Mean colony relatedness is a poor predictor of colony structure and female philopatry in the communally breeding Bechstein's bat (Myotis bechsteinii)

In order to understand why animals are social and how group members interact with each other it is important to know their relatedness. However, few studies have investigated the genealogy in complete social groups of free-living animals with low reproductive skew. This holds particularly true for bats. Although almost all bat species are social, their sociobiology is not well understood. Because they are volant, nocturnal and have a rather cryptic life-style, bats are difficult to observe in the wild. Furthermore females are generally gregarious making genetic parent-offspring assignment a challenging task. We used genetic markers in combination with knowledge about age and colony membership of individually marked bats to construct pedigrees in completely sampled maternity colonies of Bechstein's bats (Myotis bechsteinii). Despite considerable fluctuations in population size, no immigration occurred over 5 years in four colonies living in close proximity. Additionally, confrontation tests showed that females of one maternity colony were able to detect and attempted to prevent the intrusion of foreign females into a roost they occupy. Although colonies were absolutely closed, and 75% of the colony members lived together with close relatives (rS=0.25), mean colony relatedness was nearly zero (0.02). Average relatedness therefore is a poor estimator for the potential of kin selection in Bechstein's bat colonies and may be misleading when attempting to understand the social structure of animals living in groups where many members breed. Based on our results we discuss the potential adaptive value of living in closed societies with low reproductive skew.     

Low roost-site fidelity in pallid bats: associated factors and effect on group stability

Despite potential costs of changing roost or densites, many animals frequently move between roosts or dens. Pallid bats (Antrozous pallidus) change diurnal roost sites frequently and also reportedly have a variety of cooperative social behaviors, many of which are associated with the care of developing offspring. Roost switching is likely to increase the costs of maintaining the group stability expected with social cooperation. Pallid bats roosting in rock crevices in central Oregon were studied with radiotelemetry to (1) examine characteristics of day roosts, (2) determine what ecological factors were correlated with low roost fidelity, and (3) examine the temporal stability of roosting groups of pregnant and lactating bats. Pallid bats changed roosts an average of once every 1.4 days throughout the summer. The bats exhibited seasonal shifts in roost use, occupying roosts behind thin slabs of rock in cool weather and roosts in deep rock crevices in warm weather. Roost switching was not correlated with daily variations in weather conditions or with structural characteristics of the diurnal roosts, although switching may have allowed bats to maintain familiarity with several roosts that vary in microclimate. Roost switching was positively correlated with ectoparasite load. High ectoparasite levels were correlated with lower body weights in lactating females (Fig. 3), suggesting that parasites may be costly to the bats. Roost switching may be a strategy to decrease ectoparasite loads by interrupting the reproductive cycles of those parasites that spend at least part of their life cycle on the walls of the roost. Both pregnant and lactating pallid bats frequently changed their diurnal roost location, but lactating bats tended to travel shorter distances between consecutive roosts. Lactating bats were more likely to continue to associate with particular roostmates despite changes in the location of the diurnal roost (Fig. 4) and were less likely to roost alone. Although the stability of groups of lactating bats was not absolute, evidence supported the prediction that such groups are more cohesive than are those of pregnant bats. Received: 20 June 1995/Accepted after revision: 13 July 1996     

Look before you leap - individual variation in social vigilance shapes socio-spatial group properties in an agent-based model

Next to predator detection, primate vigilance also serves to keep track of relevant conspecifics. The degree of vigilance towards group members often reflects the dominance rank of an individual: subordinates pay attention to dominants. Although it has been suggested that subordinates' vigilance may result in spatial centrality of dominants, this has not been addressed in either empirical or modeling studies. Using agent-based models, we determined how social vigilance affects socio-spatial properties of primate groups. A basic model without social vigilance, where individuals avoid potential aggressors (avoidance model), was contrasted with two models that each additionally included a different type of social vigilance: a) monitoring a specific potential aggressor to remain informed on its whereabouts (monitoring model) or b) scanning the whole group to detect potential aggressors (scanning model). Adding monitoring or scanning behavior to the avoidance model reinforced spatial centrality of dominants, a pattern often observed in primates, and resulted in more spread out groups. Moreover, variation in scanning tendency alone was already sufficient to generate spatial centrality of dominants: frequently scanning subordinates could move further away from the group center than dominants, before losing sight of group members. In the monitoring model, two mechanisms caused decreased encounter frequencies among subordinates: a) increased inter-individual distances, and b) frequent monitoring of central dominants. In the scanning model, encounters among subordinates decreased due to increased inter-individual distances. This agent-based model study provides a clear indication that individual variation in social vigilance may be an important structuring feature of primate social groups.     

