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Mammals communicate socially through visual, auditory and chemical signals. The chemical sense is the oldest sense and is shared by all organisms including bacteria. Despite mounting evidence for social chemo-signaling in humans, the extent to which it modulates behavior is debated and can benefit from comparative models of closely related hominoids. The use of odor cues in wild ape social communication has been only rarely explored. Apart from one study on wild chimpanzee sniffing, our understanding is limited to anecdotes. We present the first study of wild gorilla chemo-communication and the first analysis of olfactory signaling in relation to arousal levels and odor strength in wild apes. If gorilla scent is used as a signaling mechanism instead of only a sign of arousal or stress, odor emission should be context specific and capable of variation as a function of the relationships between the emitter and perceiver(s). Measured through a human pungency scale, we determined the factors that predicted extreme levels of silverback odor for one wild western lowland gorilla (Gorilla gorilla gorilla) group silverback. Extreme silverback odor was predicted by the presence and intensity of inter-unit interactions, silverback anger, distress and long-calling auditory rates, and the absence of close proximity between the silverback and mother of the youngest infant. Odor strength also varied according to the focal silverback's strategic responses during high intensity inter-unit interactions. Silverbacks appear to use odor as a modifiable form of communication; where odor acts as a highly flexible, context dependent signaling mechanism to group members and extra-group units. The importance of olfaction to ape social communication may be especially pertinent in Central African forests where limited visibility may necessitate increased reliance on other senses.

The structure and dynamics of primate social groups are shaped by the social relationships of its members. These relationships are based on different types of interactions and vary in relation to the identity of the interactants and over time. Social network analysis tools represent a powerful and comprehensive method to characterise social interactions and recent methodological advances now allow the study of the multidimensionality of sociality via multilayer networks that incorporate multiple types of interactions. Here, we use a multidimensional network approach to investigate the multidimensionality of sociality of females in a captive group of mandrills. We constructed two multiplex networks based on agonistic, proximity and grooming interactions of 6-7 mature females to analyse the multidimensionality of relationships within two independent observation periods; and three multiplex networks (one for each interaction type) to examine how relationships changed between periods. Within each period, different individuals were the most central in each layer and at the multiplex level, and different layers (i.e., interaction types) contributed non-redundant information to the multilayer structure. Across periods, relationships based on the same interaction type also contained non-redundant information. These results indicate that female mandrills engage in multidimensional and dynamic relationships, suggesting that in order to represent the full complexity of relationships, networks need to be constructed from more than a single type of interaction and across time. Our results provide evidence for the potential value of the multilayer network approach to characterise the multidimensionality of primate sociality.

Long-lived species such as elephants, whales and primates exhibit extended post-fertile survival compared to species with shorter lifespans but data on age-related fecundity and survival are limited to few species or populations. We assess relationships between longevity, reproductive onset, reproductive rate and age for 834 longitudinally monitored wild female African elephants in Amboseli, Kenya. The mean known age at first reproduction was 13.8 years; only 5 % commenced reproduction by 10 years. Early reproducers (<12.5 years) had higher age-specific fertility rates than did females who commenced reproduction late (15+ years) with no differences in survival between these groups. Age-specific reproductive rates of females dying before 40 years were reduced by comparison to same-aged survivors, illustrating a mortality filter and reproductive advantages of a long life. Overall, 95 % of fertility was completed before 50, and 95 % of mortality experienced by age 65, with a mean life expectancy of 41 years for females who survived to the minimum age at first birth (9 years). Elephant females have a relatively long period (c. 16 years) of viability after 95 % completed fertility, although reproduction does not entirely cease until they are over 65. We found no evidence of increased investment among females aged over 40 in terms of delay to next birth or calf mortality. The presence of a mother reproducing simultaneously with her daughter was associated with higher rates of daughter reproduction suggesting advantages from maternal (and grandmaternal) co-residence during reproduction.

