The importance of individual-to-society feedbacks in animal ecology and evolution

evolution 2 3 Mauricio Cantor1,2,3,4,5†, Adriana A. Maldonado-Chaparro1,2,3†, Kristina B. Beck6§, Hanja B. 4 Brandl1,2,3§, Gerald G. Carter1,7§, Peng He1,2,3§, Friederike Hillemann8§, James A. Klarevas5 Irby2,3,9§, Mina Ogino2,3§, Danai Papageorgiou1,2,3§, Lea Prox2,10,11§, Damien R. Farine1,2,3* 6 7 1Department of Collective Behaviour, Max Planck Institute of Animal Behavior, 78315 Radolfzell, 8 Germany. 9 2Department of Biology, University of Konstanz, Germany. 10 3Centre for the Advanced Study of Collective Behaviour, University of Konstanz, Germany. 11 4Departamento de Ecologia e Zoologia, Universidade Federal de Santa Catarina, Brazil 12 5Centro de Estudos do Mar, Universidade Federal do Paraná, Brazil 13 6Department of Behavioural Ecology and Evolutionary Genetics, Max Planck Institute for 14 Ornithology, 82319 Seewiesen, Germany 15 7Department of Evolution, Ecology and Organismal Biology, The Ohio State University, Columbus, 16 Ohio, USA. 17 8Edward Grey Institute of Field Ornithology, Department of Zoology, University of Oxford, UK 18 9Department of Migration, Max Planck Institute of Animal Behavior, Konstanz, Germany. 19 10Department of Sociobiology/Anthropology, Johann-Friedrich-Blumenbach Institute of Zoology & 20 Anthropology, University of Göttingen, Göttingen Germany. 21 11Behavioral Ecology & Sociobiology Unit, German Primate Center, Göttingen, Germany 22 †These authors contributed equally 23 §Alphabetical order 24 * Corresponding author: Damien R. Farine. Max Planck Institute of Animal Behavior. Am 25

6 considered. This synthesis provides a roadmap for strengthening existing, and arising, links 122 between currently disparate research topics (Fig. 2, Box 1), which we believe will help to 123 uncover new perspectives in the study of social evolutionary ecology.

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Evolutionary changes in host or pathogen states could feed back onto social structure. Theory 180 suggests that the relationship between pathogen prevalence, virulence, and host sociality is 181 complex and, sometimes, counter-intuitive (Prado et al. 2009

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When behaviours are socially-learned, shared within subgroups of the population, and persist 266 over time, they are recognized as culture (Laland & Hoppitt 2003

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Similarly, in spotted hyenas, the amount of social support, rather than intrinsic attributes (e.g.

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We highlight existing opportunities to integrate disparate areas of research (Fig. 2)

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Feedback loops between individuals and societies mean that the evolution of social structure 799 and the state of the individuals reciprocally influence each other (Fig. 1). In this review, we look

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We first identified 14 of the common and emerging research topics in the study of animal social 806 structures, and then quantified how many publications addressed each topic together and 807 separately (Fig. 2). Although the use of network methods in the field of animal behaviour dates  1). We then generated a network depicting edges as the co-occurrence of topics in the same 822 articles (Fig. 2).

824 38
The cumulative publication networks revealed a core of well-connected research topics along 825 with a set of more peripheral topics (Fig. 2). The most studied topics in the last 12 years include 826 bottom-up drivers of individual states (e.g. 'relatedness', 'dominance'), while the top-down 827 influences are usually underrepresented (e.g. 'social stability and resilience', 'early-life 828 conditions'). Well-known cross-disciplinary research is represented by strong links (e.g.

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'relatedness'-'cooperative behaviour', and 'mating systems'-'sexual and social selection').  though the number of articles more than doubled (Fig. 2). These patterns suggest some 835 branching out from the first to the second period, and greater focus on classical topics (e.g.

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'dominance', 'cooperative behaviour'), and somewhat less exploitation of new areas in the last 837 period. Some exceptions are 'cultural transmission' and 'social stability and resilience', whose 838 degrees increased from 0 to 7 and 1 to 4, respectively.

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Importantly, our analysis reveals that there are still under-represented and weakly-connected 841 topics, revealing promising areas for further cross-disciplinary research. Among the missing 842 links, we highlight (i) the completely disconnected topic 'predator-prey dynamics'; (ii) the 843 potential links from 'social inheritance' to 'dominance', to 'relatedness' and to 'information and 844 cultural transmission'; and (iii) all potential links among the topics 'early-life conditions',

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'dispersal', 'social stability' and 'physiological interactions and stress transmission'. Among the 846 weak links, we highlight (iv) those to and from 'social stability and resilience', (v) links from 847 'social inheritance' to 'dispersal', as well as (vi) links from 'pathogen and parasite transmission' 848 39 to 'dominance' and to 'physiological interactions'. We discuss these prominent areas for future 849 research in the "Future prospects for studying individual-to-society feedbacks" section. 850 851