Societies with fission-fusion dynamics as complex adaptive systems: the importance of scale.

In this article, we argue that social systems with fission-fusion (FF) dynamics are best characterized within a complex adaptive systems (CAS) framework. We discuss how different endogenous and exogenous factors drive scale-dependent network properties across temporal, spatial and social domains. Importantly, this view treats the dynamics themselves as objects of study, rather than variously defined notions of static 'social groups' that have hitherto dominated thinking in behavioural ecology. CAS approaches allow us to interrogate FF dynamics in taxa that do not conform to more traditional conceptualizations of sociality and encourage us to pose new types of questions regarding the sources of stability and change in social systems, distinguishing regular variations from those that would lead to system-level reorganization. This article is part of the theme issue 'Connected interactions: enriching food web research by spatial and social interactions'.

Land-use change is associated with multi-century loss of elephant ecosystems in Asia.

Understanding historic patterns of land use and land cover change across large temporal and spatial scales is critical for developing effective biodiversity conservation management and policy. We quantify the extent and fragmentation of suitable habitat across the continental range of Asian elephants (Elephas maximus) based on present-day occurrence data and land-use variables between 850 and 2015 A.D. We found that following centuries of relative stability, over 64% (3.36 million km2) of suitable elephant habitat across Asia was lost since the year 1700, coincident with colonial-era land-use practices in South Asia and subsequent agricultural intensification in Southeast Asia. Average patch size dropped 83% from approximately 99,000-16,000 km2 and the area occupied by the largest patch decreased 83% from ~ 4 million km2 (45% of area) to 54,000 km2 (~ 7.5% of area). Whereas 100% of the area within 100 km of the current elephant range could have been considered suitable habitat in the year 1700, over half was unsuitable by 2015, driving potential conflict with people. These losses reflect long-term decline of non-forested ecosystems, exceeding estimates of deforestation within this century. Societies must consider ecological histories in addition to proximate threats to develop more just and sustainable land-use and conservation strategies.

Strategies of protected area use by Asian elephants in relation to motivational state and social affiliations.

Animals' space requirements may vary according to life-history and social considerations. We observed 516 wild adult Asian elephants from both sexes, over 9 years, to investigate how life-history traits and social behavior influence protected-area (PA) use at Udawalawe National Park, Sri Lanka. Male PA-use, quantified in terms of average between-sightings-interval (BSI), was significantly influenced by the interaction of age class and motivational state (i.e. reproduction vs. foraging). Musth lengthened with age, with a median of 24.5 days for ages 21-30, 32.5 days for ages 31-40, and 45 days for those > 40. A minority (11%) used it exclusively during musth, while others used it exclusively for foraging (44%) or both (45%). Males using it in both states and older musth-only males were more likely to be seen across years. There were 16 social communities containing between 2-22 adult females. Females' BSI was significantly influenced by social ties, but this relationship was weak, because members of social communities do not necessarily disperse together, resulting in high individual variation in space-use. Inter-annual variability in sightings among individuals of both sexes indicates that around ¾ of the population is likely non-residential across years, challenging the prevailing fortress-conservation paradigm of wildlife management.

Wild Sri Lankan elephants retreat from the sound of disturbed Asian honey bees.

Asian elephants (Elephas maximus) are threatened primarily by habitat loss and human-elephant conflict. In addition to establishing protected areas and corridors for wildlife, empowering farmers to protect their crops is crucial for Asian elephant conservation [1,2]. Elephants can habituate to artificial deterrents, hence natural biological alternatives are of great interest [2,3]. African elephants (Loxodonta africana) avoid African honey bees (Apis mellifera scutellata), inspiring 'beehive fences' as a successful means of small-scale crop protection [4,5]. Here, we used a recording of a disturbed hive of cavity-dwelling Asian honey bees (Apis cerana indica) and conducted sound playbacks to 120 wild elephants in 28 different groups resting under trees in Uda Walawe National Park in Sri Lanka. Elephants responded by moving significantly further away from their resting site in bee playback trials compared to controls. Elephants also increased vocalization rates, as well as investigative and reassurance behaviours in response to bee sounds, but did not display dusting or headshaking behaviour.

Research Priorities from Animal Behaviour for Maximising Conservation Progress.

Poor communication between academic researchers and wildlife managers limits conservation progress and innovation. As a result, input from overlapping fields, such as animal behaviour, is underused in conservation management despite its demonstrated utility as a conservation tool and countless papers advocating its use. Communication and collaboration across these two disciplines are unlikely to improve without clearly identified management needs and demonstrable impacts of behavioural-based conservation management. To facilitate this process, a team of wildlife managers and animal behaviour researchers conducted a research prioritisation exercise, identifying 50 key questions that have great potential to resolve critical conservation and management problems. The resulting agenda highlights the diversity and extent of advances that both fields could achieve through collaboration.

