Links between energy budgets, somatic condition, and life history reveal heterogeneous energy management tactics in a group-living mesocarnivore.

BACKGROUND: Optimal management of voluntary energy expenditure is crucial to the survival and reproductive success of wild animals. Nevertheless, a growing appreciation of inter-individual variation in the internal state driving movement suggests that individuals may follow different, yet equally optimal tactics under the same environmental conditions. However, few studies in wild populations have investigated the occurrence and demographic context of different contemporaneous energetic expenditure tactics. Here, we explore this neglected aspect of energy budgeting in order to determine the effect of life-history traits such as age and reproductive status on the co-occurrence of different energy-budgeting tactics in wild populations. METHODS: We investigated inter-individual heterogeneity in energy expenditure within a wild population of European badgers (Meles meles) by quantifying individual overall dynamic body acceleration (ODBA, from tri-axial accelerometry collars) and total daily energy expenditure (DEE, from doubly-labelled water) during 6-9 day deployments and dosing periods over six different seasons (spring, summer, and autumn) in 2018-2019. We obtained ODBA values for 41 deployments (24 unique badgers) and DEE measurements for 41 dosings (22 unique badgers). We then evaluated correlations between these energetic metrics and computed individual ratios of ODBA/DEE as a proxy for the proportion of total energy spent on activity. We measured the impact of alternative ODBA/DEE ratios on body condition, and use survival models constructed using 29 years of demographic data from the same population to situate body-condition changes in the context of age and reproductive status. RESULTS: Both ODBA and DEE were highly variable between individuals and exhibited season-specific relationships with individual body condition and life-history factors. DEE scaled allometrically with body weight, but only in summer and autumn; post-reproductive female badgers were lighter than other badgers during the spring but expended on average 350 kJ/day more than predicted from allometric scaling. Older badgers expended significantly less energy on movement during the summer than did younger adults. The ratio of ODBA to DEE (OD) provides a measure of proportional investment into movement. This ratio correlated more significantly with next-season body condition than either energetic metric did independently. However, the majority of individuals with high OD ratios were either younger badgers or reproductive females, for which lower body condition typically presented less of a mortality risk in previous analyses of this population. CONCLUSIONS: Within a single population under the same environmental conditions, we found wide inter-individual variation in both mechanical and total energy expenditure. The adoption of different tactics aligns with relationships between life-history parameters and mortality risk previously studied within the population. Crucially, younger badgers and reproductive females appeared able to tolerate energy expenditure tactics that depleted their body condition more than other badgers. These findings provide a mechanism by which differences in individual energetic context set by life history can maintain heterogeneity in wild populations, providing a wide range of potential energetic tactics under changing environmental conditions.

Early-life seasonal, weather and social effects on telomere length in a wild mammal.

Early-life environmental conditions can provide a source of individual variation in life-history strategies and senescence patterns. Conditions experienced in early life can be quantified by measuring telomere length, which can act as a biomarker of survival probability in some species. Here, we investigate whether seasonal changes, weather conditions and group size are associated with early-life and/or early-adulthood telomere length in a wild population of European badgers (Meles meles). We found substantial intra-annual changes in telomere length during the first 3 years of life, where within-individual effects showed shorter telomere lengths in the winter following the first spring and a trend for longer telomere lengths in the second spring compared to the first winter. In terms of weather conditions, cubs born in warmer, wetter springs with low rainfall variability had longer early-life (3-12 months old) telomeres. Additionally, cubs born in groups with more cubs had marginally longer early-life telomeres, providing no evidence of resource constraint from cub competition. We also found that the positive association between early-life telomere length and cub survival probability remained when social and weather variables were included. Finally, after sexual maturity, in early adulthood (i.e., 12-36 months) we found no significant association between same-sex adult group size and telomere length (i.e., no effect of intrasexual competition). Overall, we show that controlling for seasonal effects, which are linked to food availability, is important in telomere length analyses, and that variation in telomere length in badgers reflects early-life conditions and also predicts first year cub survival.

