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• W2009812036 abstract "Most bat species breed communally, but how their colonies are founded is a mystery. A recent study suggests that the formation of a new colony starts with related females splitting off from an existing colony. Most bat species breed communally, but how their colonies are founded is a mystery. A recent study suggests that the formation of a new colony starts with related females splitting off from an existing colony. How social animals form new groups strongly affects the stability of the groups as well as the reproductive success and the social behaviour of the group members [1Reeve H.K. Keller L. Partitioning of reproduction in mother-daughter versus sibling associations: a test of the optimal skew theory.Am. Nat. 1995; 145: 119-132Crossref Scopus (113) Google Scholar, 2Langer P. Hogendoorn K. Keller L. Tug-of-war over reproduction in a social bee.Nature. 2004; 428: 844-847Crossref PubMed Scopus (79) Google Scholar]. For this reason, group formation has been studied intensively, particularly in social insects and primates [1Reeve H.K. Keller L. Partitioning of reproduction in mother-daughter versus sibling associations: a test of the optimal skew theory.Am. Nat. 1995; 145: 119-132Crossref Scopus (113) Google Scholar, 2Langer P. Hogendoorn K. Keller L. Tug-of-war over reproduction in a social bee.Nature. 2004; 428: 844-847Crossref PubMed Scopus (79) Google Scholar, 3Van Horn R.C. Buchan J.C. Altmann J. Alberts S.C. Divided destinies: group choice by female savannah baboons during social group fission.Behav. Ecol. Sociobiol. 2007; 61: 1823-1837Crossref Scopus (50) Google Scholar]. Bats are highly gregarious mammals and in most species the females form colonies to raise their offspring communally [4Kerth G. Causes and consequences of sociality in bats.BioScience. 2008; 58 (in press)Crossref PubMed Scopus (188) Google Scholar]. Exactly how new colonies are formed is mysterious, however, as colony foundation is a rare and elusive event. Until recently, the available evidence was limited to population genetic data, which suggested that bat colonies are founded by groups of females that split off from an existing colony and then settle nearby [5Kerth G. Petit E. Colonization and dispersal in a social species, the Bechstein's bat (Myotis bechsteinii).Mol. Ecol. 2005; 14: 3943-3950Crossref PubMed Scopus (47) Google Scholar]. Now a paper by Metheny et al. [6Metheny J.D. Kalcounis-Rueppell M.C. Bondo K.J. Brigham R.M. A genetic analysis of group movement in an isolated population of tree-roosting bats.Proc. R. Soc. B. 2008; 275: 2265-2272Crossref PubMed Scopus (37) Google Scholar] on big brown bats (Eptesicus fuscus; Figure 1) presents the first detailed field observations of joint movements of individually marked females to a new area, in combination with genetic data on their relatedness. The study reveals that colony formation is initiated by a group of females splitting off from the original colony. The whole colony-fission process occurs over a period of four years. Intriguingly, it also shows that these founding females are closely related to one another, resulting in a higher average relatedness in the new than in the old colony. As with many other forest-living bats, big brown bat colonies almost daily switch their day roosts (tree cavities) and split into subgroups that later fuse again [6Metheny J.D. Kalcounis-Rueppell M.C. Bondo K.J. Brigham R.M. A genetic analysis of group movement in an isolated population of tree-roosting bats.Proc. R. Soc. B. 2008; 275: 2265-2272Crossref PubMed Scopus (37) Google Scholar, 7Metheny J.D. Kalcounis-Rueppell M.C. Willis C.K.R. Kolar K.A. Brigham R.M. Genetic relationships between roost-mates in a fission–fusion society of tree-roosting big brown bats (Eptesicus fuscus).Behav. Ecol. Sociobiol. 2008; 62: 1043-1051Crossref Scopus (50) Google Scholar]. Such fission–fusion behaviour is widespread among bats and probably allows females to adjust daily group sizes to changing environmental conditions, such as ambient temperature and parasite loads, while maintaining the social relationships among them [4Kerth G. Causes and consequences of sociality in bats.BioScience. 2008; 58 (in press)Crossref PubMed Scopus (188) Google Scholar]. Several studies have applied association indices and network analysis to quantify individual associations in bat species with fission–fusion behaviour [7Metheny J.D. Kalcounis-Rueppell M.C. Willis C.K.R. Kolar K.A. Brigham R.M. Genetic relationships between roost-mates in a fission–fusion society of tree-roosting big brown bats (Eptesicus fuscus).Behav. Ecol. Sociobiol. 2008; 62: 1043-1051Crossref Scopus (50) Google Scholar, 8Kerth G. König B. Fission, fusion and nonrandom associations in female Bechstein's bats (Myotis bechsteinii).Behaviour. 