Multifaceted density dependence: Social structure and seasonality effects on Serengeti lion demography.

Interactions between density and environmental conditions have important effects on vital rates and consequently on population dynamics and can take complex pathways in species whose demography is strongly influenced by social context, such as the African lion, Panthera leo. In populations of such species, the response of vital rates to density can vary depending on the social structure (e.g. effects of group size or composition). However, studies assessing density dependence in populations of lions and other social species have seldom considered the effects of multiple socially explicit measures of density, and-more particularly for lions-of nomadic males. Additionally, vital-rate responses to interactions between the environment and various measures of density remain largely uninvestigated. To fill these knowledge gaps, we aimed to understand how a socially and spatially explicit consideration of density (i.e. at the local scale) and its interaction with environmental seasonality affect vital rates of lions in the Serengeti National Park, Tanzania. We used a Bayesian multistate capture-recapture model and Bayesian generalized linear mixed models to estimate lion stage-specific survival and between-stage transition rates, as well as reproduction probability and recruitment, while testing for season-specific effects of density measures at the group and home-range levels. We found evidence for several such effects. For example, resident-male survival increased more strongly with coalition size in the dry season compared with the wet season, and adult-female abundance affected subadult survival negatively in the wet season, but positively in the dry season. Additionally, while our models showed no effect of nomadic males on adult-female survival, they revealed strong effects of nomads on key processes such as reproduction and takeover dynamics. Therefore, our results highlight the importance of accounting for seasonality and social context when assessing the effects of density on vital rates of Serengeti lions and of social species more generally.

Sex-specific social aging in wild African lions.

There is a growing interest in social behavior change with age,1,2,3,4,5 and the impacts of sociality on longevity,6,7,8 but current knowledge is broadly limited to primates, societies structured by dominance hierarchies, or single-sex studies. It is less clear how social aging patterns emerge in carnivores. The African lion (Panthera leo), a species that lives in egalitarian fission-fusion societies, presents an exceptional opportunity to examine social aging. Across felids, lions are unique in their dependence on conspecifics for many essential processes,9,10,11 and there is vast knowledge of lion behavioral ecology,10,11,12,13,14 including documented reproductive senescence in both sexes.14,15 Applying spatial-social network analyses across 30 years of data on the wild Serengeti lion population, we show that sex strongly modulates patterns of social aging and longevity. Group size increased with age for both sexes, but only males experienced significant changes in associate numbers (degree), specifically to females, which peaked in mid-life before declining. While aging females experienced declines in intra-sex connectivity (strength) and bond strength (mean strength), they peaked in both to males during mid-life. Male inter-sex strength also peaked in mid-life, while conversely their intra-sex strength and mean strength significantly dipped in mid-life. Although social associations were important for survival in both sexes, the investment diverged significantly: females' overall network connectivity was key for longevity, while the number of associates was important for males. These findings illustrate important potential effects of social aging in a wild carnivore and demonstrate how these diverge strongly between the sexes.

Interplay of competition and facilitation in grazing succession by migrant Serengeti herbivores.

Competition, facilitation, and predation offer alternative explanations for successional patterns of migratory herbivores. However, these interactions are difficult to measure, leaving uncertainty about the mechanisms underlying body-size-dependent grazing-and even whether succession occurs at all. We used data from an 8-year camera-trap survey, GPS-collared herbivores, and fecal DNA metabarcoding to analyze the timing, arrival order, and interactions among migratory grazers in Serengeti National Park. Temporal grazing succession is characterized by a "push-pull" dynamic: Competitive grazing nudges zebra ahead of co-migrating wildebeest, whereas grass consumption by these large-bodied migrants attracts trailing, small-bodied gazelle that benefit from facilitation. "Natural experiments" involving intense wildfires and rainfall respectively disrupted and strengthened these effects. Our results highlight a balance between facilitative and competitive forces in co-regulating large-scale ungulate migrations.