Gerald Kerth

Fission, fusion and nonrandom associations in female Bechstein's bats (Myotis bechsteinii

Maternity colonies of the Bechsteins bat (Myotis bechsteinii) are socially closed units that frequently split into subgroups which occupy different roosts. We analysed the social structure of one colony over three years in the field and quantified associations among pairs of females by using three association indices. Colony members exhibited marked mixing, although individual composition of subgroups was not completely random. Females associated according to the reproductive status, with lactating females preferentially roosting together. Relatedness, determined from 5 nuclear and 1 mitochondrial microsatellite, had no consistent influence on the degree of association. In combination, the indices allowed for the interpretation that nonrandom associations occur even in the absence of shared roost and group size preferences. High individual associations among reproducing female Bechsteins bats might reflect the importance of cooperation in maternity colonies. The frequent fission and fusion of subgroups suggests a flexible reaction of Bechsteins bats according to enviromental conditions.

Causes and Consequences of Sociality in Bats

ABSTRACT Bats are among the most diverse and most gregarious of all mammals. This makes them highly interesting for research on the causes and consequences of sociality in animals. Detailed studies on bat sociality are rare, however, when compared with the information available for other social mammals, such as primates, carnivores, ungulates, and rodents. Modern field technologies and new molecular methods are now providing opportunities to study aspects of bat biology that were previously inaccessible. Consequently, bat social systems are emerging as far more complex than had been imagined. Variable dispersal patterns, complex olfactory and acoustic communication, flexible context-related interactions, striking cooperative behaviors, and cryptic colony structures in the form of fission-fusion systems have been documented. Bat research can contribute to the understanding of animal sociality, and specifically to important topics in behavioral ecology and evolutionary biology, such as dispersal, fission-fusion behavior, group decisionmaking, and cooperation.

Mitochondrial DNA (mtDNA) reveals that female Bechstein's bats live in closed societies.

We present a microgeographic analysis of mitochondrial DNA (mtDNA) in Bechstein’s bats using three sources of control region sequence variability, including a novel mtDNA microsatellite, to assess individual relatedness both within and among 10 maternity colonies. Comparison of marker variability among 268 adult females revealed little genetic variability within each colony. However, most colonies were clearly distinguished by colony‐specific mitochondrial haplotypes (total n = 28). Low intracolony variability and strong haplotype segregation among colonies, was reflected by an extraordinary high FST of 0.68, indicating a very low intercolony dispersal rate of approximately one female in five generations. Haplotype distribution among 18 solitary males showed that males frequently disperse between colony locations, indicating the absence of dispersal barriers. Bechstein’s bat maternity colonies are thus closed groups that comprise 20–40 females probably belonging to only one or, at most, two matrilines. The genetic population structure of Bechstein’s bats is in agreement with the hypothesis that females seek familiar and, at least, partially related cooperation partners for raising their young. Alternatively strong philopatry might reflect the importance of profound roost or habitat knowledge for successful reproduction in female Bechstein’s bats.

Extreme sex-biased dispersal in the communally breeding, nonmigratory Bechstein's bat (Myotis bechsteinii).

Maternity colonies of the communally breeding, nonmigratory Bechsteins bat consist of closely related females that live together with unrelated females, and average colony relatedness is low despite the absence of immigration. We compared the genetic structure of both nuclear and mitochondrial microsatellites in order to quantify sex‐specific dispersal rates. More specifically, we aimed at testing whether male dispersal is able to balance the genetic effect of strong (absolute) female philopatry. Absolute female philopatry, indicated by an extreme mitochondrial DNA population differentiation of 96%, was indeed opposed by strong (possibly complete) male dispersal. Based on our knowledge of the biology of Myotis bechsteinii, we conclude that inbreeding avoidance is likely to be the crucial factor driving male dispersal in this species.

Bats are able to maintain long-term social relationships despite the high fission–fusion dynamics of their groups

Elephants, dolphins, as well as some carnivores and primates maintain social links despite their frequent splitting and merging in groups of variable composition, a phenomenon known as fission–fusion. Information on the dynamics of social links and interactions among individuals is of high importance to the understanding of the evolution of animal sociality, including that of humans. However, detailed long-term data on such dynamics in wild mammals with fully known demography and kin structures are scarce. Applying a weighted network analysis on 20 500 individual roosting observations over 5 years, we show that in two wild Bechsteins bat colonies with high fission–fusion dynamics, individuals of different age, size, reproductive status and relatedness maintain long-term social relationships. In the larger colony, we detected two stable subunits, each comprising bats from several family lineages. Links between these subunits were mainly maintained by older bats and persisted over all years. Moreover, we show that the full details of the social structure become apparent only when large datasets are used. The stable multi-level social structures in Bechsteins bat colonies resemble that of elephants, dolphins and some primates. Our findings thus may shed new light on the link between social complexity and social cognition in mammals.

Information transfer about roosts in female Bechstein's bats: an experimental field study.

