Heterothermic Migration Strategies in Flying Vertebrates.

Migration is a widespread and highly variable trait among animals. Population-level patterns arise from individual-level decisions, including physiological and energetic constraints. Many aspects of migration are influenced by behaviors and strategies employed during periods of stopover, where migrants may encounter variable or unpredictable conditions. Thermoregulation can be a major cost for homeotherms which largely encounter ambient temperatures below the lower critical temperature during migration, especially during the rest phase of the daily cycle. In this review we describe the empirical evidence, theoretical models, and potential implications of bats and birds that use heterothermy to reduce thermoregulatory costs during migration. Torpor-assisted migration is a strategy described for migrating temperate insectivorous bats, whereby torpor can be used during periods of inactivity to drastically reduce thermoregulatory costs and increase net refueling rate, leading to shorter stopover duration, reduced fuel load requirement, and potential consequences for broad-scale movement patterns and survival. Hummingbirds can adopt a similar strategy, but most birds are not capable of torpor. However, there is an increasing recognition of the use of more shallow heterothermic strategies by diverse bird species during migration, with similarly important implications for migration energetics. A growing body of published literature and preliminary data from ongoing research indicate that heterothermic migration strategies in birds may be more common than traditionally appreciated. We further take a broad evolutionary perspective to consider heterothermy as an alternative to migration in some species, or as a conceptual link to consider alternatives to seasonal resource limitations. There is a growing body of evidence related to heterothermic migration strategies in bats and birds, but many important questions related to the broader implications of this strategy remain.

Erratum: The problem and promise of scale in multilayer animal social networks.

[This corrects the article DOI: 10.1093/cz/zoaa052.].

Plasticity and repeatability in spring migration and parturition dates with implications for annual reproductive success.

In seasonal environments, animals should be adapted to match important life-history traits to when environmental conditions are optimal. Most animal populations therefore reproduce when resource abundance is highest to increase annual reproductive success. When facing variable, and changing, environments animals can display behavioural plasticity to acclimate to changing conditions. Behaviours can further be repeatable. For example, timing of behaviours and life history traits such as timing of reproduction may indicate phenotypic variation. Such variation may buffer animal populations against the consequences of variation and change. Our goal was to quantify plasticity and repeatability in migration and parturition timing in response to timing of snowmelt and green-up in a migratory herbivore (caribou, Rangifer tarandus, n = 132 ID-years) and their effect on reproductive success. We used behavioural reaction norms to quantify repeatability in timing of migration and timing of parturition in caribou and their plasticity to timing of spring events, while also quantifying phenotypic covariance between behavioural and life-history traits. Timing of migration for individual caribou was positively correlated with timing of snowmelt. The timing of parturition for individual caribou varied as a function of inter-annual variation in timing of snowmelt and green-up. Repeatability for migration timing was moderate, but low for timing of parturition. Plasticity did not affect reproductive success. We also did not detect any evidence of phenotypic covariance among any traits examined-timing of migration was not correlated with timing of parturition, and neither was there a correlation in the plasticity of these traits. Repeatability in migration timing suggests the possibility that the timing of migration in migratory herbivores could evolve if the repeatability detected in this study has a genetic or otherwise heritable basis, but observed plasticity may obviate the need for an evolutionary response. Our results also suggest that observed shifts in caribou parturition timing are due to plasticity as opposed to an evolutionary response to changing conditions. While this provides some evidence that populations may be buffered from the consequences of climate change via plasticity, a lack of repeatability in parturition timing could impede adaptation as warming increases.

Behavioural ecology at the spatial-social interface.

