Adult body mass is heritable, positively genetically correlated and under selection of differing shapes between the sexes in a bird with little apparent sexual dimorphism.

In most animals, body mass varies with ecological conditions and is expected to reflect how much energy can be allocated to reproduction and survival. Because the sexes often differ in their resource acquisition and allocation strategies, variations in adult body mass and their consequences on fitness can differ between the sexes. Assessing the relative contributions of environmental and genetic effects (i.e. heritability)-and whether these effects and their fitness consequences are sex-specific-is essential to gain insights into the evolution of sexual dimorphism and sexual conflicts. We used 20+ years of data to study the sources of variation in adult body mass and associated fitness consequences in a bird with biparental care, the Alpine swift (Tachymarptis melba). Swifts appear monomorphic to human observers, though subtle dimorphisms are present. We first investigated the effects of weather conditions on adult body mass using a sliding window analysis approach. We report a positive effect of temperature and a negative effect of rainfall on adult body mass, as expected for an aerial insectivorous bird. We then quantified the additive genetic variance and heritability of body mass in both sexes and assessed the importance of genetic constraints on mass evolution by estimating the cross-sex genetic correlation. Heritability was different from zero in both sexes at ~0.30. The positive cross-sex genetic correlation and comparable additive genetic variance between the sexes suggest the possibility for evolutionary constraints when it comes to body mass. Finally, we assessed the sex-specific selection on adjusted body mass using multiple fitness components. We report directional positive and negative selection trending towards stabilizing and diversifying selection on females and males respectively in relation to the weighted proportion of surviving fledglings. Overall, these results suggest that while body mass may be able to respond to environmental conditions and evolve, genetic constraints would result in similar changes in both sexes or an overall absence of response to selection. It remains unclear whether the weak (1%) dimorphism in Alpine swift body mass we report is simply a result of the similar fitness peaks between the sexes or of genetic constraints.

Information Gathering Is Associated with Increased Survival: A Field Experiment in Black-Capped Chickadees (Poecile atricapillus).

AbstractSampling, investing time or energy to learn about the environment, allows organisms to track changes in resource distribution and quality. The use of sampling is predicted to change as a function of energy expenditure, food availability, and starvation risk, all of which can vary both within and among individuals. We studied sampling behavior in a field study with black-capped chickadees (Poecile atricapillus) and show that individuals adjust their use of sampling as a function of ambient temperature (a proxy for energy expenditure), the presence of an alternative food source (yes or no, a proxy for risk of energy shortfall), and their interaction, as predicted by models of optimal sampling. We also observed repeatable differences in sampling. Some individuals consistently sampled more, and individuals that sampled more overall also had a higher annual survival. These results are consistent with among-individual differences in resource acquisition (e.g., food caches or dominance-related differences in priority access to feeders), shaping among-individual differences in both sampling and survival, with greater resource acquisition leading to both higher sampling and higher survival. Although this explanation requires explicit testing, it is in line with several recent studies suggesting that variation in resource acquisition is a key mechanism underlying animal personality.

Dispersal mediates trophic interactions and habitat connectivity to alter metacommunity composition.

Dispersal contributes vitally to metacommunity structure. However, interactions between dispersal and other key processes have rarely been explored, particularly in the context of multitrophic metacommunities. We investigated such a metacommunity in naturally fragmented habitats populated by butterfly species (whose dispersal capacities were previously assessed), flowering plants, and butterfly predators. Using data on butterfly species abundance, floral abundance, and predation (on experimentally placed clay butterfly models), we asked how dispersal ability mediates interactions with predators, mutualists, and the landscape matrix. In contrast to expectations, high densities of strong dispersers were found in more isolated sites and sites with low floral resource density, while intermediate dispersers maintained similar densities across isolation and floral gradients, and higher densities of poor dispersers were found in more connected sites and sites with higher floral density. These findings raise questions about how strong dispersers experience the landscape matrix and the quality of isolated and low-resource sites. Strong dispersers were able to escape habitat patches with high predation, while intermediate dispersers maintained similar densities along a predation gradient, and poor dispersers occurred at high densities in these patches, exposing them to interactions with predators. This work demonstrates that species that vary in dispersal capacities interact differently with predators and mutualist partners in a landscape context, shaping metacommunity composition.

The timing of reproduction is responding plastically, not genetically, to climate change in yellow-bellied marmots (Marmota flaviventer).

