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Cooperative Breeding Theory and an Evaluation of the Methodological Approaches to Understanding the Evolution and Maintenance of Sociality

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Cooperative Breeding Theory and an Evaluation of the Methodological Approaches to Understanding the Evolution and Maintenance of Sociality University of Wollongong Research Online Faculty of Science, Medicine and Health - Papers: part A Faculty of Science, Medicine and Health January 2017 The right tools for the job: Cooperative breeding theory and an evaluation of the methodological approaches to understanding the evolution and maintenance of sociality Martin Hing University of Wollongong, [email protected] Oya S. Klanten University of Technology Sydney, [email protected] Mark P. Dowton University of Wollongong, [email protected] Marian Y. L Wong University of Wollongong, [email protected] Follow this and additional works at: https://ro.uow.edu.au/smhpapers Recommended Citation Hing, Martin; Klanten, Oya S.; Dowton, Mark P.; and Wong, Marian Y. L, "The right tools for the job: Cooperative breeding theory and an evaluation of the methodological approaches to understanding the evolution and maintenance of sociality" (2017). Faculty of Science, Medicine and Health - Papers: part A. 5040. https://ro.uow.edu.au/smhpapers/5040 Research Online is the open access institutional repository for the University of Wollongong. For further information contact the UOW Library: [email protected] The right tools for the job: Cooperative breeding theory and an evaluation of the methodological approaches to understanding the evolution and maintenance of sociality Abstract Why do we observe so many examples in nature in which individuals routinely delay or completely forgo their own reproductive opportunities in order to join and remain within a group? Cooperative breeding theory provides a rich framework with which to study the factors that may influence the costs and benefits of emainingr philopatric as a non-breeder. This is often viewed as an initial step in the development of costly helping behavior provided by non-breeding subordinates. Despite many excellent empirical studies testing key concepts of the theory, there is still debate regarding the relative importance of various evolutionary forces, suggesting that there may not be a general explanation but rather a dynamic and taxonomically varied combination of factors influencing the ve olution and maintenance of sociality. Here, we explore two potential improvements in t he study of sociality that could aid in the progress of this field. The first addresses the fact that empirical studies of social evolution are typically conducted using either comparative, observational or manipulative methodologies. Instead, we suggest a holistic approach, whereby observational and experimental studies are designed with the explicit view of advancing comparative analyses of sociality for the taxon, and in tandem, where comparative work informs targeted research effort on specific (usually understudied) species within the lineage. A second improvement relates to the broadening of tests of cooperative breeding theory to include taxa where subordinates do not necessarily provide active cooperation within the group. The original bias toward "helpful subordinates" arose from a focus on terrestrial taxa. However, recent consideration of other taxa, especially marine taxa, is slowly revealing that the theory can and should encompass a continuum of cooperative social systems, including those where subordinates do not actively help. This review summarizes the major hypotheses of cooperative breeding theory, one of the dominant frameworks to examine social evolution, and highlights the potential benefits that a combined methodological approach and a broader application could provide to the study of sociality. Publication Details Hing, M. L., Klanten, O. Selma., Dowton, M. & Wong, M. Y.L. (2017). The right tools for the job: Cooperative breeding theory and an evaluation of the methodological approaches to understanding the evolution and maintenance of sociality. Frontiers in Ecology and Evolution, 5 100-1-100-19. This journal article is available at Research Online: https://ro.uow.edu.au/smhpapers/5040 REVIEW published: 31 August 2017 doi: 10.3389/fevo.2017.00100 The Right Tools for the Job: Cooperative Breeding Theory and an Evaluation of the Methodological Approaches to Understanding the Evolution and Maintenance of Sociality Martin L. Hing 1*, O. Selma Klanten 2, Mark Dowton 3 and Marian Y. L. Wong 1* 1 Centre for Sustainable Ecosystems Solutions, School of Biological Sciences, University of Wollongong, Wollongong, NSW, Edited by: Australia, 2 Fish Ecology Laboratory, School of the Life Sciences, University of Technology Sydney, Sydney, NSW, Australia, Trine Bilde, 3 Centre for Medical and Molecular Bioscience, School of Biological Sciences, University of