Small-scale spatial and temporal genetic structure of the Atlantic sea scallop (Placopecten magellanicus) in the inshore Gulf of Maine revealed using AFLPs

Despite long planktonic durations, many species of broadcast spawning invertebrates exhibit genetic structure at small spatial and temporal scales. Amplified fragment length polymorphisms were used to assess genetic variation in the sea scallop, Placopecten magellanicus, among four inshore and one offshore location in the Gulf of Maine and temporal genetic variation among age classes of sea scallops at one site. Our results indicated that genetic structure for P. magellanicus exists on smaller spatial scales (tens to hundreds of kilometers) than expected given the 40-day planktonic larval period. In addition, genetic differences among age classes may be influenced by inter-annual differences in larval supply or reproductive success. Future genetic studies should sample multiple age classes prior to comparison among locations.     

Timing of the evening emergence from day roosts of the grey-headed flying fox, Pteropus poliocephalus: the effects of predation risk, foraging needs, and social context

This study addresses the functional question of how variation in foraging strategy, predation risk, and social context influence the timing of the evening emergence from day roosts of the grey-headed flying fox, Pteropus poliocephalus. The onset of evening emergence was expected to vary according to the relative costs and benefits of emerging early and should, therefore, reflect an optimal trade-off between predation risks and foraging needs. The onset of the colony-wide emergence was closely correlated with the time of sunset and cloud cover. However, as expected, the onset of the colony-wide emergence was delayed when a diurnal avian predator was present, whereas the onset was advanced during lactation when presumably energetic demands are higher. The trade-off between predation risks and foraging needs was further reflected in the emergence times of individual bats: adult females emerged earlier when they had higher foraging needs as indicated by their body condition; young emerged later when they were smaller and likely to be more at risk from predation due to their less developed flying skills. However, the emergence time of adult males depended on their social status: smaller bachelor males emerged from the colony earlier than larger harem-holding males who guard their harems until the last female had left. Thus, whereas the colony-wide emergence time reflected the outcome of a trade-off between predation risks and general foraging needs, on an individual level, the outcome of this trade-off depended on sex, age, body condition, and structural size and was modified by social context.     

Do social networks of female northern long-eared bats vary with reproductive period and age?

Social structure, which is a function of the patterns of interactions among individuals, is particularly variable in fission–fusion societies. The underlying factors that drive this variation are poorly understood. Female northern long-eared bats (Myotis septentrionalis) live in fission–fusion societies where females form preferred associations within groups that vary daily in size and composition as individuals switch roosts. The goal of our study was to test the predictions that preferred associations and social networks of female northern long-eared bats vary with reproductive period and age. We also tested the prediction that preferred relationships persist across years despite movements from summer roosts to winter hibernacula. Network analyses revealed that during gestation, females roosted in smaller groups where they roosted more regularly with fewer individuals than during lactation. This variation may reflect different social strategies to mediate higher energetic costs during lactation. Females of all ages roosted more often with younger individuals, which in turn had more direct and indirect associations than all other age classes. Younger individuals may play a role in maintaining connections between individuals, perhaps as a result of younger individuals being more exploratory. Temporal analyses suggested that relationships can persist for years as some pairs roosted together for multiple summers. We suggest that the dynamic nature of fission–fusion societies is associated with individual strategies to increase fitness relative to individual characteristics, in this case reproductive condition and age.     