Spatial and social behaviour are fundamental aspects of an animal's biology, and their social and spatial environments are indelibly linked through mutual causes and shared consequences. We define the 'spatial-social interface' as intersection of social and spatial aspects of individuals' phenotypes and environments. Behavioural variation at the spatial-social interface has implications for ecological and evolutionary processes including pathogen transmission, population dynamics, and the evolution of social systems. We link spatial and social processes through a foundation of shared theory, vocabulary, and methods. We provide examples and future directions for the integration of spatial and social behaviour and environments. We introduce key concepts and approaches that either implicitly or explicitly integrate social and spatial processes, for example, graph theory, density-dependent habitat selection, and niche specialization. Finally, we discuss how movement ecology helps link the spatial-social interface. Our review integrates social and spatial behavioural ecology and identifies testable hypotheses at the spatial-social interface.

Bibliometric investigation of the integration of animal personality in conservation contexts.

Consistent individual differences in behavior, commonly termed animal personality, are a widespread phenomenon across taxa that have important consequences for fitness, natural selection, and trophic interactions. Animal personality research may prove useful in several conservation contexts, but which contexts remains to be determined. We conducted a structured literature review of 654 studies identified by combining search terms for animal personality and various conservation subfields. We scored the relevance of personality and conservation issues for each study to identify which studies meaningfully integrated the 2 fields as opposed to surface-level connections or vague allusions. We found a taxonomic bias toward mammals (29% of all studies). Very few amphibian or reptile studies applied personality research to conservation issues (6% each). Climate change (21%), invasive species (15%), and captive breeding and reintroduction (13%) were the most abundant conservation subfields that occurred in our search, though a substantial proportion of these papers weakly integrated conservation and animal personality (climate change 54%, invasive species 51%, captive breeding and reintroduction 40%). Based on our results, we recommend that researchers strive for consistent and broadly applicable terminology when describing consistent behavioral differences to minimize confusion and improve the searchability of research. We identify several gaps in the literature that appear to be promising and fruitful avenues for future research, such as disease transmission as a function of sociability or exploration as a driver of space use in protected areas. Practitioners can begin informing future conservation efforts with knowledge gained from animal personality research.

Investigación bibliométrica sobre la integración de la personalidad animal a los contextos de conservación Resumen Las diferencias individuales y constantes en el comportamiento, comúnmente llamadas personalidad animal, son un fenómeno generalizado en los taxones con consecuencias importantes para la aptitud, selección natural e interacciones tróficas. Las investigaciones sobre la personalidad animal pueden ser útiles en varios contextos de conservación, aunque falta determinar cuáles son estos contextos. Realizamos una revisión literaria estructurada de 654 estudios identificados mediante la combinación de los términos de búsqueda para la personalidad animal y varios subcampos de la conservación. Puntuamos la relevancia de la personalidad y los temas de conservación en cada estudio para identificar cuáles de estos integraron significativamente a ambos campos, contrario a las conexiones a nivel superficial o alusiones vagas. Descubrimos un sesgo taxonómico por los mamíferos (29% de todos los estudios). Pocos estudios enfocados en anfibios o reptiles aplicaron un estudio de personalidad a los temas de conservación (6% para cada uno). El cambio climático (21%), las especies invasoras (15%) y la reproducción en cautiverio y las reintroducciones (13%) fueron los subcampos de conservación más abundantes que aparecieron en nuestra búsqueda, aunque una proporción significativa de estos artículos integraron muy poco a la conservación y la personalidad animal (cambio climático 54%, especies invasoras 51%, reproducción en cautiverio y reintroducciones 40%). Con base en nuestros resultados, recomendamos que los investigadores procuren tener terminologías consistentes y de aplicación generalizada cuando describan las diferencias conductuales para así minimizar las confusiones y facilitar la búsqueda durante la investigación. Identificamos varios vacíos en la literatura que prometen ser vías fructíferas para las investigaciones en el futuro, como la transmisión de enfermedades como una función sociable o la exploración como un impulsor del uso del espacio en las áreas protegidas. Los practicantes pueden comenzar por guiar los siguientes esfuerzos de conservación con el conocimiento obtenido de las investigaciones sobre la personalidad animal.

Density-dependent plasticity in territoriality revealed using social network analysis.