With global climates changing rapidly, animals must adapt to new environmental conditions with altered weather and phenology. The key to adapting to these new conditions is adjusting the timing of reproduction to maximize fitness. Using a long-term dataset on a wild population of yellow-bellied marmots (Marmota flaviventer) at the Rocky Mountain Biological Laboratory (RMBL), we investigated how the timing of reproduction changed with changing spring conditions over the past 50 years. Marmots are hibernators with a 4-month active season. It is thus crucial to reproduce early enough in the season to have time to prepare for hibernation, but not too early, as snow cover prevents access to food. Importantly, climate change in this area has, on average, increased spring temperatures by 5°C and decreased spring snowpack by 50 cm over the past 50 years. We evaluated how female marmots adjust the timing of their reproduction in response to changing conditions and estimated the importance of both microevolution and plasticity in the variation in this timing. We showed that, within a year, the timing of reproduction is not as tightly linked to the date a female emerges from hibernation as previously thought. We reported a positive effect of spring snowpack but not of spring temperature on the timing of reproduction. We found inter-individual variation in the timing of reproduction, including low heritability, but not in its response to changing spring conditions. There was directional selection for earlier reproduction since it increased the number and proportion of pups surviving their first winter. Taken together, the timing of marmot reproduction might evolve via natural selection; however, plastic changes will also be extremely important. Further, future studies on marmots should not operate under the assumption that females reproduce immediately following their emergence.

The effect of heterospecific and conspecific competition on inter-individual differences in tungara frog tadpole (Engystomops pustulosus) behavior.

Repeated social interactions with conspecifics and/or heterospecifics during early development may drive the differentiation of behavior among individuals. Competition is a major form of social interaction and its impacts can depend on whether interactions occur between conspecifics or heterospecifics and the directionality of a response could be specific to the ecological context that they are measured in. To test this, we reared tungara frog tadpoles (Engystomops pustulosus) either in isolation, with a conspecific tadpole or with an aggressive heterospecific tadpole, the whistling frog tadpole (Leptodactylus fuscus). In each treatment, we measured the body size and distance focal E. pustulosus tadpoles swam in familiar, novel and predator risk contexts six times during development. We used univariate and multivariate hierarchical mixed effect models to investigate the effect of treatment on mean behavior, variance among and within individuals, behavioral repeatability and covariance among individuals in their behavior between contexts. There was a strong effect of competition on behavior, with different population and individual level responses across social treatments. Within a familiar context, the variance in the distance swam within individuals decreased under conspecific competition but heterospecific competition caused more variance in the average distance swam among individuals. Behavioral responses were also context specific as conspecific competition caused an increase in the distance swam within individuals in novel and predator risk contexts. The results highlight that the impact of competition on among and within individual variance in behavior is dependent on both competitor species identity and context.

Current and lagged climate affects phenology across diverse taxonomic groups.

The timing of life events (phenology) can be influenced by climate. Studies from around the world tell us that climate cues and species' responses can vary greatly. If variation in climate effects on phenology is strong within a single ecosystem, climate change could lead to ecological disruption, but detailed data from diverse taxa within a single ecosystem are rare. We collated first sighting and median activity within a high-elevation environment for plants, insects, birds, mammals and an amphibian across 45 years (1975-2020). We related 10 812 phenological events to climate data to determine the relative importance of climate effects on species' phenologies. We demonstrate significant variation in climate-phenology linkage across taxa in a single ecosystem. Both current and prior climate predicted changes in phenology. Taxa responded to some cues similarly, such as snowmelt date and spring temperatures; other cues affected phenology differently. For example, prior summer precipitation had no effect on most plants, delayed first activity of some insects, but advanced activity of the amphibian, some mammals, and birds. Comparing phenological responses of taxa at a single location, we find that important cues often differ among taxa, suggesting that changes to climate may disrupt synchrony of timing among taxa.

Demographic consequences of changes in environmental periodicity.