Overcoming racism in the twin spheres of conservation science and practice.

It is time to acknowledge and overcome conservation's deep-seated systemic racism, which has historically marginalized Black, Indigenous and people of colour (BIPOC) communities and continues to do so. We describe how the mutually reinforcing 'twin spheres' of conservation science and conservation practice perpetuate this systemic racism. We trace how institutional structures in conservation science (e.g. degree programmes, support and advancement opportunities, course syllabuses) can systematically produce conservation graduates with partial and problematic conceptions of conservation's history and contemporary purposes. Many of these graduates go on to work in conservation practice, reproducing conservation's colonial history by contributing to programmes based on outmoded conservation models that disproportionately harm rural BIPOC communities and further restrict access and inclusion for BIPOC conservationists. We provide practical, actionable proposals for breaking vicious cycles of racism in the system of conservation we have with virtuous cycles of inclusion, equality, equity and participation in the system of conservation we want.

Climate change and anthropogenic food manipulation interact in shifting the distribution of a large herbivore at its altitudinal range limit.

Ungulates in alpine ecosystems are constrained by winter harshness through resource limitation and direct mortality from weather extremes. However, little empirical evidence has definitively established how current climate change and other anthropogenic modifications of resource availability affect ungulate winter distribution, especially at their range limits. Here, we used a combination of historical (1997-2002) and contemporary (2012-2015) Eurasian roe deer (Capreolus capreolus) relocation datasets that span changes in snowpack characteristics and two levels of supplemental feeding to compare and forecast probability of space use at the species' altitudinal range limit. Scarcer snow cover in the contemporary period interacted with the augmented feeding site distribution to increase the elevation of winter range limits, and we predict this trend will continue under climate change. Moreover, roe deer have shifted from historically using feeding sites primarily under deep snow conditions to contemporarily using them under a wider range of snow conditions as their availability has increased. Combined with scarcer snow cover during December, January, and April, this trend has reduced inter-annual variability in space use patterns in these months. These spatial responses to climate- and artificial resource-provisioning shifts evidence the importance of these changing factors in shaping large herbivore spatial distribution and, consequently, ecosystem dynamics.

What lies beneath? Population dynamics conceal pace-of-life and sex ratio variation, with implications for resilience to environmental change.

Life-history and pace-of-life syndrome theory predict that populations are comprised of individuals exhibiting different reproductive schedules and associated behavioural and physiological traits, optimized to prevailing social and environmental factors. Changing weather and social conditions provide in situ cues altering this life-history optimality; nevertheless, few studies have considered how tactical, sex-specific plasticity over an individual's lifespan varies in wild populations and influences population resilience. We examined the drivers of individual life-history schedules using 31 years of trapping data and 28 years of pedigree for the European badger (Meles meles L.), a long-lived, iteroparous, polygynandrous mammal that exhibits heterochrony in the timing of endocrinological puberty in male cubs. Our top model for the effects of environmental (social and weather) conditions during a badger's first year on pace-of-life explained <10% of variance in the ratio of fertility to age at first reproduction (F/α) and lifetime reproductive success. Conversely, sex ratio (SR) and sex-specific density explained 52.8% (males) and 91.0% (females) of variance in adult F/α ratios relative to the long-term population median F/α. Weather primarily affected the sexes at different life-history stages, with energy constraints limiting the onset of male reproduction but playing a large role in female strategic energy allocation, particularly in relation to ongoing mean temperature increases. Furthermore, the effects of social factors on age of first reproduction and year-to-year reproductive success covaried differently with sex, likely due to sex-specific responses to potential mate availability. For females, low same-sex densities favoured early primiparity; for males, instead, up to 10% of yearlings successfully mated at high same-sex densities. We observed substantial SR dynamism relating to differential mortality of life-history strategists within the population, and propose that shifting ratios of 'fast' and 'slow' life-history strategists contribute substantially to population dynamics and resilience to changing conditions.