Territories are typically defined as spatially exclusive areas that are defended against conspecifics. Given the spatial nature of territoriality, it is inherently density dependent, but the economics of territoriality also depend on the distribution and abundance of defended resources. Our objectives were to assess the effects of changing population density and food availability on individually based territorial phenotypes. We developed a novel analytical framework that bridges spatially explicit territories with social network analysis to model density-dependent territorial phenotypes. Using the outputs from our data pipeline, we modelled plasticity in territory size and territory intrusion rates in a long-term study population of North American red squirrels Tamiasciurus hudsonicus. Red squirrels defend year-round territories around a central hoard (midden) of white spruce Picea glauca cones. Importantly, white spruce is a masting species that produces large cone crops every 4-7 years (i.e. mast years) in our study area interspersed with non-mast years when few cones are produced. In the spring following mast years, populations are approximately double in size, but are lower in the spring of non-mast years. We predicted that territory size and intrusion rates would decrease as resource abundance, and consequently population density, increased. By contrast, as resource abundance decreased via depletion, and therefore density decreased, territories should increase in size and intrusions should also increase. As we expected, individual territory size and territorial intrusions were negatively density dependent, such that increased density after mast years resulted in smaller territories and fewer intrusions. When considering between-individual variation in plasticity across a density gradient, individuals responded differently to changes in population density within their lifetime. Our results show that territory size and intrusion rates display negative density dependence. When food becomes available in the autumn of a mast year and density in spring of the following year increases, territories shrink in size to effectively a small area around the midden. While our findings for red squirrels are unique compared to other systems, they serve as a reminder that the direction and strength of fundamental ecological relationships can depend on the nature of the system.

Social Effects on Annual Fitness in Red Squirrels.

When resources are limited, mean fitness is constrained and competition can cause genes and phenotypes to enhance an individual's own fitness while reducing the fitness of their competitors. Negative social effects on fitness have the potential to constrain adaptation, but the interplay between ecological opportunity and social constraints on adaptation remains poorly studied in nature. Here, we tested for evidence of phenotypic social effects on annual fitness (survival and reproductive success) in a long-term study of wild North American red squirrels (Tamiasciurus hudsonicus) under conditions of both resource limitation and super-abundant food resources. When resources were limited, populations remained stable or declined, and there were strong negative social effects on annual survival and reproductive success. That is, mean fitness was constrained and individuals had lower fitness when other nearby individuals had higher fitness. In contrast, when food resources were super-abundant, populations grew and social constraints on reproductive success were greatly reduced or eliminated. Unlike reproductive success, social constraints on survival were not significantly reduced when food resources were super-abundant. These findings suggest resource-dependent social constraints on a component of fitness, which have important potential implications for evolution and adaptation.

Influence of number of individuals and observations per individual on a model of community structure.

Social network analysis is increasingly applied to understand animal groups. However, it is rarely feasible to observe every interaction among all individuals in natural populations. Studies have assessed how missing information affects estimates of individual network positions, but less attention has been paid to metrics that characterize overall network structure such as modularity, clustering coefficient, and density. In cases such as groups displaying fission-fusion dynamics, where subgroups break apart and rejoin in changing conformations, missing information may affect estimates of global network structure differently than in groups with distinctly separated communities due to the influence single individuals can have on the connectivity of the network. Using a bat maternity group showing fission-fusion dynamics, we quantify the effect of missing data on global network measures including community detection. In our system, estimating the number of communities was less reliable than detecting community structure. Further, reliably assorting individual bats into communities required fewer individuals and fewer observations per individual than to estimate the number of communities. Specifically, our metrics of global network structure (i.e., graph density, clustering coefficient, Rcom) approached the 'real' values with increasing numbers of observations per individual and, as the number of individuals included increased, the variance in these estimates decreased. Similar to previous studies, we recommend that more observations per individual should be prioritized over including more individuals when resources are limited. We recommend caution when making conclusions about animal social networks when a substantial number of individuals or observations are missing, and when possible, suggest subsampling large datasets to observe how estimates are influenced by sampling intensity. Our study serves as an example of the reliability, or lack thereof, of global network measures with missing information, but further work is needed to determine how estimates will vary with different data collection methods, network structures, and sampling periods.

