, (2) increase their vigilance, (3) exhibit reduced nutritional condition and (4) exhibit higher mortality rates. As predicted, female downy woodpeckers in treatment woodlots tended to forage in locations that were more sheltered from wind, presumably thereby reducing metabolic costs. Treatment males and females of both species significantly increased their vigilance. Finally, in the absence of parids, male nuthatches showed significantly reduced nutritional condition according to ptilochronology analysis of feathers grown during the experimental manipulation, and tended to exhibit increased mortality Copyright 1998 The Association for the Study of Animal Behaviour