Seasonal changes in the structure of rhesus macaque social networks

Social structure emerges from the patterning of interactions between individuals and plays a critical role in shaping some of the main characteristics of animal populations. The topological features of social structure, such as the extent to which individuals interact in clusters, can influence many biologically important factors, including the persistence of cooperation, and the rate of spread of disease. Yet, the extent to which social structure topology fluctuates over relatively short periods of time in relation to social, demographic, or environmental events remains unclear. Here, we use social network analysis to examine seasonal changes in the topology of social structures that emerge from socio-positive associations in adult female rhesus macaques (Macaca mulatta). Behavioral data for two different association types (grooming and spatial proximity) were collected for females in two free-ranging groups during two seasons: the mating and birth seasons. Stronger dyadic bonds resulted in social structures that were more tightly connected (i.e., of greater density) in the mating season compared to the birth season. Social structures were also more centralized around a subset of individuals and more clustered in the mating season than those in the birth season, although the latter differences were mostly driven by differences in density alone. Our results suggest a degree of temporal variation in the topological features of social structure in this population. Such variation may feed back on interactions, hence affecting the behaviors of individuals, and may therefore be important to take into account in studies of animal behavior.     

Spatial variation in population dynamics in a colonial ascidian (Botryllus schlosseri)

Recent interest in the dynamics of marine invertebrate populations has focused largely on taxa with an open population structure. However, in many colonial taxa with limited larval dispersal, settlers may be locally derived. Consequently, dynamics may vary among sites that are separated by relatively short distances. This study explored spatial variation in temporal dynamics of colonial ascidians (Botryllus schlosseri Pallas) inhabiting five sites distributed along a ≈ 17-km temperature and phytoplankton gradient in the Damariscotta River estuary, Maine, USA. Settlement and population densities and sexual reproductive status were assayed throughout the summer seasons of 1996 and 1997. Sexual reproduction and larval settlement commenced earlier in the summer in up-river populations, which subsequently underwent a seasonal population explosion that was much smalier in down-river populations. Two peaks in settlement density up-river (in early July and early September) suggest that colonies there may have completed two sexual generations, in contrast to a single generation at down-river sites. Similar spatial variation is expected among populations of other taxa with limited larval dispersal when they are distributed across environmental gradients. Published online: 18 September 2002     

Social thermoregulation exerts more influence than microclimate on forest roost preferences by a cavity-dwelling bat

Nest or roost temperature (T _roost) is thought to impact reproductive fitness in many endotherms but few studies have directly tested the hypothesis that naturally occurring variability in nest or roost microclimate is large enough to affect reproductive success. We conducted a field experiment to test whether roost selection by cavity-dwelling, reproductive female big brown bats (Eptesicus fuscus) is more strongly influenced by roost microclimate or a physical characteristic of roosts that facilitates social thermoregulation (i.e., cavity volume). We quantified spatial variability in T _roost within different-sized, unoccupied cavities and also recorded T _roost in occupied vs unoccupied roost trees. We used equations relating energy use and ambient temperature for big brown bats to calculate values of daily energy expenditure from T _roost data because energy is a currency that likely affects reproductive fitness. We found no difference between maximum and minimum T _roost, spatial variability in T _roost, or predicted energy expenditure in more-preferred vs less-preferred roosts. However, there was a significant difference between T _roost and predicted energy expenditure when we compared occupied vs unoccupied roosts. The presence of bats increased T _roost by as much as 7°C, and there was a significant positive correlation between the number of bats occupying a roost, maximum daily T _roost, and energy savings. We calculate that, on average, a normothermic individual would save about 6.5 kJ/day (roughly 9% of the daily energy budget) by roosting in an occupied cavity relative to roosting alone and that savings may increase to 40 kJ/day (about 53% of the energy budget) for an individual roosting in a group of 45 bats. Our findings suggest that variability in microclimate among potential roost or nest sites may be less important to some cavity-dwelling endotherms than has been suggested in previous studies. Our results reinforce the importance of sociality and social thermoregulation to the roosting ecology of forest-living bats and socially roosting or nesting endotherms in general.     

