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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.

The availability of food during early life has been proposed as a key proximate mechanism for the development of variation in behaviour among and within individuals.Individuals can vary amongst each other in their personality, plasticity and predictability and if an individual’s behaviour is correlated across contexts this can lead to behavioural, plasticity and predictability syndromes.In this study, we used a split brood design to raise African clawed frog tadpoles (Xenopus laevis) on a high or low diet in food availability and measured the distance they swam in a familiar and unfamiliar context eight times during their development.In a familiar context, we found that there was an increase in among individual variance in plasticity and predictability in the high food treatment. This shows that when resources are not restricted, individuals are not constrained in the expression of their behaviour at certain phenotypic levels.In an unfamiliar context, we found a different response, with an increase in individual variance in personality in the low but not the high feed tadpoles. As unfamiliar contexts may be riskier, our results highlight that individuals receiving less food may take greater foraging risks in novel contexts.Across contexts, we found a predictability syndrome in the high but not the low feed tadpoles, highlighting that cross-context behaviours can become decoupled in some developmental conditions but remain intact in others.Together our findings show that early life conditions contribute to among individual variation in behaviour but that these may only impact the phenotype at specific phenotypic levels and are context specific.We emphasise that having a fundamental understanding of how early development may promote or constrain individual variation can provide a greater understanding of how individuals and populations may respond to novel conditions brought about by anthropogenic activity.

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.

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.

Repeated social interactions with conspecifics and/or heterospecifics during early development may drive the differentiation of behaviour among individuals. This behavioural differentiation may occur through individuals behaving more different from each other on average and/or individuals behaving more consistently. 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 different behavioural traits. 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, activity, exploration and risk taking in the presence of a predator in focal E. pustulosus tadpoles six times during development. We used univariate and multivariate hierarchical mixed effect models to investigate the effect of treatment on mean behaviour and on among individual variance between and within individuals across behavioural traits. There was a strong effect of competition on behaviour, with different population and individual level responses across social treatments. Within their home tank, individuals were more consistent in their movements under conspecific competition but heterospecific competition caused more variance in the average movement among individuals. Behavioural responses were also trait specific as conspecific competition caused greater variability in movements among individuals in a novel environment. The results highlight that the impact of competition on inter-individual differences in behaviour is dependent on competitor species identity and is trait specific.

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.

Species that hibernate 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 aging suppression. Metabolic rate is also a known correlate of longevity, we thus proposed the ‘hibernation-aging hypothesis’ whereby aging 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 were modeled 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 age change 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 (2-years old). With respect to circannual patterns, the epigenetic age increased during the summer and essentially stalled during the winter. Our enrichment analysis of age-related CpG sites revealed pathways related to development and cell differentiation, while the season-related CpGs enriched pathways related to central carbon metabolism, immune system, and circadian clock. Taken together, our results are consistent with the hibernation-aging hypothesis and may explain the enhanced longevity in hibernators.

The terminal allocation and senescence hypotheses make opposite predictions about how age-specific reproductive effort should vary during old age. There is empirical support for both hypotheses, although reports on senescence are more numerous. Individual heterogeneity and selective mortality, however, decrease our ability to measure how reproductive effort varies during late life. The damage accumulation model proposes that terminal allocation and senescence could be partly age-independent. Using a reverse-age approach, we analysed an unusually complete record of annual reproductive success for 90 bighorn ewes that died between 7 and 18years of age. We estimated age-specific and age-independent variation of reproductive effort in late-life. Reproductive effort decreased in the two last reproductions, independently of age at death. Fecundity also decreased in the last 2years of life, with a steeper decline for older individuals. Our study reveals that reproductive senescence includes both age-dependent and age-independent components.