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Prolonged maternal care is vital to the well-being of many long-lived mammals. 1 The premature loss of maternal care, i.e., orphaning, can reduce offspring survival even after weaning is complete. 2-5 However, ecologists have not explicitly assessed how orphaning impacts population growth. We examined the impact of orphaning on population growth in a free-ranging African elephant population, using 19 years of individual-based demographic monitoring data. We compared orphan and nonorphan survival, performed a sensitivity analysis to understand how population growth responds to the probability of being orphaned and orphan survival, and investigated how sensitivity to these orphan parameters changed with level of poaching. Orphans were found to have lower survival compared to nonorphaned age mates, and population growth rate was negatively correlated with orphaning probability and positively correlated with orphan survival. This demonstrates that, in addition to its direct effects, adult elephant death indirectly decreases population growth through orphaning. Population growth rate's sensitivity to orphan survival increased for the analysis parameterized using only data from years of more poaching, indicating orphan survival is more important for population growth as orphaning increases. We conclude that orphaning substantively decreases population growth for elephants and should not be overlooked when quantifying the impacts of poaching. Moreover, we conclude that population models characterizing systems with extensive parental care benefit from explicitly incorporating orphan stages and encourage research into quantifying effects of orphaning in other social mammals of conservation concern., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2021 Elsevier Inc. All rights reserved.)

Over the last two millennia, and at an accelerating pace, the African elephant (Loxodonta spp. Lin.) has been threatened by human activities across its range. 1-7 We investigate the correlates of elephant home range sizes across diverse biomes. Annual and 16-day elliptical time density home ranges 8 were calculated by using GPS tracking data collected from 229 African savannah and forest elephants (L. africana and L. cyclotis, respectively) between 1998 and 2013 at 19 sites representing bushveld, savannah, Sahel, and forest biomes. Our analysis considered the relationship between home range area and sex, species, vegetation productivity, tree cover, surface temperature, rainfall, water, slope, aggregate human influence, and protected area use. Irrespective of these environmental conditions, long-term annual ranges were overwhelmingly affected by human influence and protected area use. Only over shorter, 16-day periods did environmental factors, particularly water availability and vegetation productivity, become important in explaining space use. Our work highlights the degree to which the human footprint and existing protected areas now constrain the distribution of the world's largest terrestrial mammal. 9 , 10 A habitat suitability model, created by evaluating every square kilometer of Africa, predicts that 18,169,219 km 2 would be suitable as elephant habitat-62% of the continent. The current elephant distribution covers just 17% of this potential range of which 57.4% falls outside protected areas. To stem the continued extirpation and to secure the elephants' future, effective and expanded protected areas and improved capacity for coexistence across unprotected range are essential., Competing Interests: Declaration of interests The authors declare no competing interests., (Copyright © 2021 The Authors. Published by Elsevier Inc. All rights reserved.)

Network resilience to perturbation is fundamental to functionality in systems ranging from synthetic communication networks to evolved social organization [1]. While theoretical work offers insight into causes of network robustness, examination of natural networks can identify evolved mechanisms of resilience and how they are related to the selective pressures driving structure. Female African elephants (Loxodonta africana) exhibit complex social networks with node heterogeneity in which older individuals serve as connectivity hubs [2, 3]. Recent ivory poaching targeting older elephants in a well-studied population has mirrored the targeted removal of highly connected nodes in the theoretical literature that leads to structural collapse [4, 5]. Here we tested the response of this natural network to selective knockouts. We find that the hierarchical network topology characteristic of elephant societies was highly conserved across the 16-year study despite ∼70% turnover in individual composition of the population. At a population level, the oldest available individuals persisted to fill socially central positions in the network. For analyses using known mother-daughter pairs, social positions of daughters during the disrupted period were predicted by those of their mothers in years prior, were unrelated to individual histories of family mortality, and were actively built. As such, daughters replicated the social network roles of their mothers, driving the observed network resilience. Our study provides a rare bridge between network theory and an evolved system, demonstrating social redundancy to be the mechanism by which resilience to perturbation occurred in this socially advanced species., (Copyright © 2016 Elsevier Ltd. All rights reserved.)