Comparative approaches to the evolution of primate social behavior have typically involved two distinct lines of inquiry. One has focused on phylogenetic analyses that treat social traits as static, species-specific characteristics; the other has focused on understanding the behavioral flexibility of particular populations or species in response to local ecological or demographic variables. Here, we combine these approaches by distinguishing between constraining traits such as dispersal regimes (male, female, or bi-sexual), which are relatively invariant, and responding traits such as grouping patterns (stable, fission-fusion, sometimes fission-fusion), which can reflect rapid adjustments to current conditions. Using long-term and cross-sectional data from 29 studies of 22 species of wild primates, we confirm that dispersal regime exhibits a strong phylogenetic signal in our sample. We then show that primate species with high variation in group size and adult sex ratios exhibit variability in grouping pattern (i.e., sometimes fission-fusion) with dispersal regime constraining the grouping response. When assessing demographic variation, we found a strong positive relationship between the variability in group size over time and the number of observation years, which further illustrates the importance of long-term demographic data to interpretations of social behavior. Our approach complements other comparative efforts to understand the role of behavioral flexibility by distinguishing between constraining and responding traits, and incorporating these distinctions into analyses of social states over evolutionary and ecological time.

Social structure is proposed to influence the transmission of both directly and environmentally transmitted infectious agents. However in natural populations, many other factors also influence transmission, including variation in individual susceptibility and aspects of the environment that promote or inhibit exposure to infection. We used a population genetic approach to investigate the effects of social structure, environment, and host traits on the transmission of Escherichia coli infecting two populations of wild elephants: one in Amboseli National Park and another in Samburu National Reserve, Kenya. If E. coli transmission is strongly influenced by elephant social structure, E. coli infecting elephants from the same social group should be genetically more similar than E. coli sampled from members of different social groups. However, we found no support for this prediction. Instead, E. coli was panmictic across social groups, and transmission patterns were largely dominated by habitat and host traits. For instance, habitat overlap between elephant social groups predicted E. coli genetic similarity, but only in the relatively drier habitat of Samburu, and not in Amboseli, where the habitat contains large, permanent swamps. In terms of host traits, adult males were infected with more diverse haplotypes, and males were slightly more likely to harbor strains with higher pathogenic potential, as compared to adult females. In addition, elephants from similar birth cohorts were infected with genetically more similar E. coli than elephants more disparate in age. This age-structured transmission may be driven by temporal shifts in genetic structure of E. coli in the environment and the effects of age on bacterial colonization. Together, our results support the idea that, in elephants, social structure often will not exhibit strong effects on the transmission of generalist, fecal-oral transmitted bacteria. We discuss our results in the context of social, environmental, and host-related factors that influence transmission patterns.

Flagship species are used to promote conservation and tourism. Africa's famous ‘Big five’ have become marketing flagships that fundraisers and tourism promoters emulate globally. Species can be selected systematically for marketing using characteristics such as colour, size or behaviour, but this approach can overlook unique animals or homogenise selections. Alternatively, polling the public can reveal existing preferences for animals directly. We used questionnaires with tourists in the Peruvian Amazon to identify existing biases for species and rank them for suitability for tourism and conservation marketing. Polling revealed several species that would not be considered good flagship candidates using systematic methods based on species characteristics. ‘Free listing’ tourists expressed preferences at inconsistent taxonomic levels. The response ‘monkeys’ (infraorder Simiiformes) was highest ranked, followed by ‘jaguar’ (Panthera onca), ‘Amazon dolphin’ (Inia geoffrensis), ‘sloths’ (suborder Folivora), and ‘caiman’ (subfamily Caimaninae) and ‘birds’ (class Aves). When ranking from a preselected shortlist, jaguar, Amazon dolphins and sloths (represented by Bradypus variegatus) remained popular, while vote splitting within higher taxonomic levels, in particular monkeys, made room for green‐winged macaw (Ara chloropterus) and anaconda (Eunectes murinus). When asked about their willingness to pay for excursions or donate to conservation, tourists were overwhelmingly more likely to quote larger figures for jaguars than any other species, but results for other species were more homogenous. Some popular taxonomic groups are diverse in Amazonia; up to 14 monkey species may be present at some sites Amazonia, alongside several hundred bird species. A Big five strategy obscures this diversity. Using physical characteristics as selection criteria underplays diversity and overlooks popular taxa—notably sloths for the Amazon. A strategy of polling the public to select popular species as flagships more directly identifies salient species for marketing and efficiently considers existing biases. However, diversity will trump a Big five approach in megadiverse areas. Flagship species are used to promote conservation and tourism, but selecting species purely for prescribed appealing characteristics may overlook unique flagships or homogenise selections. A strategy of polling the public can more directly identify existing preferences for species for marketing. Polling tourists in the Peruvian Amazon revealed that preferences for jaguars, Amazon dolphins and sloths are strong, but vote splitting within popular diverse groups like monkeys, leads to thier omission from Big Five rankings. In megadiverse areas, promoting diversity may trump the Big Five approach.

