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The Ontogenetic Drivers of Behavioral Variation Across Groups

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The Ontogenetic Drivers of Behavioral Variation Across Groups REVIEW ARTICLE published: 10 December 2014 ECOLOGY AND EVOLUTION doi: 10.3389/fevo.2014.00081 The development of collective personality: the ontogenetic drivers of behavioral variation across groups Sarah E. Bengston 1* and Jennifer M. Jandt 2 1 Department of Ecology and Evolutionary Biology, University of Arizona, Tucson, Arizona, USA 2 Department of Ecology, Evolution and Organismal Biology, Iowa State University, Ames, Iowa, USA Edited by: For the past decade, the study of personality has become a topic on the frontier of Ann Valerie Hedrick, University of behavioral ecology. However, most studies have focused on exploring inter-individual California Davis, USA behavioral variation in solitary animals, and few account for the role that social interactions Reviewed by: may have on the development of an individual’s personality. Moreover, a social group Elva J. H. Robinson, University of York, UK may exhibit collective personality: an emergent behavioral phenotype displayed at the Marian Yi-ling Wong, University of group-level, which is not necessarily the sum or average of individual personalities within Wollongong, Australia that group. The social environment, in many cases, can determine group success, which *Correspondence: then influences the relative success of all the individuals in that group. In addition, Sarah E. Bengston, Department of group-level personality may itself evolve, subject to the same selection pressures as Ecology and Evolutionary Biology, University of Arizona, PO Box individual-level behavioral variation, when the group is a unit under selection. Therefore, 210088 Tucson, AZ 85721-0088, we reason that understanding how collective personalities emerge and change over USA time will be imperative to understanding individual- and group-level behavioral evolution. e-mail: [email protected] Personality is considered to be fixed over an individual’s lifetime. However, behavior may shift throughout development, particularly during adolescence. Therefore, juvenile behavior should not be compared with adult behavior when assessing personality. Similarly, as conditions within a group and/or the local environment can shift, group behavior may similarly fluctuate as it matures. We discuss potential within-group factors, such as group initiation, group maturation, genetic make-up of the group, and the internal social environment, and external factors, such as how local environment may play a role in generating group-level personalities. There are a variety of studies that explore group development or quantify group personality, but few that integrate both processes. Therefore, we conclude our review by discussing potential ways to evaluate development of collective personality, and propose several focal areas for future research. Keywords: collective behavior, social groups, social insects, within-group variation, group growth DEFINING AND EVALUATING COLLECTIVE PERSONALITIES Studies measuring the development of personality tend to Non-human personality, e.g., inter-individual differences in focus on unitary organisms and the role of early experience behavior that are consistent through time, has become an increas- (DiRienzo et al., 2012), maternal effects (Groothuis et al., 2008) inglypopularareaofstudyinthepastdecade(Koolhaas et al., and persistence across life stages (Bell and Stamps, 2004). It 1999; Sih et al., 2004; Stamps and Groothuis, 2010; Dall et al., has been shown that the consistency of personality can vary 2012, 2004). This may be in part because of the seemingly ubiq- through ontogeny and upon maturation. For example, in the uitous nature of animal personalities, which have been found in dumpling squid (Euprymna tasmanica), individuals are consistent nearly every taxon in which they have been investigated, as well as in their boldness as juveniles and as adults, but are inconsis- their profound effect on behavioral plasticity and long-term fit- tent for a period as they reach sexual maturity (Sinn et al., ness consequences (Sih et al., 2004; Smith and Blumstein, 2008; 2008). In yellow bellied marmots (Marmota flaviventris), bold- Stamps and Groothuis, 2010; Wray and Seeley, 2011; Jandt et al., ness was only predictable in yearlings, while docility was con- 2014a).Forexample,ingreattits(Parus major), there are inter- sistent across all age classes (Petelle et al., 2013). On the other individual differences in exploratory behavior (“fast” vs. “slow”; hand, lake frogs (Rana ridibunda; Wilson and Krause, 2012) Verbeek et al., 1994; Dingemanse et al., 2002). This individual and firebugs (Pyrrhocoris apterus; Gyuris et al., 2012) show variation is both highly repeatable through time, and heritable consistent personality (activity and exploration, boldness and in both wild caught and lab reared populations (Dingemanse exploration, respectively) across all life stages. The flexibility of et al., 2004, 2002). Moreover, these differences affect both adult some personality traits and the persistence of others may sig- and offspring survival, as fluctuating environmental conditions nal that some traits may be adaptively fixed, while others are drive selection for different personality types from year to year plastic so as to respond to age-specific situations (Petelle et al., (Dingemanse et al., 2004). 