Distance from the queen affects workers’ selfish behaviour in the honeybee (A. mellifera) colony

It has been postulated that spatial organization of the colony, in addition to biological and social factors, influences colony life in honeybees. In this study, we examine the influence of workers’ distance from the queen on their reproductive, pheromonal and behavioural characteristics. Our results demonstrate that with increasing distance from the queen, workers increasingly develop behavioural and reproductive traits characteristic of queenlessness, which are presumably the result of significant impairment in queen pheromone transmission. Having contemplated the alternative possibilities of inadequate transmission of queen signal and voluntary escape from queen control, we concluded, based on the behavioural and physiological data, that the former is far more probable than the latter. The dose-dependent manner of the queen pheromone action and its differential influence on various worker characteristics are discussed.     

Genetic structure of camouflage grouper,<Emphasis Type="Italic"> Epinephelus polyphekadion</Emphasis> (Pisces: Serranidae), in the western central Pacific

The objective of the present study was to investigate the population genetic structure of the commercially important camouflage grouper, Epinephelus polyphekadion (Bleeker, 1849), in the western and central Pacific Ocean to improve existing management. Camouflage grouper are widely distributed in the Indo-Pacific and form brief, seasonal, spawning aggregations that are often heavily fished. The present study examined populations sampled in 1997-1998 at five sites in the western central Pacific spanning a geographic distance of ~5,000 km: New Caledonia, Great Barrier Reef, Palau, Marshall Islands, and Pohnpei (Micronesia). Primer pairs were developed to examine genetic variation at three polymorphic microsatellite loci. Cluster analysis, using genetic distance, revealed three regional groupings: (1) Palau, (2) Pohnpei and the Marshall Islands, and (3) the Great Barrier Reef and New Caledonia. Highly significant allele frequency differences were observed among sites. At Pohnpei, significant allele frequency differences in successive years were also apparent, possibly related to genetic variation among cohorts or between local spawning groups. The inter-annual differences at Pohnpei suggest that there may be further genetic structuring over relatively modest distances, a finding relative to determining management units for this commercially valuable species and suggests that future studies need to incorporate possible small-scale temporal or spatial components into study design.     

Phenotypic plasticity and “cultural transmission” of alternative social organizations in the fire ant Solenopsis invicta

Summary We investigated the process of sexual maturation in winged queens of the fire ant Solenopsis invicta, a species with two distinct forms of social organization. We found that queens of the monogynous social form (single reproductive queen per colony) differ little or not at all from queens of the polygynous form (multiple reproductive queens per colony) in weight and fat content when these are pupae or newly-eclosed adults. Furthermore, the size of a sclerotized region of the adult thorax, which is set during larval growth, does not differ between queens of the two forms. In contrast, winged queens of the two social forms differ dramatically in their physiological phenotypes once they have matured, with monogynous queens weighing more and having greater fat reserves than polygynous queens. A crossfostering experiment revealed that the different maturation processes of queens of the two forms are induced largely by the type of colony in which a queen matures (monogynous or polygynous) rather than being due to intrinsic genetic differences between the forms. However, genetic variation at a single locus does appear to play some role in determining physiological phenotype in queens of the polygynous form, providing an example of genotype-environment interaction in the expression of these physiological traits. Differences between the social forms in the mature phenotypes that are produced constrain the reproductive options of queens, so that the characteristic social organization of a colony is perpetuated by virtue of the social environment in which new queens are reared. Correspondence to: L. Keller     

Social attributes and associated performance measures in marmots: bigger male bullies and weakly affiliating females have higher annual reproductive success

Studying the structure of social interactions is fundamental in behavioral ecology as social behavior often influences fitness and thus natural selection. However, social structure is often complex, and determining the most appropriate measures of variation in social behavior among individuals can be difficult. Social network analysis generates numerous, but often correlated, measures of individual connectedness derived from a network of interactions. We used measures of individual connectedness in networks of affiliative and agonistic interactions in yellow-bellied marmots, Marmota flaviventris, to first determine how variance was structured among network measures. Principal component analysis reduced our set of network measures to four “social attributes” (unweighted connectedness, affiliation strength, victimization, and bullying), which revealed differences between patterns of affiliative and agonistic interactions. We then used these extracted social attributes to examine the relationship between an individual’s social attributes and several performance measures: annual reproductive success, parasite infection, and basal stress. In male marmots, bullying was positively associated with annual reproductive success, while in females, affiliation strength was negatively associated with annual reproductive success. No other social attributes were significantly associated with any performance measures. Our study highlights the utility of considering multiple dimensions when measuring the structure and functional consequences of social behavior.     