Information transfer among group members is believed to play an important part in the evolution of coloniality in both birds and bats. Although information transfer has received much scientific interest, field studies using experiments to test the underlying hypotheses are rare. We used a field experiment to test if communally breeding female Bechsteins bats (Myotis bechsteinii) exchange information regarding novel roosts. We supplied a wild colony, comprising 17 adult females of known relatedness, with pairs of suitable and unsuitable roosts and monitored the arrival of individuals marked with transponders (PIT-tags) over 2 years. As expected with information transfer, significantly more naive females were recruited towards suitable than towards unsuitable roosts. We conclude that information transfer about roosts has two functions: (i) it generates communal knowledge of a large set of roosts; and (ii) it aids avoidance of colony fission during roost switching. Both functions seem important in Bechsteins bats, in which colonies depend on many day roosts and where colony members live together for many years.

Group decision making in fission-fusion societies : Evidence from two-field experiments in Bechstein's bats

Group decisions are required when group coordination is beneficial, but individuals can choose between alternatives. Despite the increased interest in animal group decision making, there is a lack of experimental field studies that investigate how animals with conflicting information make group decisions. In particular, no field studies have considered the influence of fission–fusion behaviour (temporary splitting into subgroups) on group decisions. We studied group decision making in two wild Bechsteins bat colonies, which are fission–fusion societies of stable individual composition. Since they frequently switch communal roosts, colony members must regularly make group decisions over where to roost. In the two-field experiments, we provided marked individuals with conflicting information about the suitability of potential roosts. We investigated whether conflicting information led to group decisions that followed a ‘unanimous’ or a ‘majority’ rule, or increased colony fission. Individual behaviour suggests that bats considered both their own information and the behaviour of others when deciding where to roost. Group decisions about communal roosts reflected the information available to a majority of the bats roosting together, but conflicting information led to an increased fission in one colony. Our results suggest that fission–fusion societies allow individuals to avoid majority decisions that are not in their favour.

Secretions of the interaural gland contain information about individuality and colony membership in the Bechstein's bat

Mammals use chemical signals for individual and kin recognition, to establish social hierarchies, mark territories and choose mates. The nocturnal and social lifestyle of bats suggests that, besides acoustic signals, they also use scent to communicate. We investigated in the communally breeding Bechstein’s bat, Myotis bechsteinii, whether secretions of the facial interaural gland contain information that can be used for individual and colony recognition. Since female Bechstein’s bats live in closed societies and show cooperative behaviour, we predicted they would recognize colony members. We analysed interaural gland secretions, which we repeatedly sampled from 85 females belonging to four free-ranging colonies. Gas chromatography/mass spectrometry profiles were individually specific and differed between colonies. Comparing odour profiles between colonies we found a relation between chemical similarity and the mitochondrial haplotype of colony members. Within colonies there was no correlation between mass spectrometer profile similarity and genetic relatedness. Our results suggest that female Bechstein’s bats may use interaural gland secretions for individual and colony recognition but not to infer kinship directly.

High gene diversity at swarming sites suggest hot spots for gene flow in the endangered Bechstein's bat

Our study shows that endangered Bechsteinsbats utilise distinct habitats at differentstages of their reproductive cycle, a findingthat has implications how habitat should beselected for preservation. Using nuclear andmitochondrial microsatellite DNA markers wecompared gene diversity of Bechsteins batswithin breeding colonies and at potentialmating sites. Bechsteins bat is one of themost threatened European bat species. Duringsummer it depends largely on mature deciduousforests. Females exhibit strict natalphilopatry. They form demographicallyindependent breeding colonies comprisingmaternally closely related bats. Males aresolitary. Like other temperate bats,Bechsteins bats swarm at the end of summer infront of caves. Because the sexes meet there,such swarming sites are potentially importantfor gene flow. Our genetic analyses reveal thatswarming sites have greater mitochondrial DNAgene diversity than colonies. Furthermore,field data on the phenology and reproductivestatus of several hundred individuals suggestthat Bechsteins bats mate during swarming. Incombination our field and genetic data showthat swarming sites provide the opportunity forgene flow among bats originating from differentcolonies. Therefore, we suggest that swarmingsites should be strictly protected to maintainthe observed high levels of gene flow amongcolonies.

Roost selection and roost switching of female Bechstein's bats (Myotis bechsteinii) as a strategy of parasite avoidance

Ectoparasites of vertebrates often spend part of their life cycle in their hosts’ home. Consequently, hosts should take into account the parasite infestation of a site when selecting where to live. In a field study, we investigated whether colonial female Bechstein’s bats (Myotis bechsteinii) adapt their roosting behaviour to the life cycle of the bat fly Basilia nana in order to decrease their contact with infective stages of this parasite. B. nana imagoes live permanently on the bat’s body but deposit puparia in the bat’s roosts. The flies metamorphose independently in the roosts, but after metamorphosis emerge only in the presence of a potential host. In a field experiment, the bats preferred non-contagious to contagious day-roosts and hence were able to detect either the parasite load of roosts or some correlate with infestation, such as bat droppings. In addition, 9 years of observational data on the natural roosting behaviour of female Bechstein’s bats indicate that the bats largely avoid re-occupying roosts when highly contagious puparia are likely to be present as a result of previous occupations of the roosts by the bat colony. Our results indicate that the females adapted their roosting behaviour to the age-dependent contagiousness (emergence probability) of the puparia. However, some infested roosts were re-occupied, which we assume was because these roosts provided advantages to the bats (e.g. a beneficial microclimate) that outweighed the negative effects associated with bat fly infestation. We suggest that roost selection in Bechstein’s bats is the outcome of a trade-off between the costs of parasite infestation and beneficial roost qualities.