The fate of natural populations is mediated by complex interactions among vital rates, which can vary within and among years. Although the effects of random, among-year variation in vital rates have been studied extensively, relatively little is known about how periodic, nonrandom variation in vital rates affects populations. This knowledge gap is potentially alarming as global environmental change is projected to alter common periodic variations, such as seasonality. We investigated the effects of changes in vital-rate periodicity on populations of three species representing different forms of adaptation to periodic environments: the yellow-bellied marmot (Marmota flaviventer), adapted to strong seasonality in snowfall; the meerkat (Suricata suricatta), adapted to inter-annual stochasticity as well as seasonal patterns in rainfall; and the dewy pine (Drosophyllum lusitanicum), adapted to fire regimes and periodic post-fire habitat succession. To assess how changes in periodicity affect population growth, we parameterized periodic matrix population models and projected population dynamics under different scenarios of perturbations in the strength of vital-rate periodicity. We assessed the effects of such perturbations on various metrics describing population dynamics, including the stochastic growth rate, log λS . Overall, perturbing the strength of periodicity had strong effects on population dynamics in all three study species. For the marmots, log λS decreased with increased seasonal differences in adult survival. For the meerkats, density dependence buffered the effects of perturbations of periodicity on log λS . Finally, dewy pines were negatively affected by changes in natural post-fire succession under stochastic or periodic fire regimes with fires occurring every 30 years, but were buffered by density dependence from such changes under presumed more frequent fires or large-scale disturbances. We show that changes in the strength of vital-rate periodicity can have diverse but strong effects on population dynamics across different life histories. Populations buffered from inter-annual vital-rate variation can be affected substantially by changes in environmentally driven vital-rate periodic patterns; however, the effects of such changes can be masked in analyses focusing on inter-annual variation. As most ecosystems are affected by periodic variations in the environment such as seasonality, assessing their contributions to population viability for future global-change research is crucial.

RocTest: A standardized method to assess the performance of root organ cultures in the propagation of arbuscular mycorrhizal fungi.

Over the past three decades, root organ cultures (ROCs) have been the gold standard method for studying arbuscular mycorrhizal fungi (AMF) under in vitro conditions, and ROCs derived from various plant species have been used as hosts for AM monoxenic cultures. While there is compelling evidence that host identity can significantly modify AMF fitness, there is currently no standardized methodology to assess the performance of ROCs in the propagation of their fungal symbionts. We describe RocTest, a robust methodological approach that models the propagation of AMF in symbiosis with ROCs. The development of extraradical fungal structures and the pattern of sporulation are modeled using cumulative link mixed models and linear mixed models. We demonstrate functionality of RocTest by evaluating the performance of three species of ROCs (Daucus carota, Medicago truncatula, Nicotiana benthamiana) in the propagation of three species of AMF (Rhizophagus clarus, Rhizophagus irregularis, Glomus sp.). RocTest produces a simple graphical output to assess the performance of ROCs and shows that fungal propagation depends on the three-way interaction between ROC, AMF, and time. RocTest makes it possible to identify the best combination of host/AMF for fungal development and spore production, making it an important asset for germplasm collections and AMF research.

Hibernation slows epigenetic ageing in yellow-bellied marmots.

Species that hibernate generally live longer than would be expected based solely on their body size. Hibernation is characterized by long periods of metabolic suppression (torpor) interspersed by short periods of increased metabolism (arousal). The torpor-arousal cycles occur multiple times during hibernation, and it has been suggested that processes controlling the transition between torpor and arousal states cause ageing suppression. Metabolic rate is also a known correlate of longevity; we thus proposed the 'hibernation-ageing hypothesis' whereby ageing is suspended during hibernation. We tested this hypothesis in a well-studied population of yellow-bellied marmots (Marmota flaviventer), which spend 7-8 months per year hibernating. We used two approaches to estimate epigenetic age: the epigenetic clock and the epigenetic pacemaker. Variation in epigenetic age of 149 samples collected throughout the life of 73 females was modelled using generalized additive mixed models (GAMM), where season (cyclic cubic spline) and chronological age (cubic spline) were fixed effects. As expected, the GAMM using epigenetic ages calculated from the epigenetic pacemaker was better able to detect nonlinear patterns in epigenetic ageing over time. We observed a logarithmic curve of epigenetic age with time, where the epigenetic age increased at a higher rate until females reached sexual maturity (two years old). With respect to circannual patterns, the epigenetic age increased during the active season and essentially stalled during the hibernation period. Taken together, our results are consistent with the hibernation-ageing hypothesis and may explain the enhanced longevity in hibernators.

Effectiveness of 20 years of conservation investments in protecting orangutans.