Structure of the social network and its influence on transmission dynamics in a honeybee colony

Infectious processes in a social group are driven by a network of contacts that is generally structured by the organization arising from behavioral and spatial heterogeneities within the group. Although theoretical models of transmission dynamics have placed an overwhelming emphasis on the importance of understanding the network structure in a social group, empirical data regarding such contact structures are rare. In this paper, I analyze the network structure and the correlated transmission dynamics within a honeybee colony as determined by food transfer interactions and the changes produced in it by an experimental manipulation. The study demonstrates that widespread transmission in the colony is correlated to a lower clustering coefficient and higher robustness of the social network. I also show that the social network in the colony is determined by the spatial distribution of various age classes, and the resulting organizational structure provides some amount of immunity to the young individuals. The results of this study demonstrates how, using the honeybee colony as a model system, concepts in network theory can be combined with those in behavioral ecology to gain a better understanding of social transmission processes, especially those related to disease dynamics.     

Task-dependent influence of genetic architecture and mating frequency on division of labour in social insect societies

Division of labour is one of the most prominent features of social insects. The efficient allocation of individuals to different tasks requires dynamic adjustment in response to environmental perturbations. Theoretical models suggest that the colony-level flexibility in responding to external changes and internal perturbation may depend on the within-colony genetic diversity, which is affected by the number of breeding individuals. However, these models have not considered the genetic architecture underlying the propensity of workers to perform the various tasks. Here, we investigated how both within-colony genetic variability (stemming from variation in the number of matings by queens) and the number of genes influencing the stimulus (threshold) for a given task at which workers begin to perform that task jointly influence task allocation efficiency. We used a numerical agent-based model to investigate the situation where workers had to perform either a regulatory task or a foraging task. One hundred generations of artificial selection in populations consisting of 500 colonies revealed that an increased number of matings always improved colony performance, whatever the number of loci encoding the thresholds of the regulatory and foraging tasks. However, the beneficial effect of additional matings was particularly important when the genetic architecture of queens comprised one or a few genes for the foraging task’s threshold. By contrast, a higher number of genes encoding the foraging task reduced colony performance with the detrimental effect being stronger when queens had mated with several males. Finally, the number of genes encoding the threshold for the regulatory task only had a minor effect on colony performance. Overall, our numerical experiments support the importance of mating frequency on efficiency of division of labour and also reveal complex interactions between the number of matings and genetic architecture.     

Age-dependent vigilance in winter aggregations of cooperatively breeding white-winged choughs (Corcorax melanorhamphos)

Summary Winter aggregation of cooperatively breeding white-winged choughs (Corcorax melanorhamphos) was examined to elucidate the advantages for different group members. Adults have higher vigilance burdens than juveniles when foraging in their regular group. Aggregation of groups in winter enables the older members to reduce vigilance and increase the time spent foraging. Vigilance was lowest and foraging time highest in aggregations of 20 individuals but the size of cooperatively breeding groups did not exceed fifteen. Despite increased foraging time, choughs did not consume more food items when aggregating.     