The problem and promise of scale in multilayer animal social networks.

Scale remains a foundational concept in ecology. Spatial scale, for instance, has become a central consideration in the way we understand landscape ecology and animal space use. Meanwhile, scale-dependent social processes can range from fine-scale interactions to co-occurrence and overlapping home ranges. Furthermore, sociality can vary within and across seasons. Multilayer networks promise the explicit integration of the social, spatial, and temporal contexts. Given the complex interplay of sociality and animal space use in heterogeneous landscapes, there remains an important gap in our understanding of the influence of scale on animal social networks. Using an empirical case study, we discuss ways of considering social, spatial, and temporal scale in the context of multilayer caribou social networks. Effective integration of social and spatial processes, including biologically meaningful scales, within the context of animal social networks is an emerging area of research. We incorporate perspectives that link the social environment to spatial processes across scales in a multilayer context.

Personality affects dynamics of an experimental pathogen in little brown bats.

Host behaviour can affect host-pathogen dynamics and theory predicts that certain individuals disproportionately infect conspecifics during an epidemic. Consistent individual differences in behaviour, or personality, could influence this variation with the most exploratory or sociable individuals most likely to spread pathogens. We quantified exploration and sociability in little brown bats (Myotis lucifugus) and then experimentally manipulated exposure to a proxy pathogen (i.e. ultraviolet (UV) fluorescent powder) to test two related hypotheses: (i) more sociable and more exploratory individuals would be more likely to transmit infections to other individuals, and (ii) more sociable and more exploratory individuals uninfected with an invading pathogen would be more likely to acquire infections. We captured 10 groups of 16 bats at a time and held each group in an outdoor flight tent equipped with roosting-boxes. We used hole-board and Y-maze tests to quantify exploration and sociability of each bat and randomly selected one individual from each group for 'infection' with non-toxic, UV fluorescent powder. Each group of 10 bats was released into the flight tent for 24 h, which represented an experimental infection trial. After 24 h, we removed bats from the trial, photographed each individual under UV light and quantified infection intensity from digital photographs. As predicted, the exploratory behaviour of the experimentally infected individual was positively correlated with infection intensity in their group-mates, while more exploratory females had higher pathogen acquisition. Our results highlight the potential influence of host personality and sex on pathogen dynamics in wildlife populations.

Geometry of the ideal free distribution: individual behavioural variation and annual reproductive success in aggregations of a social ungulate.

Variation in social environment can mitigate risks and rewards associated with occupying a particular patch. We aim to integrate Ideal Free Distribution (IFD) and Geometry of the Selfish Herd (GSH) to address an apparent conflict in their predictions of equal mean fitness between patches (IFD) and declining fitness benefits within a patch (GSH). We tested these hypotheses in a socio-spatial context using individual caribou that were aggregated or disaggregated during calving and varied in their annual reproductive success (ARS). We then tested individual consistency of these spatial tactics. We reveal that two socio-spatial tactics accorded similar mean ARS (IFD); however, ARS for aggregated individuals declined near the periphery (GSH). Individuals near the aggregation periphery exhibited flexibility, whereas others were consistent. The integration of classical theories through a contemporary lens of consistent individual differences provides evidence for an integrated GSH and IFD strategy that may represent an evolutionary stable state.

The Ecology of Individual Differences Empirically Applied to Space-Use and Movement Tactics.