Conservation strategies are rarely systematically evaluated, which reduces transparency, hinders the cost-effective deployment of resources, and hides what works best in different contexts. Using data on the iconic and critically endangered orangutan (Pongo spp.), we developed a novel spatiotemporal framework for evaluating conservation investments. We show that around USD 1 billion was invested between 2000 and 2019 into orangutan conservation by governments, nongovernmental organizations, companies, and communities. Broken down by allocation to different conservation strategies, we find that habitat protection, patrolling, and public outreach had the greatest return on investment for maintaining orangutan populations. Given the variability in threats, land-use opportunity costs, and baseline remunerations in different regions, there were differential benefits per dollar invested across conservation activities and regions. We show that although challenging from a data and analysis perspective, it is possible to fully understand the relationships between conservation investments and outcomes and the external factors that influence these outcomes. Such analyses can provide improved guidance toward a more effective biodiversity conservation. Insights into the spatiotemporal interplays between the costs and benefits driving effectiveness can inform decisions about the most suitable orangutan conservation strategies for halting population declines. Although our study focuses on the three extant orangutan species of Sumatra and Borneo, our findings have broad application for evidence-based conservation science and practice worldwide.

Albatrosses respond adaptively to climate variability by changing variance in a foraging trait.

The ability of individuals and populations to adapt to a changing climate is a key determinant of population dynamics. While changes in mean behaviour are well studied, changes in trait variance have been largely ignored, despite being assumed to be crucial for adapting to a changing environment. As the ability to acquire resources is essential to both reproduction and survival, changes in behaviours that maximize resource acquisition should be under selection. Here, using foraging trip duration data collected over 7 years on black-browed albatrosses (Thalassarche melanophris) on the Kerguelen Islands in the southern Indian Ocean, we examined the importance of changes in the mean and variance in foraging behaviour, and the associated effects on fitness, in response to the El Niño Southern Oscillation (ENSO). Using double hierarchical models, we found no evidence that individuals change their mean foraging trip duration in response to a changing environment, but found strong evidence of changes in variance. Younger birds showed greater variability in foraging trip duration in poor conditions as did birds with higher fitness. However, during brooding, birds showed greater variability in foraging behaviour under good conditions, suggesting that optimal conditions allow the alteration between chick provisioning and self-maintenance trips. We found weak correlations between sea surface temperature and the ENSO, but stronger links with sea-level pressure. We suggest that variability in behavioural traits affecting resource acquisition is under selection and offers a mechanism by which individuals can adapt to a changing climate. Studies which look only at effects on mean behaviour may underestimate the effects of climate change and fail to consider variance in traits as a key evolutionary force.

Bigger is not always better: Viability selection on body mass varies across life stages in a hibernating mammal.

Body mass is often viewed as a proxy of past access to resources and of future survival and reproductive success. Links between body mass and survival or reproduction are, however, likely to differ between age classes and sexes. Remarkably, this is rarely taken into account in selection analyses. Selection on body mass is likely to be the primary target accounting for juvenile survival until reproduction but may weaken after recruitment. Males and females also often differ in how they use resources for reproduction and survival. Using a long-term study on body mass and annual survival in yellow-bellied marmots (Marmota flaviventer), we show that body mass was under stabilizing viability selection in the first years of life, before recruitment, which changed to positive directional selection as age increased and animals matured. We found no evidence that viability selection across age classes on body mass differed between sexes. By investigating the link between running speed and body mass, we show that the capacity to escape predators was not consistent across age classes and followed a quadratic relationship at young ages only. Overall, our results indicate that mature age classes exhibit traditional patterns of positive viability selection on body mass, as expected in a hibernating mammal, but that mass in the first years of life is subjected to stabilizing selection which may come from additional predation pressures that negate the benefits of the largest body masses. Our study highlights the importance to disentangle selection pressures on traits across critical age (or life) classes.

How social behaviour and life-history traits change with age and in the year prior to death in female yellow-bellied marmots.