1999; 136: 1187-1202Crossref Scopus (211) Google Scholar, 9O'Donnell C.F.J. Cryptic local populations in a temperate rainforest bat Chalinolobus tuberculatus in New Zealand.Anim. Conserv. 2000; 3: 287-297Crossref Google Scholar, 10Vonhof M.J. Whitehead H. Fenton M.B. Analysis of Spix's disc-winged bat association patterns and roosting home ranges reveal a novel social structure among bats.Anim. Behav. 2004; 68: 507-521Crossref Scopus (87) Google Scholar, 11Rhodes M. Wardell-Johnson G.W. Rhodes M.P. Raymond B. Applying network analysis to the conservation of habitat tress in urban environments: a case study from Brisbane, Australia.Cons. Biol. 2006; 20: 861-870Crossref PubMed Scopus (62) Google Scholar, 12Popa-Lisseanu A.G. Bontadina F. Mora O. Ibáñez C. Highly structured fission-fusion societies in an aerial-hawking, carnivorous bat.Anim. Behav. 2008; 75: 471-482Crossref Scopus (89) Google Scholar]: all of them found non-random roosting associations despite regularly changing subgroup compositions. In Bechstein's bats (Myotis bechsteinii) and big brown bats, the only two bat species with fission–fusion behaviour for which genetic data are available, related females do not roost preferentially together [7Metheny J.D. Kalcounis-Rueppell M.C. Willis C.K.R. Kolar K.A. Brigham R.M. Genetic relationships between roost-mates in a fission–fusion society of tree-roosting big brown bats (Eptesicus fuscus).Behav. Ecol. Sociobiol. 2008; 62: 1043-1051Crossref Scopus (50) Google Scholar, 8Kerth G. König B. Fission, fusion and nonrandom associations in female Bechstein's bats (Myotis bechsteinii).Behaviour. 1999; 136: 1187-1202Crossref Scopus (211) Google Scholar]. This fits well with the observation that kinship in bats is often only of secondary importance for cooperative behaviours, such as information transfer about roosts, which allows colony members to coordinate their roost switching [4Kerth G. Causes and consequences of sociality in bats.BioScience. 2008; 58 (in press)Crossref PubMed Scopus (188) Google Scholar, 13Kerth G. Ebert C. Schmidtke C. Group decision-making in fission-fusion societies: Evidence from two field experiments in Bechstein's bats.Proc. R. Soc. B. 2006; 273: 2785-2790Crossref PubMed Scopus (116) Google Scholar]. If kinship does not affect daily roosting associations, why then does it matter when females disperse together to start a new colony? The study by Metheny et al. [6Metheny J.D. Kalcounis-Rueppell M.C. Bondo K.J. Brigham R.M. A genetic analysis of group movement in an isolated population of tree-roosting bats.Proc. R. Soc. B. 2008; 275: 2265-2272Crossref PubMed Scopus (37) Google Scholar] does not answer this question, but at least two explanations seem plausible. The first involves cooperation among related colony members, as suggested by Metheny et al. [6Metheny J.D. Kalcounis-Rueppell M.C. Bondo K.J. Brigham R.M. A genetic analysis of group movement in an isolated population of tree-roosting bats.Proc. R. Soc. B. 2008; 275: 2265-2272Crossref PubMed Scopus (37) Google Scholar] for big brown bats and by other researchers for social insects and primates [1Reeve H.K. Keller L. Partitioning of reproduction in mother-daughter versus sibling associations: a test of the optimal skew theory.Am. Nat. 1995; 145: 119-132Crossref Scopus (113) Google Scholar, 2Langer P. Hogendoorn K. Keller L. Tug-of-war over reproduction in a social bee.Nature. 2004; 428: 844-847Crossref PubMed Scopus (79) Google Scholar, 3Van Horn R.C. Buchan J.C. Altmann J. Alberts S.C. Divided destinies: group choice by female savannah baboons during social group fission.Behav. Ecol. Sociobiol. 2007; 61: 1823-1837Crossref Scopus (50) Google Scholar]. Kin selection might favour dispersal with relatives over dispersal with non-relatives if, after the formation of a new bat colony, cooperative behaviours benefiting the colony as a whole are more costly for the acting individuals than in old colonies. One cooperative behaviour to which this might apply is social warming of foreign pups [4Kerth G. Causes and consequences of sociality in bats.BioScience. 2008; 58 (in press)Crossref PubMed Scopus (188) Google Scholar]. If new colonies form by fission, they will be smaller than the old ones from which they split off. Social warming in large colonies may come as a cheap by-product, as many lactating females have to stay in the roost anyway to nurse their pups. In small colonies, however, the additional presence of non-lactating females may be required to warm the pups sufficiently. Non-lactating females profit less from staying in a warm place than pregnant or lactating females and their pups, so if they are unrelated they might actually prefer to roost on their own and save energy by going into torpor [8Kerth G. König B. Fission, fusion and nonrandom associations in female Bechstein's bats (Myotis bechsteinii).Behaviour. 