We explore elephant play behaviour since (a) play has been proposed to represent a potential welfare indicator; and (b) play has been associated with long-term survival in the wild. We categorised play into four types, and investigate both social (gentle, escalated-contact) and non-social (lone-locomotor, exploratory-object) play from observations made on wild (Asian N = 101; African N = 130) and captive (Asian N = 8; African N = 7) elephant calves ranging in age from birth to five years. Social play was the most frequent type of play among immature elephants, accounting for an average of 3%–9% of active time. Non-social play accounted for an additional 1%–11% of time. The most time spent in play was seen in captive Asian calves, particularly at the ages of 1–6 months, while wild African calves spent the least time in play overall, even though they had the greatest number and most diverse range of play partners available. We assessed calf energetics using time spent suckling, resting, moving and independent feeding. Time spent playing was unrelated to time spent suckling but negatively associated with time spent independently feeding. There were no associations with time spent moving or resting. Maternal energy via lactation was unrelated to play early in life, but energy acquired independently may constrain or enable play. Play, while a potential indicator of compromised welfare for many species when absent, can act as a highly stimulating activity for captive elephants in the absence of other forms of arousal.

African elephants (Loxodonta africana) use unusual and restricted habitats such as swampy clearings, montane outcrops and dry rivers for a variety of social and ecological reasons. Within these habitats, elephants focus on very specific areas for resource exploitation, resulting in deep caves, large forest clearings and sand pits as well as long-established and highly demarcated routes for moving between resources. We review evidence for specific habitat exploitation in elephants and suggest that this represents socially learned cultural behaviour. Although elephants show high fidelity to precise locations over the very long term, these location preferences are explained neither by resource quality nor by accessibility. Acquiring techniques for exploiting specific resource sites requires observing conspecifics and practice and is evidence for social learning. Elephants possess sophisticated cognitive capacities used to track relationships and resources over their long lifespans, and they have an extended period of juvenile dependency as a result of the need to acquire this considerable social and ecological knowledge. Thus, elephant fidelity to particular sites results in traditional behaviour over generations, with the potential to weaken relationships between resource quality and site preferences. Illustrating the evidence for such powerful traditions in a species such as elephants contributes to understanding animal cognition in natural contexts.

ContextEast African ecosystems are characterized by the migrations of large herbivores that are highly vulnerable to the recent development of anthropogenic land use change.ObjectivesWe analyzed land cover changes in the Kenyan-Tanzanian borderlands of the greater Amboseli ecosystem to evaluate landscape connectivity using African elephants as an indicator species.MethodsWe used multi-temporal Landsat imagery and a post classification approach to monitor land cover changes over a 43-year period. GIS based methods were accompanied by a literature review for spatial data on land cover changes and elephant migrations.ResultsLand cover changed considerably between 1975 and 2017. Wood- and bushlands declined by 16.3% while open grasslands increased throughout the study region (+ 10.3%). Agricultural expansion was observed (+ 12.2%) occupying important wildlife habitats and narrowing migration corridors. This development has led to the isolation of Nairobi National Park which was previously part of a large contiguous ecosystem. Eight migration corridors were identified of which only one is formally protected. Two others are almost completely blocked by agriculture and three are expected to become endangered under continuing land use changes.ConclusionsLandscape connectivity is still viable for this ecosystem (except for Nairobi National Park). However, the current situation is very fragile as anthropogenic land use changes are threatening most of the identified large mammal migration corridors. Sustainable land use planning with regard to important wildlife habitats and connecting corridors is a crucial task for further conservation work to safeguard a viable future for wildlife populations in the Kenyan-Tanzanian borderlands.