Studies in natural populations are essential to understand the evolutionary ecology of senescence and terminal allocation. While there are an increasing number of studies investigating late-life variation in different life-history traits of wild populations, little is known about these patterns in social behaviour. We used long-term individual based data on yellow-bellied marmots (Marmota flaviventer) to quantify how affiliative social behaviours and different life-history traits vary with age and in the last year of life, and how patterns compare between the two. We found that some social behaviours and all life-history traits varied with age, whereas terminal last year of life effects were only observed in life-history traits. Our results imply that affiliative social behaviours do not act as a mechanism to adjust allocation among traits when close to death, and highlight the importance of adopting an integrative approach, studying late-life variation and senescence across multiple different traits, to allow the identification of potential trade-offs. This article is part of the theme issue 'Ageing and sociality: why, when and how does sociality change ageing patterns?'

Collision between biological process and statistical analysis revealed by mean centring.

Animal ecologists often collect hierarchically structured data and analyse these with linear mixed-effects models. Specific complications arise when the effect sizes of covariates vary on multiple levels (e.g. within vs. among subjects). Mean centring of covariates within subjects offers a useful approach in such situations, but is not without problems. A statistical model represents a hypothesis about the underlying biological process. Mean centring within clusters assumes that the lower level responses (e.g. within subjects) depend on the deviation from the subject mean (relative) rather than on the absolute scale of the covariate. This may or may not be biologically realistic. We show that mismatch between the nature of the generating (i.e. biological) process and the form of the statistical analysis produce major conceptual and operational challenges for empiricists. We explored the consequences of mismatches by simulating data with three response-generating processes differing in the source of correlation between a covariate and the response. These data were then analysed by three different analysis equations. We asked how robustly different analysis equations estimate key parameters of interest and under which circumstances biases arise. Mismatches between generating and analytical equations created several intractable problems for estimating key parameters. The most widely misestimated parameter was the among-subject variance in response. We found that no single analysis equation was robust in estimating all parameters generated by all equations. Importantly, even when response-generating and analysis equations matched mathematically, bias in some parameters arose when sampling across the range of the covariate was limited. Our results have general implications for how we collect and analyse data. They also remind us more generally that conclusions from statistical analysis of data are conditional on a hypothesis, sometimes implicit, for the process(es) that generated the attributes we measure. We discuss strategies for real data analysis in face of uncertainty about the underlying biological process.

Contrasting effects of climate change on seasonal survival of a hibernating mammal.

Seasonal environmental conditions shape the behavior and life history of virtually all organisms. Climate change is modifying these seasonal environmental conditions, which threatens to disrupt population dynamics. It is conceivable that climatic changes may be beneficial in one season but result in detrimental conditions in another because life-history strategies vary between these time periods. We analyzed the temporal trends in seasonal survival of yellow-bellied marmots (Marmota flaviventer) and explored the environmental drivers using a 40-y dataset from the Colorado Rocky Mountains (USA). Trends in survival revealed divergent seasonal patterns, which were similar across age-classes. Marmot survival declined during winter but generally increased during summer. Interestingly, different environmental factors appeared to drive survival trends across age-classes. Winter survival was largely driven by conditions during the preceding summer and the effect of continued climate change was likely to be mainly negative, whereas the likely outcome of continued climate change on summer survival was generally positive. This study illustrates that seasonal demographic responses need disentangling to accurately forecast the impacts of climate change on animal population dynamics.

Older mothers produce more successful daughters.

Annual reproductive success and senescence patterns vary substantially among individuals in the wild. However, it is still seldom considered that senescence may not only affect an individual but also affect age-specific reproductive success in its offspring, generating transgenerational reproductive senescence. We used long-term data from wild yellow-bellied marmots (Marmota flaviventer) living in two different elevational environments to quantify age-specific reproductive success of daughters born to mothers differing in age. Contrary to prediction, daughters born to older mothers had greater annual reproductive success on average than daughters born to younger mothers, and this translated into greater lifetime reproductive success. However, in the favorable lower elevation environment, daughters born to older mothers also had greater age-specific decreases in annual reproductive success. In the harsher higher elevation environment on the other hand, daughters born to older mothers tended to die before reaching ages at which such senescent decreases could be observed. Our study highlights the importance of incorporating environment-specific transgenerational parent age effects on adult offspring age-specific life-history traits to fully understand the substantial variation observed in senescence patterns in wild populations.

Genetic variance for behavioural 'predictability' of stress response.