1999; 136: 1187-1202Crossref Scopus (211) Google Scholar]. In colonies founded by related females, in contrast, the non-lactating females can gain indirect benefits from contributing to altruistic warming that helps related pups to grow faster and survive better. A more proximate explanation is that relatives are more likely to found colonies simply because they are more likely to share alleles that lead to increased exploratory and dispersal behaviour. Dispersal morphs and ‘explorative’ personalities have been reported in many animals, and there is some evidence that they have a genetic basis [14Wolf M. van Doorn G.S. Leimar O. Weissing F.J. Life-history trade-offs favour the evolution of animal personalities.Nature. 2007; 447: 581-584Crossref PubMed Scopus (848) Google Scholar, 15Sinervo B. Calsbeek R. Comendant T. Both C. Adamopoulou C. Clobert J. Genetic and maternal determinants of effective dispersal: the effect of sire genotype and size at birth in side-blotched lizards.Am. Nat. 2006; 168: 88-99Crossref PubMed Scopus (70) Google Scholar, 16Dingemanse N.J. Both C. van Noordwijk A.J. Rutten A.L. Drent P.J. Natal dispersal and personalities in great tits (Parus major).Proc. R. Soc. B. 2003; 270: 741-747Crossref PubMed Scopus (425) Google Scholar]. If explorative or dispersal behaviour is heritable in bats, relatives that share such traits via common descent should be more likely to found colonies together even if no fitness benefit results from increased kinship in new colonies. To discriminate between the possible explanations for colony founders being relatives, more studies are needed that evaluate the costs and benefits of living together with kin. In other taxa, differences in the way colonies are founded have been attributed to ecological factors, such as environmental conditions, and life-history traits, such as reproductive rate, that influence the likelihood of a new colony being established successfully [1Reeve H.K. Keller L. Partitioning of reproduction in mother-daughter versus sibling associations: a test of the optimal skew theory.Am. Nat. 1995; 145: 119-132Crossref Scopus (113) Google Scholar, 2Langer P. Hogendoorn K. Keller L. Tug-of-war over reproduction in a social bee.Nature. 2004; 428: 844-847Crossref PubMed Scopus (79) Google Scholar]. Experimental manipulations of the individual composition of newly founded colonies, as performed in social insects [2Langer P. Hogendoorn K. Keller L. Tug-of-war over reproduction in a social bee.Nature. 2004; 428: 844-847Crossref PubMed Scopus (79) Google Scholar], are not possible in wild bats. Instead, studies of colony foundation in bats need to be comparative, documenting the social behaviour and fitness of individual colony members as well as the fate of entire colonies under varying ecological and social conditions, including roost availability, climate, colony size, and relatedness among founding individuals. Studying the costs and benefits of living together with kin is a challenging task in bats that requires long-term studies combining pedigree analyses with field observations and experimental manipulations of behaviour [13Kerth G. Ebert C. Schmidtke C. Group decision-making in fission-fusion societies: Evidence from two field experiments in Bechstein's bats.Proc. R. Soc. B. 2006; 273: 2785-2790Crossref PubMed Scopus (116) Google Scholar, 17Kruuk L.E.B. Hill W.G. Introduction. Evolutionary dynamics of wild populations: the use of long-term pedigree data.Proc. R. Soc. B. 2008; 275: 593-596Crossref PubMed Scopus (55) Google Scholar]. Fortunately, such studies are now feasible, thanks to the development of polymorphic molecular markers, new field techniques such as automatic monitoring of individually tagged bats, and several powerful analytical tools for assessing individual associations [18Krause J. Croft D.P. James R. Social network theory in the behavioural sciences: potential applications.Behav. Ecol. Sociobiol. 2007; 62: 15-27Crossref Scopus (284) Google Scholar, 19Whitehead H. Precision and power in the analysis of social structure using associations.Anim. Behav. 2008; 75: 1093-1099Crossref Scopus (98) Google Scholar, 20Kunz T.H. Parsons S. Ecological and Behavioral Methods for the Study of Bats. John Hopkins University Press, Baltimore2009Google Scholar]. With more such studies being initiated, we can look forward to further fascinating insights into the importance of kinship for social behaviour and colony foundation in bats, and as a consequence in all social animals." @default.
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• W2009812036 title "Animal Sociality: Bat Colonies Are Founded by Relatives" @default.
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