Wollongong, Wollongong, NSW, Aarhus University, Denmark Australia Reviewed by: Michael Taborsky, Why do we observe so many examples in nature in which individuals routinely delay University of Bern, Switzerland or completely forgo their own reproductive opportunities in order to join and remain Kelly Anne Stiver, Southern Connecticut State within a group? Cooperative breeding theory provides a rich framework with which to University, United States study the factors that may influence the costs and benefits of remaining philopatric as a *Correspondence: non-breeder. This is often viewed as an initial step in the development of costly helping Martin L. Hing [email protected] behavior provided by non-breeding subordinates. Despite many excellent empirical Marian Y. L. Wong studies testing key concepts of the theory, there is still debate regarding the relative [email protected] importance of various evolutionary forces, suggesting that there may not be a general Specialty section: explanation but rather a dynamic and taxonomically varied combination of factors This article was submitted to influencing the evolution and maintenance of sociality. Here, we explore two potential Social Evolution, improvements in the study of sociality that could aid in the progress of this field. The first a section of the journal Frontiers in Ecology and Evolution addresses the fact that empirical studies of social evolution are typically conducted using Received: 23 October 2016 either comparative, observational or manipulative methodologies. Instead, we suggest Accepted: 10 August 2017 a holistic approach, whereby observational and experimental studies are designed Published: 31 August 2017 with the explicit view of advancing comparative analyses of sociality for the taxon, Citation: Hing ML, Klanten OS, Dowton M and and in tandem, where comparative work informs targeted research effort on specific Wong MYL (2017) The Right Tools for (usually understudied) species within the lineage. A second improvement relates to the the Job: Cooperative Breeding Theory broadening of tests of cooperative breeding theory to include taxa where subordinates and an Evaluation of the Methodological Approaches to do not necessarily provide active cooperation within the group. The original bias Understanding the Evolution and toward “helpful subordinates” arose from a focus on terrestrial taxa. However, recent Maintenance of Sociality. Front. Ecol. Evol. 5:100. consideration of other taxa, especially marine taxa, is slowly revealing that the theory doi: 10.3389/fevo.2017.00100 can and should encompass a continuum of cooperative social systems, including those Frontiers in Ecology and Evolution | www.frontiersin.org 1 August 2017 | Volume 5 | Article 100 Hing et al. Methodological Approaches to Study Sociality where subordinates do not actively help. This review summarizes the major hypotheses of cooperative breeding theory, one of the dominant frameworks to examine social evolution, and highlights the potential benefits that a combined methodological approach and a broader application could provide to the study of sociality. Keywords: cooperative breeding theory, evolution of sociality, methodology, kinship, ecological constraints, benefits of philopatry, life-history hypothesis, fish INTRODUCTION subordinates in groups, whether or not they actively cooperate must weigh the costs and benefits of group membership. It The animal kingdom contains many examples of species, is these costs and benefits that the hypotheses that make up including our own, which form surprisingly complex social cooperative breeding theory focus on. Thus, studies investigating structures (Munday et al., 1998; Purcell, 2011; Grueter et al., 2012; the determinants of group living need not be restricted to Chapais, 2013; Johnson et al., 2013; Taborsky and Wong, 2017). applying cooperative breeding theory only to species where The size, structure and composition of these groups can vary both helping actively occurs. within and between species, from pair-bonding monogamous Notwithstanding the excellent empirical and theoretical work partners (Kleiman, 2011; Servedio et al., 2013) to large and highly conducted in this field (e.g., Emlen, 1994; Cockburn, 1996; complex societies exhibiting social hierarchies and division of Arnold and Owens, 1998; Hatchwell and Komdeur, 2000; Pen labor (Duffy and Macdonald, 2010; Nandi et al., 2013). Such and Weissing, 2000; Buston and Balshine, 2007), the relative variation in social structure is intriguing as it suggests that there importance of the evolutionary forces at play which influence may be a great diversity of underlying social, ecological or life the decision of non-breeders to forego their own reproductive history factors that influence the evolution of stable groups and opportunities and remain within a group are still the subject their maintenance over many generations. of much discussion. Advances in understanding
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