Movement provides a link between individual behavioral ecology and the spatial and temporal variation in an individual's landscape. Individual variation in movement traits is an important axis of animal personality, particularly in the context of foraging ecology. We tested whether individual caribou (Rangifer tarandus) displayed plasticity in movement and space-use behavior across a gradient of resource aggregation. We quantified first-passage time and range-use ratio as proxies for movement-related foraging behavior and examined how these traits varied at the individual level across a foraging resource gradient. Our results suggest that individuals adjusted first-passage time but not range-use ratio to maximize access to high-quality foraging resources. First-passage time was repeatable, and intercepts for first-passage time and range-use ratio were negatively correlated. Individuals matched first-passage time but not range-use ratio to the expectations of our patch-use model that maximized access to foraging resources, a result that suggests that individuals acclimated their movement patterns to accommodate both intra- and interannual variation in foraging resources on the landscape. Collectively, we highlight repeatable movement and space-use tactics and provide insight into how individual plasticity in movement interacts with landscape processes to affect the distribution of behavioral phenotypes and potentially fitness and population dynamics.

Space use and social association in a gregarious ungulate: Testing the conspecific attraction and resource dispersion hypotheses.

Animals use a variety of proximate cues to assess habitat quality when resources vary spatiotemporally. Two nonmutually exclusive strategies to assess habitat quality involve either direct assessment of landscape features or observation of social cues from conspecifics as a form of information transfer about forage resources. The conspecific attraction hypothesis proposes that individual space use is dependent on the distribution of conspecifics rather than the location of resource patches, whereas the resource dispersion hypothesis proposes that individual space use and social association are driven by the abundance and distribution of resources. We tested the conspecific attraction and the resource dispersion hypotheses as two nonmutually exclusive hypotheses explaining social association and of adult female caribou (Rangifer tarandus). We used location data from GPS collars to estimate interannual site fidelity and networks representing home range overlap and social associations among individual caribou. We found that home range overlap and social associations were correlated with resource distribution in summer and conspecific attraction in winter. In summer, when resources were distributed relatively homogeneously, interannual site fidelity was high and home range overlap and social associations were low. Conversely, in winter when resources were distributed relatively heterogeneously, interannual site fidelity was low and home range overlap and social associations were high. As access to resources changes across seasons, caribou appear to alter social behavior and space use. In summer, caribou may use cues associated with the distribution of forage, and in winter caribou may use cues from conspecifics to access forage. Our results have broad implications for our understanding of caribou socioecology, suggesting that caribou use season-specific strategies to locate forage. Caribou populations continue to decline globally, and our finding that conspecific attraction is likely related to access to forage suggests that further fragmentation of caribou habitat could limit social association among caribou, particularly in winter when access to resources may be limited.

Testing the "Fasting While Foraging" Hypothesis: Effects of Recent Feeding on Plasma Metabolite Concentrations in Little Brown Bats (Myotis lucifugus).

Plasma metabolite concentrations can be used to understand nutritional status and foraging behavior across ecological contexts including prehibernation fattening, migration refueling, and variation in foraging habitat quality. Generally, high plasma concentrations of the ketone ß-hydroxybutyrate, a product of fat catabolism, indicate fasting, while triglycerides indicate recent feeding and fat accumulation. In recent studies of insectivorous bats, triglyceride concentration increased after feeding as expected, but ß-hydroxybutyrate also unexpectedly increased rather than decreased. An aerial-hawking foraging strategy is energetically demanding, and thus it has been hypothesized that foraging by insectivorous bats requires catabolism of stored fat. We tested this hypothesis by quantifying plasma ß-hydroxybutyrate and triglyceride concentration following feeding in little brown bats (Myotis lucifugus) that were temporarily housed in individual cages to prevent flight. We provided a fixed amount of food and collected blood samples at different intervals after feeding to produce variation in plasma metabolite concentrations. Plasma triglyceride concentration responded as predicted, but similar to previous studies and contrary to our prediction, when flight was eliminated plasma ß-hydroxybutyrate concentration responded similarly to triglyceride. Thus, it is unlikely that the unexpected plasma ß-hydroxybutyrate patterns observed in previous studies were related to flight. The mechanism underlying this unexpected pattern remains unknown, but the response has been consistent in all studies to date. Thus, plasma metabolite analysis provides an effective tool for studies of nutritional status, although more work is needed to understand why insectivorous bats respond differently than other taxa.