Genetic factors underpinning phenotypic variation are required if natural selection is to result in adaptive evolution. However, evolutionary and behavioural ecologists typically focus on variation among individuals in their average trait values and seek to characterize genetic contributions to this. As a result, less attention has been paid to if and how genes could contribute towards within-individual variance or trait 'predictability'. In fact, phenotypic 'predictability' can vary among individuals, and emerging evidence from livestock genetics suggests this can be due to genetic factors. Here, we test this empirically using repeated measures of a behavioural stress response trait in a pedigreed population of wild-type guppies. We ask (a) whether individuals differ in behavioural predictability and (b) whether this variation is heritable and so evolvable under selection. Using statistical methodology from the field of quantitative genetics, we find support for both hypotheses and also show evidence of a genetic correlation structure between the behavioural trait mean and individual predictability. We show that investigating sources of variability in trait predictability is statistically tractable and can yield useful biological interpretation. We conclude that, if widespread, genetic variance for 'predictability' will have major implications for the evolutionary causes and consequences of phenotypic variation.

Assessing seasonal demographic covariation to understand environmental-change impacts on a hibernating mammal.

Natural populations are exposed to seasonal variation in environmental factors that simultaneously affect several demographic rates (survival, development and reproduction). The resulting covariation in these rates determines population dynamics, but accounting for its numerous biotic and abiotic drivers is a significant challenge. Here, we use a factor-analytic approach to capture partially unobserved drivers of seasonal population dynamics. We use 40 years of individual-based demography from yellow-bellied marmots (Marmota flaviventer) to fit and project population models that account for seasonal demographic covariation using a latent variable. We show that this latent variable, by producing positive covariation among winter demographic rates, depicts a measure of environmental quality. Simultaneously, negative responses of winter survival and reproductive-status change to declining environmental quality result in a higher risk of population quasi-extinction, regardless of summer demography where recruitment takes place. We demonstrate how complex environmental processes can be summarized to understand population persistence in seasonal environments.

Age, state, environment, and season dependence of senescence in body mass.

Senescence is a highly variable process that comprises both age-dependent and state-dependent components and can be greatly affected by environmental conditions. However, few studies have quantified the magnitude of age-dependent and state-dependent senescence in key life-history traits across individuals inhabiting different spatially structured and seasonal environments. We used longitudinal data from wild female yellow-bellied marmots (Marmota flaviventer), living in two adjacent environments that differ in elevation and associated phenology, to quantify how age and individual state, measured as "time to death," affect body mass senescence in different environments. Further, we quantified how patterns of senescence differed between two biologically distinct seasons, spring, and late summer. Body mass senescence had an age-dependent component, expressed as a decrease in mass in old age. Overall, estimated age-dependent senescence was greater in females living in the more favorable lower elevation environment, than in the harsher higher elevation environment, and greater in late summer than in spring. Body mass senescence also had a state-dependent component, captured by effects of time to death, but only in the more favorable lower elevation environment. In spring, body mass gradually decreased from 2 years before death, whereas in late summer, state-dependent effects were expressed as a terminal decrease in body mass in the last year of life. Contrary to expectations, we found that senescence was more likely to be observed under more favorable environmental conditions, rather than under harsher conditions. By further demonstrating that senescence patterns differ among seasons, our results imply that within-year temporal environmental variation must be considered alongside spatial environmental variation in order to characterize and understand the pattern and magnitude of senescence in wild populations.

Strong social relationships are associated with decreased longevity in a facultatively social mammal.

Humans in strong social relationships are more likely to live longer because social relationships may buffer stressors and thus have protective effects. However, a shortcoming of human studies is that they often rely on self-reporting of these relationships. By contrast, observational studies of non-human animals permit detailed analyses of the specific nature of social relationships. Thus, discoveries that some social animals live longer and healthier lives if they are involved in social grooming, forage together or have more affiliative associates emphasizes the potential importance of social relationships on health and longevity. Previous studies have focused on the impact of social metrics on longevity in obligately social species. However, if sociality indeed has a key role in longevity, we might expect that affiliative relationships should also influence longevity in less social species. We focused on socially flexible yellow-bellied marmots (Marmota flaviventer) and asked whether female longevity covaries with the specific nature of social relationships. We quantified social relationships with social network statistics that were based on affiliative interactions, and then estimated the correlation between longevity and sociality using bivariate models. We found a significant negative phenotypic correlation between affiliative social relationship strength and longevity; marmots with greater degree, closeness and those with a greater negative average shortest path length died at younger ages. We conclude that sociality plays an important role in longevity, but how it does so may depend on whether a species is obligately or facultatively social.