2013). www.frontiersin.org December 2014 | Volume 2 | Article 81 | 1 Bengston and Jandt The development of collective personality Personality differences can also be observed at the level of influence over the group’s behavior, then the growth rate of the group or colony, referred to here as “collective personality.” the group (and thus, the rate of new, inexperienced individuals While human social groups have been the primary focus of col- joining the group) may be particularly important during group lective personality studies (e.g., aggression or communication ontogeny, as it would alter the ratio of experienced to inexperi- differences among human groups; Duncan, 1999; Hofmann and enced individuals. Alternatively, if collective behavior is the result Jones, 2005; Halfhill et al., 2005), social animal groups are becom- of a decentralized process, where individuals respond to chang- ing more popular research models. Collective behavior is perhaps ing conditions based upon previously established rules (innate or best understood in eusocial insects (e.g., ants, bees, wasps, and learned) or fluctuating feedback systems, this may either lead to termites that exhibit cooperative brood care, overlapping adult more stabilized (if individual behaviors are buffered by the group; generations and division of labor between reproductive and non- Dussutour et al., 2009; Sasaki and Pratt, 2011) or destabilized (if reproductive castes; Hoölldobler and Wilson, 1990). In these positive feedback processes lead to exaggeration of small effects; systems, there is selection at the colony-level (Korb and Heinze, Sinha, 2006) collective personality. 2004). As such, colony development has been studied extensively. There are few studies that investigate collective personalities However, only recently has colony-level personality been consid- (e.g., Scharf et al., 2012);mostofwhichfocusonmeasuring ered, and little work on the development of collective personality mature groups (those capable of reproducing or budding into exists. In this review, to differentiate between eusocial and social new groups; e.g., Sasaki and Pratt, 2011; Wray and Seeley, 2011; species, we will use “colony personality” or “colony behavior” Sasaki et al., 2013). This is likely because measuring the devel- when referring to social insects, and “group personality/behavior” opment of a collective unit is more complicated than measuring when referring to non-eusocial species. the development of an individual: to explore developmental fac- Stamps and Groothuis (2010) suggest that studies measuring tors affecting individual personality, measurements can readily be the development of personality fall into three categories: (1) those repeated on juvenile and adult individuals to determine the effects that measure how specific, early experiences influence adult per- of experience or ontogeny on personality types (Bell and Stamps, sonality, (2) those that consider if and how personality changes 2004; Stamps and Groothuis, 2010). However, groups tend to throughout the lifetime of an organism, and (3) how genes influ- develop at a much slower pace than individuals, and collecting ence personality development. These questions can also be asked or monitoring multiple groups over extended periods of time can at the group-level: (1) How do events during group formation be difficult. shape later group- and individual-level behavior? (2) How does Here, we discuss potential within-group (group initiation, group-level behavior change across group ontogeny or as the group maturation, genetic make-up of the group, and internal group encounters different environmental and social circum- social environment) and local environmental factors that may stances? (3) How is group-level behavior affected by the genotypes play a role in the development of collective personality in ani- that comprise it? It is important to consider these questions of mal groups. We also highlight the importance of understanding development at both group- and individual-levels for a variety of a group’s developmental phase when assessing personality type. reasons. First, individuals and groups have different lifespans, and Finally, we propose several new avenues of research that will individuals within the group
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