Morphometrics and behavior of a wild Asian elephant exhibiting disproportionate dwarfism.

BACKGROUND: Dwarfism is a condition characterized by shorter stature, at times accompanied by differential skeletal growth proportions relative to the species-typical physical conformation. Causes vary and are well-documented in humans as well as certain mammalian species in captive or laboratory conditions, but rarely observed in the wild. CASE PRESENTATION: We report on a single case of apparent dwarfism in a free-ranging adult male Asian elephant in Sri Lanka, comparing physical dimensions to those of other males in the population as well as in previous literature. The subject M459 was found to have a shoulder height of approximately 195 cm, is shorter than the average height of typical mature males, with a body length of 218 cm. This ratio of body length to height deviates from what is typically observed, which is approximately 1:1, but was similar to the attributes of a dwarf elephant in captivity documented in 1955. We report on behavior including the surprising observation that M459 appears to have a competitive advantage in intrasexual contests. We discuss how this phenotype compares to cases of dwarfism in other non-human animals. CONCLUSION: M459 exemplifies a rare occurrence of disproportionate dwarfism in a free-ranging wild mammal that has survived to reproductive maturity and appears otherwise healthy.

Demographic variables for wild Asian elephants using longitudinal observations.

Detailed demographic data on wild Asian elephants have been difficult to collect due to habitat characteristics of much of the species' remaining range. Such data, however, are critical for understanding and modeling population processes in this endangered species. We present data from six years of an ongoing study of Asian elephants (Elephas maximus) in Uda Walawe National Park, Sri Lanka. This relatively undisturbed population numbering over one thousand elephants is individually monitored, providing cohort-based information on mortality and reproduction. Reproduction was seasonal, such that most births occurred during the long inter-monsoon dry season and peaked in May. During the study, the average age at first reproduction was 13.4 years and the 50(th) percentile inter-birth interval was approximately 6 years. Birth sex ratios did not deviate significantly from parity. Fecundity was relatively stable throughout the observed reproductive life of an individual (ages 11-60), averaging between 0.13-0.17 female offspring per individual per year. Mortalities and injuries based on carcasses and disappearances showed that males were significantly more likely than females to be killed or injured through anthropogenic activity. Overall, however, most observed injuries did not appear to be fatal. This population exhibits higher fecundity and density relative to published estimates on other Asian elephant populations, possibly enhanced by present range constriction. Understanding the factors responsible for these demographic dynamics can shed insight on the future needs of this elephant population, with probable parallels to other populations in similar settings.

An Asian elephant imitates human speech.

Vocal imitation has convergently evolved in many species, allowing learning and cultural transmission of complex, conspecific sounds, as in birdsong. Scattered instances also exist of vocal imitation across species, including mockingbirds imitating other species or parrots and mynahs producing human speech. Here, we document a male Asian elephant (Elephas maximus) that imitates human speech, matching Korean formants and fundamental frequency in such detail that Korean native speakers can readily understand and transcribe the imitations. To create these very accurate imitations of speech formant frequencies, this elephant (named Koshik) places his trunk inside his mouth, modulating the shape of the vocal tract during controlled phonation. This represents a wholly novel method of vocal production and formant control in this or any other species. One hypothesized role for vocal imitation is to facilitate vocal recognition by heightening the similarity between related or socially affiliated individuals. The social circumstances under which Koshik's speech imitations developed suggest that one function of vocal learning might be to cement social bonds and, in unusual cases, social bonds across species.

The dynamics of social networks among female Asian elephants.

BACKGROUND: Patterns in the association of individuals can shed light on the underlying conditions and processes that shape societies. Here we characterize patterns of association in a population of wild Asian Elephants at Uda Walawe National Park in Sri Lanka. We observed 286 individually-identified adult female elephants over 20 months and examined their social dynamics at three levels of organization: pairs of individuals (dyads), small sets of direct companions (ego-networks), and the population level (complete networks). RESULTS: Corroborating previous studies of this and other Asian elephant populations, we find that the sizes of elephant groups observed in the field on any particular day are typically small and that rates of association are low. In contrast to earlier studies, our longitudinal observations reveal that individuals form larger social units that can be remarkably stable across years while associations among such units change across seasons. Association rates tend to peak in dry seasons as opposed to wet seasons, with some cyclicity at the level of dyads. In addition, we find that individuals vary substantially in their fidelity to companions. At the ego-network level, we find that despite these fluctuations, individuals associate with a pool of long-term companions. At the population level, social networks do not exhibit any clear seasonal structure or hierarchical stratification. CONCLUSIONS: This detailed longitudinal study reveals different social dynamics at different levels of organization. Taken together, these results demonstrate that low association rates, seemingly small group sizes, and fission-fusion grouping behavior mask hidden stability in the extensive and fluid social affiliations in this population of Asian elephants.