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Abstract

Abstract
Chapman et al.’s (Behav Ecol Sociobiol 36:59–70, 1995) ecological constraints model posits that the size and distribution of food patches place restrictions upon foraging group size. Larger groups incur increased travel costs for any given array of patches, and thus, to fulfil individual energetic and nutritional requirements, foragers should adjust group sizes to balance energy obtained against that spent on travelling. Support for this model comes from both comparative and species-specific studies but findings are contradictory, and the utility of the model has been questioned. This study provides a rigorous test, analysing measurements from distinct food patches and individual inter-patch movements, on an appropriately shorter temporal scale. Using data drawn from two social groups of a species characterised by a high degree of fission–fusion dynamics, the chimpanzee (Pan troglodytes), we show that larger parties foraged in larger food patches and for longer durations, and that larger parties were associated with further travel between patches. Overt contest competition over food increased with party size. We found no evidence of distinct sex differences in either party size or travel distances: the predictive power of forager sex was low compared to that of ecological variables. We propose that analysis at the patch level is more appropriate than a daily averaging approach that may smooth out the very variation being investigated. Our findings suggest that, despite certain limitations, Chapman et al.’s (Behav Ecol Sociobiol 36:59–70, 1995) model of ecological constraints remains a useful tool. Ecology does indeed constrain grouping patterns, and the impact of this is not necessarily differentiated by sex.

Significance statement
Foraging animals face the ‘more mouths to feed’ problem: as the numbers in a group increase, the group must travel further to find enough food, using up energy. Hence, foragers should adjust numbers to minimise these costs, but tests of this idea have proved inconclusive. We investigated the foraging behaviour of chimpanzees, a species with highly flexible grouping, considering their travel between specific patches of food. We found clear support for this proposition, with larger patches of food hosting larger numbers of foragers, and such groups having to travel further to find food. Although it is often thought that female animals should respond more strongly to foraging costs, we found little evidence of sex differences. Our results show that ecology does indeed constrain grouping patterns, and that the impact is felt equally by males and females.

Publication Type:	Article
Additional Information:	© 2022 the authors. This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article's Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article's Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creativecommons.org/licenses/by/4.0/.
Publisher Keywords:	Foraging, Patch size, Travel distance, Party size, Pan troglodytes, Chimpanzee
Subjects:	G Geography. Anthropology. Recreation > GF Human ecology. Anthropogeography G Geography. Anthropology. Recreation > GN Anthropology Q Science > QH Natural history Q Science > QH Natural history > QH301 Biology
Departments:	School of Health & Medical Sciences School of Health & Medical Sciences > Department of Psychology & Neuroscience
SWORD Depositor:	Symplectic Administrator

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