Similar is not the same: Social calls of conspecifics are more effective in attracting wild bats to day roosts than those of other bat species

Many bat species regularly need to find new day roosts as they require numerous shelters each breeding season. It has been shown that bats exchange information about roosts among colony members, and use echolocation and social calls of conspecifics in order to find roosts. However, it is unclear if wild bats discriminate between social calls of conspecifics and other bat species while searching for roosts. Furthermore, the extent that bats are attracted to potential roosts by each of these two call types is unknown. We present a field experiment showing that social calls of conspecifics and other bat species both attract bats to roosts. During two summers, we played back social calls of Bechstein’s bats (Myotis bechsteinii) and Natterer’s bats (Myotis nattereri) from different bat boxes that can serve as roosts for these species. All experimental bat boxes were monitored with infrared video to identify the approaching bat species. Three species (M. bechsteinii, M. nattereri, and Plecotus auritus) approached the boxes significantly more often during nights when bat calls were played compared to nights without playbacks. Bechstein’s bats and Natterer’s bats were both more attracted to social calls of conspecifics than of the other species, whereas P. auritus did not discriminate between calls of either Myotis species. Only Bechstein’s bats entered experimental boxes and only at times when calls from conspecifics were played. Our findings show that wild bats discriminate between social calls of conspecifics and other bat species although they respond to both call types when searching for new roosts.     

General methods for sensitivity analysis of equilibrium dynamics in patch occupancy models

Sensitivity analysis is a useful tool for the study of ecological models that has many potential applications for patch occupancy modeling. Drawing from the rich foundation of existing methods for Markov chain models, I demonstrate new methods for sensitivity analysis of the equilibrium state dynamics of occupancy models. Estimates from three previous studies are used to illustrate the utility of the sensitivity calculations: a joint occupancy model for a prey species, its predators, and habitat used by both; occurrence dynamics from a well-known metapopulation study of three butterfly species; and Golden Eagle occupancy and reproductive dynamics. I show how to deal efficiently with multistate models and how to calculate sensitivities involving derived state variables and lower-level parameters. In addition, I extend methods to incorporate environmental variation by allowing for spatial and temporal variability in transition probabilities. The approach used here is concise and general and can fully account for environmental variability in transition parameters. The methods can be used to improve inferences in occupancy studies by quantifying the effects of underlying parameters, aiding prediction of future system states, and identifying priorities for sampling effort.     

Alternative reproductive tactics in the spider Meta segmentata

Summary In the spider Meta segmentata web site quality and competitive ability determine female distributions, and ultimately male mating strategy. Large, fecund females position their webs in aggregations in high quality habitats, whereas small females live alone in lower quality sites. Competition for web sites is intense. Large females have a fighting advantage, but web ownership can offset size disadvantages of about 10%. Males compete for access to female webs. The smallest males are excluded from aggregations of females but have a very high probability of mating monogamously with females in poor habitats. The largest males move about within aggregations and by mating with many females achieve the highest reproductive gains. Moderately sized males only have a low probability of mating in an aggregation, but for those that do succeed the reproductive gains are higher than for those that mate monogamously in low quality areas. On average, moderately sized males obtain reproductive gains from the high risk, high gain strategy of defending webs in aggregations that are equal to those from definding solitary females in poor areas. That 18 of 21 moderately sized males defend females in aggregations suggests that M. segmentata behaves in a riskprone fashion.     

A spatial gradient in the potential reproductive output of the sea mussel <Emphasis Type="Italic">Mytilus californianus</Emphasis>

Spatial variation in reproductive output from different populations within a region could have important consequences for recruitment, and cascading effects on populations and communities of marine species, but is rarely examined over meso-scales (i.e., tens to hundreds of kilometers). In this study, reproduction in the dominant mid-intertidal mussel, Mytilus californianus, was examined from sites spanning Point Conception, California over a 6-month period (March–August 2000). There was a dramatic geographic pattern in the relationship between size and potential reproductive output that was qualitatively similar across all 6 months sampled. Increases in allocation to reproductive tissue with increasing body size occurred at all sites, but the slope nearly doubled at sites south of Point Conception compared to northern sites. The spatial variation in size-specific reproductive output, coupled with additional spatial gradients in mussel density and size distributions, combined to increase total reproductive output by over eightfold at southern relative to northern sites. This study highlights the need to explicitly examine spatial patterns of reproductive output at these meso-scales, in order to better understand connectivity and source–sink dynamics in marine systems.