An experimental test of effects of ambient temperature and roost quality on aggregation by little brown bats (Myotis lucifugus).

Environmental factors, such as ambient temperature (Ta) or roost/nest quality, can influence social behaviour of small-bodied endotherms because individuals may aggregate for social thermoregulation when Ta is low or select the warmest possible sites for roosting. Female temperate bats form maternity colonies in spring to communally raise pups and exploit social thermoregulation. They also select roosts with warm microclimates because low roost temperature (Troost) delays juvenile development. We studied captive female little brown bats (Myotis lucifugus) to test the hypothesis that variation in Ta and Troost influence social group size. First, we predicted that female bats would preferentially select artificially heated roosts over unheated roosts. Second, we predicted that, as Ta decreased, group size would increase because bats would rely more heavily on social thermoregulation. Third, we predicted that experimentally increasing Troost (i.e., roost quality) above Ta would result in larger group sizes due to greater aggregation in high quality roosts. We captured 34 females from a maternity colony and housed them in a flight-tent provisioned with four bat boxes. Each box was outfitted with a heating pad and thermostat. Over the course of eight-days we heated each roost box in sequence to near thermoneutral Troost for two days. Bats preferentially selected heated roosts over unheated roosts but, contrary to our prediction, group size decreased when Troost was much greater than Ta (i.e., when the benefits of a warm roost should have been highest). Our results suggest that social thermoregulation and the availability of warm roosts influence aggregation in bats and have implications for the potential of summer habitat protection and enhancement to help bat populations in the face of threats like white-nose syndrome.

Consistent individual differences and population plasticity in network-derived sociality: An experimental manipulation of density in a gregarious ungulate.

In many taxa, individual social traits appear to be consistent across time and context, thus meeting the criteria for animal personality. How these differences are maintained in response to changes in population density is unknown, particularly in large mammals, such as ungulates. Using a behavioral reaction norm (BRN) framework, we examined how among- and within-individual variation in social connectedness, measured using social network analyses, change as a function of population density. We studied a captive herd of elk (Cervus canadensis) separated into a group of male elk and a group of female elk. Males and females were exposed to three different density treatments and we recorded social associations between individuals with proximity-detecting radio-collars fitted to elk. We constructed social networks using dyadic association data and calculated three social network metrics reflective of social connectedness: eigenvector centrality, graph strength, and degree. Elk exhibited consistent individual differences in social connectedness across densities; however, they showed little individual variation in their response to changes in density, i.e., individuals oftentimes responded plastically, but in the same manner to changes in density. Female elk had highest connectedness at an intermediate density. In contrast, male elk increased connectedness with increasing density. Whereas this may suggest that the benefits of social connectedness outweigh the costs of increased competition at higher density for males, females appear to exhibit a threshold in social benefits (e.g. predator detection and forage information). Our study illustrates the importance of viewing social connectedness as a density-dependent trait, particularly in the context of plasticity. Moreover, we highlight the need to revisit our understanding of density dependence as a population-level phenomenon by accounting for consistent individual differences not only in social connectedness, but likely in other ecological processes (e.g., predator-prey dynamics, mate choice, disease transfer).

Spatial variation of mercury bioaccumulation in bats of Canada linked to atmospheric mercury deposition.

Wildlife are exposed to neurotoxic mercury at locations distant from anthropogenic emission sources because of long-range atmospheric transport of this metal. In this study, mercury bioaccumulation in insectivorous bat species (Mammalia: Chiroptera) was investigated on a broad geographic scale in Canada. Fur was analyzed (n=1178) for total mercury from 43 locations spanning 20° latitude and 77° longitude. Total mercury and methylmercury concentrations in fur were positively correlated with concentrations in internal tissues (brain, liver, kidney) for a small subset (n=21) of little brown bats (Myotis lucifugus) and big brown bats (Eptesicus fuscus), validating the use of fur to indicate internal mercury exposure. Brain methylmercury concentrations were approximately 10% of total mercury concentrations in fur. Three bat species were mainly collected (little brown bats, big brown bats, and northern long-eared bats [M. septentrionalis]), with little brown bats having lower total mercury concentrations in their fur than the other two species at sites where both species were sampled. On average, juvenile bats had lower total mercury concentrations than adults but no differences were found between males and females of a species. Combining our dataset with previously published data for eastern Canada, median total mercury concentrations in fur of little brown bats ranged from 0.88-12.78µg/g among 11 provinces and territories. Highest concentrations were found in eastern Canada where bats are most endangered from introduced disease. Model estimates of atmospheric mercury deposition indicated that eastern Canada was exposed to greater mercury deposition than central and western sites. Further, mean total mercury concentrations in fur of adult little brown bats were positively correlated with site-specific estimates of atmospheric mercury deposition. This study provides the largest geographic coverage of mercury measurements in bats to date and indicates that atmospheric mercury deposition is important in determining spatial patterns of mercury accumulation in a mammalian species.

An evolutionary framework outlining the integration of individual social and spatial ecology.

Behaviour is the interface between an organism and its environment, and behavioural plasticity is important for organisms to cope with environmental change. Social behaviour is particularly important because sociality is a dynamic process, where environmental variation influences group dynamics and social plasticity can mediate resource acquisition. Heterogeneity in the ecological environment can therefore influence the social environment. The combination of the ecological and social environments may be interpreted collectively as the "socioecological environment," which could explain variation in fitness. Our objective was to outline a framework through which individual social and spatial phenotypes can be integrated and interpreted as phenotypes that covary as a function of changes in the socioecological environment. We propose the socioecological environment is composed of individual behavioural traits, including sociality and habitat selection, both of which are repeatable, potentially heritable and may reflect animal personality traits. We also highlight how ecological and social niche theory can be applied to the socioecological environment framework, where individuals occupy different socioecological niches. Individual sociality and habitat selection are also density-dependent, and theory predicts that density-dependent traits should affect reproduction, survival, and therefore fitness and population dynamics. We then illustrate the proximate links between sociality, habitat selection and fitness as well as the ultimate, and possibly adaptive, consequences associated with changes in population density. The ecological, evolutionary and applied implications of our proposed socioecological environment framework are broad and changes in density could influence individual fitness and population dynamics. For instance, human-induced environmental changes can influence population density, which can affect the distribution of social and spatial phenotypes within a population. In summary, we outline a conceptual framework that incorporates individual social and spatial behavioural traits with fitness and we highlight a range of ecological and evolutionary processes that are likely associated with the socioecological environment.

Viral Richness is Positively Related to Group Size, but Not Mating System, in Bats.

Characterizing host traits that influence viral richness and diversification is important for understanding wildlife pathogens affecting conservation and/or human health. Behaviors that affect contact rates among hosts could be important for viral diversification because more frequent intra- and inter-specific contacts among hosts should increase the potential for viral diversification within host populations. We used published data on bats to test the contact-rate hypothesis. We predicted that species forming large conspecific groups, that share their range with more heterospecifics (i.e., sympatry), and with mating systems characterized by high contact rates (polygynandry: multi-male/multi-female), would host higher viral richness than species with small group sizes, lower sympatry, or low contact-rate mating systems (polygyny: single male/multi-female). Consistent with our hypothesis and previous research, viral richness was positively correlated with conspecific group size although the relationship plateaued at group sizes of approximately several hundred thousand bats. This pattern supports epidemiological theory that, up to a point, larger groups have higher contact rates, greater likelihood of acquiring and transmitting viruses, and ultimately greater potential for viral diversification. However, contrary to our hypothesis, there was no effect of sympatry on viral richness and no difference in viral richness between mating systems. We also found no residual effect of host phylogeny on viral richness, suggesting that closely related species do not necessarily host similar numbers of viruses. Our results support the contact-rate hypothesis that intra-specific viral transmission can enhance viral diversification within species and highlight the influence of host group size on the potential of viruses to propagate within host populations.